Jack T. Stern

Climbing to the top: A personal memoir of Australopithecus afarensis

Last autumn marked the 25 anniversary of the discovery of “Lucy.” While that certainly was a momentous event in paleoanthropology, it had no less profound an effect on my academic life, for it presaged my eventual seduction into the arena of hominid fossil interpretation. My friend John Fleagle, editor of Evolutionary Anthropology, says I may introduce this paper with a history of that experience. He assures me this is appropriate because I have now reached the age when young people in the field have no idea who I am.

Functional Morphology of Homo habilis

Olduvai hominid (O.H.) fossils 7, 8, and 35 represent the earliest species of the genus Homo dated at 1.76 million years. The O.H. 7 hand, jaw, and skull and the O.H. 8 foot come from one subadult individual, and the O.H. 35 leg are also those of Homo habilis. The skeleton represents a mosaic of primitive and derived features, indicating an early hominid which walked bipedally and could fabricate stone tools but also retained the generalized hominoid capacity to climb trees.

Body proportions, skeletal allometry and locomotion in the hadar hominids: a reply to Wolpoff

Wolpoff, 1983a , Wolpoff, 1983b ) has challenged the interpretation of relative limb proportions and locomotion offered by Jungers (1982) for Australopithecus afarensis in comparison to modern humans of similar body size. In this reply, we demonstrate that Wolpoffs conclusions are based on several misconceptions about the biomechanics and energetics of bipedalism and a misapplication of the allometric approach. Compared with modern humans (pygmies) of comparably small size (measured as cstimated body weight and as a linear variable correlated to body weight in modern humans), “Lucy” (AL 288-1) possesses very short hindlimbs outside of the known human range and which are proportioned most similarly to small-bodied African apes. By contrast, relative humerus length of Lucy can be matched precisely in a skeletal sample of human pygmics. A similar combination of relative limb lengths appears to exist in the larger individuals of A. afarensis and probably characterizes all gracile australopithecines (including STS 14). The functional implications of interlimb proportions, relative fore-and hindlimb lengths and relative foot length for locomotor biomechanics in A afarensis are also examined. The original interpretations put forth by Jungers (1982) are reaffirmed and strengthened by new data and appropriate analyses.

Electromyographic Studies of the Human Foot: Experimental Approaches to Hominid Evolution

Theories about the functions of the foot muscles have centered on their role in arch support. Previous anatomical and electromyographic studies (reviewed herein) have demonstrated that the arches are normally maintained by bones and ligaments. This study reports an electromyographic investigation of five foot muscles (flexor digito-rum longus, flexor digitorum brevis, flexor accessorius, abductor hallucis, and abductor digiti quinti) conducted on four humans. The three toe flexors act together to resist extension of the toes during the stance phase of locomotion. Despite the large flexor accessorius in humans, neither this muscle nor the flexor digitorum brevis are preferentially recruited over the flexor digitorum lon-gus for any normal posture or locomotion. The abductors affect the mediolateral distribution of pressure by positioning the forefoot. We suggest that the foot muscles play an important role in positioning of the forces on the foot in both posture and locomotion. Future electromyographic experiments on human and ape foot muscles in conjunction with detailed studies of early hominid fossils promise to elucidate the pathways of human locomotor evolution.

Preliminary electromyographical analysis of brachiation in gibbon and spider monkey

In order to refine the concept of brachiation as a locomotor mode and to examine the complex relationship between locomotor behavior and muscle morphology, we have undertaken a telemetered electromyographic (EMG) analysis of muscle recruitment in brachiating gibbons (Hylobates lar) and spider monkeys (Ateles belzebuth andAteles fusciceps) Electrical activity patterns were determined for both support and swing phases in the following muscles: cranial pectoralis major, caudal pectoralis major, middle deltoideus, short head of biceps brachii, flexor digitorum superficialis, latissimus dorsi, and dorsoepitrochlearis. Our experimental findings reinforce earlier behavioral observations that brachiation is not a discrete, stereotyped locomotor activity. EMG patterns differed most between gibbon and spider monkey in those muscles that exhibit markedly disparate morphologies in the two genera-pectoralis major (both portions) and the short head of biceps brachii. Additional recruitment differences appear related to consistent species-specific differences in the timing and mechanics of both support and swing phases, and probably to the role of the prehensile tail as a fail-safe mechanism in the spider monkey.

