The Trivers–Willard hypothesis of parental investment: No effect in the contemporary United States

The Trivers–Willard hypothesis (TWH) predicts that parents will bias their sex ratio toward sons when in good condition and toward daughters when in poor condition. Many human studies have tested the related hypothesis that parents' bias allocation of resources to existing sons and daughters according to the same principle. The present study used time diary and self-report data from the parents of 3200 children in the US to test the hypothesis that as status increases, parents will allocate more resources to sons vs. daughters. It finds no evidence that higher-status parents invest more in sons or that lower status parents invest more in daughters. This finding illustrates the specificity of situations in which the TWH effects should be expected. Only certain types of parental investment — such as protection and a bias in the sex ratio — may have been selected to vary according to parental condition. Optimal allocation of resources after the child is born, however, is achieved not by the simple bias predicted by the TWH, but by allocating resources among offspring in ways that yield the largest marginal inclusive fitness gains.     

Parental investment by sex on ifaluk

Trivers and Willard (Science 179;90–92, 1973) predict that where investment by parents in good condition increases the fitness of sons more than that of their sisters, while the opposite is true of parents in poor condition, parents with much to invest will favor sons, while those with little will favor daughters. Patterns of parent-child association on Ifaluk atoll are consistent with this prediction.     

Low serum vitamin A mothers breastfeed daughters more often than sons in drought-ridden northern Kenya: a test of the Trivers–Willard hypothesis

The Trivers–Willard hypothesis predicts that natural selection should favor unequal parental investment between daughters and sons based upon maternal condition and offspring reproductive potential. Specifically, it predicts that mothers in good condition should increase investment toward sons, while mothers in poor condition should favor daughters. Previous tests of the hypothesis in human populations overwhelmingly focused on economic resources as maternal condition indicators. We test the Trivers–Willard hypothesis using maternal nutrition—energy and vitamin A status representing macro- and micronutrition, respectively—as the indicator for maternal condition, with breastfeeding frequency recalls serving as the indicator for parental investment. Data from exclusively breastfeeding mothers (n=83) in drought-ridden Ariaal agropastoral villages of northern Kenya were used to test the hypothesis that mothers in poor condition will breastfeed daughters more frequently than sons. Poor condition was defined as having a body mass index <18.5 or serum retinol (vitamin A) concentration <1.05 µmol/l. A linear regression model was applied using breastfeeding frequency as the dependent variable and respective maternal condition, infant's sex, and the maternal condition–infant's sex interaction as the predictors, controlling for covariates. Results supported the hypothesis only in the vitamin A model which predicts that low-vitamin-A mothers breastfeed daughters significantly more frequently than sons (11 vs. 6 times/day), while vitamin-A-sufficient mothers breastfeed daughters and sons equivalently (9 times). These results indicate that maternal nutritional status, particularly micronutrient status, can contribute to the investigation of the evolutionary hypothesis of sex-biased parental investment.     

In poor families, mothers' milk is richer for daughters than sons: A test of Trivers-Willard hypothesis in agropastoral settlements in Northern Kenya

The Trivers–Willard hypothesis predicts the unequal parental investment between daughters and sons, depending on maternal condition and offspring reproductive potential. Specifically, in polygynous populations where males have higher reproductive variance than females, it predicts that mothers in good condition will invest more in sons, whereas mothers in poor condition will invest more in daughters. Previous studies testing this hypothesis focused on behavioral investment, whereas few examined biological investment. This study investigates the Trivers–Willard hypothesis on both behavioral and biological parental investment by examining breastfeeding frequencies and breast milk fat concentrations. Data from exclusively breastfeeding mothers in Northern Kenya were used to test hypotheses: Economically sufficient mothers will breastfeed sons more frequently than daughters, whereas poor mothers will breastfeed daughters more frequently than sons, and economically sufficient mothers will produce breast milk with higher fat concentration for sons than daughters, whereas poor mothers will produce breast milk with higher fat concentration for daughters than sons. Linear regression models were applied, using breastfeeding frequency or log‐transformed milk fat as the dependent variable, and offspring's sex (son = 1/daughter = 0), socioeconomic status (higher = 1/lower = 0), and the sex‐wealth interaction as the predictors, controlling for covariates. Our results only supported the milk fat hypothesis: infant's sex and socioeconomic status interacted (P = 0.014, n = 72) in their relation with milk fat concentration. The model estimated that economically sufficient mothers produced richer milk for sons than daughters (2.8 vs. 0.6 gm/dl) while poor mothers produced richer milk for daughters than sons (2.6 vs. 2.3 gm/dl). Further research on milk constituents in relation to offspring's sex is warranted. Am J Phys Anthropol , 2012. © 2012 Wiley Periodicals, Inc.     

