Reduction of variability of EEG occipital,parietal, and central alpha rhythms by visual feedback stimulation

Two experiments, each with 24 normal right-handed adults, examined variability of the response of EEG alpha rhythms during repeated visual stimulations that were contingent on the occurrence of those rhythms. Within-trial variability of alpha durations and no-alpha (alpha blocking) durations were recorded from bipolar derivations along two bilateral posterior-anterior lines. Variability was significantly lower for: (1) the contingent EEG connected to the stimulus compared to the contralateral EEG, which was recorded simultaneously but was not connected to the stimulus; (2) occipitoparietal EEGs compared to parietocentral EEGs; (3) alpha durations compared to no-alpha durations. Differences in variability among the four EEG locations on the left or the right side were significant for contingent EEGs but not for contralateral nonconnected EEGs. The results were interpreted to be a demonstration that feedback EEG method can be applied to research on the functional topography of an EEG response to sensory stimuli in terms of the reduction of variability of the response that can be achieved with feedback.     

Feedback regulation of the alpha electroencephalogram activity through control of the internal and external parameters

Summary Experiments with feedback stimulation triggered from the subject's electroencephalogram result in changing the sequential time series of intervals of occipital alpha and intervals of little or no alpha EEG activity. The rate of recurrence of alpha and no-alpha EEG can be changed by regulating the external feedback stimuli or by asking the subject to change his internal state. Four different paradigms were investigated and the results interpreted in terms of the hypothesis that oculomotor functions regulate the occurrence and nonoccurrence of alpha.This work was supported by NIGMS Grants 5 PO1 GM 14940-04 and GM 15006-03.Most of this research was conducted at the Perception Laboratory, Veterans' Administration Hospital, Bedford, Massachusetts, with the cooperation of Dr. Thomas B. Mulholland.     

Effect of feedback contingencies on the control of occipital alpha

Ten color transparencies were presented 30 times each to ten normal adults in response to changes in their occipital-parietal EEG's. By means of a feedback paradigm, detected alphas caused each slide to flash and stay on as long as alpha persisted, and then to turn off when alpha attenuated, according to four different contingency conditions. For half of the feedback trials, stimulus presentation depended on alpha detection in only one hemisphere and was not influenced by changes in the simultaneously recorded contralateral EEG. For the other feedback trials, stimulus presentation was bilaterally contingent. These contingency configurations were compared with sham feedback, a noncontingent condition during which slide presentation was controlled by a prerecorded tape. For both alpha and no-alpha, the ratio of mean duration over standard error was used as a quantification of the EEG response to visual stimulation. It was assumed that larger ratios indicated increased control of the EEG. Compared with the sham condition, all feedback contingencies produced greater ratios, and hence, improved control of the EEG. The highest ratios were obtained during unilateral feedback from the EEG in which the occurrence of alpha elicited the visual stimulus. The results show that contingency between a visual stimulus and the EEG is an important parameter with regard to experimental control of the EEG.     

Attention and regulation of EEG alpha-attenuation responses

Two experiments with 16 normal adults of both sexes tested the hypothesis that inattention to a biofeedback display is associated with increased variability of those physiological processes that had been regulated by the biofeedback. Each experiment was a repeated-measures-on-independent-subjects-design. Dependent variables were the time durations and the mean rms power of two mutually exclusive segments of the parietal-occipital EEG: alpha and not-alpha segments. Independent variables were combination of counting tasks and instructions to look at, listen to, and count visual and auditory flashes and clicks. The durations of alpha and not-alpha segments were controlled or regulated by means of an alpha-contingent visual feedback stimulus, Attention to the feedback stimulus was challenged by instructions to count other, noncontingent stimuli. Control of alpha and not-alpha segments was least for conditions of (1) “sham” feedback, and (2) feedback with instructions to count noncontingent auditory clicks, which were presented 3/sec while the feedback visual stimuli were occurring. A new EEG test of attention and distraction was suggested.     

