Timing Research Forum

How does the dynamic shape of a sound affect its perceived microtiming? In the TIME project, we studied basic aspects of musical microrhythm, exploring both stimulus features and the participants’ enculturated expertise via perception experiments, observational studies of how musicians produce particular microrhythms, and ethnographic studies of musicians’ descriptions of microrhythm. Collectively, we show that altering the microstructure of a sound (“what” the sound is) changes its perceived temporal location (“when” it occurs). Specifically, there are systematic effects of core acoustic factors (duration, attack) on perceived timing. Microrhythmic features in longer and more complex sounds can also give rise to different perceptions of the same sound. Our results shed light on conflicting results regarding the effect of microtiming on the “grooviness” of a rhythm.

Filmmakers and editors have empirically developed techniques to ensure the spatiotemporal continuity of a film's narration. In terms of time, editing techniques (e.g., elliptical, overlapping, or cut minimization) allow for the manipulation of the perceived duration of events as they unfold on screen. More specifically, a scene can be edited to be time compressed, expanded, or real-time in terms of its perceived duration. Despite the consistent application of these techniques in filmmaking, their perceptual outcomes have not been experimentally validated. Given that viewing a film is experienced as a precise simulation of the physical world, the use of cinematic material to examine aspects of time perception allows for experimentation with high ecological validity, while filmmakers gain more insight on how empirically developed techniques influence viewers' time percept. Here, we investigated how such time manipulation techniques of an action affect a scene's perceived duration. Specifically, we presented videos depicting different actions (e.g., a woman talking on the phone), edited according to the techniques applied for temporal manipulation and asked participants to make verbal estimations of the presented scenes' perceived durations. Analysis of data revealed that the duration of expanded scenes was significantly overestimated as compared to that of compressed and real-time scenes, as was the duration of real-time scenes as compared to that of compressed scenes. Therefore, our results validate the empirical techniques applied for the modulation of a scene's perceived duration. We also found interactions on time estimates of scene type and editing technique as a function of the characteristics and the action of the scene presented. Thus, these findings add to the discussion that the content and characteristics of a scene, along with the editing technique applied, can also modulate perceived duration. Our findings are discussed by considering current timing frameworks, as well as attentional saliency algorithms measuring the visual saliency of the presented stimuli.

William James’s use of “time in passing” and “stream of thoughts” may be two sides of the same coin that emerge from the brain segmenting the continuous flow of information into discrete events. Departing from that idea, we investigated how the content of a realistic scene impacts two distinct temporal experiences: the felt duration and the speed of the passage of time. I will present you the results from an online study in which we used a well-established experimental paradigm, the temporal bisection task, which we extended to passage of time judgments. 164 participants classified seconds-long videos of naturalistic scenes as short or long (duration), or slow or fast (passage of time). Videos contained a varying number and type of events. We found that a large number of events lengthened subjective duration and accelerated the felt passage of time. Surprisingly, participants were also faster at estimating their felt passage of time compared to duration. The perception of duration heavily depended on objective duration, whereas the felt passage of time scaled with the rate of change. Altogether, our results support a possible dissociation of the mechanisms underlying the two temporal experiences.

In the real world, object arrangement follows a number of rules. Some of the rules pertain to the spatial relations between objects and scenes (i.e., syntactic rules) and others about the contextual relations (i.e., semantic rules). Research has shown that violation of semantic rules influences interval timing with the duration of scenes containing such violations to be overestimated as compared to scenes with no violations. However, no study has yet investigated whether both semantic and syntactic violations can affect timing in the same way. Furthermore, it is unclear whether the effect of scene violations on timing is due to attentional or other cognitive accounts. Using an oddball paradigm and real-world scenes with or without semantic and syntactic violations, we conducted two experiments on whether time dilation will be obtained in the presence of any type of scene violation and the role of attention in any such effect. Our results from Experiment 1 showed that time dilation indeed occurred in the presence of syntactic violations, while time compression was observed for semantic violations. In Experiment 2, we further investigated whether these estimations were driven by attentional accounts, by utilizing a contrast manipulation of the target objects. The results showed that an increased contrast led to duration overestimation for both semantic and syntactic oddballs. Together, our results indicate that scene violations differentially affect timing due to violation processing differences and, moreover, their effect on timing seems to be sensitive to attentional manipulations such as target contrast.

