Jonathan Stieglitz

From the womb to the tomb: The role of transfers in shaping the evolved human life history

Humans are the longest living and slowest growing of all primates. Although most primates are social, humans are highly cooperative and social in ways that likely co-evolved with the slow human life history. In this paper we highlight the role of resource transfers and non-material assistance within and across generations in shaping low human mortality rates. The use of complex cooperative strategies to minimize risk is a necessary precursor for selecting further reductions in mortality rate in late adulthood. In conjunction with changes in the age-profile of production, the impacts of resource transfers and other forms of cooperation on reducing mortality likely played an important role in selection on post-reproductive lifespan throughout human evolution. Using medical data and ethnographic interviews, we explore several types of common risks experienced by Tsimane forager-horticulturalists, and quantify the types and targets of aid. Our results illustrate the importance of transfers in several key domains and suggest that the absence of transfers would greatly increase human mortality rates throughout the life course.

Does Blood Pressure Inevitably Rise With Age?: Longitudinal Evidence Among Forager-Horticulturalists

The rise in blood pressure with age is a major risk factor for cardiovascular and renal disease, stroke, and type 2 diabetes mellitus. Age-related increases in blood pressure have been observed in almost every population, except among hunter-gatherers, farmers, and pastoralists. Here we tested for age-related increases in blood pressure among Tsimane forager-farmers. We also test whether lifestyle changes associated with modernization lead to higher blood pressure and a greater rate of age-related increase in blood pressure. We measured blood pressure longitudinally on 2248 adults age ≥20 years (n=6468 observations over 8 years). Prevalence of hypertension was 3.9% for women and 5.2% for men, although diagnosis of persistent hypertension based on multiple observations reduced prevalence to 2.9% for both sexes. Mixed-effects models revealed systolic, diastolic, and pulse blood pressure increases of 2.86 (P<0.001), 0.95 (P<0.001), and 1.95 mmHg (P<0.001) per decade for women and 0.91 (P<0.001), 0.93 (P<0.001), and −0.02 mmHg (P=0.93) for men, substantially lower than rates found elsewhere. Lifestyle factors, such as smoking and Spanish fluency, had minimal effect on mean blood pressure and no effect on age-related increases in blood pressure. Greater town proximity was associated with a lower age-related increase in pulse pressure. Effects of modernization were, therefore, deemed minimal among Tsimane, in light of their lean physique, active lifestyle, and protective diet.The rise in blood pressure with age is a major risk factor for cardiovascular and renal disease, stroke, and type 2 diabetes mellitus. Age-related increases in blood pressure have been observed in almost every population, except among hunter-gatherers, farmers, and pastoralists. Here we tested for age-related increases in blood pressure among Tsimane forager-farmers. We also test whether lifestyle changes associated with modernization lead to higher blood pressure and a greater rate of age-related increase in blood pressure. We measured blood pressure longitudinally on 2248 adults age ≥20 years (n=6468 observations over 8 years). Prevalence of hypertension was 3.9% for women and 5.2% for men, although diagnosis of persistent hypertension based on multiple observations reduced prevalence to 2.9% for both sexes. Mixed-effects models revealed systolic, diastolic, and pulse blood pressure increases of 2.86 ( P <0.001), 0.95 ( P <0.001), and 1.95 mmHg ( P <0.001) per decade for women and 0.91 ( P <0.001), 0.93 ( P <0.001), and −0.02 mmHg ( P =0.93) for men, substantially lower than rates found elsewhere. Lifestyle factors, such as smoking and Spanish fluency, had minimal effect on mean blood pressure and no effect on age-related increases in blood pressure. Greater town proximity was associated with a lower age-related increase in pulse pressure. Effects of modernization were, therefore, deemed minimal among Tsimane, in light of their lean physique, active lifestyle, and protective diet. # Novelty and Significance {#article-title-30}

Coronary atherosclerosis in indigenous South American Tsimane: a cross-sectional cohort study

