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.

Increased physical activity is not related to markers of cardiometabolic health in two lemur species.

Insufficient physical activity is a major risk factor for cardiometabolic disease (i.e., unhealthy weight gain, heart disease, and diabetes) in humans and may also negatively affect health of primates in human care. Effects of physical activity on energy expenditure and cardiometabolic health are virtually unstudied in nonhuman primates. We investigated physical activity and metabolic markers in 15 adult ring-tailed lemurs (Lemur catta) and 11 Coquerel's sifakas (Propithecus coquereli) at the Duke Lemur Center during a period of low activity in winter when the animals were housed in buildings (with outdoor access) and a period of high activity when individuals were free-ranging in large, outdoor, forested enclosures. We compared body mass, blood glucose, triglycerides, HDL- and LDL-cholesterol, physical activity via accelerometry, and total energy expenditure (TEE) via the doubly labeled water method (in ring-tailed lemurs only) between both conditions. Both species were more active and had a lower body mass in summer. Ring-tailed lemurs had a higher TEE and lower triglyceride levels in summer, whereas sifaka had higher triglyceride levels in summer. Individuals that increased their activity more, also lost more body mass. Individuals that lost more body mass, also had a positive change in HDL-cholesterol (i.e., higher values in summer). Changes in activity were not associated with changes in markers of metabolic health, body fat percentage and TEE (both unadjusted and adjusted for body composition). Older age was associated with lower activity in both species, and decreased glucose in ring-tailed lemurs, but was otherwise unrelated to metabolic markers and, for ring-tailed lemurs, adjusted TEE. Overall, body mass was lower during summer but the increase in physical activity did not strongly influence metabolic health or TEE in these populations.

Effect of influenza vaccination on resting metabolic rate and c-reactive protein concentrations in healthy young adults.

OBJECTIVES: Chronic immune activation and severe inflammatory states are positively associated with resting metabolic rate (RMR; kcal/day), but the impacts of mild immune stimuli on metabolism are poorly understood. This study investigates the within-individual association between the inflammatory response to influenza vaccination and RMR in young adults. METHODS: We evaluated RMRs through indirect calorimetry and circulating c-reactive protein (CRP) concentrations (mg/L)-a direct measure of inflammation-via high-sensitivity immunoassays of dried blood spots (n = 17) at baseline and two- and seven-days post-vaccine. Wilcoxon matched-pairs signed-rank tests were used to evaluate the magnitude of the CRP and RMR responses. Type II Wald chi-square tests of linear mixed-effect models assessed whether those responses were correlated. RESULTS: Baseline CRP was 1.39 ± 1.26 mg/L. On day two post-vaccine, CRP increased by 1.47 ± 1.37 mg/L (p < 0.0001), representing a 106% increase above baseline values. CRP remained higher on day seven post-vaccine, 1.32 ± 2.47 mg/L (p = 0.05) above baseline values. There were no statistically significant changes in RMR from baseline to day two (p = 0.98) or day seven (p = 0.21). Change in CRP from baseline did not predict RMR variation across days (p = 0.46). CONCLUSIONS: We find no evidence that adult influenza vaccination results in a corresponding increase in RMR. These results suggest that the energetic cost of an influenza vaccine's mild inflammatory stimulus is either too small to detect or is largely compensated by a temporary downregulation of energy allocated to other metabolic tasks.

The effects of lifestyle change on indicators of cardiometabolic health in semi-nomadic pastoralists.

Background and objectives: Non-communicable disease risk and the epidemic of cardiometabolic diseases continue to grow across the expanding industrialized world. Probing the relationships between evolved human physiology and modern socioecological conditions is central to understanding this health crisis. Therefore, we investigated the relationships between increased market access, shifting subsistence patterns and cardiometabolic health indicators within Daasanach semi-nomadic pastoralists who vary in their engagement in traditional lifestyle and emerging market behaviors. Methodology: We conducted cross-sectional socioecological, demographic and lifestyle stressor surveys along with health, biomarker and nutrition examinations among 225 (51.6% female) Daasanach adults in 2019-2020. We used linear mixed-effects models to test how differing levels of engagement in market integration and traditional subsistence activities related to blood pressure (BP), body composition and blood chemistry. Results: We found that systolic and diastolic BP, as well as the probability of having high BP (hypertension), were negatively associated with distance to market, a proxy for market integration. Additionally, body composition varied significantly by socioeconomic status (SES), with significant positive associations between BMI and body fat and higher SES among adults. Conclusions and implications: While evidence for evolutionary mismatch and health variation have been found across a number of populations affected by an urban/rural divide, these results demonstrate the effects of market integration and sedentarization on cardiometabolic health associated with the early stages of lifestyle changes. Our findings provide evidence for the changes in health when small-scale populations begin the processes of sedentarization and market integration that result from myriad market pressures.

