Electromyography activity across gait and incline: The impact of muscular activity on human morphology (original) (raw)
Related papers
The musculoskeletal system of humans is not tuned to maximize the economy of locomotion
Proceedings of the National Academy of Sciences, 2011
Humans are known to have energetically optimal walking and running speeds at which the cost to travel a given distance is minimized. We hypothesized that “optimal” walking and running speeds would also exist at the level of individual locomotor muscles. Additionally, because humans are 60–70% more economical when they walk than when they run, we predicted that the different muscles would exhibit a greater degree of tuning to the energetically optimal speed during walking than during running. To test these hypotheses, we used electromyography to measure the activity of 13 muscles of the back and legs over a range of walking and running speeds in human subjects and calculated the cumulative activity required from each muscle to traverse a kilometer. We found that activity of each of these muscles was minimized at specific walking and running speeds but the different muscles were not tuned to a particular speed in either gait. Although humans are clearly highly specialized for terrestr...
The evolution of human running: Effects of changes in lower-limb length on locomotor economy
Journal of Human Evolution, 2007
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 for 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 m s À1 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 ¼ À0.69 ( p ¼ 0.002). The partial correlation between the gross cost of locomotion at 2.68 m s À1 and lower-limb length controlling for body mass was r ¼ À0.61 ( p ¼ 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.
Journal of Human Evolution, 2004
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 . and 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.
The Locomotor Energetics of Extinct Hominids
Bipedality is the defining characteristic of Hominidae and, consequently, the energetics of human bipedal locomotion has been of interest to anthropological researchers for many years. Scenarios of hominid evolution have been created to explain the rise of bipedality as predicated on such diverse factors as selection for energy efficiency (Foley, 1992), maximal thermal energy transfer (Wheeler, 1991a,b) and the use of display to maintain group cohesion (Jablonski and Chaplin, 1993). Without regard for which selective forces were at work, bipedality has been seen as a unitary adaptation, rather than as a general one with a variety of possible styles. Just as there are myriad forms of obligate and facultative quadrupedality, however, so too can there be different forms of bipedality. The style of bipedality that served the early hominids of the late Miocene was almost surely different from that of late Pliocene Homo. The fossil record has begun to reveal that hominids can possess at least two body configurations (Jungers, 1991): relatively short-legged and long-legged versions. The former is characteristic of the early hominids, the australopithecines, while the latter is limited to Homo. Australopithecines are frequently seen as a transitional group, not fully modern in their form of bipedality and, consequently, energetically inefficient. Despite warnings from a few biomechanists (Witte et al., 1991), anthropologists have often equated short legs with inefficiency [an inherent bias of Steudel (1994), Webb (1996), Jungers (1982, 1991) and McHenry (1991b), to name a few]. The implicit logic is that short legs imply short stride lengths and that short strides require a higher cadence to maintain a particular velocity (Jungers, 1991). Unfortunately for anthropological analyses that rely on this logic, energetic expenditure is not solely a matter of the number of strides taken, but also of the energy required to take a single stride. This latter variable is dependent on the configuration of the locomotor anatomy, and shorter limb length implies, among other things, a smaller mass moment of inertia, which decreases energy expenditure. Consequently, it is reasonable to question the assumption that the short-legged
Comparative energetics of mammalian locomotion: humans are not different
Debates about the evolution of human bipedality sometimes include discussion on the energy costs of terrestrial locomotion of extinct and extant hominins. However, comparative analyses of hominin transport costs conducted to date have been limited and potentially misinforming, in part because they fail to consider phylogenetic history. In the present study, we compare the measured costs of pedestrian locomotion in humans and the estimated costs for Australopithecus afarensis (an early bipedal hominin), to a database of locomotory costs for mammals. Using data for 81 species of mammal, we demonstrate significant phylogenetic signal in both log-transformed body mass (logMass) and log-transformed net cost of transport (logNCOT), but no phylogenetic signal in residuals of the relationship between logNCOT and logMass. We then used this relationship to generate a prediction line for NCOT based on body mass, and compared this prediction with published measured data for NCOT of running and walking in humans, and estimated NCOT of walking in A. afarensis. The cost of human walking was 25% lower than predicted, while the cost of running was 27% higher. The cost of A. afarensis walking was 32% lower than predicted. However, all of these data points fall within the 95% prediction interval for mammals, indicating that they are not significantly lower or higher than predicted for other mammals of similar mass. Moreover, the difference between humans and our closest living relative the common chimpanzee is comparable to differences between other similarly closely related species. We therefore conclude that there is no evidence from metabolic data that humans, or A. afarensis, have/had a reduced energy cost of pedestrian locomotion compared to other mammals in general.
