Male Traits Research Articles

Experimental immune challenges reduce the quality of male antennae and female pheromone output

Sexual signalling is a key feature of reproductive investment, yet the effects of immune system activation on investment into chemical signalling, and especially signal receiver traits such as antennae, are poorly understood. We explore how upregulation of juvenile immunity affects male antennal functional morphology and female pheromone attractiveness in the gumleaf skeletonizer moth, Uraba lugens. We injected final-instar larvae with a high or low dose of an immune elicitor or a control solution and measured male antennal morphological traits, gonad investment and female pheromone attractiveness. Immune activation affected male and female signalling investment: immune challenged males had a lower density of antennal sensilla, and the pheromone of immune-challenged females was less attractive to males than their unchallenged counterparts. Immune challenge affected female investment into ovary development but not in a linear, dose-dependent manner. While there was no effect of immune challenge on testes size, there was a trade-off between male pre- and post-copulatory investment: male antennal length was negatively correlated with testes size. Our study highlights the costs of elaborate antennae and pheromone production and demonstrates the capacity for honest signalling in species where the costs of pheromone production were presumed to be trivial.

Multiple phenotypic traits predict male mating success in a critically endangered frog

Complex sexual signals spanning multiple sensory modalities may be common in nature, yet few studies have explored how combinations of phenotypic traits influence male attractiveness and mating success. Here, we investigate whether combinations of multiple male phenotypic traits (both within and across sensory modalities) predict male mating and fertilization success in the critically endangered southern corroboree frog, Pseudophryne corroboree. We conducted breeding trials in a standardized captive environment where females were given the opportunity to choose between multiple males over the duration of the breeding season. For each male, we measured multiple call traits, aspects of coloration, body size, and age. We found that complex interactions between multiple traits best predicted male mating and fertilization success. In general, males with lower call frequency, lower call rate, and shorter call duration had the highest mating and fertilization success. Fertilization success was additionally linked to male body size and age. These findings suggest that female P. corroboree select mates based on a suite of acoustic traits, adding to a growing body of evidence that females use multiple traits to assess male quality. Our results also suggest that females may combine information from multiple signals non-additively. Moreover, our results imply that females gain direct fertility benefits from their mate choice decisions. We argue that understanding female mate choice based on various signals across multiple sensory modalities has important implications for the integration of mate choice into conservation breeding programs and needs to be considered when developing behavior-based captive breeding strategies.Significance statementSexual signals are often highly complex, yet we know little about how multiple signal components both within and across various sensory modalities predict male mating success. We investigated whether combinations of multiple phenotypic traits (within and across sensory modalities) predicted male breeding success in threatened corroboree frogs. We conducted captive breeding trials in a homogeneous environment, where females could choose between multiple males over the duration of a single breeding season. We found that interactions between multiple male traits predicted mating and fertilization success. Males with lower call frequency, call rate, and duration had higher mating success. Fertilization success was also linked to acoustic signals, body size, and age. Understanding mate choice for multiple traits further elucidates the complexity of female mate choice. This study is one of the first to consider the conservation implications of multimodal signaling in mate choice.

Don’t Look Down

Don’t Look Down

The Shape of Sex: Nonbinary Gender from Genesis to the Renaissance

The Shape of Sex: Nonbinary Gender from Genesis to the Renaissance

Male and female identity and environmental contexts influence courtship behaviour in a songbird

Behavioural variation in courtship has become a central theme in the study of sexual selection. Courtship behaviour can vary consistently between males (between-individual variation) due to inherent characteristics of individuals, but males may also plastically adjust their courtship (within-individual variation) in response to the characteristics of the potential breeding mate or the environmental contexts. However, the relative importance of these components in courtship behaviour remains poorly understood. Here, we exposed male collared flycatchers, Ficedula albicollis , to different stimulus females simulating a female's visit to the male's territory during mate choice. We measured the male's latency to explicitly invite the female to visit the nestbox (hereafter, invitation latency), as a proxy of the male's motivation to mate. Repeated measurements of focal males (exposed to different stimulus females) and stimulus females (presented to different males) allowed us to assess the relative importance of male and female identity in male courtship, while also assessing the importance of their phenotype and other environmental sources of within-individual variation in invitation latency. We found that invitation latency was determined by male and, to a lesser extent, female identity, with the latter suggesting the existence of male mate choice in the species. We failed to identify specific traits of males and females that influenced courtship behaviour, although male responsiveness to stimulus females increased as the season advanced, possibly due to the costs of delaying breeding in temperate breeding areas. Overall, these results suggest that invitation latency is a consistent courtship attribute in males that can be adjusted plastically according to perceived female quality and environmental conditions. We emphasize that understanding within- and between-individual sources of courtship variation is essential for assessing its evolutionary potential, as plasticity in the expression of sexual signals may affect their reliability and the strength of sexual selection on them. • We simulated female visits to the male's territory and measured courtship. • Courtship behaviour varied consistently between males. • Males adjusted their courtship according to perceived female identity. • Courtship behaviour was also sensitive to seasonality. • Plasticity in sexual signal expression may affect the strength of sexual selection.

