Supra-second Intervals Research Articles

Cerebellar Roles in Self-Timing for Sub- and Supra-Second Intervals.

Previous studies suggest that the cerebellum and basal ganglia are involved in sub-second and supra-second timing, respectively. To test this hypothesis at the cellular level, we examined the activity of single neurons in the cerebellar dentate nucleus in monkeys performing the oculomotor version of the self-timing task. Animals were trained to report the passage of time of 400, 600, 1200, or 2400 ms following a visual cue by making self-initiated memory-guided saccades. We found a sizeable preparatory neuronal activity before self-timed saccades across delay intervals, while the time course of activity correlated with the trial-by-trial variation of saccade latency in different ways depending on the length of the delay intervals. For the shorter delay intervals, the ramping up of neuronal firing rate started just after the visual cue and the rate of rise of neuronal activity correlated with saccade timing. In contrast, for the longest delay (2400 ms), the preparatory activity started late during the delay period, and its onset time correlated with self-timed saccade latency. Because electrical microstimulation applied to the recording sites during saccade preparation advanced self-timed but not reactive saccades, regardless of their directions, the signals in the cerebellum may have a causal role in self-timing. We suggest that the cerebellum may regulate timing in both sub-second and supra-second ranges, although its relative contribution might be greater for sub-second than for supra-second time intervals.SIGNIFICANCE STATEMENT How we decide the timing of self-initiated movement is a fundamental question. According to the prevailing hypothesis, the cerebellum plays a role in monitoring sub-second timing, whereas the basal ganglia are important for supra-second timing. To verify this, we explored neuronal signals in the monkey cerebellum while animals reported the passage of time in the range 400-2400 ms by making eye movements. Contrary to our expectations, we found that neurons in the cerebellar dentate nucleus exhibited a similar preparatory activity for both sub-second and supra-second intervals, and that electrical simulation advanced self-timed saccades in both conditions. We suggest that the cerebellum plays a causal role in the fine adjustment of self-timing in a larger time range than previously thought.

Timescale- and Sensory Modality-Dependency of the Central Tendency of Time Perception.

When individuals are asked to reproduce intervals of stimuli that are intermixedly presented at various times, longer intervals are often underestimated and shorter intervals overestimated. This phenomenon may be attributed to the central tendency of time perception, and suggests that our brain optimally encodes a stimulus interval based on current stimulus input and prior knowledge of the distribution of stimulus intervals. Two distinct systems are thought to be recruited in the perception of sub- and supra-second intervals. Sub-second timing is subject to local sensory processing, whereas supra-second timing depends on more centralized mechanisms. To clarify the factors that influence time perception, the present study investigated how both sensory modality and timescale affect the central tendency. In Experiment 1, participants were asked to reproduce sub- or supra-second intervals, defined by visual or auditory stimuli. In the sub-second range, the magnitude of the central tendency was significantly larger for visual intervals compared to auditory intervals, while visual and auditory intervals exhibited a correlated and comparable central tendency in the supra-second range. In Experiment 2, the ability to discriminate sub-second intervals in the reproduction task was controlled across modalities by using an interval discrimination task. Even when the ability to discriminate intervals was controlled, visual intervals exhibited a larger central tendency than auditory intervals in the sub-second range. In addition, the magnitude of the central tendency for visual and auditory sub-second intervals was significantly correlated. These results suggest that a common modality-independent mechanism is responsible for the supra-second central tendency, and that both the modality-dependent and modality-independent components of the timing system contribute to the central tendency in the sub-second range.

Processing of sub- and supra-second intervals in the primate brain results from the calibration of multiple neuronal oscillators via sensory, motor, and feedback processes

Processing of sub- and supra-second intervals in the primate brain results from the calibration of multiple neuronal oscillators via sensory, motor, and feedback processes

Working memory for time intervals in auditory rhythmic sequences.

