Effective detection of coupling in short and noisy bivariate data.

In the study of complex systems, one of the primary concerns is the characterization and quantification of interdependencies between different subsystems. In real-life systems, the nature of dependencies or coupling can be nonlinear and asymmetric, rendering the classical linear methods unsuitable for this purpose. Furthermore, experimental signals are noisy and short, which pose additional constraints for the measurement of underlying coupling. We discuss an index based on nonlinear dynamical system theory to measure the degree of coupling which can be asymmetric. The usefulness of this index has been demonstrated by several examples including simulated and real-life signals. This index is found to effectively disclose the nature and the degree of interactions even when the coupling is very weak and data are noisy and of limited length; by this way, new insight into the functioning of the underlying complex system is possible.

Phase-coupling of theta-gamma EEG rhythms during short-term memory processing.

Because of the importance of oscillations as a general phenomenon of neuronal activity the use of EEG spectral analysis is among the most important approaches for studying human information processing. Usually, oscillations at different frequencies occur simultaneously during information processing. Thus, the question for synchronisation of different frequencies by phase coupling and its possible functional significance is of primary importance. An answer may be given by bispectral analysis. Estimation of the (cross-) bispectrum allows to identify synchronised frequencies and possibly, the existence of non-linear phase coupling of different oscillators. Previous studies have demonstrated the simultaneous occurrence of slow (4-7 Hz) and fast (20-30 Hz) oscillations at frontal and prefrontal electrode positions during memory processing. However, interrelations between these rhythms have not been investigated up to now. In order to test short-term memory, the Sternberg task with random figures and number words was carried out on 10 female subjects. During the task EEG was recorded. Power and bispectral analyses from frontal, prefrontal and frontopolar regions were performed off-line. Increased power was found in both the theta and the gamma bands. Strong phase-coupling between theta at Fz and gamma at F3 and at Fp1, respectively, was shown for memorising number words by means of cross-bicoherence. A possible reason for this is an amplitude modulation of gamma frequencies by slow oscillations. The correspondent coherence analysis between the envelope of gamma frequencies at Fp1 and the raw EEG at Fz supports this presumption. This finding is interpreted as an EEG aspect of the functional linking between the prefrontal areas and the G.cinguli (as part of the limbic system), which are both extremely important for memory functions.

Universality in the brain while listening to music.

The human brain, which is one of the most complex organic systems, involves billions of interacting physiological and chemical processes that give rise to experimentally observed neuroelectrical activity, which is called an electroencephalogram (EEG). The presence of non-stationarity and intermittency render standard available methods unsuitable for detecting hidden dynamical patterns in the EEG. In this paper, a method that is suitable for non-stationary signals and preserving the phase characteristics and that combines wavelet and Hilbert transforms was applied to multivariate EEG signals from human subjects at rest as well as in different cognitive states: listening to music, listening to text and performing spatial imagination. It was found that, if suitably rescaled, the gamma band EEG over distributed brain areas while listening to music can be described by a universal and homogeneous scaling, whereas this homogeneity in scale is reduced at resting conditions and also during listening to text and performing spatial imagination. The degree of universality is characterized by a Kullback-Leibler divergence measure. By statistical surrogate analysis, nonlinear phase interaction was found to play an important role in exhibiting universality among multiple cortical regions.

EEG gamma-band phase synchronization between posterior and frontal cortex during mental rotation in humans.

The main purpose of the present paper was: (1) to study the phase synchronization pattern in the gamma-band while performing the classical Shepard-Metzler task of mental rotation; (2) to investigate the role of musical training; and (3) to study hemispheric differences in the degree of synchronization during mental rotation. Multivariate electroencephalograph signals from 20 male subjects (ten musicians and ten non-musicians) were recorded while performing the mental rotation task and also at resting condition. Phase synchronization was measured by a recent index, mean phase coherence. It was found that synchronization between frontal cortex and right parietal cortex was significantly increased during mental rotation with respect to rest, whereby musicians showed significantly higher degrees of synchronization than non-musicians. Left hemispheric dominance in the degree of phase synchronization, stronger in the posterior right parietal and occipital regions, was observed in musicians. Right hemispheric dominance was generally observed in non-musicians.

Long-range synchrony in the gamma band: role in music perception.

Synchronization seems to be a central mechanism for neuronal information processing within and between multiple brain areas. Furthermore, synchronization in the gamma band has been shown to play an important role in higher cognitive functions, especially by binding the necessary spatial and temporal information in different cortical areas to build a coherent perception. Specific task-induced (evoked) gamma oscillations have often been taken as an indication of synchrony, but the presence of long-range synchrony cannot be inferred from spectral power in the gamma range. We studied the usefulness of a relatively new measure, called similarity index to detect asymmetric interdependency between two brain regions. Spontaneous EEG from two groups-musicians and non-musicians-were recorded during several states: listening to music, listening to text, and at rest (eyes closed and eyes open). While listening to music, degrees of the gamma band synchrony over distributed cortical areas were found to be significantly higher in musicians than non-musicians. Yet no differences between these two groups were found at resting conditions and while listening to a neutral text. In contrast to the degree of long-range synchrony, spectral power in the gamma band was higher in non-musicians. The degree of spatial synchrony, a measure of signal complexity based on eigen-decomposition method, was also significantly increased in musicians while listening to music. As compared with non-musicians, the finding of increased long-range synchrony in musicians independent of spectral power is interpreted as a manifestation of a more advanced musical memory of musicians in binding together several features of the intrinsic complexity of music in a dynamical way.

