The prevalence of male homosexuality: the effect of fraternal birth order and variations in family size

The prevalence of male homosexuality probably varies over time and across societies. One reason for this variation may be the joint effect of two factors: (1) variations in fertility rate or family size; and (2) the fraternal birth order effect, the finding that the odds of male homosexuality increases with each additional older brother. Because of these effects, the rate of male homosexuality may be relatively high (at least in terms of sexual attraction if not behavior) in societies that have a high fertility rate, but this rate has probably declined somewhat in some, particularly western, societies. Thus, even if accurately measured in one country at one time, the rate of male homosexuality is subject to change and is not generalizable over time or across societies.     

The cause(s) of the fraternal birth order effect in male homosexuality

It has been established that the probability that a man is homosexual is positively related to his number of older brothers, but not older sisters when the brothers are accounted for. This is known as the 'fraternal birth order' effect. In the past, efforts have been made to explain this phenomenon in terms of several alternative biological hypotheses and a psychosocial hypothesis. This note examines how well these hypotheses accommodate the fraternal birth order effect. It is concluded that: (1) the evidence for the hypothesis of maternal immunoreactivity to the male fetus is weak; (2) the evidence for the intrauterine hormone exposure hypothesis is also weak; (3) the evidence for the hypothesis of postnatal learning is stronger. Lastly, there seem likely to be causes common to male homosexuality and paedophilia. They may include sexual (or quasi-sexual) experience in childhood or adolescence.     

Quantitative and theoretical analyses of the relation between older brothers and homosexuality in men

Meta-analysis of aggregate data from 14 samples representing 10,143 male subjects shows that homosexuality in human males is predicted by higher numbers of older brothers, but not by higher numbers of older sisters, younger brothers, or younger sisters. The relation between number of older brothers and sexual orientation holds only for males. This phenomenon has therefore been called the fraternal birth order effect. Research on birth order, birth weight, and sexual orientation suggests that the developmental pathway to homosexuality initiated by older brothers operates during prenatal life. Calculations assuming a causal relation between older brothers and sexual orientation have estimated the proportion of homosexual men who owe their sexual orientation to fraternal birth order at 15% in one study and 29% in another. The maternal immune hypothesis proposes that the fraternal birth order effect reflects the progressive immunization of some mothers to male-specific antigens by each succeeding male fetus and the increasing effects of such immunization on sexual differentiation of the brain in each succeeding male fetus. There are at least three possible mechanisms by which the mother's immune response could influence the fetus: the transfer of anti-male antibodies across the placenta from the maternal into the fetal compartment, the transfer of maternal cytokines across the placenta, and maternal immune reactions affecting the placenta itself. This hypothesis is consistent with recent studies showing that the quantity of fetal cells that enter the maternal circulation is greater than previously thought, and that the number of male-specific proteins encoded by Y-chromosome genes is greater than previously thought.     

The association between the fraternal birth order effect in male homosexuality and other markers of human sexual orientation

Later fraternal birth order (FBO) is a well-established correlate of homosexuality in human males and may implicate a maternal immunization response in the feminization of male sexuality. This has led to the suggestion that FBO may relate to other markers of male sexual orientation which are robustly sexually dimorphic. If so, among homosexual males the number of older brothers should strongly correlate with traits such as spatial ability and psychological gender, indicative of greater behavioural feminization, compared to heterosexual males. The present study failed to find significant associations between number of older brothers and these traits.     

Fraternal birth order and the maternal immune hypothesis of male homosexuality

In men, sexual orientation correlates with an individual's number of older brothers, each additional older brother increasing the odds of homosexuality by approximately 33%. It has been hypothesized that this fraternal birth order effect reflects the progressive immunization of some mothers to Y-linked minor histocompatibility antigens (H-Y antigens) by each succeeding male fetus and the concomitantly increasing effects of such maternal immunization on the future sexual orientation of each succeeding male fetus. According to this hypothesis, anti-H-Y antibodies produced by the mother pass through the placental barrier to the fetus and affect aspects of sexual differentiation in the fetal brain. This explanation is consistent with a variety of evidence, including the apparent irrelevance of older sisters to the sexual orientation of later born males, the probable involvement of H-Y antigen in the development of sex-typical traits, and the detrimental effects of immunization of female mice to H-Y antigen on the reproductive performance of subsequent male offspring. The maternal immune hypothesis might also explain the recent finding that heterosexual males with older brothers weigh less at birth than heterosexual males with older sisters and homosexual males with older brothers weigh even less than heterosexual males with older brothers.     

