The risks of parenthood. I. General theory and applications

Summary We use contemporary life history theory to analyze parental decisions concerned with the defense of offspring, and with the provisioning of offspring in the presence of risk. In achieving the optimal level of parental investment, the parent faces a tradeoff between present and future reproductive success. Our models, which are based on stochastic dynamic programming, lead to predictions of the following kind: (i) offspring will be defended more vigorously as they grow older; (ii) long-lived species will accept fewer risks in caring for offspring than short-lived species; (iii) parents living in permanently riskier environments will defend their offspring more vigorously than parents in less risky environments; (iv) however, temporary increases in risk will result in temporarily less vigorous defense and provisioning of offspring. The models also show that parents and their offspring have different conceptions of the optimal level of parental investment. The flexibility of our modeling approach as a method of analyzing facultative behavior is also emphasized. Finally, we apply the methods of this paper to analyze fledging behavior of Atlantic puffins.     

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.     

Fertility selection,genetic selection and evolution

The relationship between fertility selection as measured by the correlation in progeny number between parents and offspring, and selection at individual loci is investigated in humans. Estimates for the magnitude of fertility selection (0.1) and the rate of gene substitution (0.5 gene substitutions per generation per genome) are used in various mathematical models for selection. It is found that the observed magnitude of fertility selection cannot be explained by non‐epistatic directional selection at individual loci. A symmetric quantitative directional selection model is consistent with the observed data. But it is possible that fertility selection does not have a genetic basis.     

Marriage patterns in a Mesoamerican peasant community are biologically adaptive

Differential investment in offspring by parental and progeny gender has been discussed and periodically analyzed for the past 80 years as an evolutionary adaptive strategy. Parental investment theory suggests that parents in poor condition have offspring in poor condition. Conversely, parents in good condition give rise to offspring in good condition. As formalized in the Trivers-Willard hypothesis (TWH), investment in daughters will be greater under poor conditions while sons receive greater parental investment under good conditions. Condition is ultimately equated to offspring reproductive fitness, with parents apparently using a strategy to maximize their genetic contribution to future generations. Analyses of sex ratio have been used to support parental investment theory and in many instances, though not all, results provide support for TWH. In the present investigation, economic strategies were analyzed in the context of offspring sex ratio and survival to reproductive age in a Zapotec-speaking community in the Valley of Oaxaca, southern Mexico. Growth status of children, adult stature, and agricultural resources were analyzed as proxies for parental and progeny condition in present and prior generations. Traditional marriage practice in Mesoamerican peasant communities is patrilocal postnuptial residence with investments largely favoring sons. The alternative, practiced by ～25% of parents, is matrilocal postnuptial residence which is an investment favoring daughters. Results indicated that sex ratio of offspring survival to reproductive age was related to economic strategy and differed significantly between the patrilocal and matrilocal strategies. Variance in sex ratio was affected by condition of parents and significant differences in survival to reproductive age were strongly associated with economic strategy. While the results strongly support TWH, further studies in traditional anthropological populations are needed.     

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.     

Transgenerational acclimatization in an herbivore–host plant relationship

Twenty years ago, scientists began to recognize that parental effects are one of the most important influences on progeny phenotype. Consequently, it was postulated that herbivorous insects could produce progeny that are acclimatized to the host plant experienced by the parents to improve progeny fitness, because host plants vary greatly in quality and quantity, and can thus provide important cues about the resources encountered by the next generation. However, despite the possible profound implications for our understanding of host-use evolution of herbivores, host-race formation and sympatric speciation, intense research has been unable to verify transgenerational acclimatization in herbivore–host plant relationships. We reared Coenonympha pamphilus larvae in the parental generation (P) on high- and low-quality host plants, and reared the offspring (F₁) of both treatments again on high- and low-quality plants. We tested not only for maternal effects, as most previous studies, but also for paternal effects. Our results show that parents experiencing predictive cues on their host plant can indeed adjust progeny''s phenotype to anticipated host plant quality. Maternal effects affected female and male offspring, whereas paternal effects affected only male progeny. We here verify, for the first time to our knowledge, the long postulated transgenerational acclimatization in an herbivore–host plant interaction.     

