Sexual Development

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ENVOY Is a Major Determinant in Regulation of Sexual Development in Hypocrea jecorina (Trichoderma reesei)

ENVOY Is a Major Determinant in Regulation of Sexual Development in Hypocrea jecorina (Trichoderma reesei)

Although BLR1 and BLR2 can be expected to act as a complex, our data indicate that they also have individual functions. The effect in mutants lacking blr2 is in most cases stronger than that in mutants lacking blr1. In this respect a constitutive expression of BLR2 and thus higher abundance than BLR1, as suggested for their homologues in N. crassa, could be assumed. Together with the predicted importance of WC-2 for the interaction of WC-1 with FRQ (13), this could explain the more severe effect of dele- tion of blr2 for regulation of sexual development. In N. crassa, neither the pheromone receptor genes nor the pheromone pre- cursor genes or mating type genes are targets of the White Collar complex (59). Moreover, these genes also do not seem to be light induced in N. crassa (7). Consequently, we assume that in H. jeco- rina the homologues of these genes are likely to be indirect targets of BLR1 and BLR2. A flat hierarchical network (59) of transcrip- tion factors, as suggested for N. crassa, could be targeted by BLR1 and BLR2 in H. jecorina. However, since photoreceptors were shown to have a role in regulating carbon sensing and utilization (17), it cannot be excluded that altered cultivation conditions would also show binding to promoters of genes involved in sexual development.
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PRECOCIOUS GROWTH OF SEXUAL HAIR WITHOUT OTHER SECONDARY SEXUAL DEVELOPMENT; "PREMATURE PUBARCHE," A CONSTITUTIONAL VARIATION OF ADOLESCENCE

PRECOCIOUS GROWTH OF SEXUAL HAIR WITHOUT OTHER SECONDARY SEXUAL DEVELOPMENT; "PREMATURE PUBARCHE," A CONSTITUTIONAL VARIATION OF ADOLESCENCE

Variation in the pattern of sexual development described could be due to ( I) increased sensitivity of sexual hair follicles to normal levels of androgen present during. the preadolescen[r]

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Effect of the transformer 2 gene of Anastrepha on the somatic sexual development of Drosophila

Effect of the transformer 2 gene of Anastrepha on the somatic sexual development of Drosophila

The characterisation of sex determination genes in Drosophila Melanogaster has shown that the product of a gene controls the sex-specific splicing of the pre-mRNA from the downstream gene in the genetic cascade (reviewed in Sánchez et al., 2005). Sex-lethal (Sxl) is at the top of this cascade and acts as the memory device for female sexual development via its auto-regulatory function: its product controls the splicing of its own pre-mRNA (Bell et al., 1991). In addition, Sxl controls the splicing of the pre-mRNA from the downstream gene transformer (tra) (Boggs et al., 1987; Belote et al., 1989). The Tra product and the product of the constitutive gene transformer-2 (tra-2) (Goralski et al., 1989; Amrein et al., 1990) control the sex-specific splicing of the pre-mRNA of the gene doublesex (dsx), which is transcribed in both sexes (Burtis and Baker 1989). In females, the Tra-Tra2 complex binds to the female-specific exon (see Fig. 2A) and directs the splicing of the dsx pre-mRNA according to the female mode, giving rise to the female DsxF protein that promotes female sexual development. In males, in which no functional Tra protein is available, the dsx pre-mRNA follows the default male mode of splicing, giving rise to the mature dsxM mRNA, which produces male DsxM protein (Burtis and Baker 1989; Hoshijima et al., 1991; Hedley and Maniatis 1991; Ryner and Baker 1991; Tian and Maniatis 1993; Hertel et al., 1996). This promotes male sexual development.
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The Zn(II)2Cys6 Putative Aspergillus nidulans Transcription Factor Repressor of Sexual Development Inhibits Sexual Development Under Low-Carbon Conditions and in Submersed Culture

The Zn(II)2Cys6 Putative Aspergillus nidulans Transcription Factor Repressor of Sexual Development Inhibits Sexual Development Under Low-Carbon Conditions and in Submersed Culture

