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.     

The coevolution of human fertility and wealth inheritance strategies.

Life history theory concerns the scheduling of births and the level of parental investment in each offspring. In most human societies the inheritance of wealth is an important part of parental investment. Patterns of wealth inheritance and other reproductive decisions, such as family size, would be expected to influence each other. Here I present an adaptive model of human reproductive decision-making, using a state-dependent dynamic model. Two decisions made by parents are considered: when to have another baby, and thus the pattern of reproduction through life; and how to allocate resources between children at the end of the parents'' life. Optimal decision rules are those that maximize the number of grandchildren. Decisions are assumed to depend on the state of the parent, which is described at any time by two variables: number of living sons, and wealth. The dynamics of the model are based on a traditional African pastoralist system, but it is general enough to approximate to any means of subsistence where an increase in the amount of wealth owned increases the capacity for future production of resources. The model is used to show that, in the unpredictable environment of a traditional pastoralist society, high fertility and a biasing of wealth inheritance to a small number of children are frequently optimal. Most such societies are now undergoing a transition to lower fertility, known as the demographic transition. The effects on fertility and wealth inheritance strategies of reducing mortality risks, reducing the unpredictability of the environment and increasing the costs of raising children are explored. Reducing mortality has little effect on completed family sizes of living children or on the wealth they inherit. Increasing the costs of raising children decreases optimal fertility and increases the inheritance left to each child at each level of wealth, and has the potential to reduce fertility to very low levels. The results offer an explanation for why wealthy families are frequently also those with the smallest number of children in heterogeneous, post-transition societies.     

Natural selection on female life-history traits in relation to socio-economic class in pre-industrial human populations

Life-history theory predicts that resource scarcity constrains individual optimal reproductive strategies and shapes the evolution of life-history traits. In species where the inherited structure of social class may lead to consistent resource differences among family lines, between-class variation in resource availability should select for divergence in optimal reproductive strategies. Evaluating this prediction requires information on the phenotypic selection and quantitative genetics of life-history trait variation in relation to individual lifetime access to resources. Here, we show using path analysis how resource availability, measured as the wealth class of the family, affected the opportunity and intensity of phenotypic selection on the key life-history traits of women living in pre-industrial Finland during the 1800s and 1900s. We found the highest opportunity for total selection and the strongest selection on earlier age at first reproduction in women of the poorest wealth class, whereas selection favoured older age at reproductive cessation in mothers of the wealthier classes. We also found clear differences in female life-history traits across wealth classes: the poorest women had the lowest age-specific survival throughout their lives, they started reproduction later, delivered fewer offspring during their lifetime, ceased reproduction younger, had poorer offspring survival to adulthood and, hence, had lower fitness compared to the wealthier women. Our results show that the amount of wealth affected the selection pressure on female life-history in a pre-industrial human population.     

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.     

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.     

Racial and cultural differences in sensitivity to flickering light

Latin America has been registering a fast decrease in fertility rates since the mid-twentieth century. This change can be linked to the modernization process these populations have been undergoing. However, research with Latin American indigenous populations, which are undergoing relatively similar lifestyle changes, shows very different trends in fertility. The aim of this study was to analyze fertility patterns in the indigenous Toba community of Cacique Sombrero Negro, which is experiencing a rapid process of economic and social Westernization. Fertility patterns were analyzed between 1981 and 1999, the period for which the most accurate records were found. Results showed an overall increase in fertility rates and changes in the age of peak fertility across time periods. It is hypothesized that the lifestyle transition this population is experiencing leads to better access to resources that, in the absence of contraception, allow for a higher number of offspring. Nevertheless, this higher resource availability would be differential, affecting mostly the fertility of younger mothers.     

Prestige and reproductive success in man

The relationship between prestige and reproductive success is explored in ten human societies. Formally, the aim is to test the hypothesis that prestige and reproductive success are positively correlated in all populations. It is concluded that in societies at or slightly above the subsistence level prestige and reproductive success are congruent, but that with the accumulation of surplus wealth sources of prestige which have no relevance for reproductive success begin to appear, their development being facilitated by the law of diminishing returns as applied to parents' nonbiological investment in offspring, a point being reached where further investment can have no beneficial effect on reproductive success. As a result sociocultural success and reproductive success may separate. In some societies, where prestige depends upon the possession of real estate, population control is embraced by wealthy families in order to keep their estates intact, thus increasing the effectiveness of the law of diminishing returns by reducing the “fixed resources” (i.e., number of offspring) available for investment. In these conditions, which may spread to the less wealthy, sociocultural success often takes precedence over reproductive success.     

