Visuomotor synchronization in adults and seven- to eight-year-old children

A group of 22 seven- to eight-year-old children and a group of 17 adults participated in the experiment in which they synchronized their movements (pressing a button) with an isochronous sequence of visual stimuli. The period of the sequence was varied between 500 to 2000 ms at a step of 300 ms. Two consecutive phases of visuomotor synchronization were studied: the initiation phase, which corresponds to the process of transition between responding to a visual stimulus to stable synchronization with them, and the synchronization phase. The initiation phase was characterized by the shape and duration of the asynchrony time course (relaxation curve). The statistical properties of asynchrony were analyzed in terms of the phase correction of the central timer. It was shown that (1) the range of successful visuomotor synchronization was narrower in seven- to eight-year-old children than in adults (from 600–700 ms to ∼1700 ms); (2) the initiation phase duration was about the same in seven- to eight-year-old children and adults, and typical shapes of individual relaxation curves were similar in both adults and children; (3) although the statistical properties of asynchrony were comparable in children and adults, the mechanism of phase correction of the central timer operated at a lower correction gain factor in children than in adults. In children, the phase correction process was also characterized by a substantially higher level of the central and motor noise, which leads to higher asynchrony variability and more frequent and longer synchronization losses.     

Visuo-motor synchronization in children of 7-8 years old

The group of 22 children of 7-8 years old and the group of 17 adults participated in the experiment in which they were asked to synchronize their movements (pressing a button) with an isochronous sequence of visual stimuli. The period of the sequence was varied between 500 to 2000 ms with the 300 ms step. Two successive phases of visuo-motor synchronization were studied: the synchronization phase and the initiation phase which corresponds to the process of transition between reacting to a visual stimulus and the stable synchronization. The initiation phase was characterized by the shape and duration of the asynchrony time course (relaxation curve). The statistical properties of asynchrony were analyzed in the framework of the phase correction of the central timer. It is shown that (1) the range of successful visuo-motor synchronization is narrower in children of 7-8 than in adults and it spans from 600-700 ms to approx. 1700 Mc; (2) The initiation phase lasts about the same time in children of 7-8 and adults and typical shapes of individual relaxation curves are similar in both adults and children. (3) Although the statistical properties of asynchrony are comparable in children and adults, the mechanism of phase correction of the central timer operates with a lower value of the correction gain factor in children than in adults. In children, the phase correction process is also characterized by a substantially higher level of the central and motor noise which leads to a higher asynchrony variability and more frequent and longer lasting synchronization losses.     

A model of synchronization of motor acts to a stimulus sequence

A closed-loop timing model is proposed that accounts for several phenomena observed in tasks which require production of a sequence of motor acts in synchrony with a sequence of stimuli. In contrast to the previous models, variables available to the central nervous system of a subject (internal variables) and externally measurable variables are distinguished, and several physiologically justifiable internal variables are included. The model assumes the existence of (a) an internal time-keeper producing a reference interval that is used in a motor-control unit for timing of the next motor command; (b) an intrinsic (subjective) synchrony that relies on some a posteriori (feedback) information about the already executed onset of the motor act. A two-way error-corrective mechanism is hypothesized: (1) period (inverted frequency) corrections — the reference interval (period) is set at the beginning of the task according to the interstimulus-onset interval (s) and later corrected for differences between its duration and the actual duration of s; (2) phase corrections — internal synchronization errors (i.e., time gaps between the central temporal availability of internal representations of stimuli and of some feedback aspect of responses) are corrected for directly in the motor-control unit. Objectively measured systematic asynchrony of responses and stimuli is determined by the internal delays in information transduction. Finally, the model is used for making predictions of a subject's performance in some other experimental settings of the synchronization task.The core of this study was presented at the 4th Workshop on Rhythm Perception and Production, June 1992, Bourges, France (Mates 1992)     

Scene segmentation by spike synchronization in reciprocally connected visual areas. I. Local effects of cortical feedback

