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A large number of competing models exist for how the brain creates a representation of time. However, several human and animal studies point to 'climbing neural activation' as a potential neural mechanism for the representation of duration. Neurophysiological recordings in animals have revealed how climbing neural activation that peaks at the end of a timed interval underlies the processing of duration, and, in humans, climbing neural activity in the insular cortex, which is associated with feeling states of the body and emotions, may be related to the cumulative representation of time.

The "dual klepsydra model" (DKM) of internal time representation successfully models duration reproduction data, but relations between the DKM-based parameter kappa ("loss rate") and procedural variables (presentation modality) or individual characteristics (cognitive indices, age, sex) remained as yet unexplored. For that purpose, were-analyzed data from an earlier time reproduction study (N = 100), using visually or acoustically presented intervals of 1-5 sec. duration. Typical values of parameter kappa were approximately 0.03-0.04 sec.(-1), corresponding to relaxation times of internal "lossy integrators" of approximately 30 sec. Significant effects of presentation modality (smaller kappa values for the visual reproduction task) and of age (greater kappa in acoustic reproduction with increasing age) were observed. Cognitive variables (working memory, general fluid reasoning, attention) and sex of participants were not associated with kappa. Cognitive functions seem to play only a minor, if any, role at the level of time representation addressed by the DKM.

Several studies provide empirical evidence for the association between impulsivity and time perception. However, little is known about the neural substrates underlying this function. This investigation examined the influence of impulsivity on neural activation patterns during the encoding and reproduction of intervals with durations of 3, 9 and 18s using event-related functional magnetic resonance imaging (fMRI). Twenty-seven subjects participated in this study, including 15 high impulsive subjects that were classified based on their self-rating. FMRI activation during the duration reproduction task was correlated with measures of two self-report questionnaires related to the concept of impulsivity (Barratt Impulsiveness Scale, BIS; Zimbardo Time Perspective Inventory, ZTPI). Behaviorally, those individuals who under-reproduced temporal intervals also showed lower scores on the ZTPI future perspective subscale and higher scores on the BIS. FMRI activation revealed an accumulating pattern of neural activity peaking at the end of the 9- and 18-s intervals within right posterior insula. Activations of brain regions during the reproduction phase of the timing task, such as those related to motor execution as well as to the 'core control network' - encompassing the inferior frontal and medial frontal cortices, the anterior insula as well as the inferior parietal cortex - were significantly correlated with reproduced duration, as well as with BIS and ZTPI subscales. In particular, the greater activation in these regions the shorter were the reproduced intervals, the more impulsive was an individual and the less pronounced the future perspective. Activation in the core control network, thus, may form a biological marker for cognitive time management and for impulsiveness.

Recent research suggests that our sense of time intervals in the range of seconds is directly related to activity in the insular cortex, which contains the primary sensory area for interoception. We therefore investigated whether performance in a duration reproduction task might correlate with individual interoceptive awareness and with measurable changes in autonomic activity during the task. Thirty-one healthy volunteers participated in an interoceptive (heartbeat) perception task and in repeated temporal reproduction trials using intervals of 8, 14, and 20s duration while skin conductance levels and cardiac and respiratory periods were recorded. We observed progressive increases in cardiac periods and decreases in skin conductance level during the encoding and (less reliably) the reproduction of these intervals. Notably, individuals' duration reproduction accuracy correlated positively both with the slope of cardiac slowing during the encoding intervals and with individual heartbeat perception scores. These results support the view that autonomic function and interoceptive awareness underpin our perception of time intervals in the range of seconds.

It has been suggested that perception and action can be understood as evolving in temporal epochs or sequential processing units. Successive events are fused into units forming a unitary experience or "psychological present." Studies have identified several temporal integration levels on different time scales which are fundamental for our understanding of behavior and subjective experience. In recent literature concerning the philosophy and neuroscience of consciousness these separate temporal processing levels are not always precisely distinguished. Therefore, empirical evidence from psychophysics and neuropsychology on these distinct temporal processing levels is presented and discussed within philosophical conceptualizations of time experience. On an elementary level, one can identify a functional moment, a basic temporal building block of perception in the range of milliseconds that defines simultaneity and succession. Below a certain threshold temporal order is not perceived, individual events are processed as co-temporal. On a second level, an experienced moment, which is based on temporal integration of up to a few seconds, has been reported in many qualitatively different experiments in perception and action. It has been suggested that this segmental processing mechanism creates temporal windows that provide a logistical basis for conscious representation and the experience of nowness. On a third level of integration, continuity of experience is enabled by working memory in the range of multiple seconds allowing the maintenance of cognitive operations and emotional feelings, leading to mental presence, a temporal window of an individual's experienced presence.

