The Evolution of Marathon Running: Capabilities in Humans

Humans have exceptional capabilities to run long distances in hot, arid conditions. These abilities, unique among primates and rare among mammals, derive from a suite of specialised features that permit running humans to store and release energy effectively in the lower limb, help keep the body's center of mass stable and overcome the thermoregulatory challenges of long distance running. Human endurance running perfonnance capabilities compare favourably with those of other mammals and probably emerged sometime around 2 million years ago in order to help meat-eating hominids compete with other carnivores.Anthropolog

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

Endurance running and the evolution of Homo.

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. M ost research on the evolution of human locomotion has focused on walking. There are a few indications that the earliest-known hominids were bipeds 1,2 , and there is abundant fossil evidence that australopithecines habitually walked by at least 4.4 million years (Myr) ago However, although humans are comparatively poor sprinters, they also engage in a different type of running, endurance running (ER), defined as running many kilometres over extended time periods using aerobic metabolism. Although not extensively studied in non-humans, ER is unique to humans among primates, and uncommon among quadrupedal mammals other than social carnivores (such as dogs and hyenas) and migratory ungulates (such as wildebeest and horses) How well do humans run long distances? In considering human running, it helps to start from the perspective of the basic biomechanical differences that distinguish running and walking gaits in all mammals, including human bipeds. These differences are well characterized. Walking uses an 'inverted pendulum' in which the centre of mass vaults over a relatively extended leg during the stance phase, efficiently exchanging potential and kinetic energy out-of-phase with every ste

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

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

Multiple novel prostate cancer susceptibility signals identified by fine-mapping of known risk loci among Europeans

Genome-wide association studies (GWAS) have identified numerous common prostate cancer (PrCa) susceptibility loci. We have fine-mapped 64 GWAS regions known at the conclusion of the iCOGS study using large-scale genotyping and imputation in 25 723 PrCa cases and 26 274 controls of European ancestry. We detected evidence for multiple independent signals at 16 regions, 12 of which contained additional newly identified significant associations. A single signal comprising a spectrum of correlated variation was observed at 39 regions; 35 of which are now described by a novel more significantly associated lead SNP, while the originally reported variant remained as the lead SNP only in 4 regions. We also confirmed two association signals in Europeans that had been previously reported only in East-Asian GWAS. Based on statistical evidence and linkage disequilibrium (LD) structure, we have curated and narrowed down the list of the most likely candidate causal variants for each region. Functional annotation using data from ENCODE filtered for PrCa cell lines and eQTL analysis demonstrated significant enrichment for overlap with bio-features within this set. By incorporating the novel risk variants identified here alongside the refined data for existing association signals, we estimate that these loci now explain ∼38.9% of the familial relative risk of PrCa, an 8.9% improvement over the previously reported GWAS tag SNPs. This suggests that a significant fraction of the heritability of PrCa may have been hidden during the discovery phase of GWAS, in particular due to the presence of multiple independent signals within the same regio