Variation in diurnal cortisol patterns among the Indigenous Shuar of Amazonian Ecuador.

OBJECTIVES: The hypothalamic-pituitary-adrenal (HPA) axis and its primary end product, the glucocorticoid cortisol, are major components of the evolved human stress response. However, most studies have examined these systems among populations in high-income settings, which differ from the high pathogen and limited resource contexts in which the HPA axis functioned for most of human evolution. METHODS: We investigated variability in diurnal salivary cortisol patterns among 298 Indigenous Shuar from Amazonian Ecuador (147 males, 151 females; age 2-86 years), focusing on the effects of age, biological sex, and body mass index (BMI) in shaping differences in diurnal cortisol production. Saliva samples were collected three times daily (waking, 30 minutes post-waking, evening) for three consecutive days to measure key cortisol parameters: levels at waking, the cortisol awakening response, the diurnal slope, and total daily output. RESULTS: Age was positively associated with waking levels and total daily output, with Shuar juveniles and adolescents displaying significantly lower levels than adults (p < .05). Sex was not a significant predictor of cortisol levels (p > .05), as Shuar males and females displayed similar patterns of diurnal cortisol production across the life course. Moreover, age, sex, and BMI significantly interacted to moderate the rate of diurnal cortisol decline (p = .027). Overall, Shuar demonstrated relatively lower cortisol concentrations than high-income populations. CONCLUSIONS: This study expands the documented range of global variation in HPA axis activity and diurnal cortisol production and provides important insights into the plasticity of human stress physiology across diverse developmental and socioecological settings.

Reproductive inequality in humans and other mammals.

To address claims of human exceptionalism, we determine where humans fit within the greater mammalian distribution of reproductive inequality. We show that humans exhibit lower reproductive skew (i.e., inequality in the number of surviving offspring) among males and smaller sex differences in reproductive skew than most other mammals, while nevertheless falling within the mammalian range. Additionally, female reproductive skew is higher in polygynous human populations than in polygynous nonhumans mammals on average. This patterning of skew can be attributed in part to the prevalence of monogamy in humans compared to the predominance of polygyny in nonhuman mammals, to the limited degree of polygyny in the human societies that practice it, and to the importance of unequally held rival resources to women's fitness. The muted reproductive inequality observed in humans appears to be linked to several unusual characteristics of our species-including high levels of cooperation among males, high dependence on unequally held rival resources, complementarities between maternal and paternal investment, as well as social and legal institutions that enforce monogamous norms.

Bringing the lab bench to the field: Point-of-care testing for enhancing health research and stakeholder engagement in rural/remote, indigenous, and resource-limited contexts.

Point-of-care testing (POCT) allows researchers and health-care providers to bring the lab bench to the field, providing essential health information that can be leveraged to improve health care, accessibility, and understanding across clinical and research settings. Gaps in health service access are most pronounced in what we term RIR settings-rural/remote regions, involving Indigenous peoples, and/or within resource-limited settings. In these contexts, morbidity and mortality from infectious and non-communicable diseases are disproportionately higher due to numerous geographic, economic, political, and sociohistorical factors. Human biologists and global health scholars are well-positioned to contribute on-the-ground-level insights that can serve to minimize global health inequities and POCT has the potential to augment such approaches. While the clinical benefits of POCT include increasing health service access by bringing testing, rapid diagnosis, and treatment to underserved communities with limited pathways to centralized laboratory testing, POCT also provides added benefits to both health-focused researchers and their participants. Through portable, minimally invasive devices, researchers can provide actionable health data to participants by coupling POCT with population-specific health education, discussing results and their implications, creating space for participants to voice concerns, and facilitating linkages to treatment. POCT can also strengthen human biology research by shedding light on questions of evolutionary and biocultural importance. Here, we expand on the epidemiological and research value, as well as practical and ethical challenges of POCT across stakeholders (i.e., participant, community, health researcher, and trainee). Finally, we emphasize the immense opportunities of POCT for fostering collaborative research and enhancing access to health delivery and information and, by extension, helping to mitigate persistent global health inequities.

Greater chronic morbidity is associated with greater fatigue in six countries: A case of evolutionary mismatch?

