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Authors requiring further information regarding Elsevier’s archiving and manuscript policies are encouraged to visit: http://www.elsevier.com/copyright experienced emotional stimuli. time of day effects have been observed for the Circadian timing: time is measured by biological oscillations or timing of auditory and visual signals in the seconds-to-minutes range cycles that can range from milliseconds to years.g. how our sense of time is altered Interval timing: usually defined at the discrimination of durations in the seconds-to-minutes range. (ii) expressive behaviors and (iii) conscious experi. indi- logical arousal. In this sense.2]. Recent studies using refers to whether the stimulus is positive or negative. Although many studies interval procedure. conds-to-hours range.droit-volet@univ-bpclermont. aware ahead of time that the duration of an event will be relevant to are regulated by a master clock in the suprachiasmatic nucleus. directly to the amygdala.g. tive timing and includes the study of a specialized timing module or cally produce ‘perceptual. sports. mechanisms expressed in a variety of tissues.16]. modality.2.e. five main types of cognitive and affective factors have are defined by oscillations that last approximately a day (i. (decrease in clock speed) relative to control conditions.com 1364-6613/$ – see front matter ! 2007 Elsevier Ltd. hours) ranges. [54].20]. guiding behavior. CNRS (UMR 6024). Université Blaise Pascal. NC 27708. while maintaining the same PSE or peak time values [17. examples for future directions. are able to occur when the functions are flattened and shifted either up or down measure the passage of time accurately in the millise. In contrast. durations and the standard deviation of duration judgments. duration dis- eyes. www. Circadian rhythms [1. by way of the crimination is relative rather than absolute. 34 Avenue Carnot.16]. One proposal is that range. doi:10. film. Duke University. The study of time nition. Author's personal copy Opinion TRENDS in Cognitive Sciences Vol. (Sylvie. 63037. Circadian rhythms are Prospective timing: refers to situations in which the participant is a fundamental property of all higher forms of life and. Although the suprachiasmatic nucleus appears unnecessary for interval timing.g. interval timing that describes the linear relationship between target Emotion: a response of the whole organism. Functional standardized emotional material provide a unique neuroimaging and lesion-based experiments have demonstrated the crucial role of the amygdala in recognition of certain emotions opportunity for understanding the neurocognitive signaled by sensory stimuli (notably fear and anger in facial mechanisms that underlie the effects of emotion on expressions or cues predicting electric shock) that might be inde- timing and time perception in the milliseconds-to-hours pendent of the intensity of the stimulus [7. vertical shifts in timing functions have shown that humans.tics.26]. We outline how these new findings can be fear and/or anger biases attention towards threat-related stimuli by explained within the framework of internal-clock models enhancing the output from an amygdala-centered ‘pre-attentive threat evaluation mechanism’ [48]. Horizontal shifts: in timing functions occur when the entire sigmoi- dal-shaped (bisection procedure) or Gaussian-shaped (peak-interval Introduction procedure) function relating the probability of a response to signal Time and emotion are inextricably linked by the rhythm duration is shifted to the left (increase in clock speed) or to the right and tempo of a myriad of external and internal events. internal clock [1. Corresponding author: Droit-Volet. stop/pause and reset on command [1. affective valence and linguistic factors [8–15. Most Embodied emotions: refer to the fact that perceiving another research conducted on timing and time perception involves prospec- person’s emotional expressions and thinking about emotion automati. somatovisceral. a snake) produces and emotion is at a crossroads. sequence and durations of events based upon memory. to the internalization of the Earth’s light–dark cycle by molecular arousal. ears and nose bypass the cortex and go.sciencedirect.2]. In recent theories of embodied cog- tional time sharing and clock speed. involving (i) physio. cating that variability in timing behavior grows proportional to the ence.09. Interval timing is less precise than circadian timing. 1002). dance. To date. in mammals. 