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Sex‐ratio theory predicts that parents can optimise their fitness by producing offspring of the rare sex, yet there is a dearth of empirical evidence for adaptive sex allocation in response to the adult sex ratio (ASR). This is concerning, as anthropogenic disruption of the sex ratios of reproductive individuals threatens to cause demographic collapse in animal populations. Species with environmental sex determination (ESD) are especially at risk but may possess the capacity to adaptively influence offspring sex via control over the developmental environment. For example, reptiles with temperature‐dependent sex determination (TSD) could conceivably choose nest sites with thermal characteristics that produce offspring of the rare sex. To test this hypothesis, we seeded three secure outdoor ponds with different sex ratios (~M:F 3:1, 1:1, and 1:3) of adult painted turtles (Chrysemys picta), a reptile species with TSD. We then quantified nesting traits that could influence nest temperature and thus offspring sex ratio, including nesting date, nest depth, and nest canopy cover. We found no directional relationship between the ASR treatments and any measured nest traits and thus rejected our hypothesis. Interestingly, increased maternal body size was associated with reduced nest canopy cover, and this trend was more pronounced in the biased ASR treatments. If adaptive sex allocation occurs in this system, it instead may manifest via maternal epigenetic predisposition of offspring sex or in response to a phenomenon other than the ASR. [ABSTRACT FROM AUTHOR]

Small populations of imperiled species are susceptible to the negative consequences of skewed sex‐ratios. In imperiled species with environmental sex determination such as sea turtles, examining sex ratios across a range of environments and population abundance levels can provide insight into factors that influence population resilience, which can then be the foci of management plans for these species. Breeding sex ratios (the ratio of actively breeding males to females during a reproductive season; BSRs) extrapolated from genetic parentage analyses are a common approach for enumerating sex ratios in sea turtles. Such analyses also allow for the characterization of multiple paternity within sea turtle clutches, which should reflect BSRs and breeding behaviors. We characterized the first BSR for a breeding assemblage of loggerhead sea turtles (Caretta caretta) belonging to the temperate, low‐abundance Northern Gulf of Mexico Recovery Unit using genotypes of 16 microsatellite loci from nesting females and hatchlings. Unlike prior studies at both more‐tropical and more‐temperate, and higher‐abundance, Recovery Units in this region, we found a balanced BSR of 1.3:1 males:female and a low incidence (~17%) of multiple paternity. This suggests that there are relatively few males breeding at this assemblage and within this Recovery Unit. Beaches in this region are expected to produce substantial numbers of male hatchlings based on sand temperature data. The relative dearth of mature males may then be due to hydrologic disturbances that disproportionately affect the fitness and survival of male hatchlings, or due to demographic stochasticity. More work is needed to study the factors that might influence male hatchling production and fitness in this region, particularly as climate change is predicted to lead to feminization in global sea turtle populations. Our work demonstrates the broad utility of characterizing BSRs and other sex ratios across a range of populations in imperiled, environmentally sensitive species. [ABSTRACT FROM AUTHOR]

Haldane’s rule occupies a special place in biology as one of the few ‘rules’ of speciation, with empirical support from hundreds of species. And yet, its classic purview is restricted taxonomically to the subset of organisms with heteromorphic sex chromosomes. I propose explicit acknowledgement of generalized hypotheses about Haldane’s rule that frame sex bias in hybrid dysfunction broadly and irrespective of the sexual system. The consensus view of classic Haldane’s rule holds that sex-biased hybrid dysfunction across taxa is a composite phenomenon that requires explanations from multiple causes. Testing of the multiple alternative hypotheses for Haldane’s rule is, in many cases, applicable to taxa with homomorphic sex chromosomes, environmental sex determination, haplodiploidy, and hermaphroditism. Integration of a variety of biological phenomena about hybrids across diverse sexual systems, beyond classic Haldane’s rule, will help to derive a more general understanding of the contributing forces and mechanisms that lead to predictable sex biases in evolutionary divergence and speciation. [ABSTRACT FROM AUTHOR]

ABSTRACT Sex‐ratio theory predicts that parents can optimise their fitness by producing offspring of the rare sex, yet there is a dearth of empirical evidence for adaptive sex allocation in response to the adult sex ratio (ASR). This is concerning, as anthropogenic disruption of the sex ratios of reproductive individuals threatens to cause demographic collapse in animal populations. Species with environmental sex determination (ESD) are especially at risk but may possess the capacity to adaptively influence offspring sex via control over the developmental environment. For example, reptiles with temperature‐dependent sex determination (TSD) could conceivably choose nest sites with thermal characteristics that produce offspring of the rare sex. To test this hypothesis, we seeded three secure outdoor ponds with different sex ratios (~M:F 3:1, 1:1, and 1:3) of adult painted turtles (Chrysemys picta), a reptile species with TSD. We then quantified nesting traits that could influence nest temperature and thus offspring sex ratio, including nesting date, nest depth, and nest canopy cover. We found no directional relationship between the ASR treatments and any measured nest traits and thus rejected our hypothesis. Interestingly, increased maternal body size was associated with reduced nest canopy cover, and this trend was more pronounced in the biased ASR treatments. If adaptive sex allocation occurs in this system, it instead may manifest via maternal epigenetic predisposition of offspring sex or in response to a phenomenon other than the ASR.

The cladoceran crustacean Daphnia exhibits phenotypic plasticity, a phenomenon that leads to diverse phenotypes from one genome. Alternative usage of gene isoforms has been considered a key gene regulation mechanism for controlling different phenotypes. However, to understand the phenotypic plasticity of Daphnia, gene isoforms have not been comprehensively analyzed. Here we identified 25,654 transcripts derived from the 9710 genes expressed during environmental sex determination of Daphnia magna using the long-read RNA-Seq with PacBio Iso-Seq. We found that 14,924 transcripts were previously unidentified and 5713 genes produced two or more isoforms. By a combination of Illumina short-read RNA-Seq, we detected 824 genes that implemented switching of the highest expressed isoform between females and males. Among the 824 genes, we found isoform switching of an ortholog of CREB-regulated transcription coactivator, a major regulator of carbohydrate metabolism in animals, and a correlation of this switching event with the sexually dimorphic expression of carbohydrate metabolic genes. These results suggest that a comprehensive catalog of isoforms may lead to understanding the molecular basis for environmental sex determination of Daphnia. We also infer the applicability of the full-length isoform analyses to the elucidation of phenotypic plasticity in Daphnia. [ABSTRACT FROM AUTHOR]