Background and objectives: Human susceptibility to chronic non-communicable disease may be explained, in part, by mismatches between our evolved biology and contemporary environmental conditions. Disease-induced fatigue may function to reduce physical activity during acute infection, thereby making more energy available to mount an effective immune response. However, fatigue in the context of chronic disease may be maladaptive because long-term reductions in physical activity increase risks of disease progression and the acquisition of additional morbidities. Here, we test whether cumulative chronic morbidity is associated with subjective fatigue. Methodology: We constructed a cumulative chronic morbidity score using self-reported diagnoses and algorithm-based assessments, and a subjective fatigue score based on four questionnaire items using cross-sectional survey data from the Study on global AGEing and adult health, which features large samples of adults from six countries (China, Ghana, India, Mexico, Russia and South Africa). Results: In a mixed-effects linear model with participants nested in countries (N = 32 455), greater cumulative chronic morbidity is associated with greater subjective fatigue (ß = 0.34, SE = 0.005, P < 2e-16). This association replicates within each country and is robust to adjustment for key sociodemographic and physical covariates (sex, age, household wealth, physical function score, habitual physical activity, BMI and BMI2). Conclusions and implications: Fatigue is a common but perhaps maladaptive neuropsychological response to chronic morbidity. Disease-induced fatigue may mediate a self-perpetuating cycle, in which chronic morbidity reduces physical activity, and less physical activity increases cumulative chronic morbidity. Longitudinal research is needed to test whether chronic morbidity, fatigue and physical activity form a cyclical feedback loop. Lay Summary: Fatigue during acute illness may promote recovery, but persistent fatigue in the context of chronic disease may make matters worse. We present evidence from six countries that more chronic disease is associated with more fatigue. This fatigue may reduce physical activity, which increases risks of acquiring additional chronic health problems.

Low prevalence of anemia among Shuar communities of Amazonian Ecuador.

OBJECTIVE: Anemia is an important global health challenge. We investigate anemia prevalence among Indigenous Shuar of Ecuador to expand our understanding of population-level variation, and to test hypotheses about how anemia variation is related to age, sex, and market integration. METHODS: Hemoglobin levels were measured in a total sample of 1650 Shuar participants (ages 6 months to 86 years) from 46 communities between 2008 and 2017 to compare anemia prevalence across regions characterized by different levels of market integration. RESULTS: Shuar anemia rates among children under 15 years (12.2%), adult women (10.5%), and adult men (5.3%) were less than half of those previously documented in other neo-tropical Indigenous populations. Anemia prevalence did not vary between more traditional and market integrated communities (OR = 0.47, p = .52). However, anemia was negatively associated with body mass index (OR = 0.47, p = .002). CONCLUSIONS: Compared to other South American Indigenous populations, anemia prevalence is relatively low among Shuar of Ecuador and invariant with market integration. Understanding this pattern can provide valuable insights into anemia prevention among at-risk populations.

Age-related patterns of cytomegalovirus antibodies accompanying Epstein-Barr virus co-infection.

OBJECTIVE: Cytomegalovirus (CMV) infection is associated with age-related chronic disease, and co-infection with Epstein-Barr virus (EBV) may compound disease risk. We aimed to assess the frequency of CMV infection and its relationship with age among EBV seropositive individuals in an Indigenous Amazonian population. METHODS: We report concentrations of CMV and EBV antibodies in dried blood spot samples collected from 157 EBV positive Shuar participants aged 15-86 years (60.5% female) to assess CMV infection rate. We used logistic and linear regression models to examine associations among CMV, EBV, and age, adjusting for sex, geographic region, and body mass index. RESULTS: Nearly two-thirds (63.1%) of EBV seropositive participants were also CMV seropositive. A 1-year increase in age was associated with 3.4% higher odds of CMV infection (OR [95% CI]: 1.034 [1.009-1.064], p = .012), but CMV antibody concentration was not significantly associated with age or EBV antibody concentration among co-infected individuals. CONCLUSIONS: Herpesvirus-related immunosenescence may be important to understanding chronic disease risk among Shuar. Future studies should further explore the role of co-infection in shaping age-related changes in immune function.

