Davis & Panksepp (2011)

Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958Contents lists available at ScienceDirect Neuroscience and Biobehavioral Reviews journal homepage: www.elsevier.com/locate/neubiorev Review The brain’s emotional foundations of human personality and the Affective Neuroscience Personality Scales Kenneth L. Davis a,∗ , Jaak Panksepp b a b Department of Psychology, The University of North Carolina at Charlotte, Charlotte, NC 28223-0001, USA Department of Veterinary and Comparative Anatomy, College of Veterinary Medicine, Washington State University, Pullman, WA 99164-6520, USA a r t i c l e i n f o Keywords: Personality Affective neuroscience Five Factor Model Subcortical brain emotion systems Affective neuroscience Affective Neuroscience Personality Scales a b s t r a c t Six of the primary-process subcortical brain emotion systems – SEEKING, RAGE, FEAR, CARE, GRIEF and PLAY – are presented as foundational for human personality development, and hence as a potentially novel template for personality assessment as in the Affective Neurosciences Personality Scales (ANPS), described here. The ANPS was conceptualized as a potential clinical research tool, which would help experimentalists and clinicians situate subjects and clients in primary-process affective space. These emotion systems are reviewed in the context of a multi-tiered framing of consciousness spanning from primary affect, which encodes biological valences, to higher level tertiary (thought mediated) processing. Supporting neuroscience research is presented along with comparisons to Cloninger’s Temperament and Character Inventory and the Five Factor Model (FFM). Suggestions are made for grounding the internal structure of the FFM on the primal emotional systems recognized in affective neuroscience, which may promote substantive dialog between human and animal research traditions. Personality is viewed in the context of Darwinian “continuity” with the inherited subcortical brain emotion systems being foundational, providing major forces for personality development in both humans and animals, and providing an affective infrastructure for an expanded five factor descriptive model applying to normal and clinical human populations as well as mammals generally. Links with ontogenetic and epigenetic models of personality development are also presented. Potential novel clinical applications of the CARE maternal-nurturance system and the PLAY system are also discussed. © 2011 Elsevier Ltd. All rights reserved. Contents 1. 2. 3. 4. 5. Introduction: Darwin, McDougal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subcortical emotion systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Five Factor Model and the ANPS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1. Cloninger’s biologically based personality theory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2. ANPS data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2.1. Conscientiousness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2.2. Non-human personalities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3. Expansion of the FFM . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.4. Examples of studies using ANPS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Affective neuroscience trait versus theoretical expansion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1. Primary affective consciousness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.2. Could the fundamental nature of personality be affective? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3. State vs. channel functions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.1. Evidence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.2. Social rejection and the SADNESS system . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.3. Evidence summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clinical assessments from affective neuroscience trajectories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ∗ Corresponding author. Tel.: +1 336 379 9828; fax: +1 336 379 9835. E-mail address: [email protected] (K.L. Davis). 0149-7634/$ – see front matter © 2011 Elsevier Ltd. All rights reserved. doi:10.1016/j.neubiorev.2011.04.004 1947 1947 1948 1948 1949 1951 1952 1952 1952 1953 1953 1953 1953 1954 1954 1954 1954 . . . . . neurochemical. . . . . . . . . . . The capitalizations were premised on the need for a specialized terminology for the primary-process anoetic experiential processes of the brain. . . . . . . . As ruefully related in his autobiographical materials. with emotional behaviors being manifested in extreme. . . . . . . . . . with regrets. . . . . along with characteristic accompanying autonomic arousals. . . because of their capacity to engender rewarding and punishing states in the brain. 1998) – from mice to men. We sustained the capitalization convention in our attempt (i. . . . . . . . . . . . . . . . . . . . . . . although those strands of early ethological thought were never fully developed. . . . . . LUST. . separation-distress PANIC/GRIEF. . . Likewise.e. . . and hence abnormal displays. . . . . . . . . . . He included only seven behaviorally well-defined emotional instincts. . selfassertion. “simple instinctive impulse. . . and proceeded to show that. . . . . . . . . 2011). . . . 42). . . J. . . . cross-species affective neuroscientists have started to map out the continuity of primal emotional systems in subcortical-limbic circuits of selected mammalian brains (Panksepp. . . . . . . . . . . and the parental instinct. . . . . . . . . . . . . curiosity. repulsion. . 2005) synthesis of such work provides documentation for the existence of seven “blue ribbon” subcortical emotion systems in the brain. . . . . . . . . exaggerated. . . . . . . Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 1947 Affective neuroscience and therapeutic effectiveness . . which together constitute forms of affective consciousness (Panksepp. . which is consistent with certain current views of mental disorders (see below). . . . . Since it is widely believed that individual’s emotional inclinations (temperaments) are foundational for human personality development. . . . but “mental” processes such as basic feelings and perceptions (anoetic qualia). in synergy with Darwin’s vision. .4. . . . . . . .” The first was whether the emotion was “clearly displayed in the instinctive activities of the higher animals” (McDougal. see Vandekerckhove and Panksepp. . . . 2007a) This nomenclature allows the use of common emotional terms that convey the gist of the emotional meaning. FEAR. self-abasement. . . . 1955 1955 1956 1956 1956 1. . . . p. . . . . rather than RAGE and PANIC/GRIEF. . . . . pugnacity. Appendix A. . . . . . . Acknowledgements . . aspects of mind that can only really be well-studied in humans (for a coverage of such nomenclature issues. which control learning and memory (noetic secondary-processes) as well as higher mental abilities (autonoetic tertiary processes) such as thoughts. . . . . . . . . . . . . . . . Summary of affective neuroscience perspectives on the foundations of personality . . . . . . Hess resisted. . . This is the classic dilemma that subjectivity poses for modern neuroscience. His second principle was that a primary emotion should be evident in human psychopathologies. . . In contrast. . . . But before we summarize work with the Affective Neuroscience Personality Scales (ANPS). he rejected that there was any religious instinct. . copyright 2004 version . . . . . . . . can be objectively studied in a substantive phenomenological manner. . because he feared marginalization by behaviorists. . . . grief. . we briefly note historical antecedents of this way of thinking about human personality. . . . a list based on comparative observations that is arguably still relevant to personality theory a century later. . . However. In 1908 William McDougal published his classic work “Social Psychology” in which he reviewed the influence of instincts on human personality. In this classic work. . . . the theoretical implication is that these systems are critically important for the various feelings highlighted by these sci- . . . All mammalian brains may be fundamentally subjective organs – they not only generate behaviors. . . . . . . . .e. . . a recent spectrographic analysis of the tickle-induced laughter response on four species of young great apes plus humans yielded data that effectively mapped the scientifically established evolutionary branching of these closely related anthropoids (Ross et al. . . . . . . . . . . . . . .. . . feelings. . . . . . . . so to speak. .6. . the goal of the following work was to try to ground personality assessment on the affective circuits that all mammals share. . animals would escape the RAGE evoking stimulation. . . . . . . . . Scoring the ANPS 2. . . .. . . . . . 2009).L. . . . . . . . . . . . . . . . . . . . we decided to use the terms ANGER and SADNESS in this paper and in our construction of the Affective Neuroscience Personality Scales (ANPS). maternal CARE. . . . . while minimizing mereological fallacies (part-whole confusions) that are endemic to the use of emotional terms in describing animal experiences. . . The anatomical. which included a discussion of playful tickling in human children and many other descriptions of human emotional subtleties. . 1872/1967). . . . . . . Darwin’s keen observations. . 2. . . . . . . . . . . and various other debatable higherorder affective-cognitive processes. which he labeled flight. . . the ANPS) to translate anoetic (without knowledge) emotional experiences to noetic (knowledge linked) and autonoetic (thought linked). . Davis. . . We will never know how any other animal or any other human actually feels. . . 1908/1963. which remain relevant to this day. . and physical PLAY. . allowed him to extend his robust case for the “continuity of species” from body structures to the domain of emotions. . . . . . . . . . . . . . . and joy. He indicated that he always believed these irritated animals were experiencing a state akin to human anger. . . he received the Nobel Prize for his work on the central representations of the autonomic nervous system. . . . . Still. K. . . . 1998. . . Neuroscience advances will be substantially premised on our ability to conceptualize brain functions both from the external behavioral view as well as the internal psychological perspective. .” i. . . . . plans and insights. . . . . . . . . . . . . language-based evaluations of the potential influences of primal affects in the personality structures of human minds. . . and functional homologies of subcortical emotional networks strongly indicate underlying evolutionary continuities in affective principles in all mammalian brains (Panksepp and Biven. . . He was the first personality theorist to be guided by Darwin’s principle of the continuity of human and animal mental evolution. . . . . . . . . . . . 2009 and this issue). . . . . which he labeled SEEKING. . . . . . . . . In line with his views. . . . . . . . . given the opportunity. . . . Introduction: Darwin. . . . . . . . . 7. . Subcortical emotion systems Walter Hess’s seminal work on evoking emotions with localized electrical stimulation of the brain (research summarized in Hess. . . . . . thus providing some distance from dictionary definitions and everyday understandings. . . . . . . . . . based on whether the ESB (electrical stimulation of the brain) evoked ‘reward’ or ‘punishment’ effects in simple learning tasks. Panksepp (1971) later demonstrated an homologous anatomy in rats. . . any explicit discussion of emotional experiences in the animals he studied. since those are labels that are generally more understandable for most individuals. McDougal Charles Darwin provided the first modern scientific treatise on emotion in 1872 with “The Expression of the Emotions in Man and Animals” (Darwin. RAGE. . . . . . . . . . . . . . . . . . . . . . . . In 1949. . . . . Hence. . . . . . . . . with explicit discussions of the accompanying affective experiences. . . . 