Evaluation of the Phase-Dependent Rhythm Control of Human Walking Using Phase Response Curves

Humans and animals control their walking rhythms to maintain motion in a variable environment. The neural mechanism for controlling rhythm has been investigated in many studies using mechanical and electrical stimulation. However, quantitative evaluation of rhythm variation in response to perturbation at various timings has rarely been investigated. Such a characteristic of rhythm is described by the phase response curve (PRC). Dynamical simulations of human skeletal models with changing walking rhythms (phase reset) described a relation between the effective phase reset on stability and PRC, and phase reset around touch-down was shown to improve stability. A PRC of human walking was estimated by pulling the swing leg, but such perturbations hardly influenced the stance leg, so the relation between the PRC and walking events was difficult to discuss. This research thus examines human response to variations in floor velocity. Such perturbation yields another problem, in that the swing leg is indirectly (and weakly) perturbed, so the precision of PRC decreases. To solve this problem, this research adopts the weighted spike-triggered average (WSTA) method. In the WSTA method, a sequential pulsed perturbation is used for stimulation. This is in contrast with the conventional impulse method, which applies an intermittent impulsive perturbation. The WSTA method can be used to analyze responses to a large number of perturbations for each sequence. In the experiment, perturbations are applied to walking subjects by rapidly accelerating and decelerating a treadmill belt, and measured data are analyzed by the WSTA and impulse methods. The PRC obtained by the WSTA method had clear and stable waveforms with a higher temporal resolution than those obtained by the impulse method. By investigation of the rhythm transition for each phase of walking using the obtained PRC, a rhythm change that extends the touch-down and mid-single support phases is found to occur.Author Summary: Humans and animals tune their walking rhythms when motion is disturbed, such that they hesitate before making the transition from stance to swing phase. The effectiveness of rhythm control for stability has also been shown, and thus the elucidation of rhythm responses is important to understanding human strategies for walking control. In this research, how and when humans change their walking rhythm in response to disturbance is analyzed over the complete walking cycle. Phase response of human walking has previously been estimated by pulling the swing leg. The problem with this perturbation is that it hardly disturbs the stance leg, so here we apply the perturbation by changing floor velocity. However, perturbation from the floor yields another problem in that it weakly influences the swing leg, decreasing the precision of the PRC. The present research tackles this problem by introducing a new method for identifying rhythm characteristics by use of high-frequency perturbation, which allows us to obtain results with clear temporal resolution. We found that the human walking rhythm changes by lengthening the touch-down and mid-single support phases. These phase responses are compared with neural mechanisms for rhythm control, and relevance to the cutaneous and proprioceptive originated responses is shown.