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Properties of the internal clock: First- and second-order principles of subjective time.

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Abstract

Humans share with other animals an ability to measure the passage of physical time and subjectively experience a sense of time passing. Subjective time has hallmark qualities, akin to other senses, which can be accounted for by formal, psychological, and neurobiological models of the internal clock. These include first-order principles, such as changes in clock speed and how temporal memories are stored, and second-order principles, including timescale invariance, multi-sensory integration, rhythmical structure, and attentional time-sharing. Within these principles there are both typical individual differences—influences of emotionality, thought speed, and psychoactive drugs—and atypical differences in individuals affected with certain clinical disorders (e.g., autism, Parkinson’s disease, and schizophrenia). This review summarizes recent behavioral and neurobiological findings and provides a theoretical framework for considering how changes in the properties of the internal clock impact time perception and other psychological domains.

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... Human perception of time is an important dimension where people make decisions for everyday behavior and survival (Wittmann and Paulus, 2008). Time perception relates to our awareness of the passage of time; this experience is intertwined with environmental, psychological, and physiological processes (Wittmann and van Wassenhove, 2009;Wittman, 2013;Allman et al., 2014). For instance, it is a common perception that time passes more slowly when a person is bored or passes by more rapidly for adults than children (Burdick, 2017). ...
... For instance, it is a common perception that time passes more slowly when a person is bored or passes by more rapidly for adults than children (Burdick, 2017). While the neural basis for time perception is still unknown (Wittmann and van Wassenhove, 2009;Wittman, 2013), there are two predominant models used to describe the process of time perception; scalar expectancy theory, often called the pacemaker accumulator model, and the striatal beat frequency model (Allman and Meck, 2012;Allman et al., 2014). These models are used to highlight the effects of arousal (physiological or psychological) on the distortion of time (Lambourne, 2012;Jakubowski et al., 2015;Droit-Volet and Berthon, 2017). ...
... If after some time, the signal acquires some added significance (i.e., feedback, changes in the environment) then the contents of the accumulator are transferred from working memory to reference memory for long-term storage (Allman and Meck, 2012); this entails the memory stage. The decision stage occurs when the duration is experienced again on a separate trial, a ratio-decision rule compares if the current contents of the accumulator fall below, meet or exceed a threshold of similarity (Grondin, 2010;Allman and Meck, 2012;Allman et al., 2014). This process creates FIGURE 1 | The processing of information inside of the hypothesized internal clock as it relates to time perception as described by the scalar expectancy theory. ...
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The concept of time whether considered through the lenses of physics or physiology is a relative measure. Alterations in time perception can have serious implications in sport, fitness and work. Accurate perception of time is an important skill with many time constrained sports (i.e., basketball, North American football, tennis, gymnastics, figure skating, ice hockey, and others), and work environments (i.e., workers who need to synchronize their actions such as police and military). In addition, time distortions may play a role in exercise adherence. Individuals may be disinclined to continue with healthy, exercise activities that seem protracted (time dilation). Two predominant theories (scalar expectancy theory and striatal beat frequency model) emphasize the perception of the number of events in a period and the role of neurotransmitters in activating and coordinating cortical structures, respectively. A number of factors including age, sex, body temperature, state of health and fitness, mental concentration and exercise intensity level have been examined for their effect on time perception. However, with the importance of time perception for work, sport and exercise, there is limited research on this area. Since work, sports, and exercise can involve an integration of many of these aforementioned factors, they are interventions that need further investigations. The multiplicity of variables involved with work, sport, and exercise offer an underdeveloped but fruitful field for future research. Thus, the objective of this review was to examine physiological and psychological factors affecting human perception of time and the mechanisms underlying time perception and distortion with activity.
... With differences between the timing and information from the environment, the frontoparietal connection makes corrective adjustments in the next timing sequence, but individuals with amino acid sequence change from valine to methionine due to the BDNF Val66Met polymorphism, promote deficiencies in timing adjustment, and thus distort the time judgment (Wiener, Lohoff, and Coslett 2011;Marinho et al. 2018b;Lamb et al. 2015). This system may help to understand the genetic influence on temporal perception and relate our hypothesis to CNS structures which are involved as timers in time perception (Merchant, Harrington, and Meck 2013;Allman et al. 2014). ...
... In this case, it has been observed the BDNF level modulates the principle of pulse accumulation, reactions to stimuli, and motor and cognitive activities performance which are dependent on the neural excitation level and synaptic plasticity (Droit-Volet 2013; Wheeler et al. 2017). Besides, several neurological conditions caused by BDNF deficiency (Depression, Schizophrenia, and Bipolar Disorder) seem to modify the internal clock speed and soften the association with physiological and psychological aspects in the timing of stimuli (Allman et al. 2014;Wheeler et al. 2017;Lee et al. 2016). ...
... In addition, Chang et al. (2009) have shown the Met allele may be associated with the reduced severity of the schizophrenia negative symptoms. Notably, these findings support the hypothesis of perceptual ability performance, as the affected neurobiological domains are essences on the adjustment of internal clock speed demonstrated by the scalar expectancy theory and intrinsic models (Merchant, Harrington, and Meck 2013;Allman et al. 2014;Ivry and Spencer 2004). Together, these findings help us to conjecture the possibility of the BDNF Val66Met polymorphism exert an important influence on the neuronal integrity and risk of diseases that affect the time judgment. ...
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Studies at the molecular level aim to integrate genetic and neurobiological data to provide an increasingly detailed understanding of phenotypes related to the synchronization ability and brain oscillations in time perception. Genetic variation as a modifying factor at cellular and neurochemical levels permeates several neurofunctional aspects in time-lapse duration concentrating from milliseconds to hours. Thus, the review presents the BDNF Val66Met polymorphism association in a dynamic frame of brain neurotrophic factor expression in the adaptation, integrity, and neuronal synchronism processes in the ability to estimate multisensory stimuli at different time intervals. Our study aims to understand the molecular aspects involved in a neurobiological domain pertinent to the time judgment, tracing a genetic profile of association with psychometric functions and behavioral performances related to timing stimuli.
... Continuation tapping was notably less accurate for rhythms slower than 1Hz (supra-478 second rhythms) in both groups, in line with the scalar property of timing ( Allman et al. 2014;479 Gámez et al. 2019), however this was more prominent in ADHD. Previous studies have found 480 similar hastening in ADHD during continuation tapping for supra-second rhythms (Gilden and 481 Marusich 2009;Zelaznik et al. 2012), which has been attributed to a tendency to under-estimate 482 supra-second time intervals. ...
... Consistent with previous findings, our results indicate a difficultly in maintaining accurate 599 continuation tapping for supra-second rates (Allman et al. 2014). However, this was not 600 necessarily linked to individual participants' SMT/PPT. ...
... this preprint (which was not peer-reviewed) is the author/funder, who has granted bioRxiv a license to display the preprint in The copyright holder for . http://dx.doi.org/10.1101/2019.12.24.887802 doi: bioRxiv preprint first posted online Dec. 27, 2019; flexibility to sychronize to faster, but not to slower rhythms, has been previously reported even 604 in young infants (Bobin-Bègue et al. 2006), suggeting it is linked to the more general scalar 605 property of timing, predicting underestimation of supra-second intervals, rathar than specific 606 rhythmic preferences per se ( Allman et al. 2014;Gámez et al. 2019). Supporting this, when 607 continuation-tapping was inaccurate -primarily for supra-second intervals -this was not 608 systematically linked to how far the rhythm was from individual SMT/PPT. ...
Preprint
Many aspects of human behavior are inherently rhythmic, requiring production of rhythmic motor actions as well as synchronizing to rhythms in the environment. It is well-established that individuals with ADHD exhibit deficits in temporal estimation and timing functions, which may impact their ability to accurately produce and interact with rhythmic stimuli. In the current study we seek to understand the specific aspects of rhythmic behavior that are implicated in ADHD. We specifically ask whether they are attributed to imprecision in the internal generation of rhythms or to reduced acuity in rhythm perception. We also test key predictions of the Preferred Period Hypothesis, which suggests that both perceptual and motor rhythmic behaviors are biased towards a specific personal default tempo. To this end, we tested a several aspects of rhythmic behavior, including spontaneous motor tapping (SMT), perceptual preferences (PPT) and synchronization-continuations tapping in a broad range of rhythms, from sub-second to supra-second rates. Moreover, we evaluate the intra-subject consistency of rhythmic preferences, as a means for testing the reality and reliability of personal default-rhythms. Results indicate that individuals with ADHD are primarily challenged in producing self-generating isochronous motor rhythms, during both spontaneous and memory-paced tapping. However, they nonetheless exhibit a high degree of flexibility in synchronizing to a broad range of external rhythms. These findings suggest that auditory-motor entrainment is preserved in ADHD, and that the presence of an external pacer allows overcoming the inherent difficulty in self-generating isochronous motor rhythms. Interestingly, we find no advantage for performance near so-called default motor or perceptual rhythms, in either ADHD or control groups, as was suggested by the Preferred Period Hypothesis. Moreover, participants in both groups displayed large variability in their SMTs and PPTs across session, raising questions regarding the extent to which all individuals indeed have specific motor and perceptual preferences. Therefore, alongside the insights into the nature of rhythmic deficits in ADHD, this study also challenges some assumptions made previously regarding the prevalence and functional role of default rhythmic preferences in facilitating rhythmic behavior more broadly.
... MSNs are trained over successive trials by synaptic plasticity mechanisms to function as detectors of unique patterns of input that are related to specific target durations paired with reinforcement ( Dallérac et al., 2017). A major strength of the model is that it accounts for the scalar property which is the hallmark of interval timing (Allman et al., 2014;Yin et al., 2017). The first component of the scalar property requires that the mean measures of the timed behavior vary linearly, and usually accurately, with imposed temporal standards (i.e., target durations). ...
... As a consequence, the emerging view in the field is that although the timing processes governed by cortical-striatal circuits are distinct from the timing processes governed by corticocerebellar circuits, there is considerable room for integration of behavioral, systems, cellular, and molecular mechanisms. As a consequence, the proposed ICAT model assumes that these circuits work in synchrony, and contribute to distinct components of virtually all timing tasks (see Allman et al., 2014;Ohmae et al., 2017;Bareš et al., 2019;Caligiore et al., 2019). In summary, as described above, cerebellar PCs are capable of maintaining a temporal code through the time-specific pauses that occur in their millisecond spike patterns. ...
... A similar process, if verified in striatal MSNs would allow for changes in the speed of temporal integration and the coincidence detection of specific target durations in the seconds-to-minutes range (Figure 3). The proposal is that an intrinsic cellular mechanism based on microtubule dynamics (in both cerebellar and striatal ''time cells'') encodes the relevant temporal information (e.g., target duration and response thresholds) and can possibly be more effective as a read-write memory system than a circuit-based system with anatomically distinct temporal processing stages (e.g., clock, memory, and decision) as outlined by Gallistel and King (2010), Allman et al. (2014) and van . In accordance with the SBF model of interval timing, dopamine-dependent LTP leads to the encoding of both sub and supra-second target durations. ...