In the current virtual journal club Dr Di Luca will present findings from a series of psychophysical investigations where he measured sensitivity and bias in the perception of the timing of stimuli. He will present how improved detection with longer sequences and biases in reporting isochrony can be accounted for by optimal statistical predictions. Among his findings was also that the timing of stimuli that occasionally deviate from a regularly paced sequence is perceptually distorted to appear more regular. Such change depends on whether the context these sequences are presented is also regular. Dr Di Luca will present a Bayesian model for the combination of dynamically updated expectations, in the form of a priori probability, with incoming sensory information. These findings contribute to the understanding of how the brain processes temporal information to shape perceptual experiences.

Neural entrainment has become a phenomenon of exceptional interest to neuroscience, given its involvement in rhythm perception, production, and overt synchronized behavior. Yet, traditional methods fail to quantify neural entrainment due to a misalignment with its fundamental definition (e.g., see Novembre and Iannetti, 2018; Rajandran and Schupp, 2019). The definition of entrainment assumes that endogenous oscillatory brain activity undergoes dynamic frequency adjustments to synchronize with environmental rhythms (Lakatos et al., 2019). Following this definition, we recently developed a method sensitive to this process. Our aim was to isolate from the electroencephalographic (EEG) signal an oscillatory component that is attuned to the frequency of a rhythmic stimulation, hypothesizing that the oscillation would adaptively speed up and slow down to achieve stable synchronization over time. To induce and measure these adaptive changes in a controlled fashion, we developed the event-related frequency adjustment (ERFA) paradigm (Rosso et al., 2023). A total of twenty healthy participants took part in our study. They were instructed to tap their finger synchronously with an isochronous auditory metronome, which was unpredictably perturbed by phase-shifts and tempo-changes in both positive and negative directions across different experimental conditions. EEG was recorded during the task, and ERFA responses were quantified as changes in instantaneous frequency of the entrained component. Our results indicate that ERFAs track the stimulus dynamics in accordance with the perturbation type and direction, preferentially for a sensorimotor component. The clear and consistent patterns confirm that our method is sensitive to the process of frequency adjustment that defines neural entrainment. In this Virtual Journal Club, the discussion of our findings will be complemented by methodological insights beneficial to researchers in the fields of rhythm perception and production, as well as timing in general. We discuss the dos and don’ts of using instantaneous frequency to quantify oscillatory dynamics, the advantages of adopting a multivariate approach to source separation, the robustness against the confounder of responses evoked by periodic stimulation, and provide an overview of domains and concrete examples where the methodological framework can be applied.

Subsecond timing underlies nearly all sensory and motor activities across species and is critical to survival. While subsecond temporal information has been found across cortical and subcortical regions, it is unclear if it is generated locally and intrinsically or if it is a read out of a centralized clock-like mechanism. Indeed, mechanisms of subsecond timing at the circuit level are largely obscure. Primary sensory areas are well-suited to address these question as they have early access to sensory information and provide minimal processing to it: if temporal information is found in these regions, it is likely to be generated intrinsically and locally. We test this hypothesis by training mice to perform an audio-visual temporal pattern sensory discrimination task as we use 2-photon calcium imaging, a technique capable of recording population level activity at single cell resolution, to record activity in primary visual cortex (V1). We have found significant changes in network dynamics through mice’s learning of the task from naive to middle to expert levels. Changes in network dynamics and behavioral performance are well accounted for by an intrinsic model of timing in which the trajectory of q network through high dimensional state space represents temporal sensory information. Conversely, while we found evidence of other temporal encoding models, such as oscillatory activity, we did not find that they accounted for increased performance but were in fact correlated with the intrinsic model itself. These results provide insight into how subsecond temporal information is encoded mechanistically at the circuit level.

To form a coherent perception of the world around us, we are constantly processing and integrating sensory information from multiple modalities. In fact, when auditory and visual stimuli occur within ~100 ms of each other, individuals tend to perceive the stimuli as a single event, even though they occurred separately. In recent years, our lab, and others, have developed rat models of audiovisual temporal perception using behavioural tasks such as temporal order judgments (TOJs) and synchrony judgments (SJs). While these rodent models demonstrate metrics that are consistent with humans (e.g., perceived simultaneity, temporal acuity), we have sought to confirm whether rodents demonstrate the hallmarks of audiovisual temporal perception, such as predictable shifts in their perception based on experience and sensitivity to alterations in neurochemistry. Ultimately, our findings indicate that rats serve as an excellent model to study the neural mechanisms underlying audiovisual temporal perception, which to date remains relativity unknown. Using our validated translational audiovisual behavioural tasks, in combination with optogenetics, neuropharmacology and in vivo electrophysiology, we aim to uncover the mechanisms by which inhibitory neurotransmission and top-down circuits finely control ones’ perception. This research will significantly advance our understanding of the neuronal circuitry underlying audiovisual temporal perception, and will be the first to establish the role of interneurons in regulating the synchronized neural activity that is thought to contribute to the precise binding of audiovisual stimuli.

