CHRONOBIOLOGY INTERNATIONAL, 16( l), 17-32 (1999)

Minireview

TIME PERCEPTION AND TEMPORAL

PROCESSING LEVELS OF THE BRAIN
Marc Wittmann
Institut fur Medizinische Psychologie, Ludwig-Maximilians-Universitat
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Miinchen, Germany

ABSTRACT

A classification is presented to structure divergent empirical findings

on temporal mechanisms of the brain. Proceeding from our time experiences
of simultaneity, nonsimultaneity, temporal order, duration, and the subjective
present, a classification of thresholds is established that marks distinct pro-
cesses involved in time perception and the temporal control of movements.
On the basis of this classification, evidence has been collected that suggests
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that perception and action share common timing mechanisms. Furthermore,

brain structures involved in temporal aspects of behavior on different time
scales have been identified-namely, the cerebellum, the basal ganglia, and
circumscribed cortical regions. Another question under debate concerns the
representational mode in which time is represented in the brain; models sug-
gest neuronal oscillations or interval-based mechanisms. (Chronobiology Zn-
ternational, 16(1), 17-32, 1999)

Key Words: Basal ganglia; Cerebellum; Cortex; Duration judgment; lkmpo-

ral order; Temporal processing; Timing.

INTRODUCTION

Perception of events takes place over time. Motor actions are planned over time
and evolve over time. With this dynamic property of information processing explicitly
in mind, it seems plausible that there are timing mechanisms in the brain that structure
processes involved in perception and action. These processes determine the experience
of a time span or control the time interval between two related movements. Evidence
has actually been collected that refers to mechanisms on different time scales and to

Received July 14, 1998; returned for revision July 15, 1998; accepted July 15, 1998.
Address correspondence to: Institut fur Medizinische Psychologie, Ludwig-Maximilians-
Universitat Munchen, Goethestr. 3 1, 80336 Munchen, Germany. E-mail: nicosle@imp.med.
uni-muenchen.de

17
0 1999 International Society for Chronobiology www.dekker.com
Published by Marcel Dekker, Inc.
 18 WITTMANN

circumscribed neural systems. Several questions have evolved that form the core of re-
search studies that aim to identify subjective time and temporal aspects of neural pro-
cesses.
Especially for the perception of time, classifications have been formulated that
identify different processing levels that can be subjected to experimental inves-
tigation. Only with the theoretical basis of such a taxonomy do further explora-
tions of central issues in time research seem promising. A discussion of the
spectrum of time phenomena begins this review.
One of the main concerns of time research is outlined by the relationship be-
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tween perception and action. The question is whether the identified time levels
come into play both in perception and in action or are organized by separate
temporal processes.
0 To gain knowledge on the neural basis of temporal aspects in behavior, various
approaches have indicated involved brain structures and possible neural mecha-
nisms. The most striking findings come from clinical studies of patients with
injuries and malfunctions of the brain, from neuropharmacological studies, and
from intracortical recordings of neuronal activity in animals. One of the crucial
points in models of temporal processing is whether neural oscillations or other
representational mechanisms can be assumed.
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TAXONOMY OF LEVELS IN TEMPORAL PROCESSING

Two main concepts of time perception that correspond well to our subjective expe-
rience have often been described (Fraisse 1984; Block 1990)-succession and duration.
Perception of succession means that we can identify the sequential characteristic of
events, that is, their temporal order. Perception of duration applies to the persistence of
an event over time and to the interval between two events. On the basis of these two
concepts, a huge body of experimental results has accumulated. This evidence initially
appears to lack structure as the outcome is dependent on numerous factors, such as the
characteristics of the subject, the physical property of the stimuli, the contents of the
time period, task demands, and so on (Allan 1979; Nichelli 1993). Despite the divergent
factors, attempts have been made to identify numerical correspondences in experimental
results that refer to circumscribed processing levels. Starting with a taxonomy of elemen-
tary time experiences, the phenomena of simultaneity, successiveness, temporal order,
subjective present, and duration can be outlined (Poppel 1978, 1997). The range of phe-
nomena can be listed, beginning with a few milliseconds (relevant in the perceptual
differentiation between simultaneity and succession) and ending with decades (as in the
retrospective view of an individual’s life span).

