Violent men have more sons: further evidence for the generalized Trivers-Willard hypothesis (gTWH)

The generalized Trivers-Willard hypothesis (gTWH) [Kanazawa, S., 2005a. Big and tall parents have more sons; further generalizations of the Trivers-Willard hypothesis. J. Theor. Biol. 235, 583-590] proposes that parents who possess any heritable trait which increases the male reproductive success at a greater rate than female reproductive success in a given environment have a higher-than-expected offspring sex ratio, and parents who possess any heritable trait which increases the female reproductive success at a greater rate than male reproductive success in a given environment have a lower-than-expected offspring sex ratio. One heritable trait which increases the reproductive success of sons significantly more than that of daughters in the ancestral environment is the tendency toward violence and aggression. I therefore predict that violent parents have a higher-than-expected offspring sex ratio (more sons). The analysis of both American samples and a British sample demonstrates that battered women, who are mated to violent men, have significantly more sons than daughters.     

Big and tall parents have more sons: further generalizations of the Trivers-Willard hypothesis

This paper proposes the generalized Trivers-Willard hypothesis (gTWH), which suggests that parents who possess any heritable trait which increases male reproductive success at a greater rate than female reproductive success in a given environment will have a higher-than-expected offspring sex ratio, and parents who possess any heritable trait which increases female reproductive success at a greater rate than male reproductive success in a given environment will have a lower-than-expected offspring sex ratio. Since body size (height and weight) is a highly heritable trait which increases male (but not female) reproductive success, the paper hypothesizes that bigger and taller parents have more sons. The analysis of both surviving children and recent pregnancies among respondents of the National Child Development Survey and the British Cohort Survey largely supports the hypothesis.     

Offspring sex ratio produced by female guppies in the wild correlates with sexual ornaments of their sons

The attractiveness hypothesis predicts that females produce broods with male-biased sex ratios when they mate with attractive males. This hypothesis presumes that sons in broods with male-biased sex ratios sired by attractive males have high reproductive success, whereas the reproductive success of daughters is relatively constant, regardless of the attractiveness of their sires. However, there is little direct evidence for this assumption. We have examined the relationships between offspring sex ratios and (1) sexual ornamentation of sons and (2) body size of daughters in broods from wild female guppies Poecilia reticulata. Wild pregnant females were collected and allowed to give birth in the laboratory. Body size and sexual ornamentation of offspring were measured at maturity. Our analysis revealed a significant positive correlation between offspring sex ratios (the proportion of sons per brood) and the total length as well as the area of orange spots of sons, two attributes that influence female mating preferences in guppies. The sex ratio was not associated with the body size of daughters. These results suggest that by performing adaptive sex allocation according to the expected reproductive success of sons and daughters, female guppies can enhance the overall fitness of their offspring.     

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.     

Engineers have more sons, nurses have more daughters: an evolutionary psychological extension of Baron-Cohen's extreme male brain theory of autism

In his extreme male brain theory of autism, Baron-Cohen postulates that having a typically male brain was adaptive for ancestral men and having a typically female brain was adaptive for ancestral women. He also suggests that brain types are substantially heritable. These postulates, combined with the insight from the Trivers-Willard hypothesis regarding parental ability to vary offspring sex ratio, lead to the prediction that people who have strong male brains should have more sons than daughters, and people who have strong female brains should have more daughters than sons. The analysis of the 1994 US General Social Survey data provides support for this prediction. Our results suggest potentially fruitful extensions of both Baron-Cohen's theory and the Trivers-Willard hypothesis.     

Low Socioeconomic Status and Female-Biased Parental Investment: The Mukogodo Example

Hierarchies of wealth and ethnic prestige among East African herders present an opportunity to test the Trivers-Willard hypothesis that low socioeconomic status should correlate with female biases in parental investment. The Mukogodo are at the bottom of such a regional hierarchy due to their poverty and low status as former hunters. As a result of these factors, Mukogodo men have lower polygyny rates than their neighbors, and Mukogodo women have higher mean reproductive success than Mukogodo men. The data fulfill the prediction that there should be a bias in parental investment in favor of daughters. The sex ratio of the 0–4 age group and the reported sex ratio at birth are both female-biased. Although there is no evidence of infanticide, sons may be neglected in favor of daughters. Evidence from a dispensary and from a clinic run by a Catholic mission both show that the Mukogodo take daughters for treatment more often than they take sons. Also, daughters may be nursed longer than sons.     

