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We seek to shed light on the main gene regulatory networks involved in the whole sex change process by observing the cascade of expression changes, their specific patterns and their temporal autocorrelations free fish sex a transcriptomic approach.

Our time-series expression experiment provides information regarding the complete set of genes that is activated, as well free realistic sex games the dynamics and interactions between these genes. Our analysis is focused on the two main organs involved in sexual development and reproduction, the brain and gonads Gonads were histologically analyzed in twelve sex-changing individuals, two functional males and six functional females that were used as controls for the completion free fish sex sex change Table 1.

During the last steps of the transition days 40—50 after removal of the original femalesindividuals possessed highly degenerated and reduced testes, but their oocytes, although increased in size, remained in the primary-growth stage; i. All functional females were mature with vitellogenic oocytes in spawning-capable phase.

Interestingly, one fish, CF10M, who belonged to a three-member family when it was originally tagged but was left alone in the free fish sex when the female was removed since free fish sex third member had disappeared, did not undergo the expected male-to-female sex change. Individual code, time lapse days between the removal of the female and the collection of the partner undergoing sex changepercentage of male free fish sex female tissue obtained by histological analysis of the individual gonads, assigned sex-change category, corresponding individual label.

For the correlation analysis, the index of sex change of each individual as measured by the percentage of female tissue in the gonad was used. For the pairwise differential expression analysis, individuals were african love dating site into five categories according to the percentage of female tissue in the gonad Table 1: One of these groups TF has only two individuals, while it should ideally have three or more, which may impair the power for detecting differences in expression.

However, free fish sex risk of impairment is reduced due to the statistical approach taken and the free fish sex that the selection criteria of the genes is based not solely on pairwise, but also on correlation analysis.

We may take it for granted that the sex of an animal is established at birth and doesn't change. However, about species of fish change sex. In like manner, ingestion of fish produced by sex reversal does not therefore harm . of MT help ensure that tilapia is free of MT before fish reach the consumer. These changes in the relative neuronal activity in the male fish's brain are sex change in fish, using the Red Sea endemic species of clownfish, were weighed and placed in RNA-free microcentrifuge tubes containing 1.

free fish sex Evaluation of the genome for completeness identified Analysis of the reconstructed transcripts using TransDecoder revealed free fish sex total of 96, contigs potentially containing candidate protein-coding regions. Comparisons were conducted pick up women craigslist for the brain and gonadal samples, since the hierarchical clustering performed with the sample-to-sample free fish sex revealed two well-differentiated groups Supplementary Fig.

Clustering analysis of each of the tissues separately shows that expression profiles from the brain samples display higher similarity than those from the gonad samples Supplementary Fig. Therefore, more contigs were found to be female-biased than male-biased in the brain. The scatterplot of the correlations against the dispersion in expression shows that the distribution of correlation coefficients was similar on both sides of the spectrum, female and male, with largest values of 0.

Significantly correlated contigs displayed small dispersion, especially the contigs up-regulated at the end of the sex-change free fish sex, i. The largest differences in expression were observed in contigs not correlated to the index of sex change grey dots in Fig.

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Contigs are plotted by their correlation with the index of sex-change x-axis and the dispersion in expression among individuals y-axis, only values above 1 are plotted. Significant correlations are highlighted in blue for negative and in red for positive coefficients, indicating wex in males and females, respectively. Examples from selected contigs marked with a married Bottineau women regarding its free fish sex in the plot are highlighted.

In the brain, the main changes in expression occurred therefore during transitional stages, an observation that also explains the paucity of correlated seex found in the previous analysis. Five categories, based on the percentage of female tissue in the gonad, were used in the comparisons, with labels corresponding to: Thus, more free fish sex were found to be male-biased, the opposite trend than in the brain samples. The scatterplot of the correlations against the dispersion in expression Fig.

Significant positive correlations were generally smaller; the largest value was 0. Eighty-three negatively correlated contigs had coefficients above 0. Also the contigs that were up-regulated in females showed a larger range in differential free fish sex.

Strong correlation indicates a steady increase or decrease in expression, but often with small differences among individuals Fig. On the contrary there were contigs with large differences in expression, some of them not correlated with the free fish sex of sex change at the individual level grey dots in Fig.

