甲壳动物性别分化的遗传决定及外界因素的影响

本文综述了甲壳动物的性别决定机理及外界因素对性别分化的影响,绝大多数甲壳动物没有明显的性染色体,促雄腺被认为是甲壳动物性别分化的最主要的决定因子,其作用已得到了广泛的证明,由于甲壳动物幼体在早期发育过程中具有向两性发育的潜能,促雄腺可以决定个体未来发育的性别,并且通过人为摘除或移植促雄腺的方法可以使性别已经分化的个体发生性逆转,从而改变幼体的性别。虽然甲壳动物的性别是由遗传决定的,但外界的因素经如寄生,光周期,温度或激素可以改变其性比,其中以寄生的影响研究比较多,并认为是影响某些甲壳动物性别分化的主要外界因子,由于大多数养殖的甲壳动物雌雄性之间的有体重和体长的差异在水产养殖中可以利用这些特征进行全雌全雄种苗的生产,以提高产量和效益。     

甲壳动物胰岛素样促雄腺激素功能及 作用机制的研究进展

甲壳动物的雄性性别分化主要由其促雄腺（AG）分泌的胰岛素样促雄腺激素（IAG）负责调控。在罗氏沼虾（Macrobrachium rosenbergii）中,通过单个IAG的操作可以成功性反转,进而实现全雄养殖。因此,基于IAG的性别调控技术具有良好的应用潜力。目前,IAG在许多经济甲壳动物中得到研究报道,发现其表达不仅局限于促雄腺,功能也更加广泛。此外,随着RNA干扰技术在水产动物中的广泛运用,基因功能的研究更易实现,IAG如何执行其生理作用的信号机制及上游的调控网络逐渐成为学者们探究的热点。本文综述了近年来有关IAG研究的进展,从IAG的分子特征、生理功能、作用机制及上游调控机理等方面展开探讨,为深入阐明IAG的生理功能及作用机制提供基础。     

甲壳动物的性别决定

甲壳动物的性别决定吴融（浙江省海洋水产养殖研究所温州．３２５０００）甲壳动物一般为两性生殖．雌雄比率为１：１。据ｃｈａｒｎｌａｕｘ－Ｃｏｔｔｏｎ（１９６０）在端足目Ｏｒｃｈｅｓｔ。ａｇａｉｎ－ｍａｒｅｌｌａ（滨蚤）上的研究,知道性别的分化与造雄腺分泌...     

中国大鲵幼体的性别分子鉴定与性腺形态

为探明中国大鲵(Andrias davidianus)雌雄幼体的性腺发育特征,确定适合的性别分子鉴定方法,对15尾5月龄和17尾17月龄养殖个体进行形态测量、解剖观察、性腺组织切片及PCR扩增雌性特异DNA片段。结果发现,引物adf225和adf340的扩增效果好,判定5月龄个体8雌7雄;17月龄个体8雌9雄,与依据性腺形态结构区分的结果一致。体视显微镜下5月龄幼体中肾腹侧有两条半透明细条状的原始生殖嵴;组织切片显示生殖细胞形态分化不明显。17月龄卵巢波浪状弯曲,有颗粒感,精巢呈光滑的白条状,形态分化明显;组织切片显示,卵巢分化出体积较大的卵母细胞,同时保留原始卵泡,精巢分化出生精小叶和精原细胞、支持细胞。外形测量显示,5月龄与17月龄性二型不明显,不能根据外形判断性别。本研究确定了大鲵幼体性别分子鉴定的最佳引物,可用于养殖过程中雌雄选配,以节约资源。     

Rspo1对雌性脊椎动物性别分化的功能

长期以来雌性脊椎动物的性别分化被认为是一个“默认”的程序.但是近些年研究发现,Rspo1基因的突变或缺失可导致哺乳动物XX型个体性反转为雄性.Rspo1在鱼类、两栖爬行类、鸟类和哺乳类动物性腺发育的不同阶段表达,其表达在雌雄个体性别分化时期有差异,是潜在的性别调控基因.Rspo1在性别发育早期可通过Wnt/β-catenin信号通路调控性腺分化相关因子的表达,影响原始生殖细胞分裂增殖、细胞周期和生长发育,参与调控性腺中体细胞的分化.本文总结了近年来Rspo1在脊椎动物中的表达调控及其在雌性性别决定方面功能的研究进展.     

