Dietary Fiber in Human Nutrition

CRC HANDBOOK OF Dietary Fiber in Human Nutrition 3rd Edition CRC HANDBOOK OF Dietary Fiber in Human Nutrition 3rd Edition Edited by Gene A. Spiller, D.Sc., Ph.D. Director Health Research and Studies Center, Inc. SPHERA Foundation Los Altos, California CRC Press Boca Raton London New York Washington, D.C. 2387_FM_fm Page ii Sunday, May 6, 2001 6:18 PM Library of Congress Cataloging-in-Publication Data CRC Handbook of dietary fiber in human nutrition / edited by Gene A. Spiller.—3rd ed. p. cm. Includes bibliographical references and index. ISBN 0-8493-2387-8 (alk. paper) 1. Fiber in human nutrition—Handbooks, manuals, etc. I. Spiller, Gene A. QP144.F52 C73 2001 613.2'8—dc21 2001025278 CIP This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with permission, and sources are indicated. A wide variety of references are listed. Reasonable efforts have been made to publish reliable data and information, but the author and the publisher cannot assume responsibility for the validity of all materials or for the consequences of their use. Neither this book nor any part may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, microfilming, and recording, or by any information storage or retrieval system, without prior permission in writing from the publisher. All rights reserved. Authorization to photocopy items for internal or personal use, or the personal or internal use of specific clients, may be granted by CRC Press LLC, provided that $1.50 per page photocopied is paid directly to Copyright clearance Center, 222 Rosewood Drive, Danvers, MA 01923 USA. The fee code for users of the Transactional Reporting Service is ISBN 0-8493-2387-8/01/$0.00+$1.50. The fee is subject to change without notice. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. The consent of CRC Press LLC does not extend to copying for general distribution, for promotion, for creating new works, or for resale. Specific permission must be obtained in writing from CRC Press LLC for such copying. Direct all inquiries to CRC Press LLC, 2000 N.W. Corporate Blvd., Boca Raton, Florida 33431. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation, without intent to infringe. Visit the CRC Press Web site at www.crcpress.com © 2001 by CRC Press LLC No claim to original U.S. Government works International Standard Book Number 0-8493-2387-8 Library of Congress Card Number 2001025278 Printed in the United States of America 1 2 3 4 5 6 7 8 9 0 Printed on acid-free paper 2387_FM_fm Page iii Sunday, May 6, 2001 6:18 PM Dedication To Hugh Trowell and Denis Burkitt, who pioneered a new understanding of the role of plant fibers in human health. They have been to me teachers of science, medicine, and humility. They should be remembered not only as pioneers in medicine, but also for their selfless and total dedication to the prevention of disease and human suffering. 2387_FM_fm Page v Sunday, May 6, 2001 6:18 PM Preface to the First Edition The CRC Handbook of Dietary Fiber in Human Nutrition is proof of the amazing growth in the study of dietary fiber during the 1970s and 1980s. When I first edited a book on dietary fiber, Fiber in Human Nutrition, in the mid-1970s, I was happy to find at least a few good investigators who could contribute chapters to it. It would have been impossible to find a large number of scientists that could have made a major contribution to that early book. However, as work was beginning on this Handbook in 1982 and I was presenting the design at the Symposium on Fiber in Human and Animal Nutrition in New Zealand that year, not only was I able to find all the 50-plus authors I needed, but I had the sad task of omitting some outstanding names. Interest in what used to be the disregarded Cinderella nutrient of the early 20th century has grown at a rate greater than almost any other nutrient. The plant cell wall and the gums and mucilages had found their well-deserved niche in nutrition and medicine. Even though few people still believe that dietary fiber has not found the ultimate proof that makes it a required nutrient, the momentum is with the ones who have found valid uses for it in high-fiber foods in treating diseases such as type-II diabetes. The momentum is also with the epidemiologists who have found correlations with lower incidence of colorectal cancer, the ones that have found in high-fiber foods one of the best ways to prevent excessive fat and food intake. It is a list that can go on and on. Has the ultimate study on the long-range effects of dietary fiber been published? Of course not. It is probably impossible to carry out the ultimate study on the correlation of nutrition to chronic diseases under present conditions: we must accept the pieces of evidence derived from good epidemiology, from controlled human studies and, of course, animal studies. The lifetimes of many of us would be needed to satisfy the purists who hope for the ultimate study, in this or any other field in which we deal with the lifetime of a human being. There is more to complicate things: dietary fiber polymers are quite elusive, unlike vitamins that can be isolated or synthesized readily. Some of these polymers may change when torn apart from their complex structure in the cell wall. Thus, when we attempt to extract them, too often we isolate something quite different from the original material, perhaps useful, but most certainly different. This bothers many scientists who would like to use the pure form of nutrients for their investigations. There is more: dietary fiber is so interactive that other components of the diet probably vary its effect on humans. All this makes dietary fiber research so challenging and so difficult! This book is proof that there are many dedicated scientists and clinicians that have given their best efforts to dietary fiber. There are many who could not be included here, as there is a point in an effort of this kind at which the editor must sadly stop asking for contributions and recognize that the book must be a finite number of pages. I owe a very special thanks to all the authors in this book, for, after all, it is their book. The book presents a large volume of data. The reader is directed to the Table of Contents which illustrates the design of the book, a design that was conceived to make it as easy as possible to find the needed data. Chemistry, analytical methodologies, physiological and biochemical aspects, clinical and epidemiological studies, and consumption patterns are covered extensively. Tables with the dietary fiber content of various foods analyzed by different methods are given at the end of the book. Gene A. Spiller Los Altos, California — February 1985 2387_FM_fm Page vi Sunday, May 6, 2001 6:18 PM Preface to the Second Edition Dietary fiber research has seen a great deal of progress since the first edition of this Handbook. This new edition is revised and updated by individual authors and, with the exception of a few chapters, such as the Southgate and the crude fiber analytical methods, that are unchanged, new material is added. Some new authors and chapters have been added, including the new method of analysis by Englyst and Hudson (Chapter 3.3) and Bosello, Armellini, and Zamboni’s chapter on fiber consumption in Italy. Hugh Trowell passed away in 1989; his chapter has been left untouched. A major addition in the second edition is the inclusion of more extensive tables of data on dietary fiber in foods. They are prepared by various authors to give the reader a chance to compare data from different sources or methods. This new edition should give researchers, physicians, nutritionists, and other health professionals a useful and ready source of information, as a handbook should. Again, as in the first edition, we could not ask all the experts in this field to contribute, so we decided to stay with the original authors as much as possible. We hope that our efforts will make this work valuable to everyone interested in this topic. Gene A. Spiller Health Research and Studies Center, Inc. SPHERA Foundation, Los Altos, California — January 1992 2387_FM_fm Page vii Sunday, May 6, 2001 6:18 PM Preface to the Third Edition Dietary fiber research has seen a great deal of progress since the first and second editions of this Handbook. This new edition has been revised and updated by individual authors, with the exception of a few chapters that either have historical value or have data that should be available in a handbook of this nature, even though it may have been written a few years before the publication. I like to define the latter as “classic” chapters that have an essential, timeless nature. Some new authors and chapters have been added, including a new section on cereal fiber that emphasizes the crucial role of associated phytochemicals. Hugh Trowell passed away in 1989; his chapter has been left untouched. Denis Burkitt passed away soon after the publication of the second edition, in which he had written the opening chapter with me. I have chosen to expand this first chapter without changing the part published in the second edition of the Handbook. I have added a second part to that chapter to bring it up to date. This new edition should give researchers, physicians, nutritionists, and other health professionals a useful and ready source of information, as a handbook should. Again, as in the other editions, we could not ask all the experts in this field to contribute. We hope that our efforts will make this work valuable to everyone interested in this topic. One new chapter (2.8) discusses a fiber from animal sources. This is an interesting topic in fiber definitions, as dietary fiber is considered to be from plant sources. With all the great progress we have made in the past 10 years in nutrition research, it is unfortunate that the past 10 years have also seen conflicting studies on fiber, as on other topics in nutrition. In the late 1970s, Hugh Trowell, during one of our many meetings, expressed to me his concern that in the earnest desire to do more research, the picture of the benefits of fiber may become “muddled.” I would like to close this preface with an appeal to all fiber researchers to be careful in reaching sweeping conclusions before considering carefully the complexity and the interactivity of dietary fiber. Some major publications have reached conclusions on the effects of fiber after studying diets that were a long way from diets that could be called high-fiber diets. Dietary fiber is a precious gift that plants bring to us. Let’s research it and teach about it with care. Gene A. Spiller Health Research and Studies Center, Inc. SPHERA Foundation, Los Altos, California — December 2000 2387_FM_fm Page viii Sunday, May 6, 2001 6:18 PM Acknowledgments The editor wishes to thank Monica Alton Spiller for her extensive assistance in all phases of editing, from the original design to the various stages of manuscript assessment and the final proofreading of the first edition of this Handbook. For the second edition of this Handbook, the editor wishes to thank Rebecca Carr and Monica Alton Spiller for their assistance in the editing process. For the third edition of this Handbook, the editor wishes to thank Rosemary Schmele, Connie Burton, and Len Marquart for their assistance. 2387_FM_fm Page ix Sunday, May 6, 2001 6:18 PM The Editor Gene A. Spiller is the director of the Health Research and Studies Center and of the SPHERA Foundation in Los Altos, California. Dr. Spiller received a doctorate in chemistry from the University of Milan (Italy) and later a master’s degree and a Ph.D. in nutrition from the University of California at Berkeley. He did additional studies at the Stanford University School of Medicine, Stanford, California. He is a Fellow of the American College of Nutrition, a Certified Nutrition Specialist, and a member of many professional nutrition societies. In the 1970s, Dr. Spiller was head of Nutritional Physiology at Syntex Research in Palo Alto, California, where he did extensive research on dietary fiber and related topics. At the same time he edited many clinical nutrition books. He continued his work in clinical nutrition research and publishing in the 1980s and 1990s, as a consultant and as the director of the Health Research and Studies Center and of the SPHERA Foundation. Many human clinical studies, reviews, and other publications were the results of this work. Dr. Spiller has carried out clinical studies on the effect of dietary fiber and high-fiber foods including nuts, raisins, and whole grains. Other studies have focused on antioxidants and lipids. He is a lecturer in nutrition at Foothill College in the San Francisco Bay Area and earlier taught at Mills College in Oakland, California. Dr. Spiller is the editor of many clinical nutrition books on fiber and other topics at the forefront of nutrition research, such as caffeine and lipids. Among his multi-author books on fiber are Fiber in Human Nutrition (Plenum, 1975), Topics in Dietary Fiber Research (Plenum, 1978), and Medical Aspects of Dietary Fiber (Plenum, 1980), followed by the CRC Handbook of Fiber in Human Nutrition, 1st and 2nd Editions (CRC Press, 1985 and 1992). He has a special interest in lesser-known nutritional factors that may be beneficial to human health, though not essential to life, especially factors that are present in plant foods and that may work together with dietary fiber in the prevention of degenerative diseases. Dr. Spiller has been responsible for organizing international workshops on dietary fiber, such as the International Nutrition Congress in Brazil in 1978 and in Brighton (U.K.) in 1985, and has chaired many symposia and sessions on this and related topics at national meetings of various scientific societies. 2387_FM_fm Page x Sunday, May 6, 2001 6:18 PM Contributors Abayomi O. Akanji, M.D., Ph.D. Department of Pathology Kuwait University Faculty of Medicine Kuwait Sheila Bingham, Ph.D. Medical Research Council Dunn Clinical Nutrition Centre Cambridge, England Per Åman, Dr. Agr. Professor of Plant Foods Department of Food Science Swedish University of Agricultural Sciences Uppsala, Sweden Ottavio Bosello, M.D. Professor Istituto di Clinica Medica Policlinico di Borgo Roma Università di Verona Verona, Italy James W. Anderson, M.D. Professor of Medicine and Clinical Nutrition V.A. Medical Center and University of Kentucky Lexington, Kentucky Roger Andersson, Ph.D. Department of Food Science Swedish University of Agricultural Sciences Uppsala, Sweden Fabio Armellini, M.D. Istituto di Clinica Medica Policlinico di Borgo Roma Università di Verona Verona, Italy Nils-Georg Asp, M.D. Professor Department of Food Chemistry Chemical Center University of Lund Lund, Sweden Katrine I. Baghurst, B.Sc., Ph.D. Consumer Science Program Commonwealth Scientific and Industrial Research Organisation (CSIRO) Health Sciences and Nutrition Division Adelaide, Australia Peter A. Baghurst, B.Ag.Sci., Ph.D., M.Sc. Consumer Science Program Commonwealth Scientific and Industrial Research Organisation (CSIRO) Health Sciences and Nutrition Division Adelaide, Australia John H. Cummings, M.D. Department of Molecular and Cellular Pathology Ninewells Medical School University of Dundee Dundee, Scotland Hans N. Englyst, Ph.D. Medical Research Council Southampton East Leigh, Hants., England Sharon E. Fleming, Ph.D. Professor College of Natural Resources Agricultural Experiment Station Department of Nutritional Sciences University of California Berkeley, California Hugh J. Freeman, M.D. Professor of Gastrointestinal Medicine Department of Medicine University of British Columbia Vancouver, British Columbia, Canada Wenche Frølich, D.Ph. Lindebergveien 39 JAR, Norway Ivan Furda, Ph.D. Furda and Associates, Inc. Wayzata, Minnesota Daniel D. Gallaher, Ph.D. Assistant Professor Department of Food Science and Nutrition University of Minnesota St. Paul, Minnesota 2387_FM_fm Page xi Sunday, May 6, 2001 6:18 PM Sherwood L. Gorbach, M.D. Department of Family Medicine and Community Health Tufts University School of Medicine Boston, Massachusetts Mary Beth Hall, Ph.D. Assistant Professor Department of Animal Science University of Florida Gainesville, Florida Barbara F. Harland, Ph.D., R.D. Professor Department of Nutritional Sciences College of Pharmacy Nursing and Allied Health Sciences Howard University Washington, D.C. John H. Himes, Ph.D., M.P.H. Professor and Director Nutrition Coordinating Center Division of Epidemiology School of Public Health University of Minnesota Minneapolis, Minnesota Peter J. Horvath, Ph.D. Department of Animal Science Cornell University Ithaca, New York Geoffrey J. Hudson, Ph.D. Cambridge, England David R. Jacobs, Jr., Ph.D. Division of Epidemiology School of Public Health University of Minnesota Minneapolis, Minnesota Mazda Jenab, Ph.D. Department of Nutritional Sciences University of Toronto Toronto, Ontario, Canada Alexandra Jenkins, R.D., C.D.E. Clinical Nutrition and Risk Factor Modification Center St. Michael’s Hospital Toronto, Ontario, Canada David J. A. Jenkins, M.D., Ph.D., D.Sc. Department of Nutritional Sciences University of Toronto Toronto, Ontario, Canada Julie M. Jones, Ph.D. Department of Family and Consumer Science College of Saint Catherine St. Paul, Minnesota Heinrich Kasper, M.D. Professor Department of Internal Medicine Medical University Clinic University of Würzburg Würzburg, Germany Joseph Keenan, M.D. Department of Family Practice and Community Health University of Minnesota Medical School Minneapolis, Minnesota Hanako Kobayashi, M.S. Department of Nutritional Science and Toxicology University of California Berkeley, California Yves Le Quintrec, M.D. Assistance Publique Hopitaux de Paris Service de Gastro-enterologie et Pathologie Digestive Postoperatoire Hospital Rothschild Paris, France Betty A. Lewis, Ph.D. Division of Nutritional Sciences Cornell University Ithaca, New York Leonard Marquart, Ph.D., R.D. Nutrition Research Department General Mills, Inc. Minneapolis, Minnesota (Harold E.) Gene Miller, Ph.D. Principal Scientist General Mills, Inc. Minneapolis, Minnesota MS SPHERA Foundation Los Altos. Australia Donald Oberleas. Ph. Diet and Health Department AFRC Institute of Food Research Norwich. Sweden . Schneeman. Director SPHERA Foundation Health Research and Studies Center Los Altos.P. B.S.D.. California Monica Spiller. Kentucky Sally J. Randles. Ph.Organic Chemistry Department of Chemistry Swedish University of Agricultural Sciences Uppsala. Ph. Former Head Nutrition. M. Record. M.D.A. Minnesota Barbara O.2387_FM_fm Page xii Sunday.D.D.S. Ph.D. D. New York Sally F.. T. New South Wales. Professor Institute of Nutrition and Food Science Chinese Academy of Preventive Medicine Beijing. Professor Emeritus . Head Cancer Prevention Research Program Fred Hutchinson Cancer Research Center Professor of Epidemiology University of Washington Seattle.D.D. Texas Mark A. Department of Nutrition Harvard School of Public Health Department of Pediatrics Division of Endocrinology Children’s Hospital Boston.Sc. Massachusetts Janet Pettit. Schakel.D.D. Spiller. England Gene A. Database Nutritionist Nutrition Coordinating Center Division of Epidemiology School of Public Health University of Minnesota Minneapolis. Techn. Oakenfull.D. Hydrocolloids Research Wahroonga. University of Minnesota School of Medicine Department of Family Practice and Community Health Minneapolis. 2001 6:18 PM Bunpei Mori. V. Potter.D. Database Nutritionist Nutrition Coordinating Center University of Minnesota Minneapolis. Dr. Southgate.S.D. Australia James B. Ph. California Zhi-Ping Shen. Medical Center and University of Kentucky Lexington. Minnesota Joel J. R. Pins. M. M. B. May 6. Washington Kim M.A. Ph. Minnesota John D. Pereira. Ph. Department of Animal Science Cornell University Ithaca. M. Professor Department of Life Science Toita Women’s College Tokyo. Professor Emeritus Texas Tech.H. Japan David G.C. Ph..S. Consumer Science Program Commonwealth Scientific and Industrial Research Organisation (CSIRO) Health Sciences and Nutrition Division Adelaide. Professor Department of Nutrition University of California Davis. China David A. Robertson. California Olof Theander. University Lubbock. Ph. S. Department of Nutritional Sciences University of Toronto Toronto. Massachusetts Vladimir Vuksan.2387_FM_fm Page xiii Sunday. Ph. Canada Eric Westerlund. Woods. Japan Thomas M.D. Ph. Ph..D.. Human Biochemistry Research Unit The South African Institute for Medical Research Johannesburg. P. B. Italy Alexander R. Sweden Keisuke Tsuji. Associate Professor Department of Family Medicine and Community Health Tufts University School of Medicine Boston. M.Sc. Canada Peter J. Chief of Laboratory The National Institute of Health and Nutrition Tokyo.D. Istituto di Clinica Medica Policlinico di Borgo Roma Università di Verona Verona. Professor Institute of Cancer Chinese Academy of Medical Sciences Beijing. Michael’s Hospital Toronto. Clinical Nutrition and Risk Factor Modification Center St. Canada Mauro Zamboni. Walker. B.D. Ph. 2001 6:18 PM Lilian Thompson.Ch. May 6.Sc. New York Margo N. China . Ph. Wolever.D. Ontario. Ph.D. Ontario. South Africa Su-Fang Zheng. M. Ontario.D. M.D Associate Professor in Organic Chemistry Department of Chemistry Swedish University of Agricultural Sciences Uppsala. Department of Nutritional Sciences University of Toronto Toronto. Professor Animal Nutrition Department of Animal Science Cornell University Ithaca. D. Van Soest..D. D.M. . ........... Burkitt and Gene A........................................................ T.............................1 Enzymatic Gravimetric Methods ....... Southgate Chapter 2............... Methods of Analysis for Dietary Fiber Chapter 3.8 Chitin and Chitosan — Special Class of Dietary Fiber............................3 Food Components That Behave as Dietary Fiber ......... Southgate and Gene A..... May 6......... Spiller Section 2....2387_FM_fm Page xv Sunday.........51 Nils-Georg Asp Chapter 3........................................................................................ Horvath ...................15 David A.................................2 Dietary Fiber Parts of Food Plants and Algae ................................ Overview Chapter 1............................... Definitions and Properties of Dietary Fiber Chapter 2..................... Southgate Chapter 2.............45 Ivan Furda Section 3............9 Gene A........................................... Southgate Chapter 2.....................19 David A................................7 Physical Chemistry of Dietary Fiber ......... Southgate Chapter 2...........11 David A...........................6 Glossary of Dietary Fiber Components.................. Spiller Chapter 2.........................................1 Definitions of Dietary Fiber.........................33 David Oakenfull Chapter 2.... Robertson and Peter J................23 David A................................................................... 2001 6:18 PM Contents Section 1..........3 Denis P............. T.....2 Detergent Analysis of Foods........ T..................4 Food Components Associated with Dietary Fiber ........................................................5 Polysaccharide Food Additives That Contribute to Dietary Fiber...... Spiller Chapter 2.... T........1 Dietary Fiber: From Early Hunter–Gatherers to the 1990s ........................... T........................63 James B................27 David A..... ............................ T.........................5 Correlations of Transit Time to a Critical Fecal Weight (CFW) and to Substances Associated with Dietary Fiber ................................3 Dietary Fiber Analysis as Non-Starch Polysaccharides (NSPs) ...8 Determination of the Saponin Content of Foods .............................133 Daniel D..................... Spiller and Monica Spiller Chapter 4.................. and Per Åman Chapter 3........................253 Gene A........................................................................83 David A....7 Newer Methods for Phytate Analysis .................................... 2001 6:18 PM Chapter 3. Cummings Chapter 4..............................6 The Crude Fiber Method ..................................3 Effects of Dietary Fiber on Vitamin Metabolism...... Gorbach ..............................257 Margo N... Gallaher and Barbara O........5 Determination of Total Dietary Fiber and Its Individual Components by the Uppsala Method ............87 Olof Theander............... Eric Westerlund.................113 Donald Oberleas and Barbara F.127 David Oakenfull and John D............................................................................................................161 Barbara F.....1 Effect of Dietary Fiber on Protein Digestibility and Utilization ............ Roger Andersson.....4 The Effect of Dietary Fiber on Fecal Weight and Composition............................................ Schneeman Chapter 4............2 Effects of Dietary Fiber and Phytate on the Homeostasis and Bioavailability of Minerals .. Harland and Donald Oberleas Chapter 4...................................173 Heinrich Kasper Chapter 4...2387_FM_fm Page xvi Sunday........................ Physiological and Metabolic Effects of Dietary Fiber Chapter 4..............6 Influences of Fiber on the Ecology of the Intestinal Flora...111 Ivan Furda Chapter 3..183 John H........................................ Southgate Chapter 3......................................... Woods and Sherwood L........ Harland Chapter 3........... May 6.......................67 Hans Englyst and Geoffrey Hudson Chapter 3............................................ Potter Section 4........4 The Southgate Method of Dietary Fiber Analysis ...... ...435 Hugh J.......9 The Source of Dietary Fiber Influences........2 Development of the Dietary Fiber Hypothesis of Diabetes Mellitus ..................................................................................... Van Soest Chapter 4.... Trowell Chapter 5... May 6...... Jenkins..317 Hugh J......... 2001 6:18 PM Chapter 4................ S. Jenkins Chapter 5........ Walker Chapter 5........8 Effects of Dietary Fiber on Digestive Enzymes............. Akanji..............4 Fiber in the Treatment of Hyperlipidemia..........................431 Hugh J.................................................... and Peter J.. and Kim M.................363 Alexander R..373 James W..321 Thomas M.......401 Alexandra L............... Gallaher Chapter 4..1 Disease Patterns in South Africa as Related to Dietary Fiber Intake............................... Anderson................................... Mary Beth Hall..........7 Interaction between Human Gut Bacteria and Dietary Fiber Substrates............. and David J......................... Short-Chain Fatty Acid Production and Concentrations in the Large Bowel .......... Lewis........5 Human Studies on Dietary Fiber and Colon Neoplasia........... Randles Chapter 5...... Dietary Fiber in the Prevention and Treatment of Disease Chapter 5.. Freeman Chapter 5......................... Wolever and David J......3 Treatment of Diabetes with High-Fiber Diets......... A........... Fleming Chapter 4....423 Hugh J... Schneeman and Daniel D.........................11 Effect of Dietary Fiber and Foods on Carbohydrate Metabolism ... Freeman ................................ Freeman Chapter 4........ P... Abayomi O...............287 Hanako Kobayashi and Sharon E...271 Betty A.......... Freeman Chapter 5........6 Fiber and Colonic Diverticulosis ..................................... Vladimir Vuksan...............2387_FM_fm Page xvii Sunday...................... A....................................7 Fiber and Inflammatory Bowel Diseases (Ulcerative Colitis and Crohn’s Disease).. Jenkins Section 5...........10 Effects of Dietary Fiber on Fecal and Intestinal Luminal Mutagens ..............277 Barbara O...369 Hugh C.......... .............. Thompson Section 7............................ Jacobs.................. Jacobs............ May 6...... and Chronic Diseases: Experimental Evidence and Possible Biologic Mechanisms ............................... David R....... Baghurst...... Peter A........461 Mark A........ and Sally J..........441 Keisuke Tsuji and Bunpei Mori Chapter 5. Definitions and Consumption Chapter 7.................453 Gene Miller Chapter 6..........5 Phytic Acid and Cancer ..... and Antioxidants........... Baghurst................................... Jr..........3 Dietary Fiber..1 Whole Grain.............8 Disease Patterns in Japan and Changes in Dietary Fiber (1930–1980) ..... Non-Starch Polysaccharide... Spiller Section 6........ David R.................................................. Cereal Fiber.........2 Whole Grains........................... 2001 6:18 PM Chapter 5.................. Pereira...............583 Katrine I.....9 Dietary Fiber Modification of Toxin............................. and Joseph M.......................... and Resistant Starch Intakes in Australia .... Effect of Whole Grains......................2 Patterns of Dietary Fiber Consumption in Humans to 1992 ... Joel J.............. Jr..or Carcinogen-Induced Effects on Intestinal and Mammary Tissues ....................................531 Mazda Jenab and Lilian U........................ Jones Chapter 7................... and Chronic Diseases: Epidemiologic Evidence........... Mark A.....................................1 Consumption of Dietary Fiber 1992–2000....... Cereal Fiber..553 Julie M........ Fiber............ and Joseph Keenan Chapter 6...567 Sheila Bingham Chapter 7. Pereira.................... Pins.....499 Wenche Frølich Chapter 6..... Cereal Fiber............593 Zhi-Ping Shen and Su-Fang Zheng . Record Chapter 7.....481 Joel J...... Keenan Chapter 6...................... and Phytic Acid on Health Chapter 6......................4 Bioavailability of Minerals from Cereals .... Leonard Marquart....................................4 Consumption of Dietary Fiber–Rich Foods in China ............. Len Marquart.... Pins...445 Hugh J.............................2387_FM_fm Page xviii Sunday.. Freeman and Gene A................3 Whole Grains.. ..........3 Dietary Fiber Content of Selected Foods by the Southgate Methods .........................................7 Phytate Contents of Foods ........................669 Gene A.........6 Fiber Consumption in Italy...663 Wenche Frølich Table A...1 Dietary Fiber Values for Common Foods .................... and Mauro Zamboni Appendix: Tables of Dietary Fiber and Associated Substances Content in Food Table A.................. May 6........605 Ottavio Bosello......615 Sally F.................................................... and John H..673 Barbara F.............4 Dietary Fiber Content of Cereals in Norway .................................................................. Himes Table A....2 Dry Matter............................................ Cellulose........................ and Lignin Content of Selected Foods ................ Crude Protein........ Fabio Armellini. Total Dietary Fiber................................................................................9 Plant Foods That Contain Significant Levels of Saponins and Their Estimated Saponin Content..........................................6 Comparison of Analyses of Dietary Fiber and Crude Fiber..........................................................................5 Crude Fiber Values of Typical Samples ................... Neutral Detergent Residue..... Spiller Table A.......... Soluble Fiber.683 David Oakenfull and John D................... Janet Pettit..........................2387_FM_fm Page xix Sunday.... T..............665 Ivan Furda Table A....................................... Hemicellulose.....685 .......5 Consumption of Dietary Fiber in France (1950–1981) ................. Potter Index... Southgate Table A....... Spiller Table A...659 David A. 2001 6:18 PM Chapter 7...................597 Yves Le Quintrec Chapter 7.........649 James B.................................................. Robertson Table A.....................8 Tartaric Acid Content of Foods .......................................................... Harland Table A............... Ash.......... Schakel.........681 Monica Spiller and Gene A.......................................................................................................... . 2001 5:55 PM SECTION 1 Overview .2387_Sect. 1 Div_fm Page 1 Sunday. May 6. 1 Div_fm Page 2 Sunday. 2001 5:55 PM . May 6.2387_Sect. and. whose name is eponymously commemorated in anatomical anomalies and surgical instruments. According to Kliks.1_fm Page 3 Sunday.1. this coprolite has been extremely valuable in the study of human diets of civilization dating back about 10.) by the Persian physician Hakim.S. Allinson in Britain extolled the virtues of whole grains in improving health by combating constipation. Spiller The decline in plant fiber consumption by humans over tens of thousands of years is shown in Figure 1. In the early 19th century.S. The convictions of the latter culminated in the development of the breakfast cereal industry. At the same time. stalks. physiologists. Cowgill and Anderson in the 1930s published well-controlled research that proved that “fiber” was responsible for the laxative action of wheat bran. John and Harvey Kellogg were extolling the virtues of whole grains in the U. May 6. McCarrison4 drew attention to the good health of tribesmen in North India. flowers. His first reaction was to remove the colon surgically. over the past 20.. who commented on the laxative action of outer layers of cereal grains. The history of dietary fiber and health in the first 50 years of the 20th century reveals occasional interest but very few scientific publications.1. In the 1920s. often monotypic diet in which the plant foods are primarily a few cereal grains. tubers. and physicians in those years. which he attributed to the consumption of whole grains little tampered with by modern technology.1 These specimens of coprolite showed a high consumption of fibrous plant food. minerals. In more recent history.50 © 2001 by CRC Press LLC 3 .1 Dietary Fiber: From Early Hunter–Gatherers to the 1990s Denis P.2387_ch1.00+$1. Fiber was relegated to being the Cinderella of nutrients. 2001 6:19 PM CHAPTER 1. plant-based regimen of greens. an observation repeated over 1000 years later (9th century A. and legumes. Somehow the concept that a group of substances practically undigestible by human GI enzymes could be important to health did not appeal to nutritionists. and all the other digestible nutrients. Even though the study of specimens of coprolite from lower Pleistocene humans has proved difficult. seeds. the concept that coarse foods of plant origin help to combat constipation goes back to Hippocrates in the 4th century B. Shakespeare referred to the action of cereal bran in his play Coriolanus in 1610. Burkitt and Gene A. and small amounts of animal products to a more limited. pollen. the author of this figure. Remember that these were the days of major discoveries in vitamins. roots. Graham2 and Burne3 in the U. in the late 19th century and at the dawn of the 20th century.000 years. In the same decade.C.D. the British surgeon Albuthnot Lane. recognized the “dangers” of stagnant fecal content in the colon. but fortunately he subsequently appreciated that the administration of bran 0-8493-2387-8/01/$0.000 years the human diet has changed from one based on a coarse. 1978.1_fm Page 4 Tuesday. Dietary Fiber: Current Developments of Importance to Health. In the years that followed. is the early history of fiber.) was altogether simpler. in Topics in Dietary Fiber Research. Walker of South Africa was one of the first to appreciate the properties of plant fibers and to study them in a truly scientific manner.. May 8. 182. some epidemiological correlations by Cleave in Britain (1956 to 1966) and Trowell in Africa. J.12 the Royal College of Physicians wrote Medical Aspects of Dietary Fiber. Dietary Fibers: Chemistry and Nutrition.9 in 1978 by Heaton. it is worthwhile to recall some of their titles.1. more books on specific topics began to appear: Spiller and McPherson-Kay published Medical Aspects of Dietary Fiber. reported his careful studies on the effectiveness of wheat bran in treating constipation and. As books are key steps in the growth of any field of science and as they become an intrinsic part of the history of science.. in the late 1940s. (From Kliks. 2001 2:54 PM 4 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.6 Reilly and Kirsner published Fiber Deficiency and Colonic Disorders.10 and in 1979 by Inglett and Falkenhag. Some of the pioneering books appeared in 1975 to 1976: Burkitt and Trowell published Refined Carbohydrate Foods and Disease.11 In the 1980s. In the late 1930s. This. A. a young British physician. M. we could say. and Amen. and equally effective.5 attributed protective effects to unrefined carbohydrates and “bulky” foods.8 They were followed in 1977 by Spiller and Amen. G.7 and Spiller and Amen published Fiber in Human Nutrition. Dimmock.2387_ch1. with his 1960 book.13 James and Theander wrote The Analysis of . Plenum Press. Eds. Other events took place in the 1970s and 1980s: the publication of a series of multiauthor books contributed to the clarification of the role of dietary fiber.1 The decline in fiber consumption by humans. Topics in Dietary Fiber Research. With permission. 3RD EDITION Figure 1. Non-Infectious Diseases in Africa. New York. safer. as dietary fiber became well established as a topic worthy of extensive research in human nutrition. R. Spiller.. These phytochemicals. two major symposia were also held in Washington at George Washington University. at the age of 85. 1985). and this is reflected in some of the new chapters in this edition.1) and there is less than precise use of terms in some publications. In 1981 and 1984. Too often fiber purification means major structural alterations. Hugh Trowell passed away in England in 1989. It is strange that some researchers remember him more for his Burkitt lymphoma than for his work with fiber. Such phytochemicals range from many antioxidants with a variety of health benefits to tartaric acid.C. too many to list here. (1977) that appears to us now as a milestone in the history of dietary fiber.15 and Trowell and Burkitt wrote Western Diseases: Their Emergence and Prevention. functions. During the last 20 years of his life. it would have been quite difficult to find sufficient scientific and medical data to publish a Handbook such as this. who had developed many animal models for fiber research and who was an author in the first edition of this Handbook. perhaps the most important development has been the tremendous amount of research on the hundreds of phytochemicals that are present in plant foods and how many of them are closely linked with fiber foods. What follows is a brief update since 1992. 2001 6:19 PM DIETARY FIBER: FROM EARLY HUNTER–GATHERERS TO THE 1990s 5 Dietary Fiber in Food. Since the first edition of this Handbook. D. In spite of all this growth. It has a great historical value and the insight that Denis Burkitt had in all his work.14 Vahouny and Kritchevsky wrote Dietary Fiber in Health and Disease. meetings that have led to important debates on definition.. Dietary fiber is so complex that it defies the reductionist scientist’s desire to isolate a pure substance and study it in humans or animals. including several in analytical and chemical aspects of dietary fiber. dietary fiber was the most important topic in his work. there are still many controversial aspects that will become evident to the reader of this Handbook. passed away in Washington. The proceedings were published in a special supplement of the American Journal of Clinical Nutrition17 in 1978. The National Institutes of Health sponsored a major international meeting in Washington. but we should also take time to set up proper experiments and to review the literature carefully. and others began to include dietary fiber as an integral part of their programs. The pioneering days were over and dietary fiber had become an established.C. there are still conflicting views as to how dietary fiber should be defined (see Chapter 2. physicians. D.K. and even the term fiber itself. While we can study fiber concentrates.16 Many more books followed in the late 1980s and early 1990s. important factor in human nutrition and medicine. after a lifelong dedication to medicine.2387_ch1. all working together with fiber. Fast growth is exciting. Notwithstanding all this progress and widespread interest by scientists. not the least of which is the definition of dietary fiber. the field of fiber research has also suffered the loss of two great pioneers in dietary fiber work. As often happens in science. This Handbook is proof of how rapidly the field of dietary fiber research is growing. And George Vahouny. It is interesting to realize that only 4 or 5 years before the publication of the first edition of this Handbook in 1986. San Diego (1981). Since 1992. and health effects of dietary fiber. May 6. work he carried on to his very last days. Many other meetings have taken place in recent years. I have left the preceding section unchanged as it was written with him. what was true is 1992 is still true now: there are still some controversial aspects of the effects of fiber that will become evident to the reader of this Handbook. Denis Burkitt passed away in England soon after the publication of the second edition.1_fm Page 5 Sunday. and while many of them have a valuable place in health . closely interact with fiber. from antioxidants to tartaric acid. The Royal Society of New Zealand held a major symposium18 in Palmerston North in 1982 which was reported in their Bulletin 20. and health professionals on every continent. Brighton (U. International congresses of nutrition in Brazil (1978). The complexity of dietary fiber and its tremendous interactions with other food components make its study a most difficult one indeed! Since 1978 many major meetings that focused on various aspects of dietary fiber have taken place. Spiller. L. Pitman Medical. Clin. Spiller. M. Fiber Deficiency and Colonic Disorders. Soc. Royal College of Physicians. E. Topics in Dietary Fiber Research. 1980. Western Diseases: Their Emergence and Prevention. R. Burkitt. Inglett. John Libbey.. 3RD EDITION maintenance. James. W. Graham. 8. Capen. MA.. H. and McPherson-Kay. W. 2. and Mehlman. May 6. 1980... O. 1921. and Bell. J. Plenum Press. 2001 6:19 PM 6 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 10. 1978. in Topics in Dietary Fiber Research. Plenum Press. A. New York. Barrington and Haswell. 1978. P. 18. Kent. J. 1978. Plenum Press. Dietary Fiber in Health and Disease. Non-Infectious Diseases in Africa. M. Lyon and Webb. Roth. Academic Press. New York. R. P. London.. P. 1975. Harvard University Press... A. Boston. Refined Carbohydrate Foods and Disease. LA.. 1829. 1982. 1983. and Theander. 1981. U. R.. Trowell. Nutr. P. B. G. K. G.. Haswell. 1981. 1978.. New York. Medical Aspects of Dietary Fiber. Am. 17. R.. C. N. REFERENCES 1 . Frowde. 9. J. G. Hodder and Stoughton.. H. Proceedings: symposium on the role of dietary fiber. J. Eds. 31S. and Burkitt. New York. 12. Proceedings: fiber in human and animal nutrition. New York. Reilly. Academic Press. J. R... Tunbridge Wells. The Analysis of Dietary Fiber in Food. T. Paleodietetics: a review of the role of dietary fiber in preagricultural human diets. G. W. the ultimate purification of fiber often implies physical–chemical changes that need to be carefully remembered when studies are carried out. The “Fiber Hypothesis” of Trowell and Burkitt is now beyond a hypothesis: it is a truth that can help people lead much healthier lives and prevent many chronic diseases. 1960. Fiber in Human Nutrition. Heaton. D. Plenum Press. New Orleans. S. 1979. G. London. C. 5.Z. V.. Bull. and Kritchevsky. 14. Kliks. 1840.. Studies in Deficiency Disease. Wallace. Dietary Fibers: Chemistry and Nutrition. S.1_fm Page 6 Sunday. and Amen. New York. A. G. Treatise on the Causes and Consequences of Constipation... Plenum Press. Trowell.2387_ch1. 3. Marcel Dekker.. Vahouny. D. Cambridge. G. Medical Aspects of Dietary Fiber. Spiller. 181. J. R. Plenum Press.. London. Eds. A. 4. 15. Dietary Fiber: Current Developments of Importance to Health. R. 16. and Amen. 1975. and Kirsner. Edward Arnold. Spiller. 13. and Trowell. New York. I. . Lectures on the Science of Human Life. Eds. McCarrison. D. 7. A. 11. New York. and Amen. 6. H.. Burne. New York.K. and Falkenhag. 1976. H... May 6. 2001 5:59 PM SECTION 2 Definitions and Properties of Dietary Fiber .2387_Section 2_fm Page 7 Sunday. 2387_Section 2_fm Page 8 Sunday. 2001 5:59 PM . May 6. A key contribution by Trowell was to emphasize the distinction between dietary fiber and crude fiber. Southgate considers this as a physiological and philosophical definition and feels that it is necessary to produce a definition that can be translated into purely analytical terms. Dietary Fiber (Furda) Chemical definition: the sum of the plant nonstarch polysaccharides and lignin. 0-8493-2387-8/01/$0. May 6. Dietary Fiber (Southgate) The sum of lignin and the polysaccharides that are not hydrolyzed by the endogenous secretions of the human digestive tract. Spiller Term recommended and author Definition Dietary Fiber (DF) (Trowell) Plant substances not digested by human digestive enzymes. Spiller) Same as dietary fiber but defined to include other plant substances that are undigested by human digestive enzymes. including plant cell wall substances (cellulose. Physiological definition: the remnant of plant foods resistant to hydrolysis by the alimentary enzymes of humans. Largely identical to undigested (unavailable) carbohydrates plus lignin. Dietary Fiber Complex (Trowell. The early definition by Trowell of “the remnants of plant cells resistant to hydrolysis by the alimentary enzymes of man” remains as a key definition. pectin.1 Definitions of Dietary Fiber Gene A.50 © 2001 by CRC Press LLC 9 . 2001 6:21 PM CHAPTER 2.2387_ch2. and undigestible cell wall proteins.00+$1. such as waxes. cutins. This author suggests a chemical definition based on the fact that the sum of lignin and the non-α glucan polysaccharides (nonstarch polysaccharides) is the best index of dietary fiber in the diet. and lignin) as well as intracellular polysaccharides such as gums and mucilages. hernicelluloses. even though it may better apply to the dietary fiber complex defined below. These are the substances that are normally associated with and concentrated around the plant cell wall.1_fm Page 9 Sunday. Nonstarch Polysaccharides (NSP) (Englyst) The carbohydrate plant cell wall material originally called dietary fiber less the lignin.2387_ch2. the fact that some of these carbohydates are digested by bacteria in the human intestine and produce fatty acids that are actually “available” makes the term “unavailable” ambiguous. It also implies that the material is present in plants only. Neutral Detergent Residues (NDR) (Van Soest) A residue after special digestion (see Section 3) with detergents.” It includes undigestible parts of the plant cell that are not included in plantix. Edible Fiber (Trowel. 3RD EDITION Plantix (Spiller) A term designed to represent dietary fiber as defined by Trowell but that avoids the word “fiber. These products may be of pharmaceutical importance. (3) polysaccharides that are not part of traditional foods and that are not digested by human digestive enzymes. Examples are purified pectin and purified cellulose. Crude fiber values include only variable portions of the cellulose. e. . (2) synthetic or partially synthetic polysaccharides not digested by human digestive enzymes.. Unavailable Carbohydrate (McCance and Lawrence) A classical term used in nutrition for many years to distinguish between available and nonavailable carbohydrate in humans. hemicellulose. Crude Fiber The remnants of plant material after extraction with acid and alkali. hemicelluloses. analyzed in such a way as to eliminate any other plant substance that may appear as fiber in other analytical methods. This is a modification of an earlier definition that included more than one substance.. The term should not be used with reference to dietary fiber. May 6. often acting in the same way as fiber. other groups of undigested (human enzyme–undigested) polysaccharides and related substances are added: (1) animal fibers not digested by human digestive enzymes. and lignin present in dietary fiber. Often called neutral detergent fiber (NDF). Purified Plant Fiber (Spiller) Any highly purified single polymer derived from plants that is not digested by human digestive enzymes but that may be digested by microorganisms in the human intestinal tract. again to avoid the word “fiber. Methods to determine NSP must be such that any resistant starch (see below) is not taken into account.g. In fact. such as waxes.1_fm Page 10 Sunday. aminopolysaccharides. e. 2001 6:21 PM 10 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Plantix Complex (Spiller) Suggested to replace dietary fiber complex. It includes the plant polysaccharides and lignin that are not digested by human digestive enzymes. undigestible proteins. which is not a carbohydrate. and lignin. Godding) An expanded definition of dietary fiber as given above. strictly speaking. and others. be of a fibrous nature. Old definition from the 19th century. As originally defined it included lignin. essentially the sum of cellulose. cellulose is the only truly fibrous component of the plant cell wall.” since many of the substances defined may not. Furthermore. methylcelluloses. Resistant Starch (RS) (Englyst) Starch that is not digested by human digestive enzymes and that reaches the colon.g. 1 Schematic representation of the major stages in the classical scheme for the fractionation of the plant cell wall that forms the basis for the nomenclature used to describe the components of the plant cell wall. T.50 © 2001 by CRC Press LLC 11 . the plant species. May 6. 2001 6:24 PM CHAPTER 2.00+$1.2_fm Page 11 Sunday. The composition of the matrix varies with the maturity of the plant tissues.4 The nomenclature used to describe the detailed composition of the plant cell wall is based on the classical schemes for fractionating the components. The essential features of this scheme are given in Figure 2. This is a network of cellulose fibers in a matrix of non-cellulosic polysaccharides.5 Plant Tissue Extract with Aqueous Alcohol Alcohol Insoluble Material Extract with Hot Water plus Chelating Agent Pectic Substances Extract with Alkali Hemicelluloses Insoluble in Alkali α-Cellulose Figure 2.1.2 The organization and detailed composition of the plant cell wall varies with the type of tissues. although highly lignified (woody) tissues are rejected during food preparation.2387_ch2. 0-8493-2387-8/01/$0. but the essential features are common to virtually all walls. Southgate The major portion of dietary fiber in foods is derived from the plant cell walls in foods.2.1 A wide range of plant organs and types of tissue is consumed in the human diet. and the major plant grouping.2 Dietary Fiber Parts of Food Plants and Algae David A.3.2. with arabinosyl and 4-0methylglucuronyl substituents. plant tissues are highly organized structures which confer physical properties on these materials when they are consumed in foods. in addition to the rhamnogalacturonans (true pectins). Although lignin is an aromatic polymer and therefore not a carbohydrate. which would include a spectrum of polysaccharides which range from those rich in uronic acids to those poor in uronic acids.and galactomannans. galactomannans. gluco. which included. arabinogalactans. it is covalently linked to the polysaccharides and was included in the original definition of dietary fiber (see Trowell et al. 3RD EDITION The plant tissue was extracted with aqueous alcohol and the insoluble material was extracted first. and in total dietary fiber (AOAC method) Non-Structural Components Gums and mucilages Water-soluble or dispersible Galactomannans. Where a mature plant tissue that contained substantial amounts of lignin was being fractionated. These polysaccharides are linked together in what can be described as a supramolecular structure which confers additional properties on the wall. and arabinoxylans. xyloglucans Cellulose (and lignin where present) Aromatic polymer. and xyloglucans. The material insoluble in the strongest alkali was designated the “α-cellulose. It is known that the fractions obtained in the classical fractionation schemes are arbitrary and that artifacts are often produced. Plant foods contain a range of water-soluble gums and mucilages. soluble fiber . with hot water.2_fm Page 12 Sunday.8). It important to recognize that the plant cell wall is a highly organized structure. soluble in dilute alkali Insoluble in alkali Insoluble in 12M H2SO4 Arabinoxylans.” which contained cellulose and lignin. arabinoxylans Hemicelluloses Insoluble in water. This gave a “pectic substances” fraction. arabinogalactans. wide range of branched and substituted galactans Included in NCP and NSP.2387_ch2. These have structures analogous to many cell wall components and are included in the non-cellulosic polysaccharides and the non-starch polysaccharides (NSP) of Englyst et al.1. 2001 6:24 PM 12 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. The water-insoluble material was then extracted with increasing strengths of alkali to give the so-called hemicellulose fraction. arabinogalactans. This often contained a chelating agent to disrupt the polysaccharides forming the intercellular middle lamellae.1 Dietary Fiber Components of Foods Classical Nomenclature Solubility Characteristics Classes of Polysaccharide Nomenclature Used in Dietary Fiber Literature Plant Cell Wall Components Pectic substances Water-soluble Rhamnogalacturonans. β-glucans. Albersheim7 proposed that the term “non-cellulosic polysaccharides” (NCP) should be used for this fraction. β-glucans.6 Furthermore. Although modern derivatives of this method 6 have refined the alkaline extraction stages to avoid some of the degradation from the alkali.2. which included a range of xylans. Table 2. it was necessary to oxidize the lignin before the hemicelluloses could be extracted.9 The major components of dietary fiber are summarized in Table 2. not merely a collection of polysaccharides.2. May 6. non-carbohydrate α-Cellulose Lignin Included in non-cellulosic polysaccharides (NCP) and non-starch polysaccharides (NSP). soluble fiber Included in NCP and NSP Included in NSP Lignin. D. Albersheim. 31. 2. Proposed scheme of analysis. 4. May 6.. R. B. 1. N. R. 1996. Sci. Glick. J. Spiller. and Amen. Southgate. 6. Food Agric. J. 1976. 8. The chemistry of dietary fiber. and O’Neill.. Leeds... 109.. 320. Gaillard. A... New York. Am. 1958. Analyst. G. New York. Wolever. T. D. high-performance chromatographic or spectrophotometric measurement of constituent sugars. Nutr. M. Biogenesis of the cell wall. Nutr. in Plant Biochemistry. 39.. and Waldron. in Fiber in Human Nutrition.. T. J. S. D.2387_ch2. D.. E. 1976.. J... E. Gassull. and Varner. Brett. E. Englyst. 22. A.2_fm Page 13 Sunday. H. Selvendran.. 1994. Plant glycoproteins. 7. 1511.. J. London. and Hudson. 1984. 5. Dieta. Ed. A.. R. Fibre in nutrition. Eds. R. Chapman and Hall. with gas-liquid chromatographic. 32... A. G. 3. 170. A. 25. 1987. A. Quigley. 9. 9. K. Lancet. Plenum Press. J. . Clin. Selvendran. H.. Trowell. R. Southgate. 2nd ed. and Jenkins.. John Wiley. Bonner. T. P. in Methods of Chemical Analysis. 967. Eds. M. A.. Southgate. M. R. Academic Press. Dietary fibre redefined. T. Bibl. 298. Determination of dietary fibre as non-starch polysaccharides. D... 1975. 119. A detailed summative analysis of the crude fibre and nitrogen-free extractives fraction of roughages.. M. 2001 6:24 PM DIETARY FIBER PARTS OF FOOD PLANTS AND ALGAE 13 REFERENCES 1. D. The plant cell wall as a source of dietary fiber: chemistry and structure.. Physiology and Biochemistry of Plant Cell Walls. A. C. 1965. 2_fm Page 14 Sunday.2387_ch2. 2001 6:24 PM . May 6. Some authors have argued that all the indigestible components of the diet should be considered part of dietary fiber.3_fm Page 15 Sunday. used as ingredients Contaminants Processed foods Natural and synthetic Insects and crustaceans Animal tissues 15 .1 although some proteins and fats in the human diet are also indigestible. inorganic constituents Crystal structure Physically enclosed Retrograded amylose Fructo-oligosaccharides. Southgate Dietary fiber exerts a wide range of physiological effects when consumed.1 Food Components That Share Indigestibility with Dietary Fiber Category Dietary Sources Indigestible components of plant cellular tissues Lignin Cutin Suberin Waxes Protein. Table 2.4 Table 2. waxes Dyes Pigments Chitin from exoskeleton Mucopolysaccharides.00+$1. modified starches.2 By convention. galactooligosaccharides Sugar alcohols Gums.3. and its complex nature is responsible for a range of physical and chemical properties that are responsible for these physiological effects.3. May 6. especially vegetables and fruits Added to processed foods Many processed foods. 2001 6:26 PM CHAPTER 2. where it provides a substrate for the microflora of the large intestine which can degrade many of the polysaccharides present. dietary fiber has been restricted to the indigestible polysaccharides and lignin. broken Many heat-processed foods Many plants. hair Resistant starch Indigestible carbohydrates Non-structural carbohydrates Non-assimilable components 0-8493-2387-8/01/$0.50 © 2001 by CRC Press LLC Occurrence Most mature plant tissues Epidermal tissues of plants Subepidermal tissues Surfaces of fruits Virtually all plant tissues Green banana Cereal grains.3 Food Components That Behave as Dietary Fiber David A. semi-synthetic polymers Hydrocarbon oils. One characteristic property is that dietary fiber is not hydrolyzed by the endogenous enzymes of the human gastrointestinal tract.1 It therefore escapes digestion in the small intestine and passes into the large intestine.1 lists various indigestible components of food other than dietary fiber.3. T.2387_ch2. there are a range of indigestible carbohydrates in foods. These carbohydrates are soluble in 80% v/v ethanol and therefore require additional stages to the NSP methods and total dietary fiber (AOAC) to measure them.9 HEAT-PRODUCED ARTIFACTS A fifth group contains degraded carbohydrates.8 NON-STRUCTURAL POLYSACCHARIDES The fourth group includes polysaccharides which are not alpha-linked glucans (and therefore indigestible) that are present in foods but are not part of cell wall structures. Most of these escape digestion in the small intestine either because they are intrinsically nondigestible or because the cell wall structures inhibit or prevent enzymatic attack. however. and two series of oligosaccharides — those based on fructose and the galacto. These include noncarbohydrate materials which are associated with or are integral parts of the plant cell wall structure. Many starch-rich foods. and inorganic constituents. stachyose. A few foods also contain starch grains with special crystalline structures that resist enzymatic hydrolysis. These may occur naturally in plant foods or may be food ingredients of additives that have been added to processed foods. are hydrolyzed to glucose very effectively by the brush-border enzymes of the small intestine. such as lignin. RESISTANT STARCH5 This latter effect is also seen with starches that are contained within cellular structures. 2001 6:26 PM 16 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.series.2387_ch2. suberin.series: raffinose. These three types of enzymatically resistant starch are often grouped as “resistant starch. cutin. the malto.6 INDIGESTIBLE CARBOHYDRATES7 In addition to the carbohydrates in the plant cell wall. These ingredients include semi-synthetic. These include fructan.3_fm Page 16 Sunday. May 6. The other major series of oligosaccharides in foods. 3RD EDITION INDIGESTIBLE COMPONENTS OF PLANT CELLULAR TISSUES In most human diets the plant cell wall material in the diet is the major source of indigestible polysaccharides.” which refers to resistance to hydrolysis under physiological conditions. usually in the form of carbohydrate–protein complexes formed during the processing of foodstuffs. These compounds are present at significant levels in many fruits and vegetables. waxes. protein. inulin. and a range of other polysaccharides are used as food additives. Processed foods and a few raw foods also contain sugar alcohols that are only partially absorbed. and verbascose. Some exudate gums are used as ingredients in confectionery. randomized polymers of glucose. contain retrograded amylose. usually to control or influence the physical properties of a food. . plants contain other substances that are indigestible. such as polydextrose. especially those which have been processed by heat treatment in moist conditions and allowed to cool. which forms an insoluble crystalline material that is resistant to enzymatic hydrolysis. . and Cummings. J. Institut National de la Recherche Agronomique. Eds. Southgate. M. E. D. H. Eur. A. 33. Trowell.. M. and hairs. D. Nutr. AIR3CT94-2203. European Air Concerted Action. Wolever. 1. M. 1980. R. A. J.11 REFERENCES 1. Dietary fibre redefined.. Dietary fiber and resistant starch. J.. 1987. A. S. Brussels. Voeding. 2). 2001 6:26 PM FOOD COMPONENTS THAT BEHAVE AS DIETARY FIBER 17 NON-ASSIMILABLE COMPONENTS OF FOODS Finally. Trowell. A. Cambridge. Stephen. D. Contract No. and Russell. M. 1969. T. Guillon. A. 1976. J. 284. Royal Society of Chemistry. Kingman. Gudmand-Hayer.. M. Gassull. D. 9.. N.2387_ch2. Southgate. 2. D. A. 230. 3. R. AGRF/0027.” the methods of determination and the current hypotheses for the explanation of its physiological action. Some critical observations in relation to “dietary fibre. Mechanism of action of dietary fibre in the human colon.. 11. A. The significance of protein as a component of dietary fiber. C. W.. E. Clin. Euresta. S. S33 (Suppl.. there is a range of non-assimilable components of food. H. Clin.... T.. C. which was not synonymous with indigestibility. Academic Press. New York.. 1995. 46. Englyst. Hollingsworth. European Union. Leeds. 4. Saunders. Am.. 6... Nature (London). D. 1980. Dietary Fibre Analysis. T. Englyst. Methodological aspects of the in vivo methods for measurement of starch digestibility. 1978. hydrocarbon oils. T. 199. 10.. Clin.. 1998. M.10 All these substances can be said to behave as dietary fiber in the sense that they are indigestible.. T. the exoskeleton of insects and crustacea. Profibre. 283. May 6. Am. mucopolysaccharides. F. and Jenkins. Classification and measurement of nutritionally important starch fractions. 5.3_fm Page 17 Sunday. H. J. M.. and Cummings. 960. H..1992. 873. and Cummings. F. Gum Technology in the Food Industry. Many authors believe that most of these components are not considered appropriate to include in dietary fiber because they include a heterogeneous collection of materials far removed from the original conceptual definition of dietary fiber.. 967. Nantes. Non-assimilable components of foods. 33.. A. A. 7.. 46. Nutr. Southgate. Functional Properties of Non-Digestible Carbohydrates. N. degraded connective tissue components. Lancet. et al. 1992. 8. Southgate. Eds. Glicksman. in Nutritional Problems in a Changing World. J. 1973. Nutr. and Betschart. A. Hellendoorn... . H. waxes and dyes. 2387_ch2. 2001 6:26 PM .3_fm Page 18 Sunday. May 6. especially roots and tubers Plant waxes External surfaces of many fruits Inorganic constituents Virtually all walls Main Structural Features Complex aromatic polymers formed from coumaryl.4. guaiacyl.4_fm Page 19 Sunday.50 © 2001 by CRC Press LLC 19 . whereas others appear to be chance inclusions. nevertheless. ketols. and polyphenolic substances). Ca.2387_ch2.4. and it is highly probable that they contribute to the protective effects of high-fiber diets. These phytochemicals are therefore consumed with dietary fiber components. glucosinolates. Some of these are integral to the cell wall structures.00+$1.1 Food Components Associated with Dietary Fiber Component Lignins Protein Cutin Suberin Sources Found in all vascular plants. Southgate INTRODUCTION Plant foods contain a wide range of substances and many of these are biologically active (for example. May 6. Table 2. however. 2001 6:27 PM CHAPTER 2. they modify the physical and chemical properties of the wall polysaccharides and thereby can be expected to modify the physiological properties of dietary fiber when eaten. vitamins. C26 acids and alcohols with phenolics and dicarboxylic acids Complex mixtures of hydrocarbons. T. and sinapyl alcohols Often rich in hydroxyproline and linked to polysaccharides Polymeric internal esters of C16 and C18 hydroxyaliphatic acids Polymeric material from C20.1 The non-carbohydrates are usually quantitatively minor components of the plant cell wall. major component of woody tissues and some seeds Present in all cell walls External surfaces of many fruits and leaves Sub-epidermal tissues of many plants.4 Food Components Associated with Dietary Fiber David A. coniferyl. Mg salts and silicates common The plant cell wall contains or is very closely associated with a range of non-carbohydrates.1).2 0-8493-2387-8/01/$0. ketones. K. phytoestrogens. etc. and still others are closely associated with the walls on the exterior surfaces of the plants. This section is. concerned with the components of plant foods that are intimately associated with dietary fiber (Table 2. 9 CUTIN. in the main. but these are specific for the species concerned. and this accounts for the very wide range of structures observed. AND PLANT WAXES1.5 In mature woody tissues the xylem walls are completely lignified. Cutin Many external surfaces of plants are covered with a waxy layer which includes a range of complex substances containing long-chain hydroxy aliphatic acids which form internal esters.2387_ch2. The cell wall proteins in some tissues appear to have an intrinsically lower digestibility. PROTEIN Some protein is found in virtually all cell walls and may form up to 10% of the wall in immature plants. and for this reason some authors believe that these proteins should be considered part of the dietary fiber complex.6 In the Van Soest and Wine method the susceptibility to oxidation by alkaline permanganate is the basis of the determination.7 Colorimetric procedures for some grass species have been described.5 These substances are complex lipid materials present in many plant tissues. SUBERIN. guaiacyl. and as lignin is highly resistant to bacterial and enzymatic attack.4 In many food plants.3 Lignin is the name given to a complex group of aromatic polymers formed by the condensation of aromatic alcohols. and sinapyl alcohols. especially in the developed world. Lignified tissues are present in the cell walls of the seed coats of cereals.4_fm Page 20 Sunday. it is more appropriate to consider lignin as an associated component. Thus. non-enzymatic. insolubility in 12M H2SO4 is the basis of the measurement of “Klason” lignin. Analytical procedures for lignin are often nonspecific and depend on the resistance of lignin to chemical attack. and lignified seeds are widely distributed in many fruits and vegetables. the lignification is limited to the spiral and annular bands in the walls of the xylem vessels. 3RD EDITION LIGNIN Although many authors consider the lignins (a wide range of structures are found in plants) as part of dietary fiber in a formal sense. This material is highly hydrophobic and intimately associated with the cell wall at the plant surface. As the plant matures the lignification spreads through the entire wall of the xylem and extends into supporting tissues. May 6. coniferyl. 2001 6:27 PM 20 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.5 Lignification produces a hydrophobic region within the wall. coumaryl. The cell wall proteins appear to play an important structural role and are in some tissues characteristically rich in hydroxyproline. lignified tissues from foods are recoverable virtually intact from fecal material. The proportion falls as the wall matures and proportionately more polysaccharides are deposited in the wall. . This condensation is. which are consumed relatively immature. Lignification of the plant cell wall is an infiltration process of the matrix where the lignin forms a three-dimensional structure within the polysaccharide matrix and the wall expands in volume during the process.8 The proteins are often covalently linked to polysaccharide side chains. because it is only by conventional usage that lignin is included in dietary fiber. Pears contain clumps of lignified cells within their flesh. These woody tissues are most frequently not consumed as foods. A. 3. Suberin is characteristically deposited in sub-epidermal layers in many plant tissues. 5..6 Suberin Suberin is also a complex mixture with a composition analogous to cutin which includes phenolic components. J. 31. and Betschart. Procedures and Some Applications). J.S.10 Calcium is also closely involved in the structure of the pectic components of the middle lamellae between cells and the integrity of plant tissues.. 4. H.. D.. Lancet. S. 2nd ed. H.. 575. The significance of protein as a component of dietary fiber. M. Eds.. 1976. Off. Plant glycoproteins. Gassull. 8. Plant Anatomy. Eagan Press. Department of Agriculture. A. and Waldron. J. T. Dietary fibre.4_fm Page 21 Sunday. and O’Neill. 780. C. U. 1977. Selvendran. Chapman and Hall. phenolic esters. Bonner. Van Soest. Saunders. Leeds. E. 1971. Basel. May 6. Suberinized tissues are also hydrophobic and resistant to degradation in the intestine and usually are analyzed with lignin. Forage Fiber Analysis (Apparatus Reagents. Cutting. 26. but it is also found in the skins of some fruits. MN. Loewus. 1996. Edward Arnold. New York.. Determination of lignin and cellulose in acid detergent fiber by oxidation with permanganate. Brett.. in Dietary Fiber in Health and Disease. Am.. 52. and Tanner. In some analytical schemes the cutin is analyzed with the lignin unless the material is extracted with a lipid solvent. E. A. D. D. 960. 2. R. Waxes Complex hydrocarbon waxes are found in many plants in very small quantities. esters. .. F. 379. and alcohols. M. Jones.. and Jenkins. Goering. A. J. Am. and Bonfield... J. REFERENCES 1.. 1982.. Academic Press. R. H. T. K. and Van Soest. particularly in underground organs such as roots and tubers. Physiology and Biochemistry of Plant Cell Walls. London.. and Wine. J. 1980. M.. London. A.C. These waxes coat external surfaces of many fruits with a hydrophobic layer. Kritchevsky.Clin. P. 7. 1978. In some plants the inorganic material apparently has structural significance. Chem. R. D. 967. 9.. A. Eds. 1995. Agricultural Handbook No. Southgate. St. Eds. P. 1. 1970. part 2. The waxes are mixtures of ketones. Springer-Verlag. 33. this being especially true for silica and silicates. Wolever.. G. R.. H... Other inclusions of potassium and magnesium salts with phosphates and oxalates are quite common. their degradation in the intestine appears to be limited. Mineral components of plant cell walls..2387_ch2. Nutr. in Plant Carbohydrates. and Varner. T. Analyt. Clin. L. 58. K. J. A. M. Plant Biochemistry... Paul. 1968. Experiment and Interpretation. D. P. INORGANIC MATERIALS Most plant cell walls contain inorganic material. The structure of dietary fibre. 2001 6:27 PM FOOD COMPONENTS ASSOCIATED WITH DIETARY FIBER 21 Cutinized tissues are resistant to hydrolysis and degradation in the intestine and frequently can be recovered from fecal material. A. Trowell. 6. C. Although these materials are hydrolyzed by lipases in the presence of bile salts. R. A. W. 594. 2nd ed. Assoc. Nutr. Washington. Southgate. 10.. The detergent fiber procedures of Van Soest provide an extension which provides a method for measurement of cutin. 2001 6:27 PM .2387_ch2. May 6.4_fm Page 22 Sunday. therefore. Southgate A variety of polysaccharides are added to processed foods. May 6.3 Many have been.2 A few are prepared biosynthetically using microorganisms.2387_ch2. not hydrolyzed by mammalian digestive enzymes. 0-8493-2387-8/01/$0. They are added to foods to modify or control the physical properties of foods. They fall into a number of distinct categories.e. this produces water-dispersible celluloses which are used as fillers. and still are. 2001 6:29 PM CHAPTER 2. many of them have been extensively modified chemically to enhance specific properties or to reduce undesirable characteristics. but the majority are derived from plant polysaccharides and many from cell wall polysaccharides. and amidated pectins are used in low-sugar jams and as emulsifiers and stabilizers. Amidation also improves the gelling capacity. T. Cellulose A relatively simple modification involves the reduction in the chain lengths of the polymers by ball milling. they fall within the most commonly used definition of dietary fiber.00+$1.. used as models for cell wall components of dietary fiber in experimental physiological studies.50 © 2001 by CRC Press LLC 23 . While none are synthetic in the true sense.5 Polysaccharide Food Additives That Contribute to Dietary Fiber David A.5_fm Page 23 Sunday. Chemical modification involves the introduction of methyl and methoxyl groups which create a range of dispersible compounds which are used as emulsifiers and stabilizers. The polysaccharides all share one common structural feature: they do not contain α-glucosidic links (i.1. MODIFIED CELL WALL POLYSACCHARIDES These native structures have been chemically modified to improve their solubility or capacity to form gels. For this reason. Pectins The additives are selected from high-methoxyl varieties which have higher gelling properties. they are nonstarch polysaccharides) and are. are used in many milk products because of the ability to associate with milk proteins.2387_ch2. 2001 6:29 PM 24 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. guar and carob (locust) bean gums which are used as thickeners in soups and other foods. extracted from Irish moss and other algae.5%. A number of randomized glucose polymers have been prepared which are resistant to small intestinal digestion. . Fructan is an indigestible carbohydrate and contributes substrate to the large intestinal flora. These polysaccharides make a minor contribution to the total intake of dietary fiber5 but have been fed at much higher levels of intake in experimental studies. gum Arabic and gum tragacanth — are used as additives and food ingredients. Agar is used as a thickening agent in a wide range of foods. The most widely used gums are the galactomannan gums. May 6. ALGAL POLYSACCHARIDES Algae provide a range of polysaccharides used in processed foods. INDIGESTIBLE FRUCTOSE POLYMERS AND OLIGOSACCHARIDES Many vegetable foods such as Jerusalem artichokes and chicory contain the fructan inulin. Carrageenans. 3RD EDITION Modified Starches The chemical modifications include the introduction of phosphate groups or isopropyl groups and a variety of other substituents primarily aimed at preventing retrogradation in heat-processed foods where the introduced groups prevent association of the amylose chains. Modern extraction techniques have made inulin available as an ingredient. SEMISYNTHETIC POLYMERS These polymers include xanthan gum produced biosynthetically and are used as gelling agents. are also used as thickening agents.5. extracted from brown algae and modified alginates. Alginates. Some exudate gums and some randomized glucose polymers are used as ingredients at higher concentrations.4 A range of fructose oligosaccharides which are also indigestible is available as ingredients with claimed probiotic properties. Table 2.1 summarizes the major types of polysaccharides in use. and polydextrose is in use as an ingredient in “low-calorie” yogurts and similar products. In most foods the concentration of the additives is less than 1% by weight and usually less than 0. GUMS Several exudate gums — for example.5_fm Page 24 Sunday. Selective simulation of bifidobacteria in the human colon by oligofructose and iulin. A. A. Leeds. in PflanzenfasenBallastoffe in der menschlichen Ernahrung. tragacanth Complex heteropolysaccharides Guar. G. 967. with terminal pyruvic acid and acetyl groups Polydextrose is an Thermally randomized polymers example REFERENCES 1. Gastroenterology. and Jenkins. Ed. Dietary fibre redefined.. 108. M. X. methoxyl Both ether and ester derivatives are used Phosphate. D. 1998. and Cummings. D. Wirths. M. polymers sulfated galactose. 3. T. Lancet.. Institut National de la Recherche Agronomique. M. 975.2387_ch2. 5. New York. H. A. F. Academic Press... A. Southgate. 1969. Stuttgart. B. Gum Technology in the Food Industry. Nantes. 1.. S. Thieme Verlag. R.. . 1976. and sulfated anhydro-galactose Xanthan gum Backbone of glucose with alternating trisaccharide side chains of mannose and glucuronic acid.5_fm Page 25 Sunday.. E. isopropyl Cross-linked to prevent association between amylose chains Inulin Relatively low molecular weights Range of polymers Range of oligosaccharides Arabic.. Functional Properties of Non-Digestible Carbohydrates. J. 2001 6:29 PM POLYSACCHARIDE FOOD ADDITIVES THAT CONTRIBUTE TO DIETARY FIBER Table 2. Wang. 1980. 1995. Trowell. 4. Wolever. carob (locust) bean Variable proportions of glucose substituents Two major polymers Linear polymer of D-galactose and L-anhydrogalactose Salts of copolymers Salts of D-mannuronic and L-gluronic acid Range of different Polymers of anhydro-galactose. 76. Gibson...1 25 Polysaccharide Food Additives That Contribute to Dietary Fiber Major Category Modified cell wall components Polysaccharides Major Types Used Pectins High methoxyl Cellulose Modified starches Cross-linked Indigestible fructose polymers Gums Fructans Fructo-oligosaccharides Exudate gums Galactomannan gums Algal polysaccharides Agar Alginates Carrageenans Semi-synthetic polymers Bacterial synthesis Randomized glucans Main Structural Features Methoxyl esters of rhamnogalacturonans Amidated Amidated rhamnogalacturonans Methyl. 2. Glicksman. H. Guillon. Eds.5. Rottka. Gassull. W.. Beatty. Aufnahme an Hydrocolloiden in der Bundersrepublick Deutschland. May 6. H.... et al.. R.. May 6. 2001 6:29 PM .2387_ch2.5_fm Page 26 Sunday. Arabinans — Polysaccharides that give L-arabinose on hydrolysis. and Kay. which can be extracted from certain dried seaweeds. The chief use. May 6. Commercial algin is sodium alginate: it is slowly soluble in water. A. cetyl trimethylammonium bromide (CTAB). it is measured as the residue after extracting the food with a hot dilute sulfuric acid solution of the detergent. It is not hydrolyzed by mammalian digestive enzymes and is. are associated with pectin. Spiller. A.50 © 2001 by CRC Press LLC 27 . Agarose. Algal polysaccharides — The extract from the tissues of algae. M. from which it can be extracted with hot water.. Eds. Spiller Acid detergent fiber (ADF) — The cellulose plus lignin in a sample. Agar — A mixture of polysaccharides occurring as the cell wall constituents of certain red marine algae Rhodophytaceae. however. The water-soluble arabinogalactan of larch has received considerable study.00+$1. T. They are not hydrolyzed by the mammalian digestive enzymes and are therefore part of dietary fiber.. part of dietary fiber.6-anhydro-L-galactose and D-galactose as the repetitive unit.6 Glossary of Dietary Fiber Components* David A. Southgate and Gene A.2. Alginic acid — Water-insoluble algal polysaccharide. Alginic acid is not hydrolyzed by mammalian digestive enzymes and is. and precipitated by the addition of an acid. Gelidium sp. They are present in wood cellulose. therefore.5%. the main constituent. part of dietary fiber. since it is undigested by almost all of them. 0-8493-2387-8/01/$0. and have been isolated from the pectic substances of mustard seed and sugar beet. R.. New York. See Chapter 3. Medical Aspects of Dietary Fiber. is a neutral polysaccharide containing 3. polymannuronic acid. Agaropectin is a minor constituent polysaccharide and contains carboxyl and sulfate groups. T.2387_ch2.6_fm Page 27 Sunday. it is a laxative. polymers of this type are widely distributed. It is used as a stabilizer for ice cream and other food products.. Alginates — Algal polysaccharides not hydrolyzed by mammalian digestive enzymes and. 1980. as a water-soluble alginate with aqueous alkali metal hydroxides or carbonates. Plenum Press. With permission. divided into two groups: (1) reserve polysaccharides which are water soluble and (2) structural polysaccharides which are not. forming an extremely viscous solution. therefore. Arabinogalactans — Substituted galactans that form part of the hemicellulose complex in many tissues. is as a solid medium for cultivating microorganisms. e. D. * Adapted from Southgate.g. 2001 6:30 PM CHAPTER 2. therefore. It gels on cooling at a concentration as low as 0.. part of dietary fiber when used as a thickening agent in foods. G. Although most emphasis has been given to arabinogalactans from woody tissues. 6_fm Page 28 Sunday. cellulose is therefore part of dietary fiber. The ability of carrageenan to react with milk protein has led to its widespread use in preparations containing milk and chocolate. but with other residues also present. 3RD EDITION Arabinoxylans — Have a main chain composed of (1 → 4) β-D-xylopyranosyl units with an occasional branching in some preparations.. see also Chapter 2. Hardwood glucomannans appear to contain no galactose and are relatively insoluble. The method recovers 50 to 80% of cellulose. Ulceration of the cecum of both rats and guinea pigs has been demonstrated when carrageenan was added to the diet. It has been shown to be a potent cholesterol-lowering agent but. and often from the species Chrondus crispus (carrageenan. Cellulose — Best known.or double-residue side chains. and this explains its ability to increase fecal weight when added to the diet. di-. Galactans — Polysaccharides which. gums. The presence of side chains tends to make these polysaccharides more soluble in water. but the glucomannans from gymnosperms have galactose side chains and a higher mannose-to-glucose ratio (3:1). although they are uncommon in woody tissues. they have been isolated from the husks of many grains. on hydrolysis. and the ending -an signifying a polymer. give galactose. Glucomannans — Appear to be linear polymers with both mannose and glucose in the chain. from glycose. and 20% of hemicellulose. i. They occur in wood and in many algae. possibly because the side chains prevent the formation of intermolecular hydrogen bonding. Guar gum (guaram) is a representative example. Irish moss). pectins. Glucomannans are part of the hemicellulose fraction of the plant cell wall. Their resistance to digestion means they appear in the feces and may constitute a large proportion of the fecal fat. Cuticular substances are extremely resistant to digestion and in turn are thought to impair the digestibility of the other cell wall constituents. Crude fiber — Residue left after boiling the defatted food in dilute alkali and then in dilute acid. unlike almost any other plant polysaccharide. in varying proportions. Glucuronoxylans are found in the hemicellulose fraction of all land plants and most plant organs. and only truly fibrous component of the plant cell wall. Inconsistent results are obtained. hemicellulose. and lignin. They are found in the hemicellulose section of the cell wall and form the storage polysaccharides in many temperateclimate grasses. containing side chains of 4-O-methyl-α-D-glucopyranosyluronic acid and. tri-.2387_ch2. it is a lipid component of the waterproof covering and cuticle on the outer cellulose wall of plants. unmethylated D-glucuronic acid. Cutins may account for a substantial part of the increased fecal fat seen in subjects on a high cereal-fiber diet. Glucoronoxylans — Have a main “backbone” chain of (1 → 4) linked β-D-xylopyranosyl residues.4 g water per gram of cellulose).e. May 6. Arabinoxylans are widely distributed in the cell walls of many materials. Galactomannans — Polysaccharides that have both galactose and mannose in the chain. 2001 6:30 PM 28 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Dietary fiber — Includes all the polymers of plants that cannot be digested by the endogenous secretions of the human digestive tract. Cutin — A complex polymer of mono-. and it should not be used as a method for the determination of dietary fiber. Carrageenan — An algal polysaccharide chiefly composed of polymerized sulfated D-galactopyranose units. in some annual plants. . most widely distributed.4:1. It is a polymer of glucose and the glucoside linkage is β. and polyhydroxy fatty acids. has an adverse effect on the gut. The ratio of mannose to glucose is between 1:1 and 2. cellulose. mucilages. Cellulose also has the property of taking up water (0. The most important galactan is agar. The β-linkages in cellulose are not hydrolyzed by the enzymes present in man. 10 to 50% of lignin. It is the dried extract from certain red marine algae Rhodophytaceae. Glucofructans — Linear polymers with both fructose and glucose in the chain. a simple sugar. Galactomannans are part of the hemicellulose fraction of the plant cell wall. Glycan — Generic name for a polysaccharide.1. The arabinose is present as single. tri. lignin is usually classified as part of dietary fiber. Noncellulosic polysaccharides — Another term for hemicelluloses. Together with pectin. Structurally. These are not part of the cell wall structure but are generally indigestible and are thus considered a part of dietary fiber. They are found mixed with the endosperm or storage polysaccharides or in special cells in the seedcoat. The largest group consists of pentosans such as the xylans and arabinoxylans. each containing several different sugar molecules and uronic acid groups. three. They seem to be storage polysaccharides. Mucilages — Polysaccharides usually containing galactose. and mannose are all hexoses. galactose. and therefore is the structure between adjacent cell walls. Homoglycan — Polysaccharide containing only one type of sugar unit and hence on hydrolysis giving only one monosaccharide type.. a second group consists of hexose polymers such as the galactans. a leguminous vegetable cultivated in India for animal feeds. They are soluble in water to give very viscous colloidal solutions.g. The true plant gums. Middle lamella — Develops from the cell plate that forms between the daughter nuclei of the plant cell wall and extends to meet the existing wall.6_fm Page 29 Sunday. Guar gum (guaran) — A neutral polysaccharide. Since it is virtually indigestible. It is a commercial source of vanillin and other aromatic chemicals. a glycuronan. the hemicelluloses form the matrix of the plant cell wall in which are enmeshed cellulose fibers. it occurs in woody plant tissues. gum acacia and gum tragacanth. See Chapter 3. Neutral detergent fiber (NDF) — That part of food remaining after extraction with a hot neutral solution of the detergent sodium lauryl sulfate. and often xylose and arabinose. They have prefixes of di-. In small amounts it finds widespread use in the food and pharmaceutical industries as a thickener and stabilizer in. 2001 6:30 PM GLOSSARY OF DIETARY FIBER COMPONENTS 29 Glycuronans — Generic name for the polymers of uronic acids. It appears to be rich in galacturonans. and are insoluble in organic solvents. which are characteristically part of the pectic substances. Hemicelluloses — A wide variety of polysaccharide polymers. and so on to indicate the number of different types of sugar residues. Lignin — An aromatic polymer of molecular weight of about 10. a D-galacto-D-mannan. Gum (exudates and seed gums) — Complex polysaccharides. Glucose. found in the hemicellulose fraction from many cell walls. fructose.2. starch and cellulose. e. being obtained as slimy. which include all the matrix polysaccharides from the cell wall. Mannans — Polysaccharides made up of mannose units. galacturonan is a polymer of galacturonic acid and is therefore. or more monosaccharides. Hemicelluloses are those polymers extractable from plants by cold aqueous alkali. Heteroglycans — Polysaccharides that hydrolyze to two. for example. at least 250 of which are known. that is isolated from the ground endosperm of guar seed. The method for determining NDF was designed to divide the dry matter of feeds very nearly into those constituents which are nutritionally available for the normal digestive process and those which depend on microbial fermentation for their availability.. May 6.000 based on coniferyl and sinapyl alcohols.2387_ch2. The hemicelluloses are not digested in the small intestine but are broken down by microorganisms in the colon more readily than cellulose. colloidal solutions. The most abundant polysaccharides are of this type. sometimes called mucilages. The acidic hemicelluloses which contain galacturonic acid or glucuronic acid form a third group of hemicelluloses. They are not precursors of cellulose and have no part in cellulose biosynthesis but represent a distinct and separate group of plant polysaccharides. they resemble the hemicelluloses and are water soluble. e. galacturonic acid residues. salad dressing and ice cream. It is a measure of the cell wall constituents of vegetable foodstuffs. as well as in nonfood items such as toothpaste. They retain water and so protect the seed against desiccation. other than cellulose. . are the dried exudates from various plants obtained when the bark is cut or the plant is otherwise injured. Hexoses — Monosaccharides with each molecule containing six carbon atoms.g.. Guar and locust bean gums are examples of gums derived from seeds. Water-holding capacity — Amount of water that can be taken up by unit weight of dry fiber to the point at which no free water remains. They are characteristically rich in galacturonic acid and are galacturonans with a variable degree of methyl esterification. Protopectin — Term applied to the water-insoluble parent pectic substance which occurs in plants. the principal element present is often silicon. but there is no such relationship between lignin content and water-holding capacity. Pentose sugars most commonly present in human foods are L-arabinose and D-xylose. tri-. and which upon hydrolysis yields pectinic acid. It changes from an insoluble material in the unripe fruit to a much more water-soluble substance in the ripe fruit. amides. Silica has the capacity to impair the digestibility of cell wall materials. considered by the authors to be a better definition than fiber. Pectic substances — Mixtures of acidic and neutral polysaccharides that can be extracted with water from plant tissues. May 6. Pectinic acid — Groups of pectins in which only a portion of the acidic groupings are methylated. when present as glycosides. Its ability to form gels and its ion binding capacity may be important in human nutrition. The ash content of the cell wall. It is a plant lipid that cannot be extracted with a simple solvent but needs saponification before extraction. usually in the aerial part of the plant. 2001 6:30 PM 30 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. They are found in about half the known plant polysaccharides. behave like simple hydrocarboxylic acids.6_fm Page 30 Sunday. The fibrils are surrounded by an amorphous matrix of hemicellulose. Uronic acids — Present in the pectic substances and the hemicellulose portion of the plant cell wall. often acting in the same way as fiber. Dietetically. may be as high as 10%. the most common being D-galacturonic and D-glucuronic acids. Pectin is found in the primary cell wall and intracellular layer. forming metal salts. Uronic acids are derived from sugars by oxidation of the terminal −CH2OH to −COOH and. Sclerenchyma — Tissue forming the hard parts of plants such as nutshell or seedcoat. Pectin — General term designating those water-soluble pectinic acids of varying methyl ester content and degree of neutralization which are capable of forming gels with water and acid under suitable conditions. Pentoses are present in small amounts in all cell walls whether animal. either as a continuous layer or as localized thickenings or bands. 3RD EDITION Nonstarch polysaccharides (NSP) — A term suggested by Englyst and co-workers for the carbohydrate plant cell wall material originally called dietary fiber less the lignin. Silica — Deposited in the plant cell wall. or bacterial. the silica content of the soil. Plantix — A term coined from “plant” and “matrix” to replace dietary fiber to avoid the uncertain and diversified meaning of the term fiber. Oligosaccharides — Collective term for di-.2387_ch2. which are widely distributed in the polysaccharides in plants. of this. Also see pectic substances. and the maturity of the plant. . A close relationship exists between acid detergent fiber content of vegetable dietary fiber and water-holding capacity. Suberin — A cutin-like substance found in cork. Secondary cell wall — Polysaccharide in nature and apparently amorphous. Primary plant cell wall — The cellulose fibers of the primary cell wall are laid down in a random network on the middle lamella. The amount varies according to the species. It includes the same polymers found in dietary fiber from plants. the five-carbon sugars are of little importance as a source of energy for the body. particularly of wheat. Resistant starch — Starch that is not digested by human digestive enzymes and that reaches the colon. It is formed after the cell has reached maturity and is laid down inside the primary cell wall. processed foods may contain oligosaccharides with up to 9 residue. and alkyl and methyl esters. Pentoses — Monosaccharides with each molecule containing five carbon atoms. plant. and tetrasaccharides. M.. Academic Press.. Reinhold. and Kay. F. mucilages. M.. Spiller. the walls of xylem vessels are completely lignified. in Medical Aspects of Dietary Fiber. Chemistry and Biochemistry of Herbage. and Montgomery. Y.. A. R. arabinose and 4-O-methyl glucuronic acid are the most usual substituents. 1976. usually made up of cells with lignified walls. A few D-xylans are neutral molecules containing D-xylose residues only. and Kay. Budavari. 9th ed. W. 3. and Fertig. 2001 6:30 PM GLOSSARY OF DIETARY FIBER COMPONENTS 31 Water-soluble fraction — Fraction of dietary fiber soluble in water. G.. New York. 2. Spiller. M. in less mature tissues. Plenum Press. 1973. N.. REFERENCES 1. The Merck Index. Windholz. May 6. The Chemistry of Plant Gums and Mucilages and Some Related Polysaccharides. NJ. New York.. R. Eds. gums. R.. Butler. 1. 4.2387_ch2.. Smith. W.. M. White... New York.6_fm Page 31 Sunday. A. S. T. R. Eds. L. Eds. Glossary.. M. A. Southgate. it includes pectic substances. G. D. In mature woody tissues. Merck. Rahway... 1959. Xylans — Groups of polymers having a main chain of (1 → 4) β-D-xylopyranosyl residues. the lignification is partial and localized. Xylans are found in the hemicellulose portion of the plant cell wall in all land plants and in most plant organs. Xylem — Water-conducting elements of plant tissues. . and Bailey. Vol. Stroumtsos. 1980. and some polysaccharide food additives. G. 6_fm Page 32 Sunday.2387_ch2. May 6. 2001 6:30 PM . these materials can greatly enhance the consumer appeal of otherwise unattractive products.7_fm Page 33 Sunday. There are already a number of polysaccharide-based products available which their manufacturers claim can be used to formulate low-fat versions of traditionally high-fat products such as ice cream (see also. Although not usually added to foods at significantly high levels.) Most people. May 6. Through subtle adjustment to formulation and processing methods. these additives significantly boost our intake of dietary fiber. For example. for example.7 Physical Chemistry of Dietary Fiber David Oakenfull INTRODUCTION As explained in other parts of this Handbook. consumption of fiber-rich foods would have increased by far more than it actually has in response to fiber’s clear health benefits. the popularity of these products (reflected by sales in supermarkets) was inversely proportional to the comminution energy. Ward 2). This chapter outlines how physical chemistry impacts with chemical composition and influences the many different roles of dietary fiber at the various stages in its passage through the gut — from its role in influencing what we eat to its role in easing defecation. dietary fiber is a complex mixture of polysaccharides with many different functions and activities as it passes through the gastrointestinal tract. Many of these functions and activities depend on its physical chemistry. (If this were not so. 2001 6:31 PM CHAPTER 2. A positive effect of fiber on food preference can be seen in the use of non-starch polysaccharides as texture-modifying agents by the food industry. The remarkable ability of polysaccharides to thicken or gel aqueous solutions is exploited in numerous food products ranging from dessert jellies to oil-free salad dressings. FOOD PREFERENCES Dietary fiber often has a negative impact on food preference.1 Apparently. Further. we prefer foods that require little chewing. as shown in Figure 2. a great deal of attention is currently being paid in the use of polysaccharides to mimic the texture of fat.50 © 2001 by CRC Press LLC 33 . One reason for this may be that fiber-rich foods often require considerable effort to chew. for example.1. A study comparing breakfast cereals suggested that the energy required to break the product down into small pieces (the comminution energy) increases with its dietary fiber content.00+$1. prefer white bread to wholemeal bread and white rice to brown rice.7. 0-8493-2387-8/01/$0.2387_ch2. Nonetheless. decreased spontaneous energy intake in a group of 12 women.2) shows no correlation (the correlation coefficient is –0.20 Weetbix Nutri-Grain 0. because energy density has a strong effect on satiety8 and may well swamp any effect of fiber on SI.7_fm Page 34 Sunday. and the SI for all other foods is based on this standard. several investigators have speculated that the bulk associated with high-fiber foods induces a feeling of satiety and thus reduces meal size and food intake. Because distension of the stomach is believed to be one of the signals to stop eating. Some workers have found that fiber decreases energy intake3. This is perhaps not surprising.7. A plot of satiety index against dietary fiber content for a broad range of different foods (Figure 2. influences are more or less equal — and it seems that solubility and viscosity are critical factors.5. A supplement of psyllium gum. Soluble fiber reduced appetite a longer time after eating than insoluble fibers. 2001 6:31 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.11 Furthermore.). conflicting results have been reported. which is rich in soluble fiber. the high-satiety foods tend to have bulky. more dominant. others have found that it has no effect. Experimentally.00 Shredded Wheat Cornflakes 0 5 10 15 20 25 1/(Comminution Energy) Figure 2. though. May 6. White bread is given a score of 100.6 A numerical measure of the satiating effects of different foods has recently been developed called the satiety index (SI). SATIETY How Dietary Fiber Influences Satiety Not surprisingly.2387_ch2.40 Rice Bubbles All-Bran Froot-Loops Puffed Wheat 0.10 Studies in experimental animals and in humans have suggested that viscous polysaccharides can slow the rate of gastric emptying.80 Special-K 0. consumption of fiber in a meal appears to reduce energy ingested at a subsequent meal.9 Fiber appears to influence satiety when other. whereas wheat bran had no effect.12 Blundell and his colleagues13 have recently found that soluble and insoluble fiber behave differently.4. physical factors are also significant in the influence of dietary fiber on satiety and hence energy intake. but there was no difference between . fibrous.0864. or crunchy textures which make them relatively more difficult to chew or swallow.7 The SI compares how full (on average) different foods made a group of volunteers feel for the same intake of energy.1 Relative sales in Australia of breakfast cereals plotted against the reciprocal of the comminution energy as determined from the electrical energy required to grind the cereal to fine particles in a small hammer mill (see text). 3RD EDITION Relative Sales 34 1.7. are similarly unable to adopt ordered crystalline structures. Cellulose has exclusively β(1−4) linkages.7_fm Page 35 Sunday. how the two types affected total energy intake. Many different structures are possible with different linkage geometries and different monosaccharide units. Cellulose’s regularity enables it to adopt ordered crystalline structures of polysaccharide chains held together by hydrogen bonds. Polysaccharides with charged groups (COO– or SO3–). So what determines whether a particular fiber fraction is soluble or insoluble? The key to understanding why some polysaccharides are soluble in water and others are not is molecular structure.2 Dietary fiber content (g/100 g) for a range of different foods plotted against the corresponding satiety index. Solubility and viscosity both depend on the chemical composition of the fiber and the molecular structure of its component polysaccharides. Solubility Solubility is a major factor in the nutritional properties of dietary fiber. in this case because electrostatic repulsion prevents the molecules from packing close together in ordered structures. and the water-soluble β-glucans from barley and oat bran.3).7.14 Polysaccharides are composed of linked monosaccharide units. as in the arabinoxylans in wheat. Viscosity is caused by physical interactions between polysaccharide molecules in solution — in simple terms. and these compounds are also water soluble.2387_ch2. of which the most common is D-glucose (Figure 2.14. which is insoluble. the irregular structure of the β-glucans prevents the formation of ordered crystalline structure. 2001 6:31 PM PHYSICAL CHEMISTRY OF DIETARY FIBER 35 Dietary fiber (g/100g) 35 30 25 20 15 10 5 0 0 50 100 150 200 250 300 350 SI Figure 2. May 6. are also often water soluble.14 In contrast.7.14 Viscosity Almost all water-soluble polysaccharides produce viscous solutions. whereas the β-glucans have mixed β(1−4) and β(1−3) linkages. But physical properties (such as solubility) are determined more by the linkages than by the nature of the monosaccharide units.15 These ordered structures are insoluble.14 This is illustrated by comparing two forms of poly-D-glucose familiar from other chapters of this book — cellulose. such as the pectins and carrageenans. by the . generally speaking. so these polysaccharides tend to be water soluble. Branched structures. 2001 6:31 PM 36 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. or bile acids in the matrix and by slowing mixing and diffusion in the intestine. Johnson and Read20 showed that guar gum appears to inhibit absorption by resisting the convective effects of intestinal contractions.7.) The viscosity then increases sharply with concentration and becomes more dependent on the rate of shear (this means. molecules becoming entangled.23 for example. and Tietyen and colleagues24 found that .16 At low concentrations the molecules are well separated from each other and free to move independently. May 6. the rate at which the liquid is stirred). Polysaccharides that give viscous solutions seem to be those most effective in lowering plasma cholesterol concentrations (Table 2. have suggested that the cholesterol lowering observed in rats fed high β-glucan barley fractions is related to the increased viscosity in the small intestine.7.7.1). 3RD EDITION Figure 2.17.4.16 (This is shown diagrammatically in Figure 2. it is not unreasonable to suppose that viscous.21.7_fm Page 36 Sunday.3 Representations of D.19 In an in vitro model using dialysis tubing.22 Newman and her colleagues. At a critical concentration (c*) the molecules become sufficiently crowded to start to interpenetrate one another and form a tangled network.18 ABSORPTION OF NUTRIENTS FROM THE SMALL INTESTINE Effects of Viscosity Viscous polysaccharides or polysaccharides which form a gel matrix may slow absorption by trapping nutrients. digestive enzymes. soluble fiber inhibits absorption of cholesterol and bile acids from the small intestine.20.and L-glucose in open chains and α-D-glucose and β-L-glucose in ring forms. in effect. Thus.2387_ch2. Polysaccharide solutions usually show “shear thinning” — the apparent viscosity decreasing with increasing rate of shear. which is .7. in which guar gum.3). locust bean gum. May 6. 2001 6:31 PM PHYSICAL CHEMISTRY OF DIETARY FIBER 37 Low Concentration High Concentration Figure 2. reduction of plasma cholesterol was again not related to viscosity (Table 2. In a study using everted sacs of rat jejunum. and guar gum and pectin have proved beneficial in controlling hyperglycemia.19 reducing the viscosity of oat bran fiber (by treatment with enodo-β-glucanse from Bacillus subtilis) reduced its ability to lower plasma cholesterol.7. Johnson and Gee25 found that viscous gums (guar and carboxymethyl cellulose) increased the thickness of the unstirred layer overlying the mucosa (Table 2. Table 2.7. c*) they become entangled.2). at higher concentrations (greater than the critical overlap concentration. gum acacia.7. no differences between plasma or liver cholesterol concentrations were observed.26 and in an experiment in which rats were fed methylcelluloses of different viscosity grades. But the relevance of the unstirred layer has been questioned. giving very viscous solutions.27 In another study. however.2387_ch2. appear to influence glucose absorption.b a b Quantity Ingested (g/day) Change in Plasma Cholesterol (%) 16 17 15 27 25 24 30 0 +1 –11 –17 –13 –16 –19 Composed of or containing viscous polysaccharides.28 Viscosity does.4 Random coil molecules — at low concentrations the molecules move independently through the solution. and fenugreek gum were compared. Also.7_fm Page 37 Sunday. Source: Data from Chen and Anderson. high dietary intakes of pectin (6 and 8% of the diet) decrease the availability of vitamin E in rats. Would also contain saponins which have cholesterol-lowering activity.29 Ellis and his colleagues30 have shown that guar gum inhibits glucose absorption in inverse proportion to the viscosity of the digesta.1 Effect of Fiber from Different Sources on Plasma Cholesterol Concentrations Source of Fiber Cellulose Wheat bran Whole oatsa Oat brana Pectina Guar gum a Beans a. in contrast.31 But. in humans.38. gum arabic. This is obviously a complex and controversial area.27 not viscous.40 Soluble fiber .26 3. and guar gum disappear almost completely during transit. also improves glucose tolerance32 and viscosity is not predictive of glycemic response.24 Table 2. about 40% of dietary cellulose is broken down.35. which depends. Bile acids are thereby diverted from the enterohepatic cycle.12 0.46 3.7_fm Page 38 Sunday.39 FERMENTATION IN THE COLON Effects of Physical Properties and Structure on the Fermentability of Fiber Although.09 0.34 0. and the loss is made good by conversion of cholesterol into bile acids by the liver. particularly solubility. The chemical properties of the polysaccharide also appear to influence the types of microbial activity present in the large intestine. and adsorption of bile acids onto pectin is physically unlikely because both molecules are negatively charged at gut pH.36 The cholesterol-lowering effect of dietary fiber. Pectin.35.37 Moreover. Adsorption of Bile Acids Another questionable concept is the frequently cited ability of fiber to bind bile acids.40 Fermentation depends on the accessibility of the polysaccharide molecules to the microorganisms. in fact.7.7. by definition.2387_ch2. fiber is not broken down by the enzymes of the gastrointestinal tract. it is suggested. can (at least in part) be explained by adsorption of bile acids to fiber in the small intestine. pectin also causes increased fecal excretion of bile acids. in turn. 2001 6:31 PM 38 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.33 Eastwood and Morris34 have concluded that there is.2 Apparent Thickness of Mucosal Unstirred Layer in Jejunal Sacs Preincubated with and without Guar Gum or Carboxymethylcellulose Unstirred Layer Thickness (µm) Control With Polysaccharide Polysaccharide Guar gum (viscosity 16 cP) Carboxymethycellulose 317 ± 15 346 ± 12 468 ± 25 402 ± 12 Source: Data from Tietyen et al.60 3.3 Relationship between Viscosity and Cholesterol-Lowering — Comparison of Three Galactomannans and α-Cellulose in the Cholesterol-Fed Rat Fiber Viscosity α-Cellulose Guar gum Locust bean gum Fenugreek gum (Insoluble) High Medium Low Total Plasma Cholesterol (mM) 4. May 6. on chemical structure and physical properties. oat bran) can indeed increase fecal excretion of bile acids. lost by fecal excretion.09 Source: Data from Topping et al.16 ± ± ± ± 0. little evidence to suggest that viscous polysaccharides inhibit transport across the small intestinal epithelia. virtually all fiber fractions are broken down to some extent by the microorganisms in the colon. 3RD EDITION Table 2.36 Although dietary fiber (for example. adsorption of bile acids to fiber preparations in vitro is so small as to be trivial when expressed quantitatively in terms of µmoles adsorbed per gram of fiber. In contrast. The extent of breakdown may also be related to the physical structure of the plant — fiber from fruits and vegetables appears to be more fermentable than that from cereals. has the more protective effect. Thus. slowly fermented fiber. insoluble fiber fractions ferment much more slowly in a process that is continuous during transit. Some components involved in the cohesiveness of the fiber matrix may be progressively solubilized. Weaver and colleagues45 found that propionate as a fermentation product promotes the development of cancer.43. Again.41 Thus. such as guar gum and pectin. confined to the proximal colon. and it is possible that pectins have a dominant role in determining cell wall porosity.27 and short-chain fatty acids. particularly butyrate. It is often suggested that short-chain fatty acids may be factors in controlling cholesterol metabolism. Particle size may vary during transit through the gut as a result of chewing. whereas slowly fermented fiber produces a sustained release of metabolites along the full length.2387_ch2.42 In this connection. The explanation seems to be that readily fermented fiber produces a short-lived burst of metabolites. reaching the distal colon.47 . such as wheat bran. whereas butyrate was confirmed as having a protective effect. offer little protection from colorectal cancer.44 These studies are summarized in Table 2. the kinetics of fermentation appears to be a particularly important factor. Table 2. Chemical composition may be another significant factor here.44 The fermentation pattern also seems to be important. grinding in the stomach. Animal studies have shown that readily fermented fiber. In a series of rat studies with partially hydrolyzed guar gum.46 Porosity and surface area are also relevant factors in controlling susceptibility to bacterial and enzymic attack. and bacterial degradation in the colon.7. particle size before ingestion of the food is not necessarily related to the subsequent behavior of the fiber during transit.40.7_fm Page 39 Sunday.44 Particle Size and Porosity Particle size particularly affects fermentation.4. may influence the development of colorectal cancer. 2001 6:31 PM PHYSICAL CHEMISTRY OF DIETARY FIBER 39 fractions are very accessible and ferment rapidly in the proximal colon. May 6.4 Fiber Type Summary of Published Reports on Effects of Dietary Fibers on Tumorigenesis in Rat Models of Large Bowel Cancer Effect on Tumorigenesis Protective Equivocal Enhanced Total Number Poorly fermentable: Cellulose Lignin 8 2 3 0 0 0 11 2 0 16 1 3 1 3 5 1 Slowly fermentable: Wheat bran 7 9 Rapidly fermentable: Guar Pectin Oat bran 0 0 0 2 2 0 Source: Data from Young. these factors are related to origin and processing history.7. The range of particle sizes depends on the types of plant cell walls present in the foods and on their degree of processing. although some evidence suggests otherwise. physical effects have a profound effect on the kinetics of release of metabolically important metabolites such as short-chain fatty acids. ) Absorption of Small Molecules and Ions In addition to binding water. The most obvious demonstration of the ability of soluble polysaccharides to hold water is the phenomenon of gelation. (The percentage of water retained by the hydrated fiber pellet was measured.2387_ch2.52 Insoluble fibers can also absorb water but more in the manner of a sponge. 2001 6:31 PM 40 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. on cooling to room temperature the solubilized polysaccharides can reassociate or “retrograde.51 Water-Holding Capacity Polysaccharides are hydrophilic molecules. In the plant. Its authors reported that a method based on centrifugation gave tolerably consistent results across different laboratories.52 A relatively small amount of polysaccharide. However. A recent European collaborative study53 addressed this problem. amylopectin has mixed α(1−4) and α(1−6) linkages. but the results were dependent on the sample weight used. Starch is poly-D-glucose in a complex structure — a mixture of linear amylose and highly branched amylopectin. often causing considerable swelling. unable to flow away. It has been reported that absorption of zinc and iron are actually enhanced by sugar beet fiber and that the inhibition of absorption observed with wheat bran is caused by phytate. 3RD EDITION Microstructure — Resistant Starch Resistant starch (discussed in more detail in other chapters of this Handbook) behaves functionally as dietary fiber because it resists digestion by the enzymes of the stomach and small intestine. and the system has the semisolid properties characteristic of a gel.54 The number of free carboxyl groups and particularly the uronic acid content seem to be the major factors determining the ability of polysaccharides to bind metal ions.” 48 Retrograded amylose is more than 70% resistant to amylolysis in vitro 49 and restricts access of the enzyme to the starch substrate as a whole in vivo. They have shown that resistant starch has substantial segments of the amylose chains in the form of double helices loosely arranged into aggregates. leading to the formation of helical structures. Single-chain material is also present as imperfections in helices or as chains trapped between aggregates. polysaccharides have the ability to bind other polar molecules and ions. soluble and insoluble polysaccharides alike have the ability to hold water. but the majority of starch-rich foods have been processed by a combination of heat and moisture which disrupts the native granular structure and causes partial solubilization of the starch polysaccharides. The water is held within the polysaccharide matrix. May 6.55 . they have numerous free hydroxyl-groups which can form hydrogen bonds with water. the starch is mostly packaged as starch granules. can be enough to entrap the water in which it is dissolved in a three-dimensional network of polysaccharide molecules. The doublehelical conformation appears to be the primary barrier to enzyme action. that these ions are subsequently released and absorbed as fiber is broken down in the colon.41. Starch that has been gelatinized by heating in water is readily hydrolyzed by the amylolytic enzymes. though.7_fm Page 40 Sunday. It is possible. This probably depends on sample size because of a proportionate increase in the contribution from soluble material retained by the pellet at increased sample weight. The reduced mineral availability and electrolyte absorption associated with some diets high in fiber appear to be due to binding of metal ions.50 Gidley and his colleagues51 have investigated the molecular structure of resistant starch in great detail using a combination of physicochemical techniques. This is a case in which microstructure has a powerful effect on physiological properties. Consequently. Amylose has α(1−4) linkages which introduce a twist in the molecule. Measurement of water-holding capacity is surprisingly problematic for such an apparently simple parameter. They also form a hydrophilic matrix in which water is entrapped — but where the quasi-crystallinity of the polysaccharide remains and water fills the interstices. giving rise to its highly branched structure. such as 1% agarose. 58 Additionally. These functions depend to a large extent on physical properties which also change.7_fm Page 41 Sunday.5).41 The physical form of the fiber is also important.57 and it is probably for this reason that resistant starch has a mild laxative effect. we are dealing with a highly complex kinetic system which is still not well understood. and analyzes as dietary fiber by the usual methods. with rapidly fermented fiber different in its effects from slowly fermented fiber which gives a sustained release of low molecular weight (osmotically active) metabolites along the full length of the colon. The polysaccharides of which dietary fiber is mostly composed are complex structures in which the geometry of the linkage between monomer units largely defines physical properties. May 6.56 Thus.5 Effect of Fiber Supplements on Fecal Bulk Fiber Supplement Increase in Fecal Wet Weight (%) Oat bran Pectin Guar gum Apple Carrot Cabbage Cellulose Wheat bran.59. will necessarily be of significant health benefit to consumers. Coarsely ground wheat bran is a very effective fiber source in increasing fecal bulk. Although the chemistry of dietary fiber is now well defined. some laxation effects may be due to the short-chain volatile fatty acids produced by fermentation. fine 15 16–35 20 40 59 67 75 80–127 24 Source: Data from Cummings. The ability of different fiber types to increase fecal bulk depends on a complex relationship between the chemical and physical properties of the fiber and the bacterial population in the colon. coarse Wheat bran. Thus. The major components are undigested residues plus bacteria and bacterial cell debris. . 2001 6:31 PM PHYSICAL CHEMISTRY OF DIETARY FIBER 41 LAXATION The huge displays of laxatives to be seen in any supermarket or pharmacy testify to popular concern with this topic. Cereal fibers are highest in pentoses and therefore have the greatest fecal bulking power.61 CONCLUSIONS The physiological functions of dietary fiber change as it progresses through the gastrointestinal tract. the effect of fiber on laxation depends not only on the undegraded fiber residue but also on bacterial cell mass.41. whereas finely ground wheat bran has little or no effect — and may even be constipating.7.60 Osmotic effects may be important in a way that is not yet well defined. it does not in itself predict biological activity. Table 2. These form a spongelike. Kinetic effects may also be significant again here. This means that we cannot assume that any material which falls within the chemical definition of dietary fiber. Biological activity depends on physical properties which do not relate in any simple way to crude chemical composition.2387_ch2.56. water-holding matrix which conditions fecal bulk and consistency.57 This is illustrated by comparing the results from studies of increased fecal bulk in response to various fiber supplements (Table 2.7. Feces are approximately 25% water and 75% dry matter. . Smithson.. Nutr. The satiety index: a new method to measure the filling powers of foods. Adv. 1. A. R. 16. N. W. O’Sullivan. Nutr. Proc. Concentration and shear rate dependence of viscosity in random coil polysaccharide solutions. and Farmakalidis. 1703. 32. Rees.. 6. R. 1999. H. Robertson. Bryson.. The influence of guar gum on intestinal transport in the rat. and Oakenfull.. J. 1987. Effects of plant fiber in decreasing plasma total cholesterol and increasing high density lipoprotein cholesterol.. Int. Ed. 1986. 18. 27. M. I. 215. J. Fletcher.. J. and Gatty. and Atkinson. W. Wholemeal bread and satiety. 14. C. E. Russ. Nevins. W.. . C. Stevens. J. 1. Cutler. Ross-Murphey.. Millane. N. W. 1988. J. Levine. Rep. R. J. L. Mickelsen.. 1983. R. W.. Proc.. P.. 1974. G. Clin. 812.Blackburn. Gastric emptying. 24. R. Gastroenterology. S. Is fiber satiating? Effects of a high fiber diet preload on subsequent food intake of normal-weight and obese young men.. Gut.. 1995. Johnson.... 26. 675. O. R. H. Brit. S. Delargy. Effects of amount and type of dietary fibre (soluble and insoluble) on short-term control of appetite.. J. 1980. BeMiller.. Holt. S. B. 1979.. 9. Effect of psyllium gum and wheat bran on spontaneous energy intake. and Johnson.. J. The entanglement concept in polymer rheology. L.. 812. E. 1988. T. Colmey. Tietyen. J... 32. R. R.. Trimble.. Internat. Nutr. D. New York. Schoemaker. J. 67. A.. Science. C. J. 5.. J. Elsevier. J. 386. Aust. 83. 46.. D.7_fm Page 42 Sunday. 17. K..2387_ch2. Food Sci. 21. 13. Schneidecker. T. J. and Price. Am. Makdani. Hypocholesterolemic potential of oat bran treated with an endo-β-D-glucanase from Bacillus subtilis. J. Soc. J. A. J. Cotton. R. Newman. and Garrow. Hydrocolloid systems as fat mimetics in bakery products: Icings glazes and fillings. and Gray. 1995. Oakenfull. Effect of gel-forming gums on the intestinal unstirred layer and sugar transport in vitro. J. 1981. J. Intestinal diffusion barrier: Unstirred layer or membrane surface mucous coat. D. Danielson... 1977. Hum.. and Gee. 1979. 1997. 1989.. N. 515–522. rheological characterization and implications for food systems. 15. W. Carbohydrate Polym. Res.. R.. T.. Nutr. Titcomb. Am. Millar. Graessley... Haskell. R. A. 8. London. F. D. B.. 214. S.. J. Superko. J. 74. Eds. P. 1987. 11. 130.. A.. 3RD EDITION REFERENCES 1. 1988. Polysaccharide Shapes. 59.. and Illman. E. J.. Nutr.. P. L. Dore. R.. Nutr.. 1985. 1241. in Polysaccharide Association Structures in Food. 1995. 60. A. J. W. R. 3. H. N.. C. B.. Porikos. 11. E. 22. Nutr. M. fibre and absorption. 15. Rolls.. 25. L. C. A.. I... A. Nutr. 119. 398. Effects of solid and liquid guar gum on plasma cholesterol and triglyceride concentrations in moderate hypercholesterolemia. Appetite. Levitsky.. B.. Cereal Foods World. 62... 50. G. D. Newman. 2000. 22. Biol. D. S. 2. D. T.. Nutr. S. Lancet.. Brit. Topping... D. Effects of a high fiber bread diet on weight loss in college-age males.. Clin.. Chen. Marcel Decker. 113.. M. Food Aust. Cardiol. H.. J. and Read. 162. London. J. A viscous fibre (methylcellulose) lowers blood glucose and plasma triacylglycerols and increases liver glycogen independently of volatile fatty acid production in the rat. W.. 132. H. 4. Kalkwarf. van Soest. J. L. 1981. Do viscous polysaccharides slow absorption by inhibiting diffusion or convection? Eur. 21. L. G.. Polysaccharide molecular structures. 34. 310. Chapman and Hall. Brand Miller. D. N. and Blundell.. 48. 16... and Anderson. 20. Am. 7. 1998.. Gee. K. 10. Schwartz.. and Chandrasekaran. 19. Jacobs. K. 1979. Sustained pectin ingestion delays gastric emptying. O. W.. R. Lipid levels and digesta viscosity of rats fed a high-fiber barley milling fraction. A.. The role of energy density in the overconsumption of fat.. C. B. 5. Sawrey-Kubicek. R. E. and Track. 7. Johnson. 23. J.. Nutr. S. 268S. J.. in Frontiers in Carbohydrate Research — 1: Food Applications. Petocz.. Ward. S. Polysaccharide solution properties: origin. Exp. Rees. N. K. and Farquhar. Nutr. May 6. 558. J. E. R. 51. J. Walter. S.. 872. Soc... Singh.. 17. Sci. Clin. Leeds. Nutr. H. A satiety index of common foods. D. Morris. and Hagamen. D.. I. 1982. R. P. Use of high fiber diets for the out-patient treatment of obesity..M. 42. J. Holt. 42. S. 1. Eur. A. D.. 1986. and Berardinelli. 49. D..-J. J... A.. Parker. L. 307.. 12. and Roe. 2001 6:31 PM 42 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Polymer Sci. Oakenfull.. F.. 1981. Food. Comminution energy and the sales of breakfast cereals. Clin.. Morris. 162. Med. 193. and Schneeman. 51. R. M. 217. 46. P.. V. F.. 71. 31.. A.. Am. 29. 28.. O. 35. Polym.. T.. 233. Whittam. Botham. Royal Australian Chemical Institute. Ring. 29. R. G. Oakenfull. May 6. Bacterial fermentation in the human large bowel: time to change from the roughage model of dietary fibre?. L. Brit. 1968. and Barry. Adsorption of bile salts from aqueous solution by plant fibre and cholestyramine.. A. 41.. and Johnson. E.. Molecular order and structure in enzyme-resistant retrograded starch. Dietary guar gum alters colonic microbial fermentation in azoxymethane-treated rats. Food Chem. Polym.. M... Structural and physical properties of dietary fibres. 135. M.. A. S.. Oakenfull. H. 34. V. 307. and Ring. T. 266-274. Physical properties of dietary fiber that influence physiological function: a model for polymers along the gastrointestinal tract.. Nutr. Relationships between physical characteristics of sugar beet fibre and its fermentability by human faecal flora. G. 36. Nutr. and Pfeffer. Rep. Gassull. 74... T. 1996. Br. Gidley.. Russell. P.. J. 1. and Illman.. G.. Schaus. J. and Morgan. 1997. G. J. Bioavailability of vitamin E in rats fed graded levels of pectin.. D. M. Res. 310. Young. Effects of pectin on intestinal absorption of glucose and cholate in the rat... 379. ACS Symposium Series. Nutr. P.. 51.. 50. 144. M. Nutr. Goff. M. Guillon. J. J.. and Sidhu.. R. Orford.. Hoagland. A.. Internat.. 539. Nutr. J. J.. A. J.. The effect of high-molecular weight guar gum on net apparent glucose absorption and net apparent insulin and gastric inhibitory polypeptide production in the growing pig: relationship to rheological changes in jejunal digesta.. 126... Nutr. 1991. fibre analogues. 40. F. and Morris. 23.. Brit.. D. G. P. Alpern. and Fenwick.-L. E. Med. Eastwood. and Hockaday. 1437.. G. 49. E. 1985. D. Melbourne.. 436. G. Sharma. 1986. G. Internat. Schneeman. J. T. Morris... G. S.. 563. 152. S. Weaver. A.. . D. Nutr. F. and consequences of processing on human physiology. Hypoglycemic effect of gum acacia in healthy human subjects. J. Robertson. Leeds.. Noel. Carbohydr. J.. Low.. 43. G. H. Collier. 1985. Brit. 1269. de Lumen. Hood. D. A. D. 97. S. 5.. 1996. Biophys.. A. The roles of amylose and amylopectin in the gelation and retrogradation of starch. H. Polym.7_fm Page 43 Sunday.. A. L. 1992. 2001 6:31 PM PHYSICAL CHEMISTRY OF DIETARY FIBER 43 28. Eastwood. Relationship between structure and function of dietary fibre: a comparative study of the effects of three galactomannans on cholesterol metabolism in the rat. S. Jenkins. Food Ingredients J... 32. 271. 68. 1986. R. 55. J.. 52. 118. Hoffmann. A. Oakenfull. 30. A. Burkitt. 1978. Miles. Jpn. A.. Effect of soluble dietary fibre on the viscosity of gastrointestinal contents and the acute glycaemic response in the rat. D. Jenkins. methods of analysis and physiological effects. Bruxelles. R.. Carbohydr. 38. 1986. and O’Dea. Dietary fiber and cancer. Dietary fibre and bowel cancer: which fibre is best? Cereals International. 195. J. 45. and Ring. Proceedings of an International Conference held in Brisbane Australia. 48.. V... 1974. R. L. and glucose tolerance: Importance of viscosity. O. D. M. 2000. B. J. 37. Nutr. Topping. Aust. Darke. 347. M.. Cooke. 37. I.. Fishman. Thibault. P. M. Food Res. Med. Food Technol. 165. 531. R. Clin. 167. A study on the in vivo digestibility of retrograded starch. D. 48. 104. J. Evans.. 28.. Cummings. P. Nutr. G. J.. Carbohydr. 104. The Large Intestine in Nutrition and Disease. 1978. L. M. K. D.Orford. Ellis. 1988.. 299. Binding of bile salts in vivo by non-nutritive fiber. 1986. 30. 40. J.. and Sidhu. J. B. D. K. E. and Hamilton.... Parfitt. J.. Tangel.. and Jen. J.. R.. Krause. 1994. Role of pectin in binding of bile acids to carrot fiber. 1991. Reyes. J. R. P. Dietary fibres. Carbohydr.. C. A. Nutr. D. G. D. 1995. and Omaye.. M. D. 458. D.. Res. R. J. Gee. 1985. A.. 1988. E.. Resistant starch: its chemical form in foodstuffs and effect on digestibility in vitro. Guillon. D.. B. T..2387_ch2. Dietary fiber: physical and chemical properties. J. D. Biochim. 1392. and Greenwell. Studies on the adsorption of bile salts to non-absorbed components of diet. Alberti. 1996. I. L. pp. 115. J. P.... R. Carrington-Smith. Auffret. Kritchevsky. D. 39. Brit.-F... J.... M. P. S. L... and Story. G. and Champ. G. 1979. J. Eds. 1998. Acta. J. Roberts. 44. in Chemistry and Function of Pectins. Morris. M. 47. Institut Danone. 40. Chow. Nutr. D.. F. J. 1995. Oakenfull. 42. D. Gelation mechanisms. M. J. 33. and Stragand. L. A. 33. Wolever. S. 263. T. 49. D. Schneeman.. P. E. 733. T. 2000. bran and guar gum. D. M.Wiss. 547.. 60. 33.. D. 3RD EDITION 53. and Dysseler.. 59. P. Lebesnm. Aust. 39. R. A. Southgate. L. R. Cummings. The effects of sugar beet fibre and wheat bran on iron and zinc absorption in rats. and Write.. 273. Mechanism of action of dietary fibre in the human colon. Dietary fiber: physical and chemical properties. D. J. A. Thibault.. Beatty. J. A. D.. Colonic response to dietary fibre from carrot. Lancet. H. 61.. Nutr. and Cummings.. Kingman.. Southgate. J. S. R.. F. 54. apple. A. G. S. 58.. Branch. 64. 1990... methods of analysis and physiological effects. J.. M. de Monredon. Nutr. M. The nutritive value of wheat bran. D. May 6. 1986. A. Effect of isolated complex carbohydrates on cecal and fecal short chain fatty acids in stool output in the rat. Stephen.. 1990.. Nutr.. Brit. and James. H. A. Waldron. Guillon.. 75. 1978. Food Technol. Bowen. Fairweather-Tait.-F. 57. W. in Dietary Fiber: Chemical and Biological Aspects. Topping. Edwards. 288. A.2387_ch2.-Technol. Eds. 40. I. J. 55. Soluble fiber polysaccharides: effects on plasma cholesterol and colonic fermentation. Food Tech. T. Jenkins. and Eastwood. 5. 1980. Hydration properties of dietary fibre and resistant starch: a European collaborative study. Amado. Royal Society of Chemistry. Houston. 1991. A. Robertson. 1996. and Topping. A. H. J. Brit... Rev. N.. 284. 195.. K. Cummings J.. C. D.7_fm Page 44 Sunday... . L. A. Johnson... and Englyst. J. 2001 6:31 PM 44 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.. J. 56. and Fenwick. cabbage. 1987. O. I.. Nature. Digestion and physiological properties of resistant starch in the human large bowel. A. J. T. S. B.. 104. T. Bingham. 283. J.. London. Oakenfull. H. u. W.. 72. F.... shrimp.3 NUTRITIONAL AND PHYSIOLOGICAL PROPERTIES Since chitin and chitosan have the same glycosidic linkages in their molecules as cellulose. it is generally accepted that chitin is highly acetylated. while chitosan is extensively deacetylated. depending on the particle size. which usually contains 70–95% of D-glucosamine. and the reverse is true with chitosan. While the chitin contains predominantly N-acetyl-D-glucosamine units.50 © 2001 by CRC Press LLC 45 . In chitin. but it is also present in many fungal cell walls and yeasts. and other shells of shellfish. they are not hydrolyzed in the small intestine and are partially degraded by the microbial flora in the large intestine. chitosan is insoluble. while chitosan has a unique solubility profile.8 Chitin and Chitosan — Special Class of Dietary Fiber Ivan Furda STRUCTURE AND DEFINITION Chitin and chitosan are natural aminopolysaccharides which consist of linear chains containing and D-glucosamine units which are linked together by (1–4)-β glycosidic linkages. Chitin behaves as typical insoluble dietary fiber. such as Mucor rouxii. or polycationic. May 6. Although there is no sharp distinction between these two biopolymers. is the presence of amino groups in their molecules. chitosan is based primarily on D-glucosamine units.00+$1. unlike other fibers. they cannot be hydrolyzed by enzymes secreted in humans. concentration. therefore. lobsters.2 Chitosan had been also found in different fungal cell walls. In the gastrointestinal tract. while polysaccharides of common dietary fibers are either neutral or acidic (polyanionic). 2001 6:33 PM CHAPTER 2. chitin and chitosan are predominantly of animal origin but could also be of plant origin.3. Therefore. and other 0-8493-2387-8/01/$0. and a few others. they are basic.8_fm Page 45 Sunday.4 cell walls of the green alga Chlorella. Mucorales are used for preparation of numerous oriental fermented foods such as tempeh. the typical content of N-acetyl-D-glucosamine is 70–95%. Chitin is mainly present in exoskeletons of arthropods such as insects.1 N-acetyl-D-glucosamine ORIGIN AND SOURCES Unlike other common dietary fibers which are of plant origin. namely Mucorales.2387_ch2. When ingested. however. ragi. The unique structural feature of these polysaccharides. in the stomach. it becomes fully or partially solubilized. crabs. sufu.4 and others. due to low pH. 18 and act as an effective lipid binder in vivo. Biol. Lab. 1978. Hypoglycemic and hypolipidemic effect of chitosan in normal and neonatal Streptozotocin — induced diabetic mice.. Cambridge. D. chap. and Hasegawa. Fish Res.13–17. Fenton. A. Eds. it was shown in animals and in humans that chitosan can augment high-density lipoproteins13–15. 11. V..25 In order to explain chitosan activity.. 18. G. Int. R. while one-third of the subjects did not respond. J.. Enzymatic hydrolysis of chitosan. A. T. C. Lately. one lasting 4 weeks27 and one 8 weeks. The initial studies were conducted in experimental animals such as rats.8_fm Page 46 Sunday.8 rabbits. E. 214. chitosan did not show adverse effects on the morphology of the gastrointestinal tract when tested in experimental animals at moderate levels. and Seino. and Fujita. Enzymic synthesis of chitin and chitosan.. Furda. In the small intestine. Hiratsuji. A. R. These studies lasted on average 4 weeks. except for a few transient effects such as constipation. 89. this area is currently under detailed investigation.. T.10 and broiler chickens. B. in Unconventional Sources of Dietary Fiber. which lasted 12 weeks.. Y. Vol.. Oxford. Unlike strong anion exchangers such as cholestyramine.15. A. a number of clinical studies confirmed the results previously observed in experimental animals. E.19–23 The binding and reduction of absorption and utilization of lipids should lead to weight loss.. 1968.. Amer. Muzzarelli. 38. Arai.24..9. 1961. M. R. MA. in Chitin. M. chitosan precipitates and thus becomes again insoluble. Satchithanandam. Ed. Sugano.11. Pergamon Press. This effect has been observed in recent studies in which a controlled low-calorie diet in placebo and in test groups was combined with ingestion of chitosan during main meals by overweight subjects.28 In the clinical studies no adverse effects have been reported.. Cassidy.29 Since important benefits may be gained from using chitosan in oral applications. Symposium Series..24..7 mice.7. Nutr. 4. The polycationic nature and the unique in vivo solubility of chitosan may be responsible for special physiological attributes of this biopolymer. 1995. 18. D. They have been reviewed. Mihara. S. Furda. 2. 2001 6:33 PM 46 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.. 5. a number of mechanisms have been proposed and investigated..C. Bull.. Muzzarelli.17 no weight loss was observed. R.. Rotgers. S. This is similar to that of salt or sugar.2387_ch2. 3. it has been repeatedly reported that chitosan has strong hypocholesterolemic and hypolipidemic properties. Bull.12 to mention a few. Plant Cell Physiol. 56. Pharm.6. T.. Washington.. 278. T.. Muzzarelli. Y. M.. 1978. Y. 8.. I. Toxicity of chitosan. 3RD EDITION factors. Tokai Reg. Rep. Tsuura. Ed. Clin. two-thirds of the subjects showed significant weight loss. 1. 525. and Furda... chap.5 the LD50 of chitosan in laboratory mice is 16 g/kg of body weight. Vahouny.. due to the neutral pH. Lightfoot. and Eveleigh. Nutr. . Aminopolysaccharides — Their potential as dietary fiber. Usami. F.. K. and Pariser.. Davis.. H. Massachusetts Institute of Technology. There are a few plausible explanations why the studies using free-living diets did not always result in weight loss.14. 8..25 The studies in which free-living diets with no restrictions were used showed conflicting results.26 In two similar studies... 25.. 1978. Hypocholesterolemic effects of chitosan in cholesterol fed rats. and the subjects consumed 2 to 3 g of chitosan per day.13–17 In addition.. 7. 1983. According to Arai et al. M. Comparative effect of chitosan and cholestyramine on lymphatic absorption of lipids in the rat. D. I. American Chemical Society. Miura.. Kinumaki. Since the late 1970s. A. and at this time it is believed that a few mechanisms working sequentially may be involved. 1983. Fujikawa. May 6. A. Change of glucosamine content of Chlorella cells during their life cycle.. B. I. Y. REFERENCES 1... Chitin and chitosan are biopolymers of very low toxicity. In one such study. Ishida. 6. in Proceedings of the First International Conference on Chitin/Chitosan. 531. 2. and Pettersson. Y. Wuolijoki. J. 19. 29.. 17. H. Y. Sciutto. 714. J. 13.. Brit. and Marletta.... P. Han. J. Veneroni. Furda.. Nutr. Biotech.. L. Biochem. Hirano. A. Kimura. J. Atec. U. H. and Ernst. Biochem. Takiguchi.. I.. Decreasing effect of chitosan on the apparent digestibility by rats fed on a high-fat diet. Decrease in serum LDL cholesterol with microcrystalline chitosan. K. 787. W. Ed. E. L. J. Biosci. and Nagyvary. Biosci. K.. K. 65. Shizukuishi.. I. 1983. 16(4). and Hasegawa. A. K.. Nakashima.. Agric. E. 21(5). 1439. Furda. A novel use of chitosan as a hypocholesterolemic agent in rats. 1996. Ther. Ther. Hara. and Ventura.. N. Effect of a new chitosan dietary integrator and hypocaloric diet on hyperlipidemia and overweight in obese patients. and Kobayashi.. 25.. 10.. Ed. Clin. E. J.. 27. 1999. Relat.. 1999.. feed intakes. 91. 21.. 12. 23.. Nutr. K. M. 1990. and Mitsuoka. 303. 76. E. M. 721. A.2387_ch2. Kanauchi. L.8_fm Page 47 Sunday.. S. Nutr. 69. Absence of hypocholesterolemic action of chitosan in high-serumcholesterol rabbits. 2000. Clin. Weight-reducing regimens in obese subjects: effect of new dietary fiber integrator. Y.. Y. Methods Find. 22.. 18. H. S. Exp. Biotech. T. Gudmunsson. Kanauchi. 41. and Lilja. 23. S. Tsuji. Veneroni.. plasma lipid and smallintestinal bile acid concentrations in response to feeding of chitosan and pectin..023. in Chitosan Per Os: From Dietary Supplement to Drug Carrier. J. J. 1993. Lipid lowering effect of chitosan dietary integrator and hypocaloric diet in obese subjects. in Chitosan Per Os: From Dietary Supplement to Drug Carrier.. Razdan. C. Metab. S. Y. M. E. Hypocholesterolemic effect of chitosan in adult males. Y. T.223. Pharm. and Pettersson. 1980. Abbot. 1995.. Ther... Pharmacol.. 78. Clin. and Anderson.. Akimoto. Deuchi. Y. Itakura.. a chitosan with fat binding properties and potential as a weight reducing agent: a review of in vitro and in vivo experiments. Contos.S. Bridges... Imasato. A. 57.. C. J. 1999. Imasato.. 15. The binding of micellar lipids to chitosan. Biopolymer L 112. S.. 24. Pharmacol. and Ilitalo.. Muzzarelli. Atec.. 16. C. J. M.. Biochem. D. 33. 215. Y. D. Macchi. and Colombo. Sugano. C. Hirvela. M. A. Int. 379. 283. Nutr... E.. Harkness.. 387. 59(5). A new approach to the treatment of obesity: chitosan’s effect on body weight reduction and plasma cholesterol levels. and Kawakami. 1988. Guarino. A... Disorders. 17. Proc. Boleyn. Biol. Nakagawa. 1994. Effect of chitin and chitosan on nutrient digestibility and plasma lipid concentrations in broiler chickens. Y. and Okuda. Biotech. 13.. Deuchi. Pittler. Reduction of fat storage during chitin-chitosan treatment in mice fed a high-fat diet. A. 1613.. DeBernardi.. M. Chitosan as an ingredient for domestic and animal feeds. L. Broiler chicken body weights. 4. 2001 6:33 PM CHITIN AND CHITOSAN — SPECIAL CLASS OF DIETARY FIBER 47 9.. 1214.. and Akiyama. Biosci. Nonabsorbable lipid binder. Maezaki. F. D. 49. R. Thompson. Fujikawa. 1996. G. S. M. . Li. 786. Randomized.. Ther..... 10. A comparison of the lipid-lowering and intestinal morphological effects of cholestyramine. 38. 2000. Y. Lipids. 199. 17(4).. 18. 1994. 53. E. Wood. T. B. J. Muzzarelli. Food Chem. W. Exp... Heldman. May 6... Hiratsuji. Nonaka. G... R. Grottammare. J. Tripodi. 357. Hirano. R. T. 16. A. 174. Y. 26. S... 1980.. Acta Toxicol. Razdan.. Thom.. B. Ji.. Soc. Sci...-K. J. Jing.. and Kobayashi. P. 1997. 11.. Brit. O. D. 58.. Kawai. J. Tsugita.. Kanbara. T.. Akiyama. 1.. Reduction of absorption of dietary lipids and cholesterol by chitosan and its derivatives and special formulations..... 1995. Fukuda. Med.. 1995. Terada. Effect of chitosan on renal function in patients with chronic renal failure... Giustina. Seino. Obes. Amer. A. P. Patent 4. Takekawa. T.. H.. P.. 14. 189.. 53. Acta Toxicol. Acta Toxicol. and Yamaguchi. N.. F. I. Food Agric. Jennings. N. 1997. Acta Toxicol. Eur. double-blind trial of chitosan for body weight reduction.. 20.. O. chitosan and oat gum in rats. 28.. J. Wadstein. 1995. Mechanism for the inhibition of fat digestion by chitosan and for the synergistic effect of ascorbate. Nauss. Grottammare. 2001 6:33 PM .2387_ch2. May 6.8_fm Page 48 Sunday. May 6. 2001 6:00 PM SECTION 3 Methods of Analysis for Dietary Fiber .2387_Section 3_fm Page 49 Sunday. 2001 6:00 PM .2387_Section 3_fm Page 50 Sunday. May 6. These authors also introduced special centrifugation tubes with fritted glass filters for separation of the insoluble fiber from the enzyme digest. Table 3. McCance et al. In the 1930s. and vegetables by determining the residue insoluble in 80% ethanol.1 measured total unavailable carbohydrates in fruits.1. Later on.1.6 Thomas7 improved the method by adding a pancreatin step. Weinstock and Benham4 used the enzyme preparation Rhozyme S with high amylase activity.00+$1. METHODS MEASURING INSOLUBLE DIETARY FIBER Methods employing both amylolytic and proteolytic treatment and separation of an insoluble undigestible residue are listed in Table 3.1. May 6.3 The main limitation of this approach is that the protein correction gives an unacceptable error in samples with low dietary fiber and high protein content. whereas a considerable protein residue remained associated with the fiber. Elchazly and Thomas8 omitted the crude Rhozyme preparation and used amyloglucosidase or takadiastase and trypsin or pancreatin. nuts. 4 7 8 9 51 . Thus. Similar procedures have been used more recently. This was corrected for starch measured after enzymatic hydrolysis with taka-diastase and for protein.2387_ch3. starch was removed efficiently.1 0-8493-2387-8/01/$0.5.50 © 2001 by CRC Press LLC Enzymatic Gravimetric Methods Measuring Insoluble Fiber Agents for Protein and Starch Solubilization Incubation Time (h) Rhozyme S Pancreatin Rhozyme or other amylase Amyloglucosidase or takadiastase Trypsin or pancreatin Pepsin Pancreatin 24 5 18 3 18 18 18 1 Ref. 2001 6:34 PM CHAPTER 3.1_fm Page 51 Sunday.1 Enzymatic Gravimetric Methods Nils-Georg Asp INTRODUCTION Enzymatic gravimetric methods date back to the 19th century.2. 1.15 However. also measuring soluble dietary fiber components.2387_ch3.6.12 are developments of Hellendoorn et al. . 19. i. It is generally agreed that soluble components should be included in the definition and determination of dietary fiber. representing in vivo undigestible material.2 Enzymatic Gravimetric Methods Measuring Both Soluble and Insoluble Fiber Agents for Protein and Starch Solubilization Incubation Time B.12−14 and it is impossible to define exactly the degree of hydrolysis obtained in vivo. However. the sum of undigestible polysaccharides and lignin. METHODS MEASURING INSOLUBLE AND SOLUBLE DIETARY FIBER Soluble dietary fiber components constitute a considerable fraction of the total dietary fiber in mixed diets5.20 used Bacillus subtilis amylase and protease in a single overnight incubation at neutral pH. subtilis protease Amyloglucosidase 16−18 h 20 h 18 h 18 h 1h 15 min 1h 1h 3h 15−18 h 15−30 min 30 min 30 min Ref.e.21 employ long (19.. In the more recent modification of Asp et al. Two of these methods6. May 6. 2001 6:34 PM 52 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. in practice the delimitation is always related to a method involving enzymatic degradation of starch in vitro.1. This analytical method is therefore related to a physiological dietary fiber definition.2 lists developments of enzymatic methods.15−18 Table 3. pepsin plus pancreatin.11 The residues of protein and starch associated with the dietary fiber after enzymatic treatment6. the protein and starch residues are dependent upon the choice of enzymes and the conditions for enzymatic digestion.1_fm Page 52 Sunday. The methods of Schweizer and Würsch21 and Asp et al.12 enzyme incubation time was reduced to 2 h. Both insoluble and precipitated soluble fiber were separated by filtration in crucibles with glass wool as a filtering aid.10. 20 21 6 12 22 23 Furda19. and separation of both insoluble and precipitated soluble fiber — either separately or together — was carried out by filtration using Celite as a filtering aid.9 using the physiological enzymes pepsin and pancreatin. Soluble dietary fiber is precipitated with ethanol. Soluble dietary fiber components were recovered by precipitation with 4 vol of 95% (v/v) ethanol.1. The main steps of this method are shown in Figure 3. which may be more or less related to in vivo conditions.12 were even considered an advantage.9 used only physiological enzymes. 3RD EDITION Hellendoorn et al.to 38-h) incubation times and repeated centrifugations to recover the dietary fiber fractions. Table 3. subtilis amylase and protease Pepsin Pancreatin + glucoamylase Pepsin Pancreatin Termamyl Pepsin Pancreatin Amyloglucosidase Pancreatin/trypsin Termamyl B. This fact has led most workers to prefer a more “chemical” definition of dietary fiber.1.13 Saunders and Betschart14 also suggested that indigestible protein could play a significant role in the physiological effects of dietary fiber.6 and include polysaccharides such as pectins and gums with important physiological effects.. A modification of the AOAC method with further simplifications has recently been tested collaboratively by Lee et al.1 Flow diagram of the method of Asp et al. incubation times. especially the viscosity. The AOAC method23−26 is based on the common experience of three groups that have developed enzymatic gravimetric methods for the assay of total dietary fiber. . seem to be related to the properties of the soluble fiber.8 using membrane ultrafiltration to separate the soluble dietary fiber components from the enzyme digest. May 8.12.20. and the experimental conditions giving a solubility closest to that in the intestinal content have not yet been defined.1_fm Page 53 Tuesday.12. attributed to soluble fiber.12 but with other enzymes.22 developed a method based on the enzymatic method of Elchazly and Thomas. The separate determination of soluble and insoluble dietary fiber was suggested in the different enzymatic gravimetric methods6. and filtration introduced by Asp et al.1.1.21 The main steps are shown in Figure 3.25. physiological effects on plasma lipid levels and postprandial glucose and hormone response.1.2. and a detailed step-by-step description is given at the end of this chapter. This method employs the short technique.1) and has been tested collaboratively for the AOAC method.27 and approved.12 Meuser et al. Furthermore.27 The solubility of the dietary fiber polysaccharides is method dependent. 2001 2:55 PM ENZYMATIC GRAVIMETRIC METHODS 53 Figure 3.19−22 (Figure 3.2387_ch3. however.3to 0.32 simple defatting with petroleum ether at room temperature is recommended whenever the fat content exceeds 10%. 3RD EDITION Figure 3. Probably much of the fat melts at the high temperature during the starch gelatinization step. May 8.1_fm Page 54 Tuesday. Asp et al.2 Flow diagram of the AOAC method for total dietary fiber. heating in a boiling waterbath in the presence of the thermostable α-amylase Termamyl gave sufficient gelatinization and prehydrolysis of the starch. Usually.5-mm screen is recommended.1.12 demonstrated that a 15-min. It should be noted. Starch Gelatinization and Hydrolysis Whereas some authors21.31. Defatting is usually recommended and may be necessary for proper milling. but it is recognized that milling to such a small particle size may present difficulties with some materials. and any fat residue will be dissolved by the alcohol and acetone washing of the dietary fiber residue after filtration. milling to pass a 0. Even in starchy materials .2387_ch3.23−27 DISCUSSION OF VARIOUS STEPS IN ENZYMATIC GRAVIMETRIC METHODS Sample Preparation Milling to small particles is essential for proper enzyme action.23. 22 employ autoclaving for starch gelatinization. were able to analyze dietary fiber with their method12 in samples with 20% fat without defatting. that Asp et al. In the AOAC method. 2001 2:55 PM 54 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. It is essential that all enzyme preparations used are free from contaminating activities hydrolyzing dietary fiber. DMSO (“resistant starch”).20 or a combination of both types of amylase.20 at neutral pH.34 Recently it has been suggested that the term resistant starch be used in a broader sense.9 acid-labile groups of dietary fiber polysaccharides.12. This can be checked most conveniently by running samples of known polysaccharide preparations through the whole procedure.23−27 the buffer strength of the incubation medium should be kept low enough to avoid excessive ash precipitation and thus an unnecessary source of variability between samples. such as arabinose residues in cereal pentosans. To be consistent with the definition of dietary fiber as the sum of undigestible polysaccharides and lignin. The AOAC method23−27 employs the B. Polysaccharides with DP (degree of polymerization) >10 are usually precipitated. which can easily be analyzed in the filtrate to measure starch..39 Although correction for ash is recommended. 2001 2:55 PM ENZYMATIC GRAVIMETRIC METHODS 55 such as wheat flour and potatoes.5 and reducing the incubation time to 1 h. may be hydrolyzed. which are recognized as difficult in this respect.31 The resistant starch in the fiber residue is mainly retrograded amylose. only a small amount of starch available to glucoamylase remained in the dietary fiber residue. values obtained with enzymatic gravimetric methods need to be corrected for undigestible protein associated with the fiber. May 8. Recovery of Soluble Dietary Fiber Constituents Most methods employ precipitation with 78–80% (v/v) ethanol to separate soluble fiber components from the enzyme digest.12. By increasing the pH to 1. heating should be performed immediately after Termamyl addition to avoid hydrolysis of dietary fiber components. The precipitate is recovered by centrifugation21 or filtration using glass wool19. subtilis protease originally used by Furda19.9.12.28 “Resistant starch.26 similarly as found with a gas-chromatographic method30 using 1-h boiling for starch gelatinization. especially if highly branched. starch available to glucoamylase only after solubilization with 2 M KOH. Thus..37 Coprecipitation of minerals may be a problem when using alcohol precipitation.1_fm Page 55 Tuesday. The 80%-ethanol precipitation is an arbitrary delimitation between polysaccharides included in dietary fiber and oligosaccharides not included.21−27 An advantage with glucoamylase is that it degrades the starch to free glucose. inactivating contaminating activities.12 or only glucoamylase. Procedures using ultrafiltration22 or dialysis have not been documented to be more selective.” i. It may differ from that in the original protein. remained associated with the dietary fiber in bread. since the true correct factor of the undigestible protein is not known. This has been demonstrated for arabinans and pectic substances in sugar beet fiber. This can be done by analyzing the dietary fiber residue for nitrogen with the Kjeldahl method.23−27 The universal protein conversion factor 6. Protein Hydrolysis At very acidic pH (around 1) as originally used by Hellendoorn et al.2387_ch3.23 The reason why the crude Termamyl preparation can be used is the high temperature employed.12 were able to use pepsin without measurable loss of dietary fiber constituents.19.38. but in some cases larger polysaccharides will stay in solution. . Asp et al.20 or Celite12.32 This fraction is resistant to degradation also in the small intestine of both rat33 and man.e.23−27 as a filtering aid.25 should be used. The total starch remaining in the dietary fiber residue prepared with an enzymatic gravimetric method can be differentiated into one fraction available to glucoamylase without alkali solubilization (“residual starch”) and another fraction available only after solubilization in 2 M KOH or dimethylsulfoxide.35 Different methods employ either only α-amylases6. representing all the starch escaping digestion and absorption in the human small intestine. Denmark. These modifications were introduced and approved in the third AOAC study. after which the method was approved by the AOAC. Once a protein correction is accepted. Finland. 2001 6:34 PM 56 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. — have recently been calculated in the same way and compared. a somewhat higher protein residue obtained at the very short incubation time in the protease step(s) can be accepted.40 Raw and processed wheat and potato products were analyzed in another interlaboratory study. The first interlaboratory study with the AOAC method for total dietary fiber (TDF)23 was the biggest collaborative effort reported so far. The AOAC method23 and the method of Asp et al.1. and soluble fiber determination was approved as the difference between total and insoluble fiber. It is recommended in several other countries.24 A Swiss collaborative study with a slightly modified TDF method26 reported the best precision measures obtained so far. direct determination of soluble fiber was approved. and the enzymatic gravimetric method of Asp et al. Germany. Alcohol precipitation is the most rapid way to separate soluble dietary fiber components from the enzyme digest. 3RD EDITION INTERLABORATORY STUDIES Enzymatic gravimetric methods have been compared and evaluated vs. Correction for undigestible protein and ash associated especially with the soluble fiber should be made to conform with currently accepted definitions of dietary fiber. as well as some variation in the residue due to choice of enzymes. as well. the enzymatic methods gave comparatively consistent results and good agreement with gas chromatographic methods. Ultrafiltration and especially dialysis are much slower processes and have not been documented to be more selective and complete. It identified a number of problems that were corrected in the second study.28 Enzymatic methods showed good agreement with each other and with gas chromatographic methods. for both processed and unprocessed cereals and vegetables.42 CONCLUSION Enzymatic gravimetric methods using alcohol precipitation of soluble dietary fiber components are suitable for assay of total dietary fiber or insoluble and soluble fiber separately. Sweden. both with and without correction for protein. Similar results have been obtained when comparing the component analysis methods of Theander et al.37 The lower values obtained with the Englyst method can generally be explained by the fact that this method does not include lignin and resistant starch. Norway. other methods. In the EEC/IARC study conducted in 1978. Centrifugation is an alternative. Iceland. Gravimetric dietary fiber values. May 6. especially to recover very viscous types of fiber that can give filtration problems. The AOAC method has been officially approved in Switzerland. have recently been introduced in the Japanese food tables.25 Insoluble fiber determination also showed satisfactory results in that study.3). three groups participated with enzymatic methods. The enzymatic treatment removes material that might interfere in later steps.41 The gravimetric AOAC method has been tested on a wider range of foods with more variable dietary fiber content and has generally shown reproducibility values as good as or better than the Englyst method.2387_ch3. Later on.27 All the collaborative studies reported so far — both with enzymatic gravimetric methods and with the method of Englyst et al. Although incompletely developed at that time. and The Netherlands.1_fm Page 56 Sunday. Filtration in glass filter crucibles can be used to recover both insoluble and alcohol-precipitated soluble components if a filtering aid such as Celite is used. and the . Enzymatic methods are useful as preparatory steps before detailed analysis of dietary fiber composition.12 showed good agreement on these samples and also on the samples analyzed in the first AOAC collaborative study (Figure 3. and the other is incinerated at 525˚C and the ash determined. fat extracted if containing more than 10% fat.23 using different enzymes on samples in the first interlaboratory study of the AOAC method. washed with 78% ethanol. are gelatinized with Termamyl (heat stable α-amylase). 95% ethanol.2387_ch3. After drying.12 and Prosky et al.1. The efficiency of starch removal and presence of resistant starch can be checked by analyzing the dietary fiber residue.1_fm Page 57 Tuesday.3 Comparison of the methods of Asp et al. 2001 2:56 PM ENZYMATIC GRAVIMETRIC METHODS 57 Figure 3.25 Procedure Duplicate samples of dried foods. The different symbols denote two different laboratories. THE DETERMINATION OF TOTAL DIETARY FIBER IN FOODS AND FOOD PRODUCTS: APPROVED AOAC METHOD OF PROSKY ET AL. Total dietary fiber is the weight of the residue less the weight of the protein and ash present. May 8. . and acetone. then enzymatically digested with protease and amyloglucosidase to remove the protein and starch present.24. the residue is weighed. total fiber value obtained in the gravimetric assay can serve as a control of the standardization of detailed analyses. The total residue is filtered. One of the duplicates is analyzed for protein. Then 4 vol of 95% ethanol (v/v) is added to precipitate the soluble dietary fiber. 7. Celite 545 (Fisher Scientific Co. 3. W.): refrigerate the dry enzyme after each use. Reagents 1. 8. porosity #2. e. Fisher Scientific Co. Check pH.400 g of sodium phosphate dibasic anhydrous (or 1. Porosity #2 in Europe signifies pores of 40 to 90 µm. Fritted crucible. capable of weighing to 0. Balance. Crucibles indicated in the procedure may not be available in Europe.. Sodium hydroxide solution (0. Acetone—reagent grade. soak in distilled water. Course ASTM 40 to 60 µm. Pyrex. Scientific Co. Dry at 130˚C for 1 h. Beaker (tall form) 400 ml. F-24390-B or F-243-90-C. which seems to have less breakage and gives the same results. 6.5 g of Celite 545 to dried crucibles before drying to obtain constant weight. R. 2001 6:34 PM 58 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. C-85251. This can be accomplished with either a multistation shaker or multistation magnetic stirrer. 60-ml ASTM 40 to 60. 3RD EDITION Apparatus 1. Boiling water bath. Vortex mixer. 8. 350 ml of 1 M HCl to 11 with distilled water. 2. Dilute to volume with DW. Vacuum source: a vacuum pump or aspirator equipped with an inline double vacuum flask should be used to prevent contamination in case of water backup.S.68 g sodium phosphate monobasic monohydrate (or 10. 4.1_fm Page 58 Sunday. depending on size needed. ash at 525˚C.). 7. 36060 Buchner.2387_ch3. whereas it means 40 to 60 µm in the U.753 g of the dihydrate) and 9. . Several collaborators have reported breakage of crucibles when the temperature was raised to 525˚C and have recommended Corning No. Vacuum oven at 70˚C and desiccator.. and store in desiccator until used..0: dissolve to 1. Store the enzyme solution in refrigerator after each use. 2. and may be purchased from Scientific Products Co.00 g sodium hydroxide AR in approximately 700 ml DW in 1-1 volumetric flask. Constant-temperature water bath adjustable to 60˚C and equipped to provide constant agitation of the digestion flasks during enzymatic hydrolysis. fritted disk. alternatively.). 23863-040. V. analytical. Sargent Welch Co. Hydrochloric acid solution (0. Protease P-3910 (Sigma Chemical Co. 5. 95% ethanol (v/v)—technical grade. cool. Clean thoroughly. 6.g. 3. Acid washed. 08237-1A. Amyloglucosidase A-9913 (Sigma Chemical Co. 10. 5. Termamyl (heat-stable α-amylase) solution: 120 L (Novo Laboratories Inc. Equivalent to Celite C-211.. 78% ethanol: mix one volume of distilled water with four volumes of 95% ethanol.275 N): dissolve 11. Dilute to volume with DW. Add approximately 0. 9.99 g of the dihydrate) in approximately 700 ml of DW. an air oven capable of operating at 105˚C can be used. and rinse in same.08 M pH 6. 4. The fritted crucible is a Pyrex 32940. Distilled water (DW).. Dilute to 11 with DW.): keep refrigerated when not in use. 11. Phosphate buffer 0. May 6.350 N): dilute a stock solution with known titer.1 mg. 2 with 0. 18. If the surface film that develops after the addition of the sample to the Celite is broken with a spatula. Increase the length of incubation time when the number of beakers added to the boiling waterbath makes it difficult for the beaker contents to reach an internal temperature of 100˚C. 1 g of sample accurate to 0.275 N NaOH if necessary.1 mg. two 10-ml portions of 95% ethanol.1_fm Page 59 Sunday. Allow precipitate to form at room temperature for 60 min. 15. Analyze the residue from one sample of the set of duplicates for protein. trapping the liquid. Wash residue successively with three 20-ml portions of 78% ethanol. Tare crucible containing Celite to nearest 0. Add 280 ml of 95% ethanol preheated to 60˚C. Adjust to pH 6. After taring the crucible containing the Celite. and residue to nearest 0. redistribute the bed of Celite in the crucible using a stream of 78% ethanol from a wash bottle. Add 100 µl Termamyl solution. 3.4 and formula for calculating “total dietary fiber” from the data in Figure 3. Suction is then applied to the crucible to draw the Celite onto the fritted glass as an even mat. 4. and two 10-ml portions of acetone. Weigh crucible containing Celite and ash to nearest 0.2387_ch3. Run a blank through the entire procedure along with the samples to measure any contribution from reagents to the residue. With some samples a gum is formed.e. Normal suction can be applied at washing. Shake gently at 5-min intervals. 19. 2. Protein is probably most easily analyzed by carefully scraping the Celite and the fiber mat onto a suitable piece of filter paper which can then be folded shut and analyzed for protein. Step 14. Add 5 mg of protease. i. When the fiber is filtered. 5. Weigh. Add 10 ml of 0.1. 17. the Celite effectively separates the fiber from the fritted glass of the crucible.0 to 4. 13. Measure volume after heating. 14. Filter enzyme digest from Step 12 through crucible. Incinerate second sample of the duplicate for 5 h at 525˚C.275 N NaOH solution. Cool in desiccator and weigh crucible. A piece of filter paper should be analyzed to assure that it will not affect the protein value obtained. if available.0 phosphate buffer to each beaker. 9. filtration is improved. Cool in desiccator.) 59 .350 M hydrochloric acid solution to adjust pH to 4. Collaborators should use the Kjeldahl analysis as specified in “Official Methods of Analysis” of the AOAC. 12. (See flow diagram in Figure 3. 6. 7.5 ± 0. 30 min should be sufficient.6.25 for the protein factor. Cover beaker with aluminum foil and place in boiling waterbath for 15 min. 20.. Add 0. 8.1 mg (sample weight should not differ by more than 20 mg) into 400-ml tall form beakers. Adjust to pH 7. Back-bubbling of air is another way of speeding up filtrations. Incubate at 60˚ for 30 min with continuous agitation. 11. Dry crucible containing residue overnight in a 70˚C vacuum oven or a 105˚C air oven. Cool. 10.1. Use 6. Add 50 ml of pH 6. Cover with aluminum foil.5. Incubate at 60˚C for 30 min with continuous agitation. Celite. it may be preferable to make an enzyme solution just prior to use with a small amount of (about 100 µl) phosphate buffer and pipette the required amount. Cover with aluminum foil.1 by adding 10 ml of 0. allowing for easy removal of the crucible contents.1 mg. Since protease sticks to the spatula. Check pH.1 mg. in duplicate. Long filtration times can be avoided by careful intermittent suction throughout the filtration. 2001 6:34 PM ENZYMATIC GRAVIMETRIC METHODS Determination 1.0 ± 0. 16. May 6. Cool.3 ml of amyloglucosidase solution. 1.24.4 Sequences in the analysis of total dietary fiber by the official AOAC method. 2001 2:56 PM 60 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.25 . 3RD EDITION Figure 3.1_fm Page 60 Tuesday. May 8.2387_ch3. The significance of protein as a component of dietary fiber. M. Rapid enzymatic assay of insoluble and soluble dietary fiber. Atherosclerosis... Wolever. 967... Agric. A. P. M. 1936. B. Hudson. H. M. 1981. A. M. and Benham. 965. 4. 21.. Sci. C. R. B. in Dietary Fibre: Current Developments of Importance of Health. Hallmer. Southgate. 16. Crude fibre. J.. B.. D. 1975. A. Southgate. Rep. Asp.. K. 17.. P. M. A. C.. Analytical problems in the determination of dietary fibre.-G.1_fm Page 61 Tuesday. 1461. G. Sandberg. 230. C.. Cereal Chem. Enzymatische Rohfaserbestimmung in Getreideprodukten.-G. A. 1975. John Libbey. Counc. 39. T. Sci. 115. D. Ed. Enzymatic determination of the indigestible residue of human food.. A. and van de Bovenkamp. 16. 1978. Hellendoorn. Katan. R. Res. 1981. Elchazly. Am. A. 26. Noordhoff. Trowell. vegetables and nuts. B. W.. 1978.. Techniques for measuring dietary fiber: principal aims of methods and a comparison of results obtained by different techniques. N. (G.. R. Asp. . A. 173. Leeds.5 Calculation of percentage dietary fiber from data obtained by official AOAC method. 979. Johansson. in The Analysis of Dietary Fiber in Food. Clin. The use of enzyme preparations in the crude fibre determination. O. 1983. 10. Eds. 1. 4.-G. M. Res. N-G. H. A. 28.-G. 12. Theander. Heaton. Hellendoorn.1. J. O. Mehl Brot. M.. and Theander. S. R. J.) Spec. Leb-ensm. O. McCance. and Shackleton. Food Agric. W. Some critical observations in relation to ‘dietary fibre’.. in The Analysis of Dietary Fiber in Food. G. The nutritive value of fruits. Studies on dietary fibres. 32. and Betschart..... 162. E. Determination of total dietary fiber by difference and of pectin by colorimetry of copper titration. J. 1974. New York. Forsch. T.. New York. 3.25 REFERENCES 1. Hallgren. London.. H. T.. Med. 329. Serv. T. and Jenkins. 8.. Lancet. Nutr.. B. Trowell. and Slagman. 1976.. May 8. Z.. 1980.2387_ch3. 1976. Trowell.. P. 9.-S. 1979. 476. Lancet. Weinstock. H... J. J. and Siljeström. The analysis of dietary fibre—the choices for the analyst. 31.. E. Critical evaluation of some suggested methods for assay of dietary fibre. Über eine biochemische Methode zum Bestimmen der Ballaststoffe und ihrer Komponenten in pflanzlichen Lebensmitteln. Definitions of fibre. 14. and Theander.. 9.. H.B. Widdowson..24. 1978. 490. 33. Food Chem.. Agric. 5.... M.. Dietary fibre redefined. 2. W. 15. 11. Getr.. dietary fibre and atherosclerosis. and Johansson. W. the methods for its determination and the current hypotheses for the explanation of its physiological action. 213. James. Näringsforskning. P.. Asp. and Thomas. Thomas. N. 1951. 217. C. H. Marcel Dekker. L. James. Eds. and Englyst. and Hasselblad. 13. Saunders. 6. Marcel Dekker. W.. 7.. 1981. C. 1. Gassull. 29.. 132. Voeding. 97. A. T.. 503. J.. D. Food Agric. Swed. 2001 2:56 PM ENZYMATIC GRAVIMETRIC METHODS 61 Figure 3. G. 138. and Aman. 29.. 1972. Unters. Clin. Minerals and phytate in the analysis of dietary fiber from cereals. H. Siljeström.. 21. I. II... Sci.. DeVries. New York. Off.. Anal. Nyman. T.. Anal. 252. T. Prosky. Sci. and Harland. Starch. 67. Eastwood. N. 22. in Furda... J. 74.. S. 2001 2:56 PM 62 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. . in Fiber in Human Nutrition. 1984. J. Cereal Chem. Minerals and phytate in the analysis of dietary fiber from cereals. Starch/Staerke... Schweizer. Cereal.. N.. 68... W. and Asp. Schweizer.. Forsch. Schweizer. 677. James. New Developments in Dietary Fiber. Fractionation and examination of biopolymers from dietary fiber. Chem. Englyst.. Pedersen. 1981... J. Asp. M. L. Suckow. and Schweizer. J. M.-G. 227. T. 1992... Hyg.. Kritchevsky. L. 31. 1985. T. S. I. 1983. Johansson. P... I. 1976. Schweizer. Cummings. in Dietary Fiber. J.. 1017. in The Analysis of Dietary Fiber in Food. gravimetric determination of total dietary fibre in foods. I....-G. and Amen. New York. 116. C. 34. J. 1981.. 19. Lewis. 1983. Bonfield.. D. and Asp.. 4.. W. T. Andersson. Dietary fiber analysis. Marcell Dekker. What is fiber?. M. and Koivistoinen. Englyst. Off. 41. 357.. Chem. T. 26.. Nutr. and Robertsson..-G. I. Mitt. 71. W. H. Springer. Kunita. A.. W. J. 61. J.. Schweizer. Analysis of dietary fibre. Sci. 163.. 1984. Siljeström. D. N. Chem. New York. DeVries. M. Assoc. 44... 27. 1988. Unters... and Schweizer. Nishimune. On the digestibility of starch in wheat bread. 1990. Asp. MES-TRIS buffer: collaborative study. Determination of total. in Dietary Fibre—A Component of Food—Nutritional Function in Health and Disease. Assoc. N. Ichikawa. 1984. F. Collaborative study for the enzymatic.. Plenum Press. Asp.. T. B. 35. Assoc. F. F. 40. T. Z. Furda. Furda.. C. L.. Starch and non-starch polysaccharides in some cereal foods. 4.-G... 1990. Van Soest.. Furda.. Eds. and Theander. 42. Björck. 1989.. 350. 57. W.. 41.-G. Lebensmitte-lunters. Frolich. 1990.. Characterization of resistant starch from autoclaved wheat starch. J.-G.. S. Off. 147. May 8. Chem. 1984. Sumimoto. Meuser. J.. 1992.. Assoc.. Delimitation problems in definition and analysis of dietary fiber. A. J. J. 33.. DeVries. F. 395. and Theander. J.. J. L. H. 1985. 613.. C. B. 37.. Assoc. Asp. B. 66. and Björck. N. and Nakahara.. I. J. Unters. Forsch. Nutrients excreted in ileostomy effluents after consumption of mixed diets with beans or potatoes.. 32. Food Agric..1_fm Page 62 Tuesday. 39.. Lebensm. Resistant starch formation during baking—effect of baking time and temperature and variations in the recipe. N. M. and insoluble dietary fiber in foods. Anal. and Brine. Laine... DeVedovo. and Eggum. New York. A. J.. 380. Anal.. T. and Besson. 181. Walter.. Spiller. Cereal Chem. Dietary fiber and resistant starch. P.-G. Off. J. and Kulikowski. 1988. T. and Venetz. 179. Lee..2387_ch3... 933. 75.. P. C. T. 28. V. 30.-G. Eds. Anderson. F. II. Langkilde. Enzymatic-gravimetric method. B. Dietary fibre in bread and corresponding flours—formation of resistant starch during baking. Assoc. Anal. 25. Urea enzymatic dialysis procedure for determination of total dietary fiber. Jeraci. 30. Chem. Siljeström. 5. N. 677. 567. F. Cereal Foods World. and Furda. Determination of total dietary fiber in foods and food products: collaborative study. Plenum. Anal. 23. 22. A. 1989. E. and Torsdottir. P. The Analysis of Dietary Fiber in Food. Prosky. Lebensm. W. Agric. Eds. T. 1977. 1986. and Harland... Doguchi. 3RD EDITION 18. Determination of insoluble soluble and total dietary fiber in foods and food products: inter-laboratory study. J.. W. N. 24. O. Asp. 37.. 1991. and Theander.. T.. Marcel Dekker. food products and total diets: interlaboratory study. Eds. 34... T. Varo. 20. Schweizer. Plenum Press.. Obst und Gemuse. J. T. Determination of total dietary fiber in foods.. Eliasson. T.. 36. Geb.. I. Prosky. 29... 49. Off. M.. S. W... 1979. and Cummings.. Schweizer. J. O. O.. Food Agric. M. Schweizer. I. Reimann. H. 38. 1044. 79. 61. 1. R.. G. J. R. Getreide Mehl Brot. Southgate. James.. P. J. 57. A. soluble... H. C. N.. Z. J. N. Asp. dietary fibre and sugars. F. T. and Anderson.. A nutritional classification of plant polysaccharides. Frølich.. Determination of total dietary fiber in Japanese foods. B. I. Siljeström. P. Yakusiji. Prosky. Effect of heat treatment on dietary fiber: interlaboratory study. R.. 1434. Simultaneous analysis of soluble and insoluble dietary fiber. Eur.. F.. N. Furda. F. I. Analytische Bestimmung von Ballaststoffen in Brot.. and Kingman. J. N. 1983. and DeVries. Assoc. S. Off.. 24. Asp.. P.-G.. Eds.-G. Edwards. Eds.. Chem. B. L. M. and Würsch.. R. 72. London. M.. Chem.. Mauron. Off. lipid and protein interference. However.1. some of this additional residue was hemicellulose9 and pectic substances.2 Detergent Analysis of Foods James B. This buffered AD system recovered more material. Because of these problems. 2001 6:37 PM CHAPTER 3.2387_ch3.12. A semimicro method has been developed for samples of 0. Typically.4 The neutral detergent (ND) system has undergone many changes from its inception for use with ruminant forages to human foods. depends on how the ADF will be used. Starch in low concentrations is solubilized in the ND. the detergent system has been used as a gravimetric method. it is normally totally digested.2. Lignin is usually present in low amounts and can be assayed in other ways.2.2.50 © 2001 by CRC Press LLC 63 . May 6. a preliminary extraction with ND before refluxing with AD has been suggested. primarily pectic substances and soluble hemicelluloses.3 Thus.2_fm Page 63 Sunday. free from hemicellulose.15 The decision to obtain an ADF of low fermentability or an ADF composed of cellulose and lignin.00+$1.6 The primary drawback of ND is that soluble dietary fiber.13 ADF has also been used to predict cell wall digestibility. and difficulty in filtration. A complex system of detergent extractions and subsequent hydrolysis and oxidation has been developed for a gravimetric component analysis of insoluble dietary fiber.14 These “celluloses” are easily fermentable and the linear hemicelluloses are of low fermentability.1 shows a flow diagram for this system.2 g.5 to 2.20 For very small samples.5 Fecal analysis by the detergent system is not complicated by microbial mass produced during colonic fermentation. Robertson and Peter J. and for this purpose removal of the insoluble hemicellulose and recovery of the AD-soluble cellulose may not be desirable. so hemicellulose should be removed to avoid the production of artifact lignin.17−19 One problem with a gravimetric method is that small samples or components in low concentration are difficult to weigh accurately. Baker8 suggested buffering the AD (pH 1.16 Figure 3.11 Pectin can also undergo Maillard reactions. a sugar component analysis may be useful. A major advantage of the ND procedure for dietary fiber analysis is that microbial material is soluble. Sugar analysis of the NDF and ADF has been progressing. The classification of fiber components and how they can be determined is displayed in the center of Figure 3.10 ADF has usually been used as a starting material for lignin determination. is not recovered. but 0-8493-2387-8/01/$0. the detergent system has a nutritional basis for the fractionation of food components.0) to reduce cellulose loss. However. Horvath The use of detergents to isolate plant cell walls of low nitrogen content from forages was proposed in 1963. The detergent systems are neutral solutions of sodium lauryl sulfate1 for cell wall analysis and an acidic medium containing cetyltrimethylammonium bromide for isolation of the less fermentable fraction of low hemicellulose content from forages. Other problems with the detergent systems include difficulty in starch removal.7 The original acid detergent (AD) system has not changed much for human food analysis. The problem of starch removal has been explored by many workers. May 8. and lignin values. it can be omitted if only forages are being analyzed. NDF decreased with other fractions varying nonuniformly. Drying above 60˚C or in a microwave oven can increase NDF. In most cases treatment during ND extraction with an amylase from Bacillus subtilis is adequate.2_fm Page 64 Tuesday. and central nervous system.14 Cellosolve (2-ethoxy ethanol) is an eye and mucous membrane irritant and could adversely affect the kidneys.29 Freeze-drying may be the optimum way to dry the sample for grinding. Detergent-stable proteases are available to hydrolyze protein for removal. 2-ethoxy ethanol should no longer be used in the neutral detergent solution. 3RD EDITION Figure 3. 2001 2:57 PM 64 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.26 This may not be desirable because β-glucans are included in the dietary fiber complex as water-soluble unavailable carbohydrates. in starch containing feeds and foods. They found that temperature.21 Other procedures have been compared by Mascarenhas-Ferreira et a1. ADF. Heller et al. not the length of incubation. However.28.6.) certain types (modified or retrograded) and high concentrations can lead to filtration problems and an overestimation of dietary fiber (as currently defined).2. Lipids and proteins interfere when in amounts that exceed the capacity of the reagents. but Marlett and Lee23 found little difference in results from these methods.25 Difficulty with filtration can also occur when other polysaccharides such as β-glucan are present. (The quantity of each fraction is measured as either the amount of residue or the weight loss after each procedure.24. Consequently. The limits are about 10% of dry matter for lipid and 30% for protein. a volume-for-volume substitution of cellosolve with triethylene glycol . The materials measured by weight loss are shown in brackets. The reagents used for detergent extractions have undergone important changes since 1970. Preextraction with ethanol or ether will remove lipids. β-glucanase has been used to remove these glucans. Heat-stable bacterial amylase might be the fastest and easiest.27 found that as particle size was decreased. The sample is usually dried before it is ground.22 Some studies suggested a mammalian or fungal enzyme and a protease. is most important. pregnant female. It has the potential to cause adverse reproductive effects in males and females and embryotoxic effects including teratogenesis in the offspring of the exposed. Since cellosolve was included in the original reagent because it facilitates the solution of starch. liver.1 Flow diagram for fiber analysis by the detergent system.2387_ch3. Both drying and particle size can affect the values. and cool-season grasses. 8. Off. Agric. Agricultural Handbook 379. 62. Slavin. 31. C.. Bossi. J. Goering. it is recommended that hydrolysis with 72% H2SO4 without the use of asbestos30 be carried out before lignin oxidation. Camb. Assoc. J. J. H. Washington. Department of Agriculture. Assoc. Eds. J. J.. Off. W. J.2_fm Page 65 Sunday. H. Pasture quality and ruminant nutrition. Baker... 471. Houston. Windham. and Himmelsbach D. E. Mason. S. 859. 1983. Dairy Sci. P. Chem.. I. 19.. H.3 If a detailed analysis of the ADF is done. 123. 1973. Belo. 31. 17. P. J. S. R. J. Off. Nutr.Z.. Sci. 2.. 11. A. Food Chem.. Wiggins. O. 54. J. 9. W. J. The permanganate solution is essentially unchanged.. Agric. K. 1980.14 Detailed instructions and procedures are described in Robertson and Van Soest16 and Van Soest et al. R. P. Int. Van Soest. Use of detergents in the analysis of fibrous feeds. 10. A. 137. Food Chem. Forage fiber analysis (apparatus.. Sci. Hemicellulose contamination of acid detergent residues and their replacement by cellulose residues in cell wall analysis. D. Barton.. P. 1978.. J.. J. 1983. has been shown to cause a significant loss of lignin21 and should be omitted in sequential analysis. Determination of plant cell-wall constituents. P. James. A.. J. Simonetti.. Use of detergents in the analysis of fibrous feeds. 369. 14. 16. T. Porrini. 25. E. S.. 1981. Agric. REFERENCES 1. 31.S.. 29. A.. 3. M. Van Soest. Agric. 18. A rapid method for the determination of fiber and lignin. 50. B. M. 1962. 467. A rapid method for the analysis of alimentary fiber. P. in The Analysis of Dietary Fiber in Foods.. Branch. Chem.. 477.. 1981. 99. Some observations on the distribution and origin of nitrogen in sheep feces. I. 13.. Modification of the neutral-detergent fiber procedure for cereals and vegetables by using alpha-amylase. D. and Amen. May 6. High pressure liquid chromatographic analysis of component sugars in neutral-detergent fiber for representative warm. and Marlett. 829. J... Decalin has been omitted from both detergent solutions to eliminate filtering problems and the consequent high fiber values. Br. procedures and some applications). G. R. U... and Ciappellano. 50.. B. 7. J... Pectic substance content of detergent-extracted dietary fibers. and Wine. 15. and Van Soest.. and Jenkins. New York... Agric. J.. 1963. Vercesi. J. 5. G. Food Chem. W.. J. Sulfite. J. Agric. J. K. Res.. C. F. Robertson. 825. Assoc. A. and Street. Nutr. 1970. Bailey. H. Agric. J. Cummings. Off.O. 4. Sci.. D. D. J. McQueen. Agric.. J.. New York. 370. T. J. 73. J. R.. 1970. and Ulyatt. R. W. and Nicholson. Evaluation of high-performance liquid chromatography for measurement of the neutral saccharides in neutral-detergent fiber. S. Gaillard. . D. R..25 A summary of total dietary fiber and detergent values is given in Table 2 of the Appendix. J. Eds.. and Hemken. P. B. P. 31. 1. Spiller. The detergent system of analysis and its application to human foods.. J. 46. The detergent system of fiber analysis. Van Soest.. II. 12. in Topics in Dietary Fiber Research.. Cutin can be determined as the residue lost upon ashing... reagents. IV. 59. 46. Agric. S. Camb. D. J. 1983.C. Van Soest. Relative susceptibility of forages to heat damage as affected by moisture temperature and pH. L. Anal. 7. 2001 6:37 PM DETERGENT ANALYSIS OF FOODS 65 produces identical analytical values.. W. Assoc. 1979. V. Chem. Monosaccharide composition of alcohol. 6.. 639. Carbohydrate and lignin composition of detergent-extracted residues from pasture grasses and legumes. The relationship between the cell-wall constituents of roughages and the digestibility of the organic matter. G. and De Lumen. Food Chem. Bittner. 56. Preparation of fiber residues of low nitrogen content. and Van Soest. 1977. Chem. Plenum Press.. Marcel Dekker. II. Digestion of pectin in the human gut and its effect on calcium absorption and large bowel function. Southgate.. M.. Cereal Chem. 360. Morrison. 591. Food Agric.and detergent-insoluble residues in maturing reed canary grass leaves. P. H. Determining fiber in cereals.. Anal. E. which was used to reduce the nitrogen content of the NDF. 1969. and Theander. P. J. 1963.2387_ch3. 41. J. Rep. 1979. B. W. Use of detergents in the analysis of fibrous feed.. Robertson. Jr. 1982. N. E. 1967. Goering. Testolin. 676. 13.. Rivers. J. J. and nonstarch polysaccharides in relation to animal nutrition. 1991. 25. 29.. Collaborative study of acid-detergent fiber and lignin. G. 42. A. J.. and Hackler.. Roth. 254 (Abstr. Chem. Off. J. W. T. 1977. 24. Dietary fiber estimation in concentrated feedstuffs. B. J. 19. New and improved procedure for neutral-detergent fiber. Royal Society of New Zealand. Anim. Beta-glucanase as an aid in measuring neutraldetergent fiber in barley kernels. May 6. Anim. N.. B.. 1). J. 1688. Sci. Mascarenhas-Ferreira. L. 1988. B. 27.. and Wallace. Hernandez. C. P. P. Dairy Sci. 2001 6:37 PM 66 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. J.)..1983. J. Feed Sci. 74. P. Jeraci. Kerstens. 9. 1977. 28.. Assoc. A. Cereal Chem. 1965. J. G.. Marlett. 3583. 3RD EDITION 20. 1983. A. 351 (Abstr. 1).. 58. 1977. Sci. J. Anim. Van Soest. J. P... Chem. in Fiber in Human and Animal Nutrition.. 69. S.. 1980. Food Sci.. lignocellulose and hemicellulose contents of selected foods determined by modified and unmodified Vap Soest procedures. C. Robertson. and Van Soest.. Lewis. 1981. G. A semi-micro neutral-detergent fiber method for cereal products. neutral detergent fiber. H. 22. 116. S. Robertson. Methods for dietary fiber.. Darrah. Wellington. Food Sci. Van Soest. 436. and Newman. J.. and Fick. Dietary fiber. J. Sills. R.. L. 26. 23. Van Soest. C.. Van Soest.. and Van Soest. 48.. Off. Technol. Agric.. L. Microwave treatment and heat damage artifacts in forages. Agron. and Lewis. C. J.. Study of effects of heating and drying on yield of fiber and lignin in forages. P. M. B. J... and Lee. J. 56.2387_ch3. Watts. V. 66 (Suppl.. H. P. E. Anal. Robertson.2_fm Page 66 Sunday.. . H.. 785.Z. 45. A. Assoc. N. and Gasp. III. Bulletin 10... Heller. J. 1973. B.. 30. 120. 21. Use of detergents in analysis of fibrous feeds. J. W. 245.. H. J. 45(Suppl... J. The study of several modifications of the neutral-detergent fibre procedure.). N. M. J. J.. 781. Dietary fiber: effect of particle size and pH on its measurement. health professionals.4–8 The main component (approximately 90%) of the cell walls of the plant foods in the human diet is carbohydrate. non-digestible oligosaccharides and dietary fiber can be used. and drying. such as heating. which justifies their use for food labeling for dietary fiber. collectively termed nonstarch polysaccharides (NSPs). but not immutable.2387_ch3. specifically.1).11.1–3 In 1998. since the NSP content of a food is not dependent on processing.3.50 © 2001 by CRC Press LLC 67 . Englyst. scientists.3 Dietary Fiber Analysis as Nonstarch Polysaccharides (NSPs) Hans Englyst and Geoffrey Hudson INTRODUCTION There is a steadily growing body of evidence that both the amounts and the types of dietary carbohydrates are important for the promotion and maintenance of public health. Committee on Medical Aspects of Food and Nutrition Policy (COMA). The classification and measurement scheme for dietary carbohydrates shown in Table 3. including nonstarch polysaccharides (see Table 3.10 NSP values provide a reliable marker of a naturally high-fiber diet. and the compilers of food tables and databases have argued strongly for the acquisition of specific analytical values for dietary carbohydrates. Additional groupings such as polyols. the widely used McCance and Widdowson Food Tables13 have contained values for dietary fiber measured as NSP. The NSP content of cooked foods can be calculated from analytical values for the raw ingredients.14. and this has repeatedly been recommended by the U.18 and their colleagues.1 has evolved in a virtually unbroken chronological sequence from the work of McCance and Widdowson. As a consequence. are in keeping with these requirements.3.16 then Southgate.12 For more than 15 years.15 The food industry is at liberty to use appropriate values from these tables without carrying out or commissioning its own analysis. and analytical procedures are available for all of the major categories of dietary carbohydrates so far identified.00+$1. more recently. proposing “that the analysis and labeling of dietary carbohydrates. May 6.K. be based on the chemical divisions recommended.17 and. including dietary fiber.3_fm Page 67 Sunday. FAO/WHO3 presented a list of recommendations on the role of carbohydrates in nutrition. 2001 6:39 PM CHAPTER 3.9. The term “dietary fiber” was coined as a description of the plant cell wall material that was recognized as the common characteristic of the unrefined plant foods that composed a naturally “high-fiber” diet. The classification scheme is complete. nutritionists. A cornerstone of this classification 0-8493-2387-8/01/$0. polysaccharides that do not have α-glucosidic linkages. resistant starch. cooling. for whatever purpose.” The methods that we have developed for the specific measurement of chemically identified carbohydrate fractions. provided the included components are clearly defined. 3. Physiological response depends on identity (Soluble in 80% ethanol. Fructose and lactose may. galactooligosaccharides.2387_ch3. 2001 6:39 PM 68 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Encapsulate and slow absorption of nutrients.and disaccharides Glucose. and sucrose rapidly digested. RS3 (retrograded amylose) Nonstarch (NSPs) Many different types of polysaccharides Plant cell-wall NSPs Main constituents: arabinose xylose. May 6.and disaccharides Comments Partly hydrolyzed starch. RS2 (resistant granules). lactose Sorbitol. Physiological effect largely unknown (Insoluble in 80% ethanol) α-Glucans Rapidly released glucose Slowly released glucose RS1 (physically inaccessible). in part. galactitol. ≤ 2 sugar units) Sugar alcohols Mono. sucrose. escape digestion and absorption in the small intestine. > 2 sugar units) Short-chain carbohydrates Starch Glucose. Rapidly digested in the small intestine. maltose. galactose glucose. pyrodextrins.1 Classification of Dietary Carbohydrates Main Components (Soluble in 80% ethanol. Slowly digested in the small intestine. maltitol Maltodextrins α-Glucans Resistant short-chain carbohydrates (non-digestible oligosaccharides) Fructo-oligosaccharides. maltose. The amounts added to foods are known and regulated . Normally included in the measurement of starch Escape digestion in the small intestine and are fermented to different extents. fructose. polydextrose Polysaccharides Rapidly digestible starch (RDS) Slowly digestible starch (SDS) Resistant starch (RS) Poorly absorbed in the small intestine. May reach the large intestine. uronic acids Other NSPs Many types of constituents The most abundant dietary carbohydrates. The three types of RS escape digestion in the small intestine and are fermented to different extents. (Soluble in 80% ethanol. mannose.3_fm Page 68 Sunday. Minor components of the human diet. Physiological effect largely unknown Escape digestion in the small intestine and are fermented to different extents. 3RD EDITION Table 3. ≤ 2 sugar units) Free sugars Mono. inositol. mannitol. Good marker for naturally high-fiber diets for which health benefits have been shown Food additives. Some may stimulate growth of bifidobacteria. 2001 6:39 PM DIETARY FIBER ANALYSIS AS NONSTARCH POLYSACCHARIDES (NSPS) 69 scheme is the measurement of chemically identified components. which. and whole grains. This property is related to the integrity of the cell wall structure.5 was related to the health benefits consequent upon the ingestion of a diet rich in unrefined plant foods and was specifically related to the plant cell wall. i.. § 343. A naturally high-fiber diet has been suggested to be protective against (1) coronary heart diseases (CHD). which is not a characteristic of RS. was seen at an early stage to be an important property of dietary fiber. as a considerable number of studies have failed to show a protective effect of dietary fiber. Therefore. A dietary supplement that bears a claim about dietary fiber and reduced risk of colorectal cancer will be subject to regulatory action as a misbranded food under 21 U. including free sugars and starch. non-digestible digosaccharides (NDO). and this justifies the use of NSP values for food labeling for dietary fiber. (2) diabetes. the plant cell walls encapsulate and moderate the release of nutrients.” The term dietary fiber was used first by Hipsley. collectively termed nonstarch polysaccharides (NSPs). FDA has determined that health claims relating dietary fiber and reduced risk of colorectal cancer are inherently misleading.). and thus the physiological effects of other nutrients. .e. May 6. vegetables.3. Drug. may be grouped into categories of specific nutritional importance. In unrefined and minimally processed foods. On the basis of its extensive review of the scientific evidence. or any fiber supplement. specifically.2387_ch3. and Cosmetic Act. polysaccharides that do not have α-glucosidic linkages. and some studies even suggest that fiber supplementation and RS may promote colon cancer. but also provides strong evidence that such a relationship does not exist. and that (2) lack of digestion in the small intestine is not a characteristic on which to base the definition of dietary fiber. the FDA has stated the following (10 October 2000): FDA has concluded from this review that the totality of the publicly available scientific evidence not only demonstrates lack of significant scientific agreement as to the validity of a relationship between dietary fiber and colorectal cancer. this is not the case for colon cancer. those rich in fruit. fate. It is clear that (1) the material measured as dietary fiber must be a reliable marker for the type of naturally high-fiber diet for which benefit to health has been shown. However. The plant cell wall encapsulates and thus controls the release of nutrients. from the plant tissue and thereby influences the glycemic response. and (2) at least some of the benefits to health may reflect the fact that such diets tend to be rich in vitamins. and physiological properties of the various types of dietary carbohydrates shown in Table 3. NSPs therefore. There is convincing evidence and a general consensus that (1) naturally high-fiber diets.8 The dietary fiber hypothesis. is convincing. This ability to influence digestion and absorption.C. The other main properties associated with dietary fiber were its abilities to increase fecal bulk and reduce transit time. including sugar and starch. on the basis of current knowledge of the relation between dietary carbohydrates and health. Burkitt and Trowell. and (3) colon cancer. The use of such health claims is therefore prohibited by the Federal Food. are beneficial to health. The weight of the evidence for a health claim about dietary fiber and colorectal cancer is outweighed by the evidence against such a claim. CHD and diabetes. The measurement of NSP is described here. The information on the composition.3_fm Page 69 Sunday.C. which was put forward by Cleave.1 is useful when considering the definition and measurement of dietary fiber. who recommended the eating of wholemeal bread for its “salutary effects upon the bowels. and are often low in fat. provide a reliable marker of a naturally high-fiber diet for which benefit to health is shown. The main component (approximately 90%) of the cell walls of the plant foods in the human diet is carbohydrate. The link between the ingestion of unprocessed foods and good health is chronicled as far back as Hippocrates (4th century B. The evidence for the first two. minerals.S. and antioxidants. 2387_ch3.3_fm Page 70 Sunday, May 6, 2001 6:39 PM 70 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION INTERNATIONAL COLLABORATIVE TRIALS MAFF-organized trials. The Englyst procedure has been the subject of a series of international collaborative trials organized by the U.K. Ministry of Agriculture, Fisheries & Food. In the MAFF IV study,19 37 laboratories from 11 countries took part to compare the accuracy and precision of the Englyst gas-liquid chromatographic (GLC) and colorimetric procedures. This series of trials culminated in the publication of the GLC and the colorimetry techniques as MAFF-approved methods.20,21 Certification of Reference Materials EC-organized trials. As the result of a large international trial of methodology, following rigorous study of stability of the test materials, five BCR-certificated reference materials (CRMs) are available for the Englyst GLC and colorimetry NSP procedures:22 (1) dried haricot bean powder, CRM 514; (2) dried carrot powder, CRM 515; (3) dried apple powder, CRM 516; (4) full fat soya flour, CRM 517; and (5) dried powdered bran breakfast cereal, CRM 518. These CRMs can be used to check the performance of the analytical method and as quality control of analytical measurements for nutritional labeling. Methods and Principles of Measurement The Englyst procedure measures dietary fiber as NSP, using enzymatic chemical methods, and has evolved from the principles laid down by McCance and Southgate. Starch is completely removed enzymatically, and NSPs are measured as the sum of the constituent sugars released by acid hydrolysis. The sugars may be measured by gas–liquid chromatography (GLC) or by high-pressure liquid chromatography (HPLC) to obtain values for individual monosaccharides, or a single value for total sugars may be obtained by colorimetry. Values may be obtained for total, soluble, and insoluble NSP, and a small modification allows cellulose to be measured separately. The Englyst procedure allows measurement of total, soluble, and insoluble NSP in plant food products within an 8-hour working day using the colorimetric endpoint, or within 1.5 working days with the chromatography procedures. Preparation of the hydrolyzate for analysis is virtually a single-tube procedure, and no special skill or equipment is needed for the colorimetric version. The procedure as described here provides the following options: (1) GLC procedure that measures NSP as the sum of neutral sugars obtained by GLC and uronic acids measured separately; (2) HPLC procedure: measures NSP as the sum of neutral sugars and uronic acids; (3) colorimetric procedure that measures NSP as reducing sugars; and (4) colorimetric procedure with separate measurement of uronic acids. PRINCIPLE 1. Dry/defat sample if necessary 2. Disperse (DMSO) and hydrolyze starch enzymatically 3. Precipitate NSP in acidified aqueous ethanol 4. Disperse and hydrolyze NSP with sulfuric acid 5. Measure released constituent sugars by: Option 1 Option 2 Option 3 * GLC* HPLC* Colorimetry** Values for individual constituent sugars of insoluble and total NSP. Soluble NSP calculated as the difference between insoluble and total NSP. ** Single values for insoluble and total NSP. Soluble NSP calculated as the difference between insoluble and total NSP. 2387_ch3.3_fm Page 71 Sunday, May 6, 2001 6:39 PM DIETARY FIBER ANALYSIS AS NONSTARCH POLYSACCHARIDES (NSPS) 71 THE PROCEDURE Apparatus and Reagents High-purity reagents and distilled, deionized water, or water of equivalent purity, should be used throughout the method. Reagents Common to the GLC, Colorimetric, and/or HPLC Procedures All sugars used for standards should be dried to constant mass under reduced pressure with phosphorus pentoxide before use. Acidified ethanol, absolute and 85% (v/v). Add 1 mL of 5 M hydrochloric acid per liter of ethanol. Dimethylphenol solution. Dissolve 0.1 g of 3,5-dimethylphenol in 100 mL of glacial acetic acid. Dimethylsulfoxide (DMSO). Enzyme solution I. Take 2.5 mL of heat-stable amylase (EC 3.2.1.1: Termamyl), make to 200 mL with pre-equilibrated sodium acetate buffer, mix, and keep it in a 50°C waterbath. Prepare the solution immediately before use. Enzyme solution II. Take 1.2 g of pancreatin into a 50-mL tube, add 12 mL of water, vortex-mix initially, and then mix for 10 min with a magnetic stirrer. Vortex-mix again, then centrifuge for 10 min. Take 10 mL of the (cloudy) supernatant, add 2.5 mL of pullulanase (EC 3.2.1.41: Promozyme), and vortex-mix. Prepare the solution immediately before use and keep it at room temperature. Glass balls, 2.5 to 3.5 mm diameter. Sand, acid-washed, 50–100 mesh. Sodium acetate buffer, 0.1 M, pH 5.2. Dissolve 13.6 g of sodium acetate trihydrate and make to 1 L with water. Adjust to pH 5.2 with 0.1 M acetic acid. To stabilize and activate enzymes, add 4 mL of 1 M calcium chloride to 1 L of buffer. Sodium chloride–boric acid solution. Dissolve 2 g of sodium chloride and 3 g of boric acid in 100 mL of water. Sodium phosphate buffer, 0.2 M, pH 7. Adjust 0.2 M Na2HPO4 to pH 7 with 0.2 M NaH2PO4. Sulfuric acid, 12 M. Sulfuric acid, 2 M. Add 5 mL of 12 M sulfuric acid to 25 mL of water. Allow to cool to room temperature before use. Sulfuric acid, 2.4 M. Add 5 mL of 12 M sulfuric acid to 20 mL of water and mix. Reagents Used Only in the GLC Procedure Acetic anhydride. Ammonium–sodium tetrahydroborate solution. A solution of 6 M ammonium hydroxide, containing 200 mg/mL of sodium tetrahydroborate [NaBH4]. Prepare immediately before use. Benzoic acid, saturated. Prepare a saturated solution of benzoic acid at room temperature. Add 0.5 g of benzoic acid per 100 mL of water; some benzoic acid should remain undissolved after overnight stirring. The saturated solution is stable at room temperature for long periods. Bromophenol blue solution, 0.4 g/L. GLC internal standard solution, 1 mg/mL. Weigh to the nearest 1 mg: 500 mg of allose. Dissolve in water, add 250 mL of saturated benzoic acid, and make to 500 mL with water to give a 1 mg/mL solution. The solution is stable at room temperature for several months. Glacial acetic acid. GLC stock sugar mixture. Weigh to the nearest 1 mg: 0.52 g of rhamnose, 0.48 g of fucose, 4.75 g of arabinose, 4.45 g of xylose, 2.3 g of mannose, 2.82 g of galactose, 9.4 g of glucose, and 2.79 g of galacturonic acid (or 3.05 g of galacturonic acid monohydrate). Dissolve together in water, add 500 mL of saturated benzoic acid, and make to 1 L with water. The solution is stable at room temperature for several months. 1-Methylimidazole. 2387_ch3.3_fm Page 72 Sunday, May 6, 2001 6:39 PM 72 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Reagents Used Only in the HPLC Procedure HPLC neutral sugars internal standard solution. Weigh to the nearest 1 mg: 10,000 mg of deoxygalactose. Add 10 mg of thiomersal (C2H5·Hg·S·C6H4·COONa) as preservative and make to 1 L with water. HPLC neutral sugars stock sugar mixture. Weigh to the nearest 1 mg: 0.52 g of rhamnose, 0.48 g of fucose, 4.75 g of arabinose, 4.45 g of xylose, 2.3 g of mannose, 2.82 g of galactose, and 9.4 g of glucose. Dissolve together in water, add 10 mg of thimerosal as preservative, and make to 1 L with water. Store at 4°C. HPLC uronic acids internal standard solution. Weigh to the nearest 1 mg: 0.0454 g of mannuronic acid lactone. Dissolve in water, add 1 mg of thimerosal as preservative, and make to 100 mL with water. Store at 4°C. HPLC uronic acids stock sugar mixture. Weigh to the nearest 1 mg: 0.0930 g of galacturonic acid (or 0.102 g of galacturonic acid monohydrate) and 0.0233 g of glucuronic acid. Dissolve together in 2 M sulfuric acid, and make to 100 mL with 2 M sulfuric acid. Store at 4°C. Pectinase solution (EC 3.2.1.15: Novo Nordisk). Add 25 volumes of water to 1 volume of pectinase. Reagents Used Only in the Colorimetric Procedure Color reagent. Dissolve 10 g of 3,5-dinitrosalicylic acid and 300 g of sodium/potassium tartrate in approx. 300 mL of water plus 400 mL of 1 M NaOH. Dissolve by stirring (overnight) and make to a final volume of 1 L with water. Sparge for 10 min with helium or nitrogen, or degas using an ultrasonic bath. Store in well-capped opaque bottles, and keep for 2 days before use. The reagent is stable at room temperature for at least 6 months. Dimethylglutaric acid solution, 0.5 M. Add 98.5 g of 5 M NaOH to 8.0 g (weighed to the nearest 1 mg) of dimethylglutaric acid (DMG) and make to 100 mL with water at room temperature. [Note: since the purity of DMG varies between lot numbers, it is essential that the pH is checked. A portion of the DMG solution should be diluted 1:1 (v/v) with 2 M sulfuric acid; the pH should be 3.75 (±0.15) at room temperature. If the pH is greater than 3.9, add 1 mL of water to the stock solution and check the pH again. If the pH is less than 3.9, add 1 mL of 5 M NaOH to the stock solution and check the pH again. Repeat as necessary.] Color stock sugar mixture. Make the color stock sugar solution by weighing to the nearest 1 mg: 10.185 g of arabinose, 5.145 g of glucose, and 2.16 g of galacturonic acid (or 2.36 g of galacturonic acid monohydrate). Dissolve together in water, add 500 mL of saturated benzoic acid, and make to 1 L with water. The solution is stable at room temperature for several months. Pectinase solution (EC 3.2.1.15: Novo Nordisk). Add 9 volumes of water to 1 volume of pectinase. Sample Preparation All samples must be finely divided (to pass a 0.5-mm mesh) so that representative subsamples may be taken. Foods with a low water content (<10 g per 100 g of sample) may be milled, and foods with a higher water content may be homogenized wet or milled after freeze-drying. Analysis of three subsamples, A, B, and C, allows separate values to be obtained directly for total NSP, insoluble NSP, and cellulose, respectively. Soluble NSP is determined as the difference between total and insoluble NSP. Portions A and B are treated identically throughout the procedure, except in steps 3 and 4. The third portion, C, is needed only if a separate value for cellulose is required; carry it through steps 1, 2, and 3 of the procedure; then go to step 5.2. Step 1.1 Weigh, to the nearest 1 mg, between 50 and 1000 mg, depending on the water and NSP content of the sample (to give not more than 300 mg of dry matter; e.g., 300 mg is adequate for most dried foods but smaller amounts should be used for bran and purified fiber preparations), into 50–60 mL screw-top glass tubes. Add 300 (±20) mg of acid-washed sand and approximately 15 glass balls to each. If the sample is dry (85–100 g of dry matter per 100 g of sample) and contains less than 10 g of fat per 100 g of sample, proceed to step 2.1; otherwise, go to step 1.2. All analyses should be done in duplicate. 2387_ch3.3_fm Page 73 Sunday, May 6, 2001 6:39 PM DIETARY FIBER ANALYSIS AS NONSTARCH POLYSACCHARIDES (NSPS) Step 1.2 Add 40 mL of acetone, cap the tubes, and mix several times over 30 min. Centrifuge at 1000g for 10 min to obtain a clear supernatant and remove as much of the supernatant liquid as possible without disturbing the residue. Vortex-mix vigorously to ensure that the residue is dispersed thinly around the bottom 5 cm of the tube. Place the rack of tubes in a pan of water at 75°C in a fume-cupboard. Remove the tubes singly and vortex-mix vigorously at frequent intervals until the tubes and residues are dry. Isolation and Hydrolysis of NSP Dispersion and Enzymatic Hydrolysis Pre-equilibrate sufficient acetate buffer at 50°C (8 mL required per sample). Step 2.1 Add 2 mL of dimethylsulfoxide (DMSO) to the dry sample, cap the tube, and immediately mix the contents using a vortex-mixer, treating each tube in turn. It is essential that all the sample is wetted and no material is encapsulated or adhering to the tube wall before proceeding. When DMSO has been added to all the tubes, vortex-mix three or four times for 5 min. Vortex-mix and immediately place 2 tubes into a boiling waterbath. Remove after 20 s, vortex-mix, and immediately replace the tubes in the bath. Repeat this for subsequent pairs of tubes until all the tubes are in the bath; leave them for 30 min from that time. During this period, prepare enzyme solutions I and II (see Step 4.1; the volumes given are suitable for 24 samples). Step 2.2 Remove one tube at a time, vortex-mix, uncap, and immediately add 8 mL of enzyme solution I (kept at 50°C), cap the tube, vortex-mix thoroughly — ensuring that no material adheres to the tube wall — and replace it in the boiling waterbath. Leave the tubes there for 10 min, timed from the last addition of enzyme. Transfer the rack of tubes to the 50°C waterbath. After 3 min, add 0.5 mL of enzyme solution II to each tube and mix the contents thoroughly to aid distribution of the enzyme throughout the sample. Replace the tubes in the 50°C waterbath and leave them there for 30 min. Mix the contents of each tube continuously or after 10 min, 20 min, and 30 min. Transfer the rack of tubes to the boiling waterbath and leave them there for 10 min. Precipitation and Washing of the Residue for Measurement of Total NSP Only sample portion A is given this treatment. Step 3.1 Cool the samples by placing in ice water. Add 0.15 mL of 5 M hydrochloric acid and vortexmix thoroughly 2 or 3 times for 5 min with samples being replaced in the ice water. Add 40 mL of acidified absolute ethanol and mix well by repeated inversion, then leave in ice water for 30 min. Centrifuge at 1500 g for 10 min to obtain a clear supernatant liquid. Remove as much of the supernatant liquid as possible, without disturbing the residue, and discard it. Step 3.2 Add approximately 10 mL of acidified 85% ethanol to the residue and vortex-mix. Make to 50 mL with acidified 85% ethanol, mix thoroughly by repeated inversion. Centrifuge and remove the supernatant liquid as above. Repeat this stage using 50 mL of absolute ethanol. Step 3.3 Add 30 mL of acetone to the residue and vortex-mix thoroughly to form a suspension. Centrifuge and remove the supernatant liquid as described in step 3.1. Step 3.4 Place the rack of tubes in a pan of water at 75°C in a fume-cupboard or a TurboVap (Zymark Ltd.) at 65°C. Remove the tubes singly and vortex-mix vigorously at frequent intervals, to ensure that the residue in each tube is finely divided, until each tube and residue appears dry. Place the rack of tubes in a fan oven at 80°C for 10 min to remove any last traces of acetone. It is essential that the residues and tubes are completely free of acetone. Extraction and Washing of the Residue for Measurement of Insoluble NSP Only sample portion B is given this treatment. Step 4.1 After the treatment with enzymes in step 2, add 40 mL of sodium phosphate buffer. Place the capped tubes in a boiling waterbath for 30 min. Mix continuously or a minimum of three times 73 2387_ch3.3_fm Page 74 Sunday, May 6, 2001 6:39 PM 74 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION during this period. Remove the tubes and equilibrate to room temperature in water. Centrifuge and remove the supernatant liquid as described in step 3.1. Step 4.2 Add approximately 10 mL of water and vortex-mix. Make to approximately 50 mL with water and mix well by repeated inversion. Centrifuge and remove the supernatant liquid as described in step 3.1. Repeat this stage using 50 mL of absolute ethanol. Proceed as described for steps 3.3 and 3.4. Acid Hydrolysis of the Residue from Enzymatic Digestion Step 5.1 Add 5 mL of 12 M sulfuric acid to one tube and immediately vortex-mix vigorously; ensure that all the material is wetted. Repeat this for each tube in turn. Once the acid has been added to all the tubes, vortex-mix again and place all the tubes into a waterbath at 35°C. Leave the tubes at 35°C for 30 min with vigorous vortex-mixing after 5, 10, and 20 min to disperse the cellulose. Add 25 mL of water rapidly and vortex-mix. Place into a boiling waterbath and leave for 1 hour, timed from when boiling recommences; mix after 10 min. Cool the tubes in tap water. Step 5.2 A modification allowing the separate measurement of cellulose and non-cellulosic polysaccharides (NCPs). To portion C, after steps 1 through 3, add 30 mL of 2 M sulfuric acid, and mix. Place in a boiling waterbath and leave for 1 hour, timed from when boiling recommences, stirring continuously or after 10 min. The value for cellulose is obtained as the difference between glucose (measured by GLC or by glucose oxidase) for sample portions A and C. NCP is calculated as the difference between total NSP and cellulose. Breaks in the Procedure The procedure may be halted at the following stages. (1) After precipitation, washing, and drying the starch-free residue in steps 3 and 4. The residue may be stored for long periods. (2) After the hydrolysis with 2 M sulfuric acid in step 5. The hydrolysate may be kept at 4°C for 48 hours. Determination of Constituent Sugars by GLC This assay includes the measurement of neutral sugars by GLC and the separate measurement of uronic acids by colorimetry. Measurement of Neutral NSP Constituents by GLC Preparation of the standard sugar mixture. Mix 1.0 mL of the GLC stock sugar solution and 5 mL of 2.4 M sulfuric acid. Treat 2 × 1.0 mL of this standard sugar mixture for calibration in parallel with the hydrolysates from step 5 of the procedure. Prepare the alditol acetate derivatives for chromatography as follows. Add 0.50 mL of GLC internal standard (1 mg/mL allose) to 1.0 mL of the cooled hydrolysates from step 5 and to 2 × 1 mL of the standard sugar mixture; vortex-mix. Place the tubes in ice water, add 0.4 mL of 12 M ammonium solution, and vortex-mix. Test that the solution is alkaline (add a little more ammonium solution if necessary, but replace the ammonium solution if more than 0.1 mL extra is required), then add approximately 5 µL of the antifoam agent octan-2-ol and 0.1 mL of the ammonium–sodium borohydride solution; vortex-mix. Leave the tubes in a heating block or in a waterbath at 40°C for 30 min, then remove and add 0.2 mL of glacial acetic acid, and mix again. Remove 0.5 mL to a 30 mL glass tube and add 0.5 mL of 1-methylimidazole and 5 mL of acetic anhydride. Vortex-mix, then leave the tubes for 10 min for the reaction to proceed (the reaction is exothermic and the tubes will become hot). Add 0.9 mL of absolute ethanol, vortex-mix, and leave for 5 min. Add 10 mL of water, vortex-mix, and leave for 5 min. Add 0.5 mL of bromophenol blue solution. Place the tubes in ice water and add 5 mL of 7.5 M potassium hydroxide; a few minutes later add a further 5 mL of 7.5 M potassium hydroxide, cap the tubes, and mix by inversion. Leave until the separation 2387_ch3.3_fm Page 75 Sunday, May 6, 2001 6:39 PM DIETARY FIBER ANALYSIS AS NONSTARCH POLYSACCHARIDES (NSPS) 75 into two phases is complete (10 to 15 min) or centrifuge for a few minutes. Draw part of the upper phase into the tip of an automatic pipette; if any of the blue lower phase is included, allow it to separate, then run it out of the tip before transferring a portion of the upper phase alone to a small (auto-injector) vial. Inject 0.5 to 1 µL of the alditol acetate derivatives. GLC Conditions Injector temperature, 275°C; column temperature, 220°C; detector temperature, 275°C; carrier gas, helium; flow rate, 8 mL/min. Under these conditions, a GLC chromatograph fitted with flame ionization detector, auto-injector, and computing integrator, using a Supelco SP-2330 wide-bore capillary column (30 m × 0.75 mm) or a Supelco SP-2380 wide-bore capillary column (30 m × 0.53 mm), will allow accurate determination of the individual sugars in the GLC standard sugar mixture within 8 min. Carry out conventional GLC measurement of the neutral sugars. At the beginning of each batch of analyses, equilibrate with the isothermal elution conditions for at least 1 hour. Do several calibration runs to check that the response factors are reproducible. Table 3.3.2 Calibration Ratios for GLC Standard Sugar and GLC Internal Standard Combination Sugar Rhamnose Fucose Arabinose Xylose Mannose Galactose Glucose Allose (Int. std.) Actual (mg/mL) Recovery (%) Apparent (mg/mL) Calibration Ratio 520 480 4750 4450 2300 2820 9400 52 96 95 89 92 94 94 1000 500 5000 5000 2500 3000 10,000 1 0.5 5 5 2.5 10 3 In Table 3.3.2, the calibration ratios are shown for the combination of the GLC standard sugar mixture and GLC internal standard (allose). The Actual column shows the amount of each sugar in the mixture, and the Apparent column shows the values to be used for calibration, taking into account the recovery of NSP constituents. The calibration ratio column gives the ratio of sugars to the internal standard after the addition of allose to the standard sugar mixture, as described in the text. (The experimental evidence for the recovery values has been published and is discussed in detail below.) Calculation of Neutral Sugars The amount of each individual sugar (expressed as grams of polysaccharide per 100 g of sample) is calculated as A ( t ) × W ( i ) × 100 × R ( f ) × 0.89 Sugar = -----------------------------------------------------------------------------A(i) × W (t) (3.3.1) where A(t) and A(i) are the peak areas of the sample and the internal standard, respectively; W(i) is the weight (in mg; here 15: total hydrolysate 30 ml × 0.5 mg allose) of the internal standard; W(t) is the weight (mg) of the sample; R(f) is the response factor for individual sugars obtained from the calibration run with the sugar mixture and internal standard (allose) treated in parallel with the samples; and 0.89 is the factor for converting experimentally determined values for monosaccharides to polysaccharides. 2387_ch3.3_fm Page 76 Sunday, May 6, 2001 6:39 PM 76 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Measurement of Uronic Acids by Colorimetry Make the standard solutions as follows. The GLC standard sugar mixture in 2 M sulfuric acid contains, for the purpose of calibration, 500 µg/mL of galacturonic acid. To prepare the uronic acid standard solutions, take 0.5 mL, 2.0 mL, and 3.0 mL of this sugar mixture into separate tubes, and make to 10 mL with 2 M sulfuric acid to give standards of 25, 100, and 150 µg/mL of galacturonic acid. Place into separate tubes (40–50 mL capacity) 0.3 mL of blank solution (2 M sulfuric acid), 0.3 mL of each of the standard solutions, and 0.3 mL of the sample hydrolysates, diluted if necessary (with 2 M sulfuric acid) to contain no more than 150 µg/mL of uronic acids (e.g., no dilution for flour, 1:2 for bran, 1:5 for most fruits and vegetables). Add 0.3 mL of sodium chloride–boric acid solution and mix. Add 5 mL of concentrated sulfuric acid and vortex-mix immediately. Place the tubes into a heating block at 70°C for 40 min. Remove the tubes and cool to room temperature in water (the tubes may be kept in the water for up to 1 hour). Add 0.2 mL of dimethylphenol solution and vortex-mix immediately. After 15 min measure the absorbance at 400 nm and at 450 nm in the spectrophotometer against the blank solution. The timing for measurement of the absorbance of standards and samples should be identical. In practice, this is achieved by adding the color reagent at 1-min intervals. Subtract the absorbance reading at 400 nm from that at 450 nm, to correct for interference from hexoses. A straight line should be obtained if the differences in absorbance for the standards are plotted against concentration. Only the 100 µg/mL standard is required for routine analysis, and it may be kept at 5°C for several weeks. Calculation of Uronic Acids The amount of uronic acids (expressed as grams of polysaccharide per 100 g of sample) is calculated as A ( t ) × V ( t ) × D × C × 100 × 0.91 Uronic acids = --------------------------------------------------------------------------------A(s) × W (t) (3.3.2) where A(t) is the difference in absorbance of the sample solution; V(t) is the total volume of sample solution (mL, here 30); D is the dilution of the sample solution; C is the concentration of the standard (here 0.1 mg/mL); A(s) is the difference in absorbance of the 100 µg/mL standard; W(t) is the weight (mg) of the sample; and 0.91 is the factor for converting experimentally determined values for monosaccharides to polysaccharides. Calculation of NSP The amount of total, soluble, and insoluble NSP (in grams per 100 g of sample) is calculated as Total NSP Insoluble NSP Soluble NSP = Neutral sugars calculated for portion A + Uronic acids calculated for portion A = Neutral sugars calculated for portion B + Uronic acids calculated for portion B = Total NSP – Insoluble NSP It is recommended that a sample be taken for determination of the dry matter as the loss in weight after overnight incubation at 104°C. Results may then be expressed as grams of polysaccharides per 100 g of dry matter. 2387_ch3.3_fm Page 77 Sunday, May 6, 2001 6:39 PM DIETARY FIBER ANALYSIS AS NONSTARCH POLYSACCHARIDES (NSPS) 77 Breaks in the GLC Procedure The procedure may be halted at either of the following stages: (1) after acidification of the reduced samples (see Determination of Constituent Sugars by GLC section); the samples may be stored at room temperature for 2 to 3 days; or (2) the acid hydrolysate from Step 5 may be kept at 5°C for several weeks before the measurement of uronic acids. Determination of Constituent Sugars by HPLC This assay includes dilution of one subsample of the hydrolysate and direct measurement of the neutral sugars, and the separate measurement of uronic acids in a second subsample of the hydrolysate subjected to treatment with pectinase. Measurement of Neutral Sugars by HPLC Preparation of the HPLC standard sugar mixture. Mix 1.0 mL of the HPLC stock sugar mixture and 5 mL of 2.4 M sulfuric acid. Treat 2 × 0.15 mL of this standard sugar mixture for calibration of HPLC in parallel with the hydrolysates from step 5 of the procedure. To 0.15 mL of hydrolysate or the HPLC standard sugar mixture, add 5 mL of the neutral sugars internal standard solution (deoxygalactose) and mix well. Inject 25 µL for analysis. HPLC Conditions A Dionex model PAD 2 detector may be used with the following pulse potentials and durations: E1 = 0.05 V (t1 = 300 ms); E2 = 0.60 V (t2 = 120 ms); E3 = –0.60 V (t3 = 60 ms), and with a response time of 1 s and the detector output set at 1000 nA. A Dionex AG-5 guard column, an inert high-pressure valve, and a CarboPac PA-1 column are placed in series. The AG-5 column and inert high-pressure valve are used to retain and bypass sulfate ions around the analytical column. Elute with the following: 23% (v/v) solution 1 (0.020 M NaOH) from 0 to 3.5 min; a gradient from 23% to 1% (v/v) solution 1 from 3.5 to 4.5 min; and 1% (v/v) solution 1 from 4.5 to 30 min at a flow rate of 1 mL/min. Re-equilibrate with the starting conditions for at least 6 min between runs. Add 0.30 M NaOH at a flow rate of 0.5 mL/min to the column effluent before the PAD cell to minimize baseline drift and increase the analytical signal. Saturate the eluent with helium (Dionex Eluent De-gas module) to minimize CO2 absorption. Under the conditions described here, sulfate ions are retained for 80 s on the AG-5 guard column, and column switching (via the inert high-pressure valve) is applied after 60 s to prevent sulfate ions from reaching the analytical column. Sulfate ions are purged from the guard column within 19 min, well within the total run time. To regenerate the PA-1 column at the end of the chromatographic run, wash with 0.10 M NaOH–0.60 M sodium acetate for 1 hour at a flow rate of 1 mL/min. Then wash with 1 M NaOH (1 mL/min) for 1 hour but, to avoid contamination of the internal reference solution, do not allow this solution to pass through the detector. Table 3.3.3 Calibration Ratios for HPLC Standard Sugar and HPLC Internal Standard Combination Sugar Rhamnose Fucose Arabinose Xylose Mannose Galactose Glucose Deoxygalactose (Int. std.) Actual (mg/mL) Recovery (%) Apparent (mg/mL) Calibration Ratio 520 480 4750 4450 2300 2820 9400 52 96 95 89 92 94 94 1000 500 5000 5000 2500 3000 10,000 1 0.5 5 5 2.5 3 10 2 2387_ch3.3_fm Page 78 Sunday, May 6, 2001 6:39 PM 78 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Table 3.3.3 shows the calibration ratios for the combination of the standard sugar mixture and deoxygalactose and the HPLC internal standard; the Actual column shows the amount of each sugar in the mixture, and the Apparent column shows the values to be used for calibration, taking into account the recovery of NSP constituents. The amount of each individual sugar (in grams per 100 g of sample) is calculated as Sugar = A ( t ) × W ( i ) × 100 × R ( f ) × 0.89 -----------------------------------------------------------------------------A(i) × W (t) (3.3.3) where A(t) and A(i) are the peak areas of the sample and the HPLC internal standard, respectively; W(i) is the weight of the internal standard if added to the whole sample (in mg, here 25); W(t) is the weight of the sample (mg); R(f) is the response factor for individual sugars obtained from a calibration run with the sugar mixture treated in parallel with the samples; and 0.89 is the factor for converting experimentally determined values for monosaccharides to polysaccharides. It is recommended that a sample be taken for the analysis of the dry matter as the material remaining after overnight incubation at 104°C. Results may then be expressed as grams of polysaccharides per 100 g of dry matter. Measurement of Uronic Acids by HPLC To 0.5 mL of hydrolysate or calibration mixture, add 0.5 mL of the HPLC uronic acid internal standard solution (mannuronic acid lactone), 0.5 mL of dimethylglutaric acid solution (8 g/100 mL), and 2.1 mL of 1 M NaOH. After vortex-mixing, the pH must be between pH 3.5 and 4.0. (If the pH is not correct, prepare fresh DMG solution and repeat). Add 0.1 mL of pectinase solution, vortex-mix, and place the tubes into a waterbath at 50°C for 20 min. Cool the tubes, remove 0.5 mL, and add 0.1 mL of phenol red indicator solution (1 mg/mL); add sufficient (approximately 2 mL) freshly prepared 0.017 M NaOH until the pH is between 7 and 8. Inject 25 µL onto the chromatography column. Baseline separation of galacturonic acid (GalA), glucuronic acid (GlcA), and mannuronic acid is achieved isocratically within 13 min using 25% solution 2 (0.10 M NaOH, 0.6 M sodium acetate) and 75% water at a flow rate of 1.0 mL/min. Use the following ratios for the calibration mixture for the calculation of the response factors: 0.5 for GalA, 0.125 for GlcA, and 0.25 for mannuronic acid to take into account losses during hydrolysis. The values for the calculation of response factors take into account 7% loss of both GalA and GlcA and the hydrolysis of mannuronic acid lactone to mannuronic acid, as described above. The amount of uronic acids (in grams per 100 g of sample) is calculated as Uronic acids = A ( t ) × W ( i ) × 100 × R ( f ) × 0.91 -----------------------------------------------------------------------------A(i) × W (t) (3.3.4) where A(t) and A(i) are the peak areas of the test and the internal standard, respectively; W(i) and W(t) are the weight of the internal standard (in mg; here 15) and the mass of the test sample (mg), respectively; R(f) is the response factor for individual uronic acids obtained from the calibration mixture; and 0.91 is the factor for converting the experimentally determined values for monosaccharides to polysaccharides. Determination of Constituent Sugars by Colorimetry Preparation of the Standard Sugar Mixture Take 0.5 mL of the colorimetry stock sugar mixture into a glass tube, add 2.5 mL of 2.4 M sulfuric acid, and mix to give 3 mL of 3 mg/mL color sugars standard solution in 2 M sulfuric 2387_ch3.3_fm Page 79 Sunday, May 6, 2001 6:39 PM DIETARY FIBER ANALYSIS AS NONSTARCH POLYSACCHARIDES (NSPS) 79 acid. The colorimetric reaction is linear up to 3 mg/mL sugar. The absorbance of the test samples should not exceed that of the standard. Measurement of Total Reducing Sugars Place into separate glass tubes: 0.5 mL of the standard sugar solution and 0.5 mL of the hydrolysate from step 5; and place into each of two tubes (blanks 1 and 2): 0.5 mL of 2 M sulfuric acid. Add 0.5 mL of DMG solution and vortex-mix. Check the pH of 1 drop of blank 1; it should be between 3.5 and 4. If it is different from this, check the preparation of the 2 M and the 12 M sulfuric acid, and the DMG solution. If the pH is correct, add 0.1 mL of diluted pectinase solution, vortex-mix, and place all the tubes into a waterbath at 50°C for 20 min. Cool the tubes to room temperature, add 0.1 mL of 3 M sodium hydroxide, vortex-mix, and leave for 5 min. Add 1 mL of the dinitrosalicylate reagent to each tube and vortex-mix. Place all the tubes together into a briskly boiling waterbath for 5 min. Remove the rack of tubes and cool to room temperature in water. Add 10 mL of water (at room temperature) and mix well by inversion (do not use a vortexmixer at this stage). Measure the absorbance at 530 nm against blank 2. Note: Sample blanks may be prepared by diluting the hydrolysates as described above, replacing the color reagent with water, and reading the absorbance against water. Alternatively, and more conveniently, the absorbance of the undiluted hydrolysate can be measured against water and the value divided by 24.4 (the dilution of the hydrolysate after addition of the color reagent and water). The absorbance of the test sample is then calculated by subtracting this value. When the hydrolysate is colorless and the NSP content is more than 5%, the sample blank is not required. Calculation of NSP The amount of total NSP (portion A) and of insoluble NSP (portion B), in grams of polysaccharide per 100 g of sample, is calculated as NSP = A ( t ) × V ( t ) × D × F × C × 100 × 0.89 ------------------------------------------------------------------------------------------A(s) × W (t) (3.3.5) where A(t) is the absorbance of the sample solution (minus the absorbance of the hydrolysate blank if measured); V(t) is the total volume of the sample solution (in mL, here 30); D is the dilution of the sample solution (D = 1 if no dilution); F is the factor correcting the difference between the composition of monosaccharides in the standard sugar mixture and that in NSP of various types of plant foods (for the calculation of NSP in cereals (except oats), F = 0.95; for fruit and nonstarchy vegetables, F = 1.05; and for starchy vegetables, oat products, and unknown samples, F = 1; using the standard sugar mixture as specified and these factors makes corrections for the 2 to 4% hydrolytic losses); C is the concentration (in mg/mL sugars) of the standard; A(s) is the absorbance of the standard; W(t) is the weight (mg) of sample taken for analysis; and 0.89 is the factor for converting experimentally determined monosaccharides to polysaccharides. The amount of soluble NSP is calculated as the difference between total NSP and insoluble NSP. It is recommended that a sample be taken for the analysis of the dry matter as the material remaining after overnight incubation at 104°C. Results may then be expressed as grams of polysaccharides per 100 g of dry matter. Correction Based on Separate Measurement of Uronic Acids When a sample, e.g., pectin, has a high content of uronic acids, a more accurate value for NSP may be obtained if uronic acids are measured separately. The color standard sugar mixture contains 12.5% (w/v) uronic acids, and it has been determined (data not shown) that this leads to 17% underestimation of NSP when the sample contains only uronic acids (using F = 1). Correction for the underestimation is straightforward if a separate value for uronic acids is obtained. If the sample 2387_ch3.3_fm Page 80 Sunday, May 6, 2001 6:39 PM 80 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION contains 12.5% monomeric uronic acids, the color standard sugar mixture is entirely appropriate and no correction is required; otherwise, the correction required is an increment or decrement to the NSP value obtained by colorimetry equivalent to 17% of the uronic acid content that is in excess of or less than 12.5% of the sample, respectively. The percentage of the NSP value to be corrected for is the difference (∆) between the value for uronic acids (Z; expressed as a percentage of the NSP value, X) and 12.5, and it is calculated as ∆ = (Z/X × 100) – 12.5 (3.3.6) and the correction factor Y is calculated as Y = 0.17 (X/100 × ∆) (3.3.7) which may be reduced to 0.17Z – 0.021X, and the value for total NSP after correction is X + Y. Breaks in the Colorimetry Procedure The procedure may be halted at either of the following stages: (1) after precipitation, washing and drying the starch-free residue (see step 3.4); the residue may be stored for long periods; or (2) after the hydrolysis with 2 M sulfuric acid; the hydrolysate may be kept at 4°C for 48 hours. QUALITY CONTROL The certificated reference materials (CRMs) may be used as part of a complete quality control procedure. Trouble Shooting for the Common Hydrolysis Steps 1. Variation between replicate analyses may be due to inaccurate pipetting (test/calibrate dispensers by weighing replicates of water) or to incomplete removal of acetone in step 3.4. 