The Reproductive Ecology of Industrial Societies,Part II

Studies of the association between wealth and fertility in industrial populations have a rich history in the evolutionary literature, and they have been used to argue both for and against a behavioral ecological approach to explaining human variability. We consider that there are strong arguments in favor of measuring fertility (and proxies thereof) in industrial populations, not least because of the wide availability of large-scale secondary databases. Such data sources bring challenges as well as advantages, however. The purpose of this article is to illustrate these by examining the association between wealth and reproductive success in the United States, using the National Longitudinal Study of Youth 1979. We conduct a broad-based exploratory analysis of the relationship between wealth and fertility, employing both cross-sectional and longitudinal approaches, and multiple measures of both wealth (income and net worth) and fertility (lifetime reproductive success and transitions to first, second and third births). We highlight the kinds of decisions that have to be made regarding sample selection, along with the selection and construction of explanatory variables and control measures. Based on our analyses, we find a positive effect of both income and net worth on fertility for men, which is more pronounced for white men and for transitions to first and second births. Income tends to have a negative effect on fertility for women, while net worth is more likely to positively predict fertility. Different reproductive strategies among different groups within the same population highlight the complexity of the reproductive ecology of industrial societies. These results differ in a number of respects from other analyses using the same database. We suggest this reflects the impossibility of producing a definitive analysis, rather than a failure to identify the “correct” analytical strategy. Finally, we discuss how these findings inform us about (mal)adaptive decision-making.     

Optimizing Modern Family Size

Modern industrialized populations lack the strong positive correlations between wealth and reproductive success that characterize most traditional societies. While modernization has brought about substantial increases in personal wealth, fertility in many developed countries has plummeted to the lowest levels in recorded human history. These phenomena contradict evolutionary and economic models of the family that assume increasing wealth reduces resource competition between offspring, favoring high fertility norms. Here, we review the hypothesis that cultural modernization may in fact establish unusually intense reproductive trade-offs in wealthy relative to impoverished strata, favoring low fertility. We test this premise with British longitudinal data (the Avon Longitudinal Study of Parents and Children), exploring maternal self-perceptions of economic hardship in relation to increasing family size and actual socioeconomic status. Low-income and low-education-level mothers perceived the greatest economic costs associated with raising two versus one offspring. However, for all further increases to family size, reproduction appears most expensive for relatively wealthy and well-educated mothers. We discuss our results and review current literature on the long-term consequences of resource dilution in modern families.     

The demographic transition: are we any closer to an evolutionary explanation?

The radical shift in human reproduction in the late 19th century, known as the demographic transition, constitutes a major challenge to evolutionary approaches to human behaviour. Why would people ever choose to limit their reproduction voluntarily when, at the peak of the Industrial Revolution, resources were apparently so plentiful? Can the transition be attributed to standard life history tradeoffs, is it a consequence of cultural evolutionary processes, or is it simply a maladaptive outcome of novel and environmental social conditions? Empirical analyses and new models suggest that reproductive decision making might be driven by a human psychology designed by natural selection to maximize material wealth. If this is the case, the mechanisms governing fertility and parental investment are likely to respond to modern conditions with a fertility level much lower than that that would maximize fitness.     

The Trade-Off between Female Fertility and Longevity during the Epidemiological Transition in the Netherlands

Several hypotheses have been put forward to explain the relationship between women’s fertility and their post-reproductive longevity. In this study, we focus on the disposable soma theory, which posits that a negative relationship between women’s fertility and longevity can be understood as an evolutionary trade-off between reproduction and survival. We examine the relationship between fertility and longevity during the epidemiological transition in the Netherlands. This period of rapid decline in mortality from infectious diseases offers a good opportunity to study the relationship between fertility and longevity, using registry data from 6,359 women born in The Netherlands between 1850 and 1910. We hypothesize that an initially negative relationship between women’s fertility and their longevity gradually turns less negative during the epidemiological transition, because of decreasing costs of higher parities. An initially inversed U-shaped association between fertility and longevity changes to zero during the epidemiological transition. This does suggest a diminishing environmental pressure on fertility. However, we find no evidence of an initial linear trade-off between fertility and post-reproductive survival.     

