Observations on recent progress in the ﬁeld of timing and time perception !
Auditory Neuroscience Group, Department of Physiology, Anatomy & Genetics,
University of Oxford, South Parks Road, Oxford OX1 3QX, UK.
Time is an important dimension of brain function, but little is still known about the
underlying cognitive principles and neurobiological mechanisms. The field of timing and
time perception has witnessed rapid growth and multidisciplinary interest in the recent years
with the advent of modern neuroimaging, neurophysiological and optogenetic tools. In this
article, I review the literature from the last ten years (2005-2015) using a data mining
approach and highlight the most significant empirical as well as review articles based on the
number of citations (a minimum of 100 citations). Such analysis provides a unique
perspective on the current state-of-the-art in the field and highlights subtopics in the field
that have received considerable attention, and those that have not. The objective of the article
is to present an objective summary of the current progress in the field of timing and time
perception and provide a valuable and accessible resource summarizing the most cited
articles for new as well current investigators in the field.
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This article was originally written for an invited review on Auditory perception and Timing for a special issue of
Current Opinion in Behavioural Sciences on ‘Interval Timing and Skill Learning: The Multisensory Representation of
Time and Action’ guest edited by Warren Meck and Richard Ivry. The article was rejected as it did not conform to the
speciﬁc journal guidelines and is reproduced here in its original form.
This is not an editorial piece. For that, please read the editorial article for this special issue on
‘Interval Timing and Skill Learning: The Multisensory Representation of Time and Action’, by
Richard Ivry and Warren Meck, who are in the best position to comment on the various review
articles and provide a succinct summary of the ﬁeld. My objective is to present my personal
observations on the current state of research on timing and time perception, speciﬁcally from
the point of view of a postdoctoral researcher interested in building an independent research
program focused on timing.!
My advisor, Tim Grifﬁths and I were invited by the editor, Warren Meck to contribute a review
focused on ‘Auditory perception and timing’. As per the guidelines circulated to all invited
authors, the review should focus on, and highlight recent articles of note, especially those
published in the last couple of years. My own contribution to the ﬁeld, during my doctoral
research with Tim Grifﬁths, is represented by two empirical papers (Teki et al., 2011, Teki and
Grifﬁths, 2014) and a review article (Teki et al., 2012) in a special issue on timing, that was also
edited by Warren Meck. More recently, Tim and I collaborated with Warren Meck and Melissa
Allman on a review elaborating the subjective principles of time perception (Allman et al., 2014). !
The guide for authors provided by the Current Opinion in Behavioral Sciences (COBS) clearly
stipulates that - “the aim of the manuscript is to review recent articles, with particular emphasis
on those articles published in the past two years.” As the guide encourages the authors to
describe recent trends and provide subjective opinions of the topics discussed, I decided to take
the liberty to do exactly as instructed, and provide my subjective opinions of recent progress in
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A review article is meant to highlight and discuss recent trends and results on a particular topic.
Invariably, there is a tendency on the part of most authors to emphasize one’s own work at the
expense of not adequately discussing results from other research groups. At the outset, I also
outlined a possible structure for the review based on my work on duration-based and beat-
based timing (Teki et al., 2011, 2012). However, this work does not comply with the publication
timeline suggested by the journal. Thus, I decided to not discuss this work which was discussed
in a recent review article (Allman et al., 2014). Even if the restricted timeline of two years were
to be relaxed, Tim and I struggled to ﬁnd ten recent articles of note on the particular topic and
published in the last two years to emphasize and highlight in our review. After further
discussions, we decided to focus on ‘Disorders of sequence and interval timing’ based on
recent neuropsychological work from Tim’s group that examined perceptual timing abilities of
patients with striatal and cerebellar degeneration in the form of Huntington’s disease, Multiple
Systems Atrophy and Parkinson’s disease (Cope et al., 2014a, b). !
This new topic was more exciting, for the list of authors and topics provided by COBS did not
include any clinically oriented articles. Although timing abilities are known to be impaired in a
range of neuropsychological and neurological disorders (Allman and Meck, 2012) including
Parkinson’s disease, Huntington’s disease, Multiple Systems Atrophy and Schizophrenia
amongst others, the extent to which their timing performance is affected and related to the
primary neurological deﬁcits is not fully known. However, even though this ﬁeld is gaining
traction and represents a highly fruitful area of translational research, a review of the recent
literature on this topic, again, yielded few promising articles to form the basis of a constructive
review that would be beneﬁcial to the ﬁeld.!