The brain can hold information about multiple objects in working memory. It is not known, however, whether intervals of time can be stored in memory as distinct items. Here, we developed a novel paradigm to examine temporal memory where listeners were required to reproduce the duration of a single probed interval from a sequence of intervals. We demonstrate that memory performance significantly varies as a function of temporal structure (better memory in regular vs. irregular sequences), interval size (better memory for sub- vs. supra-second intervals), and memory load (poor memory for higher load). In contrast memory performance is invariant to attentional cueing. Our data represent the first systematic investigation of temporal memory in sequences that goes beyond previous work based on single intervals. The results support the emerging hypothesis that time intervals are allocated a working memory resource that varies with the amount of other temporal information in a sequence.

How many neural oscillators we need on sub- and supra-second intervals processing in the primate brain

Processing of time intervals in short tem-poral ranges, typically in sub- and supra-second intervals, is vital for the survivalof primates and humans. Research evi-dence has already shown that the process-ing of sub- and supra-second intervals issubserved by different neural mechanisms(Buhusi and Meck, 2005). In the domainof time perception, one outstanding sci-entific question remains to be solved iswhether the interval processing is imple-mented by a central clock (supra-sensory)or specific clock (neural state-specific)representation (Ivry and Schlerf, 2008).Gupta (2014) initiated theoretical explo-rations by introducing different types ofneuronal oscillators, with three key com-ponents:pacemakerneurons,tonicinputs,and synchronized excitation/inhibition ofinter-connected neurons (Gupta, 2014).Gupta further featured three characteris-ticfactorsinanintegrated model:(1).Theencoding of the time interval is reflectedby frequency modulated neuronal spikesor spikebursts affiliated with thetemporalpoints of a given interval. (2). Our brainembraces active calibration mechanismsamong multiple oscillators and modularconnections, which could to a large extentpitagainstthelossofintervaltimingfunc-tions of the brain. (3). The neuronal oscil-lators have solid foundations in neuralsubstrates that typically characterize sen-sorimotor timing, from the changes ofmembrane potential to the functioningof high-level brain structures (Teki et al.,2011; Santos et al., 2014).The concept of the building block for“neural oscillators” in Gupta (2014) inessence could be extended and interpretedin different contexts: Firstly, the “neu-ronaltemporalunits”in, Gupta (2014)in contrast to the traditional “pacemaker-accumulator” model (Treisman, 1963;Treisman et al., 1990, 1994; Wittmann,1999), is a machinery device (inherent inthebrain)thatprocessesthetimeinforma-tion. Secondly, the “oscillators” entail the“modular” nature of neuronal clocks, byemphasizing the correspondence betweenneural substrates in temporal bindingsbetween different sensory events. In thisway and from a broad perspective, thismodel dovetails the tenets of “intrin-sic model” (Ivry and Schlerf, 2008), butallows mechanistic flexibility for the inter-actions between separate clocks. Thirdly,the oscillators could demonstrate higher-level ensemble neuronal activities (butwere relatively less addressed in cur-rentmodel),includingoscillatorygamma-band responses (Herrmann et al., 2004;Tallon-Baudry and Bertrand, 1999).Movement is the expressed time (Tekiet al., 2011). The Theoretical modelfor Gupta (2014) stems from the com-mon setting of “movement control” butencourages more potential investigationstocome.Forexample,itcouldbeextendedand validated in other timing scenarios—go beyond the area of motor control—todemonstrate its robustness. Moreover, atypicalfunctionofhumantimingsystemisto predict the imminent events and hencemake efficient perceptual decision mak-ing. The current oscillation model leavesmuch room to be improved by revealinghowtheoscillatorscouldworkdirectlyinaframework of predictive coding. This pre-dictivecodingissomehowfreeoffeedbackprocesses as assumed in Gupta (2014).Furthermore, a high-level processing ofrhythm/periodicity-based upon the infor-mation of sub- or supra-second intervals,might mobilizepopulation/ensemble neu-ronal coding as well as impose more com-plexinter-connections/overlappings ofthesupposed neuronal temporal units (withmultiple neuronal feedback loops). WhatGupta (2014) proposed essentially indi-cated a potential panorama of neuraloscillators that could address the com-plexity of timing behavior. However,how many (levels of) oscillators areneeded to elucidate those timing behav-iors and how local lateral excitatory con-nectivity (Gavornik and Shouval, 2011),interacting with the global (stochastic)processingofbeattiming,warrantsfurtherexplorations.