Maternal Investment of the Virunga Mountain Gorillas

The Trivers & Willard hypothesis (TWH) predicts that females with more resources should bias their maternal investment toward offspring of the sex that is most likely to benefit from those additional resources. This paper examines the sex allocation of 61 female mountain gorillas (Gorilla beringei beringei) of the Virunga volcanoes, Rwanda from 1967 to 2004. Like most highly dimorphic, polygynous mammals, mountain gorillas are expected to show greater variance in reproductive success among males than females, so mothers in good condition should bias their investment toward sons. Using dominance rank as the indicator of maternal condition, the TWH was tentatively supported by our results with interbirth intervals (IBI). Dominant mothers had longer IBI following the birth of sons, relative to the longer IBI that subordinate mothers had with daughters. In contrast, maternal condition did not have a significant effect on birth sex ratios. We also found no significant relationships with other variables that might influence birth sex ratios (e.g., maternal age, parity, or group size), and the overall birth sex ratio was not significantly different from a 50:50 split. Collectively, our results suggest that female mountain gorillas do not control the sex ratio of their offspring at birth, but they may adjust their subsequent maternal investment. This conclusion is consistent with recurring questions about whether any adjustments in birth sex ratios occur in primates.     

Paternal reproductive success drives sex allocation in a wild mammal

Parents should bias sex allocation toward offspring of the sex most likely to provide higher fitness returns. Trivers and Willard proposed that for polygynous mammals, females should adjust sex‐ratio at conception or bias allocation of resources toward the most profitable sex, according to their own body condition. However, the possibility that mammalian fathers may influence sex allocation has seldom been considered. Here, we show that the probability of having a son increased from 0.31 to 0.60 with sire reproductive success in wild bighorn sheep (Ovis canadensis). Furthermore, our results suggest that females fertilized by relatively unsuccessful sires allocated more energy during lactation to daughters than to sons, while the opposite occurred for females fertilized by successful sires. The pattern of sex‐biased offspring production appears adaptive because paternal reproductive success reduced the fitness of daughters and increased the average annual weaning success of sons, independently of maternal allocation to the offspring. Our results illustrate that sex allocation can be driven by paternal phenotype, with profound influences on the strength of sexual selection and on conflicts of interest between parents.     

Maternal Condition Does Not Influence Birth Sex Ratios in Anubis Baboons (Papio anubis)

Trivers and Willard predicted that when parental condition has differential effects on the fitness of male and female offspring, parents who are in good condition will bias investment toward the sex that benefits most from additional investment. Efforts to test predictions derived from Trivers and Willard''s model have had mixed results, perhaps because most studies have relied on proxy measures of parental condition, such as dominance rank. Here, we examine the effects of female baboons condition on birth sex ratios and post-natal investment, based on visual assessments of maternal body condition. We find that local environmental conditions have significant effects on female condition, but maternal condition at conception has no consistent relationship with birth sex ratios. Mothers who are in poorer condition at the time of conception resume cycling significantly later than females who are in better condition, but the sex of their infants has no effect on the time to resumption of cycling. Thus, our findings provide strong evidence that maternal condition influences females'' ability to reproduce, but females do not facultatively adjust the sex ratio of their offspring in relation to their dominance rank or current condition.     

Waiting for Trivers and Willard: Do the rich really favor sons?

Parental investment theory has been put forward as a major evolutionary argument explaining male or female biased birth sex ratio, the Trivers-Willard (T-W) hypothesis, predicting that parents living in good circumstances will bias their investment to sons, whereas parents in poor circumstances will bias their investment toward daughters. Tests of the T-W hypothesis on human beings have shown limited evidence for parents appearing to differentiate their investment to sons or daughters according to the reproductive potential of each sex. The present study tests the T-W hypothesis among a large contemporary Polish sample using first birth interval and extent of breastfeeding as measures of parental investment, and economic status and level of parental education as measures of parental condition. The extents to which parental investment and markers of parental condition vary by sex of the child were examined using log-linear analysis. Weak support for the T-W effect is found among families where fathers were best educated, where a greater proportion of first-born boys are breastfed longer than girls, while the opposite trend is observed among families with fathers with lowest levels of education. Although the present study does not fully support the T-W hypothesis, it gives evidence of greater investment in female offspring at the lower extremes of income, and greater investment in males at higher levels of income.     

