TRF2 Blog Post

After a very successful TRF1 in 2017, this year saw the second bi-annual Timing Research Forum held from the 15-17th of October, 2019 in Queretaro, Mexico. The conference largely followed the format of its predecessor, with the extremely exciting addition of a Moonshot session and keynotes from Mehrdad Jazayeri, Albert Tsao and Kia Nobre. 

It would be difficult to comment on all the new ideas and findings, with so many exciting and inspiring talks/posters in one place, but some of the musings I took away were related to how we still have a long (and adventurous) road ahead of us in terms of defining what we truly mean when we speak of the specifics of timing and time perception. Specifically, several speakers proposed the idea of rethinking relative features of time such as what regularity truly means for the brain, and whether previously demonstrated functional networks account for all presentations of time as has been suggested (for example, beats and intervals), or rather whether different networks account for different aspects of time. The importance of rhythmic expectation via improved performance in behavioural measures were also touched upon, but again, extrapolating these behavioural effects to occur as a result of temporal expectation is a perilous stance that still requires further exploration (and may actually be more challenging than we initially anticipate). With every meeting dedicated to exploring time perception, it becomes increasingly obvious that we don’t all mean the same thing when we speak of time. Investigating the perception of ‘rhythms’ or ‘beat’ is distinctly different to the perception of ‘intervals’, and even that is different to the processing of ‘durations’. Perhaps, however, these differences are what make the study of time endlessly exciting and compelling – that there truly is so much more we are yet to discover (and eventually piece together). 

The Moonshot session was a novel and particularly exciting addition at this years meeting. The basic premise for the discussion was this – 

“For the next 5 years, all techniques, methods, and workforce are available to solve the question you think is essential to understand time in the brain. Which question would that be, and why?”. 

Speakers proposed several really thought-provoking and progressive advances (the session was fully livestreamed and if you didn’t get a chance to tune in, you can still catch up at Some of the suggestions included a focus on the ontology of timing and a particular need to develop a comprehensive taxonomy of time, as several accounts of interval timing actually describe pattern timing. Another suggestion was the immediate need to explore if distinct neurochemical systems mediate distinct aspects of timing and more social approaches to this question, for example, why synchronous movements with others increases social affiliation (even as early as infancy), and in a similar vein, the comparison that unlike humans, other primates (monkeys) do not spontaneously perform periodic temporal prediction, highlighting that predictive motor entrainment is intrinsically rewarding to humans (..and why exactly to species differ)?. 

While the Moonshot allowed for many exciting ideas to be brought to the forefront, I was especially excited by Molly Henry’s radical approach to the perusal of needing to delimit the now, and what exactly this means. Molly described reports on an ‘expanded now’ from individuals on LSD and other hallucinogens, and proposed that if we truly were to aim for the moon, considering the use of such stimulants to explore the extent of the ‘now’ might actually be more fruitful and eye-opening (excuse the pun!) than we realise. Whilst this leaves much to think about (and in some cases, reconsider), there is something to be said for the feeling of each of us working together to piece this puzzle in our own creative ways.

In conclusion, it is no understatement to say TRF2 was a resounding success – presenting novel insights, but also reaffirming the need to continue pushing our own assumptions of approaching the enigma of timing creatively. Building on the foundations of its predecessor, TRF2 has paved the way for more exciting work to be presented at TRF3 in Groningen (The Netherlands). Thanks again to all the organisers and attendees. See you in 2021! 

Aysha Motala is a postdoctoral fellow at the Brain and Mind Institute (Western University, Canada) working on cross-modal timing and neuroimaging of speech and rhythm processing 


Welcome Martin Wiener as our first Board of Directors!

We are excited to announce that Dr. Martin Wiener will be joining TRF’s newly instituted ‘Board of Directors’ as our first Director. 
Dr. Martin Wiener’s lab is engaged in understanding the neural mechanisms and computations underlying time, space, and action. His work employs numerous techniques, including fMRI, EEG, TMS and tES, as well as combinations of these. His work additionally explores how the neural mechanisms for time adapt to different experimental contexts. 

Martin is an active member of the timing research community and we are thrilled to have him on board and help shape TRF’s mission and support its 700+ members community. Martin will be at the TRF2 Conference in Mexico in case anyone wants to discuss anything related to TRF. 
Best wishes,TRF

Timing & Time Perception Special Issue on “Temporal Illusions”

Hosted by Fuat Balcı & Argiro Vatakis 

Decades-long research in interval timing has primarily focused on the psychophysical properties of this fundamental function typically in consideration of veridical timing behavior. Along the similar vein, generative models of interval timing mostly focus on the processing dynamics of the internal stop-watch in its default mode. Both of these approaches have largely overlooked the malleability of perceived time by exogenous factors such as stimulus intensity and endogenous factors such as physiological arousal. These very relations could actually help researchers better understand the representational constitution of subjective time and the processing dynamics of the internal stop-watch. This special issue aims to cover a wide range of empirical and theoretical work on the effects of different factors (e.g., stimulus features, physiological states, emotional states, drugs) on timing and time perception in humans and other animals.

Submission procedure:

1. Full paper submission by November 1st, 2019.

Instructions for submission: The submission website is located at: To ensure that all manuscripts are correctly identified for inclusion into the special issue it is important to select “Special Issue: Temporal Illusions” when you reach the “Article Type” step in the submission process. More details on format that must be followed in preparing your manuscripts see here

3. Standard peer review/revision process will be followed.

4. Final decisions are expected by May 15th, 2020.

PhD position in audio-visual synchrony perception at TU Eindhoven

4-years PhD position in audio-visual synchrony perception at TU Eindhoven, The Netherlands

Within the framework of the EU funded Marie Skłodowska Curie Initial Training Network VRACE (Virtual Reality Audio for Cyber Environments) a 4-years PhD position is available in the Human-Technology Interaction group of the TU Eindhoven, The Netherlands, under the supervision of Armin Kohlrausch.

