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doi: 10.1098/rstb.2009.0003 , 1955-1967364 2009 Phil. Trans. R. Soc. B
Marc Wittmann
The inner experience of time
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Review
The inner experience of time
Marc Wittmann*
Department of Psychiatry, University of California San Diego, 9500 Gilman Drive,
La Jolla, CA 92093-9116A, USA
The striking diversity of psychological and neurophysiological models of ‘time perception’
characterizes the debate on how and where in the brain time is processed. In this review, the most
prominent models of time perception will be critically discussed. Some of the variation across the
proposed models will be explained, namely (i) different processes and regions of the brain are
involved depending on the length of the processed time interval, and (ii) different cognitive processes
may be involved that are not necessarily part of a core timekeeping system but, nevertheless, influence
the experience of time. These cognitive processes are distributed over the brain and are difficult to
discern from timing mechanisms. Recent developments in the research on emotional influences on
time perception, which succeed decades of studies on the cognition of temporal processing, will be
highlighted. Empirical findings on the relationship between affect and time, together with recent
conceptualizations of self- and body processes, are integrated by viewing time perception as entailing
emotional and interoceptive (within the body) states. To date, specific neurophysiological
mechanisms that would account for the representation of human time have not been identified. It
will be argued that neural processes in the insular cortex that are related to body signals and feeling
states might constitute such a neurophysiological mechanism for the encoding of duration.
Keywords: time perception; subjective duration; emotion; interoception; insula
1. INTRODUCTION
Throughout history, philosophers have been intrigued
by the nature of time and how we, as humans,
experience its progression. The perception of time is
part of human experience; it is essential for everyday
behaviour and for the survival of the individual
organism (Po¨ppel 1997;Wittmann 1999;Buhusi &
Meck 2005). Yet, and surprisingly enough, its neural
basis is still unknown. Temporal intervals, lasting only
seconds or spanning a lifetime, are judged according to
their perceived duration—often regarded as painfully
long or, the reverse, as lasting too short. Everyday
decisions we make, as simple as either waiting for
the elevator or taking the stairs, are based on the
experienced passage of time and anticipated duration
(Wittmann & Paulus 2008). The importance of our
temporal experiences for daily living is strikingly
documented in individual neurological cases where
patients report of an accelerated progression of time
and, consequently, have troubles in adequately inter-
acting with the environment, i.e. driving a car
(Binkofski & Block 1996). Although we doubtless
have a time sense, our bodies are not equipped with
a sensory organ for the passage of time in the same
way that we have eyes and ears—and the respective
sensory cortices—for detecting light and sound. Time,
ultimately, is not a material object of the world for
which we could have a unique receptor system.
Nevertheless, we speak of the perception of time.
When we talk about time (‘an event lasted long’, ‘time
flew by’), we use linguistic structures that refer to
motion events and to locations and measures in space
(Evans 2004); a further indication that time itself is not
a property in the empirical world.
Despite a growing body of knowledge on the
psychology and on the neural basis of the experience
of time, the riddle for philosophers and scientists alike
is still unsolved: how does the mind (or, for that matter,
the brain) create time? Martin Heidegger’s paraphrase
of St Augustine’s famous quotation
1
‘In you, my spirit,
I measure times; I measure myself, as I measure time’
(Heidegger 1992) reflects a theoretical approach—
founded in western philosophical tradition—which
states that time is a construction of the self. Perceived
time, thereafter, represents the mental status of the
beholder. In terms of a functional equation, one could
state that time tis a function fof the self, where the self
stands for all possible psychological (i.e. empirical
and theoretical) properties of an individual who
perceives time,
tZfðselfÞ:ð1:1Þ
Psychological research has shown that cognitive func-
tions such as attention, working memory as well as long-
term memory determine our temporal judgements
(Brown 1997;Zakay & Block 2004;Ta a t g e n et al.
2007). Moreover, drive states, moods and emotions
(Wittmann et al.2006;Droit-Volet & Meck 2007;
Phil. Trans. R. Soc. B (2009) 364, 1955–1967
doi:10.1098/rstb.2009.0003
One contribution of 14 to a Theme Issue ‘The experience of time:
neural mechanisms and the interplay of emotion, cognition and
embodiment’.
*wittmann@ucsd.edu
1955 This journal is q2009 The Royal Society
on 31 May 2009rstb.royalsocietypublishing.orgDownloaded from
Noulhiane et al. 2007a) as well as factors of personality
(Rammsayer 1997) influence duration estimates. For
example, time intervals are judged to be longer when
we pay more attention to time and when the load of
varying experiences stored in memory is higher. Our
subjective well-being also strongly influences how time
is experienced. Time speeds up when we are involved in
pleasant activities, but it drags during periods of
boredom. Thus, our sense of time is a function of the
intricate interplay between specific cognitive functions
and of our momentary mood states.
The aforementioned psychological factors definitely
influence the processing of duration. However, a specific
neural timing mechanism—influenced by the afore-
mentioned factors—nevertheless, could account for
our ability to accurately process temporal intervals.
Especially for shorter durations up to a few seconds,
humans can accurately synchronize their movements to
regular beats (Mates et al. 1994), discriminate tones
with different durations (Rammsayer & Lima 1991)or
reproduce presented intervals ( Eisler & Eisler 1994).
Yet, there is no consensus as to which temporal
mechanisms account for these temporal-processing
abilities. Over the last decades, the most successful
models for such a mechanism have been variants of a
pacemaker–accumulator clock, where an oscillator
(a pacemaker) produces a series of pulses (analogous
to the ticks of a clock) and the number of pulses
recorded over a given timespan represents experienced
duration (Po¨ ppel 1971;Church 1984;Treisman
et al. 1990;Meck 1996;Zakay & Block 1997).