Processing of sub- and supra-second intervals in the primate brain results from the calibration of neuronal oscillators via sensory, motor, and feedback processes.

The processing of time intervals in the sub- to supra-second range by the brain is critical for the interaction of primates with their surroundings in activities, such as foraging and hunting. For an accurate processing of time intervals by the brain, representation of physical time within neuronal circuits is necessary. I propose that time dimension of the physical surrounding is represented in the brain by different types of neuronal oscillators, generating spikes or spike bursts at regular intervals. The proposed oscillators include the pacemaker neurons, tonic inputs, and synchronized excitation and inhibition of inter-connected neurons. Oscillators, which are built inside various circuits of brain, help to form modular clocks, processing time intervals or other temporal characteristics specific to functions of a circuit. Relative or absolute duration is represented within neuronal oscillators by “neural temporal unit,” defined as the interval between regularly occurring spikes or spike bursts. Oscillator output is processed to produce changes in activities of neurons, named frequency modulator neuron, wired within a separate module, represented by the rate of change in frequency, and frequency of activities, proposed to encode time intervals. Inbuilt oscillators are calibrated by (a) feedback processes, (b) input of time intervals resulting from rhythmic external sensory stimulation, and (c) synchronous effects of feedback processes and evoked sensory activity. A single active clock is proposed per circuit, which is calibrated by one or more mechanisms. Multiple calibration mechanisms, inbuilt oscillators, and the presence of modular connections prevent a complete loss of interval timing functions of the brain.

Early Attentional Modulation of the Neural Network Evoked with the Auditory Paired-click Paradigm: An MEG Study

Although numerous research studies have explored the functional attributes of the human duration perception, the spatio- temporal information on cortical networks underlying this process is still an open question. Moreover, the issue of possible differences in the nature of timing mechanisms responsible for perception of sub- and supra-second intervals requires the implementation of the functional brain imaging techniques with both high spatial and temporal resolution. Attention is very frequently used as a modulating factor for the perceived duration of a sensory stimulus. The non-attended stimuli appear to last shorter than the attended ones, increasing the perceived duration of a concurrent stimulus (Gorea, 2011). This study challenges the accepted model of early sensory responses, activated during the first 100ms after stimulus presentation, as preattentive, automatic processes which modulate the neural sensitivity to incoming stimuli (gating phenomena). We utilized Elekta Neuromag 306-channel whole-head system for magnetoencephalography (MEG) measurements and multi-dipole Calibrated Start Spatio Temporal (CSST) localization technique (Ranken et al., 2002) to investigate whether voluntary attention directed at the second tone of a pair in the standard paired-click paradigm could affect cortical networks underlying the gating out phenomenon. MEG recordings were obtained in a magnetically shielded room at the Biomagnetic Center in Jena, Germany. Two consecutive (ISI=500ms) identical short tonnes (S1 and S2; duration=20ms; f=1200Hz; ITI=8±1s) were used to evoke standard gating cortical responses in 19 healthy participants (21-38 years). In the second condition, the task was to direct attention toward the second tone and to respond to a rarely presented non-identical second tone of the pair (R=1300Hz, p(S1R)=0.3). MEG recordings showed transient early and middle latency responses with peak amplitudes over the temporo-parietal sensors followed by a peak of a sustained activity. Less prominent transient activity was recorded over the frontal sensors, accompanied with a more steady- state component from 100 until 400ms post-stimulus for both conditions. Preliminary results of the CSST spatio-temporal analyses revealed 4-6 brain regions activated during 20-500ms time interval including bilateral superior temporal gyrus (STG), bilateral and medial prefrontal (PF), bilateral parietal (PA) regions, and central motor cortex area. A cortical source underlying the steady-state component was identified in the left prefrontal region for S1 tone, and in central posterior regions (PA) for the repeated tone S2. During the standard paradigm the M50 gating suppression was observed only for bilateral STG sources while bilateral PF and PA sources did not show any reduction in the response during the first 100ms post-stimulus. Targeting the attention towards the second tone enhanced the M50 amplitude of the bilateral STG responses in respect to the repeated stimulus S2 and caused a change in the activated network evident by the emergence of a new generator in the medial PF area (orbitofrontal) instead of dorsolateral PF generators evoked in non-attended condition. These results demonstrate that voluntary attention can exert a topological and functional modulatory influence on the neural network even during early stages of auditory processing.