浅析斑马鱼性别决定的调控因素

性别决定是经典而高度保守的生物过程。在许多物种中性别决定是以遗传为基础的,个体所携带的性染色体决定了性别。然而,由于鱼类性腺发育呈现高度可变性、复杂性的特点,其性别决定机制仍未有定论。斑马鱼作为一个研究发育和疾病的重要脊椎模式动物,性别决定和分化的高度可塑性使其成为研究生理和环境因素对性腺发育影响及其作用机制的独特模型。本综述总结近年来对斑马鱼性别决定及分化过程的研究,为探索鱼类性别决定机制提供新的见解。     

Rspo1在雌性脊椎动物性别分化中的功能

长期以来雌性脊椎动物的性别分化被认为是一个"默认"的程序.但是近些年研究发现,Rspo1基因的突变或缺失可导致哺乳动物XX型个体性反转为雄性.Rspo1在鱼类、两栖爬行类、鸟类和哺乳类动物性腺发育的不同阶段表达,其表达在雌雄个体性别分化时期有差异,是潜在的性别调控基因.Rspo1在性别发育早期可通过Wnt/β-catenin信号通路调控性腺分化相关因子的表达,影响原始生殖细胞分裂增殖、细胞周期和生长发育,参与调控性腺中体细胞的分化.本文总结了近年来Rspo1在脊椎动物中的表达调控及其在雌性性别决定方面功能的研究进展.     

性畸变对腹足类生殖和种群的影响

具有内分泌干扰效应的三丁基锡能引起腹足类产生性畸变现象。在性畸变过程中 ,雌性个体会由于生殖孔口被前列腺取代或被输精管阻塞、贮精囊或卵囊腺开裂、卵囊腺内部被阻塞以及卵巢转化为精巢等多种原因而丧失生殖能力 ,甚至死亡。并由此引起雌、雄性比和幼、成年个体比的降低 ,导致种群衰退。有浮游幼体的种类可以通过外来种群的成功迁入使种群得以维持 ,而无浮游幼体的种类 ,由于幼体迁移能力差而最终导致种群的区域性灭绝。性畸变这种典型的功能效应对其他内分泌干扰物质的生态效应研究有启示作用。     

温度对爬行动物性别的影响

性别的分化可以同时看作是遗传学、胚胎学、内分泌学和生态学的问题。一般高等动物的胚胎性别发育包括下述三个步骤:1.遗传性别的决定,由精卵双方所携带的性染色体结合产生;2.性腺性别的出现,未分化的性腺发育成睾丸或卵巢;3.性腺性别转变为个体表现型性别。正常情况下,绝大多数的脊椎动物性别表现往往是由性染色体决定,即xx或zw决定雌性,xy或zz决定雄性。但是,有些爬行动物的性别表现并不完全如此,它还受着环境因素的影响。在本文中,作者将对温度与爬行动物性别表现的关系进行初步地探讨。有趣的发现     

高密度种群的社会条件对大沙鼠幼年雄鼠形态生理特征的影响:一个实例研究

在非繁殖期高密度大沙鼠(RhombomysopimusLicht)种群中,我们研究了胁迫以及幼年雄鼠的性激素浓度、腹中腺大小和体重对社群中成体(≥1年龄)存在的依赖性。用无损伤放射免疫方法,测定了于1999年秋天在野外采集的幼年雄鼠粪便样品的皮质酮和睾酮。有成体社群中的个体大于无成体社群的个体。因此,成体对幼体的影响不能独立于社群大小本身。成体的存在促进粪便睾酮的浓度和抑制了幼年雄鼠的性成熟(由雄激素依赖的较小的腹中腺来估计),但是同时促进了其生长。因此,自然存在的社群环境能影响幼年雄鼠的形态生理特征发育。     

Steroid hormone-induced male sex determination in an amniotic vertebrate.