Applications are invited (and only possible) via the WEB portal of the TU/e. More detailed information on the position and details how to appply can found at:

Important: According to the eligibility rules of the EU, specific mobility requirements apply to this position, which are described in detail on the job site. Please check before applying whether you fulfill these requirements. 

The application deadline is April 15, 2019, start of the position preferably as soon as possible after the end of the selection process.

Meta-analysis of neuroimaging during passive music listening: Motor network contributions to timing perception

We often learn about what the brain is doing by observing what the body is doing when the brain is focused on a task. This is true for investigations into rhythmic timing perception. Many insights into timing have resulted from careful observation of sensorimotor synchronization with auditory rhythms. This draws from the works of Bruno Repp that suggest that perception of auditory rhythms relies on covert action—that synchronizing with a sequence is not so different than simply perceiving a sequence without moving along with it.

More specifically, in order to synchronize a finger-tap, or any other body movement, with an auditory stream, some timing prediction is necessary in order to perform all movement planning, effector assembly and execution in time with the auditory beat instead of several milliseconds too late. If we must plan for a synchronized movement in advance, and there is some automaticity to this planning when we listen to auditory rhythms, then it is reasonable to ask whether we also perform some degree of motor planning every time we perceive a rhythm even if we do not move any body part in time with it.

The evidence suggests we do use our motor systems, or at least that our motor systems are actively being used for some purpose while perceiving rhythms when we are not synchronizing. Brain images during rhythm perception experiments consistently show activation in areas of the brain that are known to be involved in movement of the body. These areas include primary motor cortex, premotor cortices, the basal ganglia, supplementary motor area, and cerebellum. Details about covert motor activity are still being investigated, but some theories suggest covert motor activity plays an essential role in rhythmic timing perception, a theory many music cognition researchers find intriguing.

But first, what does the neuroimaging literature actually say about which motor networks are active, and which rhythm perception tasks elicit this covert action? Each study uses musical stimuli that vary on a number of features, give different instructions to the subjects on how to attend to or experience the stimuli, and these differences induce varying emotional states, arousal, familiarity, attention and memory. However, across all this stimulus variability, motor networks still robustly present themselves as players in rhythm perception. Interestingly, the stimulus variability shows up less in whether we see covert action and more in which motor networks are covertly activated.

In a recent meta-analysis of neuroimaging studies on passive musical rhythm perception, Chelsea Gordon, Patrice Cobb and Ramesh Balasubramaniam (2018) asked which covert motor activations are most reliable and consistent across studies. They used the Activation Likelihood Estimation (ALE; Turkeltaub et al., 2002), derived from peak activations in Talairach or MNI space, to compare coordinates across all PET and fMRI studies with passive music listening conditions in typically healthy human subjects. Their sample included 42 experiments that met the criteria for inclusion. As expected, the results of the ALE meta-analysis revealed clear and consistent covert motor activations in various regions during passive music listening. These activations were in premotor cortex (bilaterally), right primary motor cortex, and a region of left cerebellum. Premotor activation patterns could not be further localized to dorsal or ventral subregions of premotor cortex, but were dorsal, ventral or both dorsal and ventral. Right primary motor activations might have been excitatory or inhibitory, and were stronger in studies that asked subjects to anticipate later tapping to a beat in subsequent trials or to subvocalize humming. Most consistent across studies were premotor and left cerebellum activations, supporting predictive theories of covert motor activity during passive music listening.

One surprising aspect of these results is that the ALE meta-analysis did not find consistent activation in SMA, pre-SMA or the basal ganglia. The authors suggest that basal-ganglia-thalamocortical circuits may be specifically involved in subjects with musical training, or only in tasks with specific instructions to attend to the rhythmic timing of the stimuli instead of to listen passively.

An important concern Gordon and colleagues raised in the discussion is that of how publication bias contributes to ALE results. Also described by Acar et al. (2018), unpublished data deemed uninteresting can lead to biases in meta-analytic techniques (known as the file drawer problem), including in the ALE measure. Gordon et al. attempted to account for the file drawer problem by contacting all authors of the analyzed manuscripts to ask for the full datasets from each study to use in their ALE analysis. However, many authors did not provide this data for unreported brain activations, leading to limitations in number of explanatory contrasts that could be performed and a possible influence of publication bias on the ALE results.

The ALE technique is a powerful tool in performing large-scale neuroimaging study meta-analyses, but as with any meta-analysis technique of published results could be susceptible to the pitfalls of the file drawer problem. That being said, covert motor activity during passive music listening presents consistently across studies, even with considerable stimulus variability. This may support that timing prediction uses premotor and cerebellar networks.

Jessica M. Ross (


Gordon, C.L., Cobb, P.R., Balasubramaniam, R. (2018). Recruitment of the motor system during music listening: An ALE meta-analysis of fMRI data. PLoS ONE, 13(11), e0207213.


Acar, F., Seurinck, R., Eickhoff, S.B., Moerkerke, B. (2018). Assessing robustness against potential publication bias in Activation Likelihood Estimation (ALE) meta-analyses for fMRI. PLoS ONE, 13(11), e0208177.

Turkeltaub, P.E., Eden, G.F., Jones, K.M., & Zeffiro, T.A. (2002). Meta-analysis of the functional neuroanatomy of single-word reading: Method and validation. Neuroimage, 16, 765–780.