Event-related EEG/MEG synchronization and desynchronization: basic principles.

EEG coherency: I: statistics, reference electrode, volume conduction, Laplacians, cortical imaging, and interpretation at multiple scales

This study provides a time frame for the initial trajectory of activation flow along the dorsal and ventral visual processing streams and for the initial activation of prefrontal cortex in the human. We provide evidence that this widespread system of sensory, parietal, and prefrontal areas is activated in less than 30 ms, which is considerably shorter than typically assumed in the human event-related potential (ERP) literature and is consistent with recent intracranial data from macaques. We find a mean onset latency of activity over occipital cortex (C1(e)) at 56 ms, with dorsolateral frontal cortex subsequently active by just 80 ms. Given that activity in visual sensory areas typically continues for 100-400 ms prior to motor output, this rapid system-wide activation provides a time frame for the initiation of feedback processes onto sensory areas. There is clearly sufficient time for multiple iterations of interactive processing between sensory, parietal, and frontal areas during brief (e.g., 200 ms) periods of information processing preceding motor output. High-density electrical mapping also suggested activation in dorsal stream areas preceding ventral stream areas. Our data suggest that multiple visual generators are active in the latency range of the traditional C1 component of the ERP, which has often been taken to represent V1 activity alone. Based on the temporal pattern of activation shown in primate recordings and the evidence from these human recordings, we propose that only the initial portion of the C1 component (approximately the first 10-15 ms; C1(e)) is likely to represent a response that is predominated by V1 activity. These data strongly suggest that activity represented in the "early" ERP components such as P1 and N1 (and possibly even C1) is likely to reflect relatively late processing, after the initial volley of sensory afference through the visual system and involving top-down influences from parietal and frontal regions.

Flow of activation from V1 to frontal cortex in humans. A framework for defining "early" visual processing.

An internally or externally paced event results not only in the generation of an event-related potential (ERP) but also in a change in the ongoing EEG/MEG in form of an event-related desynchronization (ERD) or event-related synchronization (ERS). The ERP on the one side and the ERD/ERS on the other side are different responses of neuronal structures in the brain. While the former is phase-locked, the latter is not phase-locked to the event. The most important difference between both phenomena is that the ERD/ERS is highly frequency bandspecific, whereby either the same or different locations on the scalp can display ERD and ERS simultaneously. Quantification of ERD/ERS in time and space is demonstrated on data from a number of movement experiments. q 1999 Elsevier Science Ireland Ltd. All rights reserved.

Event-related EEG/MEG synchronization and desynchronization: Basic principles

A theoretical framework supporting experimental measures of dynamic properties of human EEG is proposed with emphasis on distinct alpha rhythms. Robust relationships between measured dynamics and cognitive or behavioral conditions are reviewed, and proposed physiological bases for EEG at cellular levels are considered. Classical EEG data are interpreted in the context of a conceptual framework that distinguishes between locally and globally dominated dynamic processes, as estimated with coherence or other measures of phase synchronization. Macroscopic (scalp) potentials generated by cortical current sources are described at three spatial scales, taking advantage of the columnar structure of neocortex. New EEG data demonstrate that both globally coherent and locally dominated behavior can occur within the alpha band, depending on narrow band frequency, spatial measurement scale, and brain state. Quasi-stable alpha phase structures consistent with global standing waves are observed. At the same time, alpha and theta phase locking between cortical regions during mental calculations is demonstrated, consistent with neural network formation. The brain-binding problem is considered in the context of EEG dynamic behavior that generally exhibits both of these local and global aspects. But specific experimental designs and data analysis methods may severely bias physiological interpretations in either local or global directions.

Spatial-temporal structures of human alpha rhythms: theory, microcurrent sources, multiscale measurements, and global binding of local networks

Spline generated surface Laplacians are introduced as an effective method for estimating neocortical source activity at moderate scales. The method appears to be robust to the unavoidable perturbations of measured potentials and errors of head geometry and resistivity that are certain to occur in clinical or research settings. In particular, the surface Laplacian is derived for general ellipsoidal surfaces in terms of the spline function. The spline-Laplacian accurately estimates isolated dipoles or distributed sources, is insensitive to subcortical sources and to sources which originate outside the boundaries of the electrode array, and acts as a bandpass spatial filter whose characteristics appear to provide a good match to the volume conduction of intracranial sources through human heads. As a result, spatial resolution is improved over that obtained with conventional EEG by at least a factor of three. This improvement is likely to have a significant impact on both medical and cognitive studies involving EEG. >

S.K. Law

Papers

High-resolution EEG using spline generated surface Laplacians on spherical and ellipsoidal surfaces