Analyses of neural mechanisms of duration processing are essential for the understanding of psychological phenomena which evolve in time. Different mechanisms are presumably responsible for the processing of shorter (below 500 ms) and longer (above 500 ms) events but have not yet been a subject of an investigation with functional magnetic resonance imaging (fMRI). In the present study, we show a greater involvement of several brain regions - including right-hemispheric midline structures and left-hemispheric lateral regions - in the processing of visual stimuli of shorter as compared to longer duration. We propose a greater involvement of lower-level cognitive mechanisms in the processing of shorter events as opposed to higher-level mechanisms of cognitive control involved in longer events.

Individuals are different 'chronotypes' with early 'larks' and late 'owls' forming the limits of a normal distribution in the population. We recently described that late chronotypes who suffer from a conflict between internal and external time ('social jetlag') suffer from more mental distress and are more likely to smoke than early chronotypes (Wittmann, Dinich, Merrow, and Roenneberg, 2006 . Social jetlag: mis-alignment of biological and social time. Chronobiology International, 23:497-509.). We performed a detailed analysis of the same database collected in 2002 comprising 134 daily smokers and 366 nonsmokers, scrutinizing the relationships between chronotype, smoking, and alcohol consumption as well as psychological well-being using a multiple mediation analysis. On average, smokers tend to be later chronotypes, report more sleep-associated psychosomatic symptoms, are more depressed, less balanced, and less vigilant. The mediation analysis suggests that only those late chronotypes who smoke and those who drink more suffer from increased psychological distress. We suggest that 'chronotype' is introduced as an additional factor in substance use, that is, when considering motives for smoking and drinking.

An object moving towards an observer is subjectively perceived as longer in duration than the same object that is static or moving away. This "time dilation effect" has been shown for a number of stimuli that differ from standard events along different feature dimensions (e.g. color, size, and dynamics). We performed an event-related functional magnetic resonance imaging (fMRI) study, while subjects viewed a stream of five visual events, all of which were static and of identical duration except the fourth one, which was a deviant target consisting of either a looming or a receding disc. The duration of the target was systematically varied and participants judged whether the target was shorter or longer than all other events. A time dilation effect was observed only for looming targets. Relative to the static standards, the looming as well as the receding targets induced increased activation of the anterior insula and anterior cingulate cortices (the "core control network"). The decisive contrast between looming and receding targets representing the time dilation effect showed strong asymmetric activation and, specifically, activation of cortical midline structures (the "default network"). These results provide the first evidence that the illusion of temporal dilation is due to activation of areas that are important for cognitive control and subjective awareness. The involvement of midline structures in the temporal dilation illusion is interpreted as evidence that time perception is related to self-referential processing.

In experimental studies using flight simulations subjects' duration estimates have shown to be an effective indicator of cognitive task demands. In this study we wanted to find out whether subjective time perception could serve as a measure of cognitive workload during simulated car driving. Participants drove on a round course of a driving simulator consisting of three different environments with different levels of task demands. Drivers were required to perform a time-production task while driving the vehicle. Electrodermal activity and subjective ratings of mental workload (SWAT) were recorded simultaneously. The length of produced intervals increased significantly in more complex driving situations, as did electrodermal activity and subjective ratings of mental workload. Thus, time production is a valid indicator of cognitive involvement in simulated driving and could become a valid method to measure the current mental workload of car drivers in various traffic situations.

The present paper investigates the effects of age, sex, and cognitive factors on temporal-order perception. Nine temporal-order tasks were employed using two and four stimuli presented in the auditory and visual modalities. Significantly increased temporal-order thresholds (TOT) in the elderly were found for almost all tasks, while sex differences were only observed for two tasks. Multiple regression analyses show that the performance on most temporal-order tasks can be predicted by cognitive factors, such as speed of fluid reasoning, short-term memory, and attention. However, age was a significant predictor of TOT in three tasks using visual stimuli. We conclude (1) that age-related differences can often be attributed to cognitive factors involved in temporal-order perception, and (2) that the concept of temporal-order perception is more complex than implied by the current models.

Action of a hallucinogenic substance, psilocybin, on internal time representation was investigated in two double-blind, placebo-controlled studies: Experiment 1 with 12 subjects and graded doses, and Experiment 2 with 9 subjects and a very low dose. The task consisted in repeated reproductions of time intervals in the range from 1.5 to 5s. The effects were assessed by parameter kappa of the 'dual klepsydra' model of internal time representation, fitted to individual response data and intra-individually normalized with respect to initial values. The estimates kappa were in the same order of magnitude as in earlier studies. In both experiments, kappa was significantly increased by psilocybin at 90 min from the drug intake, indicating a higher loss rate of the internal duration representation. These findings are tentatively linked to qualitative alterations of subjective time in altered states of consciousness.

