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.

Maternal and infant predictors of proinflammatory milk immune activity in Kilimanjaro, Tanzania.

OBJECTIVES: The immune system of milk (ISOM) creates a mother-infant immune axis that plays an important role in protecting infants against infectious disease (ID). Tradeoffs in the immune system suggest the potential for both protection and harm, so we conceive of two dimensions via which the ISOM impacts infants: promotion of protective activity and control of activity directed at benign targets. High variability in ISOM activity across mother-infant dyads suggests investment the ISOM may have evolved to be sensitive to maternal and/or infant characteristics. We assessed predictors of appropriate and misdirected proinflammatory ISOM activity in an environment of high ID risk, testing predictions drawn from life history theory and other evolutionary perspectives. METHODS: We characterized milk in vitro interleukin-6 (IL-6) responses to Salmonella enterica (a target of protective immune activity; N = 96) and Escherichia coli (a benign target; N = 85) among mother-infant dyads in rural Kilimanjaro, Tanzania. We used ordered logistic regression and mixture models to evaluate maternal and infant characteristics as predictors of IL-6 responses. RESULTS: In all models, IL-6 responses to S. enterica increased with maternal age and decreased with gravidity. In mixture models, IL-6 responses to E. coli declined with maternal age and increased with gravidity. No other considered variables were consistently associated with IL-6 responses. CONCLUSIONS: The ISOM's capacities for appropriate proinflammatory activity and control of misdirected proinflammatory activity increases with maternal age and decreases with gravidity. These findings are consistent with the hypothesis that the mother-infant immune axis has evolved to respond to maternal life history characteristics.

Does the immune system of milk increase activity for infants experiencing infectious disease episodes in Kilimanjaro, Tanzania?

INTRODUCTION: Multiple studies have reported that milk immune content increases for infants experiencing infectious disease (ID) episodes, suggesting that the immune system of milk (ISOM) offers enhanced protection when needed to combat ID. METHODS: To test the hypothesis that ISOM content and/or activity increases during an infant's ID episode, we characterized milk secretory immunoglobulin A (sIgA; a major ISOM constituent) and in vitro interleukin-6 (IL-6) responses to Salmonella enterica and Escherichia coli, as system-level biomarkers of ISOM activity, in a prospective study among 96 mother-infant dyads in Kilimanjaro, Tanzania. RESULTS: After control for covariates, no milk immune variables (sIgA, Coef: 0.03; 95% CI -0.25, 0.32; in vitro IL-6 response to S. enterica, Coef: 0.23; 95% CI: -0.67, 1.13; IL-6 response to E. coli, Coef: -0.11; 95% CI: -0.98, 0.77) were associated with prevalent ID (diagnosed at the initial participation visit). Among infants experiencing an incident ID (diagnosed subsequent to the initial participation), milk immune content and responses were not substantially higher or lower than the initial visit (sIgA, N: 61; p: 0.788; IL-6 response to S. enterica, N: 56; p: 0.896; IL-6 response to E. coli, N: 36; p: 0.683); this was unchanged by exclusion of infants with ID at the time of initial participation. CONCLUSION: These findings are not consistent with the hypothesis that milk delivers enhanced immune protection when infants experience ID. In environments with a high burden of ID, dynamism may be less valuable to maternal reproductive success than stability in the ISOM.

Matriliny reverses gender disparities in inflammation and hypertension among the Mosuo of China.

Women experience higher morbidity than men, despite living longer. This is often attributed to biological differences between the sexes; however, the majority of societies in which these disparities are observed exhibit gender norms that favor men. We tested the hypothesis that female-biased gender norms ameliorate gender disparities in health by comparing gender differences in inflammation and hypertension among the matrilineal and patrilineal Mosuo of China. Widely reported gender disparities in health were reversed among matrilineal Mosuo compared with patrilineal Mosuo, due to substantial improvements in women's health, with no concomitant detrimental effects on men. These findings offer evidence that gender norms limiting women's autonomy and biasing inheritance toward men adversely affect the health of women, increasing women's risk for chronic diseases with tremendous global health impact.

Labor migration is associated with lower rates of underweight and higher rates of obesity among left-behind wives in rural Bangladesh: a cross-sectional study.