24 h) due been identified that influence interval timing: attention. whereas emotional valence and drags when we are bored. France 2 Department of Psychology and Neuroscience. and motoric re-experien. Emotion might occur before cognition as the pathways from mean of the interval being estimated.008 . but can be extended to both shorter (e. Emotional intensity refers to extent that time seems to fly when we are having fun the strength of an emotional stimulus. millseconds) and longer (e. and describe how emotional arousal and valence inter. the circadian clock is where the participant is unaware of the importance of the duration of quite robust and can free run at high precision for several months an event before its occurrence and must reconstruct the temporal without external correction [1.2. whereas retrospective timing occurs in situations Although dependent upon light entrainment.21].18–26].1016/j.11 No. cing (collectively referred to as ‘‘embodiment’’) of the relevant emotion Scalar property/Weber’s law: one of the hallmark signatures of in one’s self’ (Ref. USA Our sense of time is altered by our emotions to such an thalamus. S.12 Cognitive-emotional interactions How emotions colour our perception of time Sylvie Droit-Volet1 and Warren H. social zontal shifts lead to changes in the point of subjective equality (PSE) for the bisection procedure and changes in peak time for the peak- conflict and everyday activities. Durham. perceiving a threatening object/event (e. that is time perception is like a rubber band in that it can be stretched to produce time-scale invariance across different durations [1. Meck2 1 Laboratoire de Psychologie Sociale et Cognitive. and we outline possible fear-related emotional states and remembering this object/event activates the originally experienced emotional states [54–56]. Expressed emotions: refer to the emotional responses to directly act to produce both increases and decreases in atten. Clermont-Ferrand. pp. like other animals. courtship. but has an advantage in increased Glossary flexibility in that it can run.fr). All rights reserved. These hori- which comprise music.2007. spiny neurons in the striatum serve as coincident detectors of specific patterns of oscillatory input from the cortex. [62].17]. which has loss of pacemaker pulses due to the effects of attention on the allowed us to gain a better understanding of the psycho. William James wrote: ‘our feeling of time harmonizes internal-clock models with different mental moods’ and inquired as ‘to what Inspection of internal-clock models (Box 1) suggests that cerebral process is the sense of time due?’ [4]. thus allowing the pulses generated by the pacemaker to enter the accumulator. At the onset of the event to be timed. This stimuli. circuits encode and decode time [58. the switch opens and stops the pulse transfer.65–68].12 505 by our emotions is not well understood [1–3]. Re-direction of attention can increase the latency of logical mechanisms underlying time distortions in switch closure and inattention can cause the switch to response to emotion. there remains considerable debate about how these Figure II. This article discusses the results of fluctuate between the closed and the opened states [21–23].2. cognitive psychologists have used an information-processing (IP) framework involving clock.16. However. In the SBF model as illustrated in Figure II. proces- emotion have frequently used nonstandardized emotional sing resources can be diverted away from the timer. from determine whether to respond. The accumulator thus achieves a monotonic integration of the pulses with the duration: the longer the stimulus duration. researchers are becoming increasingly aware that these accounts lack sufficient neurological specification to isolate interval-timing processes from other cognitive functions.com .60–62]. www. referred to as the striatal beat- frequency (SBF) model [1. According to these models. The striatal beat-frequency model. Adapted. the subjective experience of time is length- models of interval timing (Box 1). using comparison rules. of the processing of temporal information.64]. which make their findings inconsistent and diffi. the raw material for time representation comes from a clock stage consisting of a pacemaker that emits pulses at a given rate. More variations in our feeling of prospective time are caused by recently. As far back as Explanations of time distortions within the context of 1890. Although IP models have become highly influential within the field of interval timing. However. switch. clock perception not only involve cognition but are also inti. causes the subjective experience of time to be shorter than it cult to interpret [9–15]. Paul Fraisse has argued that timing and time different mechanisms – attentional time sharing. Although a growing body of evidence points towards the selective involvement of cortico-striatal circuits in timing and time perception [50. The current duration measured by the accumulator is compared. Consequently. Clock speed is altered in this system by the dopaminergic modulation of the cortical oscillation frequencies that are being input to the striatum. has been applied to several properties of interval timing.7]. The most prominent of these models. ened (durations seem longer than normal) by increases in Box 1. the pioneering studies on time and temporal and/or emotional events capture attention. Author's personal copy Opinion TRENDS in Cognitive Sciences Vol.63. the more pulses are accumulated. with permission. memory and decision stages to study the ability of humans and other animals to time events prospectively in the milliseconds-to-hours range [34]. speed and memory distortion – occurring at different levels mately connected to our emotional life [5].sciencedirect. the striatum is able to ‘read’ the temporal code provided by oscillating neurons in the cortex. The information-processing model. According to these information-processing models. such as attention. At the offset of this event. Ref. and the longer the duration is judged to be. from Ref. When this stimulus is timed. It has been whereas the investigation of emotion-related topics is proposed that these different mechanisms result in distinct growing rapidly [6. [34]. with permission. these new studies and integrates them with internal-clock Conversely. a variety of neurobiological frameworks for studying timing in the seconds-to- hours range have been developed to complement these IP models. and its synaptic connections are strengthened or weakened depending upon the occurrence of feedback and resulting dopaminergic input from the substantia nigra. the switch closes. with a sample drawn from a distribution of duration values stored in reference memory to Figure I. Internal-clock models of interval timing As illustrated in Figure I. memory and decision making [60]. This shortening effect can be explained by a have used standardized emotional material. analysis of the complex interplay patterns of timing behavior [1. a switch controlling how the pulses are gated and an accumulator in which the number of pulses is stored during the event(s) being timed.11 No. Adapted. Furthermore. temporal judgments also depend on memory and decision stages. several recent studies really is [18–20]. the oscillatory activity of a subset of cortical neurons is synchronized with the onset and offset of a relevant stimulus. including the effects of arousal/emotion on clock speed. between emotion and interval timing remains relatively There is now ample evidence to indicate that when non- rare [8]. In this way. This would be Angrilli et al. that negative stimuli produce a greater increase in arousal. This study is among the first to report that the effects adrenaline) versus the speed of the clock once timing has of emotion on time perception may be relatively short-lived begun (primarily DA-mediated) can also be differentiated and are maximal for sound durations around 2 s. [1. happy baby) were over- ued training under conditions in which clock speed is estimated. The pattern start the clock will shift all timing functions to the left by of results found with the IAPS appeared complex: there an absolute (e. Recently. By contrast. 3-4 s. For example. The main question is to determine strongly correlated with changes in emotion increased with which of these mechanisms (if any) are involved in the the level of arousal. At the same level of high arousal. it has www. such as haloperidol and (IAPS) [32]. that is decreasing the latency to arousal to validate the source of these effects. causing problems for interpretation [26]. The clock-speed effect is were judged to be longer than positive ones. that is increas.29]. effects on the sistent with models of interval timing that use attention to latency to initiate and/or terminate the timing process modify the allocation of processing resources to temporal involve the closure or opening of the switch that allows versus non-temporal factors [18. a consistent picture of the effects of body temperature to manipulate arousal through metabolic emotion on time perception has been difficult to discern. increases arousal mined this issue using standardized stimulus sets. smashed head) was overesti- effects on encoding and memory content [1].g. This sensitivity of clock speed to arousal has been documen. such and produces an overestimation of durations. in processes [25]. whereas that of positive pictures (erotic scenes) was on clock speed appear to be immediate.g. [24]). magnitude. clock shorter. their finding that high-arousing stimuli were larger overestimations as the target duration of the inter. as if the internal clock was running slower and arousal (low/high). and generalization. negative transient in the face of feedback. and positive pictures (pet dogs. tortion during translation [17. decreases arousal and produces an underestima. whereas DA primarily affects clock speed. emotion-induced activation temporarily increases the tional processes. increased effect of emotion on time perception as a function of the arousal accelerates the pacemaker. [33] recently found Other aspects of the internal-clock model can also be that emotional sounds were judged longer than neutral distinguished by their pharmacological profiles. was no main effect of arousal or valence. By contrast. as ascertained by heart rate. suggesting proportional to the durations being timed. for durations up to 4 s. 800 ms) rather than a proportional (e. thus causing more emotional stimuli used.17]. Arousal effects mated.g. attention to the effects of valence (unpleasant/pleasant) tion of durations. several studies have exa- such as cocaine and methamphetamine.19]. For sounds. the duration of negative pic- fications of the memory-translation constant related to tures (baby with eye tumor. rat in the dirt) were underestimated are able to recalibrate their timing functions with contin. or administered drugs that modulate arousal part because most early studies