Were fewer boys born in the United States during the early months of the COVID-19 pandemic? A test of the Trivers-Willard hypothesis.

The Trivers-Willard hypothesis predicts that mammalian parents in poor environmental conditions will favor the offspring sex with more reliable chance of reproductive success, which in humans is females. Three months following the onset of the COVID-19 pandemic in South Africa, England, and Wales, there were significant decreases in the sex ratio at birth (SRB) (male births/total live births). We analyzed this ratio with a seasonal autoregressive moving average model, and a logistic regression, using nationwide natality data for all singleton births in the United States from 2015 to 2021 (n = 25,201,620 total births). We identified no significant change in the sex ratio in either analysis. Rather, we observed marked differences in the sex ratio by maternal characteristics of race/ethnicity, age, and education, with more vulnerable groups having lower sex ratios. These findings suggest the SRB may be an important marker of reproductive vulnerability for disadvantaged groups in the United States.

Parental background and daughters' and sons' educational outcomes - application of the Trivers-Willard hypothesis.

This study uses Trivers-Willard hypothesis to explain the differences in daughters' and sons' educational outcomes by parental background. According to the Trivers-Willard hypothesis (TWH), parental support and investments for sons and daughters display an asymmetrical relationship according to parental status because of the different reproductive advantage of the sexes. It predicts that high-status parents support sons more than daughters, and low-status parents support daughters more than sons. In modern societies, where education is the most important mediator of status, the TW hypothesis predicts that sons from high-status families will achieve higher educational outcomes than daughters. Using cohorts born between 1987 and 1997 from the reliable full population Finnish register data that contain the data of over 600.000 individuals, children's educational outcomes were measured using data on school dropout rate, academic grade point average (GPA), and general secondary enrollment in their adolescence. OLS and sibling fixed-effect regression that permitted an examination of opposite-sex siblings' educational outcomes within the same family were applied. Sons with high family income and parental education, compared to daughters of the same family, have lower probability of dropping out of school and are more likely to enroll into academic secondary school track. In families with low parental education or income daughters have lower probability for school dropout and enroll more likely to academic school track related to sons of the same family. The effect of family background by sex can be interpreted to support TWH in dropout and academic school track enrollment but not in GPA.

Kangaroo Fathers Modulate Maternal Control of Offspring Sex but Not Postpartum Maternal Expenditure.

AbstractWhen sons and daughters have different fitness costs and benefits, selection may favor deviations from an even offspring sex ratio. Most theories on sex ratio manipulation focus on maternal strategies and sex-biased maternal expenditure. Recent studies report paternal influences on both offspring sex ratio and postpartum sex-biased maternal expenditure. We used long-term data on marked kangaroos to investigate whether and how paternal mass and skeletal size, both determinants of male reproductive success, influenced (a) offspring sex in interaction with maternal mass and (b) postpartum sex-biased maternal expenditure. When mothers were light, the probability of having a son increased with paternal mass. Heavy mothers showed the opposite trend. A similar result emerged when considering paternal size instead of mass. Postpartum maternal sex-specific expenditure was independent of paternal mass or size. Studies of offspring sex manipulation or maternal expenditure would benefit from an explicit consideration of paternal traits, as paternal and maternal effects can modulate each other.

Parental income inequality and children's digit ratio (2D:4D): a 'Trivers-Willard' effect on prenatal androgenization?

Income inequality is associated positively with disease prevalence and mortality. Digit ratio (2D:4D) - a negative proxy for prenatal testosterone and a positive correlate of prenatal oestrogen - is related to several diseases. This study examined the association of income inequality (operationalized as relative parental income) and children's 2D:4D. Participants self-measured finger lengths (2D=index finger, and 4D=ring finger) in a large online survey conducted in July 2005 (the BBC Internet Study) and reported their parents' income. Children of parents of above-average income had low 2D:4D (high prenatal testosterone, low prenatal oestrogen) while the children of parents of below-average income had high 2D:4D (low prenatal testosterone, high prenatal oestrogen). The effects were significant in the total sample, present among Whites (the largest group in the sample), in the two largest national samples (UK and USA) and were greater for males than females. The findings suggest a Trivers-Willard effect, such that high-income women may prenatally masculinize their sons at the expense of the fitness of their daughters. Women with low income may prenatally feminize their daughters at the fitness expense of their sons. The effect could, in part, explain associations between low income, high 2D:4D (low prenatal testosterone) and some major causes of mortality such as cardiovascular disease.