2. If values for glucose are too high for samples of known composition and/or variable for replicates, this may be due to incomplete wetting of sample with DMSO in step 2.1. Mix vigorously immediately after addition of DMSO. 3. If values for glucose and uronic acids for samples of known composition are too low and/or are variable for replicates, this may be due to incomplete wetting of samples with 12 M sulfuric acid in step 5. Vortex-mix vigorously before and after addition of sulfuric acid and at intervals during the incubation. Trouble Shooting for the GLC Procedure 1. Extra peaks on the chromatogram may be due to incomplete reduction of monosaccharides. Ensure alkaline pH before adding NaBH4. Replace old NaBH4 ; do not compensate for loss of activity by adding more NaBH4. 2. Variation between replicate analyses may be caused by nonreproducible pipetting of the internal standard or hydrolysates. Test/calibrate dispensers by weighing 1-mL replicates of water. 3. If the response factors are not reproducible, this may be due to inaccurate pipetting of the sugar mixture and/or internal standard. Test/calibrate dispensers by weighing replicates of water. 2387_ch3.3_fm Page 81 Sunday, May 6, 2001 6:39 PM DIETARY FIBER ANALYSIS AS NONSTARCH POLYSACCHARIDES (NSPS) 81 Trouble Shooting for the HPLC Procedure 1. If retention times vary during chromatography, regenerate the analytical columns as described in the section on Determination of Constituent Sugars by HPLC. Trouble Shooting for the Colorimetry Procedure 1. Variation between replicate analyses may be due to inaccurate pipetting (test/calibrate dispensers by weighing replicates of water) or to incomplete removal of acetone in step 3.4. 2. If no color is produced for standards and/or samples, this may be due to an error in the preparation of the sulfuric acid or sodium hydroxide solutions. Make new reagents. Test that the pH of the solution is between 7 and 8 before adding the color reagent solution. REFERENCES 1. Southgate, D. A. T. and Greenfield, H., Principles for the Preparation of Nutritional Databases and Food Composition Tables, Siminopoulos, A. P., Butrum, R. R., Eds., International Food Data Bases and Information Exchange, World Rev. Nutr. Diet., 68, 27, 1992. 2. Koivistoinen, P. E., Asp, N.-G., Englyst, H. N., Hudson, G. J., Hyvönen, L., Kallio, H., and Salo-Väänänen, P. P., Memorandum on terms, definitions, and analytical procedures of protein, fat and carbohydrates in food for basic composition data: issues and recommendations, Food Chem., 57, 33, 1996. 3. FAO, Carbohydrates in Human Nutrition. Report of a Joint FAO/WHO Expert Consultation, Rome, 14-18 April 1997. FAO Food and Nutrition Paper 66. Food and Agriculture Organization of the United Nations, Rome. 4. Trowell, H., Ischemic heart disease and dietary fibre, Am. J. Clin. Nutr., 25, 926, 1972. 5. Trowell, H., Dietary fibre; a paradigm, in Dietary Fibre, Fibre-Depleted Foods and Disease, Trowell, H. C., Burkitt, D., and Heaton, K. W., Eds., Academic Press, London, 1985, 1–20. 6. Cleave, T. L., Campbell, G. D., and Painter, N. S., Diabetes, Coronary Thrombosis and the Saccharine Disease, John Wright, Bristol, 1969. 7. Burkitt, D. P. and Trowell, H., Eds., Refined Carbohydrate Foods and Disease. Some Implications of Dietary Fibre, Academic Press, New York, 1975. 8. Hipsley, E. H., Dietary fibre and pregnancy toxaemia, Br. Med. J., ii, 420, 1953. 9. Englyst, H. N., Wiggins, H. S., and Cummings, J. H., Determination of the nonstarch polysaccharides in plant foods by gas–liquid chromatography of constituent sugars as alditol acetates, Analyst, 107, 307, 1982. 10. Englyst, H. N., Trowell, H., Southgate, D. A. T., and Cummings, J. H., Dietary fiber and resistant starch, Am. J. Clin. Nutr., 46, 873, 1987. 11. COMA Committee News, Dietary Fibre. Food Safety Information Bulletin, No. 97, June 1998. 12. COMA Committee News, Definition of Dietary Fibre for Labelling Purposes. Food Safety Information Bulletin, No. 109, June 1999. 13. Holland B., Welch, A. A., Unwin, I. D., Buss, D. H., Paul, A. A., and Southgate, D. A. T., McCance & Widdowson’s The Composition of Foods, 5th ed., Ministry of Agriculture Fisheries & Food and Royal Society of Chemistry, Cambridge and London, 1991. 14. Englyst, H. N., Bingham, S. A., Runswick, S. A., Collinson, E., and Cummings, J. H., Dietary fibre (nonstarch polysaccharides) in fuit, vegetables and nuts, J. Hum. Nutr. Dietet., 1, 247, 286, 1988. 15. Englyst, H. N., Bingham, S. A., Runswick, S. A., Collinson, E., and Cummings, J. H., Dietary fibre (nonstarch polysaccharides) in cereal products, J. Hum. Nutr. Dietet., 2, 253, 1989. 16. McCance, R. A., Widdowson, E. M., and Shackleton, L. R. B., The nutritive value of fruits,vegetables and nuts. Special Report Series, MRC, London, No. 213, HMSO, London, 1936. 17. Southgate, D. A. T., Determination of carbohydrates in foods. II. Unavailable carbohydrates, J. Sci. Food Agric., 20, 331, 1969. 2387_ch3.3_fm Page 82 Sunday, May 6, 2001 6:39 PM 82 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION 18. Englyst, H. N. and Hudson, G. J., The classification and measurement of dietary carbohydrates, Food Chem., 57, 15, 1996. 19. Wood, R., Englyst, H. N., Southgate, D. A. T., and Cummings, J. H., Determination of dietary fibre in foods — collaborative trials. IV. Comparison of Englyst GLC and colorimetric measurement with the Prosky procedure, J. Assoc. Publ. Analysts, 29, 57, 1993. 20. V39, Dietary fibre: Englyst procedure for determination of dietary fibre as nonstarch polysaccharides: measurement of constituent sugars by gas–liquid chromatography, J. Assoc. Publ. Analysts, 33, 127, 1997. 21. V40, Dietary fibre: Englyst procedure for determination of dietary fibre as nonstarch polysaccharides: measurement of constituent sugars by colorimetry, J. Assoc. Publ. Analysts, 33, 145, 1997. 22. Pendlington, A. W., Meuree-Vanlaethem, N., and Brookes, A., The Method Specific Certification of the Mass Fraction of Dietary Fibre in Lyophilised Haricot Beans, Carrot, Apple, Full Fat Soya Flour and Bran Breakfast Cereal Reference Materials, CRMs 514, 515, 516, 517 & 518. Office for Official Publications of the European Communities, Luxembourg, 1996. 2387_ch3.4_fm Page 83 Sunday, May 6, 2001 6:41 PM CHAPTER 3.4 The Southgate Method of Dietary Fiber Analysis David A. T. Southgate INTRODUCTION This method, strictly speaking for the measurement of unavailable carbohydrates,1,2 was developed in the late 1950s in connection with a major study for the evaluation of the energy conversion factors used to calculate the energy value of the human diet.3 In this study we were concerned about the factor which should be used for available carbohydrates1 as compared with carbohydrate “by difference”4 as used in the classical Atwater system of factors.5 The protocol for the study also required us to measure all the carbohydrates, and therefore, a method was required for unavailable carbohydrates. The conceptual basis for the method was the method used by McCance et al.6 and aimed to measure the carbohydrates in the alcohol-insoluble residue. I also decided to introduce a number of constraints: first, to develop a procedure that measured all the carbohydrates in the same analytical sample, to avoid measuring a fraction twice; second, to attempt to use specific methods for the carbohydrates; and third, to start with a sample of a sufficient size because of the problems of taking a representative sample of many foods. STAGES OF THE METHOD The stages are summarized in Figure 3.4.1. Initial Extraction of the Sample Preparation of Alcohol-Insoluble Residue (Two analytical portions of the food are taken through the whole scheme.) The analytical samples should be of sufficient size (at least 5–10 g) to be representative and thoroughly mixed. The volume of alcohol added depends on the moisture content of the mixture and is about 85% v/v with respect to methanol; the mixture is brought to a boil with constant stirring (later versions of the method used ethanol at 80% v/v7). After filtering the residue is extracted with three further portions of aqueous alcohol. The alcoholic filtrates are combined and used for the measurement of free sugars. 0-8493-2387-8/01/$0.00+$1.50 © 2001 by CRC Press LLC 83 2387_ch3. gelatinize in boiling water Cool. Wash residue with 4 portions 80% ethanol Combine and measure glucose Centrifugates Starch Hydrolyze residue with 10 ml 1 M H2SO4 At 100°C for 45 min.4. 3RD EDITION Food Sample Extract with 4 portions of boiling 80% v/v ethanol Extract Alcohol-Insoluble Residue Measure Free Sugars Wash with acetone and dry Grind thoroughly Take 100 mg Add 4 ml water. Enzymatic Hydrolysis of Starch The alcohol-insoluble residue is extracted with diethyl ether and allowed to air-dry and weighed. The residue is finely ground and portions are taken for the measurement of polysaccharides. Weight Lignin Figure 3. After cooling the pH is adjusted with acetate buffer. adjust pH to 4. Cellulose pentoses. and uronic acids NCP Wash residue with diethyl ether and dry.1 Schematic flow diagram of modified Southgate method. amyloglucosidase enzyme is added. centrifuge. May 6. 2001 6:41 PM 84 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.4_fm Page 84 Sunday. add 10 ml ethanol Centrifuge decant and wash residue 3× with 50% ethanol. pentoses and uronic acids Non-Cellulosic Polysaccharides Stand residue overnight in 12 M H2SO4 Filter through sintered filter and wash with water Filtrates Combine and measure hexoses. Centrifugates Combine and measure hexoses. and the . These portions are gelatinized with water in tubes immersed in a boiling waterbath for at least 10 min. mix.6 (acetate buffer) Incubate with amyloglucosidase overnight Add 4 vols ethanol. The centrifugates are combined and glucose is measured to give a starch value. Determination of carbohydrates in foods. provide a quantitative direct measure of those components that approximated the original concept of dietary fiber9 and showed that the composition of dietary fiber varied between the major classes of foods. T. pentoses and uronic acids.4 The method did. pentoses. J. D. The mixture is filtered through a tared sintered glass filter and washed with water.. May 6.. This preparation is no longer available. Determination of carbohydrates in foods. 2001 6:41 PM THE SOUTHGATE METHOD OF DIETARY FIBER ANALYSIS 85 mixture is incubated at 37°C overnight (about 18 h). this preparation had some proteolytic activity which assisted in starch hydrolysis. and the ether is allowed to evaporate. the residue is washed and recentrifuged with three portions of 80% v/v ethanol. The residue is taken as lignin. T. 1969. Sci. 20. The combined filtrates are analyzed for hexoses. and uronic acids are not totally specific and need to be corrected for cross-interference. 1969 2. A. The combined filtrates are made to volume and analyzed for hexoses. NCP) The residue after enzymatic hydrolysis is suspended in 10 ml 1 M H2SO4 and heated for 2. a commercial Takadiastase for analysis on talc (Parke Davis) was used. Sci. Dilute Acid Hydrolysis (for Non-Cellulosic Polysaccharides.2387_ch3. Hydrolysis in Strong Acid The residues are washed in ethanol followed by diethyl ether. In the original method. The residue is washed with three portions of 50% ethanol. A.10 REFERENCES 1. II Unavailable carbohydrates.8 The values are expressed as monosaccharides and are approximately 10% higher than if expressed as polysaccharides. Southgate. D. After removal of the centrifugate. Food Agric. 12 M H2SO4 are added and the mixture is stirred and left overnight. 331. however. Food Agric. 10 ml of ethanol are added and the mixture is filtered..5 h in a boiling waterbath. Southgate. J. . RETROSPECTIVE This method is now really only of historical interest. Four volumes of ethanol are added and the mixture is centrifuged to recover unchanged polysaccharides. pentoses. I Available carbohydrates. and uronic acids4 to give a measure of the composition of the NCP. Expression of Results The colorimetric values for hexoses. 326.4_fm Page 85 Sunday.. Measurement of Residual Lignin The residue on the filter is washed with ethanol followed by diethyl ether and dried and weighed. 20. having been superseded by more specific gas–liquid chromatographic (GLC) and high-performance liquid chromatographic (HPLC) methods for measuring the component sugars and uronic acids in the hydrolysates. . in Dietary Fibre. V. and Englyst. Rep. P. D. A. 1981.. Res.. and nuts. D. Human Nutr. J.. 1980. G. and Theander. M. O.. T. Fibre-Depleted Foods and Disease. A. 5.2387_ch3. 303. R. Widdowson... Med. 9. Southgate. J. Basis and Derivation. physical properties and analysis. G. New York.. James. and Bailey. 3RD EDITION 3. McCance.. no. 31. Southgate... and Durnin. 517. H.. . 1936.. Food Chem. and Heaton. Marcel Dekker. Merrill. 6. Department of Agriculture. Southgate. 8. and Shackleton. A guide to calculating intakes of dietary fibre. Chapman and Hall. F. E. HMSO. B. 74. 213. 2nd ed.. A. Academic Press. Calories conversion factors. 1985. Eds... May 6. Mutual interference effects in the colorimetric methods used to determine the sugar composition of dietary fibre. T. 2001 6:41 PM 86 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. A.S. T.. The Analysis of Dietary Fibre in Foods. 10. Br J. 30.. Burkitt. Eds.. Collinson. D. 1970. vegetables. Bailey. J. The nutritive value of fruits. and Watt. Ser. A. K. U. E. 1992. 4. A. A. London. R. and Walker. 201. 7. An experimental assessment of the factors used in calculation of the energy value of human diets.. 5. London. 1955. L.. 1976. D. Spec. Determination of Food Carbohydrates. B. Dietary fibre: chemistry. A.. Nutr. T. Southgate.. D. W. B. S.. B. Agriculture Handbook No. Coun. K. 24. L. H. Hudson. T.4_fm Page 86 Sunday. Trowell.. Energy Value of Foods. 13). polysaccharides. the value of Klason lignin includes native lignin but also other components such as tannins. lignin. in conjunction with this chemical definition. The Uppsala method and the AOAC enymatic gravimetric methods often give similar results because they use a similar definition for DF. Today it is often proposed to expand the definition of dietary fiber to include other components — for example. The latest version of the method. The Uppsala method for total DF analysis. and Per Åman INTRODUCTION In recent years. cutins. and 0-8493-2387-8/01/$0.K.5 Determination of Total Dietary Fiber and Its Individual Components by the Uppsala Method Olof Theander. and some proteinaceous products as well as Maillard reaction products in heat-treated foods. In the original version of our methodology. 2001 6:43 PM CHAPTER 3. Commission of the European Communities. the degree of polymerization of the nonstarch polysaccharides was intended to be the same as that defined by the International Union of Pure and Applied Chemistry (IUPAC).00+$1. greater than 10. the American Association of Cereal Chemists (AACC-method 32-25). and a U. When human foods are analyzed.4 The results showed that the five samples in the study could be accepted as standards for all four methods. oligosaccharides and fructans — in the analysis. method were evaluated in an intercomparision and certification study organized by the Community Bureau of Reference. it has become apparent that our understanding of the nutritional effects of dietary fiber (DF) has been considerably hampered by the lack of an appropriate definition of DF and. a lack of adequate analytical methods.50 © 2001 by CRC Press LLC 87 . published a gas-chromatographic method for analysis and characterization of DF. May 6. but not oligosaccharides or fructans.1 In this method. two AOAC enzymatic gravimetric methods. and the Nordic Committee on Food Analysis (NMKL-method 162).3 has been studied in a collaborative study and is now approved as an official method by the Association of Official Analytical Chemists (AOAC-method 994. AACC recently suggested DF be defined as follows: “DF consists of the remnants of edible plant cell.2387_ch3. that is.K. Roger Andersson. the Uppsala method. This original method has now gained merit as a Current Contents Citation Classic.2 Enzyme-resistant starch. while the U.5_fm Page 87 Sunday. defined as the starch that resists hydrolysis with the thermostable α-amylase and the amyloglucosidase used in the method. Eric Westerlund. consequently. Theander and Åman proposed that DF could be defined as the sum of nonstarch (amylase-resistant) polysaccharides and Klason lignin and. is included in the dietary fiber polysaccharides. procedure that only includes plant cell wall nonstarch polysaccharides in the analysis generally gives lower values. there is generally no sharp distinction between water-extractable (or removable) and water-unextractable fractions. however. method of Englyst17 are shown in Table 3. obtained by the Uppsala method. 3RD EDITION associated substances resistant to (hydrolysis) digestion by the alimentary enzymes of humans.25 A group that has been actively engaged in studies of individual fiber components — in particular.21 In a comparison between the U.8 This method. GLC determination of sugar constituents. including a critical discussion of the various steps and of alternative methods. we recently published a method for analysis of human digesta. gelatinization at pH 7 and 85 to 90°C. U.5. 2001 6:43 PM 88 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.14–17 including the development of a colorimetric procedure. BACKGROUND Our understanding of the chemical composition of DF in foods was to a large extent pioneered by Southgate. consider such enzymatically nonavailable starch as part of the DF complex.13 In both methods starch is removed enzymatically. This was demonstrated at the meeting on DF analysis in Cambridge. but with different systems. and that of the U.K. whose studies are based on the fractionation and colorimetric assay of the hexose.7 In Europe. which. May 6. so far. and the starch is incompletely removed. Another method. sugar alcohols. were also explored. as well as the scientists who have developed the enzymatic gravimetric AOAC method for analysis of DF. For several polysaccharides. and Uppsala methods. group used extraction with dimethylsulfoxide (DMSO) in combination with enzymatic treatments to remove not only the enzymatically available starch but also the so-called enzymeresistant starch.5_fm Page 88 Sunday. are added to the values for nonstarch polysaccharides found by the U.K. The Englyst method has undergone a number of modifications.1.K.2). however.9 A more detailed description of the Uppsala methodology. is that of Faulks and Timms. despite its complexity. also will include oligosaccharides. and a direct gas–liquid chromatographic (GLC) assay is used for the neutral sugar constituents of the fiber fractions. which is related to the Uppsala methodology but which uses DMSO for solubilization of starch.22 there was a rather good agreement in total DF when the values for enzyme-resistant starch and Klason lignin.5. These included two fractionation methods for soluble and insoluble fibers devised by Englyst12 and Theander and Åman. A large variability in results of the method between different laboratories was shown. The U.”5.K.28 as well as the previously cited book by James and Theander.23 Marlett’s group has made extensive studies on measurement of DF using a modification of the Uppsala methodology (see Marlett24 and references therein).26.11 In the Cambridge study. is still not specific in regard to individual sugars. although the agreement within laboratories seemed to be good.6 In accordance with a broader definition. method (Table 3. We. efforts to agree upon a common definition on DF have not been successful. enzymatic treatments with α-amylase and pullulanase. pentose.27 Their method involves extensive ball milling. in 1978.1. on the other hand. The main differences between the present method3 for total DF.18 The original method of Theander and Åman1 was later developed into two modifications19. One possible explanation for this may be that the presence of DMSO partly hampers the precipitation of soluble DF polysaccharides. a trend that these sums were slightly lower in the latter method.K. Selvendran has also published a review of methods for DF analysis which is recommended for further reading.2387_ch3. the ratio between them is dependent on the choice of conditions during the pretreatment .20 for analysis of total DF.9 where results from analyses of different types of food samples by Southgate’s and other methods were presented. informative ones. and colorimetric uronic acid determination. and uronic acid constituents.10. There was. is given below. as well as more comprehensive. which initiated several new collaborative studies and further developments in the field of DF analysis. the preparation of cell wall materials — is that of Selvendran. and fructans. besides dietary fiber polysaccharides and Klason lignin. rapid gravimetric methods. developed by the Uppsala group. Termamyl (0.7 3.4 M H2SO4 (1 h. Dimethylsulfoxide (DMSO) (0.7 0.3 0. Uppsala Methoda DF KL RS 2. KL. ice water) at pH 2 1.1 Comparison of Main Steps in Total Dietary Fiber Determination by GLC with the Latest Modifications of the Uppsla Method3 and the U. May 6.8 23.K. Termamyl (10 min. Methodb NSP NSP + KL + RS (% dry matter) 0.8 22.3 0.8 1.3 ndc nd nd U.8 20. 50°C + 10 min. Addition of internal standard myoinositol 3. Klason Lignin (KL). boiling waterbath) in acetate buffer (0.9 20. boiling waterbath) 80% (v/v) ethanol (0. Alditol acetate preparation by sodium borhydride reduction and 1-methylimidazole/acetic anhydride treatment 5.1 64.3 2. method 250–500 mg 50–300 mg Extraction with petroleum ether when content exceeds 5% Extraction with 80% ethanol when content is very high Extraction with acetone when content exceeds 10% 1.5 0.17 Table 3.08 M. 125°C) 4.5.2 67.7 2. pH 5.5.2) 3.3 Englyst and Cummings.15 Not detected. Pancreatin and pullulanase (0.6 3. boiling waterbath) in acetate buffer (0.2 14. Method. 0. 100°C) 3. boiling waterbath) 2. 12 M H2SO4 (1 h.4 15.5_fm Page 89 Sunday. pH 5.1 19.5 3.3 13. Individual correction factor for each sugar Colorimetry with 3. Amyloglucosidase (16 h.5-dimethylphenol (Scott. Alditol acetate preparation by sodium borhydride reduction and 1-methylimidazole/acetic anhydride treatment 5.2387_ch3. Addition of internal standard allose 4.5 h.5 h. and the Sum of NSP. 35°C) 80% (v/v) ethanol (1 h.2 Content of Dietary Fiber (DF). and Enzyme-Resistant Starch (RS) as Determined by the Uppsala Method. 4°C) 1.2 0. and RS for Comparison Product Corn flakes Bread crust Bread crumb Rye crisp Green peas Soybean Deskinned onion Sugar-beet fiber a b c Theander et al.1 0.8 15. 1979) Gravimetrically as Klason (sulfuric acid) lignin including ashing 2.5 h.8 2. 2 M H2SO4 (1 h.K. 2001 6:43 PM THE UPPSALA DIETARY FIBER METHOD Table 3.3 1.0 12.1 M. 12 M H2SO4 (0.0 1.K.5 h.5 h.K.0 2.0) 2.7 3. 1979) Not determined Theander et al.7 20.5 0. 60°C) 1.5-dimethylphenol (Scott. 30°C) 2.5 16. Individual correction factor for each sugar Colorimetry with 3. Method17 Procedure Uppsala method Sample size (dry matter) Fat removal Removal of low molecular weight carbohydrates Starch removal Precipitation of soluble fiber Analysis of neutral sugars Analysis of uronic acids Lignin a b 89 U.3 .2 68.8 0.3 0. Nonstarch Polysaccharides (NSP) as Determined by the U.3 Englyst et al. and samples which also have high sugar contents. and citrus pectin are recommended for such use in the Uppsala method.31. extraction with petroleum ether at room temperature is efficient for most lipids. This enzyme system has proved to be very effective for various types of starch-containing fiber sources and products. β-glucanase activity and will thus give low values for the fiber contents in. The use of DMSO in the U. and so on).23 Recently. method can be expected to increase the solubility of DF. Removal of Starch Enzymatic removal of starch is a crucial operation in the Uppsala method in order not to overestimate the DF glucans. May 6.33 We usually quantify the glucose by a glucose oxidase reagent (Megazyme Int. for example. CRITICAL DISCUSSION OF VARIOUS STEPS IN DETERMINATION OF DIETARY FIBER Removal of Free Sugars and Lipids It is important that the samples are representative and homogeneous. Under the conditions used. temperature. mixed-linkage β-glucans. neither of the enzymes liberated detectable amounts of sugars from purified barley grain β-glucan. and feces. barley straw arabinoxylan. at the high temperature used. and accurate. studies29. and pectins (including associated neutral polysaccharides) are found both as water-soluble and water-insoluble components in plant materials. are preferably extracted with 80% aqueous ethanol in an ultrasonic waterbath.3 This method has been applied to a number of divergent samples. since the reagent is an excellent solvent for hemicellulose polysaccharides. If the lipid content of the sample exceeds 5%. including foods. the incubation with amyloglucosidase is then left to stand overnight. of any nonstarch polysaccharide-degrading enzymes present. cereal samples. Ireland Ltd. oat β-glucan. Höganäs. the sample is treated with Termamyl for 30 min in a boiling waterbath. All enzyme preparations and batches should. The powerful Termamyl enzyme causes a rapid hydrolysis of starch during gelatinization. or cotton cellulose. 2001 6:43 PM 90 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.32 in which protein in the residue must be analyzed. and has proven to be rugged. Foods with low water content (<15%) are ground in a Cyclotec Sample Mill (Foss Tecator AB. Commercially available fractions of arabinoxylan. be checked for such activities. For practical reasons.3 Materials with high water contents are generally freeze-dried.K. although complete hydrolysis to glucose is reached after about 6 h.1 The specificity of the Termamyl enzyme is probably explained by a rapid inactivation.2387_ch3. however. A skilled analyst can run over 40 samples per week by this improved procedure.5.1) for routine analysis of total DF was tested in a collaborative study. like fruits and vegetables. Ireland).30 have shown that the yield and composition of soluble fibers are very dependent on extraction conditions used.5_fm Page 90 Sunday. for instance. digesta. The yield of these components can vary considerably with the fractionation conditions used (physical pretreatment. Starch is removed by incubations with a thermostable α-amylase (Termamyl) and amyloglucosidase.5-mm screen. adaptable. enzymatic treatment. the Uppsala method (Figure 3. These conditions are. It is imperative that the enzymes used are free from fiber-degrading activity. The Uppsala method is not much more time-consuming than the gravimetric enzymatic methods. never identical with in vivo conditions. reproducible. . Polysaccharides such as arabinoxylans. In the present procedure. feeds. Bray. therefore. Sweden) fitted with a 0. Thus. 3RD EDITION and solubilization procedure. Unfortunately it would appear that some commercial amyloglucosidase preparations at present have. time.. (Source: Theander. 1995. Assoc. With permission. 78.5_fm Page 91 Sunday. Chem. 1030.5. Off. O. Anal. et al. May 6.2387_ch3. 2001 6:43 PM THE UPPSALA DIETARY FIBER METHOD 91 Figure 3.1 Analysis of dietary fiber by the Uppsala method... J.) . 49 0.. Another factor contributing to the DF increase is from the Maillard reaction. 10. on the other hand. mainly as a result of increasing contents of water-soluble glucans.64 1..21 0.3).86 0. Holm et al.5.38 These modifications are. Source: Westerlund. Cereal Sci. as well as several others working in this field.09 44 0. which eliminates the phenomenon of retrogradation. branched polysaccharide structures. J. The total content of dietary fiber glucans (enzyme-resistant starch is not included).35 have studied in vitro and in vivo enzymatic digestibility of amylose–lipid complexes and found. A study40 in our laboratory has shown that the formation of enzyme-resistant starch decreased from crumb to outer crust during baking (Table 3. 2001 6:43 PM 92 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. part of the starch in foods may be resistant to α-amylase hydrolysis and that a solution of gelatinized starch may retrograde on cooling and drying. We have found that starch under food-technical conditions can also to some extent be modified nonreversibly via fragmentation to saccharides with 1. In the interlaboratory Helsinki study.65 0.5.60 0.2387_ch3. 3RD EDITION Batey34 reported that the gelatinization and solubilization of starch with a thermostable α-amylase prior to amyloglucosidase treatment resulted in higher starch values than those obtained after 8 M hydrochloric acid. May 6.70 0. This progressive decrease was probably due to water and temperature gradients formed in the bread during heating.27 In connection with DF analysis.11 0. which will be discussed below under the lignin determination.62 12 0.14 solubilized this retrograded enzymeresistant starch with 2 M potassium hydroxide solution or DMSO16 before hydrolysis with amyloglucosidase to glucose. the moisture content.14.08 0. increased from crumb to outer crust. and other factors. This observation strongly suggested that formation of nonstarch glucans had. which can via further transglucosidation form enzyme-resistant. .37.20 0. among other things.58 1.39 there was a slight but significant increase for the DF glucan values of extruded wheat flour and whole meal samples as well as for various heat-treated potato samples.1%). and so on.5_fm Page 92 Sunday. E. 1989. et al. after heat treatment. dependent on the extent of heat treatment.02 44 0. frying. 149. The two main routes whereby the amount of enzymatically available starch may decrease (and the DF glucan may consequently increase) are (1) retrogradation and (2) chemical modification of the starch structure.36 We. They have applied this technique on a large series of cereal foods and reported values of enzyme-resistant starch in the range of 0.3 Content (g/100 g dry sample) of DF Polysaccharides and Enzyme-Resistant Starch in Dough and White Bread Fractions Fraction yield Soluble DF glucan Insoluble DF glucana Soluble arabinoxylan Insoluble arabinoxylan Resistant starch a Dough Crumb Inner crust Outer crust 100 0.09 1.03 0.72 1.6-anhydroglucose end units.65 0. autoclaving. regard this enzymatically nondegradable starch as a DF constituent and call it enzyme-resistant starch. He pointed out that the good reproducibility of the analytical values using thermostable α-amylases — which is also our experience from various types of food products — is due to the rapid degradation of starch to lower molecular oligomers. DMSO or potassium hydroxide treatments.1 to 1. baking.2% (except for one breakfast cereal having 3. that the use of Termamyl in combination with pancreatin almost completely hydrolyzed the complex. occurred Table 3. among other factors.10 0. Englyst et al. The main advantage of using such enzymes — both for removal of starch in the DF determination and for direct starch analysis — is that autoclaving is not needed.30 Enzyme-resistant starch is not included. A number of other reactions and modifications of starch are conceivable during various normal heat treatments of food such as extrusion cooking. at least in part. It is well established that. 1 0. raw potato.47 They illustrate the well-known higher rates of degradation with acids for pentoses (in particular xylose) and also why we choose 6 h for the posthydrolysis step to ensure hydrolysis of sulfate ester groups (also 1 h at 125°C is enough).5 1.4 1. This decrease was partly accompanied by an increase in the content of water-soluble arabinoxylans. for example.5 0.1 0.42 we found that some polysaccharides may remain in solution after precipitation with 80% ethanol.8 1. carrot. Precipitation of Soluble Polysaccharides In previous studies on rapeseed flour.1 traces 0..2 0. et al.9 1.1 0.5.1 0.49 when .5. followed by a secondary hydrolysis in dilute acid for hydrolysis of lignocellulosic materials in combination with Klason lignin determination as originally developed by Hägglund et al.8 1.2 0. Determination of Neutral Polysaccharide Constituents Treatments with 12 M sulfuric acid aiming to effect dissolution of cellulose. show an example of the sugar yield and rate of sulfate ester removal after different times of hydrolysis for xylose and glucose. The amounts of soluble fiber polysaccharides not recovered were low: 1–6% (Table 3.2 0.43 the polysaccharide fraction (DP > 10) escaping precipitation was analyzed in various foods.7 1.4). The content of insoluble arabinoxylans decreased during baking.2 0.1 0.1 0.4 Amounts of Neutral Polysaccharide Residues Not Recovered on Precipitation of Soluble Fiber with 80% Ethanol in Samples Analyzed by the Uppsala Procedure (% of total fiber content) Sample Rhamnose Arabinose Xylose Mannose Galactose Glucose Total Corn flakes Bread crust Bread crumb Rye crisp Green peas Soybean Deskinned onion Sugar-beet fiber 0. dilution to 0. Figures 3. Recently in a detailed study using gel filtration of the ethanolic supernatant. essentially according to Sloneker46).1 1.5 0. from an investigation made at the Swedish Forest Research Laboratories on the polysaccharide analysis. May 6. e.5.2 0. This indicates that the solubility of DF polysaccharides may be changed by thermal treatment. It is important that the hydrolysis step with dilute sulfuric acid is sufficient to hydrolyze sulfate ester groups introduced in the first step. wheat flour.1 0. J.g.. particularly in the outer crust.2 0. In a preparative experiment in which the cellulose was hydrolyzed for only 4 h.3. we isolated part of the original cellulose as glucose-6-sulfate..9 1. mainly due to the formation of RS.1 0.2 0.1 0.6 3.3 0.5 0.0 3.8 0.3 0.1 0. Such a method is used for the hydrolysis of the polysaccharides in the Uppsala method.48 It is notable that Neilson and Marlett.3 0. We have previously shown20 that such precipitation losses did not exceed 3% of the total DF polysaccharides of.1 0.4 M.2 0.5 0.1 0. These losses of the total fiber content of the original sample were observed.41. 703. it is a key point to reach as complete a recovery as possible of soluble DF on their precipitation in 80% aqueous ethanol.5_fm Page 93 Sunday.2 0. probably by depolymerization. and wheat bran. The total dietary fiber content was higher in the bread fractions than in the dough. AOAC Int.2 0.1 0.1 0.5 0. 1994.1 0.1 2.45 are widely used (see below).3 0. namely pretreatment with 12 M sulfuric acid at room temperature for 2 h.2 0.3 0.6 5. with the highest values found for the most severe heat-treated sample (bread crust).0 Source: Theander.7 1.5.1 0. 77.44 and later modified by.40 Table 3.2 0. In the Uppsala method. O. and hydrolysis for 6 h under reflux1 or to gain time for 1 h at 125°C20 (in an autoclave. 2001 6:43 PM THE UPPSALA DIETARY FIBER METHOD 93 via fragmentation of starch and subsequent reactions of the fragments. Saeman et al.2 0.2387_ch3.2 and 3.. 25% were found.2387_ch3. part of the xylose. (Source: Bethge. 1 h) with 12 M sulfuric acid. et al.5_fm Page 94 Sunday.2 Yield and sulfate ester decomposition of xylose by reflux in 0. as have certain other workers in the field.4-linked glucose units interdispersed with β-1. 63B. a cellulose content of 99.4 M sulfuric acid hydrolysis after pretreatment (30°C. Communication No.3-linkages) are important components of some oat and barley products and can be determined by separate methods. since varying amounts of noncrystalline cellulose (depending on the source of the sample) may also be hydrolyzed under these conditions. found appreciable amounts of cellobiose (around 10% of total sugars) in the hydrolysates of the latter method.. Further.3) and a xylan content of 0. Kvantitativ Kolhydratbestämning — en detaljstudie.and mannose-containing polymers need pretreatment with strong sulfuric acid to be completely hydrolyzed with the dilute acid. With permission.0% (SD 1. It has been reported that for materials such as carrot. On average.14 The so-called β-glucans (containing β-1. 1971. and soybean. P. Using the autoclaving conditions. attempted to estimate the noncellulose component of the insoluble DF separately from the cellulose by using an extra heterogeneous hydrolysis step with dilute sulfuric acid. Stockholm. Swedish Forest Products Laboratory.) comparing a modified neutral detergent fiber method with a modified Uppsala procedure with only 3 h in the second hydrolysis step.22 The effect of particle size and different prehydrolysis conditions to achieve optimum hydrolysis yield has been investigated.5. May 6.51 . O.50 In the Uppsala method we have not. cabbage. 3RD EDITION 50 100 80 Yield 30 60 20 40 10 20 Yield (%) Sulfur content (µg/mg) 40 Sulfate ester content 0 0 0 4 8 12 16 20 24 Time (h) Figure 3. 2001 6:43 PM 94 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. part of the hemicellulose fraction may be difficult to get into solution. the amounts of cellobiose or other oligosaccharides in our hydrolyzed fractions are negligible. The efficiency of our conditions for hydrolysis of crystalline cellulose has been demonstrated by eight independent analyses of purified cotton linter. is the method of choice for an efficient separation and accurate determination of the sugar constituents in hydrolysates. Connors and Pandit53 introduced the use of 1-methylimidazole as an effective catalyst for rapid acetylation of polyhydroxy compounds. after acidification.54 reported the use of this catalyst for the quantitative and rapid acetylation of alditols in the presence of borate (at room temperature for 10 min). O. facilitates and shortens the derivatization step considerably. This is essential. 1 h) with 12 M sulfuric acid. May 6. The sugars are first converted to the volatile alditol acetate derivatives.52 Hydrolysates containing appreciable amounts of uronic acids must be carefully neutralized and the pH adjusted to the slightly alkaline side with. 63B. since borate forms complexes with the alditols and interferes with the subsequent acetylation with acetic anhydride and the commonly used catalyst pyridine. Communication No. Kvantitativ Kolhydratbestämning — en detaljstudie. for instance. In the original procedure (see. and later Blakeney et al. which we now generally use. not found any epimerization of hexoses or pentoses under the conditions used. which would otherwise be reduced with borohydride to alditols in the subsequent reduction step. We have. 47. but most sugars are probably transformed into alditols much quicker. References 1. With permission. 2001 6:43 PM THE UPPSALA DIETARY FIBER METHOD 95 100 Yield 40 80 30 60 20 40 10 20 Yield (%) Sulfur content (µg/mg) 50 Sulfate ester content 0 0 0 4 8 12 16 20 24 Time (h) Figure 3. A procedure for direct acetylation of aldoses in the acidic hydrolysate (without previous reduction) . however. (Source: Bethge.4 M sulfuric acid hydrolysis after pretreatment (30°C.. has usually been removed as the volatile trimethylborate by repeated evaporation with methanol. Swedish Forest Products Laboratory.) GLC.5_fm Page 95 Sunday.5..1 essentially as described by Sloneker. e. the borate from the borohydride.g. et al. 1971. This technique.2387_ch3. The reduction is left to stand at 40°C for 1 h. P.3 Yield and sulfate ester decomposition of glucose by reflux in 0. dilute ammonium hydroxide (without causing any epimerization of sugars) in order to open any uronolactones. Stockholm. and 52). particularly on capillary columns. Automated procedures using ion-exchange resins have also been successfully employed for the separation of monosaccharides. fucitol. 3. 2001 6:43 PM 96 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 4. in fact. was developed in our laboratory.5.60. As the ratio between the different sugars may influence these factors.5_fm Page 96 Sunday.90 used to convert the pentose/deoxyhexose and hexose contents. are also analyzed concomitantly.5 shows the corresponding separation from a wheat bran concentrate. rhamnitol. Capillary column chromatography has been a key factor in the separation of these complex mixtures. Comprehensive reviews56–58 on HPLC with various applications in the carbohydrate field have been published. myo-inositol is added as internal standard before the hydrolysis step with dilute sulfuric acid. imparting good thermal stability and prolonged life. reference sugar mixtures. including the use of amperometric determination59 of monosaccharides. glucitol.5. arabinitol. and Figure 3.25 mm). Capillary fused silica column (25 m × 0. xylitol. and 8. 3RD EDITION 8 7 4 Detector response 3 1 12 5 6 2 13 14 15 16 17 18 19 20 21 22 23 Retention time (min) Figure 3. myo-inositol (internal standard). galactitol.4 shows a typical GLC separation of alditol acetates derived from a reference mixture. although derivatization has.55 So far this procedure has been tested on only a limited number of samples. 2. and GLC responses for the individual sugars have been applied. mannitol. Peak 1. In these correction factors are also included the factors 0. to polysaccharides. High-performance liquid chromatography (HPLC) offers an alternative procedure for analyzing aldoses in polysaccharide hydrolysates but has not so far proved to be as powerful a technique as GLC. In the GLC separation. May 6.2387_ch3. typical for the DF composition of the food sample in question. 6. 5. and the so-called bonded phase (BP) columns. Correction factors for hydrolysis losses.5. in which the liquid phase is covalently bonded to the column.4 GLC-analysis of a monosaccharide reference mixture as alditol acetates. An advantage of HPLC is that the preparation of derivatives is obviated.61 . governed by the requirements of the investigation. but the agreement is good compared with the Uppsala method. however. often been used in HPLC in order to enhance resolution and detection of the components and to shorten the analysis time. 7. derivatization yields. Figure 3. have introduced a further degree of efficiency into the chromatographic operations. respectively. The choice of method is. which seems promising.88 and 0. They also calculated the methyl ester content from the binding of copper to the sample before and after saponification.2387_ch3.5 GLC-analysis of the monosaccharides in a hydrolysate of wheat bran concentrate as alditol acetates. The introduction of 3-phenylphenol by Blumenkrantz and AsboeHansen67 and later 3. low-molecular uronic acids may be readily analyzed by GLC in the presence of neutral sugars. interference from absorbing compounds resulting from neutral sugars. A useful technique has been developed for the separation and analysis of individual monomeric uronic acids by partition chromatography of ion-exchange resins unmodified64 (including oligomeric acids) or after transformation to aldonic acids.63 the release of these acids in acceptable yields from the polymers is complicated by the high stability of glycosyl uronic acid linkages toward acid hydrolysis — resulting often in the formation of aldobiouronic acids.65 Most workers in the DF field have used colorimetric assays and.68 The Uppsala group uses galacturonic acid for calibration because of this and the fact that galacturonic acid is more predominant in most DF samples (pectins) compared to glucuronic acid.5_fm Page 97 Sunday.4. Further. The rate of formation of 5-formyl-2-furancarboxylic acid.68 who also compared the use of different phenol derivatives for colorimetric determination of uronic acids in plant materials. Conditions and peak numbering as in Figure 3. and the inability to distinguish between individual uronic acids. the modified carbazole method according to Bitter and Muir.66 Common disadvantages with colorimetric methods are the sensitivity to reaction conditions. in particular. offers certain advantages. . In the Cambridge study. phenols.62 Although free. Methanol released by such a treatment can also be determined by GLC. the monomeric uronic acids when released are degraded to noncarbohydrate products more rapidly than the neutral sugars.5-dimethylphenol by Scott.5. Katan and Van de Bovenkamp69 used a copper-binding method for determining the uronide content. Determination of Uronic Acid Residues in DF Polysaccharides The determination of uronic acid constituents in DF by GLC presents a greater problem than in the case of the neutral sugar constituents. was shown to be faster for galacturonic than for glucuronic acids. 2001 6:43 PM THE UPPSALA DIETARY FIBER METHOD 97 Detector response 4 3 7 8 5 12 13 14 15 16 17 18 19 20 6 21 22 23 Retention time (min) Figure 3. The latter phenol reagents seem to suffer less interference from hexoses and have a greater sensitivity than the carbazole reaction. proteins. on which the colorimetric determination is based.5. May 6. and other components present. is refluxed for 30 min with hydroiodic . containing 1 to 20 mg uronic acid.6 Differences in absorbance obtained by colorimetric determination of uronic acids in autoclaved hydrolysates of citrus pectin (upper line) and galacturonic acid (lower line).62 The straight lines through the origin (Figure 3. Thus. May 6. 3RD EDITION Figure 3. 2001 6:43 PM 98 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.5. polyuronide values have to be multiplied by 0. if the calculated content of uronic acid residues is based on the degradation observed for the monomer.13. even those with low uronic acid contents.1.5_fm Page 98 Sunday. One should be aware of the fact that variations in the pectic polysaccharide structure and molecular mass as well as experimental conditions may influence the value of this factor.19 The sample. 1 h) to account for observed differences in the degradation rate70 of polygalacturonic acid residues and that of the free monomer.5.2387_ch3. Different amounts of rhamnogalacturonan from citrus with a welldetermined content of polygalacturonic acid residues (analyzed by titration and corrected for the presence of acetyl groups) and the D-galacturonic acid were subjected to acid treatment in separate experiments.88 to compensate for the lower degree of acid degradation of citrus pectin compared to that of the free monomer. We have established that the decarboxylation method developed by Bylund and Donetzhuber71 for woods and pulps also affords a rapid. The decarboxylation method for uronic acid determination does not suffer from the interference problems which are incurred with the colorimetric methods.4 M sulfuric acid.6) showed that the rate of degradation of the uronic acid–containing samples was reproducible. accurate. the degradation of D-galacturonic acid is compared with that of a well-characterized citrus pectin treated under identical acidic conditions (0. and reproducible method for DF samples. 125°C. In the Uppsala procedure. and pectic acid give the same change in conductivity per mole of hexuronic acid.20 Different types of uronic acids. the complex polymer of phenylpropane units — at least. originally introduced by Johnson et al.81 We have found a similar increase of the Klason lignin residues and their crude protein contents with the extent of heat treatment for potato and cereals37. which. chosen the sulfuric acid method. These constituents of the Klason lignin residue most likely represent food components. The carbon dioxide released is trapped in a cell containing dilute sodium hydroxide.1. It is conveniently combined with the determination of the neutral polysaccharide constituents of the DF. might also originate from protein-tannin . 2001 6:43 PM THE UPPSALA DIETARY FIBER METHOD 99 acid under nitrogen. and the lignin content is measured by dissolving the residue in 25% acetyl bromide in acetic acid and measuring the absorption at 280 nm. phenols. and the conductivity changes are registered by means of a potentiometric recorder. but which has since been further modified. and mannuronic acids. paper electrophoresis of the polysaccharide hydrolysate from the sugar analysis (even though the uronides are not completely hydrolyzed) affords a rapid. We therefore propose that the Klason lignin value will be designated the “noncarbohydrate” part of the DF. or alginate. cutin.22 When applied to human foods.1 This method. however.28 pointed out that the acetyl bromide procedure can be used only to compare the lignin contents of different organs from the same plant or samples from similar species. pyrroles. Eastwood and colleagues. D-glucuronolactone. carboxylic acids. like lignin. has been further developed and applied to forage crops by Morrison. rough estimate of the ratio between the corresponding galacturonic. Selvendran et al.77 More recently.82 It analyzed as 90% Klason lignin with a recovery of 84% of the original nitrogen. does not estimate the amount of individual uronic acid constituents but only the sum of uronic acids. which was first applied by Klason78 at the beginning of this century.73 The acetyl bromide method. The sample is extracted with water and organic solvents. are unavailable to human enzymes. Van Soest has demonstrated for some forages and feeds that high-temperature treatment increases the amount of crude protein and Klason lignin of the ADF residue. like the colorimetric methods. acidic xylan constituents. an ashing step is also included. We have found that the uronic acid determination gives essentially the same result whether it is performed before or after the extraction with organic solvent and starch removal. The lignin content determined according to Goering and Van Soest80 by KMnO4 oxidation of the ADF (acid detergent fiber) residue is probably generally closer to the amount of native lignin. A brown water-insoluble polymer was obtained (together with low molecular weight furans.72 Determination of Lignin There is no specific method for the determination of lignin. and some proteinaceous products.5. the aldobiouronic acid 2-O-(4-Omethyl-α-D-glucopyranosyluronic acid)-D-xylose. considered the method inappropriate and not directly applicable to foods. like many others working with lignified plant materials.83 The crude protein contents which one finds in the Klason lignin residue from heat-treated foods. In the present Uppsala method.79 Essentially. if one wishes to avoid laborious and time-consuming procedures. and lactones) by refluxing a slightly acidic aqueous solution of glucose and glycine. pyridines. We have. From heat-treated foods.5 for some examples). besides originating from such Maillard reaction products. since different species require different conversion factors. the residue may also contain products of the Maillard reaction and caramelization reactions. the values in most cases represent not only native lignin but also tannins. May 6. If the uronide content mainly comprises pectin. however.2387_ch3. it involves removal of extractives and polysaccharides and the gravimetric determination of the residue as lignin after washing and drying. such as D-galacturonic acid.74 for wood and wood products.39 (see Table 3. We have applied an oxidation/methylation technique to determine the lignin contents of different lightand dark-colored turnip rapeseed cultivars without interference from condensed tannins.76.75 This method has the advantage of being rapid and requires only about 50 mg of material.5_fm Page 99 Sunday. glucuronic (or its 4-O-methyl ether). 86 Cell wall proteins. May 6. the contents of these components were determined22 in some high-fiber products (Table 3..9 70. that phenolic acids. silica.2 71. 39. O.. Varo. Recently.2 2. when calculated as crude protein (N × 6.2387_ch3. Anal.3 0. Eds.5_fm Page 100 Sunday.4 0. Chem. Off. complexes.. . Stärke. So far.0 0. It is very likely. as well as lignin. and Westerlund. polyphenols other than lignin.4 1. which may be included in a broader definition of the term dietary fiber. in New Developments in Dietary Fiber.6 — — — Sources: Theander. P. J. is discussed in Theander and Åman. as determined by HPLC after removal by alkali. J. after release by alkaline treatment by HPLC. however. 273. we do not find more than 1 to 5% of the nitrogen of the original food.. 1987. In residues from unprocessed foods. cell wall protein.3 2. et al.8 4.2 0. 66.7 11.25). 1990. cutin..5 0. although generally present in only small amounts in human food.66% ferulic acid in wheat bran.5. and other nitrogen-containing components. Acetyl and phenolic acid substituents in grasses can be analyzed by GLC methods. For brans. lignin. O.6 Source: Theander. 2001 6:43 PM 100 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and other phenolic compounds may also play an important role relative to the microbial fermentation in the human gut. lignin. The acetyl constituents were quantified as 1-acetylpyrrolidin by GLC89 and phenolic acids. has been discussed mainly in connection with digestion of plant materials by the rumen microflora. C.5. and Brine. E.87 Hartley and Haverkamp88 reported the presence of 0. and compounds such as acetic acid and phenolic acids.02% p-coumaric acid and 0.84 Theander.8 0.85 and Selvendran. 3RD EDITION Table 3.3 1.5. Furda. et al.90 Table 3. the importance of such plant fiber substituents.4 37.4 2. 1983. The chemistry of DF polysaccharides. this corresponds to not more than about 10% of the Klason lignin residue.6 Content of Ester Substituents in Some High-Fiber Products (% of dry matter) Product Dietary Fiber Coumaroyl Groups Feruloyl Groups Acetyl Groups Maize bran Wheat bran Potato fiber Sugar beet fiber 76.7 1. 933.2 trace 0. phytic acid. Assoc. however.. waxes. which represent a very lignin-rich food ingredient. I.5 Klason Lignin Values in Heat-Treated Cereals and Potato Samples Klason Lignin (%) Sample Wheat Flour Untreated Extruded at 168°C Wheat Whole Meal Untreated Extruded at 180°C Potato Raw Boiled Pressure-cooked Crude Protein in the Klason Lignin (% of original protein) 0. may be included in this group. New York. and other nondigestible substances associated with plant cell walls.6). Plenum Press. which are present as ester-linked substituents to DF polysaccharides and/or lignin. 88.4 0.5 1.2 trace trace trace 2. The values determined for average total DF contents of these samples. whereas corresponding values for uronic acid residues and Klason lignin were higher. Recently.32 APPLICATIONS An obvious advantage of the Uppsala method is that the content of individual sugar residues and Klason lignin of the DF is determined. The average RSDR value was 8.1%). May 6. oat bran.2387_ch3.7). Method performance for determination of total DF was good (Table 3.6% for wheat bread to 84.8 to 64. for instance. 8-O-4′ and 8-5′ dimers dominated. and eight unknown products were analyzed for total DF and DF constituents in duplicate by each collaborator. carrot.93 In one of the oligosaccharides. when properties of DF are to be investigated and predicted. which were also released by mild acid hydrolysis of maize bran. The 5-5′. both phenolic acids of the dimer were esterified to arabinose.5. calculated on a dry matter basis.3% for wheat bran concentrate. Three main feruloylated saccharides were identified. Individual sugar residues (rhamnose. but the 8-8′ dimer was also present. mannose. galactose. i. A gravimetric method for determination of . xylose. The structure of these saccharides indicates that the heteroxylans in maize bran are covalently cross-linked through dehydrodiferulates.1–71. four from North America and five from Europe..5%. the same group has also isolated and structurally elucidated two new 5-5′-diferuloyl oligosaccharides. rye bread and wheat bread — were selected to represent a wide range in the content and composition of DF and to represent important foods with significance for the intake of DF. and Klason lignin. uronic acid residues.3 The study was completed by nine laboratories. which all had the same basic unit with an arabinofuranosyl residue esterified on position O-5 by ferulic acid. apple. green peas. which was further substituted with a 2-linked xylose residue. varied from 4. 2001 6:43 PM THE UPPSALA DIETARY FIBER METHOD 101 Markwalder and Neukom91 have isolated dehyrodiferulic acid from wheat flour. The content of neutral polysaccharide residues ranged from 3. Potex (an enriched potato fiber product).7). both for repeatability within laboratories (RSDr) and reproducibility among laboratories (RSDR).1% with an average RSDR value of 7.e.5. Such information is valuable. From this unit the other two saccharides were built by adding one xylopyranosyl residue on position 2 of the arabinose residue. arabinose.9%) and dehydrodimers of ferulic acid (2.92 By using controlled mild acid hydrolysis it was possible to release oligosaccharides with esterlinked phenolic residues still attached.8–11.8% of dry samples. These saccharides are probably side chains of the heteroxylan which have been released by the cleavage of the acid-labile glycosidic bonds of the arabinofuranosyl residues attached to the main xylan backbone.5_fm Page 101 Sunday. and in the other oligosaccharide one phenolic acid was esterified to arabinose and the other to arabinose. COLLABORATIVE STUDY OF THE UPPSALA METHOD The Uppsala method for DF analysis has been successfully applied to the analysis of a number of divergent samples and tested in a collaborative AOAC/AACC study. A collaborative study on an enzymatic gravimetric AOAC method has shown a similar range of RSDR values (0. The samples — wheat bran concentrate.5%) were released from maize bran by alkaline treatment. and glucose) generally also had good RSDR values when present above 1%. which is good considering that this enzymatic–chemical method is based on the determination of several analytes. the individual neutral sugar residues. where it is thought to be a possible cross-link in pentosans. Maize bran is rich in phenolic acid residues. and further one galactopyranosyl residue on position 4 of the xylose residue.2%) for food samples with a total DF content of 13. with generally small differences between laboratories (Figure 3.9–12.4% (range 4. Both ferulic acid (2. Assoc. (Source: Theander. 78. 2001 6:43 PM 102 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. et al. O.5_fm Page 102 Sunday. 1995. Off.) . Chem.7 Results from individual laboratories on determination of total dietary fiber by the Uppsala method... Figures given are a percentage of dry matter as an average of duplicate analysis. Anal.5. J. 1030. May 6.2387_ch3. 3RD EDITION 100 Wheat bran Potex Total dietary fiber (% of DM) 90 80 70 60 50 40 Carrot Oat bran 30 20 10 0 12 3456 78 9 12 3456 78 9 12 3456 78 9 12 3456 78 9 Laboratory 100 Total dietary fiber (% of DM) 90 80 70 60 50 40 30 Apple Green peas Rye bread Wheat bread 20 10 0 12 3456 78 9 12 3456 78 9 12 3456 78 9 12 3456 78 9 Laboratory Figure 3. This variation is due to cultivar as well as growing and processing conditions. Contents of individual dietary fiber components are presented in Table 3. The application of the Uppsala methodology on a series of fiber-rich foodstuffs.32 4. 1030. Galactose residues in Potex and arabinose residues in peas.9 11.10. Also. for products of legume seeds. were also notably high in content.8 5.6 4.9 16. is presented in Table 3.8 4.90 3. together with Klason lignin. mainly in the two latter samples. Anal.3 80.3 In concentrated wheat bran arabinoxylans (arabinose and xylose residues). The other sources have higher cellulose contents and also more pectic and associated substances (galacturonic acid.5..7 10. namely wheat bran. whole potato. cellulose (glucose residues) and Klason lignin dominate.5. cellulose (glucose residues).3 4.5_fm Page 103 Sunday. and one would therefore also expect great differences among the physical.52 2. The great differences in chemical composition among the different foods are notable.93 0.02 0..5. Assoc.7 103 Method Performance for Determination of Total Dietary Fiber by the Uppsala Method Sample Mean (% DM) sr sR RSDr RSDR Wheat bran Potex Carrot Oat bran Apple Green peas Rye bread White bread 84. rhamnose. quality control.90 0.6 9. and enzyme-resistant starch.1 Source: Theander. May 6. 1995. On a fresh weight basis.86 2.9% in enriched wheat bran.6 5.2387_ch3.9 5.7% in corn flakes to 84. rye bread. Chem. white cabbage. . O. mixed-linkage β-glucan (glucose residues) and cellulose are important DF components. In oat bran.50 1. and galactose being typical constituents) than brans. 2001 6:43 PM THE UPPSALA DIETARY FIBER METHOD Table 3. mainly in the two former samples. and nutritional properties of the different DF sources.52 1. potatoes.6 1. and products thereof are given in Table 3. pea. vegetables. carrot.48 0.1 The figures based on dry matter are in the range of 1 to 4% for water-soluble DF and 8 to 34% for insoluble DF. large variations in content were found.8 10. Off.8.16 1.4 17.43 1. and white bread arabinoxylan. and xyloglucan (glucose and xylose residues). The analyses thus indicate that brans have the highest Klason lignin values and are rich in arabinoxylans and cellulose but have a low proportion of uronic acid constituents. the brans are particularly rich fiber sources. fruits. In cereal products total DF ranged from 2. and green peas are pectin (uronic acid residues).7 8. Total DF results published from our applications during many years using the Uppsala methodology on batches and/or varieties of cereals. Major components in Potex.4 24. total DF can never provide such information. lettuce. The Uppsala method consequently provides a tool to determine the DF content and composition for the purposes of research.8 8.04 0. et al. and labeling. and fruits. vegetables. both probably part of the pectic polysaccharide complex. arabinose.7 6.0 5. The high galactan content is also typical for potatoes. carrot. The large ranges in total DF within the different sources demonstrate that care must be taken when generalizing from a single analysis of a certain DF source. rye bran.8 3.4 7.5.1 18. and apple. apple. 78. J.42 1. biological.9.6 4.58 1.88 1. 5 23.3 23.5 13.7 21.0 12.3 78.9–24.4 10.8 13.0 73.1–23.8–13.7–83.9 6.0–89.2–11.3–23.5 10.2 15.1 20.6 40.0 15.7–4.1 8.0 1 34 6 13 1 2 1 2 1 1 1 5.7 2.7 15.5 14.1 12.6 9.5–7.0 84.0 1 2 1 2.5–14.5–17.8 10.5–46.3 11.0–15.7 10.3–18.4–80.9 17.1 25.5–28.5–16.3 10.8 22.3 3.0–14.4 20.6 16.0 10.9–3.6–9.1 19.7 21.9 13.2 1 1 2 5 7 16.2387_ch3.0 23.5.9 1 5 3 49 1 1 1 15 1 1 14 10 6 2 6 2 12 18 4 2 16 8 10 8.8 Contents of Dietary Fiber (% of dry matter) in Products of Cereals. Potato Products.8 25.5 10.2 17.3 4.9–3.7 36.7 84.8–38. Legume Seeds.3 13.2 2.9 14.2 8.9 11.5_fm Page 104 Sunday.5 7.1 13.0 6.8–9.8–28.9 .8 29.5 7.1 64.7 3.6 9.4–16.5 4.2–26.9 18.8 19. and Fruits as Determined by the Uppsala Method Samples Cereal Products Corn flakes White bread crust White bread crumb White flour White bread Corn Rye bread Whole winter wheat Dehulled oats Oat bran bread Whole spring wheat Whole triticale Rolled barley flakes Rye biscuit Whole rye Rye crisp bread Dehulled barley Whole barley Oat bran Rye bran Whole oat Wheat bran Wheat bran concentrate Legume Seeds Dehulled broad bean Dehulled yellow peas Dehulled dark peas Dehulled soy bean Dehulled brown bean Green peas Soy bean Frozen green peas Mung bean Brown bean Chick pea Potato Products Raw potato Potato powder Enriched potato pulp Vegetables Deskinned onion White cabbage Lettuce Carrot Sugar beet fiber Fruits Apple pulp Apple Citrus pectin Total Dietary Fiber n Range 2.1 15. May 6.9 24.0 81.7 9.7–20.2 3.0 23.7 13. Vegetables.9–13. 3RD EDITION Table 3.2 14 1 12 68. 2001 6:43 PM 104 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.8 19. 1 1.3 — 0.8 — 28.4 0.1 15.2 25.3 0.2 11.1 0.5 1.3 8.0 tr 5.5 0. are given as a percentage of dry matter.7 2.2 9.6 16.3 3.7 17.9 0.8 2.6 Source: Theander.5 3..7 1.2 20.5 1.6 11.2 4.6 1.4 — 1.3 1.8 0.5 — 10.4 0.7 17.0 14.9 0. O.3 11.5 tr tr 28. Res.7 — 1.2 4.3 31.1 tr 1..6 0. Assoc.9 11.1 0.4 0.0 1.2387_ch3.0 69.5 2.9 11.5 28.5 tr 16.3 tr 30.5 4. and Åman.6 0.7 0.9 27. Table 3. 1995.4 tr 20.6 80.1 46.9 6.6 3.4 9.7 4.8 60.5 5.3 0.5.2 1.7 51.5.9 — 3.7 63. Chem. 1979.3 24.5 0.5 30.10 Chemical Characterization of Water-Extractable and Water-Unextractable Dietary Fibers in Various Foods Water-extractable fraction Neutral sugar residues Uronic acid residues Relative composition of neutral sugar residues Glucose Galactose Mannose Xylose Arabinose Rhamnose Fucose Water-unextractable fraction Neutral sugar residues Uronic acid residues Klason lignin Relative composition of neutral sugar constituents Glucose Galactose Mannose Xylose Arabinose Rhamnose Fucose Total DF content Wheat Bran Rye Bran Potato (whole) Carrot Pea White Cabbage 2. Apple .8 2.9 — 8.4 4.7 0.4 26.9 2.6 9.8 29.3 1.4 0.6 1. O.5 1.0 1.9 0. Off.9 105 Content and Composition of Dietary Fiber in Some Food Products Individual sugar residues Rhamnose Arabinose Xylose Mannose Galactose Glucose Uronic acids Klason lignin Total dietary fiber Wheat Bran Potex Carrot Oat Bran Apple Green Peas Rye Bread White Bread — 14.3 40.0 40.0 4.8 26.6 1.2 2.1 34.5 9.0 6.3 62.5 2.4 1. Anal.4 2.3 10.3 1..4 tr 3.5 tr tr 36.3 39.5 0.8 — 24.8 tr 39.7 2.0 5.9 tr 8.9 0.1 17.0 10. 1030.3 34.2 0.5 2.8 4.3 9.9 28. Source: Theander.1 6.0 0.5 6.2 24.3 41.4 0.5 0.1 9.9 18.1 2. Agric.3 2.9 1.4 0.5_fm Page 105 Sunday.3 20.0 0.2 15. May 6.5 1.4 0. P.3 3.0 1.9 14.8 34 tr — 36.2 15. except composition of neutral sugars.4 0.0 9.5 84.6 1.1 5.9 10.7 0. 2001 6:43 PM THE UPPSALA DIETARY FIBER METHOD Table 3. 97. J.6 2. 9.8 3.3 3.3 1.9 4.9 24.7 23.2 46.5 Lettuce Note: All figures.9 8. 78.7 1.0 0.3 21.3 17.4 3.5 1.4 15.2 tr tr 58.6 0.2 6.8 1.8 9.5 12.8 0.9 5. et al.0 1..4 6.7 1.7 2.0 22.0 tr 21.1 4.2 1.9 40.7 1.8 10.1 — 1.2 1.9 45.3 1.5 1. a Traces. Swedish J.8 — tra 38.6 3.3 tr tr 5.4 0.4 0.5 tr — 45. . T. 4. A. Sci. Commentary on results obtained by the different laboratories using the Southgate method. 1995. A review of the different analytical methods and remaining problems. 340. Asp. T. P. W. Food Agric. and Åman. J. T... H.. J. II. and Theander. Englyst.. 3RD EDITION REFERENCES 1. A comparison with gas-liquid chromatography. A. T. 9.. E. Eds. 16. New York. Miller Jones. 22. H. 63. E.. 2001 6:43 PM 106 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 1990. 1979. chap. Plenum Press. O. P. A method for the analysis and chemical characterisation of total dietary fibre. The method specific certification of the mass fraction of dietary fibre in lyophilised haricot beans. and Hudson. 1986. digestive fate & metabolic effects.. 19. Guillon. Southgate. Analyst. 33. Off.. M. P. Marcel Dekker. 20. chap. 1997. T. 1. Food Chem.. Marcel Dekker. Englyst. Assoc. and Åman.. N. Analysis and chemical characterization of water-soluble and water-insoluble dietary fibres. Am. A. R. Furda. 1030. 71. 1988. O. Gordon. O.. Eds. Furda. N. 12. Assoc.. P. B.. 307. J. apple. H. full fat soya flour and bran breakfast cereal reference materials. J. Biology & Environmental Sciences. O. Dietary fiber definition and analysis. Åman. Determination of dietary fibre as nonstarch polysaccharides with gas-liquid chromatographic. 5. and Westerlund. Cereal Foods World. 2000. Eds. 331. H. and Brine. and Brookes. N. H. 1996. Unavailable carbohydrates. O. 1991. Englyst. Englyst. 808... P. Swedish J. in New Developments in Dietary Fiber. James. C. in The Analysis of Dietary Fiber in Food. Chem. E. P. T. 34. BCR Information. and Theander. O. Res. Pendlington.. T. and Prosky. 404. H. Analyst. and Cummings. H. Eds. N. III. Analysis of dietary fibers and their main constituents.. Studies on dietary fibre. 2. in The Analysis of Dietary Fiber in Food. S. Determination of carbohydrate and its composition in plant materials. James. D. Studies on dietary fibres.. Westerlund. 9. Marcel Dekker. O. O. L. et al. and White. Colorimetric method for routine measurement of dietary fibre as nonstarch polysaccharides. Off. 688. 8.. Southgate. 1981. 21. 18. chap. James. W. O.. G. 13. 1497. D. Marcel Dekker. J.. 205. W. P. 16. Eds. Schweizer.. New AACC dietary fiber definition sets the stage for annual meeting technical round tables. Total dietary fiber determined as neutral sugar and uronic acid residues. 11. Chem. Meuree-Vanlaethem. Food. Sundberg. 3. Theander.. A. carrot. 15.. and Theander. 336. Nantes. in The Analysis of Dietary Fiber in Food. The Analysis of Dietary Fiber in Food. J. and lignin (The Uppsala method): collaborative study. 10.. 330.. May 6. Determination of carbohydrates in foods. Quigley.. J. 7.. O. and Theander. 1999. New York. in The Analysis of Dietary Fiber in Food. 1988. H. W. L. 4. and Cummings. W. function. Eds. and Pettersson. Chem. in Proceedings of the Profibre symposium: Plant polysaccharides in human nutrition: Structure. P. Englyst. New York. 17. Studies on dietary fiber. 48.. P. P. New York.. 5.. Englyst. Sci. O... Improved procedures for analysis of dietary fiber. W. M. New York. 24.. T. New York. 20. 1981. Food Agric. T. Theander. J. Reference Materials.. Nonstarch polysaccharides (dietary fiber) and resistant starch.. Agriculture. 1981. Current Contents. Marcel Dekker. Report EUR17451EN. Åman.. . F.5_fm Page 106 Sunday. H. Nutr. O.2387_ch3. 1994. and Hudson. Citation Classic. 32. Theander... James.. 1981. 78.. and Cummings. James. Theander. 1982. Agric. and Theander. 1981. Andersson. 6. and Andersson. Theander. Cereal Foods World.. Theander.. 1969. Theander.. Anal. high-performance liquid chromatographic or spectrophotometric measurement of constituent sugars. G. N. Agric. 45. 107. P. 44. R.... N. Clin. H.. J. Defining dietary fiber — A progress report. Analysis of cabohydrates in ileal excreta from humans. Informative analysis of dietary fibre. F. D. An improved method for the measurement of dietary fibre as nonstarch polysaccharides in plant foods. European Commision. and Åman. Eds. 1987. A. 119. J.. European Air Concerted Action AIR3CT94-2203. chap. 6.. D. 14. Wiggins. I... J.-G. Brussels. 1982. Anal.. 9. J.. Determination of the nonstarch polysaccharides in plant foods by gas-liquid chromatography of constituent sugars as alditol acetates.. J. and Åman. 97.. H. 46. 2001 6:43 PM THE UPPSALA DIETARY FIBER METHOD 107 22. and Graham.. J. Stärke.. Digestibility of amylose-lipid complexes in-vitro and in-vivo. Eds. Anim.. Academic Press. 1044. 23. Stärke. Ed. Andersson. J. 32. 26. Measuring dietary fiber. and Cummings. E. Agric. 39. J. R. E. E. Studies on chemical modifications in heat-processed starch and wheat flour. W. I. in Methods in Carbohydrate Chemistry. B. 1011. A rapid method for determining the carbohydrate component of dietary fibre. 10. 34. O.. 63B. Westerlund. O.. Acta Chem. May 6. J.. J. Chem. Simplified methods for the preparation and analysis of dietary fibre. Larm.. New York. Agric. Ring. 75. Kvantitativ kolhydratbestämning — en detaljstudie. A.. P.. 1983. and Westerlund.. Graham.. Sci. R. M.. Åman. Effects of baking on polysaccharides in white bread fractions. H. J... F. R. J. a solvent for hemicelluloses. AOAC Int. E. 1980. 23. 10. R. W. 1173. 1434. N. Determination of total.. M. Dimethylsulphoxide. K. . L. Anal. J. J.. soluble and insoluble dietary fiber in foods — Enzymatic gravimetric method. Marlett. Theander.. Scand.. O.. 294. Extraction of soluble dietary fiber. 1992. F. A. P. 1989.. Food Chem... New York. Longaere. Grön Rydberg. Batey. 25. Rådeström. 12. Academic Press. 1983. 1963. J. N. 273. J.. 34. and Du Pont. 41. P. I. 28. Linskens.-G.-B. 40... and Åman. S. Hägglund. 35. 42.-C. 77. Plenum Press. E. J.. E. 326.. E. C. Sugar units present.. Enzymatic/chemical analysis of dietary fiber... 1989. 1983. Starch and nonstarch polysaccharides in some cereal foods. S. A. 234. Swed. 933. and Lundquist. 38. Chemistry of Wood..-G.. 3. Stärke. Moore. and Harland. Transglycosidation reactions following heat treatment of starch. 1990. Hydrolysis and quantitative paper chromatography.. F. 1976.. in New Developments in Dietary Fiber. Holm.. F. Determination of the dietary fiber content of the EEC samples and a discussion of the various methods of analysis. Siljeström. Eds. Springer Verlag. 479. Furda... L. 1989. 67. R. New York. J.. J. and McPhersson... 34. 7. Biochem.. O.. chap. O.. J. Nutr.. Lee. Starch analysis using thermostable alfa-amylases.. Assoc. Feed Sci. Food Agric. O. Determination of total dietary fiber in foods. Recovery of soluble dietary fiber is dependent on the method of analysis. J. L. Asp.. 14.. Methods for analysis of dietary fibre... Acta Chem. B. Sloneker. P.. J. Res.. AOAC Int.. Schweizer. vol. Scand. 1994.. and Westerlund. 1975. and Brine. Cereal Sci. 33. Åman. Anal. 273. chap.. P.. Asp. 45. Prosky. S. M. and Faulks. Am. Eliasson. 125. 43. 36. 50. in Plant Fibers: Modern Methods of Plant Analysis.. Theander. M. and Bogdanske. The Uppsala method for rapid analysis of total dietary fiber. S. Selvendran.5_fm Page 107 Sunday. Larsson. T. Food Agric. L. H. and Theander. Larm. Stärke.. L. 1981. Prosky. P.. J. 1. vol. 47.. M. H. 1987. M. 1984.. Åman. Saeman.. K. Scand. Selvendran. R. Varo. H. Marcel Dekker. and Du Pont... W. 35. J. Whistler. J. E. O. Off. P. Food Chem. and Timms.. and Åman. 1160. Westerlund. V. 43. H.. and Åman.. Devries. Furda. 1951. Chesters. and Hesselman. Structural studies on a water-soluble arabinan isolated from rapeseed (Brassica napus).. James. 149. R. O.. Lindberg. 1988. Hägglund. Andersson. A. 1956. F. J. Theander. R. Off. B. Chem. R. 37. Björck.. R. 31. Englyst. C. 29.. 31.. P. M. Sci. T. Assoc. Effects on enzymic digestibility.. Acta Chem.. Theander. 135. 395. 30. 539. 494.. R. 44. 39. 30. H. J. New York.2387_ch3.. Björck. O. J. Theander. mes-tris buffer — Collaborative study.. and Graham. 1989. M. and Åman. 10. O.. Marlett. and DeVries. Berlin. Bethge.. J. Communication No. and Jackson. Laine. 29. Eds. J. and Koivistonen. Westerlund. 17. W. 24. Clin. Anal. 1971. Ostrowska. P. P. 1971 (in Swedish). J. I. S. Analysis of starch and other main constituents of cereal grains. 36. P... Selvedran.. Theander. B. R.. B... 66. V. Determination of cellulose and apparent hemicellulose in plant tissue by gas-liquid chromatography.... G. A. 1989.. 27. N. Stockholm. Faulks... 1982.. Verne.. I. Structural studies on a water-soluble arabinogalactan isolated from rapeseed (Brassica napus). S. 95.. 627. 88. Swedish Forest Products Laboratory.. V. 1985. O. 1984.. Technol. and Millet. food products and ingredients: collaborative study. L. Effect of heat treatment on dietary fiber: interlaboratory study. G. 41. in The Analysis of Dietary Fiber in Food. 703. and Theander. Carbohydr. High-performance anion-exchange chromatography for carbohydrate analysis.. and Ternrud. 261.. 62.. 140. Food Chem. B. S. N.. Hoebler. N. A simple and rapid preparation of alditol acetates for monosaccharide analysis. Biochem. Determination of aldoses and uronic acid content of vegetable fibre. Miksche. and Yasuda. Isomerisation of D-glucuronic acid in neutral aqueous solution. Academic Press. European Commision. Colorimetric determination of hexuronic acids in plant materials. New York.. Res. Acta Chem. Food Chem.5_fm Page 108 Sunday. 1988. Hicks.. 96. Anal. M. 65. Anal. 37. J. K. Liquid chromatography of sugars on silica-based stationary phases. 4. and Delort-Laval. J. 556. and Samuelson. J. A. Agric. R. Automated chromatography of uronic acids on anion-exchange resins. R. 35. O. 31. Åman. and Graham. Honda. Svensk Papperstidn. and Asboe-Hansen. Agric. L. J. Scott. Eds. R. High-performance liquid chromatography of mono.l. 1978. 1968. 1977. chap.2387_ch3. Carbohydrates.. Gas-liquid chromatography of alditol acetates. 151. and Muir... O.. 291. and Thede. Anal. Lee. 55. in The Analysis of Dietary Fiber in Food. M. New method for quantitative determination of uronic acids. J.. 505. Agric. Partition chromatography on ion-exchange resins. T. 1972. A. Anal. Carbohydr. 9. chap. Westerlund. Theander. 73. T.. 1542. T. and lignin in seed hulls of different colors from turnip rapeseed. Automated ion exchange chromatography of organic acids in acetate media. Academic Press. Chem... 54. Chim. Åman. A. 1967. 313.. H. Agric. Samuelson. R. 291. 15.. 410. 189. 270.. 115. 1342. Lehrfeld. 1. 163. 936. 51. J. T. C. Biochem. 4. 360. J. 1969.. Anal. 25. 73. 67. New York.. R. K.. Blumenkrantz. 1989. Eds. Food Chem. 60. O.. M. Hämiläinen. O. Quantitative determination of carbohydrates in cellulosic materials — losses as sulphates. 1962. J. Multivariate calibration in the determination of acetylated aldoses by g.. Popoff.. 1990. Theander. 30. Selvendran. 167. 17. 46. L.. and Marlett. N. James. Harris.. 1973. using high-performance liquid chromatography to measure neutral sugar composition. H. P. H. Adv. Chem. O.. 1990. N. S. 58. 49. A.c. M. J... Bitter. 1987. K. J. A. 50.and oligosaccharides. 71. J. in Recent Progress in the Analysis of Dietary Fibre. Determination of uronic acids as components of dietary fibre polysaccharides. (1-4)-beta-D-glukans in barley and oats. 14. Svensk Papperstidn.. Anal. M. W. New York. Food Chem. Barry. in Methods in Carbohydrate Chemistry. P. 23. Åman. G.. J. 1972. 1981. in Methods in Carbohydrate Chemistry. M. Henry. J. 52. 220. 1978. 54... F.... and Theander. Anal.. O. 1966. O.. Mopper. 1981. A modified uronic acid carbazole reaction. L. chap. J. Eds. P. and Van de Bovenkamp. Samuelson. 2001 6:43 PM 108 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and Pandit. 72... E. David. 66. 1984. 1979. 1970.. Scand. Semimicro determination of uronic acids. A.. Whistler. Verhaar. 69. J..... Biochem. Whistler. 6... I. 64. A. 85. Biochem. H.. and Kuster. Andersson.. O. Marcel Dekker. Biochem. P. vol. polyphenols. Neilson.. . B. 1983. and BeMiller. L.. Determination of total dietary fiber by difference and of pectin by colorimetry or copper titration. High-performance liquid chromatography of carbohydrates. 61. 207. P. E.. Johnson.. Differential gas-liquid chromatography method for determination of uronic acids in carbohydrate mixtures. 1979. G. 330. 53.. Connors. Sloneker.. March. Anal. 51.. S. and Theander. P. 56. 113... 63. Bylund. 1. F.. 70. B. 36. 528. Carbohydrate Res. and BeMiller... E.. Katan. S. N-Methylimidazole as a catalyst for analytical acetylations of hydroxy compounds. Brussels.. Analysis of total and insoluble mixed-linked (1-3). Chromatogr. 1... and Ring. 1983. 3RD EDITION 48. Chromatogr. vol. Acta. 484. R. Partition chromatography of sugars in ethanol.. 704. B. 50. 68. and Donetzhuber. Y. 59.. 1994. R. Carlsson. Blakeney. J. K.-L. Biochem. B. W.. May 6. Anal. J.. 282. Samuelsson. 1981. 57. and Stone.. 6. Nordkvist. Rapid acid hydrolysis of plant cell wall polysaccharides and simplified quantitative determination of their neutral monosaccharides by gasliquid chromatography. O. Improved chromatographic separations on anion-exchange resins. Chem.. C.. B. Biochem.. Hardell. and Theander.. and Theander. L. G. A comparison between detergent and non-detergent analyses of dietary fiber in human foodstuffs. Anal. O. Biochem. P. K. Aussprache. 76. 1451. McConnell. and Åman. J. J.. 1970. Klason. Saulnier. Acetyl and phenolic acid substituents in timothy of different maturity and after digestion with rumen microorganisms or a commercial cellulase. H.. 1986. Continuous changes in straw carbohydrate digestion and composition along the gastro-intestinal tract in ruminants. O. J. J. and Schweizer. Assoc.. 92.. Determination of lignin.. P. P. Review: Ferulic and diferulic acids as components of sugar-beet pectins and maize bran heteroxylans. 82... L. 85. Interscience. Theander. M.. Sci. London. 1999. Eds. M. 249. 75. A. and Neukom.. Food Agric. R15. G. M. 86. 23. W. M. D. M. P. P. Agric. G. L. and Yeoman.S.. 1974. Use of detergents in analysis of fibrous feeds. I. New York... morphology.. Government Printing Office. 82. Birch. Forage fiber analyses. chap. Van Soest. 52. Sci. Off. J. O. 793. 1983. Sci.-F. A. Hauptversammlung. E... J. Theander. G. and Thibault. Isolation and structural determination of two 5. Svensk Papperstidn. Pernemalm. U. P.... Birch. Johnson. K. Quantitative determination of O-acetyl and other O-acetyl groups in cellulosic material. 1981. O. 1978. 2001 6:43 PM THE UPPSALA DIETARY FIBER METHOD 109 74. A.. Study of effects of heating and drying on yield of fiber and lignin in forages. Diferulic acid as a possible crosslink in hemicelluloses from wheat germ. An investigation of lignin extraction from dietary fibre using acetyl bromide. Acta Chem. London.-F. New York. chap. 80.. Amado. and Åman. 88. C.. 1976. Applied Science.. 320. 315. 77. Sci. J. A. Anal. 14. 1987. Moore..... III. 91. 455. K. P.. T. in Dietary Fibre. J. in Dietary Fibers: Chemistry and Nutrition. O. Lahaye.. 81. J. Månsson. Theander. . O. in Nahrungsfasern Dietary Fibres. H. Udén..C. 32. E.. Hartley. 78. The chemistry of plant cell walls. Agricultural Handbook No. and Theander. Phytochemistry.. 1961. Reaction of D-glucose and glycine in slightly acidic. S. 1965. B. 379. 13.. K. J. B.. Garcia-Conesa. Thibault. in Verein der Zellstoff and PapierChemiker and Ingenieure. Pyrolysis-mass spectrometry of the phenolic constituents of plant cell walls. The spectrophotometric determination of lignin in small wood samples. May 6.. 84. Washington. 35. 70. L. U. 44.. Browning. Scand.. Theander. Chem. 2. J. 836. G. Olsson. 41... and Zank. E. 1999. Sci. Goering. 34. 1984. and analysis of dietary fiber components. and Eastwood. Inglett.5′-diferuloyl oligasaccharides indicate that maize heteroxylan are covalently crosslinked by oxidatively coupled ferulates. Robertson.. and Theander. Die Verfahren der Holzzellstoff-Fabrikation. Dept. Berlin. 48. 1981. P. I. P. 25. 1039. L. Advances in the chemical characterisation and analytical determination of dietary fibre components. 89.5_fm Page 109 Sunday. Res. Academic Press. 127. J. Saulnier. chap. TAPPI. M. and Parker. Agric. Food Agric. I.. 15. G. A. 1972. Markwalder. B. vol. Eds.2387_ch3.. Academic Press.. 95. Ternrud. Food Agric. Formation of aromatic compounds from carbohydrates. Food Agric.S. and Parker.. 1967. Carbohydr. and Falkehag. Sci.. O. and Van Soest.. H. Eds. J. D. V. G. 93.. M. 79. 785. 1979. D. E. Environ. J. London.-J. The chemistry. 396. 1979. R. 87. 83. Food Agric. A semi-micro method for the determination of lignin and its use in predicting the digestibility of forage crops.. R. 84.. R. and Williamson. Eds.-A. Lindberg. and Haverkamp. in Methods of Wood Chemistry. B. Selvendran. VII. 6.. J. aqueous solution. 30. 6. 90.. and Samuelsson. Crepeau.. 15. Kroon... Morrison. The chemistry of dietary fibres in different sources. M. Applied Science Publishers. in Dietary Fibre. Food Agric. 1983. R... A comparison of methods of measuring “fibre” in vegetable material. Eastwood. 1908. J. 2001 6:43 PM . May 6.2387_ch3.5_fm Page 110 Sunday. Figure 3.6 The Crude Fiber Method1 Ivan Furda Crude fiber is the loss of ignition of dried residue remaining after digestion of a sample with 1. 0-8493-2387-8/01/$0.25% sulfuric acid and 1.25% sodium hydroxide solutions under specific conditions. and any fiber-bearing material from which fat can be extracted to leave workable residue. feeds. 2001 2:58 PM CHAPTER 3.2387_ch3.6_fm Page 111 Tuesday. The principle of the method is that a finely ground air-dried sample of the food is extracted with ether to remove lipids and that this dry sample is then extracted successively with boiling acid and alkali. flours.50 © 2001 by CRC Press LLC 111 . After drying and weighing. The method is applicable to grains. cereals. The residue is filtered off and washed.00+$1.6. the residue is ashed and residual inorganic matter is measured.1 Flow chart for determining crude fiber. May 8. May 6.2387_ch3. 132.. W. Official Methods of Analysis of the Association of Official Analytical Chemists. AOAC. . Ed. 3RD EDITION REFERENCE 1.6_fm Page 112 Sunday.. Washington.C. Horwitz.. 13th ed.. 2001 6:46 PM 112 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 1980. D. This concentration of acid is just 0-8493-2387-8/01/$0. Other qualitative separations were effected by Tomlinson and Ballou10 using Dowex 1 (Cl form). Cl form. These methods have been reviewed by Oberleas. The reader interested in the history of phytate analytical methods is directed to the above references for details. A later modification of this procedure by Sobolev and Vyskrebentseva8 utilized ammonium thiocyanate as the colored ironcomplexing reagent with similar results.11. There are no known reagents specific for the identification of phytate. are necessary for the study and understanding of phytate. Anion-exchange techniques are available for the separation of mixtures of isomeric inositol polyphosphates. 200–400 mesh) with a non-linear gradient of water and 1.1 Cosgrove. Harland Good analytical methods. The chromatograms were then sprayed with an alcoholic solution of sulfosalicylic acid.13–15 A long column packed with Dowex 1 and eluted with 0.4 Oberleas. Extensive efforts were made by Cosgrove. The elution was 32 mL per hour with 1-hour fraction collection for up to 280 h. ferric sulfosalicylate formed a colorful complex. both qualitative and quantitative.1 to 0.25 × 24 cm resin bed and used a step gradient elution of 0. color did not develop.6 By current standards and technology.7 Newer Methods for Phytate Analysis Donald Oberleas and Barbara F. Nine distinct fractions were separated and identified. This paper chromatographic technique separated phosphate esters that were sprayed with a mixture of ferric ions in dilute acid.5 and Oberleas and Harland. Where free ferric ions were present. The nature of the compound with its saturated ring does not lend itself to absorption within the ultraviolet (UV) or visible spectra. 2001 6:47 PM CHAPTER 3. but this represented an impractical analytical technique. and where ferric ion was complexed with phosphate or phosphate esters.48 N HCl worked well.2.7_fm Page 113 Sunday. Anion exchange columns were employed in the separation and identification of phytate and inositol phosphates even before the principles and benefits of this technology were fully understood. May 6. many methods were tested throughout the twentieth century to achieve accurate identification and quantitation of phytate.2387_ch3. Smith and Clark9 selected a weak-anion exchange resin (60–80 mesh) in a 1.3 Maga.50 © 2001 by CRC Press LLC 113 .2 Cheryan. some methods now have little more than historic value.5 N HCl. QUALITATIVE METHODS A detection system was devised by Wade and Morgan7 that takes advantage of the strong complexation of inositol phosphates with the ferric ion.00+$1.8 N HCl.12 whose earliest efforts used Dowex 1 (X8. Therefore. by many investigators as opposed to an extraction solution. 2001 6:47 PM 114 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. whereas Averill and King20 reported slightly better results with 2% HCl with a 25/1 (v/w) ratio and a 3-hour extraction time. Finally. and with careful separation and specific reagents. Bos et al. identify. Yet. Wheeler and Ferrel23 extracted wheat and high-protein flour concentrates with 3% TCA and compared these results with 0.2% HCl for extraction from food samples. in various concentrations.13.7_fm Page 114 Sunday. the acid concentration was diluted to 0. It must then be separated from other soluble and thus extractable constituents. some roots and tubers. the addition of water-soluble . and quantify the various inositol phosphates simultaneously. they are readily soluble.66 mol/l) gave a more complete extraction in a 1-hour extraction of phytate from rapeseed. it is possible to separate.8 mol/L HCl provided consistently higher extraction rates.6 mol/l HCl with extractions of 30 to 45 min of shaking.16.2387_ch3.25 selected this extraction procedure when they attempted their pioneering studies of adapting an HPLC procedure to phytate analysis. Thus. Rather19 recommended 1. The original method reported by Heubner and Stadler18 was based on the principle that phytate was the only phosphate compound found in nature that formed an insoluble complex with ferric iron in dilute acid.22 used 5% TCA with centrifugation and re-extraction of the residue. any of the above conditions may be appropriate for a specific group of foodstuffs. and some fruits. Cilliers and van Niekerk24 reported that 3% TCA was the preferred extractant. assurance of complete extraction is the imperative first step in the analytical process.4% HCl (0. In all cases. Phytate does complex heavy metals with some specific characteristics. The quantification of phytate may be divided into three phases.17 QUANTITATIVE METHODS Quantitative analytical methods underwent a considerable evolution within the twentieth century. Tangendjaja et al. defining a dilute acidic solution. Also. Being a natural compound in plant seeds. There appears to be little agreement with regard to the type or concentration of acid appropriate for extraction.26 reported that extraction with 0. but this is not specific and can be expressed only as phytate equivalent or phytate phosphorus. it must be detectable either in toto or some component that is proportional to the concentration in the original foodstuff. Little effort has been extended toward finding an appropriate acid and concentration that can be utilized universally in all situations.6% HCl worked with isolated phytate rather than phytate extracted from foodstuffs. Though the salts are slow to dissolve in water. Heubner and Stadler18 titrated isolated phytate samples in 0. De Lange et al. With modern HPLC technology. Several other acids. Free phytic acid is not found in plants because it is too strong an acid and phytate will complex with several minerals at neutral or slightly acidic pHs. Several modifications of this original method are recorded in the reviews cited above. 3RD EDITION sufficient for the elution of inositol hexakisphosphate and the subtle differences in isomeric configuration effects the differences in separation. Titration with a standard ferric iron solution against a sensitive ferric ion indicator identified the endpoint in this method.12. Extraction Methods Phytic acid and its sodium and potassium salts are hydrophilic and hygroscopic. the properties of these metals and their complexing characteristics can be made quantitative for phytate and other inositol phosphates. This has served as a working acid concentation. Latta and Eskin21 indicated that 2. Most of these complexes are pH-sensitive. the sample must be extracted from a complex set of matrices. for accuracy in any quantitative technique. May 6. Phytate contains phosphate that can be quantified. A reliable standard must always be considered important in quantitative analytical procedures and is essential for phytate analysis.6% for titration against an ammonium thiocyanate indicator. have been used by investigators for specific extractions. this lengthy extraction time was not essential. Tangendjaja et al.66 mol/L HCl were not different. in retrospect. May 6. whereas Young31 did not address the extraction time issue for her analytical method.33 and Cilliers and van Niekerk24 each used a 2-hour extraction time with 3% TCA as the extracting reagent.4 5085. whether that be as mixed salts or associated with proteins.to 16-hr extraction time. First.1 shows the results of one effort for comparison among an almost endless variety of possible combinations. The shortest extraction time reported was the 5-min blending of potato or other plant tissue with 100 mL of 10% TCA in a Waring Blender.2387_ch3.5 and 0. Thus. Second. The phytase activity inherent in the foodstuff would be inactivated by the acid and would not become a factor in the extraction process..7 1627. however. the phytate is stable in the acid for several days at room temperature.5 6557. but extraction was with 0.9 8014.4 2965. the hydrophilic nature of phytate dictates that an aqueous acid is necessary and that the fat content of the sample may interfere with the extraction process. 10% TCA. the fat content of samples must be low or reduced before extraction with aqueous acid. In none of these studies were objective data provided indicating an ideal extraction time. Thus. Table 3.6 4717. Fat extraction may be done with diethyl ether27 or petroleum ether28 for 1 h or more. Extraction Time Extraction time is another important consideration for which very little objective data are available.1 1506. 0. Though no designed formal studies of fat content have been made.7. McCance and Widdowson30 used a 2-hour extraction period for their analyses. The important aspect of extraction is to recover as much of the phytate as possible from all possible sample matrices with the conditions utilized. once solubilized. overextension or overextraction does not occur.6 2686.1 Evaluation of Various Extraction Acids of Wheat Cereals for Phytate Analysis (mg phytate/kg air dry cereal) Sample Name Shredded Grain Bran Fortified Shredded Mini Wheat 3% TCA 5% TCA 10% TCA 0.5 mol/L HCl 0.25 Camire and Clydesdale. “vigorous” physical agitation of the mixtures does not improve the chemical solubility or separation of the phytate from the organic matrix (foodstuff). The lipophobic nature of phytate will preclude any loss of sample during fat extraction. continuous shaking.66 mol/L HCl 4798.2 1009. Regardless of the extraction method. Oberleas27 suggested that overnight extractions would represent an 8.5 N HCl.7 3775. It should be obvious that these salts would be recovered under the mildest of conditions. Table 3.2 914. extraction is the dissociation and acid solubilization of phytate from its native matrix. Since too much extraction is a non-issue. Harland and Oberleas32 considered a 2-hour extraction period appropriate. With ideal . Paired “t ” statistical analysis indicated that 3% and 5% TCA is not as effective as other extractions.8 5214.3 6379. experience has shown that fat content less than about 5% poses no problem with the acid extraction process. Only Young31 indicated subjectively that repeated extractions did not improve the analysis. When incorporating the anion-exchange concept into quantitative phytate analysis.7 1706.3 3833. Thus.7.8 6623.29 Averill and King20 analyzed a large number of foodstuffs using a 3-hour extraction period under their conditions.1 3270.7_fm Page 115 Sunday. certainly the extraction time was not a factor in the analyses.4 6299. 2001 6:47 PM NEWER METHODS FOR PHYTATE ANALYSIS 115 sodium phytate or acid-soluble barium phytate (Wheeler and Ferrel23) to samples for measurement of recoveries is totally inadequate. Graf and Dintzis28 used 2 hours. With the introduction of HPLC to the analysis of phytate.1 Notes: All extractions were for 3 h with gentle. The best extraction time remains to be determined.2 5783. Dependence on wet chemistry methods and improved technology has provided alternatives that have served well.1 were extracted with 1:10 sample-to-solvent ratios. Samples in Table 3.7_fm Page 116 Sunday. This was shown by Heubner and Stadler18 when the precipitation of white ferric phytate in their titration system made the observation of the pink titration endpoint difficult and thus compromised the accuracy and sensitivity of that original method. it is nonspecific because it detects only a change in the refractive index. This observation was repeated subsequently by Rather. Purification Methods With consideration of the multitude of possible food matrices. Tangendjaja et al. giving a sample-to-solvent ratio of between 10 and 20. it must be used in an isocratic system. unfortunately.27. In the final analysis.005 to 0.2387_ch3. and wet-ashed prior to colorimetric phosphate analysis. Averill and King20 were the first investigators to analyze a large number of foodstuffs.20 McCance and Widdowson. (2) gentle agitation for a period of at least 2 and possibly 3 hours.5 N HCl. McCance and Widdowson30 used a more casual 5–10 g of dried. 2001 6:47 PM 116 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and this was followed quickly by others who tested their method. Modifications of the iron precipitation purification were then developed. extraction with one of the appropriate extraction acids includes several acid-soluble compounds. Earley and DeTurk34 and Earley35 used a sample-to-solvent ratio of 1:20. This . Oberleas27 conducted the ferric phytate precipitation in a 50-mL centrifuge tube.7. 3RD EDITION conditions. washed the precipitate with dilute acid. They used a sample-to-solvent ratio of 1:25 of 2% HCl. First. when developing an analytical method for quantitation of phytate. a solvent concentration that would not provide consistently reliable extractions. The differential refractive index detector was the first successful detector to be applied to HPLC analysis of phytate.66 mol/l HCl or 10% TCA. depending on an estimated phytate content and extracted with 10 mL of extractant.30 and Young. either 0. this detector has several disadvantages. no useful UV or visible spectra are available.38 This caused Earley and DeTurk34 to modify the analytical method to precipitate acid-insoluble ferric phytate in dilute acid and collect the resulting precipitate on asbestos pads in Gooch crucibles which could then be dry-ashed. but. The resulting phosphate was eluted with acid and quantified colorimetrically. it appears that 2 hours is the minimum extraction time that can be recommended and 3 hours may be optimum. it is the least sensitive of the detectors available. Haug and Lantzsch36 varied the sample size from 0.5 or 0.24 Graf and Dintzis28 performed their analyses following extraction with a 1:20 sample-to-solvent ratio of 0. second. Detection Methods It has been acknowledged that because of the nature of the phytate molecule. and third.2 N HCl. This provides sample-to-solvent ratios from 167 to 2000. May 6. and (3) a sample-to-solvent ratio of at least 1:10 (w/v).25 used a 1:10 ratio with 3% TCA.19 Averill and King. HPLC has the potential to become the panacea for phytate and inositol phosphate ester analysis. finely ground material with 100 mL 0. the following considerations appear to be important for extraction of the samples: (1) an adequate concentration of an appropriate acid.32 until high-performance liquid chromatography (HPLC) entered the analytical scene.06 g of sample.25 However. they were extracting with 0. The standard appeared to be 20 for several years. Each of these compounds will react according to its chemical nature within the reaction system. Sample-To-Solvent Extraction Ratio Another factor to be considered is the ratio of the sample weight to the volume of the appropriate solvent. though some methods were nonspecific and most were laborious. both qualitative and quantitative.5 N HCl. 2387_ch3.7_fm Page 117 Sunday, May 6, 2001 6:47 PM NEWER METHODS FOR PHYTATE ANALYSIS 117 could be caused by any substance not present in the mobile phase. Using a reverse phase, C-18 column, most phytate peaks elute with the void volume in 2 to 4 min, depending on the flow rate.25,28,33,37 Lee and Abendroth38 were concerned by the lack of retention and thus added tetrabutylammonium hydroxide as an ion pair. In this system the void volume was 1.3 min, whereas phytate elution could be varied from 4.75 min to 11.3 min. The higher the concentration of the ion pair, the shorter was the elution time. The detection limit was reported at 2 ng and linearity was reported up to 40 µg of phytate. Other HPLC methodology variations include the isocratic anion exchange columns with post column derivatization.39,40 These methods were non-specific for phytate. A unique gradient HPLC method was reported by Rounds and Nielsen.41 This method provides both a specific separation and quantitative detection to about 3–5 nmol. This can be adapted to provide both qualitative and quantitative detection of phytate and other inositol phosphates. It is being described in greater detail because the described modification of this method is being pursued for submission to AOAC International for consideration as an “Official Method.” FERRIC IRON-TO-SULFOSALICYLIC ACID HPLC PHYTATE ANALYSIS METHODS Equipment The following equipment is necessary when using this method for the analysis of phytate. Brand names are not used because several brands are available that are equally effective. A typical layout of the instrumentation is shown in Figure 3.7.1. 1. A pumping unit capable of creating a gradient. 2. A UV/Vis detector capable of monitoring specific wavelengths. In this method that wavelength is 500 nm (Figure 3.7.2). This detector may be connected to a recorder, a digital printout or single computer with appropriate software, or a computer network with appropriate analytical software. 3. An autosampler is not essential but is a convenient component to the system. The autosampler usually may be programmed for injection volume and the multiple number of injections per sample. The system may be programmed for a large number of samples, allowing the analyses to be performed over an extended period of time without close monitoring. 4. Some means of recording the output for analysis is essential. This may be as simple as an analog recorder. There is computer software capable of digitizing the output data, analyzing these data, and printing the results. An analog/digital converter between the detector and the computer network allows the data to be transmitted to a centralized network server unit. 5. A secondary pump is necessary for pumping Wade’s reagent, which is used for post column reaction with phytate and inositol phosphates. Several accessory items are needed to complete the HPLC system for the analyses of phytate. 1. Inline check valve (two required). These are absolutely necessary because of the pressure differentials that may be generated between the gradient pump and the reagent pump. These check valves require very small (i.e., 1.5 psi) breaking pressure and must sustain a sizeable (i.e., 5000 psi) reverse pressure. This is necessary so that at any time, gradient reagents do not back up into the Wade’s reagent and vice versa. 2. A mixing tee, mixing cross, or similar device is needed for the initial mixing of all reagents combined to complete the analytical reaction. 3. A sample loop, mixing loop, web, etc. of the same dimensions (0.020" [0.5 mm] ID × 1/16" OD by 200 cm) as the operating tubing. This mixing loop is used to obtain thorough mixing of the post-column reactants. 4. PEEK or similar tubing, 0.020" [0.5 mm] ID × 1/16" OD, was adequate for connecting all of the necessary components of the system. 2387_ch3.7_fm Page 118 Sunday, May 6, 2001 6:47 PM 118 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Figure 3.7.1 A typical schematic layout of the equipment needed for HPLC analysis for phytate. 5. PEEK or similar fittings and ferrules with appropriate thread pattern to connect the tubing with the HPLC system. 6. An inline filter with 0.45-µm filter frits that can be easily removed and the frits exchanged as frequently as needed. The frits should be easy to replace and may be recycled by sonication with detergent or, when necessary, 30% HNO3, then rinsed by sonication with water. Clogged filter frits may create considerable noise in the analytical curves. 7. A pulse damper is necessary to smooth the flow following passage through the inline filter. This should be a low-volume damper. 8. A PL SAX (50 × 4.6 mm, particle size 8 µm, 100 nm pore type) or equivalent anion exchange column. This column is suitable for 20 µL injections of sample. A smaller column, 8 µm, 1000 Å, stainless steel, 50 × 2.1 mm, is available and may be successful with some HPLC systems. With this smaller column, 5 µL samples may be used. 2387_ch3.7_fm Page 119 Sunday, May 6, 2001 6:47 PM NEWER METHODS FOR PHYTATE ANALYSIS 119 Peak: IP6 200 Peak: IP6 150 150 100 100 50 50 400 mAU mAU 200 450 500 nm 550 600 Figure 3.7.2 A visible spectrum of the ferric sulfosalicylate complex for the analysis for phytate. Reagents 1. N-methyl piperazine (FW100.17) (corrosive, toxic, hygroscopic, combustible), pK about 4.3. 2. Sodium nitrate (FW 84.99) (strong oxidizer, toxic irritant, hygroscopic) or sodium chloride (FW 58.44) (irritation, systemic toxicity), reagent grade. 3. Ferric chloride (FW 270.30) (harmful solid, corrosive, hygroscopic), reagent grade. 4. 5-sulfosalicylic acid (FW 239.25) (corrosive), reagent grade. 5. Trichloroacetic acid, reagent grade or hydrochloric acid, reagent grade. 6. Sodium phytate (hygroscopic), 99% purified, used for standards. Obtain a Certificate of Analysis or standardize as described below. Solutions Two systems have been tested and work equally satisfactorily. One system utilizes N-methyl piperazine·HNO3 buffer with sodium nitrate as the gradient; the second system utilizes the N-methyl piperazine buffer with sodium chloride as the gradient. As a displacement anion, either nitrate or chloride ions have similar properties.42 1. 0.01 mol/L methyl piperazine·HNO3 buffer (pH 4.0). Weigh 1.0 g/L of N-methyl piperazine into a small beaker. Wash into an appropriate graduate cylinder containing about 500 mL of HPLC-grade water (milliQ water is satisfactory). Fill to volume with HPLC-grade water. Stir for about 5 min with a magnetic stirrer. The initial pH of this buffer is about 10. This should be adjusted to pH 4 with either 1 mol/L nitric acid or 1 mol/L hydrochloric acid, depending upon which system is being utilized. Filter through 0.45-µm nylon filter, degas with helium at 25 mL/min or a vacuum degasser if available with the HPLC system. Store in a stoppered clear glass bottle at room temperature. This is a stable reagent. 2. 0.5 mol/L sodium nitrate in 0.01 mol/L methyl piperazine·HNO3 buffer (pH 4.0). Weigh 42.5 g sodium nitrate, place into a 1-L graduate cylinder, make to volume with buffer. Alternatively, weigh the sodium nitrate and 1 g of N-methyl-piperazine and make to volume. Adjust to pH 4.0 with 1 mol/L nitric acid, filter through 0.45 µm nylon filter, degas as above. (With some HPLC systems it may be necessary to increase the sodium nitrate concentration to 0.6 mol/L or 51.0 g/L to effect separations within the operating framework.) The pK of N-methyl-piperazine is about 4.2–4.3. These solutions are colorless and stable. 3. Alternate: 0.7 mol/L sodium chloride in 0.01 mol/L methyl piperazine·HCl buffer (pH 4.0). Weigh 40.9 g sodium chloride, place into a 1-L graduate cylinder, make to volume with buffer adjusted with 1 mol/L HCl. Alternatively, weigh the sodium chloride and 1 g of N-methyl-piperazine and make to volume. Adjust to pH 4.0 with 1 mol/L hydrochloric acid, filter through 0.45 µm nylon filter, degas as above. The buffered solutions are prepared as above and adjusted to pH 4 with 2387_ch3.7_fm Page 120 Sunday, May 6, 2001 6:47 PM 120 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION 1 mol/L HCl. The initial pH for the piperazine solutions will be 9 to 10.5. This solution is colorless and stable when stored in a stoppered glass bottle at room temperature. 4. Wade’s reagent. Weigh 0.2 g (0.74 mmol) ferric chloride·6H2O. Weigh 1.5 g (6.87 mmol) 5-sulfosalicylic acid (SSA). Combine into a 1-L graduate cylinder, make to volume with HPLCgrade water, adjust to pH 1.8 with 1 mol/L HNO3 or HCl as appropriate, filter through 0.45-µm nylon filter, store in brown glass bottle. This is a light-sensitive reagent but is stable for several months in a stoppered brown glass bottle at room temperature. This combination provides a 9.2:1 molar ratio of SSA to Fe in the reagent solution. Wade’s reagent, ferric chloride and sulfosalicylic acid, initially should have a pH of 2.5–3.0. This solution will be reddish-purple in color; that is the reaction between ferric iron and sulfosalicylic acid. It has an absorbance maximum of 500 nm. Sulfosalicylic acid appears to have a pK of about 2–2.2 that provides some buffering in that pH range. Extracting Solutions As noted earlier, three extracting solutions are equally effective in extracting phytate, phosphate, and inositol phosphates from several food samples. 1. Trichloroacetic acid (10%): Weigh 100 g of TCA, make to 1 L with HPLC grade water. Filter through 0.45-µm nylon filter, store in a stoppered clean glass bottle. This reagent is stable for an extended period of time when stored at room temperature. 2. 0.5 M HCl: Measure 41.8 mL of concentrated HCl and make to 1 L. Filter through 0.45-µm nylon filter and store in a stoppered clean glass bottle. This is a very stable reagent for an extended period of time when stored at room temperature. 3. 0.66 M HCl: Measure 54 mL of concentrated HCl and make to 1 L. Filter through 0.45-µm nylon filter, store in a stoppered clean glass bottle. This is a very stable reagent for an extended period of time when stored at room temperature. Preparation of Standards Label information of sodium phytate as purchased is not reliable for standard preparation. A “Certificate of Analysis” based on the lot number may be available from the supplier which contains more detailed and more accurate information on each lot of sodium phytate. This may be available for download via the Internet using the lot number as the reference or may be requested from the supplier. As with many techniques, the results of analyses for phytate are determined by the care in preparation and accuracy of the standards. Dodecasodium phytate (FW 923.8) as purchased is adequate for standards with some precautions. The molecular weight of undissociated phytic acid (MW 660) has classically been used in the calculation of phytate equivalent from phosphate content of purified, ashed sodium phytate. There has been no absolute agreement that the phytate should be expressed on the basis of the undissociated phytic acid equivalent or MW 648, which is the molecular weight of the totally dissociated anion that is the active species. It makes little difference as long as everyone is consistent in this usage. Tradition has been to use MW 660 in all calculations where the specific species is not apparent. The analysis varies slightly from lot to lot and thus must be recalculated from the “Certificate of Analysis,” which provides precise information for each lot, or the phytate must be restandardized within the laboratory. Phosphorus or phosphate is one entity that can be reanalyzed by most laboratories to update the sodium phytate for use as a standard. Thus, procedures for restandardization of phytate are included later in this chapter. Calcium is variable for different lots and must be corrected individually. Water is a critical variable that may change with different climates and, in some cases, with different seasons of the year. Sodium phytate has a tendency to be hygroscopic and therefore must be stored carefully to prevent water accumulation. This can always be corrected if needed by phosphate analysis. Therefore, the weight of sodium phytate as purchased made to 2387_ch3.7_fm Page 121 Sunday, May 6, 2001 6:47 PM NEWER METHODS FOR PHYTATE ANALYSIS 121 100 mL with HPLC grade water to provide 1 g of phytic acid/100 mL is not constant and must be determined by each laboratory for the lot of sodium phytate utilized. To calculate the weight of dodecasodium phytate as purchased for your standard, utilize the percent phosphorus (g/100 g) from the “Certificate of Analysis.” Divide this by 0.282 (fractional portion of phosphorus in phytic acid; MW 660) or the percentage of phytic acid based on phosphorus analysis. To obtain 1 g phytate per 100 mL, divide 1 g by the fractional portion of phytic acid to obtain the grams of that lot necessary to obtain that 1 g, and dilute to 100 mL with 0.5 N HCl to correct for acid effects. This would be equivalent to 200 µg (303 nmol) phytate (MW 660)/20 µL injection and is the stock standard. Working standards are prepared by dilution of the stock standard, i.e., 1:4 = 50 µg (75.7 nmol); 1:8–25 µg (37.9 nmol); 1:10 = 20 µg (30 nmol); 1:25 = 8 µg (12.1 nmol); and 1:50 = 4 µg (6 nmol) per 20 µL injection. For 5-µL injection, dilutions of stock standard would be 1:2 = 25 µg (37.85 nmol); 1:4 = 12.5 µg (18.925 nmol); 1:10 = 5.0 µg (7.5 nmol); 1:25 = 2.0 µg (3.025 nmol); and 1:50 = 1 µg (1.5 nmol). The 1:50 may be below the detection limit for some instruments. Procedure The prepared solutions are placed in their appropriate locations. The N-methyl piperazine may also be used as the wash solution for washing the injection needle of the autosampler. The piperazine buffer and gradient solutions are pumped with the gradient pump; Wade’s reagent is pumped with the auxiliary pump. The piperazine buffer and Wade’s reagent are pumped until stability is reflected on the detector and system pressures are stabilized. The instrument operates at relatively low pressures, but these pressures may be different for each analytical HPLC system. Several parameters should be monitored, i.e., pressures of both pumps, pH of the final eluent, flow rate of each pump, and total flow rate. The principle of the method is to separate inorganic phosphate and the various inositol phosphates in the sample by eluting them from the column with the gradient of NaNO3 or NaCl. The buffer or buffered gradient mixed with the Wade’s reagent forms a stable baseline. When phosphate or inositol phosphates are eluted from the column, they form a more stable complex with ferric iron than does sulfosalicylic acid. Thus, they displace ferric iron from the colored complex, reducing the color produced by the complex. Ferric phytate is white and thus decreases the color produced by the ferric sulfosalicylate reaction. The decreased color is recorded as a decreased absorbance and is proportional to the concentration of phytate or inositol phosphates in the eluting sample. Negative peaks are an inherent part of this procedure but can be made positive by reversing the polarity of the recording instrument or inverting the peaks within the software of most instrument outputs. The mechanism for accomplishing this is otherwise individual for each of the software systems available. Instrumentation Conditions Equilibrate the system with 1 mL/min N-methyl piperazine buffer and 0.5 mL/min Wade’s reagent until the absorbance changes only minimally. The following are typical operating conditions, but these may vary slightly for different systems: A. Column — PL-SAX or equivalent strong anion exchange column. B. Column temperature — ambient. C. Gradient — 100% N-methyl piperazine buffer at baseline or zero time; 100% buffered salt (nitrate or chloride) at the end of the run (26 or 30 minutes). D. Injector temperature — ambient. E. Detector — Visible region, 500 nm. F. Injection volume — 5 or 20 µL, depending on column diameter. 2387_ch3.7_fm Page 122 Sunday, May 6, 2001 6:47 PM 122 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION G. Total run time — 31 or 36 min, which sometimes varies with different HPLC systems and buffered salt concentrations. H. Gradient time — 26 or 30 min, again dependent upon different HPLC systems and choice of buffered salt concentrations. I. Flow rate — 1 mL/min for the baseline buffer and gradient buffer salt solutions. 0.5 mL/min Wade’s reagent. J. Pressures — 200–400 pounds per square inch (psi) or equivalent should be appropriate for starting or baseline pressures. When pressures increase above 1000–1200 psi, this should be an indication of a need to change filter frits or locate a precipitate plug in the system. Cautions are discussed below. Extraction of Samples Three extraction solutions were shown to be comparable for extracting samples. When tested on identical samples, 10% trichloroacetic acid, 0.5 mol/L HCl, or 0.66 mol/L HCl gave results that were not different statistically. Neither 3% TCA nor 5% TCA provided comparable results when used as extracting solutions. Recovery studies that add very water-soluble sodium phytate to a sample and consider the recovery of this as a measure of completeness of recovery from the matrix are not adequate as a measure of completeness of recovery from the matrix. Food matrices are too complex for this criterion to be meaningful. Weigh approximately 1 g of sample (1.0000–1.2000 g is a good guide) and place 10 mL of one of the extracting solutions on the sample. Stir vigorously to be certain the sample is wet throughout. Place the samples on the appropriate device for gentle agitation. It is important that the samples be extracted by gentle agitation over a large surface area. This may be accomplished in several ways. A 50-mL polyethylene or polypropylene centrifuge tube with a screw cap may be satisfactory. It may be necessary to mix the tubes on a vortex before shaking and at about the midpoint, to assure that all of the sample is wet and to incorporate that which is clinging to the side of the tube into the extraction process. These tubes laid on their sides and taped to an orbital shaker or other mild shaking device should serve satisfactorily. A 125-mL Erlenmeyer flask will serve well, either on an orbital shaking platform, a dry Dubnoff shaking incubator, a radial arm shaker at low speed, or some other device that will roll or shake gently over the large surface at the bottom of the flask. Samples should be shaken for a minimum of 2 h but preferably 3 or more hours. Depending on circumstances, samples may be shaken overnight. After extraction, the extract must be removed from the matrix. This can be accomplished in several ways. If the centrifuge tubes are used for extraction, they may be centrifuged using a clinical centrifuge at about 3500 rpm (~1800 g) for 20 minutes (a high-speed centrifuge is not necessary). At this point several secondary measures may be taken. First, the extracting solution may be removed from the top of the centrifuge tube with a Pasteur pipet, placed in a micro-centrifuge tube, and centrifuged a second time using a higher-speed centrifuge for 10 min. Transfer the supernatant with a Pasteur pipet directly to a HPLC vial. Second, the extractant from the first centrifugation may be placed into a 3-mL polyethylene syringe and forced through a 0.45-µm nylon filter directly into a HPLC vial. Third, if extraction was done in an Erlenmeyer flask, the sample may be filtered with gravity or vacuum through Whatman #1, 2, or equivalent filter paper. Then place some filtrate into a micro-centrifuge tube as above or force it through a 0.45-µm filter. Precautions Ferric phytate is insoluble in dilute acid and forms a precipitate in the system. This is effectively removed by the inline filter frit. The frits must be exchanged or cleaned frequently; otherwise, pressures may increase rapidly. These frits can be readily cleaned in one of several ways. The filter can be cleaned by pumping 0.2 mol/L sodium hydroxide through the reagent side only, then flushing 2387_ch3.7_fm Page 123 Sunday, May 6, 2001 6:47 PM NEWER METHODS FOR PHYTATE ANALYSIS 123 thoroughly with water. The filter frits may be removed and replaced with clean filter frits. The plugged filter frits then can be collected and sonicated with dilute, mild detergent or, if necessary, with 30% nitric acid, rinsed by sonication with water, and recycled. Sometimes more than one procedure is needed. When the phytate reaction occurs, the ideal window of reaction should occur between pH 1.5 and 2.5. For this reaction to be most stable, all solutions are adjusted to specific pH’s of 4 for the buffered solutions and 1.8 for the Wade’s reagent. This provides a final pH of 2.1–2.2 at most times. This is not the pH of maximum precipitation for ferric phytate, which is about 1, but provides a stable environment for the reaction to occur. RESTANDARDIZATION OF SODIUM PHYTATE Though sodium phytate as purchased is quite stable and free of most contaminating phosphate compounds, there may be occasions that necessitate the analysis and standardization for use as a standard, i.e., inability to obtain a “Certificate of Analysis,” long-term storage, etc. It will be necessary to be able to analyze the inorganic phosphate in the presence of and without hydrolyzing any phytate. Total phosphate following an ashing procedure is also needed. This phosphate procedure is a modification of the Fiske–Subbarow43 method. It has a long history, has stable reagents, and can be read from 640 to 720 nm with equal sensitivity. Reagents 1. Sulfuric (sulphuric) acid (5 N). Dilute 70 mL of concentrated sulfuric acid to 500 mL. Stable reagent. 2. Ammonium molybdate reagent. Dissolve 12.5 g ammonium molybdate in 200 mL water. Transfer to a 500-mL volumetric flask, add 100 mL 5 N sulfuric acid, and dilute to volume with water. This is a stable reagent. 3. Sulfonic acid reagent (1-amino-2-naphthol-4-sulfonic acid). Dissolve 0.16 g 1-amino-2-naphthol4-sulfonic acid, 1.92 g sodium sulfite, and 9.6 g sodium bisulfite in about 90 mL water. Heat to dissolve if necessary. Quantitatively transfer to 100-mL volumetric flask. Make to volume with water. Store in brown bottle or cover flask with brown paper or aluminum foil and refrigerate. Stable for at least 2 weeks. Standard Phosphate Solution Stock phosphate solution. Prepare a standard solution by weighing 0.1757 g of dried, dessicated, potassium dihydrogen phosphate (primary standard) per liter in a volumetric flask. Add 20 mL 5 N sulfuric acid and make to volume with water. This solution contains 40 mg P (as phosphate) per liter (40 µg/mL). Dilute as necessary to obtain other standard concentrations. Procedure This can be done in 25- or 50-mL volumetric flasks, provided the standard curve is determined in the same-sized flasks. Pipette blank (water), standard, or sample (in duplicate or triplicate) into an appropriate set of flasks. Add distilled water to about 10 or 20 mL, depending on flask size. Mix thoroughly. Add 2 mL molybdate reagent; mix well. Add 1 mL sulfonic acid reagent, dilute to volume, and mix well. Wait 15 min and read at 640 nm. The peak for this reaction is about 910 nm, but the slope is so gradual as to provide stability at 640 nm, which is within the capability of all spectrophotometers. 2387_ch3.7_fm Page 124 Sunday, May 6, 2001 6:47 PM 124 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Total Phosphate After measuring the inorganic phosphate, it is necessary to hydrolyze the phytate and measure the total phosphate. There are several important considerations necessary when wet-ashing samples for phosphate analysis. Phosphoric acid has several forms which boil at fairly low temperatures. According to the Handbook of Chemistry and Physics, hypophosphoric acid decomposes at 100°C, metaphosphoric acid sublimates, and orthophosphoric acid decomposes at temperatures less than 213°C. Various forms of phosphorus acids decompose or boil below 200°C. The importance of all this is that it is necessary to hydrolyze the phytate samples completely, the wet-ash must be at as low a temperature as possible, and the heating time should be the minimum required. Otherwise, variability may be encountered in phosphate content by any or all of the above. Procedure Pipette a measured aliquot of phytate solution to be standardized into a microkjeldahl flask or 25 × 200 mm digestion tubes. Add 0.5 mL concentrated sulfuric acid to each tube. Add 2 mL of 30% hydrogen peroxide to each tube. Add two boiling beads to each tube. Heat on a microkjeldahl digestion rack or block heater until only the sulfuric acid remains. If the digestion solution is clear, digestion is complete; in the event of any color or turbidity, cool to touch, add 1 mL 30% hydrogen peroxide, and continue heating as before. Continue the process until a clear solution results. Quantitatively transfer samples to appropriate volumetric flasks and continue color development and measurement as above. Calculations The calculation of phytate concentrations from the analyses begins with the understanding of the formula from the Beer–Lambert Law as follows: Concentration Standard Concentration Sample --------------------------------------------------------------------- = ------------------------------------------------------------------Absorbance (Response) Standard Absorbance (Response) Sample Sum [Concentration Standard /Absorbance (Response) Standard ]/n Standards = Mean K Mean K × R × W × M = mg phytate/g sample where Mean K = mean of the concentrations (nmol)/response (area) for standards R5 or 20 mL = Response Sample W5 mL = 2000 (20 × 10 × 10 mL extractant/sample weight (g)) [corrects for volume and weight of sample] (5 mL × 20 = 0.1 mL) (0.1 mL × 10 = 1.0 mL) (1.0 mL × 10 mL extractant corrects for volume of extractant) W20 mL = 500 (5 × 10 × 10 mL extractant)/sample weight (g) [corrects for volume and weight of sample] (20 mL × 5 = 0.1 mL), etc. M5 or 20 mL = 660 (MW)/106 [Corrects for nmol/g to mg/g] multiply mg/g phytate by 1000 = mg phytate/kg sample multiply mg/g phytate by 100/1000 = % phytate in sample Molar Ratios When calculating this ratio, it is important that the initial units of both components be the same (i.e., mg/kg, mg/100 g, g/kg, etc.): 2387_ch3.7_fm Page 125 Sunday, May 6, 2001 6:47 PM NEWER METHODS FOR PHYTATE ANALYSIS 125 mg/kg phytate ---------------------------------660 (MW) ---------------------------------- = Phytate:Zinc molar ratio mg/kg zinc ----------------------------65.4 (At wt) When calculating [Ca][Phytate]/Zn molar ratio, which is more precise, it is necessary to convert all components to grams per kilogram and thus the appropriate definition of molar. g/kg phytate g/kg calcium ----------------------------- × ------------------------------660 (MW) 40.1 (At wt) ----------------------------------------------------------------------- = [ Calcium ] [ Phytate ]:[Zinc] molar ratio g/kg zinc ----------------------------65.4 (At wt) ACKNOWLEDGMENT The HPLC method described above was developed in the laboratories and with the assistance and cooperation of Gerber Products Company, Fremont, Michigan, and its employees. This assistance is greatly appreciated. REFERENCES 1. Oberleas, D., The determination of phytate and inositol phosphates, in Methods of Biochemical Analysis, vol. 20, Glick, D., Ed., Wiley, New York, 1971, 87. 2. Cosgrove, D. J., Inositol Phosphates, Elsevier, Amsterdam, 1980, 12. 3. Cheryan, M., Phytic acid interactions in food systems, CRC Crit. Revs. Food Sci. Nutr., 1980, 297. 4. Maga, J. A., Phytate: its chemistry, occurrence, food interactions, nutritional significance, and methods of analysis, J. Agri. Food Chem., 30, 1, 1982. 5. Oberleas, D., Phytate content in cereals and legumes and methods of determination, Cereal Foods World, 28, 352, 1983. 6. Oberleas, D. and Harland, B. F., Analytical methods for phytate, in Phytic Acid: Chemistry and Applications, Graf, E., Ed., Pilatus Press, Minneapolis, 1986, 77. 7. Wade, H. E. and Morgan, D. M., Fractionation of phosphates by paper ionophoresis and chromatography, Biochem. J., 60, 264, 1955. 8. Sobolev, A. M. and Vyskrebentseva, E. I., The problem of identification of organic acid-soluble phosphorus compounds in plants by means of regulated paper chromatography, Fiziol. Rastenii, 6, 257, 1958. 9. Smith, D. H. and Clark, F. E., Chromatographic separations of inositol phosphorus compounds, Soil Sci. Soc. Proc., 1952, 170. 10. Tomlinson, R. V. and Ballou, C. E., Myoinositol polyphosphate intermediates in the dephosphorylation of phytic acid by phytase, Biochem., 1, 166, 1962. 11. Cosgrove, D. J., The isolation of myoinositol pentaphosphates from hydrolysates of phytic acid, Biochem. J., 89, 172, 1963. 12. Cosgrove, D. J., The chemistry and biochemistry of inositol polyphosphates, Rev. Pure Appl. Chem., 16, 209, 1966. 13. Cosgrove, D. J., Ion-exchange chromatography of inositol polyphosphates, Ann. N.Y. Acad. Sci., 165, 677, 1969. 14. Cosgrove, D. J., Chemical nature of soil organic phosphorus. II. Characterization of the supposed DL chiro-inositol hexaphosphate component of soil phytate as D chiro-inositol hexaphosphate, Soil Biol. Biochem., 1, 325, 1969. 15. Irving, G. C. J. and Cosgrove, D. J., Inositol phosphate phosphatases of microbiological origin. Inositol pentaphosphate products of Aspergillus ficuum phytases, J. Bacteriol., 112, 434, 1972. 2387_ch3.7_fm Page 126 Sunday, May 6, 2001 6:47 PM 126 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION 16. Cosgrove, D. J., Inositol phosphate phosphatases of microbiological origin. Inositol phosphate intermediates in the dephosphorylation of the hexaphosphates of myo-inositol, scyllo-inositol and D-chiroinositol by a bacterial species (Pseudomonas sp.) phytase, Aust. J. Biol. Sci., 23, 1207, 1970. 17. Cosgrove, D. J., Phosphorylation of epi-inositol and muco-inositol with phosphoric acid, Carbohydrate Res., 40, 380, 1975. 18. Heubner, W. and Stadler, H., Uber eine Titration-methode zur Bestimmung des Phytins, Biochem. Z., 64, 422, 1914. 19. Rather, J. B., Determination of phytin phosphorus in plant products, J. Am. Chem. Soc., 39, 2506, 1917. 20. Averill, H. P. and King, C. G., The phytin content of foodstuffs, J. Am. Chem. Soc., 48, 724, 1926. 21. Latta, M. and Eskin, M., A simple and rapid colorimetric method for phytate determination, J. Agri. Food Chem., 28, 1313, 1980. 22. de Lange, D. J., Joubert, C. P., and du Peez, S. F. M., The determination of phytic acid and factors which influence its hydrolysis in bread, Proc. Nutr. Soc. S. Africa, 2, 69, 1961. 23. Wheeler, E. L. and Ferrel, R. E., A method for phytic acid determination in wheat and wheat fractions, Cereal Chem., 48, 312, 1971. 24. Cilliers, J. L. and van Niekerk, P. J., LC determination of phytic acid in food by postcolumn colorimetric detection, J. Agri. Food Chem., 34, 680, 1986. 25. Tangendjaja, B., Buckle, K. A., and Wootton, M., Analysis of phytic acid by high-performance liquid chromatography, J. Chromat., 197, 274, 1980. 26. Bos, K. D., Verbeek, C., van Eeden, C. H. P., Slump, P., and Wolters, M. G. E., Improved determination of phytate by ion-exchange chromatography, J. Agri. Food Chem., 39, 1770, 1991. 27. Oberleas, D., Dietary Factors Affecting Zinc Availability, Ph.D. dissertation, University of MissouriColumbia, 1964. 28. Graf, E. and Dintzis, F. R., High-performance liquid chromatographic method for the determination of phytate, Anal. Biochem., 119, 413, 1982. 29. Samotus, B. and Schwimmer, S., Phytic acid as a phosphorus reservoir in the developing potato tuber, Nature, 194, 578, 1962. 30. McCance, R. A. and Widdowson, E. M., Phytin in human nutrition. Biochem. J., 29, 2694, 1935. 31. Young, L., The determination of phytic acid, Biochem. J., 30, 252, 1936. 32. Harland, B. F. and Oberleas, D., A modified method for phytate analysis using an ion-exchange procedure: application to textured vegetable proteins, Cereal Chem., 54, 827, 1977. 33. Camire, A. L. and Clydesdale, F. M., Analysis of phytic acid in foods by HPLC, J. Food Sci., 47, 575, 1982. 34. Earley, E. B. and DeTurk, E. E., Time and rate of synthesis of phytin in corn grain during reproductive period, J. Am. Soc. Agron., 36, 803, 1944. 35. Earley, E. B., Determining phytin phosphorus: stoichiometric relation of iron and phosphorus in ferric phytate, Anal. Chem., 16, 389, 1944. 36. Haug, W. and Lantzsch, H-J., Sensitive method for rapid determination of phytate in cereals and cereal products, J. Sci. Food Agri., 34, 1423, 1983. 37. Graf, E. and Dintzis, F. R., Determination of phytic acid in foods by high-performance liquid chromatography, J. Agri. Food Chem., 30, 1094, 1982. 38. Lee, K. and Abendroth, J. A., High performance liquid chromatographic determination of phytic acid in foods, J. Food Sci., 48, 1344, 1983. 39. Fitchett, A. W. and Woodruff, A., Determination of polyvalent anions by ion chromatography, Liq. Chromat. HPLC, 1, 48, 1983. 40. Phillippy, B. Q. and Johnston, M. R., Determination of phytic acid in foods by ion chromatography with post-column derivatization, J. Food Sci., 50, 541, 1985. 41. Rounds, M. A. and Nielsen, S. S., Anion-exchange method for determination of phytate in foods, J. Chromat. A, 653, 148, 1993. 42. Snyder, L. R. and Kirkland, J. J., Introduction to Modern Liquid Chromatography, 2nd ed., John Wiley, New York, 1979, 410. 43. Fiske, C. H. and Subbarow,Y., The colorimetric determination of phosphorus, J. Biol. Chem., 66, 375, 1925. 2387_ch3.8_fm Page 127 Sunday, May 6, 2001 6:49 PM CHAPTER 3.8 Determination of the Saponin Content of Foods David Oakenfull and John D. Potter INTRODUCTION There is currently much interest in the nutritional properties of “phytochemicals” — plant materials present at relatively low levels in foods but with significant biological activity.1 Saponins are one such class of phytochemicals which are present in many fiber-rich foods, particularly legumes. Their presence is relevant to human nutrition because they have been shown to increase fecal excretion of bile acids in human2,3 and animal studies.4–6 In particular, yucca saponins2 and saponin-rich alfalfa seeds7 have been shown to lower plasma cholesterol in humans. Saponins are a structurally diverse group of triterpene or steroid glycosides.8,9 The molecules are amphiphilic, the triterpene or steroid part being hydrophobic and the sugar part hydrophilic. This gives saponins their characteristic surface activity, from which the name is derived. The structure of a typical saponin, one of those present in soybeans, is shown in Figure 3.8.1. Saponins have been identified in many hundreds of plant species, but only a few of these are used as food by humans (see Appendix, Table 9). There appear to be two mechanisms by which saponins can affect cholesterol metabolism: 1. Some saponins, with particularly defined structural characteristics, form insoluble complexes with cholesterol (e.g., the well-known precipitation of cholesterol by digitonin). When this complexation process occurs in the gut, it inhibits the intestinal absorption of both endogenous and exogenous cholesterol.10 2. Saponins can interfere with the enterohepatic circulation of bile acids by forming mixed micelles. These can have molecular weights of several millions,11 and the reabsorption of bile acids from the terminal ileum is effectively blocked.12 EXTRACTION PROCEDURE The dried plant material is first extracted with acetone, chloroform, or hexane, preferably using a Soxhlet extractor, to remove lipids, pigments, etc. The residue is then further extracted with methanol, which removes the saponins, along with many other compounds, such as simple sugars, oligosaccharides, and flavonoids. 0-8493-2387-8/01/$0.00+$1.50 © 2001 by CRC Press LLC 127 2387_ch3.8_fm Page 128 Sunday, May 6, 2001 6:49 PM 128 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION H3C CH3 OH CH3 CH3 CH3 CH3 O H3C CH2OH CO2H O HO OH H H2COH O HO OH HO CH 3 H O H HO OH Figure 3.8.1 Structure of one of the saponins from soybeans. METHODS OF ANALYSIS Saponins have proved very difficult to assay, and many of the results in the literature are, at best, only approximations. For this reason, the majority of the values given in Table 3.8.1 of the Appendix are quoted as a range — there are often major discrepancies between results from different techniques and different laboratories. The difficulty is compounded by the fact that saponin content 2387_ch3.8_fm Page 129 Sunday, May 6, 2001 6:49 PM DETERMINATION OF THE SAPONIN CONTENT OF FOODS 129 can vary considerably between different cultivars of the same plant species and can also depend on growing conditions. The major methods of analysis are thin-layer chromatography (TLC),13,14 high-performance liquid chromatography (HPLC),15,16 and gas chromatography with mass spectroscopy to identify the saponin (trimethylsilylated sapogenol) peaks (GC-MS).17 TLC The essence of the technique is to spot a TLC plate with a crude saponin extract, develop the plate with a suitable solvent system, and use one of a number of methods for estimating the quantity of saponin on the plate. Suitable solvent systems are n-butanol–ethanol–concentrated ammonia (7:2:5) or chloroform–methanol–water (13:7:2). Spots can be visualized by spraying with 10% H2SO4 in ethanol or p-anisaldehyde–H2SO4–glacial acetic acid (1:2:100) and estimated by densitometry. The method is open to criticism in that spots may be wrongly identified as saponins or may be overlapped by other compounds (such as oligosaccharides). (However, techniques such as atomic bombardment mass spectrometry have been used to unequivocally identify the spots.18) HPLC This technique has been used for the separation and analysis of both aglycones (the steroid or triterpene part of the molecule) and intact saponins. A rapid method for soybean saponins has been described16 in which defatted soy flour is boiled under reflux with 1.5 M H2SO4 in dioxane–water (1:3). This hydrolyzes the saponins, and the sapogenins can then be extracted with ethyl acetate. HPLC is carried out with a commercial silica column, eluting with light petroleum–ethanol with a gradient technique. The saponin content can be estimated by assuming that the carbohydrate-tosapogenin ratio is 1:1 (by weight). Table 3.8.1 Plant Foods that Contain Significant Levels of Saponins and Their Estimated Saponin Content David Oakenfull and John D. Potter Plant Saponin Content (g/kg dry weight) Alfalfa sprouts (Medicago sativa) Asparagus (Asparagus officinalis) Broad bean (Vicia faba) Chickpea (Cicer arietinum) Green pea (Pisum sativum) Kidney bean (Phaseolus vulgaris) Lentil (Lens culinaris) Mung bean (Phaseolus mungo) Navy bean (Phaseolus vulgaris) Oats (Avena sativa) Peanut (Arachis hypogaea) Quinoa (Chenopodium quinoa) Sesame seed (Sesamum indicum) Silver beet (Beta vulgaris) Soy bean (Glycine max) Spinach (Spinacea oleracia) Sweet lupin (Lupinus augustifolius) 80 15 3.5 0.7–60 1.8–11 2–16 0.7–1.1 0.5–6 4.5–21 0.2–0.5 0.05–16 10–23 3 58 5.6–56 47 0.4–0.7 2387_ch3.8_fm Page 130 Sunday, May 6, 2001 6:49 PM 130 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION GC-MS A method has been described in which the methanol saponin extract is evaporated to dryness and hydrolyzed by refluxing with dry HCl in methanol (5% v/v). The mixture is neutralized with ammonia and the sapogenols extracted with ethyl acetate. The dry sapogenols are derivatized by heating with bis(trimethylsilyl)trifluoroacetamide in pyridine prior to chromatography on a silica column. Quantitation is via an internal standard (cholesteryl n-decylate) and comparison with isolated, pure sapogenols. Peak identities are confirmed by mass spectroscopy.18 REFERENCES 1. Temple, N. J., Antioxidants and disease: more questions than answers, Nutr. Res., 20, 449, 2000. 2. Bingham, R., Harris, D. H., and Laga, T., Yucca plant saponins in the treatment of hypertension and hypercholesterolemia, J. Appl. Nutr., 30, 127, 1978. 3. Potter, J. D., Illman, R. J., Calvert, G. D., Oakenfull, D. G., and Topping, D. L., Soya saponins, plasma lipids, lipoproteins and fecal bile acids: a double-blind cross-over study, Nutr. Rep. Int., 22, 521, 1980. 4. Malinow, M. R., McLaughlin, P., Kohler, G. O., and Livingston, A. L., Prevention of elevated cholesterolemia in monkeys by alfalfa saponins, Steroids, 29, 105, 1977. 5. Oakenfull, D. G., Fenwick, D. E., Topping, D. L., Illman, R. J., and Storer, G. B., Effects of saponins on bile acids and plasma lipids in the rat, Br. J. Nutr., 42, 209, 1979. 6. Topping, D. L., Storer, G. B., Calvert, G. D., Illman, R. J., Oakenfull, D. G., and Weller, R. A., Effect of dietary saponins on fecal bile acids and neutral sterols, plasma lipids and lipoprotein turnover in the pig, Am. J. Clin. Nutr., 33, 783, 1980. 7. Molgaard, J., von Schenk, H., and Olsson, A. G., Alfalfa seeds lower low density lipoprotein cholesterol and apoprotein B concentration in patients with type II hyperlipoproteinemia, Atherosclerosis, 65, 173, 1987. 8. Oakenfull, D. G. and Sidhu, G. S., Could saponins be a useful treatment for hypercholesterolaemia?, Eur. J. Clin. Nutr., 65, 173, 1987. 9. Oakenfull, D. G. and Sidhu, G. S., Saponins, in Toxicants of Plant Origin, vol. 2, Cheeke, P., Ed., CRC Press, Boca Raton, FL, 1989. 10. Gestetner, B., Assa, Y., Henis, Y., Tencer, Y., Royman, M., Birk, Y., and Bondi, A., Interaction of lucerne saponins with sterols, Biochim. Biophys. Acta, 270, 181, 1972. 11. Oakenfull, D., Aggregation of saponins and bile acids in aqueous solution, Aust. J. Chem., 39, 1671, 1986. 12. Oakenfull, D. G. and Sidhu, G. S., A physico-chemical explanation for the effects of dietary saponins on cholesterol and bile salt metabolism, Nutr. Rep. Int., 27, 1253, 1983. 13. Fenwick, D. E. and Oakenfull, D. G., Saponin content of soya beans and some commercial soya products, J. Sci. Food Agric., 32, 273, 1981. 14. Price, K. R., Curl, C. L., and Fenwick, G. R., The saponin content and sapogenol composition of the seed of thirteen varieties of legume, J. Sci. Food Agric., 37, 115, 1986. 15. Kesselmeier, J. and Stack, D., High performance liquid chromatography analysis of steroidal saponins from Avena sativa L., Z. Naturforsch., 36c, 1072, 1981. 16. Ireland, P. A. and Dziedzic, S. Z., High performance liquid chromatography on silica phase with evaporative light scattering detection, J. Chromatogr., 361, 410, 1986. 17. Ng, K. G., Price, K. R., and Fenwick, G. R., A TLC method for the analysis of quinoa (Chenopodium quinoa) saponins, Food Chem., 49, 311, 1994. 18. Ridout, C. L., Price, K. R., DuPont, M. S., Parker, M. L., and Fenwick, G. R., Quinoa saponins — analysis and preliminary investigations into the effects of reduction by processing, J. Sci. Food Agric., 54, 165, 1991. 2387_Section 4_fm Page 131 Sunday. May 6. 2001 6:01 PM SECTION 4 Physiological and Metabolic Effects of Dietary Fiber . May 6. 2001 6:01 PM .2387_Section 4_fm Page 132 Sunday. Gallaher and Barbara O. species studied.00+$1. particularly starch. with incorporation of fiber by substitution. has been studied extensively. The more common method is incorporation by substitution. the first table contains information from studies using purified dietary fibers.32. the influence of digestible carbohydrates. with the second describing studies where fiber-rich sources were utilized. however.9 Dietary fiber.1 Effect of Dietary Fiber on Protein Digestibility and Utilization Daniel D. are likely to consume an equivalent amount of protein compared to control animals. there is considerable interest in the influence on protein quality of dietary components other than protein. called the dilution (or addition) method. With the second method. and the method of fiber incorporation has been included to assist in evaluating the results of each study.37 incorporation of moderate amounts of fiber usually causes an increased food intake. Three pairs of tables are presented. an indigestible carbohydrate. 2001 6:57 PM CHAPTER 4. However. time period evaluated. in which very high levels of fiber have been fed and/or weanling animals used. For each pair. information on diet composition.31. the protein content remains unchanged on a weight basis. with a fiber-diluted diet the protein-to-calorie ratio remains unchanged. the animals have not been able to increase food intake sufficiently to meet their energy needs. With the substitution method. In some studies. May 6. the fiber is added to the basal or control diet. For many years attention focused primarily on the evaluation of particular protein sources. This problem is a complicating factor in whether fiber is incorporated by substitution or dilution. The difference in the incorporation method becomes potentially significant when one considers that utilization of a protein varies with the amount of protein consumed. thus causing a dilution of all components of the diet. has also received considerable attention with regard to its effect on protein digestibility and utilization. in which the fiber is added to the diet at the expense of the digestible carbohydrate source. As animals will normally consume diets to meet their energy needs. animals will be consuming a greater proportion of their diet as protein than will animals fed a fiber-free or low-fiber diet. The method of incorporation of the fiber or fiber source into the diet is of particular interest. since it influences the protein-to-calorie ratio of the diet. Animals fed a fiber-diluted diet. One of two methods is usually employed. Hence.26 This problem has been 0-8493-2387-8/01/$0.2387_ch4. In all tables.1_fm Page 133 Sunday. For example. Schneeman Determination and prediction of the quality of dietary protein has been a research interest almost since the beginning of nutrition as a science.50 © 2001 by CRC Press LLC 133 . This chapter is a collation of data from these studies. but the protein-to-calorie ratio is increased. In contrast. either by chemical analysis or by biological assay using standardized diets. . Their results indicated that overall. whereas viscosity could reduce digestibility by slowing protein digestion and absorption in the small intestine. the protein associated with the fiber source will contribute to the fecal nitrogen. usually rats. In the case of fiber-rich sources.48 examined this possibility in cirrhotic subjects fed an animal or vegetable diet. In most cases. experiments feeding fiber sources that would allow the attribute most responsible for the reduced protein digestibility to be identified have not been published. Fiber-rich sources usually contain significant quantities of protein which can influence the digestibility data. to the presence of indigestible protein within the fiber source. Thus. however.2387_ch4.35 The two human studies reported. They found that little fecal nitrogen was associated with the fecal fiber fraction. the viscous and highly fermentable fibers. the source of fecal protein must be from incomplete digestion of the dietary protein. the secreted digestive enzymes. sloughed mucosal cells.50. since part of this protein is indigestible. For the purified fibers containing no protein (guar gum. when the diet was diluted with cellulose.7 who showed that when rats were fed cellulose in the diet by the substitution method. However. Fermentation could reduce protein digestibility by its action of promoting microbial growth. 53 This is consistent with several reports of increased protein in the small-intestinal contents when cellulose or wheat bran is fed.. Both these attributes could contribute to the effect. In contrast to the diets containing purified fibers. and microbial protein. however. Thus. such as pectin and guar gum. the soluble fraction. as they are not mutually exclusive. Thus. the protein efficiency ratio (PER) decreased with increasing cellulose.1. often contains a small amount of protein). and that the rank of the protein digestibilities of the different diets was the same for both rats and humans. have also been shown to significantly reduce protein digestibility. apparent protein digestibilities were similar between the two species. protein digestibility was lower in humans consuming the fiber-rich sources.49 have examined this question directly by measuring protein digestibility in both humans and rats fed identical diets. Two carbohydrates with fiber-like properties. a low protein digestibility may be due. As with the purified fibers. showed no effect of purified fiber on digestibility. fiber-associated protein appears to contribute little to fecal nitrogen. Most studies on fiber digestibility have been conducted in animals. At this time. The question naturally arises as to how good animals are as a model for the effect of dietary fiber on protein digestibility. 2001 6:57 PM 134 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. May 6.13 This points out a potential difference in the source of fecal nitrogen between the purified fibers and the fiber-rich sources. purified fibers. regardless of the diet. where a slight decrease was noted. However. in part. In most instances. inulin and resistant starch.1.1. only apparent digestibility was determined. the PER did not change. regardless of type. Weber et a1. the trend was similar for both. reduced digestibility significantly in animals.1_fm Page 134 Sunday.38 Protein digestibility during consumption of fiber-rich sources is shown in Table 4. at least for vegetable sources of fiber. it would appear that diet dilution is the better method of fiber incorporation. They used diets containing either fruits and vegetables. a citrus fiber concentrate. or insoluble barley fiber as the fiber sources. usually considered a purified fiber. rats would appear to be a very good model for humans in this context. Generally. in those instances where both apparent and true digestibility were reported. the presence of fiber-rich sources in the diet led to significant decreases in both apparent and true digestibility in animals for all fibers for which statistics were reported.34. Thus. except at the very highest level of cellulose.2. 3RD EDITION examined experimentally by Delorme et al. for experiments investigating the effect of dietary fiber on measures of protein utilization. The effect of purified dietary fibers on protein digestibility in animals and humans is presented in Table 4. Large increases in fecal nitrogen with the vegetable diet were due to increases in the bacterial fraction and. reduce digestibility more than the nonviscous and relatively nonfermentable fibers such as cellulose on an equal weight basis. to a lesser extent. Bach Knudsen et al. Which attribute of dietary fiber is most responsible for the reduction in protein digestibility is unclear. 3 r = 0.5) (12.9* 0. May 6.3 9.6* 2 7 7 36 19 50 19 28 46 Ref.9 90.0 20.0 ± 0.7 89.9 93.  Rats. weanling  Rats.9 16.8 74.2 94.6 ± 0.0) (14.0 99.9 89.0 2.0 5.0) (10.6 96.2 5. weanling  Rats.0) (8.0 92.1_fm Page 135 Sunday.5 5.4 92.4 9.801 92.9 85. weanling  25 → 3 5→4 10 Time on Dietb Adaptation → Balance (days) Rats.3 ± 6.5 85.6 8.0 0. weanling  Rats.6 95.0 7. weanling  4→5 21 → 7 21 → 7 21 25 → 3 4 to 56 → 5 Rats.0 12. weanling  Rats.0) Concentration in Diet Fiber Proteinc (%) (%) 92.9 0.0 0.0 ± 5.0 10.1 ± 0.0 2.6 10.5 ± 0.8 ± 1.7 ± 0.0 ± 1.6 89.5 5.0 20.7 90.0 10.5 0.5 5.1 81.9* 81.6 ± 2.0 8.7 91.0 0.0 20.3 85.0 90.2 15.7 d 100.851 89.0 89.5 r = –0.1 4.2 86.6* 86.0 (10.0) (9. 2001 6:57 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 135 .5) (9.4 ± 0.0 NR 91.0 2.3 ± 0.9 ± 0.0) (8. 2387_ch4.7eNR 86.4 11.2 ± ± ± ± Protein Digestibility Apparent True (%) (%) 0.0 10.2 27.1 0.8 89.  Rats.1.6* 93.3 (22.4 ± 0.3 NR 90.5 NR 92.Substitution Substitution Substitution Substitution Substitution Dilution Substitution Dilution Substitution Fiber Cellulose Cellulose Cellulose Cellulose Cellulose Cellulose Cellulose Cellulose Cellulose Rats.2 0.0 20. weanling  and  Species Studied Effect of Purified Dietary Fibers on Protein Digestibility Method of Incorporationa Table 4.8 0.0 ± 4.0 8.9 8.6 ± 0.5 88.0 10.0 0.0) 12.9 10.9 ± 0.8 92. 0 20. 2001 6:57 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. weanling  Species Studied Table 4. 136 Rats.0 92.5 87.6 ± 1.0 1.7* ± 6.0 10.0 0. weanling  Mice.9 ± 0.5 83.8 90.8 ± 0.4* 0.8 86.6 ± 2.0 3.0) (10.2 ± 0. May 6.0 0.0 62.  0.3 (10.5* 90.0 10.2 ± 2.1* 0. weanling  Mice.4 r = –0.4 67.5 g/d NDF 5.5 30.9 87.0 10.7* 6.0 6.0 9.1* 92.0 92.8 (10.3 89.0i 5.8 90.0 20.0 10.0 87.9 88.6* ± 2.0 2. adult.5 g/d NDFg 23.4 67.0 ± 1.0 5.8* 87.0 9.2 28.6 3.5 79.0 10.5 77.0 (10.4) 16. weanling  and  21 28 35 35 28 Rats.1_fm Page 136 Sunday.0 5.5 79.3* 92.8 ± 0.0 0.  28 Time on Dietb Adaptation → Balance (days) Mice.1 (Continued) Effect of Purified Dietary Fibers on Protein Digestibility 2387_ch4. weanling  Human.7* 0.0h 10.5 0.0) Concentration in Diet Fiber Proteinc (%) (%) ± ± ± ± ± ± ± ± 2.0 0.6 ± 3.0 23. weanling  Rats.0 20.9 74.8 10. weanling  Rats.0) 10. weanling  20 → 30 35 14 → 5 Dogs.0 0.1.0 10. 3RD EDITION .8 80.Substitution Dilution Substitution Dilution Substitution Substitution Substitution Substitution Dilution Substitution Substitution Cellulose Cellulose Cellulose Cellulose Hemicellulose Xylan Raffinose Lignin Lignin Acid detergent fiber Pectin Fiber Method of Incorporationa 10 7 → 10 Rats.0 5.0 9.0) (10.4 ± 0.8 85.7 82.59f Protein Digestibility Apparent True (%) (%) 17 14 36 21 12 12 21 38 12 3 21 Ref.4* 93.5 92.1* 82.5 84.8 92.0 74.6 ± 0.0 15.6 8.8 29.5 90.0 62.  and  Rats.9* 37.9 ± 0.8 1.5 ± 3.7 ± 0.0 8.0) (9.8 92.0h 10.7) (9.9 5.7 ± 1.0 10.5 NR 85.0 0. 1* 70.0 0.6 ± 1.4 ± 0.3 ± 1.4 ± 2.3 70.0 2.0 (22.3 ± 0.8 92.9 15 16 30 36 50 12 30 1 36 19 19 2387_ch4.7 8.0 25 → 3 21 0.0 10.0 ± 0. weanling  21 Rats.3 ± 2.3* 80.HDE)j (HMW.5 0.0 10.0 5.8 ± 1. May 6.3* 101.0) (8.3 ± 1.0 0.4 4.5 10.5 86.0 0.0* 87.0 4.8* 79.0 5.8 r = –0.0 25 → 3 Rats.0* 92. weanling  Rats.0* 92.1 94.Substitution Substitution Dilution Substitution Substitution Substitution Substitution Dilution Substitution Substitution Dilution Pectin Pectin Pectin Pectin Pectin Pectin Pectin Guar gum Guar gum Guar gum Guar gum 21 3→8 4→5 Rats.5 81.9 ± 0.4* 89.0* 91.2 87.0) (8.9 ± 1.5) (9.5 ± 1.0) (9.8 0.3 9.0 10.8 ± 1.7 ± 0.9* 92. weanling  Rats.0 20.LDE) (LMW.0 20.2* 87.5 5.1_fm Page 137 Sunday.5* 0.0 9.3 ± 0.0) 10.0 2.4 9.2 77.8 ± 1.9 ± 0.0* 92.5 ± 0.2* 89.8* 84.5 10.0) (HDE) (LDE) 10.68f 92.0 10.5 83.8 78.6 86.5 88.9 88.9* 63.1 ± 21 r = –0.4 82.4 ± 0.0 7.8 ± 0.6* 92.06* 81.78f 95.0 ± 1.0) (HMW.5* 92.0 (14.0 4.0 10.0 ± 0.7 ± 1. weanling  Rats.2 86.2 ± 5.2* 83.5) (9.8 ± 0.5) 93.1 ± 1.6 ± 0.2 78.4 ± 0.0 8.HDE) (LMW. weanling  Rats.0 0. weanling  4 to 56 → 5 35 4→5 Rats.0 10.4* 79.3* 92.4 88.9 88.4 ± 0.4 ± 0.0) (10.  Rats.0 10.  Rats.7 84.8 85.4 86.0 0.  3→7 0.9 ± 0.5 5.1 ± 1.3* 87.0 10.5 94.LDE) (10.5 78.0 ± 0.3 ± 0. weanling  9.3 79.3 ± 0.3 ± 0.0 10.0 (10.9 ± 1.0 0.0) (9. weanling  Rats.4 ± 0.0) (8.3 ± 1.0) (10.8 0.7 4.4 ± 1.0 0.1* 101.0* 81.3* 0.8 ± 1.5 91.4* ± ± ± ± 0.4 0. 2001 6:57 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 137 .0 7.8 ± 1. 2001 6:57 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.6 ± 3. Na Carrageenan Carob bean gum Ispaghula husk (Isogel) Dilution Substitution Inulin Resistant starch Dilution Substitution Guar gum Fiber 21 → 7 4 to 56 → 5 Dogs.5 75.  14 → 5 3→8 3→8 10 3→8 3→8 21 → 7 Dogs.0 8. weanling  Rats.2 88.  Species Studied 0.6* 87. adult  and  4 to 56 → 5 Time on Dietb Adaptation → Balance (days) Rats.7 ± 0.Dilution Substitution Substitution Substitution Substitution Substitution Guar gum Agar Alginate.0 10.0) Not stated 8.0 0 7 0.6) (14.5) (23.0 10.0 ± 0.0 0.6* 87. 138 Method of Incorporationa Table 4.3 ± 1.5 NR Protein Digestibility Apparent True (%) (%) 50 53 33 16 16 46 16 16 53 50 Ref.0) 10.9 ± 1.2* 92.0 10.  &  Rats.5 82.7 g)k 25 g (43.0) Concentration in Diet Fiber Proteinc (%) (%) 90.8* 88.0 10.0 (14.8* 90.2 85. weanling  Humans.2* 87.0 0.8 ± 0.5 79. 3RD EDITION . weanling  and  Rats.1_fm Page 138 Sunday.5 81.0) 8. weanling  Rats.1.0 10. May 6.5 g)k 0 7 0.0 8.8 ± 0.0 8.0 (24.3* 87.0 (10.5 87.8 ± 0. weanling  Rats.5* 89.3 89.5) (23.0 ± 0.9 ± 1.5 NR 88.5 ± 0.7 85.0 0g (19.1 (Continued) Effect of Purified Dietary Fibers on Protein Digestibility 2387_ch4.  &  Rats.7 83.6 ± 0.0 (24.0 0.0 0. Na Alginate. as the actual crude protein concentration was not reported. pectin. 65% esterified. Note: An asterisk indicates that the value is significantly different from fiber-free or low fiber control group (p < 0.4%) 2387_ch4. 32. 2. otherwise reported as grams per day or grams nitrogen (N) per day.9 g/1000 kcal 68.0* 12 52 Indicates method by which fiber or fiber source was incorporated into the diet. L. Mixed (oligofructose & sugar beet fiber. Substitution signifies addition to the diet at the expense of the digestible carbohydrate source. MW. See text for further explanation. Total dietary fiber. Where two values appear separated by an arrow. Values in parentheses indicate concentration of protein sources (usually casein) in diet.Dogs. molecular weight. Slow setting.0 9. or where only one value is present.5 83. DE. H.  Rats. indicates the length of time for which sample collections were made. In the case of dilution.2 0 5 10 10. 55% esterified. NR. the first value is the time allowed for adaptation to the experimental diet.3 ± 0. 4:1) Mixed (cellulose.1_fm Page 139 Tuesday. The correlation is statistically significant.05). resulting in a whole diet dilution.0 10. degree of esterification. no statistics reported. Dilution signifies addition to the diet.5%. xylan. NDF.8 ± 1. neutral detergent fiber. Indicates time of the experimental diet. the fiber or fiber source concentration in the diet indicates the actual concentration of fiber in the diet.8 g/1000 kcal k j i h g f e d c b a 92. 2001 2:59 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 139 . Correlation between the percentage of fiber in the diet and the apparent protein digestibility. May 8. raffinose. low.5 86.8 ± 0. weanling  Substitution Substitution 35 35 → 7 0.6 65 g/1000 kcal 66. Mean ± SEM. Fast setting. not the percentage of fiber added to the diet. The value after the arrow.2* 87. high.8 82. 0 g/d)e 12 g (53.2 2387_ch4.8 g/d)e 45 g (12.8 (10.  Pigs. 3RD EDITION .9e 12.4 1.4* 0.5* 1.3* 0.3 85.5 g N/d (15.7) (18.8 0 (3.0 12.2 i 6.13) (8.9 89.5 10.4 ± 92.0* 91 ± 3 NRf 85 ± 4 67 64* 55* 92.9 91.5 0.8 88.7) (17. weanling  Humans.0) 10.8 96.1.1_fm Page 140 Sunday.5 13.6 ± d ± ± ± ± ± ± 97. weanling  Humans.4 g N/d 16.0 10.8* 67 61* 57* 92.9 87.8 18.2 1.4 91.9 77. adult  Chickens.0) (9.0) (8.3 16.3 0.0 16.3 ± 77.7 16.1 12.5 78.6 0.5 0.1 7.01 93. May 6.0 ± 86.5 15.0 12.3 ± 89.0 ± 92.1) (18.8* 54 4 12 54 55 42 8 11 12 36 30 Ref. 2001 6:57 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.0 16.0* 97.3 95.0* 89.1) (10.6 (8.6 15.8 ± 87.0 10.4 g N/d 9.0 g/d) 4.7 ± 87.6* 101.6 0.0 0.21 6. 140 Rats.2 92.  Rats.  Rats.Substitution Dilution Substitution Dilution Substitution Dilution Substitution Substitution Substitution Dilution Substitution Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Corn bran Oat bran Oat bran Method of Incorporationa 20 → 10 7→7 7 Pigs.5) (9.8 83.90) (15.0 5.5 g/d) 0(4.2* 0. weanling  Rats.9 16.0 6.2 1.0 0.9 11.3)h 5. adult  Rats.5 0.  9→5 7 14 35 18 → 4 35 21 4→5 0.9 97.7 Protein Digestibility Apparent True (%) (%) 1.2* 92.0 22.3 15.2 96.2 20.8 0 (33.0) Concentration in Diet Fiber Proteinc (%) (%) 0.0 20.4* 0.9) (21.5 9.  Species Studied Time on Dietb Adaptation → Balance (days) Effect of Fiber-Rich Sources on Protein Digestibility Wheat bran Fiber Table 4.4) 10.4* 1.7 g N/d 10.  Chickens.8 83.38 4. 0j 93.3* 91.7 9.4 96.8 28.5 16.6 14.4 8.6 82.7 (15.0 94.6 90.0) (16.5 14.7 0.4 15.  Rats.7 0.0 10.0 7.7e 0(4.7 97.5 94.8 12.3)h 5.1 92.3e 14.0 97.9 ± 2.0 7.4 13.4 13 14 (16.7 23.4e 14.6 14.1i 6.6) 9.7 96.7 9.6 26.5 ± 2.3e 4.9) (11.2 79.9 25.0 7.1 11.9 15.  Pigs.Substitution Substitution Substitution Substitution Substitution Substitution Substitution Dilution Dilution Dilution Substitution Sorghum meal Pea fiber Barley fiber.4 ± 1.9 15.7 12. insoluble Barley hulls Barley fiber.7 21.3* 63 60* 55* 87.9 16.1_fm Page 141 Sunday.4e 14.0NR 87.6 0 12 16. May 6.1 20. adult  5→6 Humans.4 8.9 14.6 g N/d 97.2 0.5 87.0 7.9* 47 42 42 42 8 10 49 2 49 54 5 2387_ch4.1 5.0 83.3 10.8 9.6) 9.  Rats.1* 56.  7 Chickens.0 5.1* 94.0 86.7) (15.4 4.3 98 92* 65. insoluble Barley husk Barley husk Maize hulls Oat hulls Soybean hulls Soybean hulls 14 → 7 4→5 17 → 5 4→5 9→5 20 → 10 20 → 10 20 → 10 16 Rats.9 98.2 0.  Humans.3) (16.4 13. 2001 6:57 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 141 .0j 95.4) (15.7 97.9 80.8 5.  Rats.8 9.0 88.9 11.4) (15.7 91.6 0.7 21.7 9.0 7.9NR 100.6 15.7 88.8 60.  Rats.8 86.  Pigs.9* 97. adult  3.0 14.3 88.9) (11.  Pigs. 6 92.1_fm Page 142 Sunday. 142 35 Rats.7 74.4 80.0 11.5 0.5 21.2 ± 90.0j 91.0 12 0.4 93.8 ± 2. 2001 6:57 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.9 ± 88.8* 1.05 4.6 ± 83.0) Concentration in Diet Fiber Proteinc (%) (%) 84.2 0.0 10.41 14.5 ± 0.0 ± 90. weanling  Rats. May 6.5* Protein Digestibility Apparent True (%) (%) 53 30 47 18 47 12 47 42 42 51 Ref.1 12.8 19.3) 93.  Dogs.5 16.6* 2.0 7.0 ± 98 90* 92.4 13.0 10.03 7.  Dilution Lupin hulls 10 → 4 Substitution Dogs.21 13.8 g/d 0.  Dilution Pea hulls 20 → 10 Pigs.0j 89.0 10.6 101.  Substitution Bean cell wall 16 Rats.  &  Species Studied Time on Dietb Adaptation → Balance (days) Soybean hulls Fiber Method of Incorporationa Table 4.1 g/d 14.5 1.4 82.8 13 14 13.4 97.2 1.1.9 16.5) (23.8* 98 90* 92.8* 92.  Substitution Canola hulls 20 → 10 Pigs.8 85.  &  Substitution Formulatedh Substitution Substitution Dilution Soybean cell wall Soy polysaccharide Canola cell wall Sugar beet fiber Sugar beet fiber 3 → 4–5 16 3.6 14.9 15.09 6.3 g/d 13 13 (10.4 13.32 0.8 65.0 0.2 97.6 28.0 81.3 81.4 0.0 g/d 8.8 25.1 12.6 14.3 98 87* 92.9 15.2 (Continued) Effect of Fiber-Rich Sources on Protein Digestibility 2387_ch4.7 83.5 13 15 10.0 ± 1.  Rats.6 81.0g 0 7 (24.  a&nd  Rats.16 4→5 21 → 7 Humans.7 86.0 7.8 ± 84. 3RD EDITION .9 1.9 90.4 83.0 12 0.0 12 0.2 ± 1. 8 ± 3.6 g/d 17.5 86. May 6.8* 81.8 g N/d 16. wheat.001) with increased NDF intake.9 ± 0.7* 92.3 14.0 84.8 ± 0.2) 13.0 g/d 7.6NR 90.1.9 g N/d 15.5 85.3) (20.2 55. no statistics reported.1 g N/d 13.6 0. adult  Humans. adult  Humans.5 ± 0.6 N/d 14.9 81.74 g N/d 4.5 g N/d (19.1.6 4 20 49 49 25 23 22 40 57 27 2387_ch4. adult  Humans. and whole meal bran Fruits and vegetables Fruits and vegetables 1 → 14 Humans. adult  Rats.3) 26 g/d 27 g/d 10. pea fiber. See footnote b of Table 4.  Unclear 19 → 7 4 Substitution Barley.3) (19.5* 91 ± 3 NR 69 ± 2 83. adult  Rats.1_fm Page 143 Sunday.8 g N/d j i h g f e d c b a See footnote a of Table 4.7 6.65 g N/d (15.8 81.2 ± 1.4 g/d 25.1.5) (17.0 g/d 13.4 ± 0.9 g/d 10.1 72. Fruits and vegetables Fruits and vegetables Fruits and vegetables Konjac and seaweed “Guatemalan” diet Humans.0 18 g/de 3.1 91.Dilution Substitution Substitution Substitution Substitution Humans. Mean ± SEM.0 g/d 19. Neutral detergent fiber. Treated with hot water to increase the solubility of pectin.2 g/d 31.5 48.9 g N/d 14.1 ± 0.8* 86.7 g N/d 10.7 g/de 13.0 g/d 4.3 14.0 g/d 96. adult  Unclear Unclear 2→7 Human.7 g N/d 93.8 g/d 41. Enteral formulas.6 25.  14 5 17 → 5 14 → 7 21 → 7 14 → 7 5.8 g N/d 13.2 ± 3.05).1* 89. vegetables. Statistically significant linear decrease (p < 0.6 g/de 20.5 63.1* 90 ± 2 84 ± 4* 90.1. and pectin Fruits.7* 83. Note: An asterisk indicates that the value is significantly different from fiber-free or low-fiber control group (p < 0.1 ± 1. Total dietary fiber.0 g/d 1.4 ± 3.9 g/de 5.2 ± 0. adult  Human.3) (20. See footnote c of Table 4.0 ± 3.9 ± 0.8 g/d 93.1 g N/d 13.1.5 85. NR.1.9 g/d 3.6 7.  Substitution Brown rice 68. 2001 6:57 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 143 .2 g/d 16.5 90. no trends can be discerned regarding the effect of fibers on NPU.3 and 4. that this is not the same PER as described by the AOAC. However.6). such as PER and net protein utilization (NPU). as well as the method of fiber incorporation into the diet. undoubtedly due to its ease of determination. The effect of dietary fibers on several measures of protein utilization is found in the last two tables (Tables 4. comparisons between studies using the same fiber type are often not possible. However. For example.2387_ch4. It should be understood. often in an approximately linear fashion. A level of 10% or greater in the diet almost invariably led to a significant reduction in this measure of protein utilization.7 or unchanged. in most instances the decrease in urinary nitrogen did not fully compensate for the increase in fecal nitrogen. Guar gum at 10% of the diet led to a significant reduction in PER. This review covers the published literature through November 2000. such as biological value (BV). consequently. Fecal nitrogen increases are often accompanied by a decrease in urinary nitrogen.5 and 4. however. The most common measure of protein utilization is the protein efficiency ratio. Interestingly. However. the results from protein utilization studies are quite variable. it remains to be established whether dietary fiber has a detrimental effect on protein utilization. However. two other soluble fibers showed divergent effects on the PER. .5 to 10.47 than in fiber-free controls.36 whereas alginate did not.46 Few studies have investigated the effect of fibers on NPU and. one might expect a decrease in protein utilization parameters that have a digestibility component.1.1.1. 2001 6:57 PM 144 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.38.12 lower. because of differences in the protein and/or fiber concentrations of the diets.5% range. such that nitrogen balance was often significantly reduced.1. since the cellulose concentrations differed from 1% and the protein concentrations were not always in the 9. the results are too inconsistent to make any meaningful statements about the effect of fibers as a whole. It is apparent from the data that fiber consumption causes a shift in the pattern of nitrogen excretion. in rats the PER of diets containing 10 to 12% cellulose are either significantly higher. although in all cases it remained positive. Given the decrease in protein digestibility and the consequent increase in fecal nitrogen. May 6. 3RD EDITION Tables 4. since the PER is calculated in the same way (body weight gain per protein consumed).4 show the effect of fibers on nitrogen excretion and on nitrogen balance for the purified and fiber-rich sources. and an increase in parameters measuring the utilization of absorbed protein. The fiber with the most consistent effect on PER is pectin.1_fm Page 144 Sunday. From the available studies it is clear that dietary fiber and fiber-rich foods reduce protein digestibility. the term is retained. This occurred with all types of fibers and fiber sources. Unfortunately. Although a large number of studies have reported the BV of fiber-containing diets. 0 139. weanling  Rats.0 ± 3 10.1 0.6 Urinary (mg/day) 250 200 225 275 970 22.  Mice.0 g/d 16.070 0.5 ± 5.0* 10.0 g/d 0 (6.5 12.0*(12*) 47.316 0.  3→7 3→4 7→7 30 → 1 28 21 → 7 Rats.0 0.910 0.0 ± 7 50. weanling  Species Studied Time on Dietb Adaptation → Balance (days) 1.  35 → 7 21 5→4 Rats.8 Nitrogen Balance (mg/day) 24 24 41 17 21 29 29 44 36 18 Ref.0 ± 9 50.0 15.2 NR 12.0 ± 2 10.9 19.0± 12 Fecal (mg/day) Nitrogen Excretion 970* 1220 820* 1100 8.6 NR 45.4 ± 8.0*(16*) 16.0*(10*) 36.  Humans.880 7.256 0.8 ± 3.60 106.0 5.0 18.0 114.1_fm Page 145 Sunday.0 21.0* 50. adult.2cNRd 20.132 0.3 9.4 NR 25. growing  and  Monkeys.319 0.9±3.0 2.0 10.0 ± 1 20.8 g/d)i 14.1 5.0 ± 4.7 ± 1.0 2. 2001 6:57 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 145 .Substitution Dilution Dilution Substitution Substitution Substitution Substitution Dilution Dilution Dilution Cellulose Celulosef Celluloseg Celluloseh Cellulose Cellulose Cellulose Cellulose Cellulose Method of Incorporationa Cellulose Fiber 21 → 7 Rats.1 ± 1.0 g/d) 0 (6.0 20.1.4 ± 4.3 ± 6.2 g/d (21.7 181.0 0.0 0.4 NR 11.070 0.264 0.266 0.6 51. May 6.080 0.3 NR 249.0 ± 30 131.0 186.2 g/d (21.3 Effect of Purified Dietary Fibers on Nitrogen Excretion and Balance 5750 5740 1170* 5950 1140* 910 5840 163.0 g/d 0.0 ± 2 133.  and  Humans.0 20.2 ± 2.0 2. adult.0(6)e 31.0 g/d) 0.0 10.132 0.1 ± 5.289 0.188 Nitrogen Intake (g/day) Table 4.0 ± 1 10.0 20.0 17.0 ± 3 10.0 ± 2.289 0.0 10.276 0.1 22.8 g/d)i 14.287 0.4 ± 1.0 20.0 g/d 20.0 Fiber in Diet 7.0 g/d 7. weanling  Rats.0 154.0 165.0 10.0 ± 10 40. 2387_ch4.  Rats.121 0. 289 0.0*(14) 10. adult.0(10*) 40.070 7.0 g/d) 0 (6.0 6.  Rats.0 ± 6 1100 30.  Humans.850 7. 146 Humans.0 ± 2 10.0 10.3 (Continued) Effect of Purified Dietary Fibers on Nitrogen Excretion and Balance 2387_ch4. 3RD EDITION .880 0.  Mice.289 0.0 26.2 g/d (21.0 ± 5* 970 10.0 ± 4 18.0 ± 15 40.2 g/d (21.0 ± 8* 181.0 1380* 1910 860* 1220 740* 30.0 0.850 Nitrogen Intake (g/day) 10.2 g/d (21. adolescent.0 187.  Humans.0 3.  Mice. weanling. May 6.070 0.0 ± 5 5840 20.0 10.060 0.8 g/d)i 14.0 ± 4 40.2 g/d (21.0 ± 4 10. weanling.0 ± 6 1360* 1910 Nitrogen Balance (mg/day) 29 21 36 24 24 24 21 24 Ref.0 g/d) 5.0 20.0 ± 4* 5080 Urinary (mg/day) 860 Fecal (mg/day) Nitrogen Excretion 40.0 ± 7 96.275 0. 2001 6:57 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.8 g/d)i 14.0 0 (6. adolescent  Species Studied Time on Dietb Adaptation → Balance (days) Table 4.0 ± 4 20.8 g/d)i 14.0 ± 4 5570 920 20.0 5730 1110* 22.0 80.Dilution Substitution Dilution Dilution Dilution Dilution Substitution Substitution Hemicellulose Hemicellulose Hemicellulose Hemicellulose Lignin Lignin Neutral detergent fiber (wheat bran)g Method of Incorporationa Cellulose Fiber Rats. weanling  Humans.1 0 (6.0* 5080 5730 1290* 860 5750 5990 1180* 910 10. adult.0 g/d) 0.0 5.8 g/d)i 14.910 0.0 2.0 ± 6 20.1_fm Page 146 Sunday.0(6)e 29.070 0. g/d) 0 (6.050 0.264 0.0 ± 9 20.0 Fiber in Diet 0.  and  21 → 7 28 21 3→4 3→7 3→4 28 3→4 20.0 ± 5 20.297 7.1.060 7.0 ± 3 10. 0 g/d) 0 (6.0 g/d) 1.0 132.0 174.7* 30.0 90.289 0.0*(15*) 30.1_fm Page 147 Sunday.1 0.316 7.0 5840.  Rats.0 139.0 182.4 (HDE) 0.  Rats.  Humans.0* 20.0 g/d 0.0 154.0 2.910 0.  Rats.  Substitution Humans.8 g/d)i 14.0 1060.1 0.321 0.146* 7.3 (LDE)j 9. 2001 6:57 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 147 .5 23. adolescent.0* 181.0 34.0 20.0 139.319 0.0 g/d 7.0 24.0 182.0*(11*) 31.0 18.0 g/d 20.0 10.0 1100.0 154.9 122.2 g/d (21.1 0.8 g/d)i 14.1 0.0 970.2 g/d (21.0 172. adult.  3→4 3→7 Humans.  Substitution Neutral detergent fiber (wheat bran)h Acid detergent fiber (wheat bran)h Acid detergent fiber (wheat bran) Pectin 0.850 7.0 2.  Rats.8 g/d)i 14.289 0.0 97. May 6.0 2.  Substitution Substitution Substitution Substitution Substitution Dilution Dilution Dilution Dilution Dilution Pecting Pectinh Pectinh Pectin Pectin Pectin Pectin Pectin 21 7→7 Monkeys.0 5.0 g/d 13.0 25.0 118.0 181.332 5010* 1010 190 250 210 230 22. weanling.0 g/d) 0 (6.  and  3→4 21 → 7 Rats.0* 96.  and  21 → 7 Rats.319 0.0 139.0 20.319 0.288 0.0(6)e 33.1 0 (6.0 154.21 → 7 4→5 21 → 7 21 → 7 21 → 7 Rats.0 9.1 0.289 0.0 36 41 24 24 24 29 29 29 30 29 29 29 2387_ch4.0* 20.241* 0.880 0.0 910 96.2 g/d (21.0 22.0* 1830 1910 1310 1220 1090.319 0.0 154.0 25.316 0.0 13.0* 10.0 20.  Rats.0 2.0 159.0*(19*) 5080 5510 1060 860 5750 5510.191* 0.0 132.0* 18.0 0. adult.0 25.0 159.0 2.0 2.289 0. 237 Egg albumin 0.4 57.0*(12*) 44.289 0.  Rats.0* 17.0 0.0 6.0 52.0(6)e 48.8 33.0 0.0 10.0 10.Substitution Dilution Dilution Substitution Dilution Substitution Guar gum Guar guml Guar gum Guar gum Guar gum Carrageenan Method of Incorporationa 21 148 Rats.0 0.9 ±1.5 26.2% 58.6 33.6 43.8 ± 1.0 44.335 0.0 10.0% 2.0 20.0 5.0 53.0 5.0 10.0 0.6* 152.5 57.0 5.0 4.1_fm Page 148 Sunday.3 (Continued) Effect of Purified Dietary Fibers on Nitrogen Excretion and Balance 2387_ch4.6 234.0 k Nitrogen Balance (mg/day) 17 36 30 15 45 16 Ref.0 5.7 22.234 0.8* 22.5* 92. weanling. 148 Fiber Time on Dietb Adaptation → Balance (days) Table 4.259 0.109 0.142 0.0% 4.  Species Studied 0.114 0.0 0.168 0.0% 3.0 49. 2001 6:57 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.  and  4→5 Rats.0 Fiber in Diet 0.152 Nitrogen Intake (g/day) 21.217 0.3* Urinary (mg/day) Nitrogen Excretion 17.0 56.311 Casein 0.0 20.0*(17*) 61.0 18. 3RD EDITION .0 43.3 129.0*(20*) 24.9% 3.0 228.8* 10.1.2k 65.0* Fecal (mg/day) 55.5 ± 1. May 6.0 0.  Rats.0 15. weanling. weanling.0 66.  Rats.3 ± 0.0 75.0 ± ± ± ± ± ± 69.  8 3→8 Rats.5 42.0 2.0 5.8%m 3. 0k 66. adult 0.6 24. no statistics reported. h Metabolizable energy of diet was 67 kcal/day. L.2 ± 1.0 10.3 ± 4.5 ± 1.1. a Note: An asterisk indicates that these values are significantly different from the fiber-free or low-fiber control group (p < 0.05).0 10.0k 70.8 ± 0. 2001 6:57 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 149 . Substitution Carrageenan –3.1. g Metabolizable energy of diet was 57 kcal/day.7 mg/kg 106. degree of esterification.4 ± 6.0 10.1_fm Page 149 Sunday.5 ± 1.6 51.3 ± 0.0 66. k Calculated from intake and excretion means. low.2% dietary arginine groups. e Endogenous fecal nitrogen.  Rats. l Data chosen from 1.0* 51.0* 17.0 10.15 g/kg 0.154 0.3 ± 0.0 66.0 3→8 118.Substitution Substitution Substitution Substitution Carob bean gum Na-alginate Algar Vegetable Rats.7* mg/kg 85.9 ± 1.0 ± 6. i Total dietary fiber intake.3* 58. m Values are mean ± SEM for nitrogen excreted per day as a percentage of ingested nitrogen.142 0. j De.1 mg/kg 6.2 ± 1.3 ± 0.4 ± 4.142 0. May 6. then fed an energy-restricted regime with or without cellulose.6 61. H.1.5 mg/kg 27.1* mg/kg 17.4* 17.134 0.9 ± 1.0 3→8 3→8 0.6 27.6* 17.4 ± 1.6 61. Nitrogen intake and balance calculated from data presented in article.6* 11.8 58.6 28.9 ± 1.  Rats.6 ± 4.8 mg/kg 0. c Mean ± SEM. d Nr.6 49.9* 58. weanling.9 ±1. weanling.138 58.149 0.0 0.9 ± 0. uncorrected for nitrogen content of nonprotein diet.  Rats.0 45.5* b See footnote a in Table 4.0 48 16 16 16 16 2387_ch4.3 ± 0.0 66.6 38. weanling. weanling.4 ± 1. See footnote b in Table 4.5 mg/kg 66.67 g/kg 0.2 mg/kg 124.1.7 ± 1.142 0. f Rats were made obese. high.  Humans.142 0.0 3→8 3→6 0. 0% 2.  Substitution Oat brani Dilution Substitution Sorghum fiber 21 Humans.  Not stated Wheat bran 4–5 → 7 Substitution Substitution 18 → 4 Humans.5 19.0 12.7→7 Sugar beet fiber Locust bean gum Monkeys.0 87.4 Effect of Fiber-Rich Sources on Nitrogen Excretion and Balance 2387_ch4.2 ± 0.8d 19.0% 0.0* 6300 8200* 360 200 270 420 NR 280 Urinary (mg/day) Nitrogen Excretion 2020 ± 230 1470 ± 210* 1110 ± 310* 800 ± 440k NR 260 ± 320 181.0 Nitrogen Balance (mg/day) 41 30 4 5 6 43 39 11 45 29 30 Ref.9 10.9 10.0% 5.6 10.0 g Fiber in Diet 4020 4139 4730 1000 1610 ± ± ± ± ± 270 240 390 320 NR 560 26.8% 5.0% 0.289 0.7 18.  and  Humans 14 Dilution Wheat bran (largely) Wheat bran 3 → 15 Rats.0* 6.0* Fecal (mg/day) 1400 NR 2000 1400 ± 300 18.0 80.0% 0.0% 10.0% 12.0 5. 3RD EDITION .0 237.  Humans.0 185.9 1150 ± 210 NRe 1650 ± 220 10.4% 3.1_fm Page 150 Sunday.1 ± 0.287 0.0% 0.6 10.7 ± 0.0 26. adult Humans. 2001 6:57 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.  Species Studied Humans Substitution Wheat bran Method of Incorporationa 0.0 18.0% 10.3%h 4.2* 260 280 2490 ± 100* 21. adult.1.8 g/dj 12.0c 223.9 0.335 0.0 g 0 (21.8 g/d)g 31 g/d (53. May 6.5 26.8 10.5 ± 0.0 92.0 g/dl 0. 150 Fiber Time on Dietb Adaptation → Balance (days) Table 4.  4→5 Substitution Wheat fiberf 14 → 7 Rats.0% 1.  5→6 Substitution Wheat bran 4→5 Rats.6 10.2 g/d) 3.0 g/d 0.322 Nitrogen Intake (g/day) 5500 4830 4960 8390 8160 ± ± ± ± ± 96.0 g/d 15.  Dilution Wheat bran 28 → 7 Rats. adult  1 → 15 3→6 4→7 5.Substitution Substitution Vegetables Brown rice Humans.0 g/dl 3. no statistics reported. Corrected balance.  Dilution Rice hulls 7→7 Monkeys.1.8 13.6 g/dj 20.7 g/dj 13. See footnote b in Table 4. Crude fiber. Calculated from intake and excretion means. 2001 6:57 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 151 .  Dilution Oat straw 7→7 Monkeys.65 4. May 6.4 14.1 g/d 19. NR.  Dilution Soy husks 7→7 Monkeys.0 g/d 20.0 g/d 18.9 g/d 1.1 13.5 g/d 55. with one subject omitted.  Dilution Slippery elm ± ± ± ± 80 120* 140* 180* –710 ± 290 –1090 ± 330 –282 531 1090 740 1380 1840 2280 2600 27 48 43 23 22 41 41 41 41 41 2387_ch4.0 g/d 20.9 15.0 g/dl 1.4 g/d 25. Note: A single asterisk indicates that the value is significantly different from the fiber-free or low-fiber control group (p < 0.64 9.8 10.65 ± 0.05). Neutral detergent fiber.6 g/d 0.0 g/d 1.9 g/dj 10. adult (cirrhotic) Humans.74 ± 0.0 g/d 15. Not stated Cabbage Humans 14 → 7 Humans. Data from untoasted oat bran group study.1.4 13.1.0 g/dl 1.0 g/d 16.  Substitution 19 → 7 Substitution (?) Humans.0 g/dl 1.9 190 390 1500 ± 140 2410 ± 430 1400 2100 955 2291* 200 335 200 385 205 470 335 320 1350 2480 3950 ± 390 3330 ± 510 8798 7080* 11450 10810 f e d c b a See footnote a in Table 4.0 g/dl 1. Highest level fed. High protein diets.6 g/d 4.3 g/d 12. l k j i h g Total dietary fiber.  Fruits and vegetables Fruits and vegetables 7→7 Monkeys.0 g/d 18. Mean ± SEM.  Dilution Psyllium seed 7→7 Monkeys.1_fm Page 151 Sunday.1.6 14. 94 9.0 10.0 10. May 6.72 ± 0.7 ± ± ± ± ± ± 4.1 80.7 58.29 ± 0.5 5.6 (8.0 20.0 10.0 2.0 20.0 20.06 ± 0.70 ± 0.0 20.8 5.0 65.98 10.41) PER 79.1 3.8 63.1 5.5 Effect of Purified Dietary Fibers on Measures of Protein Utilization 2387_ch4.0 61.89 12.0 10.7 1.2 1.0 72.7 ± 0.3 4.Substitution Substitution Substitution Cellulose Cellulose Cellulose Substitution Substitution Cellulose Cellulose Mice Rats Rats Rats Rats Rats Species Studied 0.9 80.3 76.0 76.0 76.0 40.58 8.0) 9.24 2.89(2.95 8.11 2.3 4.7 ± 0.1_fm Page 152 Sunday.7 84.0 (10.24 ± 0.1 83.0 0.2 78.98 8.8 ± 0.21 2.0 0.5 5.4 64.0 (12.38 9.0 20.0) 9.3 2.0 2.0 5.6 60.0) 19.22 2.9 59.0 20.3 ± 0.06* 2.2 3.0) 7.2 ± 0.5) Concentration in Diet Fiber (%) Proteinb (%) 2.0 5.0 0.98 ± 0.10 2.0 4.12 2.0 10.0 BV 21 46 7 28 12 19 Ref.4 57. 152 Dilution Substitution Cellulose Fiber Method of Incorporationa Table 4.0 0.6e NRf 5.06 ± 0. 3RD EDITION .0 5.50)d 3.0 10.0 Measures of Protein Utilizationc NPU ± ± ± ± ± ± 3.19* 3.0 40.1 2.12* 1.0 20. 2001 6:57 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.05 11.0 5.0 64.0 10.7 5.2 6.82 ± 0.0) 12.0 10.4 ± 0.1.05* 2.0 0.57 ± 0.0 78.0 10.0 10.0 69.38 11.0 79.61 11.82 10.5 1.6 NR 3.7 (22.0 0.5% of calories (12.25 (10.0 0.36 ± 0.94*(3. 86 1.0 20.71 ± 0.79*(3.0 6.0 0.50)c 3.38 (1.1 10.044% N) 3.30 ± 0.0) 3.0 10.4 70.4 78.  Rats Rats Substitution Cellulose Rats Dilution Cellulose 3.0 5.15i (4.19) 75.3* 29 29 36 21 47 2 29 36 2387_ch4.0 10.96* 2.1 0.98 ± 0.0k 2.1 0.0) 3. May 6.15i (4.0 20.29 (4.75 10.89 (2.24 (3.0l 2.3 2.3 ± 0.23) 75.99 2.22 (76) 3.47) 4.2 0.98 9.7 1.19* (4.40 ± 0.0l 2.4 1.30 ± 0.83 ± 0.0 10.3 ± 0.7* 4.31 ± 0.0 3.1_fm Page 153 Sunday.0 87.0 0.1 ± 0.0 10.05 ± 0.2 ± 0.14g(87)h 3.0l 2.91 1.0k 2.4 87.86 1.2 63.78 (2.49 ± 0.50 (4.86 1.0 0.7 4.78 (2.63 ± 0.00 2.0k 2.4 77.89 ± 0.20 (3.28) 1.044% N) 12.37 (81) 10.0 (10.6 2.5* 1.78 (2.1 5.0 63.14) j 3.79 ± 0.98* 0.99* 12.9 70.0 9.1 0.21 (85) 10.95 1.3* 75.9 71.9 ± 0.28 ± 0.5% N) 13 13 9.9 74.98 ± 0.40 ± 0.48 ± 0. 2001 6:57 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 153 .0 (10.0 5.0 12 0.1 0.0 5.14) j 4.2 15.0 ± 0.90 ± 0.20 (67) 2.60 ± 0.14g(87)h 2.0 12.89 2.20) 3.89 2.91 1.28) 4.9 3.89 2.28 ± 0.31 ± 0.7 93.044% N) 3.Rats Rats Substitution Substitution Substitution Substitution Substitution Dilution Substitution Substitution Cellulose Cellulose Xylan Hemicellulose Lignin Lignin Neutral detergent fiber (wheat bran) Acid detergent fiber (wheat bran) Rats Mice Mice Rats Rats.20 (83) 20.16 ± 0.4* 95.7 63. 154 Rats Rats Species Studied Table 4.6 (8.8 ± 0.11* 3. HMW)m 10.0i 0.0 19.0 11.14)j 3.1 0.0 Concentration in Diet Fiber (%) Proteinb (%) 36 29 1 12 46 19 Ref.0 64.0 0.0 20.5* 82.0 66.9 66.6 ± 5.3 67.00 ± 0.4 ± 18.0 10.0 (HDE.0 5. LMW) 10. HMW) 10.0 83.0 78.50)d 2.83 ± 0.05 ± 0.0i 10.89 2.15i (4.47)* 2.31 ± 0.0 71.1 81.5 2.4* BV 67.0 63. 2001 6:57 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.0 10.5 5.7 ± 7.0 2.96 1.1_fm Page 154 Sunday.Substitution Substitution Substitution Substitution Substitution Dilution Pectin Pectin Pectin Pectin Pectin Method of Incorporationa Pectin Fiber Rats Rats Rats Rats 3.0 10.32 ± 0.0 2.5) PER 5.3 ± 0.0 2.98 9.0 j 0.3* 2.1. May 6.2* 5.0 (HDE.2 63.4 ± 8.38)* 3.5 (Continued) Effect of Purified Dietary Fibers on Measures of Protein Utilization 2387_ch4.0* 85.0 20.7 (22.4 4.0 (LDE.0 2.044% N) 9.1 0. LMW) 0.0 63.0) 7.0 0.37)* 75.91) 64.5 5.2 3.3 6.59 ± 0.0 76.21* (1.3 60.28 ± 0.44 ± 0.40* (2.31*(40) 10.78 (2.0 (LDE.5 5.98 ± 0.33*(66) (10.0 2.11 ± 0.0 72.86 2.0 0.0) 10.9 76.5 ± ± ± ± ± 8.0* 75.14g(87)h 3. 3RD EDITION .04* (3.9 ± 8.0 10.0) 12.0i 10.47* (3.42*(65) 1.95 (10.4 74.0 20.39 ± 0.0 66.4 78.89 (2.6 ± 0.0 Measures of Protein Utilizationc NPU 3.54 ± 0.3 2.2 ± 0. 0 0. May 6.32*(47) 20.1 79. no statistics reported.65 ± 0. NPU.57 ± 0.4 2. Metabolizable energy of diet was 57 kcal/day.0 3.50.4 (LDE) 0.3 (HDE)m 9. low. 2001 6:57 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 155 . BV.9 ± 0.4 84.45 ± 0.56)* See footnote a of Table 4.30 (4. Values in parentheses are the concentrations of the protein source (usually casein) or the percentage of nitrogen (where so indicated) in the diet.32) 4.14g(87)h 3.35 (4. NPR ÷ apparent digestibility (NPR/AD). PER. L.31 ± 0. protein efficiency ratio. H.9 ± 5.0) 5.48 ± 0.0 3.28 ± 0. molecular weight.14)j 4.98 ± 0.19 ± 0.9 ± 4.3 ± 3.0 0.1.24 ± 0.6 ± 3.Substitution Dilution Substitution Guar gum Guar gum Algin Rats Rats Rats Rats (10.1.2 2. DE.0 m l k j i h g f e d c b a 82. Mean ± SD. NR.1 4.38(79) (10.0) 3. net protein utilization.00 ± 0. Substitution Pectin 46 36 30 30 2387_ch4.33 ± 0.9 ± 4.2 ± 0.1_fm Page 155 Sunday.0 0.0) (10. Relative net protein ratio (RNPR). biological value.0 10. NPR ÷ true digestibility (NPR/TD).0 82.0 9.0) (10.05).35 ± 0.4 81. PER.64* (3. Note: An asterisk indicates that the value is significantly different from the fiber-free or low-fiber control group (p < 0. Metabolizable energy of diet was 67 kcal/day.30) 3. MW. Net protein ratio (NPR).15i (4. degree of esterification.18*(76) 10. corrected for fiber-free group = 2. high. 8 10.4 83.89 (2.17* (3.0 10.3* 90.98 10.82) 2.1_fm Page 156 Sunday.02 (2. May 6.4 80.12*(55) 20.26* (3.  Rats Rats Rats Rats Species Studied 2.57 ± 0.61) 3.50) 3.0 0.38 ± 0.14e.0 0.5% N) 2.1 Measures of Protein Utilizationc NPU 5.2* 82.5 88.3 83.3 85.22 ± 0.26 (2.9l 11.6 83.1l 7.8 9.0 j 5.22*(70) (10.3 BV 95.8 86.28 ± 0.31 ± 0.1.044% N) 2.03* 1.87 (2.Substitution Substitution Dilution Substitution Substitution Substitution Substitution Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Corn bran Barley hulls Fiber Rats Rats Rats.50 (10.89 3.22* (3.7 94.18*(64) 10.1 ± 5.0 10.92 ± 0.26*) 3. 3RD EDITION .16*) 75.07* 0.0 Concentration in Diet Fiber (%) Proteinb (%) 2 12 8 29 36 24 12 Ref.0 0(4.86 2.59 9. 156 Method of Incorporationa Table 4.4 77.8 9.9 ± 4.61 ± 0.4 87.8 9.33 ± 0.16 ± 0.1 4.f(87)g 2.0 10.2l 6.0 0.52 ± 0.98 12.6 Effect of Fiber-Rich Sources on Measures of Protein Utilization 2387_ch4.0) d PER 87.5 82.0 12.1 63.0 2.0 10.78 (2.6l 0.0 0.50)d 3.0) 9.0k 5.5* 84.14)i 3.3 83.28 ± 0.0 9.15h (4.98 ± 0. 2001 6:57 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.9 69.3)n 5.0 9.22*) 3.38 (1.0 0. 6 80.7 80. Mean ± SD.1.5* 68. corrected for fiber-free group = 2.0 10. Substitution Barley husk 84.4 83. Total dietary fiber. Metabolizable energy of diet was 57 kcal/day. PER. Metabolizable energy of diet was 67 kcal/day.8 (10.0 9.3 83. 2001 6:57 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 157 .0 10.8 9.1.7 77.3)n 5.98 11. See footnote b in Table 4. See footnote c in Table 4.7 9.7* 80. NPR ÷ apparent digestibility (NPR/AD).4 78.1_fm Page 157 Sunday.3* 10 24 10 8 2387_ch4.9i 11.50.57) 82.1.7 12.89 (2.1 92.2 ± 3.  0(4. NPR ÷ true digestibility (NPR/TD).5 83.9 ± 4.9 o n m l k j i h g f e d c b a See footnote a in Table 4.1. Neutral detergent fiber. Relative net protein ratio (RNPR).0 7.  Rats.0i 7. Treated with hot water to increase the solubility of pectin.Substitution Substitution Substitution Barley husk Sugar beet fiber Bran cell wall fiber Rats Rats Rats. Cellulose content of 0% fiber diet.43) 0.7o 0. Note: An asterisk indicates that the value is significantly different from the fiber-free or low-fiber control group (p < 0.78 9.0) 2.97 (2.05).0 67.1.50)d 2. Net protein ratio (NPR).4 ± 4.8 9.4 75.0m(7.7i 0.8 9.1.67)l 10.3* 74.1* 79.0(7.1i 6.7 9.5 83. May 6. B. Nutr. Am.. Med.. J. . D.. J. and King. D... 1979. 20. The effect of meat protein and dietary fiber on colonic function and metabolism. J. A. Nutr. J.. O. 51. feed efficiency and protein utilization. 14. V. 592. P. 4. and Arthur. 1981.. A. M. O. C. 9. C. Effects of fiber on digestibility and transit time in dogs. 31. Cereal Foods World. 1965. M. Nutr. Harmuth-Hoene.. Kaneko. Growth performance and intestinal transit time of rats fed purified and natural dietary fibers. J. and Yaphe. Bacterial metabolites in feces and urine. Atallah. and Fox. 2094. Fawley. V. Nutr. Br. J. Nutritional Studies No. 1981. 191. Nutr. W. Heymsfield. H. 1979. C. 249. Kies. Am. 1978.. and Bach Knudsen. 24. Metab. 21. J. Effect of pectin structure on protein utilization by growing rats. M. J. S. 47. Farrell. 13. C. 56. F. Aust.. E.. 11. and Akrabawi. 31. 2001 6:57 PM 158 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Kelsay. 305. 8. 54. Food Chem. Delorme. C. Biochem. 17. 22. Burrows. Am. S.. 1984.. C... 1522... 108. 1983. J. Method of addition of cellulose to experimental diets and its effect on rat growth and protein utilization... number of defecations. 32.. M. fecal weight. 1978... Osa. K. Clin. Kelsay.. A. 20. Nutrient utilization by human subjects consuming fruits and vegetables as sources of fiber. 1274. K. and Prather. R. A. C. Nutr.. W. 1978. Effect of indigestible polysaccharides on protein digestibility and nitrogen retention in growing rates. 56. M. 32. A. 1982. Nutr. J. CRC Crit.. S. Am. E. fat and calcium digestibilities and fecal cholesterol excretion. B. B. 15. 5.. Hove. Branch. and Breidenstein. J. O. 741.. 109. Effect of fiber on protein. J. Food and Agriculture Organization. K. urinary excretions of energy and nitrogen and apparent digestibilities of energy. Hill. Nutr. 1978. Clark. Behail. W. Nishida. Agric. C. Metab. J. 29. 12. D. Donangelo. L. Carrageenan as a dietary constituent for the rat: faecal excretion. Effects of dietary fiber on nutritional status of weanling mice. Bone. Clin... M. 1726... nitrogen and fat. Beames... J.. Der einfluss von guarmehl in der nahrung auf die stickstoffbilanz. S. Nutr. 7. J. Nutr. Enteral.. 1970. and Melnik. P. Yatsuda. 10. and Gordon. T. and Schwerdtfeger. J.. Galloway.. K. 22. 1978. 399. J. J. J. L. K. 23. L. A. 43... F. A. E. FAO. S... Rome.. 469.. 1979. J.. 2. M. 32. S. and Lee. Rein. G. S. L. Sci. Br. 2454. J. and Jenkins. P.. 12.. The effect of type of and level of protein fiber and starch on nitrogen excretion patterns in rats. R. Nutr.. Exp. E. Can. B. and Kies. 479. H. E. J. M. J. 23.. Girle. Nutr. L. 1984.. Nutr. Fleming. 1978. Fiber and protein nutritional status.... Clin.. E. Heller. 301.. Nutr. 1981. Effects of dietary fiber on the apparent digestibility of major food components and on blood lipids in men..... Nutr. Casper.. E.. Parenter. 113. nitrogen absorption and growth. Br.. and Eggum. and Eggum. The effect of protein quality and fibre level in the diet and microbial activity in the digestive tract on protein utilization and energy digestibility in rats. J. E. 6. 1985. Starch digestibility of foods: a nutritional perspective. Nutr. Protein and energy utilization in men given a rural Guatemalan diet and egg formulas with and without added oat bran. and Prather. R. 1982. Nutr. Wojcik. Vitaminol. Roongspisuthipong. M. S...2387_ch4.. Food Sci. W. Beames. 1979. J. 73. 16. Comparative digestibility of acid detergent fiber by laboratory albino and wild Polynesian rats. Jakubick. H. 317. Sci. and Delpeuch. Eggum. E. K. May 6. 111. B. Dreher. L. and Berry. Biol. B. 3RD EDITION REFERENCES 1. S. Rev.1_fm Page 158 Sunday. R. 112. and their contribution to energy requirement and cecal VFA in rats.. Wolstrup. J. den proteinstoff-wechsel and die transitzeit der nahrung in ratten. J. Herbst. Clin. Cereal Chem.. and Kretsch. Fiber supplementation of enteral formulas: effects on the bioavailability of major nutrients and gastrointestinal tolerance. 112. M. 2027. Bowel transit time. 23.. Effects of pectin and cellulose on growth. II. 1986.. W. 24. Nutr. 461. C. and Schelenz.. 1988. Kronfeld.. Banta.. 265.. 3.. J. Cummings. Evert. J. M. M. M. J. Hawkins.. Corms.. Harmuth-Hoene.. Garrison. 19. 1118. S. Keim. J. Amino Acid Content of Foods and Biological Data on Protein. I. 18. and Merritt. J. Effect of fiber in sorghum on nitrogen digestibility. D. and Koike.. 1149. 46. L. Comparative effects of wheat bran and barley husk on nutrient utilization in rats. T. C.. Effect of fiber from fruits and vegetables on metabolic responses of human subjects. Dreher. 34.. W. 1981. B. The influence of dietary fiber on fecal nitrogen excretion in man. 517.. 1986. J. 44.. 584.. and Bowland.. Davis. J. Peterson. and Payne. 41.. and Southgate. L. 367. Digestible energy in relation to food intake and nitrogen retention in the weanling rat. Nomani. K. H.. 1981. S. 1129. Am. M. J. 11. Dietet. 1956. Draser. and Marlett. Miyoshi. A. 71. G. Influence of dietary fiber on the growth and protein metabolism of the rat. G. J. 44. S. D. Nutr. 141A. Food and energy intake of rats fed diets varying in energy concentration and density. Effect of pectin and algin upon protein utilization. 1059. B.. 112... Spiller. Ward.. R. 31. R.. 745. M. K. The effects of amount and type of fibre on apparent digestibility. J. M. A. 38. 1979... 113. Miles... R... L. Rep. Metabolic responses to dietary supplements of bran... I. Welters. D. B. Viola. Br. Effect of graded dietary levels of plant fibers on fecal output in pig-tailed monkeys. Fresard.2387_ch4. 1979. J. 101.1_fm Page 159 Sunday.. Mahoney. T. and Gallaher. and Jacobs. P. 1069.. Br. D. 26. Brunsgaard. J. Br. H. Nutr. Nutr.. 47. 25. I.. and Eggum. Okuda. 30. J. E. T. 357. J. B. O. protein. L.. 833.. and Baird. 1980. E.. 48. Int. 29. Br. W. 1982. 1983. Hill... A. Sundaravalli. O. Stanogias. M. R.. E. 1971. 1971. Rep. 62. J. 101. 46.. N. T.. R. J. 33. D. Nutr. A. J. Soc. Daniel.. 45. M. Nutr. 1994.. Nyman. C. Wisker. Okuda. Int. L. fat and non-starch polysaccharides in mixed diets: comparative studies between man and the rat. 39.. 112. A. Nutr. Assoc.. R.. Schneeman. 116. P. digestibility of nutrients and energy in young rats. G.. and Pearce. N. 385. An experimental reassessment of the factors used in the calculation of the energy value of human diets. Weber. R. 32. S. Miller. Schneeman. B. P. The effects of a supplement of dietary fiber on fecal excretion by human subjects. The influence of protein concentration. 1970. T.. 2020. D. Branch. J... and Bradac. 47. 207. O. 41. The digestion of fibre by pigs. Nutr. Kelsay. 283. D. fat and nitrogen digestibilities. and Baumgardt. J. and Cummings. Effect of dietary fiber components on fecal nitrogen excretion and protein utilization in growing rats. 1973. J. 1075.. Nutr. 59.. and Narayana Rao. 37. 74.. Southgate. 131. Davies. Nutr. 1970. S. J. Ulnan. Nutr. Response to dietary wheat bran in the exocrine pancreas and intestine of rats. Attallah. A.. 57.. and Pellett. and Asp. Nutr. 50. B. J. H. 1961. J. 1976. M.. Richter. G.. O. Metabolism. L. 495. and Baumgardt. 1987. M. Stephen. and Mokady.. T.. Shah. 1. C.. E. Nutr. 233. 42. 513. Fashandi.. 538. and Banwell.. and Sunderavalli.. 23. Effects of brown rice on apparent digestibility and balance of nutrients in young mean on low protein diets. K. Effect of refined cellulose on apparent energy.... J.. J. K. Effect of dietary fiber on the metabolizable energy of human diets.. K. D.. A.. 1982. 1985.. Br. Nutr. Nutr. 313.. 471.. Peterson. S. Influence of level of energy demand on the ability of rats to compensate for diet dilution. Br. R. G. Berg. J. Digestibility of energy.. Calorie conversion factors. B. Nutr.. . and Durnin. and Koishi. S.. Bach Knudsen. N-G. 34. 38.. 658. M. The influence of the period of adaptation on the digestibility of diets containing different types of indigestible polysaccharides in rats. Am.. Gastroenterology... Narayana Rao. E. 110. Zimmerman. 1970. J. 40. Assoc. Inclusion of cellulose in calorierestricted diets. Nutr. 517.. Shurpalekar. 15. T. Nutr. 24. B. O.. Arginine utilization of young rats fed diets with simple versus complex carbohydrates. 1980. Dietet. 27. 2001 6:57 PM EFFECT OF DIETARY FIBER ON PROTEIN DIGESTIBILITY AND UTILIZATION 159 25.. J. W. O... 89. R. Nutr. R. C. Comparison of the effect of cell wall and hull fiber from canola and soybean on the bioavailability for rats of minerals. 110. M. 1979. May 6. Effects of vegetable diets on nitrogen metabolism in cirrhotic subjects. Feldheim. J. T. J. G. L.. J. Z.. and Fisher. J. D. Nutr. G. and Reichert. Slavin. 1982. 41. Nutr. 36. A. Southgate. Fermentation of dietary fiber components in the rat intestinal tract. Minco. Problems in the prediction of protein values of diets. 43. I. P. Changes in small intestinal digestive enzyme activity and bile acids with dietary cellulose in rats. K. A. Sci. nitrogen balance and rate of passage. and Eggum. E. 53.. Prynne. protein and lipid. 1988. 1995. J. K. Bach Knudsen. Br. Vitaminol. 35. D. R. J. D.. 28. Food Sci. 33. E. Nutr. F. J. 49. Extraneous cellulose: effect on protein utilization. J. Sibbald. W.. 1985... A.. P. V.. Proc. M. 118. and Wong. F. B. J. H.. and fibre and the biological value of protein in low. J. J. The apparent digestibility of energy. Diez. 75... Hornick.. 1997. Anim. 54. A. . Leenaars. P.. 687. P. 1995. L. digestibility and energy metabolism in broiler chickens.. 1238..2387_ch4.. R. nitrogen.. H. B. H. J. Jr. 44.. 917. Nutr.. Zhao. C. Br.. K. Patil.-Q. 91. digestibility and energy balance in rats housed in different thermal environments. L. G. L. 53. and Istasse. T. O.. May 6. S. Pettersson. The influence of sugar-beet fibre. M. Br. 57. Bach Knudsen. Merchen.. 3RD EDITION 51. 73. Cole. and Brent. 70. and Eggum. 52. M. E. J. Plant Foods Human Nutr.. Res. 1998. J.. 77. and Eggum. 56.. Baldwin. D. water consumption and plasma metabolites in healthy Beagle dogs. Soybean hulls as a dietary fiber source in dogs. P...-L. N. Hussein. Hornick.. and Moughan. Diez. Jr. Sci. J.. Baldwin. M. Sci. 1993. Jorgensen.. J. 58.. J. Murray. C. Vet. M. Res.. J. 1996. Br. Zhao.. A.1_fm Page 160 Sunday. Vet. Nutr. X. guar gum and inulin on nutrient digestibility. X.. 64. L. Am. and Istasse. 379. S... B.. The influence of dietary fibre on body composition. 55. 2001 6:57 PM 160 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.. Influence of a blend of fructo-oligosaccharides and sugar beet fiber on nutrient digestibility and plasma metabolite concentrations in healthy Beagles.. 1993. Nutr. visceral organ weight. Fahey. 187. 1999. Jørgensen.and high-fibre wheat breads. Van Eenaeme.. 127. Effects of increasing levels of sugar-beet pulp in broiler chicken diets on nutrient digestion and serum lipids.. J. R. O. and Razdan.. The influence of dietary fibre source and level on the development of the gastrointestinal tract. on the other hand. Phytate is a distinct. Other components of dietary fiber such as cellulose. identifiable. lignin. pectin. magnesium. stalks. The special exception to this may be iron. Phytate.1 This should make these essential divalent mineral elements more vulnerable to complexation.00+$1. and some hemicelluloses and the water-soluble pectin. and lignin have properties generally associated with fiber or roughage. Dietary fiber is also derived from many of these same plants as a part of the stems. fibers do not have sufficient binding characteristics to significantly alter the balance of any minerals. May 6. with the exception of adsorption of the minerals on the large cellulose structure. which has special complexing properties with phytate. phytate is a component of most human diets throughout the world. has strongly electrovalent phosphate acid groups that have stronger binding and complexing properties.2387_ch4. and quantifiable compound. In order to understand the effects of these various components of dietary fiber. these all consist of polysaccharides or poly-alcohols. if dietary phytate is to exhibit some effect.3 Therefore. Harland and Donald Oberleas Phytate is found in all plant seeds (both cereals and legumes). the greater compleximetric strength of phytate toward cations would favor phytate as having the greater effect on the homeostasis of these minerals. and leaves that support the growing plant.50 © 2001 by CRC Press LLC 161 . for this relatively small molecule there is a large capacity for the complexation of essential divalent cations. inositol is absorbable and is metabolized as glucose. many roots and tubers.4 phytate’s hydrolysis products. 2001 6:59 PM CHAPTER 4. Thus. The two major components are the water-insoluble cellulose. are absorbable. 0-8493-2387-8/01/$0. it is important to understand the physical and chemical properties of these various constituents that make up dietary fiber. The hydroxyl groups are weak binding and complexing sites and thus. and other hemicelluloses. gums. and zinc are recycled through the pancreas. There is also good evidence that calcium.2 Effects of Dietary Fiber and Phytate on the Homeostasis and Bioavailability of Minerals Barbara F.2_fm Page 161 Sunday. Also. and in fruits. Whereas phytate and inositol phosphates cannot be absorbed. the cellulose and lignin provide bulk without hydration. Independent of phosphate esters. hemicellulose. both phosphate and inositol. that effect must be limited to the gastrointestinal tract.2 All people who consume a well-balanced diet also consume a diet containing phytate. Though fiber would pass through this same region of the GI tract. gums. With the exception of lignin. but it is frequently considered a component of dietary fiber — particularly in cereal bran. Pectin and gums hydrate and imbibe water and thus tend to provide bulk and lubrication to the GI tract. 14 These observations established a definite relationship between phytate and zinc homeostasis. as in the case of non-essential metals such as cadmium and lead.0. This complexation effect may be positive.8 Phytate was not implicated in zinc deficiency until O’Dell and Savage9 demonstrated that. So little zinc was required to meet the needs of the animals.03 Ca 0. or negative. magnesium. 3RD EDITION The other characteristic chemical property of phytate is that it dissociates to a very large anion in acid solution and has a capability for complexing heavy metals.13 and humans. 14.5 pigs.5 This pH is ideal for maximum complexation and is also the approximate pH of the duodenum.6 0. and others. May 6.) ZINC HOMEOSTASIS Zinc deficiency was first described in rats by Todd et al.12 dogs. manganese. Most investigators continued to believe that the major effect of phytate was on the absorption of dietary zinc.0412 – 0. Yet zinc deficiency. 2001 6:59 PM 162 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. iron.11 rats.05 Mg 1/Atomic weight 0. r 2 = 0. Trace Elem.01 Pb 0. 1997. iron. (Source: Oberleas. 173.0034X (µmoles precipitated).2387_ch4. H.. In vitro studies have also shown that zinc complexes strongly with phytate at pH 6. it was thought that zinc deficiency was not possible with natural foodstuffs. and zinc have been studied in detail.2.2_fm Page 162 Sunday.5 More recent studies of several divalent cations in vitro demonstrated a reciprocal relationship between the atomic weights and the relative strengths of the phytate–cation complexes (Figure 4. Only calcium. The word "bioavailability" was coined in the early 1970s to describe this phenomenon. This was repeated in several species including rainbow trout.1 The correlation between atomic weight and the concentration of precipitates of each cation at a phytate-to-cation molar ratio of 1:2 at pH 6.00 0 20 40 60 80 100 µMoles Figure 4..7 This was accomplished with a diet that was chemically depleted of zinc.C. and Chan. with chickens. Several in vitro studies have been carried out that have helped to clarify the complexing characteristics with various mineral elements. With permission. with apparent adequate dietary zinc but containing plant seed protein. and to a lesser extent copper. phytic acid added to a casein-based (animal protein) diet produced the same deficiency symptoms as a soybean-based (plant seed protein) diet.2. was reported by Tucker and Salmon. Elec. D. and zinc. The correlation equation is atomic weight (Y ) = 0.1).92.02 Zn Cu 0.04 0. as in the case of essential metal elements such as calcium.10 Japanese quail. . 2. This theoretical model is represented in Figure 4. D. An equal number of animals were fed a nonphytate-containing animal protein diet. and most investigators utilize experimental diets in animal studies that contain dehydrated ovalbumin. again an animal protein deficient in zinc. With dietary zinc representing only 20 to 25% of the duodenal pool. (Source: Oberleas. Figure 4.2. a practical relationship of this type of diet to any culture or world population is nonexistent. Thus. May 6.2. 1996.2_fm Page 163 Sunday..4..18–20 Zinc is the only essential divalent trace element cation that is secreted via the pancreas. This parameter is more difficult to measure and has as yet to be adequately determined in humans.2. 62. The results of these studies are shown in Figure 4.2.15–17 Evidence in other species has continued to be developed.) . Inorg. From these results a theoretical model was developed. Biochem.21 Computer models developed to study human fluxes have either avoided or ignored this fact. Simply measuring the differential of fecal zinc excretion and comparing the ratio of radioactive excretion indicates that the major effect of phytate is not on the dietary pool. 2001 6:59 PM EFFECTS OF DIETARY FIBER AND PHYTATE ON HOMEOSTASIS/BIOAVAILABILITY OF MINERALS 163 Investigators have ignored the secretion of zinc via the pancreatic fluid in quantities as much as four times that likely to be consumed in foods each day. 231. This clearly shows that the effect of phytate on endogenous zinc is two to four times the effect on dietary zinc.26 Little effort has been made to study this zinc pool. With permission. No population group in the world consumes such a diet. J.2387_ch4. therefore. It is also obvious that much of this secreted zinc must be reabsorbed in order for zinc homeostasis to be sustained. any result must be assigned to the total duodenal zinc pool to become of any value.20 This model has been confirmed also in rats. However. as shown in Figure 4. it was possible to measure relative absorption of dietary zinc utilizing radioactive 65zinc. but rather on the duodenal pool. the effect is not dietary zinc bioavailability but zinc homeostasis. This was done using a rat model in which the endogenous pool was labeled with radioactive 65zinc (Oberleas20).2 Ratios of 65zinc excreted following intraperitoneal injection as affected by isolated soybean–casein diets and dietary calcium levels.22–25 Only recently were efforts made to understand the pancreatic pool and its importance in the maintenance of zinc homeostasis. Some of the rats were fed a phytate-containing diet using soybean protein as the native phytate source.3. 0.1 with absorption dependent upon the movement of water. 0.90 = 3. rate of water absorption. intestinal transit time. 0. The concentrations of secretory fluids in decreasing order are bile. source of magnesium. The kidney . and age. (Source: Oberleas.2 (Homeostasis) Fecal Excretion 0.80 = 3.) MAGNESIUM BIOAVAILABILITY In humans. saliva. Inorg. Secreted magnesium appears to be almost completely reabsorbed.. Magnesium is secreted into the GI tract related to the flux of water.50 = 2 0. and pancreatic fluid.15 mmol/L. With permission.50 = 2 0. 231.2_fm Page 164 Sunday. The efficiency of absorption ranges from 21 to 70%.05 mmol/L.70 mmol/L. This model assumes a daily dietary intake expressed as unity. 0.2.8 Figure 4.50 mmol/L. gastric fluid. 1996. D. May 6.20 = 0. Biochem. dietary calcium and phosphate. magnesium is absorbed in both the jejunum and the ileum. 3RD EDITION Stomach Dietary Zinc Pool Size = Unity (1) Phytate:Zinc > 10 Pancreas Pancreatic Endogenous Zinc Pool Size = 2-4 Duodenum Saliva + Stomach + Pancreas Size: Stomach = 1 Pancreas = 3 Total = 4 Absorption/Reabsorption Fractional Zinc Absorbed 0.3 A mathematical model for zinc homeostasis.4 0. 2001 6:59 PM 164 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.10 = 0. 62. and all other values are relative to the dietary intake.6 0. J. Some factors which influence magnesium absorption include dietary intake..2387_ch4. . Negretti de Bratter. Several reports of decreased magnesium absorption with human subjects have been reported using cellulose as the fiber source. C.2387_ch4. Drews et al. and Etienne.2_fm Page 165 Sunday. (Source: Oberleas. I.) is the primary site for magnesium reabsorption and regulation of homeostasis. V. and Kwun. P. Collery. John Libbey Eurotext. in Metal Ions in Biology and Medicine.27–31 and Kelsay et al. vol. Bratter. 2001 6:59 PM EFFECTS OF DIETARY FIBER AND PHYTATE ON HOMEOSTASIS/BIOAVAILABILITY OF MINERALS 165 Figure 4.. Eds. where it may be influenced by water reabsorption and parathormone. J. D. S.6 but magnesium serves well as a tertiary synergistic cation. Ten to 30% of plant cell magnesium is associated with chlorophyll. Considerable magnesium excretion may occur via sweat loss.33 reported no effect with adolescent male subjects.2. which could be a major contributor to the fiber effect.. L..4 Confirmation of zinc homeostasis with the rat as the experimental model.. Paris.. Nothing is known about the bioavailability of chlorophyll magnesium. Several studies with a rat model reported an adverse effect of dietary phytate .32 used pectin as the fiber source. May 6. utilizing either cellulose or pectin.. 1998. magnesium alone does not complex well. With permission. Much of the fecal magnesium appears to be unabsorbed dietary magnesium. With phytate. 140. 5. Khassanova. P. have been shown to alter copper absorption. magnesium. and by organic substances that may include both dietary fiber and phytate. 2001 6:59 PM 166 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. particularly the proximal colon.45 In another study. Thus. differentiating between a phytate effect and a fiber effect is most difficult. and xylose. fiber. those substances that tend to favor the absorption of iron by reducing ferric iron to ferrous can have a devastating effect on copper absorption. and some adsorption occurs in the case of fiber. under these circumstances.9 Truly. COPPER BIOAVAILABILITY In most species. but it could contribute to the synergistic co-precipitation of calcium. glucose.47 The extent of absorption is influenced by the amount and chemical form of the copper ingested.2387_ch4. that affected by other dietary divalent cations makes calcium vulnerable to phytate complexation and subsequent decreased absorption. There are many accessory dietary factors that need to be evaluated to determine the net effect of phytate on calcium absorption and bioavailability. There is little agreement on fiber having an effect on copper bioavailability. and zinc by phytate in the GI tract.39. The excretory products are very stable complexes which are not susceptible to being reabsorbed or altered by dietary fiber or phytate.52–54 and fructose or fructose-containing sugars. Bird et al. thus.6 .38. given responsibility for the synergistic secondary cation effect observed in practical animal experiments.55 The normal site of excretion of copper is via the bile. arginine. This is one reason for the variety of results that make simple conclusions questionable.43 and.42.33. Calcium absorption occurs from all segments of the small intestine. a true effect of fiber is difficult to ascertain.6. Thus.33.e. fructose. but erroneously. i. Soluble oligosaccharides such as inulin and fructooligosaccharides have been shown to enhance calcium and magnesium absorption quantifiable by increased femoral bone volume and mineral concentrations in rats. any effect of fiber or phytate on copper in the GI tract must be on dietary copper. Again.46 showed decreased large bowel calcium concentrations in pigs fed brown rice compared with boiled white rice. In adults. by the dietary level of other metal ions. The adsorption of an insoluble calcium salt onto a fiber matrix without the demonstration of a chemical complex formation is not an effect of fiber in reducing the absorption of that mineral.1 Calcium absorption is enhanced by lysine.38. copper is absorbed from the stomach and all portions of the small intestine with decreasing efficiency in descending order.39 CALCIUM BIOAVAILABILITY Calcium is normally the divalent mineral cation in greatest concentration in the diet. the synergism works both ways and.41 Phytate.2_fm Page 166 Sunday. May 6.. lactose.48–51 Copper is absorbed in the cupric state and is more easily oxidized and/or reduced than is iron. It is difficult to arrive at a uniform conclusion because most experiments used diets containing both phytate and dietary fiber. An insoluble calcium salt that coincidentally adsorbs upon a fiber matrix without chemical reaction with some fiber component would not have been absorbed if the diet were devoid of fiber.34–37 Two studies with human subjects reported a negative effect of dietary phytate on magnesium absorption.44 An important characteristic of dietary fiber or dietary fiber adjuncts in our diets is that they are fully or partially fermented in the large intestine where the lower pH fosters this activity. showing significantly greater calcium absorption from all sections of the lower bowel. dietary copper is poorly absorbed. and oxalate decrease the absorption of calcium by forming complexes.40 Calcium is also secreted into the duodenum at 30 to 40% of the dietary concentration. Ascorbic acid is renowned in this regard. sucrose. 3RD EDITION on magnesium absorption. Calcium does not complex as tightly with phytate as does copper or zinc. lead. but by virtue of its dietary concentration it is usually. 2387_ch4. It is this chemistry of ferric ions with phytate that describes the mechanism for the alteration of ferric iron availability. including the size of iron stores and particularly the extent of stores in enterocytes. However. investigators must withhold definitive conclusions. the effect of phytate on iron absorption may have already occurred in the stomach. Non-heme iron constitutes the remaining 85 to 90% of dietary iron in the developed world and maybe more in the developing countries. vulnerability toward reduced iron bioavailability.5. Heme represents approximately 10 to 15% of the total iron intake within the developed world. also in a rat model. The most active site of iron absorption is the duodenum and upper jejunum. ferric and ferrous irons form very stable. Since there is no recycling of iron via the duodenum or bile. is non-ionic and is absorbed intact and thus protected from the possible complexation by phytate. This was later confirmed by Davies and Nightingale using a rat model. May 6. with Mn+2 being the most stable. The dietary factors which affect the absorption and distribution of manganese have not been clearly identified. IRON BIOAVAILABILITY Several studies have indicated that phytate in the diet decreases the iron balances in human subjects. There are two chemical forms of dietary iron: heme and non-heme. Heme iron is present in hemoglobin from blood and myoglobin from muscle meat. Thus. this makes for considerable chemical and physiological difference. insoluble complexes. Davis et al.2.38. ferrous iron. the only logical mechanism is that iron merely gets first chance to complex and. where the acid conditions dissociate phytate from its various dietary complexes and provide the ideal environment for complexation with ferric iron. though not quite as insoluble as ferric iron. and thus. Only Davies and Nightingale37 have reported an effect of phytate on the absorption of manganese.68 More recently.72 Much of the excreted manganese may be reabsorbed in a hormonally regulated pattern. This is shown in Figure 4. 2001 6:59 PM EFFECTS OF DIETARY FIBER AND PHYTATE ON HOMEOSTASIS/BIOAVAILABILITY OF MINERALS 167 Very little interest has been shown regarding the effect of phytate or fiber directly on copper absorption or bioavailability.2_fm Page 167 Sunday.37 Human studies indicating an effect of phytate on copper absorption have been reported by Moser et al. by the presence of the heme complex. MANGANESE BIOAVAILABILITY Manganese may be found in several valence states from +1 to +7. Absorption has been reported as low as 1%69 to as high as 40%70 using rat models. starts precipitating at about pH 2.67 Non-heme iron is also found in two forms: ferrous and ferric iron. in vitro studies have been made of the complexation between phytate and ferric and ferrous ions independently. Interestingly.58–64 Some animal studies have also confirmed the effect of phytate on iron absorption to a greater37 or lesser65–67 extent. The main route of manganese excretion is via the bile. The other important characteristic of both ferric and ferrous complexes is that they remain fairly stable through pH 7. including its reaction with phytate.57 Without further information on details of the diets used in these studies. Iron absorption is dependent upon several factors. Ferric ion was declared the only ion that would complex with phytate in dilute acid conditions and thus was the basis for the first and many subsequent analytical methods for phytate. whereas other cation complexes are soluble in dilute acid. .56 reported that an isolated soybean protein diet fed to chickens decreased the bioavailability of copper. which also gives it some advantage over other essential cations for complexation. Heme iron. which are the sites of regulation of iron absorption. Though the chemical difference is one electron. the resultant complexation does not occur in the duodenum as with most cations but in the stomach.71 The homeostatic regulation of manganese in the body appears to be effected by a very efficient excretory process. 4 This is a fair conclusion for humans..78 This lack of bioavailability of phosphorus from phytate means that much of the dietary phosphorus is excreted in the feces.2 0. May 6. 558. G. Gibb.. With permission.2.8 0..76 pigs. B. in Metal Ions in Biology and Medicine.0 0.. Paris..2_fm Page 168 Sunday. also.75 Since phosphorus is absorbed as orthophosphate. 2001 6:59 PM 168 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Eds.0 1 2 3 4 PH Figure 4. P.0 0.. D. Phosphorus from phosphorus-rich chicken and pig manure has created another international problem of phosphorus pollution when applied to cropland as fertilizer . and Etienne. A. H.4 0. R.. 3RD EDITION Fraction Pptd 1.4 0. 2000. Vernet. but its non-availability to monogastric species was first shown by Common73 with young chicks as the model. C.0 1 2 3 4 5 6 7 5 6 7 PH Fraction Pptd 1.77 and recently with dogs by Schoenherr et al.5 Chemical precipitation of ferrous (top) and ferric (bottom) ions at various pHs at a 2:1 molar ratio of cations to phytate.2387_ch4. Finkelman.6 0. the conclusion was that chicks did not have the inherent ability to hydrolyze phytate.8 0. 6. Collery. This has been confirmed many times in several monogastric species reviewed by Nelson74 and more recently by Soares. Centano.38.6 0. J. J. This same conclusion has been shown for rats. John Libbey Eurotext.2 0..) At this time it is difficult to ascertain whether either phytate or dietary fiber could have a significant effect on manganese homeostasis. vol. (Source: Oberleas. PHOSPHORUS BIOAVAILABILITY Phytate is rich in phosphate. such as the pH of the GI tract (acidic in the stomach for iron and about pH 6 in the duodenum and upper jejunum). REFERENCES 1.. 2000. P. C. A. B.. and Salmon. Finkelman. the effect may be related to the relative solubility of the phosphate salts contained in the diet and the greater or lesser degree of adsorption of these salts on the dietary fiber components. 103. 304. Delaware. An added benefit from phytase is an enhancement of the bioavailability of copper. and particularly zinc. 613. J. 88..43 Here again. 1979. and Savage. H. and Harmon.. Collery. FASEB J.. 173.82 reported some decrease in phosphorus absorption. Oberleas.79. D. Nutr. 9.. 1955. Cragle.. Effect of phytic acid on zinc availability. 160. 1910. Biol. D. Fibre and mineral absorption. . B. Parakeratosis or zinc deficiency disease in the pig.2387_ch4.. vol.. 8. Fed. Med. S.80 Incorporating phytase into animal feeds can improve phytate phosphorus bioavailability by 40 to 50%. 1997. these compounds have weak binding characteristics. magnesium. H. and Etienne... However. and Chan. Exp. Proc.. Elec. 14.81 The effect of fiber (cellulose) on phosphorus bioavailability is less distinctive. 10. W. Muhrer. zinc.32. Cation complexation by phytate. 1934. L. produces outcomes that are less conclusive.. SUMMARY It is well established that phytate. A. Chem. combined with the secretion of calcium. 56. Oberleas. S. Grieti. Exptl. their reabsorption and thus homeostasis. A. create realistic metabolic deficits. E. Dynamics of mineral elements in the digestive tract of ruminants. Oberleas.. 2001 6:59 PM EFFECTS OF DIETARY FIBER AND PHYTATE ON HOMEOSTASIS/BIOAVAILABILITY OF MINERALS 169 in the Netherlands. John Libby and Sons.. 965. Maryland. H. J. whereas others indicated no differences. Centano. G. Use of Japanese quail for the study of zinc deficiency. 14. Virginia. 1960. and Morris. Soc. and Arkansas). R. are effective in improving the utilization of phytate phosphorus in both pigs and poultry.. the case is not as well defined. (North Carolina. G. N. Metal Ions in Biology and Medicine. Med. Leeds. and O’Dell. This has led to the development of microbial phytase products that. 11. 2. J. J. and Taiwan and in the U. 90. B. Harland.S. coupled with several dietary variables.. and other divalent trace minerals. Adsorption has a less reliable and less effective impact on the mineral elements and.. Todd.. London. 109. F. May 6. 6. L. Eds. 72. Schoenherr. Dietary metal-complexing agents and zinc availability in the rat. and Hart. J. W. Biol. Germany. John Libbey Eurotext. Nutr. D.. D.. L. Exp. when added to animal diets. G. The fibers provide bulk. is capable of combining with essential divalent cations. Davidson..-C..10. B. Stephens. and Harrison.. Proc. 146. B. Biol. Soc. Tucker. 5. 1985. 2000. decreasing their absorption or. R.. J.. Digestion and absorption of phytate in mature dogs. Only Godara et al. 558. Soc. F. Vernet. Ed. E. 7. M. 1964.. Proc. M. C. Biol. K. W. 6. E. Med. 116. Zinc in the nutrition of the rat. R.. R. R. O’Dell. R. vol. 1. 256. Ferrous and ferric ions with phytate in vitro. in Fibre Perspectives... 3. 107. With respect to dietary fiber.. in the case of zinc and possibly magnesium. Elvehjem. D. Trace Elem. E. H. Gibb. Wedekind. 1973. J... and pectin and gums provide osmotic potential.2_fm Page 169 Sunday.. 4.. Paris.. and thus much of their effect is limited to the adsorption of the mineral elements. Influence of dietary zinc on cataracts in rainbow trout (Salmo gairdneri ). B.. Proc. 32. Fox. Ketola. with its strong complexing bonds. The environmental conditions.. where chicken and pig production is high and the land available for spreading the manure is limited. 1966. J. Prog. Oberleas. J. 83. 1.. and Hurley. 113. J. E.. 65. B. Matel. 15. Pekas. Physiol. J. M. S. Anim.. P. J. Nutr. 407.. 57.. Am. A.. Robertson. J. and whole-body retention of zinc. L.. A. H. May 6.. L. Physiol.. American Chemical Society. 186.. P. 85... Inglett. 1979. and Burns. Clin. I. D. 34. E. Clin. 29. 30. J. 36.. copper. Wastney.. Sci. B. C. Woodhouse. Borg. Syndrome of iron deficiency anemia. P. 231. 31. Toffolo. Likuski. 1876. J. 13. M. 140. N. 1412. Morogianni. Eur. S. 34. Am. I. J. 1956. M.. Urol. 1961. A.. S. and King. Exp. Excretion of radioactive zinc (Zn-65) in bile. Effects of calcium and phytic acid on the utilization of dietary zinc. John Libbey Eurotext. 342. Montgomery. Physiol. V. . B. 33.. and Prather. Inorg.. Zinc metabolism and the zinc deficiency syndrome in the dog. 1810. 1975. L. M. 1997. M. Eds... iron and manganese in rats. 107. IV. Paris. 377. G. S. J... M. J. B.. 49. and Marlett. 243. R. Roehl. 2000. Med. 25. 294. McHale.. D. Rumble. 1980. C. Stone. 22.. Influence of refined cellulose on human bowel function and calcium and magnesium balances... R. 523. M. 1165. Marlett. C. Oberleas. hepatosplenomegaly... G. M. zinc and magnesium utilization in adolescent boys. in Nutritional Bioavailability of Zinc. zinc and phosphorus. Washington.. 823.. 230. Nutr. 14.. J. 1975. Am. Nutr. M. Athens. C. Effects of phytic acid on zinc availability and parakeratosis in swine. Woodhouse.. 1963.. Effect of cellulose phosphate on calcium and magnesium homeostasis: studies in normal subjects and patients with latent hypoparathyroidism.. and Fox.. Effect of fiber from fruits and vegetables on metabolic responses of human subjects. 1962. 20. Calcium and magnesium nutritional status of adolescent humans fed cellulose or hemicellulose supplements. A. 27. 997. J. Greece. I.. J. 37. Cobelli. Vet. 16. and Nadimi. 24. Seelig.. H. 1979. 5. Muhrer.. L.. Matel. Ismail-Beigi. S.2387_ch4. M. and Kwun. J. Am. Lowe. S. A compartmental model of zinc metabolism in healthy women using oral and intravenous stable isotope tracers. and Henkin. Nutr. 359. G.. Ed. 33. In search of a human model for zinc homeostasis. The elimination of administered zinc in pancreatic juice.. and Yudkin. C. pancreatic and duodenal secretions of the dog.. J. Am. G. Am. Prasad. C. Med. Oberleas. Lowe. and Fox. Bratter.. 23. Clin. 1893. Kies. R. Mechanism of zinc homeostasis. L. A. F. L.. Cotzias. 32.. J. Collery. Reinhold. M. Drews. S. 28. Eds. Negretti de Bratter. J. L.. C.. H.. Kies... 243.. and Abadi. Am.. Khassanova. Slavin. D. and Etienne. R. Zinc-65 metabolism: gastrointestinal secretion by the pig. Comparison of estimates of zinc absorption in humans by using 4 stable isotopic tracer methods and compartmental analysis. H. 1979. J.. Nature. J. Mineral utilization in the rat. and Henkin... D.. 211. A. 62. in Proceedings of the 2nd International Symposium on Trace Elements in Human: New Perspectives.. Saccomani.E. 24. and King. hypogonadism. Clin. S. J. The effects of phytate on intestinal absorption and secretion of zinc.. Aamodt.. J. Sci. M.. N. D. Sci. 17. 12. 3RD EDITION 12. W.. Schneeman. 2001 6:59 PM 170 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. by man. P.. M. Oberleas. 532. L. Davies. Effects of dietary fiber on copper.. T. 44. Lonnerdal. E. J. Revusova. Am.C. 202. L. 1983.2_fm Page 170 Sunday. Effects of cellulose added to diets of low and high fiber content upon metabolism of calcium.. 19. Zinc absorption in humans. G. Kelsay. Med. I. Nutr. D. and Richards. 1960. Nutr. Ermidou-Pollet.. J. 1983. Confirmation of the mechanism of zinc homeostasis... 31. Nutr. Shames. Birnstingl.. Am. Clin.. E. J. 71. M. Res. and O’Dell. B. 1977. Requirement of magnesium by the normal adult. 1964. 35. R. 18. J. Dietary phytate as a possible cause of magnesium deficiency. S. Nutr. 21. 651–659. M. Am. and Selleck. Roberts. L. J. S. M. S. Sheline.. 26.. N. M. V. Biochem.. V. 2000. J. J. L... 21. K. 1975.. 1962. and Nightingale... R. and Forbes.. R. and Pollet. J. 151. Nutr... the mouse. Br. 1998. Parfitt. T. 1965.. 185.. Food Sci.. dwarfism and geophagia. M. Clin. M. 32. and Chaikoff. J. 1966. 510. magnesium. 1932. Halsted. 1996. Faraji. Sodium cellulose phosphate-induced increment in urinary calcium/magnesium ratio. 1943. Am. Clin. Zinc and copper in rat bile and pancreatic fluid: effects of surgery. O.. 61. J.. Food Nutr. D. F. vol. 1988. Specificity of zinc pathway through the body: turnover of 65-Zn in. J. Behall... 32.. C. Development and application of a model for zinc metabolism in humans. 78.. Metal Ions in Biology and Medicine. R. Mol. B... Nutr. C. 14. duodenal juice and bile of the dog as measured by its radioactive isotope 65Zn. G... in Mineral Metabolism: An Advanced Treatise. D. E. 48. Yee. A. T. A. and Bronner. Marsono. E. W. Usiteyra. vol.. 43. Fructooligosaccharide consumption enhances femoral bone volume and mineral concentrations in rats. Nutr.. J. 1950.. B... M... Y. 115. 1980. S. T. Solomons. Zn65. C. L. Exp. 23. M. Dietary fiber and calcium excretion in diabetes. Nutr. 1st ed. Biol.. P. J. S. and Finch. and Topping. A. IIA. 101. and Rosenberg. Br. Reynolds. 175.. 1988.. L. Med..... W. 493. Acharya. Nutr.. Nutr. Jr. 185. Comar. 42. Phytic acid and iron absorption. 58.. R. Hunt. zinc. T.. 1984. Iron absorption. R. J. and Mitchell. Biochem. . 1948. J.. 62. and Kratzer. H. J. Nutr. T. R. Ohta.. W. 341.. J. A. Yamada. Biological Roles of Copper. Copper. Clin. Mineral metabolism of healthy adults on white and brown bread dietaries. and Lewis. S. Lockhart. New York. J. Reiser. 452. D. 106. and Irving. Schultze. J. Derman. and Nightingale. Bezwoda.. A. Further studies on the availability of copper from various sources as a supplement to iron in hemoglobin formation. 385. 433.. R. Simpson. 44. 1965.. Absorption. W. 87. Studies on the mechanism of copper absorption in the chick. C. Hill. A. reserpine. phytates and polyphenols on the absorption of iron from vegetables. Edgecombe. 67. C. 34... and Hart. C. G. 44... 51. Carlton. Ferretti. F... F. and Hasselbad. K.. Sandberg. Takahara. Coarse brown rice increases fecal and large bowel short-chain fatty acids and starch but lowers calcium in the large bowel of pigs. Gillooly.. 41.... J. 281.... 1964. J... B. 831.. 49.. M. F. and Henderson. 60. 86. Moser. J. C. C. R. J. 1981. R. Med. 49. Am. II. Eugene.. 53. and distribution of copper.. 1936. 46.... Studies in chickens fed a copper-deficient diet supplemented with ascorbic acid. 321. Invest. 729. McCance. Amsterdam. 59. R. J.. C. and diethylstilbesterol. Dynamics and function of calcium. Am.. J. H. 207. W. R. Howard. Liss. D. A. 77. Turnbull. Nutr. Mo99. Nutr. L. 42. The severity of copper deficiency in rats is determined by the type of dietary carbohydrate. P.. C. and Charlton... 271. 39. M.. P. The effect of phytate and other food factors on iron absorption. and Widdowson. Calcium. 52. Physiol. J.. Proc. A. M. Aro. C. 47... P. M. 1962.. Faraji. 41. The inhibitory effect of bran on iron absorption in man. J. R. Biol. Bird. 45. M. 1964. 40. 1976. A. Owen. C. W.2_fm Page 171 Sunday. Harris.. 2000. Norris. B. 54. Clin.. Nutr. and Sasa. Absorption of Cu64.... K.. 48. Clin. and Sitonen. in Absorption and Malabsorption of Mineral Nutrients. A.... 130. and Carlton. I. P. N. J. N. N.. Hasselbad. Nutr. R.. Cleton. 1980. The effect of wheat bran on the absorption of minerals in the small intestine. J. 47. Torrance. M. T. Physiol. 55.. 1984. B. M. Interference of soybean proteins with the utilization of trace minerals. Eds. A... D. Nutr. I. B. W. Excerpta Medica. H. Bronner. 331. F. Ciba Foundation Symposium 79. F. 1965. W. E. 1983.. D. F.. transport. Nutr.. Sharpe. 120. 453. A. Fields. L. J... with special reference to calcium. 87. W. May 6... M. M. Elvehjem. M.. Korhonen.. Gooden. Bremner. 57. Decreased absorption of calcium. M. 50... Reinhold. 1965. 85. S. B. Peacock. B. Fox.. and Smith. W.. 1203. 2000. Weiser. R. Sano. Morohashi. 1897... VI. 1469. R.. 39... J... 2001 6:59 PM EFFECTS OF DIETARY FIBER AND PHYTATE ON HOMEOSTASIS/BIOAVAILABILITY OF MINERALS 171 38. Lancet. Hayakawa. R. J.. 530. Davis. O. 1962. Bothwell.. E. 1780.. WacPhail. J. 1st ed. Correll. Record. and Ismail-Beigi... and selenium dietary intake and status of Nepalese lactating women and their breast-fed infants. L. McCance. C. Academic Press.. H.. Nutr. and Widdowson. magnesium. Starcher.. 130. J. J. P. M. Absorption and excretion of 64Cu-labeled copper by the rat. S. W. Walker. 41. Andon. Nutr. 1982. S. H. and Starcher. H. A. 126.-S.. Absorption of hemoglobin iron... J. 1943. Abadi. J. E.. Jr. C. Am. The effects of organic acids. 97. D. iron. A. R. 217. Cardiovascular lesions associated with experimental copper deficiency in the rabbit. 56. O. 1942.. and Fe59 from ligated segments of the rat gastrointestinal tract. Van Campen. R. 1965. 61. I... P. R. S. Soc.2387_ch4. The effect of bread rich in phytate phosphorus on the metabolism of certain mineral salts.. 1969. New York. zinc and phosphorus by humans due to increased fiber and phosphorus consumption as wheat bread. 1792. Effect of reducing agents on copper deficiency in the chick. Chem. Cooke. Br. Eds. G. Nutr. and Cook. Alan R. Mills. J... and Puyat. J. Physiol. Phytic acid and mineral metabolism in poultry. Poult. 1992. Effect of dietary calcium. R. Nelson. 90. S. Miller. G.. J. R. Factors affecting absorption and excretion of 54Mn in rats. 147. Cu.2387_ch4. 1995. Phosphorus bioavailability. 1.. C. J. A. J.. May 6. 