G-protein, we introduced a constitutively active allele of Transcription of rosA is upregulated during asexual fadA into a rosA-deletion strain (SKV8) by cotransforma- development and upon carbon starvation: The regula- tion of pYU8 with the pyr4-containing plasmid pRG1. tion of rosA expression was studied by Northern blot Among the obtained transformants ⵑ40% of the clones and RT-PCR analyses of mycelium harvested at different developmental stages. To synchronize development, hy- phae were grown in liquid culture for 20 hr and then filtered through a miracloth membrane and transferred to the surface of an agar plate. Under those conditions, asexual development is initiated immediately after trans- fer and stalks of conidiophores are visible after 6 hr; metulae, phialides, and some spores after 12 hr; and after 20 hr asexual development is completed. Sexual development is then initiated after 48 hr. The rosA tran- script was detectable only in the hyphal stage using nested PCR with cDNA as template, which was generated in a first reaction by RT-PCR (data not shown). The rosA transcript became detectable by Northern blot analysis only in early developmental stages and peaked in the 12-hr sample. These results suggest that the expression is tightly developmentally regulated (Figure 6). Since Figure 6.—Transient expression of rosA during develop-
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Temporal evaluation of commitment to sexual development in Plasmodium falciparum

Temporal evaluation of commitment to sexual development in Plasmodium falciparum

Treatment with furosemide did not completely block gametocyte production when used in combination with CT and MAS 7. Furosemide-treated cultures produced similar numbers of gametocytes to the untreated (no CT or MAS 7) control. This may be due to the presence of parasites already committed to sexual development within the sample or alternatively, due to the toxicity of furosamide to asexual stage parasites. Cultures treated with furosemide showed a decrease in asexual parasitaemia dur- ing assay period (10.5% reduction in Furosemide/ CT treated cultures and 34% reduction for Furosemide/ MAS 7 treated cultures) and it is acknowledged that toxicity over the assay period must be taken into consideration when interpreting this data. Given these experimental difficulties it is not possible to state definitely if furosemide blocks commitment to gametocytogenesis or not, but this data does raise the possibility, albeit tentatively that these com- pounds are acting on the parasite as indicated by the up- take of Dansyl labelled MAS peptides into the parasite infected erythrocyte.
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The biology of sexual development of Plasmodium: the design and implementation of transmission blocking strategies

The biology of sexual development of Plasmodium: the design and implementation of transmission blocking strategies

Although this field is developing rapidly the molecular basis underpinning the morphological, metabolic and functional maturation of gametocytes is poorly under- stood. The potential power of systematic ‘omics’ tech- nologies remains compromised by the large number ~50% of the malarial genes that remain un-annotated. Data generated to date have provided a rational explana- tion for the induction and specificity of natural immu- nity to gametocyte antigens. The fascination of understanding the genetic and ‘ environmental ’ control of sexual development; the roles of genetic master-regu- lators ’ e.g. AP2, and DOZI; the roles of individual genes; of the design, assembly and functions of molecular machines and organelles was apparent to all. Nowhere is this curiosity more challenged than in the ‘explosive’ process of male gamete formation, where rates/mechan- isms of DNA replication and axoneme assembly chal- lenge any current understanding. From, and for, such studies improved markers of the numerous stages of sexual development must emerge.
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A Mutant Defective in Sexual Development Produces Aseptate Ascogonia

A Mutant Defective in Sexual Development Produces Aseptate Ascogonia

development may serve other important functions. In ascog- enous hyphae of Sordaria humana it has been shown that the septal pores have highly complex and sometimes very elabo- rate membranous structures associated with them. These pore occlusions are of a type not found in vegetative hyphae and are highly variable structures from being relatively simple pore caps to having complex swollen rims with associated mem- brane cisternae (2, 49). These membranous structures provide continuity between the endomembrane systems of adjacent cells in ascogenous hyphae and particularly between the nu- clear envelopes of the two nuclei in the same cell and between nuclei in adjacent cells. This suggests that they play a special- ized role in communication in ascogenous hyphae that may, in some way, relate to the regulation of the dikaryotic state (2, 49). An important aspect of the plugging of septal pores is that they may also physically isolate adjacent hyphal compartments and thus allow them to undergo different modes of differenti- ation (15). In this way septation and septal pore plugging must play important roles in the differentiation of fungal multicel- lular structures such as perithecia (48). However, at present we do not know whether the septa of ascogonia possess septal pores, and if they do, whether they are plugged and by what type of septal pore occlusions. We speculate that plugged septa in ascogonia might allow for the accumulation of signaling molecules which trigger subsequent sexual development. Asep- tate ascogonia in pro22 may therefore prevent such an accu- mulation, and this could explain the sterile phenotype.
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Sex and the TEs: transposable elements in sexual development and function in animals