The basal ganglia in perceptual timing: Timing performance in Multiple System Atrophy and Huntington's disease

The timing of perceptual events depends on an anatomically and functionally connected network comprising basal ganglia, cerebellum, pre-frontal cortex and supplementary motor area. Recent studies demonstrate the cerebellum to be involved in absolute, duration-based timing, but not in relative timing based on a regular beat. Conversely, functional involvement of the striatum is observed in relative timing, but its role in absolute timing is unclear.This work tests the specific role of the basal ganglia in the perceptual timing of auditory events. It aims to distinguish the hypothesised unified model of time perception (Teki, Grube, & Griffiths, 2012), in which the striatum is a mandatory component for all timing tasks, from a modular system in which they subserve relative timing, with absolute timing processed by the cerebellum.Test groups comprised individuals with Multiple System Atrophy, a disorder in which similar pathology can produce clinical deficits associated with dysfunction of the cerebellum (MSA-C, n=8) or striatum (MSA-P, n=10), and early symptomatic Huntington's disease (HD, n=14). Individuals with chronic autoimmune peripheral neuropathy (n=11) acted as controls.Six adaptive tasks were carried out to assess perceptual thresholds for absolute timing through duration discrimination for sub- and supra-second time intervals, and relative timing through the detection of beat-based regularity and irregularity, detection of a delay within an isochronous sequence, and the discrimination of sequences with metrical structure.All three patient groups exhibited impairments in performance in comparison with the control group for all tasks, and severity of impairment was significantly correlated with disease progression. No differences were demonstrated between MSA-C and MSA-P, and the most severe impairments were observed in those with HD.The data support an obligatory role for the basal ganglia in all tested timing tasks, both absolute and relative, as predicted by the unified model. The results are not compatible with models of a brain timing network based upon independent modules.

Motor Timing Deficits in Sequential Movements in Parkinson Disease Are Related to Action Planning: A Motor Imagery Study

Timing of sequential movements is altered in Parkinson disease (PD). Whether timing deficits in internally generated sequential movements in PD depends also on difficulties in motor planning, rather than merely on a defective ability to materially perform the planned movement is still undefined. To unveil this issue, we adopted a modified version of an established test for motor timing, i.e. the synchronization–continuation paradigm, by introducing a motor imagery task. Motor imagery is thought to involve mainly processes of movement preparation, with reduced involvement of end-stage movement execution-related processes. Fourteen patients with PD and twelve matched healthy volunteers were asked to tap in synchrony with a metronome cue (SYNC) and then, when the tone stopped, to keep tapping, trying to maintain the same rhythm (CONT-EXE) or to imagine tapping at the same rhythm, rather than actually performing it (CONT-MI). We tested both a sub-second and a supra-second inter-stimulus interval between the cues. Performance was recorded using a sensor-engineered glove and analyzed measuring the temporal error and the interval reproduction accuracy index. PD patients were less accurate than healthy subjects in the supra-second time reproduction task when performing both continuation tasks (CONT-MI and CONT-EXE), whereas no difference was detected in the synchronization task and on all tasks involving a sub-second interval. Our findings suggest that PD patients exhibit a selective deficit in motor timing for sequential movements that are separated by a supra-second interval and that this deficit may be explained by a defect of motor planning. Further, we propose that difficulties in motor planning are of a sufficient degree of severity in PD to affect also the motor performance in the supra-second time reproduction task.