1981. Rev. and Zn and parameters of Zn status in piglets. Nutr. J. J. R. Parker. T.. R.. II.. Common. and Cotzias. Sci. D.. 422... M. particularly by way of the bile. Cromwell. 68. Soares.. 3RD EDITION 63. Mn. 1987. D. H.D. 75. and Cutherbertson.. 34. Influence of dietary calcium and phosphorus levels and growth rate on serum minerals. 82.. Cowan. A... 1939.. Chem. J. 51. S. Godara. Nutr. Esfahani. of iron. R. I. J. I... Anim. 66. 1974. and Steenbock. J. Lahimgarzadeh. Effect of cellulose incorporation in a low fiber diet on fecal excretion and serum levels of calcium. Biol. 65. T. Anim. Anim. P. V. University of MissouriColumbia. gilts and boars. phosphorus and iron in adolescent girls. S. Bertinchamps.. Heubner. Stahly.. Nasr.. New York. Howes. and Skanberg. Loewe... 1083. Greenburg.. 1943. 69. Kaur. Clin. Salji. D. and Johnson. K.. W. T. H. Fuhr. 141. 1988. J. in Bioavailability of Nutrients for Animals. 140. 79. Effects of purified phytate and phytaterich bread upon metabolism of zinc. 1973. Biochem. W. 323. 423. Sci... Cereal Chem. and Lewis. G. J. The distribution and excretion. 143. T. A.. 1967. 1st ed. 283. Coffey. H. Am. B. H. Effect of microbial phytase supplementation to a maizesoya diet on the apparent absorption of Mg. D. and vitamin D on the utilization of iron. G. 1964. dissertation. 749. Academic Press. 71. M. Sci.. E.. 49. 1943. Oberleas.. Kornegay. M. 1. 1.. L. 1966. H. 1971. Ph. Monegue.. 211. J.. J. Efficacy of a recombinant-derived phytase in improving the bioavailability of phosphous in corn-soybean meal diets for pigs. Baker. C.. J. .. Physiol. Brune. Chem. H.. 257. V. L. Efficacy of phytase in improving the bioavailability of phosphorus in soybean meal and corn-soybean meal diets for pigs. J. and Dyer.. S. 64.... 862. soundness scores and bone development in barrows. B. J. and Randolph. D. Ann.2_fm Page 172 Sunday. R. Fe.. 1914.. Utilization of phytate phosphorus by poultry: a review.. Z. Bioavailability of dietary iron in man. Rossander. 67. 76. 68. Nature. Hallberg.. 46. Ranhotra. 45. 1993.. H. Nutr. Anim. and Rimbach. Phytates and the inhibitory effect of bran on iron absorption in man. Anim. J. 1831.. Sci. J. H. J. Nutr. Iron absorption in man: Ascorbic acid and dose-dependent inhibition by phytate. Interdependence of routes excreting manganese. phosphorus. J. Uber eine Titration-methode zur Bestimmung des Phytins. J. 147. Eds.. G. Am.. L. Anim. and Hedayati. 81. P. 1981. H. 72.. A. 217. Effect of phytic acid on the availability of iron. 1049. 70. 64. 67. A. Cromwell.. 123. J.. Dietary Factors Affecting Zinc Availability. 1995. Effect of phytate on iron absorption in the rat. M. calcium. I. and Stadler.. J. D. L. 74. Sci. 118.. 77.. J. 379. Y. 1966. B. Am. Lee. Biol. Lancet. Diet and supplemental mineral effects on manganese metabolism in newborn calves. 78. 52.. Coffey.. and Rossander.. R. A. Pallauf.. 2001 6:59 PM 172 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 73. 71. and Bhat.... Hallberg. Clin. M. Effect of dietary phytic acid on the availability of iron and phosphorus. 1981. Am. Monegue. and Thomas. D.. L. 988. S. Ammerman. 1989.. H. Phosphorus in swine. 80.. C. H. 1509.. G. A. Clin. 32. Hallberg. L. 2000. phosphorus and nitrogen in man. and Randolph. G. Nutr. D. A. G.. Nutr. Copp... P. and Azzam.. Nutr. 59. E. L. Reinhold. D. E. and manganese. L. cobalt. 73. Hohler. dissolved in nutritional fat.8 Hydrolysis takes place in the intestinal lumen during solubilization of the vitamin esters in the mixed micelles. Polish investigators12 have shown that DF. Esters of fat-soluble vitamins. the question arises whether dietary fiber substances (DF) affect ingested fat-soluble vitamins being utilized in the GI tract.3 Effects of Dietary Fiber on Vitamin Metabolism Heinrich Kasper ABSORPTION OF VITAMINS Fat-Soluble Vitamins Starting from the fact that fecal fat excretion is increased after a high-fiber diet. DF binds components of mixed micelles.3_fm Page 173 Sunday. Vitamin A and Carotene Experiments on Test Animals Experiments on rats have shown that there is no decrease in vitamin A accumulation in the liver when vitamin A or carotene and 3% of pectin are added to the feed. disorders of fat absorption are invariably always associated with a disorder of the absorption of fat-soluble vitamins. The exact mechanism by which the intake of DF induces fecal fat excretion is not known. the authors believe that this DF substance does not affect the common steps of vitamin A and cholesterin absorption. Even the addition of 5 or 10% of microcrystalline cellulose had no effects on the postprandial serum vitamin A concentrations in rats when radioactively labeled vitamin A was administered orally. May 6.. are hydrolyzed in the intestinal lumen by pancreatic carboxylic ester hydrolases.9 Across the unstirred water layer the fatsoluble nutrients have to be moved toward the cell membrane in the form of mixed micelles. are contradictory. alfalfa. Interactions between DF and bile salts or the formation of micelles are discussed.00+$1. fatty acids. especially lipase activity.10 Due to the different concentrations of vitamin A and cholesterin when pectin is administered.50 © 2001 by CRC Press LLC 173 .7. 2001 7:02 PM CHAPTER 4. whereas the same amount of pectin has a hypocholesterinemic effect when cholesterin is added to the diet. as is the impaired activity of intestinal enzymes. and may therefore impair fat absorption in the upper jejunum. according 0-8493-2387-8/01/$0.2 Reports on the way different DFs influence pancreatic enzyme activity.2387_ch4. such as bile salts. monolene.3–6 Due to the close relationship between the absorption of triglycerides and fat-soluble vitamins. etc.1 Reduced lymphatic absorption of both cholesterol and triglycerides in rats receiving pectin. phospholipids.11 Under these experimental conditions there were no indications of DF influencing vitamin A or carotene absorption. and cellulose also supports this hypothesis. Experiments on Man Among the fat-soluble vitamins. carob seed flour. reported that carotene excretion after a high-fiber diet was approximately double that of a low-fiber diet. can have a positive as well as a negative effect on vitamin A absorption. lycopene.059). there was an increase in vitamin A absorption subsequent to the above-mentioned DF being administered. guar.3_fm Page 174 Sunday. May 6. The addition of wheat bran to a . agar.16 Wahal and co-workers19 tested the effect of wheat bran on serum vitamin A levels in healthy subjects during a 6-week trial.05) reduced by the water-soluble fibers pectin.14 using other experimental conditions. a mixed diet containing fiber from fruits and vegetables was compared with the same diet. All tested fibers significantly reduced the AUC24h of lycopene and lutein by 40–74% (P < 0. pectin.17 Single tests are inappropriate to determine whether or not long-term administration of large amounts of DF leads to the concentration of an essential serum nutrient or impairment of the supply required. indicated by accumulation in the liver and plasma concentration. The three well-known long-term studies on man on serum vitamin A concentrations subsequent to increased ingestion of DF come to different conclusions. Six healthy young women received an antioxidant mixture consisting of β-carotene. the effects of different kinds of dietary fiber on the absorption of carotenoids were investigated. it was found that serum vitamin A concentrations were higher compared to the initial figures.18 whereas in another experiment on 4 subjects after the daily intake of 15 g apple pectin or 30 g wheat bran over a period of 50 days. Carotene was added to the low-fiber diet to make the two diets as equivalent as possible in all respects except fiber. Mahle and Patton13 reported that subsequent to 8 g of a hydrophilic mucilloid laxative extracted from psyllium being administered daily.. had no negative effect on a vitamin A tolerance test in healthy subjects. Rock 1997.000 IU vitamin A palmitate were added to the meal. The mean AUC24h of β-carotene was significantly (P < 0. 3RD EDITION to individual test conditions. vitamin A has been the most extensively studied in man. guar.15 g · kg body weight–1). alginate. Because detailed information on the effect of different kinds of dietary fiber on carotenoid absorption in humans is lacking. During an experiment on 68 patients receiving two tablespoons of bran per day over a period of at least 6 months. microcrystalline cellulose. and alginate with a mean decrease of 33–43%. In healthy subjects. 2001 7:02 PM 174 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. was found when the daily vitamin A intake with feed was 30 or 90 IU and 10% methylcellulose. It is concluded that the biovailability of β-carotene. in which fruit and vegetable juices replaced the fruits and vegetables. The meal did not contain additional dietary fiber or was enriched with pectin. and the areas-under-curves (AUC24h) were calculated. In patients suffering from steatorrhea in the course of an excretory pancreatic insufficiency in chronic pancreatitis (mean daily fecal fat excretion 38 g). pectin. and canthaxanthin together with a standard meal. Increased vitamin A absorption in rats. Compared with the area under the postprandial serum concentration curve. 1998). The increases in plasma carotenoid concentrations were followed over 24 h. lycopene. cellulose or wheat bran (0.2387_ch4. With the vitamin A dose increased to 300 IU/day. Besides fat intake and the efficiency of extraction from the food matrix. In contrast to this. administered together with a formula diet.. whereas lignin had no significant effect.05).15 Barnard and Heaton16 stress the point that in healthy subjects cholestyramine added to a test meal reduced vitamin A absorption. the additional ingestion of 10 g pectin together with a standard meal had no effect on the serum vitamin A concentrations when 300. and guar.58 Wheat bran. neither the fecal excretion of vitamin A and carotene was increased nor were plasma vitamin A and carotene concentrations affected by long-term intake of hyrophilic mucilloid. the amount and type of dietary fiber in the diet seem to determine carotenoid bioavailability (Parker 1997. lutein. or wheat bran was added to the feed. the same amount of DF had a negative effect on vitamin A absorption. while serum carotene concentrations remained unchanged. a decrease in vitamin A concentrations was found. Kelsay et al. Williams et al. The dietary fiber effect on the AUC24h of canthaxanthin was almost significant (P = 0. and lutein given within a mixed supplement is markedly reduced by different kinds of dietary fiber. 30 . or wheat bran (0. if given with diets marginally deficient in vitamin D3. May 6. These findings indicate that the above-mentioned carbohydrates have a negative effect on vitamin D3 absorption. pectin. 2001 7:02 PM EFFECTS OF DIETARY FIBER ON VITAMIN METABOLISM 175 standard diet with 20. The prothrombin and thromboplastin times of diabetics who were treated with a high-fiber diet (25 to 35 g dietary fiber per 1000 kcal) for an average of 21 months remained unchanged.29 Riedl et al. This could be demonstrated with the vitamin E tolerance test when the glucomannan was added to the test meal. This indicates that the high-fiber diet does not impair vitamin K absorption. cellulose. Vitamin K It is uncertain whether there are relations between vitamin K metabolism and ingestion of dietary fiber.21 Serum or urinary calcium and phosphate concentrations and the serum alkaline phosphatase concentrations. guar.2387_ch4. are currently being discussed. Lignin combines with bile acids and increases their excretion.000 units of vitamin A significantly lowered serum vitamin A levels within 1 week. They suggest that bran in the wheat flour which forms the staple diet in some parts of India may contribute towards the vitamin A deficiency state commonly observed in this country. especially its lignin component. in combination with a low vitamin D intake and bioavailability. indicate that under this therapy vitamin D–deficiency rickets may develop.15 g · kg body weight) did not affect the plasma response curves of α-tocopherol.26–28 Experiments in Man The glucomannan konjac mannan has a lowering effect on vitamin E absorption in healthy volunteers and in diabetic patients.20 Experiments on Man Several findings indicate that a high intake of fiber from unleavened whole meal wheat flat bread is responsible for rickets occurring in population groups in Asia. and guar led to decreased fat and nitrogen utilization and. alginate.22 Experimental high-fiber diets have been reported to reduce plasma half-life of 25 (OH) D.25 Vitamin E Experiments on Test Animals There have been several fiber and vitamin E studies conducted on rats. The significance of the rich phytate concentration and the role of wheat fiber. and this trend continued over 3 weeks.23 This finding. obtained during investigations on children treated with bran for constipation.24. may explain the high incidence of rickets and low plasma 25 (OH) D concentrations in some population groups and in infants on macrobiotic diets.3_fm Page 175 Sunday. Investigations on the influence of DF on vitamin K absorption and serum vitamin K concentration are not available.58 found that a meal enriched with pectin. Vitamin D Experiments on Test Animals Investigations on chickens revealed that carboxymethylcellulose. From these studies it appears that intakes of pectin decrease vitamin E bioavailability. led to the development of rickets. The authors postulate that vitamin D probably becomes attached to the fiber/bile acid complex and is transported unabsorbed through the gut. but fiber-containing breads or cereals have only transient or no effects. 31 Water-Soluble Vitamins The underlying mechanisms of water-soluble vitamins being absorbed are more complex and less uniform than those of the fat-soluble vitamins. 3RD EDITION From the fact that the concentration of coagulation factors is changed if the fat and dietary fiber content of the diet is altered. Whereas some investigators37 found similar plasma pyridoxal-5′-phosphate concentrations in man after the intake of white and whole wheat breads. and a combination of the two on the absorption of pharmacological doses of riboflavin (30 mg) was examined with a riboflavin load test. it must be inferred that coagulation factors are not sufficiently reliable parameters for judging vitamin K absorption.33 Pyridoxine (Vitamin B6) Experiments on Test Animals A number of experiments on animals and man using natural and purified forms of dietary fiber suggest that dietary fiber has little effect on the bioavailability of vitamin B6 in foods. The authors concluded that dietary fiber accelerates GI riboflavin absorption. pectin.37 It appears that the significantly higher consumption of dietary fiber by vegetarians has no adverse effect on the availability or metabolism of vitamin B6.36 There were no uniform results in experiments on the bioavailability of vitamin B6 in bread containing varying amounts of wheat bran. Biotin No data published.38. Riboflavin (Vitamin B2) The influence of high. No effect of wheat bran on riboflavin absorption was detected. This indicates that dietary pectin supplementation has no effect on the utilization of dietary vitamin B6. wheat bran. Thiamin (Vitamin B1). cellulose.39 . 2001 7:02 PM 176 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. pectin.32 In another study the impact of psyllium gum. wheat bran.2387_ch4. Compared to a control group receiving a low-fiber diet. The plasma vitamin B6 concentrations were slightly lower subsequent to whole grain bread being ingested. such as folic acid.34 Bioassays on rats and chickens receiving cellulose.3_fm Page 176 Sunday. coarse and fine bran. Pantothenic Acid. must be transformed in the intestinal lumen prior to absorption. May 6. and mixtures of cellulose.8 to 25. and lignin indicated no inhibitory effect of these test materials on the bioavailability of pyridoxine. there was no negative effect on plasma pyridoxal-5′phosphate concentration and urinary 4-pyridoxic-acid excretion.and low-fiber diets on urinary riboflavin excretion was examined in healthy subjects. and cabbage increased urinary excretion for 8 h when 15 mg riboflavin-5′-phosphate were administered orally with breakfast.35 Experiments on Man In man receiving 15 mg pectin daily. but psyllium gum reduced the 24-h apparent absorption of riboflavin from 31. Fractional urine collections were made for 24 h. others reported significantly higher fecal B6 excretion with whole grain bread than with white bread. Some water-soluble vitamins.4%. serum folic acid levels were normal. Folic Acid According to the experiments of Luther et al.40 DF can bind pteroylmonoglutamates but not polyglutamates. there are indications that conventional nutrition in Western industrialized countries does not invariably meet folic acid requirements. administered 10 to 40 g wheat bran daily.44 it cannot be denied that ingestion of bran or isolated DF leads to folic acid requirements being impaired. Effects of the two foods were qualitatively different.3_fm Page 177 Sunday. Experiments on healthy subjects showed that there is no change in folate absorption between the low-fiber and the high-fiber bread meals. whereas beans minimized the difference between PteGlu and PteGlu 7 serum areas. No effects of these DF on plasma or hepatic folate concentrations could be demonstrated in feeding experiments on chickens. 10 g pectin. . Serum radioactivity was significantly higher when microcrystalline cellulose was added. but the product might be bound in the presence of DF.17 These fluctuations of serum folic acid concentrations are probably due to different eating habits during the various seasons rather than to the ingestion of wheat bran. Relative folate absorption was determined by measuring 24-h urinary folate excretion and serum folate levels at 0. Addition of 30 g wheat bran accelerated PteGlu absorption. constipation.2387_ch4. and irritable colon yielded opposite results. 1. or 20 g wheat bran were added to the test meal. This would suggest that naturally occurring dietary polyglutamates could be split by the normal intestinal conjugase activity. serum folic acid concentration was significantly decreased. Vitamin PP) Sina et al. and 2 h after ingestion of a formula meal containing 1. Wheat bran increased the absorption of PteGlu relative to PteGlu7. Serum activity was measured subsequent to the labeled vitamin being administered orally with or without the addition of 5 or 10 g microcrystalline cellulose to the feed. whereas PteGlu7 absorption was not significantly affected by either food. Two 6-month studies on patients with diverticulosis.13 µmol PteGlu or PteGlu7 (500 µg PteGlu equivalent). sodium alginate.. there was a significantly higher average folic acid concentration at the end of the observation period than there was at the beginning. maximum postprandial serum folic acid concentrations were obtained subsequent to 200-mg folic acid monoglutamate being ingested together with a standardized test meal. Since. As studies on healthy subjects show. Despite the fact that insoluble dietary residue of various foods was used in this study. however. The areas under the folic acid concentration curves show that total absorption was significantly decreased up to 7 h after the intake when guar and bran were added to the test meal.42 In the patients of the other study.11 studied the influence of microcrystalline cellulose on the absorption of radioactively labeled nicotinic acid in rats. other investigators41 incubated folic acid with defined DF (cellulose.30 The results of the investigation show that supply of folic acid requirements is not impaired if a mixed high-fiber diet is administered like the one recommended for prophylaxis and therapy of metabolic and GI disorders. Results were negative when folic acid monoglutamate was incubated with high-fiber bread like that consumed in Iran. 2001 7:02 PM EFFECTS OF DIETARY FIBER ON VITAMIN METABOLISM 177 Nicotinic Acid (Niacin. The above-mentioned concentrations were obtained 1 to 2 h later than in the control experiment when 5 g guar.17 Keagy and co-workers45 have examined the effect of wheat bran or California small white beans in the diet on absorption of monoglutamyl (PteGlu) and heptaglutamyl folic acid (PteGlu7) in healthy men. May 6. lignin. pectin.43 After a 21-month treatment of diabetics with a high-fiber diet. In the patients of one study receiving 3 teaspoons of wheat bran daily. and wheat bran) without observing an absorptive effect. May 6. inositol. In B12 deficiencies.30 In healthy volunteers and in diabetic patients. and hence absorbed. urinary excretion increased significantly subsequent to cellulose and pectin being ingested. 3RD EDITION Cobalamin (Vitamin B12) Experiments on Test Animals Cullen and Oace46 have examined the effects of cellulose or pectin supplements upon vitamin B12 metabolism in rats. The authors suggest that both fibers interfere with the recovery of biliary vitamin B12 by binding to a residue that is little metabolized until it passes the ileum reabsorption site.48 ENTERAL VITAMIN SYNTHESIS Due to the findings of a large number of experiments on animals. The reason for this is not known.29 Ascorbic Acid (Vitamin C) With 100 mg ascorbic acid being administered daily to healthy subjects. a significant increase in urinary excretion of the vitamin was observed when 14 g hemicellulose was added to the diet. this occurred much earlier and to a greater extent with pectin than with cellulose. sorbitol. The experiments demonstrated that the addition of either 20 to 50% cellulose or 5 to 20% pectin to the semipurified B12-deficient diet enhanced the depletion of body stores of vitamin B12. it could be definitely demonstrated that those carbohydrates which are hard to digest stimulate the intestinal synthesis of B vitamins and vitamin K. Cellulose and pectin had no effect on urinary ascorbic acid concentration. these findings might indicate an increase in intestinal ascorbic acid absorption. vitamin B12 absorption was not affected when the glucomannan konjac mannan was added to a test meal. Another explanation would be a change in the intestinal flora. 2001 7:02 PM 178 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. rats resist dietary vitamin K deficiency by eating their own feces. and cellulose. mannitol. Prevention of coprophagy in these animals leads to vitamin K deficiency after a short time. vitamin K. It is only known that there is no change in serum vitamin B12 concentrations in patients with diverticulosis on 6-month wheat bran therapy. The authors postulate that there may be greater production of propionate during the bacterial decomposition of pectin. Preliminary results of other investigators confirm these findings. Since the colon is the site of vitamin synthesis.2387_ch4. . There have been various studies.42 The serum vitamin B12 concentration of diabetics 21 months subsequent to a high-fiber diet being administered was also in the normal range. However.49 For example. with a resulting increase in B12 consumption. Contrary to all expectations. Urinary methylmalonic acid excretion was also measured and used as a parameter of vitamin B12 deficiency. pectin. Experiments on Man The influence of DF on vitamin B12 has not been studied systematically hitherto. vitamins synthesized in the intestine can probably contribute to the supply of what is required only when coprophagy takes place.50 There are findings.3_fm Page 178 Sunday.47 Postprandial serum concentrations were not measured. methylmalonic acid is produced from propionic acid in the liver. They were able to show that 5 g pectin as well as 10 g wheat bran increase serum vitamin C concentration and urinary excretion significantly when they are ingested together with a formula diet containing 400 mg ascorbic acid. which indicate that vitamin K requirements even in man are met to a large extent by the intestinal flora. however. especially on raw potato starch. Compared to the control group.51 It is not known whether during possible absorption of internally synthesized vitamin K the content of nutritional DF in man determines the amount of synthesized. 1947. L. N. J. G. F. Rabast. 20. Il.. 10. Nutr.. Gastroenterology. K. Kishore. E. Bile acids and vitamin A absorption in man: the effects of two bile acid-binding agents. and Gilat. 44. K. 37.. D. H. Carotene and vitamin E metabolism in man: the excretion and plasma level as influenced by orally administered mineral oil and a hydrophilic mucilloid. on vitamin A utilization in rat. Schneeman. D.. Nutr... Levin. Clin. Gronowska-Senger.. A. 14. Fed. Fed. M. 1980.. Harries. Poziom blonnika w diecie a wykorzystanie witaminy. REFERENCES 1. Studies on the substrate specificity of a carboxyl esterhydrolase from human pancreatic juice. E. Gut. Action of cholesterol esters and lipid-soluble vitamin esters. 389. Sommer. iron.. J. 1980. P. 7.. and Jain.2387_ch4. Weizer.. Biol. 37. 147. H. Fed. Biol. J. B.. Lipid Res. 32. and Cassidy.. M. V. and Heaton. absorption. Nutr.. Soc. 1981.. Clin. 1970... W. Biophys. Biol. May 6. 9. Proc. D... N. 27. Peters.. Graff. and Prather.. A. Roy. 11. Exp. Kasper. and Schneeman. 3500... Proc.54 It is not known to what extent bacterial vitamins synthesize in man’s small intestine and thus. L. and Vohra. 46. P.. P. and Kasper. 15. 100.52 and a free intrinsic factor is often present in the lumen of the small intestine. a hypocholesterolemic polysaccharide.. 22. G. 1981. H. 553. Biochim.. E. B. B. Soc... VI. J. 3. Bustin.. 166. F. 1967. Physicians India. 6. H. Fed. B. Dunaif. 3. Singh.. Clin. 1976. L. J.. B. Tombes. Biochem. R. L. K.. 269. 829. J.. Clin. R.. and Gallo. O. nitrogen retention and fat absorption in chickens. A high-fiber diet does not cause mineral and nutrient deficiencies. and Rubin.. Fed. and transport. and Nylund. Exp.. Am.. Friedman. Am. 18. V. cholestyramine and lignin. Roczn. J.. Mahle. 755. Effect of pectin... Exp. and Muller.. 4. 1939. J. K. M. Story. 321. V. O. P. This might account for the absence of vitamin B12 deficiency in some groups of vegetarians. and Cantone. Biol. 14.53 there has been speculation that vitamin B12 synthesized by the small intestinal flora contributes to vitamin B12 requirements being met. S. 1981. 1973. Mathias. Lombardo. D. Am. H. T. 1847. G. III. J. Exp.. Fed. H. D. 1986. 33. 9. R. R. 12.. and Guy. E. 849A. L.. 1980. V. O. J. Am. Dietary fibers. 1978.. 1980. 1979. 12. Kasper. Proc. 16. Dietary fibers. 34. B. Behall.. Gastroenterol. R. 33. J. J. The effect of polysaccharides on energy utilization. R. Phillips. The effect of dietary fiber on the pancreatic excretory function. silicon and vitamin A balances of human subjects. and Gallaher.. 1979. 8. Am. Dutta. and Brien. Soc. J. L. H. R. Cassidy.... Sina. 289. 40A. Nutr.. magnesium.. K. Barnard. E. the requirement in vitamin B12 being met can be influenced by those carbohydrates being ingested which are difficult to absorb. 12. J.. 1978. A przez organizm.. Vahouny. Kratzer.3_fm Page 179 Sunday. 19. 2. 5. A. The effect of dietary fiber on the postprandial serum vitamin A concentration in man. 316. Binding of fatty acids and monolein from mixed micelles containing bile salts and lecithin... T. B. Vahouny. Exp. unpublished data. 1981. Provenghi. W. 1489. Chudy. M. H. Hepatogastroenterology.. M. D. 17.. 1981. Kritchevsky. A. Effects of several sources of dietary fiber on human pancreatic enzymes activity in vitro. 477. and Patton.. Gastroenterology. Pzh.... Rajaguru. S. Fassl.. and Fehle. 2182. Prakash. 2001 7:02 PM EFFECTS OF DIETARY FIBER ON VITAMIN METABOLISM 179 Since the human small intestine often harbors considerable microflora. Gujral.. Effect of fiber from fruits and vegetables on calcium. B. Acta. A. 30. Effects of chronic intake on cholesterol absorption and metabolism in the rat. Intestinal fat digestion. Gallo. M. Am. 13.. N. R. W. I.. L. Effect of high fibre intake on serum vitamin A levels. T.. U. and Schrezenmeir. 80A. Assoc. Studies on the in vivo absorption of micellar solutions of tocopherol and tocopheryl acetate in the rat. Loss of lipase activity with fiber treatment. Dietary cellulose and intestinal absorption of nutrients in rats. Rattan. Biol. Proc. and Smaczny. Wahal. 611. M. .. Soc. Maheshwari. J. Kelsay. P. M. 1108. Effect of dietary fiber on pancreatic enzymes and fat malabsorption in pancreatic insufficiency. Med. Poultry Sci. . F. M. Vitamin B6 status and bioavailability in vegetarian women.. 1. Nutr. and Peffer. and Leklem. R. Nutr.. H. 27. P. W. 1978.. M. Potential complications in the use of wheat bran of constipation in infancy. Nutr. Am.. 28. T.. F. J. van den Berg. C. Vergote. Tanaka.. Pediatr. Perez-Santiago.. 35. Chow. 1. A. T. Nutr. Z. 854. R. 1990. and Compston. 29. T. deLumen... 797. Ristow. 23.... Leklem. Int. Proc. 58. Gobio-Casali... Soc. 49.). J.. Shultz. L. Roe.. van den Berg. Soc. B. J.. 1976.. S. S. 1980. Gibson. T.. 640. G. W. and Damron. 38. 1990.J. 1980. S. A. B. Shane. D. 33... Effect of hard red spring wheat bran on the bioavailability of lipidvoluble vitamins and growth of rats fed for 56 days.. Med... S. and Hautvast. J. Am. 677. D.. V. and Cecchettin. 2. E. 32.. S. A.. 41. P. 43. W. Dingjan. Ala. 111. F. Doi. 141 (Suppl.. 3. A.. Wedel. 40.. Biol.. and Stevens. Influence of citrus pectin on the bioavailability of vitamin B6 in man.. Effects of selected polysaccharides on the bioavailability of pyridoxine in rat and chickens.. R... and Humphreys. P. 46. Biochemische und haematologische Ma β stäbe zur Beurteilung des Folatstatus beim Menschen.. D. Schrijver. B. The bioavailability of vitamin B6 in foods. 80.. K. F. 391. C. J. J. 1980. D. J. Proc. Urinary ascorbic acid excretion in the human as affected by dietary fiber and zinc. I. Lowik. Mineral and vitamin status on high-fiber diets: long-term studies of diabetic patients. Exp..2387_ch4. and Butterworth. A. J. deLumen. M. Clin. Methylmalonic acid and vitamin B12 excretion of rats consuming diets varying in cellulose and pectin. Res. Nutr.. P. M. Exp. 1987. O.3_fm Page 180 Sunday. Br. J. 1819. 36. Saccomani. J. Keagy. F. I.. Med. A. Br. R. 504. K. McLain. Nguyen. Nutr. Cullen.. J. J. Med.... Lancet. 1. 799. Reyes. 1982. 26. J. Luther. Wrick. E. J. C. Deganello. Batchelor. 2001 7:02 PM 180 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Simpson. A. Gregory. Reyes. 1983. Bindra. J. 22. Rickets in Asian immigrants... Am.. 45. Chiu. 202.. 39. 29.. 115. 1978.. H. L.. L. Reduced plasma half-life of radio-labelled 25-hydroxyvitamin D3 in subjects receiving a high-fibre diet. Ismail-Beig. 24. Am. E. Bioavailability of vitamin E in rats fed diets containing pectin. Metabolic effect of bran in patients with diverticular disease. P. J... E. Hulshof.. Kawara. 9. F. Am.. Astolfi... Miller. 25. Vit. 23. Ern. B. Sci. J. 3... M... 756. 1981. J.. 39. 263. Tohoku J. 110. Proc. Rev. Exp.. M. and Oace.. Gregory. Matsuura. 1987.. 73. Fed. Am. S. Nizan. 1. Kies. G. J. Reinhold. G. M. E. and Omaye. Am. J. Med. J.. 47. C. J. O. Fed. G. High prevalence of rickets in infants on macrobiotic diets. Clin.A. 1976.. 1982. and Baba. A. Nutr. G. Food Sci. Nutr... Am. Dietetic Assoc. F. Schultz. K. H. Folate binding by insoluble components of American and Puerto Rican diets. Ferguson. 40. J. Clin.. J. 1978. 42. Nutr. J. M. A.. 51. M. Bioavailability of vitamin B6 from wheat bread in humans. Effect of fiber supplements on the apparent absorption of pharmacological doses of riboflavin. 38. T. R. Schaus. 47. H.. 241. Omaye. L. D.. 34.. Nutr... Nutr. 1988. Pietrzik. and Chow. J. 1608. 44. and Reinhold. 88. H. 37.. T. P. 210. and Fox.. E. et al. 1981. and Omaye. Effects of dietary fiber sources on riboflavin absorption. J. and Ockhuizen. B. B. R.... Diabetes Care. G. and Oace. Nutr. Santini. H. Roe.. . J. R.. The vitamin D status of East Indian Punjabi immigrants to Canada. Gastroenterol. and Draper.. E. D. D. P. Folate content of Iranian breads and the effect of their fiber content on the intestinal absorption of folic acid.. H. Nutr. T. Fed. J.. 1132. van Staveren. and Hötzel. Urban.. Brewster. Nutr.. and van Soest. and Damron. Br. Biol. 1965.. Brodribb. 49.. 389. J... 46. J. 213. 2. Effect of dietary fiber on the vitamin B6 status among vegetarian and nonvegetarian elderly (Dutch Nutrition Surveillance System). 1976. 1988. Perera. Folate bioavailability in humans: effects of wheat bran and beans.. S. K. and Leklem. 1978. 1983. Fed. T.. 39. Brit. D.. Influence of dietary fiber (konjac mannan) on absorption of vitamin B12 and vitamin E. L. B. 31. Soc. I. T.. Bioavailability of vitamin B in rats fed graded levels of pectin. 30. Am. M. F. H.. 108. 447. R. F. J. Forsch.. G. R. D. The effect of dietary fiber on folic acid bioavailability. Zoppi. Coll. Effect of high-fibre diet on haemostatic variables in diabetes.. Anderson. K.... Am. 1982. J. 424. May 6. 31. A.. J.. Lubin.. L.. 1984. K. 1403. Kalkwarf. Keltz.. Fed. 91. 1167.. R. S. M. M. 1981. 48. J. Exp. F. Clin. E. 1985. D.. J. and Kirk. Clin. 3RD EDITION 21. C. Fed.. S. Russell. Biol. 2. and Mann. Dagnelie. L. Nutr. Miller. A. Gregory. R. Thieme-Verlag... 1966.. 24. Suppl. 62. C.. S. G.. injury and vitamin K deficiency. A. and Barnes. S. May 6. 68.. Albert. Vitamin B12 synthesis by human small intestinal bacteria. Rock. W.. 781.. Williams. Barnes. H. Nutr. Mathan.. H. M. R. Der Einfluss von Ballastoffen auf die Ausnutzung von Nährstoffen und Pharmaka. 49. 53.. and Wolfram. Kasper. Proc.. R. W. 51. Gastroenterology. Exp.. 52. J. 50.. J. 51. W. Ther. S. Mathan. 1959. J. Rev... and Erdman. Nutr. V. H. 218. Ed. Shantakumarie. 106. 38. J.. Effects of the prevention of coprophagy in the rat. T. C. D. R. 2170.. Parker. E. H. 603.. 1980. G. 58. Riedl. 70. and Baker. Free intrinsic factor in the small intestine in man. Carotenoids: biology and treatment. D. Nature... Soc.. and Fiala. Pharmacol. Kapadia. P. Stuttgart. and Bauer..2387_ch4.. 1980..... Bhat. 1998. S. Contributions of the intestinal microflora to the nutrition of the host. 2001 7:02 PM EFFECTS OF DIETARY FIBER ON VITAMIN METABOLISM 181 48. 1972. 1999. H. Boileau. Hoffmann. 11. C. 55. I.. Editorial: Intestinal microflora. . 115. Mathan. J. 75. R. Eur. 185. 57.. Rajan. 56. 129. 1976.. 1980. Biovailability of carotenoids. J. 54. Some dietary fibers reduce the absorption of carotenoids in women. Nutr. J. J. J. 1. L. S86... Gastroenterology. 1997. Horm. and Kapadia. Linseisen. 283. in Pflanzenfasern-Ballastoffe in der menschlichen Ernährungs Rottka. Factors influencing the uptake and absorption of carotenoids. Kwong. M. Med... Vit. Biol.3_fm Page 181 Sunday. V. Clin. 1997. 341. Nutr. J. J. J. Bacterial flora of the gastrointestinal tract in Southern Indian control subjects and patients with tropical sprue. Hötzel. 70. G. 3_fm Page 182 Sunday.2387_ch4. 2001 7:02 PM . May 6. sterculia.00+$1. By 1909. thesis for the University of Cambridge entitled “Treatment of Habitual Constipation by the Bran Method. its content of vitamin B. hemicellulose.1 writing in 1840 in A Treatise on the Causes and Consequences of Habitual Constipation. which are made from the endosperm and germ.50 © 2001 by CRC Press LLC 183 . As a result of his own observations.2387_ch4. it not only retains moisture itself but enables the other residues in the colon to resist dehydration. and physiological studies of the use of fiber in constipation.” There was little need to convince Burne and his generation that bran stimulated colonic movement and increased fecal output. each generation of medical scientists seems to have to rediscover this fact for itself. Nevertheless. The fibre mixes intimately with the food residues in the colon. such as the presence of undigestible fiber. John Burne. agar. such as Hovis. he concluded. “My view is that bran exerts a mechanical laxative action due to its fibre content. 75 years ago fiber was being singled out as the principal component of bran responsible for its colonic effects. karaya.4pt1_fm Page 183 Sunday.3 in his M. and psyllium 0-8493-2387-8/01/$0. He discussed various current ideas about the way bran produces its laxative effect.4 The Effect of Dietary Fiber on Fecal Weight and Composition John H. biochemical.” Thus. various celluloses. and the production of volatile fatty acids from decomposition of cellulose and hemicellulose. that the substitution of whole-meal bread for white bread is a very important part of the dietetic treatment of constipation. inorganic salts. radiological.” was able to review over 50 papers reporting clinical. I believe that by so doing. It is clear. and medical and nutritional literature in the 1970s contained many papers in which little else but the laxative properties of bran were reported. May 6. recommends that “coarse brown and bran bread is very efficacious. In 1936. “One of the most valuable foods for constipation in whole-meal bread … white bread. phytin.D.” The 1930s were a time of vigorous investigation of the laxative properties of dietary fiber. Purified forms of fiber such as ispaghula. which is made of the endosperm alone and those varieties of brown bread. the bran acting as a salutory stimulus to the peristaltic action of the intestines. 2001 7:04 PM CHAPTER 4. contain only about one-fifth of the cellulose present in whole-meal bread. Cummings HISTORICAL INTRODUCTION Present-day interest in fiber stems from the middle of the nineteenth century when the preoccupation of the Victorians with their bowel habits led many physicians to declaim the virtues of bran. therefore. Sir Arthur Hertz (later Sir Arthur Hurst)2 was writing. Dimock. even when bran is taken only once a day. 4.7 (0. perhaps because of the international conflict which involved many nations at that time. which he interpreted as showing that diverticular disease of the large intestine was due to lack of bulk in colonic contents. 3RD EDITION were all in common use and were a focus of some interesting studies on their mechanism of action. xylan. 2001 7:04 PM 184 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.7 (0.3) 4 5 17 11 Comments Mainly bran.5 (0. 4. corn bran. Surprisingly. apple. and mixed sources Psyllium/ispaghula. thus requiring increased intraluminal pressures. diverticular disease could be satisfactorily treated by increasing the amount of dietary fiber in the form of wheat bran. 2. 4. banana.05) Carrot. However. of Studies (see Tables 2-15) Wheat 5. the work by Williams and Olmsted. methylcellulose. cabbage.0 (N = 9.8) 7 Oats Corn Legumes Pectin 3. Painter was concluding a series of studies in the U. not because of any new insights into the problem of constipation but as a result of epidemiological observations relating bowel habit to disease by Burkitt10−14 and studies by Painter on the etiology and treatment of diverticular disease. etc. Table 4. carrots. gum arabic. potato.7) 28 Gums and mucilages 3. He surmised from these studies that propulsion of intestinal contents was more difficult if they were dry. agar. there followed an apparently complete loss of interest in the subject and no significant developments were reported for the next 30 years.” These studies contradicted the experimental data of others and questioned the whole basis of then currently accepted beliefs about fiber. These studies stimulated new research into the mode of action of fiber in the colon. Other: traga canth.3 2. cooked 4. diluting gut contents. May 6. wheat bran.1) (0.2..2 1. It was widely believed at the time that fiber acted by virtue of its capacity to absorb and retain water in the gut.0) Also carboxymethylcellulose. inspissated. new concepts have emerged with regard to the mode of action of fiber in the colon and its role in determining colonic function. peas. and of small volume. One of these stimulating products is the lower volatile fatty acids. xanthan.4 3.9 (p < 0.4 (0. cabbage.3) (0.9 (N = 4) Oat bran or oats Corn meal or bran Soya products.15−17 In his paper on the etiology of large bowel cancer in 1971. sterculia. concluded that “contrary to the accepted belief. and favorably influencing the microbial flora. Since then. At the same time. as a result of better chemical techniques and advancing knowledge of colonic physiology.4pt1_fm Page 184 Sunday. 8.5) 27 Cellulose 3. Attention to the effect of dietary fiber on bowel habit returned around 1970. much of the early work on fiber has inevitably been repeated and many papers do little if anything to advance our knowledge or understanding of the mode of action of dietary fiber in the colon.2387_ch4. prunes.K.4−8 However.0 (N = 14). He then went on to show that.2 Source (1. Raw 7.7) 41 Fruit and vegetables 4. bassara. the effectiveness of indigestible residues is not due primarily to the mechanical stimulus of distention but rather to chemical stimuli which arise from the destruction of hemicelluloses and cellulose by the intestinal bacterial flora.9 which involved careful metabolic balance studies and fecal analysis of subjects ingesting various fibers prepared from peas.1 Average Increase in Fecal Output per Gram Fiber Fed Increase g/g Fiber ± SEM No.4.5 (N =11) Degree of methoxylation not important . Burkitt suggests that fiber affects bowel function by speeding up transit time.9 (N = 3). contrary to current medical opinion.3) (0. Despite this.29 were probably the first to recognize the significance of the extensive breakdown of fiber in the human gut. only 10% of the cabbage fiber could be recovered in feces. the greater the increase in stool weight (other factors being equal).4. minerals. Table 4.1 is a summary of the result of nearly 100 studies of the effect of the dietary fiber on bowel habit and fecal composition published between 1932 and 1984 and given in detail in Tables 4. It was initially felt that the response to pentose-containing polysaccharides was simply a reflection of the water-holding capacity of a particular fiber. the popular view has persisted that it is not degraded.9 and 5.4. and fiber from cereals survives better than that from fruit and vegetables. while 60% of bran fiber was excreted.26. Williams and Olmsted9. Because of their ability to resist dehydration. and trace elements. and bran as sources of fiber being yet more effective (4. have already emerged from experimental studies as specifically affecting the mode of action of dietary fiber. pectin for example giving an increase of about 1.4. water. Another mechanism has to be sought whereby fiber can alter colonic metabolism. Bacteria are about 80% water and. 70 to 80% of it disappears during passage through the gut. and fruit. however.4. data from the few studies where they have been measured are not included in the accompanying tables.2387_ch4. In so doing they are able to exert a physical effect on colonic function by providing both bulk to gut contents and a surface for the bacteria which allows for their more efficient metabolism. is that not all fibers are equal in this respect.2 to 4. when fiber in mixed diets is fed to healthy subjects. bacteria are more likely to retain their water .10 summarizes studies of the effect of particle size on the efficacy of wheat bran in altering colonic function. The more pentose sugars present in the dietary fiber polymers. They suggested that the fecal bulking by fiber could be accounted for by the stimulatory effect on colonic smooth muscle of the major end products of fiber breakdown. respectively).7 g/g.30 Because of this. Today. or volatile fatty acids. Large particles are more slowly degraded and so are more likely to survive passage through the gut.27 For example.9. The reasons for these differences relate to the physicochemical properties of dietary fiber. However. nitrogen.28 The cellulosic fraction tends to survive digestion better than the noncellulosic polysaccharides. these show that if exactly the same source of fiber is fed at two different particle sizes. vegetables. however.0 g/g fiber fed). Fiber digestion in man has been reported in the literature on many occasions in the past century. In general. the short chain. although there is now ample evidence to the contrary. particle size and chemical composition of individual fiber polysaccharides. a potentially important component of stools. A component of normal stools that has been neglected in studies of fiber for many years is the microflora. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 185 MODERN STUDIES Table 4. Chemical analysis of dietary fiber sources also shows a relation to fecal bulking properties. the greater particle size preparation will produce larger changes in stool output.and hexose-containing polysaccharides hold water and further research has cast doubt on the water-holding hypothesis19 as the sole explanation for the action of fiber. short-chain fatty acids are known to be rapidly absorbed from the human colon and are thought not to control laxation in man.4. therefore.3 g of stool per gram of fiber fed while pure cellulose has an effect more than double (3. All sources of fiber lead to an increase in fecal output and therefore in the components which make up feces such as fat. although its water-holding properties are still an important component of the explanation of its role in altering gut function.18 This association is an intriguing one and one which is at the present time unexplained. little has been learned from the study of their fecal excretion and.25 Since fiber is extensively degraded. therefore. What is clear from Table 4. When the digestion of cabbage and bran fiber was compared in healthy subjects taking equal doses of each material. it is not surprising that its water-holding capacity alone is insufficient to explain its physiological effects in the colon. Two factors. May 6. both pentose.1. The reason for this is thought to relate to the extent and rate of breakdown of dietary fiber in the large intestine.20−25 The most telling argument against the water-holding hypothesis is that virtually all fiber is broken down in the gut.4pt1_fm Page 185 Sunday. microbiological techniques. Many workers who have studied the effect of dietary fiber on bowel function have noted the wide range in response among individuals. in a small number of papers evenly distributed over the years. This again contributes to the change in fecal mass. An increase in lipid excretion may. and noninvasive methods of studying colonic function has allowed investigators in recent years to show that dietary fiber has. plant cell walls which resist breakdown by the microflora. and contained numerical data on changes in large bowel function. and composition published during the past 55 years. Although the literature on dietary fiber and bowel habit contains much that is repetitious. also be explained this way. Studies have been selected for inclusion provided they were conducted on man. In general. There are probably at least four distinct effects of fiber by which it brings about an increase in stool weight. most forms of dietary fiber are extensively degraded by the microflora. substances which increase bulk in the large intestine often speed up the rate of passage through the bowel. e. It is not a comprehensive survey since such a report would have been very extensive and have contained probably little more useful information than the present one. on bowel function. Table 4.4. Using this technique.4. Shortened transit time also leads to reduced water absorption by the colon and therefore wetter stools. The result of this is to stimulate microbial growth and a greater excretion of microbial products in feces. 2001 7:04 PM 186 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Increasing bulk stimulates colonic movement. The advent of better carbohydrate chemistry. Gas trapped within gut contents again adds to their bulk. fecal weight. dietary fiber is an important source of gas in the colon since the gases H2. and CO2 are some of the principal end products of fermentation.9. it would appear that in response to bran. Second. are able to exert a physical effect on intestinal bulk by retaining water within their cellular structure. These mechanisms together combine to increase stool weight. NOTES TO TABLES Selection of Papers These tables summarize essential experimental details from a selection of reports of the effect of dietary fiber.2 to 4. were reported as full papers in the English language. As transit time falls. experiments are described which demonstrate new aspects of large bowel physiology hitherto unrecognized. it will be seen that fiber almost always brings about an increase in fecal nitrogen excretion. and their role in changes in fecal composition is important. particularly . Whether this is the result or cause of the diverticular disease cannot be said but is worth further study. in part. MECHANISM OF ACTION OF DIETARY FIBER The way in which dietary fiber affects bowel habit cannot be explained on the basis of one simple hypothesis. Third. May 6.2387_ch4. Stephen and colleagues31 developed a method for fractionating human feces which allows the contribution from microbial material to be estimated. CH4.g. Fourth.. in many forms. Bacteria are therefore probably an underestimated component of human feces.4pt1_fm Page 186 Sunday. 3RD EDITION against the absorptive forces of the colonic mucosa than are the cellular skeletons of plant material that remain after cooking of food and its subsequent passage through the gut. through its breakdown in the large intestine by the microflora. these patients exhibit only about 50% of the response seen in healthy subjects. First. Studies in both man32 and animals33−34 have shown that this N is most likely to be bacterial N which increases as the result of microbial growth in the colon through fermentation of fiber. because of lignification as in bran.11 summarizes studies of dietary fiber in subjects with diverticular disease of the colon. In Tables 4.4. it can be shown that bacteria compose up to half of the fecal solids in subjects eating typical Western diets. the efficiency with which the bacterica grows improves. an important role in maintaining normal digestive function as it does in the ruminant and other hind gut fermenting species. laid the foundation for much of present-day thinking about dietary fiber. Finally.62 Thus.40. Diets are either “controlled” or ad lib.4. Alternatively. some of these studies are detailed and thoughtful and well worth consulting . TABLES 4. both types of experiments. Controlled diets are where the food is cooked and prepared for the subjects by the investigators and provides a constant background against which . whole foods and purified materials.4.61 on the mode of action of bulk laxatives. The majority of these papers. As a means of guiding the reader through this morass.29. Many early studies from the period 1920 to 195035–52 have not been included because they report only qualitative data. such as those reported by Dimock. Diet. The word “fiber” is used in many different contexts in the papers reported here and clearly means different things to different investigators. and original thought. science. All definitions and concepts are included in the tables. To say that they are solely experiments to determine the effect of dietary fiber on bowel function is an overstatement. all citations to a single experiment are recorded in this column. If these studies had received greater attention.8. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 187 fecal output and composition. Small alterations may be made to numbers of subjects or details and the work republished. however.41 together with the classic work of Williams and Olmsted. being reports of uncontrolled studies which usually show a beneficial effect of bran or other bulk laxatives. The series of papers by Fantus and colleagues in 1940 to 1942. percent changes. Column 1: Literature Sources. possibly in different journals.58. there are problems in interpreting information and extrapolating foods that are normally eaten when dealing with studies where purified materials. however. A great deal that is written about fiber and bowel habit concerns constipation. In recent years the practice has developed in which authors publish the results of a single complete study in several parts. Nevertheless. are worthless.37. or results that are otherwise difficult to convert to tabular form.9. much that is mundane and repetitious in current nutritional research would have been obviated. They have not been included in this survey unless they contain clear data on the role of fiber. Fiber Source The principal literature sources for the data given in the table are recorded in column 1. studies of young children have also been excluded. The answers to some of the questions Dimock posed have not yet been found or even looked for. The problem of the purity of fiber sources has been recognized for many years and early studies of bran. is the repeated publication of the same data in several journals. May 6.53–56 that of Cowgill and associates. only a minority of modern researchers seem to have read or taken note of these early studies.59 and the Comprehensive studies of Tainter et al. and to be decried. Unfortunately.2387_ch4. such as cellulose. since many investigators have used fiber sources such as bran or foods which contain other nutrients.2 TO 4. All this conspires to crowd the literature with irrelevant and superfluous information while apparently enhancing the reputation of the authors for industry. as have those where data are based on a single stool or only 1 day of fecal collection. Nevertheless.60. insofar as they can be identified.3 discuss at length the potential of various components of bran to induce laxation. In general.7. Most studies reported are within-subject crossover type design where each person takes both test and control diets. Some guide as to the meaning of the word in individual studies is given by an indication of the chemical methods used in the study to measure fiber intake in the experimental subjects. the age-old question “what is fiber” constantly appears when reading the literature.4pt1_fm Page 187 Sunday. are used. are needed to give a full understanding of the effects of fiber.11 An asterisk in all columns means that the value is significantly different from basal (or other appropriate) control diet. I apologize in advance to the investigators concerned. on occasion. Fiber sources are described as in the paper. Ad lib diets are self-selected by the subjects. 2001 7:04 PM 188 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. . XI.4pt1_fm Page 188 Sunday. X. Ribiero. VII. Where this was varied. Column 4: Fiber Intake (g/day) and Fiber Method Many authors have attempted to measure fiber intake using one of a number of methods (see below). and Age The number of subjects of each sex (M = male. Widdowson. VIII. 3RD EDITION to judge the effect of fiber. Column 7: Number of Subjects Collecting Feces This number may be less than the total number participating in the study either because the subjects were split into groups or some did not collect feces. Basal = control diet against which fiber supplements were compared. Sex. many studies are now carried out in this way. IX. XII. I. III. (No figure is given for ad lib diets since it is assumed to be the subject’s usual diet. Where no method is given in the paper. and Moran78 McCance.) Column 6: Fecal Collection Period (Days) This is the number of days of fecal collection pooled for analysis. Column 3: Study Period Study period describes the diet and/or fiber source relevant to the data given on that line. F = female) who took part in the whole study or part of the study are recorded together with the range of ages. These are sometimes referred to as “metabolic studies. If errors have been committed or wrong assumptions have been made by me in doing this. and Shackleton79 Katan and von de Bovenkamp80 Englyst81–82 Schweizer83–84 Angus. II. averages of several collection periods for the data. It is the minimum included since investigators have used. Column 2: Total Number of Subjects in Study. V. VI. a range is given. and Farrell 85 Column 5: Days on Diet This information appears as in the paper. IV. May 6. Crude fiber 63–64 Williams and Olmsted 193556 Southgate65–70 Paul and Southgate71 Van Soest and modifications 72–76 Crampton and Maynard77 O’Shea. an attempt has been made to calculate dietary fiber intakes using currently available food tables or analytical information. Sutherland.2387_ch4.” Commendably. It is the number which has been used to calculate standard errors in other columns. A dash indicates that no data are available. Where data are given in the paper for several periods on the same diet.× 100 total stool weight g/day Column 10: Fecal Solids (g/day) As given in the paper or calculated from data related to percent moisture of feces (± 1 SE of the mean). Values for transit time obtained are very dependent on the method used. Colored dyes give short transit times since it is difficult to detect other than the “first appearance” of marker in feces.4pt1_fm Page 189 Sunday.86–87 Where dietary fiber data are not given in the paper. in the majority of which an oral dose of inert marker is given with food and its appearance in feces is noted. IV. the time it takes a substance to pass through the gut. and investigators seeking to follow up these studies should consult the publications themselves. and Wiggins (1975 and 1976)89–90 Colored dyes — “first apperance” method91–93 51Cr sodium chromate94 Cr2O6 chromic oxide . Any errors are therefore not the fault of the original authors. Column 9: Percent Moisture If not given in the original paper. can be measured using a number of techniques. Values in this column must be interpreted. these have been calculated from available knowledge of the food and fiber source used. Hinton. V. in the light of the method used and are not directly comparable with one another. the information relating to the final period has been used in this table. this has been calculated as: total fecal weight g/day – fecal solids g/day -------------------------------------------------------------------------------------------------------. Jenkins. Column 12: Transit Time — Hours and Transit Method Transit time. while the radio-opaque pellet technique of Hinton et al. and Young (1969)88 Cummings. Methods which measure mean transit time are probably most accurate.2387_ch4. May 6. Transit Methods I. III. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 189 Column 8: Fecal Weight (g/day) Average value (± 1 SE of the mean).88 gives values which are about 20% greater than the mean transit time methods. Column 11: Apparent Increase in Fecal Weight Per Gram Dietary Fiber Fed Calculated as: fecal output with fiber source g/day – basal fecal output g/day ----------------------------------------------------------------------------------------------------------------------------------------------------g fiber fed per day Gram of fiber fed per day is always “dietary fiber” as defined in 1972 and 1976. therefore. Some of these figures are derived by calculation on the basis of data in the original papers. Lennard-Jones. II. potassium. Column 18: Comments.75 to 6.2387_ch4. chloride. phosphorus (including phytate) D = Iron. Column 15: Neutral Steroids (mg/day) See original papers for methodology which affects results especially in early (pre-1970) papers. and fatty acids and fatty acids alone (after hydrolysis). triglycerides. bicarbonate.4pt1_fm Page 190 Sunday. Millimoles converted to milligrams by factor × 386. zinc. Millimoles converted to milligrams by factor × 400. N = Hematology . Notes. and H2. ash. copper. May 6. 2001 7:04 PM 190 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. including those which measure total lipid. usually measured by calorimetry.O6. Column 17: Energy (kcal/day) Total fecal energy excretion. Other Data Available From Study A = Digestibility (fecal excretion) of fiber B = Volatile fatty acids C = Calcium. Cr2. Column 16: Acid Steroids (mg/day) See notes for Column 15. 3RD EDITION Column 13: Fat (g/day) Note that various methods are used to recover fat.25). or other electrolytes J = Microflora K = Glucose tolerance L = Bile composition and bile acid kinetics M = Breath CH4. and other trace elements E = Blood lipids F = Intestinal motility G = Polyethylene glycol. Other Data Available This column includes special notes about the studies and lists other data available from related publications of the same study. Column 14: N (g/day) This is usually measured by the Kjeldahl technique. millimoles converted to milligrams by factor × 280. Note variable conversion factors used by authors to convert fecal N to “protein” (from 5. magnesium. used as markers H = Sodium. without some additional information. This is seen in the papers on different plant gums and legumes.2 g/g raw vs. and one cannot really generalize about the whole group.4 g/g. since few investigators use these studies to cast light on the unresolved question as to how dietary fiber works. despite being a good source of insoluble fiber.220. and must include. pectin remains the least effective along with guar.255 this is hard to understand. Since they also frequently contain resistant starch.012). Fruits and vegetables remain second in the table.2387_ch4. effective as a laxative. well ahead of the rest. nature. There is clearly a lot of commercial interest in adding these preprations to food due to their potential cholesterol lowering properties. Many fruits and vegetables contain significant amounts of insoluble fiber. fruits. The impetus for these studies is partly commercial.4. Such omissions are inadvertent and the author apologizes to anyone whose work has been so excluded. These additional data have not made a great deal of difference to the overall rankings.247. fecal weights. Finally. remain unequivocally at the top of the league. Therefore. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 191 TABLES 4. by study of the physical properties of fiber in relation to its effect on fecal composition. in fact this is clearly not the case.9 cooked. Thirtyseven additional papers have been added to the original compilation of about 80. not observational. Dietary fiber is laxative. especially soya. ranked according to their effectiveness. With regard to wheat. As before. although there is clearly now an increased interest in forms of dietary fiber other than bran. guar is a poor bulker in general. Cellulose. giving increases of around 1. contain quantitative data of an experimental. There may be some publications that do not appear in these tables but which fit the criteria for inclusion. wide differences in the fecal bulking properties of gums are evident. Gums and mucilages have risen up the list to third place.248 The revised version of Table 4. Some progress has been made in this area. as a minimum.4. t = 2. . p = 0. if not more. and also include one or two additional reports omitted from the earlier survey. of course. The disadvantage of raw bran. and vegetables.4pt1_fm Page 191 Sunday. which are better. a single number on a food label for fiber is going to give the consumer very little useful information about its laxative effects. but the significant differences among the results confirm that not all fibers are the same.12 TO 4.254 These studies were mostly done with foods rather than purified forms of fiber. detailing a total of approaching 150 individual dietary studies of assorted fiber sources. For example.253. although they are well behind wheat. While many people believe fruits and vegetables contain mainly soluble fiber with little potential to alter bowel habit. May 6.220.225 while tragacanth seems to be the best at 6. usually bran. papers still continue to appear on this topic at a steady rate. and the message remains the same. and oats are all effective laxative substances and give similar effects. The legumes.15 (1986–1992) These additional tables detail papers on the effects of dietary fiber on bowel habit and fecal composition published between 1986 and 1992. but a number of important questions remain unanswered . which means that the subjects are eating intact cell walls.228 although this is only one study. 4. there are now enough studies of both raw and cooked products to show a statistically significant difference in favor of raw bran (7. although a small number of studies of methylcellulose suggest it is equally.4.253 are poor fecal bulkers. the criteria for inclusion are that the studies have been undertaken in adults. While in 1985 it appeared that everything that could be said about dietary fiber and bowel habit had been said. Some of them add little to what is already known. However.65. is its high content of phytate.0 g/g. Wheat products.1 shows average increases in fecal weight (g per day per g fiber fed) for the major sources of dietary fiber. corn. which probably contributes to the effect. with the exceptions of methylcellulose and carboxymethylcellulose. A further seven have been added for the present update as follows: XIII. although despite this the overall differences are statistically significant (by ANOVAR F 4.232 In addition. XIX.256 this creates a problem in comparing data. However.239–245 Because these methods give widely differing results for the amount of dietary fiber in a food. a further seven methods or modifications to methods have been developed over this period. the amount and type of starch in experimental diets must be controlled (as must its processing) to eliminate this as a variable.215 Theander and Westerlund 231 Asp et al.200 Prosky et al. The variability is due in part to the inherent difference in individual responses and to varying experimental designs. XIV. In the earlier part of this chapter. Anderson. XVII. .4pt1_fm Page 192 Sunday. 2001 7:04 PM 192 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.2387_ch4. What of the future? The emergence of starch as a fecal bulking agent 248–252 is probably the single most important discovery to challenge the role of fiber in the diet. we shall have to re-evaluate the whole dietary fiber story in the light of emerging evidence of the importance of resistant starch. XVIII. More than likely. Sieling. XVI.001). some of which are uncontrolled diets.78 p < 0.202–204 Slavin et al. 12 methods were listed as being used to measure fiber in the various studies. and Chen197 Mergenthaler and Scherz199 Meuser et al. XV. a major problem is lack of consistency in methodology for measurement of dietary fiber. 3RD EDITION How reliable are these rankings? Within each group there is great variability. At the very least. May 6. 4. 1–11/2 oz Days on Diet I 5. 1–11/2 oz Low fiber basal + Whole bran. 1–11/2 oz Very low fiber basal + Whole bran.4pt1_fm Page 193 Sunday.1 3.4.8) Whole bran slightly better than breakfast bran but less palatable Both brans equally effective A 2387_ch4.4. May 6. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 193 Energy (kcal/day) Acid Steroids (mg/day) Neutral Steroids (mg/day) N (g/day) Fat (g/day) Transit Time (h) and Transit Method .2 Fecal Collection Period (days) 7 7 7 7 14 14 14 14 14 14 14 14 Number of Subjects Collecting Feces 6 6 6 6 4 4 4 5 5 4 5 5 Fecal Weight (g/day) 121* (6) 187* (18) 176* (12) 78 (15) 185* (17) 189* (22) 93 (5) 112 (7) 171* (13) 93 (7) 116 (11) 193* (8) Moisture (%) 76 76 74 76 77 70 79 77 Fecal Solids (g/day) 44 22 49 27 39 28 40 27 Comments Notes Other Data Available Constipated subjects Also studied effect of fruits and vegetables (see Table 4.2 Literature Source Diet Fiber Source Cowgill and Anderson Effect of Wheat Fiber in Various Forms on Fecal Composition (See Also Table 4.(1932)58 Cowgill and Sullivan (1933)59 Controlled diet Normal foods Bran and breakfast bran (a commercial breakfast cereal) Controlled diet Normal foods Low fiber or very low fiber Bran — whole or with weak acid Total Number of Subjects in Study (sex and age) 6M 5M Study period 30−38 50 90 90 + Bran + High bran + Breakfast bran Ia 7 7 7 7 14 — 14 — 14 14 3. 1−11/2 oz Low fiber basal + Acidwashed bran.10) Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed Table 4.2 — 14 Fiber Intake (g/day) and Fiber Method Basal Very low fiber basal + Acidwashed bran.1 14 14 14 — 5. 2 Comments Notes Other Data Available Laxative properties of brown breads similar C Part of a larger study in which 9 other fiber sources fed (see Tables 4.4. prepared by washing for 24 h in warm water. 21− 42 3 M. B Noted greater fiber digestibility than in normal subjects A 194 Williams and Olmsted Transit Time (h) and Transit Method Fecal Solids (g/day) Moisture (%) Table 4. May 6. brown with dephytenized bran.4.7) A. 2001 7:04 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. or brown with dephytenized bran and mineral supplement (1942)95 Controlled diet Normal foods excluding fruit and vegetables Bran.10) 2387_ch4. brown.29.2 (Continued) Effect of Wheat Fiber in Various Forms on Fecal Composition (See Also Table 4.Literature Source Diet Fiber Source (1936) Controlled diet Bread as 40 to 50% of energy intake Bread either white. and drying McCance and Widdowson 9. 3 3M Study period Brown Brown dephytenized Brown dephytenized + minerals White Bran 6 Fiber Intake (g/day) and Fiber Method 7 7 7 10 7 7 Fecal Collection Period (days) 10 10 10 Days on Diet II + 24 Number of Subjects Collecting Feces 6 6 6 6 3 Fecal Weight (g/day) 226 224 209 116 +79b Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 3.4. extracting with hot ethanol.4.57 Total Number of Subjects in Study (sex and age) F.4pt1_fm Page 194 Sunday.4–4. 3RD EDITION Energy (kcal/day) Acid Steroids (mg/day) Neutral Steroids (mg/day) N (g/day) Fat (g/day) . 0) 2.8 (0. 28− 36 Findlay et al.2) 3.2) 1.2) 1. 21–25 2 M.11) Cellulose also studied (see Table 4.6) 2. the sec. (1975)107 Walters et al. H See original paper for comments on liquid and solid phases of gut contents F.2 12.3 9. 20 g Basal 6. IV 66 (18) 44* (4) 93 (16) 43 I 62 1.3 5. C. Cornflakes 1 oz + Bran.2) 602 (79) 820 (121) 675 (81) 646 (73) 199 (19) 279* (19) 195 (17) 199 (44) 166* (6) 108 (6) 296 (32) 352 (45) 216 (46) 207 (34) B.1) 1.4. Controlled diet Hospital low fiber Bran crisp-bread (Energen) (1977)109 McLean Baird et al.4) E 2387_ch4. 25−43 Ad lib diet Bran buds or cornflakes (1974)97 Connell and Smith Ad lib diet Bran (1973)96 Eastwood et al. 52– 69 + Fiber Basal + Bran crispbread Basal + Bran. 1 oz 10. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 195 .5 40* (4) II 58 (3) 50 (11) I. (1976)108 Southgate et al.7 (0.0* (0.5 7.4. C−E.6* (0.ond an inpatient controlled diet study of wheat bran biscuits A.16 g Basal Bran buds. 25− 45 8 M.2 (0. 3 F.7* (0.4.Controlled diet Normal foods (1975)100 Jenkins et al.7).8 (0. J Study included additional group with diverticular disease (see Table 4. 45 IV 17 28 15 + 9c — +8 IV + 7c — 21 21 12 12 35 7 28 28 21 7 7 7 7 7 7 7 7 7 7 7 6 6 5 5 6 6 8 8 8 8 228* (30) 79 (7) 166* (15) 93 (10) 120 (18) 183* (22) 218* (20) 119 (9) 107 (15) 174* (18) 80 73 77 75 74 72 76 76 45* (2) 21 (1) 38* (8) 23 (1) 46* (3) 33 (4) 41* (3) 26 (3) 5. Ad lib diet Bran (1974)98 6 M.2) 2. May 6.4pt1_fm Page 195 Sunday. E. (1976)99 Cummings et al.1 (0. G Two separate studies: one of bagasse biscuits in nuns living in a convent (see Table 4. Floch and Fuchs (1977/78)103. (1976)101 Fuchs et al. 58 to 62% of energy as bread Bread: white or basari (80 to 90% extraction) (1976)105 Reinhold et al. 2 F. C. D Eating habits showed reduced eggs.1) 2.104 Ad lib diet All Bran (Kellogg’s) (1976)102 Kahaner et al.10) 2387_ch4. May 6. and breakfast meats with increased milk and fruit during bran period Stool pH unchanged Fecal anaerobe: aerobe ratio increased by bran E. butter.3 (0.9 (0. and bran biscuits Transit Time (h) and Transit Method Table 4. 22− 47 Study period Basari.1) Comments Notes Other Data Available A.2 (Continued) Effect of Wheat Fiber in Various Forms on Fecal Composition (See Also Table 4.4. All Bran. 35 4 M.4pt1_fm Page 196 Sunday. bran. 500 g White Bread 500 g All Bran. J 196 Whole meal bread. 2001 7:04 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 24. 3 oz Basal Fiber Intake (g/day) and Fiber Method 30d 22d V + 23 IV Days on Diet 20 20 21 21 Fecal Collection Period (days) — 3 3 Number of Subjects Collecting Feces 2 2 6 6 Fecal Weight (g/day) 403 (121) 244 (92) 103 (20) 226* (45) Moisture (%) 90 91 74 73 Fecal Solids (g/day) 40 (8) 21 (5) 59 28 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 5. Total Number of Subjects in Study (sex and age) 2 M.Literature Source Diet Fiber Source Controlled diet Normal foods.3 Fat (g/day) 2. 3RD EDITION Energy (kcal/day) Acid Steroids (mg/day) Neutral Steroids (mg/day) N (g/day) .4. 0* (0.2 4.6 14 + 3.4.7* (0. H Part of a larger study in which fiber from carrot.(1980) Controlled or specified menus (1978)111 Farrell et al.6 (0.5 (0.4 — 43* (3) II 73 (10) 50 (9) 58 (6) 48* (7) 53 (17) I 65 (8) 2. E Stool weight not significantly increased by any treatment Cooked bran thought to be less effective than raw bran 2387_ch4.9 + 3.9 (0. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 197 . May 6. fruit.1) 2.1) 2. cabbage.6 4 4 7 7 6 6 5 5 5 5 5 14 14 6 6 5 5 10 10 10 10 10 — 197* (13) 95 (8) 164 (20) 125 225* 131 (17) 183 21 159 (13) 139 (9) — 76* 72 71 74 77 73 76 76 75 50* (1) 29 (1) 46* (1) 27 (1) 32 53* 35 (3) 36 (3) 38* (3) 42* (5) 30 (3) 5.8 3. 3 M. 11−41 Ad lib low fiber avoiding cereals. Controlled low fiber Normal foods White bread or coarse bran (Prewetts) made into bread and whole meal bread 14 M. apple (see Table 4. and vegetables (1976)106 Wyman et al. VII. VI.2 g All Bran. XII 33 40 III 22 9 9 21 21 14 21 14 + 5. 22−46 19 M.7 8. and guar gum (see Table 4. 30 g Basal Basal + Fiber (23–35 g bran) (1977)110 Raw bran or All Bran (Kellogg’s) Kay &Truswell IV Basal Raw bran. C−E.7). 22 g 10 M.1) 1. bread.1) 288 256 312 233 241* (6) 151 (6) A.6 2.4.5) were fed J Study design was basal-fiber-basal Steroid output also higher in second basal period C. 13. 22−27 + Fiber Basal + Bran.3e 5.7 5.0* (0. Controlled Normal food Bran prepared by extracting with ethanol and acetone 25 Stephen and Cummings (1978) 18 Cummings et al.4) 3.7 14. 20−38 3 F. 20 g All Bran. 53 V.4pt1_fm Page 197 Sunday. 12 g Raw bran.3) 1.6 14 14 + 5. 3RD EDITION Neutral Steroids (mg/day) N (g/day) Fat (g/day) .4 3. (1981)115 Sandstead et al. (1978).bran.2 3. etc. All Bran. 2001 7:04 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Fecal Collection Period (days) 12 12 12 12 7 7 Number of Subjects Collecting Feces 9 9 6 6 4 4 10 10 Fecal Weight (g/day) 151* (8) 81 (6) 99* (7) 64 (10) 210* (9) 80 (11) 255 (17) 295* (14) Moisture (%) 73 72 Fecal Solids (g/day) 56* (1) 22 (2) 5. 30 g 4 M. K No effect of protein intake on stool weight noted in other part of study C Significant effect of bran on patients with amoebic dysentery also noted in separate study 198 Normal foods All bran. May 6.4.118 Fiber Intake (g/day) and Fiber Method 28−30 + 13 Hard red spring wheat bran.119 (1979)120 Munoz et al. 20−24 10 M Study period Ad lib diet Reground bran (1978)112 Mathur et al.5 to 4.4. 19−54 High protein + Fiber Controlled diet Normal food Fiber from bran crispbread.2 (Continued) Effect of Wheat Fiber in Various Forms on Fecal Composition (See Also Table 4. on various aspects of metabolism (see Tables 4.Literature Source Diet Fiber Source Total Number of Subjects in Study (sex and age) (1978). and bran biscuits Transit Time (h) and Transit Method Table 4. 26 g 28−30 28−30 21 21 28−30 + 11 V 53 V 22 30 Days on Diet Basal Soft white wheat bran.2 4.7) C−E.4pt1_fm Page 198 Sunday. whole meal bread. soybean hulls. (1979)114 Bell et al. Basal + Bran.4. and fine bran (1979)113 High protein Cummings et al.4.10) 2387_ch4.0 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed c 50* (7) II 70 (6) 32 (2) I 37 (2) Acid Steroids (mg/day) 221 (19) 444* (27) Energy (kcal/day) 530 (21) 477 (107) 275 (40) 328 (86) Comments Notes Other Data Available Part of a larger study of corn bran.116 (1979)117. 10 M. 26 g Basal Munoz et al. whole meal bread. Controlled diet Normal food Hard and soft bran fed as bread 10 M. Ad lib but semicontrolled Normal food AACC bran (1980) 122 Huijbrechts et al.8 1.5 (0. 18−28 18 M.4) 1.126 Stasse-Wolthuis et al.6 4.5) Nitrogen balances Part of a larger study including pectin and the effects of fruits and vegetables (see Table 4.3 (3.5 g/kg (34 g) Basal 4.0 4.8 Basal V 18 35 6 7 10 16 143 (11) 166* 36 (1) 36* III 32 (2) 48* 37 22 8 17 62* (11) + Bran.(1981) 127 Yu & Miller Controlled diet Normal food Bran (1979)125. colonic pressures unchanged C. 20−35 40 M. 65–96 7.4.7 (0.5) Also transit studies with radioisotope capsule. H 2387_ch4.4. May 6.3) Blood pressure unchanged C. –  22. 38 g 89 (12) 27 62 76 42 (2) 67 (7) 16 10 10 4 74 Basal 7 5 5 4 194 (32) 273 (49) II 35 28 56 4 IX 18 +8 45 V 31 VIIIg Basal + Fiber. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 199 .5) 11.8 (1.7 (2. F.E.9 (2. 20 g + Bran 0. (1980)124 Stasse-Wolthuis AACC bran Ad lib hospital diet (1980)123 Smith et al. H Seven subjects studied for 4 weeks.4 5.5 74 (1) 4.8 kg No change in bile composition or kinetics after bran L Elderly hospitalized subjects −− also studied with ispaghula (see also Table 4.1) 629 597 18. 19 F –  1 81.9 (0. then 4 for a further 4 weeks Average weight of 4 subjects: 67.1) 254* 288 160 216 f 17.9 3. 22 F.3) 2.4pt1_fm Page 199 Sunday.2) 179 (5) f 10. low dose Coarse bran. Controlled diet Normal food Breads containing coarse and fine bran (1982)128 (1983)129 11 F.4.1* (0. Total Number of Subjects in Study (sex and age) + Bran.1) Acid Steroids (mg/day) 316 258 324 355 374 322 Energy (kcal/day) 211* (5) Comments Notes Other Data Available Constipated subjects.5 1.10) 2387_ch4.8 1. 20 g Basal Basal + Coarse bran Coarse bran. H 200 Transit Time (h) and Transit Method Table 4.0 4.6* 2.4 (0. 2001 7:04 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.10) A−E Mainly a study of mineral balance and fiber digestibility A.8 4.4pt1_fm Page 200 Sunday.4. also studied with corn bran (see Table 4. 3RD EDITION Neutral Steroids (mg/day) .1 1.AACC bran Ad lib low-fiber diet (1982) 130 Graham et al.6) which was more effective per gram material fed Also studied coarse and fine brans (see Table 4. May 6. 20−40 12 M.4. –  23 + Bran.8 1.4.4 5.6) N (g/day) 1.3 1. 15 g Study period Controlled diet Normal food AACC bran added to food and cooked 20 22 22 +8 14 14 20 20 22 35 X 20 20 V 9 22 Fiber Intake (g/day) and Fiber Method 18 Days on Diet +6 Fecal Collection Period (days) 7 7 4 4 4 4 4 4 6 Number of Subjects Collecting Feces 5 5 4 4 12 4 12 12 10 Fecal Weight (g/day) 58* 31 (9) 158 143 137 202* 77 140* 172* (12) Moisture (%) 72* 67 75 (1) Fecal Solids (g/day) 16* 10 35 34 34 46 21 34* 43* (1) 3.2 (Continued) Effect of Wheat Fiber in Various Forms on Fecal Composition (See Also Table 4.8 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 35 45 77 45* I 88 52* 29 (2) Fat (g/day) 5. high dose Coarse bran Whole meal bread Literature Source Diet Fiber Source Van Dokkum et al. 4 5.7) 3. 110 g Basal + Bran bread.3 (0.7) E Principally a study of the effect of fiber or mineral balance.8 (0.3) 4. 11 F Bran. A variety of fiber methods used (see paper). Figures are increases over basal diet period—fiber remaining in stool.2) 2.9 6.1) 220 (17) 212 (35) 263*(9) 125 (12) 136* (12) 135 (14) 162* (14) 133 (18) 163* (14) 178 (9) A dose–response study showing a linear increase in stool output with increasing doses of bran-enriched bread A.2) 1. or whole meal with added phytic acid (1983)131 Andersson et al.2) 1. White bread Brown bread Whole meal bread 5 M.5 (0.3) 2. D H 2387_ch4.7 6. 19 M.8) 4.2) 1. Assumes bran is about 44% dietary fiber. 170 g + Bran bread.5) 3. C−E A study comparing in vitro and in vivo properties of potato and bran fiber (see Table 4.3) 2.3) 2.2) 1.6 (0. X 16 114 (20) 128* (22) 130 (28) 163* (27) 110 (15) 145* (13) 173 (15) 240* (14) 183* (8) 120 (6) 137 (38) 175* (50) 236* (51) 77 79 75 74 78 78 76 75 75 72 80 79 79 54* (2) 37 (2) 36* (2) 28 (3) 36* (3) 29 (3) 30* (2) 28 (3) 46* (1) 33 (1) 47* (6) 37* (6) 29 (5) 5. 26−44 Basal 24 Eastwood et al. brown. 16 g Ad lib diet Coarse bran (1983) 9 M.3 (0.7 (0.9 (0.4 7.7 (0. and which showed no effect C.4* (0.4.g f e d c b a + Bran bread.4 (0. 60 g Basal + Bran bread.4 4. Per gram of dietary fiber.6 (0.5 (0.4pt1_fm Page 201 Sunday. Micromole per kilogram per day.2) 1.1 (0.1) 2. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 201 . Acid detergent fiber.4 7.7* (0. 1 F Controlled diet Normal food Bread either white.0 34 (3) 39 (4) 59 (7) 54* (10) 73 (15) 68 (14) 53 (12) 66 (17) 49 I 55 39 (12) 41 (13) II 46 (17) 2. May 6. Controlled Normal food Bran-enriched breads (1986)132 Basal Stephen et al. 30 g Basal 21 21 24 36 21 21 21 30 21 21 21 25 21 22 21 X 20 7 XI 19 21 24 31 27 24 24 24 6 6 6 6 6 6 6 6 7 7 6 6 6 8 8 7 7 8 8 7 7 8 8 6 6 6 Milligrams of crude fiber per kilogram body weight per day.3) 1.5 (0.5 (0. 22–25 Total Number of Subjects in Study (sex and age) Drasar and Jenkins (1976)133 Ad lib diets Pectin–Bulmers NF +32 + Pectin 35 g Basal + Pectin. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 202 Energy (kcal/day) N (g/day) . (1976)134 Literature Source Diet Fiber Source 10 M. May 6.4.3 3.6 Neutral Steroids (mg/day) 335 390* Acid Steroids (mg/day) 265 371* Comments Notes Other Data Available Study design was control-testcontrol. 21–28 12 M.6 0.8) Serum cholesterol lowered by pectin but fecal microflora unchanged E.4pt1_fm Page 202 Sunday. data from first control period reported here Serum cholesterol fell but not triglycerides E Serum cholesterol fell but not triglycerides E Also studied were guar gum and bananas (see Tables 4.5 and 4.4.3 Number of Subjects Collecting Feces 9 9 12 12 4 4 Fecal Weight (g/day) 140 168 186* (15) 150 (10) 119 (22) 148 (13) Moisture (%) 71 68* Fecal Solids (g/day) 40 52* Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 2. 5 F. 12 g +12 III +10 III III — Basal 14 Fiber Intake (g/day) and Fiber Method 14 21 21 7 14 Days on Diet Basal Study Period Effect of Purified Pectin on Fecal Composition 6 6 5 5 7 7 Fecal Collection Period (days) Table 4.9 Transit Time (h) and Transit Method III 34 37 44 (6) I 59 (9) Fat (g/day) 3.8 8. J 2387_ch4.4. 22–45 Durrington et al. 15 g + Pectin.Kay and Truswell (1977)135 Controlled diet Normal food Pectin–Bulmers NF Ad lib diet Pectin–Bulmers NF 4 M. 6 (0.9 (0.2) 669 617 642 (87) 585 (62) 398 263 239 (42) 322* (37) 537* (93) 342 (8) Subjects selected for low stool weight and slow transit Cellulose also given (see Table 4.4) 3. 10 g 14 21 Basal + Pectin.6* (0. 3RD EDITION .0 — 0. L 203 Stasse-Wolthuis (1979)125.5) 2. 4 M.5 1.2) 3. (1980)124 Spiller et al.8* 1.5 (0. 1 diverticular disease Serum cholesterol decreased but not triglycerides Bile composition unchanged E.40 M. changes in bowel habit somewhat greater than at 6 weeks 2 Normal subjects. 36 g Basal + Pectin. H Serum glutamyl transpeptidase unchanged A. 23–60 5 M.1) 2.4) 1.4. E. (1979)137 Controlled diet Normal food Pectin–Bulmers NF 21 42 35 III 15 48 VIIIa IX 18 27 Basal +Pectin.9 I 118 (13) 120 (17) 63 59 (6) II 70 (16) II 77 (18) 1. 2001 7:04 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 21–24 Cummings et al. (1980)138 Ad lib diet but excluding certain high fiber foods Pectin–Sunkist 42.4.3 (0. 6 hyperlipidemics. 18–28 Stasse-Wolthuis et al.4.4) B Part of a larger study including wheat bran and the effect of fruit and vegetables (see Tables 4.2 and 4.126 Controlled diet Normal food Pectin–Bulmers NF 5 F.4pt1_fm Page 203 Sunday. 33−60 Miettinen and Tarpila (1977)136 Controlled diet Normal food Pectin-Firmagel Bucness Ltd.8) Blood pressure unchanged Serum cholesterol reduced C.2) 1. May 6. 40–5 g 35 14 III Basal 7 7 7 7 7 7 3 3 12 12 15 15 5 5 9 9 54 (8) 55 (5) 99 89 (9) 107 (25) 123* (25) 269 (49) 200 (89) 74 75 72 73 26 22 (5) 31*(3) 27 (3) 46* (6) 36 (3) 0 1. 1 diabetic.1 (0. 25 F. 2387_ch4. C After 3 weeks on pectin.7* (0. 6 g +6 III 14 +37g + Pectin. 15 g High methoxy. 20–27 Fiber Intake (g/day) and Fiber Method Low methoxy. 15 g Study Period 22 III Days on Diet Basal Fecal Collection Period (days) 7 7 6 6 10 10 8 8 Number of Subjects Collecting Feces 126 105 Fecal Weight (g/day) 161* (21) 124 (8) Table 4.5) 6.4. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 204 Energy (kcal/day) N (g/day) . but fecal β-glucuronidase activity increased 35% while 7-αdehydroxylase activity was unchanged 2387_ch4. triglycerides unchanged High methoxy pectin produced greater fecal bulk E No significant changes in fecal excretions. May 6. 23–38 8 M.3 (Continued) Effect of Purified Pectin on Fecal Composition Moisture (%) 74 72 74 70 Fecal Solids (g/day) 39*(2) 34 (2) 33 31 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 1.Literature Source Diet Fiber Source Judd and Truswell (1982)140 Ad lib diets Pectin–Bulmers NF Either 37% (low) methoxy or 71% (high) methoxy Normal food Low fiber Pectin “slow set grade 200” Ross and Leklem (1981)139 Controlled diet Total Number of Subjects in Study (sex and age) 5 M.8 (0.2 (0.8) 4.9 5. 5 F.5 Transit Time (h) and Transit Method 23 (3) III 25 (3) 34 III 32 Fat (g/day) 6.1 Neutral Steroids (mg/day) 399 (71) 452 (60) 863 866 Acid Steroids (mg/day) 412 (31) 436 (51) 464 425 Comments Notes Other Data Available Both pectins reduced blood cholesterol to a similar extent. 15 g 21 21 IV 34 34 22 +12 +Pectin.4pt1_fm Page 204 Sunday. May 6.4 and 4.0 57 (4) 51 (7) Cellulose and lignin also studied (see Tables 4.6) Fecal pH unchanged Fecal mutagenic activity negligible in all diets A. (1983)144 Ad lib diet Pectin. 9.4 to 4.4. 10 M.5 g/kg Basal 2 2 3 3 A variety of fiber methods used.4.7) Stool pH unchanged Part of a study including cellulose. 0. 21–32 Fleming et al. corn bran. M 2387_ch4.3 g/kg. Assumes average weight of subjects 70. (1983)141 Marthinsen and Fleming (1982)142 Fleming and Rodriguez (1983)143 Pectin–Sigma 10 10 5 5 135 (21) 119 (25) 86 54 22 14 0 1.4. methoxyl a 5 M. 21–43 Hillman et al.b 9 +32b 28 28 + Pectin.4. and xylose (see Tables 4. 20 F. B.0 kg and pectin 91% dietary fiber.4pt1_fm Page 205 Sunday. 12 g +11 III 9 V 0 Basal + Pectin. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 205 . 20 g 7 7 +10 + CMC.4 2387_ch4. 4.4pt1_fm Page 206 Sunday.6. B Part of a study in which a variety of combinations of methylcellulose. and 4.4. 2001 7:04 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.7) A.0 4.4.0 6.29. magnesium oxide.4.9) Further studies were undertaken in the same subjects with a combination of psyllium and methycellulose (see Table 4. 10 g — 7 + 45 + 10 7−14 7−14 6 Fiber Intake (g/day) and Fiber Method II Days on Diet Basal Control + Methylcellulose 4.5 g Basal + Methylcellulose.0 10. and bran were tested for their laxative effect (see Table 4.2 Comments Notes Other Data Available Part of a larger study in which 9 other fiber sources fed (see Tables 4. 10 g — 8 Celluflour Study Period 3M Total Number of Subjects in Study (sex and age) Berberian et al. 4.57 Controlled diet Normal food Cellulose— celluflour commercially available Tainter (1943)60 ”Constant” diet Normal food Methylcellulose Fecal Collection Period (days) 7 7 7 7 7 7−14 7−14 7 — 8 8 — 3 Number of Subjects Collecting Feces Effect of Cellulose and Cellulose Derivatives on Fecal Composition 180 140 100 163 190 127 232 + 15 Fecal Weight (g/day) 80 80 81 82 Fecal Solids (g/day) 32 38 24 41 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 4. 3RD EDITION Energy (kcal/day) Acid Steroids (mg/day) Neutral Steroids (mg/day) N (g/day) Fat (g/day) Transit Time (h) and Transit Method .2.Literature Source Diet Fiber Source 38 Marks (1949)145 Ad lib diet Sodium carboxymethyl cellulose (CMC) + 12a +20 + CMC.4.4.4. May 6.4.5.9) An early study briefly described 206 Moisture (%) Table 4. (1952)61 Ad lib diet Methylcellulose tablets Williams and Olmsted (1936)9.5 1. 5.4.4 (0. 3 taken with either white bread or Basari (80 to 90% extraction brown bread) Slavin and Marlett (1980)147.5 16. May 6. D Bran also studied (see Table 4. corn bran. 20−39 Ismail-Beigi et al. and xylan (see Tables 4. magnesium. 16 g + 16 7 Basal 3 3 7 7 5 5 — — 7 7 5 5 13 13 7 7 2 2 3 3 4 4 106 54 97 (11) 64 (7) 130* (11) 75 (9) 280 (112) 482 (225) 223 141 (24) 152 (16) 221* (29) 70 75 74 81 42 14 39* (3) 19 (1.3) B Subjects selected for low stool weight and slow transit Fecal calcium and magnesium excretion increased Fat. C Cellulose caused increased fecal excretion of calcium. (1983)141 Marthinsen and Fleming (1982)142 Fleming and Rodriguez (1983)143 Controlled semipurified diets Cellulose— Alphacell from ICN Normal food Cellulose— Solka floc® 8 M. M Pectin also given (see Table 4.148 Controlled diet 42.5 g/kg Basal + Cellulose.2) E 2387_ch4.1) 189 (12) 103 (9) 478 (122) 396 Part of a study including pectin.3 3. 4. 16 g Basal e 21 9 + 14 V 0 9 14 — + 35 g 20−30 20−30 V 9 24 14 + 12 14 9−20 + Celluloseb 12 g Basal Basari bread + Cellulose.3 (0. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 207 .4pt1_fm Page 207 Sunday. nitrogen.3. B.8 4. 12 g 9−20 Basal white bread +12 V 21 + Cellulose.2) 1.5) 57*(6) 29 (4) 1. and energy digestibilities unaffected A.4. (1977)146 Controlled diet Normal food 60% of energy from bread Cellulose— Whatman No. 21−32 Spiller et al. (1980)138 Ad lib diet but excluding certain high fiber foods Cellulose— Solka floc® Fleming et al.6) 32 20 (1.3M 7 F.3) 2.4. 23–60 5 M. and zinc and negative balances of all three C.4 (0. (1973)96 Ad lib diet Cellulose (Whatman CFI) + Cellulose 0.5 2. 14 g Basal +Cellulose.4.8 (0. 25−43 Eastwood et al.8 6.1) 1.6) Fecal pH Fecalmutagenic activity negligible on all diets A. and 4.4.3 I 122 (12) 62 (8) 63 (10) I 102 (22) 36 I 55 3. 7) Stool pH fell 208 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed Table 4. (1978)150 Ehle et al. 21−43 Literature Source Diet Fiber Source a Total Number of Subjects in study (sex and age) Hillman et al. 20 F. Various transit methods used (see Van Soest et al. 15 g Basal 24 24 28 + 15 V 28 Fiber Intake (g/day) and Fiber Method — Days on Diet Figures are increases over basal diet period—fiber remaining in stool.4.4 2387_ch4.4.3 and 4.4. (1983)149 Van Soest et al. 13–18 g day + Cellulose. May 6.10) A. [1983]).156 10 M.4. 3RD EDITION Energy (kcal/day) Acid Steroids (mg/day) Neutral Steroids (mg/day) N (g/day) Fat (g/day) . B Pectin and lignin also fed (see Tables 4.24 Wrick et al.7 and 4.4pt1_fm Page 208 Sunday. (1982)151 Heller et al. 2001 7:04 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. (1983)155 Controlled diet Normal food Low fiber Cellulose— Solka floc® baked into bread d c b Study Period Basal + Cellulose.153 (1984)154 Van Soest et al. (1983)144 Ad lib diet Cellulose— α-cellulose from Sigma Fecal Collection Period (days) 7 7 2 2 Number of Subjects Collecting Feces 12 12 10 10 Fecal Weight (g/day) 102 137 133 (21) 208* (30) Moisture (%) (Continued) Effect of Cellulose and Cellulose Derivatives on Fecal Composition 25 33 Fecal Solids (g/day) 5.0 Transit Time (h) and Transit Method 56d 47 40*(4) II 55 (6) Comments Notes Other Data Available Part of a larger study in which coarse and fine bran and cabbage fiber were fed (see Tables 4. (1980)152 Van Soest (1981). Assumes average weight of subjects of 70 kg. Cellulose taken with 150 g apple compote (90 g apple + 60 g syrup).4. 2.4.6.57 Williams and Olmsted Total Number of Subjects in Study (sex and age) 5M 3M Study Period II +16 Basal 7 7 + Konsyl.29.4 10.4pt1_fm Page 209 Sunday.4. 5 g Agar-agar Fecal Collection Period (days) 3 3 3 3 3 3 3 3 3 7 Number of Subjects Collecting Feces 5 5 5 5 5 5 5 5 5 3 Fecal Weight (g/day) 122 (30) 96 96 118 (24) 96 135 (24) 84 68 83 (17) +111a 20 (2) 22 (6) 22 (2) 26 (6) 20 (2) 21 (2) 31 (5) 18(2) 20 (40) Fecal Solids (g/day) Percent Moisture Effect of Plant Gums. Mucilages. 5 g +5 7 7 7 6 Fiber Intake (g/day) and Fiber Method +5 Days on Diet Basal Basal + Karaya. 4. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 209 Energy (kcal/day) Acid Steroids (mg/day) Neutral Steroids (mg/day) N (g/day) Fat (g/day) Transit Time (h) and Transit Method . B A major early study of the laxative effects of plant gums including animal data and extensive in vitro studies 2387_ch4.Literature Source Diet Fiber Source (1936) Ispaghula from Plantago ovata (Konsyl Siblin) Psyllium from Plantago loeflingii (mucilose) Ad lib diet Karaya gum—sterculia urens Bassora gum (Imbicoll) Psyllium seed (back psyllium NF) (1941)7 Gray and Tainter Controlled diet Normal food Agar-agar 9.4.2 3.4.0 8.4. 4.5 Comments Notes Other Data Available Part of a larger study in which 9 other fiber sources were fed (see Tables 4.7 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed Table 4. 5 g +5 7 7 + Psyllium seed. 5 g Basal 7 Basal +5 7 + Imbicoll. May 6.4.2 4. and 4.7) A. and Other Polysaccharides on Fecal Composition 5. 4. carrot.) + Guar.5 2387_ch4. 22–25 Ad lib diet Psyllium (Metamucil) Drasar and Jenkins Literature Source Diet Fiber Source 22 M. Hercules Powder Co. 5g Basal Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed (Continued) Effect of Plant Gums. 20g Basal Cummings et al.9 1.3 and 4.4.8 Comments Notes Other Data Available Part of a larger study in which fiber from bran. 20–38 10 M.4pt1_fm Page 210 Sunday. 3RD EDITION Energy (kcal/day) Acid Steroids (mg/day) Neutral Steroids (mg/day) N (g/day) Fat (g/day) . and apple were studied (see Tables 4.1 3. Hercules Powder Co.8 2. (1978)18 + Guar gum. 2001 7:04 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.) 133 19 M. 35 g (1976) Basal +Psyllium. 12 g Basal Ad lib diets Guar gum (clear gum.Controlled Normal food Guar gum (clear gum.5 0. and Other Polysaccharides on Fecal Composition Transit Time (h) and Transit Method Table 4.8) Guar lowered blood cholesterol but did not produce a detectable change in the fecal microflora E. cabbage.4. 5 g Days on Diet 3 Fecal Collection Period (days) 7 Fiber Intake (g/day) and Fiber Method +5 Number of Subjects Collecting Feces 3 3 3 3 14 14 5 5 5 Fecal Weight (g/day) 120 (20) 139 (17) 204 (21) 100 99 (37) 52 (6) 63 (23) 110 (25) 90 Percent Moisture 74 74 Fecal Solids (g/day) 30 (3) 35 (1) 17 (4) 19 (2) 16 (4) 1.7) Also studied were pectin and banana (see Tables 4.4. May 6. J Mental hospital patients Includes one of the earliest observations of the moderating effect of fiber on blood glucose K 210 + Mucilose. Mucilages.2 and 4. 18 F Total Number of Subjects in Study (sex and age) Block (1947)157 21 21 III 22 39 14 14 7 +30 III +12 7 7 7 7 7 1 1 3 3 7 +5 7 Study Period +Siblin.4. 7 4.1) 1.1 (0. en-ergy.50. 20 g Basal + Ispaghula. May 6.4.0 (0.9 1. none of which changed significantly Great detail of nonstarch polysaccharide intakes and outputs A 2387_ch4.2 92 I 101 4. H.2 7. “double blind parallel repeated measures” design A cellulose-pectin biscuit also studied (see Table 4. 29 F.9) C. (1980)123 Ad lib hospital diet Ispaghula (Fybogel) 2M. 10 g — 44 III 20 Basal + Ispaghula. 2 F. 65–96 Smith et al. 28–31 Spiller et al. N Elderly hospitalized subjects Also studied with bran (see Table 4. 25 g Basal 28 21 21 21 21 5 5 7 7 5 5 15 15 10 10 4 4 37 75 107 58 162 (22) 242* (188) 82 77 10 20 29 19 37 (5) 43* (14) 4.2) Colonic motility (pressures) unchanged Ispaghula and bran equally effective per gram of fiber fed C. H Data also given for apparent digestibilities of N. 2001 7:04 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 211 . (1979)159 Ad lib low fiberrestricted diet Psyllium seed hydrocolloid Controlled diet Normal food Ispaghula husk (Isogel) Prynne and Southgate (1979)158 — +8 +10 + Psyllium powder.2) 104 216 Subjects selected for low stool weight and slow transit. 25–65 18 M.4.4pt1_fm Page 211 Sunday. and ash. fat. F.9 3. 10. M. May 6.5 g 3 3 5 5 5 5 Fecal Collection Period (days) Figures are increases overbasal diet period—fiber remaining in stool. 25 g 21 9 V 0 9 21 +8 +35b 7 Days on Diet — Fiber Intake (g/day) and Fiber Method Basal + Xylan. B. corn bran.5 2387_ch4. fecal pH unchanged A. E.2 Acid Steroids (mg/day) 779 699 540 699 Energy (kcal/day) 424 472 272 372 Comments Notes Other Data Available Serum cholesterol reduced and some changes in breath. (1983)161 Ad lib diet Gum Arabic Fleming and Rodriguez (1982)143 Marthinsen and Fleming (1982)142 (1983)141 5 M.5 0. 30–55 Study Period — 25 + Gum Arabic. B. Controlled semipurified diets Xylan (ICN) Ross et al.6). 0. Total Number of Subjects in Study (sex and age) 5M 5 5 5 5 Number of Subjects Collecting Feces 5 5 Fecal Weight (g/day) 161 147 65 54 139 135 Percent Moisture 73 73 16 14 38 36 Fecal Solids (g/day) 52 37 0.5 g/kg Basal Basal + Karaya. M The gum had no discernable effect on anything 212 Eastwood et al. H. Assumes average weight of 70 kg and also that xylan was 100% dietary fiber.3 0. 4. N Part of a study including cellulose. 3RD EDITION Neutral Steroids (mg/day) N (g/day) .4.4. (1983)160 Ad lib diet Gum Karaya (Sterculia urens from Nargina) Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed (Continued) Effect of Plant Gums. and pectin (see Tables 4.4pt1_fm Page 212 Sunday. and Other Polysaccharides on Fecal Composition Transit Time (h) and Transit Method Table 4. H2 excretion Extensive breakdown of gum in gut A.3. Mucilages.4.6 68 63 71 51 Fat (g/day) 3. and 4. K.9 4.b a 5 M.4. 21-32 Literature Source Diet Fiber Source Fleming et al.4. 2001 7:04 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. extracting with hot ethanol. May 6.6* (0. (1979)163 Controlled eggbased high protein formula diet Cooked oat bran. either plain or toasted Literature Source Diet Fiber Source 3M Total Number of Subjects in Study (sex and age) Williams and Olmsted (1936))9.6 Fecal Weight (g/day) 68 (10) 112* (16) 128* (29) +96a Moisture (%) 77 75 79 Fecal solids (g/day) 14 (1) 28* (3) 29* (2) Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 5. B 2387_ch4. and 4.2 (0. 72.9* (0.6 Transit Time (h) and Transit Method III 28 (10) 31 (7) 27 (6) Fat (g/day) 1. 2001 7:05 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 213 Neutral Steroids (mg/day) .4. 4.4.3) 2.4) 1.5 4.5.0 (0.4.6 M. E Part of a larger study in which 9 other fiber sources fed (see Tables 4.2.0* (0. 28–42 Calloway and Kretsch (1978)162 Kretsch et al.3) N (g/day) 1.4.8) A.2) 1.0 5.4.4.4. 4. 55 g 15 12 15 V 4 15 6 Fiber Intake (g/day) and Fiber Method 12 Days on Diet II +26 6 6 6 7 Fecal Collection Period (days) Effect of Oats and Corn Products on Fecal Composition 6 6 6 3 Number of Subjects Collecting Feces Table 4.29. and drying Study Period Oat branb (toasted) Oat brain (plain) Formula diet Corn germ meal.8 kg No difference between toasted and plain bran for any parameter Principally a study of N balance which was not affected by oat bran Blood lipids unchanged Guatemalan diet also fed (see Table 4.2) Acid Steroids (mg/day) 238 (24) 504* (37) 501 (62) Energy (kcal/day) 98 (16) 184* (27) 189* (41) Comments Notes Other Data Available Subjects average weight.7) A.57 Controlled diet Normal food but excluding fruits and vegetables Corn germ meal prepared by washing for 24 h in warm water.1) 1.4pt2_fm Page 213 Sunday.5* (0. 3RD EDITION Neutral Steroids (mg/day) Transit Time (h) and Transit Method . 24–37 Judd and Truswell (1982)164 Controlled diet Normal food Rolled oats – served as porridge and substituted for flour in bread.4pt2_fm Page 214 Sunday.5 3.4. (1979)114. cakes. data from first control period presented here Blood cholesterol fell significantly E Part of a larger study including wheat bran and soybean hulls (see Table 4.0 III Fat (g/day) 36 30 N (g/day) 3.2 and 4. 2001 7:05 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and biscuits Literature Source Diet Fiber Source Basal + Corn bran. May 6.6 (Continued) Effect of Oats and Corn Products on Fecal Composition 2387_ch4. 4 F. Basal V Basal + Oats.7) 214 Fecal Weight (g/day) Table 4. (1981)165 8 m. 110–160 g 6 M.120 Controlled diet Normal food Dry milled corn bran baked into bread 20 Fiber Intake (g/day) and Fiber Method 23 25c +24 V Days on Diet 10 21 21 28–30 28–30 3 Fecal Collection Period (days) 6 6 12 12 8 10 10 7 7 Number of Subjects Collecting Feces (14) 147 114 125 144* (10) 72 (10) Moisture (%) 74 73 Fecal solids (g/day) 29 34* (2) 35 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 5. 26 g Total Number of Subjects in Study (sex and age) 10 M 19–54 Study Period Munoz et al.1* Acid Steroids (mg/day) 230 372* Energy (kcal/day) 862 230 (68) (33) 282 315 483 (52) 337 (68) Comments Notes Other Data Available Serum cholesterol lowered Study design was control-oatscontrol.4.4 5.35–62 Kirby et al.4. 2) H Part of a study including cellulose.4.3–4.6 g/kg 14 +Corn bran. Assumes average weight of subjects of 70 kg. 20 g 10 14 17 X 43 Basal + Oat bran.6 g/kg/day. B. 21–32 Fleming et al. Oat bran fed at 0.d c b 9 9 0 + 29d Basal + Corn bran. and xylose (see Tables 4. 2001 7:05 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 215 .4pt2_fm Page 215 Sunday. pectin. 0.4.0 819 (61) 354* (28) Constipated subjects also studied with wheat bran which was less effective per gram material fed (see Table 4. K 2387_ch4. Co. Oatmeal diet contained only 2 g more dietary fiber because of substitutions—according to the authors.0 1. (1983)141 Marthinsen and Fleming (1982)142 Fleming and Rodriguez (1983)143 Controlled semipurified diets Corn bran (Quaker Oats) a 11 F. (1982)130 Ad lib low fiber diet Corn bran (G-60 grade Staley Mfg. May 6.9 28* 11 43* (2) 3. M Glucose tolerance unaffected E.) fine particle size Controlled diet Normal food Oat bran—served as muffins or hot cereal 46 14 66 65 2.4. 5 m. 20–40 Graham et al.5) Fecal pH unchanged A. 100 g 3 3 7 7 3 3 5 6 6 8 139* 54 31 (2) 82* (18) 169* (16) Figures are increases over basal diet period—fiber remaining in stool. 29.5.4. wheat bran.7 2387_ch4.4.4. and peas obtained on the open market.1 3. 21 g Carrot.4.0 Comments Notes Other Data Available A Also studied: cellulose.4. 4.4pt2_fm Page 216 Sunday. chopped and washed in warm water for 24 h. 38 g Cotton seed hulls.9 5. then extracted with hot ethanol and dried Alfalfa leaf meal. 27 g Canned peas.6) 216 Moisture (%) Table 4. and sugar beet pulp from Ralston Purina Co. cotton seed hull meal.57 Total Number of Subjects in Study (sex and age) 3M Study Period Alfalfa leaf meal. agar-agar. May 6. 2001 7:05 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. then processed as other materials Controlled diet Normal food.6 2. 25 g Sugar beet pulp.9 3. excluding vegetables and fruit Williams and Olmsted (1936)9. 4.3 4. 27 g Cabbage. and corn germ meal (see Tables 4. 3RD EDITION Energy (kcal/day) Acid Steroids (mg/day) Neutral Steroids (mg/day) N (g/day) Fat (g/day) Transit Time (h) and Transit Method Fecal Solids (g/day) .Literature Source Diet Fiber Source Carrots.2. 35g 6 6 6 6 +18 +18 +15 +12 6 6 Fiber Intake (g/day) and Fiber Method +17 Days on Diet II +13 Fecal Collection Period (days) 7 7 7 7 7 7 3 3 3 3 3 3 Number of Subjects Collecting Feces Effect of Miscellaneous Purified Forms of Fiber on Fecal Composition 104 +28 +67 +90 +16 a +40 Fecal Weight (g/day) Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 6. and 4.4. cabbage. 3 (0.5) 420 (51) 377 (13) 388 (51) 330 (87) 156 (20) 234* (16) 368 (116) Also studied on bran (see Table 4. (1983)24 Cabbage. 26 g Basal 12 28–30 7 12 12 28–30 +23 +SBH.6 2. and apple obtained commercially.4. (1978)18 Stephen and Cummings (1980)25 Controlled diet Basal +Bagasse. 20–38 Cummings et al.10 M.120 Controlled diet Soybean hulls (SBH) and textured vegetable protein (TVP) fed as bread Eastwood et al.0 5.2) E. 2001 7:05 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 217 .5) 6. 10. 26-44 Munoz et al. 30 g Basal 22 Basal 21 21 III 22 40 84 — 27 18 +Cabbage.7* (0.9) 4.4.4pt2_fm Page 217 Sunday. (1979)114. 25–72 McLean Baird (1977)109 Ad lib diet in convent Bagasse (sugar cane residue from Tate and Lyle) made into biscuits (1975)107 Walters et al. K Bran and guar gum also studied (see Tables 4.2 and 4. J 2387_ch4. D.6 3.4.4.6) Plasma cholesterol unchanged but triglycerides fell with SBH C.8 3. May 6. dried.9 (0. E. carrot.8 I 50 (11) 60 (9) 50 (6) 50 (9) 43* (6) II 80 (11) 64* (8) I 47 (6) 37 (3) 4. extracted with hot ethanol and acetone. 26 g Basal 12 7 7 7 7 28–30 V 21 21 22 44 21 42 21 7 7 7 7 Basal +Apple.5 g 20 F. 30 g Basal 19 M.4. then dried and ground 28–30 7 +4 XI 20 +TVP.4.2 and 4. 25 g +Carrot.5) J Blood lipids and fecal microflora unchanged A separate study of bran also reported (see Table 4. 7 3 3 5 5 6 6 6 6 6 6 9 9 161 (26) 68 (9) 128* (16) 92 (10) 102 (21) 117 (7) 189* (16) 141 (20) 203* (29) 88 (9) 143* (16) 88 (6) 140* (10) 75 (1) 80* 77 79* 74 75* 70 76 75 39 (5) 30 (1) 39* (2) 30 (2) 37* (3) 26 (1) 35 (6) 22 (2) 33* (1) 2.2) Also studied on bran and corn bran (see Tables 4. 19–54 9 M. 3 and 4. 3RD EDITION Acid Steroids (mg/day) Neutral Steroids (mg/day) N (g/day) . (1983)144 Lignin — pure Aspen autohydrolyzed from Stake Technology Ltd. May 6.4pt2_fm Page 218 Sunday. 12 g +12 28 III 21 7 — — 21 7 18 28 21 Fiber Intake (g/day) and Fiber Method 33 Days on Diet Basal + Boiled potato.4) Stool pH unchanged No significant effect on fecal composition or blood lipids A. Ad lib diet Potato concentrate purified by commercial company and either air or rollerdried or boiled. fed with either milk or soup Total Number of Subjects in Study (sex and age) 10 M.4.4. 2 F.1 Fat (g/day) 4.7 (0.7) 6. 20 g + Air-dried potato.8) 4.7b Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 33 (5) 48 (8) 42 (6) 35 (7) Transit Time (h) and Transit Method 4.4. 21–43 Study Period 28 + Lignin.Literature Source Diet Fiber Source Hillman et al. 2001 7:05 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 20 g Basal Fecal Collection Period (days) 2 2 7 7 7 7 7 Number of Subjects Collecting Feces 10 10 5 5 7 7 7 Fecal Weight (g/day) 139 (17) 177 (25) 231 (51) 215 (44) 170 (32) 207 (42) 204 (50) 78 (2) 76 (2) 74 (1) 75 (2) 78 (3) Moisture (%) 47 (7) 47 (6) 42 (6) 47 (6) 39 (7) 32 (10) II 50 (5) 40 (5) 3.7 2.5 (0. 20 g Basal + Roller-dried potato.9 (1.7 (Continued) Effect of Miscellaneous Purified Forms of Fiber on Fecal Composition 2387_ch4.3) 4.2 (0.8) 3.7) Energy (kcal/day) 340 (59) 340 (84) 392 (50) 392 (47) 352 (52) Comments Notes Other Data Available Cellulose and pectin also studied (see Tables 4. E 218 Fecal Solids (g/day) Table 4.2 3.5 (0. 20–30 Schweizer et al. 2001 7:05 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 219 . 26. 20–30 Tsai et al.24 Wrick et al. (1983)167 Controlled diet Normal food but low fiber Soy polysaccharidec a 2 M.2 (0.. D. 4 F. XI 14 Basal + Purified soya pulp. K Never-dried soya increased fecal deoxycholate while purified soya increased low-density lipoprotein cholesterol and altered glucose tolerance A. B Fasting blood glucose fell but lipids unchanged C.7* (1. H. E. May 6.4 and 4.4pt2_fm Page 219 Sunday. purified soya fiber obtained from soya flour by extraction 12 12 14 14 6 6 6 102 110 140 (11) 176* (16) 178* (28) 129 (16) 153* (14) 78 76 77 77 77 25 24 34 (3) 39 (4) 39* (4) 34* (4) 28 2. Soy polysaccharides “a fiber-rich product purified from soy bean cotyledon.4.8) 5.0) 764 (66) 735 (49) 726 (28) 59 (11) I 62 (4) 44 (9) 109 (8) 118 (16) 215 (17) 242 (19) 294 (23) 263 (32) 279 (23) 12 g cell wall material daily at first but increased later up to 24 g/day Bran and cellulose also fed (see Tables 4. D.4 1. 54 g Basal 7 7 7 7 7 Stool weights are increased over basal period—fiber remaining. E.5 g + Never-dried soya pulp. K 2387_ch4.156 14 M. (1983)166 Ad lib diet Soya: a never-dried pulp obtained as a by-product of milk production.3 56d 58 246 (59) 289 (49) 1. 1983). 25 g 17 21 21 14 — 34 33 V.10) A.9 (0.6 (0. (1983)149.156 Controlled diet Normal food Cabbage — ethanol extracted and incorporated into bread d c b 24 24 V — — Basal + Cabbage 17 — + Soy. Assumes fiber intake increased by about 10 g.4.2) III 26 (3) 29 (3) 7.2) 1. C.” Various transit methods used (see Van Soest et al. high fiber High fat. especially fat and fiber.Controlled diets Normal prison foods Antonis and Bersohn (1962)169. + Fruit.2* (0. 29 black 2 M.170 Controlled diet Normal food Fruit-plums. low fiber Low fat.1* (0.2) Noted greater digestibility of fiber than in healthy subjects A A chance observation in a study of mineral balance on various breads Fruit had no effect C 220 6 Cowgill and Sullivan (1933)59 Controlled diet Normal food Fruit and vegetables 7 6M Literature Source Diet Fiber Source Days on Diet I 30–38a Total Number of Subjects in Study (sex and age) Basal Effect of Foods Containing Fiber on Fecal Composition Number of Subjects Collecting Feces Table 4. low fiber High fat.4 (0. pears.2) 2. and damsons McCance and Widdowson (1942)168 58 M: 29 white.4. 3RD EDITION Neutral Steroids (mg/day) N (g/day) Transit Time (h) and Transit Method Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed Fecal Solids (g/day) Moisture (%) .4. high fiber High fat. 2 F.4pt2_fm Page 220 Sunday. May 6.3) 2.1) 4. 2001 7:05 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.1) 2. blood lipids.8 2387_ch4.2 Acid Steroids (mg/day) (35) 479 (10) 324* (31) 590 (62) 310* (27) 537 Energy (kcal/day) (28) 463 (29) 450 (38) 517 (31) 521 (31) 459 Comments Notes Other Data Available Prisoners in South Africa A very large study of the effect of diet. greengages.1) 1. on fecal lipids. 27–41 Study Period high fiber High fat. 31–183 g Basal + Fruit and vegetables +Fruit and vegetables and bran 7 108–113 22–29 22–29 15–17 4 15 22–29 4 15 22–29b I 15 14–28 7 Fiber Intake (g/day) and Fiber Method 90 Fecal Collection Period (days) 7 7 7 7 7 7 7 7 7 7 18 Wh 21 Bl 21 Bl 22 Wh 22 Wh 4 4 3 5 Fecal Weight (g/day) (13) 236 (13) 85* (6) 259 (16) 99* (6) 209 156 155 78 (15) 165a (27) 193a (10) Fat (g/day) (0. and bile acids E Constipated subjects Also studied effect of various brans (see Table 4.1 (0. 23 M.1) 1.1) 1.7* (0. J A large and detailed study of the energy value of the diet as influenced by fiber A 2387_ch4. low fiber 4 5 5 12 EW 12 EW 11 EM 11 EM 14 YW 14YW 14 YW 12 YM c 12 YM 13 Bl 13 Bl 18 Wh 121 (15) 170 (23) 208 (46) 79 (11) 140* (12) 60 (5) 130* (7) 181* (8) 91* (5) 163* (8) 47 (3) 82 (6) 100* (9) 240 (10) 97* (7) 79 75 77 77 78 73 68 76 75 18 (1.1 (0.9 (0.3) 1.1) 1.1) 1.1) 2.2* (0.7 Diet 1 7 10−14 15–17 4 7 7 15–17 15 7 10–14 15–17 4 Low fat.2 4.1) 2.4* (0.4.2 Diet 2 7 6.2* (0. 1000 g IV Basal 7 7 7 7 10−14 20.0 (0.3) 3. 26 W.3) 3.2) 6.1) 2.2* (0.4.0 (0.1) 2.4 5. low fiber Low fat.3 5.4 (0. 18-78 10 M.4 (0.3 5.1 3.4pt2_fm Page 221 Sunday.3) 5.4) 4.7) 15 (0.6 (0.0) 6. 1000 g + Plantain.8 (0. May 6.3) 4.1) 24* (1.3* (0. 22–25 Southgate and Durnin (1970)28 Controlled diet normal food Diet 1: no fruit or vegetables except potato Diet 2: contains fruit and vegetables and whole meal bread Diet 3: contains larger amounts of fruit and vegetables Drasar and Jenkins (1976)133 Ad lib diet Bananas and Plantain banana 14 14 14 +34 +58 + Banana. 2001 7:05 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 221 .1) 297* (25) 543 (51) 277* (24) 103 (5) 164* (8) 90 (5) 150* (6) 210* (7) 127* (5) 83 (4) 117 (6) 206* (6) 320* (25) 393 (18) 316* (20) Also studied pectin and guar (see Tables 4.1) 1.6 Diet 1 7 31.5 1.0) 40* (1.9 Diet 2 7 10−14 7.4* (0.3 and 4.8 (0.8) 21 (1) 1.2 Diet 1 10−14 21.2) 0.1 (0.3 Diet 2 7 10−14 9.5* (0.4 Diet 1 7 10−14 28.5 (0.6* (0.0* (0. high fiber Low fat.5) Blood lipids and fecal microflora unaffected E.5 Diet 2 7 7 10–14 III 9.9 Diet 3 10−14 16.0) 32* (1.2) 4.0) 38* (1) 15 (0.8) 28* (1.1) 1.4 8. E No change in blood lipids E. 860 g Basald Study Period Beyer and Flynn (1978)172 Controlled diet Normal food Mixed sources Ad lib diet Cooked potato Total Number of Subjects in Study (sex and age) Flynn. 3RD EDITION Neutral Steroids (mg/day) .6 11. May 6. 20–56 Literature Source Diet Fiber Source Formula diet 6 M.5 6.4* 1.4. 28-42 Calloway and Kretsch (1978)162 Kretsch et al.3) 1.7) 1.3) 3.8 (Continued) Effect of Foods Containing Fiber on Fecal Composition 2387_ch4.1* N (g/day) (0. F 222 Fecal Weight (g/day) Table 4.6 Number of Subjects Collecting Feces (89) 68 (26) 327* 51 157 125 (15) 294* (24) 149 (20) 249* (39) Moisture (%) 81 79 Fecal Solids (g/day) (10) 14 (3. 21–29 Basald (low fiber) +Potato. Beirn.9 3. 2001 7:05 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.3 (+8.1) 1.2) 62* 15 37 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 2. (1978)162 Controlled eggbased high protein formula diet Guatemalan diet Low fiber Increased fiber 6 M.48 M. 860 g + Potato.9* 2.1 8.6 Transit Time (h) and Transit Method III 48 12 62 (18) 35* (24) I 46 (20) 33 (21) Fat (g/day) (0.0 (0.2 (0.6)e 5 5 18 5 5 — Fecal Collection Period (days) 3.2* Acid Steroids (mg/day) (160) 238 (58) 502* Energy (kcal/day) (64) 98 (40) 348* 63 140* Comments Notes Other Data Available Also studied with oat bran (see Table 4.4pt2_fm Page 222 Sunday.6)e Days on Diet II 5.1 2.5f 19.5 7.6 (+8. and Burkitt (1977)171 — Fiber Intake (g/day) and Fiber Method 15 15 V 0 89 5 5 I 1.7 3 3 6 6 6 6 18 6.6) A.4. H Serum cholesterol fell Breath hydrogen excretion increased C. May 6. and banana 23 M. cheese. pumpkin. squash.1) 3. H.2 (0.8 II 55 (3) 37* (2) 55 (14) I 72 (12) 1.5 5.7) 2.5 (0. E.Controlled diet Normal food Mixed fiber sources et al. bread.7 (0. 20–27 5 High fiber Low fiber + Carrot Basal 33 21 21 VIII.1) 2.5* (0. 2001 7:05 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 223 . 23 F. E.4pt2_fm Page 223 Sunday. (1979) 174 Stasse-Wolthuis Ad lib diet Raw carrot Robertson et al. IX 12 IIIg 21 21 +6 V 21 (14–33) 3–5 3–5 7 7 43 43 5 5 69 (8) 184* (11) 177* (33) 142 (37) 76 74 76 75 18 (2) 44* (7) 42 (6) 35 (6) 3. (1979)173 Guatemalan diet contained mainly black beans and corn tortilla together with rice.4) 645 (96) 602 (73) 389 (33) 271 (29) Diets were fed at either high or low cholesterol (about 200 and 600 mg/day) Serum cholesterol fell in both groups with fiber but results confounded by changes in fat intake C. M 2387_ch4. 5) 2.9 (0.3 (0.4 (0.4.6* (0.4) 2.4.4* (0. 18–28 12 M. 3RD EDITION Neutral Steroids (mg/day) .8 (0. (1978)175–178 Controlled diet Low fiber with fruit and vegetable juices then with whole fruit and vegetables (no cereals) Kelsay et al. 22 F. D Also studied with pectin and wheat bran (see Tables 4. (1981)179 Fiber Intake (g/day) and Fiber Method 26 26 21 V 4 24 V 2 Low fiber High fiber Low fiber 35 43 35 + Fruit and vegetables. (1979)124–126 Controlled diet Normal food Fruit and vegetables Kelsay et al.4pt2_fm Page 224 Sunday.0 2.1) 2.4. 1065 g Study Period 18 g Days on Diet Low fiber Fecal Collection Period (days) 7 7 7 7 7 12 12 12 15 15 Number of Subjects Collecting Feces 87 (5) 89 (9) 209* (9) 89 (10) 138* Moisture (%) 74 75 73 Fecal Solids (g/day) 23 (1) 23 (2) 52* (2) 32* 23 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 6.5) 6.0 Transit Time (h) and Transit Method III 30 (3) III 52 (4) 38* (4) I 66 (10) 53* Fat (g/day) 4. May 6. E.3) Serum cholesterol changes not significant No change in blood pressure C.8 (Continued) Effect of foods Containing Fiber on Fecal Composition 2387_ch4.4) 4.2 (0.2) Acid Steroids (mg/day) 888 682 Energy (kcal/day) 113 (6) 255* 117 290 364 Comments Notes Other Data Available Latin square design Blood pressure unchanged except in those in whom diastolic was 80 mm + Mineral balances became “lower” A.9 N (g/day) 1.Total Number of Subjects in Study (sex and age) 40 M. 2001 7:05 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. C. 37–58 12 M.2 and 4. H 224 Fecal Weight (g/day) Table 4.1) 1. 35–49 Literature Source Diet Fiber Source Stasse-Wolthuis et al. 1) 2. EM.5 g/day).9 III 53 (8) 45 (6) 27 (2) 27 (2) 31 (4) Milligrams of crude fiber per kilogram body weight per day.6 g/day) and those consuming less (average 3.4) 6.5 4. YW. young women.5).3 (0. Dietary fiber values calculated from food tables (IV). 21–36 + Fruit and vegetables + Fruit and vegetables + Fruit and vegetables Controlled diet Low fiber with fruit and vegetable juice then increasing amounts of fruit and vegetables 5. young men. and copper balance unchanged Zinc balance “decreased” A.1) 2. + Haricot beans. Dietary fiber equivalent.g f e d c b a 21 14 26 IV 22 14 21 19 49 21 10 14 14 7 7 7 6–8 6–8 12 12 12 115 (15) 150* (14) 127* (8) 171* (12) 274* (26) 78 75 75 38 31 32* (2) 42* (2) 50* (3) 1. YM. EW.4) 6.6 (0. magnesium. May 6. elderly men.4.0 (0. 2001 7:05 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 225 .2 (0. (1982)180 Ad lib diet with substitution of meat and vegetables by canned baked beans (Haricot) 8 F.4pt2_fm Page 225 Sunday.2) 162* (10) 209* (11) 255* (16) All subjects noted marked increase in flatulence Noticeable effect of menstruation on transit time Blood pressure unchanged Calcium. elderly women.4 (0. A variety of fiber metods used (see paper).8 (0. 450 g Basal Leeds et al. Weeks on diet (data are from Table 4.3 5.8 2. D 2387_ch4. C.1) 1. Subjects divided into those consuming more than 5 g crude fiber per day (average 5. 5% MO 35% MC 61. 5 with lipid disorders.4 5. 2001 7:05 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.4pt2_fm Page 226 Sunday.2 3.1 Transit Time (h) and Transit Method I 60 (9) 35* (8) Neutral Steroids (mg/day) 505 (41) 656 (75) Acid Steroids (mg/day) 194 (23) 266 (47) Comments Notes Other Data Available Hospitalized subjects. and magnesium oxide (MO) 3 F.4 g methylcellulose and 0. 5 M.2* +3 +41/2 Days on Diet — 6 6 — 8 8 8 Literature Source Diet Fiber Source Total Number of Subjects in Study (sex and age) 8 Fecal Weight (g/day) 177 (46) 239* (35) 231 177 179 160 145 127 184 213 163 Moisture (%) 77 82 84 83 81 81 78 81 81 82 80 Fecal Solids (g/day) 32 (2) 54* (9) 36 30 35 31 31 24 35 38 32 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 1. May 6.4. (1952)61 Effects of Mixed Sources of Fiber on Fecal Composition Number of Subjects Collecting Feces Table 4.0 11.4) Also studied with pure methylcellulose (see Table 4.6 7 11.5 9 V 0 60 7 7 7−14 7−14 7−14 7−14 7−14 7−14 7−14 7−14 7 7 7 Fecal Collection Period (days) 6.9 2387_ch4.4. (1977)181 Controlled formula diets “Constant” diets.Raymond et al.2 5.4 6. 3 normals Magnesium oxide providing significant additional laxation Also studied with pure methylcellulose (see Table 4. 19−67 Low cholesterol basal + Fiber 35% MC 65% B 5g 10 g 3.7 8. bran (B).5% B 10 g 10% MO 45% MC 45% B 10 g 10% MO 90% MC 10 g Basal Tainter (1943)60 — +6 Tablets +9 Tablets Study Period Ad lib diet Tablets containing 0.4.1 g psyllium Basal Fiber Intake (g/day) and Fiber Method 7−14 7−14 7−14 7−14 28 28 6. normal foods Various combinations of methylcellulose (MC).4) 226 Berberian et al.6 7. 3RD EDITION Energy (kcal/day) N (g/day) Fat (g/day) . (1982)121 Ad lib low fiber normal food diet Low fiber bar containing mainly oats and rice. 25−65 Spiller et al. (1979)159 Ad lib but low fiberrestricted diets Cellulose-pectin biscuit Fiber muffins containing corn pericarp.9 (0. N No decrease in plasma cholesterol E 2387_ch4. carrots.5) C. “Doubleblind parallel repeated measures” design Also studied with psyllium (see Table 4. pectin. peanut butter. hydroxyethylcellulose. 50. wheat bran.4. oats. May 6. 2001 7:05 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 227 . 24−57 Spiller et al.1) 1.1) 1189 (84) 1152 (57) 423 (122) 401 (89) Considerable individual variation in responses noted Subjects selected for low stool weight and slow transit.1 (0. prunes.4pt2_fm Page 227 Sunday. 28 60 21 + Cellulosepectin 28 +2 +10 Low fiber High fiber 28 28 Basal V 21 +20 28 0 Basal High cholesterol basal + Fiber * Assumes bran about 44% fiber. soybean hulls.22. and pectin Either high (1000 mg) or low (<50 mg) cholesterol 5 5 5 7 7 7 7 11 11 11 16 16 6 6 100 (16) 114 (18) 146* (25) 106 62 192 (41) 286* (21) 77 89 31 17 22 (2) 66* (4) 2. H. etc.6 I 77 (12) 70 (12) 70 (14) 69 I 95 66 (11) 46* (9) 0. rice. High fiber bar containing soy and corn bran.2 1. 20 g III 10 Fiber Intake (g/day) and Fiber Method Basal White bread (530−630 g) Medium whole meal Fine whole meal Days on Diet 7 7 7 7 7 7 Number of Subjects Collecting Feces 21 21 21 6 6 6 Fecal Weight (g/day) 199 219 140 232* 283* 62 Fecal Solids (g/day) 69* 69* 18 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 6.0 7 14 14 +10 +9 + Coarse bran.5* 4.Total Number of Subjects in Study (sex and age) 6 9 M.0 Fat (g/day) 4.4 Energy (kcal/day) 317* 325* 99 Comments Notes Other Data Available Coarse bran increased stool weight significantly more than fine bran A study of the digestibility of the nutrients in bread Fineness of grinding made no significant difference to energy or nitrogen utilization but medium flour produced significantly greater stool bulk than fine flour 228 Fecal Collection Period (days) Table 4. 12 F Literature Source Diet Fiber Source Macrae et al.7 N (g/day) 2.3* 2.4. mainly bread (74% of energy intake) White bread and whole meal bread made from either medium or fineground flour Brodribb and Groves (1978)183 Ad lib diet with either coarse bran or same bran finely milled Study Period 10 10 I 0. 20 g + Fine bran.4pt2_fm Page 228 Sunday.9 11. 2001 7:05 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.7 12. (1942)182 Semicontrolled diet. 3RD EDITION Acid Steroids (mg/day) Neutral Steroids (mg/day) Transit Time (h) and Transit Method Moisture (%) .10 Effects of Particle Size on Fecal Composition 2387_ch4.6 8. May 6.9* 2.3* 1. (1982).6 2. 24b Smith et al.4 and 4. (1981)184 Ad lib diet Canadian red spring wheat bran (CRSW) or French soft wheat bran (FSW) in either coarse or fine form a 24 Heller et al.4. N Coarse brans both produced significantly greater effects on bowel habit Effects on motility not great F Also studied with cabbage and cellulose (see Tables 4.1 I 56 (11) 51 (12) 61 (6) 21* (9) 54 (8) 41 (5) 48 (8) 20* (7) I 44 62 1.7 1. and faster transit than fine bran.128 (1983)129 Controlled diet Normal food Breads containing coarse and fine brans 28 20 20 V 22 22 7 — +6.6 3. high moisture content. Patients with diverticular disease.9 28 7 — +59 28 +7. D. E. dietary changes during the study made comparisons with basal diet difficult THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION b Coarse bran Fine bran + Coarse FSW. C.7) Coarse bran produced greater fecal bulk. (1980)152 Controlled diet Normal food AACC white wheat bran fed whole and after fine grinding. May 6.5 3. in bread 2387_ch4. B.6 56 37 a 327 315 (See also Table 4. 20 g Basal + Coarse CRSW Basal + Fine CRSW.12 M. 23 Van Dokkum et al.5 7 4 4 7 7 7 7 7 7 7 7 4 4 4 12 4 6 6 6 6 6 6 6 6 8 8 8 126 102* 80 (6) 92 (11) 96 (6) 123* (11) 81 (11) 102 (6) 68 (9) 106* (6) 108 123 72 74 33 29 29 31 5. fiber digestibility less in coarse bran. 20 g Basal — 7 III — 28 14 +12 +58 14 +12 Coarse bran. 32 g Fine bran Basal 14 V Basal Various transit methods used (see papers).2) Fecal weight significantly lower with fine bran than coarse bran A. 2001 7:05 PM 229 .4.4.4pt2_fm Page 229 Sunday. 20 g + Fine FSW. 7 1. fruit. and whole meal bread Findlay et al.0 Acid Steroids (mg/day) 241 (85) 273 Comments Notes Other Data Available Similar value for transit by both methods Increased abdominal discomfort noted in early weeks Study included a control group of healthy subjects (see Table 4. 20 g Basal III — +12 4 4 35 9 a Fiber Intake (g/day) and Fiber Method 7 Days on Diet — Fecal Collection Period (days) 6 6 7 7 Number of Subjects Collecting Feces 21 21 7 7 Fecal Weight (g/day) 96 176* 101 (19) 84 Moisture (%) 77 78 74 70 Fecal Solids (g/day) 20 34 24 (3) 23 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 6. G 230 Fat (g/day) Table 4.4.4pt2_fm Page 230 Sunday.0 2.11 Effects of Fiber on Fecal Composition in Patients With Diverticular Disease (See Also Table 4. IV 44 37* 58* (8) I.4. 186 Out-patient semicontrolled diet Lower residue than with addition of 40 g bran. 3RD EDITION Energy (kcal/day) Neutral Steroids (mg/day) N (g/day) . F.9 I. 10 F 30−84 7 Study Period Low fiber High fiber + Bran.2) Effect of bran on fecal weight only one-third that seen in controls Intraluminal pressure in response to food reduced by bran.10) 2387_ch4. May 6.4. (1974)98 Ad lib diet Coarse bran Total Number of Subjects in Study (sex and age) 11 M. IV 93 Transit Time (h) and Transit Method 2.Literature Source Diet Fiber Source Parks (1974)185. vegetables. 2001 7:05 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 4 2.2 4.8 206 218 145 226 268 164 C. 20–40 ml 28 28 +20 28 +7 +9 28 Bran tablets (9) Basal + Bran. 7g Basal Lactulose.3 48 40 62 72 I.7 2. 30 F.4pt2_fm Page 231 Sunday.2 2. 25−85 Taylor and Duthie (1976)188 Ad lib diet High fiber diet Normacol (an ispaghula derivative) Bran tablets Brodribb and Humphreys (1976) 187 Ad lib diet Coarse bran Basal + Ispaghula. H Bran reduced postprandial motility while ispaghula increased basal pressures Bran was the most effective of the three treatments Colonic motility in response to eating reduced Patients had lower basal fiber intakes than health controls F. F. May 6. IV 88 50* I 97 (7) 76* (7) 72* (11) 56* (4) I 3. 32−84 Eastwood et al.4 3. 20 g 28 Normacol 14 14 10 10 7 7 7 7 10 5 5 20 40 40 7 7 7 7 5 5 5 28 4 High fiber diet 8b 5 ~8 +10 4 Basal + Bran. 2001 7:05 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 231 . (1978)189 Ad lib diet Coarse bran Ispaghula (Fybogel) Lactulose 10 M.8 2. K 2387_ch4. 24 g Basal 95 160* 75 108 82 103 79 (7) 102 (16) 105* (13) 121* (7) 89* 66 3.20 31. 4pt2_fm Page 232 Sunday. 11 F. 58 patients divided into 3 groups. After 8-month treatment. L 232 Tarpila et al.11 (Continued) Effects of Fiber on Fecal Composition in Patients With Diverticular Disease (See Also Table 4.7) 4.4.2 (0. 2001 7:05 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. patients showed satisfactory clinical response.2 Bran Ispaghula — Fiber Intake (g/day) and Fiber Method Placebo 12 Months control High fiber 6 Months control High fiber Time zero control High fiber Days on Diet 4 4 d 4 c 7 7 57 57 57 11 3 7 11 11 3 3 11 3 11 3 Fecal Collection Period (days) 11 Number of Subjects Collecting Feces 3 Fecal Weight (g/day) 119 (6) 137* (7) 161 (8) 167 (20) 215 (28) 155 (19) 272* (24) 179 (35) 265 (44) 33 (4) 38 (4) 29 (2) 55* (5) 37 (6) 47 (7) Fecal Solids (g/day) Moisture (%) Assumes bran about 44% fiber.2 (0. 35−64 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 6.5 22.3 (1. 3RD EDITION Energy (kcal/day) . May 6.9 Transit Time (h) and Transit Method II 50 (4) 45 (4) 47 (4) Fat (g/day) 5.1) 4. each of which took all 3 treatments in random order.0 (0. Four months on each diet.10) 2387_ch4.3 3.4) 5. (1981)191 Ad lib diet Bran crisp bread Ispaghula (Fybogel) Placebo Total Number of Subjects in Study (sex and age) 11 M.2 24. No consistent cholesterollowering effect E. Some 22 patients allocated to control or high fiber diet and followed for 12 months.4) 5.9) Neutral Steroids (mg/day) 733 (106) 816 (133) 697 (93) 883 (147) 709 (91) 541 (46) Acid Steroids (mg/day) 330 (48) 475 (82) 320 (49) 384 (61) 346 (89) 240 (39) Comments Notes Other Data Available Fiber supplement conferred no benefit on symptoms but did relieve constipation No effect on biliary cholesterol saturation Biliary deoxycholate decreased by bran.9* (1.5) 6. (1978)190 Ad lib diet Bran rusks 6 per day N (g/day) Table 4. 22 M Literature Source Diet Fiber Source Ornstein et al.5 (1.d c b a Study Period III 17.4. 4) 20 (1) 27a (1. (1984)192 Controlled diet Wheat bran Sorghum barn Fecal Collection Period (days) Table 4.4) 74 (5. (1987)195 Ad lib diet Wheat bran 11 V 8.9) II 31 (2) 30 (1) 26a (1) Fat (g/day) 2.5 Ad lib 5F 2M 59−76 Marlett et al.12 Effects of Cereal Products on Fecal Composition Number of Subjects Collecting Feces 6 6 6 7 7 28 28 28 10 10 10 Fecal Weight (g/day) 44 (12) 137 (30) 92 (27) 122 (44) 170a (52) 74 (5) 80 (4.3) Neutral Steroids (mg/day) 640 (60) 640 (48) 720 (84) Acid Steroids (mg/day) 266 (31) 243 (23) 232 (19) Comments Notes Other Data Available Study of fecal flora and enzymes.9 Wheat Total Number of Subjects in Study (sex and age) Ad lib + 2. (1986)193 Ad lib diet White or Whole meal bread 21 21 3 7 14 M 14 F 69 (50−82) Literature Source Diet Fiber Source IV 13−14 + 7.2) 3.4. 2001 7:07 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 233 Energy (kcal/day) N (g/day) . May 6.5 21 X 20 Study Period Sorghum Basal Fiber Intake (g/day) and Fiber Method 10 M 23 (22−24) Days on Diet Fedail et al.9 + Bran Mallet et al.4 3. (1986)194 Ad lib diet AACC wheat bran 182 22−23 + Whole meal bread 5 5 5 9 9 7 7 182 12−14 2 3 + White bread Eastwood et al.5 14.0 Transit Time (h) and Transit Method I 80 (5.3) 3.4 10. J.8) 101a (5.9 3.5 (0.3) Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 4.1 (0.4pt3_fm Page 233 Sunday.14) g fecal weight/g fiber based on average of 3 control periods Dietary compliance study Long-term study in the community Sudan medical students Symptom diary Stool consistency 2387_ch4.4.8) 76 (6.9) 138 (14) 173 (15) 221a (31) Moisture (%) 74 76 73 75 72 Fecal Solids (g/day) 29 (5) 44a (6) 21 (1.6 (0. Ammonia Also pectin (see Table 4.Ad lib +13 Bran 21 21 21 IV 16−21 Ad lib 3M 3F 22−26 22 20. 5) 25a (2. (1986)201 Semi-controlled Wheat bread Literature Source Diet Fiber Source 6F 23−27 8M 7F Number of Subjects Collecting Feces 24 5 5 12 12 12 12 12 36 6 6 15 15 155 (18) 95 77 79 (8) 135a (9) 73 (6) 95a (10) 139a (12) 212 (17) 144 (17) 329a (39) 151 (18) 228 (21) Moisture (%) 74 65 62 75 75 77 76 74 73 77 75 Fecal Solids (g/day) 40 (4.4.2) 4.4.3) 47a (3.1) 33 29 20 (1) 34a (2) 20 (1. (1988)205 Semi-controlled AACC Bran Fiber Intake (g/day) and Fiber Method IV.0 2.9 4.7 7 7 7 5 5 5 5 5 5 2 28 + Bread + 11g 2 Study Period + 17.9 (0. (1988)206 Controlled diet Wheat bran Melcher et al. XVI 15 (3) + 5. (1991)207 Ad lib 21 40 Bran XVI + 8.5 7. 3RD EDITION .1 Stevens et al. (1988)198 Controlled diet Wheat/Rye Total Number of Subjects in Study (sex and age) Reddy et al. May 6.8 3. K Also psyllium (Table 4.0 Transit Time (h) and Transit Method II I-V 70 (6.2 (0.14) Stool consistency Dose response study H.4 (0.2) 2.3) Neutral Steroids (mg/day) 641 (86) 568 (72) 550 (93) 492 (65) Acid Steroids (mg/day) 58 (7.2) I 77 (8) 58 (5) 58 (10) 50 (5) Fat (g/day) 2.12 (Continued) Effects of Cereal Products on Fecal Composition 2387_ch4.2) N (g/day) 1.6) 49a (3.4pt3_fm Page 234 Sunday.3 Low 21 21 XIV. and X 19.2) 38 (4) 52 (5) Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 2.9 3.9) 80 (13) 96 (14) 92 (9) Energy (kcal/day) 144 (17) 307 (41) Comments Notes Other Data Available Consistency M Letter 2 diet periods at 1 year interval A.7 (0.49 Basal + Bran 5 14 M 10 F Villaume et al. XV.5 13 13 13 D B V. XVI 21 Basal 12 F 29 22−38 7 14 13 + 28. (1987)196 Ad lib Mixed grain bread Fecal Weight (g/day) Table 4. L Study of energy metabolism Body weight A Fecal mutagenesis pH 234 Wisker et al.7 High Basal 48. 2001 7:07 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.8 6.2) 34a (2.8 18−32 Spiller et al. XIII Fecal Collection Period (days) Ad lib Days on Diet C 36 F 19.8 7 V. 4) 1.2) 28 (0.3 (0. E. (1984)209 Controlled diet Oats (and Beans) Controlled diet Miyoshi et al.0 28.2 19.2) 10.1 V 15. (1987)208 5M 20.5 14 14 8 14 14 14 14 14 14 13 5. N Blood urea Also beans (see Table 4.5 (0.3) 5.5 27 (0.1) 719 (140) 829 (9) 109 (37) 180 (43) 278 (30) 140 (12) 249a (34) 106 (13) A.9) 26 (5.6a (2.3) III 24a (1. 2001 7:07 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 235 .8 30.2 5M 21.4a (0.0 6.4 34−66 20−36 Ad lib Fiber-One (Wheat and corn bran) 11.29 18−61 20 M Ad lib diet Bran Flakes and All Bran Anderson et al.1) 2.1) 2.4pt3_fm Page 235 Sunday.4 All Bran 30 60 90 Polished rice Brown rice Brown rice Polished rice 47 Oats 14.2) III 12a (1.0 2. K Dose-response study Fecal weight on control periods varied (see original paper) 2387_ch4.6 2.2) 2.5 7. N Blood Urea Proteins C. May 6.13) Hypercholesterolemic volunteers E. (1987)210 34 M 39 F Jenkins et al.5 (0.0 (0.6 XVI + 240 Bran Flakes.3) 2. (1986)211 Controlled diet Rice Miyoshi et al.3) 27 (4) 42a (4) 74 (4. H.5 (0. E. 30 g 60 Ad lib + Fiber-One 3 3 3 3 3 3 3 3 13 5 5 5 5 24 238 (20) 154 (22) a 192a (23) 125 (19) 134 (27) 191a (20) 138 (11) 168 (7) 212 (14) 124 (12) 136 (13) 137 (5) 287a (21) 81 77 73 80 78 80 74 54 (4) 29 (3) a 51a (9) 25 (1.7 5.4.2) 2.2 V 19 Control XIII 14 9. 1) 1. 3RD EDITION Energy (kcal/day) Acid Steroids (mg/day) Neutral Steroids (mg/day) Transit Time (h) and Transit Method .8 7.4) Fecal Solids (g/day) 26 26 24 44 (4. (1986)212 Controlled diet Rice (Agar) Fecal Weight (g/day) Table 4.1) 1.8 (0.Literature Source Diet Fiber Source a 6 26−32 10 10 Corn Basal 10 + 4.3) 117 (4. fecal cholesterol C.4pt3_fm Page 236 Sunday.7) 7.5 8. May 6.0 (0. 2001 7:07 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.12 (Continued) Effects of Cereal Products on Fecal Composition 2387_ch4.8 (0.7 (0. A 236 Kaneko et al. normal protein white rice D: High fiber. Sugawara et al. normal protein brown rice pH.7 (0.3) Comments Notes Other Data Available pH Fecal ammonia and enzymes J Diets A: High fiber.4) 19 (1.8) 78 (1.2) N (g/day) 1. XVI Basal Significantly different from control.9) 41 (2.3) 79 (1.5) Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 2.5) 128 (7.4) 126 (14) 242 (15) Moisture (%) 79 (1.2 D 5 5 11.4.5) 6.6 Fat (g/day) 21 (3.5) 25 (2. low protein brown rice B: Semi-purified low protein agar C: Low fiber.0) 276 (27) 120 (9.6) 24 (1.8 C V 18.3 (0.2 5 Total Number of Subjects in Study (sex and age) B Study Period 18−20 5 Fiber Intake (g/day) and Fiber Method A Days on Diet 5F Fecal Collection Period (days) 6 6 6 5 5 5 5 Number of Subjects Collecting Feces 6 6 6 5 5 5 5 115 (4. (1991)213 Controlled diet Corn residue 16.2 V.3) 0. Fecal consistency See also Reference 216 2387_ch4.1 1.13 Effect of Legumes on Fecal Composition Number of Subjects Collecting Feces 6 6 Fecal Weight (g/day) 172 (12) 198a (12) 132 (23) 140 (14) 196 (23) 216 (31) 67 (5) 115 (13) 100 (10) 150 (20) 145 (13) Moisture (%) 78 78 77 80 86 87 Fecal Solids (g/day) 38 (21) 44 (12) 31 (7) 32 (3) 25 (5. XVII Fecal Collection Period (days) Ad lib Enrich 30 g soya Ensure 30 g soya Ensure 60 g soya Ad lib 20−34 6M Ensure Study Period 16 M Total Number of Subjects in Study (sex and age) Kurpad et al. 2001 7:07 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 237 Energy (kcal/day) Fat (g/day) .1 0.7) 3. (1985)218 Ad lib diet Red kidney beans a 20 M 34−66 Anderson et al. H Liquid diets with added soya fiber.1) 30 25 29 19 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 3.3 1. (1988)217 Ad lib diet Beans (haricot) Slavin et al.4.4pt3_fm Page 237 Sunday. M pH See also Table 4.7 1.4.6) II 55 51 57 48 V 72 N (g/day) 3.9) 33 (6. (1984)209 Controlled diet Beans (Oats) Fiber Intake (g/day) and Fiber Method V.3 1. (1985)214 Controlled diets Soya Days on Diet Table 4.9 Transit Time (h) and Transit Method 42 (3) 41 (3) 27 (5. K A.8) 29 (3.3 (0.12 (Oats) E.4 III 19 Control XIII 3 3 3 3 3 +18 5 5 5 5 5 Beans 7 10 + 25 IV 10 + 25 10 3 Significantly different from control. May 6.4) Neutral Steroids (mg/day) 878 (206) 894 (184) Acid Steroids (mg/day) 154 (37) 108a (20) Comments Notes Other Data Available B.Literature Source Diet Fiber Source 10 + 50 47 Beans 23 23 Ad lib Beans + 8. 12 M 21−35 Fleming et al.5 (0. 6 Transit Time (h) and Transit Method 51 53 51 40 51 46 Fat (g/day) 3 (1) 4a (1) Neutral Steroids (mg/day) 787 (103) 759 (96) 556 (44) 732 (100) Acid Steroids (mg/day) 346 (64) 468a (72) 289 (93) 320 (89) Comments Notes Other Data Available Data read from graphs Patients with lipid disorders Also guar study with diverticular disease patients E In vitro fermentation B Fiber by Englyst82 In vitro fermentation Data read from graphs Fiber data by Englyst82 Fiber from psyllium measured by Englyst method82 E.14 Effect of Gums and Mucilages and Other Purified Sources on Fecal Composition 2387_ch4. (1989)222 Controlled diet Plantago ovata 9 14 3 + 30 Ispaghula Lactulose 25−38 7 7 7 7 7 7 5 5 3 14 7 7 7 7 7 7 21 21 Days on Diet Ad lib + 25 X + 12.8 26−38 Number of Subjects Collecting Feces 9 7 7 7 7 7 7 7 7 Fecal Weight (g/day) 101 159 (26) 251a (39) 178 (20) 276a (15) 200 (26) 159 186 160 173 160 214a 104 (24) 223 (39) Moisture (%) 84 78 84 75 Fecal Solids (g/day) 35 (4) 39 (4) 26 (3) 36 (5) 3.3 Rasmussen et al.5 + 14.7 0. (1988)223 44 F 6M 3 11 Plantago + 25 3 Basal 3 Miettenen et al.0 Control Guar Control Xanthan X 7 Placebo Hamilton et al.8 X + 18 Control Ispaghula Psyllium Basal Fiber Intake (g/day) and Fiber Method V Fecal Collection Period (days) 6M 3F 30.4.7 3. K Retinyl esters Glucagon 238 Tomlin and Read (1988)220 Semi-controlled Guar Ispaghula Xanthan gum Total Number of Subjects in Study (sex and age) Abraham and Mehta (1988)219 Controlled diet Psyllium Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed Table 4.2 6.0 4.9 1. 3RD EDITION Energy (kcal/day) N (g/day) . May 6.Study Period + 13. 2001 7:07 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.9 1.4pt3_fm Page 238 Sunday. (1987)221 Ad lib Ispaghula Lactulose Literature Source Diet Fiber Source 6M 1F 7M 31. 8 4. (1987)195 Ad lib Pectin + 16 Pectin 21 21 IV 16−21 Ad Lib V.4. (1984)228 Ad lib Tragacanth 7 23 Ad lib 10.8 (2. M.4.4pt3_fm Page 239 Sunday.0 115 II I-V 70 (6.4. K. (1986)225 Controlled diet Guar Ad lib Methylcellulose 2387_ch4. M. (1988)205 Semi-controlled Psyllium Mallet et al. E. N Blood biochemistry B. THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION a 5M 35 26−50 Eastwood et al. N Blood biochemistry E.5 6. (1987)227 Ad lib Xanthan gum 24−58 Ad lib + 10 Guar 5M 14 7 +4 22 7 +2 Basal Anderson et al.1 (1.9 23 Xanthan Significantly different from control. 2001 7:07 PM 239 .CMC 38 12 M 20−30 12 F 29 22−38 3M 3F 22−26 Tomlin and Read (1988)229 Ad lib Ispaghula (I) Polydextrose (P) Stevens et al.8 12.4 5.12 Also bran (see Table 4.8) 7.4 to 12.2) 3.8 (0.8) 386 (66) 521 (111) 560 (80) 600 (40) 672 (120) 728 (160) 368 (36) 752 (112) 347 (39) 656 (116) 364 (77) 664 (116) Also bran (see Table 4.3) 59a (3) 33 35 36 II 54 59 59 59 36 46 56 (12) 45 (10) II I 53 (10) 46 (10) 46 (3. not means B. XVI 21 14 7 +6 + 12 5 5 7 7 10 10 10 10 10 10 +6 + 30 + 32 40 Psyllium Basal 10 10 10 10 5 5 5 5 5 5 10 10 10 10 21 Gum Basal I 7g P 30g I 2g and P 30 g Basal I 7g I 2g and P 10 g 7 Control 9. K. M.2 154a 68 (6) 76 (14) 7. N Blood biochemistry Immunoglobulins B.5) 54 (7. E. May 6.4) 7.2) 8. K.9 +15 7 14 6 6 12 12 5 5 5 5 5 5 6 6 21 20 103 (18) 93a (36) 79 (8) 163a (11) 220 254a 236a 171 180a 174a 183a 125 (18) 188a (26) 187 (27) 242 (39) 140 (33) 242 (20) 73 38 (7) 60 (5) 80 75 75 74 79 76 75 20 (1) 33a (1) 32 (5) 46a (6) 44 (3) 51 (6) 6.12) Mainly study of fecal flora See notes to Table 4. (1986)226 Ad lib diet Carboxymethylcellulose M-C 2g M-C 4g 6M 21−28 18−70 27 Perragini et al.9 (1) 7. A Also constipated subjects studied with psyllium and methyl cellulose See also Reference 224.4 4. M.8 6.4) 5.8 0.7 1.3 (0. 5M 36 21−57 Eastwood et al.0 (1. E.12) Stool consistency Medians. 15 Effect of Foods and Mixed Diets on Fecal Composition 2387_ch4. (1990)235 28 20 24 XVII 57.Forsum et al.246 Ad lib Vegetarian Fiber Intake (g/day) and Fiber Method XVIII XIX Fecal Collection Period (days) 5 3 3 3 3 2 2 Number of Subjects Collecting Feces 5 288 (62) 171 (5) 209a (7) 144 (18) 197a (10) 80 78 77 78 71 78 176a Fecal Weight (g/day) 72 Moisture (%) 118 Fecal Solids (g/day) 58 (8. (1989)230.9) 40 (1) 47a (1) 33 (1) 43a (2) 38a 33 Apparent Increase in Fecal Weight per Gram Dietary Fiber Fed 4. May 6. (1990)233 Semi-controlled Mixed foods 91 91 30 Literature Source Diet Fiber Source Vegetarian Study Period 20 6M 20 F 44 27−61 Days on Diet Ad lib Total Number of Subjects in Study (sex and age) Allinger et al. K. Na Comments Notes Other Data Available Energy (kcal/day) N (g/day) Fat (g/day) Transit Time (h) and Transit Method 240 Table 4.4pt3_fm Page 240 Sunday.3 3. 2001 7:07 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.6 5.6 Prunes A 28 XVI 18 Grape 41 M 21 56 26 High fiber Tinker et al.7 6. (1991)234 Ad lib (grape juice) Prunes 56 IV 12 Basal 34 M 50 47−55 Kesaniemi et al.8 Neutral Steroids (mg/day) 2268 (115) 2088 (102) Acid Steroids (mg/day) 501 (54) 518 (67) 1351 (83) 1391a (91) A Fecal biomass E E Biliary lipids Cholesterol kinetics Fecal pH L Urinary N. 3RD EDITION .4. 0 C Low 20 55. L Monastery A = Cereals B1. (1988)236 Semi-controlled Mixed sources Controlled diet Mixed sources 2 2 7 7 3 3 5 5 5 26 25 6 5 5 86 (26) 145a (18) 136 (15) 288a (25) 179 (19) 108 (21) 74 (9) 86 85 72 77 80 20 (1.7) 25 (1.6 (0. May 6. B2 = digestible fibers C = low fiber 2387_ch4.3 (0.4 B1 Significantly different from control.1) 115 (18) 393 (44) 249 (18) 134 (9) Also follow up at one year Fecal steroids pH E A Digestible energy Metabolizable energy Week 5 values used E.0 B2 26 42 42 20 20 71.2 12. (1988)237 Controlled Fruit and vegetables a 11 F 14 M 45.0 0. 2001 7:07 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 241 .11 F 46−67 Reddy et al.2 High High I 13.4) 3.7) 39 (3.5) 21 (2. 12 M 41 Miles et al.6 (0.8 4.2) 1.3) 2.8 1.8 47.4pt3_fm Page 241 Sunday.8 Ghoos et al.4) 3. (1988)238 Low fat/high fiber Low fat/high fiber 37 XIII 17 26 Low Ad lib 37 23.5) 36 (1.5 (0. 31. Br. Br. 819. D. Gut. 475. Varicose veins. M. Bull. 50. P. A. An investigation of the experimental conditions which could affect water-holding capacity of dietary fibre. 17. D. 1978. Nutr. Stephen. D. 1972. U. R. Dig. 3. J. 1980. M. and Durnin. 27.K.. J. The effect of cellulose. Acad. Rec. The laxative effect of a regenerated cellulose in the diet. H. J.. 315.. 21. Cummings. Br. A. Painter. Williams. Med. P. New York. D... Nutr. J. Burkitt. 32. A. Cummings. Br. Med. 1980. in Colon and Nutrition. 1981.. W. Sci. 24. W. H. D. L. H. T. Lancet. H. Br. D. London. and Cummings. A. 1980. K. 7. and Eastwood. 1983. 5. 1954. 1972. 58. Br. H. G. W. Houston.. Cancer. Diverticular disease of the colon: a disease of this century. T. Eastwood. Med. 19. 556. JAMA. Robertson. i. H. A. L. Green & Longmans.... J. D. apple. Longman.. Br. Med. Calorie conversion factors. J. J. Nutr. bran and guar gum. and Painter. and Kay. and Cummings. 20. J.D. 1979. Tainter. Robertson. J. Karaya gum as a mechanical laxative: an experimental study on animals and man.. M. N. W.. M. H. ii. T. U.. S.. J. S. 16. and Olmsted. The microbial contribution to human faecal mass.. Southgate. 284. 1982. Cambridge. Eds. Ann. J... 2. 2. 20. A. A. 1971.. 25. J.. S. M.. A method to measure the water-holding properties of dietary fibre using suction pressure.. 1981.K. Microbiol. 65.. The aetiology of appendicitis. Oxford Medical Publ. V. E. J.. 1408. J. University of Cambridge. 1936.. S. N. Burkitt. A. Colloid laxatives available for clinical use. and Tadesse. Sci. 433. 6. Unprocessed bran in the treatment of diverticular disease of the colon. Med. M. M. 995. N. Cambridge. Dietary Fibre and Human Colonic Function. 1970. and Isaacs. Dis. C. Almeida. Eds. University of Cambridge.. 12. 283. 1840. Hertz. 11.. Plenum Press. A. D. 102. thesis. 128. 130. J. L. W. A. Treatment of constipation with Normacol. 717. Epidemiology of cancer of the colon and rectum.. Stephen. A. H.. Porges. H. Am. 539. Ivy.. J... L. J. R. Nutr. cabbage. 24. D. M. 517. 26... Am. Painter.. 1938. Kasper. N. 595. J. 1936. K. Morgan. P. Dis. 37. H. 83. 1981. 10. Effect of dietary fibre on stools and transit times. 1.D. R. J. 1229. Diverticular disease of the colon. A. 29.. 3RD EDITION REFERENCES 1. J. Short chain fatty acids in the human colon. Burkitt. M. Nature (London). 1928. D... 1980.. 3. thesis.. Spiller... and its role in the causation of disease. 18. in Medical Aspects of Dietary Fiber. 438. M. 58. 9. A.. Stephen. 137. B. Robertson. A. 5. M.. Polysaccharide fermentation in the human colon.. Intern. Br. Stephen. and Olmsted. P. and MacDonald. A. M. W. ii.4pt3_fm Page 242 Sunday. P. Orme. G. The Treatment of Habitual Constipation by the Bran Method. 763.. 1968. 22... 45. 4. J. 1981. Mechanisms of action of dietary fibre in the human colon. 10. MTP Press.. A.. May 6. 23. 1972. Robertson. 1941. and Colebourne.. 13. 2. Brown. and Eastwood. Southgate. Walker. Lancaster. J. P. Constipation and Allied Intestinal Disorders. N. M. M.. Br. M. 1934. 45.. 32. and Tainter. A. Dig.. Water holding by dietary fibre in vitro and its relationship to faecal output in man. 15. D. G. 11. G. 28. . H. An experimental measurement of the factors used in the calculation of the energy value of human diets. Burkitt. Brydon. Branch.. and Eastwood. F. Lancet.2387_ch4. 2001 7:07 PM 242 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 22. The manner in which food controls the bulk of the feces. and James. and Goebell. Surg. 2. Related disease—related causes. 1971. J. Gut.. and Cummings.. 1936. Colonic response to dietary fibre from carrot. Ph.. A Treatise on the Causes and Consequence of Habitual Constipation. Med. A. Effect of fiber on colonic function. Quantitative comparisons of colloidal laxatives. A. Nutr. Med.Y. 1981.. 5. Cummings. deep vein thrombosis and haemorrhoids: epidemiology and suggested aetiology. 91. 1909. 30. Painter.. A. 247. 87. J. Ann. Measurement of water-holding properties of fibre and their faecal bulking ability in man. 1969. W.. ii. J. Williams. 28.. Lancet. 13. hemicellulose and lignin on the weight of the stool: a contribution to the study of laxation in man. O. Dimock.. Burne. 14.. H. Gray. 8. Eastwood. H. Dietary fibre. A. Cummings. 46. J. Food Agric. P... 722. J. Jenkins.. J. Burkitt. 8. 586.. An examination of factors which may affect the water holding capacity of dietary fibre. Lancet. 1969. R. A. H.. Brydon. and Buchanan. . J. A method for the rapid distillation of the lower volatile fatty acids from stools. Gastroenterology. A comparative study. J. 35. Med. L.. 275.Y. V. K. 63. 20. C. J. Machle. Kopstein. 101. V. A. C. H. N. 41. 1952. 1935. W. Med. Wozasek. G. Cowgill. Med. 21. 1941. Blake.. Stool VFA. and Steigmann. J. and Gelehrter. 279. Dis.. A. O. 39. J..2387_ch4. W. H. 60. F. F. 8. 805. Whitaker. 42. Food and bulk producing drugs in constipation. 58... 645. and Olmsted. 77. Exp.. Tainter. 56. W. Chem. 57.. Biol. 11. Streicher. and Palmer. Chem. Effect of bran upon composition of stools. Rev. Duden. 1940. W.. F. 1941.. 61. I. Roentgen study of intestinal motility as influenced by bran. 34. O. 114.. 404. H... 1929−1930.. 1942. P. D... and Bauerlein. 179. J. S. II. B. 1936. Am.. 1943. F. Dis. Dig. and Frank. Am. O. M.. 16. J.. and Quirk. 1952. W. 8. R.. Determination of the crude fibre in national flour. 423.. 795. Bastedo. 143. 1935. Sci. F. Am. 45.. 48. J... Further studies on the use of wheat bran as a laxative. B. 59. W. G. A clinical evaluation of certain bulk and irritant laxatives. Schweis. J. The influence of feeding bran. R. 44. Med. and Ivy. Soc. Gastroenterology.. Mason.. R. and Tainter.. 1932. 1943. and Clark. Olmsted. Clin. Am. 55. 276. 551. Studies on colon irritation. C. 1973. H. The use of methylcellulose as a bulk laxative. Gastroenterol. N. Cass. M. Acta Agric. 20. Methyl cellulose as a colloid laxative. 1949. Effects of hydrogels on the configuration and function of the colon. and Anderson. G. Am.. Cause of laxative effect of feeding bran pentosan and cellulose to man. Am. 1934. 99. Olmsted. Am. Mason. M. 53. Agric. G. Fantus. Some observations on the distribution and origin of nitrogen in sheep faeces. and Parker. Chem. R. Dis. 1940. W. Cowgill. Am. H. 1984. H.... C.. Assoc. Schultz.. and Quirk. Carboxymethy1cellulose as a colloid laxative. L. Dig. 73. Influence of size and shape of bran particles and of crude fiber isolated from bran. B. A method of improving function of the bowel: the use of methylcellulose. and Timm.. B. G. J. J. Rev. Analyst. 13. Soc. 98. Fantus. 336. III. 10.. A. JAMA. D.. 1822.. Curtis. The mode of action of bran. Biol. Am. Constipation: the laxative value of bulky foods. Effect of bran.. O. J. and Schmidt. Williams. May 6.Y. On the fate of ingested pectin. J. Clin. Gastroenterology. D. Lignin.. 259. 1949. 1941. L. Stein. D. 1948. Werch. 8.. 85. Biol. W. Biol. Analytical Methods Committee. 20. Studies on colonic irritation.. 1944. 62. 8... Gut. T. Dietet. 47. Fantus. Fantus. 2001 7:07 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 243 33. Gastroenterol. Olmsted. 9. Bargen.. . Biol. Dig.. Heyroth. Examination of feces. Dig. Clinical evaluation of a new laxative. Dis. Digestibility and effect on laxatives of crude fibre and cellulose in certain common foods. 26.. 15. Chem.4pt3_fm Page 243 Sunday. N. 32. A biochemical method for determining indigestible residue (crude fiber) in faeces. 653. 298. L. Streicher. 1935. F. J.. Bulk of feces. W. R. 108. 1941. W. 37. Dis. IV. L. B. and Sullivan. N. JAMA. Wozasek. L.. 48.. 50. 1943. pentosan. 46. J. 141. J. J. A. Camb. Lab. 51. 108. Proc. The fate of methyl-cellulose in the human digestive tract. Fantus.. Constipation: further clinical evidence of the use of bran as a dietary laxative agent. J. J. 68.. 54.. 115. Biol. 1969. Comparison of a plain methyl cellulose with a compound bulk laxative tablet. Studies on colonic irritation. W. JAMA.. 40. Scand. J. Curtis. Exp. 43. 153. State J. 1774.Y.. 1941. K. and Frankl. Cummings. and Timm. 100.. D. R. 277. Proc. J. Dig. A. M. 36. Dis.. Dig. and fiber to man. C. 296... H. Dig... 39. and Witherup.. Wozasek. Pauly. 7... cellulose and non-water soluble hemicellulose.. Am. K. H. 54. A preliminary report. Constipation: clinical and roentgenologic evaluation of the use of bran. S. Olmsted. Gastroenterol. A. H. R. 1866.. Hoppert. 149. J. Berberian. A. 157. C. and Steigmann. E. Cellulose and the human gut. 23. Rev. The influence of bacterial activity in the alimentary canal of rats on faecal nitrogen excretion.. The mode of action of bran. 49. Laxative effects of wheat bran and washed bran in healthy men. Williams. M. 52. 38. 141. Dig. W. 1945. A.. Dis.. 319.. 449. J. 1944. 2. 1933. Dis. and Wolf. R. and Steigmann. Med.. 1948.. O. 25. in Analysis of Dietary Fiber in Foods. 842. Soc. The Nutritive Value of Fruits. A. 15. IV. E. F.. in Analysis of Dietary Fiber in Human Foods. T. Ribiero. and Shackleton. Goering. O. 72. P. Assoc. A. New York.. 11th ed. AOAC. 2. 76. Agric. Van Soest. Use of detergents in the analysis of fibrous feeds. A. P. 1979. II. K. Agriculture Research Service.. S281. F. T. Wolever. Analysis of dietary fiber. i.. in Fiber in Human Nutrition.. T. and Wursch...2387_ch4. H. S. M. Analysis of dietary fibre. B. Agric. P. 85. Assoc. 1976. H. Official Methods of Analysis of the Association of Official Analytical Chemists. A. D. H. A guide to calculating intakes of dietary fibre. McCance. and Farrell. 1972. His Majesty’s Stationery office. J. Bailey. J. 10.. D.. 70. H. Cummings. D. 1976.. H.4pt3_fm Page 244 Sunday. D. H. Off. 1936. New York. Assoc. and Walker. P. R. Sci. 1978. Eds. Hum.. Englyst. J.. Determination of the non-starch polysaccharides in plant foods by gas-liquid chromatography of constituent sugars as alditol acetates. R.. Nutr. Agric.. J. 75. Transit through the gut measured by analysis of a single stool.. and Southgate. 1969. E. Department of Agriculture... A.. D. Simplified method for measurement of total non-starch polysaccharides by gas-liquid chromatography of constituent sugars as alditol acetates.. Development of comprehensive system of feed analyses and its application to forages. H.. 119. J. R. J. 80. 1984. Marcel Dekker. T. Washington. D. Aust. B. 1976. 69. A new method of studying gut transit times using radio-opaque markers.. M.. J. D. P.. 87. Analyst. S. Chem.... O’Shea. McCance and Widdowson’s The Composition of Foods. H.. 30.. and Maynard. T. Widdowson. P. 3RD EDITION 64. Am. and von de Bovenkamp. A. Hinton. 1982. 88. D. 1969. M. and Wursch. Sci.. P... Use of detergents in the analysis of fibrous feeds. Schweitzer.. A. L. and Van Soest. II. A. Nutr. 46. Sutherland. and Wine. Agric.. Eds. A simplified method for determining fibre in foods. 1977. James. A.. 829. J. Eds.. and Theander. Applied Science. J. J. 129. New York. Anim.. Washington. M. Res. 173. 107. 31. 1981. P. J. 65. J.. Trowell. Van Soest. U. 17. and Jenkins. R. A. A. P. 1976. 1966. 7. J. 203. T. O. 331. 307. 50. 81. T.C. N. . T. O. dietary fibre and atherosclerosis. and Theander... Paul. Van Soest.C. 49. Nutr.. Clin. New York. Ed. and Moran. and Wiggins. J. 613. A. Marcel Dekker. Atherosclerosis.. Southgate. M. Crampton. B. 303. H. F. 20.. Lennard-Jones. Southgate. D. 68. D. 79. 1968. 50... 967. Off. 1970. Gassull. M. Crude fibre. M. James. Food Agric. Agriculture Handbook No. 73. Leeds. 1970. E. and Cummings. Southgate. Wiggins.. 383.. P. Determination of Food Carbohydrates.. in The Analysis of Dietary Fiber in Food. G. Lancet. 219. 379. 2001 7:07 PM 244 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 1938. T. Nutr. P. T. Spiller. Her Majesty’s Stationery Office. Use of the Southgate method for unavailable carbohydrates in the measurement of dietary fiber. A. Gut... A. 1981. 26. Gut. 71. Forage Fiber Analysis: Apparatus. Angus. 82. S. The analysis of dietary fibre.. Southgate.. 86. London. Food Agric. 1981. J.. The relation of cellulose and lignin content to the nutritive value of animal feeds. J. James. W. Van Soest. Determination of plant cell wall constituents. Collinson. and Cummings. J. A.. 30. AOAC. 89... T. 84. Dietary fibre redefined. London. Determination of total dietary fiber by difference and of pectin by calorimetry or copper titration.. Chem. Off.. 109. 74. 1. C. and Walker.. T. F. T. Analyst. J. Collinson. D. 67. Unavailable carbohydrates. 1975. Vegetables and Nuts. 90. L.. and Young. W. Sci. J. D. Trowell. J. R. J. Southgate. J. Relationships between digestibility (in vitro). B. 1978. Reagents. do V. Englyst. H. Southgate.. A. and Theander. 1963. J. A. Ir.. 1967.. London. Non-nutritive residues. Katan. 73.. May 6. E.. Marcel Dekker.. A rapid method for the determination of fiber and lignin. Horwitz. A. 77... A. Plenum Press. H. 16. T. Dietary fiber analysis tables. 66. R. 78. Procedures and some Applications. Determination of carbohydrates in foods. W. Eds..S. Southgate.. W. A system for the replacement of crude fibre. 546. 138. 1. A. M. crude fibre and cellulose content of some animal feeds. Schweitzer. W. 83. Chem. Bailey. E. J. 1967. T. A. 937. and Amen. Proc.. A. Wyman. 1974. 28. Modification of eating habits. N. Br. Changes in fecal composition and colonic function due to cereal fiber. 29. J. J. Nutr. J.. Lancet. B.. 108. Hansky. 2086. H.. Jenkins. Hill. Intermittent and continuous faecal markers in short-term metabolic balance studies in young women. 1974.. Mitchell. 1978. 26. Br.. Biol. Effects of dietary fibre on the apparent digestibility of major food components and on blood lipids in men. A. Hill.. Vancouver. Exp. Drasar. and Wiggins. A.2387_ch4. Hill. Effects of unprocessed bran on colon function in normal subjects and in diverticular disease. The effect of wheat fibre on plasma lipids and faecal steroid excretion in man. Walters. J. E. A. M. Usefulness of chromic oxide as an internal standard for balance studies in formula-fed patients and for assessment of colonic function. 1977. 1443. 17. 117. II. D. 91. J. M. D. A. M.. Nutr. J. 304. J... Davies. Mineral metabolism on dephytenized bread. W. 833. D.. May 6. D. McCance. The effect of dietary fiber supplementation in man. Clin. D. Nutr.. 32. 109.. Kay. A.. B. GI Motility. A. 392... J. J. Baird. Walters.. J. H. Davies. S. J. A. P. 1979. 3. P. Mitchell. zinc and phosphorus by humans due to increased fiber and phosphorus consumption as wheat bread... 1976. M. Floch. J. 107. Effect of wheat fiber on blood lipids. J. Am. M.. and Baird. M. 112. H. Mathur.. J. Jenkins. Hill. M. R.. Jivraj. J.. S. A. T. The effect of meat protein and dietary fiber on colonic function and metabolism.. 98. 489. J.. D. and Southgate. B. Nutr. 4th Int. A..4pt3_fm Page 245 Sunday.. S. J. 2001 7:07 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 245 90. 1977. Am. and Hamilton.-M. M. 30. Proct. 30.. and fecal bile acids in normal subjects. J. J. Reinhold. R. W.. M... F. H. 99. and Floch. S. Floch. 469. C. L. H. 1129. M. 103. V. W. Clin. J. S. Nurr. Am. L. Dorfman. Metabolism.. Am. I. 101. Hill.. J.. W. 1975. Davignon. 31. 25. and Wicks..... Changes in bowel habit. Am. M. Nutr. Nutr. Clin. S. 1977. M... M. Nutr. P.. J. F. 105. 1978. H. J.... 106. S. W. 1976. 102. and Smith.. A. Drasar. The effect of intestinal transit and the feces of raw and cooked bran in different doses.. Measurement of the mean transit time of dietary residue through the human gut.. B. 1973. Med. I. magnesium.. J. and Connell. M. Med. Br. J. and Widdowson. and Floch.. E. 1976. J. Fuchs. 210. Effect of dietary fiber supplementation in man. P. Green. A. 100. R. Drasar. Southgate. A. 1976.. bile acid excretion and calcium absorption. 1976. M. and Ismail-Beigi. J. Clin. J. 37. G. Kirkpatrick. Effects of two types of dietary fibre on faecal steroid and lipid excretion. H. B. J.. Effect of bran on intestinal transit time in normal Indians and in intestinal amoebiasis.. 1942. J. Physiol. H. 1968. J. 365. 493. Am. Mulinos. J. M.. Clin. M. D. P. 187. M. 97. and Cummings. R. H. Symp. 1962. M. Decreased absorption of calcium. 1935. 47. Baird.. Colon Rect. Effects of dietary supplements of wheat bran and cellulose on faeces and bowel function. Br... 1976.. 113. A.. Abadi.. M.. Am. Branch.. 29. 1970. I. M... and Chadda. H. Farrell. 1978.. I. R. B. L. S... The effect of dietary fiber supplementation in man. Am. 110.. . 29. Branch. T.. Surg. 1976. 127. M. 29.–M. 2. Hill. Walters. Anderson. Girle. A. Sci. S. D. and Morgan. L. T. Kahaner.. and Fuchs. J. K. Clin.. Gut.. 29. R. Fuchs. B. Nutr. S.. The effects of two dietary fiber supplements on gastrointestinal transit. Canada. H. Rev. 94. S. J. Cummings.. Smith. Houston. Gastroent.. and Wiggins. Bone. 111. Simmonds. iv. L. Metabolic responses to dietary supplements of bran. 1474. 292. J.-M. A. H. R. J. 95. J. N.. 92.. Jenkins.. S. Med. Modifications of stool content by increased bran intake. and Truswell. J.. J.. S185. Davies. 104. The value of elective drugs in the treatment of constipation.. Manning.. fecal steroid excretion and serum iron. 101. A. Invest. 1975. 146.. Cummings. Cummings. T. The effect of dietary fibre on transit time. Clin. 1468. and Ahrens. and Jenkins. 24. J.. Faradji. 106. Findlay.. S. Sharpe. B. and Robinson. H. 93. 96.. Aust. D. J. J. Am. Gastroenterol. Alteration in fecal physiology and bacterial flora. 56. Mitchell Press. Connell. Gut. Nutr. H. M. Ram. H. H. 1408. C... and bacterial flora. and Eastwood . G. in Proc.. i..... and Arthur. W. 1437. H... Measurement of gastrointestinal transit using radioactive chromium.. S. 536. A. Eastwood. J. Pearson. 2. Clin. Clin. Heaton.. D. T.. I. and Fuchs. J. Nutr. Metabolism. stool weight and frequency.. Southgate. 227. J. . 1071. 117. L.. A. D. J.. W. M. Hectors.. Logan. C. R. Am. New York. Nutr. A. 122. Graham.. M. G. G.. M. Sandstead. C. W. Effect of dietary fiber on glucose tolerance of normal men. and Cummings. Stasse-Wolthuis. 2001 7:07 PM 246 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and Wiggins. F. 128. P.... J. J.. H. C.. Inglett. J.4pt3_fm Page 246 Sunday. H.. C. 61. Wayman. M. fecal lipids... B. Sandstead. A.. 10... J. Nutr. 34. Yu. H. F. Am. 216. Increase of serum cholesterol in man fed a bran diet. N.. Durrington.. Effects of dietary fiber from wheat. Jacob. G... and Kelvay. Clin. G. Reek. Wesstra. corn and soy hull bran on excretion of fecal bile acids in humans. 147... and Eastwood. F.. M. A. Bacteria. and Sandstead. Logan... Jacob.. Effects of a standard wheat bran preparation on biliary lipid composition and bile acid metabolism in young healthy males.. B. Wong. G. 1983. and Schippers. 56. J. M. S.. 9. I. J. 31. J. Navert. H. Am. Englyst. E. E.. Food Sci. G. J. Sandstead. L. Munoz. Stasse-Wolthuis. 1978. G.. S. Clin. Johnson. 1983. M. Physiological effects of fibre-rich types of bread. Correlation of gastrointestinal transit time to fecal weight in adult humans and the levels of fiber intake. M. 32. H. Sandstead. J. W. F. W. M... J. Dintzis. G. C.. and stool weight. J. von Schaik. 1976. Diabetes. A. Br.. and Hautvast.. Hautvast.. Dintzis. 394. Physiological effects of fibre-rich types of bread. F.. J.. 1979. I. H. Dintzis. 127. Jacob. R. The effect of dietary fibre from bread on the mineral balance of young men.. Inglett. A. 134.. H. R.. S180. Clin. 33. S. E. S. R. Klevay.2387_ch4. Nutr. E. Spiller. G. Effect of pectin on serum lipids and lipoproteins.. The effect of bran on bowel function in constipation. J. H. G... Sandstead. Age Aging. W. B.. R. J. 1981. Nutr. 933.. G. Rowe. Eur. Katan. J. J. S. 599. Logan. M. Effect of citrus pectin on blood lipids and fecal steroid excretion in man. Br. A.. Influence of dietary fiber from vegetables and fruits. M. and Shuey. J. Hermus. Van Dokkum. 47. A... Klevay. and Truswell.. 1986. Inglett. A.. Mineral requirements: influence of fiber and protein. Jr. M. Eastwood. A. Huijbregts.. M. P. G. and Scala.. Invest. J.. 34. G.. Jacob. 31. H. M. 116. J. J. J. A.. A. Reck. Lancet. 1980. N. 28. S. and Hartog. Br. M. M. R. Effects of dietary fiber and protein level on mineral element metabolism. 1976. G. Influence of dietary fiber on trace element balance. H. M... Van Dokkum.. 451. J.. J. Nutr. The effect of age. Munoz.. 29.. 171. Pikaar. A.. 50. 3RD EDITION 114. 267. R.. L. M. J. M.. M. 696... H. 1745. and Falkehag.. Brydon.. Atherosclerosis. J. Stephen. Am. 31.. 1982. R. Influence of cooked wheat bran on bowel function and fecal excretion of nutrients.. H. 32. S. Munoz. 130. May 6. and van der Werf. Cummings... Nutr. Am. in Dietary Fibers: Chemistry and Nutrition. G. Gastroenterol... Qual.. M. Moser. Stasse-Wolthuis. 451. 1979. L.. Nutr. P.. Br. Andersson.. Nutr. J. Effects of dietary fiber on plasma lipids of normal men. Rep. S.. S.. 126. Am. Clin. P. Munoz. 1981. Munoz. 1979. . J. Whittam.. C. J. J... A.. W. Albers.. J. 496. D. J.. J. H. A. Clin. Wil de Jong. N. Jacob. M. and Shuey. Nutr. Reck. and Thissen. E. 1980. M. W. E. R. L. M. J. Clin. Clin. II. A. and Miller. Klevay. J. Drasar. diet and large bowel cancer. F.. Int..... Smith. van Jeveren. 1982. Academic Press. H.. Eds. T. and Estes.. M. H. N... Kay. L.. bran or citrus pectin on serum lipids. H.. and Jenkins. S. 30.. H. A. 1979. M. and Brydon.. 1982. 349. Klevay. L. D. The effects of breads containing similar amounts of phytate but different amounts of wheat bran on calcium. A. H. 120. 132. Katan. E. 1410. zinc and iron balance in man. Nutr. and colonic function. K. T. H.. J.. van Berge-Henegouwen. W. Effect of a natural high fibre diet on blood lipids and intestinal transit in man. . 1979. Plant. J. whole-gut transit-time. R.. J. C. 133.. W.. Drummond. Inglett. 25. M. and Jacob. 118.. 1979. Nutr. G. Clin. Am. 503. 1980. A.. Dintzis.. N. Logan. H.. N. 87. G. B. A.. Smith. Clin. 125. H. 29. M. 50. Manning. M. Am. Klevay. F. and Inglett.. H. A study of bulking agents in elderly patients.. 129. M. 720. 135. L. Hermus. 115. 119. Nutr. A. A. 580. Nutr. M.. Munoz. 124. J. 1978.. sex and level of intake on the colonic response in man to wholemeal bread with added wheat bran. Effects of some cereal brans and textured vegetable protein on plasma lipids.... L. Am. H. Dietary fibre from bread: digestibility by the intestinal microflora and water-holding capacity in the colon of human subjects. E. J.. R. 123. Englyst. 23. J. Bolton. B. 2. Bingham. 131. 1977. J. 77. and Shuey. R. Sandstead. R. Emken. 121. Bell.. and a low-residue diet on fecal volatile fatty acids. L. A. 141. J. J. 145. C. Unconventional Sources of Dietary Fiber. and Kuhnlein.. cellulose and lignin on stool pH. I. 7: Gastrointestinal Cancer: Endogenous Factors.. 154.. S.. J. K.. L. 1982. Hillman. N. R.. 147. Am. Horrath. Nutr. P. 64. Van Soest. L... J. pectin.. Dietary fiber: the effect of particle size of wheat bran on colonic function in young adult men. Critique and evaluation of markers for use in humans and farm and laboratory animals.. M. A. 215. Chim. Comparison of the effects of high. 17. 5. A.. and Van Soest.. Branch.. 2535. and Abadi. 1932. K. 477... 150. J. Excretion of breath and flatus gases by humans consuming high fiber diets.. Fleming.. Ismail-Beigi. Nutr. L. M. Nutr.. Roe. 16. 1983. Br. and Tarpila. P. 1983. B. H. P. 1949. Nutr. 146. Nutr. Slavin. 135. Effect of purified cellulose. A. F. Management of constipation with a refined psyllium combined with dextrose. transit time and weight. 152. 27. Some physical characteristics of dietary fibre and their influence on the microbial ecology of the human colon. 33. Colonic function and fermentation in men consuming high fiber diets. and Southgate. Nutr. D. P. Influence of refined cellulose on human bowel function and calcium and magnesium balances. J.. and Leklem. D.. 1982.. J. Heller. Influence of dietary fiber on fecal excretion of volatile fatty acids by human adults. transit time. J. Dis. A. Block. 25. R. J. Nutr. 1983. R. 1464. 151. J.. C... 510. 2001 7:07 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 247 136. 451. Cellulose esters in the treatment of constipation. Correa. Van Soest. 1734. Houston. J... J. 33. R.. Canada. I. and Marlett. Lewis. J. Reinhold.. Nutr.. Nutr. R.. Bruce.. Am. Symposium Series 214. J. Van Soest. 112. B. J. Van Soest. E.. Robertson. Int. W. Nutr. Am. 33. Nutr.. A. A.... 1978. D.. Fisher. Uden. Br.. P. E. J... A. Nutr. 1981. G. S. P. D.. Soc. 158. A. J. Furda. B. Rivers. J. 110.... M. J. 138. Hill.. in Proc. A. Nutr.. May 6.. 754. H. 144. 471. Judd. 155.. Rivers. Some factors influencing the ecology of gut fermentation in man. The effects of a supplement of dietary fibre on faecal excretion by human subjects.. Proc. D.. J. Jenkins. P. Robertson. Lewis. A. A. 79.. Br. McBurney. H. J.. 1984. and Hackler. B. 1980. B... T. in Banbury Report No. 142.. and fecal weight in humans. The digestion of pectin in the human gut and its effect on calcium absorption and large bowel function... Cummings. D. and Pomare. fecal bile acids and biliary lipids in normolipidemic and hyperlipidemic individuals. The role of dietary fiber in human nutrition. C. P. 113. Clin.. 158. Southgate. P. J. 41. 153. 50. 1982. Van Soest.. Effect of refined cellulose on apparent energy. Am. A. M. S.. N. Van Soest. 1983. J.. J. R.and low-methoxyl pectins on blood and faecal lipids in man. Prynne. T. E. 1983. J. R. and Rodriguez. 14. American Chemical Society. and Hill. Robertson. Nutr. 2020.. J...4pt3_fm Page 247 Sunday. Lewis. Effect of pectin on serum cholesterol. Rep. 113. 43. magnesium. Peters. Faraji. Acta. Spiller. 1613.. 137. Roe.. and Hackler. Influence of dietary fibers on fermentation in the human large intestine. Eds. J. Roe... E.. 143. B. The influence of dietary fiber source on human intestinal transit and stool output.. 1980. D. J. 1947. Jivraj. Marks. T. Dis. 189. J.. 61. fat and nitrogen digestibilities. Differing effects of pectin... Br.. G. B. and Truswell. The effect of dietary citrus pectin on the excretion of human fecal neutral and acid steroids and the activity of 7 α−dehydroxylase and β−glucuronidase. Effects of cellulose added to diets of low and high fiber content upon the metabolism of calcium. J.. Cold Spring Harbor. 41. 1977. 113. W. Clin. J. J. Ed. M. L. 139. Rivers. 1133. zinc and phosphorus by man.. Cornell Nutrition Conference for Feed Manufacturers. J.. H. Marthinsen. P.2387_ch4. 1979. A. A. Slavin. J.. 1980. Wrick. S. D. J. J. Am. Nutr. 1981. Nassar. J. A. T.. L. D. M. J. Hackler. A. E.. L. Dig. 157. Chernoff. Van Soest.. S. Lipkin.C. Ehle.. T. and Robertson. and Wilkins. Fleming. M... 34.. and Fleming. Clin. S.Y. Tannenbaum.. 1979. E. W. 148. 1977. 149. 1980. 112. M... J.. Am. P. F. 1983. J. 107. A. P. M. and Allen. Clin... J. 495. J. L. Cold Spring Harbor Laboratory.. Marthinsen. Miettinen. . Nutr. J.. D. P. J. A. Nutr. Clin. and Marlett. K. J. Wiggins. 48.. S. H. 140. S. and Shipley. S. 2068.. and Wrick. Dig. Washington. J. 156. M. S. Ross. E. J.. Gates. J. Nutr. L. number of defecations. L. Nutr. 143.. J. 179. and Steinke. Clin. A. A. Clin. Herbst. 1977.. S. P. B. D. J. G. K.. 1849. M. E. 32... D. A..... Tadesse. 11. S. Clin... J. 824. 44. A. Clin. I. Clin. Nutr. . C. and colonic function. J. Bekhechi.. Pometta. and Anderson.. Nutr. W.. A. J. Am. A. J. F. S. E. Behall. P.and low-fiber diets on human feces. S.. Miller. W. The effects of dietary gum karaya (sterculia) in man. G. 461. Nutr. J. 1983. J. A. H. Nutr. P. H. H. A. II. D. 1983. and Calloway.. J.. J. Agric.. and Kay. 1982. J. A. Sieling. R. J. 38. Beyer. R. G. nitrogen and fat. J. D. G. 33.. R. Zinc. Judd. J. 2061. copper and phosphorus balances. Bulk laxative efficacy of a psyllium seed hydrocolloid and of a mixture of cellulose and pectin.. Antonis. E. Am. steroid excretions... Clin. W. J. Am. E. Clin. Brydon.... Am. The influence of diet on serum lipids in South African white and Bantu prisoners.. M. L. K. Khumalo. 165.. G. Ir. H.4pt3_fm Page 248 Sunday. G. 1983. Bennick. Am. 1962.. 37. M.. M. and Bersohn.. Rees. E. 171. Lett. Mineral metabolism of healthy adults on white and brown bread dietaries. B. E. D. 175. May 6.. 169. W. K. Lin Chen. Kelsay.. 1118. S. Effect of fiber from fruits and vegetables on metabolic responses of human subjects. J. S. K.. J. 31. and Bersohn.. C. Nutrient utilization by human subjects consuming fruits and vegetables as sources of fiber. 10. 166.. I.. Clin. W.. 1942.. 3RD EDITION 159. and Brydon. 180. 32. Sci. 1979. 1981. Schweizer.. Clin.. Velema. 38. C. J. S. D. Am. M. Clin. 1981. and other parameters in humans. RietbergBrussard. F. fecal excretions and apparent digestibilities. E. Kirby. Nutr. M. L. Anderson. Reimann. J. 162. Calloway.-J. 1979. H. transit time and stool weight. 168. 1978.. Nutr. and Flynn.. and Cooper. M. and Truswell. 1983. serum lipids.. Kelsay. Am. M. and Prather. III.. Am. Nutr. Eastwood. Jacob. 485. Plant Foods. M. Am. 174.. J. McLean. Ndaba. T. L. and Prather.. Eastwood. O. 19. W. 161. The potato as a source of fiber in the diet. 32. Dietet. and Anderson. A. Am. G. 1982. and Bron. F. 1979. D.. F. Am... Am. Effect of fiber from fruit and vegetables on metabolic responses of human subjects: fiber intakes. The effect of raw carrot on serum lipids and colon function. and Kretsch. Am. 271. J. S. 1492.. 32. Effect of fiber from fruits and vegetables on metabolic responses of human subjects. M. and Lincoln.. Koellreutter.. M. A study of the effects of dietary gum arabic in humans. Oat-bran intake selectively lowers serum low-density lipoprotein cholesterol concentrations of hypercholesterolemic men.. 177. and Prather... Food Chem. E. 504. A. M. Spiller. J. R. S.. The influence of diet on fecal lipids in South African white and Bantu prisoners. Effect of fiber from fruits and vegetables on metabolic responses of human subjects. 176. Leeds. 1979.. 31... Kelsay. 1978. T. iron and silicon balances. Haricot beans. 29. Nutr. Kretsch.. Physiol. Toxicol. 285. 1. E. Bausch. Am. J. Robertson. Nutr. J. 313. Metabolic effects of dietary fiber from dehulled soybeans in humans. Zondervan. B. magnesium. 34.... J. 34. M. J. Clin. M... L. and Prather. Effects of high. W. M. Katan. J. K. G. and Widdowson. B. 164.. Brydon. 178. Tsai. nutrient balance. D. J. J. G. J. Mott. M. Protein and energy utilization in men given a rural Guatemalan diet and egg formulas with and without added oat bran.. 167.. Crawford. A. R. Med. 163. Nutr. J. A. Hermus. 173. The effect of rolled oats on blood lipids and fecal steroid excretion in man. and Eastwood. Nutr.. Owen. fecal lipids. W. Clin. R. Assoc. Wenham. P.. 4. 1978... Bowel transit time.. P. 159. J. Chernoff. W. M. A. Stasse-Wolthuis. L. M. 1149. Walls.. Goering. I. Behall. W. Calcium. J. Am.2387_ch4.. Flynn.... 268. Antonis. 2307. 172. Brydon. W. L. Pharmacol. J. Clin. 1981. 1962. R.. J.. Some aspects of bile acid and urobilinogen excretion and fecal elimination in men given a rural Guatemalan diet and egg formulas with and without added oat bran. R. Eastwood. A. B. urinary excretions of energy and nitrogen and apparent digestibilities of energy. 34. A. J.... 101.. Behall. Kelsay. Nutr. Anderson.. R.. E. M.. 1979. A. J. A. The effect of a natural high-fiber diet on serum lipids. and Prather. J. Clin. A.. Kelsay. Lo. Clin.. McCance.. D. Clin. and Burkitt. 1876. R. Beirn. Effects of soy polysaccharide on gastrointestinal functions. M. Am. Shipley. Nutr. 2001 7:07 PM 248 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.. fecal weight.. E. 146. H. 160. Ross. 72. glucose tolerance. H... 1979.. 170. 1881. Clark.. J... J.. A.. Hautvast.. 17. 32. 1889. 945.. Relation to other disorders and fibre intake. Brodribb. J. 1065. Gut. Eastwood. Hyg. L. C. 19. 1977. Rowland. Nunes. Clin. Warner. III. D.. and McDougall. Med. A. 185. 1981. 189. T. Nutr.. 116. and total diets: interlaboratory study. F.-G. R. 2001 7:07 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 249 181. 1974. Gut. 40. 644. Effects of bran on serum cholesterol. Comparative digestibility of wholemeal and white breads and the effect of the degree of fineness and grinding on the former. Smith. E. Prosky... Engle. 1987. S. and Feldheim. O. A. 67. A. 1986. Am. 1984. fat. 191.. 4th Int. Vancouver... Furumoto. Diet and diverticular disease. 196.. Meuser. Petro.. and Chen. and Pritchard. G. M. M.. J. Am. C. Anal... Johnson. 19. H. M. Camb. J.. Nutr. W. M. Z. Beitrage zur Analytik von als Lebensmittelzusatzstoffen verwendeten Polysacchariden. Story. A. 282... M. N. R. Fedail. Are fibre supplements really necessary in diverticular disease of the colon? A controlled clinical trial. 1980. 778. 677.. W. E. Littlewood. J. R. GI Motility. W. W. Clin. N. 162. J.. 343. Sieling.. and neutral sterols. J. Natl. 19.. Vitamins and other nutrients. 129 (Suppl. J. A.-J. Elton. 1942. Anderson.. E. K. 193. Brydon. The effect of low and high residue diet on the rate of transit and composition of the feces. 1429. 1988. 202. S. 197. and Farthing. Bacon... J. Lexington. F. D. 137. The interaction of dietary fibers and cholesterol upon the plasma lipids and lipoproteins. H. G.. D. Puska. Balasubramanian. Med. S. Ornstein. and Korpela.... E.. M. J. and Scala. T.. D. and Musa. F. Whittam. N. Comparison of bran. Schweizer. 1037. 988. transit time. S. 184.. Br. Maltz..... 195. M. J. 1986. W. R. M.. in Proc. Assoc. faecal mass... R. Gut. E. Chem. A. D.. H. L. 1144. Wisker.. bile acids and neutral sterols. Eastwood. 101.. and Smith.. N. W. 1. 1974.. Miettinen. Parks.. Tarpila. 190. Sharma. Clin. B.. 181. P. Furda.. 1986. I. J... B. Determining compliance with a dietary fiber supplement. B. A... and Draper. 1985.. 1976. J. 60. Lebens. and Groves. J.. McLean. J. M.. H. Irwin.. Br. Baird. S. I. 1353. Cancer Res. North..4pt3_fm Page 249 Sunday. R.. J. Am. I.. 118. F.. H. 1978. 1978. 34. fecal bile acids. Spiller.. Badi. S. Assoc. M.. Canada. A... Soc. Asp. R. Wong. Effect of bran particle size on stool weight. Connor. S. and Duthie. 1984. Taylor. A. A.. 776. 423. Lin. Mitchell Press. Parks.. Nutr. Mergenthaler. Schweizer... J. and biliary lipids in patients with diverticular disease. F. A. T. sterol balance. J. and bowel function in human subjects. Med. 76. Determination of total dietary fiber in foods and food products: collaborative study. L. T. Off. 424. Diverticular disease: three studies.. A. J.. L. S.. 43. Metabolic effects of bran in patients with diverticular disease.. Invest. Asp. Reddy. W. 67. A. N. Clin. II. 1976.. Forsch. and Cox.. M. Mallett. R.. Brodribb. C. The effect of coarse and fine Canadian Red Spring Wheat and French Soft Wheat bran on colonic motility in patients with diverticular disease. Br. 68. S. 60. K. Lebens. 198.2387_ch4. I. Smith. G. J. Dietary modification of intestinal bacterial enzyme activities—potential formation of toxic agents in the gut.... J. The effects of sorghum and wheat bran on the colonic functions of healthy Sudanese subjects. 201. Bran tablets and diverticular disease. Metabolizable energy of diets low or high in dietary fiber from cereals when eaten by humans. Anal. fats. Raymond. Treatment with bran. T.. M. Fowler. S. 251. G. Long term effect of whole meal bread on stool weight.. A. I. Prosky... J. and Scherz. J. food products. J. E. Vitamins and other nutrients. L.. 22). Gastroenterol. Suckow. P. E. 1987. L. G. 188. Chem. J. ispaghula and lactulose on colon function in diverticular disease. 194. 192... 182. Forsch. minerals and steroids and gastrointestinal transit time in healthy young women. 1044. Nutr.. Determination of total dietary fiber in foods. J. W.. A. Laakso.. G. Z. Drummond. and Connor. M. Proc. J. 203. 183.. A. 186. and Harland. R. Hutchinson.. 199. and Kulikowski.. L. Macrae. Simi. T. L. I. B. Alton. A. I. . 1. Plant Fiber and Food. 25. 1985. and Eastwood. Devries... Symp. Unter.-G. 2460.. J. Verfahren zur Bestimmung von unloslichen and loslichen Ballaststoffen in Lebensmittein. J. Furda. Off. M. J. Metabolic epidemiology of colon cancer: effect of dietary fiber on fecal mutagens and bile acids in healthy subjects. Devries.. Marlett. Effect of increasing levels of hard wheat fiber on fecal weight. J. 187. W. 1981. J. 42. 1978. and Metsaranta.. J.... B. 1976. 47.. J. University of Kentucky.. E. Cancer Inst.. S. Med. Nutr. May 6. and Humphreys. E. 200... F. T. Fry. and Harland. Unters. Scand. J. 211. The influence of ispaghula husk and lactulose on the in vivo and the in vitro production capacity of short-chain fatty acids in humans. 1986. J. ammonia and bacterial enzyme activities in healthy humans.. 253. 1985. Am. J. K. Fleming. J.4pt3_fm Page 250 Sunday. R. 2353. Indian J. W. W. 373. 214. B. M.. 1986.. 37. (Tokyo).. Schweizer... Gastroent. Okuda. (Tokyo). A... A. et al. H. J. J. Anderson.. 333. Vilaminol.. and Perman.. A. Krag.. Clin. 1. J. H.. 1988.. and Mortensen. Nutr. 1991. 1985. D. 1017.. Diabete Metab.. 113. Burdick. J.... and Mejean. 71... Miyoshi. J. The dietary effects of sodium carboxymethylcellulose in man. S. Methane production and bowel function parameters in healthy subjects on low and high fiber diets. Ment. A. 222. Petro. and Levitsky. Y.. Sci. L. Effects of brown rice on apparent digestibility and balance of nutrients in young men on low protein diets... W. Br. W... and carbohydrate absorption in men. 205. Nutr. C. 88. 199. and Mitsuoka. Assoc. Holmes. fecal steroid excretion. 7. Tomlin... 1987. Parent. Clinical evaluation of methylcellulose as a bulk laxative. H. 82. Nishida.. 67. W. 220. Okuda. (Tokyo).. I. 213. N. J. L. B. J..... Sci. E. J. K. M. L. Chem. 226. L. J. Hall. Marlett. and Story. The effects of dietary fibre in a liquid diet on bowel function of mentally retarded individuals. 209.. Hamilton.. 317. 1988. Penagini. soluble and total dietary fiber in foods and food products: interlaboratory study... and Slavin. L.. J.. Influence of frequent and long-term bean consumption on colonic function and fermentation. Determination and apparent digestibility of neutral detergent fiber monosaccharides in women... P. Chen. Chem..... nitrogen and fat. Vigorelli. T. Slavin. Castagnone. Am. Asp. V. 1146.. M. M. P. G. 581. E. M. J. Gastroenterol. Effect of corn fiber residue supplementation on fecal properties.. and Koishi. and degradation of neutral detergent fiber in young men.. T. Gastroenterol. fat and calcium digestibilities and fecal cholesterol excretion. 206. and Shetty. T. and Koike. L. J.. Res. Effect of fiber on protein. Physico-chemical properties of wheat bran and long term physiological effects in healthy man. et al. Miettinen.. Bam. Comparative laxation of psyllium with and without senna in an ambulatory constipated population.. A. Wagner.. 81. 1983. J. Stevens. Off. P.. Dig. J.. Nutr. . J. J. L. Oi.. Gastroenterol. Clin.. Adkins. 219. 16. H. Holtug. and Neilson. J. Melcher. P. Osa. 32. 323.. 909. Effects of rice fiber on fecal weight. T. N.. Nelson. T.. (Tokyo). Villaume. 1988. J... 1988. Peterson. Comparison of the effects of psyllium and wheat bran on gastrointestinal transit time and stool characteristics.... Am. Am. T. McNamara.. 3RD EDITION 204. Sato. 123. J. S. The effect of dietary guar on serum cholesterol. H. and Tarpila. Wheat fiber and laxation: dose response and equilibration time. J.. and Bianchi. 1987... Bowel function of healthy men consuming liquid diets with and without dietary fiber.. Kurpad. U. Anal. Defic. L. 218. K. B. Entr. P.. and Brydon. Okuda. D. J.. Sci. 1987. R. Bozzani.. S. T. Nutr. R. Nutr.. 85. Anderson. A. Nutr. Sieling. M. M. and Mehta. Eastwood. Sci. Fischer. Diet. W. Z. G. 223.. Serum lipids and cholesterol metabolism during guar gum. 1986. 1985. 1988. J. J. Yatsuda. H. J. 1984. 29. 32. Gastroent. Cancer. Clin. 2001 7:07 PM 250 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.. A. Velio.. R.. W. Levitt. Vitaminol. 40. Li.. A. A. Scand. Am. 41. S.. The relation between bacterial degradation of viscous polysaccharides and stool output in human beings.-G. 82. Rasmussen. S. 208. O’Donnell. 1986.. Nutr. plantago ovata and high fibre treatments. D.-J. E. 1991. K. J. W. and fecal output in man.. 47.. 33. J. 1987. D.2387_ch4. Sugawara. C. Robertson. B. 212. 183. A. intestinal transit. L. 1988. et al. 207. Dis. M... Andersen. 225. Clin. and Koishi. 1259... May 6. Slavin. J. W. 1989. Hypocholesterolemic effects of oat-bran or bean intake for hypercholesterolemic men. 216. Assoc. J. 9. D. A. Food Hydrocolloids. Vitaminol. A. and Bass.. K.. N. 221. 109. Yokoyama.. Kaneko. Abraham. J. Am. 215. 664. U. 406. 37.. Van Soest. J. J. J. 33.. Am. P. Three-week psyllium husk supplementation: effect of plasma cholesterol concentrations. Acta. Nutr. P. 60. Thorne.. Vitaminol. 224. Story. Nutr. Jenkins. 207. Patrow. 1988. and Furda. A.. J. Devries. J. 317. Nutr. flora.. J. 467... A.. D. 210. and Bass. Effect of fibre supplementation and activated charcoal on faecal parameters and transits in the tropics. 22. Determination of insoluble. D. S.. Marlett. F... Miyoshi. Prosky. 217. S. Nutr. apparent digestibility of energy. and Read. 14. M. Ranzi. Y. E. 993. May 6. J. Davis. A. and Cardozo. N. 243. Eds. 52. New York. J. Schneeman. A.... 249.. Clin. Consumption of prunes as a source of dietary fiber in men with mild hypercholesterolemia. A. Furda. Johansson. Marlett. P. E.. J. Dietary fiber: in vitro methods that anticipate nutrition and metabolic activity in humans. G. The effects of dietary gum tragacanth in man. and Brassard.. using high-performance liquid chromatography to measure neutral sugar composition. Eds.. New York. and Anderson. B. G.. and Hicks. 250. 231. Reddy. 210. Lett. Johansson. R. Adiotomre. 51. J. N. 1987. J. J. Rutgeerts. Li. and Brine. U. Composition of faeces from human subjects consuming diets based on conventional foods containing different kinds and amounts of dietary fibre. A comparison with gas-liquid chromatography. New York. 128.. 2001 7:07 PM THE EFFECT OF DIETARY FIBER ON FECAL WEIGHT AND COMPOSITION 251 227. A. M. L. 244. W.. Brydon. R.. Contam. A comparison between detergent and nondetergent analyses of dietary fiber in human foodstuffs.. G.. 24. Eastwood... N. and Brine. A. and Wynder. F. 1989. 17. 1990. T. T. R. Nutr. 1991. D. and Van Soest.. and Wong. 2.. in New Developments in Dietary Fiber. C. W. C. A.. D. V. Simi. 27. Prev... R. A. Clin. 476... 245. Am... P. Agric.. Tomlin. 238.. 283. 18. Clin. G. L. Therap. 3. Shift from a mixed to a lactovegetarian diet: influence on acidic lipids in fecal water—a potential risk factor for colon cancer. J. A. 992. J. Nutr. O. H. Eds. Rapid enzymatic assay of insoluble and soluble dietary fiber. 63. 1986. E. M. V.... M. and Shetty. 240. 1990. and Waggoner. B. J. Food Chem. and Schurmans. 230. 1990. 17. high fiber diet on fecal bile acids and neutral sterols. Colorimetric method for routine measurement of dietary fibre as nonstarch polysaccharides. M. J. G. Eur. Am.. Studies on dietary fiber. M.. Effects of antimicrobial therapy on faecal bulking. Gut... 589... Johansson. W. and Sohostrom. Br.. 64. 1990.. W. Barnard. Eriksson. W. 64. Y.. 73. C. V. M. J. . Kesaniemi. 1988.. Effect of low fat. J. 513. S. 239. Hiele. in New Developments in Dietary Fiber. The effect of long-term fibre and starch intake by man on faecal bile acid excretion. M. 51.. Nutr. J. A. Cancer.. C. Gallaher. 43. 1990... 14. Improved procedures for analysis of dietary fiber. 235.. Nutr.. M. 1984. Food Chem. H. Plenum Press.. J. and Westerlund. 242. Increasing starch intake in the human diet increases fecal bulking. K. Nutr. A.. Food Chem. Vantrappen.. W. and fecal lipids in middle aged men. 4. Am. 21. Analysis of dietary fiber. W. 246.. 232. 1259. J. J. B. Eastwood.. L.. L. Neilson. 53. H. 118. Nutr.2387_ch4. J. D. I. Improved methods for analysis and biological characterization of fiber. Engle. Englyst. A.. Furda. and Kurpad. Asp. 50. Med.. and Read. Allinger. Mongeau. J. Toxicol. Agric.. L. Plenum Press. Kurpad. and Marleft. Shetty. J.-G. Tinker.. P. S. Ghoos.. and Brydon. 1986. 1990. Animal Feed Sci. K. O’Brien. Food Sci.. 23. 432. 330. 1007. 1988.-G. 248. C.. D. C. Forsum. Tomlin. Pharmacol. 1987... A. 1986.. The effect of resistant starch on colon function in humans. 1983. A comparative study of the effects on colon function caused by feeding ispaghula husk and polydextrose. Pritikin. 34.. Food Chem. Simplified method for the determination of total dietary fiber and its soluble and insoluble fractions in foods. and Hudson. 234. G. N. 229. Hallmer. 1990. C.. high carbohydrate. Shift from a mixed diet to a lactovegetarian diet: influence on some cancer-associated intestinal bacterial enzyme activities. 171. and Anderson. Miles.. J. and Read. Gustafsson.. Modifications of the AOAC total dietary fiber method. 237. and Rafter. M. 228. 1333. 55. J. Low vs high dietary fiber and serum. S. N. Eastwood. P.. 1998. Goranzon. Nutr. Food Add. Plenum Press. O. J.. J. Clin... Am. L.. 31.4pt3_fm Page 251 Sunday. Clin.. 245. and Miettinen. 1342. J. A. Y. 1990. Jeraci. 1. W. N. P. J. 31. Effect of dietary fiber on the metabolizable energy of human diets. Am... 1075. Aliment. J. Tech. A. A. J. A rapid method for the determination of soluble and insoluble dietary fiber: comparison with AOAC total dietary fiber procedure and Englyst’s method.. Nutr. Nutr. Ottova. S. 128... 236. in New Developments in Dietary Fiber... J.. C. 239. Lee. The dietary effects of xanthan gum in man. J. P. and Siljestrom. biliary. Invest. Tarpila. W. I. Theander. C. Agric. 237. Br. J.. Edwards. and Gustafsson.. E. Kelsay. B. 1989. Clin.. 1983.. 233. 1986. I. 247. G... Furda. J. and Brine. A. 241. S. G. S. Brydon. P. 1988. P. N. Wood. Determination of dietary fibre—collaborative trials Part IV.. 247. R. S. Nutr. insoluble and total dietary fibre. J.. P.. F... N.. 256. D. 1989. A548... N. 2. and Cummings. and Cummings. S. 1992 (in press). Nutr.. H. 252. Kingman. A comparison of the Englyst and Prosky procedures for measurement of soluble. 90. J. Collinson. 1992 (in press).. A.. J. S. and Rambaud. N... A. H. and Cummings. S.. Dietet. Classification and measurement of nutritionally important starch fractions. Collinson. H. Dietet.. M.. Publ. 111. E.2387_ch4. Assoc. S.. H. Flourie. J. 1. H. 253. T. A. J. Englyst. H.. Southgate. Kingman.. Bingham. Hum. May 6. J. Colonic metabolism of wheat starch in healthy humans. 102.. Florent. Cummings. Beatty. H. vegetables and nuts. R. Bingham. Englyst. B. 255. J. Clin. 1986. Analysts.. 1992.. Etanchaud. Hum. C. J..4pt3_fm Page 252 Sunday. S. H. Runswick. A. Runswick. J. N. . E.. Gastroenterology. H. Englyst. Laxative properties of resistant starches. and Cummings. European J.. S. Thivend. E. Gastroenterology. A... Dietary fibre (non-starch polysaccharides) in fruit. Englyst. 3RD EDITION 251. Jouany. and Englyst. 2001 7:07 PM 252 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION... Dietary fibre (non-starch polysaccharides) in cereal products. 254. H. J. 253.. Bingham. C. Nutr.. Effects on fecal outputs and clinical symptoms. 1988. A. 5.1 and 4. EPIDEMIOLOGICAL CORRELATIONS A well-executed epidemiological study6 on two Scandinavian populations showed lower colon cancer incidence in the population with an average FWW of 200 g/d compared to the group with an average of 150 g/d. When data correlating TT to FWW are presented for individual subjects (Figure 4. and that a more predictable colon function is preferable.5 Correlations of Transit Time to a Critical Fecal Weight (CFW) and to Substances Associated with Dietary Fiber Gene A.5. 2001 7:09 PM CHAPTER 4.2). and that the CFW is a basic physiological concept.2 summarize these concepts. May 6. This supports the concept that there is a CFW. there is a critical fecal wet weight (CFW) of about 200 g/d. The line correlating TT to FWW becomes practically asymptotic to the abscissa for this FWW.00+$1. as it gives a simple method to help individuals determine whether their diet supplies enough fiber and enough of the right kind of fiber. that its value is in the range of 200 g/d for adults.5_fm Page 253 Sunday. Spiller and Monica Spiller THE CONCEPT OF CRITICAL FECAL WEIGHT (CFW) The correlation of transit time (TT) to fecal wet weight (FWW) is important. While we do not know the ideal transit time for humans.5. one finds a wide scatter of points correlating TT to FWW below FWW values of about 180 g/d. In a more general way. many values for TT are as high as 4 or 5 days. Several studies1–5 have shown that (1) TT decreases rapidly as FWW increases to about 200 g/d and (2) beyond FWWs of about 200 g/d there is little change in TT as FWW increases.50 © 2001 by CRC Press LLC 253 . This indicates that as FWW values increase. 0-8493-2387-8/01/$0. beyond which practically no TT is greater than 2 days. it is useful in arriving at desirable recommendations for fiber intake on various diet patterns and for different populations. Figures 4.2387_ch4. it may be reasonable to assume that extremely long transit times are not desirable. subjects’ TT decreased from 42 hours at baseline to 28 hours after a week of adaptation to the raisin diet. 2001 7:09 PM 254 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. which is present in grapes and their products such as sun-dried raisins and in tamarinds. When 120 g of raisins were fed.5.2 curve B is from Connell.5_fm Page 254 Sunday. The question of how much of the water-soluble fibers we need in relation to their effect on lipid and carbohydrate metabolism has not yet been answered. but it is certainly logical to assume that a fraction of the fiber should be of the watersoluble type. Pharmacological uses of dietary fiber and their effective levels are not part of these considerations. may decrease the amount of fiber needed to achieve a desirable TT. When the same subject consumed the amount of tartaric .7 and tartaric acid appears to play an important role. especially wheat-type fiber. May 6..8 and curve C is from Burkitt et al.1 Correlation of transit time to fecal weight in three different studies.1 CORRELATION OF CFW TO LEVELS OF DIETARY FIBER AND ASSOCIATED COMPOUNDS The amount of fiber needed to achieve the CFW appears to be in the range of 35 to 45 g/d. The effect of various amounts of raisin on TT is greater than expected by their fiber content. A typical example of such compounds in common foods is tartaric acid. Wheattype fibers are high in water-insoluble fractions. TARTARIC ACID: EXAMPLE OF FACTORS THAT WORK WITH DIETARY FIBER TO AFFECT COLON FUNCTION Many plant foods contain substances other than dietary fiber that affect colon function.2387_ch4. perhaps 30 to 40%. Curve A is from Spiller et al. 3RD EDITION 180 160 140 RANGE OF FECAL WEIGHTS AT WHICH LINES BECOME ASYMPTOTIC TO ABSCISSA TRANSIT TIME (TT) (hours) 120 100 80 60 A 40 B 20 0 C 30 60 90 120 150 200 250 300 400 500 FECAL WEIGHT (g/day) Figure 4. as long as enough cereal fiber is included (probably 60 to 70%). such as tartaric acid in grapes and raisins. It also appears that other substances present in plant foods. which was increased by the raisins. it did not significantly affect FWW.5.5_fm Page 255 Sunday. In this way. on both the raisins and the tartaric acid. whole grains such as brown rice. the TT decreased from 42 to 31 hours. as presented in the Appendixes of this Handbook. In addition. 2001 7:09 PM CRITICAL FECAL WEIGHT AND TRANSIT TIME 255 Figure 4. whole corn and oats. . This example was used to emphasize the importance of considering dietary fiber when present in whole foods as part of a complex matrix of bioactive compounds. OVERALL DIETARY PATTERN AND TRANSIT TIME Part of any recommendation on the intake of dietary fiber to obtain ideal transit times should include the importance of eating a variety of dietary fibers.2 Scatter of points for individual subjects below and above a critical threshold (CFW). While tartaric acid affected TT.2387_ch4. More studies are needed to determine other non-fiber substances in plant foods that may work together with fiber to normalize colon function. The data on dietary fiber content of various foods. acid equivalent to that present in the raisins. indicate that it is difficult to obtain sufficient dietary fiber from a single food (except for concentrated sources) and that the easiest way to obtain a reasonable amount of dietary fiber in the daily diet is to consume a diversity of unrefined plant foods. feces were softer and easier to eliminate. in the form of breads and prepared cereals. such as whole grain products. both water-insoluble and water-soluble components will be represented. May 6. J. 5.5_fm Page 256 Sunday. while beans and oats deserve a special mention as a source of good mixtures of water-soluble and water-insoluble fiber. as there is a tremendous diversity of breads on the market.. REFERENCES 1. J. 207.. 2001 7:09 PM 256 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. G. 1977. 536. 659. Connell. G. A. G. Bruce. A. Eds. J.. Whole grain wheat deserves a special mention as a high source of insoluble dietary fiber.. J. 7. M.. Petro. and Spiller. R. Reily. 25. E.. Spiller. M. Dietary fiber and disease. In bread selection great care should be taken. 8. J.. International Agency for Cancer Research. G. Wong.. 1068.. New York. G. and Scala. J. and Kirsner. A. H. the amount and type of dietary fiber may vary greatly according to the level of extraction and type of flours used. H. S.. Spiller. Clin. A.. A. 2. 10. Story. minerals and steroids and gastrointestinal transit time in healthy young women. Am. B.. and fruits.. Beigler. A.. P.. Nutr. A. Wong. Whittam. Shiplet. J. A. and Briggs. Coll. and Scala. Am. R. Walker. 1974. J. 1977. 81. JAMA.. Spiller. P. Nutr.. Whittam. J. Nutr.. 3. Dietary fiber. W. in Fiber Deficiency and Colonic Disorders. M. i. 39.2387_ch4. 4.. 229. . A.. G. 14. M. 1982.. Nutr. Can fecal weight be used to establish a recommended intake of dietary fiber (plantix)?.. J. Rev. E.. 1986. S. H. Burkitt. Food Sci. 6. D. bulk and colonic activity. transit time.. Fiber. Lancet. vegetables. 3RD EDITION beans. Nunes. 1978. 778. 31. M. L. Rep. 116. J. Nutr... A. and Blake.... M.. Spiller. G. Effect of increasing levels of hard wheat fiber on fecal weight.. 1995. May 6.. The use of concentrated sources such as wheat bran (for the wheat-type fiber portion) or oat bran offers additional assurance of a sufficient daily intake of particular fiber types. fecal bacteria and colon cancer in two Scandinavian populations. A. Plenun Press. Alton. A. Int. nuts. Effect of tartrate and dietary fiber in sun-dried raisins on colon function in healthy adults. E. Chernoff. Spiller.. C. Furarnoto. 23. Some Asian populations obtain additional dietary fiber from sea vegetation. 1975. D. B. N. L. J. G. Shipley. and Painter. CRC Crit. Recent progress in dietary fiber (plantix) in human nutrition.. Correlation of gastrointestinal transit time to fecal weight in adult humans at two levels of fiber intake. The upper bowel.00+$1. and jejunum. Bacterial interactions and competition 9. Fiber has the potential to influence most of the factors listed above. Woods and Sherwood L. substrata. 2001 7:27 PM CHAPTER 4. and drugs) 8. but the major concentration reside in the large bowel. 3. The composition of the flora itself can affect many of these factors. intestinal antibodies. This complex interactive system is only now being explored. 5. The bacteria are washed down in a wave-like fashion. May 6.2387_ch4. Intestinal motility Individual microbial species vary in their sensitivity to changes in these parameters. has a sparse microflora derived largely from the oral cavity. including the stomach. Diet. Gorbach INTRODUCTION A variety of environmental and physiological conditions are known to influence microbial composition and metabolic activities of the intestinal microflora. 2.6_fm Page 257 Sunday. Across 0-8493-2387-8/01/$0.50 © 2001 by CRC Press LLC 257 . duodenum. bacteriocides. with a flora containing elements from the upper intestine. along with some coliforms and anaerobes from the colon.6 Influences of Fiber on the Ecology of the Intestinal Flora Margo N. and nutrient availability Redox potential Gas composition Acidity or pH Osmotic and ionic effects Surface tension and liquid flow Endogenous and exogenous substances that may inhibit bacterial growth (bile salts. up to 105 colony-forming units (CFU) per milliliter. The ileum is a transitional zone.1–3 Among the recognized factors within the bowel lumen are 1. 4. The organisms are found in the upper bowel in relatively low concentrations. 7. 6. INTESTINAL BACTERIA Bacteria are distributed throughout the intestine. volatile and nonvolatile fatty acids. and only partial data are available in humans. along with saliva. 2387_ch4. Total concentrations are 1011 to 1012 CFU per gram. Lactobacillus. (5) encephalopathy. coli Streptococci Bacteroides a Fusobacteria a Enterococci Bifidobacteria a Lactobacilli Eubacteria a Source: From Gibson. and (6) increased cancer risk. The majority of microorganisms present in the large intestine have a strictly anaerobic metabolism. 1401. (3) stimulation of immune functions. (3) infection. (4) improved absorption of nutrients (calcium.1 Morphologic Characteristics of Human Intestinal Microflora • Gram-negative anaerobic nonsporing rods Bacteroides Fusobacterium • Gram-negative facultative rods Enterobacteriaceae • Gram-positive cocci Streptococcus – facultative bacteria Peptostreptococcus – anaerobic bacteria • Gram-negative anaerobic cocci – minor contribution to flora in man • Gram-positive nonsporing rods Bifidobacterium – anaerobic bacteria Lactobacillus – facultative or anaerobic bacteria Eubacterium – anaerobic bacteria • Gram-positive spore-forming anaerobic rods Clostridium When evaluating the composition of the intestinal flora.4 Some of the general harmful effects that can be caused by bacteria include (1) production of toxins. J. neutral/potentially harmful. and (5) synthesis of B-vitamins that are available to the host. Neutral.6.4..4 The first four are saccharolytic. May 6. M. Nutr.6.4 A number of health-promoting actions have also been associated with some intestinal flora. R. or Beneficial Harmful/Pathogens (Some Species) Ps-Aeruginosa Proteus Staphlococci Clostridia Veillonellae Neutral/Potentially Harmful Healthful/Beneficial Peptostreptococci E.9 Another way of describing the intestinal flora is by morphological characteristics. 3RD EDITION the ileocecal valve there is a marked increase in numbers and types of bacteria. and healthful/beneficial flora. it is useful to keep in mind some general considerations regarding classifications of the most typical intestinal flora as harmful/pathogens. Bifidobacteria and Lactobacillus are the two common microflora most often described as desirable. Table 4. indicating that they can break down various complex carbohydrates present in the intestine and colon. 2001 7:27 PM 258 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and the number of facultative anaerobes are several orders of magnitude lower than those of the obligate anaerobes. B.1. approaching the theoretical limit of forms that can be accommodated in that mass.6_fm Page 258 Sunday. (2) decreasing gas formation.6. zinc. five genera account for the majority of the viable forms: Bacteroides. 125. With permission.8. Although over 400 species of anaerobic bacteria are present in the large intestine of a human being. (4) liver damage.2 Division of Common Bacteria into Harmful. which remain fairly consistent throughout the large bowel.. and iron). and Roberfroid. and Gram-positive cocci. Eubacterium. Table 4. Bifidobacterium. and they include (1) inhibition of growth of harmful bacteria. G. (2) diarrhea/constipation. including an example of typical species found in the human intestine.5–7 Bacteria account for 35 to 50% of volume of the contents of the human colon or 41 to 57% of the dry weight. Six major categories are listed in Table 4. 1995.20 . a Genera that account for the majority of viable bacterial forms found in the large intestine. A number of studies have looked at the advantageous physiological characteristics of Bifidobacteria and Lactobacillus. which results in lower levels of secondary bile acids in the intestinal lumen.18.13 It has also been reported that Bifidobacteria excrete a metabolic end product inhibitory to a number of pathogenic bacteria.3. B.3 Percentage of Microflora from Humans on Two Different Diets Bacterial Group Bacteroides Bifidobacteria Fusobacteria Clostridia Sucrose Oligofructose 72% 17% 9% 2% 16% 82% 1% 1% Note: The four groups represented were taken as 100% of the fecal flora. and during days 16–30. 1998.4 An example of the changes observed in the predominant bacterial groups in eight volunteers fed a strictly controlled diet for 45 days is shown in Table 4. Nutr.. intestinal bacterial substrates. as stated earlier in this text. M. During days 0–15. al. 11.6.12 Bifidobacteria produce strong acids and metabolic products (acetate.4 Certain dietary fibers have been labeled prebiotics.17 These secondary bile acids have been reported to be co-carcinogens. “potentially harmful” intestinal flora provides many useful research questions.16. lactate) that lower gut pH and exert an antibacterial effect directly. and food intake patterns that affect type and prevalence of intestinal organisms offer a second approach to enhancing beneficial bacteria. Probiotics are defined as live microbial food supplements which beneficially affect the host animal by improving its intestinal microbial balance.”23 Food components that are included in this definition are presented below: • Resistant starch • Non-starch polysaccharides (NSP) • Cellulose • Hemicellulose • Pectin • Gums • Non-carbohydrate aromatic polymers • Lignin • Non-digestible oligosaccharides • Fructooligosaccharides • Galactooligosaccharides • Glucooligosaccharides . et. as reviewed by Goldin and Gorbach.21.11 Bifidobacteria may constitute up to 25% of the total population in the gut of adults. the supplementation was 15 g of oligofructose. it is generally agreed to include the components of plant cells which are “resistant to hydrolysis by the digestive enzymes of humans.20 Table 4. Source: From Roberfroid.19 Alternatively. J.22 Probiotics are reported to have many positive effects on numerous medical conditions.6_fm Page 259 Sunday.11.2387_ch4.2 lists some common bacteria in each of the three categories of desirability of the microflora..10.19 Understanding the factors that encourage the presence of these “beneficial” vs. 2001 7:27 PM INFLUENCES OF FIBER ON THE ECOLOGY OF THE INTESTINAL FLORA 259 Table 4. the diet was supplemented with 15 g of sucrose. DIETARY FIBER Definitions of fiber are broad but. Enhancing beneficial bacteria has become a new food technology and market opportunity termed probiotics.14.6.15 An increase in Bifidobacteria also results in lower levels of the bacterial enzymes glucosidase and β-glucuronidase in the bacterial milieu. 128. A non-digestible food ingredient that beneficially affects the host by selectively stimulating the growth and/or activity of one or a limited number of bacteria in the colon and thus improve host health is termed a prebiotic. With permission. investigations into intestinal conditions. May 6. notably the fructooligosaccharides.6. The binding capacity of fiber may determine its availability for metabolism by the flora. dietary fiber includes a heterogeneous mixture of materials. and lignin. 4.” An alternative approach is the investigation of specific. and inulin (fructooligosaccharides). and legumes. . and transit time in the bowel. Pectin is used widely in the food industry to obtain certain properties of texture and binding. however. They are available under the product names of RAFTILOSE® (oligofructose) and RAFTILINE® (inulin). water-holding properties. the addition of these fiber components to standard food products would allow labeling of their health benefits and advertising as “functional foods. This is in contrast to information on hemicellulose.25 Foods in our usual diet that are high in dietary fiber include fruits. have higher water-holding capacity. 2001 7:27 PM 260 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Insoluble fibers. 3. Cation-exchange characteristics: The ability to bind endogenous and exogenous materials in the intestine is related to the ionic binding properties of the fiber. and legumes.24 while nondigested carbohydrates account for 10–60 g/day. 3RD EDITION Most of these food components or undigested residues are good substrates for intestinal flora. Studies on fructooligosaccharides. There is great variation in fiber amount and type within each food group.S. May 6. It is of interest to note that the direction of this research appears to be determined by the availability of pure or standardized forms of the dietary components and their usefulness to the food industry. vegetables. Among these properties are the following: 1. If benefits are observed.2387_ch4. Digestibility or degradability: This depends on the composition of the fiber. Hemicellulose — 50 to 80% digested by the flora D.6_fm Page 260 Sunday. cellulose. commonly consumed foods that contain high levels of these specific types of fiber and their role in supporting and/or maintaining health or markers of health. These are more prevalent in grains. banana. which are readily available. Pectins and hemicellulose are generally considered soluble fiber. Particle size: Particle size determines the surface area of the fiber and alters its susceptibility to digestion.25 They are defined by the degree of polymerization (DP). They are naturally found in such foods as garlic. Inulin is on the “Generally Recognized As Safe” (GRAS) list. Water-holding capacity: The swelling and permeability properties can influence the ability of bacteria to infiltrate and digest the fiber by changing the surface area available for digestion and the speed of transport through the intestine. whole grains. and are derived from the tougher structural components of plants. and are in higher quantities in fruits. have been used for some time in food manufacturing. artichoke. as such. Inulin is obtained by hot water extraction of chicory root and. Oat bran is a good example of a soluble fiber. onion. A. As can be seen. is an ingredient in a number of food products due to its fat-like properties and its sweet taste (1/3 the sweetness of sugar). Lignin — not digested by the flora B. Pectin and gums — 90 to 100% digested by the flora E. some of which can influence the intestinal flora. then they are termed oligofructose. have a lower waterholding capacity. hemicellulose. cellulose. form gels in the intestines. and with an average DP of 12 (but up to 60). The fructooligosaccharides are the most widely studied of the oligosaccharides.25 and oligosaccharides provide 1–10 g of substrate per day. binding properties. Cellulose — 30 to 50% digested by the flora C. 2. and wheat. Its physiological effects depend on specific properties. Standardized forms of wheat bran have also been developed to facilitate studies investigating their potential benefits. are not water-soluble. Non-digestible oligosaccharides — 95 to 100% digested by the flora Most of the research investigating the effect of dietary fiber on intestinal and fecal flora has been carried out with wheat bran. vegetables. cellulose and lignin.3 Resistant starch provides 8–40 g/day of substrate to the large intestine. they are termed inulin. or lignin. which appear to be influenced by the amount of lignin present. These provide pectins. which are difficult to obtain in their pure form with their physiological properties intact. Estimates of daily intake of dietary fiber in the U. as NSP + lignin are 8–18 g/day. If the DP is <9. leek. Wheat bran is an example of an insoluble fiber. asparagus. pectin. 6. A rating of 3+ would indicate a higher level or increase in SCFA. The increase in SCFAs is generally agreed to be beneficial. have been studied. Fructooligosaccharides have also been studied in a reasonable number of controlled experiments (N = 8 studies). 4. due to their association with decreased absorption of intestinal cholesterol and other steroids. May 6. or high-risk diet (high fat.19 A rating of 5+ would indicate a lower level or decrease in one or both of these enzymes. . low fiber) reported higher levels of β-glucuronidase or dehydrogenation or steroids and increased secondary bile acids. 2.4 indicates that the Western-type or high-risk diets (high fat. In the first approach.6. since it would indicate a decrease in formation of secondary bile acids (lithocholic and deoxycholic acids). Increase in the bacteria generally recognized as desirable.6.) 5. Therefore. and serve as substrates for colonic cells. the intestinal flora has been measured in people eating customary diets with different levels of fiber intakes (Table 4. Production of short-chain fatty acids (SCFAs).” A rating of 1¯ would indicate the reverse. “undesirable. 2001 7:27 PM INFLUENCES OF FIBER ON THE ECOLOGY OF THE INTESTINAL FLORA 261 In order to evaluate the potential benefit or harm of specific dietary fibers on the intestinal flora. and 20 would indicate no difference. For example. A decrease in 7-α-hydroxylase would also be considered a positive observation. omnivore diet.4). Production of immunostimulants. A rating of 2¯ would indicate a higher pH. and 30 would indicate no difference.6.14. or Western and Japanese people eating their customary diet. A rating of 4+ would indicate that the results of the study supported the excretion or presence of a metabolite that decreased known microbial pathogens. A decrease in bacterial glucuronidase and/or glycosidase would be considered a positive change. and 50 would indicate no change. The Western diet. A rating of 3– would indicate a lower level or decrease. and 10 would indicate no difference. such a change caused by the presence of specific fibers or fiber in general would be considered a beneficial change. and proprionic. will be the focus for the evaluation and rating of the cited literature. It has been reported and previously stated that Bifidobacteria excretes a metabolic product that results in the decrease in pathogenic flora. An increase in Bifidobacteria or Lactobacillus and/or the decrease of bacteria generally recognized as “neutral or potentially harmful” (Bacteroides and Fusobacteria) is considered desirable.5 supplied data on this topic. there is more information on the effect of wheat bran on the intestinal flora. low fiber) generally showed higher levels of Bacteroides and sometimes lower levels of Bifidobacteria. Four studies reported on bacterial enzymes. vegetarians and non-vegetarians.4 or 4.5). Change in intestinal pH.15 A decrease in known pathogens could be an indicator of this capability. A decrease in the pH of the intestinal contents is deemed to support an increase in bifidobacteria and to generally decrease pathogens. 3. which represent our current understanding of the biochemistry and physiology of intestinal flora.” The factors given below. The most typical acids formed are acetic.2. lactic. act as antibacteriacidals. Thus. (No study reported in Tables 4. one needs to identify criteria that are generally accepted as “beneficial” vs. Two approaches have been taken in studies of fiber and its effect on the intestinal microflora. since they decrease the pH. A rating of 5¯ would indicate a higher level or increase in the enzyme. Evaluation of data from Table 4.6. A rating of 1+ indicates a higher level or increase of one or both of the “desirable” microflora and/or a lower level or decrease in the “undesirable. which are considered co-carcinogens. A reasonable number of human experiments have been performed with fiber sources such as wheat bran (N = 8 studies).2387_ch4. Justification of this position was previously stated and is related to the ability of Bifidobacteria to decrease pathogenic organisms and promote physiological properties such as increased immune response. The second approach involves a defined diet or a controlled diet in a study population to which the fiber of interest is added as a supplement (Table 4. A rating of 2+ would indicate a lower intestinal or fecal pH.6_fm Page 261 Sunday. 1. butyric. Changes in bacterial enzymes. 3RD EDITION . Lactobacilli. May 6. Copenhagen No change in predominant fecal flora English had 30-fold more Bacteroids and 4-fold more Bifidobacterium. Hawaiian (20) Low-risk colon cancer Seventh-Day Adventists British (91). Vegetarians (17) High risk. Wales (65). Scotland (11). and Lactobacillus) Fecal Microflora Characteristics 36 1+ 39 40 5+ 38 1+ 1° 37 — 1° 35 1° 34 33 1+ 1° 32 30 5+ 5+ 29 5+ 31 28 1+ — 26 27 Ref. rural Finns High-risk colon cancer. South Indians. Western diet eliminated Eubacterium contortum No consistent pattern between types or numbers in fecal samples. Enterococi. 7-α-dehydroxylase was higher in omnivores compared to lacto-ovo vegetarians Western population had greater numbers of Clostridium paraputrificum than non-Western groups A positive sign indicates more “desirable” microflora characteristics in the Non-Western Diet group compared to the Western diet group. 1° 1+ Scorea 262 Nonvegetarian-Seventh-Day Adventists Vegetarian Ugandans (48) Western English Non-Western Diet (N) Fecal Microflora in Different Human Populations Western Diet (N) Table 4. Seventh-Day Adventists (11).6_fm Page 262 Sunday. 2001 7:27 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Chinese (11) Japanese diet in Japanese (1) Low risk Traditional Japanese (15) Japanese. and azoreductase were higher in the omnivores compared to the vegetarian groups.4 2387_ch4. American (34) British Western (17) American diet in Japanese (1) High risk Western Japanese (18) North American High-risk colon cancer Non-Seventh-Day Adventists Low risk. Ugandans had significantly greater numbers of Strepotococci. β-glucuronidase.Ugandans. nitroreductase. and yeasts British and Americans had greater numbers of Bacteroides and a higher ratio of anaerobes Vegetarians had fewer isolates of Bacteroides that could dehydrogenate bile acids Westerners had greater conversion of bile acids to secondary bile acids and 2. U.S.S. Clostridium septum. U. Africa Lacto-ovo vegetarians (13). Streptococcus. higher Streptococcus faecalis. population.S. great differences between individuals High-risk group had more anaerobes (Bacteroides and Fusobacterium).6. tertim. urban Danes. C. and Fusobacterium were lower in the Adventist group Adhering and nonadhering Seventh-Day Adventists showed little differences High-risk group had higher ratio of anaerobes due to decrease in aerobes (Enterococcus. and fewer Bifidobacterium. Hong Kong.S. Uganda England (36). (15) Western population had higher levels of Bacteroides and Bifidobacterium Fecal bacterial enzymes. U. (8) a Low risk colon cancer. Japan. American vegetarians (12).5 times more β-glucuronidase activity Western diet resulted in disappearance of Fusobacterium (10-month study) High-risk populations had higher levels of Clostridium paraputrificum which was able to dehydrogenate steroids Traditional diets have more aerobes and facultative bacteria. low-risk had more Eubacterium Seventh-Day Adventists had higher Lactobacillus and lower Clostridium perfringens than the U. Japanese Strict vegetarians Japanese. this suggests an increase in bacteria capable of digesting carrots or an induction of enzymes present in the flora No significant difference in qualitative or quantitative count of bacteria.18 g DF 18 g DF 30 g 30 g British diet plus Wheat bran Cabbage Western diet plus Bran Wheat germ — — — — American diet plus Laminarin Xylan Guar gum Psyllium hydrocolloid 200 g. Plant sources cited resulted in induction of enzymes in Bacteroides species that were capable of degrading the polysaccharides Usual Diet with Fiber Supplement In vitro incubation of colon flora 10 14 3 3 11 6 astronauts Low-Residue Diets No. 2001 7:27 PM INFLUENCES OF FIBER ON THE ECOLOGY OF THE INTESTINAL FLORA 263 . and Enterococci Disappearance of Bifidobacterium and increase in E. decrease in Enterococci in lowresidue diet Diverse microbial population reduced to bacteroids. of subjects 43 1– 5+ — — 1° 5° 1° — 40 50 49 48 47 46 45 44 42 1– — 41 Ref. raw — Low-residue diet British diet plus Carrots — Elemental-low residue 1 oz — Chemically defined diet American diet plus All Bran 9 days — 1–2 months 3 weeks 3 weeks 3 weeks 7 days 12 days 10 days 15 days — Low-residue diet (glucosebase) Vivonex low-residue diet 21–56 days — Duration Low-residue diet Amount Effect of Various Fiber on Fecal Microflora Diet and Fiber Table 4.6. coli. ratio of anaerobes to aerobes appeared to increase. 1° Scorea 2387_ch4. no change was seen until 10 days after the start of eating carrots. Enterococci and other lactic acid bacteria decreased in number Decrease in Enterococci.5 4 5 6 5 6 No significant change in fecal flora. decrease in very oxygen-sensitive anaerobes Enterobacteria increased. May 6. no change in total anaerobes. aerobes. or coliforms No change in fecal flora Fecal Microflora Characteristics Addition of fiber led to a reduction of fecal 7-α-dehydroxylase activity Cabbage stimulated growth of intestinal flora Total breath hydrogen increased twofold by the third week.6_fm Page 263 Sunday. coliforms. 6 (p < .01). 7-α-dehydroxylase increased and cholesterol dehydrogenase decreased. 3RD EDITION . slight increase in Streptococcus during fiber supplement. slight decrease in Lactobacillus during fiber supplementation No change in relative number of the following: Enterobacteria. Lactobacillus. 21-day test diet 12 weeks 3 weeks 3 weeks 14-Mar 14–70 days Duration 1 oz 12–26 g Amount 6 5 20 20 4 4 3 5 4 6 7 24 # of subjects Pectin caused a decrease in anaerobic/aerobic ratio from 3. increased volatile fatty acids were observed in fecal samples of persons on the fiber diets for more than two weeks.5 g 25 g 15 g British diet plus Wheat bran Western diet plus Bran Bagasse British diet plus Gum arabic (GA) American diet plus Pectin 21 days 7-day control. increase in total bacteria excreted associated with increase in stool weight Increase in anaerobes in high-fat diet. Fecal Microflora Characteristics 5– — 1° 58 57 56 55 54 1– 1° 53 52 51 Ref. Bifidobacterium. Enterococcus. but neither reached significance Flora not tested — effect on metabolism tested.6_fm Page 264 Sunday. no change in ability to digest bran Fecal samples from persons on fiber-supplemented diets showed no difference in the in vitro digestibility of the individual fiber.6. Clostridium.5 (Continued) Effect of Various Fiber on Fecal Microflora 2387_ch4. May 6.6 to 2. Eubacterium.2 weeks 35 g/day 35 g/day 1000 g/day 1000 g/day 3 oz/day (27 g dietary fiber) 11. GA caused a significant increase in breath hydrogen after chronic but not acute treatment with GA. or Streptococcus No significant difference in composition of flora No change in flora. this might suggest that endogenous sources of nutrients and not diet are the energy source of flora. this suggests increases in number of flora capable of metabolizing GA or induction of bacterial enzymes No change in composition of the bacteria. no change in aerobes. 2001 7:27 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Bacteroides.7 g of crude fiber 39 g 10. however. 1° 1° 3+ Scorea 264 Western diet plus Wheat bran British diet plus Pectin Guar Plantains Bananas American diet plus All Bran British diet plus Coarse bran Diet and Fiber Table 4. only BFM alone.6_fm Page 265 Sunday. perfringens Oligosaccharides (Fructooligosaccharides) 12 females 5 (M) 8 (M) 1+ 1+ 1+ 1˚.5 0. no change in pH.6 15 g (30–40 g 1 day) 18-day cycles SCFA production Baseline NC NC Decrease in total SCFA Pectin decreased fecal pH 35% increase of β-glucuronidase on pectin diet No effect of pectin on 7-α-dehydroxylase (question of adequacy of fiber supplementation and wash-out times) Only pectin decreased fecal pH Indirect effect on flora via changes of pH which can affect species and enzymatic activity 8 8 4 12 Hyperlipidemics __ Increase of Bifidobacteria. 30% fat and 0 gm DF) plus Cellulose Pectin Xylan Corn Bran Controlled diet (3×3 design) Low-fiber diet High-fiber/high-protein High-figer/low-protein Barley fiber 18 g/day 15 g/day 15 g/day 4 g/day 8 g/day days days days days days 12 days 15 15 15 38 12 5 weeks 5 weeks weeks weeks weeks weeks 8 g/day 3 3 3 3 2 weeks g g g g 20 34 36 23 wgt wgt wgt wgt 9-day cycles 8 g/day g/kg g/kg g/kg g/kg 0.placebo Sucrose (placebo) Typical Western diet plus Oligofructose Controlled diet plus Oligofructose Inulin Typical diet (France) +BFMb Inulin Japanese diet plus Oligofructose (OF) (neosugar) Japanese diet plus Oligofructose Basal diet (12% protein. decrease in Bacteroides Increase in Bifidobacteria with BFM No increase in Bifidobacteria with BFM and inulin. 260 mg cholesterol. May 6. 5 g NDF) plus Pectin Basal diet (. 33% fat.8 g kg wgt of egg albumin. decrease in C.5 0. nitrate reductase. 5˚ 66 65 64 63 1+ 1+ 62 61 60 59 1+ 3° 3° 3– 2+ 2+ 2+ 5– 5° 2387_ch4.5 0. 2˚. nitro-reductase or azoreductase Significant increase in Bifidobacteria with oligofructose Fivefold increase in Bifidobacteria after oligofructose and decrease in Bacteroides and Eubacteria Tenfold increase in Bifidobacteria. 2001 7:27 PM INFLUENCES OF FIBER ON THE ECOLOGY OF THE INTESTINAL FLORA 265 . decrease in Bacteroides Increase of Bifidobacteria. 266 a Controlled diet — baseline plus Fn type oligosaccharide (RAFTILOSE®) Usual diet plus oligofructose Controlled diet (CD) plus oligofructose (OF) (neosugar) Lactore Typical diet (American) plus Inulin Diet and Fiber Table 4.b Increased Bifidobacteria on CD Increased Bifidobacteria by one log OF decreased β-glucuronidase OF decreased glycocholic acid hydroxylase OF resulted in no change in nitroreductase 1–6 days 7–32 days 4 g/day A positive sign indicates that the tested fiber supplement restulted in more “desirable” microflora characteristics. decreased pH Increased Bifidobacteria. decreased pH Decreased Lactobacilli and increased Enterococci No change in pH or β-glucosidase or β-glucuronidase 3 weeks Increased Bifidobacteria and decreased Enterobacteria Fecal Microflora Characteristics 8 g/day 10 10 15 15 8 (M+F) # of subjects 8 g/day g g g g Duration 1–8 days 9–19 days 1–8 days 9–19 days 2 weeks 2 weeks 20 40 20 40 Amount 1+ 1+ 5+ 5+ 5˚ 1+.5 (Continued) Effect of Various Fiber on Fecal Microflora 2387_ch4. 3RD EDITION . May 6. 5˚ 1+ Score 69 68 67 Ref. 2001 7:27 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.6_fm Page 266 Sunday. 2+ 1+. 12 Increased Bifidobacteria. 2+ 1– 2˚. Bifidoborterium of fermented milk.6. 2387_ch4.6_fm Page 267 Sunday, May 6, 2001 7:27 PM INFLUENCES OF FIBER ON THE ECOLOGY OF THE INTESTINAL FLORA 267 In Table 4.6.5, the use of low-residue diets caused a decrease in Bifidobacteria and a general indication of a rise of less desirable microflora. Studies of fiber supplementation with wheat bran indicated no change or a decrease in beneficial bacteria (Lactobacillus). One study with bran40 reported a reduction of fecal 7-α-dehydroxylase. Studies using oligosaccharides (N = 8) resulted in seven of the eight studies reporting an increase in the desirable over the undesirable microflora. The one study that did not report an increase in Bifidobacteria with inulin was designed to deliver inulin in a probiotic of Bifidobacteria. The addition of inulin with the probiotic did not increase Bifidobacteria over the level achieved with the probiotic alone. Four studies reported on changes in pH: two saw a decrease, and two failed to see a decrease. Two studies of fructooligosaccharides reported on fecal enzymes, and only one study69 reported a decrease in β-glucuronidase. Additional studies are needed to clarify the effect of fiber, including oligosaccharides, on bacterial enzymes, to determine the amount necessary to elicit desirable effects and the equivalent intake of usual foods (whole wheat, wheat bran, fruit, vegetable, etc.) that produce similar results. REFERENCES 1. Simon, G. L. and Gorbach, S. L., Intestinal flora in health and disease, Gastroenterology, 86, 174, 1984. 2. Goldin, B. R., Lichtenstein, A. H., and Gorbach, S. L., The role of intestinal flora, in Modern Nutrition in Health and Disease, 8th ed., Shils, M. and Young, V., Eds., Lea & Febiger, Philadelphia, 1994, chapter 38, 569–582. 3. Cummings, J. H. and Macfarlane, G. T., A review: the conditions and consequences of bacterial fermentation in the human colon, J. Appl. Bacteriol., 70, 443, 1991. 4. Gibson, G. R. and Roberfroid, M. B., Dietary modulation of the human colonic microbiota: introducing the concept of prebiotics, J. Nutr., 125, 1401, 1995. 5. Finegold, S. M., Flora, D. J., Attebury, H. R., and Sutter, L. V., Fecal bacteriology of colonic polyp patients and control patients, Cancer Res., 35, 3407, 1975. 6. Moore, W. E. C. and Holdeman, L. V., Identification of anaerobic bacteria, Am. J. Clin. Nutr., 25, 1306, 1972. 7. Reddy, B. S., Weisburger, J. H., and Wynder, E. L., Effects of high risk and low risk diets for colon carcinogenesis on fecal microflora and steroids in man, J. Nutr., 105, 878, 1975. 8. Salyer, A. A., Energy sources of major intestinal fermentative anaerobes, Am. J. Clin. Nutr., 32, 258, 1979. 9. Stephen, A. N. and Cummings, J. H., The microbial contribution to human fecal mass, J. Med. Microbiol., 13, 45, 1980. 10. Tamura, Z., Nutriology of bifidobacteria, Bifidobacteria Microflora, 2, 3, 1983. 11. Fuller, R., Ed., Probiotics, The Scientific Basis, Chapman and Hall, London, 1992. 12. Finegold, S. M., Sutter, V. L., and Mattisen, G. E., Normal indigenous intestinal flora, in Human Intestinal Microflora in Health and Diseases, Hentges, D. J., Ed., Academic Press, New York, 1983, 3–31. 13. Rasic, J. L., The role of dairy foods containing bifido and acidophilic bacteria in nutrition and health, N. Europ. Dairy J., 4, 80, 1983. 14. Gibson, G. R. and Wang, X., Inhibitory effects of bifidobacteria on other colonic bacteria, J. Appl. Bacteriol., 77, 412, 1994. 15. Gibson, G. R. and Wang, X., Bifidogenic properties of different types of fructooligosaccharides, Food Microbiol., 11, 491, 1994. 16. Marteau, P., Pochart, P., Flourie, B., Pellier, P., Santos, L., Desjeux, G. F., and Rombaud, J. C., Effects of chronic indigestion of a fermented dairy product containing Lactobacillus acidophilus and Bifidobacterium bifidum on metabolic activities of the colonic flora in humans, Am. J. Clin. Nutr., 52, 685, 1990. 17. Kulkami, N. and Reddy, B. S., Inhibitory effect of Bifidobacterium longum cultures on the azoxymethane-induced aberrant crypt foci formation and fecal bacterial β-glucuronidase, Proc. Soc. Exp. Biol. Med., 207, 278, 1994. 2387_ch4.6_fm Page 268 Sunday, May 6, 2001 7:27 PM 268 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION 18. Mallett, A. K. and Rowland, I. R., Factors affecting the gut microflora, in Role of the Gut Flora in Toxicity and Cancer, Rowland, I. R., Ed., Academic, London, 1988, 347–382. 19. Goldin, B. R. and Gorbach, S. L., Probiotics for humans, in Probiotics. The Scientific Basis, Fuller, R., Ed., Chapman and Hall, London, 1992, 3555–376. 20. Roberfroid, M. B., Van Loo, J. A. E., and Gibson, G. R., The bifidogenic nature of chicory inulin and its hydrolysis products, J. Nutr., 128, 11, 1998. 21. Fuller, R., Probiotics in man and animals, J. Appl. Bacteriol., 66, 365, 1989. 22. Fuller, R., Probiotics in human medicine, Gut, 32, 439, 1991. 23. Trowell, H., Definition of fiber, Lancet, 1, 503, 1974. 24. Federation of American Societies for Experimental Biology, Life Sciences Research Office, For the U.S. population, daily dietary fiber intake in grams, on 1 day, by sex, age, and race/ethnicity, in Third Report on Nutrition Monitoring in the United States: Volume 2, U.S. Government Printing Office, Washington, D.C., 1988–1991, 154. 25. Van Loo, J., Coussement, P., Leenhees, L. D., Huebrege, H., and Smits, G., On the presence of inulin and oligofructose as natural ingredients on the Western diet, Crit. Rev. Food Service Nutr., 35, 525, 1995. 26. Moore, W. E. C., Cato, E. P., and Holdeman, L. V., Anaerobic bacteria of the gastrointestinal flora and their occurrence in clinical infections, J. Infect. Dis., 119, 641, 1969. 27. Aries, V., Crowther, J. S., Drasar, B. S., Hill, M. J., and Williams, R. E. O., Bacteria and aetiology of cancer of the large bowel, Gut, 10, 334, 1969. 28. Hill, M. J., Drasar, B. S., Aries, V., Crowther, J. S., Hawksinth, G., and Williams, R. E. O., Bacteria and aetiology of cancer of the large bowel, Lancet, 1, 95, 1971. 29. Aries, V. C., Crowther, J. S., Drasar, B. S., Hill, M. J., and Ellis, F. R., The effect of a strict vegetarian diet on the faecal flora and faecal steroid concentration, J. Pathol. Bacteriol., 103, 54, 1971. 30. Reddy, B. S. and Wynder, E. L., Large bowel carcinogenesis: fecal constituents of populations with diverse incidence rates of colon cancer, J. Natl. Cancer Inst., 40, 1437, 1973. 31. Ueno, K. et al., Comparison of characteristics of gram-negative anaerobic bacilli isolated from feces of individuals in Japan and the United States, in Anaerobic Bacteria: Role in Disease, Balowz, A., DeHaan, R. M., Dowell, V. R., Jr., and Gage, L. B., Eds., Charles C. Thomas, Springfield, IL, 1974, 135. 32. Hill, M. J., Steroid nuclear dehydrogenation and colon cancer, Am. J. Clin. Nutr., 27, 1475, 1974. 