Analysis of Spatial Patterns of Phase in Neocortical Gamma EEGs in Rabbit

Abstract

Arrays of 64 electrodes (8 × 8, 7 × 7 mm) were implanted epidurally on the surface of the visual, auditory or somatosensory cortex of rabbits trained to discriminate conditioned stimuli in the corresponding modality. The 64 electroencephalographic (EEG) traces at all times displayed a high degree of spatial coherence in wave form, averaging >90% of the variance in the largest principal components analysis component. The EEGs were decomposed with the fast Fourier transform (FFT) to give the spatial distributions of amplitude and phase modulation (AM and PM) in segments 128 ms in duration. Spatial (2-dimensional) and temporal (1-dimensional) filters were designed to optimize classification of the spatial AM patterns in the gamma range (20–80 Hz) with respect to discriminative conditioned stimuli. No evidence was found for stimulus-dependent classification of the spatial PM patterns. Instead some spatial PM distributions conformed to the pattern of a cone. The location and sign (maximal lead or lag) of the conic apex varied randomly with each recurrence. The slope of the phase gradient varied in a range corresponding to that of the conduction velocities reported of axons to extend parallel to the cortical surfaces. The durations and times of recurrence of the phase cones corresponded to those of the optimally classified spatial AM patterns. The interpretation is advanced that the phase cones are manifestations of state transitions in the mesoscopic dynamics of sensory cortices by which the intermittent AM patterns are formed. The phase cones show that the gamma EEG spatial coherence is not due to volume conduction from a single deep-lying dipole generator nor to activity at the site of the reference lead on monopolar recording. The random variation of the apical sign shows that gamma AM patterns are self-organized and are not imposed by thalamic pacemakers. The half-power radius of the phase gradient provides a useful measure of the soft boundary condition for the formation and read-out of cooperative cortical domains responsible for binding sensory information into the context of prior experience in the process of perception.