The provisioned primate: patterns of obesity across lemurs, monkeys, apes and humans.

Non-human primates are potentially informative but underutilized species for investigating obesity. I examined patterns of obesity across the Primate order, calculating the ratio of body mass in captivity to that in the wild. This index, relative body mass, for n = 40 non-human primates (mean ± s.d.: females: 1.28 ± 0.30, range 0.67-1.78, males: 1.24 ± 0.28, range 0.70-1.97) overlapped with a reference value for humans (women: 1.52, men: 1.44). Among non-human primates, relative body mass was unrelated to dietary niche, and was marginally greater among female cohorts of terrestrial species. Males and females had similar relative body masses, but species with greater sexual size dimorphism (male/female mass) in wild populations had comparatively larger female body mass in captivity. Provisioned populations in wild and free-ranging settings had similar relative body mass to those in research facilities and zoos. Compared to the wild, captive diets are unlikely to be low in protein or fat, or high in carbohydrate, suggesting these macronutrients are not driving overeating in captive populations. Several primate species, including chimpanzees, a sister-species to humans, had relative body masses similar to humans. Humans are not unique in the propensity to overweight and obesity. This article is part of a discussion meeting issue 'Causes of obesity: theories, conjectures and evidence (Part II)'.

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.

Lifestyle and patterns of physical activity in Hadza foragers.

OBJECTIVES: Physically active lifestyles are associated with several health benefits. Physical activity (PA) levels are low in post-industrial populations, but generally high throughout life in subsistence populations. The Hadza are a subsistence-oriented foraging population in Tanzania known for being physically active, but it is unknown how recent increases in market integration may have altered their PA patterns. In this study, we examine PA patterns for Hadza women and men who engage in different amounts of traditional foraging. MATERIALS AND METHODS: One hundred and seventy seven Hadza participants (51% female, 19-87 years) wore an Axivity accelerometer (dominant wrist) for ~6 days during dry season months. We evaluated the effects of age, sex, and lifestyle measures on four PA measures that capture different aspects of the PA profile. RESULTS: Participants engaged in high levels of both moderate-intensity PA and inactivity. Although PA levels were negatively associated with age, older participants were still highly active. We found no differences in PA between participants living in more traditional "bush" camps and those living in more settled "village" camps. Mobility was positively associated with step counts for female participants, and schooling was positively associated with inactive time for male participants. CONCLUSIONS: The similarity in PA patterns between Hadza participants in different camp types suggests that high PA levels characterize subsistence lifestyles generally. The sex-based difference in the effects of mobility and schooling on PA could be a reflection of the Hadza's gender-based division of labor, or indicate that changes to subsistence-oriented lifestyles impact women and men in different ways.

Applying an evolutionary mismatch framework to understand disease susceptibility.

Noncommunicable diseases (NCDs) are on the rise worldwide. Obesity, cardiovascular disease, and type 2 diabetes are among a long list of "lifestyle" diseases that were rare throughout human history but are now common. The evolutionary mismatch hypothesis posits that humans evolved in environments that radically differ from those we currently experience; consequently, traits that were once advantageous may now be "mismatched" and disease causing. At the genetic level, this hypothesis predicts that loci with a history of selection will exhibit "genotype by environment" (GxE) interactions, with different health effects in "ancestral" versus "modern" environments. To identify such loci, we advocate for combining genomic tools in partnership with subsistence-level groups experiencing rapid lifestyle change. In these populations, comparisons of individuals falling on opposite extremes of the "matched" to "mismatched" spectrum are uniquely possible. More broadly, the work we propose will inform our understanding of environmental and genetic risk factors for NCDs across diverse ancestries and cultures.

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.

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.

Impact of energy availability and physical activity on variation in fertility across human populations.

Human reproduction is energetically costly, even more so than other primates. In this review, we consider how the energy cost of physical activity impacts reproductive tasks. Daily energy expenditure appears to be constrained, leading to trade-offs between activity and reproduction expenditures in physically active populations. High workloads can lead to suppression of basal metabolic rate and low gestational weight gain during pregnancy and longer interbirth intervals. These responses lead to variation in fertility, including age at first reproduction and interbirth interval. The influence of energetics is evident even in industrialized populations, where cultural and economic factors predominate. With the decoupling of skills acquisition from food procurement, extrasomatic resources and investment in individual offspring becomes very costly. The result is greater investment in fewer offspring. We present a summary of age at first reproduction and interbirth interval trends across a diverse, global sample representing 44 countries and two natural fertility populations. While economic factors impact fertility, women in energy-rich, industrialized populations are capable of greater reproductive output than women in energy-stressed populations. Thus, energetic factors can be disentangled from cultural and economic impacts on fertility. Future research should focus on objective measurements of energy intake, energy expenditure, and physical activity in a broader sample of populations to elucidate the role of energetics in shaping reproductive outcomes and health.