Article
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The majority of studies in the field of timing and time perception have generally focused on sub- and supra-second time scales, specific behavioral processes, and/or discrete neuronal circuits. In an attempt to find common elements of interval timing from a broader perspective, we review the literature and highlight the need for cell and molecular studies that can delineate the neural mechanisms underlying temporal processing. Moreover, given the recent attention to the function of microtubule proteins and their potential contributions to learning and memory consolidation/re-consolidation, we propose that these proteins play key roles in coding temporal information in cerebellar Purkinje cells (PCs) and striatal medium spiny neurons (MSNs). The presence of microtubules at relevant neuronal sites, as well as their adaptability, dynamic structure, and longevity, makes them a suitable candidate for neural plasticity at both intra- and inter-cellular levels. As a consequence, microtubules appear capable of maintaining a temporal code or engram and thereby regulate the firing patterns of PCs and MSNs known to be involved in interval timing. This proposed mechanism would control the storage of temporal information triggered by postsynaptic activation of mGluR7. This, in turn, leads to alterations in microtubule dynamics through a “read-write” memory process involving alterations in microtubule dynamics and their hexagonal lattice structures involved in the molecular basis of temporal memory.
... While working memory stores the current amount of pulses generated by the pacemaker, reference memory stores the earlier amount of pulses that have been learnt per unit of time. In the decision phase, pulses in the working and reference memories are compared to decide whether they correspond to the same time interval [74]. The internal clock theory offers an explanation about how animals learn a duration in a fixed-time interval operant conditioning procedure (FI) [75], where an animal learns to press a button In a classical duration comparison task, the agent is asked to decide which stimulus is longer or shorter (t 1 and t 2 ). ...
... Thus, it gives priority to the prospective estimation of time. Finally, the localization of the internal clock in the brain is still a matter of debate (for candidate brain areas, refer [74]). Internal clock theory, despite its limitations, supports an intuitive mechanism. ...
Article
Animals exploit time to survive in the world. Temporal information is required for higher-level cognitive abilities such as planning, decision making, communication, and effective cooperation. Since time is an inseparable part of cognition, there is a growing interest in the artificial intelligence approach to subjective time, which has a possibility of advancing the field. The current survey study aims to provide researchers with an interdisciplinary perspective on time perception. Firstly, we introduce a brief background from the psychology and neuroscience literature, covering the characteristics and models of time perception and related abilities. Secondly, we summarize the emergent computational and robotic models of time perception. A general overview to the literature reveals that a substantial amount of timing models are based on a dedicated time processing like the emergence of a clock-like mechanism from the neural network dynamics and reveal a relationship between the embodiment and time perception. We also notice that most models of timing are developed for either sensory timing (i.e. ability to assess an interval) or motor timing (i.e. ability to reproduce an interval). The number of timing models capable of retrospective timing, which is the ability to track time without paying attention, is insufficient. In this light, we discuss the possible research directions to promote interdisciplinary collaboration in the field of time perception.
... In physics (Layzer, 1975) (Veneziano, 1999) (Lebowitz, 1993) (Parrondo, Van den Broeck, & Kawai, 2009) and psychology (Allman, Teki, Griffiths, & Meck, 2014) (Ghaderi, Moradkhani, Haghighatfard, Akrami, Khayyer, & Balcı, 2018) (Ivry & Schlerf, 2008) (Meck W. H., 2006) (Meck W., 2014), a vast literature has been devoted to describing these two timing systems. However, just a few studies have addressed the relationship between subjective and physical time (Buonomano D. , 2017) (Buzsáki & Llinás, 2017) (Ghaderi, Moradkhani, Haghighatfard, Akrami, Khayyer, & Balcı, 2018). ...
... Previous studies in physics have shown that physical time (objective time) is a vector parameter with direction and magnitude (Layzer, 1975) (Stephani, 2004). However, a considerable number of studies in psychology predicted a modulated biological clock in the brain (modulated models) that can create time with a cumulative property as a scalar (Meck W., 2014) (Allman, Teki, Griffiths, & Meck, 2014) (Ivry & Schlerf, 2008). According to these descriptions, internal and external time systems have completely different natures which raises the concern of finding a logical relationship between subjective and objective time. ...
... In physics [18,28,19,26] and psychology [1,12,16,21,20], a vast literature has been devoted to describing these two timing systems. However, just a few studies have addressed the relationship between subjective and physical time [3,2,12]. ...
... Previous studies in physics have shown that physical time (objective time) is a vector parameter with direction and magnitude [17,27]. However, a considerable number of studies in psychology predicted a modulated biological clock in the brain (modulated models) that can create time with a cumulative property as a scalar [20,1,16]. According to these descriptions, internal and external time systems have completely different natures which raises the concern of finding a logical relationship between subjective and objective time. ...
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The neural basis of demanding mathematical problem solving is currently indeterminate and unclear. Mathematical problem solving engages higher order cognition and a complex associative activity of functional neural networks occurs during demanding problem solving. Twenty right handed subjects (mean age: 24.6 years; SD =3.97 years; 50% female) participated in this study. An arithmetic logic puzzle was used as a demanding mathematical task. EEGs were recorded in the eye open rest and eye open task conditions. To clarify functional connectivity of brain networks, clustering coefficient, transitivity, global efficiency, degree and entropy were investigated in two conditions. During problem solving, disrupted brain connectivity and decreased brain segregation were observed in the alpha band. However, in the beta band, increased connectivity, transitivity and clustering associated with higher modularity were observed. Theta exhibited unaltered brain network function. In the demanding problem solving task, decreased local alpha coupling may suggest that default mode network activity is interrupted. Since there is no significant difference within the theta network, the central executive network may not be as strongly involved. Increased segregation of functional brain network (without increasing of integration level) can be discussed in relation of demanding aspects of mathematical problem. We suggest a complex network may involve in the real situation of demanding problem solving.
... Although several studies in the recent decades have been conducted on the effects of emotions on individual's perception of time (29), no study was found on the effects of parents' feelings on children's understanding of time and the results of this disorientation in the formation of the psychological structure of children. However, studies have shown that the internal clock of each person can be specific and affect his psychological function (30) and several neural systems are involved in regulating this internal clock, including globus pallidus external, globus pallidus internal, supplementary motor area, substantial nigra pars compacta, ventral tegmental area, hippocampal, cortex (30). However, the following questions still remain unanswered: What is the relationship between parents' personality disorder and time per-ception disorder in children? ...
... Although several studies in the recent decades have been conducted on the effects of emotions on individual's perception of time (29), no study was found on the effects of parents' feelings on children's understanding of time and the results of this disorientation in the formation of the psychological structure of children. However, studies have shown that the internal clock of each person can be specific and affect his psychological function (30) and several neural systems are involved in regulating this internal clock, including globus pallidus external, globus pallidus internal, supplementary motor area, substantial nigra pars compacta, ventral tegmental area, hippocampal, cortex (30). However, the following questions still remain unanswered: What is the relationship between parents' personality disorder and time per-ception disorder in children? ...
... While working memory stores the current amount of pulses generated by the pacemaker, reference memory stores the earlier amount of pulses that have been learnt per unit of time. In the decision phase, pulses in the working and reference memories are compared to decide whether they correspond to the same time interval [74]. The internal clock theory offers an explanation about how animals learn a duration in a fixed-time interval operant conditioning procedure (FI) [75], where an animal learns to press a button In a classical duration comparison task, the agent is asked to decide which stimulus is longer or shorter (t 1 and t 2 ). ...
... Thus, it gives priority to the prospective estimation of time. Finally, the localization of the internal clock in the brain is still a matter of debate (for candidate brain areas, refer [74]). Internal clock theory, despite its limitations, supports an intuitive mechanism. ...
Preprint
Animals exploit time to survive in the world. Temporal information is required for higher-level cognitive abilities such as planning, decision making, communication and effective cooperation. Since time is an inseparable part of cognition, there is a growing interest in artificial intelligence to time, which has a possibility of advancing the field. This study aims to provide researchers with an interdisciplinary perspective on time. Firstly, we briefly discussed the necessary information from psychology and neuroscience, such as characteristics and models of time perception and related abilities. Secondly, we investigated the emergent computational and robotic models of time perception. As a result of the review, we observed that most timing models showed a sign of dedicated time processing like the emergence of clock-like mechanism from the neural network dynamics and revealed a relationship between embodiment and time perception. We also noticed that most models of timing developed for either sensory timing, the ability of assessment of an interval, or motor timing, ability to reproduce an interval. Additionally, the number of timing models capable of retrospective timing, which is the ability to track time without paying attention, is insufficient. In this light, we discussed possible research directions to promote interdisciplinary collaboration for time perception.
... The central timing mechanism, also known as the dedicated clock model (e.g., Allman et al., 2014), stemmed from Treisman's (1963) work. Decades of research into the human timing mechanism were based on this model and have assumed that timing is a specific cognitive module, hypothetically located across the global neural network (e.g., Allman & Meck, 2012). ...
Article
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The current review addresses two internal clock models that have dominated discussions in timing research for the last decades. More specifically, it discusses whether the central or the intrinsic clock model better describes the fluctuations in subjective time. Identifying the timing mechanism is critical to explain and predict timing behaviours in various audiovisual contexts. Music stands out for its prominence in real life scenarios along with its great potential to alter subjective time. An emphasis on how music as a complex dynamic auditory signal affects timing accuracy led us to examine the behavioural and neuropsychological evidence that supports either clock model. In addition to the timing mechanisms, an overview of internal and external variables, such as attention and emotions as well as the classic experimental paradigms is provided, in order to examine how the mechanisms function in response to changes occurring particularly during music experiences. Neither model can explain the effects of music on subjective timing entirely: The intrinsic model applies primarily to subsecond timing, whereas the central model applies to the suprasecond range. In order to explain time experiences in music, one has to consider the target intervals as well as the contextual factors mentioned above. Further research is needed to reconcile the gap between theories, and suggestions for future empirical studies are outlined.
... This design enabled to address the possibility of internal timing or duration being explicitly available to awareness through self-referential metacognition (Block, 1995;. Because current neuroscientific models posit that internal dynamics in timing are mediated by oscillatory or statedependent network dynamics (Laje & Buonomano, 2013;Karmarkar & Buonomano, 2007;Buhusi & Meck, 2005;Merchant et al., 2013;Allman et al., 2014;Gu et al., 2015;van Wassenhove, 2016;Bueno et al., 2017), we combined magneto-and electroencephalography (M/EEG) to quantify the dynamics of oscillatory brain responses when participants engaged in self-generated and self-assessed timing. Using a temporal production task to assess metacognition provided several paradigmatic and conceptual benefits. ...
