Total daily energy expenditure and elevated water turnover in a small-scale semi-nomadic pastoralist society from Northern Kenya.

BACKGROUND: Pastoralists live in challenging environments, which may be accompanied by unique activity, energy, and water requirements. AIM: Few studies have examined whether the demands of pastoralism contribute to differences in total energy expenditure (TEE) and water turnover (WT) compared to other lifestyles. SUBJECTS AND METHODS: Accelerometer-derived physical activity, doubly labelled water-derived TEE and WT, and anthropometric data were collected for 34 semi-nomadic Daasanach adults from three northern Kenyan communities with different levels of pastoralist activity. Daasanach TEEs and WTs were compared to those of other small-scale and industrialised populations. RESULTS: When modelled as a function of fat-free-mass, fat-mass, age, and sex, TEE did not differ between Daasanach communities. Daasanach TEE (1564-4172 kcal/day) was not significantly correlated with activity and 91% of TEEs were within the range expected for individuals from comparison populations. Mean WT did not differ between Daasanach communities; Daasanach absolute (7.54 litres/day men; 7.46 litres/day women), mass-adjusted, and TEE-adjusted WT was higher than most populations worldwide. CONCLUSIONS: The similar mass-adjusted TEE of Daasanach and industrialised populations supports the hypothesis that habitual TEE is constrained, with physically demanding lifestyles necessitating trade-offs in energy allocation. Elevated WT in the absence of elevated TEE likely reflects a demanding active lifestyle in a hot, arid climate.

Total Energy Expenditure and Nutritional Intake in Continuous Multiday Ultramarathon Events.

Continuous multiday ultramarathon competitions are increasingly popular and impose extreme energetic and nutritional demands on competitors. However, few data have been published on energy expenditure during these events. Here, we report doubly labeled water-derived measures of total energy expenditure (in kilocalories per day) and estimated physical activity level (PAL: total energy expenditure/basal metabolic rate) collected from five elite and subelite finishers (four males and one female, age 34.6 ± 4.9 years)-and nutritional intake data from the winner-of the Cocodona 250, a ∼402-km race in Arizona, and from a fastest-known-time record (one male, age 30 years) on the ∼1,315-km Arizona Trail. PAL during these events exceeded four times basal metabolic rate (Cocodona range: 4.34-6.94; Arizona Trail: 5.63). Combining the results with other doubly labeled water-derived total energy expenditure data from ultraendurance events show a strong inverse relationship between event duration and PAL (r2 = .68, p < .0001). Cocodona race duration was inversely, though not significantly, associated with PAL (r2 = .70, p = .08). Water turnover varied widely between athletes and was not explained by PAL or body mass. The Cocodona race winner met ∼53% of energy demand via dietary intake, 85.6% of which was carbohydrate, while ∼47% of energy demand was met via catabolism of body energy stores. Together, these results illustrate the energetic deficits incurred during competitive continuous multiday ultramarathon efforts and implicate macronutrient absorption and/or storage as key factors in ultramarathon performance.

Quantifying physical activity energy expenditure based on doubly labelled water and basal metabolism calorimetry: what are we actually measuring?

PURPOSE OF REVIEW: Physical activity impacts energy balance because of its contribution to total energy expenditure. Measuring physical activity energy expenditure (PAEE) is often performed by subtracting the estimated 24 h expenditure on basal metabolism (called basal energy expenditure or BEE) from the total energy expenditure (TEE) measured by doubly labelled water minus an estimate of the thermic effect of food (TEF). Alternatively it can be measured as the ratio of TEE/BEE, which is commonly called the physical activity level (PAL). RECENT FINDINGS: PAEE and PAL are widely used in the literature but their shortcomings are seldom addressed. In this review, we outline some of the issues with their use. SUMMARY: TEE and BEE are both measured with error. The estimate of PAEE by difference magnifies these errors and consequently the precision of estimated PAEE is about 3× worse than TEE and 25-35× worse than BEE. A second problem is that the component called PAEE is actually any component of TEE that is not BEE. We highlight how the diurnal variation of BEE, thermoregulatory expenditure and elevations of RMR because of stress will all be part of what is called PAEE and will contribute to a disconnect between what is measured and what energy expenditure is a consequence of physical activity. We emphasize caution should be exerted when interpreting these measurements of PAEE and PAL.

