The internal clock: Electroencephalographic evidence for oscillatory processes underlying time perception

It has been proposed that temporal perception and performance depend on a biological source of temporal information. A model for a temporal oscillator put forward by Treisman, Faulkner, Naish, and Brogan (1990) predicted that if intense sensory pulses (such as auditory clicks) were presented to subjects at suitable rates they would perturb the frequency at which the temporal oscillator runs and so cause over- or underestimation of time. The resulting pattern of interference between sensory pulse rates and time judgments would depend on the frequency of the temporal oscillator and so might allow that frequency to be estimated. Such interference patterns were found using auditory clicks and visual flicker (Treisman & Brogan, 1992; Treisman et al., 1990). The present study examines time estimation together with the simultaneously recorded electroencephalogram to examine whether evidence of such an interference pattern can be found in the EEG.

Alternative models for the organization of a temporal system consisting of an oscillator or multiple oscillators are considered and predictions derived from them relating to the EEG. An experiment was run in which time intervals were presented for estimation, auditory clicks being given during those intervals, and the EEG was recorded concurrently. Analyses of the EEG revealed interactions between auditory click rates and certain EEG components which parallel the interference patterns previously found. The overall pattern of EEG results is interpreted as favouring a model for the organization of the temporal system in which sets of click-sensitive oscillators spaced at intervals of about 12.8 Hz contribute to the EEG spectrum. These are taken to represent a series of harmonically spaced distributions of oscillators involved in time-keeping.     

On the Relation Between Time Perception and the Timing of Motor Action: Evidence for a Temporal Oscillator Controlling the Timing of Movement

Studies of time estimation have provided evidence that human time perception is determined by an internal clock containing a temporal oscillator and have also provided estimates of the frequency of this oscillator (Treisman, Faulkner, Naish, & Brogan, 1992; Treisman & Brogan, 1992). These estimates were based on the observation that when the intervals to be estimated are accompanied by auditory clicks that recur at certain critical rates, perturbations in time estimation occur. To test the hypothesis that the mechanisms that underlie the perception of time and those that control the timing of motor performance are similar, analogous experiments were performed on motor timing, with the object of seeing whether evidence for a clock would be obtained and if so whether its properties resemble those of the time perception clock. The prediction was made that perturbations in motor timing would be seen at the same or similar critical auditory click rates. The experiments examined choice reaction time and typing. The results support the hypothesis that a temporal oscillator paces motor performance and that this oscillator is similar to the oscillator underlying time perception. They also provide an estimate of the characteristic frequency of the oscillator.     

The internal clock: evidence for a temporal oscillator underlying time perception with some estimates of its characteristic frequency

Evidence for the proposition that human time perception is determined by an internal clock is largely indirect. It would strengthen the case for this hypothesis if a model for the internal clock were available from which predictions could be derived and tested, and if the basic parameter of such a model, the frequency at which the clock runs, could be estimated. A model for an internal temporal pacemaker is briefly described and its properties are explored by computer simulation. Results are obtained that provide a basis for predicting that, under appropriate conditions, interference between an imposed rhythm and the frequency of a temporal oscillator may cause perturbations in temporal judgment which are related to the characteristic frequency of that oscillator. Experimental data are reported which appear to demonstrate such an interference pattern. These results allow some estimates of the characteristic frequency of the temporal oscillator to be obtained.     

The contribution of rapid visual and auditory processing to the reading of irregular words and pseudowords presented singly and in contiguity

This study examined the relative involvement of rapid auditory and visual temporal resolution mechanisms in the reading of phonologically regular pseudowords and English irregular words presented both in isolation and in contiguity as a series of six words. Seventy-nine undergraduates participated in a range of reading, visual temporal, and auditory temporal tasks. The correlation analyses suggested a general timing mechanism across modalities. There were more significant correlations between the visual temporal measures and irregular word reading and between the auditory measures and pseudoword reading. Auditory gap detection predicted pseudoword reading accuracies. The low temporal frequency flicker contrast sensitivity measure predicted the accuracies of isolated irregular words and pseudowords presented in contiguity. However, when a combined speed-accuracy score was used, visible persistence at both low and high spatial frequencies and auditory gap detection were active in the reading of pseudowords presented in contiguity. Sensory processing skills in both visual and auditory modalities accounted for some of the variance in the reading performance of normal undergraduates, not just reading-impaired students.     

