Two year postdoctoral fellowship: Adults’ and children’s understanding of time

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Post-Doctoral Research Fellow

School of Psychology

Queen’s University Belfast

This post is funded for two years from 1st January 2018 as part of the interdisciplinary AHRC project “Time: Between Metaphysics and Psychology” led by Professor Christoph Hoerl (Philosophy, Warwick) and Professor Teresa McCormack (Psychology, Queen’s University Belfast). The Research Fellow will be based at Queen’s University Belfast, and will work with Professor Teresa McCormack to design and carry out novel studies of adults’ and children’s understanding of time. This is an opportunity for a researcher interested in working within an interdisciplinary team to develop an original and exciting programme of empirical work.

Informal enquiries may be directed to Professor Teresa McCormack, t.mccorma=<>, (++44) (0)2890974174

Anticipated interview date: Monday 30th October

Closing date: 4th October

TRF Newsletter – September 2017

Dear all,
We are pleased to share the September 2017 Newsletter of the Timing Research Forum. We have a number of exciting and important updates and announcements about the upcoming TRF Conference in Strasbourg, which is just a month away!
1. TRF Conference
– Registration
– Conference Program
– Business Meeting
– Accommodation
– TRF1 Speakers Q&A
– Poster instructions
– Open content (optional)
– Call for Social Media Outreach
– Call for Photographers
2. TRF Membership
3. TRF Blogs
4. TRF Mailing List
5. Blog your Paper
6. Blog your Conference
7. Contribute to TRF
8. Feedback for TRF
Dates: October 23-25, 2017
Contact: Anne Giersch –
If you are planning to attend the TRF Conference, and haven’t registered yet, we would encourage you to do so at the earliest!
See the following link to complete your registration –
In case you no longer plan to attend the conference, can you let us know ASAP (email Anne) so we can make arrangements accordingly.
We’ve updated the details and schedule for all symposia, oral sessions as well as the poster blitz session on the website. We encourage all attendees to use this to plan their conference itineraries in advance –
We will organize a TRF Business Meeting from 9-10AM on Oct. 25 that is open to all TRF members. We have several announcements to make during the meeting, including the exciting location of the 2nd TRF Conference in 2018!
We would like to remind everyone to book their accommodation for Strasbourg if you haven’t already, as hotels are being booked out quickly for the conference dates. If you would like to share rooms/airbnb with other attendees, please let us know at trf@timingforum.orgso we can pair you according to your preferences.
TRF is pleased to launch a new initiative – Speakers Q&A, where select Speakers share their thoughts on the current state-of-the-art in timing research and their presentations during the TRF Conference.
The first Speaker Q&A column features Dean Buonomano, who is organizing a symposium, ‘Timing, Neural Dynamics, and Temporal Scaling’ –
The second Speaker Q&A column features Warren Meck, one of the Keynote Speakers –
We’ve many more columns planned in the weeks leading up to the TRF conference, and we hope that enjoy reading these Q&A columns.
Participants who have been selected to present their work on a poster can find the relevant instructions here –
TRF supports open science and would like to encourage all participants to share their work (slides/posters etc.) with all conference attendees. It is completely optional and we hope that you would join us in openly celebrating the science that is presented at the TRF Conference.
Please share your final slides/posters in advance of the conference by emailing us at /
TRF has a very strong presence on a variety of social networks including ResearchGate, Twitter and Facebook.
We encourage all conference attendees to support our social media outreach by tweeting before/during the conference by tagging @timingforum and using the hashtag #TRF1. We hope this will inform the wider scientific community who will not be at the TRF conference about the science and the conference.