BACKGROUND Conventional coronary artery disease risk factors might potentially explain at least 90% of the attributable risk of coronary artery disease. To better understand the association between the pre-industrial lifestyle and low prevalence of coronary artery disease risk factors, we examined the Tsimane, a Bolivian population living a subsistence lifestyle of hunting, gathering, fishing, and farming with few cardiovascular risk factors, but high infectious inflammatory burden. METHODS We did a cross-sectional cohort study including all individuals who self-identified as Tsimane and who were aged 40 years or older. Coronary atherosclerosis was assessed by coronary artery calcium (CAC) scoring done with non-contrast CT in Tsimane adults. We assessed the difference between the Tsimane and 6814 participants from the Multi-Ethnic Study of Atherosclerosis (MESA). CAC scores higher than 100 were considered representative of significant atherosclerotic disease. Tsimane blood lipid and inflammatory biomarkers were obtained at the time of scanning, and in some patients, longitudinally. FINDINGS Between July 2, 2014, and Sept 10, 2015, 705 individuals, who had data available for analysis, were included in this study. 596 (85%) of 705 Tsimane had no CAC, 89 (13%) had CAC scores of 1-100, and 20 (3%) had CAC scores higher than 100. For individuals older than age 75 years, 31 (65%) Tsimane presented with a CAC score of 0, and only four (8%) had CAC scores of 100 or more, a five-fold lower prevalence than industrialised populations (p≤0·0001 for all age categories of MESA). Mean LDL and HDL cholesterol concentrations were 2·35 mmol/L (91 mg/dL) and 1·0 mmol/L (39·5 mg/dL), respectively; obesity, hypertension, high blood sugar, and regular cigarette smoking were rare. High-sensitivity C-reactive protein was elevated beyond the clinical cutoff of 3·0 mg/dL in 360 (51%) Tsimane participants. INTERPRETATION Despite a high infectious inflammatory burden, the Tsimane, a forager-horticulturalist population of the Bolivian Amazon with few coronary artery disease risk factors, have the lowest reported levels of coronary artery disease of any population recorded to date. These findings suggest that coronary atherosclerosis can be avoided in most people by achieving a lifetime with very low LDL, low blood pressure, low glucose, normal body-mass index, no smoking, and plenty of physical activity. The relative contributions of each are still to be determined. FUNDING National Institute on Aging, National Institutes of Health; St Lukes Hospital of Kansas City; and Paleocardiology Foundation.

Leadership in an Egalitarian Society

Leadership is instrumental to resolution of collective action dilemmas, particularly in large, heterogeneous groups. Less is known about the characteristics or effectiveness of leadership in small-scale, homogeneous, and relatively egalitarian societies, in which humans have spent most of our existence. Among Tsimane’ forager-horticulturalists of Bolivia, we (1) assess traits of elected leaders under experimental and naturalistic conditions and (2) test whether leaders impact or differentially benefit from collective action outcomes. We find that elected leaders are physically strong and have more kin and other exchange partners. Their ranks on physical dominance, kin support, and trustworthiness predict how well their groups perform, but only where group members have a history of collaborative interaction. Leaders do not take more of the spoils. We discuss why physically strong leaders can be compatible with egalitarianism, and we suggest that leaders in egalitarian societies may be more motivated by maintaining an altruistic reputation than by short-term rewards of collective action.

Learning, menopause, and the human adaptive complex.

This paper presents a new two‐sex learning‐ and skills‐based theory for the evolution of human menopause. The theory proposes that the role of knowledge, skill acquisition, and transfers in determining economic productivity and resource distribution is the distinctive feature of the traditional human ecology that is responsible for the evolution of menopause. The theory also proposes that male reproductive cessation and post‐reproductive investment in descendants is a fundamental characteristic of humans living in traditional foraging and simple horticultural economies. We present evidence relevant to the theory. The data show that whereas reproductive decline is linked to increasing risks of mortality in chimpanzees, human reproductive senescence precedes somatic senescence. Moreover under traditional conditions, most human males undergo reproductive cessation at the same time as their wives. We then present evidence that after ceasing to reproduce, both men and women provide net economic transfers to children and grandchildren. Given this pattern of economic productivity, delays in menopause would produce net economic deficits within families.

The goals of direct paternal care among a South Amerindian population

Human males are remarkable among mammals in the level of investment they provide to their wives and children. However, there has been debate as to the degree to which men actually invest and through which fitness pathways the benefits of familial investment are realized. Much of the previous research exploring these issues has focused on mens roles as providers, but few have explored correlates of mens direct parental care. Although this is reasonable given mens parental emphasis on provisioning, the providing of direct care is more straightforward with a clear provider and recipient and little ambiguity as to the care-givers intent. Here, we explore contextual correlates of mens direct care among the Tsimane of Bolivia to determine the extent to which such care is patterned to enhance its effectiveness in increasing child wellbeing and the efficient functioning of the family. We also explore whether Tsimane fathers provide care in ways that enhance the positive effect it has on the wifes perception of the care provider. Overall, we find that Tsimane men appear responsive to the needs of children and the family, but show that there is little evidence that men respond to factors expected to increase the impact that mens care has on their reputations with their wives.

