Forward dynamic simulation of Japanese macaque bipedal locomotion demonstrates better energetic economy in a virtualised plantigrade posture

A plantigrade foot with a large robust calcaneus is regarded as a distinctive morphological feature of the human foot; it is presumably the result of adaptation for habitual bipedal locomotion. The foot of the Japanese macaque, on the other hand, does not have such a feature, which hampers it from making foot–ground contact at the heel during bipedal locomotion. Understanding how this morphological difference functionally affects the generation of bipedal locomotion is crucial for elucidating the evolution of human bipedalism. In this study, we constructed a forward dynamic simulation of bipedal locomotion in the Japanese macaque based on a neuromusculoskeletal model to evaluate how virtual manipulation of the foot structure from digitigrade to plantigrade affects the kinematics, dynamics, and energetics of bipedal locomotion in a nonhuman primate whose musculoskeletal anatomy is not adapted to bipedalism. The normal bipedal locomotion generated was in good agreement with that of actual Japanese macaques. If, as in human walking, the foot morphology was altered to allow heel contact, the vertical ground reaction force profile became double-peaked and the cost of transport decreased. These results suggest that evolutionary changes in the foot structure were important for the acquisition of human-like efficient bipedal locomotion.