Winter resource wealth drives delayed dispersal and family-group living in western bluebirds

Delayed dispersal, where offspring remain with parents beyond the usual period of dependence, is the typical route leading to formation of kin-based cooperative societies. The prevailing explanations for why offspring stay home are variation in resource wealth, in which offspring of wealthy parents benefit disproportionately by staying home, and nepotism, where the tendency for parents to be less aggressive and share food with offspring makes home a superior place to wait to breed. These hypotheses are not strict alternatives, as only wealthy parents have sufficient resources to share. In western bluebirds, Sialia mexicana, sons usually delay dispersal until after winter, gaining feeding advantages through maternal nepotism in a familial winter group. Experimentally reducing resource wealth (mistletoe) by half on winter territories caused sons to disperse in summer, even though their parents remained on the territory during the winter. Only 8% of sons remained with their parents on mistletoe-removal territories compared to 50% of sons on control territories (t(9,10)=3.33, p<0.005). This study is the first to demonstrate that experimentally reducing wealth of a natural food resource reduces delayed dispersal, facilitating nepotism and family-group living. The results clarify the roles of year-round residency, resource limitation and relative wealth outside the breeding season in facilitating the formation of kin-based cooperative societies.     

Resource exploitation and relatedness: implications for offspring size variation within broods

Much of the theory on offspring size variation within a brood relies on unequal maternal allocation of resources to each embryo. However, maternal allocation strategies are subject to an inherent conflict between mothers and offspring: individual offspring, being more closely related to themselves than to their siblings, should always prefer a larger share of the available resources than that which is optimal from their mother's perspective. Thus, in species where mothers cannot unilaterally impose a resource allocation strategy, offspring can respond to this conflict by competing for more resources than is maternally optimal. Here we show that variation in offspring size within a brood can arise as a by‐product of competition between siblings over a common resource, even when 1) there are no competitive inequalities within families, and 2) maternal investment per brood is fixed. Moreover, we show that size variance among offspring increases with increasing levels of competition, brought about by decreasing relatedness among siblings. Conflict thus offers a simple, testable and, potentially general, explanation for the wide variability in offspring size seen in nature. This extends explanatory hypotheses for offspring size variation beyond those of maternal effects, under which most explanations have been subsumed to date.     

Ecological demography: A synthetic focus in evolutionary anthropology

The interests of evolutionary anthropologists, behavioral ecologists, and demographers converge on the ecology of human fertility. Ecological conditions influence the optimum pattern of maternal effort. Patterns of abortion, neglect, and infanticide vary with mothers' ability to invest in their children and children's ability to use that investment. As in most other mammals, the ecology of human fertility varies between the sexes: status and resource control are important for males, whereas reproductive value is crucial for females. In pre-industrial societies, and even in monogamous societies in demographic transition, wealthy men had more children than did poorer men. This correlation, often assumed to have disappeared, persists today, with richer men still having more sexual access than others. Sex differences in the ecology of fertility mean that sex of the offspring, as well as birth order, influences parental investment. Because individual fertility varies with environment, it is not surprising that “natural” (uncontrolled) fertility varies across societies or that demographic transitions proceed locally, with occasional reverses, as individuals strive to maximize their lifetime reproductive success in changing, competitive, conditions.     

Fertility decline and the changing dynamics of wealth,status and inequality

In the course of demographic transitions (DTs), two large-scale trends become apparent: (i) the broadly positive association between wealth, status and fertility tends to reverse, and (ii) wealth inequalities increase and then temporarily decrease. We argue that these two broad patterns are linked, through a diversification of reproductive strategies that subsequently converge as populations consume more, become less self-sufficient and increasingly depend on education as a route to socio-economic status. We examine these links using data from 22 mid-transition communities in rural Poland. We identify changing relationships between fertility and multiple measures of wealth, status and inequality. Wealth and status generally have opposing effects on fertility, but these associations vary by community. Where farming remains a viable livelihood, reproductive strategies typical of both pre- and post-DT populations coexist. Fertility is lower and less variable in communities with lower wealth inequality, and macro-level patterns in inequality are generally reproduced at the community level. Our results provide a detailed insight into the changing dynamics of wealth, status and inequality that accompany DTs at the community level where peoples'' social and economic interactions typically take place. We find no evidence to suggest that women with the most educational capital gain wealth advantages from reducing fertility, nor that higher educational capital delays the onset of childbearing in this population. Rather, these patterns reflect changing reproductive preferences during a period of profound economic and social change, with implications for our understanding of reproductive and socio-economic inequalities in transitioning populations.     