Numerical quantity affects time estimation in the suprasecond range

Of stimuli differing in the magnitude of their numerical information, the one with the larger numerosity is perceived to last longer than that with the smaller numerosity. This numerosity–time interaction is proposed to be due to a shared neural representation for numerical magnitude and time intervals in the parietal cortex. Neuroimaging studies of temporal processing suggest that subsecond and suprasecond intervals could be mediated by distinct neural substrates. However, whether the numerosity–time interaction occurs independently of the time intervals used in the tasks remains unknown. Here we show that numerical information interacts with time estimation in the suprasecond range in females, but not in males. Our results suggest that the numerical magnitude and suprasecond intervals have shared representations in the human brain, but the associative strength between these dimensions might be different between males and females.

The influence of tempo upon the rhythmic motor control in macaque monkeys

We examined behavioral features of isochronous repetitive movements in two macaques. The monkeys were required to press a button repetitively in response to external cues. If the cue-intervals were constant (isochronous) and sub-second, the reaction time was shorter than in random-interval condition. In contrast, in the supra-second isochronous conditions, the reaction time was not different from random-interval condition. The results suggest that the monkeys can acquire isochronous rhythms if the intervals are sub-second, probably depending on the automatic timing system. However, the conscious timing system for supra-second intervals is not well developed in monkeys, unlike humans.

Perception of Short Time Scale Intervals in a Hypnotic Virtuoso1

Previous studies showed that hypnotized individuals underestimate temporal intervals in the range of several seconds to tens of minutes. However, no previous work has investigated whether duration perception is equally disorderly when shorter time intervals are probed. In this study, duration perception of a hypnotic virtuoso was tested using repeated standard temporal generalization and duration estimation tasks. When compared to the baseline state, hypnosis affected perception of intervals spread around 600 ms in the temporal generalization task but did not alter perception of slightly longer intervals spread around 1000 ms. Furthermore, generalization of temporal intervals was more orderly under hypnosis than in the baseline state. In contrast, the hypnotic virtuoso showed a typical time underestimation effect when perception of longer supra-second intervals was tested in the duration estimation task, replicating results of the previous hypnosis studies.

Double Dissociation of Dopamine Genes and Timing in Humans

A number of lines of evidence implicate dopamine in timing [Rammsayer, T. H. Neuropharmacological approaches to human timing. In S. Grondin (Ed.), Psychology of time (pp. 295-320). Bingley, UK: Emerald, 2008; Meck, W. H. Neuropharmacology of timing and time perception. Brain Research, Cognitive Brain Research, 3, 227-242, 1996]. Two human genetic polymorphisms are known to modulate dopaminergic activity. DRD2/ANKK1-Taq1a is a D(2) receptor polymorphism associated with decreased D(2) density in the striatum [Jönsson, E. G., Nothen, M. M., Grunhage, F., Farde, L., Nakashima, Y., Propping, P., et al. Polymorphisms in the dopamine D(2) receptor gene and their relationships to striatal dopamine receptor density of healthy volunteers. Molecular Psychiatry, 4, 290-296, 1999]; COMT Val158Met is a functional polymorphism associated with increased activity of the COMT enzyme such that catabolism of synaptic dopamine is greater in pFC [Meyer-Lindenberg, A., Kohn, P. D., Kolachana, B., Kippenhan, S., McInerney-Leo, A., Nussbaum, R., et al. Midbrain dopamine and prefrontal function in humans: Interaction and modulation by COMT genotype. Nature Neuroscience, 8, 594-596, 2005]. To investigate the role of dopamine in timing, we genotyped 65 individuals for DRD2/ANKK1-Taq1a, COMT Val158Met, and a third polymorphism, BDNF Val66Met, a functional polymorphism affecting the expression of brain-derived neurotrophic factor [Egan, M. F., Kojima, M., Callicott, J. H., Goldberg, T. E., Kolachana, B. S., Bertolino, A., et al. The BDNF val66met polymorphism affects activity-dependent secretion of BDNF and human memory and hippocampal function. Cell, 112, 257-269, 2003]. Subjects were tested on a temporal discrimination task with sub- and supra-second intervals (500- and 2000-msec standards) as well as a spontaneous motor tempo task. We found a double dissociation for temporal discrimination: the DRD2/ANKK1-Taq1a polymorphism (A1+ allele) was associated with significantly greater variability for the 500-msec duration only, whereas the COMT Val158Met polymorphism (Val/Val homozygotes) was associated with significantly greater variability for the 2000-msec duration only. No differences were detected for the BDNF Vall66Met variant. Additionally, the DRD2/ANKK1-Taq1a polymorphism was associated with a significantly slower preferred motor tempo. These data provide a potential biological basis for the distinctions between sub- and supra-second timing and suggest that BG are integral for the former whereas pFC is implicated in the latter.

