Striding bipedalism is a key derived behaviour of hominids that possibly originated soon after the divergence of the chimpanzee and human lineages. Although bipedal gaits include walking and running, running is generally considered to have played no major role in human evolution because humans, like apes, are poor sprinters compared to most quadrupeds. Here we assess how well humans perform at sustained long-distance running, and review the physiological and anatomical bases of endurance running capabilities in humans and other mammals. Judged by several criteria, humans perform remarkably well at endurance running, thanks to a diverse array of features, many of which leave traces in the skeleton. The fossil evidence of these features suggests that endurance running is a derived capability of the genus Homo, originating about 2 million years ago, and may have been instrumental in the evolution of the human body form.

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Endurance running and the evolution of homo

The extracellular matrix (ECM), and especially the connective tissue with its collagen, links tissues of the body together and plays an important role in the force transmission and tissue structure maintenance especially in tendons, ligaments, bone, and muscle. The ECM turnover is influenced by physical activity, and both collagen synthesis and degrading metalloprotease enzymes increase with mechanical loading. Both transcription and posttranslational modifications, as well as local and systemic release of growth factors, are enhanced following exercise. For tendons, metabolic activity, circulatory responses, and collagen turnover are demonstrated to be more pronounced in humans than hitherto thought. Conversely, inactivity markedly decreases collagen turnover in both tendon and muscle. Chronic loading in the form of physical training leads both to increased collagen turnover as well as, dependent on the type of collagen in question, some degree of net collagen synthesis. These changes will modify the mechanical properties and the viscoelastic characteristics of the tissue, decrease its stress, and likely make it more load resistant. Cross-linking in connective tissue involves an intimate, enzymatical interplay between collagen synthesis and ECM proteoglycan components during growth and maturation and influences the collagen-derived functional properties of the tissue. With aging, glycation contributes to additional cross-linking which modifies tissue stiffness. Physiological signaling pathways from mechanical loading to changes in ECM most likely involve feedback signaling that results in rapid alterations in the mechanical properties of the ECM. In developing skeletal muscle, an important interplay between muscle cells and the ECM is present, and some evidence from adult human muscle suggests common signaling pathways to stimulate contractile and ECM components. Unaccostumed overloading responses suggest an important role of ECM in the adaptation of myofibrillar structures in adult muscle. Development of overuse injury in tendons involve morphological and biochemical changes including altered collagen typing and fibril size, hypervascularization zones, accumulation of nociceptive substances, and impaired collagen degradation activity. Counteracting these phenomena requires adjusted loading rather than absence of loading in the form of immobilization. Full understanding of these physiological processes will provide the physiological basis for understanding of tissue overloading and injury seen in both tendons and muscle with repetitive work and leisure time physical activity.

Role of Extracellular Matrix in Adaptation of Tendon and Skeletal Muscle to Mechanical Loading

Locomotion results from complex, high-dimensional, non-linear, dynamically coupled interactions between an organism and its environment. Fortunately, simple models we call templates have been and can be made to resolve the redundancy of multiple legs, joints and muscles by seeking synergies and symmetries. A template is the simplest model (least number of variables and parameters) that exhibits a targeted behavior. For example, diverse species that differ in skeletal type, leg number and posture run in a stable manner like sagittal- and horizontal-plane spring-mass systems. Templates suggest control strategies that can be tested against empirical data. Templates must be grounded in more detailed morphological and physiological models to ask specific questions about multiple legs, the joint torques that actuate them, the recruitment of muscles that produce those torques and the neural networks that activate the ensemble. We term these more elaborate models anchors. They introduce representations of specific biological details whose mechanism of coordination is of interest. Since mechanisms require controls, anchors incorporate specific hypotheses concerning the manner in which unnecessary motion or energy from legs, joints and muscles is removed, leaving behind the behavior of the body in the low-degree-of-freedom template. Locating the origin of control is a challenge because neural and mechanical systems are dynamically coupled and both play a role. The control of slow, variable-frequency locomotion appears to be dominated by the nervous system, whereas during rapid, rhythmic locomotion, the control may reside more within the mechanical system. Anchored templates of many-legged, sprawled-postured animals suggest that passive, dynamic self-stabilization from a feedforward, tuned mechanical system can reject rapid perturbations and simplify control. Future progress would benefit from the creation of a field embracing comparative neuromechanics.

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Templates and anchors: neuromechanical hypotheses of legged locomotion on land.

Identifying recently bottlenecked populations (populations severely reduced in size) is important because bottlenecks can increase demographic stochasticity, rate of inbreeding, loss of genetic variation, and fixation of deleterious alleles and, thereby, reduce adaptive potential and increase the probability of population extinction (Frankel & Soulé 1981; Lande 1988, 1994; Leberg 1990; Hedrick & Miller 1992; Mills & Smouse 1994; Frankham 1995 a , 1995 b ; but see Bryant et al. 1986; Goodnight 1987). Unfortunately, it is usually difficult to determine if a population has recently experienced a bottleneck because historical population sizes and levels of genetic variation are seldom known. We developed a statistical test (a sign test for heterozygosity excess) for detecting recent historical bottlenecks using allele frequency data (Cornuet & Luikart 1996). The test requires no data on historical population sizes or levels of genetic variation; it requires only measurements of allele frequencies from 5 to 20 polymorphic loci in a sample of approximately 20-30 individuals. The test has reasonable statistical power when applied to allele frequency data sets generated by computer simulations (Cornuet & Luikart 1996). The performance of the test, however, must be evaluated by means of empirical data from natural populations before it can be used with confidence. Our objectives were to (1) explain to conservation biologists the principle of the sign test for detecting heterozygosity excess and (2) evaluate the reliability of the test by analyzing 56 allozyme and 37 microsatellite data sets from bottlenecked and nonbottlenecked natural populations.