 To investigate scene segmentation in the visual system we present a model of two reciprocally connected visual areas using spiking neurons. Area P corresponds to the orientation-selective subsystem of the primary visual cortex, while the central visual area C is modeled as associative memory representing stimulus objects according to Hebbian learning. Without feedback from area C, a single stimulus results in relatively slow and irregular activity, synchronized only for neighboring patches (slow state), while in the complete model activity is faster with an enlarged synchronization range (fast state). When presenting a superposition of several stimulus objects, scene segmentation happens on a time scale of hundreds of milliseconds by alternating epochs of the slow and fast states, where neurons representing the same object are simultaneously in the fast state. Correlation analysis reveals synchronization on different time scales as found in experiments (designated as tower, castle, and hill peaks). On the fast time scale (tower peaks, gamma frequency range), recordings from two sites coding either different or the same object lead to correlograms that are either flat or exhibit oscillatory modulations with a central peak. This is in agreement with experimental findings, whereas standard phase-coding models would predict shifted peaks in the case of different objects. Received: 22 August 2001 / Accepted in revised form: 8 April 2002     

Variability in 5- and 8-year-olds' memory for duration: an interfering task in temporal bisection

The purpose of this experiment was to investigate 5- and 8-year-olds' long-term memory for stimulus duration in a bisection task. Children were trained to discriminate between a short and a long standard duration presented as visual stimulus for 2 and 8 s, respectively. They had then to decide whether an intermediate stimulus duration was more similar to the short or to the long standard in two identical testing phases separated by an interfering task lasting for 15 min (immediate test vs. deferred test). The results showed that the 5- and the 8-year-olds produced orderly psychophysical functions. However, the 8-year-olds produced psychophysical functions, which increased more abruptly with the increasing stimulus duration. Nevertheless, whatever the age of children tested, the psychophysical function curves were flatter in the deferred test than in the immediate test, and more particularly in the 5-year-olds. Furthermore, the 5-year-olds produced fewer 'long' responses in the deferred than in the immediate test. Modeling of the data suggests that the variability of the representation of the standard in long-term memory was higher in the 5- than in the 8-year-olds and that the interfering task increased this memory variability.     

Evidence for dissociation between the perceptual and visuomotor systems in humans

When a visual stimulus is continuously moved behind a small stationary window, the window appears displaced in the direction of motion of the stimulus. In this study we showed that the magnitude of this illusion is dependent on (i) whether a perceptual or visuomotor task is used for judging the location of the window (ii) the directional signature of the stimulus, and (iii) whether or not there is a significant delay between the end of the visual presentation and the initiation of the localization measure. Our stimulus was a drifting sinusoidal grating windowed in space by a stationary, two-dimensional, Gaussian envelope (sigma=1 cycle of sinusoid). Localization measures were made following either a short (200 ms) or long (4.2 s) post-stimulus delay. The visuomotor localization error was up to three times greater than the perceptual error for a short delay. However, the visuomotor and perceptual localization measures were similar for a long delay. Our results provide evidence in support of the hypothesis that separate cortical pathways exist for visual perception and visually guided action and that delayed actions rely on stored perceptual information.     

Reduced variability of ongoing and evoked cortical activity leads to improved behavioral performance

Sensory responses of the brain are known to be highly variable, but the origin and functional relevance of this variability have long remained enigmatic. Using the variable foreperiod of a visual discrimination task to assess variability in the primate cerebral cortex, we report that visual evoked response variability is not only tied to variability in ongoing cortical activity, but also predicts mean response time. We used cortical local field potentials, simultaneously recorded from widespread cortical areas, to gauge both ongoing and visually evoked activity. Trial-to-trial variability of sensory evoked responses was strongly modulated by foreperiod duration and correlated both with the cortical variability before stimulus onset as well as with response times. In a separate set of experiments we probed the relation between small saccadic eye movements, foreperiod duration and manual response times. The rate of eye movements was modulated by foreperiod duration and eye position variability was positively correlated with response times. Our results indicate that when the time of a sensory stimulus is predictable, reduction in cortical variability before the stimulus can improve normal behavioral function that depends on the stimulus.     

