Electromyography activity across gait and incline: The impact of muscular activity on human morphology.

The study of human evolution depends upon a fair assessment of the ability of hominin individuals to gain access to necessary resources. We expect that the morphology of extant and extinct populations represents a successful locomotory system that allowed individuals to move across the environment gaining access to food, water, and mates while still maintaining excess energy to allocate to reproduction. Our assessment of locomotor morphology must then incorporate tests of fitness within realistic environments--environments that themselves vary in terrain and whose negotiation requires a variety of gait and speeds. This study assesses muscular activity (measured as the integrated signal from surface electromyography) of seven thigh and hip muscle groups during walking and running across a wide range of speeds and inclines to systematically assess the role that morphology can play in minimizing muscular activity and thus energy expenditure. Our data suggest that humans are better adapted to walking than running at any slope, as evidenced by small confidence intervals and even trends across speed and incline. We find that while increasing task intensity unsurprisingly increases muscular activity in the lower limb, individuals with longer limbs show significantly reduced activity during both walking and running, especially in the hip adductors, gluteus maximus, and hamstring muscles. People with a broader pelvis show significantly reduced activity in the hip adductor and hamstring muscles while walking.

Optimal running speed and the evolution of hominin hunting strategies.

Recent discussion of the selective pressures leading to the evolution of modern human postcranial morphology, seen as early as Homo erectus, has focused on the relative importance of walking versus running. Specifically, these conversations have centered on which gait may have been used by early Homo to acquire prey. An element of the debate is the widespread belief that quadrupeds are constrained to run at optimally efficient speeds within each gait, whereas humans are equally efficient at all running speeds. The belief in the lack of optimal running speeds in humans is based, however, on a number of early studies with experimental designs inadequate for the purpose of evaluating optimality. Here we measured the energetic cost of human running (n=9) at six different speeds for five minutes at each speed, with careful replicates and controls. We then compared the fit of linear versus curvilinear models to the data within each subject. We found that individual humans do, in fact, have speeds at which running is significantly less costly than at other speeds (i.e., an optimal running speed). In addition, we demonstrate that the use of persistence hunting methods to gain access to prey at any running speed, even the optimum, would be extremely costly energetically, more so than a persistence hunt at optimal walking speed. We argue that neither extinct nor extant hominin populations are as flexible in the chosen speeds of persistence hunting pursuits as other researchers have suggested. Variations in the efficiency of human locomotion appear to be similar to those of terrestrial quadrupeds.

The effects of body proportions on thermoregulation: an experimental assessment of Allen's rule.

Numerous studies have discussed the influence of thermoregulation on hominin body shape concluding, in accordance with Allen's rule, that the presence of relatively short limbs on both extant as well as extinct hominin populations offers an advantage for survival in cold climates by reducing the limb's surface area to volume ratio. Moreover, it has been suggested that shortening the distal limb segment compared to the proximal limb segment may play a larger role in thermoregulation due to a greater relative surface area of the shank. If longer limbs result in greater heat dissipation, we should see higher resting metabolic rates (RMR) in longer-limbed individuals when temperature conditions fall, since the resting rate will need to replace the lost heat. We collected resting oxygen consumption on volunteer human subjects to assess the correlation between RMR and lower limb length in human subjects, as well as to reexamine the prediction that shortening the distal segment would have a larger effect on heat loss and, thus, RMR than the shortening of the proximal segment. Total lower limb length exhibits a statistically significant relationship with resting metabolic rate (p<0.001; R(2)=0.794). While this supports the hypothesis that as limb length increases, resting metabolic rate increases, it also appears that thigh length, rather than the length of the shank, drives this relationship. The results of the present study confirm the widely-held expectation of Allen's rule, that short limbs reduce the metabolic cost of maintaining body temperature, while long limbs result in greater heat dissipation regardless of the effect of mass. The present results suggest that the shorter limbs of Neandertals, despite being energetically disadvantageous while walking, would indeed have been advantageous for thermoregulation.

The evolution of human running: effects of changes in lower-limb length on locomotor economy.

Previous studies have differed in expectations about whether long limbs should increase or decrease the energetic cost of locomotion. It has recently been shown that relatively longer lower limbs (relative to body mass) reduce the energetic cost of human walking. Here we report on whether a relationship exists between limb length and cost of human running. Subjects whose measured lower-limb lengths were relatively long or short for their mass (as judged by deviations from predicted values based on a regression of lower-limb length on body mass) were selected. Eighteen human subjects rested in a seated position and ran on a treadmill at 2.68 ms(-1) while their expired gases were collected and analyzed; stride length was determined from videotapes. We found significant negative relationships between relative lower-limb length and two measures of cost. The partial correlation between net cost of transport and lower-limb length controlling for body mass was r=-0.69 (p=0.002). The partial correlation between the gross cost of locomotion at 2.68 ms(-1) and lower-limb length controlling for body mass was r=-0.61 (p=0.009). Thus, subjects with relatively longer lower limbs tend to have lower locomotor costs than those with relatively shorter lower limbs, similar to the results found for human walking. Contrary to general expectation, a linear relationship between stride length and lower-limb length was not found.

Energetics in Homo erectus and other early hominins: the consequences of increased lower-limb length.

