Energy constraint and compensation: Insights from endurance athletes.

The Constrained Model of Total Energy Expenditure predicts that increased physical activity may not influence total energy expenditure, but instead, induces compensatory energetic savings in other processes. Much remains unknown, however, about concepts of energy expenditure, constraint and compensation in different populations, and it is unclear whether this model applies to endurance athletes, who expend very large amounts of energy during training and competition. Furthermore, it is well-established that some endurance athletes consciously or unconsciously fail to meet their energy requirements via adequate food intake, thus exacerbating the extent of energetic stress that they experience. Within this review we A) Describe unique characteristics of endurance athletes that render them a useful model to investigate energy constraints and compensations, B) Consider the factors that may combine to constrain activity and total energy expenditure, and C) Describe compensations that occur when activity energy expenditure is high and unmet by adequate energy intake. Our main conclusions are as follows: A) Higher activity levels, as observed in endurance athletes, may indeed increase total energy expenditure, albeit to a lesser degree than may be predicted by an additive model, given that some compensation is likely to occur; B) That while a range of factors may combine to constrain sustained high activity levels, the ability to ingest, digest, absorb and deliver sufficient calories from food to the working muscle is likely the primary determinant in most situations and C) That energetic compensation that occurs in the face of high activity expenditure may be primarily driven by low energy availability i.e., the amount of energy available for all biological processes after the demands of exercise have been met, and not by activity expenditure per se.

Human energetic stress associated with upregulation of spatial cognition.

OBJECTIVES: Evolutionary life history theory has a unique potential to shed light on human adaptive capabilities. Ultra-endurance challenges are a valuable experimental model allowing the direct testing of phenotypic plasticity via physiological trade-offs in resource allocation. This enhances our understanding of how the body prioritizes different functions when energetically stressed. However, despite the central role played by the brain in both hominin evolution and metabolic budgeting, cognitive plasticity during energetic deficit remains unstudied. MATERIALS: We considered human cognitive plasticity under conditions of energetic deficit by evaluating variability in performance in three key cognitive domains. To achieve this, cognitive performance in a sample of 48 athletes (m = 29, f = 19) was assessed before and after competing in multiday ultramarathons. RESULTS: We demonstrate that under conditions of energetic deficit, performance in tasks of spatial working memory (which assessed ability to store location information, promoting landscape navigation and facilitating resource location and calorie acquisition) increased. In contrast, psychomotor speed (reaction time) remained unchanged and episodic memory performance (ability to recall information about specific events) decreased. DISCUSSION: We propose that prioritization of spatial working memory performance during conditions of negative energy balance represents an adaptive response due to its role in facilitating calorie acquisition. We discuss these results with reference to a human evolutionary trajectory centred around encephalisation. Encephalisation affords great plasticity, facilitating rapid responses tailored to specific environmental conditions, and allowing humans to increase their capabilities as a phenotypically plastic species.

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.

A Life History Perspective on Athletes with Low Energy Availability.

The energy costs of athletic training can be substantial, and deficits arising from costs unmet by adequate energy intake, leading to a state of low energy availability, may adversely impact athlete health and performance. Life history theory is a branch of evolutionary theory that recognizes that the way the body uses energy-and responds to low energy availability-is an evolved trait. Energy is a finite resource that must be distributed throughout the body to simultaneously fuel all biological processes. When energy availability is low, insufficient energy may be available to equally support all processes. As energy used for one function cannot be used for others, energetic "trade-offs" will arise. Biological processes offering the greatest immediate survival value will be protected, even if this results in energy being diverted away from others, potentially leading to their downregulation. Athletes with low energy availability provide a useful model for anthropologists investigating the biological trade-offs that occur when energy is scarce, while the broader conceptual framework provided by life history theory may be useful to sport and exercise researchers who investigate the influence of low energy availability on athlete health and performance. The goals of this review are: (1) to describe the core tenets of life history theory; (2) consider trade-offs that might occur in athletes with low energy availability in the context of four broad biological areas: reproduction, somatic maintenance, growth, and immunity; and (3) use this evolutionary perspective to consider potential directions for future research.

Time in Nature Associated with Decreased Fatigue in UK Truck Drivers.

Heavy goods vehicle (HGV) driving is recognised as a highly hazardous occupation due to the long periods of sedentary behaviour, low levels of physical activity and unhealthy food options when working. These risk factors combine with shift work and concomitant irregular sleep patterns to increase the prevalence of fatigue. Fatigue is closely linked with stress and, subsequently, poor physiological and psychological health. In parallel, a wealth of evidence has demonstrated the health and wellbeing benefits of spending time in nature. Here, we sought to examine whether spending time in nature was associated with lower levels of fatigue, anxiety and depression in HGV drivers. 89 long-distance drivers (98.9% male, mean ± SD age: 51.0 ± 9 years, body mass index: 29.8 ± 4.7 kg/m2) participating in a wider health promotion programme reported time spent in nature (during and before the Covid-19 pandemic) and symptoms of occupational fatigue, depression and anxiety. After controlling for covariates, truck drivers who visited nature at least once a week exhibited 16% less chronic fatigue prior to the pandemic, and 23% less chronic fatigue and 20% less acute fatigue during the pandemic. No significant differences were observed for either anxiety or depression. As fatigue has a range of physical and mental health sequelae, we propose that increased exposure to natural settings may make a valuable contribution to interventions to promote the health and wellbeing of this underserved group.

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

OBJECTIVES: Life 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. MATERIALS: In 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). RESULTS: Measurement 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). DISCUSSION: The 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.

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.

Nonfreezing Cold Injuries Among Long-Distance Polar Rowers.

INTRODUCTION: Nonfreezing cold injury (NFCI) is a peripheral cold injury that occurs when the extremities are exposed to cold temperatures, at or near the freezing point, for sustained periods of time (48-96 h at temperatures of usually around 0 to 6°C with associated wind chill). Although NFCI often goes unreported and may be underdiagnosed, it is a cause of significant morbidity in those working in cold conditions, particularly those in the military. Thus, further research into the prevention, recognition, and treatment of NFCI is warranted. METHODS: The height, body weight, and body composition of 6 rowers taking part in 1 or 2 legs of the 2017 Polar Row expedition were measured. The weather conditions of the 2 legs of the journey were recorded, and symptoms relating to NFCI were documented. RESULTS: All incidences of NFCI occurred during Leg 2 of the expedition, which was colder and wetter. Of the Leg 2 rowers, those who developed NFCI had a trend toward higher pre-row body weight and body mass index and a trends toward losing more weight and body water relative to those who did not. CONCLUSIONS: The main factor contributing to the incidence of NFCI appeared to be weather; NFCI only occurred during the colder and wetter leg of the expedition. We also tentatively suggest that nutrition and dehydration may be linked to the incidence of NFCI as predisposing factors. More work, with sample sizes greater than those reported here, is required to investigate these associations to further characterize risk factors.

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.

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.

Tandem Androgenic and Psychological Shifts in Male Reproductive Effort Following a Manipulated "Win" or "Loss" in a Sporting Competition.

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.

Short-term resource allocation during extensive athletic competition.

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.