Influenced by two mechanisms of genetic sex determination and environmental sex determination, the sex control of fish is of great significance for aquaculture. Compared with other vertebrates, fishes exhibit great plasticity in sex differentiation and gonadal development. There are significant differences in the size and growth rate of male and female teleosts. Cultivating and breeding large and fast-growing unisexual populations is beneficial to improve economic benefits. Mastacembelus armatus is a kind of freshwater aquaculture fish with high nutritional and economic values. Male individuals grow faster and have a larger body size than females, therefore, cultivating all-male M. armatus has higher economic benefits.In the wild environment, the male to female ratio of the M. armatus is approximately 1∶1, however, there is a high female rate under artificial breeding conditions. At present, it has not been clear what factors cause the feminization of M. armatus, and there have been no successful reports of inducing physiological female sex reversal in the M. armatus, which greatly restricts the development of the M. armatus breeding industry and the improvement of economic benefits. At present, researches on the M. armatus mainly focus on morphology, physiology, reproductive biology, genetic diversity, and other aspects. The research on sex determination and differentiation of the M. armatus is relatively weak, and it is necessary to explore ideas from the sex determination mechanisms, sex regulation methods and sex control breeding techniques of other teleosts. Based on the mechanism of sex determination in teleosts, and the review summarizes several ways of sex control in teleosts, mainly including genetic factors regulation like sex determining genes on sex chromosomes and autosomes, as well as environmental factors regulation, such as temperature, exogenous hormones and fish feed. It aims to provide ideas for analyzing the causes and potential mechanism forhigh feminization of artificial cultivation of the M. armatus, and to provide important theoretical basis for the reproduction and maintenance of unisexual populations, the selection of excellent varieties, and the protection of germplasm resources of the M. armatus, to better understand the gonadal development and reproductive mechanism of the M. armatus, solve the bottleneck of low male rate in currently cultivated M. armatus and provide guarantees for the sustainable development of the M. armatus aquaculture industry.

Phenotypic plasticity is often posited as an avenue for adaptation to environmental change, whereby environmental influences on phenotypes could shift trait expression toward new optimal values. Conversely, plastic trait expression may inhibit adaptation to environmental change by reducing selective pressure on ill-adapted traits. While plastic responses are often assumed to be linear, nonlinear phenotype–environment relationships are common, especially in thermally sensitive traits. Here we examine nonlinear plasticity in a trait with great ecological and evolutionary significance: sexual phenotype in species with environmental sex determination (ESD). In species with ESD, development switches between male and female at an environmental threshold (the inflection point). The inflection point is a key trait for adaptive responses to changing environments and should evolve toward the new optimum in order to maintain evolutionarily stable sex ratios. We used an individual-based theoretical model to investigate how two forms of plasticity in the ESD reaction norm—the nonlinear slope of the reaction norm and a linear shift in the inflection point—influence the evolution of the inflection point under climate warming. We found that steeper reaction norm slopes (high nonlinear plasticity) promoted evolution toward new optimal phenotypes (higher inflection points). In contrast, increased linear plasticity in the inflection point (shift) hindered adaptive evolution. Additionally, populations in moderate warming scenarios showed greater adaptive evolution of the inflection point compared with populations in extreme warming scenarios, suggesting that the proximity of existing phenotypes to new optimal phenotypes influences evolutionary outcomes. Unexpectedly, we found greater population persistence under high climate variability, due to the increased production of rare-sex individuals in unusually cold years. Our results demonstrate that different forms of phenotypic plasticity have crucially different effects on adaptive evolution. Plasticity that prevented sex ratio bias hindered the evolution of the inflection point, while plasticity that exacerbated sex ratio bias promoted adaptation to environmental change. [ABSTRACT FROM AUTHOR]

Influenced by two mechanisms of genetic sex determination and environmental sex determination, the sex control of fish is of great significance for aquaculture. Compared with other vertebrates, fishes exhibit great plasticity in sex differentiation and gonadal development. There are significant differences in the size and growth rate of male and female teleosts. Cultivating and breeding large and fast-growing unisexual populations is beneficial to improve economic benefits. Mastacembelus armatus is a kind of freshwater aquaculture fish with high nutritional and economic values. Male individuals grow faster and have a larger body size than females, therefore, cultivating all-male M. armatus has higher economic benefits. In the wild environment, the male to female ratio of the M. armatus is approximately 1∶1, however, there is a high female rate under artificial breeding conditions. At present, it has not been clear what factors cause the feminization of M. armatus, and there have been no successful reports of inducing physiological female sex reversal in the M. armatus, which greatly restricts the development of the M. armatus breeding industry and the improvement of economic benefits. At present, researches on the M. armatus mainly focus on morphology, physiology, reproductive biology, genetic diversity, and other aspects. The research on sex determination and differentiation of the M. armatus is relatively weak, and it is necessary to explore ideas from the sex determination mechanisms, sex regulation methods and sex control breeding techniques of other teleosts. Based on the mechanism of sex determination in teleosts, and the review summarizes several ways of sex control in teleosts, mainly including genetic factors regulation like sex determining genes on sex chromosomes and autosomes, as well as environmental factors regulation, such as temperature, exogenous hormones and fish feed. It aims to provide ideas for analyzing the causes and potential mechanism forhigh feminization of artificial cultivation of the M. armatus, and to provide important theoretical basis for the reproduction and maintenance of unisexual populations, the selection of excellent varieties, and the protection of germplasm resources of the M. armatus, to better understand the gonadal development and reproductive mechanism of the M. armatus, solve the bottleneck of low male rate in currently cultivated M. armatus and provide guarantees for the sustainable development of the M. armatus aquaculture industry. [ABSTRACT FROM AUTHOR]

Sex determination (SD) is a crucial developmental process, but its molecular underpinnings are very diverse, both between and within species. SD mechanisms have traditionally been categorized as either genetic (GSD) or environmental (ESD), depending on the type of cue that triggers sexual differentiation. However, mixed systems, with both genetic and environmental components, are more prevalent than previously thought. Here, we show theoretically that environmental effects on expression levels of genes within SD regulatory mechanisms can easily trigger within-species evolutionary divergence of SD mechanisms. This may lead to the stable coexistence of multiple SD mechanisms and to spatial variation in the occurrence of different SD mechanisms along environmental gradients. We applied the model to the SD system of the housefly, a global species with world-wide latitudinal clines in the frequencies of different SD systems, and found that it correctly predicted these clines if specific genes in the housefly SD system were assumed to have temperature-dependent expression levels. We conclude that environmental sensitivity of gene regulatory networks may play an important role in diversification of SD mechanisms. [ABSTRACT FROM AUTHOR]

Following Parker's 1970 publication, sperm biologists began to consider how sperm competition may drive the evolution of sperm cell morphology [[20]], a line of inquiry that is ongoing and continues to add new insight into sperm evolution to this day. This complexity is further highlighted by the work of Hook and colleagues [[36]], which explores how sperm head shape can influence the cell's migration and finds that sperm motility is influenced by the apical hook - a hook structure on the head of sperm cells that is a common feature of rodent sperm and has been hypothesized to facilitate sperm-sperm interactions. 10.3390/cells10061358 36 Hook K.A., Wilke L.M., Fisher H.S. Apical Sperm Hook Morphology Is Linked to Sperm Swimming Performance and Sperm Aggregation in Peromyscus Mice. This has resulted in massive leaps in our understanding of, among other things, sperm form and function, reproductive strategies, sperm development, and how the environments in which sperm operate shape sperm evolution. [Extracted from the article]

