Neuromusculoskeletal computer modeling and simulation of upright, straight-legged, bipedal locomotion of Australopithecus afarensis (A.L. 288-1)

The skeleton of Australopithecus afarensis (A.L. 288-1, better known as "Lucy") is by far the most complete record of locomotor morphology of early hominids currently available. Even though researchers agree that the postcranial skeleton of Lucy shows morphological features indicative of bipedality, only a few studies have investigated Lucy's bipedal locomotion itself. Lucy's energy expenditure during locomotion has been the topic of much speculation, but has not been investigated, except for several estimates derived from experimental data collected on other animals. To gain further insights into how Lucy may have walked, we generated a full three-dimensional (3D) reconstruction and forward-dynamic simulation of upright bipedal locomotion of this ancient human ancestor. Laser-scanned 3D bone geometries were combined with state-of-the-art neuromusculoskeletal modeling and simulation techniques from computational biomechanics. A detailed full 3D neuromusculoskeletal model was developed that encompassed all major bones, joints (10), and muscles (52) of the lower extremity. A model of muscle force and heat production was used to actuate the musculoskeletal system, and to estimate total energy expenditure during locomotion. Neural activation profiles for each of the 52 muscles that produced a single step of locomotion, while at the same time minimizing the energy consumed per meter traveled, were searched through numerical optimization. The numerical optimization resulted in smooth locomotor kinematics, and the predicted energy expenditure was appropriate for upright bipedal walking in an individual of Lucy's body size.