BACKGROUND: Among Bangladeshi men, international labor migration has increased ten-fold since 1990 and rural to urban labor migration rates have steadily increased. Labor migration of husbands has increased household wealth and redefined women's roles, which have both positively and negatively impacted the health of wives "left behind". We examined the direct and indirect effects of husband labor migration on chronic disease indicators and outcomes among wives of labor migrants. METHODS: We collected survey, anthropometric, and biomarker data from a random sample of women in Matlab, Bangladesh, in 2018. We assessed associations between husband's migration and indicators of adiposity and chronic disease. We used structural equation modeling to assess the direct effect of labor migration on chronic disease, undernutrition, and adiposity, and the mediating roles of income, food security, and proportion of food purchased from the bazaar. Qualitative interviews and participant observation were used to help provide context for the associations we found in our quantitative results. FINDINGS: Among study participants, 9.0% were underweight, 50.9% were iron deficient, 48.3% were anemic, 39.6% were obese, 27.3% had a waist circumference over 35 in., 33.1% had a high whole-body fat percentage, 32.8% were diabetic, and 32.9% had hypertension. Slightly more women in the sample (55.3%) had a husband who never migrated than had a husband who had ever migrated (44.9%). Of those whose husband had ever migrated, 25.8% had a husband who was a current international migrant. Wives of migrants were less likely to be underweight, and more likely to have indicators of excess adiposity, than wives of non-migrants. Protection against undernutrition was attributable primarily to increased food security among wives of migrants, while increased adiposity was attributable primarily to purchasing a higher proportion of food from the bazaar; however, there was a separate path through income, which qualitative findings suggest may be related to reduced physical activity. CONCLUSIONS: Labor migration, and particularly international labor migration, intensifies the nutrition transition in Bangladesh through increasing wealth, changing how foods are purchased, and reducing physical activity, which both decreases risk for undernutrition and increases risk for excess adiposity.

High-altitude adaptations mitigate risk for hypertension and diabetes-associated anemia.

BACKGROUND: Human populations native to high altitude exhibit numerous genetic adaptations to hypobaric hypoxia. Among Tibetan plateau peoples, these include increased vasodilation and uncoupling of erythropoiesis from hypoxia. OBJECTIVE/METHODS: We tested the hypothesis that these high-altitude adaptations reduce risk for hypertension and diabetes-associated anemia among the Mosuo, a Tibetan-descended population in the mountains of Southwest China that is experiencing rapid economic change and increased chronic disease risk. RESULTS: Hypertension was substantially less common among Mosuo than low-altitude Han populations, and models fit to the Han predicted higher probability of hypertension than models fit to the Mosuo. Diabetes was positively associated with anemia among the Han, but not the Mosuo. CONCLUSION: The Mosuo have lower risk for hypertension and diabetes-associated anemia than the Han, supporting the hypothesis that high-altitude adaptations affecting blood and circulation intersect with chronic disease processes to lower risk for these outcomes. As chronic diseases continue to grow as global health concerns, it is important to investigate how they may be affected by local genetic adaptations.

The evolution of inequality.

Understanding how systems of political and economic inequality evolved from relatively egalitarian origins has long been a focus of anthropological inquiry. Many hypotheses have been suggested to link socio-ecological features with the rise and spread of inequality, and empirical tests of these hypotheses in prehistoric and extant societies are increasing. In this review, we synthesize several streams of theory relevant to understanding the evolutionary origins, spread, and adaptive significance of inequality. We argue that while inequality may be produced by a variety of localized processes, its evolution is fundamentally dependent on the economic defensibility and transmissibility of wealth. Furthermore, these properties of wealth could become persistent drivers of inequality only following a shift to a more stable climate in the Holocene. We conclude by noting several key areas for future empirical research, emphasizing the need for more analyses of contemporary shifts toward institutionalized inequality as well as prehistoric cases.

Breastfeeding over two years is associated with longer birth intervals, but not measures of growth or health, among children in Kilimanjaro, TZ.