However, competing models assume neuronal system
properties for encoding duration not related to a simple
pacemaker–accumulator system (Matell & Meck 2004;
Wackermann & Ehm 2006;Karmarkar & Buonomano
2007), or they propose that memory decay processes
are involved in time perception (Staddon 2005;
Wackermann & Ehm 2006). Related to this unsolved
issue, the question of which areas of the brain
process duration has also not yet been answered
definitely. Among other regions, most prominently,
the cerebellum (Ivry & Spencer 2004), the right
posterior parietal cortex (Bueti et al.2008a), the
right prefrontal cortex (Rubia & Smith 2004;
Lewis & Miall 2006) as well as fronto-striatal circuits
(Harrington et al. 2004a;Hinton & Meck 2004)have
been implicated as the neural substrates of a potential
timekeeping mechanism.
In summary, as this brief outline suggests, there is
still considerable uncertainty on how (regarding
psychological and neurophysiological models) and
where in the brain time is processed. This paper has
several goals that are related to the issues raised: (i) to
give an answer to the question of why so many different
brain regions have been assigned to the neural basis for
our experience of time, i.e. to explain some of the
variation found across models, and (ii) to describe
recent developments in the research on time percep-
tion, which are indicative of a strong involvement of
emotions and mood states. These developments could
be described as an ‘emotive turn’ in this area of
research, which might follow decades of focusing on
cognitive aspects of time perception. (iii) Recent
conceptualizations and empirical findings, which have
led to this emotive turn, might develop into a neural
theory of time perception that will encompass sub-
jective feeling and interoceptive (within the body)
states. Specific neurophysiological processes in circum-
scribed regions of the brain, as they are related to these
feeling states, might constitute a mechanism for
encoding duration.
2. COGNITIVE AND NEURAL MODELS OF
TIME PERCEPTION
Investigations in the fields of experimental psychology,
clinical neuropsychology and neuroimaging have
resulted in an extensive literature on the mechanisms
and underlying neural systems of temporal processing.
Over the last decades, certain cognitive and neural
models have dominated time perception research, but
alternative models exist and the number of potential
theories has to date increased considerably. To
summarize the status of the research field in general
terms: there is a lot of conflicting evidence and there are
several competing conceptualizations. In the following,
a short review of (i) predominant cognitive models
of time perception and (ii) related neural theories of
interval timing is provided. These models and theories
will be contrasted with alternative conceptualizations
and empirical evidence in order to provide an overview
of the heterogeneity of ideas concerning mechanisms of
time perception.
Cognitive models distinguish two fundamental
perspectives in time estimation: prospective and retro-
spective time estimation ( James 1890;Zakay & Block
2004).
2
In the former, an observer judges the duration
of an interval that is being presently experienced. In
retrospective time estimation, by contrast, an observer
estimates a timespan that has already been passed and
to which he is only now paying attention. Models of
prospective time estimation assume an ‘internal clock’
with a pacemaker producing a sequence of time units
that are fed into an accumulator (Church 1984;
Treisman et al. 1990). In a variant of these pace-
maker–accumulator models, the attentional-gate
model (Zakay & Block 1997), the time units produced
are only registered when attention is directed to time.
Thus, prospective timing is always a dual task since an
observer has to divide attention between temporal and
non-temporal processes (Grondin & Macar 1992;
Taatgen et al. 2007). The number of units that have
been recorded during a physical time period (being
stored in working memory) is then compared (in a
decision process) with long-term memory of stored
representations of time periods, which can be verbalized
as seconds or minutes (Pouthas & Perbal 2004;Wea r d e n
2004). Thus, in addition to the pacemaker (the actual
clock component), several cognitive processes such as
working memory, long-term memory, attention and
decisions are involved in prospective time perception.
In retrospective time estimation, the duration of a
time interval that has already elapsed has to be judged.
Then, an observer has to estimate a given duration in
retrospect from the amount of processed and stored
memory contents; that is, duration has to be recon-
structed from memory (Ornstein 1970;Flaherty et al.
2005;Noulhiane et al. 2007b). The more changing
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experiences we have during a certain timespan—which
are stored in memory and later retrieved—the longer
the duration is subjectively experienced (Bailey & Areni
2006). In retrospect, routine activity when compared
with novel activity leads to the perception of shorter
time intervals (Avni-Babad & Ritov 2003). Thus, the
subjective impression of a long time interval depends
on the activity of a person with diverse experiences and
recruits the activation of areas such as the medial
temporal cortex, known to be involved in episodic
memory ( Noulhiane et al.2007b). Retrospective
duration judgements are based on temporal intervals
spanning a few seconds (short-term memory) to, in
principle, a whole lifetime (long-term memory)
(Wittmann & Lehnhoff 2005). Pure prospective
duration judgements, by contrast, are only conceivable
over a limited and shorter time range where an observer
attends to time for a period of seconds to minutes.
Prospective timing studies in animals and humans
have yielded the general hypothesis that fronto-striatal
circuits consisting of recurrent loops between frontal
cortex (SMA), caudate–putamen, pallidum and
thalamus, which are modulated by the dopamine
system, are critical for the processing of duration
(Hicks 1992;Harrington et al. 2004a). Support for the
anatomical hypothesis comes from investigations in
patients with brain lesions and from neuroimaging
studies. For example, individuals with structural
damage to the frontal lobes ( Nichelli et al. 1995;
Kagerer et al. 2002) or traumatic brain injury
predominantly affecting frontal areas (Pouthas &
Perbal 2004) show impaired estimates of temporal
intervals. Neuroimaging studies with healthy volun-
teers find that the perception of duration is linked to
activation in right prefrontal and striatal regions
(Ferrandez et al. 2003;Nenadic et al. 2003;Coull
et al. 2004). Regarding the involvement of neurotrans-
mitter systems, D2 dopaminergic antagonists (such as
haloperidol) impair duration discrimination abilities in
healthy subjects (Rammsayer 1999). Moreover, studies
with animals and humans indicate that both dopamin-
ergic agonists, e.g. methamphetamine, and antagonists
influence timing processes, presumably by increasing
and decreasing clock speed, respectively (Mohs et al.