Geckos (Gekkota), the species-rich clade of reptiles with more than 2200 currently recognized species, demonstrate a remarkable variability in diploid chromosome numbers (2n = 16–48) and mode of sex determination. However, only a small fraction of gekkotan species have been studied with cytogenetic methods. Here, we applied both conventional (karyotype reconstruction and C-banding) and molecular (fluorescence in situ hybridization with probes for rDNA loci and telomeric repeats) cytogenetic analyses in seven species of geckos, namely Blaesodactylus boivini, Chondrodactylus laevigatus, Gekko badenii, Gekko cf. lionotum, Hemidactylus sahgali, Homopholis wahlbergii (Gekkonidae) and Ptyodactylus togoensis (Phyllodactylidae), in order to provide further insights into the evolution of karyotypes in geckos. Our analysis revealed the presence of interstitial telomeric repeats in four species, but we were not able to conclude if they are remnants of previous chromosome rearrangements or were formed by an accumulation of telomeric-like satellite motifs. Even though sex chromosomes were previously identified in several species from the genera Hemidactylus and Gekko by cytogenetic and/or genomic methods, they were not detected by us in any examined species. Our examined species either have poorly differentiated sex chromosomes or, possibly, environmental sex determination. Future studies should explore the effect of temperature and conduct genome-wide analyses in order to identify the mode of sex determination in these species. [ABSTRACT FROM AUTHOR]

The article presents a study which explores about the genomic imprinting-like monoallelic paternal expression determines sex of channel catfish. It mentions the mechanism of sex determination provides insights into the plasticity of genetic sex determination in lower vertebrates and its connection with temperature sex determination where DNA methylation as broadly involved.

Simple Summary: Environmental sex determination is a modality of sex determination related to external factors and that has implicated determinants such as climatic conditions, which act on the embryo after fertilization and deposition of the egg. For reptiles, the temperature is the main element for sex determination; this factor affects laid eggs in different ways. Details remain to be elucidated concerning the temporal gene expression and the functions of their protein products. Therefore, the aim of the present work was to determine the genetic determinants differentially represented during the embryonic development of a model species already known in temperature-dependent sex determination, the leopard gecko Eublepharis macularius. Following this investigation, new data were acquired on genes expressed in the sexual differentiation of E. macularius. In addition, new genes potentially involved in the mechanisms of tissue and metabolic sexual differentiation of the embryo of this species have been identified. This study could bring new useful information in order to correctly interpret the regulatory pathway underlying the determination of sex in vertebrates. During development, sexual differentiation results in physiological, anatomical and metabolic differences that implicate not only the gonads but also other body structures. Sex in Leopard geckos is determined by egg incubation temperature. Based on the premise that the developmental decision of gender does not depend on a single gene, we performed an analysis on E. macularius to gain insights into the genes that may be involved in gonads' sexual differentiation during the thermosensitive period. All the genes were identified as differentially expressed at stage 30 during the labile phase of sex differentiation. In this way, the expression of genes known to be involved in gonadal sexual differentiation, such as WNT4, SOX9, DMRT1, Erα, Erβ, GnRH, P450 aromatase, PRLand PRL-R, was investigated. Other genes putatively involved in sex differentiation were sought by differential display. Our findings indicate that embryo exposure to a sex-determining temperature induces differential expression of several genes that are involved not only in gonadal differentiation, but also in several biological pathways (ALDOC, FREM1, BBIP1, CA5A, NADH5, L1 non-LTR retrotransposons, PKM). Our data perfectly fit within the new studies conducted in developmental biology, which indicate that in the developing embryo, in addition to gonadal differentiation, sex-specific tissue and metabolic polarization take place in all organisms. [ABSTRACT FROM AUTHOR]

Long-day length and high temperature inhibit sex expression in pumpkin (Cucurbita moschata Duch.), and therefore directly impact the production potential. In this study, female flowering patterns in photoperiod-insensitive (PPIS) and photoperiod-sensitive (PPS) germplasms differed significantly in a moderately long day and high temperature environment. However, both germplasms exhibited a similar response in short day with either low temperature or high temperature environment. Photoperiod sensitivity led to this difference in sex expression between the germplasms. For the traits of 1st female flowering node (FFFN) and number of female flowers (NFF), high-density linkage map construction and quantitative trait locus (QTL) mapping were performed using SLAF-seq technology and 162 F2 individuals generated from PPIS and PPS. In total, 4 655 SLAFs were selected and mapped on 20 linkage groups (LGs). The total map length was 2 502.01 cM with an average interval distance of 0.75 cM. Major QTLs for both FFFN and NFF were detected on LG6 with intervals of 7.89 and 17.67 cM and PVE values of 30.5% and 22.9%, respectively. Further analyses of the major locus for FFFN revealed 73 protein-coding genes. Among them, 4 were related to sex expression, photoperiod flowering, and hormone response. An InDel (insertion and deletion) marker partially correlated with FFFN of the F2 population was also developed. Our study identified the QTL for the sex expression response to environmental factors using the high-density linkage map. The identified candidate genes and markers will provide useful information about the molecular interaction between the environment and sex expression and for marker-assisted selection of pumpkin environment-insensitive resources. [ABSTRACT FROM AUTHOR]

Sexual vs. asexual reproduction—unisexual vs. bisexual populations—diploid vs. polyploid biotypes—genetic vs. environmental sex determination: all these natural phenomena are associated with the genus of teleost fish, Carassius. This review places emphasis on two Carassius entities with completely different biological characteristics: one globally widespread and invasive Carassius gibelio, and the other C. carassius with a decreasing trend of natural occurrence. Comprehensive biological and cytogenetic knowledge of both entities, including the physical interactions between them, can help to balance the advantages of highly invasive and disadvantages of threatened species. For example, the benefits of a wide-ranged colonization can lead to the extinction of native species or be compensated by parasitic enemies and lead to equilibrium. This review emphasizes the comprehensive biology and cytogenetic knowledge and the importance of the Carassius genus as one of the most useful experimental vertebrate models for evolutionary biology and genetics. Secondly, the review points out that effective molecular cytogenetics should be used for the identification of various species, ploidy levels, and hybrids. The proposed investigation of these hallmark characteristics in Carassius may be applied in conservation efforts to sustain threatened populations in their native ranges. Furthermore, the review focuses on the consequences of the co-occurrence of native and non-native species and outlines future perspectives of Carassius research. [ABSTRACT FROM AUTHOR]

Medaka (Oryzias latipes) is a teleost fish with an XX/XY sex determination system. Sex reversal from female-to-male (masculinization of XX fish) can be induced through cortisol elevation from exposure to environmental stress such as high temperature during sexual differentiation. However, the effects of oxidative stress, generated via metabolic reactions and biological defense mechanisms, on the sexual differentiation of medaka are unclear. Here, we investigated the effect of oxidative stress on medaka sexual differentiation using hydrogen peroxide (H2O2), which induces oxidative stress in vertebrates. H2O2 treatment from 0 to 5 days post-hatching induced masculinization of wild-type XX medaka, but not of gonadal soma-derived growth factor (gsdf) or peroxisome proliferator-activated receptor alpha-a (pparaa) knockout XX fish. Co-treatment with an oxidative stress inhibitor caused masculinization recovery but co-treatment with a cortisol synthesis inhibitor did not. H2O2 treatment significantly upregulated gsdf and pparaa expression in XX medaka. However, H2O2 did not elevate cortisol levels in medaka larvae during sexual differentiation. These results strongly indicate that oxidative stress induces masculinization of XX medaka without causing elevation of cortisol. [ABSTRACT FROM AUTHOR]