Oxycodone lengthens reproductions of suprasecond time intervals in human research volunteers

Oxycodone, a popularly used opioid for treating pain, is widely abused. Other drugs of abuse have been shown to affect time perception, which, in turn, may affect sensitivity to future consequences. This may contribute to continued use. This study evaluated the effect of oxycodone on time perception in normal healthy volunteers. For this within-subject, double-blind design study, participants performed a temporal reproduction task before and after receiving placebo or oxycodone (15 mg, orally) over six outpatient sessions. Participants were first trained with feedback to reproduce three standard intervals (1.1, 2.2, and 3.3 s) in separate blocks by matching response latency from a start signal to the duration of that block's standard interval. During testing, participants were instructed to reproduce the three intervals from memory without feedback before and after drug administration. Oxycodone significantly lengthened time estimations for the two longer intervals relative to placebo. These results suggest that opioids alter temporal processing for intervals greater than 1 s, raising questions about the effect of these drugs on the valuation of future consequences.

Left hand dominance affects supra-second time processing.

Previous studies exploring specific brain functions of left- and right-handed subjects have shown variances in spatial and motor abilities that might be explained according to consistent structural and functional differences. Given the role of both spatial and motor information in the processing of temporal intervals, we designed a study aimed at investigating timing abilities in left-handed subjects. To this purpose both left- and right-handed subjects were asked to perform a time reproduction of sub-second vs. supra-second time intervals with their left and right hand. Our results show that during processing of the supra-second intervals left-handed participants sub-estimated the duration of the intervals, independently of the hand used to perform the task, while no differences were reported for the sub-second intervals. These results are discussed on the basis of recent findings on supra-second motor timing, as well as emerging evidence that suggests a linear representation of time with a left-to-right displacement.

Temporal Discrimination of Sub- and Suprasecond Time Intervals: A Voxel-Based Lesion Mapping Analysis

We used voxel-based lesion-symptom mapping (VLSM) to determine which brain areas are necessary for discriminating time intervals above and below 1 s. VLSM compares behavioral scores of patients that have damage to a given voxel to those that do not on a voxel-by-voxel basis to determine which voxels are critical for the given behavior. Forty-seven subjects with unilateral hemispheric lesions performed a temporal discrimination task in which a standard stimulus was compared on each trial to a test stimulus. In different blocks of trials, standard stimuli were either 600 or 2000 ms. Behavioral measures included the point of subjective equality, a measure of accuracy, and the coefficient of variation, a measure of variability. Lesions of the right middle and inferior frontal gyri were associated with decrements in performance on both durations. In addition, lesions of the left temporal lobe and right precentral gyrus were associated exclusively with impaired performance for subsecond stimuli. In line with results from other studies, these data suggest that different circuits are necessary for timing intervals in these ranges, and that right frontal areas are particularly important to timing.

