Predictors of Grandparental Investment Decisions in Contemporary Europe: Biological Relatedness and Beyond

Across human cultures, grandparents make a valued contribution to the health of their families and communities. Moreover, evidence is gathering that grandparents have a positive impact on the development of grandchildren in contemporary industrialized societies. A broad range of factors that influence the likelihood grandparents will invest in their grandchildren has been explored by disciplines as diverse as sociology, economics, psychology and evolutionary biology. To progress toward an encompassing framework, this study will include biological relatedness between grandparents and grandchildren, a factor central to some discipline''s theoretical frameworks (e.g., evolutionary biology), next to a wide range of other factors in an analysis of grandparental investment in contemporary Europe. This study draws on data collected in the Survey of Health, Ageing and Retirement in Europe from 11 European countries that included 22,967 grandparent–child dyads. Grandparents reported biological relatedness, and grandparental investment was measured as the frequency of informal childcare. Biological and non-biological grandparents differed significantly in a variety of individual, familial and area-level characteristics. Furthermore, many other economic, sociological, and psychological factors also influenced grandparental investment. When they were controlled, biological grandparents, relative to non-biological grandparents, were more likely to invest heavily, looking after their grandchildren almost daily or weekly. Paradoxically, however, they were also more likely to invest nothing at all. We discuss the methodological and theoretical implications of these findings across disciplines.     

Grandparental investment and reproductive decisions in the longitudinal 1970 British cohort study

There has been a recent increase in interest among evolutionary researchers in the hypothesis that humans evolved as cooperative breeders, using extended family support to help decrease offspring mortality and increase the number of children that can be successfully reared. In this study, data drawn from the 1970 longitudinal British cohort study were analysed to determine whether extended family support encourages fertility in contemporary Britain. The results showed that at age 30, reported frequency that participants saw their own parents (but not in-laws) and the closeness of the bond between the participant and their own parents were associated with an increased likelihood of having a child between ages 30 and 34. Financial help and reported grandparental childcare were not significantly positively associated with births from age 30 to 34. Men's income was positively associated with likelihood of birth, whereas women's income increased likelihood of birth only for working women with at least one child. While it was predicted that grandparental financial and childcare help would increase the likelihood of reproduction by lowering the cost to the parent of having a child, it appears that the mere physical presence of supportive parents rather than their financial or childcare help encouraged reproduction in the 1970 British birth cohort sample.     

How Grandparents Matter

Low birth rates in developed societies reflect women’s difficulties in combining work and motherhood. While demographic research has focused on the role of formal childcare in easing this dilemma, evolutionary theory points to the importance of kin. The cooperative breeding hypothesis states that the wider kin group has facilitated women’s reproduction during our evolutionary history. This mechanism has been demonstrated in pre-industrial societies, but there is no direct evidence of beneficial effects of kin’s support on parents’ reproduction in modern societies. Using three-generation longitudinal data anchored in a sample of grandparents aged 55 and over in 1992 in the Netherlands, we show that childcare support from grandparents increases the probability that parents have additional children in the next 8 to 10 years. Grandparental childcare provided to a nephew or niece of childless children did not significantly increase the probability that those children started a family. These results suggest that childcare support by grandparents can enhance their children’s reproductive success in modern societies and is an important factor in people’s fertility decisions, along with the availability of formal childcare.     

The effect of paternal investment on female fertility intention in South Korea

Life history theory views reproduction as an outcome of resource allocation. The allocation of resources such as parental investments of time, energy and material resources involves trade-offs between number of offspring and timing of reproduction. Within the framework of mammalian parental investment, the outstanding feature of human reproduction is the high level of paternal care. Although empirical evidence suggests that human paternal investment may have evolved as a reproductive strategy to reduce infant and child mortality rates, the effects of actual paternal investment, including allocating time to child care, on female reproductive decisions have received relatively little attention. We examined the trade-off from two perspectives using a representative sample of married South Korean women aged 20–44 in 2005 (n=977). First, paternal investment in domestic labor, including child care and housework, was expected to be associated with women's preference regarding future reproduction. Second, relative paternal investment was expected to increase women's preference for future reproduction, especially among employed women. We found that increased paternal investment in child care and housework remarkably enhanced women's intention to have a second child, especially among employed women. In addition, although family members provide a low percentage of child care in South Korea, such help is likely to be a useful resource for second childbirth among employed women. Somewhat expectedly, older age and longer time since first birth had negative effects on women's second-child intention. There is growing evidence that, in the lowest fertility societies, paternal investment may be an essential resource for promoting future reproductive behavior of women, especially employed women.     