Pathogen disgust sensitivity protects against infection in a high pathogen environment.

Disgust is hypothesized to be an evolved emotion that functions to regulate the avoidance of pathogen-related stimuli and behaviors. Individuals with higher pathogen disgust sensitivity (PDS) are predicted to be exposed to and thus infected by fewer pathogens, though no studies have tested this directly. Furthermore, PDS is hypothesized to be locally calibrated to the types of pathogens normally encountered and the fitness-related costs and benefits of infection and avoidance. Market integration (the degree of production for and consumption from market-based economies) influences the relative costs/benefits of pathogen exposure and avoidance through sanitation, hygiene, and lifestyle changes, and is thus predicted to affect PDS. Here, we examine the function of PDS in disease avoidance, its environmental calibration, and its socioecological variation by examining associations among PDS, market-related lifestyle factors, and measures of bacterial, viral, and macroparasitic infection at the individual, household, and community levels. Data were collected among 75 participants (ages 5 to 59 y) from 28 households in three Ecuadorian Shuar communities characterized by subsistence-based lifestyles and high pathogen burden, but experiencing rapid market integration. As predicted, we found strong negative associations between PDS and biomarkers of immune response to viral/bacterial infection, and weaker associations between PDS and measures of macroparasite infection, apparently mediated by market integration-related differences. We provide support for the previously untested hypothesis that PDS is negatively associated with infection, and document variation in PDS indicative of calibration to local socioeconomic conditions. More broadly, findings highlight the importance of evolved psychological mechanisms in human health outcomes.

Childhood Daily Energy Expenditure Does Not Decrease with Market Integration and Is Not Related to Adiposity in Amazonia.

BACKGROUND: Childhood overweight and obesity (OW/OB) is increasingly centered in low- and middle-income countries (LMICs) as rural populations experience market integration and lifeway change. Most explanatory studies have relied on imprecise estimates of children's energy expenditure, restricting understanding of the relative effects of changes in diet and energy expenditure on the development of OW/OB in transitioning contexts. OBJECTIVES: This study used gold-standard measurements of children's energy expenditure to investigate the changes that underlie OW/OB and the nutrition/epidemiologic transition. METHODS: Cross-sectional data were collected from "rural" (n = 43) Shuar forager-horticulturalist children and their "peri-urban" (n = 34) Shuar counterparts (age 4-12 y) in Amazonian Ecuador. Doubly labeled water measurements of total energy expenditure (TEE; kcal/d), respirometry measurements of resting energy expenditure (REE; kcal/d), and measures of diet, physical activity, immune activity, and market integration were analyzed primarily using regression models. RESULTS: Peri-urban children had higher body fat percentage (+8.1%, P < 0.001), greater consumption of market-acquired foods (multiple P < 0.001), lower concentrations of immune activity biomarkers (multiple P < 0.05), and lower REE (-108 kcal/d, P = 0.002) than rural children. Despite these differences, peri-urban children's TEE was indistinguishable from that of rural children (P = 0.499). Moreover, although sample-wide IgG concentrations and household incomes predicted REE (both P < 0.05), no examined household, immune activity, or physical activity measures were related to children's overall TEE (all P > 0.09). Diet and energy expenditure associations with adiposity demonstrate that only reported consumption of market-acquired "protein" and "carbohydrate" foods predicted children's body fat levels (multiple P < 0.05). CONCLUSIONS: Despite underlying patterns in REE, Shuar children's TEE is not reliably related to market integration and-unlike dietary measures-does not predict adiposity. These findings suggest a leading role of changing dietary intake in transitions to OW/OB in LMICs.

Variability of C-reactive protein in first-generation Ecuadorian immigrants living in the United States.