used idiosyncratic by altering the effective levels of dopamine (DA) in the brain emotional stimuli. ceived durations) until clock speed returns to baseline after The latency to begin timing (primarily mediated by nor. which is as the International Affective Digital Sounds system characteristic of an increase in clock speed. the high-arousal condition.11 No. Noulhiane et al.sciencedirect. pulses to be transferred to the accumulator. example. In addition.com . Of behaviorally. did positive pictures. Author's personal copy 506 Opinion TRENDS in Cognitive Sciences Vol. negative pictures elicited a stronger orienting reaction than To summarize. Using the IADS. [35] used the IAPS stimuli to investigate expected to produce an absolute effect on the horizontal the effects of valence and arousal on time perception and placement of psychophysical functions regardless of the used converging measures of attention and physiological duration being timed. According to the internal-clock models. In the low-arousal condition. speed of emer. perceived to be shorter than low-arousing stimuli is con- val being timed increases. interpreted these intriguing data within ergic drugs have their primary effects on memory for time the context of scalar timing theory [34] and proposed that [17]. but relatively underestimated. which affects atten. In the case of low arousal. This memory mechanism for low arousal and an automatic mechanism ‘corruption’ appears to be relatively permanent even in the related to motivational-survival systems for high arousal. regardless of arousal. The effect of emotional stimuli (sounds and pictures) ted in studies that have presented a repetitive series of rated for valence and arousal on timing and time auditory clicks or visual flickers designed to increase arousal perception before the timed event (for example. ing the rate of pulse accumulation leads to larger and In addition. altered As mentioned earlier. We focus on these studies here.12 arousal. skin conductance measures that are or memory) are affected. attention. the administration of psychostimulants. face of corrective feedback due to the multiplicative dis.30]. cholin. The release of noradrenaline. distortions in the content of the arousal level suggests that two different mechanisms temporal memory emerge gradually as new clock readings are triggered by arousal levels: a controlled-attention are translated and encoded into memory. whereas the (IADS) [31] and the International Affective Picture System administration of antipsychotics. the direction. This opposite effect of valence as a function of modified. appears to impact time perception by speed of an internal clock (thereby leading to longer per- modifying the latency to close or open the switch [27]. more attention was paid to negative gence and robustness of temporal distortions vary as a than to positive pictures and the former was judged as being function of which specific mechanisms (e. but there was a 20%) amount of time. significant interaction between these two dimensions. In Clock-speed effects can also be distinguished from modi. with particular pimozide. see Ref. because the participants pictures (dead cow. pulses to accumulate within the same physical unit of time. Noulhiane et al. provided that several durations are used in particular interest is their report that negative sounds the same experiment [28. [35] and lished)) on the processing of emotional facial expression Noulhiane et al. These data also suggest Time and facial expression that negative images in the high arousal condition activate Faces are often used in studies on emotions. given arousal/unpleasant pictures for the 2 s duration.. As suggested by Angrilli and col. with a basic fore. attention-related mechanisms presumably prevail pictures that elicit disgust? Consistent with theories at longer durations because of the expected diminution of suggesting a close relationship between emotion and action the autonomic response within a few seconds of the stimulus readiness [38. escape). [35] found a temporal overestimation of high. This causes [18. For example. escape and attack. www. the findings of Angrilli et al. fewer pulses accumulated. but not for the same high-arousal level. distortion similar for pictures that elicit fear to that for leagues.. consequences for temporal processing. and their Angrilli et al. a description of associated with a speeding up of the clock. in. Adapted.D. in comparison to perceiving other people’s emotional states is essential for Figure 1. autonomic nervous system activity (e. the capture of attention by the features ing. Consequently. Gil and S. copulation and care giv. Author's personal copy Opinion TRENDS in Cognitive Sciences Vol. the internal clock will run relatively faster motivational systems: one defensive and one appetitive. under high arousal conditions for negative images and lead As stated by Bradley [36]. that define the emotional valence of the stimulus diverts which require defensive motor behaviors. and a basic behavioral the low arousal condition will be underestimated because repertoire is built on withdrawal. there- cluding sustenance.12 507 been proposed that emotions are organized around two positive images. Con. negative pictures in activated in threatening contexts. high-arousal/unpleasant conditions.