Demographic studies enhance the understanding of evolutionarily (mal)adaptive behaviors and phenomena in humans: a review on fertility decline and an integrated model

Recently, statistical analyses of demographic datasets have come to play an important role for studies into the evolution of human life history. In the first part of this paper, I highlight fertility decline, an evolutionarily paradoxical phenomenon in terms of fitness maximization. Then, I conduct a literature review regarding the effects of socioeconomic status on the number of offspring, especially in modern developed, (post-)industrial, and low-fertility societies. Although a non-positive relationship between them has often been recognized as a general feature of fertility decline, there actually exists a great deal of variation. Based on the review, I discuss the association between socioeconomic success and reproductive success, and tackle an evolutionary question as to why people seek higher socioeconomic success that would not directly lead to higher reproductive success. It has been suggested that, in modern competitive environments, parents should set a higher value on their investment in children, and aim to have a smaller number of high-quality children. Also, parents would maintain higher socioeconomic status for themselves so as to provide high-levels of investment in their children. In the second part, I broadly consider seemingly evolutionarily (mal)adaptive outcomes besides fertility decline, including child abuse, menopause, and suicide. The integration of the major three approaches to human behavioral and psychological research (behavioral ecology, evolutionary psychology, and cultural evolution) could lead to a deeper understanding. I provide a model for the integrated approach. Rich empirical evidence accumulated in demographic studies, especially longitudinal and cross-cultural resources, can assist to develop a theoretical framework.     

The Reproductive Ecology of Industrial Societies,Part I

Is fertility relevant to evolutionary analyses conducted in modern industrial societies? This question has been the subject of a highly contentious debate, beginning in the late 1980s and continuing to this day. Researchers in both evolutionary and social sciences have argued that the measurement of fitness-related traits (e.g., fertility) offers little insight into evolutionary processes, on the grounds that modern industrial environments differ so greatly from those of our ancestral past that our behavior can no longer be expected to be adaptive. In contrast, we argue that fertility measurements in industrial society are essential for a complete evolutionary analysis: in particular, such data can provide evidence for any putative adaptive mismatch between ancestral environments and those of the present day, and they can provide insight into the selection pressures currently operating on contemporary populations. Having made this positive case, we then go on to discuss some challenges of fertility-related analyses among industrialized populations, particularly those that involve large-scale databases. These include “researcher degrees of freedom” (i.e., the choices made about which variables to analyze and how) and the different biases that may exist in such data. Despite these concerns, large datasets from multiple populations represent an excellent opportunity to test evolutionary hypotheses in great detail, enriching the evolutionary understanding of human behavior.     

Variation in wealth and educational drivers of fertility decline across 45 countries

Fertility decline in human populations is an inherent evolutionary puzzle with major demographic, socio-cultural and evolutionary consequences. The individual level predictors of fertility decline are numerous, but the way these effects vary by country and how they are causally mediated by other factors has received relatively little attention. Here we take a multilevel approach to compare similarities and differences in the primary predictors of contemporary fertility declines—wealth and education—across 45 countries in Africa, Asia, Central and South America, the Caribbean, and the Middle East using Demographic and Health Survey (DHS) data collected from 2003 to 2015. We use multilevel models to understand variation in the slopes of these predictors on fertility, and structural equation models to examine the causal pathways by which they take their effects, focusing on four mediating variables: local mortality and birth rates, women’s work status, and contraceptive use. We find that associations between wealth and fertility differ substantially across populations, while associations between education and fertility are consistently negative. The mediators also vary: community-level birth rates and women’s contraceptive use are important mediators between education, wealth and the number of children born across a wide variety of countries, but community-level mortality rates and women’s work status are not. We discuss our results in the context of different causal pathways that reflect cultural and biological evolutionary dynamics as simultaneous and interacting drivers of fertility decline.     

Why pigs are important in Papua? Wealth,height and reproductive success among the Yali tribe of West Papua

Many studies have investigated how different variables influence the reproductive success (RS) in the populations of natural birth control. Here, we tested hypotheses about positive relationship between wealth, height and several measures of RS in an indigenous, traditional society from West Papua. The study was conducted among the Yali tribe in a few small, isolated mountain villages. In this tribe, a man's wealth is measured by the number of pigs he possesses. We found that wealth was related to fertility and number of living children, but not to child mortality in both men and women. Additionally, child mortality increased with the number of children in a family. Finally, we did not observe any relationship between height and reproductive success measures or wealth. We provide several possible explanations of our results and also put forward hypothetical background for further studies of indigenous populations.     

