Walter Gruber

A shift of visual spatial attention is selectively associated with human EEG alpha activity.

Event‐related potentials and ongoing oscillatory electroencephalogram (EEG) activity were measured while subjects performed a cued visual spatial attention task. They were instructed to shift their attention to either the left or right visual hemifield according to a cue, which could be valid or invalid. Thereafter, a peripheral target had to be evaluated. At posterior parietal brain areas early components of the event‐related potential (P1 and N1) were higher when the cue had been valid compared with invalid. An anticipatory attention effect was found in EEG alpha magnitude at parieto‐occipital electrode sites. Starting 200 ms before target onset alpha amplitudes were significantly stronger suppressed at sites contralateral to the attended visual hemifield than ipsilateral to it. In addition, phase coupling between prefrontal and posterior parietal electrode sites was calculated. It was found that prefrontal cortex shows stronger phase coupling with posterior sites that are contralateral to the attended hemifield than ipsilateral sites. The results suggest that a shift of attention selectively modulates excitability of the contralateral posterior parietal cortex and that this posterior modulation of alpha activity is controlled by prefrontal regions.

Brain Oscillatory Substrates of Visual Short-Term Memory Capacity

The amount of information that can be stored in visual short-term memory is strictly limited to about four items. Therefore, memory capacity relies not only on the successful retention of relevant information but also on efficient suppression of distracting information, visual attention, and executive functions. However, completely separable neural signatures for these memory capacity-limiting factors remain to be identified. Because of its functional diversity, oscillatory brain activity may offer a utile solution. In the present study, we show that capacity-determining mechanisms, namely retention of relevant information and suppression of distracting information, are based on neural substrates independent of each other: the successful maintenance of relevant material in short-term memory is associated with cross-frequency phase synchronization between theta (rhythmical neural activity around 5 Hz) and gamma (> 50 Hz) oscillations at posterior parietal recording sites. On the other hand, electroencephalographic alpha activity (around 10 Hz) predicts memory capacity based on efficient suppression of irrelevant information in short-term memory. Moreover, repetitive transcranial magnetic stimulation at alpha frequency can modulate short-term memory capacity by influencing the ability to suppress distracting information. Taken together, the current study provides evidence for a double dissociation of brain oscillatory correlates of visual short-term memory capacity.

Are event-related potential components generated by phase resetting of brain oscillations? A critical discussion.

The event-related potential (ERP) is one of the most popular measures in human cognitive neuroscience. During the last few years there has been a debate about the neural fundamentals of ERPs. Two models have been proposed: The evoked model states that additive evoked responses which are completely independent of ongoing background electroencephalogram generate the ERP. On the other hand the phase reset model suggests a resetting of ongoing brain oscillations to be the neural generator of ERPs. Here, evidence for either of the two models is presented and validated, and their possible impact on cognitive neuroscience is discussed. In addition, future prospects on this field of research are presented.

Theta synchronization during episodic retrieval: neural correlates of conscious awareness.

The neural correlates of conscious awareness during successful memory retrieval were examined. In a recognition test, subjects indicated whether they consciously recalled the event in which a word was earlier presented (Remembering), or whether they recognized it on the basis that it was familiar in the absence of recollection (Knowing). An early EEG synchronization in the theta band predicted knowing, and a later remembering. Moreover, early and late event-related potentials were also found to predict knowing and remembering, respectively. The results indicate that the temporal dynamics of theta synchronization are related to the particular conscious experiences associated with memory retrieval.

A short review of slow phase synchronization and memory: evidence for control processes in different memory systems?

An integrative theoretical approach about memory related oscillations is presented. The basic assumptions are that memory related oscillations are probably confined to theta and upper alpha and that other frequencies particularly in the gamma range are important for memory primarily because they become coupled to lower frequencies and/or because they play a specific role for a high precision timing of neural events (including phenomena such as LTP or LTD). In contrast to previous studies, where we related theta and upper alpha to a variety of different memory processes, we suggest here that these oscillations are associated with top-down control processes in two large storage systems, working memory (WM) and long-term memory (LTM). These systems may have their own types of top-down processes that control access to and/or manipulation of stored information. The hypothesis, suggested here is that theta and upper alpha reflect these processes which can be best studied when analyzing phase.