In many reptiles, sex is determined by the temperature at which the eggs are incubated (i.e., temperature-dependent sex determination, or TSD). Past studies have shown that exogenous steroid hormones can override the effects of temperature and induce female sex determination. However, past attempts to induce male sex determination have consistently failed. In the present study, sex determination was studied in a turtle with TSD. By utilizing an incubation temperature regimen that resulted in approximately a 1:1 sex ratio in the control group, sex determination was shown to be sensitive to both exogenous androgen and estrogen treatments: androgen induced the production of male hatchlings, whereas estrogen induced the production of female hatchlings. This is the first report of an amniotic vertebrate in which an exogenous steroid hormone induces male sex determination.     

Development of the testes in female domestic fowls submitted to an experimental sex reversal during embryonic life

The sex differentiation of the female chick embryo can be totally inverted toward the male sex by an early extraembryonic testis grafting. This sex reversal remains permanent, as shown by three adult fowls described in this paper. They possess two testes associated with normally differentiated male excretory ducts and their Müllerian ducts have regressed. The development of male sex characteristics such as external features, behavior and complete spermatogenesis is evidence that these cocks have endocrine and exocrine capabilities similar to those of normal cocks. Although these cocks were able to mate with female fowls, they were sterile. A mechanism is discussed by which grafted testes induce such modifications in females. Hypotheses considering the heterogametic sex (female in birds) as exerting a dominant influence on the phenotypic sexual differentiation can be discarded in light of our results because a homogametic testis provokes the definitive sex inversion of a female.     

Effects of androgens on sex differentiation of the urodele Pleurodeles waltl

In nonmammalian vertebrates, steroids have been hypothesized to induce somatic sex differentiation, since manipulations of the steroidal environment of gonads have led to various degrees of sex reversal. Whereas the critical role of estrogens in ovarian differentiation is well documented, studies on androgens have produced a perplexing variety of results depending upon species variations and nature of androgens used. In this way, testosterone induces masculinization of females in some species but provokes paradoxical feminization of males in many other species such as the urodelan Pleurodeles waltl. In reptiles this phenomenon could be interpreted by conversion of exogenous testosterone to estradiol by aromatase. Treatments of Pleurodeles larvae with nonaromatizable androgens bring support to this hypothesis and suggest a role of androgens in sex differentiation. Dihydrotestosterone (DHT) could not induce the paradoxical feminization of ZZ larvae. In addition, DHT as well as 11beta-hydroxy-androstenedione could drive a functional male differentiation of ZW larvae. Moreover, other 5alpha reduced androgens also induced sex reversal of female larvae. Yet, the 5alpha reductase inhibitor CGP 53133 and antiandrogens such as flutamide or cyproterone acetate did not exert any effect on male sex differentiation of ZZ larvae. Though the precise role of androgens is still unknown, especially for 11-oxygenated androgens, our results suggest an implication in male sex differentiation. In this way, testosterone could play a pivotal role in being metabolized either into other androgens during testis differentiation or into estradiol during ovarian differentiation.     

Genetic mechanisms underlying male sex determination in mammals

Genetic control of gonadal development proceeds through either the male or female molecular pathways, driving bipotential gonadal anlage differentiation into a testis or ovary. Antagonistic interactions between the 2 pathways determine the gonadal sex. Essentially sex determination is the enhancement of one of the 2 pathways according to genetic sex. Initially, Sry with other factors upregulatesSox9 expression in XY individuals. Afterwards the expression ofSox9 is maintained by a positive feedback loop withFgf9 and prostaglandin D₂ as well as by autoregulative ability of Sox9. If these factors reach high concentrations, then Sox9 and/or Fgf9 may inhibit the female pathway. Surprisingly, splicing, nuclear transport, and extramatrix proteins may be involved in sex determination. The male sex determination pathway switches on the expression of genes driving Sertoli cell differentiation. Sertoli cells orchestrate testicular differentiation. In the absence of Sry, the predomination of the female pathway results in the realization of a robust genetic program that drives ovarian differentiation.     