Genetic analysis of sexual dimorphism in growth of Jamunapari goats of India.

A genetic study of sexual size dimorphism (SSD) in Jamunapari goats was carried out to identify differences between sexes in genetic control of body weight at birth and at 3 (weaning), 6, 9 and 12months of age. A total of 6,687 kids out of 264 sires and 1,704 dams were used in the study. Estimates of SSD were derived from male:female body weight ratios. Males were 9.9, 6.8, 9.3, 13.7 and 16.8% heavier than females at birth and at 3, 6, 9 and 12months of age, respectively, demonstrating modest SSD. Phenotypic standard deviations were larger for male kids and tended to be proportional to body weight means. However, males also had somewhat greater phenotypic coefficients of variation and total resemblance among relatives. Additive direct genetic correlations between body weights of males and females exceeded 0.96 at birth and weaning. Corresponding additive maternal correlations exceeded 0.99. Additive direct correlations between sexes were somewhat less than unity for postweaning weights but exceeded 0.80 in multi-trait models. Our results indicated that body weights could be treated as the same trait in males and females, but the use of different phenotypic variances for the two sexes would improve accuracies of breeding value predictions. High genetic correlations between body weights in males and females suggest limited opportunity to use sex-specific selection to create or modify SSD or create divergent body weight phenotypes between sexes.

Very short mountings are enough for sperm transfer inLittorina saxatilis

ABSTRACTConflict over reproduction between females and males exists because of anisogamy and promiscuity. Together they generate differences in fitness optima between the sexes and result in antagonistic coevolution of female and male reproductive traits. Mounting duration is likely to be a compromise between male and female interests whose outcome depends on the intensity of sexual selection. The timing of sperm transfer during mounting is critical. For example, mountings may be interrupted before sperm is transferred as a consequence of female or male choice, or they may be prolonged to function as mate guarding. In the highly promiscuous intertidal snail Littorina saxatilis, mountings vary substantially in duration, from less than a minute to more than an hour, and it has been assumed that mountings of a few minutes do not result in any sperm being transferred. Here, we examined the timing of sperm transfer, a reproductive trait that is likely affected by sexual conflict. We performed time-controlled mounting trials using L. saxatilis males and virgin females, aiming to examine indirectly when the transfer of sperm starts. We observed the relationship between mounting duration and the proportion of developing embryos out of all eggs and embryos in the brood pouch. Developing embryos were observed in similar proportions in all treatments (i.e. 1, 5 and 10 or more minutes at which mountings were artificially interrupted), suggesting that sperm transfer begins rapidly (within 1 min) in L. saxatilis and very short matings do not result in sperm shortage in the females. We discuss how the observed pattern can be influenced by predation risk, population density, and female status and receptivity.

Hybrid Seed Set in Relation with Male Floral Traits, Estimation of Heterosis and Combining Abilities for Yield and Its Components in Wheat (Triticum aestivum L.).