OBJECTIVES: Establish the variability of C-reactive protein (CRP) within a population of first-generation immigrants living in the United States. Prior work has theorized that individuals with high levels of childhood pathogen exposure may have lower CRP levels in adulthood, and therefore that for these individuals, CRP may not be as accurate an index of chronic disease risk related to low-level inflammation as is presumed based on data from wealthy populations. This potentially has major implications for the interpretation of CRP as a biomarker of chronic inflammation. METHODS: This longitudinal study collected a total of 125 dried blood spot (DBS) samples from 31 participants (median 4 samples each) and CRP levels in these DBS were assayed by enzyme-linked immunosorbant assay. Surveys were administered to characterize childhood pathogen exposure, and current illness. Variance was estimated using mixed effects regression models. RESULTS: On average, participants were adults (mean = 41.9 years old) who had immigrated to the United States nearly 20 years prior to the study and had nearly universally experienced childhood helminth infection and other major pathogen exposures. Median serum-equivalent CRP was 0.77 mg/L. Individuals reliably differed in subacute CRP levels, and, depending on whether untransformed or log-transformed CRP was the outcome variable, 45% or 62% of variance in CRP was attributable to between-individual differences. CONCLUSIONS: The variability of CRP levels in individuals with relatively high childhood pathogen exposure is comparable to previously reported studies in North America and Europe. However, CRP values are relatively low. CRP is an appropriate measure of subacute inflammation in this sample.

Market integration and soil-transmitted helminth infection among the Shuar of Amazonian Ecuador.

BACKGROUND: Soil-transmitted helminth (STH) infections have many negative health outcomes (e.g., diarrhea, nutritional deficiencies) that can also exacerbate poverty. These infections are generally highest among low-income populations, many of which are also undergoing market integration (MI; increased participation in a market-based economy). Yet the direct impact of MI-related social and environmental changes on STH infection patterns is poorly understood, making it unclear which lifestyle factors should be targeted to better control disease spread. This cross-sectional study examines if household infrastructure associated with greater MI is associated with lower STH burdens among Indigenous Ecuadorian Shuar. METHODS: Kato-Katz fecal smears were used to determine STH infection status and intensity (n = 620 participants; 308 females, 312 males, aged 6 months-86 years); Ascaris lumbricoides (ascarid) and Trichuris trichiura (whipworm) were the primary infection types detected. Structured interviews assessing lifestyle patterns (e.g., measures of household infrastructure) measured participant MI. Multilevel regression analyses and zero-inflated negative binomial regression models tested associations between MI measures and STH infection status or intensity, controlling for individual and community characteristics. RESULTS: Participants residing in more market-integrated households exhibited lower infection rates and intensities than those in less market integrated households. Parasite infection status and T. trichiura infection intensity were lower among participants living in houses with wood floors than those with dirt floors, while individuals using well or piped water from a spring exhibited lower A. lumbricoides infection intensities compared to those using river or stream water. Unexpectedly, latrine type was not significantly related to STH infection status or intensity. These results suggest that sources of exposure differ between the two helminth species. CONCLUSIONS: This study documents associations between household measures and STH infection among an Indigenous population undergoing rapid MI. These findings can help healthcare programs better target interventions and reduce STH exposure among at-risk populations.

Validation of an enzyme-linked immunoassay assay for osteocalcin, a marker of bone formation, in dried blood spots.

OBJECTIVES: Investigating factors that contribute to bone loss and accretion across populations in remote settings is challenging, particularly where diagnostic tools are scarce. To mitigate this challenge, we describe validation of a commercial ELISA assay to measure osteocalcin, a biomarker of bone formation, from dried blood spots (DBS). METHODS: We validated the Osteocalcin Human SimpleStep ELISA kit from Abcam (ab1951214) using 158 matched plasma and DBS samples. Passing-Bablok regression analysis assessed the relationships between plasma and DBS osteocalcin concentrations. Dilutional linearity and spike and recovery experiments determined if the DBS matrix interfered with osteocalcin measurement, and intra- and inter-assay coefficients of variation (CVs) were calculated. Limit of detection, analyte stability, and specific forms of osteocalcin measured by the kit were also investigated. RESULTS: Mean plasma osteocalcin value was 218.2 ng/mL (range 64.6-618.1 ng/mL). Linear relationships existed between plasma and DBS concentrations of osteocalcin, with no apparent bias in plasma vs DBS concentrations. There was no apparent interference of the DBS matrix with measurement of osteocalcin in DBS. Intra-assay CV for DBS was ~8%, while average inter-assay CV was 14.8%. Limit of detection was 0.34 ng/mL. Osteocalcin concentrations were stable in DBS stored at -28°C and room temperature, but not those stored at 37°C. This ELISA kit detects total osteocalcin. CONCLUSIONS: Osteocalcin, a bone formation biomarker, can be measured from DBS. Combined with a previously validated DBS assay for TRACP-5b, a bone resorption biomarker, these assays have the potential to help researchers disentangle the many factors contributing to bone strength.