Maternal stress and sex ratio at birth in Sweden over two and a half centuries: a retest of the Trivers-Willard hypothesis.

STUDY QUESTION: Is there a negative relationship, as predicted in the Trivers-Willard hypothesis (TWH), between the intensity of maternal stress and sex ratio at birth (SRB)? SUMMARY ANSWER: Using a comprehensive data set with multiple indicators of maternal stress, most measures of stress show no statistically significant association with SRB over a period spanning 243 years, indicating no support for the TWH. WHAT IS KNOWN ALREADY: Evolutionary biologists have proposed a widely discussed hypothesis that women in poor and stressful conditions during pregnancy are more likely to give birth to girls, and exposure to stressful events may therefore lead to a reduction in sex (male-to-female) ratio at birth. The empirical evidence so far is mixed. STUDY DESIGN, SIZE, DURATION: Annual time series data, spanning 243 years between 1749 and 1991 for Sweden at the national level, were drawn from multiple sources. The outcome is defined as the percentage of male births relative to all births in Sweden in a given year. The covariates include a set of economic and climatic variables as proxies for maternal stress. PARTICIPANTS/MATERIALS, SETTING, METHODS: We conduct a series of ARIMA (autoregressive integrated moving average) models to examine the relationship between maternal stress and SRB during three periods: 1749-1991, 1749-1861 and 1862-1991. MAIN RESULTS AND THE ROLE OF CHANCE: In 1749-1991, economic proxies for maternal stress showed no statistically significant association with SRB. In 1749-1861, two indicators were significantly associated with SRB, but the coefficients were opposite in direction to the TWH. In 1862-1991, five out of six covariates showed no significant association with SRB. An additional analysis found no significant correlation between sex ratio of stillbirths and all covariates in 1862-1991. Our results are incompatible with the TWH and suggest that previous findings in support of the TWH are not robust. LIMITATIONS, REASONS FOR CAUTION: This study provides population-level evidence that may not necessarily reflect the nature of all individuals due to the ecological fallacy. The time series analysed in this study are annual data, and we cannot examine the potential seasonality due to the lack of disaggregated monthly data. Our findings may not be generalised to the contexts of extreme maternal stress conditions such as famine and war. WIDER IMPLICATIONS OF THE FINDINGS: The results from existing studies in this topic may be speculative, and additional research with more comprehensive design, data and covariates is needed to reconsider the robustness of previous findings. STUDY FUNDING/COMPETING INTEREST(S): The author receives no external funding and has no conflict of interest to declare. TRIAL REGISTRATION NUMBER: N/A.

Richardson's ground squirrel litter size-sex ratio trade-off reveals conditional adaptive sex allocation.

Trivers and Willard proposed that female mammals should adjust their investment in male versus female offspring relative to their ability to produce high-quality offspring. We tested whether litter size-sex ratio trade-offs predicted by Adaptive Sex Allocation (ASA) theory occur among Richardson's ground squirrel (Urocitellus richardsonii) dams over 10 distinct breeding years in a population where individuals experienced variability in food availability and habitat disruption. Litters of primiparous dams became increasingly female-biased with increasing litter size, but that trend waned among second litters born to dams, and reversed among third litters, with larger litters becoming more male-biased, suggesting that ASA is a product of interacting selection pressures. Trade-offs were not associated with habitat disruption, the availability of supplementary food, or dam age. An association between habitat disruption and male-biased sex ratios, the prevalence of litter size-sex ratio trade-offs and placental scar counts exceeding the number of juveniles at weaning in our population, but not in a geographically distinct population of conspecifics exposed to different environmental conditions reveal that the expression of ASA varies among populations and among years within populations, illustrating the conditional nature of ASA.

Biological, environmental and socioeconomic determinants of the human birth sex ratio in the Czech Republic.

