Human athletic paleobiology; using sport as a model to investigate human evolutionary adaptation

The use of sport as a conceptual framework offers unprecedented opportunities to improve our understanding of what the body does, shedding new light on our evolutionary trajectory, our capacity for adaptation, and the underlying biological mechanisms. This approach has gained traction over recent years. To date, sport has facilitated exploration not only of the evolutionary history of our species as a whole, but also of human variation and adaptation at the interindividual and intraindividual levels. At the species level, analysis of lower and upper limb biomechanics and energetics with respect to walking, running and throwing have led to significant advances in the understanding of human adaptations relative to other hominins. From an interindividual perspective, investigation of physical activity patterns and endurance running performance is affording greater understanding of evolved constraints of energy expenditure, thermoregulatory energetics, signaling theory, and morphological variation. Furthermore, ultra-endurance challenges provoke functional trade-offs, allowing new ground to be broken in the study of life history trade-offs and human adaptability. Human athletic paleobiology-the recruitment of athletes as study participants and the use of contemporary sports as a model for studying evolutionary theory-has great potential. Here, we draw from examples in the literature to provide a review of how the use of athletes as a model system is enhancing understanding of human evolutionary adaptation

A trade-off between cognitive and physical performance, with relative preservation of brain function

Debate surrounds the issue of how the large, metabolically expensive brains of Homo sapiens can be energetically afforded. At the evolutionary level, decreased investment in muscularity, adiposity and the digestive tract allow for a larger brain. Developmentally, high neo-natal adiposity and preferential distribution of resources to the brain provide an energetic buffer during times of environmental stress. Through an experimental design, we investigated the hypothesis of a trade-off involving brain and muscle at the acute level in humans. Mental performance was measured by a free-recall test, and physical performance by power output on an indoor rowing ergometer. Sixty-two male student rowers performed the two tests in isolation, and then again simultaneously. Paired samples t-tests revealed that both power output and mental performance reduced when tested together compared to in isolation (t(61) = 9.699, p < 0.001 and t(61) = 8.975, p < 0.001). Furthermore, the decrease in physical performance was greater than the decrease in mental performance (t(61) = -2.069, p = 0.043). This is the first investigation to demonstrate an acute level trade-off between these two functions, and provides support for the selfish brain hypothesis due to the relative preservation of cognitive function over physical power output. The underlying mechanism is unclear, and requires further work

Can persistence hunting signal male quality? A test considering digit ratio in endurance athletes

Various theories have been posed to explain the fitness payoffs of hunting success among hunter-gatherers. 'Having' theories refer to the acquisition of resources, and include the direct provisioning hypothesis. In contrast, 'getting' theories concern the signalling of male resourcefulness and other desirable traits, such as athleticism and intelligence, via hunting prowess. We investigated the association between androgenisation and endurance running ability as a potential signalling mechanism, whereby running prowess, vital for persistence hunting, might be used as a reliable signal of male reproductive fitness by females. Digit ratio (2D:4D) was used as a proxy for prenatal androgenisation in 439 males and 103 females, while a half marathon race (21km), representing a distance/duration comparable with that of persistence hunting, was used to assess running ability. Digit ratio was significantly and positively correlated with half-marathon time in males (right hand: r = 0.45, p<0.001; left hand: r = 0.42, p<0.001) and females (right hand: r = 0.26, p<0.01; left hand: r = 0.23, p = 0.02). Sex-interaction analysis showed that this correlation was significantly stronger in males than females, suggesting that androgenisation may have experienced stronger selective pressure from endurance running in males. As digit ratio has previously been shown to predict reproductive success, our results are consistent with the hypothesis that endurance running ability may signal reproductive potential in males, through its association with prenatal androgen exposure. However, further work is required to establish whether and how females respond to this signalling for fitness

Tandem androgenic and psychological shifts in male reproductive effort following a manipulated “win” or “loss” in a sporting competition