33. Finegold, S. M., Attebury, H. R., and Sutter, V. L., Effect of diet on human fecal flora: comparison of Japanese and American diets, Am. J. Clin. Nutr., 27, 1456, 1974. 34. Moore, W. E. C. and Holdeman, L. V., Human fecal flora: the normal flora of 20 Japanese-Hawaiians, Appl. Microbiol., 27, 961, 1974. 35. Peach, S., Fernandez, F., Johnson, K., and Drasar, B. S., The non-sporing anaerobic bacteria in human faeces, J. Med. Microbiol., 7, 213, 1974. 36. Finegold, S. M., Sutter, V. L., Sugihara, P. T., Elder, H. A., Lehmann, S. M., and Phillips, R. L., Fecal microbial flora in Seventh-Day Adventist population and control subjects, Am. J. Clin. Nutr., 30, 1781, 1977. 37. International Agency for Research of Cancer, Intestinal Microbiology Group, Dietary fibre, transittime, faecal bacteria, steroids, and colon cancer in two Scandinavian populations, Lancet, 2, 207, 1977. 38. Drasar, B. S., Goddard, P., Heaton, S., Peach, S., and West, B., Clostridia isolated from faeces, J. Med. Microbiol., 9, 63, 1976. 39. Moore, W. E. C., Cato, E. P., and Holdeman, L. V., Some current concepts in intestinal bacteriology, Am. J. Clin. Nutr., 31, 533, 1978. 40. Goldin, B. R., Swenson, L., Dwyer, J., Sexton, M., and Gorbach, S. L., Effect of diet and Lactobacillus acidophilus supplements on human fecal bacterial enzymes, J. Natl. Cancer Inst., 64, 255, 1980. 41. Cordaro, J. T., Sellers, W. M., Bull, R. J., and Schmidt, J. P., Study of man during a 56-day exposure to an oxygen-helium atmosphere at 258 mmHg total pressure, Aerospace Med., 37, 594, 1966. 42. Winitz, M., Adams, R. F., Seedman, D. A., Davis, P. N., Jayko, L. G., and Hamilton, J. A., Studies in metabolic nutrition employing chemically defined diets. II. Effects on gut microflora populations, Am. J. Clin. Nutr., 23, 546, 1970. 43. Attebery, W. R., Sutter, V. L., and Finegold, S. M., Effect of a partially chemically defined diet on normal human fecal flora, Am. J. Clin. Nutr., 25, 1391, 1972. 44. Crowther, J. S., Drasar, B. S., Goddars, P., Hill, M. J., and Johnson, K., The effect of a chemically defined diet of the faecal flora and faecal steroid concentrations, Gut, 14, 790, 1973. 2387_ch4.6_fm Page 269 Sunday, May 6, 2001 7:27 PM INFLUENCES OF FIBER ON THE ECOLOGY OF THE INTESTINAL FLORA 269 45. Burnside, G. and Deurode, G. J., Effects of an elemental diet of human fecal flora, Gastroenterology, 66, 210, 1974. 46. Bounous, G. N. and Cohn, I., Stability of normal human fecal flora during a chemically defined, low residue liquid diet, Ann. Surg., 181, 58, 1975. 47. Salyer, A. A., Palmer, J. K., and Wilkins, T. D., Degradation of polysaccharides by intestinal bacterial enzymes, Am. J. Clin. Nutr., 31, S128, 1978. 48. Floch, M. J. and Fuchs, H. M., Modification of stool content by increased bran intake, Am. J. Clin. Nutr., 31, S185, 1978. 49. Robertson, J., Brydon, W. G., Tadesse, K., Wenham, P., Wallks, A., and Eastwood, M. A., The effect of raw carrot on serum lipids and colon function, Am. J. Clin. Nutr., 32, 1889, 1979. 50. Stephen, A. M. and Cummings, J. H., Mechanism of action of dietary fiber in the human colon, Nature (London), 284, 283, 1980. 51. Ehle, F. R., Robertson, J. B., and Van Soest, P. J., Influence of dietary fiber on fermentation in the human large intestine, J. Nutr., 112, 158, 1982. 52. Hirschberg, N. and Fantus, B., Mode of action of bran: III. Bacterial action on bran, Rev. Gastroenterology, 9, 370, 1942. 53. Drasar, B. S. and Jenkins, D. J. A., Bacteria, diet and large bowel cancer, Am. J. Clin. Nutr., 29, 1410, 1976. 54. Fuchs, H. M., Dorfman, S., and Floch, M. H., The effect of dietary fiber supplementation in man. II. Alteration in fecal physiology and bacterial flora, Am. J. Clin. Nutr., 29, 1443, 1976. 55. Drasar, B. S., Jenkins, D. J. A., and Cummings, J. H., The influence of a diet rich in wheat fibre on the human faecal flora, J. Med. Microbiol., 9, 423, 1976. 56. McLean-Baird, I., Walters, R. L., Davies, P. S., Hill, M. J., Drasar, B. S., and Southgate, D. A. T., The effect of two dietary fiber supplements on gastrointestinal transit, stool weight, and frequency, and bacterial flora, and fecal bile acids in normal subjects, Metabolism, 26, 117, 1977. 57. McLean Ross, A. W., Eastwood, M. A., Anderson, J. R., and Anderson, D. M. W., A study of the effects of dietary gum arabic in humans, Am. J. Clin. Nutr., 37, 368, 1983. 58. Doyle, R. B., Wolfman, M., Vargo, D., and Floch, M. H., Alteration in bacterial flora induced by dietary pectin, Am. J. Clin. Nutr., 34, 635, 1981. 59. Ross, J. K. and Leklem, J. E., The effect of dietary citrus pectin on the excretion of human fecal neutral and acid steroids and the activity of 7-α-dehydroxylase and β-glucuronidase, Am. J. Clin. Nutr., 34, 2068, 1981. 60. Fleming, S. E., Marthinsen, D., and Kuhnlein, H., Colonic function and fermentation in men consuming high fiber diets, J. Nutr., 113, 2535, 1983. 61. Daniel, M., Wisker, E., Rave, G., and Feldheim, W., Fermentation in human subjects of non-starch polysaccharides in mixed diets, but not in a barley fiber concentrate, could be predicted by in vitro fermentation using human fecal inocula, J. Nutr., 127, 1981, 1997. 62. Hidaka, H., Eida, T., Takizawa, T., Tokunaga, T., and Tashiro, Y., Effects of fructooligosaccharides on intestinal flora and human health, Bifid Microflora, 5, 37, 1986. 63. Hidaka, H., Tashiro, Y., and Toshiaki, E., Proliferation of bifidobacteria by oligosaccharides and their useful effect on human health, Bifid Microflora, 10, 65, 1991. 64. Williams, C., Witherly, S., and Buddington, R., Influence of dietary neosugar on selected bacterial groups of the human faecal microbiota, Microb. Eco. Health Dis., 7, 91, 1993. 65. Gibson, G. R., Beatty, E. R., Wang, S., and Cummings, J. H., Selective stimulation of Bifidobacteria in the human colon by oligofructose and inulin, Gastroenterology, 108, 975, 1995. 66. Bouhnik, Y., Flourie, B., Andriew, C., Bisetti, N., Briet, N., and Rambaud, J.-C., Effects of Bifidobacterium sp. fermented milk ingested with or without inulin on colonic bifidobacteria and enzymatic activities in healthy humans, Eur. J. Clin. Nutr., 50, 269, 1996. 67. Kleesen, B., Sykura, B., Sunft, H.-J., and Blaut, M., Effects of inulin and lactose on fecal microflora, microbial activity, and bowel habit in elderly constipated persons, Am. J. Clin. Nutr., 65, 1397, 1997. 68. Menne, E., Guggenbuhl, N., and Roberfroid, M., Fn-type chicory inulin hydrolysate has a prebiotic effect in humans, J. Nutr., 130, 1197, 2000. 69. Buddington, R. K., Williams, C. H., Chen, S.-C., and Witherly, S. A., Dietary supplement of neosugar alters the fecal flora and decreases activities of some reductive enzymes in human subjects, Am. J. Clin. Nutr., 63, 709, 1996. 2387_ch4.6_fm Page 270 Sunday, May 6, 2001 7:27 PM 2387_ch4.7_fm Page 271 Sunday, May 6, 2001 7:28 PM CHAPTER 4.7 Interaction between Human Gut Bacteria and Dietary Fiber Substrates Betty A. Lewis, Mary Beth Hall, and Peter J. Van Soest Dietary fiber plays an important physiological and nutritional role in human diets. It encompasses a wide range in its chemical composition and physical characteristics.1–3 Each source of dietary fiber has an intrinsic property that is based on its chemical composition and physical properties and which determines the biological and fermentive properties, i.e., the maximum limit for the rate and extent of fermentation for a specific substrate under optimal environmental conditions.4,5 The effects of different dietary fibers on human intestinal bacteria are further complicated since these dietary fibers also affect environmental conditions throughout the GI tract.5–8 Attempts to understand interactions between dietary fiber and intestinal bacteria also have to consider how responses of the GI tract will affect the fermentation.8 The presence of dietary fiber in the human diet could affect intestinal bacteria directly through catabolite regulation and indirectly through physical changes in the GI tract environment. The amount of fiber fermented, the amount of microbial organic matter produced, and the amount of water held by each fraction must be considered in order to predict the effect of dietary fiber on colonic contents.9 References 10–12 give more detailed information about the bacteria and the intestinal environment. Intrinsic properties of cereal brans have received considerable attention, but other sources of dietary fiber, which include vegetables, fruits, plant gums, bacterial and wood-derived polysacharides, and chemically and physically modified polysaccharides, offer a wide variety of properties.13–15 Mucopolysaccharides from sloughed epithelial cells and secreted mucin glycoproteins are carbohydrates supplied by the host that are available to intestinal bacteria.16,17 However, in vitro studies with Bacteroides species suggest that these host substances are not an important energy source for the bacteria.18,19 The Bacteroides, numerically a major bacterial genus in the colon, may utilize low levels of a variety of polysaccharides in vivo rather than a single source. Some of the predominant anaerobic species found in the human intestine which have been studied in pure culture with carbohydrate substrates are summarized in Table 4.7.1. It is apparent that the colonic bacterial species vary in their specificity for different polysaccharides.20 Thus, some Bacteriodes species can utilize different types of polysaccharides, while other species are more limited. In a study based on DNA homologies of fecal bacteria, the colon of each person evaluated had a bacterial population specific for that individual.21 The mechanism by which colonic bacteria digest a particular polysaccharide depends on the structure of the polysaccharide and the bacterial species.20 Several polysaccharide-degrading enzymes have been isolated from colonic Bacteroides species, revealing aspects of their mode of 0-8493-2387-8/01/$0.00+$1.50 © 2001 by CRC Press LLC 271 2387_ch4.7_fm Page 272 Sunday, May 6, 2001 7:28 PM 272 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Table 4.7.1 Some of the Predominant Anaerobic Bacterial Species Found in the Human Intestine and Studied in Pure Cultures with Selected Carbohydrates Substrate Degradability Cellulose Methyl and carboxymethyl cellulose Hemicellulose Pectin Cereal gums Guar gum, locust bean gum Arabinogalactans Maillard polymer Algal gum Mucopolysaccharide Mucin glycoprotein Partially fermentable Partially fermentable or unfermentable Partially fermentable Highly fermentable Highly fermentable Highly fermentable Partially fermentable Unfermentable Unfermentable Highly fermentable Partially fermentable a Bacterial Speciesa 1 N.K. 3,4,5,8,11,13 4,5,6,8,10,15 2,6,7,9 5,7,11,16 5,6,7,8,9,10,14 N.K. N.K. 4,5,6,10 Few Bacteroides strains: 12, 17 Adapted from Reference 20: 1, Bacteroides sp.; 2, B. distansnis; 3, B. eggerthii; 4, B. fragilis subspecies; 5, B. ovatus; 6, B. thetaiotaomicron; 7, B. uniformis; 8, B. vulgatus; 9, B. “T4-1”; 10, B. “3452A”; 11, Bifidobacterium adolescentis; 12, B. bifidum; 13, B. infantis; 14, B. longum; 15, Eubacterium eligens; 16, Ruminococcus albus; and 17, R. torques. N.K., not known. action. Thus, B. thetaiotaomicron degrade starch,22 laminaran,23 polygalacturonic acid,24 and chondroitin sulfate25 by induced enzymes that are cell-associated rather than extracellular. These glycanases, either soluble or membrane bound, appear to be located in the periplasmic space of the cell. They degrade the polysaccharides directly to mono- and oligosaccharides. Proteins bound to the outer cell surface of the bacteria may act as receptors to bind and facilitate transport of the polysaccharide into the cellular periplasmic space where the glycanases are located.26 Although the neutral polysaccharides are hydrolyzed to mono- and oligosaccharides, the acidic polysaccharides are broken down by lyase enzymes which cleave glycosidic bonds by a β-elimination reaction, giving rise to C4-C5-unsaturated uronic acid oligosaccharides. In both cases the oligosaccharides are further digested to the sugars by intracellular glycosidases. Not all polysaccharides are digested to mono- and oligosaccharides by the glycanases, however. The viscous guar galactomannan was digested only to high molecular weight fragments,27 which may be related to the distribution of branches or to the high viscosity. Aspects of the mechanisms by which colonic bacteria utilize polysaccharides have been reviewed.28 Person-to-person variation in the microbial fermentation of dietary fiber as shown in Table 4.7.2 was the most consistent result reported when mixed cultures of human fecal bacteria were incubated with various dietary fibers.29,30 Variation in the microbial fermentation of different sources of dietary fiber among people reflects responses by the bacteria and responses by the intestinal tract in a particular individual. This variation was dramatic with Solka floc as the source of dietary fiber,29–31 with fermentation ranging from 0 to 40%. Solka floc is a processed wood cellulose, which has been used as a source of dietary fiber for human and animal experimental diets and is commercially available for food-processing applications. The chemical and biological properties of Solka floc are different from native vegetable and forage celluloses.32 Mixed rumen bacteria incubated with Solka floc usually demonstrate an 18- to 24-h lag before significant fermentation is observed.4,5 Rumen bacteria and human fecal bacteria that ferment dietary fiber have many similarities in respect to nutritional requirements, volatile fatty acid production, and fermentation balances.33–36 However, human fecal, bovine ruminal, and equine cecal bacteria differed in their ability to ferment fiber from various sources (Table 4.7.3).4 Some humans also produce methane gas.36 The source of fiber supplement can have both short- and long-term effects on digestibility of other fiber sources. Ehle et al.30 reported that the source of dietary fiber (coarse white wheat bran, AACC-certified food grade R07-3691; cabbage; and Solka floc) in the diet was a significant factor 2387_ch4.7_fm Page 273 Sunday, May 6, 2001 7:28 PM INTERACTION BETWEEN HUMAN GUT BACTERIA AND DIETARY FIBER SUBSTRATES Table 4.7.2 Concentration (g/g) of Various Neutral Detergent Fibers Remaining after Incubation with Inoculum from Three Human Donors at Various Times in Batch Culture Substratea Hours 1 SD Human Donor 2 SD Cabbage Wheat bran Alfalfa Cabbage Wheat bran Alfalfa Cabbage Wheat bran Alfalfa Solka floc® 6 6 6 24 24 24 48 48 48 48 0.58b 0.69 0.85 0.10 0.56 0.61 0.05 0.56 0.59 1.00 0.03 0.02 0.01 0.01 0.04 0.03 0.02 0.02 0.04 0.01 0.56 0.86 0.83 0.20 0.53 0.65 0.07 0.53 0.60 0.99 a b 3 SD 0.39 0.72 0.89 0.17 0.53 0.59 0.14 0.53 0.54 0.77 0.05 0.02 0.02 0.02 0.04 0.05 0.05 0.01 0.03 0.02 Effect of Inoculum Source on the Fermentation of Neutral Detergent Fibers of Various Substrates Inoculum: Source: Substrate Alfalfa Timothy Wheat brana Cabbageb Wheat straw b 0.02 0.01 0.02 0.03 0.03 0.02 0.01 0.03 0.04 0.03 Ethanol-extracted cabbage, coarse white wheat bran (AACC-certified RO7 3691), Solka floc® (food grade cellulose), and alfalfa hay. n = 3, replicates. Table 4.7.3 a 273 Human Feces 38 1 53 91 0 Fermentation (%) Bovine ConcentrateHay-Fed Hay-Fed Fed Rumen Rumen Cecum 53 40 56 91 31 57 50 71 91 42 49 32 — — — Equine Hay- and Grain-Fed Cecum 50 27 — — — Coarse white wheat bran (AACC-certified food grade RO7-3691). Ethanol-extracted cabbage, see Ehle et al.30 Source: Jeraci, J. L., Interactions between Rumen or Human Fecal Inocula and Fiber Substrates, Master’s thesis, Cornell University, Ithaca, NY, 1981. With permission. influencing the in vitro fermentation of coarse white wheat bran, ethanol-extracted cabbage, Solka floc, and alfalfa hay when incubated with the respective human fecal bacteria. No long-term microbial adaptation to these fiber-supplemented diets was observed, since fermentation values obtained from 12 individuals who received a single fiber supplement for an extended period of time (approximately 70 days) did not differ from fermentation values obtained in another 12 individuals who received a different fiber supplement every 2 weeks. In the same study, the apparent in vivo fermentation of neutral detergent fiber (NDF) was significantly affected by the source of dietary fiber in the fiber-supplemented diets. Estimates of the apparent in vivo fermentation were confounded by the quantity of microbial organic matter and the presence or lack (for cabbage) of unfermented residue from the diet in the feces. However, the true calculated in vivo fermentation of NDF was affected by the source of dietary fiber. Regardless of the dietary fiber source, ingestion of Solka floc in any 2-week period resulted in the carryover effect of depressed fermentation of NDF in all future periods. The observation that previous exposure to Solka floc depressed the fermentation of other dietary fiber substrates was confirmed with in vitro techniques.8,37 Fermentations with human fecal bacteria showed a 36-h depression of pectin fermentation in batch culture only when the bacterial inoculum came from a continuous culture that received Solka floc as the substrate. A selective depression of 2387_ch4.7_fm Page 274 Sunday, May 6, 2001 7:28 PM 274 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Solka floc fermentation in the batch culture flasks was not observed when bacteria received Solka floc or pectin as the continuous culture substrate. This would suggest that certain modified or highly crystalline sources of cellulose may generate unexpected artifacts in fiber digestion studies. Although colonic fermentation of the hydrated and less lignified cellulose of plant foods has been well established for humans,32,38 fermentation of the more crystalline isolated celluloses (Solka floc, filter paper) by fecal bacteria has been demonstrated for only 20 to 30% of the subjects studied.30–32,39 Resistance of polysaccharides to fermentation has also been related to binding to other components in the plant cell wall and to branching patterns of polysaccharides, in addition to intraand intermolecular H-bonding with consequent lack of hydration and solubility. Significant amounts of starch (20%) may survive digestion and absorption in the small intestine.40 The proportion of dietary starch passing the ileum is related to the plant source, foodprocessing effects, diet portion, and individual variation. This resistant or nondigested starch, like other nondigestible polysaccharides, is fermented to short-chain fatty acids (SCFA) and the gases hydrogen, methane, and carbon dioxide. Recent studies have shown differences among human subjects in the rate at which the fecal bacteria ferment the starch and in the relative proportions of the individual SCFA formed.41,42 Weaver et al.43 observed significantly higher proportions of acetate and lower proportions of butyrate to total SCFA in subjects with polyps or colon cancer. Butyrate, the predominant energy source for colonocytes, may play a regulatory role in the pathogenesis of colon cancer. In vitro systems have been evaluated as models for fermentation of fiber in the large intestine.8,44 Human feces afford an adequate and representative inoculum for in vitro systems and have been used in many studies of colonic fermentation.9,45,46 In an in vitro system, SCFA production from fermentation of ileal effluents was significantly correlated with SCFA production from dietary fiber isolates but not with the SCFA production from whole foods.47 REFERENCES 1. Eastwood, M. A., The physiological effect of dietary fiber: an update, Ann. Rev. Nutr., 12, 19, 1992. 2. Selvendran, R. R. and MacDougall, A. J., Cell-wall chemistry and architecture in relation to sources of dietary fibre, Eur. J. Clin. Nutr., 49, S3, S27, 1995. 3. Van Soest, P. J. and Robertson, J. B., Chemical and physical properties of dietary fibre, in Dietary Fibre, Proc. Miles Symp. 1976, Halfax, Nova Scotia, Nutr. Soc. Canada, Hawkins, W. W., Ed., Miles Laboratories, Ltd., Rexdale, Ontario, 13. 4. Jeraci, J. L., Interactions between Rumen or Human Fecal Inocula and Fiber Substrates, Master’s thesis, Cornell University, Ithaca, NY, 1981. 5. Van Soest, P. J., Nutritional Ecology of the Ruminant, 2nd ed., Cornell University Press, Ithaca, NY, 1994. 6. Stevens, C. E., Physiological implications of microbial digestion in the large intestine of mammals: relation to dietary factors, Am. J. Clin. Nutr., 31 (suppl.), S161, 1978. 7. Spiller, G. A., Chernoff, M. C., Hill, R. A., Gates, J. E., Nassar, J. J., and Shipley, E. A., Effect of purified cellulose, pectin, and a low-residue diet on fecal volatile fatty acids, transit-time, and fecal weight in humans, Am. J. Clin. Nutr., 33, 754, 1980. 8. Jeraci, J. L., Use of the High Fiber Chemostat System to Study Interactions among Gut Microflora, Ph.D. thesis, Cornell University, Ithaca, NY, 1984. 9. McBurney, M. I., Horvath, P. J., Jeraci, J. L., and Van Soest, P. J., Effect of in vitro fermentation using human faecal inoculum on the water-holding capacity of dietary fibre, Br. J. Nutr., 53, 17, 1985. 10. Roth, H. P. and Mehlman, M. A., Eds., Symposium on role of dietary fiber in health, Am. J. Clin. Nutr., 31 (suppl.), 1978. 11. Hentges, D. J., Ed., Human Intestinal Microflora in Health and Disease, Academic Press, New York, 1983. 12. Cummings, J. H. and Macfarlane, G. T., The control and consequences of bacterial fermentation in the human colon, J. Appl. Bacteriol., 70, 443, 1991. 2387_ch4.7_fm Page 275 Sunday, May 6, 2001 7:28 PM INTERACTION BETWEEN HUMAN GUT BACTERIA AND DIETARY FIBER SUBSTRATES 275 13. Cummings, J. H., Hill, M. J., Jenkins, P. J. A., Pearson, J. R., and Wiggins, H. S., Changes in fecal composition and colonic function due to cereal fiber, Am. J. Clin. Nutr., 29, 1468, 1976. 14. Van Soest, P. J., Horvath, P. J., McBurney, M. I., Jeraci, J. L., and Allen, M., Some in vitro and in vivo properties of dietary fibers from noncereal sources, in Unconventional Sources of Dietary Fiber, Series 214, Furda, I., Ed., American Chemical Society, Washington, D.C., 1983. 15. Horvath, P. J., The Measurement of Dietary Fiber and the Effects of Fermentation, Ph.D. thesis, Cornell University, Ithaca, NY, 1984. 16. Savage, D. C., Factors involved in colonization of the gut epithelial surface, in Symp. on Role of Dietary Fiber in Health, Roth, H. P. and Mehlman, M. A., Eds., Am. J. Clin. Nutr., 31 (suppl.), S131, 1978. 17. Salyers, A. A., O’Brien, M., and Schmetter, B., Catabolism of mucopolysaccharides, plant gums, and Maillard products by human colonic Bacteroides, in Unconventional Sources of Dietary Fiber, Series 214, Furda, I., Ed., American Chemical Society, Washington, D.C., 1983. 18. Salyers, A. A. and McCarthy, R. E., Assessing the importance of host-derived polysaccharides as carbon sources for bacteria growing in the human colon, Anim. Feed Sci. Tech., 23, 109, 1989. 19. Salyers, A. A., Activities of polysaccharide-degrading bacteria in the human colon, in Dietary Fiber: Chemistry, Physiology, and Health Effects, Kritchevsky, D., Bonfield, C., and Anderson, J. W., Eds., Plenum Press, New York, 1990, 187. 20. Salyers, A. A., Energy sources of major intestinal fermentative anaerobes, in 5th Int. Symp. on Intestinal Microecology, Luckey, T. D., Ed., Am. J. Clin. Nutr., 32, 158, 1979. 21. Johnson, J. L., Specific strains of Bacteroides species in human fecal flora as measured by deoxyribonucleic acid homology, Appl. Environ. Microbiol., 39, 407, 1980. 22. Smith, K. and Salyers, A. A., Purification and characterization of enzymes involved in starch utilization by Bacteroides thetaiotaomicron, J. Bacteriol., 173, 2962, 1991. 23. Salyers, A. A., Palmer, J. K., and Wilkins, T. D., Laminarinase (β-glucanase) activity in Bacteroides from the human colon, Appl. Environ. Microbiol., 33, 1118, 1977. 24. McCarthy, R. E., Kotarski, S. F., and Salyers, A. A., Location and characteristics of enzymes involved in the breakdown of polygalacturonic acid by Bacteroides thetaiotaomicron, J. Bacteriol., 161, 493, 1985. 25. Salyers, A. A. and O’Brien, M., Cellular location of enzymes involved in chondroitin sulfate breakdown by Bacteroides thetaiotaomicron, J. Bacteriol., 143, 772, 1980. 26. Cheng, Q., Yu, M. C., Reeves, A. R., and Salyers, A. A., Identification and characterization of a Bacteriodes gene, csuF, which encodes an outer membrane protein that is essential for growth on chondroitin sulfate, J. Bacteriol., 177, 3721, 1995. 27. Gherardini, F. C. and Salyers, A. A., Characterization of an outer membrane mannanase from Bacteriodes ovatus, J. Bacteriol., 169, 2031, 1987. 28. Salyers, A. A., Polysaccharide utilization by human colonic bacteria, in New Developments in Dietary Fiber, Furda, I. and Brine, C. J., Eds., Plenum Press, New York, 1990, 151. 29. Jeraci, J. L., Robertson, J. B., and Van Soest, P. J., A human fecal inoculum in the in vitro fermentation procedure, J. Anim. Sci. Suppl., 51, 206, 1980. 30. Ehle, F. R., Robertson, J. B., and Van Soest, P. J., Influence of dietary fibers on fermentation in the human large intestine, J. Nutr., 112, 158, 1982. 31. Betian, H. G., Linehan, B. A., Bryant, M. P., and Holdeman, L. V., Isolation of a cellulolytic Bacteroides sp. from human feces, Appl. Environ. Microbiol., 33, 1009, 1977. 32. Van Soest, P. J., Jeraci, J. L., Foose, T., Wrick, K., and Ehle, F. R., Comparative fermentation of fibre in man and other animals, in Proc. Dietary Fibre in Human and Animal Nutrition Symp., Wallace, G. and Bell, L., Eds., The Royal Society of New Zealand, Palmerston North, N.Z., 1982. 33. Bryant, M. P., Nutritional features and ecology of predominant anaerobic bacteria of the intestinal tract, Am. J. Clin. Nutr., 27, 1313, 1974. 34. Miller, T. L. and Wolin, M. J., Fermentations by saccharolytic intestinal bacteria, in 5th Int. Symp. on Intestinal Microecology, Luckey, T. D., Ed., Am. J. Clin. Nutr., 32, 164, 1979. 35. Smith, C. J. and Bryant, M. P., Introduction to metabolic activities of intestinal bacteria, in 5th Int. Symp. on Intestinal Microecology, Luckey, T. D., Ed., Am. J. Clin. Nutr., 32, 149, 1979. 36. Wolin, M. J. and Miller, T. L., Carbohydrate fermentation, in Human Intestinal Microflora in Health and Disease, Hentges, D. J., Ed., Academic Press, New York, 1983, 147. 2387_ch4.7_fm Page 276 Sunday, May 6, 2001 7:28 PM 276 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION 37. Jeraci, J. L. and Horvath, P. J., In vitro fermentation of dietary fiber by human fecal organisms, Anim. Feed Sci. Tech., 23, 121, 1989. 38. Cummings, J. H., Microbial digestion of complex carbohydrates in man, Proc. Nutr. Soc., 43, 35, 1984. 39. Wedekind, K. J., Mansfield, H. R., and Montgomery, L., Enumeration and isolation of cellulolytic and hemicellulolytic bacteria from human feces, Appl. Environ. Microbiol., 54, 1530, 1988. 40. Stephen, A. M., Haddad, A. C., and Phillips, S. F., Passage of carbohydrates into the colon, Gastroenterology, 85, 589, 1983. 41. Weaver, G. A., Krause, J. A., Miller, T. L., and Wolin, M. J., Constancy of glucose and starch fermentations by two different human faecal microbial communities, Gut, 30, 19, 1989. 42. Scheppach, W., Fabian, C., Sachs, M., and Kasper, H., The effect of starch malabsorption on fecal short-chain fatty acid excretion in man, Scand. J. Gastroenterol., 23, 755, 1988. 43. Weaver, G. A., Krause, J. A., Miller, T. L., and Wolin, M. J., Short chain fatty acid distributions of enema samples from a sigmoidoscopy population: an association of high acetate and low butyrate ratios with adenomatous polyps and colon cancer, Gut, 29, 1539, 1988. 44. Macfarlane, S., Quigley, M. E., Hopkins, M. J., Newton, D. F., and Macfarlane, G. T., Polysaccharide degradation by human intestinal bacteria during growth under multi-substrate limiting conditions in a three-stage continuous culture system, FEMS Microbiol. Ecology, 26, 231, 1998. 45. McBurney, M. I. and Thompson, L. U., Effect of human faecal inoculum on in vitro fermentation variables, Br. J. Nutr., 58, 233, 1987. 46. McBurney, M. I. and Thompson, L. U., Dietary fiber and energy balance: integration of the human ileostomy and in vitro fermentation models, Anim. Feed Sci. Tech., 23, 261, 1989. 47. McBurney, M. I., Thompson, L. U., Cuff, D. J., and Jenkins, D. J. A., Comparison of ileal effluents, dietary fibers, and whole foods in predicting the physiological importance of colonic fermentation, Am. J. Gastroenterol., 83, 536, 1988. 2387_ch4.8_fm Page 277 Sunday, May 6, 2001 7:30 PM CHAPTER 4.8 Effects of Dietary Fiber on Digestive Enzymes Barbara O. Schneeman and Daniel Gallaher Dietary fibers affect the functioning of the GI tract as indicated by a lower digestibility and availability of nutrients from high-fiber diets. Assimilation of nutrients from diets requires the movement of the bolus of food through the gut, the enzymatic hydrolysis of complex nutrients to simpler compounds, and absorption of these compounds into and through the small intestinal cells. The presence of dietary fibers can alter these processes, resulting in a slower rate of nutrient absorption and a shift in the site of absorption to the more distal areas of the small intestine. In this chapter, the effects of dietary fibers on digestive enzyme activity and on the intestinal contents are reviewed. Table 4.8.1 presents the effects of various fiber sources on amylase, lipase, trypsin, chymotrypsin, or pepsin activity in vitro. The enzymes were derived from human samples or from commercial enzyme preparations. In general, commercial lipase or amylase is of porcine origin, and -trypsin and chymotrypsin are of bovine origin. Discrepancies are most likely due to the enzyme source or to the method of incubating the fiber and enzyme and reporting values. As shown in Table 4.8.2, lipase inhibitory activity is associated with several cereals. Addition of fiber sources to an in vitro protein digestibility test can lead to reductions in the percentage of digestible casein (Table 4.8.3), indicating that fibers can interfere with proteolytic enzyme activity. Other in vitro data indicate that certain foods which contain fiber may slow starch or carbohydrate hydrolysis (Table 4.8.4). For both carbohydrates and protein, the change in in vitro digestibility was dependent on the source of fiber (Tables 4.8.3 and 4.8.4). The data in Table 4.8.4 indicate that the physical state of the plant cell wall rather than simply the presence of dietary fiber may be important in slowing the rate of carbohydrate hydrolysis. Grinding brown rice or lentil samples significantly increased the percentage of starch hydrolyzed in a 30-minute period.11 These results indicate that the physical state of the food or of the cell wall layers can slow the penetration of digestive enzymes.11,13 Tinker and Schneeman14 demonstrated in vivo that the consumption of the viscous polysaccharide, guar gum, slows the disappearance of starch from the small intestine of rats. An interference with starch hydrolysis by amylase may contribute to this effect. These studies suggest that slowing enzymatic hydrolysis in the intestine could contribute to the apparent slower rate of nutrient assimilation associated with certain fiber-rich foods or diets. The activity of digestive enzymes in vivo has been estimated (Table 4.8.5). Both rat studies, collecting total intestinal contents, and human studies based on a sample of duodenal aspirate indicate that within the intestinal contents pancreatic enzyme activities are either similar or significantly higher when a fiber supplement is added to a basal diet. In pancreatic duct cannulated rats, instillation of a pectin test diet reduced amylase and chymotrypsin secretion; however, infusion of 0-8493-2387-8/01/$0.00+$1.50 © 2001 by CRC Press LLC 277 2387_ch4.8_fm Page 278 Sunday, May 6, 2001 7:30 PM 278 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Table 4.8.1 Percent of Control Enzyme Activity In Vitro Fiber Source Amylase Alfalfa Alfalfa Oat bran Wheat bran Wheat bran Wheat bran Rice bran Safflower meal Xylan Xylan Cellulose Cellulose Pectin Pectin Pectin-HMa Pectin-LMb Pectin Guar gum Guar gum Guar gum Carob bean Gum Na-alginate Agar-agar Carrageenan Carrageenan Carrageenan Carrageenan Psyllium Lignin Lipase Trypsin Chymotrypsin 48.6 72.8 83.83 63.0* 29.0* 94.9 93.9 51.6* 71.3 82.1 85.9 70.0 44.6* 31.0* 20.8* 4.6* 9.9* 113.0 123.0 20.0 5.0 96.1 93.8 92.1 56.3* 11.2* 91.8 55.3* 87.5 105.0 100.0 40.0 12.0 90.8 76.2 90.1 90.1 20.0* 98.5 52.9* 78.3* 105.0 129.0* 20.0 25.0 87.3 72.6 8.0 66.9* 32.7* 20.4* 148.0* 50.0 0.0 Pepsin 98.0 54.0 60.0 45.0 90.0 74.0 121.0 95.0 89.0 101.0 74.0 Inhibits Inhibits 59.0 59.0 100.0 Ref. 1c 2d 2 1 1 2 1 1 2 1 2 1 1 2 4+e 4+ 5+c 3+e 4+ 6+c 6+ 6+ 6+ 5+ 7+c 8+c 6+ 3+ 3+ Note: An asterisk means that there was a significant change from control. A plus sign in the reference column indicates that a statistical analysis was not reported in these studies. a High methoxy. Low methoxy. c Commercial enzyme source was used. d Human pancreatic juice was used. e Human duodenal juice was used. b Table 4.8.2 Pancreatic Lipase Inhibition by Cereals, In Vitro Cereal Source French soft wheat French durum wheat Scandinavian soft wheat North American hard wheat Barley (8 varieties) White sorghum (5 varieties) Red sorghum (4 varieties) Millet (6 varieties) a Lipase Inhibitory Activitya (IU/g) 25.6 33.6 43.3 37.9 23.5 22.4 3.3 37.3 ± ± ± ± ± ± ± ± 1.1 1.0 4.1 1.1 2.0 4.8 1.5 3.3 Defined by Borel et al. (1989) as the mg of material that decrease lipase activity by 50% under their experimental conditions. Source: Adapted from Cara et al.9 2387_ch4.8_fm Page 279 Sunday, May 6, 2001 7:30 PM EFFECTS OF DIETARY FIBER ON DIGESTIVE ENZYMES Table 4.8.3 Decrease in In Vitro Casein Digestibilitya Due to Fiber Addition Fiber Source Holocellulose Lignin Citrus pectin Xylan Karaya gum Wheat bran Brown rice Cooked broccoli Cooked blackeyed peas Canned corn a 279 Decrease Digestibility of Casein Fiber: Protein (wt:wt) 1.66 1.36 4.03 5.34 4.42 0.15 7.64 8.93 9.35 14.86 0.4 0.4 0.4 0.4 0.4 0.5 0.5 0.5 0.5 0.5 Enzyme mixture was trypsin, chymotrypsin, and peptidase followed by a bacterial protease. Source: Adapted from Gagne, C. M. and Acton, J. C., J. Food Sci., 48, 734, 1983. With permission. Table 4.8.4 In Vitro Carbohydrate Hydrolysis of Foods Product Wheat Rolled Cooked, rolled White bread Whole meal bread Whole meal bread Rice Cooked, brown Cooked, ground brown Cooked, white Cooked, ground white Rye, rolled Barley, rolled Oats, rolled cooked Maize, cooked rneal Brown lentils Whole Ground Lentils, ground cooked Soybeans, ground cooked a b % Starch Hydrolyzed in 30 min % Carbohydrate Released in 3 h Ref. 27 11a 11a 11a 11a 12b 5.1 45.8 77.6 80.1 17.6 68.2 11a 11a 30.8 71.8 11a 11a 11.9 13.5 68 71.5 11a 11a 11a 11a 12.1 60.9 15 13a 13a 12b 6 12b Enzyme source is commercial α-amylase and amyloglucosidase. Enzyme source is human saliva and jejunal juice. a pectin alone (i.e., not in a test diet) did not change basal pancreatic protein secretion from basal values.17 In contrast, two viscous polysaccharides, konjac mannan and Na-alginate, have been reported to increase pancreatic output,18 although this response could be due to long-term adaptation to continual feeding and not a change in the acute response. The data in Table 4.8.5 illustrate an interesting point: although various fibers may interfere with digestive enzyme activity based on in vitro evidence, they do not reduce the total amount of measurable digestive enzyme activity in the gut contents. The only reported significant decrease in enzyme activity occurred in pancreatectomy 2387_ch4.8_fm Page 280 Sunday, May 6, 2001 7:30 PM 280 Table 4.8.5 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION In Vivo Pancreatic Enzyme Activity: Values Expressed as Percentage of Fiber-Free Treated Controls Fiber Source Amylase Lipase Trypsin Chymotrypsin Protease Species Ref. Rat 18 Rat 15 Rat 19 Human 16 Human 20 Rat 16 Rat 18 Rat 17 Total Units in Intestinal Contents (% Control) Apple pectin Carrageenan Na-alginate Locust gum Xanthan gum Guar gum Cellulose Wheat bran Pectin Guar gum 119 121 111 104 109 115 142 71 152* 141* Pectin Wheat bran 883 488 Total Units in Ileostomy Fluid (% Control) 359* 151 187 180* Pectin Carob flour Pectina Wheat branb 175* 179* 53* 48* Units per ml of Intestinal Fluid (% Control) 144 154 130 218* 228 163* 37* 37* 45 65 Pectin Carob flour Carrageenan Guar gum Pectin Carrageenan Na-alginate Locust gum Xanthan gum Guar gum Cellulose Pectin 193* 203* 208* 167* 303* 321* 89.5 172* 216* 218* 142* 140* 136* 134* 142* 161* 114 101 240* 69.6 80.4 138* Rate of Pancreatic Secretion (% Control) 47.8* 61.5 91.4 108 56.8 51.4 67.2 114 122 125 115 132 155* 109 153* 160 173* 155 232* 186* No change in pancreatic protein secretion from baseline 159* 44.9* 63.6 51.3 102 185 217* 170 208* 205* 250* Note: * The value was reported as different from the fiber-free treated control. a b Pancreatectomized patients who received a pancreatic enzyme supplement. Chronic pancreatitis patients who did not receive an enzyme supplement. or pancreatitis patients, who cannot respond effectively by increasing secretion.20 Enzyme activity from intestinal contents is typically measured under in vitro conditions that optimize activity and thus may not reflect the physiologically available activity within the gut contents. To answer the question of whether fiber sources cm slow the rate of substrate hydrolysis, in vivo studies have been conducted that estimate the rate or extent of lipid and carbohydrate disappearance from the gastrointestinal tract. Table 4.8.6 summarizes several studies conducted in humans and rats to determine if fiber supplements will delay the absorption of lipid. The physiological importance of in vitro lipase inhibition by wheat bran and wheat germ has been studied extensively.9,25 Borel et al.25 reported that feeding wheat bran or wheat germ decreases gastrointestinal lipolysis of fats, resulting in lower intestinal absorption of cholesterol and fatty acids. Experimental evidence suggests that in vitro inhibition of lipase by cereal fractions may be due to a protein fraction.9 However, in vivo reductions In altering digestion and absorption in the small intestine. another effect of dietary fiber could be to alter the composition of the intestinal contents. 20 21 20 22 23 33 25 19 25 33 24 33 19 24 24 33 of lipid digestion may be due to interference with micelle formation as well as direct inhibition of lipolytic activity. This apparent discrepancy is most likely due to factors such as feeding very high levels of fiber. Table 4. not fasting the animals prior to killing them for collection of intestinal mucosa. One study reported that thymidine kinase activity was higher than the fiberfree control in rats fed alfalfa.1 presents the viscosity of gastric and intestinal .8_fm Page 281 Sunday. only cholesterol appearance was lower at 24 h Intestinal disappearance of cholesterol and triolein was 20–25% lower than cellulose at 2 h after meal Lymphatic appearance of triolein and cholesterol was lower than fiber-free adapted rats at 4 h after lipid infusion. In Table 4.8.8. appearance of cholesterol was lower at 4 h but not at 24 h after lipid infusion Delayed the disappearance of triglyceride from the gastrointestinal tract 1. May 6. or differences in the site of tissue sampling.8. The results summarized in Table 4. guar gum. and two studies reported decreases with pectin treatment. we have summarized a variety of reasons that will lead to different observations when examining digestive enzyme adaptation and activity. 2001 7:30 PM EFFECTS OF DIETARY FIBER ON DIGESTIVE ENZYMES Table 4. and Figure 4.2387_ch4. use of weanling animals that may be unable to adapt to high fiber intakes. Two of the studies tend to report increases in enzyme activity due to the fiber supplement where a change in activity was observed. or psyllium.8. but not in those fed cellulose or pectin.9 gives the effect of several fibers on viscosity of human duodenal juice.8.6 281 Effect of Fiber Sources on Lipid Digestion and Absorption Fiber Source Dose Pectin 5g Pectin Wheat bran 15 g/day 20 g Wheat bran 16 g/day Cellulose 20% Cellulose 10% Wheat bran 10% Wheat bran 20% Wheat germ 10% Guar gum 5% Guar gum 5% Pectin 5% Pectin 5% Konjac mannan 5% Chitosan 5% Psyllium husk 5% Results Human Studies Triolein breath test indicated a 30% reduction in lipid digestibility in pancreatectomized patients receiving enzyme replacement Fat excretion in ileostomy fluid increased by 36% Triolein breath test indicated up to a 30% reduction in lipid digestibility in pancreatitis patients No change in fat excretion in ileostomy fluid Animal Studies Intestinal disappearance of triolein reduced by 20–30% for 5-h period after meal compared to fiber-free group Lymphatic appearance of triolein was not lower than fiber-free adapted rats.7.6-fold nonsignificant increase in the excretion of fat in ileostomy fluid Delayed the disappearance of triglyceride from the gastrointestinal tract Lymphatic appearance of triolein and cholesterol was lower than fiber-free adapted rats at 4 h after lipid infusion.6 indicate that sources of viscous polysaccharides reduce the overall rate of lipid digestion.8. The change in the activity of small intestinal brush order enzymes with fiber treatments in animals is shown in Table 4. only cholesterol appearance was lower at 24 h Threefold significant increase in the excretion of fat in ileostomy fluid Intestinal disappearance of cholesterol and triolein was 40% lower than cellulose at 2 h after meal Intestinal disappearance of cholesterol and triolein did not differ from cellulose at 2 h after meal Lymphatic appearance of triolein and cholesterol was lower than fiber-free treatment at 4 and 24 h after lipid infusion Ref.8. milligrams of tissue. Table 4. it has been reported that addition of fiber to a basal diet will increase the volume and weight of intestinal content in rats.5 g%) HM-pectin (1. 2001 7:30 PM 282 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.34 Pasquier et al.8 Reasons for Differences in Enzyme Values Use of weanling vs. NC = no change. a b Proximal intestine. In addition. – = not determined. contents of rats fed different fiber sources.8. G..8. Gastroenterology. or DNA Table 4.8_fm Page 282 Sunday.b 26 27 28 29 30 26 27 28 30 26 31 30 30 30 Note: ↑. May 6. Distal intestine.7 Small Intestinal Enzyme Activity: Change in Enzyme Activity Fiber Source Peptidase Sucrase Lactase Maltase Thymidine Kinase Ref.31.36 have demonstrated in vitro that viscous polysaccharides increase the size of lipid droplets under conditions that mimic the stomach or small intestine.32 Sandberg et al.8. The lipase enzymatic systems act at this interface to release free fatty acids. 918. a threefold increase in droplet size was associated . 15 g%) Wheat bran (1.5 g%) Guar gum (0. 3RD EDITION Table 4. The amount of surface area available is determined by the size of lipid droplets and/or micelles. The greater volume or viscosity of intestinal contents will have an impact on the interaction of substrates and enzymes as well as on the movement of nutrients to sites for absorption. 82. 1982. adult animals Level and type of fiber added Protein content of the diet Fasting or fed state at time of killing Methodology differences relative to sample preparation and analysis Method of expressing units relative to volume of sample. protein. Hydrolysis of triglycerides in the stomach and small intestine is related to the surface area available.15. ↓ = direction of change from control.21. and monoglycerides.22 reported that the wet weight of ileostomy fluid was increased by 94 g/d due to wheat bran and by 314 g/d due to pectin.35. et al.2387_ch4.5 g%) 30 40 200 500 90 Source: Adapted from Isaksson..b ↑a. Under conditions that mimic gastric contents. Pectin Pectin Pectin Pectin Pectin Cellulose Cellulose Cellulose Cellulose Oat bran Wheat bran Alfalfa Guar gum Psyllium ↑ – – ↓a – ↑ – – – NC NC – – – NC ↓ ↑ – NC NC NC NC NC NC NC – ↑a ↑a – ↓ ↑ – – – NC NC – – – – – – – NC ↑ – – – NC NC – – NC – – – – – – – NC – – – NC – – ↑b ↑a. diglycerides.9 Effect of Dietary Fiber on Viscosity of Duodenal Juice (mPa) Duodenal juice LM-pectin (2. 34 REFERENCES 1. W. 634. C.. Schneeman.18) with reduced surface area and about a 30% reduction in the extent of triglyceride hydrolysis. and Schulz. Am. The effect of dietary fiber on human pancreatic enzyme activity in vitro.. 1960. . Effect of plant fiber on lipase. Dunaif. Effect of dietary fiber on pancreatic enzyme activity in vitro. Clin. J.8_fm Page 283 Tuesday.. 82. and Schneeman.. 43. The inhibition of pepsin and peptic ulcers. J. Food Sci. triglyceride hydrolysis was inversely correlated with viscosity and an increase in viscosity resulted in larger lipid droplets.. 2001 3:00 PM EFFECTS OF DIETARY FIBER ON DIGESTIVE ENZYMES 283 Figure 4.. 29.. 1982. 34. Gastroenterology. Bhayana. B. Lundquist. 4. 5. *Value differs significantly from the fiber-free control group (p < 0. and chymotrypsin activity. I. B. (Data adapted from Ikegami et al. E.. G..8. 918. T. Likewise. O. J. The effect of dietary fiber on pancreatic amylase activity in vitro. trypsin. 3. 196. I. J. Hansen. 10349. 2.. 1982. with conditions that mimic the duodenum in vitro..1 Gastric and intestinal viscosity in rats fed different fiber sources. 1978. 1981. O. and Ihse. Gastroenterology.. Hepato-Gastroenterology. Isaksson. 1990. Nutr. May 8. G.05). 39. The effects of viscous polysaccharides on in vitro lipid emulsification and hydrolysis have been confirmed in vivo in rat studies. 157. and Lee.2387_ch4. G. Houck. Br. Armaud. 171. Fiber constituents and fibrous food residues effects on the in vitro enzymatic digestion of protein. 1985. I. . N. B. Scand.2387_ch4. L. 14.... 112. 14. Milling and processing of wheat and other cereals affect their capacity to inhibit lipase in vitro. Lundquist....-S.. S. 10.. Pharm. Wong. J. Borel.. Int. C. H.. 13.. and Holtzapple. Digestion. E. 20.. 1979. Hasselblad. Ahderinne. Isaksson. 1982. J. and Lafont.. Tinker. Jenkins. O. L. Nutr.. and O’Dea. Tsuchihashi.-G. 281. Am. 1980. and Losowsky. 23. A.. in press. J. 1983. 1981. The Royal Society of New Zealand. The effects of citrus pectin on the absorption of nutrients in the small intestine. 1972. Effects of pectin and wheat bran on intraluminal pancreatic enzyme activities and on fat absorption as examined with the triolein breath test in patients with pancreatic insufficiency. 1979. C. G. K. and Tasmen-Jones. 45. Gagne.. 25. 29. A.. 1990. Effect of dietary cellulose on site of lipid absorption. L. B. Andersson. P. Isaksson.. Wheat bran and wheat germ: effect on digestion and intestinal absorption of dietary lipids in the rat. H. J. B. in Fiber in Human and Animal Nutrition.. Akesson. H. and Kasper. 353... 28. D. Senft.. G. Kelleher.. C. H. O. Dietary fiber decreases cholesterol and phospholipid synthesis in rat intestine.. Clin. J. and Gallaher. 29.. 120.. 1980. 477.. and Hultén. 27.. C.. 34.. 467. O.. Med. Harmuth-Hoene. FEBS Lett. 1983. Nutr. 66. Effects of dietary fiber on pancreatic enzyme activities of ileostomy evacuates and on excretion of fat and nitrogen in the rat. 11. 1100. 1981. Bulletin 20. S. L. Nutr. 403. Nutr.. G.. Food Sci. 119. Anderson.. B. 1983.. Schneeman. 139. cholesterol. HepatoGastroenterology. K. phospholipids. and O’Dea. Hasselblad. G184. J. Food Sci. R. cellulose and pectin.. 357. Am. and Stute. K... May 6. 1979. 715. 1192. 17. 2001 7:30 PM 284 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. J. Hallgren. R. J. Res. 1983.. Effect of indigestible polysaccharides on protein digestibility and nitrogen retention in growing rats. 18. J. 1989. Clin.. pectin and oat bran on the small intestine in the rat. S. 49. 23. The effect of dietary fiber on the pancreatic excretory function. J. 12. J. K. Br. Lafont.. Baille. Nutr. A. Br. P.. 1315.. 37.. 20.. P. D. Enzymatic reactions in the presence of polymers. 9. Lesgards.. 19. 1984. 45. 26.-S. C. The effect of guar gum or wheat bran on the disappearance of 14C-labelled starch from the rat gastrointestinal tract.. Starr. 42. 22. P. 1968. Sandberg. D.. B. Rep. and Ihse. The competitive inhibition of trypsin by A-carrageenan. Disaccharide levels of the rat jejunum are altered by dietary fibre. J. A.. 119. J.. O.. R. J. K. and Lairon.. R... M. Lipid.. 734. Backman.. Pancreatic and intestinal enzyme activity in rats fed various fiber sources.. F. 399. 746. Importance of physical form rather than viscosity in determining the rate of starch hydrolysis in legumes. and Schwerdtfeger.. 1982. H. Farness. J. I. O. Chantan. Snow. 16. and Ihse. A. D. 18. 19. 48. and Schneeman. 15. W.. E. 7. Asp. 252. B. Effects of viscous indigestible polysaccharides on pancreatic biliary secretion and digestive organs in rats. Effects of dietary cellulose. L. B. Nutr. and Schneeman. Clin. Isaksson. Nutr. Schwartz. Andersson. Gastroenterol.. Rate of digestion of foods and postprandial glycaemia in normal and diabetic subjects. and Schneeman. J. L.. The effect of pectin on the structure and function of the rat small intestine. 24. Pharmacol. 3RD EDITION 6... M. Nutr. L. 1989.. 2721. S. 20. 28. Gatfield. Wallace. and Innami.. D. E. Sommer. 249. and Schneeman. J. Am. 23. J. Physiol. Ebihara. Nutr.. and Harthill. Forman. J. 417. M..8_fm Page 284 Sunday. Thomsen... Acute pancreatic and biliary response to protein.. Am. Human Nutrition: Clinical Nutrition. Nutr. H. G. J. Nutr. and Acton.. Metab. J. L. H.. J. P. N. M. 1983. Lairon. Experimental model for in vivo determination of dietary fibre and its effects on the absorption of nutrients in the small intestine.. Hallgren. G. Schneeman. I. Eds. and Bell. B.. and triglycerides with dietary fibers in the small intestine of rats. Cara.. Isaksson.. 37C. 1983. 21. Sandberg. Peptic inhibition by macroanions. Gastroenterol. I. Interaction of bile acids.. P. Scand. 27.. and Hulten. B... Ikegami. B. O. 283. Gallaher. 24. Factors affecting the rate of hydrolysis of starch in food.. Brown. M. 1989.. S. B. L. Harada. 8. S. E. Borel.. Clin. Emulsification and lipolysis of triacylglycerols are altered by viscous soluble dietary fibres in acidic gastric medium in vitro. Biochem. A. J. Guillon. Stock-Damgé. Pasquier. Borel.... Armand. J. Satchithanandam.. Cassidy.. 2001 7:30 PM EFFECTS OF DIETARY FIBER ON DIGESTIVE ENZYMES 285 30. Kritchevsky. J... F. 7. 47. Nutr. D. O. R. 41... S.. D. Barry. M. and Schneeman. D.. 427. P. Poksay.. B. Pasquier.8_fm Page 285 Sunday. Clin.. May 6. Tepper. G. C. L.. 33.. J. Am. Effects of wheat bran on the exocrine pancreas and the small intestinal mucosa in the dog. 269–275.. 1249. Exp. O. B. M. B. C. and Bouchet. 31... Borel.. Viscous soluble dietary fibres alter emulsification and lipolysis of triacylglycerides in duodenal medium in vitro. 201.. M. Dietary fiber and intestinal adaptation: effects on lipid absorption and lymphalic transport in the rat. Humbert. Castelain.. F. Satchithanandam. Pancreatic and intestinal response to dietary guar gum in rats.. Vahouny. 99–108.2387_ch4. G. J. 113. M.. Biochem. J. and Lairon. S. H. 114.. Armand. . Chen. 1984. Castelain.. Soluble fiber and dietary lipids.. B. 35. Lairon. V. V. 1076. C... Am.. 1544. I. F. 1997... 1985. Pieroni.. J. 1996. Lightfoot. Adv. 34.. D. Calvert. 32. Nutr. Med.. M. 1988. F. 293–302. Nutr. 314.. and Vahouny. K. Schneeman.. G. Biol. G. Dietary fiber and intestinal adaptation: effects on intestinal and pancreatic digestive enzyme activities. 1983. 36. and Cassidy. and Lairon. B.. Guillon. Aprahamian. Nutr. B. M.. Nutr... Lafont. S.. Raul. S. W. 2387_ch4. May 6.8_fm Page 286 Sunday. 2001 7:30 PM . 3 Decreased fecal excretion may be due to increased SCFA absorption.9 The Source of Dietary Fiber Influences — Short-Chain Fatty Acid Production and Concentrations in the Large Bowel H. absorption. propionate. In the previous edition.9_fm Page 287 Sunday.9. This chapter summarizes data which show the effects due to dietary fiber on SCFA production and excretion. dog. May 6. also referred to as volatile fatty acids. which compose more than 80% of all SCFAs. we extended the data from studies conducted from 1990 to the present.50 © 2001 by CRC Press LLC 287 . and increased SCFA absorption. They are produced from carbohydrates or dietary fiber which cannot be degraded by human digestive enzymes. and rabbits are excluded due to important differences from humans in their intestinal physiology and functions. since they stimulate SCFA production.47 Carbohydrate and dietary fiber affect colonic metabolism and the luminal environment. pigs.00+$1. and butyrate. In this edition. absorption. These earlier data showed also that the age of animal used in experiments might affect SCFA production. DURATION OF FEEDING DIETARY FIBER The studies reported in the previous edition of this review48 generally showed that increasing the duration of dietary fiber consumption increased SCFA concentration in the intestinal lumen. and excretion which. increased SCFA digesta pool size. and excretion. cats. and humans are included.48 the summarized data illustrated the importance of intestinal microflora on SCFA production and of having internal consistency within each experiment regarding gender of animal due to differences between gender in SCFA production. absorption. and excretion. Kobayashi and Sharon E. and on fecal pH. Data from ruminant animals. affect characteristics such as luminal and fecal pH.2387_ch4. in turn. as shown in an earlier rat study. primarily include acetate. but which are hydrolyzed and fermented by intestinal microflora in the large bowel. two studies showed increases in SCFA concentration in cecal digesta production with increasing duration of dietary fiber consumption. animal models are widely used.48 0-8493-2387-8/01/$0. Fleming Short-chain fatty acids (SCFA). 2001 7:31 PM CHAPTER 4. Since SCFA production cannot easily be measured directly in humans due to difficulties obtaining human colonic contents.1.2 while one study showed a possible decrease in SCFA excretion in feces with increased exposure to fiber. Of the three more recent studies (Table 4.1). Data from various strains of rats. 5 14.8 16.1 ± 0.7 16 wk 15.0 6 mo 47. then ↓ [Bu] at 27 wks ↑ duration → ↑ [Pr] at 27 wks ↑ duration → ↑ [Ac] for FOSb at 27 wks Comments 3 2 1 Ref.9. male Sprague-Dawley 40–60 g starch starch starch Cecal contents (Mean.1 ± 3.2 ± 2.3 ± 1.a b Significantly different from control group p <0.4 ± 0.2 21.3 25.4 ± 0.0 23.8 a 12.7 26.3 ± 1.9 53. FOS = fructooligosaccharides.3 ± 2.05.6 69. 2001 3:02 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.9 ± 2. n = 10.2 ± 4.6 94 a 19.0 14.0 64.2 103.7 20.3 40.0 11.5 a 19.3 5. male Wistar. µmol/g wet contents) Acetate Control Fructooligosaccharide 55.2 a 60.3 a 73. 115 g Specimen and Diet Influence of the Duration of Feeding on SCFA Concentrations in Luminal Contents and Feces Animal Model Table 4.2 162.7 32. 288 Rats.5 ± 5.5 ± 2.5 5.0 90.2 17.3 a 27 wk 47.3 a 8 wk 55.3 ± 10.5 16.3 18.1 ± 2.5 ± 4.7 ± 1.3 31.4 ± 5.9 a 12.1 7.3 a 62.4 2 mo 55.4 ± 13.8 2.3 23.5 mo 44. µmol/g wet) Acetate Wheat bran Oat bran Propionate Wheat bran Oat bran Butyrate Wheat bran Oat bran Total SCFA Wheat bran Rats.1 12.7 15.9 66.2 a 62. n = 6.4 a 73.9 28.8 4 wk 24.5 ± 2.5 a 2 wk 44.9 13. male Wistar.4 115.4 ± 1.7 45.0 83.1 2387_ch4.1 4.9 ± 1.9 37.0 ↑ duration → may ↓ SCFA excretion Differences between diets were statistically significant ↑ duration → ↑ [total SCFA] ↑ duration → ↑ [Bu] ↑ duration → ↑ [Pr] ↑ duration → ↑ [Ac] ↑ duration → ↑ [total SCFA] ↑ duration → may ↑ [Bu] for FOS at 8 wks.7 ± 3. Oat bran Feces (Mean ± SEM.4 5.5 a Duration of Feeding 46. 115 g Rats. µmol/g wet) Acetate Control Resistant potato Propionate Control Resistant potato Butyrate Control Resistant potato Total SCFA Control Resistant potato starch Control Fructooligosaccharide Total SCFA Butyrate Control Fructooligosaccharide Control Fructooligosaccharide Propionate Cecal contents (Mean.3 ± 2.6 ± 2.7 4.6 7.8 2.9_fm Page 288 Tuesday.0 55. May 8.2 a 16.6 ± 1.0 12.6 4.2 25.4 a 79.7 ± 4.4 ± 0.4 4.7 ± 1.7 0.5 2.4 8.9 8 wk 17.4 94.8 4.5 ± 2.9 a 79.1 ± 1. 3RD EDITION .9 4.3 4.5 12 wk 14.8 ± 3.5 16.3 4. n = 6.2 ± 1.9 11.4 5. when compared to a lowfiber diet. cellulose were consistent among the cecum. and distal colon.4). SCFA concentrations were highest in the contents of the cecum and lowest in feces. Based on earlier studies. oat bran.3. As one example. however. caution should be used in extrapolating these findings to physiological situations.2 and differences for pectin and oat bran vs. concentration in the diet..12 Some inconsistencies among studies are evident.11 studies. wheat bran.9. In most. potato starch increased the concentrations of acetate and butyrate in the intestinal contents of both the proximal and distal colon of rats. it is possible to use data from one segment to predict the relative differences among dietary fibers in another segment. this increase was offset by a decrease in the proportion of propionate.12 with only one exception. acetate constituted a greater proportion of the three SCFAs in contents of the distal colon than in the cecum or proximal colon. a high-fiber diet caused total SCFA concentrations to be higher in lumenal contents of both the cecum and colon of rats.2. Specifically. proximal colon.3. The reader is referred to the earlier review.5 Both studies were conducted for only 1 week.9_fm Page 289 Sunday.48 In several recent studies.10.10 was similar for treatments including potato starch. but not the wheat bran. thus. the molar ratios of propionate and butyrate increased. insufficient data are available to conclude that differences in fecal SCFA concentrations can equally well predict dietary differences in the luminal contents of the cecum or colon.11 but not all.2.9. it was noted that feeding cellulose resulted in lower SCFA production than pectin. proximal colon.6–8.4 With both of these fiber sources.10.2). Although not the purpose of this review. By comparing data within the study. the effects of a dietary fiber on SCFA concentrations were consistent along the length of the large bowel. since the effects of concentration of fiber on SCFA production and excretion was more thoroughly studied in the previous years.9 oat bran.3).9 and was lower in the cecum than the distal colon for treatments including rolled oats12 and wheat bran. fecal SCFA concentrations were increased by increasing the vegetable fiber. that differences between segments in molar percentages of the three SCFAs were not consistent. whereas butyrate percentages were unchanged. it appears that the source of dietary fiber may play a role in determining the longitudinal gradient in molar percentages among the three major SCFAs. and distal colon contents of rats. Unfortunately. and psyllium.2387_ch4.6–9.8 and pectin. 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 289 CONCENTRATION OF FIBER IN DIET The influence of the amount of fiber ingested was evaluated by increasing the concentration of fiber in the diet (Table 4.12 Calculations performed on original data indicated.3 and 4. are reported for several fiber sources (Table 4.7 Similarly.9. May 6.9 Differences among segments in the molar ratios of the three major SCFAs are worth noting. THE SOURCE OF FIBER INFLUENCES SCFA PRODUCTION Comparisons among fiber sources have been made most commonly by feeding diets that contain equivalent concentrations of fiber but which differ in the source from which the fiber is derived (Tables 4. In one of the two recent studies.8 As a result.3. data suggest that cellulose consumption may depress cecal SCFA concentrations below those .9. the proportion of acetate was higher in the cecum than the distal colon for treatments including cellulose. In the one human study. and in excreted feces.2 wheat bran. whereas the molar ratio of acetate decreased with increasing dietary fiber concentrations. THE EFFECTS OF FIBER ON SCFA CONCENTRATIONS ARE OBSERVED ALONG THE LENGTH OF THE LARGE BOWEL The SCFA concentrations in intestinal segments including the cecum. SCFA concentrations in rat cecal contents were increased in a dose-dependent manner with increasing dietary concentrations of both inulin and resistant starch. In the one study where these were reported.8 pectin.8 and psyllium. SCFA conc, mM Molar Percentage (Ac/Pr/Bu) Fecal dyalysate fluid Mean ± SEM, n = 9 SCFA conc, mM Molar Percentage (Ac/Pr/Bu) SCFA conc, mM Molar Ratio (Ac:Pr:Bu) SCFA conc, mM Molar Ratio (Ac:Pr:Bu) Cecal contents Mean ± SEM, n = 10 a Significant differences from fiber-free group. Note: Ac = acetate; Pr = propionate; Bu = butyrate. Veg fiber Wheat bran Resistant starch Inulin Specimen/Data 58.2 ± 3.6 55/23/9 Fiber free 101 ± 7 69:24:07 Fiber free Fiber free 101 ± 7 69:24:07 Vegetable fiber (10g) 74.5 ± 4.1 58/22/10 Wheat bran (10g) 99.3 ± 3.6 55/22/14 Resistant starch (5%) 114 ± 6 61:26:13 Inulin (3.75%) 104 ± 6 60:29:11 Vegetable fiber (30g) 90.0 ± 4.1 60/22/12 Wheat bran (30g) 75.5 ± 3.4 52/22/16 Resistant starch (10%) 149 ± 9a 56:27:17 Inulin (7.5%) 117 ± 6 50:33:17 Source (and Level) of Fiber in Diet Resistant starch (20%) 173 ± 10a 53:30:17 Inulin (15%) 129 ± 8a 43:37:20 ↑ [fiber] → ↑ [SCFA] No change in molar percentage No change in [SCFA] No change in molar percentage ↑ [fiber] → ↑ [SCFA] ↑ [fiber] → ↓ %Ac, ↑ %Bu ↑ [fiber] → ↑ [SCFA] ↑ [fiber] → ↓ %Ac, ↑ %Bu Comments 5 4 Ref. 290 Human, fed 8 days Rats, male Wistar, 200 g, fed 7 days Fiber Influence of the Dietary Fiber Concentration on SCFA Concentration in Cecal Contents and Fecal Fluid Animal Model Table 4.9.2 2387_ch4.9_fm Page 290 Sunday, May 6, 2001 7:31 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Rats, male Wistar, 115 g Rats, male Wistar, fed 18 mo Rats, male Wistar, 150 g, fed 28 days Luminal contents Mean, n = 6, µmol/g wet Acetate Proximal colon Distal colon Propionate Proximal colon Distal colon Butyrate Proximal colon Distal colon Total SCFA Proximal colon Distal colon Colon Potato starch 66.3* 52.7* 16.6 10.9 30.5* 25.2* 114.5* 89.5* Basal 29.6 15.5 11.9 6.8 2.8 1.7 45.3 24.5 135.9 ± 12.1* 296.6 ± 18.4* 60.2 ± 10.3 24.7 ± 3.5 High fiber (133 g NSP/kg) 120.4 ± 12.1 218 ± 24.9 380 ± 23.4 Wheat Bran (10%) Low fiber (17 g NSP/kg) 158.5 ± 15.7 Distal colon Luminal contents Mean ± SEM, n = 10, µmol/g dry Total SCFA Cecum 201 ± 16.5 441 ± 17.9 Basal 180 ± 16.4 326 ± 20.2* 517 ± 26.7* Ispagula (5%) Source (and Level) of Fiber in Diet Proximal colon Total SCFA Cecal Luminal contents Mean ± SEM, n = 9–20, µmol/g dry Specimen/Intestinal Segment Comments Potato starch increases the conc. of each SCFA in both the proximal and distal colon [Total SCFA] higher for high-fiber diet in cecum [Total SCFA] higher for high-fiber diet in colon [Total SCFA] higher for ispagula in cecum [Total SCFA] higher for ispagula in prox. colon No differences among fibers in distal colon Influence of the Source of Dietary Fiber on SCFA Concentrations and Proportions in Various Intestinal Segments Animal Model Table 4.9.3 2 7 6 Ref. 2387_ch4.9_fm Page 291 Sunday, May 6, 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 291 Luminal contents Mean ± SEM, n = 19–20, µmol/g Acetate Cecum Proximal colon Distal colon Propionate Cecum Proximal colon Distal colon Butyrate Cecum Proximal colon Distal colon Total SCFA Cecum Proximal colon Distal colon † 7.9a 6.7a 3.9 ± 0.3b 1.3b 1.25b 1.9b 62.8 ± 2.6a 40.8 ± 1.7a 21.4 ± 1.0b 5.9b 4.2b 1.9c 1.7b 0.8b 0.6c 37.9 ± 1.3b 20 ± 1.25b 9.7 ± 0.3c 53.5 ± 4.0 53.5 ± 4.6 45.4 ± 4.1 38.7 ± 4.1 52.3 ± 2.0a 31.7 ± 1.7a 13.6 ± 0.6b 17.2 ± 1.7 19.0 ± 2.0 9.6 ± 1.8 8.2 ± 1.8 28.8 ± 1.3c 13.3 ± 0.8b 5.8c 6.7 ± 0.6 7.0 ± 0.7 4.9 ± 0.6 4.6 ± 0.6 Pectin (6%DF) 27.1 ± 2.5 23.8 ± 2.9 29.2 ± 2.6 24.2 ± 2.6 Cellulose (6%DF) Oat bran Wheat bran 57.6 ± 2.6a 42.5 ± 1.7a 33.1 ± 1.3a 11.8a 5.4a 5.2 ± 0.3a 7.2a 6.7a 4.9 ± 0.3a 36.6 ± 2.6b 28.8 ± 1.7a 21.4 ± 1.0a Oat bran (6%DF) Source (and Level) of Fiber in Diet [total SCFA] for pectin and oat bran > cellulose at all three sites [Bu] for oat bran > cellulose and oat bran at all three sites [Pr] for pectin and oat bran > cellulose at all three sites [Ac] for pectin and oat bran > cellulose at all three sites [SCFA] are similar for wheat and oat bran at both the proximal and distal colon Comments 8 3 Ref. 292 Rats, male Sprague-Dawley, 270–320 g, fed 4 wk Colonic contents Mean ± SEM, n = 11, µmol/g Acetate Proximal colon Distal colon Propionate Proximal colon Distal colon Butyrate Proximal colon Distal colon Total SCFA Proximal colon Distal colon Specimen/Intestinal Segment Rats, male Sprague-Dawley 40–60 g Animal Model Table 4.9.3 (Continued) Influence of the Source of Dietary Fiber on SCFA Concentrations and Proportions in Various Intestinal Segments 2387_ch4.9_fm Page 292 Sunday, May 6, 2001 7:31 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Rats, male Luminal contents and feces Sprague-Dawley, Mean, µmol/g wet 208–234 g body wt, Acetate fed 2 wk Cecum Proximal colon Distal colon Fecal Propionate Cecum † Proximal colon Distal colon Fecal Butyrate Cecum Proximal colon Distal colon Fecal Rats, male Luminal contents Sprague-Dawley, Mean ± SEM, n = 10, mmol/l 200–230 g body wt, Acetate fed 3 wk Cecum Proximal colon Distal colon Propionate Cecum Proximal colon Distal colon Butyrate Cecum Proximal colon Distal colon Total SCFA Cecum Proximal colon Distal colon † 9.2 ± 4.9 13.3 ± 7.4a 5.1 ± 1.5a 2.5 ± 0.8b 3.1 ± 2.0b 3.1 ± 1.4b 65 ± 23a,b 110 ± 40a 43 ± 7 9.5 ± 3.6 1.5 ± 0.8b 0.3 ± 0.1b 4.7 ± 1.0b 1.6 ± 0.8b 0.4 ± 0.4b 45 ± 13a 60 ± 10b 45 ± 10 4a 3.6 1.8 0.9a 4a 3.1 2.2 0.4a 10.3a 14.2b 5 7.5 5.3b 29.3a 30 23.3 23.6b 35.3b 26 13.3 12.6a 10.7 4.6 1.1a with Psyllium Alone 92 ± 18b 97 ± 20a 53 ± 13 1.6 ± 3.3a 12.4 ± 2.4a 11.6 ± 2.7a 9.0 ± 2.1 10.8 ± 3.1a,b 3.1 ± 1.5a,b 64.2 ± 14.0 72.6 ± 14.0a,b 37.7 ± 8.4 Wheat bran 9.3b 7.1 5.3 2.7b 10.7 8.9 5.3b 14.2b 34b 31.3 16.6 18a,b Alone 8.7b 4.9 6.4 5.3c 4.6 6 8.5b 12.4b 29.3a 26 23.3 22.6b with Psyllium High Amylose Starch 53.0 ± 19.5 92.1 ± 11.2a 36.3 ± 5.6 39.1 ± 11.2 55.8 ± 11.2b 44.7 ± 11.2 Low Amylose Starch Pectin (8% DF) Fiber-free (8% DF) Psyllium had little or no effect on [Bu] at all sites With low amylose, psyllium → ↓ [Pr] in cecum ↑ [Pr] in feces Psyllium → ↓ [Ac] in cecum ↑ [Ac] in feces In prox. colon only, [total SCFA] for pectin and wheat bran > fiber-free At all 3 sites, [Bu] for wheat bran > fiber-free and pectin In prox. and distal colon, [Pr] for pectin > fiber-free In prox. colon only, [Ac] for pectin > fiber-free 10 9 2387_ch4.9_fm Page 293 Sunday, May 6, 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 293 † Pigs, fed 34 days Rats, male Sprague-Dawley, 40–60 g, fed 38 wk 51/40/6 55/31/6 Alone 44/40/6 55/31/8 51/40/7 51/36/7 Wheat Flour w/ Aleurone w/ Pericarp 3.86 ± 0.28 Distal colon Luminal contents Mean, n = 4 Molar percentage (Ac/Pr/Bu) Cecum Proximal colon 4.68 ± 0.30 4.22 ± 0.19 56/36/7 56/34/8 w/Bran 4.36 ± 0.26 3.01 ± 0.15 4.22 ± 0.21 4.79 ± 0.39 2.68 ± 0.34 29.1 ± 3.6 15.3 ± 1.3 50.7 ± 5.1 50b 35.6 35.6 36.7c 21.8 ± 2.32 57.8c 48.9 31.1 26.7b Pectin (6 g/100 g diet) 43.3a 37.8 32.2 24.4b Cellulose (6 g/100 g diet) 53.3b,c 40 21.1 14.4a Source (and Level) of Fiber in Diet Distal colon Butyrate Proximal colon Distal colon Propionate Proximal colon Luminal contents Mean + SD, n = 5 , µmol/g wet Acetate Proximal colon Total SCFA Cecum Proximal colon Distal colon Fecal Specimen/Intestinal Segment Adding wheat bran to wheat flour did not, substantially change the percentages of the three SCFAs [Bu] for pectin and cellulose similar at both sites [Pr] for pectin and cellulose similar at both sites [Ac] for pectin > cellulose at both sites Psyllium → ↓ [total SCFA] in cecum ↑ [total SCFA] in feces Comments 12 11 Ref. 294 Animal Model Table 4.9.3 (Continued) Influence of the Source of Dietary Fiber on SCFA Concentrations and Proportions in Various Intestinal Segments 2387_ch4.9_fm Page 294 Sunday, May 6, 2001 7:31 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Pigs, fed 42 days 55/34/6 60/23/8 69/14/6 Molar percentage (Ac/Pr/Bu) Cecum Proximal colon Distal colon 50/38/8 57/26/8 66/15/9 52/38/7 52/34/9 58/22/9 133 143.8 75.7 100.5 91.9 54.1 117.8 108.1 51.9 Alone 63/23/7 Wheat Flour Alone & Oat Bran 59/26/6 Luminal contents Mean, n = 4 Total SCFA (mmol/l) Cecum Proximal colon Distal colon 70/18/6 * Significantly different from basal group p < 0.05. † Data was extrapolated from the original graphs. a,b,c Values in a row with different superscripts are significantly different at p < 0.05. † Distal colon 53/31/11 55/28/12 61/19/10 144.9 121.1 60.5 Rolled Oats & Oat Bran 69/19/7 Adding oat bran did not consistently influence percentages of the three SCFAs In comparison to the cecum, the distal colon tended to have higher % Ac and lower % Pr Adding oat bran to wheat flour or rolled oats tended to increase [SCFA] in cecum, but decrease [SCFA] in distal colon contents In comparison to the cecum and prox. colon, the distal colon appeared to have higher % Ac and lower % Pr 12 2387_ch4.9_fm Page 295 Sunday, May 6, 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 295 2387_ch4.9_fm Page 296 Sunday, May 6, 2001 7:31 PM 296 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION observed for animals fed a fiber-free diet.15,16 More recent studies confirmed previous observations48 that, in comparison to cellulose, pectin ingestion increased luminal SCFA concentrations and pool size.8,15 This study extended our knowledge by showing that cecal SCFA concentration and pool size was higher also for pea fiber than cellulose.14 In comparison to a cellulose diet, the cereal fibers oat bran and corn bran resulted in higher luminal propionate concentration in pigs21 and higher concentrations of each of the three major SCFAs when this comparison was made using rats.8 When compared to methylcellulose consumption, cecal SCFA concentrations in rats were higher for diets containing both rice bran and wheat bran.17 The results of these most recent studies show that water-dispersible fibers such as inulin, pectin, indigestible dextrins, and ispagula provide higher luminal SCFA concentrations than do cereal fibers or brans from wheat, oats, and corn.6,14,18,19 Furthermore, it appears that fiber from chicory, carrots, and cocoa might respond similarly to inulin.14,18 Among the cereal fibers, wheat bran, corn bran, and oat bran may support similar luminal SCFA concentrations, while rice bran might support higher concentrations.3,14,17,21 Dietary Fiber Source Influences Fecal SCFA Excretion The generalizations that were made earlier regarding the influence of various sources of dietary fibers on luminal concentrations of SCFA appear to apply, but to a more limited extent to fecal excretion of SCFA (Table 4.9.5). In particular, there is evidence that fecal excretion of SCFA is increased when water-dispersible compounds such as ispagula,6,27 inulin,24 galactooligosaccharides,24 senna,25 and resistant starch26 are consumed. By contrast, wheat bran had little or no effect on fecal SCFA excretion.6,22,23,25 Many Dietary Fiber Sources Reduce Luminal pH The effect of the dietary fiber on luminal pH has been studied most commonly in rats, although fecal pH data are also available for humans (Table 4.9.6). In general, wheat bran did not have a consistent significant effect on either luminal or fecal pH using rats,9,28 pigs,12 or humans.25 Also, cellulose consumption did not appear to lower luminal pH.29 Luminal and fecal pH, however, appeared to be reduced following consumption by rats of dietary fibers including fructooligosaccharides,1 galacto-oligosaccharides,29 inulin,4,14 pectin,8,9 psyllium,21 oat bran,8,12 and following consumption of diets high in resistant starch.4,13,26 Results using inulin and resistant starch suggest that there is a dose–response relationship such that increasing fiber consumption shows further decreases in luminal pH.4 Results of another study suggest that there may be a threshold, however, after which further increases in fiber intake do not further influence fecal pH.30 Overall, the pH values of control treatments, even when compared within a specific site, varied considerably among the experiments. If one focuses simply on the absolute pH values, rather than on the significance of differences between a specific fiber treatment and a control, then it appears that luminal pH values of 6 or less can be attained by feeding water-dispersible but fermentable constituents including inulin, galactooligosaccharides, resistant starch and, in some cases, fructooligosaccharides. In Vitro SCFA Production Varies with the Source of the Dietary Fiber Since it is difficult to examine the fermentation dynamics within the intestine of humans or animals, in vitro fermentation systems have been developed and utilized to study the influence of dietary fiber (substrate) on SCFA production (Table 4.9.7). Many research groups presented data for SCFA as a function of fermentation time.6,21,22,34,35,37,40–46 In general, time-dependent increases in SCFA production were observed. Since SCFA production tended to level off after a 24-hour incubation period, data are presented only for this later time point in an attempt to simplify this presentation. Rats, male Sprague-Dawley, 150–200 g, fed 4 wk Rats, male Fischer, 10 wk, 280 g, fed 8 wk 48 ± 6a 72/21/4 SCFA pool, µmol Molar percentage (Ac/Pr/Bu) 37.1 ± 2.3a 48.9 ± 3.4c 18.4 ± 1.4c 3.3 ± 0.3b,c 70.6 ± 4.9c Butyrate, µmol/animal Total SCFA, µmol/animal 33.1 ± 4.6d 1.7 ± 0.3c Cellulose (30% DF) 15.7 ± 2.7b 21.9 ± 2.8d 9.5 ± 1.6d Pea 23.91 ± 1.22b 9.44 ± 0.26 3.12 ± 0.60b 37.62 ± 1.84b Wheat bran 28.40 ± 1.53a,b 9.42 ± 0.41 4.14 ± 0.33b 43.20 ± 2.11b Acetate Propionate Butyrate Total SCFA Control 46.02 ± 3.66a,b 55.83 ± 2.69a Total SCFA Cecal contents Mean ± SEM, n = 5 Total SCFA, mmol/l Acetate, µmol/animal Propionate, µmol/animal 8.80 ± 0.55 3.66 ± 0.27b 32.66 ± 3.04a 31.42 ± 1.91a 8.65 ± 0.26 14.83 ± 0.88a Carrot Inulin 69/19/5 Propionate Butyrate Cecal contents Mean ± SEM, n = 6, µmol/g wet Acetate 76 ± 5a,b 58 ± 4a 78 ± 10a Pea starch Control Rats, male Wistar, 75 ± 5 g, fed 12 days 515 ± 78a 7.1 ± 2.5b Pectin (30% DF) 45.9 ± 3.2a 386.4 ± 58.2a 121.4 ± 20.6a Oat 23.27 ± 1.29b 9.15 ± 0.35 3.20 ± 0.29b 36.81 ± 1.65b 45.99 ± 0.97a,b 8.71 ± 0.21 4.16 ± 0.30b 32.16 ± 0.65a Cocoa 71/15/10 150 ± 19b 85 ± 4b Potato starch Source of DF (Level) Cecal contents Mean ± SEM, n = 7 SCFA conc. mmol/l Specimen/Data 244 ± 12.8b 25.4 ± 1.6a Pea fiber (30% DF) 46.3 ± 3.6a 167.4 ± 8.7b 51.3 ± 5.7b 72/16/8 153 ± 15b 61 ± 4a Resistant starch Influence of the Source of Dietary Fiber on SCFA Concentration, or Quantity, in Cecal Contents Animal Model Table 4.9.4 Cellulose had lowest conc. of all SCFAs Pectin had greatest [Ac] and [Pr] Pea fiber had greatest [Bu] [Bu] higher for inulin than others [Ac] for Inulin, carrot, and cocoa > others [SCFA] for potato starch > resistant starch SCFA pools for potato and resistant > pea starch Molar ratios not different Comments 15 14 13 Ref. 2387_ch4.9_fm Page 297 Sunday, May 6, 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 297 Rats, male Sprague–Dawley, 152 g, fed 4 wk Butyrate Cecal contents Mean, n = 8, mmol/g wet Acetate Propionate Butyrate Propionate Cecal contents Mean ± SEM, n = 8, mmol/l Acetate Rats, male Sprague-Dawley, 150–200 g, fed 4 wk Pool, µmol/ cecum Acetate Propionate Butyrate Butyrate Cecal contents Mean ± SE, n = 6 Conc, µmol/ g Acetate Propionate Specimen/Data Cecal contents Mean, n = 8 Molar percentage (Ac:Pr:Bu) Rats, male Sprague-Dawley, 125–150 g, fed 22 days Animal Model 7.1 ± 1.2c 9.1 7.8 41.6 28.0b,c 55.2 24.8c 1.6 ± 0.3a 5.3 ± 0.6a Chicory (1%) 4.9 ± 0.3a 23.5 ± 1.5c Fiber-free Methylcellulose 13.1 ± 0.7a Low Amylose/ Psyllium (1.5%) 68:24:9 Fiber-free 70.0 ± 3.9c Low Amylose Starch 66:26:8 85.7 ± 6.9b 39.4 ± 2.7b 27.4 ± 1.7b 10.7 ± 0.6b 33.9 ± 5.3 17.3 ± 1.8b 41.1 ± 6.9c 24.0 ± 2.7c 10.3 ± 1.7c Maize husk (100 g/kg) 32.7 ± 1.8 16.1 ± 0.6b Fiber-free 9.0 51.5 33.6a Chicory (5%) 18.0 ± 1.0b 16.0 ± 1.0b Parboiled rice bran 67.0 ± 3.0c High Amylose Starch 59:25:16 116.6 ± 6.9a 68.6 ± 6.9a 37.7 ± 3.4a 14.3 ± 0.6a Acid-treated Maize husk (100 g/kg) 40.5 ± 2.4 23.2 ± 2.4a Source of DF (Level) 8.0 40.9 31.5a,b Inulin (5%) 15.2 ± 0.9b 13.6 ± 0.9b Coarse WB 32.9 ± 2.0b High Amylose/ Psyllium (1.5%) 58:25:17 [Pr] for chicory and inulin > fiber-free group Low levels of all SCFAs were seen for methylcellulose [Pr] was significantly higher for no-fiber group [Bu] rice and wheat bran > others High amylose starch had lower % Ac and higher % Bu Higher amount of SCFA for maize husk groups Acid treating husks increased SCFA concs. and pools Comments 18 17 10 16 Ref. 298 Rats, male Sprague-Dawley, 208–234 g, fed 2 wk † Table 4.9.4 (Continued) Influence of the Source of Dietary Fiber on SCFA Concentration, or Quantity, in Cecal Contents 2387_ch4.9_fm Page 298 Sunday, May 6, 2001 7:31 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Cecal contents Mean ± SD, n = 18, µmol/g Acetate Propionate Butyrate Total SCFA Rats, male Sprague-Dawley, 5 wks old, fed 28 wk † e,f,g a,b,c,d 52.9 ± 3.8b 39.0 ± 4.0a,b Values in a row with different superscripts are significantly different at p < 0.05. Values in a row with different superscripts are significantly different at p < 0.001. Data was extrapolated from the original graphs. Total SCFA 27.9 ± 2.1 17.0 ± 2.3b 5.5 ± 0.7b 25.2 ± 2.3 10.0 ± 1.2a,b 2.6 ± 0.5a SCFA pool, mmol/animal Acetate Propionate Butyrate 55:33:12 67:26:07 9.2 ± 2.3 81.4 ± 8.6a,b 42.6 ± 3.6 25.7 ± 3.0b 37.9 ± 1.7 14.9 ± 1.0a 3.9 ± 0.5 58.5 ± 3.5a Oat bran (7%) ± ± ± ± Cellulose (7%) 169.4 51.0 59.0 279 43.9f 18.9f 16.4f 72 ± ± ± ± 31.0f 10.9g 10.5g 43 148.3 31.3 29.1 208 Casein/Wheat 93.4 ± 7.1b 40.5 ± 2.2b 19.7 ± 1.6b 48.8 ± 7.8a 18.5 ± 1.5a 8.4 ± 1.1a Casein/Starch Indigestible dextrin (5%) Control Molar ratio (Ac:Pr:Bu) Butyrate Total SCFA Pigs, male, Cecal digesta Hanford Miniature, Mean ± SEM, n = 5 ~5 mo old, 40 kg SCFA conc., mmol/l body wt Acetate Propionate Cecal contents Mean ± SEM, n = 7–9 µmol/cecum Acetate Propionate Butyrate Rats, male Sprague- Dawley, 6 wks old, fed 8 wk ± ± ± ± 48.0e 17.8e 19.0e 66 34.6 ± 7.3a 21.8 ± 5.2 9.1 ± 2.0a,b 2.5 ± 0.7a 65:28:07 6.4 ± 0.7 88.5 ± 7.2b 55.4 ± 4.3 23.7 ± 3.1b Corn bran (7%) 231.8 78.2 81.5 390 Chickpea/Wheat 96.8 ± 13.5b 45.2 ± 8.1b 21.2 ± 3.5b Pectin (5%) 33.3 ± 4.6a 22.1 ± 3.5 9.0 ± 1.2a 2.7 ± 0.5a 64:27:08 5.9 ± 1.0 72.6 ± 8.0a,b 47.1 ± 3.7 20.1 ± 2.8a,b Corn bran (13%) 64.1 ± 7.8a 22.6 ± 2.7a 12.7 ± 2.3a Corn fiber (5%) Bu and total SCFA pools for oat bran > corn bran [total SCFA] for 7% corn bran > cellulose Oat bran had lower % Ac and higher % Bu than others [Pr] for oat and corn bran > cellulose Chickpea had highest [Ac], [Pr], and [Bu] Output of each SCFA for dextrin and pectin > others 21 20 19 2387_ch4.9_fm Page 299 Sunday, May 6, 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 299 Feces Mean ± SEM, n = 24 Concentration (mmol/l) Acetate Propionate Butyrate Output (mmol/d) Acetate Propionate Butyrate Feces Mean ± SD, n = 12 mmol/100 g dry Acetate Humans, 12 male, 12 premenopausal female, 33 ± 2 yr old Humans, 12 male, 23 ± 3 yr, fed 3 wk 671 ± 297 313 ± 177 795 ± 300 401 ± 179 1181 ± 355* 854 ± 541 707 ± 297 318 ± 105 Fructooligosaccharides (15 g/d) 1058 ± 444 16.82 ± 2.13* 3.86 ± 0.60* 5.35 ± 0.66* 10.21 ± 1.62 2.34 ± 0.38 3.07 ± 0.51 Inulin (15 g/d) 64.2 ± 3.9 14.3 ± 1.2 21.3 ± 1.9* 60.7 ± 4.6 14.2 ± 1.2 19.2 ± 2.4 Control Wheat bran (30 g DF/d) Oat bran (93g NSP/kg) 270 ± 62.7 57/21/9 High-amylose starch 70/21/11 Ispagula (5%) 572.6 ± 54.7 Elemental Low-fiber (56g NSP/kg) 248 ± 61.6 60/21/6 Low-amylose and psyllium 76/21/3 Wheat bran (10%) 287 ± 51.3 Source (and Level) of Fiber in Diet Psyllium reduced % Ac and increased % Pr or Bu 679 ± 293 369 ± 199 Galactooligosaccharides (15 g/d) 1246 ± 698* Acetate excretion for inulin and GOS > control and FOS Wheat bran → ↑ fecal output of Ac, Pr, and Bu Wheat bran → ↑ [Bu] Wheat bran (102g NSP/kg) 252 ± 66.2 63/21/6 Oat bran gave significantly higher % Bu High amylose and psyllium 66/24/14 SCFA excretion for ispagula > wheat bran > basal Comments 24 23 22 10 6 Ref. 300 Propionate Butyrate Feces Mean ± SD, n = 5 Total SCFA (mmol/kg dry) Molar percentage (Ac/Pr/Bu) Pigs, barrow Cross-bred, 50–85 kg, fed > 7 days Low-amylose starch 90/6/3 127.4 ± 11.4 Total SCFA, µmol/day Feces n=8 Molar percentage (Ac/Pr/Bu) Basal Feces Mean ± SEM, n = 9–20 Rats, male Sprague-Dawley, 220 g, fed 2 wk Rats, male Wistar, 150 g, fed 28 days Specimen/Data Influence of the Source of Dietary Fiber on Fecal Excretion of SCFA Animal Model Table 4.9.5 2387_ch4.9_fm Page 300 Sunday, May 6, 2001 7:31 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION 7.5 ± 1.3 7.3 ± 1.0 44.7 ± 4.4 25.7 ± 2.3 Placebo 24.9 ± 5.4 11.1 ± 2.0 100.9 ± 12.6 Ispagula (18g/d) 60.1 ± 6.3 * Significant difference from control at p < 0.05. Note: NSP = non-starch polysaccharide; GOS = galactooligosaccharides; FOS = fructodigosaccharides. Propionate Butyrate Total SCFA Fecal water Mean ± SEM, n = 7, mmol/l Acetate Humans, 5 male, 2 female, 21–25 yr old, fed 15 days Low-resistant starch (fresh maize porridge) 65.8 24.8 17.6 116.1 Highly resistant starch (stale maize porridge) 93.9* 43.1 35.1* 182.6* Loperamide 51.6 16.6* 6.0* 82.0* Control 79.8 27.2 24.8 152.0 Acetate Propionate Butyrate Total SCFA Feces Mean, n = 10, mmol/kg dry Acetate Propionate Butyrate Total SCFA Senna 138.2* 40.3* 59.1* 202.0* Control 63.9 18.4 16.6 111.0 Acetate Propionate Butyrate Total SCFA 72.9 24.0 69.0 95.0 Wheat bran (28.3 ± 8.7 g/d) 81.0 21.9 79.4 113.0 Control Feces Mean, n = 13, µmol/g wet Acetate Propionate Butyrate Total SCFA Humans, 10 male, fed 4 days Humans, 3 male, 10 female, fed 9 days Ispagula increased fecal [Ac] and [Pr] Highly resistant starch treatment had higher excretion of Ac and Bu Loperamide → ↓ fecal [Pr] and [Bu] Senna → ↑ fecal [Ac], [Pr], and [Bu] 27 26 25 2387_ch4.9_fm Page 301 Sunday, May 6, 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 301 Cecal contents Mean pH, n = 6 0.5 mo 6 mo Cecal contents Rats, male, Wistar, 115 g, fed 0.5 or 6 mo Rats, male Wistar, 75 ± 5 g, fed 12 days Cecal contents Mean pH ± SEM, n = 10 Cecal contents Mean pH, n = 39 Rats, male Wistar, 200 g, fed 7 days Rats, male Wistar, 8 wk old, fed 9 mo Medium-Fat Low-GOS High-GOS 6.2 5.8** High-GOS 5.8** Medium-Fat Low-Cellulose High-Cellulose 6.6 6.4 5.99 ± 0.09* 6.58 ± 0.06* Res starch (20%) 5.61 ± 0.08* Low-Fat Low-GOS 6.3 Res starch (10%) Res starch (5%) Inulin (15%) 5.34 ± 0.08* Resistant starch from wheat 7.0 ± 0.1b Low-Fat Low-Cellulose High-Cellulose 6.5 6.5 Inulin (7.5%) 5.93 ± 0.13* 6.5 ± 0.1a Potato starch Resistant starch from potato 6.4 6.4 Wheat Bran (200 g/kg diet) 5.9 6 FOS (9 g/100 g) 6.6 Inulin (3.75%) 6.40 ± 0.11* 7.2 ± 0.0b 7.8 ± 0.1c Fiber-free 7.02 ± 0.08 Pea starch Control 7.5 7.5 Basal 6.1 5.9 Basal 7.5 Basal Source (and Level) of Fiber in Diet GOS → ↓ pH in cecum at all fat levels Cellulose did not influence cecal pH at any fat level ↑ Resistant starch → ↓ pH in cecum ↑ Inulin → ↓ pH in cecum Each starch → ↓ pH in cecum Resistant potato starch → ↓ pH in cecum FOS → may ↓ fecal pH at all timepoints Comments 29 4 13 2 28 1 Ref. 302 Mean pH ± SEM, n = 7 Cecal contents Mean pH, n = 4 4 h post-feeding 16 h post-feeding Mean pH, n = 6 Cecal contents Rats, male, fed 17 days Rats, male Wistar, 115 g, fed 27 wk Specimen/Data Influence of the Source of Dietary Fiber on Luminal and Fecal pH Animal Model Table 4.9.6 2387_ch4.9_fm Page 302 Sunday, May 6, 2001 7:31 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION, 3RD EDITION Rats, male Sprague-Dawley, 40–60 g, fed 38 wk 7.27 ± 0.05 Distal colon Luminal contents Mean pH ± SE, n = 5 Proximal colon Distal colon 6.98 ± 0.08a Proximal colon Pectin 6.67 6.81 Cellulose 6.66 6.85 7.03 ± 0.03 6.16 ± 0.03b 7.35 ± 0.05a Fiber-free Pectin (8% DF) 6.34 ± 0.05c Luminal contents Mean pH ± SEM, n = 10 Cecum Rats, male Sprague-Dawley, 200-220 g, fed 8 wk † 7.5b 7.7 7.7 7.5c Low-amylose and psyllium 7.3b 7.4 ± 0.1 7.0 ± 0.1 7.3b 7.05 ± 0.03a 7.22 ± 0.03a Distal colon Low-amylose 6.36 ± 0.06b 6.72 ± 0.03a Proximal colon Feces and colonic digesta Mean pH ± SEM Cecum Proximal colon Distal colon Feces 6.67 ± 0.06b 7.11 ± 0.08a Rats, male Sprague-Dawley, 208–234 g, fed 2 wk Pectin (6%DF) Pea 6.3 ± 0.1b Wheat bran 6.3 ± 0.1b Cellulose (6%DF) Carrot 6.3 ± 0.1b Inulin 5.9 ± 0.1a Luminal content Mean pH ± SEM, n = 19–20 Cecum † Rats, male Sprague-Dawley, 270–300 g, fed 4 wk Cecal contents Mean pH ± SEM, n = 6 Rats, male Fischer, 280 g, fed 8 wk High-Fat Low-Cellulose High-Cellulose 6.5 6.4 7.27 ± 0.05 6.55 ± 0.18a,b Wheat bran (8% DF) 6.74 ± 0.08b 6.9a 6.9 6.6 7.3b High-amylose 6.97 ± 0.11b 6.30 ± 0.04b 6.57 ± 0.06b Oat bran (6%DF) Oat 6.5 ± 0.1b Cocoa 6.5 ± 0.1b High-amylose and psyllium 7.0a 6.8 ± 0.2 6.3 ± 0.1 6.7a High-Fat Low-GOS High-GOS 6.2 5.8** Pectin → ↓ pH in cecum and prox. colon Wheat bran → ↓ pH in cecum only Psyllium → ↓ in fecal pH Cecal pH for pectin and oat bran < cellulose Prox. colon pH for pectin and oat bran < cellulose Distal colon pH for oat bran < pectin and cellulose Cecal pH was lowest for the inulin-fed group 11 9 21 8 14 2387_ch4.9_fm Page 303 Sunday, May 6, 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 303 65 7.48 6.43 6.14 ± 0. 304 † Pigs.09 Wheat Bran 6.91 6. fed 42 days † Luminal content Mean. 3RD EDITION .47 6.83 ± 0. n = 5 ~5 months old.09 Control 6.47 ± 0.Pigs.9.14 ± 0.28 Rolled oats 6.08 ± 0.82 ± 0.92 ± 0. 10 female.37 6. fed 34 days 6.27a 6.and distal colon Oat bran.77 Wheat flour + wheat bran 5.52 6. May 6. may lower pH at all three sites Cecal pH was lower for oat than corn bran at 0–4 hr Comments 25 12 12 21 Ref.85 ± 0.91 5.51 ± 0.88 ± 0.b Wheat flour Oat bran Cellulose 5.48 6.39 7. male Cecal digesta Hanford miniature.6 (Continued) Influence of the Source of Dietary Fiber on Luminal and Fecal pH Animal Model 2387_ch4.26 6.91 6.28 Senna may lower pH at all sites Wheat bran may lower pH in mid.48 6.23b Corn bran Source (and Level) of Fiber in Diet Rolled oats and oat bran 5.31±0.39 ± 0.91 6.45 6.99 ± 0.91 6.52 6.35 Control 6. n = 13 fed 9 days Mid-colon Distal colon Feces Pigs.52 Senna 6.59 ± 0.66 ± 0.80 ± 0. Mean pH ± SEM.30a. n = 4 Cecum Proximal colon Distal colon Mid-colon Distal colon Feces Humans. Feces and colon 3 male.77 Wheat flour 5.67 Wheat flour and oat bran 5.9_fm Page 304 Sunday. 0~4 h postprandial 40 kg 8~12 h postprandial Specimen/Data Table 4. Mean pH ± SD.78 5.04± 0.48 7.52 6. when added to wheat flour.73 6. 2001 7:31 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.70 ± 0. n = 4 Cecum Proximal colon Distal colon Luminal content Mean pH.71 6.97 6.66 5. 23 ± 3 yr old.37 7.46 Medium-fiber (30 g/d) 6.17 g/da Fresh 6. n = 18-22 Mean pH.82)a White vegetarian 29.05.89 ± 0. n = 14 Feces Mean pH ± SEM.6 ± 0.22 ± 0.70 ± 0.40–6. female Feces Mean pH ± SD.5 ± 0.23 High-fiber (42 g/d) 6. Significantly different from fiber-free group at p < 0.96 ± 0. male.11 ± 0.Feces Humans.18 (5. male.48 Fecal pH was lowest for Indian vegetarians ↑ Starch resistance → ↓ fecal pH Fiber appears to have a threshold effect on pH 31 26 24 30 2387_ch4.1b Maize porridge Indian vegetarians 16.65 (6. Note: GOS = galactooligosaccharides.05. fed 3 wk Mean pH.b.53 Different superscripts within row indicate significant differences between diets at p < 0. Significantly different at p < 0. FOS = fructooligosaccharides.9_fm Page 305 Sunday.01. n = 9 Humans.1b Loperamide 6. May 6. fed 4 wk Mid-colon Distal colon Feces GOS (15 g/d) 6.97 g/da 6.57 g/db Stale 5.25 6.89 ± 0. Data was extrapolated from the original graphs. n = 12 Humans.c 6.9 ± 0.91* FOS (15 g/d) 6. fed 3 days Feces Mean fiber intake ± SEM.46 7.3 ± 2.2 ± 1.45)b 6.91–6. Humans.39 7. 22 ± 1 yr old.15 ± 0.19 Control Low-fiber (16 g/d) 6. male.55 (6.42 † * ** a. 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 305 .6 ± 0.1a Control 6.83 ± 0.27–6.72 ± 0.91)a White omnivore 16.88 ± 0.7 Inulin (15 g/d) 6. (95% CI) 6. 9 ± 0. µmol Feces Mean.0 ± 2.3 5. produced more SCFA than others.5 4.6 ± 0. 24 h incubation Rats. male Sprague-Dawley.0 ± 0.6 1.2 0.20 ± 0.02 0.2 ± 1.8 2.23c Citrus pulp Ileal digesta from pigs consuming Oat bran Wheat bran 1617 3319a 49:05:56 Citrus pectin (5%) 10.2 0.4 2.92c 7.02c Citrus pectin 64:11:25 Corn fiber (5%) 1. Humamil. 306 Rats.4 ± 0.41b 0.97b 1.7 ± ± ± ± Leo 0. May 6.5 ± 0.3 5.02 0.03a 0.93b 0. n = 8 Total SCFA.37a Control Humamil 9.2 0.02 11. Comments 34 33 18 32 Ref. 6 wk old.4 0. 50 kg body wt.2 ± 0.7 2387_ch4.62b Fybogel 7.1 0.00a 0. n = 6 µmol/mg substrate Acetate Propionate Butyrate Total SCFA ± ± ± ± 0.2 0.8 ± 0.8 8.78c 6.48b 0.54b 4.3 0. 24 h incubation Inoculum Influence of the Source of Dietary Fiber on SCFA Production In Vitro Animal model Table 4. n = 5 Total SCFA. n = 6 mol/g substrate organic matter Acetate Propionate Butyrate Total SCFA Rats.9 2.1 0.0 ± 9.5 ± 0. female Sprague-Dawley.1 0.4 Metamucil Beet pulp 2. 2001 7:31 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. barrows.3 4.9 ± 2.46a Fibra 6.88 ± 0.6 ± 0.9.19c Fibraplan 6. male Wistar.5 5.5 0.97c 0.83 ± 0. 3RD EDITION .4 ± 0.04 0.1 0.3 ± 0.1 ± 0. Fybogel. and Fibraplan.0 ± 0.4 ± 0. 24 h incubation Pigs. 190 g.2 38:19:43 Indigestible dextrin (5%) 7.5 0.04 13.1 0.2 0.Cecal contents Mean.04 13.08 0.5 Agiolax Source (and Level) of Fiber in Diet ± ± ± ± 0.1 15.9 3.59 ± 0.2 ± 1. 24 h incubation † Molar ratio (Ac:Pr:Bu) Cecal contents Mean ± SEM.0 Fibra Kneipp Cellulose had the lowest SCFA production SCFA for oat bran > wheat bran SCFA for control and pectin > corn and dextrin % of the 3 SCFAs differed greatly among treatments Metamucil.2 0.5 7.6 6.3 4.9 ± 1.97c 1.03a 0.9_fm Page 306 Sunday.13c 1. which are mainly soluble fiber components.3 5. mM Acetate Propionate Butyrate Total SCFA Cecal content Mean ± SD.06a 3.5 Cenat Cellulose 15:27:58 10. 04b 0.b 0.d 6.2 7.b 0.45a 0.b Gum karaya 5.d:25e Wheat bran fiber 1. oat fiber.27a. mmol/l Acetate Molar percentage (Ac/Pr/Bu) Feces Mean ± SEM.77a 62a.68c.b:21c.86 0.9_fm Page 307 Sunday.45 5.b 0. 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 307 .c 78c:16b.b 61a. sugar beet fiber.90d.99a.57 Psyllium 12.6 ± 2.b 3. May 6.e 0.c.Humans.02a 0.77 1.09a 0.6 20 Williamson oat fiber 17.48a 0.16a. mmol/g ± ± ± ± 0.49 0.32b 0.23a.b:22d:16c.12 1.d:17c.52b. 28-42 yr old.03a 0.59b.16a.7 68/16/13 Pea fiber Apple fiber ± ± ± ± 0.c:7a 0.8 12. and xanthan gum.05c 0.71a 62a.48a 56a.24a.21b 0.d Corn bran 11. n = 3 mmol/g original org.7 75.22 1. 24 h incubation Molar ratio (Ac:Pr:Bu) Acetate Propionate Butyrate Total SCFA Feces Mean.86d 0.23 8.2 48 Pea Production of SCFA and ratios among the 3 major SCFAs were significantly influenced by the fiber source [Bu] wheat bran and sugar beet > maize and pea [Ac] and [Pr] were highest for sugar beet Mixed fiber contains pea fiber.b:13b:26e 33.b:19c.28d. Total SCFA production for β-glucan > psyllium > mixed fiber SCFA production from apple fiber > pea fiber 38 37 36 35 2387_ch4.89 0. 24 h incubation Humans.c 0.50 Mixed fiber 66/18/11 54.6 18 88 5.05b 0.03c 0. n = 2.05a 0.2 ± 5.c 1.12a 0.24 44 0.d 0. n = 4 Total SCFA.d Xanthan gum 3.b.60 4.11a 0.91b.28a 0.e 0.10a 79c:12b:9a. n = 12 mmol/g organic matter Acetate Propionate Butyrate Total SCFA Feces Mean ± SEM. 24 h incubation † Humans. matter Acetate Propionate Butyrate Total SCFA Molar ratio (Ac:Pr:Bu) Propionate Butyrate Feces Mean.05c Sugar beet ± ± ± ± β-Glucan Wheat bran 2.27b Canadian Harvest oat fiber 0.3 24 Maize 3.13a 0. 24 h incubation Humans.03c 0. 78c.d 1.32f 0.d Gum arabic 6.01 1.66i 2.08 0.99j Pea 0.08b. May 6.11 ± 0.42 Fig Canadian Harvest oat fiber Acetate Propionate Butyrate Total SCFA Molar ratio (Ac:Pr:Bu) Williamson oat fiber Citrus pectin 5. 3RD EDITION .55 ± 0.e 4.37d 77c:6a:17c.45a-d 1.d 2.22 ± 0.11 0.16 0.55i Soy fibrium 5.08 1.09e.24e 1.38 1.27 0.26 0.c 0.46 ± 0.19 ± 0.9.82c.64e 66b:21c.41 Carboxymethylcellulose Gum arabic and soy fiber produced the highest amounts of SCFA Comments 40 39 Ref. 2001 7:31 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.d 62a.93 8.d:18c.e 0.d:13a.9_fm Page 308 Sunday.95d. 24 h incubation Feces Mean ± SEM.19 Gum arabic Guar gum 3.e.b. 35–61 yr old.b:20c.85d 53a:33e:14b.52 0.72 0.d:22d.09 0.d.f 6.18 3.22f 7.87 0.02f 1.71 1.c Acetate Propionate Butyrate Total SCFA Molar ratio (Ac:Pr:Bu) Feces Mean.7 (Continued) Influence of the Source of Dietary Fiber on SCFA Production In Vitro 2387_ch4.42 ± 0.e Inoculum 1.24f 9.92 5. 308 Humans.38 ± 0.c 59a.06 0. n = 6 mmol/g of dietary fiber Acetate Propionate Butyrate Total SCFA Acetate Propionate Butyrate Total SCFA 1.4b. 4 male.87 0. 3 male.04 ± 0. 25–41 yr old.01 2. 24 h incubation Animal model Table 4.01 0.01 0.10 0.96 ± 0.05 0.f 6.05 0.59c.05 0.b: 19c.00 Soy fiber 3.e 0.14 ± 0.01 0.89d.92h Psyllium 2.c Source (and Level) of Fiber in Diet Oat bran fiber 2.33 ± 0.19 0.d 1.12 0.75 Oat 0.Humans.32 ± 0.01 2.7d.38e 2.07 0.002 2.14c. n = 6 mmol/g dry substrate matter Acetate Propionate Butyrate Total SCFA Soy fiber 3. 44 ± 0.01a 0.11 ± 0.66 ± 0.05 3.05 0.b 0. 35–61 yr old.79b 1.002 0.21 ± 0.54* β-glucan Pear 0.01a –0.07 0.23 ± 0.01a 0.41 ± 0.81c Sugarbeet fiber –0.41 ± 0.76 ± 0.02a 0.24 ± 0.003 1.36 ± 0. Mean.48c 0.b 0.95m Ac.72b Pectin 5. n = 6 mmol/g polysaccharide Acetate Propionate Butyrate Total SCFA Humans. and Bu production for citrus pectin > pea and sugarbeet There were significant differences among substrates for production of individual and total SCFA.01a 0. 3 donors.01 0.83b 0.65b 0. but differences not significant In comparison to uncooked (whole) starch.04a Soy fiber Methycellulose 1500 1.06 1.04 1.9_fm Page 309 Sunday.01 1.02 6.02 1.09l 2387_ch4.01 0.06 ± 0. total SCFA production was lower for resistant starch and higher for pectin and β-glucan Viscosity for methylcellulose 15 > 1500 > 4000 There were significant differences among substrates for total SCFA production. 24 h incubation Acetate Propionate Butyrate Total SCFA 1074b 77a 9a 1385b 400b 158b.00a 0.16 ± 0. n = 9 24 h incubation mmol/g organic matter Acetate Propionate Butyrate Total SCFA Humans.01a –0.20 0.51* Pectin Wheat 0.06b –0.09 ± 0.00 ± 0.01 0.53d 7.13l Starch (whole) Apple 1.b Pea fiber Solka Floc ® 0. and it appeared that differences were present also for each of the the 3 major SCFAs 42 41 40 THE SOURCE OF DIETARY FIBER INFLUENCES 564a 127a 70a. Feces 24 h incubation Mean.69 ± 0.01 1. Pr. 43 yr old.02a.04 0.39 ± 0.42* 2.30 ± 0.96 ± 0.01 0.03a.03 0.c Psyllium 0. 4 male.01 0. 2001 7:31 PM 309 . production was highest for pectin Negative values indicate that SCFA production was lower than in controls.002 1.02 0.02 0.01 5.24c 1.18 ± 0.89 ± 0.Feces Mean ± SEM.76 ± 0.02 4.04a –0. May 6.57 ± 0. n = 3.56k 5765c 453b 182c Citrus pectin 2.08a 0.00a 0. Feces av.03a 0.00a Methycellulose 4000 5.35 Methycellulose 15 Resistant Starch Corn 0.62 ± 0. µmol/g substrate Acetate Propionate Butyrate Acetate Propionate Butyrate Total SCFA Humans. 3 donors.05 ± 0. 01* Feces Mean ± SE.3 ± 0. All treatments (except FOS) → ↑ Pr prod.6* 21.2* 14.7 ± 0.9_fm Page 310 Sunday. and inulin Comments 44 43 42 Ref.3 5620b 793b 472b Apple pectin Source (and Level) of Fiber in Diet Psyllium Husk 12.2 ± 2.5 ± 0.4 ± 0.1* Acetate Propionate Corn Fiber Fractions Total dietary fiber residue 1. n = 2.7 Methyl Cellulose 22. > TDF fraction All treatments → ↓ Ac prod.1 ± 0.5 43. 310 Human. 3 male. increased by soy.3 ± 0.8 ± 0.60a 2.3* 11.6 ± 0.5* 41.7 ± 1.2* Butyrate Feces Mean.2* 9.9 ± 0.03* 54.02 38.4 ± 0.8* 10. 2001 7:31 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.01 8.3* 16.7 6.4 Acetate Propionate Butyrate Total SCFA Cellulose 30.7 Hydrolyzed Inulin 21. 24 h incubation Humans.5* 43. guar.1 44.74 11.5 ± 0.7 (Continued) Influence of the Source of Dietary Fiber on SCFA Production In Vitro 2387_ch4.9 ± 0.75e 0.6 ± 0.8 ± 0.24d.1* 10.0 ± 0.58a 0.3* 14.9* 11. mg/ml Acetate Propionate Butyrate Total SCFA Oat fiber Gum arabic Cell wall polysaccharides 5. 28–40 yr old.1 33.5 ± 1. psyllium.5* 3.3 ± 1. mmol/g substrate Acetate Propionate Butyrate Inulin 19.8 ± 1.9 Hydrolyzed Guar Gum 19.5 ± 0.38 41.0 ± 1. 48 hr incubation 44.4 ± 0.Inoculum Human.0 ± 0.2 ± 0.8 ± 0. µmol/g substrate Acetate Propionate Butyrate Animal model Table 4.4 ± 0.21 Total SCFA Original material 2.e 0. 3RD EDITION .56g 0.6 ± 0.6 ± 0. May 6.50g 0.2* 7.3 ± 0.7 ± 1.8 ± 0.9.4* 13. Feces 24 h incubation Mean. n = 3. n = 6.2 Fructooligosaccharide 22.4 ± 0.1 4.2* 12.1 Soy Oligosaccharide 17.2* 13.0 ± 0. Bu prod.1 ± 0.5 ± 0.6 Glucose 68a 55a 10a 4593b 1758c 585b Powdered Cellulose 11.7* 401a 188a 47a Corn fiber Ac and Pr from cell wall polysacc.6 ± 2.6 8.4 38.7 ± 0.3* 15.7 ± 0. 3 donors. 3 donors.4* 19.4 ± 0. 03a 0.00a Himanthalia elongata 39.32b Citrus pulp Cell wall polysaccharides 4.43b 1.02a 0.28d.17b.6 ± 1. 32 ± 5 yr old.d 0.65b 0.c 75/14/9 2.27b 1.1a 66/19/12 Undaria pinnatifida 46.5d — 10.85 Feces Mean ± SE.f 0.80b.6b 80/14/5 Total SCFA.43b –0.60b.c Citrus pectin In comparison to the nofiber treatment. May 6. mmol/l Molar % (Ac/Pr/Bu) No fiber 2.85c. 24 h incubation Acetate Propionate Butyrate Acetate Propionate Butyrate Original material Sugar beet 73.Fucans 11. mmol/l Molar % (Ac/Pr/Bu) Feces Mean.6 ± 1.6c 73/15/10 Humans.6 ± 3.04b 0. 24 h incubation Beet pulp Laminarans 87.3 ± 3. 3 male.96 Original material 4.84 Wheat bran fractions Total dietary fiber residue 2.c 1.68b 1. n = 6 mmol/g substrate org matter Acetate Propionate Butyrate Total SCFA Humans. n = 2–7 Total SCFA. all fibers except for fucans produced more total SCFA SCFA prod.93a.9_fm Page 311 Sunday.15d. > TDF fraction Pr from Cell wall polysacc.04b 0.e 0.c 1. > TDF fraction 45 34 2387_ch4.97b 0. 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 311 .g 0.96d 4.72 Cellulose 2.27c.c 1.00a 0.e 0.91 Oat bran fractions Total dietary fiber residue 3.58f.2 ± 2.0b 69/19/9 2.d 0.7d — Alginates 54.7b.5 ± 0.61c 4.b 1.43 Cell wall polysaccharides 4.6a 60/23/16 Laminaria digitata 54.81b. for cellulose < other treatments Ac and Pr from cell wall polysacc. 2 donors.42b 3.4 ± 1.05b 0.91e. d 6.42 ± 0.b 0. 2001 7:31 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.20b 6.78 ± 0.9_fm Page 312 Sunday.82b 1.23 ± 0.46 ± 0. 3RD EDITION .04b 3.03 ± 0. 1.e.f.17a. for oat and wheat > corn and rice Comments 46 Ref.01 ± 0.i.92b 3. n = 6 mmol/g organic matter Acetate Inoculum SCFA production vegetables > corn and rice Ac production for oat and wheat > corn and rice Pr production for oat and wheat > corn Bu production for oat and wheat > corn and rice Total SCFA prod. * Significant difference from control treatment.84b 1.d.l..87b 1.07c 1.49b 1.53b 1.55 ± 0.37 ± 0.15 ± 0.21a Rice Solka Floc®.82 ± 1.b.28c. registered trademark of Fiber Sales and Development Corp.78 ± 0.m Different superscripts within row (or within individual or for Total SCFA) indicate significant difference between diets at p < 0. Louis.k.15a.09c.38 ± 0.29b Butyrate Total SCFA Acetate Propionate Butyrate Total SCFA Carrot 4.c.14 ± 0.b Lettuce 4.1.23a.53 ± 0.60 ± 0.02 ± 0. † h.06a.45a 0.22b 1.03b 2.c.21b 1.7 (Continued) Influence of the Source of Dietary Fiber on SCFA Production In Vitro 2387_ch4. between diets at p < 0.09b 1.27 ± 1.20 ± 0.05.d 6.j.21a Wheat a.d 6. Oat Source (and Level) of Fiber in Diet Corn Feces Mean ± SEM.45 ± 0.12b 1.45 ± 0.g 3.59b 0.22 ± 0.58 ± 0. St.01.10 ± 0.89 ± 0.75 ± 0.11b Data were extrapolated from the original graphs.63 ± 0. 3 donors.07b 1.c.10a.90 ± 0. MO.9. 312 Note: Humans.31 ± 0.08b 0. Different superscripts within row indicate significant differences.24 ± 0.28a Broccoli 4.42 ± 0. May 6.33b Cabbage 4.d 7.21 ± 0. 24 h incubation Animal model Table 4.d 6.09a Propionate 1.01 ± 0. 6..9_fm Page 313 Sunday. Enhancement of butyrate production in the rat caecocolonic tract by long-term ingestion of resistant potato starch.. including differences in residence time. Prolonged intake of fructo-oligosaccharides induces a short-term elevation of lactic acid-producing bacteria and a persistent increase in cecal butyrate in rats. Zoran. et al. While a standard in vitro system is useful to determine SCFA production under defined and rigorously controlled conditions. fibers derived from citrus or sugar beets produced more SCFA in vitro than did cellulose or corn bran. B. 2231. SCFA absorption will increase along with increased SCFA production.. 2217. A.. P. 1997. and Eastwood. or wheat. 33. 1999. Edward. Gut. Am. 127. M. M. Fermentable carbohydrate exerts a urea-lowering effect in normal and nephrectomized rats. 3. 1997.. SCFA production appears to be higher for water-dispersible sources of dietary fiber than for fibers composed of a heterogeneous mix of water-dispersible and -indispersible compounds. 1994.38.. A. REFERENCES 1. J.2387_ch4. J. Nutr. 1999.. H. et al.42 Also. C. 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 313 In vitro. Wheat bran diet reduces tumor incidence in a rat model of colon cancer independent of effects on distal luminal butyrate concentrations. and prevention of cancer development. 1522. Nutr. and Lupton. As more information becomes available. 7. Balay. G515. luminal SCFA concentrations will be attenuated by absorption. 419.. Nutr. . 33. Younes. et al. LeBlay. Physiol. changes in luminal concentration will almost certainly underestimate the effect of the dietary component on SCFA production in vivo. Dietary fibers stimulate colonic cell proliferation by different mechanisms at different sites. J. D. Fredstrom.42 Results with three different types of cereal suggest that SCFA production arises primarily from the cell wall polysaccharides of the brans rather than from the total dietary fiber residue. etc. a range of plant gums appeared to produce more SCFA in vitro than did cereal fibers derived from oats.44 In a separate study. 8. 22. Wilson. G.. 5.. et al. J. 272.. A. Comparison of the effects of ispaghula and wheat bran on rat caecal and colonic fermentation. G. C. 267..38. 1076. 4. R.39. 2. Zhang.34. and luminal pH. Since SCFA transport from the lumen to the mesenteric and portal blood supply is strongly dependent on luminal SCFA concentration. it appears that foods rich in water-dispersible cell wall polysaccharides support greater SCFA production than do foods rich in cellulose.. Taken together. S. 9. and Kurtz. Gut. Apparent fiber digestibility and fecal short-chain fatty acid concentrations with ingestion of two types of dietary fiber. corn. 129.. 1992. J. 14. 1229. L. Relationship of colonic luminal short-chain fatty acids and pH to in vivo cell proliferation in rats. L. L. Nutr. et al. Edwards. Lupton. 1992. G. 123. J..18. R. J. or other water-indispersible but structurally important components. A.41. and Eastwood. 82. Nutr. 18. vegetables were shown to support higher SCFA production in vitro than cereals. Hanlon. Effect of the dietary fibre content of lifelong diet on colonic cellular proliferation in the rat. Cancer. this approach is unable to consider fully aspects of the physiological situation. P. Thus.. J. Much interest has been focused on the physiological roles of SCFA for the intestine and also for other tissues and cells. differences among fibers will be able to be more effectively exploited to optimize health and prevent disease. May 6. SUMMARY It is now evident that the quantity and source of fiber affect SCFA production and absorption. 1993.. identifying the precise mechanism by which dietary fiber and SCFA exert their physiological effects. 1994. Given these kinetics. R. lignin. interaction effects with other luminal constituents. JPEN. More specifically. Br. The importance of fiber in the diet evolved from viewing this as a nonnutritional and noncaloric substance to valuing dietary fiber as an important substance able to effect the health of the intestine as well as lipid metabolism. carbohydrate metabolism. J. May 6. and Tagny. The influence of dietary fiber on proliferation of intestinal mucosal cells in miniture swine may not be mediated primarily by fermentation. Morita. P. and Takeda. 1994. A. Kishimoto. 1521. Clin. M.. short chain fatty acids.. 14. Nutr. Gastrointestinal implications in pigs of wheat and oat fractions. and DeVries. Bjorck. 53. 1994. J. 1991. Christensen. 33. 2000. McIntosh. 13. E. and Madar. 128. and Royle.. et al... J. J. W. Am. S. Kim.. I. J. blood lipid concentrations and glucose absorption in young healthy male subjects. P. 241. Integration of ileum cannulated pigs and in vitro fermentation to quantify the effect of diet composition on the amount of short-chain fatty acids available from fermentation in the large intestine. Jenkins. Wakabayashi. et al. 123. 245. Eur. S. 130.. Nutr. Fleming. Psyllium shifts the fermentation site of high-amylose cornstarch toward the distal colon and increases fecal butyrate concentration in rats. A. A... Goni. Nutr. Effect of nondigestible oligosaccharides on large-bowel functions. 17. J. Nutr. G. Roland. A comparison of the effects of dietary cellulose and fermentable galactooligosaccharide. 1999. Nutr. J. Dokkum. A. K. 615. 20. Monsma. blood lipids and glycemic index. 18. 1731. The water-soluble extract of chicory influences serum and liver lipid concentrations.. 71. 804. Y. J. 1998. Sci. J. 129.. J. 25. N. et al. 12. . J. Nutr. 906. K. 755. Br. 41. 26. Br. Stark.. 585.. J. S. et al. A. Nutr. J. et al. S. L. cecal short-chain fatty acid concentrations and fecal lipid excretion in rats.. Carcinogenesis. H. Diet and carcinogen alter luminal butyrate concentration and intracellular pH in isolated rat colonocytes.. 233. Hypocholesterolemic effect of dietary fiber: relation to intestinal fermentation and bile acid excretion. Nutr. I. 65. Wang Y. 24. E. Evaluation of fermentability of acid-treated maize husk by rat caecal bacteria in vivo and in vitro. V. et al. S.2387_ch4. V. J. 22. Faecal pH. M. 21. Br. 1747. Fermentation of dietary starch in humans. Gut. H. J. Physiological effects of resistant starches on fecal bulk. K... D.. N. Gut. Folino. Nutr. et al. et al.. 1999..... J.. D. Nutr. Br. M. A diet containing chickpeas and wheat offers less protection against colon tumors than a casein and wheat diet in dimethylhydrazine-treated rats. Nutr.. and Nyman. Zoran. 30. 71. Hara. Gastroenterology. Diurnal changes in large-bowel metabolism: short-chain fatty acids and transit time in rats fed on wheat bran. C. D. 18. 651. 74. 128. Fitch. Sci. 1998. 31. 2000. et al. 1077. M. Res. 1997. Haack. H. Am. Br. 1998. 11. Comparative study of the fermentative characteristics of inulin and different types of fibre in rats inoculated with a human whole faecal flora. 35. 23. 16.. 1995. 20. 68. Ahmed. J.. In vitro fermentation and hydration properties of commercial dietary fiber-rich supplements. J. and Shin. 609.. 2001 7:31 PM 314 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Nutr. 68. M.. and Young G. 2166. 719. 32. et al. D.. et al. 1998. Wijnands. 495. et al. 1992. Food Agric. H. S. 209.. Sci. J. 3RD EDITION 10.. J. Nutr.... Berggren. 122. Vitaminol. Nutr.. L. G. M. et al. 95. 27. 29. 1993. J.. 27. 1995. 151.. 1998. 1999.. In vitro fermentation of swine ileal digesta containing oat bran dietary fiber by rat cecal inocula adapted to the test fiber increases propionate production but fermentation of wheat bran ileal digesta does not produce more butyrate. 1999. E. W. H. 222. 1995. 125. Reddy.. Nutr. 2081. Lewis. Food Agric. et al.. Nutr. F. V. Mathers. D... 1998. R.. bile acid and sterol concentrations in premenopausal Indian and white vegetarians compared with white omnivores. Am. 79. 15. J. Z. Nutr. Short-chain fatty acid content and pH in caecum of rats fed various sources of starch. 17. 79.9_fm Page 314 Sunday.. In vitro production of short-chain fatty acids by bacterial fermentation of dietary fiber compared with effects of those fibers on hepatic sterol synthesis in rats. J. 239. 1995. Marteau... 1997.... 1994. 28. M. Increasing amounts of dietary fiber provided by foods normalizes physiological response of the large bowel without altering calcium balance or fecal steroid excretion. H. and Heaton. 41. P. M. Dietary fibers differ in their effects on large bowel epithelial proliferation and fecal fermentation-dependent events in rats. E. Increasing butyrate concentration in the distal colon by accelerating intestinal transit. Nutr. 1. Digestibility and bulking effect of ispaghula husks in healthy humans. T. J. Clin. 1017.. J. Bachknudsen. in a rat model of colorectal carcinogenesis: fermentable fibre confers greater protection than non-fermentable fibre in both high and low fat backgrounds. Cancer. 19. Coll. J. Velazquez. Boca Raton. J. J. 521. 2463. Sunvold.. in CRC Handbook of Dietary Fiber in Human Nutrition. Anaerobe. 36. 122. and potential water-holding capacity of various substrates. 1995. C. 43. et al. A. 35.. Food Agric. E. L. 70. 1508. Y. Sci. 16.. 38.. M.. J. Food Chem. U. 46. et al. 1997. E. 47. In vitro fermentation of cellulose. 14. Sci. Scand.9_fm Page 315 Sunday. 44. et al.. 73. and Thompson. horses. citrus pulp. 3639. Vanderhoof. 1993. Br. L.. J. Gastroenterology. I. J. In vitro fermentation of various food fiber fraction. L. 6. 2001 7:31 PM THE SOURCE OF DIETARY FIBER INFLUENCES 315 34. Guillon. Nutr.. Nutr. 1119. Res.. Nutr. 1993.. Oles. 87. 37. Jr. In vitro fermentation by human faecal bacteria of total and purified dietary fibres from brown seaweeds. L. Casterline. Bourquin.. Fermentation of various dietary fiber sources by human fecal bacteria. J. 2nd ed. Short-chain fatty acid production and fiber degradation by human colonic bacteria: effects of substrate and cell wall fractionation procedures. et al. Nutr. Fermentability of various fiber sources by human fecal bacteria in vitro. Psyllium and methylcellulose fermentation properties in relation to insoluble and soluble fiber standards. I. et al. 48. 2000. . or excretion of short chain fatty acids in humans. 1993. 45. 41. 421.. humans. J. Fleming. beet pulp. Nutr.. J.. D.. D. 7.. absorption. 619. 1995. Clin. L. J. G. et al. 17. Immunonutrition: the role of carbohydrates. 53. Potential water-holding capacity and short-chain fatty acid production from purified fiber sources in a fecal incubation system. Characterisation of residual fibres from fermentation of pea and apple fibres by human faecal bacteria. M. Br.2387_ch4. 263. A.. Campbell. Nutr.. May 6. 189. Bourquin. Spiller. Res. 40. and pigs and ruminal fluid from cattle. F. M. et al. et al. Am. 75. FL.. M.. 249. 68. CRC Press. 1997. V. Titgemeyer. Ed. Anim. 387. 1991. D. Michel. C. J. Effect of oligosaccharides and fibre substitutes on short-chain fatty acid production by human faecal microflora. Agric. 1418. Nutr. 28. 13.. Influence of dietary fiber on the production. 271. 45. 42. S. C. Cancer. J. 1996. Fermentative characteristics of cereal brans and vegetable fibers. G. Nutr. 39. McBurney. and citrus pectin using fecal inoculum from cats. dogs. D.. and Ku.. 1992.... Nutr.. 1991. 1996.. 1998. Bourquin. Salvador.. et al. short-chain fatty acid production from.. Fermentation of dietary fibre by human colonic bacteria: disappearance of. Sugar composition of dietary fibre and short-chain fatty acid production during in vitro fermentation by human bacteria. McBurney.. et al.. 1990. 595. 2001 7:31 PM .9_fm Page 316 Sunday.2387_ch4. May 6. these are based on a qualitative relationship between carcinogenicity and DNA damage. Using a different approach. urban blacks. methods employed to measure mutagenic activity have involved bacterial mutation assays. the precise nature of the compound or compounds in feces causing bacterial mutations was not known. Generally. poultry. It was noted that the chromosome aberrations seen were comparable to those in mammalian cells exposed to 250 to 400 rads of Xrays as well as to several known carcinogens. and this activity may be modulated by the diet. and fiber but less animal fat than New Yorkers.e.7 Initially. Canada.00+$1. or meat) with a low colon cancer risk. it was suggested that bacterial flora are involved in the formation of this mutagenic activity.2387_ch4. the presence of mutagenic activity was shown in feces from healthy human male volunteers consuming a “Western diet. Finland with a low risk. For example. No Adventist sample was mutagenic. In Vancouver. had a significantly higher percentage of mutagen-positive stools compared to two low-risk black populations. Freeman Human feces contain mutagens and have been assessed by a variety of short-term in vitro methods. genotoxic effects of human feces were demonstrated using induction of chromosome damage in cultured Chinese hamster ovary cells as the indicator of mutagenic activity.9 however.50 © 2001 by CRC Press LLC 317 . dairy products. Later.10 Effects of Dietary Fiber on Fecal and Intestinal Luminal Mutagens Hugh J. a high-risk group.17 Because this bacterial assay method is extremely sensitive compared to the method 0-8493-2387-8/01/$0. urban whites. however. fecal mutagens were determined in strict vegetarians. and rural blacks) with differing colon cancer risks. and nonvegetarians16 using the fluctuation test for weak mutagens. 2001 6:51 PM CHAPTER 4.14 Others15 examined fecal mutagens in three groups: Seventh-Day Adventists living in New York consuming an ovo-lacto vegetarian diet (milk and milk products but no fish.10_fm Page 317 Sunday.1–6 Fecal mutagenic activity has been postulated to influence colon cancer risk. fecal mutagens were defined in a single stool sample from each individual in groups of South Africans (i. inhabitants of Kuopio. They are sensitive and rapidly performed.”1 It was subsequently claimed that stool samples contain both volatile and nonvolatile N-nitroso compounds8 as well as nitrite and nitrates.11 Subsequent studies focused on changes induced in mutagen levels of feces by altered diets or lifestyles.12 Later. May 6. ovo-lacto vegetarians.10 Later.. a high colon cancer risk population. ingestion of antioxidants such as ascorbic acid or tocopherol caused a reduction in fecal mutagenic activity followed by a gradual return to control values over a 1-month period. and omnivorous New Yorkers.13 Urban whites. positive activity was observed in 13% of Finnish samples compared to 22% of New Yorkers. fecal extracts from both carnivorous and herbivorous animals were demonstrated11 with high chromosome damaging potency observed in three carnivorous species examined. The Finns consumed more milk. Subsequently. distal small bowel. a potent mutagen present in human feces related to fecapentaene fiber adsorption.5 or 9%) or a mixture of cellulose and pectin (4. such as pure cellulose.2387_ch4. More recent studies have shown that fecal mutagen levels are lower in a rural low-risk colon cancer population (i. Although no significant change was seen in the high-fat group. Previous studies using chemically induced animal models demonstrate protective effects for some dietary fibers in rats exposed to exogenously administered chemical carcinogens (see Chapter 5.26 Extracts were obtained from fecal pellets and the luminal contents along the length of the GI tract in rats fed either a chemically defined fiber-free diet or nutritionally and calorically equivalent diets with differing amounts of purified cellulose or pectin (4. and colon after 8 weeks on the fiber-free diet revealed that the distal small bowel had a several-fold-higher mutagen concentration than any other site. they are presented to the cecum and colon in high concentrations. Over a 6-week period only the highcellulose diet influenced fecal mutagen levels.10_fm Page 318 Sunday. a shift from a well-balanced mixed diet to a lacto-vegetarian diet reduced direct-acting mutagen activities in human urine and feces.20 Multiple types of mutagens may be present. low-fiber “mixed” Western diet.22 Human studies have demonstrated that only specific sources of ingested dietary fibers. both wheat bran fiber supplementation and phytic acid (inositol hexaphosphate) resulted in decreased formation of aberrant crypt foci. and the presence of at least two fecal mutagens was suggested. there were significant differences between subjects on identical diets. followed by 2 weeks on a high-risk beef diet. Later. as shown in a New York population consuming a high-fat. Surrogate endpoint biomarkers for colon cancer risk in both rats and humans have been evaluated.5% each)..23 In addition.34. possibly related to higher water content in feces from dilution of fecal mutagens. These have revealed that different fiber sources including corn bran. Mutagenicity was measured using the fluctuation test for weak mutagens17 with Salmonella typhimurium TA 98 and TA 1535 as tester strains. Finland). Kuopio. Analysis of the contents of the stomach. Fecal mutagenic activities were determined after a period of supplementation with high fat and high fiber. Finland) compared to an urban high-risk colon cancer population (i.27–30 Alternatively. significant reduction was observed with the TA 1535 strain. however.18 The effects of short-term dietary modification on fecal mutagens were also assessed:9 two dietary regimes were used.34 In these studies.25 Studies specifically focused on the effect of dietary fibers per se are limited. such as wheat bran or cellulose. lignin. in experimental colon cancer31 as well as in humans on highfiber diets.e. high-fiber feeding reduced fecal mutagenic activities.12–15 virtually all samples showed mutagenic activity. some recent studies have explored the effects of luminal mutagenic activities on the colonic mucosa per se. a low colon cancer risk non-meat diet was consumed for 2 weeks.e.24 Finally. in vitro studies have shown a dietary fiber–induced reduction in fecapentaene excretion. it was shown that formula-fed individuals with consistent nutrient levels had fecal mutagenic activities that did not vary significantly within an individual. Some studies have explored possible mechanisms for these dietary effects on mutagens. wheat bran. Ovo-lacto vegetarians and strict vegetarians had significantly lower levels of fecal mutagens than nonvegetarians. Within a 2-week period. reduce the production and/or excretion of fecal mutagens. have been shown to alter colonic bacterial enzyme activities important in mutagen metabolism.32 This differed from effects of the purified fiber.. These activities were significantly reduced by ingestion of either the cellulose or pectin single-fiber diet. that did not lead to reduced levels of colon cancer in chemically treated rats. and alfalfa directly bind carcinogens in a pH-dependent manner. 2001 6:51 PM 318 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.33 Finally. May 6. 3RD EDITION used in earlier studies.1. cecum. This study further extended these observations to endogenous mutagenic activities and provided strong evidence that these mutagens are formed within the small bowel. optimally in the pH range of 4 to 6 found in the human GI tract and apparently through a mechanism of cation exchange.9). suggesting that long-term dietary and possibly genetic factors influence mutagen levels.35 . measured with the fluctuation assay for weak mutagens. Helsinki. mutagen levels increased on the highmeat diet. specific types of dietary fibers. D. A. 511.. Walker. 199. Potter. Fecal mutagens in high. T. Int. R. Baltimore. 113. V. Bosworth.. 72. Laakso. and Engle. Stich.. Cancer. 23. 2001 6:51 PM EFFECTS OF DIETARY FIBER ON FECAL AND INTESTINAL LUMINAL MUTAGENS 319 REFERENCES 1. and Williamson. A. and Yamasaki. Cold Spring Harbor Conferences on Cell Proliferation.. Eds. Kuhnlein. in Colonic Carcinogenesis. 1985.. and neutral sterols in healthy subjects. 280. A.. Pilot study of the effect of diet on the mutagenicity of human faeces. Varghese. B. K.. 1978. J. 353... and Wilkins. Ames. Stich.. H. 15.. 33. L. Choi. and Bell. Characterization of a mutagenic bacterial product in human feces. Bergstrom.. J... 78. 284. A. Nutr. D. Reddy. 4. H. J. Eds. and Acton.. C.. T. H.. 24. Venitt. 79... W. P. Erhrich. May 6... Mutat. R. A.. and Reddy. J. W. D. R. Bartram. 1. 119. S. 9.. 1. K. N. 21. A. University Park Press. 17. Mutagens in feces from vegetarians and nonvegetarians. Mutat. 18. E. 1981. A. 16. Cancer Lett.. Tannenbaum. Volatile nitrosamines in normal human feces. Clin. P. M. Kakizoe. L... . Lederman. and Wynder. 1979. H. C. MTP Press. R. NY.. R. C.. 1975. K. Simi. Cancer. Wei. Mutagenesis. Bruce. 200. V. 12.. Laakso. 2.. P. Fett. Kuhnlein. 2513. W. 1641. Nitrite and nitrate are formed by endogenous synthesis in human intestine.. C. 1983. E. Mutat. R.. Am. U. S. L.. Engle. J. van Tassell. Finland.. Mutagens in feces of 3 South African populations at different levels of risk for colon cancer. 113.. Katsifis. and Wilkins.. de Vet. C. 1981. Reddy. H.. vol. 6. Young. Rep. Biochemical epidemiology of colon cancer: effect of types of dietary fiber on fecal mutagens. M. I. 64. T. and Muriel. Sharma. P. 20.. B. D. Diet and DNA-modifying activity in human fecal extracts. 23. J. W. L. Nature (London). Varghese. Mutat. 4629. R. 347. H. 38. and Aldrick. V.. J... A mutagen in feces of normal humans. 1487. 115.and low-risk populations for colon cancer. and Henion. 1989. Effect of dietary fried meat on fecal mutagenic and co-mutagenic activity in humans. Kuhnlein. R.. Reddy.. S. and Weller. H.. 1981. Green. and Lam. P. C... Wilkins. Mutat. Furrer. R. Mutat.. West.. D.. F. Book C.... Puska. R. 85. P.. Methods for detecting carcinogens and mutagens with the Salmonella/mammalian microsome mutagenicity test.. 231. Res.. The endogenous production of nitroso compounds in the colon and cancer at that site. J. B. P. J. Metabolic epidemiology of large bowel cancer. 1979. Cold Spring Harbor.. C. and Kuhnlein. Wang. Nutr. L. B. Res. C. L. Res. Mutagens in feces from subjects on controlled formula diets.. 3. 8. Mutat.. and Kuhnlein. Venitt.. 1977. and Bruce... S. N. S. E.. Mutat. 10. 1979. T. 13.. J... B. Chromosome breaking activity of human feces and its enhancement by transition metals... Cancer Res. 22. Miller.2387_ch4. 49. in Naturally Occurring Carcinogens — Mutagens and Modulators of Carcinogenesis..... S. C. H. C. Aswell. 135.. L. Richardson.. F. S. A. 105. 1980... U. L. 11. Res. 1978. Nutr. D. 1980. 97. R. Faecal mutagens in the aetiology of colonic cancer.. E. 5.. Falk Symposium 31. Mutagenicity of fecal extracts from carnivorous and herbivorous animals. T. 1982. Engle. Cold Spring Harbor Laboratory.... Science. D. 2. Mutat. R. J. Rep. 59. L. U. Fecal mutagens from subjects consuming a mixed-Western diet... A. G. 276. 23. Nader. Kuhnlein... B. and Land. Res... Res. A.. S. Stich. R.. S. and Mahan.. Lancaster. Res. C. van Tassell. W. Mutagen testing using TRP+ reversion in Escherichia coli.. 1984. 221. and Dion... J. Res. 1978. 653. Sharma. M. D. 152.. 1981. in Origins of Human Cancer. Watson. 5. P. F. Dion. R. N. 31. B. Nutr. J... Int. Int. McCann. and Kuhnlein. The need for a mammalian test system for mutagens: action of some reducing agents. B. Land... Lederman. Ed. England. and Winsten. The effect of short-term dietary modifications on human fecal mutagenic activity. Darby. H. Malt. 1. Hiatt. S. R.. W. A.. Sharma.. A. 1976. R. 11. E. 1986. P. Res. 19.. Wang. Mutat. R.. Kingston. In vitro production of human fecal mutagen. J. A. W. Reddy. and Bruce. 7. Stich.10_fm Page 319 Sunday... Perrino. F. Metabolic epidemiology of colon cancer: fecal mutagens in healthy subjects from rural Kuopio and urban Helsinki.. 3. H. van Tassell.. D... Mathews. M.. U. H.. 4. P. Erhrich. C. H. Sharma. Res. and Korpella. Varghese. S. 1980. Furrer. 1980. 14. Mathews. Bruce. acid. A. Mutat. Hara. B.5-f)quinoline to dietary fibers.. 1992. Reddy. W. B... M. R.. 77. T. 1986. M. M... and Goldman.. 34. de Kok... L. 1993. The nonfermentable dietary fiber lignin alters putative colon cancer risk factors but does not protect against DMH-induced colon cancer in rats. Nutr. H. Barnes. E. 1997. M. 2001 6:51 PM 320 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and Mutai. Invest.. 26.. Hollands... S. . 28. Effects of purified cellulose and pectin fiber diets on mutagenicity of feces and luminal contents of stomach. 195. C. 757. Hardman. A. 102. Gallagher. T. H. 1475. 29. 30..... and Gustafsson. 35. 302.... Effects of differing purified cellulose. Wassen. J.. Morotomi. 153. and Alberts. 3681. Adsorption of mutagens by refined wheat bran. W. 32S. 33. 28.10_fm Page 320 Sunday. In vitro binding of potent mutagenic pyrolysates to intestinal bacteria.2-dimethylhydrazine-induced rat colon carcinogenesis. Simi. J.. and Heitman... Takeuchi. U. 32. J. Protective role of wheat bran fiber: data from marker trials. L. In vitro study on the effects of fecal composition on fecapentaene kinetics in the large bowel. J.. In vitro binding of the food mutagen 2-amino-3methylimidazo(4. 263. Freeman.. J. Earnest. 244. and Ferguson. Med. J. A... D. The effect of a shift from a mixed diet to a lacto-vegetarian diet on human urinary and fecal mutagenic activity. M. S. 46.. Hogstedt. 3RD EDITION 24. Natl. Cancer Inst. 173. Res. Med. 31. and Kada.. J.. G. S. May 6. D.. Engle. R. 5529.. Mutat. Effect of dietary fiber on colonic bacterial enzymes and bile acids in relation to colon cancer. Johansson. 1999. Res.2387_ch4. T. Inoue. A model system for studying the adsorption of a hydrophobic mutagen to dietary fibre.. Harris. M.. 1986. ten Hoor. A. 170. 70. Holmen. 6. small and large bowel in rats. Ottova.. W.. Anticancer Res. D. J. L.. F. M. Res. and Weisburger. B. 27. 253.. Carcinogenesis. 1983. Cancer. J. Cancer Inst. 106.. Prevention of colon carcinogenesis by components of dietary fiber.. 1990. 25. H. Natl. Cameron. Roberton. Am. L. M. P. S.. Clin. G. Gastroenterology. J. and hemicellulose fiber diets on fecal enzymes in 1. 1999... pectin. H.. Persson. Mutat. 103.. L. Reddy. 1992. Cancer Res. 1988. L. Kuhnlein. R. 204. J. and Freeman. J. I. 1983. Einspahr. C. and Kleinjans. van Iersel. 19. Maiello.. 13. 1.11.1) or psyllium (1 study.1 through 4.11. p < 0.11. May 6. 2001 6:53 PM CHAPTER 4. ns.01. pectin.01.1).2).11. which have little or no acute effect (Tables 4.11_fm Page 321 Sunday. when guar (4 studies. Wolever and David J.3).11. the blood glucose response was reduced by an average of 44% (p < 0. The degree of flattening of the glycemic response does not appear to be related to the dose of guar or pectin.00+$1. pectin reduced the glucose response by an average of 29% (n = 10. there was no effect on blood glucose responses. Table 4. ADDITION OF FIBER TO SINGLE TEST MEALS There is evidence that the viscosity of purified dietary fiber is directly related to its effect on glucose tolerance. Table 4.2387_ch4.11. Figure 4.2).11. guar and other viscous fibers must be mixed with the carbohydrate in a hydrated form. p < 0. Table 4. when guar gum was adequately mixed with a test meal. S. By contrast. food form.3). 6 other gelling agents by 23% (n = 8. A. and 7 other nongelling fibers by only 17% (n = 10. The necessity of adequate mixing of fiber with carbohydrate for 0-8493-2387-8/01/$0.50 © 2001 by CRC Press LLC 321 . Table 4. Necessity of Adequate Mixing of Fiber with Food To be effective. In 24 groups of subjects in 15 studies.11. psyllium.001).4. p < 0. Table 4. Table 4. The addition of pectin or hemicellulose did not enhance the effect of guar alone (Table 4.1). wheat bran by 27% (n = 6.11.2.11. and the so-called “antinutrients” all combine to produce the glycemic response typical of the whole food. Thus. Table 4. p < 0. possibly due to differences in experimental design.11. Jenkins INTRODUCTION This chapter will review the relationship between gastrointestinal events and carbohydrate metabolism in the short and long terms.3) was sprinkled dry onto foods or taken before the main carbohydrate portion of the test meal.4). and other viscous fibers have been shown to flatten postprandial blood glucose and insulin responses more consistently than wheat bran and other nonviscous fibers. psyllium by 29% (n = 13.16 Guar.01. beginning with the effects of dietary fiber and including the extension of this to more recent work on the glycemic index of foods where fiber.3).001.11.11 Effect of Dietary Fiber and Foods on Carbohydrate Metabolism Thomas M. but a linear dose-response relationship is seen with psyllium (Figure 4.11. Table 4. and milk Bread.5 G15 G10 G 5. egg.P 5 G 5. GD = gestational diabetes. * = p < 0.16.11.D N N N N N N N N D N D AN N.5 G14. butter. bread. butter.025. On the other hand.D D N D % Change with Fibera Glucose Insulin –24% ns –34%* 0% ns –41% + + + –54%*** –2% ns – 53% –36% + + – 32%* –49% –58% –23%* –48%** –58%** –78%* –68%*** –42%** –5% ns –9%* –80%** –29% ns –50%** –38%** –27%* –0% ns –68%** –2% ns –61%* –60%* –41%* – 2% ns –23% –51%** –7% ns –22% –64%*** –65%** –44% + –66%** –37% + + + –50%*** –20% ns Ref.213–215 although delayed gastric emptying may also play a role.01. milk. butter. margarine. the consumption of fruit juice .11. egg. effectiveness is important because the mechanism of action of viscous fiber is related to its ability to reduce the rate of diffusion of nutrients from the lumen of the small intestine. and milk Bread. cheese Guar pasta.2387_ch4. + + = p < 0. bread. 1 2 3 4 5 5 6 –59%** –65%*** –40% ns 7 8 8 9 9 3 10 10 10 5 11 0% ns 11 –37%** –68% ns –48%** –45%** –55% + + + –54%** –60%** –58%*** +28% ns –6% ns –16% ns 0% ns –51% –50%** –1% ns –71% 12 12 13 2 14 15 15 16 17 18 19 19 19 20 21 17 22 % change in area under the curve (or peak rise) of blood glucose and insulin (lack of a figure means that the data is not available).2).11. and Insulin Responses to Single Test Meals Fiber (g) G 2.P10 G15.P10 G 4. jam.11_fm Page 322 Sunday.5 G 2. sucrose. May 6. *** = p < 0. egg. AN = diabetes with autonomic neuropathy.001. egg. and butter Soup. butter.5).26.05. and milk Bread.P10 G16. ** = p < 0.5 G5 G5 G5 G5 G10 Available Carbohydrate (g) Source 50 50 50 50 49 49 85 G8 G15 G15 G7 G7 G9 G10 G10 G10 G10 G5 58 100 100 42 42 50 52 52 47 46 23 G5 23 G16 G20 G23 G14. + + + = p < 0.5 G14. jam.77. 3RD EDITION Table 4. honey.30 Effect of Food Refining on Postprandial Glucose Responses Removal of cereal fiber from foods by refining has no effect on the postprandial glucose and insulin responses (Table 4. honey. honey.P 5 G16. and milk Bread. Subjects: N = normal.30. butter. egg. 2001 6:53 PM 322 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and jam (guar sprinkled on cereal) Biscuits Glucose Glucose Bread and cheese Bread and soybeans Glucose Soup.5 G14. and butter Soup and bread Guar bread and guar soup Mashed potato (guar mixed with meal) Mashed potato (guar taken before meal) Guar pasta.005.5 G14.14. bread.216 Nevertheless.D N D D N. This is consistent with the lack of effect of cereal fiber when added to carbohydrate test meals (Table 4. egg.P 5 G 5. + = p < 0. D = diabetic. ns = not significant. and milk Glucose (after fiber) Bread Subb N N N N N N D D N GD D D N N D N. ham Sucrose Glucose Glucose Glucose (mixed with guar) Glucose (taken after guar) Glucose Glucose (taken after guar) Oatmeal porridge Bread. butter.H 4 a b 66 83 80 50 50 50 50 50 50 20 72 72 72 102 106 50 25 Glucose Glucose Glucose Glucose Guar bread and soup Bread and guar soup Cornflakes.002. viscous fiber has not been demonstrated to cause carbohydrate malabsorption.1 Effect of Adding Guar Gum (G) Alone or with Hemicellulose (H) or Pectin (P) on Blood Glucose. cheese. beef. Grams pectin and glucose are per square meter of body surface.5 P14. losses were reduced by 41% after guar (p < 0. and jam (pectin sprinkled on cereal) 60 Bread. rice.01. paradoxically. LONGER-TERM EFFECTS OF DIETARY FIBER SUPPLEMENTS Normal Volunteers Nonviscous fibers have been added to the diets of normal individuals more often than viscous fibers (Table 4. but in seven studies with nonviscous fibers. disruption of food form.005. 25 26 17 27 16 28 29 30 17 31 32 33 16 34 34 % change in area under the curve (or peak rise) of blood glucose and insulin (lack of a figure means that the data is not available). corn.2387_ch4.2g/kg B20 B36 B41.001. meat.2 Fiber (g) P10 P10 P10 P7 P15 P10. ns = not significant. may be important.001). as opposed to whole fruit is associated with a somewhat greater blood glucose response and markedly enhanced peripheral insulin levels. rice. lactulose 50 Glucose 50 Glucose 50 Glucose Subb %Change with Fibera Glucose Insulin N N N –59%** –4% ns –23% ns –22% ns +11% ns –18% ns 23 23 6 D –12%* –35%* 24 D PGS N IGT N PGS PGS N N N D N N D/D D/O –44% –23%** +5% ns –25% + –11% ns –20%** –40%*** –55% + +7% ns –37%** –11% ns –54% ns –27% ns –13% ns –21% + –56% +6% –11% –2% + ns ns ns –76%*** +11% ns +6% –16% –8% –34% 0% nsd ns ns ++ ns Ref. tomato 50 Glucose.5 B10 B0.11. rather than removal of fiber per se. Treatment of Diabetes There have been more than 20 studies using guar gum or other viscous fibers to treat diabetes and about half that number using various nonviscous fibers (Tables 4. + + + = p < 0. cheese. 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM Table 4. Subjects: N = normal.11.50–52 In these cases. milk.002.05.11_fm Page 323 Sunday. In 18 studies where urinary glucose was measured. sucrose. * = p < 0.8).025. there was a mean increase of 34% (ns). + + = p < 0. PGS = postgastric surgery: IGT = impaired glucose tolerance with reactive hypoglycemia.6).11. D/D = diabetic on diet alone. These studies show. ** = p < 0. *** = p < 0.11. that nonviscous fibers have more effect than viscous fibers in improving fiber-free glucose tolerance tests given after a period of fiber supplementation (Table 4. bread.5 B50 B50 a b c d 323 Effect of Adding Pectin (P) Alone or Wheat Bran (B) on Blood Glucose. D = diabetic.3). bread.11. D/O = diabetic on oral agents. May 6. and Peach 105 Bread and jam 75 Glucose 50 Glucose (after pectin) 45 Glucosec 50 Glucose 50 Glucose 50 Glucose 50 Glucose 50 Glucose (after bran) 1 g/kg Glucose 90 Beef. Represents change in insulin requirements of Type 1 diabetics on an artificial pancreas.11. and Insulin Responses to Single Test Meals Available Carbohydrate (g) Source 100 Glucose and beef 100 Glucose 85 Cornflakes.5 P14. + = p < 0.5 P14.6). Fasting blood glucose was reduced by a mean of .7 and 4. The improvements in diabetic control after the addition of viscous fibers were larger and more consistent than after the addition of nonviscous fibers (Figure 4.5 P10 P9 P14. meat. D = diabetic. 3RD EDITION Table 4.35 Liquid formula diet Sub c N D IGT N N N N N DO D % Change with Fiberb Glucose Insulin –26% ns –6% –22% ns +16% ns +24% ns –14% ns –65%* –39% ns –21% –18% ns –24% ns 0% ns 0% –24% –19% 0% –7% ns ns ns ns ns Ref. + + = p < 0.11_fm Page 324 Sunday.1 S12. % change in area under the curve (or peak rise) of blood glucose and insulin (lack of a figure means that the data is not available).5 X/L2. beef. 16 25 27 31 31 39 33 33 40 41 Q = cholestyramine.8 S15. Subjects: N = normal.01.05. cheese. A = agar.11.05. D = diabetic.01.2g/kg S20 S22 F30 P10 P10 b c Subc Glucose Insulin Ref. milk Bread.2 S16. Table 4. sucrose. G = bagasse. lactulose 58 Noodles.025. jam Glucose Glucose (psyllium as Fibogel) Bread. * = p < 0. ns = not significant. L = Locust bean gum. May 6.005.5 L10 X2.3 Effect of Adding Various Gelling Agents on Blood Glucose and Insulin Responses to Single Test Meals % Change with Fiberb Fibera (g) T14. egg. This figure represents the percent reduction of GIP response. + = p < 0. lactulose 50 Glucose. F = pea fiber. margarine 67. milk. beef.7 S15.5 M14. * = p < 0. K = Konjac mannan.5 B15 C9 C 0.6 S6. + + + = p < 0. butter. % change in area under the curve (or peak rise) of blood glucose and insulin (lack of a figure means that the data is not available). milk.002. M = Methyl cellulose. *** = p < 0. bread. milk. Subjects: N = normal.025.5 A10 K5 S3. juice. bread.11. cheese Glucose (psyllium as Fibogel) Glucose (psyllium as Fibogel) Glucose (psyllium as Metamucil) Psyllium-enriched flaked bran cereal Psyllium-enriched flaked bran cereal Psyllium-enriched flaked bran cereal Psyllium-enriched flaked bran cereal Psyllium-enriched flaked bran cereal Psyllium-enriched flaked bran cereal Psyllium sprinkled on bran flakes Psyllium taken just before bran flakes N N N D N N D N N D D N D N N N N N N D N N –34%** –29%* –29% ns –15% ns –41%* –28% ns –7% ns 0% ns –33% ns –33%* –29% –12% ns –17% ns 0% ns –12% ns –24% ns –20% ns –46%** –46%** –51%* –56%** –4% ns –10% ns –18% ns –19% ns –11% ns –37%** –64%** +17% ns –28%* –39% ns –19%*d 16 16 1 6 1 1 6 35 2 36 37 2 2 2 38 38 38 38 38 38 38 38 –28% ns –13% ns T = Gum tragacanth. + = p < 0.005. IGT = impaired glucose tolerance. P = soy polysaccharide. 2001 6:53 PM 324 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.001.002.7 S16.2 a b c d 50 50 50 85 50 50 85 80 50 50 50 50 50 50 50 50 50 50 50 50 50 Fiber (g) Q14. jam Glucose Glucose Cornflakes.5 L2.2387_ch4. sucrose. ** = p < 0. Glucose Glucose Glucose Cornflakes. *** = p < 0. + + + = p < 0.4 a Available Carbohydrate (g) Source Effect of Adding Various Nongelling Fibers on Blood Glucose and Insulin Responses to Single Test Meals Available Carbohydrate (g) Source 50 Glucose 105 Bread and jam 45 Glucose 1 g/kg Glucose 1 g/kg Glucose 86 50 g sucrose. butter. S = sugar beet fiber. B = barley bran.6 S7 S7 S7 S3. + + = p < 0. ** = p < 0. S = psyllium. ns = not significant. C = cellulose.7 S16. 45 g flour 50 Glucose.9 S8. DO = obese diabetic.5 S3. .2g/kg G 0.001. ns). 2001 3:03 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM Figure 4.1 through 4.8% after nonviscous fibers ( p < 0. 11% after guar ( p < 0.2387_ch4.11.01) and 3.11.02). Serum cholesterol fell by 11% after guar ( p < 0.11. These fibers are associated with short. 05) but not after nonviscous fibers (+1%.001) and 3.and long-term beneficial changes in carbohydrate metabolism. Insulin doses tended to be reduced on guar (–11%. Glycosylated hemoglobin fell by a mean of 5.11_fm Page 325 Tuesday.2 Effect of psyllium dose on acute glycemic responses (data from Table 4. Summary — Long-Term Effects of Dietary Fiber Supplements Those types of fiber with the most marked short-term effects also appear to have the greatest ability in the longer term to improve diabetic control.4). p < 0.11.4% (ns) after nonviscous fibers. Figure 4.3). May 8. ns). such as the rate of absorption of glucose. Viscous forms of dietary fiber such as guar gum are most able to alter events occurring within the gastrointestinal tract.1 325 Effect of various types of purified dietary fiber on acute glycemic responses: proportion of studies with a statistically significant reduction (data from Tables 4. .0% after guar ( p < 0.11.001) but not after nonviscous fibers (–1%. Study length: D = days.005.01.05.e. W = weeks. + + + = p < 0.05. *** = p < 0. *** = p < 0. 2-h postglucose blood glucose values. % change in area under the curve (or peak rise) of blood glucose and insulin (lack of a figure means that the data is not available).0 16. 26 Corn bran.002. May 6.0 4. Removal of Fiber) on Blood Glucose and Insulin Responses Test Meal Contents (g) Dietary Fiber Whole Refined Available Carbohydrate Food Wheat bread Wheat bread Wheat bread Wheat bread Wheat bread Rice Rice Spaghetti Maize meal Potato Oranges Oranges Oranges Grapes Apples a b 50 50 58 and 60 50 25 50 75 50 50 50 50 50 30 60 60 10.005.8 2.6 a –18% ns Ref. (D) = diabetic).002. + + = p < 0. + + + = p < 0.2 0. 12 Xanthan gum. ** = p < 0. D = diabetic. * = p < 0. Subjects: N = normal.5 1. 20 Wheat bran. 3RD EDITION Table 4. + + = p < 0.2 10. 36 Wheat bran.6 0. 12 Wheat bran. + = p < 0. CHO PRO FAT Comments Glucose Insulin Ref. 26 Apple/carrot. Table 4. 26 Wheat bran.3 2. ** = p < 0.0 0.5 Effect of Refining Food (i.01. ns = not significant.2387_ch4. 39 15 42 +100% ns ns 53 47 14 36 FRD. 26 Soy hulls. + = p < 0. 21 Soy hulls.11.001. 24 Wheat bran.001. .8 3. M = months. 25 c +50% + + + Effect of Dietary fiber on Fiber-Free Glucose Tolerance Testsa Diet Constituents b +14% ns 42 43 44 45 46 42 47 42 48 49 49 50 51 50 52 % change in area under the curve (or peak rise) of blood glucose and insulin (lack of a figure means that the data is not available)..025. 15 6W 4W 7W 6M 30 D 30 D 30 D 3W 3W 30 D 30 D 6W 6 W(D) 2W 2 W(D) d +32% + +51%** Change after Fiberc b Gum arabic.11_fm Page 326 Sunday.0 8. 26 Xanthan gum.2 Subb 2. 15 Guar gum. 2001 6:53 PM 326 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. ns = not significant. 21 Soy hulls. 12 Guar gum.e.11.0 N D N N D N D N N N N N N N N % Change with Refininga Glucose Insulin – 6% ns – 2% ns –14% ns – 5% ns + 7% ns +11% ns + 6% ns +11% ns + 4% ns +14% ns +15% ns ns +6% ns +28%* ns Fiber Dose (g/day) Study Length 21 D Pectin. all fiber in morning MD FRD MD FRD MD MD MD FRD undried DF FRD dried DF MD MD FRD FRD FRD FRD +11% ns ns +36% + 54 55 56 45 57 57 57 58 58 57 57 59 59 60 60 40 42 19 19 40 37 +18% ns ns – 32%* –15%** – 17% +24% – 56%* +5% ns +59%* – 72%* – 28%* – 25% nsd – 31%* – 5% ns –4% ns ns ns ns +10% ns –7% ns ns ns –6% ns – 4 % ns +13% ns Subjects are normal except where indicated after study length (i. * = p < 0..4 7.025. In this study. UG = urinary glucose. jam. NC = poorly compliant. CAP = capsules.05). O = obese.11.05.In 18N 9I 9N.Child 20N 2I.PC 8I. LOW = weight-reducing.6N 12I 5I 5N 6I. and insulin receptor number by 50% (p < 0. N = noninsulin-dependent. SW = snacks and sweets. *** = p < 0.ND. EC = excellent control. 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 327 . jelly. HD = hospital diet.11N 10N 10I. fruit bar).O. BR = bread.01.1N 6I.NC 20N. HbA1c = glycoylated hemoglobin.025).PC 14N 22I. Insulin = daily insulin dose.3N 6I.O.B10 G21 G22 G15 G15 G14−26 G14−26 a Treatment of Diabetes with Guar (G) or Guar plus Wheat Bran (B) Dose (g) Table 4. B = brittle.45g/kg G20−25 G15 G15 G15 G20 G15 G8 G12−19 G10 G 10−15 G15 G 15 G20.7 2387_ch4.O. guar increased peripheral insulin sensitivity by 70% (p < 0. P = pasta. + = p < 0.025).025. SU = controlled on sulfonylureas. Child = children. MT = minitablets. BI = biscuits. * = p < 0. Diets: MD = metabolic diet.EC 12N. PC = poorly controlled.001.2 G20 G15−25 G. CB = crispbread. Represents fall in fasting serum insulin. FP = flavored powder taken in water before meals.11_fm Page 327 Sunday. Subjects: I = insulin−dependent. Form = type of guar formulation.SU 38N.3N 8N. R = rusks. TG = serum triglycerides. May 6.Child 6I 41N.PC 9N 10I. insulin binding to monocytes by 28% (p < 0. FBG = fasting blood glucose. CHOL = serum cholesterol. VF = guar incorporated into various foods (bread. ND = newly diagnosed. GR = granulate. + + = p < 0. 5D 5D 7D 7D 2W 2W 3W 4W 4W 4W 4W 4W 4W 4W 4W 6W 6W 8W 8W 8W 8W 8W 3M 3M 3M 3M 6M 12M a Length G25 G23 G10 G24 G 9.2 G 9. ** = p < 0.002. low-calorie (results for guar compared to a separate control group on a low-calorie diet not given guar).O 29N.O. In = controlled on insulin.3N 2I.B 6N.d c b Subjects 6I. FRD = free-range diet.2N MD MD FRD HD FRD FRD FRD FRD FRD FRD FRD FRD FRD FRD FRD FRD FRD FRD MD LOW FRD FRD FRD FRD FRD FRD FRD FRD BI BI GR GR BR SW MT MT GR GR GR BR VF GR P GR MT GR GR R CAP GR CB CB CB BI Commentsb Diet Form –3% ns –24%** –16% ns 0% ns – 39%+ –17%+ –19% ns 0% ns –10% ns +4% ns –25%* ns –11%* ns –22%** –20%* –7% –8% ns –18%** –10%* +13% ns FBG –5%ns –3% ns –7% ns +2% ns –10%* +2% ns –7% ns ns –9%*** –9%*** –11% ns –3%* –7% ns – 44%* –73%* –60% –15%*** –32% ns –66%+ –10% ns –51%** –8% –49% ns –50%* –14%* –32% –80%** –62%** –50%** –38%+ + –28%*** –26%** –14%+ –3% ns –5%+ 0% –4% ns % Change after Fiber Treatmentc HbA1c UG Loss Insulind –9%*** –10%* – 9% – 9%* –10%* –10%* – 21%+ –8% ns ns –11%+ 0% ns –16%* –2% ns –10%*** –30%+ +11% ns –18%** –14%* –14%** CHOL –4% ns +8% ns +8% ns –40% ns +5% ns –17%* –12%* –2% ns +13% ns –25% ns 0% ns ns TG 61 62 7 63 7 7 64 65 66 67 68 68 69 70 71 72 73 74 12 75 76 77 78 79 80 81 82 82 Ref. 001. Subjects: N = noninsulin-dependent diabetes. *** = p < 0.002. Insulin = daily insulin dose.11. CHOL = serum cholesterol. I = insulin-dependent diabetes.6 7. W = weeks.8 3. MM = “Metamucil".8 2387_ch4. FRD = free-range diet. 3RD EDITION . May 6.2 3.01.6–7 c b a 9N 13N 9I 20N 20N 8I 17I 18N 38IGT 10N 10N 10N 10N FRD MD FRD FRD FRD MD MD FRD FRD FRD FRD FRD FRD FRD FRD FRD MU MM MM MM BR BR –38%* –29%+ –6% ns –11% ns Psyllium Cellulose –13% ns –2% ns Wheat Bran –2% ns –1% ns Soy Hulls –2% ns –8%* Corn Bran +2% ns +3% ns –8%** Apple Fiber HbA1c Xanthan Gum Konjac Mannan +6% ns –19%** –18%** –6% ns 0% ns –5% ns –10% ns –1% ns –6% ns +2% ns –2% ns –2% ns –8% ns FBG –36%** + 5% ns + 7% ns +26% ns +67% ns +49% ns +119% ns Reduced 0% 0% +2% ns –13%* –11% –5%* –5%* 0% ns –9%*** 0% ns +3% ns –2% ns +1% ns –5% ns +6%* −5% ns % Change after Fiber Treatmentc UG Loss Insulind CHOL –25% ns +7% ns –10%* +9% ns +3% ns +1% ns –4% ns +5% ns +2% ns 0% ns TG 59 89 35 88 87 87 85 69 86 83 83 83 83 83 83 84 Ref. M = months. Length: D = days. TG = serum triglycerides. UG = urinary glucose. BR = bread. * = p < 0.4W 4W 10–15D 4W 4W 10D 7D 2M 2M 30D 26 52 14 15 20 15 10. + = p < 0. IGT = impaired glucose tolerance. 6W 4W 4W 26 52 10N 10N 12N Subjects Commentsb Diet Form 328 12 4W 4W 7W Length a Treatment of Diabetes with Miscellaneous Types of Dietary Fiber Nonviscous Fibers 26 52 15 Dose (g) Table 4. HbA1c = glycoylated hemoglobin. Diets: MD = metabolic diet.11_fm Page 328 Sunday.025.05. 2001 6:53 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. FBG = fasting blood glucose. ** = p < 0. MU = fiber in muffins. + + = p < 0. Form = type of fiber formulation. 11..224. the GI makes it possible to compare the blood glucose responses of foods tested in different groups of subjects. 2001 3:04 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM Figure 4. have the lowest GI of all starchy foods. amylopectin). Tables 4. again in the context of the whole food.219 and enzyme inhibitors.11_fm Page 329 Tuesday.220.11. Among these are antinutrients such as phytate.221 macronutrient interactions such as protein-starch222 and lipid-starch.3 329 Effect of purified dietary fiber in the treatment of diabetes. May 8.9 shows . Table 4.223 the nature of the starch (amylose vs. Proportion of studies with statistically significant improvements in fasting blood glucose (FBG).11.11.217 lectins.2387_ch4.225 gastric emptying rate226 food form (e.. BLOOD GLUCOSE RESPONSE TO FOODS In whole foods. so that the relative importance of each of the factors can be determined. many factors in addition to dietary fiber may alter digestion and absorption of nutrients and hence the glycemic response.g.158 Area under the glucose curve after a food × 100% Glycemic Index = -----------------------------------------------------------------------------------------------------------------------Area under the glucose curve after white bread In this way. fiber) to be assessed.11. and also to compare the full range of foods. and serum cholesterol concentration (data from Tables 4.7 and 4. The Glycemic Index of Foods The glycemic index (GI) of foods is an expression which permits the blood glucose response to different foods to be compared directly and so allows the relevance of their constituents (e.105.9 and 4. It therefore becomes important in any discussion of glycemic response to be able to assess the collective interaction of all these factors in terms of the glycemic response to the whole food.218 tannins. The methods used to determine the glycemic response and calculate the area under the curve can have a major influence on the results obtained. bread as opposed to spaghetti). glycosylated hemoglobin (HbA1c).g. urinary glucose excretion. and particle size. as a class.10 show the wide range of Gl values for different foods. It is defined as the incremental blood glucose response following consumption of a 50-g-available carbohydrate portion of a food expressed as a percent of the response after a standard 50-g-available carbohydrate portion of white bread taken by the same individual. The legumes.11. cooking.8).227 exact GI methods are given elsewhere. 93. 96. B U. 66. May 6. J. 41 25. W A. 65 10. 32. 77. 86. Tn. K. A D. Z. 77. 70. A. 16 39. M CVc 15% 6% 4% 16% 23% 9% 43% 21% 60 15 ± ± ± ± 5 13 14 17 9% 16% 23% 24% 26 ± 11 138 ± 10 45% 7% . 46. Z 67 ± 16 23% 31. 32. B. 58. A. C. 119. S 82 100 Root Vegetables Carrots Potato (instant mashed) Potato (mashed) Potato (new/white boiled) Potato (Russett. D. A. Pi M.2387_ch4. A. C. W. 72. 95. U. Tn. A. 68. 100. 60. B. 3RD EDITION Glycemic Index Values of Foods Tested in Several Studies Studiesb Mean ± SD 90. G. 137 ¢. G. Q. W. 76 107. A Ti. # 95 98 ± 14 68 100 100 ± 6 46. O. 83. L. L. 119 58. β. B. 132.9 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. A C. 97.67. 72. 158 V. V 52 82 59 71 Food Breads Rye (crispbread) Rye (whole meal) Rye (whole grain-pumpernickel) Wheat (white) Wheat (whole meal) Pasta Spaghetti (white) Cereal grains Barley (pearled) Buckwheat Rice (polished) Rice (parboiled) Sweet corn Legumes Baked beans (canned) Bengal gram dal Butter beans Chick peas (dried) Green peas (dried) Green peas (frozen) White beans (dried) Fruit Apple Banana Orange Orange juice Sugars Fructose Glucose Individual GI Valuesa 48. A. B. 75. A. 90. E. 145. 84 27. 56. J J. 43. dried) Lentils (red. 78 66. M C. 112. M J. D 36 78 ± 11 81 ± 12 14% 15% 69 ± 9 80 ± 11 12% 13% Breakfast cereals “All Bran” Cornflakes Porridge oats 71. A U. B. £. 80 52 43. 56. H. A @n. A. A. 89. 116. 74 18. B J. P E. H. 139 71. 92. @h. 27. baked) 96. A. A 70 ± 10 12 46 47 ± 4 50 65 45 ± 20 Kidney beans (dried) Lentils (green. B. C V. 99 73 90 9. J C. 75. L. 88. C. 46. B. D 114 120 ± 5 98 80 ± 13 116 ± 26 14% 43 ± 16 36 39 ± 8 38% Pinto beans (dried) Peanuts 55. 106. 105. 30. Y. 45. 73. 104. B. A. A A. 90 F. 35 122. W. 96. 42. 121. Z. # C. 90 68. 69. 137. #. Xi M. K E. 100 67. 68 55. C. 74. V. 108 C. 59. L.11. 31. 46. Xi Xn. Ti. 108 B. A C. 78. Xn. 2001 6:53 PM 330 Table 4. K. 106. 7. 100 82. 96 B. 88 I. 126 96. W 74 ± 3 121 ± 13 89 ± 10 3% 11% 12% Cookies Digestive Plain crackers (water biscuits) 77. 29. A Y. E. £. A. P. B. U. P. 31. 45 57 81. 66. A. 140. Z R. C. dried) 60. A C. N L. Pn. A. 41 70. S. 78 100 (Defined) 93. W U. 101 80. 52. 74. 52 44. 131. 134. 87. 43. 78. 67. U J. C. Xi. J. 30. 104 58. 44. J.11_fm Page 330 Sunday. Y Y. 65 42. 85. A A. 96. 19 A. 45. 141. 131 116. 86 91. G A–K. $. N(101).238 Large amounts of protein enhance insulin secretion (Table 4. Z. fat taken at one meal may impair carbohydrate tolerance in the subsequent meal. J(97). H(95). K V.11) but have little effect on the blood glucose response in the smaller amounts normally eaten. only in large amounts (45 to 65% of total test meal calories) does it significantly reduce postprandial glucose and insulin responses (Table 4. the mean GI values for foods tested in different groups of subjects and by different investigators. K. tested 3 different foods 4 times each.111 In a recent study.2387_ch4. there is reasonable agreement between the values derived in different centers. instant potatoes) for unequal amounts of carbohydrate. £(116).g. in one study. 12 subjects with diabetes. G(94). as opposed to whole grains.99.106. Q(104). S(106). U(108). Y(112). including individuals with IDDM and NIDDM. C(46). L(99). @h. V(109).49.12). T(107). B(90). Most of the variability in GI values was due to day-to-day variability within subjects with no significant difference in GI values between the different individuals. E(92). J. regardless of whether the subjects tested were normal or diabetic Insulin-dependent diabetic subjects (IDDM) tend to show slightly higher GI values than subjects with noninsulin dependent diabetes.136 and the whole grain rye breads popular in Northern .11.11. @n.11. and that soluble fibers have differing viscosities and hence variable effects in impeding glucose absorption. in the bread increases. The GI values of foods are related to their fat and protein contents. 92. the mean glycemic response to bread was greater than that to rice. n = NIDDM.9 (Continued) Glycemic Index Values of Foods Tested in Several Studies Individual GI Valuesa Studiesb 45. P(103). β(114). was given. 49 A.126 However.111 This may be a reflection of the high degree of withinindividual variability of blood glucose responses in IDDM to the same meal rather than to true differences between foods tested in these individuals. Where glucose.236 Surprisingly.123. 39. May 6. 54. L 77 Food Lactose Sucrose a b c Mean ± SD CVc Individual mean values from groups of subjects.11. 86 26. 100 V. X(111). A 69 ± 19 36 ± 10 27% 29% Snack Foods Potato chips 74. The studies are listed in the same order as the values: A(49). For 27 foods tested by 3 or more groups of subjects. &(120): h = healthy subjects. @n 57 86 ± 12 14% Dairy Products Ice cream Whole milk 52.9).135 finely ground flour produces higher glycemic responses than coarse flour. @(119). W(110).11.235 However. S @h. 29. $(115).11_fm Page 331 Sunday. Values have been adjusted proportionately so that the glycemic index (GI) of white bread = 100. C. the strongest correlation between food GI and components of fiber was with cellulose and the uronic acid fraction of insoluble fiber. Z(113). In addition. K(98). A @h. M(100). The lack of relationship between food GI and soluble fiber is probably due to the fact that many other factors in foods influence digestibility. R(105). which. the mean SD of the mean Gl values is 12 (Table 4. breads.228 The GI values of foods relate strongly to the rate of digestion of foods in vitro. i = IDDM. and not significantly to soluble fiber. 85.11). O(102). 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 331 Table 4. 86. D(91). F(93). bread with cheese. was greater than that to spaghetti. @n. #(118). S.4). T. 91. in turn. CV = coefficient of variation (100 × SD/Mean). peas.38 (the GI of glucose/100).49 Fat delays gastric emptying237 and enhances GIP responses.90. The glycemic response to wheat and barley breads increases linearly as the proportion of flour. Values have also been adjusted proportionately (studies C.229–234 but only weakly to total fiber content. 69 61. the GI values were multiplied by 1. In general. increasing the soluble fiber content of a meal with no change in its GI resulted in a major reduction of the postprandial glycemic response in subjects with NIDDM. I(96). In all 12 subjects.. e. major components of the cell wall (Figure 4. 84.11.103 Particle size is particularly important in determining the responses of starchy foods (Table 4. but not white bread. 87. ¢(117). 79 A. May 6. data from 11 other groups worldwide (Lund. As the number of tests done increases. a similar phenomenon was observed.11.240 In a meta-analysis of data from the literature (Table 4.11.11_fm Page 332 Sunday. Table 4. and the method of calculation of area under the curve must be the same as that used for the GI. russet potato.11.94 Increased particle size is probably related to the reduced glycemic effect after swallowing foods without adequate chewing. Melbourne. respectively [ns]. Variables such as food variety (e. with the weighting based on the proportion of the total carbohydrate contributed by each food (Table 4. Differences in food form may also be of great importance in determining the glycemic index of a food. in the absence of increased fiber. the degree of ripeness (e. Clinical Utility of the Glycemic Index The GI of mixed meals can be estimated from the weighted mean of the GI values of the individual carbohydrate foods in the meal. Toronto.g. fiber appears to be more effective at higher rather than lower carbohydrate intakes (Table 4.160.5). rice.241 However. Table 4. Naples. To be of predictive value.7).e.11.13).2387_ch4. For small numbers of subjects.11.11.g.. When guar gum was added to the diets of diabetic patients for 5 days. 3. who have greater variability of glycemic responses.15. the safety of highcarbohydrate diets in diabetes has been questioned.9. are not consistently effective in reducing fasting blood glucose and serum cholesterol levels (mean reductions.11. and Sydney) are grouped around the line of identity (Figure 4.161.. new vs. 2001 6:53 PM 332 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.11. High-Fiber Diets High-carbohydrate diets.15) and may increase postprandial blood glucose and insulin responses.8). Aarhus.11. the GI value of mixed meals predicts almost exactly the relative differences between their mean glycemic responses (Figure 4. Bristol. in subjects with IDDM.14).11. 34 is the predictive difference). Table 4.228 For a subject with NIDDM. For these reasons.9). Minneapolis (2 different groups). and differences in cooking and processing (e. Morgantown. the expected difference in meal GI. insulin.11. However. the predictive difference is reduced by a factor of 1/ n.10).11. In our hands. The use of high-fiber foods in the diet.11. the predictive difference is 50. and the number of subjects or repeat tests done. Table 4. Tables 4.9). at least in the short term (2 to 6 weeks. cooking and processing of foods markedly affect their in vitro digestibility.242 Those patients whose diets contained more than 40% carbohydrates had a larger reduction of their urinary glucose output on guar than those patients whose diets contained less than 40% carbohydrates (64% compared with 33%. the most appropriate use of the GI is to help in determining which of two meals is likely to produce the greater glycemic response. and triglycerides and enhance their cholesterol-lowering effects.16.2%. p < 0. The chance of a correct prediction depends upon the day-to-day variability of glycemic responses in the subjects being tested. where overall carbohydrate content is kept constant. In the limited number of studies done in this situation. banana. 3RD EDITION Europe have relatively flat glycemic and insulinemic responses. the calculation of meal GI must take into account all of the carbohydrate foods in the meal.4 ± 2. 4..02). increase fasting triglyceride levels.6). the use of foods rich in dietary fiber may offset the deleterious effects of high-carbohydrate diets on blood glucose.g. 161 Treatment of Diabetes with High-Carbohydrate.9) may therefore explain some of the differences in GI values obtained for the same food by different investigators.11. as exemplified by the similar composition but very different GI values of white bread and spaghetti (Table 4.16). and reduce HDL cholesterol.14).239 In addition. improves diabetic blood glucose and lipid control. . accurate GI values for the foods fed must be used. and these changes are reflected in similar alterations of metabolic responses in vivo (Table 4. there is a 95% chance of a correct prediction for a GI difference of 34 (i.11. the Stanford group has been unable to predict mixed meal glycemic responses using the GI of foods. but only when the increase in fiber comes from low-glycemic-index foods (Table 4..0% and 3. London (Canada).11. Figure 4.4 ± 2. Figure 4. at least in part.16.2387_ch4.254 low-glycemic-index foods.g.265–268 which influence carbohydrate and lipid metabolism269–271 and may be part of the mechanism of action of high-carbohydrate/high-fiber diets. increased meal frequency. Figure 4.8).253.11_fm Page 333 Sunday. In these studies. but this was achieved often at the expense of increased postprandial blood glucose and serum triglyceride levels (Table 4.245. This suggests that the long-term metabolic effects of high-carbohydrate/high-fiber diets depend. the use of low-glycemic-index foods. . and lower insulin dosage. 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 333 The combined use of high-carbohydrate/high-fiber diets has benefited both IDDM and NIDDM patients in terms of improved blood glucose control. In studies where the carbohydrate and fiber intakes were increased using high-GI.272–274 The long-term consequences of these morphologic and absorptive changes are likely to be important and remain to be explored..and noninsulin-dependent diabetes. blood glucose and lipid lowering effects similar to those produced by high-carbohydrate/high-fiber diets were seen with no changes in carbohydrate or fiber intake. but only in those studies where the increase in carbohydrate and fiber was achieved with low-GI foods such as legumes and pasta (Table 4.11.17.247–252 Increased starch consumption. May 6.7).244 CONCLUSIONS Long-term benefits have been demonstrated using high-carbohydrate/high-fiber diets in both insulin.11.246 Fiber has been shown to alter gut morphology in experimental animals.11.16. upon the ability to reduce the rate of carbohydrate absorption and hence reduce the acute postprandial blood glucose response. A number of dietary trials have been undertaken where the GI of the diet has been reduced without changing its overall macronutrient or fiber content (Table 4. cereal products (e. Slow-release carbohydrate may be obtained by the addition of viscous types of dietary fiber. Figure 4.244 and is associated with significant reductions in serum total and LDL cholesterol and apolipoprotein B levels. Recent studies where the rate of absorption was reduced by nibbling small meals at regular intervals throughout the day instead of consuming exactly the same food in three large meals indicate that nibbling reduces mean day-long insulin levels by about 30%243. reduced serum lipid levels. fasting blood glucose and cholesterol levels were improved. and butyric acids.11. or pharmacologic intervention with amylase inhibitors such as Acarbose. propionic. The mechanism for the lipid-lowering effect of slow absorption may relate to reduced insulin secretion and hence reduced stimulus to lipid synthesis. This will result in the formation and absorption of the short-chain fatty acids and acetic. These are associated with.11.8). Figure 4. slowing the rate of carbohydrate absorption within the gastrointestinal tract. whole meal bread).255–260 and dietary fiber261–264 also increase the amount of fermentable carbohydrate which enters the colon. and may be due to.11. boiled 5′) Spaghetti (protein-enriched) Spirali (durum) Star pasta (white. boiled 15′) Spaghetti (white. boiled 5′) Rye kernels Unrefined maize meal porridge Refined maize meal porridge Wheat kernels Wheat kernels (quick-cooking) 65 103 65 121 58 54 47 71 74 63 75 G A H L H H 6 J J G S Breakfast Cereals Muesli Puffed rice Puffed wheat Shredded wheat “Weetabix” 96 132 110 97 109 A L S A A Cookies Oatmeal “Rich Tea” Shortbread 78 80 88 A A S Miscellaneous Mars bar Corn chips Tomato soup 99 99 55 A L A 93 113 99 73 68 70 74 75 A & & & J A A J 60 74 74 64 48 24 U U U U A J Food Breads Wheat (French baguette) Wheat (puffed crispbread) Pasta Fettuccine (nondurum) Ravioli (durum) Macaroni (white.11_fm Page 334 Sunday. canned) Pinto beans (canned) Black-eyed peas Brown beans . boiled 5′) Vermicelli (nondurum) Root Vegetables Beetroot Cassava (flakes) Cassava (Eba) Cassava (Lafun) M’fino Sweet potato Yam Pumpkin Legumes Chick peas (canned) Kidney beans (canned) Lentils (green. 2001 6:53 PM 334 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. boiled 5′) Spaghetti (brown.11.10 Glycemic Index (GI) of Foods Tested Only Once GI Ref. boiled 5′) Rice (parboiled. May 6.a 131 112 Z S 44 54 64 61 45 38 59 54 48 # # I I I I # I # Cereal Grains Bulgur (cracked wheat) Millet Rice (instant. boiled 6′) Rice (polished. boiled 1′) Rice (instant. 3RD EDITION Table 4.2387_ch4. 2387_ch4.11.4 See legend for Table 4. Relationship between the glycemic index values of 25 foods and their content of total dietary fiber. 2001 3:04 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 335 Table 4.11_fm Page 335 Tuesday.a Soybeans (canned) Soybeans (dried) 20 22 A A 45 59 90 32 36 62 40 47 34 93 V $ $ C C C C C C A 126 152 A A 59 46 52 S A A Fruit Apple juice Banana (slightly underripe) Banana (slight overripe) Cherries (raw) Grapefruit Grapes Peaches Pears Plums Raisins Sugars Honey Maltose Dairy Products Custard Skim milk Yogurt a Figure 4.10 (Continued) Glycemic Index (GI) of Foods Tested Only Once Food GI Ref.11. . and selected fiber fractions (grams/50 g available carbohydrate portion.9 for identification of references. data from Reference 235).11. May 8. May 6.11. 3RD EDITION .11_fm Page 336 Sunday.CHO (g) 50 50 50 50 50 50 50 75 75 50 50 50 50 50 50 50 50 25 25 50 50 50 50 50 50 50 Flummery ± fat Flummery ± fat Flummery ± fat Potatoes ± olive oil Rice ± olive oil Lentils ± olive oil Bread ± butter Potato ± butter Lentils ± butter Potato ± corn oil Glucose ± avocado oil Potato ± intralipid (ileum)c Potato ± intralipid (duodenum)c Potato ± butter Glucose ± lean hamburger Bread ± cottage cheese Bread ± skim milk cheese Potato ± tuna Spaghetti ± tuna Glucose ± beef Glucose ± turkey Glucose ± gelatin Glucose ± egg white Glucose ± cottage cheese Glucose ± fish Glucose ± soy Suba N NID ID N N N NID N N N NID N N N –49%* 0% ns 0% ns +40% +108% –27% –16% ns –58%* –52%* –85%** –12% ns –87%** –85% –52%++ 10 12 12 25 25 25 25 25 25 25 25 25 3 2 2 1 1 8 3 0 0 0 1 0 NID NID NID NID NID NID NID NID NID NID NID NID +9% ns –2% ns –21% ns –8%* +9% ns –10% ns –20%* –33%* 0% ns –34%* –10% ns –33% ns Effect of Adding Protein Alone 10 10 10 18 18 18 23 38 38 28 40 41 41 50 Glucose +59%*** +69%*** +230%* +230%* +260%* +190%* +360%* +230%* +220%* +25% ns +640%** +650%** –9% ns –26% ns –89%** +14% ns –79%*** –63% −7% ns +800%+ + +100% ns +400%** +700%* GIP –8% ns +5% ns % Changeb Insulin 127 90 103 128 128 129 129 129 129 129 129 129 121 121 121 122 122 122 103 123 123 124 125 125 125 126 Ref. 2001 6:53 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 336 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Effect of Adding Fat Alone Added (g) PRO FAT Effect of Fat and Protein on Metabolic Responses to Foods Test Meal Constituents Table 4.11 2387_ch4. + = p < 0.12 14 14 25 25 30 50 50 50 25 25 50 c a 8 8 21 21 21 4−20 50 50 50 50 50 50 Bread. Change in incremental area under curve or peak rise compared to low protein/fat. The fat was infused into the intestine and the potato taken by mouth. or barley ± cheddar cheese Flummery ± fat and protein Flummery ± fat and protein Flummery ± fat and protein Bread. N = normal.025. spaghetti. NID = noninsulin-dependent diabetes.01. ns = not significant. spaghetti. 25 25 25 26 26 40 11 11 10 10 10 19 Effect of Adding Fat and Protein 30 42 42 42 30 50 16 25 34 50 50 50 50 50 50 50 58 58 58 58 Glucose ± casein Potatoes ± veal Rice ± veal Lentils ± veal Glucose ± lean hamburger Glucose ± lean hamburger Sugars ± milk/soy protein Sugars ± milk/soy protein Sugars ± milk/soy protein Sugars ± milk/soy protein 2387_ch4. or beans ± cheese and butter Bread ± skim cheese and butter Bread ± peanut butter Bread ± peanut butter Potato ± tuna and margarine Spaghetti ± tuna and margarine Glucose ± casein and avocado oil 0 1 1 1 9 15 tr tr tr tr NID N N N NID NID N N N N −37% +29% +52% +12% –9% ns –35%** –40%** –51%** –70%** –78%** NID NID ID NID NID NID NID ID N NID ID NID –4% ns –27%** +2% ns –27%*** –7% ns –60%*** +3% ns +25%* –89%* 0% ns +63%** –20% ns +70%*** +85%*** +16% ns +56%* +70%* +185%** +100%* +207%** +102%** +117%** +115%** +129%** +99%** +340%** +600%** +1000%* 103 103 103 128 128 125 111 111 121 121 121 113 125 122 122 122 127 127 130 130 130 130 Sub = subjects. ID = insulin-dependent diabetes.05. * = p < 0.005.001. rice. ** = p < 0. *** = p < 0.11_fm Page 337 Sunday. May 6. potato. 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 337 . rice. + + = p < 0. 2001 6:53 PM 338 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 50. Wheat: cracked. N = normal. fine. 75. 0. ** = p < 0. 0. 50. parboiled wheat) plus various proportions of whole meal flour.005. Varied proportions of whole grain and flour in breads compared to whole grain alone. cooked pearled barley. flour. 50. Wheat breads made with bulgur (cracked. 50. cracked. Oats. 50. 3RD EDITION Table 4. Change in incremental area under the curve or peak rise.11_fm Page 338 Sunday.2387_ch4.2% retained on 710-µm sieve. 50. 50. 52% retained on 710-µm sieve. 17% retained on 2200-µm sieve. 50. 75.001. 75.025. each kernel broken into 23 pieces. 0% retained on 710-µm sieve. 50.05. + + = p < 0. 50. pumpernickel bread (bread containing whole kernels).01. barley bread vs. Rye kernels vs. *** = p < 0. 50. 50. 131 128 132 132 133 134 135 135 135 135 135 135 94 94 94 94 136 136 136 136 136 136 136 Sub = subjects.12 Effect of Grinding on Metabolic Responses Carbohydrate (g) and Source 75. Corn. 50. * = p < 0. coarse. 50. 50. a b c d e White rice Brown rice Brown rice Brown rice Cooked lentils Lentils Cracked wheatc Cracked wheat Cracked wheat Barleyc Barley Barley Wheat kernels Wheat kernels Rye kernels Rye kernels Wheate Wheat Wheat Corne Corn Oatse Oats Degree of Grinding Suba Glucose Fine Fine Fine Fine Fine Fine 25% Flour 50% Flour 100% Flour 25% Flour 50% Flour 100% Flour 100% Flour Cracked (bulgur) 100% Flour Pumpernickeld Cracked grains Coarse flour Fine flour Cracked grains Flour Rolled oats Flour N N D N N N D D D D D D D D D D N N N N N N N +38%** +46%*** +67%* +48%* +6% ns +80%+ +5% ns +26%* +39%* 0% ns +59%* +146%* +52%* +3% ns +89%* +66% +7% ns +34% ns +32% ns −21% ns −3% ns +29% ns +75% ns % Changeb Insulin +138%+ + +95%*** +46%* +54%** −11% ns +16% ns +42%* +95%*** +18% ns +89%** +53% ns +19% ns GIP +60%+ +54%* Ref. each kernel broken into 6 pieces.11. 50. 50. 50. . flour. + = p < 0.2% retained on 710-µm sieve. 50. D = diabetic. ns = not significant. May 6. 025. 141 –39%* (8) 142 0% ns (12) –28% + (12) +5% ns (13) –8% ns (22) 143 144 145 –31%* (7) 146 0% ns (8) 147 –31%* (8) –31%* (8) –20% ns (8) . Salad and apple 96 100 +4% 1. canned Home cooked vs. Pinto beans 50. boiled 15 min Boiled 5 min vs.01. 20 min Boiled 20 min. boiled 25 min Boiled 5 min vs. Parboiled rice 50.2387_ch4. boiled 15 min Boiled 11 min vs. raw Roasted vs. canned Home cooked vs. canned Home cooked vs. Cornflakes. bread. Rice and corn 110 85 –22% 1. May 6. High-fiber 1. canned Home cooked vs. D = diabetic. boiled Roasted vs. Spaghetti 50. Parboiled rice 50. Tested in different groups of nonstandardized subjects. + = p < 0. Table 4. Polished rice 50. N = normal. banana Cornflakes. Corn starch 50. 133 133 +133%** +8% ns 0% ns +100%** 137 138 138 139 139 95 95 95 96 140 140 116 108 108 108 108 108 Sub = subjects. White beans 50. ground and dried 12 h at 250°F vs. juice Potato. juice Griddle cakes.14 Utility of Glycemic Index of Foods in Predicting the Relative Blood Glucose Responses of Single Mixed Meals Meal GI GI 89 56 –38% 1. Spaghetti 50. Green lentils 50. Baked beans 50. 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM Table 4. boiled 16. *** = p < 0.11. ns = not significant. Spaghetti 50. canned N N % Changeb Glucose Insulin +24% ns +78% + N +160%** N N D D D D D D N N N D D D D D +15% ns –2% ns +196% ++ +3% ns +43% ** +24% ns +22% ns +2% ns +10% ns –22% ns +50% + +86%* +139%** +16%** +36% ns +23% ns Ref.11_fm Page 339 Sunday. Carrots 50.001. White bread 2. boiled 20 min Raw vs. boiled 20 min Boiled 5 min vs. Chick peas 50. * = p < 0. Whole grain bread 1. Kidney beans a b c 339 Cooking/Processing Procedures Suba Boiled: 1 h vs. Red lentils 50. boiled Raw vs. 2.005. canned Home cooked vs. boiled 22 min Home cooked vs. Low-fiber 2. juice % Differencea BG–NIDDM BG–Normal BG–IDDM Ref.5 min Boiled 11 min vs. Potatoes 25. Plantainc 50.11.05. ** = p < 0. boiled 15 rain Boiled 5 min vs. Brown rice and beans 117 74 –37% 84 85 +1% 113 90 88 99 –21% –27% –12% Text Meal 1. Flummery 2. + + = p < 0. Red lintils 1 g/kg.13 Effect of Cooking/Processing on Metabolic Responses Carbohydrate (g) and Source 50. 4. Change in incremental area under the curve or peak rise. Formula 2. boiled 20 min and ground Heated in water at 68 to 70°C until gelled vs. 3. Potato and sucrose 2. Yamc 50. Candy bar. 152. Mashed potato. for studies 147 and 148 in (102). 3. Cornflakes. peanuts 1.2387_ch4. Sweet potato. Glucose. 5. 3. BG = blood glucose response incremental area. 5. The meal GI calculations for studies 141 through 146 are presented in (159). rice Potato starch. Beans. for study 149 in (160). 2001 6:53 PM 340 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Potato meal 3. cherries 92 66 53 –29% –42% 1.14 (Continued) Utility of Glycemic Index of Foods in Predicting the Relative Blood Glucose Responses of Single Mixed Meals Text Meal GI % Differencea BG–NIDDM BG–Normal 1. Mashed potato meal 2. Spaghetti meal 3. “All Bran. 4. Spaghetti meal 75 67 60 –11% –20% 1. 4. Potato meal Bread meal Rice meal Spaghetti meal Barley. Figures in brackets = number of subjects. 2. 3. rice Fructose. Potato meal 2. NIDDM = noninsulin-dependent diabetic subjects. potato chips Raisins. for studies 152 and 153 in (161). Meal GI for references 141 through 149. barley meal 1. and for study 156 in (162). IDDM = insulin-dependent diabetic subjects. Lentils meal 71 48 34 –32% –52% 105 41 –61% –61%* (5) 95 68 –28% –24%* (11) 110 105 87 75 −4% −21% −32% −7% ns (10) −2% ns (10) −51% ns (10) 69 69 65 60 40 38 0% −6% −13% −42% −45% −27% −15% −30% −40% −53% 1. 4. lentils meal 97 86 76 65 47 –10% –22% –33% –52% –13% ns (6) –26%** (6) –44%** (6) –56%** (6) 1. Potato meal Rice meal Spaghetti meal Lentils meal 123 89 77 62 –28% –37% –50% –23%** (8) –31%** (8) –43%** (8) 1. 148 –78%* (10) –60%* (10) –56%* (10) –25% ns (10) –55% –16% –6% –8% ns ns ns ns (10) (10) (10) (10) 149 150 151 –5% ns (12) –9% ns (12) –11% ns (13) –44% ns (13) 152 –12% –41%** (7) 153 –49%***(7) 154 –13% ns (9) –19% ns (9) –6% ns (6) –6% ns (6) 155 156 157 158 (8) (8) (8) (8) (8) Percent difference from meal 1 for each study for GI = glycemic index. 5. 3. rice Sucrose.11_fm Page 340 Sunday. peanuts Banana. 2. Rice meal 2. 3RD EDITION Table 4. 2. Bread meal 2. 6. tea Cola. rice Wheat starch. The GI values for the meals in study 157 have not been presented previously. May 6. orange 3.” juice a Meal GI 1. 4. 156. Lentils. . 153. juice 2. Lebanese meal Western meal Chinese meal Indian meal Italian meal Greek meal BG–IDDM Ref. 2. 4. rice 111 53 82 92 94 –52% –26% –17% –16% –46%* (12) –23% ns (12) –7% ns (12) 0% ns (12) 1. GI values have been adjusted for unequal carbohydrate in meals. bread 2. and 157 were not reported in the original references. 2. 3. Spaghetti meal 80 66 –17% 1.11. ns). Figure 4.14.18 × ED. n = 41.001). dotted line = line of identity.168.01 × % difference in GI). 2001 3:04 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 341 Figure 4. r = 0.11.3 + 1. r = 0. observed difference (OD) = percent difference of glycemic response of each meal from that of the meal with the highest glycemic index. Dashed line. solid line.6 Prediction of mixed meal glycemic response using the glycemic index using the data from Table 4. n = 9.11_fm Page 341 Tuesday.690.11. dotted line = line of identity. closed circles = regression equation of data from 11 other groups worldwide (OD = –6 + 0. May 8.11. p < 0. .88 × ED. open circles = regression equation of data from the Stanford group (OD = –13 + 0. Expected difference (ED) = percent difference of the glycemic index of each mixed meal from that of the meal with the highest glycemic index.150.5 Prediction of mixed meal glycemic response using the glycemic index by Wolever et al.155 Solid line = regression equation (% difference in glycemic response = 3.2387_ch4. May 6. TG = serum triglyceride. * = p < 0. GTT = area under blood glucose curve after a standard oral glucose load (G) or while actually eating the diet (D). D = combined ID and NID. NID = noninsulin−dependent diabetes. ID = insulin-dependent diabetes.NF OPD OPD OPD OPD OPD 7D 10D 10D 7−10D 7−10 10D 10D 10D 4W 6W 6W 40W 12M 12M 13M LENc IGt N N UD TD IGT D ID NID NID NID NID NDN IDC ID SUBd –15%* –2% ns 0% ns +7% ns –13%+ –10%*** –2% ns +6% ns –10%* +2% ns –14% ns 0% ns +3% ns +1% nse 0% ns G –8%* D +3% ns D +36%*** D +11%+ D +65%*** G –7% ns D 0% ns D 0% ns +5% ns –2% ns 0% ns +2% ns –2% ns ns –11%* –6%* – 21%*** 0% ns –2% ns ns 0% ns +33%* –17% ns ns +19% ns +11% ns +25%*** +36%** +42%*** +49%*** % Change on High-Carbohydrate Dietb FBG GTT CHOL TG 163 164 165 166 166 167 168 169 170 171 172 173 174 175 176 Ref. → → → → → → → → → → → → → → → Diet Compositiona FAT Comment Metabolic Effects of High-Carbohydrate. NDN = newly diagnosed. CHOL = serum cholesterol.NF OPD OPD MD.NF FD HD. MD = metabolic diet. ** = p < 0. OPD = outpatient diets. 3RD EDITION .e d c b a 77% 55% 60% 85% 85% 85% 53% 60% 60% 60% 55% 56% 54% 58% 64% 36 45 41 40 40 40 37 41 47 40 39 43 40 33 42 → → → → → → → → → → → → → → → 16% 30% 21% 0% 0% 0% 30% 20% 22% 20% 29% 28% 26% 29% 20% MD. TD = treated diabetics. % change of: FBG = fasting blood glucose. UD = untreated diabetics. Subjects: N = normal.001.11_fm Page 342 Sunday. 2001 6:53 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. M = months. *** = p < 0.15 2387_ch4. Change in HbA1c.11. Study length: D = days. Low-Fiber Diets 342 44 40 40 45 45 45 42 41 35 40 42 41 40 53 41 CARB Table 4. % of energy as carbohydrate (CARB) or fat on low → high carbohydrate diet. + = p < 0.05.NF FD FD FD MD.01. NF = normal foods. IGT = impaired glucose tolerance.025. IDC = insulin-dependent diabetic children. W = weeks. noninsulin-requiring diabetes. FD = formula diet. 11_fm Page 343 Tuesday. postprandial blood glucose (PPBG). 2001 3:05 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM Figure 4.11. and serum cholesterol (CHOL) and triglyceride (TG) levels (mean ± SEM of data from Table 4.11. .18). Right: effect of decreasing dietary glycemic index with no change in carbohydrate or fiber intake on glycosylated hemoglobin or albumin (GlyPr). May 8.15).2387_ch4.7 343 Left: effect of increasing dietary carbohydrate with no change in fiber on fasting (FBG) and postprandial (PPBG) blood glucose and serum cholesterol (CHOL) and triglyceride (TG) levels (mean ± SEM of data from Table 4.11. 2001 6:53 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. ** = p < 0. PPBG = postprandial blood glucose levels. Grams dietary fiber per 1000 Kcal. Study length: D = days. % change of: FBG = fasting blood glucose. ns. TG = serum triglyceride. W = weeks. Percent of calories from carbohydrate (CHO). protein (PRO). Crude fiber. + = p < 0. Subjects: D = diabetic. 21 17 19 23 16 47 59 44 44 53 30 30 40 30 47 46 30 FIBER SUBc 344 a 17 17 20 15 17 13 53 53 39 55 35 40 53 CHO LENb Metabolic Effects of Increasing Fiber Intake from Whole Foods with no Change in Available Carbohydrate Diet Compositiona PRO FAT Table 4.01.05. Change in HbA1c. NID = noninsulin−dependent diabetic. *** = p < 0. * = p < 0. and increase in dietary fiber (FIBER) from low to high fiber diet.11_fm Page 344 Sunday. Increase in fiber of 14 to 24 g/day. CHOL = serum cholesterol.025. and fat.005.16 2387_ch4. May 6. 3RD EDITION . CSII−ID = on continuous insulin infusion therapy.h g f e d c b 34 23 37 33 28 19 11 + 14 16 30 16 16 2 19 18 18 14 54 54 20e 56 33 36 23 42 27f 24 gg 44 41 → → → → → → → → → → → → 8D 14D 8D 10C 16D 10CSII 25NID –9% ns –34%* –6% ns –24%+ + –38%** –37%** –24%+ + –18%+ + –30%*** –20%* +1% ns –5%h ns 3W 4W 4W 8W 8W 14NID 6NID 12ID 14NID 42NID –6%* +10% ns –14% –6%** –7%h –13%** ns –15% +6% ns Fiber Increased with HIGH Glycemic Index Foods 10D 10D 15D 6W 6W 6W 6W Fiber Increased with LOW Glycemic Index Foods FBG –37%* –10% ns –76%** –19% ns –82%+ +11% ns +8% ns +10% ns –7% ns –4%+ –6% ns –26% ns –12%* ns TG +2% ns +5% ns 0% ns –5% ns –13%* –14%* –20%** % Change on High-Fiber Dietd PPBG UGO CHOL 182 183 184 185 186 177 168 178 179 180 181 172 Ref. UGO = urinary glucose output. + + = p < 0.11.001. C = diabetic children. Right: increased fiber from foods with low GI. Top: effects of increasing fiber intake with no change in available carbohydrate intake (data from Table 4.11_fm Page 345 Tuesday.2387_ch4. May 8.11. Left: increased fiber from foods with high glycemic index (GI).17). 2001 3:05 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM Figure 4. Bottom: effects of increasing both fiber and available carbohydrate intakes (data from Table 4. .11.11.8 345 Effect of high-carbohydrate.16). high-fiber diets on fasting (FBG) and postprandial (PPBG) blood glucose and serum cholesterol (CHOL) and triglyceride (TG) levels. R NID NID H NID NID ID NID ID NID NID − 13% ns − 7% ns 0% ns − 26% + + − 7% ns − 5%* + 5% ns − 15%*** − 38% ns − 23%* − 5% − 25%*** − 18%* − − − − 10% + 2% ns 19%* 12%** U-64% ns I-90%*** U-11% ns U-94%* U-80%* U-56% ns B-15% + B-20%* I-56% + I-58%*** → → → → → → → → → → 64 57 50 78 78 50 42 32 65 35 g g g g g g g g g g 14D 3W 4W 6W 6W 6W 12W 4M 4M 6M ID NID ID NID ID ID NH ID D. + change of: FBG = fasting blood glucose. TG = serum triglyceride.01. % of energy as carbohydrate (CARB). 3RD EDITION .025. H = healthy. Subjects: D = diabetic. P = pregnant. NH = hypertensive NID. PC = NID in poor control.17 2387_ch4. High-Fiber Diets Diet Compositiona CARB FAT FIBER Table 4. May 6.001. ID = insulin−dependent diabetes. ** = p < 0. HbA1c = glycosylated hemoglobin. NID = noninsulin−dependent diabetes. UGO = urinary glucose output (U). Grams of crude fiber. fat. 2001 6:53 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. unspecified or mixed types. Grams dietary fiber per 1000 Kcal. mean blood glucose on the diet (B).11. + = p < 0.05. Study length: D = days.f e d c b a 23% 21% 20% 23% 23% 20% 25% 34% 15% 41% 22 21 28 36 36 28 20 20 20 19 10D 10D 10D 2W 16D 3−4W 5W 6W 6W 6W 2M 15M 48M D D D. and dietary fiber (FIBER) on low → high carbohydrate diet. W = weeks.P PC 11% ns 18%** 1% ns 12%** 36%** 1% ns + 17%** − − − − − − − 11%** + 1% ns + 2% ns − 15%*** + 19%** − 30% + 6% ns I-65%* B-+7% B.ns U−33% ns U−14% ns R-+26% + I-6%** I-0% ns Fiber and Carbohydrate Increased with HIGH Glycemic Index Foods 20 → 54 g 20 → 54 g 22 → 65 g 5 → 14e 26 → 65 g 17 → 78 g 9 → 19f 18 → 97 g 18 → 97 g 8 → 23 g 30 → 70 g 26 → 40 g ? → 25 g − − − − − − − − − − − − − − − − − − − 10%*** 22%* 14%*** 10%*** 22%* 2% ns 19%*** 16%*** 18%* 24%*** 29%*** 22%** 10% ns 14%** 15%* 15%* 4% ns 5% ns 13%* + + − − + − 10% ns 20%** 6% ns 9% ns 61%** 17%* + 11% ns − 43% + + − 15%* + 23% ns ns − 6% ns − 15% ns − 2% ns + 21% ns ns − 14% ns − 18% ns TG 196 197 198 199 200 201 202 203 204 205 177 168 187 188 189 190 191 192 192 172 193 194 195 Ref. R = chronic renal failure. M = months. ID = insulin dose (I) or rate of glucose disposal during euglycemic insulin clamp (R). *** = p < 0. CHOL = serum cholesterol. 35 49 40 50 50 40 40 43 40 45 → → → → → → → → → → 60% 60% 65% 61% 61% 65% 53% 45% 65% 40% → → → → → → → → → → FBG % Change on High-Carbohydrate Dietb HbA1c UGO/ID/BG CHOL 346 46 28 45 34 34 45 47 38 40 35 37 → 37% 37 → 30% 51 → 38% 34 → 9% 37 → 9% 42 → 14% 42 → 21% 39 → 18% 39 → 18% 39 → 32% 40 → 30% 34 → 27% ? → 13% 42 → 53% 42 → 53% 40 → 50% 43 → 75% 43 → 70% 43 → 68% 43 → 65% 40 → 61% 40 → 61% 42 → 50% 40 → 58% 43 → 55% ? → 66% SUBd Fiber and Carbohydrate Increased with LOW Glycemic Index Foods LENc Metabolic Effects of High-Carbohydrate.11_fm Page 346 Sunday. * = p < 0. FAT = fat as % of energy. children. DF = dietary fiber in grams per day. TG = serum triglycerides. PPBG = postprandial blood glucose high-GI diet vs. GlyPr = glycosylated protein (F = fructosamine. SUBJ = subjects (N = normal. NID = noninsulin-dependent diabetes.11. IDC = insulin-dependent diabetes. 24 g/d). CHO = carbohydrate as % of energy.18 2387_ch4. HC = type 2a hypercholesterolemia. adults. Significant increase in dietary fiber on low-GI diet (39 vs.11_fm Page 347 Sunday. H = hemoglobin). CHOL = serum cholesterol. PRO = protein as percent of energy. 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 347 . LEN = study length (W= weeks. May 6. but in some cases there were small differences in macronutrients. Significant decrease in dietary fiber on low-GI diet (28 vs.d c b 29 21 22 22 31 32 31 26 19% 21% 19% 19% 21% 17% 17% 21% g gc gd g g g g g 2W 1M 1M 1M 4W 2W 6W 3W 3M Diet Compositiona PRO DF LEN 6N 12HT 24HT 6HC 24N + I 8NID 7IDC 8IDA 16NID SUBJ –41(39%) –13(16%) –11(13%) –11(13%) –7(12%) –23(26%) –12(15%) –14(23%) –13(14%) Diet GI Long-Term Effects of Low-Glycemic-Index (GI) Diets H –0% ns H –3% ns F – 27%* F –22% H –11%* –13% –14%* F – 2% ns PPBG –37%** 0% ns I –14% ns I –9%* C –39%** Change on Low GI Dietb GlyPr Ins/CP CHOL –3% ns –9%** 0% ns 0% ns –11%* –9%** –9%** –3% ns TG 0% ns –3% ns –16%* +6% ns –13% ns –16%*** –19%** –1% ns 206 93 207 207 208 209 210 211 212 Ref. 61% 50% 49% 49% 60% 54% 48% 46% 45% a FAT 20% 29% 29% 29% 20% 25% 35% 36% 31% CHO Table 4. IDA = insulin-dependent diabetes. low-GI diet. Ins/CP = Insulin dose (I) or urinary c-peptide output (C). These studies were designed to have equivalent dietary composition during the high and low GI periods. N + I = NID and IDA. HT = hypertriglyceridemic. M = months). 34 g/d). Craig-McFeely.. B. Marks. Bacarese-Hamilton... 11. J.. Roseman.. 515.. Johnson.. Wolever. G. M. 41. Davison.. H. G. J. J. Griffin. D.. Guar and gastric emptying in non-insulin dependent diabetes. Guar in NIDD: effect of different modes of administration on plasma glucose and insulin responses to a starch meal. Edwards. The effect of ispaghula (Fybogel and Metamucil) and guar gum on glucose tolerance in man. Leatherdale.. M. and Alberti.. Practical Diabetes. 1985. 2. 46. N. U. Jaris. J. L.. and Bailey. 11. Gatti. and Bloom. G. A... 12. L.... D. Green. C. 1982. Williams. L. S. James. Brook. A.. 6.. G. Gassull. A. Wolever. 16. Unabsorbable carbohydrates and diabetes: decreased postprandial hyperglycaemia. M. and Barzilae. W. N. Cohen.. Metab. 3. E. Acta Diabetol. Jenkins. S.. S. Goff... 1. Sive. fibre analogues and glucose tolerance: importance of viscosity. Med.. Med. 329... 1980.. A.. Adrian. 196. Fibre and diabetes. A. D. A.. M... Levitt. The effect of guar gum on carbohydrate.. R.. 1979. M. 9. 17. 143.. and Read. N. J. M... H...... S. Sugden. Tsiolakis. G. Dietary fibres. P. D. Torri. Craigen.11_fm Page 348 Sunday.. 258.. 19. Wolever. E... Morgan. Donaldson. Hanning. C. Holgate. Effects of guarenriched pasta in the treatment of diabetes and hyperlipidemia. Taylor. Improved glucose tolerance and insulin response in obese and diabetic patients on fiber-enriched diet. Harding. W. Tomlin. A. 1984. D. Kanter. M. Walquist. 172. J. R. M. Aust. A. N. P. W.. O.. J. 1978. Nutr. Diabetologia. A.. S. J. I. 20. J. L. 281. Diabetes Care. V. J. S. Blackburn. A. 434. The effect of dietary fiber on glucose tolerance in healthy males. Nutr. 467. K.. Child. 15.. R. Dilawari. Jenkins. 9.. J.. M. R.. A. K. A. M. R... N. N. Fuessl. Absorption characteristic of breakfast determines insulin sensitivity and carbohydrate tolerance for lunch. 1982. J. L.. A. Clore. M.. G.. G. D.. T... 3RD EDITION REFERENCES 1. M. 2. V.. D. Effect of guar crispbread with cereal products and leguminous seeds on blood glucose concentrations of diabetics. The mechanism of action of guar gum in improving glucose tolerance in man. Redfern... and Jackson. 14.. Obstet. Bond. The effect of dietary fiber on glucose and hormone responses to a mixed meal in normal subjects and in diabetic subjects with and without autonomic neuropathy. 1984. J. Brit. M. 1981. J. G. and Jackson. 1976. 505. Lancet. The effects of unabsorbable carbohydrate on gut hormones: modification of postprandial GIP secretion by guar. Nineham. and Jenkins. and Snook. V. 1980. . and Marks. T. S. 1982.. W. G... L. 85. liquid food and both.. A. K. and Schwartz. 514.. Denegri.. and Morgan. T. N. Med. Tredger. 66. Diabetes Care. Refhman. E. Jarjis... E. Viscosity of food gums determined in vitro related to their hypoglycemic actions. 1987. 3(5). and Evans. I. 154. P. I. J. 21. M. J. Effect of dietary fiber on the oral glucose tolerance test in pregnancy. Jenkins. 1248. Brit.Z. Viscosity differences between various guar gums. 3. J. T. O. Clin. V. 8. J. A. Johnson. A. J. Dietary fibre supplementation of a normal breakfast administered to diabetics. A. Ist. May 6.. Y. Madden. G. B. A... M. Leeds. Nineham. and Sirtori. J.and protein-stimulated gut hormone secretion: modification of postprandial gastric inhibitory polypeptide and gastrin responses. Alberti. 1979.. Brit. E. D. J. C. D. 53. T.. Goff. L. K. Camisasca. Sci. 51. N. D. Bariascini. Scarpello. Morris. Blackburn. S. Vinik. A. H.. S... 16. 371. Redfern.. P. R. Sci. fat.. V. R. R.. I. G. 5. 612. Metz.. C. Afr. A. Gondler. Littlejohn. Tredger. 1. Am... Guar gum and reduction of post-prandial glycaemia: effect of incorporation into solid food.. Wolever. Catenazzo. Annu. 4. R.. M. Clin. Barker.. 18.. S. Diabetologia. Morgan. M. I. 1980. Leigh. Nutr.. D. 379. Smith. Guar biscuits in the diabetic diet. K. N.. 28.. 1988. Nurr. 7.. J.. U. A. T.. and Blackard.. H. Leeds. J. 2001 6:53 PM 348 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. D. M. Blackburn.. J. W.. K. S. A. and Alberti.. 20.. J. Nutr. Diabetologia. 19. Gynecol..... J. T. 1392. 1979. J.. Eitan. and Alberti. and Read. Fielden. T. R. H. Levitt.. W. Craddock.. D. D. 1980. Brit. P. M. 1984... L. and Hockaday. N. and Read. 1. Lancet.. A.. W. J. J. 1979. J. N. M. Gabbe. J. Jenkins. 17. O. and Jackson. 1986. 755. S.. Nestler. W. 61. E... P. 18. 72. 1. 339. 13. B. K. R. O’Connor. Jenkins.. T. S.. Med. Kwasowski. 10. Gassull. Lat. S. T. P. A.. A...2387_ch4. T. T. M. A. N. Am. Med. J. C. Brit. H.. Lack of effect of refining on the glycemic response to cereals. McMurry.. Influence of indigestible fibers on glucose tolerance. and Combeaux. Carter. and Jenkins.. Monnier. R. I. 1978. Scand. Hamberg. A. and Josse. Hanssen. Gut. S. Diabetes Care. C. J. Nutr. 1977.. H.. Sheard. Diabetes Care. J. 37.11_fm Page 349 Sunday. Clin. Hall.... G. 577. D. Am. Ardvidssonlenner. J. J. H. 1989. Ralphs. McCourt. Metab. Nutr. 29... M.. J... M. J.. Aguirre. 36. Lancet. 1982. Colette. J. J.. H.. Reduction of post-prandial insulin needs by pectin as assessed by the artificial pancreas in insulin-dependent diabetics. T. G. Vinik. 50. M. Dilawari. G.. and Gassull. The effect of Metamucil on postprandial blood glucose and plasma gastric inhibitory peptide in insulin-dependent diabetics.. Sarson. D.. J.. Leeds. J. Carlstrom. Jenkins. S.. T.. V. and Lo. S. Proc. 8.2387_ch4. G. 44. C. A. Slavin..)... and Blendis. insulin.. L.. S. 217. 656 (Suppl. G. Heading. F. 1982... J. 1981. F. 1988. A.. 574. Pectin: an examination in normal subjects. Pectin in the dumping syndrome: reduction of symptoms and plasma volume changes. R.. R. L. Rep. Effect of dietary fiber on complications of gastric surgery: prevention of postprandial hypoglycemia by pectin. T. Lasichak. A. Diabetes Care. Clin. 45. Rumessen... J.. Restored synergistic entrohormonal response after addition of dietary fiber to patients with impaired glucose tolerance and reactive hypoglycemia. T. Wolever. Leeds. 24. P. Blood glucose response to pea fiber: comparisons with sugar beet fiber and wheat bran. D.. 1979. S. Vaaler. B. The effect of bran on glucose kinetics and plasma insulin in noninsulin-dependent diabetes mellitus. Acta Med. Gastroenterology. Metz.. L. F. P. Sartor.. and Tothill. S. glucagon. 155.. A. Scand. D.. 1980. Clin. A high-wheat bran diet in insulin-treated diabetes mellitus: assessment with the artificial pancreas. and Merimee. Metz. and Alberti. T. A. G. K. Col.. Peterson.. 168. O. L. G. 1982. 25. Barker. L. Pectin and post-gastric surgery complications: normalisation of postprandial glucose and insulin responses.. 50.. Jefferys. 212. Diabete Metab. Ebihara. M. Laine. and Kiriyama. Vaaler. J. somatostatin. 33. S. Lee. The effect of dietary fibre on the response to orally administered glucose.. Diabetes Care. R.. F. Ebied. Glucose and insulin response in diabetic subjects: acute effect of carbohydrate level and the addition of soy polysaccharide in defined formula diets. K.. S. 215. 596. K. L. C. L. 24.. Jenkins.. and Tchobroutsky. 32.. 26. Nutr.. a water-soluble dietary fiber on plasma glucose and insulin responses in young men undergone (sic) glucose tolerance test. 520. 1. Plasma glucose and insulin responses to orally administered carbohydrate-rich foodstuffs. A. S. 1980. 1974. J. and Aagenaes. 28.. 1980. Jenkins. T. and Goetz. 83. 1981. S. 1980. Hanssen... Blood glucose and insulin levels in normal subjects following a meal with and without added sugar beet pulp. Storey. Soc. and triglyceride in obese diabetic patients. R. Acta Med. Effects of soy polysaccharide on postprandial plasma glucose... G. T. 3. A. Wolever. Orsetti.. M.. 40. Wong. Glycemic response to wheat products: reduced response to pasta but no effect of fiber.. 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 349 22. H. J. Masuhara.. Gold. C.. 43. D. Thomas.. 7. M. 72.. V.. Leeds. 1983.. F. 211. R. Lancet. G. A. Poynard. 33. . R. 30. H. Nutr.. G. Taylor. L. B. Gassull. Tsai.... F.. Metz. Aguirre. pancreatic polypeptide. K.. Nutr.. Slama. C... R. Orsetti. Diabete Metab... C. D. A. Int. 23. 1075. 7. Fielden.. E... J. 1984. 1. G. 324. and Schersten. P. Am. Albuquerque.. P. 509. 31. 1980. 1987.. 3. C. B. S.. Nutr. S. Am.. D. and Baumgardner. 23... O. 208. 1981. and Gudmand-Høyer. M... 4. 1981. R. B. 237. L. Diabete Metab. R. O. H. M.. Effect of Konjac mannan. Diabetes Care. J. Effect of method of administration of psyllium on the glycemic response and carbohydrate digestibility. Jenkins. H. 41. Nutr. 169.. 6. M. Jorde. G. Holt. A. Scand. 42. R. D. S.. R. and Burhol. and Hetenyi. Pham.. 11A. 27. 187. 1. 1048.. A. A. M. 35. D. L. Wolever.. R. L. D. 48. Bloom. in press.. Effect of different kinds of fibre on postprandial blood glucose in insulin-dependent diabetics..... M.. D. and Mirouze. and Marks. May 6. 8. A... 636.. Tredger. N. Bolton. Jenkins. D.. Prescott. J. Effect of gel fibre on gastric emptying and absorption of glucose and paracetomol. 34. 38. Dietary supplementation of fiber (lunelax) as a means to reduce postprandial glucose in diabetics. and Aagenase. Chao. A. E. Monnier. E. 39. J... L. A. L.. 51. C. Am. 1981. Acta Med. and Dilawari. M. Diabetes Care.. Florholmen. Spadafora. 21. L. Eshuis. A. Vuksan. B.. 389.. B. Wolever. Otto. and satiety.. and Niklas.. 36C. Munoz. Carroll.. Carbohydrate tolerance in man after six weeks of pectin administration. Clin. T. Glycaemic index of South African foods determined in rural blacks — a population at low risk to diabetes. T... R. Treatment of diabetes with guar gum... Metz.. The role of dietary fiber in satiety.. M. A. Alberti. glucose. S. B. H. blood pressure. Effect of a modified guar gum preparation on glucose and lipid levels in diabetics and healthy volunteers. Metabolic effects of dietary fiber from dehulled soybeans in humans. and Bron. Eur. 1981. M. Brit.. Am. J. and insulin. H. P. Molsted-Pedersen.. J. Johansen. 137. M.. Anderson. Laube.. 1977. 505. 3RD EDITION 45. Clin. 1983. Pediatr. R. 292... Nineham. G. G.. 1990. J. J. Guar gum and glycemic control of pregnant insulindependent diabetic patients. 93.. J. Apling. Effects of guar on plasma viscosity and related parameters in diabetic children. Kestin.. A. Nestel.. M. W. Moss. Glycemic index of foods: a Physiological basis for carbohydrate exchange.. Leeds.. R. and Nestel. Long-term evolution of the effect of bran ingestion on meal-induced glucose and insulin response in healthy man. 661... D. 215. 59. Am. Klimm. and Brauning. Food form.. 52. P.. post-prandial glycemia. Guar gum is not a panacea in diabetes management. Guar and its effects on metabolic control in type II diabetic subjects.. and Walker. and Holm. Scand.. Kay. Wahl.. 87.... H. Lancet. Sandstead. V. H. A.. Diabetes Care. Diverticular disease: three studies.. R. Differences d’action sur la glycemie d’aliments contenant des hydrates de carbone. Hum. P. S. S. Bowling. M. Diabetes.. 1.. Nutr. Clin. A. R. R. Murphy. Eastwood. 51. M.. Am. J. S. S. A. 152. 738... K. Proc. M. Osilesi. K. M. D. Heaton.. J. Trout. Am. P. and Hegewisch. 597. H. Am. Atherosclerosis.. and Burroughs. Bolton. Effects of dietary fiber on glucose tolerance of normal men. D... D. Guar crispbread in the diabetic diet. 34. H.. J. 34. 1981.. and Anderson. L. Nutr. A. E.. Jenkins. E. Harper. Glover.. 1984. J. M. Heaton.. J. M. Beck. M. Nutr. L.. K.. V. and Mako.. 64. 496. Nutr. G. Clin. E. M... Metabolic effect of bran in patients with diverticular disease. A.. McLean Ross. and insulin: studies with fruit and fruit juice. B.. . and Dilawari. and glucose metabolism in mildly hypercholesterolemic men. 1985.. Relationship between changes in GIP concentrations and changes in insulin and c-peptide concentrations after guar gum therapy.. Smith... 60. Nutr.. 1984. Jacob.. R. B. J. T. Clifton. D. Clin. J. M... 63. Jenkins.. L.. L. Lab. J. E.. O’Dea. Knick. Clin. H. 7.. 57. O’Dorisio.. 46. W. Dykes. Clin. T. Brydon. and Burroughs. 53.. G. 58. 1744. L.. R. F..2387_ch4... Haworth. Hockaday. Clin. 1. A. 1982. F. Johnson. A. Consequences pour le traitment dietetique du diabete sucre. D. L. Irsigler. 362.. NZ Med. 1977. M. and Tartt. D. J.. and Antonoff. T. M. K. J. 54. 2. 68. T. and Cummings. H. 7... 368. F. 1981. J. 67. Am. K. Kuhl. 50. R. Groop. U. 679. Taylor. D. A. Groop. L. Najemnik. W. A. 1977. B. 215. K. 61. J. P. K. 38. J. Villaume. 60A.. A. Bacon. S... 1980. Comparative effects of three cereal brans on plasma lipids. H. T. Am. Baldwin. Nutr. Hyg. 40. 1983. B. T. Med.. Behall. R. J. 1978. D. Brodribb. J. G. Nutr. Wolever. C. J. Soc. 1979. H.. Totterman. Clin. 1981. Barker. Med.. D. Newman. 37. Depletion and disruption of dietary fibre: effects on satiety. and Humphreys. D. Gariot. A.. 3424. Decreased urinary glucose excretion and plasma cholesterol level in non-insulin-dependent diabetic patients with guar. R.. 52. R. 2001 6:53 PM 350 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. L. Part III. Clin. 31.. 46. E. 65. M. A... S. Vollmar. 47. Am. 62. J. 33. Am. O. R. and Debry. C. J. F. D.. Bacon. 1–11.. 55. 56. and Hornnes. 1986. Haber. 428. Koepp. Brit. G. W. J.. Fielden. 38.. Pometta. Walker. C. L. 42. 1023. J. Physical factors influencing postprandial glucose and insulin responses to starch. Taylor. Desalme. 760. Nutr. L.. J. K. Koellreutter. Jenkins. 1980. D. Kritz.. H. R.. 211. Nutr. B. Use of xanthan gum in dietary management of diabetes mellitus. 28.. W. Clin. M.. P.. and Goff. C. A. Diabete Metab. 36... 779.. Jenkins. Houston. C. Leeds. A study of the effects of dietary gum arabic in humans. J. 1984. 48. 2. Diabetes Care.11_fm Page 350 Sunday. J.. D. C.. Med.. J. Nutr.. 1981. and Hodgson. P. Wolever. Jenkins. H.. 1983. A. K.. P. Nut.... L. Reimann.. 1978. C. 31. R. 2. Invest. plasma-glucose. and Hockaday. H.. Bekhechi. G. Lancet. J. Hinks. M. P. P. 45. Koh... 6.. 49. Schweizer. May 6. 1976.. H. 66.. H. L... Nutr. and Fyhrquist. S. F. 376.... T. soy hulls. Monnier.. 1981. J. M. P. T. Nutr. J. 151. D.. E. J.. 49. M. Wolever. The influence of guar-gum brerad on the regulation of diabetes mellitus type 11 in elderly patients. Aimellini. J. J. H. Med.. 1147. Nutr. Gormley.. The effects of a bulk laxative (metamucil) on fasting blood glucose. M.. R... G.. Effects of dietary fibre supplementation in stable and labile insulin-independent diabetics. 73.. Effect of consuming fiber from corn bran. S. Lee.. K. M. 59. S.. Practice. and Simell. P.. A.. E.. 1988. U.. H.. Doi. L.. Sels. Current Therapeutic Res. plasma lipids and gut hormones in non-insulin dependent diabetic patients. 1978. J.. Reichert. Intern. Brit. K. 1984. J. and Siitonen. Med.. Ann. Ir. and Kiovisto.. D. J... J. R. 82. 257. 1980. Nineham. low fat diet confer any benefit to the management of newly diagnosed overweight type Il diabetics?. Am. Brit. M. 1992.. 345. Med. Am.. Blotman. and Boden. A. Beattie. 76. Korhonen. J. Eberling.. 1981. Steemson.. Kawara.. Cullen. C. Kalmusky. Miranda. J. T. R.... Ward... Taylor. 20. P. Lousley... Scand. Sinisalo. 1983. Bosello. 57. Inman. M.. and Hockaday.. J. J. Penttila.. Soc. T... Nutr. A. Yki-Jarvinen. Treatment of diabetes with glucomannan (Konjac mannan). D. D. 98.. J. P. and Ludovico. M.. J..11_fm Page 351 Sunday.. 29. and Mirouze. D. Diabetes Care.. R. . K. 1980. 1. D. 333. Fuessl. T. 1981. M. M. 1988. Longterm effects of dietary fiber on glucose tolerance and gastric emptying in noninsulin-dependent diabetic patients. and Taylor. M. Diabetes Res. Ray.. 72. 1987. Flendrig. 1979. 88. Slaughter. Jenkins. J. Clin. Mahalko. Glucose tolerance and blood lipids in bran fed patients with impaired glucose tolerance. May 6. Metabolic and nutritional effects of long-term use of guar gum in treatment of noninsulin-dependent diabetes of poor metabolic control. Jelfs. Reynolds. A.. 37. 39. and Slabbert.. O.. D. P. R. J.. Leong. Does adding fibre to a low energy. Carter. T.. H. I. O.. A. Savolainen. Glucose and lipid metabolism and insulin sensitivity in type I diabetes: the effect of guar gum. K. and O’Moore. P. Matsuura. A. Guar sprinkled on food: effect on glycaemic control. Salmon. M... high carbohydrate. R. 48. L.. A. Williams. Fagerberg. T.. Thorne.. Stenman. M... 75. Sandstead. 46. Warner. V. J. Monnier. Micciolo. H.. A. and Postmes. E. L. 79. 21.. Aust. A. Diabetic glucose control. H.. The effect of apple fibre on diabetic control and plasma lipids. S. L. P. and other variables in constipated patients with noninsulin-dependent diabetes. 24.. The glycaemic index of foods tested in diabetic patients: a new basis for carbohydrate exchange favouring the use of legumes. Diabetes Care.. 482. A. A. Glycosylated haemoglobin. and Wong. S. J. H. Mayne. N. 1. serum lipids. J.. W.. C.. Edwards. Beneficial effects of palatable guar and guar plus fructose diets in diabetic children. K. R. J. T. Helve.. 71. Darling. S... R. C. K.. E. 1987. Jenkins. S.. 25. Johnson. 296.. O.. J. Clin. J. R. E. Al-Hussary.. 987. Steyn. Sci.. Julian. M.. V. and Martin. Owen. J. 1980. A. D. J. or apple powder on glucose tolerance and plasma lipids in type II diabetes. R. W. Nutr. S. Knip. 70. J.. L.. Clin. I. O. R. Acta Paediatr. 83. 1989. Lancet. 87. 68. R. N. 74..... 36. W. S. R. and trace elements on long term guar. Wolever. Med.2387_ch4. and Turner. Holman. H.. 463.. F. Am. and Horwitz. D.. 78... and Bloom. 177. Improved diabetic control and hypocholesterolemic effect induced by long-term dietary supplementation with guar gum in type 2 (insulin-independent) diabetes.. S.. O. Akerblom. D.. L.. R. E. Tomplin... A. P. Silvasti. F. Vontilainen.. A.. Cohen. K. 3. F.. Jenkins. Knight. H. 1987.. The treatment of poorly controlled non-insulindependent diabetic subjects with granulated guar gum. and Baba. 86. 153. 76. Diabetologia. S. Maenpaa. 1353. H. Siitonen. Am. Botha.. 84. No glycemic benefit from guar administration in NlDDM. Ostuzzi. Hosker. Uusitupa. Med. J. J. V.. 90. 89. R. 78.. Milne. G. Aro. C. 1985. C. Hersio. and Mann. G.. 1987. Clin. and Parviainen. Diabetologia.. High fiber diets in the treatment of diabetes mellitus. 80. R. Clin.. Paganus. R. 3... 85. T.. A.. Med. H. Jones. Nutr. 59. M. Aro. G. 81. K. G. Afr.. 88.. Atkins.. 1983. P. 1.. Low-dose guar improves diabetic control. D. T. E. Diabetologia. 166. 76. R. Uusitupa. 31.. R. blood glucose and serum cholesterol levels in diabetics treated with guar gum. 546. McGill. B. 4. B. Colette... Med. J. A. Brit. Esterhuysen. M. Malmud. 10. R. Adrian. 12. lipids.. Diabetic Med. and Read. Mansell.. R. L. M. The role of guar and dietary fibre in the management of diabetes mellitus. Roy. and Haunz. I. 1987. 1982. A.. 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 351 69. M. 77. R... S. 1985. A. Clin. Hum.. Potter. M. 604. 1335. S. T. C. 135. D.... P. W.. R. A. The serum insulin and plasma glucose responses to milk and fruit products in Type 2 (non-insulin-dependent) diabetic patients. S.. Nuttall. Kamath.. M. D. U. Nuttall.. G. P. Giordano. Effect of canning on the blood glucose response to beans in patients with type 2 diabetes. P. 9. Clin. Thompson. S. Clin.. 631. J. 1986. M.. 1986. Q. Clin.. Postprandial hormonal responses to different types of complex carbohydrate in individuals with impaired glucose tolerance. 93. T. Billington. A. Lam.. Thompson. G. P. C. and Wong. Jenkins. S. 108... 1985. Sperling. J. A. G. 205. Josse. P. A.. R. J. J. D. S. Nutr. M. J. D. and Wong. M. J.. Kalmusky. S. G. J. S. Diabetes Care. M. Nutr. R. Diabetes Care. Am. 1981. 107.. D. 215. Y. P. Brand.. J. Am. Csima. Hum. J. Josse.... G. Nuttall. Am. G.. Gannon. 8.. Thorburn. Jenkins. May 6. Wong... and Mantere. L. P. H. 1986. 2450. S. Jenkins. G. Wolever. M. 1987. C. Insel. J.2387_ch4. Crapo. A. 1178.. L.. G. J.. U.. Am. A. Dilawari.. Nutr. S.. 1980. Kalmusky. Ross. 401. Glycemic response to pasta: effect of food form... Wolever.. 105. Comparison of regular and parboiled rices: explanation of discrepancies between reported glycemic responses to rice. Nutr.... A. G. K. Krall. Nutr. A... Am. R. 34. J.. L.. P. 106. Wolever. Giordano. 34. Josse. R. Kalmusky.. and Josse. C. J. S. Clin. W. 101... L. Hum. A. S. Low glycemic index food in the management of hyperlipidemia... S. T. J. 94. 29. S. S.. 295. Diabetes Care. Krezowski. 39A.. Kenshole. 1984. 6. C. Josse. 103. Wong. S. A. J. J. S. F. G.. A.. J. M. J. S. T. M. 1985. Lee. Bird. and Parker. 5.. J. J.. Appl. M.. A. and Olevsky. Wolever. 2001 6:53 PM 352 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. U. G. 6.. 43. and Raghavan. R.. A. R. J.. cooking and protein enrichment. 41C. S. S. A. Glycaemic index of South African foods determined in rural blacks — a population at low risk to diabetes. Glycemic index of processed wheat products. L. Patten. Prediction of the relative blood glucose response of mixed meals using the white bread glycemic index.. and Little. G. D. Jenkins. 108. J. The glycemic index: similarity of values derived in insulin-dependent and non-insulin-dependent diabetic patients. J. Clin.. Josse. J. Giudici. A. Nutr. K. 1723. S. Giudici.. 111. L. C. S. C. J. G... A.. Diabetes.. Jenkins.. A. 102. 1984. and Kolterman. Jenkins. Nutr. G. 95. and Truswell.. Jenkins.. R. Kalmusky. insulin and glucagon responses to different types of complex carbohydrate in noninsulin-dependent diabetic patients. P. T. Wong. Wong.. 1986. Billington. Am. J.. K. T. and Walker. U. 1981. Am. R.. 110. S. and Parker. Low glycemic response to traditionally processed wheat and rye products: bulgur and pumpernickel bread. 1983. 36C. 184. Thompson. 3RD EDITION 91. A. Res. and Olefsky. D.. L.. 38. G. U.. 1985. 40. Wolever... and Jenkins. Reid. Col. Clin... Clin. 1987. The effect of phytic acid on in vitro rate of starch digestibility and blood glucose response. M. Thorburn. Yoon. Nutr.. Giordano. J. Jenkins. 349.. G. Comparison of serum glucose. Crapo.. B. Wolever. 1981. M. 1985. S... D. Nutr.. Nutr. Effect of test meals of varying dietary fiber content on plasma insulin and glucose response.. T.. C. J. Buckley. and Lee. M. S. Thompson. C. R.. Food processing and the glycemic index. 34. Lactose in the diabetic diet: a comparison with other carbohydrates. 104. A. D.. G. Heinonen. Wong. 328... Am. G... M.. J. Hall. Nutr. G. G. A.. 1977.. and Lam.. R. 100... Josse. Clin.. N. B.. J. Wong.. R. 33. Am.. Josse. 46.. M. 26. Kolterman. and Jenkins. and Jenkins. 516. 109. Giudici. 98. Clin. Nutr. L. Nutr. Batta. C. and Truswell.. Giordano. J. Nutr.. A. A. H. Wolever. J. Insulin and glucose responses to various starch-containing foods in type II diabetic subjects. Q.. 418. T... Wolever. Clin. S. R. J. Am.11_fm Page 352 Sunday.. J. Krezowski. O. Kenshole. S.. Gannon. C.. Walker. D. Nicholson. J... Coffman.. G. Jenkins. R. 971. J. G. L.. M. 99. J. S. Giudici. Mukewar. Clin. L. Nutr. S. G. Post-prandial plasma-glucose and -insulin responses to different complex carbohydrates. 784. S. 42. F. J. . Y. Reaven. R. A.. O.. 1987.. Q.. Jenkins. 1192.. R. T. Glycemic responses to foods: possible differences between insulin-dependent and noninsulin-dependent diabetics. A. and Albrink.. Reaven.. 92.... 1987. P. Nutr. Crapo. F. J. 42. S. W. Nutr. Nutr.. L. Diabetologia. The effect of different types of Finnish bread on postprandiaI glucose responses in diabetic patients. Wolever.. 97. Res. Thompson. 96. Am. Korpela. Brand. and Wong. 835... M. A.. Reduction of postprandial plasma glucose by bengal gram dal (Cicer arietnum) and rajmah (Phaseolus vulgaris). S.. Nutr. Waldeck. A.. 10. Ravnick. S. G. Nutr. Costagliola. 1990. Effects of co-ingestion of fat and protein upon carbohydrate-induced hyperglycemia. 760. J.. G.. T. H... 114. Nutr. Clin. Gannon. W. 133. M. Brand. J. Fukayama. O’Dea. J. Siskoudis. 773.. P. T. L.. Insulinogenic and glycemic indexes of six starch-rich foods taken alone and in a mixed meal by type 2 diabetics. 113. Akanji. 449. 123. Bucalossi. D. C. The glycaemic index: effect of age in insulin dependent diabetes mellitus. Collier.. S. Whittam. R. K. 1988. J. Duodenal and ileal lipid suppresses postprandial blood glucose and insulin responses in man: possible implications for the dietary management of diabetes mellitus. Bicknell. 1982. Sci. 1980. 33.. Clin. and O’Dea.. M.. 1986. J. 1984. 363. 474. A. Clin.. Metab. 137. D. L. Billington. Camelon.. and Westphal. L. Am. Fontvielle. D. 115. A. T. Ehrlich.. and Slama. G. Welch.. Collier. Eur. 71. Nutr. and Antonoff. A. S... S.. Food glycaemic index or meal glycaemic response?. J. 121. Nutr. Effect of physical form of carbohydrate on the post-prandial glucose. M. A.. 1987... M.. Jensen. T. J... 1988. K. Nutr. G. C.. Plasma glucose and thiocyanate responses to different mixed cassava meals in non-diabetic Nigerians. A. J. Frangaki. Am. Kalmusky. Mooradian. 26. A...... Jenkins. Josse. I.. 232.. 42. 116. 50. 435(letter). A. Marsilii. F.. J. 44. May 6. 1984.. 390. J.. 282. S. K. A. Wolever. 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 353 112. Clin. 941.. Clin. A. V. and O’Dea.. Diab. Katsilambros. P. C. A. L. S. insulin and gastric inhibitory polypeptide responses to carbohydrate and protein. Chuck. 132. J. Clin. S.. J. J. Atley. Petruzzi.. Wolever. Nutr. Macronutrients have different metabolic effects in nondiabetics and diabetics. Estrich. Clin. E. 1989.. Neil. J.. S. M. 465. A.. 122. E.. G... Reichert... Wolever. G. D.. Diabetologia. and Maranes. 7. C. and Osotimehin. Giudici. C.. D. Crossman. Jenkins. 72. 1988. Nutr... E. A. Glycaemic response to carrot in Type II diabetic patients. 130.. C. M... 1985. Giordano. R.. 71.. Diab.. M. Nutr. Piazza. Schlierf. and Lee. Walqvist. P. S. S. Calle-Pascual. Metab. Thorne... D. Kofotzouli. 1983.. A.. Adeyefa. Food. S. G. insulin and gastric inhibitory polypeptide responses in type 2 diabetes. Rao. A.. 209. Inst. S. C. H. 120.. The effect of coingestion of fat on the glucose.. H. J. D. F. K. P. Nutr. Effect of protein dose on serum glucose and insulin response to sugars. A. 37. Nestel. 1986 and 41A.and low-glycemic-index carbohydrates in noninsulin-dependent diabetic subjects. . and Scala.. and O’Dea. Bordiu. Galanopoulos. and Krezowski.. Metabolism. Bruce. Philippides... Sci... Nutr. K. Pattison. F. N. 16. 1990.. Blayo. and Pulini. Differential effect of protein and fat ingestion on blood glucose responses to high. U. Gannon.. 40A. 10.. Commercial canning increases the digestibility of beans in vitro and postprandial metabolic responses to them in vivo. Nutr. Hill. Romeo.2387_ch4. J. J. D.. W. Romero... R. 143... M. Appl. R. Conti. P. Glycaemic and insulinaemic responses to different carbohydrates in Type II (NIDD) diabetic patients. 44. 1967. A. K. Foster.. Physical factors influencing postprandial glucose and insulin responses to starch. and Scarpello. Collier. and Read. 1. Am.11_fm Page 353 Sunday. G. A. O... K. R. 45. Clin. T. K. J. Vuksan.. 128. Effect of co-ingestion of fat on the metabolic responses to slowly and rapidly absorbed carbohydrates. The glycaernic and insulin indices of realistic meals and rye breads tested in healthy subjects. 124. Nutr. G.. F. 36. Bornet. 127. T. C... 126. L.. Wong. Clin.. P... xxxv. 1982. Am. Can. Am. A. L... S. J. N.. 36.. Tchobroutsky. M.. Metab. 1987.. Nutr. Kalmusky. R.. 50. and TrusweIl. Nuttall.. Spiller. A. Lombardo. J. and Francis. D. Jenkins. P. Clin. M. Jenkins. Clin... Clin. Letanoux. B.. G. Q. J. E. A. Nutr. 1985. Diabetes Res. M. Am. McLean. Thompson. 3. Diabetes Care. Haardt.. Guilliford. and O’Dea. Am.. Taylor. J. Traianedes. 118. Am. 131. O. 1987... J. Nuttall... 18. Hum. G. 1988. Nutr. 85. Gastroenterol. Diab. 119.. McDonald. Nutr.. 46. L. Am. and Outch. M. Charles-Davies.. 3. Peppas. 37. and Josse. Effect of processing on digestibility and the blood glucose response: a study of lentils. Effect of ripeness on the glycaemic response to banana. J.. Simpson. 1081. J. G.. 1093. Q. C. Martin-Alvarez. J. 1987. J. The insulin and glucose responses to meals of glucose plus various proteins in type II diabetic subjects. K. L. R.. B. 7. V... Wong. and Sfikakis... S. E.. I. 3. Collier. M. J.. J. 1990.. Diabetes... Technol. J. J. 129. and Jenkins. A. Effect of protein ingestion on the glucose and insulin response to a standardized oral glucose load.. R. 117. 125. G.. R. Effect of processing/preparation on the blood glucose response to potatoes. 588.. C. and Kinsell. Am. L. Williams.. Vaaler. Heaton. and Davidson. Albrink. Colonna. A. 7. Comparison of predictive capabilities of diabetic exchange lists and glycemic index of foods.. 1984. Med.-D. Rasmussen.. C. W. Nutr. 656 (Suppl. T.. and Onwuameze. Arnfred. Glycemic effects of spaghetti and potato consumed as part of mixed meal on IDDM patients. Enger. M. Hanssen.. J. F. A.. C. 401. V. and Parker. Clin. G.. 149. J. M.. J. 47. Giudici.. Nutr.. M.. S. G. Kraemer. Johansson. G. C.. G. Nutr. D. Wolever. Different glycaemic responses to pasta. R. 32. Dietary fiber in type II diabetes.. 154. 38. 147.... M. D. Barry. Effect of differences in source of dietary carbohydrate on plasma glucose and insulin responses to meals in patients with impaired carbohydrate tolerance. insulin.. 141. 139. C. J. and Reaven. Effect of high..-G. Am... C. Med. M. R.. 1981. Clin. S.). 6. Nutr.. J. Diabetes Care. O’Dea. and Calmiche. A. M. and Bantle. Cloarec. 144. Mazzaferri. Acta Med.. E. J.. and Rivellese. Emmett. 3RD EDITION 134. Thomas. Am. 1. Effect of source of dietary carbohydrate on plasma glucose and insulin responses to mixed meals in subjects with NIDDM. and Lundquist. G.. C. R. S.. and potatoes in diabetic patients. O. and MacDonald. T. Particle size of wheat. Clin. O.. Med. D.. and fat content as calculated using the ADA exchange lists... S. S. G. May 6. 48. J. Williams. 34. bread. A. 2001 6:53 PM 354 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 395. Jenkins. 150. W... Clin. Giacco. Hollenbeck.. J. D. Simpson.. Clin. and Collier.. Diabetic Med. and Schmitz.. 1985. J.. Marcus. F.G. Ahren. P.. M. M. C. and Wong. F. R. Am. 374. S. P. J.. 44. B. Clin. 136.... F. 153. R. Aust. J. 1032. Thomas. S.. I P. Castle... Geog. Laine. A. T. Br. Jenkins. Lack of effect of dietary fiber on serum lipids. D. M. M. A. Tobey.11_fm Page 354 Sunday.. J. 7. PostPrandial glucose and insulin responses to meals containing different carbohydrates in normal and diabetic subjects. Gouilloud. New Engl. 1279. 1980.. Effect of cooking on serum glucose and insulin responses to starch. 10. Ulirich.. A. 146. G.. Levitt. Nutr. Liu. H. 309.and low-fiber diets on plasma lipids and insulin. 40.. Diabetes Care. M. Am. 1988.. 297. G. Effect of naturally occurring dietary fibre in Western foods on blood glucose. Oli. 1983. Newman. Am. Rao.. Trop. and oat test meals: effects on plasma glucose and insulin responses and on the rate of starch digestion in vitro. and gastric inhibitory polypeptide responses to test meals in subjects with noninsulin-dependent diabetes mellitus. 1982. and Josse. DeMarais. 1981. Nutr. Csima.. McDonald. 33.. N. F. E. Diabetes Care.. J. Clin. G.. S. B. A. O. P. 421. M.. 1988. J. Clin.. A.. 1981. 675.... 432. G.. R. Wesson. Balaza. J. D. Med. 1982. J. Hermansen. G. 155.. 135. B. 137. 10. P.. S. 349. 1979. M. Am. N. S. Am.. Q. N.. 1987. M. C. J.. Jenkins. V. J... 51.. 1984.. 1981. Mooradian.. Effect of source of dietary carbohydrate on plasma glucose. T. 47.... 1983. G. 145. carbohydrate.. C. maize. J. and Schersten... Prediction of glycemic response to mixed meals in non-insulin dependent diabetic subjects. Collings. 152. Dencker. Clin. M. Wahlqvist. Riccardi. 484.. and Dunlop. Parillo. 36. Am... 34. Bantle. Nutr. 2716. Wong. 382.. Greenfield. S. Diabetes Care. and Bolton. M. H. A. Pacioni. G.. M. D. J. P.. 142. and Wong. Hoogwerf. R. J. T. I. B.. K. Brit. Kalmusky. 965. Wolever. 1987. 282.. Sartor. 2. Second-meal effect: low-glycemic-index foods eaten at dinner improve subsequent breakfast glycemic response. M. 221. C. Tokay... Coulston. glucose.. C. Coulston. R. Med. Swislocki. G. 143. NZ J.. J. Wolever. W. The rate of starch hydrolysis in vitro does not predict the metabolic responses to legumes in vivo. K. 1486. 140. I.. Nutr. I. 11.. H. J.. and insulin in healthy young men fed high starch diets. A.. J.. 148. M.. A. A. and Reaven. The glycemic effect of different meals approximately isocaloric and similar in protein. M.. L. J. I... 317... M. L. D. G. W. 1997. M. Agardh. B.. Coulston. K. and Aagenaes. W. R. Winther. Ikeakor. The effect of cooking upon the blood glucose response to ingested carrots and potatoes. Ocana. J.. 1983. P. Scand. L. D. Blood glucose responses to common Nigerian foods. Josse... Bornet. 387. F. J.. Nutr. Coulston. Collier. Clin. Nuttall. and Reaven.. Diabetes Care. and Albrink. A. .. Am. C. M. Wholemeal versus wholegrain breads: proportion of whole or cracked grain and the glycaemic response. B. 1990. 10. M. Laval. 1041. K... 1988. Pasta cooking time: influence on starch digestion and plasma glucose and insulin responses in healthy subjects.. S.. Asp. S. 138. 151.. Nutr. and Goetz. Effect of source of dietary carbohydrate on plasma glucose and insulin responses to test meals in normal subjects. Greenfield.2387_ch4. Starich. Hollenbeck.. and Reaven. A. Nyman. R.. 1986. 958. J. J. Laine. A. M. insulin secretion and blood lipids. G. Metabolism. 354. Med.. T. insulin and lipid responses to high carbohydrate low fat diets in normal humans. 359.. D. 8. A. Ed. M. Olefsky. Anderson. 1971.. Wolever... Clin. . De La Garza. and Connor. 8. 50. 30. M. Kaiser.. 45.. Effect of dietary fiber and foods on carbohydrate metabolism. Brit. 357. 170. 176. G... M. Effect of carbohydrate restriction and high carbohydrate diets in men with chemical diabetes. C. and Bierman. 2. Endocrinol. J. M. 1976. J. R. 127. 80. Clin. Pacioni.. Dis. 1982. and Mancini..). and Reaven. 235. A-L. L. Digestible carbohydrate—an independent effect on diabetic control in type II (non-insulin-dependent) diabetic patients?. B. Wolever. S. 2.. Boca Raton. I. Assal. S. Am. R. high carbohydrate diet upon the serum lipids in diabetic patients. Genovese. Diabetes. Tumini.. R. 164. J. G.. A. 171. 52. 53. A. A.. M. W. Improved glucose tolerance with high carbohydrate feeding in mild diabetes. Nutr. 169. Effect of a fat free. and Jenkins. Clin. Brunzell. H. and Baum. Persistence of hypertriglyceridemic effect of low-fat high-carbohydrate diets in NIDDM patients. 157. and Jenkins. J. Brand. 23. L. Riccardi. 165. 168. Carter. 1990. G. Separate influence of dietary carbohydrate and fibre on the metabolic control in diabetes.. 167. 26. 160. High and low carbohydrate diets in diabetes mellitus. and Ovalle-Berumen. 173. M.. Stevens. C. 1974. 167. Application of glycemic index to mixed meals. Plasma glucose. J.. W. 138. Chew. 177.. Jenkins... Col.. 1980. Emmett.. L. J..... T. A. 1986. B. 23. 51.. A. Chiarelli.. R. M. Nutr. L. S. A. Am. 42. Diabetologia. A. Santoro. Liu. corn flakes: plasma glucose and insulin responses in young females. and Mancini. 1982. 1988. 1113. D. Verrotti. P. 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 355 156. A. S. G.. Thorburn.. E. D. Stone.. Prospective comparison of modified-fat-high-carbohydrate with standard low-carbohydrate dietary advice in the treatment of diabetes: one year follow-up study. R. C. Ann. Hockaday. and Reaven. Am.. S. T. Genovese. L. Genovese. 105. 1985. Giacco. 1984. Diabetologia. J. C.. 447. and Dupre.. Am. Diabetes Care. Wolever.. A. Spiller. Arch. Jenkins. 1983. Med. Dieta alta in fibras preparada con alimentos regionales como complemento in il control de la diabetes. All bran vs.. Diabetes Care.. Coulston. P. I.. J. Garcia. 402. 1984. D. A.. 521. Invest.11_fm Page 355 Sunday. Nutr. 1989. Induction of hypertriglyceridemia by a low fat diet. T. R.... 332. 163.. D. Angus. Espinosa-Campos.. F. Nutr. and Reaven. D. M. G. Clin... 172. J. and Morgese. J.. Rivellese. Nutr. L. Seeman... G. S... Crapo. 34. 1986.. A. J.. M.. H. G. G.. N. Burgess. S. J. D. J. Csima... Lousley. Prolonged effects of a low cholesterol. J.. 178. Mann. Simpson. A.. D. Porte. D. Metab. Oettle... Coulston. Nutr. W. J. 57. M. Behme. Effects of normal carbohydrate/low fat diet on lipid metabolism in insulin-dependent diabetes mellitus in childhood. 7. S. Mattioli. D. 47. S. G. 1965. D.. E. 285. Wong.. Whole foods and increased dietary fibre improve blood glucose control in diabetic children. 86.. and Riccardi. P. Ginsberg. A. R. 1240. 1989. and Truswell. Kimmerling. T. 162. 12.. J. and Gleason. C. D. Wolever. 159. P. J.. S. M.. A. R. Perrotti. Lerner. M. P. D. Soeldner. G. P. Am. Study of effects on diabetic control. Child. J. and Josse... and Bierman. New Engl.. Diabetes. D... S. Weinsier. Rev. 175. A.. 1987.. Hollenbeck. Riccardi. 174. 161.. high carbohydrate diet on diabetic subjects with fasting hyperglycemia. J. M. Hockaday.. and Heaton. 12. Kinmonth. Glucose and insulin responses to manufactured and whole-food snacks.. E. Clin. Nutr... and Turner. 1989. 1989. R. J. L. Pancioni. M. S. Mastranzo.. A. 1974. 43. 166.. J. Lerner.. Metab. L. 32. G. Effect of digestible carbohydrates on glucose control in insulin-dependent diabetic patients.. Clin. Rivellese. M. 94.... 1977. Diabetes Care. T. B.. A. Nutr. R. M. 187. A.. M.. 87. 116. Brunzell.. Lancet. R. C. G. May 6. 235. D. Effect of dietary fibre on glucose control and serum lipoproteins in diabetic patients. 1978. Giacco. 179.. Glycemic index and mixed meals. Diab. The glycemic index: variation between subjects and predictive difference. F. and Mann. Hazard. 729. S. 1982. 158. 39. Garza. Rivellese. Clin. Smith. W.. Am. J. J. M. M... E.. M. R.. Outpatient management of diabetes mellitus with patient education to increase dietary carbohydrate and fiber. I. Henera. Swislocki. Intern. J... W. S.... (Mex.2387_ch4. FL. in CRC Handbook of Dietary Fiber in Human Nutrition... Nutr. Am. 284. K. Clin. CRC Press. Porte. Use of the glycemic index in predicting the blood glucose response to mixed meals.. and Sheppa. J. J. J. G. M. 12.. G. and Atley.... Marescotti.. Hockaday.. Massey. and Hockaday. M. (Paris). 185. N. A. 48.. Long-term effects of high carbohydrate. de Jager.. Abbott. 268.. A.. Lancet. Geekie. Coulston. Clin.. and Rivellese. 10. Pacioni. R. Asp. and Ward. Diabetologia.. Long-term use of a highcomplex-carbohydrate. lipoprotein. Henry. W. Hagander. S. K. Hageman. M. and Reaven. R. F. and Huisman. 195.. Asp.. A. Prac. 1988. A. N. 16.. 34. Diabetes Care. and Mann. 190. K. C... 184. Nutr. Silvis. J. J. Lindsay. Bruttomesso. L. C. N. W. 1984. D. D.. DeMarco. B. 1981. Eaton. M. Moore. high fiber diets on glucose and lipid metabolism: a preliminary report in patients with diabetes. 1979... J. R. Nutr. 32. and Tiengo. V.. W. and apoprotein concentrations in insulin-dependent (Type I) diabetes mellitus. A. Effects of high-bran bread on blood glucose control in insulin-dependent diabetic patients. Simpson.. 189... J... 6.. 52. and Werner.. K. Diabete Meiab. G. K. V. 63. high fiber diets for insulin treated men with diabetes mellitus. I. 194.. N. J. D. W. J. Carter. M. . Clin. Am.. The medium-term effect of natural or extractive dietary fibers on plasma amino acids and lipids in type 1 diabetes. Lavagnini. M. 2. high-fiber diets increase peripheral insulin sensitivity in healthy young and old adults. and Minaker. T.. Isernia. Rev.. G. R. 524. Am. Young. Anderson. Riccardi. C. J. Giorato. Kiehm. Nutr. 272.. H. 200. 47. Lousley. Cherny.. Connor. O’Brien. M. T... The effects of a high-carbohydrate low-fat cholesterol-restricted diet on plasma lipid. I.... Balasz. 181. 1988. and Rallinson. 6. and Hockaday. Am. Diabetologia. D. C.. G.. L.. E. Diabetes Care. Venhous. R.. V.. Diabetes Care. High carbohydrate... D. H. B. K. M.. J... N. Beneficial effects of a high carbohydrate high fiber diet in hyperglycemic men. C.... H. 43. 191. Diabetes Care. G. 35.. Hollenbeck.. B. J... J. 1752.. Clin. 1984. B. 193. 31. To what extent does increased dietary fiber improve glucose and lipid metabolism in patients with non-insulin-dependent diabetes mellitus (NIDDM)?. Effects of an increased content of cereal fibre in the diet of Type 2 (non-insulin-dependent) diabetic patients. M. Mollentzer. 255. Clin... Effects of replacing saturated fat with complex carbohydrate in diets of subjects with NIDDM. 2001 6:53 PM 356 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Hollenbeck. W. Wahlqvist. B. 895. B. Nutr. Hartog. 1990. Clin.. McDonald. W. L... 1988. D. 7.. 1986. B. R. K. Parillo. S. Self-selected unrefined and refined carbohydrate diets do not affect metabolic control in pump-treated diabetic patients. Anderson. W. C. R. Anderson. and Howard.. Briani. L. F. J. Vorster. 196.. 1. 1988.. Simpson.. Nutr.. C. 1985. J. 1978. R.. N.. 39. I. Efendic. 852. I.. Nutr. Brit.. 3RD EDITION 180. J. I.. R. Am. 197. Hallmans. 1984. L. 77.. R.. C. Nutr. J. J. 1990. J. and Lithner. High-carbohydrate. 187. low-fat diet and exercise in the gratement of NIDDM patients. Anderson. J. High carbohydrate diets in insulin-dependent diabetes. and Ward. T. L. 10.. R. 104. Vessby. H.. and Scherstén. Nilsson-Ehle.. 102. Vitale. 186. Am. high-fiber. F.. W. H. Jarrett. 149. 1979. Clin. Iovine. Simpson.. R.. Karlström. G. 48.. Brit. T. Clin. H. 1983. J. L. W..11_fm Page 356 Sunday. A. Bilardo. W. Sissons.. Hardy.. and Heaton. Barnard. W. high-fiber diet in children with type I diabetes mellitus. E. 26. Eaton. J.. 559. Gustafsson.. 1. 183. 1. Mann. J. G. and Ward. M. 192. Clin. E. P. 29.. Grundy.. 99. R. Manhire.. W. Res. and Chantelau. Nutr. Int. Hum. F. T. C. 1979. Mann... N.. J. Temporal study of metabolic change when poorly controlled non insulin-dependent diabetics change from low to high carbohydrate and fiber diet. Duner. E. 182. and Leklem. S. M.. E. Am. 1976. Alaupovic.. Fukagawa. 198. Med. High-carbohydrate. high-fiber diet to diabetic patients with chronic kidney failure. R. Nygren.. Med. Unrefined carbohydrate and dietary fibre in treatment of diabetes mellitus. Metabolic consequences of feeding a high-carbohydrate. Dietary fiber decreases fasting blood glucose levels and plasma LDL concentration in non-insulin-dependent diabetes mellitus patients. Contaldo.... Simpson. 1981. 153. 199. A high carbohydrate leguminous fibre diet improves all aspects of diabetic control.2387_ch4. D. G.... 1989. Boberg... C. Simpson. 1989. Clin.. Nutr. N. Carter.. Perrotti.. Am. 2312. 1. Diab. 523. Metabolic and haemostatic consequences of dietary fibre and n-3 fatty acids in black type 2 (NIDDM) diabetic subjects: a placebo controlled study. R. Improved glucose control in maturity onset diabetes treated with high carbohydrate-modified fat diet.. Riddle.. G. P. L. J. Metabolism. M. T. W. 188. Carter. B. May 6. G. Lithell. S. and Pritikin. J. Boyce. A. 362. Nutr. Nutr. 205. R. J.. Harding. low-fat diet. J. L.. G. Soc. C. Brand. Med.. S. Diabetes Care. Comparison of high fibre diets.. 250. G. A moderate switch from high to low glycaemic-index foods for 3 weeks improves metabolic control of Type I (IDDM) diabetic subjects.. P.. R. J.. 5. Decreased insulin requirement and improved control of diabetes in pregnant women given a high-carbohydrate... D.. Jenkins. B.... L. 214. and Caspary... S. L. and Josse. Collier. and Bailey.. V. N. M. Lectins in foods and their relation to starch digestibility. J. Nutr. 207. K.. I. Thompson. 968. Nutr.. J.. Meyer... M. R. 1988. G... Wolever.. A. W. Colagiuri. and Truswell. W.... R. 1975. 5. Sufke. D. A.11_fm Page 357 Sunday. S. 210. Effect of dietary fiber on pancreatic enzyme activity in vitro: the importance of viscosity. Roberts.. 217. 373. A. Isaksson. A. Clin. J. J. J. and Jenkins.. J. A. C. 14. 203. M. Wesson. A. G. low fat and low sodium diet for mild hypertension in Type 2 (non-insulin-dependent) diabetic patients. 208. Rea.2387_ch4.. L... S. Diabete Metab. T. Riddle. E. I. Bird. and Lauda. and Tattersall. A. R. S. Jenkins. Biol. D. 204. Ehrlich. and flexible insulin treatment for noninsulin dependent (type II) diabetics poorly controlled with sulphonylureas. Wolever. D. Heiman. 1991... J. A. Mitchell. R. Wolever.. J. Leatherdale. A.. 1985. M. C. G. S.. 206. Sci. and Jenkins. R. and Tattersall. Nutr. Diabetologia. Proc. Collier. Am. Jenkins. A. and Cousins. Guar and gastric emptying in non-insulin dependent diabetes. J.. Low. Hollenbeck. Jeffcoate. Kubicki. Griffin. S.. 41.... Y. T. Nutr. A. J. A. ionic strength. C. 216.. L. L. 529. . Metab. Leon. Buckley. R.. Giudici.. 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 357 201. U. 19.... 1987. 202. M. 1. 219. 139. K. 213. 1. Low-glycemic index diet in hyperlipidemia: use of traditional starchy foods.. V. H. J... J. 48A.. R. pH. 28. (Paris). G.. K.. Diabetes Care. Rizkalla. 66. Green.. Pacy. Lat... L... 1984. 919.. Diab. L. Csima. J. J. A.. B. and Jenkins. J. Marshall. Nutr. May 6. Am. 220. Gastroenterology. R. 1982. P. Bird.. L. C. J... Giudici. Wolever. Rao. Jenkins. Bornet. C. T. 629.. Giudici. Letanoux. J. Chem. S.. Low glycemic index foods improve long term glycemic control in non-insulin-dependent diabetes mellitus. 218. U.. D. S. Giordano. and Slama. 1987. J. A controlled trial of a high fibre.. J. L. Y. 215. D. Dodson.. Clin. A. 95. Fletcher.. D. R.. Crossman... B. 339. Yoon. Wolever. and time of incubation. A... K. and Leklem. G. L. U.. Kalmusky. 27. T. and Bordiu. J. 211.. and Taylor. R. J. Phytic acid and calcium affect the in vitro rate of navy bean starch digestion and blood glucose response in humans. L. A. Nutr. Nutr... I. A. an inhibitor of alphaamylase from the kidney bean phaseolus vuloaris. E. 293. A prospective comparison of I conventional and high carbohydrate/high fiber/low fat diets in adults with established Type 1 (insulin dependent) diabetes. F.. Relationship between polyphenol intake and blood glucose response of normal and diabetic individuals. J. Fletcher. Scott. Low glycaemic index starchy foods improve glucose control and lower serum cholesterol in diabetic children.. T. Ocana... 