Optimal Temporal Risk Assessment

Time is an essential feature of most decisions, because the reward earned from decisions frequently depends on the temporal statistics of the environment (e.g., on whether decisions must be made under deadlines). Accordingly, evolution appears to have favored a mechanism that predicts intervals in the seconds to minutes range with high accuracy on average, but significant variability from trial to trial. Importantly, the subjective sense of time that results is sufficiently imprecise that maximizing rewards in decision-making can require substantial behavioral adjustments (e.g., accumulating less evidence for a decision in order to beat a deadline). Reward maximization in many daily decisions therefore requires optimal temporal risk assessment. Here, we review the temporal decision-making literature, conduct secondary analyses of relevant published datasets, and analyze the results of a new experiment. The paper is organized in three parts. In the first part, we review literature and analyze existing data suggesting that animals take account of their inherent behavioral variability (their “endogenous timing uncertainty”) in temporal decision-making. In the second part, we review literature that quantitatively demonstrates nearly optimal temporal risk assessment with sub-second and supra-second intervals using perceptual tasks (with humans and mice) and motor timing tasks (with humans). We supplement this section with original research that tested human and rat performance on a task that requires finding the optimal balance between two time-dependent quantities for reward maximization. This optimal balance in turn depends on the level of timing uncertainty. Corroborating the reviewed literature, humans and rats exhibited nearly optimal temporal risk assessment in this task. In the third section, we discuss the role of timing uncertainty in reward maximization in two-choice perceptual decision-making tasks and review literature that implicates timing uncertainty as an important factor in performance quality. Together, these studies strongly support the hypothesis that animals take normative account of their endogenous timing uncertainty. By incorporating the psychophysics of interval timing into the study of reward maximization, our approach bridges empirical and theoretical gaps between the interval timing and decision-making literatures.

P3a and P3b of event-related potentials in persons with Asperger disorder

Although numerous research studies have explored the functional attributes of the human duration perception, the spatio- temporal information on cortical networks underlying this process is still an open question. Moreover, the issue of possible differences in the nature of timing mechanisms responsible for perception of sub- and supra-second intervals requires the implementation of the functional brain imaging techniques with both high spatial and temporal resolution. Attention is very frequently used as a modulating factor for the perceived duration of a sensory stimulus. The non-attended stimuli appear to last shorter than the attended ones, increasing the perceived duration of a concurrent stimulus (Gorea, 2011). This study challenges the accepted model of early sensory responses, activated during the first 100 ms after stimulus presentation, as preattentive, automatic processes which modulate the neural sensitivity to incoming stimuli (gating phenomena). We utilized Elekta Neuromag 306-channel whole-head system for magnetoencephalography (MEG) measurements and multi-dipole Calibrated Start Spatio Temporal (CSST) localization technique (Ranken et al., 2002) to investigate whether voluntary attention directed at the second tone of a pair in the standard paired-click paradigm could affect cortical networks underlying the gating out phenomenon. MEG recordings were obtained in a magnetically shielded room at the Biomagnetic Center in Jena, Germany. Two consecutive (ISI=500 ms) identical short tonnes (S1 and S2; duration=20ms; f=1200 Hz; ITI=8±1s) were used to evoke standard gating cortical responses in 19 healthy participants (21-38 years). In the second condition, the task was to direct attention toward the second tone and to respond to a rarely presented non-identical second tone of the pair (R=1300 Hz, p(S1R)=0.3). MEG recordings showed transient early and middle latency responses with peak amplitudes over the temporo-parietal sensors followed by a peak of a sustained activity. Less prominent transient activity was recorded over the frontal sensors, accompanied with a more steady- state component from 100 until 400 ms post-stimulus for both conditions. Preliminary results of the CSST spatio-temporal analyses revealed 4-6 brain regions activated during 20-500 ms time interval including bilateral superior temporal gyrus (STG), bilateral and medial prefrontal (PF), bilateral parietal (PA) regions, and central motor cortex area. A cortical source underlying the steady-state component was identified in the left prefrontal region for S1 tone, and in central posterior regions (PA) for the repeated tone S2. During the standard paradigm the M50 gating suppression was observed only for bilateral STG sources while bilateral PF and PA sources did not show any reduction in the response during the first 100 ms post-stimulus. Targeting the attention towards the second tone enhanced the M50 amplitude of the bilateral STG responses in respect to the repeated stimulus S2 and caused a change in the activated network evident by the emergence of a new generator in the medial PF area (orbitofrontal) instead of dorsolateral PF generators evoked in non-attended condition. These results demonstrate that voluntary attention can exert a topological and functional modulatory influence on the neural network even during early stages of auditory processing.