Wealth,fertility and adaptive behaviour in industrial populations

The lack of association between wealth and fertility in contemporary industrialized populations has often been used to question the value of an evolutionary perspective on human behaviour. Here, we first present the history of this debate, and the evolutionary explanations for why wealth and fertility (the number of children) are decoupled in modern industrial settings. We suggest that the nature of the relationship between wealth and fertility remains an open question because of the multi-faceted nature of wealth, and because existing cross-sectional studies are ambiguous with respect to how material wealth and fertility are linked. A literature review of longitudinal studies on wealth and fertility shows that the majority of these report positive effects of wealth, although levels of fertility seem to fall below those that would maximize fitness. We emphasize that reproductive decision-making reflects a complex interplay between individual and societal factors that resists simple evolutionary interpretation, and highlight the role of economic insecurity in fertility decisions. We conclude by discussing whether the wealth–fertility relationship can inform us about the adaptiveness of modern fertility behaviour, and argue against simplistic claims regarding maladaptive behaviour in humans.     

Fitness and fertility among Kalahari !Kung

In this paper we develop a model that examines fertility and childhood mortality patterns and their relationship to environmental variables. Interactions among environmental variables can account for different fertility patterns and different mixes of these variables can produce similar patterns of fertility. Our model attempts to quantify the idea that there is a trade-off between producing a few children likely to survive to reproductive age and producing a greater number of children with lower chances for survival. The optimum mix of these strategies depends on environmental characteristics. We use the model to make predictions about fertility and mortality patterns among two Bushmen populations of southern Africa--the Ghanzi and Ngamiland !Kung--using data collected by Harpending in 1967-1968. The results do not support explanations of the low fertilities observed among !Kung Bushmen women, in whom it is thought that fitness is maximized by limiting fertility, and show no relationship between mortality and family size in either !Kung population. Instead, the number of offspring reaching reproductive age in both populations increases as their completed family size increases. We examine the effects of sex, birth order, and paternal investment on mortality. No sex ratio differences and no differences in mortality by sex or birth order are present. Infant mortality among women who married more than once is significantly higher than among women who married once, suggesting that paternal care has a significant effect.     

Maternal risk of breeding failure remained low throughout the demographic transitions in fertility and age at first reproduction in Finland

Radical declines in fertility and postponement of first reproduction during the recent human demographic transitions have posed a challenge to interpreting human behaviour in evolutionary terms. This challenge has stemmed from insufficient evolutionary insight into individual reproductive decision-making and the rarity of datasets recording individual long-term reproductive success throughout the transitions. We use such data from about 2,000 Finnish mothers (first births: 1880s to 1970s) to show that changes in the maternal risk of breeding failure (no offspring raised to adulthood) underlay shifts in both fertility and first reproduction. With steady improvements in offspring survival, the expected fertility required to satisfy a low risk of breeding failure became lower and observed maternal fertility subsequently declined through an earlier age at last reproduction. Postponement of the age at first reproduction began when this risk approximated zero-even for mothers starting reproduction late. Interestingly, despite vastly differing fertility rates at different stages of the transitions, the number of offspring successfully raised to breeding per mother remained relatively constant over the period. Our results stress the importance of assessing the long-term success of reproductive strategies by including measures of offspring quality and suggest that avoidance of breeding failure may explain several key features of recent life-history shifts in industrialized societies.     

Why do large mothers produce large offspring? Theory and a test

To explain the general tendency of large mothers to produce large offspring, we developed two models in which either the rate at which each single offspring extracts resources from the mother or the rate at which the mother supplies resources to all the offspring is limited (terminal- or upper-stream-limitation on resource transport, respectively). We also reanalyzed the data of Erythronium japonicum to test the models. The terminal-stream-limitation model predicted that the optimal offspring size that maximizes the fitness of the mother increases with an increase in the maximum rate of resource extraction by each single offspring. Thus, large mothers produce large offspring if the maximum resource extraction rate is high in those mothers. The upper-stream-limitation model predicted that the optimal offspring size decreases with an increase in the maximum rate of resource supply by the mother to all the offspring. In E. japonicum, the maximum growth rate of a seed was independent of the number of seeds of a plant, suggesting that the resource extraction rate is limited at the individual seed level. The maximum growth rate was high in large plants and had a strong positive effect on final seed mass. Thus, the results were consistent with the terminal-stream-limitation model.     

Determinants of family size in rural Glacia (Spain)

The determinants of complete family size and offspring survival were assessed by means of family reconstitution data from a 19th & 20th century rural population of Northwest Spain. The reproductive performances of families were submitted to a stepwise regression analysis in which the number of children born alive and those surviving to reproductive age per family were taken as dependent variables, while 13 other variables were defined as independent. Infant mortality appears to explain most fertility variation; an earlier marital age and at first maternity leads to an increased family size. Survival seems to be the result of the apposite interaction of both fertility and mortality in early childhood.     