Sex determination mechanisms evolve surprisingly rapidly, yet little is known in the large nematode phylum other than for Caenorhabditis elegans, which relies on chromosomal XX-XO sex determination and a dosage compensation mechanism. Here we analyze by sex-specific genome sequencing and genetic analysis sex determination in two fungal feeding/plant-parasitic Bursaphelenchus nematodes and find that their sex differentiation is more likely triggered by random, epigenetic regulation than by more well-known mechanisms of chromosomal or environmental sex determination. There is no detectable difference in male and female chromosomes, nor any linkage to sexual phenotype. Moreover, the protein sets of these nematodes lack genes involved in X chromosome dosage counting or compensation. By contrast, our genetic screen for sex differentiation mutants identifies a Bursaphelenchus ortholog of tra-1, the major output of the C. elegans sex determination cascade. Nematode sex determination pathways might have evolved by "bottom-up" accretion from the most downstream regulator, tra-1. In most species, sex is determined by genetic or environmental factors. Here, the authors present evidence that sex determination in Bursaphelenchus nematodes is instead likely to be regulated by a random, epigenetic mechanism. [ABSTRACT FROM AUTHOR]

The article presents a study which explores how truncated jarid2 and kdm6b transcripts are associated with temperature-induced sex reversal during development in a dragon lizard. It mentions that the work verifies that chromatin-modifying genes are involved in highly conserved temperature responses and can also be transcribed into isoforms with new sex-determining roles.

Discussion Our study demonstrated that the low (24 °C) and high (32 °C) incubation temperatures yielded significantly female-biased hatchlings, while the medium temperatures (around 28 °C) produced male-biased hatchlings or an even sex ratio of hatchlings in I G. japonicus i . Keywords: geographic variation; reptile; sex determination; sex ratio; thermal environment EN geographic variation reptile sex determination sex ratio thermal environment N.PAG N.PAG 10 04/26/22 20220415 NES 220415 1. The dataset provided daily mean air temperatures, daily maximum air temperatures, and daily minimum air temperatures of 30-year means: (a) mean air temperature; (b) maximum air temperature; (c) minimum air temperature. In our species, maternally derived yolk steroid hormones interact with incubation temperatures to affect offspring sex ratio, with substantial effects of yolk hormones on offspring sex ratio at the medium temperature of 28 °C [[14]]. [Extracted from the article]

Medaka (Oryzias latipes) is a teleost fish with an XX/XY sex determination system. Sex reversal from female-to-male (masculinization of XX fish) can be induced through cortisol elevation from exposure to environmental stress such as high temperature during sexual differentiation. However, the effects of oxidative stress, generated via metabolic reactions and biological defense mechanisms, on the sexual differentiation of medaka are unclear. Here, we investigated the effect of oxidative stress on medaka sexual differentiation using hydrogen peroxide (H2O2), which induces oxidative stress in vertebrates. H2O2 treatment from 0 to 5 days post-hatching induced masculinization of wild-type XX medaka, but not of gonadal soma-derived growth factor (gsdf) or peroxisome proliferator-activated receptor alpha-a (pparaa) knockout XX fish. Co-treatment with an oxidative stress inhibitor caused masculinization recovery but co-treatment with a cortisol synthesis inhibitor did not. H2O2 treatment significantly upregulated gsdf and pparaa expression in XX medaka. However, H2O2 did not elevate cortisol levels in medaka larvae during sexual differentiation. These results strongly indicate that oxidative stress induces masculinization of XX medaka without causing elevation of cortisol.

The cladoceran crustacean Daphnia produces only females by parthenogenesis in a healthy population. However, in response to environmental declines such as crowding and lack of foods, it produces eggs destined to become males that are genetically identical to females. During the development of the sexually committed eggs, DM domain-containing transcription factor Doublesex1 (Dsx1) orchestrates male trait formation globally both in somatic and gonadal tissues. Recent studies have revealed that Dsx1 expression is tightly controlled at transcriptional, post-transcriptional, and epigenetic levels to avoid sexual ambiguity. In this review, together with basic information on Dsx1 structure and expression, we introduce the multi-layered Dsx1 regulation and discuss how each regulation is interconnected for controlling male development in environmental sex-determining Daphnia.

The factors affecting sex determination remain largely unknown for most bivalve species. Environmental factors, such as temperature, are known to influence sex determination in some species, and this has been hypothesized for the mussel Mytilus edulis , but not experimentally validated yet. Adult exposure to different environmental conditions during gametogenesis, which occurs seasonally, may also affect offspring phenotype, including sex determination. Intergenerational carryover effects have been reported in bivalves, but the impact of parental exposures on offspring sex determination has not been examined so far. To address these questions, artificial fertilizations were performed on individuals collected in three different years and their embryos were reared at three different temperatures to specifically test if the environment influences offspring sex ratio through effects on parental developing gametes and/or on developing embryos. The analysis of 1938 embryos from 25 crosses revealed a significant effect of both temperature treatments and sampling years on the proportion of females. However, the overall variation is mostly explained by the "temperature treatment × sampling years" interaction. Taken together, our results suggested that interannual environmental variations experienced by the parents (during gametogenesis) and embryo (during development) can influence the sex of the offspring in M. edulis. [ABSTRACT FROM AUTHOR]

Sex ratio theory predicts both mean sex ratio and variance under a range of population structures. Here, we compare two genera of phoretic nematodes (Parasitodiplogaster and Ficophagus spp.) associated with 12 fig pollinating wasp species in Panama. The host wasps exhibit classic local mate competition: only inseminated females disperse from natal figs, and their offspring form mating pools that consist of scores of the adult offspring contributed by one or a few foundress mothers. By contrast, in both nematode genera, only sexually undifferentiated juveniles disperse and their mating pools routinely consist of 10 or fewer adults. Across all mating pool sizes, the sex ratios observed in both nematode genera are consistently female-biased (approx. 0.34 males), but markedly less female-biased than is often observed in the host wasps (approx. 0.10 males). In further contrast with their hosts, variances in nematode sex ratios are also consistently precise (significantly less than binomial). The constraints associated with predictably small mating pools within highly subdivided populations appear to select for precise sex ratios that contribute both to the reproductive success of individual nematodes, and to the evolutionary persistence of nematode species. We suggest that some form of environmental sex determination underlies these precise sex ratios. [ABSTRACT FROM AUTHOR]

Sexual systems are surprisingly diverse, considering the ubiquity of sexual reproduction. Sequential hermaphroditism, the ability of an individual to change sex, has emerged multiple times independently across the animal kingdom. In molluscs, repeated shifts between ancestrally separate sexes and hermaphroditism are generally found at the level of family and above, suggesting recruitment of deeply conserved mechanisms. Despite this, molecular mechanisms of sexual development are poorly known. In molluscs with separate sexes, endocrine disrupting toxins bind the retinoid X receptor (RXR), activating ectopic male development in females, suggesting the retinoid pathway as a candidate controlling sexual transitions in sequential hermaphrodites. We therefore tested the role of retinoic acid signaling in sequentially hermaphroditic Crepidula snails, which develop first into males, then change sex, maturing into females. We show that retinoid agonists induce precocious penis growth in juveniles and superimposition of male development in females. Combining RXR antagonists with retinoid agonists significantly reduces penis length in induced juveniles, while similar treatments using retinoic acid receptor (RAR) antagonists increase penis length. Transcripts of both receptors are expressed in the induced penis. Our findings therefore show that retinoid signaling can initiate molluscan male genital development, and regulate penis length. Further, we show that retinoids induce ectopic male development in multiple Crepidula species. Species-specific influence of conspecific induction of sexual transitions correlates with responsiveness to retinoids. We propose that retinoid signaling plays a conserved role in molluscan male development, and that shifts in the timing of retinoid signaling may have been important for the origins of sequential hermaphroditism within molluscs. [Display omitted] • Retinoid agonists induce penis growth in sequentially hermaphroditic snails. • Blocking RXR or retinoic acid synthesis decreases penis length. • RXR , RAR , and CYP2 6 mRNA are expressed in the induced penis. • Shifts in retinoid timing may be involved in repeated evolution of hermaphroditism. [ABSTRACT FROM AUTHOR]