The Trivers-Willard Hypothesis (TWH) states that parents in good conditions bias the sex ratio towards sons and parents in poor conditions bias the sex ratio towards daughters. This study used data from a large nationwide population dataset (N=1,401,851) from the Czech Republic - a modern contemporary society. The study included air pollution and property prices in the TWH estimation, and had a more detailed focus on stillbirths than previous studies. Using official natality microdata from the Czech Statistical Office for years between 1992 and 2010 and data on levels of air pollution in the country over the same period, the study assessed whether the biological and socioeconomic status of mothers and environmental factors affected the sex of children. The results were largely insignificant and not robust across specifications. The presented epidemiological evidence suggests that stillbirths are randomly distributed in the Czech Republic and that the sex ratio is not affected by the socioeconomic status of mothers or by environmental characteristics.

Is there a Trivers-Willard effect for parental investment? Modelling evolutionarily stable strategies using a matrix population model with nonlinear mating.

The Trivers-Willard hypothesis (TWH) states that parents in good condition preferentially produce the sex with a higher variation in reproductive success, whereas parents in bad condition favour the opposite sex. Theorists distinguish two variants of the TWH: (a) a biased sex-ratio at birth and (b) biased parental investment after birth. It has been argued before that the conditions stated by Trivers and Willard (good condition is inherited and affects reproductive success more strongly for one of the sexes) are sufficient for the sex-ratio version but insufficient for the investment version of the TWH. However, it has not yet been investigated how these conditions affect parental investment in high and low quality parents, depending on the life-cycle of a species. The present study aims to fill this gap by introducing a multi-stage matrix population model with nonlinear mating to describe the effects of parental investment after birth on the reproductive values of male and female individuals. Using methods from adaptive dynamics and evolutionary invasion analysis, evolutionary trajectories and evolutionarily stable strategies are derived for different parameterizations of the model. Simulation results demonstrate that the conditions given by Trivers and Willard produce a general bias of parental investment towards the sex with higher variance in reproductive value. This bias is stronger for low-quality parents than for high-quality parents and matches the expected marginal offspring reproductive values for parental investment.

Maternal condition and previous reproduction interact to affect offspring sex in a wild mammal.

Trivers and Willard proposed that offspring sex ratio should vary with maternal condition when condition, meant as maternal capacity to care, has different fitness consequences for sons and daughters. In polygynous and dimorphic species, mothers in good condition should preferentially produce sons, whereas mothers in poor condition should produce more daughters. Despite its logical appeal, support for this hypothesis has been inconsistent. Sex-ratio variation may be influenced by additional factors, such as environmental conditions and previous reproduction, which are often ignored in empirical studies. We analysed 39 years of data on bighorn sheep (Ovis canadensis) that fit all the assumptions of the Trivers-Willard hypothesis. Production of sons increased with maternal condition only for mothers that weaned a son the previous year. This relationship likely reflects a mother's ability to bear the higher reproductive costs of sons. The interaction between maternal condition and previous weaning success on the probability of producing a son was independent of the positive effect of paternal reproductive success. Maternal and paternal effects accounted for similar proportions of the variance in offspring sex. Maternal reproductive history should be considered in addition to current condition in studies of sex allocation.

The evolutionary ecology of early weaning in Kilimanjaro, Tanzania.

Public health recommendations promote prolonged breastfeeding of all children; however, parental investment (PI) theory predicts that breastfeeding will be allocated among a mothers' offspring to maximize her reproductive success. We evaluated PI in terms of risk for weaning before age two among 283 children in Kilimanjaro, Tanzania. Results demonstrate: (i) a Trivers-Willard effect--high socioeconomic status (SES) females and low SES males were more likely to be weaned early; (ii) later-born children were less likely to be weaned early; (iii) higher birthweight children were less likely to be weaned early, and (iv) no effect of cattle (a source of supplementary milk) ownership. These associations were largely independent and remained significant in models controlling for potential confounders; however, the inverse association between early weaning and birth order lost significance in the model containing birthweight. These patterns were observed despite public health recommendations encouraging breastfeeding for at least two years.

Pandemic influenza and gender imbalance: Mortality selection before births.