Preprint
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Metacognition, the ability to know about one’s thought process, is self-referential. Here, we combined psychophysics and time-resolved neuroimaging to explore metacognitive inference on the accuracy of a self-generated behavior. Human participants generated a time interval and evaluated the signed magnitude of their temporal production. We show that both self-generation and self-evaluation relied on the power of beta oscillations (β; 15−40 Hz) with increases in early β power predictive of increases in duration. We characterized the dynamics of β power in a low dimensional space (β state-space trajectories) as a function of timing and found that the more distinct trajectories, the more accurate metacognitive inferences were. These results suggest that β states instantiates an internal variable determining the fate of the timing network’s trajectory, possibly as release from inhibition. Altogether, our study describes oscillatory mechanisms for timing, suggesting that temporal metacognition relies on inferential processes of self-generated dynamics.
... The neural mechanisms by which we perceive and measure the passage of time largely remain a mystery [1][2][3][4][5][6][7] . For the past century researchers have developed behavioral paradigms to address questions such as: Is time in our brain represented linearly like distance, logarithmically like pitch, or qualitatively like color? ...
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How our brains measure the passage of time is still largely open for debate. One behavioral task commonly used to study how durations are perceived is the Temporal Bisection Task, in which subjects categorize time durations as either “short” or “long.” The duration equally likely to be categorized as short or long is known as the bisection point. It has been consistently demonstrated that for humans, the bisection point is near the arithmetic mean of the longest and shortest durations the subject was trained on. In contrast, for non-human subjects it has been consistently found near the geometric mean. This difference implies that humans may process or represent temporal durations differently than other species. Here we present a behavioral model that reconciles the differences by demonstrating that rats’ performance on this task is driven not only by their noisy estimates of duration, but also by the temporally-discounted value of future rewards. The model correctly predicts shifts in the bisection point induced by unequal rewards and explains otherwise-paradoxical psychometric reversals documented three decades ago. Furthermore, as predicted by the model, we found that modifying the Temporal Bisection Task to eliminate the temporally-discounted reward component shifted the rats’ bisection point from the geometric mean to the arithmetic mean, thus bringing the rat results into line with the human results. We therefore propose that humans and rats (and perhaps other non-human subjects as well) process temporal information similarly, and that the difference between them in the Temporal Bisection Task may be simply due to rats weighing temporal discounting of future rewards more strongly than humans.
... The idea that we time sensory signals via a single Bcentralised^and Bamodal^clock has dominated the field of temporal cognition for decades (Gibbon et al. 1984). However, alternative positions propose that we have multiple timing mechanisms 'distributed' across brain areas or circuits and that the engagement of each single mechanism depends on the psychophysical task, sensory modality, and lengths of time intervals (Allman et al. 2014;Finnerty et al. 2015;Ivry and Schlerf 2008;Mauk and Buonomano 2004;Merchant et al. 2013;van Rijn et al. 2014). ...
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Understanding the neural and cognitive mechanisms underlying time estimation remains a challenge. Transcranial electric stimulations, such as transcranial random noise stimulation (tRNS), are useful tools to interfere with brain activity and identifying brain areas involved in temporal processing. Here, the aim is to investigate the specific role of primary sensory cortices (either V1 or A1) in temporal processing and to further investigate if the stimulation acts on either perceived duration or temporal sensitivity. Forty-eight university students were included in the study. Twenty-four participants were stimulated over A1 and 24 participants were stimulated over V1. All participants performed a time bisection task, either in a visual or auditory modality, involving standard durations lasting 300 ms (short) and 900 ms (long). When tRNS was delivered over A1, an effect of stimulation was observed on perceived duration (temporal over-estimation) under random stimulation compared to sham in both visual and auditory modalities. When tRNS was delivered over V1, the effect of stimulation was observed only in the visual modality (temporal over-estimation). No effect of stimulation was observed on temporal sensitivity in any condition. Our results showed for the first time that tRNS acts modulating individual’s perceived duration, but not on temporal sensitivity.
... 26 Explicit models assume monitoring of signal lengths of discrete temporal units and comparison of preceding lengths to the average of those stored in memory. 38,39 Instead, implicit timing models propose that timing abilities are dependent on neural oscillations, which are coupled to external signals. 40 Behavioural responses to conspecific or experimentally gen- erated stimuli can be used to infer whether an organism's tim- ing mechanism is proepisodic or homoepisodic. ...
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Time is one crucial dimension conveying information in animal communication. Evolution has shaped animals’ nervous systems to produce signals with temporal properties fitting their socio-ecological niches. Many quantitative models of mechanisms underlying rhythmic behaviour exist, spanning insects, crustaceans, birds, amphibians, and mammals. However, these computational and mathematical models are often presented in isolation. Here, we provide an overview of the main mathematical models employed in the study of animal rhythmic communication among conspecifics. After presenting basic definitions and mathematical formalisms, we discuss each individual model. These computational models are then compared using simulated data to uncover similarities and key differences in the underlying mechanisms found across species. Our review of the empirical literature is admittedly limited. We stress the need of using comparative computer simulations – both before and after animal experiments – to better understand animal timing in interaction. We hope this article will serve as a potential first step towards a common computational framework to describe temporal interactions in animals, including humans.
... Thus, depending on the task used, stimuli with higher speeds could, for instance, speed up the pacemaker, thereby leading to longer perceived durations as a result of a higher number of pulses being registered per unit time in the accumulator ( Zakay and Block, 1997;Wearden, 1999). On the other hand, a similar stimulus may lead to inadvertent attentional lapses which may lead to some of the pulses not getting registered in the accumulator (e.g., Penney, 2003;Kars ˛ılar and Balcı, 2016), thereby leading to shorter perceived durations (for a review see Allman et al., 2014). Examples of increases in the pacemaker rate (in addition to those mentioned above) have been shown in response to fast click-trains presented before timing a duration (Penton-Voak et al., 1996), higher body temperature (Wearden and Penton-Voak, 1995), emotional stimuli (DroitVolet et al., 2004), auditory as opposed to visual timing stimuli (Wearden et al., 1998), physical activity/motion ( Sayalı et al., 2018), as well as those manifested in terms of drug effects (see Coull et al., 2011 for a review). ...
Article
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The physical properties of events are known to modulate perceived time. This study tested the effect of different quantitative (walking speed) and qualitative (walking-forward vs. walking-backward) features of observed motion on time perception in three complementary experiments. Participants were tested in the temporal discrimination (bisection) task, in which they were asked to categorize durations of walking animations as “short” or “long.” We predicted the faster observed walking to speed up temporal integration and thereby to shift the point of subjective equality leftward, and this effect to increase monotonically with increasing walking speed. To this end, we tested participants with two different ranges of walking speeds in Experiment 1 and 2 and observed a parametric effect of walking speed on perceived time irrespective of the direction of walking (forward vs. rewound forward walking). Experiment 3 contained a more plausible backward walking animation compared to the rewound walking animation used in Experiments 1 and 2 (as validated based on independent subjective ratings). The effect of walking-speed and the lack of the effect of walking direction on perceived time were replicated in Experiment 3. Our results suggest a strong link between the speed but not the direction of perceived biological motion and subjective time.
... Two major theoretical approaches, among several, have been suggested to account for the mechanisms behind human timing (Wing and Kristofferson, 1973a,b;Getty, 1975;Meck, 1996;Church, 1999;Grondin, 2001Grondin, , 2010Mauk and Buonomano, 2004;Karmarkar and Buonomano, 2007;Ivry and Schlerf, 2008;Allman et al., 2014;Merker, 2014). The most influential and empirically tested psychoacoustic model is the "scalar expectancy theory" (Wearden, 1991;Allman and Meck, 2011). ...
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One curious aspect of human timing is the organization of rhythmic patterns in small integer ratios. Behavioral and neural research has shown that adjacent time intervals in rhythms tend to be perceived and reproduced as approximate fractions of small numbers (e.g., 3/2). Recent work on iterated learning and reproduction further supports this: given a randomly timed drum pattern to reproduce, participants subconsciously transform it toward small integer ratios. The mechanisms accounting for this “attractor” phenomenon are little understood, but might be explained by combining two theoretical frameworks from psychophysics. The scalar expectancy theory describes time interval perception and reproduction in terms of Weber's law: just detectable durational differences equal a constant fraction of the reference duration. The notion of categorical perception emphasizes the tendency to perceive time intervals in categories, i.e., “short” vs. “long.” In this piece, we put forward the hypothesis that the integer-ratio bias in rhythm perception and production might arise from the interaction of the scalar property of timing with the categorical perception of time intervals, and that neurally it can plausibly be related to oscillatory activity. We support our integrative approach with mathematical derivations to formalize assumptions and provide testable predictions. We present equations to calculate durational ratios by: (i) parameterizing the relationship between durational categories, (ii) assuming a scalar timing constant, and (iii) specifying one (of K) category of ratios. Our derivations provide the basis for future computational, behavioral, and neurophysiological work to test our model.
... This result implicates serotonin in suprasecond human interval timing (see also Rammsayer 1989;Wackermann et al. 2008), potentially through 5-HT 2A -mediated inhibition of dopamine (De Gregorio et al. 2016), which is believed to play an important mechanistic role in the perception of time (Allman and Meck 2012;Coull et al. 2011;Matell and Meck 2004;Rammsayer 1999;Soares et al. 2016;Terhune et al. 2016b;Vatakis and Allman 2015;Wiener et al. 2011) (for a review, see Coull et al. 2011). Given the role of interval timing across a range of psychological functions (Allman et al. 2014;Matthews and Meck 2016;Merchant et al. 2013), distorted timing under LSD may contribute to, or underlie, broader cognitive and perceptual effects of this drug. Therefore, elucidating its impact on interval timing is likely to inform neurochemical models of interval timing as well as our broader understanding of the effects of LSD on cognition and perception. ...
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Rationale Previous research demonstrating that lysergic acid diethylamide (LSD) produces alterations in time perception has implications for its impact on conscious states and a range of psychological functions that necessitate precise interval timing. However, interpretation of this research is hindered by methodological limitations and an inability to dissociate direct neurochemical effects on interval timing from indirect effects attributable to altered states of consciousness. Methods We conducted a randomised, double-blind, placebo-controlled study contrasting oral administration of placebo with three microdoses of LSD (5, 10, and 20 μg) in older adults. Subjective drug effects were regularly recorded and interval timing was assessed using a temporal reproduction task spanning subsecond and suprasecond intervals. Results LSD conditions were not associated with any robust changes in self-report indices of perception, mentation, or concentration. LSD reliably producedover-reproduction of temporal intervals of 2000msand longerwiththese effects mostpronounced in the 10μgdosecondition.Hierarchical regression analyses indicated that LSD-mediated over-reproduction was independent of marginal differences in self-reported drug effects across conditions. Conclusions These results suggest that microdose LSD produces temporal dilation of suprasecond intervals in the absence of subjective alterations of consciousness
... This result implicates serotonin in suprasecond human interval timing (see also Rammsayer 1989;Wackermann et al. 2008), potentially through 5-HT 2A -mediated inhibition of dopamine (De Gregorio et al. 2016), which is believed to play an important mechanistic role in the perception of time (Allman and Meck 2012;Coull et al. 2011;Matell and Meck 2004;Rammsayer 1999;Soares et al. 2016;Terhune et al. 2016b;Vatakis and Allman 2015;Wiener et al. 2011) (for a review, see Coull et al. 2011). Given the role of interval timing across a range of psychological functions (Allman et al. 2014;Matthews and Meck 2016;Merchant et al. 2013), distorted timing under LSD may contribute to, or underlie, broader cognitive and perceptual effects of this drug. Therefore, elucidating its impact on interval timing is likely to inform neurochemical models of interval timing as well as our broader understanding of the effects of LSD on cognition and perception. ...