With or without sex chromosomes, sex determination is a synthesis of many molecular events that drives a community of cells towards a coordinated tissue fate. In this review, we will consider how a sex determination pathway can be engaged and stabilized without an inherited genetic determinant. In many reptilian species, no sex chromosomes have been identified, yet a conserved network of gene expression is initiated. Recent studies propose that epigenetic regulation mediates the effects of temperature on these genes through dynamic post-transcriptional, post-translational and metabolic pathways. It is likely that there is no singular regulator of sex determination, but rather an accumulation of molecular events that shift the scales towards one fate over another until a threshold is reached sufficient to maintain and stabilize one pathway and repress the alternative pathway. Investigations into the mechanism underlying sex determination without sex chromosomes should focus on cellular processes that are frequently activated by multiple stimuli or can synthesize multiple inputs and drive a coordinated response. This article is part of the theme issue 'Challenging the paradigm in sex chromosome evolution: empirical and theoretical insights with a focus on vertebrates (Part I)'. [ABSTRACT FROM AUTHOR]

Amniotes possess astonishing variability in sex determination ranging from environmental sex determination (ESD) to genotypic sex determination (GSD) with highly differentiated sex chromosomes. Geckos are one of the few amniote groups with substantial variability in sex determination. What makes them special in this respect? We hypothesized that the extraordinary variability of sex determination in geckos can be explained by two alternatives: 1) unusual lability of sex determination, predicting that the current GSD systems were recently formed and are prone to turnovers; and 2) independent transitions from the ancestral ESD to later stable GSD, which assumes that geckos possessed ancestrally ESD, but once sex chromosomes emerged, they remain stable in the long term. Here, based on genomic data, we document that the differentiated ZZ/ZW sex chromosomes evolved within carphodactylid geckos independently from other gekkotan lineages and remained stable in the genera Nephrurus , Underwoodisaurus , and Saltuarius for at least 15 Myr and potentially up to 45 Myr. These results together with evidence for the stability of sex chromosomes in other gekkotan lineages support more our second hypothesis suggesting that geckos do not dramatically differ from the evolutionary transitions in sex determination observed in the majority of the amniote lineages. [ABSTRACT FROM AUTHOR]

According to the IUCN, most sea turtles fall into one of the endangered categories. Since, sea turtles, like many other reptiles, present an unusual developmental process, marked by the determination of the sex of the offspring by environmental factors, more specifically by temperature. In the temperature sex determination (TSD) system the temperature of an embryo's environment during incubation period will dictate the embryo's sex development. This developmental process, together with the complex mating and nesting behavior and the vulnerability of sea turtles to threats of a natural or anthropogenic nature, naturally lead to the study of the population dynamics of the species. For this reason, in this paper, we have developed a continuous model given by a system of three ordinary differential equations to study the dynamics of the green sea turtle population long-term, focusing the mathematical simulations on the data obtained for the nesting species of Galapagos Islands. Through the qualitative analysis of the model, the following is demonstrated: 1) The flow induced by the system is positively invariant on the region of biological interest (); and 2) The given condition on f is necessary and sufficient for the unique nontrivial equilibrium point (I) to be globally asymptotically stable in that region. When implementing the estimated values for our parameters in the numerical simulations, it was observed that indeed the population of Galapagos green sea turtles complies with the condition for which the nontrivial critical point (I) is globally asymptotically stable; that is, the asymptotic stabil- ity is maintained for any initial value within the set ν. In contrast, when altering the estimated values of the parameters so that the established condition is not met, the trivial critical point (I0) becomes globally stable, and the population falls towards extinction regardless of the values taken within the positively invariant set. [ABSTRACT FROM AUTHOR]

Abstract Background Sex-determining systems may profoundly influence the ecology, behaviour and demography of animals, yet these relationships are poorly understood. Here we investigate whether species with temperature-dependent (TSD) and genetic sex determination (GSD) differ in key demographic traits, using data from 181 species representing all major phylogenetic lineages of extant reptiles. Results We show that species with TSD exhibit significantly higher within-species variance in sex ratios than GSD species in three major life stages: birth or hatching, juvenility and adulthood. In contrast, sex differences in adult mortality rates do not differ between GSD and TSD species. However, TSD species exhibit significantly greater sex differences in maturation ages than GSD species. Conclusion These results support the recent theoretical model that evolution of TSD is facilitated by sex-specific fitness benefits of developmental temperatures due to bimaturism. Our findings suggest that different sex-determination systems are associated with different demographic characteristics that may influence population viability and social evolution.

Species that have sex determined by environmental conditions during development (i.e., environmental sex determination [ESD]) are especially vulnerable to environmental change, including altered stress levels, habitat loss, and species translocations. These factors can produce multigenerational trends in population size and eco‐evolutionary dynamics not captured by existing theory based on lifetime reproductive success (R0). Here, we extend ESD theory to use per capita growth rate r as a more appropriate measure of evolutionary success (fitness), and we demonstrate the importance of this change when males and females can differ in maturation times and when maturation times vary with local conditions (plasticity). In these cases, we show that primary and secondary sex ratios may be strongly biased; that optimal maturation times, when locally plastic, depend on the balance between mortality and growth effects; and that plasticity of maturation times can ameliorate fitness costs of increasing environmental stress. [ABSTRACT FROM AUTHOR]

Aim: Species with temperature‐dependent sex determination (TSD) are particularly vulnerable to climate change because a resultant skew in population sex ratio can have severe demographic consequences and increase vulnerability to local extinction. The Australian central bearded dragon (Pogona vitticeps) has a thermosensitive ZZ male/ZW female system of genetic sex determination (GSD). High incubation temperatures cause reversal of the ZZ genotype to a viable female phenotype. Nest temperatures in the wild are predicted to vary on a scale likely to produce heterogeneity in the occurrence of sex reversal, and so we predict that sex reversal will correlate positively with inferred incubation conditions. Location: Mainland Australia. Methods: Wild‐caught specimens of P. vitticeps vouchered in museum collections and collected during targeted field trips were genotypically and phenotypically sexed to determine the distribution of sex reversal across the species range. To determine whether environmental conditions or genetic structure can explain this distribution, we infer the incubation conditions experienced by each individual and apply a multi‐model inference approach to determine which conditions associate with sex reversal. Further, we conduct reduced representation sequencing on a subset of specimens to characterize the population structure of this broadly distributed species. Results: Here we show that sex reversal in this widespread Australian dragon lizard is spatially restricted to the eastern part of the species range. Neither climatic variables during the inferred incubation period nor geographic population genetic structure explain this disjunct distribution of sex reversal. The main source of genetic variation arose from isolation by distance across the species range. Main conclusions: We propose that local genetic adaptation in the temperature threshold for sex reversal can counteract the sex‐reversing influence of high incubation temperatures in P. vitticeps. Our study demonstrates that complex evolutionary processes need to be incorporated into modelling biological responses to future climate scenarios. [ABSTRACT FROM AUTHOR]