Breeding hybrids with maximum heterosis requires efficient cross-pollination and an improved male sterility system. Renewed efforts have been made to dissect the phenotypic variation and genetic basis of hybrid floral traits, although the potential of tailoring the appropriate flower design on seed setting is less known. To this end, elite wheat genotypes were crossed using a chemical hybridizing agent at different doses. A total of 23 hybrids were developed from a partial diallel design; and planted in an alpha lattice design with their parents at two locations in Morocco, for two years, to evaluate for yield components, heterosis and combining abilities. The 13.5 L ha−1 dose induced a maximum level of sterility (95%) and seed set showed large phenotypic variation and high heritability. In parallel, seed set showed tight correlation with pollen mass (0.97), visual anther extrusion (0.94) and pollen shedding (0.91) (p < 0.001), allowing direct selection of the associated traits. Using the combined data, mid-parent heterosis ranges were −7.64–14.55% for biomass (BM), −8.34–12.51% for thousand kernel weight (TKW) and −5.29–26.65% for grain yield (YLD); while best-parent heterosis showed ranges of −11.18–7.20%, −11.35–11.26% and −8.27–24.04% for BM, TKW and YLD, respectively. The magnitude of general combining ability (GCA) variance was greater than the specific combining ability (SCA) variance suggesting a greater additive gene action for BM, TKW and YLD. The favorable GCA estimates showed a simple method to predict additive effects contributing to high heterosis and thus could be an effective approach for the selection of promising parents in early generations.

Mapping sex differences in the effects of protein and carbohydrates on lifespan and reproduction in Drosophila melanogaster: is measuring nutrient intake essential?

Understanding how diet affects reproduction and survival is a central aim in evolutionary biology. Although this relationship is likely to differ between the sexes, we lack data relating diet to male reproductive traits. One exception to this general pattern is Drosophila melanogaster, where male dietary intake was quantified using the CApillary FEeder (CAFE) method. However, CAFE feeding reduces D. melanogaster survival and reproduction, so may distort diet-fitness outcomes. Here, we use the Geometric Framework of Nutrition to create nutrient landscapes that map sex-specific relationships between protein, carbohydrate, lifespan and reproduction in D. melanogaster. Rather than creating landscapes with consumption data, we map traits onto the nutrient composition of forty agar-based diets, generating broad coverage of nutrient space. We find that male and female lifespan was maximised on low protein, high carbohydrate blends (~ 1P:15.9C). This nutrient ratio also maximised male reproductive rates, but females required more protein to maximise daily fecundity (1P:1.22C). These results are consistent with CAFE assay outcomes. However, the approach employed here improved female fitness relative to CAFE assays, while effects of agar versus CAFE feeding on male fitness traits depended on the nutrient composition of experimental diets. We suggest that informative nutrient landscapes can be made without measuring individual nutrient intake and that in many cases, this may be preferable to using the CAFE approach. The most appropriate method will depend on the question and species being studied, but the approach adopted here has the advantage of creating nutritional landscapes when dietary intake is hard to quantify.

Frequent origins of traumatic insemination involve convergent shifts in sperm and genital morphology.

Traumatic insemination is a mating behavior during which the (sperm) donor uses a traumatic intromittent organ to inject an ejaculate through the epidermis of the (sperm) recipient, thereby frequently circumventing the female genitalia. Traumatic insemination occurs widely across animals, but the frequency of its evolution, the intermediate stages via which it originates, and the morphological changes that such shifts involve remain poorly understood. Based on observations in 145 species of the free‐living flatworm genus Macrostomum, we identify at least nine independent evolutionary origins of traumatic insemination from reciprocal copulation, but no clear indication of reversals. These origins involve convergent shifts in multivariate morphospace of male and female reproductive traits, suggesting that traumatic insemination has a canalizing effect on morphology. We also observed sperm in both the sperm receiving organ and within the body tissue of two species. These species had intermediate trait values indicating that traumatic insemination evolves through initial internal wounding during copulation. Finally, signatures of male‐female coevolution of genitalia across the genus indicate that sexual selection and sexual conflict drive the evolution of traumatic insemination, because it allows donors to bypass postcopulatory control mechanisms of recipients.

Genome-wide association study of beef bull semen attributes

BackgroundCattle production is dependent upon fertility because it results in producing offspring to offset production costs. A number of semen attributes are believed to affect fertility and are frequently measured as part of routine breeding soundness exams or semen collection procedures. The objective of this study was to perform a single-step genome-wide association study (ssGWAS) for beef bull semen attributes. Beef bull fertility phenotypes including volume (VOL), concentration (CONC), number of spermatozoa (NSP), initial motility (IMot), post-thaw motility (PTMot), three-hour post-thaw motility (3HRPTMot), percentage of normal spermatozoa (%NORM), primary abnormalities (PRIM), and secondary abnormalities (SEC) were obtained from two artificial insemination (AI) centers. A total of 1819 Angus bulls with 50,624 collection records were used for ssGWAS. A five-generation pedigree was obtained from the American Angus Association and consisted of 6521 sires and 17,136 dams. Genotypes on 1163 bulls were also obtained from the American Angus Association and utilized in ssGWAS.ResultsA multi-trait animal model was used for the estimation of single nucleotide polymorphism (SNP) effects. Significant SNP were those with a -log10P-value threshold greater than 4.0. Volume, CONC, NSP, IMot, PTMot, 3HRPTMot, %NORM, PRIM, and SEC have five, three, six, seven, two, six, six, and two genome-wide significant SNP, respectively.ConclusionsSeveral significant SNP were determined to be near or within quantitative trait loci (QTL) associated with beef bull semen attributes. In addition, genes associated with fertility were found to contain or be near the significant SNP found in the study. The results indicate there are regions of the genome that impact fertility, proving inclusion of genomic information into genetic evaluation should be advantageous for genetic improvement of male fertility traits.

