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An internal clock model of interval time perception. A graphical representation of the components of information processing that occurs while perceiving short durations. Pulses are counted by an accumulator. The output of the accumulator is transformed into a memory representation. The comparator judges the current memory trace against previously formed representations to make a temporal discrimination. 

An internal clock model of interval time perception. A graphical representation of the components of information processing that occurs while perceiving short durations. Pulses are counted by an accumulator. The output of the accumulator is transformed into a memory representation. The comparator judges the current memory trace against previously formed representations to make a temporal discrimination. 

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Models of time perception often describe an "internal clock" that involves at least two components: an accumulator and a comparator. We used functional magnetic resonance imaging to test the hypothesis that distinct distributed neural networks mediate these components of time perception. Subjects performed a temporal discrimination task that began...

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... fMRI studies of interval timing have been moti- vated by variations of an information processing "internal clock." However, their designs and conclusions are not often tied to the components of these models. As originally proposed by Gibbon (1977), this class of models proposes three compo- nents in perceiving interval timing durations: an accumulating clock component that produces the timing signal, a memory component where the output of the accumulator is stored, and a comparator component where decision processing occurs (Fig. 1). A switch or gate is proposed to signal the accumulator that a task-relevant event is beginning and ending (Buhusi and Meck 2006). Attention is one mechanism hypothesized to influence the switch's functioning (Meck 1984;Meck and Benson 2002;Penney et al. 2000). Internal clock models have been successful in accounting for much of the psychophysical data relating to interval timing. Unfortunately, functional im- aging studies have not consistently dissected these compo- nents. Therefore little is known about the neuroanatomical structures underlying these component ...

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... /fncom. . species (Harrington et al., 1998;Malapani et al., 1998;Hinton and Meck, 2004;Matell and Meck, 2004;Wencil et al., 2010;Coull et al., 2011;Adler et al., 2012;Merchant and de Lafuente, 2014;Emmons et al., 2016Emmons et al., , 2017Emmons et al., , 2020Dallerac et al., 2017) trained to discriminate long and short intervals show significant fMRI activation in the caudate in particular (Rao et al., 2001;Pouthas et al., 2005). Several studies have employed striatal inactivation by lesions or pharmacological and genetic manipulations and have shown that doing so disrupted the animal's estimate of elapsed time and impaired performance (Drew et al., 2003(Drew et al., , 2007Meck, 2006;Gouvea et al., 2015). ...
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... Great attention has been given to the SMA which, in connection with the striatum, might provide the neural substrate of the accumulation process commonly postulated in models of time processing . This putative role is supported by a high number of fMRI studies showing that SMA activity during timing perceptive task increases with physical duration Wencil et al., 2010), perceived duration (Tipples et al., 2013), or the amount of attention allocated to the duration of stimulus (Coull et al., 2004;Henry et al., 2015;Herrmann et al., 2014). Furthermore, Coull et al. (2015) suggested that SMA selectively codes the accumulation of temporal magnitude by showing that its activity increased incrementally as a function of both physical and perceived duration, but not as function of distance. ...
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... However, one alternative interpretation for the successful decoding performance is the existence of firing-rate coding of the stimulus duration, which may correspond to the accumulator in the clock-counter model of time perception 27 . The existence of an accumulator predicts greater overall BOLD responses as the stimulus duration increases [28][29][30] . To examine this possibility, the values of correlation coefficients (i.e., betas) for the S1 offset estimated by a GLM (refer to GLM analysis on main scans in the online Methods for details) were extracted and averaged across all voxels within each ROI (Fig. 4e-h). ...
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Precise time estimation is crucial in perception, action and social interaction. Previous neuroimaging studies in humans indicate that perceptual timing tasks involve multiple brain regions; however, whether the representation of time is localized or distributed in the brain remains elusive. Using ultra-high-field functional magnetic resonance imaging combined with multivariate pattern analyses, we show that duration information is decoded in multiple brain areas, including the bilateral parietal cortex, right inferior frontal gyrus and, albeit less clearly, the medial frontal cortex. Individual differences in the duration judgment accuracy were positively correlated with the decoding accuracy of duration in the right parietal cortex, suggesting that individuals with a better timing performance represent duration information in a more distinctive manner. Our study demonstrates that although time representation is widely distributed across frontoparietal regions, neural populations in the right parietal cortex play a crucial role in time estimation.