Trivers-Willard at birth and one year: evidence from US natality data 1983-2001

Trivers & Willard (TW) hypothesized that evolution would favour deviations from the population sex ratio in response to parental condition: parents in good condition would have more sons and parents in poor condition would have more daughters. We analyse the universe of US linked births and infant deaths to white mothers 1983-2001, covering 48 million births and 310,000 deaths. We find that (i) married, better educated and younger mothers bore more sons and (ii) infant deaths were more male if the mother was unmarried and young. Our findings highlight the potential role of offspring sex ratio as an indicator of maternal status, and the role of infant mortality in shaping a TW pattern in the breeding population.     

Sex ratio manipulation within broods of house wrens, Troglodytes aedon

Trivers & Willard (1973, Science, 179, 90-92) developed an economic theory of parental investment to explain how the relative profitability of sons and daughters varies under specific ecological conditions. In their maternal condition hypothesis they proposed that in polygynous species, the sex of an offspring should be associated with the amount of parental care likely to be made available to it. In these species, the amount of parental investment directed towards offspring may differentially influence the fitness of male and female offspring because males in better than average condition as adults may enjoy larger fitness gains than a female would if she were in better than average condition, while the reverse may be true when conditions are poor. I tested this hypothesis by determining the sex of specific offspring within house wren broods. Because hatching is asynchronous and fledging is synchronous in this polygynous species, last-hatched young fledge having received less parental care than their broodmates. I predicted that last-hatched offspring would be more likely to be female. I found that these young were indeed more likely to be females, were more likely to have hatched from last-laid eggs and were fledging in poor condition relative to their broodmates. I propose that female house wrens behave in a manner consistent with the predictions of the Trivers & Willard hypothesis by producing female offspring last in the laying sequence of their clutches. Copyright 2000 The Association for the Study of Animal Behaviour.     

Sexual selection favors female-biased sex ratios: the balance between the opposing forces of sex-ratio selection and sexual selection

In a verbal model, Trivers and Willard proposed that, whenever there is sexual selection among males, natural selection should favor mothers that produce sons when in good condition but daughters when in poor condition. The predictions of this model have been the subject of recent debate. We present an explicit population genetic model for the evolution of a maternal-effect gene that biases offspring sex ratio. We show that, like local mate competition, sexual selection favors female-biased sex ratios whenever maternal condition affects the reproductive competitive ability of sons. However, Fisherian sex-ratio selection, which favors a balanced sex ratio, is an opposing force. We show that the evolution of maternal sex-ratio biasing by these opposing selection forces requires a positive covariance across environments between the sex-ratio bias toward sons (b) and the mating success of sons (r). This covariance alone is not a sufficient condition for the evolution of maternal sex-ratio biasing; it must be sufficiently positive to outweigh the opposing sex-ratio selection. To identify the necessary and sufficient conditions, we partition total evolutionary change into three components: (1) maternal sex-ratio bias, (2) sexual selection on sons, and (3) sex-ratio selection. Because the magnitude of the first component asymmetrically affects the strength of the second, biasing broods toward females in a poor environment evolves faster than the same degree of bias toward males in a good environment. Consequently, female-biased sex ratios, rather than male-biased sex ratios, are more likely to evolve. We discuss our findings in the context of the primary sex-ratio biases observed in strongly sexually selected species and indicate how this perspective can assist the experimental study of sex ratio evolution.     

Life-history analysis of the Trivers and Willard sex-ratio problem

Phenotypic quality, such as condition or size, often variesbetween individuals. For species with extensive maternal care,the quality of offspring may partially be determined by thequality of their mother. Trivers and Willard (1973) predictedthat high quality females should prefer offspring of the sexwhose reproductive success is most strongly influenced by maternalcare, which in many cases will be sons. Correspondingly, lowquality females should prefer daughters. However, this predictionis not based on a proper analysis of variation in reproductivevalue. Using state-dependent life-history theory, I show herethat high quality females should prefer offspring of the sexwhose reproductive value is most strongly influenced by maternalcare. I also show that when offspring quality is strongly determinedby their mother's quality, but not influenced by their father'squality, high quality females can have higher reproductive valuethan high quality males, even though their reproductive successmay be much lower. In such cases, high quality females shouldprefer daughters and, correspondingly, low quality females shouldprefer sons.[Behav Ecol 7: 316–325 (1996)]     

