Thalamocortical theta coherence in neurological patients at rest and during a working memory task.

We simultaneously recorded the local field potential (LFP) in the thalamus and the electroencephalogram (EEG) on the scalp of 5 patients suffering from neurogenic pain, epilepsy and movement disorders. In an earlier study [], we have investigated the slowing of EEG and the high thalamocortical coherence in the framework of thalamocortical dysrhythmia, the common underlying pathophysiology. The current study focuses on the effects of different cognitive conditions. When patients rested with eyes closed, a theta peak dominated the EEG spectra. The peak height was reduced upon opening the eyes, reminiscent of the classical alpha blocking. This peak reduction also appeared in the thalamic LFP recording. When patients activated their working memory by counting backwards, the theta peak increased in scalp EEG or in the LFP recorded in thalamic nuclei VA/VL. The coherence estimates between EEG and LFP ranged between 21% and 76% for different patients and cognitive conditions (mean: 50%). The involvement of both cortex and thalamus in working memory and the high thalamocortical coherence underline, in addition to cortico-cortical interactions, the importance of thalamocortical modules in the generation of higher cognitive functions.

Different frequencies for different scales of cortical integration: from local gamma to long range alpha/theta synchronization.

Cortical activity and perception are not driven by the external stimulus alone; rather sensory information has to be integrated with various other internal constraints such as expectations, recent memories, planned actions, etc. The question is how large scale integration over many remote and size-varying processes might be performed by the brain. We have conducted a series of EEG recordings during processes thought to involve neuronal assemblies of varying complexity. While local synchronization during visual processing evolved in the gamma frequency range, synchronization between neighboring temporal and parietal cortex during multimodal semantic processing evolved in a lower, the beta1 (12-18 Hz) frequency range, and long range fronto-parietal interactions during working memory retention and mental imagery evolved in the theta and alpha (4-8 Hz, 8-12 Hz) frequency range. Thus, a relationship seems to exist between the extent of functional integration and the synchronization-frequency. In particular, long-range interactions in the alpha and theta ranges seem specifically involved in processing of internal mental context, i.e. for top-down processing. We propose that large scale integration is performed by synchronization among neurons and neuronal assemblies evolving in different frequency ranges.

Top-down processing mediated by interareal synchronization.

Perception and cortical responses are not only driven "bottom-up" by the external stimulus but are altered by internal constraints such as expectancy or the current behavioral goal. To investigate neurophysiological mechanisms of such top-down effects, we analyzed the temporal interactions of neurons on different levels of the cortical hierarchy during perception of stimuli with varying behavioral significance. We found that interareal interactions in a middle-frequency range (theta and alpha; 4-12 Hz) strongly depend on the associated behavior, with a phase relationship and a layer specificity indicating a top-down-directed interaction. For novel unexpected stimuli, presumably processed in a feed-forward fashion, no such interactions occurred but high-frequency interactions (gamma; 20-100 Hz) were observed. Thus corticocortical synchronization reflects the internal state of the animal and may mediate top-down processes.

Bi-directional interactions between visual areas in the awake behaving cat.

The study of the cooperativity among cortical areas is essential to our understanding of brain functioning. Here we investigated the relative contributions of top-down and bottom-up directed interactions between area 17 and area 7 of the cat visual system. Bipolar local field potentials were recorded while the animals performed a go/no-go task or were in a quiet resting state. The data were analyzed by applying measures of interaction based on the Wiener-Granger causality concept. We found that during the visual task top-down directed interactions were of a similar magnitude as the bottom-up component. Second, interareal couplings tended to increase in conditions requiring a discriminative effort. Third, during behaviors not dominated by visual processing non-directed interactions increased.

Synchronization between temporal and parietal cortex during multimodal object processing in man.