1957) demonstrated the importance of discrete subcortical areas of the brain for generating specific emotional-behavioral episodes. . . . . and he used two principles for determining whether an emotion should be considered a “primary emotion. . . . . . . . . . . . Panksepp’s (1982. . . . . . He never inquired whether the animals found such evoked states of anger to constitute aversive experiences. . Darwin documented cross species commonalities that occurred for many primary emotions such as fear. anger. Panksepp and some others continued these lines of research. . . References . The FFM has provided a personality assessment standard. but three (CARING. while learning and higher brain functions can be deemed to be secondary and tertiary processes.g. FEAR and LUST). as our understanding of brain affective systems has become more resolved. we will only contrast our work on the ANPS. 1990).. emotional responses from each of the primary process emotions can be activated by localized subcortical ESB or chemical brain stimulation. with the cross-species emotion systems of mammalian brains.1948 K. Each of these seven emotions can be evoked by stimulating distinct but partly overlapping subcortical regions of mammalian brains. Specifically. (1992) and consisted of an analysis of over 500 adjectives placed on five orthogonal dimensions as well as numerous off-axis.” such as “angry” emerging as a “blend” of low Emotional Stability and low Agreeableness.1. there are reasons to believe the SEEKING system is foundational for aspects of all of the other primal emotions (e.. studies of primary-process emotional aspects of personality may provide a heuristic intersection for the Darwinian “continuity of species”. Agreeableness. The two individuals who have attempted to construct neurobiologically based personality assessment instruments in the past are Cloninger (1986) who based his thinking on functional animal work on some of the first well characterized neurochemical systems of all mammalian brains – dopamine. during FEAR intensification. As such. and double capitalized BrainMind to reflect our monistic ontology).g. the shift from freezing to flight may be due to recruitment of SEEKING related dopaminergic psychomotor drive). The present work is an extension of this strategy. That decortication of young animals generally leaves the expression of these emotions intact (Deyo et al. 1998). 1986. the ANPS was hypothesized to provide affective underpinnings to the widely used Five Factor Model (FFM). 1990) and have been replicated in several languages (Saucier and Goldberg. It is a major supposition of this paper that these basic emotions are fundamental powers of the human BrainMind. pharmacological. 1994). (see Panksepp. Neuroanatomical. and hence potentially of critical importance for understanding the foundations of personality (we intentionally conflate ‘brain’ and ‘mind’. CARE.. It is our premise that these emotions form an important foundation of personality. as in the ANPS (Davis et al. 1998). that could also open up a common terminology for study personality across species. which was conceptualized largely as a research tool aspiring to. Panksepp et al. 2006). 2011).L. However. Emotional Stability. situating adult human temperaments within the complexities of primary-process affective space shared by all mammals. two-factor “blends. 3.. 1990. but surely with antecedents in “lower” vertebrates. In contrast. see Huber et al. Also. (5) The integrity of these seven systems is demonstrated by the ability to elicit coherent specific emotional responses and/or the associated affects with localized brain stimulation – as evaluated by the capacity of the subcortical arousals to mediate ‘reward’ and ‘punishment’ functions that control learning. 1992). and the resulting sensitivity to threatening situations. 1998. neurochemical. and serotonin (see below). and Intellect or Openness to Experience (Hofstee et al. see Panksepp. 2003). and SADNESS. Several of these emotions have evolutionally deep reptilian roots (SEEKING. We assumed these brain systems could serve as an empirical roadmap for advances in the study of personality and serve as a template for emotional personality assessment. with the popular Cloninger assessment tool following a brief introduction of the five factor personality model. norepinephrine. 2011 for detailed reviews). and the need for neurobiologically meaningful psychological assessment of human temperaments/personalities. However.. which while not universally accepted (Block. But this is not the first attempt to bridge basic neuroscience and human personality evaluation. 1987) and the Tridimensional Personality Questionnaire (TPQ). The FFM has origins in adjective descriptors of personality (Allport and Odbert. personality assessment can be informed by these subcortical affects. 1998. the brain’s presumed punishment system (which is highly debatable. reward and punishment) systems in the brain. Each of these “temperament” traits was seen as highly heritable and stable across the developmental stages of life. the ANGER system energizes states commonly labeled as ‘irritability’ and ‘anger’. 1947). ANGER. 2011 for fuller descriptions). each instinctual emotional system engenders affective valence since animals terminate the stimulation of ANGER. primal affects that accompany those responses. 2005. and PLAY system arousals (Panksepp. they typically work interactively in concert to increase the adaptiveness of individual feelings. thoughts. while actively working to obtain SEEKING. the first four dimensions have been more robust with Openness to Experience typically accounting for the least variance of the five factors (Goldberg. These five dimensions have been consistently derived from a variety of descriptive data using factor analysis (Digman.. (4) These brain systems generate instinctual behavioral responses that are closely linked to the raw. perceptions. 2005. Panksepp and Biven. In summary. A comprehensive presentation of the five factor “descriptive” model was provided by Hofstee et al. Because of space constraints. 1936) and the early history of factor analysis (Cattell. these emotion systems. However. and perhaps capable of. and behaviors. and (3) Reward Dependence featuring behavioral maintenance and sensitivity to reinforcement contingencies with norepinephrine as the major neuromodulator. LUST. The Five Factor Model and the ANPS At the very outset of this work. SADNESS. these systems remain relatively unscathed in animals whose neocortices were surgically removed in early development (see Panksepp. .. further reinforces the subcortical nature of these emotion systems. Carver and White (1994) developed the Behavioral Activation and Behavioral Inhibition Scales on the basis of Jeffrey Gray’s (1981) early brain research recognizing the existence of behavioral approach/arousal and withdrawal/inhibition (aka. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 entific terms – e. (3) These basic emotions also have similar chemistries in all mammals. Panksepp and Biven. Davis. 2002). There are several principles basic to understanding these seven brain systems: (1) These emotion systems are subcortical networks and lower brain regions have evolutionary primacy in generating these basic emotions and their affects. 2005). Cloninger’s biologically based personality theory Cloninger was the first to attempt an integration of personality and brain systems with his biogenic amine-based “biosocial” theory of personality (Cloninger. situated in ancient brain regions.g. and physiological brain research have helped to more precisely define each of these seven emotion systems (see Panksepp. (6) Lastly. and PLAY) are more uniquely mammalian adaptations. are largely homologous in all mammals. The five personality dimensions are usually labeled Extraversion or Surgency. (2) Harm Avoidance. (2) To the best of our knowledge. Conscientiousness. FEAR. If so. perhaps invertebrates also (e. 1995) has offered a robust personality model in general population and clinical research (Costa and Widiger. which included three scales: (1) Novelty Seeking based on a postulated dopaminergically based behavioral activation system that promotes interest in new experiences. J. There is strong evidence for these seven well-defined emotion systems in all mammalian brains. 3. postulated to primarily reflect brain serotonin activity. designed to estimate six affects relevant for defining temperamental/personality dimensions. homologous with feelings experienced by other mammals.. We excluded LUST..30 was ANGER.0 with FEAR. 1999). 80) that higher order factors such as Positive Affect. A factor analysis of just the six primary ANPS scales yielded two eigenvalues greater than 1. and thinking about loved ones and past relationships including crying. were reported ranging from . as already noted. the ANPS offers personality primes that align with the Big Five model but target the six specific primary emotion brain systems that have been well documented with comparative brain research. 1988) and generally replicates Gray’s (1970) Behavioral Inhibition and Approach Systems. 2008a. see Appendix A). it is also important to note Goldberg and colleague’s argument (Ashton et al. Other research found that the Self-Directedness character scale also responded to serotonin treatments (Peirson et al.. The SEEKING system not only promotes exploration.. 1985) or pharmacologically as well as through brain damage research (e. The PLAY scale focused on playing social games with physical contact plus laughter. SADNESS was conceptualized as feeling social separation distress. Wai and Bond. 3.2. which also loaded −. and ANGER on the first component and PLAY. (5) SADNESS with Emotional Stability (r = −. tertiary-process type BrainMind functions). 2003. have shown all reliabilities now over . 1 (for details. Since there were only four eigenvalues > 1.. (3) PLAY and Extraversion.65) as well as with Agreeableness (r = −.68 being the lowest primary factor loading.K. and losing sleep.46).L. and hence did not support a distinction between “temperament” and “character” dimensions. The Spirituality scale focused on feelings of connectedness with all of life and oneness with creation. When factor analyzing ANPS scales with FFM scales. and expressing anger verbally or physically. Cloninger subsequently expanded his assessment instrument adding an additional temperament dimension – Persistence – and three controversial “character” traits – Self-Directedness.. The only scale having a loading on a second factor greater than . 2009. and accounted for the revised TCI scales with five factors (Farmer and Goldberg. In summary. CARING centered on nurturing tendencies including liking to care for others. Panksepp. (4) FEAR with Emotional Stability (r = −. investigation and foraging but probably ‘energizes’ all basic emotional systems with forms of appetitive and anticipatory arousal (e.0. Based on our first study using these scales. Davis. 