Recent findings suggest that human cortex is more active during steady-speed unperturbed locomotion than previously thought. However, techniques that have been used to image the brain during locomotion lack the temporal resolution necessary to assess intra-stride cortical dynamics. Our aim was to determine if electrocortical activity is coupled to gait cycle phase during steady-speed human walking. We used electroencephalography (EEG), motion capture, and a force-measuring treadmill to record brain and body dynamics while eight healthy young adult subjects walked on a treadmill. Infomax independent component analysis (ICA) parsed EEG signals into maximally independent component (IC) processes representing electrocortical sources, muscle sources, and artifacts. We calculated a spatially fixed equivalent current dipole for each IC using an inverse modeling approach, and clustered electrocortical sources across subjects by similarities in dipole locations and power spectra. We then computed spectrograms for each electrocortical source that were time-locked to the gait cycle. Electrocortical sources in the anterior cingulate, posterior parietal, and sensorimotor cortex exhibited significant (p<0.05) intra-stride changes in spectral power. During the end of stance, as the leading foot was contacting the ground and the trailing foot was pushing off, alpha- and beta-band spectral power increased in or near the left/right sensorimotor and dorsal anterior cingulate cortex. Power increases in the left/right sensorimotor cortex were more pronounced for contralateral limb push-off (ipsilateral heel-strike) than for ipsilateral limb push-off (contralateral heel-strike). Intra-stride high-gamma spectral power changes were evident in anterior cingulate, posterior parietal, and sensorimotor cortex. These data confirm cortical involvement in steady-speed human locomotion. Future applications of these techniques could provide critical insight into the neural mechanisms of movement disorders and gait rehabilitation.
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