Strong spatial population structure shapes the temporal coevolutionary dynamics of costly female preference and male display

Female mating preferences for exaggerated male display traits are commonplace. Yet, comprehensive understanding of the evolution and persistence of costly female preference through indirect (Fisherian) selection in finite populations requires some explanation for the persistence of additive genetic variance (V a) underlying sexual traits, given that directional preference is expected to deplete V a in display and hence halt preference evolution. However, the degree to which V a, and hence preference‐display coevolution, may be prolonged by spatially variable sexual selection arising solely from limited gene flow and genetic drift within spatially structured populations has not been examined. Our genetically and spatially explicit model shows that spatial population structure arising in an ecologically homogeneous environment can facilitate evolution and long‐term persistence of costly preference given small subpopulations and low dispersal probabilities. Here, genetic drift initially creates spatial variation in female preference, leading to persistence of V a in display through “migration‐bias” of genotypes maladapted to emerging local sexual selection, thus fueling coevolution of costly preference and display. However, costs of sexual selection increased the probability of subpopulation extinction, limiting persistence of high preference‐display genotypes. Understanding long‐term dynamics of sexual selection systems therefore requires joint consideration of coevolution of sexual traits and metapopulation dynamics.

Association of the MAOA-uVNTR polymorphism with psychopathic traits may change from childhood to adolescence.

Psychopathic traits can lead to violence, making it a serious public health concern. Genetic factors contribute to the aetiology of psychopathy. We examined whether monoamine oxidase A (MAOA-uVNTR) was associated with psychopathic traits measured quantitatively from controls through clinically aggressive youth (n = 336). Subjects were sub-categorized into at or above, andbelow age 13years. Results reveal that males below age 13 were more likely to display psychopathic traits with the MAOA long variant, whereasmales above age 13yearsweremore likely to display withthe short variant. This suggests that developmental factors may be crucial for understanding the role of the MAOA polymorphism in psychopathic traits in males.

Evidence for niche conservatism in alpine beetles under a climate‐driven species pump model

AbstractAimPast glacial cycles have generated lineage diversity in alpine habitats, acting as a climate‐driven species pump. It is not clear how much this process contributes to ecological diversification of alpine species. To examine this problem, we test patterns of genetic and phenotypic divergence in two co‐distributed species complexes of flightless alpine ground beetles. Greater differentiation in functional traits would indicate that ecological selection is an outcome of oscillating climate change, whereas greater differentiation in non‐ecological traits would indicate niche conservatism.LocationThe Cascades Range and Trinity Mountains of western North America.TaxonNebria paradisiandN.vandykeispecies complexes (Insecta: Coleoptera: Carabidae: Nebriinae).MethodsWe generated genome‐wide single nucleotide polymorphism data and mitochondrial sequence data, as well as morphological and physiological data, to compare populations spanning the range of both species. Phylogenetic and population genetic analyses were used to infer relationships among taxa and populations in each species complex, as well as historical demography. Support vector machines were used to test for classification of taxa and populations based on ecomorphological, ecophysiological and male reproductive traits. Mantel tests were then used to assess associations between phenotypic and genetic divergence among populations.ResultsTheN.vandykeiandN.paradisispecies complexes are each comprised of genetically distinctive populations exhibiting long‐term demographic declines. Each phylogeny supports multiple monophyletic groups with geographical cohesion. By examining phenotypic traits among populations in both species’ complexes, we show that reproductive trait divergence can discriminate species and population status more effectively than ecomorphological or ecophysiological traits. Reproductive and genetic divergence are significantly correlated in theN.vandykeispecies complex.Main ConclusionsWe found limited evidence of ecological selection acting on functional traits. Instead, reproductive and genetic divergence evolved among isolated populations in both species complexes, suggesting niche conservatism may be a common outcome in alpine species diversification.