Humeral retractor EMG during quadrupedal walking in primates

The mammalian humeral retractors latissimus dorsi, teres major and caudal parts of the pectoral muscles are commonly thought to contribute to forward impulse during quadrupedal locomotion by pulling the body over the supporting forelimb. While most electromyographic studies on recruitment patterns for these muscles tend to support this functional interpretation, data on muscle use in chimpanzees and vervet monkeys have suggested that the humeral retractors of nonhuman primates are largely inactive during the support phase of quadrupedal locomotion. In the chimpanzee and vervet monkey, in contrast to what has been documented for other mammals, the contributions of latissimus dorsi, caudal pectoralis major, and teres major during quadrupedal locomotion are restricted to slowing down the swinging forelimb in preparation for hand touchdown and/or retracting the humerus to help lift the hand off the substrate at the initiation of swing phase. Based on these results, it has been proposed that unique patterns of shoulder muscle recruitment are among a set of characteristics that distinguish the form of quadrupedalism displayed by nonhuman primates from that of other nonprimate mammals. However, two primate taxa is a limited sample upon which to base such far-reaching conclusions. Here we report on the activity patterns for the humeral retractors during quadrupedal walking in an additional eight species of nonhuman primates. There is some variability in the activity patterns for latissimus dorsi, caudal pectoralis major and teres major, both between and within species, but in general the results confirm that the humeral retractors of primate quadrupeds do not contribute to forward impulse by pulling the body over the supporting forelimb.

A three-dimensional musculoskeletal model of the chimpanzee (Pan troglodytes) pelvis and hind limb

SUMMARY 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.

Electromyography of wrist and finger flexor muscles in olive baboons ( Papio anubis )

SUMMARY Some non-human primates use digitigrade hand postures when walking slowly on the ground. As a component of an extended limb, a digitigrade posture can help minimize wrist joint moments thereby requiring little force production directly from wrist flexors (and/or from the assistance of finger flexors) to maintain limb posture. As a consequence, less active muscle volume would be required from these anti-gravity muscles and overall metabolic costs associated with locomotion could be reduced. To investigate whether the use of digitigrade hand postures during walking in primates entails minimal use of anti-gravity muscles, this study examined electromyography (EMG) patterns in both the wrist and finger flexor muscles in facultatively digitigrade olive baboons (Papio anubis) across a range of speeds. The results demonstrate that baboons can adopt a digitigrade hand posture when standing and moving at slow speeds without requiring substantial EMG activity from distal anti-gravity muscles. Higher speed locomotion, however, entails increasing EMG activity and is accompanied by a dynamic shift to a more palmigrade-like limb posture. Thus, the ability to adopt a digitigrade hand posture by monkeys is an adaptation for ground living, but it was never co-opted for fast locomotion. Rather, digitigrady in primates appears to be related to energetic efficiency for walking long distances.

Electromyography of crural and pedal muscles in tufted capuchin monkeys (Sapajus apella): Implications for hallucal grasping behavior and first metatarsal morphology in euprimates.

A hypertrophied peroneal process of the hallucal metatarsal, as seen in prosimians, has been linked to a powerful hallucal grasp via the contraction of the peroneus longus (PL) muscle causing adduction of the big toe. Electromyography (EMG) studies of lemurs and lorises, however, have concluded that PL is not substantially recruited during small branch locomotion when powerful hallucal grasping is needed most, and have suggested that there is no link between PL activity and peroneal process size. If this is correct, then we should also observe no change in PL activity when strong hallucal grasping is required in anthropoids because they have a relatively smaller peroneal process for PL to act on. This study addresses this hypothesis by evaluating EMG of crural and pedal muscles in capuchins (Sapajus apella) walking on substrates of different diameters. During locomotion on the narrow substrate (3.1 cm) that should elicit a strong hallucal grasp, we observed an intense increased recruitment of adductor hallucis, but only sustained, rather than markedly increased, PL activity. This indicates that PL is not involved in powerful hallucal grasping in capuchins, and confirms similar findings previously documented in prosimians. We continue to reject the hypothesis that a large peroneal process is an adaptation for powerful grasping and further argue that its morphology may not be related to PLs ability to adduct the hallux at all. In addition, the morphology of the peroneal process should not be used to assess hallucal grasping performance in fossils.

Function of the Subclavius Muscle in a Nonhuman Primate, the Spider Monkey (Ateles)

Within the primate order, the morphology of the subclavius muscle is known to be unique among the prehensile-tailed South American monkeys. 3 spider monkeys, Ateles, were monitored electromyographically to determine the recruitment of this muscle during various locomotor and postural activities. Rather than indicating a static stabilizing function, which has typically been inferred from classical anatomical studies, results from this study suggest that the subclavius performs more as a dynamic element in movements of the pectoral girdle during brachiation, vertical climbing, pronograde quadrupedalism and leaping. Complementary activity patterns were also identified between the subclavius and the caudal fibers of the trapezius indicating that the subclavius is used when the animal must depress, or resist cranial displacement, of the protracted shoulder girdle, while the caudal trapezius is recruited when the girdle is retracted on the chest wall.