Species displaying temperature-dependent sex determination (TSD) are especially vulnerable to the effects of a rapidly changing global climate due to their profound sensitivity to thermal cues during development. Predicting the consequences of climate change for these species, including skewed offspring sex ratios, depends on understanding how climatic factors interface with features of maternal nesting behaviour to shape the developmental environment. Here, we measure thermal profiles in 86 nests at two geographically distinct sites in the northern and southern regions of the American alligator’s (Alligator mississippiensis) geographical range, and examine the influence of both climatic factors and maternally driven nest characteristics on nest temperature variation. Changes in daily maximum air temperatures drive annual trends in nest temperatures, while variation in individual nest temperatures is also related to local habitat factors and microclimate characteristics. Without any compensatory nesting behaviours, nest temperatures are projected to increase by 1.6–3.7°C by the year 2100, and these changes are predicted to have dramatic consequences for offspring sex ratios. Exact sex ratio outcomes vary widely depending on site and emission scenario as a function of the unique temperature-by-sex reaction norm exhibited by all crocodilians. By revealing the ecological drivers of nest temperature variation in the American alligator, this study provides important insights into the potential consequences of climate change for crocodilian species, many of which are already threatened by extinction. [ABSTRACT FROM AUTHOR]

Premise: Plant sex is usually fixed, but in rare cases, sex expression is flexible and may be influenced by environmental factors. Theory links female sex expression to better health, but manipulative work involving the experimental change of health via injury is limited, particularly in sexually plastic species. A better understanding of mechanisms influencing shifts in sex is essential to our understanding of life history theory regarding trade‐offs in sex allocation and differential mortality. Methods: We investigated the relationship between physiological stress and sex expression in sexually plastic striped maple trees (Acer pensylvanicum) by inflicting damage of various intensities (crown pruning, defoliation, and hydraulic restriction). We then monitored the sex expression of injured and control individuals for 2 years to assess the extent to which injury may cue changes in sex expression. Results: We found that severe damage such as full defoliation or severe pruning increased odds of changing sex to female and decreased odds of changing to male. In fact, no pruned male trees flowered male 2 years later, while all males in the control group flowered partially or fully male. After full defoliation, trees had 4.5 times higher odds of flowering female. Not all injury is equal; less‐severe physical trauma did not affect the frequency of sex change to femaleness. Conclusions: This work demonstrates that physical trauma in striped maple appears to exhibit a threshold effect in which only the most stressful of physiological cues instigate changes in sex expression, a phenomenon previously unknown, and that damage stress is strongly correlated with switching to femaleness. These findings have implications for population sex ratios and sustainability within an increasing stressful climate regime. [ABSTRACT FROM AUTHOR]

Premise: Environmental sex determination (ESD) is a rare sex determination system in which individuals may switch sex expression throughout their lifetimes in response to environmental factors. In sexually stable species, individuals usually bear more female flowers if the plants are larger, have greater access to limiting resources, or are in better condition. Research regarding sexually plastic species with ESD and how resources correlate with sex expression is limited. Furthermore, most research investigates resources at the population level, failing to account for resources available to individuals for growth, maintenance, or reproduction. Methods: Acer pensylvanicum is a species that is known to switch sex. Using twig samples collected during 2014–2016 in December and May, we analyzed resource status in the form of stored nonstructural carbohydrates (NSCs) and compared this with expressed sex. Results: We found that females had higher sugar concentrations than males. Furthermore, males changing expression to female had higher sugar concentrations during the prior winter than did males remaining male. We found that size was not a key predictor: neither male nor female‐flowering individuals increased NSC concentrations with size. Dying female trees had high concentrations of NSCs throughout the dying process and only manifested reduced NSCs once dead. Conclusions: This is the first study showing significant correlations between NSCs and sex expression in a plant species with ESD. These findings support the hypothesis that sex switching could be a consequence of increased resource availability and that the high female mortality of A. pensylvanicum populations is likely not a direct result of carbon starvation. [ABSTRACT FROM AUTHOR]

In mammals, including humans, males and females differ not only in physical appearance but in every cell of their body: male cells have a tiny Y-chromosome which females lack. Females instead have two X-chromosomes while males have only one. It is not universally true though, as majority of fishes, frogs, lizards and turtles have no sex chromosomes. Their sex is generally determined based on the environment (e.g. temperature) in which the eggs grow. Though, the Y-chromosome gene, Sry, triggers male development, it alone is not enough to differentiate the two sexes; orderly expression of a number of genes, generally present on the autosomes, is required to ensure differentiation of a specific gonad - testis or ovary. Individuals bearing testes become male while those having ovaries become female. Excepting the Y-chromosomal Sry, almost all other genes in this cascade are evolutionarily conserved throughout vertebrates. Mutually antagonistic interactions of the male and female pathway genes lead to the formation of the gonads that eventually determine the sex of the individual. Most disorders of sexual development occur due to mutations in any of these genes. [ABSTRACT FROM AUTHOR]

Sex determination, due to the obvious association with reproduction and Darwinian fitness, has been traditionally assumed to be a relatively conserved trait. However, research on teleost fishes has shown that this need not be the case, as these animals display a remarkable diversity in the ways that they determine sex. These different mechanisms, which include constitutive genetic mechanisms on sex chromosomes, polygenic constitutive mechanisms, environmental influences, hermaphroditism, and unisexuality have each originated numerous independent times in the teleosts. The evolutionary lability of sex determination, and the corresponding rapid rate of turn-over among different modes, makes the teleost clade an excellent model with which to test theories regarding the evolution of sex determining adaptations. Much of the plasticity in sex determination likely results from the dynamic teleost genome, and recent advances in fish genetics and genomics have revealed the role of gene and genome duplication in fostering emergence and turn-over of sex determining mechanisms.

We compiled information from the literature on the taxonomic distributions in extant teleost fishes of alternative sex-determination systems: male-heterogametic (XY) gonochorism, female-heterogametic (ZW) gonochorism, hermaphroditism, unisexuality, and environmental dependency. Then, using recently published molecular phylogenies based on whole-genomic or partial mitochondrial DNA sequences, we inferred the histories and evolutionary transitions between these reproductive modes by employing maximum parsimony and maximum likelihood methods of phylogenetic character mapping. Across a broad teleost phylogeny involving 25 taxonomic orders, a highly patchy distribution of different sex-determination mechanisms was uncovered, implying numerous transitions between alternative modes, but this heterogeneity also precluded definitive statements about ancestral states for most clades. Closer inspection of family-level and genus-level phylogenies within each of four orders further bolstered the conclusion that shifts in sex-determining modes are evolutionarily frequent and involve a variety of distinct ancestral-descendant pathways. For possible reasons discussed herein, the evolutionary lability of sex-determining modes in fishes contrasts strikingly with the evolutionary conservatism of sex determination within both mammals and birds. © 2006 The Linnean Society of London.