© 2018, The Author(s). Male-male competition is involved in inter- and intrasexual selection, with both endocrine and psychological factors presumably contributing to reproductive success in human males. We examined relationships among men’s naturally occurring testosterone, their self-perceived mate value (SPMV), self-esteem, sociosexuality, and expected likelihood of approaching attractive women versus situations leading to child involvement. We then monitored changes in these measures in male rowers (N = 38) from Cambridge, UK, following a manipulated “win” or “loss” as a result of an indoor rowing contest. Baseline results revealed that men with heightened testosterone and SPMV values typically had greater inclinations toward engaging in casual sexual relationships and a higher likelihood of approaching attractive women in a hypothetical social situation. As anticipated, both testosterone and SPMV increased following a manipulated “victory” and were associated with heightened sociosexuality, and increased expectations toward approaching attractive women versus individuals who would involve them in interacting with children after the race. SPMV and self-esteem appeared to mediate some of the effects of testosterone on post-race values. These findings are considered in the broader context of individual trade-offs between mating and parental effort and a model of the concurrent and dynamic androgenic and psychological influences contributing to male reproductive effort and success

Ultra-endurance athletic performance suggests that energetics drive human morphological thermal adaptation

Both extinct and extant hominin populations display morphological features consistent with Bergmann's and Allen's Rules. However, the functional implications of the morphologies described by these ecological laws are poorly understood. We examined this through the lens of endurance running. Previous research concerning endurance running has focused on locomotor energetic economy. We considered a less-studied dimension of functionality, thermoregulation. The performance of male ultra-marathon runners (n = 88) competing in hot and cold environments was analysed with reference to expected thermoregulatory energy costs and the optimal morphologies predicted by Bergmann's and Allen's Rules. Ecogeographical patterning supporting both principles was observed in thermally challenging environments. Finishers of hot-condition events had significantly longer legs than finishers of cold-condition events. Furthermore, hot-condition finishers had significantly longer legs than those failing to complete hot-condition events. A degree of niche-picking was evident; athletes may have tailored their event entry choices in accordance with their previous race experiences. We propose that the interaction between prolonged physical exertion and hot or cold climates may induce powerful selective pressures driving morphological adaptation. The resulting phenotypes reduce thermoregulatory energetic expenditure, allowing diversion of energy to other functional outcomes such as faster running

Short-term resource allocation during extensive athletic competition

© 2017 The Authors. American Journal of Human Biology Published by Wiley Periodicals, Inc. Objectives: Following predictions from life history theory, we sought to identify acute trade-offs between reproductive effort (as measured by psychological arousal) and somatic maintenance (via functional measures of innate immunity) during conditions of severe energetic imbalance. Methods: Sixty-six male ultramarathon runners (ages 20 to 37 years) were sampled before and after a lengthy race. Saliva and sera were collected for testosterone and immunological analyses (hemolytic complement activity and bacterial killing ability). Lean body mass was assessed by bioelectrical impedance, and libido was measured using a slideshow of arousing and neutral images. Results: Following predictions, there was a significant decrease in salivary testosterone levels (109.59 pg/mL versus 97.61 pg/mL, P <.001) and arousal scores in response to provocative images (5.40 versus 4.89, P =.001) between prerace and postrace time points. Additionally, participant bacterial killing ability (P =.035) and hemolytic complement activity (P =.021) increased between prerace and postrace. Conclusions: Decreased libido and testosterone with concomitant heightened innate immune responses suggest a shift in energetic priorities away from reproduction and toward maintenance/defense during a period of energetic stress

Patterns of energy allocation during energetic scarcity; evolutionary insights from ultra-endurance events

Exercise physiologists and evolutionary biologists share a research interest in determining patterns of energy allocation during times of acute or chronic energetic scarcity. Within sport and exercise science, this information has important implications for athlete health and performance. For evolutionary biologists, this would shed new light on our adaptive capabilities as a phenotypically plastic species. In recent years, evolutionary biologists have begun recruiting athletes as study participants and using contemporary sports as a model for studying evolution. This approach, known as human athletic palaeobiology, has identified ultra-endurance events as a valuable experimental model to investigate patterns of energy allocation during conditions of elevated energy demand, which are generally accompanied by an energy deficit. This energetic stress provokes detectable functional trade-offs in energy allocation between physiological processes. Early results from this modelsuggest thatlimited resources are preferentially allocated to processes which could be considered to confer the greatest immediate survival advantage (including immune and cognitive function). This aligns with evolutionary perspectives regarding energetic trade-offs during periods of acute and chronic energetic scarcity. Here, we discuss energy allocation patterns during periods of energetic stress as an area of shared interest between exercise physiology and evolutionary biology. We propose that, by addressing the ultimate “why” questions, namely why certain traits were selected for during the human evolutionary journey, an evolutionary perspective can complement the exercise physiology literature and provide a deeper insight of the reasons underpinning the body's physiological response to conditions of energetic stress