208. J.. M. M.. Clin.. 467. J. E.. S.. J. G. Tchobroutsky. Connor. Allen. Kalmusky. high-fiber. G. Nutr. Wong. Patten. and Zebrowska. Brit.. 209. 39. 1988. Hollingsworth. Lundquist. U.. 918. T. 1987... D. Attenborough.. B. 707. Foods with a low glycemic index do not improve glycemic control of both type I and type 2 diabetic patients after one month of therapy. 46. G. Clin. A.. 1982.... 1988. 1982. K.. 46.. S.. Elsenhaus. Y. G. Am. E. M. P. 522. P. J. S. U. Acosta.. C.. J. M. Effect of guar gum on glucose absorption from isolated loops of jejunum in conscious growing pigs. Wong. Bal. Calle-Pascual. Buckley. A. 1988. Clin. K. Blume. Metab. A. Low glycemic-index starchy food in the diabetic diet. G. 1985. Hall. S... Purification and properties of phaseolamin.. Am.. 1985. C. 46. F. The influence of carbohydrate gelling agents on rat intestinal transport of monosaccharides and neutral amino acids in vitro. Diabetologia. F.. Button. Thompson. 82. M. Diab. A.. Am.. 39. Thompson. 248. W. 11. and Ihse. David.. D. D. 212. 1988. 1982. The metabolic effects of a low glycemic index diet.. Kalmusky. Ambler. G. The effects of subject-selected high carbohydrate.. M. W. Fontvieille.. Buckley. S. B. R. and Little.. 1984. 14.. A. D. 297. Res.. R. adsorption. D. Acta Diabet. M.. basal insulin supplements. and Jenkins. Am. Rainbird.. G. 48. D. McCulloch. E.. Jenkins. Lam. A. J. V.. L. 1980. Patten. low fat diets on glycemic control in insulin dependent diabetes mellitus. Ney. 745. Peacock. Clin. and Thompson. 8030. J. Gomez. Lam. M. 59.. Clin. D. Goddard.. I. D.. P. M. Am. Diabetic diets: high carbohydrate combined with high fiber. 1988.. G.. D.. M. 11... S.. 759. S. A. Scholfield. Wolever. The effect of starch-protein interaction in wheat on the glycemic response and rate of in vitro digestion. 234. 47. J. A. 453. L. J. 50. 1989. Colonna. et al. M. Krobicka.. J. Brit. Bjork.. Med.. and Lintas. 1984.. 226. M. Am.. Nutr. N. R. Chan. A. 946. The effect of amylose content on insulin and glucose responses to ingested rice. Clin. and Hockaday.. Wolever. A. Clin. M.. Diabetes Care. Marcelli. T. Starch/Starke.. 72. J.. S. P.. H.. W. and Jenkins. C. G. C. Starchy foods and the glycemic index. Austen. Collier. Jenkins. C. Eliasson. Holm. A. T. G. and Jenkins. 243. Metabolism.. 241. J. Corey. 1991.... J. Q. 39. A.. T. M. Mechanics and regulation of gastric emptying. G. Thorburn.. R. Am.. Reaven. M. J. C. R. A. Nutr. S. M.. J. D. 13.. J. Jenkins.. S. S. M. G... A.. R. C. Vuksan.. Clin. 1990.2387_ch4. J. F. Am. M.. A. R. A. Gannon.. 22. T. J. Ostrowska. Jenkins. M.. 149. Jenkins. G. T. Moran. 1980. 224.. 3RD EDITION 221. Larsson.. M... D. Effect of starch structure on glucose and insulin responses in adults. 1988. New Engl.... Ramdath.. 1035... a simple way to reduce postpranidal glycaemia. 862. T. Del Toma. Wolever. Diabetes Care. 48. Antoine.. M. Food fiber choices for diabetic diets. A. Swallowing food without chewing. 236.. A. 23... Digestibility of amylose-lipid complexes in-vitro and in-vivo. Lees. and Nestel. Bird. 1984. A. Couet.. Nutr. 1729. C... J. W. A. 45. 43. S. Clementi. 14. G. 229. L.. D. Wong. 1983. Snow.. C.. and Jenkins. G. H. M. Jenkins... R. 242. Jenkins. Jenkins. S. Clin. M.. M. Wolever. A.. P.. Nutr. Clin.. N. M. S. H. J. L. May 6. Bornet.. L. 1987.. G. Brit. Mourot. 230. A. Fielden. Wolever. Diabetes Care. K. 10. and Truswell. M. Wolever. Thorne.. and Jenkins. 45. T. 223.. H. J. Taylor. 244. J. Cappelloni. Am. 39. Taylor. A. 1987. A. Barker. Relationship between the rate of gastric emptying and glucose and insulin responses to starchy foods in young healthy adults.. 35. Bacon.. M... H. 428. M. Ghafari. M. 34. 388. New Engl. Dietary therapy for non-insulin-dependent diabetes mellitus. Rao. J.. Wolever.. Nutr.. V. J. 37. 1989. Med. J. Nutr. 55... A. and Bowling. Love. Brighenti. J. Nutr. 232. Josse. Nineham. J.. Plant Foods. Welch. J. Clin. A... K. Jenkins. 231. Blayo. Rate of digestion of foods and postprandial glycaemia in normal and diabetic subjects... Insulin and glycemic responses in healthy humans to native starches processed in different ways: correlation with in vitro alpha-amylase hydrolysis. 1990. Rev.. and Debry. L. W. Jenkins. Nutr. 450. and Slama. Barker. Nutr. The relationship between glycemic response.. 126. L. L... digestibility. 98. 240. U.. Asp. A. Thorne. R. A. 40. Relationship between the rate of digestion of foods and postprandial glycaemia... M. Cunnane. “Nibbling versus gorging”: Metabolic advantages of increased meal frequency. R.... H. S. and Csima. S. Nutr. A.. Brand. Clin. D. The in vitro and in vivo anti-amylase activity of starch blockers... 228. T. Nutr. 1957. G. Concurrent ingestion of fat and reduction in starch content impairs carbohydrate tolerance to subsequent meals. 237. Clin. A. 321. V.. 1982.. Med. J. 33. Josse... Rate of starch hydrolysis in vitro as a predictor of metabolic responses to complex carbohydrate in vivo. J.. C. Wolever. 5. C. V. 947.. 1988. Am. R. J. Rizkalla. 2.. 1986. 239. 319. 1980. 1988. P. M. 294. Am..... I. Wong.. and Thompson. S. R. 233. Clin.. Metabolic effects of continuous feeding. Buckley.. 238. 222. J. 37.11_fm Page 358 Sunday.. D. P. 315... J. Thouvenot. S. C. 1987. 1175. and factors influencing the dietary habits of diabetics... 2001 6:53 PM 358 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Am. T.. T. S. D. O’Dea. Fontvieille. J. Rowden. J. D. Am. A.. J. Singer. Clin.. Slowly digested and absorbed carbohydrate in traditional bushfoods: a protective factor against diabetes?. J. Factors affecting interpretation of postprandial glucose and insulin areas. Am. L. E. Young. and Lundquist. Wolever.. 1990. 45. J. J. S. 1980.. Mercier. 963. G. Nutr.. Ghafari. J. and Nuttall. Jenkins. . F.. T. A. J. Read. A. Jenkins. Clin. 929. S. J. Relationship between dietary fiber content and composition in foods and the glycemic index.. J. C. I. J. Physiol. 235. A. D. and Canary. Am. and Josse. Clin. Wolever. Vuksan. V.. Wolever. T. R.. 227.. G. 1981. 225.. S.. Jenkins. J. R. Nutr. D. L. K. 667.. S. and Jenkins. S. 1987. Wolever. Am.. S. 1983. E. Law. J. Jenkins. J. D. J. Rao.. H. Thomas. D. F. M. Behall.. R. Patten. M. T. 51. A. Glycemic index in individual subjects. Diabetologia. M.. and Marcus. J. Houston. dietary fibers. A.. and Jenkins.. 83. Lüthke. 260. Starch malabsorption and breath gas excretion in healthy humans consuming low. and Prokipchuk.. A. Knowland. 1983... and Kay.. 265. and Lee. Hanefeld. and whole foods in predicting the physiological importance of colonic fermentation.. R. J. M. J. M. 536. 519. D. A. Tasman-Jones. 347. 589. L. Cohen. E.... 251. Modulation of mucosal cell proliferation in the intestine of rats fed a wheat bran diet. Am. 1979. 1980. Clin. S. J. 1987. Gray. Effects of dietary fiber on the structure and function of the small intestine. The digestion of pectin in the human gut and its effect on calcium absorption and large bowel function. Bisalli.. Jenkins. J. Gastroenterology. Dig. Thompson.. H. 1981. D. and Spengler. 82. 1983.. A. Stephen. 41.. Am. Gastroenterology. in vitro digestibility.. and Levitt. Effects of dietary wheat bran on rat colonic structure and mucosal cell growth. Fetzer. H. D. Am. 248. R. 1988. L.. and Thompson.. and Phillips. A. Am. Nutr. Effects of dietary fiber on mucosal growth and cell proliferation in the small intestine of the rat: a comparison of oat bran. Clin. 1983. Diab. Food Sci.. and Levine. U. 1988. K. M. 92. 111. M. D. Holloway. Gastroenterol. Am. L.. Branch.. Nutr. 927. 67. L. Passage of carbohydrate into the colon: direct measurements in humans. New York. Z.. A. M. C. 444. Buchwald.. W. L. J. Fate of soluble carbohydrate in the colon of rats and man. D. Amsterdam. M. 1229. 85. T. E. U. Spiller.. .. Dis. 210. J. Nutr. Pectin digestion in humans. A. 263. Jenkins. Effect of chronic intake of dietary fibers on the ultrastructural topography of rat jejunum and colon: a scanning electron microscopy study.. 1983. 1987. Gastroenterology. 1980. T. C. Colonic conservation of malabsorbed carbohydrate. E. 247. V. Jivraj. and Jones.. 250. in Medical Aspects of Dietary Fiber... J. Semipurified dietary fiber and small bowel morphology in rats. S. J. Sheahan. 81. W. Am. 1982.. Nutr. Brit. 1990..... Nutr. Clin. McBurney. 259. 1981. M. 256. S.. Plenum Medical Books. M... Nutr.. 798.. J.. J. 954. 264. L. 1158. Jenkins. Am. B. May 6. 34. O. 1989. Cuff.. 255. J. J. Nutr. J. M. Florent.. Currier. J. D. Sci. Thompson. Tasman-Jones. Res. Haddad.. D... P. L.. Am. Wiggins. Cummings. 254. and Maher. 253.. U. 1981... 1981. Wolever. B. I. Gastroenterol. M. H2 excretion after ingestion of complex carbohydrates. Clin. J. 37. and Levitt.. Ch.. D. 1986. 54.. S. A. Potential use of acarbose as first line drug in non-insulin dependent diabetes mellitus insufficiently treated with diet alone. Owen. 27. Tasman-Jones.. M.. and Schneeman. F. Nutr. 258. McBurney. H. 1976. E. 78. Southgate. G. J. Jacobs.-C.. Clin. Clin. Short chain fatty acids in the human colon.. Thompson. 262. 383. J. 22. Oct. S. 218.. Leblond. 43. 3 (Suppl. A.. L. and Jenkins. Proceedings of first international symposium on acarbose. Increased starch intake in the human diet increases fecal bulking. Bond. A. and Vahouny. P.11_fm Page 359 Sunday.. Flourié. Cummings. M. Metab. Schulze. F. A. 249. Nutr. R.. D.. Excerpta Med... Bond. S. Prokipchuk... Digestion of certain fractions of dietary fiber in humans. M. Colonic fermentation of some breads and its implication for energy availability in man. 261. Invest. T. Jacobs. Story. and guar with total fiber derpivation.. 356. In vitro fermentabilities of purified fiber supplements. S. Holloway.. Montreux. Comparison of ileal effluents. A. 246. 1982. Wolever. C. 267. Lightfoot. M. C. J.. A. B.. 37. Eds. Ileal loss of available carbohydrate in man: comparison of a breath hydrogen method with direct measurement using a human ileostomy model. 1986. H. Hirsh. 57... J. Cuff. Gastroenterol. 95. Fischer. Ed. D. Thorne.. R. J. and Hill. Gut. 252. and Rambaud. Kritchevsky. A. J. Shetty. A. Jacobs. and glycemic response.. I. J... Nutr. 257.. J. Clin. F.. J. J. Creutzfeldt.. A. and White. J.. I.. M. G. Levitt. M. P. 31. T. pectin. L. Rautureau. S. D.. Gastroenterology... Thompson.. 266. J. J. G. 2001 6:53 PM EFFECT OF DIETARY FIBER AND FOODS ON CARBOHYDRATE METABOLISM 359 245. 709. Digestibility of carbohydrate foods in an ileostomate: relationship to dietary fiber. 763. D... 51. McBurney. A.and high-starch diets. W. U. M...2387_ch4. J. V. R. 945. 1987. A. Tasman-Jones.. 1978. 477. 1).. H. C.. U. D. D. W. Am... C. and Jenkins. L. J. 37. 253. D. M. H. 115. A.. L. and Kurpad. 7. M.. C. Cohen.. Z.. Cassidy. Influence of propionic acid on the cholesterol metabolism of pigs fed hypercholesterolemic diets. Wolever. Short chain fatty acids in human large intestine. Royall. Soc. 32. E. Gut. Nutr. and Eshuis.. J. and Bowland. 1990. Clinical significance of colonic fermentation. B. Pomare.. P. J. Royall. J. P. P. J... Am.. Biol. 1989. Am. Taylor... Topping. Interaction between colonic acetate and propionate in man. E.. 1981. R. A. G. L. 1307. Thacker.. 272. F.. A. Naylor. 270. 2001 6:53 PM 360 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Proc.11_fm Page 360 Sunday. D. and MacFarlane. Anim. Sci.. Cummings.. Ann. N. T. Gastroenterol. D. 3RD EDITION 268. J. P.... T. S. 215. T. Nutr. Salomons. Q. 1984.-J. and Cheng. P. 175.. 53.. O. F. 969. 681.. 269. 61. 85. Anderson. L... W. W. Spadafora. Metab. 274... 1221. M. McIntosh. T. L. Trimble. Storer. G. 1988. Aherne. H. M. D.. 271. X... Med.. and Jenkins. Illman. M. J. Exp. Clin..... Brighenti. B.. S. J. C.. May 6. 1027. L. Chen. G. Branch. 273. R. J. Hypocholesterolaemic effects of dietary propionate: studies in whole animals and perfused rat liver. A.2387_ch4. 84. P. Wolever. 1991. Effect of rectal infusion of short chain fatty acids in human subjects. 97. D. W.. 28. Gastroenterol... hepatic and venous blood. Am. 1987. M. Can. S. Propionate may mediate the hypocholesterolemic effects of certain soluble plant fibers in cholesterol fed rats. and Jennings. Jenkins. K.. J. Wolever. and Jeejeebhoy. H. W. H. Milligan. J. D. . portal. N.. M. May 6.2387_Section 5_fm Page 361 Sunday. 2001 6:02 PM SECTION 5 Dietary Fiber in the Prevention and Treatment of Disease . May 6. 2001 6:02 PM .2387_Section 5_fm Page 362 Sunday. e.. of total protein (especially of animal protein). To exemplify the magnitude of changes. Thus. coloreds (Euro-AfricaMalay) (3 million). are related to differences in environmental factors.17 stresses (particularly those linked with urbanization and rise in income). These populations exhibit considerable differences in patterns of diseases. especially diet. and with animal foods as a whole. and fat. 2001 7:32 PM CHAPTER 5.16 extent of smoking practice. and of total fat (especially of animal fat). Indians (1 million). however. rises are occurring in all these disorders and diseases. although not all. Most of the differences. the frequency of hypertension (WHO criteria) now exceeds that in whites11 (Tables 5. in contexts of indigence. Indeed. as juxtaposed populations. Walker In South Africa there are four ethnic populations: blacks (25 million). of protein intake (especially animal protein). especially the respective moieties of animal origin. Each of these components has a variable although often powerful influence in the regulation of the frequencies of the diseases mentioned. In contrast. between them they afford probably greater contrasts. The first pattern of diet.10 Among urban black adults.1. first it must be appreciated that patterns of health and disease are determined not only by diet but also by genetic and by nondietary factors. and coloreds.. The latter include degree of physical activity.1.g. P.1. the second is associated with very high or high frequencies of diseases of prosperity (Table 5. and the availability and utilization of medical services. May 6. and coronary heart disease (CHD).4). As to the precise assessment of the protective role of dietary fiber intake.1–3 Among rural blacks. is well nigh invariably associated with very low or low frequencies of degenerative disorders and diseases (although high frequencies of infections). diabetes. with changes 0-8493-2387-8/01/$0.g. the teeth of urban black children are now inferior to those of white children. Thus. the diet of whites.14) is characterized by a relatively low intake of energy.00+$1. colon. In this respect there are great contrasts in the patterns consumed by the different populations and their segments.3).50 © 2001 by CRC Press LLC 363 .1 through 5. dietary fiber intake is low. hypertension.1 Disease Patterns in South Africa as Related to Dietary Fiber Intake Alexander R. yet their diet is high in fiber intake. appendicitis and certain cancers (e. than are encountered elsewhere in the world. and whites (5 million). conversely. Asians. it must be appreciated that the levels of a number of dietary components are powerfully correlated with level of fiber. is high in energy. Next. there are very low frequencies of dental caries.5–9 Among urban blacks. At the one extreme. breast). the latter is inversely correlated with level of fat intake (especially of animal fat).2387_ch5. total protein. hypertension.1_fm Page 363 Sunday.15 and that of the more privileged segments of the other populations. The Asian population exhibits high frequencies of obesity. CHD. the diet of rural blacks12 (which in pattern resembles that of ancestors of Western populations13. noninfective bowel diseases. May 6. Table 5. Total Carbohydrate. Frequency of occurrence related to some regional but not nationwide populations. and Fiber Intakes Rural Blacks Energy from fat (%) Energy from carbohydrate (%) Dietary fiber (g) a + + + + + + + Implies that occurrence is rare. Table 5.3 Frequencies of Noninfective Bowel Diseases in South African Populations Rural Blacks Hemorrhoids Appendicitis Ulcerative colitis Irritable bowel syndrome Diverticular disease Colon cancer a – + – – – – Urban Blacks ++ ++ + ++ + + Coloreds + + + + + + + + Indians + + + + + + + + + + + Whites + + + + + + + + + + + + + + + + + + + + + + + + + + + + + Implies that occurrence is rare.2387_ch5.1.4 Dietary Patterns Respecting Fat. Table 5.1 Frequencies of Some Diseases of Prosperity in South African Populations Rural Blacks Dental caries Femoral fractures Obesity Hypertension Diabetes CHD Stroke a Urban Blacks + + + + + –a + + + + + + + + ++ +++ ++ + + + + + + + + + Indians + + + + + + + + + ++ ++ + + + + + + + + + Whites ++ + + + + + + + + + +++ +++ + + + + + + + + + +++ + + + +++ Cancer Patterns in South African Populations Rural Blacks Lung Breast Colon Stomach Pancreas Liver Esophagus Cervix a b Urban Blacks a – –a – – – + +b + + + +b ++ + + + + + + + + Coloreds + + + + + + + + + + + ++ +++ + + + + + + + + Indians + + + + + + + + + + + + Whites + ++ + + + + + + + + + + + + + + + ++ +++ +++ + Implies that occurrence is rare.1_fm Page 364 Sunday.2 a Coloreds ++ 10–15 70–75 20–25a Urban Blacks Coloreds Indians Whites 20–30 65–75 10–20 30–35 60 15–20 35–40 60 15–20 35–45 55 15–20 Intake depends on the season.1. 2001 7:32 PM 364 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 3RD EDITION Table 5. .1.1. seasonal fruits and “spinaches” are high in dietary fiber. the staple food of the black population.18.19 There were associated changes in disease pattern. they are still far lower than those in the white population. there is little doubt that changes in intakes of fat and of fiber are particularly influential. or fruit).48 Should this be confirmed. and that if fiber exerts its anticancer effect by being fermented into short-chain fatty acids. evidence indicates that maize. the extent of the specific involvement of dietary fiber remains a subject of uncertainty. legumes.37 diverticular diseases. obesity. Heaton22 noted that there are many ways in which a high fiber intake “ought to” protect against colorectal cancer.2387_ch5.46 Perhaps starch intake matters as much as fiber intake in terms of protection from cancer. the magnitude of the alterations required is too great to win widespread public acceptance. circumstances caused diets in certain countries to change. tenfold. among church groups such as SeventhDay Adventists. Equally great problems are latent in endeavors to.g. such as those which took place from the time of our ancestors to the present. then any carbohydrate that enters the colon and is similarly fermented could be protective. it must be stressed that although ameliorative decreases in degenerative diseases can be accomplished by changes in diet. therefore. say. Briefly.40 In fairness. holistically.27–29 Additionally. it is this phenomenon which contributes to maintaining blacks’ still faster transit time51 and lower fecal pH value52 and so protects them in measure against the development of chronic bowel diseases. since a considerable amount of the starch consumed escapes digestion and enters the colon.45 He suggested that fiber is only a part of the story. is malabsorbed. while their frequencies of chronic bowel diseases have risen slightly. there are similarities in the patterns of diet and disease exhibited by ancestors of Western populations and the patterns displayed by rural third-world dwellers. having said this. it is important to keep in mind that while different sources of fiber have different physiological actions.1_fm Page 365 Sunday. However. atherosclerotic lesions of aorta and coronary vessels.34 diabetes.30.20–22 which does not lend itself to ready resolution..19. On the positive side. 2001 7:32 PM DISEASE PATTERNS IN SOUTH AFRICA AS RELATED TO DIETARY FIBER INTAKE 365 in diet.38 cancers of the breast39 and colon.19 propounders of etiological hypotheses must be on their guard against overclaiming and overblaming.35 CHD.24–26 Probably the epidemiological evidence of the character already referred to provides the most persuasive support for a relationship.42 Can this be explained? In his recent review.18.”41 A perplexing fact in the present local situation in South Africa is that among urban blacks. As evidence of this. double intake of dietary fiber. diabetes. Only a very small proportion of populations have changed their eating habits and adopted a “prudent diet. there are lower frequencies of a number of diseases32 — obesity. involving. prosperous Western populations are having enormous difficulties in seeking to reduce their energy intake from fat to that level recommended of 30%. Account must be taken of changes and of contexts. reductions in fat intake and increase in fiber intake. people with colonic polyps are unusually efficient at digesting starch. therefore.50 This implies that a variable. there were falls in dental caries. rather than the focusing of attention and explanations almost exclusively on the changes that occurred in one or more food components. . nor of people who actually have it. and also which occurred when such changes became somewhat reversed as prevailed in certain wartime populations.31 and osteoporosis.49 In South Africa. How much starch reaches the colon? This varies enormously.43 yet low fiber intakes are not a consistent feature of populations prone to this cancer.31 and among vegetarians whose diet usually contains much more dietary fiber than the diet of omnivorous eaters. where it is rapidly fermented. such as South African blacks.33 hypertension.23 as do many short-term studies on humans. constipation. possibly a large proportion of “resistant” starch enters the colon and is therefore available for fermentation. e.44. from person to person. chronic bowel diseases are uniformly rare or very uncommon. then possibly cancer might be prevented by people eating more starch and by eating it in a less digestible form. Moreover. vegetables. despite the fact of the now relatively low fiber intake of urban dwellers.36 appendicitis.18. indeed.30. experimental studies on animals support the validity of a relationship. Conceivably. in World War II. irrespective of the source of fiber (cereals. May 6. Understandably. it must be recognized that in rural Africa. and appendicitis. inter alia. In this type of context.47 According to one small study. .. 76. L. J. 1990... N. Med. P. D. F. H. Dyer. 1989. 21. III and Rigotti. J. Ferreira.. Walker. Dietary fibre: After 21 years of study the verdict remains one of fruition and frustration. Med. 6. Serv.. J. 1989. 1410. and Joubert.. P. Afr.. F.. H. 1479. Khan. and Lesser. starch and cellulose on fecal bile acids in mice. 123.. treatment. Pezold. J. 7. 162. Afr.. Nutritional value of diets of blacks in Ciskei. Hollingsworth. 7. S. S. S. 16. J.. L. A. Schettler... 1984. Med. Caderni. 1988. J. Med.. M.. ii. J. 1988. Langenhoven. 1990.. M. 48. N. A. A.. Changing patterns of food consumption in Britain.. Seedat. Fleisch. 4. 8. and Engelbrecht. 1946. D.. Cardiovascular disease during and after World War II: a comparison of the Federal Republic of Germany with other European countries. van Heerden. R.. Armstrong. Aust. Nestel. Miller. 2. 129. 109. 43. Walker.. and Gould. 3. Sucrose consumption and dental caries in twelve-year-old children residing in Cape Town. Trowell. Risk factors for coronary heart disease in the Indians of Durban. H. 26. 3RD EDITION REFERENCES 1... D. F. K... J. Med. 2395. Health Publ. 2001 7:32 PM 366 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.. S. Darmstadt.. and Joubert.. A dietary survey of freeliving middle-aged white males in the Western Cape. A. E. S. R. 3. Du Plessis. M. Med. Dolara. risk factors. and Walker. 1974. Funani. 19. Postgrad. and Jordaan. 32. 119. M. Assoc. L. Physical fitness and all-cause mortality: a prospective study of healthy men and women.. S. S. J. Walker. P. D. 29. Somers. 12.. B. Med.. and Seedat. Motala. 78.. 17. A. S. J. Bull. and Magee. Ornish. Colorectal cancer. Y.. J. 1985. Med. Prev. A. R. S. Effect of enemy occupation on the state of health and nutrition in the Channel Islands. J. A. R. and Walker. Swanepoel. E. and Hackland. Am.. Mayet. Health. 33. P. D. 2. A.993 rural Zulus. 1987.. 1984. 73. 924. 1974. 81. South Africa... Rajput. S. Adv. J. 42. B.. Steven. 20. R. The prevalence of diabetes mellitus in a large group of South African Indians. Coronary heart disease in blacks in underdeveloped populations. and Gibbons. 262. S. Food and longevity in 18th century Scotland.. C. Afr. H.. Walker. N. P. 62. Can lifestyle changes reverse coronary heart disease?. 1990. R. Fourie.. Br. 9. 1591. Dent. Seedat. M. J. Brown. J. 43. Kohl. A.. Trans. 10. K. 73. 2528. 581. 22. Some aspects of epidemiology.. 316. M. I. . 1989. Med. 998. 27. 1290. Walker.. G. Walker. 82. C.. H. L. 581.. K. M.. 336. Nutrition in Switzerland during the war. Appendicectomy in South African inter-ethnic school pupils. R. 1974.. J. Richter. J.1_fm Page 366 Sunday.. Trans. R.2387_ch5. F. C. Cooper.. Bianchini. Regression of atherosclerosis in humans. May 6. R. S. Y. L. Hyg. T. J.. K. 61. Clark. 1988. S. 13.. K. Y.. W. 431. Nutr.. 18. Arteriosclerose and Ernahrung. Afr. R. The health consequences of cigarette smoking and the internist’s role in smoking cessation. F. K. J. Med. Effect of wheat fiber and vitamins C and E on rectal polyps in patients with familial adenomatous polyposis. C. Y. E. P. Kirkeeide. B. Heart J. Cancer Inst. E. Malinow. C.. 1959. Banks. Trop. Stokes. P. 1945. Lab.. W. R. Soc.. and Walker.. 24. Schweiz. DeCosse. Gastroenterol. B. 1985. Seedat. M. Dison. E.. M. McLanahan. An inter-racial study of the prevalence of hypertension in an urban South African population. Steinkopf. B. Dietary fiber and health.. A..... T. Diabetes mellitus death-rates in England and Wales 1920–70 and food supplies... Lancet.. D. Ports. 28. The Cancer J. 184. A. 1982.. Rossouw. Heaton.. J. K.. 15. Med. Seedat. Paffenbarger. 5. 153. Dental caries and malnutrition in rural South African Black ten to twelve-year-olds. S. Blair. Med. Min.. Dent.. G. Med. 785. W. L. Nutr. 300. A. JAMA. 1982. 16.. 1979. S. T. L. M. 965. 1987. Dietary fibre. J. M. Council on Scientific Affairs. Mon. J. J. 447.. 1983. A. 23.. 16.. 65. Afr. S. Hyg. W.. K. Nutr. and Albertse.. L. 232. Seedat.. H. and Segal. Afr. 67.. Lifestyle and disease: hypertension and ischaemic heart disease in Indian people in South African and in India... A. 1989. Brand. 1988. F.. 78.. Omar.. 219. Soc.. J.. P. M... H. 1617. F. R. 8. Fanetti. and Kriebel. Walker. screening and survival. The prevalence of hypertension in 4. Am. L. Assoc. P. 11. J. Characteristics of black women with breast cancer in Soweto. 338. H. G. 1990. 353. Med. 1990. R. A.. Afr. Rev.. Wochenschr. 14. Lancet. P.. JAMA. Trop. Intern.. J. Steyn. and Walker. Aft. A. 889. P. J. S. G. Effect of dietary fat. B. F. A. Scherwitz. W. 653. Nat.. Billings. G. J. 25. 118. B.... Role of life-style and dietary habits in risk of cancer among Seventh-Day Adventists. G. R. A. Hum. J.. Cancer.. P. B. 1987. J. C. E. Filley. 1983. Clin.. Clin. S. P. 35. J. A. 48. A. Margetts. 3513. M. P. Nutr.2387_ch5. Manetsi. M. L. J. 26. Lancet. S. Dietary habits and colorectal cancer incidence in a seconddefined kibbutz population. N. S. and Walker. O. S. Segal. Rouse. J. K.. 47. Horwitz.1_fm Page 367 Sunday. Med. M. G. 47. 1992.... J.. J. Heart disease in British vegetarians. J. Segal... 33. 18. Leger. Clin. 1987.. J.. Beillin. 10.. M. Particle size of wheat. Marcus. M. J... N. L. Tlotetsi. 51. Rozen. Food Science Nutr. R. 1988. 806. Snowdon.. May 6.. Dis. Thornton. and hormone concentrations in vegetarian and nonvegetarian postmenopausal women. and Losowsky.. Lelake.. K. K. Appendicectomy and dietary fibre. Ware.. J. 1990. R. Nuir. M. diabetes. B.. Fiber and colon cancer. 511... J. 1988. S. 543. Malabsorption of carbohydrate foods by blacks in Soweto. 50. 1987. 42.. Cancer News. T.. Riedel. and De Beer. diet. Mann. Marsh. 49.. I. 48. Gastroenterology. Animal product consumption and mortality because of all causes combined. W.. 267. Nutr. C. 31. Afr. Daya. J. Med. D. 45. M. H. Intra. I. Barbosa. 2001 7:32 PM DISEASE PATTERNS IN SOUTH AFRICA AS RELATED TO DIETARY FIBER INTAKE 367 30.. 589. 1989. 17. Emmett. stroke. 1980. Burr. 1243. 48. J. A. Aft.. 185. Health aspects of vegetarian diets. fallacies. J. Nutr. 39.. A. Dryden.. H. Nolan. Johannesburg. Hämäläinen. Nutr.. F. 46. 43. The relationship among adiposity. Vegetarian lifestyle and bone mineral density... J.. R. in press. L. 49.. 80. Verardi. Cummings. Shultz. Cummings. Am. 1988.. . Nutr. Am. P.. P.. and Burr. Am. Passage of carbohydrate into the colon. C. S. F. I. Fursdon. A. M. 1988. R. J. and Bolton. 8.. T. K. L. J... J. S. A. Gorbach.. facts. 40. 1987.. Fermentation in the human large intestine and the available substrates. 601. Anderson. 48. Clin. 34. Am... N. D. Afr. Direct measurements in humans. Dietary fibre. 34.. I.. Clin. Gastroenterol. 9. Intern. J. 1986. J. and Brodribb. I. Chaffee. Symptomless diverticular disease and intake of dietary fibre. South Africa.. B. and Dwyer. Walker.. P. 830... Cancer Res. S. Nutr... 1988.. R. fermentation and large bowel cancer. and Nieman. Nutrition and cancer. A risk factor for colonic neoplasia?. Am. A. South Africa.. and Phillips. coronary heart disease. Am. B.. Clin. 1988. 67. 35. M. M. A. J.. Jacobs. 177. Sci. A. Vessey. and Walker. S. and Tabenkin. A. C. D. C.. Nutr. W. F. Naik. L... Nutr. M.. Am. R. 1991. 36. J. 712. 433. Clin.. Woods. Kelleher. 37. 6. Clin. 38. M. F. and cancer in Seventh-Day Adventists.. 837. A. A.. N. Sanchez. Anonymous. T. and Walker. 1980.. J... Clin. J.. T. V. 1975. and Bingham. 1987. Haddad. Nutr.. Adlercreutz. Phillips. Walker. H. M. 1088. Am. 32. I. Vegetarian diet and blood pressure levels: incidental or causal association?. A.and inter-individual variations in serial faecal pH values in South African interethnic schoolchildren. and Vandongen. Super-efficient starch absorption.. H. Gear. Clin. and Fagal. The role of dietary carbohydrate and fibre in the control of diabetes. 45. Am. R. 739. Nutr. B. 52. J. 747. H. P.. 48.. 675. St.. maize and oat test meals: effects on plasma glucose and insulin responses and on the rate of starch digestion in vitro.. 41. A. 1988. and Butland. P. D. L. North Am.. R. Armstrong.. C. Cancer. 32. 44. C. L. 4(l). Diet and plasma androgens in postmenopausal vegetarian and omnivorous women and postmenopausal women with breast cancer. and Englyst. Heaton... 51 Walker. Michelsen. i. 1979. S. S. Goldin. Low-fat intake with falling fiber intake commensurate with rarity of noninfective bowel diseases in Blacks in Soweto. S. Summer. Westlake. and ACS activities. Stephen. Transit time and fibre intake in black and white adolescents in South Africa.. B. J. Food Science Nutr. Dwyer.. Nutr. L. B. 798.. S. Cancer Surv. Dig. Adv. 48. Walker. A. 85. May 6.1_fm Page 368 Sunday.2387_ch5. 2001 7:32 PM . High fiber high extraction national flour became mandatory for the entire British population from 1942 until 1953. A comprehensive British government report on human food supplies from 1938 through 1958 allowed a more detailed reexamination of this problem. however. but death rates continued falling for another 5 years. he attributed falling death rates to reduced fat intakes.50 © 2001 by CRC Press LLC 369 . Trowell passed away in 1989. African diets are usually high in their fiber content but in towns refined flours.3 In 1966. large diabetes clinics in all towns. and fell until 1953. Diabetes death rates started to rise again in 1954. many grossly obese African blacks in urban streets and second.”1 No earlier reference has been traced to any connection between fiber and diabetes. This historical chapter is unchanged from the 1986 edition of this handbook. On returning to East Africa in 1970 I was amazed to see two new phenomena: first.2_fm Page 369 Sunday. Fat and sugar supplies. lasted until 1953. but in 1942.00+$1. 0-8493-2387-8/01/$0. in 1930. Cleave and Campbell republished these mortality and food data but concluded that reduced sugar intakes caused the falling death rates. sugar.2387_ch5. mainly in middle-aged and elderly groups. 55% in men and 54% in women. and fats form a large part of the diet which may contain little fiber.4 Postwar food regulations. After teaching medicine for nearly 30 years in East Africa. This reported that diabetes death rates started falling not in 1940. 2001 7:33 PM CHAPTER 5. however.2 HIGH FIBER BRITISH NATIONAL FLOUR In 1948. During the years of the national * Dr. not only until 1948. Kenya. Himsworth published diabetes mortality death rates in British women during the period of the Second World War (1940 to 1945) and the postwar food rations. May 6. I reviewed the rising incidence of diabetes in sub-Saharan urban blacks and suggested that “their high-carbohydrate low-fat diets are protective and that low-carbohydrate high-fat diets predispose. rose in 1949 to prewar levels and continued thus thereafter.2 Development of the Dietary Fiber Hypothesis of Diabetes Mellitus* Hugh C. Trowell DIABETES AND OBESITY BECAME COMMON IN EAST AFRICAN BLACKS (1930–1960) Diabetes mellitus and obesity were extremely rare diseases in East African blacks when I started treating medical patients in Nairobi Hospital. then ceased and low fiber white bread was eaten. coronary heart disease.5 This provided the basis of the hypothesis that high fiber high starchy carbohydrate diets are protective against maturity-onset noninsulin-dependent Type II diabetes mellitus. they reported improved diabetic control. and lowered enteroglucagon responses. should be fiber-rich unprocessed starchy foods. Possibly viral infection or other noxious agents damage the pancreatic cells in Type I diabetes. rheumatoid arthritis is certainly uncommon.S. They recommended increasing starch carbohydrates to 50% total calories by increased consumption of fiber-rich whole wheat and brown bread and potatoes. they are probably contributory etiological factors. NEW HIGH FIBER HIGH STARCH DIETS In the U.10 IMPROVED DIABETIC DIETS In 1979..11 In 1981. diabetes. bread consumption rose about 25%. One overall aim was to reduce fat consumption and hopefully in the long term to reduce the risk of cardiovascular disease. reduced GIP.2_fm Page 370 Sunday.9 Jenkins studied purified fiber supplements such as pectin and guar gum. these slowed digestion and absorption. the (British National) Health Council recommended guidelines for nutritional changes in the whole population to decrease the incidence of modern metabolic diseases such as overweight and obesity. is not the same as a high carbohydrate high fiber diet because the latter might be a diet containing much white flour and sucrose supplemented by much wheat bran. containing much fat and sucrose. the British Diabetic Association made similar recommendations. High energy diets. which resembles the ancient tranditional diet of most peasants.13 . thereby increasing dietary fiber intake by 25%. are the main causative factors in the production of Type II diabetes mellitus. The original hypothesis carries a corollary that low fiber starchy carbohydrates. wherever possible. Mann and colleagues treated both Type I diabetes and Type II diabetes with a comparable diet. pernicious anemia and Hashimoto’s thyroiditis are extremely rare. 3RD EDITION flour.8 In Britain. they recommended that sugar intake be decreased to 50% and fat decreased by 25%. IMPROVED DIETS FOR THE WHOLE COMMUNITY In 1983. and gallstones. the American Diabetes Association recommended that carbohydrate intakes of Type I diabetic patients be increased to 50 to 60% of total calories. GENETIC AND AUTOIMMUNE FACTORS Genetic factors predispose strongly to Type II diabetes but are weak in Type I diabetes.12 These also recommended that simple sugar intakes be restricted and that carbohydrate. such as white wheat flour and white rice. Many autoimmune diseases are certainly very rare in African blacks: multiple sclerosis has not yet been definitely reported. Anderson and colleagues pioneered high fiber high starch carbohydrates in the treatment of diabetes. May 6.2387_ch5. both of which contain no fiber. Total energy intakes rarely decreased below 3000 kcal/day per head and were stationary during all these years. This variety is an autoimmune disease. 2001 7:33 PM 370 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.7 Unknown dietary factors or lifestyle protect sub-Saharan blacks from all or almost all autoimmune diseases. encourage overweight and obesity.6 This protective diet. J. Proc. Diabetologia. i. Lancet. 520. Hum. D. 11. The diabetic diet. R. chap 3. Bristol. 1982. 445. T.. 1983. 2001 7:33 PM DEVELOPMENT OF THE DIETARY FIBER HYPOTHESIS OF DIABETES MELLITUS 371 A consensus of opinion is emerging about desirable changes in modem Western diets in order to decrease the incidence of many diseases characteristic of modern affluent communities. P. J. H.. Uncertainty about the degree of change and the desirable rate of change will continue for a decade or more. 89. 782. Trowell.. Nutr. 1975. S. Edward Arnold.. 1974. D.. Diabetes. Non-Infective Disease in Africa. John Wright.. Dietary recommendations for diabetes for the 1980s. Western Diseases.. 42.. 762. 835. Am. ii. Campbell. Their Emergence and Prevention. Trowell.. and Burkitt. K. 9. C. H.. and Painter. L. Nutr. 2. G. Clin. Diabetes mellitus and dietary fiber of starchy foods. Nutr.2387_ch5. Diet in the aetiology of human diabetes. N. Lancet. C.. National Advisory Committee on Nutrition Education.. . 1.2_fm Page 371 Sunday.. H. Diabetes. Coronary Thrombosis. P. 323. 2. Trowell. Mann. Taylor. 217. Principles of nutrition and dietary recommendations for individuals with diabetes mellitus. 1981. and the Saccharine Disease. dietary carbohydrates and differences in digestibility. L. 1980. Do. Diabetes mellitus death rates in England and Wales 1920–70 and food supplies. 4. Trowell. 1969. T. Diet and diabetes.. 378. 8. 6. S. 10. London. Dietary-fiber hypothesis of the etiology of diabetes mellitus. 477. Himsworth H. 2. Long-term effects of high carbohydrate.. J. 1979. Jenkins. 36A. 24. 14. S53. Med. H. Cambridge. 719. H.. 23. 3. Anderson. 1978. 218. Diabetes Care. A. 12. Harvard University Press. 7. W. Lancet. D. M. 31. R. and Ward. 1949. 902.. J. 303. 13. and Wolever.14 REFERENCES 1. Trowell. 439. Diabetologia. Cleave. R. Eds. British Diabetic Association Medical Advisory Committee. H. Committee of the American Diabetes Association on Food and Nutrition Special Report. 1960. May 6. 998. 18. Diabetes Care. Appl. Soc. 77. 1982. 1983. MA.. Proposals for nutritional guidelines for health education in Britain. Prospects for prevention.. high fiber diets on glucose and lipid metabolism. 5. 1982. 2_fm Page 372 Sunday. 2001 7:33 PM . May 6.2387_ch5. High-fiber diets may decrease the emergence of obesity and resultant diabetes. drawn from the following sources: (1) computer (Medline) search on diabetes and fiber. blood pressure. (2) an excellent bibliographic survey of dietary fiber.63. and Index Medicus. Randles INTRODUCTION Diabetes mellitus is emerging as a major health problem throughout the world. The frequency of post-meal hypoglycemica is reduced in “chemical diabetes” on a high-fiber diet.226 6. High-fiber diets may specifically prevent the “carbohydrate-induced hypertriglyceridemia” associated with low-fiber. Akanji.10 and (3) review of references in cited articles and reviews.3 Treatment of Diabetes with High-Fiber Diets James W. 0-8493-2387-8/01/$0. Since 1976. Nutritional Abstracts.4 many basic and clinical investigators have documented the therapeutic benefits of fiber in diabetes and its complications.156.219 and obesity. and current evidence suggests that increased fat intake and decreased fiber intake may contribute. 2001 7:34 PM CHAPTER 5. high-carbohydrate intakes for diabetic subjects.50 © 2001 by CRC Press LLC 373 . and Kim M.121 3.115. While the clinical utility of dietary fiber in the treatment of diabetes is well established.3_fm Page 373 Sunday.144. up to and including the year 1986.1. May 6.143. including Scientific Citation Indexes.2387_ch5. report that 1.2 the role of fiber in the prevention of diabetes and reduction of risk for atherosclerotic complications is not established and data are still emerging.44.215.226 These and other observations led Trowell1 to postulate that diabetes was a fiber deficiency disorder.143. High-fiber diets may reduce risk factors for atherosclerotic cardiovascular disease — serum lipids (see tables).159. Anderson. High-fiber diets have favorable effects on blood rheology and hemostatic variables in diabetic subjects.25 2. however.144 5.3.00+$1. Abayomi O.229 4.125.134.12.134. The tables are an extension of our earlier report225 and include material in the medical literature between 1976 and 2000.229 The tables summarize the clinical studies of diabetic subjects using either fiber supplements or diets generous in high-fiber foods (herein termed “high-fiber diets”). High-fiber diets may ameliorate the frequency of discomfort associated with intermittent claudication in diabetic subjects. Some studies. 3). To improve palatability.70 Indian fenugreek seeds. and it is now generally accepted that diabetic subjects derive distinct advantages from both HCHF and HF diets. the high-carbohydrate. Corn bran hemicellulose198 and oat bran199 were also shown to lower the serum insulin response.79. Many of the studies78. random order.100 were adequately controlled and used random allocation.156 which investigated the effects of varying proportion of carbohydrate.2).76 Mexican nopal leaves. High-Fiber Diets Table 5. 2001 7:34 PM 374 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.89 in IDDM demonstrate the effects of guar supplementation on long-term (3-month) glycemic control and blood cholesterol levels. and psyllium extract clearly have greater effects on the glycemic response than do water-insoluble fibers such as cellulose and wheat bran.3_fm Page 374 Sunday. or barley — had greater effects than high-glycemic foods such as white bread or potatoes. 3RD EDITION CLINICAL STUDIES Fiber-Supplemented Meals Table 5.190 apple powder. or apple fiber.109. were incorporated into everyday foods such as bread.39 The glucose loads or meals supplemented with these soluble fibers are followed by lower glycemic responses than are control loads without fiber. these fiber supplements. The responses of normal subjects to similar test meals are described in a separate chapter. fiber. although some tested high-fiber (HF) diets that were similar in carbohydrate content to the control diets.57. moderately high-fiber diet.3. Our initial studies3.102 in NIDDM and Vaaler et al.76. The controlled studies of Uusitupa et al. Many of the more recent studies were well controlled using random allocation and crossover techniques.95.3.90 Most studies suggest that fiber supplements lower average blood glucose and cholesterol (especially LDL) levels and reduce requirements for insulin or oral hypoglycemic agents.88 Japanese glucomannan. the former probably being more beneficial. moderately high-fiber diet does not improve glycemic control in patients with mild NIDDM as compared to a low-carbohydrate. In fact.19. These findings could be interpreted to suggest that fiber supplements may improve long-term glycemic control. many investigators examined the response of diabetic subjects to fiber-supplemented diets (Table 5.1 summarizes the reported responses of diabetic subjects to glucose loads or meals with or without fiber supplements of high-fiber foods. and fat on metabolic control in NIDDM and concluded that high-carbohydrate diets for diabetic patients should select carbohydrates that are unrefined and high in fiber.110. Water-soluble fibers such as guar. Several studies suggest that wheat bran supplements11. A particularly useful study is that reported by O’Dea et al.105 with HCHF diets documented that these diets improved glycemic control.206 and xanthan gum. especially guar. legumes. Most investigators used guar or another fiber source rich in soluble fiber such as glucomannan.48. Fiber-Supplemented Diets To extend the meal studies. Other fiber supplements investigated in this context include cottonseed fiber. lowered insulin requirements.99 soy fiber. and hence acceptability. May 6. Another study216 indicated that a high-carbohydrate diet without a concomitant increase in fiber intake results in deterioration of glycemic .199 biscuits..2387_ch5.46.68. It should be noted that one study217 found that a high-carbohydrate. or crossover techniques.3 summarizes the responses of diabetic subjects to high-fiber diets developed from high-fiber foods rather than fiber supplements. pectin.40.3.89 also improve glycemic control and reduce the requirements for insulin and/or oral hypoglycemic agents.212 and they have been confirmed by many other groups (Table 5.15. highfiber (HCHF) diets. Most investigators used high-carbohydrate.3.92.83 High-fiber foods with a low glycemic index — oat bran.89. psyllium.27 and chocolate bars. moderately high-fiber diet actually increased plasma triglyceride and VLDL cholesterol concentrations. and reduced blood lipid levels. We have extended these studies106. 218 Use of Dietary Fiber in Distinct Diabetic Groups The various investigators (Tables 5. High-carbohydrate.30.80.123. 2001 7:34 PM TREATMENT OF DIABETES WITH HIGH-FIBER DIETS 375 control and increased plasma triglyceride and VLDL levels.3.168. high-carbohydrate diets reduce plasma cholesterol and triglyceride levels.1.94 and incorporation of natural high-fiber foods rather than fiber-enriched supplements. low-fiber diets.88 guar and glucomannan48. as was absorption of simultaneously administered drugs. High-Fiber Diets and Lipid Levels Many studies have shown a clear correlation between a high-fiber diet and reduced plasma lipid levels. various studies120.140.22.174.170 although the latter appears quite rare.175 High-fiber foods also are rich in phytochemicals and phytoestrogens that may favorably affect glucose metabolism and insulin sensitivity.130.and triglyceride-reducing properties.196 have chronic renal failure.126 report increased insulin binding to monocytes and adipocytes in subjects on HF diets.101. state of diabetic control.40. Special groups such as children. May 6.23. and trace element levels were generally unaffected.85 Mechanism of Effect of Dietary Fiber High-fiber diets.169 A potential problem is with acceptability and palatability of the diets.132 pregnant women. propionate.193 that fiber taken at a meal could improve glycemic response to subsequent meals.3_fm Page 375 Sunday.71. both high-fiber and high-fiber.101 fenugreek seeds.52 improving glycemic and lipidemic control.26.157 The consensus is that there are no significant long-term nutritional risks for patients on these diets. and 5.134.2.75. or degree of compliance to treatment. In the long term.139.3.94.164. especially in light of abdominal discomfort routinely experienced by many patients.152. Other possible mechanisms of action of fiber may be via modulation of the secretion of gut hormones13. Dietary fiber confers additional benefits in the management of diabetic patients who are hypertensive.196. Vitamin.222 Long-Term Safety of Fiber Preparations Many investigators have reported on the long-term efficacy and safety of high-fiber diets.3. especially those with a high carbohydrate content.91 oat bran.150 or have hepatic encephalopathy from liver cirrhosis.25. including patients on the insulin pump and artificial pancreas treatment. The major potential problems with prolonged fiber intake remain hypertriglyceridemia.194 or the intermediary metabolic effects of short-chain fatty acids — acetate.3) confirm that fiber confers distinct advantages to both IDDM and NIDDM subjects.89.96. reports on insulin clamp studies to assess insulin sensitivity in these subjects are not consistent.86.30.197 Wheat bran.155 and small bowel obstruction.95 also benefit from fiber supplementation.101. irrespective of body weight (lean or obese). However.28. 5.117. This is obvious from the reduced need for antidiabetic medication or insulin in subjects on these diets. probably exert their effect on glycemic control by improving insulin sensitivity.119 and geriatric subjects61.162 or improved.83 beet fiber.102. Plantago psyllium has been shown to reduce both total and LDL cholesterol levels while raising HDL cholesterol levels. high-fiber diets have also been shown to improve peripheral insulin sensitivity in healthy individuals.213 . This has partially been obviated by the use of low-dose guar preparations.206 also demonstrate cholesterol.63.68 xantham gum. mineral.2387_ch5. and butyrate — derived from the colonic fermentation of fiber in the diet.100.199 apple fiber.90. being variously reported as unchanged98.212 However. especially in high-carbohydrate. It has also been reported in both normal subjects and diabetics165–167.118.93 impregnation of guar with fructose for children.143. May 6. and pyruvate similar after control and test meals Fiber decreased reactive hypoglycemia Serum GIP lower. serum insulin. lactate.1 2387_ch5. GIP unchanged These fiber-supplemented meals were unpalatable Whole grain bread and apples compared to white bread and apple juice — Blood glucose and insulin response similar after control and test meals Exercise did not affect blood glucose Glucose tolerance normalized Gastric emptying time similar after control and test meals Variable changes in serum insulin Serum alanine. of Subjects Table 5. 376 6 IDDM “Chemical” diabetes — 6 IDDM 6 IGT — Special Group Effects of Dietary Fiber on Glycemic Responses to Single Meals for Diabetic Subjects 11 3 IDDM 8 NIDDM No. glucagon unchanged after test compared to control meal Metabolic clearance of radioactive glucose unaffected by bran High-fiber foods used Serum insulin lower after test than after control meal 25 22 23 24 20 21 19 17 16 15 14 13 12 11 4 Ref.— — 14 NIDDM 6 Autonomic neuropathy — — — — Children Gestational diabetes “Chemical” diabetes 12 NIDDM 13 NIDDM 8 NIDDM 12 NIDDM 12 NIDDM 21 IDDM 4 GDM 10 DM 8 IGT 1 IDDM 5 NIDDM — Nature of Meal Glucose Mixed Glucose solution Oatmeal Mixed meal Mixed Breakfast Breakfast Breakfast Breakfast Glucose solution Mixed Breakfast Breakfast Breakfast Type of Fiber Pectin or cellulose phosphate or cellulose High-fiber foods Guar Guar Psyllium High-fiber Guar or pectin or agar or locust bean gum High-fiber foods Guar and pectin Guar or lentils or soybeans Wheat bran Guar Guar and hemicellulose Guar Guar and pectin Glycemic Response Lower for pectin and cellulose phosphate.3.3_fm Page 376 Sunday. 3RD EDITION . 2001 7:34 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. glucagon. unchanged for cellulose Lower Lower Lower Lower Unchanged Lower All unchanged Lower or unchanged All lower Lower or unchanged Lower Lower Lower Lower Comments Fiber did not affect response of patients with autonomic neuropathy. somatostatin reduced Glycemic effect persisted through late postprandial period Insulin requirement unchanged with artificial pancreas Xylose absorption reduced. potato Mixed (and beans) Guar Wheat flour: extrusion vs. pancreatic polypeptide.3_fm Page 377 Sunday. 2001 7:34 PM TREATMENT OF DIABETES WITH HIGH-FIBER DIETS 377 . insulin. insulin levels unchanged Absorption of lactose from milkcontaining foods reduced Insulin response lowest with conventional whole grain bread — Postprandial C-peptide. May 6. GIP response reduced Insulin response unchanged HF breakfast had no effect on glucose tolerance to lunch (insulin pump treatment) — Insulin needs with artificial pancreas lower after test than control meals Guar biscuit used 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 2387_ch5. and GIP reponses reduced after beans despite similar fiber content to potato Insulin unchanged. baking Rye bread Guar Wheat bran Wheat bran Psyllium Mixed and rye Guar Guar Guar Pectin Lower Lower with beans Lower with beans Lower Lowest with whole grain bread Lower Lower Lower Unchanged Unchanged Lower Lower Lower Lower — Postprandial insulin reponse lowest with beans.Lean Pasta breakfast 5 — 7 IDDM — 19 8 IDDM 11 NIDDM Obese subjects — 22 20 NIDDM 2 IGT 7 NIDDM — 37 16 IDDM 21 NIDDM Obese subjects — 9 NIDDM 4 NIDDM — 10 NIDDM 5 NIDDM OGTT — 14 NIDDM Mixed breakfast Breakfast Breakfast Breakfast or lunch Breakfast Breakfast Breakfast Breakfast Lunch or supper Breakfast 8 NIDDM 2 IDDM 3 NIDDM Breakfast and lunch Glucose solution Mixed meal 6 IDDM — — 9 3 IDDM 6 NIDDM — 4 IDDM Soy polysaccharide Beans vs. nondiabetic subjects Glucose oxidation rates. serum triglycerides. glucagon. effect of processed food may be different in diabetic vs. 378 13 NIDDM 5 IGT No. fruit Guar Fiber + lactulose Plantago psyllium Bran Mixed high-fiber Mixed high-fiber Mixed high-fiber Type of Fiber No change No change Lower Lower Lower Lower Lower Lower Lower Lower Unchanged Lower Lower Glycemic Response Platelet adhesiveness reduced when fiber was given with glucose Lower AUC-glucose. May 6. no reduction in postprandial plasma C-peptide levels Lower area under curve for glucose and insulin.3. and insulinic index Lower glucose and insulin responses to meals Lower postprandial rise in blood glucose. 3RD EDITION .3_fm Page 378 Sunday. AUC-insulin. wheat bran. of Subjects Table 5.— Mildly hypertriglyceridemic — — — — — 8 NIDDM 8 NIDDM 14 NIDDM 14 NIDDM 14 NIDDM 10 NIDDM Obese — 14 NIDDM 4 NIDDM 12 NIDDM Indian population — 55 DM — 22 10 IDDM 12 NIDDM North Indians Special Group Breakfast Breakfast Breakfast Breakfast Breakfast Breakfast Breakfast Breakfast and lunch Breakfast OGTT Breakfast Breakfast OGTT Nature of Meal Lupin Insoluble maize-cob Oat bran Oat bran enriched with β-glucans Locust bean Oat bran. soluble:insoluble ratio did not affect glycemic response Lower AUC-glucose and postprandial insulin level Decreased glucose response.1 (Continued) Effects of Dietary Fiber on Glycemic Responses to Single Meals for Diabetic Subjects 2387_ch5. lower plasma insulin and plasma GIP. lower insulin response in patients with BMI > 30 Glucose and insulin levels unchanged Glucose and insulin levels unchanged Reduced postprandial plasma glucose and insulin Insulin requirement with artificial pancreas reduced in IDDM but not in NIDDM Typical North Indian diet improves glucose tolerance Postprandial insulin responses lower Comments 185 185 186 185 184 183 182 181 180 44 43 42 41 Ref. 2001 7:34 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. high-carbohydrate. 10 NIDDM Youths Excellent metabolic control Excellent metabolic control — 6 NIDDM 30 IDDM 10 NIDDM — 6 NIDDM Well-controlled — 8 NIDDM 7 NIDDM — 10 NIDDM 2387_ch5. HF. noninsulin-dependent (type II) diabetes mellitus. no effect on blood glucose when psyllium taken in water before the cereal Reduced postprandial glucose elevation and serum insulin concentration. IDDM. rice. May 6. categorized as second-meal effect Lower postprandial rises in blood glucose. oral glucose tolerance test. low-carbohydrate. 2001 7:34 PM TREATMENT OF DIABETES WITH HIGH-FIBER DIETS 379 . moderatefiber. GDM. slower gastric emptying No change in blood sugar levels Lower AUC-glucose vs. low-fiber. and plasma C-peptide. MFLC. high-carbohydrate. insulin-dependent (type I) diabetes mellitus. pasta. NIDDM. high-fiber. LFHC. pasta or rice 214 195 214 194 193 192 190 189 188 187 186 Note: DM. impaired glucose tolerance (formerly called chemical or latent diabetes). reduced postprandial glucose elevation after lunch. high-fiber. IGT. and barley Lower AUC-glucose vs. gestational diabetes mellitus. diabetes mellitus (type not indicated). serum insulin.— — — 8 NIDDM 6 NIDDM 18 NIDDM Pasta Pasta Pasta Breakfast Breakfast and dinner Breakfast Breakfast Spaghetti Breakfast Dinner Breakfast Barley Guar Beet fiber Algae-isolate Psyllium Psyllium flake cereal Soya Guar Vegetable Food fiber Wheat farina + oat gum Lower No change Lower Lower Lower Lower Lower No change Lower Higher Lower Reduced postprandial plasma glucose and insulin Greater postprandial glycemic response compared to equicaloric higher-fat and lower-fiber meal Blood glucose levels inversely related to quantity of vegetable fiber in test diet No change in integrated postprandial glucose or C-peptide response Lower glycemic profile as compared to that produced by equivalent weight of cellulose Lower blood glucose. OGTT. HCHF.3_fm Page 379 Sunday. cholesterol lower Insulin requirements lower Serum cholesterol. serum cholesterol lower Glycosuria reduced more with higher than lower carbohydrate intake Glucose and insulin responses lower — Insulin requirements. 49 45 Ref. of Subjects Table 5. May 6.— 9 — 7 IDDM — — — — — — 19 IDDM 11 5 IGT 6 NIDDM 11 8 IDDM 3 NIDDM 14 IDDM 10 IDDM 6 IDDM Poorly controlled. 2001 7:34 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 380 8 IDDM 13 NIDDM 7 IDDM 2 NIDDM — Special Group Response of Diabetic Subjects to Fiber-Supplemented Diets 9 DM No. poorly compliant — — 6 DM 38 IGT 22 NIDDM — — Type of Fiber Wheat bran Guar Guar Guar Guar + pectin Guar Guar or wheat bran Guar Guar Wheat bran Cellulose in bread Glucomannan Guar Guar 14–28 91 28 183–365 3 5 91 1 56 30 10 90 5 7 Duration (days) Lower Unchanged Lower Unchanged Lower — Unchanged Unchanged Lower Lower Lower — — Glycemic Response Comments Insulin requirements lower Serum cholesterol lower Serum cholesterol 9% lower Insulin requirements.3. triglycerides lower Insulin needs with artificial pancreas lower in 5 out of 7 patients No beneficial effects Glycosuria with guar supplements less than half of control values Glycosuria 38% lower with guar crispbread than with control diet 57 59 60 56 55 54 53 52 50 51 47 48 46. 3RD EDITION .3_fm Page 380 Sunday.2 2387_ch5. LDL cholesterol. drug requirements reduced Improvements with guar persisted after guar was stopped Plasma cholesterol lower on guar Body weight. triglycerides reduced Urine glucose excretion 31% lower Glycohemoglobin and serum cholesterol lower Serum cholesterol lower Serum cholesterol lower Body weight. bran increased cholesterol Glycohemoglobin. poorly controlled subjects Children Geriatric subjects Stable and labile diabetic subjects 10 NIDDM 14 NIDDM 17 IDDM 12 NIDDM — 10 NIDDM Pregnant — 8 NIDDM 12 IDDM — 8 IDDM Guar Pectin Guar or wheat bran Guar + wheat bran Guar Boiled nopal leaves (Mex. cholesterol. May 6. triglyceride.Subjects taken off all drugs Constipated patients — — — 10 NIDDM 40 NIDDM 12 NIDDM 17 6 IDDM 11 NIDDM 7 DM — — 17 IDDM 5 2 IDDM 3 NIDDM — 28 IDDM Obese. total. glycated albumin unchanged Total cholesterol. 2001 7:34 PM TREATMENT OF DIABETES WITH HIGH-FIBER DIETS 381 .) Apple fiber Guar Guar Psyllium Guar Guar Wheat bran Guar Wheat bran Guar 14 91 91 61 7 10 49 7–21 122 28 61 10–15 28 91 91 Lower Unchanged Lower Lower Unchanged Lower Lower Lower Lower — Lower Lower — Unchanged Unchanged Guar lowered glycohemoglobin.3_fm Page 381 Sunday. cholesterol lower Oral hypoglycemic agent doses and cholesterol decreased Glycosuria and cholesterol decreased Insulin requirements decreased Serum cholesterol lower Improved glycemic control 29 74 73 72 71 70 68 69 66 67 63 64 65 62 61 61 2387_ch5. insulin requirements. HDL cholesterol lower. and GIP responses lower Insulin response. May 6.— — — — — 20 8 IDDM 12 NIDDM 8 NIDDM 13 8 IDDM 5 NIDDM 9 NIDDM 10 NIDDM Guar Xanthan gum Guar Guar Guar (low dose) Wheat bran — 42 42 98 112 61 42 28 42 28 14 Apple powder Guar Rye bran Cellulose — 79 38 IDDM 41 NIDDM 28 Corn bran Duration (days) 28 Type of Fiber Soy hull — — 8 NIDDM Lower Lower — Unchanged Unchanged Lower Lower Lower Unchanged Lower Unchanged Lower Glycemic Response Comments Total. triglyceride.3_fm Page 382 Sunday. VLDL.3. antidiabetic medication reduced Glycosuria. HDL cholesterol lower. HDL cholesterol increased VLDL cholesterol. blood chemistry. blood/urine chemistry unchanged 84 83 82 81 80 78 79 77 76 Ref. triglyceride unchanged Insulin requirements lower Total. total. 3RD EDITION . LDL cholesterol. of Subjects Table 5. glycohemoglobin. glycohemoglobin lower Total. insulin sensitivity and receptor binding greater. phospholipids. total. gastrin. HDL. trace elements unaffected Serum cholesterol lower. 382 12 IDDM 4 IDDM Special Group No. insulin need greater with cellulose Glycohemoglobin. insulin need lower with wheat bran Glycohemoglobin. 2001 7:34 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. glycosuria reduced. LDL cholesterol higher Serum cholesterol. glycohemoglobin unchanged. VLDL triglyceride. hepatic glucose production unchanged Hematologic parameters. triglycerides.2 (Continued) Response of Diabetic Subjects to Fiber-Supplemented Diets 2387_ch5. insulin sensitivity unchanged. LDL lower in men. cholesterol triglyceride. LDL. PSE parameters unchanged 97 98 95 94 93 92 90 91 89 88 87 86 85 2387_ch5. triglyceride unchanged. noncompliant subjects 16 NIDDM 7 NIDDM — Asian Indians 5 NIDDM 28 IDDM Obese — 11 NIDDM 8 NIDDM Subjects with portal systemic encephalopathy (PSE) from liver cirrhosis 8 NIDDM Guar gum Guar Guar Guar Guar Guar 112 28 183 42 42 56 183 30 91 Wheat bran Guar Guar 91 21 183 14 15 Guar Fenugreek seeds Guar Guar Vegetable protein and psyllium Unchanged Lower Lower — Unchanged Unchanged — Lower Lower Lower Lower — Unchanged Lower Glycohemoglobin. total cholesterol. insulin response lower Glycohemoglobin.Geriatric Obese Lean patients on insulin pump treatment 14 NIDDM 17 NIDDM 9 IDDM — 16 NIDDM Children Subjects with near normal fasting plasma glucose levels 29 NIDDM 22 IDDM — Poorly controlled. total. erythrocyte insulin binding. LDL cholesterol lower Tolbutamide dose reduced. LDL cholesterol lower HDL cholesterol total cholesterol ratio higher Nutritional parameters unchanged Glycosuria. fasting insulin lower. May 6. triglyceride reduced Glycosuria. insulin. HDL cholesterol. triglycerides unchanged No additional benefit Glycohemoglobin.3_fm Page 383 Sunday. C-peptide responses unchanged Serum triglyceride higher. effect not maintained over 6 months Glycosuria. serum cholesterol. serum calcium. LDL cholesterol lower Serum cholesterol. C-peptide. 2001 7:34 PM TREATMENT OF DIABETES WITH HIGH-FIBER DIETS 383 . total cholesterol reduced C-peptide lower. glycohemoglobin increased. GIP response lower. OGTT and glycohemoglobin unchanged Glycohemoglobin. bowel movements greater. insulin needs. HDL cholesterol. glycohemoglobin. neurotensin higher Lower fructosamine. pancreatic polypeptide and motilin lower in obese. and LDL:HDL Hypoglycemic effect No change in basal or aftermeal glycemia No change in basal or aftermeal glycemia 200 200 200 199 198 197 196 104 103 102 101 100 99 Ref. insulin enteroglucagon. Zn. Ca. vitamins A and E lower. Mn. LDL. GIP. LDL.2 (Continued) Response of Diabetic Subjects to Fiber-Supplemented Diets 2387_ch5. glucose and insulin unchanged Lower fasting plasma glucose. 2001 7:34 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. triglycerides lower. 3RD EDITION . 20 obese.3_fm Page 384 Sunday. glucagon. Cu. May 6. hypertensive 11 NIDDM 28 NIDDM — 12 NIDDM Obese — 12 NIDDM 19 NIDDM — 8 NIDDM Poorly controlled obese — 12 NIDDM Type of Fiber Cellulose pulp Wheat bran Lignin-enriched bran Oat bran Soluble corn bran hemicellulose Plantago psyllium Glucomannan Guar Guar Guar Beet fiber Guar Cotton seed fiber 28 28 28 84 180 42 21 14 91 396 56 183 30 Duration (days) No change Lower No change Lower Lower Lower Lower Lower Lower Unchanged Unchanged — Lower Glycemic Response Comments Insulin response to test meal. 384 33 NIDDM Special Group No. Mg) balance unchanged Systolic BP.Free-living men Hyperlipidemic Hyperlipidemic Hyperlipidemic 8 NIDDM 100 NIDDM 100 NIDDM 100 NIDDM — 125 IDDM Mild IGT. PYY unchanged. cholesterol. and fasting glucose Lower mean glycemic and insulin responses. higher HDL Lower serum insulin response. HDL increased.5 mmol/l) Serum cholesterol. triglyceride lower. LDL cholesterol lower (if serum cholesterol > 6. postprandial insulin. of Subjects Table 5. glycohemoglobin unchanged Serum total cholesterol.3. cholesterol. and blood pressure. serum lipids unchanged Mineral (Fe. 8 non-obese Hyperlipidemic. plasma Zn higher Total. total cholesterol. and triglycerides. decreased HbA1c . lower total cholesterol. no change in blood glucose or HbA1c . concentrations of all other substrates unchanged 191 208 207 207 206 205 204 203 202 201 200 TREATMENT OF DIABETES WITH HIGH-FIBER DIETS Note: See Table 5. no change in blood glucose or HbA1c .— — — Mildly hypercholesterolemic 72 NIDDM 10 NIDDM 10 NIDDM 17 IDDM Guar Guar Guar Wheat bran Konjac food Sweet lupine hull Beet fiber Guar Guar Guar Citrus pectin 90 42 90 90 65 60 42 90 56 336 28 Lower Lower Lower Lower Lower No change No change No change Lower Lower No change No change in fasting blood glucose. Lower postprandial plasma insulin.1 for definition of abbreviations. serum triglycerides. or body weight Blood glucose increased and cholesterol decreased during first month of supplementation. decreased triglyceride levels in hypertriglyceridemic subjects Lower postprandial plasma insulin. 2001 7:34 PM 385 . — — 16 NIDDM 40 33 NIDDM 7 IDDM — 13 NIDDM Moderately poor metabolic control — 16 NIDDM 7 IDDM — Hyperlipidemic 15 NIDDM 100 NIDDM 2387_ch5. insulin response unchanged Lower fasting insulin and HbA1c concentrations Lower cholesterol. Decreased fasting blood glucose and HbA1c Lower HbA1c and cholesterol No change in basal or aftermeal glycemia Improved long-term glycemic control. postprandial blood glucose. May 6. blood glucose decreased to control levels during second month Reduced fasting blood glucose. C-peptide response increased.3_fm Page 385 Sunday. and lipid concentrations.3. and HbA1c . HbA1c . postprandial glucose tolerance. 3 2387_ch5. blood lipids lower Insulin requirements.27 9 IDDM 18 NIDDM 16 10 IDDM 6 NIDDM 5 NIDDM — Geriatric nursing home residents — — 8 4 IDDM 4 NIDDM — Obese 11 4 IDDM 7 NIDDM 21 12 IDDM 9 NIDDM 14 NIDDM 11 IDDM — — — 10 7 IDDM 3 NIDDM Lean — 14 3 IDDM 11 NIDDM HCHF MFLC HF HCHF HCHF HCHF HCHF HCHF HCHF HCHF HCHF HCHF Nature of Diet 42 42 14 10 18–26 12–42 42 42 16 182–1095 1478 12–28 Duration (Days) Lower Slightly lower Lower Lower Lower Lower Lower Unchanged Lower Lower Lower Lower Glycemic Response Comments Serum cholesterol lower. blood lipids reduced Glycohemoglobin. blood lipids reduced 114 113 112 111 110 109 107 108 106 105 96 3 Ref. 2001 7:34 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. serum lipids lower Insulin dose. 386 20 IDDM — Special Group Response of Diabetic Subjects to High-Fiber Diets Developed from High-Fiber Foods 13 8 IDDM 5 NIDDM No.3. 3RD EDITION . increasing fiber from 18 to 33 g/day did not change glycemic control Diabetic control improved Serum cholesterol lower HCHF and LFHC diets had equivalent insulin dose-reducing effects Insulin requirements. good long-term outpatient compliance Insulin or oral hypoglycemic agent needs. triglycerides lower. serum cholesterol.3_fm Page 386 Sunday. cholesterol lower Insulin requirements. May 6. of Subjects Table 5. serum cholesterol lower Insulin requirements. HDL:LDL ratio higher With low carbohydrate. LDL. body weight lower Urinary C-peptide excretion lower on high-bean diet than on high-bran diet Insulin sensitivity unchanged Glycosuria decreased Insulin requirements decreased 130 129 128 127 126 125 124 123 121 120 117 118 119 116 115 2387_ch5. 2001 7:34 PM TREATMENT OF DIABETES WITH HIGH-FIBER DIETS 387 . May 6. fasting insulin unchanged. but not in Asians Insulin requirements lower.3_fm Page 387 Sunday. mean blood pressure lower Serum cholesterol lower by 15%. postheparin plasma lipoprotein lipase activity unchanged Total. triglyceride. blood lipids. improved insulin receptor binding and sensitivity Serum triglycerides lower Insulin binding to circulating monocytes increased Diet and exercise sustained lower serum lipids Blood pressure lower in whites and blacks. VLDL cholesterol lower Glycosuria reduced. hypertensives 35 1 IDDM 34 NIDDM Pritikin program and exercise — 7 NIDDM 69 NIDDM — 21 10 IDDM 11 NIDDM Very obese Children Pregnant 7 NIDDM 10 IDDM 20 10 IDDM 10 NIDDM — Obese 8 NIDDM 20 IDDM Pritikin program with exercise 60 NIDDM HF HCHF HCHF HF HCHF HCHF (low sodium) HCHF HCHF HCHF HCHF HCHF HF HCHF HF HCHF 91 10 42 91 21 30 730–1095 42 42 28 7 42 28–210 2 26 — Lower — Lower Lower — Lower — Slightly lower Unchanged Unchanged Lower Lower Lower Lower Glycohemoglobin. 9 Asian.— — Mild hypertension 14 6 IDDM 8 NIDDM 25 NIDDM Long-term outpatient study 10 IDDM 10 2 IDDM 8 NIDDM 9 NIDDM — 16 white. 10 black. insulin binding to circulating monocytes increased Plasma-clotting factors reduced Insulin or oral hypoglycemic agent needs. hematologic variables unchanged Frequency of hypoglycemic reactions and insulin dose unchanged Triglyceride. total cholesterol unchanged Glycohemoglobin. fecal fat. HDL. LDL. LDL cholesterol. no drug treatment 6 NIDDM 7 NIDDM — Poorly controlled 13 IDDM 22 NIDDM Self-selected foods used — 4 IDDM 6 IDDM — Mild hypertension 25 5 IDDM 20 NIDDM Nature of Diet HCHF HCHF HCHF HF HCHF HCHF HF HCHF HCHF (low-fat. lipase activity in muscle or adipose tissue. 3RD EDITION . same blood pressure reduction as bendrofluazide. insulin requirement. HDL higher 143 142 141 139 138 137 136 135 134 132 131 Ref. body weight higher Insulin sensitivity improved. HDL cholesterol lower Glycosuria. glycohemoglobin unchanged compared to MCLF (high-fat) diet Health education is probably the best method for achieving optimal control in IDDM subjects No deleterious effect on blood urea and nutritional status. May 6.Chronic renal failure (hospitalized) 5 IDDM Lean Moderate hypertension. glycohemoglobin. body weight lower LDL. plasma creatinine reduced Total.3 (Continued) Response of Diabetic Subjects to High-Fiber Diets Developed from High-Fiber Foods 2387_ch5. total. 2001 7:34 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. HDL. glycosuria. 388 14 6 IDDM 8 NIDDM Children — 14 NIDDM 12 IDDM Special Group No.v. lowsodium) HCHF HF 91 28 42 10 580 42 42 10 91 14 21 Duration (Days) — Unchanged Lower Lower Worsened Unchanged Lower Lower — Unchanged Lower Glycemic Response Comments Glycohemoglobin.3. Zn). total cholesterol reduced. blood pressure.3_fm Page 388 Sunday. total cholesterol lower. HDL cholesterol lower when diet complements intensive patient education Insulin. insulin need. triglycerides. HDL. body weight lower. of Subjects Table 5. i. fat tolerance. trace elements (K. fatty acids. symptom frequency reduced. geriatric Chronic renal failure. LDL cholesterol. and -fiber diet without need for insulin Total. C-peptide. insulin. cholesterol. glycosuria reduced on low-GI diet. glycohemoglobin. least compliance was to carbohydrate recommendation Blood pressure. triglycerides.Poorly controlled in metabolic ward study — Obese. nitrogen balance. blood lipids unchanged Subjects complied best to fiber recommendation. body weight unchanged 158 156 154 153 152 151 150 149 147 146 145 144 2387_ch5. glucagon reduced. insulin. Doppler ankle/arm ratios unchanged Glycohemoglobin. total. serum phosphate increased Glycohemoglobin. insulin needs. OGTT improved Glycohemoglobin. body weight lower. low-sodium) 84 14 700 42 21 183 10 56 30 21 365 91 Lower on low-GI diet Lower — Unchanged Lower Elevated Lower Lower Lower Lower Unchanged — Body weight. body mass index. somatostatin unchanged Total cholesterol lower. -legumes. insulin sensitivity unchanged Control throughout pregnancy with highcereal. moderate protein restriction Poorly controlled Poorly controlled Patients on insulin pump treatment Retrospective evaluation of three studies 15 NIDDM 14 NIDDM 6 IDDM 28 NIDDM 13 NIDDM 10 IDDM Outpatient study To compare low glycemic index (GI) with high-GI foods 10 NIDDM 16 NIDDM 40 19 IDDM 21 NIDDM 1 GDM — Intermittent claudication 35 15 IDDM 20 NIDDM 17 6 IDDM 11 NIDDM About 45% carbohydrate HCHF HCHF HF HCHF HF HCHF HF Mixed HF HF (legumes) HCHF HF (low-fat. glycohemoglobin. lipoproteins unchanged Glycosuria lower.3_fm Page 389 Sunday. triglyceride. insulin. 2001 7:34 PM TREATMENT OF DIABETES WITH HIGH-FIBER DIETS 389 . insulin receptor binding unchanged Body weight. glucagon. LDL cholesterol. HDL cholesterol. nutritional status unchanged. May 6. LDL cholesterol. total. total. triglyceride. and HDL unchanged Glycosuria. renal function. insulin responses reduced. LDL:HDL ratio lower. glycemic control and other lipid fractions unchanged Better glycemic control.1 for definition of abbreviations. triglyceride. highly motivated Early NIDDM HF HCHF HCHF HCHF HF HF Nature of Diet 42 30 28 540 — 12 Duration (Days) Lower Lower Lower No change Lower Lower Glycemic Response Comments Lower postprandial capillary blood glucose Lower basal insulin and body weight. lower blood glucose.3. May 6.3 (Continued) Response of Diabetic Subjects to High-Fiber Diets Developed from High-Fiber Foods 2387_ch5. of Subjects Table 5. reduction of insulinemia and hyperglycemia in glucose tolerance test Improvement in glycemic control during recruitment phase Reduced basal insulin requirements. HbA1c . and fructosamine.3. 2001 7:34 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and AUC-insulin. increase in insulin immediate pool. glucose excretion. AUC-glucose. and VLDL 215 213 212 211 210 209 Ref. lower total cholesterol and HDL. 390 NIDDM 9 IDDM No. 13 NIDDM Obese — 10 IDDM 84 NIDDM — 70 NIDDM Well controlled. decreased total cholesterol and triglycerides with no effect on HDL Lower preprandial plasma glucose. lower total cholesterol. amelioration of clinical symptoms. 3RD EDITION .Special Group — Note: See Table 5.3_fm Page 390 Sunday. 7. These diets also lower blood glycohemoglobin levels.223. The bran layer of cereal fibers is rich in fiber and phytochemicals that may have specific antioxidant.220 A high fasting insulin concentration has been linked to an increased incidence of heart disease.227 We recommend three servings daily. Diets high in carbohydrate but not fiber content. we recommend an intake of 800 IU daily for adults with diabetes. This amount should be reduced in individuals with nephropathy. plasma cholesterol levels (20 to 30%).222.226 1. these diets can reduce insulin requirements by 25 to 50% and improve glycemic control.171 For obese persons with diabetes. intake of salt and vitamin and mineral supplements. For lean individuals with type 1 diabetes. and other effects that provide these benefits.177 with less than 10% saturated fat and daily cholesterol intake of less than 200 mg.148. We routinely recommend a prudent diabetic diet containing the following:7.176.179 a high–monounsaturated fat diet has been shown to increase HDL cholesterol levels and reduce the risk of hypertriglyceridemia associated with high-carbohydrate. 2001 7:34 PM TREATMENT OF DIABETES WITH HIGH-FIBER DIETS 391 High-Fiber Diets and Epidemiological Studies High fiber intakes appear to reduce risk for developing diabetes.161.226 Because of the cardioprotective and anti-inflammatory effects of vitamin E.178 4.8–1. Two independent studies have confirmed an inverse correlation between a high-fiber diet and the risk for NIDDM in both men and women. with the increase mainly from monounsaturated fats.222. phytoestrogen. may adversely affect blood lipid levels. especially in relation to total caloric intake.226 3.2 g/kg desirable body weight to a maximum of 90 g/day). 55 to 60% of energy from carbohydrate (two-thirds derived from polysaccharides). 5. use of alternative sweeteners. The specific benefits of whole grain intake deserve special mention. these diets can lower antidiabetic drug needs by 50 to 100% and often eliminate the need for insulin.219 Further observational studies suggest that high fiber intakes may protect from development of obesity. fiber-deficient diets. as well as alcohol ingestion. have been reviewed elsewhere. as indicated above. and decreasing blood lipid concentrations.227.221 Several epidemiological studies have indicated a strong link between a high-fiber diet.229 CONCLUSIONS High-fiber diets provide many benefits for diabetic patients.224 Insulin resistance appears to be a forerunner or risk factor for development of diabetes. consistent with the American Heart Association recommendations. a high-fiber diet is associated with a lower fasting insulin concentration. and the prevention of coronary heart disease. reducing postprandial insulin levels and antidiabetic drug requirements. Other features of the diabetes diet. anti-inflammatory. especially a generous intake of whole grains. these diets associated with high satiety promote weight loss and usually provide reasonable glycemic control without specific antidiabetic medication.228 High fiber intakes also appear to protect from development of hypertension. to include soluble and insoluble fibers from commonly available foods. by lowering blood glucose concentration. total fat intake could be increased to 30 to 40% energy. While still controversial with yet unclear long-term effects.222 In non-diabetic subjects. Dietary fiber of about 35 g/d (or 15 to 25 g/1000 kcal).230 . and blood pressure (average of 10%).3_fm Page 391 Sunday.155. May 6. triglycerides (slightly to moderately).2387_ch5.171 For lean individuals with type 2 diabetes. 2. Whole grains appear to increase insulin sensitivity and reduce risk for developing diabetes and coronary heart disease. 12 to 16% of total calories from protein (or daily intake of 0. When increased carbohydrate intake cannot be tolerated or is impractical. Less than 30% of total calories from fat.135. Lancet. 1984. 1981. G. Nutr. Gabbe. G. 1982. Reviews and Bibliography. L. Simpson.. S. J. Levitt. Arch Dis. 40. Carlstrom. M. et al... 9. Jenkins.. 1. Gatti. and Tchobroutsky.. E. 1982. 656 (suppl... 8. 4. W. Med. Effect of different kinds of fiber on postprandial blood glucose in insulin-dependent diabetics. New perspectives in nutrition management of diabetes mellitus. Guar biscuits in the diabetic diet. 2. 18. and Schersten. S. Child. J.. Leeds. Monnier. Intern. The effect of bran on glucose kinetics and plasma insulin in noninsulin-dependent diabetes mellitus. 28. W. 29. and Hetenyi. 17. Clin. Scand. J.. 45. high fiber diet on hyperglycemic diabetic men. 143. . Anderson. 1. and Geil. Diabetes. The role of dietary carbohydrate and fiber in the control of diabetes. 20. 23. Diabetes Metab. 1980.. P. 8. 28. Unabsorbable carbohydrates and diabetes: decreased postprandial hyperglycaemia. Smith.). vol. and Jenkins. Anderson. Boca Raton.. Acta Med. 25. 389. Diabetologia.. T. J. Effect of dietary fiber on the oral glucose tolerance test in pregnancy. 27. 1988. A. J. J. A.. D. J. 14. Diabetes Metab. 15. Dietary fiber in management of diabetes. 281. 1978. H. 1981. 159. J. Nutr.. 83. K.. J. 58.. Asp. Restored synergistic enterohormonal response after addition of dietary fiber to patients with impaired glucose tolerance and reactive hypoglycaemia. 19. Aust. J. Diabetes Care. 7.. M. L. 85.. J. T. 542. E. 1978. 3RD EDITION REFERENCES 1. O. H. 1248. N. 16. E. M.. L. A. 172. 762. J. London.. Council on Scientific Affairs.. M. et al. 656 (suppl... S. et al.. Acta Med. G. Dietary Fibre Perspectives. American Medical Association. 1979. 11. 2. Beneficial effects of a high carbohydrate. Br. Wahlquist. et al. Anderson. 1975. G. Kiehm. L.. 47.. D. J. R. 187. B. in CRC Handbook of Dietary Fiber in Human Nutrition. et al.. Vaaler. 1980. G. 2.. 67. FL. 1980. J.. A. 1982. 1982. et al. 1990. Med. Ed. M. Med.). C. Dietary fiber hypothesis of the etiology of diabetes mellitus. K. Effect of naturally occurring dietary fiber in Western foods on blood glucose. Nutr. Afr. JAMA. 208. Diabetes Care. S. W.. et al.. and Jenkins. 351. Effect of guar crispbread with cereal products and leguminous seeds on blood glucose concentrations of diabetics. 3. 85... B. F. Nutr.. 339. 29. 2001 7:34 PM 392 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 1980.. Alberti. 1981.. Dietary fiber and health. and Evans. Clin. Gynecol. 484... 1989. 196. T. W.. et al. Dietary fiber supplementation of a “normal” breakfast administered to diabetics. Med. Slama. Leatherdale. P. Sartos. 3. K. D. D. Acta Med... T. 1984. Eds.. et al. W. and Ward. Reduction of postprandial insulin needs by pectin as assessed by the artificial pancreas in insulin-dependent diabetics. Diabetes Care. Anderson. W.. 1987. Dietary supplementation of fibre (Lunelax) as a means to reduce postprandial glucose in diabetics. 1232. The effect of unabsorbable carbohydrate on gut hormones. et al. 17. 11. J. 6. 520. 51. Med. 262. Am. et al. J. 26. Spiller.. 515. J. 5. Chenon. 1982. Am. May 6. Scand. Effect of added fiber on the glucose and metabolic response to a mixed meal in normal and diabetic subjects. 1976. 1982. Bolton. G. 217. Adv. Diabetes Care. 514. 11. A. A. Diabetologia.. G. Vinik. Dietary fiber in type II diabetes. 160.. T. D.. H. Lat. I. 24.Z. 1980. S. and Aagenase.. Jenkins. 13. R. Trowell. James. I. 18. Obstet. et al. Morgan. Clin.. G. S.. Acta Diabetol.. Scand... 1988. J. 3... 8. 21. and Burley. G. The effect of dietary fiber on glucose and hormone responses to a mixed meal in normal subjects and in diabetic subjects with and without autonomic neuropathy...3_fm Page 392 Sunday.. 26. N. Effects of dietary fiber on postprandial glycemic profiles in diabetic patients subjected to continuous programmed insulin infusion. 379. Diabetes Care. Metab.. B.2387_ch5. Effects of guar-enriched pasta in the treatment of diabetes and hyperlipidemia. Am. et al. Baumer. 22. 1984. et al. Goulder. J. 24.. V. 905. N. Am. 61. H. Poynard. 1. D. R. Am. Hanssen. A. Ann. Hall. Williams.. 10. Influence of indigestible fibers on glucose tolerance. 1976. 895. Guar and gastric emptying in noninsulin dependent diabetes. Effects of dietary fiber and exercise in mid-morning diabetic control — a controlled study. 349. Dietary fiber and carbohydrate metabolism. 19. John Libbey. 12.. 1980. Monnier. Treatment of diabetes with high fiber diets. 57. CRC Press. A. Adv. Kanter. J. Effects of four meals with different kinds of dietary fiber on glucose metabolism in healthy subjects and non-insulin-dependent diabetic subjects. O. A. 119. J. Diabetologia. 26. 3.. India. and Mantere... et al. et al. Cohen.. et al. pancreatic polypeptide. 1984. 32. 1980. Uusitupa. Aust. J. 31. et al. S. 1980. ii. 1985... et al. Phys. 779. M. McMurry. A high wheat bran diet in insulin treated diabetes mellitus: assessment with the artificial pancreas. C. Sci. Nutr. 1729. S. C-peptide and GIP responses in non-insulin-dependent (type II) diabetics and controls. Diabetes Care. M. 2. 30. Dietary fiber and ketone bodies: reduced urinary 3-hydroxybutyrate excretion in diabetics on guar. B. Am. 1977. et al. 48. A.. 66. 462. 34. Br.. Glucose tolerance in North Indians taking a high fibre diet. Y. Effects of high fiber breakfasts on glucose metabolism in non-insulin-dependent diabetics.. 1978. Lancet. J. et al. 88. J.. Nutr.. McIvor.2387_ch5. 41. D. Schrezenmeir. Improved glucose tolerance and insulin response in obese and diabetic patients on a fiber-enriched diet. 39A.. R. 37. Appl. 1980. Diabetes Care. Jenkins. J. et al. 1989.. Nutr. 659. 42.. 47. Simpson. The effect of different types of Finnish bread on postprandial glucose response in diabetic patients. A. 211... 45. 46. and Horwitz. L.. J. Diabetic diets: high carbohydrate combined with high fiber. et al. Assoc. 46. Nutr. Jenkins. type 1 and type 2 diabetic man.. H. R. 33. et al. 1980. 1987.. Heinonen. 42. D. 1984. Scand. 1985. Biol. J. Christiansen. Am. Food physical factors have different metabolic effects in nondiabetics and diabetics. Jenkins. plasma immunoreactive insulin. et al. 1984. Med. J. R. 1985. Glucose tolerance and blood lipids in bran-fed patients with impaired glucose tolerance. 1978. 275. Clin. insulin. Khan. Med. et al. Bosello. Wochenschr. W... Miranda. 59. Clin.. Med. Flattening postprandial blood glucose responses with guar gum: acute effects. Comparison of glycaemic response and insulin requirements after mixed meals of equal carbohydrate content in healthy.. B.. glucagon. somatostatin. and Baumgardner. 1988. Eur. 45. Isr. Jarjis. Acta Med. et al. Ann. Hormonal and metabolic responses to breakfast meals in NIDDM: comparison of white and whole-grain wheat bread and corresponding extruded products. Nutr. Jenkins. Role of guar and dietary fibre in the management of diabetes mellitus. 205. May 6. et al. 44. Am. et al. 8. 274. J. 987. Hum. 1988. A. 219. 51.. Effect of dietary fiber on blood glucose.. et al. Role of dietary fiber on platelet adhesiveness. Nutr. 1979. 1.3_fm Page 393 Sunday. Appl. M. Jenkins. J. et al. 40. Treatment of diabetes with guar gum.. High fiber diets in the treatment of diabetes mellitus. B. Hagander. Doi. Am. J. 215. D.. et al.. K. 40. Clin. Br. Clin. 2001 7:34 PM TREATMENT OF DIABETES WITH HIGH-FIBER DIETS 393 30. J... 519. Tappy. Dietary fiber and diabetic therapy: a progressive effect with time. 482. M. A. 54. et al.. Nutr.. 42. Guar crispbread in the diabetic diet.. H. A. 55. Klin. . Diabetes Care.. J. Clin.. 53. Med. 453. Exp.. 67. 51. 2. 371. J.. Nutr. and triglyceride in obese diabetic patients. Bhatnagar. Med. 39. 108. Metabolic effect of pre-cooked instant preparations of bean and potato in normal and in diabetic subjects. Tsai. B. Hagander. A. Nutr. Clin. P. et al. Effect of guar gum on 24-hour insulin requirements of insulin-dependent diabetic subjects as assessed by the artificial pancreas. 1985. D.. J. Intern. 42. Eur. 36. L. J. 596. J. 50. 7. L.. Med. 985. S. 16. 1984. Effects of soy polysaccharide on postprandial plasma glucose. Treatment of diabetes with glucomannan (Konjac mannan). 1. D. 38. 39A. 3. 1984. 43. 1986. 52. 1555. D. 114. D. Reduction of urinary glucose loss in diabetics. J. F.. Diabetes Care. 1979. 1980... E. 38. Nutr. 1744. 49. 1990. Clin. Nutr. Nutr. Br. Korpela. 33.. J.. Hum. 1979. Parsons. Karlstrom. 35. Lancet. S.. 1023. Am. Blood glucose and serum insulin responses to breakfast including guar gum and cooked or uncooked milk in type 2 (non-insulin-dependent) diabetic patients. J. 43. 1. The effect of ispaghula (Fybogel and Metamucil) and guar gum or glucose tolerance in man. et al. E. Med. Am. lipids.. Effects of dietary fiber supplementation in stable and labile insulin-dependent diabetics. and Larsen. A. R. B. Smith. 1984. 1985. Improvement in long-term diabetic control after high fibre (bran and guar) diets. et al. L. 37. C. 68. 1982. The effects of a bulk laxative (metamucil) on fasting blood glucose. Aro. J. W. 41. 1981... Long-term effects of dietary fiber on glucose tolerance and gastric emptying in noninsulin-dependent diabetic patients. Low dose guar improves diabetic control. et al. 238. Gardner.. et al. I.. F. Am. R. J. A. Clin. P. 58. et al. R. Mayne. 1985. D. Glycosylated hemoglobin. J. P. 1981.2387_ch5. Acta Endocrinol. Br. Sci.. Botha. J. 1983. D. 20. A. S. Najemnik. 74. Br. 7. 46. 25. Bull. Soc. et al. G. H. M. F. High fiber and low fat diets in diabetes mellitus.. Laursten. Nutr. 83.. T. Dykes. Ther. J. Nutr. 200. 63. 872. 117. A.... O. et al.3_fm Page 394 Sunday. 78. D.. 1983. J. T. 78. R. and Lahikainen... 1983.. P. G. Clin.. Afr. Ford.. Med. 59. soy hulls of apple powder on glucose tolerance and plasma lipids in type II diabetes. 292. 61. S. J.. P. P. 41. Jones. Effects of nopal (Opuntia sp. Diabetologia. 30. 12. K.. M. Monnier.) on serum lipids. 6. et al. 57. C. Pediatr. G. Diabetes Metab.. Diabetes Care. and Hodgson. 85. 546. 137... 77. U.. McNaughton. D. 7. L. D. Guar gum and glycemic control of pregnant insulin dependent diabetic patients. et al. Eur.. 1353. 67. et al. N. 76. Assoc. F. Guar and its effects on metabolic control in type II diabetic subjects.. P. Med. J. Hallmans. 31. A. J. 1985... 1980. 66. 93. 1984. Nygren. H.. 1985. and other variables in constipated patients with non-insulin dependent adult diabetes. 1980. Diabetologia. 21.. Am Diet. Nygren. 1981.. Nutr. 73. 25. Molsted-Pederson. 70. 66. Fagerberg.. Med. 1985. Diabetic glucose control. Diabetes Care. and Milner. Int. 1981. D. S. 39. Frati-Munari. Diabetes Care.. K. Koepp. McIvor.. 376. 29. 1981. serum lipids. 143.. Dietary pectin and glycemic control in diabetes. J.. Nutr. A. A controlled trial of a high dietary fiber intake in pregnancy — effects on plasma glucose and insulin levels. 2001 7:34 PM 394 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Nutr. 1982. 75.. B. 81. J. 166. Kuhl. 1981. Kyllastinen.. 3RD EDITION 56. 1981. Effect of guar on plasma viscosity and related parameters in diabetic children... May 6. Long-term ingestion of guar gum is not toxic in patients with non-insulin-dependent diabetes mellitus. 60.. C. Decreased urinary glucose excretion and plasma cholesterol level in non-insulindependent diabetic patients with guar.. and Lithner. Nutr. Vaaler. Carroll. and trace elements on long-term guar.. H.. 7. 36. Ir. Am. et al. Ther. Diabetologia. et al. 45. Rep. S. R. J.. 79. The effect of a high bran diet on diabetes in mice and humans. (Mex. Effects of high bran bread on blood glucose control in insulin-dependent diabetic patients. Dan. et al. Effect of consuming fiber from corn bran. Med. M.. 80. Med. Diabetologia. 289.. Curr. 59. 65. 1981. J. 10. 1. and Holm. Jenkins. E. G. 31.. et al. J. 333. 151. blood glucose and serum cholesterol levels in diabetics treated with guar gum. 87. 19X3. Stokholm. C.Z. Am.. 152. A.).. 597. 42. Osilesi. 1984. 64. 94 (S 237). 25. Med. Res. 28.. Fraser. 72. . 1. Mahalko. 69. et al. and Hornnes.. Reduced glycosuria during guar gum supplementation in non-insulin-dependent diabetes. Changes in total serum cholesterol levels of diabetics fed five grams guar gum daily.. Effect of dietary fiber in insulin-dependent diabetics: insulin requirements and serum lipids. 1455. 31. 891. Harold. 1981.. C. 39. K. J. C. and Mendeloff. Ray. Curr. Improved diabetic control and hypocholesterolaemic effect induced by long-term dietary supplementation with guar gum in type II (non-insulin-dependent) diabetes... J. H. Diabete Metab. M. 82.. 62. Invest. et al. Clin. Effect of a modified guar gum preparation on glucose and lipid levels in diabetics and healthy volunteers. Clin. Cummings. glycemia and body weight. C.. and Hegewisch. Atherosclerosis. 505. 71. Guar gum is not a panacea in diabetes management. et al. 215. J. Use of xantham gum in dietary management of diabetes mellitus. Res.. Arch. 14. et al. 1983. 1984. Long-term dietary supplementation with a fiber product (guar gum) in elderly diabetics. Dodson. B. Johansen. The effect of apple fiber on diabetic control and blood lipids.. 1982. . Med. 57. 1105. Am. Lalor. A. Nutr. 1980. M. et al. Nutr.. Nutritional risk of high-carbohydrate.. 2. L. 1987. Med. Am. Hypolipidemic effects of high-carbohydrate.. R. et al. 1990. 3. Improved glucose control in maturity onset diabetes treated with high carbohydrate. 523.. 1989. Diab. No glycemic benefit from guar administration in NIDDM.2387_ch5. J. Paganus. 35. 77.. 380. O. et al... 4. Holman. 100. Atherosclerosis. Med. 105. 1985. J. Br.. 901. 48. Assoc.. 1986.. W. D. Metabolic and nutritional effects of long-term use of guar gum in the treatment of non-insulin-dependent diabetes of poor metabolic control.. 1980. Flendrig. 60.. et al. 7. K. A. J. P. K. K. 1990. 1147. 92. T. Am. 1986. high fiber diets for insulin-treated men with diabetes mellitus. 1. J.. et al. W. J.. 1980. Adherence to high carbohydrate. 106. J.. G. C.. Effects of alpha-glucosidase inhibition and viscous fiber on diabetic control and postprandial gut hormone responses. and Sieling.. Ebeling. Glucose and lipid metabolism and insulin sensitivity in type I diabetes: the effect of guar gum. Br. lipoprotein profile and gut hormones in NIDDM patients.. Metabolism. 230.. Anderson. et al. Moderate guar gum addition to usual diet improves peripheral sensitivity to insulin and lipaemic profile in NIDDM. P. Tagliaferro. and Postmes. 1985. B. Uttenhal. et al. 153. J. Uusitupa. 90. Diab. 88. R. 11. Beneficial effects of palatable guar and guar plus fructose diets in diabetic children.. et al. 1987. 1986. Beattie. and Sieling. Long-term effects of high carbohydrate. Groop. Sels. Lancet. 99. V. et al.. 1353.. 98.. M. 9. 109. 102. Effect of guar gum on mineral balances in NIDDM adults. Br.. Story. Van Duyn. 3. et al. guar gum dietary supplementation in non-insulin-dependent diabetes mellitus... Diabetic control is improved by guar gum and wheat bran supplementation. B. W. Beneficial effect of vegetable protein diet supplemented with psyllium plantago in patients with hepatic encephalopathy and diabetes mellitus. Low dose guar in a novel food product: improved metabolic control in noninsulin-dependent diabetes... 296. Nutr. 10. Dietary fiber enrichment. R.. Diab. J. Med. 345. Clin. P. 1979.. Chen. B. High carbohydrate.. 111. 1989. W. W.. et al. A. J. M. 1985. A. Res. et al. Med. et al. Z. N. 89. K. 1989. D. High fiber diets for obese diabetic patients. J.. et al. Eur. Long-term effects of guar gum on blood lipids. 505. Nutr... Rivellese.. 1988. J. 9.. 1979. et al. 93. blood pressure... Clin. Diabetes Care. Clin. Effect of dietary fiber on glucose control and serum lipoproteins in diabetic patients. and Ward. Diab. 357. L. 103. Br. 447. J. 1753. S.... and Taylor. The treatment of poorly controlled non-insulin dependent diabetic subjects with granulated guar gum. Obesity/Bariatric Med. Does adding fiber to a low energy. Vaaler. Clin. 177. and Bloom. Relationship between changes in GIP concentrations and changes in insulin and C-peptide concentrations after guar gum therapy. M. Diabetes Care. et al. 68. W. 108. 104. 7. J. M. R. high fiber diets on glucose and lipid metabolism: a preliminary report on patients with diabetes. Al-Hussary. J. 87. et al. Diabetes Care. 95.. 1979.. Effect of fenugreek seeds and leaves on blood glucose and serum insulin responses in human subjects. modified fat diet. J. 1. Requejo. Behall. et al. Clin. 1988. 1987.3_fm Page 395 Sunday. Diabetes Metab. Gastroenterology. Med. 101. The influence of guar gum bread on the regulation of diabetes mellitus type II in elderly patients. 85. Nutr. 110.. Anderson. Simpson. et al. and Ward. W. 32. S. J.-H. 1988. Uribe. J... 49. Med. Effects of cottonseed dietary fiber on metabolic parameters in diabetic rats and noninsulin-dependent diabetic humans. high fiber diets: long-term studies of non-obese diabetic men. 2001 7:34 PM TREATMENT OF DIABETES WITH HIGH-FIBER DIETS 395 84. Am.. 109. 107. Invest. Nutr. Diet. 7. R. High carbohydrate diets and insulin dependent diabetes. Placebo-controlled trial of the effects of guar gum and metformin on fasting blood glucose and serum lipids in obese. 97. Pract. 6. Lab. A. Diabetes Care. 85. 43. Madar. 98. 1987. 91. B.. 46. 1987. McIvor.. 29.. Res. Atkins... 118.. 76. T. L. May 6. 88. 1986. S. 1143. R. Anderson. F. Diab. 1978. Scand. ii. E. 551. 497. 242. 515. Clin. Sharma. Scand. Anderson. type 2 diabetic patients. 96. 76. 94. W. . Hagander. high carbohydrate. Nutr. Simpson. B.. et al. A. low fat diet confer any benefit to the management of newly diagnosed and overweight type II diabetics?. high fiber diets. 2312. Acta Paediatr. 12. 86. Peterson. J. 1986. 111. J. Hollenbeck. J.. Hum. W.. Whole foods and increased dietary fiber improve blood glucose control in diabetic children. 12. B. H. et al.. et al. Ney.. Diabetes Care. 116. Diabetes Care. et al. Med. Fiber and health: an overview. G.. 132. 44. 1982. 133.. 27. 2001 7:34 PM 396 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Pedersen. 1982. P. Riccardi. J. and Jackson. 127. Diabetologia. high starch. 121. W. non-insulin dependent diabetics. 1983.. 131.. 1983. et al. 2902. 272. Clin. Kay. N. Proc. et al.. 1982.. C. J. Serum lipids and lipoproteins in insulin dependent diabetic subjects during high carbohydrate. M. high fiber.. The effect of long-term high fiber diets in diabetic outpatients. low fat diets on glycaemic control in insulin-dependent diabetes mellitus. 5. Lindsay. R.. Rivellese. 1982. 26. Diabetes Care. Hum. 137. 1981. 123. W. 1984. Smith. et al. Comparison of the hypotensive and metabolic effects of bendrofluazide therapy and a high fiber. 63. A controlled trial of a high fiber. high fiber. 36C. O. C.. i. A.. and Allus.. 1983.. 114. J. Assessment of the metabolic effects of dietary carbohydrate and fiber by measuring urinary excretion of C-peptide. Separate influence of dietary carbohydrate and fiber on the metabolic control of diabetes. 522. 5.. Hollingsworth. Dodson. The effects of subject-selected high carbohydrate. 116. high fiber diet in children with type I diabetes.. Med. et al.. S.. Child. 641. Am. Taskinen. R. Barnard. 373. 135. Diets rich in natural fibre improve carbohydrate tolerance in maturity-onset. A. D. J. 5. 113. S. 1983. 1981. May 6. Physiologic and metabolic effects of dietary fiber. 7. C. Hjollund. 1983. 119.. 284. Eur. Hypertension. 187. High carbohydrate. Kinmonth. high fiber diet. et al. R. 41. Fed. et al. 235. Arch.. 1982. E. Diabetes Care. Response of non-insulin dependent diabetic patients to an intensive program of diet and exercising... J. 310..3_fm Page 396 Sunday. M.. low sodium diet in diabetic subjects with mild hypertension. Diabetologia. A. M. M. Am. low fat diet. Diabetes Care. Gastroenterol. L. 136. low fat and low sodium diet for mild hypertension in type 2 (non-insulin-dependent) diabetic patients. Hoffman. Simpson. Diabetologia. Invest. Med.. 1984. 215. R. Manhire... 32. 1986. 81. et al. 1985. H. 93. J. 1984. 122. et al. The effects of a high fiber. H. Clin. Effects of an increased content of cereal fiber in the diet of type II (non-insulindependent) diabetic patients.. Postgrad. Pacy. A. G. Prev. 6. E. 293. J. Burke. 6. et al. J. 26. 126. Long term use of a high complex carbohydrate. Diabetologia. 1982. 124. high carbohydrate diet in very obese diabetic individuals. 117. A. and Cousins.. et al. low fat.. Br. 892. Grobin. 120. Barnard. Dodson. 23. Ward. Anderson. 370.... 1984. 57. Unrefined carbohydrate and dietary fibre in the treatment of diabetes mellitus. Simpson. 12. U. 1984. 63. P. M. L. low fat diet and exercise in the treatment of NIDDM patients. et al. Anderson. 1982. J.. J. 5. . Decreased insulin requirement and improved control of diabetes in pregnant women given a high carbohydrate. 1982. D. C. Dis. C. et al. Simpson. N. 3RD EDITION 112. J. et al. 20. 128. P. W. J...2387_ch5. 118. Metabolism.. P. B. 35.. 1981. Nutr. 18. A high carbohydrate leguminous fibre diet improves all aspects of diabetic control. 134. Nikkila.. 59.. 1608. 99. J. Reduction of risk factors for atherosclerosis in diabetic patients treated with a high-fiber diet. 129. 2. Med. 224. Increased insulin receptor binding to monocytes from insulin dependent diabetic patients after a low fat. 115. Digestible carbohydrate — an independent effect on diabetic control in type 2 (non-insulin-dependent) diabetic patients?. Nutr. A. et al. Short-term effects of a high fiber. 605. Diabetes Care. 130. high fiber diet. 268. Clin. and Track. M.. et al. 1. 1983. 1982.. Karlstrom. 1067. Diabetologia. J. Nutr. Rosman. B. S. Diabetes Care. 529. Effect of high fiber diet on haemostatic variables in diabetes. Insulin receptor binding increased by high carbohydrate low fat diet in non-insulindependent diabetics. low fat and low sodium dietary regime on diabetic hypertensive patients of different ethnic groups. 125. 284. et al. 128. R. 1985. et al. Afr. Nutr. Lancet. R. C. Increased insulin binding to adipocytes and monocytes and increased insulin sensitivity off glucose transport and metabolism in adipocytes from non-insulin-dependent diabetics after a low-fat/high starch/high fiber diet.. R. Nutr. Garg. Am. 7. 208. Adherence to high carbohydrate. W. Am. Diabetes Care. Am.. P. et al. May 6. 140. 3rd ed. 64. 1986. 151.. low sodium diet on diabetics with intermittent claudication. J..3_fm Page 397 Sunday. Med. Anderson. Comparison of a high carbohydrate diet with a high monounsaturated fat diet in patients with non-insulin-dependent diabetes mellitus. 16. A high fiber diet in gestational diabetes-wheat fiber. 5. Stevens. 149. 829. Diabetes Care. C.. 48. K. 15. Obesity. M. Paisley. Teuscher. P. low fat. Med. J. Karlstrom. G. F. Clin. Coulston. A. Hollenbeck. 28. 143. 1985. and fiber on carbohydrate and lipid metabolism in type II diabetes. 343. Nutrition management of diabetes mellitus. Nutr. Nutr.. Dietary fiber: diabetes and obesity. Nutr. P. M.. A prospective comparison of “conventional” and high carbohydrate/high fiber/low fat diets in adults with established type I (insulin dependent) diabetes. 199. 152. 1988.. Dodson. et al. Diabetes Care. W. 48. 154. Diabetologia. 1988. 94. Philadelphia.. Coulston. 158. leguminous fiber or both?. B. Effects of a high-starch diet with low or high fiber content on postabsorptive glucose utilization and glucose production in normal subjects. 1988. Hagander. Eds. Appl. and Bryant. Diabetes Care.. N.. Pract. Res. A. Anderson. V. basal insulin supplements. J. 145.. and Chantelau. Diabetologia.. A fiber-rich diet for the treatment of diabetic patients with chronic renal failure. et al. 255. New York. D. 160. L. Scott. E. 139. M. J. A. R. 41A.. G. 1989. 89.. 146. 156. J. B. A. 1990. 116. Clin. 1991.. To what extent does increased dietary fiber improve glucose and lipid metabolism in patients with noninsulin-dependent diabetes mellitus (NIDDM). P. 144. 429. Effect of a high carbohydrate. Clin... Lea and Febiger. 1007. K. J. high fiber diets. Am.. J. J. M. 153. 8.. Pacy. 153. Clin. and Taylor. Dodson. 8. Temporal study of metabolic change when poorly controlled noninsulindependent diabetics change from low to high carbohydrate and fiber diet. Hartog. 1985. Venhaus. 1985. Wechenschr. Metab. M. 359. M. J. et al. 157. Comparison of high fiber diets.. high-fiber diet to diabetic patients with chronic kidney failure. Nestel. Rivellese.. Clin. C. J. R. 159. Engl. Persistence of hypertriglyceridemic effect of low-fat high-carbohydrate diets in NIDDM patients. 81. 142. 148. Clin.. in Modern Nutrition in Health and Disease. . and flexible insulin treatment for non-insulin dependent (type II) diabetics poorly controlled with sulphonylureas. 147. 162. et al. and Reaven. Br. 620. et al. low fat diet: effect on lipid and carbohydrate metabolism. Nutr.. B. A. et al. W. 1987. 1986. 707. 14.. Diabetes Care. et al. 898. Outpatient management of diabetes mellitus with patient education to increase dietary carbohydrate and fiber.. Dan. 1988. Effects of dietary carbohydrates on metabolism of calcium and other minerals in normal subjects and patients with non-insulin-dependent diabetes mellitus. Simpson.. 1988. Nutr. Dietary fiber in nutrition management of diabetes. V.. and Fletcher. 43. B. J.. Med. A. 319. Self-selected unrefined and refined carbohydrate diets do not affect metabolic control in pump-treated diabetic patients. 95. A. 1985. et al.. 1986. Parillo. J. 150. Am. 155. J. 1076. W.. Brand. et al.. Diab. Garg. A. carbohydrate. 313. et al. 1988. Am. 1989. 104. et al. 32. and Kritchevsky. Effects of leguminous seeds in a mixed diet in NIDDM (non-insulin-dependent) diabetic patients. R. Assoc. 10. 70. N. J. Int. 1988. Gastroenterol. Anderson.. GIP and insulin secretion in diabetics.. 1986. Bull... M. et al. 1987. 282. et al... 1986. J. Hum..2387_ch5. J. Pacy. C. 31. et al. 2001 7:34 PM TREATMENT OF DIABETES WITH HIGH-FIBER DIETS 397 138. and Young.. 1986. J. 161.. 43.. 12.. P. 146. P. in Dietary Fiber (Basic and Clinical Aspects). and Savage.. High carbohydrate. Anderson. low sodium and low fat diet in type 2 diabetics with moderate hypertension.. Diet. E. Schweiz. Die kohlenhydrate und nahrungsfasern in der diabetesdiat.. Sestoft. R. The effect of a high fiber. 1989.. D.. W. Vahouny.. The effects of diet differing in fat. 141. Br. Diabetes Educator.. Eds. J.. J. Low glycemic index foods improve long-term glycemic control in NIDDM. M. G. Metabolic consequences of feeding a high-carbohydrate. 852. Endocrinol. McCulloch... J. Dietary fiber decreases fasting blood glucose levels and plasma LDL concentration in noninsulin-dependent diabetes mellitus patients. and Gustafson. 1984. Plenum Press. Shils. 46. 47. K. O’Dea. A. J. 297. Med. 207. Diabetic Medicine. 1126. 168. Ciavarella. Diabetic Medicine. 77. 289. 49. M. W. 171. 137. Pathol. 358. J. 29. J.. 1984. 1334.. Nutr.. Blanchi. 300. Pharmacol. J. Pharmacol. 263. P. 176. Diabetes Care. D. Akanji. Ann.. 717. J. Commun. 1996. Short-chain fatty acid fermentation products of plant fiber affect glucose metabolism of isolated rat hepatocytes. R. 1995. Proc. W. Nutr. E.. 1996.. Effects of changing amount of carbohydrate in diet on plasma lipoproteins and apoproteins in type II diabetic patients. Effect of lactulose on carbohydrate metabolism and diabetes mellitus. Rev. Rivellese. Change in plasma acetate levels in diabetic subjects on mixed high fiber diets. 222. Chaturvedi. High carbohydrate. Frati-Munari. 13. K. R. KY. Ronchi. 1990. Fibre in the management of diabetes. Med. 187. 174. The effect of guar gum on the acute metabolic response to glyburide. 44.. and Hockaday. American Heart Association. 333.. 1336. 1989. P.. 1988. A. Chem. Diabetic Medicine. N. CRC Crit. Feldman. A. J. Trinick. Judd. Plasma glucose and insulin.. J. Fibre in the management of diabetes.. Med. Brit. Lexington. Effects of breakfast cereals containing various amounts of betaglucan fibers on plasma glucose and insulin responses in NIDDM subjects. 10.. et al. 188. G. Effect of vegetable fibre on postprandial glycemia. K.. and Chaturvedi. Am. Gugolz.. 165. insulin.. Postprandial glucose. 831. et al. 74. May 6. L. B. R. Gastroenterol. 184. Absorption characteristic of breakfast determines insulin sensitivity and carbohydrate tolerance for lunch. Glycaemic responses in type 2 diabetic patients to various mixed meals taken at home. S. Effect of guar on second-meal glucose tolerance in normal man. Am. Clin. and Akanji. High beta-glucan oat bran and oat gum reduce postprandial blood glucose and insulin in subjects with and without type 2 diabetes. Dietary fiber and coronary heart disease. 1993. Nutr. 1990. Miller. Metab. J. et al. 180. Braaten. 54. E. 33. Diabetes Care. and Mansell.. et al. Sci. Food Sci. hypertriglyceridemic subjects. 13. 1990. 177. 1988. 185.. 1987. Guevin. 71. 62. 2001 7:34 PM 398 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.. 524. I. Tappy. 183. Biol. Arch. 10. 14. Exp.. A. J.. . and Marchesini. et al. J. D. Nutr. Enrichment of an Israeli ethnic food with fibres and their effects on the glycaemic and insulinaemic responses in subjects with non-insulin-dependent diabetes mellitus. N. O. Gatenby. 29. A. Res. Anderson. Clin. R. Nutrition Management of Metabolic Conditions. J. T.. Br. Circulation. 137. N. Effect of partially depolymerized guar gum on acute metabolic variables in patients with non-insulin-dependent diabetes. 312. 1990. Scand. and Wursch. S. J. et al. 173.. Hunpponen. 172. J. Benefits of fibre itself are uncertain. 166... 1990.. A. Nestler.. 179. 19. Soc. 813. D. Diabetes Care. 28. Fukagawa... 177. 1991. 1989. W.... 755. A.. high fiber diets increase peripheral insulin sensitivity in healthy young and old adults. 654. Dietary fiber — an overview. E...3_fm Page 398 Sunday. et al. 175. Harnden. Tattersall. J. Hunpponen. S. 1996. F. et al. 3RD EDITION 163. et al.. P. Diabetes Care. 11. J. 167. J.. Plant Foods Human Nutr. Anderson.. C.2387_ch5. 681. Reduced albuminuria after dietary protein restriction in insulin-dependent diabetic patients with clinical nephropathy. Diabetes Care.. S. 11. 446. 1986. II. 84.. J. Dietary guidelines for healthy American adults. et al. et al. 95.. J. W. 372. 1986. Hockaday. Res. et al. 1994. Effect of guar gum on glipizide absorption in man. 71. and Bridges. T. G... O. Br. T. Clin. Am. P.. 1997. R. 1990.. Coll. Position statement. 181.. Lowering glycemic index of food by acarbose and Plantago psyllium mucilage. I. Gastroenterol. (suppl. Nutr. Anderson. 1365.. Natural fibre useful as part of total dietary prescription. Med. and Dekker. Miller. urinary catecholamine and cortisol responses to test breakfasts with high or low fiber content: the importance of the previous diet. Small bowel obstruction from bran cereal. T. 721A. Sundell. 170. A. 178. and Seppala. 182. Eur. 1998. 164. L. 1985. HCF Diabetes Research Foundation. and lipid responses to four meals containing unpurified dietary fiber in non-insulin dependent diabetes mellitus (NIDDM). P. 186. et al. 169.). 1993. A.. 52. JAMA. R. 300.. Anderson. 15.. 407. 1981. Med. Karhuvaara.. Assoc. 11. 49 (suppl.. J. . 972. total cholesterol and triglycerides in patients with diabetes mellitus. Clin. Vuksan. 35. 192. Am. W. P. Anderson. 1992. 111. Tox. Nutr. 58.. Diabetes Care. J. Effects of different glycaemic index foods and dietary fibre intake on glycaemic control in type I diabetic patients on intensive insulin therapy. L... M. Huang.. 15. Voprosy Pitaniia. A. (suppl. 123. Clin.. Clin.. 118. et al. 61. Rev. wheat bran and placebo on carbohydrate and lipid metabolism in type II diabetics. S.. and Tsagikian. Int. 213. 1995. M. 1996. 402. D. S. 1358. Metabolic effects of hypocaloric high-carbohydrate/highfibre diet in non-insulin dependent diabetic patients. Clin. et al. Pick. Comparison of guar gum. 1990. J.. Biosci. 52. et al. J. 17. 22. 196. 203. 1997. A randomized controlled metabolic trial. Sinisalo. 1990.. Guerrero-Romero. S242. and Koivisto. Chuang.. 1992. 2001 7:34 PM TREATMENT OF DIABETES WITH HIGH-FIBER DIETS 399 189. Hanai. Am. Biotech. et al. 333. M. et al. Long-term comparison of three dietary prescription in the treatment of NIDDM.. 121. 195. and Gianino. Diabetes Care. D. 13. 96. O. J.. and Lazcano-Burciaga. Biomed. 201. and Mogila. et al. J. Nutr. The effects of a dietary fiber (white lupine bran) in the treatment of non-insulindependent diabetes. J. Long-term effects of guar gum in subjects with non-insulin-dependent diabetes mellitus. Effect of soya and cellulose fibers on postprandial glycemic response in type II diabetic patients. 211.. 56. 1998.. 54. Minerva Pediatrica. Schweiz. et al. et al. H. 513. 191.. Librenti. V. 913.. 190. et al. 1991.. 1999. 198. Carra. 209. Vuorinen-Markkola. et al. Stahl. 1991. Milne. 1998... 15. Metabolic effects of high-carbohydrate. Clin. J. Kh. 194. Krashenitsa. Sharafetdinov. Kirsten. Diabetic Medicine. 3. Am...... 15. et al. 208. 1992. 26. Am. 1056.. Therapeutic effect of guar gum in patients with non-insulin-dependent diabetes mellitus. Sci. 12. 10. E. A small dose of soluble alginate-fiber affects postprandial glycemia and gastric emptying in humans with diabetes. 1994. et al.. Coll. M. Diaz. 24. Eur. 3. Diabetes Res. et al.... Comi. M. Nutr. Effectiveness of increased contents of dietary fiber in early stages of non-insulin dependent diabetes mellitus. Nutr. 273.. et al. et al. 212.. Oat bran concentrate bread products improve long-term control of diabetes: a pilot study. high-fiber diets for insulin-dependent diabetic individuals.. Clin. M.. 65. Diabetes Care. Diabetes Complications. A. Bruttomesso. 193. 24. A. Effect of Konjac food on blood glucose level in patients with diabetes. 30. Prac. 205. and Berger. Am. 204. 9. et al. I.. Psyllium fiber reduces rise in postprandial glucose and insulin concentrations in patients with non-insulin-dependent diabetes. W. Pastors. C. 362. et al. Prac.. 1992. Diabetes Res. L. Ther. M. Clin..and glucose-lowering efficacy of Plantago psyllium in type II diabetes.. M. Groop. J. 200. 1990. 1991.. 206.3_fm Page 399 Sunday. Voprosy Pitaniia. M.. 1990. Blood sugar response to administration of bran or guar added to pasta in children with type I diabetes. R. Am. 1991. C. 53. 199. J. 42. Guar gum in insulin-dependent diabetes: effects on glycemic control and serum lipoproteins. G. A.. J. Nutr. M.. Konjac-mannan (glucomannan) improves glycemia and other associated risk factors for coronary heart disease in type 2 diabetes. 210. T. Nutr.2387_ch5. T. G. 1254. Absence of guar efficacy in complex spaghetti meals on postprandial glucose and Cpeptide levels in healthy control and non-insulin-dependent diabetes mellitus subjects. 1994. Formosan Med. 1992. Assoc.. 74. et al.. No effects of high-fiber diets on metabolic control and insulin-sensitivity in type 1 diabetic subjects. Y. 197. 1431.. G. V. J. Influence of two guar preparations on glycosylated hemoglobin. Biochem. 15. V. Chile.). Effect of method of administration of psyllium on glycemic response and carbohydrate digestibility... 207. Rodriquez-Moran. H. 4. H. Clin. 1993. Wschr. 1991.. Res. M. and Efendic. 3). 1993. Long-term effects of water-soluble corn bran hemicellulose on glucose tolerance in obese and non-obese patients: improved insulin sensitivity and glucose metabolism in obese subjects. Sels. Med. 795. Lafrance. Armyr. J. Diet. Hormone Metab.. L. Nutr. 936. May 6. P. Plotnikova. Karlander.. J. Env. Botvineva.. J. Lipid. Metabolic effects and clinical value of beet fiber treatment in NIDDM patients.. Brugnani. Comparative efficiency of various food fibers in the correction of carbohydrate and lipid metabolism in patients with type II diabetes mellitus. A.. 582. F. 202. Med. R. Torsdottir. 91. Wolever. I. 1991. Pharm. 785. 334. Am.. J... Dietary fiber. Anderson. 1999. W. Diabetologia.. glycemic load. Effect of dietary fiber and protein intake on blood pressure: a review of epidemiologic evidence. Med. 1392. Am. Clin. Nutr. et al. M. 16. 1999. W. H. Hyperinsulinemia as an independent risk factor for ischemic heart disease. Nutr. G. 1997. Lintas.. 18.. Antioxidant supplementation effects on low-density lipoprotein oxidation for individuals with type 2 diabetes. J. M. J. Ed. 277.. 219. 226. 307. Am. 343. Comparison of effects of high and low carbohydrate diets on plasma lipoproteins and insulin sensitivity in patients with mild NIDDM. 20. 215. A.. K. 223. Exp. M. 291S. 451. He. High-carbohydrate. 1278. New Engl. D.. in CRC Handbook of Dietary Fiber in Human Nutrition. Clin. Dietary fibre. M. 49 (suppl. K. et al. 52. Glycaemic and insulinaemic responses in NIDDM patients. W.. S264. Chandalia. Med. Beneficial effects of high dietary fiber intake in patients with Type 2 diabetes mellitus. Coll. R. Am. N. Am.. Hypertension. Diabetes Care. Garg. 2000. Stampfer. Dietary fiber. 430. J. Anderson. Med. Despres. et al. O. C. Dietary fiber. et al. and risk of non-insulin-dependent diabetes mellitus in women. 229.. glycemic load and risk of NIDDM in men. J. Garg. Engl. J. CRC Press. Bessesen. R. 1994. J. Nutr. et al. Boca Raton.. J.. 40. Whole grains and protection against coronary heart disease: what are the active components and mechanisms?. 230. N. Med. J. and Hanna. A. J. Anderson. A.. Assoc.. Clin. 1999. weight gain. 21. Baltimore. 70. 1990. 222.2387_ch5. et al. 221. 224. 1996. Treatment of diabetes with high fiber diets. 524. Primary prevention of coronary heart disease in women through diet and lifestyle.. 1421. and Whelton. et al. 282. Med. H. 3RD EDITION 214. et al.. 1539.. E. Lugwig. Whole grain foods and CHD risk.. Diabetes. W.. Effects of varying carbohydrate content of diet in patients with non-insulin-dependent diabetes mellitus. Clin... in Modern Nutrition in Health and Disease.. A. FL. J. 220. J.. 2000.. et al. J. J. Spiller.3_fm Page 400 Sunday. 1992.. J. 545..-P. Salmeron. J. 1997. 1999. high-fiber diets increase peripheral insulin sensitivity in healthy young and old adults. P. Williams & Wilkins.. 271. and Unger. 2000. Am. Am. and Hamman. J. 1993. T. Nutr. 19. 1997. Shils. 952. 342. Grundy. and Akanji. et al. Assoc. S. J. 3). Nutr.. Med. Anderson. Shike. J. et al. 41. Salmeron. .. May 6. Anderson. D. Assoc. M. Engl. A. J. resistant starch and in vitro starch digestibility of cereal meals. 1365. Nutritional management of diabetes mellitus. Coll. 227. 1999. Eur.. High saturated fat and low starch and fibre are associated with hyperinsulinaemia in a non-diabetic population: the San Luis Valley Diabetes Study.. 216. Eds. and cardiovascular risk factors in young adults. 217. et al. J. 228. 1995... 472. W. Marshall. 218.. 443. J. J. 225.... Fukagawa. F. N. Olson. A. 2001 7:34 PM 400 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 4. Table 5. % reduction) 5 5 5 5 10 5 5 40% Depending on the sterol and stanol.2387_ch5. May 6.00+$1. and plant sterols — into a “cholesterol-lowering diet portfolio” may permit a diet to reduce serum lipids to a similar extent to a starting dose of a statin.4_fm Page 401 Sunday. Source: Adapted from Jenkins et al.1). Furthermore. for cholesterol lowering and cardiovascular risk reduction.S. Reduce trans-fatty acids to as close to zero as possible.3 One is tempted to speculate that studies of viscous fibers opened the way for the systematic assessment of food components with cholesterol-lowering properties.2.4 Fiber in the Treatment of Hyperlipidemia Alexandra L. The others include soy proteins and most recently plant sterols. barley.4.50 © 2001 by CRC Press LLC 401 . a combination of these four elements — viscous fibers. Assuming the effects are additive. the currently preferred treatment for cholesterol reduction (Table 5. and legumes) as part of the strategies to reduce the risk of CHD. Vladimir Vuksan. These two fiber sources were the first of the four food components to be permitted by FDA to make health claims. Food and Drug Administration (FDA) has allowed two viscous fiber health claims. Soy protein Plant sterolsa Dietary cholesterol Saturated fatb Body weight Low-glycemic-index diet Total a b c Amount 5–10 g/day 25 g/day 1–3 g/day <200 mg/day <7% of calories Lose 10 lb Reduce by 10 GI units Full portfolioc Reductions in LDL (approx. and David J. A.151 0-8493-2387-8/01/$0.1 Dietary Factors Portfolio for Cholesterol Reduction Dietary Strategy Viscous fibers: Psyllium Guar Konjac Beta-glucan.1 Indeed. low-glycemic-index foods. 2001 7:37 PM CHAPTER 5. the U. vegetable protein. and national agencies concerned with cardiovascular health also endorse the use of soluble fiber foods (oats. one for oats and the other for psyllium. Jenkins. Jenkins INTRODUCTION It is now well accepted that viscous types of fiber may lower serum cholesterol levels. etc. 23 The nature of the fermentation and the type of fiber may therefore be important in determining the final outcome.7 mmol/l/g of psyllium. .14. Increased Fecal Sterol Losses From the beginning. 3RD EDITION META-ANALYSIS There are a number of meta-analyses of the effects of viscous fibers in lowering serum cholesterol. when colonic fermentation is increased using the non-absorbable sugar lactulose. soy protein.4. this meta-analysis did not include all the studies which were used by the FDA in determining health claims. 2001 7:37 PM 402 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and further data are therefore urgently required. such as incorporating viscous fibers.15 Studies have been limited in number due to the unsavory nature of this line of work. looked at four viscous fibers: pectin. none of which are mutually exclusive in providing an explanation of the mechanism for an individual food. human-feeding studies of propionate20.4.12. On the other hand. and different mechanisms are likely to predominate depending on the fiber.5 alone found reductions in LDL-cholesterol of 0. Propionate has also been demonstrated in humans to inhibit.21 have not demonstrated a clear effect in reducing LDL-cholesterol levels. acutely. plant sterols. Indeed. Implementing a range of dietary strategies. which are absorbed.4_fm Page 402 Sunday. Increased Short-Chain Fatty Acid Generation Bacterial fermentation of fiber in the colon gives rise to short-chain fatty acids (SCFA). the acetate-induced rise in serum cholesterol after rectal infusion. it is likely that for a given food. May 6.7 may result in clinically significant reductions in serum cholesterol (Table 5. Other meta-analyses on psyllium4. The authors emphasized the smallness of the reduction of LDL-cholesterol (0. Together with this mechanism of action.13 but the effect of fiber in foods is less clear. there are possibly three other broad reasons why fiber lowers serum cholesterol levels. has been shown in pigs16 and rats17 to reduce serum cholesterol levels and to inhibit cholesterol synthesis in liver in vitro. it represents a potential building block for a dietary portfolio (Table 5.22 Another mechanism through which SCFAs may reduce cardiovascular risk factors is through their effects on clotting factors. oat bran.23 Increasing acetate levels by feeding either pectin or acetate increases permeability and lysability of fibrin networks while at the same decreasing the tensile strength. propionate. DIETARY FIBER AND POSSIBLE MECHANISMS OF ACTION A number of possible mechanisms are likely to be involved in the hypocholesterolemic effect of dietary fiber.1).57 mmol/l/g fiber) with no major differences between the fibers.1). The pioneer studies of Kritchevsky and colleagues demonstrated that a number of fiber sources were capable of binding bile acids in vitro 8–10 and provided a rationale for the increased bile acid losses seen in vivo.8 There is general agreement that purified viscous fiber11 administration increases bile acid outputs by 20 to 80%. and psyllium.6 However.2387_ch5. it was recognized that increased fecal sterol losses provided one explanation for the lipid-lowering effect of fiber. This area also requires further studies for its definition.4–6 The meta-analysis by Brown et al. Although this reduction is not equivalent to reductions seen with standard drug therapies. guar. and low-glycemic-index foods together with a National Cholesterol Education Program (NCEP) step 2 diet. LDL-cholesterol levels appear to rise rather than fall. One of the SCFAs. more than one mechanism is operative.23 This effect is seen despite total plasma fibrinogen levels remaining unchanged.19 However. in the longer term. reduction in carbohydrate intake may minimize differences in glycemic response and. received this degree of scrutiny. only those with the E3 phenotype have shown an increase in apparent fat absorption. May 6.33. may alter small intestinal morphology and lipid absorption. these are simply speculations to be explored. and bile acid excretion. The literature relating to healthy volunteers has already been reviewed. however.53–57 There are exceptions to this generalization. as yet. presumably due to an increase in the bile acid pool and chylomicron formation.29 Early studies demonstrated that hepatic cholesterol synthesis in the rat increased during periods of maximum insulin secretion. a rate-limiting step in cholesterol synthesis. 2001 7:37 PM FIBER IN THE TREATMENT OF HYPERLIPIDEMIA 403 Reduced Insulin Levels Increased insulin levels have been linked with CHD.e.4_fm Page 403 Sunday. multicenter population-based cohort study. The present discussion will therefore focus on the therapeutic use of fiber. however. fiber foods which induce a bile salt loss might be more effective if the bile salt pool is expanded through greater intakes of dietary fat.37 Insoluble fiber does not seem to alter fat absorption. cholesterol absorption. On the other hand. the suggestion that lignin may be hypocholesterolemic by virtue of its bile acid–binding ability resulted in two conflicting clinical studies.41–46 this genetic classification may be particularly useful in predicting responses to altered fat and fiber intakes. i. Vitamin A tolerance tests with added fiber indicate that some fibers appear to enhance chylomicronemia. and .2). viscous fibers have proved useful in lowering serum lipids..37 Added to this are the genetic differences which may make fiber more or less effective. In the absence of studies. and vegetable protein39.47–52 while nonviscous fibers have for the most part been without effect. fiber consumption predicted insulin levels and other cardiovascular risk factors more strongly than did total or saturated fat consumption in a 10-year.4. EXPERIENCE WITH SPECIFIC FIBERS The early studies described the hypolipidemic effects of fibers in healthy volunteers before they were tested on patient groups. including a prudent diet. differences in the apo E genotype and dietary change.28. Hypothetically.30 The cholesterol-lowering effect associated with reduced insulin levels has been confirmed using a model of altered food frequency (“nibbling”)31 to mimic slow absorption.37 Other genetic markers have not.40 have attracted much attention and may influence the response to drugs such as gemfibrozil.59 The dosages were small (Table 5.38 In view of the association of E genotype with differences in remnant particle uptake. In general.58 Lignin and Cellulose Early on. Among genetic variants which influence serum lipids. Viscous fiber seems to equally effective in lowering serum cholesterol in both E3 and E4 phenotype.2387_ch5. No detailed studies have been carried out at different levels of dietary fat to assess the effect on the different possible mechanisms of action of dietary fiber.32 Altered Lipid Absorption and Genetic Factors Fiber delays the rate of nutrient absorption29 and.24–27 A common effect of the viscous fibers and high-fiber foods which reduce serum cholesterol levels is that they produce relatively flat postprandial glucose and insulin responses.34 Alteration in the rate and site of lipid absorption may alter the pattern of lipoprotein secretion35 and catabolism. hence. irrespective of phenotype.36. Finally.30 The explanation was that insulin induced an increase in activity of hepatic HMG-COA reductase. dietary cholesterol. whether some mechanisms are more or less effective at different levels of dietary fat intake. mechanisms which relate to altered insulin secretion.54. Note: Abbreviations: TG.6 NS –5. 3RD EDITION . low-density lipoprotein cholesterol. total-C. 2001 7:37 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. NC. HLP. HC. triglycerides. high-density lipoprotein cholesterol.8 NS 6. hypercholesterolemia.7 NS 0 –2. Cellulose Lignin Fiber Effects of Lignin and Cellulose on Serum Lipid Concentrations 404 Table 5.6 NS NC TG (% change) NC in FBG. total cholesterol.2 2387_ch5. HDL-C. hyperlipoproteinemia.4_fm Page 404 Sunday.4. LDL-C.6 NS +8 NC NC –21 Total +6 NS NC –10. May 6. or HbA1c –22% change in total-C on test and control Celluline duplicated effect or maintained effect of cholestyramine Comments 55 55 56 53 56 54 59 Ref.3 NS Cholesterol (% change) LDL VLDL HDL +8 NS 1. no change.15 15 2 12 60 4 Dose (g/day) Soyhull Normal diet Normal diet Normal diet Normal diet Cholestyramine Control 3 mo 12 wk 4 wk 4 wk 4 wk 2–5 mo Duration 14 hyperlipidemia Type 2 diabetes 10 healthy 7 hyperlipidemia 10 healthy 60 healthy 6 hyperlipidemia Subjects NC –11. 47. Many studies have been conducted to explore the lipid-lowering effects of psyllium.5.60–82 including three meta-analyses4.88–93 Nevertheless.4.0 7. Psyllium consumption of 9. No changes were seen in serum triglycerides.2 4. baked into conventional breads.2387_ch5. all were equally effective.47.4.48 A clear dose–response is not evident from these studies.88–93 The cholesterol-lowering results with guar were materially the same as those observed with pectin.4.55 cellulose.6 Improvements in the ratio of Apo B to Apo A-1 are also observed. but the reduction was significant only when the guar was incorporated into very low fat. Psyllium has long been accepted as a safe and reliable bulk laxative. May 6. in a study in which the same guar was added in powder form to fruit juices and soup.54. Similarly.3). to support the earlier work87 (Table 5. Palmer and Dixon demonstrated that little cholesterol-lowering effect could be seen in healthy individuals taking 6 g or less of pectin daily. since greatly differing effects were reported by different investigators when doses of guar of the order of 15 g/d were given.4. and hence needs to be mixed intimately with the food.2).49.4 7. 2001 7:37 PM FIBER IN THE TREATMENT OF HYPERLIPIDEMIA 405 the work has not been repeated. the particulate fiber.86 However. psyllium too seems to exerts its greatest effect only when mixed with foods. a substantial lowering of serum total and LDL-cholesterol levels was observed even when as little as 12 g of pectin daily were taken. The observation that the resulting falls in serum cholesterol levels were associated with only modest increases in fecal sterol loss suggested that increased sterol excretion was likely to be only one of a number of mechanisms responsible.37 4.89 These findings suggested that prehydration was not a prerequisite for the hypolipidemic action .4_fm Page 405 Sunday.3 Meta-Analysis on LDL Reduction by Psyllium No. such as crisp bread or spaghetti. in general. Consistent with the postulated mechanism of action that fiber delays the rate of nutrient absorption. or incorporated into a dry crisp bread or melba toast-type formulation.49.4. with much less or no change in the HDL-cholesterol fraction.5.4.1–10. Triglyceride levels were reduced. 9.2 g/day is associated with reductions in LDL-cholesterol of 6–7. with no change in HDL-cholesterol level. The interest in pectin has continued.79 0. Guar Again following observations in healthy volunteers. The effect was predominantly reflected in the LDL-cholesterol fraction.4). starchy carbohydrate foods.52. was without effect on serum cholesterol or triglyceride levels (Table 5.83 Table 5.4 10. Psyllium Psyllium is a concentrated source of soluble fiber derived from the husks of blonde psyllium seed.19 6.2%.4).33 4.151 Pectin The early studies of Palmer and Dixon84 in normal volunteers were followed by those of Miettinen and colleagues85 on hyperlipidemic patients who consumed relatively large doses of pectin (40 to 50 g/d) (Table 5.6 (Table 5.1 9.70 6 4 5 Source: Adapted from Jenkins et al.2 0.89 The physicochemical nature of the guar and the formulation in which it is provided may be very important factors.84 Subsequent studies in hyperlipidemic patients confirmed this observation.66 0.47. of Subjects Taking Psyllium 479 209 384 Average Dose (g/day) Mean LDL Starting Value (mmol/l) LDL Reduction (%) Reduction per Gram Psyllium Ref. studies testing the effects of guar were undertaken in hyperlipidemic patients. 4 9–12 Metamucil Ground husk Cereal Vi-Siblin Metamucil Psyllium Granules 57 12 5–15 6 6 15 15 40–50 Dose (g/day) wk wk wk wk 26 wk 8 wk 2 wk 6 wk 24 wk 4 wk 4 wk 90 d 12 wk 8 wk 8 wk 3 wk 6 wk 11 d 13 wk 5 wk 6 wk 4 wk 8 wk 6 6 3 4 2 wk Duration 250 HLP 34 type 2 diabetes.3 –23.0 6.7 –9. May 6.4 –19.2 1. 3RD EDITION .7 10.7 45 NC 1.8% 50 subjects acted as control group –6% MUFA diet –12% MUFA diet LDL:HDL decreased 13.6 NS 20.6 –26.1 NS NC 3. 2001 7:37 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.2 15 10.0 –7. 406 Cereal Citrus K-Pectin + Gum Arabic Lemon Apple Citrus Grapefruit Pectin Fiber Table 5.5 NS –8.2 NS –5 NS HDL Effects of Soluble.4.2 2.4 NS Cholesterol (% change) LDL VLDL NC NC –3.6 –3.8 NS –10.5 2.2 –1.2 22.2 –8.3% 8.5 +5 –4 NS 18 –7.2 Cereal Metamucil Metamucil Husk Cereal 11 21 57 30 20.7 –8.6 –13 Total –6.7 NS +0.8% fall in Apo B on psyllium Weight loss with psyllium Fiber was incorporated into apple juice 9.7 –63 –15 –7.2 10.4 NS –2.7 –20.7 NS –11.5 –23.9 –6.9 NS –6.1 –14.3 –4.4 g/MJ 1.7 NS 3.9 NS –12.9 –6 NS –10.6 NS –10.7 2.10.6 –27.7 –9 –5.4 g/MJ 7 6.1 NS 28.7 NS +36 NS TG (% change) LDL:HDL decrease by 4. Purified Fibers on Lipid Concentrations in Normal and Hyperlipidemic Subjects NC NC NC –25 NC NS –16.2 –14.4 Metamucil Plantago Seed husk Cereal 6.1 –18 –5.4_fm Page 406 Sunday.7 –13 –7.5 NS NC –1.7 NS 4.4 2387_ch5. male 125 type 2 diabetes 70 HLP 32 HLP 27 HLP 24 HC type 2 diabetes 50 healthy children (6–11 y) 42 HLP 20 HC 14 HC 7 healthy 58 HC males 9 HLP 27 HC 9 healthy 58 hyperlipidemia 10 healthy 110 hyperlipidemic 7 hyperlipidemia 2 healthy 33 hyperlipidemia 33 hyperlipidemia 10 healthy 27 hyperlipidemia Subjects –4.8% decline in LDL:HDL Comments 81 82 80 78 79 75 76 77 62 63 52 73 64 74 60 73 56 149 86 86 51 56 85 Ref.1 NS 2.9 –6.3 15.8 –16 –5.8 –12. 4 10 Guar bars Crispbread Pasta Gel Powder Granules Minolest (guar + psyllium) 8 wk 12 wk 11 d 21 d 24 mo 3 mo 16 16 Powder Gel 60 d 6 wk 15 Granulate 2 wk 8 wk 12 wk 14–19 3–12 mo 18 wk wk mo mo 2 8 4 2 15 13 15 16 In pasta Crispbread Granules Granules Guar type 2 diabetes HLP HLP.6 –26 –5.1 NC NC –13. corn bran 50 wk 24 wk 2 wk 4d Guarita 20 14 mo 26–40 11. but not sustained Apo B decreased by 20% Preparation with gel inhibitor 103 102 100 97 57 64 127 98 99 96 95 106 107 93 94 92 47 49 88 89 90 91 2387_ch5.9 NS –21 NS 17 NS NC NC NC –2.8 NS –6 NS –22 –10.7 NS –3 –23. 2001 3:06 PM FIBER IN THE TREATMENT OF HYPERLIPIDEMIA 407 . females HLP 58 HLP 16 type 2 diabetes 20 obese.2 –10 3 NS –16 –11.2 NS NC –1.5 NS –23 17.8 NS –6 NS –0.4 NC NC –9. May 8.8 –10.4_fm Page 407 Tuesday.9 NS 9 1. females 32 HLP 22 type 2 diabetes 8 diverticular disease 13 HLP 20 with carotid stenosis 83 healthy/HLP 23 HC 8 type 2 diabetes 6 healthy.20–30 15 15 40 15 15 16.6% Significant changes seen at 6 weeks.1 NS –1. males 17 HLP 10 11 32 12 –8.8 –14.9 –10 –11 –32 –6 –8 NS –13 –11 –14 –3 NS –7 –12.4 17.6 –9.2 NS –9 NS 2 –1. soy.2 –17 NC –9.8% FBG decreased 9.5 NC NC –5.9 NS 2.8 NS –40 –27 –4 NS 8 NS 13 NS –13 NS –3 NS 28 NS Weight gain same in control and test FBG + HbA1c decreased significantly Diabetes control improved Increase in ApoA1/ApoB by 11.5 20 Granules Cracker 8 wk 36 wk Guar.6 –10 –17 –3. male 10 obese women 12 HLP 10 type 2 diabetes 8 HLP 5 HLP 14 HC. pea.9 –16 NS –10 –18 –14 NS –18 –14 –16 –3.4 NS –26 NS 0 –13 NS NC NC NC 12 NS NC –2. pectin.1 NS 1. pectin. high-density lipoprotein cholesterol. 3RD EDITION . low-density lipoprotein cholesterol. guar Locust Bean Gum Fiber 408 Table 5.4_fm Page 408 Tuesday.3 –6 to 19 Total –10. LDL-C. triglycerides.4 –10 to 19 –10 to 19 Cholesterol (% change) LDL VLDL NC NC 0 to –17 HDL NC NC –10 TG (% change) Comments 99 58 50 Ref. psyllium. 10 healthy Subjects –6. NC.4 –8.4. total cholesterol. guar Locust. Note: Abbreviations: TG. May 8.5 –12. no change. HDL-C. Purified Fibers on Lipid Concentrations in Normal and Hyperlipidemic Subjects 2387_ch5. Locust. HC.4–12 wk 6 mo 15 8 wk Duration 15 18–30 Dose (g/day) 18 HC adults/children. hypercholesterolemia. hyperlipoproteinemia. total-C. psyllium. pectin.4 (Continued) Effects of Soluble. 2001 3:06 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. HLP. positive effect of guar..4.106. This included low-fat.57.4. However. water-soluble fiber which has been shown to lower blood lipids. On the other hand. The purified powder (Konjac mannan) from the tuber is a viscous.112 Wheat Bran Wheat fiber appears to protect from cardiovascular disease despite115.114 Improvements in the ratio of Apo B to Apo A were also observed.98.122 (Table 5.9%) in subjects with diabetes.4).7%). Results from long-term studies of guar have been conflicting. After 3 weeks of Konjac supplementation. in pharmacological terms. 2001 7:37 PM FIBER IN THE TREATMENT OF HYPERLIPIDEMIA 409 of this viscous fiber.104. almost all studies which have used other wheat bran preparations have failed to show significant reductions in blood lipid levels of hyperlipidemic individuals118.49. when the maximum effect has been achieved with clofibrate. or their analogs.49.95. since the bile acid .112 and glycemia112–114 when taken as a supplement.g. These results were accompanied by a 5% reduction in serum fructosamine levels. it is possible that any further effect of the fiber may be reduced. study design.5).94–103 possibly because of differences in dietary formulations of the guar supplement. a marker of glycemic control. with or without cholestyramine. Traditionally. since both substances are galactomannans.89.91 Further work has contributed support to previous evidence (Table 5.94–97. although there is one report of a significant rise in HDL-cholesterol. of the 13 studies cited. etc. The physiologically active component is a high-molecular-weight glucomannan polymer. the ratio of total to HDL cholesterol (10%). the falls in total and LDL-cholesterol were comparable. However.64. clofibrate. only hard red spring wheat bran has been convincingly shown to lower serum cholesterol levels in normal humans. Konjac mannan is made into a rubbery jelly and has been used as a food and a remedy. 9 reported a sustained. direct comparative studies must be undertaken before any statement about their relative efficacy can be made. low-cholesterol diets. Its advantage has been claimed to be its superior taste (or lack of taste) in comparison with guar.117–121 Of the different wheat brans.89 This indicates that. following a 3-week Konjac-supplemented diet compared to the control. cholestyramine).63.4). subjects with insulin resistance syndrome showed decreases in total and LDL-cholesterol by 12 and 22%. Konjac Mannan Konjac-mannan fiber is obtained from the tuber of the perennial Amorphophallus Konjac k and has been known in Japan for over 1000 years.120 The lack of consistent effect of wheat bran on blood lipids is of interest from the standpoint of mechanisms. It is not possible at present to say whether the mechanism or action of fiber overlaps with those of the established hypolipidemic drugs and whether specific combinations might bestow an advantage.116 its lack of consistent effects on serum lipids. it is likely that the mechanisms of fiber will complement those of the bile salt–binding (anion exchange) resins (e. In 2-week studies in which the maximum acceptable dose of guar given in crisp bread form was compared with the maximum acceptable dose of cholestyramine in the same patients. fiber was added to the patients’ preexisting diet and/or drug therapy.105 and guar104 compared with specific drugs.100–103 In the majority of the guar and pectin studies.119.4_fm Page 409 Sunday. May 6. 65.89.108–111 systolic blood pressure.107 Locust Bean Gum This viscous fiber has also been used successfully in a range of hyperlipidemic patients to lower serum cholesterol50 (Table 5.2387_ch5.114 Konjac also reduced serum fructosamine (5.94.57. In view of the relatively small bile acid losses seen with pectin85. respectively. the effects of viscous fiber on lipid metabolism might have significant clinical utility. which was then maintained constant. and systolic blood pressure (6.117 As with normal volunteers.4. the taste of guar depends on its purity and. 104 which consistently lowers serum cholesterol. 3RD EDITION losses in the stool following bran consumption have been shown to be comparable to those following pectin.132 These may result in a chronically reduced stimulus to hepatic triglyceride synthesis and hepatic lipid synthesis in general.4_fm Page 410 Sunday. notably soy protein.124 In addition. The other fractions remained low throughout the maintenance treatment period and for the four patients who were followed for 2 years.137 The reasons for the effects. also in healthy volunteers. Oat Bran Since the early studies of DeGroot et al.69. beneficial effect on serum lipids (Table 5. The effect does not appear to be related to associated fiber or saponins.4.5).2387_ch5.139 Legume Protein In addition to legume fibers (e. One investigator also recorded falls in serum triglyceride comparable to those seen with oat bran of equivalent soluble fiber content (20 g/day). especially triglycerides.126.135(Table 5. that 24-h urinary C-peptide outputs were reduced on high-legume diets. and unlikely to provide more than a small part of the explanation of the hypocholesterolemic action of oat bran.124 Despite early enthusiasm. Evidence for this hypothesis has been drawn together in the studies of Albrink et al.126 Diets which increased the total fiber intake by increasing not only the intake of legumes but also of whole grains. the increase in fecal acidic steroids was small. it has been proposed that the volatile fatty acids from oat bran and other viscous fibers.138 in healthy volunteers and is supported by the observation.5). with interest in their effect of improving glucose tolerance132 and other aspects of diabetic control. and nuts and seeds have also been shown to be effective in reducing cardiovascular risk factors.130 although not of young student volunteers. the effects on blood lipids appear to be sustained for 4 months to 2 years. the HDL-cholesterol level increased slowly to almost approximately the starting value by 24 weeks. then apparent despair.5).136.127–129 Dried Legumes Cooked. however. have been shown to reduce cholesterol levels of hyperlipidemic patients (total and LDL)140–146 (Table 5. 2001 7:37 PM 410 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.126 As with oat bran. Increases in fecal output on 115 g of beans are small and not significant in hyperlipidemic individuals. vegetables and fruits. some legume proteins.133 high-legume diets have been studied in types IIa and IV hyperlipidemic patients124.63. . may be related to the flatter postprandial glucose and insulin responses elicited by legumes. oat fiber contains an appreciable proportion of viscous fiber (beta glucan).4. increases in fecal acidic steroid losses were not noted.. and it is likely that this constituent may be one of its active hypolipidemic ingredients. Unlike wheat bran. May 6. the body of evidence supports that oat bran will have a significant. Although there were highly significant falls in all cholesterol fractions together with serum triglyceride during the initial 3 weeks of fiber treatment. Nevertheless. the falls in blood lipids. In this respect. All studies have shown falls in serum cholesterol irrespective of the class of hyperlipidemia studied.6).65. where recorded. Studies of Anderson and co-workers have demonstrated the lipid-lowering effect of oat bran given to hyperlipidemic patients (predominantly Types IIa and IIb) for 10 days to 2 years124–126 (Table 5.126 Again. although small. Further studies are required to explore this relationship.g. may produce metabolic changes which favor reduced cholesterol synthesis. dried legumes have been shown to lower serum cholesterol levels of middle-aged men. are not clear.134.. in proportion to the increase in fecal bulk. guar and locust bean gum).123 it was realized that oat constituents may have hypocholesterolemic effects. which arise from colonic fermentation of fiber and are subsequently absorbed.131 More recently.4. It has been suggested that perhaps the displacement of carbohydrate by wheat gluten on a high–wheat bran diet122 may be responsible for the positive relationship of wheat bran and cardiovascular disease.4. HC.1 5. hypercholesterolemia.5 2387_ch5. glucose.9 NC –6 NS –4 NC 0. LDL-C.4 3 –2 NS 0. healthy 12 HC. total-C. triglycerides. HLP.2 NS –17. male 24 HC males Subjects –4 –8 –7 –10 –12 –10.3 NS 23 3.8 –5.8 Total –2 –8. male 5 HLP 8 HLP. male 17 HLP 7 HLP 10 HLP 13 Healthy.8 NS NC –2 NS –3.5 35 Dose (g/day) Effects of Fiber-Rich Whole Foods and Supplements on Serum Lipids Fiberform Bread.5 –18 –24 –28 1. 6 Healthy 30 HLP. Beans/oat bran Baked beans Beans Mixed Cookies Instant oats Oatmeal Bread.6 56.8 30 NS HDL –6 –25 NS –25 –3 NS 7.5 –2 NS 0.7 0 5 NS –24 3.3 NS NC 13–32 NS –9 NS 8.5 NS –10. muffins Powder Oat bran Flakes 50 50 10. muffins Wheat bran Fiber Table 5. BP Comments 126 135 135 124 68 70 126 66 63 127 124 150 128 65 67 129 134 125 69 118 119 120 63 Ref.4 –23 –6.24 wk wk mo wk wk wk wk 28 d 4 wk 21 d 3 wk 5–11 mo 8 wk 8 wk 8 wk 6 wk 3–6 mo 17 95 77 100 52 2. May 6.1 TG (% change) 4 subjects were followed for 99 weeks Apo B decreased by 9.4_fm Page 411 Sunday.9 –23 –25 –26 –9.1 –14 –8.8% NC insulin. high-density lipoprotein cholesterol. total cholesterol. male 24 HLP 10 HLP. no change.7 NS 0.2 –2 NS LDL 1.3 –3 NS –13 –17 –9 –5 NS –4. male 12 HC.3 –30 NS 20 NS VLDL Cholesterol (% change) 17 NS –15 NS –12 –14. male 19 Healthy 24 HC. males 42 HC 113 Healthy 13 HLP 136 HLP 106 8 HLP. Note: Abbreviations: TG.3 –19 –22 –13 –5.4 –37 3.7 56 84 50 (100 g dried) 110 dried 140 dried 115 450 120–162 100 beans or oats 50 oats or 134 beans 2 4 3 2 3 3 3 mo 2 wk mo mo wk wk 100 25 3 2 8 4 Duration 10 HLP. male 13 moderate HLP 10 HLP 31 HLP. NC.8 –21 –8 NS –3.4.8 –6. 2001 7:37 PM FIBER IN THE TREATMENT OF HYPERLIPIDEMIA 411 .4 –23 –5 –24 –9 NS –6. hyperlipoproteinemia. low-density lipoprotein cholesterol.9 –3. HDL-C.9 –20 2 NS 2. guar. have been shown to reduce serum total and LDL-cholesterol levels by 10 to 20% and with a lesser fall in HDL-cholesterol levels. demonstrated a 13% reduction in LDL cholesterol for a soy protein intake of 43g/d. oat bran.136 However. crisp bread.4_fm Page 412 Sunday.71. since fava protein isolate failed to achieve the desired effect. the greatest lowering of cholesterol and triglyceride levels of all the studies to date was seen. in addition to the background diet.” i. CONCLUSION Viscous fibers such as pectin. the levels of fiber in the oat bran– and bean–containing diets have also been of this order of magnitude.6). since soluble fiber composes approximately 20% of the dry weight of both beans and oat bran.e. the capacity to lower blood lipids may not be a universal property of all legume proteins. . The effects of soy protein in lowering serum cholesterol are well recognized. reducing or destroying the viscosity will eliminate its beneficial effects. However. The mechanisms of action of fiber are likely to be complex and possibly include increased bile salt loss.2387_ch5. and enhanced colonic synthesis and uptake of SCFAs. the effectiveness of the supplement in hyperlipidemia may be determined by its formulation. when Gatti and colleagues enriched spaghetti with guar. With respect to the whole bean.145 The mechanisms for the hypolipidemic action of soy protein remain obscure but may relate to the amino acid profile148 or the presence of specific pharmacologically active peptides liberated during digestion. and high-fiber foods such as oat bran and dried beans. the total-to-HDL cholesterol ratio was reduced by 6.47 Spaghetti is already recognized as a slowly digested carbohydrate form which causes an unexpectedly low rise in blood glucose. significant falls in serum triglyceride levels have also been observed. or beans. appear to lie in finding the most effective food vehicles in which to incorporate fiber.150 This effect is likely to have been greatly enhanced by the addition of guar. locust bean gum. The addition of guar would likely not only have enhanced the reduction of glycemic and insulinemic responses to the pasta but may. The viscosity of viscous fibers is responsible for the slowing of absorption in the small intestine. Interestingly. presumably because of the low viscosity of the apple juice mixture. Data continue to accumulate supporting the lipid-lowering effect of legume protein (Table 5. The choice by Gatti and co-workers of spaghetti as the vehicle for delivery of the fiber may have been in large measure the reason for the success of their trial. since it encompassed many of the mechanisms responsible for reducing blood lipids. A large meta-analysis of studies up to 1997 by Anderson et al.3%. altered site and rate of absorption. therefore.4. in effect. therefore.149 On the other hand.72 Effective Fiber Dosage and Formulation In general. resulting in the creation of a very effective sustained release carbohydrate source. the effective dose of the viscous fibers required to lower serum cholesterol levels has been of the order of 12 to 30 g/d. have resulted in a proportion of the pasta starch being converted to “fiber. May 6. Konjac mannan. other factors will have to be uncovered to explain the general hypolipidemic effect of legumes in hyperlipidemic individuals.. all providing 12 to 30 g fiber daily. although specific fiber and protein effects may be relevant to the action of some beans.147 In a study with hyperlipidemic subjects where both the soy protein and the soluble fiber intake were increased. 3RD EDITION since it is also found after administration of soy isolate. Incorporation of a small quantity of pectin into apple juice failed to reduce cholesterol levels. reduced hepatic lipogenesis secondary to reduced postprandial glucose and insulin responses. 2001 7:37 PM 412 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. One of the important directions for future development in this field would. carbohydrate which is unavailable for small intestinal absorption but which acts as an additional source for synthesis of SCFAs in the colon. When the fiber was provided in a starchy food such as pasta. 6 2387_ch5.8 –26 –9 –0. HLP. LDL-C. hyperlipoprotemia. triglycerides. NC.2 NS 13 0 –12 NS NC HDL –69 NS 11 NS 3 NS –11 –17 NC TG (% change) Changes sustained over 18 months 500 mg cholesterol added Comments 71 144 143 144 141 153 140 140 Ref. HDL-C. no change.8 –21 –10 –3 NS –20 –26 –19 –23 Total –25. 1 healthy 21 HC 127 HLP 27 HLP 20 HLP 8 HLP Subjects –20.60–100 70–80 “Cholsoy” Mixed food Mixed meals “Cholsoy” 4 wk 4 wk 2 wk 4 wk 8 wk 4 wk (18 mo) 3 wk 6 wk Duration 19 HLP 6 HLP 21 HLP.4. total-C. 2001 7:37 PM FIBER IN THE TREATMENT OF HYPERLIPIDEMIA 413 . May 6. total cholesterol. low-density lipoprotein cholesterol. Note: Abbreviations: TG. 60–100 60–100 Granules “Cholsoy” Dose (g/day) Effects of Legume Protein (± Saponins) on Serum Lipid Concentrations of Hyperlipidemic Subjects Textured soybean protein Granules Granules Fiber Table 5.4_fm Page 413 Sunday.6 NS –17 3 NS 15 NS Cholesterol (% change) LDL VLDL –7. high-density lipoprotein cholesterol. hypercholesterolemia. HC.8 NS –33 –18 32. Exp.. Cholesterol-lowering effects of psyllium intake adjunctive to diet therapy in men and women with hypercholesterolemia: meta-analysis of 8 controlled trials. Hypercholesterolemic effects of soluble fibers. J.. J. S. J. 175. 13. Story. Tombes... Dietary fibers. Krauss. A.. and Hengehold. Anim. Lawrence. D. M. R. J. Gassull. V. REFERENCES 1. and Truswell. et al. D. Kritchevsky. Interim final rule for food labeling: health claims. phospholipids and cholesterol from mixed micelles by bile sequestrants and dietary fibers. L. W. Biol.. Salomons.. F. J. Leeds. Sci... J. D. Hypocholesterolemic effects of oat-bran or bean intake for hypercholesterolemic men.. 3RD EDITION Further developments to enhance the clinical utility of this approach should include not only a search for effective fiber types but also the appropriate vehicles in which to deliver them. 57699-57733.. Gastroenterol.. M. 1986. A. M. W. J. 6. S. Clin. M. Lin. 1012.. Morris. J.. J. W. D. J. Truswell. B. P. A. D. S. 9. D. V.. D. S. M. Rippe. Houston. L. 1974.. L. M. Am. Goff. 104. B. Goff... S. Rosner. 5. H. . M. et al. 1977. in Dietary Fiber: Basic and Clinical Aspects. M. W. Clin. Willett. Nutr. E. J.. T. M. D. J. V. Proc. Becker. The Federal Register 64 (October 26). 1973–80. and Story... J. 15. 3015–3023. Chen. H. C. 458. T. S. Nutr. B. 2000. M. Nutr. C. Gassull. 2000.. W. D. Wolever.... Jenkins. W. 30–42... L. Anderson. L. B. Influence of propionic acid on the cholesterol metabolism of pigs fed hypercholesterolemic diets. Soc.. J.. L. Cassidy. 2. Jenkins. J. S. Fulgoni. M. A. M. Invest.. X. 15. and Sacks. D. 71. J. Anderson. 127... Nutr. K. W. Thacker.. Anderson. C.. Clin. A. Clin. 30. JAMA. Allgood. Anderson. Sieling.. A. Dietary fibres. Jerdack. 175. J. L. 969. The interaction of dietary fibers and cholesterol upon the plasma lipids and lipoproteins.. Binding of bile salts in vitro by nonnutritive fiber.2387_ch5. Metz. J. Conner. J. LaRosa. B. Br. 1988. In vitro binding of bile acids and bile salts.. Kay. Kritchevsky. 69... 1146. Can. Sci. J. V. L. S. M. Wolever.. Dilawari. G. and Story. Hunninghake. in hypercholesterolemic adults: results of a meta-analysis. Summary of the Second Report of the National Cholesterol Education Program (NCEP) Expert Panel on Detection.. 171. 51. P. 28. 102. 1997. 1977... Chen.. Med. F... Serum short chain fatty acids after rectal infusion of acetate and propionate in man. Effect of citrus pectin on blood lipids and fecal steroid excretion in man. New York. Food and Drug Administration. 1993. Jenkins.. Nutr. Brown. Fry.. 1999.. 4... 19. 18. Wolever. W. Clin. Summerbell. 40(b). L. 17. A. and Jennings.. L. and Gallo. J. and Jenkins. 305. D. M. R. Vahouny. 3. D. W. and Vahouny. J. G.. Leeds. M. 65 (September 8). J.. J.. Am. G. Haisman. 2284-2299. Olson. Altringer. 1981. O. 12. 1978. Med. 16.. 60. and Anderson. Plenum Press. 2000. Anderson... A. 8. M.. 275. D. Am. 3. 1999. Final rule for food labeling: health claims: soy protein and coronary heart disease. 10. Binding of bile salts. 1372. Raymond. J.. W.. 1976. Aherne. Nutr. Mol. Roberts. G. L. D. D. F.. Lipids.. J.. M.. A. Stoy. P.. Revision 200: A statement for healthcare professionals from the nutrition committee of the American Heart Association. Chen.. A. R. AHA dietary guidelines. Am. R. Warner. 1984. and Story. J. Clin. 54685-54739... Kritchevsky. Nutr. The cholesterol lowering properties of guar and pectin. 2001 7:37 PM 414 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. S. 1984. W. but not HDL cholesterol. Petro. and Alberti..... 1980.. H.. and Connor... Med. Brighenti. Circulation. Evaluation and Treatment of High Blood Cholesterol in Adults. L. T.4_fm Page 414 Sunday. 215.. D. Kritchevsky.. (Adult Treatment Panel II). M. 11. V. Clin. J. Cholesterol-lowering effects of dietary fiber: a meta analysis. A. 42. Federal Register. Nutr. 1429. 61.. Food and Drug Administration. M. D. A.. R. sterol balance and bowel function in human subjects. and Hill.. plant sterol/stanol esters and coronary heart disease. A. T. Eds. D.. D. 14. Psyllium enriched cereals lower blood total cholesterol and LDL cholesterol. 8. L. J. Am. M.. Propionate may mediate the hypocholesterolemic effects of certain soluble plant fibers in cholesterol-fed rats (41791). 7. G. May 6. 1975.. R. V. fibre analogues and glucose tolerance: importance of viscosity. 269. A. Clin.. 1. 472–9. . J. 1990. 549. R. 1392. Clin. Slattery. 40.. A.. Relationship of glucose tolerance and plasma insulin to incidence of coronary heart disease: results from two population studies in Finland. 66... J.. Nepokroeff. Wolever. et al. A. and Rosselin. 1976. G.. 1990.. Am. L.. J. 35.. Ann. Nair. A. Dugan. G.. J. J. in Dietary Fiber: Basic and Clinical Aspects.. M. G. Specific types of colonic fermentation may raise low-density-lipoprotein-cholesterol concentrations.. J. and Furumoto. L. Dietary treatment of familial hypercholesterolemia — differential effects of soy protein according to the Apolipoprotein E phenotypes. J. Lakeshmanan. J. Plenum Press. 34. Metabolism. F. Jenkins. R. Eschwege. Nutr. T.. J. Res. Wolever. Dilawari. A. 564. Ocana.. J. Br. Nutr.. 253. M. A. S. F. L. Schaefer. C. R. 1595. V. Relationship of plasma insulin levels to the incidence of myocardial infarction and coronary heart disease mortality in a middle-aged population... W. Veldman... Cohen. E.. JAMA. 253–64.. 282. K. D. Ravalia. Gatti. Fassi. J. Corey. G. Leeds.. M. Nutr. Clin. V. Dietary fibers.. P. M. Ransom... G. B. M. J.. and Lefevre. C. 2001 7:37 PM FIBER IN THE TREATMENT OF HYPERLIPIDEMIA 415 20. Lipid Res.. S.. Todesco. Rao. Metabolic effects of reducing rate of glucose ingestion by single bolus versus continuous sipping. Rao. 50. Ducimetiere. 36. Diabetes. Goff. K. Vuksan. 141.. U. G. and Manninen. M. K. J. 80.. Diabetes Care.4_fm Page 415 Sunday. W. and Wearne. Story... 2. 1.. J. Ehnholm. Am. G... 32.. M. M. Biophys. Modification of steroid excretion in response to dietary fiber. E. P.. A. K. T. Welborn. 54.. W. Possible mechanisms through which dietary pectin influences fibrin network architecture in hypercholesterolemic subjects. Eds. Dietary fiber and intestinal adaptation. 24. 41. Plenum Press.. J. 1988. J. J. Clin. Kasper. P. E. R. P. S. Richard. 1991. Gmull. M. Ludwig... S. Story. C. 1979. New York.. A. Hutlunen. S. H.. T. Connelly. H.... M. Gregg. Nutr. 54. F. Jenkins.. H. Diabetologia. Composition of high density lipoproteins in rats. J. 1991. Am.. 86... C. H.. C. Int. and Sirtori. 775.. R... Vahouny. 19. H.. V.. 30.. Vermaak.. weight gain. Jr. S.. J. 578.. D. O. 27. T. R. A.. Ness. and Alberti. Kesaniemi. O. Vorster. Papoz... Commun. A. P. Pereira. J. Kroenke. 1980. Stimulation by insulin of rat liver Beta-hydroxy-Beta-methylglutaryl coenzyme A reductase and cholesterolsynthesizing activities. Vuksan. 27.. B. C. Biochem. 40. V. T... E. Bosello.. Clin. Vorster. Rabast... C. W. M. Banting Lecture 1988. 23. 1191. J. Schneeman. 21.. Thromb. E. C. Vezina. 1986. Diabetes Care.. Am. 38. and Kritchevsky.. 39. 1977. M. H. A.. G. M... 31. 33... .. and Venter. A... Intestinal cholesterol absorption efficiency in man is related to Apoprotein E phenotype. S. and Kritchevsky.. and Fehle. V. Nutr. D. M. Clin.. Dietary fiber. Apolipoprotein E metabolism in normolipoproteinemic human subject. G. and Jacobs. 1991. K.. 154. 1997. 1539–46. A. C. T... and Cummings.. H. J.. D.. A. Coronary heart disease incidence and cardiovascular mortality in Busselton with reference to glucose and insulin concentrations. V. 28. 1999. 1999..2387_ch5. D. 1978.. A. A. 93. W. L. Effects of dietary propionate on carbohydrate and lipid metabolism in healthy volunteers. Manttani. J. and Alberti. Jenkins. Jerling.. 181.. 1979. A. R. Wolever. Venter. Role of insulin resistance in human disease. H. A. L. 1167. Osthuizen. 1979. Jenkins. P. C. and Brewer. Propionate lowers blood glucose and alters lipid metabolism in healthy subjects. Zech. Cimmarusti. M. and Miettinen.. 584–90.. Descovich. 53. 205. B. Am. A. M. Clin. V. Leeds. R. C.. J... R. and Cassidy. 1987. Apolipoprotein E polymorphism influences the serum cholesterol response to dietary intervention. Matteucci. Decrease in postprandial insulin and glucose concentrations by guar and pectin. Gassull.. M. Claude. Pyorala. S. 1847.. and Jenkins. D. S. 45. M. T. Brighenti. Med.. May 6. G. A. and cardiovascular disease risk factors in young adults. Clanocchi... D. 20. 217. Koskinen. 2. A. fiber analogues and glucose tolerance: importance of viscosity. L. A... G. M. Wolever.. 22. 29. Cochet. Downes. D. R. 37. Vahouny. The effect of dietary fiber on the postprandial serum vitamin A concentration in man. 32. Y. Gastroenterol. J. A.. A.. Reaven. M. Nutr. J. and Porter. Hegele. 1984. Invest.. 1973. 25. Connelly. 1986. Cunnane. and Patten. Ehnholm.. D.. New York. Ocana. 1991. C. Med. H.. Res. Buckley. Diabetes. L. A. Am. J. 25. 39. T. Rimondi. 131. V. 26. 37. A. B. H. M. J. G. J.. Vahouny. V.. Van Horn. L. Eds.. 85(5).. Long-term effect of soluble-fiber foods on postprandial fat metabolism in dyslipidemic subjects with apo E3 and apo E4 genotypes.. 704.. Jenkins. D. in Dietary Fiber: Basic and Clinical Aspects. Metz. 50. Med. 48... L. A. A. M. P. Keinanen-Kiukaanniemi. healthy population. New Engl. 427. 1988.. Nutr.. Nutr. 661. B. Cholesterol absorption: regulation of cholesterol synthesis and elimination and within-population variations of serum cholesterol levels.. P. and Nestel. Hughes. 61.. I. S. 148. Nakamura. J.. L. S. and Schneider. 42. Schwandt. 1989.. Liu. 1982.. Cholesterol lowering effects of psyllium hydrophilic mucilloid for hypercholesterolemic men. Nutr. M. R. plantago ovata and high fiber treatments.2387_ch5. Simons. Torri.. Clin. Kern. J. Arch. Kuba. K. Hillman... Clin.. E. Kestin. J. L. Gayst. Miettinen. W. W.. M. L. Balm... R. Hanson.. 64. 43. Zettwoch. K. 1. Fisher. O.. Niemi. Jensen. S. 44.. Reynolds. and Moller. 1975.. I. 1989.. P. Gastroenterol. JAMA.. and Neureuther. 57. N... S. 15. 1991. Huth. E. C. Am.4_fm Page 416 Sunday. 165.. 1989... 1983. The hypocholesterolemic effect of locust bean food products in hypercholesterolemic adults and children. A.. and Floch. Cholesterol intake and serum cholesterol level. Van Horn. 67. Lancet. Comparison with cholestyramine treatment in Type IIa hyperlipoproteinemia. K. B.. F. 52. 148. and Stamler. L.. and Fettel... 2. J.. Apo E modulates the metabolism of apo B containing lipoproteins by multiple mechanisms. and Tarpila. 1984. Lignin: a cholesterol-lowering agent?... Vargo. J.. 47.. Comparative effects of three cereal brans on plasma lipids. JAMA... Clifton. 456. Traitement de l’hyperlipoproteinemie essentielle de type II par un nouvel agent pierapeutique.. 46. Role of water-soluble dietary fiber in the management of elevated plasma cholesterol in healthy subjects. 45. Moss. Colonic bacterial flora and serum cholesterol: alterations induced by dietary citrus pectin. and Bishop.. Tietyen-Clark. C.. G. K... Moag-Stahlberg. L.. Elmer. Clin. J. Assoc. E. 896.. P. D. Anderson. J. D. 2.. C. Lindner. blood pressure and glucose metabolism in mildly hypercholesterolemic men. Am. Belanger. Med.. Cabelli. Abraham. Tada. H. 324. Med. M.. Am. Zavoral. Kagami. 80(5). Kalparik. Three-week psyllium-husk supplementation: effect on plasma cholesterol concentrations fecal steroid excretion. V. 1985. Gregg. G. Long-term effects of guar gum and microcrystalline cellulose on glycemic control and serum lipids in type II diabetes. Richter. 103. Am.. and Mehfa. . and Ruys. J. 1989. Bell. K. K. A. Springer Verlag. Public Health. G. 207. M. W... M. Atherosclerosis. and Pomare. R. S.. Berlin.. J. J. D. Am. Kondo. 34. Balasubramaniam.. Atherosclerosis. and carbohydrate absorption in men.. 433–9. 1991. la celluline. 52.. 38. A. 44.. A. E.. 285. 1985. E. Gates. Clin. J. M. Feldman.. 60. P. C. J. Long-term treatment of hypercholesterolaemia with a new palatable formulation of guar gum. T. Hectorne. D. C. 240. Miyazima. II. 1982. G. Nutr. J. 361. Can. 51. Effects of guarenriched pasta in the treatment of diabetes and hyperlipidemia. 81(2).. Am. Med. 223. H. P.. 66. 58. Clin. 101. R. L. The effects of the fiber components pectin. Cholesterol-lowering effects of psyllium hydrophilic mucilloid. 1970.. 299. 261(23). Am. K. 28.. Clin.. Cholestyramine plus pectin in treatment of patients with familial hypercholesterolemia.. P. 55. Int.. Lancet. West J. 49. Ballew. Oat bran as a cholesterol-reducing dietary adjunct in a young. 45. K. Fields. M. J. and Takeyama. 1259.. A. Briones. W. 253. and Nelson. and Davidson. Thiffault. Pharmacol.. Z... 2001 7:37 PM 416 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 1988.. B. T. Breslow. 59. Bran and blood lipids. Nutr. E. J. A. N.. A. Clinica Chim.. M. D.. W. P. Rep. H. Am. Eur. Peters. and Salmela. New Engl. Int. A. 53. H... R. J... P. Frantz. J. H. J. Ann. 1989. Weisweiler. Gotto.. Serum lipids and cholesterol metabolism during guar gum. J. Palumbo. and Pouliot. D.. P. in Cholesterol Transport Systems and Their Relation to Atherosclerosis... 183. C. 1973.. Effect of several kinds of dietary fibers on serum and lipoprotein lipids... T. A. E. M. 63. J. T. A. Genetic basis of lipoprotein disorders. Steimetz. 629. 292. 84. Eds. Oeltgen. 1990. Spiller. M. Ishikawa. and Huninghake. Gold. R. R.. N. 56. Gatti. High fiber diet in hyperlipidemia. D. 49. and Jacobs. and Gates. Normal plasma cholesterol in an 88-year old man who eats 25 eggs a day. 1988. Cardiol.. cellulose and lignin on serum cholesterol levels. and Brewer. T. B. Acta. Nutr. May 6. 324.. Miettinen. 1982. 3RD EDITION 42. Y. Ruth. 54. 3419. 912.. Ellis. E. T. Effects on serum lipids of adding instant oats to usual American diets. K. C. R. Nutr. and Kesaniemi.. J.. Invest. 65. 1978. Mechanisms of adaptation. P. Hannan. 1988. Denegri. C... H.. 379. Camisasca.. 1991.. 215... Catenazzo. Med. S. K. J. J. Zech.. 26(2). 373.. 1992.. J. and Sirtori. Clin. Haskell. 183. A. 62. Doyle... 47. J... Metab. Mueller.. Soluble fiber enhances the hypercholesterolemic effect of step 1 diet in childhood.. Davidson... Reynolds. 71.. 30.. Wolever.. M. J.. 269–73. King. J. H. Davidson. Oeltgen. D. Tietyen. and de Gery. Nutr. 14.2387_ch5. Clin. R. Acta. S. J. S.. Rimondi.. P. 100.. C. T.... Am.. Conelly. M.. P.. Ginsberg. Jenkins.. 81.. M. 77. Bircher. P. J. 75. D. C. J. Blonde. Setchell. D. D.. P. K.. Ravaglia. Fulgoni. K. Singh. R. F. Nutr.. Gagne. B. Effect of pectin dose on serum-cholesterol levels. Boctor. and Ghatak. W.. T. Cunnane. A. 1988.. D. D. J. Prev.. 463.. J. Nutr. Vidgen. 78. H. K. Patten. 83. Clin. J. 83(10). S.. 1433–8. 72. H. Am. M.... A. Mehling. Relle. Williams. May 6. L. Ciarrocchi. M. T... Spark.. Patten. 79. W. D. Nutr. Am... Med. 1995. and Balm. C. T. Ransom. 1977. and Lupien. Baliew.. Van Horn. S... D. Am. 1990. M. Chao. N.. 237–41.. Rondier. M. L. Effect of pectin on serum cholesterol. Clin.. H.. J. E. Walker. 1191.. Serum lipid response of hypercholesterolemic men to single and divided doses of canned beans.. V.. L. South Med. 2000. J. N. D. C. J.-J.. Neal. Guerrero-Romero. Cholesterol lowering effects of soluble fiber cereals as part of a prudent diet for patients with mild to moderate hypercholesterolemia. J. 1994. Long-term cholesterol-lowering effects of psyllium as an adjunct to diet therapy in the treatment of hypercholesterolemia. and Lazcano-Burciaga. 54. 53. Method of administration influences the serum cholesterol-lowering effect of psyllium. J. 1994.. A. Nutr... H. 61.. A. Gustafson. Anderson. J.. A. Anderson.. A. S. 1013. J. K. S. H. J.. G... and Puder. 85. S. Nutr. B. Allgood. C. L. Howard.. J. D. Steinke. Clin. Brun. 65. Fulgoni.. P. Wolever. G. G. Boctor. K. Tredger... Effects of a soy-protein beverage on plasma lipoproteins in children with familial hypercholesterolernia. H. Nutr.. L... M. Chao. Sci. 466–73. D. Clin.. N. J. Patten. Rodriguez-Moran. G. Anderson. 70. C. C.. J.. D. K.. M. and Sirtori. The effect of daily baked bean (Phaseolus vulgaris) consumption on the plasma lipid levels of young. L. P. Lipid-lowering efficacy of psyllium hydrophilic mucilloid in non insulin dependent diabetes mellitus with hyperlipidaemia. 73. Drennan. W.. M. and Crey. Res. J.. Br. A. S.. 1994.. 67. Spencer. V. K. J. Liu. J. C. Jenkins... 495. H. 2001 7:37 PM FIBER IN THE TREATMENT OF HYPERLIPIDEMIA 417 67. Teitel.. M. 437. G. Mueller. 1977. 1990 74. V. Effects of psyllium on glucose and serum lipid responses in men with type 2 diabetes and hypercholesterolemia. and Fulgoni. Clin. Laurin... W. Allgood. J. S.. Synergistic effects of psyllium in the dietary treatment of hypercholesterolemia. 251–7... Am. V.. 12. Anderson. 59... Moorjani. Nutr. Coll. W. 1990. J. S. Kendall.. 1991.. and Tarpila. 1990. Chim. 257. C. D. A. S. J. fecal bile acids and biliary lipids in normolipidemic and hyperlipidemic individuals.. J... C.. 70.. J. Diabetes Complications. 71. 18.. L. P. Jenkins. Clin. 1991. L. 1999..4_fm Page 417 Sunday. Agrawal. 1055–9. and Olson. Hamilton. T. and Bryant. W. R. A. L.. K. G. Matteucci. Ranson. and Daggy. 471. Nutr. 68. Clin. Clin. 1524–33. A. Spencer. L.. Indian J. 84. Lipid. Am. Tietyen. L. C. W... normo-cholesterolemic men. H. Clin. C. Smith. 1998. K. Serum lipid response to a fat-modified oatmeal-enhanced diet.. Bell.. Saldanha. Liao. J. Bryant.. Descovich. S. J. 1966. Nutr. Jacques. A. Am. T. D. Dugan. P. Dietary treatment for familial hypercholesterolemia — differential effects of dietary soy protein according to the apolipoprotein E phenotypes. 367–76. V. 98.. Psyllium reduces blood lipids in men and women with hyperlipidemia. Gaddi. A. 272–8. 1988. Anderson..and glucose-lowering efficacy of Plantago psyllium in type II diabetes. Shutler. 51. Oat-bran cereal lowers serum total and LDL cholesterol in hypercholesterolemic men... Am. Delbarre... Garvey.. and Low. 307.. 86.. L.. A. 79... E.. Hall. B... J. 52.. B. 76. A.. Palmer. P.. Maki. 82. H.. J. G. F.... Am. Am. B.. J. C. S. J. McMillan. Clin. Ransom.. Am. R. Bollella. and Hunninghake.. and Weingand.. Am.. Effect of psyllium in hypercholesterolemia at two monounsaturated fatty acid intakes. J. 1020. Hectorn. C. G. Med. Lack of effect of two pectins in ideopathic or gout-associated hyperdyslipidemia hypercholesterolemia. Torri.. S. 377. W. A. C. L. V.. Kong. 17. 1131. P.. 52. M.. Turner. C. Am. W. B. F. G. Clin.. T. R. Nutr. C. J. and Dixon. W. and Fulgoni. Mueller. . G. Nutr... B. Clin. M. and Oeltgen.. R. Emidy. D. Am. J. and Stamler. Med.. Miettinen. F. McMillan. 1997. Wolever. Brown.. Lau. 80. Nutr. F. D. O’Conell.. T.. R. E. Gupta.. Long-term effects of consuming foods containing psyllium seed husk on serum lipids in subjects with hypercholesterolemia. Morgan. 1998.. G. 69. K. Nutr. F. D. and Edelman.. H. 72. Rozzani.. J. 39–43. T. Leo. 3RD EDITION 87. V. J... Slavin.. S. and Vigorelli. and Uusitupa. Dujovne. Chuang. Cravero. Voutilainen.. A. 1985.. and Saris. Jenkins. G. 51. 16. 23.. 1997. Treatment of familial hypercholesterolemia with a combination of bezafibrate and guar. Kwiterovich. Ranzi. G. W.. H. J. Haskell W. Application of soluble dietary fibres in treatment of hyperlipoproteinemias. A. R. 15–9. diabetes and hyperlipidemia. 105.. M. and Lin. 1979. Bianchi. M. Leeds. 107. T. 1988. Graham.. 2001 7:37 PM 418 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.. 98. G. Koistinen. C.. D.. Long-term effects of water-soluble dietary fiber in the management of hypercholesterolemia in healthy men and women. 575. Nutr. Effects of solid and liquid guar gum on plasma cholesterol and triglyceride concentrations in moderate hypercholesterolemia. J. 1997. H. P. M. Miccoli. P.. Clin. M. 1980. O. A. Cassader.. J. A. 1972.. E.. L.. 6. M. 99. S.. M.. Metabot. Tuomilehto. I. 585–89. Reynolds.. Insull. Jenkins. and Groop L. 1999. 208. H. A..... Am... 33.. Van Duyn. 1992.. Muls.. Yoshida. R.. J. 45. 231–36. Bozzo.-G. 104. C.. Bruno. C. Davidson.. and Korhonen.. Melvor.. Relat. 32. 34–7.. Margolis.4_fm Page 418 Sunday. I. Jenkins.. D. Penagial. Long term treatment of severe hypercholesterolemia with guar gum.. 39.. A..... J. 17. O.. Superko. Acta Med. Karttunen. J.. 79. A. J.. 513–8.. Arvill. Cavallo-Perin. U. 101.. Gref. K. Therapeutic effect of guar gum in patients with non-insulin-dependant diabetes mellitus. 89. Rep. P. E. Effect of guar gum on body weight and serum lipids in hypercholesterolemic females. 61. BMJ. and Homan. R. Assoc... D. 39.. 911. Long-term effects of guar gum in subjects with noninsulin-dependent diabetes mellitus. Pisu.. P. K.. R. R. and Shimahara. M. 1988. E. Nutr. R.. 58. Groop P. 1982. Am. 1984. T. 23. 81(2). Harju. L... 1995. Tagliaferro. H. 94. M. Disord. 548–55. R. 310. 91. J. Slavin. 40(b).. Marena. Clin. Clin. Cerri.. 1984. (Plovdiv). 91. L. J.. A.. Krachanova. W. K. Uusitupa. J. Med. and Whittam. W. Middlehoff. J. 291. A. J.. L. 1986. A. G. Jokela. B. Behall. 380. Kay. Am.. F. M. S... Aro. H. Clin. H. S. Zocca. C.. Atherosclerosis. Effect of short-term ingestion of Konjac glucomannan on serum cholesterol in healthy men. 21. Huang. T. Cardiol. Castagnone.. J. and Krachanov. H. The effectiveness of long-term fibre supplementation on weight maintenance in weight-reduced women. Scand.J. A. B. Clin. 96. Nutr. E. Cummings. 1165. and Bodin L. Nutr.. M.. Tognarelli. Guar-pasta: a new diet for obese subjects. Hypercholesterolemic activity and molecular weight of Konjac-mannan. 60... Atherosclerosis. 1287. . van Ree. Cominacini. 109. S... C. J. J. D. W. Y. and Navalesi. J. Acta Diabetol. Dietary fiber and blood lipids: treatment of hypercholesterolemia with guar crispbread. 1993.. L.. Mann. E. D. Salenius. J. 18–23. Riekkinen. Clin. A. R. 157. Michnowski. Dietary fibre. Zavoral. M. Huttunen. M.. A. A.. Diab. 123.. Westerterp-Plantenga. and Farquhar. P. L. Intl. J. Moderate guar-gum addition to usual diet improves peripheral sensitivity to insulin and lipemic profile in NIDDM.. Am.. A. H. S... E. E. E. S. 45. Jou. V. Jenkins.. 62. Metab. J. 1982. 108.. Obes. Effects of guar gum on plasma lipoproteins and apolypoproteins C-II and C-III in patients affected by familial combined hyperlypoproteinemia. Knopp. 95. Tuomilehto. Gruveda-Popova.. C... 1995. Gastroenterol. 11. O.. F.. 88... 97. Voutilainen. 77.. Suhiama.. A. Velio.. Nutr. intestinal transit and fecal output in man. 1986. Sawrey-Kubicek. J.... M. B.. 90. W. Wu. J. A.. .. P.. and Silvasti.2387_ch5.. J. Int... Med. 1986. M. Am. R. Wirth. Superko. Ferrari.. 106. J. 1997. 1980.. R.. A. O’Connor. 102.. and Pagano. Effects of guar gum in male subjects with hypercholesterolemia. 100.. 103. H. Aro A. A.. Long term effects of guar gum on lipid metabolism after carotid endarterectomy.. 92. Pasman W... J.. Kiriyama S.. Garbin. and Jepson..... C. E. Tai. Nutr. A. J. Yang. Long-term effects of guar gum on blood lipids.. Am.. Dietary fiber and blood lipids: reduction of serum cholesterol in type II hyperlipidemia by guar gum. Am. D. The effect of dietary guar on serum cholesterol. Atherosclerosis. May 6. and Davoli. M. and Mendeloff. Jensen C.. and Schlierf. ii. Stenman S.. Haskell. N. H. Vinni.. Nutr. Am. Bosello. Am. Formos. Enishi. Long-term blood cholesterol-lowering effects of a dietary fiber supplement. Folia Med. Djurdjev.. 93. Am. Vansant. Dietary fiber. H. R. Lipid Res.. Leeds. Lancet. H. J. T. A. C. Ehnholm. Prev. 221. 7.. Brauning.. M.. Aro. 95–96. Cardiol.. M. and Jepson. A.... Giampietro. A.. 1979. R. Silvasti. Pyorala. J. P. 41F. 2. Chen..4_fm Page 419 Sunday. J. F... M. Reck. J. P. Kirby. 115. F. 126. and Uusitupa.. J. W.. Jenkins. B. W. 574. The effects of dietary fiber component Konjac-glucomannan on serum cholesterol levels of hypocholesterolemic subject. V. Am... Owen. J. 753. J. D. Parker. 1999. Connelly.. Third. N... 1979. 1987. J. Jenkins. Am. A.. and Fulgoni. 40(b). F. A. S.. Jacob. L. M. F. 1999. Lancet. 23–28. C. 601–8. R. W. 111. 128. 124. 129. J. The effects of Konjac flour on blood lipids in elderly subjects. D. and Third. Vidgen.. and Pikaar. B. Coll. F. Erkilla. H. Lee.... 1998–2004. C. DeGroot. 1976. Inglett. Leiter. Vidgen. Bran in hypertriglyceridaemia: a failure of response.. Dietet. V.. Stampfer. and Larsson. Cookies enriched with psyllium or oat bran lower plasma LDL cholesterol in normal and hypercholesterolemic men in Northern Mexico. Vorster. R.. Am. T. Dietary associates of serum total.. C.. J. J. Mehling. 119. 1998. A. L. Dintzis. 140. Am.. J. 128. 114.. M. S. Josse.. Z. 35. J. Effect of guar gum and oat fiber source on plasma lipoproteins and cholesterol in hypercholesterolemic adults. Seyler.. Anderson. C. Food Sci.. Sandstead. Logan. Assoc.. T. S. Diabetes Care. Clin. Wolk. E.. D. and O’Connor D. E. 824.. 113. E. 1146. H. Kendall. 120. Miller. Vuksan. 55–61. 127... L.. Matsuura. Culhane... B.. and Kay.. Story.. C.. Effect of wheat bran on serum lipids: influence of particle size and wheat protein.. R. Cholesterol-lowering effect of rolled oats. Clin. C. Brooks. S. Kruger. S. Owen. Vidgen.. E.. Prev.. H.. Long-term intake of dietary fiber and decreased risk of coronary heart disease among women.. E.. and Chen. J.. V. 1984. H. Oat-bran intake selectively lowers serum low-density lipoprotein cholesterol concentrations of hypercholesterolemic men. Effects of a concentrated bran fibre preparation on HDL-cholesterol in hypercholesterolaemic men. L...2387_ch5. 1963.. and DeVilliers. Augustin. Faulkner. A. V. Clin.. T. and Shuey. Serfonrein.. J.. L. 123. 122. M. G. Med. 1981.. and Nichols. Full-fat rice bran and oat bran similarly reduce hypercholesterolemia in humans. hypertriglyceridaemia and urate clearance. and Baba.. J. Beneficial effects of viscous dietary fiber from Konjac-mannan in subjects with the insulin resistance syndrome: result of a controlled metabolic trial. Manson. A. Jenkins. 2. Am. W.... F.. A. L. M. 159–65. Josse. 1979. F. F... W. Farquhar. C. 2001 7:37 PM FIBER IN THE TREATMENT OF HYPERLIPIDEMIA 419 110. Xu.. Clin. C. W. V. W.. 45. 865–9. L.. E.. Brighenti. 38C. 303. H. J.... J. Am. 226–30. 112. M. L. Nutr. 580. L. Luyken.. Spiller. Nutr. W. S. E. 11... Lehto. Hum. K. Ubbinik. C. W. Rees. Sievenpiper. Kawara. D. Ransom. Nutr. L. May 6. Klevay. P. Sieling.. J. A. and Lawrie. A. J. J. Brooks. Anderson.. M. L. R. E. J.. Lindegarde. Brighenti. Oshima. Swilley.. 23.. R. Vuksan. 281(21). M. Josse.. Arteriosclerosis Thrombosis. and Gallo. M. Med. L.. Nutr. Romero... and Fernandez. L. P. 1984. R. J. Br... D. R. Jenkins. Coll. D. R. Petro. K. G. Gerhardt. G. Munoz. J. J... P. 1999... K. Jr. S. 1999. L. Konjac-mannan (glucomannan) improves glycemia and other associated risk factors for coronary heart disease in Type 2 diabetes: a randomized controlled metabolic trial.. S. Effects of some cereal brans and textured vegetable protein on plasma lipids. Aust. Nutr.. N. Nutr. 28. A.. 1979.. Nutr.. W. Sieling. Guar gum and plasma cholesterol. Story. I. Bremner. Nutr. 118. Hypocholesterolemic effects of oat-bran or bean intake for hypercholesterolemic men. E. Galaviz. JAMA. 558–565. Terasawa. Can. 22. B. Hum.. G.. 1991. Nutr. 9–14. H. and Story.-J. R. Gates. S.. Hypocholesterolemic effects of high-fibre diets rich in water-soluble plant fibres. E. G. Anderson.. and Bruce-Thompson. Tsuji. Tsuji. S. S... 125.. 1204. Nutr. 1998. J... F. Sieling.. W.. H. Hennekens.. M.. W.. M. E. Vidgen. D. E. P. T. Leiter. Suzuki.. and Seki. Spadafora.. E. A. Treatment of diabetes with glucomannan Konjacmanna. Chen. 1984. 1. 2000. M.. J.. 117... Venter.. 17. B. Spadafora... and Willett. A. S. Lancet...... J. W. L. T. L. 1999. J. R. B. Vuksan. Bran. Doi.. J. J. H. R.. and Novokmet. Colditz Hu. Clin... W. Vuksan. Nutr.. M. Bremner. B.. P. . 121. H. 18. L. 2. F.. 1975. R.. 3. 2. 987–988. K. J.. Nutr. J. A novel source of wheat fiber and protein: effects on fecal bulk and serum lipids. S. 34. Romero.. 116. Sarkkinen.. Med. J. Ng.. W. Leiter. A. V. L. 39. C... Sievenpiper. Speizer. 69. 37. 913–919. Diabetes Care. LDL and HDL cholesterol and triglycerides in patients with coronary heart disease. Pacioni. M. Noseda. Diabetes Care. and Lenzi. Jenkins.. R. Grossi. Conti. 1981. Kalmusky. 137. 1982. Glycemic response to wheat product: reduced response to pasta but no effect of fiber.. B. Wolever. Metabolic effects of lecithinated and non-lecithinated textured soy protein treatment in hypercholesterolemia.. J. T... and Werner. Metabolism. Diabetologia. Nutr.. Coletta. Garsetti. 1999... F. D. J. S. A.. 49. L. S.. M. F.. 146. J.. Gaddi.. Cunnane. G. 1978. 1965. Nutr.. 1982.. 67–72. Piazzi. Jenkins. A. Anderson... Elsevier. 144.. T..2387_ch5.. Z. A.. Carter. Vessby. W.. 36C. 143. M.. C. J. Ceredi.. 24. R... 149. D. Chinese Med. and Hood. S. soy and vegetable protein foods... 747. Am. Evaluation of the hypocholesterolemic effect of vegetable proteins. L. Nutr. L. Agarwal. 45. L. Calvert. G. Effects of bean meal on serum cholesterol and triglycerides.. Simpson. i.. 86. fecal bile acids and neutral sterols in hypercholesterolemic men. R. A.. G. Newman. C. 1977. M. 333.. and Rongjue. May 6. Jenkins. 275. The glycemic index of foods tested in diabetic patients: a new basis for carbohydrate exchange favouring the use of legumes. 279.. Bird. C. B. Multicentre study of soybean protein diet for out-patient hypercholesterolemic patients. Moro. Lonsley. M. Rao A.. Seyler. F. O.. Lancet. 134. S. 138. C. Hum. R.. S.. Kendall. L. Burke. H. Med. 132. C. Br.C. 94. and Gatti. 1486. Lancet. Influences of soy protein and cascin on atherosclerosis in rabbits. Soybean-protein diet in the treatment of Type II hyperlipoproteinemia. L. ii. DiBiase.. 1981. B. R. Wong. 36Z. F. Gustafsson. Leiter. Jenkins. J. Clin. and Hooper. Clin. R. Lee. D. Eds. et al.. and Story. H. K. Buckley.. S. 1. R. Grasso. Mancini. Nutr.. Wolever. L..... K. Am.. Combined effect of vegetable protein (soy) and soluble fiber added to a standard cholesterol-lowering diet. 1979. 136. . W. C. R. 36.. Cappelli.... D. 1983. J. and Mann. S. Jenkins.. S. C... 147. I. D.. G... 709. V.. 179. and Keys. S. Stocki. G.. and Fidanza. Songiorgi. C. A. 455. wheat and potatoes on serum cholesterol concentration in man. 38. 257. Grande. M. 48. Wong. Med. R.. S. M... D. S. C.. 1974. V. and Cook-Newell. Hartog. and Davidson. 567. 27. Heston... Gaddi.. 1983. Contaldo. 1980.. 277. Nutr.. Nutr. P.and low-fiber diets on plasma lipids and insulin. Johnstone... Vidgen. E. J. E. J. McLeod... A. L. 3RD EDITION 130. A... D. Atherosclerosis. A trial of the effects of soya-bean flour and soya-bean saponins on plasma lipids. Nutr. D... 135. T. G. Patten. J.. G. O. McGuire. J. 2001 7:37 PM 420 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 7. Hall.. Maki. Mannino. Z. G. Fumagalli. J. Hockaday. J. Nutr. Leguminous seeds in the dietary management of hyperlipidemia. Karlstrom. G.. A.. Griffin. R. D.. B. J. Reichert. A. 148. Amsterdam. Meta-analysis of the effects of soy protein intake on serum lipids.4_fm Page 420 Sunday.. 1982. Kritchevsky. Agradi.. A. Am. A low-viscosity soluble fiber fruit juice supplement fails to lower cholesterol in hypercholesterolemic men and women. T.. A.. T.... Lithell. R. Reichert. Proc. Exp. Faulkner. B. Clin. G.. Zhaofeng. 37. Bingwen. 2000. A high carbohydrate leguminous fibre diet improves all aspects of diabetic control. Rubel... 133. S. Lee. Giacco. Appl. G. Mantero. A. W. Mehling.. Simpson. 313. C.. 1983. 32. Nutr. Biol... 142. R. C. 379. Sirtori. J.. Anderson. D. Davidson. W. Wanzhen. et al. and Little. W. F. Nutr. R. J.. Geekie. J.. Tepper. Am. Lancet. S. Fragiacomo.V. M. L. S... 141. Descovich. 1981. P. and Josse. 6. Cotter. Connelly. and Paoletti. L. J. B. F.. L.. C. 139. Holmes... Prevent. E. 140. E.. M. T.... 1983. 809–816. Soc. I. 128. The effect on serum lipids and oxidized low-density lipoprotein of supplementing self-selected low-fat diets with soluble-fiber. Assessment of the metabolic effects of dietary carbohydrate and fibre by measuring urinary excretion of C-peptide.. New Engl.. J. and Blight. A.. J. R. in Lipoproteins and Coronary Atherosclerosis. J. Fed. M... H. D. J.... Benassi. M. J. 150. W.. and Wong. and Keys. 1. 373. R. and Josse. Anderson. M. Clin. Jenkins. C. 1927–1932.. Sucrose and various carbohydrate containing foods and serum lipids in man.. 155. 131. Thorne. 1995... 1043. Fed.. M. Parker. I.. M.. Jenkins. Effect of high. M... S. J. T. P. W.. Albrink. J. Dicklin. Grande.. D. M. Comparison of the effect of dietary meat versus dietary soybean protein on plasma lipids of hyperlipidemic individuals. Hum. 1980. 276–82. 1998. Dugan. and Hoersten K.. M. 145. Vuksan. 138. 12. J. Descovich. Effect of carbohydrates and leguminal seeds. R. G.. A.. A. H. Ryan. Metabolism. The effects on lipid and carbohydrate metabolism of replacing some animal protein by soy-protein in a lipid-lowering diet for hypercholesterolemic patients.. 153. 247.4_fm Page 421 Sunday. D. 152. M.. 1984. R.. M. 1982..2387_ch5. R. and Vuksan V. C. A. Fumagalli. Jenkins D.. Leguminous seeds and their constituents in the treatment of hyperlipidemia and diabetes.. Clin... N... . E... Jenkins. in Lipoproteins and Coronary Atherosclerosis. Kendall C. May 6. Noseda. Viscous fibers. Am. B. 2001 7:37 PM FIBER IN THE TREATMENT OF HYPERLIPIDEMIA 421 151.. 50.. G. B. 401–2. Eds. S. B. Elsevier. and Paoletti. C.. and Jepson. Marenah. Miller. Nutr. A multifactorical diet in the management of hyperlipidemia.. W. Jackson. J. Katon. E. Choudhury. P.. J. Cortese. and Lewis. Fragiacomo. 93. Atherosclerosis. Amsterdam. C. 2000. J. health claims and strategies to reduce cardiovascular disease risk. 71.. 2001 7:37 PM . May 6.2387_ch5.4_fm Page 422 Sunday. In large part. colon cancer was observed to be uncommon in many developing countries but relatively common among age-matched inhabitants of most Western nations. the role of these dietary substances. EARLY FIBER STUDIES IN HUMANS Following the development of the fiber hypothesis. but not beef or other meats.10 Although this dietary characteristic is thought to be shared with certain African tribes. A casecontrol study from Israel6 described a highly significant inverse correlation between colon cancer and ingestion of certain fiber-containing foods. in the pathogenesis of colon cancer has not been defined.00+$1. Later. 2001 7:39 PM CHAPTER 5.7 A report by Graham and associates8 suggested that the frequency of ingestion of certain vegetables. if any. despite a diet low in plant fiber. Freeman ORIGIN OF THE FIBER HYPOTHESIS IN COLON CANCER The possible role of dietary fiber in human colon cancer pathogenesis became of particular interest following epidemiologic studies in different populations. are not unusually prone to colon cancer. Certain groups of Inuit in the Canadian Arctic. where the usual diet contains less cellulose.9 yet the Inuit eat large amounts of animal connective tissues composed of apparently nondigestible aminopolysaccharides. however.3–5 and the possible role of fiber in prevention of the disease was emphasized. was lower in white males with an increased colon cancer incidence.5% fiber. Seventh-Day Adventists in Loma Linda. Black patients from the San Francisco Bay area reported less frequent consumption of foods containing 0. either supporting or refuting the importance of dietary fiber in colon cancer pathogenesis.2387_ch5. Interviewers were not aware of which patients had histologically confirmed malignant disease of the large bowel. Malhotra2 subsequently found a low incidence of colon cancer in northern India. and broccoli. where the usual diet apparently contains large amounts of dietary fiber. a number of epidemiologic reports appeared. California. are reported to eat vegetarian diets11 and have a low incidence of colon cancer.12 However. and a high incidence of colon cancer in southern India. the fiber hypothesis was widely popularized by Burkitt and colleagues.50 © 2001 by CRC Press LLC 423 . 0-8493-2387-8/01/$0.5 Human Studies on Dietary Fiber and Colon Neoplasia Hugh J. especially cabbage. sprouts.5_fm Page 423 Sunday. the hypothesis that fiber consumption may prevent subsequent colon cancer development appears to have emerged subsequent to the report by Higginson and Oettle1 on studies in the Bantu of rural South Africa. although methods used to determine fiber content were not indicated. May 6. . the mean intakes of total dietary fiber as well as specific fiber polymers (cellulose and lignin) were significantly less in Copenhagen. drew attention to studies showing a protective role for vegetable intake. Although preliminary. Some of these have attracted significant media publicity.15 Haenszel and colleagues16 found a significant positive association between colon cancer and the frequency of ingestion of fiber containing legumes in Hawaiian Japanese. Most provided evidence for a protective effect. especially cruciferous vegetables rich in indoles. cabbage was reported to be protective.5. such as familial polyposis. Although 121. Over 16 years. 3RD EDITION other malignant diseases also occur at a low incidence in this group. follow-up studies included questions related to specific dietary constituents. differences in dietary fiber consumption within that country were correlated with its apparent protective role in colon carcinogenesis. cohort studies.700 nurses were initially enrolled.18 on the other hand. and precise measurements of the content of individual fiber polymers in their diets have not been performed. However. Although the proportions of dietary fiber polymers were similar. international and within-country correlation studies. in other words. Review is critical to further refine the direction of future research.13 In another. In the past decade. In a recent study by Fuchs et al. discrepancies that emerged may have also reflected inadequate definition of the precise dietary fiber composition of foods ingested by the different groups.65 nurses who completed a retrospective dietary semiquantitative food frequency questionnaire in 1980 were evaluated.. both conditions that predispose to colon cancer.5_fm Page 424 Sunday. Finland) and an area with a high incidence (Copenhagen) were analyzed.000 were excluded from the final analysis because of recognition of a potential high risk factor.64 the effective risk reduction was less for those with a family history of colon cancer. while some revealed only equivocal results or no evidence of a protective effect for fiber in colon cancer. 2001 7:39 PM 424 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. several epidemiologic studies appeared (Table 5. All of these recent studies have aided in definition of critical variables (Table 5. such as inflammatory bowel disease (see Chapter 5. or loss of follow-up. In another study from Japan. no other quantitative data were provided. Using the Southgate method of fiber measurement. almost 33.17 Graham and Mettlin. Other earlier studies from Europe20.19 While it was difficult to conduct studies on the basis of retrospective dietary histories dependent on respondent recall.2) for subsequent studies important in further definition of dietary fiber in colon cancer. diets from population samples in an area with a low incidence of colon cancer (Kuopio. vegetables. RECENT STUDIES AND FUTURE DIRECTIONS These early studies served to stimulate considerable interest in the role of dietary fiber in colon cancer pathogenesis. or fruit.1). Nurses were .21 further attempted to examine the fiber question from this perspective. Some international and in-country studies were also done soon after this hypothesis was developed.14 In a report from the U. Indeed. grains. Particular emphasis may be needed for prevention studies in those with colonic disorders. Similar observations supporting a possible protective role for vegetables were also previously noted. these included case-control studies. there may be individual genetic factors that influence the potential benefits of dietary fiber and other environmental factors in risk reduction.5. no correlation between crude dietary fiber consumption and colon cancer mortality was observed. measured intakes of specific fiber polymers were documented for the first time and differed in human populations with different incidence rates of colon carcinoma. May 6. In one. and timetrend studies concerning colon cancer and fiber. In total.K. cereal consumption appeared to be negatively correlated with colon cancer incidence data. Clearly. genetic factors may be so significant in some highly predisposed individuals that it may be difficult to define any potential beneficial role for dietary fiber. 787 cases of invasive adenocarcinoma and 1012 cases of adenoma in the distal colorectum were detected.7) and colon polyps. In a recent study.2387_ch5. several studies have been published. 1988 Graham et al.. No difference in colon cancers or polyps was detected between different fiber groups.. 1989 West et al.5_fm Page 425 Sunday. Several issues were not addressed in this highly publicized study. 1994 Tsuji et al.. 1997 Fuchs et al.. 1988 Lyon et al. 1990 Giovannucci et al. 1981 Tuyns et al.2387_ch5. 1998 Sellers et al. 1999 a 43 44 45 46 47 48 49 50 51 52 53 54 58 60 65 Meta-analyses of other published studies.. 1988 La Vecchia et al. 2001 7:39 PM HUMAN STUDIES ON DIETARY FIBER AND COLON NEOPLASIA Table 5. 1984 Rozen et al. May 6.. Evidence for Protective Effect Tuyns... 1983 Willett et al. 1985 Potter and McMichael. 1988 Helms et al.. 1994 McKeown-Eyssen et al.. Food-frequency questionnaires often fail to provide a true estimate of dietary macronutrient or ... 1996 Hardman et al. 1988 Young and Wolf.. 1986 Miller et al. The highest fiber diet was about 25 g per day. 1987 Slattery et al. 1984 Bingham et al.. 1997 Franceschi et al.. First. 1987 Bristol et al... 1990 Olsen et al... Year Ref.1 425 Studies on Fiber in Colon Neoplasia Author. 1986 Kune et al. 1997 Ghadirian et al.... 1985 Rosen et al. 1988 Heilbrun et al... 1984 Tajima and Tominaga... 1981 Pickle et al.. the measures used to collect data were imprecise and based on retrospective diet recall. 1999 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42a 55 56 57 59 61 62 63 64 66 Equivocal or Lack of Protective Effect Martinez et al. 1989 Freudenheim et al.. 1990 Trock et al. 1983 Boing et al... categorized into quintiles depending on computed fiber intake from the retrospective dietary data.. 1983 Macquart-Moulin et al.. 1985 Powles et al.5. 1989 Lee et al. 1981 Jensen.. 1982 Hirayama. 1985 Phillips and Snowdon... 1998 Macrae. 1985 Tajima et al. 1994 Steinmetz et al. 1985 McKeown-Eyssen and Bright-See.. 1994 Alberts et al. 1986 Manousos et al. Second. an future research.5.. particularly in relation to individual fiber components. Cancer incidence in Bantu and “Cape coloured” races of South Africa. 24. 8. B. HLA type. Cancer Inst. 15. including the methods used to measure the endpoints of data collection.2 Variables for Fiber Studies Dietary Factors • • • • • • Individual fiber components (cellulose.g. 1976. 3RD EDITION Table 5.. J. A. ulcer drugs) Medications for prophylaxis Colonic Factors • • • • Colonic site-specific effects (e. etc. J. N. A. 1972. and Oettle. may be missed because of the reported time-lag effects of dietary fiber in colon cancer. Modan. Dales.. hemicellulose. endoscopic evaluations and pathology reports may differ depending on levels of expertise. Am.. R. and Burkitt.) Digestion products of fiber (short-chain fatty acids. Walker.) Environmental variables (smoking. but specific components of fiber in animal fiber studies have been shown to be critical for cancer prevention.) Fiber–micronutrient interactions (trace elements.. with a maximum 23-year lag. Total dietary fiber measurements are of interest. Lubin. Natl. D.. 910. 1975. more definitive studies. . Dig. Greenberg. A. Cancer Inst. Geographical distribution of gastrointestinal cancers in India with special reference to causation. K. etc. lignin. P. 1960.5-fold difference in estimated total dietary fiber between the highest and lowest quintile. Moreover. P. L.. Ury. vegetables.e.. J. estimated to demonstrate a maximal negative correlation after a 15. Cancer. P. 55. 1967.) Source of dietary fiber components (grains. S. D. May 6. R. L. D. protein. dietary prophylaxis. there was only a 2. V. Finally. Report of a cancer survey of the Transvaal (1953-1955). Friedman. G... 361. Natl. R. P.. Epidemiol.. A case-control study of relationships of diet and other traits to colorectal cancer in American blacks. 28. Moreover. etc. Colonic cancer — hypothesis of causation.. J. Walker. etc. such as cellulose. vitamins. and Williams. 2. 7. 5. P. Gut.) micronutrient intakes. Grossman. rectosigmoid) Preneoplastic disorders (ulcerative colitis.) Food preparation methods Patient or Population Factors • • • • • • Sex and age Geographic locale Genetic variables (blood type.2387_ch5.5_fm Page 426 Sunday. Burkitt.68 REFERENCES 1. etc. Low-fiber intake as an etiologic factor in cancer of the colon.) Fiber–macronutrient interactions (fat. cecal vs. Dis. S.. Crohn’s disease) Other colonic neoplastic disorders (colon polyps. 3. and Graham. etc. 132. 1971. etc. 2.59 Clearly. other important concerns have been subsequently raised. J. S. A. 4.. 10. including dietary fiber. D. Malhotra. S... 1408.. H.. important differences. Barell. high-fiber foods may not be equivalent. even if present. S. M. polyposis syndrome) Underlying gastrointestinal or other diseases (diabetes. 6. G.. F.. Higginson. 1979.67.to 27-year delay. Am. Effect of dietary fiber on stools and transit-times. i..) Medications for treatment (antibiotics. and its role in the causation of disease. 21. Modan. Epidemiology of cancer of the colon and rectum. likely not sufficient to detect any differences.. R. etc. pectin. 2001 7:39 PM 426 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and Painter. 589. Lancet. the evaluation of this study raised many criticisms that need to be considered in the design of future.. Burkitt. G. 1474.. J. P. D. S. 1988.. J. 492. International Agency for Research on Cancer... R. W. R. Gardner. Bingham. Cancer Inst. T.. J. J. Diet and colon cancer. R. 486.. Dietary fibre and regional large bowel cancer mortality in Britain... Schaefer. and Karn.. and Wilkinson. Eugen. T. 1977. 81.. 31. Dietary fibre. 167. Natl. 1042. 78. 29.5_fm Page 427 Sunday.. J.. H. 38. 1765.. Cancer. 16. B. W.. 61. 1979. K. 27. Med. R.. Environmental factors in cancer incidence and mortality in different countries with special reference to dietary practices. 21. 64. transit time.. Viewpoints Dig. fat and the risk of colorectal cancer. French. Praventivemed. Graham.. Can. 2001 7:39 PM HUMAN STUDIES ON DIETARY FIBER AND COLON NEOPLASIA 427 8. Marshall. J. Large bowel cancer in Scandinavia in relation to diet and faecal characteristics. Clin. 1489. Int.. Cancer. M.. 5. Br. 5. Wahrendorff. 167. K. S. J. 396. W. E.. Walden. D. 490. Natl. A case-control study of diet and colorectal cancer in northern Italy. F.. Lemon. 1. S. 31.. 34. 1987. L. 40. J. Assoc.. Mcquart-Moulin. J. 3. Case-control study on colorectal cancer and diet in Marseilles. and Locke. Int. Kritchevsky. Fiber bibliographies and terminology. Haenszel. S. Natl. 1983. Med.... R. C. H. Case control study of dietary etiological factors: the Melbourne colorectal cancer study. 13. Preliminary report of a population study. Natl. P. 28. Sorenson. Riboli. Bingham. P. Nutr. dietary and family histories of 450 cancer patients and an equal number of control patients.... 12. Mahoney. J. Williams. 1964. 853. and Cummings. Kune. Negri. S. A. E. Mahoney. F. J. Soz. Swanson.. Intestinal Microecology Group. M. 1985. Manousos. A. L. J. Epidemiol. Diet and colorectal cancer: a case control study in Greece.. Large bowel cancer in Hawaiian Japanese. S.. R... Colon cancer connection: beef. Cancer. T. O. G. Cancer Inst. C.. Cancer Inst.. 24. J.. 19. and Spiller. Environmental factors in cancer of the colon and breast. Nutrition and nutritional deficiencies. and Bright-See. Cancer. J.. 20. 1988. home life.. L. 81. 26. 10. Haenszel. and Watson. Charnay. 15. W. 4. O.. D. B. W. and Heaton. J. 41.. and Irving. 183. J. 21. G. 23... D. 291.. 1989). W. 1984. Br. 9. Int. Dis. Cancer. 207. Cancer. Berg. steroids. and Gorbach. W. Zielezny. West.. 10. 1933. J. 80... 52. A. W. Am. Lyon. 9. 15. Segi.. J. H.. G.. Goldin. Br. 128. and Mettlin.. 237. J. Draser. K. Armstrong. Graham. 1987. C. Wilkinson. Dietary factors in colon cancer: international relationships.... J. and IARC. 127. T. Diet and cancer mortality in the counties of Sweden.. 1982.. M. Am. 1975. and Wolf. R. 1978. A. M. M. 1973. Young. W. Day.. Large Bowel Cancer Group. M. 1973. Natl. 25. Nutr. Natl.. J. Case control study of proximal and distal colon cancer and diet in Wisconsin. E. J. 1959. Bristol. Mittelmann. Kune. Ann.. Lancet.. Emmett. 22. Cancer.. W. 42. II. D.. K. Cancer Inst.. and Segi. and Trichopoulos. Swanson. A case control study of large bowel cancer in Japan. A. 1.. Berthezene. M.. L. 6.. T. Am.. Nutr. A. J. 456. 32. M. Energy intake: its relationship to colon cancer risk. 51.. Nystrom. D. A cooperative study of the habits. B. MacLennan. and Stanish. 1988. M. and West. B. B. 17. and Doll. Cancer Inst. Cancer. W.. and Decarli. Tuyns. 30. L.. 1986. T. 2. G. M. N. A. 31. 11. Cancer. Cancer. 14.2387_ch5. An update. B.. Dietary epidemiology of cancer of the colon in western New York.. Br. N. Diet and colon cancer: assessment of risk by fiber type and food source (published erratum appears in J. J. Sorenson. faecal bacteria. A. and Day. D. A.. A. E. Godding.. Nutr. A case-control study on colorectal cancer in Belgium. Sugar.. 1985... and James. 27. 81. 109. bran. bile and bacteria. Cancer. Locke.. Haughey. . G. 1986. J. 1988. and Street. Int. Williams. J. 617.. E.. S. 32. F. 42.. W. May 6. E.. 1979. and colon cancer in two Scandinavian populations.... 18. O. and Wall. D. Cancer. 1980.. Cornee. B. Cancer of the lung and mouth in Seventh-Day Adventists. Trowell. J. F. P. M. Am. Smith. Byers. F. 160. Medical observations and problems of the Canadian Arctic.. 1978. Cole. Int.. 1467. S. Mittelman. Jensen. Epidemiol. M. B. 1988. 399. McKeown-Eyssen. and Woods.. R. 33.. 17. Diet in the epidemiology of cancer of the colon and rectum.. Epidemiol. G. Dietary fibre consumption in Britain: new estimates and their relation to large bowel cancer mortality. R.. W.. S.. A. Stocks. 709.. Dayal. Rosen. J. Cancer Inst. Slattery. Graham. Kurihara.. R. La Vecchia. B. 17. J. 1978. J. 54. 44. J. 557. Colditz. 3RD EDITION 35. 307. P. Marshall. and Sorensen. Trends in bowel cancer in selected countries in relation to wartime changes in flour milling. Japan.. Nomura. Z. 13. 70.. Oncol. 1981.. 1985. A. 47. 307. 650. P. 38. L. Natl. 50. S.. and Potter. J. 1990. Denmark. M. 47. and Nakagawa. F. Int. Dietary risk factors for cancer and adenomas of the large intestine. 1988.. 1985. Vegetables.. 155. Dietary intake and colon cancer: sex. B. W. Cold Spring Harbor Laboratory. Epidemiol. L. and Snowdon.. and Haelterman. E. Jpn. J. Helms. 1994. and Horwitz. Kaaks. Horvath. W.. Estieve. R.. K. Ford. E. J. L. Natl. A. O. Greene.and anatomic site-specific associations. Jazmaji. F. P.. R. L.. R. and Ewertz. A. Folsom. 1989.. Diet and colorectal cancer with special reference to fiber intake. R. vegetables. J. Engl. 1985. J. Jensen... 48. Natl... 705. 56. 1983. 1011. Rozen. H. M. Cancer Inst... Colorectal cancer and diet in an Asian population — a case-control study among Singapore Chinese. Kronborg. 525. 1983. Haughey. Bruce. fat. Torres.. H. 40. Cancer Res.. J. Phillips. K. 76.. Lyon. M. M. K. Mahoney. Colorectal cancer and the consumption of foods: a case control study in Belgium. K. Cancer Res. A. B. 37. Dietary relationships with fatal colorectal cancer among SeventhDay Adventists. V.. 6. and fiber intake to the risk of colon cancer in a prospective study among women.. 43. G. 130. 1984. J. J. 74.. Epidemiol. Freudenheim. 76.. Cancer. 1989. 44.. T. A. Jorgensen. Robison. W. Olsen. J. Colorectal cancer in rural Nebraska. L.... Clin Epidemiol. Rosner. N.. 1989. 1007. J. and Saibil. W..... 717.. and colon cancer in the Iowa Women’s Health Study. Lanza. Hirose. G. Lynggaard. J. R. Miller. I. J. J. 55. and Williams. A. J. Martinez... W. 53. M. 53. Bright-See. H. Urban-rural difference in the trend of colo-rectal cancer mortality with special reference to the subsites of colon cancer in Japan. R.. M. L. 409. 323.. R.. Potter. M. 42. W. Cancer. Hankin. B.. 189. 1986. Regional nutritional pattern of cancer mortality in the Federal Republic of Germany.. in Gastrointestinal Cancer: Endogenous Factors. P. J. G... A. Dietary patterns in Them and Copenhagen. West. H. 76. Cancer Inst.. S. Latinoam. A.. A. D. N. Cancer Inst. E. and Wilkinson... Risks associated with source of fiber and fiber components in cancer of the colon and rectum. and McMichael. D. A randomized trial of a low fat high fibre diet in the recurrence of colorectal polyps. J... J.. Kushi. Correa.. J. Hellerstein. S. Eur. K. Diet and cancer of the colon and rectum: a case-control study. C. G... Cancer. 46. 139. L. 1990. 49. J.. P. 1. J. 30A. 1984. Clin. Dietary fiber. Graham.. L.. R. Cancer Inst.. 57. P. R. 41. E. L. W. Nutr. Cancer. L. A case-control study within a screening trial in Denmark. Bostick. and Harrison. and Tominaga. R. Lee. S. Tajima. 1994. D. R.. Howe.. E. 1981. and Paerregaard. Duffy.. 883.. D. 1. A. N. H.. M.. G. Int. Am. 43. Lee. M.. J... Cancer. C. 1. May 6. G. Powles. Jpn. NY. 1981. and Frentzel-Beyme.. J. Relation of meat... 4.. 40. Boing... 13.. Eds. B. Cancer. 48. L. Heilbrun. Natl.. 44. Natl. K. Tajima.. 39. 1664. Willett. The low incidence of colorectal cancer in a “high risk” population: its correlation with dietary habits. 1982.. . Am. 82. M.. 1985. Stampfer. A. K. J. 363.. Toronto Polyp Prevention Group (published erratum appears in J.. M. Bruce. Cancer Res. Hirayama. and Frias. W.. Rev. Tuyns. McKeown-Eyssen. Epidemiol. J.. B.5_fm Page 428 Sunday.. and Day.. M. Nutr. 48. 1990. J. Gourley. Factors associated with adenocarcinomas of the large bowel in Puerto Rico. H. Clin. 45. Schuman.. Int. and colon cancer: critical review and meta-analyses of the epidemiologic evidence.. 1995). Cancer Res. 52. J. Dietary habits and gastro-intestinal cancers: a comparative case-control study of stomach and large intestinal cancers in Nagoya. and Lipkin. C. P. A... O. 2692.. 34. Cancer.. J. Steinmetz. Craib.2387_ch5.. J.. 2001 7:39 PM 428 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. relationships with fatal colorectal cancer among Seventh-Day Adventists. Cohen. Trock. 51. 1994.. Cancer. Slattery. Pappas.. D. N. and Speizer. 74.. A large-scale cohort study on the relationship between diet and selected cancers of the digestive organs. D. Cold Spring Harbor. W. Jain.. 11. 32.. L. Med. Food items and food groups as risk factors in a case-control study of diet and colo-rectal cancer. Martinez. 36. Cancer risk among Danish male Seventh-Day Adventists and other temperance society members. L. Cancer Inst. and Stemmermann. R. Nutr. and Greenwald. M.. 50.. I. S. J. and Ziegler. Pickle. fruit. 3295. . 1997.. S. Wheat bran fiber and development of adenomatous polyps: evidence from randomized controlled clinical trials. E. N.. Sci. 1999. Colditz. Cummings. Role of high fibre foods in the prevention of colorectal neoplasia. C. and Shirataka. 1924. K. P.... 1998. Harashima. A.. P... Ritenbaugh... Perret. Kushi. M. and Lang. and Willett.. and Clark. and Willett... Time-lag effect of dietary fiber and fat intake ratio on Japanese colon cancer mortality. 80. A... Anderson. 1999. N. Cancer Epidemiol. M. J...5_fm Page 429 Sunday. P. meat. Ghadirian. Giacosa. 9. Atwood. Dietary fiber and the risk of colorectal cancer and adenoma in women. 67.. D. Nakagawa. 66.. Giovannucci. Alberts.. E. Cancer Causes Control.. 340. 38S. 65. 858.... J. J. J. G. 2390. T. Sellers. Lewis. G. S19.. Stampfer. J. Engl.. Med... Quebec. W. S. Bernard. L. Potvin. D.. H. Bellapravalu. Madar. M. E. J. Italian study on colorectal cancer with emphasis on influence of cereals. S. 54. ... C. A.. Einspahr.. H. Favero.. Gravel.. S. 379. 13. A. Speeg. A. 59. Hardman. C.. and Folsom. J. Cancer. S. Canada. W. 68. Cancer Res. Aickin. C. J. M. and La Vecchia. Transforming growth factor alpha distribution in rectal crypts as a biomarker of decreased colon cancer risk in patients consuming cellulose. A. Am. Med. Southgate. Diet and risk of colon cancer in a large prospective study of older women: an analysis stratified on family history (Iowa. D. P.. N.. F. D. Ravin.2387_ch5.. W. Med. M. Kadakia. and Camire. 1997. Parpinel... Franceschi... K. H.. 1999.. 1999.. Cameron. 61.. 169. 60. Intake of fat... E. C.. 7 (suppl. E. E. W. R. D.. C. Ramanujam. A. 1997. F. E. Biomarkers Prev.. 161. T. Biomarkers Prev. C. Nutritional factors and colon carcinoma: a case-control study involving French Canadians in Montreal. Stampfer. 340. M. L. L. 1996... K. Giovannucci. J.. J. May 6. 6.. A. Rimm. Olson. Fuchs. 63. A. L. E. Canad. K.. J.. J... 223. B. C. and Boyle. Rosner. United States).. 1998. Emerson. S.. A. Speizer. D. N. Dietary fiber and colorectal cancer. 6. 106 (1A). J. Bostick. Z. A. Ascherio. M. L. Cancer Epidemiol. Heaton. E. Environ. Y.. Urata.. Lacroix.. R. B.. G.. 357. J. The effect of wheat bran fiber and calcium supplementation on rectal mucosal proliferation rates in patients with resected adenomatous colorectal polyps.. K... Rees-McGee. J. Eur. I. Lazovich.. and fiber in relation to risk of colon cancer in men. Mason-Liddil. Mohandras... Bettinger. 64. 62. Freeman... A. Phelps. S. K. M.. 9. S. Colditz. J.. Cancer Prev. 1994. Gastroenterol. 2001 7:39 PM HUMAN STUDIES ON DIETARY FIBER AND COLON NEOPLASIA 429 58. S. Tsuji. Hunter D. Maisonneuve.. Beer. Bazyk. Macrae. Biomed. Engl. Stark. Patel. W.. V. E. H. 633. 2). May 6. 2001 7:39 PM .2387_ch5.5_fm Page 430 Sunday. 1.2387_ch5. Painter and Burkitt.4 Subsequently.5%. sepsis. indeed. but these techniques were not available to earlier clinicians. Barium enema and colonoscopy are the most common current methods of diagnosis.5 Later.7 suggested that diverticulosis is rare in many parts of Africa. Brodribb has cited several autopsy studies reporting very high prevalences of diverticular disease (Graser in 1899. using a variety of anecdotal sources. therefore.9 To date.50 © 2001 by CRC Press LLC 431 . While precise figures are not available. except among Europeans. and hemorrhage.7 precise analyses are not available. and uncommon in the Indian subcontinent. More interesting information comes from geographic studies. historical data providing strong support for an increased prevalence of colonic diverticulosis at the present time are limited and controversial. especially the sigmoid colon. and South America. Painter and Burkitt published their hypothesis that diverticular disease was caused by a reduced intake of dietary fiber.3 Dietary fiber was recommended for symptomatic diverticular disease as early as 1929 by Spriggs.2 Although there are different anatomical forms of colonic diverticulosis. Sudsuki in 1900. 30%).8 Interestingly. Middle East. this is a herniation of mucosa and submucosa through the colonic muscle wall.6_fm Page 431 Sunday.6 Fiber and Colonic Diverticulosis Hugh J. and Mourges in 1931. the basic abnormality observed in most North American and European populations is the pseudodiverticulum. May 6. 37. 2001 7:40 PM CHAPTER 5. Far East. 0-8493-2387-8/01/$0. but only a small minority will endure more serious complications including diverticulitis. this “fiber hypothesis” was examined in carefully matched population groups from Oxford. obstruction. all are economically developing areas with cereal-based diets typically high in fiber. these are multiple and involve the left side of the colon.1 In developed nations. radiographic appearances were not described in detail until 1925. Most often.6 A variety of epidemiologic studies on the relationship between dietary fiber consumption and diverticular disease have been done both from a historical as well as a geographic perspective.00+$1. the disorder is extremely common and prevalence correlates well with increasing age. This contrasts with the well-developed countries in Europe and North America where diets tend to be highly refined and fiber-depleted and where a high prevalence of diverticular disease is observed. 64%. it has been estimated that about 20% of patients with diverticulosis will develop symptoms and signs of illness. Although it has been suggested that the prevalence of diverticular disease has increased over the past century. diverticulosis was observed to be significantly more frequent in non-vegetarians compared to vegetarians. Freeman Colonic diverticular disease is an acquired deformity of the colon that is generally irreversible but usually asymptomatic. 25 have also explored factors that may influence the appearance of increased numbers of colonic diverticulae in rats administered a fiber-deficient diet. the most comprehensive and best-conducted study. indeed.6. including Japan. Additional studies. many dietary fibers are more extensively degraded in these animals than in humans. especially for cellulose fiber.1). the role of bran was examined in five controlled trials (Table 5. In one prospective study in American males. but the presence of a large cecum in these animals raises doubts regarding the applicability of these results to humans. It appears that a high-fiber diet may be effective treatment for symptoms in some patients with uncomplicated diverticular disease. it is unknown how accurately these reflect the true frequency in the population as a whole. 5.11 but not in the Orient. a major weakness of such correlative studies relates to the dietary component of the equation and. In one interesting report. a number of uncontrolled trials suggested that added fiber in the form of bran supplements may be therapeutically beneficial. and 0 g per day. especially if symptoms are severe. some supportive evidence for a role for deficient dietary fiber in the pathogenesis of diverticulosis comes from animal studies. the same investigators noted that maternal diet during gestation. Finally. May 6. Brodribb et al. In one report. including fiber intake. As early as 1937.20 fed low-fiber diets to rats and showed the development of diverticulae. methods used to precisely calculate the fiber content of the diet or specific fiber components.13 Similar observations have been reported from Europe. the true incidence of diverticular disease in these populations is not known. ispaghula alone31 and methylcellulose alone32 have been studied in controlled trials. are still required to determine if high-fiber diets can alter the natural history of diverticular disease over the long term or prevent its complications. Similar observations have been reported in rats and rabbits. but the data remain limited.34 In a subsequent report of a prospective study of different dietary fiber types. 2001 7:40 PM 432 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. low-fiber diet compared to diets with low fat and higher fiber content. recent studies on the distributional pattern of diverticular disease contrast with the increased frequency in the left colon in Western communities compared to the right colon in Oriental populations. Hyland and Taylor33 described 100 patients that had been retrospectively reviewed after 5 to 7 years on a high-fiber diet — over 90% remained symptom-free.12 Indeed. Lubbock et al.24 a high-fiber diet appeared to protect against collagen cross-linking with a resultant reduction in appearance of diverticulae.25 Subsequently. Although a definitive role for fiber-deficient diets in the pathogenesis of colonic diverticulosis has not been proven.6_fm Page 432 Sunday.29 largely using patients with mild symptoms. showed no major benefit in pain scores but there was improvement in constipation.2387_ch5.23 Using an experimental model. in part because of variable diagnostic methods and availability of accurate postmortem studies.35 . In those populations with available data.17–19 In addition.22 chose the stub-tailed monkey with a gastrointestinal anatomy more similar to humans as a model for colonic studies. subjective improvement was reported with ispaghula but no effect was noted with methylcellulose. and subsequent nutrition of progeny may play a critical role in development of diverticulae. Later. the incidence of symptomatic colonic diverticular disease was lowest in those with high-fiber diets. He found that colonic intraluminal pressure increased as the amount of fiber in the diet decreased gradually from 20 to 15. 10. some studies24. 3RD EDITION This relationship between decreasing dietary fiber and increasing prevalence of diverticular disease has also been reported in urban South African blacks10 and Hawaiian Japanese. More recent studies in adult female vervet monkeys revealed that diverticulosis frequently developed in those administered a Western-type high-fat.26–30 In addition.14–16 Although intriguing. while only 4% of rats fed a psyllium seed supplement developed diverticulae. this inverse relationship with measurable insoluble fiber was confirmed. Moreover. specifically. Carlson and Hoelzel21 found that about 57% of a colony of Wistar rats more than 100 weeks old had diverticulae in the proximal colon. especially in humans. . Spiller. R. Yamagata.. J. placebo (either wheat crispbread or refined wheat) Coarse bran vs. 6. 72. A. 1977. J. 511. 4. 11. Wheat vs. 75. J. I. .. and Hunt. Med. Brodribb. T. J. 31. The distributional pattern of diverticular disease. R. 13. Clin. D. 19. but diagnostic method not defined Sterculia or bran × 4 wk Bran crispbread vs.. 450. J. and Yatani. hyoscyamine vs.. M. D... 302. Eds. 2. bran crispbread 18 with X-ray diagnosis No.6. pain scores subjective 26 Wheat bran (coarse) vs. Clin. Clin. 7. Watanabe. E. 215. and Kay.. and Marxer. Solomon... S. New York. S. A.. Gear. A. 569. 32. 4. placebo 58 with X-ray diagnosis controlled trial × 16 wk Double-blind randomized crossover Symptoms relieved to an equivalent extent in all groups. Gastroenterol. sterculia with or without antispasmotic 20. Tohoku J. near normal stool weights and transit times 29 Symptom scale based on ability to work 30 REFERENCES 1. P. Brodribb. 1980. Diverticular disease of the colon. A. J. 4. and Howell. isphagula vs... 3.. 1260. N. S. Br.. Mann. 43.. in Medical Aspects of Dietary Fiber. Dis. Painter. Intestinal diverticula.. Segal. N. 1980. and Leibowitz. Exp. 1975. Tajima. Munakata. E.. Nolan. 2001 7:40 PM FIBER AND COLONIC DIVERTICULOSIS Table 5. Gastroenterol.6_fm Page 433 Sunday.. I. Cancer. N. Stemmermann. G. and Vessey. 147. obstruction and bleeding. 324. D. Parks. 227. Spriggs. Br. patients Double-blind control trial × 12 wk Study Protocol Reduced symptom pain scores.. A. 8. 53. Lancet. T. P. 1979. Symptomless diverticular disease and intake of dietary fiber. Q. 1973. P. 115. F. A. Med. 9. but bran or sterculia with antispasmotic better than sterculia alone for pain relief Pain score not improved but improvement in constipation 105 with X-ray diagnosis Control trial up to 52 wk Improved symptom scale for bran and hyoscyamine 27 28 Pain symptoms mild in degree. 1925. 1971.. 1975. Emergence of diverticular disease in the urban South African black. T. P. Diverticulosis and polyps of the large intestine: a necropsy study of Hawaii Japanese. Fursdon. Plenum Press. Spriggs. M. lactulose 31 with X-ray diagnosis Control trial × 4 wk Coarse bran vs... 10. and Burkitt. May 6. 1. Diverticular disease of the colon. N. I. a 20th century problem. 5. Complications of diverticular disease: inflammation. 12.. G. M. Natural history of diverticular disease of the colon. Painter. B. Diverticular disease of the colon: a deficiency disease of western civilization.. Statistical analysis of diverticulosis of the colon. D. 4. Colon Rectum. and Matatsunaga. Almy. Narasaka. J. Hughes. J. and Burkitt.2387_ch5. Ware. 2. 1. Gastroenterol.. L. Segal.. G. 3. Med... Med. 271. early placebo effect noted Symptoms moderate to severe. Engl. L.. A.. M. P. isphagula vs. H.. G.. 1989. Gastroenterology. J. 2.. Diverticulitis. O. S. Med..1 433 Controlled Clinical Trials of Bran in Colonic Diverticular Disease Fiber Type and Form Results Comments Ref. I. 1929. Dietary fiber in diverticular disease of the colon. only bran reduced colon motility and pressures Equivalent improvement in constipation. A. T. 1975. E. J.. . Diverticular disease: three studies. Thomson. MTP Press.. G.. Am.. Ornstein.. is it important in diverticular disease?.. Eastwood. C. J. 39. North. H. G. Am. 315. A. Hodgson. 714. W. J. Chir. 1994. Giovannucci... Brodribb. J. J. 77. 60. Epithelial overgrowth and diverticula of the gut. A. 1252. Suppl. Edwards. Controlled studies with dietary fibre in the therapy of diverticular disease and irritable bowel syndrome. S. 1977. W.. The placebo effect.. 28... I. Weinreich. J. Ewerth.. B. J. 18.. Gastroenterology.. Treatment of symptomatic diverticular disease with a high fiber diet. A. Med. Scand. D. 1976. B... 30. 423. 10. 239. Gut. Unprocessed bran in treatment of diverticular disease of the colon. S.. and Miller. and Hoelzel. Lancet. and DeCosse. Med. and Woodroof. Nutr. Br.. A. Goebell. and Humphreys. Almeida. D. 33. Collagen alteration in an animal model of colonic diverticulosis. J... W. ispaghula.. A. and Francis.. D. W. C.. Busuttil. 1971. 49. 25. Wess. P. C. . A prospective study of diet and risk of symptomatic diverticular disease in men. U... Edwards. B.. Wess. Holmstrom.. Busuttil. Lancaster. Brydon. 21. 1996. H.. 128. Med. H. 1976. Fowler. L. 1996. and Kasper. Rockett. Gastroenterol. J. R. 2001 7:40 PM 434 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 1986. Painter. 753.. 167. L.. 1937. Plumley. J. 664. K. and Taylor.. Br. D. England.. 108. 148. 126.. P. J. L. Rossouw. N.. Aldoori.. and Painter... 34. 19. A.. Eastwood. L... 500. The incidence of colonic diverticulosis in Finland and Sweden. K. in Colon and Nutrition. Acta.2387_ch5. 1998. 2. T. N. Baird. Brodribb. 1949. Fincham. L.. J.. Persson. 336. and Pritchard. 67. Giovannucci. Are fibre supplements really necessary in diverticular disease of the colon? A controlled clinical trial.. Pathol. 1963. E.. J.. Am. 1. M... Jaskiewicz. J.. A.. Br. 1972. Falk Symposium 32. and Miller. and Cox. C. M. V. W. J. 29.. Trichopoulos. Smith. Aldoori. Littlewood. J. R. E. M. 1.. Nutr. 15. 67. 1973.. E... C. B. Gut. Br. J.6_fm Page 434 Sunday. Med. 1969.. Hyland. Kritchevsky. Condon. R. Br. M. 361. 157. 137. J. Br. and Uden. Scand. Acta. 1977. Exp. N. A. 31. 1979. A. S. I. Gastroenterology. Havia.. 19.. 16. L. 282. 1981. 38. 1353. 22. 1982. E. An association between maternal diet and colonic diverticulosis in an animal model. E. Surg. F. 12. Acta. 35. The influence of diet and dimethylhydrazine on the small and large intestine of vervet monkeys. Effect of dietary fiber on intraluminal pressure and myoelectrical activity of the left colon in monkeys. Smith. and Colebourne. V. Srivastava. M.. R.. and lactulose on colon function in diverticular disease.. 63. Rimm. Eds. 20. M. and Garry. 1. M. J. Rimm. 27. Eastwood. W. 137. A. F. 26.. Comparison of bran. A. J. Ahlberg. 23. A. Chir. Diverticulosis of the colon. 70. and and Willett.. 1980. Relationship of diet to diverticulosis of the colon in rats.. van Rensburg. M. 701. J. Z. H. A...... 3RD EDITION 14. W. E.. M. N. Sampson. 1978. C. Influence of symptoms and transit time of Vi-Siblin in diverticular disease. M. S. W. J. Lubbock. Scand. J. R. Sterculia bulk-forming agent with smooth muscle relaxant versus bran in diverticular disease. Clin. Wing. A. 32. J. E. H. B. M.. S. 17. Gut. W.. 24.... 1. R. G. 1980. Med. Carlson. Gut. A. L.. Brodribb. J. A. E. W. 67. 424... Chir. E.. Dietary management of diverticular disease. H. 77. 1144. May 6. J. Br.. Assoc. Post-mortem survey of diverticular disease of the colon. Hughes. Does a high fibre diet prevent the complication of diverticular disease?. S. Cowles. Kohler. J. 527. and Willett. Dietet.. R.. A prospective study of dietary fiber types and symptomatic diverticular disease in men. 7. May 6.2387_ch5. but significantly.5 While the median duration of symptoms in the CD patients was only 15 months.2 0-8493-2387-8/01/$0. a consistent dietary difference between patients with CD and controls is the high refined-carbohydrate intake observed in CD.3 g/d). 2001 7:41 PM CHAPTER 5. from 1 to 92 months. The low incidence of Crohn’s disease in less-industrialized third-world countries and its apparently increasing incidence in more industrialized Western countries have led to speculation that dietary changes in the Western world that have developed in the past few decades may be partly responsible. In common usage.3 FIBER STUDIES IN CD Fiber consumption in CD has also been examined in some studies (Table 5. more fiber than control subjects (26.50 © 2001 by CRC Press LLC 435 . Of interest was the finding that the CD patients consumed only about 25% of the raw fruit and vegetable fiber as the controls (0. Freeman Inflammatory bowel disease (IBD) refers to that group of conditions in which inflammation involves the small or large intestine or both. IBD is restricted to those conditions whose etiology is unknown and generally includes Crohn’s disease (CD) and ulcerative colitis (UC). have been proposed to play an important role in pathogenesis. dietary factors.6 vs. This increase was largely in the form of a significantly increased consumption of non-cellulose polysaccharide (17. For example.7 Fiber and Inflammatory Bowel Diseases (Ulcerative Colitis And Crohn’s Disease) Hugh J.2. 2. the range was wide. Thornton and co-workers reported that pre-illness dietary fiber intake of 30 patients with CD were significantly less than that of 30 controls. 14.3 g). Kasper and Sommer employed an experienced dietitian to perform dietary histories in CD patients and controls over 7 successive days.00+$1. a finding which was first reported in 19761 and which has subsequently been confirmed.7_fm Page 435 Sunday.6 vs. As with many other intestinal conditions of undetermined or uncertain etiology. this confirmed the wellappreciated phenomenon that the validity of recall in some patients with longstanding symptoms may not be precise.4 They reported that patients with CD consumed slightly.5 g). In contrast.3 vs. A further study by Mayberry and co-workers found no difference between the dietary fiber intakes of patients with CD compared with normal controls.1). including fiber. 22. nutritional status. While the study reports higher refined carbohydrate intake in controls compared to CD patients on the diet.. hospitalizations. The CD patients on the modified high-fiber. 2001 7:41 PM 436 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.6 reported the effects of treating 32 CD patients with a fiber-rich diet for a period of 4 years and 4 months. but the significance of the difference in the total fiber intake (13 vs.3 (0.8 in which 190 patients with CD received the high fiber “Bristol diet” while 162 received the “low-fiber” diet. The number of portions of fiber consumed by those on the “normalized” diet was significantly higher than the number taken by those on the “usual” diet. led to several further studies (Table 5. Patients in the “high-fiber” group also restricted their intake of refined carbohydrate.2b) 19.1 No. Raw fruit and vegetable fiber. or postoperative recurrence.2). Significantly more patients in the high-fiber group withdrew from the trial for reasons other than disease deterioration. complications. the statistical significance of this was not defined.2387_ch5. Patients Fiber Intake in Crohn’s Disease Disease Duration 35 Crohn’s 70 Control 1 year (av.7 Thirty patients with CD placed on a “normalized” diet were compared to 28 patients given the “usual” low-residue diet prescribed in Italy.3 (14. 39 90 11 (111 d) 18 (533 d) 1 5 6 4 5 7 7 14 8 9 92 18 21 Low refined carbohydrate. 3RD EDITION Table 5.7.5a) 17. especially raw fruit and vegetable fiber. No differences were found in clinical outcome over the two-year study period.7 Daily Sugar (g) Patient Admissions Operations Ref.2 (2. randomized 30 Normalized 28 Low-residue 190 High-fiber 162 Low-fiber a Study Duration Daily Fiber (g) 52 mo 33. 5). May 6. The apparent success of this “Bristol diet” led to a prospective study by Levenstein et al.4 29 mo 13a 3 2 yr 27. a population-based case-control study on dietary habits of patients with inflammatory bowel disease in Stockholm was reported.3b) 4 5 14 20 2 Non-cellulose polysaccharides. high vegetable. and fruit fiber (“Bristol diet”). operations. 3 g) was not stated.7.7 ) 22. no figures for dietary fiber intake in controls were given. there were no differences in outcome with respect to symptoms. Table 5. More recently. while those in the “low-fiber” group were encouraged to eat refined carbohydrate. a 26.2 Fiber Trials in Crohn’s Disease No. The observations by Thornton et al.6 (17.7.9 Retrospective dietary habits over a five-year period .7_fm Page 436 Sunday. Heaton et al. CD patients who were not provided with dietary instruction served as retrospective controls. low-sugar diet had significantly fewer hospital admissions and total days in hospital compared to the control patients.9a 15.) 30 Crohn’s 30 Control 5 mo (range: 1–92) 16 Crohn’s 16 Control Not stated a b Fiber Intake (g/day) Pre-Illness Current Ref. Over a period of 29 months.5 that the pre-illness diet of Crohn’s patients may have been low in fiber. While the number of operations in those patients on the special diet was fewer (1 vs. A second prospective randomized controlled study was reported by Ritchie et al. Patients Retrospective controls 32 Fiber 32 Control Prospective. 5 yr Daily Fiber (g) Pre-Illness Trial 19.3 Fiber in Ulcerative Colitis No. Table 5. possibly.15 In a study from Seattle. SHORT-CHAIN FATTY ACIDS IN IBD While only limited data are available to support a prominent role for fiber in the treatment of IBD. (rate. and vegetables. No differences were detected in refined carbohydrate or fiber intakes between these UC patients and 30 control subjects. The cumulative relapse rates of the sulfasalazine group and the fiber group were 20 and 70%. H2. The relapse rate on the highfiber diet was similar to that expected in UC patients treated with placebo.7. respectively. randomized 15 Sulfasalazine 20 High-fiber a Disease Duration 2 mo 8. CH4) and SCFAs (predominately butyrate. whole wheat bread.3). over 6 months. no definite effects were documented on pouch function with either pectin or methyl cellulose. the effects of different forms of fiber have been evaluated in UC patients that have been surgically treated with an ileal pouch–anal anastomosis. in the colon may be very relevant in disease pathogenesis and. and a supplement of 25 g bran supplied as Kellogg’s All Bran or Allinson’s Bran Plus. FIBER STUDIES IN UC Only limited studies have examined the effects of dietary fiber in UC (Table 5. short-chain fatty acids (SCFAs).7.7_fm Page 437 Sunday.18 The SCFAs which are produced have been .13 reported that ispaghula husk may relieve gastrointestinal symptoms in patients with UC.9 18.10 observed that the mean daily intake of fiber was significantly lower in Crohn’s disease patients than in controls from the Netherlands. i. Geerling et al. %) 97 96 13a 3 Ref. Patients Retrospective controls 30 UC 30 Control Prospective. Later. The increased fiber was taken in the form of whole wheat bread. while 20 of the 24 patients who tolerated a highfiber diet were continued on that diet with sulfasalazine being stopped. anaerobic bacteria metabolize a varying amount to gases (CO2. Finally.e. The relative risk of Crohn’s disease was decreased in those consuming a high intake of fiber (15 g or more per day).3 Daily Sugar (g) Relapse No. Several studies in ileostomy subjects have shown that about 90 to 100% of orally admininstered fiber is recoverable in the ileostomy effluent. being equivalent to mesalamine. a commonly prescribed anti-inflammatory pharmacologic agent. renewed interest has resulted from the recognition that important metabolic by-products of fiber. vegetables.17 When the fiber reaches the colon. Hallert et al.11 Fifteen patients continued on sulfasalazine and their regular diet. Davies and Rhodes divided 39 patients with UC in remission on sulfasalazine into two groups. propionate. May 6. and acetate). In a recent randomized clinical trial of dietary fiber (administered in the form of Plantago ovata seeds). 2001 7:41 PM FIBER AND INFLAMMATORY BOWEL DISEASES (ULCERATIVE COLITIS AND CROHN’S DISEASE) 437 were evaluated in 152 patients with Crohn’s disease and compared to 145 ulcerative colitis patients and 305 controls.. Later.16. A second study by Thornton et al.12 examined the “pre-illness” diet of 30 patients with UC diagnosed within the previous 3 months. therapy. 