... This result implicates serotonin in suprasecond human interval timing (see also Rammsayer 1989;Wackermann et al. 2008), potentially through 5-HT 2A -mediated inhibition of dopamine (De Gregorio et al. 2016), which is believed to play an important mechanistic role in the perception of time (Allman and Meck 2012;Coull et al. 2011;Matell and Meck 2004;Rammsayer 1999;Soares et al. 2016;Terhune et al. 2016b;Vatakis and Allman 2015;Wiener et al. 2011) (for a review, see Coull et al. 2011). Given the role of interval timing across a range of psychological functions (Allman et al. 2014;Matthews and Meck 2016;Merchant et al. 2013), distorted timing under LSD may contribute to, or underlie, broader cognitive and perceptual effects of this drug. Therefore, elucidating its impact on interval timing is likely to inform neurochemical models of interval timing as well as our broader understanding of the effects of LSD on cognition and perception. ...
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Rationale Previous research demonstrating that lysergic acid diethylamide (LSD) produces alterations in time perception has implications for its impact on conscious states and a range of psychological functions that necessitate precise interval timing. However, interpretation of this research is hindered by methodological limitations and an inability to dissociate direct neurochemical effects on interval timing from indirect effects attributable to altered states of consciousness. Methods We conducted a randomised, double-blind, placebo-controlled study contrasting oral administration of placebo with three microdoses of LSD (5, 10, and 20 μg) in older adults. Subjective drug effects were regularly recorded and interval timing was assessed using a temporal reproduction task spanning subsecond and suprasecond intervals. Results LSD conditions were not associated with any robust changes in self-report indices of perception, mentation, or concentration. LSD reliably produced over-reproduction of temporal intervals of 2000 ms and longer with these effects most pronounced in the 10 μg dose condition. Hierarchical regression analyses indicated that LSD-mediated over-reproduction was independent of marginal differences in self-reported drug effects across conditions. Conclusions These results suggest that microdose LSD produces temporal dilation of suprasecond intervals in the absence of subjective alterations of consciousness.
... Considerando que los GB son parte del CETC, los cuales también se encuentran relacionados con distorsiones fisiopatológicas de la percepción del tiempo (para recientes revisiones ver Allman & Meck, 2011;Allman, Teki, Griffiths & Meck, 2014;Merchant, Harrington & Meck, 2013;Schwartze & Kotz, 2013), Kotz y Schmidt-Kassow (2015) propusieron que un déficit de procesamiento temporal generalizado puede afectar el procesamiento de señales lingüísticas (sintaxis) y no lingüísticas (patrones musicales y lingüísticos bajo el control de la atención (Kotz & Gunter, 2015;Kotz, Gunter & Wonneberger, 2005;Kotz & Schwartze, 2010;Kotz, Schwartze & Schmidt-Kassow, 2009). Estos déficits de procesamiento temporal descritos parecen afectar no solo el comportamiento perceptivo, sino también la conducta motora. ...
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La disminución de dopamina en los ganglios basales (GB) es la principal causa de la enfermedad de Parkinson (EP). Además de las disfunciones motrices, los individuos con EP presentan alteraciones cognitivas que incluyen déficits de control atencional y temporal, de percepción del ritmo de la música y del habla, y de procesamiento del lenguaje. El circuito estriado-tálamo-cortical (CETC), participa en la comprensión musical y del habla. Personas con EP presentan problemas en distinguir las estructuras rítmicas por verse afectado el funcionamiento de los GB y, por lo tanto, el control de su pulso interno en el CETC. Sin embargo, se ha encontrado que personas con EP pueden mejorar su sincronización temporal interna mediante el acompañamiento de señales auditivas externas temporalmente predecibles. Estudios argumentan que señales de este tipo pueden compensar la vía disfuncional CETC mediante una interface con el circuito cerebelo-tálamo-cortical (CCTC) el cual es sensible a la codificación de eventos temporales.
... For example, people experience time as dilated, or lengthened, when viewing emotionally engaging pictures (Effron et al., 2006;Droit-Volet et al., 2004). Several models of time perception have been proposed (for reviews, see Allman et al., 2014;Buhusi and Meck, 2005). Although the debate whether the perception of time relies on dedicated or distributed intrinsic neural brain systems remains open (Ivry and Schlerf, 2008;Buhusi and Meck, 2005), evidence indicates that the subjective sense of time is not a recent achievement in brain evolution. ...
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Time perception depends on an event's emotional relevance to the beholder; a subjective time dilation effect is associated with self-relevant, emotionally salient stimuli. Previous studies have revealed that oxytocin modulates the salience of social stimuli and attention to social cues. However, whether the oxytocin system is involved in human subjective time perception is unknown. The aim of the present study was to investigate whether increased oxytocin levels would induce a time dilation effect for self-relevant, positive social cues. In a double-blind, placebo-controlled, between-subject design, heterosexual men were administered intranasal oxytocin or placebo. After about 50 min, participants completed a time-bisection task in which they estimated lengths of exposure to happy female faces (self-relevant positive stimuli, based on sexual orientation), emotionally neutral and negative female faces (control), and happy, neutral, and negative male faces (control). Oxytocin induced a subjective time dilation effect for happy female faces and a time compression effect for happy male faces. Our results provide evidence that oxytocin influences time perception, a primary form of human subjectivity.
... One can consider several brain regions as candidates to supply this information to the amygdala. The striatum is one, shown to underlie and pace short timescales at hundreds of milliseconds 46 , but also to signal longer durations 47,48 , yet mostly in specific dedicated time tasks 35,49 . However, the striatum does not project directly to the BLA, yet the BLA does project to parts of the striatum 50,51 . ...
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Associative learning forms when there is temporal relationship between a stimulus and a reinforcer, yet the inter-trial-interval (ITI), which is usually much longer than the stimulus-reinforcer-interval, contributes to learning-rate and memory strength. The neural mechanisms that enable maintenance of time between trials remain unknown, and it is unclear if the amygdala can support time scales at the order of dozens of seconds. We show that the ITI indeed modulates rate and strength of aversive-learning, and that single-units in the primate amygdala and dorsal-anterior-cingulate-cortex signal confined periods within the ITI, strengthen this coding during acquisition of aversive-associations, and diminish during extinction. Additionally, pairs of amygdala-cingulate neurons synchronize during specific periods suggesting a shared circuit that maintains the long temporal gap. The results extend the known roles of this circuit and suggest a mechanism that maintains trial-structure and temporal-contingencies for learning.
... Thus, beta desynchronization (possibly in co-operation with lower frequencies) in passive auditory tasks may encode top-down predictive timing of rhythms and elevate gain in neural processing by constraining auditory processing (Arnal, 2012;Arnal & Giraud, 2012;Arnal, Wyart, & Giraud, 2011). This type of prediction might be expected when listening to music (Allman, Teki, Griffiths, & Meck, 2014) or speech (Skipper, 2014;Skipper, Devlin, & Lametti, 2017;Skipper, Nusbaum, & Small, 2005). Similar interpretations related to prediction have been made when beta suppression is recorded in action-observation studies (Avenanti, Annela, & Serino, 2012;Press, Cook, Blakemore, & Kilner, 2011;Schippers & Keysers, 2011;Schutz-Bosbach & Prinz, 2007;Urgesi et al., 2010;Wilson & Knoblich, 2005). ...
... By comparison, learning of the temporal dimension has been less investigated. Mainly, time processing has been assessed either by the sequential order of events (Fortin et al., 2002;Kesner et al., 2002), or by time intervals (the duration of a segment of time) (Allman et al, 2014;Church et al., 1976;Holder & Roberts, 1985; for a review see Buhusi & Meck, 2005). In a Pavlovian task combining reinforcement time with locations and reinforcement probabilities, a recent study indicates that mice could learn to associate time intervals and locations in order to optimize their behaviour (Tosun et al., 2016). ...
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Time and space are commonly approached as two distinct dimensions, and rarely combined together in a single task, preventing a comparison of their interaction. In this project, using a version of a timing task with a spatial component, we investigate the learning of a spatio-temporal rule in animals. To do so, rats were placed in front of a five-hole nose-poke wall in a Peak Interval (PI) procedure to obtain a reward, with two spatio-temporal combination rules associated with different to-be-timed cues and lighting contexts. We report that, after successful learning of the discriminative task, a single Pavlovian session was sufficient for the animals to learn a new spatio-temporal association. This was seen as evidence for a beneficial transfer to the new spatio-temporal rule, as compared to control animals that did not experience the new spatio-temporal association during the Pavlovian session. The benefit was observed until nine days later. The results are discussed within the framework of adaptation to a change of a complex associative rule involving interval timing processes.
... En otra revisión, Allman et al. 40 reportaron un resumen de los recientes trabajos en neuroimagen que estudian la percepción del tiempo en el modelo animal y en seres humanos. Los resultados también corroboran la implicación de un circuito fronto-estriatal, que sería la base del funcionamiento del reloj interno. ...
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There are three purposes to this theoretical review. First, to present the main cognitive models of time perception based on the idea of an internal clock, as a core mechanism for making temporal judgments. Secondly, presenting the studies of cognitive neuroscience that provide evidence for cortical and subcortical structures involved this models. Specifically, we will focus on the role of the supplementary motor area (SMA) as an accumulator. Finally, we propose this model as a new framework to better understand the deficit in temporal processing in patients diagnosed with attention deficit and hyperactivity disorder. RESUMEN Este artículo de revisión teórica tiene tres objetivos. Primero, presentar una revi-sión de los principales modelos cognitivos de la percepción del tiempo basados en la idea de la existencia de un reloj interno como mecanismo necesario para la es-timación de juicios temporales. Segundo, presentar los estudios de neurociencias cognitivas que aportan evidencia de las estructuras corticales y subcorticales que participan en el funcionamiento de este reloj. En particular, el rol del área moto-ra suplementaria (AMS) en la acumulación del tiempo. Finalmente, proponer el modelo del reloj interno, como una alternativa para una mejor comprensión del déficit en el procesamiento temporal en pacientes diagnosticados con déficit por trastorno de atención con hiperactividad (TDAH). Key words: model of the internal model clock, time perception, temporal information processing, cognitive neuroscience, attention deficit and hyperactivity disorder. Palabras claves: Modelo del reloj interno, percepción del tiempo, procesamiento temporal de la información, neurociencias cognitivas, trastorno de atención e hiperactividad.