1980;Buhusi & Meck 2002;Cevik 2003). Individuals
dependent on cocaine and methamphetamine, who
have abnormal brain metabolism and structural
changes involving dopaminergic target areas such as
the striatum and the frontal cortex, exhibit impaired
time processing on several timing tasks ( Wittmann
et al. 2007b). Additional evidence for the involvement
of the dopamine system in time perception comes
from patients with Parkinson’s disease who have
decreased dopaminergic function in the basal ganglia
and show deficits discriminating and reproducing
temporal intervals ( Hellstrom et al. 1997;Harrington
et al. 1998). Further evidence comes from a study
in which subthalamic deep brain stimulation in
patients with Parkinson’s disease levelled temporal
reproduction performance to that of control subjects
(Koch et al. 2004). Thus, intact dopamine neurotrans-
mission within striatal and frontal areas of the brain is
an important prerequisite for accurate temporal-
processing abilities.
However, some neuroimaging studies, revealing
areas of activity while subjects estimate durations or
time their movements, suggest that a core temporal-
processing mechanism is located in the right prefrontal
cortex (dorsolateral and ventrolateral areas) for both
sub- and supra-second intervals (Rubia et al. 1998;
Brunia et al. 2000;Lewis & Miall 2003a;Smith et al.
2003). In these (and other) studies, the basal ganglia
did not show activation, thus leading to the conclusion
that dopamine modulation in right prefrontal areas
might be the basis for a primary timekeeping
mechanism (Lewis & Miall 2006). Furthermore, and
pointing to yet another different brain region, since
patients with cerebellar dysfunctions are impaired in
the precise timing of movements (Ivry et al. 1988)as
well as in the sensory discrimination of duration
(Ivry & Keele 1989), the cerebellum seemingly plays
an important role in the processing of time. It has been
speculated that separate, i.e. non-overlapping, neural
elements in the cerebellum that have different delay
properties could potentially encode duration (Ivry &
Spencer 2004). Last but not least, the right posterior–
inferior parietal cortex has been implicated (i) in
integrating duration information as represented in the
sensory modalities and (ii) as an interface between
the perception of duration and timed actions (Bueti
et al. 2008a,c). Repetitive transcranial magnetic
stimulation (rTMS) selectively disrupts duration
discrimination in the visual and auditory modality if
the respective sensory cortices are affected, but
stimulation of the right parietal cortex disrupts timing
in both the visual and auditory domains (Bueti et al.
2008a,b). Bueti and colleagues thus support the idea of
distributed, modality-specific, timekeeping processes
that converge in the parietal cortex. Neurophysiological
studies in monkeys complement these findings by
showing that specific increasing (ramp-like) neuronal
activity in the posterior parietal cortex encodes
duration (Leon & Shadlen 2003). However, similar
neural activity in the monkey brain and related to the
timing of events has been recorded in the premotor and
motor cortex as well (Lebedev et al. 2008).
One difficulty in deciding on which regions are the
primary target areas for a suspected timekeeping
mechanism comes from neuroimaging studies where,
typically, multiple brain regions show activation during
time perception tasks ( Lewis & Miall 2003b). The
involvement of all these brain areas is probably due to
different cognitive processes not subtracted out in the
contrasts between primary time perception and control
tasks, the principal method used in functional
magnetic resonance imaging (fMRI ). Many active
areas are not primarily related to the encoding of
duration (i.e. an internal clock) but, nevertheless, take
part in a complex timing system representing atten-
tion, working memory and decision-making processes
(Rubia & Smith 2004;Livesey et al. 2007).
A consequence of the complex architecture of an
assumed timekeeping system is that disruptions in any
component can result in timing impairments. That
could explain why so many different patient groups
with damage to the brain, i.e. lesions or degenerative
processes in the basal ganglia, cerebellum or right
parietal, as well as frontal cortex, can be impaired
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(Wittmann 1999). Likewise, it cannot be clearly
decided whether experimental disruptions with
rTMS are due to the transient impairment of a neural
clock or of other processing components. For example,
rTMS applied to the dorsolateral prefrontal cortex
reliably impairs temporal reproductions of a few
seconds as well as multiple second intervals, but it is
unclear whether this accounts for the disruption of a
neural clock or of working memory ( Koch et al.
2003,2007;Jones et al. 2004). rTMS applied to
the cerebellum transiently impairs the temporal
reproduction of intervals with durations of approxi-
mately half a second ( but not approx. 2 s), a finding
that could be interpreted by the disruption of either
a millisecond timer or of a motor programme involved
in millisecond timing ( Koch et al. 2007).
Two further complicating factors in this discussion
arise from neuropsychological studies with brain-injured
patients. First, some studies report negative findings in
which patients do not differ in their timing abilities from
control subjects. Second, timing deficits, if registered,
although significant from a statistical point of view, are
not necessarily dramatic. For example, and contrasting
earlier findings, patients with cerebellum lesions
following a stroke were not impaired in a duration
discrimination task and only mildly impaired in a motor-
timing task (Harrington et al. 2004b). Patients with focal
basal ganglia lesions, although impaired in performing
with maximum tapping frequency, were as accurate in
motor timing as were controls (Aparicio et al.2005).
Moreover, although these patient groups, as shown in
several studies, on average, and according to the
inferential statistics, differed from the performance of
control subjects, the deviations were often minimal
(Ivry & Keele 1989;Harrington et al. 1998;Kagerer
et al. 2002;Wittmann et al. 2007b).