Sex determination of Daphnia pulex is decided by environmental conditions. We established a suitable experimental system for this study using D. pulex WTN6 strain, in which the sex of the offspring can be controlled by photoperiod. Long-day conditions induced females and short-day conditions induced males. Using this system, we previously found that methy farnesoate (MF), which is a putative innate juvenile hormone molecule in daphnids, is necessary for male sex determination and that protein kinase C (PKC) is a candidate factor of male sex determiner. In this study, we demonstrated that a PKC inhibitor [bisindolylmaleimide IV (BIM)] application strongly suppressed male offspring induction in the short-day condition. Moreover, co-treatment of BIM with MF revealed that PKC signaling acts upstream of MF signaling for male sex determination. This is the first experimental evidence that PKC is involved in the male sex determination process associated with methyl farnesoate signaling in daphnid species.

Epigenetics integrates genomic and environmental information to produce a given phenotype. Here, the model of Conserved Epigenetic Regulation of Sex (CERS) is discussed. This model is based on our knowledge on genes involved in sexual development and on epigenetic regulation of gene expression activation and silencing. This model was recently postulated to be applied to the sexual development of fish, and it states that epigenetic and gene expression patterns are more associated with the development of a particular gonadal phenotype, e.g., testis differentiation, rather than with the intrinsic or extrinsic causes that lead to the development of this phenotype. This requires the existence of genes with different epigenetic modifications, for example, changes in DNA methylation levels associated with the development of a particular sex. Focusing on DNA methylation, the identification of CpGs, the methylation of which is linked to sex, constitutes the basis for the identification of Essential Epigenetic Marks (EEM). EEMs are defined as the number and identity of informative epigenetic marks that are strictly necessary, albeit perhaps not sufficient, to bring about a specific, measurable, phenotype of interest. Here, we provide a summary of the genes where DNA methylation has been investigated so far, focusing on fish. We found that cyp19a1a and dmrt1 , two key genes for ovary and testis development, respectively, consistently show an inverse relationship between their DNA methylation and expression levels, thus following CERS predictions. However, in foxl2a , a pro-female gene, and amh , a pro-male gene, such relationship is not clear. The available data of other genes related to sexual development such as sox9 , gsdf , and amhr2 are also discussed. Next, we discuss the use of CERS to make testable predictions of how sex is epigenetically regulated and to better understand sexual development, as well as the use of EEMs as tools for the diagnosis and prognosis of sex. We argue that CERS can aid in focusing research on the epigenetic regulation of sexual development not only in fish but also in vertebrates in general, particularly in reptiles with temperature sex-determination, and can be the basis for possible practical applications including sex control in aquaculture and also in conservation biology. [ABSTRACT FROM AUTHOR]

Background and Aims The ability of individuals to change sex during their lifetime is known as environmental sex determination (ESD). This represents a unique life history trait, allowing plants to allocate resources differentially to male and female functions across lifetimes, potentially maximizing fitness in response to changing environmental or internal cues. In this study, Acer pensylvanicum , a species with an unconfirmed sex determination system, was investigated to see what patterns in sex expression existed across multiple years, if there were sex-based differences in growth and mortality, and whether this species conformed to theoretical predictions that females are larger and in better condition. Methods Patterns of sex expression were documented over 4 years in a phenotypically subdioecious A. pensylvanicum population located in New Jersey, USA, and data on size, mortality, health and growth were collected. A machine-learning algorithm known as a boosted classification tree was used to develop a model to predict the sex of a tree based on its condition, size and previous sex. Results In this study, 54 % of the trees switched sex expression during a 4-year period, with 26 % of those trees switching sex at least twice. Consistently monoecious trees could change relative sex expression by as much as 95 %. Both size and condition were influential in predicting sex, with condition exerting three times more relative influence than size on expressed sex. Contrary to theoretical predictions, the model showed that full female sex expression did not increase with size. Healthy trees were more likely to be male; predicted female sex expression increased with deteriorating health. Growth rate negatively correlated with multiple years of female sex expression. Populations maintained similar male-skewed sex ratios across years and locations and may result from differential mortality: 75 % of dead trees flowered female immediately before death. Conclusions This study shows conclusively that A. pensylvanicum exhibits ESD and that femaleness correlates with decreased health, in contrast to prevailing theory. The mortality findings advance our understanding of puzzling non-equilibrium sex ratios and life history trade-offs resulting from male and female sex expression. [ABSTRACT FROM AUTHOR]

Fisherian sex-ratio theory predicts sexual species should have a balanced primary sex ratio. However, organisms with environmental sex determination (ESD) are particularly vulnerable to experiencing skewed sex ratios when environmental conditions vary. Theoretical work has modeled sex-ratio dynamics for animals with ESD with regard to 2 traits predicted to be responsive to sex-ratio selection: 1) maternal oviposition behavior and 2) sensitivity of embryonic sex determination to environmental conditions, and much research has since focused on how these traits influence offspring sex ratios. However, relatively few studies have provided estimates of univariate quantitative genetic parameters for these 2 traits, and the existence of phenotypic or genetic covariances among these traits has not been assessed. Here, we leverage studies on 3 species of reptiles (2 turtle species and a lizard) with temperature-dependent sex determination (TSD) to assess phenotypic covariances between measures of maternal oviposition behavior and thermal sensitivity of the sex-determining pathway. These studies quantified maternal behaviors that relate to nest temperature and sex ratio of offspring incubated under controlled conditions. A positive covariance between these traits would enhance the efficiency of sex-ratio selection when primary sex ratio is unbalanced. However, we detected no such covariance between measures of these categories of traits in the 3 study species. These results suggest that maternal oviposition behavior and thermal sensitivity of sex determination in embryos might evolve independently. Such information is critical to understand how animals with TSD will respond to rapidly changing environments that induce sex-ratio selection. [ABSTRACT FROM AUTHOR]

Daphnia spp., a keystone genus in freshwater lentic habitats, are subject to environmental sex determination wherein environmental conditions dictate offspring sex and whether they reproduce asexually or sexually. The introduction of males into a population denotes the first step in the switch from asexual parthenogenetic reproduction to sexual reproduction. We tested the hypothesis that photoperiod and temperature co-regulate male sex determination and that these environmental stimuli would activate elements of the male sex determination signaling cascade. The results revealed that photoperiod was a critical cue in creating permissive conditions for male production. Further, under photoperiod-induced permissive conditions, male sex determination was temperature dependent. The two daphnid species evaluated, Daphnia pulex and Daphnia magna, exhibited different temperature dependencies. Daphnia pulex produced fewer males with increasing temperatures between 16 and 22°C, and D. magna exhibited the opposite trend. We found consistent expression patterns of key genes along the male sex-determining signaling pathway in D. pulex independent of environmental stimuli. mRNA levels for the enzyme responsible for synthesis of the male sex-determining hormone, methyl farnesoate, were elevated early in the reproductive cycle, followed by increased mRNA levels of the methyl farnesoate receptor subunits Met and SRC. Environmental conditions that stimulated male offspring production significantly increased Met mRNA levels. The results indicate that male sex determination in daphnids is under the permissive control of photoperiod and the regulatory control of temperature. Further, these environmental cues may stimulate male sex determination by increasing levels of the Met subunit of the methyl farnesoate receptor. [ABSTRACT FROM AUTHOR]