... This research traces the role of the cerebellum from projection neurons of the cerebellar cortex, through the dentate nucleus, and into thalamocortical-striatal circuits [104,105]. As a consequence, the cerebellum may not be limited to timing sub-second durations but may in fact play an important role in the timing of discrete intervals in the millisecond-to-minutes range [15,106]. ...
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Time perception is an essential element of conscious and subconscious experience, coordinating our perception and interaction with the surrounding environment. In recent years, major technological advances in the field of neuroscience have helped foster new insights into the processing of temporal information, including extending our knowledge of the role of the cerebellum as one of the key nodes in the brain for this function. This consensus paper provides a state-of-the-art picture from the experts in the field of the cerebellar research on a variety of crucial issues related to temporal processing, drawing on recent anatomical, neurophysiological, behavioral, and clinical research. The cerebellar granular layer appears especially well-suited for timing operations required to confer millisecond precision for cerebellar computations. This may be most evident in the manner the cerebellum controls the duration of the timing of agonist-antagonist EMG bursts associated with fast goal-directed voluntary movements. In concert with adaptive processes, interactions within the cerebellar cortex are sufficient to support sub-second timing. However, supra-second timing seems to require cortical and basal ganglia networks, perhaps operating in concert with cerebellum. Additionally, sensory information such as an unexpected stimulus can be forwarded to the cerebellum via the climbing fiber system, providing a temporally constrained mechanism to adjust ongoing behavior and modify future processing. Patients with cerebellar disorders exhibit impairments on a range of tasks that require precise timing, and recent evidence suggest that timing problems observed in other neurological conditions such as Parkinson’s disease, essential tremor, and dystonia may reflect disrupted interactions between the basal ganglia and cerebellum. The complex concepts emerging from this consensus paper should provide a foundation for further discussion, helping identify basic research questions required to understand how the brain represents and utilizes time, as well as delineating ways in which this knowledge can help improve the lives of those with neurological conditions that disrupt this most elemental sense. The panel of experts agrees that timing control in the brain is a complex concept in whom cerebellar circuitry is deeply involved. The concept of a timing machine has now expanded to clinical disorders.
... Moreover, qualitative or quantitative genetic changes promote underestimation or overestimation of time according to the Scalar Expectancy Theory (SET) [5,13,14]. This is accounted by the interference in the number of oscillations captured per time unit from the internal clock [15,16], and judgments of time intervals may result from changes in the pulse flow from an internal pacemaker in the presence of an event [17]. Consequently, genetic polymorphisms increase or decrease the speed of the internal clock, modulating the There was neither an association between the 5HTTLPR genotype and cognitive tasks, but there might be a tendency for better performance of SL as compared with SS carriers for fEP [24]. ...
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Background: Studies at the molecular level aim to integrate genetic and neurobiological data to provide an increasingly detailed understanding of phenotypes related to the ability in time perception. Main text: This study suggests that the polymorphisms genetic SLC6A4 5-HTTLPR, 5HTR2A T102C, DRD2/ANKK1-Taq1A, SLC6A3 3'-UTR VNTR, COMT Val158Met, CLOCK genes and GABRB2 A/C as modification factor at neurochemical levels associated with several neurofunctional aspects, modifying the circadian rhythm and built-in cognitive functions in the timing. We conducted a literature review with 102 studies that met inclusion criteria to synthesize findings on genetic polymorphisms and their influence on the timing. Conclusion: The findings suggest an association of genetic polymorphisms on behavioral aspects related in timing. However, order to confirm the paradigm of association in the timing as a function of the molecular level, still need to be addressed future research.
... First, attentional and arousal accounts of the OE have been suggested based on the pacemaker-accumulator model, which assumes the existence of a specialized internal clock mechanism for timing processes. The pacemaker-accumulator model is the dominant paradigm in the timing literature (Gibbon and Church, 1984;Gibbon, 1991;Church, 2003;Buhusi and Meck, 2005), mainly because timing ability shares many characteristic hallmarks with sensory perception, such as scalar properties (Wearden and Lejeune, 2008;Allman et al., 2014). The second major explanatory theory is the predictive coding account, centered on the neural coding efficiency framework. ...
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Perception of time is susceptible to distortions; among other factors, it has been suggested that the perceived duration of a stimulus is affected by the observer’s expectations. It has been hypothesized that the duration of an oddball stimulus is overestimated because it is unexpected, whereas repeated stimuli have a shorter perceived duration because they are expected. However, recent findings suggest instead that fulfilled expectations about a stimulus elicit an increase in perceived duration, and that the oddball effect occurs because the oddball is a target stimulus, not because it is unexpected. Therefore, it has been suggested that top-down attention is sometimes sufficient to explain this effect, and sometimes only necessary, with an additional contribution from saliency. However, how the expectedness of a target stimulus and its salient features affect its perceived duration is still an open question. In the present study, participants’ expectations about and the saliency of target stimuli were orthogonally manipulated with stimuli presented on a short (Experiment 1) or long (Experiment 2) temporal scale. Four repetitive standard stimuli preceded each target stimulus in a task in which participants judged whether the target was longer or shorter in duration than the standards. Engagement of top-down attention to target stimuli increased their perceived duration to the same extent irrespective of their expectedness. A small but significant additional contribution to this effect from the saliency of target stimuli was dependent on the temporal scale of stimulus presentation. In Experiment 1, saliency only significantly increased perceived duration in the case of expected target stimuli. In contrast, in Experiment 2, saliency exerted a significant effect on the overestimation elicited by unexpected target stimuli, but the contribution of this variable was eliminated in the case of expected target stimuli. These findings point to top-down attention as the primary cognitive mechanism underlying the perceptual extraction and processing of task-relevant information, which may be strongly correlated with perceived duration. Furthermore, the scalar properties of timing were observed, favoring the pacemaker-accumulator model of timing as the underlying timing mechanism.
... Although converging evidence has demonstrated crossmodal interaction and the transfer of information between different quantitative dimensions (see Alards-Tomalin, Leboe-McGowan, Shaw, & Leboe-McGowan, 2014), informationprocessing models aimed at explaining these interactions fall short of formally accounting for their representational overlaps, at both the neural and cognitive levels (but see Meck & Church, 1983;Meck et al., 1985). The most prominent of such models of temporal processing employs a pacemaker-accumulator theoretical approach, with three stages: (1) a pacemakeraccumulator component, which generates and temporarily stores durations in the form of pulses; (2) a memory component, in which the total number of pulses from the previous component is stored permanently; and (3) a decision/comparison stage, in which a random sample from the memory is compared to the value currently stored in the accumulator (e.g., Bshorter/ longer response^; Gibbon, Church, & Meck, 1984; see Allman, Teki, Griffiths, & Meck, 2014, for a review). ...
Article
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The perception of quantities has been suggested to rely on shared, magnitude-based representational systems that preserve metric properties. As such, different quantifiable dimensions that can characterize any given stimulus (e.g., size, speed, or numerosity) have been shown to modulate the perceived duration of these stimuli—a finding that has been attributed to cross-modal interaction among the quantity representations. However, these results are typically based on the isolated effects of a single stimulus dimension, leaving their potential combined effects uncharted. In the present study we aimed to investigate the joint effects of numerical magnitude and physical size on perceived time. In four complementary experiments, participants categorized six durations as “short” or “long,” which were presented through combinations of Hindu–Arabic numerals in three font sizes, as well as with simple shapes (rectangles) and unfamiliar symbols (Klingon letters), the sizes of which corresponded to the font sizes of the Hindu–Arabic numerals. Our results showed temporal underestimation for the smallest numeral in the set (3), with no effects of font size on perceived duration. The perceived durations were longest for the physically smallest geometric stimuli (i.e., a rectangle), and the font size of symbol-like stimuli (i.e., Klingon letters) was not found to have an effect on perceived time. Finally, presenting only one numeral (6) instead of the rectangle once again eliminated the relationship between physical size and perceived time, suggesting an overshadowing of physical-size-based influences on temporal choice behavior, presumably by perceived symbolism. Our results point at the complex nature of the interaction between different magnitude representations.
... Time perception is referred to as the subjective experience of time, and is often quantified by perception of the duration of elapsed time of a past event [1], moreover, time perception is critical for survival, given the needs to conceptualise the temporal course of events in preparation and planning for further actions (e.g., decision-making) [2]. Thus, the time duration is subjectively perceived as shorter or longer than its actual passage due to dynamic interaction between personal experience and environment conditions [3,4]. ...
Article
Several studies have demonstrated that stroke subjects present impairment of functions related to decision-making and timing, involving the information processing in the neural circuits of the cerebellum in association with the prefrontal cortex. This review is aimed to identify the gaps, and demonstrate a better understanding of decision-making and timing functions in the patients with stroke. Electronic literature database was searched and the findings of relevant studies were used to explore the mechanisms of decision-making and timing in patients with stroke, as well as the circuit connections in timing mediated by prefrontal cortex and cerebellum. A literature review was conducted with 65 studies that synthesized findings on decision-making and time perception in individuals with stroke. Types of neurobiological modalities in this study included: Relationships among decision-making, time perception, related cognitive aspects (such as discrimination tasks, verbal estimation, bisection tasks, time production and motor reproduction), and motor control. We demonstrate that the timing processes are important for the performance in cognitive tasks and that the cerebellum and prefrontal cortex are involved in decision-making and time perception. In the context, the decision-making is impaired in stroke patients has a great impact on executive functions, and this seems to be important in determining neurobiological aspects relevant to the time interval interpretation.
... Oscillatory hierarchy (Pöppel 1972(Pöppel , 1997 or dynamic attending theory (Jones 1976;Large and Jones 1999 Timekeeping mechanisms also come with a number of first-and second-order principles (Allman et al. 2014): the first-order principles apply to durations and typically involve the accuracy and precision with which a duration is being timed. ...
... The memory stage is considered as a place for storing accumulated pulses in working memory for comparison with the content of reference memory. According to internal clock model of time perception (Meck 1983, Block, Zakay et al. 1998, Allman, Teki et al. 2014, time production task involves comparing duration experience with the duration stored in memory (reference memory). The reference memory is a long-term memory which represents number of pulses accumulated in previous experience. ...
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A growing body of research suggests that space, time and number are represented within a common system. Other studies have shown this relationship is related to the mathematical competency. Here we examined the influence of the mathematical capacities of 8-12 years old children, grouped into high (n=63) and low (n=58) on performance in mental number line, time knowledge and time perception. The results revealed that mathematical competency influences mental number line and time knowledge, but with regard to time perception the effects were only observed in time production task. In addition, the results of correlation analysis revealed interaction between time knowledge, time production (but not reproduction) and mental number line. Finally, the findings are discussed within the framework of the recent theories regarding representation of space, time and number.