The developmental origins of risk and time preferences across diverse societies.

Risk and time preferences have often been viewed as reflecting inherent traits such as impatience and self-control. Here, we offer an alternative perspective, arguing that they are flexible and environmentally informed. In Study 1, we investigated risk and time preferences among children in the United States, India, and Argentina, as well as forager-horticulturalist Shuar children in Amazonian Ecuador. We find striking cross-cultural differences in behavior: children in India, the United States, and Argentina are more risk-seeking and future-oriented, whereas Shuar children are more risk-averse and exhibit more heterogeneous time preferences, on average preferring more today choices. To explore 1 of the socioecological forces that may be shaping these preferences, in Study 2, we compared the behavior of more and less market-integrated Shuar children, finding that those in market-integrated regions are more future-oriented and risk-seeking. These findings indicate that cross-cultural differences in risk and time preferences can be traced into childhood and may be influenced by the local environment. More broadly, our results contribute to a growing understanding of plasticity and variation in the development of behavior. (PsycINFO Database Record (c) 2020 APA, all rights reserved).

Disparities in bone density across contemporary Amazonian forager-horticulturalists: Cross-population comparison of the Tsimane and Shuar.

OBJECTIVES: This study investigates bone density across the life course among Bolivian Tsimane and Ecuadorian Shuar of Amazonia. Both groups are rural, high-fertility forager-horticulturalists, with high lifetime physical activity levels. We test whether Tsimane and Shuar bone density patterns are different from each other, and if both groups are characterized by lower osteoporosis risk compared to U.S. references. METHODS: Anthropometric and calcaneal bone density data, obtained via quantitative ultrasonometry (QUS), were collected from 678 Tsimane and 235 Shuar (13-92 years old). Population and sex differences in QUS values (estimated bone mineral density, speed of sound, broadband ultrasound attenuation) by age group were assessed using Mann-Whitney U tests. Age-related change and age at peak QUS value were determined using polynomial regressions. One-way analyses of covariance assessed population-level differences in QUS values by age group adjusting for body mass index. Participants aged 50+ years at elevated osteoporosis risk were identified using a T score < -1.8; binomial tests assessed risk compared to U.S. references. RESULTS: Shuar males and females <50 years old have QUS values 3-36% higher than Tsimane, with differences evident in adolescence. Among Tsimane and Shuar, 49 and 23% of participants aged 50+ years old, respectively, are at high risk for osteoporosis, compared to 34% of Americans; Shuar osteoporosis risk is comparable to Americans, while Tsimane risk is elevated. CONCLUSIONS: Disparate patterns in QUS values are documented for Tsimane and Shuar, with pronounced differences early in life. Potential explanations for differences include gene-environment interactions and/or degree of market integration, which influences diet, activity profiles, pathogen exposures, and other lifestyle covariates. As Tsimane osteoporosis risk is greater than in the United States, findings point to alternative risk factors for low bone density that are not readily discernible in industrialized populations.

Measuring subjective social status in children of diverse societies.

Subjective Social Status (SSS) is a robust predictor of psychological and physiological outcomes, frequently measured as self-reported placement on the MacArthur Scale of Subjective Social Status. Despite its importance, however, there are still open questions regarding how early into ontogeny SSS can be measured, and how well SSS measures can be extended to non-Western and small-scale populations. Here, we investigate the internal consistency of responses to the MacArthur ladder across four cultures by comparing responses to more explicit social comparison questions. We conduct these comparisons among children and adolescents, ages 4 to 18, in India, the United States, and Argentina, in addition to those in two indigenous communities of the Ecuadorean Amazon marked by differing degrees of market integration (total N = 363). We find that responses are consistent in all populations, except for the more remote forager-horticulturalist Ecuadorian community. We also find that, consistent with findings among American adolescents, SSS declines with age. We then assess the test-retest reliability of the MacArthur Scale across two time-points: a subset of Indian participants (N = 43) within one week, and a larger, second sample of Indian participants after one year (N = 665). We find that responses are highly correlated within one week (ρ = 0.47), and moderately correlated after one year (ρ = 0.32). These results suggest that responses to the MacArthur ladder are internally consistent and reliable among children across a range of diverse populations, though care must be taken in utilizing these measures among children of non-industrial, small-scale societies.