Birth sex ratios relate to mare condition at conception in Kaimanawa horses

Several hypotheses have been proposed to explain variation inbirth sex ratios, based on the premise that variation is expectedwhen the profitability of raising sons and daughters variesbetween individual parents. We tested the Trivers-Willard hypothesisthat mothers in better condition produce relatively more sonsand that mothers in poorer condition produce relatively more daughterswhen male reproductive success is more variable. We examinedbirth sex ratios in relation to mare body condition at conceptionin horses in which male reproductive success is differentiallyhelped by slight advantages in condition. Horses meet the assumptionsof the Trivers-Willard hypothesis better than many species onwhich it has been tested and in which sex ratio biases are notconfounded by sexual size dimorphism such that one sex is more likelyto die in utero in females in poor condition. Mares that hada female foal were in poorer condition at conception than thosethat had a male foal, and mares that had foals of differentsexes in different years were in significantly poorer conditionwhen they conceived their female foal. There was no relationshipbetween offspring sex and mid-gestation condition, and therewas no difference in foaling rates in relation to body conditionat conception. Consequently, sex ratio deviations are not explainedby fetal loss in utero. Furthermore, differential fetal lossof the less viable sex cannot explain the greater proportionof males produced by mares in better condition. Therefore, ourresults suggest that sex ratio modification occurs at conceptionin wild horses.     

Brothers and sisters

Data from the Kipsigis of Kenya are used to test two models for how parents invest in offspring, the Trivers-Willard and local resource competition/enhancement hypotheses. Investment is measured as age-specific survival, educational success, marital arrangements, and some components of property inheritance, permitting an evaluation of how biases persist or alter over the period of dependence. Changes through time in such biases are also examined. Despite stronger effects of wealth on the reproductive success of men than women, the survival of sons and daughters is not related to parental wealth. However, a Trivers-Willard effect characterizes educational investment: poor families show a greater concern for daughters’ (vis-à-vis sons’) schooling than do rich families, a trend that has increased over time. In regard to models of local resource competition and enhancement, men’s reproductive success decreases with number of brothers and increases with number of sisters; this pattern of competition with same-sex sibs and cooperation with opposite-sex sibs is not found among women. As predicted from these observations, parents show reduced investment in sons with a large number of brothers, and increased investment in sons with a large number of sisters. By contrast, investment in daughters is entirely unaffected by number of sisters and is influenced only in subtle ways by number of brothers. Levels of investment in relation to sibship size (irrespective of siblings’ sex) are highest for younger children of large sib sets. Discussion of the results in relation to those from other studies leads to three conclusions. First, predictive models for how investment biases vary across societies must consider a broad range of socioecological factors constraining parental options and payoffs. Second, the timing of investment biases within societies will be affected by the value of children and the costs of parental investment. Third, measures of investment appropriate for between-sex and between-class comparisons need careful attention. Each of these issues is brought to bear on the question of why, in contrast to so many other parts of the world, sex preferences are so muted in Africa.     

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.     

No evidence for adjustment of sex allocation in relation to paternal ornamentation and paternity in barn swallows

Sex allocation theory predicts that parents should adjust investment in sons and daughters according to relative fitness of differently sexed offspring. In species with female preference for highly ornamented males, one advantage potentially accruing to parents from investing more in sons of the most ornamented males is that male offspring will inherit characters ensuring sexual attractiveness or high-quality genes, if ornaments honestly reveal male genetic quality. Furthermore, in species where extra-pair fertilizations occur, offspring sired by an extra-pair male are expected to more frequently be male than those of the legitimate male if the latter is of lower quality than the extra-pair male. We investigated adjustment of sex ratio of offspring in relation to ornamentation of the extra-pair and the social mate of females by direct manipulation of tails of male barn swallows Hirundo rustica . Molecular sexing of the offspring was performed using the W chromosome-linked avian chromo-helicase-DNA-binding protein (CHD) gene while paternity assessment was conducted by typing of hypervariable microsatellite loci. Extra-pair offspring sex ratio was not affected by ornamentation of their biological fathers relative to the experimental ornamentation of the parental male. Experimental ornamentation of the parental males did not affect the sex ratio of nestlings in their broods. Female barn swallows might be unable to bias offspring sex ratio at hatching according to the quality of the biological father. Alternatively, fitness benefits in terms of sexual attractiveness of sons might be balanced by the cost of compensating for little parental care provided by highly ornamented parental males, if sons are more costly to rear than daughters, or the advantage of producing more daughters, if males with large ornaments contribute differentially more to the viability of daughters than sons.     