The plot of the free fish sex for the gonads was similar to that for plenty of fish sex brain although generally there were larger differences among individuals. Contigs are plotted by their correlation with the index of sex change x-axis and the dispersion in expression among individuals y-axis, only values above 1 are plotted. The majority were negatively correlated while 64 were positively correlated.

The majority of the significant DECs were found in comparisons involving mature females Table 2indicating that expression patterns during the last phase of the sex change are clearly distinct from all the previous stages. The PCA and cluster analysis performed on the selected contigs from the brain revealed a clear clustering of individuals into two well-defined groups: The difference between transitional males and transitional females is considerably larger than differences between any groups in the non-transitional cluster.

Although males and mature females form two distinct groups, they cluster together with immature females and are clearly separated from transitional males and females. Hierarchical clustering of the transcripts resulted in three main clusters Free fish sex.

Two of the clusters grouped transcripts that were either expressed in all the reproductive stages tested BG3 or were slightly down-regulated in transitional fish BG2.

The expression of selected genes for each south korean dating sites is free fish sex in Fig. Expression patterns include up-regulation in males popdc3, rab41, nt5marked free fish sex in transitional stages hdgfrp2tenm2, foxp4, sox6up-regulation in transitional stages tspan8 and up-regulation in females adamts16, cyp19a1b, f13a, phactr4. Gene expression changes were investigated free fish sex A Principal Component Analyses performed on normalized RNA-Seq data of a selected set of transcripts from the brain left and transcripts from the gonad right.

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B Free fish sex dendrogram clustering on the same data. The clustering of transcripts results in three brain: BG1 to BG3 and five gonad: GG1 to GG5 clearly differentiated expression profiles.

M — males dark blue ; TM — transitional males light blue ; TF — transitional females; FI pink — immature females purple ; and FM — mature females brown. The description of each gene is found horny virgins French Polynesia Table 3and the free fish sex statistics in Supplementary Tables S2 and S3. Clustering of the selected contigs from the gonads resulted in a completely fisu arrangement Fig. The PCA and cluster analysis resulted in three major clusters.

Males and transitional males clustered together forming a separate group from free fish sex female stages.

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Within the female stages, two other distinct free fish sex can be observed: Hierarchical clustering of transcripts revealed five clusters Fig. In between, a group of transcripts that, while still up-regulated in all male categories, were also expressed in free fish sex females TF, FI, GG2 and black female swingers females GG3.

Another group of transcripts were found not to be expressed in males, but showed signs of up-regulation in transitional fish and females GG4. These clear expression patterns allow the identification shemale woods genes that likely play key roles during each stage of sex change.

Expression profiles of a typical gene from each group are shown in Fig.

We also summarise the major approaches used to control sex in fish and discuss their application in commercially important species. These changes in the relative neuronal activity in the male fish's brain are sex change in fish, using the Red Sea endemic species of clownfish, were weighed and placed in RNA-free microcentrifuge tubes containing 1. Sex for fish sometimes referred to as "fish for sex" is a phenomenon in which female traders . This simple advancement allowed women to be free from sexual exploitation in order to secure their fish supply. No sex for fish aimed to change the.

Out of the 47, contigs, Free fish sex E-value distribution of the top hits revealed that Only two were shared between these two sets and From the gonadal transcriptome a total of contigs were selected Supplementary Fig. The vast majority To aid the overall functional interpretation of the response to sex change, we fizh genes to groups according to their gene ontology GO free fish sex for biological quizzes for single women.

At the brain level, hierarchical clustering revealed similar patterns of expression between two well-supported groups, transitional stages transitional males and females se non-transitional stages males, immature and mature females Fig. Analysis of differentially expressed contigs that were specifically up-regulated in transitional stages resulted tish nine over-represented GO terms that had significant GO enrichment for regulation agency dating line free fish sex G-protein-coupled receptor protein signaling pathway and activation of meiosis involved in free fish sex development Supplementary Fig.

S4Supplementary Table S4.

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At the gonadal level, our hierarchical clustering revealed three well-supported groups, with male stages forming a clearly separated cluster from the two female groups Fig. GO enrichment analysis was performed on differentially expressed contigs which showed negative and positive correlation to the index of sex change for details on selected contigs, see Supplementary Table S3. Their putative functions include reproduction usa male models, interestingly, eye development lens development in camera-type eye, iris morphogenesis, lens morphogenesis, embryonic retina morphogenesis in camera-type eyesuggesting free fish sex possibility of novel additional functions of these genes related to the sex change mechanism Supplementary Fig.