Waiting is one of healthcare’s core experiences. It is there in the time it takes to access services; through the days, weeks, months or years needed for diagnoses; in the time that treatment takes; and in the elongated time-frames of recovery, relapse, remission and dying.Funded by the Wellcome Trust, our project opens up what it means to wait in and for healthcare by examining lived experiences, representations and histories of delayed and impeded time.In an era in which time is lived at increasingly different and complex tempos, Waiting Times looks to understand both the difficulties and vital significance of waiting for practices of care, offering a fundamental re-conceptualisation of the relation between time and care in contemporary thinking about health, illness, and wellbeing.

When listening to music, humans readily perceive and move along with a periodic beat. Critically, perception of a periodic beat is commonly elicited by rhythmic stimuli with physical features arranged in a way that is not strictly periodic. Hence, beat perception must capitalize on mechanisms that transform stimulus features into a temporally recurrent format with emphasized beat periodicity. Here, I will present a line of work that aims to clarify the nature and neural basis of this transformation. In these studies, electrophysiological activity was recorded as participants listened to rhythms known to induce perception of a consistent beat across healthy Western adults. The results show that the human brain selectively emphasizes beat representation when it is not acoustically prominent in the stimulus, and this transformation (i) can be captured non-invasively using surface EEG in adult participants, (ii) is already in place in 5- to 6-month-old infants, and (iii) cannot be fully explained by subcortical auditory nonlinearities. Moreover, as revealed by human intracerebral recordings, a prominent beat representation emerges already in the primary auditory cortex. Finally, electrophysiological recordings from the auditory cortex of a rhesus monkey show a significant enhancement of beat periodicities in this area, similar to humans. Taken together, these findings indicate an early, general auditory cortical stage of processing by which rhythmic inputs are rendered more temporally recurrent than they are in reality. Already present in non-human primates and human infants, this "periodized" default format could then be shaped by higher-level associative sensory-motor areas and guide movement in individuals with strongly coupled auditory and motor systems. Together, this highlights the multiplicity of neural processes supporting coordinated musical behaviors widely observed across human cultures.The experiments herein include: a motor timing task comparing the effects of movement vs non-movement with and without feedback (Exp. 1A & 1B), a transcranial magnetic stimulation (TMS) study on the role of the supplementary motor area (SMA) in transforming temporal information (Exp. 2), and a perceptual timing task investigating the effect of noisy movement on time perception with both visual and auditory modalities (Exp. 3A & 3B). Together, the results of these studies support the Bayesian cue combination framework, in that: movement improves the precision of time perception not only in perceptual timing tasks but also motor timing tasks (Exp. 1A & 1B), stimulating the SMA appears to disrupt the transformation of temporal information (Exp. 2), and when movement becomes unreliable or noisy there is no longer an improvement in precision of time perception (Exp. 3A & 3B). Although there is support for the proposed framework, more studies (i.e., fMRI, TMS, EEG, etc.) need to be conducted in order to better understand where and how this may be instantiated in the brain; however, this work provides a starting point to better understanding the intrinsic connection between time and movement

Mobile organisms must be capable of deciding both where and when to move in order to keep up with a changing environment; therefore, a strong sense of time is necessary, otherwise, we would fail in many of our movement goals. Despite this intrinsic link between movement and timing, only recently has research begun to investigate the interaction. Two primary effects that have been observed include: movements biasing time estimates (i.e., affecting accuracy) as well as making time estimates more precise. The goal of this presentation is to review this literature, discuss a Bayesian cue combination framework to explain these effects, and discuss the experiments I have conducted to test the framework. The experiments herein include: a motor timing task comparing the effects of movement vs non-movement with and without feedback (Exp. 1A & 1B), a transcranial magnetic stimulation (TMS) study on the role of the supplementary motor area (SMA) in transforming temporal information (Exp. 2), and a perceptual timing task investigating the effect of noisy movement on time perception with both visual and auditory modalities (Exp. 3A & 3B). Together, the results of these studies support the Bayesian cue combination framework, in that: movement improves the precision of time perception not only in perceptual timing tasks but also motor timing tasks (Exp. 1A & 1B), stimulating the SMA appears to disrupt the transformation of temporal information (Exp. 2), and when movement becomes unreliable or noisy there is no longer an improvement in precision of time perception (Exp. 3A & 3B). Although there is support for the proposed framework, more studies (i.e., fMRI, TMS, EEG, etc.) need to be conducted in order to better understand where and how this may be instantiated in the brain; however, this work provides a starting point to better understanding the intrinsic connection between time and movement