Simultaneity, Succession, Temporal Order

With a simple experimental setting, two thresholds can be marked that separate the
experiences of simultaneity, nonsimultaneity, and temporal order: fusion threshold and
temporal-order threshold. If a subject is presented with two acoustic stimuli, such as
clicks via headphones-one to each ear or both to one ear with a certain interstimulus
interval-the correct physical order is not necessarily perceived. Simultaneity of two
 TIME PERCEPTION AND TEMPORAL PROCESSING 19

acoustic stimuli is detected with time intervals below approximately 2 to 3 ms. Only
above this temporal threshold, the so-called fusion threshold, are they perceived as tem-
porally separate (Hirsh 1959; Lackner and Teuber 1973; Moore 1993). The detection of
a temporal order of events, however, is only possible when events are separated by
intervals of at least 20 to 40 ms (Hirsh 1959; Steinbuchel 1995; Pastore and Farrington
1996). Dependent on the psychophysical methods used, thresholds vary, but are usually
situated in the given range; the maximum mean order threshold in literature for click
pairs was measured with 60 ms (Mills and Rollman 1980). Only above the order thresh-
old can a subject reliably indicate which of two stimuli came first. As approximately the
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same order threshold is assessed over different sensory modalities-clicks or tones in

audition, light flashes in vision, or tactile stimulation in the somatosensory domain (Hirsh
and Sherrick 1961)-a central time-organizing system has been postulated (Poppel 1994,
1997). In contrast, fusion threshold is considered more dependent on peripheral mecha-
nisms as it varies over sensory modality. On this temporal level, auditory fusion threshold
is lowest, somatosensory thresholds are slightly higher, and resolution in the visual sys-
tem has the least acuity (Poppel 1978; Hirsh and Sherrick [1961] discuss the results of
Exner’s [ 18751 results; more recent experimental findings on sensory-dependent differ-
ences can be found in Artieda and Pastor [1996]).
The results of a temporal-order threshold of 20 to 40 ms have been interpreted as
an indication of a discontinuous mode of the high-frequency processing of iriformation
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(Poppel 1994, 1997). Minimal time intervals of approximately 30 ms are required for the
identification of temporally segregated events. The underlying cortical processes could
reflect an organizational principle to generate elementary units of perception over time
in steps of 30 ms. Temporal-order identification is not totally independent of stimulus
parameters (Pastore et al. 1982), but thresholds seem to converge roughly with values in
the given range. One must, however, remember that this basic value is primarily restricted
to the detection of two successive events; second, the threshold is assessed with very
short stimuli (e.g., in audition, clicks of 1 ms). When more than two considerably longer
stimuli are presented, different processing capacities on a different time scale come into
play. Warren and Obusek (1972) discovered that subjects could only detect the temporal
order of four consecutively presented unfamiliar sounds when the duration of the single
stimuli exceeded 300 ms. Despite this finding, the temporal threshold of 30 rns remains
the lower limit for the perception of clearly separated events in time.

Duration and Subjective Present

Experiencing durations is a daily routine. Durations we experience can range from
a fraction of a second (the sound of a door bell), to several seconds (a car approaching
while we are crossing a street), or years (the time that elapsed since we left school). Not
a single timing process, however, can be assumed that is involved in the processing of
such a variety of time lengths. Fraisse (1984), for example, established the categorical
distinction between perception of duration and estimation of duration. He refers to James
( I 890), who introduced the notion of the psychological present (“specious present”) for
our feeling of “nowness” in subjective experience, which extends over a few seconds.
Durations of a few seconds are thereafter processed by means that are based on the actual
perception as a whole. Events with durations that exceed this limit of about 3 seconds
cannot be perceived as a subjective temporal gestalt; memory processes must be involved
that link moments in the past with the present. Thus, durations below 3 seconds are
 20 WITTMANN

perceived, whereas durations above 3 seconds are estimated; these notions refer to dis-
tinct processing modes. The width of the subjective present is controversial, with an
upper limit of 7 or 8 seconds postulated (Michon 1978), but an average of 2 to 3 seconds
seems a realistic value (Poppel 1978, 1997; Fraisse 1984; Block 1990). Experimental
evidence points to this time constant in information processing, which structures our
perception of objects and events. When a subject continuously gazes at the Necker cube,
the two perspectives of the ambiguous figure alternate every few seconds-with a mode
of about 3 seconds (G6mez et al. 1995; Steinbuchel 1998). When listening to a continu-
ous string of acoustic stimuli produced by a metronome, subjective structures of inte-
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grated beats are usually perceived by accentuating every xth stimulus (e.g., 1-2-3, 1-2-3,
. . .), with the integration units not exceeding 3 seconds (Szelag et al. 1996).