We would like to invite volunteers who are amateur/expert photographers and are happy to wield their DSLRs/smart phone cameras to capture a variety of moments during the conference. Please email us at to register your interest. All photos will be made available in a shared folder for all conference attendees to view and download.
To become a member of TRF and join a community of ~ 600 timing researchers, please fill in the form here –
ResearchGate: 313 followers (+15.5%)
Twitter: 321 followers (+25.9%)
Facebook: 377 followers (+25.7%)
We have a number of new blog articles reviewing recent papers on timing by a number of promising early career researchers.
Please read, share, comment and discuss!
Bowen Fung, University of Melbourne:
1) Dopamine encodes retrospective temporal information
2) The P3 and the subjective experience of time
3) Time perception in Schizophrenia
Bronson Harry, MARCS Institute, University of Western Sydney:
1) Perceptual reorganisation in deaf participants can high level auditory cortex become selective for visual timing
Mukesh Makwana, Centre of Behavioural and Cognitive Sciences, University of Allahabad:
1) Intended outcome appears longer in time
2) Olfactory visual sensory integration twists time perception
3) What language you speak shapes your subjective time
Molly Henry, University of Western Ontario:
1) Review of a number of rhythm and timing conference in the summer of 2017
2) Implicit variations of temporal predictability shaping the neural oscillatory and behavioral response
3) Sequence learning modulates neural responses and oscillatory coupling in human and monkey auditory cortex
If you would also like to contribute as a TRF blogger, please get in touch:
Everyone is invited to share any items related to timing related positions, grants, news, or anything that concerns timing research with the TRF community via our mailing list.
Make sure to use plain text when sending these messages (i.e. no attachments or fancy formatting is allowed). Please keep in mind that the mailing list is monitored, and only the the items approved by the mailing list moderators will be circulated to our community. Looking forward to your emails!
Please email your items directly to
We invite TRF members to submit short summaries of their recently published articles on timing. Articles should be no longer than 500 words and not include more than one representative figure.
Please submit your entries after your paper is published by emailing us at Submissions are open anytime and will be featured on the TRF blog page –
We invite TRF members to blog about their experience of a timing conference/meeting/workshop that you have recently attended. Submissions can highlight prominent talks/papers presented, new methods, trends and your personal views about the conference. Pictures may also be included.
Please submit your articles (no longer than 1000 words) to within two months from the date of the conference you intend to highlight.
TRF aims to host pertinent timing related resources, so that the TRF website acts as the definitive platform for everything related to timing research. The current resources listed on the TRF website include: (1) all members’ publications, (2) timing related special issues, (3) books on timing, (4) list of meetings focused on timing, (5) list of timing related societies/groups, (6) as well as code and mentoring resources.
TRF ecnourages open science and supports sharing of relevant information and knowledge between its members, with the aim to advance the field of timing research. We therefore invite you all to contribute to these resources. Please email us ( your suggestions for new resources for the timing community.
As an open academic society, we hope that you participate freely and support the TRF community in achieving its mission. As we like to repeatedly emphasize, TRF’s aim is to serve all timing researchers through open exchange of ideas, information and resources to advance the timing research community. We are open to receiving your suggestions or ideas that will help TRF grow and continue to deliver on its mission. We look forward to your feedback!
With best wishes,
Sundeep Teki
University of Oxford
Argiro Vatakis
Cognitive Systems Research Institute