1999. which was consistent with the hypothesis that these six brain emotion systems form a foundation for the adult five-factor personality model. p.80 (Davis et al.55 and all secondary loadings less than . even though they figure heavily in the arousal dimensions of all personality control systems. as seen in Table 1. ANGER. In addition. individual psychotherapy practitioners) without need to obtain permission of the copyright owners (Davis and Panksepp).53 on the Agreeableness factor. Each ANPS dimension corresponded to one of the “blue ribbon” primal emotions identified by cross-species ESB-based affective neuroscience research.48) (Davis et al. We suspect that serotonin and norepinephrine may function too broadly to be isolated as specific contributors to very specific types of affects the way Cloninger proposed.. and thus may contaminate frankness on the other scales. and feeling softhearted towards animals and people in need. heritabilities did not differentiate the temperament and character scales (Gillespie et al. and generally having fun. Those wishing to use it for commercial use are requested to contact the first author. computed as Cronbach’s alpha. revealed that each of the ANPS scales except Spirituality correlated strongly with at least one of the FFM scales as follows: (1) SEEK with Openness to Experience (r = . ANGER. Multiple data sets using the revised ANPS 2. and (4) SEEK and Openness to Experience loading on four separate scales with . feeling tense. (3) CARE with Agreeableness (r = . Although Cloninger’s earlier TPQ factors have been related to many genetic variables. which seems less relevant to current conceptualizations of human personality and we also suspected that it may potentially be an affective factor that people would not wish to be frank about. 2003. Norepinephrine promotes attention by amplifying signalto-noise processing in most sensory-perceptual channels and also promotes emotionality in general. and Self-Transcendence – hypothesized to have low heritabilities and be more sensitive to individual development (we would conceptualize these as higher-order. The data supported strong relationships between primary emotions and the most widely accepted model of human personality. many of Cloninger’s later theoretical claims have not been supported. ANPS data The Affective Neuroscience Personality Scales (ANPS) were introduced to explore the potential primary-process brain emotional system foundations of personality. feeling lonely. Cooperativeness. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 Our affective neuroscience perspective would align the Novelty Seeking dopaminergic brain incentive dimension with the SEEKING system. De Fruyt et al. We feel this is a striking demonstration of the multifactorial nature of any general positive and negative valence type of measurement tool. difficulty making decisions. These scales are provided for free use in all scientific endeavors and non-commercial clinical use (e. CARE.86 with the PLAY and SEEK scales below . Bechara et al. 2000). 2004). The scales were rationally defined to potentially provide biological underpinnings for the descriptive FFM of personality (Davis et al. a Spirituality scale was included to acknowledge the importance of this factor in drug addiction (Panksepp et al.. 12-step programs) and what might be considered the highest human emotion. SADNESS.50).. humor.65 to . serotonin dampens and restricts neural processing in the brain. a detailed description of the scales and scoring instructions are provided in the appendix at the end of this paper. Contrasting self-rating data for ANPS and FFM scales. 2003). Cloninger. Our view is that these personality dimensions escape the problem of definitional circularity since comparative research has demonstrated how each primary dimension can be manipulated in humans as independent variables including experientially (Eisenberger et al. In contrast..g.b).g.. The revised instrument was called the Temperament and Character Inventory (TCI. reliabilities for the ANPS scales.20.70 (unpublished data). 1999. SEEKING was defined as anticipating new positive experiences including being curious. and (6) ANGER with Emotional Stability (r = −. In this vein. (2) CARE and Agreeableness.4 items are listed in Fig.. worrying.47). being easily irritated and frustrated. Again. Others also reported considerable convergence between Cloninger’s revised TCI (Cloninger.. Negative Affect. being drawn to young children and pets.g.4. 2004) and many drug addiction recovery programs (e. The revised ANPS 2.. ANGER included feeling hotheaded. liking to strive for solutions to problems. and overall Emotional Stability “may represent artifacts rather than substantive dimensions of personality. 2001).70 and the FEAR. 1999). 2003). seeking of safety when FEAR is aroused). Cloninger.68). corresponds nicely to other scales which simply try to parse human personality into positive and negative affective dimensions (Watson et al.75). 2003). (2) PLAY with Extraversion (r = . The FEAR scale incorporated experiencing anxiety.g. and Emotional Stability. and SEEK on the second with all primary factor loadings greater than . 1986). we reported a five factor solution with the FFM Conscientiousness scale on a factor by itself. ruminating. This clustering.” . J. resulting generally in reduced emotionality (Panksepp. Factor analysis placed the temperament Harm Avoidance and the character Self-Directedness on the same factor.. In other words. we removed the Conscientiousness scale and published a four factor solution with (1) FEAR SADNESS. 1999) and the FFM (Ball et al.. and Spirituality scales above . and gener- 1949 ally liking to explore.. 27. 36. 23. and correlations with the FFM. I am often spiritually touched by the beauty of creation. I enjoy anticipating and working towards a goal almost as much as achieving it. if I have nearly met my goals. 46. My friends would probably describe me as being too serious. Feeling a oneness with all of creation helps give more meaning to my life.g. I often have the feeling that I am going to cry. FEAR. People who know me would say I am a very fun-loving person. Almost any little problem or puzzle stimulates my interest. I often feel like swearing.anps. I generally do not like vigorous games which require physical contact. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 Pahlavan et al. 15. Fig. J. I often feel sad. I love being around baby animals. I seem to be affected very little by personal rejection. 4. 54. I usually feel little eagerness or anticipation. 29. If my peers have outperformed me.4 items and response scale. For me. 56. factor structure. Davis. I never stay irritated at anyone for very long. 2008a. Affective Neuroscience Personality Scale 2. 11. Caring for a sick person would be a burden for me. 42. . 38. When I get angry. 26. CARE. Through personal correspondence. Inc. Berthoz and colleagues (personal communication) replicated the Pahlavan et al. 51. 13. 40. I think it is ridiculous the way some people carry on around baby animals. Contemplating spiritual issues often fills me with a sense of intense awe and possibility. I am known as one who keeps work fun. I usually avoid activities in which I would be the center of attention.. Ph. I usually get angry. spirituality is not a primary source of inner peace and harmony. 16. 14. 31. I rarely rely on spiritual inspiration to help me meet important challenges. People who know me well would say I am an anxious person.. For a while Martin Reuter (University of Bonn) had all the original scales readily available on the web. When someone makes me angry. 17. I dislike humor that gets really silly. experiencing a connection to all of life is an important source of inspiration. 5. 45.D. My friends would probably describe me as hotheaded. For me. 1. 19. I often think of what I should have done after the opportunity has passed. 10. 9. I cannot remember a time when I became so angry that I wanted to break something. and has already proved its utility in evaluating certain psychiatric problems (e. 20. and the comparative fit index = . see below). When I play games. mdanps24 47321 Copyright © 2004. I tend to remain fired up for a long time. I often think about people I have loved who are no longer with me. 6.4 Name: Age: Sex: Please mark bubbles like this Disagree Agree Str Disagree Str Agree 1. All rights reserved. 39. 30. When I am frustrated. Savitz et al. I like to joke around with other people. 37. I always tell the truth. and also for data collection purposes (www. 28.L. (2008) findings on a large sample (n = 830) and also performed an item-level confirmatory factor analysis obtaining an acceptable fit modeling 6 factors (standardized root mean square residual = . 25. I would not describe myself as a worrier.b. 49. I will gossip a little at times. 32. Feeling like a part of creation is not an important source of meaning for my life. 52. 21. (a) Affective Neuroscience Personality Scale 2. I have very few fears in my life. but this site has been closed. I feel softhearted towards stray animals. 35. 47. Seeking an answer is as enjoyable as finding the solution. 53. 50. 34. Kenneth L.1950 K. I rarely become sad. 12. ANGER. Sometimes I feel like swearing. The ANPS items are arranged in fourteen blocks using the following item sequence: SEEK. I often feel lonely. When I am frustrated. 33.78). I am usually not highly curious. 43.06.D.. 41. I like to be the one in a group making the decisions. 2. (2008) confirmed the psychometric properties of the ANPS in a French population including scale reliabilities. 8. Jaak Panksepp. I am a person who is easily amused and laughs a lot. I sometimes cannot stop worrying about my problems. root mean square error of approximation = . and Spirituality. SEEK. 24. I never become homesick. SADNESS as described in Appendix A. 44. I rarely become angry.039. I am not frequently jittery and nervous. I would still be happy. Pegasus International. I do not get much pleasure out of looking forward to special events. (b) ANPS continued: see description in above figure legend and Appendix A. Davis. I often cannot fall right to sleep because something is troubling me. PLAY.de).. 7. Ph. it is important for me to win. I really enjoy looking forward to new experiences. 3. we know the scale has been translated into a dozen languages. 18. I like taking care of children. 48. (2008c) obtained significant evidence of heritability for four of the ANPS scales: CARE. I do not especially like being around children. FEAR. 55. I often feel a strong need to take care of others. 22. Savitz et al. 112. Fear of embarrassment often causes me to avoid doing things or speaking to others.24). I do not particularly enjoy kidding around and exchanging "wisecracks.L. Congdon and Canli’s (2008) view of impulsiveness as a “top down” lack of behavioral inhibition involving the inferior frontal cortex and subthalamic nucleus fits well with King’s (2007) description of chimpanzee Conscientiousness – characterized by predictable. Ph. Feeling a connection with the rest of humanity motivates me to make more ethical choices. All rights reserved. My sense of significance and purpose in life does not come from my spiritual beliefs. 106. I almost never lose sleep worrying about things. 63. (Continued ). When I play games. I often worry about the future." 62. 97. and SADNESS (r = −. There are very few things that make me anxious. it seems unlikely that a single primary brain affect serves as the foundation for Conscientiousness. 89.K. Conscientiousness Conscientiousness was the only FFM dimension that did not correlate strongly with the ANPS. 64. 110. I feel sorry for the homeless.. My curiosity often drives me to do things. Kenneth L. 67. 