Some of the most long-standing questions in paleoanthropology concern how and why human bipedalism evolved. Over the last century, many hypotheses have been offered on the mode of locomotion from which bipedalism originated. Candidate ancestral adaptations include monkey-like arboreal or terrestrial quadrupedalism, gibbon- or orangutan-like (or other forms of) climbing and suspension, and knuckle-walking. This paper reviews the history of these hypotheses, outlines their predictions, and assesses them in light of current phylogenetic, comparative anatomical, and fossil evidence. The functional significance of characteristics of the shoulder and arm, elbow, wrist, and hand shared by African apes and humans, including their fossil relatives, most strongly supports the knuckle-walking hypothesis, which reconstructs the ancestor as being adapted to knuckle-walking and arboreal climbing. Future fossil discoveries, and a clear understanding of anthropoid locomotor anatomy, are required to ultimately test these hypotheses. If knuckle-walking was an important component of the behavioral repertoire of the prebipedal human ancestor, then we can reject scenarios on the origin of bipedalism that rely on a strictly arboreal ancestor. Moreover, paleoenvironmental data associated with the earliest hominins, and their close relatives, contradict hypotheses that place the agents of selection for bipedality in open savanna habitats. Existing hypotheses must explain why bipedalism would evolve from an ancestor that was already partly terrestrial. Many food acquisition and carrying hypotheses remain tenable in light of current evidence.

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Origin of human bipedalism: The knuckle-walking hypothesis revisited.

The human gluteus maximus is a distinctive muscle in terms of size, anatomy and function compared to apes and other non-human primates. Here we employ electromyographic and kinematic analyses of human subjects to test the hypothesis that the human gluteus maximus plays a more important role in running than walking. The results indicate that the gluteus maximus is mostly quiescent with low levels of activity during level and uphill walking, but increases substantially in activity and alters its timing with respect to speed during running. The major functions of the gluteus maximus during running are to control flexion of the trunk on the stance-side and to decelerate the swing leg; contractions of the stance-side gluteus maximus may also help to control flexion of the hip and to extend the thigh. Evidence for when the gluteus maximus became enlarged in human evolution is equivocal, but the muscle's minimal functional role during walking supports the hypothesis that enlargement of the gluteus maximus was likely important in the evolution of hominid running capabilities.

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The human gluteus maximus and its role in running

The telemetered electromyographic activity (EMG) of select hindlimb muscles of unrestrained cats during standing, walking, trotting, and galloping have been recorded. Simultaneous cinematographic records permitted close correlation of muscle activity and locomotor behavior. In general, the pattern of extensor activity of the ankle, knee, and hip during locomotion is fairly consistent, while that of the flexors is more variable.



Changes in basic EMG patterns from walk, to trot, to gallop are most evident in the two-jointed muscles associated with the knee and hip. Progressively greater variation of activity onset and cessation can be seen among extensor muscle groups from the walk, to trot, to gallop. Co-activation of the joint extensors and flexors, especially of the hip, at the end of the stance phase (E3) is slight in the walk, moderate in the trot, and considerable in the gallop. These EMG changes are necessary to meet the demands imposed upon the musculature at the faster gaits, particularly galloping, which include limb rigidity as related to loading, momentum as related to the limb's directional change from the stance phase to the swing phase, and lower spinal movements.



The peroneal muscles of the ankle and the gluteal muscles of the hip show extensor activity and act as joint stabilizers during locomotion. Both biceps femoris anterior muscle and biceps femoris posterior muscle show consistent hip extensor patterns at all gaits. During quiet standing, extensor activity about the knee, ankle, and metatarsophalangeal joints is evident; but the hip extensor and flexor musculature is remarkably silent.



EMG data for unrestrained cats are compared to those of dogs on a treadmill (Tokuriki, '73a,b, '74; Wentink, '76) and those recorded from decerebrate cats (mesencephalic preparation) during controlled locomotion (Gambaryan et al., '71). The EMG patterns from decerebrate cats are more consistent at the walk and gallop within functional groups of muscles at the ankle, knee, and hip than the EMG patterns observed in unrestrained cats or animals moving on a treadmill.

