The speed and phase of locomotion dictate saccade probability and simultaneous low-frequency power spectra

Every day we make thousands of saccades and take thousands of steps as we explore our environment. Despite their common co-occurrence in a typical active state, we know little about the coordination between eye movements, walking behaviour and related changes in cortical activity. Technical limitations have been a major impediment, which we overcome here by leveraging the advantages of an immersive wireless virtual reality (VR) environment with three-dimensional (3D) position tracking, together with simultaneous recording of eye movements and mobile electroencephalography (EEG). Using this approach with participants engaged in unencumbered walking along a clear, level path, we find that the likelihood of eye movements at both slow and natural walking speeds entrains to the rhythm of footfall, peaking after the heel-strike of each step. Compared to previous research, this entrainment was captured in a task that did not require visually guided stepping - suggesting a persistent interaction between locomotor and visuomotor functions. Simultaneous EEG recordings reveal a concomitant modulation entrained to heel-strike, with increases and decreases in oscillatory power for a broad range of frequencies. The peak of these effects occurred in the theta and alpha range for slow and natural walking speeds, respectively. Together, our data show that the phase of the step-cycle influences other behaviours such as eye movements, and produces related modulations of simultaneous EEG following the same rhythmic pattern. These results reveal gait as an important factor to be considered when interpreting saccadic and time-frequency EEG data in active observers, and demonstrate that saccadic entrainment to gait may persist throughout everyday activities.