Warren Meck: TRF1 Speaker Q&A


Warren Meck obtained a B.A. degree in psychology from the University of California, San Diego, a Ph.D. in experimental psychology from Brown University, and has been a professor at Brown University, Columbia University, and now Duke University.

His publications are accessible at Google Scholar and can be downloaded at ResearchGate, which also hosts preprints and descriptions of current research projects.


How can we determine the brain’s code for time?

It will take well-designed psychophysical studies in combination with neuroimaging, optogenetic stimulation, and electrophysiological recording techniques (triangulation) to break the code. Evaluating subjects with selective lesions and/or genetic backgrounds will continue to be important as well.


What will your talk at the 1st Timing Research Forum Conference focus on?

My talk will focus on the pervasiveness of timing abilities across animal species and the idea that a common timing mechanism is used that co-evolved with motor systems, i.e., to move is to time.


What according to you are the most pressing and fundamental questions in timing research?


a) To map out the “temporal connectome” for time, whereby central timing mechanisms can monitor and synchronize satellite timing mechanisms.

b) To better understand the relationship between intelligence/working memory capacity and timing accuracy/precision.


What current topics/techniques or new advances in timing research are you most excited about?

Optogenetics, i.e., selective stimulation of specific types of neurons and/or pathways thought to be involved in controlling the speed of the “internal clock” as well as its mode of operation (e.g., run, pause, and reset).


What advice do you have for students and postdoctoral researchers interested in investigating the brain’s code for time?

I would first recommend that students keep in mind the inspiration provided by Robert Rousseau (Laval University) in his forward to the book Functional and Neural Mechanisms of Interval Timing (CRC Press, 2003).

“For more than a century, time has been an object of study in experimental psychology. In his Experimental Psychology, Titchener (1905) wrote, “A student who knows his time sense … has a good idea of what experimental psychology has been and of what it has come to be.” At the dawn of the 21st century, I believe that Titchener’s judgment about the status of timing and time perception in psychology is still appropriate. As was the case a century ago, knowledge of the current research on timing gives a sense of what cognition, cognitive psychology, and cognitive neuroscience have come to be and will become.”

I would also advise students to learn as much as they can about the different levels of analysis that can be applied in the study of timing and time perception in humans and other animals. For me, this would involve comparative neuroanatomy, electrophysiology, and computational modeling.

Dean Buonomano: TRF1 Speaker Q&A

Dean Buonomano is Professor at the Departments of Neurobiology and Psychology, University of California Los Angeles. At the 1st TRF Conference, he is the Organizer of a symposium on ‘Timing, Neural Dynamics, and Temporal Scaling‘. He regularly tweets about time at @deanbuono.


How can we determine the brain’s code for time?

I don’t think there will be a single code for time any more than there is a single code for space in the brain. I think there be will be a number of ways the brain represents and tracks time, depending on the time scale and task at hand. Timing is simply to integral to the brain’s fundamental computations to rely on a single strategy. We have increasingly compelling evidence that in some cases temporal information is encoded in dynamically changing neural activity patterns (population clocks) or ramping of firing rates. The challenge will be to understand the mechanisms by which these codes are generated, and the domain in which different coding and timing strategies are relevant (a problem related to the Taxonomy of Time, see #3 below).


What will your talk at the 1st Timing Research Forum Conference focus on?

A striking ability we have at the both the sensory and motor level is to recognize and generate temporal patterns at different speeds—such as the tempo of music or the speed of speech. Along with Hugo Merchant and Mehrdad Jazayeri my talk will focus on the problem of temporal scaling: the ability to produce simple or complex temporal motor patterns at different speeds.


What according to you are the most pressing and fundamental questions in timing research?

I think the most pressing question in the timing field may be defining what exactly we mean by the timing field. Specifically, there is an increasing recognition that we need a Taxonomy of Time. A taxonomy of memory (e.g., Procedural x Declarative) was in many ways one of the most important advances in the study of learning and memory in the 20th century. The timing field is severely hampered by our inability to define and pinpoint the different forms, and time scales, of timing and temporal processing.


What current topics/techniques or new advances in timing research are you most excited about?

To date most studies have primarily focused on the activity or contribution of a given brain area in a timing task. But the brain is one big “which came first the chicken or the egg” problem when it comes to cause and effect. So I’m excited about improvements in our ability to record from hundreds of neurons in multiple different brain areas simultaneously. I think focusing on the transformations that happen between areas and the differences in representations will provide a powerful tool to understand timing and temporal processing.


What advice do you have for students and postdoctoral researchers interesting in investigating the brain’s code for time?

Read, and try to seek out opportunities to write reviews and perspectives.

Dopamine encodes retrospective temporal information

A new study published in Cell Reports shows that midbrain dopamine neurons are sensitive to previously experienced time intervals, and that this is likely to be important in terms of reward processing. Midbrain dopamine neurons are frequently discussed in terms of their roles in reward, motivation, and certain forms of learning. However, within the time perception literature, we commonly associate dopamine as modulating the rate of the internal pacemaker. Naturally, these functions of dopamine are not exclusive, and this study makes important progress in integrating them.

