Niko A. Busch

Size matters: effects of stimulus size, duration and eccentricity on the visual gamma-band response

OBJECTIVE The effects of stimulus size, duration and eccentricity on the visual gamma-band response (GBR) in human EEG were investigated and compared to visual evoked potentials (VEPs) in order to differentiate in future (and past) experiments whether changes in GBRs are due to stimulus-related (exogenous) or cognitive effects. METHODS EEG was recorded from 23 subjects while they performed a simple choice reaction time task requiring discrimination of squares and circles. In separate blocks stimulus size, duration, and eccentricity were manipulated. EEG was recorded from 64 electrodes. A wavelet transform based on Morlet wavelets was employed for the analysis of gamma-band activity. RESULTS Amplitude of the GBR was diminished for small and peripheral stimuli. With short stimulus durations ON and OFF responses of the GBR merged into one peak. In comparison, VEP amplitudes were less susceptible to stimulus features. In contrast to VEP latencies, however, GBR latency did not show a lateralization for eccentric stimuli. CONCLUSIONS In addition to previous experiments which have shown a modulation of the GBR by various cognitive processes, the present results demonstrate the susceptibility of the GBR in human EEG to exogenous factors, as numerous intracortical recordings in non-human primates have shown before. The results suggest that the human GBR resides in early visual areas. SIGNIFICANCE The demonstration of the susceptibility of the GBR to stimulus properties implies that studies aimed at exploring the involvement of the GBR in information processing have to be designed carefully. It also constrains the localization of the human GBR.

A cross-laboratory study of event-related gamma activity in a standard object recognition paradigm

This study proposes a standard paradigm for the investigation of visual information processing by means of gamma activity and presents a novel set of stimuli with a broad range of complex, coloured familiar real world and unfamiliar nonsense objects which are well matched with respect to physical stimulus properties. In order to demonstrate that the paradigm and stimulus set yield reliable results both were employed in two electrophysiological investigations in two independent laboratories. Participants were required to discriminate familiar from unfamiliar stimuli. The pattern of results was very consistent across laboratories. Early event-related potentials were not influenced by the stimulus type suggesting that physical stimulus properties did not confound object familiarity. Induced gamma band activity was stronger for familiar than for unfamiliar objects, supporting the notion of gamma activity as a signature of cortical networks underlying object representations.

Time-frequency analysis of target detection reveals an early interface between bottom-up and top-down processes in the gamma-band.

The early visual gamma-band response is an oscillatory signal evoked approximately 100 ms after stimulation. While some studies have found effects of various cognitive processes on this signal, such effects could not be replicated in other studies. Accordingly, some authors have claimed that evoked gamma-band activity reflects merely sensory functions. To resolve these conflicting positions, we conducted a target detection experiment in which the feature that defined the target could be distributed over a large or a small part of the entire stimulus. Only targets covering a larger area of the entire stimulus evoked stronger gamma-band activity than standards although the over-all stimulus size was identical for all stimuli. This increase in evoked activity resulted from stronger oscillatory power and not exclusively from stronger phase-locking. In contrast, N1 and P3 amplitudes were larger for target stimuli irrespective of the distribution of the relevant stimulus feature. These results are consistent with the notion that early gamma-band activity is generated by feature-selective neural assemblies the activity of which can in fact be modulated by top-down processes. This interaction, however, may be only detectable in scalp-recorded EEG if it affects a sufficient number of neural assemblies.

Electrophysiological evidence for different types of change detection and change blindness

Numerous studies have demonstrated that observers often fail to notice large changes in visual scenes, a phenomenon known as change blindness. Some experiments have suggested that phenomenological experience in change blindness experiments is more diverse than the common distinction between change detection and change blindness allows to resolve. Recently, it has been debated whether changes in visual scenes can be detected (“sensed”) without a corresponding perception of the changing object (“seeing”) and whether these phenomena build on fundamentally different perceptual processes. The present study investigated whether phenomenologically different perceptual processes such as sensing and seeing rely on different or similar neural processes. We studied ERP effects of visual change processing (as compared to change blindness) when observers merely detected the presence of a change (“sensing”) and when they identified the changing object in addition to detection (“seeing”). Although the visual awareness negativity (VAN)/selection negativity was similar for detection with and without identification, a change-related positivity and the N2pc contralateral to changes were found exclusively when the change was fully identified. This finding indicates that change identification requires perceptual and neural processes that are not involved in mere detection. In a second experiment, we demonstrated that the VAN and N2pc effects are similar to effects of selective attention in a visual search task. By contrast, the change-related positivity was specific for conscious processing of visual changes. The results suggest that changes can be detected (“sensed”) without perception of the changing object. Furthermore, sensing and seeing seem to rely on different neural processes and seem to constitute different types of visual perception. These findings bear implications for how different categories of visual awareness are related to different stages in visual processing.