Sex and the TEs: transposable elements in sexual development and function in animals

Depending on the animal lineage, sexual development and particularly sex determination can show very different evolutionary dynamics. Some SD systems are ancient and at least 100 million years old, such as the mammalian male heterogamety system driven by the Y-linked gene Sry [54] or the avian female heterogametic determination controlled by the Z-linked dmrt1 gene [40]. In other line- ages, for instance in teleost fish, sex determination is much more labile, with a frequent switch between and even combination of ESD and GSD, and an important turn-over of sex chromosomes and master sex- determining genes in GSD [55, 56]. For example, the gen- etic sex determination system is not conserved in the genus Oryzias: while O. latipes, O. curvinotus, O. luzonen- sis and O. dancena use a XX/XY system, O. javanicus de- termines sex through ZW/ZZ female heterogamety [57]. Strikingly, Oryzias species with a XX/XY system generally have different sex chromosomes and even different master sex-determining genes: sex is controlled by dmrt1bY (aka dmy) in O. latipes and O. curvinotus, gsdfY in O. luzonen- sis and sox3Y in O. dancena [57]. Hence, the control of sexual development can be considered as a fast-evolving trait in this clade. Beyond the initiation of sex differenti- ation, the downstream molecular pathways also appear variable among animals: a comparison of genes expressed in medaka fish and mammalian gonads revealed substan- tial differences [58]. Very interestingly, the control of sex- ual development sometimes experiences convergent evolution: in both therian mammals (non-egg-laying pla- cental mammals and marsupials) and Oryzias dancena for instance, the master sex-determining gene evolved from the Sox3 gene [59]. This happened independently in the two lineages, 148 to 166 million years ago in a common ancestor of therian mammals, and less than 20 million years ago in Oryzias dancena. Another striking example is the dmrt1 gene in birds and in the tongue sole. This gene was ancestrally located on the vertebrate linkage group A, which became the Z chromosome independently in both lineages [60].
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Effect of the gene transformer of Anastrepha on the somatic sexual development of Drosophila

Effect of the gene transformer of Anastrepha on the somatic sexual development of Drosophila

ABSTRACT The gene transformer (tra) is the key regulatory memory device for sex determination in tephritid insects. The present manuscript addressed the question about the functional conser- vation of the tephritid Anastrepha Transformer protein to direct somatic sexual development in Drosophila (Drosophilidae). The transformer cDNA of Anastrepha encoding the putative full- length Tra protein was cloned in pUAST and introduced into Drosophila melanogaster. To express this protein, the GAL4-UAS system was used. The Anastrepha Tra protein induced the female- specific splicing of both dsx and fru pre-mRNAs in Drosophila XY male flies, so that these became transformed into females, though this transformation was incomplete (the sexually dimorphic foreleg basitarsus and the external terminalia were monitored). It was found that the degree of female transformation directly depended on the dose of Anastrepha tra and Drosophila trans- former-2 (tra-2) genes, and that the Anastrepha Tra-Drosophila Tra2 complex is not as efficient as the Drosophila Tra-Tra2 complex at inducing the female-specific splicing of Drosophila dsx pre- mRNA. This can explain why the Anastrepha Tra protein cannot fully substitute for the endog- enous Drosophila Tra protein.
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The clp1 Gene of the Mushroom Coprinus cinereus Is Essential for A-Regulated Sexual Development

The clp1 Gene of the Mushroom Coprinus cinereus Is Essential for A-Regulated Sexual Development

