Gait-phase-dependent and gait-phase-independent cortical activity across multiple regions involved in voluntary gait modifications in humans

Modification of ongoing walking movement to fit changes in external environments requires accurate voluntary control. In cats, the motor and posterior parietal cortices have crucial roles for precisely adjusting limb trajectory during walking. In human walking, however, it remains unclear which cortical information contributes to voluntary gait modification. In this study, we investigated cortical activity changes associated with visually guided precision stepping using electroencephalography source analysis. Our results demonstrated frequency- and gait-event-dependent changes in the cortical power spectrum elicited by voluntary gait modification. The main differences between normal walking and precision stepping were as follows: (a) the alpha, beta or gamma power decrease during the swing phases in the sensorimotor, anterior cingulate and parieto-occipital cortices, and (b) a power decrease in the theta, alpha and beta bands and increase in the gamma band throughout the gait cycle in the parieto-occipital cortex. Based on the previous knowledge of brain functions, the former change was considered to be related to execution and planning of leg movement, while the latter change was considered to be related to multisensory integration and motor awareness. Therefore, our results suggest that the gait modification is achieved by higher cortical involvements associated with different sensorimotor-related functions across multiple cortical regions including the sensorimotor, anterior cingulate and parieto-occipital cortices. The results imply the critical importance of the cortical contribution to voluntary modification in human locomotion. Further, the observed cortical information related to voluntary gait modification would contribute to developing volitional control systems of brain–machine interfaces for walking rehabilitation.