Energetic costs of testosterone in two subsistence populations.

OBJECTIVE: Testosterone plays a role in mediating energetic trade-offs between growth, maintenance, and reproduction. Investments in a high testosterone phenotype trade-off against other functions, particularly survival-enhancing immune function and cellular repair; thus only individuals in good condition can maintain both a high testosterone phenotype and somatic maintenance. While these effects are observed in experimental manipulations, they are difficult to demonstrate in free-living animals, particularly in humans. We hypothesize that individuals with higher testosterone will have higher energetic expenditures than those with lower testosterone. METHODS: Total energetic expenditure (TEE) was quantified using doubly labeled water in n = 40 Tsimane forager-horticulturalists (50% male, 18-87 years) and n = 11 Hadza hunter-gatherers (100% male, 18-65 years), two populations living subsistence lifestyles, high levels of physical activity, and high infectious burden. Urinary testosterone, TEE, body composition, and physical activity were measured to assess potential physical and behavioral costs associated with a high testosterone phenotype. RESULTS: Endogenous male testosterone was significantly associated with energetic expenditure, controlling for fat free mass; a one standard deviation increase in testosterone is associated with the expenditure of an additional 96-240 calories per day. DISCUSSION: These results suggest that a high testosterone phenotype, while beneficial for male reproduction, is also energetically expensive and likely only possible to maintain in healthy males in robust condition.

The fire of evolution: energy expenditure and ecology in primates and other endotherms.

Total energy expenditure (TEE) represents the total energy allocated to growth, reproduction and body maintenance, as well as the energy expended on physical activity. Early experimental work in animal energetics focused on the costs of specific tasks (basal metabolic rate, locomotion, reproduction), while determination of TEE was limited to estimates from activity budgets or measurements of subjects confined to metabolic chambers. Advances in recent decades have enabled measures of TEE in free-living animals, challenging traditional additive approaches to understanding animal energy budgets. Variation in lifestyle and activity level can impact individuals' TEE on short time scales, but interspecific differences in TEE are largely shaped by evolution. Here, we review work on energy expenditure across the animal kingdom, with a particular focus on endotherms, and examine recent advances in primate energetics. Relative to other placental mammals, primates have low TEE, which may drive their slow pace of life and be an evolved response to the challenges presented by their ecologies and environments. TEE variation among hominoid primates appears to reflect adaptive shifts in energy throughput and allocation in response to ecological pressures. As the taxonomic breadth and depth of TEE data expand, we will be able to test additional hypotheses about how energy budgets are shaped by environmental pressures and explore the more proximal mechanisms that drive intra-specific variation in energy expenditure.

Variability in energy expenditure is much greater in males than females.

In mammals, trait variation is often reported to be greater among males than females. However, to date, mainly only morphological traits have been studied. Energy expenditure represents the metabolic costs of multiple physical, physiological, and behavioral traits. Energy expenditure could exhibit particularly high greater male variation through a cumulative effect if those traits mostly exhibit greater male variation, or a lack of greater male variation if many of them do not. Sex differences in energy expenditure variation have been little explored. We analyzed a large database on energy expenditure in adult humans (1494 males and 3108 females) to investigate whether humans have evolved sex differences in the degree of interindividual variation in energy expenditure. We found that, even when statistically comparing males and females of the same age, height, and body composition, there is much more variation in total, activity, and basal energy expenditure among males. However, with aging, variation in total energy expenditure decreases, and because this happens more rapidly in males, the magnitude of greater male variation, though still large, is attenuated in older age groups. Considerably greater male variation in both total and activity energy expenditure could be explained by greater male variation in levels of daily activity. The considerably greater male variation in basal energy expenditure is remarkable and may be explained, at least in part, by greater male variation in the size of energy-demanding organs. If energy expenditure is a trait that is of indirect interest to females when choosing a sexual partner, this would suggest that energy expenditure is under sexual selection. However, we present a novel energetics model demonstrating that it is also possible that females have been under stabilizing selection pressure for an intermediate basal energy expenditure to maximize energy available for reproduction.