Do grandparents favor granddaughters? Biased grandparental investment in UK

Differential grandparental investment in grandchildren is often explained with paternity uncertainty. The asymmetric inheritance of the sex chromosomes, especially of the X chromosomes, may also bias grandparental investment. Recent studies show that ignoring the sex of the grandchild can mask important differences in the investment patterns of the same grandparent category, but this has not been tested in contemporary societies with nationally representative data. With 17 variables from the Involved Grandparenting and Child Well-Being 2007 survey, we tested differential grandparental investment as reported by British and Welsh adolescents and compared predictions based on X-chromosomal relatedness with predictions based on paternity uncertainty. The theories are expected to differ with regard to grandmaternal investment in grandsons and granddaughters. We test whether paternal grandmothers invest (H1) more in granddaughters than in grandsons, (H2) more in granddaughters than maternal grandmothers do and (H3) less in grandsons than maternal grandmothers do. In addition, following the suggestion that paternal grandmothers may reduce sibling competition between girls and boys by harming grandsons, we study whether (H4) paternal grandmothers channel more noninvestment into grandsons than into granddaughters. The results show no convincing support for the type of sex discrimination of grandchildren that is predicted by X-chromosomal relatedness theories, but do provide support for the paternity uncertainty theory. X-chromosomal relatedness does not appear to shape grandparental behavior in developed societies.     

Studying parents and grandparents to assess genetic contributions to early-onset disease

Suppose DNA is available from affected individuals, their parents, and their grandparents. Particularly for early-onset diseases, maternally mediated genetic effects can play a role, because the mother determines the prenatal environment. The proposed maximum-likelihood approach for the detection of apparent transmission distortion treats the triad consisting of the affected individual and his or her two parents as the outcome, conditioning on grandparental mating types. Under a null model in which the allele under study does not confer susceptibility, either through linkage or directly, and when there are no maternally mediated genetic effects, conditional probabilities for specific triads are easily derived. A log-linear model permits a likelihood-ratio test (LRT) and allows the estimation of relative penetrances. The proposed approach is robust against genetic population stratification. Missing-data methods permit the inclusion of incomplete families, even if the missing person is the affected grandchild, as is the case when an induced abortion has followed the detection of a malformation. When screening multiple markers, one can begin by genotyping only the grandparents and the affected grandchildren. LRTs based on conditioning on grandparental mating types (i.e., ignoring the parents) have asymptotic relative efficiencies that are typically >150% (per family), compared with tests based on parents. A test for asymmetry in the number of copies carried by maternal versus paternal grandparents yields an LRT specific to maternal effects. One can then genotype the parents for only the genes that passed the initial screen. Conditioning on both the grandparents' and the affected grandchild's genotypes, a third log-linear model captures the remaining information, in an independent LRT for maternal effects.     

Ancestral feeding and survival of offspring in European corn borer

Experiments were conducted to evaluate the influence of ancestral adult feeding by European corn borer, Ostrinia nubilalis (Hübner) (Lepidoptera: Crambidae), on larval survival in the field and neonate movement behavior in laboratory. Larval survival was higher when either the grandparental or parental generation had fed, but the feeding sites of the surviving larvae were not affected by ancestral feeding condition. This is the first evidence that grandparental feeding could influence larval survival in the field. Larval movement was observed in the laboratory. Silking speed of neonates was faster when either grandparents or parents had fed, while walking speed was faster only when parents had fed. No broad-sense genetic correlation was found between silking speed and walking speed. Broad-sense heritability among feeding histories were not significant for silking speed, but was significantly greater than zero for walking speed when grandparents fed and parents did not. These intergenerational effects could induce complex population dynamics in this species.     