Cognitive Models of Duration Judgment

Four main methods in duration judgment tasks can be employed: verbal estimation
(or perception) of a period of time, reproduction of a stimulus duration, production of a
time span, and comparison between two stimulus durations (Allan 1979; Nichelli 1996;
Zakay and Block 1997). Relations between duration judgments and other factors are
generally examined. For example, subjects’ performances in the judgment of duration
are investigated by comparing empty (only the onset and offset is perceivable) and filled
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(a continuously presented tone is perceived) time periods. With this method, Rammsayer
and Lima (Rammsayer and Lima 1991), for example, disclosed different processing
modes for shorter (50-ms) and longer (1-second) durations.
The underlying models of duration judgment can be categorized into timing-with-
a-timer models and timing-without-a-timer models (for a discussion of this categonza-
tion, see Block 1990). In timing-with-a-timer models, a temporal pacemaker produces a
series of pulses with a stable frequency that are fed into a pulse counter. The number of
pulses that has been recorded for a physical time period is then compared with a memory
store in which correspondences of pulse counts with representations of time periods that
can be verbalized (as seconds or minutes) are stored. The whole system can be referred
to as the internal clock (Treisman et al. 1990; see Zakay and Block 1996 for a discussion
of similar timing-with-a-timer models). An attentional gate model complements these
cognitive models. It assumes that, in prospective time judgment, an attentional gate is
opened when an external signal starts a time period. When an external signal indicates
the end of the time period, the gate is closed, and the number of pulses is sent to a
reference memory (Zakay and Block 1997). When subjects follow a temporal task and,
at the same time, are confronted with a nontemporal distraction task, the attentional gate
is opened less wide as attention resources are diverted to the nontemporal task. As a
result, fewer pulses are recorded, creating the subjective impression that less time has
passed (Zakay 1993).
Timing-without-a-timermodels reconstruct time in reference to the amount of in-
formation processed and stored. The more information the subject has collected during
a time span, the longer the durations are experienced in retrospect (Ornstein 1969; Block
1990). The two models, however, are not exclusive. Regarding the distinction between
prospective and retrospective time estimation, they can be understood as representing
different cognitive processes that are active under certain conditions. In prospective tem-
poral judgment-when the subject knows he or she has to estimate a duration from that
point in time-the opening of the attentional gate is responsible for the experience of
 TIME PERCEPTION AND TEMPORAL PROCESSING 21

time. When subjects are instructed to estimate a time period in retrospect, after the rele-
vant duration has elapsed, the cognitive load (the amount of information stored in mem-
ory) leads to the impression of the time span.
All these aforementioned components involved in time estimation can account for
the time paradox everybody has experienced-waiting half an hour for a bus can be
unbearably long, but when we talk to a person to whom we are attracted, the same time
span is experienced as passing very quickly. Both cases are examples of a prospective
time judgment. In the former case, our attention is strongly attached to the (slow) passing
of time. In the latter case, our attention is not directed to time as it is solely directed to
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the conversation with the person (and, thus, time flies). However, when looking back at
the half hour that has passed, not much has happened during the time spent waiting for
the bus, and, owing to the lack of information stored in our memory, the experienced
time span shrinks to a negligibly small temporal portion of our life. In conlrast, the
intensive half hour spent with our friend may have been filled with so many thoughts
and feelings that were exchanged that, in retrospect, this time span extends to a consider-
ably longer time unit.