104. not impulsive or erratic. 80. Sometimes little quirky things people do really annoy me. I am very playful. 93. 98. I sometimes feel like kicking or hitting something. Davis. 75.30) suggested that while Conscientiousness may be associated with the regulation of negative affects. 60. I am usually not interested in solving problems and puzzles just for the sake of solving them. 107. 72. 1999). From animal FFM studies.. Conscientiousness only appeared in chimpanzees (Gosling and John. Inc. 108. FEAR (r = −. 101. I do not mind losing. 105. 87. People who know me well would say I almost never become angry. 94. . 66. J. Being embarrassed or looking stupid are among my worst fears. 109. 61. I tend to get irritated if someone tries to stop me from doing what I want to do. My choices are not guided by a sense of connectedness with all of life. When things do not work out the way I want.2. 99. I hardly ever become so angry at someone that I feel like yelling at them. I am not satisfied unless I can stay ahead of my peers. 103. I rarely feel the need just to get out and explore things. I am a person who strongly feels the pain from my personal losses. I see life as being full of opportunities to have fun. 59.30). which suggests that Conscientiousness is a more cerebral dimension emerging late in mammalian evolution. 82. 88. I would generally consider pets in my home to be more trouble than they are worth. I rarely have the feeling that I am close to tears.D. I often feel like I could accomplish almost anything.D. I do not tend to see the humor in things many people consider funny. Striving to be better than my peers is not important for me. 78. Playing games with other people is not especially enjoyable for me. 76. 65. 84. 71. I like to explore the area and get a better feel for my surroundings. Lower correlations with ANGER (r = −. Spiritual inspiration helps me transcend my limitations. I like having authority over others. 73. I do not like to feel "needed" by other people. I am the kind of person that likes to touch and hug people. 90. 79. Whenever I am in a new place. I have never intentionally told a lie. I rarely feel the urge to say nasty things to them. 58. 81. I am not the kind of person that likes probing and investigating problems. I rarely worry about my future. 83. 70. 1. I am not particularly affectionate. There have been times in my life when I was afraid of the dark. 85. I like all kinds of games including those with physical contact. Fig.1. It would not bother me to spend the holidays away from family and friends. 95. I am a person who strongly feels the pain of other people. I frequently feel downhearted when I cannot be with my friends or loved ones. behaviors. Ph. 91. It does not particularly sadden me when friends or family members are disapproving of me. My friends would say that it takes a lot to frighten me. Davis. Pegasus International. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 Please mark bubbles like this and not like this or Disagree Str Disagree 1951 Agree Str Agree 57. I tend to think about losing loved ones often. When working on a project. 111. The goals I set for myself are not influenced by my spirituality. I rarely get angry enough to want to hit someone. 74. 92. 100. 86. 69. I often feel nervous and have difficulty relaxing. I rarely think about people or relationships I have lost. 77. 68. 96. I often worry about whether I am making the correct decision. 47321 mdanps24 Copyright © 2004. I do not especially want people to be emotionally close to me. When people irritate me. I am not an extremely inquisitive person. 3. Jaak Panksepp. I have never "played sick" to get out of something. I do not frequently ask other people to join me for fun activities. 102. J.50* −. which makes Emotional Stability equivalent to Negative Affect.46* . nurturing feelings on the positive pole and hostile. 2003). we interpret the ANPS scales as tertiary (thought-mediated) approximations of the influence of the various primary emotional systems in people’s lives. These results provide encouraging evidence that the FFM reflects primary subcortical affects. MRI data on a subsample of subjects revealed that ANGER scores correlated negatively with gray matter volume in the left amygdala. The construct validity of the ANPS ANGER scale was also supported by Reuter et al.001.. two-tailed. Examples of studies using ANPS Several of the ANPS scales have been further validated by independent research. 3. The most consistent FFM temperament dimensions observed were Extraversion and Emotional Stability. one should be able to identify these same FFM dimensions in non-human mammals.75* −. 2003).65* −. with CC genotypes having significantly lower ANGER scores. * p < .47* . 32 kDa (DARPP-32) is a key regulatory molecule in the dopaminergic signaling pathway. 1872/1967.and cAMP-regulated phosphoprotein. and anger (Vytal and Hamann. 1997)).. demanding feelings on the negative pole. only the SADNESS scale continued to show significant differences across groups.07 −. 3. and lowest for the two control groups. which could lead to statistical lumping.48* −.12 . and higher ANGER scores than unaffected family members. Savitz et al. but that does not mean it is desirable to combine three of our most powerful emotional drivers into a single dimension. since all self-report assessments must include cognitive reflection.08 −.” (Darwin. Savitz et al.01 . the evidence for Openness to Experience was weaker than for the previously named three dimensions.21 .13 . 1993)) or mania (Altman Self-Rating Mania Scale (Altman et al. 121 females). (2008b) also studied the distribution of hypomanic. p.01 . Consistent with human data (Goldberg. and ANGER.30* . the two scales correlated in opposite directions with the FFM Agreeableness scale.. SADNESS. (2009) investigating links between ANGER and the rs907094 (aC → T single nucleotide polymorphism of the DARPP-32 gene).25 −.17 Student sample: n = 170 (50 males. human’s closest evolutionary relative. Agreeableness was the next most common. can generate individual differences in normal personality as well as the affective imbalances characterizing mental disorders.1952 K. 2010). SEEK. Our view is that while the FFM is an important theory-free. Despite the fact that the ANPS ANGER and CARE did not correlate (r = −..05 −. Although ANPS items attempt to address primary affects directly.01 −. the ANPS SADNESS and FEAR scores trended highest for individuals diagnosed as BPD I or II. At the tertiary process level. certainly is one of degree and not of kind. As noted previously. Reuter’s group has also recently observed that the homozygous long-variant of the serotonin transporter polymorphism and the TT variant of the single nucleotide polymorphism rs2268498 on the oxytocin receptor gene showed significantly lower scores on the personality dimensions FEAR and SADNESS of the ANPS.19 −.3. and ANGER. Davis. studying families with a member having bipolar disorder (BPD).04 −. fear. After controlling for age.68* . Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 Table 1 ANPS Scale correlations with Five Factor Model scales. A recent meta-analysis of past brain imaging studies also provided strong evidence that distinct anatomical systems in human brains can be related to basic affective processes including support for sadness. than carriers of the other serotonin transporter and oxytocin gene variants (Montag et al.17 −. Human minds seem to generate a descriptive “love/hate” Agreeableness construct with caring. In a review of 19 studies of personality in non-human animals. Dopamine. PLAY SEEK CARE FEAR ANGER SADNESS Spirituality Extraversion Agreeableness Conscientiousness Emotional stability Openness to experience . .14 .L. 2005).036 in Davis et al.15 . and hostile traits in BPD probands and their families. ANGER also correlated with Agreeableness.2. it reflects a tertiary cognitive-type reconfiguration of the underlying primary brain emotion systems. BPD II patients had significantly higher ANPS ANGER scores than individuals with only a single previous depressive episode.13 . selecting 14 markers for each Big Five dimension (data adapted from Davis et al. Conscientiousness was only observed in chimpanzees. (2008a) found that the ANPS SADNESS scale was significantly higher in BPD I diagnosed individuals than two control groups made up of unaffected family members or family members with a DSM-IV diagnosis other than depression such as alcoholism or generalized anxiety disorder.29* −.06 −. where the foundations of emotions reside. However. cyclothymic. Consistent with the quantitative genetic model of bipolar spectrum illness (Evans et al. These findings support the Darwinian concept of continuity of species that “the difference in the mind between man and the higher animals. Non-human personalities If subcortical primary-processes are foundational for the FFM. and mania. As an example of ANPS research.13 . The C-allele is more common in Sub-Saharan Africa but rather infrequent in European populations. lower for individuals with recurrent major depression or a single depressive episode.. humans may not always differentiate well between the distressful feelings associated with FEAR. Gosling and John (1999) found consistent cross-species evidence for the FFM in mammals ranging from rats to chimpanzees.00 −. Expansion of the FFM These ANPS results also revealed FFM limitations such as FFM Emotional Stability associating with all three of the ANPS negative affects. it is our working hypothesis that the subcortical primary-processes neural systems. depression. FEAR. 2011). as well as on the overall super-factor of Negative Emotionality. after controlling for self-rated depression (Beck Depression Inventory (Beck and Steer. great as it is.26* .The Five Factor Model was measured using Goldberg (1992) adjectives. The genetic analysis of German subjects without psychopathology showed that carriers of the T-allele had significantly higher ANPS ANGER scores.24 −. However.00 . Savitz et al. SADNESS.4. 104) 3.2. 1990). (2008b) also reported that Beck Depression Inventory scores were significantly correlated with higher ANGER scores and lower SEEK and PLAY scores while the Altman Self-Rating Mania Scale significantly correlated with higher ANGER. gender.12 −.09 . empirical step forward in parsing personality.30* −. and PLAY scores. persistent systems is pre-human. 1999) including the capacity to play (Panksepp et al. even more important may be helping to advance personality theory from trait and situationist approaches to interactionist and ontogenetic personality models (see Ballantyne. 1998) suggests an enhanced neuroscientifically premised personality research model. from pain to joy. such longitudinal effects of primary-process experiences can be tested with animal models that incorporate variables and controls that would not be possible with human subjects. Pleasurable experiences result in finding similar situations attractive in the future with traumatic or painful experiences having the opposite effect. This active selection of environments becomes relatively more important as individuals mature and are more able to choose the circumstances and experiences they find attractive. This primary-process affective consciousness may include various sensory/perceptual feelings but especially includes ancient emotional/motivational experiences all mammals share that provide the basis of a “primordial evaluative system” (Kahneman.L. affective neuroscience helps five-factor theory escape circular reasoning by suggesting brain mechanisms that can be manipulated to influence the expression of personality.1. much more ancient than “neomammalian” cortex. there is also an evoked effect. However. Focusing on the PLAY system. which reflect imbalances in these intrinsic value systems of the BrainMind. Early experience epigenetic models have already been tested in rats. and perhaps some other cortically well-endowed animals. Scarr and McCartney. 