Spatial structure has been shown to favor female‐biased sex allocation, but current theory fails to explain male biases seen in many taxa, particularly those with environmental sex determination (ESD). We present a theory and accompanying individual‐based simulation model that demonstrates how population structure leads to male‐biased population sex ratios under ESD. Our simulations agree with earlier work showing that the high productivity of female‐producing habitats creates a net influx of sex‐determining alleles into male‐producing habitats, causing larger sex ratio biases, and lower productivity in male‐producing environments (Harts et al. 2014). In contrast to previous findings, we show that male‐biasing habitats disproportionately impact the global sex ratio, resulting in stable male‐biased population sex ratios under ESD. The failure to detect a male bias in earlier work can be attributed to small subpopulation sizes leading to local mate competition, a condition unlikely to be met in most ESD systems. Simulations revealed that consistent male biases are expected over a wide range of population structures, environmental conditions, and genetic architectures of sex determination, with male excesses as large as 30 percent under some conditions. Given the ubiquity of genetic structure in natural populations, we predict that modest, enduring male biased allocation should be common in ESD species, a pattern consistent with reviews of ESD sex ratios. [ABSTRACT FROM AUTHOR]

• A time-course gonadal RNA-seq was conducted using Mauremys reevesii. • Sexual differentiation genes in males were activated at an earlier stage than the ones in females. • Turtle-alligator comparative analysis revealed novel candidate TSD genes. The mode of sex determination in vertebrates can be categorized as genotypic or environmental. In the case of genotypic sex determination (GSD), the sexual fate of an organism is determined by the chromosome composition with some having dominant genes, named sex-determining genes, that drive the sex phenotypes. By contrast, many reptiles exhibit environmental sex determination (ESD), whereby environmental stimuli drive sex determination, and most notably temperature. To date, temperature-dependent sex determination (TSD) has been found in most turtles, some lizards, and all crocodylians, but commonalities in the controlling processes are not well established. Recent innovative sequencing technology has enabled investigations into gonadal transcriptomic profiles during temperature-sensitive periods (TSP) in various TSD species which can help elucidate the controlling mechanisms. In this study, we conducted a time-course analysis of the gonadal transcriptome during the male-producing temperature (26℃) of the Reeve's turtle (Chinese three-keeled pond turtle) Mauremys reevesii. We then compared the transcriptome profiles for this turtle species during the TSP with that for the American alligator Alligator mississippiensis to identify conserved reptilian TSD-related genes. Our transcriptome-based findings provide an opportunity to retrieve the candidate molecular cues that are activated during TSP and compare these target responses between TSD and GSD turtle species, and between TSD species. [ABSTRACT FROM AUTHOR]

Bluehead wrasses (Thalassoma bifasciatum) follow a socially controlled mechanism of sex determination. A socially dominant initial-phase (IP) female is able to transform into a new terminal-phase (TP) male if the resident TP male is no longer present. TP males display an elaborate array of courtship behaviors, including both color changes and motor behaviors. Little is known concerning the neural circuits that control male-typical courtship behaviors. This study used glutamate iontophoresis to identify regions that may be involved in courtship. Stimulation of the following brain regions elicited diverse types of color change responses, many of which appear similar to courtship color changes: the ventral telencephalon (supracommissural nucleus of the ventral telencephalon [Vs], lateral nucleus of the ventral telencephalon [Vl], ventral nucleus of the ventral telencephalon [Vv], and dorsal nucleus of the ventral telencephalon [Vd]), parts of the preoptic area (NPOmg and NPOpc), entopeduncular nucleus, habenular nucleus, and pretectal nuclei (PSi and PSm). Stimulation of two regions in the posterior thalamus (central posterior thalamic [CP] and dorsal posterior thalamic [DP]) caused movements of the pectoral fins that are similar to courtship fluttering and vibrations. Furthermore, these responses were elicited in female IP fish, indicating that circuits for sexual behaviors typical of TP males exist in females. Immunohistochemistry results revealed regions that are more active in fish that are not courting: interpeduncular nucleus, red nucleus, and ventrolateral thalamus (VL). Taken together, we propose that the telencephalic-habenular-interpeduncular pathway plays an important role in controlling and regulating courtship behaviors in TP males; in this model, in response to telencephalic input, the habenular nucleus inhibits the interpeduncular nucleus, thereby dis-inhibiting forebrain regions and promoting the expression of courtship behaviors. • Microiontophoretic stimulation of the brain for the first time in a sex-changing fish • Stimulation elicited coloration and motor behaviors typical of male courtship • A brain circuit for controlling male-typical courtship displays is presented • The interpeduncular nucleus is proposed to gate male-typical courtship behaviors Bluehead wrasses exhibit socially controlled sex determination, in which females can transform to males. Luong et al. present a neuronal circuit for controlling male-typical courtship behaviors and show that it exists in females. The interpeduncular nucleus is postulated to gate the expression of the male behavioral sex phenotype. [ABSTRACT FROM AUTHOR]

Although socially controlled sex transformation in fishes is well established, the underlying mechanisms are not well understood. Particularly enigmatic is behavioural transformation, in which fish can rapidly switch from exhibiting female to male-typical courtship behaviours following removal of 'supermales'. Bluehead wrasses are a model system for investigating environmental control of sex determination, particularly the social control of sex transformation. Here, we show that the onset of this behavioural transformation was delayed in females that occupied low-ranking positions in the female dominance hierarchy. We also establish that expression of maletypical courtship behaviours in competent initial-phase (IP) females is facultative and gated by the presence of terminal-phase (TP) males. Dominant females displayed reliable TP male-typical courtship behaviours within approximately 2 days of the removal of a TP male; immediately following reintroduction of the TP male, however, females reverted back to female-typical behaviours. These results demonstrate a remarkable plasticity of sexual behaviour and support a 'priming/gating' hypothesis for the control of behavioural transformation in bluehead wrasses. [ABSTRACT FROM AUTHOR]

Abstract: The pejerrey possesses a genotypic sex determination system driven by the amhy gene and yet shows marked temperature‐dependent sex determination. Sex‐reversed XY females have been found in a naturally breeding population established in Lake Kasumigaura, Japan. These females could mate with normal XY males and generate YY “supermale” individuals that, if viable and fertile, would sire only genotypic male offspring. This study was conducted to verify the viability, gender, and fertility of YY pejerrey and to develop a molecular method for their identification. Production of YY fish was attempted by crossing a thermally sex‐reversed XY female and an XY male, and rearing the progeny until sexual maturation. To identify the presumable YY individuals, we first conducted a PCR analysis using amhy‐specific primers to screen only amhy‐positive (XY and YY) fish. This screening showed that 60.6% of the progeny was amhy‐positive, which suggested the presence of YY fish. We then conducted a second screening by qPCR in order to identify the individuals with two amhy copies in their genome. This screening revealed 13 individuals, all males, with values twice higher than the other 30 amhy‐positive fishes, suggesting they have a YY complement. This assumption as well as the viability, fertility, and “supermale” nature of these individuals was confirmed in progeny tests with XX females that yielded 100% amhy‐positive offspring. These results demonstrate that qPCR can obviate progeny test as a means to identify the genotypic sex and therefore may be useful for the survey of all three possible genotypes in wild populations. [ABSTRACT FROM AUTHOR]