Increases in regional brain volume across two native South American male populations.

Industrialized environments, despite benefits such as higher levels of formal education and lower rates of infections, can also have pernicious impacts upon brain atrophy. Partly for this reason, comparing age-related brain volume trajectories between industrialized and non-industrialized populations can help to suggest lifestyle correlates of brain health. The Tsimane, indigenous to the Bolivian Amazon, derive their subsistence from foraging and horticulture and are physically active. The Moseten, a mixed-ethnicity farming population, are physically active but less than the Tsimane. Within both populations (N = 1024; age range = 46-83), we calculated regional brain volumes from computed tomography and compared their cross-sectional trends with age to those of UK Biobank (UKBB) participants (N = 19,973; same age range). Surprisingly among Tsimane and Moseten (T/M) males, some parietal and occipital structures mediating visuospatial abilities exhibit small but significant increases in regional volume with age. UKBB males exhibit a steeper negative trend of regional volume with age in frontal and temporal structures compared to T/M males. However, T/M females exhibit significantly steeper rates of brain volume decrease with age compared to UKBB females, particularly for some cerebro-cortical structures (e.g., left subparietal cortex). Across the three populations, observed trends exhibit no interhemispheric asymmetry. In conclusion, the age-related rate of regional brain volume change may differ by lifestyle and sex. The lack of brain volume reduction with age is not known to exist in other human population, highlighting the putative role of lifestyle in constraining regional brain atrophy and promoting elements of non-industrialized lifestyle like higher physical activity.

Women's subsistence strategies predict fertility across cultures, but context matters.

While it is commonly assumed that farmers have higher, and foragers lower, fertility compared to populations practicing other forms of subsistence, robust supportive evidence is lacking. We tested whether subsistence activities-incorporating market integration-are associated with fertility in 10,250 women from 27 small-scale societies and found considerable variation in fertility. This variation did not align with group-level subsistence typologies. Societies labeled as "farmers" did not have higher fertility than others, while "foragers" did not have lower fertility. However, at the individual level, we found strong evidence that fertility was positively associated with farming and moderate evidence of a negative relationship between foraging and fertility. Markers of market integration were strongly negatively correlated with fertility. Despite strong cross-cultural evidence, these relationships were not consistent in all populations, highlighting the importance of the socioecological context, which likely influences the diverse mechanisms driving the relationship between fertility and subsistence.

Gains to cooperation drive the evolution of egalitarianism.

What conditions favour egalitarianism, that is, muted hierarchies with relatively equal distributions of resources? Here, we combine the hawk-dove and prisoner's dilemma games to model the effects of economic defensibility, costs of competition and gains from cooperation on egalitarianism, operationalized as the absence of hawks. We show that a 'leveller' strategy, which punishes hawkishness in the hawk-dove game with defection in the prisoner's dilemma, can be evolutionarily stable provided that the gains from cooperation are high relative to the benefits of hawkishness. Under these conditions, rare mutant levellers select for hawks that acquiesce to punishment by playing dove. If these 'acquiescent hawks' become common, levellers outperform hawks and establish a new egalitarian equilibrium. An analysis of human foraging groups corroborates these results, as groups with a greater reliance on cooperation are more egalitarian. Cooperation fosters greater equality when individuals can withhold its benefits from would-be dominant individuals.

Very Low Prevalence and Incidence of Atrial Fibrillation among Bolivian Forager-Farmers.

Background: Atrial fibrillation is the most common arrhythmia in post-industrialized populations. Older age, hypertension, obesity, chronic inflammation, and diabetes are significant atrial fibrillation risk factors, suggesting that modern urban environments may promote atrial fibrillation. Objective: Here we assess atrial fibrillation prevalence and incidence among tropical horticulturalists of the Bolivian Amazon with high levels of physical activity, a lean diet, and minimal coronary atherosclerosis, but also high infectious disease burden and associated inflammation. Methods: Between 2005-2019, 1314 Tsimane aged 40-94 years (52% female) and 534 Moseten Amerindians aged 40-89 years (50% female) underwent resting 12-lead electrocardiograms to assess atrial fibrillation prevalence. For atrial fibrillation incidence assessment, 1059 (81% of original sample) Tsimane and 310 Moseten (58%) underwent additional ECGs (mean time to follow up 7.0, 1.8 years, respectively). Findings: Only one (male) of 1314 Tsimane (0.076%) and one (male) of 534 Moseten (0.187%) demonstrated atrial fibrillation at baseline. There was one new (female) Tsimane case in 7395 risk years for the 1059 participants with >1 ECG (incidence rate = 0.14 per 1,000 risk years). No new cases were detected among Moseten, based on 542 risk years. Conclusion: Tsimane and Moseten show the lowest levels of atrial fibrillation ever reported, 1/20 to ~1/6 of rates in high-income countries. These findings provide additional evidence that a subsistence lifestyle with high levels of physical activity, and a diet low in processed carbohydrates and fat is cardioprotective, despite frequent infection-induced inflammation. Findings suggest that atrial fibrillation is a modifiable lifestyle disease rather than an inevitable feature of cardiovascular aging.