1999. this homologous foundation consists of instinctual emotion action systems shared by all mammals. Gosling. the evoked effect continues throughout an individual’s life.. Furthermore. It is our position that in these evolutionally older parts of the brain. as well as behavioral reactive guidance. Could the fundamental nature of personality be affective? A key question from a purely cognitive perspective is how non-humans with their more limited cerebral cortical capacities compared to humans can have distinct personalities at all. channel functions The influence of these basic emotion systems is pervasive. p. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 Reuter et al. there is also an active effect in which the individual selects from different environments. individuals experience more passive environmental effects through the environment provided by their parents. one could also test Scarr and McCartney’s (1983) theory that the personalities of fraternal and adoptive siblings would become less similar as they mature whereas genetically 1953 identical siblings (whether adopted or not) would converge over time. they were built into the BrainMind as ancestral “memories”. 2000). thoughts. At the primary emotion level of consciousness.. This primary level makes itself felt throughout our lives but is soon augmented by secondary consciousness reflecting basic conditioning-learning. Shewmon et al. Davis. superior maternal nurturing positively contributed to the stress tolerance of offspring (Francis et al. and numeric creativity tests. However. State vs.. In shorterlived species. since infant individual differences can influence the kind of care infants receive from their parents. (2005) also demonstrated a link between the ANPS SEEK scale and creativity.. 2011 as well as other contributions to understanding psychic pain in that book edited by MacDonald and Jensen-Campbell) would find such distressing situations more aversive – perhaps at lower thresholds – and exhibit more avoidant behavior after those experiences than individuals who were less sensitive to social pain. 1981. This genotype-environment interaction model can be tested with an affective neuroscience approach to personality. By providing a putative physiological basis for FFM traits. 1999). 701). These “ancient tools for living” engender a within-brain type of emotional affective valence.K. 4. but the evolutionary origin and foundational power of these discrete. The mechanism mediating these old brain influences may lie in the distinction between “channel” and “state” brain functions (Mesulam. 4. One example prediction would be that individuals who were more sensitive to social pain (for a full discussion of this construct. 4. These influences determine the “state” of information processing rather than the content being transmitted along the point-to-point “channels. Since personality traits can also be assessed in animal subjects (Gosling and John. awareness of awareness. 2007. it is the primaryprocess level of affective consciousness that encodes biological values through a diversity of raw positively and negatively valenced emotional experiences. affective neuroscience (Panksepp. conditioned place preferences or aversions provide a mechanism for defining the attractiveness of environments. However. each cortical area receives inputs arising from limbic neurons. 2003). The theory predicts that earlier in life. that may be foundational for personality as well as disorders of personality. let alone personalities defined by the well accepted FFM dimensions (see Gosling and John. While the passive effect is hypothesized to decrease with ontogenetic development. 1995 for a more detailed discussion of personality theory levels). Thus. One ontogenetic model is Scarr and McCartney’s (1983) theory of genotype-environment effects hypothesizing that individuals make their own environments. verbal. Affective neuroscience trait versus theoretical expansion In addition to suggesting a reprioritized selection of personality traits in the FFM. However. 2009 and in this issue). subjects with high SEEK scores had higher numeric creativity scores and were significantly superior on figural and verbal creativity.2. thoughts about how external events relate to internal events yield even higher tertiary forms of consciousness consisting of thoughts about thoughts. At the secondary process level characterized by simple conditioning and learning. and general physical adaptation to survival challenges faced by our ancestors for millions of years. 1994).3. and the linguistic/symbolic transformation of experience at which humans excel (for overview. Our theory is related to the idea that the affective foundations of personality lie in MacLean’s (1990) sub-neocortical “limbic” and “reptilian” areas of the central nervous system. SEEK scores also explained more than 15% of the variance of total creativity. and memories. 1999). Using figural. For “state” modulation. J. affecting not only our actions and reactions but also our perceptions. Centering on the CARE system. 2003. 2008). Using intelligence tests as a covariate indicated that the relationship of SEEK to creativity was not moderated by intelligence. These “primary” systems are undoubtedly elaborated during human development by “secondary” conditioning and “tertiary” thoughts and self-reflections. 4. it is clear that young animals and humans without a neocortex are still conscious creatures with primitive affective and perceptual capacities (Merker. extra provisioning of roughand-tumble play decreased ADHD-type impulsiveness (Panksepp et al. one finds the most important evolutionary “roots” of personality. see Panksepp. In humans.. see Vandekerckhove and Panksepp. 1983). which can modulate activity in the entire cerebral cortex. which accounts for adopted siblings not resembling each other’s personalities by late adolescence (Scarr et al. Primary affective consciousness Another affective neuroscience concept related to personality is the multi-tiered framing of consciousness.” These modulatory “corticopetal” projections to the cortex are not . 5. Furthermore..3. 1105). during the final scan subjects were socially excluded when the other players (actually a computer program) stopped tossing the ball to the subject.3. Davis. a fixation rest condition showed decreased amygdala activation to neutral relative to fixation stimuli. differences in subcortical “states” biased perceptions and generated greater reactive tendencies. and my have distinct higher brain regions of influence where raw affects are blended with cognitive information-processing types of strategies. Canli and Lesch proposed a “tonic activation” model that suggested short variant 5HTT carriers exhibit elevated amygdala activity at rest compared to noncarriers. 2002). Individuals with different levels of responsiveness in these primary brain systems not only react differently to the same stimuli. while the FEAR and ANGER systems are closely related anatomically in lower brain regions. Furthermore. . neutral pictures or matching fearful and angry faces vs. FEAR. Coronarybypass patients with more social support reported less pain and took less pain medication (Kulik and Mahler. since affective neuroscience would predict that. . 1989). Evidence Evidence supporting such “state” modulation comes from many sources. (1991) report that patients with generalized anxiety disorder exhibited stronger temporal and frontal cortex PET responses to a passive viewing task compared to normal controls. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 balanced by corresponding “corticofugal” connections from the cerebral cortex to the subcortex.L. 1998). Subjects showed significantly more dACC activation during the social exclusion condition compared to inclusion.. 2010). In addition. college students who were told they had performed poorly on important exams also reported higher pain ratings to a cold pressor task (Levine et al. they remain distinctly separate systems. Women with continuous social support reported less labor pain (Kennell et al. In other words. 1986. subjects were linked in with “virtual” participants supposedly in other scanners and “included” in the virtual ball-tossing game. (2003) did an fMRI analysis of brain changes as she manipulated “painful” social exclusion experienced by participants supposedly playing a virtual ball-tossing game with other participants. Eisenberger and Lieberman (2005) have also linked social and physical pain arguing that increasing or decreasing social harm will correspond to increased or decreased sensitivity to physical pain. mixing fearful and angry face stimuli makes it difficult to integrate results into primary brain emotion systems.” (p. Evidence summary The above studies link primary emotions to limbic brain activity. and SEEKING are foundational for personality expression as well as the emergence of mental anguish and pathology. psychometric approaches to psychopathology (Livesley. and many other brain functions such as cortical processing (see Panksepp. Analysis of emotionally negative or neutral stimuli vs. they will experience these stimuli differently and develop different conditioned response tendencies and ongoing personal preferences. CARING. 1993. Higher baseline amygdala activity levels of short variant 5HTT carriers suggested these short variant carriers may experience a different more threatening world than their noncarrier counterparts. 4. ostensibly because “technical difficulties” prevented them from participating. it is thought to be elicited by social rejection by peers and general social isolation. Conversely. van den Hout et al. Eisenberger et al. Social rejection and the SADNESS system Similar fMRI research has focused on the SADNESS system and the relationship of dorsal anterior cingulate cortex (dACC) activity in response to social pain. individuals whose brain systems have higher medial cortical “resting-state” levels of activity (see Northoff et al. Clinical assessments from affective neuroscience trajectories The FFM is a factor analytic approach to personality. 1987.. (2006) found that short variant 5HTT carriers showed increased rumination in response to life stress compared to noncarriers. the general brain neurotransmitter serotonin is not thought to play a specific role in modulating any of the primary emotional responses but functions as a general modulator of all of the primary affective systems.. social support and social activities were associated with less cancer pain (Zaza and Baine. and SADNESS) along with the positive affect systems of PLAY. [and] alter the tone. these higher scoring BDI subjects also showed greater amygdala reactivity when viewing fearful or angry faces. For extensive discussions of the social-pain construct. Canli et al. J. Brown et al. During an initial control scan.2. Canli and Lesch (2007) have analyzed the influence of the serotonin transporter gene (5-HTT) short variant on fMRI measured activity of amygdala and related brain areas and found that welladjusted and phobic-prone short variant carriers showed greater amygdala activity during tasks such as passive viewing of negative vs. which they supported with a perfusion imaging study that measured absolute amygdala blood flow at rest. Also. The SADNESS system during infancy is characterized by distress calls in response to separation from parents and group members. coloring. For example.. personality theorists and practitioners must distinguish between distress reflecting social harm avoidance and physical harm avoidance likely linked with the SADNESS or FEAR systems respectively. when compared to subjects with lower BDI scores. 