Daniel Lenz

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.

Memory-matches evoke human gamma-responses

BackgroundHuman brain activity in the gamma frequency range has been shown to be a correlate of numerous cognitive functions like attention, perception and memory access. More specifically, gamma activity has been found to be enhanced when stimuli are stored in or match with short-term memory (STM). We tested the hypothesis that gamma activity is also evoked when stimuli match representations in long-term-memory (LTM). EEG was recorded from 13 subjects performing a choice reaction task. Visual stimuli were either known real-world objects with a memory representation or novel configurations never seen before.ResultsAll stimuli evoked an early gamma response which was maximal over occipital electrodes. This evoked gamma activity was significantly larger for items that matched memory templates.ConclusionsTherefore, we argue that gamma activity results from the feedback from memory into perception systems. This assumption seems to be true for STM as well as LTM.

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.

Enhanced gamma-band activity in ADHD patients lacks correlation with memory performance found in healthy children

Previous electrophysiological as well as imaging research has contributed to the understanding of impairments in attention, executive functions, and memory in patients with attention-deficit/hyperactivity disorder (ADHD). However, there is a lack of studies investigating ADHD related differences in the gamma range of human electroencephalogram (EEG), although gamma activity is strongly associated with cognitive processes impaired in ADHD patients and is also modulated by dopamine polymorphisms linked with ADHD. To close this gap, the present study compared gamma activity in ADHD children with that of healthy controls and correlated it with memory performance. EEG was recorded from 13 ADHD patients as well as 13 healthy control subjects during the encoding phase of a visual memory paradigm. In a subsequent recognition test, participants had to judge pictures as being old or new. Analysis of evoked gamma-band responses (GBRs) during stimulus encoding revealed a strong task-related enhancement for ADHD patients in parieto-occipital areas. Interestingly, this augmentation was not associated with recognition performance, whereas healthy subjects exhibited a strong positive correlation between evoked gamma activity during stimulus encoding and subsequent recognition performance. We interpret this finding as evidence of enhanced excitation levels and unspecific activation of processing resources in ADHD patients. Furthermore, enhanced GBRs in ADHD could also indicate a decrease of neuronal signal-to-noise ratio, partially caused by the genetic variations within the dopaminergic pathway of ADHD patients. The involved genetic polymorphisms have been shown to modulate evoked GBRs, which therefore could be a possible marker of impaired neurotransmission in ADHD.

Human EEG very high frequency oscillations reflect the number of matches with a template in auditory short-term memory.

Auditory perception comprises bottom-up as well as top-down processes. While research in the past has revealed many neural correlates of bottom-up processes, less is known about top-down modulation. Memory processes have recently been associated with oscillations in the gamma-band of human EEG (30 Hz and above) which are enhanced when incoming information matches a stored memory template. Therefore, we investigated event-related potentials (ERPs) and gamma-band activity in 17 healthy participants in a Go/NoGo-task. They listened to four frequency-modulated (FM) sounds which varied regarding the frequency range traversed and the direction of frequency modulation. One sound was defined as target and required a button press. The results of ERPs (N1, P2, N2, and P3) were consistent with previous studies. Analysis of evoked gamma-band responses yielded no significant task-dependent modulation, but we observed a stimulus dependency, which was also present in a control experiment: The amplitude of evoked gamma responses showed an inverted U-shape as a function of stimulus frequency. Investigation of total gamma activity revealed functionally relevant responses at high frequencies (90 Hz to 250 Hz), which showed significant modulations by matches with STM: Complete matches led to the strongest enhancements (starting around 100 ms after stimulus onset) and partial matches resulted in intermediate ones. The results support the conclusion that very high frequency oscillations (VHFOs) are markers of active stimulus discrimination in STM matching processes and are attributable to higher cognitive functions.

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.

Altered evoked gamma-band responses reveal impaired early visual processing in ADHD children.

Neurophysiological studies yield contrary results whether attentional problems of patients with attention-deficit/hyperactivity disorder (ADHD) are related to early visual processing deficits or not. Evoked gamma-band responses (GBRs), being among the first cortical responses occurring as early as 90ms after visual stimulation in human EEG, have been assigned a pivotal role in early visual processing. In particular, they are involved in memory matching processes and are enhanced when known stimuli are processed. The current study examined whether evoked GBR patterns during early memory matching processes could be indicative of an early visual processing deficit in ADHD patients. EEG was recorded from 13 young ADHD patients as well as 13 age-matched healthy participants. Both groups performed a simple forced choice reaction task employing line drawings of either known real-world items with representations in long-term memory or physically similar unknown items without such representations. Evoked GBRs of ADHD patients did not differentiate between known and unknown items. However, in healthy children, evoked GBRs were enhanced when stimuli matched a representation stored in memory. This finding indicates disadvantages at early visual processing stages in ADHD patients: In contrast to healthy participants, ADHD children lack an early memory based classification, possibly resulting in an impaired ability to rapidly reallocate attentional resources to relevant stimuli. These findings suggest that impaired early automatic stimulus classification in ADHD patients could be involved in deficits of selective and sustained attention.

Altered evoked gamma-band responses as a neurophysiological marker of schizophrenia?

Evoked gamma-band responses (GBRs) were shown to be involved in different aspects of human cognition and behavior. They have been linked to the integration and processing of incoming information leading to an adequate behavioral outcome. Consequently, altered evoked GBRs have been associated with impaired cognitive and behavioral states present in a variety of psychiatric disorders. However, to the best of our knowledge, there are no reports directly comparing evoked GBRs of different clinical groups in the same experimental setting. Thus, the purpose of the present study was to shed light on the question, whether evoked GBRs, as a kind of a neurophysiological biomarker of pathological states, might serve for characterization and distinguishing of groups suffering from diverse psychiatric disorders. We measured EEG during a passive auditory oddball-paradigm. Participants were patients diagnosed with schizophrenia, mood disorder, and personality disorders as well as a fourth group consisting of healthy participants. Our results indicate that evoked GBRs significantly differed from healthy participants only in schizophrenic patients whereas no difference could be observed for the other clinical groups. Our findings support the notion that early evoked GBRs could be indeed a trait variable of schizophrenia and are not a general marker of pathological brain states.

Resonance phenomena in the human auditory cortex: individual resonance frequencies of the cerebral cortex determine electrophysiological responses.

The brain can be considered a dynamical system which is able to oscillate at multiple frequencies. To study the brain’s preferred oscillation frequencies, the resonance frequencies in the frequency response of the system can be assessed by stimulating the brain at various stimulation frequencies. Furthermore, the event-related potential (ERP) can be considered as the brain’s impulse response. For linear dynamical systems, the frequency response should be equivalent to the frequency transform of the impulse response. The present study test whether this fundamental relation is also true for the frequency transform of the ERP and the frequency response of the brain. Results show that the spectral characteristics of both impulse and frequency response in the gamma frequency range are significantly correlated. Thus, we speculate that the resonance frequencies determine the frequency spectrum of the impulse response. This, in turn, implies that both measures are determined by the same, individually specific, neuronal generator mechanisms.