Multiple strategies to correct errors in foot placement and control speed in human walking

Neural feedback plays a key role in maintaining locomotor stability in the face of perturbations. In this study, we systematically identified properties of neural feedback that contribute to stabilizing human walking by examining how the nervous system responds to small kinematic deviations away from the desired gait pattern. We collected data from 20 participants (9 men and 11 women). We simultaneously applied (1) small continuous mechanical perturbations, forces at the ankles that affected foot placement, and (2) small continuous sensory perturbations, movement of a virtual visual scene that produced the illusion of change in walking speed, to compare how neural feedback responds to actual and illusory kinematic deviations. We computed phase-dependent impulse response functions that describe kinematic and muscular responses to small brief perturbations to identify critical phases of the gait cycle when the nervous system modulates muscle activity. In addition to the known foot-placement strategies that counteract kinematic displacement, such as the modulation of the hamstring muscle group during swing, we identified phase-specific muscle modulations that compensated for the perturbations. In particular, our results suggested that an early-stance modulation of anterior leg muscles (i.e., dorsiflexors and quadriceps) is a general control mechanism that serves to control forward body propulsion and compensates for errors in foot placement. Another detected general compensatory strategy was the late-stance modulation of the rectus femoris and gastrocnemius muscles, which controls walking speed in the next cycle.