Integration of the Head and Forelimb in Bipedal Hominids

Integration, a fundamental property of organisms, occurs via multiple mechanisms and for diverse reasons. Although there has been substantial work on the genetic and epigenetic mechanisms by which developmental integration occurs, we have less of an understanding of the evolutionary relationships between functional and developmental integration. In this respect, human evolution provides an interesting test case. In quadrupedal mammals, there is considerable functional integration among and between the limbs, but little functional integration between the limbs and the skull. The evolution of bipedalism in hominids, however, provided new opportunities for novel forms of integration by emancipating the forelimbs from any major role in locomotion. Here we consider how the forelimb and head become increasingly integrated in the genus Homo because of the biomechanical challenges of running. While the arm and the head interact little during walking, we have found that, during running, the stance side arm acts as a counterbalance to the head, stabilizing it against impulsive pitching forces generated by the heel strike transient. Moreover, the functional properties of this linkage may have driven several developmental changes in the proportions of the arm and the anatomy of the shoulder girdle during human evolution. Thus, evolutionary changes in arm and head morphology during human evolution may be more integrated than previously considered.AnthropologyHuman Evolutionary Biolog

A new look at the Dynamic Similarity Hypothesis: the importance of swing phase

Summary The Dynamic Similarity Hypothesis (DSH) suggests that when animals of different size walk at similar Froude numbers (equal ratios of inertial and gravitational forces) they will use similar size-corrected gaits. This application of similarity theory to animal biomechanics has contributed to fundamental insights in the mechanics and evolution of a diverse set of locomotor systems. However, despite its popularity, many mammals fail to walk with dynamically similar stride lengths, a key element of gait that determines spontaneous speed and energy costs. Here, we show that the applicability of the DSH is dependent on the inertial forces examined. In general, the inertial forces are thought to be the centripetal force of the inverted pendulum model of stance phase, determined by the length of the limb. If instead we model inertial forces as the centripetal force of the limb acting as a suspended pendulum during swing phase (determined by limb center of mass position), the DSH for stride length variation is fully supported. Thus, the DSH shows that inter-specific differences in spatial kinematics are tied to the evolution of limb mass distribution patterns. Selection may act on morphology to produce a given stride length, or alternatively, stride length may be a “spandrel” of selection acting on limb mass distribution

The Evolution of Endurance Running and the Tyranny of Ethnography: A Reply to Pickering and Bunn (2007)

Endurance running (ER) poses a conundrum for paleoanthropologists. As summarized in Bramble and Lieberman (2004), human ER capabilities, which are unique among primates, either match or exceed those of mammals adapted for running (cursors), including dogs and equids. Because many of the biomechanical and physiological challenges of human ER are so different from those of walking, we can conclude that human ER capabilities did not arise merely as a by-product of selection for walking. Instead, the available evidence suggests that an array of features that improve ER performance were selected in the genus Homo, and they were probably present to some extent by the appearance of Homo erectus at approximately 1.9 Ma. Yet, ER is no longer necessary for human survival, even among extant foragers such as the Hadza or the Bushmen. Thus, a puzzle that paleoanthropologists must solve is identifying what past behaviors - behaviors no longer common among living foragers - favored the evolution of ER. Pickering and Bunn’s (2007) criticisms of the ER hypothesis center on two issues: first, that early Homo lacked the tracking abilities necessary for successful pursuit hunts, and second, that recent ethnographic evidence suggests that modern hunter-gatherers rarely use ER to either hunt or scavenge. These arguments are based on a presumptive link between modern human- like cognition and tracking abilities, as well as the notion that the modern ethnographic record provides an adequate reflection of past behaviors. Both of these assumptions are flawed. Although tracking is complex, there is little evidence to suggest that early hominids lacked the tracking abilities of much less encephalized carnivores. Additionally, as noted by Marlowe (2005), comparatively recent inventions, such as the bow and arrow, the spear thrower, nets, and even the spear point, fundamentally altered how humans hunt and scavenge. A strict reliance on the recent ethnographic record, what Wobst (1978) termed the ‘‘tyranny of ethnography,’’ is a fundamentally problematic way of testing hypotheses of past hunting behavior. Even so, a review of the ethnographic evidence reveals errors in Pickering and Bunn’s contentions.Anthropolog

Control and Function of Arm Swing in Human Walking and Running

We investigated the control and function of arm swing in human walking and running to test the hypothesis that the arms act as passive mass dampers powered by movement of the lower body, rather than being actively driven by the shoulder muscles. We measured locomotor cost, deltoid muscle activity and kinematics in 10 healthy adult subjects while walking and running on a treadmill in three experimental conditions: control; no arms (arms folded across the chest); and arm weights (weights worn at the elbow). Decreasing and increasing the moment of inertia of the upper body in no arms and arm weights conditions, respectively, had corresponding effects on head yaw and on the phase differences between shoulder and pelvis rotation, consistent with the view of arms as mass dampers. Angular acceleration of the shoulders and arm increased with torsion of the trunk and shoulder, respectively, but angular acceleration of the shoulders was not inversely related to angular acceleration of the pelvis or arm. Restricting arm swing in no arms trials had no effect on locomotor cost. Anterior and posterior portions of the deltoid contracted simultaneously rather than firing alternately to drive the arm. These results support a passive arm swing hypothesis for upper body movement during human walking and running, in which the trunk and shoulders act primarily as elastic linkages between the pelvis, shoulder girdle and arms, the arms act as passive mass dampers which reduce torso and head rotation, and upper body movement is primarily powered by lower body movement.AnthropologyHuman Evolutionary Biolog

Is arm swing active or passive during human walking and running?