Previous studies of daily energy expenditure (DEE) in hominin fossils have estimated locomotor costs using a formula that was based on six species, all 18 kg or less in mass, including no primates, and that has a number of other problems when applied in an ecological context. It is well established that the energetic cost of human walking is lower than that of representative mammals, particularly for individuals with long lower limbs. The current study reevaluates the daily energy expenditures of a variety of hominin species using more appropriate approaches to estimating locomotor costs. To estimate DEE for primates, I relied on published data on body mass, day range, and the percentage of time spent in various activities. Based on those data, I calculated a value for nonlocomotor DEE. I then used a variant of a method that I have suggested elsewhere to calculate the daily cost due to locomotion (DEEL) and summed the two to calculate total DEE. The more up-to-date methods for calculating the cost of travel result in lower estimates of this aspect of the energy budget than seen in previous studies. Values obtained here for DEE in various representatives of Australopithecus are lower than reported previously by around 200 kcal/day. Taking into account the greater economy of human walking, particularly the effect of the longer lower limbs found in many later Homo species, also results in lowered estimates of DEE. Elongation of the lower limbs in H. erectus reduced relative travel costs nearly 50% in comparison to A.L. 288-1 (A. afarensis). The present method for calculating DEE indicates that female H. erectus DEE was 84% greater than that of female Australopithecus; this disparity is even larger than that suggested by previous workers.

The application to bipeds of a geometric model of lower-limb-segment inertial properties.

Drawing inferences about locomotor energetics from limb morphology, especially in regard to small differences between individuals, depends critically on valid estimates of lower-limb inertial properties. While there are numerous options for such estimations in the literature, geometric models that involve simple measures and straightforward mathematics combined with the ability to capture individual variation are rare. In this research, we apply a method, originally developed for quadrupeds, that models limb segments as elliptical columns. When the elliptical model is applied to bipeds, it provides a means of estimating limb-segment inertial properties accurately enough to test differences between individuals of similar stature and mass, but with variation in mass distribution and limb length. We test the method against commonly used equations and are able to show the validity of the method for thigh and shank segments.

Froude number corrections in anthropological studies.

The Froude number has been widely used in anthropology to adjust for size differences when comparing gait parameters or other nonmorphological locomotor variables (such as optimal walking speed or speed at gait transitions) among humans, nonhuman primates, and fossil hominins. However, the dynamic similarity hypothesis, which is the theoretical basis for Froude number corrections, was originally developed and tested at much higher taxonomic levels, for which the ranges of variation are much greater than in the intraspecific or intrageneric comparisons typical of anthropological studies. Here we present new experimental data on optimal walking speed and the mass-specific cost of transport at that speed from 19 adult humans walking on a treadmill, and evaluate the predictive power of the dynamic similarity hypothesis in this sample. Contrary to the predictions of the dynamic similarity hypothesis, we found that the mass-specific cost of transport at experimentally measured optimal walking speed and Froude number were not equal across individuals, but retained a significant correlation with body mass. Overall, the effect of lower limb length on optimal walking speed was weak. These results suggest that the Froude number may not be an effective way for anthropologists to correct for size differences across individuals, but more studies are needed. We suggest that researchers first determine whether geometric similarity characterizes their data before making inferences based on the dynamic similarity hypothesis, and then check the consistency of their results with and without Froude number corrections before drawing any firm conclusions.

The effect of lower limb length on the energetic cost of locomotion: implications for fossil hominins.

The consequences of the relatively short lower limbs characteristic of AL 288-1 have been widely discussed, as have the causes and consequences of the short limbs of Neanderthals. Previous studies of the effect of limb length on the energetic cost of locomotion have reported no relationship; however, limb length could have accounted for as much as 19% of the variation in cost and gone undetected (Steudel and Beattie, 1995; Steudel, 1994, 1996). Kramer (1999) and Kramer and Eck (2000) have recently used a theoretical model to predict the effect of the shorter limbs of early hominids, concluding that the shorter limbs may actually have been energetically advantageous. Here, we took an experimental approach. Twenty-one human subjects, of varying limb lengths, walked on a treadmill at 2.6, 2.8, 3.0 and 3.2m.p.h., while their expired gases were analyzed. The subjects walked for 12 minutes at each speed and their rates of oxygen consumption (VO2) over four minutes were averaged to estimate VO2. We also measured each subject's height, weight and lower limb length. Lean body mass and % fat were determined using dual-energy x-ray absorptiometry. ANCOVA with total VO2 at either speed as the dependent variable and total lean mass, % fat and lower limb length as covariates resulted in all three covariates having a significant positive effect on VO2 at p<0.01. Subjects with relatively longer lower limbs had lower locomotor costs. Thus the short lower limbs characteristic of some hominid taxa would have resulted in more costly locomotion, barring some physiological anomaly. The magnitude of this effect is substantial; Neanderthals are estimated to have had locomotor costs 30% greater than those of contemporary anatomically modern humans. By contrast the increase in lower limb length seen in H. erectus would have mitigated the increase in locomotor costs produced by the increase in body size.

The energetic cost of locomotion: humans and primates compared to generalized endotherms.

A wide range of selective pressures have been advanced as possible causes for the adoption of bipedalism in the hominin lineage. One suggestion has been that because modern human walking is relatively efficient compared to that of a typical quadruped, the ancestral quadruped may have reaped an energetic advantage when it walked on two legs. While it has become clear that human walking is relatively efficient and human running inefficient compared to "generalized endotherms", workers differ in their opinion of how the cost of human bipedal locomotion compares to that of a generalized primate walking quadrupedally. One view is that human walking is particularly efficient in comparison to other primates. The present study addresses this by comparing the cost of human walking and running to that of the eight primate species for which data are available and by comparing cost in primates to that of a "generalized endotherm". There is no evidence that primate locomotion is more costly than that of a generalized endotherm, although more data on adult Old World monkeys and apes would be useful. Further, human locomotion does not appear to be particularly efficient relative to that of other primates.