Evolutionary mismatch and the role of GxE interactions in human disease.

Globally, we are witnessing the rise of complex, non-communicable diseases (NCDs) related to changes in our daily environments. Obesity, asthma, cardiovascular disease, and type 2 diabetes are part of a long list of "lifestyle" diseases that were rare throughout human history but are now common. A key idea from anthropology and evolutionary biology-the evolutionary mismatch hypothesis-seeks to explain this phenomenon. It posits that humans evolved in environments that radically differ from the ones experienced by most people today, and thus traits that were advantageous in past environments may now be "mismatched" and disease-causing. This hypothesis is, at its core, a genetic one: it predicts that loci with a history of selection will exhibit "genotype by environment" (GxE) interactions and have differential health effects in ancestral versus modern environments. Here, we discuss how this concept could be leveraged to uncover the genetic architecture of NCDs in a principled way. Specifically, we advocate for partnering with small-scale, subsistence-level groups that are currently transitioning from environments that are arguably more "matched" with their recent evolutionary history to those that are more "mismatched". These populations provide diverse genetic backgrounds as well as the needed levels and types of environmental variation necessary for mapping GxE interactions in an explicit mismatch framework. Such work would make important contributions to our understanding of environmental and genetic risk factors for NCDs across diverse ancestries and sociocultural contexts.

Early childhood growth in Daasanach pastoralists of Northern Kenya: Distinct patterns of faltering in linear growth and weight gain.

OBJECTIVES: Investigations of early childhood growth among small-scale populations are essential for understanding human life history variation and enhancing the ability to serve such communities through global public health initiatives. This study characterizes early childhood growth trajectories and identifies differences in growth patterns relative to international references among Daasanach semi-nomadic pastoralist children living in a hot, arid region of northern Kenya. METHODS: A large sample of height and weight measures were collected from children (N = 1756; total observations = 4508; age = 0-5 years) between 2018 and 2020. Daasanach growth was compared to international reference standards and Daasanach-specific centile growth curves and pseudo-velocity models were generated using generalized additive models for location scale and size. RESULTS: Compared to World Health Organization (WHO) reference, relatively few Daasanach children were stunted (14.3%), while a large proportion were underweight (38.5%) and wasted (53.6%). Additionally, Daasanach children had a distinctive pattern of growth, marked by an increase in linear growth velocity after 24 months of age and relatively high linear growth velocity throughout the rest of early childhood. CONCLUSIONS: These results identify a unique pattern of early childhood growth faltering among children in a small-scale population and may reflect a thermoregulatory adaptation to their hot, arid environment. As linear growth and weight gain remain important indicators of health, the results of this study provide insight into growth velocity variations. This study has important implications for global public health efforts to identify and address sources of early growth faltering and undernutrition in small-scale populations.

Gray squirrels consume anthropogenic food waste most often during winter.

Urban habitats provide wildlife with predictable, easily accessible and abundant food sources in the form of human food waste. Urban eastern gray squirrels (Sciurus carolinensis) are commonly observed feeding in trash bins, but we lack data regarding the type, quantity and seasonal changes in food waste usage. We observed five trash bins on an urban university campus during four different observation periods. We recorded the time squirrels spent on and inside trash bins and type of retrieved food items. We also recorded ambient temperature, human presence and trash bin filling. Moreover, we determined changes in squirrel population density in a natural and three anthropogenic habitats during the same periods. Trash bins were fuller when human presence was higher. The higher human presence, the more squirrels went on and inside the bin, but there was no effect on number of retrieved food items. Trash bin usage by squirrels decreased when ambient temperature and bin filling increased. Most food items were retrieved during the coldest observation period, a period of high human presence, and the majority of retrieved food items were starchy foods (e.g., bread, French fries). The relationship between the number of squirrels observed along transects and a measure of urbanization, the normalized difference built-up index, was negative in periods with high ambient temperatures and positive in periods with low ambient temperatures, indicating winter may be less challenging in urban areas, likely facilitated by the availability of anthropogenic food sources, allowing a higher level of activity throughout winter. Supplementary Information: The online version contains supplementary material available at 10.1007/s42991-022-00326-3.

Variation in human water turnover associated with environmental and lifestyle factors.

Water is essential for survival, but one in three individuals worldwide (2.2 billion people) lacks access to safe drinking water. Water intake requirements largely reflect water turnover (WT), the water used by the body each day. We investigated the determinants of human WT in 5604 people from the ages of 8 days to 96 years from 23 countries using isotope-tracking (2H) methods. Age, body size, and composition were significantly associated with WT, as were physical activity, athletic status, pregnancy, socioeconomic status, and environmental characteristics (latitude, altitude, air temperature, and humidity). People who lived in countries with a low human development index (HDI) had higher WT than people in high-HDI countries. On the basis of this extensive dataset, we provide equations to predict human WT in relation to anthropometric, economic, and environmental factors.