The cat step cycle: Electromyographic patterns for hindlimb muscles during posture and unrestrained locomotion

Kinematic analysis has shown a near-stationary proximal carpal row during the dart-thrower's motion, which is believed to provide a stable platform for the generation of force and accuracy during certain power and precision grip activities. This finding is consistent with evidence in the human hand of adaptations that enabled effective manipulation of stones, cylindric wood, and bone tools for throwing and clubbing. There are at least two possible explanations for the observed human proximal carpal row kinematics. One is that it is retained from a previous common ancestor with great apes and previously adapted to some form of foraging or locomotor behavior involving the hands, but was recruited for tool use after we diverged from the apes. The second is that it evolved after our divergence from apes, in synchrony with adaptations in the human hand to the manipulation of tools, and central to the development of the human's unique ability to aim and accelerate tools and weapons.

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The dart-throwing motion of the wrist: is it unique to humans?

The partial skeleton of Australopithecus from the Hadar Formation, Ethiopia, is reconstructed and compared with other primates. It is demonstrated that the skull of A.L. 288-1 is not as chimplike as it was proposed for Australopithecus afarensis and that the cranial fragments do not differ from Australopithecus africanus. Structural features like the funnelshaped thorax and the pelvis, with its broad iliaca for insertion of musculus latissimus dorsi and the long lever arm of the pubic muscular attachments, invoke a high level of suspensorial behavior. In our opinion, A. africanus is a generalized hominid primate that differs from the specialized African apes, the living pongids beeing too derived to represent a model of a primitive hominoid or hominid ancestor.

Eine Rekonstruktion des Skelettes von A.L. 288-1 (Hadar) und deren Konsequenzen

Bipedality not only frees the hands for tool use but also enhances tool use by allowing use of the trunk for leverage in applying force and thus imparting greater final velocity to tools. Since the weight and acceleration of the trunk and forelimbs on the hindlimbs must be counteracted by muscles such as m. gluteus maximus that control pelvic and trunk movements, it is suggested that the large size of the cranial portion of the human gluteus maximus muscle and its unique attachment to the dorsal ilium (which is apparent in the Makapan australopithecine ilium) may have contributed to the effectiveness with which trunk movement was exploited in early hominid foraging activities. To test this hypothesis, the cranial portions of both right and left muscles were investigated in six human subjects with electromyography during throwing, clubbing, digging, and lifting. The muscles were found to be significantly recruited when the trunk is used in throwing and clubbing, initiating rotation of the pelvis and braking it as trunk rotation ceases and the forelimb accelerates. They stabilize the pelvis during digging and exhibit marked and prolonged activity when the trunk is maintained in partial flexion during lifting of heavy objects.

Gluteus maximus muscle function and the origin of hominid bipedality.

Eight highly skilled male distance runners were filmed while running on level and inclined treadmills. During the filming sequence, EMG measurements were collected on vastus medialis (VM) and semimembranosus (SM) of the right thigh. The initiation and duration of electrical activity of the two muscles was extracted from the recordings relative to the step cycle events. Analysis of covariance for repeated measures was utilized as a statistical treatment to examine the effect of treadmill grade (– 10%, 0%, and + 10%) upon step cycle and EMG data. Results indicate that treadmill grade had a significant effect on the relative duration of both stance and swing phases of the running step cycle and total step cycle duration. Total step cycle time and relative swing phase duration decreased as the treadmill grade was changed from a negative grade to a zero grade to a positive grade while the duration of the stance phase increased. EMG measures indicated decreases in relative duration of SM activity durin...

Electromyographic Patterns During Level and Inclined Treadmill Running and Their Relationship to Step Cycle Measures

Kinematics of locomotion in cats with partially deafferented spinal cords: the spared-root preparation.

Two recurring problems hampering investigations of muscular activity patterns during locomotion are electrical “noise” and movement artifact in the electromyographic (EMG) signal and proper synchronization of behavioral events to the EMG. A relatively simple technique for achieving these ends is presented. Vigorous locomotor movements of unrestrained animals are recorded on 16-mm film and synchronized with one or more EMGs from selected muscles with little electrical noise or movement artifact interference. The system employs telemetry which is compared to hardwire techniques.

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Stanley A. Rasmussen

Papers

Gluteus maximus muscle function and the origin of hominid bipedality.

Electromyographic Patterns During Level and Inclined Treadmill Running and Their Relationship to Step Cycle Measures

Kinematics of locomotion in cats with partially deafferented spinal cords: the spared-root preparation.

A simple technique for correlating film and EMG in locomotion studies

Mirrored Three-View Cinematography in Small Animal Locomotion Studies