... Il englobe généralement différentes capacités qui se réfèrent à l'utilisation d'informations temporelles telles que les durées 1 L'entraînement est entendu ici dans son acception neurophysiologique : un stimulus périodique (i.e., répétitif) est perçu par un ensemble de neurones spécifiquement dédiés à cette fonction, lesquels génèrent une activité électrique décelable par des techniques d'enregistrement électroencéphalographique. Des travaux récents font état de cette propriété remarquable d'entraînement neuronal à des stimuli auditifs, qui serait à l'origine de la perception de patterns rythmiques externes et de la capacité du système perceptivo-moteur à se synchroniser aux propriétés métriques des stimuli externes (Fujioka et al., 2012 ;Nozaradan, 2014). ou le rythme (Allman, Teki, & Griffiths, 2014;Repp & Su, 2013;Schirmer, Meck & Penney, 2016). Dans ce cadre, il s'agit donc de regrouper sous le terme timing des capacités de traitement perceptif d'occurrences temporelles périodiques ou isolées, qu'elle qu'en soit la nature (e.g., visuelle, auditive, tactile), et des possibilités de réponse à ces évènements temporels (e.g., se synchroniser à des stimuli rythmiques, émettre un jugement sur la durée d'un stimulus, etc.). ...
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Au-delà des symptômes cardinaux qui caractérisent la maladie de Parkinson (MP) – tremblement, akinésie, et rigidité – des déficits rythmiques se manifestent dans différents domaines de coordination motrice, comme au niveau du membre supérieur, de la sphère oro-faciale, ou de la marche. Des altérations rythmiques sont également mises en évidence sur des tâches de perception de rythme (i.e., sur des tâches n’impliquant pas de production motrice). Face à l’étendue des dysfonctionnements rythmiques dans la MP, l’hypothèse d’une dysrythmie généralisée a été formulée. Cette hypothèse implique que l’ensemble des altérations rythmiques qui s’observent au travers de diverses tâches et dans différents systèmes effecteurs partage des mécanismes causaux communs. Néanmoins, cette proposition n’a pas été confirmée à ce jour, et nombre de questions demeurent, tant sur le plan théorique que clinique : les déficits rythmiques caractéristiques de la MP sont-ils réellement liés ? Une source commune aux manifestations rythmiques déficitaires est-elle envisageable ? Si tel est le cas, quels en sont les corrélats cérébraux, et les retombées cliniques ? Élaborée autour de deux principaux axes de recherche, cette dissertation avait pour objectif principal de tester l’hypothèse d’une dysrythmie généralisée dans la MP, au travers de deux questions : i) existe-t-il des liens entre trois domaines de production rythmique (i.e., coordinations oro-faciale, manuelle, et de marche) et un domaine perceptif dans la MP ?; et ii) quel est l’impact d’un entraînement rythmique d’un domaine moteur (i.e., coordination rythmique manuelle) sur d’autres domaines de coordination motrice (i.e., oro-faciales et de la marche) ? L’ensemble des résultats confirme l’hypothèse d’une dysrythmie généralisée dans la MP, et l’existence très probable d’altérations de mécanismes en lien avec une fonction prédictive générale qui, lorsqu’elle est la cible d’un entraînement rythmique, pourrait permettre de réduire certains troubles moteurs dans la MP.
... timing | embodiment | internal clock T he ability of animals to adapt their behavior to periodic events is critical for survival, as the appearance of a sensory cue can predict the timing of food availability, predator attack, or mating opportunity (1)(2)(3)(4). It has been postulated that humans and other animals use a dedicated internal clock to evaluate the duration of behaviorally relevant time intervals and sensory cues, or to produce well-timed movements (5)(6)(7)(8)(9)(10). However, time is a critical parameter for a wide range of behaviors engaging distinct brain regions. ...
Article
How animals adapt their behavior according to regular time intervals between events is not well understood, especially when intervals last several seconds. One possibility is that animals use disembodied internal neuronal representations of time to decide when to initiate a given action at the end of an interval. However, animals rarely remain immobile during time intervals but tend to perform stereotyped behaviors, raising the possibility that motor routines improve timing accuracy. To test this possibility, we used a task in which rats, freely moving on a motorized treadmill, could obtain a reward if they approached it after a fixed interval. Most animals took advantage of the treadmill length and its moving direction to develop, by trial-and-error, the same motor routine whose execution resulted in the precise timing of their reward approaches. Noticeably, when proficient animals did not follow this routine, their temporal accuracy decreased. Then, naïve animals were trained in modified versions of the task designed to prevent the development of this routine. Compared to rats trained in the first protocol, these animals didn’t reach a comparable level of timing accuracy. Altogether, our results indicate that timing accuracy in rats is improved when the environment affords cues that animals can incorporate into motor routines.
... M uch information that the brain processes and stores is temporal in nature. Therefore, understanding the processing of time in the brain is of fundamental importance in neuroscience (1)(2)(3)(4). To predict and maximize future rewards in this ever-changing world, animals must be able to discover the temporal structure of stimuli and then flexibly anticipate or act correctly at the right time. ...
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To maximize future rewards in this ever-changing world, animals must be able to discover the temporal structure of stimuli and then anticipate or act correctly at the right time. How do animals perceive, maintain, and use time intervals ranging from hundreds of milliseconds to multiseconds in working memory? How is temporal information processed concurrently with spatial information and decision making? Why are there strong neuronal temporal signals in tasks in which temporal information is not required? A systematic understanding of the underlying neural mechanisms is still lacking. Here, we addressed these problems using supervised training of recurrent neural network models. We revealed that neural networks perceive elapsed time through state evolution along stereotypical trajectory, maintain time intervals in working memory in the monotonic increase or decrease of the firing rates of interval-tuned neurons, and compare or produce time intervals by scaling state evolution speed. Temporal and nontemporal information is coded in subspaces orthogonal with each other, and the state trajectories with time at different nontemporal information are quasiparallel and isomorphic. Such coding geometry facilitates the decoding generalizability of temporal and nontemporal information across each other. The network structure exhibits multiple feedforward sequences that mutually excite or inhibit depending on whether their preferences of nontemporal information are similar or not. We identified four factors that facilitate strong temporal signals in nontiming tasks, including the anticipation of coming events. Our work discloses fundamental computational principles of temporal processing, and it is supported by and gives predictions to a number of experimental phenomena.
... The polymorphisms active in neurotransmission predisposed participants to different behavioral phenotypes, and consequently it is inferred to inadequate recruitment, decrease or increase in neurotransmitter levels, which causes changes in neural inputs during the time intervals timing [9,30,40]. This is reflected by the interference in the number of oscillations captured per unit of time from the internal clock [41,42], and thus, changes occur in the flow of pulses produced by an internal pacemaker in the presence of an event [10]. Consequently, the molecular bases increase or decrease the internal clock speed, and this modulates the neurotransmission of Review Juliete Bandeira pulses and reactions to stimuli that determine the synchronism in cognitive and motor actions through visual and/or auditory stimuli [6,24,30]. ...
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Research at the molecular level aims to integrate neurobiological information in a more detailed manner on behavioral phenotypes associated with the judgment of time intervals. The genetic polymorphisms SLC6A4 5-HTTLPR and 5HTR2A T102C as modulators of serotoninergic levels are associated with several neurobiological aspects inbuilt in sub-second and supra-second timing. In this sense, a state-of-the-art review was performed with 60 studies, among them experimental and reviews, to synthesize the main findings on the action of genetic bases of the serotoninergic system in timing. The findings indicate that the level of expression of serotonin receptors and transporters change neural inputs in neurobiological domains related to the time perception.
... One can consider several brain regions as candidates to supply this information to the amygdala. The striatum is one, shown to underlie and pace short time scales at hundreds of miliseconds 55 , but also to signal longer durations 56,57 , yet mostly in specific dedicated time tasks 39,58 . However, the striatum does not project directly to the BLA, yet the BLA does project to parts of the striatum [59][60][61] . ...
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Associative learning forms when there is temporal relationship between a stimulus and a reinforcer, yet the inter-trial-interval (ITI), which is usually much longer than the stimulus-reinforcer-interval, contributes to learning-rate and memory strength. The neural mechanisms that enable maintenance of time between trials remain unknown, and it is unclear if the amygdala can support time scales at the order of dozens of seconds. We show that the ITI indeed modulates rate and strength of aversive-learning, and that single-units in the primate amygdala and dorsal-anterior-cingulate-cortex signal confined periods within the ITI, strengthen this coding during acquisition of aversive-associations, and diminish during extinction. Additionally, pairs of amygdala-cingulate neurons synchronize during specific periods suggesting a shared circuit that maintains the long temporal gap. The results extend the known roles of this circuit and suggest a mechanism that maintains trial-structure and temporal-contingencies for learning. It further suggests a novel model for maladaptive behaviors.
... L'altération des processus temporels -ou capacités de timing -dans le TDA/H est une donnée qui s'est imposée progressivement depuis les années 1990 sur la base de nombreuses preuves expérimentales (voir [17] pour une revue). Le timing englobe généralement différentes dimensions qui se réfèrent à la capacité à percevoir des événements temporels, et à s'ajuster à eux sur le plan comportemental, de manière implicite ou explicite [18,19]. Le timing renvoie également à la capacité à considérer les conséquences futures de nos actes, et à anticiper ce qui pourrait découler de nos décisions actuelles [17]. ...
Article
Résumé But de l’étude L’objectif de cette étude était de proposer une prise en charge psychomotrice basée sur des mises en situation rythmique et de tester son impact sur les capacités attentionnelles et exécutives d’enfants TDA/H. Patients et méthode Vingt-et-un enfants âgés de 8 à 13 ans (moyenne : 9,4 ans ± 1,8 ans ; 3 filles) diagnostiqués avec un TDA/H ont bénéficié d’une prise en charge psychomotrice (8 à 12 séances) axée sur des mises en situation rythmique (exercices de synchronisation à des rythmes externes de type métronome ou musiques). Un design test-retest a permis d’évaluer le niveau des fonctions attentionnelles et exécutives à l’aide de batteries d’évaluation neuropsychologiques avant et après l’intervention. Résultats Les performances attentionnelles et exécutives ont été améliorées après la mise en place du programme : capacités d’inhibition, maîtrise de l’impulsivité cognitive, et mémoire de travail visuo-spatiale. Le niveau de l’attention divisée s’est légèrement amélioré également, alors que l’attention soutenue auditive, l’attention sélective visuelle, et l’aversion au délai sont restées inchangées. Conclusion Cette étude établit des liens entre un entraînement des capacités rythmiques et l’amélioration de plusieurs fonctions attentionnelles et exécutives chez des enfants TDA/H. Ce type d’intervention psychomotrice apparaît donc prometteur dans l’accompagnement thérapeutique des enfants souffrant de TDA/H, en complément ou alternativement à un traitement par méthylphénidate chez certains sujets.
... Much information that the brain processes and stores is temporal in nature. Therefore, to understand the processing of time in the brain is of fundamental importance in neuroscience [1,2,3,4]. Working memory, the ability to maintain and manipulate information over a period of seconds, is a core cognitive function for planning and executing tasks [5,6]. In this study, we focused on the working memory of a basic temporal pattern: time interval. ...