Home Range Size Variation in Female Arctic Grizzly Bears Relative to Reproductive Status and Resource Availability

The area traversed in pursuit of resources defines the size of an animal’s home range. For females, the home range is presumed to be a function of forage availability. However, the presence of offspring may also influence home range size due to reduced mobility, increased nutritional need, and behavioral adaptations of mothers to increase offspring survival. Here, we examine the relationship between resource use and variation in home range size for female barren-ground grizzly bears (Ursus arctos) of the Mackenzie Delta region in Arctic Canada. We develop methods to test hypotheses of home range size that address selection of cover where cover heterogeneity is low, using generalized linear mixed-effects models and an information-theoretic approach. We found that the reproductive status of female grizzlies affected home range size but individually-based spatial availability of highly selected cover in spring and early summer was a stronger correlate. If these preferred covers in spring and early summer, a period of low resource availability for grizzly bears following den-emergence, were patchy and highly dispersed, females travelled farther regardless of the presence or absence of offspring. Increased movement to preferred covers, however, may result in greater risk to the individual or family.     

Calcium Availability Influences Litter Size and Sex Ratio in White-Footed Mice (Peromyscus leucopus)

The production of offspring typically requires investment of resources derived from both the environment and maternal somatic reserves. As such, the availability of either of these types of resources has the potential to limit the degree to which resources are allocated to reproduction. Theory and empirical studies have argued that mothers modify reproductive performance relative to exogenous resource availability and maternal condition by adjusting size, number or sex of offspring produced. These relationships have classically been defined relative to availability of energy sources; however, in vertebrates, calcium also plays a critical role in offspring production, as a considerable amount of calcium is required to support the development of offspring skeleton(s). We tested whether the availability of calcium influences reproductive output by providing female white-footed mice with a low-calcium or standard diet from reproductive maturity to senescence. We then compared maternal skeletal condition and reproductive output, based on offspring mass, offspring number and litter sex ratio, between dietary treatments. Mothers on the low-calcium diet exhibited diminished skeletal condition at senescence and produced smaller and strongly female-biased litters. We show that skeletal condition and calcium intake can influence sex ratio and reproductive output following general theoretical models of resource partitioning during reproduction.     

How should parents adjust the size of their young in response to local environmental cues?

Models of parental investment typically assume that populations are well mixed and homogeneous and have devoted little attention to the impact of spatial variation in the local environment. Here, in a patch‐structured model with limited dispersal, we assess to what extent resource‐rich and resource‐poor mothers should alter the size of their young in response to the local environment in their patch. We show that limited dispersal leads to a correlation between maternal and offspring environments, which favours plastic adjustment of offspring size in response to local survival risk. Strikingly, however, resource‐poor mothers are predicted to respond more strongly to local survival risk, whereas resource‐rich mothers are predicted to respond less strongly. This lack of sensitivity on the part of resource‐rich mothers is favoured because they accrue much of their fitness through dispersing young. By contrast, resource‐poor mothers accrue a larger fraction of their fitness through philopatric young and should therefore respond more strongly to local risk. Mothers with more resources gain a larger share of their fitness through dispersing young partly because their fitness in the local patch is constrained by the limited number of local breeding spots. In addition, when resource variation occurs at the patch level, the philopatric offspring of resource‐rich mothers face stronger competition from the offspring of other local mothers, who also enjoy abundant resources. The effect of limited local breeding opportunities becomes less pronounced as patch size increases, but the impact of patch‐level variation in resources holds up even with many breeders per patch.     

Intergenerational transmission of fertility in Spain among cohorts born during the first half of twentieth century

It is known that historically fertility is correlated between generations of the same family. These links tend to be explained either in terms of the biogenetic determinants of reproduction or by the transmission of intra-familial values associated with reproduction and family life. Less is known about the micro-determinants of these links or about the extent to which the progressive modernization of reproductive outcomes over the past century has affected behavior. In this paper, we will address these issues for Spain with data from the Socio-Demographic Survey (SDS) carried out in 1991 and including data on cohorts born between 1900 and 1946. These data enable us to explore the micro determinants of fertility at different points of time during this period. Our results point to the existence of a significant correlation between intergenerational reproductive outcomes that persists and strengthens throughout this period of demographic change. Results confirm the importance of birth order in large family groups where firstborn offspring are more likely to have larger families than subsequent siblings. There is also evidence that the strength of these intergenerational ties increases with the onset of more modern demographic behavior characterized by sharply declining fertility. The results presented here promise to condition future debates on the subject.