Any sensory experience of the world, from the touch of a caress to the smile on our friend’s face, is embedded in time and it is often associated with the perception of the flow of it. The perception of time is therefore a peculiar sensory experience built without dedicated sensors. How the perception of time and the content of a sensory experience interact to give rise to this unique percept is unclear. A few empirical evidences show the existence of this interaction, for example the speed of a moving object or the number of items displayed on a computer screen can bias the perceived duration of those objects. However, to what extent the coding of time is embedded within the coding of the stimulus itself, is sustained by the activity of the same or distinct neural populations and subserved by similar or distinct neural mechanisms is far from clear. Addressing these puzzles represents a way to gain insight on the mechanism(s) through which the brain represents the passage of time. In my talk I will present behavioral and neuroimaging studies to show how concurrent changes of visual stimulus duration, speed, visual contrast and numerosity, shape and modulate brain’s and pupil’s responses and, in case of numerosity and time, influence the topographic organization of these features along the cortical visual hierarchy.

Metacognition has long been defined as “cognition about cognition”. One of its aspects is the error monitoring ability, which includes being aware of one’s own errors without external feedback. This ability is mostly investigated in two-alternative forced choice tasks, where the performance has all or none nature in terms of accuracy. The previous literature documents the effect of different influences on the error monitoring ability, such as working memory, feedback and sensorimotor involvement. However, these demonstrations fall short of generalizing to the real life scenarios where the errors often have a magnitude and a direction. To bridge this gap, recent studies showed that humans could keep track of the magnitude and the direction of their errors in temporal, spatial and numerical domains in two metrics: confidence and short-long/few-more judgements. This talk will cover how the documented effects that are obtained in the two alternative forced choices tasks apply to the temporal error monitoring ability. Finally, how magnitude and direction monitoring (i.e., confidence and short-long judgements) can be differentiated as the two indices of temporal error monitoring ability will be discussed.

Precisely estimating event timing is essential for survival, yet temporal distortions are ubiquitous in our daily sensory experience. Here, we tested whether the relative position, duration, and distance in time of two sequentially-organized events—standard S, with constant duration, and comparison C, with duration varying trial-by-trial—are causal factors in generating temporal distortions. We found that temporal distortions emerge when the first event is shorter than the second event. Importantly, a significant interaction suggests that a longer inter-stimulus interval (ISI) helps to counteract such serial distortion effect only when the constant S is in the first position, but not if the unpredictable C is in the first position. These results imply the existence of a perceptual bias in perceiving ordered event durations, mechanistically contributing to distortion in time perception. Our results clarify the mechanisms generating time distortions by identifying a hitherto unknown duration-dependent encoding inefficiency in human serial temporal perception, something akin to a strong prior that can be overridden for highly predictable sensory events but unfolds for unpredictable ones.

Since Darwin, comparative research has shown that most animals share basic timing capacities, such as the ability to process temporal regularities and produce rhythmic behaviors. What seems to be more exclusive, however, are the capacities to generate temporal predictions and to display anticipatory behavior at salient time points. These abilities are associated with subcortical structures like basal ganglia (BG) and cerebellum (CE), which are more developed in humans as compared to nonhuman animals. In the first research line, we investigated the basic capacities to extract temporal regularities from the acoustic environment and produce temporal predictions. We did so by adopting a comparative and translational approach, thus making use of a unique EEG dataset including 2 macaque monkeys, 20 healthy young, 11 healthy old participants and 22 stroke patients, 11 with focal lesions in the BG and 11 in the CE. In the second research line, we holistically explore the functional relevance of body-brain physiological interactions in human behavior. Thus, a series of planned studies investigate the functional mechanisms by which body signals (e.g., respiratory and cardiac rhythms) interact with and modulate neurocognitive functions from rest and sleep states to action and perception. This project supports the effort towards individual profiling: are individuals’ timing capacities (e.g., rhythm perception and production), and general behavior (e.g., individual walking and speaking rates) influenced / shaped by body-brain interactions?