Energetics as a driver of human morphological thermal adaptation; evidence from female ultra-endurance athletes

Functional benefits of the morphologies described by Bergmann’s and Allen’s rules in human males have recently been reported. However, the functional implications of ecogeographical patterning in females remain poorly understood. Here, we report the findings of preliminary work analysing the association between body shape and performance in female ultramarathon runners (n = 36) competing in hot and cold environments. The body shapes differed between finishers of hot and cold races, and also between hot race finishers and non-finishers. Variability in race performance across different settings supports the notion that human phenotype is adapted to different thermal environments as ecogeographical patterns have reported previously. This report provides support for the recent hypothesis that the heightened thermal strain associated with prolonged physical activity in hot/cold environments may have driven the emergence of thermally adaptive phenotypes in our evolutionary past. These results also tentatively suggest that the relationship between morphology and performance may be stronger in female vs. male athletes. This potential sex difference is discussed with reference to the evolved unique energetic context of human female reproduction. Further work, with a larger sample size, is required to investigate the observed potential sex differences in the strength of the relationship between phenotype and performance

Fluctuating asymmetry, a marker of poor growth quality, is associated with adult male metabolic rate

ObjectivesLife history theory, a branch of evolutionary theory, predicts the existence of trade‐offs in energetic allocation between competing physiological functions. The core metabolic cost of self‐maintenance, measured by resting metabolic rate (RMR), represents a large component of human daily energy expenditure. Despite strong selective pressures for energetic frugality and high observed interindividual variation in RMR, the link between RMR and energetic allocation to life‐history traits remains understudied in humans.MaterialsIn a sample of 105 (m = 57, f = 48), we investigated the relationship between adult RMR and investment in growth quality, as measured by fluctuating asymmetry (FA).ResultsMeasurement of RMR and FA in university rowers revealed a significant positive correlation amongst males (n = 57, r = 0.344, p = 0.005, 1‐tailed; standardized 95% CI, 0.090 to 0.598). Convincing evidence for a correlation among females was not found (n = 48, r = 0.142, p = 0.169, 1‐tailed, standardized 95% CI, −0.152 to 0.435).DiscussionThe data suggest that low‐quality asymmetrical growth is associated with later‐life metabolic inefficiencies in males. Energetic investment in processes (likely concerning the stress‐response) unrelated to growth during childhood may thereby trade‐off against adult metabolic efficiency. We suggest that the presence of a relationship between RMR and FA in males but not females may be explained by the additional metabolic strain associated with larger body size and increased male muscularity, which may amplify the inefficiencies arising from low‐quality growth.</div

Alternative metabolic strategies are employed by endurance runners of different body sizes; implications for human evolution

Objective: A suite of adaptations facilitating endurance running (ER) evolved within the hominin lineage. This may have improved our ability to reach scavenging sites before competitors, or to hunt prey over long distances. Running economy (RE) is a key determinant of endurance running performance, and depends largely on the magnitude of force required to support body mass. However, numerous environmental factors influence body mass, thereby significantly affecting RE. This study tested the hypothesis that alternative metabolic strategies may have emerged to enable ER in individuals with larger body mass and poor RE. Methods: A cohort of male (n = 25) and female (n = 19) ultra-endurance runners completed submaximal and exhaustive treadmill protocols to determine RE, and V̇O2Max. Results: Body mass was positively associated with sub-maximal oxygen consumption at both LT1 (male r=0.66, p2Max (male r=0.60, p=0.001; female r=0.41, p=0.046). Additionally, sub-maximal oxygen consumption varied positively with V̇O2Max in both male (LT1 r=0.54, p=0.003; LT2 r=0.77, p<0.001) and female athletes (LT1 r=0.88, p<0.001; LT2 r=0.92, p<0.001). Conclusions: The results suggest that, while individuals with low mass and good RE can glide economically as they run, larger individuals can compensate for the negative effects their mass has on RE by increasing their capacity to consume oxygen. The elevated energy expenditure of this low-economy high-energy turnover approach to ER may bring costs associated with energy diversion away from other physiological processes, however