The effects of age on sensory thresholds and temporal gap detection in hearing, vision, and touch

Differences in sensory function between young (n 5 42, 18—31 years old) and older (n 5 137, 60—88 years old) adults were examined for auditory, visual, and tactile measures of threshold sensitivity and temporal acuity (gap-detection threshold). For all but one of the psychophysical measures (visual gap detection), multiple measures were obtained at different stimulus frequencies for each modality and task. This resulted in a total of 14 dependent measures, each based on four to six adaptive psychophysical estimates of 75% correct performance. In addition, all participants completed the Wechsler Adult Intelligence Scale (Wechsler, 1997). Mean data confirmed previously observed differences in performance between young and older adults for 13 of the 14 dependent measures (all but visual threshold at a flicker frequency of 4 Hz). Correlational and principalcomponents factor analyses performed on the data from the 137 older adults were generally consistent with task and modality independence of the psychophysical measures.     

Auditory and visual differences in time perception

The present experiment assessed intersensory differences in temporal judgments, that is, auditory stimuli are perceived as longer than physically equivalent visual stimuli. The results confirmed the intersensory difference. Auditorially defined intervals were experienced as longer than visually defined intervals. Auditory boundaries were perceived as longer than visual ones. An interaction of boundary modality and interval modality was obtained which suggested that auditorially defined intervals provided more temporal information about events occurring in close temporal proximity than visually defined intervals. It was hypothesized that cognitive factors, specifically stimulus complexity, would affect the auditory and visual systems differentially. This hypothesis was not substantiated, although highly complex stimuli were experienced as longer than those of low complexity.     

The rhythm aftereffect: support for time sensitive neurons with broad overlapping tuning curves

Ivry [Ivry, R. B. (1996). The representation of temporal information in perception and motor control. Current Opinion in Neurobiology, 6, 851-857.] proposed that explicit coding of brief time intervals is accomplished by neurons that are tuned to a preferred temporal interval and have broad overlapping tuning curves. This proposal is analogous to the orientation selective cells in visual area V1. To test this proposal, we used a temporal analog to the visual tilt aftereffect. After adapting to a fast auditory rhythm, a moderately fast test rhythm (400 ms between beats) seemed slow and vice versa. If the speed of the adapting rhythm was made too disparate from speed of the test rhythm the effect was diminished. The effect occurred whether the adapting and test stimuli were presented to the same or different ears, but did not occur when an auditory adapting rhythm was followed by a visual test rhythm. Results support the proposition that explicit time information is coded by neural units tuned to specific temporal intervals with broad overlapping tuning curves. In addition, it appears that there is a single timing mechanism for each incoming sensory mode, but distinct timers for different modes.     

Audiovisual beat induction in complex auditory rhythms: Point-light figure movement as an effective visual beat

This study investigated whether explicit beat induction in the auditory, visual, and audiovisual (bimodal) modalities aided the perception of weakly metrical auditory rhythms, and whether it reinforced attentional entrainment to the beat of these rhythms. The visual beat-inducer was a periodically bouncing point-light figure, which aimed to examine whether an observed rhythmic human movement could induce a beat that would influence auditory rhythm perception. In two tasks, participants listened to three repetitions of an auditory rhythm that were preceded and accompanied by (1) an auditory beat, (2) a bouncing point-light figure, (3) a combination of (1) and (2) synchronously, or (4) a combination of (1) and (2), with the figure moving in anti-phase to the auditory beat. Participants reproduced the auditory rhythm subsequently (Experiment 1), or detected a possible temporal change in the third repetition (Experiment 2). While an explicit beat did not improve rhythm reproduction, possibly due to the syncopated rhythms when a beat was imposed, bimodal beat induction yielded greater sensitivity to a temporal deviant in on-beat than in off-beat positions. Moreover, the beat phase of the figure movement determined where on-beat accents were perceived during bimodal induction. Results are discussed with regard to constrained beat induction in complex auditory rhythms, visual modulation of auditory beat perception, and possible mechanisms underlying the preferred visual beat consisting of rhythmic human motions.     

节律性时间期待效应不受注意控制的影响

采用双任务范式探讨当听觉节律刺激序列以较慢速度呈现时,其诱导产生的时间期待效应是否受到同时进行的视觉工作记忆任务的影响。结果发现,无论目标刺激是呈现在听觉通道还是视觉通道,双任务和单任务条件下目标刺激出现在规律听觉刺激序列之后被试的反应时均快于目标出现在非规律听觉刺激序列之后,即节律性刺激序列诱导产生的时间期待效应不受工作记忆任务的影响。该结果表明节律性时间期待效应不受注意控制的影响。     

Evidence for auditory temporal distinctiveness: modality effects in order and frequency judgments

Two new, long-lasting phenomena involving modality of stimulus presentation are documented. In one series of experiments we investigated effects of modality of presentation on order judgments. Order judgments for auditory words were more accurate than order judgments for visual words at both the beginning and the end of lists, and the auditory advantage increased with the temporal separation of the successive items. A second series of experiments investigated effects of modality on estimates of presentation frequency. Frequency estimates of repeated auditory words exceeded frequency estimates of repeated visual words. The auditory advantage increased with frequency of presentation, and this advantage was not affected by the retention interval. These various effects were taken as support for a temporal coding assumption, that auditory presentation produces a more accurate encoding of time of presentation than does visual presentation.     