"Trans-generational immune priming": specific enhancement of the antimicrobial immune response in the mealworm beetle, Tenebrio molitor

Encounters with parasites and pathogens are often unpredictable in time. However, experience of an infection may provide the host with reliable cues about the future risk of infection for the host itself or for its progeny. If the parental environment predicts the quality of the progeny's environment, then parents may further enhance their net reproductive success by differentially providing their offspring with phenotypes to cope with potential hazards such as pathogen infection. Here, I test for the occurrence of such an adaptive transgenerational phenotypic plasticity in the mealworm beetle, Tenebrio molitor. A pathogenic environment was mimicked by injection of bacterial lipopolysaccharides for two generations of insects. I found that parental challenge enhanced offspring immunity through the inducible production of antimicrobial peptides in the haemolymph.     

Parental quality of life in the framework of paediatric chronic gastrointestinal disease

Chronic gastrointestinal diseases, such as inflammatory bowel disease (IBD) and celiac disease (CD), might have impact not only on the affected child but also on their parents since inside the family framework, a change in each member may have influence on the whole system. The aim of this study was to test the hypothesis that parents whose children have IBD or CD will have a lower quality of life (QoL) compared to parents of healthy children, as well as that lower QoL will be found among parents who perceive lower health related quality of life (HRQOL) of their child. 55 parents of children with IBD, 49 of CD and 53 parents of healthy children were included in the study. Children were aged 7-18 years. In order to measure parental QoL, a Croatian version of the WHOQOL-BREF questionnaire was administered, with four domains: physical health, psychological health, social relationship and environment. The Total QoL was calculated as a sum of all domain items. The parent-proxy report of children's HRQOL, PedsQL 4.0 Generic Core Scales was used. Main result shows that parents of children with IBD report a significantly lower psychological health, compared to parents of other children, and significantly lower physical health compared to parents of healthy children. Among parents of children with CD and parents of healthy children, better parental Total QoL was significantly correlated with better parental report of children's HRQOL, while for parents of children with IBD those associations were not found. Results of study show association between presence of the disease in offspring with poorer parental QoL, thus highlight the importance of supporting not only children with chronic disease but also involving their parents in psychosocial interventions, as well as supporting the patient's association groups which gather both children with chronic disease and their parents.     

Transnational families and the subjective well-being of migrant parents: Angolan and Nigerian parents in the Netherlands

Studies on transnational families argue that the subjective well-being of migrant parents is negatively affected by living separated from their children. Most studies employ qualitative methods without a control group and hence are not able to distinguish whether the effects found are associated with separation only or with other factors. This paper investigates the association between parental subjective well-being and parent–child separation by comparing migrant parents who have at least one child in their country of origin with those who live with all their children in the Netherlands. The paper further investigates whether the same associations are found between migrant groups from Angola and Nigeria. Results indicate that transnational parents indeed report lower subjective well-being, as measured by happiness, life satisfaction and mental health for both groups and additionally, self-assessed health for Angolans. However, legal status, socio-economic status and the quality of the parent–child relationship are found to be important mediators.     

The delineation of fertility strategies in a tribal population of India: the Koyas of Koraput District,Orissa

Demographic data collected for a tribal population of India, the Koyas of Koraput District, Orissa, were examined in light of 2 models of reproductive behavior associated with the economic value of children: the replacement effect and son survivorship motivation. Both models are united in the concept that infant/child mortality affects subsequent fertility. The database consists of retrospective fertility histories of Koya women who had completed their reproductive period. The total number was 260, with the total offspring numbering 1407. 2 distinct cohorts of women were formed for the purpose of analysis, separated only by the criterion of offspring survival: women who had experienced infant child mortality (129 women with 739 children); and women who completed their reproductive period without suffering offspring loss of this nature (132 women with 668 children). The cohort without child loss had a mean parity of 5.10, lower than the average parity of 5.73 recorded for the cohort whose reproductive histories included at least 1 infant/child death. Age specific marital fertility and birth interval analyses indicated that this differential was because of biological, not behavioral, factors. The age pattern of fertility of females suffering offspring mortality failed to demonstrate a high rate of childbearing in the later age intervals of the reproductive period, a characteristic pattern of couples attempting to "replace" lost offspring. Birth interval analysis pointed to biological "interval effect," whereby infant/child mortality caused a cessation of lactation and hence a shortening of postpartum amenorrhea. Computer simulation further indicated that the higher fertility differential of the cohort experiencing offspring loss still did not result in high son survivorship values. The findings agree with earlier studies indicating that for predemographic transitional populations, economically motivated fertility strategies are ineffectual.     