Sexual development in the mushroom Coprinus cinereus is under the control of the A and B mating- type loci, both of which must be different for a compatible, dikaryotic mycelium to form between two parents. The A genes, encoding proteins with homeodomain motifs, regulate conjugate division of the two nuclei from each mating partner and promote the formation of clamp connections. The latter are hyphal configurations required for the maintenance of the nuclear status in the dikaryotic phase of basidiomycetes. The B genes encode pheromones and pheromone receptors. They regulate the cellular fusions that complete clamp connections during growth, as well as the nuclear migration required for dikaryosis. The AmutBmut strain (326) of C. cinereus, in which both A- and B-regulated pathways are constitutively activated by mutations, produces, without mating, dikaryon-like, fertile hyphae with clamp connections. In this study we isolated and characterized clampless1-1 (clp1-1), a mutation that blocks clamp formation, an essential step in A-regulated sexual development, in the AmutBmut background. A genomic DNA fragment that rescues the clp1-1 mutation was identified by transformations. Sequencing of the genomic DNA, together with RACE experiments, identified an ORF interrupted by one intron, encoding a novel protein of 365 amino acids. The clp1-1 mutant allele carries a deletion of four nucleotides, which is predicted to cause elimination of codon 128 and frameshifts thereafter. The clp1 transcript was normally detected only in the presence of the A protein heterodimer formed when homokaryons with compatible A genes were mated. Forced expression of clp1 by promoter replacements induced clamp development without the need for a compatible A gene combination. These results indicate that expression of clp1 is necessary and sufficient for induction of the A-regulated pathway that leads to clamp development.
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Histopathological spectrum of disorders of sexual development: a case series of seven cases

Histopathological spectrum of disorders of sexual development: a case series of seven cases

Disorders of sexual development (DSD) refer to cases in which there is a discordance among at least two of the following; genetic sex, gonadal sex, genital tract sex and phenotypic sex. DSDs are quite rare with reported incidence varying from 1 in 4,500 to 1 in 5,500. Ovotesticular disorder is amongst the rarest variety of DSD comprising only to 3-10% of all cases of DSD with only 500 cases reported till now worldwide. Frequency of MRKH syndrome is 1 in 4,500 cases and is the cause of amenorrhoea in 15% of cases of primary amenorrhoea. Authors present a case series of seven cases of DSDs with three cases diagnosed as androgen insensitivity syndrome, two cases of true ovotesticular DSD (true hermaphrodite), one case each of mixed gonadal dysgenesis and Mayer-Rokitansky-Kuster Hauser (MRKH) syndrome. Authors received the histopathology specimen of these cases in this department which was extensively sampled to study the gonads and the other derivatives of Mullerian and Wolffian duct and to rule out presence of any malignancy.
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MORN1 Has a Conserved Role in Asexual and Sexual Development across the Apicomplexa

MORN1 Has a Conserved Role in Asexual and Sexual Development across the Apicomplexa

With the exception of Plasmodium species, where the sexual stages can be observed in the insect blood meal or the process of gamete development can be stimulated in vitro, our knowl- edge of sexual development has almost exclusively been lim- ited to in vivo morphology and a few immunohistological stud- ies (23, 37). To date, the sexual stages of all coccidian parasites have proven refractile to cell-culturing techniques, which has made applying modern molecular techniques difficult. Within the Coccidia, sexual development involves the formation of distinctive micro- and macrogametes that fuse to form a zy- gote. The zygote is the only diploid stage and immediately undergoes meiotic division, making all the asexual stages hap- loid. Within the definitive host, after a number of asexual cycles, merozoites entering new host cells can develop into either microgametocytes or macrogametocytes rather than un- dergoing another round of asexual proliferation. Since the parasite is haploid and there are no sex chromosomes, this development must represent a phenotypic change. However, the factors controlling the conversion to sexual development and the factors controlling whether a merozoite will develop into a micro- or macrogametocyte are unknown. The micro- gametocyte gives rise to a number (ranging from 30 to many hundreds) of microgametes, which bud from the surface of the mother cell (7, 11, 33). The motile microgametes are unique within the coccidian life cycle in possessing flagella. In contrast, macrogametogony involves an increase in size, but no nuclear division, and the synthesis of material required for oocyst wall formation (wall-forming bodies) and storage (polysaccharide granules and lipid droplets) to allow the mature macrogamete to develop into the oocyst and undergo sporulation in the external environment (8, 12, 34). In the present study, while examining the asexual development in the definitive host, we also exam- ined the expression of MORN1 during sexual development in T. gondii and in the three species of Eimeria. An unexpected finding was the observation that MORN1 was involved with the budding process during microgamete formation and was also present within the mature microgamete. Thus, MORN1 ap- pears to play a conserved, possibly structural role in both asexual and sexual development.
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Genetics of Sexual Development: An Evolutionary Playground for Fish

