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.     

Breast-feeding in Bangladesh

In this paper the duration of breastfeeding is examined in relation to demographic and socioeconomic characteristics of women and households. 98% of Bangladesh mothers breastfeed their children from birth. In data from the Bangladesh Fertility Survey conducted in 1975-1976 on 4998 live births, the mean duration of breastfeeding was 27.3 months. Duration of breastfeeding was positively related with the age of women. Female children were breastfed for periods about 5 months shorter than male children. Children born to urban mothers were breastfed for shorter durations than children born to rural mothers of all age groups. The duration of breastfeeding decreased with the increase of education of the mother. Promotion of breastfeeding should be a primary responsibility of family planning clinics. Further decline in the duration of breastfeeding would increase levels of fertility and infant and child mortality.     

Son preference and its effects on Korean lactation practices

Using retrospective data from Korea, multivariate log-linear analyses were used to determine if the sex of a child influences the probability of it being breastfed, or if it influences the number of months he/she was breastfed. In Korea the likelihood of a woman not breastfeeding a child is very small. The fact that a woman is without living sons increased the average odds of not breastfeeding by 1.55 times. The absence of living sons has the strongest effect of any of the predictor variables used. The highest predicted conditional odds of not breastfeeding are for women with female infants, who live in rural areas, have less than 3 children of which none are boys, and who do not use oral contraceptives. The lowest predicted conditional odds of a woman not breastfeeding are for a city dweller, with 3 or more children of which at least 1 is a boy, who uses the pill, and whose infant is a boy. Although oral contraceptive use is significant for deciding how long to breastfeed, the direction of its effect is not what was expected had it affected lactation physiologically. Apparently, social factors are operative which make pill users more likely to breast feed than non-users. 1 possible explanation for this is that Korean women are aware of the fertility-suppressing effects of lactation and are using breastfeeding as a deliberate form of birth control. The fact that the effect of pill use becomes progressively stronger the longer that women breastfeed lends further support to this explanation. The effect of sex of child is not significant in determining the duration he/she will be breastfed. This study indicates support for the contention that Korean women are aware of the contraceptive effects of lactation, and that their decisions of breastfeed are affected by their preference for sons.     

Partnership status and the human sex ratio at birth

If two-parent care has different consequences for the reproductive success of sons and daughters, then natural selection may favour adjustment of the sex ratio at birth according to circumstances that forecast later family structure. In humans, this partnership-status hypothesis predicts fewer sons among extra-pair conceptions, but the rival 'attractiveness' hypothesis predicts more sons among extra-pair conceptions, and the 'fixed-phenotype' hypothesis predicts a constant probability of having a son, regardless of partnership status. In a sample of 86 436 human births pooled from five US population-based surveys, I found 51.5% male births reported by respondents who were living with a spouse or partner before the child's conception or birth, and 49.9% male births reported by respondents who were not (chi(2)=16.77 d.f.=1 p<0.0001). The effect was not explained by paternal bias against daughters, by parental age, education, income, ethnicity or by year of observation, and was larger when comparisons were made between siblings. To my knowledge, this is the first direct evidence for conditional adjustment of the sex ratio at birth in humans, and could explain the recent decline in the sex ratio at birth in some developed countries.     

High initiation and long duration of breastfeeding despite absence of early skin-to-skin contact in Karen refugees on the Thai-Myanmar border: a mixed methods study

Background

Early skin-to-skin contact (SSC) after birth is recommended as part of the United Nations Children’s Fund (UNICEF) baby friendly health initiative to promote optimum breastfeeding. This paper reports rates of breastfeeding initiation and duration in a low resource environment, where early SSC is not practised, and explores views of pregnant women and midwives surrounding breastfeeding and swaddling.

Methods

Data from records from a single hospital on the Thai-Myanmar border where refugee women gave birth during a one-year period (2010) were used to determine breastfeeding initiation rates and the time of the first breastfeed, and duration of breastfeeding of the previous alive child in multigravidae. Focus group discussions (FGD) were conducted to obtain information from pregnant women attending antenatal care about their intended or previous duration of breastfeeding and views on breastfeeding. Interviews with local midwives explored reasons for high rates of breastfeeding in this setting and the practice of newborn swaddling.