Visual discrimination performance is related to decreased alpha amplitude but increased phase locking

This study investigated the question whether good and bad performance in a visual discrimination task is related to resting alpha power in a different way as it is known from memory tasks. The results show that good perceptual but not memory performance is related to low alpha amplitudes. In addition, we found that large phase resetting in the alpha band, and enhanced early components in the ERP are related to good performance in the discrimination task. The conclusion of this study is that in contrast to memory performance which is related to large resting alpha activity low alpha amplitudes are an indicator for good perceptual performance.

Cross-frequency phase synchronization: A brain mechanism of memory matching and attention

Spatial attention amplifies the neural response, i.e. spike rates, brain metabolism, and oscillatory activity at gamma frequency (beyond 30 Hz). In this study we show that when a visual target is attended enhanced synchrony between gamma phase (30 to 50 Hz) and theta phase (4 to 7 Hz), representing bottom-up and top-down activity, respectively, can be observed. This is interpreted as memory matching between incoming visual information and stored (top-down) information. The results highlight the function of oscillatory brain activity in the integration of memory and attention processes. This seems to be true in particular for theta oscillations showing increased interregional phase-coupling. We conclude that memory information is stored within a distributed theta network and it is matched with an incoming sensory trace at posterior brain areas.

Theta coupling in the human electroencephalogram during a working memory task.

The role of coupling between prefrontal and temporo-parietal brain areas within the theta frequency range of the human electroencephalogram was explored in a working memory task. During encoding of visual information higher theta amplitudes were observed in the right compared to the left hemisphere. Retrieval of visuospatial and verbal information elicited a more bilateral activation pattern. These effects were accompanied by theta coupling between dorsolateral prefrontal and right posterior temporal electrode sites during encoding. During retrieval prefrontal and bilateral temporo-parietal brain areas were coupled. These results support the idea of working memory functions being dependent on distributed prefrontal-temporal networks.

The interplay between theta and alpha oscillations in the human electroencephalogram reflects the transfer of information between memory systems

The exchange of information between the working and long-term memory system (WMS and LTMS) was investigated. We analyzed evoked theta and upper alpha desynchronization in a special memory task, designed to study the transfer of information between both memory systems. The results show that during attempts to retrieve information from the LTMS, evoked theta oscillations spread from anterior to posterior recording sites. When information actually is retrieved, the direction reverses and theta spreads to frontal sites. This time point--when direction reverses--varies between subjects to a large extent but is significantly correlated with memory performance and the onset of upper alpha desynchronization. We conclude that this phenomenon reflects the transfer of information between the WMS and LTMS

Theta oscillations and the ERP old/new effect: independent phenomena?

OBJECTIVES The hypothesis is examined whether a memory-related change in induced band power (oscillatory old/new effect) is functionally related to a memory-related increase in ERP positivity (ERP old/new effect). METHODS In order to avoid a confounding on the measurement level, induced band power (IBP) was used as a measure that is devoid of the influence of evoked components. The EEG was recorded during a recognition memory task. RESULTS The results show that compared to correctly rejected words, targets (remembered words) elicit a significantly larger P300. An oscillatory old/new effect was found for the delta and theta but not for the alpha band. It is manifested by an increase in delta and theta IBP which is significantly larger for targets than for correctly rejected words. It can be observed during the same time interval and shows the same topographic distribution as the ERP old/new effect. Most importantly, however, the ERP old/new effect (as well as the P300 itself) is generated by very slow frequencies which lie below the delta band. CONCLUSIONS These findings demonstrate that the two types of old/new effects are functionally related. Possible physiological mechanisms underlying this relationship are discussed in terms of a threshold change in the cortex (generating the P300) that occurs during an increase in hippocampal theta activity (generating an increase in induced theta power).