Human total, basal and activity energy expenditures are independent of ambient environmental temperature.

Lower ambient temperature (Ta) requires greater energy expenditure to sustain body temperature. However, effects of Ta on human energetics may be buffered by environmental modification and behavioral compensation. We used the IAEA DLW database for adults in the USA (n = 3213) to determine the effect of Ta (-10 to +30°C) on TEE, basal (BEE) and activity energy expenditure (AEE) and physical activity level (PAL). There were no significant relationships (p > 0.05) between maximum, minimum and average Ta and TEE, BEE, AEE and PAL. After adjustment for fat-free mass, fat mass and age, statistically significant (p < 0.01) relationships between TEE, BEE and Ta emerged in females but the effect sizes were not biologically meaningful. Temperatures inside buildings are regulated at 18-25°C independent of latitude. Hence, adults in the US modify their environments to keep TEE constant across a wide range of external ambient temperatures.

Balancing the scales: Preliminary investigation of total energy expenditure and daily metabolizable energy intake in Matschie's tree kangaroo (Dendrolagus matschiei).

Matschie's tree kangaroo (Dendrolagus matschiei) is an endangered arboreal marsupial native to Papua New Guinea. Detailed field studies of its behavior and ecology are scarce due largely to its occupation of remote cloud forests and cryptic nature. Although this species has been in human care since the 1950s, much of its biology is still unknown. The current ex situ population is not sustainable due to health and reproductive problems, believed to stem largely from issues with diet and obesity. To better assess potential discrepancies between energy requirements and energy intake, we sought to 1) quantify total energy expenditure (TEE) of two zoo-housed Matschie's tree kangaroos (body mass = 9.0-9.7 kg) on a diet composed largely of leafy browse; 2) quantify food and macronutrient intake, apparent dry matter macronutrient digestibility, and metabolizable energy (ME) intake over a 14-month period; and 3) test for seasonal changes in ME intake due to seasonal differences in the varieties of leafy browse offered. Using the doubly labeled water method, we determined TEE for the female (288 kcal day -1) and male (411 kcal day -1). Resulting mean TEE was well below the expected value for marsupials and macropods (i.e., ~60% of the expected value based on body mass). The mean calculated ME intakes for the female and male were 307 kcal day-1 and 454 kcal day-1, respectively. There were significant seasonal differences in ME intakes, driven by reduced intake in the autumn. These results demonstrate that Matschie's tree kangaroos can be maintained at healthy body weights and conditions on fiber-rich and browse-heavy diets. Our findings contribute important insights into tree kangaroo energetics and physiology and can be applied to help reformulate the diet of Matschie's tree kangaroos at captive facilities to improve population health and sustainability.

Balancing growth, reproduction, maintenance, and activity in evolved energy economies.

Economic models predominate in life history research, which investigates the allocation of an organism's resources to growth, reproduction, and maintenance. These approaches typically employ a heuristic Y model of resource allocation, which predicts trade-offs among tasks within a fixed budget. The common currency among tasks is not always specified, but most models imply that metabolic energy, either from food or body stores, is the critical resource. Here, we review the evidence for metabolic energy as the common currency of growth, reproduction, and maintenance, focusing on studies in humans and other vertebrates. We then discuss the flow of energy to competing physiological tasks (physical activity, maintenance, and reproduction or growth) and its effect on life history traits. We propose a Ψ model of energy flow to these tasks, which provides an integrative framework for examining the influence of environmental factors and the expansion and contraction of energy budgets in the evolution of life history strategies.

Reindeer herders from subarctic Finland exhibit high total energy expenditure and low energy intake during the autumn herd roundup.