Soil-transmitted helminth infection and intestinal inflammation among the Shuar of Amazonian Ecuador.

OBJECTIVES: Little research exists documenting levels of intestinal inflammation among indigenous populations where exposure to macroparasites, like soil-transmitted helminths (STHs), is common. Reduced STH exposure is hypothesized to contribute to increased prevalence of elevated intestinal inflammation in wealthy nations, likely due to coevolutionary histories between STHs and human immune systems that favored anti-inflammatory pathways. Here, we document levels of intestinal inflammation and test associations with STH infection among the Shuar of Ecuador, an indigenous population undergoing socioeconomic/lifestyle changes that influence their hygienic environment. We predict that fecal calprotectin (FC; a measure of intestinal inflammation) will be lower in STH infected individuals and that FC will be negatively associated with infection intensity. METHODS: Stool samples to analyze FC levels and STH infection were collected from 69 Shuar participants (ages 5-75 years). Children (<15 years) and adults (15+ years) were analyzed separately to understand the role of exposure in immune system development and the intestinal inflammatory response. RESULTS: Two species of STH were present: Ascaris lumbricoides and Trichuris trichiura. The relationships between infection and intestinal inflammation were age- and species-specific. While no significant relationships were found among adults, children who were singly infected with T. trichiura had lower FC levels than uninfected children. Infection intensity was not significantly associated with FC in children or adults. CONCLUSIONS: These preliminary results provide limited support for our hypotheses, documenting tentative age- and species-specific associations between FC and infection status. Findings may point to the importance of species-specific STH exposure during immune system development.

A dried blood spot-based method to measure levels of tartrate-resistant acid phosphatase 5b (TRACP-5b), a marker of bone resorption.

OBJECTIVES: A number of basic questions about bone biology have not been answered, including population differences in bone turnover. In part, this stems from the lack of validated minimally invasive biomarker techniques to measure bone formation and resorption in field-based population-level research. The present study addresses this gap by validating a fingerprick dried blood spot (fDBS) assay for tartrate-resistant acid phosphatase 5b (TRACP-5b), a well-defined biomarker of bone resorption and osteoclast number. METHODS: We adapted a commercially available enzyme-linked immunosorbent assay (ELISA) kit from MyBiosource for the quantitative determination of TRACP-5b levels in serum and plasma for use with DBS. We used a rigorous process of assay modification and validation, including the use of a matched set of 189 adult plasma, fDBS, and venous DBS (vDBS) samples; parameters evaluated included precision, reliability, and analyte stability. RESULTS: Plasma and DBS TRACP-5b concentrations showed a linear relationship. There were no systematic differences in TRACP-5b levels in fDBS and vDBS, indicating no significant differences in TRACP-5b distribution between capillary and venous blood. Parallelism and spike-and-recovery results indicated that matrix factors in DBS do not interfere with measurement of TRACP-5b levels from DBS using the validated kit. Intra- and interassay CVs were 5.0% and 12.1%, respectively. DBS samples should preferably be stored frozen but controlled room temperature storage for up to a month may be acceptable. CONCLUSIONS: This DBS-based ELISA assay adds to the methodological toolkit available to human biologists and will facilitate research on bone turnover in population studies.

High energy requirements and water throughput of adult Shuar forager-horticulturalists of Amazonian Ecuador.