On the impact of sex and birth order on contact with kin

Previous research indicates that birth order is a strong predictor of familial sentiments, with middleborns less family-oriented than first- or last-borns. In this research, effects of sex and birth order on the actual frequency of contact with maternal and paternal kin were examined in two studies. In Study 1, one hundred and forty undergraduates completed a questionnaire relating to the amount of time they spent in contact with specific relatives, while in Study 2, one hundred and twelve undergraduates completed the same questionnaire with the addition of two questions relating to the subjects’ parents’ birth orders. Subjects were more likely to have frequent contact with maternal, as opposed to paternal, kin and women experienced more frequent contact than men with relatives in general. The birth order of subjects did not appear to have a significant influence on contact but the birth order of the subjects’ parents did, with the offspring of middleborn mothers having relatively little contact with maternal grandparents and the offspring of middleborn fathers having relatively little contact with paternal grandparents. These sex and birth order differences are discussed in relation to possible differences in how women and men use kinship ties and in terms of how birth order may influence parental solicitude. Catherine Salmon recently received her Ph.D. in psychology from McMaster University in Hamilton Ontario. Her interest in kinship and family relationships has grown out of her own large extended family and many visits to Utah as well as exposure to evolutionary thinking about the family in the lab of Martin Daly and Margo Wilson. Her other current research interests focus on female sexuality and the evolutionary study of literature.     

Changes in the eruption order of the first permanent tooth and their relation to season of birth in Japan

In order to examine the eruption order of the first two permanent teeth, kindergarten children 5 to 6 years old were examined in Hakone, Japan. Among a total of 817 children examined from 1976 to 1984, 349 were determined as I-type children, whose mandibular first incisor erupted earlier than the mandibular first molar, and 183 were as M-type children, whose first molar erupted earlier in the mandible. The mandibular I-type rate, i.e., the proportion of the I-type among a total of I- and M-type children, was 66% (349/532). In 1983-1984, the I-type rate was 70% among boys and 62% among girls, but the overall I-type rate did not differ significantly by the sex of the subject or by the year of examination. The I-type rate varied significantly with the season of subject's birth. Those born in October or November showed a significantly lower I-type rate (33%) than the other subjects (P less than 0.001), in spite of a shift of birth season with low I-type rate toward winter in 1983-1984. The difference of the I-type rate according to birth season suggests that the causes responsible for this change are primarily environmental and act at the prenatal or perinatal stage of life.     

Birth order and male androphilia in Samoan fa'afafine

The manner in which male androphilia is publicly expressed varies cross-culturally. As such, it is unclear whether distinct or common underlying causal processes characterize male androphilia in different cultures. Establishing the existence of cross-cultural universals in male androphilia is one means of ascertaining whether common biological bases underlie this phenomenon despite its culturally distinct forms. The evidence that the number of older biological brothers increases the odds of androphilia in later-born males has been well documented for Western samples (i.e. the fraternal birth order effect); but there is little evidence for this effect in non-Western samples. Here, we compare the birth order of androphilic males (i.e. fa'afafine) and gynephilic males from the politically autonomous Polynesian nation of Independent Samoa. Results indicate that relative to gynephilic males, fa'afafine tend to have more siblings and are generally later born when birth order is quantified using Slater, fraternal and sororal indices. More specifically, fa'afafine tend to have a greater number of older brothers, older sisters and younger brothers. We discuss the observed effects in relation to the differing reproductive patterns exhibited by the mothers of fa'afafine and gynephilic males, and to existing social and biological theories for sexual orientation.     