This study uses data from the 1918-1920 influenza pandemic in Japan along with newly digitized and complete census records on births and infant deaths to analyze mortality selection in utero. I find that fetal exposure to the influenza pandemic during the first trimester of the pregnancy decreases the proportion of males at birth. The results from mechanism analysis suggest that this decline in male births is associated with the deterioration of fetal and infant health. This result supports a wide range of existing literature on the long-run adverse effects of pandemic influenza.

The effects of parental age difference on the offspring sex and fitness of European blackbirds.

BACKGROUND: Many studies of birds have indicated that offspring sex ratios can vary with environmental and parental traits. On the basis of long-term research, we first evaluated the possible influence of parental age difference and brood characteristics on offspring sex and fitness in multi-brooded Blackbirds Turdus merula. METHODOLOGY: The study was conducted in the city-centre Stefan Zeromski Park in Szczecin, NW Poland, where the local population of Blackbirds has been studied since 1996. Data on the offspring sex and fitness were collected in five years, 2005-2007 and 2016-2017. During the breeding season we inspected the study area to locate the pairs' territories and to track their nests and clutches. RESULTS: We found that the overall sex ratio did not differ statistically from 50:50, but that younger females bonded with older mates did tend to produce more sons, probably because of the greater fitness of male descendants. Accordingly, the sons' breeding success increased with the father's age, but this relationship was close to non-linear, which may indicate that the transgenerational effect of paternal senescence could negatively affect progeny fitness despite the high-quality of older fathers. Older females mated with younger males produced more daughters, which could have been due to the lesser attractiveness of the males and the mothers' poorer condition caused by accelerated senescence. We found that neither offspring hatching sequence nor hatching date or clutch sequence were significant for sex determination. CONCLUSIONS: We consider that in our Blackbird population, parental age could make a more significant contribution to shaping offspring sex and reproductive success.

Parental investment in the Columbian ground squirrel: empirical tests of sex allocation models.

Parental allocation of resources into male or female offspring and differences in the balance of offspring sexes in natural populations are central research topics in evolutionary ecology. Fisher (Fisher, R. A. 1930. The genetical theory of natural selection, Clarendon Press, Oxford, UK) identified frequency-dependent selection as the mechanism responsible for an equal investment in the sexes of offspring at the end of parental care. Three main theories have been proposed for explaining departures from Fisherian sex ratios in light of variation in environmental (social) and individual (maternal condition) characteristics. The Trivers-Willard model (Trivers, R., and D. Willard. 1973. Natural selection of parental ability to vary the sex ratio of offspring. Science 179:90-92) of male-biased sex allocation by mothers in the best body condition is based on the competitive ability of male offspring for future access to mates and thus superior reproduction. The local resource competition model is based on competitive interactions in matrilines, as occur in many mammal species, where producing sons reduces future intrasexual competition with daughters. A final model invokes advantages of maintaining matrilines for philopatric females, despite any increased competition among females. We used 29 yr of pedigree and demographic data to evaluate these hypotheses in the Colombian ground squirrel (Urocitellus columbianus), a semisocial species characterized by strong female philopatry. Overall, male offspring were heavier than female offspring at birth and at weaning, suggesting a higher production cost. With more local kin present, mothers in the best condition biased their offspring sex ratio in favor of males, and mothers in poor condition biased offspring sex ratio in favor of females. Without co-breeding close kin, the pattern was reversed, with mothers in the best condition producing more daughters, and mothers in poor condition producing more sons. Our results do not provide strong support for any of the single-factor models of allocation to the sexes of offspring, but rather suggest combined influences of relative maternal condition and matriline dominance on offspring sex ratio.

Bias of Calf Sex on Milk Yield and Fat Yield in Holstein Crossbreed Cows.

In order to examine the biased milk production depending on the sex of calves, data on calving and milk yield characteristics of 15,181 Holstein type cows in PK Belgrade, Serbia were analyzed. A total of 30,362 lactations that were realized in the period from 1985 to 2017 were analyzed. Data were prepared and analyzed using the SAS software package (SAS Institute Inc. Software License 9.3, 2012). The expression and variability of investigated traits were determined using the PROC MEANS procedure, while the effect of individual factors on milk yield traits was analyzed using the PROC GLM procedure. Obtained results deviate from the views of the Trivers-Willard (TW) hypothesis. The results indicate that mothers invest more in female offspring by producing a higher milk and fat yield in the first and second lactation compared to male offspring. This is especially emphasized under better environmental conditions. The highest milk yield (7788 kg) and fat yield (271 kg) in the second lactation were achieved in the combination with two consecutive female calves in the group of higher-than-average milk production farms, and lowest in the combination of two consecutive male calves (6783 kg for the MY and 243 kg for the FY), respectively.