-V. muscles contract) that would be the basic function of emotions. pupils dilate.g. the appetitive system is activated in contexts. with permission. unpub- Taken together.19]. [33] emphasize the importance in time suggest that the urgency of action for incoming events is studies of investigating the temporal dynamic of emotions. is the magnitude of time the longer durations. recent studies conducted by Droit-Volet presentation. the capture of attention by those pictures means that less versely.-V. Consequently. because the effects of emotion might change in a systematic fashion with the passage of time. the defensive system is primarily to their overestimation. involve the ability processing resources away from the timing system itself to produce a rapid reaction (attack. An example of one of the female faces with the neutral expression (center) and the six emotions tested (upper and lower rows). procreation and nurturing. This might explain why different adaptive functions of emotions [37]. Indeed.11 No. the (valence and arousal) is insufficient to account for the readiness for action is more rapid. and her collegues ([40–44] and (S. attention will be allocated to the timing system and. By contrast. blood pressure Time perception thus appears to be a sensitive index of increases. arousal conditions. because the defensive system.sciencedirect. from (S. one of the crucial factors affecting our perception of time. In other words. when emotional space involving only two major dimensions the passage of time is experienced as passing faster. Gil and S.D. unpublished). under low behavioral repertoire of ingestion.com .39]. However. [26].sciencedirect. Open circles indicate rats trained under continuous footshock stress (FSS) and tested without the stressor (control).12 Figure 2. (a) Median proportion ’long’ response as a function of signal duration during seven-signal duration bisection training (left panel) and testing (right panel). Failure to divide attention results in the sequential www. (b) The role of the amygdala in the ability to divide attention between two signals and time them simultaneously was investigated under both appetitive and aversive conditions using peak-interval procedures. from Ref. with permission.com . Author's personal copy 508 Opinion TRENDS in Cognitive Sciences Vol. Adapted.11 No. closed circles indicate rats trained without the stressor and tested under continuous FSS. In the same manner. [13] have also shown that blindfolded actions [45].e. unpublished) ation because the partner could become aggressive. Author's personal copy Opinion TRENDS in Cognitive Sciences Vol. especially arousing. respectively. 60 s) due to the effect of selective attention. These other threatening situations. In this task. whereas rats with frontal cortex lesions demonstrated sequential processing under all conditions. [40] investigated how the participants overestimated time more when moving on a perception of facial expressions of emotion affects our platform towards a drop (danger) than when moving away experience of time by using the duration bisection task from it (safety). the temporal pattern was differ- similar horizontal shift to the left in bisection functions ent. The similar patterns of time spent in an eye-contact task was judged longer with temporal bias obtained in different threatening contexts an angry than with a friendly partner. speed and. By contrast. followed by happy and then sad faces.com . duration dimension of those stimuli associated with the indicating that the presentation duration of angry faces fear-provoking event (i. Droit-Volet et al. which indicates a temporal underestimation. this is a similar result to that observed after summarize. Single vertical lines indicate when the peak time was at the programmed time of reinforcement for the visual signal (e.-V. this temporal bias with angry faces occurs at an Although arousal is clearly a very important factor in early age (3 years).11 No. increased their temporal overestimation. emotion [48]. same context). The results showed a an increase in arousal that leads to acceleration in clock significant leftward horizontal shift of the bisection func. suggest that the internal clock speeds up when the emotion amine [26] (Figure 2a – right-hand panel). because methamphetamine produces similar result is expected because shame.. These provoking stimulus can be abolished by inactivation of the findings are consistent with those previously reported by amygdala. radiofrequency lesions of the amygdala. an automatic. judged longer by arachnophobic participants than by other mediate value) as being more similar to the short or to the non-arachnophobic individuals. In (Figure 3). it has been systematically results are consistent with the idea that anger tends to be shown that humans and other animals tend to overesti. Gil and S. there is expression (as shown in Figure 1). www. However. long standard durations. Impairments in simultaneous temporal viewing angry faces distorts time perception. and less