... These results are also in line with the hypothesis that supplementary motor area acts as an "accumulator" of temporal information (Macar et al., 1999;Macar and Vidal, 2002;Casini and Vidal, 2011;Coull et al., 2015). Indeed, consistently with previous EEG studies reporting a positive correlation between Contingent Negative Variation amplitude and subjective duration (Macar et al., 1999;Macar and Vidal, 2002;Pfeuty et al., 2003), recent evidence showed that supplementary motor area activity increases as a function of stimulus duration (Wencil et al., 2010), subjective temporal dilation (Tipples et al., 2013), and sequential integration of temporal information (Coull et al., 2015).Finding that the basal ganglia are consistently activated in the context of both internally-based and externally-cued tasks is not surprising, considering that this region has long been implied in fMRI studies of timing, both in the context of internally-based (Rao et al., 2001;Belin et al., 2002;Ferrandez et al., 2003;Bueti et al., 2008a;Bueti and Macaluso, 2011) and externally-cued timing tasks (Dhamala et al., 2003;Grahn and Brett, 2007;Grahn and McAuley, 2009;Teki et al., 2011;Grahn and Rowe, 2013). Indeed, one of the most influential theories on the anatomical substrates for temporal processing -the Striatal Beat Frequency model -acknowledges cortico-striatal interaction as crucial for timing, with striatal medium spiny neurons playing a pivotal role in integrating cortical oscillatory inputs and detecting oscillation patterns coincidence (Matell et al., 2003;Matell and Meck, 2004). ...
... The interpretation of the role of inferior frontal gyrus in general timing processes is less straightforward: indeed, the specific contribution of frontal regions to timing and time perception has not been fully elucidated yet (Cerasa et al., 2005;Coull et al., 2011). Activation level of bilateral inferior frontal gyrus, similarly to supplementary motor area, has been shown to increase with stimulus duration (Wencil et al., 2010;Harrington et al., 2010). It has thus been proposed that inferior frontal gyrus collaborates to the accumulation of temporal information, possibly interrupting the accumulation process in SMA at the end of the to-be-timed intervals (Harrington et al., 2010). ...
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... Los resultados también corroboran la implicación de un circuito fronto-estriatal, que sería la base del funcionamiento del reloj interno. En este circuito se destaca el rol de la corteza frontal dorsolateral derecha 41-43 y el del área motora suplementaria y pre-suplementaria, como contador o acumulador del tiempo [44][45] . ...
... Estudios que utilizan técnicas muy diferentes proponen que el área motora suplementaria (AMS) está relacionada con la función de acumulador del tiempo 34,[45][46][47] . ...
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... 3, 4 Under pacemaker-accumulator models of time perception, neural signals are collated, counted and compared to stored representations of interval durations to which future durations and time estimations are based (see Figure 1. for depiction of internal time perception model). 5 ...
... This depiction is from a study by Pencil, Coslett, Aguirre and Chatterjee (2010) and was created to discuss the neural bases for interval timing. 5 The authors concluded that the neurophysiological effects of high state-anxiety disrupted the attentional resources of the participants because the participants' attentional resources were directed to negative thought processes, thus leading to their performance changes. 9 Although this was not a time perception study, because time perception is hypothesized to be a process utilizing attentional resources it is reasonable to suggest that if emotional arousal altered attentional processes time perception processes would also have been altered. ...
Article
Participants in this study were randomly assigned to one of three interval groups of either one, three, or five minutes. All participants were asked to estimate a group-assigned time interval and complete the state portion of the State Trait Anxiety Inventory (STAI). It was hypothesized that higher levels of state anxiety would cause participants to overestimate the passage of time. It was also hypothesized that shorter interval durations would be estimated more accurately than longer interval durations. Results of a t-test did not support the first hypothesis. Results of a linear trend analysis did support the second hypothesis (P < .05). These results indicate that state-anxiety does not cause the passage of time to be overestimated and that interval duration length affects how accurately time is estimated. Implications of this study are important to the understanding of human time-management ability and time estimation errors. KEYWORDS: Time estimation; Perception; State-anxiety; Attention; Working memory; Emotion
... A meta-analysis study reported the presence of brain regions that are activated in both sub-and suprasecond timing (Wiener et al., 2010). Several functional magnetic resonance imaging (fMRI) studies have reported that BOLD activities in various brain regions correlate with event durations at timespans ranging from the milliseconds to seconds range (Morillon et al., 2009;Wencil et al., 2010;Coull et al., 2015). These fMRI studies, however, assumed a monotonic change of BOLD activity across sub-and supra-second timing, and did not directly investigate the distinction between sub-and supra-second timing. ...
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Recent neuroimaging studies have revealed that distinct brain networks are recruited in the perception of sub- and supra-second timescales, whereas psychophysical studies have suggested that there are common or continuous mechanisms for perceiving these two durations. The present study aimed to elucidate the neural implementation of such continuity by examining the neural correlates of peri-second timing.We measured neural activity during a duration reproduction task using fMRI. Our results replicate the findings of previous studies in showing that separate neural networks are recruited for sub- versus supra-second time perception: motor systems including the motor cortex and the supplementary motor area for sub-second perception, and the frontal, parietal, and auditory cortical areas for supra-second perception. We further found that the peri-second perception activated both the sub- and supra-second networks, and that the timing system that processed duration perception in previous trials was more involved in subsequent peri-second processing. These results indicate that the sub- and supra-second timing systems overlap at around 1 second, and cooperate to optimally encode duration based on the hysteresis of previous trials.
