Shorter distal forelimbs benefit bipedal walking and running mechanics: Implications for hominin forelimb evolution.

OBJECTIVES Brachial index is a skeletal ratio that describes the relative length of the distal forelimb. Over the course of hominin evolution, a shift toward smaller brachial indices occurred. First, Pleistocene australopiths yield values between extant chimpanzees and humans, with further evolution in Pliocene Homo to the modern human range. We hypothesized that shorter distal forelimbs benefit walking and running performance, notably elbow and shoulder joint torques, and predicted that the benefit would be greater in running compared to walking. MATERIALS AND METHODS We tested our hypothesis in a modern human sample walking and running while carrying hand weights, which increase the inertia (mass and effective length) of the distal forelimb, simulating a larger brachial index. RESULTS We found longer distal forelimbs and the added mass increased elbow muscle torque by 98% while walking and 70% in running, confirming our hypothesis that shorter distal forelimbs benefit walking and running performance. Shoulder muscle torque similarly increased in both gaits with the addition of hand weights due to elongation of the effective forelimb length. Normalized elbow torque, which accounted for the effect on shoulder torque caused by the experimental manipulation, increased by 16% while walking but 52% while running, indicating that shorter distal forelimbs provide a greater benefit for running by approximately three-fold. DISCUSSION Selection for economical bipedal walking in Australopithecus and endurance running in Homo likely contributed to the shift toward relatively smaller distal forelimbs across hominin evolution, with modern human proportions attained in Pleistocene Homo erectus and retained in later species.