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To predict and maximize future rewards in this ever-changing world, animals must be able to discover the temporal structure of stimuli and then take the right action at the right time. However, we still lack a systematic understanding of the neural mechanism of how animals perceive, maintain, and use time intervals ranging from hundreds of milliseconds to multi-seconds in working memory and appropriately combine time information with spatial information processing and decision making. Here, we addressed this problem by training neural network models on four timing tasks: interval production, interval comparison, timed spatial reproduction, and timed decision making. We studied time-coding principles of the network after training, and found them consistent with existing experimental observations. We reveal that neural networks perceive time intervals through the evolution of population state along a stereotypical trajectory, maintain time intervals by line attractors along which the activities of most neurons vary monotonically with the duration of the maintained interval, and adjust the evolution speed of the state trajectory for producing or comparing time intervals. Spatial information or decision choice preserves the profiles of neuronal activities as functions of time intervals maintained in working memory or flow of time, and is coded in the amplitudes of these profiles. Decision making is combined with time perception through two firing sequences with mutual inhibition. Our work discloses fundamental principles of the neuronal coding of time that supports the brain for flexible temporal processing. These principles facilitate generalizable decoding of time and non-time information.
... It has previously been assumed that the freely chosen tapping frequency is directly related to the frequency of the internal clock ). The internal clock is assumed to be located in the brain (Ivry & Keele, 1989), as described in a review paper on the subject (Allman, Teki, Griffiths, & Meck, 2014). However, it has been mentioned that it constitutes a challenge that knowledge about the location and about physiological details of the internal clock are sparse ( Baer et al., 2015). ...
Article
These hypotheses were tested: (1) Freely chosen frequency in unilateral index finger tapping is correlated between the two index fingers, and (2) A 3-min bout of unilateral index finger tapping followed by 10 min rest results in an increase of the freely chosen tapping frequency performed by the contralateral index finger in a second bout. Thirty-two adults participated. Freely chosen tapping frequencies from first bouts were 167.2 ± 79.0 and 161.5 ± 69.4 taps/min for the dominant and non-dominant hand, respectively (p=.434). These variables correlated (R=.86, p<.001). When bout one and two were performed with the dominant and non-dominant hand, respectively, the frequency increased by 8.1%±17.2% in bout two (p=.011). In opposite order, the frequency increased by 14.1%±17.5% (p<.001), which was not different from the ∼8% (p=.157).
... On another note, the human being's perception of time is crucial in humans for everyday activities. All stimuli and actions have temporal extent, and judgment of duration is essential for certain basic behaviors such as combining motor sequences to achieve a goal, and for more complex acts such as arriving on time for appointments, temporally structuring speech, or foreseeing future events (Allman, Teki, Griffiths, & Meck, 2014;Buhusi & Meck, 2005;Matthews & Meck, 2016). ...
Article
Prepotent response inhibition and temporal perception abilities were explored in a sample of individuals with cerebral palsy relative to typically developing peers. The extent to which inhibitory control difficulties might affect temporal processing was also investigated. For this purpose, two inhibitory control tasks and two duration estimation tasks were given to the groups of cerebral palsy and typically developing children. Results showed inhibition and temporal perception problems in the group with cerebral palsy. A relationship was found between inhibition and temporal estimation performances, which indicates that inhibitory control contributes, at least partially, to acquisition of the temporal processing ability.
... Firstly, time estimates increase linearly with a rise in the duration to be judged. Secondly, the variation of the estimate is proportional to the duration (Allman, Teki, Griffiths, & Meck, 2014). These two properties, and also the effect of emotion on time perception, are considered in the framework of the PA model (Gibbon et al., 1984;Treisman, 1963). ...
Article
Time sensitivity is affected by emotional stimuli such as fearful faces. The effect of threatening stimuli on time perception depends on numerous factors, including task type and duration range. We applied a two‐interval forced‐choice task using face stimuli to healthy volunteers to evaluate time perception and emotion interaction using functional magnetic resonance imaging. We conducted finite impulse response analysis to examine time series for the significantly activated brain areas and psychophysical interaction to investigate the connectivity between selected regions. Time perception engaged a right lateralised frontoparietal network, while a face discrimination task activated the amygdala and fusiform face area (FFA). No voxels were active with regards to the effect of expression (fearful versus neutral). In parallel with this, our behavioral results showed that attending to the fearful faces did not cause duration overestimation. Finally, connectivity of the amygdala and FFA to the middle frontal gyrus increased during the face processing condition compared to the timing task. Overall, our results suggest that the prefrontal‐amygdala connectivity might be required for the emotional processing of facial stimuli. On the other hand, attentional load, task type, and task difficulty are discussed as possible factors that influence the effects of emotion on time perception. This article is protected by copyright. All rights reserved.
... Oscillatory hierarchy (Pöppel 1972(Pöppel , 1997 or dynamic attending theory (Jones 1976;Large and Jones 1999 Timekeeping mechanisms also come with a number of first-and second-order principles (Allman et al. 2014): the first-order principles apply to durations and typically involve the accuracy and precision with which a duration is being timed. ...
... Critically, there was a dissociation within the iCNV signal between tasks where the amplitude for temporal estimates covaried with the length of the interval the subject was planning on reproducing, while no difference in the iCNV for spatial estimates was observed. A central framework of timing continues to be the pacemaker-accumulator model in which the brain contains a pacemaker whose pulses are integrated by an accumulator to process measures of time ( Allman et al., 2014 ). Included within this theory is the notion of climbing of neural activity over time that is associated with the SMA and is indexed by the CNV ( Coull et al., 2016 ). ...
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The perception and measurement of spatial and temporal dimensions have been widely studied. Yet, whether these two dimensions are processed independently is still being debated. Additionally, whether EEG components are uniquely associated with time or space, or whether they reflect a more general measure of magnitude quantity remains unknown. While undergoing EEG, subjects performed a virtual distance reproduction task, in which they were required to first walk forward for an unknown distance or time, and then reproduce that distance or time. Walking speed was varied between estimation and reproduction phases, to prevent interference between distance or time in each estimate. Behaviorally, subject performance was more variable when reproducing time than when reproducing distance, but with similar patterns of accuracy. During estimation, EEG data revealed the contingent negative variation (CNV), a measure previously associated with timing and expectation, tracked the probability of the upcoming interval, for both time and distance. However, during reproduction, the CNV exclusively oriented to the upcoming temporal interval at the start of reproduction, with no change across spatial distances. Our findings indicate that time and space are neurally separable dimensions, with the CNV both serving a supramodal role in temporal and spatial expectation, yet an exclusive role in preparing duration reproduction.
... Under this scenario, temporal processing is governed by MPC neural population clocks that switch from temporal scaling of their state dynamics during interval timing to amplitude modulation in their tangent circular trajectories during rhythmic timing. Importantly, because MPC is part of both the corticobasal ganglia and the cortico-cerebellar circuits, it can play an important role in both interval and rhythmic timing and can act as a synergistic context-dependent element within the two core timing systems, as suggested previously [46][47][48][49]. Evolving patterns of activation. A. Neural activation periods for the second produced interval (second and third taps as white vertical lines) during SC for the target interval of 850 ms. ...
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Our motor commands can be exquisitely timed according to the demands of the environment, and the ability to generate rhythms of different tempos is a hallmark of musical cognition. Yet, the neuronal underpinnings behind rhythmic tapping remain elusive. Here, we found that the activity of hundreds of primate medial premotor cortices (MPCs; pre-supplementary motor area [preSMA] and supplementary motor area [SMA]) neurons show a strong periodic pattern that becomes evident when their responses are projected into a state space using dimensionality reduction analysis. We show that different tapping tempos are encoded by circular trajectories that travelled at a constant speed but with different radii, and that this neuronal code is highly resilient to the number of participating neurons. Crucially, the changes in the amplitude of the oscillatory dynamics in neuronal state space are a signature of duration encoding during rhythmic timing, regardless of whether it is guided by an external metronome or is internally controlled and is not the result of repetitive motor commands. This dynamic state signal predicted the duration of the rhythmically produced intervals on a trial-by-trial basis. Furthermore, the increase in variability of the neural trajectories accounted for the scalar property, a hallmark feature of temporal processing across tasks and species. Finally, we found that the interval-dependent increments in the radius of periodic neural trajectories are the result of a larger number of neurons engaged in the production of longer intervals. Our results support the notion that rhythmic timing during tapping behaviors is encoded in the radial curvature of periodic MPC neural population trajectories.
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It has been proposed that animals rely on internal representation of time to measure supra-second long intervals and adapt their behavior accordingly. If this was the case, accurate timing should be largely independent of variations in external factors (e.g., changes in the environment). Here, we used a task in which rats, freely moving on a motorized treadmill, could obtain a reward if they approached it after a fixed interval. By manipulating several task parameters, such as the speed of the treadmill, we found that the animals accurately timed their approaches only if they could perform a stereotyped motor sequence inside the confined space of the treadmill. Reinforcement learning-based models further suggested that the strategy of the animals was incongruent with that of agents using an internal representation of time. We conclude that the emergence of well-timed behaviors critically depends on the ability of animals to develop motor routines adapted to their environment.
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We consider the task of measuring time with probabilistic threshold gates implemented by bio-inspired spiking neurons. In the model of spiking neural networks, network evolves in discrete rounds, where in each round, neurons fire in pulses in response to a sufficiently high membrane potential. This potential is induced by spikes from neighboring neurons that fired in the previous round, which can have either an excitatory or inhibitory effect. We first consider a deterministic implementation of a neural timer and show that $\Theta(\log t)$ (deterministic) threshold gates are both sufficient and necessary. This raised the question of whether randomness can be leveraged to reduce the number of neurons. We answer this question in the affirmative by considering neural timers with spiking neurons where the neuron $y$ is required to fire for $t$ consecutive rounds with probability at least $1-\delta$, and should stop firing after at most $2t$ rounds with probability $1-\delta$ for some input parameter $\delta \in (0,1)$. Our key result is a construction of a neural timer with $O(\log\log 1/\delta)$ spiking neurons. Interestingly, this construction uses only one spiking neuron, while the remaining neurons can be deterministic threshold gates. We complement this construction with a matching lower bound of $\Omega(\min\{\log\log 1/\delta, \log t\})$ neurons. This provides the first separation between deterministic and randomized constructions in the setting of spiking neural networks. Finally, we demonstrate the usefulness of compressed counting networks for synchronizing neural networks.
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Recent sensory history plays a critical role in the perception of event duration. For example, repetitive exposure to a particular duration leads to the distortion of subsequent duration perception. This phenomenon, termed duration adaptation, induces a robust repulsive duration aftereffect. In particular, adaptation to relatively long sensory events shortens the perceived duration of a subsequent event, while adaptation to relatively short sensory events lengthens the perception of subsequent event durations. This phenomenon implies the plasticity of duration perception and offers important clues for revealing the cognitive neural mechanism of duration perception. Duration aftereffect has received more and more attention in recent years. In this review, we introduce recent research advances in our understanding of duration aftereffect, especially with regards to its manifestations, origin, and cognitive neural mechanisms. We also propose possible directions for future research. In sum, we posit that studies on the duration aftereffect phenomenon are helpful in understanding general duration perception, and as such, should receive more attention in future.