Results

Of 1404 live births in 2010 in Maela refugee camp there were 982 evaluable mother-newborn pairs, including 80 infants born before 37 weeks gestation. Initiation of breastfeeding within the first hour after birth and exclusive breastfeeding at discharge in term mother-newborn pairs was 91.2% (823/902) and 99.3% (896/902); and before 37 weeks gestation, 48.8% (39/80) and 98.8% (79/80). Reported duration of previous breastfeeding was 19 (range 2 to 72) months.During FGD all primigravidae (n?=?17) intended to breastfeed and all multigravidae (n?=?33) had previously breastfed; expected or previous duration of feeding was for more than one year or longer. The major theme identified during FGD was breastfeeding is “good”. Women stated their intention to breastfeed with certainty. This certainty was echoed during the interviews with midwifery staff. SSC requires a delay in early swaddling that in Karen people, with animistic beliefs, could risk loss of the spirit of the newborn or attract malevolent spirits.

Conclusions

In a population with a strong culture of breastfeeding and robust breastfeeding practices, high rates of initiation and duration of breastfeeding were found despite a lack of early skin-to-skin contact. Local preferences, traditions and practices that protect, support and maintain high rates of breastfeeding should be promoted.

     

Lack of Sex-Biased Maternal Investment in spite of a Skewed Birth Sex Ratio in White-Headed Langurs (Trachypithecus leucocephalus)

Numerous hypotheses have been developed to explain sex allocation. In male-dispersing, female cooperatively breeding species, the local resource competition model predicts male-biased birth sex ratio, the local resource enhancement model predicts female-biased birth sex ratio, and the population adjustment model predicts that biased birth sex ratio should not be favored if the two sexes are equally costly to rear. The male quality model predicts that, in polygynous species, females in better physical condition will either produce more sons than daughters or invest more heavily in sons than in daughters. White-headed langurs are a female philopatry and female cooperatively breeding species. During a 11-yr study, a total of 133 births were recorded, among which birth sex ratio (M:F = 73:49) was significantly male-biased. This is consistent with the prediction of the local resource competition model. On the other hand, if mothers balanced their investment between the two sexes, according to Fisher's population adjustment model, males should be the less-costly-to-rear sex. However, we found no sex difference for infant mortality (12.3% in males and 12.2% in females), and sons induced slightly longer interbirth interval (son: 26.4 ± 1.1 mo, daughter: 24.1 ± 0.6 mo) and lactational period (son: 20.9 ± 1.0 mo, daughters: 19.6 ± 0.5 mo) for their mothers. Thus, the population adjustment model was not supported by this study. The local resource enhancement model was not supported because birth sex ratio did not bias to females who provided more reproductive assistance. On the individual level, probit regression showed no relation between birth sex ratio and group size. Because the group size was considered to be negatively related to female physical condition, our study did not support the male-quality model. We suggested several possibilities to explain these results.     

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.     

Effects of maternal birth season on birth seasonality in the Canadian population during the seventeenth and eighteenth centuries

Birth records of the French-Canadian population for the period 1621-1765 were analyzed retrospectively to examine the effect of maternal birth season on the seasonal distribution of births. Preliminary examination indicated that there was a bimodal pattern in birth seasonality: a major peak in early spring, a trough in early summer, a minor peak in autumn, and a trough around December. Because this seasonality was strongly biased at the level of the first birth by the month of marriage, which was concentrated in November, the seasonality of nonfirst births (n = 32,926) was examined in relation to the four seasons of maternal birth. Mothers born in May-July showed a flatter monthly distribution of nonfirst births at a maternal age of 28 years or more. Analysis of marriage-first birth intervals indicated that mothers who married in August-October showed a lower percentage of immediate conception (intervals of 8-10 months), whereas those mothers born in May-July had a higher percentage of immediate conception. This difference in birth seasonality shown by mothers born in May-July is similar to results from early twentieth-century Japan. Some seasonal infertility factors could have affected the embryos at the earliest stage of pregnancy, modifying a part of the seasonal variation in birth rate.     