Auditory stimulation dishabituates anti-predator escape behavior in hermit crabs (Coenobita clypeatus)

Responses to innocuous stimuli often habituate with repeated stimulation, but the mechanisms involved in dishabituation are less well studied. Chan et al. (2010b) found that hermit crabs were quicker to perform an anti-predator withdrawal response in the presence of a short-duration white noise relative to a longer noise stimulus. In two experiments, we examined whether this effect could be explicable in terms of a non-associative learning process. We delivered repeated presentations of a simulated visual predator to hermit crabs, which initially caused the crabs to withdraw into their shells. After a number of trials, the visual stimulus lost the ability to elicit the withdrawal response. We then presented the crabs with an auditory stimulus prior to an additional presentation of the visual predator. In Experiment 1, the presentation of a 10-s, 89-dB SPL noise produced no significant dishabituation of the response. In Experiment 2 we increased the duration (50 s) and intensity (95 dB) of the noise, and found that the crabs recovered their withdrawal response to the visual predator. This finding illustrates dishabituation of an antipredator response and suggests two distinct processes—distraction and sensitization—are influenced by the same stimulus parameters, and interact to modulate the strength of the anti-predator response.     

Response dynamics of bullfrog ON-OFF RGCs to different stimulus durations

Stimulus duration is an important feature of visual stimulation. In the present study, response properties of bullfrog ON-OFF retinal ganglion cells (RGCs) in exposure to different visual stimulus durations were studied. By using a multi-electrode recording system, spike discharges from ON-OFF RGCs were simultaneously recorded, and the cells’ ON and OFF responses were analyzed. It was found that the ON response characteristics, including response latency, spike count, as well as correlated activity and relative latency between pair-wise cells, were modulated by different light OFF intervals, while the OFF response characteristics were modulated by different light ON durations. Stimulus information carried by the ON and OFF responses was then analyzed, and it was found that information about different light ON durations was more carried by transient OFF response, whereas information about different light OFF intervals were more carried by transient ON response. Meanwhile, more than 80 % information about stimulus durations was carried by firing rate. These results suggest that ON-OFF RGCs are sensitive to different stimulus durations, and they can efficiently encode the information about visual stimulus duration by firing rate.     

A feedforward model of suppressive and facilitatory habituation effects

 Simple exposure to repeatitive stimulation is known to induce short-term learning effects across a wide range of species. These effects can be both suppressive and facilitatory depending on stimulus conditions: repeatitive presentation of a weak stimulus decreases the strength of the response (habituation), whereas presentation of a tonic stimulus following a series of weak stimuli transiently increases the response strength (dishabituation). Although these phenomena have been comprehensively characterized at both behavioral and cellular levels, most existing models of nonassociative learning focus exclusively on the suppressive or facilitatory changes in response, and do not attempt to relate cellular events to behavior. I propose here a feedforward model of habituation effects that explains both suppressive and facilitatory changes in response relying on the interaction between excitatory and inhibitory processes that develop in parallel on two different timescales. The model's properties are used to explain the rate sensitivity property of habituation and recovery and stimulus dishabituation. Received: 1 June 2001 / Accepted in revised form: 4 December 2001     

Functional aspects of evoked alpha and theta responses in humans and cats

A systems theoretical approach was used to compare possible functional roles of theta (4--7 Hz) and alpha (8--15 Hz) response components of brain evoked potentials. These response components were described earlier by Ba\c sar (1980). We recorded EEG and evoked potentials (EPs) from occipital scalp locations in 11 subjects. We used auditory and visual stimuli as inadequate and adequate stimuli, respectively (``cross-modality' measurements). The combined EEG-EP epochs were analysed in frequency domain with fast Fourier transform and adaptive digital filters. Alpha (8--15 Hz) response components turned out to be dependent on whether the stimulus was adequate or not (median amplitude with inadequate vs. adequate stimulation: vs. ). Theta (4--7 Hz) response components were less dependent on stimulus modality (inadequate vs. adequate stimulation: vs. ). In EP recordings the occipital alpha response almost disappeared in the first 250 ms following auditory stimulation. Comparable behaviour was observed in similar experiments with recordings from the cat visual cortex (area 17) and with occipital magnetoencephalographic recordings. Taking into account the above-mentioned previous reports on intracranial recordings in primary sensory areas of the cat brain and preliminary results of magnetoencephalographic measurements, we propose the following hypothesis: alpha responses in a time window of about 250 ms after stimulation might predominantly reflect primary sensory processing whereas the theta responses in the first 250 ms after stimulation might be more involved in supra-modality -- or cross-modality -- associative-cognitive processing. Received: 25 February 1994 / Accepted in revised form: 5 August 1994     