Evolutionary dynamics of the Trivers-Willard effect: A nonparametric approach.

The Trivers-Willard hypothesis (TWH) states that parents in good condition tend to bias their offspring sex ratio toward the sex with a higher variation in reproductive value, whereas parents in bad condition favor the opposite sex. Although the TWH has been generalized to predict various Trivers-Willard effects (TWE) depending on the life cycle of a species, existing work does not sufficiently acknowledge that sex-specific reproductive values depend on the relative abundances of males and females in the population. If parents adjust their offspring sex ratio according to the TWE, offspring reproductive values will also change. This should affect the long-term evolutionary dynamics and might lead to considerable deviations from the original predictions. In this paper, I model the full evolutionary dynamics of the TWE, using a published two-sex integral projection model for the Columbian ground squirrel (Urocitellus columbianus). Offspring sex ratio is treated as a nonparametric continuous function of maternal condition. Evolutionary change is treated as the successive invasion of mutant strategies. The simulation is performed with varying starting conditions until an evolutionarily stable strategy (ESS) is reached. The results show that the magnitude of the evolving TWE can be far greater than previously predicted. Furthermore, evolutionary dynamics show considerable nonlinearities before settling at an ESS. The nonlinear effects depend on the starting conditions and indicate that evolutionary change is fastest when starting at an extremely biased sex ratio and that evolutionary change is weaker for parents of high condition. The results show neither a tendency to maximize average population fitness nor to minimize the deviation between offspring sex ratio and offspring reproductive value ratio. The study highlights the importance of dynamic feedback in models of natural selection and provides a new methodological framework for analyzing the evolution of continuous strategies in structured populations.

The effects of toxoplasmosis on sex ratio at birth.

Toxoplasmosis affects about one third of human population worldwide. It has a wide range of effects on the health, immunity, behaviour, and both prenatal and postnatal outcomes of infected hosts, including humans. Among these effects, stage of infection-specific shifts in secondary sex ratio were described about ten years ago both in humans and in artificially infected mice. In both women and female mice, in the early stage of infection the probability of giving birth of sons significantly increases, up to 260 sons to every 100 daughters. In the late stages of infection, the probability of giving birth to sons markedly decreases to as low as 78 to every 100 daughters. An ecological correlation study shows that the effect of latent toxoplasmosis on human population biology and demography can be large. In fact, the effect of prevalence of toxoplasmosis on a nationwide sex ratio was the third strongest effect from the effects of 15 factors included in the analysis. It has been suggested that toxoplasmosis-associated concentration of steroid hormones or glucose may be the proximal cause in the sex ratio shift. A more parsimonious explanation of the upward secondary sex ratio shift is found in a lower stringency of quality control of embryos, whose side-effect is increased survival rate of the more immunogenic male embryos in immunosuppressed infected females. The most parsimonious explanation of the downward secondary sex ratio shift relies on the Trivers-Willard hypothesis, which predicts an adaptive shift to more daughters in females with impaired health or lower socioeconomic status.

Maternal RhD heterozygous genotype is associated with male biased secondary sex ratio.

BACKGROUND: Previous studies suggest that RhD positive heterozygotes express better health status than RhD positive homozygotes and especially RhD negative subjects. This also applies to pregnant women. According to the Trivers-Willard hypothesis, women in better physical condition should have more sons. AIM: To test the hypothesis that RhD positive heterozygous mothers have a male-skewed sex ratio. STUDY DESIGN: Cross-sectional study. The data was analysed using Chi-Square test for all women, separately for RhD positive and RhD negative women, and separately for primiparous and multiparous women. The effects of maternal weight as a continuous predictor and the RhD phenotype of newborn as a categorical predictor of newborn sex were evaluated by the generalized linear model (GLZ) separately for RhD positive and RhD negative women using binomial distribution and logit link function. OUTCOME MEASURES: Clinical records comprised maternal weight before pregnancy, number of previous deliveries, sex of the newborn, maternal RhD phenotype, and RhD phenotype of the newborn. SUBJECTS: We analysed data from 5655 women who gave birth between 2008 and 2012 in General University Hospital in Prague. RESULTS: Secondary sex ratio was significantly higher (P = 0.028) in RhD positive mothers who had RhD negative newborns, i.e., in heterozygotes (SR = 1.23), than in RhD positive mothers who had RhD positive newborns, i.e., in a mixed population of heterozygotes and homozygotes (SR = 1.00), especially in primiparous women (P = 0.013; SR = 1.37 and 0.99 resp.). CONCLUSION: The sex ratio at birth was significantly higher in RhD positive mothers who had RhD negative newborns than in RhD positive mothers who had RhD positive newborns.