... Modern theories about internal time representation share the view that the processing of temporal information is mediated by a distributed network with varying engagement of its individual components depending on the task requirements [3]. However, there is substantial disagreement on the effects reported in various studies, providing no definitive anatomical or cognitive model [3][4][5]. Several regions, including the basal ganglia, cerebellum, posterior parietal cortex, and frontal cortex, have been implicated as relevant to interval timing; however, their precise role remains shrouded. ...
... However, recent advances point towards an alternative view offered by distributed timing models, deriving temporal information from the coincidental activation of different neural populations [1]. Although diverse, regarding the hypothesized specific neuropsychological mechanisms of time processing, this broad spectrum of models basically postulate the representation of time as ubiquitous in more networks that also encode other 2 Neural Plasticity stimulus properties [4,9]. Thus, temporal information may be encoded in the entire activity pattern of a neuronal mechanism, as suggested in the state-dependent network model [9,10]. ...
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Time perception is an essential part of our everyday lives, in both the prospective and the retrospective domains. However, our knowledge of temporal processing is mainly limited to the networks responsible for comparing or maintaining specific intervals or frequencies. In the presented fMRI study, we sought to characterize the neural nodes engaged specifically in predictive temporal analysis, the estimation of the future position of an object with varying movement parameters, and the contingent neuroanatomical signature of differences in behavioral performance between genders. The established dominant cerebellar engagement offers novel evidence in favor of a pivotal role of this structure in predictive short-term timing, overshadowing the basal ganglia reported together with the frontal cortex as dominant in retrospective temporal processing in the subsecond spectrum. Furthermore, we discovered lower performance in this task and massively increased cerebellar activity in women compared to men, indicative of strategy differences between the genders. This promotes the view that predictive temporal computing utilizes comparable structures in the retrospective timing processes, but with a definite dominance of the cerebellum.
... SMA activity varies with the duration of the sensory stimulus itself, with greater activity for longer durations [13], and correlates both with individual differences in the propensity to overestimate stimulus duration [14] and individual susceptibility to temporal illusion [15 ]. Indeed, Coull et al. [16 ] showed that SMA activity increases both as a function of objective physical stimulus duration as well as subjectively perceived stimulus duration ( Figure 2). ...
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Traditionally, the Supplementary Motor Area (SMA) is linked to motor control, including voluntary movement selection and response inhibition. However, it is also implicated in timing, whether a timed motor response is required or not. We synthesize recent neuroimaging, electrophysiological and stimulation studies of timing and action control and reconcile these distinct literatures by proposing a functional gradient for voluntary action within SMA: preSMA controls the voluntary selection and inhibition of stimulus-specified action whereas SMA proper instigates spontaneous, self-generated action. In addition, developmental and learning studies indicate that we develop and refine our sense of time through action, potentially explaining why even non-motor, perceptual forms of timing recruit SMA. We suggest that SMA is engaged through action control to build up a sensory representation of time.
... In the temporal discrimination paradigm, in which the duration of a probe stimulus is compared to that of a prior standard, encoding and storage of duration into WM occur during presentation of the standard stimulus, whereas retrieval and comparison of stimulus duration occur during presentation of the probe. Event-related fMRI studies of temporal discrimination paradigms, in both the auditory (Harrington, Zimbelman, Hinton, & Rao, 2010;Rao et al., 2001) and the visual Wencil, Coslett, Aguirre, & Chatterjee, 2010) domains, have found that the basal ganglia are activated selectively during timing of the initial standard, whereas the frontal and temporal cortices are activated during timing of the probe. These convergent findings strongly suggest that the basal ganglia mediate the shortterm representation of time in WM. ...
... These convergent findings strongly suggest that the basal ganglia mediate the shortterm representation of time in WM. Incidentally, common to timing of both standard and probe stimuli was the SMA ( Harrington et al., 2010) whose activity has also been shown to increase as a function of stimulus duration (Morillon et al., 2009;Wencil et al., 2010). Given that timing of a currently elapsing duration is the only process common to both stages of the task, these data strongly support a role for SMA in timing an ongoing duration. ...
... For example, timing-induced activity in the putamen and right inferior frontal cortex is higher in those individuals with better temporal discrimination performance (Wiener et al., 2014), while activity in the SMA and right inferior frontal cortex is greater in participants who subjectively perceive time as lasting longer (Tipples, Brattan, & Johnston, 2013). The latter finding in particular confirms and extends previous findings that activity in the SMA and inferior frontal cortex also increases in line with the objective duration of a stimulus (Pouthas et al., 2005;Wencil et al., 2010), underlining the critical role of these areas in the 'accumulation' of information over time. ...
Article
Timing the duration of events recruits the supplementary motor area (SMA), right inferior frontal cortex (around the operculum and anterior insula), and basal ganglia. This frontostriatal circuit represents duration, at least in the range of hundreds of milliseconds to seconds, in an amodal, context-independent manner. Distorting the subjective perception of time, through visual illusion, emotional context, or pharmacological manipulation, modifies activity within this circuit. Moreover, distinct nodes appear functionally specialized: The SMA is implicated in accumulating temporal information, the inferior frontal cortex in updating temporal information in working memory, and the putamen in encoding temporal information into memory (potentially contributing to rhythm perception).