Feedback delay and amplitude threshold and control of the occipital EEG

Six normal adults looked at 12 color transparencies that were presented 30 times each in response to the occurrence of alpha in the occipital EEG and that were not presented when no-alpha occurred. The time delay between the occurrence of recorded EEG alpha and the feedback stimulus was systematically varied, as was the threshold value of alpha amplitude required for feedback stimulus to occur. As time delay increased, the replicative reliability of the temporal association between alpha and the visual stimulus decreased. The measures of the EEG response were: the mean durations of alpha and of no-alpha durations ; an estimate of random scatter or variability in a single trial (S _e ); and the ratio of these , which is the reciprocal of the coefficient of variation. The latter ratio defines control of the EEG response. Control was greatest for the minimum feedback delay (.25 sec) and a feedback threshold of 25% of the maximum peak alpha amplitude recorded under a prior eyes-closed. resting condition. Control of the EEG decreased as time delays increased and at lower and higher feedback thresholds. It was proposed that the measurement of control for various combinations of values of relevant experimental parameters could be used to select that combination which gave the best control. This optimum combination of values of parameters could be chosen for astandardized method.This research was supported in its entirety by the Veterans Administration Medical Research Program.     

How does the toad's visual system discriminate different worm-like stimuli?

Behavioral experiments show that toads exhibit stimulus- and locus-specific habituation. Different worm-like stimuli that toads can discriminate at a certain visual location form a dishabituation hierarchy. What is the neural mechanism which underlies these behaviors? This paper proposes that the toad discriminates visual objects based on temporal responses, and that discrimination is reflected in different average neuronal firing rates at some higher visual center, hypothetically anterior thalamus. This theory is developed through a large-scale neural simulation which includes retina, tectum and anterior thalamus. The neural model based on this theory predicts that retinal R2 cells play a primary role in the discrimination via tectal small pear cells (SP) and R3 cells refine the feature analysis by inhibition. The simulation demonstrates that the retinal response to the trailing edge of a stimulus is as crucial for pattern discrimination as the response to the leading edge. The new dishabituation hierarchies predicted by this model by reversing contrast and shrinking stimulus size need to be tested experimentally.     

On the interactivity in the human visual cortex caused by specific and nonspecific inflows

Summary An interactivity (Sato, 1969) with respect not only to the interstimulus intervenal between the conditioned and test stimuli but also with respect to the time after the test stimulus was defined as a new generalized recovery function.The contour maps of the above interactivities of the cerebral evoked potentials elicited by the paired flash stimuli and by the conditioned cutaneous and test flash stimuli, respectively, were obtained in the adult human visual cortex.It was suggested to a certain degree by the autocorrelograms of background occipital EEGs and average VEPs that the more prominent VEPs are elicited, the more predominant are the alpha waves in the occipital background EEG, and vice versa. In the contour maps of the interactivity by the paired flash stimuli, cyclic changes of facilitory mountains and occlusive valleys were regular and conspicuous in the subject who exhibited predominant alpha waves in his background EEG, whereas the cyclic changes were less regular and less marked in the subject with few alpha waves in the background EEG.The cyclic changes in the former subject appeared at an interval of about 100 msec at an interstimulus interval of around 100 msec. This acts favorably in the contribution of the cyclic change in interactivity via natural stimulations to the generation of alpha waves in spontaneous EEG. The main facilitory mountains and occlusive valleys in the maps were arranged in a line parallel to the ordinate (axis of the interstimulus interval), which suggest that the conditioned flash will affect in a specific fashion each response compoment elicited via specific afferent inflows in the VEP timelocked to the test flash stimulus. On the contrary, those in the maps obtained by the conditioned cutaneous and test flash stimuli were diagonal to the abscissa (time after the test stimulus) and ordinate, and/or the peaks of facilitory mountains and bottoms of occlusive valleys tended to form a line parallel to the abscissa, which indicates that as the cutaneous somatosensory afferent inflow converges to the visual cortex nonspecific interactivities are exhibited in each component in the VEP.     

Alpha ERD and human visual selective attention

We compared the alpha band EEG depression (event-related desynchnization, ERD) level in two tasks, involving activation of different attentional processes: visual search for a deviant relevant stimulus among many similar ones and visual oddball. Control data for the visual search task consisted of simple viewing of several stimuli being of the same shape as the relevant stimulus in the search trials. Gaze position was verified by eye tracking method. We interpreted alpha band ERD as a correlate of activation of attentional processes. Fixating the target in visual search task caused a significantly larger ERD than fixating the same stimuli in control trials over all leads. We suppose this to be related with task and visual environment complexities. The frontal ERD domination may indicate attentional control over voluntary movements execution (top-down attention). The caudal ERD may be related with updating of visual information as a result of search process (bottom-up attention). Both relevant and irrelevant stimuli in the oddball task also induced alpha band ERD, but it was larger in response to relevant one and reached maximum level over occipital leads. Domination of caudal ERD in oddball task is supposed to indicate bottom-up attention processes.     