The effects of the minimum threshold condition for breeding on offspring sex-ratio adjustment in the lesser kestrel

We propose a model for sex-ratio adjustment complementary to that of Trivers and Willard. In addition to the three basic assumptions of the Trivers-Willard model, our model assumes that the sex with more variable reproductive success (normally male) is also the sex less constrained for reproduction. This assumption seems realistic, because several studies have demonstrated that poor-condition males may adopt alternative mating strategies and sire some offspring, whereas females have physiological constraints for gestation or egg production that cannot be avoided. Thus, under these circumstances, sons of both poor and good condition would be more valuable for parents than daughters, whereas daughters would be relatively more valuable than sons at intermediate condition. This model predicts, therefore, a U-shaped relationship between parental condition and offspring sex ratio. We present a case study for the monogamous lesser kestrel (Falco naumanni) that fulfills the assumptions and predictions of the model. The minimum body condition for breeding, measured as pectoral thickness, was lower for sons than for daughters. Below this minimum, males had a higher chance of breeding than females. Above this minimum, however, the lifetime reproductive success was condition dependent in males but not in females. Thus, males in better body condition attain, on average, higher reproductive success than females. Offspring sex ratio varied with the size of the father's ornaments and mother condition according to the U-shaped pattern predicted by the model.     

Maternal investment in relation to sex ratio and offspring number in a small mammal – a case for Trivers and Willard theory?

1.  Optimal parental sex allocation depends on the balance between the costs of investing into sons vs. daughters and the benefits calculated as fitness returns. The outcome of this equation varies with the life history of the species, as well as the state of the individual and the quality of the environment.
2.  We studied maternal allocation and subsequent fecundity costs of bank voles, Myodes glareolus , by manipulating both the postnatal sex ratio (all-male/all-female litters) and the quality of rearing environment (through manipulation of litter size by −2/+2 pups) of their offspring in a laboratory setting.
3.  We found that mothers clearly biased their allocation to female rather than male offspring regardless of their own body condition. Male pups had a significantly lower growth rate than female pups, so that at weaning, males from enlarged litters were the smallest. Mothers produced more milk for female litters and also defended them more intensively than male offspring.
4.  The results agree with the predictions based on the bank vole life history: there will be selection for greater investment in daughters rather than sons, as a larger size seems to be more influencial for female reproductive success in this species. Our finding could be a general rule in highly polygynous, but weakly dimorphic small mammals where females are territorial.
5.  The results disagree with the narrow sense Trivers & Willard hypothesis, which states that in polygynous mammals that show higher variation in male than in female reproductive success, high-quality mothers are expected to invest more in sons than in daughters.     

Trivers-willard rules for sex allocation

We present a quantitative model of sex allocation to investigate whether the simple “rules of thumb” suggested by Trivers and Willard (1973) would really maximize numbers of grandchildren in human populations. Using demographic data from the !Kung of southern Africa and the basic assumptions of the Trivers-Willard hypothesis, we calculate expected numbers of grandchildren based on age- and sex-specific reproductive value. Patterns of parental investment that would maximize numbers of expected grandchildren often differ from the Trivers-Willard rules. In particular, the optimum parental behavior is sensitive to population dynamics, type of parental investment, and, most important, relative ages of sons and daughters. It is doubtful whether a parent blindly following the simple Trivers-Willard rules would maximize numbers of expected grandchildren, on average. In addition, we show that sex-specific infanticide will almost never achieve the goal of maximizing expected numbers of grandchildren.     

Human Prenatal Investment Affected by Maternal Age and Parity

This paper investigates the effects of mother's age and parity on the prenatal allocation of resources in sons and daughters using a sample of 6994 in 20- to 34-year-old mothers from Granada (Spain). This study tests the Trivers–Willard allocation model, which predicts that, depending on the mother's condition, there can be a bias from 50/50 expected investment in the sexes. The results show that mothers of 25–29 age class produce relatively heavier sons compared to younger mothers, suggesting a bias in the allocation of resources before birth favoring neonate boys. This extra investment in boys appears to be related more to the mother's age than to her parity. Offspring of multiparous mothers had shorter gestation length, but multiparous mothers of the 25–29 age class had boys with a longer gestation than the younger ones. The 25–29 age class mothers are in the peak of fertility and they are presumably in better condition than the other mothers in the study sample. The results suggest that Granada women performed a sex-biased prenatal investment favoring sons by prolonging the gestation period. An index of boys' extra birth weight is proposed as a suitable variable to measure the maternal prenatal investment.     