One suggested solution to the problem of not being
able to pin down specific regions of the brain as
representing a clock-type timing mechanism is to
assume distributed neural networks where neural
populations within each region would encode duration
(Mauk & Buonomano 2004). If several neural units
would possess intrinsic temporal-processing proper-
ties, many different brain areas could contribute to the
perception of time depending on the modality and
the type of task. These network or state-dependent
models do not incorporate a dedicated timing system
(with a centralized clock) but rather time-dependent
neural changes, such as short-term synaptic plasticity
(Karmarkar & Buonomano 2007). However, such
intrinsic mechanisms would be limited to short time
intervals up to several hundred milliseconds. Time
perception in the range of seconds would still require
additional processes (Ivry & Schlerf 2008). Similarly, it
is conceivable that temporal-processing functions are
embedded in several specialized and interacting neural
circuits, where the timing function would not be the
primary process in those brain regions (through a
dedicated mechanism for temporal processes) but, for
example, motor systems would implicitly be involved in
the temporal processing of intervals ( Nobre & O’Reilly
2004). Alternatively, an idea that has been very recently
stated suggests that the effort made when we perceive
and act might determine experienced duration
(Marchetti 2009). An intuitive example fitting this
idea is the phenomenon that novel experiences seem to
last longer than routine events, which can be explained
by the greater demand of mental activities involved in
performing a task or analysing a situation for the first
time. The idea of mental effort as a determinant of time
experience is compatible with the notion of ‘fluency’,
the subjective experience of ease or difficulty of a
mental task (Oppenheimer 2008). The experience of
time, according to this view, would not rely on clock
processes but would be an epiphenomenon evolving
from cognitive and emotional responses during a time
interval, where a cognitively demanding task (a filled
interval) and doing nothing (an empty interval, i.e.
when sitting in the doctor’s waiting room) would
likewise lead to the impression of a slow passage of time
(the act of waiting without any distraction can be filled
with painful emotions) ( Flaherty 1999).
Further conceptualizations, which argue against the
idea of dedicated timing systems, add to the diversity of
time perception models. Just as is argued for the
retrospective perspective on duration, prospective
timing could be governed by memory processes, where
the decreasing memory strength over time, specifically
modelled using leaky integrators, leads to the
impression of passing time and, eventually, of duration
(Staddon 2005). In a different model, the ‘dual
klepsydra’ model, inflow and outflow processes of a
leaky accumulator form properties of a timekeeping
mechanism (Wackermann & Ehm 2006). Subjective
duration is defined by the state of a leaky integrator
that depends on inflow units (accumulation over time
during the encoding of stimulus duration) and outflow
units (loss over time). Parameters of that model fit
well to individual temporal reproduction responses
as well as to changes in the timing of behaviour
induced by experimental manipulations ( Wackermann
et al. 2008).
In summary, a multitude of incompatible conceptu-
alizations exists on how time is processed. Furthermore,
there is no agreement on which brain areas or brain
systems—dedicated to time or not—might underlie our
perception of duration. The important conclusion so
far is that different processing components are involved
when we perceive duration or time our movements,
which are not related to an internal clock. Nevertheless,
through experimental manipulation or in certain
patient populations, changes in these processes can
affect perception and behaviour related to the time
domain. In the following, one further factor will
be discussed that may explain some of the variation
found in studies on time perception, i.e. why so
many different brain regions have been assigned with
a central timekeeping function. The main point will be
that different neural systems are involved in temporal
processes (and related experiences) depending on the
duration of the processed interval.
3. TIME SCALES IN THE PERCEPTION OF TIME
It is intuitively most unlikely that one mechanism or
one neural system would be responsible for all possible
durations that an organism has to process. Different
temporal processing mechanisms must be involved on
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different time scales ( Trevarthen 1999;Wittmann
1999;Mauk & Buonomano 2004;Buhusi & Meck
2005). Experimental interventions have repeatedly
shown duration-specific effects of psychopharmacolo-
gical agents on interval timing. For example, the
dopamine receptor antagonist haloperidol as well as
the benzodiazepine midazolam impair duration
discrimination of intervals ranging approximately 1 s,
whereas processing of 50 ms intervals is affected by
haloperidol only (Rammsayer 1999). According to
these results, any pharmacological treatment that
affects working memory capacity (e.g. midazolam)
would interfere with temporal processing of intervals
above 1 s. However, intervals with a length of up to a
few hundred milliseconds (such as the 50 ms interval)
are supposed to be processed based on brain
mechanisms outside of motor and cognitive control
and reflect pure timing processes (thus, not being
influenced by midazolam). According to this view,
additional processes such as attention and working
memory, not related to time per se, come into play with
intervals exceeding several hundred milliseconds in
length and complementing dopamine-driven pro-
cesses, which are involved both in shorter and longer
time intervals ( Rammsayer et al. 2001). A similar
theoretical proposal, derived from a meta-analysis on
neuroimaging data, suggests two distinct neural timing
systems: (i) an automatic timing system for shorter
intervals up to approximately 1 s, which recruits motor
systems of the brain (SMA, basal ganglia and
cerebellum) and (ii) a cognitively controlled timing
system for supra-second intervals connected mainly to
right prefrontal and parietal cortical areas (Lewis &
Miall 2003b). This separation of time perception
systems is to some extent mirrored by findings
in motor-timing studies, where qualitative changes in
tapping performance occur with inter-tap intervals of
approximately 1 and 1.5 s duration (Madison 2001).
Time ranges between 0.45 and 1.5 s seem to be
automatically processed, i.e. not strongly affected by
attentional demands, whereas attention and working
memory processes (stimulated by secondary tasks)
affect intervals in the range between 1.8 and 3.6 s
(Miyake et al. 2004).
The notion that perception and motor behaviour are
processed in discrete windows or processing epochs has
been conceptualized for some time ( White 1963;
Po¨ppel 1970;Dehaene 1993;VanRullen & Koch
2003;Fingelkurts & Fingelkurts 2006). These
temporal integration units fuse successive events into
a unitary experience, ‘snapshots of experience’ or
‘psychological presents’ (Blumenthal 1977), which
are characterized by co-temporality, meaning that
events within this time zone have no temporal
relationship (Ruhnau 1995). For example, the percep-
tion of temporal order of short stimuli in different
modalities is only possible if the individual events are
separated by at least 20–60 ms ( Exner 1875;Hirsh &
Sherrick 1961;Kanabus et al. 2002;Fink et al. 2006).