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At this juncture, it appeared that the two topics of specialization investigated in Tim’s lab could
not be covered in a detailed and topical review for COBS. At this point, I wondered whether the
lack of recent signiﬁcant breakthroughs is true of the ﬁeld of timing and time perception as a
whole, or is it just a representation of the speciﬁc topics we focused on? !
II. Key papers on timing and time perception!
To obtain a representative picture of the ﬁeld, I decided to examine recent papers by the authors
invited to contribute to this special issue (75 authors). As the list of invited authors was a fairly
small sample, I extended the list by considering all members of the recently concluded
European COST Action - Timely which included members from all over the world. This list
provided another 129 authors who were not already invited to contribute to the COBS special
issue, thus resulting in a reasonable sample size of 204 authors.!
A number of metrics are used to evaluate the quality and impact of research articles including
impact factor, h-index, i-10 index amongst others. Although none of these metrics are accepted
as standard across the scientiﬁc community, I decided on the number of citations as a metric as
it indicates the impact of a paper and how well the idea is accepted and circulated in the ﬁeld by
other researchers. It is not a perfect measure, for the number of citations an articles receives is
often skewed by the impact factor of the journal where it is published but good ideas tend to
circulate well no matter where the ideas are published. Furthermore, to draw reasonable
conclusions about progress in the timing ﬁeld, I focused on a period of 10 years and considered
all articles that were indexed in Google Scholar and published by the above set of authors from
2005 onward. In order to restrict my sample of publications and consider only the most impactful
papers (ideas), I used a threshold of a minimum of 100 citations. This timeframe is also not
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ideal, for it is biased towards older papers than more recent articles which have not had the
same time to accumulate as many citations. !
Using these search criteria, 66 papers (an average of 1 article per every 3 authors) were found
as described in Table 1. These papers covered research on topics related to perception of time,
rhythm, music, inter-sensory synchrony amongst others and used techniques including
psychophysics, neuroimaging, electrophysiology and modeling. Out of these 66 papers, 24
papers were review articles (marked with an asterisk next to the number of citations) that
received an average of 223.5 citations (Z-score ranged from - 0.65 to 4.44), i.e. one of out three
prominent articles on timing in the last ten years were review articles that discussed the current
state of research. The remaining empirical papers, 42 in all, received an average of 181.6
citations per paper (Z-score ranged from - 0.82 to 3.70). !
There are several conclusions to be drawn from Table 1, for instance that review articles tend to
dominate the overall citations in the ﬁeld while only an average of four signiﬁcant empirical
papers are published a year. Although many of these reviews are now ‘classic’ in the ﬁeld, even
the most recent article in the table is a review (Merchant et al., 2013). Among other things, this
suggests that either the ﬁeld is still in an embryonic stage where review articles by established
researchers are needed to set the precedent on certain topics or that the ﬁeld of timing is too
diverse, at an intersection of various ﬁelds including time perception, rhythm perception, music
perception, temporal coding, inter sensory asynchrony, motor timing and coordination, that is
reﬂected in the diversity of topics covered by the review articles. Although not evident from the
list, there has been a recent proliferation of review articles given the rise of specialist open
access journals like Frontiers that encourage researchers (repeatedly and frequently) to
commission special issues covering empirical work as well as reviews. It is not known whether a
similar analysis of the most recent and highly cited papers in other ﬁelds like memory, vision, or
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decision-making will yield the same ratio of reviews to empirical studies but one could make a
reasonable null hypothesis that this ratio may be smaller than for the ﬁeld of timing.
Alternatively, compared to ﬁelds like vision and memory that have been intense topics of
investigation for several decades the ﬁeld of timing is more nascent and does not boast a large
research community as evident by the number of participants at specialist meetings in such
ﬁelds, for instance, the annual Vision Science Society conferences. !