Their putative functions include regulation of hormone levels and steroid metabolic free fish sex Supplementary Fig. To confirm gay skype topix forum expression changes detected by RNA-seq, we performed qPCR on six genes exhibiting broad expression ranges: Expression patterns free fish sex consistent between both methods Supplementary Fig.

A few recent studies have analyzed the transcriptome of non-model fish species to shed light on sex differentiation and sex-specific gene expression.

Remarkable differences in gene expression profiles of gonads and brain have been ladies seeking sex Orogrande New Mexico between males and females. Such studies include hermaphrodite species e. Thalassoma bifasciatum 18 and gonochoristic species displaying a dominance hierarchy that is similar to that of clownfish e.

African cichlids However, little previous free fish sex has examined differences in gene expression across a dynamic life history process. This study addresses the transcriptome analysis of sex change in a species with a strong social hierarchy. Our study is the first to highlight the rapid and complex genomic responses of the brain and gonad associated with sex change in the natural environment. Furthermore, our work examines relative gene expression across multiple groups—rather than just two contrasting conditions—allowing us to explore the differences in expression patterns, including those of intermediate stages, during the whole process.

This response is subsequently transmitted to the gonads where differential expression and histological changes are clearly observed after three to four weeks. The main transcriptional response driving sex change at the brain level was found to be completed 30 days after removal of the original female, while differential expression is still detected in the gonads 50 days after removal of the original female, although sex change is completed when males become immature females.

We hypothesize that changes in brain expression are transmitted to free fish sex gonad through the hypothalamic-pituitary-gonadal axis.

Receptors on the gonadal tissue receive the hormonal signals resorbing the testicular tissue and developing the ovary to complete the gonadal sex change. Our study has revealed an important number of genes involved in these processes, some of well-known and several novel genes potentially playing a role in sex change, both at the brain and gonadal levels summarized in Table 3expression profiles are shown in Supplementary Fig.

Our analysis shows that functional males day 0 and males whose female partners were removed 5 and 11 days earlier display very similar expression profiles, while no histological signal of sex change was observed at the gonadal level. Reorganization of gonadal tissue is costly, not only energetically but in terms of the time spent on sex change, a period during which the fish are dex able to reproduce.

In the case of A. It has aex shown in other Amphiprion species that staying male and re-pairing with a new larger mate who has emigrated from another free fish sex is a more affordable mate acquisition strategy 9. Thus, the disappearance of the female would seex trigger the sex change process at the brain level until it becomes clear to the male that moving up in the hierarchy is beneficial given the lasting free fish sex of free fish sex female. Consistent with this hypothesis, we found the first clear changes in brain expression profiles of transitional males days 15 to 30 after original female removal compared to males 0—11 days.

These individuals showed a marked down-regulation in differentially expressed genes that was maintained also in transitional females. On the other hand, females displayed very similar expression patterns to males, indicating subtle sex baoding Baoding free sex porno massage in overall gene expression between both mature stages.

Accordingly, most differentially expressed genes were detected in transitional stages and only twelve showed a significant correlation to the index of sex change. To tease out the molecular basis of sex change in clownfish at the brain level, we examined differences in neural expression of both candidate and highly differentially expressed novel genes and assessed their gene expression patterns during the whole sex change process. Free fish sex very well-known gene that plays an important fiwh in sex change in teleosts both at the gonad and free fish sex level is cyp19a1 also known as aromatase or ParomA.

In non-mammalian blind dating site, cyp19a1 is a key steroidogenic free fish sex operating free fish sex the female pathway and converting androgens into estrogens 20hence controlling the balance of sex steroids.

This gene followed a clear increasing trend of expression values free fish sex the female categories, resulting in significant correlation and free fish sex differential expression between male stages and mature females gish our experiment.

Two possible candidates to be involved in the mechanism of sex change showed a significant down-regulation during transitional stages: Sox6 is a transcription factor that has been found to be involved in the sex developmental pathway of vertebrates by regulating spermatogenesis FoxP4 pittsburgh seniors swingers.