SEX-RATIO MANIPULATION IN COLOR-BANDED POPULATIONS OF ZEBRA FINCHES

Significant correlations were found between attractiveness of leg-band color (determined by preference tests [Burley et al., 1982]) and sex ratio of offspring in two long-term breeding experiments involving zebra finches. In both experiments, birds with attractive band colors produced more same-sex offspring, while birds with unattractive band colors produced more opposite-sex offspring. The results of these experiments are consistent with those of a previous experiment (Burley, 1981). To explain the earlier results, I hypothesized that parents adjust their allocation to sons and daughters to produce offspring they “expect” to be most attractive. The purpose of such sex-ratio manipulation is to enhance fitness by the production of offspring with superior mate-getting opportunities. Two alternative hypotheses are presented here. One is that sex ratios change with parental age and/or experience. Evidence does not support this hypothesis. There were no temporal trends in sex ratio independent of band color. A second possibility is that sex ratios reflect differential parental ability to rear sons and daughters. This hypothesis cannot be conclusively tested on the basis of present evidence, but available evidence does not support it. Within color classes, weights of sons and daughters did not differ. Evidence indicates that parents effect secondary sex-ratio manipulation through the selective rejection of young, usually within six days of hatching. There is no evidence of manipulation prior to egg-laying. The costs associated with brood reduction probably set limits on the extent to which secondary manipulation can be profitably employed.     

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.     

Sex ratio bias in the dung beetle <Emphasis Type="Italic">Onthophagus taurus</Emphasis>: adaptive allocation or sex-specific offspring mortality?

Sex allocation theory predicts that females should adjust the sex of their offspring when the fitness returns of one sex are higher than the other. However, biased sex ratios may also arise if mortality differs between the sexes. Here, we examine whether offspring sex ratio bias in the dung beetle, Onthophagus taurus, represents adaptive sex allocation by females or is due to sex-specific mortality. First, we re-analyze an existing data set to show that females produce an excess of daughters when mating to smaller, less attractive males and near equal sex ratio with large, more attractive males. We show, that this results from females adjusting larval provisions after mating to males of variable attractiveness which in turn influences the likelihood that sons die during development. Second, we conduct a manipulative experiment varying the quantity and quality of larval provisions and show that the mortality of sons increased when larval provisions were reduced. Collectively, our work demonstrates that offspring mortality is contingent on the amount of resources provisioned by females and that sons have greater nutritional demands than daughters during development, leading to higher mortality. Our results therefore demonstrate the importance of considering sex-specific offspring mortality in studies of sex ratio evolution.     

Offspring are lighter at birth and smaller in adulthood when born after a brother versus a sister in humans

In mammals, including humans, it is more costly to produce sons than it is to produce daughters, with maternal survival and subsequent reproductive success diminished more by producing male over female offspring. It is therefore predicted that offspring who are produced by mothers who have previously produced sons versus daughters will be compromised by the relatively high cost their mother incurred in the previous reproductive episode. Such effects are potentially important because characters that determine offspring survival and fecundity ultimately contribute to maternal fitness. Using questionnaire-based data from a contemporary human population, I show that birthweight (irrespective of their sex) is lower in individuals born after an elder brother than in those born after an elder sister. In addition, I show that both men and women who were born after a male versus a female sibling have reduced adulthood height, a known correlate of reproductive success in both sexes. The results suggest that producing sons may have a negative effect on the fitness of subsequent offspring, which has implications for calculations of maternal fitness and for optimal sex allocation.     