Neuropsychological studies in brain-injured patients with aphasia and children with specific language-learning deficits have shown the dependence of language comprehension on auditory processing abilities, i.e. the detection of temporal order. An impairment of temporal-order perception can be simulated by time reversing segments of the speech signal. In our study, we investigated how different lengths of time-reversed segments in speech influenced comprehension in ten native German speakers and ten participants who had acquired German as a second language. Results show that native speakers were still able to understand the distorted speech at segment lengths of 50 ms, whereas non-native speakers only could identify sentences with reversed intervals of 32 ms duration. These differences in performance can be interpreted by different levels of semantic and lexical proficiency. Our method of temporally-distorted speech offers a new approach to assess language skills that indirectly taps into lexical and semantic competence of non-native speakers.

Some authors have suggested separate mechanisms for the processing of temporal intervals above versus below 2-3s. Given that the evidence is mixed, the present experiment was carried out as a critical test of the separate-mechanism hypothesis. Subjects reproduced five standard durations of 1-5s presented in the auditory and visual modalities. The Corsi-block test was used to assess effects of working-memory span on different interval lengths. Greater working-memory span was associated with longer reproductions of intervals of 3-5s. A factor analysis run on mean reproduced intervals revealed one modality-unspecific factor for durations of 1-2s and two modality-specific factors for longer intervals. These results are interpreted as further indications that two different processes underlie temporal reproductions of shorter and longer intervals.

Hallucinogenic psilocybin is known to alter the subjective experience of time. However, there is no study that systematically investigated objective measures of time perception under psilocybin. Therefore, we studied dose-dependent effects of the serotonin (5-HT)2A/1A receptor agonist psilocybin (4-phosphoryloxy-N, N-dimethyltryptamine) on temporal processing, employing tasks of temporal reproduction, sensorimotor synchronization and tapping tempo. To control for cognitive and subjective changes, we assessed spatial working memory and conscious experience. Twelve healthy human volunteers were tested under placebo, medium (115 microg/kg), and high (250 microg/kg) dose conditions, in a double-blind experimental design. Psilocybin was found to significantly impair subjects' ability to (1) reproduce interval durations longer than 2.5 sec, (2) to synchronize to inter-beat intervals longer than 2 sec and (3) caused subjects to be slower in their preferred tapping rate. These objective effects on timing performance were accompanied by working-memory deficits and subjective changes in conscious state, namely increased reports of 'depersonalization' and 'derealization' phenomena including disturbances in subjective 'time sense.' Our study is the first to systematically assess the impact of psilocybin on timing performance on standardized measures of temporal processing. Results indicate that the serotonin system is selectively involved in duration processing of intervals longer than 2 to 3 seconds and in the voluntary control of the speed of movement. We speculate that psilocybin's selective disruption of longer intervals is likely to be a product of interactions with cognitive dimensions of temporal processing -presumably via 5-HT2A receptor stimulation.

OBJECTIVE: The experience of time is strongly related to our momentary mood states. Patients with a life-threatening illness experience an extreme change in mood and suffer from psychological distress that can develop into clinically relevant psychiatric disorders, like anxiety and depression. The aim of this study was to investigate the associations among the subjective perception of time, psychological distress, and quality of life in patients with hematological malignancies. METHODS: Eighty-eight inpatients with hematological malignancies rated how fast time passes subjectively on a visual analog scale and prospectively estimated a time span of 13 min. The Hospital Anxiety and Depression Scale (HADS) self-report measures of health-related quality of life (FACT-G) and spiritual well-being (FACIT-Sp) were employed to assess psychological distress and quality of life. RESULTS: Those patients who reported a lower quality of life, less spiritual well-being, and more anxiety experienced a slower passage of subjective time and overestimated the 13-min time interval. SIGNIFICANCE OF RESULTS: Our interpretation of the results is that patients with a life-threatening illness who show symptoms of psychological distress draw attention away from meaningful thoughts and actions and, thus, experience time as passing more slowly. An altered sense of time can be a sign of mental suffering, which should be addressed within psycho-oncological interventions. As this is the first study to demonstrate this relation in cancer patients, further research is needed to investigate the experience of time and its relation to meaning as an issue in clinical diagnostics.

Standard diagnostic procedures for assessing temporal-processing abilities of adult patients with aphasia have so far not been developed. In our study, temporal-order measurements were conducted using two different experimental procedures to identify a suitable measure for clinical studies. Additionally, phoneme-discrimination abilities were tested on the word, as well as on the sentence level, as a relationship between temporal processing and phoneme-discrimination abilities is assumed. Patients with aphasia displayed significantly higher temporal-order thresholds than control subjects. The detection of an association between temporal processing and speech processing, however, depended on the stimuli and the phoneme-discrimination tasks used. Our results also suggest top-down feedback on phonemic processing.