Swinging. a member of the FoxP gene family of transcription regulators whose functions and targets have yet to be determined The fox genes free bbw butts important roles free fish sex various biological ifsh, including sexual development.

A transcriptomic analysis revealed very high expression levels of foxp4 in the gonads of Nile Tilapia 23whereas sex-dependent changes dree expression gree foxp4 affecting only male mammals were found We propose that cyp19a1b plays a central role in the mechanism of sex change in the brain of A.

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Solid arrows represent the known regulations free fish sex genes involved in sexual differentiation in A. Some genes encoding key neural regulators that free fish sex been shown to play major roles during sexual development and maturation in fish at the brain level such as gonadotropin-releasing hormone GnRHarginine-vasotocin AVTKisspeptin Feee or steroid hormones and their receptors, were not detected in our analysis. Note, however, that our study examined free fish sex brain as a whole, whereas it is well established that sex-specific flsh in gene expression are present in fish when only free fish sex brain regions are considered GnRH, AVT and Kiss have been shown to be involved during sex change in other Amphiprion species, but all the studies were restricted to the hypothalamus 2627 Similarly to our results, no significant differences in the expression levels of estrogen receptors were detected in the forebrain of bluehead wrasses Our analysis revealed a considerably higher number of transcripts showing both significant correlation to the index of sex change and significant differential expression in the gonads of A.

Cluster analysis of these differentially expressed transcripts revealed three well-differentiated groups: The main changes in expression related to sex change at the gonadal level started 24 days after the removal of the original female and were observed until the end of the experiment day Histological analysis indicated that male tissue was completely resorbed after 50 days sed therefore the sex change process was completed, although none of the new females were spawning capable yet, as opposed to the original mature females.

Moreover, our study revealed a higher number of over-expressed transcripts at the gonadal level during the first stages of the sex change, when the transitional individuals were still acting as males, than at the end free fish sex the transition to becoming females. This is in accordance with the fact of A. Up-regulation or expression of a higher number of genes is required for the maintenance of the male status of the gonads ambisexual gonad when compared to that required for the female status ovarian-only.

This male-bias has been observed in thailand guys fish free fish sex at the gonadal level 1929 Sexual free fish sex platonic friends 24 Modesto 24 teleosts has been more extensively studied at the gonad than at the brain level. It is a major process that takes place after sex determination and involves the actual free fish sex of testes or ovaries from the undifferentiated gonad.

Gene regulatory cascades consisting of a complex network feee transcription factor interactions and signaling molecules control gonad differentiation and identity maintenance In contrast, free fish sex components operating during these processes appear to be evolutionarily more conserved To dissect the molecular basis of sex change at the gonadal level of A.

Among the free fish sex genes, the cyp19a1a ssx is one of the best-studied genes known to play an important role in sex change in teleosts, as mentioned earlier.

In our experiment, cyp19a1a showed a strong up-regulation in all female categories, similar to the trend found in the brain, resulting in significant correlation and significant differential expression free fish sex male and female stages.

At the gonadal level, cyp19a1a is considered essential for ovarian differentiation and development in fish 34amphibians and reptiles It has been widely characterized in gonads of sex-changing fishes, including the yellowtail clownfish 16 male seeking male in chennai, and its pivotal role in gonadal sex change in teleost fish has been extensively studied The Foxl2 transcription factor also plays a decisive role in the ovarian differentiation process in various vertebrate species, including fishes.

It is involved in the regulation of estrogen synthesis via direct modulation of aromatase expression and possibly the entire steroidogenic pathway This protein is free fish sex required to maintain the identity free fish sex ovarian granulosa cells In our experiment, foxl2 was markedly up-regulated in mature females, displaying a expression profile parallel to cyp19a1.

A positive free fish sex loop regulating these two genes in fish 36 has been suggested, since foxl2 regulates cyp19a1while estrogens up-regulate foxl2 Besides the cyp19a1a gene, another target of foxl2 in the ovaries is the steroidogenic acute regulatory protein Star whose transcriptional activity is repressed by Foxl2 in mice In our experiment, a transcript encoding Star was markedly up-regulated in mature females, resulting in differential expression compared to all the other categories.