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Measurement of perceived time to complete tasks may be influenced by the real time needed, subjective distortions, and by the measurement instruments used. To disentangle these sources the present study examined an Internet-based achievement task. Participants were asked to solve logical problems in sequence. After each problem, participants estimated their completion time. This estimation was either given on a Likert-type scale or visual analogue scale (VAS). To test for an expected learning effect, the difference between very similar problems was calculated for both scale types, both for objective reaction time and estimation of time. We further manipulated between subjects the fluency of the logical problem, i.e. the ease of processing of task relevant information. The sample consisted of 571 participants. Correlations between objective reaction time and subjective perception of speed for correctly solved items were moderate, mean r=.14. The associations were higher for VAS scales than Likert-type scales, r=.27 versus r=.19, respectively. The results for absolute changes in reaction times were mixed. When predicting only the direction of change, VASs did show a higher sensibility than Likert-type scales. Correct responses to logical problems in the fluent condition were more frequent (54%) than for the same problems in the non-fluent condition (42%), χ2(4)=22.54, p<.001. Objective reaction times did not differ between the conditions, but subjective estimates tended to be faster for fluent problems. In conclusion, findings suggest that subjective time estimations vary depending on the type of response scale used for the assessment as well as fluency of the task.
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A large number of competing models exist for how the brain creates a representation of time. However, several human and animal studies point to 'climbing neural activation' as a potential neural mechanism for the representation of duration. Neurophysiological recordings in animals have revealed how climbing neural activation that peaks at the end of a timed interval underlies the processing of duration, and, in humans, climbing neural activity in the insular cortex, which is associated with feeling states of the body and emotions, may be related to the cumulative representation of time.
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Contrary to data showing sensitivity to nontemporal properties of timed signals, current theories of interval timing assume that animals can use the presence or absence of a signal as equally valid cues as long as duration is the most predictive feature. Consequently, the authors examined rats' behavior when timing the absence of a visual or auditory stimulus in trace conditioning and in a "reversed" gap procedure. Memory for timing was tested by presenting the stimulus as a reversed gap into its timed absence. Results suggest that in trace conditioning (Experiment 1), rats time for the absence of a stimulus by using its offset as a time marker. As in the standard gap procedure, the insertion of a reversed gap was expected to "stop" rats' internal clock. In contrast, a reversed gap of 1-, 5-, or 15-s duration "reset" the timing process in both trace conditioning (Experiment 2) and the reversed gap procedure (Experiment 3). A direct comparison of the standard and reversed gap procedures (Experiment 4) supported these findings. Results suggest that attentional mechanisms involving the salience or content of the gap might contribute to the response rule adopted in a gap procedure.
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The past few decades have seen an explosion in studies exploring the effects of emotion on time 17 judgments. The aim of this review is to describe the results of these studies and to look at how they 18 try to explain the time distortions produced by emotion. We begin by examining the findings on time 19 judgments in affective disorders, which allow us to make a clear distinction between the feelings of time distortion that originate from introspection onto subjective personal experience, and the ef- 20 fects of emotion on the basic mechanisms involved in time perception. We then report the results of 21 behavioral studies that have tested the effects of emotions on time perceptions and the temporal pro- 22 cessing of different emotional stimuli (e.g. facial expressions, affective pictures or sounds). Finally, 23 we describe our own studies of the embodiment of timing. Overall, the different results on time and 24 emotion suggest that temporal distortions are an indicator of how our brain and body adapt to the 25 dynamic structure of our environment.
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The terms 'autism' and 'autistic' derive from the Greek word autos meaning self. This is appropriate to describing the autistic behavioral phenotype in which there is a pathological impairment in socialization and verbal and nonverbal communication, in addition to behavior and interests that are often highly restricted and repetitive (the triad; American Psychiatric Association, 1994). The autistic individual often appears isolated, and unable to make sense of the world around them. They often reveal an inability to predict and understand the behavior of others, and perceptions of the world remain fragmented and are not embedded into a coherent pattern or structure. Time is part of the fundamental intellectual structure in which we make sense of the events in our lives. 'Timing and time perception allow us to unite action sequences and events occurring separately in time, to adapt to
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Medical College of Wisconsin, Milwaukee, Wisconsin Weber’s law applied to interval timing is called thescalar property. A hallmark of timing in the secondsto-minutes range, the scalar property is characterized by proportionality between the standard deviation of a response distribution and the duration being timed. In this temporal reproduction study, we assessed whether the scalar property was upheld when participants chronometrically counted three visually presented durations (8, 16, and 24 sec) as compared with explicitly timing durations without counting. Accuracy for timing and accuracy for counting were similar. However, whereas timing variability showed the scalar property, counting variability did not. Counting variability across intervals was accurately modeled by summing a random variable representing an individual count. A second experiment replicated the first and demonstrated that task differences were not due to presentation order or practice effects. The distinct psychophysical properties of counting and timing behaviors argue for greater attention to participant strategies in timing studies.
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Time is embedded in any sensory experience: the movements of a dance, the rhythm of a piece of music, the words of a speaker are all examples of temporally structured sensory events. In humans, if and how visual cortices perform temporal processing remains unclear. Here we show that both primary visual cortex (V1) and extrastriate area V5/MT are causally involved in encoding and keeping time in memory and that this involvement is independent from low-level visual processing. Most importantly we demonstrate that V1 and V5/MT come into play simultaneously and seem to be functionally linked during interval encoding, whereas they operate serially (V1 followed by V5/MT) and seem to be independent while maintaining temporal information in working memory. These data help to refine our knowledge of the functional properties of human visual cortex, highlighting the contribution and the temporal dynamics of V1 and V5/MT in the processing of the temporal aspects of visual information.
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We examined the effect of temporal context on discrimination of intervals marked by auditory, visual and tactile stimuli. Subjects were asked to compare the duration of the interval immediately preceded by an irrelevant "distractor" stimulus with an interval with no distractor. For short interval durations, the presence of the distractor affected greatly the apparent duration of the test stimulus: short distractors caused the test interval to appear shorter and vice versa. For very short reference durations (≤100 ms), the contextual effects were large, changing perceived duration by up to a factor of two. The effect of distractors reduced steadily for longer reference durations, to zero effect for durations greater than 500 ms. We found similar results for intervals defined by visual flashes, auditory tones and brief finger vibrations, all falling to zero effect at 500 ms. Under appropriate conditions, there were strong cross-modal interactions, particularly from audition to vision. We also measured the Weber fractions for duration discrimination and showed that under the conditions of this experiment, Weber fractions decreased steadily with duration, following a square-root law, similarly for all three modalities. The magnitude of the effect of the distractors on apparent duration correlated well with Weber fraction, showing that when duration discrimination was relatively more precise, the context dependency was less. The results were well fit by a simple Bayesian model combining noisy estimates of duration with the action of a resonance-like mechanism that tended to regularize the sound sequence intervals.
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The brain's ability to tell time and produce complex spatiotemporal motor patterns is critical for anticipating the next ring of a telephone or playing a musical instrument. One class of models proposes that these abilities emerge from dynamically changing patterns of neural activity generated in recurrent neural networks. However, the relevant dynamic regimes of recurrent networks are highly sensitive to noise; that is, chaotic. We developed a firing rate model that tells time on the order of seconds and generates complex spatiotemporal patterns in the presence of high levels of noise. This is achieved through the tuning of the recurrent connections. The network operates in a dynamic regime that exhibits coexisting chaotic and locally stable trajectories. These stable patterns function as 'dynamic attractors' and provide a feature that is characteristic of biological systems: the ability to 'return' to the pattern being generated in the face of perturbations.
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Duration estimation is known to be far from veridical and to differ for sensory estimates and motor reproduction. To investigate how these differential estimates are integrated for estimating or reproducing a duration and to examine sensorimotor biases in duration comparison and reproduction tasks, we compared estimation biases and variances among three different duration estimation tasks: perceptual comparison, motor reproduction, and auditory reproduction (i.e. a combined perceptual-motor task). We found consistent overestimation in both motor and perceptual-motor auditory reproduction tasks, and the least overestimation in the comparison task. More interestingly, compared to pure motor reproduction, the overestimation bias was reduced in the auditory reproduction task, due to the additional reproduced auditory signal. We further manipulated the signal-to-noise ratio (SNR) in the feedback/comparison tones to examine the changes in estimation biases and variances. Considering perceptual and motor biases as two independent components, we applied the reliability-based model, which successfully predicted the biases in auditory reproduction. Our findings thus provide behavioral evidence of how the brain combines motor and perceptual information together to reduce duration estimation biases and improve estimation reliability.
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In most species, interval timing is time-scale invariant: errors in time estimation scale up linearly with the estimated duration. In mammals, time-scale invariance is ubiquitous over behavioral, lesion, and pharmacological manipulations. For example, dopaminergic drugs induce an immediate, whereas cholinergic drugs induce a gradual, scalar change in timing. Behavioral theories posit that time-scale invariance derives from particular computations, rules, or coding schemes. In contrast, we discuss a simple neural circuit, the perceptron, whose output neurons fire in a clockwise fashion (interval timing) based on the pattern of coincidental activation of its input neurons. We show numerically that time-scale invariance emerges spontaneously in a perceptron with realistic neurons, in the presence of noise. Under the assumption that dopaminergic drugs modulate the firing of input neurons, and that cholinergic drugs modulate the memory representation of the criterion time, we show that a perceptron with realistic neurons reproduces the pharmacological clock and memory patterns, and their time-scale invariance, in the presence of noise. These results suggest that rather than being a signature of higher-order cognitive processes or specific computations related to timing, time-scale invariance may spontaneously emerge in a massively-connected brain from the intrinsic noise of neurons and circuits, thus providing the simplest explanation for the ubiquity of scale invariance of interval timing.
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Accurate timing is a ubiquitous aspect of mental processes. How does the central nervous system solve the demands involved in the temporal aspects of information processing? One solution would be that timing is handled by subsystems specialized for domain-specific processing. For example, the timing required for producing wellarticulated speech would be solved by areas involved in speech production, whereas the timing demands for the coordination of manual actions would be controlled by brain areas that also control the force and spatial aspects of these movements. Alternatively, humans are capable of producing rather arbitrary behaviors that exhibit accurate timing. We can produce periodic movements over a considerable range of durations. These actions can be achieved with different parts of the body and, indeed, do not even require overt actions; we can covertly maintain an internal beat. We can also detect and judge rhythmicity in a wide variety of sensory signals. While our sense of rhythm may be most accurate for auditory events, we can readily detect temporal perturbations in a sequence of visual or tactile events. Thus, there likely exists some general system specialized to represent temporal information, a system that is recruited for tasks that require this form of computation.
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