To determine the relative safety of onboard display positions while driving, participants performed a lane-keeping task in a driving simulator. Concurrently, they reacted to a light by pushing the brake pedal. A secondary task was projected onto a display at one of the seven different locations in the cockpit. Behavioral data, eye movements, and subjective rating scales showed that the manipulation of display information during driving disturbed drivers' performance exponentially as a function of distance between the line of sight to the outside primary task and the onboard display position. Vertical eccentricity had a greater detrimental effect than horizontal distance. Under a certain condition with a high secondary task load, reaction time of pushing the brake to the outside stimulus nearly doubled with a diagonal eccentricity of 35 degrees as compared to lower eccentricities. Subjective workload measures complement the behavioral data of clear detrimental effects with eccentricities of at least 35 degrees .

Humans show large differences in the preferred timing of their sleep and activity. This so-called "chronotype" is largely regulated by the circadian clock. Both genetic variations in clock genes and environmental influences contribute to the distribution of chronotypes in a given population, ranging from extreme early types to extreme late types with the majority falling between these extremes. Social (e.g., school and work) schedules interfere considerably with individual sleep preferences in the majority of the population. Late chronotypes show the largest differences in sleep timing between work and free days leading to a considerable sleep debt on work days, for which they compensate on free days. The discrepancy between work and free days, between social and biological time, can be described as 'social jetlag.' Here, we explore how sleep quality and psychological wellbeing are associated with individual chronotype and/or social jetlag. A total of 501 volunteers filled out the Munich ChronoType Questionnaire (MCTQ) as well as additional questionnaires on: (i) sleep quality (SF-A), (ii) current psychological wellbeing (Basler Befindlichkeitsbogen), (iii) retrospective psychological wellbeing over the past week (POMS), and (iv) consumption of stimulants (e.g., caffeine, nicotine, and alcohol). Associations of chronotype, wellbeing, and stimulant consumption are strongest in teenagers and young adults up to age 25 yrs. The most striking correlation exists between chronotype and smoking, which is significantly higher in late chronotypes of all ages (except for those in retirement). We show these correlations are most probably a consequence of social jetlag, i.e., the discrepancies between social and biological timing rather than a simple association to different chronotypes. Our results strongly suggest that work (and school) schedules should be adapted to chronotype whenever possible.

OBJECTIVES: The study aimed to clarify whether cognitive and psychomotor performance, which are important for occupational and traffic safety, are impaired by working permanent night shifts (NSs) compared with early-late two shifts (TSs) and whether age and chronobiological type influences the relationship between shift and performance. METHODS: The study included 44 male automobile workers, 20 working TSs and 24 working NSs. Chronobiological type was determined by questionnaire (D-MEQ). Each subject was tested at the beginning and end of the shift for alertness [by a visual analogue scale (VAS)]; feeling of well-being (Basler); concentration and accuracy (d2); reaction speed, orientation and reaction to stress (Vienna System). RESULTS: TS workers were more frequently morning types whereas the NS workers were more frequently evening types. In the performance tests, the TS and NS workers did not differ at shift start or shift end. Over the course of the shift, concentration and accuracy improved in both groups, as did reaction to stress. Chronobiological type alone or in combination with shift type had no effect on performance. CONCLUSIONS: The results of this study indicate that-if chosen voluntarily-working NSs has no immediate negative effects on cognitive and psychomotor performance when compared with working TSs. There was no indication of an increased risk of accidents after working NSs. The unequal distribution of the circadian types in the shift groups may indicate selection.

PURPOSE: The relationship between auditory temporal-order perception and phoneme discrimination has been discussed for several years, based on findings, showing that patients with cerebral damage in the left hemisphere and aphasia, as well as children with specific language impairments, show deficits in temporal-processing and phoneme discrimination. Over the last years several temporal-order measurement procedures and training batteries have been developed. However, there exists no standard diagnostic tool for adults that could be applied to patients with aphasia. Therefore, our study aimed at identifying a feasible, reliable and efficient measurement procedure to test for auditory-temporal processing in healthy young and elderly adults, which in a further step can be applied to patients with aphasia. METHODS: The tasks varied according to adaptive procedures (staircase vs. maximum-likelihood), stimuli (tones vs. clicks) and stimulation modes (binaural- vs. alternating monaural) respectively. A phoneme-discrimination task was also employed to assess the relationship between temporal and language processing. RESULTS: The results show that auditory temporal-order thresholds are stimulus dependent, age related, and influenced by gender. Furthermore, the cited relationship between temporal-order threshold and phoneme discrimination can only be confirmed for measurements with pairs of tones. CONCLUSION: Our results indicate, that different norms have to be established for different gender and age groups. Furthermore, temporal-order measurements with tones seem to be more suitable for clinical intervention studies than measurements with clicks, as they show higher re-test reliabilities, and only for measurements with tones an association with phoneme-discrimination abilities was found.