OBJECTIVES: Breastfeeding has been associated with numerous health and well-being benefits for both children and their mothers, including prolonging the birth interval to the subsequent sibling. The clearest associations between breastfeeding and health outcomes, per se, reflect exclusive breastfeeding in the first months of postnatal life and are most evident during infancy. Fewer studies explore the consequences of breastfeeding for multiple years. In this article, we ask whether breastfeeding for more than 2 years is associated with discernible health and well-being benefits to children. METHODS: Data were collected from 315 children, aged 2 to 7, and their caretakers residing in Kilimanjaro, Tanzania. Basic demographic and health information was solicited, and anthropometric and blood markers of health were evaluated. RESULTS: Our results indicate a strong positive relationship between breastfeeding for 2 or more years and interbirth interval, but little evidence for a relationship between prolonged breastfeeding and several indicators of child growth and health. CONCLUSIONS: We suggest that these relationships may support the recently rekindled birth spacing hypothesis, positing selection for longer interbirth intervals, rather than, or in addition to, more direct health benefits associated with breastfeeding for 2 or more years. Our results may indicate attenuating health benefits associated with longer breastfeeding.

Palmar Creases: Classification, Reliability and Relationships to Fetal Alcohol Spectrum Disorders (FASD).

A normal human palm contains 3 major creases: the distal transverse crease; the proximal transverse crease; and the thenar crease. Because permanent crease patterns are thought to be laid down during the first trimester, researchers have speculated that deviations in crease patterns could be indicative of insults during fetal development. The purpose of this study was twofold: (1) to compare the efficacy and reliability of two coding methods, the first (M1) classifying both "simiana" and Sydney line variants and the second (M2) counting the total number of crease points of origin on the radial border of the hand; and (2) to ascertain the relationship between palmar crease patterns and fetal alcohol spectrum disorders (FASD). Bilateral palm prints were taken using the carbon paper and tape method from 237 individuals diagnosed with FASD and 190 unexposed controls. All prints were coded for crease variants under M1 and M2. Additionally, a random sample of 98 matched (right and left) prints was selected from the controls to determine the reliabilities of M1 and M2. For this analysis, each palm was read twice, at different times, by two readers. Intra-observer Kappa coefficients were similar under both methods, ranging from 0.804-0.910. Inter-observer Kappa coefficients ranged from 0.582-0.623 under M1 and from 0.647-0.757 under M2. Using data from the entire sample of 427 prints and controlling for sex and ethnicity (white v. non-white), no relationship was found between palmar crease variants and FASD. Our results suggest that palmar creases can be classified reliably, but palmar crease patterns may not be affected by fetal alcohol exposure.

Correction: Betel quid use is associated with anemia among both men and women in Matlab, Bangladesh.

[This corrects the article DOI: 10.1371/journal.pgph.0001677.].

The evolutionary ecology of early weaning in Kilimanjaro, Tanzania.

Public health recommendations promote prolonged breastfeeding of all children; however, parental investment (PI) theory predicts that breastfeeding will be allocated among a mothers' offspring to maximize her reproductive success. We evaluated PI in terms of risk for weaning before age two among 283 children in Kilimanjaro, Tanzania. Results demonstrate: (i) a Trivers-Willard effect--high socioeconomic status (SES) females and low SES males were more likely to be weaned early; (ii) later-born children were less likely to be weaned early; (iii) higher birthweight children were less likely to be weaned early, and (iv) no effect of cattle (a source of supplementary milk) ownership. These associations were largely independent and remained significant in models controlling for potential confounders; however, the inverse association between early weaning and birth order lost significance in the model containing birthweight. These patterns were observed despite public health recommendations encouraging breastfeeding for at least two years.

Does gender structure social networks across domains of cooperation? An exploration of gendered networks among matrilineal and patrilineal Mosuo.

Cooperative networks are essential features of human society. Evolutionary theory hypothesizes that networks are used differently by men and women, yet the bulk of evidence supporting this hypothesis is based on studies conducted in a limited range of contexts and on few domains of cooperation. In this paper, we compare individual-level cooperative networks from two communities in Southwest China that differ systematically in kinship norms and institutions-one matrilineal and one patrilineal-while sharing an ethnic identity. Specifically, we investigate whether network structures differ based on prevailing kinship norms and type of gendered cooperative activity, one woman-centred (preparation of community meals) and one man-centred (farm equipment lending). Our descriptive results show a mixture of 'feminine' and 'masculine' features in all four networks. The matrilineal meals network stands out in terms of high degree skew. Exponential random graph models reveal a stronger role for geographical proximity in patriliny and a limited role of affinal relatedness across all networks. Our results point to the need to consider domains of cooperative activity alongside gender and cultural context to fully understand variation in how women and men leverage social relationships toward different ends. This article is part of the theme issue 'Cooperation among women: evolutionary and cross-cultural perspectives'.