attention is directed toward the timing judged a particular intermediate duration as either ‘short’ of other neutral or appetitive events occurring within the or ‘long’) did not change with the facial expression. or aspiration lesions of the frontal cortex. which suggests that it reflects a factor. Further. for other facial expressions. the amygdala is crucial for the emergence of fear-evoked increments in selective attention leading to deficits in simultaneous temporal processing. Adapted. They were then required to classify reported that a 45 s period spent watching a spider is comparison durations (equal to the standards or of inter. the imminence of the danger [46. increased attention to the tion for angry faces compared with the neutral faces. These results support the proposal that the frontal cortex exerts primary control over the allocation of attentional resources but that.D. faces produced the most temporal bias.12 509 the determination of appropriate behavior in social inter. inactivation of the amygdala blocked this fear-related impairment and allowed rats to time both signals simultaneously. cating that the lengthening of time estimates resulted from as described by evolutionary theories of anxiety and an arousal-related acceleration of clock speed. unconscious program of responses to danger. Droit-Volet and colleagues [40] found that angry basic phenomenon of adaptation in social interactions. 50 s) and double vertical lines indicate when the peak time was shifted to the right by 10 s (e. such that processing resulting from selective attention to the fear- event durations are judged to be longer than normal. Gil and Droit-Volet (unpublished) observed no change when electric footshocks are administered during a test in bisection functions for disgust. timing the specific events associated with the fear eliciting sensitivity to time (the consistency with which participants stimulus. Langer et al. To sessions. another emotion. Further analyses of in both rats and humans at different ontogenetic stages the data indicated that this effect was multiplicative (i. happiness or sadness) and a neutral-baseline confronted with angry faces or electric footshock. mate the duration of events. these results showed that a period of and illustrated in Figure 2b. in some cases. while the other signal is selectively attended to. from Ref.e. control and amygdala- lesioned rats were able to divide attention and time both signals simultaneously. Furthermore. which is a social–moral results (Figure 2a – left-hand panel). there is a growing body of evidence indi- presented in the form of faces with three different emotions cating that in threatening situations. adults. that is a short greater when they stopped at 15 rather than 20 feet from (400 ms) and a long duration (1600 ms). for the short durations tested. rats show a such as disgust or shame. For example. it is not the only stages of development [42].g. more attention is directed toward was judged to be longer than for neutral faces. for example when (anger. suggest that these responses to fear-eliciting stimuli reflect overestimations were greater for longer durations). which was nate between two standard durations.sciencedirect. participants were trained to discrimi. This effect appears to be due to increased shame. By contrast. and a shift to the right for phase [26]. control rats were impaired in their ability to divide attention and were able to time only one signal at a time. more. Thus. The redirection of attention would lead to a functions when the footshock was discontinued during test loss of pulse accumulation and lower clock readings [19]. The effects of amygdala or frontal cortex lesions were compared in rats trained to time both a 50 s visual signal paired with food and an embedded 10 s auditory signal that was paired with either food (top row) or footshock (bottom row). This arousal. these results haloperidol injections or the withdrawal of methamphet. and does not change across different the effects of emotion on time perception. presented in the the drop. These comparison durations were In summary. In human perceived in the other individual involves readiness for processing of the signals and the suspension of timing for one of the signals.g.47]. indi. By contrast. captures attention by a reflexive activity of self- group of rats trained to time under the stressful footshock awareness that directs attention away from the stimulus to condition exhibited a shift to the right in their timing be timed [49]. whereas when the embedded auditory signal was paired with footshock. under stressful conditions. were a close second (S. Watts and Sharrock [14] form of a pink oval. and fearful faces Seeing an angry face might present a threatening situ. Columns (left to right) represent groups of rats with control surgeries. with permission. When both signals were paired with food. The 10 s auditory signals (filled circles) were randomly embedded within the 50 s visual signals (open circles) and are plotted at their average time of occurrence. as demonstrated by Meck and MacDonald [20] Schiff and Thayer [10. [20]. motor output [50] as well as in the processing of anger [51]. happiness (non-Duchenne smile) or shame. Gil and S. with permission. Author's personal copy 510 Opinion TRENDS in Cognitive Sciences Vol. [40]. The effect of emotional faces on time perception in children and adults. www. It is thus reasonable to suppose that shown that dopaminergic function in the basal ganglia the urgency to produce an immediate action (avoidance/ plays an important role in the efficiency of timing and withdrawal) is higher in the presence of an angry individ. Recent studies have action (angry faces). the disgusted faces did not produce any time distortions. d’ is the difference between the proportion of ‘long’ responses for an emotional facial expression and that for the neutral expression. Contrary to the other facial emotions.sciencedirect. but only with the Duchenne smile (shown in Figure 4). The panels show the results found by Gil and Droit-Volet (unpublihsed) with children aged 5 and 8 years old.-V. The results are consistent with those found by Droit-Volet et al. disgust. the emotional stimulus attracting attention away from the processing of temporal information. although the 5-year-olds over- reacted to sadness compared with the older participants. Further analyses revealed that the temporal overestimation was linked to the emotional expressions that increased the level of arousal. A d’ greater than zero indicates that the emotional expression is overestimated compared with the neutral expression. Adapted. They showed that anger and fear produced an overestimation of time with a magnitude that did not vary with age. shame produced an underestimation of time. the participants had to categorize as ‘short’ or ‘long’ the presentation durations (between 400 and 1600 ms) of faces expressing a specific emotion or neutrality. except for the angry faces that were relatively less overestimated in the 8-year-olds. the facial expression of sadness also produced an overestimation of time. the temporal underestimation was linked to the attention- related processes. In a duration bisection task. sadness. and a d’ shorter than zero indicates that the emotional expression is underestimated compared with the neutral expression. action.11 No. except in the youngest children.com . unpublished). By contrast. and not with the non-Duchenne smile. The happy faces also produced an overestimation of time. [40] and (S.12 Figure 3. from Ref. By contrast.. and does so to a greater extent in the case of urgent ual than of a disgusted individual. thus in turn increasing the rate of the pacemaker of an internal clock. fear. To a lesser extent. as well as with adults.D. when the participants were presented with faces expressing neutrality and a specific emotion in the same session: anger. [54]). [41]. the neural system that is activated of time going faster might make one happier with the performance of in the observer of an emotion is just as if they were actually a web browser. Mondillon et al. for reaction time. The figures show the d’-score plotted against the stimulus durations. It has been shown that individuals automatically imitate perceived facial expressions and that this voluntary adoption of emotional facial expressions produces autonomic changes that are correlated with the emotional states [57].sciencedirect. by an imitation process. with permission. In an elegant demonstration of the effect of embodied emotion on time perception. and a d’ shorter than zero indicates that the emotional over the course of the duration of an event remains to be determined. Variation in current emotions might also alter the perception of past events. clock? The expression of an emotion might elicit a different including their durations. In the inhibited imitation diminish within a few seconds. [43] also found an ‘in-group’ advantage with a temporal bias occurring when Caucasian partici- pants were presented with Caucasian. that is a shift of clock speed in the opposite direction from the initial effect when the emotional stimulus is removed. The effect of emotional faces on time perception in adults. In this duration ships as a selective filter or as an inhibitory mechanism of bisection task. arousal and valence. the perceptual systems relay information to neurocognitive mechanisms underlying emotional experience via an unconscious motor simulation of viewed expressions [54–56]. is an extremely fruitful Time and embodied emotion avenue for future research that might challenge the types of internal- How can the perception of an emotion. Outstanding questions for an emotional facial expression and that for the neutral expression. a video) are uncertain as is the occurrence of rebound Adapted. and motor control centers [48. these results point to a faces with two different emotional expressions (anger and happiness) and a neutral- baseline expression. we might assume that it is because we feel in our own body the anger of another. The nature of the social relationship with other individuals (mother–child. facial expressions. effects. various decreases in clock speed as well as the stimulus features controlling attentional time sharing and/or clock speed. that increases clock speed. The results Current evidence suggests