Genetics of Sexual Development: An Evolutionary Playground for Fish

Until recently, the steps following the initiation of sex determination were thought to follow a rather well-orchestrated and conserved cascade of genes (some of them candidates for the limited options) controlling sexual development mainly via the action of steroid hormones (Lange et al. 2002; Nakamura 2010; Angelopoulou et al. 2012; Moroha- shi et al. 2013). Among the most prominent examples of apparently conserved genes of sexual development are dmrt1 (Matson and Zarkower 2012) and sox9 (Morrish and Sinclair 2002; Kobayashi et al. 2005) for testis forma- tion and wnt4 (Smith et al. 2008) and the aromatase cyp19a1 (Valenzuela et al. 2003; Diotel et al. 2010) for ovary development. However, recent data from various fish species suggest that, in addition to the flexibility at the top of the cascade, even these key elements are not as conserved as previously assumed. This is evidenced by varying and species-specific expression patterns between sexes and throughout development (see, e.g., Vizziano et al. 2007; Ijiri et al. 2008; Hale et al. 2011; Herpin et al. 2013; and Table 1). For example, sox9 and the aromatase cyp19a1 show expres- sion patterns in East African cichlid fishes that are not con- sistent with conserved testis and ovary functions, respectively (Böhne et al. 2013).
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4.    The Process of Sexual Development and Gender Identity

4.    The Process of Sexual Development and Gender Identity

Sexual development is defined as maturing sexually, embracing and behaving in accordance with one’s gender identity; controlling one’s sexual desires or satisfying them through appropriate means and taking pleasure from sexuality, and as possessing the knowledge and skills to overcome any problems encountered with respect to sexuality. Sexuality and sexual development is biologically, psychologically and socially important for human beings. Biologically it ensures reproduction, having children and thus the continuation of posterity. Psychologically it ensures taking pleasure from sexuality itself and the satisfaction of some of the most fundamental needs of humans such as being in love and being loved. Socially it involves a myriad of processes and dimensions of human life such as being a man or woman, various social roles attached based on gender in a given society, gender identity, positive and negative expectations, emotions, sexual confidence and social status of an individual in a society [1]. In order comprehend the process of sexual development in its entirety, a number of concepts should be explained. The first among those concepts is the concept of gender. Gender describes an individual as either male or female both perceptually and physically [2]. Gender identity, on the other hand, involves acceptance of one’s gender and behaving accordingly [3]. The concept of sex typing emphasizes the nature of the traditionally designated roles of males and females. For instance, while fighting is more associated with the male sex type, crying is considered to be more suitable to the female sex type [4]. Topics such as sex, sexuality, sexual development, gender identity, social gender have been hotly debated for centuries and various different theories put forward by various theoreticians. When taken individually, none of the theories put forward so far is fully capable of accounting for sexual development on its own [5]. For this reason, the evaluation of theories in tandem with each other is highly important for better analyzing the concept of sexual development.
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Sexual development genes of Neurospora crassa.

Sexual development genes of Neurospora crassa.

The expression of most of the sdv genes also required a functional A mating type product, even under crossing growth conditions, suggesting that this product functions [r]

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Direct targets of the D  melanogaster DSXF protein and the evolution of sexual development

Direct targets of the D melanogaster DSXF protein and the evolution of sexual development

A second theme in modern considerations of the evolution of sex is that sex evolves rapidly when compared with other developmental processes (Schütt and Nöthiger, 2000). This view derives in part from the wide diversity in sex determination mechanisms used in animals, and findings that the primary sex determination mechanisms can differ in closely related species (Schütt and Nöthiger, 2000; Bull, 1985; Gempe and Beye, 2011). Furthermore, and probably independently, at a developmental level evidence for the rapid evolution of sex comes directly from the myriad of secondary sexual characteristics that frequently distinguish closely related species, which otherwise have nearly identical body plans (Darwin, 1871). Our data, cited immediately above, provides a relatively broad look at the molecular genetic level at the frequencies with which there are potential changes between sex-specific and sex-nonspecific patterns of expression of downstream effector genes. Consistent with relatively rapid evolutionary changes in the sex-specificity of expression of some of the downstream genes in the Drosophila sex hierarchy, we found that ~70% of the 23 D. melanogaster genes associated with DSX- binding sites are not associated with such a site in one or more of the other 11 sequenced Drosophila species.
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In humans, early cortisol biosynthesis provides a mechanism to safeguard female sexual development