Fisherian and “good genes” benefits of mate choice: how (not) to distinguish between them

“Good genes” models of mate choice are commonly tested by examining whether attractive males sire offspring with improved survival. If offspring do not survive better (or indeed survive less well), but instead inherit the attractiveness of their father, results are typically interpreted to support the Fisherian process, which allows the evolution of preferences for arbitrary traits. Here, I show that the above view is mistaken. Because of life‐history trade‐offs, an attractive male may perform less well in other components of fitness. A female obtains a “good genes” benefit whenever males show heritable variation in quality, even if high‐quality males invest so much in sexual advertisement that attractiveness has no positive correlation with any other life‐history trait than male mating success itself. Therefore, a negative correlation between attractiveness and viability does not falsify good genes, if mating with a high‐quality male results on average in superior offspring performance (mating success of sons included). The heritable “good genes” benefit can be sustained even if sexually antagonistic genes cause female offspring sired by high‐quality males to survive and reproduce less well. Neglecting the component of male mating success from measurements of fitness returns from sons and daughters will bias the advantage of mating with a high‐quality male downwards. This result may partly account for the rather weak “good genes” effects found in a recent meta‐analysis.     

Long-term fitness benefits of egg sex modification by the Seychelles warbler

Sex-ratio theory states that if the fitness costs to the parents of producing one offspring's sex relative to the other are higher, parents should discount these costs by producing fewer individuals of the more costly sex. In the co-operatively breeding Seychelles warbler (Acrocephalus sechellensis) mothers adaptively modify the sex of their single egg toward daughters, the helping sex, when living on territories with rich resources where helpers increase parental reproductive success, but toward sons, the dispersing sex, when living on territories where resources are scarce and/or no helping benefits accrue. By modifying offspring sex ratio, parents maximize their inclusive fitness benefits. Pairs in high-quality territories gained significantly more inclusive fitness benefits (through helping and reproducing offspring) from the production of daughters than from sons, and vice versa in low-quality territories (through reproducing offspring). Experimental manipulation of the offspring's sex shows that the consequences of sex allocation are adaptive for parents on high-quality territories. On high-quality territories with female production, breeding pairs raising step-daughters gained significantly higher inclusive benefits (through indirect and direct fitness gains) than by raising step-sons.     

Big mothers invest more in daughters – reversed sex allocation in a weakly polygynous mammal

How mothers allocate resources to offspring is central to understanding life history strategies. High quality mothers are predicted to favour investment in sons over daughters when to do so increases inclusive fitness. This is the case in ungulates with polygynous mating systems, where reproductive success is more variable among males than females, but information is scarce on sex allocation in less polygynous species. Here, for the weakly dimorphic roe deer, we show that as maternal capacity to invest increases, mothers increase allocation to daughters more than to sons, so that relative allocation to daughters increases markedly with increasing maternal quality. This cannot be explained by a between sex difference in growth priority, hence we conclude that this is evidence for active maternal discrimination. Further, we demonstrate that condition differences between offspring persist to adulthood. For high quality mothers of weakly polygynous species, daughters may be more valuable than sons.     

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.     

Absence of seasonal variation in great tit offspring sex ratios

When the timing of breeding affects the reproductive value of sons and daughters differently, parents are expected to increase their fitness by changing the offspring sex ratio during the course of the breeding season. Previous studies have shown that in great tits Parus major hatching date has a stronger effect on the fitness of juvenile males than on that of juvenile females. We tested whether this difference was reflected in a seasonal decline in the proportion of sons per breeding attempt. Although offspring sex ratio was more variable than would be expected from a binomial distribution, there was no significant relationship between the proportion of sons and the laying date of the clutch. Moreover, individual females did not adjust the sex ratio of their offspring following an experimental delay of breeding. This study therefore fails to demonstrate adaptive seasonal variation in great tit offspring sex ratios.     

Offspring sex ratios in tree swallows: females in better condition produce more sons

Organisms are expected to adjust the sex ratio of their offspring in relation to the relative fitness benefits of sons and daughters. We used a molecular sexing technique that amplifies an intron of the CHD1 gene in birds to examine the sex ratio at egg-laying in socially monogamous tree swallows (Tachycineta bicolor). We examined all individuals in 40 broods (210 young), including all unhatched eggs and nestlings. Thus, the sex ratio we measured was the same as the sex ratio at laying. Overall, the mean sex ratio per brood (+/- SD) was biased significantly towards males (57 +/- 2% male). Within broods, male-biased sex ratios were associated with females in better body condition, and these females were more likely to produce sons in better condition. Tree swallows have one of the highest known levels of extra-pair paternity in birds (38-76% extra-pair young), and, as a consequence, variance in male reproductive success is greater than that of females. Thus, in tree swallows, investment in sons has the potential for higher fitness returns than investment in daughters, assuming that sons in better condition have greater reproductive success.