Humans habitually swing their arms in phase with the contralateral leg during walking and running. This arm motion is generally thought to counteract the torque about the body’s vertical axis (i.e. yaw moment) that is generated by the legs as they swing with each step. Thus it has been argued that the motion of the arms is a tuned, habitual, active response that is critical for maintaining stability during human locomotion, especially running. In this study, we investigated whether arm swing is in fact an active behavior, or is instead a passive response that follows solely as a consequence of our anatomical design. Human subjects walked and ran on a treadmill under different arm- and leg-weighting conditions, and without armswing, while kinematic and surface EMG data were recorded. A modeling study was also performed to determine the inherent effect of leg swing on arm movement in a human-like biped. Results of both studies suggest that arm swing is largely a passive response, and is not entirely an active, tuned behavior. Arm swing may therefore be an emergent property of human bipedalism, with the arms acting largely as passive damping mechanisms that decrease whole-body yawing.AnthropologyHuman Evolutionary Biolog

The Human Gluteus Maximus and its Role in Running

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 stanceside 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.Anthropolog

Hunter-gatherer energetics and human obesity

Western lifestyles differ markedly from those of our hunter-gatherer ancestors, and these differences in diet and activity level are often implicated in the global obesity pandemic. However, few physiological data for hunter-gatherer populations are available to test these models of obesity. In this study, we used the doubly-labeled water method to measure total daily energy expenditure (kCal/day) in Hadza hunter-gatherers to test whether foragers expend more energy each day than their Western counterparts. As expected, physical activity level, PAL, was greater among Hadza foragers than among Westerners. Nonetheless, average daily energy expenditure of traditional Hadza foragers was no different than that of Westerners after controlling for body size. The metabolic cost of walking (kcal kg(-1) m(-1)) and resting (kcal kg(-1) s(-1)) were also similar among Hadza and Western groups. The similarity in metabolic rates across a broad range of cultures challenges current models of obesity suggesting that Western lifestyles lead to decreased energy expenditure. We hypothesize that human daily energy expenditure may be an evolved physiological trait largely independent of cultural differences

Primate Energy eExpenditure and Life History

Humans and other primates are distinct among placental mammals in having exceptionally slow rates of growth, reproduction, and aging. Primates’ slow life history schedules are generally thought to reflect an evolved strategy of allocating energy away from growth and reproduction and toward somatic investment, particularly to the development and maintenance of large brains. Here we examine an alternative explanation: that primates’ slow life histories reflect low total energy expenditure (TEE) (kilocalories per day) relative to other placental mammals. We compared doubly labeled water measurements of TEE among 17 primate species with similar measures for other placental mammals. We found that primates use remarkably little energy each day, expending on average only 50% of the energy expected for a placental mammal of similar mass. Such large differences in TEE are not easily explained by differences in physical activity, and instead appear to reflect systemic metabolic adaptation for low energy expenditures in primates. Indeed, comparisons of wild and captive primate populations indicate similar levels of energy expenditure. Broad interspecific comparisons of growth, reproduction, and maximum life span indicate that primates’ slow metabolic rates contribute to their characteristically slow life histories

Exercise-induced endocannabinoid signaling is modulated by intensity

Abstract Endocannabinoids (eCB) are endogenous ligands for cannabinoid receptors that are densely expressed in brain networks responsible for reward. Recent work shows that exercise activates the eCB system in humans and other mammals, suggesting eCBs are partly responsible for the reported improvements in mood and affect following aerobic exercise in humans. However, exercise-induced psychological changes reported by runners are known to be dependent on exercise intensity, suggesting that any underlying molecular mechanism should also change with varying levels of exercise intensity. Here, we examine circulating levels of eCBs following aerobic exercise (treadmill running) in recreationally fit human runners at four different intensities. We show that eCB signaling is indeed intensity dependent, with significant changes in circulating eCBs observed following moderate intensities only (very high and very low intensity exercises do not significantly alter circulating eCB levels). Our results are consistent with intensity-dependent psychological state changes with exercise and therefore support the hypothesis that eCB activity is related to neurobiological effects of exercise. Thus, future studies examining the role of exercise-induced eCB signaling on neurobiology or physiology must take exercise intensity into account