If the two events are separated by smaller intervals, an
observer is not able to tell which of the two appeared
first. Moreover, since stimulus properties of speech
stimuli are perceptually segmented into these
sequential processing units, temporal information
within a segment of a speech sound may not be
relevant for decoding (Kiss et al. 2008). A longer time
frame of approximately 200 ms determines the integra-
tion of auditory–visual input in speech processing
(van Wassenhove et al. 2007). Fusion percepts were
reported if the onset of lip movements and heard
syllables did not exceed this time lag. Threshold values
of approximately 250–300 ms have long been
suggested to represent a specific integration process
in perception (Mu
¨nsterberg 1889;White 1963).
A minimum stimulus duration (or minimum inter-
onset interval) of 200–300 ms is necessary for detecting
the temporal sequence offour acoustic events (Warren &
Obusek 1972;Shrivastav et al. 2008)andforoptimal
effects in an oddball task that leads to the subjective
expansion of duration (Tse et al.2004). Temporal
integration mechanisms in a time frame of approximately
250 ms also seem to be involved in sensorimotor
processing, distinguishing maximum tapping speed
from a personal, controlled motor speed (Peters 1989;
Wittmann et al. 2001).
On a different time scale, a perceptual mechanism
seems to exist that integrates separate successive events
into a unit or perceptual gestalt (see Po¨ ppel 2009). We
do not just perceive individual events in isolation, but
automatically integrate them into perceptual units with
a duration of approximately 2–3 s ( Fraisse 1984;
Po¨ ppel 1997). For example, while listening to a
metronome at a moderate speed, we do not hear
a train of individual beats, but automatically form
perceptual units, such as 1–2–3, 1–2–3, etc. These are
mental constructs—physically speaking, they do not
exist. The duration of this temporal integration
mechanism, referred to as the subjective present,
seems to be limited to 2–3 s (Szelag et al. 1996;
Wittmann & Po¨ppel 2000). Phenomenological
approaches have revealed that temporal intervals
shorter than 2 s are perceived qualitatively as different
from longer intervals (Benussi 1913;Nakajima et al.
1980). Typically, intervals up to 2–3 s are reproduced
accurately, whereas longer intervals tend to be under-
estimated ( Ulbrich et al.2007;Noulhiane et al. 2008). In
one study measuring event-related potentials (ERPs)
during the reproduction of visual stimuli ranging in
duration between 1 and 8 s, accurate reproductions up to
3 s were accompanied bya contingent negative variation-
like slow negative shift in the ERP signal. This shift was
reduced or absent when durations exceeded 3 s (Elbert
et al. 1991). Duration discrimination thresholds with
base durations up to 2 s show constant Weber fractions
(Dt/t, where Dtisthe difference between the base duration
tand the length of the comparison interval at which a
duration difference is just noticeable), but with longer
durations, Weber fractions rise rapidly
3
(Getty 1975).
Subjects can accurately synchronize their motor actions
to a sequence of tones presented with a frequency of
approximately 1–2 Hz. The ability to synchronize these
tones becomes more difficult with increasing inter-tone
intervals and finally breaks down when intervals exceed
durations of approximately 2–3 s (Mates et al.1994;
Miyake et al. 2004). With respect to even longer intervals,
it is also conceivable that distinct processes are involved
for different durations. For example, the estimation of a
1 h interval seems to be related to the duration of wake
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time of an individual, whereas the estimation of seconds
and minutes is related to body temperature (Aschoff
1998).
Experimental interventions selectively affect
intervals of specific time ranges, i.e. the psychopharma-
cological substance psilocybin has an effect on duration
longer than 2 s ( Wittmann et al. 2007a), rTMS of the
cerebellum affects solely millisecond timing and rTMS
of the right dorsolateral frontal cortex only affects
timing in the seconds range ( Jones et al. 2004;Koch
et al. 2007). Nevertheless, a reliable correlation
between time scales and neural systems has not been
accomplished so far. Also, individual neuroimaging
studies often find similar brain areas activated in time
perception tasks employing different durations
(Pouthas et al. 2005;Jahanshahi et al. 2006).
To summarize §§ 2 and 3, several factors contribute
to the diversity of different models on the question of
how (which mechanism) and where (which neural
system) in the brain duration is processed. First, there
are several cognitive processes that are entangled with
our perception of time (independent of a potential
internal clock), which require the integrative proces-
sing of multiple modules distributed across the brain.
Experimental manipulations of task load (in attention
and working memory, for example), clinical studies
with selected patients with dysfunctions in different
brain areas (i.e. cerebellum, basal ganglia, frontal and
parietal cortex) as well as neuroimaging studies show
that many parts of the brain and multiple cognitive
processes contribute to the perception of duration.
Moreover, there is no single area of the brain on which
functioning our temporal experiences would
completely rely on; that is, patients with damage to
the brain may be affected in the processing of duration
but their performance hardly ever breaks down
completely. Second, different neural mechanisms are
most probably responsible for temporal processes and
time experiences depending on the duration involved.
4. THE EMOTIONAL EXPERIENCE OF TIME
Time does not pass with a steady-paced flow.
Perceptual time is not isomorphic to physical time,
meaning that the subjective passage of time and
estimates of duration vary considerably. In uneventful
or unpleasant situations, such as when nervously
waiting for something to happen, we experience a
slower passage of time and overestimate its duration.
By contrast, if we are entertained and focus on
rewarding activities, time seems to pass more quickly
and duration is more likely to be underestimated.
These examples of time judgements are inherently
emotional. They point to an aspect of time perception,
although part of everyday experience, which has been
neglected in research over the past few decades. Only
very recently, a body of evidence has accumulated
which is indicative of the intricate interplay between
mood states and perceived duration. In most cases, the
influence of emotions (often leading to longer time
estimate) is explained by the standard cognitive model of
prospective time perception in which emotions
affect the degree of attention to time or increased physio-
logical arousal levels lead to a higher pacemaker rate
(Droit-Volet & Meck 2007;Wittmann & Paulus 2008).
Both increased attention to time and a higher pace-
maker rate of an assumed internal clock would lead to
the accumulation of more temporal units during a
given timespan.
