Cortical involvement during upright walking is not well-studied in humans. We analyzed non-invasive electroencephalographic (EEG) recordings from able-bodied volunteers who participated in a robot-assisted gait-training experiment. To enable functional neuroimaging during walking, we applied source modeling to high-density (120 channels) EEG recordings using individual anatomy reconstructed from structural magnetic resonance imaging scans. First, we analyzed amplitude differences between the conditions, walking and upright standing. Second, we investigated amplitude modulations related to the gait phase. During active walking upper μ (10-12 Hz) and β (18-30 Hz) oscillations were suppressed [event-related desynchronization (ERD)] compared to upright standing. Significant β ERD activity was located focally in central sensorimotor areas for 9/10 subjects. Additionally, we found that low γ (24-40 Hz) amplitudes were modulated related to the gait phase. Because there is a certain frequency band overlap between sustained β ERD and gait phase related modulations in the low γ range, these two phenomena are superimposed. Thus, we observe gait phase related amplitude modulations at a certain ERD level. We conclude that sustained μ and β ERD reflect a movement related state change of cortical excitability while gait phase related modulations in the low γ represent the motion sequence timing during gait. Interestingly, the center frequencies of sustained β ERD and gait phase modulated amplitudes were identified to be different. They may therefore be caused by different neuronal rhythms, which should be taken under consideration in future studies.
Keywords: brain mapping; electroencephalography (EEG); gait; magnetic resonance imaging; motor cortex.