Perceptual echoes at 10 Hz in the human brain

The occipital alpha rhythm (～10 Hz) is the most prominent electrophysiological activity in the awake human brain, yet its functional role and relation to visual perception are little understood. Transient stimuli normally elicit a short series of positive and negative deflections lasting between 300 and 500 ms: the visual-evoked potential (VEP). Alpha oscillations, on the other hand, are generally suppressed by transient visual input; they only augment in response to periodic ("steady-state") inputs around 10 Hz. Here, we applied reverse-correlation techniques to the visual presentation of random, nonperiodic dynamic stimulation sequences and found that the brain response to each stimulus transient was not merely a short-lived VEP but also included a strong ～10 Hz oscillation that lasted for more than 1 s. In other words, the alpha rhythm implements an "echo" or reverberation of the input sequence. These echoes are correlated in magnitude and frequency with the observer's occipital alpha rhythm, are enhanced by visual attention, and can be rendered perceptually apparent in the form of ～10 Hz flicker. These findings suggest a role for the alpha rhythm in the maintenance of sensory representations over time.     

Topography of alpha and theta oscillatory responses upon auditory and visual stimuli in humans

Brain resonance phenomena and induced rhythms in the brain recently gained importance in electroencephalographic, magnetoencephalographic and cellular studies (Ba\c sar and Bullock 1992). It was hypothesized that evoked potentials are superpositions of induced rhythms caused by resonance phenomena in neural populations (Ba\c sar et al. 1992). According to Ba\c sar (1972), such resonance phenomena are reflected in the main peaks of the amplitude frequency characteristics computed from EEG responses. The present study is based on a frequency domain approach for the evaluation of topography- and modality-dependent properties of oscillatory brain responses. EEG and evoked potentials were recorded from vertex, parietal and occipital scalp locations in 24 volunteers. Two combined methods were applied: (1) amplitude frequency characteristics were computed from the transient evoked responses, and (2) frequency components of the transient responses were obtained by adaptive digital filtering. Our main goal was to investigate theta (4--7 Hz) and alpha (8--15 Hz) response components. (1) Amplitude frequency characteristics. Auditory stimuli elicited theta-alpha compound responses in the 4--11 Hz frequency band (e.g. typical peaking frequency around 7 Hz for vertex recordings). Visual stimuli elicited alpha responses (e.g. typical peaking frequency for vertex recordings around 9--12 Hz). Frequency maxima for visual stimuli thus had main peaks at higher frequency values than frequency maxima for auditory stimuli. (2) Digital filtering confirmed these results: for vertex recordings, theta vs. alpha response amplitudes were 9 vs 6 for auditory stimuli and 5 vs 5 for visual stimuli, thus confirming a shift towards higher frequencies, i.e. a more prominent contribution of the alpha range, in the case of visual stimulation. We hypothesize that these properties might reflect site- and modality-specific features of stimulus encoding in the brain in which resonance properties of neuron populations are involved. Furthermore we emphasize the utility of the systems theory approach for a better understanding of brain function by means of EPs. Received: 25 February 1994 / Accepted in revised form: 5 August 1994     

Configurational pattern discrimination responsible for dishabituation in common toads Bufo bufo (L.): Behavioral tests of the predictions of a neural model

Summary Recently, a neural model of visual pattern discrimination for stimulus-specific habituation was developed, based on previous behavioral studies which demonstrated that toads exhibit a dishabituation hierarchy for different worm-like stimuli. The model suggests that visual objects are represented by temporal coding and predicts that the dishabituation hierarchy changes when the stimulus/background contrast direction is reversed or the stimulus size is varied. The behavioral experiments reported in this paper were designed to test these predictions, (1) For a pair of stimuli from the contrast reversal prediction, the experimental results validated the theory. (2) For a pair of stimuli from the size reduction prediction, the experimental results failed to validate the theory. Further experiments concerning size effects suggest that configurai visual pattern discrimination in toads exhibits size invariance. (3) Inspired by the Groves-Thompson account of habituation, we found that dishabituation by a second stimulus has a separate process from habituation to a first stimulus. This paper serves as an example of a fruitful dialogue between experimentation and modeling, crucial for understanding brain functions.Abbreviations a-h worm-like stimulus patterns - AT anterior thalamus - ERF excitatory receptive field - IRF inhibitory receptive field - RF receptive field - R2 to R4 retinal ganglion cell types - vMP posterior ventromedial pallium     

