Exploring the effects of skeletal architecture and muscle properties on bipedal standing in the common chimpanzee ( Pan troglodytes ) from the perspective of biomechanics.

It is well known that the common chimpanzee, as both the closest living relative to humans and a facultative bipedal, has the capability of bipedal standing but cannot do so fully upright. Accordingly, they have been of exceeding significance in elucidating the evolution of human bipedalism. There are many reasons why the common chimpanzee can only stand with its hips-knees bent, such as the distally oriented long ischial tubercle and the almost absent lumbar lordosis. However, it is unknown how the relative positions of their shoulder-hip-knee-ankle joints are coordinated. Similarly, the distribution of the biomechanical characteristics of the lower-limb muscles and the factors that affect the erectness of standing as well as the muscle fatigue of the lower limbs remain a mystery. The answers are bound to light up the evolutional mechanism of hominin bipedality, but these conundrums have not been shed much light upon, because few studies have comprehensively explored the effects of skeletal architecture and muscle properties on bipedal standing in common chimpanzees. Thus, we first built a musculoskeletal model comprising the head-arms-trunk (HAT), thighs, shanks, and feet segments of the common chimpanzee, and then, the mechanical relationships of the Hill-type muscle-tendon units (MTUs) in bipedal standing were deduced. Thereafter, the equilibrium constraints were established, and a constrained optimization problem was formulated where the optimization objective was defined. Finally, thousands of simulations of bipedal standing experiments were performed to determine the optimal posture and its corresponding MTU parameters including muscle lengths, muscle activation, and muscle forces. Moreover, to quantify the relationship between each pair of the parameters from all the experimental simulation outcomes, the Pearson correlation analysis was employed. Our results demonstrate that in the pursuit of the optimal bipedal standing posture, the common chimpanzee cannot simultaneously achieve maximum erectness and minimum muscle fatigue of the lower limbs. For uni-articular MTUs, the relationship between muscle activation, relative muscle lengths, together with relative muscle forces, and the corresponding joint angle is generally negatively correlated for extensors and positively correlated for flexors. For bi-articular MTUs, the relationship between muscle activation, coupled with relative muscle forces, and the corresponding joint angles does not show the same pattern as in the uni-articular MTUs. The results of this study bridge the gap between skeletal architecture, along with muscle properties, and biomechanical performance of the common chimpanzee during bipedal standing, which enhances existing biomechanical theories and advances the comprehension of bipedal evolution in humans.