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.

Neuromechanical linkage between the head and forearm during running.

OBJECTIVES: The main objective was to test the hypothesis of a neuromechanical link in humans between the head and forearm during running mediated by the biceps brachii and superior trapezius muscles. We hypothesized that this linkage helps stabilize the head and combats rapid forward pitching during running which may interfere with gaze stability. MATERIALS AND METHODS: Thirteen human participants walked and ran on a treadmill while motion capture recorded body segment kinematics and electromyographic sensors recorded muscle activation. To test perturbations to the linkage system we compared participants running normally as well as with added mass to the face and the hand. RESULTS: The results confirm the presence of a neuromechanical linkage between the head and forearm mediated by the biceps and superior trapezius during running but not during walking. In running, the biceps and superior trapezius activations were temporally linked during the stride cycle, and adding mass to either the head or hand increased activation in both muscles, consistent with our hypothesis. During walking the forces acting on the body segments and muscle activation levels were much smaller than during running, indicating no need for a linkage to keep the head and gaze stable. DISCUSSION: The results suggest that the evolution of long distance running in early Homo may have favored selection for reduced rotational inertia of both the head and forearm through synergistic muscle activation, contributing to the transition from australopith head and forelimb morphology to the more human-like form of Homo erectus. Selective pressures from the evolution of bipedal walking were likely much smaller, but may explain in part the intermediate form of the australopith scapula between that of extant apes and humans.

Straight arm walking, bent arm running: gait-specific elbow angles.

Stereotypically, walking and running gaits in humans exhibit different arm swing behavior: during walking, the arm is kept mostly straight, while during running, the arm is bent at the elbow. The mechanism for this behavioral difference has not been explored before. We hypothesized that a mechanical tradeoff exists between the shoulder joint and the elbow joint. Bending the elbow reduces the radius of gyration of the arm and reduces shoulder muscle torque, but at the price of increasing elbow torque. We predicted that the mechanical tradeoff would result in energetics that favored straight arms during walking and bent arms during running. The hypothesis was tested experimentally by having eight subjects walk and run with both straight arms and bent arms while recording arm swing mechanics, and oxygen consumption in a subset of six subjects. The mechanical tradeoff hypothesis was confirmed, with bent arms reducing normalized shoulder muscle torque in both gaits (walking: -33%, running: -32%) and increasing normalized elbow muscle torque in both gaits (walking: +110%, running: +30%). Bent arms increased oxygen consumption by 11% when walking, supporting our prediction that energetics favor straight arms during walking. However, oxygen consumption was equivalent for the straight and bent arm running conditions, and did not support our running prediction. We conclude that straight arms are stereotyped in walking as a result of optimal energetics, while the mechanism leading to bent arms during running remains unknown.