Causal parenthood and the ethics of gamete donation

According to the causal theory of parenthood, people incur parental obligations by causing children to exist. Proponents of the causal theory often argue that gamete donors have special obligations to their genetic offspring. In response, many defenders of current gamete donation practices would reject the causal theory. In particular, they may invoke the ‘too many parents problem’: many people who causally contribute to the existence of children – for instance, fertility doctors – do not thereby incur parental obligations. This article argues that the conclusions commonly drawn by causal theorists, and by their critics, are premature. Causal theorists have a promising response to the too many parents problem. This response, however, defuses the moral concern that many causal theorists have raised about gamete donation. A similar point, it is argued, applies to Rivka Weinberg's ‘Hazmat Theory’.     

Carry-over effect of host nutritional quality on performance of spruce budworm progeny

The effect of host nutritional quality on spruce budworm (Choristoneura fumiferana (Clemens)) parental and offspring performance was studied using field and laboratory rearing experiments, and foliar chemical analyses. Foliage of balsam fir (Abies balsamea (L.) Mill.), white spruce (Picea glauca (Moench) Voss) and black spruce (P. mariana (Mill.) BSP) was used to rear the parental generation in the field, whereas an artificial diet was used to rear the progeny under laboratory conditions. Important differences in the food quality were provided by the three hosts. Black spruce foliage had higher concentrations of certain monoterpene deterrents and total phenolics, together with stronger seasonal declines in nutrients such as N, P and Mg, compared with the other hosts. We hypothesise that this trend may be related to poor performance and survival of the progeny. Laboratory rearing showed that progeny of parents that fed on black spruce exhibited longer developmental times and greater mortality, and had lower pupal mass than progeny of parents fed on the other hosts. Further, artificial food-fed progeny of insects reared on black spruce reached sixth-instar later, with lower mass, and exhibited higher relative growth rate (RGR) than progeny of parents fed on the other hosts. These results suggest nutritionally-based parental effects. These results also confirmed that the quality of food consumed by the parents can influence the fitness of the next generation.     

Leaf-rust resistance in rye (<Emphasis Type="Italic">Secale cereale</Emphasis> L.). 2. Genetic analysis and mapping of resistance genes <Emphasis Type="Italic">Pr3</Emphasis>, <Emphasis Type="Italic">Pr4</Emphasis>, and <Emphasis Type="Italic">Pr5</Emphasis>

The conventional way to drive modifications in old forest tree seed orchards is to establish progeny trials involving each parent tree and then evaluate its contribution to the performance of the progeny by estimating its general and specific combining ability (GCA and SCA). In this work, we successfully applied an alternative parent selection tactic based on paternity testing of superior offspring derived from a hybrid seed orchard established with a single Eucalyptus grandis seed parents and six E. urophylla pollen parents. A battery of 14 microsatellite markers was used to carry out parentage tests of 256 progeny individuals including two independent samples of selected trees and one control unselected sample, all derived from 6-year-old forest stands in eastern Brazil. Paternity determination was carried out for all progeny individuals by a sequential paternity exclusion procedure. Exclusion was declared only when the obligatory paternal allele in the progeny tree was not present in the alleged parent tree for at least four independent markers to avoid false exclusions due to mutation or null alleles. After maternity checks to identify seed mixtures and selfed individuals, the paternity tests revealed that approximately 29% of the offspring was sired by pollen parents outside the orchard. No selfed progeny were found in the selected samples. Three pollen parents were found to have sired essentially all of the offspring in the samples of selected and non-selected progeny individuals. One of these three parents sired significantly more selected progeny than unselected ones (P0.0002 in a Fisher exact test). Based on these results, low-reproductive-successful parents were culled from the orchard, and management procedures were adopted to minimize external pollen contamination. A significant difference (P<0.01) in mean annual increment was observed between forest stands produced with seed from the orchard before and after selection of parents and revitalization of the orchard. An average realized gain of 24.3% in volume growth was obtained from the selection of parents as measured in forest stands at age 2–4 years. The marker-assisted tree-breeding tactic presented herein efficiently identified top parents in a seed orchard and resulted in an improved seed variety. It should be applicable for rapidly improving the output quality of seed orchards, especially when an emergency demand for improved seed is faced by the breeder.Communicated by D.B. Neale     

WHAT DETERMINES FITNESS WHEN DISPERSAL IS LIMITED?

Simple evolutionary models typically assume a homogeneous environment in which all individuals have equal access to resources. However, when dispersal is limited this assumption is unlikely to correspond to reality. Instead, the offspring of relatively fecund parents can be expected to find that resources are scarce because of competition from their relatively large number of siblings. We show that these factors lead to selection for decreased variation in the number of offspring produced. We investigate the relationship between fitness and variation in fertility for a variety of different circumstances. Our results indicate that, in some cases, selection for reduced variation in fertility can be very substantial.     