In order to drive more experimental work, it is clear that the ﬁeld of timing needs to attract more
young researchers and ensure a a bright future for the ﬁeld, and this would need concerted
efforts from the entire timing community. A recent positive step in this direction is represented by
the launch of a specialist journal for timing, Timing and Time Perception as well as its
corresponding review journal, Timing and Time Perception Reviews. Another step forward would
be the launch of an academic society exclusively for researchers in timing that would promote
interdisciplinary exchange of ideas amongst researchers with diverse interests in timing via
annual conferences that draw on a range of methods from purely behavioral to
neurophysiological and neuroanatomical measures, and from neurostimulation and
neuropsychological approaches to animal work and computational models; share pertinent
news and information like grant funding calls, new papers, job opportunities for doctoral and
postdoctoral candidates, workshops and training opportunities; and promote career
development of young researchers through grants for short cross-disciplinary collaborations or
exchange visits, funding for attending conferences and general mentoring support. Although
there already exist a few scientiﬁc societies and communities relevant to timing like the Society
for Music Perception and Cognition (SPMC: http://www.musicperception.org), Rhythm
Perception and Production Workshop (RPPW: http://rppw.org), European Society for Cognitive
Sciences of Music (ESCOM: http://escom2015.org), Society for Education, Music and
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Psychology Research (SEMPRE: http://www.sempre.org.uk), Deutsche Gesellschaft fur
Musikpsychologie (DGM: http://www.music-psychology.de), Asia-Paciﬁc Society for the
Cognitive Sciences of Music, Fondazione Mariani (http://fondazione-mariani.org/) that organizes
the NeuroMusic conferences, their scope is limited to music perception and psychology, and do
not cover the interests of the entire ﬁeld of timing and time perception. Society for Neuroscience
(SfN) represents the primary venue where researchers gather together for structured mini- or
nano-symposia on human and animal timing research but the scientiﬁc interaction and
discussions are limited given the hectic nature of SfN meetings. A recent example of such a
successful academic organization is the Society for the Neurobiology of Language (http://
www.neurolang.org/) funded by the National Institutes of Health, which since its inception in
2009, attracts more than 400 researchers for its annual conferences that are held alternatively
in the USA (as a satellite meeting of SfN) and Europe.!
III. Future directions!
Organizational considerations apart, there are several new scientiﬁc directions that the ﬁeld can
and should embrace to achieve a more comprehensive understanding of the neurobiology of
timing in natural environments. Animal models of timing focused on core timing networks
including the basal ganglia, cerebellum, premotor and parietal cortex (Grahn 2012; Teki et al.,
2012; Schneider and Ghose, 2012; Merchant et al., 2013; Allman et al., 2014; Hayashi et al.,
2015) will be key to understanding the encoding of time by neuronal ensembles. Such a line of
work has been recently pioneered by Hugo Merchant colleagues in rhesus macaques that
combines timing behaviors and the examination of the underlying neuronal code in the basal
ganglia (Merchant et al., 2011, 2013; Bartolo et al., 2014; Bartolo and Merchant, 2015). Recent
work by Mello et al. (2015) further demonstrated that a population code for time exists in the
striatum that scales with the interval being timed and multiplexes information about action as
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well as time. Optogenetic approaches in speciﬁc target cells in animal models will yield further
crucial insights into the causal role of such mechanisms and their impact on timing behaviors
(Grosenick et al., 2015). For instance, a recent study by Chen et al. (2014) reported rapid
modulation of striatal activity by the cerebellum via a disynaptic pathway which has implications
for the coordinated processing of temporal information in the two core timing areas.!
The other dominant view of timing is that it is not a computation of speciﬁc dedicated circuits but
rather the output of intrinsic neuronal dynamics (Karmarkar and Buonomano, 2007; Ivry and
Schlerf, 2008). In this respect, the activity of sensory areas including auditory, visual and
somatosensory cortices merits further attention. Combining optogenetics and single-unit
recordings in primary visual cortex (V1), Hussain Shuler and colleagues have recently provided
beautiful insights into how V1 responses predict and drive the timing of future actions
(Namboodiri et al., 2015) and recruit basal forebrain and cholinergic input within V1 to encode
the timing of visually cued behaviors (Liu et al., 2015). !
In order to obtain a fundamental understanding of timing, it is also imperative to use stimuli and
paradigms that mimic timing behaviors in the natural world. Such naturalistic sequences that go
beyond the use of single intervals that have been traditionally used will offer additional insights
on encoding of time as well as associated motor behaviors (Kornysheva and Diedrichsen,
2014). Table 1 and the reviews therein highlight that timing is not mediated by a single brain
area but rather involves a distributed network (Meck, 2005) in cortical and subcortical areas
including prefrontal, parietal, premotor and sensory cortices, insula, basal ganglia, cerebellum,
inferior olive amongst others. To form a clear picture of how timing is mediated by these
structures, it is also important to understand the core functions of these areas and what
particular aspect of timing they mediate, whether it is related to attention, memory or perception.