OBJECTIVES: We measured total energy expenditure (TEE; kcal/d) and water throughput (L/d) among Shuar forager-horticulturalists from Amazonian Ecuador to compare their daily energy and water demands to adults in other small-scale and industrialized populations. METHODS: TEE and water throughput were measured using the doubly labeled water method among 15 Shuar adults (eight women, seven men; age range 18-60 years) living in a relatively remote village. We used multiple regression to assess the effects of anthropometric variables (body size, fat free mass, age, and sex) on TEE and water throughput. We also compared Shuar TEE and water throughput to those of other small-scale and industrialized societies. RESULTS: TEE among Shuar adults (men: 4141 ± 645 kcal/d, women: 2536 ± 281 kcal/d) was most strongly correlated with fat free mass. Estimated physical activity levels (PAL) calculated as (TEE/estimated BMR), were greater for men (2.34 ± 0.29) than women (1.83 ± 0.14, P < 0.001). Water throughput was also greater among Shuar men (9.37 ± 2.34 L/d) than women (4.76 ± 0.36 L/d, P < 0.001). Shuar TEE and water throughput were elevated compared to adults in industrialized populations. DISCUSSION: TEE and PAL of Shuar men are among the highest recorded during normal daily life, and likely reflect both high levels of physical activity and cultural dietary practices. Drinking large amounts of chicha, a traditional carbohydrate-rich drink made from manioc, likely contributes to the high levels of water throughput among Shuar men, and may contribute to elevated TEE.

Constraint and trade-offs regulate energy expenditure during childhood.

Children's metabolic energy expenditure is central to evolutionary and epidemiological frameworks for understanding variation in human phenotype and health. Nonetheless, the impact of a physically active lifestyle and heavy burden of infectious disease on child metabolism remains unclear. Using energetic, activity, and biomarker measures, we show that Shuar forager-horticulturalist children of Amazonian Ecuador are ~25% more physically active and, in association with immune activity, have ~20% greater resting energy expenditure than children from industrial populations. Despite these differences, Shuar children's total daily energy expenditure, measured using doubly labeled water, is indistinguishable from industrialized counterparts. Trade-offs in energy allocation between competing physiological tasks, within a constrained energy budget, appear to shape childhood phenotypic variation (e.g., patterns of growth). These trade-offs may contribute to the lifetime obesity and metabolic health disparities that emerge during rapid economic development.

Cross-cultural invariances in the architecture of shame.

Human foragers are obligately group-living, and their high dependence on mutual aid is believed to have characterized our species' social evolution. It was therefore a central adaptive problem for our ancestors to avoid damaging the willingness of other group members to render them assistance. Cognitively, this requires a predictive map of the degree to which others would devalue the individual based on each of various possible acts. With such a map, an individual can avoid socially costly behaviors by anticipating how much audience devaluation a potential action (e.g., stealing) would cause and weigh this against the action's direct payoff (e.g., acquiring). The shame system manifests all of the functional properties required to solve this adaptive problem, with the aversive intensity of shame encoding the social cost. Previous data from three Western(ized) societies indicated that the shame evoked when the individual anticipates committing various acts closely tracks the magnitude of devaluation expressed by audiences in response to those acts. Here we report data supporting the broader claim that shame is a basic part of human biology. We conducted an experiment among 899 participants in 15 small-scale communities scattered around the world. Despite widely varying languages, cultures, and subsistence modes, shame in each community closely tracked the devaluation of local audiences (mean r = +0.84). The fact that the same pattern is encountered in such mutually remote communities suggests that shame's match to audience devaluation is a design feature crafted by selection and not a product of cultural contact or convergent cultural evolution.

The Perception of Spontaneous and Volitional Laughter Across 21 Societies.

Laughter is a nonverbal vocalization occurring in every known culture, ubiquitous across all forms of human social interaction. Here, we examined whether listeners around the world, irrespective of their own native language and culture, can distinguish between spontaneous laughter and volitional laughter-laugh types likely generated by different vocal-production systems. Using a set of 36 recorded laughs produced by female English speakers in tests involving 884 participants from 21 societies across six regions of the world, we asked listeners to determine whether each laugh was real or fake, and listeners differentiated between the two laugh types with an accuracy of 56% to 69%. Acoustic analysis revealed that sound features associated with arousal in vocal production predicted listeners' judgments fairly uniformly across societies. These results demonstrate high consistency across cultures in laughter judgments, underscoring the potential importance of nonverbal vocal communicative phenomena in human affiliation and cooperation.