Locomotor energetics and leg length in hominid bipedality
Journal of Human Evolution, 2000
Locomotor energetics and leg length in hominid bipedality Because bipedality is the quintessential characteristic of Hominidae, researchers have compared ancient forms of bipedality with modern human gait since the first clear evidence of bipedal australopithecines was unearthed over 70 years ago. Several researchers have suggested that the australopithecine form of bipedality was transitional between the quadrupedality of the African apes and modern human bipedality and, consequently, inefficient. Other researchers have maintained that australopithecine bipedality was identical to that of Homo. But is it reasonable to require that all forms of hominid bipedality must be the same in order to be optimized? Most attempts to evaluate the locomotor effectiveness of the australopithecines have, unfortunately, assumed that the locomotor anatomy of modern humans is the exemplar of consummate bipedality. Modern human anatomy is, however, the product of selective pressures present in the particular milieu in which Homo arose and it is not necessarily the only, or even the most efficient, bipedal solution possible. In this report, we investigate the locomotion of Australopithecus afarensis, as represented by AL 288-1, using standard mechanical analyses. The osteological anatomy of AL 288-1 and movement profiles derived from modern humans are applied to a dynamic model of a biped, which predicts the mechanical power required by AL 288-1 to walk at various velocities. This same procedure is used with the anatomy of a composite modern woman and a comparison made. We find that AL 288-1 expends less energy than the composite woman when locomoting at walking speeds. This energetic advantage comes, however, at a price: the preferred transition speed (from a walk to a run) of AL 288-1 was lower than that of the composite woman. Consequently, the maximum daily range of AL 288-1 may well have been substantially smaller than that of modern people. The locomotor anatomy of A. afarensis may have been optimized for a particular ecological niche-slow speed foraging-and is neither compromised nor transitional.
Biomechanical and physiological aspects of legged locomotion in humans
European Journal of Applied Physiology, 2003
Walking and running, the two basic gaits used by man, are very complex movements. They can, however, be described using two simple models: an inverted pendulum and a spring. Muscles must contract at each step to move the body segments in the proper sequence but the work done is, in part, relieved by the interplay of mechanical energies, potential and kinetic in walking, and elastic in running. This explains why there is an optimal speed of walking (minimal metabolic cost of about 2 J.kg–1·m–1 at about 1.11 m.s–1) and why the cost of running is constant and independent of speed (about 4 J.kg–1.m–1). Historically, the mechanical work of locomotion has been divided into external and internal work. The former is the work done to raise and accelerate the body centre of mass (m) within the environment, the latter is the work done to accelerate the body segments with respect to the centre of m. The total work has been calculated, somewhat arbitrarily, as the sum of the two. While the changes of potential and kinetic energies can be accurately measured, the contribution of the elastic energy cannot easily be assessed, nor can the true work performed by the muscles. Many factors can affect the work of locomotion - the gradient of the terrain, body size (height and body m), and gravity. The partitioning of positive and negative work and their different efficiencies explain why the most economical gradient is about –10% (1.1 J.kg–1.m–1 at 1.3 m.s–1 for walking, and 3.1 J.kg–1.m–1 at between 3 and 4 m·s–1 for running). The mechanics of walking of children, pigmies and dwarfs, in particular the recovery of energy at each step, is not different from that of taller (normal sized) individuals when the speed is expressed in dynamically equivalent terms (Froude number). An extra load, external or internal (obesity) affects internal and external work according to the distribution of the added m. Different gravitational environments determine the optimal speed of walking and the speed of transition from walking to running: at more than 1 g it is easier to walk than to run, and it is the opposite at less than 1 g. Passive aids, such as skis or skates, allow an increase in the speed of progression, but the mechanics of the locomotion cannot be simply described using the models for walking and running because step frequency, the proportion of step duration during which the foot is in contact with the ground, the position of the limbs, the force exerted on the ground and the time of its application are all different.
Journal of Human Evolution, 2003
We have previously reported that predictive dynamic modeling suggests that the 'bent-hip, bent-knee' gait, which some attribute to Australopithecus afarensis AL-288-1, would have been much more expensive in mechanical terms for this hominid than an upright gait. Normal walking by modern adult humans owes much of its efficiency to conservation of energy by transformation between its potential and kinetic states. These findings suggest the question if, and to what extent, energy transformation exists in 'bent-hip, bent-knee' gait.