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Concept: Sexual selection

165

Conclusive evidence for sexual dimorphism in non-avian dinosaurs has been elusive. Here it is shown that dimorphism in the shape of the dermal plates of Stegosaurus mjosi (Upper Jurassic, western USA) does not result from non-sex-related individual, interspecific, or ontogenetic variation and is most likely a sexually dimorphic feature. One morph possessed wide, oval plates 45% larger in surface area than the tall, narrow plates of the other morph. Intermediate morphologies are lacking as principal component analysis supports marked size- and shape-based dimorphism. In contrast, many non-sex-related individual variations are expected to show intermediate morphologies. Taphonomy of a new quarry in Montana (JRDI 5ES Quarry) shows that at least five individuals were buried in a single horizon and were not brought together by water or scavenger transportation. This new site demonstrates co-existence, and possibly suggests sociality, between two morphs that only show dimorphism in their plates. Without evidence for niche partitioning, it is unlikely that the two morphs represent different species. Histology of the new specimens in combination with studies on previous specimens indicates that both morphs occur in fully-grown individuals. Therefore, the dimorphism is not a result of ontogenetic change. Furthermore, the two morphs of plates do not simply come from different positions on the back of a single individual. Plates from all positions on the body can be classified as one of the two morphs, and previously discovered, isolated specimens possess only one morph of plates. Based on the seemingly display-oriented morphology of plates, female mate choice was likely the driving evolutionary mechanism rather than male-male competition. Dinosaur ornamentation possibly served similar functions to the ornamentation of modern species. Comparisons to ornamentation involved in sexual selection of extant species, such as the horns of bovids, may be appropriate in predicting the function of some dinosaur ornamentation.

Concepts: Sexual dimorphism, Sexual selection, Ronald Fisher, Sexuality, Dinosaur, Jurassic, Stegosaurus, Morrison Formation

107

We analyzed one decade of data collected by the Programme for International Student Assessment (PISA), including the mathematics and reading performance of nearly 1.5 million 15 year olds in 75 countries. Across nations, boys scored higher than girls in mathematics, but lower than girls in reading. The sex difference in reading was three times as large as in mathematics. There was considerable variation in the extent of the sex differences between nations. There are countries without a sex difference in mathematics performance, and in some countries girls scored higher than boys. Boys scored lower in reading in all nations in all four PISA assessments (2000, 2003, 2006, 2009). Contrary to several previous studies, we found no evidence that the sex differences were related to nations' gender equality indicators. Further, paradoxically, sex differences in mathematics were consistently and strongly inversely correlated with sex differences in reading: Countries with a smaller sex difference in mathematics had a larger sex difference in reading and vice versa. We demonstrate that this was not merely a between-nation, but also a within-nation effect. This effect is related to relative changes in these sex differences across the performance continuum: We did not find a sex difference in mathematics among the lowest performing students, but this is where the sex difference in reading was largest. In contrast, the sex difference in mathematics was largest among the higher performing students, and this is where the sex difference in reading was smallest. The implication is that if policy makers decide that changes in these sex differences are desired, different approaches will be needed to achieve this for reading and mathematics. Interventions that focus on high-achieving girls in mathematics and on low achieving boys in reading are likely to yield the strongest educational benefits.

Concepts: Male, Female, Sexual dimorphism, Sexual selection, Gender, Sex, Student, Mating

81

Compelling evidence from many animal taxa indicates that male genitalia are often under postcopulatory sexual selection for characteristics that increase a male’s relative fertilization success. There could, however, also be direct precopulatory female mate choice based on male genital traits. Before clothing, the nonretractable human penis would have been conspicuous to potential mates. This observation has generated suggestions that human penis size partly evolved because of female choice. Here we show, based upon female assessment of digitally projected life-size, computer-generated images, that penis size interacts with body shape and height to determine male sexual attractiveness. Positive linear selection was detected for penis size, but the marginal increase in attractiveness eventually declined with greater penis size (i.e., quadratic selection). Penis size had a stronger effect on attractiveness in taller men than in shorter men. There was a similar increase in the positive effect of penis size on attractiveness with a more masculine body shape (i.e., greater shoulder-to-hip ratio). Surprisingly, larger penis size and greater height had almost equivalent positive effects on male attractiveness. Our results support the hypothesis that female mate choice could have driven the evolution of larger penises in humans. More broadly, our results show that precopulatory sexual selection can play a role in the evolution of genital traits.