12 3 (15%) 15 (75%) 11 Fiber added as bran cereal.2387_ch5. a Spanish group14 described its effectiveness in maintaining remission in UC. M. Ritchie. C. J. Treatment of Crohn’s disease with an unrefinedcarbohydrate. Controlled multicentre therapeutic trial of an unrefined carbohydrate.. Ahlbom. 2.21 Diversion of the fecal stream as occurs. 295. Thornton. S.. 999. G. Wochenschr. E.19 In addition. 762. Br. and Brummer. 367. H. J. following colostomy. 2. would increase the osmotic load and provide a mechanism for diarrhea in IBD. 323.. K. J. SCFAs stimulate colonic sodium and water absorption. Metabolism of fiber entering the colon to SCFAs and lactic acid.. Dietary fiber and nutrient intake in Crohn’s disease.. Br.. M. 293. in whom fecal concentrations were similar to those of normal controls. 11. R. ulcerative colitis or irritable bowel syndrome. e. Heaton.. . Increased consumption of refined carbohydrates in patients with Crohn’s disease.24 have shown that fecal lactic acid concentrations are increased in both Crohn’s colitis and UC. 6.. Vernia et al. Gut. In addition to a possible role for fiber in the generation of SCFAs in various colonic disorders. and Heaton.. and Brandes. controlled trial will need to be conducted.. P. 5. Thornton. Geerling. Luzi. Br. 2001 7:41 PM 438 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Sci. J. A. 1978. Nine were judged to be much improved. J. 1976. 1980.. Med. Twelve patients were treated with SCFA rectal irrigation over a period of 6 weeks in a non-blinded fashion.. W. totally undigested fiber probably has minimal effects. 47. 764. Thornton. Wadsworth... and Allan. Maintenance of remission in ulcerative colitis with sulfasalazine or a high-fibre diet: a clinical trial. Diet and ulcerative colitis. Jarnmark.. 1983. Levenstein.. H. Diet and Crohn’s disease: characteristics of the pre-illness diet. Persson. J.. E. C. Martini. 54.. If SCFAs are found to be better than placebos. however. but that fecal weights correlate with lactic acid concentrations only in UC. J. R.. Dig. since fiber has very little osmotic activity. Badart-Smook. Mayberry. P. A. K. and Rhodes. Emmett. 1998. J. G.20.. J.. A. 26.27 to perform rectal irrigation of SCFAs in patients with distal UC. Of course. G. 1524. Rhodes. Lennard-Jones. as noted above. the importance of these substances in UC will be confirmed. R. 1979. Emmett. and their stimulation of sodium and water absorption. may result in an inflamed distal excluded segment. called “diversion colitis. W. and Nilsson. 7. unabsorbed complex carbohydrates per se have been hypothesized to play a part in the diarrhea pathogenesis in IBD. G. 1.. W.... Br. Kasper. 9. Scand. The abnormalities in SCFAs in UC led Breuer et al. A. their use as a colonic fuel. 919. 3RD EDITION shown to be important sources of energy for colonocytes..” Harig and co-workers22 have reported improvement in diversion colitis using rectal SCFA irrigations. S. 1981. J. 1987. and Heaton. J. P. 12. J. 444. May 6. J. Nutr. M. 1985. Consumption of refined sugar by patients with Crohn’s disease. REFERENCES 1. and Sommer. However. and Emmett. Digestion. Diet and inflammatory bowel disease: a case-control study. J.. SCFA production may not lead to diarrhea due to their rapid absorption by the colon. P.. Low residue or normal diet in Crohn’s disease: a prospective controlled study in Italian patients. and Rogers. Two studies of current and previous habits in newly diagnosed patients.. R. Clin. Med. Med J. Diet in Crohn’s disease. 1979. K.. Med. W. Epidemiology. Davies. 67. 280. R. Stockbrugger.. other investigators have found higher SCFA levels in severe UC when compared with normal controls... K. randomized.25 Indeed.. Comprehensive nutritional status in patients with long-standing Crohn disease in remission. 989.. 10. J.. 20. 2. Dis.. Med. 18.. Prantera. Holtug and coworkers have reported that some of the changes in SCFA pattern in UC may be due to bacterial fermentation of blood. J.. J. F. W. Klin. K.. Fecal SCFA concentrations were found to be much lower in patients with UC than in those with Crohn’s colitis..2387_ch5. Am.7_fm Page 438 Sunday. Br. B. 517. a blinded. 3. 4. 26.g. 3. Gastroenterol. fibre-rich diet. Clearly. and D’Ubaldi... Janerot. 8. P. J. I.26 The reasons for the discrepancy in these studies are not clear but may relate to differences in methods or severity of disease. and Hellers. fibre-rich diet in Crohn’s disease. 1992.23. 1979. R.. . K. E. Sanchez-Lombrana. 45.2387_ch5. V. Med. M. 1988... Gastroenterol.. 2001 7:41 PM FIBER AND INFLAMMATORY BOWEL DISEASES (ULCERATIVE COLITIS AND CROHN’S DISEASE) 439 13. H.. Wood. 78. Hinojosa.. G. 18... 1982. 95. Dig.. J. Gastroenterology. E. J.. 1982. M. Harig. Pathol. P. C. H. 1988. I. Br... Engl. Heyworth. Roediger. 19.. J. A.. Am. F.. Pectin and methyl cellulose do not affect intestinal function in patients after ileal pouch-anal anastomosis. W. L. K. Surg.. 23. Gastroenterol. and Truelove. Fecal lactate and ulcerative colitis. Holtug. P. Roediger. Riera. Gastroenterol. S. J.. M. and Christ. 22.. and Kelly. S.. Magliocca. R. 48. 1986. Short chain fatty acids in the human colon. J. Soergel.. 24. Moles. J. R. Digestion of the carbohydrates of banana (Musa paradisiacal sapientum) in the human small intestine. 94.. 26. 44. Kaldma. Nutr. H. Sci. Barbetti. Gomollon.. Garcia-Puges. Englyst. Gonzalez-Huix. Rectal irrigation with short chain fatty acids for distal ulcerative colitis. 1982. W. 423. Dig. M. Am. J. . 35. F.. Moore... and Schmitt. N. A... Invest. 320.7_fm Page 439 Sunday. C. and Cummings. Organic anions and the diarrhea of inflammatory bowel disease. H.... 23. 15. Dominguez-Abascal. 17. Gastroenterology. Ruppin. The effect of bacterial fermentation of blood.. Gonzalez-Lara. Digestion of polysaccharides of potato in the small intestine of man. 21. Luminal ions and short chain fatty acids as markers of functional activity of the mucosa in ulcerative colitis. and Cummings. J. M. 1991.. 763. L. 1999. 1564. 1987. 69.. and Cittadini. R. A.. Cummings.. Scand. and Gassull. Scand. Gastroenterology. Sci.. Spanish Group for the Study of Crohn’s disease and ulcerative colitis. W.. Latella. Piris.. D. 33..M.. and Petersson. N. and Rae. J. G. I.. A. W. 1353. 27... Vernia. M. Sorgel. Martinez-Salmeron. 1989. J. Am.. 427. Clin. E. and Breuer. W. and Woods.. 424.. A. F... 1980. Gut. Utilization of nutrients by isolated epithelial cells of the rat colon. J. 42. Nutr. Hallert. Trophic effect of short chain fatty acids on mucosal handling of ions by the defunctioned colon. 83. Komorowski. Randomized clinical trial of Plantago ovata seeds (dietary fiber) as compared with mesalamine in maintaining remission in ulcerative colitis. Am. 424. M... J.. E. M. J. 323. and Mortensen. 23.. R.. H. Breuer. M. Clin. H.. H.. S. P. Treatment of diversion colitis with short chain fatty acid irrigation. J.. 1988. C. A. J. R. 1981.. J.. J. J. B.. K.. 14. Gine. Clin. Dis. 22. C. H. J. 667. Clin. G. Roediger. Gnaedinger.. Absorption of short-chain fatty acids by the colon. 16. F. Thirlby. K. 92. Hauck. F. Willoughby. H. 747. N. 99. J.. Bar-Meir. B. Vernia. J. Lab. Englyst. P. 1991. Rasmussen. 185.. Ispaghula husk may relieve gastrointestinal symptoms in ulcerative colitis in remission. Fernandez-Banares. W. Buto. 20. W. May 6. 25. Navarro. Caprilli. Short chain fatty acids in inflammatory bowel disease.. R. C.. Dis. 26. 36. F. 1997. S.. F. 2387_ch5. 2001 7:41 PM . May 6.7_fm Page 440 Sunday. 1 Dietary fiber (DF) intake by the Japanese people since 1930. 2001 3:08 PM CHAPTER 5.8_fm Page 441 Tuesday.8.70%. May 8.00+$1.1 DF content values are rice. 1.02%.2 wheat.2387_ch5. Food intake values are from Food Balance Tables of Japan.8 Disease Patterns in Japan and Changes in Dietary Fiber (1930–1980) Keisuke Tsuji and Bunpei Mori Figure 5. 1.3 and barley. where DF intake (grams per capita per day) = food intake × DF content.4 0-8493-2387-8/01/$0.86%.50 © 2001 by CRC Press LLC 441 .18 to 2. 4. 7.2 Dietary fiber (DF) intake by the Japanese people from potatoes.2 seaweeds. 2001 3:08 PM 442 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 80%. 0.6%.54%. pulses. vegetables.77%.5 potatoes. 1.96%.01%. Food intake values are from Food Balance Tables of Japan.2 total fruits.1 DF content values are sweet potatoes. 0.2 and konjac flour.8_fm Page 442 Tuesday.5 total vegetables. 23. 1.14%.2387_ch5. May 8. 3RD EDITION Figure 5. and seaweeds since 1930 where DF intake (grams per capita per day) = food intake × DF content.6 .8.5 pulses as soybeans. fruit. 8.8_fm Page 443 Tuesday. May 8.7 443 .2387_ch5.3 Changes of death rates from adult disease in Japan. 2001 3:08 PM DISEASE PATTERNS IN JAPAN AND CHANGES IN DIETARY FIBER (1930–1980) Figure 5. Nutr. 35. H.. .. T. J. et al. and Southgate. 15. Jpn. and Takehitsa. 1930−1980. Ayano. 1982.. 6.. 2. McCance and Widdowson’s The Composition of Foods. Japan Patient Survey (1955−1982). Int.2387_ch5. 8. 1982.. 1978..8_fm Page 444 Tuesday. 159. 1980. London. 3. Japan Vital Statistics (1900−1980). May 8.4 Number of patients in Japan suffering from adult diseases8 and from gallstones. unpublished data. Med. 26. A. F. Food Balance Tables of Japan.9 REFERENCES 1. Kameta. Chori Kagaku. A. D. Rep. 133. 16. 5. 4.. 1982. 3RD EDITION Figure 5. Insoluble dietary fiber contents of foodstuffs. Paul.. Y. 7. B. Hoshi. 2001 3:08 PM 444 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Mori. Eiyo to Shokuryo..8. K. 9. et al. 4th ed. 2924. Contents of dietary fiber in some Japanese foods and the amount ingested through Japanese meals. Tsuji. 24. A. Her Majesty’s Stationary Office. S. 9 Dietary Fiber Modification of Toxinor Carcinogen-Induced Effects on Intestinal and Mammary Tissues* Hugh J. The long-term effects of low fiber compared to high fiber intakes on the toxic effects of food components. more specifically. After the hypothesis that colon cancer may be less common in populations consuming a high-fiber diet was proposed (see Chapter 5. Differences in research protocols. It is interesting to note that some dietary fibers or dietary fiber-rich foods are protective for some toxic substances but not for others. The interaction between dietary fiber polymers and other components of the diet means that the composition of the entire diet is a key factor in carcinogenesis and toxicity studies. * This chapter is an updated version of Chapter 7. Other examples of the specificity of the antitoxic effect of high-fiber substances are given in Tables 5. This complex function of the fiber polymers. Many of the early studies were done by Erschoff. the possible protective role of dietary fiber in carcinogenesis has been the subject of many studies. which is hypocholesterolemic but does not prevent cyclamate toxicity in rats when cyclamate is fed at the 5% level.2. These studies emphasize that the effect of dietary fiber depends on the composition of its fiber components. food contaminants. Spiller The possible antitoxic effect of dietary fiber or fiber-rich foods and. and.5). and Erschoff showed that other beneficial effects such as the hypocholesterolemic effect of some fibers is often not connected to their antitoxic effects.9. specifically those that might be induced by carcinogens.00+$1. 2001 7:45 PM CHAPTER 5.2 focuses on more recent studies on carcinogenesis and the possible protective effect of dietary fiber polymers or high-fiber foods. 0-8493-2387-8/01/$0. the number of laboratory animal model studies on chemically induced carcinogenesis in the presence of various types of dietary fiber increased rapidly.9.1 summarizes some of the studies of protection against toxicity in general and some early carcinogenesis studies. diet composition. their limitation is the extrapolation to humans from animal studies. A problem instrinsic to these animal studies is that the same toxicity experiments cannot be done in humans. As a result. as other non-fiber components of food might be responsible for the antitoxic or cancer-preventing effect. environmental toxic substances. Table 5.8 An example is locust bean gum.11 who demonstrated that otherwise-toxic levels of various substances in foods became either less toxic or non-toxic if the subject was fed with foods high in dietary fiber or some of the fiber polymers.2387_ch5. can have major implications on the etiology and/or pathogenesis of some diseases. in general. or high-fiber foods. and many other factors are certainly responsible for the differences in some results obtained. Reddy and Gene A.50 © 2001 by CRC Press LLC 445 .9. Caution is required in the interpretation of results with high-fiber foods.4.9. is an important one. Freeman and Gene A. Spiller.9_fm Page 445 Sunday. by Bandaru S. May 6. Modification by Dietary Fiber of Toxic or Carcinogenic Effects.10.9. Table 5.1 and 5. in the second edition of the Handbook.1. starch.5-Di-t-butylhydroquinone Chlorazanil hydrochloride Sodium cyclamate Tween 60 FD & C Red #2 Calcium (CdCl2) Tween 60 2-Acetylaminofluorene Dietary Fiber Source Amount in Diet (%) Soybean meal Alfalfa meal Alfalfa meal Rye grass Wheat grass Fescue grass Orchard grass Purified cellulose Alfalfa meal Stock diet b vs. without specific studies on tumorigenesis.). 2001 7:45 PM 446 Table 5. Amount corresponding to the alfalfa meal fed. Rats Rats Rats 3 4 5 Rats + + + + + + + + – + Rats + 7 Rats – – + + + + + + + + + – + + + + 8 Rats Rats Rats 10 10 10 5 5 Mice 10 10 Rats + + + + + – – – + 6 8 9 10 11 12 Note: This table summarizes only selected results of the cited studies. May 6. many other high-fiber substances and polymers were tested. alfalfa.9_fm Page 446 Sunday.5 or 5 10 10 10 10 10 10 10 Animal Effecta Ref. The term “stock diet” refers to diets based on natural foods (grains. In many of these studies.2387_ch5. . purified diet Alfalfa 15 20 10 10 10 10 10 10 20 Locus bean gum Purified cellulose Psyllium seed powder Alfalfa meal Carrot root powder Gum karaya Sodium alginate Psyllium seed Alfalfa meal Rice straw Carrot root powder Purified cellulose Alfalfa meal Watercress powder Parsley powder Increased fiberc but differing diets Alfalfa meal Wheat grass meal Rye grass meal Sodium alginate Agar Alfalfa juiceb Locust bean gum Apple powder Stock dietsc vs. etc. purified diets 10 10 10 20 20 10 15 2.1 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.9. and glucose. a b c “+” indicates a general protective effect (such as weight maintenance or survival) when compared to controls on fiber-free or lower-fiber diets. 3RD EDITION Studies on the Antitoxic Effect of Dietary Fiber or High-Fiber Foods and Early Carcinogenesis Studies in Experimental Animals Toxic Substance Tween 60 Glucoascorbic acid (4%) Tween 20 Tween 60 Span 20 2. These diets were usually compared to diets based on purified ingredients such as casein. “–” indicates no protective effect. 2-DMH 1.2-DMH 1.5 9 6. SD rats SD rats SD rats Fischer rats SD rats Fischer rats Fischer rats Fischer rats SD rats SD rats SD rats Fischer rats SD rats SD rats Balb/c mice Fischer rats Fischer rats Fischer rats Fischer rats Fischer rats Balb/c mice Fischer rats Fischer rats Fischer rats Balb/c mice SD rats SD rats Fischer rats Fischer rats SD rats Fischer rats SD rats Fischer rats Wistar rats Wistar rats SD rats Wistar rats Wistar rats Wistar rats Wistar rats Wistar rats Wistar rats Wistar rats Wistar rats Fischer rats SD rats SD rats –– –– –– +++ +++ +++ +++ +++ +++ +++ +++ +++ –– –– –– –– +++ +++ +++ – – – – – – +++ +++ – – +++ +++ – +++ – – – – – +++ +++ +++ +++ +++ +++ +++ +++ +++ 26 26 27 28 13 14 15 16 17 17 18 19 20b 18c 21d 14e 40f 15 16 22 21d 19 19 19 21d 17 17 14 14c 33 34a 20b 14 23 23 20b 35 35 36 24 23 23 35 35 25 17 17 Colon Studies 1.2-DMH 1.2-DMH 1.2-DMH AOM 1.2-DMH 1.2-DMH AOM 1.5 9 4.2-DMH 1.2387_ch5.2-DMH 1.2-DMH 1.2-DMH 1.5 4. 2001 7:45 PM DIETARY FIBER MODIFICATION OF TOXIN.2-DMH 1.2-DMH 1.2-DMH MNU AOM AOM 3.2-DMH 1.9.2-DMH 1.5 4.2-DMH 1.2 447 Studies on the Protective Effect of Dietary Fiber or High-Fiber Foods in Chemical Carcinogenesis Studies Carcinogen Dietary Fiber Source (%) Amount in Diet (%) Animal Effecta Ref.2-DM-4-ABP 1.2-DMH 1.2-DM-4-ABP AOM AOM 1.2-DMH 1.5 9 40 20 30 .2-DM-4-ABP 3.2-DMH 1.2-DMH 1.5 9 4.2-DMH 1.5 4.2-DMH AOM 1.2-DMH AOM AOM AOM MNU 1.OR CARCINOGEN-INDUCED EFFECTS Table 5.2-DMH AOM AOM 3.2-DMH AOM AOM AOM Oat bran Guar gum Guar gum Phytic acid Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Wheat bran Citrus fiber Citrus fiber Corn bran Corn bran Corn bran Rice bran Soybean bran Soybean bran Alfalfa Alfalfa Alfalfa Alfalfa Barley fiber Coffee fiber Carrot fiber Pectin Pectin Pectin Pectin Pectin Pectin Hemicellulose Cellulose Cellulose Cellulose Cellulose Cellulose Cellulose Cellulose Cellulose 20 10 5 1 20 15 15 15 20 30 20 20 20 20 20 15 15 15 15 20 20 20 20 20 20 30 15 15 10 20 15 4. May 6.2-DMH 1.9_fm Page 447 Sunday. 3. with associated reduction in nitric oxide synthase and total cyclooxygenase activities (COX-1. H. 1. 1960.. J. 27... H. 1977. 2001 7:45 PM 448 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Biol. 1975. J. Nutr. T. W. Occup.. 857. Nutr. f Study showed lipid portion of wheat bran critical for protective effect.2-DMH 1.. 1957. Ershoff. 4. Comparative effects of a purified and stock diet on DBH (2. c Effect of wheat bran on stage of initiation was studied. Biol. 1974. Exp. Ershoff. 36. . Fischer rats Fischer rats SD rats SD rats SD rats SD rats SD rats – +++ +++ – +++ +++ +++ 25 22 37 37 38 38 39 +++ 26 Mammary Studies 10 Fischer rats Note: Listed studies refer to chemically induced carcinogenesis. Proc. 656. Ershoff.. 49. Fed. May 6.2387_ch5.. Med.. H. and Thurstun. Ind. during. Am. 1950. Clin.41 a “+++” indicates a protective effect against tumorigenesis in the cited study (compared to fiber-free or lower-fiber diets). Nutr. Med. Wheat bran was fed to rats during carcinogen treatment only. and Marshall. b Experimental fiber diets were fed to animals 3 days before. 1. Beneficial effect of alfalfa meal on chlorazanil hydrochloride toxicity in the rat. H. F.. C.. d Animals fed the control diet or fiber-free diet had very low colon tumor incidence. Ershoff. 10. B. 1395. J.2-DMH Cellulose Lignin Cellulose Lignin Cellulose Cellulose Cellulose MNU Wheat bran Amount in Diet (%) 20 7. e MNU. Effects of diet on amaranth (FD & C Red No. 6. B.. Proc. Fed. Med. 70. Burnett. Other abbreviations used for indirect-acting carcinogens include 1.. H. R. H.2-DMH 1. J.. Importance of diet in studies of chronic toxicity.5 5 15 10 Animal Effecta Ref. Am. H.2) toxicity in the rat. AOM.. 1959. Food Sci. 17. 73. Arch. and 14 days after carcinogen treatment. B. Ershoff. Soc. J. Soc. methylnitrosourea is a direct-acting carcinogen. Ling. 40.. 104. Hyg. Soc. Chow. E. M.. 563.2-DMH 1. 7. 937. 484.5-di-t-butylhydroquinone). J.2 (Continued) Studies on the Protective Effect of Dietary Fiber or High-Fiber Foods in Chemical Carcinogenesis Studies Carcinogen Dietary Fiber Source (%) AOM 3. Biol.2-dimethyl-4-amino-biphenyl. “– –” indicates enhancing effect in the cited study. 95.2-dimethylhydrazine. but genetic model of intestinal polyposis (Apc knockout mice) shows protective role of high-fiber-containing diet for small and large intestinal neoplasia.2-DMH 1. Wilson. L.2-DM-4-ABP. 9. azoxymethane. W. Exp.. Ershoff. Kritchevsky. H.. 357. 1953. Beneficial effect of alfalfa and other succulent plants on glucoascorbic acid toxicity in the rat. Beneficial effect of alfalfa meal and other bulk-containing or bulk-forming materials on the toxicity of nonionic surface-active agents in the rat. B.2-DM-4-ABP 1. F. COX-2).9_fm Page 448 Sunday. 2. Antitoxic effects of plant fibers. 8. 3.. Exp. and De Eds. E. 141. 204. 3. Ershoff. “–” indicates no protective effect in the cited study. toxicity in the rat. Med.. They were transferred to fiber-free diet until termination of the experiment. 1972. 3RD EDITION Table 5. B. Surg. and Barrows. Effect of basal diet on the response of rats to certain dietary non-ionic surface-active agents. Modification by fiber of toxic dietary effects. Exp. 5. D. Protective effect of dietary fiber in rats fed toxic doses of sodium cyclamate and polyoxyethelene sorbitan monostearate (Tween 60)... B. REFERENCES 1. Nutr..9. 1692.2-DMH. 1974. B. B. The animals were then transferred to standard rat pellets and fed this diet until termination of the experiment. Proc. Clin. Reddy.2-dimethylhydrazine administration. and Royle. Pilch. E. 30. 1727. M. and Hernandez H. S. L. 4057. B. 1986... M.. and Shock T. D. 1983.. and Campbell. 1952. G.. E. A... 1980. M. 29. and soybean bran on 1. J.2-dimethylhydrazine-induced rat colonic neoplasia. Scott. G. and Copeland. M.. 16.. Nutr. A. H. Cancer Inst. 141. and colon carcinogenesis in 1.2-dimethylhydrazine-induced large bowel tumorigenesis. H. L.. 28. Natl. Effect of dietary fiber on azoxymethane-induced intestinal carcinogenesis in rats. 176. cell proliferation. Reddy. Nutr.. pectin. B. D. P.. 46. Y. Effect of dietary fiber on induction of colorectal tumors and fecal B-glucuronidase activity in the rat. Maeura. A double-blind study on the effects of purified cellulose dietary fiber on 1.. 71. 20. Dahlquist. D. M... D.. 25. S. S.2-dimethyl-4-aminobiphenyl-induced intestinal carcinogenesis in male F344 rats. and Rose... R. B. H. Spiller. Cancer.. Carcinogenesis. Nyman. Dietary fiber and colon carcinogenesis: a critical review. G. Y... The potential of an insoluble dietary fiber-rich source from barley to protect from DMH-induced intestinal tumors in rats. E.. Cancer Res. Bethesda. and Fredlund. Cancer Inst. R. B. and Kritchevsky... 21. 13. G. 1981. and Lupton.2-dimethylhydrazine initiation of colon tumors and on fecal B-glucuronidase activity in the rat. Relationship between colonic luminal pH. and Kritchevsky. J. Cancer Inst. Cancer. Natl. P. and Nicolais. S. H. Asp. R. Modulation of N-nitrosomethylurea-induced mammary tumor promotion by dietary fiber and fat. 1959. Am. J.. Natl. Effects of wheat... 31. 2001 7:45 PM DIETARY FIBER MODIFICATION OF TOXIN. W.. 2661.. Y. P. and Wayman. 1990..V. 1983. Freeman.. S. 40. Protective action of stock diets against the cancer-inducing action of 2-acetylaminofluorene in rats. J. D. M. and Oste.. J. J. Clapp. N. A double-blind study on the effects of differing purified cellulose and pectin fiber diets on 1. Cancer Res. May 6. S. Cancer Res. Fredlund. E.. Asp. Plenum Press. Wilson. Watanabe. 66. Meschter. R. Bauer. Cancer Res. P. Freeman. H. M.. R. 1983. B..2-dimethyl4-aminobiphenyl-induced intestinal carcinogenesis in F344 rats.. Nigro.. 1979. 26.. G. A. Bauer. Cancer Res.. 41. 2219. soybean and wheat bran. rice. 39.. 213. 1978. Spiller.or methynitrosourea-induced colon carcinogenesis in F344 rats.2-dimethylhydrazine-induced rat colonic neoplasia. R.. 1991. Cancr Inst.. 5. Effect of dietary corn bran and autohydrolyzed lignin on 3. H. 1977. Enhancement of rat colon carcinogenesis by wheat bran consumption during the stage of 1.. 1980. Effect of two kinds of pectin and guar gum on 1. Klopfer.. Cancer Inst. N.. G.. and Berry. in Medical Aspects of Dietary Fiber. Vahouny. H. . L. Spiller. Enhancement of 1. Dahlquist.. Kendall. L. S.. Ullah. and Kim. Bull. Freeman. Jorensen. 24. 1973. A... Jacobs. L. 23. Jacobs. N. Cancer Inst.. Experimental animal studies in colon carcinogenesis and dietary fiber. Cancer Res.. B. 83. G. Dimethylhydrazine-induced colon tumors in rats fed diets containing beef fat or corn oil with and without wheat bran. D. MD.. Oste.9_fm Page 449 Sunday... 1979. Federation of American Societies for Experimental Biology. M. 1984. K. A. 12.. R. Reddy. Effect of dietary wheat bran and dehydrated citrus fiber on azoxymethane-induced intestinal carcinogenesis in Fischer F344 rats. N. K. McIntosh. K. R. in Dietary Fiber in Health and Disease.. Cellulose dietary bulk and azoxymethane-induced intestinal cancer. 19. and wheat bran on azoxymethane. N. 77. J. 1097. 12. Eds.. B. 2912. P. 11. 2518.. S.. 265. 30. A.. and Weisburger. 21. 1993. 43. S. S. A. 14. New York. F. Cancer Res.. Dose-dependent inhibition of large intestinal cancer by inositol hexaphospate in F344 rats. and Mori. Natl. 1987. J. Yamamoto. Clapp. A. C. 553. W.. R. B. Reddy.. S. 62. L. 17. G. 22. 83.2387_ch5.. A.. 2. J. 172. Engel. Beneficial effects of alfalfa meal and other bulk-containing or bulk-forming materials on symptoms of Tween 60 toxicity in the immature mouse. 19. 419. Weisburger. 713.. H. H.. L..2-dimethylhydrazineinduced large bowel tumorigenesis in Balb/c mice by corn. 6. Life Sciences Research Office. M. and Wideman. 18.2-dimethylhydrazine treated rats fed high fiber diets. Reddy. D.OR CARCINOGEN-INDUCED EFFECTS 449 11. 27.. J. 1982.. 1. J. Barnes. S.. Cohen.. A. A. 496. and Kim. 33. Zang. 1979. 69. J. J. D. 1981. Henke. J.. Physiological Effects and Health Consequences of Dietary Fiber... J. Nutr. 15. H. Ershoff. B. Carcinogenesis. H. 51.. G. Pak.. 32. 3752. 38. Ward. Natl. and Shamuddin. New York. 63. 1981. 211. Cancer. corn. Effect of dietary alfalfa. Hutcheson. London... Natl. Effect of dietary wheat bran and dehydrated citrus fiber on 3. H. Eds. Nutr.. A. J.. Mori. J. Nutr. Plenum Press. and McPherson-Kay. .. . Nakaji.. D.. 5581. Sugawara. The effect of the fiber components cellulose and lignin on experimental colon neoplasia. D. 110. Richards.. 35. 5529. and Taketo. 40. 261.. Shivapurkar. Carcinogenesis. K. R. inulin and pectin. N. 1986. S.. H..2-dimethylhydrazine-iduced rat colon carcinogenesis. 2001 7:45 PM 450 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Carcinogenesis. M. 1989.. H. Alabaster. 52. Simi. G.. 60. 1998. Y. D. Freeman. 1984. Spiller. Reddy. M.. H. B.. I. O.. and Reddy. V. Suppression of intestinal polyp development by low-fat and high-fiber diet in Apc (delta716) knockout mice. 38. 1815.2-dimethylhydrazine-induced colon carcinogenesis in rats.2387_ch5. B. Gastroenterology. 49. Oshima. Murray. Sakamoto. Iwane..... 1996. Oshima... Prevention of colonic aberrant crypt foci and modulation of large bowel microbial activity by dietary coffee fiber.. M. S. D. Preventive potential of wheat bran fractions against experimental colon carcinogenesis: implications for human colon cancer prevention. Cohen. A. Freeman. Cooma. K. D. May 6. 41. L..9_fm Page 450 Sunday. Effect of high hemicellulose corn bran on 1. 36... 3RD EDITION 34. B. A. Cancer Res. Simi.. Effect of dietary cellulose on cell proliferation and progression of 1. K. Ord. 5. K.. J.. A. Effects of differing purified cellulose. and Cameron. L. 1997. J. C. Oncol. 4792. Hioki. B.. Hunter. 116. M. 19.. Surg. 46. and Brown. V. 37.2-dimethylhydrazine-induced colonic carcinogenesis in rats. 18.... Comparison of resistant starch with cellulose diet on 1. S. Carcinogenesis. 1993. Cancer Res. J. Chou. Hertman. Sloan. I. S.2dimethylhydrazine-induced rat intestinal neoplasia. 2000. W. and Munakata. Ku. and Kim. V. 77.. H. 39. pectin and hemicellulose fiber diets on fecal enzymes in 1. Fleiszer. A.. and Rao. Cancer Res.. E.. C. A... Hirose. J. S.. Y. 1863. G. Rao. A.. and Phytic Acid on Health . 2001 6:03 PM SECTION 6 Effect of Whole Grains. Cereal Fiber. May 6.2387_Section 6_fm Page 451 Sunday. 2001 6:03 PM . May 6.2387_Section 6_fm Page 452 Sunday. 10–13 Still. Cinnamic acids. In fact. flavonoids. whole grain products.21 Many are common among plants.1–6 Several decades of research have focused on identification of food components and mechanisms to explain the positive health effect.2387_ch6. Examples of a few grain antioxidant structures are shown in Figure 6. Lignans have been the subject of considerable study. there are many other antioxidant compounds that occur in lesser amounts in grains.and diol-esters of ferulic and caffeic acids and alkyl resorcinols. Avenanthramides are unique to oats. and other chronic diseases of aging is reduced by diets high in fruits.1. are abundant in most grains. where heart disease is relatively low for a wine-drinking population with high saturated fat intake. 2001 7:46 PM CHAPTER 6. In spite of sometimes-ambiguous results from experiments to test ability of pure compounds to inhibit cancer or coronary heart disease. Experimental studies with individual compounds validate their importance but do not explain completely the efficacy of fruits and vegetables.22. heart disease.16–18 Natural antioxidants such as vitamin C.23 These esters are about 90% ferulate and 10% coumarate. Such studies demonstrate the benefits of grain fiber.14. it is generally accepted that natural antioxidants are important to good health. and Antioxidants Gene Miller INTRODUCTION Considerable scientific data suggest that the risk of cancer.00+$1. Tocotrienols. and β-carotene were identified as potential active components. long-chain mono. which together constitute about 1% of the grain 0-8493-2387-8/01/$0. The French paradox observation.50 © 2001 by CRC Press LLC 453 .7–9 but for grain products. it has been suggested that fiber alone does not explain the full effect of whole grain products. benzoates. May 6. vitamin E.11. most research centered on fiber as the efficacious agent. Fiber.1_fm Page 453 Sunday.15 Epidemiological studies regarding fruits and vegetables clearly indicate their positive impact on health regarding chronic.1 Whole Grain. but some are unique. and vegetables. In addition. degenerative disease.19 draws attention to the wide variety of phenolic antioxidants that may act to prevent disease.1. flavonoids were recognized by SzentGyörgyi20 in 1937 to have vitamin-like activity. and vegetables in diverse structural combinations. and sound mechanisms have been proposed to explain biological function. WHOLE GRAIN ANTIOXIDANTS Whole grains contain numerous antioxidants of different structural types. fruits. These compounds vary in solubility from freely soluble in water to fat soluble. Insoluble antioxidants include phenolic acids covalently bound by ester linkages to the arabinose side chains of arabinoxylan fiber polymers of cell walls. and tocopherols are basic antioxidant compounds common to grains. 2001 7:46 PM 454 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. results from studies on physiological effects of diets that contain raw grain and bran or intact cooked grain could differ from studies using the same grains but that have been processed in ways that disrupt cellular structure. seed coat.1 Plant antioxidants. dependent on microflora activity.2387_ch6.1_fm Page 454 Sunday. 3RD EDITION R3 CH3 CH3 HO C=C CH3 COOH CH3 CH3 CH O CH3 CH3 R1 R2 CH3 Cinnamic Acids Tocopherols OH R O HO O OH O OCH NH OH OH COOH O Avenanthramides Flavonoids HO O HO HO OH (CH2)nH O HO O RO O Diol Esters OH Alkyl Resorcinols Figure 6. Seven different diferulates have been identified that could be formed by a radical mechanism. A substantial part of ferulate esters dimerizes to form diferulate bridges between fiber strands and give structural strength to the cell walls.1. Components of grain contained in thick-walled bran cells are hindered from absorption. In this respect. .25 Antioxidants esterified to grain fiber are not available for absorption unless freed by enzymatic hydrolysis in the colon. The majority of grain antioxidants are found in outer layers of grain seeds. as are most other phytonutrients. May 6. Even soluble materials may be carried to the colon with bran. spinach and beets being familiar examples.24 Fiber-bound phenolic esters are common to grains and grasses but are found in only a few of the dicots. where they may or may not be released for absorption. This may partially explain why ORAC values on aqueous extracts of cereal products were similar to DPPH results of the whole product. it was of interest to estimate total activity in grain products. This may be due to differences in products analyzed.1. The ORAC method utilizes a high-energy free radical (2. May 6.1. vegetables. Fruits. For example. but average activity for vegetables was similar to fruits by the ORAC method.28 A significant part of the antioxidants in grains is not readily soluble in 50% aqueous methanol.30 The same kinds of fruits and vegetables were analyzed in both studies. Because of this. and ingredients were analyzed for antioxidant activity using the DPPH method. stable free radical that reacts slowly with some antioxidants.27. White rice with no bran has significant antioxidant activity but is considerably lower than brown rice with bran. unfortunately.1_fm Page 455 Sunday. there are significant differences in activity between different grains. DPPH reacts very slightly with β-carotene and only slowly with vanillic acid. is a minor grain in most Western diets. antioxidants have a wide range of activity. Similar results were reported for aqueous extracts of foods using ORAC (oxygen radical absorption capacity) technology. grains. fruits. FIBER.26 DPPH has been thoroughly studied as an analytical reagent for antioxidants. It is apparent that whole grain products average higher in antioxidant activity than fruits or vegetables on an as is basis.1 Average Antioxidant Activity for Food Products Food Material Average TE/100 g Common vegetables Common fruits Whole wheat bread Whole grain breakfast cereals 400 1200 2000 2800 Antioxidant activity of different fruits. or methodology.2387_ch6.1diphenyl-2-picrylhydrazyl (DPPH).1. from highly reactive to very weak activity. and vegetables varies considerably. AND ANTIOXIDANTS 455 ANTIOXIDANT CONTENT OF FOOD PRODUCTS Since grains contain many different antioxidants with potential for health benefit. but a variety of compounds and activities offer potential protection for a range of human disease mechanisms. Table 6. As for fruits and vegetables. Rye is exceptionally high in antioxidant activity but.29 This procedure measures soluble antioxidants and also allows the DPPH to react with insoluble antioxidants distributed throughout the food matrix. Results shown in Table 6. Antioxidant activity for common grains is shown in Table 6. and not at all with very weak proton radical donors.2′-azobis(2-amidinopropane) dihydrochloride). The significance to human health of different antioxidant compounds that have diverse chemical properties is unknown. as previously reported. is trapped in coarse cells of the bran. DPPH was reacted directly with finely ground sample in warm aqueous methanol for 4 hours. Analysis of antioxidant activity was done using a stable free radical. or is covalently bound to cell walls. 1. in 50% aqueous methanol. 2001 7:46 PM WHOLE GRAIN. Consumption of a variety of foods is important to ensure a balanced intake of phytonutrients including antioxidants. grains. and vegetables. whole grain products are substantially higher in antioxidant activity than commonly consumed fruits and vegetables.1 are expressed as Trolox equivalents per 100 g (TE) of sample. which can react with weak antioxidants.26 Vegetables range from 1400 TE for red cabbage to 50 TE for celery. solvent systems. The relative amount of antioxidant activity in these foods is an indication of potential contribution to disease reduction by antioxidant mechanisms. Annual consumption of rye is . In addition to solubility issues. fruits vary from 2200 TE for red plums to 100 TE for melons. processed products. Pearled barley with partially removed bran is higher in antioxidant activity than red wheat.2. Antioxidant activity was 3000 TE for a whole wheat cereal and 1300 TE for a cereal made from refined rice. DPPH is a bulky. Red wheat is higher in antioxidant activity than white wheat or oat groats. Regardless. or whole grain breakfast cereal are compared in Table 6. whole wheat bread. Tannins.1. antioxidant intake could be increased significantly by modest diet changes such as eating whole grain breakfast cereals and whole grain breads in place of refined grain products. It is not possible from these results to determine how much of total bound or insoluble antioxidants is measured by DPPH. Antioxidant activities calculated for an average serving of fruit. vegetable.1. but it is reasonable to assume a significant part is from phenolic acids esterified to fiber. and it is believed to have important health benefits due to lignans and other phytonutrients.1. Whole grain cereal that was extracted with 50% aqueous methanol had about 65% of starting activity. but DPPH analysis indicates that there is significant antioxidant activity associated with fiber of cereal grains. Table 6.3. There is not a convenient way to evaluate the relative health significance of antioxidant mixtures in different foods. which have very high antioxidant activity. Insoluble antioxidants have not been completely identified. Fruits. and Vegetables Product Serving (g) TE Activity Whole grain breakfast cereals Whole wheat bread Average fruit Average vegetable 50 50 100 100 1400 1000 1200 630 Antioxidant activity is compared for grain fractions and grain products in Table 6.7.1. Antioxidants are concentrated in bran fractions.1_fm Page 456 Sunday. 2001 7:46 PM 456 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.31 All whole grains contain substantial antioxidant activity when compared to an average value for common fruits or vegetables. Table 6. white (dry) 900 11000 2500 6000 1700 1600 2050 1100 .32 and lignins may explain some of the activity.2387_ch6. The amount of antioxidants from a serving of whole grain product is similar to that from an average serving of fruit and greater than an average serving of vegetables. but it is reasonable to assume that very low activity foods have less potential for benefit than the medium or high activity foods. Even oat hulls that were extracted with aqueous alkali retained high antioxidant activity. May 6. although the endosperm has significant activity. 3RD EDITION Table 6.4.1. MeOH ext’d wheat cereal White bread (dry) Bread. Since Western diets typically contain relatively little whole grain products and are low in fruits and vegetables.4 Antioxidant Activity of Grain Fractions and Products Ingredient TE/100 grams Red wheat flour Red wheat bran Whole red wheat Alkali extracted oat hulls Aq.3 Antioxidant Content For Average Servings of Whole Grain Products. crust (dry) Bread.2 Antioxidant Activity of Grains Grain TE/100 grams White rice Whole white wheat Brown rice Oat groats Whole red wheat Pearled barley Rye 700 1400 1500 1800 2500 3100 4700 high in Finland. 4). In addition. For example. Potential health benefit of unique antioxidants. The crust is about double in antioxidant activity compared to the crust-free part. A similar effect is seen for breakfast cereals. wine. it was determined that 19% of the label was excreted in the feces. these compounds may also participate in other mechanisms suggested for specific antioxidants. This activity is consistent with reports for phenolic content in bran fractions. In addition. This suggests that there is little change in antioxidants in the interior of the bread loaf during baking. This is demonstrated by comparison of crust antioxidant activity to that of crust-free white bread. antioxidant activity in baked bread is higher than the starting flour (Table 6.46 This suggests that availability of phytonutrients in whole grain will be reduced unless the grain is processed to open bran cells and colon microflora are healthy. Antioxidant activity increases gradually during cooking steps of breakfast cereal processing. It appears that there is essentially no loss of natural antioxidants while there is formation of new antioxidant activity. AND ANTIOXIDANTS 457 PROCESSING AND ANTIOXIDANTS Another aspect of grain antioxidant activity is that it does not appear to decrease during bread making or processing for breakfast cereals.43–45 Arabino-ferulates and free phenolic acids are released by microorganism fermentation. are bioavailable.37–42 Further. but other constituents would not be available at all except after fiber degradation by bacteria in the colon.48 Grain fiber material has significant antioxidant activity. 20% was excreted in urine. Bran fiber is of interest because of how it may effect the distribution and absorption of phytonutrients. High temperature conditions at the loaf surface promote crust browning. including Maillard reaction products. High-protein and fatty diets reduce the abundance of colon microorganisms as compared to a high-fiber diet. It is reasonable to assume that many of the grain and grain product antioxidants are bioavailable and can function as free radical scavengers in the body. From experiments with rats fed 14C-labeled bound phenolics in spinach. Diferulate release is possible but is more difficult because of attachment to two fiber chains. the contents of cells will be much less available for absorption. antibiotics can inhibit growth of colon microflora that ferment fiber for over 30 days. after release in the colon by enzymatic hydrolysis. bioavailability of grain antioxidants in food products has not been determined specifically. but enzymatic hydrolysis is possible in the colon.2387_ch6. The crust-free part is about the same as flour on an equal moisture basis.45 These results demonstrate that bound phenolics.1. most likely Maillard reaction products.44. and over 34% of the label was retained in body tissues after 18 hours.33–35 And it is known that antioxidants released in the colon by microbial action can be absorbed. FIBER. but the biggest increase is from toasting. ANTIOXIDANT ABSORPTION AND METABOLISM It has been shown that natural antioxidants are bioavailable and that the level of antioxidant activity in blood increases on a diet high in fruits and vegetables or after consumption of foods high in antioxidants such as tea. Covalently bound phenolics are resistant to digestion in the stomach and small intestine. If the bran of whole grains is essentially intact when it is consumed. cell apoptosis. Reductone products are characteristic of the Maillard reaction and may explain the increase in crust antioxidant activity. and induction of detoxification enzymes. or fruits. vasorelaxation. antioxidants in the digestive tract have potential to react with nitrites and free radicals existing in food when consumed or generated by biological processes. May 6.22. readily soluble in water.1_fm Page 457 Sunday.36 However.47. is not known. such as reducing platelet adhesion. with consideration for natural . Fiber-bound phenolics were identified many years ago and have been the subject of considerable study in recent years. may diffuse out of the bran cells. Small molecules. 2001 7:46 PM WHOLE GRAIN. It is essential that the everyday diet is relatively high in fiber to help sustain a microflora that is optimum for obtaining maximum health benefit from grain phytonutrients. It may be that corn bran contains high levels of ferulate.000 TE by DPPH analysis. deliver a unique functionality to the colon and are an important part of the overall health benefit provided by whole grain products. 2001 7:46 PM 458 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. In the case of antioxidants. in synergy with fiber.400.000 TE. Diet and Health. This activity.1. along with the protective effect of fiber and fiber fermentation products.000 TE. 2. Washington. to be released by fermentation in the presence of appropriate microorganisms. and water-soluble antioxidants can be absorbed in the small intestine. Washington. sphingolipids. Antioxidants covalently bound to cell wall fiber and compounds contained within cellular structures are transported to the large intestine. Food. including significant amounts of phenolic acids esterified to cell wall fiber. Although high. Implications for Reducing Chronic Disease. Whole grain antioxidants represent a variety of structural types. tocotrienols.1% ferulate.2387_ch6. DC. Based on the activity measured for tannic acid (1. National Academy Press. equivalent to 2.9% ferulic acid if all activity is due to ferulates. DC. Whole grain products provide equal or greater antioxidant activity compared to fruits or vegetables on a per-serving basis. Covalently bound antioxidants are transported to the colon along with fiber and remain functional agents for an extended time period. including antioxidants. REFERENCES 1. there are several ways they can be utilized by the body. lignans. analyzed for 160. In addition. In this manner. Transport of antioxidants to the colon is characteristic to whole grain materials because of covalently bound phenolics but is possible for only a few vegetables and fruits. World Cancer Research Fund and American Institute for Cancer Research. 1997. This calculates for 3. Fat-soluble antioxidants can be absorbed through the lymphatic system. and minerals.32 The sample of very pure red wheat bran (Table 6.50 It is of interest that carob hulls. sterols. this is in the range of ferulate concentrations reported for wheat bran. Enzymatic hydrolysis in the colon can free phenolics for absorption into the epithelium. phytates.49.32. Epithelial cells of the colon absorb free phenolics and can benefit from their antioxidant activity. there is time-delayed activity present in the digestive tract.1_fm Page 458 Sunday. It is hypothesized that bound antioxidants. Antioxidants released in this manner are available to scavenge free radicals present in the colon. carob could contain roughly 11% tannic acid that is reactive with DPPH in 50% aqueous methanol. . Nutrition and Prevention of Cancer: A Global Perspective. May 6.44.4) was 11. vitamins. AICR. which are known to be high in tannin compounds. CONCLUSION It is well established that grain fiber is important to human health. The health contribution of antioxidants in grain products has received little attention.44. and it may contain small amounts of highly active compounds such as tannins. whole grain antioxidants can act as free radical scavengers through the entire digestive tract and in colon tissues. including antioxidants that are concentrated in the bran. It is possible that there is a synergistic relationship with fiber that goes beyond a simple combination of chemical compounds. may provide unique protection that is not possible by any single component. 1989. there are many other potential health-supporting phytonutrients in whole grains. When fiber carries bound antioxidants to the colon. Processed whole grain products contain considerable antioxidant activity in the form of phenolic acids covalently bound to cell wall fiber.000 TE).49 Antioxidant activity for a very pure sample of corn bran was 20. National Academy of Sciences. Grain products provide many phytonutrients. These hypothetical mechanisms offer additional possibilities to explain the efficacy of whole grain products to reduce chronic disease. 3RD EDITION variability and activity from insoluble polyphenolics such as lignin or tannins. F. J. G. Speizer. Chem. H. G. E. Ed. fruit and vegetables.. J. 61. 84. L. Long-term intake of dietary fiber and decreased risk of coronary heart disease among women. Acad. N... 3.. J. Meyer. J. 79. and Hatfield. I. 1996. A diet high in whole and unrefined foods favorably alters lipids. L.. and Young. Soc. 2001 7:46 PM WHOLE GRAIN. Ferulic acid and diferulic acid as components of sugar beet pectins and maize bran heteroxylans.. S. and Folsom... J.. CRC Handbook of Dietary Fiber in Human Nutrition. chap. Fd. Fukumoto. Anderson. Natl. F. Perkin Trans. Spiegleman.. G. 1999. E. Am. R. Saulnier. 281(21). Gey.. 85. Antioxidants and atherosclerosis: a current assessment. Cancer. H. Does cereal reduce the risk of cancer?. and Kanter. Am.. A. F. Ralph.. Lancet. 1937. 3597. B. Steinberg. Webster. Chem. 1189. K. Davin. Use of a free radical method to evaluate antioxidant activity. Bentsath.. R. 7. L.. College Nutr. Wolk. and Lewis. H.. K. Paul. S. 1995. 307. G. M.. fruit and cereal fiber intake and risk of coronary heart disease among men. Prakash. 5... platelets and the French paradox for coronary heart disease. C. FIBER. M.. M. S. 27. 1999. H.. Plant lignans and health: chemoprevention and biotechnoloical opportunities. Greenberg. M. JAMA. Marquart.. Agric. N. Jacobs. L. American Association of Cereal Chemists. Jacobs. Eds. E. Rusznyak... 8. M. 90. H. F.. Antioxidant content of whole grain breakfast cereals. Ed. 1999.. 26. W. F. W. Hennekens. A. 89. 19(S). S. P. chap. Rimm. epidemiology and mechanisms. in Oat Chemistry and Technology. R. 1997.. L. 19(1). Spiller. of Nutr. Kushi. J. E. 44(2). and Sporn.. 6. Klevay.. Renaud. 20. C. A. 24. J. Adlercreutz.. Sci. Ed. 1986.. 2000. A. Morphological and chemical organization of the oat kernel. A. and Fürst. . J... A. Med. 2nd ed. 10. Shigena. 2000. H.. B.... USA. Ames.. Whole grains and protection against coronary heart disease: what are the active components and mechanisms.. A. 21. Med. F. 18. Nature. JAMA.. 19.. antioxidants and the degenerative diseases of aging.. T. FL. 2000.. and Mazur. P. and de Lorgeril. 1999. and Serra. Fd. A. Am. Agric. St. Nutr. 275. L. Nutr.. Oxidants.. Kardianal. Ascherio. American Association of Cereal Chemists. W... Assessing antioxidant and prooxidant activities of phenolic compounds. Ford. 17. M. 125(6). E. Kluwer Academic/Plenum Publishers. Oat phenolics: structure. R. Soc.. 1993. health benefits and safety concerns. Pharmacology. Ecology. Whole grain intake and cancer: an expanded review and meta-analysis. 15. M. M. 1995. Phyto-oestrogens and western diseases. the Iowa Women’s Health Study. Boca Raton. Vitamin P. Office for Official Publications of the European Communities. 16.. 1041. D. Dietary fibers differ in their effects on large bowel epithelial proliferation and fecal fermentation-dependent events in rats. Coll. Giovannucci. G. Eds.. Slavin.. Brand-Williams.. Folino. and Berset. Food Chem.) varieties. 1999. Nutr. Circulation. 48. 3125. 18. Cereal Foods World. 1996. J. AND ANTIOXIDANTS 459 3. E. T. 1999. J. and Hanson.. Christensen.. 95. B. L. F. J. 2000. D. Stampfer. 30(2).. K. 447. B. Cuvelier. K.. S. 76. S. W. 326. Public Health. Grabber. Bruce. 1990. F. 1994. B. Stampfer. The antioxidant hypothesis of cardiovascular disease. S.. L. Rigelhof. G. 339.. 675–694. 1999. P. Meyer.. occurrence and function. Content of phenolic acids and ferulic dehydrodimers in 17 rye (secale cereale L. 7915. Webster. B. A. 1992. 1986. 14... and Hanna. Biology. 25. 1523. antioxidant defenses and colon function.2387_ch6. J. A. BauschGoldbohm. D. 1993. R. in Plant Polyphenols 2: Chemistry. and Hagen. 1998. H. W. in Oat Chemistry and Technology. 322. 4. L. Spiller.. and Mazza. 396. 25. Miller. Proc.. Vegetable. 28. 1420. R. and Kushi. A. G. C.. St... Phyto-estrogens: exposure.. R. cancer and cardiovascular disease.. Andlauer. alcohol. Trans. and Szent-Györgi. J.. Is whole grain intake associated with reduced total and cause-specific death rates in older women. Sci. 3485.. COST 916... McIntyr. 29.C.. Antioxidant vitamins. bioavailability.. 70. F. Clin. and Gallagher. D... Lebensm-Wiss U-Technol.. 13. W. W.. 48. Agric. Andreasen. 23... A. 9. Luxembourg. J.. New York. J. J. Am. Eng.. 12. and Thibault. J. Hu. 9. Quideau. 1998. D. Biochem. 22. Ann.. 11. 1521. Identification and synthesis of new ferulic acid dehydrodimers present in grass cell walls. CRC Press. 1991.. May 6. Marquart. Fulcher. and Willet. M. M. Manson.. Gross et al. J. Colditz. Collins. F. E.1_fm Page 459 Sunday.. M. 334. 2837. M.. and Willet. A. 139. 28. Paul.. Wine. H. A. H. 1996. 427-435. Frank.. cholesterol.. 2000. A. Kroon.. 35(4). Adlercreutz. Y. Protection against cancer by wheat bran: role of dietary fibre and phytochemicals. Wallace. L. R. Chem. A. J. and Eastwood. B. R. May 6.. Nutr. P. 1999. 4595. Eur. J.. High molecular weight polyphenolics (tannins) as biological antioxidants. B. C. F. Papers Am. Fd. 118.. 441-447. Marquart.. 411–413. 32... 49. 8. Meyer. Kamal-Eldin. 38. 1999.. 37.. Nutr. A.. A. Nutr. Ferguson. S. P..... Agric.. A. Hydroxycinnamic acids in the digestive tract of livestock and humans. 1998. Res. Jacobs. personal communication. 36. Bioavailability of flavanoids from tea. S. 2383... In vivo release of 14C-labeled penolics groups from intact dietary spinach cell walls during passage through rat intestine. 2000. 57(8). N.1_fm Page 460 Sunday. Faulds. J. A. Abstr. C. spinach. Hollman. 373. Kuchi. C.. C. Fd.. 56(11). Fd. K. Rye: a Nordic focus.. 65.. Fd. N. A. 1999. H. L. and Cai. Teugblad. M. Razdan. Phillipson. Res. G. Richardson. Agric. Y. Russel. J. dietary source.. Shiyi. Lischner... Sci. L. and Marquart. and Vessby. Miller. 2000. G. 46. S.. Release of hydroxycinnamic and hydroxybenzoic acids in rye by commercial plant cell wall degrading enzyme preparations. M.. Chem. 1998. W. C. and Rrior. 241. Agric. Rev. H.2387_ch6. and Kanter. Prior. 2001 7:46 PM 460 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION... R.. T. Sci. G.. Agric. and Williamson. Bravo. Buchanan. Antioxidant content of cereal grain ingredients and commercial breakfast cereals. Commun. O. Rigelhof. Bowery.. Cao. G.. Nutr. J.. 17.. F. 31. J.. S. L. F. Biochem. Kroon. C. 46.. 41. Christensen. R. C. 1664.. L. Robertson. 44. Chesson. and Stewart. M. 43. G. 59. Serum antioxidant capacity is increased by consumption of strawberries. 45. Smith. J. Sci. Rong.. . and Inglett. 79. 1999.. 34. 44.. 1998. A. 47. R. Release of covalently bound ferulic acid from fiber in the human colon. J. F.. Plausible mechanisms for the protectiveness of whole grains. J. M... Clin. Russell.. Agric. K. Semboku. L. G. P. Chem. G. A study on OH radical scavenging activity of dietary fibre from wheat bran. 216. J. 1887. Slavin. J. P. Provan. J. J. J. Cancer Prev. A. 317. 48. K.. Fd. and fatty acids in rats. J. and Williamson. 50. 45(2). 79. Lipids. Poutanen. Fd. J. Plant polyphenols: biologically active compounds or nonselective binders to protein?.. M. Andreasen. and Yang. 128. and Sevanian. Hydroycinnamates in plants and food: current and future perspectives. N. and Rice-Evans. P. Neutraceut. metabolism and nutritional significance. J. 719. 30. T. 3. Tubaro. Martini. R.. Crit.. E.. Fd.. A. and Harris.. A. Zhu.. and Liukkonan. Jones.. 1997. E. 45. P. 661... Bourne. A.. Whole grain products and antioxidants. M. K. 1997. Optimization of nutrition: polyphenols and vasculer protection. Basu.... 253. 1998.. and Hartzfeld. S. 191.. 70(5). Carb. Antioxidant activity of corn bran cell wall fragments in the LDL oxidation system. N.. 35. 1997. Effects of dietary phenolic compounds on tocopherol. M.. 40.. Greengrass. 37. M. and Kongron. 1997. Ohta. 2000. Chem. Sovik. Agric. and Hansen. Sci. Soc. 1983. 21(2). L. Fry.. and Hartley. 222. Cao. Fd. Polyphenols: chemistry. A.. Func.. 30.. 1999. D. Tijburg. 1999. 71. 1999. G. Yan.. C. J. 42.. 39... Agric. 119. L. and Sanada. 79. Egashira. C. R. Ryden. Hagerman.. J. Occurrence and nature of ferulic acid substition of cell-wall polysaccharides in graminaceous plants.. H.. Bioavailability of ferulic acid. L.. Phytochemistry... 459. M.. M. Riedl. Am. B.. Ursini.. 1998.... Biophys. L. C. Nutr. 45(3). Sci. Prakash.. J.. 355. D. L. Rev. Ritchard. red wine or vitamin C in elderly women. B.. E.. N. Acta Nutrimenta Sinica. H. Fd. Scobbie.. 33. Cereal Foods World. Rev. M. 3RD EDITION 29. W. L. GRAIN INTAKE AND DEFINITIONS FOR EPIDEMIOLOGIC STUDIES At the base of the Food Guide Pyramid. Leonard Marquart. British Medical Journal. Recent epidemiologic findings on whole grain foods allow us to evaluate the food source of fiber. and confirmation from other studies in similar and different populations.2_fm Page 461 Sunday.50 © 2001 by CRC Press LLC 461 . and Joseph Keenan INTRODUCTION The finding on cereal fibre illustrates one of the uses of epidemiology — in exploration.2 exploratory findings for cereal fiber intake and CHD risk were in need of support by candidate mechanisms. with at least 95% being stripped of its bran and germ. Pereira. May 6.3. important epidemiologic observations have been made on the topic of dietary fiber and chronic disease risk. This hypothesis came from whole plant foods. Joel J. J. in association with chronic disease risk.2 Whole Grains. As indicated by the quote above by Morris and colleagues. and less attention being given to whole grain foods. grains compose approximately 25% of the food supply in the U. 1977 The hypothesis that dietary fiber may reduce the risk for chronic diseases was extrapolated from the ecological observations of Burkitt et al.2387_ch6. very little grain is available in whole or minimally refined form. which we describe herein.. we have developed a nontraditional hypothesis that fiber alone is only one potentially efficacious component of whole grains and other appropriately processed plant foods. As such.S. and Chronic Diseases: Epidemiologic Evidence Mark A.5 It has been estimated that average intake of whole grain 0-8493-2387-8/01/$0. Jr. N. which we describe in Chapter 6. Pins... with fruits and vegetables historically and contemporaneously receiving much public health attention.1 who compared and contrasted the diets and disease patterns of Westernized and non-Westernized cultures. with its nutrientrich complex still somewhat intact. 2001 7:47 PM CHAPTER 6. The goal of this chapter is to review the observational epidemiologic evidence for the potential role of cereal fiber and its primary food source — whole grains — in reducing risk for chronic diseases.00+$1. Jacobs. Cereal Fiber. Morris et al. David R.3 However. Over the past few decades. thus fueling a reductionist approach rather than a “whole food” one.4. The loss of critical nutrients in the bran and germ during the refining process is acknowledged by the enrichment of refined grains with many vitamins and minerals by the food industry in an attempt to replace natural nutrients. and cancer (see Chapter 6. For a thorough review of nutritional epidemiologic methods. These include commercially available “dark” or “mixed grain” breads. Furthermore. Prospective studies have the capability to exclude those with the disease of interest at baseline. and follow them through time to track the number of cases (e. perhaps synergistically.g. Table 6. whole grain foods. may recall dietary and other exposures differently . May 6. unsaturated fatty acids. Given the wealth of longitudinal studies on fiber. We have hypothesized that these nutrients present in grains in their natural form may act together. or time 0. and many micronutrients and phytochemicals.3). however.6 In observational epidemiologic studies based on self-reported dietary intake. and brand-name breakfast cereals with a refined grain as the first ingredient and less than 2 g of fiber per serving. may help to provide further impetus for modification of the current dietary guidelines in Westernized countries. Cases. Case-control studies have commonly been done for cancer. we will not thoroughly review the cross-sectional studies. 2001 7:47 PM 462 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. because of their disease status.10 Study Design Prospective studies are stronger than cross-sectional. white rice. A quick review on the interpretation of the studies described in these tables and the accompanying text may help in judging strengths and weaknesses of the literature. thereafter conducting analyses to determine if self-reported diet at baseline is associated with incidence of disease.9 Epidemiologic evidence reviewed in this chapter. we are limited to rather crude differentiation between whole and refined grain intake.7.2387_ch6. diabetes. and CVD and type 2 diabetes. including some unidentified. because with cross-sectional studies it is not possible to establish temporality — that is.1 summarizes the foods contributing to whole and refined grain intake for epidemiologic studies utilizing databases of self-reported intake.2_fm Page 462 Sunday. Case-control studies are particularly prone to bias. whole grains. is ~0. especially recall bias and other biases related to the selection of controls and interviews that probe for exposure status. 3RD EDITION products in the U.5 eating occasions per day. are not able to be replaced. the presence of frank disease in cross-sectional studies may result in reverse causality — real changes in diet due to secondary prevention efforts or consequences of illness or medication use — or self-report bias in dietary recall due to knowledge of diet and health. and experimental evidence of the accompanying chapter. refined grains are defined for epidemiologic purposes as those foods that are known or strongly suspected to have no bran or germ due to the refining process and only the fiber-poor. and chronic disease risk. enrichment is insufficient when one considers that fiber. as this study design is more efficient for less common diseases. BRIEF METHODOLOGIC OVERVIEW OF NUTRITIONAL EPIDEMIOLOGY The methodologic nuances of nutritional epidemiology should be reviewed prior to embarking on a critical evaluation of the published studies. in the body to modify risk for chronic diseases such as cardiovascular disease. or brand-name breakfast cereals with a whole grain (or whole grain flour) as the first ingredient or elsewhere on the ingredient list with at least 2 g of fiber per serving.2. Foods coded as whole grain are those that we know or suspect to include bran and/or germ and therefore are a potentially nutrient-rich fiber complex. the reader is referred to Willett. These include white bread. nutrientpoor endosperm present.. incidence of myocardial infarction). We have arranged tables in this chapter to guide the reader through the salient features of many studies of dietary fiber.8 In contrast. brown rice. However. that the exposure (diet) precedes disease onset. with total grain products (whole + refined) being less than 3 per day in comparison to the recommended intake of 6 to 11 servings per day.S. brown. AND CHRONIC DISEASES: EPIDEMIOLOGIC EVIDENCE 463 .Whole wheat pancakes and waffles Made from whole wheat or other whole grain Brown and wild rice Popcorn. tortillas.5.30 and Pereira et al. cakes. rolls.1 2387_ch6.8 Pizza. and croissants Regular pancakes and waffles Made from white or semolina flour or other refined grain flour White rice White wheat flour crackers and pretzels.2. May 6. corn chips. donuts. mixed grain bread Source: Adapted from Jacobs et al. pasta. cookies and brownies Refined grain or sugar as first ingredient White bread Foods 3% 45% 8% 10% 4% 30% Estimated Relative Contribution to Food Group Refined Grain Description and Relative Consumption of Common Whole and Refined Grain Foods in Epidemiologic Studies Food Group Table 6. rye. and pretzels 25% Whole grain or bran as first ingredient Whole grain or bran elsewhere in ingredient list AND at least 2 g fiber per serving Oatmeal and other hot whole grain cereals Whole wheat. etc. and bran muffins 1% 13% — 1% 60% Dark.2_fm Page 463 Sunday. 2001 7:47 PM WHOLE GRAINS. Rice Snacks and desserts Other breakfast items Bread and bread products Breakfast cereal Foods Estimated Relative Contribution to Food Group Whole Grain Whole grain or bran elsewhere in ingredient list AND less than 2 g of fiber per serving Cream of wheat and other hot refined grain cereals Regular white flour muffins. whole wheat or rye crackers. CEREAL FIBER. Therefore. thus biasing associations toward the null hypothesis of no association between diet and disease. depending on the sampling strategy and age. such a sample has internal validity due to the comprehensive and careful methods.g. as will be . The high response rate and very large sample size in the Nurses Health Study also contribute to the enhanced likelihood of detecting important associations between diet and health and generalizing these findings to middle-aged and older women who have similar characteristics.10 depending on the nature and quality of the measurement error adjustment and. Within the physically active group of participants there may be a broad distribution of physical activity and dietary fiber intake.2387_ch6. However. Adjustment for Other Potential Confounders Adjustment for confounders is especially critical with such self-reported behaviors as intake of fiber-rich foods. This potential problem would be minimized if a study had a very detailed measure of physical activity. Indeed. vitamin supplement use. and racial composition of the sample. 2001 7:47 PM 464 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. the Nurses Health Study). May 6. Measurement of Diet There is a large degree of error in self-reported diet in epidemiologic studies. and increasing the probability of missing a true association (type II error). as reviewed in Chapter 6. gender. the more likely the sample is to have a broad distribution of dietary intake and the greater the number of disease endpoints that will be accumulated over time. the association is likely to be real. if a certain study of fiber and risk for coronary heart disease has a crude measure of physical activity. In fact.2_fm Page 464 Sunday. However. which correlates strongly with important lifestyle behaviors such as cigarette smoking. National Health Interview Survey) or a population subgroup (e. In addition. 3RD EDITION (overestimating or underestimating exposure) than controls. and other dietary factors. Although female nurses may be a relatively homogenous group. into two categories — sedentary vs. when one finds a statistically significant and biologically meaningful association in support of some a priori hypothesis.g. such as classifying participants. leaving open the possibility of confounding within subgroups of physical activity.3. Many studies discussed in some detail in this chapter have rigorously controlled for potential confounders. and the discrepancies between these case-control studies and the few prospective studies of fiber and cancer will be discussed. especially after control for potentially confounding factors (see below). Not only must epidemiologists control for these potential confounders in their statistical modeling. some comprehensive metaanalyses of whole grain and cancer have considered these biases in their methods. Sample Size and Characteristics The larger the sample size and the more representative the sample is of the overall population. if one also finds support for physiologic effects from animal experiments and human trials. of course. but they must have measured these potential confounders accurately to minimize susceptibility to residual confounding. such as a continuous metabolic index derived from a series of questions about type. physical activity. residual confounding is less likely to be operating to explain the observational findings. frequency. This error is generally agreed by nutritional epidemiologists to be random in nature. a larger sample size may not always result in an unbiased and representative sample of the population in terms of dietary intake and characteristics. For example. there is evidence that adjustment for this measurement error may sometimes strengthen associations. whether there is a true association as hypothesized. These factors will increase the likelihood of detecting an association between exposure (fiber or food intake) and endpoint (risk factor or incidence rate of disease)... and intensity of activity. The overall population from which the participants are recruited may be an entire country (e. active — physical activity level could still confound associations between fiber and incidence of coronary disease. a 20% reduction in risk may have public health relevance. generalizable amounts of a nutrient or food. Insulin may promote the growth of tumors. Such effects of dietary fiber may operate systemically.2387_ch6. while a more thorough review of associations between fiber (and its food sources) and actual incidence of frank disease will follow.16 have identified possible mechanisms to explain their associations between intake of crude fiber and coronary heart disease incidence (included in the next section) in hypercholesterolemic middle-aged men. There are a few biomarkers reflecting risk for CVD and type 2 diabetes that also may be important in the etiology of certain cancers. systemic inflammation. 2001 7:47 PM WHOLE GRAINS. AND CHRONIC DISEASES: EPIDEMIOLOGIC EVIDENCE 465 discussed below. type 2 diabetes. impaired glucose metabolism. markers of inflammation may reflect an immune response. one might ask: What is the relative risk reduction for coronary heart disease for each increase in cereal fiber of five grams per day — or the equivalent of one or two servings of whole grain breakfast cereal? If a dose-response curve is also observed across increments of increasing nutrient intake (whereby risk for the disease is decreased with increasing intake). in part because cancer is a much more heterogenous set of diseases than CVD and type 2 diabetes. have the potential to affect all of these risk factors for the three main chronic diseases — cardiovascular disease. For example. and antioxidant capacity. hypofibrinolysis. Humble et al.12–15 This association appears to be independent of other dietary and lifestyle factors. then the findings are more likely to be real rather than spurious or confounded. few studies have provided good detail of fiber–risk factor associations. and central fat patterning. with such common diseases as cardiovascular disease.11 Risk factors for cancer are less clear. One of the most promising fiber–risk factor associations is that of glucose control and insulin sensitivity. especially if this risk reduction is observed between groups within the study population consuming realistic. Across quintiles of increasing fiber intake at baseline of the study. hypertension. because these diseases have many risk factors in common. The few cross-sectional and longitudinal studies of dietary fiber and fasting insulin and other markers of insulin sensitivity have revealed that fiber appears to be favorably associated with insulin sensitivity. obesity. Fiberrich foods. for reasons discussed in the accompanying chapter along with a review of the experimental evidence. Indeed. Instead. The crosssectional studies of dietary fiber reviewed herein do not include frank cases of disease. including hyperinsulinemia. some nutritional findings from the Nurses Health Study and another study of postmenopausal women from the state of Iowa (the Iowa Women's Health Study) are consistent.001). Magnitude and Nature of Association Generally. Although the cholesterol-lowering effect of dietary fiber appears to be modest and dependent on its solubility (as discussed in the accompanying chapter).2_fm Page 465 Sunday. because of the difficulty in measuring diet by self-report and the expense of collecting good risk factor data in large samples of individuals for epidemiologic purposes. CEREAL FIBER. Risk factors for cardiovascular disease and type 2 diabetes will not be differentiated. However. resulting in benefits on blood pressure and hemostatic factors through the clustering of related abnormalities as mentioned above. there are many other potential mechanisms whereby dietary fiber–rich foods may reduce risk for chronic diseases. a strong inverse linear trend was observed for post-challenge glucose concentration (p = . and reduced antioxidative capacity may indicate susceptibility to free-radical damage of tissue or DNA. and certain cancers. with a weaker . These may include blood insulin concentration. dyslipidemia. these diseases are thought to have a common etiology in the insulin resistance syndrome. DIETARY FIBER AND CHRONIC DISEASE RISK FACTORS In this section we will provide a brief overview of observational studies of dietary fiber or whole grain intake and risk factors for chronic disease. these studies have examined associations between dietary fiber and risk factors or biomarkers for chronic disease. May 6. antioxidants. red meat and its processing) related to a high-saturated-fat dietary pattern. and dietary fiber — were each shown to explain some of the observed association between whole grain intake . HDL cholesterol.18 These findings from the Family Heart Study18 are consistent with the PAI-1 and fibrinogen findings for our human experiments discussed in the accompanying chapter. Furthermore. As such. it may not be the saturated fat itself that is the most pernicious factor in the Westernized diet. such as minerals. carbohydrate. 3RD EDITION inverse (p = . even after adjustment for all known potential cofounders as well as body mass index.17 Those in the highest 20% of dietary fiber intake (>21 g/2000 kcal) gained approximately 8 fewer pounds of weight over the 10-year period than those in the lowest 20% of dietary fiber intake (<12 g/2000 kcal).22 In fact.20 and that the association between saturated fat intake and coronary disease risk may in fact be confounded by intake of dietary fiber. were all associated with dietary fiber intake in a dose–response manner in whites. This is a classic example of confounding that has often been overlooked with respect to the classic diet–heart hypothesis. protein. and some contributions from nuts and legumes. and insulin was not measured. and carbohydrate in this regard.06) and positive (p =. and protein.. It should also be noted that these associations were not adjusted for potential confounding factors. magnesium (abundant in whole grains and potentially an important nutrient for type 2 diabetes evolution). Fasting and 2-hour post-glucose insulin concentrations demonstrated graded inverse associations with dietary fiber intake in both black and white women and men. While the above study did not examine intake of whole foods. a graded inverse association between fiber intake and PAI-1 concentration was reported. supporting a mechanistic role of insulin sensitivity in the effect of fiber intake on CVD risk factors. (Weaker but consistent associations were observed in blacks. Study participants were 2909 young black and white adults enrolled in the Coronary Artery Risk Development in Young Adults (CARDIA) Study.11) trend for body mass index and HDL cholesterol. LDL cholesterol. May 6. No association was found for systolic blood pressure. Dietary fiber intake was consistently associated in a protective dose response manner with 10-year body weight gain.17 not only were these associations for dietary fiber and risk factors stronger than those for saturated fat. fruits. unsaturated fat. 2001 7:47 PM 466 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. but the few associations that were observed between total or saturated fat and body weight or insulin were considerably attenuated and no longer statistically significant after adjustment for dietary fiber intake. Therefore. a previous report from the CARDIA Study described inverse dose–response associations between whole grain intake and fasting plasma insulin concentration. A prospective study was recently undertaken to assess the relative capacity of the major macronutrients to predict changes in cardiovascular disease risk factors over a 10-year period (1985/6 to 1995/6).19.g. as well as cross-sectional associations with fibrinogen (a marker of hemostasis as well as inflammation). In addition to comparing fiber to saturated fat.8 These associations were independent of many other lifestyle factors associated with whole grain intake and fasting insulin. and phytoestrogens. 10-year changes in blood pressure. Factors potentially on the causal pathway between whole grain intake and fasting insulin—body mass index (through satiety or insulin itself).2_fm Page 466 Sunday. Similar results were found for the waist–hip ratio. these authors hypothesized that dietary fiber may exert its beneficial effects on attenuating atherogenesis through its effects on circulating insulin concentrations.17 Total dietary fiber from all sources was used. unsaturated fat.21 Dietary fat and fiber intake are inversely and moderately correlated — high-saturated-fat diets almost invariably include little dietary fiber.) Most of these associations could be explained by adjustment for fasting insulin concentration. In comparison. the dietary fiber consumed by this population came predominantly from whole grains. but rather the lack of fiber and associated nutrients. such as physical activity habits. Previous studies have demonstrated that fiber may have stronger associations with risk for coronary disease than saturated fat or cholesterol. and triglycerides. In the study described above. and vegetables. as source of fiber was not available for analyses. the findings may also apply to type 2 diabetes and certain hormonally mediated cancers in which insulin levels may be important.19.2387_ch6. while a recent cross-sectional study found no association between dietary fiber intake and fibrinogen. or other factors (e. 57. and regress these events on categories fiber intake derived from self-reports at baseline. In support of this possibility of a spurious finding. there have been a series of studies with very consistent findings for risk of CHD according to intake of cereal fiber and whole grain foods. Morris and colleagues admitted that they didn't have an a priori hypothesis for this finding.26 which did not include type of fiber in the analyses. . while white (refined) bread was found to have no correlation with CHD incidence. this finding may have been spurious due to the small number of events. such as age.26 observed a nonsignificant increased risk of CHD with increasing total fiber consumption.8 Only when all three of these variables were added to the model simultaneously was all of the association explained. Kushi et al.24 and Khaw and Barrett-Connor25 in the mid-1980s examined CHD deaths over 23 and 13 years. and physical activity. The studies are heterogenous in that they span a time period of 23 years and include small and large numbers of men and women. This is a very small study sample of English men (n = 337) whom the authors had to follow for 20 years in order to have the minimal number of events (45) for which to conduct their analyses. or a 43% reduction in risk of CHD in comparison to the reference group — men in the lowest tertile of cereal fiber intake.. brown breads were found to be the most influential. then total dietary fiber should have a weaker association with coronary disease incidence than cereal fiber. The findings were startling. whereas Khaw and Barrett-Connor25 observed a 15% (not significant) reduction in men and a 33% reduction in women. differing in characteristics from male British men2 to U. these authors reported no association between dietary fiber intake and allcause mortality.2 describes the salient features of epidemiologic studies of dietary fiber and coronary heart disease. They proceeded to investigate this finding further by stratification for potential confounders as one alternative to multivariate adjustment. Upon examination of the foods that contribute to dietary cereal fiber.S. cigarette smoking.24 observed a 43% reduction in CHD risk for the highest tertile of fiber intake. Studies by Kushi et al. lowest quintile of intake.2 were real. Those men in the highest tertile of cereal fiber intake had an 82% lower risk than those in the lowest tertile. for several reasons.23 The studies are relatively homogenous in their dietary and analytic methods. If the observations of Morris et al. Humble et al.2_fm Page 467 Sunday. with a nonsignificant 8% reduction for the highest vs. body mass index. the same inverse association persisted for cereal fiber and incidence of coronary heart disease.2387_ch6. May 6. Although Mann et al. a limitation of these studies was that they did not examine the type of dietary fiber. AND CHRONIC DISEASES: EPIDEMIOLOGIC EVIDENCE 467 and fasting insulin. the p-value for this observation was very low. The early study by Morris et al. All studies have adjusted for potential confounders. regressed on total dietary fiber intake in their respective populations. DIETARY FIBER AND INCIDENCE OF CHRONIC DISEASE Cardiovascular Disease Table 6. Still. They reported a graded inverse association between intake of cereal fiber and incidence of CHD events over the 20-year period such that men in the middle tertile of cereal fiber intake had a relative risk of 0. suggesting that the chances of this finding being spurious were 5 out of 1000. Although these estimates of relative risk have wide confidence intervals (not shown) due to the small sample size of this study. From 1996 to the present. The one exception is the study by Mann et al. or intake of fatty acids and cholesterol. Some studies have adjusted for additional dietary factors — a particular strength of those studies — such as fruit and vegetable intake when regressing heart disease incidence on whole grain intake or cereal fiber. CEREAL FIBER.2. energy intake. alcohol intake. respectively.2 is particularly intriguing. nurses.16 reported weaker findings for quintiles of fiber intake. However. For every stratum of smoking status and occupation and for duration of follow-up (years since baseline in intervals of 5 and 10 years). having followed healthy individuals over time to accumulate cases and deaths of coronary heart disease. 2001 7:47 PM WHOLE GRAINS. S.007 .0 1. vegetable.05 <. and fruit Highest and lowest quintiles of fiber intake Quintiles of fiber intake from cereal.94 0. May 6.0 1.0 1.83 0.71 1.08 0.93 0.005 NS Relative Risks per Increasing Category of Intake 1 2 3 4 5 p Value Note: All studies adjusted for potentially confounding variables.06 1.93 1.25 21.86 1.757 U.57 1. including demographics.68 0.92 <. p-values are for tests of linear trend for relative risks across categories of intake for studies with three or more categories.18 0. women 1801 U. Cases = fatal + nonfatal events.09 0.782 U. Todd et al. pulses. men Kushi et al.00 0.66 1.28 43.60 0.0 1. and lifestyle factors.10 NS .802 U.0 1..66 1.S.2.0 1.96 1.85 0.99 0.63 .0 1. male health professionals Humble et al.11 0. other dietary factors.13 0.20 503 U.77 .10 .0 1.2 2387_ch6. and nuts 1. 2001 7:47 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.78 0.05 .16 0.26 0.84 0.10 1.S.05 0.83 2.27 337 English men Sample Prospective Epidemiologic Studies of Dietary Fiber and Risk of Coronary Heart Disease Morris et al.92 0.001 .0 1.67 0.0 1. veg.88 1.00 0.94 0.90 0.24 45 cases over 20 years Outcome 468 Pietinen et al.23 64 deaths over 13 years 296 cases 97 cases over 9 years 591 cases over 10 years 734 cases over 6 years 249 cases over 10 years 635 deaths over 6 years 110 deaths over 23 years 65 deaths over 13 years Quintiles of fiber intake from cereal. female nurses Mann et al.08 . and fruit Quartiles of dietary fiber intake Tertiles of fiber intake Quintiles of fiber intake from cereal.S.91 0.2 Study Table 6.25 0.76 1.01 . vegetable.71 0.10. vegetable. body mass.0 1.92 0. men Khaw et al.0 1.K.39 0.82 0.81 0.64 0.83 0.15 <.51 Not given .0 1. 3RD EDITION .2_fm Page 468 Sunday.0 1.88 0.0 1.56 0.57 0.89 1.0 1.70 0. and fruit Difference in fiber intake of 6 g Tertiles of fiber intake Tertiles of fiber intake Exposure Cereal Vegetable Fruit Cereal Vegetable Fruit Cereal Vegetable Fruit Cereal Fruit.86 1.930 Finish smoking men 1001 Irish/American men 356 U.29 0.0 1.26 Wolk et al.05 <.88 0.16 Rimm et al.S. men and women 5754 Scottish men 5875 Scottish women 68.0 1. Wolk et al. Furthermore. These authors found a reduced rate of death with increasing quintile of cereal fiber intake that reached a 26% reduction in risk for men in the highest quintile (consuming an average of 26 g of cereal fiber per day) relative to those in the lowest quintile of intake (consuming an average of 9 g per day). Also. Todd et al.23 reported particularly robust associations between intake of cereal fiber and CHD risk in a large cohort (n = 68. May 6. with more statistical power due to a larger number of cases. In comparison to women in the lowest quintile of intake.7 g per day — similar to the lowest quintile from the ATBC Study. with some of the association being explained by intake of antioxidant vitamins.S. fruit. Alternatively.2. suggesting that it may not be the fiber in fruit that is responsible for reducing risk. women in the highest quintile were observed to have a relative risk of CHD of 0.1.833 women and 4.20 reported findings from the Health Professionals Follow-up Study in the U. and fruit fiber intake among 21.2387_ch6. 2001 7:47 PM WHOLE GRAINS. were statistically significant for each of the second through fourth quartiles of fiber intake when compared to the lowest quartile. the amount of fiber obtained from fruit may be insufficient in most diets to observe an association with disease risk. fruit. CEREAL FIBER. the average intake in this quintile was only 9. it was estimated that risk of heart disease . caution should be used in such endeavors for cultural and methodologic reasons. indicating a 34% reduction in risk of coronary heart disease. From the Scottish Heart Health Study. Finnish smoking men generally eat few fruits and vegetables and a lot of whole grain rye bread. the protective associations were entirely explained by antioxidant intake. Although cross-cultural comparisons are of interest in evaluating consistency among studies. because fruits and vegetables contain a great deal of water and relatively little fiber per unit volume. Shown in Table 6.28 reported the relative risk of CHD incidence for men and women. For example.66. was relatively similar across food groups (vegetable. with a reduced risk of 29% (95% confidence interval = 46% to 8%) in the highest quintile of intake relative to the lowest quintile. vegetable fiber. and this may partly explain why findings were more robust for cereal fiber. very similar to those just described from the ATBC Study. these authors observed a consistent decrease in CHD risk with increasing quartile of dietary fiber intake that reached a magnitude of 44% reduced risk for the highest quartile of women.782) of women from the Nurses Health Study.930 male Finnish smokers enrolled in the ATBC Study.25 The findings for men.2 are the relative risk estimates from this study from a regression model which simultaneously included cereal. and vegetable fiber and a long list of potential confounding variables. This possible gender difference is consistent with the findings of Khaw and Barrett-Connor. The distribution of dietary fiber intake. such as male smokers in Finland. in this population intake of cereal fiber was much greater than for fiber from vegetables and fruits. the reduced risk across quintiles was graded and appeared to be stronger than the association in men. and the method of quantifying dietary intake in the ATBC Study is much more comprehensive than the food frequency questionnaire used in the HPFS Study. Following this line of thought.27 compared the rate of CHD deaths across quintiles of cereal fiber. Rimm et al. Interestingly. unlike the ATBC Study. Although the confidence interval around this estimate was wide and not statistically significant.2. AND CHRONIC DISEASES: EPIDEMIOLOGIC EVIDENCE 469 Pietinen et al. Although the sample size (n = 3.036 men available for analysis) and number of events (n = 97 women) were small in comparison to the other studies in Table 6. the findings were strongest and statistically significant for cereal fiber. findings were weaker and not statistically significant when the endpoint used was all major coronary events (fatal and nonfatal). although total intake appeared to be lower than in the ATBC Study. Although all three types of fiber appeared to reduce risk for myocardial infarction. A possible explanation for this discrepancy is that dietary fiber intake reduces the case-fatality rate in men with coronary disease and has less impact on the incidence of coronary disease in a population at high risk. Findings for vegetable fiber appeared to be somewhat weaker. Although fruit fiber also appeared to be protective for coronary death.2_fm Page 469 Sunday. for every increase of 5 g of cereal fiber intake. The authors did note that age-adjusted findings were stronger for fatal than for non-fatal myocardial infarction but that fatal and nonfatal categories were grouped together due to a limited number of cases. and cereal). 29 had made simple observations regarding specific whole grain breads.5 expanded these analyses to include all CVD deaths (as well as deaths from cancers and all causes). Morris and colleagues described the relative contribution of various grain products to dietary cereal fiber intake and their qualitative association with CHD risk. is apparent. no association was found between intake of other grains and white bread and CHD incidence. From the Seventh-Day Adventists Study. Brown bread and some whole meal breads and breakfast cereals were found to be strongly associated with cereal fiber intake and were therefore presumed to explain the robust association between intake of cereal fiber and CHD risk...2. 47 g per day).2. in 75. Recently.20 and Pietinen et al.. Whereas the previous studies2. fruits and berries. In comparison. While rye products were associated with a reduced risk for CHD death of 25% in the highest (172 g per day) in comparison to the lowest (16 g per day) quintile.30 published studies of whole grain intake and mortality from cardiovascular disease and other causes.2. The consistent reduction in risk for cardiovascular disease incidence or mortality.” These findings are summarized in Table 6. and other cereal products. is displayed in Table 6. This association was attenuated but could not be entirely explained by adjustment for selected constituents of whole grain (dietary fiber. In their study of fiber and coronary disease incidence in the ATBC Study.521 women in the Nurses Health Study for endpoints of coronary disease31 and incident ischemic stroke. no such apparent protection was observed for other cereal products (215 vs.29 supported the earlier findings by Morris et al. However.27. although this finding was not statistically significant. Using data on postmenopausal women from the Iowa Women’s Health Study. vitamin E. 3RD EDITION was reduced by 37% (95% confidence interval = 51% to 19%). for which the same dietary questionnaire and analytical methods were used. 2001 7:47 PM 470 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. folic acid. Pietinen et al. magnesium. whereas vegetables. May 6. and rye products in particular were all associated with reduced risk for coronary death in a dose–response manner.23 Table 6. whereas no such protection was observed for refined grain intake.32 using the same food groups for whole and refined grain. observations from epidemiologic cohort studies reported over the past few years are novel and important in two respects: (1) whole grain foods appear to offer protection from .31 was observed for whole grain breakfast cereals. phytic acid.32 Most of the protective association of whole grain intake with CHD risk in the study by Liu et al. Reductions in risk for CHD mortality were observed within increasing whole grain intake. brown rice. primarily consumed in whole grain form in Finland.27 the corresponding relative risks for fruit and vegetable fiber were weaker and not statistically significant in the Nurses Health Study.3 for this study.3 describes findings from prospective epidemiologic studies of whole and refined grain intake and risk for coronary heart disease as well as total cardiovascular disease. The only food group that appeared to offer protection from CHD incidence was vegetables. Jacobs et al. Women who reported consuming more than one serving of whole grains per day had a 30% lower risk (95% confidence interval = 50% to 2%) of ischemic heart disease death than those women who reported one-half or fewer servings per day. These findings were confirmed by Liu et al. which probably included a lot of refined grain foods.2_fm Page 470 Sunday.2 demonstrating a 43% reduction in risk of CHD for men and women who endorsed whole grain bread intake relative to those endorsing white bread consumption.31.3.5.30 considerable CHD risk reduction was also observed specifically for intake of “dark bread. and manganese).30 developed an algorithm for breakfast cereals according to whole grain content described earlier and incorporated these cereals along with breads and many other types of whole grain foods into food groups of whole and refined grains. The contrast between rye products. Jacobs et al.2387_ch6. In summary. Fraser et al. Jacobs et al. 10 g of cereal fiber per day appeared to be necessary to reduce the risk of CHD by a similar amount in the men of the Health Professionals Follow-up Study. as well as the lack of any association between refined grain intake and these endpoints.5..27 examined dietary intake of various food groups and food sources which contribute fiber as well as many other nutrients. whereas in the study by Jacobs et al. and bran. Those endorsing mixed grain breads also appeared to have a reduction in risk of 41% compared to white bread eaters.20 Consistent with the findings of Rimm et al. iron. 0 1.0 1. Largest differences across tertiles of cereal fiber intake were observed for brown and whole meal breads and breakfast cereals. AND CHRONIC DISEASES: EPIDEMIOLOGIC EVIDENCE 471 .79 1.96 0.78 0.86 0.03 0.097 CVD deaths over 9 years 761 CHD cases over 10 years 352 ischemic stroke cases over 12 years 134 CHD cases 45 CHD cases over 20 years Outcome Quintiles of whole and refined grain intake Quintiles of whole grain intake Rye products Other cereal products Quintiles of whole and refined grain intake Quintiles of whole and refined grain fiber intake Whole wheat and mixed grain bread relative to white bread Intake of brown and whole meal breads.94 0. 1.02 .32 Liu et al.75 0.75 1.0 1. 2001 7:47 PM WHOLE GRAINS.57 . Cases = fatal + nonfatal events.740 U.93 0.02 .64 1.69 0.72 1.59 NS 1 Note: All studies adjusted for potentially confounding variables.56 <.93 0.82 1.521 U.04 0.01 1. female nurses 75.0 1. Study Table 6. breakfast cereals.0 1.58 .30 31.31 Jacobs et al.09 .930 Finnish smoking men 34. and white bread Exposure Whole Refined Whole Whole Refined Whole Refined Whole Mixed p-Value 1.01 .18 0.29 Morris et al.94 0.83 0.78 1.02 . May 6. Seventh-Day Adventists 21.S. p-values are for tests of linear trend for relative risks across categories of intake for studies with three or more categories. body mass. CHD = coronary heart disease. CEREAL FIBER.S.2.14 0. female nurses 635 CHD deaths over 6 years 438 CHD deaths over 9 years 1.12 0. cakes.0 0.97 0.0 1.92 0. No association was observed between white bread and other refined carbohydrates and CHD incidence.5 Jacobs et al.60 0.99 1.85 1.521 U. including demographics. women 337 English men Sample Relative Risks per Increasing Category of Intake 2 3 4 5 Prospective Epidemiologic Studies of Whole and Refined Grain Intake and Risk of Cardiovascular Disease Pietinen et al.71 1.87 0.05 0.2_fm Page 471 Sunday.83 .03 0.27 Fraser et al. Liu et al.11 0. CVD = cardiovascular disease (heart disease + stroke and other events).208 U.S.08 .08 0. and lifestyle factors.3 2387_ch6.0 0.96 0.492 U.70 1.70 Analyses were described qualitatively. biscuits.0 1.S.0 0.2 75. other dietary factors. women 38.01 1.S.0 1. Therefore. Type 2 Diabetes As discussed above. may be associated with coronary heart disease more consistently and strongly than fiber from fruits and vegetables. Meyer et al. The results from the Nurses Health Study37 suggested that cereal fiber and the dietary glycemic load were additive in their association with type 2 diabetes. with fiber intake appearing to protect women from type 2 diabetes at all levels of the glycemic load.2387_ch6. These recent observations may have great public health importance because of the rising incidence of type 2 diabetes in the Westernized world.36 However.2_fm Page 472 Sunday. cardiovascular disease and type 2 diabetes share common etiologies in which diet appears to play an important role.35 An early study reported no differences in dietary fiber intake or foods providing dietary fiber between women who did and did not develop type 2 diabetes in the future. Findings from the Nurses Health Study. An interesting aspect of these studies was that the association between cereal fiber and type 2 diabetes incidence was independent of the dietary glycemic index or glycemic load. The stronger associations of cereal fiber.37 the Health Professionals Follow-up Study. coming primarily from whole grain foods.4). Although the Health Professionals Follow-Up Study confirmed this additive finding for cereal fiber and glycemic load on risk of type 2 diabetes. 2001 7:47 PM 472 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.5 times more likely to develop diabetes over the 6-year period in comparison to women in the highest tertile of cereal fiber and lowest tertile of glycemic load.38 and the Iowa Women's Health Study39 have revealed that cereal fiber. is inversely associated with risk of type 2 diabetes in a dose–response manner in both women and men (Table 6. and Finland had suggested that cereal fiber. could possibly be due to a synergy of nutrients in the bran and germ that cannot be explained by quantifying the fiber itself or a few common nutrients in the diet. The magnitude of the risk reduction observed for the highest quintile of cereal fiber intake (median intake of ~7–10 g/day) was 30–35% relative to the lowest quintile (median intake ~2–4 g/day). However. May 6.39 observed no association between glycemic load or glycemic index and incidence of type 2 diabetes in the Iowa Women’s Health Study. 3RD EDITION cardiovascular disease above and beyond the effects that may be explained by fiber or dietary fat. which is an estimate of potential for the diet to increase blood glucose acutely. the apparent protection conferred by whole grain foods was observed for virtually every whole grain food item . it is not surprising that recent epidemiological evidence from prospective studies suggests that intake of dietary fiber.39 and an additional report from the Nurses Health Study40 have confirmed that whole grain intake is inversely associated with risk of type 2 diabetes. given the evidence described above for cereal fiber. and especially cereal fiber and whole grain foods. Previous studies in the U.S. but not fruit and vegetable fiber.2. this study used a single 24-hour recall of diet. This risk reduction was approximately 20%39 and 25%40 for women in the highest quintile of whole grain intake in comparison to those in the lowest quintile in these two studies. In the Nurses Health Study. if real. women in the lowest tertile of cereal fiber intake and the highest tertile of glycemic load were 2. whole grains. and (2) the benefits of dietary fiber and whole grain foods have been observed in women with magnitudes of association that may be stronger than those in men. there may be many yet-to-be identified biologically active constituents in these parts of the plant that tend to be retained in whole grain foods. is inversely associated with risk for developing type 2 diabetes. and heart disease risk. which is known to be a poor measure of habitual dietary intake and would have considerably biased any real association toward the null hypothesis. the study by Meyer et al. the nature of the joint associations of cereal fiber and glycemic load on risk for type 2 diabetes is not clear from these three studies. whereas glycemic load had no association with diabetes in those men in the top two tertiles of cereal fiber intake. Finally. while refined grain intake is not associated with risk of type 2 diabetes. primarily through amount and type of carbohydrate reported.33 As discussed in the chapter on mechanisms and experimental evidence. it also appeared that cereal fiber had no association with risk for type 2 diabetes in those men in the lowest tertile of glycemic load.34. 94 1.87 1. other dietary factors.36 <.0 1.07 0.26 Relative Risks per Increasing Category of Intake 1 2 3 4 5 p-Value Note: All studies adjusted for potentially confounding variables.37 Meyer et al.01 .93 0.72 0.521 U. body mass. female nurses 35. p-values are for tests of linear trend for relative risks across categories of intake.11 .38 Study Table 6.0 1. female nurses Salmeron et al.97 0. Whole Grain Foods.39 Liu et al.87 1.01 .98 1.0 1.85 0. 42.01 .759 U.87 0.06 1.96 0.94 1.17 0.173 U.95 1.14 1.02 0. men Sample Prospective Epidemiologic Studies of Dietary Fiber.12 1. AND CHRONIC DISEASES: EPIDEMIOLOGIC EVIDENCE 473 .75 1.74 1. May 6.0 1.08 .92 0.98 0.988 U.01 1. women 75.65 <.S.91 1.12 0.40 0.0 1.0 1.99 0.14 1.S.0 1. and lifestyle factors.23 0.141 cases over 11 years 915 cases over 6 years 523 cases over 6 years Outcome Quintiles of whole and refined grain intake Quintiles of fiber and whole and refined grain intake Quintiles of cereal fiber intake Quintiles of fiber intake Exposure Cereal Fruit Vegetable Cereal Fruit Vegetable Cereal Fruit Vegetable Whole Refined Whole Refined 1. including demographics.2.4 2387_ch6.40 1.01 1.01 .2_fm Page 473 Sunday. CEREAL FIBER.82 0.10 0.S.77 .22 0.39 .88 1.65.10 0.70 1.91 1.12 0.01 .29 0.879 cases over 10 years 1.0 1.01 0.79 0.98 0.S.10 0.77 1.0 1.64 1.0 1.12 0.0 1.89 1. 2001 7:47 PM WHOLE GRAINS.0 1.81 0. and Risk of Type 2 Diabetes Salmeron et al.54 <.95 0.68 .27 0.14 1.0 1. brown rice. It is the main exposure of interest that would be most susceptible to recall or interviewer bias in these studies.39 This is not to say that there aren’t other possible mechanisms or constituents of whole grains for which cereal fiber and magnesium serve as markers in epidemiologic data.0 for whole grain eaters in comparison to non-eaters for 17 of 18 cancers (the one exception being thyroid cancer).50 Using data from an integrated series of case-control studies in northern Italy from 1983 to 1996. popcorn. because dietary fiber has been thought to reduce risk for colorectal cancer through actions in the colon that are described in the accompanying chapter. Jacobs et al. cereal fiber and dietary magnesium intake appeared to explain most of the protective associations between whole grain intake and risk of type 2 diabetes.012 adenomas of the distal colon and rectum were identified by endoscopy. found no association between dietary fiber intake and the incidence of colorectal cancer in 88. Therefore. The most rigorous prospective epidemiologic analysis on this topic to date.40 In the Iowa Women’s Health Study.3.66. The pooled odds ratio for all cancers was 0. These results were similar after excluding six instances of design/reporting flaws or low intake and were also similar when examined in studies that adjusted for the most covariates. A meta-analysis by Chatenoud et al. As described earlier in this chapter. Cancer Studies of dietary fiber and cancer have revealed mixed findings. They examined types of fiber (fruit. wheat germ. 3RD EDITION (dark bread. This finding is probably not explained by inaccuracies in dietary reporting because.50 took advantage of existing dietary data in the published reports where whole grain was not a main focus of any of the studies. The reader is referred to Chapter 6. cooked oatmeal. non-neoplastic . most of the case-control studies included in this meta-analysis did not have a hypothesis about whole grain intake or a specific hypothesis about diet in general. bran. A further strength is the inclusion of many types of cancers in this meta-analysis. Case-control and prospective studies of dietary fiber and colorectal cancer have been inconsistent. which is important to remember when interpreting studies because most of the studies on fiber and chronic diseases have adjusted the associations for body mass index. and other whole grains).49 These authors attempted to uncover many methodologic barriers that may have masked a real association. vegetable. there have been strong associations between cereal fiber and whole grains with risk of heart disease and type 2 diabetes in this same study. which provides insight into the experimental evidence on this important topic.50 revealed that individuals who reported habitual diets including whole grain foods were at significantly reduced odds of having 18 types of cancer. and cereal) and many subgroups of women. these authors observed odds ratios less than 1. May 6. as discussed above. or a 34% reduction in odds (95% confidence interval = 40% to 28%) for whole grain eaters compared to non-eaters. from the Nurses Health Study. Each of these factors may be on the causal pathway between whole grain intake and glucose control. Whole grain intake was observed to reduce the odds of having colorectal cancer by 21% (95% confidence interval = 31% to 11%). 2001 7:47 PM 474 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.757 women followed for 16 years. but no association emerged between intake of fiber and risk for colorectal cancer. Therefore.2_fm Page 474 Sunday. The findings from the meta-analysis of Jacobs et al. and body mass index.51 revealed findings very similar to those reported by Jacobs et al. Pereira et al. this area of nutrition research remains very controversial. in which 787 cases of colorectal cancer were documented and 1.41–49 Few studies have explored other types of cancer. whole grain breakfast cereal. Jacobs and colleagues have conducted a meta-analysis of 40 case-control studies of cancer in order to estimate the odds of having various cancers contingent upon dietary whole grain intake.8 observed an inverse association between whole grain intake and fasting insulin concentration in young adult men and women that was explained by fiber intake. Until recently there has been little development of biological plausibility for fiber and other cancers.2387_ch6. It is important to note that the authors selected 7990 controls who were admitted to the hospitals for acute. magnesium intake.50 Although case-control studies of diet and cancer may be particularly susceptible to recall bias. such as weeks. be true even if there is no benefit from these foods for cancer risk. Certainly there have been secular trends toward improving diet in the U. and Minnesota to examine associations between dietary eating patterns and risk of developing colon cancer among 1993 cases and 2410 controls.53 used data from a population-based study conducted in Northern California. intermediate. 30% reduction in older women. tobacco use. was associated with higher levels of vigorous leisure-time physical activity. there are many constituents. If cereal fiber really does protect against colorectal cancers.52 Dietary fiber intake was associated with a 50% reduction in the odds of having colon cancer for older men. findings of cereal fiber. and therefore foods providing fiber. such that diet as reported for the past month or year may not reflect the habitual diet of 5 years or 10 years or longer. whole grains.53 In summary. have more accurate reports of long-term diet. with the clearest evidence for a potentially protective association coming from casecontrol studies of whole grains. May 6. there is a need to examine the association between whole grain intake and risk for colorectal cancer prospectively. one would expect that studies which have included all-cause mortality as an endpoint would reveal that dietary fiber. or alcohol. 47% reduction in odds of colon cancer. over the past few decades. 45% reduction in odds.2_fm Page 475 Sunday. in fact. AND CHRONIC DISEASES: EPIDEMIOLOGIC EVIDENCE 475 conditions. 2001 7:47 PM WHOLE GRAINS. women. Slattery et al. unlikely to have been caused by diet. CEREAL FIBER.52 Among men. These observations were strengthened by the documentation of a dose–response association in comparing odds of cancers across low. 95% confidence interval = 62%–13%) and among people with proximal tumors (men. examined the association between types of dietary fiber and whole grain foods in association with the risk of all-cause mortality. Total Mortality Given the epidemiologic studies reviewed in the previous sections. The . Some of the studies described above have. as well as the animal experiments and a few randomized trials in humans on this topic.3 discusses in greater detail these possible mechanisms. and risk of colorectal cancer have been inconsistent. Only fiber and colorectal cancer have been examined in prospective studies. Utah. Risk factors for cardiovascular disease and type 2 diabetes may be modified over relatively short periods of time. nutrients. 95% confidence interval = 57%–8%. than prospective studies which may span only a decade or two at most. Perhaps case-control studies. with the strongest associations being observed among people diagnosed prior to age 67 years (men. why hasn’t this been borne out in prospective studies as it has for heart disease and type 2 diabetes? There may be etiological as well as methodologic reasons for this discrepancy.9 That the case-control metaanalyses by Jacobs et al.50 and Chatenoud et al. Total fruit intake was not associated with colon cancer risk. would appear to extend life.52 A “prudent” dietary pattern. 42% reduction in odds. 95% confidence interval = 62%–20%.2387_ch6. Perhaps most importantly. higher levels of whole grain intake were associated with a 40% reduction in the odds of having colon cancer (95% confidence interval. and 50% reduction for women with proximal tumors. and phytochemicals present in whole grain foods that may offer protection from colorectal cancer. because the inverse associations with cardiovascular disease and type 2 diabetes are quite consistent across various populations with different study methods.51 both demonstrate decreased risk for many types of cancer in addition to colorectal provides evidence that whole grains are offering much more than just cereal fiber.S. whereas slow-growing colon polyps may be less mutable and may require longer periods of dietary stimulus to affect their growth and malignant potential. women.52. Colorectal cancer may depart from heart disease and type 2 diabetes in its long latency period. identified by factor analysis. and high consumption of whole grain intake. 36% reduction in odds. potentially including early adulthood. smaller body size. Beyond fiber.53 This dietary pattern appeared to confer protection from colon cancer. 40% reduction for men with proximal tumors. of course. dietary fiber. and higher intakes of dietary fiber and folate. This would. 60%–10%). although susceptible to recall bias and interviewer bias (important issues discussed in some detail above). Chapter 6. 95% confidence interval = 55%–8%). and cancer. we submit that. this trial used a unique approach: partitioning cereal fiber eaters into those consuming cereal fiber from refined grains vs. may improve the quality (delaying disease incidence) and the quantity (delaying death. and cereal fiber with risk for coronary disease. and perhaps whole grains. with one group consuming predominantly whole grain cereal fiber (71% of cereal fiber. consumption of fruit and vegetables has if anything increased. A subset of 11. and since we ourselves are only now and painfully learning about “fibre”. we shall not speculate.33 From 1986 to the end of 1997. POTENTIAL APPLICATIONS. then why would it matter if the fiber came from the starchy nutrient-poor endosperm exclusively (refined grain) or also from the nutrient-packed bran and germ (whole grain)? As discussed earlier.8 g/day). vegetable. those consuming cereal fiber from whole grains. One might ask that. after adjustment for potentially confounding factors including demographics. death rates from coronary disease and other cardiovascular diseases as well as cancer also appeared to be lower for the whole grain fiber consumers compared to their refined-grainconsuming counterparts. and chronic disease risk? Do we now know. 3RD EDITION findings have been generally consistent.and nutrient-dense structures of certain plants. Recently. alcohol use. So. the link should be with the fibre in cereals. in fact. so different physiological effects are to be expected. other dietary factors. The results of this study were very enlightening and. physical activity. fiber and fiber-rich foods have been more closely examined in studies examining cardiovascular disease incidence in older women. especially due to cardiovascular diseases) of life.040 women was selected on their similar consumption of cereal fiber (total = 5. AND FUTURE RESEARCH NEEDS What have we learned by reviewing the epidemiology of dietary fiber.31 in that fiber itself does not explain all of the association between whole grains and risk for CVD.33 Although this subset analysis had limited statistical precision to evaluate cause-specific mortality. Taken together. the epidemiologic findings for disease incidence and for death suggest that a diet rich in minimally or appropriately processed plant foods. To those who wilt at the possibility of yet another behavioural risk factor for CHD it may be said that what is presently known by no means accounts for the occurrence of heart attack . May 6. Those reporting consumption of mostly whole grain fiber had an average death rate that was 17% lower (95% confidence interval = 27% to 6%) than those consuming mostly refined grain fiber. meanwhile. may be particularly efficacious for controlling cardiovascular disease risk factor evolution. It is the intake of that which has fallen. the observation makes sense: if dietary fibre is related to the “modern epidemic” of CHD.2387_ch6. no mechanisms can be postulated of the present observation (which could be tested fairly quickly).30 and Liu et al.… The physical and chemical properties of cereal and fruit and vegetable fibre differ substantially. although not as strong as for cardiovascular disease and type 2 diabetes. with this wealth of data from studies around the world. while fiber itself may have many important physiologic effects that may modulate risk for coronary disease. potatoes apart. body habitus measures. CONCLUSIONS. the fiber. n = 3559). fiber-rich foods. confirmed the findings of Jacobs et al. what Morris and his colleagues2 felt that they knew 23 years ago? The finding on cereal fibre illustrates one of the uses of epidemiology — in exploration.2_fm Page 476 Sunday. if the previous studies were able to explain all of the associations between whole grain intake and coronary disease by fiber intake. 1341 deaths were observed among these women.… The linoleate is only one possibility that cereal fibre may be the vehicle for other effectual nutrients. 2001 7:47 PM 476 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.5. Historically. In the Iowa Women’s Health Study. smoking. and hormone replacement therapy. we asked: “Does the association between cereal fiber and risk for disease depend on its source?” While this issue has been partly addressed in studies evaluating associations of fruit. such as the germ and bran of grains. n = 7481) and the other consuming predominantly refined grain cereal fiber (77% of cereal fiber. Results have been inconsistent. January 29. J. Marr. We have also. 1998. BMJ. M. 1974. and.2387_ch6. R. D. 231.. Consumption of grain and whole-grain foods by an American population during the years 1990 to 1992. 37.S... Nutr. with heart disease. 1992. Am. and the progression of cancer may be less mutable in response to behavioral modification. and Allshouse. 1998. 1997. AND CHRONIC DISEASES: EPIDEMIOLOGIC EVIDENCE 477 and immunity from it. May 6. Habitual dietary intake and insulin sensitivity in lean and obese adults. 1595. As such... E. D. D... Oxford University Press. G. Department of Health and Human Services. 1068. 70. Clin. but it is difficult to make recommendations specific to cancer when benefit has not been consistently observed in observational studies of free-living humans. 95. S. and expenditures. Cancer may be more etiologically heterogenous than cardiovascular disease and type 2 diabetes. Diet and physical activity as determinants of hyperinsulinemia: the Zutphen Elderly Study.. U.. 5.. and cereal fiber and whole grain foods in particular. to promote the consumption of foods rich in dietary fiber — especially whole grains. J. 55. 1994.” (FDA Docket #99P-2209). MC92-1-20D.. 1998.. Nutritional Epidemiology. etiologically. Food and Drug Admistration allowed the following health claim: “Diets rich in whole-grain foods and other plant foods and low in total fat. 2. 2. Pub. Jr.. Office of Disease Prevention and Health Promotion).. R. Jacobs.2 So what is new in 2001? We have confirmed Morris’s findings2 for coronary disease and extended them to the newer epidemic of type 2 diabetes — a disease with much in common. JAMA. Food consumption. Jr. Risk for type 2 diabetes and coronary disease. 12. J. S. J. prices. Grain Mill Products. that this one would be easier to put right than some. W. J. Diet. Am. et al. M. C. And we are setting population goals to strive toward increased whole grain consumption (Healthy People 2010.S. Whole-grain intake and cancer: an expanded review and meta-analysis. without sufficient support for either increased or decreased risk for cancer according to intake of dietary fiber. on the other hand. Diet and heart: a postscript. may reduce risk for these diseases. Am. 1307.. we are making progress toward changing dietary patterns in the population by specifying whole grain intake rather than simply total grain intake. Burkitt. et al. New York. Epidemiol. Diabetes. J. 8. Plausible mechanisms for the protectiveness of whole grains. 85. 4. J. et al.2 in conducting experiments in animals and humans to identify many promising and possible biological mechanisms through which dietary fiber. 30. Walker A. saturated fat and cholesterol may reduce the risk for heart disease and certain cancers. 1988. 13. 10. CEREAL FIBER.. USDA Economic Services Report. which has lagged far behind the promotion of fruit and vegetable intake.2_fm Page 477 Sunday. The association of whole grain intake and fasting insulin in a biracial cohort of young adults: the CARDIA Study. Lovejoy. in 1999 the U. J. and Kromhout.. 1. N. Is whole grain intake associated with reduced total and cause-specific death rates in older women? The Iowa Women’s Health Study. Personal communication. 1999. 2001 7:47 PM WHOLE GRAINS. P. Nutr. Role of insulin resistance in human disease. While the jury is still out on cancer. 2nd ed.. 1995. . and Tobelmann. 350. USDA. J. Loeber. J. 1977. L... 1970-94. J. 3. The potential mechanisms are many. Morris. C. Slavin. Albertson. heeded the call of Morris et al. Pereira. D. CVD Prevention. Annual data. Dietary fiber and disease. W. 1174.. based on Census of Manufacturers. Putnam... 1999. 1996. Willett. makes good sense in itself. 6. E. M.. et al. J. J. Feskens. Statistical Bulletin #928. Clin. J. 229. REFERENCES 1. Jacobs D. Nutr. as in so much of today’s health and prevention. A. Assoc.. Am. primarily on the part of government and industry.. 703.3. and DiGiroloama. Health. findings for cardiovascular disease and diabetes and total mortality are sufficiently strong to warrant continued public health efforts. G. Am. and Painter N. Cancer. Finally. 89. M.. 7. Industry Series. 140. R. 11. may be effected in weeks or less in response to diet or exercise. and Clayton. J. A. as described in Chapter 6. Reaven. 322. Gerrior. 459S. One area of continued controversy is the role of dietary fiber in the etiology of colorectal cancer. 9. . 277. 19. D. R. J. Med. 284. Jr. Nutr. A. Am. Dietary habits and incidence of noninsulin-dependent diabetes mellitus in a population of women in Gothenburg. 282. The Ireland-Boston DietHeart Study. Epidemiol. and risk of noninsulin-dependent diabetes mellitus in men.. L. Clin. S. A possible protective effect of nut consumption on risk of coronary heart disease. JAMA. 708. A. P.. J. 1821.. Long-term intake of dietary fiber and decreased risk of coronary heart disease among women. 30. Intake of dietary fiber and risk of coronary heart disease in a cohort of Finnish Men. Dietary fiber. Nutr. 1534. 27. 313. 921. E.. Khaw.2_fm Page 478 Sunday. Clin... D. 1989. et al. 37. H. et al. Wolk. 876. and cereal fiber intake and risk of coronary heart disease among men... glycemic load. 94.. Clin. 1014. 1997. Marshall. S. 46. 35. Jr. et al. Dietary fiber. J. 568. et al. and Barrett-Connor. 248. 33. 1985. 1996. Rimm. 26. 39. et al.. 2000. 1997. Beta-Carotene Cancer Prevention Study. and Blood Institute Family Heart Study. L.. J. J. 152.. 1993. 1416. F. JAMA. J. and risk of noninsulin-dependent diabetes mellitus in women. et al. Dietary antioxidant vitamins and fiber in the etiology of cardiovascular disease and all-cause mortality: results from the Scottish Heart Health Study. J. Am. Engl. et al. 1998.. et al. 450. J. 78. Meyer. Sweden. is inversely associated with all-cause mortality in older women: the Iowa Women’s Health Study. P.. Am. Intern. Pietinen. H. 1999. T. Vitelli. 24. Arch. Diabetes Care. 1998. Diabetologia. Carbohydrates. J. Diet and 20-year mortality from coronary heart disease.. D. 2001 7:47 PM 478 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Dietary fiber and coronary heart disease in middle-aged hypercholesterolemic men. Dietary fiber and weight gain [letter to the editor]. 6. S. JAMA. Lundgren. Vegetable. et al. and Hamman. JAMA. 25. L.. A. Am. et al. J. 49. 18.. 32.. Burke. E. women. et al. K. Whole grain intake may reduce risk of coronary heart disease death in postmenopausal women: the Iowa Women’s Health Study. Am. 1998. 71. M. J.. 9. et al. 19. Lung. Todd.. Epidemiol. J. 1997. Pereira. et al. 68. 1409.. 1093. fruit. D. Dietary fiber and reduced ischemic heart disease mortality rates in men and women: a 12-year prospective study. I. JAMA. 16. . BMJ. M. R. Prev. 1073. 28.. A. J. Malarcher. Am. S. Pietinen. JAMA.. Dietary determinants of ischaemic heart disease in health conscious individuals. and Tyroler. Fiber from whole grains. Epidemiol.. Am.. 145.. 90. S... and incident type II diabetes mellitus in older women. 2720. 1987. J. 2000. Liu. 2000. 1450. Salmerón. Med. C. The Alpha-Tocopherol. Med. but not refined grains. J.. Mann. 2000. 283. 29. A. Nutr.. J. Djousse. 126. H. Dis. Association of dietary composition with fasting serum insulin level: the ARIC Study. 1997. Relationship between dietary fiber consumption and fibrinogen and plasminogen activator inhibitor type 1: the National Heart. Clin. et al. Fraser.S. 472. 1997. Am.. G. Ludwig. Am. A. 811. P.. E. 1996. 3RD EDITION 14. 326S. 1996. High saturated fat and low starch and fibre are associated with hyperinsulinemia in a non-diabetic population: the San Luis Valley Diabetes Study. A prospective study of whole-grain intake and risk of type II diabetes mellitus in U. The Alpha-Tocopherol. et al. S. Intake of fatty acids and risk of coronary heart disease in a cohort of Finnish men.. Bessesen. N. 22. A.. H. Am. 1999. Whole grain consumption and risk of ischemic stroke in women. 38.. 1999. weight gain. 84. Clin. Nutr. Jacobs D. L. 31. Trends in the prevalence and incidence of self-reported diabetes mellitus–US 1980–84.. Pub. et al. Nutr. MMWR. 70. Nutr. Coll. 31. et al.2387_ch6. 1998. Heart.. Humble. Beta-Carotene Cancer Prevention Study. 17. R. et al. 40. Am. 1997. Circulation. Health. Whole-grain consumption and risk of coronary heart disease: results from the Nurses’ Health Study. et al.. B. 281. 36. G.. 275. 412. Dietary fat and risk of coronary heart disease in men. 150. 1539. 21. J... Liu. K. Salmerón. 20. 40. 430.. May 6. 34.. 68. Ascherio. Kushi. 545. and Jacobs. Nutr. 1996. Med. 23. 159. Jacobs. Intern. et al. 447.. Metab. 1999. Centers for Disease Control and Prevention. 20. Liu.. and cardiovascular disease risk factors in young adults. et al. 15. 197. Ludwig. Dietary fiber. Cardiovasc. dietary fiber.. 194. J. 1999. Rapid rise in the incidence of type II diabetes from 1987 to 1996. Arch. glycemic load. 2000. et al. D. 1992. New Engl. D.2_fm Page 479 Sunday. Thun.. G. et al. Whole grains intake and cancer: an expanded review and meta-analysis. Dietary intake of fiber and decreased risk of cancers of the colon and rectum: evidence from the combined analysis of 13 case-control studies. 1998. Dietary fiber and the risk of colorectal cancer and adenoma in women. meat.. Fuchs et al. Natl. 54. et al.. Am. and Riboli. 95. CEREAL FIBER. 42. Cancer Inst. et al. 5. 53.2387_ch6. 1. 1997. 44. Giovannucci. Steinmetz. Nutr. Chatenoud. Epidemiol. et al.. 49. et al. 1992. M. 28.. et al. J. 8. E. 169. 51. 45. 47. Jacobs. 30.. 1997. and colon cancer in the Iowa Women's Health Study. M.. Cancer Causes Control. Whole grain food intake and cancer risk. 84..] 43. Slattery. 276. 1994. Cancer. 77. 1999. Plant foods and colon cancer: an assessment of specific foods and their related nutrients (United States). Prospective study of diet and female colorectal cancer: the New York University Women's Health Study.. 1998.. Kato.. 1994. et al. Am. L. 1989. R. A. Eating patterns and risk of colon cancer. Risk factors for fatal colon cancer in a large prospective study.. Intake of fat.. Vegetables. M. 1. J.. 4. Goldbohm. Epidemiology. Cancer. 385.. 85. Nutr. K. 1887. Howe... 139. L. May 6. 148. 1994. et al. E. R. R. J. Cancer. 1994. I. 44. et al. Prospective study on alcohol consumption and the risk of cancer of the colon and rectum in the Netherlands. Natl. 1491. and fiber in relation to risk of colon cancer in men.. 24. F. 2001 7:47 PM WHOLE GRAINS. Int. Friedenreich. Jr. Influence of methodologic factors in a pooled analysis of 13 case-control studies of colorectal cancer and dietary fiber. J. J. L. 340. J. 46. 2390. Slattery. Diet and colorectal cancer with special reference to fiber intake. Heilbrun. 575.. 1994. 66. Epidemiol. [Erratum. Epidemiology. L. . AND CHRONIC DISEASES: EPIDEMIOLOGIC EVIDENCE 479 41. Cancer Causes Control. Cancer.. 52.. M. J. et al. 50.. A. 5. C.. J. et al. K. 1998. fruit. 5. J. Int. 84. Brant. Cancer Res. Med. Cancer Inst. 48. 2001 7:47 PM .2387_ch6.2_fm Page 480 Sunday. May 6. 3_fm Page 481 Sunday. Len Marquart. http://www. with some recovery since then. David R. Dietary Guideline Advisory Committee Rationale. This chapter will provide a working definition of whole grains as well as explore several possible biological mechanisms linking whole grain intake with coronary heart disease (CHD). and Chronic Diseases: Experimental Evidence and Possible Biologic Mechanisms Joel J. Pins.. More research is warranted to understand how whole grain foods might produce health benefits. 2001 7:48 PM CHAPTER 6. Jacobs. Mark A. 29–30.S.50 © 2001 by CRC Press LLC 481 . 0-8493-2387-8/01/$0. Jr. the USDA Dietary Guidelines for Americans were amended to include specific mention of whole grains: “Eat a variety of grains daily. grain consumption in the U. Recent data indicate that grain intake currently composes about 25% of total energy consumption. fell throughout the 1900s until 1972. especially whole grains” (pp. Few clinical trials have been done investigating the causal relationships and the potential biological pathways by which whole grains might modify chronic disease and possibly mortality. but it likely is a more realistic model for testing outcomes. Historically.00+$1. Pereira.ars. and Joseph M.pdf ).2387_ch6.gov/dgac/dgac_ration. The current research on whole grain intake and health tends to be nonspecific as to mechanisms.4 The “whole food” approach does make studying biological mechanisms challenging.S. May 6. Keenan INTRODUCTION Despite the efforts of early health advocates. type II diabetes. with whole grain intake providing only 1% of our total energy. evidence continues to amass supporting the health benefits of increased whole grain intake.3 Whole Grains. Such methodology is built on many assumptions and fails to recognize the possible interactive and/or additive effects of all whole grain components. Based on the same evidence. The evidence was sufficiently strong that in 1999 the U. most of the research in this field has focused on the “magic bullet” approach — testing a single grain component rather than foods rich in whole grains. Cereal Fiber. Food and Drug Administration (FDA) allowed the following health claim: “Diets rich in whole-grain foods and other plant foods and low in total fat. saturated fat and cholesterol may reduce the risk for heart disease and certain cancers” (FDA Docket #99P-2209).1–3 Nevertheless.usda. and cancer. and phytochemicals are disproportionately concentrated in the bran and germ and are therefore lost in milling. and bran. most nutrients. modification of the harvested kernel occurs in almost all cases prior to human consumption. germ.2387_ch6. milling. the FDA has adopted a definition in the health claim mentioned above that whole grain foods must consist of at least 51% whole grain flour (containing bran. most of the research focuses on the lipid-modifying effects of soluble fibers. parboiling.8–10 Many of the protective effects of whole grains have been attributed to their dietary fiber content. Whole grains must then. 2001 7:48 PM 482 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. to increase shelf life and palatability. whole grain pasta. and corn.” 7 Furthermore. hypertension. with minor grains including oats. ground. In milling. flaking. Lipid-Modifying Effects Since the publication of the first oat-feeding trial in the early 1960s.5 Prior to processing. Examples of foods made with whole grain flours include some breakfast cereals. These fiber effects may be related to the nondigestible constituents themselves. The percentage of bran and germ varies between grains. 3RD EDITION WHOLE GRAINS: WHAT ARE THEY? In terms of production. .6 Based on this information. but the endosperm consistently represents a little more than 80% of the kernel. such as hemicellulose. wheat and rice account for over 50% of total grain production. heat extraction. Antioxidant intake has been shown to decrease the incidence of CHD by slowing the rate of low-density lipoprotein (LDL) oxidation and exhibiting other vaso-protective properties.3_fm Page 482 Sunday.S. the major cereal grains are wheat. rice. the beneficial cholesterollowering effects of soluble fiber have been well documented: especially among hyperlipidemics.5 Though minor in terms of pounds of production. barley. phytoestrogens. Unfortunately. by definition. the American Association of Cereal Chemists has created the following working definition: “Whole grains shall consist of the intact. Recent data indicate that refinement of whole grains results in significant reduction of the content of phenolic acids.” Whether the bran and the germ are removed. cracked. and a few commercial breads. grain foods that are unrefined (retain the bran and germ) can be classified as “whole. Refined flour products contribute a significant percentage of total carbohydrate intake in the U. dietary fibers have been shown to directly and indirectly affect other CVD-related factors such as hyperinsulinemia. More specifically. obesity. This section of the chapter will focus on the CHD-altering effects of the fiber and antioxidant components of whole grains. and bran layer. Additionally. dietary fiber. one of the more recently researched whole grain phytochemicals is a family of compounds named phenolic acids. and other techniques. in fact. Moreover. the bran and germ are stripped from the endosperm and the latter is ground into flour. cooking. Such processing includes. CORONARY HEART DISEASE As described in our accompanying chapter. hyperglycemia. whose principal anatomical components. and hemostatic factors. triticale. oats are generally eaten whole and contribute disproportionately to whole grain consumption. the starchy endosperm.5 For example. but is not limited to. sorghum. and millet. puffing. whole grains are also rich in antioxidants such as tocotrienols. germ. be minimally processed grains. numerous epidemiologic studies have demonstrated that increased whole grain consumption is associated with reduced CHD. May 6.5 In developed countries. the most common of the refinement processes listed above. and phenolic acids. and endosperm) by weight. extruding. germ. or flaked caryopsis. all grains are similar in structure — composed of an endosperm. rye. most grains undergo considerable refinement. but they are equally likely related to the many phytochemicals that are found intermixed with the nondigestible constituents. For example. are present in substantially the same relative proportions as they exist in the intact caryopsis. Whereas no lipid changes occurred during the wheat phase.17 The primary mechanism (see Figure 6.21 A final hypothesis is that decreased absorption of dietary fats and cholesterol due to increased intestinal viscosity by soluble fibers might result in serum cholesterol reduction. serum LDL cholesterol. The hypocholesterolemic effects of whole grains rich in dietary fibers and oligosaccharides appear to work via one or more of four primary pathways. Though not all fibers affect each of these pathways equally. and undigested starches in the large intestine by indigenous bacteria.3.3_fm Page 483 Sunday. which accounts for approximately 60% of the total fiber content. where they inhibit hepatic cholesterol synthesis by limiting the action of HMG-CoA reductase.1) by which some viscous fibers reduce serum cholesterol is by binding bile acids in the small intestine and thereby altering enterohepatic bile acid recycling. This effect was observed only in men. were observed in the rye phase. Leinonen et al.15 examined the lipid-lowering effects of unrefined rye bread — a rich source of dietary fiber including arabinoxylan. May 6. AND CHRONIC DISEASES 483 they reduce total and LDL cholesterol in serum. refined grains.16. Results from a recent trial suggest that inulin and oligofructose (the most common fructans) might independently improve serum lipid concentration.13 Isoflavones from soy foods have also been shown to beneficially affect serum lipids. Some of these SCFAs (primarily propionate) enter the system via portal circulation and are delivered to the liver. the total lipid-modifying effects of whole grains might be due to the combined effect of the fiber. hepatic cholesterol.19 A recent animal trial demonstrated increased bacterial propionate production after feeding of poorly digestible resistant starches. which the authors suggest may be due to the difference in total rye bread consumed by men vs. Consider. that hydrolyzed guar gum.20 Viscous soluble fibers have also been suggested to slow gastric emptying and the digestion/absorption of carbohydrates in the small intestine. During each trial phase. a viscous soluble fiber. as well as the mechanism by which they might do so. Such action results in the reduction of postprandial insulin and a decrease in the activity of HMG-CoA reductase. respectively. LDL and total cholesterol reductions of 12% and 10%. butyrate. has not yet been fully established.14 Analogous phytoestrogens in whole grains might then also produce beneficial effects on serum triglyceride and HDL cholesterol concentrations. very few human and animal trials have suggested that wheat and other cereal brans significantly reduce serum triglyceride concentrations. To do so. Whether cereal bran and phytoestrogens improve lipid profile. for example. the liver must either up-regulate endogenous sterol synthesis or increase LDL particle uptake from circulation. One of the by-products of this process is short-chain fatty acids (SCFAs). Unfortunately. which does not possess the viscous properties of unmodified guar gum. without lowering high-density lipoprotein (HDL) cholesterol or significantly altering serum triglyceride (Tg) concentrations. and hepatic triglyceride in rats. In a recent crossover clinical trial. because they resist digestion in the upper gastrointestinal tract and are fermented in the large intestine by indigenous bacteria. oligosaccharides classified as dietary soluble fibers. and propionate.2387_ch6. Hypotensive Effects The blood pressure–modifying effects of whole grains and their associated components have not been studied as extensively as their hypocholesterolemic effects. women. oligosaccharides. the healthy young participants were asked to consume either refined wheat or whole rye bread in addition to their normal diet.18 A second mechanism of action is stimulated by the fermentation of fibers. and non-viscous soluble fibers such as gum arabic are ineffective in reducing serum cholesterol. 2001 7:48 PM WHOLE GRAINS. CEREAL FIBER. Some epidemiologic studies . This decreases the bile acid pool and requires more endogenous hepatic bile acid synthesis. and resistant starches as well as the bran fraction and the phytoestrogens. very few clinical trials have been conducted directly testing the lipid-altering effects of whole vs. all mechanisms generally rely on the viscous nature of these fibers. oligosaccharides. including acetate.12 Additionally.22 As mentioned above. Elevated levels of propionic acid resulted in reduced serum total cholesterol.11 Whole grains are also rich in fructans. . Both conditions result in increased vasoconstriction and further sodium retention. Legend: + signifies an increase in the variable after the pathway has been initiated.1 Whole grains and CHD/diabetes. Vuksan et al.signifies a decrease in the variable after the pathway has been initiated. Limited clinical trials have been conducted with blood pressure as the primary outcome variable. 2001 7:48 PM 484 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. while other studies have failed to show an effect.Coagulation factors + Fibrinolysis + Serum HDL-C .Body weight .LDL oxidation + Hepatic lipoprotein uptake . In a follow-up trial.3.27 have reported that hyperinsulinemia causes increased renal tissue sodium reabsorption. and phytochemicals Colonic fermentation to SCFA Increased intraluminal meal viscosity Portal circulation . It has been proposed that the increase in insulin sensitivity induced by soluble fiber results in lower circulating insulin levels.HMG-CoA reductase .Serum triglycerides .4 mmHg (6.3_fm Page 484 Sunday.3. have suggested that increased dietary fiber intake is associated with a decreased incidence of hypertension as well as lower blood pressure. leading to increased blood pressure. Participants were crossed to a wheat fiber control after 3 weeks of intervention.Postprandial glucose and insulin + Satiety . micronutrients. enhances stimulation of the sympathetic nervous system.5 mmHg.9%). the soluble fiber–rich intervention reduced systolic blood pressure (sBP) 9. Keenan et al. our group demonstrated that whole grain oat-based cereals significantly reduced the need for antihypertensive medication compared to a refined grain cereal control.2387_ch6.25 The mechanism (see Figure 6.23 added a viscous soluble fiber (glucomannan) to the AHA Step 2 diet of type 2 diabetics. reduced insulin levels have been shown to result in improved endothelial cell function and reduced blood pressure. . when compared to refined wheat-based cereals.Free fatty acids + Insulin sensitivity .Serum insulin concentration Hepatocyte Effects + Glucose oxidation + Rate of bile acid excretion .5 mmHg and 5.Serum LDL-C + LDL particle size + Insulin clearance .24 reported that oat-based cereals — oats that are rich in the soluble fiber beta-glucan — reduced both sBP and diastolic blood pressure (dBP) by 7. using whole grains rich in fiber or fiber supplements as the sole intervention.Cholesterol synthesis Figure 6. Scherrer et al. theoretical model of mechanisms of action. 3RD EDITION Whole grains rich in dietary fiber. Compared to the control period. which subsequently increases plasma volume.Rate of gastric emptying and macronutrient absorption . In animal models. In a recent study among diabetics. and induces hypercatecholaminemia.26 Additionally. respectively.1) by which soluble fibers exert their hypotensive effect is poorly understood. May 6.Blood pressure Up-regulation of cell surface insulin receptors . e. Specifically.28 PAI-1 concentration was significantly reduced at weeks 2. Numerous trials to date have investigated the effects of dietary fibers or whole grains on satiety and/or body weight.2387_ch6. was elevated for a longer time after the high-soluble-fiber meal compared to the low-fiber meal. fibrinogen levels were consistently lower during all weeks of the feeding of whole vs. though these differences did not reach statistical significance.3. the effect of whole grains and dietary fiber on these risk factors is less clear. a recent trial investigating the satiating effects of a hydrolyzed guar gum (non-viscous fiber) found no effect on various measures of satiety. AND CHRONIC DISEASES 485 Antithrombotic Effects Various studies have noted a strong association between rates of coagulation and fibrinolysis and CHD. to an improvement in insulin sensitivity during the whole grain diet. Pereira et al.4 g/day) in addition to the normal diet of healthy volunteers.30 recently investigated the satiating effects of a psyllium supplement (7. up-regulation of cell surface insulin receptors or suppression of hepatic free fatty acid release). the biological effects of whole grains rich in soluble fibers might largely be attributed to their viscous nature and their slowing of gastric emptying.1) relating whole grain and dietary fiber consumption to greater feelings of fullness is likely related to the rate of gastric emptying as well as the rate of total carbohydrate digestion and absorption. 4. refined grains. and the non-grain part of the diets was matched between treatments. In a controlled feeding study of whole vs. cholecystokinin (CCK). At the end of the 4-week study. The mechanism (see Figure 6. even though both diet phases were designed to maintain constant body weight. there was a nonsignificant weight loss of 2 pounds in the whole grain group. the psyllium treatment reduced feelings of hunger by 13% and decreased energy intake by 17%. a refined grain wheat product.1 ng/ml.29 Body Weight Effect Excessive body fatness is a primary predictor of CHD as well as type 2 diabetes. Rigaud et al. refined grains. This effect appeared to be due. Additionally. and factor VIIc increase CHD risk. 2001 7:48 PM WHOLE GRAINS.3. and 6 during treatment with whole grains (mean difference = 28. Pins et al. p < .3_fm Page 485 Sunday. Bourdon et al.31 recently investigated the satiety effects of a whole grain barley product vs. rice.. Here again.001). Grain intake included 6 to 10 servings per day of breakfast cereal.32 reported that participants had a tendency to be less hungry between meals on the whole grain diet. participants in the whole grain group reported a much greater feeling of fullness and being less hungry. Additionally. CEREAL FIBER. there was a nonsignificant trend toward weight loss on the whole grain diet when compared to the refined grain diet. This finding might help explain the mechanism by which dietary fibers increase satiety and ultimately facilitate weight loss. . plasminogen activator inhibitor type 1 (PAI-1). bread. however. and various snacks. The washout period was 6 to 9 weeks.33 recently demonstrated that carbohydrate digestion and absorption were significantly slowed in response to a high-soluble-fiber meal. the condition may inhibit hepatic synthesis of certain coagulation factors. Compared to a placebo. May 6. Animal trials have indicated that increased dietary fiber intake might reduce both PAI-1 and fibrinogen.1) by which whole grains and dietary fibers increase fibrinolytic activity is not well elucidated. refined grain diets in 11 overweight insulinresistant adults. at least in part. The mechanism (see Figure 6. After 1 week on the whole grain supplement treatment. It has been proposed that regardless of the pathway to increased insulin sensitivity (i. Some have suggested that increased insulin sensitivity is pivotal in the relationship. which reduces the rate of gastric emptying. research suggests that increased concentrations of plasma fibrinogen. Our recently completed crossover feeding experiment compared whole vs. pasta. Additionally. This enzyme complex is responsible for the clearance of lipid hydroperoxides (i. as well as endotheliummodulated dilatation. 2001 7:48 PM 486 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 3RD EDITION Antioxidant Effects and Other Vasoprotective Effects Various experimental and clinical data support the hypothesis that modification (oxidation or glycation) of the LDL particle increases the atherogenicity of the particle and is a major step in CHD. which have been more extensively studied. decrease blood pressure. at least to the extent that isovolumic replacement of refined grain flour with whole grain flour leads to a 15–20% reduction in energy intake from the grain food. The mechanism (see Figure 6. which reduces risk for CHD. Nevertheless. In comparison to unmodified particles. allowing it to function as a co-factor for glutathione peroxidase. improve fibrinolysis. it is apparent that whole grains and their many constituents may reduce the risk of CHD via their effects on multiple variables.35–37 including vitamins. Whole grains also improve insulin sensitivity. plant lignans (secoisolariciresinol and matairesinol) are converted to mammalian lignans (enterolactone and enterodiol) via the action of endogenous gut bacteria.41. decreased platelet aggregation. exogenous estrogen has been shown to improve vascular response after an injury as well as vasodilatation. . In a recent animal trial. Vitamin E is incorporated into the LDL particle and can reduce oxidative modification of LDL via its activity as a reducing agent. yet limited information is available on food concentrations..34 Whole-grain foods are a rich source of antioxidants. The exact mechanisms.42 Whole grains are rich in folate. Moreover.45 In a more recent animal trial. May 6. improve antioxidant status.40 Homocysteine is another known risk factor for CHD and might be reduced by increased consumption of whole grains.3_fm Page 486 Sunday.e. data indicate that urinary excretion is associated with fiber intake.46 Rabbits in the treatment group experienced a significant reduction in various measures of lipid oxidation. it is well accepted that whole grains improve serum lipids. Various phytic acids have also been shown to exhibit antioxidant properties in vivo. need much further exploration. soy isoflavones were shown to significantly reduce the development of atherosclerosis in disease-free rabbits.44 It is known that once ingested. suggesting that they are located in the outer layer of the grain. via its functions as an antioxidant or in other capacities. Based on these data. which is the primary agent responsible for lowering homocysteine levels.S. decreased activation of protein kinase C. This aspect will be discussed in the next section on type 2 diabetes. Antioxidants are known to delay the onset of lipoprotein oxidative modification or at least slow the rate at which it occurs. specifically tocotrienols and (depending on the soil content) selenium. and various nonnutrients such as phenolic acids. as well as provide beneficial effects on eicosanoid metabolism. phytoestrogens.38 Whole grains are especially rich sources of vitamin E. has also been shown to modify other risk factors for CHD such as smooth muscle cell proliferation. Another important contributor to CHD is abnormal vascular reactivity.3. Some whole grain foods and all refined grain flours are now fortified with folate in the U. as well. malondialdehyde).43 Recent data indicate that some plant estrogens share various chemical properties with estradiol. The lignans of whole grains share chemical properties with soy isoflavones. oxidatively modified particles are known to be major contributors to plaque formation in the subendothelial space and subsequent CHD. Arterial response to acetylcholine was enhanced. and the extent to which they are operative.2387_ch6. as well as improve vascular reactivity. as well as 37% less cholesterol deposition in the aortic arch when compared to controls. trace minerals. which are known to be atherogenic.34 Vitamin E. and phytic acid. rhesus monkeys with known atherosclerotic disease were treated for 6 months with soy isoflavones. Whole grain may help in weight loss. Certain plant estrogens called lignans are found in whole grains.1) by which antioxidants protect against CHD is fairly well understood. However.39 Vitamin E also keeps selenium in the reduced state. in each case. glucomannan. other fibers have been investigated for their glucose. Only the intact oats resulted in a significant reduction in postprandial glucose and insulin compared to white bread. it is just as likely that an unknown component of whole grains or an unexpected interaction between grain components might be responsible for the reduced risk of diabetes observed in the epidemiologic studies.and insulin-modifying properties. Recent studies have shown that certain foods possess a low glycemic index. in our recent feeding trial. Reductions were .0% and 32. Braaten et al.2387_ch6. the lower glycemic index of foods that contain whole grain particles might partially explain the beneficial effects of these foods in terms of type 2 diabetes. was reported to improve long-term glycemic control.49. In another study. The following section will focus on the glycemic and insulinemic effects of whole grains as well as their fermentable components. May 6. a recent epidemiologic study47 (described in detail in the accompanying chapter) found that insoluble fiber was as predictive of reduced diabetes incidence as soluble fiber.48 Moreover. boiled rolled oats. Additionally.54 tested the effects of raw rolled oats. significantly reduced both glucose and insulin responses. that is. Dietary fiber-rich interventions lower serum insulin and glucose concentrations as well as improve insulin sensitivity. when added to wheat farina. This same group showed that both oat gum and oat bran.7%) was significantly reduced by the arabinoxlyan soluble fiber. more specifically. Of interest. Fourteen healthy individuals were asked to consume three isoenergetic breakfast meals — containing either 0.3 and 5. psyllium supplementation produced analogous reductions in all-day and postprandial glucose levels when added to the traditional diabetic diet in men with type 2 diabetes (n = 34). were one of the first to demonstrate a significant reduction in serum glucose and insulin when oat gum was added to a glucose load. reducing serum fructosamine levels by 5. For example. 6. The acute effects of soluble fibers on glucose and insulin have been fairly well established. AND CHRONIC DISEASES 487 TYPE 2 DIABETES As is the case with CHD. the area under the curve for glucose (20. soluble fibers have been shown to exert significant glycemic and insulinemic effects. CEREAL FIBER.23 Whole grains rich in total and soluble fiber can also improve glucose and insulin metabolism. respectively. a less frequently studied soluble fiber.9 mmol/l for participants on the 6 g and 12 g fiber plans. The arabinoxylan-rich meals reduced peak postprandial glucose from 7. 2001 7:48 PM WHOLE GRAINS.2% and 41.2 mmol/l to 6.7% after a 3-week intervention at 0. and boiled intact oat kernels vs. they have less postprandial effects toward glycemia and insulinemia. some of which was provided by whole grains.52 investigated the postprandial effects of arabinoxylan in a normoglycemic population. Whole grains and/or their components have been shown to exert a significant effect on glucose and insulin metabolism.53 In a third study.55 demonstrated significant glucose and insulin effects in a diabetic population after increasing dietary fiber intake to 25 g of soluble and 25 g of insoluble fiber daily.4%) and insulin (17. Lu et al. Thus. Chandalia et al. Such findings suggest that many “unknowns” might at least in part help to explain the beneficial effects of whole grains in terms of diabetes. Glycemic and Insulinemic Effects Due to their viscous nature and gastrointestinal effects.51 More recently. white bread in older men. or 12 g of soluble fiber — on nonconsecutive days after an overnight fast. Similarly. we noted a significant improvement in insulin sensitivity in response to a whole grain diet rich in foods with a moderate to high glycemic index. numerous plant lignans were just discovered in whole grain rye at many times the concentration of those traditionally thought to be the most prevalent of these phytoestrogens. the fermentable fiber content of whole grains. The majority of this effect might be due to fiber content or. as illustrated by recent trials.2).3_fm Page 487 Sunday. epidemiologic cohort studies have shown that whole grains reduce the risk of type 2 diabetes (see Chapter 6. Granfeldt et al.7 g per 100 kcal.50 More recent trials have studied the long-term glycemic effects of purified fibers. 65–67 Yet various animal experiments and epidemiologic studies suggest that whole grains provide significant protection against gastrointestinal cancers and possibly other systemic cancers as well. Reduced serum glucose levels decrease the amount of insulin needed to clear the glucose load. The first pathway has to do with the glycemic index of whole grain foods.3_fm Page 488 Sunday. thereby increasing insulin sensitivity. as well as 24-hour glucose (10%) and insulin (12%) areas under the curve. decrease free fatty acid release. 2001 7:48 PM 488 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. it is likely that other whole grain constituents or interactions between known and unknown constituents beneficially modify risk for diabetes via the mechanisms highlighted above or via other currently unexplored pathways.05). soluble fibers decrease the rate of gastric emptying and slow the digestion of macronutrients. again.57 One of the primary predictors of the glycemic index of a whole grain is the structure of food components and soluble fiber content.32 also tested the insulin-modifying effects of a whole grain vs. Two primary mechanisms (see Figure 6. a refined grain diet. The following section will look at the anticarcinogenic effects of various .58 Any processing.62 The second pathway involves the fermentation of soluble fibers.24 As described above. Pereira et al.61 Independent of their delivery medium.3. May 6. although most of the whole grain foods consumed were made from whole grain flour.and insulinimproving actions of whole grains rich in fiber. There is some evidence that hepatocytes. such as refinement. the treatment group experienced a significant reduction in the amount of insulin needed to clear a glucose load.56.3 g). with a moderate to high glycemic index. recently published a study showing that simply altering the structure of starch-rich foods (decreasing glycemic index) resulted in a 30% reduction in area-under-the-curve for glucose and insulin. However. and increase insulin clearance — all of which might improve insulin sensitivity. Results of euglycemic hyperinsulinemic clamp tests in 9 of the 11 subjects confirmed that the average insulin sensitivity of these subjects was improved by the whole grain diet. suggesting improved insulin sensitivity. CANCER It has long been thought that dietary fiber is very protective against the incidence of colon cancer. whole grains (not whole meal) tend to reduce the glycemic response. over time. when exposed to an increase in SCFAs. By 2 weeks into the whole grain diet. that disrupts the food structure and/or decreases the soluble fiber content results in increased glucose and insulin responses. producing SCFAs which then enter the portal circulation.2387_ch6. oligosaccharides. 3RD EDITION observed in mean daily glucose (13 mg/dl) and urinary glucose (1.64 In closing.63. Insulin levels remained suppressed and significantly lower than the refined grain diet through 6 weeks (p < . the reduced ambient insulin levels may result in an up-regulation of cell surface insulin receptors. after 6 weeks of whole grain cereal supplementation. Therefore. These phenomena result in a significant reduction in postprandial glycemia. The study diets included six to ten daily servings of grains.1) have been proposed for the glucose. as well as a significant decrease in fructosamine and PAI-1. In addition to the blood pressure reductions noted above. Jarvi et al. fasting insulin was lower. findings from both a recent prospective epidemiologic study and two clinical trials of polyp recurrence prevention have raised serious questions about this relationship. with grains richest in soluble fiber having the lowest indexes. Such findings suggest that other whole grain components or a synergy between components provide protection against carcinogenesis.60 Intact whole grains have lower glycemic indexes than refined grains. and resistant starches by bacteria in the large intestine. may increase glucose oxidation. The glycemic index is the elevation in plasma glucose after the consumption of a specific carbohydrate-rich food as compared to white bread or a glucose solution.62 A recent animal trial suggested that soluble fibers may also directly increase the translocation of GLUT 4 receptors to the cell surface. Our research group conducted a trial comparing whole grain oat cereal intake with refined grain cereals. resulting in improved insulin sensitivity without affecting PI3 kinase activation.59 In fact. increased intake of fermentable carbohydrate results in corresponding increases in the concentration of bifidobacteria and decreases in E. such as the lipid fraction or oligosaccharides. 2001 7:48 PM WHOLE GRAINS.2) help explain the protective nature of whole grain components against colon cancer. Total and secondary bile acid concentrations were significantly lower during the rye bread period. to a lesser extent.. More recently other whole grain components.3_fm Page 489 Sunday.72 Additionally. Such actions decrease the opportunity for fecal mutagens to interact with intestinal epithelial cells. may be a protective factor in colon carcinogenesis. Gastrointestinal Effects Most of the research to date has investigated the effects of dietary fiber on colorectal cancer. thereby reducing the likelihood of cellular mutation. one of the end products of bacterial fermentation is increased formation of SCFAs. Certain bacteria exhibit differential enzyme activity (i. refined grain) are known to exert various gastrointestinal effects.2387_ch6. Test breads provided a minimum of 20% of daily energy intake and were consumed for two 4-week feeding periods separated by a 4-week washout. May 6. the preferred fuel of colonocytes. insoluble lignified fiber found in the outer bran of whole grains is only slightly degraded but greatly increases fecal bulk due to its physicochemical properties and water-holding capacity. Dietary fibers and.68 Moreover.74 Diacylgycerol content is further increased in the presence of secondary bile acids. multiplicity.76 compared the effects of whole-meal rye vs. increased fecal bulk has also been shown to greatly reduce intestinal transit time. coli and clostridia. Compared to the white bread control. Additionally. suggesting that dietary fibers might affect the cancer process by encouraging the colonization of certain types of bacteria. AND CHRONIC DISEASES 489 components in whole grains. only one human trial has been conducted to investigate the effects of whole grains on risk for colon cancer. most of which appear to be protective against colorectal cancer. Additionally. a recent interesting animal trial demonstrated that feeding male F344 rats a 10% wheat bran diet fortified with 2% bran oil and dephytinized resulted in a significant decrease in colon tumor incidence. whole-meal rye bread significantly increased fecal output and frequency as well as reduced mean intestinal transit time in both men and women. beta-glucuronidase) whose by-products might be carcinogenic. For example. oligosaccharides dilute colonic contents by increasing fecal weight. Moreover. The lipid fraction of whole grains is known to contain omega-3 fatty acids. and volume. As indicated above.73 Moreover.70 Highly fermentable carbohydrates such as soluble fibers and oligosaccharides also affect fecal weight by affecting bacterial metabolism and increasing the formation of bacterial biomass. which have been suggested to promote the tumorigenesis process related to colon cancer. CEREAL FIBER.69 Total fiber and oligosaccharides (which are known to be more concentrated in whole vs. and increase fecal frequency.75 To date. intestinal bacteria convert primary bile acids into secondary bile acids. It has been suggested that butyrate.e. the cumulative effect of whole grains on insulin metabolism will be explored. suggesting that whole grains improve gastrointestinal function and reduce colon cancer risk. Much of the action has to do with the fermentability of these agents. Several mechanisms (see Figure 6.3. the fermentable fraction of whole grains increases SCFA . and the feeding of a diet containing 5% energy from n-3 fatty acids has been shown to decrease the incidence of aberrant crypt foci in rats. antioxidants. Colonocyte proliferation is promoted by secondary bile acids. Dietary fibers protect colonocytes by binding or diluting secondary bile acids. including the fermentable fraction.71 Importantly. Hyperproliferation increases the opportunity for DNA mutations and subsequent replication of abnormal cells. have been shown to decrease risk of this cancer. white wheat bread on some putative colon cancer risk markers in 17 healthy Finnish study participants. and phytoestrogens. The findings from this human trial are congruent with many animal and in vitro trials. accelerate intestinal transit time. Grasten et al. gut bacteria increase intestinal diacylglycerol content via the catabolism of dietary fatty acids and phospholipids. apoptosis. such as bifidobacteria.80 The lipid fraction of whole grain includes n-3 fatty acids.testosterone-related cancers Figure 6.testosterone production . These fatty acids may exert their anticarcinogenic effects via their influence of arachidonic acid metabolism. micronutrients.signifies a decrease in the variable after the pathway has been initiated. and phytochemicals Colonic Effects + antioxidant status + phytoestrogen intake + formation of fermentation by-products Improved gut ecology + fecal bulk/weight .82 n-3 fatty acids decrease arachidonic acid concentration by competitive substitution and thereby decrease the production of the PGE2 series prostaglandins. increase while others decrease.2387_ch6. Legend: + signifies an increase in the variable after the pathway has been initiated.81. increasing subsequent cancer risk.nitrosamines Neutralize free metals .glycemic load + micronutrients + phytochemicals .LH production . possibly via its effects on bacterial metabolism. However. 2001 7:48 PM 490 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and end products of this enzyme have been suggested to be carcinogenic. concentrations.MAP kinase & IGF-1 . Antioxidant Effects Antioxidants from a variety of food sources have been shown to decrease the incidence of various types of cancer. The type of bacteria in the colon also affects intestinal concentrations of diacylglycerols. Diacylglycerols have been shown to activate protein kinase C isozymes.formation of secondary bile acids .PGE2 series prostaglandin formation . Bifidobacteria possess a low beta-glucuronidase activity.79 Wheat bran compared to a low-fiber diet has been shown to decrease the total intestinal diacylglycerol concentration.intestinal diacyglycerol content . These fatty acids have also been shown to alter oncogene expression.GTR production + apoptosis . and intracellular signal transduction pathways.3.PTEN & PI-3 kinase .estrogen-related cancers . which lowers colonic pH and results in a decreased conversion of primary to secondary bile acids. The PGE2 series of prostaglandins have been shown to increase colon cancer cell proliferation and inhibit apoptosis. This 20-carbon fatty acid is the precursor of the dienoic prostaglandins.2 Whole grains and cancer. In doing so.colonocyte proliferation .77 Other bacterial enzymes including urease (produces ammonia from urea) have been implicated in colon cancer due to the tumor-promoting actions of their end products.cell proliferation + 2:16alpha-OHE1 ratio .DNA mutagenesis . May 6. Fermentable agents also alter gut ecology.oxidation reactions . theoretical model of mechanisms of action. 3RD EDITION Whole grains rich in dietary fiber. the effect of whole grains on these enzymes is unknown.fecal mutagens . Whole grains which contain a variety of antioxidants such as vitamin E.colonocyte AA content + n-3 fatty acid concentration (from whole grains) . . .intestinal transit time .3_fm Page 490 Sunday.estrogenic activity .78 and different isozymes stimulate cell proliferation. This is one of the anticarcinogenic effects of n-3 fatty acids.serum insulin concentration . concentrations of certain bacteria. oats. This enzyme has been shown to inhibit cell proliferation and.1 and 6.. selenium is considered a suppressing agent when it comes to its anticarcinogenic activity.5). Vitamin E. Whole grains are thought to be particularly rich sources of phenolic acids. This vitamin has also been shown to inhibit nitrosation reactions in the gut. in general.3_fm Page 491 Sunday. and coumestans (phytoestrogens) are compounds found in plants that exhibit estrogenic activity due to their structural similarity with endogenous estrogens. They appear to act by preventing the formation of carcinogens from precursor compounds and by blocking the interaction of carcinogens with critical subcellular components.83 Phytoestrogen Effects Lignans. Antioxidants function to reduce or delay the rate of oxidation reactions. and biological activity. isoflavones. metabolism. If such mutations become part of the permanent genetic code. especially ferulic acid. This suggests that only whole grains. AND CHRONIC DISEASES 491 some trace minerals. and antinutrients such as phytic acid. the action of glutathione peroxidase is a function of the availability of selenium.e. Under numerous experimental conditions. and not refined grains. polyunsaturated fatty acids and genetic material) from oxidation reactions. phenolic acids. in addition to its general antioxidant capacities. phytic acid greatly reduces the oxidant load in the gastrointestinal tract. thereby decreasing the availability of nitrosamines for absorption. Moreover. are a significant dietary source. certain phenolic acids (caffeic and ferulic acids) have been defined as inhibitors of cancer cell initiation. less oxidative damage occurs in the cell and the likelihood of forming cancer cells is reduced. antioxidants protect genetic material from oxidative damage and subsequent mutation. 2001 7:48 PM WHOLE GRAINS. phenolic acids protect against carcinogenesis via the induction of detoxification systems.2387_ch6. Suppressing agents prevent the expression of neoplasia in cells that have previously been exposed to a carcinogen. which might be reduced by phytic acid. vitamin E has been shown to protect electron-dense cellular components (i. and phytoestrogens.83 By chelating metals.84 Lignans are known to be associated with the fiber components of grains and therefore are thought to be lost in the refining process. specifically the Phase II conjugation reactions. May 6. Phytic acid is another antioxidant. In general.2) by which the antioxidants in whole grains provide protection against cancer are many. have also been shown to effect the carcinogenesis process (see Chapters 6. the concentration of both compounds is significantly reduced by the refining process. However. Phytic acid is often classified as an antinutrient due to its ability to bind various minerals in the intestinal tract. thereby making then unabsorbable. Certain whole grains such as wheat. Phytic acid. such activity might explain the anticancer properties of phytic acid. In doing so. The antioxidant enzyme glutathione peroxidase requires selenium as a cofactor. Grains are also potentially rich sources of vitamin E and selenium. In addition to their general antioxidant properties.83 As mentioned above. Metals such as iron and copper have been shown to initiate oxidation reactions via the formation of reactive oxygen or nitrogen species such as the hydroxyl radical. which is concentrated in whole grains. with its known chelating properties. might decrease the availability of metals and subsequent oxidation reactions. has been classified as a cancer inhibitor due to its ability to inhibit the formation of carcinogens from precursor compounds such as nitrosamines.3. and compromised selenium status has been shown to decrease the activity of this enzyme. Colonic bacteria also produce a significant amount of reactive nitrogen species. The mechanisms (see Figure 6.84 . However. and rye are known to be concentrated sources of lignans. the initiation stage of cancer is said to have begun. Wheat bran and readyto-eat cereals containing bran have been shown to exhibit significant antioxidant activity.35–37 Phenolic acids are concentrated in the bran fraction of the grain. Phytoestrogens have been shown to exhibit certain cancer-protecting properties. CEREAL FIBER. At least some of these effects are due to their effect on endogenous sex hormone production. Such activities reduce the oxidant exposure of intestinal epithelial cells and possible genetic mutation. Iron has been shown to be in its unbound form in the intestinal tract and thereby capable of initiating such reactions. It has been proposed that phytoestrogens result in decreased hypothalmic production of gonadotrophin-releasing hormone. thereby explaining their powerful cancer-reducing effects. the spontaneous development of prostate cancer was significantly reduced in male LobundWistar rats after the feeding of isoflavones for 22 months.94–97 As mentioned above.2387_ch6. The end products of these hydroxylation pathways are 2-hydroxylated and the 16-alpha-hydroxylated metabolites. a higher glycemic index diet has been shown to decrease insulin sensitivity and results in increased serum insulin levels.88 Similarly. and specifically lignans. changes in testosterone metabolism have been suggested to protect men against hormone-related cancers. For example. thereby decreasing the production of the powerful androgen dehydrotestosterone from testosterone. refined grains on average exhibit a higher glycemic response than whole grains.91 In men. Conversely.93 Phytoestrogens.89 The mechanisms (see Figure 6.2) by which insulin is related to the carcinogenesis process. Over time.92 Additionally.2) by which phytoestrogens might protect against the occurrence of hormone-related cancers are poorly understood. changes in endogenous estrogen metabolism products might explain the chemoprotective effects of lignans.98 Therefore. The phytoestrogen family of isoflavones has been studied to this end.3_fm Page 492 Sunday. Modification of the glycemic index has been shown to affect satiety. which ultimately might affect weight gain and subsequent insulin resistance. Due to the different biological activities of these products. it has been proposed that an elevated ratio of 2-hydroxyestrogen to 16-alphahydroxyestrone (2:16alpha-OHE1) would reduce the risk of hormone-related cancers in women. that has been the delivery agent studied most often. very little empirical data exist to explain the mechanism (see Figure 6. Estradiol and estrone are metabolized along two irreversible and competing pathways. emerging evidence suggests that increased whole grain consumption might protect against future cancers. genistein. This may explain in part why obesity is a risk factor for cancer. probably exhibit many other anticarcinogenic properties. the 16-alpha-OHE1 demonstrates significant estrogenic activity and subsequently increases risk of breast cancer.89. significant lengthening of the overall cycle duration has been shown to reduce the risk of hormone-related cancers. Insulinemic Effects Epidemiologic studies have reported that higher serum insulin levels are associated with increased risk of colon. daily consumption of 10 g of flaxseed significantly increased the urinary 2:16alpha-OHE1 ratio in premenopausal and postmenopausal women. has been shown to reduce the activity of 5-alpha reductase. In two recent human trials. but currently these are unknown. a specific phytoestrogen. and possibly other cancers. breast. Additionally. phytoestrogens act as agonists for the action of testosterone in the prostate and related tissues and thereby possibly reduce carcinogenesis. Due to the concentration of lignans in flaxseed. As might be expected. Either directly or indirectly. Flaxseed powder supplementation has been shown to increase the average luteal phase length of the menstrual cycle in premenopausal women.85 Additionally. whole grains might reduce the risk of certain types of cancer by preventing acute and chronic states of hyperinsulinemia.86. 2001 7:48 PM 492 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. such as prostate cancer. In a recent animal trial.87 Flaxseed has also been shown to delay the progression of mammary tumorigenesis in a recent animal trial as well as reduce early markers of risk for mammary and colon carcinogenesis. 2-OHE metabolites are thought to exhibit very little estrogenic activity and might even possess antiestrogenic properties.3. It is also known that increased deep visceral adiposity decreases insulin sensitivity. Most of the evidence comes from animal .3.90 The ratio of these metabolites is a fair measure of total estrogenic activity and is decreased in women with breast cancer. 3RD EDITION Modification of estrogen metabolism has been suggested to alter the etiology of breast and other hormone-related cancers. Preliminary reports indicate that lignans directly inhibit the growth of human mammary tumor cells as well as reduce mammary and prostate tumor initiation. in women. which in turn inhibits the production of luteinizing hormone in the anterior pituitary with a subsequent decrease in the production of circulating testosterone by the Leydig’s cells of the testes. May 6. 412. Jacobs. 337. and Kruger. Gerrior. Knudsen. 2. Am. Finally.. 70. Statistical Bulletin 928. S. J. 1999. and Eggum. 5. Chronic hyperinsulinemia and related metabolic consequences might increase risk of future breast. and (2) more studies testing whole grains as the primary intervention rather than a whole grain component are greatly needed. Washington D. January. Other metabolic effects (elevated triglycerides and free fatty acids) may explain the carcinogenic effects associated with states of hyperinsulinemia. Am. Epidemiol. and polyphenol oxidase activity. 1026. Department of Agriculture.3_fm Page 493 Sunday. B. R. AACC whole grain ingredient definition. 1997. type 2 diabetes. Am... 1997. CEREAL FIBER. Nutr. O. S. Am... possibly through the mitogen-activated protein (MAP) kinase pathway. et al. D. and Allshouse. It is very possible that gastrointestinal and/or systemic interactions occur between dietary components to. serum insulin growth factor-I (IGF-I) levels are elevated in response to insulin. Whole grain intake may reduce risk of coronary heart disease death in postmenopausal women: the Iowa Women’s Health Study. 1996. American Association of Cereal Chemists.. 60. Public Health. 68. but it is imperative that more studies are conducted using whole grains as the primary intervention. More mechanistic work is certainly needed. Potter. as well as other diseases. Nutritive value of cereal products with emphasis on the effect of milling. REFERENCES 1. Liu. J. D. 1989. R. Nutr. 1998. E.. 9. however. prostate. IGF-I has also been shown to increase cell proliferation and reduce apoptosis. Clin. most of these pathways are currently poorly understood. 4. In terms of the latter issue. Food Consumption. J. B. Pedersen. 6.. J. Written communication. 79. explain their physiological effects. Simple phenolic acids in flours prepared from Canadian wheat: relationship to ash content.99 Decreased apoptosis has also been associated with elevated insulin levels. Cereal Foods World. and modification of the phophatidylinositol 3-kinase and PTEN pathways might explain this effect.. Cereal Chem. this will allow a clearer discussion of the relationship between whole grain intake and CHD. 1999.2387_ch6. Clin. 248. J. B. 8. et al.. Prices and Expenditures. 2000. but future elucidation of this relationship is necessary. and cancer. Jacobs. 45. World Rev. Food and phytochemicals. and the recent health claims and public health recommendations to increase whole grain intake appear justified. two things are apparent: (1) the possible mechanisms by which whole grains and their components modify risk for CHD and type 2 diabetes are better elucidated than are mechanisms which might relate to cancer. Whole-grain consumption and risk of coronary hear disease: results from the Nurses’ Health Study. Hatcher. it has become clear in recent years that the “magic bullet” approach for investigating the disease-modifying effects of diet is not the best model.S. In the future. Putnam.100 Additionally. J. E. Diet. 1997.. J. D.C. W. more animal work is needed to determine the possible mechanisms by which insulin and related variables might induce carcinogenesis. Hyperinsulinemia has been shown to induce cell proliferation. Jr. it is clear from this review that the “whole” is greater than the sum of its parts. Nutr. U. 322. Is whole grain intake associated with reduced total and cause-specific death rates in older women? The Iowa Women’s Health Study. Finally.. 2001 7:48 PM WHOLE GRAINS. 3. at least in part. May 6. D.. K. 7. More prospective epidemiologic data are needed as well as human trials investigating the relationship between insulin and risk markers for these cancers. Jr. E.. 144.. AND CHRONIC DISEASES 493 studies investigating the effect of insulin on colon carcinogenesis. et al. 89. . 74. 1996. and colon cancer. CONCLUSIONS Based on this review. 1. magic bullets and measurement error: a commentary. J. color. J. and Macfarlane. Effects of dietary inulin on serum lipids. V. H. W. J. Am. 55. Rigaud. and Talbott. sympathetic overactivity and cardiovascular morbidity. Fermentation of resistant rice starch produces propionate reducing serum and hepatic cholesterol in rats. Pins. 1999. Biol. Food Chem.. E. Diet Assoc. et al. 13.. et al. 1994. 14... May 6. 1996.. blood lipids and plasma glucose levels (Abstract #39).. 1998. fruit. M. M. 1979. Leinonen. et al. Rev. A. 2000. Nutr... K. 1998. Colonic microflora: nutrition and health. M. 1450. and Pomare. B. J. Physiol. Wurch. J.. Miller. A.. 407. M. G. Pharmacology of dietary fibre. Vuksan. Lipid Res. J. Abnormalities of coagulation and fibrinolysis in insulin resistance. Meta-analysis of effects of soy protein intake on serum lipids in humans. Am. J. Spiller. A. and cholecystokinin responses in men fed barley pasta enriched with β-glucan. H. J. 1474S. insulin. 2000. 1774. Navab. 1. J. 1927. Pereira.. Rimm. S. K. Miller H.. . 94. S. C. 129. 35. 13. P. Soluble fiber and serum lipids: a literature review. Arterioscler. 1990. Nutr.. R.. U. Oat ingestion reduces systolic and diastolic blood pressure among moderate hypertensives: a pilot trial. 1. et al. et al. Am. 1997. JAMA. Cummings. 275. et al. Davidson. Katakam. C.. Nutr. Decker. J. 1991. 18. J. Food Sci. 27. The role of viscous soluble fiber in the metabolic control of diabetes. 276. Clin. FASEB J. J. Prev.. Cheng. Thromb. et al. 11.. A comparison of barley and wheat products on blood lipids. Antioxidant effect of aqueous extracts from wheat based ready-to-eat breakfast cereals. Effect of oral supplementation with d-alpha-tocopherol on the vitamin E content of human low density lipoproteins and resistance to oxidation. Marlett. and Mykkanen. Diabetes Care. J. 14. A203.. 20. Engl.. Johnstone. 52. fruits and vegetables. I. 15. hunger feeling and food intake in normal volunteers: a double blind study. 22. Endocrinol. 59. Whole-grain products and antioxidants.. 19.... S. J. Cereal Foods World. B. W. and cereal fiber intake and risk of coronary heart disease among men.. J. A. Evaluation of the health aspects of using partially hydrolyzed guar gum as a food ingredient. et al.. et al.. A. Effect of psyllium on gastric emptying. 10. Clin. 39. 21.. D. 32. Bourdon. Heaton. 275. J. R. 1998. Defective nitric oxide synthesis: a link between metabolic insulin resistance. 8. Diabetes Care. N. et al. Evidence for a common antecedent?. 3RD EDITION 10.. 69. Crit. J.. Antioxidants and hormone-mediated health benefits of whole grains.. Yudkin. 686. Endothelial dysfunction preceeds hypertension in diet induced insulin resistance. M. 2000.2387_ch6. Dieber-Rotheneder. E. 19. Rye bread decreases serum total and LDL cholesterol in men with moderately elevated serum cholesterol. 1994. W. Prac. 142.. Whole grain cereals reduce antihypertensive medication need. 447. H.. A low-viscosity soluble-fiber fruit juice supplement fails to lower cholesterol in hypercholesterolemic men and women. Nutr.. Thompson.3_fm Page 494 Sunday. Scherrer. Baublis. 25. 26.. 239. 38. Nutr. 17. C. 16. L. and Lai.. T.. Am.. S. and Maki. et al. 2001. 1992. 16. Mechanism of serum cholesterol reduction by oat bran. M.. The effect of whole grains on inflammation and fibrinolysis (Abstract #3). 130. Pereira. Am Coll. 20. 2000. A. Keenan. 2001 7:48 PM 494 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.. 23. 36. in press. E.. 1999. 22. 1325. M. Anderson. 18. 1997.. H. The yin and yang of oxidation in the development of the fatty streak. Pins. satiety. 1999. Eur.. 31.. Hepatology. Lancet. Fisher. U. J. J. M. Bethesda.. CVD Epi. 20. and Pi-Sunyer. 34.. 2000.. et al. Diabetes Care. K. 40. et al. Poutanen. et al. 45. 62. The effect of whole grains on insulin sensitivity in overweight adults (Abstract #150. J. D. 315. et al. Vasc. H. Nutr. J. and Sartori. M. 2000. 22. 30. 32. R788. 34. Konjac-mannan (Glucomannan) improves glycemia and other associated risk factors for coronary heart disease in type 2 diabetes. E. 1994. F. J. M.. E. Nutr. Med. Anderson. and Cook-Newell. 19. 1996. and markers of oxidation (Abstract #49). 913. H.. Davidson.2).. 312S. J. 128. 333.. Federation of American Societies for Experimental Biology. J. 2000. 473. Prog. 425. Postprandial lipid. K. J. 29. 1994. Coll. 37. S... X. 1995. Antioxidant content of whole grain breakfast cereals. 33. M.. Eur. Pharmacol Ther. 12.. Effect of bran on blood lipids and calcium. Vegetable. MD. Coll. 831. 529. 1999. K. Am. Life Sciences Research Office. glucose. Glore. et al. 164. 28.. Fam. Nutr. 49. F. E. J. C25... Nutrition. 476. et al... Nutr. 2000. 24. 130. J. Nutr. 1999. P. M. and Clydesdale. M. 1999. Braaten. 1988. AND CHRONIC DISEASES 495 40.. 688S. 759. J. W. 2000. 45. 27. Adlercreutz.. Jenkins. 63. J. T. E.. N. 1999. 43.. Pharmacol. Nutr. Determination of urinary lignans and phytoestrogen metabolites. and Clarkson. Y. 971. J. et al. high-fiber diets for insulin-dependent diabetic individuals. 1999.. Anderson. Nutr. 47. Wholemeal versus wholegrain breads: proportion of whole or cracked grain and the glycaemic response. J. Med. J. High beta-glucan oat bran and oat gum reduce postprandial blood glucose and insulin in subjects with and without type 2 diabetes. Vitamin E and atherosclerosis. J. Data presented at American Association of Cereal Chemists annual meeting.. Y. Coron. E. C. 1990. H. J... W. M. J. Thorburn. Chandalia. 363. A. 53. 459. A. et al. K. J. 43.. J. 60. Clin. Am. H.. B. Arabinoxylan fiber. Liukkonen. Mendelsohn. Chan.. Nutr. Improved glycemic control and lipid profile and normalized fibrinolytic activity on a low-glycemic index diet in type 2 diabetic patients. Diabetes Care. S. Jenkins. 1392. 59.. et al. Soluble dietary fibre improves insulin sensitivity by increasing muscle GLUT-4 content in stroke-prone spontaneously hypertensive rats. D. J. 51.2387_ch6. Intern. 1991. 2000. Kansas City. 56. Med. Clin.. et al. T.. Med... Formation of enterolactone during in vitro fermentation of rye. Beneficial effects of high dietary fiber intake in patients with type 2 diabetes mellitus. and incident type 2 diabetes in older women. 130. et al. H. Vorster. J. R. G. Fuchs. Nutr. 1994. Lu. T. Reaven. K. Nutr. M. Drews. 1300. et al.. and Cummings. D. Clin. J. 49. J. 1968. N.. J. C.. Granfeldt. Carbohydrates. X. 41. J. et al. 169. I. et al. 1009. 1999. 2000. R. J. 1998. 1801. Am. 340. Arepas made from high amylose corn flour produce favorably low glucose and insulin responses in healthy humans. 466. Metabolism. a byproduct of wheat flour processing. S. 1986. dietary fiber. 57. 1994. 921. Diabet. Williams.. 846. K. 41. et al. Reduction of plasma homcyst(e)ine levels by breakfast cereals fortified with folic acid in patients with coronary heart disease. 312. Metabolic effects of high-carbohydrate. Am. 128. W. 65.. C. Venter. et al. Nutr. 356. A. J. 1995.. et al.. Fukagawa.. Meyer... et al. J. Nutr. Engl... 340... and Miller.. 46. 52. 1998. J. Calcium and fibre supplementation in prevention of colorectal adenoma recurrence: a randomised intervention trial. Steroid Biochem. 1986. 516. A. et al. Effects of dietary propionate on carbohydrate and lipid metabolism in healthy volunteers. Dietary fiber and the risk of colorectal cancer and adenoma in women. 71. CEREAL FIBER. Nutr. Bjorck. 1990. et al. Clin. J. 42. 2000. N. 780... Yamakoshi. Nov. Lancet. 55. et al. Clin. 71. 936... 560. 59S. Artery Dis.3_fm Page 495 Sunday. H. Clin. J. 2000.. 42. J. N. et al.. H. Am. Nutr. Jarvi. Engl. 125. 297.. Med. Am. 62. 1123... J. A. 1887. 524. Carbohydrate fermentation decreases hepatic glucose output in healthy subjects.. 958.. Med. Wolever.. Engl. J. Brit... A. Bonithon-Kopp. 22. 1593. 11. and Proietto. 1999. Med. Ann. 2001 7:48 PM WHOLE GRAINS. . 1998. Dietary soy isoflavones inhibit in-vivo constrictor responses of coronary arteries to collagen-induced platelet activation. M. N. J... Muir. Med. Engl.. May 6. MO. 66. 58. J. 17. 64.. Anderson. and Bjorck. 9.. 54. 54. 1999. S. et al. 50. R. Am. 1993. M. et al. et al. Clin. reduces the postprandial glucose response in normoglycemic subjects. Clin. Physiol. The glycemic index: methodology and clinical implications. Am. Malinow.. J. Am. Exp.. Nutr. Am. C. The protective effects of estrogen on the cardiovascular system. Food properties affecting the digestion and absorption of carbohydrates. 61. 2000. Gastroenterol. Homocyst(e)ine and cardiovascular disease: a critical review of the epidemiologic evidence.. 44. 2000. Isoflavone aglycone-rich extract without soy protein attenuates atherosclerosis development in cholesterol-fed rabbits. A. 25. 338. High-carbohydrate. J. Diabetes. K. 48. potential antiestrogens and anticarcinogens in urine of women on various habitual diets. I. et al. 549. Eikelboom. and Karas. 131. 10. 1991. 54. Z. 85. J. high-fiber diets increase peripheral insulin sensitivity in healthy young and old adults. 342. 52. Nutr. Song. 70. Clin.. Effects of psyllium on glucose and serum lipid responses in men with type 2 diabetes and hypercholesterolemia. Low glycemic response to traditionally processed wheat and rye products: bulgur and pumpernickel bread. Study of the relationship between glucose and insulin responses to an oral glucose load in man.. Haggans. Acad. 1. H. and Cummings. 649.. et al. 75. and Glitso. Engl. Med. 294. Preventive potential of wheat bran fractions against experimental colon carcinogenesis: implications for human colon cancer prevention. 1997. B. Phyto-oestrogens and Western diseases. J. et al. Biochemical epidemiology of colon cancer: effects of types of dietary fiber on colonic diacylglycerols in women. Nutr. J. 29. an activator of protein kinase C. colon cancer risk and dietary intake of non-starch polysaccharides (dietary fiber). 33. Gastroenterology. 1995. Cancer Epidemiol. et al. G. Grasten.. L. E. 72. Science. Haggans. Griffith. Rickard.2387_ch6. 50. May 6. and Mazur. Bradlow. M.. Med.. 60. Academic Press. W. et al. Microbiol.. and Wolter. Cummings. Oxidative metabolism of estradiol. J. Inhibition of 5α reductase and 17β hydroxysteroid dehydrogenase in genital skin fibroblasts by dietary lignans and isoflavinoids.. H. 3539. 1994. Ann.... Cancer Res.. in Functional Properties of Non-Digestible Carbohydrates... 85. G.. Schneider. 3047. 1995. Selected eicosanoids increase the proliferation rate of human colon carcinoma lines and mouse colonocytes in vivo. Hormones. Simi.. 81. et al. G. S. 3RD EDITION 67. Cancer Res. B. Morotomi. Sci. J. 1156. 2215. Chem. Bartoli. Biophys. H. M. M. Dietary fibers and fats alter rat colon protein kinase C activity: correlation to cell proliferation. 92. T. et al. Proc. R. Med. Nutr.. Cancer. Acad. Fishman. Qiao. J. 73. Reddy. 883. Hawksworth. 45. 1980. J. Stephen.. B. M. 1990. 84. 3104. W. et al. and Litwack. 513. Reddy. C. 90. 1997. Guillon. 1992. J. S. K. 74. Lack of effect of a high-fiber cereal supplement on the recurrence of colorectal adenomas. 13. 608. 191. Med. Urinary 2/16-hydroxyestrone ratio: correlation with serum insulin-like growth factor binding protein-3 and a potential biomarker of breast cancer risk. N. 9. Rye bread improves bowel function and decreases the concentrations of some compounds that are putative colon cancer risk markers in middle-aged women and men. 295. Effect of olive oil on early and late events of colon carcinogenesis in rats: modulation of arachidonic acid metabolism and local prostaglandin E2 synthesis.3_fm Page 496 Sunday. R.. Pollard. 79. E. Cancer Res. S. 70. 101. Evans. N. 58. D. H. 80. L. and Hill. and Morton. by human intestinal flora. 1982.. 1997. Bach. S. 2000.. 1521.. Nutr... 77. M.. Y. 147. V... Natl. 2000. 451. 46. Prostate. S. 86.. The effect of flaxseed and wheat bran consumption on urinary estrogen metabolites in premenopausal women.. Production of diacylglycerol. 78. Singapore. Fecal weight. Scheppach.. 27. C. 1985. 82. and Engle. Dose effects of flaxseed and its lignan on N-methyl-N-nitrosourea-induced mammary tumorigenesis in rats. 4. Reddy. 69. 45... and Watanabe.. B. 2000.. Norman. 2000. W. France. A. Biol. F. 4792. and Gallagher. et al. S. Endocrinology.. 39. The microbial contribution to human fecal mass. Abnormal oxidative metabolism of estradiol in women with breast cancer. Gut.. 106. Acta. Drasar. Modulation of apoptosis and Bcl-2 expression by prostaglandin E2 in human colon cancer cells. K. 1999. A. 87... Biochem. Nantes. Effect of cholesterol metabolites and promoting effect of lithocholic acid in colon carcinogenesis in germ-free and conventional F344 rats. 1998... et al.. Prevention of spontaneous prostate-related cancer in Lobund-Wistar rats by a soy protein isolate/isoflavone diet. 1971. Cancer Res. W. Intracellular signaling by hydrolysis of phospholipids and activation of protein kinase C. 1960. Butyrate and the epithelium of the large intestine. Chapkin. 1998.. 79. 71. 1999. 1998. Cancer. Ho.. H. et al. 130. Med. et al. H. Wattenberg. 362. 89. K. 235. 215. 342. J. Sheng.. S.. 45. 123. Johansen. A. Eds. 1258. H. 50. Biomarkers Prev. 1992. 93. Adlercreutz. 91. et al. et al. 88. J. Chemoprevention of cancer. 72. 95. 215. Microbiol. J. USA. Cereal Foods World. 1993... K. et al. J. 2001 7:48 PM 496 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 690. F. 83. 2000. 2000. A. Rye dietary fiber and fermentation in the colon. L. 35. 258. Alberts. Ann. S. Nutr. . M. Cancer Res. Intestinal bacteria and the hydrolysis of glycoside bonds. 1979.. Nishizuka. 103. 76. B. 1783.. Gastroenterology. S. New York. W.. 188. J.. J. J. 719.... 68. et al. B. Effect of flaxseed consumption on urinary estrogen metabolites in postmenopausal women. R. et al. 1994. 1995. Cancer Causes Control. Schoen. Krazer. G. 2001 7:48 PM WHOLE GRAINS. 1994. 43. Insulin resistance. McKeown-Eyssen. E. CEREAL FIBER. Insulin and colon cancer. E. 1999. 1999. 687..3_fm Page 497 Sunday. 1245. JNCI. Trends Cell Biol. 98. insulin like growth factor I and breast cancer: a hypothesis. Metabolism. 1066. 96.. and the cause of type II diabetes. 1147.. 6.. T. J. 3. 164. Giovannucci. E. 99. Cancer Epi. Diabetes. Insulin action. ..2387_ch6. body size and incident colorectal cancer. R.. R.. 125. 97.. C. R. 1998. 1995. Cancer. 62. Banting Lecture. Insulin sensitivity in women at risk of coronary heart disease and the effect of a low glycemic diet. Bio. 9. G. Prev. Kahn. J. Increased blood glucose and insulin. Epidemiology of colorectal cancer revisited: are serum triglycerides and/or plasma glucose associated with risk?. 91. diabetogenes. PTEN: a tumor suppressor that functions as a phospholipid phosphatase. 100. 403. Int. Maehama. 95. May 6. et al. Frost. 47. and Dixon.. AND CHRONIC DISEASES 497 94. May 6.2387_ch6.3_fm Page 498 Sunday. 2001 7:48 PM . chromium.2387_ch6. and nickel. to a certain extent. May 6.4_fm Page 499 Sunday.00+$1. been defined as a part of the dietary fiber complex. An increased consumption of flours with high extraction rates of these compounds is therefore a nutritional aim in many countries. Due to the chelating properties of the dietary fiber components and the phytic acid. and it seems that currently only a small fraction of these problems are recognized and corrected for. MINERALS STUDIED Bioavailability problems of practical significance are known for some minerals of long established nutritional importance such as calcium. where dietary fiber and phytic acid are also recovered. but there seem to be similar problems for other minerals and trace elements such as selenium.4 Bioavailability of Minerals from Cereals Wenche Frølich INTRODUCTION Whole grain cereals and cereal products are some of the best sources not only for dietary fiber. there has been a lot of concern about the effect of an unrefined high-fiber diet with respect to mineral bioavailability. An increasing knowledge of the complexity of nutrient interactions and the number of dietary components that can influence bioavailability makes this field very complicated. copper. but also for minerals and trace elements in our diet. Most minerals and trace elements in the cereals are closely related to the outer layers.50 © 2001 by CRC Press LLC 499 . The complexity of interactions that may take place between minerals and different components in the dietary fiber complex makes it very difficult to predict the bioavailability of minerals in whole grain cereals by chemical determinations of minerals and trace elements present in the cereal products. MINERALS AND UNREFINED CEREAL PRODUCTS It has been claimed that the bioavailability of minerals and trace elements in diets rich in whole grain products can be reduced in comparison with diets rich in refined cereal products. New knowledge of bioavailability is constantly being discovered. 2001 7:49 PM CHAPTER 6. and iron. especially the aleurone cells. All these components have therefore. zinc. Great 0-8493-2387-8/01/$0. This includes both the difficulties in the analysis of dietary fiber and the various physical and chemical conditions which may alter the interactions between the nutrients. have been connected with the chelating properties of dietary fiber vs. It is also important to stress that the refining of cereals leads to a marked fall in the mineral content. 3RD EDITION controversies. This type of enrichment study has given quite different results from studies with the corresponding amount of the mineral naturally occurring in the food. and P is available. both human and animal studies. The interpretation of these studies is difficult and extrapolation to normal diets is often impossible. probably due to the various chelating properties of the different fiber components. Zn. as has been done in various enrichment studies. or both that are responsible for the decreased absorption of minerals from whole grain cereal products. an inhibition of the percent uptake of minerals and trace elements from the whole grain cereal products does not necessarily mean a decreased absolute intake. Dietary fiber is often divided into a soluble and an insoluble fraction due to relative solubilities and other chemical properties. 2001 7:49 PM 500 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. the results in the literature are conflicting. BINDING BEHAVIOR OF MINERALS The binding behavior of different minerals to cereal fibers seems to be quite variable.4_fm Page 500 Sunday. . It is important to realize that addition of isolated components to the diet only indicates the effect of the same amount of native dietary fiber or phytate present in the original food item. ENRICHMENT STUDIES When a single mineral is added to the food. In many studies. it is difficult to distinguish between the effect of the different dietary fiber components and the phytic acid. The binding behavior of different minerals to cereal fibers seems to be extremely variable. Therefore.2387_ch6. In earlier studies it was claimed that the phytic acid present in the whole grain cereal products was the component chiefly responsible for the chelating of divalent minerals. even if the same amounts of dietary fiber and phytic acid were present in the studies. and no agreement exists for the different minerals studied. in terms of mineral interactions. the balance between this mineral and the rest of the minerals and trace elements present is changed. Fe. in addition to reduced transit time. Little information on minerals other than Ca. Cu. The solubilities of the minerals may be decreased as a result of chelation to fiber components. The various cereals also have quite different dietary fiber compositions with different chemical structure and binding capacities. Much of the older work is still relevant today and has been taken into consideration when conclusions have been drawn. Dietary fibers could increase the viscosity and reduce the rate of migration of minerals. In many studies isolated fiber components and phytic acid have been added to the diet to study the effects of the individual components on the mineral bioavailability. making the chelating properties of the same mineral special for each cereal. because the chelating properties are not the same in different cereals due to different amounts of fiber. More recently it has been suggested that phytic acid is not the component which is solely responsible for the decreased mineral availability. different fiber composition. Generalization about the bioavailability of minerals can be misleading. due to the considerably higher amount of minerals present in the whole grain. As dietary fiber and phytic acid most often occur together in the cereal foods. May 6. which. and it is still debatable whether it is the fiber components. could result in changes in the bioavailability of minerals. and different amounts of phytate present in the different cereals. However. probably due to various chelating properties of the minerals. the phytic acid. phytic acid. Fiber itself or other polysaccharides seem to a great extent to complex with minerals. dosages of dietary fiber and phytic acid which are not physiological have been used. which is a measure of the degree of both absorption and utilization. citrate.g. seems to depress iron absorption. studies on children and pregnant women. (2) pH in the solution. the composite diet should be taken into consideration and not only a single food component or fraction of the food. while animal protein improves both zinc and iron absorption. Ascorbic acid seems to be the most potent enhancer of iron absorption. The ability of minerals to bind or chelate to the different cereal components or isolated fiber fractions seems to be different in an in vitro system than when ingested alone or together with a composite diet. oxalic acid. OTHER FACTORS INFLUENCING BIOAVAILABILITY The bioavailability. INFLUENCE OF PROCESSING The heat treatment during processing of cereal foods. Several in vitro studies have been published concerning dietary fiber components that are responsible for the mineral association. On the other hand. direct extrapolations from animals to humans do not give correct answers in all cases. that many observations carried out in animals do have relevance to man.g.4_fm Page 501 Sunday. and amino acids. This is important to bear in mind when conclusions concerning the final cereal products (e. Components in the diet are the extrinsic factors and could have both promoting and inhibitory effects. and before any definite conclusions can be drawn. and several factors must be taken into consideration when interpreting the results: (1) presence of different chelators such as phytic acid. It is of importance to bear in mind that when studying mineral bioavailability. tannins. Only limited studies on minerals connected with deficiency states can be carried out in humans. It is also difficult to carry out long-term studies. The individual needs for the mineral due to nutritional and health status. both during the milling procedure and baking of bread. This is mainly due to the breakdown of phytic acid. e. The studies made with respect to the influence of other food items on mineral absorption from cereals are mainly done with respect to iron. (3) concentration of minerals. Studies on a single food item. (5) presence of type and amount of various dietary fiber components. . is shown to be affected by both intrinsic and extrinsic factors. May 6. Tea. (4) heat treatment that may affect the binding. The different isolated fiber components have been shown to have different cation exchange potentials. with only a few studies on zinc. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 501 HUMAN STUDIES/ANIMAL STUDIES Much less is known about the mineral requirement and bioavailability in humans than in animals. sex. human studies are needed. can be used to identify the dietary components that are responsible for the changes in the mineral absorption. changing the chelating properties of these components. bread) are to be drawn on the basis of the cereal ingredients like flour and bran.. Fermentation and leavening seem to improve the utilization of the minerals. It is important to realize that great care is needed when extrapolating results from in vitro studies to in vivo conditions. also influences the bioavailability of the minerals present in the final product. on the other hand. Evidence suggests. but reorganization of dietary fiber components is also a factor. on the other hand. and age are intrinsic factors. IN VITRO/IN VIVO STUDIES The mechanisms by which dietary fiber chelates to the minerals are largely unknown.2387_ch6. and (6) concentration of minerals.. however. and studies which might cause risks to humans cannot be performed from an ethical point of view. In countries where the intake of minerals is limited and whole grain cereals account for the main part of energy. animal. Such a diet is usually well balanced with a good quality standard. In experiments with a duration longer than 3 to 4 weeks. May 6. however. In diets with extremely high dietary fiber and phytic acid content. deficiencies in both zinc and iron are likely to occur. the somewhat decreased absorption could be of importance. if the increase in the dietary fiber content is due to increased intake of isolated fiber components. and animal products are a natural part of the diet.4_fm Page 502 Sunday. 3RD EDITION LONG-TERM STUDIES NEEDED Adaptation to a diminished availability of minerals is probably of importance after continuous addition of fiber from unrefined cereals. there is no additional intake of minerals. and it is therefore difficult to draw any definite conclusions about the inhibitory/promoting factors until more long-term studies have been performed. There is.2387_ch6. Most of the studies published are. no evidence that the dietary fiber intake from a fiber-rich Western diet will interfere sufficiently with mineral absorption to cause deficiency in a healthy population. The chelating properties of these isolated dietary fiber components could have an influence on the utilization of minerals and trace elements. and in vitro studies on the effect of dietary fiber on mineral bioavailability. There is a general agreement in the literature that further research is needed to clarify a number of questions still not solved. no change in mineral availability from whole grain cereals is observed. CONCLUSIONS There seems to be an inhibitory effect on the bioavailability of some minerals due to some of the components in the dietary fiber complex in cereals. much work is still needed in this field. as fiber and minerals are located together in the cereals. However. It is important to stress that a natural high-fiber diet contains more minerals and trace elements than a low-fiber diet. short-term experiments. On the other hand. 2001 7:49 PM 502 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. more controversy than consensus is found. In the numerous human. however. Different cereals have different chelating properties due to different fiber components present in the various cereals. . The studies completed so far only give us an idea of the complexity of the mineral bioavailability from cereals. 87. 81. 113. 89. 19. 35. 55. 96. 135. 184 .2387_ch6. 158. 186 182 4. 30. 100. 48. 38. 96. 14. 97. 16. 104. 106. 82. 7. 108. 171. 178. 107. 106. 88.1 References to Minerals Studied with Respect to Bioavailability from Cereals Mineral Increased Absorption. 112. 22. 158. 160. 99. 82. 106. 85. 89. 58. 179 64. 48. 186. 36. 43. 74. 72. 84. 158. 183. 94. 40. 182. 132. 183. 59. 173. 51. 122. 184 P 151 87. 71. 88. 68. 101. 115. 13. 20. 103. 89. 23. 105. 172. May 6. 2. 138. 115. 24. 78. 152. 56. 117. 81. 145. 5. 12. 17. 88. 65. 178 4. No Change in Availability. 47. 63. 37. 89. 64. 66. 70. 276 12. 159 83. 123. 42. 84. 177. 15. 3. 136. 44. 121. 33.4_fm Page 503 Sunday. 184. 113 12. 152. 130. 114. 120. 182. 75. 182. 133. 49. Ca Decreased or Low Absorption. 115. 123. 97. 130 6. 154. 121. 64. 10. 159. 11. 106. 111. 73. 125. No. 140. 91. No. 156. 175. 176. 87. 32. 160. 152. 69. 93. 48. 175. 96. 11. 97. 39. 156. 9. 79. 75. Ref.160 69. 149. 106. 89. 167. 66. 114. 182 4. 159. 106. 113. 115. 88. 183. 28. 101. 19. 16. 157. 87. No. 182 5. 153. 159. 8. 169. 109. 116. 61. 113. 83. 137. 106. 152. 65. 131. 16. 115 16.4. 163. 82. 168 Zn 52. 119. 187 1. 53. 95. 175. 82. 18. 110. 177. Ref. 97. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 503 Table 6. 54. 27. 31. 96. 13. 47. 159. 34. 86. 184 Mn Cu Mg Se Fe 21. 99. 45. 82. 16. 115. 80. 158. 57. 114. 177. 159. Ref. 99. 115. 5. 43. 134. 177. 124. 158. 46. 90. 77. 99. 186 4. 177. 98. 96. 42 children Human. Ca. food intrinsically labeled with 55Fe Radio-Fe erythrocyte utilization method Radio-Fe. 1 meal Controlled. Cu Fe Fe Fe Fe. Fe not affected No significant effect of bran. 66 people Human. but mean balance positive Lowered availability. Ca Fe Fe Human.3%) Low absorption Very low absorption (3. 66 people Human. Cu absorption better because of higher intake Not affected Affected by phytic acid Affected. 26 g wheat bran Unpolished rice (27 oz) Maize Maize. 12 people Fe Fe Human. Mg. Mg retention improved.Rat Human. Mg. 1 meal Controlled. Mg Hgb-measurements Balance Serum iron measurements Balance Radioassay.4_fm Page 504 Sunday. 3–5 weeks 2-week period 15 days 49–91 days Time Controlled. Mg absorption lowered on unpolished rice for 3 weeks. Mg. 116 people Human. serum iron. 2001 7:49 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 1 meal Controlled. erythrocyte utilization method 59Fe-labeled maize. 504 Fe Human. absorption negative after 4 weeks. 3 people Fe. P not affected. iron from bread less available than inorganic iron Increased absorption with fortification with FeSO4 Decreased Fe absorption Result 43 1 24 97 16 2 101 22 100 106 55 39 80 107 Ref. 1 meal Controlled. 8 people Subject Description of Mineral Bioavailability Studies from Cereals Fe Mineral Table 6. 40–50% of energy from white flour Diet Wheat bread Different cereals Whole wheat bread 26 g corn bran. IBC radioassay. Ca. 154 people Human Fe Fe Human. 5 people Human Human Human Fe. whole body counter Balance Hgb. 56Fe Balance Radioisotope technique Hgb-relation Balance Exp.8%) Fe. Zn. meal Controlled. Ca absorption positive after 8 weeks Low availability Decreased iron absorption by brown bread. Design 28–30 days 3 weeks 14 days 7-day period. 59Fe. 3RD EDITION .2 2387_ch6. long term (18 weeks): Ca.4. wheat Maize meal porridge Rice Brown bread (1 lb) Corn Rice Whole wheat meal (extraction rate 92%) replaces white bread Whole wheat bread Fiber Source Ca. May 6. possible binder phytic acid Low absorption (4. Rolls containing 10 or 40% bran 2.2 g fiber/kg Wheat bread.3 to 10% bran 36 g bran of wheat Chapathi from whole meal wheat flour or white flour Low availability. Zn. 2 g fiber. Rolls containing 0. Mg Human. possible binder phytate Decreased Fe absorption 104 32 30 10 99 75 78 49 7 27 2387_ch6. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 505 . decreased Fe balance No differences in Fe absorption Decreased absorption with bread Not affected by wheat bran. 28 people Human. 9 people Fe Fe. whole grain.7%). Ca Rat Human.4_fm Page 505 Sunday. serum iron 1. 130. 6 people. whole meal. unrelated to phytic acid or fiber Serum iron fell.Human. enriched with ferrum reductum or ferric ammonium citrate Bread/white. 22 g fiber Oatmeal 20 g wheat bran Whole grain wheat bread 1. 6 people Rat Human. hemoglobin decreased Affected.2% contra 1. 15 g fiber. 221. brown bread. 14 people Human. 10 people Fe Fe Fe Fe. May 6. Low phytate 2. Ca. serum Fe measurement Hgb-repletion test Radioisotope Radioisotope technique. Ca unaltered Serum Fe level fell. brown. possible binder phosphor Serum Fe decreased.2 g fiber/kg. 60. Chemical balance measurement 2. Radioassay Radioassay Blood samples. but not significantly absorption higher from white flour than whole grain (3. blood samples 14 days 6–12 weeks 6 weeks 3 weeks repletion 3 weeks 10 days 17 days Bread 60% of energy Semisynthetic.5 g fiber/kg. serum mineral levels Serum measurements Metabolic balance. 21 people Fe Balance experiment. 2 people Fe Fe Fe Human. Normal phytate level Bread. white bread. 310 g bread/kg diet Self-controlled Self-controlled Metabolically controlled Self-selected 20 g wheat bran 1. phytic acid no influence Fe utilization negatively affected. no correlation with phytate. small effect on Zn retention due to particle size No differences in Fe absorption Geometric mean wheat Fe absorption 5. 12 g bran Monoferric phytate from bran Bran Whole meal from wheat Rice White bread. fecal and urinary Zn Hemoglobin repletion Double radioisotope method.2 (Continued) Description of Mineral Bioavailability Studies from Cereals 2387_ch6. meal + bread 200 g rice (2 mg iron) Controlled. 28 g bread Diet Wheat bran 20% of diet Bran Corn. 3RD EDITION . Zn Fe Hemoglobin repletion Isotope dilution technique. not phytic acid.Rat Human. 506 Fe Fe Human. Fe intake constant Semisynthetic diet. but the soluble fraction more than insoluble Fe four times better absorption from milled than unmilled rice Absorption not influenced by phytic acid Fe better absorption from FeSO4 than bread Low Fe absorption Lowered Fe absorption in wheat Fe from bran Result 35 14 37 51 57 8 105 68 9 46 54 63 28 Ref. pancakes Self-selected. in one iron deficient 19% Relative biological value of Fe the same from bran and ferrous ammonium sulfate Decreased Fe absorption 51 to 74%. exchanged for whole meal bread Bread. 11 people Human. May 6. 180 g bran/kg diet Controlled. 100 g bread Controlled Self-selected. wheat Fiber Source Significantly lower Fe absorption from whole meal bread Fe 50% less available than ferrous sulfate No differences in Fe absorption due to particle size. 60 people Human Dog Human Rat Rat Pig Fe Fe Fe Fe Fe Fe.4. Design 7–9 weeks 24 days 10–15 days Time Controlled.4. NFD up to 15% Whole meal bread Monoferric phytate Unmilled rice Whole wheat bran. 4 people Rat Fe Fe Fe Human. meal. 2001 7:49 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.1%. 1/4 of that of FeSO4 Fe absorption 6. total body counting Radioisotope technique Isotope technique Hemoglobin regeneration technique Double isotope technique Radioiron absorption Radioiron absorption Repletion experiment Radioactivity measurements Exp. 4 people Subject Fe Mineral Table 6.4_fm Page 506 Sunday. 42 people Human. plasma Growth measurements Balance. absorption in femur and serum Balance Balance Phytate Mg 20 days 15 weeks 7 days 4 weeks 20 days 6 days 28–32 days 4 weeks 41 days. adaptation 85 42 72 38 84 70 87 90 81 69 5 93 2387_ch6. Zn negatively affected by fiber and phytate Not affected. Ca. bread. 14 g Whole meal bread (25 g fiber) Whole meal bread. 2 people Human. 7 people Zn Zn. 2-week adaptation period 32 days Controlled. probably due to completely hydrolyzed phytate during breadmaking Negative effect on Zn May be affected. Ca. P Balance experiment Rat Zn. 7 people Zn. 28 people Human. 500 g bread (60% of energy) Controlled. 4 people Human. cellulose. Mg. 40% of energy. feces Growth response. P Cu absorption not affected. metabolic ward. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 507 .Rat Human. P Negatively affected Depressed growth due to phytic acid Negative Zn balance. Mg Human. plasma Balance Balance Balance. 50% of energy from bread Controlled Self-selected + bran Controlled. urine. negative Ca balance but not significant. feces. Zn. 3 people Human Swine Human Human. hemicellulose Wheatflakes. favored P retention. Ca Zn Ca. corn. P Balance Balance. P Zn Zn. 10% and 15% Bread Negative balance of Ca. Zn. Ca. Zn Zn Zn. unleavened Whole meal bread Wheat bread Rice. May 6. caused by phytate No adverse effect on mineral absorption.4_fm Page 507 Sunday. wheat bran Whole wheat bread Wheat bran. wheat Wheat bran. Mg. binder is cellulose Not significantly affected by bran Tendency to increase fecal Zn loss Negative effect on Ca. Zn intake constant Controlled. phytate content 680 and 1040 mg/day Bread. Cu Zn. 350 g Controlled Controlled Controlled Wheat bran (10−20 g). bread and dough Wheat bran 50 g fiber/kg Rice. decreased zinc absorption (phytate possible binder) Bread better Zn source than unfermented dough. Design Table 6.4_fm Page 508 Sunday. phytic acid not the only factor responsible for decreased availability Result 52 34 31 20 103 71 98 Ref. possible binder phytic acid Barley poorer source than mixed infant cereal (wheat. ratio of Zn to phytate important. 3RD EDITION . 66 people Chicken Rat Rat Rat Rat Rat Zn Zn Zn Zn Zn Zn Subject Zn Mineral Growth response Weight gain.Human. no correlation between Zn and fiber/phytate Phytic acid inversely related to Zn availability.2 (Continued) Description of Mineral Bioavailability Studies from Cereals 2387_ch6. May 6. possible binder phytic acid Decreased absorption.oat). rice poorer than standard. 508 Repletion experiment and growth response Femur Zn measurement Growth response Radioisotope technique. more inhibition of phytic acid in cereals than legumes Availability of Zn in grain better than in legumes. Zn availability. corn. wholebody counter Exp. no correlation between Zn availability and fiber/phytate Reduced growth. 2001 7:49 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.4. only 60% Zn utilized. barley Corn Whole grain bread Fiber Source Decreased percent absorption but increased total absorption. femur Zn content 28 days 4 weeks Time Controlled Controlled Controlled Diet Grain Selected cereals Iranian flat bread. no differences on dough with different fermenting time. 4_fm Page 509 Sunday. Ca Zn Ca. K. 27 people Human Human Human Human Human Zn Zn. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 509 . depressed growth. Zn adaptation Not affected Phytic acid naturally occurring in food. 10%. phytate-to-Zn ratio major factor affecting Zn availabilty No significant changes 108 33 18 83 6 74 43 106 11 19 73 4 67 45 2387_ch6.5%. bran (22 g → 53 g fiber) Whole grain Bran (10−20 g) Whole meal bread Different cereals. growth response Balance experiment and 65Zn kinetic 7 weeks 3 weeks 6 weeks 19 weeks 4–9 weeks 5–6 weeks 3 weeks Controlled. Ca. Na. but probably due to higher Mg intake No change or adaptation Negatively affected Ca absorption. possible binder phytic acid Decreased Ca absorption Lowered Ca in serum Apparent Zn absorption and retention in absolute values. Ca. higher from flour with high extraction rate Decreased Ca absorption. May 6. corn Cereal Decreased Ca absorption Decreased Ca absorption Negative Ca balance even if increased Ca intake Decreased Ca absorption. Cl. CI small changes. whole meal) Whole meal bread Whole meal bread Wheat bran 0. 2 people Human.Rat Rat Rat Human. fecal Mg increased. fiber intake increased from 17 to 45 g/day (bran. 6 people Human Human. Na. 15% Rye Breakfast cereals: rice. Mg. K Ca Zn. biological response not directly correlated to dietary fiber in cereals. Mg Ca Ca Ca Ca Ca Ca Ca Rat Zn Balance Balance Balance Balance Balance Serum measurements Balance studies Balance Balance experiment Balance studies Metabolic balance Balance. 230 g bread Controlled Controlled Wheat whole meal bread. unleavened Whole meal bread chapathi Whole meal bread Whole meal bread. 19 people Human. affects Zn metabolism to the same extent as Na phytate If molar ratio > 15. May 6. 14 people Human. 510 Table 6. Design Balance (serum) Balance (serum) Balance Balance Balance Balance Time 5 weeks 4–9 weeks 11 days 3 weeks 28 + 32 days 9 month periods: 2–3 weeks Diet Controlled Controlled Controlled Controlled Flour. 24 g bran per day Wheat fiber. P Subject Human. 3RD EDITION . high phytate intake can cause negative disturbance in Zn and Ca adaptation for serum Fe and serum Zn Negative Ca balance 109 44 58 17 89 59 Ref. unleavened or leavened Whole grain flour (extraction rate 92%) Result No changes in serum Ca No changes in plasma Ca Negative Ca balance Negative Ca balance. Ca. 6 people Human. 31 g fiber Whole grain bread. 3 people Zn. possible binder phytate. 25 people Ca Ca Ca Ca Human. 123 g (standard 106 g) Wheat bran. dephytinization gives better absorption Unleavened bread: negative Ca and Zn balance. 38 g bran per day Wheat bran.2 (Continued) Description of Mineral Bioavailability Studies from Cereals 2387_ch6.4_fm Page 510 Sunday. 2001 7:49 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.Human. 9 people Mineral Ca Exp. 4 people Human. 40–50% of energy Fiber Source Whole meal wheat bread fiber.4. Fe. bran does not cause deficiency No effect on mineral absorption except Zn Mg available Mg available Negative Zn balance. 60% of energy from bread Self-selected Wheat bran. P. negative Mg balance Decreased absorption Ca adaptation 96 82 77 79 110 21 48 94 2387_ch6. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 511 . 16 g/day 3 tsp bran per day Whole grain wheat bread Whole grain wheat flour Whole grain wheat bread Phytic acid Whole meal bread. 12. Cu. Zn. 2 people Zn. May 6. 8 people (ileostomy) Mg Mg Ca. fiber. 55% Mg. negative Ca balance. Mg P. Zn. Mg. Zn.Balance Rat Rat Rat Human. Mg Mg Ca. children Human. Mg. P Ca 2–3 weeks 4 weeks 3 weeks 1940–1948 Controlled Self-selected Wheat flour. the rest of Mg from magnesium carbonate Controlled. 68 people Human. higher absorption from flour than from the same amount of added salt No influence on mineral balance.6 g/day Whole meal bread Mg available. Ca. Fe Metabolic balance technique 13 months Balance Balance Balance Balance Balance Rat Survey.4_fm Page 511 Sunday. wheat bran Wheat bran. fruit enhanced Increased Fe excretion Result 145 156 134 122 112 183 120 119 159 132 124 187 116 Ref. Zn Fe Subject Fe Mineral Whole body radioassay procedure Balance study Hgb regeneration Balance technique Metabolic balance and tissue conc Balance Metabolic balance Isotope study. but no difference within the breads. Ca Fiber not the major determinant of food Fe availability. 50% less available than FeSO4 No effect on absorption of Fe. beans Baked/unbaked wheat bran muffins Extruded maize Bran cereal. whole meal bread 22% DF Cereals. correlation between soluble Fe and repletion.4. Cu. 3RD EDITION . brown bread 13% DF. Ca Fe. vegetables increased from 20–30 g Different wheat varieties Whole wheat. Mg.4_fm Page 512 Sunday. Cu. Fe from corn tortillas. Ca Fe Fe. decreased absorption for Zn. ± juice Diet Wheat bran White bread 6% DF. no effect on Fe bioavailability No change in Fe or Zn absorption Digestibility lowest with wheat bran. Ca Fe Fe Fe Fe. Zn. corn bran. soybean husk Wheat bran Muffins baked with wheat bran Three levels of phytate Maize. Mn. but Fe absorption lowered in bran muffins. Zn. Mn. Design Table 6. 512 Balance study Hgb repletion Hgb regeneration assay in weanling anemic rats Whole body counter Hgb regeneration Exp. fruits.2a Description of Mineral Bioavailability Studies from Cereals (References After 1984) 2387_ch6.Rat Rat Rat Rat Human Human Human Chicks Rats Rat Rat Human Rat Fe Fe Fe Fe. May 6. fiber itself no effect Minerals not affected Baking + organic acids increased bioavailability Extrusion no effect on Fe absorption Cereals significant contribution to available Fe in diet Fe availability reduced with 15% NDF. Mg. 2001 7:49 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. absorption test 12 weeks 10 days 18 days 26 days 45 days Time Controlled Controlled Controlled Controlled Controlled Controlled Controlled Corn tortillas and cooked beans Controlled. no significant relation between Fe and phytate absorption No change Fe absorption higher in control. Zn. corn meal Fiber source Selection of wheat for high protein content. Fe Zn Zn.4_fm Page 513 Sunday. whole grain. Mg. and crisp bread Endosperm. triticale. P Zn Zn Zn Zn Zn. limit availability of Zn to minor degree when Zn is needed Retention lowered by presence of fiber. barley. bone-Zn deposition Whole body counter Whole body counter Balance Isotopic retention from fecal extraction Retention study on rat intestine Erythrocytes utilization of radioactive Fe 9–12 days 3 weeks 25 weeks 4 weeks 4–7 Days Controlled Meals based on 60 g cereals + milk Controlled ad lib Controlled 40 g extruded/ not extruded cereal + milk Dosa + milk Browned and unbrowned corn products Wheat grain Wheat bran. Zn retention. Se Whole body isotope study Radioisotope technique Balance study. Ca. femur consistency of Zn. Fe. Ca. no effect on absorption 146 154 121 173 140 138 149 150 111 123 133 171 2387_ch6. Zn concentration in serum and bone Growth response. oatmeal. 33 people Rat Rat Human Rat Human Rat Fe Fe Zn. polymerization Polyphenol and phytate decreased Fe absorption. barley husk Rye. May 6. Mg. probably due to Maillard products Se and Zn have antagonistic effects on each other Zn absorption higher in plain dosa diet than in raw dosa mixture The total effects of extrusion on bioavailability are small but observable. P Whole body counter Growth response.Rat Human Human Rat Human Human. bran-enriched crisp bread 20 or 30% wheat bran Extruded high-fiber cereals Dosa (rice and dal) Wheat bran and flour Sorghum Maize and wheat fiber Lowered Zn absorption when high phytic acid/Zn ratio. Ca. wheat bread. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 513 . no effect on Fe Zn absorption improved when extrusion was performed after phytate reduction DF and phytate from bran. improvement by enrichment of Zn Food preparation that decreases the phytic acid improves Zn absorption Both sources had more negative effect on Zn than Ca absorption. P Zn Zn. hemicellulose and lignin possible inhibitors of Fe absorption Extrusion cooking. phytate did not appear as a major factor affecting mineral absorption in barley husk Reduced Zn absorption in browned products. whole wheat Whole meal. negative effect on Zn. soybean husk Wheat bran bread. rise mainly due to phytate Result 177 153 158 182 176 113 175 169 178 125 152 136 Ref. 2001 7:49 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. May 6. Cu. Mg. Fe. P. whole body monitor Whole body retention using radioisotopes Metabolic balance Rise in plasma zinc Exp. Cu. P.2a (Continued) 2387_ch6. Fe. wheat bran. + 15% fiber Controlled Controlled Controlled ad lib Controlled ad lib Controlled Controlled Diet 5. tissue Zinc tolerance test. Cu. Fe. except Mg which increased No change in serum or urinary concentration of any of the minerals No effect on mineral utilization. Na. same as milk diet No change in mineral balance Decreased Zn. 15% processed oat husk Hard red wheat bran baked in yeast leavened bread. Mg Ca.4 g/kg/ day) Wheat bran Fiber source Increased loss of Ca. Zn balance lower with oat husk: Fe balance higher with soya husk No effect on Ca absorption Diets did not appreciably affect mineral levels Wheat bran leads to significant reduction in Zn absorption Fe highly available.4. corn bran at 4 of 14% TDF Oat bran Wheat bran Whole wheat bread (enriched with Ca) Sorghum products Wheat bran. bioavailability measured by TIBIA. Fe. Fe Ca Ca. Zn. Mg Zn Zn. guar gum bread Cellulose. Fe Ca Ca. Zn. Na. Mg Balance study Balance Balance study Balance serum. feces. urine Balance Metabolic balance 6 weeks 26 days 3 months 72 days 7 months 27 days 19 days 4 weeks 6 hours Time Controlled Controlled Controlled. K. Zn more available from fermented products with lower phytic acid Ca bioavailability 95% from bread. P. Se Ca. or Mg for the increased DF No impact on Ca intake Fecal excretion for minerals not changed. Cu. 514 Balance study. 3RD EDITION .4_fm Page 514 Sunday. Zn. Mg Ca Ca. phytatereduced bran rice (lowphytate. urine. but not of Zn. 10. oat bran. low-fiber) Wheat bran (0. Mg. 20–45 g DF/day Oat husk.Human Human Human Rat Rat Rat Human Human Human Pig Rat Human Zn. Ca. Ca content reflecting retention Balance study. Zn. Design Description of Mineral Bioavailability Studies from Cereals (References After 1984) Subject Zn Mineral Table 6. Ca absorption influence negative by Zn supplement. Fe. wheat. May 6. Mg. bread w/barley fiber.7 times higher in wheat than corn due to phytase activity Phytic acid in rice bran decreased Ca absorption No effects on mineral balance. wheat — increased loss of minerals. a source of available P and Mg. but might have an unfavorable effect on the utilization of Ca and Zn. Zn P Balance Balance study Balance study Balance study Balance study. Zn P P. Se P. 100 mg/kg) Controlled wheat diets Controlled cereals/milk Controlled Wheat or corn + 0. triticale and wheat higher in phytase than corn Wheat bran. the greater the phytate P availability and the lower the bone mineral disorder.Human Human Pig Pig Pig Pig Pig Ca Ca. oat — higher absorption. Mn. no change of intake. Low-methoxylated pectin has deleterious effect on mineral balances P utilization 1. Mg. oat. vegetables Wheat. improved absorption of Fe.3% P Wheat bran. barley Rice bran In comparison with mixed cereal diet: barley — decreased loss due to lower content of minerals. Mg. urine.4_fm Page 515 Sunday. Zn. addition of Zn (20 vs. 20% DF Corn triticale wheat Barley. Ca.5 and 0. fruits. Zn supplement affected phytase activity in small intestine.35%). corn Fiber 50 g/day. Fe Ca. bread w/whole wheat bran. especially Fe. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 515 . Ca Absorption of P high in wheat. Zn 168 114 167 151 130 186 163 2387_ch6. feces Balance Urinary Ca excretion 6 weeks 3 weeks 6 weeks 2 weeks 1–36 months 21 days Controlled Controlled Marginal Ca and P (0. Ca. mineral absorption different in different cereals The higher the phytase activity of the diet. Cu. Zn. excretion significance greater when consuming coarse than fine bran bread P balance lower in brown rice. Zn. rice. Cu Balance value for Mg. Cu. Mg. other minerals not affected No effect on mineral bioavailability Ref. soybean meal + fiber Diet Whole or dephytinized wheat bran Brown rice Coarse bran bread (35 g NDF. 22 g NDF) Wheat bran. Mg. 160 157 184 115 516 Mineral Table 6. Ca. 3RD EDITION .Human Human Human (10 men) Mg. lower concentration of Zn. Rat P. Design Balance study Balance Metabolic balance Balance Note: OH = oat husk. P. Fe.4. rice bran cellulose Fiber source Description of Mineral Bioavailability Studies from Cereals (References After 1984) Result Phytate more reduced in wheat bran diet than rice. and Cu did not differ by type of bread.2a (Continued) 2387_ch6. cellulose no effect on serum or tissue mineral concentration. K. May 6. Fe. Mg. Ca balance was negative. Cu Subject 15 days 14 days 22 days Time Controlled Controlled Controlled corn. Zn. rice bran no effect on serum levels of Ca. Ca. NDF = neutral detergent fiber. 2001 7:49 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.4_fm Page 516 Sunday. P Various minerals Exp. Zn Na. Fe Mineral Table 6. Zn affected dietary Cu binding Bran high binding capacity for Zn and Fe. EDTA. toasting for 1 h: 19% destruction of phytic acid. percentage ionizable iron was lower in the parboiled rice than in raw rice.4. rice Wheat. Cu > Zn > Fe. cereal. phytic acid. Fe.4_fm Page 517 Sunday. metal binding: pH dependent Result 66 64 104 91 26 32 41 84 12 86 76 47 65 13 Ref. but actual amount the same. Metals released with enzyme incubation. minerals released with incubation for 16 h No correlation with phytate and Fe absorption. oatmeal Fe Fe Fe Zn Zn Zn Zn. twofold increase of ionizable iron when germinating. possible binder: phytate. boiling for 1 h no effect on phytic acid. starch. the only soluble metals of wheat bran associated with phytic acid. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 517 . Ca Fe Fe. and calcium. Hemicellulose. phosphate? Lignin. Cu. Mg Mg.Whole meal wheat bread Wheat. Mg. can to some extent explain decreased availability of Fe and Zn Ionizable iron nearly twice as high when compared with that from chapathi (wheat). hemicellulose: low availability Minerals available for absorption. complexes with wheat fiber. 