Individual differences in vulnerability to subjective time distortion1

Time duration is perceived to be longer when accompanied by dynamic sensory stimulation than when accompanied by static stimulation. This distortion of time perception is thought to be due to the acceleration of an internal pacemaker that has been assumed to be the main component of temporal judgments. In order to investigate whether the function of the internal pacemaker is modality dependent or independent, we examined the correlation of visual flicker and auditory flutter effects on a temporal production task. While seeing a 10‐Hz visual flicker or hearing a 10‐Hz auditory flutter, participants estimated a duration of 2500 ms as accurately as possible by pressing a button. The results showed a significant within‐individual correlation between the time distortion due to visual flicker and that due to auditory flutter. Additionally, we found that time distortion due to auditory flutter tended to be larger in female participants than in male participants. These results suggest that the mechanisms underlying subjective time dilation are similar between vision and audition within individuals, but that they vary across individuals.     

Intersensory redundancy guides attentional selectivity and perceptual learning in infancy

This study assessed an intersensory redundancy hypothesis, which holds that in early infancy information presented redundantly and in temporal synchrony across two sense modalities selectively recruits attention and facilitates perceptual differentiation more effectively than does the same information presented unimodally. Five-month-old infants' sensitivity to the amodal property of rhythm was examined in 3 experiments. Results revealed that habituation to a bimodal (auditory and visual) rhythm resulted in discrimination of a novel rhythm, whereas habituation to the same rhythm presented unimodally (auditory or visual) resulted in no evidence of discrimination. Also, temporal synchrony between the bimodal auditory and visual information was necessary for rhythm discrimination. These findings support an intersensory redundancy hypothesis and provide further evidence for the importance of redundancy for guiding and constraining early perceptual learning.     

Visual beats: Preliminary observations of perceived rate as a function of retinal locus stimulated

Visual beats produced by simultaneous presentation of two different flicker frequencies were used to determine the temporal processing of various retinal areas. The dichoptic presentation of two different flicker frequencies to the center and periphery of the retina resulted in a perceived beat rate that equalled the physical beat frequency. When the stimuli were presented to more discrete retinal areas (temporal vs temporal and nasal vs temporal), the perceived beat rate also equalled the physical beat frequency. The data indicate that different flicker frequencies stimulating divergent retinal loei can be represented accurately and simultaneously in the visual system: Differences in temporal resolution at the different retinal loei do not occur for this repetitive stimulation.     

Auditory rhythms are systemically associated with spatial-frequency and density information in visual scenes

A variety of perceptual correspondences between auditory and visual features have been reported, but few studies have investigated how rhythm, an auditory feature defined purely by dynamics relevant to speech and music, interacts with visual features. Here, we demonstrate a novel crossmodal association between auditory rhythm and visual clutter. Participants were shown a variety of visual scenes from diverse categories and asked to report the auditory rhythm that perceptually matched each scene by adjusting the rate of amplitude modulation (AM) of a sound. Participants matched each scene to a specific AM rate with surprising consistency. A spatial-frequency analysis showed that scenes with greater contrast energy in midrange spatial frequencies were matched to faster AM rates. Bandpass-filtering the scenes indicated that greater contrast energy in this spatial-frequency range was associated with an abundance of object boundaries and contours, suggesting that participants matched more cluttered scenes to faster AM rates. Consistent with this hypothesis, AM-rate matches were strongly correlated with perceived clutter. Additional results indicated that both AM-rate matches and perceived clutter depend on object-based (cycles per object) rather than retinal (cycles per degree of visual angle) spatial frequency. Taken together, these results suggest a systematic crossmodal association between auditory rhythm, representing density in the temporal domain, and visual clutter, representing object-based density in the spatial domain. This association may allow for the use of auditory rhythm to influence how visual clutter is perceived and attended.     