Interaction of recovery cycles of evoked potentials in the cerebral cortex of the cat in response to stimulation of the superior colliculi and pulvinar

Similar character of recovery cycles of evoked potentials in the visual cortex to electric stimulation of the superior colliculi (SC) and pulvinar was found in chronic experiments on alert cats irrespective of stimuli presentation order. In the association cortex preceding SC stimulation facilitated the response to test stimulation of pulvinar almost at all delays between the stimuli. If the pulvinar stimulation was applied as a conditioned stimulus, then the response to SC stimulation under intervals of 20-200 ms was depressed. The obtained data point to equivalence of the inputs from SC and pulvinar to the visual cortex, to different informational value of inputs from SC to the association and visual cortex, and to mutual function dependence of the inputs from SC and pulvinar to the association cortex.     

Event-related synchronization and desynchronization of EEG during appraisal of threatening and pleasant visual stimuli in high anxious subjects

The 62-channel EEG was recorded while low (LA, n = 18) and high (HA, n = 18) trait-anxious subjects viewed sequentially presented neutral, threatening and pleasant IAPS stimuli. Event-related desynchronization (ERD) and synchronization (ERS) were studied in the delta, theta1, theta2, alpha1, alpha2, beta1, beta2, beta3, and gamma frequency bands. Between-group differences, related to stimulus emotionality, were linked to theta1 and theta2 bands. In the low theta at prefrontal sites in the test period of 100-700 ms after stimulus onset HA exhibited relative predominance of the left hemisphere in response to both threatening and pleasant stimuli, whereas LA yielded larger right than left hemisphere activity in response to all the three stimulus categories. In the upper theta band between group differences were associated with posterior cortical regions and the test period of 0-1000 ms after stimulus onset: HA exhibited the largest ERS to threatening, whereas LA prompted the largest ERS to pleasant stimuli. Finally, according to the ERD data, in the alpha1 band HA participants in comparison with LA revealed enhanced left hemisphere activation in response to all the stimulus categories. It is suggested that as it is indexed by theta-ERS relative predominance of the left hemisphere at prefrontal sites along with the largest bilateral activity of posterior cortical regions (i.e., enhanced higher order visual processing) to threatening stimuli could form the basis for general bias towards threatening information in HA at the very early stages of emotional processing.     

Opposite Distortions in Interval Timing Perception for Visual and Auditory Stimuli with Temporal Modulations

When an object is presented visually and moves or flickers, the perception of its duration tends to be overestimated. Such an overestimation is called time dilation. Perceived time can also be distorted when a stimulus is presented aurally as an auditory flutter, but the mechanisms and their relationship to visual processing remains unclear. In the present study, we measured interval timing perception while modulating the temporal characteristics of visual and auditory stimuli, and investigated whether the interval times of visually and aurally presented objects shared a common mechanism. In these experiments, participants compared the durations of flickering or fluttering stimuli to standard stimuli, which were presented continuously. Perceived durations for auditory flutters were underestimated, while perceived durations of visual flickers were overestimated. When auditory flutters and visual flickers were presented simultaneously, these distortion effects were cancelled out. When auditory flutters were presented with a constantly presented visual stimulus, the interval timing perception of the visual stimulus was affected by the auditory flutters. These results indicate that interval timing perception is governed by independent mechanisms for visual and auditory processing, and that there are some interactions between the two processing systems.     

Non-linear population firing rates and voltage sensitive dye signals in visual areas 17 and 18 to short duration stimuli

Visual stimuli of short duration seem to persist longer after the stimulus offset than stimuli of longer duration. This visual persistence must have a physiological explanation. In ferrets exposed to stimuli of different durations we measured the relative changes in the membrane potentials with a voltage sensitive dye and the action potentials of populations of neurons in the upper layers of areas 17 and 18. For durations less than 100 ms, the timing and amplitude of the firing and membrane potentials showed several non-linear effects. The ON response became truncated, the OFF response progressively reduced, and the timing of the OFF responses progressively delayed the shorter the stimulus duration. The offset of the stimulus elicited a sudden and strong negativity in the time derivative of the dye signal. All these non-linearities could be explained by the stimulus offset inducing a sudden inhibition in layers II-III as indicated by the strongly negative time derivative of the dye signal. Despite the non-linear behavior of the layer II-III neurons the sum of the action potentials, integrated from the peak of the ON response to the peak of the OFF response, was almost linearly related to the stimulus duration.