2387_ch6. iron in wheat and maize strongly inhibited Ascorbic acid inhibits complete binding of ferrous iron. at least 70% of phytate does not exist as Ca5Mg-phytate Only weakly binding. Zn binding pH dependent. Mg Ca Ca. Binding pH dependent. increased binding with lowered pH (pH under 5) Cellulose. iron binding by fiber is strongly inhibited by ascorbic acid. adding of phytic acid lowered the ability to bind Zn Soluble components responsible for 39% of total binding power of wheat bran. and components of bran Wheat bran Wheat bran Cell wall fraction Rice bran Phytate. pectin. high metal-binding capacity. Zn. May 6. Fe Fiber Source Wheat bran and fractions of dietary fiber Rice bran In Vitro Binding Studies Ca. Fe. probably to hemicellulose sequence of binding. phosphate. cellulose. maize Wheat bran Rye bread and whole wheat bread Whole wheat bread Bran. Zn Zn. boiling 1 h in 1 N HCL: 36% destruction of phytic acid Higher availability with increased rising time Dephytinized fiber did bind Zn. Mn. pectin contribute only 10% of total binding capacity Zn. cooking of rice/wheat did not influence ionizable iron Neutral detergent fiber accounts for all binding capacity of iron. no important contribution to Ca binding. lignin and two fractions of hemicellulose high capacity for Zn. citric acid.3 All three metals bound. oatmeal low availability. cysteine. Ca. fiber responsible more than phytic acid Binding to cellulose and lignin not affected by cooking. coprecipitation due to isolation procedure of fiber partly responsible. not affected by toasting. Zn affected significantly by pH and type of cooking (toasting had no effect). hemicellulose. 518 Mineral Table 6. K. cellulose.3 (Continued) In Vitro Binding Studies 2387_ch6. 6.4_fm Page 518 Sunday. P Ash Zn Fe Fiber Source Cellulose important binder. decreasing binding by boiling pH 7 Cornflakes bound more Cu and less Fe than corn grits at pH 5. Mg. Fe Zn. starch. Zn. boiling no effect at pH 6. Cu. dependent on ionic strength and different buffers Ash associated with soluble fiber components Soluble components responsible for 37% of total binding power of wheat bran. Fe Na. the differences found between iron sources prior to baking vanished in final baked product Result 56 36 92 50 102 88 13 Ref. 7 Ash associated with soluble fiber. Ca. Fe Ca. possible binder phytate. Mg. May 6. pH-dependent Ca binding diminished by boiling. pectin contribute only 10% of total binding capacity Insoluble iron generated due to baking process. 3RD EDITION . Zn.4.Bread Wheat bran Corn Wheat bran Different cereals Wheat bran Wheat bread Ca. 2001 7:49 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 60% Ca. phytate potent inhibitor in wheat. 2387_ch6. germ reduced availability. naturally occurring soluble ligands important controlling factors for bioavailabilities 70% of minerals present in kernel associated with soluble fiber fractions. citrate + malate. Cu. malate. Cd. alone and in combination. Cu Ca. citrate + lactate. Zn. isolation procedures Extraction precipitation Isolation procedure In vitro procedure simulating gastro intestinal digestion Solubility study Solubility boiling Diffusing across a semipermeable membrane Digestion under similar physical conditions Solubility during sequential pH treatment Measurements of ionizable iron Table 6. Zn. processing affects availability Germination. 24% Zn. Mg. May 6. Mg. most probably phytic acid/glucans. Ca. Cu Ca. mainly due to refining of maize Degradation of inositol hexa. Zn Fe. lactate + malate. proteinaceous components increases soluble Zn Result 118 127 155 131 147 126 131 161 172 179 135 142 117 Ref. and citric acid prevented precipitation.4. his. Fe and Cu the only minerals that are associated with insoluble fraction Citric acid increased solubility of Zn and Ca.4_fm Page 519 Sunday.3a In Vitro Binding Studies (References After 1984) Minerals associated with soluble fiber fractions. enhanced Zn solubility. potentiometric methods. only 10% Fe associated with phytic acid. Mn. oat Different cereal products Whole oat grain Ca-fortified wheat flake Zn Zn. but not Ca. 60% Fe associated with insoluble fractions. P. fermentation increased soluble Fe Availability of Fe same as for soluble Fe Increased availability after extrusion-cooking procedure. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 519 . Mg. Fe.Wheat flake cereal Cereals with 0–973 mg tannins Wheat bran and whole wheat flour Fe Fe Fe Polished rice Wheat. glucan did not affect Zn and Ca solubility Wheat bran. cyst. filtration of soluble Zn components 31P-NMR spectroscopy. 9% Mg associated with phytic acid More than 30% of rice zinc-solubilized.and penta-phosphates significantly reduced inhibitory effect on Fe availability Acid incubation to form an organic acid chelate with iron improved bioavailability with Fe fortification of cereals Tannins as well as phytates may be responsible for low absorption of iron in Indian diets The different minerals chelated to different DF components.. soak. rye. barley. EDTA. P Wheat roll Fe Fe Fe Fiber Source Breakfast cereals w/wheat bran and germ Sorghum Maize Maize Fe Mineral Method Solubility during sequential pH treatment (pH 2–7) Isolation procedures In vitro digestion. K. separation of gel. but not in the other cereals studied Organic ligand. Zn Ca. between pH 3. May 6. cellulose Barley. rye. Zn Mineral Wheat bran. inolaxol.Cu. the ability to bind was in order Cu (II) > Zn (II) > Cd (II) for cereals barley > oats = rye. Cd.5–5. phytic acid important chelator for oats. 520 Table 6. 2001 7:49 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.4_fm Page 520 Sunday. Zn Cu. Cd. breakdown of phytic acid in oats same as for wheat Phytic acid in wheat interacted strongly with minerals Result 129 166 165 Ref. 3RD EDITION .4. oat Soluble fiber of wheat bran. pure β-glucan from barley no complexing agent Phytase present in oats.3a (Continued) In Vitro Binding Studies (References After 1984) 2387_ch6. whole wheat grain. oat Fiber Source 31 P-NMR Potentiometric method Potentiometric method Method Considerable association found between minerals and soluble fractions of all three cereals. Corn Bread Rice Maize Maize. ascorbic acid increased absorption 10 times. 100 g fish. Fe diffusibility was enhanced by fruit juices Decreased availability Improved availability Tea reduced absorption from 3. egg yolk Fruit juice. 150 g papaya (66 mg vitamin C) Fish or food of animal origin Tea Ascorbic acid Tea. egg Fish Orange juice Ascorbic acid Protein (animal) Milk Various beverages and condiments Fish Vitamin C Fruit. not from wheat Improved nonheme iron absorption Both improved absorption Tea reduced to half. corn flakes Rice Fe Fe Fe Fe Fe Fe Zn Fe Fe Fe Fe Fe Zn Fe Fe Fe Fe Rice Corn maize.8 to 2. orange juice.4_fm Page 521 Sunday. wheat Cereals Whole grain bread Whole grain bread Whole meal bread Rice Wheat bread Whole meal bread Flat bread Breakfast cereal foods Whole wheat flour Processed foods of white and whole meal bread.1%. 2387_ch6. egg no effect Absorption improved with fish Improved absorption Enhanced absorption Could prevent complexation of Zn with phytic acid Enhanced bioavailability Choice of beverage and condiments influence Fe availability in either reducing or enhancing Increased availability Increased absorption Increased absorption Absorption increased 5 times Absorption increased 2 times with meat and fish. maize meal. meat 50 g meat. wheat bran. milk (Ca) Tea. egg yolk inhibitor Fruit juice enhanced absorption. 5 times with papaya Result 150 143 28 3 10 51 31 174 144 101 29 71 95 23 100 22 55 39 15 40 53 Ref. May 6. orange juice increased 2 times. inhibitory effect of tea overcome if ascorbic acid given Enhanced absorption from maize. ascorbic acid Food Addition Effect of Addition of Other Foods on Bioavailability of Minerals Fe Fe Fe Fe Mineral Table 6. bread Rice Maize Fiber Source Rats fed ad libitum Human Human Human Human Rat In vitro In vitro (diffusion across a semipermeable membrane) In vitro (diffusion across a semipermeable membrane) Human Human Human Human Human Human Human Human Human Human Human Human Subject Various carbohydrates Fruit juices Ascorbic acid Ascorbic acid Animal protein.4 Bioavailability higher in diets containing starch and lactose Processing increased bioavailability compared to raw materials.4. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 521 . Soluble Fe increased with milk 2.4 (Continued) Effect of Addition of Other Foods on Bioavailability of Minerals 2387_ch6.4. In vitro solubilization 2. 3RD EDITION . orange juice.Breakfast cereals Cereal-based Wheat bran Corn germ Corn germ (0. orange juice dramatically enhanced. citric acid Food Addition Improved absorption with citric acid Iron absorption inhibited 50% by phytate.5% phytate) Fe Fe Zn Zn Fiber Source Fe Mineral 1. No change in Fe absorption Result 164 164 137 181 117 Ref. ascorbic acid/meat counteract the inhibition Citric acid improved Zn availability Sugar slight increase. May 6. growth development Depletion experiment with rats In vitro Subject 1% citric acid Citric acid Ascorbic acid or meat Milk Sugar. 522 Table 6.4_fm Page 522 Sunday. citric acid more potent enhancer than ascorbic acid 1. Isotope experiments on humans Isotope experiments on humans Rat depletion experiments. 2001 7:49 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Nutr.. Cannell. bile acid excretion and calcium absorption. 4. and Walker. 47. T. Hallberg... 24. J. and Walker.. Effect of wholemeal and white bread on iron absorption. 1977. 20. 22. D. B. Jenkins. and Wickramanayaka. 50. 46. N. 7. Cook. 12. 11. Am. J.. Nutr. R. E. Absorption of iron from maize (Zea mays L. M. Bedouin osteomalacia due to calcium deprivation caused by high phytic acid content of unleavened bread. R. Pearson. 1296.. Clin. S. S... and Wiggins. E. Nutr.. T. Lynch. L. R.2387_ch6.. Br. F.. E. 58. Nutr. Lancet. and Warner. Kies. 24.4_fm Page 523 Sunday. Nutritional implications of dietary fiber.. R. Charlton. 1641.. 367.. 31.. and Mayet... Berlyne. Med.. 29. M.. W. A. 2086. Campell. 1170. A. D. Am. H. G. E. 207. Bothwell. J.. Bothwell.. Nutr. 38. Iron absorption from South East Asia rice based meals. and Clydesdale. M.. D.. Branch. A. and Gueguen.. 14.. Changes in bowel habit. Am. N. Walker. J.. 10. 5. Phytate rather than fibre in bran as the major determinant of zinc availability to rats. Nutr.. Aung-Than-Batu. J. Isotopic exchange of iron between labelled foods and between a food and iron salt. V. Hill. 1977. Charlton.. and Baird.. T. H. 23.. 1968. L. 47.. 1981.. Food Sci.. T. Nutr. 467. F. P. 1976. and Shainkin. Lynch. W. 1977. 13. magnesium utilization by adolescent boys. Rep. R. J. A.. Changes in fecal composition and colonic function due to cereal fiber. The effect of meat protein and dietary fiber on colonic function and metabolism. The effect of prolonged consumption of whole meal bread upon metabolism of calcium. Med. 548. A. J. Callender.. Pahlavi Med.. The effect of heat treatment and particle size of bran on mineral absorption in rats. Milner. Nutr. Nord. Iron absorption from bread.. M. 219.. Br. zinc.. 25. 16.. J. 101. Camire. 16. C. J. M. R. .. Am. M. Effect of pH and heat treatment on the binding of calcium. S.. 21. T.. 32. iron. T. 1981. J. 2. Jivraj.. 235. I.. Br. 261. R.. L.. Nutr. J. W. T. and Monsen.. Iron absorption from cereal based meal containing cane sugar fortified with ascorbic acid. L. I. T. copper. 2047.. Cullumbine. 2. J. Res. Rep... J. M. Davis. 9..) and soya beans (Glycine hispida Mx) in Jamaican infants. 21. 1950. R . Nutr. Clin. J. J. Gut.. 112. and Thave-Toe. 546. P. R. G. S. 34. Am. R. 29. R.. and Venkatachalam.. D. G. 269.. the common cold and iron absorption. R. Davis. E. J. H. 1981. and Flett. May 6. 1893. 26.. S.. J. Drews.. F. 1982. J. Effects of wheat bran on the metabolism of 46Ca and 65Zn in rats. S21. M.. 1. Torrance. 1976. R.. D. 1982. 48... G. Dobbs. J. magnesium. and Jenkins.. Bagheri. J. 26.. J. Food Sci. J. I. 1978. 15. J. B. Hill. V. Derman. S. Effect of dietary fiber on copper. M. 32. Clin. Thein-Than. W. Nutr. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 523 REFERENCES 1. 35.. Br. Caprez. J. 3. G... 4.. The effect of tea on iron absorption. Cummings. Björn-Rasmussen. Houston.. Nutr. magnesium. 1974.. M. 19. Le Motte. and Fox. Nutr. Hristic. Ind. M. Ari. 18. J. Bagheri. Clin. M.. 1982. Relation between phytic acid and trace metals in wheat bran and soy bean.. 6. J. Iron absorption from brown bread. Am. Cummings. Camire. Clin. Clin. J. Mineral metabolism on rice diets. 8. phosphorus and zinc upon the absorption of these minerals in rats. 17. and Warner.. M. A. J. F.. Influence of various amount of bran. Influence of wheat bran diets containing unequal amounts of calcium. magnesium zinc and iron to wheat bran and fraction of dietary fiber. M. S. R. Clin.. 101. L. Nutr. 26. and Nightingale. J. and Gueguen. B. Clin. The effects of phytate on intestinal absorption and secretion of zinc and whole body retention of zinc. 516. T. and Clydesdale.. L. Waterlow. and Mayet. Björn-Rasmussen. J. 1970.. Ellis. J. C. Basnayake. Ashworth. J. J. S. Nutr. V. 15. 910. 1975.. Disler. Br. S. A. Br. 1. J. 16. and Nourmana. 1976. zinc and phosphorus of two young American adults.. 1973. 1468. and Fairweather-Tait. Nutr.. J. J. J.. Sayers. A. J. 1977. 7. Nutr. Cummings. P. 1979. S. Inc. Iron absorption in human volunteers using high phytate cereal diet. M. 193. S.. Reinhold. Cereal Chem. 1962.. Metab. Iron absorption from wheat bread. M. 1979. 1311. Callender. F. Vitamin C. T. 1973. B. Clin. manganese in rats. 1975. Am. Apte. Nutr. 29.. and Morris.. 243. Interactions of soluble iron with wheat bran. 1973. Am. . Int. M. J. Am. Br. 37. 3RD EDITION 27. Johnson. C. J. white and whole wheat breads. Reinhold... Nutr. 48. Cereals Chem.. Biol. Nutr. 464.2387_ch6. E. 53. K. R. Clin. Franz. Hill. Fairweather-Tait. and Nickel. D. Reinhold.. 1977. Med... 243. 31. F. 37. 27. B. W. Bioavailability of iron from wheat bran in pigs.. 446. V. iron and copper in humans. 1974. J... G. Binding of zinc and iron to wheat bread. fecal steroid excretion arid serum iron. 23. Am.. Eakins. 52. J. Las Vegas. P. 1978. Lykken.... E. May 6... Clin. House. 1977. Food Chem. 40.. 1974. Nutr. W. J. Arbeitskries Teir ernährungsforgschung Weihen stephan (ATW) Freising-Weihenstephan. 3. L. J. Elwood. W. 1983. J. Proc. P. zinc and phosphorus by man. Benjamin. Brown. J. 33.. C.. Natur. Iron absorption from South-East Asian diets and the effect of iron fortification.. Fed. J.. Erdman. Suwanik. Biol. W.. 35. G. Clin. Zinc balance studies during wheat bran supplementation. B. 1982.. G. Iron absorption from South-East Asian diets. K. Faradji.. Exp. 50. 724. and Shah. V. Am. 1267. 1. D. and Harland. Food Sci. K.. Lower plasma zinc in humans ingesting wheat bran products. Fed. E. P... 510.. Guthrie. T. Nutr. Am. Frølich.-A. W. Bioavailability of nutrients from food. Br.. E.. 110. 51. J. Iranian flat breads: relative bioavailability of zinc. S. Clin. J. 29. G. 267. F. 1978. W. Colhoun. and Bjorn-Rasmussen.. Am. 1970. J. Br. Ed. J. 1974. D. J. magnesium. M. 1721. A. 21. L. Effect of fiber in corn tortillas and cooked beans on iron availability. C. 1. 1403. J. 31.. B.. 1978. J. 1976. and Dunningan. 43.. Zn availability in grains and legumes. Nutritive value of food iron. J. Walker. Nutr.. D. 1981. 1978. A. R. Hallberg. M. A. M. Exp. Kobza. Effect of bran of blood lipids and calcium. Clark. and Finch.. 1977. G. and Lysø. Nutr. J. Nutr. 1968. F.. 28. M. J. F.. Relative bioavailability of zinc from selected cereals and legumes using rat growth. 254. D. and Abadi. 55.. B.. NV... H. Finney.. Effect of phytic acid on the absorption. D. G... 1981.. 107. Effect of wholemeal and white bread on iron absorption in normal people. Sanstead. 37.. W. 49. N. Garby. 1982. C. Fermentative reduction of phytate in rye. Nutr. 1983. L. 1162.. Biochemical response of late rickets and osteomalacia to a chapathi-free diet. and Hartog. I. 1972. M.. J. J. E. M.. B. Elwood. 2001 7:49 PM 524 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Bjorn-Rasmussen... Harland. J. Schenkel.. M. E. 38. P. Fed. J. W. 23. W.. Eakins. Lantzsch. J.. A. and Asp. and Heggie.. 226. West Germany.. J.. 30. 107. Clin. Fed. 1982. 32. Absorption of iron from chapathi made from wheat flour. M. L. J. 27. 42. F. A. F. Measurement of the total daily dietary iron absorption by intrinsic tag model. Nutr. and Abadi. M. 36. J.... M. Am. and Cummings. 1965. 47. . Mahalko. 30. 47. Clin. P. distribution and endogenous excretion of zinc in rats. Jr.. Nutr.. presented at ACS Meeting... Med... 2272. Nutr. and Steenblock.. Cereal Chem. J. 45. A. 941. 28... J.. Am. The effect of different levels of wheat bran on iron absorption in rats from bread containing similar amounts of phytate. E. Dietary fiber content in cereals in Norway. and Gonzales. Hallberg. 1975. Heaton. Milne. P. and Brown. Nutr.. J. Clin.4_fm Page 524 Sunday. and Wyatt.. Int. D. 1980. Nutr. H. Lancet.. J... Am. 16. M. Am. Stringfellow. Harland. D. Ismail-Beigi. 35. G.. Ismail-Beigi. G. Faradji. Clin. L.. 826. B. and Garby. Soc. S. J. Contemp. B. Effect of wheat fiber on blood lipids. and Feller.. M. P. Am.... P. 112. Manning. B.. Kennedy. and Rubenthaler.. H. and Robinson.. Garcia.. R. H. H. The effect of browned and unbrowned corn products on absorption of zinc. 1980. J. and Pomare. E. 548. Am. McIntosh.. 152. De Kock. B... Fry. W. Faradji. Frølich. 1641.. 44. wheat bran and their components. C. Am. Inman. Layrisse. Effect of cellulose added to diets of low and high fiber content upon the metabolism of calcium. 38. Agric. I.. 58. Layrisse. S. 41. J. K. B... Soc. Heaton. A. P. Hussain.. Proc.. Martinez-Torres. R... C. Kirchgessner.. 3. Human studies of iron absorption from fiber-rich Iranian flatbreads. Absorption of iron from bread. M.. 1. A. F. Vol.. D. H. J.. 48. Welch. Faridi. Nutr.. 31. and Reinhold.. Ford. L. 3. 524.. A... Nutr. Newton. 46. Jenkins. 57.. Rep. 39. 49. 1408. and Van Campen. Garcia-Lopez. 30. D. L. in Trace Element Metabolism in Man and Animals. E. C. R. and Inglett. Foster. 34. Connor. 1982. Br. 460.. Lack of effect on blood lipids and calcium concentrations of young mean on changing from white to wholemeal bread. 1979. D. . Food iron absorption: iron supplementation of food. 185. rice and milk containing various levels of phytate. Sci.. J. Gerontol. Nutr. Rep. 1983. 56. 30. R. Preg. N.. and Clydesdale. 7. and Walsham. 22.. 29. Food Sci. R. Bioavailability of iron in high-cellulose bread. P. C. Nutr. S. 334... Sci. 1976. J. 203. Int. 1979.. 25. K. Clin. and Puyat... Nutr. Mod. D. Cook. and Ellis.. 66. 1981. 45. Ory.. R. M. Bioavailability of zinc in soy-fortified wheat bread. I. J. I. Food Sci. A. Layrisse.. and Eggum. J. 1154. R. B. 24. Complexing of Cal2+ and Mg2+ by cell wall fraction and organic acid. J. In vitro interaction of rice hemicellulose with trace minerals and their release by digestive enzymes. K. 156. Layrisse. J. 851. Clin. 106. 1175.. Effect of domestic preparation of cereals and legumes on ionizable iron. Prabhavathi. L. Persson. E.. J.. Cereal Chem.. 1981. B. Nutr. Oberleas. 1315. B.. Obizoba. M. 80. 1979. I. D.. Lee. 70. McCance. L. 1976. L. Miller. and Fox.. C. 3. J. 18. C. . K. R. Graff. Ranhotra. Qual. 397.. and Normand. Anim. G. 351. The effect of baking on the iron in iron enriched flour... N. E. Food Agric. Nutr. F. E. Isolation of monoferric phytate from wheat bran and its biological value as an iron source to the rat. 1942.. B. and Gelroth. J. 61. Anti-calcifying action of phytate. Int.. M. H. L. M. proteins. T. Lancet. L. 75. Effect of prolonged bran administration on serum levels of cholesterol.. 57.. Rattan. The influence of milling on the nutritive value of flour cereal grains. 32. 1981. and Widdowson.4_fm Page 525 Sunday. ionized calcium and iron in the elderly.... E. Martinez-Torres. A. R. E. J. R. Nutr. 21. Agric. 60. 53. Morris. purines. J. 19. Weizer. Miller. S. 55.. 597. Exp. 21. A. M. M.. Soc. 770. C. Calcium and magnesium nutritional status of adolescent humans fed cellulose and hemicellulose supplement. V. 1977. S. 1500. Fonns-Beck.. 1982. M.. 1412. L. Am. Some effects of phytic acid on zinc availability and physiology of swine. N. P.. Effect of dietary components upon zinc availability. Morris.. K. Hematol.. G.. Study of experimental conditions for most reliable estimates of relative biological value of iron in bread. 63. Fønt-Beck. Effect of heat processing diet mixtures on bioavailability of iron.. 58. Ory.. Cereal Chem. 79. S. J. R. and Ellis. J. Zinc and copper metabolism of human adults fed combinations of corn. 1979. A. and O’Dell. 73. 2. J... 44. Ranhotra. A. Morris. A high fiber diet does not cause mineral and nutrient deficiences. Nutr.. R. J. G.. 363. J. 137.. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 525 54. 64. 38. J.. Fed... and Richards.. Effect of interaction of various foods on iron absorption.. Physiol. E. 1976.. Nutr. 57. 68. A. Plant Foods Hum. Pedersen... 74.. M. Rep. fat and calcium in wholemeal wheaten bread. 1969... R. B. Expanded cereal fortification: bioavailability and functionality (flavor) of magnesium in bread. 1975. 2. Proc... 3. 81. Br. Am. F. Food Chem. 56. and Narasinga Rao. 488.. Phytate-zinc molar ratio of breakfast cereals and bioavailability of zinc to rats. 1980. 1978.. 22. 59. 45.... 101. and Martinez-Torres.. and Morris. 78. J. Ranhotra. 1949. Gastroenterol. H. 1983. L. Molloy. wheat. Simpson.. R. 1980. R. Zinc and magnesium utilization as affected by graded levels of hemicellulose.. J. L. M. J. and Roche. 1981. 59. M. Agric. beans. 753. C. 44. J. H.. and Gilat.. and Jensen. 487... Kies. Persson. R. Lee. Nutr.. C. The digestibility and absorption of calories. 538. Biol. 1979.. Cereal Chem. C. O. I. N. 58. 1971.. cellulose and wheat bran. A. Am. 26. G. 1968. J. 339. P. 69. 65. J. J. 82. C. 449. 67.. and Gelroth. J. J. 109. Food Chem. Bioavailability of magnesium from wheat flour and various organic and inorganic salts.2387_ch6. 1208. Rep. D.. Bioavailability of iron in some commercial variety breads. and Gelroth. Lee. Food Sci. E.. Mineral metabolism of healthy adults on white and brown bread dietaries. Lee. II. Raby.. 915. 109. L. McCance.. Int. 1948. Muhrer. Rye. Agric. 154.. C. Fed. and Fox. Ranhotra. Cereal Chem. Soc. Chemical properties and interactions of rice hemicellulose with trace minerals in vitro. C.. T.. Absorption of monoferric phytate by dogs. E.. 1962. Res. 77. Mod. S. Clin... 72. 549. F. and Jensen. 1979. Am. Food. Bran and blood-lipids... May 6. Lipschitz. S. Loewe. Physiol. and Gelroth. A. G. Kies.. and Normand. O’Dell. Complexing of calcium and magnesium by organic constituents of Yorkshire Fog (Holcus lanatus).. F. Ranhotra. Lee.. E. 42... E. 1971. Mellanby. 71. 62... Raby. Levin.. R.. Morris. M. 76. M. Papakyrikos. Nutr. McHale.. 1979. . 1983. 93. 209. J. Br. 1950. 89. J. L. Morris.. F.. 1981. Int. Nutr. G. A. 102. J. 283. Rep. W. Peacock.. B..... W. and Besson.. L. P.. Nutr. 61(2). Nature (London).. 1973. M. Å.. C... J. 106.. 302... and Björn-Rasmussen. Reinhold. Nasr. J. 84. and Harris. Eds. S.. Andersson. R. P. The effect of phytate and other foods on iron absorption. calcium. J. G. 97. Del Vedovo. and Faradji. S. 283.. 24... 1.. Cooke. 295. J.. 75. Br. F. 1469. J. and Nasr. 157. M. Hasselblad. Shah.. A. Lancet. Reinhold. and Melkersson. M. Hasselblad. 90. 32. I... R. Ecol.. Vol. 1975. H. Robertson. 493. Schweizer. 1. K. J... Jacob. B. 33. Am. 1973. J. B. J. G. J.. A. 95. J. Klevay. M. Academic Press.. A.. and Mayet.. H. Reinhold. soya.. S. The significance of wheat extraction rate. M. The inhibitory effect of bran on iron absorption in man. W. B. R. The role of cereals in the aetiology of nutritional rickets. 31. 1981. Göransson. J. Rendelman. H. Zinc. S. Sandström. G. Absorption of iron from breakfast meals. R.. 104. Bothwell. L. and Ismail-Beigi. 98. Am. 163. Zinc absorption from composite meals. A. 2001 7:49 PM 526 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.. and Björn-Rasmussen. 101. The effect of wheat bran on the absorption of minerals in the small intestine. and Yudkin. Metab. and Hedayati. Cereal complexes: binding of zinc by bran and components of bran. S. Logan... Nasr. and Grobe.-S. A... J. Sandstead. Roberts. Am.. T. J. 91. Dietary phytate as a possible cause of magnesium deficiency. M..... and Hultén.. A. the lesson of the Irish National Nutrition Survey 1943–1948... Clin. The effect of wheat bran on the intake of energy and nutrients and on serum mineral levels in constipated geriatric patients. and Cook. and Garzon. Influence of dietary fiber on trace elements balance. W. J. T. K. New York. T. B. Dintzis. E.. 23. Falkheden. and Mayet. Lahimgarzadeh. W. G. E. Nutr. M.. Reck. S. R.. 92. Food Nutr. F. F. E. 1980. G. calcium. J.. Faradji. R... B. zinc and phosphorus by humans due to increased fiber and phosphorus consumption as wheat bread. 85. Abadi. G. The effects of ascorbic acid supplementation on the absorption of iron in maize. Minerals and phytic acid in the analysis of dietary fiber from cereals. Clin. Reinhold. Am. Nutr. A. J.. I. 17. Inglett. 88. 1983. 103. B. Lahimgarzadeh. 823. B... 31. 99. E. Iron absorption from rice meals cooked with fortified salt containing ferrous sulfate and ascorbic acid. Nutr. K. F. phosphorus and nitrogen in man.. P.. Lynch. A. Br. Walker. Abadi. C. G. calcium and protein content in meals based on bread. Reinhold.. Binding of zinc to fiber and other solids of whole-meal bread. A. 86.. 100. Cederblad. 1. 1384.. 105. Cereal Chem. and Oberleas. C. Frølich. Sharpe.. J. Haematol.. M...2387_ch6. 94. Lancet.. Hedayati.. and Belonje. M.. Clin. A.. 185. F. Br. 12. 286.. Clin. W. 48. Nutr. Nutr. K. G.. Rickets in Asian immigrants. Chariton. Bosaeus. Ismail-Beigi. D. Arvidsson. H.. Lahimgarzadeh. Am. G.. Bothwell. 1982.. in Trace Elements in Human Health and Disease.. J.. 1982. 367.... B. 1974. Charlton.. Rendelman. H.. Reinhold. Fiber vs phytate as determinant of the bioavailability of calcium. B. 87... M. Sandström.. Cereal Chem. S180. Bioavailability of zinc in infant cereals. Effects of purified phytate and phytate rich bread upon metabolism of zinc. Munoz. M. Clin. 1976. F. Nutr. Clin. 2484. Hallberg. Prasad. 2. R.. M. S.. R. Nutr. zinc and iron of bread stuff.. L... 1979. R. 435. 59. 1973. Ford. Nutr. H.. zinc. Sayers. 34. 1981. B. 1982. Reinhold. Binding of iron by fiber of wheat and maize. I. S. and Hedayati. J. J. 34. 37. Sayers. G. and Ismail-Beigi. wheat. 59. Jacob. 41. Nutr. Cereal complexes: binding of calcium by bran and components of bran.. 96.. 116.4_fm Page 526 Sunday. 1132. J.. Nutr. 45... A.... May 6... Hum. calcium. Faradji.. phosphorus and nitrogen in man. Giroux.. 185. C. Decreased absorption of calcium. 310. 1960. Lancet. Cereal Chem. Garcia.. G. M. . M. Sandberg.. 1976. J. 1978. McIntosh. 3RD EDITION 83. T. Nutr. maggnesium. Nutr. Simpson. and Dunnigan. 1973.. R. 1979. P. D. K. 2. J. Lynch. Rossander. Effect of purified phytate and phytate rich bread upon metabolism of zinc. M. R.. 739. phosphorus and nitrogen balances of Iranian villagers following a change from phytate-rich to phytate-poor diets. and Shuey. M. J. Reinhold. 1976. and Ranhotra. Minerals. A. Fairweather-Tait. and Asp. 1988.. Mineral content of brans passed through the human G. 48.. Effect of wheat bran on zinc absorption. A. 38. 126. M.. A. J. 433. 238. Vanderstoep. Cereal Sci. T. 114. J. J. J. 1983. Food Proc. and Gueguen. 117. magnesium and zinc by the growing pig. Cereal Sci. D. Begum. L. Frølich. A comparison between 31P-NMR spectroscopy and a ferric ion method. Widdowson. 1. Effects of product formulation.. K. and Russel. Food Sci.4_fm Page 527 Sunday.. C. Technol. O. Mills. N. T.. L.. T. J. 269. Studies on protein quality and availability of zinc from Dosa. 38. C. J. Res.. N.. K. 1987. 7. S. F. G. 60. M. Am. M. Relative bioavailability of magnesium from mineral and soyfortified breads. Widdowson. Sci. 1942. 151. 73.... 1211. 115. Food Sci. Nutr. L. N. F. 125. R. 38. M. Minerals and phytate in the analysis of dietary fiber from Cereals.. J.2387_ch6.. Food Agric. L. Iron exchanges of adults on white and brown bread diet.. 120. 129. R.. . 1986.. but different amounts of wheat bran on calcium. 1985. Effectiveness of organic chelators in solubilizing calcium and zinc in the fortified cereals under simulated gastrointestinal pH conditions. 1983. Wahlgren. Rep. Technol.. Hum. 123... Ballam.. Weinreich. 25.. R. J. 118. Nutr. S. D. 121. and Minski. P. Comparative effects of wheat bran and barley husk on nutrient utilization in rats.. 112. Acta Med. Nutr. Carlson. 49. 116. 653. Effects of heat treatment and organic acids on bioavailability of endogenous iron from wheat bran in rats. Frølich. J. 23. Englyst. Effect of wheat bran and pectin on the absorption and retention of phosphorus. Preserv. and Petersburg. Nutr. calcium and phosphorus. Br. III. Int. Interactions of dietary calcium with phytates. Clin. 30. S. W. J.. E. 243. N. Brit. Can. 1988... 50. M.. W. R.. L. Dintzis... 119. Noble. 452. J. M.. H. A. Cereal Chem.. Ajit Kaur Labana and Kawatra. and Asp.. G. J. Nutr. Bingham. H... J. Nelson. J. 7. Dieta. Farah. Role of bran in normals. H. D.-G. S. zinc and iron balance in man. and Irving. L. J. 15.. Smith. 1988. and Miller. tract. T.. 1948. R. D. S. Studies in human mineral metabolism. Nutr.. 108. Br. G. Gastroenterology.. Enzymic degradation of phytate (myo-inositol hexaphosphate) in whole grain flour suspension and dough. Food Sci. and Johnson. 1987. Clydesdale. 124. 161. Scand.. B. R.. M. 56... 109.. II. Donangelo. J.. Symss. N. E. Eagles. 1983... 833. M. 705.. Buchowski. processing and meal composition on in vitro estimated iron availability from cereal-containing breakfast cereals. Symss. L.-G. B. 85. Cook. Zinc. Cereal Sci. H.. calcium. J. 202. Watson. and Nyman. 1984. T. Pedersen. 113. 47. 224. Winterringer.. D. Nutr.. E... Nutr. 47.I. Cereal Chem. The effects of breads containing similar amounts of phytate. J. Andersson. and Sandstead. O.. Rev. Fairweather-Tait. J. B. Morck. Fairweather-Tait... 13. I. 1986. 1977. The effect of bread rich in phytate phosphorus on the metabolism of certain mineral salts with special reference to calcium. J. Studies on phytate in oats and wheat using 31P-NMR spectroscopy. S. D.. and Cummings. 588. 1354.. Nutr. 1986. Am. B. 1989. 1983. J.. The effect of phytate and fiber source on phytate hydrolysis and mineral availability in rats. Walker. Lancet. P. 21. 4. S. 1089. 128. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 527 106. J.. J... 1985. Clin. I. Fox. Clin. Inst. C.. Portwood. 122. S. and Sattar. T. 125. Drakenberg. R. 110. May 6.. L. 73. The effect of different levels of wheat bran on iron absorption in rats from bread containing similar amounts of phytate. A. phytate and dietary fiber in different fractions of oat grain. 1983. 1970. J. W. A. 62. 8. I. and Dinesen. Reprod. A. D. phosphates and fats.. 503. Effect of dietary phytate on bioavailability of iron. W. Bagheri. 113. 1982. S. and Drakenberg. W. H. 107.. 307. 1984. Akhtar. D. The effect of extrusion cooking on iron absorption from maize and potato. and Eggum.. 14. M. Iron and zinc absorption in human subjects from a mixed meal of extruded and non-extruded wheat bran and flour.. J. Lynch. and McCance. Navert. 111..... P. and Kitts.. J. Effect of fiber on nonheme iron absorption. F. 1989... L. Nutr. Nutr... 325.. Frølich. Frølich. 127. Hall. L.. and Kirby. 42. Biochem. J..-J. whole grain and bran-enriched wheat crispbreads fed to rats on a zinc-deficient diet. Scheuermann. E. 365. J. D. 83. W. P. 51. Nutr. C. Harrned. Nutr. I. 152. 116. Torrance. 1297. 7955. Hall. and Wyatt. Nutr. 133. J. 189. 476.. and Inglett.. P. The role for mineral availability. P.. H. Cederblad. 1989. Nutr. House. 148. Exp. J. 145. M. 143. D. K. Agric. Ca and Mg in diabetics. Effects on apparent absorption of zinc. 119. Gordon. W. Lopez-Guisa. dietary fiber and minerals in wheat. P.. 165. 135.. 139. Davidsson. J. Anim. Scand. Wheat fiber..... I. J. Z. D. Wetter. and Morris.. 1982. Kivistö. A.. New York. I. E. D. Hazell. Novelli. Dintzis. R.-H. 953. F... R. C. S. J. 132.. Technol. Nutr. Sandstead. R. London. S. D. J. Ink. H. Hallmans. W. Eds.. Bothwell. 1989. S.. L. 1987. J.. S.. 2001 7:49 PM 528 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.. calcium and zinc metabolism of young pigs at different dietary levels. Milne. Br... T. Bezwoda. O. Liu. Garcia. L. and Welch. May 6. L. R. J. Cu. 1990. and Sandström. W. 4. S.. 1984. Nutr. and Johnson. 1989. 1987.. J. Nutr. 144. Johnson. vegetables and fruit. calcium. 1987... and Meuser. Dietary fiber and bioavailability of essential trace elements... Z. 138. M. Relationship of components in wheat bran and spinach to iron availability in the anemic rat. 60. Iron and its availability in maize (Zea mays L. 724. J. a controversial topic. 46. and Welch. Med. 146.. J. 142.. 49. 1986. E.. A.. J. Frølich. Nutr.. and Mayet. Characterization of zinc components on in vitro enzymatic digestion of foods... 1989. Physiol... and Chao. magnesium and phosphorus in humans. C. Influence of food processing on iron availability in vitro from extruded maize-based snack foods. G. and Johnson. Lykken. Bodwell. 1988. Bioavailability of and interactions between zinc and selenium in rats fed wheat grain intrinsically labelled with 65Zn and 75Se. J. and Menke. Sci. G. Morrall. D. 55. 1988. Derman.. C. iron. W. H. G. and Wing. 25. B. Sandberg. Gastroenterol. 72. B... Investigation of the effect of a diet with wheat bran on the metabolic balances of Zn. M. L. 117. Br. S. 30. 741. H. Sandberg. 154. 27.. Harmuth-Hoene.. 43. W. 108. Chung Hua Nei Ko Tsa Chih.... Effect of browned and unbrowned corn products intrinsically labeled with zinc-65 on absorption of zinc-65 in humans.. Reviews and Bibliography. Harland. 1984. Anim. Biol. Food Sci. Kivistö. I. 1989. 37. T. Fiber and mineral absorption. A. 38. 150. T. Hallberg. W. Rao.. Processed oat hulls as potential dietary fiber sources in rats. F..)... Ernaehrungswiss. Chelating properties of dietary fiber and phytate.-S. 129. Extrusion cooking of a high-fiber cereal product. Dubielzig. Influence of various sources on the phosphorus. and Johnson. 118. and Farah. Fiber. in Dietary Fibre Perspectives. L.. J. J... 131. T. B. 526. Improvement of the bioavailability of zinc in wholemeal bread and crispbread. Marcel Dekker.. M. Food Chem.. Food Agric. 1988. 147.. 1987. House. Walter de Gruyter. Eur. H.. 916. 114. 255.2387_ch6.4_fm Page 528 Sunday. Ene... 3RD EDITION 130. and Wu. F. M.. The influence of beverages and condiments on in vitro estimate of iron availability from wheat flour and potato. R. Food Chem.. Mahalko. H. Bioavailability to rats of iron in six varieties of wheat grain intrinsically labeled with radio iron. C. Gillooly. B. W. Downs. MacPhail... Harmuth-Hoene.. Proc. Effect of reduced phytate wheat bran on zinc absorption. Mineral and fiber-vitamin interaction. E.. Gupta... S. Br. 1986. and Erdman. 270. 140. H.. Q.. 151. B. 1987. Berlin. J.. 179. 10. phytase and iron absorption. in Trace Element—Analytical Chemistry in Medicine and Biology. Factors affecting the absorption of iron from cereals. J. . Cederblad. G. Sci. T.. Suppl.... 327. J.. E.. 149. 136. A. 141. T. et al. 137. T. 1988.. W. Hazell. Ed. II. 1989. E. 134. S. Ikeda. 73. Anderson. Adv. F. 244. Garcia-Lopez. A. 62. M. 23rd Nordic Cereal Conf. and Sandström. R. A.. 1988. Chao. 1988. M. Leeds. Effect of meal composition and phytate content on zinc absorption in humans from an extruded bran product. 4431. K. Effect of fiber in corn-tortillas and cooked beans on iron availability. Phytic acid. J. Nutr. 1984. R.. 73. 153. Food Sci.. 27. Vol. J.. The availability of zinc in endosperm. 151. 1988. J. T. oats and barley. Sjöström. 28. John Libbey. and Marlett. A. Lantzsch. Effects of food processing and fruit juices on in vitro estimated iron availability from cereals. Food Agric. 1985. Hazell.. R.. 253. L. A. A. Charlton.. Clin. R. Cereal Sci. 146. E. J.. A.. J. R. in Nutrient Interactions.. Nutr. M. M. Frølich. D. J. 26. Effects of brown rice apparent digestibility and balance of nutrients in young men on low protein diets. Nyman. spinach and non-fat dry milk in rats. P. D. Effects of inositol tri-. 1898. Spiller. 1990. M. N. 846. J. Shah.. Kitagawa.. 1987. J.. and Miniski.. 172. N.. 29. 40.. Rep. J. and Asp. R. A. Judd. Effect of dietary cereal brans on the metabolism of calcium. 241.. (Tokyo). M. 1007.. M.. A. 778.... Importance of cereal phytase activity for phytate phosphorus utilization by growing pigs fed diets containing triticale or corn. 27.. B. S. E.2387_ch6.. 1986. S. almond powder.. and Garzon. M. Urol... Food Chem.. 1015. penta. Ernaehrungswiss. E. J. Bioavailability of calcium from sesame seeds. J. Morris. F. G. Food Sci.. Binding of mineral elements by dietary fibre components in cereals—in vitro. 139. R. 175.. T. 63. 1987. Petro. bioavailability and loss. B. 1987. E. Nunes. Station. and Prabhavathi. 6. Res. Malcolm... 1986. J.. A. Effect of increasing levels of hard wheat fiber on fecal weight. Nutr. Binding of mineral elements by some dietary fibre components in vitro. T. Story. barley. J. Persson. 39. J. 165. S. M. B. Res. B... and Moser. Cottrell.. B. R.. Okuda. 1988. Derler. A. Sandström. J. Effect of different carbohydrates on the availability of iron from rice. Am. 164. 1117. 29. S. L. 33. A. 33. Johnson. 1984...and hexaphosphates on in vitro estimation of iron availability. Rep. and State Univ. Pointillaert. H. Nutr. B. Nutr. Br. Hill. S. and Frølich. J.. 116. 170... S.. Nutr. R. Fed. S. Reddy.. and Reddy. 907. 163. Alton. Fourdin. Nutr.. Effect of citric acid supplementation on the bioavailability of zinc from corn germs.. phosphorus and magnesium in a long term rat study. D. Frølich. Ellis. Y. 160. Pointillart.. 1982. 1989.. T. 2001 7:49 PM BIOAVAILABILITY OF MINERALS FROM CEREALS 529 155. D. M. Nutr. Nutr. Nadeau. May 6. W... B. E.. 169. Phytate phosphorus utilization and intestinal phosphatases in pigs fed low phosphorus: wheat or corn diets.. Proc.-G. Ebisuno. The effect of whole milk. Mineral balance of adult men consuming whole or dephytinized wheat bran. Calcium requirement. M. L. A. A.. Almgren. Nair. P. Önning. R. and Moser.. G. 1989.. Exp. U. N. 445. J. Mason.. A. Okuda. 166.. H.. 8.. J. Spencer. B. tetra-.. B. 473. Effect of acid pretreatment on the stability of citric and malic acid complexes with various iron sources in a wheat flake cereal. 167. and Osis. 159. 178. J... and Svanberg. Mg... and Thomasset. 176. 889. M.. 1987. J. M. A. R.. Studies on the solubility of Ca. H. G. Nutr. Nyman. 717. Food Biochem. 162.. J. 1987. III. Miyazaki. K. Res. J. Trick. Virginia Polytech. S. oatmeal. Stuart. 43. and Yasukawa. 190. 161.. Nutr. H.. Persson. S. 177. G. Brassard. P.. S. B. S. Sandberg. Absorption of zinc and iron by rats fed meals containing sorghum food products.. Rice bran treatment for patients with hypercalciuric stones: experimental and clinical studies. Virginia Agric. G. Pallauf. Saxena. B..... P. milk protein and some constituent amino acids on the in vitro availability of iron from cereal meals. A. Nutr. 157. Food Agric. Inter. Animal Sci.. M. Food Chem. Effect of dietary fiber level and duration of feeding on fiber digestibility and mineral utilization by growing pigs fed high-fiber diets. J. 1990. W. G. Trace elements nutriture of adult men consuming three levels of phytate... 6. Fairweather-Tait.. Wong. Steele. 132... J. Sci. Miyoshi. Eagles. Steele. E. P. 171. and Monge. P.. Inst.. Vitaminol. Fed. Morris. 631.. J. P.. Hamaker. 117. S.. D.. Garcia Estrada. 116. Nutr. vegetables and fruit) for 12 weeks on iron and zinc metabolism... Markwitan.. K.. 159. D. 158. Morimoto. 54.. 81. Ohkawa... and Kornegay.. P. A.. 1985. 96. Proc. M. ascorbate and citrate. J.. 1986. 8.. and Erdman. 46. H. 1984. The effect of moderately increased intakes of complex carbohydrates (cereals. M. Furumoto. H. and Fountaine. D. Zn and Cu in cereal products.. Cereal Sci. J. 169. Lyon. and Clydesdale. Moore. . Poneros-Schneier.. G. 1140. J. 156.. H. 31. Int. K. Narasinga Rao... whole wheat bread. Belonje. II.. 10. 1990. minerals and steroids and gastrointestinal transit time in healthy young women. 174. Zinc absorption in humans from meals based on rye. Nutr.. 173.. Retention of iron by rat intestine in vivo as affected by dietary fiber. Ellis. Whittam. 168. Z. A. N. Kivistö. M. A. Tannin content of foods commonly consumed in India and its influence on ionizable iron. R. triticale and whole wheat... T.. and Cederblad. 207. P. J. Kraemer. and Kirleis. 1991. 1984.. R. 1984.. 54. and Scala. Sci. Fontaine. 1987. 150. Garcia.. 10. Food Sci. and Ebel.. A. Liljeberg. P. J. 1986/1987. and Koishi... J. Reinhold... Clin. 117. J. A.. and Seshadri. T. Moy.. 597. 1988.. A. Carlsson.4_fm Page 529 Sunday. N. 1986. I. Van der Aar. 2001 7:49 PM 530 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. S. J. TEMA-5. Zhang.. I. W. Special Publication No. A. 6.. G.. 1982.. P.. Bricke.. U... D. Nutr.. Effects of dietary fiber on mineral balance in humans. C.. .. Proceedings of a workshop held in Narirobi. Chemical and Biological Aspects.. G. J. 652.. Aspects Med. D. G. 47. and Curtis. Physiological effects of fibre-rich types of breads. R. 142. Hanssen. R. Br. M. Milk’s effect on the bioavailability of iron from cereal-based diets in young women by use of in vitro and in vivo methods.. D. F. Am. W. Nutr.. 180.. 184. Clin. R.. 10G9.. Waldron. E. Ødegaard. Royal Society of Chemistry. W. P.. Svanberg. Wesstra. in Dietary Fibre. 373. J. and Shah.. T. and Berger.. 426. Frølich. G.. 113.. 1985.... L.. 1987. K.. 653. Improved iron availability in weaning foods through the use of germination and fermentation. and Agenaes. Turnlund.. G. Dahl-Jørgensen. 1985. Norwich. Vaaler. B. A. A. P. Walker. Trace Element Metabolism in Man and Animals. 186. van Dokkum. A. C. May 6. 23. D.. S. Molec. Aaseth. and Feldheim.. A..2387_ch6. Ø. Household level food tech.. 1983.. Southgate. Effects of dietary fibers on mineral status of chicks. K. T. R.. Henricks. S. J. 83. Nutr. Kenya. Food Sci... A. The effect of dietary fibre from bread on mineral balance of young men. spinach.. Allen. Smith. W. R. A. B.. 181. 69. and Schippers. Wisker. J. Kretsch. Dietary fibre and mineral metabolism. J. FSTA 22. W. Bioavailability of iron in green peas. Keyes. F. Toma. bran cereal and corn meal fed to anemic rats. 187. 111. J. 182. Johnson. Fahey. 50. 1990. and Sandberg. and Cornforth. Effect on mineral bioavailability. S. F. Symp. K. Mahoney. D. R. J. Schweizer.4_fm Page 530 Sunday.. 3RD EDITION 179. 1987. L. and Fenwick. J. Eds.. 183. 185. T. Food Technol. 451. E. 1990. Trace elements in serum and urine of diabetes patients given bread enriched with wheat bran or guar gum. in Int. It is myoinositol with six phosphate moieties attached and is a ubiquitous plant component often complexed with calcium. May 6. proteins.28 and many ways of removing it from foods have been suggested.25–27 For this reason. not all epidemiological findings are able to establish a clear. legumes. PA constitutes up to 1 to 5% of the weight and serves as the chief storage form of phosphorus.29–32 However. Thompson INTRODUCTION Many epidemiological studies.22–24 Because of its highly negatively charged phosphorylated structure. will be reviewed. PA has traditionally been considered an antinutrient. that has been suggested to have cancer-protective effects.50 © 2001 by CRC Press LLC 531 . and oilseeds. A majority of the in vivo studies were conducted in animal models using purified PA added to either the diet or drinking water. 2001 7:58 PM CHAPTER 6. of the inconsistencies.33–37 In this chapter. PA is able to bind with many divalent cations. along with its potential mechanisms of action. potassium. in vitro and in vivo studies on the effect of PA on cancer risk.21 In many cereals and oilseeds. in the last two decades. nuts.16–20 Differences in environmental factors. Phytic acid (PA) is one fiber-associated component. or magnesium in cereals or a crystalline globule inside protein bodies in legumes.1–3 meta-analyses. Only a few studies have been conducted comparing the effect of purified PA (exogenous) to that of the naturally occurring PA (endogenous) in foods. The limitations of these different experimental approaches to prove the effects of PA are also discussed. However. in part.00+$1. particularly the reduction of cancer risk. and models used may account for some. but not all. several studies suggest that this same reactivity may confer on PA some health-beneficial effects. suggesting that some dietary fiber–associated components rather than dietary fiber per se may.4–6 and reviews7–15 suggest that diets high in fiber and low in fat may be associated with a decreased risk of many cancers. be responsible for the suggested cancer protective effects of high-fiber diets. and starch and consequently reduce their bioavailability. found in high concentrations in cereals.2387_ch6.5_fm Page 531 Sunday. and oilseeds.24. 0-8493-2387-8/01/$0. experimental design. definitive role for all dietary fibers in cancer prevention.5 Phytic Acid and Cancer Mazda Jenab and Lilian U. time periods.44 In the latter experiment. In a number of the above studies.54 Mammary Cancer Cell proliferation in the mammary gland was reduced by dietary PA supplementation (1. 3RD EDITION IN VITRO STUDIES Table 6.46 suggesting that the antineoplastic action of PA may involve the direct modulation of genes controlling the growth and maturation of the cell.38–41 PA caused a reversion of malignant phenotype.51 When provided in the drinking water. a tumor suppressor gene.41 have shown a dose-dependent decrease. tumor size.41 human rhabdosarcoma.55 subsequent experiments36. supplemented in the drinking water. Eggleton45 has shown that PA does not directly activate neutrophil inflammatory events but does enhance immune cell response to inflammatory stimuli.52 When administered to AOM-treated rats up to 5 months post-initiation. PA (2%.. pure PA has been shown to reduce the rate of colonic cell proliferation at early (2–14 weeks)33.53. Colon Cancer In colon cancer studies.57 .43 and estrogen receptor positive MCF-7 and estrogen receptor negative MDA-MB-231 human breast cancer cells.2%).1 summarizes the in vitro cell culture studies showing the antineoplastic action of purified PA on various cell types.2).5_fm Page 532 Sunday.5.50 or in the drinking water. and endpoints ranging from preneoplastic markers such as cell proliferation and aberrant crypt foci (ACF) parameters to tumorigenesis (summarized in Table 6.2).42 human HepG2 liver cancer. supplied in the drinking water) reduced the number of colon tumors (by 58%) and the tumor volume (by 76%). 1% PA has been shown to significantly decrease both the number of tumors and the tumor volume of azoxymethane-treated (AOM-treated) rats when treatment was commenced in the pre-initiation phase. significant inhibition was achieved with 1mM PA after 6 h of treatment. decrease in expression of tumor markers within the cells (HT-29 cells) or change to a “normal” cellular appearance (K-562 erythroleukemia cells). Similar effects have been observed with pure PA provided in the diet. Similar colon cancer protective effects were observed when PA was provided in the diet.49 time points and to reduce various ACF parameters when provided in the diet37. In addition.2387_ch6. a growth inhibitor.53.34. Although an initial study showed only a non-statistically significant decrease in mammary tumors.49 suggesting that PA can have protective effects at both the initiation and promotion stages of colon carcinogenesis. while other studies39.38. with the reductions being more dramatic when the PA was added to diets supplemented with high levels of iron and calcium35 (Table 6. May 6.40 human PC-3 prostate adenocarcinoma. and mitotic rate when compared to the control group.5.47 and late (36–40 weeks)48. i. supplied in the drinking water) again significantly reduced the number of colon tumors. PA inhibited cell growth and increased cell differentiation and maturation of HT-29 human colon carcinoma cells.39 K-562 human erythroleukemia cell lines. and p21waf1/cip1.37. models. 2001 7:58 PM 532 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.51 also found that post-initiation administration (1 week) of PA (2%. ANIMAL STUDIES A strong cancer-protective effect of purified PA has been shown in a large number of animal studies done under various experimental conditions. pure PA added to HT-29 cells has been shown to upregulate the expression of p53.35.56 have shown that pure PA. Pretlow et al. can effectively decrease mammary tumor incidence in dimethylbenzanthracene (DMBA)-treated rats.5.e. 3. increased differentiation and reversion to normal phenotype.40 Neutrophils adherent to plastic or laminin 100.66–10. MDA-MB-231 human breast cancer cells Vucenik et al. no effect of PA on non-stimulated cells PA decreased growth and increased differentiation.0.2387_ch6.0 mM Dose-dependent decrease in growth with resumption after removal of PA from media Blood Eggleton 45 Shamsuddin et al. inhibition of PA breakdown within the cell did not alter growth inhibition Breast.1–10.0 mM Sakamoto et al38 HT-29 human colon carcinoma cell line HT-29 human colon carcinoma cell line 0. increase in differentiation. 8.43 HepG2 human liver cancer cells 1–5. increase in cell differentiation . 2001 7:58 PM PHYTIC ACID AND CANCER Table 6.0 mM Yang and Shamsuddin39 0. stimulated cells increased interleukin-8 production and sustained assembly of F-actin.05% and 0. Mammary Shamsuddin et al.5_fm Page 533 Sunday.0 µM K-562 erythroleukemia cell line 0.3 and 5 mM PA at 3 and 6 days Decrease in cell proliferation and markers of cell differentiation Decrease in cell growth and proliferation at 1 and 5 mM. decrease in expression of mutant p53 protein and increase in p21waf1/cip1 Soft Tissue Vucenik et al.42 Rhabdomyosarcoma (soft tissue sarcoma) cells 0.0 mM Dose-dependent decrease in growth.44 MCF-7.33–20. increase in differentiation Liver 0.25–5.0 mM Increase in expression of p53 and p21waf1/cip1 with 3.1% In presence of PA. 250. intracellular Ca and lower IP concentrations Prostate Shamsuddin and Yang41 PC-3 human prostate cancer cells 0.1 533 In Vitro Studies on the Effects of Phytic Acid on Various Cell Lines Reference Amount of PA Used Study Details PA Effect Colon Saied and Shamsuddin46 HT-29 human colon carcinoma cell line 3.0 mM Dose-dependent decrease in growth.0. decrease in DNA synthesis.0 mM Dose-dependent decrease in growth.5.1–1. May 6. 5. 0% Cell proliferation Cell proliferation Amount of PA Given Colon — Early Risk Markers Disease Parameter Studied Decrease in tumor number. 33 Study Details In Vivo Studies on the Effects of Phytic Acid on Various Cancers Reference Table 6. 14.0% Diet Diet Drinking water Diet N/A Drinking water Diet Diet 1. and 36 wk Adenoma patients Tumor.70 Thompson and Zhang35 Pretlow et al. 36 months Shamsuddin et al. decreased colonic mitotic rate Decrease in tumor number Decrease in number of ACF with multiplicity ≥ 4 at 12 wk No association between fecal PA levels and colon cell proliferation.48 Male F344 rats.2 2387_ch6. AOM. volume (min.0% 1. mineral levels. AOM. 4. 63%). May 6.6.51 Nielsen et al. and volume. 2001 7:58 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. cell proliferation Mode of PA Administration 1. AOM.0% 2. 4 wk Shivapurkar et al. mitotic rate Tumor. treatment commenced 20 wk after carcinogen induction Drinking water Diet Drinking water. tumor Cell proliferation 1. no carcinogen. MNU. SIM ACF.47 Female SD rats. 24 wk Male F344 rats. AOM.5_fm Page 534 Sunday. treatment commenced prior to carcinogen Colon — Tumorigenesis ACF ACF.Male F344 rats.53 Male Wistar rats.5. 2. 13 wk Female SD rats. 40 wk (20 wk of PA treatment) Male CD-1 mice. and colonic mitotic rate. treatment commenced after 12 wk of carcinogen Drinking water. AOM. 3 wk Male F344 rats.0% ACF. or lipid content Decrease in total number of ACF and ACF with multiplicities ≥ 4 Decrease in incidence of ACF Decrease in proliferation of normal crypts but not of ACF Decrease in number of ACF and SIM ACF. DMBA.25% 1. cell proliferation ACF Variable amounts of dietary PA 2. size.7%).34 Shamsuddin et al. 9 and 30 wk Male F344 rats. AOM.0% Drinking water.3 wk Jenab and Thompson37 Challa et al. 3RD EDITION . decrease in mitotic rate Decrease in tumor frequency (34.54 Shamsuddin and Ullah49 Shamsuddin et al.50 Corpet et al. DMH. decrease in cell proliferation and crypt height Decrease in colon cell proliferation Decrease in colonic cell proliferation PA Effect 534 Owen et al.0% 2.2% 0. AOM. 24 wk Male SD rats.0% 2. 19 and 36 wk Nelson et al. AOM. 1.0% 2. DMH.2. decrease in hydroxyl radical formation by PA Inhibit the colon cancer–promoting effect of iron Decrease in tumor number. mitotic rate Tumor Tumor Tumor ACF.0 and 2. 2 wk Female C57BL/6J mice. 34 Male F344 rats.12.0% 0. 5_fm Page 535 Sunday.58 Vucenik et al. decrease in promotional effect of Ca and Fe Decrease in tumor number and volume at 36 wk Decrease in tumor prevalence and size with 0. AOM.4% 2. 8. PA provided prior to carcinogen administration.Female SD rats. increase in number of surviving animals Decrease in tumor incidence.1% PA. 58 Vucenik et al. 1. 38 wk Male F344 rats.0% PA reduced tumor frequency and size 2387_ch6. 1. multiplicity.56 Hirose et al. 35 wk Female SD rats. nuclear aberration ACF.12. DMBA. DMBA.51 Male F344 rats. 45 wk Female SD rats.53 Vucenik et al.36 Vucenik et al. and 36 wk Ullah and Shamsuddin52 Tumor Tumor Tumor Tumor Tumor Tumor Tumor 2.57 Male BALB/c nude mice inoculated with HepG2 human liver cancer cells.0% Drinking water Drinking water Diet Liver NA 40 mg/kg of body weight 0. DMBA.0% 15 mM 2.1% and 1. 4.7 wk Male BALB/c nude mice inoculated with HepG2 human liver cancer cells pretreated with PA. MNU. 71% tumor incidence in mice receiving untreated cells Decrease in tumor weight Decrease in tumor incidence and number Decrease in size of palpable tumors.0% 15 mM N/A Intra-tumoral injection Drinking water Diet Drinking water Diet Drinking water Mammary — Tumorigenesis 1. AOM. tumor Tumor No tumors in mice receiving cells pretreated with PA vs. 3 wk Thompson and Zhang35 Pretlow et al.55 Shivapurkar et al. DMBA. 18 wk Female SD rats.9 wk Female C57BL/6J mice. DMBA. and size Decrease in mammary cell proliferation and nuclear aberrations. PA provided prior to carcinogen administration.0% 0. decrease in tumor multiplicity and burden when PA combined with 15 mM inositol Decrease in number of tumors Non-statistically-significant reductions in tumor frequency. 2001 7:58 PM PHYTIC ACID AND CANCER 535 .2% Mammary — Early Risk Markers Cell proliferation. May 6. 9 and 30 wk Female SD rats. AOM. 29 wk Vucenik et al. 25% Soft Tissue Amount of PA Given Drinking water Peritumoral injection i.0% pentapotassium dimagnesium phytate (equiv. 4 wk Male NIH athymic nude mice inoculated with rhabdomyosarcoma cells.0% Skin 40 mg/kg of body weight 0.61 Vucenik et al.60 Male C3H/Jsed mice with transplanted tumor from FSA-1 fibrosarcoma cells.60 Jariwalla et al. 2001 7:58 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. decreased cell proliferation after 1 wk Decrease in time of tumor appearance. cell proliferation Tumor Metastatic tumor Tumor Tumor Disease Parameter Studied 2. with FSA-1 fibrosarcoma cells. to 8.p. prolonged survival Decrease in incidence and growth rate of cell transplant–induced fibrosarcomas PA Effect 536 Table 6. injection i. injection Diet Mode of PA Administration Decrease in number of papillomas and number of tumor-bearing mice at 3 wk.5. 3RD EDITION . 3 and 22 wk Ishikawa et al.2 (Continued) In Vivo Studies on the Effects of Phytic Acid on Various Cancers 2387_ch6. 2 and 5 wk Female F344 rats injected with fibrosarcoma cells Study Details Vucenik et al.Female ICR mice.9 wk Male C3H/Jsed mice injected i. up to 6. and growth Decrease in number of pulmonary metastases Decrease in tumor growth. DMBA.p. tumor incidence.42 Vucenik et al. May 6.5_fm Page 536 Sunday.p. 59 Reference Papillomas.25% 12.9% PA) 0. 25. May 6. F344 – Fisher344. small intestine. 2. EHEN. 36 wk treatment commenced during initiation stage of multi-organ carcinogenesis Hirose et al. weak decrease in neoplastic lesions in liver and pancreas. forestomach. BBN. ACF – aberrant crypt foci.2′-dimethyl-4-aminobiphenyl. Wide Spectrum. DMBA – dimethylbenzanthracene.5% Drinking water Necrosis and calcification of renal papillae 100–108 wk with papillomas in some of the high-dose rats Takaba et al.0% Diet Sodium-PA increased development of pre. 2001 7:58 PM PHYTIC ACID AND CANCER 537 . DMAB – 3. BBN in incidence or multiplicity but decrease in diameter of mammary tumors.18 dione. esophagus. Mg-PA and pure PA had no effect Takaba et al.and neoplastic urinary bladder lesions. SD – Sprague-Dawley.Male F344 rats TTAD. EHEN – N-ethyl-N-hydroxy-ethylnitrosamine. PA – phytic acid. increase in bladder carcinogenesis with sodium salt of PA Note: Abbreviations: AOM – azoxymethane.0% Drinking water Decrease in hepatic tumors. Bladder.0% Diet No modifying effects when dietary EHEN. kidney.5_fm Page 537 Sunday. DMAB. Tumor 2. DMAB. 62 2387_ch6. Hirose et al. no difference EHEN. colon. 32 wk Tumor 2. no effect in lung.0% Diet Multi-Organ. MNU – methylnitrosourea. DHPN. SIM – sialomucin producing. and Adverse Increase in urinary bladder papillomas.120 Male and female F344.72 Male F344 rats. DHPN. DHPN – 2. Tumor 1.73 Male F344 rats.76 Male F344 rats. BBN – N-butyl-N-(4-hydroxybutyl)nitrosamine. DHPN.2′-dihydroxy-di-n-propylnitrosamine. Tumor 1. 32 wk Tumor 2. N/A – not applicable. and thyroid gland Hiasa et al. TTAD – n-tritriacontane-16. DMAB. 5.5_fm Page 538 Sunday. animals fed 1% PA in the diet may be receiving different amounts of PA than animals receiving 1% PA in the drinking water. Very few of the studies listed in Table 6. making it less available for absorption or interaction with other dietary components or the colonic mucosa. Nevertheless.42 Supplementation of PA (2%) in the drinking water has protective effects in skin 2-step papillomas.36. in the drinking water. HepG2 cells treated with PA prior to inoculation into the mice developed substantially fewer tumors than untreated cells. in such cases it is very difficult to effectively compare the effects of the endogenous PA in wheat bran to exogenous pure PA given in the drinking water. Since most of these studies show positive effects of PA on cancer prevention (Table 6. or starch components in the diet. 2001 7:58 PM 538 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.63 have attempted to make a comparison between endogenous and exogenous PA. Thus. Vucenik et al. in immunodeficient mice. 3RD EDITION Other Cancers Injection of pure PA directly into HepG2 liver cancer tumors inhibited their growth in immunodeficient mice. However. proteins or minerals. as the intake depends on how much they eat or drink.4). May 6. the cancer-protective effect of PA may theoretically differ depending on whether it is provided in the pure form (exogenous) or as a natural component (endogenous) of a high-fiber food such as wheat bran.5. high-fiber diets provide many more phytochemicals than PA and are also associated with reduction in risk of diseases other than cancer.60 Furthermore. Thus. However. may already be tightly bound to the fiber. even though PA was not provided to the mice. Therefore. it is difficult to compare effective PA levels in studies that provide PA in the diet vs.2 provide an assessment of overall amount of PA consumed or information about amount of food or drinking water intake. although the authors matched the level of pure PA in the drinking water of one group to the level of PA present in the 20% wheat bran diet of the other group.61 while dietary supplementation of the same amount has inhibited hepatocellular carcinomas in the liver as well as eosinophilic foci. PA delivered by injection directly into the tumor inhibited growth of human rhabdomyosarcoma.58 PA reduced the growth of rat fibrosarcoma59 as well as subcutaneously transplanted mice fibrosarcoma cells and the number of pulmonary metastases established after their injection.62 Mode of PA Administration Diet vs. to date only a few studies36. Pure (Exogenous) vs. Suggestions have been made that to reduce cancer risk.37.64 However. When given in the drinking water. while an amount of PA equivalent to that in 20% wheat bran (0. Distilled Water A majority of the studies on PA and cancer used pure PA at the level of 1–2% given either in a low-fiber diet or in the drinking water (Table 6. . Hence. supplementing a diet with pure PA may be better than eating fiber-rich foods.5. which are putative preneoplastic lesions.3). an aggressive soft tissue tumor cell line. pure PA may form complexes with the mineral. they provided no data on the total amount of PA consumed by each group nor gave any assurance that the total amounts consumed were equal.36 showed that levels of wheat bran up to 20% were ineffective in reducing rat mammary tumorigenesis. protein. such as wheat bran. as discussed earlier. On the other hand. the ability of PA to bind with other dietary components depends on whether it is consumed concurrently with diet or at different time periods than the diet. endogenous PA in a high-fiber food source.4%) added in pure form to the drinking water caused a significant reduction in tumor incidence.58 Also.2387_ch6. it can be argued that either mode of PA administration is effective. in the pancreas. Food (Endogenous) PA When administered as part of the diet. Table 6.5. the actual intake of pure PA (exogenous) was the same as the intake of endogenous PA. Results showed a significant reduction in the number of sialomucin-producing (SIM-producing) ACF.5). or dephytinized wheat bran with 1.5.0% pure PA (equivalent to the amount of PA in the 25% wheat bran diet).2387_ch6. by all the treatment diets (Table 6. PA by determining the effect of diets supplemented with either 25% wheat bran.5. 1. Injection Amount of PA Used in Study <1% 1 to <2 % 2% >2% 15 mM 40 mg/kg body weight Total 1 6 6 1 0 0 14 2 4 6 1 2 0 15 0 0 0 0 0 2 2 Note: Values represent number of studies using a certain level of PA with each mode of PA administration. Some studies with more than one level of PA intake were repeated in the count. 25% dephytinized wheat bran (wheat bran with the endogenous PA removed). it is still more practical to eat diets containing high-fiber foods than a refined diet supplemented with just one phytochemical. an important early biomarker of colon cancer risk.3 539 Number of In Vivo Studies Using Different Levels and Modes of Phytic Acid (PA) Administration Number of Studies Using Each Mode of PA Administration Intratumoral or Diet Drinking Water I. The effect of endogenous PA was indicated by a comparison of wheat bran and dephytinized wheat bran. In addition. 2001 7:58 PM PHYTIC ACID AND CANCER Table 6. Jenab and Thompson37 tried to differentiate the role of endogenous versus vs. exogenous .0% pure PA. while the effect of exogenous PA in a high.5_fm Page 539 Sunday.P.or low-fiber diet was indicated by a comparison of the 1% PA and the 25% dephytinized wheat bran plus 1% PA diets. In this study. May 6.4 Summary of All In Vivo Studies on Phytic Acid and Cancer Number of Studies Organ and Endpoint Colon Early risk markers Tumor Mammary Early risk markers Tumor Liver Tumor Soft tissue Tumor Bladder Necrosis/calcification Tumor Skin Tumor Multi-organ (except bladder) Total Protective Effects No Effects/Equivocal Promotive/Adverse Effects 8 7 7 7 1 0 0 0 1 4 1 3a 0 1 0 0 2 2 0 0 4 4 0 0 1 4 0 0 0 0 1 4 1 2 1 0 0 2 0 1 Note: Values represent number of studies. a One study was effective only when phytic acid was combined with inositol. Some studies with more than one timepoint or endpoint were repeated in the count. Carcinogenesis. Table 6. DWBPA = 25% dephytinized wheat bran plus 1.57 3.5. 1087. 1998. to its PA. May 6. The removal of PA from the wheat bran caused an increase in the rate of cell proliferation (Table 6.46ab 4.48bc 9. 2001 7:58 PM 540 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.33 15. However.83b 2.69 ± 0.5. they may be acting through different mechanisms.4%) is much less than the amount (1–2%) in most of the other studies listed in Table 6. Reddy et al. in part. addition of 0.64 18. Likewise.91cd 12. suggesting that the colon cancer–protective effects of wheat bran may be associated with many of its components and not just its endogenous PA. DWB = 25% dephytinized wheat bran diet. This differential effect of endogenous and exogenous PA suggests that.01b Number of Sialomucin ACF Labeling Index in the Top 40% ± ± ± ± ± 18. values with different superscripts are significantly different.97 ± 0.58a 1. the control diet.37ab 3.59a 2.0% added PA diet.13b Note: Values are means ± SEM on a sample size of 15 rats per group for aberrant crypt foci data and 24 crypts per rat with 6 rats per group for labeling index of cell proliferation data. L.64 18. the amount of PA tested in this study (0.4% exogenous PA to a 10% dephytinized and defatted wheat bran diet significantly reduced the number of adenocarcinomas vs.4%. U.2.67 51.64b 3.09 54. control group. the control diet.45 ± 1.05. as well as in the Jenab and Thompson37 study cited above.5). although both are effective. and Thompson. but wheat bran. suggesting that the colon cancer–protective effects of wheat bran may be due.55 12.19bc 34. since removal of endogenous PA and addition of exogenous PA did not significantly affect colon tumorigenesis. the results do suggest that PA may not be a strong factor in the protective effects of wheat bran. with its endogenous PA. BD = basal diet. was significantly more effective than the other diets (Table 6.63 found that a diet supplemented with 10% defatted and dephytinized wheat bran had an effect on colon tumorigenesis which did not differ from that of a diet supplemented with a similar amount (10%) of wheat bran with its endogenous PA and oil intact. PA = 1.5_fm Page 540 Sunday.69 ± 1.5.2387_ch6.0% added PA diet.38a 7. p < 0. because the amount of oil added in this study was largely in excess of the amount found in 10% wheat bran. The rate of cell proliferation with the 1% PA diet was significantly lower relative to the control group but significantly higher than the wheat bran group and not different from the other diets. Perhaps the most important finding of this study is that removal of both the oil and PA components of wheat bran caused a significant increase in the number of adenocarcinomas. the 10% wheat bran diet. may be required for reduction in risk of colon cancer. addition of 2% wheat bran oil (an amount in excess of that found in 10% wheat bran) alone or with 0.99d 13. suggesting that wheat bran oil may also be an important colon cancer–protective component of wheat bran.55 54. M. Source: Adapted from Jenab.5 Groups BD WB DWB DWBPA PA Number of Aberrant Crypt Foci and Labeling Index of Cell Proliferation in the Distal Colon of Rats on Wheat Bran or Phytic Acid Diets Number of ACF 64.4% exogenous PA to a dephytinized and defatted wheat bran (10%) diet did not cause any significant changes in colon tumorigenesis compared to a 10% wheat bran diet. In the same study.5).5.82 44. making direct comparisons with the wheat bran is difficult.28ab 3..71 ± 1. However. Since most of the colon cancer–protective effects of wheat bran have been found at levels of intake much higher than 10%. which is the amount in a 10% wheat bran diet. 3RD EDITION PA added to the low-fiber diet significantly reduced the number of ACF and size of ACF vs. 19.65–69 levels of PA greater than 0. WB = 25% wheat bran diet. All the treatment diets also significantly reduced the labeling index of cell proliferation in the top 40% of the crypt vs.93 ± ± ± ± ± 4.16b 3. Although this study was not designed to compare the role of endogenous and exogenous PA. . However. PA can potentially bind dietary proteins and digestive enzymes such as trypsin. thus inhibiting protein digestion and absorption.82 PA may also reduce the rate of .62. HUMAN STUDIES Limited clinical studies have been conducted on the role of PA in cancer prevention. 2001 7:58 PM PHYTIC ACID AND CANCER 541 The results of the above studies suggest that endogenous PA derived from wheat bran. particularly iron. the potential negative effects of PA. particularly diets supplemented with sodium-PA.74 or it may directly affect transitional cell epithelial growth in the bladder. Interactions with Protein and Starch — Enzyme Inhibition and Malabsorption PA may potentially bind important proteins and enzymes within the cell and thus alter its growth characteristics.4). such as wheat bran.5. it is unclear from this study whether the colonic sampling for the rate of cell proliferation was performed at the same time as fecal collection or whether the patients were maintained on the same diets as they were at the time of fecal collection.5. PA has a cancer-protective effect on different tissues.2387_ch6.73 Since most of these studies use the sodium salt of PA.5_fm Page 541 Sunday. It is possible that it is the PA that is broken down within the colon to lower inositol phosphates or the PA that is absorbed by the colonocytes that may be having protective effects and not the essentially unreactive PA that is complexed with other dietary components that is excreted in the feces.70 observed a strong correlation between fecal PA and fecal minerals. the mechanisms behind the action of PA are not clear. Table 6. POTENTIAL MECHANISMS OF ACTION Evidently. the effects of PA may be different in vivo.72 and Takaba et al.75 Furthermore. depending on the concentration of PA. Nonetheless.2) have found that these bladder cancer–promotive effects are observed only with sodium-PA and not with unchelated PA or other PA chelates. as seen in various cancer models under different experimental conditions. ADVERSE EFFECTS While most of the studies on PA show a protective effect (Table 6.81.70 did not relate fecal PA levels to dietary PA intake.78–80 However. an exception is an adverse effect of PA on bladder cancer.71 However. They also highlight the point that the cancer-protective effects of high-fiber diets may stem from more than just one important component. may be cancer protective. it may be possible that the sodium-PA alters urinary sodium ion concentration or pH. Any effects of dietary PA on cell proliferation probably require longterm exposure to constant levels of dietary PA and. as well as exogenous PA added back to wheat bran or to a low-fiber diet. therefore. Thus. PA has been shown in vitro to inhibit the activity of several enzymes such as serine/threonine protein phosphatases77 as well as a number of intracellular proteins.5. clinical experiments in this area will require long-term consumption of controlled diets with known levels of PA and minerals. need to be further studied prior to any recommendations on its intake. there may be little risk from consuming food sources rich in PA. but no relationship between the rate of cell proliferation and fecal PA content in adenoma patients.76 (Table 6. with negative consequences for the bladder.62.72. They attributed the lack of an effect possibly to the presence of adenomas affecting colorectal cell kinetics.2. Owen et al. Hirose et al. factors which may influence the relationships. Within the digestive tract. Owen et al. although several have been suggested. May 6. 5_fm Page 542 Sunday.86 The undigested and unabsorbed starch reaches the colon and contributes to increased fecal bulk or is fermented to short-chain fatty acids (SCFAs) and decreases pH. Iron has been associated with both the initiation and promotion stages of carcinogenesis by enhancing oxidative damage. the inhibitory effects of PA on starch digestibility and absorption have been reversed by the addition of calcium.33 suggesting that chelation of zinc by PA was inhibiting DNA synthesis. perhaps via chelation with PA. indicating that PA is capable of binding iron within the digestive tract. 2001 7:58 PM 542 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. for which zinc is required.2387_ch6. PA is a particularly effective chelator of iron since it does not allow the iron to be soluble.82. These. However.99 Such oxygen-derived free radicals. This is of consequence not only to diabetes but also to colon cancer in light of the insulin hypothesis of colon carcinogenesis.96 The PA is then unable to bind to or interact with starches in the ingested food.. although at different ratios of PA to iron.93–95 The extent of the protein–starch–PA interactions depends on the ability of PA to also interact with other dietary compounds. whereby lesser insulin response is related to decreased tumor growth promotion. 3RD EDITION digestion and absorption of starches. Individuals consuming less dietary fiber have been shown to produce significantly more hydroxyl radicals. may have protective effects on the colon.23 A correlation has been observed between fecal PA and iron content.104 PA has also been shown to completely inhibit iron-catalyzed lipid peroxidation23 and the production of reactive oxygen species103 in vitro.92 Any starch that is not digested and absorbed in the small intestine would also lessen the glucose absorption and potentially decrease the insulin response. May 6. added to a diet or drinking water may be different from the same amount of PA present within the matrix of a foodstuff.e. making it completely bio-unavailable22 and incapable of participating in freeradical-generating pathways.101 The amount of iron necessary for such reactions can be diet-derived. This may also be a mechanism in the recently observed protective effects of PA on human rhabdomyosarcoma42 and HepG2 liver cancer cells in athymic mice. such as iron. the effects of reactive pure PA.24 The SCFA butyrate has been shown in several studies to reversibly prolong the doubling time and to slow down the growth rates of human colorectal cancer cell lines. in turn.102 Thus. Interactions with Minerals and Antioxidative Effects PA can chelate polyvalent cations. whereby PA was injected directly into the tumor daily. re-addition of PA to dephytinized navy bean flour caused a reversion of the increased rate of digestion and blood glucose response seen upon dephytinization. generated in fecal material close to colonic epithelium.84 either by hydrogen binding to starch.91 and to suppress colon tumor formation. and zinc.85 suggesting reduced absorption of dietary starch with increasing levels of PA. indirectly by binding to proteins that starch is bound to. zinc supplementation has been shown to reverse inhibition of colonic epithelial cell proliferation. i. which is preferentially chelated by the PA.89 to induce apoptosis in human colon cancer cell lines90 and in rats fed butyrate pellets.87. removal of iron. PA also catalyzes the oxidation of ferrous to ferric iron.97 which are required for vital cellular functions such as cofactors for enzymes and metalloproteins involved in gene regulation and expression.. may inhibit the production of hydroxyl radicals and may be protective.22. suggesting that .84 or by binding amylase or enzyme cofactors such as Ca2+. i. the latter has only been shown to be true for EDTA-depleted systems.88 to reduce DNA synthesis. Furthermore. may play a role in the etiology of colon carcinogenesis by causing either direct cellular and genetic damage or by promoting the conversion and oxidation of procarcinogens to carcinogens or mitogenic tumor promoters. thus removing the substrate for the Fenton reaction. which may already be bound to other food components. endogenous PA. calcium.103 In vitro addition of PA to a superoxide radical-generating system inhibits the hydroxyl radical formation.98 For example. exogenous PA.100 since the formation of hydroxyl radicals within the colon depends on iron for catalysis.e. For example.70.24 A negative correlation has been observed between PA intake level from cereal and legume foods and the glycemic index. Thus.83. 48 Similar observations have been made in metastatic and mammary models of cancer.106 it is also possible that the cancer-protective effects of PA are being mediated through the lower inositol phosphates or myoinositol itself.108 In light of these observations. and with the mostly anaerobic environment of the colon.55. Since there was no significant difference between any of the treatment groups. in the in vivo situation. with so many roles within the cell. Inositol produced from PA breakdown may also participate in the IP pool.56. and differentiation. This is important. For example. secondary messenger roles.2) have shown that PA can reduce the rate of colonic cell proliferation. This possibility is strengthened by the observation that myoinositol protects against mammary cancer56 and lung cancer.33–35. since it shows that the inhibitory effects of PA may be strong only in the absence of other chelating agents. and since PA can be converted to lower inositol phosphates within mammalian cells. Rimbach and Pallauf 104 observed no effect of PA on liver oxidant or antioxidant status even in a high-iron situation. contribute to lower IP production. although Porres et al. Interestingly. respectively. the antioxidant effects of PA seen in vitro may not be strongly reproducible in vivo.2387_ch6. Jenab and Thompson110 have recently shown that both the endogenous PA in 25% wheat bran and exogenous PA (1%) added either to dephytinized wheat bran or to a low-fiber diet can increase the colonic rate of cell apoptosis and the degree of differentiation. and modulate signal transduction mechanisms.38 If PA can enter the cell. These studies suggest that part of the beneficial effects of PA can be modulated through alterations in cell proliferation. the parent compound of PA. if dietary iron is already chelated or bound. It is evident that PA may act as a preventive antioxidant by chelating and insolubilizing iron to prevent its participation in free-radical-generating pathways in vitro. 2001 7:58 PM PHYTIC ACID AND CANCER 543 EDTA may be a stronger chelator of iron than PA. Participation in the Inositol Phosphate Pool PA has been shown to be rapidly absorbed and to be converted to lower inositol phosphates in various murine and human cells in vitro106 and in vivo.47. . Other Mechanisms A number of in vitro (Table 6. it is conceivable that the dietary modulation of inositol phosphates by PA may play an as-yet undiscovered role in colon cancer prevention or inhibition. indicating both absorption and conversion of PA. endogenous and exogenous PA as well as wheat bran fiber can all be considered to equally affect apoptosis and cell differentiation. PA has been shown in vitro to be absorbed and converted to IP3 by WRK-1 rat mammary epithelial cells.105 have shown that breakdown of endogenous PA in a corn–soy diet by addition of phytase enzyme can cause an increase in colonic lipid peroxidation. May 6.37.52 suggesting that PA affects tumor growth through changes in the rate of cell division. Thus.1) and in vivo studies (Table 6. it can participate in the inositol phosphate pool and be converted to lower inositol phosphates (IP1–5 ).40 Similarly. apoptosis. number. and multiplicity than PA or inositol alone. These measures are important prognostic indicators of colon cancer risk and tumor development111 and thus provide a mechanism whereby endogenous and exogenous PA may be protective of early biomarkers of colon cancer risk.5.5. Thus.109 The full functions.62 with the PA + myoinositol group showing significantly greater reduction of tumor burden. PA added to the media of K-562 erythroleukemia cells in vitro results in a 41% increase and 26% decrease in intracellular IP3 and IP2 .107 The colon cancer–protective effects observed with PA supplementation are strengthened in the presence of myoinositol. but more study is necessary to determine whether this is applicable in vivo.5_fm Page 543 Sunday. and intracellular importance of inositol phosphates still need much further clarification. 139. Negri.. possibly by PA. R. Fiber intake and risk of colorectal cancer. J. L. 636. A. indicating that IGF receptor inhibition. F. Biomarkers Prev.118 Manousos et al. Epidemiol.. 6. J.. R.. C. Kushi. For example. P. 1887. G. Merzenich.5_fm Page 544 Sunday. B. May 6. 2000. fruit. and Greenwald. Robertson. vegetables. 1990. H. Gandini. may be a favorable event.. Meta-analysis of studies on breast cancer risk and diet: the role of fruit and vegetable consumption and the intake of associated micronutrients. REFERENCES 1. 24. and colon cancer in the Iowa women’s health study. Dietary intake of fiber and decreased risk of cancers of the colon and rectum: evidence from the combined analysis of 13 case control studies. D.. Cancer. S. P. J. E. Negri. Chatenoud. and gene expression. Howe. 18. J.. and minerals such as the pro-oxidant iron. Franceschi. Benito. Tavani. 77. Jr. A. A. 1. it appears that endogenous PA may be in part responsible for the cancer-protective effects of high-fiber foods.119 CONCLUSIONS It is evident from in vitro and animal studies that purified exogenous PA provided either in the drinking water or the diet has cancer-protective effects on a variety of tissues. Eur. Steinmetz. M.. 2001 7:58 PM 544 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Cancer Inst..113 Enhanced natural killer cell activity has been shown in carcinogen-treated mice fed with PA and in vitro when splenocytes from normal mice were treated with PA. Folsom. P. J. Trock.. Int. K.. Cancer Inst. J.118 suggest that IGF receptor activation plays a role in later stages of carcinogenesis. Dietary fiber. including its ability to bind starch.2387_ch6. 2. E.. Parpinel.112 PA has also been shown to block TPA or epidermal growth factor–induced cell transformation in JB6 mouse epidermal cells by inhibiting phosphatidylinositol-3 (PI-3) kinase. Cancer. Natl. La Vecchia. and involvement in signal transduction... and Gallagher. R. Whole grain food intake and cancer risk. PA may modulate cellular response at the receptor level. Castelleto. and Potter. PA may have an important role in the reduction of cancer risk by high-fiber diets advocated by major health and government agencies.116. H.. 3. Cancer Epidemiol.114 Such immune system effects may be another means by which PA can exert its actions.. and Levi.. Well-designed. J. and La Vecchia. 4. an enzyme involved in the phosphorylation and hydrolysis of lipids involved in cellular signaling. proteins. 1992. and colon cancer: critical review and meta-analyses of the epidemiologic evidence.. R. PA has also been shown to bind the basic fibroblast growth factor receptor (bFGF). Natl.. E. R.. C. E. 5.. D. 84.. exogenous PA. 36. sterically blocking its binding domain and thus disrupting its signals. Several mechanisms have been suggested for the cancer-protective effects of PA.117 Increased IGF-II levels have been linked to an increased risk for colorectal cancer.... Cornee. J. 667. S. C.. From the limited number of studies conducted on the role of endogenous vs. Am. potential participation in cellular inositol phosphate pools. more research is needed to further distinguish the role of endogenous PA present within the matrix of a foodstuff and exogenous PA added in pure form either to the diet or drinking water and to relate experimental studies to data to be obtained from various human populations. particularly wheat bran.115 which is over-expressed in certain tumors and cell lines. 3RD EDITION PA may also affect the process of tumor promotion by inhibiting 12-O-tetradecanoylphorbol13-acetate-induced (TPA-induced) ornithine decarboxylase activity in HEL-30 keratinocytes and CD-1 mouse skin... 1998. cell-signaling cascades. 1994. . Jacobs. L. Vegetables. a rich source of PA... 650. and Boyle. 1998.. Esteve. R. 7. Lanza. PA has been shown to compete for binding sites on the IGF-II receptor. Bostick.. prospective clinical studies still need to be conducted to validate the effects of PA observed in animal studies.. However. enzymes. M. epidemiology and experiment. S. Technology of removal of unwanted components of dry beans. 26. A.. Jacobs. Levine. U.. M. 75.. E. 1991. and Thompson. 1991. Diet.. 1995. D. 6. 342. 1984.. 436. Hunter. Eur. F. S. 14.. and Hallberg. Crit. 25. D. Nutr. retinol. Nasr.. Martinez. C. 2000. and Eaton.... Chem. R. 1984. Block.. W. Adv. and Sheridan. Lancet.. additives. F. J. J. Verhoeven. Assen. Lancet. 23. 1987. S. high-fiber diet on the recurrence of colorectal adenomas.. Nutr. T. 35. and Sampliner.. and Eddy. P. M. Med. E. J.. J. and Cahill. Plenum Press.. 9.. R. R. Cancer Res. Fuchs. Nutrition and human cancer. 149.. 1149.. 283. 1984. S. L.. Vargas. Effects of phytate on mineral bioavailability in mice. Sathe. R. Cancer. M.. R. Rev. 1. Engl. S. Cooper. 11647. 31. Zhou. J.. Reddy. 1979. Polyp Prevention Trial Study Group. B. E.. J. A. Diplock. L. and Hedayati.). R. J.2387_ch6.. 49. T. Marnett. Cancer.. 4. K. Cancer prevention. Food Sci.. Crit. Med. A. 7. J. 294.. Food Sci. Cancer Lett. S. 1156. Y. J. L. W. Assoc. D. and van den Brandt.. R. J. 16. Nutr. Dietary factors in colon cancer: international relationships. J. D. J. B. Cancer Prev. A. and Salunkhe... J. 1. Lack of effect of a low-fat. vol. F.. Clin. J. M.. B. Kikendall. American Chemical Society. E. 1222. L. P. Lanza. Phytic acid in health and disease.. N. E. .. E.. supplements and drugs. Graf. 110.. E. Lahimgarzadeh. E. Shahidi. A.. J. B. Helzlsouer. 1992. 1994. 54(suppl. and Cho.. P. 1980. J. E. Greenwald. Earnest. D. J. R. Nutr. May 6. R. Stampfer. W.. 1987. Food Sci. Iron absorption: no intestinal adaptation to a high-phytate diet. Sathe. Food Res. 340. G.. J. K. Ritenbaugh. MacLennan. Marshall. J.. 24.. and Eaton. U.. Interactions and biological effects of phytic acid. Cancer Causes Control. R. 2000. van Leeuwen. L. 1.. 2001 7:58 PM PHYTIC ACID AND CANCER 545 7. 1995. minerals and phytic acid in rapeseed flour and protein concentrate. J. Blumberg. Potter. Effect of acylation upon extraction of nitrogen. 127. D.. A. Alfin-Slater. Rev. 747.. 49... 70. and Erdman. 1989. 1. D. 1997. Guillen-Rodriguez. M. F. Nutr. R. 28. 29.. J. 13. Sci. J. Lipworth. Reddy.. 1.. Roe. 15. Cheryan. 12. G... 1997. 17. J. Goldbohm. Crit. 1999. A. Dietary fiber in the reduction of colon cancer risk. Hermus. 169. Giovannucci. Vitamins C and E. Food Sci.. C... and Bright-See. and Newcombe. Rossander.. J. N. Thompson. and Kritchevsky. Burt. Gastroenterol.. 22. 19. 2044s. Summary of the round table discussion on strategies for cancer prevention: diet. and Saura-Calixto. 1. Graf. G.. Reid. S. 114. Wynder. R. 87. 17. Effects of purified phytate and phytate rich bread upon metabolism of zinc. Br. J.. 263. J. 21. A. Nutrition and colorectal cancer.. Washington... Empson. Rev.. and Willett. Epidemiology of breast cancer. Rickard. 1982. P. L. M. E. food. Biol. 21. Nutr. Spence. J. L. 1178... Am.. Bhattacharyya. Engl. Brune. Environmental factors in colorectal cancer: some unresolved issues. Alberts. 1192. B. E. D.. H. 11. L. in Human Nutrition: A Comprehensive Treatise.. 160. Lack of effect of a high-fiber cereal supplement on the recurrence of colorectal adenomas. Rev. Diet and large bowel cancer in three socioeconomic groups in Hong Kong. 32. K. R....... calcium. 542. van’t Veer. New York.... 18.... and Ellis. N. J.. J. L. H. 262. J. Nutr. 1984. J. Schulplein. Rodriguez.... Phytic acid: a natural antioxidant. E.C. J. K.. T.. Cancer.5_fm Page 545 Sunday. R. 7... E. K. Eds. B. Med. Weissfeld. 10. Crit. L. Torre. M.. Reinhold. N.. 2000. M. 114. 27.. Iber.. J.. 1996. Speizer. and Salunkhe. 1980. North Am.. 1988. Hill. G. A. D.. 342.. W. E. Clin. Phytates in legumes and cereals.. Ed. M. Schatzkin. M. Am. and Weisburger. D. Potter. 20. M. 8. in Antinutrients and Phytochemicals in Food. Rosner. Caan.. Lanza. M.. J. 7. Fiber and colon cancer. beta-carotene and dietary fibre in relation to breast cancer risk: a prospective cohort study. J. W. 297. J. Shike. R. K.. K. Higginson.. 30. Colditz. 7. 28.. M. Corle.. Phytic acid interactions in food systems. McKeown-Eyssen. Dietary fiber and the risk of colorectal cancer and adenoma in women. Sturmans. and Simic.. G. A... Engl. C. Morris. 1997. Food. Effects of dietary fiber and phytic acid on mineral bioavailability.. 13. Lance. N. 1973. F.. D.. Bioavailability to rats of iron and zinc in wheat bran: response to lowphytate bran and effect of the phytate/zinc molar ratio. 771. L. J. phosphorus and nitrogen. 495. Nutr. Dorant. D. P. E. Carcinogenesis.. Colon tumor promotion: is it a selection process? Effects of cholate. L. Anticancer Res... G.. and Yang. G... . Yang... Corpet. A. I. 195. 12. Suppression of large intestinal cancer in F-344 rats by inositol hexaphosphate. and Shamsuddin. 95... B. L. 50. 1997. Tache.. 16. M.. 114. 2023. 35. A. P. Vucenik. Y.2387_ch6. Tang. J... and Zhang. and Alabaster. K. Sakamoto.. phytate and food restriction in rats on proliferation and apoptosis in normal and aberrant crypts. Cancer Lett. May 6... Novel anticancer function of IP6: growth inhibition and differentiation of human mammary cancer cell lines in vitro. U. Aberrant crypts correlate with tumor incidence in F344 rats treated with azoxymethane and phytate.. 1992.. 39. G. I. Yang. M. Effect of IP6 on human neutrophil cytokine production and cell morphology. C. 37. 577. Saied. M. Cancer Lett. M.. Y. Cancer. 9. and Pretlow. Carcinogenesis. Phytic acid and minerals: effect on early markers of risk for mammary and colon carcinogenesis. 1975. 1997. T. Zhang. Cancer Lett. Carcinogenesis... Nielsen. S. 1990. A. and Ullah. L. Growth inhibition and differentiation of HT29 cells in vitro by inositol hexaphosphate (phytic acid). 18. Saied. 4083.. 34.. 625. Anticancer Res. Carcinogenesis. A.. M. 1989. A. M. M. 1995. 16. 1993. 1996.. M.. and Shamsuddin. Nutr.. 1989. Shamsuddin. 1989.. G. I. Bansal. A. T. 2041. 45.. G. 11. Carcinogenesis. A rapid dual organ rat carcinogenesis bioassay for evaluating the chemoprevention of breast and colon cancer. Eggleton.. I... K.. Anticancer Res. N. P. A. Z. Vucenik. B.. M... I. Amini. S.. 37. A. O.. U.. 53. Vucenik. I. 7. M. S. 2219. Frost. Interactive suppression of aberrant crypt foci induced by azoxymethane in rat colon by phytic acid and green tea.. K. 16... 1993. A. Rao. Thompson. T. Tantivejkul.. 1997. A. Cole. R. 51. A. A. M. 54. 1055.. Z. M.. 3287. A... Y. 44. and Thompson. A. Pretlow. E. A. 1987. Shamsuddin. Sakamoto. 41. IP6-induced growth inhibition and differentiation of HT-29 human colon cancer cells: involvement of intracellular inositol phosphates. 1992. 46.. N. 10.. and Chakravarthy. Shamsuddin. G. and Reddy. and Shamsuddin. 55. 135. G. Andrianopoulos. Carcinogenesis. Comparison of pure inositol hexaphosphate and high-bran diet in the prevention of DMBA-induced rat mammary carcinogenesis. and Lalwani. Jenab. Carcinogenesis. R. Tanure. I.. Y. 28. 1995. 1087. G. Effects of inositol hexaphosphate on growth and differentiation in K-562 erythroleukemia cell line.. E. 1998.. 1377. M. Effect of phytic acid on colonic epithelial cell proliferation. A.. Anticancer Res.. Anticancer Res. 18. Shamsuddin. Challa.. 1477. 14... and Shamsuddin. M. and Ullah. Yoo. Up-regulation of the tumor suppressor gene p53 and WAF1 gene expression by IP6 in HT-29 human colon carcinoma cell line. 64. 47.. A. Elsayed. S. Shivapurkar. 19. Tantivejkul. 38. 48. Baten. and Shamsuddin. O’Riordan. 36. Inositol hexaphosphate and inositol inhibit DMBAinduced rat mammary cancer.. 1991. K. and Shamsuddin.. 1996. 1988.. Somich. 43. 52.. 169.. 2479.. Shamsuddin.. M. S. Vucenik. Inhibition of rat mammary carcinogenesis by inositol hexaphosphate (phytic acid). 1998. D. G. Carcinogenesis. A.. A.5_fm Page 546 Sunday. and Shamsuddin. Carcinogenesis. Shamsuddin. M.. 40. G. 10.. Carcinogenesis. 3RD EDITION 33.. T... Nelson. Novel anticancer function of inositol hexaphosphate: inhibition of human rhabdomyosarcoma in vitro and in vivo. R. A. L. A. 13. The effect of iron on experimental colorectal carcinogenesis.. 15.. 42. IP6 inhibits growth and reverses transformed phenotype in HepG2 human liver cancer cell line. Thompson. 3711. P. and Shamsuddin. M. I.. 56. 1815. Inositol hexaphosphate inhibits large intestinal cancer in F344 rats 5 months after induction by azoxymethane. 2001 7:58 PM 546 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Ullah. Vucenik. A. IP6 in treatment of liver cancer. 18. D. A pilot study. and Peiffer. 18. M. 1461. K. I. A. Inositol hexaphosphate inhibits growth and induces differentiation of PC-3 human prostate cancer cells. 75. 1998. and Shamsuddin. Kalebic. 100. The influence of phytic acid in wheat bran on early biomarkers of colon carcinogenesis. A. B. 1995. Anticancer Res. and Bird. K. Cancer Lett. L. 1479. Inositol and inositol hexaphosphate suppress cell proliferation and tumor formation in CD-1 mice. U. Dose-dependent inhibition of large intestinal cancer by inositol hexaphosphate in F344 rats. D. 317. Yang. 19. and Vucenik. Ullah. Cancer Lett. M. Carcinogenesis. Anticancer Res. Y. A. A. 1998. G. 1509.. 9. 49.. A. and Misumi. Yang. 1999... Venkatraman. 51. M. 1992. S. A. 75.. J. K. Ito. K. B. Hirose. Shamsuddin. 4792. 1999. A. 3665. Nutr.. Carcinogenesis. 65. May 6. N. 1999. R. H.. Hoshiya. . and Shamsuddin.. M. 59. corn and soybean bran on 1. Modifying effects of phytic acid and gamma-oryzanol on the promotion stage of rat carcinogenesis. and Bartsch... and Goldsmith. 1978. Effect of dietary wheat bran and dehydrated citrus fiber on 3. Effect of dietary phytic acid (phytate) on the incidence and growth rate of tumors promoted in Fisher rats by magnesium supplement.. and Ito. 4091.. L. A. 813. Tamano. K. rice. D. Protective effects of hard red wheat versus hard white wheats in chemically induced colon cancer in CF1 mice. 1989.. 19. 70.. Barnes. 67. D. M.. Toxicol. and Rao.. T.. Kagawa.. Weisgerber. and Leipold. A. U. 18-dione in a rat wide-spectrum organ carcinogenesis model.. Fukushima. M. 60. Imaida. 69.. M. A.. K. 1988... P. 359. Simi. Yoshida. and Shirai. H. 5.. Jariwalla.. 1981. 2000 64. M. Cooma. Pharmacol. D.2-dimethylhydrazine-induced large bowel tumorigenesis in F344 rats. Fukushima. 176. S.. S. Kimura. R. Prender.. Hirose. Modification of n-butyln(4-hydroxybutyl)nitrosamine-initiated urinary bladder carcinogenesis in rats by phytic acid and its salts... Dimethylhydrazine induced colon tumors in rats fed diets containing beef fat or corn oil with or without wheat bran. M. S. Zhang. 2001 7:58 PM PHYTIC ACID AND CANCER 547 57. Cancer. and Shirai. 9. and Shamsuddin. Appl. V. N. S. 39. and Vucenik. 83. Zarkovic. 1997. and Fukushima. and Wideman. Colonic protection from dimethylhydrazine by a high fiber diet.. A.. 149. T. J. and Ito. Klopfenstein. Shirai... Cohen. 63. Cancer Res.. Andrews. curcumin. F. 68. M.. Risio. M. 37.2387_ch6. 1996. B. 1917. 71. S. Gut.. M. Changes in urine composition. S.. 1994.. B. 76. A... Coverlizza.. Hakoi. Intra-tumoral injection of IP6 regresses pre-existing human liver cancer xenotransplanted in nude mice. and Ito... Takaba. 30. and DNA synthesis in male F344 rats in response to ingestion of bladder tumor promoters. A.. Scott.. Vucenik. Gyn. T. Clapp. D. Y. 58.. A. M. Surg. C.. I.. C. F. Clin. 9. Spiegelhalder. M. I.. 499.. H. Lipkin. Anticancer Res. L. Cancer Res. M. 38. Cancer Lett. Bloch.. Kurata. F. Effects of n-tritriacontane16.. S.. H. 503.. and Herman. S. W.2-dimethyl4-aminobiphenyl induced intestinal carcinogenesis in F344 rats. Candelaresi. L. Nakatsuru. Lawson. 19. K. bladder epithelial morphology. S. 32. 19. Anticancer Res. Fabian. Faecal phytic acid and its relation to other putative markers of risk for colorectal cancer. V. Y. 21. B. W. 1999. 1988.. Inhibition of skin cancer by IP6 in vivo: initiation-promotion model. 74.. J.. T. Futakuchi. Hirose.. Preventive potential of wheat bran fractions against experimental colon carcinogenesis: implications for human colon cancer prevention. Cancer Lett.18-dione. D.. 1994.. Wilson. W.. A pilot study.... Food Chem. Res.. 18.. V. Carcinogenesis. tannic acid and phytic acid on the initiation stage in a rat multi-organ carcinogenesis model. and Fukushima. and Mori. Nutr. The role of urinary pH and sodium ion concentration in the promotion stage of two-stage carcinogenesis of the rat urinary bladder.12-dimethylbenz[alpha]anthracene. 1983. 71. Ogawa. R. R. Carcinogenesis.. Hutcheson. 12.. J. Effects of wheat. M. Antitumor activity of phytic acid (inositol hexaphosphate) in murine transplanted and metastatic fibrosarcoma. 66. M. Y. Owen. IP6 in treatment of liver cancer.... M.. 591. and Shamsuddin. 2. Hirose. B.. Nutr. B. N. S. H. 61. chlorphyllin. 1977. S.. 1917. Takaba. P. Mammary tumor inhibition by IP6: a review. Anticancer Res. Takaba. K.. II. Modifying effects of the naturally occurring antioxidants gamma oryzanol.. 147.. 1203. Correlations between rectal mucosa cell proliferation and the clinical and pathological features of non-familial neoplasia of the large intestine. M. Yamada... Oberst. 3671. Patchevsky. G. Shibata. Maziya-Dixon. 1.. 72... S. T. Bertone. Inhibition of mammary gland carcinogenesis by green tea catechins and other naturally occurring antioxidants in female Sprague-Dawley rats pretreated with 7.. N. S. Obs.. Ito. Reddy.. Chen. I. S. Anticancer Res... Toxic... Cereal Chem. B... Vucenik... Ozaki. Ishikawa. A. Fukushima... A. N. M. tannic acid and n-tritriacontane-16. 1991. 1998. 1991. N. Hirose. Z. Z. 99. K. K. Shibata.. Akagi. M. 8. Sabin. 60.5_fm Page 547 Sunday. I.. 1994. 65. dihydroguaiaretic acid. S. 73. M. Hirose.. D.. phytic acid.. Hirose. Tanaka. G... Tomazic.. 62. Reddy. and Berry. 113.. and Rossini. Y.. 3749.. K. Am.. Cancer Lett. 175. 1027... 1986.. Hart. New York. May 6. J. G. Sjoholm. K. The colonic protein fermentation and insulin resistance hypothesis for colon cancer etiology: experimental tests using precursor lesions. Thompson. Nilsson. W... Gut. Caderni. Eds. 92.. E. 93. 498... H.. C. L. Med.. J. 88. Clin. 27838. 471.. 410. A. M. Coradini. Duden. Kinsella. 1993. McKeown-Eyssen. Dietary iron and colorectal cancer risk. S. Transition metals in control of gene expression. B.. U. Eur. 1993. Antioxidant functions of phytic acid. 99. J. Nutr. Larsson. and Feistein. Am. and Mikoshiba. Ikenaka. 85. Cancer... and Dolara. A. 95.. K.. Bendich. O. 1993. 173. 1982. J. G. 38. J. McIntyre. in Dietary Fiber in Health and Disease. Nutr. and Soucie. T. Yamada. Tessitore. D. and Berggren. Cell Prolif... The interaction of coatomer with inositol polyphosphates is conserved in Saccharomyces cerevisiae.. Nutr. 1998. 1996. N.. P. Champaign. L. Borenfreund.. 46. Minneapolis. 1983... 835.. 215. Nutr.. 1987.. V. U. G. 1989. American Oil Chemists Society. 109. J. D. L. and Corpet. 33. Pellizzaro. and Jenkins. O. J. Stimulatory effect of short chain fatty acids on epithelial cell proliferation in rat intestine: a possible explanation for trophic effects of fermentable fiber. Nutr. H. 164. 84... Butyrate production from dietary fibre and protection against large bowel cancer in a rat model. Biochem. J.. Bruce. Giovannucci. Hanlon. J. W. Button. Matthews... Thompson. Yoon.. 96. 89. IL. M. 161. Honkanen.. J. Khetarpaul. Epidemiology of colorectal cancer revisited: are serum triglycerides and/or plasma glucose associated with risk?. A. Nelson. 90. gut microbes and luminal trophic factors. J.5_fm Page 548 Sunday. J. L. O’Rourke. L. 34.. Gupta.. 95. 1995.. E. Slow-release pellets of sodium butyrate increase apoptosis in the colon of rats treated with azoxymethane. Hague. Zwiler. Manning. A. P. E. L. 279. 127. 799. Plant Foods Hum. Science. 41. 3. A diet rich in fat and poor in dietary fiber increases the in vitro formation of reactive oxygen species in human feces. 305. 1996. C. Singh. Bembenek. E. 467.. Eds. R. T. A. 3). H. S.. Egan Press. Biol.. without affecting aberrant crypt foci and colonic proliferation..2387_ch6. in Phytic Acid: Chemistry and Applications.. E.. M. R.. C. 1995. E. D. 2000. 2. O’Haloran.. C. A. and Chauhan. Phytic acid and other nutrients: are they partly responsible for health benefits of high fiber foods. C. Niinobe. Cancer Prev. 8. A. Ali.. Phytic acid and calcium affect the in vitro rate of navy bean starch digestion and blood glucose response in humans. 55. E. 278. 6. 1980.. Huschtscha. Sodium butyrate modulates cell cycle-related proteins in HT29 human colonic adenocarcinoma cells. N.. J. Am. L... Science. 127.. Biochem. Kritchevsky. Free Rad. and Franke.. 3RD EDITION 77. Thompson.. J..... D. M. 1990.. Yamaguchi. R. Abolafio. R. U. MN.. Bode. D. P. Y. B. R.. F. 1995... G. J. P. E. and Krikorian. 139. G. but not DM-20. Ed.. 271. 687. L. 1996. 2001 7:58 PM 548 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. E. Barker. 42. Sakata. 1987. E. U. Myelin proteolipid protein (PLP). S. 58. and Shears. 98. 87... R. 81. M... 12. L. and Jenkins. Pilatus Press. Marimpietri. Inhibition of trypsin activity in-vitro by phytate. D. 1992. Nutritional and physiological effects of phytic acid. 86. 261. Isolation of InsP4 and InsP6 binding proteins from human platelets: InsP4 promotes Ca2+ efflux from inside-out plasma membrane vesicles containing 104 kDa GAP1IP4BP protein. 94. is an inositol hexakisphosphate-binding protein.. 83. Luceri. M. 91. Rabadi fermentation of wheat: changes in phytic acid content and in vitro digestibility. Nutr.. Med.. D. Graf. Carlqvist. Schmid. Insulin and colon cancer. Inhibition of phosphatases and increased Ca2+ channel activity by inositol hexakisphosphate. 97. 100.. Thompson. Mayr. 715. Biol.. B. 386.. Cancer. Free Radical Biol. 80. 61. 30... A. 79. Erhardt. 1997. Cancer Epidemiol. and Young.. L. Constitutive aggregates of intermediate sized filaments of the vimentin and cytokeratin type in cultured hepatoma cells and their dispersal by butyrate.. G. 78. S. Sodium butyrate induces apoptosis in human colonic tumor cell lines in a p53-independent pathway: implications for the possible role of dietary fiber in the prevention of large bowel cancer.. G. Cell Res. .. C.. Exp. 315 (pt. Phytic acid: a factor influencing starch digestibility and blood glucose response. D. J.. Lancioni.. Lim. A. 82. Michell.. M. 1994.. and Eaton. J. C.. and Daidone.. Thompson. I. U. 310 (pt. and Bonfield. W. T. 1992. Chem. The effect of phytic acid on in vitro rate of starch digestibility and blood glucose response. 1997. H. 1). W.. J. Int. and Paraskeva. Biomarkers Prev. C. and Bode. 30. Ag. Graf. Bertorello. W. K... J.. 706. M. Clin. Food Chem... in Food Proteins. G. Gibson.. Br. Cancer Causes Control. Med. reactive oxygen species and colorectal cancer. Cheng.. C... Yamaguchi. Shi. Dong. Trans. 3733. M. 1989. J.. W. H... 116... L. Kitahori.. 110. J. 2001 7:58 PM PHYTIC ACID AND CANCER 549 101. Babbs. Phytic acid inhibits free radical formation in vitro but does not affect liver oxidant or antioxidant status in growing rats. PI-3 kinase in signal transduction. and Estensen. 56. M. Kar. Adami. C. V. Barker. a natural food additive. A. L. B.. 23... 221. H. 721. 861.. Stahl. Exp. cell transformation. 1547. Eur. E. Soc. D. 191. H. J. 783. N. and Lupton. Food Chem. Carcinogenesis. Degalle. 169s. E. Y. and Shamsuddin. P. 103. J. Dietary intrinsic phytate protects colon from lipid peroxidation in pigs with a moderately high dietary iron intake. Chang.3-trisphosphate is a product of InsP6 dephosphorylation in WRK-1 rat mammary epithelial cells and exhibits transient concentration changes during the cell cycle. J.. S.. Predictive value of proliferation. D. 30A. 30... 140.. O. Carcinogenesis. 1997... Biol. Free radicals and the etiology of colon cancer.. Ullah. J. 21.. Z. Shamsuddin. Jenab.. Huang. M. 107.-Y.. 1999. Suppression of colonic cancer by dietary phytic acid. 11. W. Free Rad. Phytate.2387_ch6. W. Inositol-phosphate-induced enhancement of natural killer cell activity correlates with tumor suppression. 106. Tomazic.... Sullivan.. R. S. and Ma. Inositol hexaphosphate reduces 12-O-tetradecanoylphorbol-13acetate-induced ornithine decarboxylase independent of protein kinase C isoform expression in keratinocytes. Exp. 1999. and Shamsuddin. 105. J. T. and Brison. and as a target for chemoprevention of cancer.. 19. C. Lamonerie.. Porres. IGF-I and IGF-II in relation to colorectal cancer. M. Fu. Res. 104. A. V. Toxicol.... K. 1995..... Dev.G. and Palczewski. Tzonou. G. H. 128. Hanash.. R. 1999. . C..... J. and Eaton.. A. F. Int. Carcinogenicity study in rats of phytic acid ‘Daiichi’. and Mitchell. Wattenberg. [3H]Inositol hexaphosphate (phytic acid) is rapidly absorbed and metabolized by murine and human malignant cells in vitro. 115. and Pallauf. 1990. I. R. R. R. A. O. M. Biol. L. 18. A. 5132. W. A. J. S. Quirion. O. F. Proc. 80. L. Anticancer Res. Biochem. 1994. 2000. 1995. Cancer Res. Nutr. V. 1993. Bosetti. A.2. McKeehan. C.. Cancer Lett.. Cancer. Nutr. Binoux. 118... Neuroreport. 1950. Rudnicka-Nawrot. Cancer. Roneker. C... 1790. S. Insulin-like growth factor II in the pathogenesis of human neuroblastoma. 1995. suppl. 83. Carcinogenesis. Spiegelhalder. W. 117.... 3743. Manousos. C. Kirk.. C. 124. H. 1999. 5. X. A. and Lei. J. Pond. and Bartsch. K. Cell. Morimoto. 108... 1999. Wright. J. Med. and Nishioka.. 625.. Biol.. 114.. 111. R. 1994. J. Rimbach.. 1595. K.. 102. R. Graf. Trichopoulos. 1992.. 147. Vucenik. 19.. Kan. Souglakos. U. E. and Belury. Y. Inositol 1. Pathol. Constitutive or inducible overexpression of the IGF-2 gene in cells of a human colon carcinoma cell line. J. 117. C.. Chapkin. Anticancer Res. Hiasa. W. An interaction between inositol hexakisphosphate (IP6) and insulinlike growth factor II receptor binding sites in the rat brain. M.. and Mantzoros. Baten. Nutr. Am.5_fm Page 549 Sunday. M. S. Cancer Prev. K. 216. Chatzidakis. J. H. In Vitro Cell. 8. and Thompson.. 113. 112. 109. G. Soc. 119.. Metabolism and cellular functions of IP6: a review. Chemopreventive effects of myo-inositol and dexamethasone on benzo[a]pyrene and 4-(methylnitrosoamino)-1(3-pyridyl)-butanone-induced pulmonary carcinogenesis in female A/J mice.. 1998. cell differentiation and apoptosis.. 105. 342. Morrison. M.... and Parent. 10. 15. A.. Castle. W. P. 2. W. Owen. Phytic acid in wheat bran affects colon morphology.... R. 1998. Nakaoka. and Feldman. J. M. M. 19. Y. 1996. 120. B. May 6. 1994. Konishi. Inositol hexaphosphate (InsP6): an antagonist of fibroblast growth factor receptor binding and activity.. S41.. Nickel. Lavialle. differentiation and apoptosis as intermediate markers for colon tumorigenesis. May 6. 2001 7:58 PM .2387_ch6.5_fm Page 550 Sunday. 2387_Section 7_fm Page 551 Sunday. 2001 6:06 PM SECTION 7 Definitions and Consumption . May 6. 2387_Section 7_fm Page 552 Sunday. May 6. 2001 6:06 PM . fiber was given no more than a single line in nutrition textbooks and was simply referred to as “roughage” and measured as crude fiber. probably for all the wrong reasons.2 Thus. there have been nearly 8000 Medline entries on the subject of dietary fiber. fiber was launched from merely being roughage to a substance with many possible therapeutic and preventative roles. In 1984.50 © 2001 by CRC Press LLC 553 . fiber’s role in the prevention of colon cancer has been seriously eroded by three recently published studies8. Unfortunately. Painter. 2001 7:59 PM CHAPTER 7.2387_ch7. All previous studies carried out on populations with elevated serum lipids were summarily discounted and forgotten by the public. 0-8493-2387-8/01/$0. when excerpts from a study published in the New England Journal of Medicine 7 appeared on the front page of the New York Times and showed that the addition of oat bran to the diets was ineffectual. In a like manner. there were several attempts by the industry to increase the fiber content of the diet through addition of fiber to products. Kellogg’s took the bold move of using advice on the intake of dietary fiber from the US National Cancer Institute and placing it on the back of its bran cereal boxes. as well. Claims were disseminated through advertising and through a best-seller entitled The 8-Week Cholesterol Cure.1 In 1970 the dietary fiber hypothesis was born with the return from practicing in rural Africa of three British physicians.4 Furthermore. the scientific community.1_fm Page 553 Sunday.10 which said that fiber did not affect polyp recurrence and did not prevent cancer in women. These men observed that the rural Africans did not have many of the diseases that plagued the West. news about the soluble fibers in oat bran also reached the public. During this time.9. the so-called “oat bran craze” was crushed instantly. In the 1970s and 1980s. May 6. and Trowell. Burkitt.1 Consumption of Dietary Fiber 1992–2000 Julie M.3. and industry has waxed and waned. The messages not only increased consumption of Kellogg’s cereals. 6 Oat bran became sought by the public and was being added to a vast array of food products in a wide range of doses from the insignificant to the efficacious. but it also had the effect of increasing consumption of all bran cereals. However.5 In this same time frame in which wheat brans and bran cereals were touted as a cancer preventative.00+$1. the widely touted study had used subjects with stellar diets and normal blood lipids. and they theorized that the high-fiber diet not only exerted positive effects on the gut directly but had many other systemic effects. Jones Prior to 1965. This campaign to increase consumer awareness and increase bran cereal sales was highly effective. fiber’s popularity with the public. In the 30 years since this hypothesis was proposed. Dietary fiber was not commonly found as an item in Medline prior to 1970 but was described by Hipsley in 1953. it eventually ushered in the era of allowed health claims and nutrition label changes. Dietary Guidelines for Australians86 — Eat plenty of breads and cereals (preferably whole grain). hemicelluloses such as arabinoxylans and arabinogalactans. gums. and associated substances that are resistant to digestion in the alimentary tract of humans. as fiber data for various countries may not be comparable. Table 7.2387_ch7. cereals.81 American Heart Association — 25 to 30 g per day from foods.1).82 American Cancer Society — Choose most of the foods you eat from plant sources. the latest definition includes all nonstarch polysaccharides resistant to digestion in the small intestine and fermentable in the large intestine. obesity. there were many attempts to develop methods that would accurately measure what was not digested in the human small intestine.80 The average American currently eats 12–17 g of fiber a day. including both soluble and insoluble fiber. lignin.29.87 FIBER DEFINITION While there is agreement that fiber is important nutritionally because of its physiological function and its non-absorbability. National Cancer Institute (NCI)84 — 25 to 35 g/d.1 North American Organizations and Their Fiber Recommendations Organizations Which Specify a Fiber Amount American Dietetics Association — Americans should eat 20–35 g of fiber each day. modified celluloses. cancer. Eating 3 g a day of soluble fiber from oats or 7 g a day of soluble fiber from psyllium has been shown to lower blood cholesterol levels. (This fact is important. This added materials such as modified celluloses. how it can be measured. May 6. and non-insulin-dependent diabetes mellitus in the United States.” After the advent of the fiber hypothesis. and what most closely approximates what actually happens in the human gut. the only measure of dietary fiber was “crude fiber. and .1. mucilages. pasta. such as breads. oligosaccharides. Non-starch polysaccharides include celluloses. therefore. despite the fact that most health promotion groups and government agencies recommend that fiber intake be at the very minimum 20 g/day and some recommend up to 35 g/day (Table 7. Part of the difficulty stems from lack of agreement on what is to be included. lignin. grain products. gums.83 Eat five or more servings of fruits and vegetables each day. The US Preventive Services Task Force — Recommends that Americans increase fiber and fruit/vegetables intake as well as lower dietary fat and cholesterol to reduce prevalence of heart disease. hemicellulose. to ensure nutrient adequacy. The definition then came to be limited to those materials measured by AOAC method 985.1_fm Page 554 Sunday. American Diabetes Association — Daily consumption of a diet containing 20–35 g of dietary fiber from a wide variety of food sources is recommended. or beans several times each day (no specified quantity). especially whole grains. definitions as well as methods of analysis vary from country to country. cutin. 2001 7:59 PM 554 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and suberin. modified cellulose. and fruits. and pectins and associated minor substances such as waxes.1. not supplements. vegetables (including legumes). oligosaccharides. Thus. and polyfructans such as inulin. Organizations Which Recommend Fiber-Containing Foods But Do Not Specify Amounts USDA Dietary Guidelines 200085 — Choose a variety of grains daily. there has been no agreement as to its definition. stroke. 3RD EDITION Thus. Eat other foods from plant sources. pectins. Only about 1/4 of this is soluble fiber. there has been a change in consumer beliefs about including fiber in the diet. In the 1970s the agreed-upon definition in the US was as follows: Dietary fiber was defined as the remnants of edible plant cells including polysaccharides. rice.11 In 1976 the definition was widened to include all indigestible polysaccharides.) When the fiber hypothesis was in its infancy. In 2000 a committee of members of the American Association of Cereal Chemists (AACC) agreed upon a definition that expanded somewhat the various compounds that are not absorbed and exert a physiological effect. hypertension. the average American is eating only 3–4 g of soluble fiber — below the recommended amount of 5–10 g. Thus. This definition defined a macro constituent of foods which includes cellulose. 2001 7:59 PM CONSUMPTION OF DIETARY FIBER 1992–2000 555 mucilages.15 Intakes did not vary across age groups in NCFS. Furthermore. using Southgate values. Dietary fiber includes polysaccharides.2387_ch7. and associated plant substances. The other important difference between the earlier definitions and the AACC definition is that a physiological component is included. Diets high in fiber usually have more vitamins. FIBER INTAKES Fiber Intake as a Marker Many studies indicate that fiber is a marker of a good diet. The new definition allowed for materials that behave analogously to dietary fiber to be included. The committee wanted the physiological measures to be included. cutin. oligosaccharides. US General Population Depending on the specific population group and the actual method of calculating dietary fiber. non-polysaccharide material that is bound to the plant cell wall is also part of the definition. the average dietary fiber intake for adults in the US is 11 to 15 g/d. those individuals with high intakes of fiber or whole grain diets are more likely to exercise. and suberin. studies conducted by the National Cancer Institute (NCI) determined the mean dietary fiber intake in the US adult population (>19 years of age) to be 13.1 g/d were determined if fiber values compiled from the literature were used. minerals.5 g/1000 kcal) at every age. As with the earlier definition. and less likely to smoke. it may mean that the diet is low in saturated fats and higher in fats deemed beneficial. the term “analogous carbohydrates” was added to the 2000 definition. and/or blood cholesterol attenuation. Thus.5 g/1000 kcal) than men (5. The wider definition will require a change in the method in order to capture substances such as the oligosaccharides.15 . because that way any newly formed analogous substance would not only need to be structurally similar to existing substances but would also need to have a beneficial physiological effect in order to be included in the fiber label analysis. attributing that fiber itself is a preventative agent is indeed difficult. one of the reasons that teasing out the role of fiber in disease prevention is so difficult is that there are too many confounders. women consumed more dietary fiber (6. Also. lignin. which showed that the average American woman ate 13. Fiber may have more functions than those listed in the current definition. but the evidence for these other functions is still inconclusive. Dietary fibers promote beneficial physiological effects including laxation. more likely to use supplements.12 Analysis of data13 from The National Food Consumption Survey (NCFS) of 1987–1988 showed an average intake from 3-day food records of over 9000 respondents at 12–13 g/day. and phytochemicals. the NCI data showed some differences in intake by group. vegetables. May 6. In fact. Any natural food diet that is high in fiber is also high in plant foods. and/or blood glucose attenuation. Such material includes lignin waxes. and cereals. Lower mean intakes of 11.14 Likewise. On a per 1000 kcal basis.1_fm Page 555 Sunday.3 g/d. A nearly identical value was determined for intakes in the National Health and Examination Study (NHANES II). Other functions can be added in subsequent iterations when the data present a clearer picture. Thus. most populations fail to meet the recommendations. Oligosaccharides of various lengths are also included in this definition. The AACC definition is as follows: Dietary fiber is the edible parts of plants or analogous carbohydrates that are resistant to digestion and absorption in the human intestine with complete or partial fermentation in the large intestine.13 In contrast. Despite overwhelming data that diets high in fiber and plant foods are useful.2 g/d. fruits. 3% in the free-living elderly. fiber intake was calculated to range from a median of 12. 3RD EDITION The net analysis of all these studies is best captured by a histogram analysis of the NCI data. which showed that 90% of the population failed to meet the bare minimum fiber intake of 20 g/d. In epidemiological data from 43.19 Eating food not prepared at home may also be a contributor to the low fiber intakes. US Army recruits ate approximately 4. and nuts and seeds.21 Data from Specific US Cohorts Fiber intakes of various population segments have been assessed by a variety of methods. potassium. Table 7. 20 g more fat. and 400 mg more sodium in the high-eating-out group (6–13 times per week) (Table 7. Breakfast cereals made a smaller contribution to the total fiber intake. vegetables. Currently over half of the food dollar is spent on food eaten away from home.1.6 2903 Source: Clemens et al. Breads and cereals contributed 33% of the fiber intake. It was noted that it would be possible to more than double this amount to 9. those who ate the recommended minimum of 20 g or more of dietary fiber per day had diets that contained more iron. 2001 7:59 PM 556 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. the overall mean intake appears to be slightly higher than the population as a whole and would not be consistent with the finding that there was little difference in intakes across population age groups as indicated by the NCFS data.5 3299 1769 60.18.1_fm Page 556 Sunday. especially potatoes.5 g/1000 kcal if all whole grain choices were made. magnesium. according to data derived from analysis of a 6-day weighed military ration.2387_ch7. May 6. The rank-order of sources of fiber for the free-living elderly also varied from that of all adults sampled in NCFS. followed by bread and fruit. selenium. Those Who Don’t Mean calories Mean fat (g) Mean sodium Eat Out Often Eat Out Seldom 2057 79. 7%. and zinc. Elderly In a sample specifically assessing fiber intakes of free-living elderly aged 60–78 years. Data gathered for the US Army showed that although higher fiber choices are possible. the mean reported fiber intake was 18. As anticipated. many diners tenaciously hold onto the idea that eating out is a treat and that it is not necessary to be concerned about nutrition. A recent study20 showed that there was no increase in fiber. they are not selected. despite often gargantuan portion sizes which delivered an average of nearly 300 more calories. were the principal sources of dietary fiber. A factor may be that these individuals were free-living and perhaps were healthier and had greater control over their food choices than the segment of elderly in institutions.20 Despite the common occurrence that eating away from home has become. 22%.4 g/d in . legumes. copper.2 Comparison of Intakes of Selected Nutrients of Consumers Who Dine Out Frequently vs. NHANES II data showed that vegetables.3 g/d.000 male health professionals who filled out food frequency questionnaires of 131 foods. 21%.17 Fiber Intake in Institutions and Restaurants Those living in situations of institutional feeding or using food service outlets may have differing fiber intakes.1 g of fiber per 1000 kcal. Legumes were important sources for some populations and ranked first for several age–sex–race categories. fruits.1.16 In this sample.2). more popular cereals do not offer significant increases in dietary fiber. fiber intake as assessed by dietary records24 in the normal weight group was 18. and less carbohydrate. where vegetables and fruits provided the most fiber in 1977–78 and bread provided the most in 1987–88. However. In one study.7 g/d and lean men consumed 27. but the total amount of fiber was lower.25 Another study of 203 men showed that those weighing the most ate more fat.29 In this case. The intervention goal was to increase this number to 18 g/1000 kcal.29 Several studies show that the diet contains a smaller percentage of fiber as children age. The intake gap between males and females widened as age increased. the fiber intake of US children also fails to meet the various recommendations.2 g/d in the lowest quintile and 9. 2001 7:59 PM CONSUMPTION OF DIETARY FIBER 1992–2000 557 the lowest quintile to 28. or 5 g of dietary fiber per 1000 kcal. with intakes varying from 1.2 g/d in the lowest quintile to 7.8 g and 24. 45% of the 4–6-year-olds met the age + 5 rule and only 32% of the 7–10-year-olds.22 As was seen in the NCI data for the general population.9 g/d in the highest quintile. Similar patterns were seen in the Nurses’ Health Study with over 88. According to analysis of data from NFCS on over 2500 children.2 g/d and the highest quintile 11. While the total intake increased slightly from age 10 to 15 years. Large cohorts of various-age children in the Bogalusa Heart Study found the average fiber intake by 24-hour recalls was 12 g/d.1_fm Page 557 Sunday. vegetables provided the most fiber.48 g/d for 10–17-year-olds.8 g.2387_ch7.7 servings of fruit and vegetable per day. In the National Cancer Institute (NCI) Polyp Prevention Trial. Obese Fiber intake and obesity seem to be inversely related. Lean women consumed 22.31 the Bogalusa Heart Study29 and the NFCS sample28 males were more likely than females to have higher fiber intakes.1 g/d.29 . had higher fiber intakes per 1000 kcal than white children.7 g/1000 kcal. Black children. In this cohort.28 In that time period.9 g/d in the highest quintile.7 g/d).5 g of dietary fiber per kg of body weight for children and adolescents. A similar trend with respect to changing food patterns was seen in the NFCS.28 In the North Dakota 15-year-olds. Their obese counterparts consumed 15.26 Children and Adolescents The American Academy of Pediatrics recommends 0. over 90. Grains and ready-to-eat cereal consumption did increase during this period.8 g/d.28 55 to 90% of American children failed to meet AHF requirements for dietary fiber. the participants ate on average 3. the dietary fiber intake decreased as the body weight increased.7 g/d and 20. or an average of 9.28 The 1991 US Department of Agriculture Continuing Survey of Food Intakes by Individuals (CSFII) generated similar findings. The American Health Foundation (AHF) recommends that children follow the age + 5 rule for the minimum dietary fiber intake and no more than age + 10 for the safe range.0 g/d.23 the average baseline dietary fiber intake was reported to be 10–20 g/d.30 In the Bogalusa study the actual intakes varied from 10. while it was lower in both the moderately obese (13.8 g and cereal fiber contributing 1. Comparison with past data on fiber intake showed a decrease in intake from the 1977–1978 CSFII survey to the survey 10 years later. an 8-year-old should try to get 8 + 5 or 13 g/d of dietary fiber. While 60. many of the newer.2% of the children aged 2–5 years in the NFCS study failed to meet the age + 5 rule. there was a decrease in the consumption of fruits and vegetables. Thus. respectively.3% of the girls aged 12–18 years failed to meet this rule.65 to 13.9 g/d.3 g/d) and the severely obese (13.000 subjects. May 6. Another study showed the same trend but higher fiber intakes for all groups. Fruit provided the least dietary fiber. less fiber. As seen in adults. aged 10 to 17. Cereal foods provided 2. with the lowest quintile ingesting a median of 3. the total amount of fiber decreased by 17 years of age.27 For example.7 g/d in the highest quintile. with median daily dietary fiber intake ranging between 9.0 g to 4. 3 g/d.2387_ch7. These substantially higher values may reflect differences in the sample and measurement techniques and changes in dietary patterns. niacin. Spain.31 Similar results occurred in North Dakota teens participating in the 24-hour recall collection. folate. These NSP intakes were reported as higher than those from a 1977 survey but fell short of the recent government-recommended population mean of 18 g/d. cereals. Mean intakes in the UK are higher than mean intakes in the US Part of the reason for a higher intake is the tendency to eat more bread and cereal products. and consumed less fiber. which includes Denmark. of the 8677 girls and 7525 boys aged 9 to 14 years. nuts. A study in the UK using 7-day weighed food records showed that the fiber intake of 472 dietitians and adult members of their households was 38 g/d. Italy.38 . the values are far lower than values reported in 1985 based on a 7-day food record from a small sample of omnivores and vegetarians. For women aged 25–69 years. The NSP intake of a sample of 739 men aged 40–69 was 15. but diets that satisfied FGP recommendations also tended to satisfy nutrient requirements. so values are not always comparable. folacin. In this sample there were no clear differences by age group.33 Fiber Intakes in Europe It should be pointed out that values for fiber are defined differently in various countries.37 There appeared to be a difference in the readiness of various subgroups to adopt nutrition information and change their fiber intake. and seeds.31 Surveys of college students do not show that they fare much better. and iron. iron. Fiber intake of dietitians and members of their households was reportedly much higher than of the population as a whole. UK and Ireland Until 2000. Those who ate according to the age + five rule consumed more breads.5–16. B12. May 6. B6.4 g/d using a validated food-frequency questionnaire that focused on carbohydrates. 17% ate together with their families never or only on some days. Family behavior also impacted fiber intake. In the UK a 78% extraction rate is used rather than a 70% extraction rate used in the US Some differences may be due to measurement. phosphorus. where just over a fifth of the respondents reported consuming more than 20 g dietary fiber daily. fruits. and C.36 In this sample omnivores consumed an average of 23 g of dietary fiber per day.3–15. fiber in the UK was measured as nonstarch polysaccharide (NSP). France. thiamin. the NSP intake was 14. ate fewer fruits and vegetables. Greece.35 However. zinc.32 Of those surveyed. This group of children and adolescents ate foods with higher glycemic loads. Only 8% of the students consumed the minimum recommended number of servings for all food groups in the Food Guide Pyramid (FGP). In the group with higher fiber intakes. vegetables. In the families involved in the Nurses’ Health Study. according to the European Prospective Investigation into Cancer and Nutrition (EPIC). riboflavin. 2001 7:59 PM 558 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Black housewives living in the UK were more likely to respond to various public health programs and messages and report change in dietary fiber intake than their white counterparts. Great Britain. including vitamins A. These diets were shown to be higher in vitamins A and E. Asians were least likely to change their dietary intake of components such as fiber with public health campaigns and publicity. diets were more nutrient dense and were associated with greater likelihood of adequate intake of several key nutrients. and calcium.34 Differing methods of measurement may also be used. magnesium. magnesium. Vegetarians were reported to eat an average of 37 g/d and vegans 47 g/d. Over half consumed less than 15 g of fiber daily. legumes. Germany.1_fm Page 558 Sunday. 3RD EDITION Several studies have shown that those children who met the fiber recommendations had better diets than those who failed to meet the recommendations. 45% failed to meet the fiber recommendations. and Sweden. A small difference may also be due to differences in the extraction rates commonly used for bread flour. the Netherlands. 46 Increased socioeconomic status in Finland did not improve diets or fiber intake. called ASPCC. below the recommended range of 30–40 g for French adults. Thus. 2001 7:59 PM CONSUMPTION OF DIETARY FIBER 1992–2000 559 Teens and Children in the UK and Ireland In a study of Irish secondary school youths (ages 12–18). The foods that were predominant contributors of dietary fiber were bread and potatoes. Irish diabetics ate more dietary fiber than the general population. was done in 1993–1994.0 g/d based on the 7-day weighed record and 18. the first national dietary survey.39 Thus.1 g/d. Dietary data collected in 1992 and 1993 on nearly 500 children were compared with records on diets of children collected in the 1950s. One study compared diets of nearly 500 men and women in the UK and France.1_fm Page 559 Sunday. these diets would have higher mean fiber intakes than UK adults. but this is lower than recommended by the European Association for the Study of Diabetes (EASD). and dietary fiber. Overall. making those diets more in line with current recommendations on healthy eating. In both countries. they ate less bread.43 In France. the average daily consumption of dietary fiber of French adults is about 16 g. as French respondents scored significantly better for indices for fat. However. May 6.40 Special Populations According to the findings of the Irish National Nutrition Survey. women had healthier diets in both countries. Children of the 1950s ate substantially more bread and vegetables and less sugar and soft drinks. Respondents in the UK reported eating more beans and pulses than French respondents but less cereal and fewer fruits and vegetables.41 Although a survey of Irish attitudes about diet placed nutrition/healthy eating in the top five most frequently selected factors affecting food choice. The median intake of dietary fiber was 26 g/d in the cohort of 22. In Sweden the estimated mean consumption of dietary fiber was 19. the story for children seems to be different in England.3 g/d based on the food-frequency . The intake of fruit and vegetables was particularly low for younger people and manual workers. The cohort with the highest intake consumed a median intake of 35 g/d and the quintile with the lowest intake consumed 16. fruits and vegetables. and Wales.6–25 g/d for boys and 17 g/d for girls. actual diets selected were not congruent with nutrition and health as being among the most important factors. Sweden Intakes in Sweden appeared to be higher than in the US or some studies in the UK but were less than those observed in Finland. the southern French diet was healthier. the diets of children 50 years ago had higher starch and fiber contents.42 Continental Europe France Using the statistical data on French food consumption published by the French National Institute of Economic Studies.2387_ch7. Scotland. the mean dietary fiber intakes were approximately 19.44 Some studies have compared fiber intake of those in Europe with those in the UK.45 Finland Finland appears to have the highest fiber intakes of any Western diet.000 Finnish smokers in the Alpha-Tocopherol Beta Carotene Study. making the fiber intake in this group low.47 While those in higher socioeconomic groups ate more fruits and vegetables. Intake of dietary fiber in this population correlated with antioxidant intake and inversely correlated with meat intake.5 g/d in summer. The Netherlands The average intake of dietary fiber in the Netherlands of 22 g/d was higher than in some other European countries.56 In a study with German insulin-dependent diabetics (EURODIAB IDDM Complications Study). studies of food offered in 20 nursing homes in the German state of Hessan had dietary fiber contents over 30 g/d.48 Thus. Individuals with low fiber intakes ate less fruit. bread.58 Austria Diets of 63 youth with insulin-dependent diabetes mellitus were compared with those of other Austrian youth and were found to be very similar.61 . diets of adolescents in Valencia offer an excess of proteins and saturated fat.54 A study of 627 healthy German children and adolescents between the ages of 1 and 18 years found the mean fiber intake was 1.57 Poland In Poland.60 Spain Fiber intakes in Spain were reported as highly variable according to region and other factors.5 g/d in winter and 8.000 East Germans aged 35–65 years. dietary fiber intakes ranged from 23.88/2. the average daily dietary fiber intake was 1.51 Germany According to the German National Health Interview and Examination Survey of 4030 participants.29 g/MJ). poultry. Those ingesting higher amounts of fiber also ate more fruits and vegetables. fiber intake was deemed suboptimal.53 In a survey of over 10. and other markers of a healthy diet. antioxidants. fiber intake was below the desirable level.17 vs.52 However. In 10-year-old boys.9 g/MJ. In teens. and cheese. with lower intakes among the less educated (1. These values exceeded that recommended by the German Association for Nutrition (Deutsche Gesellschaft für Ernährung). high sugar intake was associated with low dietary fiber intakes. while complex carbohydrates and dietary fiber are scarce. Studies of diets in three villages showed that dietary fiber and carbohydrate were 29–35% below recommended levels.03/2. fish. 3RD EDITION questionnaire for 92 randomly selected middle-aged Swedish men. In this cohort.1_fm Page 560 Sunday.8 g/MJ.59 A study of over 4000 adults in Warsaw showed that fiber intake met the RDA for men but not for women.7 g/d in the lowest quartile. 2.0 g/d in the highest quartile to 13. 2001 7:59 PM 560 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. For example. about 20% of the men and 27% of the women ate the recommended levels of dietary fiber. but the total carbohydrate was too low and fat and cholesterol were deemed too high.2387_ch7. According to the Dutch National Food Consumption Survey of 1987–1988 of 5595 subjects. higher fiber intakes were associated with supplement use. May 6. the method of determining the mean dietary fiber intake had only a small effect on the values. Dietary fiber intakes were near the recommendations. dietary fiber content of 12 standard hospital diets showed that fiber was derived mainly from grains and gave a mean dietary fiber intake of 11.49 There was a difference in fiber intake by education in the Rotterdam study (a cohort of over 5000 men and women over 55). 8%) and the majority of this fiber (71%) is insoluble. vegetables were the primary fiber source. For male Japanese medical students.3 g/d using Brazilian food tables. 2001 7:59 PM CONSUMPTION OF DIETARY FIBER 1992–2000 561 The differences in fiber intake between men and women were observed in the six countries studied in EURALIM. the dietary fiber intake was 19. Mexican-Americans born in Mexico consumed significantly less fat and significantly more fiber than those born in the US. South Africa Nearly 800 free-living Indian men and women (15–69 years of age) living in Durban were interviewed using a 24-hour dietary recall. with vegetarians consuming more than omnivores. Africa. as fruit and vegetable consumption is fairly limited.6 g/d using AOAC values and 17.4 g/d by the . 9.0 g per 4. Beans were the most important dietary fiber source in the population diet.2387_ch7. females consumed an average intake of 19 g/d and males 25 g/d. with an average consumption of insoluble fiber of 17 g/d and soluble fiber of 7 g/d.66 In general.65 The very high dietary fiber intake could be attributed to the intake of tortillas and beans and is not related to high intakes of fruits and vegetables. the average consumption was 20 g/d and among men 29 g/day. May 6. A study of 686 Japanese medical students showed the intake to be less that the average Japanese adult.62 Some Data from Latin America. 31% from vegetables. Dietary fiber density was significantly higher in women than in men.69 In New Zealand. In Japanese diets.0 and 11. Japan The average adult in Japan was reported to have a dietary fiber intake of 22–24 g/d. A study of 52 children (mean age of 6. Australia and New Zealand The average Australian consumes 22 g/d of dietary fiber.68 The average consumption of dietary fiber was 24 g/d. The children with normal bowel habits ate a median of 12.000 in 1991 to 1995) may be attributed to components of the grains and legumes. The diet is particularly high in insoluble fiber and phytate.8 years) with chronic constipation was ageand gender-matched with 52 children with normal intestinal habits. Among women.67 Brazil Dietary fiber intake of 559 adults (>20 years old) in Sao Paulo was assessed by the dietary history.7 g/d using AOAC values and 13.1_fm Page 561 Sunday.72 Furthermore. the typical Sonoran diet is high in dietary fiber (7. For example. the low colon cancer rates (20/100. this study showed that the method of analysis made a difference in the actual numbers reported. Seventh-Day Adventists in New Zealand consumed 27–40 g of dietary fiber per day.70 About 33% came from cereals.8 g/d using Brazilian food tables. Median dietary fiber intakes were low and varied between 8. Thus.71 according to 351 1-day weighed diet records. Constipated children consumed less. The fiber content of the diet was evaluated with a 24-hour dietary recall. and 20% from fruits. and the South Pacific Mexico Dietary fiber intakes in some parts of Mexico are extremely high for both children63 and adults64 and have been recorded as high as 94 g/d.2 kJ. Another reason for low fiber intakes is that Americans and Europeans responded that their diets are quite good and do not need changing. Countries such as Mexico and Brazil have higher fiber intakes.3 g/d and 16. Knowledge did affect dietary fiber intake in some people. On any survey day. From the five servings of fruits and vegetables. 3RD EDITION Southgate method and 15.000 respondents. especially coarse brown breads. the survey population failed to eat according to the FGP. Of the perceived barriers to eating a healthful diet. From the meat group. the fiber intake was 18. diets do not seem to be improving over time. showed that the consumption of fruits and vegetables remained constant over that time period.75 Some studies show that diets are actually getting worse.78 CONCLUSIONS Individuals living in Western countries have fiber intakes below recommended levels.2387_ch7.74 The proportion of the population consuming at least the desired number of servings from each of these food groups was 29% for the grain group. The new dietary fiber definition is just one . the National Health Interview Survey.1_fm Page 562 Sunday. or simply did not wish to change. many responded that they did not want to give up the foods they liked. fiber.76 Increasing the median fiber intake by an average of 9–12 g to the level at which it could be protective would require selection of at least three to four more servings of high-fiber products. knowledge. are good for them. May 6. ease of eating a healthful diet was most strongly and consistently predictive of intake. and 61% for the vegetable group.9 g/d by the Prosky method. If individuals followed the FGP. According to NHANES II data. preferred the low-fiber foods. fiber intakes from the 6–11 servings of the bread and cereals group could range from 6 to 30 g/d. many also prefer the taste and texture of their low-fiber counterparts. Between 1987 and 1992. Furthermore.9 g/d.73 Only 10% of the population ate at least five servings of fruits and vegetables. Among the Western countries. 5% of the population failed to consume any foods from the grain group. Another reason for a poor showing in terms of fiber intake is that many popular. for females.77 Furthermore. 0–6 g/d of dietary fiber could be obtained if just one serving of nuts or legumes was selected from the meat group. 29% for the fruit group. Only 3–6% of the US population consumed diets that fulfilled the USDA Pyramid recommendations on any survey day. REASONS FOR FAILURE TO MEET FIBER INTAKES One reason for the low fiber intakes in the US and in other countries is the fact that many fail to eat according to country recommendations or according to food guides such as the USDA Food Guide Pyramid (FGP). While surveys suggest that the average consumer believes that high-fiber foods. fat. Thus. 2001 7:59 PM 562 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. good-tasting foods contain < 2 g of dietary fiber per serving. and 18% of the population failed to consume any foods from the vegetable group. health professionals need to use every means possible to institute programs that can increase the fiber content of the diet. 46% of the population failed to consume any foods from the fruit group. Specifically. respectively. and fruit and vegetable intakes more closely approximated dietary recommendations for persons with more cancer-prevention knowledge. it is clear that the intake of fiber-containing foods is very beneficial in the prevention of disease. with 10. The strength of the associations between attitudes. While it is not clear that fiber itself is preventative of many diseases. Finland has the highest intakes and the US has the lowest. and dietary behavior varied in some cases according to level of education and perceived barriers to eating a healthful diet. 4–12 g/d of dietary fiber could come from fruit and 4–12 g/d of dietary fiber could come from vegetables. D... Med. S. 5. 545.. 7. 2000. E. and risk of NIDDM in men. Hunter. Diabetes Care. European Cancer Prevention Organization Study Group. May 6. Stampfer. F. Ascherio. W. E. Cooper. D. 8 Week Cholesterol Cure.. L. 46. W. 790. S. R. J. Schatzkin. Cancer Epidemiol.. 1994.. 1997. Shike. Rouse. J. Dietary fiber. N. Walker. Dietary fiber and disease. A.. Curley. and Sacks. Chicago. Harper & Row. B. 2000. 3. Shike.. Rimm. 23. 5. Levy.. Food group intake patterns and associated nutrient profiles of the US population. Polyp Prevention Trial Study Group. L. 1068. 2001 7:59 PM CONSUMPTION OF DIETARY FIBER 1992–2000 563 way to raise consumer and industry awareness so that steps may be taken to encourage the eating of more fiber-containing foods and to strengthen the demand for more fiber in food products. Lanza.. 1996... Kronborg. M. Rimm. 46. Slattery. E. D. Jones. 1996. Lanza. Ascherio. Fruit and vegetables in the service member’s diet: data from military institutional feeding studies. J. M. J.. 106. Med. Clemens. J. and Faivre. and Willett... J. 229... Hermann.. Ganji. Elder. 49. 22. Iber. 91.. JAMA. 1999. high-fiber diet on the recurrence of colorectal adenomas. 17. 169. B... Dietary fiber content of a six-day weighed military ration. 240. Dietary fiber and the risk of colorectal cancer and adenoma in women.. Colditz. F. C. 438.. The brave new world of FDA nutrition regulation — some thoughts about current trends and long-term effects. Mil. Giovannucci. M. D. 15. A. R. Engl... The effect of eating out on quality of diet in premenopausal women. 1953. 1987. Inst. 356.. O. Nutr. M.. E.. and Lee. Am. C. 322.. J. H. 147. Schatzkin. B. Med. discussion 50. L. Ziegler.. R. L. Implementation of dietary modifications. Ballard-Barbash.. 20. J... Block.. Colditz. Lance. Warber. A. 14.. 1987.. E. R. A. Restaurants and Institutions. Spiegelman. 1999. N. E. J. G... D. A. 99. G.. 2. J. and Cline. fiber and sodium intakes of US population: evaluation of diets reported in 1987-88 Nationwide Food Consumption Survey.. 21. and Kopel. 1532. Corle. F. Kikendall.. and Cahill. J. and Nestle. REFERENCES 1. N. 1990. Diet Assoc. 1999. and Stokes. 21... and Kessler.. B. M. Speizer. Salmerón.. H. Nutr. M. fruit. Rosner. Brit.. 20. 447. Kant.. P. A.. Engl. McNamara. Caan. Mil.. Hodgkin. Med. Lancet. Burkitt... Clin. Eur. 102.. R. 1991. Schatzkin. DeVries. Boquist.. C. and Daston... G. 2000.... Nutr. Weissfeld. J. 215. G. Y. 160. J. E.. M. Haddad. 11. Warber.. Rath.... C. A.. Slavin. and cereal fiber intake and risk of coronary heart disease among men. Am. J. Vegetable.. Clifford. Am. D. Hammond... Kowalski.. N. Lanza. J. C. L.. and Klesges.. V. 2. J. and Willett. C. . V. 1999. U. Paskett.. and Betts. J. 442. Dietary “fibre” and pregnancy toxemia. H.. J. 275.. 9.. 915. J. Corle. Fuchs. D. 13. D. W. Stampfer. Public Health.. J.. Am. Block. J. 1300. 398. 1149. W. Caan. Annual Meeting. Med.. A. J.. A. S. H. glycemic load. J. Freimuth. Spiegelman. A. 16.. A. Matters of choice. Stampfer.. S.2387_ch7. 8. 10.. Jenkins.. R. Giacosa.. G. Weissfeld. Burt. 19. R. L. D.. C. 1987. J. Mateski. Health advertising: prevention for profit. cholesterol. E. Calcium and fibre supplementation in prevention of colorectal adenoma recurrence: a randomised intervention trial. 1988. E. E. Hipsley. Effects of a health promotion advertising campaign on sales of readyto-eat cereals.. 6. D. Fat.. 4. M. Biomarkers Prev.. Food Tech. C. Comparison of the effects of oat bran and low-fiber wheat on serum lipoprotein levels and blood pressure. H. JAMA. E. Giovannucci.. 340. and Willett. Am. N. R. Wing. S. C. K. 22. 385. 34(2).1_fm Page 563 Sunday. C. J. J. and Painter. 342... E. J.. B. 14. C. 78. L. 468.... Food Sci. and Stein. Dietary Fiber Symposium. P. Lack of effect of a low-fat. Clin. Engl.. 1997. 18. 1995. 1995.. Diet Assoc. J. 1974. 12.. Rev. Crit. D. Nutr. I.. Public Health Rep. G... Med. A.. 162. A. M. F. B. Fiber intake of older adults: relationship to mineral intakes. M. R... Bonithon-Kopp. 73. J. Slawson. 1992.. A. Hanson.. Swain... B. S. 557. P. Med.. Bote. 11. S. J. E. Dietary fiber intake in the US population. Sheridan. Hayes. The polyp prevention trial II: dietary intervention program and participant baseline dietary characteristics. D. F.. K. 30. 988. Clin. J. Nutr. 1994. G. S. 1995.. H. Southgate.. and Dickerson. R. 1995. 11. 1986.. M. Public Health Nutr. J. Eur. Evaluation of lifestyle. 1995. and Riboli. P. Nutr.. W. Nelson. Y. sugar. J. 31. C. Fam. Hum. Luscombe. 219. L.. and Wynder. Slimani. Gunner. J.. 38. 1999. and Colditz.. G. I. M.. Public Health Nutr. G. W.. Rimm.. J.. G. Pediatrics. C... 25. A. 27. K. Saldanha. S. 1999. Betts. Sociodemographic determinants of perceived influences on food choice in a nationally representative sample of Irish adults. 32. moderately obese and severely obese subjects.. Beardsworth. G.. and Tucker. C. 9. Wallace.. and Anderson. H... C.. 541. and Starr. K... G.. and Beevers. Leverkus. E. Food and nutrient intake of a national sample of 4-year-old children in 1950: comparison with the 1990s. Am. A... Am... Day. Prynne. 77. 2720... and Heiss. C. 21. Crowder. Shultz. M. and Debry. Pediatrics 96. J. Frazier. Ann. 79. Scali. 53. Grace. Hampl. Pediatrics. J. 1996. 2. 96. E. E. Hurson.. Diabet Med. L. Korhonen. C.. M. and Berenson. Cole-Hamilton. A comparison of dietary behaviour in central England and a French Mediterranean region. Niederpruem. Ethnic differences in public health awareness. and Gustafson. Appl. 46. 225. Nutr.. M.. 47. 771. Circulation. A. Family dinner and diet quality among older children and adolescents.. 985. L. 3RD EDITION 24.. Cronin. M. M. A.. Haslam. 612. D. R. L. 96. B.. Br.. Nutr.. J. Metab. Jr. Dietary fat. Eur. 2000.. E. Clin. Coll. 29. 36. Charrondière. Am. 139. A. M. G. A. J.. JADA. Health. Nutr.. The ‘age+5’ rule: comparisons of dietary fiber intake among 4.. S. M. Malik. J. Health.1_fm Page 564 Sunday. food consumption and nutrient intake patterns among Irish teenagers.. and Heaton. Hilder. May 6.. Field. A.. 98.. 28. M.. A.. 1985.. P. Nicklas. 537... Humphreys.. Rockett. L.. C. L. Fiber intake of normal weight. Myers. and Corish. S. Albanes. 2001 7:59 PM 564 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and Wadsworth. A study among dietitians and adult members of their households of the practicalities and implications of following proposed dietary guidelines for the UK British Dietetic Association Community Nutrition Group Nutrition Guidelines Project. 26.. 39. C. 69. W. D. Sci. C. Dietary fiber intake of children: the Bogalusa Heart Study. Nicklas.. E. J.. Estimation of the daily dietary fiber intake in France. M. Paul. 39. 94. Avallone. W. Pomerleau.to 10-year-old children. Diet composition related to body fat in a multivariate study of 203 men. A. and Verger. Appl. Willett. Diet Assoc. J. 1993. G.. 2000. M.. C. McCarry. Beta-Carotene Cancer Prevention Study. 209. European Prospective Investigation into Cancer and Nutrition. B. Beerman.. Nutr. J. 1. J. 45. Rifas-Shiman. Ethn. 34.. E. 1998. . C.. C. D. 1997. 1418.. S. Bollella. Symes. and Virtamo. 47.. Nutr. C. Nutr. Davies. J. 33.. 20.. Are the nutritional recommendations for insulin-dependent diabetic patients being achieved?. A. 42.. J. 105. J. M.. 235. The AlphaTocopherol. L. L. Emmett. H. Intake of dietary fiber and risk of coronary heart disease in a cohort of Finnish men.. 1994.. A. P.. 95. Dunne. 2000. S. G. N. Alfieri.. J. Dietary intake and sources of nonstarch polysaccharide in English men and women. K. W. O’Neil.. L. 43. and Sherratt. and fiber predict body fat content. Sources of error and nutritional adequacy of the food pyramid. 40... A.2387_ch7. Holdsworth. A.. A. A new recommendation for dietary fiber in childhood. 94. 3. Clin. 1996. Kearney.. Eur. 96. Fiber in the diet of US children: results of national surveys.. T. Gillman. I. 1995. and Gibney. Obes. M. M. Barry. D. 166. M. M. 34. J. 3. M. K. 2000... Deharveng. Hartman.. E... M. 81.. Hum.. Res. C. J... 60. N.. B.. U. Pietinen. N... L.. 1990. G. 2. Lip. Myers. J. Am. A. 41.. C.. W.. 365. L. 57. 994.. Gerber. Recent national French food and nutrient intake data.. M. 530. Diet Assoc. 47. P.. A. Nutr.. 54. Miller. Pediatrics. Williams. Med. Camargo. Berkey. G. Bagheri. L.. 40. 37. 35. 1996. Impact of dietary fat and fiber intake on nutrient intake of adolescents. Diet Assoc. Med. Ir. T. health perceptions and physical exercise: implications for heart disease prevention. P.. Volatier. J. Tavelli. C. and Ferriss. and Benes. 1998. 44. and Lindeman. S. Comparison of nutrients in the food composition tables available in the nine European countries participating in EPIC. Arch. Kearney.... 1997. Dietary fiber intakes of individuals with different eating patterns. Price. Ribas.. P. 2000.. W... Giampaoli. Clin. J.. R. 1996. A. J. Buyken.. Dixon. Kulesza. 55.. EURODIAB IDDM Complications Study Group. P. 37. Wyatt. Dietary patterns in six European populations: results from EURALIM. 2000. Nutrient intake and growth of preschool children from different socioeconomic regions in the city of Oaxaca. F... 2000. 56. Carlsson.. A. sodium.. A. J. Food frequency questionnaire versus 7-day weighed dietary record information on dietary fiber and fat intake in middle-aged Swedish men... and food intakes in a US sample of Mexican-American women and men: findings from the Third National Health and Nutrition Examination Survey... and Roma. Farre. Kistemaker.. 58. Rywik. M. 1991. G. M.. P. J. 2000. 1998. J. K... P... Sucrose intake in Germany. and Navarro. 54. Coll. S. van Rossum. Löwik. Galan. Nutr. G. 51.. and Martins. S. Preziosi. S. S. F. 225. C. J. Cabrera-Pacheco. R. U. Am. and Fuller.. 1999. K.. O. P. Hig. Macronutrient intake of 1 to 18 year old German children and adolescents. Epidemiol. Vitek.. Sundquist.. S. L. L. and Triana Tejas. 65. Intake of dietary fiber... Nutr. 303. Vitelli.. A. Schober. Prättälä. 66. Yarnell. 46.. Hulshof. Grobbee. Ann. F. Gastroenterol. Nutr.. M.. R. Morabia. Z. S. 64.. Diabetologia. R. R. Modern and healthy? Socioeconomic differences in the quality of diet. 29.. Rev. Publica. G. Med. B. E.. M.. Nutr. Dietary fiber consumption in an adult population. Arch. B.. J. J. G. Bernstein. Mensink. J.. P.. E. B. P. 40... C. Stehle. J.1_fm Page 565 Sunday. Saude Publica.. 1993. G. Schneijder. McCrum.. and Ockhuizen. Roos...... U. . Eur. Gesundheitswesen. Dietary assessment of adults in three villages in Warsaw. L. and Schoch. Kleemola. 163. Nutr... T. S. 54. S. E. E. L. 161... Van Poppel. Ostenson. Dietary habits of the middleaged Warsaw population in 1984 relative to nutritional guidelines. 1998. Valencia. A.. Sygnowska. R. Nutr. Nutritional intake of daily diets in nursing homes for the aged.. and Winkleby. Ernahrungswiss. and calcium and its relation with arterial blood pressure in normotensive adult men... 48. M. E. 35. 50.. Perers. I. G. Kroke.. Hercberg. and Pietinen. 62. and Kok.. C. C.. J. J.. protein. 2001 7:59 PM CONSUMPTION OF DIETARY FIBER 1992–2000 565 47. 16. Differences in energy. and Muskat. van de Mheen. 2000. and Wallen. Eur.. J.. 241. 26. M. Kersting. Z. Wyatt. a collaborative European data harmonization and information campaign.. Scand. The usual diet of a group of adolescents from Valencia. 57. 50. Klipstein-Grobusch. The Rotterdam Study. A. C. and Waskiewicz.. N. 37. Persson.. G. Frasquet. H. Brzozowska. C. ten Hoor.. 61. Gareis. M. Clin. Heitkamp. J. and Puzio-Dbska. 37. Sichert-Hellert.. J. Nutr. and Haas. Beer-Borst. B. J. J. nutrient. 75.. Manz. 60. Zakl. M. M. Nutr. Pediatr.. F. H. 41. Eur. Taschan. Relation of fiber intake to HbA1c and the prevalence of severe ketoacidosis and severe hypoglycaemia.. M. Thamm... Scherbaum. J. L. Arch. Wyatt. K... E.. and Mackenbach. H. I. C. 1998.. 14. Appetite. Panico. Z. Ernahrungswiss. Ballesteros-Vasquez. 223. Ernahrungswiss. E. M. A. 49. C. P. 66. Nutr. Comparison of dietary intake data with guidelines: some potential pitfalls (Dutch nutrition surveillance system). F. Evaluation of the composition of the regional diet in Sonora. M. Galasso.. Education and nutrient intake in Dutch elderly people. Hermus. 1999.. Serra-Majem. 882. Rocz. Ernahrungswiss... V. Toeller.. Mexico. México: incidence of colon cancer. Vescio. Nutr. M. R. B.. 52. G. Voss. Pardo. 53.. W. Martinez. J. 1998. Schrijver. 68. 1999. A. I.. R.. 548. Metab. Salud.. R. and Boeing. Pietruszka. 1998. 12. 38.. 159. Mex. 152.. 1998. M. 48.. Saucedo-Tamayo. H. Löwik. E.... S.. May 6. A. M. T. 176... 1988-1994. K. E. Mattos. Grill. Lundgren. F. C. S.. 67. P.. Am. Hagman. 253. Soc. Colon cancer in rats and diet in the Sonoran desert region of Mexico.. J. Dietary intake of Austrian diabetic children 10 to 14 years of age. and Rami. Lahelma. S.. 49.. 50. Nutrition in Germany 1998. Linseisen. E. 34. Rupprecht. carbohydrate and fiber. M. Winter. 1991. Latinoam. Alexy. Karg. 200. Influence of lifestyle on the use of supplements in the Brandenburg nutrition and cancer study... 144. Langergraber. 1996. B. 753. Dorado... 61. G. A. Radom and Biaa Podlaska districts. Witteman. 63. and Wolfram. Nutritional status and food consumption in 10-11 year old Dutch boys (Dutch Nutrition Surveillance System). 54. I. and Northridge.. A. M. 44. W.... Panstw. Z... S. I — Energy... Hosp. 33. 11. 1998.. fat. 1998. potassium. 1996.2387_ch7. 14.. 59. and Grijalva-Haro. 219. Stelz.. E. 252.. Latinoam. I. Clin. Gedrich. Br.. A. Clin. and dietary fiber in medical students. Breslow. 2000. Assoc. K. American Heart Association. C. J. H. Block. A. and Block. M. and Brand. Am. Lane..org/nfs/nfs88.. C. M.. 78. Hasegawa... 71. 2000. 48. J. Nutr. Dietary intake and iron status of Australian vegetarian women. L.. NHANES II. 97. Subar. T.. 519.. 1997. 1996. Ziegler. Ikeda. 72. and Jenson. 1997. Clin. 79. G.. 353. L. 86... A. United States Department of Agriculture.. Canberra. 85a..org/diabetescare/supplement/s16. S. Sugawara.... 1996. 48. and Cheung. J. Dietary guidelines and the results of food consumption surveys.. Am. E. and Baghurst. 1999. Nutrition. C. Dietary Guidelines for Australians. . Marlett. J. Dietary diversity in the US population. Am. S. 18. lipids. J. United States Department of Health and Human Services.. 75. http://www.2387_ch7. 2. and Nestle. G.. http://www. 133. Diet. A. Kant. Dietary Guidelines for Older Australians. J. Canberra. Clin. A. A rapid food screener to assess fat and fruit and vegetable intake.. 1976-1980. Am. 1997. A. 356S. 1139. 76. 888. 632.. National Health and Medical Research Council. M. National Health and Medical Research Council. T. Australian Government Publishing Service. Subar. G. Diet. USDA. 1991. Clifford. 82. R. 2001 7:59 PM 566 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Kearney.. J. and attitudes to cancer prevention dietary behavior. and Lanza. Nutr. 95. Kitagawa. Harnack. F. Block. Trends in food intake: the 1987 and 1992 National Health Interview Surveys. I. 70. and Cancer Prevention Guidelines.. NCI dietary guidelines: rationale... energy. 83. N. 70. P. 957. Diet. Br.. 43.. K. American Cancer Society. 28..html 80. Diet. and McElhone.diabetes. Nutr. 1988. suppl. F. May 6. G. 1997. minerals and dietary fiber in middleaged Japanese. Block.. Nutr. Schatzkin. S. Association of cancer prevention-related nutrition knowledge. Med. B.. 84. G.1_fm Page 566 Sunday. Aoki.. Am. 1999. Australian Government Publishing Service. J. R. Rosenbaum. M. Circulation... and Bartlett.. J... 2701.. 53. 1999. Ball. Cancer. Butrum. M.. Imaeda. Eat Well for Life. 97. Sci. Baghurst. (Tokyo). Assoc. 85. Am. Assoc. Vitaminol. A.. 77. beliefs. Fujiwara.htm 81.... 45. A. Nutr. Endo. S. K. N. 284. Prev. K. J. Patterson. G.. 74. Block. Nakaji. Dietary patterns and intake of nutrients. R. H. Gillespie. Dietary Guidelines for Americans. H.. Fiber. a statement for healthcare professionals from the Nutrition Committee. and coronary heart disease. 1999. 1994. J. Nutr. S. Am. E.. I. Perceived barriers in trying to eat healthier — results of a pan-EU consumer attitudinal survey. Nippon Koshu Eisei Zasshi. 91. 1991. 81. 86. Tokudome. 1526. Y... Dietary fibre. Database and quick methods of assessing typical dietary fiber intakes using data from 228 commonly consumed foods. and Tokudome. R. M. J. 3RD EDITION 69.. Van Horn. Foods contributing to absolute intake and variance in intake of selected vitamins. R. Supplement Food Austral. 73. and Totsuka.eatright. 50 © 2001 by CRC Press LLC 567 . and the majority of information about fiber consumption worldwide is based on these analyses. they are doubtless more accurate and therefore are discussed first in this chapter.7. which can introduce artifactual increases in “fiber” seen in some other methods. the majority of expert committees have recommended an increase in the fiber content of Western diets because there is accumulating evidence that fiber is important in the prevention of a large number of bowel disorders. METHODS Analytical There are various methods in existence for the measurement of “fiber” in foods. using tables of food composition.00+$1. storage.2387_ch7. are also lower than formerly.33. in fact little is known of the patterns of fiber consumption worldwide. showing a reduced level using these figures compared with former estimates. 2001 8:00 PM CHAPTER 7. UK food tables published in 19784 included values for fiber using mainly the Southgate method. and preservation of food. incorporation of “fiber” values into food tables is a difficult problem and most government-sponsored food tables do not include values for it. estimates based on the table of Lanza and Butrum.8 which incorporate data using four different methods. Although the data on these new estimates are limited.1–3 However. 0-8493-2387-8/01/$0. Some information on NSP consumption is beginning to emerge.2_fm Page 567 Sunday.36 These do not suffer from analytical problems encountered from the cooking. Hence. Crude fiber analyses are not relevant to human population studies. especially cooked foods. all of which give different values for particular foods.2 Patterns of Dietary Fiber Consumption in Humans to 1992 Sheila Bingham INTRODUCTION In recent years. recent supplements and editions now incorporate the analysis of nonstarch polysaccharides (NSP) from the analyses of Englyst. May 6. although it is assumed that fiber intakes are greater in areas where these disorders are rare. However. This is largely due to methodological problems in both the collection of food consumption data from representative population samples and in converting that data into grams of fiber eaten per day. In the US. The use of these markers in nutritional epidemiology is discussed elsewhere.2 or less. is used. published estimates of dietary fiber consumption are available for some countries using these data and are reported below. It should be borne in mind. but these statistics are not available for earlier years.1. All of these except one. Definitive information on food consumption is obtained from studies of individuals where the amount of food eaten is determined directly. calculated from food balance sheets. that based on the US NHANES II study. probably by different amounts in different populations.2_fm Page 568 Sunday.20 and 24-h recalls in the US NHANES II7 and Canadian students study.17 interviews in South African urban blacks. see Table 7. and these included the AOAC method. May 6. from 11 g in the US and Japan to 18 g in men in rural Finland. which is 1. and they give no indication of the amounts of fiber consumed by different age. that the estimation of the food intake of free-living individuals is no easy undertaking since all methods rely on information supplied by the subjects themselves. with no apparent loss of body weight is unphysiological. The Japanese diet seems to differ from that of the West with its low content of pentose (xylose and arabinose) polysaccharides . however. such as 24-h urine N from validated urine collections.10 and the data from individuals can be aggregated into group estimates. In the NHANES II study.14 the UK National Household Food Survey.2. Overall. A figure for Finnish NSP intakes. They are also only valid indicators of trends in countries with stable and well-documented population bases. or an independent index. and average intakes estimated on the same population using one method may be up to 30% greater or less than those obtained using a different method.8 The food consumption values in the table also incorporate different methods of obtaining food consumption data.13 In the remaining studies — in Scandinavia. and occupational groups within the population. neutral detergent fiber. estimates of energy consumption from national statistics are some 25% greater than the estimated energy requirement. there is only a small range in average intakes worldwide. and Englyst values. There are many sources of error. unless actual food intakes can be observed by investigators in certain limited situations. and supplies from vegetables usually predominate over cereals. using the publication of Heller and Hackler for 1973 to 1975 to NSP. is also shown. Insoluble fiber is consumed in rather greater quantities than soluble.9 Nevertheless.6 times the BMR in sedentary populations. ranging from 7-day weighed records in UK men and women. 2001 8:00 PM 568 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. which have been reported in some populations. sex. which may not be correct. Urine N should be 80 ± 5% of calculated dietary N intake in populations consuming amounts of fiber commonly found in Western-type diets.19 Recalculated data21 from the US food balance sheets.12 NEW ESTIMATES OF NONSTARCH POLYSACCHARIDE (NSP) CONSUMPTION Estimates of NSP and fiber using newer values are currently available for seven countries. There are many different ways in which this is done.22 and Japan18 — duplicate diets were analyzed directly for NSP.16 16-day weighed records in UK women volunteers. National per capita statistics also overestimate food consumption. 3RD EDITION Food Consumption In some countries.7 several different methods were used to compile a table of fiber values for foods. per capita data on food consumption from national statistics and household surveys are available for the re-analysis of long-term trends in dietary fiber consumption over this century. for example.4 to 1. In Britain.15 Loma Linda University volunteers.11.12 Energy intake to BMR ratios of 1.2387_ch7. used the Englyst values for the NSP content of foods. are also shown.10 The correct result is usually open to debate.11 A second validity check for populations is to calculate basal metabolic rate (BMR) from body weight using published equations and to compare energy intake from the dietary survey with estimated energy expenditure. and hence the validity of this dietary data is questionable. 4 — 5.0 14.0 4.2 2.6 — — — — — — — 5.0 3.2 5.4 3.3 5.6 3.1 5.5 — — — — From Vegetables .6 — 3.9 Uronic Acids } — — — 5.1 — — — — — — — 3.8 4.5 13. 2001 8:00 PM PATTERNS OF DIETARY FIBER CONSUMPTION IN HUMANS TO 1992 569 .4 6. 2387_ch7.3 — 2.1 7.5 4. — — — — 13 9 17 12 5 5 — 14 13 South Africa — urban black M urban black F — — 17 12 US — National Statistics 1973–75 US — NHANES II Men Women US — volunteers Loma Linda Univ.4 UK— National Food Survey UK — Men UK — Women UK — Women volunteers — 7.8 — — — — Soluble 8 6 — — — — 7 6.8 — — — — From Cereals 22 22 7 21 20 19 18 17 16 15 13 14 14 14 14 Ref.2.7 3.2 Cellulose — — — — — — — 2.5 Pentoses — — — — — — — 3.8 4.1 — 11.8 6.7 3.7 — 4.5 15.2 — — — — Insoluble Nonstarch Polysaccharide and Fiber Intake (grams per day) Country Table 7.5 4.6 Hexoses — — — — — — — 1.4 6.8 Japan — 12.4 4.5 — 7.7 2.a = daily range.0 18.2_fm Page 569 Sunday.1 5.8 — 4.2 9.2 3.5 3.9 Canada — students 3.4 5.0 1.1 3. May 6.8 16.6 5.1 1.2 Finland Rural Finland Rural Denmark Helsinki Copenhagen SD Total — — — — — 1–81a — 5–19 7–25 1–68a — — — — — — Range 9 6 1–48a — — — 5 4.9 2.2 12.8 6.2 — 3.4 18.6 — — — — — — 6.8 — 1.9 2.2 2. Men Women 10.4 — 9.0 — 1.1 11. 2. 333. However.1 Distribution of individual nonstarch polysaccharide intakes in 63 men and women.3 summarizes results from a number of surveys of between 30 and 2200 individuals that have been carried out in Europe. Comparisons are possible because the samples were all randomly selected from population or electoral registers and response rates were high. mainly due to the low intake of 14 g found in Iceland.2. 3RD EDITION and higher intake of uronic acids. In the small UK study.2. with the exception of the small study in the UK. With permission. Nevertheless. and cereals contribute more than vegetables. with an equal contribution from vegetables and cereals. J.2. is still greater than the range in average intake worldwide. 2001 3:09 PM 570 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. May 8. individual intakes were averaged over 7 days. starch consumption in this area is high and would have led to an overestimate in previous analyses. see Figure 7.. (From Bingham et al. show that fiber intakes are greatest in the Mediterranean countries. data on 23 countries from Bright-See and McKeown-Eyssen23 have been collated into 4 regions of Europe. There is a very large individual range in NSP intake. Nutr. However.1. usually around 80%. These data. South African blacks consume remarkably little NSP compared with previous estimates using Southgate’s analyses. while other estimates are used in UK volunteers and in the US NHANES study. In Scandinavia. based on food analyses of Southgate. this is probably due to the fact that single-day estimates are reported in these studies.2.) DIETARY FIBER (SOUTHGATE ANALYSIS) CONSUMPTION Europe National Statistics In Table 7. for example. from 7 to 25 g/d. Dietet. intakes are lowest. Hum. Individual Surveys Table 7. Figure 7.2_fm Page 570 Tuesday. In North and Central Europe. food consumption was assessed by asking subjects to keep records . 1990. vegetables are the major source. Men generally seem to consume more NSP than women. up to 81 g/d in Canadian students. East European intakes follow a similar pattern. In all these surveys.. the individual range in NSP intake.2387_ch7. 3. when the apparent range was reduced. Czechoslovakia. compared with an average of 45% in the EEC. 1966.2 Area a Scandinavia North/Centralb Mediterraneanc Eastd a b c d 571 (5) (8) (5) (5) Total From Cereal From Vegetable From Fruit 21 25 38 30 10 9 16 14 7 11 16 12 4 5 6 4 Denmark. and approximately 30 families were studied per month over a period of 3 years. but in one survey in Scotland. Spain. but it is estimated from records of food purchases kept by the housewife for 1 week. a largely urban area. whereas intakes in urban areas were also around 20 g/d. as with NSP. Ireland. Most subjects were asked to keep a record for 1 week. whereby food entering a household over a certain period. from 8 to 32 g/d in the Welsh and British surveys. recorded an intake as high as 25 g in men.24 However.2_fm Page 571 Sunday.3. Food entering the household is not weighed. and these are summarized in Table 7. Household Surveys A commonly used method of assessing the dietary intake of populations is the household survey. which has been carried out every year in Britain since 1940. Yugoslavia. This does not give information about the distribution of intakes between different individuals. Thus. data on the distribution of dietary fiber intakes of individuals within the overall average are published. Greece. Italy. Belgium. even though dietary fiber intakes in Europe are comparatively low.39 Intakes in rural Finland and Denmark and in Holland and Yugoslavia were rather greater.38 Fiber intakes in the UK were measured at around the 20-g mark.2387_ch7. Norway. and the amounts of food eaten were in most cases weighed by the subjects themselves. Poland. Hungary. estimated from this survey in a comparative year. using values for British foods together with data from another household survey. K. Romania. Iceland.2. Table 7. thus eliminating the need to make corrections for inedible wastage. Total dietary fiber intakes in Britain. the National Food Survey. each individual was studied for 1 month. . 2001 8:00 PM PATTERNS OF DIETARY FIBER CONSUMPTION IN HUMANS TO 1992 Per Capita Dietary Fiber Supply in Europe 1972–1974 (grams per day)23 Table 7. The largest survey. Individual Variation In some of these surveys in Europe. Austria.2. is estimated and the total food available is divided by the number of people living in the household.40 In this survey. Switzerland. Coefficients of variation (standard deviation/mean %) are on average 35% and the range between individuals is greater than that found between populations. Portugal. May 6. of over 2000 individuals. France. for example. with lowest intakes recorded in Scotland. individuals appear to vary markedly within a population. usually 1 week. U. each family weighed all their food after preparing it. Finland. Sweden.2. were 21 g/d. modified values for the fiber content of bread were used in these surveys. The majority of the results were calculated using Southgate’s method of analysis in the British food tables. In this. although the staple food in all of the areas was white bread. Bulgaria. randomly selected from each of the nine standard regions in England. about 6500 households are studied per year. All the areas studied were rural.4 summarizes fiber intakes calculated from the published values on food consumption. and Scotland. Germany. Netherlands. This is less than the rural areas of Europe with only 32% derived from cereals. of food actually consumed. but additional information about dietary fiber consumption in European populations is provided from a survey carried out in the EEC from 1963 to 1965 to assess levels of radioactive contamination. although this was supplemented in Denmark and Finland with Englyst’s values for rye flour26 and in the Netherlands with data from the method of Hellendoorn37 and Karen. Wales. Sample Randomly selected from electoral registers.8 20.0 2. 572 70 Cooperation Rate (%) TABLE 7. 2001 8:00 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.6 — — Fruit DF (g/d) — — 32 34 27 26 36 36 TDF (CV %) — — 6–36 8–32 10–32 18–32 10–45 7–33 Range (g) 28 27 26 25 24 24 Ref.1 25 19 Total DF (g/d) — — 7.5 19.7 6.3 Dietary Fiber Consumption in Europe: Individual Surveys 2387_ch7.6 9.2. 16–64 years Men Women Randomly selected from electoral registers.9 19.0 17. May 6. 28-d estimated record 7-d weighed record 7-d weighed record 7-d weighed record 7-d weighed record Survey Method S S S S SM Analytical Method 16.8 7.5 15.6 — — Cereal DF (d/f) — — — — 8.0 5.6 — — Vegetable DF (g/d) — — — — 2. 20–80 years Men Women Randomly selected men from prevalence study of heart disease Randomly selected men aged 40 years Randomly selected men and women aged 21–69 years Men Women Country Great Britain England Wales Scotland Scotland 84 78 89 82 58 16 27 107 119 32 31 1087 1110 No.2_fm Page 572 Sunday. 3RD EDITION . 7 1. men aged 50–59 years Rural Copenhagen Randomly selected men. Randomly selected from electoral registers.5 23. men aged 50–89 years Rural Helsinki Randomly selected from population registers. Zutphen Randomly selected from population registers. Ireland — 63 75 83 74 — 67 74 49 30 30 30 29 44 56 871 258 334 7-d weighed record 4-d weighed record 4-d weighed record 7-d estimated record Diet history 7-d weighed record S S E S E K S H ? S 25.5 4.7 4. E36. 35 32.7 13.5 21. H37.8 — — 3.Randomly selected from electoral registers.5 12.3 30. May 6.5 27.8 — 7.7 1. K38.6 6. 33 32.6 21.4 2. 7 countries heart study Netherlands Finland Denmark Yugoslavia Note: S4.6 1.1 — 9.6 — 3. 33 32 30 29 2387_ch7.2_fm Page 573 Sunday.3 6.0 26.6 — — 29 37 36 42 42 28 22 32 37 32 9–46 10–55 2–30 5–45 10–40 — — — — 34. SM39.7 — — 13. 25–65 years Men Women Men. 16–64 years Men Women N.4 21 23 8. 2001 8:00 PM PATTERNS OF DIETARY FIBER CONSUMPTION IN HUMANS TO 1992 573 .6 19.5 10.4 18. 42 In all three European countries. greater than the variations between areas.1 7.0 3. Dietary surveys to quantitate intakes of other nutrients have been done.8 29.4 Sources of Dietary Fiber in 11 Regions of the European Economic Communitya Areab Cereal Vegetable Fruit. in fact.1 10. H.2 7.1 2.6 28.1 6.2 1.6 24.7 — 22.3 24.5 13. These are summarized in Table 7. Germany.5.2 8.6 1.34 although there are no national statistics available earlier than 1909.3 4.8 2.0 6.6. Seasonal variations in dietary fiber estimated from these two surveys have been reported elsewhere.3 6. such as Britain. No dietary surveys have been done specifically to measure fiber intake and nothing is therefore known of its main sources and composition. Nuts Potato Total Friesland (H) Gent (B) Liege (B) Luxembourg Hessen (G) Normandy (F) Bretagne (F) Vendee (F) Fruili (I) Campania (I) Basilicata (I) Britain (1966) 8. to be taken into account when comparing dietary fiber intake of individuals or populations. I.50 Previous data44 showing intakes of 150 g in Uganda and 130 g in the Kikuyu resulted from food analyses of 6% in plantain and 10% in maize flour.7 5.1 a b c Values are given as grams per person per day.5 6.8 23.9 12.30 some long-term trends in dietary fiber consumption over the 20th century have been calculated from national statistics.9 8. G.2 10.1 27. cereal dietary fiber intake has probably declined.7 13. May 6.6 9. F.7 5.2_fm Page 574 Sunday. Cereal fiber intakes may have been greater in 1860. although in largely urban countries. and the most striking change in Britain was the doubling of fiber intakes during the period 1940 to 1953. The decline in cereal fiber consumption has been offset somewhat by increases in vegetable sources of dietary fiber.9 4.4 13. together with calculations of the dietary fiber content of diets consumed by farming households in Denmark. thus more than doubling the fiber content of bread and cereal fiber intakes during and after World War II. Includes potato. At the same time.7 5.2 10. France. suggest intakes of 63 g in urban areas and 69 g in rural ones. These are summarized in Table 7. Africa While much has been written about dietary fiber intakes in Africa. Studies on dietary fiber intake.8 1.3 12.8 21.3 3. and these have been used as a basis for obtaining some preliminary figures for fiber intakes. in some cases. Belgium.34 Long-Term Trends In Britain42 and the Netherlands.7 2.2387_ch7.0 7.9 2.2. therefore.6 0.2.8 1. so that more bread was eaten. B. quarterly seasonal variations are not as marked and are related to consumption of different types of vegetables. bread and vegetable consumption was encouraged while other foods such as meat and cheese were in short supply. This was the result of the raising of flour extraction rates. Italy.6 4.8 6. Holland.7 4. 3RD EDITION Table 7. using 7-day weighed records and values for the NDF content of foods in 300 young Nigerian women.9 26. 2001 8:00 PM 574 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.3 11.6 2.3c 2.3 10.9 26.4 These high values were almost certainly .9 23.1 14.8 29. seasonal changes from month to month were substantial and.3 2.5 4. very little is known about it.7 10. although by only 2 to 3 g since 1909 in Britain.2.9 5.34 In the rural areas. Possible seasonal variations need. Southgate et al. 48 } 49 50 Not available. 4. starch (including resistant starch) intakes are high and may account for the absence of Western bowel diseases.6). 1978. J.1). although there is a trend for increased consumption of brown bread... 324.23 . 45 46 47. Despite these low fiber intakes. Intakes are still comparatively high. 1982. 2001 8:00 PM PATTERNS OF DIETARY FIBER CONSUMPTION IN HUMANS TO 1992 Table 7. Assoc.2. however. nuts 86 32 54 —a 0 25 55 60 70 63 69 0 25 40 40 0 23 10 0 — 10 19 67 40 59 0 — 4 1 3 — — 23 42 22 32 18 36 10 22 — — — — — — — — Ref. 274. Data from another African country. 1982.7. were obtained from food balance sheets by Bright-See and McKeown-Eyssen23 and are shown in Table 7.2. In Soweto.5. 40 to 50 g daily in rural blacks. Mauritius. Dietet. Nature (London).6 Dietary Fiber Intakes in Africa and India (grams per head per day) Africa Kenya Kikuyu Masai Warriors Women Malawi (foothill village) Swaziland (middle veld) Uganda (Buganda) Nigeria (urban) (rural) India Andhra Karnataka Kerala Tamil Nadu a Total Cereal Vegetable Fruit.2387_ch7. bowel disease is rare. about 10 g/d. and that the current intake from unpublished surveys is 25 to 35 g/d.. such as Japan. much lower so that total intakes are apparently some of the lowest in the world. 80. Helms et al. with only Iceland having a lower value at 14 g/d. compared with present day estimates of 14 g in 76 urban Sowetan black males and 13 g in 113 females. The staple cereal in Mauritius is rice. Table 7..2.6 The data have therefore been recalculated and result in intakes of 70 g and 90 g. Intakes of vegetables and fruit dietary fiber are. May 6. 2% and 4%.2_fm Page 575 Sunday. respectively. contaminated with starch.2. Nutr.2. and consumption of vegetables is low due to their high cost. However. and recently published values are much lower. Legume consumption has decreased. in these areas (Table 7. obtained by Segal and Walker20 (Table 7. Cancer. Am.5 Long-Term Trends in Dietary Fiber Consumption in Europe (grams per head per day) Britain42 Total Cereal Year 1909–1913 1927 1933 1938 1944 1950–1951 1957–1960 1970–1976 30 42 43 575 24 — — 22 32–40 — 23 23 Netherlands30 Total Cereal 11 — — 9 19–25 — 9 8 — — — — — 27 24 25 Denmark43 Total Cereal — — — — — 14 10 8 — 34 32 — 37 31 25 23 — — — — — — — — Van Staveren et al. 51. 23. refined maize meal is consumed. respectively. They state that fiber intake in the past was higher. an urban area. so that country’s intakes of cereal dietary fiber are similar to those of other rice-eating areas. however. 8 g/d of dietary fiber came from cereals. Total Cereal Vegetable Fruit Ref.6). May 6. Australasia A number of reports of the dietary fiber intake in New Zealand. and Australia are available and are summarized in Table 7. Singapore (rice) 28 10 14 4 25 12 10 3 33 19 9 5 North America United States Canada 23 22 6 6 11 11 6 5 Other Israel Mauritius 36 17 16 10 12 6 8 1 Area America Chile. 53 51. where additional cereals such as millet are eaten. 65% of the dietary fiber coming from taro. In Kerala.2. Tokelau (Table 7.2_fm Page 576 Sunday. Uruguay (wheat and rice) Costa Rica. Mexico (maize) India Dietary fiber intakes have been calculated using data from a number of dietary surveys carried out by the National Nutritional Monitoring Bureau in 1980 in India.49 These show a twofold range in fiber intake from 22 g/d in predominantly rice-eating states such as Kerala to 42 g/d in Karnataka.9 g/d in Karnataka (Table 7.2.7 Per Capita Dietary Fiber Supply 1972–1974 (grams per day) Total From Cereal From Vegetables From Fruit Australasia Australia and New Zealand (wheat) 24 9 10 5 Japan. Hong Kong. 9. 52 51. 52. All suggest that dietary fiber intakes are low and similar to those of Europe.2. but surprisingly low on another Polynesian island. Cuba. 52 54 54 55 55 . 18 ± 5 16 ± 6 19 ± 6 6 6 5 8 8 10 4 3 4 51.2.8).7. 52 72 ± 29 15 16 1 — — 48 — — 22 — — 20–22 19–20 — — — — — — 51.2. and breadfruit.2387_ch7.8 Dietary Fiber Intakes in Australasia (grams per head per day) New Zealand Europeansa Maoria New Zealand Tongansa Polynesia Tongaa Tokelaua Tokelaub Australia Adelaidea Adelaideb a b Women. Polynesia. Men. Intakes are three times higher in Tongan women. 2001 8:00 PM 576 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. cassava. compared with 35. Table 7. 3RD EDITION Table 7. Trinidad. around 20 g/d in New Zealand women and in Australia. 0 5.0 54.9 Secular Changes in Major Nutrient Intakes.57 In the ten largest cities in 1979. particularly since these patterns of disease change within one or two generations in Japanese who migrate.56 Traditional Japanese diets are known to be low in fat and high in carbohydrate.7b 7.2 19.4 — — — 7. together with pickles. salted fish. Carbohydrate and dietary fiber intakes were also lower. 18.6 50. The Japanese diet was thus reported to contain only 19.7 36.7 g/d in these areas.53 Total dietary fiber from rice has fallen from 7. 32 g/d. 296 and 17.1 g/d in 1979.0 35.6 g/d.9).1 Protein from Animal Food (g) 25. However.2 g/d. less protein from animal sources.5 g/d of fat were eaten. These estimates using Southgate values for fiber are less than those for NSPs. reported the dietary fiber content of two student meals to be 14. and this data has been used to calculate dietary fiber intakes using the British Food Tables together with neutral detergent fiber plus pectin values for some seaweed-based foods. 57. which are 11 g/d (Table 7. and a 25% increase in the amount of protein derived from animal products since 1950. Only 12 g/d of fat was eaten.3 g/d.4 g/d in 1979. (per capita per day) in Japan Year Fat (g) Carbohydrate (g) 1950 1955 1960 1965 1970 1975 1979 18 20. yearly nationwide household surveys are carried out.8 418 411 399 384 368 337 315 a b Dietary Fiber Total From Rice (g) (g) —a — — 21.2b 20. There are no household statistics available prior to 1950. 50. and soy sauce.2_fm Page 577 Sunday. a threefold increase in fat intake. 1966.2.7.58 Estimates from food balance sheet data are rather greater.7 g/d in 1965 to 5. on which basis dietary fiber intakes might be expected to be high. Long-Term Trends In Japan. rice was eaten three times a day.2 g/d in 1965 to 19. since rice contains so little dietary fiber.2.1).2. 2001 8:00 PM PATTERNS OF DIETARY FIBER CONSUMPTION IN HUMANS TO 1992 577 Japan With its highly urbanized society yet a disease pattern which contrasts with Western countries.5 it might be predicted that the fiber intake of populations where rice is the staple cereal would be low. These changes are small compared with a 70 g/d decline in carbohydrate intake since 1965. spring onions.2 20.0 32.5 g/d. Table 7.3 24.2. although at that time rice supplied 11. radishes. 1950–1979.3 Data not available.4 g/d in 1979 (Table 7. but the dietary fiber content was the same as today.1 48. May 6.8 and 19. and 52% of protein was derived from animal foods.7 5. soybean curd.57 Mori. . Japan is of special interest to the epidemiologist.5 52. but a dietary record of an artist’s family (all adults) in 1925 showed that on a typical day.4 40. and overall total dietary fiber intake has fallen from 21.0 46. using his own technique for analyses of Japanese foods. This compares with less fat.0 44.52.57 Present-Day Intakes Geographic comparisons within Japan show that these changes in diet have been most marked in the urban areas.23 The estimates of Far East countries are shown in Table 7.2387_ch7. 2. 20. Previous studies suggested an intake of about 20 g/d from individual surveys (Table 7.1) are some 25% lower for food balance sheet estimates. 332 g. In wheat and rice areas. 11 g on average compared with 13 g using Southgate analyses alone.61 Table 7.7.2. Uruguayan intakes were 20 g/d.21. and range (from 24-h recall data) from 1 to 46 g/d. although Bright-See and McKeown-Eyssen23 calculated fiber intakes from food balance sheets which are summarized in Table 7.8 Recalculations of data on long-term trends in food intake showed a 33% decrease in total dietary fiber and a 55% decrease in cereal dietary fiber in the US from 1909 to 1975.23 The Middle East Data from food balance sheets and individual surveys are available for Israel. more carbohydrate. 2001 8:00 PM 578 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.2%.2_fm Page 578 Sunday. depending on the size of the city.751 American female nurses ranged from 14 to 17 g/d. of which 7 g is derived from vegetables and 6 g from cereals. Men Women Simulated diet NHANES II Menb Womenb Female nurses Third quintile American-Japanese men Canada Menb Women a Total Cereal Vegetable Fruit Ref.2. in towns and villages. 3RD EDITION 48. Calculated intakes were higher in the two maize-eating areas.7 Intakes assessed by a questionnaire of the third quintile of 88. although the recent reanalysis of the NHANES II suggests intakes lower than this. 20 ± 10 13 ± 6 19 7a 4a — 9 5 — 5 3 — 60 60 61 16 11 — — — — — — 7 14–17 12 4–5 (range 1–46 g) 5–7 2–3 64 65 18 ± 9 19 ± 7 9±6 — — — — — 62 63 Includes fiber from legumes. Intakes assessed by . at 38 g/d (Table 7. NHANES estimates using the fiber table of Lanza and Butrum are 20% less.10 Dietary Fiber Intake in North America (grams per head per day) U. North America Fiber intakes calculated using Southgate analyses from the food balance sheets of Canada and the US show almost identical intakes of 22 to 23 g per capita per day.7). 13 g on average. NSP estimates (Table 7. whereas those in Chile were 35 g.7. 17 g/d in the US.2387_ch7.7). S. and more dietary fiber.34 South America No individual surveys to measure dietary fiber consumption have been published in South American countries. 12 g/d.10). 11 g for women and 16 g for men. fiber intakes were similar to North America.2.2 g. Food balance sheet data suggest that intakes are comparatively high. 6 g from cereals and 11 g from vegetables (Table 7. There is a clear trend between these extremes.2. although there was a wide variation.60 Intakes of 8000 American-Japanese men were lower. May 6. with 5 to 7 g obtained from vegetables and 4 to 5 g from cereals. b Food consumption estimated for 24-h recall.2. Costa Rica and Mexico. ... Cancer. R. Herbs and Spices. the US.. 1982. R. S. H. Rev. Cereals and Cereal Products. 1991. Subj. 1990. most of the Middle East. J. Laine. Nutr. 1991. Varo. Kok. Annual Reports of National Food Survey Committee. K. Letchworth.. and Southgate. 7. Block. Bingham. with average intakes of 11 to 12 g in Japan. 1987.2387_ch7. London. R.. Kok. Williams. McCance and Widdowson’s The Composition of Foods. Her Majesty’s Stationery Office. Relatively little data is available. 56. A critical review of fiber analysis and data. 399. Nutrition and Chronic Diseases. Bingham. J. Evidence from Japan and India suggests that in areas where rice is the staple cereal.. I. J. 6. 4. D. Diet. Holland. D. and van’t Voer. Denmark. 3.. S. H.. Sci. 797. 4. and Koivistoinen. Y.. Unwin. Clin. Her Majesty’s Stationery Office. 1991. 5th Suppl. 57. Ministry of Agriculture. Agric. Assoc. namely a decline in cereal fiber consumption which has been offset to some extent by an increase in vegetable consumption. Rep. 1968. 3rd Suppl. Validations of dietary assessment through biomarkers. Recommended Dietary Allowances 10th Edition. Bingham. intakes of dietary fiber are likely to be similar to those in the West. C. 11. 12. 52. Pero. 705. and Butrum. Paul. Eds. 4th ed. Rep.. MRC Spec.. F. Dietary fiber intakes in the US population.. E. 2001 8:00 PM PATTERNS OF DIETARY FIBER CONSUMPTION IN HUMANS TO 1992 579 interviews of 250 adults aged over 40 years were lower. A.. B. D. Dietary fibre and available carbohydrates in Finnish cereal products. 732. D. World Health Organization. H.. Nutr. Her Majesty’s Stationery Office.. D. London. A. Set. Diet. Vegetables. in Biomarkers of Dietary Exposure. S.. Energy expenditure as a biomarker of energy intake: workshop report. P. D.. UK. vegetables and fruits. 16.. The dietary assessment of individuals. Br.. J. 790. National Academic Press. J. and Japan are similar to those reported in the US and Britain. R. Geneva. and Buss. 3. S. in Biomarkers of Dietary Exposure. 1986. IARC.. Smith-Gordon. and Westerterp. Am. A. A. REFERENCES 1. T. S.. Food and Nutrition Board Sub-Committee on the Tenth Edition of the RDAs.. and Russia.. 10. 14. UK. 1990. there is no data at all due partly to continuing problems in the analysis of dietary fiber and the assessment of food consumption. 3. 9. Trends in dietary fiber intake in Holland. More carefully standardized and validated studies to assess dietary fiber intake worldwide are clearly needed. Tech. P. B. Nutr. Report of the Second IARC Coordinated International Study on Diet and Large Bowel Cancer in Denmark and Finland. K. 1989. Nutr. L. H. 333.. 39. 297. Veijalainen. Household Food Consumption and Expenditure 1966. London. 5. 1985. E. P... 1991. 46.. . 86. J. Jones. newer analyses show that there is only a small range in intake to a maximum of 16 to 18 g/d in Finland. Eds. such as parts of Asia. Lanza.. 1978.. Smith-Gordon. Holland. 15. Finland. Royal Society of Chemistry. and Cummings... Unwin.... 1987. D. Cancer. Am. Fisheries and Food. A.2_fm Page 579 Sunday. and for some areas. Bingham. Dietet. J. Letchworth.. R. and the UK. and Kessler. 1988. G.66 CONCLUSIONS Current evidence indicates that average dietary fiber intake is in the range of 20 to 40 g/d in the majority of populations studied throughout the world. 19 ± 6 g in Tel Aviv. I. Hum. Health Soc. Pett. Bingham. S. Non-starch polysaccharide intake of a representative sample of British adults. and Buss. and Day. Abs. Dietary fibre consumption: new estimates and their relation to large bowel cancer mortality in Britain. Royal Society of Chemistry. 1984. The most accurate data using newer values for the fiber analysis of foods confirm this. F. 8. Worldwide. and 31 ± 11 g and 23 ± 8 g in two rural Kibbutz areas. 2. 13. May 6. however. Lanza.. WHO. Rep. 41. K. J. and van’t Voer. COMA Report on DRV for Food Energy and Nutrients for the UK. P.. London. K. 1981.. C. London. Clin. 1983. 31. 1982. Voeding. Sci.. J. P. 38.. Smart. R.. M. Sci.) 22. J. A. Tyler. L. Brodavec. A. M.. (Data recalculated by S.. R. E. 443. L. Fisheries and Food. J.. Diets of rural families and heads of families in two regions of Yugoslavia.. and Coulander. M. and Cummings. 32.. Logan. E. Bingham. G. M. Bright-See. N. F. 27. 33. J. H.. Changes in the crude fibre content of the American diet... P.. W. N... 824.. C. J. M. Stephen. and Horvat. 1985. Appl. W... R. Milbank. Intakes and sources of dietary fibre in man. 40. 821. 1990. Kuratsune.. J. W. Thompson. T. 272. J. S.. 1983.. Eds.. EUR. G.. and Hackler.. 36A. Englyst. 36. J. unpublished. Choolun..C. E. James. M. Assoc. 24. 1510. B. G.. Buzina. Ledermann.. and Walker. R. A. A dietary survey of 11 areas of the E. 33.. B. Bosschieter. T.. A. and Theander. D.. O. Southgate. and Agater. Ecol. L. and Lacourly. 30. 34.. Food Agric. Sweetnam.. S. Nutr. Dietary survey in 40 year old Edinburgh men.. J. Nutr. A. 324. lifestyle and health in Northern Ireland. Royal Society of New Zealand. 29. Individual variation in dietary fibre intake in different populations. Gregory. OPCS: Her Majesty’s Stationery Office.E. and Seppanen. H. May 6. Noordhoff. Hellendoorn. James. Helms.... 25. Bingham. Englyst. personal communication. 914. 23.. S. in Fibre Hum. Nutr. Cancer Res. and van Oosten-van der Goes.. Marcel Dekker. Arthur. Nutr. Kromhout. Foster. Nutr. E. 97. Laboratory analysis of 3 day composite food samples. A. S. 113. W.. Nutr. Eds.. in Medical Aspects of Dietary Fibre. 41. 51. W. H. Estimates of dietary fibre supply in 38 countries. R. 518. G. Dietet. 36. Van Staveren. et al. R. 12. and Calkins. I. M. Helms. 1978. T. R. N. E. H. 1950–1981.Z. R. with Englyst method (Englyst.. Intakes and sources of dietary fiber in the British population.... Nutr. Low fat intake with falling fiber intake in Soweto. 1986. Hum. H. Food and nutrient intakes on Westray in the Orkney Islands. A. 1969. 42. A. G. Wenlock. A. Non-starch polysaccharide consumption in four Scandinavian populations. A. and Kay. H. Cancer. 736. S. S.. 39. J. Lancet. P. 2. J.. 32. G.. Diet. Seppanen. New perspectives on carbohydrates. B. Eds.. Cole. 31.. E.. A. Keys. Bingham. 629. A. S. 26. Bulletin 20. N. Segal. Fehilly.. 1981. 18. and Cummings. G. 1461..2387_ch7. 26... 80. Katan. 1982.2_fm Page 580 Sunday.. Ferber. P. University of Ulster. and Judson. E. van Montfort.. Appl.... Household Food Consumption and Expenditure. 39. Clin. O. New York. and Wiseman. P. 40.. Dietary fibre in the British diet. Anderson. J. 1985. R. Am. Ireland. 35. 103. 1986. 41. J.... A. J. Nutr.. 8. Heller. Marcel Dekker.. E. Food. 2001 8:00 PM 580 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.. Nature (London). Sharman.. 274.. M. Lockerbie.. D. Am. A short dietary questionnaire for use in an epidemiological study. B. H. Gardiner. E. Spiller. C. L. H. N. N. Reanalysis of composites... Jpn... Cancer. Cummings. T. Day. M.. S. and Theander. Determination of total dietary fiber by difference and of pectin by titration or copper titration. Dietary fiber in the Japanese diet as investigated in connection with colon cancer risk. London. Food Agric. Nutr. Nutr. H. Bingham. T. Bingham. Collinson. and Oliver. 37. Agneletto. J. Rider.. and Walker. A. The dietary and nutritional survey of British adults. . T. Riemersma. Lombardo. M. A. The determination of carbohydrate and its composition in plant material. R. 19. Can. K. 1982. 1313.. Cresta. and Slagman. 1953-1983. J. October. Pharm. M. I. Dietary fiber and 10 year mortality in the Zutphen study. A. H. S. 4. Ministry of Agriculture. Cummings. in The Analysis of Dietary Fiber in Food. G... 21. Kingman. Bull.. and McKeown-Eyssen. Englyst.. J. Yarnell.. S. and van de Bovenkamp. Annual reports of the National Food Survey Committee. L. 50. 28. J. Dietary fiber fractions in cereal products in Britain. Hum. 1984. 1975.. 1989.. Hautvast. B. Her Majesty’s Stationery Office. 1979. 20. J. D. Am. personal communication). 185. M. 1982. Plenum Press. 37A... 1978. A. Wiggins. Enzymatic determination of the indigestible residue content of human food. in The Analysis of Dietary Fibre in Food. Nutr.. Sivell. M. Euratom.. Clin. 25. C.. 261. J. New York. A. in order to determine levels of radioactive contamination.. V. J.. Honda. Am. K. 3RD EDITION 17. P. M. M. M. R. H.. 1980. 1985. Barker.. W. and Cummings.. Katan.. P... Wellington. N.. L. Clin.4218F.. H.. New York. Nutr. 77. Am. 1990. Southgate.. A.. Dietary fiber consumption in an adult Dutch population. Bingham. 1964. J. N. and McNeil. and Robertson. 1982. 1980. A. S. 1291. Food. and Stemmerman.Z. London. C. M. 1983. H. Hum. Nutr. A. Marlett. 323. 44. 51. Wellington. M.. Food. J.. 1982. Am. S. Am. W. 1981.. C. 61. 59. S.. J... 49. Annual Conf. J. S. A. Tanaka. Nutr. Hankin. H.. Thomas.. Studies of Japanese migrants mortality from cancer and other diseases among Japanese in the United States. Diet and colorectal cancer with special reference to fiber intake. F. 64. Colditz. N. Minowa. Dietary intakes and stool characteristics of patients with the irritable bowel syndrome. Clin.. A. C. Dietary fibre intakes in India. Spiller... P. 60. Horwitz. G.. 36. Nutr. P.. K. Nutr. Salmond.. 1981.. 73. N. M. I. Rep. 37A. 33.. L. Jones.. Kay. Platt. Bulletin 20. 335. 34. H. Yamamoto.. W.. 44. 40. Davidson.. and Boucher. 177.. H. Dietet. Unpublished report on Nyasaland held at London School of Hygiene and Tropical Medicine. Wellington.. E. L. Nutr. Assoc. Dietary fibre content of foodstuffs for diabetes. A. Pomare. Shetty. Relationship between calculated and crude fiber intakes of 200 college students. Cancer. Cancer Inst. 89. 55. 1982. 113.. and Pomare. C. L.. B.. 47. Rosner.. J. Helms. A. 97. 155. Am. Clin. Spec... C. Boca Raton. A. F. J. G. Stace. R. 9.. Gastroenterology. 54. Poulson. and Bokram. Orr. 35. W. J. I. fat and fiber intake to risk of colon cancer.. A. 1. J. Proc. and Fisher. W. A. Gastrol.. D. CRC Press. R. London. S. N.. L. Nutr.. Nutr. N. H. Stace. Heilbrun. Int... Bulletin 20. 34. Ruitishauser. Report for the University of Natal for the Swaziland Government (not dated). J. and Kurihara.. Sabry. 225. Yoshi. B. A. and Izumi. McMichael. 1977.. Am. 58. 66. The Physique and Health of Two African Tribes. Rozen. Willett. K. Stampfer. Ser. 34. M. S. J. Am. D.. 613. R. J.. Her Majesty’s Stationery Office. J. Soc. E. Takenkin. Royal Society of New Zealand. 1552. and Aragane.... L. 1986.2387_ch7. 1983... 1989. S. J. Natl. L. Med. and Mosbech. 52. 56. N. Cancer.. N. 2566.. 48. A Study of Swazi Nutrition.. 45... J. Royal Society of New Zealand. E. 63. 53.. 48. E.. J. A. Cancer. A. 1981. and Omolulu. Nutr. M. J. Bingham. 241. Atinmo. 1931. Ed. Denmark.. Dietary intakes. 71. 1981. Fisher... J. 57. . Natsushita. Mbofung. N. Trivandrum. Jorgensen... W. Z. I. 65. 1983. Clin. May 6. L.. 1938/39. C.. FL. Nutr. L. in CRC Handbook of Dietary Fiber in Human Nutrition. Ron. F. C.. B.. Nakamura.. Ind. G. and Speizer. personal communication. and Cummings. 1664. and Bonnett. E. J. Gibson. Bingham. Nutr. 46... Multivariate analysis of diet and serum lipids in normal men. 4. Ahrens. Nutr. 43. C. Mori. I. Pomare. H.. S. Dietary fibre intake in Japan. 23. Peters. 1990. 1981. New Engl.. E. The composition of a simulated American diet. J.. J. C. Studies of Nutrition. Z. 4. Tamura..2_fm Page 581 Sunday. J. 626. 50. C. A.. Paerregaard. 33.. Haenszel. Dietary patterns in Them and Copenhagen. 1981. The Pukapuka and Tokelau Island studies.. A. Bjerrum.. P.. Dietary fiber in the diets of urban and rural Nigerian women. in Fibre in Human and Animal Nutrition.. On the determination of dietary fibre... Potter. and Czochanska.. Hillman. and Fisher... Res. Dietary habits and colorectal cancer incidence in a second-defined Kibbutz population. in Fibre in Human and Animal Nutrition. and Csima. 1984.. A case control study of colorectal cancer in South Australia. G. S. 1982. Patterns of dietary fiber consumption in humans.. stool characteristics and biliary bile acid composition in four South Pacific populations. T. 1968. Trace element intakes of women. and Katz. and Gilks. Br. M. E. Nomura. 34.Z.. 1978. Prior. and Scythes. Stace. R. A. Clin. Relation of meat.. A. B. 62. 2001 8:00 PM PATTERNS OF DIETARY FIBER CONSUMPTION IN HUMANS TO 1992 581 43. 1987. Peters. 2_fm Page 582 Sunday. May 6.2387_ch7. 2001 8:00 PM . and resistant starch intake. Many of the studies used traditional techniques 0-8493-2387-8/01/$0. for example. non-polymeric polyphenols. 2001 8:02 PM CHAPTER 7. randomized surveys of the Australian population have been undertaken17–23 and. Since 1983.3 Dietary Fiber. In the absence of a consensus. Record INTRODUCTION Until the early to mid-1980s. where assessed. and Resistant Starch Intakes in Australia Katrine I. Baghurst. Peter A.” As pointed out by Southgate. saponins. the Life Sciences Research Office of the Federation of American Societies for Experimental Biology1 adopted a definition of dietary fiber as “the endogenous components of plant materials in the diet which are resistant to digestion by enzymes produced by humans. and inorganic constituents. and Sally J. METHODOLOGIES The subjects and methodologies employed in the surveys described here are shown in the relevant tables alongside the relevant intake data.3 It can be considered to include some components of what is now known as “resistant starch. May 6. furthermore.3_fm Page 583 Sunday.00+$1. so survey analyses relied on British food databases. nonstarch polysaccharide. Nonstarch Polysaccharide. and usually small. subpopulations and. The earlier surveys of the Australian population4–16 were limited to an estimate of total dietary fiber using the definition of the time. a de facto working definition of the term “dietary fiber” appears to be emerging which includes all nonstarch polysaccharides and lignin from plants. Australian databases were limited. Baghurst.2 this definition is virtually identical to that for “unavailable carbohydrates” as originally defined by McCance and Lawrence in 1929.2387_ch7. In 1987. the only nationally based information about dietary fiber intake in the Australian population came from the apparent consumption or food disappearance data collected by the Australian Bureau of Statistics. those forms of resistant starch that arise as a consequence of cooking and processing techniques. the data is given for dietary fiber.50 © 2001 by CRC Press LLC 583 .” One difficulty with the word “endogenous” in this definition is that it excludes. lectins. The only data available from direct survey of individuals was on selected. several national or statewide. It also excludes substances which are intimately associated with the major components of dietary fiber and which are capable of having important nutritional and/or physiological effects such as phytates. as assessed using the Southgate . Figure 7. the intake of NSP mirrored that of total dietary fiber. in particular those conducted by the CSIRO.0 g at 4–7 years.4 and 7. In general.3. May 6. Between 1988 and 1993. to a lesser extent. there was a small increase in the number of people reaching the target of 30 g/day recommended by the Commonwealth Department of Health. fruit. When comparing the CSIRO National Dietary Surveys of 1988 and 1993. the techniques for sampling varied widely. if the more generally representative data from the earlier period are used for comparative purposes.17–23 Assessment of trends over time is difficult because of the non-random nature of earlier surveys and the variety of measurement techniques employed.8 g at 8–11 years.2387_ch7.5 show the intakes and major food sources of total nonstarch soluble and insoluble polysaccharides in the CSIRO National Dietary Surveys of Adults (1988 and 1993). the percentage of fiber provided by vegetables had decreased. intakes of women increased with age and occupational status.3. Breakfast cereals were a more important source of dietary fiber in older people and for people of upperoccupational status. 2001 8:02 PM 584 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. as detailed in the tables. However. In the 1993 survey. but a large number of the surveys employed a variation of the quantitative food-frequency technique developed by the CSIRO Division of Human Nutrition. and 20% by fruit (Table 7. The more recent random surveys of adults indicate that approximately 33% of dietary fiber is currently provided by cereals. Data from the CSIRO National Dietary Surveys (adults only. while fiber continued to increase over the period. 3-day records. there was a paucity of data on dietary fiber intakes of young children in Australia apart from smaller. and the processed meat categories increased. while the yield from breakfast cereal. the increases were much smaller than had been observed from the late 1970s to late 1980s. RESULTS AND DISCUSSION Tables 7. The figures for adults indicate an increasing relative contribution to total fiber intake from the cereal group over the last decade and a lessening of the relative contribution of vegetables and.4 g/day in 2–3-year-olds rising to 16.0 g/day at 16–18 years of age. or diet history which will not be detailed here.24 As with the methodologies used. 21. the 1995/6 National Nutrition Survey of the Commonwealth Department of Health and Aged Care showed intakes of 13.3. 31% by vegetables. from 17–18 g/day in the late 1970s to 24–28 g/day in the mid-1990s. it does appear that dietary fiber intake has increased over the past 20 years or so by about 50%. used a random selection procedure based on the Electoral Rolls of the States of the Commonwealth.4 g at 12–15 years.1 shows the percentiles of intake of dietary fiber as well as NSP and resistant starch in the 1993 CSIRO survey. Until recently. The increase was most marked in the older age groups.3. well under half the population still failed to reach recommended intakes. for women in the highest occupational group. non-random population groups. 18.3).3. 3RD EDITION such as 24-hour recall or record. NONSTARCH POLYSACCHARIDES Tables 7.2 show the dietary fiber intakes and the group mean fiber-to-energy ratios from selected non-randomized surveys from the late 1970s and early 1980s. However.3_fm Page 584 Sunday. However. particularly for people in the top occupational group. aged 18 yr plus) of 1988 and 1993 showed that. and 23.3.4–16 and from randomized population surveys carried out by the Commonwealth Government and the CSIRO Division of Health Sciences and Nutrition since 1983.1 and 7. and for men from the lowest occupational group. Many of the random surveys. but this was not evident in men. fruit and fruit juice. 4 31.8 12. method.0 15. and a number of inconsistencies in existing databases. NON-STARCH POLYSACCHARIDE AND RESISTANT STARCH INTAKES IN AUSTRALIA 585 Mean Daily Fiber and Energy Intakes in Selected.6 8.0 19.6 8.8 13.8 24.7 4-day record 8.9 17.0 9.0 20.1 22.5 9. Non-Random Australian Population Samples from 1977 to 1995 Energy (MJ/day) Fiber (g/day) Fiber/Energy Ratioa 9.3 21. Because of the small amount of available data. and soluble accounted for some 9 g. record 200 24-hr.5 18.0 7.6 41.4 27.1 11.53 44.2 24.6 21.5 21.0 27.3 32.1 24. the varying techniques used to assess resistant starch. record 8. 2001 8:02 PM DIETARY FIBER.5 8.2 9.0 15.4 19.0 15.0 21. May 6.0 8.0 21.5 20.4 7.1 7.3 37.2 8.9 23.5 25.1 15. a food database was constructed using analytical data presented at the EURESTA25 meeting and from published figures.7 22.3.1 24.76 20.1 21.2 30.6 Survey Sex No.3 20.2 8.4 6.4 9.0 21.5 9.9 39. record 82 24-hr.2 31.0 24. Insoluble NSP accounted for about 12 g of the total.3 10.3_fm Page 585 Sunday.4 26.9 11.4 20. urban SA8 11 yr 13 yr 15 yr Children urban WA9 11 yr 12 yr Rural children SA10 aged 13 yr Urban adolescents SA 14–15 yr4 Pregnant women11 Elderly Community12 Institutionalized13 Nursing home14 Religious groups15 Adventist (vegetarian) Adventist (omnivore) Mormon (omnivore) Vietnamese migrants16 a FFQ FFQ FFQ FFQ M F M F M F M F F 161 172 72 69 91 M and F M and F M and F 27 87 30 Hist.1 18. record M F M F M F F } } } FFQ FFQ 3-day record } } } Table 7.5 40.4 7.7 23.2 19. record 113 24-hr.7 15.5 10.8 7.0 15.0 16.4 22.4 19.5 21.7 26.4 9.9 6.5 7.9 7.0 7. It is likely .83 7.3 24-hr.0 11.4 33.2 23.0 19 17 24.5 20. RESISTANT STARCH To estimate current resistant starch intakes in Australia.2387_ch7.9 22. with a mean intake in 1993 of some 21 g/day for both adult men and women.0 7.1 Group mean ratio g fiber/10 MJ.6 10. 3-day record 98 24-hr. Method Urban community4 (random within age bands) Fitness intervention group5 M F M F M F M F M and F 481 441 581 454 142 60 300 300 300 FFQ M F M F M F 124 106 121 116 104 104 Public servants6 Parents of 10-yr-olds7 10-yr-old children of above group7 Children.4 20.9 24.6 7.8 22. care should be taken in interpreting the data.3 19.7 22.7 25.9 20.9 21. 7 shows the estimate of NSP plus resistant starch from the 1993 survey.5 10.6 24.4 26. Resistant starch density in 1993 was some 5. rising from 4.)21 1988 1993 National Elderly Survey (55–75 yr.0 10.1 8.1 31.9 32.3 8.0 21.0 25.0 25.1 37.9 20. recall 24-hr.4 8.3 10.0 23.1 9.7 g/day in men and 26.4 22.48 23.3 7.0 26.8 32.0 g/day for women in 1993.2 29.6 10.2 7. Method Fiber (g/day) Fiber/Energy Ratioa (fiber/10 MJ) 11.2 27.4 g in 1988 for women (Table 7.3 33.7 32.3 21.6). recall CSIRO Surveys Victorian State surveys (18+ yr.5 8.0 7.4 32.7 11.3 g/day of resistant starch for men and 5.1 7.2 18.0 8.4 8.9 g in 1988 for men and 4. with approximately 4% of starch in men and 4.1 23.4 g per 10 MJ in men and 6.8 31.4 7.)16 National schoolchildren’s survey 198517 National Nutrition Survey 1995/6 (19+ yr. May 6.2 g in women.2 28. The main .2 Random Surveys of Australian Population Samples from 1983 to 1995 Survey Sex No.8 27.4 17.5 15.) 1989/9022 a b M F M F 1321 1595 1321 1595 FFQ M F M F M F 445 456 445 456 445 456 FFQ M F M F M F 1115 1200 736 997 606 707 FFQ FFQ FFQ FFQ FFQ Group mean ratio.7 26.9 21.3.1 25.8 Energy (MJ/day) Commonwealth Government Surveys National Adults Surveyb 1983 (25–64 yr.)19 1985 1990 South Australian State surveys (18+ yr.3 g per 10 MJ in women.2387_ch7.2 10.1 25.3.3 27.3 7. record 24-hr.0 28. There was little evidence of any age.)18 M F M (10–11 yr) M (12–15 yr) F (10–11 yr) F (12–15 yr) M F 3021 3234 912 1719 925 1668 5079 5770 24-hr.3 21. Table 7.7 28.4 34.3 32. 3RD EDITION Table 7.7% in women being resistant.3 10.3_fm Page 586 Sunday.3 10.7 18.0 7.3.3 26.)20 1988 1990 1993 National Dietary Surveys (18+ yr.9 27.9 26. Metropolitan sample only.5 34.9 27. The analysis of the 1988 and 1993 CSIRO National Dietary Surveys gave a mean figure of about 5. etc.7 20. The composite food database was used to estimate intakes using data collected from the CSIRO Food & Nutrition Surveys of 1988 and 1993 and the Commonwealth Department of Health’s National Dietary Survey of Adults undertaken in 1983 and of Schoolchildren undertaken in 1985.or occupation-related trends in the density of resistant starch in the diet. 2001 8:02 PM 586 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. that some of the analyses based on chemical assessment of resistance might underestimate resistance in the human gut and that much individual variation will occur with respect to physiological functions such as degree of chewing foods.9 26. showing an average for these combined components of 26.3 27.0 9.4 25.8 27. 2% of starch being resistant. Total NSP (g/day) 30 20 10 0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 Percentile 10 Minimum intake 0. For children. . sources of resistant starch found in the 1988 and 1993 CSIRO surveys of adults are shown in Table 7. Resistant starch (g/day) 8 6 4 2 0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 Percentile of intake Figure 7.12 g Maximum intake 17. there was no consistent age trend. In girls. May 6. nonstarch polysaccharides. for adults. and resistant starch in the Australian population. This equated in both girls and boys to about 3.1 g per 100 g starch.8.35 g Maximum intake 60.2387_ch7. with intakes ranging from 3. About 42% of resistant starch was provided by cereals.39 g c.3_fm Page 587 Sunday. At that time.0 g in older men and from 3.7–4. NON-STARCH POLYSACCHARIDE AND RESISTANT STARCH INTAKES IN AUSTRALIA 587 50 Minimum intake Maximum intake 6.8 g per 10 MJ energy.1 Percentiles of intake for dietary fiber.2 g/day in younger men to 4.3. with 26% from vegetables and 22% from fruit and fruit juice. the density of resistant starch seems to have approximated 4.7 g/day in older women in 1983.3. Dietary fiber (g/day) 40 30 20 10 0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 Percentile 40 Minimum intake 4. with a resistant starch density of some 4.2 g/day in younger women to 2. 2001 8:02 PM DIETARY FIBER.7 g a.2 g 78.2 g per 10 MJ or 3.7 g/day across age categories.4 to 3.6 g/day in 15-year-old boys. intakes rose from 4 g/day in 10-year-old boys to 5.07 g b. A re-analysis of the 1983 and 1985 CDH National Dietary Surveys of Adults and Schoolchildren gave estimates of resistant starch intakes ranging from 5. 0 2.4 9.1 26.8 28.2 10. 3RD EDITION Sources of Dietary Fiber in the Australian Population Percentage of Total Dietary Fiber Derived from Food Groups Vegetables Fruit Brassica and Legumes.0 9.4 17.3 8.3 6.8 27.2387_ch7. Peas.1 25.3 17. 2001 8:02 PM 588 Table 7.2 23.4 CSIRO National Dietary Surveys — Dietary Fibera and Nonstarch Polysaccharide (NSP) Intakes and Densities Men Intake (g/day) Dietary fiber Total NSP Soluble NSP Insoluble NSP Density (g/10 MJ) Dietary fiber Total NSP Soluble NSP Insoluble NSP a 1988 1993 Women 1988 1993 27.6 15.1 8.9 30.2 15.5 Othera Food Sources of Nonstarch Polysaccharides from CSIRO National Dietary Surveys of Australian Adults 1993 (percentage contribution of food groups) Food Sources Total NSP Soluble NSP Insoluble NSP Breakfast cereals Breads and crackers Rice and pasta Fruit and fruit juice Vegetables Processed meats and cooked meat dishes Confectionery.0 9.8 27.1 12. Table 7.6 20.5 27.7 8. beverages.7 37.7 13.2 12. May 6.5 5.5 As measured by Southgate method.8 20.1 11. snack foods. cakes.3.9 Includes composite dishes such as pies. and seeds..5 26.3.6 8.6 32. Source: Department of Community Services and Health (DCSH).8 14.5 34.3 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.0 20.2 6. nuts. etc.2 4.8 11. pizzas.2 9.5 12.9 11.4 25.3 33. and biscuits Takeaways and snacks 8 18 5 20 34 5 7 16 3 24 24 5 9 20 6 17 34 5 7 7 7 3 4 3 .3.9 20.8 4. Green-Leafy and Beans Total Cereals Survey Urban community samples 1978–83 (CSIRO) DCSH National Adults 1983 DCSH National Schoolchildren’s 1985 CSIRO National Survey 1993 a 32.9 39.3_fm Page 588 Sunday.9 8.7 21.6 11.5 32. Table 7.2 37.4 22.3 21.9 20. 3_fm Page 589 Sunday.05 4.08 6.63 6.63 5.6 21.2387_ch7.38 5.3.8 2.58 5.41 5.91 4.5 26.3.7 26.8 27.91 5.34 5.61 3.9 25.78 4.0 F 1993 .8 8.57 6.6 Age 18–29 30–39 40–49 50–59 60+ Occupation Top 2nd 3rd 4th 5th Resistant Starch Intakes and Densities by Demographics from CSIRO National Surveys M 1988 Intake (g/day) M F 1993 1988 F 1993 M 1988 5.28 4.17 4.81 4.4 28.01 5.8 4.6 26.48 5.66 4.40 5.26 5.96 5.6 24.8 24.94 4.6 2.72 4.1 0.8 25.73 6.5 26.1 0.12 4.35 6.76 5.5 0.12 6.98 4.3 0.7 26.05 5. May 6.5 0.31 5.3 5.74 5.56 Table 7.8 Density (g/10 MJ) M F 1993 1988 Men (g/day) Women (g/day) 26.4 0.6 2.4 0.2 26.92 4.57 5.1 22.45 4.2 0.41 4.30 4.7 Sum of Intake of Resistant Starch and NSPs from CSIRO National Survey 1993 Total Age 18–29 30–39 40–49 50–59 60+ Occupation Top 2nd 3rd 4th 5th Table 7.69 4.50 4.36 6.7 4.20 4.05 4.52 5.11 5.63 4.67 4.9 26. NON-STARCH POLYSACCHARIDE AND RESISTANT STARCH INTAKES IN AUSTRALIA 589 Table 7.10 5.53 4.2 26.38 4.25 6.3 24.70 4.94 4.51 4.46 6.26 5.3 11.99 4.35 5.73 5.30 5.19 5.13 4.0 24.95 5.85 4.92 5.26 5. 2001 8:02 PM DIETARY FIBER.7 26.6 5.16 5.3.87 6.16 5.7 3.40 5.30 5.88 4.12 4.5 24.17 5.25 5.5 25.09 5.2 28.7 26.0 0.06 5.0 0.4 Sources of Resistant Starch % Contribution to Total Intake Breads and crackers Vegetables Fruit and fruit juice Rice and pasta Breakfast cereals Takeaways and snacks Processed meats + Confectionery + Dairy and ice cream Chicken and fish + Drinks Spreads and sauces Red and organ meat 1988 1993 26.5 26.1 27.2 3.1 28.63 5. . English. 4. S. nonstarch polysaccharide. 19. Elsevier Science. 1998. in selected Australian subpopulations. 18. and Magnus.. G.. 9. Stud. Aust.. Australia. Southgate. I. D. Nutr. 5. 1993. Wahlquist. Essex. Aust. 1. 15. L. Aust. K. S. 1987. Wallace. et al. Nutritional profile of recruits to a fitness program. A more detailed review of dietary fiber. A. J. R. Syrette. A. and Boulton. 3RD EDITION COMPARISONS OF AUSTRALIAN RS INTAKE WITH OTHER COUNTRIES Estimates of resistant starch intake in developed countries vary widely in the literature. May 6. R. Parish. MRC Special Report Series no.. J. 1989. M.. Macronutrient and micronutrient intakes at ages 11. K. A. and Bennett.. J. The estimates of Australian intakes show them to be at the upper end of the European range. Nutrient Intakes and Physical Measurements. J. 10. M. L. 13. 2. Cashel. 1983. Waters. Food Nutr. Dietary intakes of public servants. What are Australians eating? Results from the 1985 and 1990 Victorian Nutrition Surveys. 14... and Bell. D. Royal Society of New Zealand. K.. J. J. Intake and sources. and Record. J. Syrette. K. Baghurst. K. and 15 years: age and sex differences. Berzins. Australia. and Baghurst. and Strohm. Dietary intake of ten-year-old children and their parents with respect to occupational status.. The relationship of blood pressure to diet and lifestyle in two religious populations. 11.. and Syrette. 17. Ash.. Commonwealth Department of Health and Aged Care and Australian Bureau of Statistics.. 9. Eds. 1985. 12. P. A. Hlth.5 to 6.. AGPS. 1987.. A. of dietary constituents implicated in the aetiology of chronic disease.. Record. Nutr... Lewis. 48 (9).. R. 715. Nutr. 2. 2001 8:02 PM 590 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Dietary fiber intakes of elderly Australians. R. A. 20. Rouse. 40. 1994. Cashel. 1988. I. Comm. Life Sciences Research Office. M. Nutr. Intakes of selected nutrients in Year 7 Western Australian schoolchildren. J. MD. 135. 65. Physiological Effects and Health Consequences of Dietary Fibre.. I. Med. Estimates for developing countries range from 9–10 g/day to 30–40 g/day for countries with high starch intake. 1987. 1991. S. Jenner. E. English. and Record. Baghurst. J.. A. 11. Diet. I. A. Hypertension. K. and Beilin. CSIRO. K. A. T. AGPS.. Soc.K. 99. Armstrong.. Federation of American Societies for Experimental Biology. and Record. and resistant starch intakes in Australia and New Zealand was published in 1996 and is available from the authors.. 1929. Res. Determination of Food Carbohydrates.26 REFERENCES 1. A.-M. J. I. E.. and Tran. National Dietary Survey of Adults: 1983. Nutrient Intakes. A. 3. 1. M. and Smith. National Nutrition Survey 1995... 6. Baghurst. U. Adelaide. Canberra. K. Baghurst. unpublished data. Diet.. J. Bennett. comparisons between weekdays and relationships with socioeconomic status. others attempt to include an estimate of all possible forms of resistant starch.. Baghurst. Australia. I. unpublished data. Canberra. Magarey. National Dietary Survey of Schoolchildren (Aged 10-15 Years): 1985.. I.. W. 13. unpublished data. Aust. I.. The Carbohydrate Content of Foods.3_fm Page 590 Sunday.2387_ch7. 2.0 g/day for a number of European countries. J. 188. Dietary intakes of children in Port Pirie. 51:3. Proc. 50. McCance. 33. unpublished data. K. Bethesda. K. Baghurst. Waters. 1983.. I. 1983. M... S. Baghurst. K.. HMSO. P. T. Hope. 7. Dietary profile of Vietnamese migrant women in Australia. Flint. Australia. J. 1991.. 16. D. A recent data-based analysis reported at the EURESTA summary meeting in 1994 gave intakes ranging from 3. Nutrient Intakes. J. The Adelaide Nutrition Study. J. The range of estimates reflects the fact that some estimates relate only to retrograded starch. Canberra. . S.. 12. Flint. S. 160. J. in Fibre in Human and Animal Nutrition.. M.. L. K. J.. B. 111. 8. Baghurst. S.. Dietary fibre intake in a group of institutionalised elderly and the effect of a fibre supplementation program on nutrient intake and weight gain. D.. 1985. and Miller.. 1991.-M. 143... with figures ranging from 3–4 g/day to 15–20 g/day. and Lawrence. A. Sedgwick.. and Down.. D. London. 2.. ABS. 1983. K. Dietary fibre.. . May 6. Syrette. S. Food Australia. and Record. Nutrition in South Australia from 1988 to 1993. Record. and Baghurst. P. A. Syrette. 1988 and 1993. and Powis.2387_ch7.. S. Health Stud. I. Adelaide. P.. J. I. J.. S. Baghurst.. S. 11. 26. Results from the CSIRO Australian Food and Nutrition Surveys. I.. Baghurst.. G.. Baghurst. 2001 8:02 PM DIETARY FIBER. K. I. March suppl. A. 22. A.. P. 1995. 1989. CSIRO. J. 1984. K. Baghurst.. J. J. A. K. K.. 8. A. 1996.. EURESTA summary meeting. 24. 25. Adelaide. P. nonstarch polysaccharides and resistant starch: a review.. CSIRO. s1-36.. Comm. S. 1991. Food and Nutrition in Australia — Does Five Years Make a Difference.3_fm Page 591 Sunday. Australia.. Baghurst. A. J. Record. NON-STARCH POLYSACCHARIDE AND RESISTANT STARCH INTAKES IN AUSTRALIA 591 21. France. CSIRO. Australia. Australia. J. 23.. 1994. National Dietary Survey of the Elderly. and Record.. I. Baghurst. A computerized dietary analysis system for use with diet diaries or food frequency questionnaires. Record. and Baghurst. Adelaide. Baghurst.. May 6.2387_ch7.3_fm Page 592 Sunday. 2001 8:02 PM . 2 indicate the contribution of crude fiber from various foods in north China peasant diets1 as calculated by Zheng from the annual dietary surveys in 1979. The data in Table 7.4.2 The age-adjusted death rate for colorectal cancer in China was 5. Chinese diets are rich in vegetables. However.3 lists the crude fiber contents of processed rice and wheat. The supply of staple and nonstaple food varies in amount and varieties with season. Table 7. The amount and varieties of plant subsidiary foods are dependent on the climate.4.00+$1. The crude fiber content of the Chinese daily diet is much higher than the daily crude fiber intake in England. sorghum. May 6. which was estimated to be about 4 g per capita.4. which provide about 80% of the daily caloric intake. as shown in Table 7. National nutrition surveys carried out in China in 1959 and 1982 indicate that the consumption pattern of plant foods and crude fiber intake differ in different parts of China.4. the secondlowest in the world.4 The comparison between crude fiber and neutral detergent fiber contents in selected Chinese foods is listed in Table 7.49 per 100. In general.1. 2001 8:04 PM CHAPTER 7.50 © 2001 by CRC Press LLC 593 . millet. Crude fiber is mainly from cereal grains and vegetables.5 0-8493-2387-8/01/$0.3 The Chinese government has stipulated the standard of rice milling and wheat flour extraction in order to retain more nutrients. the predominant grain is rice and in the north.4 Consumption of Dietary Fiber–Rich Foods in China Zhi-Ping Shen and Su-Fang Zheng China is a big country with great varieties of foodstuffs. etc.4_fm Page 593 Sunday. the most important components of the daily diets are grains and cereals.000 in 1979.4. the main staple foods are wheat and coarse grains including corn. In south China.2387_ch7. 66 0.2387_ch7.02 — 0.3 0.3 11 173 344 352 21 305 13 1 11.44 0.2 0.03 0. Long Grains Crude Fiber (g/100 g) 0. dried Vegetables Fruits Nuts Crude fiber Table 7.1 Consumption Pattern of Fiber-Containing Foods and Crude Fiber Intake in Different Parts of China (Grams/Day/Capita) Food Rice Wheat flour Coarse grains Starchy tubers Legumes.47 2.4 0.2 4.8 37. 1st Standard.07 — 2.4.81 3.2 Shanghai South Jiangxi South Hebei North Shanxi North 90 314 125 35 8 440 82 4 8.46 4.1 6.4.5 463 44 — 43 4 326 9 1 6. Henan Crude Consumption Fiber Contribution (g) (g) (%) Food Rice Wheat flour Coarse grains Starchy tubers Legumes Vegetables Fruits Nuts Seaweeds Table 7.06 0.4 0.42 4.6 .2 34.2 26.5 Extraction Whole 75% extraction 81% extraction 85% extraction Wheat Flour Crude Fiber (g/100 g) 2.6 Crude Fiber Contents of Processed Cereal Grains in China Grade Husked Fine Standard. 3RD EDITION Table 7.3 Gansu Northwest 3 610 49 149 5 147 34 — 6.8 — 0.06 1.29 2.76 0.2 0.9 558 13 — 110 9 383 1 1 7. Henan Crude Consumption Fiber Contribution (g) (g) (%) 66 456 225 11 20 830 85 — 1 0.02 0.4.2 0. 2nd Rice.4_fm Page 594 Sunday.6 Contribution of Crude Fiber from Various Foods in North China Peasant Daily Diets Linxian County.52 — 0.2 3.3 Beijing North 13 250 300 646 21 583 3 1 — 0.3 0.2 — Xianxiang County.7 24. May 6.4 0.4 26.5 11.2 21. 2001 8:04 PM 594 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.7 13 350 252 97 3 280 30 2 9.7 0. 2001 8:04 PM CONSUMPTION OF DIETARY FIBER–RICH FOODS IN CHINA Table 7.89 6..04 14.50 1.11 2. H. 1979. yellow Vegetables Cabbage.24 0.41 7. May 6.42 8. Dai.57 2.13 0. Zheng. 3.53 18.57 8.30 17. Comparison between crude fiber and dietary fiber contents in foods. Zao. The crude fiber intake of peasants in north China.. Nutr.66 21.24 9.43 10. 4. J.59 1. Am..03 3. Beijing.55 18. 2.94 7. Ed.2387_ch7.66 7. X.81 7. Chinese Cabbage. common Celery stem Chive Garlic green Garlic shoot Onion shallot Spinach Roots Carrot Lotus root Potato Yam CF NDF (g/100 g dried edible portion) 1. S.04 3.40 10.4..70 0.79 8.4 595 Comparison Between Crude Fiber and Neutral Detergent Fiber Contents in Selected Chinese Foods Food Cereals Corn Corn meal Millet Sorghum Rice. unpublished.21 18.23 11. 417.38 11.19 0. 29. Colorectal cancer. Z. Clin.42 4. glutinous Rice.37 10.12 8.38 6. celery Cabbage. Definition of dietary fiber and hypothesis that it is a protective factor in certain diseases. in The Investigation of The Cancer Mortality in China (in Chinese).03 8.17 0.50 8. Acta Nutrimenta Simica.36 1.. 5..16 10.88 0. People’s Medical Publishing House.54 3.70 0. 9. Trowell. 1981.99 11.48 4.49 0.20 14. 333. Chinese Academy of Medical Sciences. long-grain Rice.90 4.74 3.28 0. Institute of Health.74 3. dried Adsuki bean Kidney bean Mung bean Soybean. 3rd ed. Food Composition Table (in Chinese). round-grain Whole wheat flour Legumes.70 3. 1989.4_fm Page 595 Sunday.90 11. green Soybean. . 1976..09 REFERENCES 1. Beijing. People’s Medical Publishing House. The National Cancer Control Office of the Ministry of Health. May 6. 2001 8:04 PM .2387_ch7.4_fm Page 596 Sunday. 61 8.50 © 2001 by CRC Press LLC 12.5.0 28.3 Ref.00+$1.4 4 597 .64 ± 0.6 ± 4. May 6.0 ± ± ± ± 0. The dietary fiber consumption can then be calculated by tables using the PaulSouthgate method1 to obtain the total fiber consumption and not just the crude fiber consumption.1 which is based on English alimentation and. 2 3 6. The results are identical in two studies. but in the third4 the consumption of dietary fibers is far more important. while the consumption of cereal fiber is virtually the same in the three studies.5_fm Page 597 Sunday. the content of dietary fiber was calculated with a new analysis of fiber content carried out by Southgate himself on French bread and flour which showed a higher proportion of fiber in French products (Table 7.70 10.5.6 ± 0. the total consumption of fiber is higher in the study by Macquart-Moulin et al.8 11.5. In this chapter we will make a distinction between direct and indirect methods for evaluating this consumption.1).8 Cereal Fiber (g/day) 3. English bread. Therefore. Three studies of this type have been published in France (Table 7.4 Table 7.70 1.81 ± 1.08 7. DIRECT METHOD OF EVALUATION This method is based on dietary recall in a sample of people as representative as possible of the general population.0 11.7 ± 5.2387_ch7. 2001 8:05 PM CHAPTER 7.2). in particular. the calculation of the dietary fiber content was made with Southgate’s table. in the third study.1 Average Daily Consumption of Dietary Fiber in France Calculated from Dietary Recall Total Dietary Fiber (g/day) Females Females May October Males May October Females Males 0-8493-2387-8/01/$0.34 28. more generally.0 8.88 8. about food consumption. This difference can be partly explained: in the first and second studies.6 ± 6.5 Consumption of Dietary Fiber in France (1950 –1981) Yves Le Quintrec INTRODUCTION Little has been published about the consumption of dietary fiber in France and. consumption means the gross amount of foodstuffs made available at the wholesale stage. We have. it remains high in the agricultural population and constitutes a large percentage of the total awayfrom-home consumption of vegetables. Still. The consumption of bread has been decreasing for many years. Agricultural Statistics of the CEE (European Economic Community) or Eurostat. except for citrus fruits. because no account is taken of losses at the retail trade stage or at the household level. but this tendency seems to have now stopped. two major sources of information in France: 1. CEREAL CONSUMPTION (TABLES 7.0 INDIRECT METHODS OF EVALUATION These methods consist of measuring the total consumption of a given food in a given population and then dividing it by the number of individuals to obtain an evaluation of the average per capita consumption.5.5. statistically representative of the total French population. Bread consumption appears to be higher in the agricultural population than in the urban (Table 7. in all forms: direct. the study being completed in the course of 1 year.71–3. jam. 3RD EDITION Table 7.5.7) The total cereal consumption has been quite stable for the last 10 years.8 TO 7. May 6. The consumption of dried pulses is very small. the consumption of fruits (including bananas and citrus fruits) is certainly higher in the urban population than in the highest classes of society (Table 7. Unfortunately. Consequently the consumption is overestimated. still. food consumption is always underestimated.93 3.7 3. are each examined for 1 week. preserved. therefore. etc. bread remains the chief source of cereal fiber in the French diet.5.5. The major cereal in France is wheat (soft wheat).2387_ch7. etc. which represents at least 10% of the total consumption of vegetables and about 15% of the fruit consumption.96 2. is no longer on the rise.). only the “at-home” consumption is estimated and not the away-from-home consumption (including restaurants. It also includes the awayfrom-home consumption.11).5. CONSUMPTION OF FRUITS AND VEGETABLES (TABLES 7. 