Oscillatory Activities in Multiple Frequency Bands in Patients with Schizophrenia During Motion Perception.

Patients with schizophrenia show impairment in binding stimulus features into coherent objects, which are reflected in disturbed oscillatory activities. This study aimed to identify disturbances in multiple oscillatory bands during perceptual organization of motion perception in patients with schizophrenia. EEG was recorded from healthy controls and patients with schizophrenia during continuous presentation of a motion stimulus which induces reversals between two exogenously generated perceptions. This stimulus was used to investigate differences in motion binding processes between healthy controls and patients with schizophrenia. EEG signals were transformed into frequency components by means of the Morlet wavelet transformation in order to analyse inter-trial coherences (ITC) in the delta (1-4 Hz), theta (4-7 Hz), alpha (8-12 Hz), and gamma (28-48 Hz) frequency bands during exogenous motion binding. Patients showed decreased delta-ITC in occipital and theta-ITC in central and parietal areas, while no significant differences were found for neither alpha nor gamma-ITCs. The present study provides one of the first insights on the oscillatory synchronizations related with the motion perception in schizophrenia. The ITC differences revealed alterations in the consistency of large-scale integration and transfer functions in patients with schizophrenia.

What does the broken brain say to the neuroscientist? Oscillations and connectivity in schizophrenia, Alzheimer's disease, and bipolar disorder.

The application of the concept and methods of brain oscillations has been an important research area in neurosciences. In the last decades, besides the application in cognitive processes, the study of changes in brain oscillations in diseases has also become an important focal point of research. In the present paper, some remarkable examples in three different diseases are taken into consideration: 1) schizophrenia (SZ), 2) Alzheimer's disease (AD), 3) bipolar disorders (BD). In the current literature, decreased oscillations in cortical recordings are observed in most of the pathologies. For example, decrease of gamma activity in SZ, decrease of delta activity in almost all diseases, as well as frequency shifts in alpha and the lower frequencies were recorded. However, there are also paradoxical cases in which an increase of oscillatory activities is observed. In BD, whereas alpha activity is greatly decreased, a huge increase of beta activity is observed. Or, in SZ, a paradoxical increase of gamma activity can be observed during cognitive loading. We also observed paradoxical changes in the analysis of connectivity. In AD, we find that alpha, delta, and theta coherences between distant parts of the cortex are greatly decreased, whereas in the gamma band, event-related coherences attain very high values. The comparison of the results and paradoxical changes in diseases may lead to important conclusions related to the web of oscillations and neurotransmitters. In turn, we could gain new insights to approach "brain function", in general.

New rather than old? For working memory tasks with abstract patterns the P3 and the single-trial delta response are larger for modified than identical probe stimuli.

Memory-guided decision making is dynamic and context-dependent, even though many studies describe an enhancement of the P3 for recognized items in memory tasks ("old-new effect"). This study utilized a delay-dependent working memory task during which decision making could be optimized by focusing attention on detected changes instead of recognized similarities. Mean P3 amplitude and delta activity were analyzed from participants who classified probe stimuli as identical or modified. The P3 amplitudes were larger for modified than for identical probes, even when the probe occurred 4,000 ms after the primary stimulus. Enhanced single-trial amplitude, trial-by-trial consistency, and frontoparietal phase coherence of delta activity contributed to the larger P3 for the modified probe. Thus, context-dependent attentional resource allocation supporting memory-guided decisions might explain the enhancement of the P3 for specific probe types.

Event-related delta and theta brain oscillations reflect age-related changes in both a general and a specific neuronal inhibitory mechanism.