Fraternal birth order and birth weight in probably prehomosexual feminine boys

The purpose of this study was to confirm a previous finding that homosexual males with older brothers weigh less at birth than do heterosexual males with older brothers. The subjects comprised 250 feminine boys referred to a child psychiatry service because of extreme cross-gender wishes or behavior and assumed, on the basis of previous research, to be prehomosexual, plus 739 control boys and 261 control girls referred to the same service for reasons unrelated to sexual orientation or gender identity disorder and assumed, from base-rate probabilities, to be preheterosexual. The feminine boys with two or more older brothers weighed 385 g less at birth than did the control boys with two or more older brothers (P = 0.005). In contrast, the feminine and control boys with fewer than two older brothers did not differ in birth weight. This finding suggests that the mechanism by which older brothers increase the odds of homosexuality in later-born males operates prior to the individual's birth. We hypothesize that this mechanism may be immunologic, that antimale antibodies produced by human mothers in response to immunization by male fetuses could decrease the birth weight of subsequent male fetuses as well as increase their odds of homosexuality.     

Testosterone control of male courtship in birds

A sequence of behaviours which we call courtship initiates reproduction in a large number of species. In vertebrates, as a component of male sexual behaviour courtship is strongly influenced by testicular androgen. Here I will review some salient issues about the regulation of courtship by testosterone in birds. The first section will briefly summarize the first 100 years of research on this topic. The specific role of testosterone or its oestrogenic metabolites in the control of different components of courtship will be the subject of the second section. Then, I will discuss how behavioural patterns can be recruited into courtship and modified in their structure by testosterone action. In the following section, the role of sexual selection and female choice in shaping the link between testosterone and courtship will be addressed. The problematic nature of the quantitative relationships between testosterone and behaviour will be topic of the fifth section. Finally, I will discuss how courtship traits that are activated by testosterone can be apparently independent of hormone blood concentrations. These issues will be examined in an evolutionary perspective, in an attempt to understand how natural and sexual selection have shaped the links between the hormone and the behaviour.     

Consistent handedness of microtubule helical arrays in maize and Arabidopsis primary roots

Summary The orientation of cortical microtubules in plant cells has been extensively studied, in part because of their influence on the expansion of most plant cell types. Cortical microtubules are often arranged in helical arrays, which are well known to occur with a specific pitch as a function of development or experimental treatment; however, it is not known if the handedness of helical arrays can also be specified. We have studied the handedness of helical arrays by using Vibratome sectioning of maize primary roots and confocal microscopy of Arabidopsis primary roots. In cortical cells of maize roots, the helical array was found to have the same handedness at a given position, not only for the cells of a single root, but also for the cells of more than one hundred roots examined. Quantification of angular distribution of apparent individual microtubules showed that defined regions of the root were composed of cells with highly uniform microtubule orientation. In the region between transverse and longitudinal microtubules (5–10.5 mm from the tip), the array formed a right-handed helix, and basal of cells with longitudinal microtubules (11.5–15 mm from the tip), the array formed a left-handed helix. Similarly, in epidermal cells of Arabidopsis roots right-handed helical arrays were found in the region between transverse and longitudinal microtubules. These results suggest that, in addition to the orientation of microtubules, the handedness of helical microtubule arrays is under cellular control.Abbreviations Cy3 indocarbocyanine - PBS phosphate-buffered saline - PIPES piperazine-N,N-bis-[2-ethanesulfonic acid]     

Effects of cohort,sex, and birth date on body development of roe deer (Capreolus capreolus) fawns

We investigated the effects of the year of birth (cohort effect), sex and birth date on the variations observed in birth weight and postnatal growth rate in 209 newborn fawns marked during eight consecutive fawning seasons from a highly productive roe deer population under good nutritional conditions. Roe deer exhibited a fast body development with a birth weight of 1628 g and a postnatal growth rate of 139 g/day constant over the first 20 days of their life. As previously reported for dimorphic and polygynous ungulates, there was a marked cohort effect on the rate of body development. This variability was partly explained by climatic conditions during late gestation. When fewer than 5 days with rainfall over 5 mm occurred in April, fawns were heavier. High temperatures during April and during the winter could also be involved in fast body development of roe deer fawns. Sex of fawn did not affect roe deer growth pattern. This was expected on the basis of low sexual dimorphism in size and low polygyny level characteristic of roe deer. Lastly, the date of birth did not affect the body development of roe deer fawns. High constancy of mean birth dates and high synchrony of births observed in this population could account for this result.     