Status competition,inequality, and fertility: implications for the demographic transition

The role that social status plays in small-scale societies suggests that status may be important for understanding the evolution of human fertility decisions, and for understanding how such decisions play out in modern contexts. This paper explores whether modelling competition for status—in the sense of relative rank within a society—can help shed light on fertility decline and the demographic transition. We develop a model of how levels of inequality and status competition affect optimal investment by parents in the embodied capital (health, strength, and skills) and social status of offspring, focusing on feedbacks between individual decisions and socio-ecological conditions. We find that conditions similar to those in demographic transition societies yield increased investment in both embodied capital and social status, generating substantial decreases in fertility, particularly under conditions of high inequality and intense status competition. We suggest that a complete explanation for both fertility variation in small-scale societies and modern fertility decline will take into account the effects of status competition and inequality.     

Kin and birth order effects on male child mortality: three East Asian populations, 1716–1945

Human child survival depends on adult investment, typically from parents. However, in spite of recent research advances on kin influence and birth order effects on human infant and child mortality, studies that directly examine the interaction of kin context and birth order on sibling differences in child mortality are still rare. Our study supplements this literature with new findings from large-scale individual-level panel data for three East Asian historical populations from northeast China (1789–1909), northeast Japan (1716–1870), and north Taiwan (1906–1945), where preference for sons and first-borns is common. We examine and compare male child mortality risks by presence/absence of co-resident parents, grandparents, and other kin, as well as their interaction effects with birth order. We apply discrete-time event-history analysis on over 172,000 observations of 69,125 boys aged 1–9 years old. We find that in all three populations, while the presence of parents is important for child survival, it is more beneficial to first/early-borns than to later-borns. Effects of other co-resident kin are however null or inconsistent between populations. Our findings underscore the importance of birth order in understanding how differential parental investment may produce child survival differentials between siblings.     

Adaptive transgenerational plasticity in an annual plant: grandparental and parental drought stress enhance performance of seedlings in dry soil

Stressful parental (usually maternal) environments can dramatically influence expression of traits in offspring, in some cases resulting in phenotypes that are adaptive to the inducing stress. The ecological and evolutionary impact of such transgenerational plasticity depends on both its persistence across generations and its adaptive value. Few studies have examined both aspects of transgenerational plasticity within a given system. Here we report the results of a growth-chamber study of adaptive transgenerational plasticity across two generations, using the widespread annual plant Polygonum persicaria as a naturally evolved model system. We grew five inbred Polygonum genetic lines in controlled dry vs. moist soil environments for two generations in a fully factorial design, producing replicate individuals of each genetic line with all permutations of grandparental and parental environment. We then measured the effects of these two-generational stress histories on traits critical for functioning in dry soil, in a third (grandchild) generation of seedling offspring raised in the dry treatment. Both grandparental and parental moisture environment significantly influenced seedling development: seedlings of drought-stressed grandparents or parents produced longer root systems that extended deeper and faster into dry soil compared with seedlings of the same genetic lines whose grandparents and/or parents had been amply watered. Offspring of stressed individuals also grew to a greater biomass than offspring of nonstressed parents and grandparents. Importantly, the effects of drought were cumulative over the course of two generations: when both grandparents and parents were drought-stressed, offspring had the greatest provisioning, germinated earliest, and developed into the largest seedlings with the most extensive root systems. Along with these functionally appropriate developmental effects, seedlings produced after two previous drought-stressed generations had significantly greater survivorship in very dry soil than did seedlings with no history of drought. These findings show that plastic responses to naturalistic resource stresses experienced by grandparents and parents can "preadapt" offspring for functioning under the same stresses in ways that measurably influence realized fitness. Possible implications of these environmentally-induced, inherited adaptations are discussed with respect to ecological distribution, persistence under novel stresses, and evolution in natural populations.     

Alternatives to the Grandmother Hypothesis

We conducted a meta-analysis of 17 studies that tested for an association between grandparental survival and grandchild survival in patrilineal populations. Using two different methodologies, we found that the survival of the maternal grandmother and grandfather, but not the paternal grandmother and grandfather, was associated with decreased grandoffspring mortality. These results are consistent with the findings of psychological studies in developed countries (Coall and Hertwig Behavioral and Brain Sciences 33:1-59, 2010). When tested against the predictions of five hypotheses (confidence of paternity; grandmothering, kin proximity, grandparental senescence, and local resource competition), our meta-analysis results are most in line with the local resource competition hypothesis. In patrilineal and predominantly patrilocal societies, the grandparents who are most likely to live with the grandchildren have a less beneficial association than those who do not. We consider the extent to which these results may be influenced by the methodological limitations of the source studies, including the use of retrospective designs and inadequate controls for confounding variables such as wealth.     