that the changes in clock speed after the showed that. not simply the percep. For the motor simulation version of the embodi- www. clock models reviewed in the current article because of the need to expressed by another individual accelerate our internal recreate these emotions and events from memory. It remains to be determined whether this social factor regulates emotion–time relation- Figure 4. However. On the basis of these results on time perception. The d’ is the difference between the proportion of ’long’ responses (stimulus duration judged as more similar to the ’long’ than to the ’short’ standard) Box 2. but not with Chi- nese. colour. see layout and dynamics of graphical content) are likely to influence attentional timing sharing and/or clock speed [69]. lines of evidence indicate that there is a close connection ! How time perception can affect emotional satisfaction is an area between the neural substrates of emotion perception and that is particularly ripe for investigation. stimuli (e.g. For example. in the spontaneous imitation condition (upper panel). expression is underestimated compared with the neutral expression. especially with regard to autobiographical memories.g. and that this overestimation was greater for anger than for happiness. Mondillon et al. it might be fear. the presentation duration (between 400 and 1600 ms) of imitation.com . such as anger. ! The study of emotion and retrospective time perception.52]. the participants had to categorize as more similar to a ’short’ or a ’long’ standard duration. to explain differences in the emotional reaction to increases or tion of anger. experiencing this emotion themselves (for a review. but the effects of more dynamic condition (lower panel). [41] found that the overestimation of the duration of presentation of angry faces was not observed when the imitation of facial expression was inhibited by a pen held between the participants’ lips (as illustrated in Figure 4). probably related to the general role of DA photographs or faces versus simple lights or sounds) will help to on the rapid access of relevant incoming events to timing address the issue of how attentional time sharing is influenced by the nature of the stimulus – including social aspects. Author's personal copy Opinion TRENDS in Cognitive Sciences Vol. and various aspects of the browser (e. the presentation presentation of a static emotional stimulus (e. In a similar study. boss–employee) that motivates us to imitate them might thus modulate the effects of facial expressions on timing and time perception. emotion in an observer.g.g. that we also become angry and/or aroused. Ref. Whatever the case. the d’ was close to zero indicating no time distortion. an expression of ! The role of time in the regulation of social behavior and cognition anger is more likely to elicit fear than anger in the observer as a function of normal aging or/and different pathologies might help [53]. from Ref. Effron et al.12 511 ment theory of emotions. [43] determined an empathy quotient and found a reduced temporal overestimation for angry faces in the low empa- thy compared with the high empathy participants. For example.11 No. the perception emotional experience. A d’ greater than zero indicates that the emotional expression is overestimated compared with ! How the effects of emotion on timing and time perception evolve the neutral expression. a photograph) duration of angry and happy faces was significantly overestimated. ! Determination of the variation in emotional effects on interval This finding suggests a common arousal-based mechanism timing as a function of the nature of the stimulus events (e. Consequently. 269–287 International Affective Picture System. (2005) What makes us tick? Functional 30 Meck. J. Endeavour 21.eds). L. (2007) Anger and temporal perception in children. Brain Res. 8) (Carterette. W. (1996) Neuropharmacology of timing and time perception. Exp. R. and Meck. 6. Perform.D. Cogn. S. 4. 43 Mondillon. 2. 849–858 positive. D. R. Behav. J. 195–205 29 Droit-Volet. 1. Skills 26. J.H. 59. For example. A. 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Trends Cogn.J. M. Locate the article with the required figure on ScienceDirect and click on the ‘Full text + links’ hyperlink 2. Brain Res. 156–161 functional link between mesolimbic. and Adolphs.com/locate/permissions www. 307–316 interval timing: coincidence-detection of oscillatory processes.. M. Copy the image and paste it into an image-display program Permission of the publisher is required to re-use any materials from Trends. (2004) Frontal-striatal circuitry activated dopaminergic systems. 1109. W. (2004) Waiting for the web: how screen color affects difficulty dissociates clock functions from other cognitive demands. Psychol. V. (2007) A further analysis of temporal Française 50. Brain Mapp. and Craighero. (2006) The substantia nigra pars compacta and 55 Gallese.J. (2006) Neuroanatomical localization of an internal clock: a Curr. S.H. S. G. Brain Res. and mesocortical 63 Hinton. W. A. (2007) Time perception: manipulation of task 69 Gorn.elsevier. J. Bioessays 22. et al. 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