Flicker-induced color and form: interdependencies and relation to stimulation frequency and phase

Our understanding of human visual perception generally rests on the assumption that conscious visual states represent the interaction of spatial structures in the environment and our nervous system. This assumption is questioned by circumstances where conscious visual states can be triggered by external stimulation which is not primarily spatially defined. Here, subjective colors and forms are evoked by flickering light while the precise nature of those experiences varies over flicker frequency and phase. What's more, the occurrence of one subjective experience appears to be associated with the occurrence of others. While these data indicate that conscious visual experience may be evoked directly by particular variations in the flow of spatially unstructured light over time, it must be assumed that the systems responsible are essentially temporal in character and capable of representing a variety of visual forms and colors, coded in different frequencies or at different phases of the same processing rhythm.     

Transient increase of intact visual field size by high-frequency narrow-band stimulation

Three patients with visual field defects were stimulated with a square matrix pattern, either static, or flickering at frequencies that had been found to either promote or not promote blindsight performance. Comparison between pre- and post-stimulation perimetric maps revealed an increase in the size of the intact visual field but only for flicker frequencies previously found to promote blindsight. These changes were temporary but dramatic – in two instances the intact field was increased by an area of ∼30 deg² of visual angle. These results indicate that not only does specific high-frequency stimulus flicker promote blindsight, but that intact visual field size may be increased by stimulation at the same frequencies. Our findings inform speculation on both the brain mechanisms and the potency of temporal modulation for altering the functional visual field.     

A new visual illusion: flickering fields are localized in a depth plane behind nonflickering fields

Flickering regions of the visual field are perceived to lie well behind regions which are not flickered. The depth segregation is not due to luminance differences since the average temporal luminance across all the regions was equal. This depth effect produced by flicker is not dependent on the texture of the visual field; nor does it depend on a specific configuration of the flickering and nonflickering areas. It is optimal at a temporal frequency around 6 Hz, which suggests that visual channels responding maximally to high temporal frequencies are involved in the segregation of perceptual regions in depth.     

Twice upon a time: multiple concurrent temporal recalibrations of audiovisual speech

Audiovisual timing perception can recalibrate following prolonged exposure to asynchronous auditory and visual inputs. It has been suggested that this might contribute to achieving perceptual synchrony for auditory and visual signals despite differences in physical and neural signal times for sight and sound. However, given that people can be concurrently exposed to multiple audiovisual stimuli with variable neural signal times, a mechanism that recalibrates all audiovisual timing percepts to a single timing relationship could be dysfunctional. In the experiments reported here, we showed that audiovisual temporal recalibration can be specific for particular audiovisual pairings. Participants were shown alternating movies of male and female actors containing positive and negative temporal asynchronies between the auditory and visual streams. We found that audiovisual synchrony estimates for each actor were shifted toward the preceding audiovisual timing relationship for that actor and that such temporal recalibrations occurred in positive and negative directions concurrently. Our results show that humans can form multiple concurrent estimates of appropriate timing for audiovisual synchrony.     

Hearing what the eyes see: auditory encoding of visual temporal sequences

When the senses deliver conflicting information, vision dominates spatial processing, and audition dominates temporal processing. We asked whether this sensory specialization results in cross-modal encoding of unisensory input into the task-appropriate modality. Specifically, we investigated whether visually portrayed temporal structure receives automatic, obligatory encoding in the auditory domain. In three experiments, observers judged whether the changes in two successive visual sequences followed the same or different rhythms. We assessed temporal representations by measuring the extent to which both task-irrelevant auditory information and task-irrelevant visual information interfered with rhythm discrimination. Incongruent auditory information significantly disrupted task performance, particularly when presented during encoding; by contrast, varying the nature of the rhythm-depicting visual changes had minimal impact on performance. Evidently, the perceptual system automatically and obligatorily abstracts temporal structure from its visual form and represents this structure using an auditory code, resulting in the experience of "hearing visual rhythms."     

Brightness enhancement and the Talbot level in stationary gratings

At low spatial frequencies, the perceived brightness of the light phase of a stationary square-wave grating is greater than the brightness of a solid field of equal physical luminance. That increase in the perceived brightness of a grating at low spatial frequencies is analogous to the brightness enhancement observed in a flickering light at low temporal frequencies. At or above the critical spatial frequency—the visual resolution threshold—the brightness of a grating is determined by its space-average luminance, just as the brightness of a flickering light at or above the critical flicker frequency is determined by its time-average luminance in accordance with Talbot’s law. Thus, Talbot’s law applies in the spatial as well as the temporal domain. The present study adds to the evidence that temporal and spatial frequency play analogous roles in some aspects of brightness vision.