From perception to action: phase-locked gamma oscillations correlate with reaction times in a speeded response task

BackgroundPhase-locked gamma oscillations have so far mainly been described in relation to perceptual processes such as sensation, attention or memory matching. Due to its very short latency (≈90 ms) such oscillations are a plausible candidate for very rapid integration of sensory and motor processes.ResultsWe measured EEG in 13 healthy participants in a speeded reaction task. Participants had to press a button as fast as possible whenever a visual stimulus was presented. The stimulus was always identical and did not have to be discriminated from other possible stimuli. In trials in which the participants showed a fast response, a slow negative potential over central electrodes starting approximately 800 ms before the response and highly phase-locked gamma oscillations over central and posterior electrodes between 90 and 140 ms after the stimulus were observed. In trials in which the participants showed a slow response, no slow negative potential was observed and phase-locked gamma oscillations were significantly reduced. Furthermore, for slow response trials the phase-locked gamma oscillations were significantly delayed with respect to fast response trials.ConclusionThese results indicate the relevance of phase-locked gamma oscillations for very fast (not necessarily detailed) integration processes.

EEG oscillations in the gamma and alpha range respond differently to spatial frequency

Physical properties of visual stimuli affect electrophysiological markers of perception. One important stimulus property is spatial frequency (SF). Therefore, we studied the influence of SF on human alpha (8-13 Hz) and gamma (>30 Hz) electroencephalographic (EEG) responses in a choice reaction task. Since real world images contain multiple SFs, an SF mixture was also examined. Event related potentials were modulated by SF around 80 and 300 ms. Evoked gamma responses were strongest for the low SF and the mixture stimulus; alpha responses were strongest for high SFs. The results link evoked and induced alpha and evoked gamma responses in human EEG to different modes of stimulus processing.

Circles are different: the perception of Glass patterns modulates early event-related potentials.

Glass patterns are randomized dot arrays that generate the perception of a global structure. They consist of correlated dot pairs which are generated by geometric transformations. The present study employed behavioral and event-related brain potential (ERP) measures to characterize the underlying neuronal processing when such patterns are perceived. Stimuli were circular, parallel, and randomized Glass patterns presented in two isoluminant colors using a choice reaction paradigm. Sixteen subjects were instructed to differentiate between colors with a button-press response. The N170 component increased in amplitude for circular patterns, and this effect was most pronounced bilaterally over occipito-temporal areas. The results suggest that the global perception of form generated by Glass patterns occurs at a stage of visual processing past area V1.

Time pressure modulates electrophysiological correlates of early visual processing

Background Reactions to sensory events sometimes require quick responses whereas at other times they require a high degree of accuracy–usually resulting in slower responses. It is important to understand whether visual processing under different response speed requirements employs different neural mechanisms. Methodology/Principal Findings We asked participants to classify visual patterns with different levels of detail as real-world or non-sense objects. In one condition, participants were to respond immediately, whereas in the other they responded after a delay of 1 second. As expected, participants performed more accurately in delayed response trials. This effect was pronounced for stimuli with a high level of detail. These behavioral effects were accompanied by modulations of stimulus related EEG gamma oscillations which are an electrophysiological correlate of early visual processing. In trials requiring speeded responses, early stimulus-locked oscillations discriminated real-world and non-sense objects irrespective of the level of detail. For stimuli with a higher level of detail, oscillatory power in a later time window discriminated real-world and non-sense objects irrespective of response speed requirements. Conclusions/Significance Thus, it seems plausible to assume that different response speed requirements trigger different dynamics of processing.

Anticipation of natural stimuli modulates EEG dynamics: physiology and simulation

In everyday life we often encounter situations in which we can expect a visual stimulus before we actually see it. Here, we study the impact of such stimulus anticipation on the actual response to a visual stimulus. Participants were to indicate the sex of deer and cattle on photographs of the respective animals. On some trials, participants were cued on the species of the upcoming animal whereas on other trials this was not the case. Time frequency analysis of the simultaneously recorded EEG revealed modulations by this cue stimulus in two time windows. Early