OBJECTIVE: High levels of total energy expenditure (TEE, kcal/day) have been documented among numerous human populations such as tropical climate horticulturalists and high-altitude agriculturalists. However, less work has been conducted among highly physically active cold climate populations. METHODS: In October 2018, TEE was measured using the doubly labeled water (TEEDLW , N = 10) and flex-heart rate methods (TEEHR , N = 24) for 6-14 days among reindeer herders (20-62 years) in northern Finland during an especially physically demanding, but not seasonally representative, period of the year for herders-the annual reindeer herd roundup. Self-reported dietary intake was also collected during TEE measurement periods. TEE was then compared to that of hunter gatherer, farming, and market economies. RESULTS: During the herd roundup, herders expended a mean of 4183 ± 949 kcal/day as measured by the DLW method, which was not significantly different from TEEHR . Mean caloric intake was 1718 ± 709 kcal/day, and was significantly lower than TEEDLW and TEEHR (p < .001). Herder TEEDLW was significantly higher than that of hunter gatherer (p = .0014) and market (p < .0014) economy populations; however, herder TEEDLW was not different from that of farming populations (p = .91). CONCLUSION: High TEE and low caloric intake among herders reflect the extreme demands placed on herders during the annual herd round up. Although TEEDLW was similar between cold climate herders and hot climate farming populations, there are likely differences in how that TEE is comprised, reflecting the local ecologies of these populations.

Total energy expenditure of bottlenose dolphins (Tursiops truncatus) of different ages.

Marine mammals are thought to have an energetically expensive lifestyle because endothermy is costly in marine environments. However, measurements of total energy expenditure (TEE; kcal day-1) are available only for a limited number of marine mammals, because large body size and inaccessible habitats make TEE measurements expensive and difficult to obtain for many taxa. We measured TEE in 10 adult common bottlenose dolphins (Tursiops truncatus) living in natural seawater lagoons at two facilities (Dolphin Research Center and Dolphin Quest) using the doubly labeled water method. We assessed the relative effects of body mass, age and physical activity on TEE. We also examined whether TEE of bottlenose dolphins, and more generally of marine mammals, differs from that expected for their body mass compared with other eutherian mammals, using phylogenetic least squares (PGLS) regressions. There were no differences in body mass or TEE (unadjusted TEE and TEE adjusted for fat-free mass) between dolphins from the two facilities. Our results show that adjusted TEE decreased and fat mass increased with age. Different measures of activity were not related to age, body fat or adjusted TEE. Both PGLS and the non-phylogenetic linear regression indicate that marine mammals have an elevated TEE compared with that of terrestrial mammals. However, bottlenose dolphins expended 17.1% less energy than other marine mammals of similar body mass. The two oldest dolphins (>40 years) showed a lower TEE, similar to the decline in TEE seen in older humans. To our knowledge, this is the first study to show an age-related metabolic decline in a large non-human mammal.

Daily energy expenditure through the human life course.

Total daily energy expenditure ("total expenditure") reflects daily energy needs and is a critical variable in human health and physiology, but its trajectory over the life course is poorly studied. We analyzed a large, diverse database of total expenditure measured by the doubly labeled water method for males and females aged 8 days to 95 years. Total expenditure increased with fat-free mass in a power-law manner, with four distinct life stages. Fat-free mass-adjusted expenditure accelerates rapidly in neonates to ~50% above adult values at ~1 year; declines slowly to adult levels by ~20 years; remains stable in adulthood (20 to 60 years), even during pregnancy; then declines in older adults. These changes shed light on human development and aging and should help shape nutrition and health strategies across the life span.

Effects of Evolution, Ecology, and Economy on Human Diet: Insights from Hunter-Gatherers and Other Small-Scale Societies.