Market integration, income inequality, and kinship system among the Mosuo of China.

Increased access to defensible material wealth is hypothesised to escalate inequality. Market integration, which creates novel opportunities in cash economies, provides a means of testing this hypothesis. Using demographic data collected from 505 households among the matrilineal and patrilineal Mosuo in 2017, we test whether market integration is associated with increased material wealth, whether increased material wealth is associated with wealth inequality, and whether being in a matrilineal vs. patrilineal kinship system alters the relationship between wealth and inequality. We find evidence that market integration, measured as distance to the nearest source of tourism and primary source of household income, is associated with increased household income and 'modern' asset value. Both village-level market integration and mean asset value were associated negatively, rather than positively, with inequality, contrary to predictions. Finally, income, modern wealth and inequality were higher in matrilineal communities that were located closer to the centre of tourism and where tourism has long provided a relatively stable source of income. However, we also observed exacerbated inequality with increasing farm animal value in patriliny. We conclude that the forces affecting wealth and inequality depend on local context and that the importance of local institutions is obscured by aggregate statistics drawn from modern nation states.

Betel quid use is associated with anemia among both men and women in Matlab, Bangladesh.

Anemia accounts for 8.8% of total disability burden worldwide. Betel quid use among pregnant women has been found to increase anemia risk. Betel quid is prepared by wrapping the betel (or areca) nut, with spices and other additions, in betel or tobacco leaf and it is chewed or placed in the mouth. We explored the association between betel quid use and anemia among men and non-pregnant women. We collected data from a random sample of women and their husbands in Matlab, Bangladesh. Participants reported their current betel quid use and individual characteristics. We assessed hemoglobin (a biomarker of anemia) with a hemoglobinometer and soluble transferrin receptor (a biomarker of iron deficiency) and C-reactive protein (a biomarker of inflammation) in dried blood spots via enzyme immunoassay. We estimated logistic regression models to evaluate the association between betel quid use and anemia and structural equation models (SEM) to evaluate mediating roles of iron deficiency and elevated inflammation. A total of 1133 participants (390 men and 743 non-pregnant women) were included. After controlling for important confounders, any betel quid use was positively associated with anemia among men (OR: 1.80; 95% CI: 1.12, 2.89). Among women, betel quid use was associated with anemia only among the most frequent users (OR: 1.62; 95% CI: 1.03, 2.53). SEM did not reveal indirect paths through inflammation or iron deficiency. Betel quid use may contribute to the burden of anemia among adults in Bangladesh. Our findings suggest the burden of disease attributed to betel quid use has been underestimated.

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'.

Tradeoffs in milk immunity affect infant infectious disease risk.

Background and objectives: The human immune system has evolved to balance protection against infection with control of immune-mediated damage and tolerance of commensal microbes. Such tradeoffs between protection and harm almost certainly extend to the immune system of milk. Methodology: Among breastfeeding mother-infant dyads in Kilimanjaro, Tanzania, we characterized in vitro proinflammatory milk immune responses to Salmonella enterica (an infectious agent) and Escherichia coli (a benign target) as the increase in interleukin-6 after 24 h of incubation with each bacterium. We characterized incident infectious diseases among infants through passive monitoring. We used Cox proportional hazards models to describe associations between milk immune activity and infant infectious disease. Results: Among infants, risk for respiratory infections declined with increasing milk in vitro proinflammatory response to S. enterica (hazard ratio [HR]: 0.68; 95% confidence interval [CI]: 0.54, 0.86; P: 0.001), while risk for gastrointestinal infections increased with increasing milk in vitro proinflammatory response to E. coli (HR: 1.44; 95% CI: 1.05, 1.99; P: 0.022). Milk proinflammatory responses to S. enterica and E. coli were positively correlated (Spearman's rho: 0.60; P: 0.000). Conclusions and implications: These findings demonstrate a tradeoff in milk immune activity: the benefits of appropriate proinflammatory activity come at the hazard of misdirected proinflammatory activity. This tradeoff is likely to affect infant health in complex ways, depending on prevailing infectious disease conditions. How mother-infant dyads optimize proinflammatory milk immune activity should be a central question in future ecological-evolutionary studies of the immune system of milk.

