Origin, structure, and role of background EEG activity. Part 2. Analytic phase.
OBJECTIVE
To explain spontaneous EEG through measurements of spatiotemporal patterns of phase among beta-gamma oscillations.


METHODS
High-density 8 x 8 intracranial arrays were fixed over sensory cortices of rabbits. EEGs were spatially low pass filtered, temporally bandpass filtered and segmented in overlapping windows stepped at 2 ms. Phase was measured with the cosine as the temporal basis function, using both Fourier and Hilbert transforms to compensate for their respective limitations. Spatial patterns in 2D phase surfaces were measured with the geometric form of the cone as the spatial basis function.


RESULTS
Two fundamental state variables were measured at each digitizing step in the 64 EEGs: the rate of change in phase with time (frequency) and the rate of change in phase with distance (gradient). The parameters of location, diameter, duration, and phase velocity of the cone of phase were derived from these two state variables. Parameter distributions including recurrence intervals extending into the low theta range were fractal; the mean values varied with window duration and interelectrode distance.


CONCLUSIONS
The formation of spatial amplitude patterns began with state transitions that were documented by phase discontinuities and phase cones. The multiplicity of overlapping cones indicated that sensory neocortices maintained a scale-free state of self-organized criticality (SOC) in each hemisphere as the basis for its rapid integration of sensory input with prior learning stored in cortical synaptic webs. Further evidence came from the fractal properties of the phase parameters and the self-similarity of phase patterns in the ms/mm to m/s ranges.


CONCLUSIONS
These EEG data suggest that neocortical dynamics is analogous to the dynamics of self-stabilizing systems, such as a sand pile that maintains its critical angle by avalanches, and a pan of boiling water that maintains its critical temperature by bubbles that release heat. Beta-gamma oscillations stem from the ability of neocortex to maintain its stability under continuous sensory bombardment. Modeling implies that the critical parameter of neocortex (analogous to angle of repose or temperature) is the mean firing rates of neurons that are homeostatically regulated by refractory periods everywhere at all times in cortex. The advantage of SOC in perception may be the ability it gives neocortex to generate instantaneous global state transitions (avalanches, bubbles) large enough to include the multiple sensory areas that are necessary to form gestalts (multisensory percepts).
Freeman
Walter J
WJ
Department of Molecular and Cell Biology, LSA 142, University of California, Berkeley, CA 94720-3200, USA. wfreeman@socrates.berkeley.edu
D000818:Animals; D003213:Conditioning, Psychological / Q000502:physiology; D004192:Discrimination, Psychological / Q000502:physiology; D004569:Electroencephalography; D005583:Fourier Analysis; D008959:Models, Neurological; D019579:Neocortex / Q000502:physiology*; D011817:Rabbits