This protein controls the rate-limiting step in steroid hormone synthesis and is required for normal ovarian steroid production in mice and humans free fish sex Similarly, hsd17b1 was also found to be significantly up-regulated in mature females, showing significant differential expression compared to all other categories and significant correlation to the index of sex change.

All these hungaryan girls are key players that regulate ovarian steroidogenesis. On the other hand, our analysis revealed three genes known to have a clear role in sex determination, testicular differentiation and spermatogenesis in teleosts: These three genes were strongly up-regulated in males compared to females in our study, showing a strong negative correlation to the index of sex change. Recent work has revealed that Sox8 is an important determinant for the maintenance of testis cell identity any Gurdon Arkansas aged dudes up horny mice 45 as well as a critical regulator of adult Sertoli cell function and male fertility Sox8 regulates the expression of amh in mice 47which encodes a member of the transforming growth-factor-beta gene family that mediates male sexual differentiation and development.

In our study, free fish sex different transcripts encoding Amh were detected amh1 free fish sex a mh2both displaying a steady decrease in expression values throughout the sex change. It has been proposed that Amh is an important hormone involved in the sex change of the protandrous black porgy In zebrafish, estrogen-induced alterations had a suppressive effect on the expression of amh and dmrt1 and this older women dating younger men statistics found to be associated with a disruption in male gonadal sex development On the other hand, dmrt1 had a marked correlation with the index of sex change with a steady decrease in expression values.

The dmrt1 gene family has been free fish sex to be involved in sexual differentiation in organisms as phylogenetically divergent as corals, Caenorhabditis elegansDrosophilafishes, frogs, birds and mammals In fishes, either male-restricted or strong male-biased expression has been reported free fish sex at least twenty gonochoristic species see go fast and green hotel get together 50 and references.

In sequential hermaphrodites, its expression has been shown to parallel the development and regression of the testis, both in protandrous 51 and protogynous species A recent study gangbang party club highlighted the important role of Dmrt1 and Foxl2 in maintaining sexual cell identity.

The loss of Dmrt1 in male adult mice can cause sexual transdifferentiation of testicular Sertoli cells to ovarian granulosa cells. Similarly, dmrt1 expression in the ovary of mice, even in the absence of the testis-determining gene sox8silenced foxl2 and reprogrammed adult granulosa cells into Sertoli-like cells Based on our results, we postulate that cyp19a1 and foxl2 play a pivotal role in the activation of the female free fish sex driving the gonadal transformation from testis to ovary during sex change in A.

All these genes have shown consistent expression profiles in previous studies of other sex-changing species, including protandrous sharpsnout seabream 29Asian seabass 54black porgy 55 and protogynous fish bluehead wrasse The proposed mechanism involves a complex regulatory loop combining transcriptional regulation with steroid hormonal activity Fig.

Similarly to the brain, some well-known gene candidates involved in the regulatory network of sexual development were absent from our analysis.

Wnt free fish sex has been implicated in free fish sex reproductive role in teleost, promoting ovarian differentiation through the upregulation of free fish sex aromatase Our data did not reveal significant differences in expression throughout sex change of key members of the Wnt-signaling cascade, such as wnt2, wnt4, ctnnb1 beta-catenin 1 free fish sex, or rspo However, a female-specific function of the Wnt pathway does not seem to be a general pattern in fishes In addition, some key genes implicated with testicular differentiation in other fish species like wt1, gsdf or nr5a2 showed no sex-biased expression in our study.

These genes are likely to represent early testis genes 15859 and might therefore not be involved in sex change in this protandrous species. However, we cannot exclude the possibility of having missed these genes in the annotation process why do men only want to have sex with me free fish sex a mutation, sequencing free fish sex or simply due to the limited genomic information available in the public databases for the non-model organism under study.

Several genes with a potential role in the sex change process were found to be down-regulated during free fish sex transitional stages, among them the gene encoding teneurin-2 tenm2. Teneurins are a unique family of transmembrane proteins conserved from C. Interestingly, a recent study proposed that Tenm2 may have an effect on the sexual maturity of chickens since the expression of genes in the nervous system can influence the age when chickens lay their first egg The up-regulation of teneurin-2 expression observed in our experiment in mature stages both mature males and females may indicate that it plays a role in the mature gonads in A.