Influence of the Circadian Clock on Time Estimation

In a comprehensive review of mechanisms that are involved in temporal experi-
ence, circadian rhythms must also be considered. Diurnal variations on the basis of en-
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dogenous oscillations can be detected in numerous physiological and psychological func-

tions (Aschoff and Wever 1981) and also exist in temporal mechanisms of perception
and action (e.g., in the production of time intervals, when subjects are instructed to press
a button for the assumed duration of 10 seconds) (Poppel and Giedke 1970). There is also
evidence that circadian rhythms as an internal clock contribute to our time experience. In
isolation laboratories, subjects can be deprived of all external temporal cues (such as
external light, sound, temperature, etc.) for a period of weeks. Under these conditions,
subjects are able to make long-term time estimations (from l h to 24h), but estimates are
characterized as being rather sloppy (Campbell 1990). A relatively inexact time estima-
tion (in the range of hours) has been attributed to the sleep-wake cycle in which, in a
time-free environment (as in the isolation laboratory), sleep is not restricted to a stable
cycle, but can occur at any time of day. The association between the duration estimation
of hours and the sleep-wake cycle was assessed by Aschoff (1985), who found lhat time
estimation of l h intervals correlated with the duration of wakefulness and the length of
the circadian cycle. A correlation did not exist, however, for time estimation in the order
of minutes. The difference between long-term (in the range of hours) and short-term
(seconds to minutes) time estimation is further emphasized by the fact that light intensity,
which influences the subjective time experience, only affects the production of second
intervals and not the production of l h intervals (Aschoff and Daan 1997).

TIMING IN PERCEPTION AND ACTION

In the taxonomy of temporal processing levels, evidence for mechanisms in percep-

tion have been disclosed. However, as actions evolve over time, motor control depends
on processes that determine when exactly to move. Consider a simple finger-tapping task
for which taps with constant time intervals are initiated or consider the complex perfor-
 22 WITTMANN

mance of a drummer, who moves his arms and feet in a fixed time scheme to produce a
rhythm. One could ask whether the same timing mechanisms identified in perception
also underlie motor behavior. This question is integral to the debate whether perception
and movement share common processes (common coding) or are organized indepen-
dently (separate coding) (see Prinz 1992 for a discussion that concludes in favor of the
common-coding view). Certain phenomena indicate a common timing mechanism: we
often automatically move to a rhythm we hear, with the temporal structure of the beat
effortlessly being copied by foot tapping or movement of the whole body. A common
timing system would be an efficient way of controlling sensory-dependent behavior and
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anticipating changes in the environment. With this assumption in mind, it is tempting to

look for indicators of temporal processes of the described taxonomy.
Frequency distributions in motor tasks show several modes with intermodal dis-
tances of 30 ms in reaction-time tasks (Harter and White 1968) and in oculomotor move-
ments (Poppel and Logothetis 1986; Ruhnau and Haase 1993). These intermodal dis-
tances have been interpreted as indications of a discrete technique of motor processing
in steps of 30 ms and are thought to complement the findings of temporal-order threshold
that, taken together, could point to a central timing mechanism (Poppel 1994, 1997).
Results of motor performance tests also suggest a temporal segmentation of 2 to 3 sec-
onds to be a central feature of neural integration controlling behavior in humans and
other mammals (for a detailed overview, see Schleidt and Kien 1997). In humans, every-
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day movements (such as chopping, wiping, sewing, scratching) were analyzed in five
cultures and found to be segmented into action units of 2 to 3 seconds (Schleidt et al.
1987). An impressive effect is achieved in sensorimotor synchronization when tones with
a fixed interstimulus interval have to be reproduced by tapping a key. The task can be
carried out accurately, although the registered tap usually precedes the tone by some tens
of milliseconds (Aschersleben and Prinz 1995). Subjects are not aware of this slight
asynchrony. If, however, the interstimulus interval exceeds a certain length, the ability
to synchronize the presented beats breaks down. The critical interval varies interindividu-
ally and is about 2 seconds (Mates et al. 1994). It seems as if the motor system is only
capable of properly synchronizing to the consecutively presented sensory events in time
units that correspond to the subjective present.
Further insight into common timing derives from comparisons between perfor-
mance of timing in perception and in motor skills. Based on animal research that suggests
the pacemaker of an internal clock to be dependent on the dopamine level (Church 1984),
Rammsayer (1989) showed that the oral intake of a dopamine antagonist by healthy
volunteers impaired the ability of duration discrimination and increased choice reaction
time. Treisman and colleagues (1990) estimated the frequency of a temporal oscillator
involved in perception by observing disturbances of time estimation when clicks were
presented at certain rates; the frequency was estimated at 49.5 Hz. The same frequency
of just below 50 Hz was also calculated for a hypothesized oscillator underlying choice
reaction hand movements. These complementing results were then discussed as evidence
of a common internal clock for both perception and action (Treisman et al. 1992). In a
different study, subjects’ perceptual acuity in a duration judgment task (comparing two
acoustic intervals, one of which had a duration of 400 ms, and the other was longer or
shorter) was associated with the temporal variability in a tapping task for which intertap
intervals of 400 ms had to be produced (Keele et al. 1985). In a similar approach, further
evidence for a common timing mechanism was collected by showing that performance
 TIME PERCEPTION AND TEMPORAL PROCESSING 23

