Locomotor kinematics and EMG activity during quadrupedal versus bipedal gait in the Japanese macaque

Abstract

Several qualitative features distinguish bipedal from quadrupedal locomotion in mammals. In this study we show quantitative differences between quadrupedal and bipedal gait in the Japanese monkey in terms of gait patterns, trunk/hindlimb kinematics, and electromyographic (EMG) activity, obtained from 3 macaques during treadmill walking. We predicted that as a consequence of an almost upright body axis, bipedal gait would show properties consistent with temporal and spatial optimization countering higher trunk/hindlimb loads and a less stable center of mass (CoM). A comparatively larger step width, an ~9% longer duty cycle, and ~20% increased relative duration of the double-support phase were all in line with such a strategy. Bipedal joint kinematics showed the strongest differences in proximal, and least in distal, hindlimb joint excursions compared with quadrupedal gait. Hindlimb joint coordination (cyclograms) revealed more periods of single-joint rotations during bipedal gait and predominance of proximal joints during single support. The CoM described a symmetrical, quasisinusoidal left/right path during bipedal gait, with an alternating shift toward the weight-supporting limb during stance. Trunk/hindlimb EMG activity was nonuniformally increased during bipedal gait, most prominently in proximal antigravity muscles during stance (up to 10-fold). Non-antigravity hindlimb EMG showed altered temporal profiles during liftoff or touchdown. Muscle coactivation was more, but muscle synergies less, frequent during bipedal gait. Together, these results show that behavioral and EMG properties of bipedal vs. quadrupedal gait are quantitatively distinct and suggest that the neural control of bipedal primate locomotion underwent specific adaptations to generate these particular behavioral features to counteract increased load and instability. NEW & NOTEWORTHY Bipedal locomotion imposes particular biomechanical constraints on motor control. In a within-species comparative study, we investigated joint kinematics and electromyographic characteristics of bipedal vs. quadrupedal treadmill locomotion in Japanese macaques. Because these features represent (to a large extent) emergent properties of the underlying neural control, they provide a comparative, behavioral, and neurophysiological framework for understanding the neural system dedicated to bipedal locomotion in this nonhuman primate, which constitutes a critical animal model for human bipedalism.