The free fish sex encoding the hepatoma-derived growth-factor-related protein 2 hdgfrp2 was also down-regulated during transitional stages, resulting in significant differential expression between transitional females and all the other categories. This gene is expressed in a wide range of tissues suggesting a function in cells of the central nervous system, but very little is known about the function of Hdgfrp2 Free fish sex, over-expression free fish sex hdgfrp2 was detected in male free fish sex tissue of E.

The transcript encoding coagulation factor XIII A chain-like f13a showed the highest positive correlation to the index of sex change in our study and strong up-regulation in mature females, resulting in significant differential expression compared to all the other categories. Female-biased expression of f13 was also detected in a transcriptomic analysis of the guppy brain F13 is best known for its role in fibrin stabilization during coagulation, but recent advances have proposed a novel function in neuronal regeneration in sex women xxx Owa Aledimma Similarly, the transcript encoding the disintegrin and metalloproteinase with thrombospondin motifs 16 adamts16 showed a strong correlation to the index of sex change.

Adamts16 is a member of a family of metalloproteinases, expressed at high levels in free fish sex brain and ovary in humans A recent study has revealed that it is essential for normal development of the testis in rats Nt5 represents the major enzyme responsible for the formation of extracellular adenosine, involved in the regulation of neurotransmitter release Gender-dependent changes have been shown for nt5 activity and nucleoside transport in rodents 68 while gender differences in nucleoside levels in the brain have been detected in humans Popdc3 belongs to a family that codes for membrane proteins that are expressed in neuronal cells 70 of unknown function, while Ras proteins are involved in signal transduction pathways of several cellular functions, as cell proliferation, differentiation and apoptosis The expression of tetraspanin 8 tspan8 was significantly up regulated in transitional females in our study.

Free fish sex is known to mediate signal transduction events that contribute to the regulation of cell development, activation, growth and motility Interestingly, genetic ablation of tspan8 in mice resulted in a reduction in the body weight of males fed a normal diet, suggesting that it may have free fish sex role in the neural control of metabolism This last function might explain the up-regulation observed in the transitional stages in our study, since sex change implies the rapid growth of the sex-changing individual who is freed from the strict social constraints previously imposed by the dominant female pair mate.

Finally, the transcript encoding phosphatase and actin regulator 4A phactr4a showed significant differential expression between male and female stages and positive correlation to the index of sex change in our experiment. Phactr4a is critical during neurulation and eye development Its role in the brain of teleosts remains unexplored. Several other known candidate genes that may play a role free fish sex gonadal development were detected in our study.

Free fish sex

A recent study has demonstrated direct regulation of Ras proteins by free fish sex receptor 75 while their activation in granulosa cells has been fjsh to be crucial for normal follicle development Similarly, several genes with immune-related function were fosh in A.

It has been proposed that immune molecules may play an essential role in the orchestration of gametogenesis and the maintenance of gonad tissue homeostasis vree fish Rish study is important for understanding how genes get turned off and on during free fish sex in all animals including humansand how the environment can influence this process. Co-lead author PhD candidate Oscar Ortega-Recalde, also sex dating in Breesport the University of Otago, said the amazing transformation also appears possible through changes in cellular "memory.

Our study is free fish sex because it shows that sex change involves profound changes in these free fish sex marks," Mr Ortega-Recalde said.

La Trobe's Professor Jenny Graves said the project links to studies of sex reversal in Australian dragon lizards that she is collaborating on with researchers at the University of Canberra. Materials provided by La Free fish sex University. Content may be edited for style and length. Science News. Story Source: Journal Reference: Erica Dex. Lamm, Kim M. Rutherford, Hugh Cross, Michael A. Black, Olga Kardailsky, Jennifer A.

Marshall Graves, Timothy A. OpenLearn works with other organisations by providing free courses and resources that support our mission of opening up educational opportunities to more people in more places. All rights reserved. The Open University is authorised and regulated by the Financial Conduct Authority in relation to its secondary activity of fisg broking. Skip to content Study with The Open University. Search for free courses, interactives, videos and more!

Free Learning from The Open University. Featured content. Free courses. All content. Why do some fish change sex? Updated Thursday 26th October How does sex change occur?

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Copyright information. Publication details Originally published: Thursday, 26th October Leave flsh comment. Sign in or free fish sex your OpenLearn account to join the discussion.