Sexual selection theory meets disease vector control: Testing harmonic convergence as a “good genes” signal in Aedes aegypti mosquitoes

BackgroundThe mosquito Aedes aegypti is a medically important, globally distributed vector of the viruses that cause dengue, yellow fever, chikungunya, and Zika. Although reproduction and mate choice are key components of vector population dynamics and control, our understanding of the mechanisms of sexual selection in mosquitoes remains poor. In “good genes” models of sexual selection, females use male cues as an indicator of both mate and offspring genetic quality. Recent studies in Ae. aegypti provide evidence that male wingbeats may signal aspects of offspring quality and performance during mate selection in a process known as harmonic convergence. However, the extent to which harmonic convergence may signal overall inherent quality of mates and their offspring remains unknown.Methodology/Principal findingsTo examine this, we measured the relationship between acoustic signaling and a broad panel of parent and offspring fitness traits in two generations of field-derived Ae. aegypti originating from dengue-endemic field sites in Thailand. Our data show that in this population of mosquitoes, harmonic convergence does not signal male fertility, female fecundity, or male flight performance traits, which despite displaying robust variability in both parents and their offspring were only weakly heritable.Conclusions/SignificanceTogether, our findings suggest that vector reproductive control programs should treat harmonic convergence as an indicator of some, but not all aspects of inherent quality, and that sexual selection likely affects Ae. aegypti in a trait-, population-, and environment-dependent manner.     

Intralocus sexual conflict,adaptive sex allocation,and the heritability of fitness

Intralocus sexual conflict arises when selection favours alternative fitness optima in males and females. Unresolved conflict can create negative between‐sex genetic correlations for fitness, such that high‐fitness parents produce high‐fitness progeny of their same sex, but low‐fitness progeny of the opposite sex. This cost of sexual conflict could be mitigated if high‐fitness parents bias sex allocation to produce more offspring of their same sex. Previous studies of the brown anole lizard (Anolis sagrei) show that viability selection on body size is sexually antagonistic, favouring large males and smaller females. However, sexual conflict over body size may be partially mitigated by adaptive sex allocation: large males sire more sons than daughters, whereas small males sire more daughters than sons. We explored the evolutionary implications of these phenomena by assessing the additive genetic (co)variance of fitness within and between sexes in a wild population. We measured two components of fitness: viability of adults over the breeding season, and the number of their progeny that survived to sexual maturity, which includes components of parental reproductive success and offspring viability (RS^V). Viability of parents was not correlated with adult viability of their sons or daughters. RS^V was positively correlated between sires and their offspring, but not between dams and their offspring. Neither component of fitness was significantly heritable, and neither exhibited negative between‐sex genetic correlations that would indicate unresolved sexual conflict. Rather, our results are more consistent with predictions regarding adaptive sex allocation in that, as the number of sons produced by a sire increased, the adult viability of his male progeny increased.     

Better‐surviving barn swallow mothers produce more and better‐surviving sons

Sex allocation theory predicts that parents are selected to bias their progeny sex ratio (SR) toward the sex that will benefit the most from parental quality. Because parental quality may differentially affect survival of sons and daughters, a pivotal test of the adaptive value of SR adjustment is whether parents overproduce offspring of the sex that accrues larger fitness advantages from high parental quality. However, this crucial test of the long‐term fitness consequences of sex allocation decisions has seldom been performed. In this study of the barn swallow (Hirundo rustica), we showed a positive correlation between the proportion of sons and maternal annual survival. We then experimentally demonstrated that this association did not depend on the differential costs of rearing offspring of either sex. Finally, we showed that maternal lifespan positively predicted lifespan of sons but not of daughters. Because in barn swallows lifespan is a strong determinant of lifetime reproductive success, the results suggest that mothers overproduce offspring of the sex that benefits the most from maternal quality. Hence, irrespective of mechanisms causing the SR bias and mother–son covariation in lifespan, we provide strong evidence that sex allocation decisions of mothers can highly impact on their lifetime fitness.     

Parental age difference, educationally assortative mating and offspring count: evidence from a contemporary population in Taiwan

Using contemporary population data from Taiwan, we examine the relationships between parental age difference, educationally assortative mating, income and offspring count. Controlling for women's reproductive value (measured by age at first birth), we find that an older husband is associated with fewer offspring, whereas a husband with similar or higher education is associated with more offspring. Concerning resources, we find that women's income is negatively associated with fertility and husband's income is positively associated with fertility among highly educated women. These results are consistent with the view that women compensate for trade-offs between education, income generation and childbearing by seeking mates with a higher status.     

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.