Brain volume, energy balance, and cardiovascular health in two nonindustrial South American populations.

Little is known about brain aging or dementia in nonindustrialized environments that are similar to how humans lived throughout evolutionary history. This paper examines brain volume (BV) in middle and old age among two indigenous South American populations, the Tsimane and Moseten, whose lifestyles and environments diverge from those in high-income nations. With a sample of 1,165 individuals aged 40 to 94, we analyze population differences in cross-sectional rates of decline in BV with age. We also assess the relationships of BV with energy biomarkers and arterial disease and compare them against findings in industrialized contexts. The analyses test three hypotheses derived from an evolutionary model of brain health, which we call the embarrassment of riches (EOR). The model hypothesizes that food energy was positively associated with late life BV in the physically active, food-limited past, but excess body mass and adiposity are now associated with reduced BV in industrialized societies in middle and older ages. We find that the relationship of BV with both non-HDL cholesterol and body mass index is curvilinear, positive from the lowest values to 1.4 to 1.6 SDs above the mean, and negative from that value to the highest values. The more acculturated Moseten exhibit a steeper decrease in BV with age than Tsimane, but still shallower than US and European populations. Lastly, aortic arteriosclerosis is associated with lower BV. Complemented by findings from the United States and Europe, our results are consistent with the EOR model, with implications for interventions to improve brain health.

Navigating cross-cultural research: methodological and ethical considerations.

The intensifying pace of research based on cross-cultural studies in the social sciences necessitates a discussion of the unique challenges of multi-sited research. Given an increasing demand for social scientists to expand their data collection beyond WEIRD (Western, educated, industrialized, rich and democratic) populations, there is an urgent need for transdisciplinary conversations on the logistical, scientific and ethical considerations inherent to this type of scholarship. As a group of social scientists engaged in cross-cultural research in psychology and anthropology, we hope to guide prospective cross-cultural researchers through some of the complex scientific and ethical challenges involved in such work: (a) study site selection, (b) community involvement and (c) culturally appropriate research methods. We aim to shed light on some of the difficult ethical quandaries of this type of research. Our recommendation emphasizes a community-centred approach, in which the desires of the community regarding research approach and methodology, community involvement, results communication and distribution, and data sharing are held in the highest regard by the researchers. We argue that such considerations are central to scientific rigour and the foundation of the study of human behaviour.

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.

Low mineral density of a weight-bearing bone among adult women in a high fertility population.

Evolutionary theories of aging posit that greater reproductive effort causes somatic decline given a fundamental trade-off between investing energy in reproduction and repair. Few studies in high fertility human populations support this hypothesis, and problems of phenotypic correlation can obscure the expected trade-off between reproduction and somatic condition. This cross-sectional study investigates whether greater reproductive effort is associated with reduced calcaneal bone mineral density (BMD) among female Tsimane forager-farmers of lowland Bolivia. We also investigate whether female Tsimane BMD values are lower than sex- and age-matched US reference values, despite the fact that Tsimane engage in higher physical activity levels that can increase mechanical loading. To measure calcaneal BMD, quantitative ultrasonography was performed on 130 women (mean ± SD age = 36.6 ± 15.7, range = 15-75) that were recruited regardless of past or current reproductive status. Anthropometric and demographic data were collected during routine medical exams. As predicted, higher parity, short inter-birth interval, and earlier age at first birth are associated with reduced BMD among Tsimane women after adjusting for potential confounders. Population-level differences are apparent prior to the onset of reproduction, and age-related decline in BMD is greater among Tsimane compared with American women. Greater cumulative reproductive burden may lower calcaneal BMD individually and jointly with other lifestyle and heritable factors. Fitness impacts of kin transfers in adulthood may determine the value of investments in bone remodeling, and thus affect selection on age-profiles of bone mineral loss.

Cross-cousin marriage among Tsimane forager-horticulturalists during demographic transition and market integration.

Although still prevalent in many human societies, the practice of cousin marriage has precipitously declined in populations undergoing rapid demographic and socioeconomic change. However, it is still unclear whether changes in the structure of the marriage pool or changes in the fitness-relevant consequences of cousin marriage more strongly influence the frequency of cousin marriage. Here, we use genealogical data collected by the Tsimane Health and Life History Project to show that there is a small but measurable decline in the frequency of first cross-cousin marriage since the mid-twentieth century. Such changes are linked to concomitant changes in the pool of potential spouses in recent decades. We find only very modest differences in fitness-relevant demographic measures between first cousin and non-cousin marriages. These differences have been diminishing as the Tsimane have become more market integrated. The factors that influence preferences for cousin marriage appear to be less prevalent now than in the past, but cultural inertia might slow the pace of change in marriage norms. Overall, our findings suggest that cultural changes in marriage practices reflect underlying societal changes that shape the pool of potential spouses.

Socio-economic predictors of Inuit hunting choices and their implications for climate change adaptation.

In the Arctic, seasonal variation in the accessibility of the land, sea ice and open waters influences which resources can be harvested safely and efficiently. Climate stressors are also increasingly affecting access to subsistence resources. Within Inuit communities, people differ in their involvement with subsistence activities, but little is known about how engagement in the cash economy (time and money available) and other socio-economic factors shape the food production choices of Inuit harvesters, and their ability to adapt to rapid ecological change. We analyse 281 foraging trips involving 23 Inuit harvesters from Kangiqsujuaq, Nunavik, Canada using a Bayesian approach modelling both patch choice and within-patch success. Gender and income predict Inuit harvest strategies: while men, especially men from low-income households, often visit patches with a relatively low success probability, women and high-income hunters generally have a higher propensity to choose low-risk patches. Inland hunting, marine hunting and fishing differ in the required equipment and effort, and hunters may have to shift their subsistence activities if certain patches become less profitable or less safe owing to high costs of transportation or climate change (e.g. navigate larger areas inland instead of targeting seals on the sea ice). Our finding that household income predicts patch choice suggests that the capacity to maintain access to country foods depends on engagement with the cash economy. This article is part of the theme issue 'Climate change adaptation needs a science of culture'.

Gender disparities in material and educational resources differ by kinship system.

Contemporary inequality exists at an unprecedented scale. Social scientists have emphasized the role played by material wealth in driving its escalation. Evolutionary anthropologists understand the drive to accumulate material wealth as one that is coupled ultimately to increasing reproductive success. Owing to biological caps on reproduction for women, the efficiency of this conversion can differ by gender, with implications for understanding the evolution of gender disparities in resource accumulation. Efficiency also differs according to the type of resources used to support reproductive success. In this paper, we review evolutionary explanations of gender disparities in resources and investigate empirical evidence to support or refute those explanations among matrilineal and patrilineal subpopulations of ethnic Chinese Mosuo, who share an ethnolinguistic identity, but differ strikingly in terms of institutions and norms surrounding kinship and gender. We find that gender differentially predicts income and educational attainment. Men were more likely to report income than women; amounts earned were higher for men overall, but the difference between men and women was minimal under matriliny. Men reported higher levels of educational attainment than women, unexpectedly more so in matrilineal contexts. The results reveal nuances in how biology and cultural institutions affect gender disparities in wealth. This article is part of the theme issue 'Evolutionary ecology of inequality'.

The complex life course of mobility: Quantitative description of 300,000 residential moves in 1850-1950 Netherlands.

Mobility is a major mechanism of human adaptation, both in the deep past and in the present. Decades of research in the human evolutionary sciences have elucidated how much, how and when individuals and groups move in response to their ecology. Prior research has focused on small-scale subsistence societies, often in marginal environments and yielding small samples. Yet adaptive movement is commonplace across human societies, providing an opportunity to study human mobility more broadly. We provide a detailed, life-course structured demonstration, describing the residential mobility system of a historical population living between 1850 and 1950 in the industrialising Netherlands. We focus on how moves are patterned over the lifespan, attending to individual variation and stratifying our analyses by gender. We conclude that this population was not stationary: the median total moves in a lifetime were 10, with a wide range of variation and an uneven distribution over the life course. Mobility peaks in early adulthood (age 20-30) in this population, and this peak is consistent in all the studied cohorts, and both genders. Mobile populations in sedentary settlements provide a productive avenue for research on adaptive mobility and its relationship to human life history, and historical databases are useful for addressing evolutionarily motivated questions.

Inheritance, ecology and the evolution of the canoes of east Oceania.

We consider patterns in the evolution of canoe technology in the eastern Pacific relative to three general processes: movement of canoe traits along the Polynesian settlement sequence, adaptations to local island environment, and post-settlement interaction between island groups. Using model selection methods on the distributions of canoe technology, we show that social and ecological covariates together consistently outperform each considered individually, though knowledge of island area and post-settlement trading spheres does not add explanatory power. In particular, decorative canoe traits are not effectively explained by either our ecological or transmission models. We also estimate negative effects from both settlement sequence and island geomorphology, consistent with the die-off of particular canoe designs on resource-rich high island groups such as Hawaii and New Zealand. This decline in measured traits may be owing to the lifting of ecological constraints on population size or building materials.

Entropy trade-offs in artistic design: A case study of Tamil kolam.

From an evolutionary perspective, art presents many puzzles. Humans invest substantial effort in generating apparently useless displays that include artworks. These vary greatly from ordinary to intricate. From the perspective of signalling theory, these investments in highly complex artistic designs can reflect information about individuals and their social standing. Using a large corpus of kolam art from South India (N = 3139 kolam from 192 women), we test a number of hypotheses about the ways in which social stratification and individual differences affect the complexity of artistic designs. Consistent with evolutionary signalling theories of constrained optimisation, we find that kolam art tends to occupy a 'sweet spot' at which artistic complexity, as measured by Shannon information entropy, remains relatively constant from small to large drawings. This stability is maintained through an observable, apparently unconscious trade-off between two standard information-theoretic measures: richness and evenness. Although these drawings arise in a highly stratified, caste-based society, we do not find strong evidence that artistic complexity is influenced by the caste boundaries of Indian society. Rather, the trade-off is likely due to individual-level aesthetic preferences and differences in skill, dedication and time, as well as the fundamental constraints of human cognition and memory.

Limited Scope for Group Coordination in Stylistic Variations of Kolam Art.

In large, complex societies, assorting with others with similar social norms or behaviors can facilitate successful coordination and cooperation. The ability to recognize others with shared norms or behaviors is thus assumed to be under selection. As a medium of communication, human art might reflect fitness-relevant information on shared norms and behaviors of other individuals thus facilitating successful coordination and cooperation. Distinctive styles or patterns of artistic design could signify migration history, different groups with a shared interaction history due to spatial proximity, as well as individual-level expertise and preferences. In addition, cultural boundaries may be even more pronounced in a highly diverse and socially stratified society. In the current study, we focus on a large corpus of an artistic tradition called kolam that is produced by women from Tamil Nadu in South India (N = 3, 139 kolam drawings from 192 women) to test whether stylistic variations in art can be mapped onto caste boundaries, migration and neighborhoods. Since the kolam art system with its sequential drawing decisions can be described by a Markov process, we characterize variation in styles of art due to different facets of an artist's identity and the group affiliations, via hierarchical Bayesian statistical models. Our results reveal that stylistic variations in kolam art only weakly map onto caste boundaries, neighborhoods, and regional origin. In fact, stylistic variations or patterns in art are dominated by artist-level variation and artist expertise. Our results illustrate that although art can be a medium of communication, it is not necessarily marked by group affiliation. Rather, artistic behavior in this context seems to be primarily a behavioral domain within which individuals carve out a unique niche for themselves to differentiate themselves from others. Our findings inform discussions on the evolutionary role of art for group coordination by encouraging researchers to use systematic methods to measure the mapping between specific objects or styles onto groups.

Do wealth and inequality associate with health in a small-scale subsistence society?

In high-income countries, one's relative socio-economic position and economic inequality may affect health and well-being, arguably via psychosocial stress. We tested this in a small-scale subsistence society, the Tsimane, by associating relative household wealth (n = 871) and community-level wealth inequality (n = 40, Gini = 0.15-0.53) with a range of psychological variables, stressors, and health outcomes (depressive symptoms [n = 670], social conflicts [n = 401], non-social problems [n = 398], social support [n = 399], cortisol [n = 811], body mass index [n = 9,926], blood pressure [n = 3,195], self-rated health [n = 2523], morbidities [n = 1542]) controlling for community-average wealth, age, sex, household size, community size, and distance to markets. Wealthier people largely had better outcomes while inequality associated with more respiratory disease, a leading cause of mortality. Greater inequality and lower wealth were associated with higher blood pressure. Psychosocial factors did not mediate wealth-health associations. Thus, relative socio-economic position and inequality may affect health across diverse societies, though this is likely exacerbated in high-income countries.