Social support, nutrition and health among women in rural Bangladesh: complex tradeoffs in allocare, kin proximity and support network size.

Malnutrition among women of reproductive age is a significant public health concern in low- and middle-income countries. Of particular concern are undernutrition from underweight and iron deficiency, along with overweight and obesity, all of which have negative health consequences for mothers and children. Accumulating evidence suggests that risk for poor nutritional outcomes may be mitigated by social support, yet how social support is measured varies tremendously and its effects likely vary by age, kinship and reproductive status. We examine the effects of different measures of social support on weight and iron nutrition among 677 randomly sampled women from rural Bangladesh. While we find that total support network size mitigates risk for underweight, other results point to a potential tradeoff in the effects of kin proximity, with nearby adult children associated with both lower risk for underweight and obesity and higher risk for iron deficiency and anaemia. Social support from kin may then enhance energy balance but not diet quality. Results also suggest that a woman's network of caregivers might reflect their greater need for help, as those who received more help with childcare and housework had worse iron nutrition. Overall, although some findings support the hypothesis that social support can be protective, others emphasize that social relationships often have neutral or negative effects, illustrating the kinds of tradeoffs expected from an evolutionary perspective. The complexities of these effects deserve attention in future work, particularly within public health, where what is defined as 'social support' is often assumed to be positive. This article is part of the theme issue 'Multidisciplinary perspectives on social support and maternal-child health'.

Patterns of paternal investment predict cross-cultural variation in jealous response.

Long-lasting, romantic partnerships are a universal feature of human societies, but almost as ubiquitous is the risk of instability when one partner strays. Jealous response to the threat of infidelity is well studied, but most empirical work on the topic has focused on a proposed sex difference in the type of jealousy (sexual or emotional) that men and women find most upsetting, rather than on how jealous response varies1,2. This stems in part from the predominance of studies using student samples from industrialized populations, which represent a relatively homogenous group in terms of age, life history stage and social norms3,4. To better understand variation in jealous response, we conducted a 2-part study in 11 populations (1,048 individuals). In line with previous work, we find a robust sex difference in the classic forced-choice jealousy task. However, we also show substantial variation in jealous response across populations. Using parental investment theory, we derived several predictions about what might trigger such variation. We find that greater paternal investment and lower frequency of extramarital sex are associated with more severe jealous response. Thus, partner jealousy appears to be a facultative response, reflective of the variable risks and costs of men's investment across societies.

The expendable male hypothesis.

Matriliny is a system of kinship in which descent and inheritance are conferred along the female line. The theoretically influential concept of the matrilineal puzzle posits that matriliny poses special problems for understanding men's roles in matrilineal societies. Ethnographic work describes the puzzle as the tension experienced by men between the desire to exert control over their natal kin (i.e. the lineage to which they belong) and over their affinal kin (i.e. their spouses and their biological children). Evolutionary work frames the paradox as one resulting from a man investing in his nieces and nephews at the expense of his own biological offspring. In both cases, the rationale for the puzzle rests on two fundamental assumptions: (i) that men are in positions of authority over women and over resources; and (ii) that men are interested in the outcomes of parenting. In this paper, we posit a novel hypothesis that suggests that certain ecological conditions render men expendable within local kinship configurations, nullifying the above assumptions. This arises when (i) women, without significant assistance from men, are capable of meeting the subsistence needs of their families; and (ii) men have little to gain from parental investment in children. We conclude that the expendable male hypothesis may explain the evolution of matriliny in numerous cases, and by noting that female-centred approaches that call into doubt assumptions inherent to male-centred models of kinship are justified in evolutionary perspective. This article is part of the theme issue 'The evolution of female-biased kinship in humans and other mammals'.