variability increased comparably as a function of duration in both a perceptual judgment

and a tapping task (Ivry and Hazeltine 1995).

NEURAL BASIS OF TEMPORAL PROCESSING

Temporal Binding and Temporal Separation
Fusion thresholds differ substantially among sensory modalities, with values lowest
in audition and highest in vision. Temporal-order thresholds, in contrast, do not vary
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essentially between assessed modalities. This difference between the two basic thresholds
regarding constancy over sense modalities has led to the hypothesis that the detection of
simultaneity and nonsimultaneity is based on peripheral mechanisms such as transduction
times in the receptor organs, whereas the detection of temporal order is based on more
central mechanisms (Hirsh and Sherrick 1961). Transduction times vary over sensory
modality: the relevant membrane time constants in the receptor organs were measured at
about 0.1 ms in auditory hair cells and 10 ms to 50 ms in the photoreceptors of the retina
(Torre et al. 1995). Clinical findings, however, suggest a more systemic view of the
underlying neural structures. Auditory fusion thresholds are only slightly elevated after
cortical injury (in a study by Lackner and Teuber [1973], from ca. 1.5 ms in control
subjects to ca. 2 ms in patients with left-hemispheric lesions). A more dramatic increase
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in auditory fusion threshold (by the factor of 10) is seen in patients with lesions to the
thalamus or the basal ganglia (Hosokawa et al. 1981). Patients with Parkinson’s disease
were affected in the recognition of separation of two stimuli in all three (auditory, tactile,
visual) tested sensory modalities (Artieda et al. 1992). Despite the plausible hypothesis
of a dependency of fusion thresholds on peripheral mechanisms, subcortical (especially
basal ganglia) structures also seem to be involved in temporal processing at this level.
The identification of temporal order indeed seems to be organized mainly by corti-
cal processes, more specifically, by processes in the left hemisphere that are associated
with language processing as patients with brain injuries in these regions and aphasia
demonstrate much higher order thresholds (Efron 1963; Swisher and Hirsh 1972; Stein-
buchel et al. 1996). A fine auditory temporal resolution for the order of spectral compo-
nents in the speech signal is necessary for identification (Pastore and Farrington 1996;
Bishop 1997). As both a phonological decoding deficit and higher temporal-order thresh-
olds are measured in patients with left hemispheric injuries and aphasia, one factor of a
disturbance in the language domain may be explained by an underlying temporal-process-
ing deficit (Tallal and Newcombe 1978; Steinbuchel and Poppel 1993).
Poppel (1994, 1997) proposed a model of a central oscillatory processing system
that explains the experience of two temporally ordered events over time. In this model,
a first stimulus can initiate a neuronal oscillation with periods of 30-ms duration. If the
second stimulus is processed within the same period of such an oscillation (e.g., the
interstimulus interval is below 30 ms), the two events are treated as cotemporal, ,which
means that temporal order is not evaluated. These system states of 30-ms duration could
be responsible for the binding of intra- and intermodal information (Poppel et al. 1990).
Due to this temporal-binding property, two spatial events have to be represented with an
interval longer than about 30 ms to be perceived in their temporal order.
Interestingly, theoretical calculations of internal pacemakers involved in time per-
ception and recordings of rhythmic neural activity exist that converge at about the same
 24 WITTMANN