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The use of comparative paradigms in healthy human volunteers as well as clinical populations
that show timing deﬁcits such as patients with Parkinson’s, Huntington’s, Schizophrenia
amongst others will provide a more uniform understanding of timing functions and dysfunctions
in health and disease. An identical approach (and even the use of similar paradigms) in animal
models via use of control animals as well as lesion or knock-out models will complement
ﬁndings from the human literature and provide a more generic understanding of fundamental
mechanisms of timing.!
Irrespective of the present state of affairs, the ﬁeld of timing and time perception represents a
promising and highly active ﬁeld of research that is growing every year in terms of number of
researchers and scientiﬁc output and one where new students and researchers can ﬁnd a niche
topic and leave a signiﬁcant mark on the ﬁeld. !
Page of 923
Table List of 66 papers on timing, rhythm and music perception from 2005 to present sorted
according to the number of citations (minimum of 100 citations) in Google Scholar collated
during the period from August 30 to September 16, 2015 (see section II for more details).
Asterisks next to the number of citations denote review articles.!
Buhusi & Meck
Nat Rev Neurosci
Time is represented in a distributed
manner through coincidental
activation of cortico-striatal
Casasanto & Boroditsky
Spatial information affects
judgments about duration but not
Wittmann et al.
Social jetlag, i.e. the discrepancy
between social and biological
timing affects wellbeing and
Grahn et al.
J Cogn Neurosci
Basal ganglia and Supplementary
Motor Areas mediate beat
perception, in addition to motor
Review of recent behavioral and
neuroscientiﬁc studies of timing.
Karmarkar & Buonomano
Cortical networks can read out time
as a result of intrinsic network
Ivry & Schlerf
Trends Cogn Sci
Dedicated models of timing are
preferred over intrinsic models.
Shuler & Bear
Primary sensory cortex, like V1,
mediates reward-timing activity.
Coull et al
Review of neuroimaging,
psychopharmacological aspects of
Morrone et al.
Short intervals of time between two
successive perisaccadic visual
stimuli (but not auditory) are
Page of 10 23
Chen et al.
Passively listening to rhythms
recruits motor regions of the brain.
Droit-Volet & Meck
Trends Cogn Sci
Review of how emotional arousal
and valence modulates attentional
time-sharing and clock speed.
Patel et al.
Snowball, a cuckatoo, can
spontaneously synchronize its
movements to a musical beat.
Wittman & Paulus
Trends Cogn Sci
Review of how impulsivity affects
perception of time and decision
Winkler et al.
Proc Natl Acad Sci
Newborn infants show beat
MacDonald et al.
Hippocampal time cells encode
successive moments during a
sequence of events.
Brain & Cognition!
Review of timing that suggests a
distributed representation of time
across multiple neural systems.
Meck et al.
Review that proposes striatum
serves as a core timer, as part of a
distributed timing system.
Wiener et al.
Meta analysis that suggests distinct
for perceptual vs. motor timing;
SMA and right IFG are most
commonly activated in various
Coull & Nobre
Review that suggests basal ganglia
is key for explicit timing while
parietal and premotor areas
mediate implicit timing.
Nobre et al.
Review that describes how
temporal expectations modulate
perception and action, and the
underlying neural mechanisms.
Patel et al.
Beat perception and
synchronization show modality
speciﬁc beneﬁts for auditory vs.
visual beat patterns.
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Chen et al.
J Cogn Neurosci
Musicians show greater prefrontal
cortex activity vs. non-musicians
while tapping to complex auditory
Grahn & Rowe
Putamen, SMA and premotor cortex
are important for internal generation
of the beat and auditory motor
coupling during beat perception.
Lewis & Miall
Trends Cogn Sci
Dorsolateral prefrontal cortex
mediates working memory as well
Dopamine depleting lesions in
different parts of the basal ganglia
shows dissociable effects on
Noesselt et al.
Temporal correspondence between
auditory and visual streams
modulates activity of multisensory
STS as well as unisensory cortices.