4. Measures of self-reported distress also correlated positively with dACC activity supporting the relationship between dACC activity and felt emotional distress during the social exclusion condition. While studies like Canli and Lesch demonstrate the capacity of limbic structures to modulate cognitive interpretations of neutral or negative stimuli.3. see MacDonald and Jensen-Campbell (2011). individuals screened for lack of a mental health treatment history but with higher Beck Depression Inventory (BDI) scores exhibited heightened resting fMRI amygdala activity to neutral visual stimuli (Way et al. 4. right ventral prefrontal cortex (RVPFC) was also more active during social exclusion compared to inclusion conditions but negatively associated with self-reported emotional distress suggesting that the RVPFC may play a role in regulating the dACC. 1990) have demonstrated that personality disorder (PD) . (2003) found that undergraduates reported less pain performing the cold pressor task in the presence of a non-evaluative supportive friend or stranger than when alone or just with an “interactive” control participant. 2011) are more reactive to neutral stimuli and respond differently to events. It is our position that individual differences in such higher affective as well as lower primary-process aversive affective brain systems (RAGE. 2000). and interpretation of experience rather than its content” (Mesulam. However. In the second scan. 2000. In medical settings. pp. which was “interpreted as indicating elevated amygdala activation during the processing of emotionally undefined stimuli (our italics) in short variant carriers. which has been derived from studying “normal” populations. 1991). To create a sense of social exclusion. Also. In older individuals. Given these findings.1. Such tendencies would likely result in observed individual differences in the many behavioral dimensions of individual lives. 78–79). Furthermore. This asymmetry allows the limbic system to “rapidly shift information processing states throughout the cerebral cortex . geometric shapes.3. For example. subjects just watched other participants play the virtual ball-tossing game.1954 K. and we just touch on a few: Wu et al. Clark. These two themes have also been shown to be important for the success of cognitive behavior therapy (CBT).K. Hostility. ANGER. getting them to move around and physically interact with the therapists may help promote a shift in affective balance. a challenge for the FFM may be identifying facets that are congruent with neuroscience research rather than relying on an arbitrary number of rationally defined facets. just like other mammalian young. could emerge as newly defined brain emotions in animals that can be used to guide higher-order affective personality scale construction.L. If a person is frozen in an emotional state. and having separate dimensions for the ANGER and CARE systems might contribute . Consequently. 1996. Panksepp (2009) has emphasized the critical nature of the CARE system for effective therapy and has even considered how oxytocin administration might be used to enhance a supportive nurturing therapist demeanor.. 2008). 2011). Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 diagnosis can be conceptualized in terms of the FFM (Clark and Livesley. and that those more skilled in mindfulness enhanced the effect (Creswell et al. 2007b. For example “empathy” (Chen et al. 2009). 499). In our estimation. affective neuroscience helps reconnect personality theory back to our evolutionary roots by showing how closely our personalities are linked to the ancestral affective forces we share with other mammals and by relating psychopathology to disturbances in the primary emotional subcortical brain systems. 2010. 2002. Affective neuroscience and therapeutic effectiveness Affective Neuroscience also has important implications regarding therapy for affective imbalances (see Coenen et al. tertiary-process awareness level may be therapeutically beneficial. whose emotionality is closely linked to motor action systems.. and creativity of real “rough-and-tumble” PLAY.g. Shedler (2010) has summarized data on the long-term effectiveness of psychodynamic therapy and likewise emphasized the importance of a nurturing “working alliance” between the patient and the therapist. it is most important to be very clear about the primary-process issues that undergird personality development. In a second point. Play urges in adults may be reenergized by physical activities like sports or dance accompanied by music that stimulates the rhythmic motor impulses of the body (Panksepp and Trevarthen. recognizing that our emotional instincts are fundamentally action systems suggests the importance of incorporating bodily activity into therapy. The NEO PI-Revised (Costa and McCrae. which suggests that the FFM does represent a meaningful continuum of behavior extending into the realm of clinical pathology. Widiger and Costa (2002) reviewed 56 studies that related the FFM to PD symptomology. In a review of evidence-based explanations for psychotherapeutic interventions. 1997. 1996) that linked poor therapeutic outcomes to rigid insensitive implementation of therapies. this can result in receiving psychostimulant drugs.. Of these studies. Dyce and O’Connor. One advantage of distinct emotional dimensions is their underlying unipolar nature. 1998).” In one sense. There is good reason to believe that one of the main functions of the PLAY system is the epigenetic construction of the social brain (Panksepp... p. While adult brains are not as plastic as those of children. primary-process emotional systems of the brain suggests that the FFM itself needs further clarification. it does not seem to be the cognitions as originally proposed. which dealt with CBT’s emphasis on cognitive distortions. a finding that was replicated by Hayes et al. bringing the body and emotional actions on to the therapeutic field.. and SADNESS into a general Neuroticism factor detracts from their unique contributions to personality expression and mental disorder. 80).” (p. effective therapeutic interventions may typically impact the affective dimensions and levels of the BrainMind more than cognitive. lies in the sensitive exploration of troubling feelings and reactive patterns and perhaps not as much in changing distorted cognitions. 109) and in which it may be more productive to use functional neuroscience rather than factor analysis to inform and identify new trait dimensions. playfulness can be a valuable resource for redirecting adults onto a more adaptive life track. Thus. 2007). Play deprived human children. 2003). 6.” An exploration of cross-species. A third theme. ANGER and CARING need to remain distinct dimensions rather than fusing them onto opposite poles of a scientifically confusing Agreeableness factor. Shedler also emphasized the importance of “labeled experience” structured so the client “gains awareness of previously implicit feelings and meaning” (Shedler. In adult psychotherapy. p. which has been previously associated with emotional regulation (Lieberman et al. spontaneity. We may be at a point “in which software capabilities now exceed both the quality of data and the scope of conceptualization” (Wiggins and Pincus. develop heightened motivations to play. 2007). This suggests that a better understanding of primary-process affective feelings at a thoughtful. In fact the therapeutic aim of cognitive change predicted poorer therapeutic outcomes. Furthermore. 7. some of which may be best defined as factor “blends. 104. Effective treatment of behavioral disorders. 1955 In addition to the importance of the CARE system being displayed by therapists. was not supported in these studies. For many people. information-processing ones. J. Activating the PLAY system requires the bodily vigor. 1992) includes six facets for each of the FFM dimensions. 2006. p. 2010). 2009). In humans. Some feel that facet level analyses are required for optimal discrimination between PDs (Axelrod et al. He also discussed research (see Castonguay et al. Davis. Robust physical activity by itself may be as effective an antidepressant as the medications that dampen emotionality (see Watt and Panksepp... quoted from Castonguay et al. Lieberman showed that affect labeling decreased amygdala response that was inversely correlated with the right ventrolateral prefrontal cortex activity. which may be most easily applicable to children. Ample physical play time for children may be one of the best ways to protect them against ADHD as well as depression.” e. After all we are dynamically ‘embodied’ creatures. Summary of affective neuroscience perspectives on the foundations of personality Our scientific understanding of personality began with Darwin and McDougal with their focus on emotion and instinct and the recognition that the differences between humans and their nonhuman ancestors were of “degree” and not of “kind. However. The authors concluded that “much of this research indicates strong support for understanding PD symptomatology as maladaptive variants of the personality traits included with the FFM” (Widiger and Costa. 30 used clinical populations. 2009) and “jealousy” (Panksepp. p.. In support of this approach. which are effective inhibitors of physical play urges. 2002. Combining the key affective dimensions of FEAR. (1996). De Fruyt et al. 2002). the PLAY system may have considerable untapped potential for helping patients reintegrate troublesome emotional experiences towards more adaptive and emotionally comfortable affective trajectories. and the other socially constructed emotions. Kazdin (2007) concluded “whatever may be the basis of changes with CBT. interactive though they are with other emotions. the PLAY system could have powerful and underutilized therapeutic effects as well (Panksepp. may open up many opportunities for change that would not exist as long as therapy consists largely of “talking heads. and long-term therapeutic change. 2009). Animal models confirm that ADHD impulsivity can be reduced with rough-and-tumble play during early development (Panksepp et al. 8). J.E. if the person responded with “Disagree”.A. FEAR. W. Perhaps linking human personality to our ancestral emotional urges will bring further clarity to the understanding of human personality. K. Ball. L. Castonguay. Spielberger. Psych. Psychol.A. Soc. Trull.L. Sheffield. P. Neural correlates of epigenesis. 64.R.A. The Altman Self-Rating Mania Scale. S.. 2009. Cons. P. Note: Parkinson’s disease has been linked with distinctive personality differences that include an increased risk for depression. 171. Psychiatry 42. Ph. Amin. 2007. Assessment 11 (4). 1993. 17. 2003. Spirituality (with only 12 items). American Psychological Association.. A unified biosocial theory of personality and its role in the development of anxiety states. behavioral activation.. 2. 13. Psychol..B. Neurosci. Brown. In: Costa. It would have been interesting to have supplemented the Neuroticism measure with measures of SADNESS. 57. M. and ventral striatum–all very ancient brain areas.. 