Changes in plasma,pituitary and brain α-MSH content in rats from birth to sexual maturity

Immunoreactive α-MSH was measured in plasma, pituitary and brain of male and female rats on the day of birth and at intervals afterwards up to 55–70 days of age. Plasma α-MSH concentrations on the day of birth were 528 ± 111 pg/ml and 406 ± 137 pg/ml in famel and male rats, respectively. Plasma α-MSH concentrations then fell and remained low until the onset of sexual maturity when they again rose reaching 406 ± 38 pg/ml in 70 day old females and 312 ± 46 pg/ml in 55 day old males. Pituitary α-MSH concentrations also changed with age and in male rats generally reflected the changes in plasma α-MSH concentrations. In females, on the other hand, pituitary α-MSH concentrations showed a gradual increase with age. Concentrations of α-MSH in the hypothalamus and brain increased with age and as with plasma and pituitary reached peak values in sexually mature animals. These findings are consistent with the idea that both piutiary and brain α-MSH have a role in sexually mature rats. Although α-MSH may have a role in sexual behaviour there was no evidence of any change in brain α-MSH throughout the estrous cycle.     

Fetal testosterone surge: specific modulations induced in male rats by maternal stress and/or alcohol consumption

Plasma testosterone (T) was measured in control male and female rats on gestational days 16, 17, 18, 19, and 20 and on days 17-20 in males from dams who were fed ethanol and/or were stressed during pregnancy. Circulating T in control males showed an earlier rise, yielding a longer period of prenatal T elevation, than was reported previously (Endocrinology 106 (1980)306). Compared to control males, exposure to alcohol-alone augmented T on days 18 and 19, stress-alone attenuated prenatal T, and the combination of stress and alcohol completely blocked the normal rise in T between days 17 and 18. When these prenatal alterations in T are viewed along with effects these same treatments have on the postparturient T surge (Horm. Behav. 41 (2002) 229), a possible explanatory mechanism emerges for the uniquely different behavioral patterns of sexual behavior differentiation induced in males by prenatal exposure to alcohol, stress, or both factors. Whereas the potential for feminine behavior is retained to the extent that either the prenatal or the neonatal T surge is attenuated, the male potential is more sensitive to reductions in the fetal surge and is maximally disrupted if both the prenatal and the postparturitional T surges are suppressed.     

Hand preferences for coordinated bimanual actions in 777 great apes: implications for the evolution of handedness in hominins

Whether or not nonhuman primates exhibit population-level handedness remains a topic of considerable scientific debate. Here, we examined handedness for coordinated bimanual actions in a sample of 777 great apes including chimpanzees, bonobos, gorillas, and orangutans. We found population-level right-handedness in chimpanzees, bonobos and gorillas, but left-handedness in orangutans. Directional biases in handedness were consistent across independent samples of apes within each genus. We suggest that, contrary to previous claims, population-level handedness is evident in great apes but differs among species as a result of ecological adaptations associated with posture and locomotion. We further suggest that historical views of nonhuman primate handedness have been too anthropocentric, and we advocate for a larger evolutionary framework for the consideration of handedness and other aspects of hemispheric specialization among primates.     

Neural correlates of sexual arousal in heterosexual and homosexual women and men

Most men have a category-specific pattern of genital and subjective sexual arousal, responding much more strongly to erotic stimuli depicting their preferred sex than to erotic stimuli depicting their nonpreferred sex. In contrast, women tend to have a less specific arousal pattern. To better understand this sex difference, we used neuroimaging to explore its neural correlates. Heterosexual and homosexual women viewed erotic photographs of either men or women. Evoked neural activity was monitored via fMRI and compared with responses to the same stimuli in heterosexual and homosexual men. Overall, a network of limbic (as well as the anterior cingulate) and visual processing regions showed significantly less category-specific activity in women than men. This was primarily driven by weaker overall activations to preferred-sex stimuli in women, though there was also some evidence of stronger limbic activations to nonpreferred-sex stimuli in women. Primary results were similar for heterosexual and homosexual participants. Women did show some evidence of category-specific responses in the visual processing regions, although even in these regions they exhibited less differential activity than men. In the anterior cingulate, a region with high concentrations of sex-hormone receptors, subjective and neural category specificity measures correlated positively for women but negatively for men, suggesting a possible sex difference in the role of the anterior cingulate. Overall, results suggest that men tend to show more differentiated neural responses than do women to erotic photographs of one sex compared to the other sex, though women may not be entirely indifferent to which sex is depicted.     