Grandparental investment as a function of relational uncertainty and emotional closeness with parents

Several theoretical perspectives have generated research on grandparental investment, notably socialization and evolutionary psychological perspectives. Using data collected from more than 200 older adults (mean age 67 years), we test three hypotheses derived from socialization and evolutionary perspectives about grandparents’ relationships with and investment in grandchildren. Results indicate that (1) emotional closeness with both children and children-in-law is positively related to reports of emotional closeness with grandchildren; (2) maternal grandmothers invest more in grandchildren than do other grandparents; and (3) grandparents invest more in daughters’ children than in sons’ children. Discussion addresses limitations of self-report methodology in this context and situates the current findings within the body of research indicating the existence of psychological mechanisms designed to guide differential nepotism.     

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.     

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.     

Kin Relationships and the Caregiving Biases of Grandparents, Aunts, and Uncles

Paternity certainty and matrilineal family ties have been used to explain the asymmetric caregiving of grandparents and aunts and uncles. The proximate mechanisms underlying biased kin investment, however, remain unclear. A central question of the study presented here was whether the parent-kin relationship is an important link in the caregiving. In a two-generational questionnaire study, we asked subjects to estimate the intensity of their relationships to parents, grandparents, aunts, and uncles (emotional closeness, investment received in childhood). In addition, the subjects’ parents rated their emotional closeness to their parents and siblings. We found that the parent-kin relationship was closely linked to the relatives’ child care and could partly explain asymmetric caregiving. Maternal aunts played a special role as caregivers. Especially the mother’s younger or last-born sister cared intensively for nieces and nephews, regardless of her closeness to the subjects’ mother.     

Whether to have a second child or not? An integrative approach to women's reproductive decision-making in current China

The transition to low fertility worldwide has led to introduction of diverse frameworks across disciplines to understand its causes and consequences. Previous attempts to compare the relative importance of the key factors influencing women's fertility decision-making largely focused on a single rather than multiple steps of decision-making—an important problem if different factors are associated with different steps. Furthermore, insufficient attention has been paid explicitly to husband's and already-born children's influences, two potentially important factors. Here we introduce a framework covering three steps of reproductive decision-making—ideal family size, fertility desire and fertility intention—and test it using multi-level survey data collected from Chinese one-child mothers. Mother's attitudes towards having two children were paramount factors underlying her ideal family size, and husband's and the firstborn child's attitudes were critical to her desire to have a second child, which in turn played a decisive role in her intention to have a second child. Although husband's attitude was related to all steps, most factors were only relevant to one step; e.g., perceived child mortality and value for old-age security predicted ideal family size, admiration—a prerequisite for social learning—for two-child families predicted fertility desire, and physical/economic constraints primarily predicted fertility intention. Our study emphasizes multiple decision-makers in family reproduction; indicates the relative importance of fertility-influencing factors could vary with steps of decision-making; and has important implications for population policy in low-fertility societies.     

Father Absence and Reproduction-Related Outcomes in Malaysia,a Transitional Fertility Population

Father absence is consistently associated with children’s reproductive outcomes in industrialized countries. It has been suggested that father absence acts as a cue to particular environmental conditions that influence life history strategies. Much less is known, however, about the effects of father absence on such outcomes in lower-income countries. Using data from the 1988 Malaysian Family Life Survey (n?=?567), we tested the effect of father absence on daughters’ age at menarche, first marriage, and first birth; parity progression rates; and desired completed family size in Malaysia, a country undergoing an economic and fertility transition. Father absence during later childhood (ages 8 to 15), although not during earlier childhood, was associated with earlier progressions to first marriage and first birth, after controlling for other confounders. Father absence does not affect age at menarche, desired family size, or progression from first to second birth. The patterns found in this transitional population partly mirror those in developed societies, where father absence accelerates reproductive events. There is, however, a notable contrast between the acceleration in menarche for father-absent girls consistently found in developed societies and the lack of any association in our findings. The mechanisms through which father absence affects reproduction may differ in different ecological contexts. In lower-income contexts, direct paternal investment or influence may be of more importance in determining reproductive behavior than whether fathers act as a cue to environmental conditions.     