Paternal provisioning results from ecological change.

Paternal provisioning among humans is puzzling because it is rare among primates and absent in nonhuman apes and because emergent provisioning would have been subject to paternity theft. A provisioning "dad" loses fitness at the hands of nonprovisioning, mate-seeking "cads." Recent models require exacting interplay between male provisioning and female choice to overcome this social dilemma. We instead posit that ecological change favored widespread improvements in male provisioning incentives, and we show theoretically how social obstacles to male provisioning can be overcome. Greater availability of energetically rich, difficult-to-acquire foods enhances female-male and male-male complementarities, thus altering the fitness of dads versus cads. We identify a tipping point where gains from provisioning overcome costs from paternity uncertainty and the dad strategy becomes viable. Stable polymorphic states are possible, meaning that dads need not necessarily eliminate cads. Our simulations suggest that with sufficient complementarities, dads can emerge even in the face of high paternity uncertainty. Our theoretical focus on ecological change as a primary factor affecting the trade-off between male mating and parenting effort suggests different possibilities for using paleo-climatic, archaeological, and genomic evidence to establish the timing of and conditions associated with emergence of paternal provisioning in the hominin lineage.

Voluntary collective isolation as a best response to COVID-19 for indigenous populations? A case study and protocol from the Bolivian Amazon.

Indigenous communities worldwide share common features that make them especially vulnerable to the complications of and mortality from COVID-19. They also possess resilient attributes that can be leveraged to promote prevention efforts. How can indigenous communities best mitigate potential devastating effects of COVID-19? In Bolivia, where nearly half of all citizens claim indigenous origins, no specific guidelines have been outlined for indigenous communities inhabiting native communal territories. In this Public Health article, we describe collaborative efforts, as anthropologists, physicians, tribal leaders, and local officials, to develop and implement a multiphase COVID-19 prevention and containment plan focused on voluntary collective isolation and contact-tracing among Tsimane forager-horticulturalists in the Bolivian Amazon. Phase 1 involves education, outreach, and preparation, and phase 2 focuses on containment, patient management, and quarantine. Features of this plan might be exported and adapted to local circumstances elsewhere to prevent widespread mortality in indigenous communities.

Greater wealth inequality, less polygyny: rethinking the polygyny threshold model.

Monogamy appears to have become the predominant human mating system with the emergence of highly unequal agricultural populations that replaced relatively egalitarian horticultural populations, challenging the conventional idea-based on the polygyny threshold model-that polygyny should be positively associated with wealth inequality. To address this polygyny paradox, we generalize the standard polygyny threshold model to a mutual mate choice model predicting the fraction of women married polygynously. We then demonstrate two conditions that are jointly sufficient to make monogamy the predominant marriage form, even in highly unequal societies. We assess if these conditions are satisfied using individual-level data from 29 human populations. Our analysis shows that with the shift to stratified agricultural economies: (i) the population frequency of relatively poor individuals increased, increasing wealth inequality, but decreasing the frequency of individuals with sufficient wealth to secure polygynous marriage, and (ii) diminishing marginal fitness returns to additional wives prevent extremely wealthy men from obtaining as many wives as their relative wealth would otherwise predict. These conditions jointly lead to a high population-level frequency of monogamy.

Information transmission and the oral tradition: Evidence of a late-life service niche for Tsimane Amerindians.

Storytelling can affect wellbeing and fitness by transmitting information and reinforcing cultural codes of conduct. Despite their potential importance, the development and timing of storytelling skills, and the transmission of story knowledge have received minimal attention in studies of subsistence societies that more often focus on food production skills. Here we examine how storytelling and patterns of information transmission among Tsimane forager-horticulturalists are predicted by the changing age profiles of storytellers' abilities and accumulated experience. We find that storytelling skills are most developed among older adults who demonstrate superior knowledge of traditional stories and who report telling stories most. We find that the important information transmitted via storytelling typically flows from older to younger generations, and stories are primarily learned from older same-sex relatives, especially grandparents. Our findings suggest that the oral tradition provides a specialized late-life service niche for Tsimane adults who have accumulated important experience and knowledge relevant to foraging and sociality, but have lost comparative advantage in other productive domains. These findings may help extend our understanding of the evolved human life history by illustrating how changes in embodied capital predict the development of information transmission services in a forager-horticulturalist economy.