Different maternal investment strategies for male and female calves in a polygynous mammal.

Maternal effects occur when the phenotype of the mother influences that of the young to the detriment of her survival, growth or fitness. The investment of the mother can be affected by maternal body condition and/or experience. Trivers-Willard Hypothesis (TWH) and Local Resource Competition Hypothesis (LRCH) are the main hypotheses used to explain bias in birth sex-ratios in mammals, as well as for sex-biased maternal investment. Both hypotheses suggest that a different amount of investment must be expected according to the sex of the young. However, recent studies suggest that these differences are not in quantity but in the strategies: mechanisms and objectives may differ for each sex. We studied how maternal characteristics (age, body mass, body condition, and dominance status) influence relevant aspects of the birth and early growth of the calf (birth date, birth body mass, body mass at weaning, and body condition at weaning) separately for each sex; and how that investment is mediated by milk production and composition (lactose, fat, and protein). One hundred eighty-eight newborns from 75 captive red deer hinds aged from 2 to 19 years were analyzed. The main differential investment observed was related to birth date: when producing a female, hinds give birth earlier in the season only if they have a good body condition; however, when gestating a male it is the older hinds those which deliver earlier. Subsequently, milk production and composition are correlated with birth body mass in female calves, but to weaning body mass in males. Thus, only hind body mass affects the weaning body mass of female calves, compared with age and hind body mass in males. These results suggest that while TWH fits the maternal investment strategy found for male calves, it is LRCH which correlates with the maternal investment patterns observed for females.

Parental human capital and child health at birth in India.

Health at birth shapes an individual's well-being over her life cycle. We categorize the Indian states into high and low infant mortality regions to capture the diverse disease environment and analyze the nature of the association between parental human capital and child survival and nutrition measures at birth. We restrict our analysis only to firstborns to avoid confounding from a number of factors including sex-selective abortions in the higher birth orders. We broadly find that parental human capital, especially maternal health, is a strong and significant predictor of a child's birth outcomes under adverse disease environment. In the rural areas of the high infant mortality states, a 10-centimeter increase in maternal height is associated with 1.7% lower probability of a child dying as a neonate and 5% increase in birth weight around the mean. These estimates suggest that an investment in human capital of the mothers from this region could accompany large gains in survival and nutrition outcomes of their children.

Sex Differences in Intergenerational Income Transmission and Educational Attainment: Testing the Trivers-Willard Hypothesis.

From an evolutionary point of view, sex differences in intergenerational transmission of income may be influenced by the Trivers-Willard (T-W) effect: Low status parents should invest more in daughters, whereas high status parents are expected to invest more in sons. This bias in parental investment may result in status-dependent sex biased parental support for higher education and educational attainment and should therefore affect the level of intergenerational income transmission for the sons and daughters. We used the data from the Wisconsin Longitudinal Study (WLS) to model the effect of parental financial investment on the child's income and educational attainment controlling for the number of siblings. The observed sex differences in intergenerational income transmission demonstrate that sons profited more from parental income and education in terms of their own income than daughters. Furthermore, we showed that fathers with a high socioeconomic index (SEI) invest more in their sons' education in terms of completed years of education and financial support during college. In contrast daughters of low SEI fathers completed more years of education and received more financial support than sons of low SEI fathers. However, the pattern in intergenerational income transmission might be better explained as a product of sociological factors and reproductive trade-offs in later life rather than as a consequence of the T-W effect.