Interval timing disruptions in subjects with cerebellar lesions

The cerebellum has long been implicated in time perception, particularly in the subsecond range. The current set of studies examines the role of the cerebellum in suprasecond timing, using analysis of behavioral data in subjects with cerebellar lesions. Eleven cerebellar lesion subjects and 17 controls were tested on temporal estimation, reproduction and production, for times ranging from 2 to 12s. Cerebellar patients overproduced times on both the reproduction and production tasks; the effect was greatest at the shortest duration. A subset of patients also underestimated intervals. Cerebellar patients were significantly more variable on the estimation and reproduction tasks. No significant differences between normal and cerebellar patients were found on temporal discrimination tasks with either sub- or suprasecond times. Patients with damage to the lateral superior hemispheres or the dentate nuclei showed more significant impairments than those with damage elsewhere in the cerebellum, and patients with damage to the left cerebellum had more significant differences from controls than those with damage to the right. These data suggest that damage to the middle-to-superior lobules or the left hemisphere is especially detrimental to timing suprasecond intervals. We suggest that this region be considered part of a network of brain structures including the DLPFC that is crucial for interval timing.

Short-term Memory of Duration: Subjective Shortening or Lengthening

Although many studies revealed a subjective shortening in the human's short-term memory of time,the conclusions remain ambiguous. The present study aimed to explore whether when the different duration (suband supra-second intervals) is retained in short-term memory,the duration becomes subjective shortening or lengthening as retention interval increased. Based on the method of Wearden and Ferrara,Two experiments ( Experiment 1,2) were used to investigate the retention effect of temporal intervals from 350ms to 650ms and from 1000ms to 2000ms respectively,and each experiment included two sub-experiments which separately examined the duration retention of the auditory presentation (Experiment 1a,2a) and that of the visual presentation (Experiment 1b,2b). 2(stimulus type:filled duration and unfilled duration) ×4 (delay time:1s,4s,8s and 12s) ×3 (trial type:short trial,equal trial and long trial) within-subject factorial design was taken separately for every sub-experiment (Experiment 1a,2a,1b and 2b). All participants were required to respond by pressing one of the three keys after judging whether the comparison was longer,shorter,or of the same duration as the standard duration. The results of the two experiments showed that,the short-term retention of durations from 350ms to 650ms (Experiment 1) indicated subjective shortening both in auditory (Experiment 1a) and visual conditions (Experiment 1b),whereas the short-term retention of durations from 1000ms to 2000ms (Experiment 2) showed subjective lengthening both in auditory (Experiment 2a) and visual conditions (Experiment 2b),and there was neither effect of modality nor the effect of stimulus type that influenced the retention effect. Conclusions were drawn from this study. For Experiment 1 (several hundred milliseconds),the shortening trend resulted from the interaction of shrink of the positive time order error (TOE) and the effect of subjective shortening. By contrast,for Experiment 2 (the retention of durations above 1s),the lengthening trend resulted from the interaction of shrink of the negative TOE effect and the effect of subjective lengthening. According to these results,the present study compensated and perfected the relative duration hypothesis and the partial processing feature of the scalar timing model. However,apart from duration length and stimulus type,other factors such as experimental methods are likely to affect time duration retention as well,and whether these factors are independent or interacted needs to be investigated in future research.