Empirical Evaluation of a Test for Identifying Recently Bottlenecked Populations from Allele Frequency Data

We demonstrate that an irreducibly simple, uncontrolled, two-dimensional, two-link model, vaguely resembling human legs, can walk down a shallow slope, powered only by gravity. This model is the simplest special case of the passive-dynamic models pioneered by McGeer (1990a). It has two rigid massless legs hinged at the hip, a point-mass at the hip, and infinitesimal point-masses at the feet. The feet have plastic (no-slip, no-bounce) collisions with the slope surface, except during forward swinging, when geometric interference (foot scuffing) is ignored. After nondimensionalizing the governing equations, the model has only one free parameter, the ramp slope gamma. This model shows stable walking modes similar to more elaborate models, but allows some use of analytic methods to study its dynamics. The analytic calculations find initial conditions and stability estimates for period-one gait limit cycles. The model exhibits two period-one gait cycles, one of which is stable when 0 < gamma < 0.015 rad. With increasing gamma, stable cycles of higher periods appear, and the walking-like motions apparently become chaotic through a sequence of period doublings. Scaling laws for the model predict that walking speed is proportional to stance angle, stance angle is proportional to gamma 1/3, and that the gravitational power used is proportional to v4 where v is the velocity along the slope.

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The simplest walking model: stability, complexity, and scaling.

Energy can be saved in terrestrial locomotion in many different ways. The maximum shortening speeds (Vmax) of the muscles can be adjusted to their optimum values for the tasks required of them. The moments exerted by the muscles at different joints can be adjusted to keep the ground force in line with the leg so that muscles do not work against each other. The joints of the legs can be kept as straight as possible, minimizing muscle forces and work requirements. Walking gaits should be selected at low Froude numbers (a dimensionless speed parameter) and running gaits at high Froude numbers. Tendon and other springs can be used to store elastic strain energy and to return it by elastic recoil. This paper aims to show how these energy-saving mechanisms work and to what extent mammals exploit them. Arguments based on our rather limited knowledge of the relationship between the mechanical performance of muscle and its metabolic energy consumption are used throughout. They suggest that muscles that are optimally adapted for their tasks in running should do positive work with constant efficiency.

Energy-saving mechanisms in walking and running

Simple mathematical models capable of walking or running are used to compare the merits of bipedal gaits. Stride length, duty factor (the fraction of the stride, for which the foot is on the ground) and the pattern of force on the ground are varied, and the optimum gait is deemed to be the one that minimizes the positive work that the muscles must perform, per unit distance travelled. Even the simplest model, whose legs have neither mass nor elastic compliance, predicts the changes of duty factor and force pattern that people make as they increase their speed of walking. It predicts a sudden change to running at a critical speed, but this is much faster than the speed at which people make the change. When elastic compliance is incorporated in the model, unnaturally fast walking becomes uncompetitive. However, a slow run with very brief foot contact becomes the optimum gait at low speeds, at which people would walk, unless severe energy dissipation occurs in the compliance. A model whose legs have mass as well as elastic compliance predicts well the relationship between speed and stride length in human walking.

A model of bipedal locomotion on compliant legs.

Humans, bushbabies, frogs, locusts, fleas and other animals jump by rapidly extending a pair of legs. Mathematical models are used to investigate the effect muscle properties, leg design and jumping technique have on jump height. Jump height increases with increased isometric force exerted by leg muscles, their maximum shortening speeds and their series compliances. When ground forces are small multiples of body mass (as for humans), countermovement and catapult jumps are about equally high, and both are much better than squat jumps. Vertebrates have not evolved catapult mechanisms and use countermovement jumps instead. When ground forces are large multiples of body mass, catapult jumps (as used by locusts and fleas) are much higher than the other styles of jump could be. Increasing leg mass reduces jump height, but the proximal-to-distal distribution of leg mass has only a minor effect. Longer legs make higher jumps possible and additional leg segments, such as the elongated tarsi of bushbabies and frogs, increase jump height even if overall leg length remains unchanged. The effects of muscle moment arms that change as the leg extends, and of legs designed to work over different ranges of joint angle, are investigated.

Leg design and jumping technique for humans, other vertebrates and insects

Wallaby tail tendons fail after repeated application of stresses much lower than would be needed to break them in a single pull. We show that this a fatigue phenomenon, distinct from the creep rupture that occurs after prolonged application of a constant stress. The two phenomena are disctinguished by experiments in which tensile stress is cycled at different frequencies, ranging from 1 to 50 Hz.

Fatigue rupture of wallaby tail tendons

High jumpers run at moderate speeds and set down the foot, from which they take off, well in front of the body. Long jumpers run up much faster and place the foot less far forward, with the leg at a steeper angle. A simple model, which takes into account the mechanical properties of muscle, predicts optimum take-off techniques that agree well with those used by athletes. These predictions are remarkably insensitive to the numerical values assigned to the physiological parameters.

R. M. Alexander

Papers

Energy-saving mechanisms in walking and running

A model of bipedal locomotion on compliant legs.

Leg design and jumping technique for humans, other vertebrates and insects

Fatigue rupture of wallaby tail tendons

Optimum Take-Off Techniques for High and Long Jumps