The environmental-endocrine basis of gynandromorphism (intersex) in a crustacean

Commensurate with the decline in many crustacean populations has been an accumulation in reports of sexually ambiguous individuals within these populations. The cause of gynandromorphism or intersex among crustaceans is unknown. We show that gynandromorphism in the branchiopod crustacean Daphnia magna is initiated by the sex-determining hormone methyl farnesoate when levels of the hormone are intermediate between low levels that stimulate the production of broods containing all female offspring and high levels that stimulate the production of broods of all male offspring. The incidence of hormonally-induced gynandromorphism was low (0.14% at the maximum stimulatory hormone concentrations) but was significantly increased (46-fold) when the animals were hormone-treated at 30 degrees C. Some environmental chemicals also can stimulate the gynandromorphic phenotype as we demonstrated with the insecticide pyriproxyfen. Gynandromorphism occurs due to inadequate signaling of male-sex determination since: a) gynandromorphs did not occur in a population that was producing only female offspring; and, b) conditions that stimulated gynandromorphism also reduced the incidence of male offspring. We suggest that male sex determination normally occurs prior to the first embryonic cleavage. Elevated temperature may alter the timing of sex determination such that methyl farnesoate signaling occurs after the first embryonic cleavage and bilateral gynandromorphism occurs as a consequence of signaling to only one of the daughter cells. These results demonstrate that environmental factors can cause aberrant sex determination via perturbations in methyl farnesoate signaling.     

Zebrafish sex determination and differentiation: Involvement of FTZ-F1 genes

Sex determination is the process deciding the sex of a developing embryo. This is usually determined genetically; however it is a delicate process, which in many cases can be influenced by environmental factors. The mechanisms controlling zebrafish sex determination and differentiation are not known. To date no sex linked genes have been identified in zebrafish and no sex chromosomes have been identified. However, a number of genes, as presented here, have been linked to the process of sex determination or differentiation in zebrafish. The zebrafish FTZ-F1 genes are of central interest as they are involved in regulating interrenal development and thereby steroid biosynthesis, as well as that they show expression patterns congruent with reproductive tissue differentiation and function. Zebrafish can be sex reversed by exposure to estrogens, suggesting that the estrogen levels are crucial during sex differentiation. The Cyp19 gene product aromatase converts testosterone into 17 beta-estradiol, and when inhibited leads to male to female sex reversal. FTZ-F1 genes are strongly linked to steroid biosynthesis and the regulatory region of Cyp19 contains binding sites for FTZ-F1 genes, further linking FTZ-F1 to this process. The role of FTZ-F1 and other candidates for zebrafish sex determination and differentiation is in focus of this review.     

Sexual differentiation in higher plants

As in all organogenetic studies, sex differentiation implies three fields of research: the identification of macsomalecular markers specific for stamens or carpels }expression program); the analysis of the signals (such as phytohormones{ inducing them: and finally the knowledge of the regulator genes (sex detsrmination genes).
In dioecious plants, sex determination (male and female genes or combinations of heterochromosomes) occurs at the fertilization stage. These regulators probably act early by means of inducers; their action is perceptible mainly when reproductive organs develop, and the existence of target cells able to respond to the inductive message is obvious. Experimental control of sex expression by phytohormones leads to the repression of the normal organogenetic program (induced by the presence of the sex genes) arrd to the induction of the opposite program (normally incompatible with these genes).
Analogous mechanisms occur in monoecious plants, but here the male program is always expressed before the female one. Experimental control of sex expression suggests that the successive induction of first male and then the female program results from inverse gradients of male and female signals. Sex differentiation studies of monaeciousness are more difficult than studies on dioeciousness since, in all the cells. not only male and female programs co-exist as in dioeciousness, but also male and female sex penes and signals.
The present model has been tested in the dioecious Mercurialisannua (2n = 16) and some other species, and the results that have been established are described.     

Gonadal sex reversal in triploid rainbow trout (Oncorhynchus mykiss).