In humans, early cortisol biosynthesis provides a mechanism to safeguard female sexual development

In humans, sexual differentiation of the external genitalia is established at 7–12 weeks post conception (wpc). During this period, maintaining the appropriate intrauterine hormone environment is critical. In contrast to other species, this regulation extends to the human fetal adrenal cortex, as evidenced by the virilization that is associated with various forms of congenital adrenal hyperplasia. The mechanism underlying these clinical findings has remained elusive. Here we show that the human fetal adrenal cortex synthesized cortisol much earlier than previously documented, an effect associated with transient expression of the orphan nuclear receptor nerve growth factor IB-like (NGFI-B) and its regulatory target, the steroidogenic enzyme type 2 3b-hydroxysteroid dehydrogenase (HSD3B2). This cortisol biosynthesis was maximal at 8–9 wpc under the regulation of ACTH. Negative feedback was apparent at the anterior pituitary corticotrophs. ACTH also stimulated the adrenal gland to secrete androstenedione and testosterone. In concert, these data promote a distinctive mechanism for normal human development whereby cortisol production, determined by tran- sient NGFI-B and HSD3B2 expression, provides feedback at the anterior pituitary to modulate androgen biosynthesis and safeguard normal female sexual differentiation.
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Changes in lipid composition during sexual development of the malaria parasite Plasmodium falciparum

Changes in lipid composition during sexual development of the malaria parasite Plasmodium falciparum

RBCs containing mature gametocytes showed more than six times as much total lipid as uninfected cells whereas the total lipid content in trophozoite infected RBCs almost triples compared to uninfected RBCs [see Additional file  4]. Neutral lipids especially contribute to this increase in total lipids during P. falciparum game- tocyte development (60-fold compared to uninfected RBCs). The proportion of neutral lipids compared to total lipids increases considerably from 3 % in uninfected RBCs to 18  % in trophozoite-infected RBCs consistent with previous reports [58, 61], and amounting to 27 % in mature gametocyte-infected RBCs (Fig. 3b). The change in phospholipids was less dramatic, but still substantial: relative to the uninfected RBCs the increase in phos- pholipids amounts in trophozoite-infected and gameto- cyte-infected RBCs were 4.8- and 6.9-fold, respectively, which might reflect the extensive proliferation of parasite membranous system [see Additional file  4]. However, Cholesterol
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In humans, early cortisol biosynthesis provides a mechanism to safeguard female sexual development

In humans, early cortisol biosynthesis provides a mechanism to safeguard female sexual development

In humans, sexual differentiation of the external genitalia is established at 7–12 weeks post conception (wpc). During this period, maintaining the appropriate intrauterine hormone environment is critical. In contrast to other species, this regulation extends to the human fetal adrenal cortex, as evidenced by the virilization that is associated with various forms of congenital adrenal hyperplasia. The mechanism underlying these clinical findings has remained elusive. Here we show that the human fetal adrenal cortex synthesized cortisol much earlier than previously documented, an effect associated with transient expression of the orphan nuclear receptor nerve growth factor IB-like (NGFI-B) and its regulatory target, the steroidogenic enzyme type 2 3b-hydroxysteroid dehydrogenase (HSD3B2). This cortisol biosynthesis was maximal at 8–9 wpc under the regulation of ACTH. Negative feedback was apparent at the anterior pituitary corticotrophs. ACTH also stimulated the adrenal gland to secrete androstenedione and testosterone. In concert, these data promote a distinctive mechanism for normal human development whereby cortisol production, determined by tran- sient NGFI-B and HSD3B2 expression, provides feedback at the anterior pituitary to modulate androgen biosynthesis and safeguard normal female sexual differentiation.
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ABNORMALITIES OF SEXUAL DEVELOPMENT

ABNORMALITIES OF SEXUAL DEVELOPMENT

Ftc. Example of female from Group 5. The general appearance of the external genitalia is normal i)tmt close inspection showed only a single orifice, which proved to be a urogenital sinus[r]

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