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Primary Publication
O'Neill MC, Lee LF, Larson SG, Demes B, Stern JT Jr, Umberger BR. 2013. A three-dimensional musculoskeletal model of the chimpanzee (Pan troglodytes) pelvis and hind limb. Journal of Experimental Biology 216, 3709-3723. (2013)  View

Musculoskeletal models have become important tools for studying a range of muscle-driven movements. However, most work has been in modern humans, with few applications in other species. Chimpanzees are facultative bipeds and our closest living relatives, and have provided numerous important insights into our own evolution. A chimpanzee musculoskeletal model would allow integration across a wide range of laboratory-based experimental data, providing new insights into the determinants of their locomotor performance capabilities, as well as the origins and evolution of human bipedalism. Here, we described a detailed three-dimensional (3D) musculoskeletal model of the chimpanzee pelvis and hind limb. The model includes geometric representations of bones and joints, as well as 35 muscle-tendon units that were represented using 44 Hill-type muscle models. Muscle architecture data, such as muscle masses, fascicle lengths and pennation angles, were drawn from literature sources. The model permits calculation of 3D muscle moment arms, muscle-tendon lengths and isometric muscle forces over a wide range of joint positions. Muscle-tendon moment arms predicted by the model were generally in good agreement with tendon-excursion estimates from cadaveric specimens. Sensitivity analyses provided information on the parameters that model predictions are most and least sensitive to, which offers important context for interpreting future results obtained with the model. Comparisons with a similar human musculoskeletal model indicate that chimpanzees are better suited for force production over a larger range of joint positions than humans. This study represents an important step in understanding the integrated function of the neuromusculoskeletal systems in chimpanzee locomotion.

Related Publications
ONeill MC, Lee L-F, Demes B, Thompson NE, Larson SG, Stern JT Jr & Umberger BR. 2015. Three-dimensional kinematics of the pelvis and hind limbs in chimpanzee (Pan troglodytes) and human bipedal walking. Journal of Human Evolution 86, 32-42. (2015)  View

The common chimpanzee (Pan troglodytes) is a facultative biped and our closest living relative. As such, the musculoskeletal anatomies of their pelvis and hind limbs have long provided a comparative context for studies of human and fossil hominin locomotion. Yet, how the chimpanzee pelvis and hind limb actually move during bipedal walking is still not well defined. Here, we describe the three-dimensional (3-D) kinematics of the pelvis, hip, knee and ankle during bipedal walking and compare those values to humans walking at the same dimensionless and dimensional velocities. The stride-to-stride and intraspecific variations in 3-D kinematics were calculated using the adjusted coefficient of multiple correlation. Our results indicate that humans walk with a more stable pelvis than chimpanzees, especially in tilt and rotation. Both species exhibit similar magnitudes of pelvis list, but with segment motion that is opposite in phasing. In the hind limb, chimpanzees walk with a more flexed and abducted limb posture, and substantially exceed humans in the magnitude of hip rotation during a stride. The average stride-to-stride variation in joint and segment motion was greater in chimpanzees than humans, while the intraspecific variation was similar on average. These results demonstrate substantial differences between human and chimpanzee bipedal walking, in both the sagittal and non-sagittal planes. These new 3-D kinematic data are fundamental to a comprehensive understanding of the mechanics, energetics and control of chimpanzee bipedalism.

ONeill MC, Umberger BR, Holowka NB, Larson SG, & Reiser PJ. 2017. Chimpanzee super strength and human skeletal muscle evolution. Proceedings of the National Academy of Sciences. 114, 7343-7348. (2017)  View

Chimpanzee “super strength” has been widely reported since the 1920s although a critical review of the available data suggests that the chimpanzee–human muscular performance differential is only ∼1.5 times. Some hypothesize that this differential reflects underlying differences in muscle mechanics. Here, we present direct measurements of chimpanzee skeletal muscle properties in comparison with those of humans and other terrestrial mammals. Our results show that chimpanzee muscle exceeds human muscle in maximum dynamic force and power output by ∼1.35 times. This is primarily due to the chimpanzee’s higher fast-twitch fiber content, rather than exceptional maximum isometric force or maximum shortening velocities. We suggest that muscular performance capabilities declined during hominin evolution in response to selection for repetitive, low-cost contractile behavior.

ONeill MC, Demes B, Thompson NE, & Umberger BR. 2018. Three-dimensional kinematics and the origin of the hominin walking stride. Journal of the Royal Society Interface 15, 20180205. (2018)  View

Humans are unique among apes and other primates in the musculoskeletal design of their lower back and pelvis. While the last common ancestor of the Pan–Homo lineages has long been thought to be ‘African ape-like’, including in its lower back and ilia design, recent descriptions of early hominin and Miocene ape fossils have led to the proposal that its lower back and ilia were more similar to those of some Old World monkeys, such as macaques. Here, we compared three-dimensional kinematics of the pelvis and hind/lower limbs of bipedal macaques, chimpanzees and humans walking at similar dimensionless speeds to test the effects of lower back and ilia design on gait. Our results indicate that locomotor kinematics of bipedal macaques and chimpanzees are remarkably similar, with both species exhibiting greater pelvis motion and more flexed, abducted hind limbs than humans during walking. Some differences between macaques and chimpanzees in pelvis tilt and hip abduction were noted, but they were small in magnitude; larger differences were observed in ankle flexion. Our results suggest that if Pan and Homo diverged from a common ancestor whose lower back and ilia were either ‘African ape-like’ or more ‘Old World monkey-like’, at its origin, the hominin walking stride likely involved distinct (i.e. non-human-like) pelvis motion on flexed, abducted hind limbs.