No evidence for sex biases in milk macronutrients,energy, or breastfeeding frequency in a sample of filipino mothers

Maternal reproductive investment includes both the energetic costs of gestation and lactation. For most humans, the metabolic costs of lactation will exceed those of gestation. Mothers must balance reproductive investment in any single offspring against future reproductive potential. Among mammals broadly, mothers may differentially invest in offspring based on sex and maternal condition provided such differences investment influence future offspring reproductive success. For humans, there has been considerable debate if there are physiological differences in maternal investment by offspring sex. Two recent studies have suggested that milk composition differs by infant sex, with male infants receiving milk containing higher fat and energy; prior human studies have not reported sex‐based differences in milk composition. This study investigates offspring sex‐based differences in milk macronutrients, milk energy, and nursing frequency (per 24 h) in a sample of 103 Filipino mothers nursing infants less than 18 months of age. We found no differences in milk composition by infant sex. There were no significant differences in milk composition of mothers nursing first‐born versus later‐born sons or daughters or between high‐ and low‐income mothers nursing daughters or sons. Nursing frequency also showed no significant differences by offspring sex, sex by birth order, or sex by maternal economic status. In the Cebu sample, there is no support for sex‐based differences in reproductive investment during lactation as indexed by milk composition or nursing frequency. Further investigation in other populations is necessary to evaluate the potential for sex‐based differences in milk composition among humans. Am J Phys Anthropol 152:209–216, 2013. © 2013 Wiley Periodicals, Inc.     

Breeding performance of a captive colony of rhesus macaques (Macaca mulatta)

For most of the 18 years recorded, fewer than 50% of the adult females gave birth in any one year. The colony, of 6 social groups, showed a clear-cut breeding season. Female parity and dominance had no effect on breeding rate, though 1st infants were born earlier in the year than 2nd-born ones. Only when females gave birth in successive years were the months of giving birth correlated. Mothers and daughters may tend to give birth closer in time within a breeding season than do other females.     

Puerperal morbidity: A neglected area of maternal health in Sri Lanka

Abstract

Data on women with at least two children are used to examine how the breastfeeding experience with the first child affects whether subsequent children are breastfed. Our results indicate that women most often repeat with later children the feeding decision they made with their first child. That is, those who breastfed their first child are very likely to breastfeed a later child and those who did not breastfeed their firstborn are unlikely to breastfeed a later‐bom. Among those who did not breastfeed their first child, education beyond high school increases the likelihood that they will switch to breastfeeding with a later‐born. Those who breastfed their first child are less likely to breastfeed a later‐born if the first breastfeeding experience was of short duration or was perceived to be unsuccessful or unsatisfactory or if the woman had not gone beyond high school or received anesthesia at the later birth. Hence, educational differences are greater at higher parities than at first parity.     

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.     

Variation of sex ratio and ABO blood group composition by month of birth of blood donors in Tokyo

Sex ratio of 17,273 blood donors born during the period between 1925 and 1935 was examined according to their month of birth and ABO blood groups in comparison with 5,810 healthy non-blood donors born in the 1900s to 1930s. The sex ratio of the blood donors and the non-blood donors varied similarly according to their month of birth with a prominent peak in summer births and a trough in winter births. This birth season with a high sex ratio was different from that of the general births during the period between 1921 and 1935, in which a maximum sex ratio was found in November. A possible explanation for the difference is the different rate of male and female infant deaths according to birth month. Variation of the sex ratio according to season of birth was not similar among the four ABO blood groups. Sex ratio of the donors with blood group B showed no elevation among the summer births. Non-blood donors with blood group B, on the contrary, showed a higher sex ratio than the others in the summer births. This difference can not be explained by infant or juvenile deaths. A possibility is that a tendency to become a blood donor is modified by the season of one's birth differently according to gender and ABO blood groups.     

Human sex ratio at birth and mother's birth season: multivariate analysis.