On the impact of sex and birth order on contact with kin

Previous research indicates that birth order is a strong predictor of familial sentiments, with middleborns less family-oriented than first- or last-borns. In this research, effects of sex and birth order on the actual frequency of contact with maternal and paternal kin were examined in two studies. In Study 1, one hundred and forty undergraduates completed a questionnaire relating to the amount of time they spent in contact with specific relatives, while in Study 2, one hundred and twelve undergraduates completed the same questionnaire with the addition of two questions relating to the subjects’ parents’ birth orders. Subjects were more likely to have frequent contact with maternal, as opposed to paternal, kin and women experienced more frequent contact than men with relatives in general. The birth order of subjects did not appear to have a significant influence on contact but the birth order of the subjects’ parents did, with the offspring of middleborn mothers having relatively little contact with maternal grandparents and the offspring of middleborn fathers having relatively little contact with paternal grandparents. These sex and birth order differences are discussed in relation to possible differences in how women and men use kinship ties and in terms of how birth order may influence parental solicitude. Catherine Salmon recently received her Ph.D. in psychology from McMaster University in Hamilton Ontario. Her interest in kinship and family relationships has grown out of her own large extended family and many visits to Utah as well as exposure to evolutionary thinking about the family in the lab of Martin Daly and Margo Wilson. Her other current research interests focus on female sexuality and the evolutionary study of literature.     

Matriliny as daughter-biased investment

From an evolutionary perspective, matriliny presents a puzzle because men in matrilineal societies transmit wealth to their sisters' sons, to whom they are only half as related as to their own sons. It has been argued that such systems would only maximise fitness under unrealistically high levels of paternity uncertainty. In this paper, we propose that matriliny can arise from daughter-biased investment by parents and/or grandparents. We show that daughter-biased investment is adaptive if the marginal benefit of wealth to sons (compared to daughters) does not outweigh the risk of nonpaternity in sons' offspring. We argue that such conditions will be rare where resource-holding polygyny is prevalent but could otherwise be widespread under even moderate levels of paternity uncertainty. The daughter-biased investment model explains two well-known characteristics of matrilineal descent: (a) matriliny's association with high levels of paternity uncertainty and (b) matriliny's ecological correlates, including its association with horticulture, its rarity in pastoralist and agro-pastoralist societies, and the tendency for matriliny to be replaced by son-biased inheritance during economic development. We present data on wealth, sex, and reproductive success (RS) in two African societies, the matrilineal Chewa in Malawi and patrilineal Gabbra in Kenya, which support the daughter-biased investment theory.     

Causes and consequences of increase in child survival rates: ethnoepidemiology among the Hmong of Thailand.

The Hmong "hill tribe" minority in Thailand has much higher exposure to factors usually associated with risk of child mortality (high fertility, low status of women, low education, less use of modern medical care for births, exposure to warfare, economic and physical disruption, and poor hygienic conditions) than the rural ethnic Thai population. Nonetheless, infant mortality has declined from over 120 per 1000 to under 50 per 1000 live births among both these populations in the past 30 years. The reason for the rapid increase in child survival among the Hmong appears to be better access to and more use of modern curative and preventive medical care associated with road construction rather than major changes in social or hygienic conditions. Conventional wisdom suggests that high fertility is both a cause and a consequence of high infant and child mortality and that parents will not reduce fertility until they see that mortality has declined. Most Hmong parents recognize the decline in child mortality and attribute it to better access to modern medical care. Most Hmong parents also say that, if they were starting to have children now, they would want to have fewer children. Fear of child death is infrequently mentioned as a motive for having more children, and the perceived decline in child mortality is rarely mentioned as a reason for reduced fertility. Most Hmong parents explain their desired family size in terms of economic conditions rather than perceived risk of child mortality. Results of this study suggest that fertility and child mortality can vary independently of one another and that major reductions in child mortality can be accomplished without waiting for major social changes (e.g., improved education or status of women) or major reductions in fertility.