Experimental tests of the tolerated theft and risk-reduction theories of resource exchange.

Here we report the results of an experiment that tests the reciprocal risk reduction 1 and 'tolerated theft' or taking hypotheses 2 for why the human species is unique in having extensive exchange of resources among non-kin. We designed an experiment to determine whether, in response to variance of resource acquisition, people exchange food resources via taking or, alternatively, form reciprocal relationships based on giving. In the experiment, subjects forage individually, experience variation in resource acquisition, and then consume either by actions in which resources are taken from, or moved to, others in a group environment. The key feature of the experimental design is that individuals can transfer resources to others, attempt to take resources from others and defend against take-away attempts. Many subjects begin by attempting to take resources from others, who can resist those attempts at a cost to each. Over time, subjects shift to a cooperative strategy of voluntary reciprocal giving, a response not suggested by the instructions. These results provide evidence that people are independently able to overcome initial use of taking strategies, evaluate gains from exchange, and create endogenous reciprocal trading relationships as a response to unsynchronized variance in resource availability.

Conflict or congruence? Maternal and infant-centric factors associated with shorter exclusive breastfeeding durations among the Tsimane.

Six months of exclusive breastfeeding (EBF) is considered optimal for infant health, though globally most infants begin complementary feeding (CF) earlier-including among populations that practice prolonged breastfeeding. Two frameworks for understanding patterns of early CF emerge in the literature. In the first, maternal and infant needs trade-off, as "maternal-centric" factors-related to time and energy demands, reproductive investment, cultural influences, and structural barriers- favor supplanting breastfeeding with earlier and increased CF. A second framework considers that "infant-centric" factors-related to infant energetic needs-favor CF before six months to supplement breastfeeding. We apply these two frameworks in examining early CF among the Tsimane-a high-fertility, high-mortality, forager-horticulturalist population residing in the Bolivian Amazon. Data were collected from a mixed-longitudinal sample of 161 Tsimane mother-infant pairs from August 2012-April 2013. Tsimane mothers generally reported introducing CF because of perceived infant needs. However, CF is introduced with continued intensive breastfeeding, and generally coupled with premastication. Risks of earlier CF relative to the minimum hazard (estimated at 5 births) were elevated for lower and higher parity mothers, but were significantly greater only after 9 births. Seventeen percent of mothers reported introducing CF because of low milk supply. Introducing CF because of low milk was most common from 0 to 3 months of age and among higher parity mothers, which may reflect physiological constraints. Maternal reproductive trade-offs and perceived infant needs may help explain the low prevalence of EBF to six months among other populations in which breastfeeding is not structurally or culturally constrained.

Cardiovascular disease and type 2 diabetes in evolutionary perspective: a critical role for helminths?

Heart disease and type 2 diabetes are commonly believed to be rare among contemporary subsistence-level human populations, and by extension prehistoric populations. Although some caveats remain, evidence shows these diseases to be unusual among well-studied hunter-gatherers and other subsistence populations with minimal access to healthcare. Here we expand on a relatively new proposal for why these and other populations may not show major signs of these diseases. Chronic infections, especially helminths, may offer protection against heart disease and diabetes through direct and indirect pathways. As part of a strategy to insure their own survival and reproduction, helminths exert multiple cardio-protective effects on their host through their effects on immune function and blood lipid metabolism. Helminths consume blood lipids and glucose, alter lipid metabolism, and modulate immune function towards Th-2 polarization - which combined can lower blood cholesterol, reduce obesity, increase insulin sensitivity, decrease atheroma progression, and reduce likelihood of atherosclerotic plaque rupture. Traditional cardiometabolic risk factors, coupled with the mismatch between our evolved immune systems and modern, hygienic environments may interact in complex ways. In this review, we survey existing studies in the non-human animal and human literature, highlight unresolved questions and suggest future directions to explore the role of helminths in the etiology of cardio-metabolic disease.

High resting metabolic rate among Amazonian forager-horticulturalists experiencing high pathogen burden.