Review of work on rhythmic
categorization which argues that
timing is distinct from rhythm.
Wittmann et al.
Exp Brain Res
Posterior insula mediates delayed
gratiﬁcation of reward while
striatum encodes time delay.
Kotz & Schwartze
Trends Cogn Sci
Review which suggests that
temporal and speech processing is
processed by cortical and
subcortical systems associated with
Burr et al.
Short visual events are encoded by
visual neural mechanisms with
localized receptive ﬁelds rather
than by a centralized supramodal
Vroomen & Kreetels
Review that focuses on
intersensory timing and
mechanisms that encode
Phil Trans R Soc B
Review that discusses different
models of time perception with a
particular focus on the insula as a
Page of 12 23
McAuley et al.
J Exp Psychol:
Event timing proﬁles for a battery of
perceptual-motor timing tasks vary
across the life span (4-95 years
Chen et al.
Metrical structure of musical
rhythms modulates functional
connectivity between auditory and
dorsal premotor cortex.
Boroditsky et al.
English and Mandarin speakers
think about time differently.
Ta at g en et al .
A time perception model based on
adaptive control of thought-rational
can explain effects of attention and
learning during time estimation.
Correa et al.
Review that focuses on how
temporal attention modulates the
amplitude and latency of ERPs like
N2 and P300 components.
Matlock et al.
Fictive motion inﬂuences temporal
Allman & Meck
Review that focuses on distortions
of time perception and timed
performance in various neurological
and psychiatric conditions.
Block et al.
Meta analysis that focuses on the
effects of cognitive load on
prospective and retrospective
Kotz et al.
Review that focuses on the non-
motor functions of basal ganglia
with particular emphasis on
prediction in speech and language.
Teki et al.
Perception of relative and absolute
time is mediated by distinct
networks based in the basal ganglia
and the cerebellum respectively.
Lewis & Miall
Dorsolateral prefrontal cortex
mediates working memory and
posterior parietal cortex and
anterior cingulate attentional
aspects of timing.
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Rubia et al.
Phil Trans R Soc B
Review that suggests that
impulsivity in ADHD is related to
compromised timing functions and
Styns et al.
Hum Mov Sci
Walking speed is modulated by the
tempo of musical and metronome
Kanai et al.
Temporal frequency of a stimulus
serves as the clock for perceived
Noulhiane et al.
Emotional stimuli are judged longer
than neutral stimuli, when balanced
for the levels of arousal.
Fuhrman & Boroditsky
Temporal judgments in nonlinguistic
tasks are inﬂuenced by culturally
speciﬁc spatial representations.
Grahn & Brett
Parkinson’s patients show selective
deﬁcits in discrimination of beat-
Eagleman et al.
Review of timing based on
imaging and computational
Keller et al.
Consc & Cogn
Action simulation in ensemble
musicians like pianists underlies
synchronization and self-
Correa et al.
Psychon Bull &
Temporal orienting enhances
Merchant et al.
Ann Rev Neurosci
Review that highlights the role of a
core timing mechanism in the basal
ganglia and its interaction with
context dependent areas.
Grahn & McAuley
Individual differences in beat
perception exist and modulate
activity in auditory and motor areas.
Zarco et al.
Performance of rhesus monkeys
and humans is compared on a
number of sub-second interval
Page of 14 23
Wearden et al.
Q J Exp Psychol
Decreasing arousal affects
performance on time perception
Vatakis & Spence
discrimination performance is better
for audiovisual stimuli of lower
Nozaradan et al.
EEG frequency tagging reveals
neural entrainment to beat and
Wassenhove et al.
Multisensory interactions inﬂuence
perception of time: vision can
impact auditory temporal
Ishihara et al.
A mental time line exists from left to
right along the horizontal axis in
Review that addresses cognitive
processes underlying joint action in
Casasanto et al.
Spatial information inﬂuences
temporal judgments more than time
affects spatial judgments in children
as well as adults.
Iversen et al.
Ann NY Acad Sci
Beta-band activity inﬂuences
auditory rhythm perception.
Droit-Volet & Gil
Phil Trans R Soc B
Review that addresses the role of
emotional context on timing.
Jahanshahi et al.
Basal ganglia and cerebellum are
involved in reproduction of both
short and long intervals.
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