1936. R. E.R. it may be helpful to have more precise tools to systematically assess the location of experimental subjects and therapeutic clients in affective space. 1997. Confirmation and clarification of primary personality factors. 497–504. Damholdt et al. Pers.4 (and compare scores to the original norms published in the 2003 paper) is to use a 3 for the highest responses and a 0 for the lowest responses. A. S. Using this 4-point scale with 1 for “Strongly Agree” and 4 for “Strongly Disagree” (but computing scores equivalent to hand scoring above with a base score of 0) the following formulas were used for “computer scoring” the 6 primary ANPS scales plus Spirituality: SEEK score = (+21 −ans1 −ans17 −ans33 −ans49 −ans65 −ans81 −ans97 +ans9 +ans25 +ans41 +ans57 +ans73 +ans89 +ans105). Nat. doi:10. 276–283. 948–955. Jaak Panksepp. Med. Copyright © 2004. Ph.. Davis.4 was originally set up for scoring with a “scanner” that by default numbered the bubbles from left to right 1... pp. each of the negatively worded SEEK scale items in the even blocks (numbers 9. 151–157.” Thus. In: Lubek. and 105) would be reverse scored as follows: Strongly Agree = 0. Nat.. K. References Allport. H.. ANGER score = (+21 −ans4 −ans20 −ans36 −ans52 −ans68 −ans84 −ans100 +ans12 +ans28 +ans44 +ans60 +ans76 +ans92 +ans108). L. Clark. Robinson. 243–266. SADNESS score = (+21 −ans6 −ans22 −ans38 −ans54 −ans70 −ans86 −ans102 +ans14 +ans30 +ans46 +ans62 +ans78 +ans94 +ans110). T. Psych.4. C. Social support and experimental pain. T.A. (2011) collected FFM data in a study of Parkinson’s disease and found that Parkinson’s patients with depression displayed lower Extraversion and higher Neuroticism scores. C. Spirituality score = (+18 −ans7 −ans23 −ans39 −ans55 −ans71 −ans87 +ans15 +ans31 +ans47 +ans63 +ans79 +ans95).. J. 47 (1). 67 (2). Acad. 3. M.. and affective responses to impending reward and punishment: the BIS/BAS scales.). second ed. Carver. 103 (43). A contrarian view of the five-factor approach to personality description. White. from the Hope for Depression Research Foundation. All rights reserved. Kranzler. J. Different contributions of the human amygdala and ventromedial prefrontal cortex to decision-making. Davis.1956 K. 49. Hayes. Inc. Two approaches to identifying the dimensions of personality disorder: convergence on the five-factor model. Ballantyne... ANGER. Kenneth L. Sci. Chen. . a psycho-lexical study.L. vol... 514–524. Recent Trends in Theoretical Psychology. Above all. Correspondingly. Congdon. Damasio.L. vol.. FEAR.. Advances in Personality Assessment. C. and 4.E. 1999. Proc. 10 (9). Hedecker. 1103–1109. 1996. Clin.. Canli.. Pers. H. and an “Agree” response was given a value of 2. D. Personality Disorders and the Five-Factor Model of Personality. 2009..).. 65.P. 5473–5481. 2002. This work was supported by a grant to J. P. Appendix A. 33. Davis.R. Bechara. Lesch. the ANPS 2. 25. Lee.. FEAR score = (+21 −ans2 −ans18 −ans34 −ans50 −ans66 −ans82 −ans98 +ans10 +ans26 +ans42 +ans58 +ans74 +ans90 +ans106). Steer. Odbert. . USA. 65. 81. 6 were designed as “social desirability” or “unlikely virtue” items.. Disagree = 1. G.B.J.. R. M. 1994. Block.. Predicting the effect of cognitive therapy for depression: a study of unique and common factors.P.T. M.R. Manual for the Beck Depression Inventory. J. 1990. A. 8. Constable. J. 1999..L. Livesley. Factor replicability and validity of the temperament and character inventory in substance-dependent patients. U. L.C. Remy et al. For example. T. each of the positively worded SEEK scale items (numbers 1. and Strongly Disagree = 0. the “hand scoring” and “computer scoring” procedures are identical.D.R.P. In: Butcher.T. SADNESS. D.. followed by a filler research question. Empathy is moderated by genetic background in mice. and a “Strongly Disagree” response was given a value of 4. and 3 were written to measure “social anxiety” (+ans24 +ans104 +ans112). J.J.L. Wiser.. J. 1995. thalamus. S. Leary... Acknowledgements We appreciate all the investigators who have taken an interest in the Affective Neuroscience Personality Scales. Psychiatric Dev. Alternatively.. amygdala. Springer.. e4387. E. Agree = 2. Maolin. Pegasus International. Agree. Bull. Widiger. 2006...G. E. San Antonio.4: 7 were designed as “Dominance” items (+ans8 +ans40 −ans56 −ans72 +ans88 +ans103 −ans111).. T. 1947.” the scanner gave them value of 1.. They hypothesized that depression and anxiety in Parkinson’s patients were associated with loss of dopamine and noradrenaline activity in the limbic system. pp.. Psych.. (Eds. (Ed.L. Omura.S. Trait-names. pp. Beck. 197–220. New York... Cloninger. Lee. DC. Corbitt.. Cattell. Washington.A. Pers. The columns were correspondingly labeled “Strongly Agree. p16033–16038. Psychometrika 12 (3). Herrmann. H. Z. CARE score = (+21 −ans3 −ans19 −ans35 −ans51 −ans67 −ans83 −ans99 +ans11 +ans27 +ans43 +ans59 +ans75 +ans91 +ans107). Biol.D.. 117 (2).. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 to clearer primary-process personality thinking. Goldfried. and Strongly Disagree. G. The Psychological Corporation. K. T.S.F.. Raue. Canli. Agree = 1. There are 16 filler items in the ANPS 2. 319–333. And that was our original motivation for developing the ANPS. Goldberg. We hope it will be useful for many other research and practical endeavors. 4.W.D.. M..P. Peterson. R. Psychosom.M. Disagree. Q. J. J.S. B. Tennen. Disagree = 2. Algebraically. 187–215. 161–176. 1995.A. Behavioral inhibition. PLAY.. A. Widiger.). copyright 2004 version The ANPS items are arranged in 14 blocks using the following item sequence: SEEK. Ashton. This procedure allows for the possibility of a low score of zero on each scale.... The easiest way to “hand score” the ANPS 2.pone.1371/journal. Panksepp. Q. I. T. From initial abstractions to a concrete concept of personality. PLAY score = (+21 −ans5 −ans21 −ans37 −ans53 −ans69 −ans85 −ans101+ans13 +ans29 +ans45 +ans61 +ans77 +ans93 +ans109). L. Psychol. CARE. 89. PlosOne 4 (2). 79–91. Soc. Monogr. (2005) used PET analysis to investigate the brain correlates of depression in Parkinson’s patients and matched controls and found that depressed Parkinson’s subjects had lower activity in the locus coeruleus. and 97) would be scored as follows: Strongly Agree = 3. 1986. In effect.. the items are being scored on a scale running from 0 to 3 rather than 1 to 4. 19.. Neurosci.W.. J. 167–226. 297–313.. Higher order factors of personality: do they exist? Rev. Assessment 69 (2). Relations of Five-Factor Model antagonism facets with personality disorder symptomatology. G. The items in the even blocks are reverse scored. Lahvis. if a person responded to an item with “Strongly Agree. 3. Lesch.R. Towards a consensual set of symptom clusters for assessment of personality disorder.. Damasio. anterior cingulate cortex. 73. 41.M. R. 1997.P.. which can cautiously be used as an indication of deceptive responding (−ans16 +ans32 −ans48 +ans64 −ans80 +ans96). and Strongly Disagree = 3. Long story short: the serotonin transporter in emotion regulation and social cognition..M.T. Haas. Scoring the ANPS 2. J. (Eds. the scanner gave them a value of 3. Axelrod. Clark. and ANGER to assess likely links to discrete brain systems. H.A. Likewise.R. de Vries. Altman.. J. K. A.0004 387. 3. Normansell. Psychol.. Farmer. Soc. Lincoln.F. 2005. P. 2197–2201. The ‘resting-state hypothesis’ of major depressive disorder-A translational subcortical-cortical framework for a system disorder.R. L. Direct.. Cloninger’s psychobiological model of temperament and character and the five-factor model of personality.L. Lieberman.J. J.A.R. Way. USA.M. Personality disorders and the five-factor model: a test of facet-level predictions. 2006. A. Zenasni. Neurosci. 2007. (Eds. 69–75.. Personality in non-human animals. 1999. L. Oxford University Press. Alcaro. 1 (45): Clinical Neuropsychology. R. Livesley. Psychol. N.. Handbook of Clinical Neurology (Revised Series). W.. Heath. N. MacDonald. Center for Psychobiology of Personality. FL.P. 2011.M. de Raad. Res.. Way.. R. pp. 63–81. Normansell. 76 (6). 1990. The Social Outcast: Ostracism. Affective Neuroscience: The Foundations of Human and Animal Emotions.A.. American Psychological Association.. 8 (2). M. Nebraska. Psychiatry 69. Hofstee. and the construction of the social brain: should the first class each day be recess? Am. Curr. K. N.L. pp. 407–467. Behav. Sci. Burgdorf. 1216–1229.. Play 1. Norton & Company. J.D. Panksepp... Brain emotional systems and qualities of mental life. 2010 Dec 28 [Epub ahead of print]. L.R.E. J. Pers. Keck Jr. Van Heeringen. J. Psychologist 58 (9). New York.. Eisenberger..A. O’Connor.. Remick.).I.J. Huber. Department of Psychiatry.. Ostergaard.. 2002. French validation of the Affective Neuroscience Personality Scales (ANPS). J. S. Physiol. The Expression of the Emotions in Man and Animals. UK. Ann. The affective neuroscience personality scales: normative data and implications.. 2007b. Washington. 153–168. Rejection. Panksepp.. MO (available from C. 231–244. Rev.B.I. NY. Panksepp. J. N. Coenen. 1987. A.D.. Disord.. B. P. 2010. Am.. T.. Kirsch. J. Compass 2. B. Social Pain: Neuropsychological and Health Implications of Loss and Exclusion. 985–1001. H. Physiol.. 1998. second ed.F. 1957. 69 (6). Primate Perspectives on Behavior and Cognition. B. C. Goldberg. Psychological Assessment Resources.).. Edinborough..F. P. Gray. Integration of the big five and circumplex approaches to trait structure.T. C.. D. The Functional Organization of the Diencephalons. Physiol. McElroy. Panksepp. J. De Fruyt. Assessment 20 (3). Panksepp. Panksepp.). D.... 2003.D. C. 601–603. Mood changes... 1–27.. J. L.A. Consulting Clin. 77– 107. In: Eysenck.W. 1971. A systematic method for clinical description and classification of personality variants. A neurogenetic approach to implusivity. M.. 2007a.. Merker. L.D.. Great Britain. G.. J.. Widiger. Larsen. Chichester. Research. American Psychological Association. 2003. A systematic approach to the delineation of personality disorders. Handbook of Jealousy: Theory.. Diff. J. 249–266. T. Rev. S. Science 302. Morrison & Gibb. Vol. In: Velmans. L... J. Oxford University Press.. (Eds. Neuroevolutionary sources of laughter and social joy: modeling primal human laughter in laboratory rats. Ind. C. 2000. 2011. 265.. 1992. R. MacDonald. 2000. Wiley-Blackwell. Gen.. Cross-species affective functions of the medial forebrain bundle-Implications for the treatment of affective pain and depression in humans. Arch.. 2008... Psychiatry 144.). Eisenberger. 2007. Neural correlates of dispositional mindfulness during affect labeling... pp. 30. De Clercq. 11–51. S... C. Goldberg. Kennell.M.. 30–80. J. (Ed. J. 1970. Pers. 281–291. 85–126. Bruyn. New York. Therapy 8.J. Panksepp. P... Vol5 (3). Neurosci. 182. Nocjar. Oxford University Press. B. London. 1998. Introduction: Personality Disorders and the FiveFactor Model of Personality. 57–69. N. 2007. Feinberg. Rev. Psych. 1155–1158.. McCrae. Solomon.M.J. Principles of Behavioral and Cognitive Neurology. J. Am.. The Archaeology of Mind: Neuroevolutionary Origins of Human Emotions.. The Triune Brain in Evolution. Panksepp. P.D. 3-14. West Sussex. Rev. B. The psychophysiological basis of introversion-extraversion. Interaction of 5-HTTLPR and a variation on the oxytocin receptor gene influences negative emotionality. G. and Multidisciplinary Approaches. Soc. G. L. L. Washington.. 246–276. 1872/1967. Oxford University Press. Goldfried. J. F.. .. 50th Nebraska Symposium on Motivation: Motivational Factors in the Etiology of Drug Abuse.R. Aggression elicited by electrical stimulation of the hypothalamus in albino rats. 2004. L. Biobehav. Liu. Pers. pp. Brain Sci. Krass. Panksepp. Berlin. L. 48 (1). W. 1993. Van De Wiele. Dyce.M. 1981.. 