Parental investment and the optimization of human family size

Human reproductive behaviour is marked by exceptional variation at the population and individual level. Human behavioural ecologists propose adaptive hypotheses to explain this variation as shifting phenotypic optima in relation to local socioecological niches. Here we review evidence that variation in fertility (offspring number), in both traditional and modern industrialized populations, represents optimization of the life-history trade-off between reproductive rate and parental investment. While a reliance on correlational methods suggests the true costs of sibling resource competition are often poorly estimated, a range of anthropological and demographic studies confirm that parents balance family size against offspring success. Evidence of optimization is less forthcoming. Declines in fertility associated with modernization are particularly difficult to reconcile with adaptive models, because fertility limitation fails to enhance offspring reproductive success. Yet, considering alternative measures, we show that modern low fertility confers many advantages on offspring, which are probably transmitted to future generations. Evidence from populations that have undergone or initiated demographic transition indicate that these rewards to fertility limitation fall selectively on relatively wealthy individuals. The adaptive significance of modern reproductive behaviour remains difficult to evaluate, but may be best understood in response to rising investment costs of rearing socially and economically competitive offspring.     

Resources and reproductive success in women with an example from the Kipsigis of Kenya

Contradictory results regarding the relationship between resources and reproductive success of women have led some social scientists to conclude that evolutionary biological models are inappropriate to the study of human social behavior. This paper suggests instead that the variability across societies in this relationship reflects an inadequate specification of the nature and availability of the resources critical to reproduction as well as a failure to understand the mechanisms whereby resources confer reproductive success in traditional, developing, preindustrial, and modern societies. These methodological and conceptual issues are illustrated through use of data on the association between wealth and reproductive success from the Kipsigis, a polygynous agropastoralist population in southwestern Kenya. In this society, land is owned by men, and women gain access to land through marriage. In 3 of the 5 marriage cohorts studied, women with access to larger land plots had higher lifelong reproductive success than their poorer counterparts both in terms of enhanced fertility and survivorship of offspring. This association was independent of confounding factors such as education, age at menarche, husband's age, or occupation. Moreover, wealthy women were found not to make greater use of modern medical child health services when their children were sick than poor women. The Kipsigis data indicate that wealthy women had more nutritional resources than poor women and were able to introduce more suitable weaning foods, leading to a lower incidence of episodes of illness in offspring. Overall, the findings suggest that wealth-related differences in the nutrition and health of mothers and children are important factors in reproductive differentials in Kipsigis society.     

Ecological variation in wealth–fertility relationships in Mongolia: the ‘central theoretical problem of sociobiology’ not a problem after all?

The negative wealth–fertility relationship brought about by market integration remains a puzzle to classic evolutionary models. Evolutionary ecologists have argued that this phenomenon results from both stronger trade-offs between reproductive and socioeconomic success in the highest social classes and the comparison of groups rather than individuals. Indeed, studies in contemporary low fertility settings have typically used aggregated samples that may mask positive wealth–fertility relationships. Furthermore, while much evidence attests to trade-offs between reproductive and socioeconomic success, few studies have explicitly tested the idea that such constraints are intensified by market integration. Using data from Mongolia, a post-socialist nation that underwent mass privatization, we examine wealth–fertility relationships over time and across a rural–urban gradient. Among post-reproductive women, reproductive fitness is the lowest in urban areas, but increases with wealth in all regions. After liberalization, a demographic–economic paradox emerges in urban areas: while educational attainment negatively impacts female fertility in all regions, education uniquely provides socioeconomic benefits in urban contexts. As market integration progresses, socio-economic returns to education increase and women who limit their reproduction to pursue education get wealthier. The results support the view that selection favoured mechanisms that respond to opportunities for status enhancement rather than fertility maximization.     

Understanding variation in human fertility: what can we learn from evolutionary demography?

Decades of research on human fertility has presented a clear picture of how fertility varies, including its dramatic decline over the last two centuries in most parts of the world. Why fertility varies, both between and within populations, is not nearly so well understood. Fertility is a complex phenomenon, partly physiologically and partly behaviourally determined, thus an interdisciplinary approach is required to understand it. Evolutionary demographers have focused on human fertility since the 1980s. The first wave of evolutionary demographic research made major theoretical and empirical advances, investigating variation in fertility primarily in terms of fitness maximization. Research focused particularly on variation within high-fertility populations and small-scale subsistence societies and also yielded a number of hypotheses for why fitness maximization seems to break down as fertility declines during the demographic transition. A second wave of evolutionary demography research on fertility is now underway, paying much more attention to the cultural and psychological mechanisms underpinning fertility. It is also engaging with the complex, multi-causal nature of fertility variation, and with understanding fertility in complex modern and transitioning societies. Here, we summarize the history of evolutionary demographic work on human fertility, describe the current state of the field, and suggest future directions.     