Regarding durations of multiple seconds to minutes,
it has been shown that patients with depression (Bschor
et al. 2004) and cancer patients with high levels of
anxiety (Wittmann et al. 2006) report a slowing down
of the pace of time and overestimate temporal intervals.
It would appear that the psychological distress these
individuals suffer from directs attention away from
meaningful thoughts and actions to the passage of time.
Also, boredom-prone individuals estimate time inter-
vals to last longer than persons with low levels of
boredom (Danckert & Allman 2005). Similarly,
subjects who were socially rejected in a psychological
experiment overestimated intervals of multiple
seconds, a finding that was interpreted as stemming
from a state of reactive emotional and cognitive
deconstruction, which in turn resulted in a stronger
attentional focus on the present ( Twenge et al. 2003).
Positive correlations of general anxiety levels with
duration estimates of multiple seconds have been
reported in undergraduate students (Siegman 1962)
and in patients with psychiatric diagnoses (Melges &
Fougerousse 1966). Spiderphobics who had to look at
spiders for 45 s also overestimated this exposure time
when compared with controls (Watts & Sharrock
1984). These overestimations were interpreted as
resulting from an arousal-induced increase in an
internal pacemaker. Moreover, emotional stress such
as the anticipation of a mild electric shock ( Falk &
Bindra 1954), when compared with control conditions,
led to an overestimation of duration.
Generally speaking, time distortions are stress
related as they are often experienced during dangerous
or life-threatening situations such as road accidents or
encountered violence (Hancock & Weaver 2005). In
movies, scenes depicting combat are sometimes shown
in slow motion to portray what the involved protagonist
subjectively experiences. To date, two studies
experimentally tested this phenomenon. Novice sky-
divers, for example, who reported a stronger fear
during their first tandem jump, also experienced its
duration (subjective estimates approx. 30 min) as
lasting longer than did less fearful novice skydivers
(Campbell & Bryant 2007). A similar time dilation
effect was also seen on a shorter time scale. Subjects
who experienced a free fall of 31 m before they landed
safely in a net overestimated retrospectively the
duration of that fall when compared with when
watching others fall (Stetson et al. 2007). However, a
slow-motion effect was not detected when probed with
a special chronometer that had to be watched during
the fall. Even in more conventional and less frightening
experimental situations, subjects overestimate the
duration of high arousing pictures with emotional
valence (depicting angry faces or accidents), which
last only several hundred milliseconds to a few seconds
(Angrilli et al. 1997;Droit-Volet et al. 2004;Gil et al.
2007). These effects seem to be strongly tied to the
embodiment of emotions. Participants seem to over-
estimate emotional faces only when they are able to
1960 M. Wittmann Review. The inner experience of time
Phil. Trans. R. Soc. B (2009)
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spontaneously imitate the perceived emotions on the
faces which they have to judge, thus showing that affect
is embodied (Effron et al. 2006). To be more precise,
however, paradoxical effects can occur in the way that
emotional stimuli can cause over—as well as under-
estimations of time. For example, in one study (Angrilli
et al. 1997), low-arousing and emotionally negative
pictures led to an underestimation of duration (inter-
preted as resulting from the subjects’ distraction from
time and attention to the emotional content of the
stimuli), whereas high-arousing emotional pictures led
to an overestimation of duration (interpreted as
resulting from an increased pacemaker). In another
study (Noulhiane et al. 2007a), unpleasant sounds
(e.g. sobs, a crying women) were judged to last longer
than pleasant (e.g. laughs) or neutral stimuli. This
finding was again interpreted through an increase in
pacemaker rate. Furthermore, and contrasting with the
reported findings above, high-arousing sounds were
judged to be shorter than low-arousing sounds
(interpreted as resulting from the distraction from time).
In summary, increased attention to time (such as in
waiting situations) and an increase in physiological
arousal (such as under stress) can lead to longer time
estimates when judging intervals in the range of
milliseconds to seconds and minutes. However,
paradoxical effects of an underestimation of duration
in emotional situations can occur, which are discussed
as stemming from a distraction from time. It is difficult
to decide which mechanism, attention related or
activation induced, actually affects the sense of time
(and in which direction). Physiological measurements
would have to complement the employed time
perception measures. For example, in one study,
acute marijuana effects in healthy subjects corre-
sponded with underproductions of time intervals
(interpreted as acceleration of a pacemaker) and an
increase in heart rate, indicative of an increase in
physiological arousal (Tinklenberg et al. 1976). The
two proposed mechanisms influencing time experience
are not necessarily exclusive but could contribute in an
additive way (Burle & Casini 2001). For example,
overestimations of time intervals of approximately
1 min in duration as detected in many impulsive
patient groups could be due to an increase in
physiological arousal as well as caused by an increase
in attention to time (Berlin & Rolls 2004;Wittmann
et al. 2007b).
The impact of our emotional states on the
experience of time is usually discussed in the framework
of the standard cognitive model of prospective time
perception, which proposes a pacemaker–accumulator
that is embedded in a system of cognitive components.
However, on reviewing the empirical evidence and as
argued above, the basic questions of how our sense of
time is created are still unresolved. The recent upsurge
in empirical investigations on the relationship between
emotions and time, perhaps signifying an emotive
turn, might result in a new psychological and neural
theory of time perception. So far, emotions and
physiological states, similar to cognitive functions such
as attention and working memory, have been treated as
modulators of an assumed neural clock. What if
mood processes and the representation of body
sensations themselves function as a timekeeper? Since
emotions and physiological states seem so fundamental
to the experience of time, it is tempting to assign a
pivotal role to these processes related to a core time-
keeping system.
Since the late nineteenth century, based on the
theory by James and Lange, it has been argued that
affective states as well as experienced emotions are
inseparable from autonomic responses (e.g. cardiovas-
cular activity, abdominal sensations and breathing
patterns; Saper 2002;Pollatos et al. 2005). According
to this notion, bodily signals, visceral and somatosen-
sory feedback from the peripheral nervous system,
enact emotions (Damasio 1999;Wiens 2005).