OBJECTIVES: Resting metabolic rate (RMR) reflects energetic costs of homeostasis and accounts for 60 to 75% of total energy expenditure (TEE). Lean mass and physical activity account for much RMR variability, but the impact of prolonged immune activation from infection on human RMR is unclear in naturalistic settings. We evaluate the effects of infection on mass-corrected RMR among Bolivian forager-horticulturalists, and assess whether RMR declines more slowly with age than in hygienic sedentary populations, as might be expected if older adults experience high pathogen burden. MATERIALS AND METHODS: RMR was measured by indirect calorimetry (Fitmate MED, Cosmed) in 1,300 adults aged 20 to 90 and TEE was measured using doubly labeled water (n = 40). Immune biomarkers, clinical diagnoses, and anthropometrics were collected by the Tsimane Health and Life History Project. RESULTS: Tsimane have higher RMR and TEE than people in sedentary industrialized populations. Tsimane RMR is 18 to 47% (women) and 22 to 40% (men) higher than expected using six standard prediction equations. Tsimane mass-corrected TEE is similarly elevated compared to Westerners. Elevated leukocytes and helminths are associated with excess RMR in multivariate regressions, and jointly result in a predicted excess RMR of 10 to 15%. After age 40, RMR declines by 69 kcal/decade (p < .0001). Controlling for lean mass and height accounts for 71% of age-related RMR decline, and adding indicators of infection minimally affects the age slope. The residual level of age-related decline from age 40 is 1.2% per decade. CONCLUSION: High pathogen burden may lead to higher metabolic costs, which may be offset by smaller body mass or other energy-sparing mechanisms.

Status competition, inequality, and fertility: implications for the demographic transition.

The role that social status plays in small-scale societies suggests that status may be important for understanding the evolution of human fertility decisions, and for understanding how such decisions play out in modern contexts. This paper explores whether modelling competition for status--in the sense of relative rank within a society--can help shed light on fertility decline and the demographic transition. We develop a model of how levels of inequality and status competition affect optimal investment by parents in the embodied capital (health, strength, and skills) and social status of offspring, focusing on feedbacks between individual decisions and socio-ecological conditions. We find that conditions similar to those in demographic transition societies yield increased investment in both embodied capital and social status, generating substantial decreases in fertility, particularly under conditions of high inequality and intense status competition. We suggest that a complete explanation for both fertility variation in small-scale societies and modern fertility decline will take into account the effects of status competition and inequality.

Skills, division of labour and economies of scale among Amazonian hunters and South Indian honey collectors.

In foraging and other productive activities, individuals make choices regarding whether and with whom to cooperate, and in what capacities. The size and composition of cooperative groups can be understood as a self-organized outcome of these choices, which are made under local ecological and social constraints. This article describes a theoretical framework for explaining the size and composition of foraging groups based on three principles: (i) the sexual division of labour; (ii) the intergenerational division of labour; and (iii) economies of scale in production. We test predictions from the theory with data from two field contexts: Tsimane' game hunters of lowland Bolivia, and Jenu Kuruba honey collectors of South India. In each case, we estimate the impacts of group size and individual group members' effort on group success. We characterize differences in the skill requirements of different foraging activities and show that individuals participate more frequently in activities in which they are more efficient. We evaluate returns to scale across different resource types and observe higher returns at larger group sizes in foraging activities (such as hunting large game) that benefit from coordinated and complementary roles. These results inform us that the foraging group size and composition are guided by the motivated choice of individuals on the basis of relative efficiency, benefits of cooperation, opportunity costs and other social considerations.

Skill ontogeny among Tsimane forager-horticulturalists.

OBJECTIVES: We investigate whether age profiles of Tsimane forager-horticulturalists' reported skill development are consistent with predictions derived from life history theory about the timing of productivity and reproduction. Previous studies of forager skill development have often focused on a few abilities (e.g. hunting), and neglected the broad range of skills and services typical of forager economies (e.g. childcare, craft production, music performance, story-telling). MATERIALS AND METHODS: By systematically examining age patterns in reported acquisition, proficiency, and expertise across a broad range of activities including food production, childcare, and other services, we provide the most complete skill development study of a traditional subsistence society to date. RESULTS: Our results show that: (1) most essential skills are acquired prior to first reproduction, then developed further so that their productive returns meet the increasing demands of dependent offspring during adulthood; (2) as postreproductive adults age beyond earlier years of peak performance, they report developing additional conceptual and procedural proficiency, and despite greater physical frailty than younger adults, are consensually regarded as the most expert (especially in music and storytelling), consistent with their roles as providers and educators. We find that adults have accurate understandings of their skillsets and skill levels -an important awareness for social exchange, comparison, learning, and pedagogy. DISCUSSION: These findings extend our understanding of the evolved human life history by illustrating how changes in embodied capital and the needs of dependent offspring predict the development of complementary skills and services in a forager-horticulturalist economy.