numerical values of oscillatory periods with 20-ms to 40-ms lengths. The frequency of
an assumed temporal oscillator in the internal clock that underlies time judgments (see
Cognitive Models of Duration Judgment section) was estimated at 49.5 Hz (Treisman et
al. 1990). On this time scale, brain measures on the cellular level and with electro- and
magnetophysiological recordings have revealed rhythmic brain activity of around 40 Hz
(the gamma band) that are considered to be fundamental building blocks in sensory and
cognitive processing (for a comprehensive overview, see BaHar-Eroglu et al. 1996). For
example, in the visual cortex of the cat, these oscillations are recorded and interpreted
as a binding mechanism for feature-selective cells that are spatially distributed (Singer
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1993). Magnetoencephalograph(MEG) recordings in humans suggest that 40-Hz activity

is involved in the perceptual separation of acoustic events over time. The perception of
two temporally separated clicks corresponded to the appearance of two 40-Hz waves.
Only one 40-Hz wave was recorded when, below a certain threshold for the interstimulus
intervals, subjects could not separate the two clicks over time (Joliot et al. 1994). These
specific MEG recordings may result from a coherent 40-Hz resonance between thalamo-
cortical loops responsible for temporally binding neural processes involved in cognition
(Llinis and Ribary 1993).
Independent of these detected oscillatory binding operations, neurophysiological
recordings in the auditory cortex of the cat revealed cortical fields that are temporally
selective for certain monaurally presented stimulus frequencies (Schreiner and Urbas
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1988). The selectivity of a field for temporal variations of a given stimulus amplitude
can be described as its temporal resolution. The highest temporal resolution of a field (in
the anterior auditory field) was recorded for stimulus frequencies of about 30 Hz, with
this value corresponding to the temporal-order threshold of 20 ms to 40 ms usually
measured in humans. This finding, of course, cannot account for temporal-order detection
in a binaural task with the presentation of one click to each ear. In this case, one has to
assume a central timing mechanism that evaluates the time gap between the arriving
events. The notion of a central evaluator becomes even more plausible in intermodal
stimulation, for which the temporal order of a light flash and a click has to be judged
(Jaskowski et al. 1990). To conclude, these findings from measurements on the cellular
and macrocellular level all point to mechanisms that could be interpreted as a principle
of temporal processing of the brain.

Processing of Time Intervals

Involvement of the Cerebellum and the Basal Ganglia
In addition to an association of temporal-order judgment with cortical processes,
two main neural systems have been identified as being involved in temporal aspects of
behavior-the basal ganglia and the cerebellum (Ivry 1996; Gibbon et al. 1997). Korn-
huber (1971) once postulated that two separate neural clocks might be implemented by
these two neural structures. A clock integrated in a cerebellum-cortex loop would be
involved in ballistic and maximum speed movements, and a clock integrated in a basal
ganglia-cortex loop would be activated in smooth movements under voluntary control.
Empirical findings support this hypothesis: lesions to the cerebellum can lead to dysdia-
dochokinesia, the inability to alternate agonist and antagonist muscles in maximum speed
(Dichgans 1984). With various effectors, such as the finger, wrist, forearm, or foot,
healthy subjects perform repetitive movements in maximum speed with a frequency of
 TIME PERCEPTION AND TEMPORAL PROCESSING 25