Bull fertility and semen quality are not correlated with dairy and production traits in Brown Swiss cattle

Undisturbed reproduction is key for successful breeding of beef and dairy cattle. Improving reproductive ability can be difficult because of antagonistic relationships with other economically relevant traits. In cattle, thorough investigation of female fertility revealed unfavorable genetic correlations with various production phenotypes. However, the correlation between male reproductive ability and production traits remains poorly understood. Here, we investigated the genetic relationships among and between male fertility characteristics and economically relevant traits in a population of Brown Swiss cattle. We performed GWAS with imputed genotypes at nearly 12 million sequence variants for semen quality (sperm head and tail anomalies, motility, concentration, and volume), male fertility, and 57 production phenotypes. Allele substitution effects were then correlated on a trait-by-trait basis to estimate genetic correlations. Correlations between male reproductive characteristics and traits of economic value were small and ranged from −0.0681 to 0.0787. Among the semen quality parameters, sperm motility was negatively correlated with anomalies (head: r = −0.7083 ± 0.0002; tail: r = −0.7739 ± 0.0002) and volume (r = −0.1266 ± 0.0003), whereas volume was negatively correlated with concentration (r = −0.3503 ± 0.0002). Sire nonreturn rate was negatively correlated with sperm anomalies (head: r = −0.1640 ± 0.0002; tail: r = −0.1580 ± 0.0002) and positively correlated with motility (r = 0.1598 ± 0.0002). A meta-analysis of male reproductive traits identified 2 quantitative trait loci: a previously described region on chromosome 6 showed pleiotropic effects and a novel region on chromosome 11 was associated with sperm head anomalies. In conclusion, our results suggest that selection for economically important dairy and production phenotypes has little impact on semen quality and fertility of Brown Swiss bulls.

The significant role of post-pairing male behavior on the evolution of male preferences and female traits

Existing sexual selection theory postulates that a sufficiently large variation in female fecundity or other direct benefits are fundamental for generating male mate choice. In this study, we suggest that, in addition to pre-pairing preferences, choosy males can also have different post-pairing behaviors, a factor which has been comparatively overlooked by previous studies. We found that both male preferences and female traits could evolve much more easily than previously expected when the choosy males that paired with unpreferred females would allocate more efforts to seeking additional post-pairing mating opportunities. Furthermore, a costly female trait could evolve when there was a trade-off between seeking additional mating and paternal care investment within social pair for choosy males. Finally, a costly male preference and a costly female trait might still evolve and reach a stable polymorphic state in the population, which might give rise to a high variability in male choice and female traits in nature. We suggest that male mate choice may be even more common than expected, which needs to be verified empirically.

Relationships between male secondary sexual traits, physiological state and offspring viability in the three-spined stickleback

BackgroundSexual signals produced by males play a central role in sexual selection, but the relationship between these traits and the quality of the bearer are often ambiguous. Secondary sexual traits may represent genetic quality of the bearer, resulting in positive relationships with physiological state, or may be costly to produce, showing trade-off with physiological state. A number of studies have explored the relationships between secondary sexual traits and other functional traits, but few have studied their fitness consequences. We studied the link between diverse physiological traits and both morphological and behavioural sexual traits and examined how their interplay influences offspring viability in the three-spined stickleback.ResultsMale sticklebacks showing nest building and courtship behaviour were smaller than those not investing in reproductive activities. There was no evidence that the expression of red nuptial colouration and the quality of courtship behaviour of males are positively related to their metabolic rates, swim ability, oxidative damage and mtDNA copy number. However, individuals showing larger red nuptial colour areas had higher levels of oxidative DNA damage in their sperm. Male courtship behaviour and aggressiveness, but not red colour area, were good predictors of offspring hatching and survival.ConclusionsOur results suggest that, in our study population at the southern edge of the species’ distribution, sexual colouration of male sticklebacks was not a good indicator of their body state, but both courtship quality and aggressiveness during the courtship are reliable cues of their gamete quality, influencing the viability of their offspring. Thus, females that choose mates based on their courtship behaviour will have high fitness. In the study population, which represents a fast pace-of-life with high reproductive rate and short lifespan, sexual ornaments of males may not honestly signal their physiological and physical state because they invest at maximum in a single reproductive season despite high costs.