Dopamine in reinforcement learning

While early research implicated dopamine as the principle neurotransmitter responsible for the hedonic nature of “liking” something, the contemporary view conceptualises dopaminergic activity as a reinforcement signal that facilitates learning, rather than directly causing pleasure. This is in part due to the classic finding that phasic dopamine activity in the mesolimbic pathway constitutes a reward prediction error (the difference between expected and received reward), commensurate with prescriptive models of reinforcement learning.

During learning, dopamine responses gradually transfer to the earliest predictors of a reward, and after this associative pairing is established, response to the reward itself is reduced or absent. Importantly, this means that these response dynamics are fundamentally sensitive to the expected time of reward delivery.

Further to this, if rewards are delivered at different delays, the phasic responses of dopamine neurons to cues signalling these rewards depend on the duration of the delay (as well as reward probability, magnitude and type). This decreased response to longer reward delays typifies the economic principle of temporal discounting: rewards are devalued as a function of delay until their receipt. In reflecting the reduced value of delayed rewards, these neural responses demonstrate sensitivity to timing and appear to encode the intervals between cues and prospective (i.e. future) rewards.

Dopamine and time perception

In addition to its associations with motivation and reward, as a pharmacological agent, dopamine has been routinely acknowledged to play a significant role in time perception, in what some refer to as the ‘dopamine clock hypothesis1. Two sets of evidence in particular highlight this.

Firstly, non-human animal studies have pharmacologically manipulated dopamine during time perception tasks. When given dopamine agonists (e.g. methamphetamine) during a peak interval procedure, rats’ response rates peak earlier, as if their internal pacemaker was accelerated. When given dopamine antagonists (e.g. haloperidol), peak responses are later, commensurate with a slowing of the pacemaker2.

Secondly, electrophysiological and optogenetic studies of neurons in the substantia nigra (which produces dopamine and has inputs to the striatum) have shown that optogenetic activation or suppression of these neurons result in later and earlier timed responses, respectively. These results respectively reflect a slower or faster internal pacemaker, which is the opposite pattern of results seen in the pharmacological studies.

The present study

From the background above, we can see that dopamine appears to be involved in both time perception and reward processing. However, dopamine neurons have previously only been shown to encode elapsing and future delays. The study from Fonzi et al. questioned whether dopamine signals could also convey information related to retrospective, past delays. For example, do dopamine responses to a reward cue encode how much time has already been invested in the pursuit of the reward?

The researchers developed a Pavlovian conditioning paradigm with two reward cues that provided identical information about an upcoming reward, but differed in terms of how much time had elapsed since the previous reward. One cue was only presented after a 15–25 s wait time (“short cue”), while the other was only presented after a 65–75 s wait time (“long cue”). The researchers trained rats with this design while simultaneously using fast-scan cyclic voltammetry to record dopamine concentration in the nucleus accumbens core. If the dopamine responses to the short and long cues did not differ, then it would seem that dopamine activity only encodes prospective information. On the other hand, if the dopamine response to the long cue was larger than that of the short cue, this could be said to reflect the sunk cost of time. Conversely, if the signal to the long cue was decreased relative to that of the short cue, this could be said to reflect the rate of reward3.

The results showed that within this simple experimental design, dopamine responses to the long cue were decreased relative to short cue, suggesting that dopamine in the nucleus accumbens encodes reward rate. An alternative possibility was that this differing dopamine response reflected differing expectations about the time of delivery – the response to the long cue could be decreased because as time elapses, it is increasingly likely that the cue will be shown (i.e. a change in hazard rate). However, there was no relationship between the dopamine response and the time elapsed within each cue type. Furthermore, when another cohort of rats was trained with only a single cue for both short and long wait conditions, no differences were seen in the cue-evoked dopamine response for different wait times. Both of these results speak against the possibility that the dopamine response reflected the changing likelihood of reward delivery over time.