Between nurture and nature: the shifting determinants of female fertility in Danish twin cohorts

Behaviors related to fertility constitute primary candidates for investigating the relevance of evolutionary influences and biological dispositions on contemporary human behaviors. Using female Danish twin cohorts born 1870-1968, we document important transformations in the relative contributions of "nurture" and "nature" to within-cohort variations in early and complete fertility, and we point toward a systematic relation between the socioeconomic context of cohorts and the relevance of genetic and shared environmental factors. This transformation is most striking for early fertility where genetic factors strengthen over time and are consistent with up to 50 percent of the variation in early fertility in most recent cohorts. Understanding this emerging relevance of genetic factors is of central importance because early fertility constitutes an important determinant of complete fertility levels in low-fertility societies, and because teenage motherhood and early childbearing are often associated with negative life-cycle consequences. Moreover, our results emphasize the need for socially and contextually informed analyses of nature and nurture that allow both factors to influence human reproductive behavior over time.     

Morph‐dependent fitness and directional change of morph frequencies over time in a Dutch population of Common buzzards Buteo buteo

How genetic polymorphisms are maintained in a population is a key question in evolutionary ecology. Previous work on a plumage colour polymorphism in the common buzzard Buteo buteo suggested heterozygote advantage as the mechanism maintaining the co‐existence of three morphs (light, intermediate and dark). We took advantage of 20 years of life‐history data collected in a Dutch population to replicate earlier studies on the relationship between colour morph and fitness in this species. We examined differences between morphs in adult apparent survival, breeding success, annual number of fledglings produced and cumulative reproductive success. We found that cumulative reproductive success differed among morphs, with the intermediate morph having highest fitness. We also found assortative mating for colour morph, whereby assortative pairs were more likely to produce offspring and had longer‐lasting pair bonds than disassortative pairs. Over the 20‐year study period, the proportion of individuals with an intermediate morph increased. This apparent evolutionary change did not just arise from selection on individual phenotypes, but also from fitness benefits of assortative mating. The increased frequency of intermediates might also be due to immigration or drift. We hypothesize that genetic variation is maintained through spatial variation in selection pressures. Further studies should investigate morph‐dependent dispersal behaviour and habitat choice.     

Resources and reproductive success in women with an example from the Kipsigis of Kenya

Contrary to findings in studies of many mammalian species, the relationship between resources and reproductive success of women is highly variable across societies. This has led some social scientists to suggest that evolutionary biological models are inappropriate to the study of human social behaviour. This paper proposes that variability in the relationship between resources and reproductive success arises from an inadequate specification of the nature and availability of resources critical to reproduction and a failure to understand the mechanisms whereby resources confer reproductive differentials in different environments. Data are presented from the Kipsigis of Kenya showing how land ownership affects female reproductive success; reasons why the effects of landholding are changing over time are examined.     

Evolutionary model of protein secondary structure capable of revealing new biological relationships

Ancestral sequence reconstruction has had recent success in decoding the origins and the determinants of complex protein functions. However, phylogenetic analyses of remote homologues must handle extreme amino acid sequence diversity resulting from extended periods of evolutionary change. We exploited the wealth of protein structures to develop an evolutionary model based on protein secondary structure. The approach follows the differences between discrete secondary structure states observed in modern proteins and those hypothesized in their immediate ancestors. We implemented maximum likelihood-based phylogenetic inference to reconstruct ancestral secondary structure. The predictive accuracy from the use of the evolutionary model surpasses that of comparative modeling and sequence-based prediction; the reconstruction extracts information not available from modern structures or the ancestral sequences alone. Based on a phylogenetic analysis of a sequence-diverse protein family, we showed that the model can highlight relationships that are evolutionarily rooted in structure and not evident in amino acid-based analysis.     