2.9) and INSEE (Table 7.12) The consumption of potatoes has been steadily decreasing for many years.5. 10. but this result may be due to the lack of information about away-from-home consumption (lunch is frequently taken in a restaurant or a canteen by the urban population). The consumption of rice is increasing but is still very low.000 households.67–5.5.6). Therefore. Nevertheless.7 According to the data of INSEE. canteens. interesting comparisons between different population subgroups can be made.5.6 Here.5_fm Page 598 Sunday. and processed products.3 TO 7.2 Dietary Fiber Content (g/%) of Bread and Flour from Southgate French Bread4 English Bread1 French Flour4 English Flour1 5. The estimation by Eurostat probably is more realistic because it reflects all forms of consumption including preserves. . 2001 8:05 PM 598 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.11). There is a great discrepancy between the data on this consumption collected by Eurostat (Table 7. The consumption of other vegetables and fresh fruits. Inquiries by INSEE (National Institute of Statistic and Economic Studies5). the consumption of bread and potatoes has been decreasing to a considerable extent.65 4. 2001 8:05 PM CONSUMPTION OF DIETARY FIBER IN FRANCE (1950–1981) 599 Total Human Consumption of Cereals in France (1000 t/year) (Eurostat’s Data6) Table 7. but this increase cannot compensate for the lack of cereal fiber.93 INTAKE OF DIETARY FIBER An estimate of average daily fiber consumption can be made from these data. with a large margin of error.16 5.32 37. From Eurostat (Table 7. May 6. For the last century (Figure 7.5 Table 7. the at-home consumption is surprisingly not much lower: 16 to 20 g/day.5.78 3. From INSEE.2387_ch7.9 .5 11 12. including 116 g of cereal fiber.85 to 24.5.1).5.3 Year Total Cereals (excluding rice) Total Wheat Soft Wheat Other Cereals (excluding wheat and rice) Rice 1970−1971 1975−1976 1980−1981 5089 5016 5344 4978 4898 5180 4444 4426 4650 111 118 164 160 211 278 Table 7. the “average Frenchman’s” total dietary fiber intake ranges from 17. Therefore.14). the consumption of fruits and vegetables has been increasing. who probably take the greatest number of meals at home. the values are a little higher: 22 to 26 g/day and 19 to 24 g/day.5.60 g/day.00 5.5.57 78.37 3.13).5_fm Page 599 Sunday.5. the intake of dietary fiber currently is a great deal lower than in the years 1900 to 1930.6 Average Annual At-Home Consumption of Cereal Foods (Kilograms per Capita) (INSEE’s Data 19805) Total population Agricultural population Nonagricultural population Population of the greater Paris area Bread (total) Noodles Wheat Flour Rice 48.5.72 5.49 7.28 4.5 Average Daily Consumption of Wheat and Rice (Grams per Capita) Calculated from Annual Consumption Year Total Wheat Wheat in Flour Equivalent Bread Rice 1970–1971 1975–1976 1980–1981 268 257 261 206 198 201 199 182 148 5.77) (Eurostat’s Data6 ) Year Total Cereals (excluding rice) Total Wheat Soft Wheat Other Cereals (excluding wheat and rice) Rice Before 1970 1975–1976 1980–1981 80 73 76 78 79 75 70 64 67 2 2 1 2 4 4 Table 7. Conversely.4 Average Annual Consumption of Cereal (Kilograms per Capita) in Flour Equivalent (= Grain × 0.86 2.41 5. If one calculates the quantity of dietary fiber in the diets of farm owners and the unemployed (Table 7.57 45.98 3.58 4. 41 4.13 Unemployed 600 Table 7.11 6.00 Medium-Level Executives 39.5.5_fm Page 600 Sunday.87 2.96 4.12 Farm Owners 64. May 6.55 Farm Laborers 42.22 5.92 2.75 Blue-Collar Workers Average Annual Consumption of Cereal Foods by Occupation Group (Kilograms per Capita) (INSEE’s Data 19805) 48.53 4.38 3.17 4.85 Upper-Level Executives 38.43 4.12 5.7 2387_ch7.32 2. 2001 8:05 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.97 Self-Employed Professionals 32.03 6.39 3.99 3.27 2.86 White-Collar Workers 51. 3RD EDITION .65 2.63 4.58 3.75 5.52 4.49 3.Bread Noodles Wheat flour Rice 81.50 5.53 3. 38 63. In both groups.87 22. the total dietary fiber intake (12. Meyer found a negative correlation between consumption of fruits and mortality due to colonic cancer8 but did not calculate the total dietary fiber intake or even the crude fiber intake. an increased intake of dietary fiber was correlated with a diminished relative risk of colorectal cancer. 2001 8:05 PM CONSUMPTION OF DIETARY FIBER IN FRANCE (1950–1981) Table 7.64 77.10 Average Daily Consumption of Fruits and Vegetables (Grams per Capita) Calculated from Annual Consumption Year Potatoes Dried Pulse Other Vegetables Total Fresh Fruits Citrus Fruits 1970–1971 1975–1976 1980–1981 266 222 202 — 5.54 54. .69 1.44 22.2387_ch7. Recently.5.69 63. one consisting of women suffering from constipation and the other of controls. We did not find any difference in the dietary fiber intake between two groups of women.5. among other dietary factors.82 29.3 and 3.93 1.55 37.9 601 Potatoes Dried Pulse Other Vegetables Total Fresh Fruits Citrus Fruits 4897 4790 3970 — 113 73 6632 6257 6168 2819 2890 2779 856 998 1015 Average Annual Consumption of Fruits and Vegetables (Kilograms per Capita) (Eurostat’s Data6) Year Potatoes Dried Pulse Other Vegetables Total Fresh Fruits Citrus Fruits 97 81 74 — 2. May 6.4 and 12.1 1.29 40.45 64.38 DIETARY FIBER INTAKE AND MORBIDITY Statistics on this problem are very rare in France.10 Therefore.81 45.5.8 Total Human Consumption of Fruits and Vegetables (100 t/Year) (Eurostat’s Data 6) Year 1970–1971 1975–1976 1980–1981 Table 7.7 3.46 40. Macquart-Moulin et al.4 found that.79 1.5.5 62.47 2.50 14. the lack of dietary fiber in alimentation cannot be considered the only cause of constipation.7 g/day) and the cereal fiber intake (3.11 Average Annual At-Home Consumption of Fruits and Vegetables (Kilograms per Capita) (INSEE’s Data 19805) Total population Agricultural population Nonagricultural population Population of the greater Paris area Potatoes Dried Pulse Other Vegetables Metropolitan Fresh Fruits Bananas and Citrus Fruits 55.5_fm Page 601 Sunday.4 129 118 115 56 55 52 17 19 19 Before 1970 1975–1976 1980–1981 Table 7.8 353 323 315 153 151 142 47 52 52 Table 7.39 40.6 g/day) were virtually the same. 5 0.9 5 0.5–2 97 0.8 133 7.50 17.2 2 52 1 Citrus Fruits White-Collar Workers Table 7.41 28.42 21.8 133 7.8 222 12.9 175 2.70 52.5.55 13.5 11 0.13 21. 3RD EDITION .83 2.7 68 1.77 Self-Employed Professionals 3.1 14 0.97 36.03 1.85 Unemployed 602 Table 7.3 9.8 110 0.7–2.2 Rice 2 160 3.24 73.0 22.8 0.8 Bread ? 19 ? 15 ? 15 ? Noodles 3.5–2 140 0.13 Farm Owners 50.8 13 0.68 1.71 20.8 150 3 Potatoes 15 6 0.63 Blue-Collar Workers Bananas 2 35 0.6 Total 69.42 27.75–3 Fruits 42.01 60.70 35.42 2.29 34.5_fm Page 602 Sunday.57.12 Farm Laborers 48.77 83.85–24.54 54.2 Vegetables 64.85 Upper-Level Executives Average Consumption of Fruits and Vegetables by Occupation Group (Kilograms per Capita) (INSEE’s Data 19805) 16.88 60.35–20.4 11 0.5–3 200 4–6 230 4.6–6.5–2 145 0.14 Average Daily Fiber Intake by Occupation Groups (INSEE’s Data) Calculated from Southgate’s Analyses % of total that is dietary fiber (g/day) Dietary fiber (g/day) Cereals in Flour Equivalent 29. May 6.2 190 3.35 58.45 57.12 1.5–9 Vegetables 44.6 2 200 4 Potatoes 1. 2001 8:05 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.3 60 1.5–3 300 4.18 24.55–2.12 2387_ch7.4 Wheat Flour 2 5.24 23.68 40.8 200 7.74 1.78 % of total that is dietary fiber Farm owners (g/day) Dietary fiber Unemployed (g/day) Dietary fiber General average (g/day) Dietary fiber 5.6 Dried Pulse 1.71 Medium-Level Executives 0.24 58.20 33.34 1.5.13 Average Daily Fiber Intake (Eurostat’s Data6) Potatoes Dried pulse Vegetables Fresh fruits Bananas and citrus fruits Table 7.24 0.6 19.65–24.6–5.2 Fruits 0.7–26.1 Total 17.12 42.75 4 0.5. . Les fruits et 1égumes frais dans la restauration collective et communale. Biol.F. 1981. Mercier. Biol.9 REFERENCES 1. Clin. Clin. Presse Med. Gastroenterol. 1977. Berthezene. 1982. F. 10. 277. Gastroenterol. 2479. Y. Fibres alimentaires et constipation. M. 1. A. 1983. 3. F. 1983. Statistiques de production agricole de la Communauté Européenne. Publiees par O.. Rapports entre fibres alimentaires et constipation.. J. Paul. Biol. Centre technique interprofessionel des fruits et légumes. Clin..2387_ch7.. S.. Meyer. T. Clin. 8. 27.. 2001 3:10 PM CONSUMPTION OF DIETARY FIBER IN FRANCE (1950–1981) 603 Figure 7. Le Quintrec. A. Vlandas. Documents. Relations alimentation—cancer en France. Evolution de l’alimentation des Français 1781−1972. 4.. M. Durbec.. .1 Evolution of dietary fiber in France. Evaluation de la ration en fibres alimentaires dans la région MidiPyréneés et en Nouvelle Calédonie.. 9. 7. Meyer. and Southgate. Biol. J. A. Gastroenterol. 2. No. 1977. Alimentation et cancer recto-colique. 7. Biol. 1... Clin. 1970 a 1980. London. Corvee. Her Majesty’s Stationery Office. 4.. A. 6. M.5_fm Page 603 Tuesday.. Y. Vol.5. A. Meyer. 10. and Allegret. Frexinos. 5..E. 1. 4. Nouv. Vol. 1043. and Meyer. G.. Consommation et lieux d’achat des produits alimentaires en 1980. D. INSEE (Ser.. Gastroenterol. 1. EUROSTAT. Macquart-Moulin. F. 1978.C. 739. and Le Quintrec. T.. McCance and Widdowson’s: The Composition of Foods. D. 174A (summary). 1980. F. 971. P. P.. and Southgate. J. 1980. 99). May 8. 9. Gastroenterol.. May 6. 2001 8:05 PM .5_fm Page 604 Sunday.2387_ch7. 00+$1. and a clear drop in carbohydrate consumption (Table 7. Systematic sampling was performed on the resident population in the city and each of its suburbs. in the period from 1980 to 1984. considered a representative sample of the city of Trento in northern Italy: 1697 people. with total quantity evaluation for a 7-day period. carried out sample monitoring and registration5 of family food consumption in many zones of Italy (180 towns and cities.4) The INN. * Unpublished information from an epidemiological cross-sectional study by M. Research involved about 10.4) has shown a further increase in consumption of animal-origin products and a reduction in those of plant origin. has dropped to 1964−1966 values.6 Fiber Consumption in Italy Ottavio Bosello.6. distributed over 9 regions).8 SUBJECTS AND METHODS The Trento Study (Table 7. Research by the Istituto Nazionale della Nutrizione (National Institute of Nutrition: INN) for the period 1980 to 1984 (Table 7.2387_ch7. Trento Hospital) on a randomly selected sample of the population of Trento (northeast Italy) in 1989. on the other hand.4 Italian INN Study (Table 7.2) This research. between 25 and 69 years of age. has also involved a decrease in dietary fiber consumption and is more accentuated in northern Italy and less so in the south. Fabio Armellini. Nutritional and fiber values were calculated using the food composition tables published by the INN.6_fm Page 605 Sunday. was then checked and discussed with the subject by a dietitian when the diary was picked up. May 6. Food consumption was monitored by simultaneous recording of the food consumed during meals. Miori (Departmentof Internal Medicine. previously sent to the subject’s residence. Caloric intake.50 © 2001 by CRC Press LLC 605 .6. a doubling of animal-origin proteins and animal and plant fats.3 The diary.1).6. 0-8493-2387-8/01/$0. going from the Mediterranean model2 to a Western diet. and Mauro Zamboni INTRODUCTION Epidemiological and alimentary studies on the population of Italy from 1954 to 19781 have shown a substantial increase in daily energy intake. This gradual change in Italy’s traditional eating habits. 2001 8:06 PM CHAPTER 7.6.* carried out in 1989. 809 males and 888 females.000 families selected by random sampling of the civic lists of voters. These results show that Italian fiber intake from potatoes is low (0.0 49.7) This study* was performed in 1989 on 11 members of the vegetarian community in Verona (northeastern Italy).3 65.3 Data were subsequently checked by a dietitian. Comparisons were made on a one-to-one basis with a group of subjects of equal age. thus avoiding the need to make corrections for waste. The table also includes both total and percentage values for daily energy intake from each of the three main nutrients. using British food composition tables.9 33.6 The weekly eating habits of 30 family units were studied each month for 3 years. Nutritional and dietary fiber values were calculated using the INN tables. Zamboni about dietary fiber consumption in lacto-ovovegetarians and omnivores living in Verona (northeast Italy) in 1989. Nutritional parameters were obtained using a diary with simultaneous registration of foods consumed during meals.6 26. Fiber intake is indicated. Research in Italy involved three regions: Friuli Venezia Giulia in northeastern Italy.1 Energy MJ/day Total 9.6.4 CONSUMPTION PATTERNS Table 7.4 97.7 52.8 48.1 58. for 7 days.7 on the food intakes as measured.6. There is a tendency for greater fiber consumption in older age groups.5 48. research was performed in Europe between 1963 and 1965. May 6. Nutritional and dietary fiber values were calculated using the INN tables.4 to 14.6 54.3) For the European Economic Community (EEC) Dietary Survey. employing specifically trained dietitians. Other European countries show a totally opposite picture.8 * Unpublished information from a case-control study by M.8 70.4 11.9 13.2 gives a few results from the Trento study.8 g per person per day) and that from grains is high (13.7 g per person per day).4 European Economic Community Survey (Table 7.2 89. Fiber intake was calculated.5 428.7 to 3. 3RD EDITION Table 7.6. 17 ± 8 g/day in the youngest).4 114. as part of studies to determine environmental radioactive contamination levels.6.8 24. measured by quantity. 17 ± 8 g/day in the youngest) and for females (20 ± 9 g/day in the oldest. Campania in south-central Italy.2387_ch7.1 37.3 Data were collected by a mixed weight and inventory method.3 gives total and food-group fiber consumption measured during the EEC study.1 Changes in Food Consumption from 1954 to 1978 in Italy1 1954–1956 1964–1966 1976–1978 Proteins g/day Plant Animal Total 45. separately for males and females and for three age groups. both for males (21 ± 9 g/day in the oldest age group.6. and Basilicata in southern Italy. Each family weighed the food after preparation but before cooking.2 Carbohydrates g/day Total 389.9 451. 2001 8:06 PM 606 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and sociocultural level. Table 7. weight. Verona Vegetarian Study (Table 7. .6 88.6_fm Page 606 Sunday.4 Lipids g/day Plant Animal Total 32. body mass index. degree of physical activity. Sex Number of Subjects Mean (± SD) Daily Energy. Miori (Department of Internal Medicine.2 2387_ch7.0 ± 2. Trento Hospital) on a randomly selected sample of the population of Trento (northeast Italy) in 1989.3 9. in 1989a Age (Years) Table 7.1 9.256 172 222 25−39 40−54 55−69 Females a 227 204 177 25−39 40−54 55−69 Males 8.8 8.6. Nutrient (as % Daily Energy).6_fm Page 607 Sunday. 2001 8:06 PM FIBER CONSUMPTION IN ITALY 607 .6 7.5 ± 2. and Dietary Fiber Consumption in an Urban Northern Italian Population by Sex and Age.3 ± 3.8 9.2 ± 2.7 ± 2.6 Energy MJ/day 45 ± 10 44 ± 10 45 ± 10 48 ± 10 45 ± 10 44 ± 11 Carbohydrates (% Daily Energy) 39 ± 9 38 ± 10 38 ± 9 35 ± 9 34 ± 9 35 ± 10 Lipids (% Daily Energy) 15 ± 4 16 ± 4 16 ± 3 16 ± 4 16 ± 3 16 ± 3 Proteins (% Daily Energy) 17 ± 8 17 ± 8 20 ± 9 17 ± 8 20 ± 9 21 ± 9 Dietary Fiber (g/day) Unpublished information from an epidemiological cross-sectional study by M.9 ± 2. May 6. 3 3.1 ± 0. in Three Italian Regions. A look at four geographic areas shows moderate variations (Table 7.9 ± 0. The vegetarians. For animal fats.09 Energy (MJ/day) 11. Table 7.6.1 7. Plant protein consumption compared with the national average (38.6.04 Protein (g/day) Animal Plant Total 59.3 ± 0.6.8 Table 7. 1963–19656 Region Friuli Venezia Giulia (northeast) Campania (central) Basilicata (south) Cereals Vegetables Fruits and Nuts Potatoes Total 14. range from +9% in northeastern Italy to −8% in southern Italy.5 g/day) ranges from +7% in southern Italy to −17% in northwestern Italy.5 5.8 ± 0.1 ± 0. 3RD EDITION Dietary Fiber Intake from Food (g per person per day). May 6. Table 7. 1980−19845 Fiber (g/day) 20.3 ± 2.07 97.9 3. Nutrients.9 26.6. whereas plant fat consumption compared with the national average (54.1 ± 0.5 ± 0.9 23.6 ± 0.6 g/day). showed about twice the fiber intake (31.7 gives results from a study on vegetarians (lacto-ovo-vegetarians) and nonvegetarians (omnivores) living in a northern Italian town.6 illustrates this same territorial variability phenomenon in terms of consumptions expressed as food preferences.3 g/day). consumption varies from +21% in northeastern Italy to −16% in southern Italy.12 38. and this is true for fiber intake.7 24.16 108.4 gives mean values for fiber consumption and total and percentage energy intake per nutrient for Italians as indicated by the Italian INN study.7 g/day) given in Table 7.2 g/day). The latter demonstrate eating habits practically identical to the population at large.7 1.4 Mean (± SE) Fiber. as well. The fiber intake of these northern Italian vegetarians is greater even than that measured in 1980 to 1984 in the southern regions of Italy (22.7 5.13 54.3 4.8 ± 0.6.7 13.6_fm Page 608 Sunday.1 g/day) ranges from +8% in southern Italy to −13% in northwestern Italy.9 g/day) show greater consumption in central (+4%) and southern ( +6%) Italy than in the north (−12 and −13 %).5) from one area to another of Italy. . on the other hand.6.6.4. these same consumption values vary much more when they are subdivided into plant and animal protein and lipid values.21 Alcohol (g/day) 17. see Table 7.2387_ch7.6.54 89.5.6 0.5 gives interesting data regarding energy and nutrients.4 13. 2001 8:06 PM 608 Table 7.9 Table 7. Whereas average total protein and lipid consumption values show only small percentage differences with respect to totals between geographic areas. when compared with the national average (53.4 g/day) compared to the nonvegetarians (17.9 ± 4.7 ± 0. Data on fiber consumption compared with the national average (20. compared to the national average (59.8 1.02 Table 7.22 Carbohydrates (g/day) Available Soluble 325.15 Lipids (g/day) Animal Plant Total 53.3 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Percentage differences in animal protein consumption.6 ± 0.6. and Energy Intake in Italy. 5 609 Percentage Intake of Fiber. Starting in 1970. pasta. g/day. CONCLUSIONS Fiber consumption in Italy is higher in central than in northern regions of Italy and is even greater in the south. increased physical activity.2387_ch7.7) South 106 (22. in comparison with other Italian regions.11. 2001 8:06 PM FIBER CONSUMPTION IN ITALY Table 7. it is practically impossible to evaluate mortality trends for cardiovascular diseases and ischemic heart diseases. These southern and central areas seem to still be fairly well tied to the Mediterranean alimentary model. a continuous increase in fat consumption and cigarette smoking has been counterbalanced by better control of high blood pressure.10 On the other hand. This is because many risk factors have undergone simultaneous changes. May 6.2) Center 104 (21. fruits. Compared with the Whole of Italy (intake = 100). foods that are especially rich in fiber and complex carbohydrates. have a protective effect with regard to stomach cancer.6. where the source of proteins and lipids is predominantly from plants and where there is an evident preference for the consumption of vegetables.12 FIBER CONSUMPTION AND CANCER IN ITALY One study13 performed on 206 women with endometrial cancer and on 206 controls showed that the risk for contracting this disease is higher for subjects consuming the more fat-rich diets that are also poor in green vegetables. for example. Given in Parentheses9 Northwest 88 (18.6_fm Page 609 Sunday. there have been studies in Italy that show the beneficial effects that fiber has on the main coronary risk factors.4) Northeast 87 (18. Actual Fiber Intake. 1980−1984. on the other hand. The risk for contracting endometrial cancer appears inversely correlated to beta-carotene and fiber intake. and pizza. the influence . and Energy in Four Geographic Areas. Nutrients. and improved health care for patients with coronary disease. in particular. such as whole wheat bread and whole wheat pasta.2) Protein Animal Plant Total 106 87 99 109 89 101 104 100 102 92 108 98 Lipids Animal Plant Total 120 83 102 121 95 108 103 100 102 84 107 95 Carbohydrates Available Soluble 95 98 99 109 99 100 103 97 Alcohol 97 121 105 88 Energy 98 103 101 99 Fiber FIBER CONSUMPTION AND CARDIOVASCULAR DISEASE IN ITALY In Italy. bread. The same authors14 have performed a similar case-control study on stomach cancer. and whole foods. In northern Italy. 6 Food Preferences in Four Italian Geographical Areas. and pasta. Rice Potatoes Giblets Low-fat cheese Plant fats and oils Sugar and honey Soft drinks Beer Cerealsa Citrus fruit Dry fruit and nuts Beef Pork Salted meats Poultry Nonfat milk Low-fat milk Yogurt Medium-fat cheese Animal fats Pasta sauces Soup preparations Biscuits and cakes Wine Spirits . 1980−19849 Northwest Northeast Center South More Frequently Consumed Food Rice Fresh fruit Salted meats Giblets Yogurt Medium-fat cheese High-fat cheese Biscuits and cakes Artificial sweeteners Dietetic products Soft drinks Cerealsa Potatoes Dry fruit Fresh pork Poultry Milk Plant fats and oils Pasta sauces Sugar Soft drinks Wine Beer Spirits Tomatoes Vegetables Citrus fruit Beef Preserved fish Animal fats Soup preparations Pizza Pasta Dry legumes Meat Fresh fish Low-fat cheese Less Frequently Consumed Food Pasta Dry legumes Vegetables Fresh fish Preserved fish Whole milk a Bread and pizza Tomatoes for sauces Fresh fruit Meat Low-fat cheese Eggs Olive oil Snacks Excluding bread. May 6.2387_ch7. but healthier. Table 7. A recent study that shows that. who are presumably still tied to older. 3RD EDITION of the Western diet is much stronger and fiber intake is low.6. 2001 8:06 PM 610 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.6_fm Page 610 Sunday. eating habits gives further confirmation of this change toward the so-called Western diet which is relatively low in fiber. pizza. in the same northern-Italian urban environment. there is lower fiber consumption in the younger age groups compared to the elderly. Lombardo.. Napoli.. G. Boca Raton. O. L... 24.0 ± 37. De Carli. 88. Glucose tolerance and blood lipids in bran fed patients with impaired glucose tolerance. A.7 1.. 12.. Fisheries and Food. A. Prev. Ledermann..6_fm Page 611 Sunday. Rivellese.7 Carbohydrates (g/day) Total Fiber Energy (MJ/day) a ± ± ± ± 5. F.. 1950–1981. 6. 3.. F... Situazione ed evoluzione dei consumi alimentari in Italia. Bingham. A. La Vecchia.. F. S.5 ± ± ± ± 4. 484.E. G. G.6 28. Tabelle di composizione degli alimenti. Recent trends in coronary heart disease and other cardiovascular diseases in Italy.. 1981.. A. CRC Press.1 ± 9. Litho Delta. G.. 13. Idelson.3 4. Micciolo.9 ± 3. Nutrients. 1989. A. E. Her Majesty’s Stationery Office.8 60. Spiller. and Lacourly. and Liguori. 1986.3 3.7 611 Mean (± SE) Fiber. The Mediterranean Diet in Health and Disease. Fidanza.1 4.. E..2 ± 4. 5.. Farchi. A. Eds. Mistura. Cialfa. A. A case-control study of diet and gastric cancer in northern Italy. .. 1969.2 69. 1980. I.6 7.7 48. Cancer. La Vecchia. and Mariani Costantini. in Nutrizione Umana... and Pasquali. Cialfa. et al.9 9. and Energy Intake by Lacto-ovo-vegetarians and Omnivores Living in a Northern Italian Town.6 378.3 9.6 ± 1. May 6. 128. Saba. 57. 1981. 1989a Lacto-Ovo-Vegetarians Omnivores Protein (g/day) Animal Plant Total 17.. Nutrition and diet in the etiology of endometrial cancer. C. C..4 320. Gardiner. Annual reports of the National Food Survey Committee. 10. Istituto Nazionale della Nutrizione. A. MacDonald.8 ± 0. Napoli. Cardiology.8 42. A.. Diabetes Care. 40. and Scuro. De Carli. and Franceschi.. 72. F. F... 1991. 3. G. Ed.0 29. A dietary survey of 11 areas of the E.. 1987. and Liguori.4218F.8 13. in order to determine levels of radioactive contamination.. and Cialfa. Zamboni about dietary fiber consumption in lacto-ovo-vegetarians and omnivores living in Verona (Northeast Italy) in 1989. D’Avanzo. R. A..1 ± 42. 12(1).6 ± 6. A. 1990. 8. A national food survey... 19(4). Fasoli. 1953−1983.5 9. Indagine nazionale sui consumi alimentari delle famiglie 1980−84. M. A.3 ± 2. Reduction of risk factors for atherosclerosis in diabetic patients treated with high-fiber diet.. R. 53. 7. Human Nutrition Reviews (ILSI). Tecniche di rilevamento delle abitudini e dei consumi alimentari. S. Household Food Consumption and Expenditure.3 23.6. Med. Negri..9 ± 4.3 10. A. Menotti. Riccardi. Turrini.8 Lipids (g/day) Monounsaturated Polyunsaturated Saturated Total 32. G.1 ± 5.. J. R. 46. 11. and Lintas.C.5 75. Euratom. Idelson... FL. Ed. 2001 8:06 PM FIBER CONSUMPTION IN ITALY Table 7. 1248. Cancer. in CRC Handbook of Dietary Fiber in Human Nutrition. Int. Armellini. Van Nostrand Reinhold. 1985. edizione 1988. Milano..4 17. Fidanza. E.7 ± 7... 2. Fidanza. M. REFERENCES 1. 1983.. 9. E. S. B.3 77. Cresta. E.. M. Eds. Giacco. Ministry of Agriculture. Ostuzzi. New York. 14.. La Rivista della Societa Italiana di Scienza dell’ Alimentazione. Food balance sheets and other methodologies: a critical overview.. EUR..4 31. London. C.2 Unpublished information from a case-control study by M.2387_ch7. Turrini. 4. Alcuni importanti risultati. Capocaccia. Bosello. and Gentile. G... in Nutrizione Umana. Spiller. Patterns of Dietary Fiber Consumption in Humans.0 32. A. 1991. 1986. May 6.2387_ch7. 2001 8:06 PM .6_fm Page 612 Sunday. May 6. 2001 6:07 PM APPENDIX Tables of Dietary Fiber and Associated Substances Content in Food .2387_Section App_fm Page 613 Sunday. 2001 6:07 PM .2387_Section App_fm Page 614 Sunday. May 6. or (3) calculation of recipes or formulations.00+$1. 0-8493-2387-8/01/$0.1 Dietary Fiber Values for Common Foods Sally F. Where only two of the three fiber values were known for a food. Janet Pettit. the third was calculated using the following formula: Total Dietary Fiber = Insoluble Fiber + Soluble Fiber.2387_chzAppendix_T1_fm Page 615 Sunday. Himes The dietary fiber table developed by the Nutrition Coordinating Center (NCC) contains values compiled from scientific literature. the three fiber values were obtained from different references and. Estimated values were derived from (1) a different form of the same food (e. For some foods. 2001 8:09 PM Appendix — Table A. Schakel..g. (2) a similar food. therefore. raw to cooked). and estimated data.135 manufacturer’s information. the sum of the insoluble and soluble fractions may not equal the value for total dietary fiber.50 © 2001 by CRC Press LLC 615 . May 6. the USDA Nutrient Database for Standard Reference. and John H. 35 0.31 0.20 1.57 3.00 31.08 1. oatmeal Bread.99 1.00 26. pumpernickel Bagel.00 25.48 1.Dietary Fiber Values for Common Foods 1 Medium 1 Medium 1 Medium 1 Medium 1 Medium 1 Medium 1 Medium 1T 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Slice 1 Medium 1 Square Bread and Other Related Products Amount a 57. reduced-calorie and high-fiber Bread.96 0.59 0.54 0.75 45.00 57.40 0.00 26.00 84.00 6. cracked wheat Bread.40 0.54 1.25 0.55 0.09 1.45 0. raisin Bread.62 0.61 1.30 Insoluble Fiber (g) 0.26 0.00 37.54 0.1 2387_chzAppendix_T1_fm Page 616 Sunday. cheese Bread.22 0.47 0.26 0. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.00 26.42 0. oat bran Bagel.26 0.23 0.76 1.40 1. wheat.00 25.00 28.00 27. egg Bread.31 2.12 1.73 0.07 0.12 1.00 28.78 0.00 Weight (g) 1. cracked wheat. Boston brown Bread.67 0.27 1.36 0.00 25.03 3.76 1.22 0. oat bran Bread. egg Bagel.65 0. rye (light or dark) Bread.51 0.00 25.45 Soluble Fiber (g) 616 Bagel.86 0.34 0. wheat Bread.48 1. whole wheat Biscuit.58 0.11 0.03 3.94 0.57 0. multigrain or granola. sourdough Bread. reduced-calorie and high-fiber Bread.76 0.41 0.00 28.73 1.32 0.31 2. reduced-calorie and high-fiber Bread.67 0. Italian Bread.15 1. whole wheat Breadsticks.58 0. baking powder or buttermilk Bread crumbs.78 0.42 0. white.15 0.62 0.51 0.00 26.32 1.35 0.00 57.31 1. plain Bread.79 3.00 57.25 1.34 0. rye Bagel.60 1. bread type Coffee cake.06 2. reduced-calorie and high-fiber Bread.00 70.10 26.78 0.92 0.86 0.34 0. white Bread.16 2.00 26.00 57.18 0.61 2.49 0.67 0. 3RD EDITION . focaccia Bread. multigrain or granola Bread.00 29.00 28.35 20.00 26.59 0.92 0. hovis Bread.73 0. cinnamon swirl Bread.69 0.92 Total Dietary Fiber (g) 0.00 57. quick-bread or crumb type Table A.51 1. French Bread.00 23. pumpernickel Bread. white flour Bagel.30 0.58 1.71 0. May 6.25 0.36 0.50 25.49 0.67 1.51 0.35 32.80 1.27 1. plain Croissant Croutons Danish pastry Doughnut. raised or yeast French toast Muffins.45 1.00 60.31 0. hamburger.08 2387_chzAppendix_T1_fm Page 617 Sunday.60 0.42 0. white Pita.08 3. whole wheat Muffins.59 0.83 0.93 0.08 2.23 1.00 36.00 45.56 1.15 1.26 0. English muffin.00 65.27 0.16 1.56 0.32 0.64 0.23 1.41 1.24 0.00 45.38 1.86 0.97 1.00 0.42 1. whole wheat Pita.00 57. rye 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Square Piece Medium Medium C Medium Medium Medium Slice Medium Medium Medium Medium Each Each Medium Each Each Each Each Medium Medium Each Medium Each Medium Medium Medium Medium Medium Regular Medium Medium Medium Medium Medium Medium Bread and Other Related Products 39. blueberry Muffins.64 1.52 0.46 0. white Rolls.21 1.00 43.33 0.49 0.39 0.01 0.00 36.00 28. white flour. Kaiser Rolls. white Muffins. oatmeal Rolls.Coffee cake. white flour. corn Muffins.52 0. wheat Rolls.10 0. yeast type Cornbread Crepe. hard Rolls.23 0.03 0.39 0.78 1.00 60.77 2.77 0.90 57.10 0.00 57.32 0.33 0.66 0.62 0. crescent (refrigerated dough) Rolls.00 43. 2001 8:09 PM APPENDIX — TABLE A. hot dog Rolls.00 50.15 1. oat bran or oatmeal Pancake.00 59.43 0.45 0.27 0.00 30.49 0.39 0.46 0.00 50. carrot Muffins.22 0.90 36.83 1.17 0.00 57.48 0.49 1.07 0.00 57.00 47. May 6.00 38.34 0.45 1. cake Doughnut. white flour.16 2.19 0. prepared from a mix Pancake. hamburger.00 38. whole wheat Popover Rolls.58 0.30 0. frozen Pancake.45 0.53 0.47 1.32 0. French or Vienna Rolls. oat bran Rolls.09 2.99 3.29 0.00 36.09 0. bran Muffins.54 0.00 1.00 66. hamburger.89 2.53 1.36 0. whole wheat Rolls.00 45.31 0.63 0.76 0.00 36.00 44.64 0.00 43.01 1.08 1.33 0.00 54.24 0. English muffin.39 0. cracked wheat Rolls.89 1.59 0.25 0.83 0.00 57.21 0. pumpernickel Rolls.52 1.37 0.15 1.50 2.66 0. multigrain Rolls.23 1.95 1.23 0.54 4.65 0.00 36.00 57.25 0.1 617 .53 1.45 2.00 43.62 0.00 62.00 36.17 0.38 0. prepared from a recipe Pancake.27 0.38 0. 46 0.Cornmeal Cracked wheat Cream of rice Cream of wheat.49 1.13 2.00 39.48 0.48 3.00 242. regular cooking Rolled wheat Waffles.67 3.85 1.44 0.00 43.73 0.00 Weight (g) 2. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.27 0.05 1.59 0.00 242.99 0.42 0.14 0.64 2.85 0.21 2.48 0. instant cooking. instant cooking Creamed wheat.78 1.94 0.00 26.74 1.30 0.09 1.37 0. home recipe Amount a 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C C C C C C C C C C C C C C C C Cereals — Cooked Medium Medium Medium Medium Medium Medium Medium Regular Regular Each Each Each – 4 in.57 0.51 0. plain Oat bran Oatmeal.65 1.1 (Continued) 2387_chzAppendix_T1_fm Page 618 Sunday.64 0.97 1.88 0. flavored Mix ’N Eat Cream of Wheat.35 1.30 52.00 36.02 0.00 95.00 36.00 234.98 3.60 0. diameter 1 1 1 1 1 1 1 1 1 1 1 1 Bread and Other Related Products Dietary Fiber Values for Common Foods 240. wheat Rolls.01 1.00 239. frozen Table A.47 1.39 0.53 0.35 0.00 241.00 45.45 2.05 0.58 0.00 242.43 0.81 0. diameter 1 Each – 7 in.00 42.38 0.39 0.17 0.51 1.87 0.00 244. white flour.82 5. quick cooking Oatmeal. 3RD EDITION .84 3.63 1.53 0. corn Tortilla. plain Oatmeal. multigrain flavored Creamed wheat. regular cooking Grits Mix ’N Eat Cream of Wheat.82 1.62 5.06 0. submarine or hoagie Rolls.43 0.25 0. white Rolls.93 1.76 0.06 1.42 0. white flour.08 2.21 0. May 6.39 0.00 241.36 2.08 1.02 1.00 234.00 219. whole wheat Scone Sweet roll Taco shell. hard Toaster pastry or Pop-Tart Tortilla.68 2.06 2.38 0. instant cooking.22 Insoluble Fiber (g) 0. sourdough Rolls.00 241.13 3. flavored Oatmeal. quick cooking Creamed wheat.28 0.00 13.53 Total Dietary Fiber (g) 2.32 3.40 0.98 3.00 241.32 Soluble Fiber (g) 618 Rolls.00 251. white flour Waffles.00 57.00 234.00 94.29 1.14 3.00 36.00 234.12 0.85 0.00 1.64 6.10 0. 09 52.35 6.72 1.04 3.65 0.67 36.34 1.00 30.12 2387_chzAppendix_T1_fm Page 619 Sunday.00 1.00 34.00 40.39 4.93 0.75 3.03 2.00 40.29 1.33 0.08 0.02 0.29 2.37 1.40 0.35 0.00 40.00 32.00 41.00 33.00 55.00 30.00 40.95 26.04 1.21 0.65 5.59 1.00 40.14 2.69 0.00 40.25 0.12 0.00 36.17 2.40 18.24 0.41 1.00 30.00 40.15 1.06 0.37 1.00 65.78 20.16 1.67 1.23 1.67 6.44 28.42 3.64 0.00 40.07 0.41 2.38 9. May 6.14 0.44 1.35 0.20 24.24 2.40 0.25 0.00 55.51 1.77 0.69 6. 2001 8:09 PM APPENDIX — TABLE A.69 2.04 1.10 0.70 1.00 33.60 0.52 0. Sweetened (Arrowhead Mills) Bite Size Shredded Wheat (Arrowhead Mills) Body Buddies (General Mills) Booberry (General Mills) Bran Cereal with Apples & Cinnamon (Health Valley) Bran Flakes (Arrowhead Mills) Bran Flakes (Malt-O-Meal) Bran Flakes (Post) Cap’n Crunch (Quaker) Cap’n Crunch’s Crunch Berries (Quaker) Cap’n Crunch’s Peanut Butter Crunch (Quaker) Cheerios (General Mills) Cinnamon Life (Quaker) Cinnamon Toast Crunch (General Mills) Cocoa Blasts (Quaker) Cocoa Comets (Malt-O-Meal) Cocoa Frosted Flakes K-sentials (Kellogg’s) Cocoa Krispies K-sentials (Kellogg’s) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C Cereals — Ready-to-Eat 87.77 0.70 0.07 0.32 0.47 5.59 7.00 50.1 619 .100% Bran (Nabisco) All-Bran (Kellogg’s) All-Bran Bran Buds (Kellogg’s) All-Bran with Extra Fiber (Kellogg’s) Alpha-Bits (Post) Amaranth Flakes (Arrowhead Mills) Apple & Cinnamon Toasty O’s (Malt-O-Meal) Apple Cinnamon Cheerios (General Mills) Apple Jacks K-sentials (Kellogg’s) Apple Zaps (Quaker) Basic 4 (General Mills) Berry Berry Kix (General Mills) Berry Colossal Crunch (Malt-O-Meal) Betty Crocker Cinnamon Streusel (General Mills) Betty Crocker Dutch Apple (General Mills) Bite Size Shredded Wheat.95 2.08 0.63 1.21 1.33 41.00 49.10 0.03 35.07 1.68 0.92 0.34 2.39 11.61 0.15 0.00 34.09 62.24 0.19 1.00 40.00 30.25 6.33 29.47 1.12 1.04 4.73 3.00 55.57 1.30 0.54 22.09 0.21 0.00 90.11 1.33 24.01 6.64 0.87 0.03 0.00 1. 97 2.00 40. Mills) Erewhon Poppets (U.87 6.40 0.12 Insoluble Fiber (g) 0.51 1. Mills) Erewhon Crispy Brown Rice (U.99 0.00 30.13 0.01 0.00 38.83 1.1 (Continued) 2387_chzAppendix_T1_fm Page 620 Sunday.13 0.43 0.S.77 0.75 0.00 30.00 20. Mills) Erewhon Banana-O’s (U.00 24.77 4.10 6.12 0.17 2.41 0.S.11 0.15 0.64 1.06 0. Mills) Erewhon Wheat Flakes (U.99 0.S.09 0. No Salt Added (U.17 0.05 1.51 2.00 2.28 0.00 65.00 40.16 Total Dietary Fiber (g) 0.17 0. 3RD EDITION .S.00 0. May 6.99 Soluble Fiber (g) 620 Cocoa Pebbles (Post) Cocoa Puffs (General Mills) Colossal Crunch (Malt-O-Meal) Complete Oat Bran Flakes (Kellogg’s) Complete Wheat Bran Flakes (Kellogg’s) Corn Bursts (Malt-O-Meal) Corn Chex (General Mills) Corn Flakes (Arrowhead Mills) Corn Flakes (Kellogg’s) Corn Flakes (Malt-O-Meal) Corn Pops K-sentials (Kellogg’s) Count Chocula (General Mills) Country Corn Flakes (General Mills) Cracklin’ Oat Bran (Kellogg’s) Crispix (Kellogg’s) Crispy Puffs (Arrowhead Mills) Crispy Rice (Malt-O-Meal) Crispy Wheaties ’N Raisins (General Mills) Crunchy Bran (Quaker) Double Chex (General Mills) Erewhon Apple Stroodles (U.00 36.S.67 31.S.00 35.18 0.31 0.19 1.00 36.57 3.18 1.29 0.S.77 1.04 0.00 30. Mills) Erewhon Corn Flakes (U. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.00 31.15 0. Mills) Erewhon Crispy Brown Rice.00 53.45 7.18 6. Mills) Erewhon Aztec (U.78 1.S.00 52. Mills) Erewhon Raisin Bran (U.25 5.00 30.35 0.59 0.S.00 4.83 6.26 5.96 1.10 1.00 Weight (g) 0.14 1.74 0.89 0.00 28.48 0.45 0.34 1.15 1.23 3.S.63 0.29 8.00 36.14 0.00 30.66 3.97 1.28 0.64 0.03 1.54 0.29 6. Mills) Erewhon Fruit’n Wheat (U.01 6.00 1.44 0.58 0.33 29.00 40.11 0.10 0.04 0.10 1. Mills) Erewhon Galaxy Grahams (U.S.04 0.78 0.44 0.43 2.36 1.00 30.04 0.34 0.33 50.00 70.S.00 40.Amount a 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C Cereals — Ready-to-Eat Dietary Fiber Values for Common Foods 38.52 0.35 0. Mills) Erewhon Kamut Flakes (U.20 1.66 0.41 6.78 0.00 30.67 37.49 0.00 30.06 0.01 5.67 30.83 0. Mills) Fat-Free 10 Bran O’s.48 5.82 1.93 5.00 55. Apple Cinnamon (Health Valley) Table A.98 30. 59 0.13 0.00 30.48 0.24 1.02 0.81 1.29 0.00 41.88 1.01 0.00 30.00 55.10 0.81 0.00 36.55 0.95 0.1 621 .33 28.56 5.38 1.63 28.45 0. Almond (Health Valley) Fat-Free Honey Clusters & Flakes.27 1.Fat-Free Honey Clusters & Flakes.00 36.01 0.00 40.83 1.99 0.50 0. May 6.97 1.39 1.72 0.09 0.96 4.59 0.59 2387_chzAppendix_T1_fm Page 621 Sunday.61 1.67 58.72 6.33 40.55 0.71 0.76 0.63 5.84 0.82 1.33 41. Honey Crunch (Health Valley) Fiber One (General Mills) Frankenberry (General Mills) French Toast Crunch (General Mills) Froot Loops K-sentials (Kellogg’s) Frosted Cheerios (General Mills) Frosted Flakers (Quaker) Frosted Flakes (Malt-O-Meal) Frosted Flakes K-sentials (Kellogg’s) Frosted Oats (Quaker) Frosted Shredded Wheat Bite Size (Nabisco) Frosted Toasty O’s (Malt-O-Meal) Fruit & Fiber — Dates.84 1.31 0.45 0. Apple Cinnamon (Health Valley) Fat-Free Honey Clusters & Flakes.54 0.41 0.47 12.63 5.53 0.36 1.95 1.94 1.46 0.00 0.33 60.08 1.42 0.88 4.08 1.00 31.18 1.60 0.00 41.33 41. 2001 8:09 PM APPENDIX — TABLE A.04 0.28 3.40 0.88 0.00 44.00 100.46 0.00 40.57 3.00 55.33 54.01 0.44 0.00 1.14 9.52 0.18 0.62 3.69 0.01 2.66 1.00 1.38 1. Raisins & Walnuts (Post) Fruitangy Oh’s (Quaker) Fruity Pebbles (Post) Golden Crisp (Post) Golden Flax (Health Valley) Golden Grahams (General Mills) Golden Puffs (Malt-O-Meal) Grainfield’s Brown Rice (Weetabix) Grainfield’s Corn Flakes (Weetabix) Grainfield’s Crispy Rice (Weetabix) Grainfield’s Raisin Bran (Weetabix) Grape-Nuts (Post) Grape-Nuts Flakes (Post) Healthy Choice Almond Crunch with Raisins (Kellogg’s) Healthy Choice Golden Multi-Grain Flakes (Kellogg’s) Healthy Choice Toasted Brown Sugar Squares (Kellogg’s) Honey & Nut Toasty O’s (Malt-O-Meal) Honey Bunches of Oats Honey Roasted (Post) Honey Bunches of Oats with Almonds (Post) Honey Crunch Corn Flakes (Kellogg’s) Honey Frosted Wheaties (General Mills) Honey Graham Oh’s (Quaker) Honey Nut Cheerios (General Mills) Honey Nut Clusters (General Mills) Honey Nut Oats (Quaker) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C 41.00 36.76 4.00 40.29 0.00 4.33 1.00 41.94 1.41 0.81 0.00 40.15 0.30 1.19 9.89 4.56 3.36 8.06 4.00 30.00 5.00 38.24 3.00 30.76 26.40 1.11 3.33 40.00 41.00 30.16 0.08 0.00 30.31 4.72 0.34 0.00 52.94 1.07 1.00 30.78 1.42 5.77 1.20 0.83 0.19 0.10 0.46 3.15 1.32 0.00 116.94 1.48 0.69 0.00 30.76 4.04 1.09 0.22 3.00 36.00 28.49 0.00 32. 74 2.48 0.43 0.63 0.93 0.02 5.51 134.46 30.17 0.45 0.81 2.00 53.05 0.00 73.86 1.20 2.81 4.00 1.60 3.77 1.92 Insoluble Fiber (g) 0.00 55.67 82.00 Weight (g) 4.28 3.67 66.22 0.01 0.39 0.44 0.50 0.00 20.37 42.96 70.41 1.22 3.60 4.1 (Continued) 2387_chzAppendix_T1_fm Page 622 Sunday.23 1.98 0.73 5.10 0.20 6.56 0.11 0.73 1.92 2.86 1.87 4.67 59.00 56.65 0.85 0.00 30.01 5.Amount a 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C Cereals — Ready-to-Eat Dietary Fiber Values for Common Foods 52.21 1.Raisin (Kellogg’s) Mini-Wheats .00 46.22 0.Apple Cinnamon (Kellogg’s) Mini-Wheats .55 0.61 1.46 80.56 4.63 0.40 4.64 5.88 5.05 0.75 0.60 3.79 0.00 35.88 6.59 4.00 70.00 40. May 6.14 1.Blueberry (Kellogg’s) Mini-Wheats .86 0.00 55.07 Soluble Fiber (g) 622 Honey Nut Shredded Wheat .33 72.Strawberry (Kellogg’s) Morning Traditions Banana Nut Crunch (Post) Morning Traditions Blueberry Morning (Post) Morning Traditions Cranberry Almond Crunch (Post) Morning Traditions Great Grains Crunchy Pecan (Post) Morning Traditions Great Grains Raisins.00 44.49 5.99 0.45 2.67 0.41 0. Dates & Pecans (Post) Mueslix — Apple & Almond Crunch (Kellogg’s) Mueslix — Raisin & Almond Crunch with Dates (Kellogg’s) Multi Grain Cheerios Plus (General Mills) Multi Grain Flakes (Arrowhead Mills) Multi-Bran Chex (General Mills) Nature O’s (Arrowhead Mills) Table A.95 0.46 1.99 3.67 30.67 22.20 2.00 55.02 5.00 6.66 0.00 Total Dietary Fiber (g) 3.24 1.86 0. 3RD EDITION .45 6.00 2.44 1.39 5.44 0.00 32.35 1.00 24.00 21.76 60.00 79.60 1.61 0.56 0.56 0.00 6.88 5.52 2.04 1.99 0.00 29. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.25 5.Bite Size (Nabisco) Honeycomb (Post) Just Right Fruit & Nut (Kellogg’s) Just Right with Crunchy Nuggets (Kellogg’s) Kaboom (General Mills) Kamut Flakes (Arrowhead Mills) King Vitaman (Quaker) Kix (General Mills) Kretschmer Honey Crunch Wheat Germ (Quaker) Life (Quaker) Lucky Charms (General Mills) Maple Corns (Arrowhead Mills) Marshmallow Alpha-Bits (Post) Marshmallow Blasted Froot Loops K-sentials (Kellogg’s) Marshmallow Mateys (Malt-O-Meal) Marshmallow Safari (Quaker) Mini-Wheats .Frosted Original (Kellogg’s) Mini-Wheats .56 0.03 1.00 30.07 0.40 32.39 6.39 11.34 5.61 14.00 6.Frosted Bite Size (Kellogg’s) Mini-Wheats .39 2.64 5. 45 0.78 2.00 14.00 22.58 0.67 0.88 0.78 3.99 8.00 15.85 4.20 1.29 3.02 1.21 0.43 0.42 1.06 0.00 32.56 4.63 1.33 37.74 1.87 4.30 2.00 56.34 0.Nutri-Grain — Almond Raisin (Kellogg’s) Nutri-Grain — Golden Wheat (Kellogg’s) Oat Bran Flakes (Arrowhead Mills) Oat Bran Ready-To-Eat (Quaker) Oatmeal Crisp — Almond (General Mills) Oatmeal Crisp — Apple Cinnamon (General Mills) Oatmeal Crisp — Raisin (General Mills) Oatmeal Squares — Cinnamon (Quaker) Oatmeal Squares (Quaker) Oreo O’s (Post) Organic Amaranth Flakes (Health Valley) Organic Blue Corn Flakes (Health Valley) Organic Bran with Raisins (Health Valley) Organic Crisp Brown Rice (Health Valley) Organic Fiber 7 Flakes (Health Valley) Organic Healthy Fiber Flakes (Health Valley) Organic Oat Bran Flakes (Health Valley) Organic Oat Bran Flakes with Raisins (Health Valley) Organic Oat Bran O’s (Health Valley) Organic Raisin Bran Flakes (Health Valley) Post Toasties (Post) Product 19 (Kellogg’s) Puffed Corn (Arrowhead Mills) Puffed Corn (Health Valley) Puffed Kamut (Arrowhead Mills) Puffed Millet (Arrowhead Mills) Puffed Rice (Arrowhead Mills) Puffed Rice (Malt-O-Meal) Puffed Rice (Quaker) Puffed Wheat (Arrowhead Mills) Puffed Wheat (Malt-O-Meal) Puffed Wheat (Quaker) Raisin Bran (Arrowhead Mills) Raisin Bran (Kellogg’s) Raisin Bran (Malt-O-Meal) Raisin Bran (Post) Raisin Nut Bran (General Mills) Real Oat Bran — Almond Crunch (Health Valley) Reese’s Peanut Butter Puffs (General Mills) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C 39.21 0.09 0.12 0.24 2.00 16.22 0.41 1.34 4.11 0.97 1.00 30.00 15.85 5.89 0.03 4.68 1.32 0.60 55.07 5.93 4.46 4.04 7.72 1.06 8.52 2.03 2.00 96.33 66.60 7.87 2.52 4.77 2.00 54.41 5.10 5.04 3.92 5.35 1.31 0.26 0.00 12.00 34.57 1.00 23.00 25.00 55.79 4.33 37.53 0. 2001 8:09 PM APPENDIX — TABLE A.22 0.21 1.00 36.33 41.1 623 . May 6.30 0.33 37.00 61.26 4.39 0.71 0.00 60.57 0.01 1.66 0.95 1.99 0.01 2.61 2.00 59.14 5.14 5.00 37.33 44.27 0.33 0.56 4.00 61.54 2.50 9.37 1.13 8.36 0.54 0.29 7.00 55.49 2.55 0.06 9.02 0.53 4.67 0.11 1.00 55.67 37.26 1.56 5.26 0.22 0.96 1.18 2.02 1.12 2387_chzAppendix_T1_fm Page 623 Sunday.40 0.00 45.35 0.09 7.15 7.65 4.10 2.52 2.33 37.45 3.67 42.00 28.01 0.09 0.24 5.00 25.00 40.04 0.12 0.68 0.17 7.35 0.62 0.00 3.73 4.56 5.13 2.20 40. 31 0.50 64.00 31.96 6. Mills) Wafflers — Vanilla Nut (U.09 0. May 6.Amount a 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C C C BIS BIS C C C C C C C C C C C C C C C C C C C C C C C C BIS C C C Cereals — Ready-to-Eat Dietary Fiber Values for Common Foods 31.35 1.54 4.18 0.1 (Continued) 2387_chzAppendix_T1_fm Page 624 Sunday.00 0.37 0.98 44.05 1.10 Total Dietary Fiber (g) 0.22 4.65 2.00 36.00 40. Mills) Weetabix (Weetabix) Wheat Bran (Arrowhead Mills) Wheat Chex (General Mills) Wheaties (General Mills) Table A.00 0.67 30.75 0.38 0.54 2. Mills) Waffle Crisp (Post) Wafflers — Cinnamon (U.00 49.78 0.31 0.83 0.75 0.01 1.26 2.06 1.00 Weight (g) 0.89 1.S.S.99 0.00 0.40 2. Mills) Wafflers — Maple (U.73 3.88 1.00 30.00 44.98 44.17 0.00 41.00 66.83 1.80 6.03 0.33 30.62 0.56 5.25 2.98 17.00 30.00 0.34 0.21 0.89 1. Mills) Skinner’s Raisin Bran (U.00 55.58 1.96 3.03 0.45 1.00 50.25 0.31 0.70 7.S.06 4.33 2.18 2.00 30. 3RD EDITION .S.82 0.72 Soluble Fiber (g) 624 Rice Chex (General Mills) Rice Krispies K-sentials (Kellogg’s) Rice Krispies Treats (Kellogg’s) Shredded Wheat (Nabisco) Shredded Wheat (Quaker) Shredded Wheat Spoon Size (Nabisco) Shredded Wheat’n Bran (Nabisco) Skinner’s Corn Flakes (U.00 26.34 2.00 49.28 1.29 2.29 0.98 44.40 0.52 0.43 1.98 27.00 30.98 7.98 55.31 0.00 3.52 0.12 0.00 3. Mills) Wafflers — Original (U.58 1.94 2.01 1.58 0.15 0.S.87 1.69 25.92 1.89 1. Mills) Smacks (Kellogg’s) Smart Start (Kellogg’s) S’Mores Grahams (General Mills) Special K (Kellogg’s) Spelt Flakes (Arrowhead Mills) Sweet Crunch (Quaker) Toasted Oatmeal — Honey Nut (Quaker) Toasted Oatmeal — Original (Quaker) Toasty O’s (Malt-O-Meal) Tootie Fruities (Malt-O-Meal) Total (General Mills) Total Corn Flakes (General Mills) Total Raisin Bran (General Mills) Trix (General Mills) Uncle Sam (U.83 1.11 0.08 0.40 40.02 5.89 1.31 0.51 55.72 4.21 1.00 40.28 0.38 1.32 2.12 0.39 0.89 1.07 1.00 27.00 22.S. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.72 9.20 41.84 0.36 0.39 5.00 23.58 1.12 1.00 21.58 1.00 47.38 Insoluble Fiber (g) 0.97 2.S. 72 4.00 82.73 4.14 138.33 2.11 5.13 4.01 9.67 3.40 7.06 6.39 9.09 98.33 7.76 2.00 89.41 2.S.24 2.67 3.21 3.46 90.00 41.88 4.14 150.92 6.50 10.64 2.64 11.64 3.46 82.19 1.70 1.94 5.80 2387_chzAppendix_T1_fm Page 625 Sunday.63 12.33 41.48 2.21 3.49 2.46 114.14 147. Mills) Familia Swiss Muesli — No Added Sugar Familia Swiss Muesli — Original Recipe Fat-Free Granola O’s — Almond (Health Valley) Fat-Free Granola O’s — Apple Cinnamon (Health Valley) Fat-Free Granola O’s — Honey Crunch (Health Valley) Heartland Granola — Original (Pet) Heartland Granola — Raisin (Pet) Low Fat Alpen (Weetabix) Low Fat Granola (Kellogg’s) Low Fat Granola with Raisins (Kellogg’s) Mountain House Granola with Blueberries (Oregon Freeze Dry) Nature Valley Low Fat Fruit Granola (General Mills) Sun Country Granola with Almonds (Quaker) Sun Country Granola with Raisins & Dates (Quaker) Vita Crunch Granola — Almond (Organic Milling Company) Vita Crunch Granola — Light & Crunchy 7 Grain (Organic Milling Company) Vita Crunch Granola — Raisin (Organic Milling Company) Vita Crunch Granola — Tropical (Organic Milling Company) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C Granola and Muesli 110.72 4.28 4.46 82.02 5. May 6.00 4. Mills) Erewhon New England Style Granola — Maple (U.88 12.S.33 41.19 2.46 82.09 5. Mills) Erewhon Spiced Apple Granola (U.1 625 . 2001 8:09 PM APPENDIX — TABLE A.49 3.00 3.08 12.00 90.21 3.00 110.94 5.90 3. Mills) Erewhon New England Style Granola — Honey Almond (U.20 5.15 141.18 1. Honey & Raisins (Grist Mill) 100% Natural Low Fat Granola with Raisins (Quaker) 98% Fat-Free Granola — Date Almond (Health Valley) 98% Fat-Free Granola — Raisin Cinnamon (Health Valley) 98% Fat-Free Granola — Tropical (Health Valley) Alpen (Weetabix) Erewhon #9 Granola with Bran (U.14 11.64 2.30 2.15 114.00 73.S.01 3.00 120.09 144.01 3.73 1.16 7.56 2.20 5.93 7.96 110.84 7.33 73.74 7.32 5.00 128. Mills) Erewhon New England Style Granola — Date Nut (U.67 8.20 5.74 7.93 1.19 3.00 120.80 1.04 3.66 1.93 7.S.43 5.S.92 5.52 4.78 5.24 3.20 6.94 8.22 4.48 2.20 2.69 1.01 2.84 8.28 5.14 144.100% Natural Granola — Oats & Honey (Grist Mill) 100% Natural Granola — Oats.93 4.33 128.00 82.35 5.72 5.S.21 6.48 5.33 73.19 1.63 7. Mills) Erewhon Sunflower Crunch Granola (U.74 7.07 15.33 73.69 1.77 7.64 3.88 5. 04 0.00 44.02 0.03 0.09 0.83 0.00 1.00 11.44 3.00 10.78 70.01 0.47 0.08 0.00 11.35 84.30 0.30 0.92 0.05 0.25 0.15 0.60 1.72 0.14 1.50 0.80 4.27 0.41 0.00 2.08 0.35 5.05 1.04 0.1 (Continued) 2387_chzAppendix_T1_fm Page 626 Sunday.00 3.00 1.37 0.35 15.85 0.65 1.36 0. whole wheat Wheat Whole wheat Zwieback Table A.01 0.11 0.04 0.55 1.13 1.00 4.35 28.25 0. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.02 0. peanut butter Onion flavored rings Popcorn cake Round Round Wafer Each Each Each Each Rectangle Each Wafer Wafer Each Each Square Each Each 1 1 1 1 1 1 1 1 1 1 1 1 1 1 oz Bar C C oz oz C oz Bar Bar Bar Bar oz Each Snacks and Chips 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Crackers Amount a 28.01 0.Dietary Fiber Values for Common Foods Bagel chip Cereal bar Cereal party mix.77 0.03 0. white Saltine or soda.17 0.37 0.05 0.08 Total Dietary Fiber (g) 0.50 0.07 0.66 0.00 27.12 0.00 1.39 0.35 10. plain Matzo or matzoh.00 5.42 0.08 28.38 1.05 1.13 0.50 28.00 3.35 37.50 28.00 0. seasoned Saltine or soda.05 0. plain bar (no coating).03 0. plain bar (no coating).35 28.00 Weight (g) 1. egg Matzo or matzoh.80 28.99 1.16 0.06 0.09 0.39 0. May 6.00 7.07 0.07 0.16 0.05 Soluble Fiber (g) 626 Butter Cheese Crispbread — rye Flatbread Matzo or matzoh. flavors other than peanut butter Granola bars. puffs or twists Corn chip Corn nuts Fried pork rinds Fruit bar Granola bars. homemade Cheese balls.36 0.01 0.80 23.12 0.23 0.75 0.80 0.75 0.11 0.03 0.00 3.36 5.85 0.04 0.06 0.35 28.00 0.37 0. plain Rye wafer.09 0.18 0.64 0.03 Insoluble Fiber (g) 0.27 0.37 1. 3RD EDITION .37 0.89 0. chocolate-coated Granola bars. commercial Cereal party mix.50 23.00 3. whole wheat Melba toast Oyster Rye wafer.35 5. hot-air-popped Popcorn.22 1. dry Couscous. dry Bran. microwave popped from package Potato chip Potato sticks Pretzels.50 12.99 10.95 99.00 60. peanut.61 1.08 0.00 164.00 11.04 0.48 24.00 28. not “buttered” Popcorn.33 9.89 25.00 120.62 0.04 0.66 1.96 14.09 1. and Other Grains 1 1 1 1 1 1 1 1 1 1 1 195.50 10.22 0.68 8.00 162. barley bran Flour.00 173.1 Amaranth.26 0.31 4.68 20. cooked Couscous.13 0.69 95.35 9.18 1.00 58.38 1.48 1.71 21.55 21.00 157.00 114.80 0.00 76.20 11.35 29.22 1.09 4.91 0.70 0. defatted Flour.03 0. wheat (unprocessed) Buckwheat groats.78 24. arrowroot Flour.61 2. cooked Barley.67 9.21 2.17 1. dry Flour. dry Bran. chickpea Flour. cheese-flavored Popcorn. dry Bran.60 6. dry Barley. cake or pastry Flour.31 1. buckwheat Flour.19 1.83 1.00 94.00 1.87 10.94 9.01 APPENDIX — TABLE A.37 3. home-popped.02 0.86 2. baking mix Flour.84 6. home-popped.92 2. barley Flour.87 7.51 1.59 1.60 7.04 0.00 200.94 10.00 128.52 0. caramel or sugar-coated Popcorn.00 157.77 3.00 138.00 156.99 1.35 1.13 2.13 1.09 6.94 0.36 1. canned Bulgur.56 0.53 31.48 1.94 2. dry Bulgur. 2001 8:09 PM 627 . rice.80 9.80 16.00 148.21 1.00 28.26 4.88 14.28 0. dry Bulgur.00 60. oat bran.00 182.85 6.38 2. Rice.C C C C C C oz C oz Each oz 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C C C C C C C C C C C C C C C C C C C C C C C C C C Pasta.17 0.34 0.00 140. May 6. hard type Rice cake Taco or tortilla chips 2387_chzAppendix_T1_fm Page 627 Sunday.33 0.29 2.62 5. dry Cornmeal.75 0.67 24.65 0.13 1. commercially popped (prepopped).40 148.20 64.12 0.89 6. popped in fat Popcorn.00 113.65 4.95 7.70 8.10 11.97 11.00 28. dry Bran.47 14.00 135.87 1.48 9.35 36. peanut.64 8.59 22. barley malt Flour.00 2. corn Flour.04 0.00 12.31 4. corn.87 6.00 92.16 6.80 1.00 118.71 4.61 0.21 0.00 137.07 6.40 61.00 8. low-fat Popcorn.00 35. home-cooked Corn grits. cooked Pasta.00 140. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.75 1.89 17.62 0.98 18.00 192.00 195.00 160. noodles.76 1.Amount a 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C Pasta.95 10. whole wheat Hominy.82 3.17 8.19 0.98 3. noodles.63 26.34 1.00 100. white. cooked Sorghum Table A. dry Pasta.00 102.34 4.23 0. low-fat Flour. parboiled (converted).00 165. instant (precooked).00 175.82 3. dry Rice.44 7.00 6. white.22 3.92 9.00 120. soy. rye Flour.00 158. full-fat Flour.00 140. chow fun rice. and Other Grains Dietary Fiber Values for Common Foods 160. white. cooked Pasta.00 2.88 3.00 160.79 14.00 140.84 0.37 14. 3RD EDITION .12 1.52 0.48 2.36 3.23 0.47 Insoluble Fiber (g) 5.80 0.76 4. brown. Chinese. whole grain Flour. white.92 1.70 0.54 3. crisp type Pasta. noodles. cooked Pasta.12 0. cellophane. dry Millet.1 (Continued) 2387_chzAppendix_T1_fm Page 628 Sunday.00 Weight (g) 9. cooked Millet. triticale.57 13.00 168.08 1.00 84.07 0. whole wheat.88 1.43 15. noodles.72 3.12 0. cooked Oatmeal.34 1. potato Flour.32 6.27 3.66 0. soy.59 1. cooked Quinoa. chow mein.00 158.00 140.23 0.00 200. cooked Pasta.00 130.00 176.51 0. Rice.84 3.91 9. macaroni noodles. cooked Pasta.28 0.29 1.06 8. cooked Rice.48 1.70 3. cooked Quinoa.48 8.00 176.00 45.82 5. cooked Pasta.00 240.67 3.06 Soluble Fiber (g) 628 Flour.51 0. defatted Flour.44 3.41 0.78 0. white all-purpose Flour. spaghetti noodles.00 158. canned Kasha. soft type.04 0. cooked Pasta.70 3. brown Flour.96 1.48 2.84 1. rice.60 2.00 158.32 10. cooked Pasta. noodles. soy. cooked Rice.26 1.18 0. egg. rice. cooked Rice.13 5. white Flour.64 4.29 17. noodles.00 125.36 0.00 170.96 1.98 7.37 0.46 18.06 8. whole wheat.53 Total Dietary Fiber (g) 4. white.48 1.00 165.50 8.37 12.63 5. cooked Pasta. noodles. macaroni noodles.08 8.00 160.00 88.63 4.16 1.19 2. regular cooking. whole wheat.26 4. spaghetti noodles.97 1.48 0.18 1. rice. May 6.71 0.00 81.03 3.60 1.00 160. canned.00 173. sprouted Wild rice. fresh Apricot.21 8. California type Banana.19 0.32 1. water pack Apricot. fresh Figs.00 145.77 1.12 0. with skin.1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Fruit 1 1 1 1 1 1 1 1 Medium Medium C C C C C C C C C C Each Medium C C C C C C C C C C C C C C C C C C C C C C C 138.77 16.72 4. fresh.00 199. dried.12 178.61 4.54 2. canned.01 2.00 258. canned. gluten Wheat.00 8.19 2.00 152.00 1. 2001 8:09 PM 629 .04 10.70 8. cooked.68 1.71 1.01 2.62 2.42 13.43 1. heavy syrup pack Figs.13 11. unsweetened Apple.37 26.94 5. dried. fresh Cherries.00 128.62 2. cooked 2387_chzAppendix_T1_fm Page 629 Sunday.00 187.00 237.15 5. black skin.00 253. with skin Apple.28 2.70 24.44 11. heavy syrup pack Apricot.30 1.01 13.91 1.15 5.15 3.00 164. fresh Cranberries.55 5. fresh Cantaloupe.04 0.00 259.00 190.90 1.48 3.11 0.00 413.43 3.99 6.99 120.72 14. without skin Apple. maraschino cherries Cranberries. canned.00 130.40 4.48 0.00 108.19 3. unsweetened Avocado.14 1.00 120.00 108. spelt Wheat.58 12.00 3.00 113.1 Apple.00 161. fresh Cherries.01 1.97 0.35 10. canned.00 155.96 APPENDIX — TABLE A.89 4.56 2.58 5.00 244.70 2.00 198. dried (Craisins) Dates Elderberries.85 5.00 255.95 0.00 95.66 3.00 135.83 2.73 16.84 5.01 2.00 145.02 3.47 14.96 1. fresh Figs.92 3.72 3.22 21. canned.01 1.00 243. heavy syrup pack Cherries.33 1. with skin. fresh Carambola (starfruit).93 7.76 1. uncooked Fruit cocktail. dry Teff.00 160. unsweetened Apricot. baked or scalloped.00 118.44 2. uncooked Apricot.65 2. baked or scalloped.91 3.62 1.00 145.42 7.04 1. canned.06 2. dried.32 2.00 144.24 2. fresh Blueberries.21 2.56 0.12 6. uncooked Applesauce or stewed apples.63 3.68 1.33 3.43 4.97 0.93 3.14 0.66 1.96 0.32 1.44 0. cracked whole wheat Wheat. water pack Tapioca.00 86.98 3.28 2.43 0.00 250. fresh or ripe Blackberries.73 2.45 3. May 6. sweetened Applesauce or stewed apples.98 2.33 12.83 7.09 2.79 1. dried. germ Wheat.00 177. fresh. sweetened Apple.94 4. dry Wheat.37 1.47 0.93 2. 47 2.00 150. fresh Mandarin orange. cooked.27 0.20 1.46 1.78 2. white Grapefruit. fresh Guava.98 2.00 170. pink or red Grapefruit.74 1. fresh Table A.00 256. fresh Pear.41 13. fresh Nectarine. water pack Pear.61 6.19 1. fresh.84 0.Dietary Fiber Values for Common Foods 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Fruit C C C C Medium Medium C C C C C Medium Each Medium C C C Each Medium Medium C Each Medium C C C C Medium C C C C Each Amount a 237.90 4.61 0.22 2.18 3.30 0.36 1. fresh Passion fruit.30 2.53 0.76 3. May 6.12 1.12 6.97 3.29 5.60 8.00 58. water pack Grapefruit.64 1. unsweetened Pear. fresh Mandarin orange.00 76.58 1. canned. canned.96 7.87 1.37 2. fresh Gooseberries. uncooked Pear.79 Soluble Fiber (g) 630 Fruit cocktail.00 166.00 266. juice pack Fruit cocktail. canned.1 (Continued) 2387_chzAppendix_T1_fm Page 630 Sunday.51 0. fresh Lemon.00 244. canned.98 0.00 98.96 3.45 6.48 0.31 1. fresh Orange.18 0.48 1.75 0.00 255. fresh Loganberries.34 0. dried.26 13.32 6.20 4. fresh Papaya. dried.00 19.00 258. cooked.02 2.18 1.02 1.04 3.00 Weight (g) 2.48 6.97 8.99 5.05 Total Dietary Fiber (g) 0.00 160.83 1.00 248. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 3RD EDITION .00 252.00 140.10 4.47 5.52 1.00 244.53 1.05 2. fresh Kiwi Kumquat.51 0.02 2.73 2.00 207.00 136.78 1.00 256. canned.62 7. canned Grapefruit.00 244.26 Insoluble Fiber (g) 1.98 3. canned. canned.82 1.14 2.00 195.04 7.80 0.49 1. syrup pack Mango. fresh common Honeydew melon.03 0.73 1.26 2. heavy syrup pack Pear. canned.95 0.68 1.00 180.98 1. dried.40 8.17 3.00 147.00 262.00 160.34 0.10 7. fresh Peach.50 16.82 0.00 254.82 0.00 18.42 0.01 0. heavy syrup pack Gooseberries. water pack Peach.50 1. fresh Peach.00 131.37 0.73 1.16 0.25 1. syrup pack Grapes.25 1.00 252. heavy syrup pack Peach.00 168. dried.08 3.42 1.36 6. fresh.91 1.00 165. unsweetened Persimmon.61 0.35 2.83 3. uncooked Peach.82 2. canned. 75 0.97 0.50 0.00 258.21 0.15 0.20 4.49 1. unsweetened Raisins.16 9.50 0.00 250.27 0.00 250.17 0.58 0.15 0.77 1. fresh Strawberries. light syrup pack Plantains.00 247.31 4. sweetened Rhubarb.48 0.53 0.28 0.00 244.00 253.37 6.00 252.20 8.75 5. heavy syrup pack Pomegranate Prune.25 0.86 1.89 0.00 286. dried. cooked from fresh. frozen.37 0.56 0.00 155.50 2.00 0.00 154.79 0.22 1.51 1.52 0.75 0.84 1.00 249.47 2.97 1.94 1.77 1. uncooked Prune. fresh Plum.36 11.86 0.00 248.00 170.20 1.43 0.37 0.25 1.00 250.99 2.57 0. canned.28 0.25 1.15 1.32 1.49 1.60 2.1 631 . fresh Pineapple.73 0.08 0. fresh Strawberries.07 16.25 0.00 242.00 236. sweetened Watermelon. cooked.50 9.00 240. canned.05 0.36 13.00 251.07 0.02 3.25 0. canned.44 1.00 144.70 1.92 12.00 249.00 123.00 249.46 4.24 0. boiled or baked Plum.80 3. dried.00 165.00 253.18 0.00 170. sweetened Rhubarb.20 0.61 12.57 2387_chzAppendix_T1_fm Page 631 Sunday.49 7.40 0.00 247.15 4.00 155. unsweetened Sapodilla.75 3.45 1.69 0. fresh Raspberries.93 0. 2001 8:09 PM APPENDIX — TABLE A. uncooked Raspberries.00 245.45 3.50 1. juice pack Pineapple.48 0.24 0.52 1.25 1.17 6.54 2. ripe.25 0.86 1.00 243.Apple juice Apricot nectar Black currant juice Carrot juice Cranberry juice Cranapple juice Grape juice Grapefruit juice Lemon juice Orange juice Papaya juice Passion fruit juice Peach nectar Pineapple juice Prune juice Tomato juice Vegetable juice Pineapple. frozen.00 250.00 240.00 154.00 256.94 3.35 1.00 4. fresh 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Juices 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C C C C C C C C C C C C C C C C C C C C C C C Each C C C C C C C Each C C Slice 248.25 1.05 0. cooked from fresh. May 6.01 2.25 0.49 0.70 0.10 0.50 1.86 1.98 0.00 255.25 0.00 0.15 0.51 2. 65 11.41 4.00 180.00 177.87 1.00 262.72 4. cooked from fresh Beets.76 4. baked beans with pork in tomato sauce Beans.75 0.00 135.00 135.27 2.71 13. green or string.46 3.00 20.18 4.94 2. cooked from frozen Beans. raw Asparagus.44 13.05 1.53 0. wax or yellow.02 7.75 0.10 7.02 Soluble Fiber (g) 2.40 13.83 3. soybeans. tepary.75 4. wax or yellow.06 4. green or string. pinto. cooked from dried Beans.00 171.55 5. cooked from fresh Beans. cooked from frozen Beans. garbanzo.00 252.36 1.14 13.00 182. cooked from dried Beans.00 135. cooked Arugula.15 14. green or string.29 1.16 9.62 1.96 2.27 1.99 2.05 Total Dietary Fiber (g) 6. raw Table A.00 164. soybeans. cooked Beans.83 0.37 2.31 Insoluble Fiber (g) 632 Artichoke. drained Beet greens.18 2. roasted Beans.62 1.36 10. lima.14 7.32 10. cooked Broccoflower (green cauliflower).89 1.88 3. cooked Beans.81 1.Amount a 1C 1C 1C 1C 1C 1T 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C Vegetables and Legumes Dietary Fiber Values for Common Foods 168.90 1.00 253. canned Beans.07 0.60 0.75 4. canned.29 1. kidney.76 12.89 1.32 2.05 2. green or immature. refried beans. fresh Beans.20 7.66 6.65 253. canned.64 3.44 0.00 172.86 11.20 5. cooked from frozen Beans. cooked Beans.56 30.32 7.12 2.75 3. cooked from dried Beans.66 6. May 6.00 242.69 1.47 0.40 2. canned. canned.00 144.00 172. northern.94 9.97 0.36 0. drained Beans.36 3.00 Weight (g) 9.00 125.00 170.57 3.00 170.00 131.29 0.39 2. cooked Asparagus. navy.00 125.28 8.00 254. cooked from dried Beans.42 1.62 1.12 2. bayo.84 2.00 38.33 13.00 188.00 180.16 12. lima. baked beans.00 172.94 5. drained Bamboo shoots. regular globe.62 5.04 1. baked beans with pork in brown sugar Beans. soybeans. cooked from dried Beans. drained Basil.05 1. raw Beet greens. cooked from dried Beans.1 (Continued) 2387_chzAppendix_T1_fm Page 632 Sunday.68 4.44 2.49 9. cooked from dried Beans.00 170.75 2.52 5. wax or yellow.56 11.06 5.00 2. drained Beets.08 8.10 12. cooked from fresh Beans.00 135.16 16. 3RD EDITION . vegetarian Beans.00 64. canned. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.00 177. 18 0.26 2387_chzAppendix_T1_fm Page 633 Sunday.00 132.00 150.33 1.95 2.46 0.00 119.21 0. raw Collards.53 1.00 156.71 1.32 1.52 6.67 0.60 100.04 1.73 0.15 4. raw Collards.02 0.05 0.45 1.73 0. raw Carrots.32 1.06 2.94 3. cooked Cabbage.00 170.88 0. Pak-choi.00 89.21 1. May 6.62 0.83 0.64 1. cooked from fresh Carrots. cooked Chinese vegetables.05 3.00 180.00 29.66 0. canned.02 0.30 5. Pe-tsai. cooked Coriander leaf.38 1.61 0.00 125.00 175.36 3.49 0. raw Cabbage.07 1.37 4. whole kernel Corn. raw without peel Dill weed.21 0.35 2.44 2.01 2.60 2.88 0.28 0.01 1.82 1. cooked from fresh. cooked Cabbage.00 119.69 2. Pak-choi.08 0.Broccoflower (green cauliflower). cooked Chard. fresh Coriander leaf.00 70.90 0.61 0.77 0.00 2.00 155. canned.00 89.00 100.00 120.00 156.65 3.23 0.00 146. cooked Chinese cabbage.64 4.02 2. cooked from fresh Broccoli. drained Cucumber. fresh Eggplant. cooked Cauliflower.18 2.54 0. cooked from frozen Celeriac or celery root. green.40 3. canned. red.78 3.65 2. cooked Carrots. raw Cauliflower.30 5. raw 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1T 1C 1C 1T 1 TS 1 Medium 1C 1C 1C 1C 1T 1C 1C 82.46 1.00 184.30 3.56 99.00 36.00 155. red.88 0.32 1.34 2. cooked Broccoli. raw with peel Cucumber. 2001 8:09 PM APPENDIX — TABLE A.00 190. raw Cabbage.50 0.00 104.49 0.86 1.72 2.83 0.48 1.78 3.69 0.86 2.03 0.04 2.00 3.90 0.00 146.49 3.00 0.47 0.95 0.00 164.00 88.96 2.76 3.26 0.34 2.00 76. Pe-tsai.80 3.00 150.88 0.90 1. cooked Endive (curly).82 2.00 150.92 1.09 0. dried Corn. cooked from frozen.70 1.50 3.90 1.65 2.71 2.1 633 . cooked Celery.68 1.00 2.00 164.75 0. raw Celery.42 0. raw Broccoli.53 2. cooked from frozen Carrots.60 0.32 0.33 0.00 125.96 1. raw Chinese cabbage.53 0.70 2.98 1. green. cooked Chinese cabbage. drained Cassava (yuca).04 0.87 1.50 3. cooked from frozen Brussels sprouts.05 0.00 0.00 110.76 2. cooked from fresh Cauliflower. cob Corn. drained Chives.62 0.51 2.68 5.72 0.78 0. raw Chinese cabbage.19 1. 34 3. green beans.00 182. yellow.92 3. or red.00 135. cooked Kohlrabi. cooked from dried Lettuce.80 7.74 0.75 156.01 Total Dietary Fiber (g) 2387_chzAppendix_T1_fm Page 634 Sunday. drained Mushrooms.00 184.00 2.00 172. cowpeas.12 6. canned. yellow.14 0.00 196. yellow or green.00 156.26 3.00 160.00 160.48 16. cooked Peas.94 0. split peas.31 2. lima beans. fresh Parsnip. cooked Fennel bulb.09 3.00 182.19 4.76 2.45 0. cooked Peas.56 0.96 0.37 1.Amount a 130.85 1. peas. cooked Lentils. cooked Kale.82 3.25 1. green peas. canned.12 9. or red.00 160.70 3. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.48 0. raw Garlic.81 15.74 5.31 0.20 0.11 1. iceberg Lettuce. and carrots).00 87.48 0.80 6. raw Onion.24 8.86 1.12 3.02 2. cooked from frozen Mixed vegetables (corn.00 160.01 Insoluble Fiber (g) 634 5.00 146. raw Kohlrabi. fresh Ginger root. canned Jicama or yambean. cooked Parsley. snow peas or edible pea pods.00 165.88 2.10 3.97 11.16 1. peas.24 6.1 (Continued) 4.00 165.00 210.15 2. canned. drained Peas.18 8.66 0.00 170. cooked from frozen Mixed vegetables (peas and carrots). green peas.46 1. 3RD EDITION .00 160.02 1.12 2.30 1.10 0. lima beans. cooked from dried Table A.68 1. romaine or cos Mixed vegetables (corn. drained Okra. and carrots).00 55.37 2.70 1.25 4.60 4. cooked from fresh Mushrooms.11 1.62 4.27 3.00 156. raw Mushrooms. white.00 135.50 6. cooked Peas.51 0.64 0. drained Mixed vegetables (peas and carrots).84 3.43 3.11 0.56 14. green beans.36 3.20 130.43 3. canned.95 8.89 1.59 2.05 Soluble Fiber (g) 3.07 1.17 1. white.83 96.06 1. cooked from dried Peas.13 1.38 1.77 0.00 130.84 0.46 5.19 2. May 6. cowpeas.00 3.43 3.28 0.65 6. raw Jicama or yambean.00 70.00 1C 1C 1C 1C 1C 1C 1C 1C 1C 1T 1C 1C 1C 1C 1C 1C 1C Weight (g) 1C 1C 1 TS 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C 1C Vegetables and Legumes Dietary Fiber Values for Common Foods Endive (curly).00 198.12 0.96 4.36 14. cooked from fresh Peas.70 0. cooked Onion.19 0.57 2.38 0.00 56. raw Hearts of palm. 93 1. drained Sprouts. boiled. cooked from fresh Spinach. jalapeno pepper.98 1. cooked Peppers. canned.82 0.25 1. cooked Peppers. green. hot chili.98 2.00 70. red.10 0. skin not eaten Potato.11 1.16 1.87 1. red — sweet. alfalfa Sprouts. raw Peppers.77 10.78 3.04 1.00 156.00 30.63 1.85 1.00 236.54 2. canned.00 170.00 100. hot chili.08 1. yellow or green. cooked from fresh Peppers.36 6.00 0.57 0.00 184.00 245. raw Peppers.96 3.00 125.71 0. hot chili.73 2.20 0.34 1.36 2.21 1. raw Radish.50 0.18 1.36 1.54 2.31 1.81 4.57 4.52 0.79 1.00 136.12 1. raw Scallions or spring onions.00 150. baked. canned.42 1.36 2.15 5. cooked from fresh Sprouts. canned.54 2. jalapeno pepper.00 90. canned. drained Pumpkin.00 136. drained Potato. without skin Potato.Peas.92 2387_chzAppendix_T1_fm Page 635 Sunday.68 0.00 180.85 1.00 149. raw Squash.09 1.99 6. jalapeno pepper.60 1. 2001 8:09 PM APPENDIX — TABLE A.00 156.18 1.38 0.22 1. red — sweet.94 0. drained Peppers.00 127. cooked from fresh Peppers.25 2. cooked 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C 196.28 0.00 16. with skin Potato.85 0. baked.86 3.00 184.40 6.92 3.21 2. mung bean.47 1. split peas.00 136.44 14.62 0. cooked from frozen Spinach.87 0.06 5. drained Peppers.68 2.05 1.32 4.37 0.25 2. acorn.31 0.60 5.41 4.52 3. hot chili. hot chili. canned.00 122.36 1.63 0. cooked Peppers.39 2.00 37.00 136. boiled.1 635 .23 1.53 0.00 124.63 0. raw Peppers. cooked Sauerkraut Scallions or spring onions.04 1. cooked from fresh Peppers.99 1. soybean.74 1.00 205.00 136. butternut. cooked Squash.47 0.98 0.14 0.00 240. green pepper — sweet.24 1.00 40. drained Sprouts.00 116.63 0. canned.81 5.20 0. raw Peppers.00 219. hot chili. yellow — sweet.00 180.61 3. raw Spinach.46 0.01 2.52 7.62 2.27 2.00 149. raw Peppers. green. cooked Spinach.92 0. drained Peppers. yellow — sweet.55 3.49 0. skin eaten Potato.82 1. green.60 1.38 1. raw Peppers.76 1.64 0.00 214.00 150.90 3. green pepper — sweet.90 2.35 1. canned Radicchio.77 10.59 2.00 149.51 2. mung bean.00 139.00 245. hot chili.17 0.04 2. red.87 0.13 0.00 33. raw Rutabaga.29 0. red.18 0.01 2.15 1. May 6. sun-dried Peppers.00 184. 42 10.97 3.98 2. sun-dried. canned Watercress.24 0.00 139.00 110.58 2.98 2. textured vegetable protein Breakfast strips. green. textured vegetable protein Amount a C C C C C C C C C C C C C C C C C C C C C C C C C C C C 1T 1 Strip Meat Substitutes 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Vegetables and Legumes Dietary Fiber Values for Common Foods 7.54 3.52 1.96 0. canned. hubbard.25 2.98 1.83 0.10 5.63 1. cooked Water chestnuts. cooked Squash. canned Tomato.00 200. dry pack Tomato.25 0.1 (Continued) 2387_chzAppendix_T1_fm Page 636 Sunday.44 1.00 160.58 0.38 0.00 155.12 3. summer-type (green or yellow).00 255.80 0.00 Weight (g) 0.00 8.00 180.00 144.84 2.65 5.48 6. zucchini.59 3.00 236.08 0.93 0.72 7.40 0.00 180. drained Sweet potato.88 4.00 200. vacuum-packed Tomatillo.51 1.00 250.00 132.89 1. paste Tomato.28 1.08 0.83 0.80 3.16 0.64 6.24 4.34 0. canned. cooked Squash.17 1.00 180.80 3.88 3. 3RD EDITION .18 0. yellow. canned.60 0. syrup-packed.80 1. drained Tomato. spaghetti.38 Soluble Fiber (g) 636 Squash.40 1.21 4. sun-dried. canned.00 262.00 240. raw Yams.52 6.00 196. drained Yams. cooked Yams.22 2.84 2. raw Squash. oil pack. syrup-packed. summer-type (green or yellow).56 1.00 54.63 1.44 1.38 0. raw Tomato.80 1.84 2.76 1. raw Tomato.52 2.18 0.36 2. winter-type (dark green or orange).60 Total Dietary Fiber (g) 0.13 2.60 2. raw Turnip.00 113.88 3. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.00 155.78 1. cooked Squash.00 158.22 Insoluble Fiber (g) 0.30 8.51 7.79 1. orange. zucchini.04 5.00 180.00 240. cooked Turnip greens. vacuum-packed Table A.Bacon bits.28 1. raw Squash.08 1.00 6.74 5.22 0.98 2. raw Tomato.56 0.24 1. cooked Squash.15 2.65 5. May 6.00 0.00 140.00 255.12 2.00 196.61 2.00 34. chayote. raw Tomato. cooked Sweet potato. cooked Squash.04 3. puree Tomato.00 113. cooked Sweet potato.96 3.00 6.09 0.25 2.27 2. 82 0. breakfast or brown-and-serve.29 13. soy product Hamburger/ground beef substitute — meatless Hot dogs.00 130.97 3.00 114.00 130.00 140.00 15.80 227. seeds. fresh Filberts or hazelnuts Flax seeds Trail mix (nuts. link Sausage.35 28.00 144.92 31.20 0.08 8.44 0.13 108.56 0.21 0.85 0.35 28.68 12.50 0. or wieners substitute — meatless Sausage.00 16.73 3.19 140.33 13.86 11.03 0.93 0.09 0.24 2.74 0.55 2.04 35.06 0.00 28.10 1.78 7.46 11.56 1.46 2.08 0.05 0.63 0. and dried fruit) Hickory nuts Macadamia nuts Mixed nuts with peanuts Mixed nuts without peanuts Peanuts Peanut butter Pecans Pine nuts — pignolias Pine nuts — pinyon Pistachio nuts Poppy seeds Pumpkin or squash seeds Sesame seeds Tahini (sesame butter) Canadian-style bacon.05 11. 2001 8:09 PM APPENDIX — TABLE A.92 9. breakfast or brown-and-serve.82 14.00 16.63 14.13 7.35 28.80 2.00 134.69 6. dried (shredded or flaked).55 13.80 0.Almonds Almond butter Almond paste Brazil nuts Cashews Cashew butter Chestnuts Coconut.92 10.68 1.50 1.48 10. sweetened Coconut.88 9.28 1.42 8.61 6.83 1.62 1.59 13.56 4.00 397.22 6.00 80. frankfurters.76 4.04 0. May 6.05 28.68 1.03 0.31 2.35 28.27 14.39 0.95 8.32 2.16 3.34 0.91 13. dried (shredded or flaked).07 0.46 12.00 15.43 2.00 128.74 4.38 13.35 28.92 13.05 1.08 1.23 25.00 0.73 0.06 12.03 3.18 1.73 8. from frozen Tofu (soybean curd) Vegetable burger oz oz oz oz oz oz C oz oz oz 1C 1T 1T 1C 1C 1T 1C 1C 1C 1 Medium 1C 1C 1C 1C 1C 1C 1C 1C 1T 1C 1C 1C 1C 1C 1C 1C 1T Nuts and Seeds 1 1 1 1 1 1 1 1 1 1 142.35 28.00 150.12 3.77 7.42 2387_chzAppendix_T1_fm Page 637 Sunday.21 14.00 120.94 0.00 0.1 637 .57 9.37 0.00 150.51 0. patty Tempeh (fermented soybean product) Textured vegetable protein.10 0.90 0.58 0.00 136.08 1.89 2.13 0.29 6.32 16.00 74.00 143.35 28.43 3.52 0.59 0.21 0.00 142.61 10. substitute — meatless. unsweetened Coconut.57 0.70 5.12 0.00 144.06 0.35 15. substitute — meatless.00 135.26 1.25 0.35 94. from dry Textured vegetable protein.09 5.85 11. 76 5.91 0.00 248.03 5.51 0.13 0.21 0.43 0. prepared from ready-to-serve can Bean with bacon.00 248.36 0. prepared from ready-to-serve can Clam chowder.00 238. undiluted Clam chowder.70 0.15 0.00 240.31 10. undiluted Cream of broccoli. prepared from ready-to-serve can.88 0.38 0.00 252.61 0.76 Total Dietary Fiber (g) 0.44 1.00 248.00 239.13 0.00 248.44 1. undiluted Black bean.92 0.38 2. undiluted Chicken with noodles or pasta.96 0.55 1. ham.26 2.17 0. prepared from ready-to-serve can Beef with noodles or pasta.35 0.38 1. prepared from ready-to-serve can Chicken with rice.80 Soluble Fiber (g) 638 Bean with bacon.88 0.28 5.66 0.72 0.26 0.10 0.00 C C C C C C 1 1 1 1 1 1 1C 1C 1 1 1 1 1 1 C C C C C C 252.67 0.07 1.15 0.00 C C C C C C C C C C 238.45 0.Dietary Fiber Values for Common Foods Soup 252.12 0.75 0. undiluted Sunflower seeds Sunflower butter Walnuts Table A.00 241.49 1.00 252.55 3.00 128. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.00 252. prepared from ready-to-serve can Black bean.33 0.00 120. Manhattan (tomato base).00 242.29 0. prepared from ready-to-serve can Chicken with noodles or pasta. prepared from ready-to-serve can Chicken with vegetables. undiluted Cream of onion.24 2.22 0.00 256.00 238. or pork.80 0. prepared from ready-to-serve can Clam chowder.25 0. May 6.93 0.1 (Continued) 2387_chzAppendix_T1_fm Page 638 Sunday.40 0.35 1. undiluted Cream of celery.43 3. undiluted Chicken with rice.00 16.65 0.62 0.57 1. undiluted Beef with noodles or pasta.79 3.80 3. New England (cream base).01 0.02 1. 3RD EDITION .00 252.27 0. Manhattan (tomato base).01 0. undiluted Cheese.00 248.74 0. New England (cream base).97 2.95 7.30 6.67 0. undiluted Broccoli cheese.22 2.00 240.21 0. chunky-style Chicken with noodles or pasta.71 1.30 1. undiluted Cream of mushroom.96 Insoluble Fiber (g) 0.00 248.45 1.12 1.00 252.00 Weight (g) 1 1 1 1 1 1 1 1 1 1 1C 1T 1C Nuts and Seeds Amount a 2. undiluted Cream of chicken.67 13.76 0.00 248.47 0.00 240.76 2.94 5.69 0.67 1. undiluted Chicken with vegetables. undiluted Clam chowder.25 0.38 10. undiluted Split pea.82 0.35 28.35 28.35 28. prepared from ready-to-serve can Tomato. May 6.35 28.00 248.80 0.35 28.28 0.57 3.00 256.19 1.00 252. undiluted Pea (green).41 0.43 0.13 0.11 0. undiluted Onion.21 16.62 0.31 0.08 1.21 0.35 28.00 245.96 0.57 0.23 0.99 0.71 7.10 2.05 14.19 0.71 0.20 2.39 2.35 28.00 241.35 28.00 243.23 0.00 245. vegetarian.35 28. undiluted Vegetable.34 1.20 0.52 3. ham.91 1.36 1.74 0.56 0.33 6. undiluted Vegetable beef.15 0.77 1.43 0.00 241.44 2. chunky-style Vegetable beef.15 3.78 1.00 252.08 1.45 0.00 0. prepared from ready-to-serve can.23 0.35 28.20 0.35 28.29 0.00 252.69 1. clear (French style).35 28. prepared from ready-to-serve can Split pea with ham or bacon.41 0.12 1.65 0.32 0.27 1.18 0.00 256.35 28.35 28.02 0.20 0.77 0.63 1.34 0.10 17.59 6.35 28.35 28.33 1.00 252. prepared from ready-to-serve can Minestrone.60 3.33 1.1 Bridge mix Candy-coated almonds Candy-coated chocolate Candy-coated peanuts and chocolate Caramel.00 0. and nougat Chocolate-covered cherry Chocolate-covered coconut Chocolate-covered cream Chocolate-covered fondant Chocolate-covered marshmallow Chocolate-covered peanut butter Chocolate-covered peanuts Chocolate-covered raisins Chocolate-covered toffee 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Candy and Sweets Lentil.35 28. undiluted 2387_chzAppendix_T1_fm Page 639 Sunday.07 0.18 APPENDIX — TABLE A.18 7.61 2.00 244.00 0.79 1.35 28.48 0.89 1.44 1.46 8.66 2.02 0.68 15.22 0.41 0. 2001 8:09 PM 639 .67 0.00 245.33 0. prepared from ready-to-serve can Minestrone.35 245.18 1.61 0. plain (vegetarian).17 2.19 1.38 1.C C C C C C C C C C C C C C C 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 oz oz oz oz oz oz oz oz oz oz oz oz oz oz oz oz oz oz oz oz oz oz 28. undiluted Split pea with bacon.35 28.21 0. prepared from ready-to-serve can Tomato.76 1.18 3.03 1. or pork.56 8.31 0.07 0.15 0.53 0.20 4.28 3.12 0.05 0.35 28.18 0.35 28.10 2.27 0.96 0.31 0.56 1.71 2.01 0. plain Carob Carob-coated peanuts Carob-coated raisins Chocolate-covered almonds Chocolate-covered caramel Chocolate-covered caramel and nougat Chocolate-covered caramel and peanuts Chocolate-covered caramel. prepared from ready-to-serve can Vegetable beef.13 0. prepared from ready-to-serve can Vegetable.39 0.09 0.08 2. peanuts.13 7. 21 0.57 Total Dietary Fiber (g) 0.96 1.28 0. 3RD EDITION .00 1. and Miscellaneous Dietary Fiber Values for Common Foods 0.50 0.26 0.45 1.19 0.22 0.00 1.35 28.14 0.18 0.06 1.37 1.35 28.26 0.61 0.18 0.04 0.33 15.35 28.35 28.76 0.79 1.35 50.23 1.00 1.37 0.35 28.19 1.00 28.25 0.27 0.28 1.16 0.35 28.52 0. May 6.15 0.99 0.07 1.16 0.43 0.00 0.41 Insoluble Fiber (g) 640 Chocolate-covered toffee with nuts Dark chocolate Fruit leather or rolls Fruit snacks Halvah.23 28.35 28.54 0.39 0.35 1.17 0.16 Soluble Fiber (g) 4.00 2.62 0.35 28.10 0.17 0.28 0.39 0.60 2.00 2.93 2.72 0.60 0. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.1 (Continued) 2387_chzAppendix_T1_fm Page 640 Sunday. plain Salted nut roll White chocolate Yogurt-covered almonds Yogurt-covered peanuts Yogurt-covered raisins Table A.01 0.48 0.01 0.33 0.35 28.35 Weight (g) 1 TS 1 TS 1 oz 1 TS 1 TS 1 TS 1 TS 1 TS 1T 1 TS 1 TS 2.35 28.35 28.63 Spices. plain Licorice Malted milk balls Marshmallow Milk chocolate with almonds Milk chocolate with cereal Milk chocolate with peanuts Milk chocolate.17 0.68 0.50 0.00 0.Allspice (ground) Anise seed Baking chocolate Basil (ground) Bay leaf Caraway seed Cardamom (ground) Carob powder Catsup Celery seed Chervil (dried) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 oz oz oz oz oz oz oz C oz oz oz oz oz oz oz oz oz Candy and Sweets Amount a 28.05 1.07 0.35 28.33 4.00 1.88 0.78 0.85 0.28 0.05 0.35 28.15 0.35 28.26 0.53 0.67 0. Condiments.93 0.25 0. 01 0. stuffed Onion powder Oregano (ground ) Paprika Parsley (dried) Pepper (ground). without seeds Mace (ground) Marjoram (dried) Miso (soybean paste) Mustard Mustard.72 1.91 0.67 0.80 2.13 0.76 0.1 641 .04 0.22 0.57 0. red Pepper (ground).27 2.21 1.40 4.77 0.16 0.28 0.00 2.37 160. fresh Saffron Sage (ground) 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 Medium 1 Medium 1 Medium 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1 TS 1C 1C 1C 1T 1 TS 1 TS 1 TS 1 TS 2.12 0.12 0.62 1.00 160.00 3. black Olives. May 6.23 0.57 0.70 0.28 0.19 0. powder Ginger (ground) Hot chili pepper — mature red (dried).00 12.70 2.57 5.15 0.77 0.36 1.93 3.50 2.47 0.77 2.08 0.07 0.70 0.49 0.10 0. with seeds Hot chili pepper — mature red (dried).04 0.Chili powder Cinnamon (ground) Cloves (ground) Cocoa powder.46 0.21 0.09 2387_chzAppendix_T1_fm Page 641 Sunday.50 2.70 0.47 0.22 0.73 5.03 0.13 0.20 1.05 0.13 0.67 2. 2001 8:09 PM APPENDIX — TABLE A.00 0. unsweetened Coriander seed Cumin seed Curry powder Dill seed Dill weed.33 4.59 0.00 1.10 2. dried Rosemary.13 1.06 0.00 143.64 0.00 2. white Pickles.48 0.43 2.17 1.16 0.14 0.07 1.22 0.80 1.06 0. bread-and-butter Pickles.20 1. black Pepper (ground).76 1. powder Nutmeg Olives.85 1.03 1.36 0.36 0.36 1.50 2.75 0.23 0.48 0.10 0.36 0.13 0.00 0.40 0.79 1. dill Pickles.00 2.30 0.27 0. green Olives.23 0.02 0.03 0.00 1.08 0.40 0.18 0.03 0.31 0. dried Fennel seed Fenugreek seed Garlic. sweet — gherkins Pimiento Rosemary. 1300 South Second Street.00 Total Dietary Fiber (g) Insoluble Fiber (g) Soluble Fiber (g) a Abbreviations: T = tablespoon. BIS = biscuit. fresh Thyme (ground) Turmeric (ground) Yeast.00 0. Condiments. contact The Nutrition Coordinating Center.60 2. This table has been revised since the 1994 2nd edition.43 2. 2001 8:09 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 1 TS 1 TS 1T 1 TS 1 TS 1 Cake 1T 1T Spices. Suite 300.1 (Continued) 2387_chzAppendix_T1_fm Page 642 Sunday.27 17.40 1. For further information regarding specific values. TS = teaspoon. and Miscellaneous Dietary Fiber Values for Common Foods 642 Savory (ground) Tarragon (ground) Thyme. C = cup.38 2. Minnesota 55454-1015.67 0.Amount a Weight (g) 1. baking — compressed Yeast. 3RD EDITION . Brewers Table A. May 6.00 12.53 0.48 1.52 3. School of Public Health. oz = ounce.34 0.12 0. baking — active dry Yeast. Note: Includes both analytic and estimated values. Minneapolis. Division of Epidemiology.00 15. based on the availability of improved data.47 1. . Brillouet. physical. 81. D. Hoebler. Fiber contents of some Chinese vegetables and their in vitro binding capacity of bile acids. Boothby.-M.. 1983. J. 1997. Candlish. .. 62.. Burritt... A.. J. 112.. 2001 8:09 PM APPENDIX — TABLE A. 49. Lehtonen. F. 396. C. Agric.. The determination of fiber in processed cereal foods by near-infrared reflectance spectroscopy. W. E. S.. W. 11. and Jeltema. Nutr. 244. M.. Sci. 26. Food Comp. Effects of processing on the dietary fiber content of wheat bran. 1982. 87. 49. Food Agric.. J. D. Leung. 22. 33. 60. Carre. 1981. Y.. 24. Baker. J. J. 26. M. E. Insoluble dietary fibre content of some local foods. W.R. 1988. 21. 65.. 1988. Total dietary fiber content of selected nuts by two enzymatic-gravimetric methods. 27. 7. B. 476. Nutr. Fiber in breakfast cereals. Pectic substances in developing and ripening plum fruits. Barry. Proc. 1117.. 47. 41. Assoc.. J. X. H. 1982. HCF Nutrition Research Foundation. Composition of dietary fiber: neutral and acidic sugar composition of the alcohol insoluble residue from human foods. M. Food Agric. Diabetes Care. Am. Food Sci. 1189. Lexington. and Essatara. 23. 1978. 37. 440. T. J. R. 284.. Chemical. Cardozo. 77. Dietary fiber content of selected foods.. D. Brandt. Prep. J. 1988. Angus. K. Byrant. and Clydesdale. 1370. and Holden... J.. Food Chem. 373. M. 1994. 557. Agric. H. and Bridges. 1983. A new method for determination of insoluble cell walls and soluble nonstarchy polysaccharides from plant materials. 46. J.. Fiber in wheat foods. J. C. J. 25.. Building fiber into foods.. Dietary fiber and diabetes: a comprehensive review and practical application. L.. J. Total dietary fiber analysis of selected baked and cereal products. N. 161. Cereal Chem.. L. Cereal Foods World. Food Chem.. P. Anderson. Am. 1.. 23. R. R. 32. W. J. Tietyen. Nutr. 5. M. S.. 4. D. 31. D. Babcock. Jeltema. 10. Pectic substance content of detergent-extracted dietary fibers. Rep.. Cereal Foods World. 1987. Plant Fiber in Foods. Sci.. S. 1981. 1976. 31. Gourley.. P. L. Chen.. 1981. Nutr.. Food Sci. Anal. Anderson. Dietary fiber and starch in some Southeast Asian fruits. 36. Bell. C. Baker. 28. 7.. Johansson. B. A. Soc. Anal. J. 20.... 19. J.. 1533. Aalto. 1985. P. Cereal Foods World. L. 1990. D. Aust. 352... A. Reassessment of some fruit and vegetable pectin levels. 1987. 17... 18. Food Sci. and Guoo. Gustafson.. KY. Food Chem... 225. J.. N. S. O. J. K. M. B. L. M. Foods. Long-term effects of high-carbohydrate. M. and Street. 29. Belo.. Food Comp. A rapid method of dietary fibre estimation in wheat products. J. 3. Balasubramaniam.2387_chzAppendix_T1_fm Page 643 Sunday. H. Rice bran as a source of dietary fiber. 45. 15. Clin. and Baranowski. Nutr.. Chen... and Tietyen-Clark. Food Sci. J.. J..1 643 REFERENCES 1. 65. S. and Li. Am. 6. 47. 13. G.-G. W. Chang. Cereal Chem. and Farrell. J. 1978.. E. M. Baker. A. Int. B. Cardozo.. and Varo. 14. J.. 1983. J. Anderson.. Rep. 34. Asp. R.. D.... and baking properties of apple fiber compared with wheat and oat bran. Anderson. and de Lumen. 1469. 16. 217. C. S.. 1987. 414.. Clin. R. Rouau. Dietet. Anderson. W.A. K. 36. 1980.. N. Moser. J. J. and Ward.. Food Sci. Alaoui. Bridges. Anderson. Hallmer. May 6. 9.. Food Sci. J. pureed green beans. 538. Dietary fiber and phytic acid levels in the major food items consumed in Morocco. July. 1053. Dietary fiber content of simulated American diet and selected research diets. and carrots. Rubenthaler. 1989.. E. M.. Sutherland.. 900. B. Polysaccharides of the kernel of maturing and matured coconuts. Agric.. 1985..... C. Bittner. 1988. Rapid enzymatic assay of insoluble and soluble dietary fiber. M. and Siljestrom. high-fiber diets on glucose and lipid metabolism: a preliminary report on patients with diabetes. 815.. T. and Lorta.. 6. 8. J. S... 1984. Zabik. J. and Lee. and Gustafson. M. J. J. M. Best. Effects of cooking in solutions of varying pH on the dietary fiber components of vegetables. 1988. H. D. J.. Baker. Dietary fiber content of barley grown in Finland. 12. J. N. J. D. Cereal Chem. K.-G.-L. 969. 1987. 1. and Eitenmiller. Int. 469.. 2.. J. 30. M. 38.. 50. C. in The Analysis of Dietary Fibers in Food.. 1998. A.. Eds. H. Chem. J. 1987. 1077.. J. W.. 808. R. 37. W. O. Swarner. Rep. E.2387_chzAppendix_T1_fm Page 644 Sunday. Englyst. L.. Englyst. Femenia. J. 305. 273. 82. 47. 687. Detergent fiber analyses on thirty foodstuffs ingested by man. 1975. S. G. T. Assoc.. 1983. 48. Hellendoorn. 54. J. Dreher. Horvath. hemicellulose and lignin content of raw and cooked processed vegetables. 30. Faulks. and Rasco. 274. 1756. Marcel Dekker. 1981. 77.. and lignin contents of distillers’ dried grains with solubles... A. B. 1461. and Cummings. Improved method for measurement of dietary fiber as nonstarch polysaccharides in plant foods. L. 44. Breedon. W.. 1965. Am.. 1983. 1981. W. and Aman. 57. J. N... J. 420.... N. Dong. 820. 40. Food Sci. S.. 1981. Partially hydrolyzed guar gum as a source of fiber. W. J. 1927. Pomare.. M. 1980. 34. W. Anal. Vidal-Valverde. 46.. 1981.. J. 60. 33. 403. V. Composition of fruit.. Holloway. 17. 197. H. H. Nutr. Cereal Foods World.. 1996. 24. 42. 1434. J. P. 1339.. 55. 1236. and Timms. W.. 46. 85. Anal. Heckman.. 28... Evans. D. New York.. 34. D.. E. Food Agric.. J. Cauliflower (Brassica oleracea L.. G. crude fiber. Monro. The neutral detergent fiber. 34. A study of relationships between flatus potential and carbohydrate distribution in legume seeds. Food Agric.. J.. 48. . Graham. B. J. 46. Herranz. H. Sci. 53. J. Food Chem. J. and Slagman. May 6. Foy. Tirtohusodo.. Holloway. A. M. 52. J. Extraction and fractionation of insoluble fiber from five fiber sources. Cellulose.. hemicellulose and lignin content of raw and cooked Spanish vegetables.. Food Agric. and Wyatt.. 31. 463. Extraction of soluble dietary fiber. Int. S. 36. Dietary fiber content in cereals in Norway. Food Sci... 1988. N. James.. M. R. 703. Dietet. H. 34. and Zabik.. A. J. 31. Influence of the composition of potatoes on their cooking kinetics. Harada. 71. and Anderson. 1996. and Orr. K.. Effect of fiber in corn tortillas and cooked beans on iron availability.-G. H... 524. Greenberg. B.. Food Chem.. 35. The measurement of dietary fiber and the effects of fermentation. Fleming. and Weber. Carbohydrates in foods. C. Idouraine. Off. 794. Herranz. N. S. Food Agric. J. 64. 32. Food Sci. Med.. A. Jeltema. J.. Hardinge. J. Assoc. Cornell University. 36.. 511. 1985. Food Chem. K. C. Enzymatic determination of the indigestible residue (dietary fibre) content of human food. Noordhoff. C. 3RD EDITION 29. Frolich. A.. A rapid method for determining the carbohydrate component of dietary fibre. M.. and Stace.. 50. and Hestangen. 1983. 1988.. H. D. P. and Lane. 494.. and Asp.. 50. Chen. Cereal Chem. J. F. Sci. and Rojas-Hidalgo. Agric. 58. 43.. M. 1983. 41. J. 49. 2001 8:09 PM 644 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. vegetable and cereal dietary fibre. E. globe artichoke (Cynara scolymus) and chicory witloof (Cichorium intybus) processing by-products as sources of dietary fibre. and Wohlt.. A.Z.. 1981. W.. 52. 1981. Food Sci. Holloway. Tasman-Jones... Determination of carbohydrate and its composition in plant materials. Agric.. Waldron. Assoc. Am. and Paulus. 1984. M. Percent starch hydrolysis and dietary fiber content of chipped and baked potatoes. H. Food Agric.. S. Int. S. J. B. G.. J. Cellulose. Rep.. W. Sci. M. and Selvendran. L. Cereal Chem.. J. and Theander. 575. Soluble and insoluble plant fiber in selected cereals and vegetables.-J. and Rojas-Hidalgo. J.. and Sellman. 1981. and Maher. P.). Revised method for quantitating dietary fibre components. Nutr. 1977. Frolich. Anderson. Food Sci. and Crooks. L. Dietary fiber and other constituents of some Tongan foods.. 46.. W. E. K.. Comparison of dietary fiber methods for foods. Clin. W. acid detergent fiber. Sci. Nutr. W.. D. N. A. Thesis. 39. 1985... Sci.. Claye. N. 1983. Englyst. J..-B. H. E. Food Sci.. 26... Off. Dietary fiber content of different cereal products in Norway. Starch and non-starch polysaccharides in some cereal foods. 43. Rydberg. 51. E. and Cummings. C. 1982. Towards an accurate measurement of dietary fibre. J. P. Garcia-Lopez. J. T. C. R. J. Robertson. W. E. C. M. 45. Vidal-Valverde. J.. M. Gurnsey. Chem... J. 724. 1985. H. Food Chem. . Food Chem. 76. Characterization of starch and fiber of banana fruit. V. Food Agric. M. Potatoes. Flentje. M. P. 94. 42. 1990. P. Assoc. D. J Food Comp.. J. Food Chem. R. B. J. F. . L. 1981. Measuring dietary fiber in human foods. M. Kamath.. L. and Van de Bovenkamp. D. Agric.2387_chzAppendix_T1_fm Page 645 Sunday. Analyse van het totale voedingsvezlgehalte en van het pectine-aandeel hierin in Nederlandse voedingsmillelen. Effect of variety and growing year on the constituents of durum bran fiber. 75. J. S. 1983. G. 5. 42. 1979. Database and quick methods of assessing typical dietary fiber intakes using data for 228 commonly consumed foods.-F. Lanza. A. Aust. Agric. 1982. D.. 78. 57. Jones.. M.. A. 3944.S. 1986.. 143.. D. B. 1981. L. 146. Kunerth.. and Chesters. Marlett. 83. Briggs. and Belavady. Assoc. K. J. Lund. Johnston. G. 44. 8. 1995. and Wang. E. Dietet.. 1984. G.. P. Nutr. A. C. Prediction of total dietary fiber by near-infrared reflectance spectroscopy in cereal products containing high sugar and crystalline sugar. J. Windham.. Van Soest.. Y. A critical review of food fiber analysis and data. J. Agric. S. 71. A. Jones. 73. and Youngs. and Cheung. Jeraci. J. 1995. Chem. and Cappelloni. 1982. and Dorrian. Food Technol. 1983. 733. Dietary fibre content of Australian foods. 27. and Zabik.. G. Cereal Foods World. M. M.). 46. 175. and Sayre. P.. R. J. L.. 61.. Am.. A. E.. 77. Food Sci. Briggs. M. 5. Assoc. M. May 6... J. N. B. Food Chem. L.. W. F. Lund.. L.. 81. T. 1999. 30. 1989. Simplified enzymatic-gravimetric method for total dietary fiber in legumes compared with a modified AOAC method. Food Chem. Food Chem. Sci. W. 1997... E. Content and composition of dietary fiber in 117 frequently consumed foods. M. J.. Dietary fiber content of Italian fruit and nuts. Enzyme neutral detergent fiber analysis of selected commercial and home-prepared foods. L. 1... 59.. Unavailable carbohydrates of commonly consumed Indian foods. Lewis.. Wahlqvist. 790. Cereal Chem. J.. and Oliver.. M. 1982. Smoot.. Hudson. Dietet. J. A. and Himmelsbach. 417. J. D. E. Jones.. B. Manthey.. Losses of pectic substances during cooking and the effect of water hardness.. and Kessler. 1139. Kays. 69.. Dietary fiber intake in the U. Lanza.. Block.1 645 56.. Lintas. A. 58. W.. Marlett. Effect of β-glucan barley fractions in high-fiber bread and pasta. D. 86... Cereal Chem. P. 350. 60. J. 72.. 1993. 26. M. 194. W. Dietary fibre content of Australian foods. J. 1997. J. Li. 1013. 66. Voeding..-L.. L. 34. M. L. Food Aust. Leeds. J. Jones.. 1987. 677. and Cardozo. Sarhan. Urea enzymatic dialysis procedure for determination of total dietary fiber. S2.. Li. B.. J. 410.. Sci.. E.. Wahlqvist. Wahlqvist. Z. 924. N. 79. R. R. Jwuang. 68. V. C. B. R. Anal. 64. 81. 70. M. Dietary fiber content of some tropical fruits and vegetables. Off. W. R. Kelly. Dietary fibre content of Australian foods. Psyllium — a superior source of soluble dietary fibre.. W. Flentje. 45. 61.. Soluble and insoluble dietary fiber content and composition in oat. Food Technol. B.. Pectins and fibres in processing tomatoes. O. M. Anal. Longe. Katan. Anal. Changes in the composition and carbohydrate constituents of okra (Abelmoschus esculentus.. Dietet. 67.. 36. 42. 1982.. and Robertson. and VanSoest... 8.. Am. E. and Smoot.. M. and Akenova... J. 76.. 62. 1123.. P. D. and Butrum. Food Aust. Assoc. 80. Briggs. J. 72.. G. M. Knuckles. and Huseby. 1990. G. Hood... F. D. T. Food Comp. J. 27. B. E. Chiu. 70. B. B. 1992. 82... 1980. Luh. population. Food Sci... Comparison of three methods and two cooking times in the determination of total dietary fiber content of dried legumes. J. J. Barton. P. H. S.. E. 1992.. 47 (suppl. S. Kayisu. D. 99.. 153. 1985. and Flentje.. and Shiell. Am. 58. A.. Johnston. Marlett. 31. Effect of boiling on the carbohydrate constituents of some non-leafy vegetables.. J. 6.. Aust. Clin. E. Food Aust. T. 3. J. 65. 1885.. 929. 46.. 2. 2001 8:09 PM APPENDIX — TABLE A. Linn) with age.. The influence of cooking technique on dietary fibre of boiled potato. G. R. O. J.. and Shiell. 97. Vegetables. Fetuga. Hareland. 31. 1984. D. 1985. Longe. Food Technol. 1997. E. 17. 732. 63. Dietary fiber content of eleven tropical fruits and vegetables. M. 50. B. E.. J. G.. D. L. 92.. A.. Food Sci. J.. Am. and Hall. J. L. 1. 74. 37. J. Fruits and fruit products. P. Food Sci. Food Agric. R. 1980. 159.. using high-performance liquid chromatography to measure neutral sugar composition. A. R. and Harland. L. and Verdier. 36. A. lignocellulose and hemicellulose contents of selected foods determined by modified and unmodified Van Soest procedures. J. and Marlett. Marlett. Chem.. Ichikawa. 545. and Kumpulainen. 1988. N.. 677. 1994. 1979. Food Sci. J. Neilson... G. M. 100.. 1979. C. soluble. T.. J. P.. Devries. 67. Cereal Chem. Modification of the neutral-detergent fiber procedure for cereals and vegetables by using a-amylase. J. 53. 45.. Prosky. fruits and berries consumed in Finland. 1999. Food Sci. 2001 8:09 PM 646 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. W. G. R. 86. 1984. Schweizer. J. November. J.. 627. 1982.. Chem. R. Sumimoto. Chem.. and Brassard... 1994. meat and fish dishes. 311.. A. Nakahara. 65. L. and total diets: interlaboratory study. S. J. 1978. Am.. 1013. 682. T. Pritz. Furda. 1988. Determination of total dietary fiber in foods and food products: Collaborative study. 51. 85. 103. 33. 83. 36. Paul. P. 91. Brassard. 99. 1991. Taguchi.. 1342. R. Devries. 71. 1983. 1988. N. Schweizer.. J.. Mongeau. Prosky. M... and Marlett. and Furda. D.. Comparison of gravimetric and chemical analyses of total dietary fiber in human foods. Am... McCance and Widdowson’s “The Composition of Foods”: Dietary fibre in egg. C.. W. 104. Food Sci. D. Marlett.. 101. G. J. R.. A. B.. Dietet. and Vollendorf. Total dietary fibre contents in vegetables.. 317. T. T. D. pastas. J. A. McCormick. S. Food Agric. T. and Navis. 106. 159. and Lee. J. F. A. Asp.. A comparison between detergent and nondetergent analyses of dietary fiber in human foodstuffs. P. storage. Contents of dietary fiber in some Japanese foods and the amount ingested through Japanese meals. 1344. Agric. 93. Nelson. N. Clin. 77. 98.. Food Chem. J. Rep. hemicellulose.... J. J. 45. B.-G. B. 1986. Assoc.. Agric. Furda.. R. 31. . J.. Food Agric. J. F. 1982.. 47. Marlett. Monro. Food Comp.. and Garcia L. Anal. S. 97. 109. Anal.. Foods. Nutr... J. B. Food Comp.. N. Navarro. J. Prep... W. C.. 1985. Neri. Anal. Nutr. E. October. 68.. Schweizer. Food Chem. Harding. Characteristics of California navel orange juice and pulpwash.-G. 794. Int. 88. 59.. 1988. Soluble and insoluble dietary fiber contents of various breads.. Food Sci. J.. R.. 39. 96. and Damiani. 48. Nutr. A rapid method for the determination of soluble and insoluble dietary fiber: comparison with AOAC total dietary fiber procedure and Englyst’s method. 12. May 6. and Russell.. Effect of cultivar. W. E. Assoc. 550.. Am. 1983. and Sistrunk. Dietary fiber and some elements in nuts and wheat brans. and cooking method on carbohydrates and some nutrients of sweet potatoes. R. A. 65. Park. Byers... and Southgate. Bertelli. Determination of insoluble.. Chem... Plessi. J. Dietary fibre of coconuts from a Pacific atoll: Soluble and insoluble components in relation to maturity. Soluble and insoluble fiber in soda crackers. W. L. Function and nutrition guide fiber ingredient selections. and Gelroth.. J. 1988.. Dietary fiber content and composition of different forms of fruits. food products. McQueen.. DeVries. J. S. A..... Y. Anal. 1986. Mongeau. Chem. M. T. Konishi. Matthee. D.. A. Anal. N. Variability in the dietary fiber content of wheat and mixed-grain commercial breads. 1992. 676.. Soluble and total dietary fiber in white bread.. 3RD EDITION 84. 1985 94.. and Appledorf. J. Assoc. 1988.-G.. V. Off.. Determination of total dietary fiber in foods. J. Smolensky. I. and total dietary fiber in foods and food products: interlaboratory study. T. S. C. N. Nishimune. Hum. A.. 414... 90. Reinhold. Assoc. T. 62.. 1990–1991. J. Asp. Przybyla.. 95. 43. J. F. Sci. Anal. L. R. and Brassard. Ranhotra. J.. J. Effect of cooking on vegetable fiber. and rye flours on the Finnish market. 1326. L. 2. J. Plaami. 1989. Plaami. cellulose and lignin content. and Kunita. 51. R. 32. and Vandercook. C. 155. Food Comp. 1979. and Harland. Kumpulainen.. H. E. Ranhotra. Formulating fiber into foods. 7. Assoc.. E. Monzani. 92. J. Off. 87. F. M.. Asp. W. 105. Off. A. Yakushiji. A.. Fiber of the maize tortilla. and Nicholson. Food. and Gelroth. J. 26. 91. J.. L. 110. D.. G. J. Nutr. Cereal Chem. Simonetti. 1044. S. and Tahvonen. 134. T.. size. F.. A. 108.. 107. Dietary fiber. 89. I. N. J. 1688. L. J. Determination of neutral detergent fiber in breakfast cereals: pentose. Food Sci. J. Patrow. Reddy. Measurement of dietary fiber in a total diet study. J. 102. I.. Food Eng.2387_chzAppendix_T1_fm Page 646 Sunday. Prosky. J.. Food Sci. C. G.. Off. 1333. 1017. 335. 86. Mori. Glycemic response and fiber content of some foods. J. W. Sci.. Clin. J. 1980. Anal. S. J. Mongeau. . P. Food Sci. A. J. E. L.. 132. 1985. 34. Reistad... Res. Munguia. Sci. Cereal Foods World. Varo. 29.. I. and Rojas-Hidalgo. J. 64. Agric. 1978. Sci. and Koivistoinen. J.. 124.. 2001 8:09 PM APPENDIX — TABLE A. 1983.. Varo. 21. 133. 949. R. 293. Pectic substances in raw and cooked. and Perlmutter. Department of Agriculture. The analysis of dietary fibre — the choices for the analyst.. 1980. Am. San Buenaventura. J. Anal. Prunes.. 727. Seibert. J. O.. J. G. E. 31. A. 1354. and Rojas-Hidalgo. Andelic. Nutrient Data Laboratory. Reistad. Schaller. Assoc. Siddiqui. 37. Dietary fiber constituents of selected fruits and vegetables.. P. 189. dried. G. 1164. C. R.. and Aman.. 1979. 128. and DuPont... N.. C. T. and Hagen. 485. S. and Aman. P. P. . 140.... 120. L. Dietet.. J.. 31. Skurray. Sci. Effect of heat treatment on the dietary fibre contents of potato and tomato. Lara. Agricultural Research Service.. T. sorghum... Southgate. L. 31. Lopez. 1982. 116. 114. K. S. Pectic substances in fresh. D. Torp. Veijalainen.. J. Vidal-Valverde. 127.. Dietary fibre in raw and cooked potatoes. I. E. and distillers’ dried grain with solubles. A.. Slavin. Agric. Pero.. Theander. 1997.. Improved procedures for analysis of dietary fiber. R. Sci. J. J. J.. USDA Nutrient Database for Standard Reference. J. R. O... 1987. T. C. English. J.. The non-starch polysaccharide content of Mexican foods. 166. Vidal-Valverde. H. 39.. Blanco. C. 115. G. B. Studies on dietary fibres 1.. E.. B. M. 1989. G. 613. W. J. T. 1978. 47.. S99. Agric. Dietet. Clin.. A. Hudson. Blanco. Schweizer. Dietary fiber: physical and chemical properties. 19. 119. 537. 1840. 330. J. 31. 112. desiccated and oleaginous Spanish fruits. Agric. Analysis and chemical characterization of water-soluble and water-insoluble dietary fibres. J. Simplified methods for the preparation and analysis of dietary fibre. 113. 1984. F. J. 1994. and Westerlund.. J. R. D.. M.. 303. Nutr. K.. 1982. K. Food Agric. Am. Food Chem. 30.. Clin. methods of analysis and physiological effects. Fin. J.. Bailey. 126. 85. S. N. H... P. 3.... Assoc. Food Agric. 137... Swedish J. I. 17. 647. and Walker. I. Agric. 33. and Rasco. 39.. W.. February.. 979... C. Food Chem.. and Wursch. M. Englyst. O.. J. Dietet. Selvendran.1 647 111. R. P. K.. R. Dietary fiber values for various breads are higher using enzymatic analysis rather than detergent analysis.. de Lourdes Solano. 340. B. 1987. A.. Am. 94. F. Studies on dietary fiber. P. Laine. A. M. Food Chem. J. Muralikrishna. and Nguyen. E. 117. barley.. 135. P. T. 60.. 1984. Nutr. Sanchez-Castillo. The total dietary fiber content of wheat. 56. 1979. Food Chem. 138.. Fiber content and structure in foods. 541. 265. 104. D. 1980. 30. The plant cell wall as a source of dietary fiber: chemistry and structure. Agric. A. B. J. 1984. G. Sci. A guide to calculating intakes of dietary fibre. 1986.. F. L. S.. 32. 1987. J. Hudson. J. 139. 30. and Malleshi. Am. 1173. Oat bran as a source of soluble dietary fiber. Schneeman. Rice bran as a source of dietary fibre in bread. Food Comp. corn. Ross. Dietary fiber: analysis and food sources. 87. Herranz. J. C. 135.2387_chzAppendix_T1_fm Page 647 Sunday. Food Technol. Southgate. 134. chemical analyses and food sources. Food Agric.. 121.. Analysis of dietary fibre. P... R. Suhasini. Am. and James. U. Sci.. Nutr.. Dietary fiber: classification.. Am.. 1985. 130. 1986. 136. A. Somogyi. May 6. J. 123. Southgate. and Englyst. A method for the analysis and chemical characterisation of total dietary fibre. Theander. 32. Human Nutr.. Collinson. 129. Food Chem.. E.. 9. Studies on vegetables: fiber content and chemical composition of ethanol-insoluble and -soluble residues. a fiber-rich ingredient. Ross.. J. 1978.. Food Agric. 131. Food Technol. M. J.... Dietary fibre in some Norwegian plant foods during storage. 122. M. 118.. J. Food Technol. and Koivistoinen. 552. Theander. Selvendran. P. 1987. Dietary fibre and available carbohydrates in Finnish cereal products. Steen. Studies on dietary fibre. Food Chem. Food Agric. F. 1986. Release 13. 19. M. O. P. Assoc. A... Vidal-Valverde. 320. and Rojas-Hidalgo.. 1111. and Rogeberg. J. 31. R... E. Dietary fiber in Spanish fruits. 1976. Clin. Wooldridge. 12. J. D. 1999. S107. 832. Dong. Cereal Chem. Cereal Foods World. Veijalainen.. 1982. Food Chem.. D. Variation in the concentration of major carbohydrates in the grain of some spring barleys.. 1999. 125. 97. K. 1986.fresh or processed Spanish vegetables. Free sugars and non-starch polysaccharide contents of good and poor malting varieties of wheat and their malts. H. 223. Chem. 152. Peas and beans. J. R. R. 39.. Aust. K. fruits and fruit juices as served. Wills. S. A. 142.. T.. K. Food Technol. 1986. Inconsistent differences between neutral detergent fiber and total dietary fiber values of fruits and vegetables. Sci.. and Greenfield. Anal.. 1985.. Food Sci. and Hagen. C. Scriven. M. 36. 1988. 512. H. J... Agric. H. K. 1984.. H... 2001 8:09 PM 648 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. H. I. 39.. J. and Greenfield. J. Composition of Australian foods. Wills. . 1987.. Elkins. Aust. A. Dietary fibre fractions in cereal and cereal-containing products in Britain. M. B. Zyren. Off. Cereal Foods World. 40. 36.. Nutrient composition of Chinese vegetables. Lim. Composition of Australian foods. and Marlett. B. S. J... W. 1987. 413. 48. 69. and Agater.. 33. 1984. 1987. C.. H. 600. Wills. F. R.. Wenlock. Anal. Wills. H. 146. E. Food Chem. 148. Wong. H.. 39. B. J. and Greenfield.. Lim. 6. R. L.. and Greenfield. 1983. F. H. Aust.. B. and Gurnsey. 565. N. S. Composition of Australian fresh fruit and vegetables. W. Brassica vegetables. 147. Scriven. 149. 488. Food Agric. Vegetable fruits... R. Tuber. 1988. Comparison of two methods of fiber analysis of 58 foods. J.. Lim. 203. S. B. Food Technol. Aust.. R. A.. 32. What is dietary fiber and what is starch?. Wills.. R. R.. B.. Food Comp.. Wills. E. B. J. 39... 42. Lim... R.. 3RD EDITION 141. Lim. J. Aust. Composition of Australian foods. J. Vollendorf. H. and Greenfield. R. 523. F. and Greenfield. Composition of Australian foods. 23. J. Wills. Visser. 36. K. Assoc. May 6. K. 1984. J. 384.2387_chzAppendix_T1_fm Page 648 Sunday.. W. Sivell. root and bulb vegetables. Fiber contents of selected raw and processed vegetables.. S. 143. Food Technol. Evans. 38. R. Food Technol. Aust. H. H. H. J. Canned fruits. 1993.. Waslien. Composition of Australian foods. 312. 144. 113. B.. Food Technol. M.. 145. Food Technol. 150. K. 151. Dudek. 25.. J. 176. 00+$1.2 Dry Matter. May 6. Hemicellulose. Ash. 2001 8:10 PM Appendix — Table A. Robertson 0-8493-2387-8/01/$0. and Lignin Content of Selected Foods James B.50 © 2001 by CRC Press LLC 649 . Crude Protein.2387_chzAppendix_T2_fm Page 649 Sunday. Soluble Fiber. Cellulose. Neutral Detergent Residue. Total Dietary Fiber. toasted Apple honey wheat Bagel Bagel.01 0.32 9.25 1.79 88.96 17. white Bread.63 3.50 5.58 1.01 52.83 ND –1.33 0.82 69.00 0.24 5.65 2.42 1. wheat Bread.41 2.10 0.90 2.35 2.2 2387_chzAppendix_T2_fm Page 650 Sunday.19 26.39 2.14 0.36 TDF –0.09 9.72 91.76 4.13 ND 2.10 CP 5.46 3.11 ND 9.05 5.96 9.22 10.26 0.53 92.56 1.67 4.Description Apple honey wheat.17 2.03 0.72 5.19 6.04 0.22 1.44 3.29 0. self-rising Flour.94 2.99 3.54 –0.24 11.98 1. recipe.39 4.37 89.99 4.51 7. May 6.64 64.95 2.60 12. 3RD EDITION .77 93.92 12.26 0.53 3.87 3.41 1.23 63.72 9.20 94. corn.89 2.49 13. white Bread.81 1. buttermilk Biscuits.31 91.33 20.99 61.50 0.30 72.76 3.06 0.85 0. corn. reg.18 1.34 51.89 67. corn Bread.21 14.72 0. yellow Cookie.06 0.99 1.57 2. corn Muffins.78 91.67 2.94 ND ND –0.41 4.30 0.07 5.66 0.08 89. toasted Norwegian flatbread Oatmeal bread Pancake mix Brownberry Brownberry Kavli Pepperidge Farm Quaker Quaker Stroehmann Dr.08 ND 1.57 1.19 73.61 1. plain Biscuit mix. French Bread.35 5.33 –0.23 12.12 5.33 14.49 0.43 4.55 2.11 1.62 6.97 –1.77 2.19 3.52 4.14 7.07 0. enriched.48 13. baked Bread mix. plain Cookies.31 6.31 0.18 Ls 650 Cornell Quaker Brownberry Brownberry Maker Dry Matter Content of Selected Foods Breads.47 0.64 18.17 1.98 1.51 13.15 ND ND 8.31 2. plain Natural bran Natural bran. 2001 8:10 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.98 4.24 3.68 1.09 1.08 2.40 ASH 13.32 16.97 3.17 1.01 95.17 3.22 8. white Bread.57 –0.55 4.75 10. Olindo Quaker Pepperidge Farm Homemade Less Millbrook Less Quaker Quaker Homemade 76.37 0.24 5.19 1. white Bread mix.22 10.18 5.76 11.02 5.46 1.39 4.43 2. Oreo Cookies.06 70.89 92.35 0.33 2.13 1.25 0.95 5.46 0.70 27.00 5.22 0.20 1.15 14.05 8. etc.94 11.01 ND ND 3.21 19.71 3.31 14.29 19.38 1.32 0.95 In Dry Matter (%) SF NDR 4.63 62.46 1.13 1.52 18.28 18.25 3.42 5.52 2.33 4.25 90. shortbread Fibre Goodness Bread Fibread Flour.20 2.44 8.70 8. Table A.19 0.30 3.28 0.33 3.07 0.39 64.06 4.09 6.66 0.28 0.73 0.25 1.52 1.74 3.13 57.60 ND ND –0.03 2.63 3.62 95. ginger snaps Cookies.18 1.20 ND ND 9.37 3.81 56.99 14.26 20.69 3.76 6.54 0.58 0.00 2. white Graham crackers Muffin mix.68 3.39 % DM ND ND 2.71 14.29 6.85 3.79 1. whole wheat Cake. blueberry Muffin mix.47 15.41 62.80 4.25 –0.50 0.70 7.70 3.06 6.87 8.16 7.11 0.50 0.17 2.38 0.35 1.10 1.69 2.61 1.22 59.39 13.24 HC 2. yellow Bread.21 90.77 19.16 0.40 6.99 ND 12.99 19.13 0.21 4.40 3.13 –0.84 5.46 0.21 0.13 7.64 16.65 52.56 3.17 2.25 3. Crackers.71 1.05 5.69 0.96 88.36 2.34 1.17 2.00 0.52 CE 1.65 3.19 2.97 1.82 3.98 0.52 1.13 8.06 17.22 1. 61 12. egg.05 5.53 0.35 1.90 3.07 4.79 1.92 2.40 10.92 5.87 0.12 0.63 3.26 90.69 0.90 38.03 6.14 1.77 8.49 9. sweet.70 7.49 0.67 4.29 3.73 10.77 69.63 0.76 94.25 12.30 2.03 0.01 4.14 0.03 0.96 4. cinnamon Rye cakes Saltine crackers Sesame crackers Snackbread Snackbread.75 2. whole wheat Swedish rye crispbread Taco shells Taco shells Triscuit Wheat cakes Wheat thins White extra fiber.31 3.05 0.57 2.67 13.89 2.17 0. 2001 8:10 PM 651 .57 0.21 11.87 ND ND 9.50 15.78 8.00 2.55 11.65 4.50 1.94 90.04 2. whole wheat Pasta Pancake mix.64 9.41 3.60 7.93 96.41 4.80 19.30 12.68 15.47 2.72 16.57 1.76 12.92 2.92 93.66 7.79 96.88 2.69 –1.58 9.2 Kellogg’s Nabisco Nabisco Nabisco Quaker Goia Goia Nabisco Quaker Nabisco Arnold’s Arnold’s Ak-Mak Ryvita Ryvita Siljans Old El Paso Quaker Quaker Quaker Nabisco Wege Wege Friehofer Quaker 2387_chzAppendix_T2_fm Page 651 Sunday.95 1.79 4.45 3.44 95.42 2.45 12.50 18.88 97.46 1.40 1.78 96.55 4.93 12.16 3.21 6.49 12.85 12.86 97.81 3.20 8.82 9. May 6.04 2.28 16.07 2.80 6.18 22.87 5.96 9.37 0.15 1.33 0.39 4.93 2.30 7.82 10.87 4.78 0.31 1.00 0.01 24.10 4.49 7.50 1.63 8.69 2.59 0.76 21. whole wheat Premium saltines Pretzels.74 3.42 13.20 0.16 32.54 15.61 0.63 32.68 1.55 6.42 96.47 –0.25 2.33 90.87 12.36 5.35 1.79 1.67 10.55 4. buckwheat Pancake mix.86 60.30 1. Cooked Noodles.16 36.55 7.27 0.61 0.53 4.17 0.14 0.18 30.35 1.53 0.92 6.09 10.88 10.84 1.42 97.60 95.50 2.67 34.00 1.12 4.86 1.53 4.57 0.18 2.07 3.18 0.76 27.45 16.24 3.27 2.77 12.72 1.48 0.15 8.64 0.56 3.88 5.00 1.92 0. whole wheat Prograin bread Rice cakes Rolls.16 8.60 33.80 2.38 91.94 4.40 35.46 0.16 22.54 16.32 2.78 19.37 0.42 1.75 15.25 0.64 5.02 11. cooked Spaghetti Spaghetti.36 3.92 73.41 3.55 5.74 3.54 6.29 11.57 5.51 2.46 2. cooked Spaghetti.22 0.70 3.71 95.58 8.77 7.19 6.37 0.63 1.48 2.45 6.76 2.45 6.78 1. hard Pretzels.100% Bran 100% Bran 100% Bran 100% Natural 40% Bran Flakes Bran Flakes All Bran Cereals Macaroni Macaroni.43 9.18 17.12 6.62 0.79 96.41 7.06 5.72 0.00 3.18 0.15 7.73 1.47 2.89 9.30 6.42 7.85 2.05 16.66 1.13 5.26 1.13 0.61 96.26 5.92 –0.19 94.07 ND ND 3.76 4.64 1.36 11.98 1.55 7.84 95.10 15. toasted White extra fiber 95.37 1.28 5.44 1.50 1.64 6.44 37.32 1.78 93.44 11.70 5.49 3.06 0.01 28.27 2.28 0.45 2.34 6. cooked Spaghetti w/sauce.91 10.50 4.62 1.11 13.57 ND 1.67 10.71 2.60 92.89 5.66 13.04 63.07 5.96 94.57 APPENDIX — TABLE A.61 8.33 5.00 12.02 37.17 1.21 3.96 15.47 4.12 26.86 12.24 1.90 5.57 90.41 95.69 1.53 95.14 3.75 19.36 3.88 0.67 2.11 2.84 7.20 7.46 2.87 2. 75 4.03 2.43 0.21 0.93 2.24 95.19 0.63 94.22 2.00 6.71 3.15 97.62 3.76 10.28 25.82 0.29 6.94 3.46 47.28 10.35 94.14 1.50 0.12 1.07 3.69 20.20 96.33 3.30 3.76 45.30 4.33 4.10 47.21 0.54 ND ND 3.06 12.59 4.11 1.21 1.54 7.62 11.07 96.39 6.12 7.77 9. -0.22 0.78 0.30 ND ND ND 21.58 0.08 In Dry Matter (%) SF NDR 22.83 % DM 6.47 4.36 0.56 6.60 ND 2. 3RD EDITION .49 4.64 0.25 1.25 7.09 0.43 4.14 4.59 2. Raisin Date Cereals Description Table A.82 94.65 3.70 3.67 6.03 3.49 2.44 –2.09 89.19 2.88 0.54 ND ND ND 8.99 1.66 96.30 2.39 5.27 2.17 5.59 97.23 1.61 16.24 1.75 6.09 34.61 95.55 5.66 12.83 0.34 0.98 7.35 0.78 2.45 15.00 ND 14.98 2.18 2.87 4.33 87.71 2.94 4.10 6.13 4.94 1.46 1.14 33.59 18.14 4.09 4.81 1.37 2.68 0.27 4.72 0.90 2.15 5.97 90.44 94.34 7.96 ND ND ND 7.76 1.46 49.76 11.26 50. Cinnamon Natural Cereal Natural Cereal.12 3.04 3.39 11.38 16.26 8.74 7.18 3.06 0.Quaker Quaker Quaker Quaker Quaker Quaker Quaker Quaker Quaker Quaker Quaker Quaker Ralston Purina General Mills General Mills Quaker Quaker Kellogg’s Kellogg’s Kellogg’s Kellogg’s General Mills Kellogg’s Kellogg’s Kellogg’s Kellogg’s Kellogg’s Quaker General Mills General Mills Maker 94.21 2.35 0.41 38.42 96.41 1.72 10.74 44.93 0.46 0.71 0.03 11.97 1.65 ND ND ND 2.84 1.45 5.04 2.48 94. cooked Creamy Wheat. instant Honey Graham Oh’s King Vitamin Life Life Life.87 0.76 40.66.93 ASH 12.99 15.59 ND ND ND ND ND CP 37.19 0.98 5.48 2.19 1.60 6.78 1.70 0.43 97.69 0.78 10.08 1.04 0.45 96.56 2.72 5.19 95.47 10.47 3.22 33.24 2.32 8.82 1.92 8.74 2.39 CE 4.07 ND ND ND ND ND 35.22 0.66 0.04 4.47 3.95 1.14 0.98 98. Apple Cinnamon Natural Cereal.35 9.47 1.39 1.09 1.04 13.42 48.91 3.32 9.00 6.78 13.95 0.38 3.08 48.57 6.78 ND ND 7.30 9.73 6. 2001 8:10 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.24 6.82 97.59 11.93 95.76 4.74 4.61 4.69 6.31 TDF 6.63 12.30 3.79 0.73 9.90 3. quick Crunchberries Crunchy Nut Oh’s Double Chex Fibre One Fibre One Grits Grits.87 7.52 1.36 95.87 2.40 ND ND 8.10 9.82 12.34 4.78 HC 8.97 2.47 93.38 96.55 8.28 3.93 95.01 1.74 0.03 6.82 0.79 11.91 33.46 0.32 3.14 0.68 3.80 11.46 5.51 1.81 4.59 95.2 (Continued) Dry Matter Content of Selected Foods 2387_chzAppendix_T2_fm Page 652 Sunday.94 3.99 10.65 2. May 6.25 2.35 3.39 95.84 96.50 4.56 6.73 94.91 Ls 652 All Bran All Bran All Bran w/ Extra Fiber All Bran w/ Extra Fiber Bran Flakes Cap’n Crunch Cheerios Cheerios Cheerios Choco Crunch Corn Bran Corn Flakes Corn Flakes Corn Flakes Corn Flakes Corn Flakes Cream of Wheat.95 1.95 6.50 10.40 95.08 1.40 4. 38 0.71 17.33 1. Instant.09 4.57 7.31 5.81 ND ND ND ND 2. Instant Oats. Instant.13 9.83 91.77 6.21 ND ND 8.04 14.39 10.25 0. Instant.86 93.70 95.40 3. One-Minute.65 Quaker General Mills 2387_chzAppendix_T2_fm Page 653 Sunday.00 0.91 8.02 10. Instant.71 0.71 1.56 4.73 7.75 8.12 7.80 2.23 2.02 13.71 3.66 4.72 0.79 1.28 2.15 0.82 7.62 5.54 2.90 2.43 13.64 5.42 4. Instant.50 1.10 93.08 10.48 0.05 3.88 1.94 95.76 0.61 2.95 91.36 91.01 3.78 2.35 96.30 ND ND ND ND ND ND ND ND ND 9.01 1.56 0.30 17.18 0.67 3.62 11.21 12.21 0. One-Minute. Strawberries and Cream Oats.80 11.21 2.70 4.64 3.30 1.16 1.86 16.93 0.16 2.91 0.69 0.34 12. Old Fashioned Oats.97 10.60 3.82 1.21 13.38 0.81 3.76 0.45 97. Apple Cinnamon Oats.49 1.25 12.17 93.23 2.89 1.10 20.82 2.24 2.61 3.39 1.39 0.47 1.98 6.25 89.35 5.57 92.85 92.12 96.88 3.73 2.42 8.50 3.43 95.80 9.49 14.42 3.81 9.83 1.82 9.47 22. Blueberries and Cream Oats.19 95.37 0.74 3. Apple Raisin Spice Oats.16 3. Peaches and Cream Oats.50 1.41 9.30 92.66 1.68 0.50 5.65 10.12 1.56 3.Natural Cereal.81 94.36 0. May 6.41 90.32 ND ND ND ND ND 4.00 2.30 19.23 2.39 0. Raisin Walnut Oats.51 11. Raisin Cinnamon Peanut Butter Crunch Popeye Sweet Puffs Post Toasties Product 19 Product 19 Puffed Rice Puffed Rice Puffed Wheat Puffed Wheat Rice Krispies Rice Krispies Rice Krispies Shredded Wheat Shredded Wheat Shredded Wheat Shredded Wheat N’ Bran Special K Sweet Crunch 91.79 1.24 2.12 95.37 95.14 3.15 0.35 6.92 6.45 1. 2001 8:10 PM 653 .50 12.21 3.99 5.88 2.28 8.15 3.71 3.37 0.08 1.81 6.86 0.17 1.80 13.45 10.74 2.28 ND ND ND ND ND 5.40 2.35 0.02 90.81 2.13 1.35 0.62 92. Apple Raisin Oats.90 9.97 0.57 0.36 1.79 3.84 1.38 0.50 1.00 1.18 8.76 6.12 ND ND ND ND 6.79 1.29 6.76 3.51 13.48 0.25 1.68 0.05 0.52 0.11 4. Instant.23 2.88 0.83 95.28 5.04 5.21 94.77 3.61 6. Instant.12 2.04 1.26 3.35 3.47 5.17 0.08 0.23 4.13 2.98 7.79 94.66 1.31 1.35 2.38 4.20 ND 1.50 10.93 0.28 94.12 92.85 1.26 2.78 2.10 5.26 1.30 93.63 0.90 0.93 1.27 1.38 1.97 1.91 3.87 ND ND 3.74 96.07 12.24 1.66 0.64 11.08 9.87 16.24 9.72 6.2 Nabisco Nabisco Kellogg’s Quaker Kellogg’s Kellogg’s Quaker Quaker Quaker Quaker Quaker 88.91 ND ND ND ND ND ND ND ND ND 6.66 0.69 1.16 2.54 0.98 3.08 97.00 0. Raisin Date Walnut Oats.39 5.64 10.88 3.96 13.75 6.13 12.45 1.01 2. Whole Wheat Nature Valley Granola Nature’s Harvest Granola Nutrigrain Oat Bran Oat Bran Creamy Hot Cereal Oats.59 7.89 1.56 4.80 1.08 4.10 0.13 3.20 7.77 4.22 90.61 3.20 Quaker Quaker Quaker Quaker Quaker Quaker Quaker Quaker Quaker General Foods Kellogg’s Kellogg’s Quaker Quaker Quaker Quaker Kellogg’s Kellogg’s Kellogg’s Nabisco 2.41 97.38 APPENDIX — TABLE A.38 3. 97 3.83 0.84 4.85 1.99 1.15 0.20 2.90 7.24 3.79 TDF –1.79 8.61 4.24 1.35 2.83 5.36 14.87 1.83 0.02 1. 3RD EDITION .06 0.80 2.14 1.73 14.13 ND ND ND ND 7.06 6.43 12.16 12.81 11.32 6.96 5.25 1.25 2.39 4.04 7.66 1.16 1.63 2.12 1.45 95.58 11.75 0.92 5.54 20.78 26.77 1.11 7.13 14.66 0. tart Canned Grapes.83 10.19 3.33 2.95 16.35 7.97 85. Florida.76 11.92 3.35 30. peeled Fresh Oranges.2 (Continued) Dry Matter Content of Selected Foods 2387_chzAppendix_T2_fm Page 654 Sunday. Red Delicious.14 3.82 4.66 5.17 10.97 0.91 1.Apple Juice Apple Sauce Apple.43 0.27 95.79 13.25 0.77 % DM 10.25 5. California Banana Blueberries Blueberries Fresh Canteloupe Fresh Cherries.00 4. cored and peeled Avocado.35 7.01 1.29 1. Red Delicious.10 17.53 1.05 15.40 4.67 4.32 3.22 4.55 8.68 1.59 5.78 0.46 2.72 12.49 3.28 17.70 0.88 ND ND 13.56 3.26 4.46 4.15 9.68 8.88 0. Thomson Fresh Orange juice Orange.31 2.78 3.21 3.63 2.41 1.81 3.35 32.32 2.54 10.24 2.75 15.25 1.83 17.52 0.17 1.77 13.94 2.73 15.56 13.12 22.91 2.13 0. 2001 8:10 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.31 0.07 2.64 10.98 10.54 4.96 12.73 22.10 4.37 7.06 ASH 10.37 ND 7.95 3.20 ND 11.19 0.59 ND 6.82 CP 25.02 1.77 94.62 5.32 4. mandarin Canned Peaches Canned Pear.50 0.25 5.63 1.60 8.35 0.98 4.25 2.22 13.71 11.79 1.96 1.88 0.10 0.53 2.51 11.38 CE 9.27 1.22 5. peeled Fresh Orange.28 4.66 18.72 11.27 4.74 9.93 4.32 7.00 5.91 6.78 2.64 13.65 5.61 1.69 3.82 7.02 4.38 1. Bartlett.76 10.48 1.85 1. cored Fresh Pears Canned Pineapple Canned Plum.41 2.38 2.81 2.03 1.47 0. friar Fresh Raisins.83 2.11 5.10 27.19 0.71 Ls 654 Fruits Team Total Wheat Germ Wheat Germ Wheaties Cereals Description Table A.86 HC 10.85 8.12 4.94 0.70 17.26 6. seedless Fresh Strawberries Wegmans Cornell Cornell General Mills Nabisco General Mills Kretschmer Maker 2.63 0. May 6.70 10.27 0.14 8.49 9.95 21.43 4.88 1.87 3. cored Apple.09 1.35 0.04 In Dry Matter (%) SF NDR 7.85 18.91 13.51 2.98 4. navel.57 17.05 1.74 5.25 6.15 0.35 1.28 96.80 5.25 5.83 9.32 1.90 95.73 0.73 5.51 8.63 3.06 3.69 1.75 6.63 0.54 8.33 3.75 26.91 2.68 7.11 2. frozen Brussels sprouts.01 5.42 11.65 9.43 6.70 37.10 36.40 5.03 8. May 6. frozen Broccoli.85 0.90 45.73 25.83 7.08 32.64 18.05 3.75 21.66 37.25 0.66 11.64 11.59 2.54 1.18 ND 18.10 1.76 13.98 25.17 7.10 ND ND ND 10.90 ND 10. lima.66 6.18 11.61 8.20 8.40 48. green.46 27.29 ND 15.80 0.77 2.50 18.47 1.24 0.07 0. pork.45 4.94 ND ND 12.20 20. cooked Broccoli.10 22.29 7.27 3.51 8.82 ND 6. 2001 8:10 PM APPENDIX — TABLE A.62 14. cooked Cauliflower.69 ND ND 17.35 5.18 ND ND 16.64 1.22 6.83 43.80 1.34 15.48 7.62 8.63 5.57 6.60 24.30 ND 18.24 21.00 11.31 10. wax. frozen Asparagus.83 32.53 34.62 1.22 37.10 10.53 28.92 2.89 10.36 1.41 10.69 32. frozen Cabbage.18 43.26 ND 19.89 5.86 0.52 0.67 9.10 ND 23.11 23. frozen Beans. cooked Brussels sprouts.70 11.58 40.01 3.86 35.35 21.85 20.43 11.34 3.52 10. boiled Beans.70 9.29 10.95 14.64 1.37 16.74 2.93 39. canned Broccoli. green.42 34.20 18.97 15.90 ND 20.18 12.21 4.21 10.96 2.40 8.53 6.64 21. kidney.17 ND 6.10 3.50 7. navy.59 6.30 33.30 7.34 24. canned Beans.41 ND ND 20. frozen Asparagus spears.88 1.93 2. canned Beans.10 1.53 5.10 8.01 1.79 3.31 30.30 8.10 8.79 7.31 ND ND 7.33 13.70 3.14 17.99 18.27 15.91 6. red.49 8.67 34.16 0.50 2.13 4.25 27.60 39. canned Bean sprouts.12 9.68 3.20 15.15 11. boiled Cabbage Cabbage.49 14.12 9. canned Beans.14 3. peeled Watermelon Fresh Fresh Fresh General Foods General Foods General Foods General Foods General Foods General Foods General Foods General Foods 12.57 10.58 1.50 9. microwaved Carrots.61 13.75 16.12 18.08 35.90 18.47 6.68 ND ND ND ND 15. boiled Carrots. frozen Cauliflower Vegetables Strawberries Tangerine.69 17.84 33.08 23.07 2. cooked Cauliflower.19 7.01 15.25 0.01 ND ND 11.78 1. microwaved Beet root. canned Beans.49 36.61 29.30 14.50 6.37 ND ND 7.40 36.30 34.30 9.19 24. lima.81 16.08 11.98 28.76 17.10 2.76 4.50 15.37 12.26 35.48 6.88 1.78 4.91 28.09 34.49 1.68 1.89 19.16 9.26 ND 14. frozen Beans.50 1.93 ND ND ND 5.63 6. wax.36 ND ND 10.58 3.59 ND ND 4.70 30.Artichoke hearts.89 ND 9.60 17.96 3.43 5.86 27.36 24.54 0. frozen Beans.70 ND 7.93 5.80 1.78 3.85 8.59 12.40 ND ND 3.00 23.67 1.40 3.58 0.71 19.42 11.62 ND 6.40 22. green. peeled Carrots.91 ND 23.00 4.34 10.51 29.43 22.22 16.73 ND ND ND 10.32 35.70 24.32 ND ND 10.89 13.64 ND ND 5.50 1.60 36.2 655 .91 14.75 ND 4.36 0. cooked Beans.23 28.60 13. microwaved Beans. green.52 4. kidney. boiled Beans.92 33.49 18.80 13.41 30.87 11.34 27.25 2.17 6.60 2387_chzAppendix_T2_fm Page 655 Sunday. lima.49 27.17 23. cooked Cauliflower.30 10.76 4.17 9.83 5. green.83 13.22 54.05 0.77 15.31 7.20 12.69 18. frozen Beans.19 20.01 12.24 2.10 9.74 7.18 36. canned Beans.38 14.50 12.20 47. boiled Beans. 06 0.17 0. black-eyed.10 1.93 0.21 40. stuffed Onions.23 5. boiled Peas. boiled Onions.34 30.93 –5.25 2.96 11.10 26.32 CP 40.10 7.80 18.09 1. with skin Cucumber.51 14. dill Potatoes.30 7.61 10.95 30.11 CE 1.65 9.16 ND ND 4.24 3.00 20.01 0. frozen Okra.2 (Continued) Dry Matter Content of Selected Foods 2387_chzAppendix_T2_fm Page 656 Sunday.90 ND 5.55 1.65 7.64 15.66 6.26 17.86 19.68 0.75 35.10 0.21 4.06 1.25 3.62 18.17 9.02 6. frozen Corn.71 0.58 7.48 18.56 2.60 11. boiled Potatoes.37 1.90 4.90 17.95 12.00 6.88 1.36 8.88 23.60 5.05 7.73 1.20 ND 20.71 44.94 0.95 20.65 16.52 25.32 1.60 9.18 10.75 In Dry Matter (%) SF NDR 1.57 9.32 27.65 21.24 1.49 34.21 0.49 7.24 27.61 9.00 ND ND 23.00 41.Celery Collard greens.31 ND ND ND ND ND ND 10.08 40. canned Peas.93 12.99 4.28 0.07 7. green seeded Pickles.69 3.29 25.58 4.49 20.87 11.00 33.70 24.10 4.91 0. peeled.30 4. black-eyed.00 11.85 % DM 16.80 1.90 21.23 ND ND ND 5.16 4.68 6.94 13. black Olives.83 7.95 ASH 11.51 29.77 4.20 ND ND ND 8.47 10.43 1.79 0.09 2. 3RD EDITION .50 6. w/skins. green.73 9. frozen Pepper.86 –2.58 18.68 13.30 19.28 9.13 27.88 7.63 11.04 18.41 2. microwaved Peas. mashed Potatoes.55 ND 1. green Peas.41 13.74 9.60 21.96 20.34 32.54 31.10 11.39 14. boiled Vegetables Description Ore-Ida General Foods General Foods General Foods General Foods General Foods 5.60 9.26 1.35 28.91 11.10 2.31 1.55 12.50 43.29 18.30 7.90 13.43 16.21 3.24 26. baked Radishes Rice.80 2.93 0.40 9. canned Corn.79 0.30 2.91 14.09 7.00 ND 6.12 32.00 3.17 ND ND ND ND ND ND –6.69 20.21 19.88 2.36 23.10 7. boiled.97 2.67 ND 9.49 7.40 3.07 3.07 4.14 7.86 27.89 3.60 12.62 0.44 16.67 2.83 7.70 ND ND 31. frozen Cucumber.65 27. 2001 8:10 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. frozen Onions.83 12.97 8.91 32.24 8.78 1.02 6. canned Peas.84 12.66 3. frozen Olives.79 2.87 10.40 40.80 10.17 12.58 1. black-eyed. french fries Potatoes.05 3.09 TDF 19.11 13. peeled Kale.92 6.98 3.90 11.25 0.74 10.50 18.60 7.15 0.01 2.86 7.00 19. boiled Mushrooms Mustard greens.35 0.20 5.98 ND 10.75 26.29 3.03 61.16 0.05 6.00 14.09 0.95 9.40 4.72 27.85 7. peeled Onions.20 8.18 ND ND 8.40 2.43 ND 10.40 26.74 14.39 11.84 19.80 3.69 1. green.60 7.70 5.39 7. May 6.58 20.90 1.96 6.10 2.80 HC 17.96 0.31 14.90 6.83 Ls 656 General Foods General Foods General Foods Maker Table A. cooked Rice.22 3. frozen Lentils.71 19.96 20.20 5.83 36.09 12. green.50 3. frozen Peas.93 3.30 20.40 4.69 9. frozen Okra. green.08 ND 21.38 25.96 ND 3.36 0.17 ND 8.28 13.20 5.60 0. 76 0.90 0.38 1.73 0.63 95.47 8.73 8.34 0.70 93.93 ND 0.46 12.48 0.05 0.86 4.46 0.10 8.77 8.93 3.88 95.08 14.73 53.48 5.67 0.00 4.51 12.48 5.27 74.12 6.69 2.94 9.98 10.00 0.43 1.57 84.53 28.96 5.40 14.23 1.43 10.99 8.35 10.34 1.48 19.28 6.16 1.49 2.40 1.64 ND ND ND ND ND ND 11.37 0.00 0.91 8.96 13. 2001 8:10 PM 657 .57 16.40 0.00 4. baked Tomato catsup Tomatoes.57 83.32 6.36 1.60 1.47 35.04 33.84 5.10 96.60 3.41 0.08 6. frozen Squash.41 20.05 3. summer.47 34.19 7.60 23.65 12.60 3.12 ND 11.45 9.42 23. microwaved Squash.58 2.86 65.06 2.10 32.28 21.91 1.90 2.86 27.80 58.52 15.56 15.70 1.09 21.63 ND 13.85 14.46 8.20 5.00 0.70 0.00 14.23 1. frozen Squash.27 1.35 2.00 6.11 6. May 6.41 14. frozen Vegetable soup.82 72.59 2.99 3.12 19.12 80.54 9.14 9.28 47.74 1.38 5.86 1.79 95.50 7.63 2.16 2.51 7.2 Fiber 88 Fiber Diet Fiber Diet Fiber Diet Fiber Filler Fiber Full Fiber-Off Fiberall Orange Sugarfree Fiberall Regular Sugarfree Fiberall Wafers Fiberguard Fiber Supplements Fruit Wrinkles (Fruit Punch) Fruit Wrinkles (Strawberry) Other Peanuts Walnuts Nuts Rutabaga.02 3.32 0.20 9.80 5.12 52.99 0.18 6.89 4. frozen Sweet potatoes Sweet potatoes.40 1.82 10. canned Yams 2387_chzAppendix_T2_fm Page 657 Sunday.47 4.90 21.72 ND ND ND 20.48 ND 48.45 3.80 54.05 11. canned Turnip Turnip greens.27 ND ND ND ND ND ND 11.10 1.77 41.15 9.32 0.15 95.27 4.45 94.00 0.58 ND 17.00 0.55 51.21 7.48 7.91 12.90 12.60 9. cooked Squash.75 12.62 14.31 17.93 3. boiled Spinach.82 ND 2.32 16.82 1.67 29.37 0.24 37.77 ND 2.11 26.50 99.33 0.42 7.80 1.49 20.15 1.51 19.57 8.63 2.55 1.40 APPENDIX — TABLE A.48 ND 29.23 21.70 1.37 62.59 93.83 22.70 0.32 4.40 8.27 45.28 12.16 10.60 11. microwaved Squash. zucchini.25 6.16 94.15 8.46 2.87 4.70 11.90 4.33 15.14 13.60 ND 19.60 20.27 0.60 4.34 8.81 ND 4.28 24.39 35.64 93.70 4.60 16.70 2.15 15.36 39.52 28. summer. frozen Turnip greens.13 0.47 2.27 11.33 ND 6.49 91.07 89.36 6.12 10.16 1.90 31.77 33.00 3.88 2. zucchini Squash.21 21.91 2.30 28.50 8.18 89.40 13.74 2.53 26.93 11.36 21.27 21.96 ND 1.99 0.80 4.00 52.63 29. summer.24 6. Your Life Solar Nutrition Nutrition Headquarters Rydelle Laboratories Rydelle Laboratories Rydelle Laboratories Ayerst General Foods General Foods General Foods General Foods 94.34 47.09 13.30 14.00 12.92 6.00 0.39 64.Fresh Fresh Natrol Puritan’s Pride Vitamin World BQE Vit & Suppl.95 3.29 11.43 18.87 2.94 ND 48.47 ND 13.72 33.65 6.80 1.00 29. 47 89. Samples provided mainly by J.66 11. (B) Neutral detergent residue and its components by the methods of Van Soest: Robertson. and Van Soest. Van Soest.07 12. Schweizer.11 23. Williams Co Nature’s Bounty Hi-Health Maker 658 Fiberguard Fibermed Wafers Fibretrim Fibretrim Fibretrim w/Calcium Full of Fiber Grain ‘N Citrus Hi-Fiber Metamucil Regular Metamucil Regular Sugarfree Serutan.85 19.-G.10 97. (C) Soluble fiber determined directly or by the difference (TDF-NDF corrected for residual protein).) 1.02 1. P. and Harland. L. T.43 4. Furda. HC = Hemicellulose. Asp. A.58 49. 67.50 11.25 13. Marllett.19 8.95 TDF 23. DeVries.53 9. New and improved procedure for neutral-detergent fiber.39 21.2 (Continued) Dry Matter Content of Selected Foods 2387_chzAppendix_T2_fm Page 658 Sunday. D. J. The detergent system of analysis and its application to human foods.72 4. N. Ls = Klason lignin. W..04 17.. W. AOAC. New York.-G.40 6.50 43.86 92. 1984.. James.54 CP 59. F. I. J.26 2. N. 351.70 51..18 93. and P.18 25. soluble.97 14.69 44.83 ASH 13. A.18 9. O.92. Regular Serutan.36 0.02 2.09 5. and total dietary fiber in foods and food products.58 3. L. 66 (Suppl.93 12. May 6.68 3. J.76 1. J.36 6.41 25.04 40.10 % DM 15. Sci.12 0. Prosky.80 94.40 0.20 8.07 6.54 HC 19. J. H..96 52. and Theander. 71. in The Analysis of Dietary Fiber in Food. Toasted Slim with Fiber Ultra Plan Fiber Supplements Description Table A.03 6. B.34 44.13 92..14 6.86 23.82 49. B.26 16.51 36.22 44. Determination of total dietary fiber in foods. P.89 8.76 24.64 In Dry Matter (%) SF NDR 18.21 3. B. Jeraci. J.72 45. 1988. J.55 10.01 35. W. Williams Co J..88 8. I.. Asp.43 28.51 44.00 8. J.18 5. and Van Soest.08 17.27 38. Eds..59 98. SF = Soluble fiber.75 6.20 5.85 3...83 90.28 14. M. 2001 8:10 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.52 6.92 2.48 35.31 84. 1981.03 6.32 8.34 42. Anim.85 6.03 6.65 9.99 95. CE = Cellulose.40 2.51 14.25): TDF = Total dietary fiber. Robertson. F.25 34.16 4. T.05 95. food products and total diets: interlaboratory study. NDR = Neutral detergent residue (insoluble fiber)..80 9. 4. 123. Interlaboratory study.31 10.53 5.51 52. The majority of these analyses were supported by the National Cancer Institute Contract N01-CN-45182. AOAC.65 16.58 6.58 4...40 16. T.. L. METHODS: (A) Total dietary fiber values obtained by the methods of: Prosky. T.07 0.09 1. 3.71 48.32 1.20 15. Marcel Deckker. CP = Crude protein (N × 6. DM = Dry matter. Schweizer.36 16.66 93.00 41..86 90.. 1044.40 2.41 12.33 21. J. W. B.. Ayerst Purdue Fredrick Schering Schering Schering Jameson Nutritional Hilstone Walgreen Laboratories Searle Searle J.25 CE 3.80 94.04 30.45 8. Lewis..57 5.14 0. 1017.77 9. 3RD EDITION . DeVries.91 3.88 46. W.05 –0. and Furda. P.26 42. Hernandez.71 92.98 46. J. 1988.85 Ls Notes: 1. Determination of insoluble. 2. J. Hurt. Southgate 0-8493-2387-8/01/$0. T.3 Dietary Fiber Content of Selected Foods by the Southgate Methods David A. 2001 8:12 PM Appendix — Table A. May 6.00+$1.2387_chzAppendix_T3_fm Page 659 Sunday.50 © 2001 by CRC Press LLC 659 . 07 0.86 2.3 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.09 0.16 1.20 11.99 2.09 2.13 1.55 0.21 3.00 7.54 8.50 3.90 2.99 1.73 4.48 1.22 2.92 1.26 5.02 1.80 5.70 Tr Tr Tr Tr 3.19 5.28 0.49 2.90 2. 18 6.71 1.48 3.53 2.32 Tr .40 2.14 5.70 6.28 7.10 2.01 2.33 0.35 1.55 1.72 5.47 1.42 2.51 2.69 1.07 0.47 15. bread-making Brown Whole meal Breads White Brown Hovis Whole meal Breakfast cereals All Bran Cornflakes Grapenuts Readibreak Rice Krispies Puffed Wheat Sugar Puffs Shredded Wheat Special K Swiss breakfast (mixed brands) Weetabix Cookies and crispbreads Chocolate digestive (half-coated) Chocolate (fully coated) Crispbread.63 3.75 6.30 5.60 4.34 1.32 0.99 3.03 0.63 0.70 4.85 4. young (boiled) Parsnips (raw) Swedes (raw) Turnips (raw) Potato Main crop (raw) Chips (fries) Crisps Canned Total Dietary Fiber Noncellulosic Polysaccharidesa Celluloseb Ligninc 3.72 17.78 1.39 3.74 0.19 0.60 1.32 1.07 4.66 0.23 1.15 0.59 0.36 8.87 9.85 9.35 4.03 0.00 8.72 3.24 0.38 Tr Tr Tr 7.58 1.11 4.79 3.47 2.31 1.42 1.13 1.01 3.47 0.22 0.05 10.83 1.82 7. May 6.78 2.40 0.08 12.50 1.08 0.30 0.03 0.80 0.83 1.28 Tr 0.42 1.70 0. wheat Ginger biscuits Matzo Oatcakes Semisweet Short-sweet Wafers (filled) Leafy vegetables Broccoli tops (boiled) Brussels sprouts (boiled) Cabbage (boiled) Cauliflower (boiled) Lettuce (raw) Onions (raw) Legumes Beans.99 0.30 1.66 1.94 0.36 2.20 6.45 7.41 12.09 11.84 1.67 1. baked (canned) Beans.88 1. 3RD EDITION Dietary Fiber Content of Selected Foods by the Southgate Methods (Grams Per 100 g Edible Part) Flours White.50 2.15 0.40 1.35 2.68 5.47 1.33 3.51 2.15 7.00 7.46 0.52 5. 2001 8:12 PM 660 Table A.31 Tr 0.51 3.75 0.85 5.31 1.01 0.21 0.76 1.33 1.60 2.41 1. frozen (raw) Garden (canned)d Processed (canned)d Peanuts Root vegetables Carrots.95 0.61 1.29 2.13 0.44 0.24 26.70 11.80 1.22 3.67 0.18 0.76 0.06 0.69 0.42 1.2387_chzAppendix_T3_fm Page 660 Sunday.62 2.04 1.12 1.59 0.13 0.19 3.80 2.31 9.79 0.47 10. rye Crispbread.27 3.43 0.11 0.05 0.26 5.22 0.10 2.77 1.72 1.45 2.35 7.20 2.74 1.00 2.41 6. runner (boiled) Peas.25 0.55 0. 89 1.04 0.03 0.3 (Continued) Dietary Fiber Content of Selected Foods by the Southgate Methods (Grams Per 100 g Edible Part) Peppers (cooked) Tomato Fresh Cannedd Sweet corn Cooked Cannedd Fruits Apples Flesh only Peel only Bananas Cherries (flesh and skin) Grapefruit (canned)d Guavas (canned)d Mandarin oranges (canned)d Mangoes (canned)d Peaches (flesh and skin) Pears Flesh only Peel only Plums (flesh and skin) Rhubarb (raw) Strawberries Raw Cannedd Sultanas Nuts Brazil Preserves Jam Plum Strawberry Lemon curd Marmalade Fruit mincemeat Peanut butter Pickle Dried soups (as purchased) Minestrone Oxtail Tomato Beverages (concentrated) Cocoa Drinking chocolate Coffee and chicory essence Instant coffee Extracts Bovril (beef extract) Marmite (yeast extract) a b c d Total Dietary Fiber Noncellulosic Polysaccharidesa Celluloseb Ligninc 0.85 0.03 0.91 0.91 0.70 0.73 3.34 Tr 1.73 15.30 0.94 1.20 0.65 1.02 0.01 0.15 Tr 0.46 0.33 1.21 1.40 0.62 2.45 0.05 0.37 0.19 7.93 0.04 0.03 0.32 3.69 4.42 3.08 1.00 4.64 0.97 0.03 0.04 1.12 0.09 5.96 1.41 11.04 43.50 0.75 1.64 2.10 0.3 661 Table A.93 0.17 1.48 2.20 0.64 0.60 1.27 8.44 8.37 0.14 0.01 0.07 0. Expressed as glucose.98 0.61 0.28 0.45 0.91 2.32 0.84 3.48 1. Drained material.40 0. .22 0.83 0.60 2.12 1.94 2.18 0.31 0.55 1.67 0.80 0.33 0.69 0. This value includes heat-induced artefacts analyzing as lignin.29 1.52 1.59 0.24 0.74 5.95 1.45 2.49 0.79 16.06 Expressed as the sum of the component monosaccharides.12 6.53 0.15 2.17 7.40 0.59 1.16 0.71 3.64 0.13 1.50 Tr 0.85 2.67 0.43 0.53 27.17 0.03 4.23 0.60 0.65 0.85 0. May 6.34 1.92 0.91 0.71 1.32 4.03 0.12 0.61 3.25 2.12 0. 2001 8:12 PM APPENDIX — TABLE A.81 0.31 4.02 0.72 0.44 3.55 4.06 0.67 2.99 0.03 0.33 0.01 0.25 0.01 0.96 0.33 0.30 0.26 0.11 0.20 0.60 2.2387_chzAppendix_T3_fm Page 661 Sunday.90 4.78 1.20 0.18 0.80 0.00 2. . REFERENCE Southgate. J.. Collinson. D. T. . Nutr. 1976. 2001 3:11 PM 662 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.3.1 Percentage composition of the noncellulosic polysaccharides in wheat flours. A.. A. and Walker. 3RD EDITION Figure A. E.2387_chzAppendix_T3_fm Page 662 Tuesday. Hum. May 8. B. and fruits.. F. vegetables. Bailey. 303.. A guide to calculating intakes of dietary fiber. 30. 00+$1.50 © 2001 by CRC Press LLC 663 .4 Dietary Fiber Content of Cereals in Norway Wenche Frølich 0-8493-2387-8/01/$0.2387_chzAppendix_T4_fm Page 663 Sunday. May 6. 2001 8:13 PM Appendix — Table A. 9 9.7 9.1 3.9 18.7 5.2 23.7 7.8 8.9 43.6 10.3 3. wheat flour Macaroni.2387_chzAppendix_T4_fm Page 664 Sunday.7 22.1 11.2 3.4 3. ..7 15.3 11.0 23.5 13.5 5.1 8.1 3.5 11.2 10.7 10.0 3.7 4.1 16. health shop Kavli flatbread Kavli breakfast flatbread Kavli flatbread with bran Kavli thin crispbread Ideal flatbread Ideal homemade Ideal fiber flatbread Spaghetti.0 3.4 2.5 9. whole wheat flour Rice.7 50.5 38.2 2. round-grain rice (health shop) Unpolished.6 22.6 21.2 4.4 8.0 21.7 10.4 6.2 25.8 3.7 5.1 6.4 8.4 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. Rapid enzymatic assay of insoluble and soluble dietary fiber.1 12.0 4.5 18.7 8.0 10. May 6. long-grain rice (health shop) Nature.7 15.1 18.4 3. whole wheat flour Macaroni.6 8.7 4. 476.9 47. and Siljeström.-G.4 10. Food Chem.1 15.1 4.7 10. M..6 9. parboiled Rice.3 2. 2001 8:13 PM 664 Table A.0 7.8 3.1 10..0 REFERENCE Asp.1 10.5 4.2 3. Johansson.5 19..0 4.-G.0 45.9 35.2 6. 1983.1 9. brown rice Unpolished.2 24. barley Oat flakes Bran of wheat Bran-germ of wheat Germ of wheat Semolina Triticale Everyday breakfast cereal 4-Grain breakfast Bran-cracker type I Bran-cracker type II Bran-cracker type III Bran-cracker. nature rice Rice.9 3.6 2.4 2.8 34. porridge Rice. rye Barley flour (50% extraction) Barley flour (70% extraction) Whole grain.6 17.6 11.0 20. Agric.0 3. C.2 4.9 3.5 10.6 15.6 7.9 42. H.5 4.9 14. 3RD EDITION Dietary Fiber Content of Cereals in Norway White wheat flour (78 to 80% extraction) Whole grain flour Rye flour (mixed with 15% white wheat flour) Whole grain flour..8 13.8 11.3 13.8 10.9 9.8 11.4 4.9 7. Hallner. quick rice Rice. 31.2 4.4 10.0 10.1 10.0 36.7 57.1 3.5 8.2 11.7 11.9 12. wheat flour Spaghetti. biodynamic-grown rice (health shop) Dietary Fiber (wet weight %) Moisture (%) Dietary Fiber (dry weight %) 3. precooked Rice.9 8.0 7. N.0 10.6 10.9 11. 50 © 2001 by CRC Press LLC 665 . May 6.5 Crude Fiber Values of Typical Samples Ivan Furda 0-8493-2387-8/01/$0. 2001 8:14 PM Appendix — Table A.00+$1.2387_chzAppendix_T5_fm Page 665 Sunday. raw Onions.1 67.1 4.7 4.6 2.7 90.9 2.7 2.4 0. raw Crude Fiber (g/100 g) Moisture (%) Crude Fiber (moisture-free basis) (%) 1.2 0. raw Oatmeal.0 1.7 11.7 13. lima Beans. whole-ground Grapefruit.9 2.0 86. dried Bean flour.8 1.7 2. raw Plums.5 13.9 78.1 92.7 1. bitter Cocoa. edible podded. raw Peppers.4 0.0 1.6 10.7 10.5 90.3 88.2 1. defatted Pears.5 1.3 83.5 0.3 78.6 5.31 0.2 4. raw peeled fruit Peaches Peanuts.0 91. raw prune type Popcorn.2 1.6 0.6 0. fresh.2 9.4 4. raw Pineapple.4 0.4 1.7 3.1 1.5 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.1 5.3 7.4 0.6 91.5 4.5 84.5 1.2 10.6 4.9 8.4 0.6 3.4 2.0 12.8 74. raw Macadamia nuts Malt.0 12.3 0. lima. raw Carrots Raw Dehydrated Cauliflower.4 8.5 8.2 2.5 0.5 6. raw Olives.3 2.2 0. raw without skins Peanut flour.6 3.3 3.0 35.0 36.0 79.6 1.2 16.0 12.3 9. raw Carob flour Chocolate.3 1. dry powder Corn grits.4 81.6 35.2 21.3 0. green Onions.0 5. raw Bananas Breads White Rye French Pumpernickel Whole wheat Broccoli.5 0.1 88. raw Artichokes.7 2.8 1.2 0. raw Cabbage.0 2.1 1. field.5 4.1 . raw (including skin) Peas.4 2.5 5. plain Potatoes. popped. dry Mushrooms. 2001 8:14 PM 666 Table A.0 85. raw Grapes.3 1. whole grain.0 5.0 13.8 0.3 1.4 89.2 89. immature.0 2.4 2. dry Okra.7 2.6 74. raw Apples. raw Cornmeal.0 5.8 2. raw Horseradish.0 88.2387_chzAppendix_T5_fm Page 666 Sunday.2 83.8 7. sweet.8 7.4 7. May 6.5 9.4 1.0 6.4 2.5 30. dehydrated Oranges. green. raw Potato flour Pumpkin.5 6.6 1.7 2.6 4.5 8.4 2.7 1.4 85.6 2.2 1.0 1.5 0.6 1.4 11.2 0.6 2. 3RD EDITION Crude Fiber Values of Typical Samplesa Sample Avocados.2 2.6 8.3 1.0 1.4 0.0 9.6 34.0 89. raw Almonds. dry Corn flour Corn.3 93.6 5. 7 10.5 1.5 5.0 11.3 1.5 2.5 11.5 6.6 0. soft wheat Bread flour (enriched) Wheat bran (crude.7 2.0 73.4 2.3 0. whole grain Hard Red Spring White Wheat flours Whole (from hard wheats) 80% extraction Straight.3 2.6 93. 8.0 10. raw Walnuts.2387_chzAppendix_T5_fm Page 667 Sunday.0 12.9 11.5 2.0 90.3 9. D. Eds. Department of Agriculture.7 11.S. raw Rice bran Rye (whole grain) Rye flour (light) Rye flour (dark) Rye wafers (whole grain) Soybeans (mature seeds). enriched.0 11.5 2.1 0. raw Strawberries.0 9. B. brown. commercially milled) Wheat germ (crude.5 0. raw Rice.1 0.4 2.0 11. U. black Wheat.9 13.0 12.4 Adapted from Composition of Foods.C. and Merrill.4 2. raw Sweet potatoes.8 3.9 1. dry.7 0.3 5. raw Turnips..1 2. white (milled or polished).2 0. debittered) Yam.0 6.4 7.5 667 Table A.0 12.3 10.5 10. Washington. L.9 12.7 0.5 91.0 12.6 2.7 2.0 0.7 94.5 3. 2001 8:14 PM APPENDIX — TABLE A. raw Squash.5 0.9 12.1 2.45 0.7 8.0 8.9 70.8 1.5 0.6 2.5 (Continued) Crude Fiber Values of Typical Samplesa Sample Rice. A.3 1.6 1.9 2.2 4.0 12.6 1.0 11.3 0. commercially milled) Yeast (Brewer’s.. Watt. Agricultural Research Service. 1963.4 2. May 6. raw a Crude Fiber (g/100 g) Moisture (%) Crude Fiber (moisture-free basis) (%) 0.8 2.. Agriculture Handbook No. ripe. . raw Tomatoes. tuber.0 0.4 0. raw Soybean flour (full-fat) (defatted) Spinach. K.0 89.3 0. May 6.2387_chzAppendix_T5_fm Page 668 Sunday. 2001 8:14 PM . 50 © 2001 by CRC Press LLC 669 .6 Comparison of Analyses of Dietary Fiber and Crude Fiber Gene A.00+$1. Spiller 0-8493-2387-8/01/$0.2387_chzAppendix_T6_fm Page 669 Sunday. 2001 8:14 PM Appendix — Table A. May 6. 50 1.12 1.20 1. 3RD EDITION Table A.50 7. raw Parsnips. fresh Fruits Apples.50 0. baked Beans.50 1.53 2.60 7.15 9.40 0. white Bread.40 0.70 4.24 2. whole wheat Bran Bread. frozen.52 2.00 2.27 3.51 1.60 2.90 1. flesh only Bananas Cherries.80 0.40 0.30 1.50 0.74 1. flesh and skin Strawberries. raw Tomatoes. white Flour.70 0.42 1. canned Potatoes.41 0.15 44.40 0.75 1.6 Comparison of Analyses of Dietary Fiber and Crude Fiber Foods Vegetables Beans.4) and crude fiber for various foods showing lack of correlation between the two measurements. 2001 3:12 PM 670 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.70 11. green Carrots. Chapter 3.55 0. whole wheat All Bran cereal Corn Flakes Figure A. sweet. raw Onions.00 0.30 2.70 3.1 Crude Fiber1 (%) Total Dietary Fiber2 (%) 1. .6. flesh and skin Pears.10 4.30 9. May 8.75 7. raw Nut-like products Peanut butter Grains and grain products Flour. raw Peas.00 1.90 7.00 Data for dietary fiber (Southgate method.35 3.60 1. cooked Lettuce.07 3. flesh and skin Peaches.2387_chzAppendix_T6_fm Page 670 Tuesday.90 7. flesh only Plums. raw Peas.60 0.60 0. cooked Corn.50 26.28 2.72 8.00 1.44 1. Handbook Number 8. A.2387_chzAppendix_T6_fm Page 671 Sunday. ...S.6 671 REFERENCES 1. Collinson. 2. D. 30. Dietary fiber data from Southgate.. 1963. Crude fiber data from Composition of Foods. 303. May 6. Nutr. Washington.C. U. 1976. Department of Agriculture. Bailey. E. J. F. and Walker. D.. 2001 8:14 PM APPENDIX — TABLE A. A. B. T.. A guide to calculating intakes of dietary fiber. Hum.. 2001 8:14 PM .2387_chzAppendix_T6_fm Page 672 Sunday. May 6. 2001 8:15 PM Appendix — Table A. Harland 0-8493-2387-8/01/$0.2387_chzAppendix_T7_fm Page 673 Sunday. May 6.7 Phytate Contents of Foods Barbara F.50 © 2001 by CRC Press LLC 673 .00+$1. Jerusalem. mg 2387_chzAppendix_T7_fm Page 674 Sunday.5 10. lima. chicken-flavored (Wyler’s) Brazil nuts Bread.5 7.3 25.7 1/2 c 1 1 bud 1 Tbsp 1 1 1 Tbsp 1 Tbsp 1 Tbsp 1 oz 1/2 c 1/2 c 1c 1/2 c 1/2 c 1/4 c 1/2 c 1/2 c 1/2 c 1/2 c 1/2 c 1/2 c 1/2 c 2 cubes 2 cubes 1/2 c 1 sl 1 sl 1 sl 1 1 1 sl 1 sl 1 sl 2 sl Serving (Uc) 71 150 380 15 120 201 15 15 15 28 120 120 124 92 55 40 85 62 55 62 85 72 115 8 8 70 35 28 27 10 35 32 25 25 10 Size (g) 909 94 110 70 11 2 196 27 11 251 197 22 112 282 124 404 294 564 122 56 2 7 3 7 3 1259 6 65 21 65 43 34 14 39 18 Phytate per Serving (mg) 1280 63 29 468 9 1 1310 180 73 897 164 18 90 307 226 1010 346 910 222 91 3 10 3 88 32 1799 17 232 79 654 123 107 58 155 183 EPd. green. navy. canned.6 36. pinto. lima.4 80.7 82.5 91. high-fiber. white (Fresh Horizons) Bread.0 67.9 67. casserole with cheddar cheese Beans.7 70. beef-flavored (Wyler’s) Boullion cubes.9 90. raisin (Giant) Bread. canned.0 4.4 35. mature. dry. broad.3 4.1 5. May 6.0 7. dry.0 10. immature. raw Beans. pearl. canned.0 35. whole (S & W) Avocado Bacon chips.g green. raw.Moistureb (%) 5. American (Giant) Bread. buttermilk (Bimix.2 10. wheat (Fresh Horizons) Bread.0 34.5 6. dry (Gerber) Barley.3 35. Borden) Phytate Contents of Foodsa Food Table A. canned. 2001 8:15 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. high-fiber. boiled Beans.6 83.9 90. navy. drained Beans. boiled Beans. dry.5 30. instant cooking. raw Beans. mature. not pared Artichoke. sliced (Del Monte) Blackberries Blueberries. wheat (Melba Toast. drained Boullion cubes. pumpernickel (Giant) Bread. drained Beans.3 69. raw Beans. buttermilk (Bisquick) Barley. Norwegian flat (Kauli) Bread.0 84. drained Beets. raw Beans.0 Almonds (Taylor’s Sunshine Colony) Apples. flour Artichoke hearts. Jerusalem.0 72. 3RD EDITION . rye. Martha White) Baking mix.5 10.3 69.0 8.8 4. snap.0 66. boiled Artichoke. kidney. Betty Crocker) Baking mix. pita (Giant) Bread.0 69. mature. sweetened. mg 347 404 146 525 88 3 1379 195 79 1000 539 110 300 990 695 1126 1116 1021 684 1123 30 56 11 92 33 1966 24 365 123 683 164 162 89 240 195 2h 1h 3h 2h ± 5e ± ± ± ± ± 4h ± 2e ± 2e ± 7e ± 18e ± 5f ± 2h ± 9f 24f 2c 2c 9e 14e 7e 674 ± ± ± ± ± ± ± 17c 100 g Dry Wt. French Bread. imitation (Bacos. infant cereal. boiled. enriched (Giant) Bread.8 12.7 90 56 20 52 28 38 20 28 80 60 40 225 225 5 28 42 180 180 180 104 28 8 81 70 70 80 100 62 27 28 24 28 44 52 120 2387_chzAppendix_T7_fm Page 675 Sunday. without icing (Duncan Hines Swiss Choc. chocolate. fresh Bun.8 33. dry powder (Hershey) Coconut meat. instant (Tasters Choice 100% Freeze-Dried) Coffee substitute. honey-roasted (Eagle Snacks) Chestnuts Chickpeas (garbanzos).2 5. baked (Duncan Hines) Cookie mix.8 2.7 10 17 127 1019 9 1866 950 47 730 208 69 390 334 365 18 82 531 2e 2e 0e 0e 6h ± 9h ± 31h ± 20e ± 12h ± 11h ± ± ± ± ± 340 40 38 200 368 104 161 219 ± ± ± ± ± ± ± ± 10h 6h 3e 6h 10e 3e 3e 7e 112 ± 9f 161 ± 3e 190 ± 6h 284 ± 10e 280 ± 3e 1960 237 64 ± 6h 368 37 526 21 128 1078 76 1968 1000 97 817 670 107 613 529 397 164 124 701 APPENDIX — TABLE A. dry.9 8. Nabisco) Cookie (Nilla Wafers. Sara Lee) Coffee.9 5.8 11.2 3. Nabisco) Cookie. white.0 4.0 sl sl sl sl oz bud 1 1 1 1 1 1 1 35. oatmeal and raisin (Pepperidge Farm) Cookie (Peanut Butter Nut. May 6.3 22.1 4.5 88. Nabisco) Cookie.1 3. Duncan Hines) Cookie (Toddler Biter Biscuits.0 98.7 10. mature seeds.2 294 20 7 101 92 38 31 55 10 89 74 187 169 94 65 21 12 1 18 18 36 81 8 1306 665 38 730 129 18 109 80 102 8 43 637 327 35 37 194 328 100 153 197 12 148 186 83 75 1880 230 50 7 0.1 98.1 98.2 5. whole wheat (Oroweat) Breading mix (Shake n’ Bake) Broccoli. brewed (Folgers Flaked) Coffee. drained Chili con carne.3 24.0 2. plain. no beans (Wolf) Chili con carne.0 51.0 88. shredded Coffee cake (Butter Streusel. dry.4 36. peanut butter (Duncan Hines) 1 sl 1 oz 1 Tbsp 1 1/2 c 1/2 c 1/2 c 1/2 c 1/2 c 1c 1c 1 Tbsp 1 oz 1 sl 6 oz c 6 oz c 6 oz c 1c 4 4 10 4 2 4 2 2 1 1 oz 20. raw Cashew nuts Cashew nuts.7 73. chocolate chip (Chips Ahoy. 2001 8:15 PM 675 . boiled. lightly iced (Duncan Hines Mix) Candy.Bread.1 88. hamburger (United Premium Quality) Cake. coconut bar (Rippin Good) Cookie.0 10. Mix) Cake. grain and spice (Celestial Seasonings) Collard greens. white. fig bar (Fig Newtons. dried. milk chocolate (Hershey) Caraway seeds Carrots.7 69.9 8. whole wheat (Giant) Bread.8 36.0 0.0 70. raw Chickpeas or garbanzos. brownie. Nabisco) Cookie. sweetened. Gerber) Cookie mix. chocolate sandwich cream-filled (Oreos.0 11. w/ beans (Wolf) Cocoa.7 2. raw Cookie. 0 4.9 Moistureb % 78 28 122 180 80 15 95 1260 311 344 2 11 1 412 26 65 20 26 20 65 178 142 142 2048 754 107 20 8 50 11 19 95 79 15 117 5 343 956 175 489 Phytate per Serving mg 1615 1112 282 1 14 7 434 31 232 70 94 73 232 635 506 507 1707 943 711 16 84 179 103 172 317 141 28 366 4 385 1620 625 627 EPd. chopped Granola. uncooked Filberts (hazelnuts). Nabisco) Cracker.0 54. Nabisco) Croutons. sugar-coated (Mrs.0 7.6 3.8 3. tomato (Heinz) Lentils.3 87. saltine (Nabisco) Cracker (Wheat Thins.9 12. instant (Swiss Miss Hot Cocoa Mix) Kale.3 6.0 5.0 12.2 Serving Uc 53. cake.3 10. whole ground corn meal (Washington Combread Mix) Corn. animal (Barnum’s. drained solids Corn cereal.0 3. raw Food Table A. toasted (Brownberry) Cucumber.0 4.4 6. Nabisco) Cracker. 3RD EDITION . dried. ready-to-eat (Corn Bran.9 3. canned. stoneground (Stone Mountain) Corn puddingi Cracker.8 6.7 65. Nabisco) Cracker (Ritz. ready-to-eat (Corn Pops. mg 2e 3e 6h 6h 0e ± ± ± ± ± 3e 2e 5e 15e ± 57h ± ± ± ± ± 3e ± 5e ± 3e 16h 3e 10e 3e 18e ± ± ± ± ± 1670 1240 2186 ± 38h 53 128 ± 7f 20 ± 3e 494 129 240 72 97 78 244 660 538 570 1940 1072 808 24 89 190 107 180 337 152 62 481 38 500 1720 650 1360 100 g Dry Wt. graham (Honey Maid. degermed.0 11. cooked (corn grits). 2001 8:15 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.1 88. mg 676 82 28 28 28 28 28 28 28 28 120 80 15 122 10 28 11 11 30 56 52 32 122 89 59 28 78 Size g 2387_chzAppendix_T7_fm Page 676 Sunday. Kellogg’s) Corn chips (Cheetos) Corn chips (Cornquistos. raw Ketchup.0 6. shelled.0 89. and raisins (Nature Valley) Hickory nuts High-protein infant cereal. whole-kernel. coconut.7 (Continued) Phytate Contents of Foodsa 1 sl 1/2 c 1 oz 1 oz 1 oz 1 oz 1 oz 1 oz 1 oz 1 oz 1c 1/2 c 1 Tbsp 1/2 c 4 4 4 4 2 2 oz 1/2 c 1 1/2 c 1/2 c 1/2 c 1 oz 1/2 c 1 oz 1/2 c 6 oz c 1c 1 Tbsp 1/2 c 75. assorted nuts.0 34.0 12.8 3.0 5. May 6. enriched Hot chocolate. Quaker) Corn cereal.1 24.Cornbread.0 12. Snack Master) Corn chips (Doritos) Corn chips (Fritos) Corn chips (Nacho Dorritos) Corn germ flour (Quaker) Corn meal (Quaker) Cornmeal. Baird’s) Farina. regular.5 23.5 2.1 98. instant dry form (Gerber) Hominy. cooked (Cream of Wheat) Figs. yellow.0 6. raw Doughnut. Kellogg’s) Corn cereal. unbolted. ready-to-eat (Corn Flakes. 2001 8:15 PM APPENDIX — TABLE A.0 66.Macaroni. shelled Piecrust stick (Betty Crocker) Plantain. Spanish w/pimiento Olives. chow mein (La Choy) Oatmeal or rolled oats. dry form (Gerber) Okra. drained solids Peas.2 2. instant. green. toasted.6 2.9 10. instant. blackeyed.0 35.8 1.9 74. dry (Quaker) Oatmeal. ready-to-eat (Special K. raw Peach pie (Giant) Peanuts.3 2. green. sliced Pancakes.0 86. Nabisco) Mixed grain cereal.5 6. raw Peas. plain Popcorn. unpopped (Orville Redenbacher’s Gourmet) 1/2 c 1/4 c 1 oz 1 oz 1 oz 1 oz 1 oz 1 oz 1 oz 1 1 1 oz 1/2 c 1 oz 1 oz 1/2 c 1/2 c 2 2 1 1c 1 sl 1 oz 1 Tbsp 1/2 c 1/2 c 1/2 c 1/2 c 1/2 c 1/6 1/2 c 1c 1/8 c 10.4 4.9 85.7 3.1 50.5 11.4 19. cooked (Quaker) Oatmeal.8 67 118 28 16 100 100 85 100 54 53 50 6 6 28 28 66 40 28 120 28 28 80 48 40 48 28 28 28 28 43 25 28 28 21 4 261 200 815 851 24 664 793 62 16 37 34 65 203 48 199 114 133 264 251 4 15 1 1 244 76 76 45 112 124 143 50 32 3 933 1252 815 851 28 664 1468 117 32 614 561 232 726 73 498 409 111 943 897 5 32 3 2 871 272 272 162 260 494 510 177 ± ± ± ± 5e 8h 7e 0e 137 6 999 1280 911 982 151 752 1519 144 95 640 622 240 809 120 767 413 822 1035 1000 88 286 11 15 1763 2e 4e 3e 2e ± 13e ± 3e ± 16f ± 2h ± 7h ± 22e ± 3e ± ± ± ± ± 6h ± 8h ± 2e 278 ± 8e 281 ± 2e 166 ± 5e 290 560 517 181 2387_chzAppendix_T7_fm Page 677 Sunday. infant cereal. Post) Mixed grain cereal.5 13. dry Pecans. salted (Tom’s) Peanut butter (Jif. dry form (Gerber) Muffin. elbow.0 9. ready-to-eat (Team.6 76.2 3. Kellogg’s) Mixed grain cereal. infant. split. dried.8 88. immature. canned. ready-to-eat (Alpha Bits. Kellogg’s) Mixed grain cereal. ready-to-eat (Froot Loops. English (Thomas) Muffin.9 47. dry Mixed grain cereal. raw Peas. May 6. raw Popcorn.2 10. popped. extra crunchy) Peas. ready-to-eat (Product 19. cooked (Trappey’s) Okra.3 94. dried. enriched (Kroger) Millet. wheat bran (Duncan Hines Mix) Noodles.4 11. wheat germ (Aunt Jemima Mix plus 1 teaspoon of wheat germ/pancake) Parsnips.7 677 .5 8. Kellogg’s) Mixed grain cereal.1 1. Kellogg’s) Mixed grain cereal.3 2.3 39.3 81. ready-to-eat (Apple Jacks.5 3.2 10. ripe. raw Olives. 0 7. mg 2580 1360 1260 1420 1630 2316 200 401 181 488 111 1998 108 577 159 284 2200 139 193 150 212 980 93 1081 1710 270 120 62 951 1151 2e 2e 2e 10e ± ± ± ± ± 2h ± 14h 6e 15h 15e 22h 7e ± ± ± ± ± ± 1h ± 13h ± 15h 100 g Dry Wt. mature seeds. dry form (Gerber) Roll.0 Moistureb % 2387_chzAppendix_T7_fm Page 678 Sunday. 3RD EDITION . instant. drained. brown. wild Rice cereal.0 7. Post) Rice cereal. white.0 7.6 10. General Foods) Rice. dry Rice.0 7. Deli) Pretzels. fully milled or polished Rice. 2001 8:15 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.0 5. pared Potatoes.0 5.Poppy seeds Potato chips Potatoes. mg 678 7.0 7. boiled in skin.4 15. infant.5 2. ready-to-serve. May 6. dry form.5 94. ready-to-eat (Fruity Pebbles. Kellogg’s) Rice cereal.7 (Continued) Phytate Contents of Foodsa 10. ready-to-eat (Cocoa Pebbles.9 2.3 10.8 44.5 1.3 9. butter (Seyferts) Pumpkin seeds Radish.0 87. dry. uncooked (Minute. French fries Potato salad w/ egg (Giant.5 7.8 79. Concentrate) Soy-based TVP bacon (Archer Daniels Midlands) Soy-based TVP bacon and vitamins (Archer Daniels Midlands) Soy-based TVP beef (Archer Daniels Midlands) Soy-based TVP ham (Archer Daniels Midlands) Soy-based TVP pork (Archer Daniels Midlands) Soy-based TVP unflavored and vitamins (Archer Daniels Midlands) Soybeans.3 12. plain (Giant) Rye flour (Pillsbury) Sesame seeds Soy-based chicken analog (General Mills) Soy-based ham analog (General Mills) Soy-based infant formula (Advance. Post) Rice cereal.0 2. regular.0 oz oz oz oz 1/2 c 1 1 1 1 1 Tbsp 1 oz 1/2 c 10 1/2 c 10 1 oz 1 1/4 c 1/4 c 1/4 c 1/2 c 1 oz 1 oz 1 oz 1 oz 1 oz 1 1c 1 Tbsp 1 oz 1 oz 4 oz 1 oz 1 oz Serving Uc 105 28 28 28 28 8 28 78 50 75 60 28 45 25 45 25 100 28 28 28 28 28 28 128 8 28 28 120 28 28 Size g 2438 354 328 370 424 175 55 63 50 63 63 529 3 130 65 64 1936 38 53 41 58 246 18 1176 129 70 31 10 248 300 Phytate per Serving mg 2322 1265 1172 1321 1516 2189 196 81 100 84 105 1889 6 518 144 255 1936 136 189 146 207 879 64 919 1616 251 112 8 884 1070 EPd. long-grain. ready-to-eat (Cocoa Krispies.3 31. raw Elton variety Food Table A.1 2. ready-to-eat (Rice Krispies.8 5.7 82. Kellogg’s) Rice cereal.0 7. fresh Rice.4 10.0 5. 2 3. brewed (Kaffree. ready-to-eat (Bran Flakes. ready-to-eat (Raisin Bran. black.8 93. General Mills) Strawberries. Kellogg’s) Wheat cereal. instant (Nestea 100% Instant) Tomato seeds Tomato soup (Campbell’s) Tomatoes. Post) Wheat cereal. ready-to-eat (Frosted Flakes. shredded (Nabisco) Wheat cereal.2 3. Post) Wheat cereal. frozen.5 oz oz oz oz 1 oz 1c 1c 1 oz 1/2 c 1 Tbsp 1 1 6 oz c 6 oz c 1 Tbsp 1/2 c 1/2 c 1 Tbsp 1/2 c 1/2 c 1/2 c 1 oz 1 oz 1 oz 3. Bran Chex (Ralston Purina) Wheat cereal. raw Walnuts. ready-to-eat. ready-to-eat.6 28 28 28 28 28 28 28 28 28 28 28 137 137 28 128 8 180 20 180 180 8 122 120 15 65 62 50 28 28 28 411 77 22 184 415 186 51 151 298 16 375 1915 1689 399 8 128 9 110 3 2 148 8 8 90 4 1226 380 843 305 887 1467 274 80 659 1481 666 182 541 1066 58 1341 1398 1233 1425 6 1606 5 549 2 1 1847 6 6 597 6 1977 760 3011 1088 3168 ± 10e ± 2h ± 0f ± 3e ± 8h ± 8e ± 2h ± 4h ± 11h ± 11h 1520 278 ± 7e 83 ± 0e 719 ± 2e 1530 690 ± 10h 188 ± v8e 560 ± 3e 1099 ± 2e 60 ± 2e 1390 ± 22h 1520 1340 1500 21 1699 17 572 333 167 1955 49 95 670 47 2040 993 3360 1122 3286 2387_chzAppendix_T7_fm Page 679 Sunday. dried leaves) Tea. fortified (Special K. ready-to-eat (Super Sugar Crisps.4 5.Soy flour (Ralston Purina) Soy isolate (Ralston Purina) Soy.3 3.1 23. sweetened.j Wheat cereal. General Mills) 1 oz 1 oz 1 oz 1 oz 1 1 1 1 1 oz 1 oz 3.5 1.7 679 . English. textured concentrate (Patti Pro. shelled Wheat bran.0 5.5 10. May 6. 2001 8:15 PM APPENDIX — TABLE A.7 11. shelled Walnuts.5 8. Kellogg’s) Wheat cereal. drained Sunflower seeds Sweet potato. ready-to-eat (Grape Nuts Flakes. raw Taco shells (Paco’s) Tea. crude AACC ref.5 70. Kellogg’s) Wheat cereal.0 99. ready-to-eat. ready-to-eat (40% Bran.0 3.5 3. ready-to-eat (Honey Smacks. Kellogg’s) Wheat cereal.0 87.5 3. Kellogg’s) Wheat cereal. ready-to-eat 100% bran (All Bran.4 99.2 3.0 71. ready-to-eat (Post Toasties) Wheat cereal.3 5. Kellogg’s) Wheat cereal.4 3.0 8. Post) Wheat cereal. canned solids and liquid Triticale flour Turnips.0 4.4 3.5 87. ready-to-eat (Wheaties.0 2.0 8. and Oberleas.5 73. unbleached (Gold Medal. F. World Review of Nutrition and Dietetics.j i h g f e d c b a 1c 1 Tbsp 1 Tbsp 1 1/4 oz 12. Mean ± SE of triplicate determinations by Harland et al. baker’s dry Food Table A. bake at 350°F for 30 min. Mean ± SE of triplicate determinations by Harland and Oberleas (1986). and sl = slice. 1c 1 Tbsp Serving Uc 12.0 120 6 15 180 7 137 15 Size g 1014 244 38 94 35 386 20 Phytate per Serving mg 845 4071 252 52 495 282 136 EPd. One c whole-kernel corn. General Mills) Wheat flour. drained. 1/2 c grated cheddar cheese.0 11. Mix ingredients. c = 8-oz cup. (1986). 235. 1 c sour cream. 1987. May 6. This food has been certified to contain 3 ± 0. Two c snap green beans. 52. whole wheat (Pillsbury) Wheat germ (Kretchmer) Wheat gluten flour Yam. 1 can Campbell’s Cream of Mushroom Soup. If no value was given. mg Table 1 is taken in part from Harland.0 5.0 12.2% or 3348 ± 223 mg phytate/100 g (dry weight). Basel.7 (Continued) Phytate Contents of Foodsa 2387_chzAppendix_T7_fm Page 680 Sunday. 3340 Pilot Knob Road. MN 55121). 1/2 c melted butter. Karger. B. 2001 8:15 PM CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. raw Yeast. Tbsp = tablespoon.0 Moistureb % 680 Wheat flour. St. 3RD EDITION . mg 960 4600 276 ± 4f 193 521 ± 7f 320 154 ± 2e 100 g Dry Wt. 2 c cream-style corn. all-purpose (General Mills) Wheat flour.5 8. a "best estimate" was calculated from existing data. Mean ± SE of triplicate determinations by Oberleas and Roy (1985). Edible portion. oz = ounce. Mix and heat. Phytate in Foods.. Vol. 1 pkg Jiffy cornbread mix. D. AACC Certified Food Grade Wheat Bran (10-4-77) used as a reference material during phytate analyses (American Association of Cereal Chemists. 2 beaten eggs. S. Paul. enriched. 0-8493-2387-8/01/$0. 2001 8:16 PM Appendix — Table A.2387_chzAppendix_T8_fm Page 681 Sunday. In sun-dried raisins all of the tartaric acid of the grape is present. as it affects colon function.8 Tartaric Acid Content of Foods Monica Spiller and Gene A.00+$1.50 © 2001 by CRC Press LLC 681 . In temperate climates only grapes and their products have reasonable amounts of tartaric acid. while in wine making some of the tartaric acid is lost. amounts that may have physiological significance. Spiller Few foods contain more than minimal quantities of tartaric acid (C4H6O6). May 6. Tartaric acid is of interest in fiber studies and in health. 2 g/100 ml 1.5–0.7 g/100 ml 2.0–3.55–0.8–2.9 g/100 g 0.2 g/100 g .5–2. May 6. 2001 8:16 PM 682 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.5 g/100 g 1.2387_chzAppendix_T8_fm Page 682 Sunday. 3RD EDITION Food Amount Grapes (fresh) Wine Sun-dried raisins Raisin juice concentrate Raisin paste 0. 9 Plant Foods That Contain Significant Levels of Saponins and Their Estimated Saponin Content David Oakenfull and John D.50 © 2001 by CRC Press LLC 683 . Potter 0-8493-2387-8/01/$0. May 6. 2001 8:17 PM Appendix — Table A.00+$1.2387_chzAppendix_T9_fm Page 683 Sunday. 2001 8:17 PM 684 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION.5 0.05–16 10–23 3 58 5.7–60 1.2387_chzAppendix_T9_fm Page 684 Sunday.1 0.4–0. May 6.7 .5 0.8–11 2–16 0. 3RD EDITION Table A.9 Plant Alfalfa sprouts (Medicago sativa) Asparagus (Asparagus officinalis) Broad bean (Vicia faba) Chickpea (Cicer arietinum) Green pea (Pisum sativum) Kidney bean (Phaseolus vulgaris) Lentil (Lens culinaris) Mung bean (Phaseolus mungo) Navy bean (Phaseolus vulgaris) Oats (Avena sativa) Peanut (Arachis hypogaea) Quinoa (Chenopodium quinoa) Sesame seed (Sesamum indicum) Silver beet (Beta vulgaris) Soy bean (Glycine max) Spinach (Spinacea oleracia) Sweet lupin (Lupinus augustifolius) Saponin Content (g/kg dry weight) 80 15 3.2–0.5–6 4.7–1.6–56 47 0.5–21 0. 287–313. 63–65 Englyst procedure for nonstarch polysaccharides. 138 Agarose. 279 protein digestibility and. 446. 277. 90 Uppsala method for total dietary fiber. 90. See also Phytate. 482 cardiovascular protective effects. 87–105 detergent systems. 557 American Association of Cereal Chemists (AACC). 567. 282 fecal composition studies. 554 American Diabetes Association. 16. 2001 8:07 PM INDEX A Acetate. 27 Almond. 391. 33 African dietary and morbidity patterns. 447 digestive enzymes and. polysaccharide. 97 phytate analysis. 51–60. 63 nitrogen excretion and balance effects. 24. 567. 198 Amylases. See also Agar. Alginates. 87–105. 88. 53–60. 52. 317 food product content. See also Starch Amylose.2387_Index_fm Page 685 Sunday. 84–85. See also Uppsala procedure Animal connective tissues. 324 fecal composition studies. 453–458. 99 Acid detergent fiber (ADF). 363–365. 570. 88 insoluble and soluble fiber. 27 antitoxic effects. See Enzymatic gravimetric methods historical perspectives. 554–555 American Cancer Society. 147 protein digestibility and. 24. 52–53 lignin. 483. 554 Aminopolysaccharide. 40. 554 American Diatetics Association. 455–456 685 . 54. 24. retrograde. 431. 278. Carrageenans Alginates. 216 fecal mutagens and. 557 American Heart Association. 20 modified carbazole method. 90–92 sugar and lipid removal. 637. 83–85. 55 reagent toxicity. 278. See Sprouts Algal polysaccharides. 90. 155 short-chain fatty acids and. 88. 64. 40 Amylose. 17 Animal fibers (chitin and chitosan). 666. 627 American Academy of Pediatrics. 54 saponins. 674 Amaranth. 27. 40. 136 protein utilization effects. 92 Amylopectin. 574–576 Agar. analytical methods protein hydrolysis. 23–25. 554 American Health Foundation (AHF). 138 protein utilization and. 153 Additives. May 6. 27 Agiolax. 569. 446 digestive enzymes and. 52. 27 antitoxic effects. 51. See also Southgate method starch. 27. 45–46 Antioxidants. 127–130 Southgate method for unavailable carbohydrates. 144. 486 fecal mutagens and. 45–46 Amoebic dysentery. 306 Alfalfa antitoxic effects. 369. See also Resistant starch Analytical methods AOAC approved methods. See also Short-chain fatty acids Acetyl bromide method. 209 nitrogen excretion and balance effects. 318 sprouts. 149 protein digestibility and. 561. 83–84 sample preparation. 280 Amyloglucosidases. 446 blood glucose and insulin responses and. 64 sample extraction. 87. 55. 70–81. 423. See also Englyst procedure enzymatic gravimetry. 278. 311 Alginic acid. 113–125. 446 certificated reference materials for Englyst procedures. 309 Uppsala method applications. 101. 324. 28. 260 glycemic index values. 447 carbohydrate hydrolysis. 12. 185–186. 374. 560 Autoimmune diseases. 660. 15. 365 Appetite and satiety. 317 iron absorption and. 632 Banana. 684 Atherosclerotic disease. 104 Basic fibroblast growth factor (bFGF) receptor. 101. 521. 35. 445–448 Association of Official Analytical Chemists (AOAC) approved methods. 55 Asparagus fiber values. 370 Avenanthramides. undesirable fiber effects. 260 phytate content. 632. 616 Bamboo shoots. 330. 339 glycemic index values. 335. 654. 453 Avocado. 482 antioxidants. 513. 670 diabetes treatment effects. 330 glycemic/insulinemic response and. 629 Arabinans. 576. 411 fecal composition studies. 307. 3RD EDITION phytic acid anticarcinogenic effects. 655. 257–267 antioxidants and. 330. 561. 684 Beet fecal composition studies. 217 fiber values. 519. 363. 650–658 Ash precipitation. 93. 544 Basil. 493. 179 Bacterioides. 339 glycemic/insulinemic response and. 328 fecal composition studies. 217 fecal microflora studies. 674 short-chain fatty acids and. 670 glycemic index values. 674 Arugula. 279 digestive enzymes and. 52. 70 crude fiber/dietary fiber comparison. 340 fecal microflora studies. 627 Artichoke fiber values. 261–262 effects of different fiber types. 2001 8:07 PM 686 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 156–157 Uppsala method applications. 339 phytate content. 129. 264 Barley. 258–259 morphological characteristics. 65 Ascorbic acid. neutral. 491 vitamin B12 and. 655 fructooligosaccharides. 258 probiotics and prebiotics. 453–454 Antitoxic effects. 457 whole grain anticarcinogenic effects. 629 Apricot. 263. 674 B Bacillus subtilis enzymes. 259 reactive nitrogen products. 263–266 enzymes. 364. 661. 237 fiber values. 12. 501 mineral bioavailability and. 338. 666. 508. 55. 34–36 Apple antitoxic effects. 103 Applesauce. 27 Arabinose. 143 protein utilization effects. 104 Uppsala method collaborative study. 335. 326. 374 mineral bioavailability studies. 265 Bagasse blood glucose and insulin responses and. selected foods. 27 Arabinogalactans. 87–105 Apoptosis. 632 . 260 saponin content. 53–60. 330. 522 Ash content. 666 fructooligosaccharides. 595 cholesterol-reducing effects. 334. See also Legumes Chinese consumption patterns. 178. 631. 129. and beneficial species. 666. 632. 216 fiber values.2387_Index_fm Page 686 Sunday. 583–590 Austria. 489–490 dietary characteristics and. 455 antitoxic effects. 674 saponin content. 515. 543–544 Appendicitis. 258. 264. 483. 670 glucose/insulin responses. 655. 381. 632. 486 Australia. 324 fecal composition studies. 632 Asbestos. 457 beneficial vs. 490–491 whole grain components. 654. 103. 261 harmful. 261 carcinogenesis and. 670 fructooligosaccharides. 55 Bacteria (fecal or intestinal microflora). 674 protein digestibility and. 278 glycemic index values. May 6. 262. 629. 261. 632 Beans. 520 phytate content. 264 Bagels. See also Vitamin C fecal mutagens and. 661 crude fiber/dietary fiber comparison. 141. 382 phytate content. processing effects. 105 Arabinoxylans. 487–488 Arrowroot. 666. 542 processing and. 436 Broccoli cancer risk and. Whole-grain bread. P. See Oat bran.2387_Index_fm Page 687 Sunday. 330 Bulgur fiber values. 656 glycemic index values. 633. 233–234 fiber values. 312 Brown rice. 661. glycemic index values. 492. 509. 306. 678 protein digestibility and. 482. 330. 667 Bristol diet. 330. 629. 385 nitrogen excretion and balance effects. 667 glycemic index. 595 colon cancer risk and. 33 French consumption patterns. See also White bread. Rice bran. 103 Breadsticks. 307. specific grains carbohydrate hydrolysis. 628. 235 fiber values. 670 dry matter content. 654. 261. 637. 334 Burkitt. 409–410 dietary fiber and adsorption/excretion. 142 protein utilization effects. 150 phytate content. 684 short-chain fatty acids and. 258. 409. 616 Breakfast cereals antioxidants. 670 food preferences. 235 food preferences. 483 intestinal microflora and. 311 Benzoates. 402. 485. See also Short-chain fatty acids cancer risk and. 660 Buckwheat. 457 crude fiber/dietary fiber comparison. 616–618. 490 Bile acids bran and fecal losses. 667 fecal composition studies. 678. 259. 101. 423 casein digestibility and. 456. 453 Berries fiber content. 674 Bifidobacteria. 660 phytate content. 679 US consumption patterns. 322 687 Uppsala method applications. 334. 561 Brazil nuts. 666. 674 Body weight. 650–651 fat and protein effects on glycemic responses. 33 antioxidants. 675 short-chain fatty acids and. 384. 264–267. 4. 664. 483–484 Blueberries. 630. 504–514. 36 Bioavailability. 674 protein digestibility and. 326 soluble fiber effects on glucose and insulin responses. 261 mineral excretion and. 674–675 refining and glucose and insulin responses.. 2001 8:07 PM INDEX glycemic index values. 521 neutral polysaccharides not recovered by Uppsala procedure. 431. 655. 183–184 Brans. 627 glycemic index values. D. 556 Breast cancer. 334 glycemic/insulinemic response and. 455 Chinese consumption patterns. 483. 492 Bowel habit. 674 Breads. See Mineral bioavailability Biotin. 334 phytate content. 489. 654 phytate content. historical interest in fiber. 369–370 dry matter content. 176 Blackberries. 279 diabetes and national flour changes. 262. 666 fiber values. 674 Black-eyed peas casein digestibility and. 279 crude fiber values. 660. 677 Blood pressure. 461 Butyrate. 33 glycemic index values. 104 Uppsala method collaborative study. 324. Wheat bran Brazil. 336–337 fecal composition studies. 127 viscosity effects on intestinal absorption. 618–625. 677. 631. 309. 166. 650–651. 598 glycemic index values. 5. 279 dry matter content. 633. 287–313. 143. 339 mineral bioavailability studies. 466. 279 crude fiber values. 151 phytate content. May 6. 184. 516 nitrogen excretion and balance effects. 651–654. 655. 339 grinding effects on glucose and insulin responses. 629. 660. 288 C Cabbage antioxidants. chronic disease risk and. 338 mineral bioavailability studies. 519 phytate content. 259. 228–229. 334 mineral bioavailability studies. 167 saponins and fecal excretion. 651–654 fecal composition studies. 38 fiber lipid-modifying effects and. 39 dietary fiber feeding duration and. 423 . 69. 532 Brewer's yeast. 93 phytate content. mineral. 157 saponin content. 279 Brussels sprouts. 455 carbohydrate hydrolysis. 670 dietary fiber values. 633. 317 fiber fermentation and. 517–519 phytate effects. 339 diabetes treatment. 104 Case-control study design. 485 short-chain fatty acids and. 148–149 protein digestibility and. 3RD EDITION fecal composition studies. 491–492 Candy and sweets. 499. 445–448. 363. 326 glycemic index values. 138 vitamin A absorption effects. 69 fecal mutagens and. cancer and short-chain fatty acids and. 312 Uppsala method applications. 321–322 particle size effects and. See also Cancer Cardiovascular diseases (CVDs). 97 Carbohydrate metabolism. 321–347. 217. 553 whole grain and fiber effects. 481. 279. See also Diabetes. 346. 24 crude fiber. 308 viscosity effects and nutrient bioavailability. 569. 21. 505. 223 fecal/luminal pH and. 477 dietary fiber and risk. 175 Carcinogens. 138 solubility. 24. 104 Cadmium. high-fiber diets. 332–333. 323–329 long-term effects of low-glycemic index diets. 303 fecal microflora study. 173–175. 174 Carotene. 142 Cantaloupe. 465 South African risk. 216. 462 Casein. 37 vitamin D absorption effects. 542 Canada. 217. 161. 330 short-chain fatty acids and. 35 Carrot antitoxic effects. 347 mixing fiber with food.2387_Index_fm Page 688 Sunday. Glycemic index. 2001 8:07 PM 688 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 363 whole grain and fiber effects. 655. 519–520 Caffeic acids. 491–492 FDA findings regarding dietary fiber health claims. 334. 239 laxative effectiveness. 345. 654 Caraway seeds. 507. 633 . 191 short-chain fatty acids and. 481. 453 Carrageenans. 509–516 in vitro binding studies. 339–341 whole grains and diabetes. 504. 474–475 estrogen metabolism and. 280 nitrogen excretion and balance effects. Insulin response blood glycemic responses to foods. 660. 280 nitrogen excretion and balance effects. See also Diabetes fat and protein effects. 370 long-term effects of fiber supplements. 670 glucose and insulin responses and. 467–472 risk factors. 391 Carbohydrates. 312 Uppsala method applications. 639–640. 299. 338 high-carbohydrate. dietary fiber effects. 103. 16 Carboxymethylcellulose fecal composition studies. 336 Cashew. 69 prospective epidemiologic studies. See Phytic acid. 666 nitrogen excretion and balance effects. 317–318. 342. 277. 69 phytic acid and. 103. 336–337 fiber and single test meals. 322–323 grinding effects. 70 Chinese consumption patterns. Glycemic response. specific neoplasms controversial issues. See also Coronary heart disease dietary fiber and risk and mortality. Glucose tolerance. 216. 487–488 Carbohydrate intake recommendations. 660. 675 Cassava. 489–490 insulinemic effects. 488 utility of glycemic index. 476 Italian study. protein digestibility and. 492–493 phytoestrogen effects. 655. 490–491 gastrointestinal effects. 39 Italian study. 570 Cancer. dietary fiber consumption patterns. 446. 531–544. 263 fiber values. 149 protein digestibility and. 488–493 antioxidant effects. 447 certificated reference materials for Englyst procedures. 666 digestive enzymes and. See also Colon cancer. May 6. 467–472 fiber type and. 675 Carbazole method. 151 short-chain fatty acids and. 331–332 satiety and. 321 food refining effects. 263 fiber values. 219 fecal microflora study. 630. 278. 370. 278. 28 digestive enzymes and. 595 fecal content and composition. 329–333 cooking/processing effects. 609 naturally high-fiber diets and. 482–486 Carob. 675 Canola. 503 bioavailability studies. 321–324 fiber viscosity and. 323–329. 633. 489 South African patterns. 491 Calcium. indigestible. 364 studies indicating no effect of dietary fiber. 166. 461–477. 609 naturally high-fiber diets and. 661. 616–617. 94. 178 Cocoa. 33 Chickpeas glucose/insulin responses. 520 national consumption patterns. 260 digestive enzymes and. 380. 661. 292. 403–412. 324. 675 College students. 36–38 Cholestyramine. dietary fiber consumption patterns. See Cancer. 173 Cell wall composition. 281. 306. 262. 39 Cereals. 135–136 protein utilization effects. 45–46. 298 Children and adolescents. 569 nicotinic acid absorption effects. 281 Chives. 303 fecal microflora and. 661. 655. 134. 412 insulin levels and. 481–493. 401–414. 675 Coffee. 661. See also specific fiber components viscosity effects on intestinal absorption. 593–595 Chinese vegetables. 265. 264. 294. 374. 280 Cinnamic acids. 650–658 food values by Southgate methods. 177 nitrogen excretion and balance effects. 324 Chromium. whole grain. 629. 35 Uppsala method applications. 676 Cholecystokinin (CCK). 253–256 dietary fiber and risk. 173 Cholesterol-modifying effects. 206–208 fecal/luminal pH and.2387_Index_fm Page 689 Sunday. 9. 489 . 595 fiber values. 461–477. 145–146 plant cell wall composition. 445 early fiber studies. 446. 633. breakfast. 280. 94 Cellulose. 278. 330. Uppsala procedure for measurement. 296. 637. 175–176 Cobalamin. 485 Coagulation factors. 447. 28 diabetes treatment effects. chronic diseases and. 423–426. 675 saponin content. bioavailability studies. 557–558. 499 Chronic diseases. 282 fecal bulk and composition effects. Coronary heart disease. 127 specific fiber types. 103 vitamin A absorption effects. 670 Chestnut. 660–662 glycemic/insulinemic response and. 559 China. 63 Cherry. 403–405 mineral bioavailability studies. 632–633. 455 Chinese consumption patterns. 152–153 pyridoxine bioavailability and. See Breakfast cereals Cereals. 522 Clostridium. 633 Chloride. 176 short-chain fatty acids and. 318 food values. specific grains Cetyltrimethylammonium bromide. 289. 558 Colon cancer. 660 Celery antioxidants. 11–12 protein digestibility and. Colorectal cancer critical fecal weight correlations. 675 Coffee cake. 423–424 fecal bulk and transit time and. 516. May 6. 633 Chitin. 311 solubility. "whole food" approaches. 675 Chewing. 299 Uppsala method applications. 448 crude fiber composition. See specific grains Cereals. 447. 328 dietary fiber definitions and. 260 Cauliflower. 339 glycemic index values. 24. 401 effective fiber dose and formulation. 45–46 Chitosan. Inflammatory bowel disease "magic bullet" vs. See also Lipid-modifying effects dietary factors portfolio. 475 dietary fiber anti-toxic effects. 482–483. See also Cancer. 509 Chocolate. 475–476 whole grain and cereal fibers and. 184. 28 additives. 675 Collard greens. 191. 303. 277. 485 Cholesterin. 278. 507. 656. 684 689 short-chain fatty acids and. 178 vitamin metabolism and. 104 Chicory. 489 Coagulation effects. 334 phytate content. 310. See Whole-grain foods. 297. 633. 2001 8:07 PM INDEX Cation-exchange. chronic diseases and Chymotrypsin. 465–468 total mortality. 656 Cellobiose. 174 vitamin B12 and. 384 lipid-modifying effects. mineral bioavailability and. Cardiovascular diseases. 23 antitoxic effects. 453 Citric acid. 402 potential mechanisms. 554 digestibility by microflora. Diabetes. 11–12 Cell wall porosity. 335. 265 fecal mutagens and. See Whole-grain foods. 403 meta-analysis. 481 risk factors. 402–403 saponins and. 514. 141 short-chain fatty acids and. 104 Cornbread. 69 lack of evidence from prospective studies. 330. 184 fiber values. 306. 578 Cotton seed fecal composition studies. See also Breakfast cereals dietary fiber and crude fiber analysis. 481. 461–477. 374. 279 Chinese consumption patterns. 309. 660 glycemic index. 660 glycemic index values. 328 fecal composition studies. 474 FDA findings regarding dietary fiber health claims. 553 vegetarian diet and. 187 South African morbidity patterns and. 438 Constipation. 382. 618 phytate content. 339 Coronary Artery Risk Development in Young Adults (CARDIA) Study. 676 refining effects on glycemic responses. 363 whole grain and fiber effects. 265 fecal mutagens and. 512. 191. 100 protein digestibility and. 426 French mortality. 3RD EDITION fecal mutagens and. 299. 465–466 prospective epidemiologic studies. 485 hypotensive effects. 601 intestinal microflora and. 374. 539. 489–490 naturally high-fiber diets and. 617. 488–493 antioxidant effects. 491–492 Colon diverticulosis. 518. 482–483 Costa Rica. 334 grinding effects on glucose and insulin responses. 482–486 antioxidant and phytochemical effects. 279 casein digestibility and. 326 Corn starch. 216 glycemic/insulinemic response and. 652 fiber values. 482 antitoxic effects. 69 possible mechanisms for fiber protective effects. 363. 541–544 recent fiber studies. 299. 670 dry matter content. 2001 8:07 PM 690 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 214. 503 bioavailability studies. 483–484 lipid-modifying effects. 365 Continuing Survey of Food Intakes by Individuals (CFII). 475 latency period. 488 Colorectal cancer Chinese mortality. 365 Corn bran diabetes treatment effects. 304 fecal microflora and. See Bacteria Colon polyps. 183. 307. 486 antithrombotic effects. 521 phytate content. 633 Corn (maize). 70. 166–167 Coprolite. 307. 236 fiber values. 97 Colostomy effects. 504. 513. 507. processing effects on metabolic responses. 627 glycemic/insulinemic response and. 69 origin of fiber hypothesis. 489–490 phytoestrogen effects. 557 Cookies fiber content. 490–491 gastrointestinal effects. 364 studies indicating no effect of dietary fiber. 326. May 6. 466 Coronary heart disease (CHD) dietary fiber and risk and mortality. 519. 298. 339 Uppsala method applications. 447 carbohydrate hydrolysis. 661. 542 South African patterns. 492–493 mechanisms. 467–472 naturally high-fiber diets and.2387_Index_fm Page 690 Sunday. 215 fecal/luminal pH and. 633. 78–80. 384 . 477 dietary fiber and risk. 423 phytic acid and. 504. 424–426 short-chain fatty acids and. 39 Colorimetry. 334 phytate content. 431–433 Colonic microflora. 330. 424. 670 glycemic index values. 506–509. 3 Coriander. 330. 618. 517–520 phytate effects on bioavailability. 676 Cornmeal. 675 Copper. 338 mineral bioavailability studies. 467–472 South African risk. 310 Corn flakes. 514–516 in vitro binding studies. 213. 595 fecal bulk and composition effects. 93. 489. 312 South African morbidity patterns and. 384 phenolic acid residues. 317 fiber research issues. 650. 485 body weight effects. 475 short-chain fatty acids and. 489–490 insulinemic effects. 511. 318 fecal output and. 156 short-chain fatty acids and. 512. 676 protein digestibility and. 140 protein utilization effects. 593 controversial issues. 532–535. 499. 81. 423 whole grain and fiber effects. purification effects. 666–667 defined. 369–371 dietary fiber and risk. 386–390 Diacylglycerol. 472–474 genetic and autoimmune factors. 664 phytate content. 660 candy and sweets. 629 Critical fecal weight (CFW). Non-insulin-dependent diabetes mellitus British national flour and. 654–655. 267 Denmark. 522 Dates. 259–260. 56 Dietary carbohydrates. 339–341. 619–625. resistant starch. 370. 161. 676 Cutin. ready to eat. 481. 618 cereals. 332–333. 370. See Analytical methods Dietary fiber. 28 common food values. Cancer. 27–31 Dietary fiber. 331. 670 fruit. 306 Diabetes. 3–6. 33 laxation. 363. 323–329 recommended diets. purity of sources. 627 breads. Coronary heart disease. 521. 19. 38–39 food preferences. 370 high-fiber. 40 solubility. 666–667. 664. 101 7-α-Dehydroxylase. 391 risk factors. 55. 63–65 Dextrin. 185. 375 lipid levels and. definitions and terminology. 187 Dietary fiber. 555 Cyclamate. 676 Cranberry. 41 microstructure. 335 mineral bioavailability and. high-fiber diets and. 20–21. 346. 489. 364 Detergent analysis. 253–256 Crohn's disease. cooked. 67–69 Dietary fiber. high-fiber diets. 345. 615–642. 661. 34–36 small molecule and ion adsorption. 373–391 epidemiological studies. 627–628. 40 nutrient absorption. 67. 639–640 cereals. 374–375. high-starch treatment. 5 pioneering literature. 465 South African patterns. 573 Dental caries. 664 fiber supplements. 626. 629 Decalin. 462 Crude fiber. 650–651. Diabetes. 375 long-term safety. 670 Crude protein. 650–658 Cucumber. 330. 345. 15–21. 4–5 Dietary fiber. 342. 40. short-chain fatty acids and. 490 Dialysis. 52. 370 glycemic index values and. values for selected foods. physical chemistry of. 656. 100 Coumestans. 5–6 Dietary fiber. 554. 583 glossary. selected foods. See Antioxidants. 299. 69. plant tissue composition. 670 . 28. 487–481 Diabetes. 435–437 Cross-sectional study design. Glycemic response Dietary fiber. 488 insoluble fiber treatment effects. 375 natural (unsupplemented) high-fiber diets. 657–658 flours. See also specific foods brans. 346. 374. 650–651. transit time and dietary fiber correlations. 554–555. See also Fecal bulk and composition fermentation. See also Waterholding capacity Dietary fiber. methods of analysis. historical contexts. 342.2387_Index_fm Page 691 Sunday. Cardiovascular diseases. 332. 660. See also Viscosity water-holding capacity. classification of. 38 fecal output and. 617–618. Blood pressure. 651–654. 491 Couscous. 65 Dehydroferulic acid. 260 bile acid adsorption. 9–10. 370 IDDM and NIDDM subgroups. 391 fiber-supplemented diets. associated food components. Fecal bulk and composition. 650–651. 380–385 high-carbohydrate. May 6. 375 mechanism of effect. 69. 33–41. 10 total dietary fiber comparison. 667. 36–38 satiety. 626. 35 viscosity. 629–631. 11–12 Dietary fiber. 183–184 meetings and congresses. 65. possible protective effects. 87–88. 2001 8:07 PM INDEX 691 Coumarate. 259–260 Dietary fiber. 365 whole grain and fiber effects. 453 p-Coumaric acid. See also Insulin-dependent diabetes mellitus. 445 D Dairy products glycemic index values. 627 Crackers dietary fiber values. 35–36. 332–333. 571. 369–370 development of dietary fiber hypothesis. See also Breakfast cereals crackers. 597–603. 88–89 HPLC. 70 methods and measurement principles. 3RD EDITION grains. 556–557 US general population. 78–80. 627–629 juices. 233–236 diverticular disease and. 370 historical background. 92 Dodecasodium phytate. 637–638. 561. 240–241 fruit and vegetable effects. 655–657. 670 pasta. 461–462 Dietary fiber supplements. 92 Diversion colitis. 577 "magic bullet" vs. 81 international collaborative trials. 628 snacks and chips. 567–579 public health patterns. 277 lipid absorption and. 617. 661 Southgate method data. 237 mixed-fiber source effects. 660–662 spices. 553–563 Africa. 558–561. 431–433 DMSO. 657–658 Dietary recall. 226–227 oat and corn effects. 81 GLC. 568–570 European patterns. specific supplements dry matter content. 636–637 nuts and seeds. 216–219 purified pectin studies. 80–81 reagents. 555 Middle East. 562 specific US cohorts. 554 patterns to 1992. 556 Endive. 578 obesity and. 260. 633 3. 90. 633 Elderberries. 70. 660. 278–280 in vivo measurements. 676 Dry matter content. 76. 120–121 Doughnuts. 632–636. 262. 79 NSP isolation and hydrolysis. 88. 576–577. 558 reasons for failure to meet recommended intakes. 213–215 particle size effects. 78 Enteral vitamin synthesis. 555–556 whole grain foods. 80. 230–232 foods and mixed diets effects. 2001 8:07 PM 692 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. and miscellaneous. 576. 651 rice. 264. 605–611. 571 institutions and restaurants. 76. 553 household surveys. 266 Enzymatic gravimetric methods. 263. 584. 661. 220–225 gum and mucilage effects. 567 certificated reference materials. 578 long-term trends. 587 direct and indirect evaluation methods. 638–639. defined. 87 . 633–634 Endometrial cancer. 72–73 Uppsala method for total dietary fiber and.5-Dimethylphenol. 631 meat substitutes. 90. 209–212. 640–642 vegetables and legumes. 277–283 cell wall characteristics and. 556 Latin America. 628. 597–598 elderly. 74–76. condiments. 186. 626–627 soup. 277. 88–89 uronic acid determination. 630 Elderly dietary fiber intake. 583–590 children and adolescents. "whole food" approaches. 557–558. 70 colorimetry. 667. 280–281 in vitro measurements. See also specific countries fat and protein effects. 178–179 Enterobacteria. 73–74 quality control. 574. 556 estimates of nonstarch polysaccharide consumption. 481 as marker. 184. 70. 570–574. 557 organizations and their recommendations. 230. 670 Dietary fiber complex.2387_Index_fm Page 692 Sunday. 597 Diferulates. 363–365. 561. 542–543 Eggplant. 77–78. 571 individual variation. 280 Digitonin. 666–667. 561. 238–239 legume effects. 609 Energy excretion cellulose effects. 127 Dill. 657. 650–658 E Edible fiber. 193–201 Englyst procedure. 70. 202–205 wheat fiber studies. 228–229 purified fiber effects. See also Metamucil. 10 EDTA. 441–442. May 6. 97 Dimethylsulfoxide (DMSO). 454 Digestive enzymes. 438 Diverticular disease. defined. 593–595 Australia and New Zealand. 568 guidelines for whole-population nutritional changes. 574–576 Asia. 72 sample preparation. 666. 70–81. 88. 51–60. 264. 70. 70 NSP calculation. 561–562. 578–579 North America. 9 Dietary fiber intake. 336–337 food consumption data. 206–208 cereal product effects. 266 Enterococci. 576. See Phytoestrogens Estrone. 559. 220–225 gum and mucilage effects. 100–101. 216–219 purified pectin studies. 492 Estrogens. 336–337 Fat absorption. 238–239 legume effects. phytate content. 202–205 wheat fiber studies. 629 phytate content. 228–229 purified fiber effects. 45–46 Extraction methods phytate. 122 saponins. 296. 90 Fecal bulk and composition. 52 protein hydrolysis.2387_Index_fm Page 693 Sunday. 206–208 cereal product effects. 637 . 446 Fibra Kneipp. May 6. 51–52 interlaboratory studies. 569. 676 short-chain fatty acids and. 17. 676 Finland. 403. 462–465 Estradiol. 226–227 oat and corn products. 183–241 correlations of transit time. 302–305 Fecal short-chain fatty acids. 634 Fenugreek. short-chain fatty acids and. 637. 230–232 foods and mixed diets effects. 37. 237 mixed-fiber source effects. 253–256 dietary fiber mechanism of action. 264 European Economic Community (EEC) Dietary Survey. 391 Fat removal detergent systems and. 209–212. 216–219 purified pectin. 67 Farina. 144–151 Fecal pH. 186–187 wheat fiber. 501 particle size and porosity effects. dietary fiber and chronic diseases. 206–208 cereals. 318 Fennel. and dietary fiber. 258. See also Lipid-modifying effects Fat excretion cellulose effects. 38–39. 187–192 fiber-containing food. 55 sample preparation. 185–186 particle size. 228–229 purified forms of fiber. 453 Flax seed. 100 carcinogenesis and. 374. 453–454. 57–60 insoluble and soluble fiber. 185 historical background. 485 Fig fiber content. 209–212. 41. 186 fiber physicochemical properties and. 54–55 Epidemiologic studies. See Transit time Fecal microflora. 402 Fecapentaene. 573 Fish. 186. 491–492. 228–229 percent moisture. critical fecal weight. 39 products. 489 mineral bioavailability and. 233–236 explanation of table terms. 457–458 Ferulic acid. effects on metabolic responses to foods. 55 recovery of soluble dietary fiber constituents. 195. 240–241 gums. fiber lipid-modifying effects and. 64 Eubacterium. 491 Fescue. 571. 202–205 report selection. 240–241 fruit and vegetable effects. 238–239 legumes. 568. See Short-chain fatty acids Fecal sterol losses. 64 Uppsala protocol. 189 studies. 56 protein and starch solubilization agents. 486. 262. 237 mixed-fiber sources. 134 Ferulate. 306 Fibrinolytic activity. and other polysaccharides. 54 starch gelatinization and hydrolysis. 127 F FAO/WHO recommendations. 308 Filberts. 190 transit time. 233–236 diverticular disease and. 213–215 particle size effects. 461–477 methodology. 220–225 fiber type and effectiveness. fiber and mineral bioavailability and. 383 Fermentation. 51. 521 Flavonoids. antitoxic effects. 606 Exoskeleton. 213–215 particle size. 114–116. 2001 8:07 PM INDEX 693 approved AOAC protocol. 492 2-Ethoxy ethanol. 193–201 total energy excretion. See Bacteria Fecal nitrogen excretion. 191 foods and mixed diets. See also Short-chain fatty acids protein digestibility and. 676 Fat. 52–53 insoluble fiber. mucilages. 226–227 oat and corn effects. 185–186 cellulose. 402. 183–184 microflora and. 197. 198 Fat intake recommendations. 196. 330. 93. indigestible components. 595 fiber values. 310 Glucose oxidase. 331–332 phytic acid and. 64. 481 Food processing. 482 Germany. saponin analysis. 191. 56 Theander and Åman method. 492 high-carbohydrate. methods of analysis. 601. 667. 220–225. 336–337. 326 US consumption patterns. 650. 95–96 Gelation. 64 β-Glucans. 654–655. 55 Glucofructans. 88–89 Uppsala procedure. 28 Glucosamine. 472 particle size effects and. phytate content. short-chain fatty acids and. 300. 455–456 Australian consumption patterns. 12. 37–38. 15–17 Foods. 267 GLUT 4 receptors. 259. 35 Uppsala procedure and. 680 starch gelatinization and hydrolysis. 260 Gas chromatography (GC). 542 . 33 Food and Drug Administration (FDA). 401. 151 protein digestibility and. 670 fecal bulk and composition effects. 54 Flummery. 491 Glycan. 311 Fungal walls. 28 Galactooligosaccharides. 660. values for common foods. 594 colon cancer risk and. 240–241 fecal output and. 259 short-chain fatty acids and. 307. 335 heart disease risk and. 598. 634 Glucanases. 492 Germ. 129. dietary fiber-associated components. 345. 309 solubility. 261. See also specific fruits antioxidants. See also Antioxidants Fructans. 296. 634 fructooligosaccharides. 560 Ginger. 334–335. 90 β-Glucuronidase. 130 Gas-liquid chromatography (GLC) Englyst procedure. 487. See Dietary fiber. 584 Chinese consumption patterns. 94 short-chain fatty acids and. 666–667. 305 Galactose. 3RD EDITION Flours dietary fiber values. 557 Fucans. 329–332. 488 cancer risk and. 28 Glucomannans. 488 Fructose. Glycemic response. 602 glycemic index values. allowed fiber health claims. 260. 40 Gemfibrozil. 2001 8:07 PM 694 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. See also Carbohydrate metabolism. 28 Glycemic index. 321. 305 fecal microflora and. 262 G Galactans. 166 prebiotics. 333. polysaccharide. dietary fiber values. 629–631. 409. 184.2387_Index_fm Page 694 Sunday. 177 Food additives. 403 Genistein. 453. 260 fecal/luminal pH and. FDA. 261. 330. 670 French consumption patterns. 105 Garlic Chinese consumption patterns. 339 Folic acid. 597–603 Free radical scavengers. 69. 300. 80. 340 Fruit cocktail. 321 short-chain fatty acids and. 512 nitrogen excretion and balance effects. 70. 347 NIDDM risk and. 321–347 fiber viscosity effects. 674–680 Food service outlets. 469 mineral bioavailability studies. 556 France. 310 Fructosamine. 45 Glucose glycemic index values. 265 mineral absorption and. 87. 143 refining effects on glycemic/insulinemic responses. See Analytical methods Foods. 19–21 Foods. Insulin response dietary fiber effects. 87 Gas chromatography-mass spectroscopy (GC-MS). 23–25. 384 Glucoronoxylans. 16. 340 Uppsala method and. 94 Glucoamylase. See Processing effects Foods. 666–667. 559. 346 long-term effects of low-GI diets. 664. 330. 93. 185 fiber content by Southgate method. 74–76. 488 Glutathione peroxidase. 556. 457. 670 phytate content. 28. specific foods Foods. 661 fiber values. 627–628. 629 Fruits. 45 Fusobacteria. 296. 483 Fructooligosaccharides. 28 Galactomannans. May 6. 380. 661. 475 crude fiber values. 105 Glucose metabolism. high-fiber diets and. 24. 483 short-chain fatty acids and. 630. 666 glycemic index values. 345. 261 Glycuronans. 654. 264 fermentation. 277. 374. 324 fecal bulk and. Diabetes. May 6. See Peas Grits. Insulin response cooking/processing effects. 544 Heme iron. 661 Gum acacia. Inflammatory bowel disease Health Professionals Follow-up Study. 137–138 protein utilization effects. 374. 29 Gum arabic. 391 fiber-supplemented diets. 240 fiber values. 38–385 lipid-modifying effects. Coronary heart disease. of dietary fiber or naturally highfiber diets. Fecal bulk and composition. 278. 625 phytate content. 346 high-fiber diets and. 666 glycemic index values. 323–329 fat and protein effects. 373–391. 380–385 fiber-supplemented meals. See Cardiovascular diseases. 339 diabetes treatment. 209–212. 177 food additives. 191. high-fiber diets and. 637. 370. 36–38 H Hairs. 38. 191. 24. Cancer. colon cancer risk and. 339–341 Glycemic response. 264 glucose and insulin responses and. 424 Hazelnut. 405. 310 Gum karaya antitoxic effects. 472 Heart disease. 260 digestive enzymes and. 24 viscosity effects and nutrient bioavailability. 412 high-carbohydrate. 407. See Antioxidants. See Glycemic index grinding effects. 38 fecal composition studies. 661. high-fiber diets and epidemiological studies. 386–390 particle size effects. See also specific types digestibility by microflora. 148 protein digestibility effects. 280. 483 nitrogen excretion and balance effects. 326. 327. 488 whole grain effects. 307 Gum tragacanth. 29. 627 Guar gum. 409. 2001 8:07 PM INDEX utility of. 29 Gooseberries. 374. 375 mechanism of effect. 487–488 anticarcinogenic effects. 238. 17 Haricot bean. 279 short-chain fatty acids and. 332 digestive enzymes and. 676 Health benefits. 281. 335 Grapes fecal composition effects. 70. 375 long-term safety. 681–682 Green peas. Diabetes. 326 lipid-modifying effects. 191 fecal composition studies. See specific grains Gram-positive cocci. 554. 333. 308. 626 Grapefruit fiber values. 376–379 IDDM and NIDDM subgroups. 335 refining and glucose and insulin responses. 36–37 vitamin A absorption effects. 184 food additives. 212 protein digestibility and. 630. 447 diabetes treatment effects. 374–375. 167 . 24 695 glycemic/insulinemic response and. 212 fecal microflora studies. 263. 310 viscosity and nutrient absorption effects. 469–470. 280 fecal composition studies. 254. 29 antitoxic effects. 39 folic acid and. 336–337 glycemic index of foods. 238–239 fecal output and. 225 Hashimoto's thyroiditis. dietary fiber values. 492–493 Glycosidase. 161. 375 lipid levels and. 282 fecal composition studies. Cardiovascular diseases. 332. 175 Guava. 278. 338 guar additives and. Glycemic response. 308. 239 fecal microflora studies. 331–332 short-chain fatty acids and. 210. 69. 375 natural (unsupplemented) high-fiber diets. 239 Gums. 29 blood glucose and insulin responses and. 321–323. 28. 630 Grains. 144. 618. See also Diabetes. 676 Granola bars. 342. 258 Granola dietary fiber values. 326 tartaric acid content. 446 fecal composition effects. 134.2387_Index_fm Page 695 Sunday. 155 short-chain fatty acids and. See also Carbohydrate metabolism. Coronary heart disease HEL-30 cells. 370 Hawaiian Japanese. 174 vitamin D absorption effects. 29. 51–53 glucose and insulin responses and. 333. 136 protein utilization effects. See also Phytate anticarcinogenic activity. 345. 81 phytate analysis method. 507 nitrogen excretion and balance effects. 465–467 cooking/processing effects. high-fiber diets and. 370. 96 Hippocrates. 332 high-carbohydrate. 15–17 Inflammatory bowel disease. high-fiber diets. 113. 346. 322 crude fiber composition. 335 Horseradish. 370 glycemic index values and. 279 Hominy. Diabetes. 220–225 high-carbohydrate. 374. 532. 560 genetic and autoimmune factors. 533 7-α-Hydroxylase. 374 mineral bioavailability studies. 492–493 Intestinal microflora. high-starch diets. See Cholesterolreducing effects Hypolipidemic activity. 485 apoptosis and. 554 antitoxic effects. 63 digestibility by microflora. 637. 424. fiber in treatment of. See also Diabetes. 386–390 high-fiber. 321–347. 161. 569 values for common foods.2387_Index_fm Page 696 Sunday. 533. 391. 342 high-fiber diets and. 161–162. 435–437 short-chain fatty acids and. 615–642 Institutional diets. See also Cholesterol-modifying effects heart disease risk and. 321–347 whole grains and. 260 glycemic/insulinemic response and. 146 plant tissue composition. 29 Hexoses. 178 HepG2 cells. 676 High-carbohydrate. 436 diabetes treatment. 486. 339–341 Insulin-like growth factor-I (IGF-I). 77–78. high-fiber diets and fecal composition studies. 28 detergent-based analysis and. 339 fat and protein effects. 2001 8:07 PM 696 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 261 Indigestible food components. 435. 282. 69 transit time and. glycemic index values. 342. 321--347 lipid-lowering effects. 3. 69 HMG-CoA reductase. 437–438 ulcerative colitis. 423 . 70. 538. 409. 493 Insulin-like growth factor-II (IGF-II) receptor. See also under Glycemic response viscous fiber effects. 373–391. 483 Holocellulose. 465. 321–324 fiber hypotensive effects and. See also Diabetes. 261 Hyperlipidemia. 113–125 saponin measurement. 544 Insulin resistance. 404–413. 336–337 fiber and single-test meals. 436 possible protective effects. 543 Insoluble dietary fiber. 338 high-fiber diet and. 129 Uppsala procedure. 487–488. See also Diabetes Austrian youth diets. 370 inflammatory bowel disease and. See pH Inuit. See Lipid-modifying effects I Iceland. 342. See also Carbohydrate metabolism. Lignin analytical methods. 402–413. 532. See Lipid-modifying effects Hypertension. 386–390 High density lipoprotein (HDL) cholesterol chitin and chitosan and. casein digestibility and. 374. diabetes treatment and. 484 fiber lipid-modifying effects and. 484 Hypocholesterolemic activity. 370. 12 protein digestibility and. 401–414. 466 High-fiber diets Bristol diet. 403–405 national consumption patterns. 370. 437 Immunostimulant production. 374. 332–333. 332–333. 331. 493 chronic disease risk factors. 35. high-fiber diets. 9. See specific sugars Hickory nut. 437 Inositol phosphates. high-fiber diets and utility of glycemic index. 29 Honey. 666 HT-29 cells. See also Cellulose. See Bacteria Intestinal pH. 676 Homoglycan. 363. 255–256 High-pressure liquid chromatography (HPLC). 46 fiber effects on blood levels. 94 vitamin C and. 345. 492 Insulin response. 153 selected food values. 29. 370. 650–658 Uppsala procedure and. 435–438 Crohn's disease. 542 Heteroglycans. 3RD EDITION Hemicellulose. Glycemic response antithrombotic effects and. 447 carbohydrate metabolism and. 403 grain grinding effects. 575 Ileostomy effluent. 373–391. 403. 556 Insulin-dependent diabetes mellitus (IDDM). 346. May 6. 166 protein digestibility and. 311 Laminarin. 290. 339 . 279 glycemic/insulinemic response and. 476 Ireland. 183–184 Leavening. 474. 40 factors affecting absorption. 441–442. 262. See also Lignin Kohlrabi. 656. 684 Lemon curd. 385 lipid-modifying effects. 259. 541. 191 lipid-modifying effects. 28 Iron. 631 glycemic index values. 339 saponin content. 330. 87. 634 carbohydrate hydrolysis. 99–101. 279 dry matter content. 237 fiber values. phytate content. 24 lipid-modifying effects. 443 Jerusalem artichokes. 432 fecal composition studies. 260 Legumes. 492 Ispaghula diverticular development and. 178 vitamin E bioavailability effects. 655–657. 184 glycemic index values. 517–519 phytate and. 330. 335 mineral bioavailability and. 410–413 processing effects. 303 food additives. 470. 134. 266 Laminaria digitata. 674 processing effects on glycemic/insulinemic response. 666 fecal composition studies. 553 Ketchup. 374 laxative effectiveness. 211. 374. 143 vitamin B12 and. See also Fecal bulk and composition fiber mechanism of action. 409 protein digestibility and. 167. 260. 296. 239 protein digestibility and. 499. 138 short-chain fatty acids and. 301 Israel. 3–4. 129 Kjeldahl method. 335 phytate content. 264. 300. 438 Lactobacillus. 575. 424 dietary patterns. 561–562. 441–444 mortality rates. 300. 334. 634 Konjac mannan diabetes treatment effects. 3 Laxative effects.2387_Index_fm Page 697 Sunday. 676 Kellogg's. 605–611 J Japan colon cancer risk. 634. 330. See also specific types Chinese consumption patterns. 661 Lentils. 340 glycemic/insulinemic responses fat and protein effects. 656 glycemic index values. 532. 261. 559. 24. 24 Jicama. 310 Iowa Women's Health Study. 676 K562 cells. 573 Irish moss. 263 Lane. 336–337 grinding effects. 576. 296. 267 mineral absorption and. mineral bioavailability and. 503 bioavailability studies. 266. 267 Lactore. 260. 41. 302. 666 dietary fiber values. 578–579 Italy. 191 historical interest in fiber. 330. Albuthnot. 533 Kidney beans fecal composition studies. 175 L Lactic acid. 511–516 dietary fiber and absorption. 334. 138 short-chain fatty acid excretion and. 594–595 crude fiber values. 328 digestive enzymes and. 483 microflora and. 483. 472. 655 glycemic index values. 296. 634 Juices fiber content. 339. 501 in vitro binding studies. 324. May 6. 501 Leek. 632–636. 577 fecal microflora. 504–507. 266. 186 fiber type and effectiveness. 281. 568. 265 morbidity patterns. 334 glycemic/insulinemic response and. 338 processing effects. 16. 339 saponin content. 542 Isoflavones. 20. 237 fecal output and. 570. 260–261. 161. 554 fecal/luminal pH and. 298. 491. 2001 8:07 PM INDEX 697 Inulin. 660. 238. 522 K Kale. 329. 55 Klason lignin. 258. 129. 260. 651. 482 dietary fiber effects. 504. 575 . 103. 407. 664 glycemic index values. refined grains. 517 phytate effects on bioavailability. 335 Mammary cancer. 448 Manganese. 218 lipid-lowering effects. 661 Mastication. 277. 21. 409 insoluble fibers. 512 in vitro binding studies. 305 Low density lipoprotein (LDL) cholesterol antioxidants and. 403–405 nitrogen excretion and balance effects. 29. 401–414 bile acid binding. 412 heart disease and. 403–412. See also Cholesterol-reducing effects high-carbohydrate. 161. glycemic index values. 176 selected food values. 153 pyridoxine bioavailability and. 409 nitrogen excretion and balance effects. 661 Mannans. 405. 628. 410–413 locust bean gum. 486 Lipid-modifying effects. 486. 487. 676 saponin content. 532. 403–405 Konjac mannan. 511–516 diabetes risk and. 150 Loperamide.2387_Index_fm Page 698 Sunday. 406 wheat bran. 104 Lignans. 554. See also specific fiber components guar. 136. 637. 173 whole vs. 19. 456. 402 particle size and. 507. 539 Mammary studies. 33 Matairesinol. 650–658. 301. 660–662 sulfite and. See Corn Malondialdehyde. high-fiber diets and. 517–519 phytate effects on bioavailability. 503 bioavailability studies. 282–283 possible mechanisms. 46 digestive enzymes and. 65 Uppsala procedure. 129 Lettuce crude fiber/dietary fiber comparison. 448 crude fiber composition. 164–166 Magnesium oxide. 24. 677 Magnesium. 167–168 Mango. 406 psyllium. 532. 278. 453. 20. 445. phytic acid and. 279 protein utilization effects. 146 protein digestibility and. 28 dietary fiber definitions and. 385 protein digestibility and. 409 oat bran. 2001 8:07 PM 698 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 103 vitamin A absorption effects. 402 specific fiber types. 175 Lipases. 410 pectin. 334 phytate content. 482–483 Lupin digestive enzymes and. 486 Maltose. 403 insulin effects. 492 Lignin. 105 Marmalade. 333 vitamin E and. 503 bioavailability studies. 411 vitamin metabolism and. 280–281 effective dosage and formulation. 538. 20 antioxidant activity. 660–662 Southgate method. 408. 402 fiber effects on blood levels. 403 lipid absorption alterations. 37 antitoxic effects and. 9 digestibility by microflora. 174 vitamin D absorption effects. May 6. 402. 99–101. 666 Macaroni fiber values. 535. 280. 402 genetic factors. 458 antitoxic effects. 280 glycemic/insulinemic responses and. 409–410. 278 glycemic response and. phytic acid and. 405. 85. 533. 446 digestive enzymes and. 142 saponin content. 129 M Macadamia. 634. 405. 533. 486 whole grain and fiber effects. 483 chitin and chitosan. 483 Liver cancer. 93. 409 legumes and legume proteins. 260 fecal composition studies. 312 Uppsala method applications. 29 Mannose. 660 short-chain fatty acids and. 324 lipid-lowering effects. 280 Lipid hydroperoxides. 173. 226 Maize. 474 in vitro binding studies. 538. 482–483 meta-analysis. 402–413. 670 fiber values. 408. 539 Locust bean gum. 509. 505. 486 Mauritius. 3RD EDITION phytate content. 555 analytical procedures. 535. 403 short-chain fatty acid generation. 161. 402–403 fecal sterol losses. 571. 547 Nitrates. 226–227 . 543 N National Cancer Institute (NCI). 220–225 gum and mucilage effects. Chronic diseases. 240–241 fruit and vegetable effects. 541. 144–151. 310 N-Methyl piperazine buffer. whole grains and. 573. 629. 17 Mucorales. 504–516 enrichment studies. 501 in vitro binding studies. 29. 533 MDA-MB-231 cells. 96. 555–556. 532. 568 National Health Interview Survey. 576 Niacin. 475–476 Mucilages. 502 other food additions and. 650–658 defined. 501 Mineral(s). 167–168 phosphorus. 499–500 Mineral coprecipitation. 306 Methylcellulose diverticular development and. 161. 237 mixed-fiber source effects. 168–169 zinc. 432 fecal composition studies. 482. 177 Nickel. 338. 406 short-chain fatty acids and. 129 Mushrooms. 191. 184. 177 Nigeria. 500 cereals studies. 55 Modified carbazole method. 161. 666 Mustard greens. Inflammatory bowel disease. 162–164 processing effects. 554. 226. 542–543 anticarcinogenic effects. 161. 206–208 cereal product effects. 164–166 manganese. 97 Modified starch additives. 334 Microflora. 634. 654 Metamucil dry matter content. 63 nitrogen excretion and balance effects. 104. 206. dietary fiber effects. 24 699 Morbidity. 324 short-chain fatty acids and. 562. 578 M'fino. 533 Meat substitutes. 501 references to minerals studied. 45 Muesli. 209–212. 191. Diabetes. 317 Nitrites. 166–167 iron. 309. 656 Mustard seed.2387_Index_fm Page 699 Sunday. 29. 334 Milling effects. 29 Milk. 278 glycemic index values. 40. 555. See Dairy products Millet. 500 unrefined cereal products and. 503 solubility effects. 238–239 legume effects. 499 Nicotinic acid. 10 New Zealand. 500. 542 calcium. 298. 630 Netherlands. 560. 532. 121 Mexico. Coronary heart disease. chronic diseases and Mortality (all causes). 167 magnesium. 40. Cardiovascular diseases. 238–239 Mucopolysaccharides. 636–637 Melon antioxidants. 161. 544 Nectarine. 230–232 foods and mixed diets effects. 119. 557 National Food Consumption Survey (NCFS). 531. 500 human studies/animal studies. 517–520 in vitro/in vivo studies. 317 Nitrogen analysis. 562 Natural killer cells. 2001 8:07 PM INDEX McCance and Widdowson Food Tables. 658 lipid-modifying effects. 501 intrinsic and extrinsic factors. See Bacteria Middle lamella. 161–169. 555 National Health and Examination Study (NHANES II). 445–448 Myoinositol. 521–522 phytate and. 554 fecal composition studies. 499–522 binding behavior. 501 long-term studies needed. 209–212. See Cancer. 317–318. 153 Neutral detergent residues (NDR) common foods. 27 Mutagens. 625 Multiple sclerosis. 595 digestive enzymes and. 190 cellulose effects. 55 Nitrogen excretion and balance. 239 glycemic/insulinemic responses and. 67 MCF-7 cells. 144 Neutral detergent fiber (NDF). 574 Net protein utilization. 334. May 6. 561. 370 Mung bean. 233–236 diverticular disease and. 21 Mineral bioavailability. 186. 146–147 protein utilization effects. Whole-grain foods. 656. 464. 166 copper. 455 fiber values. 630. 561. 482 Chinese consumption patterns. 481. 151 Oat bran antitoxic effects. 369 cancer risk and. 30. 482 antioxidants. 660. 374. 312 straw. 365 Oils. 17 Okra fiber values. 307. 292. 677 Obesity African dietary changes and. 24 Olives. 10 Englyst procedure for determining. 650 mineral bioavailability studies. 260. 304 fiber values. 228–229 purified fiber effects. 631. 627 glycemic/insulinemic responses and. 288. 653 FDA-allowed health claims. 469–470. 101. 85 selected food content by Southgate methods. 453 carbohydrate hydrolysis. 666–667. 213. 487–488 Nonstarch polysaccharides (NSPs). 410 mineral bioavailability studies. 265. 216–219 purified pectin studies. 332–333. 311 Uppsala method applications. 295. 213–215 particle size effects. 661. 677 Omega-3 fatty acids. 637–638. 338 hypotensive effects. 12. 666. 374. 675. high-fiber diets and utility of glycemic index. 489. See also Diabetes dietary fiber and risk. 557 Nutrition Coordinating Center (NCC). 487–488 historical interest. 87 food additives. 370 high-fiber diets and. 278. 67. 615 Nuts and seeds crude fiber values. 660–662 Non-digestible oligosaccharides. 104 Uppsala method collaborative study. 664 Nurses Health Study. 260 Non-insulin-dependent diabetes mellitus (NIDDM). 37 Oatmeal crude fiber values. 235 fecal/luminal pH and. 492 fiber intake and. 295. 472–474 epidemiological studies. 484 lipid-modifying effects. 491 N-Nitroso compounds. high-fiber diets and. 279 dietary fiber values. 88–89. nitrogen excretion and balance effects. 670 dry matter content. 519. 30. 666–667. 297. See also Diabetes. 666 glycemic index values. 289. 488 high-carbohydrate. 661. 214. 677 Oligofructose. 654. See also Englyst procedure marker of a naturally high-fiber diet. 292. 670 fiber values. 332–333. 618. 282 fecal composition studies. 140 short-chain fatty acids and. 70–81. 370.2387_Index_fm Page 700 Sunday. 342. 656 fiber values. 29. 656. 68. 129. 567 defined. 308. dietary fiber table. 553 lipid-lowering effects. 513. 296. 191. 568–570. 370 glycemic index and. 447 digestive enzymes and. 618. 472. 630. 299. 676 O Oat. 330 . 670 fructooligosaccharides. 634. 472. 554 dietary fiber definitions. 69 national consumption patterns. 490 Onion crude fiber/dietary fiber comparison. 657. 583–585 Noodles. 384. 483 Oligosaccharides. 464. 303. 342. 373–391. 391 genetic and autoimmune factors. 184 fecal/luminal pH and. 332. 684 short-chain fatty acids and. 670 phytate content. 514 nitrogen excretion and balance effects. 517 phytate content. 304 grinding effects on glucose and insulin responses. 656. 634. 339–341 whole grain and fiber effects. 193–201 Nitrosamines. 202–205 wheat fiber studies. 656. 401 mineral binding studies. 260 neutral polysaccharides not recovered by Uppsala procedure. 317 Non-cellulosic polysaccharides. 215 fecal output and. May 6. 674. 666 phytate content. 103 viscosity treatment. 288. 520 saponin content. 401 fecal bulk and composition effects. 310. 266. 474. 666. See Pasta Nopal leaves. 3RD EDITION oat and corn effects. 666 dietary fiber values. 557 South African morbidity patterns and. 300. 381 Norway. digestibility by microflora. 2001 8:07 PM 700 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 558. 150 protein digestibility and. 93 Orange fiber values. 308. 51. 265 nitrogen excretion and balance effects. 630 pH. 308. 330. 446. 326 Peach crude fiber/dietary fiber comparison. 657 fiber values. 30 Peppers. 129. 634–635. 97 Phosphates. 380. 677 Particle size fecal composition studies. 677 Peanut dry matter content. 63 Pectin. 278 Pernicious anemia. See also Digestive enzymes. 303 fecal mutagens and. 637. 684 Pear. bioavailability studies. 661. 104 viscosity effects and nutrient bioavailability. 381. 141. 447 cell wall porosity and. 228–229 fecal output and. 176 Parsley. 670. 309 Peas. 173. 670 glycemic index values. 277–278. 173. 292. 374. 93 phytate content. 660. 154–155 pyridoxine bioavailability and. 23 antitoxic effects. 281. 177 glycemic/insulinemic response and. 297. 280. 293. 630. 617. specific enzymes Pancreatin. 660. See also specific compounds phytic acid and. 12. 631 mineral bioavailability and. 409 Pancake. 303 fiber values. 175 Pectinic acid. 511. 405. 12 protein digestibility and. 513–516 in vitro binding studies. 185 fiber effects on glycemic response and. 144. 174 vitamin B12 and. 104 Uppsala method collaborative study. 176 short-chain fatty acids and. 178 vitamin C and. N. 628 fiber values. 656. 136–137. 30. 27. 510. 278. 299. 666. 307. 677 Pancreatic enzyme activity. 324 neutral polysaccharides not recovered by Uppsala procedure. 123–125 Phosphorus. 634 Parsnips. 330 phytate content. 486 Persimmon. 261. 370 Peroxides. 280. 330 glycemic/insulinemic responses and. 294. 664 glycemic index values. 654. 101. 302–305 Phenolic acids or esters. 185. G. 264. 20 fiber values. 287. 334 guar additive and glycemic response. 666. 35 Uppsala method applications. 554. 531 restandardization for phytate analysis. 100–101. 650–651. Spaghetti dietary fiber values. 37 vitamin A and. May 6. 635. 657. See also Uronic acid residues additives. 55. 277. 21 P Painter. 161. 335 short-chain fatty acids and. 52. 670 glycemic index values. 103. 309 Uppsala method applications. 302. 306. 630. 660. 184. 651. 661. 30 Pentoses. 628. 282 fecal bulk and composition effects. 666 Pepsin. 38 folic acid and. See also Macaroni. 2001 8:07 PM INDEX 701 refining effect on glycemic/insulinemic responses. 670 glycemic index values. 202–205. 503. 482. 239 fecal/luminal pH and. 216 fecal/luminal pH and. 191 lipid-modifying effects. 318 fermentation. 142 saponin content. 326. See also Legumes fecal composition studies. 677 Pectic substances. 654. 289. 308–311 solubility. 507. 147 plant tissue composition. 296. 326 fiber content. 297. 670 fiber content. 51. 412 phytate content.. 522 Oxalates. 321–323. 335 phytate content. 518. 260. 168–169 . 129. 684 short-chain fatty acids and. 39 digestive enzymes and. 282–283 Pasta. 677 refining and glycemic/insulinemic responses. 670 glycemic index values.2387_Index_fm Page 701 Sunday. 519 phytate effects. 677 protein digestibility and. 661. 453–458. 677 saponin content. 279 protein utilization and. 296. 52 Pantothenic acid. 33. 21. 491 3-Phenylphenol. 178 vitamin D absorption effects. 9. 431 Palatability. 331–332 lipid hydrolysis and. 30 detergent-based analysis and. 385 laxative effectiveness. 406 microflora and. 103 Pecan. 661. 485. 24 Polyfructans. 541–542 inositol phosphate pool. 113. 374 fat and protein effects. 406 Plantain. 538. 164–166 manganese. See Beans Pistachio. 326 phytate content. 533 liver. 500. 541–544 antioxidative effects. 114 restandardization. 678 short-chain fatty acids and. 541–543 binding studies in vitro. 532. 660. 538. 177 Polymers. 654. 661 Pineapple. 539 soft tissue. 516 in vitro binding studies. 539 mammary. 532–535. 539 human studies. 30 Plasminogen activator inhibitor type 1 (PAI-1). 541 animal studies. 488 Plum antioxidants. 514. 33 Pomegranate. 161. 656. 68 Polysaccharide food additives. 291. 54 Uppsala method applications. semisynthetic food additives.2387_Index_fm Page 702 Sunday. 542 antiproliferative activity. 166–167 iron. 104 Potato chips fiber values. defined. physical chemistry of Phytase. 670 glycemic index values. 122–123 procedure. 167 magnesium. 539. 329 Phytate. 120–121 Phytate. 340 glycemic/insulinemic response and. mineral bioavailability and. See Phytic acid. 677 Plantic complex. 543–544 enzyme inhibition and malabsorption. 330. 531–544 administration mode vs. 11–12 Plantago. 114–116. 166 copper. 161. May 6. 509. 336–337 processing effects. 113–125 detection methods. 626–627. 113–114 quantitative methods. 539–541 adverse effects. 264. 490–492 Phytoestrogens. 2001 8:07 PM 702 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. cancer and antinutrient effects. 656 fecal composition studies. 161–169. 532. 168–169 zinc. 519 Potato dry matter content. 40. 491–492 Pickles. 340 Potex. 666 Pinto beans. 531 anticarcinogenic effects. 23–25. 543 mineral interactions. 10 Plantix. 537. 511. 339 refining effects. 674 Plant tissue composition. 339. 654. 288. 631. 491. 486 whole grain anticarcinogenic effects. 482. 670 glycemic index values. 103 . 169 Phytate (and phytic acid). effectiveness. 162–164 Phytic acid. 666. 3RD EDITION Physical chemistry. 297 starch gelatinization and hydrolysis. 631 Popcorn. 103. 666. 531 antioxidative effects. 161. 486 in whole grain. 666. 512. 330. analytical methods. 218. 121 supplies. 532. 677 Poppy seeds. See Dietary fiber. 222 fecal/luminal pH and. 116–117 extraction. 560 Polydextrose. 539 mode of administration. 122 gradient HPLC method. 121–122 precautions. 536. 533 colon. 486. 661. See also specific substances cardiovascular protective effects. 466. 535. 533. 627 glycemic index values. 518. 637 Pita. 541 immune response effects. 617. 635. 533. 533 skin. 539 Phytochemicals. 10. 537. 538–541 multi-organ and bladder. 576 Polysaccharide classification. 532 in vitro studies. 631. 167–168 phosphorus. 542–543 prostate. 482. 24 Polynesia. cancer and. 535. 101. 116 qualitative methods. 117–125 instrumentation. 541 potential mechanisms. 515 anticarcinogenic effects and. 533. 335 Poland. 117–120 purification. 161. 678 Porosity. 536. 127. 39 Potassium. 674–680 glycemic response and. 339. 554 Polyglutamates. 532. 302 fiber values. 517 calcium. 123–125 standards. 532–541 blood. 542 food content. 21 bioavailability studies. 238. 455 fiber values. 55 intake recommendations. 260. 585–590 defined. 134 fiber-diluted diet. 133–144. 16 mineral bioavailability and. 300. 635. 51. 491 Reactive oxygen species. 310 vitamin A absorption effects. 321. selected food values.2387_Index_fm Page 703 Sunday. 226. 278. 5–6 Purified plant fiber. 138 . 177 Public health programs. 384 lipid-modifying effects. 238. 402. 678 Purification effects. 653. 631 Reactive nitrogen species. 654 fiber values. 288 fiber lipid-modifying effects and. 134 in vitro studies. 55 fecal/luminal pH and. 631 Psyllium antitoxic effects. 277. 661 Pretzels. 30. 340 tartaric acids. glycemic index values. 307. 133–134 fiber-rich sources and. 16 protein digestibility and. 374. 298. 34. 533 Proteases. 328 digestive enzymes and. 338–339 heat-produced artifacts. See also Short-chain fatty acids cancer risk and. 19. 289. crude. 239 fecal/luminal pH and. 259 Preserves. 259 Processing effects. May 6. 174 Pteroylmonoglutamates. 623 Raisins. 254–255. 136 Raisin bran cereals. 240 juice. 383. 55 Resistant starch. 144 hydrolysis. antioxidants and. 406 riboflavin absorption and. 492. 482 antioxidants and. 401. 279 animal studies. 462 Prostaglandins. 391 metabolic responses to foods and. 52. 135–139 substitution method. 176 satiating effects. 501 postprandial glycemic responses and. 129 R Radicchio. 296. 405. 30 Prune. 152–155 Protopectin. 282 diverticular development and. 144–151 general fiber influences. 457 glycemic/insulinemic responses and. 40. 628 saponin content. 678 Raffinose. 52 Protein. 335. 302 protein digestibility and. 293. 631. 336–337 solubilization agents. 192. 446 diabetes treatment effects. 381. 16. 140–143 fiber effects on nitrogen excretion and balance. 666 Pumpkin seeds. 678 Probiotics. 309. 651. 20 efficiency ratio. 156–157 fiber effects on nitrogen excretion and balance. 656 fiber content. 281. 133–134 Protein utilization effects of fiber-rich sources. 176 Q Quinoa fiber values. 631 fecal composition effects. 134. 485 short-chain fatty acids and. 144–151 purified dietary fiber effects. 322–323 Propionate. 490 Prostate cancer. 583 Australian consumption patterns. 635 Radish dry matter content. 2001 8:07 PM INDEX Prebiotics. dietary fiber values. 303 fecal microflora study. 558 Puffed grain cereals. 681–682 Raspberries. 10 enzymatic gravimetry. 39 dietary fiber feeding duration and. See Antioxidants Refined flour. 650–658 Protein digestibility. 263 703 glycemic/insulinemic responses and. 287–313. defined. 635 phytate content. 432 FDA-allowed health claims. 532. 660 Puffed rice. 296. 209–211. 10 Pyridoxine. 482 Residual starch. 401 fecal composition studies. 654 glycemic index values. 334. 278 purified dietary fibers and. 64 Protein. 324. 87. 623. 334 Pumpkin. dietary fiber values. 483 Prospective study design. 261. 334 lignans. 45 Short-chain fatty acids (SCFAs). 678 protein digestibility and. 423–424 heart disease risk. 517. 278 fiber values. 542 Rhamnogalacturonan. 486. 3RD EDITION short-chain fatty acids and. 338 processing effects. 98 Rhamnose. 289 fiber feeding duration and. 487 lipid-modifying effects. 538. 290 fiber effects and intestinal segments. 515 Semisynthetic polymers. 517 Uppsala method applications. 289. 306–313 glycemic and insulinemic effects. 297. 483 mineral bioavailability studies. 678 Uppsala method applications. 683–684 analytical methods. 304 short-chain fatty acid excretion and. 127. 506. 143. 533. 628. 128–130 Satiety. 402. 627 short-chain fatty acids and. 469 Secoisolariciresinol. Propionate animal models and. 664. Butyrate. 129. 309 South African morbidity patterns and. 447 digestive enzymes and. 446 carbohydrate hydrolysis. 235–236 fiber values. 470 Shear thinning. 513. 365 colon cancer risk. 499. May 6. 127–130 extraction. 678. 521 national consumption patterns. 34–36. 631. 296. 519. 2001 8:07 PM 704 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 93. 503. 515. 93 phytate content. 278 Sample preparation. 51 Rhubabrb. 279 Chinese consumption patterns. 279 crude fiber values. 301 Sesame seed. 287 cancer and. 330. 16. 509. 513. 30 Scotland. 513. 508. 340 glycemic/insulinemic responses fat and protein effects. 455–456 antitoxic effects.2387_Index_fm Page 704 Sunday. 667 fecal output and. 312 Rice bran antitoxic effects. 301. 290. 287–288 fiber lipid-modifying effects and. 363–365 Rutabaga. 317. 297–299 fecal excretion. 33 glycemic index values. 101. 54. 176 Rice. 482 antioxidants. 593–594 crude fiber values. 514. 667 food preferences. 39. 520 neutral polysaccharides not recovered by Uppsala procedure. 571. 664 Senna fecal/luminal pH and. 507. 36 Shellfish. 455 antitoxic effects. See also Acetate. 339 mineral bioavailability studies. 556 Retrograde amylose. 637. 103. 104 S Safflower. 635 Sclerenchyma. 542 fiber concentration effects. 151 phytate content. 291–295 in vitro production. 485 Satiety index (SI). 488 . 572 Scottish Heart Health Study. 365 Restaurant foods. 491. 446 carbohydrate hydrolysis. 482. 339. 532. 291–313 cecal concentrations. 300–301 fecal/luminal pH. 298 Rolls. 334. 24 Semolina. 103 Rye bran glycemic response and. 577 nitrogen excretion and balance effects. 104 Uppsala method collaborative study. 296. 302–305 intestinal segment concentrations. 504. White rice antioxidants. 39. 34 Sauerkraut. 370 Rhozyme S. 105 Rheumatoid arthritis. 575. 486 Secondary cell wall. 412. 262. 464–465 Sapodilla. 279 short-chain fatty acids and. 288. 631 Saponins. 519. 516. 289. 185. 489 glycemic index values. food content. 287–313. 678 Rural African morbidity patterns. 617–618. 72–73 Sample size issues. 661 Riboflavin. 667 dry matter content. 656. 330. 635. 382 mineral binding studies in vitro. 129 methods of analysis. 336–337 grinding effects. See also Brown rice. 489. 30 Selenium. 657 Rye. 684 Seventh Day Adventists. 656 fecal composition studies. See also Resistant starch Rhabdomyosarcoma. 40. 483 fiber source and. 264 Study design. 569 values for common foods. 279. 27. See also Alginates antitoxic effects. 52. 657 fiber values. 595 crude fiber values. 447 carbohydrate hydrolysis. See also Hemicellulose. 482 Chinese consumption patterns. 278 fecal composition studies. 336–337 guar additive and. 150 protein digestibility and. 279 certificated reference materials for Englyst procedures. 631. 654–655. 335 705 glycemic/insulinemic responses and. phytic acid and. 640–642 Spinach crude fiber values. 410. copper bioavailability and. 667. 638–639. 536. 635 saponin content. 167 protein digestibility and. 514. 413 protein. 141–142 saponins. 539 Solka Floc. 90–92 Strawberries fiber values. 83–85. 308. 539 Slipper elm. May 6. 339 refining effects. Pectin cholesterol-lowering effects. 334 Silica. 518 Sodium alginate. 667 glycemic index values. 684 short-chain fatty acids and. 278. 280 protein digestibility and. 363–365. 233 fiber values. 334.2387_Index_fm Page 705 Sunday. 635–636. bioavailability studies. 661. 309. 287 Shredded wheat cereals dietary fiber values. 514. 533. 83–84 starch hydrolysis. 623. 569. 462–464 Suberin. 653 glycemic index values. 88 solubilization agents. 30. 513. 93 phytate content. 382 neutral polysaccharides not recovered by Uppsala procedure. 401–414 FDA-allowed health claims. 24 protective effects. 661 South Africa. fiber values. 54–55 hydrolysis. 670 phytate content. 330. 554. 651 glycemic index values. 326 Spain. 40 fecal bulking and. 70 Chinese consumption patterns. 667 Stachyose. 340 glucose/insulin responses fat and protein effects. 570. 597 acid hydrolysis for non-cellulosic polysaccharides. 650–658 Sorghum. 88. 51. 628. 129. 584–585. 628 mineral bioavailability studies. 667 dry matter content. 85 sample extraction. 423 fiber values. 19. 30 Skin cancer. 412 processing effects. 635. 667 diabetes treatment effects. 615–642. 260. 374. 21. 624. 84–85 Soybeans (and soya products) antitoxic effects. 216 . 127. 632. 657. 437–438 in vitro systems. 657. 401 national consumption patterns. cholesterol-reducing effects. 365 residual. dietary and morbidity patterns. 678–679 protein. 63 Soft tissue cancer. 192 gelatinization. 262. 328 fecal composition studies. 509. 567. 667 saponin content. 326. 129. 54–55. 561. 35 Soluble fiber. 219. 324. 21. 88 Uppsala protocol for total dietary fiber. 141 Soup. 138 Sodium lauryl sulfate. 40 fecal composition studies. 679 Streptococci. 27. 660–662 interlaboratory studies. 55 resistant. 88 residual lignin measurement. 68 Sugar beet. 85 dietary fiber content of selected foods. 446. 628. 5560 Spices and condiments. 536. 595 digestive enzymes and. 129. phytic acid and. 104 Spaghetti fiber values. 309 Solubility. 35. 555 Sugar alcohols. 310 Uppsala method applications. 296 luminal/fecal pH and. 423. 151 Sodium. 621. 516. 657. 519 nitrogen excretion and balance effects. 570 Southgate method. 237 fiber values. 684 Sprouts colon cancer risk and. 412. 16 Starch. 2001 8:07 PM INDEX inflammatory bowel disease and. 446 digestive enzymes and. See Resistant starch Starch removal detergent systems and. 84–85 modified additives. 684 Squash. 63–64 Englyst and Uppsala methods. 93 nitrogen excretion and balance effects. 254 Tangerine. 51. 661 Sunflower seed. 482 mineral bioavailability studies.2387_Index_fm Page 706 Sunday. 238–239 legume effects. 411. 657. 482. 142 protein utilization effects. May 6. 157 short-chain fatty acids and. 667. 667. 254–255 wheat fiber studies. 660. 309. See Non-insulin-dependent diabetes mellitus U Uganda. 413. See also Lipid-modifying effects Triticale. 629 Tartaric acid. 55. 571. 657. 281 Thyroid cancer. 486 Tokelau. dietary fiber modification effects. 667. 660 Sweet potato fiber values. 445–448 Toxic reagents. 681–682 Tea iron absorption and. 618. 280 Turnip. 492 Theander and Åman method. 679 Teff. 679 Tween. 340. fiber effects on blood levels. 101–105 collaborative study. 90 Testosterone. 679 Sweden. 515 phytate content. 87–105. 254–255. See Insulin-dependent diabetes mellitus Type 2 diabetes. 129 Thymidine kinase. See specific sugars Sulfite. 3RD EDITION glycemic/insulinemic responses and. 101 GLC. 334 phytate content. 456. 458 Tapioca. 559–560 Swedes. 483. 317 Tocotrienols. 278. 501 mineral bioavailability and. 567–570. 51 Tamarinds. 638. 521 phytate content. 489 cellulose effects. 95 . H. 230–232 foods and mixed diets effects. 576 Tortilla. 307. 175. 2001 8:07 PM 706 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 255–256 particle size effects. 679 Uppsala method applications. 226–227 oat and corn effects. 667 glycemic index. 679 T Taco shells. 52. 216–219 purified pectin studies. 193–201 Triethylene glycose. 679 Takadiastase. 435. 240–241 fruit and vegetable effects.. 629 Termamyl. 661. 65 Sultana. 446 Type 1 diabetes. 311 Sugars. 670 dry matter content. 655 Tannins. 83–85. 410. 324 neutral polysaccharides not recovered by Uppsala procedure. 651. 453. 282–283. 636. 679 Tonga. 202–205 tartaric acid and. 618 Total dietary fiber. 150 protein digestibility and. See also Vitamin E fecal mutagens and. 572. 233–236 diverticular disease and. 189 carcinogenesis and. 474 Tocopherols. 64 Transit time. 636. 237 mixed-fiber source effects. 369–370 Uppsala procedure. 574 flour changes and morbidity patterns. 4 Trypsin. saponin measurement. 209–212. 55. 437 Ultrafiltration. 54. 558–559. 576 Tomato crude fiber values. See Uppsala procedure Thiamin. 95–96 hemicellulose fraction. 311 United Kingdom fiber consumption patterns. C. 176 Thin-layer chromatography (TLC). 574 Ulcerative colitis. 104 Trolox equivalents. 636. analytical methods for. 657 fiber values. 670 phytate content. 206–208 cereal product effects. 228–229 purified fiber effects. 513. 583 Southgate method for measuring. 68. 52–53 Toxicity. 277. See also Southgate method Undaria. 56 Unavailable carbohydrate. 220–225 gum and mucilage effects. 213–215 overall dietary pattern and. 676. 10. 52. 404–408. 87–105 applications. 455 Trowell. 64 Triglycerides. 241. See also Legumes. 37. 412 duodenal juices. 638. 490. 21. 455 antitoxic effects. 176–179 Vitamin PP. 578 USDA Dietary Guidelines 2000. 470 Verbascose. 178. 317. 262. 632–636. specific vegetables antioxidants. 365. 282–283 Vitamin A. 175 Vitamin K. 330. 184. 611 Seventh Day Adventists. 481 US National Cancer Institute. 660. 287. 667. 178 Vitamin metabolism. 173–179 enteral synthesis. 281–282 fiber effects on carbohydrate metabolism and. 93 starch removal. 469 mineral bioavailability studies. 69. 608. 260 anticarcinogenic effects. 185 fiber values. 260. 554 USDA Dietary Guidelines for Americans. 631. 401. 334 heart disease risk and. 657. 569 Uppsala procedure determination. 553 US Preventive Services Task Force. 177 Volatile fatty acids. 134 triglyceride hydrolysis and. 240 fecal/luminal pH and. See also Tocopherols fiber effects on bioavailability. 486. See also Lipid-modifying effects. 481 USDA Nutrient Database for Standard Reference. 317–318 Italian study. fiber effects on blood levels. 338 mineral bioavailability studies. 655–657. 176 Vitamin B6. 670 French consumption patterns. 78 national consumption patterns. allowed fiber health claims. 99–101 neutral polysaccharide constituents. 40. 16 Very low density lipoprotein (VLDL) cholesterol. 423–424 fecal composition effects. 504–514 Wheat bran. 305 fecal microflora and. 447 . phytic acid and. 101 Uronic acid residues. See Short-chain fatty acids W Wales. 636 Watercress. 404. 554 V Vegetables. 179 Vitamin C. 411. 46 Wheat. 598. 334. 90 uronic acid determination. 572 Walnut. 70. 584 Chinese consumption patterns. Obesity chitin/chitosan and. 97–99. 629 glycemic index values. 602 glycemic index values. 615 US Food and Drug Administration (FDA). 185. 175–176. 76. 446 Water-holding capacity. 17. 539–541 antioxidants. 176 Vitamin B2. 30. 312 US consumption patterns. 486. 35–36. 453. See also Pectin Englyst procedure determination. 666–667. 178. 482 antioxidants. 173–176 water-soluble. 178–179 fat-soluble. 31 Waxes. 584. 88–89 soluble polysaccharide precipitation. 557 Vegetarian diets colon cancer and. 93–96 Englyst method and. 101 Uruguay. 512 national consumption patterns. 36–38 protein digestibility and. 30. 655 Water-soluble fraction. 446 crude fiber values. 554 Weetabix.2387_Index_fm Page 707 Sunday. 601. 173–175 Vitamin B1. 624. 453. 175 Vitamin E. 220–225 fecal output and. 594–595 colon cancer and. 406–408. 670 fecal bulk and composition effects. May 6. 667 fiber values. 176 Vitamin B12. 191. 455–456 Australian consumption patterns. 413 Viscosity. 606. 161–162 fecal mutagens and. 90–92 sugar and lipid removal. 260 Watermelon. 19. 149. 556. 666–667. 97–99. 423–424. 569. 491 antitoxic effects. 594–595 nitrogen excretion and balance effects. 423–424 crude fiber values. 660 Weight loss. 143 short-chain fatty acids and. 521 Vitamin D. See also Ascorbic acid fiber and mineral bioavailability and. 334 grinding effects on glycemic/insulinemic responses. 151 protein digestibility and. 621. 453. antitoxic effects. 679 Water chestnut. 2001 8:07 PM INDEX 707 lignin determination. 321 nutrient absorption effects. 177 glycemic/insulinemic responses and. 174 vitamin C and. 321 crude fiber values. 147. 176 riboflavin absorption and. 411 microflora and. 279 cardiovascular disease correlations.2387_Index_fm Page 708 Sunday. 504–514. 233 fiber content by Southgate methods. 280 fecal composition studies. 101 vitamin A absorption effects. 616 fecal composition studies. 264 mineral bioavailability studies. 385 historical interest. 153. 279 cardiovascular disease correlations. 667 fecal microflora study. 521 phytate content. chronic diseases and. 481–493 antioxidants in. 482–486 antioxidant and phytochemical effects. 485 body weight effects. 103 Uppsala method collaborative study. 681–682 X Xanthan gum. 183–184 lipid-modifying effects. 339 mineral bioavailability studies. 474–475. 462. 228–229 food preferences. 455 crude fiber values. 482 diabetes risk and. 490 phytochemicals in. 176 short-chain fatty acids and. 150 phytate content. 553 laxative properties. 339 glycemic response and. 481. 233 fecal/luminal pH and. 303. 517. 382. 491–492 definitions. 470. 488–493 antioxidant effects. 469. 482 cancer and. 304 fecal mutagens and. 183–184. 467–472 disease risk factors and. 302. 667. 380. 680 White bread carbohydrate hydrolysis. 488 Wine. 228–229. 487–488 disease incidence and. 678 Whole-grain bread carbohydrate hydrolysis. 296. 382 short-chain fatty acids and. 2001 8:07 PM 708 CRC HANDBOOK OF DIETARY FIBER IN HUMAN NUTRITION. 486 antithrombotic effects. 485 hypotensive effects. 675 Whole-grain foods. 184. 309. White bread. 664. 301. 279 protein utilization effects. 260. 326. 33 grinding effects on glycemic/insulinemic responses. 40. 667. Whole-grain bread Wheat flour. 374. 318 fiber values. 465–468 epidemiologic evidence. 330. 288–294. 482 weight loss effects. 482–483 insulin sensitivity effects. May 6. 483–484 lipid-modifying effects. 261. 24 diabetes treatment effects. 490–491 gastrointestinal effects. 263. 278. 486 whole grain and refined cereal comparison. 263 fiber content. 307 . 185. 486 mortality from all causes and. 338 phytate content. 519–520 nitrogen excretion and balance effects. 670 diabetes treatment effects. 326. 409–410 fecal bulk and composition effects. 627. 472–474. 667 fiber values. 282 fecal bile losses and. 311 Uppsala method applications. 140. 467. 33 glycemic index. 481. 489–490 phytoestrogen effects. 156 pyridoxine bioavailability and. 384. 330. 489–490 insulinemic effects. 461–477 heart disease and. 374 phytate content. 280. 191. See Breads. 475–476 omega-3 fatty acids. 307. 178 Wheat bread. 664 lipid digestion/absorption effects. 239 glycemic/insulinemic responses and. 281. 41. 467 dietary fiber values. 628 food preferences. 193–201. 461–477. See Flours Wheat germ crude fiber values. 670 folic acid and. 670 fecal composition studies. 3RD EDITION carbohydrate metabolism and. 374. 471 crude fiber/dietary fiber comparison. 504–514. 660 food preferences. 3–4. 281 phytate content. 328 digestive enzymes and. 492–493 mechanisms. 300. 675 White rice antioxidants. 381. 409–410. 33 glycemic index values. 489. 327 digestive enzymes and. 679 protein digestibility and. 12. 633 Yugoslavia. 334 fiber values. 31 digestive enzymes and. 45 crude fiber values. 680 Yucca. 667 glucose/insulin responses. 103 Xylose. 661 phytate content. 278 fecal composition studies. 517–520 phytate effects on bioavailability. processing effects on. 162 dietary fiber and absorption. 93. 573 Z Zinc. 40 in vitro binding studies. 636. 667 fiber values. 263. 31 Xyloglucan. 279 protein utilization effects. 105 Y Yam. 503 bioavailability studies. 571. 265 protein digestibility and. 162–164. 339 phytate content. 212 fecal microflora and. 136. 162–164 . 504–516 deficiencies. May 6. 127.2387_Index_fm Page 709 Sunday. 499. 657. 680 Yeast. 153 Xylem. 2001 8:07 PM INDEX 709 Xylans. 161. 2001 8:07 PM .2387_Index_fm Page 710 Sunday. May 6.