For example, people’s heartbeat detection ability is
positively related to subjective ratings of emotional
pictures as more arousing (Pollatos et al. 2007a).
Interoceptive awareness, as tested with heart rate
detection tasks, is predicted (among other regions) by
right anterior insula activity (Critchley et al. 2004;
Pollatos et al. 2007b). The insular cortex of primates,
structurally embedded in the extended limbic system,
is considered as the primary receptive area for visceral
input, i.e. for physiological states of the body (Craig
2002;Saper 2002). The capacity for the awareness of
emotions is probably built on the anatomical organi-
zation and a progressive integration of information in
the insula (Craig 2003). A posterior-to-anterior
progression of representations in the human insular
cortex is the basis for the sequential integration of body
states and internal autonomic responses with cognitive
and motivational conditions, the latter being instan-
tiated by distributed neural processes across the brain.
This progression culminates in the anterior insula and
leads to the conscious awareness of complex feeling
states. A direct link between the perception of time and
physiological processes has been proposed by Craig
(2008), who claims that our experience of time relates
to emotional and visceral processes because they share
a common underlying neural system, the insular cortex
and the interoceptive system. He suggests that the
insula, through the temporal integration of signals from
within the body, produces a series of ‘emotional
moments’ in time. The perception of duration there-
after would be defined by these successive moments of
self-realization, formed by information originating
within the body (see also Craig 2009).
Several attempts have been made to directly relate
rhythms of the body to temporal processes in perception
and action. For example, Mu
¨nsterberg (1889), being his
own subject, reported that when the onset and the offset
of temporal intervals to be reproduced and ranging in
duration between 6 and 60 s coincided with him starting
to inhale, his temporal reproductions were more
accurate than when temporal intervals started at
other points in time not systematically related to his
breathing cycle. Mu
¨nsterberg, who did not count in
his experiments, concluded that the sense of time
relied on the sensation of tension in different organs
which are caused by muscle contractions. Fraisse
(1982) highlights findings showing that the period-
icities of the heart, of walking, of the preferred tapping
tempo as well as of preferred acoustical rates are of
the same order of magnitude of 500–700 ms. However,
Review. The inner experience of time M. Wittmann 1961
Phil. Trans. R. Soc. B (2009)
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he concludes that ‘we cannot assume that one
phenomenon can be explained by the other. There is
only a narrow range of frequencies of natural or
voluntary rhythms and of preferred tempo’ ( p. 154).
Despite this conclusion, however, one study showed
that participants’ preferred tempo of successive tones
was in a harmonic relation (with a ratio of 1, 1.5 and 2) to
individual heart rates as measured during the presen-
tation of the tone sequences (Iwanaga 1995).
Other studies, which attempted to relate heart rate
(in some cases also breathing rate) to time estimates in
the range of several seconds to minutes, some testing
subjects before and after physical exercise, found no
relationship (Schaefer & Gilliland 1938;Bell & Provins
1963) or associations that were weak or difficult to
interpret (Lediett & Tong 1972;Osato et al. 1995).
One study, nevertheless, employing shorter intervals
with duration up to 4 s, found that drugs, which
either stimulate or inhibit the central or peripheral
sympathetic nerve system, lead to an increase or
decrease in heart rate and breathing rate and an
accompanying relative under- or overproduction of
intervals, respectively ( Hawkes et al. 1962). A relative
underproduction of intervals can be interpreted as
resulting from an increase in clock speed. Although this
latter result points to the role of cardiac and respiratory
cycle rates in the judgement of duration, one has to
keep in mind that proponents of the standard cognitive
models of time perception could still argue that it is not
the heart rate per se that functions as an internal clock
but that generally increased arousal levels affect
the pacemaker of an internal clock in the brain
(Wittmann & Paulus 2008). Especially regarding the
short durations tested in the study by Hawkes et al.
(1962) of just a few seconds, a heart rate cycle of
typically approximately 700 ms could not accurately
represent different temporal intervals.
Although hardly any convincing evidence exists that
would show how specific physiological cycles function
as an internal clock for judging time, body states as a
whole could, nevertheless, form the building blocks of a
timekeeping mechanism. Ultimately, interoceptive
processes as registered in the insula encompass the
physiological status of all body tissues and organs such
as the skin, muscles and the viscera (Craig 2008). It has
beenshownthattheself-regulationofemotions
(Vohs & Schmeichel 2003) and of autonomic para-
meters in biofeedback procedures (Cohen 1981) lead
to longer time percepts. One attempt to explain these
and other findings is that the insular cortex, which
integrates body signals, is the anatomical basis for the
creation of emotions and the sense of time (Craig
2008). Being more strongly aware of one’s own
emotions and body processes would, at the same
time, lead to a prolongation of subjective duration.
When individuals are experimentally deprived of
sensory stimuli (auditory–tactual–visual) their overall
sense of duration over several days gets strongly
impaired. However, these subjects, who have to rely
solely on their inner sense, report that time passes
painfully slowly (Schulman et al. 1967). Experienced
practitioners of mindfulness meditation who concen-
trated ‘on the self across time and the present moment’
in an fMRI study showed stronger activity in a
right lateralized network including the insula and
somatosensory cortex (Farb et al. 2007).
An explicit assignment to a functional role for the
insula in temporal processing was made by Ackermann
et al. (2001), who showed in an fMRI study that a linear
increase in left anterior insula activity was a function of
presented click train rates (increasing up to 6 Hz). By
contrast, a linear increase in right anterior insula
activity was recorded when click rates slowed down to
2 Hz. This and other findings have led to the idea that
the insula is an essential component in the sequencing
of sounds and the perception of rhythm in music
(Bamiou et al. 2003). Moreover, an involvement of the
insular cortex has repeatedly been shown in neuroima-
ging studies on duration processing with different
timing tasks (Brunia et al. 2000;Rao et al. 2001;
Lewis & Miall 2003b;Pouthas et al. 2005;Livesey et al.
