Ingo Fründ

Human gamma-band activity: a review on cognitive and behavioral correlates and network models.

Gamma-band oscillations (roughly 30-100 Hz) in human and animal EEG have received considerable attention in the past due to their correlations with cognitive processes. Here, we want to sketch how some of the higher cognitive functions can be explained by memory processes which are known to modulate gamma activity. Especially, the function of binding together the multiple features of a perceived object requires a comparison with contents stored in memory. In addition, we review recent findings about the actual behavioral relevance of human gamma-band activity. Interestingly, rather simple models of spiking neurons are not only able to generate oscillatory activity within the gamma-band range, but even show modulations of these oscillations in line with findings from human experiments.

Inference for psychometric functions in the presence of nonstationary behavior

Measuring sensitivity is at the heart of psychophysics. Often, sensitivity is derived from estimates of the psychometric function. This function relates response probability to stimulus intensity. In estimating these response probabilities, most studies assume stationary observers: Responses are expected to be dependent only on the intensity of a presented stimulus and not on other factors such as stimulus sequence, duration of the experiment, or the responses on previous trials. Unfortunately, a number of factors such as learning, fatigue, or fluctuations in attention and motivation will typically result in violations of this assumption. The severity of these violations is yet unknown. We use Monte Carlo simulations to show that violations of these assumptions can result in underestimation of confidence intervals for parameters of the psychometric function. Even worse, collecting more trials does not eliminate this misestimation of confidence intervals. We present a simple adjustment of the confidence intervals that corrects for the underestimation almost independently of the number of trials and the particular type of violation.

PyMVPA: A Unifying Approach to the Analysis of Neuroscientific Data

The Python programming language is steadily increasing in popularity as the language of choice for scientific computing. The ability of this scripting environment to access a huge code base in various languages, combined with its syntactical simplicity, make it the ideal tool for implementing and sharing ideas among scientists from numerous fields and with heterogeneous methodological backgrounds. The recent rise of reciprocal interest between the machine learning (ML) and neuroscience communities is an example of the desire for an inter-disciplinary transfer of computational methods that can benefit from a Python-based framework. For many years, a large fraction of both research communities have addressed, almost independently, very high-dimensional problems with almost completely non-overlapping methods. However, a number of recently published studies that applied ML methods to neuroscience research questions attracted a lot of attention from researchers from both fields, as well as the general public, and showed that this approach can provide novel and fruitful insights into the functioning of the brain. In this article we show how PyMVPA, a specialized Python framework for machine learning based data analysis, can help to facilitate this inter-disciplinary technology transfer by providing a single interface to a wide array of machine learning libraries and neural data-processing methods. We demonstrate the general applicability and power of PyMVPA via analyses of a number of neural data modalities, including fMRI, EEG, MEG, and extracellular recordings.

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.

Inter- and Intra-Individual Covariations of Hemodynamic and Oscillatory Gamma Responses in the Human Cortex

The time course of local field potentials (LFPs) displaying typical discharge frequencies in the gamma frequency range highly correlates with the blood oxygen level dependent (BOLD) signal in response to rotating checkerboard stimuli in animals. In humans, oscillatory gamma-band responses (GBRs) show strong inter-individual variations in frequency and amplitude but considerable intra-individual reliability indicating that individual gamma activity reflects a personal trait. While the functional role of these GBRs is still debated, investigations combining electroencephalography–functional magnetic resonance imaging (EEG–fMRI) measurements provide a tool to obtain further insights into the underlying functional architecture of the human brain and will shed light onto the understanding of the dynamic relation between the BOLD signal and the properties of the electrical activity recorded on the scalp. We investigated the relation between the hemodynamic response and evoked gamma-band response (eGBR) to visual stimulation. We tested the hypothesis that the amplitude of human eGBRs and BOLD responses covary intra-individually as a function of stimulation as well as inter-individually as a function of gamma-trait. Seventeen participants performed visual discrimination tasks during separate EEG and fMRI recordings. Results revealed that visual stimuli that evoked high GBRs also elicited strong BOLD responses in the human V1/V2 complex. Furthermore, inter-individual variations of BOLD responses to visual stimuli in the bilateral primary (Area 17) and secondary (Area V5/MT) visual cortex and the right hippocampal formation were correlated with the individual gamma-trait of the subjects. The present study further supports the notion that neural oscillations in the gamma frequency range are involved in the cascade of neural processes that underlie the hemodynamic responses measured with fMRI.

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.

Impairments of Gestalt perception in the intact hemifield of hemianopic patients are reflected in gamma-band EEG activity

Gamma-band responses (GBRs) are associated with Gestalt perception processes. In the present EEG study, we investigated the effects of perceptual grouping on the visual GBR in the perimetrically intact visual field of patients with homonymous hemianopia and compared them to healthy participants. All observers were presented either random arrays of Gabor elements or arrays with an embedded circular arrangement. For the hemianopic patients, the circle was presented in their intact hemifield only. For controls, the hemifield for the circle presentation was counterbalanced across subjects. The participants were instructed to detect the circle by pressing a corresponding button. A wavelet transform based on Morlet wavelets was employed for the calculation of oscillatory GBRs. The early evoked GBR exhibited a larger amplitude and shorter latency for the healthy group compared to hemianopic patients and was associated with behavioral measures. The late total GBR between 200 and 400ms after stimulus onset was significantly increased for Gestalt-like patterns in healthy participants. This effect was not manifested in patients. The present findings indicate deficits in the early and late visual processing of Gestalt patterns even in the intact hemifield of hemianopic patients compared to healthy participants.

Early gamma-band responses reflect anticipatory top-down modulation in the auditory cortex.

For efficient and fast encoding of our complex acoustic environment, not only aspects of bottom-up processing are significant, but rather top-down influences such as attention, memory, and anticipation promote specific behavior and perception. Neural oscillatory activity in the gamma-range (30-80 Hz) is discussed as a conceivable candidate to represent very rapid modulations of top-down factors. We investigated effects of anticipation on early gamma-band responses (GBRs) of the EEG and event-related potentials (ERPs) in response to tone sequences. These sequences were composed of six sinusoidal tones, which could be either regularly ascending or descending in frequency. Thus, the sequences reflected a good continuation of pitch, which also resulted in the buildup of strong expectancies for the upcoming stimulus within the sequence. However, some of the tone sequences contained a violation of the good continuation of pitch at the third or fifth tone position. The early phase-locked portion of the gamma-band activity was significantly increased when tones were in line with the good continuation of sequences compared to deviant tones. Further, a pronounced early negative ERP response, starting at 150 ms, was elicited by deviant tones at the third and fifth position. Our results support the notion that gamma-band oscillations reflect perceptual grouping processes of concurrent sounds and anticipatory top-down modulation, which involves some of the first stages of auditory information processing.