Notably, the principle finding above relied on a single analysis, and the relative difference between the short and long cues. The authors of the study thus performed a follow up analysis to determine whether this retrospective temporal information could be encoded when the animals were not able to directly compare cues. To do this, they trained an independent cohort of rats with short trials and long trials in separate sessions. Even in these scenarios, the short cue evoked a larger dopamine response than the long cue, which suggested that the encoding of retrospective delays was context-independent.

However, once these rats were exposed to both cues in a mixed session, the response to the short cue was increased. While for most of the above experiments there were no differences in behaviour between the two conditions, this increase in dopamine response to the short cue in this intermixed session was also accompanied by an increase in behavioural responding. This implies that (while elapsed wait times can be learnt independently) the dopaminergic encoding of retrospective delays is not entirely context-independent. It also shows that while there are not generally behavioural differences between the short and long cues, there appear to be changes in behaviour when there are also changes in dopamine response.

In a final analysis, the researchers also investigated the effect of the previous trial type, and the tonic dopamine signals over the waiting time. Firstly, for rats recently switch from the separate sessions to an intermixed session, they found that dopamine responses to short cues were significantly increased when the preceding trial was a long cue trial, compared when the preceding trial was a short cue trial. Similarly, dopamine levels were increased during the waiting period after long cue trials, relative to short cue trials (but only up to 25 s, before the identity of the current trial was known). From around the point that the identity was known (25 s), conditioned responding decreased when the preceding trial was a long cue trial, relative to when it was a short cue trial. One possible implication here is that a decrease in wait time dopamine could promote increased anticipatory responding. This would be consistent with the electrophysiological and optogenetic evidence that reducing dopamine increases pacemaker rate (see above).

It is important to reiterate that the results in the former two paragraphs only applied to the experiments where rats where moved from separate training on the short and long cues to an intermixed schedule. These results therefore represent peculiarities in how these animals learnt and adapted to their new context. Overall, the results of the first experiment are the most important here: phasic dopamine responses encode previous durations and appear to constitute a signal of previous reward rate.

This study compellingly demonstrates how even simple experimental designs can lead to novel and valuable findings. The fact that nucleus accumbens dopamine responses encode reward rate suggests a potential mechanism that could normalise value signals for future rewards, and provide contextual information such as the sunk cost of time.

If cue-evoked dopamine responses have to encode durations over a large range of timescales (potentially over 15 orders of magnitude) one interesting future avenue for research would be to describe the mapping between these dopamine responses and the duration of the delays preceding them, in order to precisely understand how durations are represented. More work needs to be done to comprehensively understand the functions of tonic and phasic dopamine and how they relate to perceived and experienced durations, but this study makes substantial progress toward this goal.

Source paper:

Fonzi, K. M., Lefner, M. J., Phillips, P. E. M., & Wanat, M. J. (2017). Dopamine Encodes Retrospective Temporal Information in a Context. Cell Reports 20(8), p. 1774. doi: 10.1016/j.celrep.2017.07.076

  1. It should be noted that much research into the neurobiology of reward and motivation typically focuses on the mesolimbic dopamine pathway. This is in contrast to time perception research, which is more often related to the nigrostriatal pathway (this is also commonly associated with movement). However, these pathways are not independent and the nigrostriatal pathway has also been shown to be critical for reward processing. ↩︎
  2. When both drugs were delivered simultaneously, rats’ peak responses are similar to that of a control condition. ↩︎
  3. Previous research has suggested that longer timescale tonic dopamine activity encodes reward rate. ↩︎

Perturbing and Enhancing Perception and Action using Oscillatory Neural Stimulation

The workshop “Perturbing and Enhancing Perception and Action using Oscillatory Neural Stimulation” (PEPA ON Stimulation), scheduled for the 18th/19th January 2018 at the MRC Cognition and Brain Sciences Unit (CBU) in Cambridge, UK. Abstract submission is open.

The workshop might be of interest for some researchers working in the field of timing and rhythm.

Please refer to our website for detailed information: