Adipose development is consistent across hunter-gatherers and diverges from western references.

Despite agreement that humans have evolved to be unusually fat primates, adipose patterning among hunter-gatherers has received little empirical consideration. Here we consider the development of adiposity among four contemporary groups of hunter-gatherers, the Aka, Savanna Pumé, Ju'/Hoansi and Agta using multi-level generalized additive mixed modelling to characterize the growth of tricep skinfolds from early childhood through adolescence. In contrast to references, hunter-gatherers show several consistent patterns: (i) children are lean with little fat accumulation; (ii) no adiposity rebound at 5 years is evident; (iii) girls on average have built 90% of their body size, and reach menarche when adiposity is at its maximum velocity; and (iv) a metabolic trade-off is evident in young, but not older children, such that both boys and girls prioritize skeletal growth during middle childhood, a trade-off that diminishes during adolescence when height velocity increases in pace with fat accumulation. Consistent results across hunter-gatherers living in diverse environments suggest that these patterns reflect a general forager pattern of development. The findings provide a valuable baseline for adipose development not apparent from reference populations. We emphasize both generalized trends among hunter-gatherers, and that inter-populational differences point to the plasticity with which humans organize growth and development.

Uncertainty in a globalizing world. Livelihood and fertility variance increases in response to rapid change.

OBJECTIVES: The extreme condition that we address in this special issue is how people adapt to rapid change, which in this case study is instigated by globalization and the process of market integration. Although market integration has been underway for centuries in some parts of the world, it often occurs precipitously in small-scale societies, initiating an abrupt break with traditional ways of life and fostering a keen sense of uncertainty. METHODS: Using cross sections from 30-years of data collected in a Yucatec Maya subsistence farming community, we test the expectation that when payoffs to pursue new livelihood and reproductive options are uncertain, variance in social, economic, and reproductive traits will increase in the population. Our data span the transition from subsistence farming to a mixed economy, and bridge the transition from natural to contracepting fertility. Exposure to globalizing and market forces occurred when a paved road was built in the early 2000s. RESULTS: We find that livelihood traits (a household's primary economic strategy, amount of land under cultivation, amount of maize and honey sold), become more variable as new, but uncertain options become available. Variance in levels of education and family size likewise immediately increase following the road, but show signs of settling back down a decade later. Rather than replacing one way of life with another, Maya farmers conservatively adopt some new elements (family planning, wage labor), until the tradeoffs to commit to smaller families and the labor market become clearer. CONCLUSION: Our findings highlight that in rapidly changing environments when the payoffs to assimilate new options are uncertain, some households and individuals intensify what they know best, while others adopt new opportunities, driving variance up in the population.

Sex differences in adipose development in a hunter-gatherer population.

OBJECTIVE: Humans are unusually sexually dimorphic in body composition, with adult females having on average nearly twice the fat mass as males. The development of adipose sex differences has been well characterized for children growing up in food-abundant environments, with less known about cross-cultural variation, particularly in populations without exposure to market foods, mechanized technologies, schooling, vaccination, or other medical interventions. METHODS: To add to the existing cross-cultural data, we fit multiple growth curves to body composition and anthropometric data to describe adipose development for the Savanna Pumé, South American hunter-gatherers. RESULTS: (1) Little evidence is found for an adiposity 'rebound' at the end of early childhood among either Savanna Pumé girls or boys. (2) Rather, fat deposition fluctuates during childhood, from age ~4 to ~9 years, with no appreciable accumulation until the onset of puberty, a pattern also observed among Congo Baka hunter-gatherers. (3) Body fat fluctuations are more pronounced for girls than boys. (4) The age of peak skeletal, weight, and adipose gains are staggered to a much greater extent among the Savanna Pumé compared to the National Health and Nutrition Examination Survey (NHANES III) reference, suggesting this is an important developmental strategy in lean populations. CONCLUSION: Documenting growth patterns under diverse preindustrial energetic conditions provides an important baseline for understanding sex differences in body fat emerging today under food abundance.

Balancing fertility and livelihood diversity in mixed economies.

Mixed economies provide a unique context for testing theories of fertility change. Because they have a stake in two traditions, mixed-economy households balance the demands of both a labor-based subsistence economy, which benefits from a large family, and a wage-labor economy, which benefits from reduced fertility. Additionally, household size changes over the course of its life-cycle and shapes available economic opportunities. Here we argue that in mixed economies, fertility may reflect opportunities for livelihood diversity rather than simply responding to the restricted socioeconomic benefits of small families. While low fertility may in some cases have an economic benefit, low fertility can also limit the livelihood diversity of a household which is a key strategy for long-term economic success. We test this prediction with longitudinal data from a Maya community undergoing both a sustained decline in fertility and rapid integration into the market economy. Using household-level fertility, number of adults, and livelihood diversity at two time points, we find that household size is positively related to livelihood diversity, which in turn is positively related to household income per-capita. However, household size also has a negative association with income per capita. The results reflect a balancing act whereby households attempt to maximize the economic diversity with as few members as possible. Broadly, these results suggest that theories of fertility decline must account for how households pool resources and diversify economic activities in the face of increasing market integration, treating fertility as both an outcome and an input into economic and reproductive decision-making.

Disentangling basal and accrued height-for-age for cross-population comparisons.

OBJECTIVES: Current standards for comparing stunting across human populations assume a universal model of child growth. Such comparisons ignore population differences that are independent of deprivation and health outcomes. This article partitions variation in height-for-age that is specifically associated with deprivation and health outcomes to provide a basis for cross-population comparisons. MATERIALS AND METHODS: Using a multilevel model with a sigmoid relationship of resources and growth, we partition variation in height-for-age z-scores (HAZ) from 1.5 million children across 70 countries into two components: (1) "accrued HAZ" shaped by environmental inputs (e.g., undernutrition, infectious disease, inadequate sanitation, poverty) and (2) a country-specific "basal HAZ" independent of such inputs. We validate these components against population-level infant mortality rates and assess how these basal differences may affect cross-population comparisons of stunting. RESULTS: Basal HAZ differs reliably across countries (range of approximately 1.5 SD) and is independent of measures of infant mortality. By contrast, accrued HAZ captures stunting as impaired growth due to deprivation and is more closely associated with infant mortality than observed HAZ. Assessing stunting prevalence by accrued HAZ suggest that populations in West Africa and Haiti suffer much greater levels of stunting than indicated by observed HAZ. DISCUSSION: Current universal standards may dramatically underestimate stunting in populations with taller basal HAZ. Relying on observed HAZ rather than accrued HAZ may also lead to inappropriate cross-population comparisons, such as concluding that Haitian children enjoy better conditions for growth than do Indian or Guatemalan children.

Identifying the limits to socioeconomic influences on human growth.

Contemporary humans occupy the widest range of socioeconomic environments in their evolutionary history, and this has revealed unprecedented environmentally-induced plasticity in physical growth. This plasticity also has limits, and identifying those limits can help researchers: (1) parse when population differences arise from environmental inputs or not and (2) determine when it is possible to infer socioeconomic disparities from disparities in body form. To illustrate potential limits to environmental plasticity, we analyze body mass index (BMI) and height data from 1,768,962 women and 207,341 men (20-49 y) living in households exhibiting 1000-fold variation in household wealth (51 countries, 1985-2017, 164 surveys) across four world regions-sub-Saharan Africa, South Asia, Latin America, and North Africa and the Middle East. We find that relationships of environmental inputs with both mean height and BMI bottom out at roughly 100-700 USD per capita household wealth (2011 international units, PPP), but at different basal BMIs and basal heights for different regions. The relationship with resources tops out for BMI at around 20 K-35 K USD for women, with growth potential due to environmental inputs in the range of 6.2-8.4 kg/m2. By contrast, mean BMI for men and mean height for both sexes remains sensitive to environmental inputs even at levels far above the low- and middle-income samples studied here. This suggest that further work integrating comparable data from low- and high-income samples should provide a better picture of the full range of environmental inputs on human height and BMI. We conclude by discussing how neglecting such population-specific limits to human growth can lead to erroneous inferences about population differences.