Enhanced phase synchrony in the electroencephalograph gamma band for musicians while listening to music.

Multichannel electroencephalograph signals from two broad groups, 10 musicians and 10 nonmusicians, recorded in different states (in resting states or no task condition, with eyes opened and eyes closed, and with two musical tasks, listening to two different pieces of music) were studied. Degrees of phase synchrony in various frequency bands were assessed. No differences in the degree of synchronization in any frequency band were found between the two groups in resting conditions. Yet, while listening to music, significant increases of synchronization were found only in the gamma-frequency range (>30 Hz) over large cortical areas for the group of musicians. This high degree of synchronization elicited by music in the group of musicians might be due to their ability to host long-term memory representations of music and mediate access to these stored representations.

Musicians and the gamma band: a secret affair?

While listening to music, a significant high degree of phase synchrony in the gamma frequency range globally distributed over the brain was found in subjects with musical training (musicians) compared with subjects with no such training (non-musicians). No significant differences were found in other EEG frequency bands. Listening to neutral text did not produce any significant differences in the degree of synchronization between these two groups. For musicians, left-hemispheric dominance was found during listening to music. The right hemisphere was found to be dominant for non-musicians in text listening. The high degree of synchronization in musicians could be due to their high ability to retrieve musical patterns from their acoustic memory, which is a cogent condition for both listening to and anticipating musical sounds.

Interdependencies in the spontaneous EEG while listening to music.

We studied the patterns of interdependency between different brain regions during the performance of higher cognitive functions. Our goal was to check the existence in these patterns of both task-related differences (e.g. listening to music vs. rest) and training-related differences (musicians vs. non-musicians). For this purpose, a non-linear measure, called similarity index (S.I.), was used to detect asymmetric interdependencies between different brain regions by means of EEG signals. Relatively active and passive regions of the brain were found where the degree of activity was represented by excited degrees of freedom. The S.I. obtained during listening to different kinds of music was compared statistically with the S.I. with eyes closed, and significant changes (P< or = 0.05) were entered into schematic brain maps. A topographical representation of the S.I. yielded differences in the interdependency while performing different cognitive tasks. The results demonstrate the occurrence of task-related differences in both groups of subjects. Furthermore, subjects with musical training possessed significantly higher degrees of interdependencies than such without musical training while listening to music but not to text. We conclude that the new measure can be successfully applied for studying the dynamical co-operation between cortical areas during higher cognitive functioning.

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.

[Contribution of spontaneous EEG to understanding cognitive functions]. / Der Beitrag des Spontan-EEGs zum Verständnis kognitiver Funktionen.

Probability mapping of amplitude and coherence based on spectral analysis of the on-going EEG turned out to yield information on the cooperation of brain areas during cognitive processes. The method is based on comparing these two parameters during the performance of a mental task with the respective values of the EEG at rest before and after the task. For each of six frequency bands, spanning and entire EEG spectrum, the signs of significant changes of these parameters at the recording electrodes (19) and between all recording sites (19 x 18) are entered into schematic maps, representing the brain as seen from both sides and from the top. The results have proved to be largely task-specific for mental processes in both groups and individual studies. They are considered as electrical manifestations of differential attention. Since they may encompass EEG periods of even several minutes, these maps allow observations of average EEG characteristics even in complex, long-lasting mental accomplishments. The usefulness of the method is illustrated by four examples: doing mental arithmetic, listening to and composing music, simultaneous interpretation and contemplating and memorizing pictures.

Persistent patterns of brain activity: an EEG coherence study of the positive effect of music on spatial-temporal reasoning.

Motivated by predictions from the structured trion model of the cortex, behavioral experiments have demonstrated a causal short-term enhancement of spatial-temporal reasoning in college students following exposure to a Mozart sonata, but not in control conditions. The coherence analysis of electroencephalogram (EEG) recordings is well suited to the neurophysiological investigation of this behavioral enhancement. Here we report the presence of right frontal and left temporo-parietal coherent activity induced by listening to Mozart which carried over into the spatial-temporal tasks in three of our seven subjects. This carry-over effect was compared to EEG coherence analysis of spatial-temporal-tasks after listening to text. We suggest that these EEG coherence results provide the beginnings of understanding of the neurophysiological basis of the causal enhancement of spatial-temporal reasoning by listening to specific music. The observed long-lasting coherent EEG pattern might be evidence for structured sequences in cortical dynamics which extend over minutes.