Concepts: Sexual selection, Penis, Charles Darwin, Physical attractiveness, Sex organ, Human penis size, Secondary sex characteristic, Baculum

66

Post-copulatory sexual selection (PSS), fuelled by female promiscuity, is credited with the rapid evolution of sperm quality traits across diverse taxa. Yet, our understanding of the adaptive significance of sperm ornaments and the cryptic female preferences driving their evolution is extremely limited. Here we review the evolutionary allometry of exaggerated sexual traits (for example, antlers, horns, tail feathers, mandibles and dewlaps), show that the giant sperm of some Drosophila species are possibly the most extreme ornaments in all of nature and demonstrate how their existence challenges theories explaining the intensity of sexual selection, mating-system evolution and the fundamental nature of sex differences. We also combine quantitative genetic analyses of interacting sex-specific traits in D. melanogaster with comparative analyses of the condition dependence of male and female reproductive potential across species with varying ornament size to reveal complex dynamics that may underlie sperm-length evolution. Our results suggest that producing few gigantic sperm evolved by (1) Fisherian runaway selection mediated by genetic correlations between sperm length, the female preference for long sperm and female mating frequency, and (2) longer sperm increasing the indirect benefits to females. Our results also suggest that the developmental integration of sperm quality and quantity renders post-copulatory sexual selection on ejaculates unlikely to treat male-male competition and female choice as discrete processes.

Concepts: Male, Evolution, Sexual dimorphism, Sexual selection, Sex, Ronald Fisher, Evolutionary psychology, Charles Darwin

47

In many species, male secondary sexual traits have evolved via female choice as they confer indirect (i.e. genetic) benefits or direct benefits such as enhanced fertility or survival. In humans, the role of men’s characteristically masculine androgen-dependent facial traits in determining men’s attractiveness has presented an enduring paradox in studies of human mate preferences. Male-typical facial features such as a pronounced brow ridge, a more robust jawline may signal underlying health while beards may signal men’s age and masculine social dominance. However, masculine faces are judged as more attractive for short-term relationships over less masculine faces, while beards are judged as more attractive than clean-shaven faces for long-term relationships. Why such divergent effects occur between preferences for two sexually dimorphic traits remains unresolved. In the present study, we used computer graphic manipulation to morph male faces varying in facial hair from clean-shaven, light stubble, heavy stubble and full beards to appear more (+25% and +50%) or less (-25% and -50%) masculine. Women (N=8520) were assigned to treatments wherein they rated these stimuli for physical attractiveness in general, for a short-term liaison or a long-term relationship. Results showed a significant interaction between beardedness and masculinity on attractiveness ratings. Masculinized and, to an even greater extent, feminized faces were less attractive than unmanipulated faces when all were clean-shaven, and stubble and beards dampened the polarising effects of extreme masculinity and femininity. Relationship context also had effects on ratings, with facial hair enhancing long-term, and not short-term, attractiveness. Effects of facial masculinisation appears to have been due to small differences in the relative attractiveness of each masculinity level under the three treatment conditions and not to any change in the order of their attractiveness. Our findings suggest that beardedness may be attractive when judging long-term relationships as a signal of intra-sexual formidability and the potential to provide direct benefits to females. More generally, our results hint at a divergence of signalling function, which may result in a subtle trade-off in women’s preferences, for two highly sexually dimorphic androgen-dependent facial traits. This article is protected by copyright. All rights reserved.

Concepts: Male, Sexual selection, Gender, Interpersonal relationship, Man, Physical attractiveness, Facial hair, Beard

43

Hippocampal atrophy is found in many psychiatric disorders that are more prevalent in women. Sex differences in memory and spatial skills further suggest that males and females differ in hippocampal structure and function. We conducted the first meta-analysis of male-female difference in hippocampal volume (HCV) based on published MRI studies of healthy participants of all ages, to test whether the structure is reliably sexually dimorphic. Using four search strategies, we collected 68 matched samples of males' and females' uncorrected HCVs (in 4418 total participants), and 36 samples of male and female HCVs (2183 participants) that were corrected for individual differences in total brain volume (TBV) or intracranial volume (ICV). Pooled effect sizes were calculated using a random-effects model for left, right, and bilateral uncorrected HCVs and for left and right HCVs corrected for TBV or ICV. We found that uncorrected HCV was reliably larger in males, with Hedges' g values of 0.545 for left hippocampus, 0.526 for right hippocampus, and 0.557 for bilateral hippocampus. Meta-regression revealed no effect of age on the sex difference in left, right, or bilateral HCV. In the subset of studies that reported it, both TBV (g=1.085) and ICV (g= 1.272) were considerably larger in males. Accordingly, studies reporting HCVs corrected for individual differences in TBV or ICV revealed no significant sex differences in left and right HCVs (Hedges' g ranging from +0.011 to -0.206). In summary, we found that human males of all ages exhibit a larger HCV than females, but adjusting for individual differences in TBV or ICV results in no reliable sex difference. The frequent claim that women have a disproportionately larger hippocampus than men was not supported.