OBJECTIVE: ERPs may be limited in validity when investigating inhibitory functions in later adulthood, as age-related increases in intraindividual variability and changes in EEG-oscillations are not considered. The present study compared averaged ERP peak and single trial time-frequency (TF) data analysis. METHODS: Go/NoGo ERP waves amplitude/latency measures were compared with a TF analysis estimating single trial event-related EEG spectral power enhancement and intertrial phase-locking (ITC) in delta and theta band. RESULTS: Age-related larger ITC was found for theta oscillations in the N2-P3 time range during NoGo, only. Discrepancies between N1/N2 ERP and TF results were obtained. Go/NoGo-P3 amplitude reductions in elderly were not related to an increased delta latency jitter. CONCLUSIONS: Discrepancies between ERPs and TF results challenge conclusions made about age-related changes in Go/NoGo-N2. Earlier reports of age-related changes in P3 are supported by the present results. The study implies age-related impairments in a general neuronal inhibition mechanism and a specific response inhibition mechanism. SIGNIFICANCE: The study indicates long-range communication impairments in the aged brain and the results are discussed considering hypotheses on increases in neural noise.

Altered oscillatory alpha and theta networks in schizophrenia.

In the present study we used a simple visual evoked potential and a visual oddball paradigm to investigate alterations in the temporal integration of different frequency components such as alpha and theta oscillations in patients with schizophrenia. We found that neither the amplitude enhancement after stimulus onset nor the intertrial phase coherence was generally reduced in patients, but that the topography of the neural response was altered. While healthy controls elicited their maximum early alpha as well as late theta response over posterior electrode sites, the maximum response in patients was shifted to anterior electrode positions. This result was not found for the late theta response for targets as target processing was accompanied with frontal theta amplitude enhancement in healthy controls as well. The change of the topographical response pattern was mirrored by the intertrial phase coherence in both frequency bands. The findings imply that schizophrenia is related to multiple alterations in oscillatory networks. Even during simple tasks without high cognitive demands dysfunctional mechanisms of temporal and regional coordination appear to be of importance in schizophrenia.

Event-related theta oscillations during working memory tasks in patients with schizophrenia and healthy controls.

Altered frontal lobe activity and executive control associated with working memory (WM) dysfunction are recognized as core deficits in schizophrenia. These impairments have been discussed as being associated with deficits in self-regulated action monitoring and anticipatory action plan generation. To study electrophysiological correlates of executive control -- specifically action monitoring and action rule switching -- under varying WM load, we used a paradigm derived from classic N-back (WM) tasks and requiring monitoring of simple actions. We focused on event-related changes in post-stimulus theta oscillatory activity during varying cognitive and WM demand in healthy controls and schizophrenia patients. The results show significant WM load and rule-switching-related increases of post-stimulus theta amplitude at fronto-central locations in controls. In patients with schizophrenia, there was no such modulation, but -- apart from an increased early theta at left temporal locations -- generally reduced late theta responses in all tasks and at all locations. Furthermore, the patients with schizophrenia showed significant differences in their error patterns, which imply differences in automation and anticipation of actions between controls and patients. These findings suggest that theta oscillations are involved in mediating frontal lobe activity and functions related to enhanced executive control. We conclude that the patients with schizophrenia showed deficits in acquiring a mental task set which appear to be associated with impairments in action monitoring and task-specific regulation of executive control.

Event-related oscillations are 'real brain responses'--wavelet analysis and new strategies.

The EEG consists of the activity of an ensemble of generators producing rhythmic activity in several frequency ranges. These oscillators are active usually in a random way. However, by application of sensory stimulation these generators are coupled and act together in a coherent way. This synchronization and enhancement of EEG activity gives rise to 'evoked' or 'event-related oscillations'. The compound evoked potential manifests as superimposition of evoked rhythms in the EEG frequencies ranging from delta to gamma ('natural frequencies of the brain'). The superimposition principle is described with efficient strategies and by utilization of an efficient algorithm. The wavelet analysis confirms the results of the combined analysis procedure obtained by using the amplitude frequency characteristics (AFCs) and digital filtering. The AFC and adapted digital filtering methods are based on the first approach to analyze average evoked potentials. In contrast, the wavelet analysis is based on signal retrieval and selection among a large number of sweeps recorded in a given physiological or psychological experiment. By combining all these results and concepts, it can be stated that the wavelet analysis underlines and extends the expression that alpha-, theta-, delta-, and gamma-responses described in this report are the most important brain responses related to psychophysiological functions. The wavelet analysis confirms once more the expression 'real signals' which we attribute to EEG frequency responses of the brain. It will be demonstrated that the delta, theta, and alpha responses (i.e. the rhythms 'predicted' by digital filtering) are real brain oscillations. The frequency components of the event-related potential vary independently of each other with respect to: (a) their relation to the event; (b) their topographic distribution; and (c) with the mode of the physiological measurements.