Synchronization of neuronal assemblies in reciprocally connected cortical areas

Summary To investigate scene segmentation in the visual system we present a model of two reciprocally connected visual areas comprising spiking neurons. The peripheral area P is modeled similar to the primary visual cortex, while the central area C is modeled as an associative memory representing stimulus objects according to Hebbian learning. Without feedback from area C, spikes corresponding to stimulus representations in P are synchronized only locally (slow state). Feedback from C can induce fast oscillations and an increase of synchronization ranges (fast state). Presenting a superposition of several stimulus objects, scene segmentation happens on a time scale of hundreds of milliseconds by alternating epochs of the slow and fast state, where neurons representing the same object are simultaneously in the fast state. We relate our simulation results to various phenomena observed in neurophysiological experiments, such as stimulus-dependent synchronization of fast oscillations, synchronization on different time scales, ongoing activity, and attention-dependent neural activity.     

The role of phase synchronization in memory processes

In recent years, studies ranging from single-unit recordings in animals to electroencephalography and magnetoencephalography studies in humans have demonstrated the pivotal role of phase synchronization in memory processes. Phase synchronization - here referring to the synchronization of oscillatory phases between different brain regions - supports both working memory and long-term memory and acts by facilitating neural communication and by promoting neural plasticity. There is evidence that processes underlying working and long-term memory might interact in the medial temporal lobe. We propose that this is accomplished by neural operations involving phase-phase and phase-amplitude synchronization. A deeper understanding of how phase synchronization supports the flexibility of and interaction between memory systems may yield new insights into the functions of phase synchronization in general.     

Brightness induction: rate enhancement and neuronal synchronization as complementary codes

In cat visual cortex, we investigated with parallel recordings from multiple units the neuronal correlates of perceived brightness. The perceived brightness of a center grating was changed by varying the orientation or the relative spatial phase of a surrounding grating. Brightness enhancement by orientation contrast is associated with an increase of discharge rates of responses to the center grating but not with changes in spike synchronization. In contrast, if brightness enhancement is induced by phase offset, discharge rates are unchanged but synchronization increases between neurons responding to the center grating. The changes in synchronization correlate well with changes in perceived brightness that were assessed in parallel in human subjects using the same stimuli. These results indicate that in cerebral cortex the modulation of synchronicity of responses is used as a mechanism complementary to rate changes to enhance the saliency of neuronal responses.     

A quantitative synchronization model for smooth pursuit target tracking

We propose a quantitative model for human smooth pursuit tracking of a continuously moving visual target which is based on synchronization of an internal expectancy model of the target position coupled to the retinal target signal. The model predictions are tested in a smooth circular pursuit eye tracking experiment with transient target blanking of variable duration. In subjects with a high tracking accuracy, the model accounts for smooth pursuit and repeatedly reproduces quantitatively characteristic patterns of the eye dynamics during target blanking. In its simplest form, the model has only one free parameter, a coupling constant. An extended model with a second parameter, a time delay or memory term, accounts for predictive smooth pursuit eye movements which advance the target. The model constitutes an example of synchronization of a complex biological system with perceived sensory signals. Cognitive and Neurobiological Research Consortium in Traumatic Brain Injury (CNRC-TBI).     

Directed corticocortical functional connectivity at the early stages of serial learning in adults and seven- to eight-year-old children

Serial learning at its earlier stages, presumably involving the working memory, was studied in adults and seven- to eight-year-old children during the reproduction of a sequence of discrete movements following the order specified by a sequence of visual stimuli. In both age groups, the learning curves (latent time vs. trial number) were qualitatively similar in shape. The overall shape of the learning curve depended on the relative proportion of the fast vs. slow phases of latent time reduction. Comparison of the corticocortical functional connectivity patterns in the prestimulus period in the sequence reproduction task vs. the simple visuomotor reaction task showed a general tendency of an increase in the influence of postcentral cortical areas accompanied by the reduced influence of prefrontal and central cortical areas. In particular, it was typical of adults to show an increase in the directed influence of temporo-parieto-occipital (TPO) cortical areas, while the children also showed an increase in the directed influence of the parietal cortex. Comparison of the subgroups with different shapes of learning curves in the prestimulus period has shown the difference in their patterns of directed functional connectivity. The results are discussed with a special emphasis on the role of the working memory retaining the internal representations of sequences being learned.     

