Neuron phase shift adaptive to time delay in locomotor control

Based on neurophysiological evidence, theoretical studies have shown that walking can be generated by mutual entrainment of oscillations of a central pattern generator (CPG) and a body. However, it has also been shown that the time delay in the sensorimotor loop destabilizes mutual entrainment, and results in the failure to walk. Recently, it has been reported that if (a) the neuron model used to construct the CPG is replaced by physiologically faithful neuron model (Bonhoeffer–Van der Pol type) and (b) the mechanical impedance of the body (muscle viscoelasticity) is controlled depending on the angle between two legs, the phase relationship between CPG activity and body motion could be flexibly locked according to the loop delay and, therefore, mutual entrainment can be stabilized. That is, locomotor control adaptive to the loop delay can emerge from the coupling between CPG and body. Here, we call this mechanism flexible-phase locking. In this paper, we construct a system of coupled oscillators as a simplified model of a walking system to theoretically investigate the mechanism of flexible-phase locking, and to analyze the simplified model. The analysis suggests that the following are required as the essential mechanism: (i) an asymptotically stable limit cycle of the coupling system of CPG and body and (ii) a sign difference between afferent and efferent coupling coefficients.