Selectively distributed gamma band system of the brain.

Experiments from the cat brain demonstrate that gamma responses can be recorded in cortex, hippocampus, cerebellum, formatio reticularis and thalamus, independent of the mode of sensory stimulation. These results support the hypothesis of a selectively distributed gamma system of the brain.

Gamma response of the brain: a multifunctional oscillation that represents bottom-up with top-down processing.

This paper deals with the functional correlates of the gamma response of the brain. A critical review of the literature findings reveals the existence of two types of gamma responses: an early gamma that fulfills sensory functions and a late gamma that fulfills perceptual-cognitive functions. However, even the early gamma shows individual differences. Such a finding points to the existence of top-down influences on sensory processes and to a parallel-processing model for brain function.

Event-related theta oscillations: an integrative and comparative approach in the human and animal brain.

This report provides a synthesis of results in both cat and human brains in order to point out the importance of theta responses during cognitive processes and P300 paradigms. The unique features of this report consisted of the fact that human and cat data during several cognitive paradigms were compared. The results open the way to formulate the selectively distributed theta system in the brain as analyzed by Basar, Schürmann and Sakowitz (this issue).

Topological distribution of oddball 'P300' responses.

This report describes the frequency response of the oddball paradigm upon auditory stimuli. Other reports related to wavelet analysis of the same ERPs (Demiralp et al., 1999) and the application of visual signals (Schürmann et al., this volume) indicate that the P300 response has a dominant delta response oscillation, independent of the modality of the stimulation. Moreover, the adaptive digital filtering and the wavelet analysis lead to very similar results, confirming that delta responses are real brain responses as already mentioned, by Basar et al. (this volume). The theta response has a second late response window in comparison to auditory evoked potentials. Moreover, the functional significance of the selectively distributed theta and delta systems of the brain will be clearly demonstrated. Signal detection, short-term memory, and decision-making processes are discussed.

Wavelet analysis of oddball P300.

The comparative wavelet analysis presented in details by Demiralp et al. (1999), Ademoglu (1995) and by Basar et al. (2001) will be now applied to oddball P300 results (see Basar-Eroglu et al., 2001). The results obtained basically confirm those obtained by using adaptive digital filtering: The delta response dominates the P300 potential while the theta response is prolonged in a second late window.

Delta responses and cognitive processing: single-trial evaluations of human visual P300.

Visual P300 responses were recorded by using checkerboard-type stimuli. (1) High amplitude P300-delta responses were visible even in single trial ERPs. (2) An algorithm for efficient selection of P300 single trials (based on evaluation of delta responses) is introduced. (3) The 'universal' character of the P300-delta response demonstrated in this report may open new avenues for the understanding of functional ERP components.

Gamma, alpha, delta, and theta oscillations govern cognitive processes.

The increased interest in gamma oscillations, now widely regarded as functionally relevant signals of the brain, underlines the importance of the concept of event-related oscillations for bridging the gap between single neurons and neural assemblies. Taking this concept further, we review experiments showing that oscillatory phenomena such as alpha, theta, and delta responses to events are, just as the gamma band, strongly interwoven with sensory and cognitive functions. This review argues that selectively distributed delta, theta, alpha and gamma oscillatory systems act as resonant communication networks through large populations of neurons. Thus, oscillatory processes might play a major role in functional communication in the brain in relation to memory and integrative functions.

Reversal-rate dependent differences in the EEG gamma-band during multistable visual perception.