Challenging the Inevitability of Prostate Enlargement: Low Levels of Benign Prostatic Hyperplasia Among Tsimane Forager-Horticulturalists.

BACKGROUND: Often considered an inevitable part of male aging, benign prostatic hyperplasia (BPH) is the most common non-life threatening disease to affect men in Western populations. We examine age-related change in prostate size and BPH risk and related serum biomarkers among the Tsimane Amerindians of the Bolivian Amazon who live a traditional lifestyle of hunting and small-scale horticulture. The Tsimane are a critical case study for understanding the etiology of BPH as they have low levels of obesity and metabolic syndrome, as well as lower levels of testosterone than age matched U.S. males, factors associated with BPH in previous research. METHODS: Ultrasounds were conducted on 348 men aged 28-89 years (median age 56 years). Testosterone, prostate specific antigen, sex hormone binding globulin, and glycosylated hemoglobin were examined in relationship to prostate size and BPH. RESULTS: Tsimane have less than half of the BPH prevalence experienced by U.S. men, and prostate volumes 62.6% smaller. While Tsimane have low levels of testosterone and subclinical levels of metabolic syndrome compared to U.S. men, Tsimane with high testosterone were more likely to experience BPH, as were those with higher glycosylated hemoglobin, suggesting targets for clinical interventions to reduce BPH. CONCLUSIONS: These results have clinical significance for the growing number of men taking testosterone supplementation; even at low levels the additional testosterone exposure could be placing these men at higher risk of BPH. Overall, these data suggest that BPH may not have been an inevitable part of male aging throughout human evolutionary history.

Salivary oxytocin increases concurrently with testosterone and time away from home among returning Tsimane' hunters.

Oxytocin, testosterone and cortisol can have opposing effects on social behaviour, yet few studies have examined their interactions. We measured changes in salivary oxytocin, testosterone and cortisol among Tsimane' men returning home after hunting, an ancient context of male status competition, parental investment and cooperation. Contra normal diurnal rhythm, oxytocin increased relative to baseline and this increase was positively associated with duration of the hunt and change in testosterone, but not cortisol, social context, hunting outcome or physical activity. The concurrent increase in endogenous peripheral oxytocin and testosterone is unexpected given their opposing independent effects on social cognition and behaviour, and has not been observed before. We discuss the potential significance of these effects for the biology of pair-bonding, parenting and social foraging in humans and other species.

Inclusive fitness and differential productivity across the life course determine intergenerational transfers in a small-scale human society.

Transfers of resources between generations are an essential element in current models of human life-history evolution accounting for prolonged development, extended lifespan and menopause. Integrating these models with Hamilton's theory of inclusive fitness, we predict that the interaction of biological kinship with the age-schedule of resource production should be a key driver of intergenerational transfers. In the empirical case of Tsimane' forager-horticulturalists in Bolivian Amazonia, we provide a detailed characterization of net transfers of food according to age, sex, kinship and the net need of donors and recipients. We show that parents, grandparents and siblings provide significant net downward transfers of food across generations. We demonstrate that the extent of provisioning responds facultatively to variation in the productivity and demographic composition of families, as predicted by the theory. We hypothesize that the motivation to provide these critical transfers is a fundamental force that binds together human nuclear and extended families. The ubiquity of three-generational families in human societies may thus be a direct reflection of fundamental evolutionary constraints on an organism's life-history and social organization.

Political influence associates with cortisol and health among egalitarian forager-farmers.

BACKGROUND AND OBJECTIVES: Low social status increases risk of disease due, in part, to the psychosocial stress that accompanies feeling subordinate or poor. Previous studies report that chronic stress and chronically elevated cortisol can impair cardiovascular and immune function. We test whether lower status is more benign in small-scale, relatively egalitarian societies, where leaders lack coercive authority and there is minimal material wealth to contest. METHODOLOGY: Among Tsimane' forager-horticulturalists of lowland Bolivia, we compare informal political influence among men with urinary cortisol, immune activation (innate and acquired), and morbidity as assessed during routine medical exams. RESULTS: After controlling for potential confounds, we find that politically influential men have lower cortisol, and that this association is partly attributable to access to social support. Cortisol is positively associated with men's income, which may reflect chronic psychosocial stress from market involvement. Greater influence is also associated with lower probability of respiratory infection, which is a frequent source of morbidity among Tsimane'. Among men who lost influence over a 4-year period, cortisol and probability of respiratory infection were higher the greater the decline in influence. CONCLUSIONS AND IMPLICATIONS: Deleterious effects of low status on health are not merely 'diseases of civilization' but may result from how (even subtle) status differences structure human behavior.