Sex determination in salmonids is primarily governed by sex chromosomes; however, phenotypic expression and successful development of the gonads may be influenced by additional factors. Exposure to exogenous steroids during the critical period of gonadal differentiation will reverse the expected phenotypic sex of both female and male trout. Triploidy, a viable condition in rainbow trout (RBT), alters the degree of gonadal development in a gender-specific manner. Males produce testes with similar morphology and function as diploid fish, but females produce underdeveloped ovaries devoid of growing oocytes. One possible explanation for this observed gender difference is that the timing of meiotic initiation may influence ovarian/testicular development in triploid RBT. To determine whether the early entrance of germ cells into meiosis results in the lack of ovarian development in triploid females, the objective of this study was to sex-reverse genotypic triploid female RBT (XXX) into phenotypic males and genotypic triploid male RBT (XXY) into phenotypic females. Male fish were exposed to estradiol-17beta (E(2)) and females were exposed to the non-aromatizable androgen 17alpha-methyldihydrotestosterone (MDHT). Over 90% of the male fish treated with exogenous E(2) developed gonadal structures indistinguishable from the gonads of triploid females. Triploid female RBT treated with MDHT developed testes; however, not all fish treated with this androgen were completely sex reversed. The results of this investigation are consistent with the hypothesis that the failure of ovarian development in triploid RBT is due to the early onset of meiosis and does not appear to be due to genotypic sex. J. Exp. Zool. 284:466-472, 1999.     

脊椎动物性别决定和分化的分子机制研究进展

哺乳类性别决定是多种转录因子和生长因子相继表达和相互调控的结果。SRY的表达启动雄性通路并诱导下游雄性特异基因SOX9、AMH等的表达。FOXL2在雌性未分化性腺表达,WNT-4和DAX1也在雌性性别决定或分化时期表达,表明雌性通路也是受特定基因调控的,而并非“默认通路”。鸟类的性别也是由遗传基因决定的,EFT1(雌性)和DMRT1(雄性)可能是性别决定候选基因。爬行类为温度性别决定的典型,温度可能通过调节雌激素水平和控制性别特异遗传基因表达决定性别。大部分两栖类性别受环境因素影响,但发现DMRT1和DAX1可能与其精巢发育有关。鱼类性别决定和分化方式差异很大,多种因素(遗传基因、环境因素、类固醇激素等)参与了这一过程。从青Q鳉Y染色体定位克隆的DMY,被认为是第一个非哺乳类脊椎动物雄性性别决定基因。所有这些表明脊椎动物性别决定和分化机制是多样化的。     

Sex determination and differentiation in Asparagus officinalis L.

The paper summarizes the coordinated researches conducted by three Italian groups in the area of sex determination and differentiation in the dioecious species Asparagus officinalis. Morphological evidence indicates that sex differentiation in Asparagus consists essentially of selective abortion of gynoecium or androecium of initially hermaphroditic floral primordia occurring in genotypically determined male and female individuals. Abortion occurs in pollen-mother cells and anthers in females and in megaspore-mother cells but not in the vegetative tissues of the ovary in males. The differential developmental pathway is accompanied by changes in relative abundance of auxin and cytokinins. The genetic ssytem controlling abortion of male or female organs is apparently monogenic (possibly a bipartite gene) with factor(s) associated with the homomorphic chromosome pair L5. Other genes influence the development of reproductive structures as indicated by the presence of genetic factors controlling stylar growth in male plants. The presence of extensive polymorphism in isoenzyme and DNA restriction fragment length patterns (RFLP) allows the search for markers associated with ‘sex genes’: a locus encoding a malic dehydrogenase (MDH) isoenzyme has been found about 20 cM from sex genes implying that chromosomes in which sex factors are located could pair and recombine. Searches for messages specifically expressed in reproductive structures were conducted by 2D-electrophoresis of existing and newly synthesized polypeptides or of in vitro translation products of poly(A) ⁺RNA from male and female flowers and by isolating specific monoclonal antibodies against sex specific floral antigens.