We review the evolutionary origins of the human diet and the effects of ecology economy on the dietary proportion of plants and animals. Humans eat more meat than other apes, a consequence of hunting and gathering, which arose ∼2.5 Mya with the genus Homo. Paleolithic diets likely included a balance of plant and animal foods and would have been remarkably variable across time and space. A plant/animal food balance of 50/50% prevails among contemporary warm-climate hunter-gatherers, but these proportions vary widely. Societies in cold climates, and those that depend more on fishing or pastoralism, tend to eat more meat. Warm-climate foragers, and groups that engage in some farming, tend to eat more plants. We present a case study of the wild food diet of the Hadza, a community of hunter-gatherers in northern Tanzania, whose diet is high in fiber, adequate in protein, and remarkably variable over monthly timescales.

Determinants of climbing energetic costs in humans.

Previous studies in primates and other animals have shown that mass-specific cost of transport (J kg-1 m-1) for climbing is independent of body size across species, but little is known about within-species allometry of climbing costs or the effects of difficulty and velocity. Here, we assessed the effects of velocity, route difficulty and anatomical variation on the energetic cost of climbing within humans. Twelve experienced rock climbers climbed on an indoor wall over a range of difficulty levels and velocities, with energy expenditure measured via respirometry. We found no effect of body mass or limb proportions on mass-specific cost of transport among subjects. Mass-specific cost of transport was negatively correlated with climbing velocity. Increased route difficulty was associated with slower climbing velocities and thus higher costs, but there was no statistically significant effect of route difficulty on energy expenditure independent of velocity. Finally, human climbing costs measured in this study were similar to published values for other primates, suggesting arboreal adaptations have a negligible effect on climbing efficiency.

Hotter and sicker: External energy expenditure and the tangled evolutionary roots of anthropogenic climate change and chronic disease.

BACKGROUND: The dual crises of climate change and chronic, or non-communicable, disease (NCD) have emerged worldwide as the global economy has industrialized over the past two centuries. AIMS: In this synthesis I examine humans' dependence on external (non-metabolic) energy expenditure (e.g., fire, fossil fuels) as a common, root cause in these modern crises. MATERIALS AND METHODS: Using fossil, archeological, and historical evidence I show that the human lineage has been dependent on external energy sources since the control of fire in the Paleolithic. This reliance has grown with the development of agriculture, the use of wind- and water-power, and the most recently with industrialization and the transition to fossil fuels. To place industrialization in context I develop a Rule of 50, whereby individuals in industrialized economies consume roughly 50-times more external energy and manufacture roughly 50-times more material than do hunter-gatherers. RESULTS: Industrialization and mechanization, powered by fossil fuels, have promoted centralization and processing in food production, reduced physical activity, and increased air pollution (including greenhouse gas emissions). These developments have led in turn to NCD and climate change. DISCUSSION AND CONCLUSION: Climate change and NCD are connected both to one another and to our species' deep evolutionary dependence on external energy. Transitioning to carbon-free energy is essential to reduce the existential risks of climate change, but will likely have only modest effects on NCD. With the impending exhaustion of oil, coal, and natural gas reserves, developing replacements for fossil fuels is also critical to maintaining our species' external energy portfolio.

Gendered movement ecology and landscape use in Hadza hunter-gatherers.

Understanding how gendered economic roles structure space use is critical to evolutionary models of foraging behaviour, social organization and cognition. Here, we examine hunter-gatherer spatial behaviour on a very large scale, using GPS devices worn by Hadza foragers to record 2,078 person-days of movement. Theory in movement ecology suggests that the density and mobility of targeted foods should predict spatial behaviour and that strong gender differences should arise in a hunter-gatherer context. As predicted, we find that men walked further per day, explored more land, followed more sinuous paths and were more likely to be alone. These data are consistent with the ecology of male- and female-targeted foods and suggest that male landscape use is more navigationally challenging in this hunter-gatherer context. Comparisons of Hadza space use with space use data available for non-human primates suggest that the sexual division of labour likely co-evolved with increased sex differences in spatial behaviour and landscape use.

Childhood Daily Energy Expenditure Does Not Decrease with Market Integration and Is Not Related to Adiposity in Amazonia.