2007;Stevens et al. 2007); however, the significance of
insula activation in the context of time perception is
seldom discussed. Recently, however, evidence of
neurophysiological activity in the posterior insula
of the human brain has shown to be involved in the
encoding of multiple-second durations. Time–activity
curves of neural activation derived from event-related
fMRI during a time reproduction task showed acti-
vation in bilateral posterior insula that linearly built up
when subjects were presented with 9 and 18 s tone
intervals ( Wittmann et al. 2008). This build-up of
neuronal activation peaked at the end of the respective
intervals. Related to the functional role of the dorsal
posterior insula, this finding of accumulator-type
activity was, therefore, interpreted as representing the
registration of physiological changes over time that
eventually leads to the representation of duration. The
flow of time, thereafter, corresponds to the flow of
interoceptive signalling from the body as sensed in the
dorsal posterior insula.
A shift in attention to emotional processes and
insular cortex activity in the search for the underlying
mechanisms of time perception has only recently
started. The greatest obstacle to a sound theory on
the neural bases of time perception has been the lack of
evidence for a neurobiological mechanism. The finding
on accumulator-type activity during the processing of
multiple second intervals represents such a potential
mechanism. This finding is also compatible with the
pacemaker–accumulator model of time perception
where pulses emitted by a pacemaker are transiently
stored in an accumulator and the number of pulses
defines duration (Pfeuty et al. 2005). In line with this
conceptualization, it is conceivable that the number
and rate of body signals accumulated in the posterior
insula over a given timespan creates our perception of
duration. To disclose mechanisms on different time
scales, i.e. milliseconds and a few seconds, will be the
challenge for future research. Interval processing over
different time ranges may be controlled by different
neurobiological mechanisms.
5. CONCLUSION
Despite the fact that time is an essential factor for
understanding complex behaviour, the processes
underlying the experience of time and the timing of
1962 M. Wittmann Review. The inner experience of time
Phil. Trans. R. Soc. B (2009)
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action are incompletely understood. Too many contra-
dictory theories exist—in psychology and neuroscience
alike—that aim at explaining how we judge duration.
One could argue that the conceptualization presented
here of relating body processes to the experience of
time represents yet another alternative approach
among so many others. Only future empirical evidence,
based on strong theoretical grounds, will tell whether
a dedicated neural system actually exists for the
representation of time and which neurobiological
mechanism causes the experience of time.
One argument of plausibility as to why our inner
sense, our body awareness, might be related to
the sense of time should be mentioned here. The
recent proposal of an intimate relationship between
time and the awareness of an emotional and visceral self
(see Craig 2008 2009) seems to be in line with a
western philosophical tradition proposing that time is a
creation of the self. ‘Inner time or duration is virtually
indistinguishable from the awareness of the self, the
experience of the self as an enduring, unitary entity that
is constantly becoming’ (Hartocollis 1983, p. 17).
Thereafter, the experience of our self is only possible as
an entity across time. One attribute of the self is time,
or differently expressed, the self is defined by its
extension over time, a succession of moments that
constitutes duration.
4
This idea is probably most
explicitly expressed by Henri Bergson who noted that
‘psychic states [.] unfold in time and constitute
duration’(Bergson 1913). According to his view, the
phenomenal self creates the sense of duration.
In neurobiological terms, we perceive signals from
the body which create a ‘material me’ that has
subjective feelings and is self-aware (‘I feel it’s me!’;
Craig 2002). Since the ascending pathways to the
insular cortex inform us about the ongoing status of
the body, a pacemaker-type signal (accumulating
successive states of the material self) would be continu-
ously present that could be employed in a timekeeping
system. An accumulation of physiological states
over time would be registered in the insula. This sketch
of a processing model is, of course, borrowed from
the cognitive models with a pacemaker–accumulator unit
and is, to date, speculative. To summarize the main lines
of argument put forward in this review:
(i) Although the perception of time is an essential and
inextricable component of everyday experience,
no conclusive answers to the questions of which
neural substrates and what kind of neurophysio-
logical processes could account for the experience
of duration have been established. That is, several
areas of the brain have been identified as potential
contributors to timekeeping (e.g. cerebellum,
frontal cortices and basal ganglia), but none
have been specifically implicated for this process
and there is no consensus as to the precise neural
mechanisms accounting for our sense of time.
(ii) Philosophical wisdom beginning in antiquity has
related the experience of time to the feeling of a
self. The body self and emotional self in modern
biological terms is based on insular cortex activity.
The signalling of body states, which define the
material me and contribute to the ‘feeling me’, is a
permanent and ongoing process over time and,
thus, could function as an inner measure of
duration by matching external temporal intervals
with the duration of physiological changes.
(iii) The strong relationship between effect and time is
not only an everyday experience of everyone but it
is also well documented by many empirical
studies. Since affective states are entangled with
body states, insular cortex activity (the primary
sensory area for visceral signals) may, therefore,
cause the experience of the passage of time.
ENDNOTES
1
Book 11 of St Augustine’s Confessions: In te, animus meus, tempora
metior [.]ipsam metior, cum tempora metior (electronic edition: http://
ccat.sas.upenn.edu/jod/augustine/).
2
A vivid description of our experience of time from the prospective and
retrospective perspectives can be found in Thomas Mann’s novel The
magic mountain. In his chapter entitled Excursus on the sense of time,
Thomas Mann narrates some of the basic mechanisms of time
perception as described in the psychological sciences of today.
3
From the beginning of psychophysical investigations following
Fechner’s pioneering work, it has been know that, across the time
range between milliseconds and several seconds, the auditory
time sense does not follow the Weber–Fechner law, i.e. Dt/tis not a
constant (Mach 1865). Mach discovered that, in auditory duration,
discrimination, the highest acuity is achieved with base durations
approximately 300–400 ms.
4
In this context,Borges (1999), hintingat the existential aspectof time,
formulated: ‘Time is the substance of which I am made. Time is a river
that sweeps me along, butI am the river; it is a tiger thatmangles me, but
I am the tiger; it is a fire that consumes me, but I am the fire’ (p. 332).
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