Sex Allocation in a Colobine Monkey

In polygynous, sexual dimorphic species with higher variance in male reproductive success compared with females, females are expected to invest more heavily in sons than daughters within the constraints imposed by their physical condition (Science 1973; 179:90). Mothers in good condition, usually those of high rank, should produce more sons than females in poor condition or of low rank. We investigated sex allocation and sex‐biased maternal investment in a population of wild Hanuman langurs using rank and group size as approximations of female physical condition. Our results show that reproductive costs of sons were higher with both significantly longer interbirth intervals following male births and longer lactational periods for sons. Not in all groups did analyses of rank‐dependent sex allocation reveal the expected pattern of high‐ranking mothers producing more sons. However, sex ratio was significantly influenced by group size, with females from larger groups, i.e., in worse physical condition, producing a daughter‐biased sex ratio. In fact, only females of population‐wide superior physical condition can be expected to produce sons, because in Hanuman langurs males disperse and compete population‐wide. Thus, our results support the Trivers–Willard model and may explain the mixed evidence accruing from studies of single groups. We present a graphical model of how group size and dominance‐related differences in energy gain may influence sex allocation under different competitive regimes relative to overall resource availability. Tests of adaptive sex allocation models should consider whether reproductive competition of the preferred sex takes place primarily within a group or within the population.     

The Primary Parental Investment in Children in the Contemporary USA is Education

This paper tests the Trivers-Willard hypothesis that high-status individuals will invest more in sons and low-status individuals will invest more in daughters using data from the 2000 to 2010 General Social Survey and the 1979 National Longitudinal Survey of Youth. We argue that the primary investment U.S. parents make in their children is in their children’s education, and this investment is facilitated by a diverse market of educational choices at every educational level. We examine two measures of this investment: children’s years of education and the highest degree attained. Results show that sons of high-status fathers receive more years of education and higher degrees than daughters, whereas daughters of low-status fathers receive more years of education and higher degrees than sons. Further analyses of possible mechanisms for these findings yield null results. We also find that males are more likely to have high-status fathers than females.     

Social rank and sex ratio of captive long-tailed macaque females (Macaca fascicularis)

Variation in birth sex ratios in primates can be accounted for by two hypotheses: the local resource competition hypothesis [Silk: American Naturalist 121:56–66, 1983] and the hypothesis of Trivers & Willard [Science 179:90–92, 1973] concerning the maternal effect on the quality of a male. We examined the effects of female dominance rank on aspects of reproduction in three well-established captive groups of long-tailed macaques (Macaca fascicularis). High-ranking females produced a higher proportion of sons than low-ranking females, and factors other than rank did not have significant effects on birth sex ratios. Interbirth intervals following daughters were longer than those following sons, but they were independent of the mother's rank. The sons of high-ranking mothers had better survival prospects than sons of low-ranking mothers in some of the groups; no such difference was found for daughters. Overall, there was no sex difference in survival up to 5 years of age. These results support the Trivers-Willard hypothesis rather than the local resource competition hypothesis. An analysis of interbirth intervals suggested that the deviation in birth sex ratio is already established at conception.     

Gender and Risk in a Matrifocal Caribbean Community: A View from Behavioral Ecology

Matrifocality is a feature of Caribbean communities in which mothers and adult daughters often form the household core. I argue that daughter-biased parental care underlies matrifocality. Parental investment (PI) theory predicts sex-biased care, but factors promoting daughter preference are not always clearly specified. If sons are more likely than daughters to experience unpredictable hazards, then parents may bias their efforts toward daughters. In this study, I examine gender differences in rural Dominica and test PI predictions. Men were more likely to be poor and develop alcoholism and less likely to migrate or attend high school than women were. Educational outcomes showed a Trivers-Willard effect: Boys from unfavorable family environments were less likely to receive secondary education than were other boys, but there was no association for girls. PI variables generally accounted for less variance in men's outcomes than women's, suggesting that unpredictable hazards for sons may promote daughter preference and matrifocality.