Neural synchrony in brain disorders: relevance for cognitive dysfunctions and pathophysiology

Following the discovery of context-dependent synchronization of oscillatory neuronal responses in the visual system, novel methods of time series analysis have been developed for the examination of task- and performance-related oscillatory activity and its synchronization. Studies employing these advanced techniques revealed that synchronization of oscillatory responses in the beta- and gamma-band is involved in a variety of cognitive functions, such as perceptual grouping, attention-dependent stimulus selection, routing of signals across distributed cortical networks, sensory-motor integration, working memory, and perceptual awareness. Here, we review evidence that certain brain disorders, such as schizophrenia, epilepsy, autism, Alzheimer's disease, and Parkinson's are associated with abnormal neural synchronization. The data suggest close correlations between abnormalities in neuronal synchronization and cognitive dysfunctions, emphasizing the importance of temporal coordination. Thus, focused search for abnormalities in temporal patterning may be of considerable clinical relevance.     

Fractal Gait Patterns Are Retained after Entrainment to a Fractal Stimulus

Previous work has shown that fractal patterns in gait can be altered by entraining to a fractal stimulus. However, little is understood about how long those patterns are retained or which factors may influence stronger entrainment or retention. In experiment one, participants walked on a treadmill for 45 continuous minutes, which was separated into three phases. The first 15 minutes (pre-synchronization phase) consisted of walking without a fractal stimulus, the second 15 minutes consisted of walking while entraining to a fractal visual stimulus (synchronization phase), and the last 15 minutes (post-synchronization phase) consisted of walking without the stimulus to determine if the patterns adopted from the stimulus were retained. Fractal gait patterns were strengthened during the synchronization phase and were retained in the post-synchronization phase. In experiment two, similar methods were used to compare a continuous fractal stimulus to a discrete fractal stimulus to determine which stimulus type led to more persistent fractal gait patterns in the synchronization and post-synchronization (i.e., retention) phases. Both stimulus types led to equally persistent patterns in the synchronization phase, but only the discrete fractal stimulus led to retention of the patterns. The results add to the growing body of literature showing that fractal gait patterns can be manipulated in a predictable manner. Further, our results add to the literature by showing that the newly adopted gait patterns are retained for up to 15 minutes after entrainment and showed that a discrete visual stimulus is a better method to influence retention.     

Spatial and temporal structure of phase synchronization of spontaneous alpha EEG activity

Spatiotemporal characteristics of spontaneous alpha EEG activity patterns are analyzed in terms of large-scale phase synchronization. During periods with strong phase synchronization over the entire scalp, phase patterns take either of two forms; one is a gradual phase shift between frontal and occipital regions and the other is a stepwise pattern with a sudden phase shift in the central region. The former is regarded as a traveling wave of electrocortical activity, of which the direction of propagation is predominantly from anterior to posterior in three out of four subjects, and opposite in the remaining one. The other activity pattern observed may correspond to a standing wave composed of two traveling waves propagating in opposite directions. The duration distributions of these patterns have similar forms within a subject, which suggests that they share the same mechanism for their generation.     

Oscillatory synchronization in large-scale cortical networks predicts perception

Normal brain function requires the dynamic interaction of functionally specialized but widely distributed cortical regions. Long-range synchronization of oscillatory signals has been suggested to mediate these interactions within large-scale cortical networks, but direct evidence is sparse. Here we show that oscillatory synchronization is organized in such large-scale networks. We implemented an analysis approach that allows for imaging synchronized cortical networks and applied this technique to EEG recordings in humans. We identified two networks: beta-band synchronization (~20 Hz) in a fronto-parieto-occipital network and gamma-band synchronization (~80 Hz) in a centro-temporal network. Strong perceptual correlates support their functional relevance: the strength of synchronization within these networks predicted the subjects' perception of an ambiguous audiovisual stimulus as well as the integration of auditory and visual information. Our results provide evidence that oscillatory neuronal synchronization mediates neuronal communication within frequency-specific, large-scale cortical networks.     

Visual working memory load-related changes in neural activity and functional connectivity

Background

Visual working memory (VWM) helps us store visual information to prepare for subsequent behavior. The neuronal mechanisms for sustaining coherent visual information and the mechanisms for limited VWM capacity have remained uncharacterized. Although numerous studies have utilized behavioral accuracy, neural activity, and connectivity to explore the mechanism of VWM retention, little is known about the load-related changes in functional connectivity for hemi-field VWM retention.