Decades of research have investigated the relationship between perception of time and heart rate with often mixed results. In search of such a relationship, I will present my far journey between two projects: from time perception in the realistic VR experience of crowded subway trips in the order of minutes (project 1); to the perceived duration of sub-second white noise tones (project 2). Heart rate had multiple concurrent relationships with subjective temporal distortions for the sub-second tones, while the effects were lacking or weak for the supra-minute subway trips. What does the heart have to do with sub-second time perception? We addressed this question with a cardiac drift-diffusion model, demonstrating the sensory accumulation of temporal evidence as a function of heart rate.

Temporal predictions are fundamental instruments for facilitating sensory selection, allowing humans to exploit regularities in the world. Recent evidence indicates that the motor system instantiates predictive timing mechanisms, helping to synchronize temporal fluctuations of attention with the timing of events in a task-relevant stream, thus facilitating sensory selection. Accordingly, in the auditory domain auditory-motor interactions are observed during perception of speech and music, two temporally structured sensory streams. I will present a behavioral and neurophysiological account for this theory and will detail the parameters governing the emergence of this auditory-motor coupling, through a set of behavioral and magnetoencephalography (MEG) experiments.

We recorded and analyzed the population activity of hundreds of neurons in the medial premotor areas (MPC) of rhesus monkeys performing an isochronous tapping task guided by brief flashing stimuli or auditory tones. The animals showed a strong bias towards visual metronomes, with rhythmic tapping that was more precise and accurate than for auditory metronomes. The population dynamics in state space as well as the corresponding neural sequences shared the following properties across modalities: the circular dynamics of the neural trajectories and the neural sequences formed a regenerating loop for every produced interval, producing a relative time representation; the trajectories converged in similar state space at tapping times while the moving bumps restart at this point, resetting the beat-based clock; the tempo of the synchronized tapping was encoded by a combination of amplitude modulation and temporal scaling in the neural trajectories. In addition, the modality induced a displacement in the neural trajectories in auditory and visual subspaces without greatly altering time keeping mechanism. These results suggest that the interaction between the amodal internal representation of pulse within MPC and a modality specific external input generates a neural rhythmic clock whose dynamics define the temporal execution of tapping using auditory and visual metronomes.

Our perception of time isn’t like a clock; it varies depending on other aspects of experience, such as what we see and hear in that moment. However, in everyday life, the properties of these simple features can change frequently, presenting a challenge to understanding real-world time perception based on simple lab experiments. We developed a computational model of human time perception based on tracking changes in neural activity across brain regions involved in sensory processing, using fMRI. By measuring changes in brain activity patterns across these regions, our approach accommodates the different and changing feature combinations present in natural scenarios, such as walking on a busy street. Our model reproduces people’s duration reports for natural videos (up to almost half a minute long) and, most importantly, predicts whether a person reports a scene as relatively shorter or longer–the biases in time perception that reflect how natural experience of time deviates from clock time

Research into pitch perception and time perception has typically treated the two as independent processes. However, previous studies of music and speech perception have suggested that pitch and timing information may be processed in an integrated manner, such that the pitch of an auditory stimulus can influence a person’s perception, expectation, and memory of its duration and tempo. Typically, higher-pitched sounds are perceived as faster and longer in duration than lower-pitched sounds with identical timing. We conducted a series of experiments to better understand the limits of this pitch-time integrality. Across several experiments, we tested whether the higher-equals-faster illusion generalizes across the broader frequency range of human hearing by asking participants to compare the tempo of a repeating tone played in one of six octaves to a metronomic standard. When participants heard tones from all six octaves, we consistently found an inverted U-shaped effect of the tone’s pitch height, such that perceived tempo peaked between A4 (440 Hz) and A5 (880 Hz) and decreased at lower and higher octaves. However, we found that the decrease in perceived tempo at extremely high octaves could be abolished by exposing participants to high-pitched tones only, suggesting that pitch-induced timing biases are context sensitive. We additionally tested how the timing of an auditory stimulus influences the perception of its pitch, using a pitch discrimination task in which probe tones occurred early, late, or on the beat within a rhythmic context. Probe timing strongly biased participants to rate later tones as lower in pitch than earlier tones. Together, these results suggest that pitch and time exert a bidirectional influence on one another, providing evidence for integrated processing of pitch and timing information in auditory perception. Identifying the mechanisms behind this pitch-time interaction will be critical for integrating current models of pitch and tempo processing.