BACKGROUND: Childhood overweight and obesity (OW/OB) is increasingly centered in low- and middle-income countries (LMICs) as rural populations experience market integration and lifeway change. Most explanatory studies have relied on imprecise estimates of children's energy expenditure, restricting understanding of the relative effects of changes in diet and energy expenditure on the development of OW/OB in transitioning contexts. OBJECTIVES: This study used gold-standard measurements of children's energy expenditure to investigate the changes that underlie OW/OB and the nutrition/epidemiologic transition. METHODS: Cross-sectional data were collected from "rural" (n = 43) Shuar forager-horticulturalist children and their "peri-urban" (n = 34) Shuar counterparts (age 4-12 y) in Amazonian Ecuador. Doubly labeled water measurements of total energy expenditure (TEE; kcal/d), respirometry measurements of resting energy expenditure (REE; kcal/d), and measures of diet, physical activity, immune activity, and market integration were analyzed primarily using regression models. RESULTS: Peri-urban children had higher body fat percentage (+8.1%, P < 0.001), greater consumption of market-acquired foods (multiple P < 0.001), lower concentrations of immune activity biomarkers (multiple P < 0.05), and lower REE (-108 kcal/d, P = 0.002) than rural children. Despite these differences, peri-urban children's TEE was indistinguishable from that of rural children (P = 0.499). Moreover, although sample-wide IgG concentrations and household incomes predicted REE (both P < 0.05), no examined household, immune activity, or physical activity measures were related to children's overall TEE (all P > 0.09). Diet and energy expenditure associations with adiposity demonstrate that only reported consumption of market-acquired "protein" and "carbohydrate" foods predicted children's body fat levels (multiple P < 0.05). CONCLUSIONS: Despite underlying patterns in REE, Shuar children's TEE is not reliably related to market integration and-unlike dietary measures-does not predict adiposity. These findings suggest a leading role of changing dietary intake in transitions to OW/OB in LMICs.

Sex differences in respiratory and circulatory cost during hypoxic walking: potential impact on oxygen saturation.

Energy expenditure (EE) during treadmill walking under normal conditions (normobaric normoxia, 21% O2) and moderate hypoxia (13% O2) was measured. Ten healthy young men and ten healthy young women walked on a level (0°) gradient a range of speeds (0.67-1.67 m s-1). During walking, there were no significant differences in reductions in arterial oxygen saturation (SpO2) between the sexes. The hypoxia-induced increase in EE, heart rate (HR [bpm]) and ventilation ([Formula: see text] [L min-1]) were calculated. Using a multivariate model that combined EE, [Formula: see text], and HR to predict ΔSpO2 (hypoxia-induced reduction), a very strong fit model both for men (r2 = 0.900, P < 0.001) and for women was obtained (r2 = 0.957, P < 0.001). The contributions of EE, VE, and HR to ΔSpO2 were markedly different between men and women. [Formula: see text] and EE had a stronger effect on ΔSpO2 in women ([Formula: see text]: 4.1% in women vs. 1.7% in men; EE: 28.1% in women vs. 15.8% in men), while HR had a greater effect in men (82.5% in men and 67.9% in women). These findings suggested that high-altitude adaptation in response to hypoxemia has different underlying mechanisms between men and women. These results can help to explain how to adapt high-altitude for men and women, respectively.

Extreme events reveal an alimentary limit on sustained maximal human energy expenditure.

The limits on maximum sustained energy expenditure are unclear but are of interest because they constrain reproduction, thermoregulation, and physical activity. Here, we show that sustained expenditure in humans, measured as maximum sustained metabolic scope (SusMS), is a function of event duration. We compiled measurements of total energy expenditure (TEE) and basal metabolic rate (BMR) from human endurance events and added new data from adults running ~250 km/week for 20 weeks in a transcontinental race. For events lasting 0.5 to 250+ days, SusMS decreases curvilinearly with event duration, plateauing below 3× BMR. This relationship differs from that of shorter events (e.g., marathons). Incorporating data from overfeeding studies, we find evidence for an alimentary energy supply limit in humans of ~2.5× BMR; greater expenditure requires drawing down the body's energy stores. Transcontinental race data suggest that humans can partially reduce TEE during long events to extend endurance.