The reflection of cognitive tasks in EEG and MRI and a method of its visualization.

Up until recently, neurology was dominated by localisatory thinking. Language and other so-called "centers" were considered to be centers of command controlling the respective functions. Today, there is general agreement that, instead, for every brain function numerous brain regions must act together. For the exploration of these manifold topographic cooperations produced by cognitive tasks, coherence of long-term EEG periods proved to be a proficient parameter for the representation of functionally essential connections. Because of the unequivocal meaningfulness of absolute coherence values, instead, only the signs of significant differences between coherence values during cognitive tasks and periods of EEG at rest before and after the task were considered for all possible electrode pairings and charted on schematic maps of the brain. In addition, the signs of significant changes of amplitude were entered. This procedure was performed for each of 6 frequency bands and for the 19 electrodes of the 10/20 system, thus yielding 171 possible plus or minus values for coherence and 19 for amplitude, respectively. The positions of the electrodes were marked by an MRI contrast medium. After the EEG, MRI examination was performed. The MRI data were segmented and the cortex was mapped onto a plane using a method similar to cartography. The exact electrode positions are registered from a similarly obtained map of the scalp and the electrode position pattern is used as basis for the coherence graphs. A detailed map of the cortex based on the segmented MRI data with the electrode positions marked is provided as a reference enabling allocation of the electrodes to the cortical structures. The usefulness of this procedure is demonstrated with a single subject by means of different cognitive tasks including musical thinking.

Approaches to verbal, visual and musical creativity by EEG coherence analysis.

Our approach to coherence analysis of the on-going EEG yields data on the cooperation between all possible electrode sites of the 10/20 system. This procedure compares epochs during mental activity with epochs of EEG at rest and takes into account only significant differences which are represented as lines between the respective electrodes on maps of the brain surface. The patterns thus obtained reflect the temporal average of the changes of the overall coherence pattern caused by any mental task. They are interpreted as reflecting differential attention required for the achievement of the mental task in question. Acts of creative thinking, be it verbally, visually or musically, are characterized by more coherence increases between occipital and frontopolar electrode sites than any other mental tasks. The results are interpreted by a stronger involvement of long cortico-cortical fibre systems in creative tasks.

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.

Sources of electric brain activity: intracortical current dipoles.

Intracortical evoked potentials after electrical stimulation of the optic nerve were simultaneously recorded from the different layers of the rabbit's visual cortex. The averaged potentials were subjected to current source density analyses. The results show that the first event after the stimulus is a current sink in layer VI according to the excitatory terminations of the specific afferent fibres of the geniculate body. Since the largest part of the specific afferents terminates in layers IV and V, the sink in those layers may be due to excitatory synaptic events there. The corresponding current sources are situated in the upper layers and are most probably passive sources due to the active sinks. This example demonstrates that the generation of intracortical electric potentials can be explained by vertically oriented current dipoles according to the anatomical peculiarities.

Brain electrical mechanisms of bilingual speech management: an initial investigation.

This exploratory study deals with EEG changes in 3 professional interpreters while mentally interpreting from their mother language into foreign languages and vice versa. EEGs were recorded while interpreting and compared with the periods at rest between these periods of interpreting. Significant (P < 0.05) changes of coherence between all pairs of electrodes with respect to the averaged EEG at rest were computed for 5 frequency bands between 4 and 32 Hz. The verbal tasks were control-compared with comparable coherence measures for mental arithmetic and listening to music. Interindividual differences predominated, but certain common characteristics of the EEG measures were also found. The temporal regions were most involved in interpreting and particularly in the uppermost beta band (24-32 Hz). More coherence increases--particularly in the right hemisphere--were found while interpreting into the foreign than into the native language. Coherence changes were found to accumulate in certain regions of the scalp as pivots or focal areas which apparently have functional significance for the task in question as nodal points of information exchange and/or transfer. Such pivots were found in T3 more than in T4 (in the right-handers) and vice versa in a left-hander. Theta and alpha bands behaved differently and did not show such clear-cut differences. The results during mental arithmetic and listening to music were different from the ones while interpreting. The results give support to the conception of the cortex as a network serving the greatest possible divergence and convergence of signals.

Thinking with images or thinking with language: a pilot EEG probability mapping study.

This pilot EEG mapping study was designed to explore thinking processes using complex mental imagery and thought processes. EEG was recorded with 19 electrodes (10/20 system against averaged ear lobe signals) while volunteers (n = 42) performed two separate tasks: visualization of an abstract concept and interpretation of a painting. Average spectral parameters such as amplitude, local and interhemispheric coherences were computed for five frequency bands (theta, alpha, beta 1, 2 and 3). Results indicate that the frontal regions are strongly involved during these tasks as evidenced by coherence changes. Changes are also present in temporal, parietal and occipital regions and are discussed in relation to information processing with the frontal regions considering the different cognitive functions required by the tasks.