Methodology/Principal Findings

In this study, we recorded electroencephalography (EEG) from 14 normal young adults while they performed a bilateral visual field memory task. Subjects had more rapid and accurate responses to the left visual field (LVF) memory condition. The difference in mean amplitude between the ipsilateral and contralateral event-related potential (ERP) at parietal-occipital electrodes in retention interval period was obtained with six different memory loads. Functional connectivity between 128 scalp regions was measured by EEG phase synchronization in the theta- (4–8 Hz), alpha- (8–12 Hz), beta- (12–32 Hz), and gamma- (32–40 Hz) frequency bands. The resulting matrices were converted to graphs, and mean degree, clustering coefficient and shortest path length was computed as a function of memory load. The results showed that brain networks of theta-, alpha-, beta-, and gamma- frequency bands were load-dependent and visual-field dependent. The networks of theta- and alpha- bands phase synchrony were most predominant in retention period for right visual field (RVF) WM than for LVF WM. Furthermore, only for RVF memory condition, brain network density of theta-band during the retention interval were linked to the delay of behavior reaction time, and the topological property of alpha-band network was negative correlation with behavior accuracy.

Conclusions/Significance

We suggest that the differences in theta- and alpha- bands between LVF and RVF conditions in functional connectivity and topological properties during retention period may result in the decline of behavioral performance in RVF task.     

Dynamics of induced EEG in the intervals between a target and trigger stimuli during formation of a set to emotionally negative facial expression

A visual set was used as a model to study the influence of the increased memory load on the recognition of facial expression in 70 healthy adults. In order to additionally load the working memory, we lengthened the time gap between target (faces) and trigger stimuli. Such a lengthening from 1 to 8 s resulted in an increase of set plasticity (fewer mistakes in facial expression recognition). It also led to a reduction of the reaction time and less number of contrast illusions in recognition. We analyzed theta- and alpha-band EEG changes during individual segments of the time gap and suggested that repeated trials with a certain fixed interval between stimuli formed an inner representation of the interval duration. This inner representation up-regulates the visual attention in case of anticipation of a relevant event (stimulus) and down-regulates the attention when the stimulus is not expected. In case of the plastic set, the induced EEG synchronization in the alpha band is stronger in the trials with correct recognition in the middle of the inter-stimulus time gap. We think this synchronization reflects the action of the top-down cognitive control that suppresses the influence of irrelevant information on the brain activity. Theta-band dynamics in the inter-stimulus time gap can be associated with the emotional strain caused by the fact that a person had to retain in memory (for several seconds) the result of facial expression recognition.     

Characteristics of Human Luminance Discrimination and Modeling a Neural Network Based on the Response Properties of the Visual Cortex

Reaction time (RT) and error rate that depend on stimulus duration were measured in a luminance-discrimination reaction time task. Two patches of light with different luminance were presented to participants for ‘short’ (150 ms) or ‘long’ (1 s) period on each trial. When the stimulus duration was ‘short’, the participants responded more rapidly with poorer discrimination performance than they did in the longer duration. The results suggested that different sensory responses in the visual cortices were responsible for the dependence of response speed and accuracy on the stimulus duration during the luminance-discrimination reaction time task. It was shown that the simple winner-take-all-type neural network model receiving transient and sustained stimulus information from the primary visual cortex successfully reproduced RT distributions for correct responses and error rates. Moreover, temporal spike sequences obtained from the model network closely resembled to the neural activity in the monkey prefrontal or parietal area during other visual decision tasks such as motion discrimination and oddball detection tasks.     

Biased competition through variations in amplitude of <Emphasis Type="Italic">γ</Emphasis>-oscillations

Experiments in visual cortex have shown that the firing rate of a neuron in response to the simultaneous presentation of a preferred and non-preferred stimulus within the receptive field is intermediate between that for the two stimuli alone (stimulus competition). Attention directed to one of the stimuli drives the response towards the response induced by the attended stimulus alone (selective attention). This study shows that a simple feedforward model with fixed synaptic conductance values can reproduce these two phenomena using synchronization in the gamma-frequency range to increase the effective synaptic gain for the responses to the attended stimulus. The performance of the model is robust to changes in the parameter values. The model predicts that the phase locking between presynaptic input and output spikes increases with attention.