It is an often reported observation in the literature on multistable perception that the reversal rate within a given observation time is subject to a high interindividual variability. Recently, we reported frontal gamma-band enhancement during multistable visual perception. The aim of the present study was to investigate whether changes in the gamma-band correspond to the variability of the reversal rate. Therefore, a total of 25 observers were divided into two subgroups according to their reversal rate during a 400-s observation period of a reversible pattern based on apparent motion. Subjects with more than 40 reversals within the 400-s were defined as high-rate switchers (HRS). Subjects with a reversal rate below 40 switches were defined as low-rate switchers (LRS). EEG was recorded from frontal, central, parietal, and occipital locations of both hemispheres. The results showed significantly higher gamma activity for the HRS in both phase-locked and non-phase-locked oscillations. Both subgroups showed the highest gamma amplitudes at frontal locations. The results support the involvement of attentional top-down processing in figure reversal. It is concluded that the higher gamma activity for the HRS reflects states of higher arousal, alertness and/or attention according to their fast reversal rate.

Electroencephalogram alpha (8-15 Hz) responses to visual stimuli in cat cortex, thalamus, and hippocampus: a distributed alpha network?

To investigate possible functional correlates of alpha (8-15 Hz) oscillations in the electroencephalogram (EEG) intracranial recordings in cats (from thalamus, occipital cortex, and hippocampus) were performed. In response to visual stimuli, event-related alpha oscillations were observed. Such alpha responses were found not only in a specific sensory (visual) pathway but also in the hippocampus, hinting at a possible distributed alpha response system.

Alpha activity decreases during the perception of Necker cube reversals: an application of wavelet transform.

Since the first observation of perceptual reversal by Necker, many theoretical approaches have been proposed. In a previous study, we showed that a positive wave appeared approximately 250 ms prior to the button press of the subjects, indicating perceptual reversal during the observation of the Necker cube figure. A basic difficulty in this type of study is the possible jitter in the latency of the button press due to the variability of the subjects' reaction time during a recording session. To overcome this difficulty, a pattern selection method based on the wavelet transform was proposed in the previous study. A dominant positive wavelet coefficient in the delta band was found to represent the perceptual-reversal-related positivity. In the present study, we aim to analyze the changes in the alpha frequency band during perceptual reversal by using the Necker cube. The RMS values of the alpha frequency band were measured for two time periods: +/- 3 SD around the mean peak latency of the perceptual-reversal-related positivity and a time window of the same length before the positive wave. We found significantly increased delta power and decreased alpha power during the perceptual-reversal-related positivity.

Brain oscillations in perception and memory.

Gamma oscillations, now widely regarded as functionally relevant signals of the brain, illustrate that the concept of event-related oscillations bridges the gap between single neurons and neural assemblies. Taking this concept further, we review experiments showing that oscillatory phenomena such as alpha, theta, or delta responses to events are strongly interwoven with sensory and cognitive functions. This review argues that selectively distributed delta, theta, alpha, and gamma oscillatory systems act as resonant communication networks through large populations of neurons. Thus, oscillatory processes might play a major role in relation with memory and integrative functions. A new 'neurons-brain' doctrine is also proposed to extend the neuron doctrine of Sherrington.

VEP-interhemispheric transfer time in 20-32 Hz band in man.

This research aimed to investigate callosal transfer in the different frequency bands of VEP to lateralized reversal of checkerboard pattern as stimuli. The chosen band pass filters (4-8 Hz, 8-15 Hz, 15-20 Hz, 20-32 Hz) were applied to the VEPs of subjects, and four different components for each VEP were obtained. Latency differences between hemispheres for digitally unfiltered and filtered VEPs were computed to estimate IHTT. Different IHTTs in the theta (17 ms), alpha (9 ms) and beta1 (7 ms) bands from right to left and from left to right were estimated, supporting the previous report. Furthermore, a transfer within 4.5 ms from right to left in the 20-32 Hz band for the occipital lobe was found. In conclusion, our findings suggest that the frequency analysis of VEP to lateralized stimuli give us additional information, relating to the types of callosal fibers.