Why do women have more children than they want? Understanding differences in women's ideal and actual family size in a natural fertility population

We develop and test a conceptual model of factors influencing womens ideal family size (IFS) in a natural fertility population, the Tsimane of Bolivia. The model posits affects of socioecology, reproductive history, maternal condition, and mens IFS. We test three hypotheses for why women may exceed their IFS despite experiencing socioeconomic development: (H1) limited autonomy; (H2) improved maternal condition; and (H3) low returns on investments in embodied capital.

The evolutionary fitness of personality traits in a small-scale subsistence society.

article i nfo Article history: Initial receipt 31 May 2013 Final revision received 7 September 2013 The maintenance of personality variation remains an unexplained puzzle in evolutionary biology. Despite evidence among non-humans that personality variation affects fitness, few data exist to assess the personality- fitness relationship in humans. Among Tsimane forager-horticulturalists (n = 632), we test whether personality traits (assessed using a 43-item Big Five Inventory administered orally in native language) predict fertility, offspring survivorship, age of first reproduction, and other fitness correlates (extramarital affairs, conflicts, social visitation, food production, and several health measures). Among men, several personality factors associate with higher fertility, more time spent producing food and social visitation. Among women, the relationship between personality and fitness varies across regions of Tsimane territory. The only case of an intermediate personality level associated with highest fitness was found for Industriousness in men. We find that personality factors positively associated with fitness do not associate with greater health costs, although greater Extraversion and Openness may lead to more conflicts among men. Factor heritability ranges from 60% for Prosociality and Extraversion to 8% for Neuroticism. We interpret our results in light of evolutionary modelsthat explain maintenance of personality variation,including incomplete directional selection,mutation- selection balance, condition-dependent reaction norms and fluctuating selection based on sex or spatial variability in selection pressures.

Helminth infection, fecundity, and age of first pregnancy in women

Parasitic worms influence human fecundity Parasitic worms infect 2 billion people globally. Mostly, such infections are symptomless and individual worm burdens are low. Blackwell et al. monitored the fecundity of Tsimane women in Bolivia. These women have on average of 10 children in their lifetimes. However, if they had successive hookworm infections, lifetime births dropped to 7. Surprisingly, if the women were chronically infested with roundworm, they had as many as 12 children. These effects may relate to the balance of immune responses that the different worms induce, rather than to the physiological costs of parasitism. Science, this issue p. 970 Roundworm and hookworm infections are associated with contrasting effects on human reproduction. Infection with intestinal helminths results in immunological changes that influence co-infections, and might influence fecundity by inducing immunological states affecting conception and pregnancy. We investigated associations between intestinal helminths and fertility in women, using 9 years of longitudinal data from 986 Bolivian forager-horticulturalists, experiencing natural fertility and 70% helminth prevalence. We found that different species of helminth are associated with contrasting effects on fecundity. Infection with roundworm (Ascaris lumbricoides) is associated with earlier first births and shortened interbirth intervals, whereas infection with hookworm is associated with delayed first pregnancy and extended interbirth intervals. Thus, helminths may have important effects on human fertility that reflect physiological and immunological consequences of infection.

Immune function in Amazonian horticulturalists

Abstract Background: Amazonian populations are exposed to diverse parasites and pathogens, including protozoal, bacterial, fungal and helminthic infections. Yet much knowledge of the immune system is based on industrialised populations where these infections are relatively rare. Aim: This study examines distributions and age-related differences in 22 measures of immune function for Bolivian forager-horticulturalists and US and European populations. Subjects and methods: Subjects were 6338 Tsimane aged 0–90 years. Blood samples collected between 2004–2014 were analysed for 5-part blood differentials, C-reactive protein, erythrocyte sedimentation rate (ESR) and total immunoglobulins E, G, A and M. Flow cytometry was used to quantify naïve and non-naïve CD4 and CD8 T cells, natural killer cells, and B cells. Results: Compared to reference populations, Tsimane have elevated levels of most immunological parameters, particularly immunoglobulins, eosinophils, ESR, B cells, and natural killer cells. However, monocytes and basophils are reduced and naïve CD4 cells depleted in older age groups. Conclusion: Tsimane ecology leads to lymphocyte repertoires and immunoglobulin profiles that differ from those observed in industrialised populations. These differences have consequences for disease susceptibility and co-vary with patterns of other life history traits, such as growth and reproduction.