around 5 Hz to 6 Hz (Keele and Hawkins 1982; McManus et al. 1986). In contrast, some
patients with Parkinson’s disease cannot perform with a pace that corresponds to self-
paced speed with a tapping frequency of around 2 Hz (two finger taps per second). When
required to perform in this tempo, some patients speed up, showing a maximum fre-
quency above 5 Hz. This hastening phenomenon in patients with Parkinson’s disease has
been reported for finger tapping (Nakamura et al. 1978) and for repetitive articulatory
movements (Ackermann et al. 1997). This phenomenon could not be replicated in every
study, but reduced accuracy in timing is still detectable (Freeman et al. 1993). A further
timing dependency was registered in a study on cerebral blood flow changes during
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finger movements in which activity of the cerebellum was predominantly seen when
movements were performed quickly; during slower movements, activity in basal ganglia
was mainly increased (Sabatini et al. 1993).
To separate the functions of the two neural systems, one could hypothesize differ-
ent time ranges. The cerebellum may be involved in the processing of relatively brief
durations, presumably being bound to the 3-second integration level (Clarke et al. 1996).
Patients with lesions of the cerebellum exhibited deficits in duration discrimination, with
standard intervals defined by two tones with 400 ms onset asynchrony between the pair
(Ivry and Keele 1989). In patients with cerebellar degeneration, deficits in perceptual
timing in that time range have been reported by Nichelli, Alway, and Grafman (1996). In
contrast, an internal clock linked to dopamine function in the basal ganglia is considered
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responsible for time durations from seconds to minutes (Meck 1996). In search of the
neural substrate of cognitive timer models (see Cognitive Models of Duration Judgment
section), neuropharmacological experiments on lesioned rats identified the substantia ni-
gra as the pulse generator and the dorsal striatum as the accumulator of those pulses
(Meck 1996). Patients with Parkinson’s disease underestimated the duration of time inter-
vals in seconds when counting and, by pressing a key, reproduced time intervals longer.
These results were interpreted as indications of a slowed-down internal clock (Pastor et
al. 1992). Differential results were reported in patients with lesions to the basal ganglia,
who performed with either acceleration or deceleration when estimating time intervals
up to 60 seconds. Here, lesions to the posterior part of the supralenticular white matter
were identified as responsible for the alterations (Rubia et al. 1997).
Despite these unequivocal results, other investigations indicate that the involvement
of the cerebellum and the basal ganglia cannot be differentiated by the time range (Gib-
bon et al. 1997). Attempts have been made to disclose the variance of a central timer
and a motor implementation system in finger tapping, referring to the model by Wing
and Kristofferson (1973). Patients with cerebellar lesions are differentially impaired in
one of the two variability components, depending on the lesion site, when synchronizing
their tapping performance to consecutively presented tones with interstimulus intervals
of 550 ms (Ivry et al. 1988). Patients with Parkinsons’s disease in an “off-medication’’
condition were investigated with a similar paradigm and showed greater variances in
both components (O’Boyle et al. 1996). Timing stability of the central timer in this motor
behavior apparently relies on both neural structures.
A further question is under debate concerning the characterization of the way time
is represented in the brain. Two possible mechanisms are discussed-oscillatory models
and interval-based models (Ivry 1996). In oscillatory models such as the clock-counter
models described above, an oscillatory mechanism produces output with a constant fre-
quency, with the number of accumulated pulses in a counter representing the time
elapsed. In these models, one mechanism is responsible for different durations, for in-
 26 WITTMANN

stance, durations of 500 ms and lo00 ms are represented by different numbers of pulses
in a counter (Treisman et al. 1990; Meck 1996). In interval-based models, however, time
is represented by nonoscillatory mechanisms, for which intervals of different durations
are processed by distinct sets of interval timers that are specific for certain durations
(Ivry 1996). One interval timer, for example, would be activated for durations of 500
ms, another for durations of loo0 ms. The representation of short time durations is
distributed over distinct neural mechanisms that are thought to be localized in the cerebel-
lum (Ivry 1996).
To sum, two distinct ways of processing could categorize timing in the cerebellum
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and the basal ganglia: the time level and the representational mode. First, the time level
would associate temporal processing of fractions of a second up to a few seconds to the
cerebellum and temporal processing from the seconds to minutes range to the basal
ganglia (Hazeltine et al. 1997). Second, an interval-based timing mechanism would apply
to the cerebellum and a clock-counter system to the basal ganglia.
Involvement of the Cortex
As both aforementioned neural systems are integrated in the cortex, cortical injur-
ies can cause distortion in temporal control of behavior (Ivy and Keele 1989). Patients
with unilateral lesions of the premotor cortex and the supplementary motor area (SMA)
showed deficits when requested to reproduce rhythms (Halsband et al. 1993). Monkeys
with lesions in the supplementary motor cortex were impaired in tasks that required
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raising the arm in a self-paced tempo (Chen et al. 1995). A study assessing cortical
activation in healthy volunteers identified the circuit comprising the basal ganglia, thala-
mus, and SMA as involved in timekeeping in tapping (Rao et al. 1997). In cortical
control of timed movements-analogous to the detection of temporal order-a hemi-
spheric asymmetry for the organization of repetitive-movement patterns could be the
case. In two patients after infarction of the corpus callosum, slow tapping to consecu-
tively presented beats with the left hand that, due to the special lesion, was deafferenti-
ated from the left hemisphere, was shown to be irregular. Tapping with the right hand,
controlled by the left hemisphere, was normal. This lateralization in performance did not
show in a tapping task with maximum speed. The authors (Kashiwagi et al. 1989) con-
cluded, therefore, that the left hemisphere is involved in the voluntarily controlled timing
of repetitive patterns. Temporal processing asymmetries between the hemispheres are
discussed both for perception and motor behavior, with the higher temporal resolution of
the left hemisphere being assumed as prerequisite to the sequential organization of lan-
guage perception and production (Tzeng and Wang 1984; Nicholls 1996).
What makes the understanding of involved neural structures more complicated is
that, in addition to the cerebellum taking part in duration processing, cortical areas have
been disclosed that are sensitive to specific durations of acoustic stimuli (He et al. 1997).
Specific areas in the dorsal zones of the auditory cortex in the cat were selectively activated
for presented durations ranging from 30 ms to 500 ms, thus lying in a time range for
duration processing that was associated with the cerebellum (Ivry 1996). In a lesion study
in which only patients with right-hemispheric lesions (compared to patients with left-hemi-
spheric lesions and controls) showed deficits in time perception with standard intervals of
300 ms and 600 ms, a right-hemispheric prefrontal-inferior-parietalnetwork in timing was
postulated (Harrington et al. 1998). Future research will have to separate more clearly
neural components involved in timing at different levels of the taxonomy and account for
their fine interplay to comprise all the empirical evidence gained.
 TIME PERCEPTION AND TEMPORAL PROCESSING 27