Poverty is bad for health. People living in poverty are more likely to struggle to afford nutritious food, lack access to health care, or be overworked or stressed. This may make them susceptible to chronic diseases, contribute to faster aging, and shorten their lifespans. In high-income countries, there is growing evidence to suggest that a person's 'rank' in society also impacts their health. For example, individuals who have a lower position in the social hierarchy report worse health outcomes, regardless of their incomes. But it is unclear why living in an unequal society or having a lower social status contributes to poorer health. One possibility is that inequalities in society are creating a stressful environment that leads to worse physical and mental outcomes. It is thought that this stress largely comes from how humans evolved to prioritize reaching a higher social status over having a long and healthy life. If this is the case, this would mean that the link between social status and health would also be present in non-industrialized communities where social hierarchies tend to be less pronounced. To test this, Jaeggi, Blackwell et al. studied the Indigenous Tsimane population in Bolivia who live in small communities and forage and farm their own food. The income and relative wealth of 870 households from 40 Tsimane communities were compared against various outcomes, including symptoms associated with depression, stress hormone levels, blood pressure, self-rated health and several diseases. Jaeggi, Blackwell et al. found poverty and inequality did not negatively impact all of the health outcomes measured as has been previously reported for industrialized societies. However, blood pressure was higher among people with lower incomes or those who lived in more unequal communities. But because the Tsimane people generally have low blood pressure, the differences were too small to have much effect on their health. People who lived in more unequal communities were also three times more likely to have respiratory infections, but the reason for this was unclear. This shows that social determinants such as a person's wealth or inequality can affect health, even in communities with less rigid social hierarchies. In industrial societies the effect may be worse in part because they are compounded by lifestyle factors, such as diets rich in fat and sugar, and physical inactivity which can also increase blood pressure. This information may help policy makers reduce health disparities by addressing some of the social determinants of health and the lifestyle factors that cause them.

APOE4 is associated with elevated blood lipids and lower levels of innate immune biomarkers in a tropical Amerindian subsistence population.

In post-industrial settings, apolipoprotein E4 (APOE4) is associated with increased cardiovascular and neurological disease risk. However, the majority of human evolutionary history occurred in environments with higher pathogenic diversity and low cardiovascular risk. We hypothesize that in high-pathogen and energy-limited contexts, the APOE4 allele confers benefits by reducing innate inflammation when uninfected, while maintaining higher lipid levels that buffer costs of immune activation during infection. Among Tsimane forager-farmers of Bolivia (N = 1266, 50% female), APOE4 is associated with 30% lower C-reactive protein, and higher total cholesterol and oxidized LDL. Blood lipids were either not associated, or negatively associated with inflammatory biomarkers, except for associations of oxidized LDL and inflammation which were limited to obese adults. Further, APOE4 carriers maintain higher levels of total and LDL cholesterol at low body mass indices (BMIs). These results suggest that the relationship between APOE4 and lipids may be beneficial for pathogen-driven immune responses and unlikely to increase cardiovascular risk in an active subsistence population.

Genes contain the instructions needed for a cell to make molecules called proteins, which perform various roles in the body. Different variants of a gene can affect how the protein works, and in some cases, can increase a person's risk to develop certain diseases. For example, people who carry a version of the apolipoprotein E gene called APOE4 have a greater risk of developing Alzheimer's disease or heart disease. Individuals with two copies of this genetic variant have a 45% higher risk of heart disease and 12 times higher risk of Alzheimer's disease. Studies in industrialized countries suggest this increased risk may be the result of higher cholesterol and inflammation in people with APOE4. But if APOE4 is harmful, why does it continue to be so common worldwide? One potential explanation is that APOE4, which has been around since before modern humans, may be beneficial in some contexts. Cholesterol is essential for many vital tasks in the body. In physically demanding environments where parasitic infections are common ­ conditions similar to those experienced by early humans ­ APOE4 might be beneficial. Under those circumstances, having more cholesterol might help fuel metabolic activities, fight infections, or reduce inflammation caused by infections. Garcia et al. investigated the link between the APOE4 genetic variant, cholesterol and inflammation in 1,266 Indigenous Tsimane people from 80 villages in Bolivia. Tsimane people live an active lifestyle foraging and farming for food. Parasite infections are a common problem in their communities, but obesity rates are very low. Garcia et al. found that Tsimane people with at least one copy of the APOE4 have lower levels of inflammation and higher levels of cholesterol than those who have two copies of the APOE3 version of the gene. Very lean people with APOE4 had especially high levels of the so called "bad" low density lipoprotein (LDL) cholesterol compared to people with APOE3 only. However, in this situation, storing a little extra cholesterol may not be so bad. The findings contradict other studies that have linked obesity to higher LDL levels and APOE4 to higher levels of inflammation. For the majority of human history, humans lived in more physically strenuous and calorically restrictive environments, with less access to clean water. Garcia et al. suggest that the harmful effects of APOE4 seen in studies in more industrialized societies ­ where people tend to be more sedentary and have less exposure to pathogens ­ may reflect a mismatch between a person's environment and their genes. More studies that capture the diversity of environmental conditions under which people live will help clarify the role of APOE4 health and disease.

Differences between sons and daughters in the intergenerational transmission of wealth.

Persistent interest lies in gender inequality, especially with regard to the favouring of sons over daughters. Economists are concerned with how privilege is transmitted across generations, and anthropologists have long studied sex-biased inheritance norms. There has, however, been no focused cross-cultural investigation of how parent-offspring correlations in wealth vary by offspring sex. We estimate these correlations for 38 wealth measures, including somatic and relational wealth, from 15 populations ranging from hunter-gatherers to small-scale farmers. Although small sample sizes limit our statistical power, we find no evidence of ubiquitous male bias, at least as inferred from comparing parent-son and parent-daughter correlations. Rather we find wide variation in signatures of sex bias, with evidence of both son and daughter-biased transmission. Further, we introduce a model that helps pinpoint the conditions under which simple mid-point parent-offspring wealth correlations can reveal information about sex-biased parental investment. Our findings are relevant to the study of female-biased kinship by revealing just how little normative descriptors of kinship systems, such as patrilineal inheritance, capture intergenerational correlations in wealth, and how variable parent-son and parent-daughter correlations can be. This article is part of the theme issue 'The evolution of female-biased kinship in humans and other mammals'.

Reciprocal Exchange Patterned by Market Forces Helps Explain Cooperation in a Small-Scale Society.

Social organisms sometimes depend on help from reciprocating partners to solve adaptive problems [1], and individual cooperation strategies should aim to offer high supply commodities at low cost to the donor in exchange for high-demand commodities with large return benefits [2, 3]. Although such market dynamics have been documented in some animals [4-7], naturalistic studies of human cooperation are often limited by focusing on single commodities [8]. We analyzed cooperation in five domains (meat sharing, produce sharing, field labor, childcare, and sick care) among 2,161 household dyads of Tsimane' horticulturalists, using Bayesian multilevel models and information-theoretic model comparison. Across domains, the best-fit models included kinship and residential proximity, exchanges in kind and across domains, measures of supply and demand and their interactions with exchange, and household-specific exchange slopes. In these best models, giving, receiving, and reciprocating were to some extent shaped by market forces, and reciprocal exchange across domains had a strong partial effect on cooperation independent of more exogenous factors like kinship and proximity. Our results support the view that reciprocal exchange can provide a reliable solution to adaptive problems [8-11]. Although individual strategies patterned by market forces may generate gains from trade in any species [3], humans' slow life history and skill-intensive foraging niche favor specialization and create interdependence [12, 13], thus stabilizing cooperation and fostering divisions of labor even in informal economies [14, 15].

Understanding the nature of wealth and its effects on human fitness.

Studying fitness consequences of variable behavioural, physiological and cognitive traits in contemporary populations constitutes the specific contribution of human behavioural ecology to the study of human diversity. Yet, despite 30 years of evolutionary anthropological interest in the determinants of fitness, there exist few principled investigations of the diverse sources of wealth that might reveal selective forces during recent human history. To develop a more holistic understanding of how selection shapes human phenotypic traits, be these transmitted by genetic or cultural means, we expand the conventional focus on associations between socioeconomic status and fitness to three distinct types of wealth-embodied, material and relational. Using a model selection approach to the study of women's success in raising offspring in an African horticultural population (the Tanzanian Pimbwe), we find that the top performing models consistently include relational and material wealth, with embodied wealth as a less reliable predictor. Specifically, child mortality risk is increased with few household assets, parent nonresidency, child legitimacy, and one or more parents having been accused of witchcraft. The use of multiple models to test various hypotheses greatly facilitates systematic comparative analyses of human behavioural diversity in wealth accrual and investment across different kinds of societies.