Analysis of joint force and torque for the human and non-human ape foot during bipedal walking with implications for the evolution of the foot

The feet of apes have a different morphology from those of humans. Until now, it has merely been assumed that the morphology seen in humans must be adaptive for habitual bipedal walking, as the habitual use of bipedal walking is generally regarded as one of the most clear-cut differences between humans and apes. This study asks simply whether human skeletal proportions do actually enhance foot performance during human-like bipedalism, by examining the influence of foot proportions on force, torque and work in the foot joints during simulated bipedal walking. Skeletons of the common chimpanzee, orangutan, gorilla and human were represented by multi-rigid-body models, where the components of the foot make external contact via finite element surfaces. The models were driven by identical joint motion functions collected from experiments on human walking. Simulated contact forces between the ground and the foot were found to be reasonably comparable with measurements made during human walking using pressure- and force-platforms. Joint force, torque and work in the foot were then predicted. Within the limitations of our model, the results show that during simulated human-like bipedal walking, (1) the human and non-human ape (NHA) feet carry similar joint forces, although the distributions of the forces differ; (2) the NHA foot incurs larger joint torques than does the human foot, although the human foot has higher values in the first tarso-metatarsal and metatarso-phalangeal joints, whereas the NHA foot incurs higher values in the lateral digits; and (3) total work in the metatarso-phalangeal joints is lower in the human foot than in the NHA foot. The results indicate that human foot proportions are indeed well suited to performance in normal human walking.