We used a population-based historical French Canadian database to examine the effects of mother's birth season on sex ratio at birth. Non-first births in the database (n = 127,658) were analyzed for their sex, parish size (2 large parishes of Montreal and Quebec or the other smaller parishes), time period (births up to 1719 or those from 1720), maternal age (< or = 24, 25-29, 30-34, 35+ years), sex of the preceding sibling (male or female), and birth seasons of the child and his or her parents (February-April, May-July, August-October, November-January). Season of child's birth significantly affected the sex ratio (chi 2 = 11.507, d.f. = 3, p = 0.009), with the births in February-April or May-July showing a lower sex ratio. Season of mother's birth also contributed highly significantly to the variation of sex ratio (chi 2 = 15.196, d.f. = 3, p = 0.002); mothers born in February-April had a low sex ratio among their children (sex ratio = 1.013). In contrast, season of father's birth did not affect the sex ratio (chi 2 = 0.618, d.f. = 3, p = 0.892). When a multiple logistic model was applied to the data, mother's birth season was the single most significant factor. The lower sex ratio from mothers born in February-April was observed consistently for every maternal age and delivery season. Seasonal influences on female fetuses seem to have changed their future reproductive characteristics.     

A retrospective study of infant mortality of cotton-top tamarins (Saguinus oedipus) in captive breeding

Fifty of 156 (32%) colony bred cotton-top tamarins were stillborn, and 31 (20%) died within the first week after birth. The stillbirth rate was related to litter size and parity in captivity. A higher percent of single births were stillborn (69%) compared to those of twin births (26%) (p less than 0.005). A higher percent of infants born of the first litter in captivity (42%) were stillborn than those of subsequent litters (23%) (p less than 0.025). Stillbirths were not related to season of birth or sex of infant. Survival of live infants was unrelated to season of birth, sex, litter size, or number of litters.     

Birthdate, mass and survival in mountain goat kids: effects of maternal characteristics and forage quality

In temperate environments, early-born ungulates may enjoy a longer growth period before winter, and so attain a higher body mass and an increased probability of survival compared to late-born ones. We assessed the effects of maternal characteristics, forage quality and population density on kid birthdate, mass and survival in a population of marked mountain goats (Oreamnos americanus) in Alberta. The duration and timing of the birth season were similar in all years. Births were highly synchronised: 80% of kids were born within 2 weeks of the first birth. Maternal age, maternal social rank and density did not affect kid birthdate or mass. Previous breeding experience was not related to kid birthdate, but kids born to pluriparous mothers were heavier during summer than kids born to primiparous mothers. Male and female kids had similar mass and accumulated mass linearly during summer. Early-born kids were heavier than late-born kids. Faecal crude protein (FCP) in late spring and maternal mass were positively related to kid mass. Survival to weaning appeared higher for males (90%) than for females (78%), but survival to 1 year was 65% for both sexes. FCP in late spring, density, birthdate and mass did not affect kid survival to weaning in either sex. Survival to 1 year increased with FCP in late spring for females, but not for males. Survival to 1 year was independent of birthdate for both sexes, but heavy females survived better than light ones. Multiple logistic regression revealed a positive effect of mass on survival to 1 year when the sexes were pooled. Our results suggest that mountain goats are constrained to give birth in a short birth season synchronised with forage productivity.     

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.     

Patterns of low birth weight in the Bengali newborns

This paper examines the distribution of low birth weight (2500 g or less) by gestation time, sex, maternal age, parity (birth order), socioeconomic conditions, and season of birth among 5117 single live births born to Bengali mothers at the Ramakrishna Mission Seva Pratisthan Hospital in Calcutta, India. Preterm infants have low birth weight significantly more often than their full term counterparts. Female infants have low birth weights significantly more often than male infants. The infants of poor mothers have lower birth weights in higher order births more often than infants of higher orders born to well-off mothers. Teenaged mothers produce low birth weight babies significantly more often than older mothers. Although the relationship is not significant, low birth weight infants occur more often among 1st and late born infants and less often among 2nd born infants. The season of birth is not significantly associated with birth weight. Less than 10% of low birth weight infants are pre-term, while the rest are full term. The great majority of low birth weight infants are small-for-gestational-age; the minority are small due to curtailed gestational age. The proportion of infants weighing less than 2001 g is only 9%; this figure tallies closely with earlier studies of India.     

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.