Concepts: Human, Male, Female, Sexual dimorphism, Sexual selection, Gender, Hippocampus, Sex

41

Gene expression differences between the sexes account for the majority of sexually dimorphic phenotypes, and the study of sex-biased gene expression is important for understanding the genetic basis of complex sexual dimorphisms. However, it has been difficult to test the nature of this relationship due to the fact that sexual dimorphism has traditionally been conceptualized as a dichotomy between males and females, rather than an axis with individuals distributed at intermediate points. The wild turkey (Meleagris gallopavo) exhibits just this sort of continuum, with dominant and subordinate males forming a gradient in male secondary sexual characteristics. This makes it possible for the first time to test the correlation between sex-biased gene expression and sexually dimorphic phenotypes, a relationship crucial to molecular studies of sexual selection and sexual conflict. Here, we show that subordinate male transcriptomes show striking multiple concordances with their relative phenotypic sexual dimorphism. Subordinate males were clearly male rather than intersex, and when compared to dominant males, their transcriptomes were simultaneously demasculinized for male-biased genes and feminized for female-biased genes across the majority of the transcriptome. These results provide the first evidence linking sexually dimorphic transcription and sexually dimorphic phenotypes. More importantly, they indicate that evolutionary changes in sexual dimorphism can be achieved by varying the magnitude of sex-bias in expression across a large proportion of the coding content of a genome.

Concepts: Gene, Genetics, Male, Evolution, Sexual dimorphism, Sexual selection, Gender, Sex

40

Opposing forces influence assortative mating so that one seeks a similar mate while at the same time avoiding inbreeding with close relatives. Thus, mate choice may be a balancing of phenotypic similarity and dissimilarity between partners. In the present study, we assessed the role of resemblance to Self’s facial traits in judgments of physical attractiveness. Participants chose the most attractive face image of their romantic partner among several variants, where the faces were morphed so as to include only 22% of another face. Participants distinctly preferred a “Self-based morph” (i.e., their partner’s face with a small amount of Self’s face blended into it) to other morphed images. The Self-based morph was also preferred to the morph of their partner’s face blended with the partner’s same-sex “prototype”, although the latter face was (“objectively”) judged more attractive by other individuals. When ranking morphs differing in level of amalgamation (i.e., 11% vs. 22% vs. 33%) of another face, the 22% was chosen consistently as the preferred morph and, in particular, when Self was blended in the partner’s face. A forced-choice signal-detection paradigm showed that the effect of self-resemblance operated at an unconscious level, since the same participants were unable to detect the presence of their own faces in the above morphs. We concluded that individuals, if given the opportunity, seek to promote “positive assortment” for Self’s phenotype, especially when the level of similarity approaches an optimal point that is similar to Self without causing a conscious acknowledgment of the similarity.

Concepts: Sexual selection, Choice, Similarity, Physical attractiveness, Assortative mating

34

Sperm competition theory predicts that animals face a trade-off between investment in weaponry and investment in ejaculate composition. Within the Madagascan giant hissing cockroaches (Tribe Gromphadorhini) differences in morphology exist that may indicate differing strategies of male-male competition. We compared relative pronotal horn length using high-resolution X-ray CT scanning data, relative testes mass, and male-male agonistic behaviour between two species of hissing cockroaches, Gromphadorhina oblongonota and Aeluropoda insignis. The gross morphology and behaviour of these two species indicated that G. oblongonota is selected for pre-copulatory mate acquisition and that A. insignis is selected for post-copulatory sperm competition. We found evidence for a trade-off when investing in testes mass vs. horn length between the species. The large, aggressive G. oblongonota follows a strategy of greater investment in weapons at the expense of testes mass while the smaller, less-aggressive A. insignis invests in relatively greater testes mass and less in pronotal weapon length. We also found evidence of a trade-off within each species, where individuals invest more heavily in weapon length at the expense of testes mass. These findings support the predictions of pre- and post-copulatory competitive investment trade-offs for a relatively understudied Tribe of cockroaches.

Concepts: Sperm, Sexual selection, Investment, Selection, Sperm competition

33

Sex differences in some preferences and motivations are well established, but it is unclear whether they persist in selective sub-populations, such as expert financial decision makers, top scientists, or elite athletes. We addressed this issue by studying competitiveness in 1,147 varsity intercollegiate distance runners. As expected, across all runners, men reported greater competitiveness with two previously validated instruments, greater competitiveness on a new elite competitiveness scale, and greater training volume, a known correlate of competitiveness. Among faster runners, the sex difference decreased for one measure of competitiveness but did not decrease for the two other competitiveness measures or either measure of training volume. Across NCAA athletic divisions (DI, DII, DIII), the sex difference did not decrease for any competitiveness or training measure. Further analyses showed that these sex differences could not be attributed to women suffering more injuries or facing greater childcare responsibilities. However, women did report greater commitment than men to their academic studies, suggesting a sex difference in priorities. Therefore, policies aiming to provide men and women with equal opportunities to flourish should acknowledge that sex differences in some kinds of preferences and motivation may persist even in selective sub-populations.

Concepts: Natural selection, Male, Sexual dimorphism, Sexual selection, Gender, Sex, Motivation, Mating