Father's physique influences mate preferences but not the actual choice of male somatotype in heterosexual women and homosexual men

Body constitution plays an important role in human mate choice. Cross-cultural research reports that women on average prefer men with muscular physique. It is still unclear, however, what mechanisms influence the inter-individual variation in mate preferences and choices of partner's physique. In this study, we tested the mechanisms of an imprinting-like effect (similarity between father and an ideal and actual partner) and of homogamy (similarity between self and an ideal and actual partner) for male physique in heterosexual women and homosexual men. To assess the variation in male physique, we employed somatotype paradigm which characterizes body constitution using three components: endomorphic (heavy-set), mesomorphic (muscular), and ectomorphic (lean). In total, 149 homosexual men and 769 heterosexual women from the Czech Republic indicated the somatotype of their father, ideal and actual partner, and in homosexual men also their own somatotype. In line with previous research, the somatotype most preferred by both men and women was the mesomorphic, followed by the ectomorphic and the endomorphic one. Women's preferences for an ideal partner somatotype weakly correlated with their fathers' somatotype, especially in women who reported a positive relationship with their fathers during childhood. Among homosexual men, we found imprinting-like preferences only for the ectomorphic somatotype component and no significant association with the quality of their relationships with their fathers. We also found no significant relationship between the fathers' and actual partners' somatotype in either heterosexual women or homosexual men. Our research indicates that fathers have a rather weak influence on mate preference for somatotypes and no influence on actual mate choice.     

Effects of testosterone on the development of neuromuscular systems and their target tissues involved in courtship and copulation in green anoles (Anolis carolinensis)

Male green anole lizards court females using a red throat fan (dewlap) and copulate by intromitting one of two penises (hemipenes). These structures begin sexually monomorphic, but by adulthood males have larger dewlaps, only males have hemipenes, and many of the neuromuscular components of both systems show male-biased dimorphisms. We hypothesized that testosterone (T), which increases in juvenile males but not females about a month after hatching, facilitates masculinization. To test this idea, on post-hatching day 30, gonadally intact females received either a blank or T implant, and males were either castrated or sham-castrated. At day 90, juveniles were euthanized and the length of the cartilage and cross-sectional areas of the muscle fibers and motoneurons required for dewlap extension were examined. We also measured the cross-sectional areas of the hemipenes and associated muscle fibers and motoneurons, and counted the motoneurons. T-treated females had longer cartilages and larger dewlap muscle fibers compared to those with blank implants. No effects on motoneurons were detected, and no females possessed hemipenes or associated musculature. In males, castration produced shorter dewlap cartilages and smaller hemipenes; other measures were not affected by treatment. These data indicate that components of the dewlap system differentiate relatively late in development, that T likely mediates the process, and that although components of the copulatory system are plastic in juvenile males, sexual differentiation of peripheral features is complete before day 30. The data also suggest that target structures (dewlap cartilage and hemipenes), compared to their neuromuscular effectors, are particularly sensitive to developmental T exposure.     

Estrogen treatment during development alters adult partner preference and reproductive behavior in female laboratory rats

There is broad acceptance for the idea that during development estradiol ‘organizes’ many aspects of reproductive behavior including partner preferences in the laboratory rat. With respect to partner preference, this idea is drawn from studies where estrogen action was in someway blocked, either through aromatase or estrogen receptor inhibition, during development in male rats. The lack of estrogens neonatally results in a decrease in the male rat's preference for females. In this study, the effect of early postnatal estradiol treatment on the partner preferences of female rats was examined as a further test of the hypothesis that male-typical partner preference is dependent upon early exposure to estrogens. Our principal finding was that increased postnatal estradiol exposure during development affected partner preference in the expected direction, and this effect was seen under several adult hormonal and behavioral testing conditions. Female rats that received exogenous estradiol during development spent more time with an estrous female and less time with a sexually active male than did cholesterol treated females. The estradiol treatment also disrupted normal female sexual behavior, receptivity, and proceptivity.