Sociobiology's relevance to modern society: commentary on two articles published here

This article looks at two articles recently published here, one by R. Hopcroft [“Sex, status, and reproductive success in the contemporary United States,” Evolution and Human Behavior 27 (2006) 104–120] the second by M. Fieder and S. Huber [“The effects of sex and childlessness on reproductive effort in modern humans,” Evolution and Human Behavior 28 (2007) 392–398]. They both report a positive relationship between male income and male fertility, which marks a real advancement in our knowledge. Hopcroft speculates at the end of her article that the father's status and female fertility will be positive, too. At this point, this will depend on her and others' reaction to the efforts of S. Huber, F. Bookstein, and M. Fiedler. (2010). Socioeconomic status, education, and reproduction in modern women: an evolutionary perspective. American Journal of Human Biology, 22, 578-587. M. Fieder and S. Huber. (2007). The effects of sex and childlessness on the association between status and reproductive output in modern society. Evolution and Human Behavior, 2, 392-398. are not explicit about this relationship; their closing lines just leave open the possibility that it might be positive. The conclusion is that sociobiology is not really relevant yet to the understanding of modern, industrial society, urban civilization in general, except to show us the various restraints imposed by inborn human nature.     

Trade-offs in modern parenting: a longitudinal study of sibling competition for parental care

Evolutionary and economic models of the family propose that parents face a fundamental trade-off between fertility and investment per offspring. However, tests of this hypothesis have focused primarily on offspring outcomes rather than direct measures of parental investment. Existing studies of parenting also suffer a number of methodological problems now recognized as common sources of error in sociodemographic studies. Here, we present a more definitive picture of the effects of family structure on parental care by analyzing an extensive longitudinal dataset of contemporary British families (the Avon Longitudinal Study of Parents and Children). Unlike other studies, we simultaneously track maternal and paternal behaviors within the same family and consider variation both across time and between distinct population subgroups. Parental investment was measured as frequency of engagement in key care activities over the first decade of life. For both parents, larger family size was traded off against investment per offspring, representing the strongest explanatory variable considered in our analysis. However, contrary to the predictions of traditional quantity–quality trade-off models, increasing family socioeconomic status did not alleviate this effect. In fact, for paternal care in particular, increases in wealth and education created stronger trade-offs. We also demonstrate that large sibships were particularly costly for later-born offspring. Sex of siblings did not influence parental care, however maternal investment was biased towards daughters and paternal investment biased towards sons. Unrelated father figures were also associated with lower investment from both parents. Results are discussed in relation to parental investment theory and evolutionary models of modern low fertility.     

A test of genetic models for the evolutionary maintenance of same-sex sexual behaviour

The evolutionary maintenance of same-sex sexual behaviour (SSB) has received increasing attention because it is perceived to be an evolutionary paradox. The genetic basis of SSB is almost wholly unknown in non-human animals, though this is key to understanding its persistence. Recent theoretical work has yielded broadly applicable predictions centred on two genetic models for SSB: overdominance and sexual antagonism. Using Drosophila melanogaster, we assayed natural genetic variation for male SSB and empirically tested predictions about the mode of inheritance and fitness consequences of alleles influencing its expression. We screened 50 inbred lines derived from a wild population for male–male courtship and copulation behaviour, and examined crosses between the lines for evidence of overdominance and antagonistic fecundity selection. Consistent variation among lines revealed heritable genetic variation for SSB, but the nature of the genetic variation was complex. Phenotypic and fitness variation was consistent with expectations under overdominance, although predictions of the sexual antagonism model were also supported. We found an unexpected and strong paternal effect on the expression of SSB, suggesting possible Y-linkage of the trait. Our results inform evolutionary genetic mechanisms that might maintain low but persistently observed levels of male SSB in D. melanogaster, but highlight a need for broader taxonomic representation in studies of its evolutionary causes.     

How contemporary human reproductive behaviors influence the role of fertility-related genes: the example of the p53 gene

Studies on human fertility genes have identified numerous risk/protective alleles involved in the occurrence of reproductive system diseases causing infertility or subfertility. Investigations we carried out in populations at natural fertility seem to suggest that the clinical relevance that some fertility genes are now acquiring depends on their interaction with contemporary reproductive behaviors (birth control, delayed childbearing, and spacing birth order, among others). In recent years, a new physiological role in human fertility regulation has emerged for the tumor- suppressor p53 gene (P53), and the P53 Arg72Pro polymorphism has been associated with recurrent implantation failure in humans. To lend support to our previous observations, we examined the impact of Arg72Pro polymorphism on fertility in two samples of Italian women not selected for impaired fertility but collected from populations with different (premodern and modern) reproductive behaviors. Among the women at near-natural fertility (n = 98), the P53 genotypes were not associated with different reproductive efficiency, whereas among those with modern reproductive behaviors (n = 68), the P53 genotypes were associated with different mean numbers of children [Pro/Pro = 0.75相似文献