2009. R. New York. Gillespie. Pers.). Panksepp. In: Bevins.. 1985. An Introduction to Social Psychology.P.R. W. Psychiatry 44. McGrath. 63 (1). J. Psychiatry 143. 18 (5).J. 772–777. Panksepp.. (Eds. Maedler. 2004.A. R. C.. Biobehav. Louis. Brain Res. 1987.. Cloninger.R.. Psychosom. pp.D. Psychology Press. P. Akiskal. K. W. Kazdin.. W. DC. Psych. Abbott. C. 48–55.I. ADHD. 560–565.E. Turner.. Lieberman. pp. C. J.. Biol. 74. Behav. S. Panksepp. 114–129. New York. 14. MacLean. Perinatal decortication impairs performance on an 8-arm radial maze task. Behav. Social Pain: Neuropsychological and Health Implications of Loss and Exclusion. 1990.. In: Fosha. von Hippel. Livesley. Damholdt. 429–443. 37. Northoff.. Cloninger. Reuter..M. Padawer. 2011. Panksepp. MO 63110).E. 573–588. Eisenberger. Sci. St.T. Borghammer.. J. 271–285.. Dimensions of the Ape Mind: adding personality to behavior and cognition. 6. 146–163. K. J. 2011... Pers. Social support and recovery from surgery.). The genetic and environmental relationship between Cloninger’s dimensions of temperament and character. Francis. 1989. Psychol. Am. An alternative description of personality: the big-five factor structure. O.. H. 31–45. Box 8134. Mahler. C. Grune and Statton. M. DC.. Lieberman..R.D.A. D.. J... W.. W. Eisenberger. The Healing Power of Emotion. Evans. Van Heeringen. Jensen-Campbell. Hayes. L. Nathaniel. New York. Elsevier Science Publishers. 2003. T. 155–163.. J..O.I. A critique of Eysenck’s theory of personality. 300–304.. J..J.. (Eds. F.. Pers. 2003. N. 2008b. Med.R... Panksepp. In: Williams...R. M. M. J. MA. 1996. Personality dimension in non-human animals: a crossspecies review. King. Castonguay. Revue européenne de psychologie appliquée 58. J. 1994. M.J.. Mouchiroud.. M. P. Health Psych. Costa. Kahneman. M. 1999.M. 421–428. Pahlavan. J. McDougal. Brain modules. R. In: Washburn. J.D. and the equally implausible distinction between TCIR “Temperament” and “Character” Scales: reply to Cloninger (2008). Digman. Montag. Oxford. 109–127. 1908/1963.E. Washington.M.. Panksepp. 697–720. J. hospital: a randomized control trial. 290–292. Panksepp. D. Costa. Schneider. Williams. American Psychological Association.H. F. Washington University. 55–60. A. Soc. personality layers. H. Davis. Brain Sci. G.. J..... The Blackwell Companion to Consciousness. The neurochemistry of behavior. The neurobiology of social loss in animals: Some keys to the puzzle of psychic pain in humans. Psych. J. De Fruyt. S. Does rejection hurt? An fMRI study of social exclusion. 2008..M. J. Panksepp. 2008a. D. J. Pers. Schlaepfer. Behav. 1999. Toward a general psychobiological theory of emotions.. 35. C. Creswell. Inc. 221–238. (Eds. Biven. R. Hinkley. Panksepp. Drug-sensitive reward in crayfish: An invertebrate model system for the study of SEEKING. 1991.. 2003. Gosling. D.J. 2005..A. Siviy. Burgdorf. John. Hess. Norton. Klawans.A. 97–105. Behav.T.. K. 2011.A.R. J. In: Hart. Ind.D. (Eds.A. 59 (6)...C. Personality Disorders and the Five-Factor Model of Personality... J. 2011.S. Rev. M.F.M.A. Psychol. R. 2007. C.A. M. Behav. Pers. Mediators and mechanisms of change in psychotherapy research. M. 1990. Tom.. Amsterdam. J.. Consciousness without a cerebral cortex: a challenge for neuroscience and medicine.. Springer-Verlag. 728–732.. Goldberg.. A Psychometric Evaluation of the Revised Temperament and Character Inventory (TCI-R) and the TCI-140. 56. Panksepp..R. L. Nongenomic transmission across generations of maternal behavior and stress responses in the rat. 55–79... Effectiveness of targeting the vulnerability factors of depression in cognitive therapy. Blackwell Publishing. Panksepp.. addiction. F.A. 12 (1).. Putting feelings into words: affect labeling disrupts amygdala activity in response to affective stimuli. J.L. Washington University School of Medicine. spiral structures.K. Med. J. Social Exclusion... Revised NEO Personality Inventory (NEO PI-R) and NEO Five Factor Inventory (NEO-FFI) Professional Manual. Personality structure: emergence of the five-factor model. C. Martin. J. Sadovnick. 85. Oxford. 23. A perspective on judgment and choice: mapping bounded rationality. Lieberman... B. pp. DC. Siegel.. A. Odessa. Robertson. Psych. Pain 54. 101–120. 47–60.). 417–440.).P. H. S. J.. 441–452. Bardo. 2010 Dec 22 [Epub ahead of print]... Congdon. 2010 Dec 21 [Epub ahead of print]. Cox. The validity of cloninger’s psychobiological model versus the five-factor model to predict DSM-IV personality disorders in a heterogeneous psychiatric sample: domain facet and residualized facet descriptions. Fiebach. USA.S. 479–510. Neuropersonality Clin... Assessment 20 (3). Crockett. T.). J. Forgas. J. pp.. J. LLC. N. Kulik. Consciousness Cogn. 41. M. New York. S. DC. Ann. Familiarity of temperament in bipolar disorder: support for a genetic spectrum. Ltd. M. F. Diff. J. Pfeifer. Cloninger.. Neurosci. S. 29. Darwin... St. 2008. 1447–1483. Taylor & Francis Group... 311–316. In: Vinken. Psych. L. A. 52.. 335– 340. Oxford. Science 286. Why it hurts to be left out: the neurocognitive overlap between physical and social pain. 623–627. W. J.. planes of being. Traits and behavioral prototypes of personality disorder. 1992. Canli... Biobehav. S. J. 2005.. 2011. Gosling. Mesulam. and Bullying. Am. Kelsoe. 1–26.F. Van De Wiele. Modeling ADHD-type arousal with unilateral frontal cortex damage in rats and beneficial effects of play therapy. Panksepp.. Assn. K. V.. Panksepp.. Levine. Diorio. J. Wiebking.G.. and withdrawal. A Model for Personality. 1990. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 Cloninger. Clin. T. 2011... Huber. 1986.. Ann. (Ed. Meaney. 64 (3). Plenum Press. Panksepp..J. pp. 1986.. The parkinsonian personality and concomitant depression. Panksepp. Goldberg. The role of emotional systems in addiction: a neuroethological perspective. Behav.. B.. L. pp. M. E. Klaus. Brain Cogn. Feeling Good: The Science of Well-Being. Louis. S. Psych. P. 2007. Gray. Malden. The Temperament and Character Inventory—Revised.. B. C. Psych... Deyo. (Eds. J. 1957 reward. Play. Washington.R. Affective Disord.J. Affective consciousness: core emotional feelings in animals and humans. Neuropsychoanalysis 5 (1). American Psychological Association.. Effects of neonatal decortication on the social play of juvenile rats. J.B. Failure hurts: the effects of stress due to difficult tasks and failure feedback on pain report.A..L. Continuous emotional support during labor in U. Affective consciousness..W. 2008.. 1931–1946. Cloninger. M. Legerstee. NY. J. In: Jensen-Campbell. The evolutionary sources of jealousy: cross-species approaches to fundamental issues. Rev. J. 8 (3). 1982. Farmer. J. Neurosci. Gordon. J. Davis. Cloninger.. Creswell. A.. M. 424–435. Weinberg. In: Malloch. C. D. Scarr. Wu. J.R. T. M.. 2008a.C. The flow of anoetic to noetic and autonoetic consciousness: a vision of unknowing (anoetic) and knowing (noetic) consciousness in the remembrance of things past and imagined futures. Eur. J. van den Hout. Vlaeyen. J. Child Neurol. Washington. DC. 19. 2000.. S. Hamann.. Serotonergic involvement in the psychosocial dimension of personality. L. Lees. Byrne. Panksepp. M.. 12–24. 15 (3). Personality Disorders and the FiveFactor Model of Personality. USA. Hazlett. Peirson. M. Wittig. Owen. Ross. N.. caractère et tempérament: La structure translinguistique des traits.T. Neuropsychoanalysis 11. N. character and temperament: The cross-language structure of traits. Genes Brain Behav. 1063–1070.. (Eds. Cogn. 2010. 2009.L. J.. Dysthymic and anxiety-related personality traits in bipolar spectrum illness. In: Costa. pp. Psych. 179–183... 59–87. P.. 7 (8). R. Emotion 10 (1). L.. Psychologie Franc¸aise 51 (3). van der Merwe. Scarr. M.. peptidergic. USA.. Davis. Depression in Parkinson’s disease: loss of dopamine and noradrenaline innervation in the limbic system. Turjanski.. J. 29–37.. Does failure hurt? The effects of failure feedback on pain report..H. 869–876. Behav. J. 2010. J. Consciousness in congenitally decorticate children: developmental vegetative state as self-fulfilling prophecy.G. Personality resemblance among adolescents and their parents in biologically related and adoptive families. Baine. 22. Zaza. A. Neurosci. S. Tellegen.T. Widiger.. (Eds. The neuroscience of emotion in music. L. 1181–1199. The efficacy of psychodynamic psychotherapy. Personality and biological markers of creativity..D. B. 1991. J. 2006.). C. J. J. Depression: An evolutionarily conserved mechanism to terminate separation-distress?: a review of aminergic. T.R.W. Trevarthen.. J. Reconstructing the evolution of laughter in great apes and humans. a theory of genotype. 335–346. M. Schnabel.. Biol.. Montag. 195–198. Goldberg. Pincus.. Fiebach. Eur. pp. J. C. M. 2009. . 1981. S.... Pain 4. J. C.J. Widiger. PET in generalized anxiety disorder.). Psychiatry Res. J. Reuter.. Pers. 89. R.1958 K. P. second ed.A. Widiger. 83–95. How people make their own environments. 98–109. van der Merwe. M. P. 1988.. 41. A.. M.. N. cyclothymic and hostile personality traits in bipolar spectrum illness: a family-based study. pain tolerance and pain avoidance.. . Engelhard.. Pers. 2008b.. R. K. In: Costa. D. 2005. Watson. Zimmermann. Affective Disord. Wiggins. Am.. T.. N.. J. Holmes. Saucier. R.. 1999.. 2010..D. J.J. 1983. Five-factor model personality disorder research. I.. Psychiatry 29. G. Remy... 54. Weber. Neuroimaging support for discrete neural correlates of basic emotions: a voxel-based meta-analysis. J. 1314–1322. Eisenberger. J. 2009. 42.. L.. . A. Buchsbaum. Webber..F. Brain Res. 885–898. Dev. Development and validation of brief measures of positive and negative affect: the PANAS scales. S. DC. Hypomanic.A.L.A. Hershey. Shedler. Personnalité. J. 265–284.. J. Consciousness Cogn. 202. Pain Symptom Manage. Washington. Panksepp. Berk.. L... 2001. Kellerhoff. E. 1–22. D. Curr. L. 1018–1028. Soc. 1106–1111.. G.. Lieberman... Ramesar. pp. J. Way. Plein. American Psychological Association. McCartney. American Psychological Association. Savitz. second ed.L.. Bond..S. Shewmon. T. Personality endophenotypes for bipolar affective disorder: a family-based genetic association analysis. Johnson.A. 18. D. Savitz..S... J. 2009. Psychiatric Res. 2002.. 2008c.C. 920–929. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 Panksepp. 2005.. P. Child Dev. Biol. environment effects..L.I.. 19. 40 (5).. Watt.. Doder. 24 (5)..D.. A.. and neural network perspectives.. 1999. Elger. Oxford University Press. 2002. Psychologist 65 (2).M. Trevarthen..T. Relationship between serotonin and the Temperament and Character Inventory. Ramesar. Ramesar.S.). 5–104.A. Savitz. Vytal. Pers. (Eds. Panksepp. N. UK. P. C.. J. Psych. Cancer pain and psychosocial factors: a critical review of the literature. S.A. Vandekerckhove. Oxford. Clark. Reuter.C.. Heuchert. Personality Disorders and the Five-Factor Model of Personality. C. 526–542. 109. Thomala.. 364–374.A../Personality. J.. M. Sicotte.. 103–124. 2009. Costa. Communicative Musicality. Brooks. C.W. Peters.. Soc. 105–146. van den Hout. H. P. Dispositional mindfulness and depressive symptomalogy: correlations with limbic and selfreferential neural activity during rest.. M. N.. Brain 128... Personality structure and the structure of personality disorders.R. J. P.T.J. Hennig. Pharmacol. The biological basis of anger: associations with the gene coding for DARPP-32 (PPP1R1B) and with amygdala volume. 54.. Kempel.. J. E. K.L. van der Merwe. M. 305–311. Wai. 2002. B.A.A.M.