CONCLUSION

On the basis of a taxonomy of time experiences--comprising simultaneity, nonsi-

multaneity, temporal order, and duration-several thresholds have been identified that
refer to distinct temporal processes in the brain. With this classification, explorations in
relevant topics, such as the common timing hypothesis for perception and motor behavior
or the neural basis for temporal processes, seem promising. Here, cortical areas, the basal
ganglia, and the cerebellum have been identified as predominantly involved. Models
still have to be developed that account for the functional interaction of the three neural
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components.
In recent years, research into the topics has not only augmented basic scientific
knowledge, but also has led to clinical application of the findings. Based on the diagnos-
tic findings that patients with aphasia (Efron 1963; Tallal and Newcombe 1978; Steinbu-
chel et al. 1996), children with general language-learning impairments (Tallal and Piercy
1973), and children and adults with dyslexia (Tallal 1980; Hari and Kiesila 1996; for an
overview, see Farmer and Klein 1995) show impairments in temporal processing of rap-
idly presented stimuli (e.g., have increased order thresholds), training methods in tempo-
ral processing have been developed that have improved not only the ability in temporal
discrimination, but also have significantly improved basic language competence (Stein-
buchel 1995; Merzenich et al. 1996; Steinbiichel et al. 1996). As complex cognitive
For personal use only.

functioning is dependent on underlying temporal constraints (Steinbuchel and Poppel

1993), more empirically detected associations between timing functions and language
competence will probably be detected. Patients with Broca’s aphasia, for example, can
be disturbed in the task of subjectively grouping metronome beats to units of 2 to 3
seconds (Szelag et al. 1997).
The rather neglected topic of temporal processing will probably become more
prominent in the future. A considerable body of knowledge has been accumulated.
On the one hand, this knowledge contributes to our understanding of the mechanisms in
the experience of subjective time; on the other hand, the knowledge will provide informa-
tion about temporal properties as underlying constraints in cognitive functioning, such
as in language processing or in basic motor control. Furthermore, diagnostic and thera-
peutic tools may be developed to treat disturbed temporal mechanisms in various patient
populations with injuries and malfunctions of the brain.

ACKNOWLEDGMENT

I want to thank Ernst Poppel, Nicole von Steinbiichel, Till Roenneberg, and Bar-
bara Herzberger from the Institute of Medical Psychology in Munich and Elzbieta Szelag
from the Nencki Institute of Experimental Biology in Warsaw for their critical and help-
ful remarks on earlier versions of the manuscript.

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