Sex-specific associations of adiposity with cardiometabolic traits in the UK: A multi-life stage cohort study with repeat metabolomics.

BackgroundSex differences in cardiometabolic disease risk are commonly observed across the life course but are poorly understood and may be due to different associations of adiposity with cardiometabolic risk in females and males. We examined whether adiposity is differently associated with cardiometabolic trait levels in females and males at 3 different life stages.Methods and findingsData were from 2 generations (offspring, Generation 1 [G1] born in 1991/1992 and their parents, Generation 0 [G0]) of a United Kingdom population-based birth cohort study, the Avon Longitudinal Study of Parents and Children (ALSPAC). Follow-up continues on the cohort; data up to 25 y after recruitment to the study are included in this analysis. Body mass index (BMI) and total fat mass from dual-energy X-ray absorptiometry (DXA) were measured at mean age 9 y, 15 y, and 18 y in G1. Waist circumference was measured at 9 y and 15 y in G1. Concentrations of 148 cardiometabolic traits quantified using nuclear magnetic resonance spectroscopy were measured at 15 y, 18 y, and 25 y in G1. In G0, all 3 adiposity measures and the same 148 traits were available at 50 y. Using linear regression models, sex-specific associations of adiposity measures at each time point (9 y, 15 y, and 18 y) with cardiometabolic traits 3 to 6 y later were examined in G1. In G0, sex-specific associations of adiposity measures and cardiometabolic traits were examined cross-sectionally at 50 y. A total of 3,081 G1 and 4,887 G0 participants contributed to analyses. BMI was more strongly associated with key atherogenic traits in males compared with females at younger ages (15 y to 25 y), and associations were more similar between the sexes or stronger in females at 50 y, particularly for apolipoprotein B–containing lipoprotein particles and lipid concentrations. For example, a 1 standard deviation (SD) (3.8 kg/m2) higher BMI at 18 y was associated with 0.36 SD (95% confidence interval [CI] = 0.20, 0.52) higher concentrations of extremely large very-low-density lipoprotein (VLDL) particles at 25 y in males compared with 0.15 SD (95% CI = 0.09, 0.21) in females, P value for sex difference = 0.02. By contrast, at 50 y, a 1 SD (4.8 kg/m2) higher BMI was associated with 0.33 SD (95% CI = 0.25, 0.42) and 0.30 SD (95% CI = 0.26, 0.33) higher concentrations of extremely large VLDL particles in males and females, respectively, P value for sex difference = 0.42. Sex-specific associations of DXA-measured fat mass and waist circumference with cardiometabolic traits were similar to findings for BMI and cardiometabolic traits at each age. The main limitation of this work is its observational nature, and replication in independent cohorts using methods that can infer causality is required.ConclusionsThe results of this study suggest that associations of adiposity with adverse cardiometabolic risk begin earlier in the life course among males compared with females and are stronger until midlife, particularly for key atherogenic lipids. Adolescent and young adult males may therefore be high priority targets for obesity prevention efforts.

Toward a Nonbinary Model of Gender/Sex Traits

AbstractI argue against the exclusive female/male divide, referring to the phenomenon of epistemic injustice in the cases of people with nonbinary gender identities and people with intersex traits. Such people have traits that are counterexamples to the binary female/male model. I have separated female and male traits into nine basic layers, five of which belong to sex (chromosomes, gonads, internal sex organs, external genitals, and secondary sex characteristics) and four to gender (gender identity, legal gender, external gender presentation, and gender pronouns). In every layer, I have found traits that are neither female nor male, and the application of the model to individuals provides examples of clusters of traits for which one layer is male and another female. Such traits and clusters of traits create the category of the nonbinary. Table 1 provides a sketch of a nonbinary model. The nonbinary category takes its name from the existing category of nonbinary gender identity; however, in the current model, it is a third category of traits, not of people. Under the nonbinary model, the basic gender concepts do not disappear.Sis a woman ifSis a human being with enough female traits, and the trait of having self-determined female gender identity is sufficient but not necessary.

Effects of artificial selection for moving activity on male reproductive traits and leg length in the red flour beetle Tribolium castaneum

Effects of artificial selection for moving activity on male reproductive traits and leg length in the red flour beetle <i>Tribolium castaneum</i>