A series of recordings in cat visual cortex suggest that synchronous activity in neuronal cell ensembles serves to bind the different perceptual qualities belonging to one object. We provide evidence that similar mechanisms seem also to be observable in human subjects for the representation of supramodal entities. Electroencephalogram (EEG) was recorded from 19 scalp electrodes (10/20 system) in 19 human subjects and EEG amplitude and coherence were determined during presentation of objects such as house, tree, ball. Objects were presented in three different ways: in a pictorial presentation, as spoken words and as written words. In order to find correlates of modality-independent processing, we searched for patterns of activation common to all three modalities of presentation. The common pattern turned out to be an increase of coherence between temporal and parietal electrodes in the 13-18 Hz beta1 frequency range. This is evidence that population activity of temporal cortex and parietal cortex shows enhanced coherence during presentation of semantic entities. Coherent activity in this low-frequency range might play a role for binding of multimodal ensembles.

Synchronization between prefrontal and posterior association cortex during human working memory.

We measured coherence between the electroencephalogram at different scalp sites while human subjects performed delayed response tasks. The tasks required the retention of either verbalizable strings of characters or abstract line drawings. In both types of tasks, a significant enhancement in coherence in the theta range (4-7 Hz) was found between prefrontal and posterior electrodes during 4-s retention intervals. During 6-s perception intervals, far fewer increases in theta coherence were found. Also in other frequency bands, coherence increased; however, the patterns of enhancement made a relevance for working memory processes seem unlikely. Our results suggest that working memory involves synchronization between prefrontal and posterior association cortex by phase-locked, low frequency (4-7 Hz) brain activity.

The possible meaning of the upper and lower alpha frequency ranges for cognitive and creative tasks.

This study is aimed at verifying the functional independence of two frequency bands within the alpha range. It is based on experiments that examined the role of these two hands with regard to the amount of local electrogenesis (amplitude) and the cooperation of brain areas (coherence) in mental tasks concerning: (1) visual perception and imagery; (2) listening to and composing music; (3) verbal and visual creativity; and (4) aspects of mood. In all experiments EEG were recorded for at least 1 min during each task, separated one from another by at rest periods of at least equal lengths. EEG electrodes were pasted according to the 10/20 system (averaged ear lobes as reference). After FFT power was calculated for all 19 electrodes, coherence was estimated for all possible electrode pairs (i.e. 171). This was done for six frequency ranges between 1.5 and 31.5 Hz, the alpha range having been divided into two (7.5-9 Hz and 9.5-12.5 Hz). The spectral parameters obtained during each task were compared with those of the merged EEG at rest, significant changes (P < or = 0.01-P < or = 0.05) were entered into schematic maps of the brain. Generally, fewer differences were found for amplitude than for coherence. In all four tasks concerning visual perception the clearest differences were found in single person studies. But also in group studies more or less distinct differences were found between alpha 1 and 2. Also in the series with music the two alpha bands did not behave uniformly, nor were uniform features found in the two series of musically trained and untrained subjects. Distinct discrepancies were also found in a verbal and visual imagery task. With respect to mood, only elevated mood was correlated with a decrease of coherence in alpha 2 and an increase of amplitude in alpha 1. This study though hinting at a different functional significance of these two alpha bands, however, does not allow to draw any conclusions as to their distinct functional meanings. Generally, the long-term coherence changes observed under these different mental tasks support the idea that part of information processing in the brain is reflected by the EEG. Structural peculiarities and microelectrode recordings of the cortex support this conclusion.

EEG aspects of mentally playing an instrument.

This pilot study examines the possibility to detect activities of SMA by means of EEG coherence analysis in a female professional violoncellist. The proband was asked, for 5 min each, to listen to a piece of music (she knew by heart), to imagine playing this piece and to imagine playing scales. The experiment was repeated after 5 days. Consistent significant coherence changes with respect to the averaged EEG at rest were plotted as probability maps. For each of these three tasks different coherence patterns emerged. Among the electrodes next to SMA, Fz was most involved while playing scales, less while imagining playing the same piece and still less while just listening to it.