Early gonadal development and primary males in the protogynous epinepheline, Cephalopholis boenak

Early gonadal development of the protogynous epinepheline, Cephalopholis boenak, was examined histologically in 289 specimens with standard length (L_S) of 42–130 mm, collected from May 2000 to April 2002 in Hong Kong waters, to determine male developmental pathways and establish its sexual pattern. All juvenile gonads developed an ovarian lumen with primary‐growth stage oocytes and scattered spermatogenic tissue prior to sexual differentiation and first sexual maturation. From this bisexual phase containing both female and male tissues, some gonads differentiated as ovaries with further oocyte growth to cortical‐alveolus and vitellogenic stages, the rest differentiated as testes with the proliferation of spermatogenic tissue and the formation of a sperm sinus. All testes retained the lumen and primary‐growth stage oocytes, and sperm sinuses ran within the gonad wall. Unlike most protogynous species, among functional males it was impossible to distinguish those resulting from juveniles through sexual differentiation (i.e. primary male) from those resulting from functional females through sex change (i.e. secondary male) based solely on testicular morphology. A proportion‐spermatogenic‐tissue index (I_ST) was, therefore, developed and determined to be a reliable quantitative indicator for distinguishing differentiating, primary males before a sperm sinus was evident, from differentiating females during sexual differentiation. Since sexually transitional specimens with the concominant appearance of degenerating vitellogenic, or later, stage oocytes and spermatogenic tissue in the gonads were previously noted from Hong Kong, diandric, protogynous hermaphroditism is confirmed in C. boenak. For species, such as this and other epinephelines, in which all males have the same testicular morphology, a complete analysis of a wide range of body sizes from juveniles to adults is necessary for understanding male developmental pathways, and determining sexual pattern.     

Sexual differentiation of the zebra finch song system: Positive evidence,negative evidence,null hypotheses,and a paradigm shift

Permanent sex differences in the brain are found in many vertebrates, and are thought to be induced by sex differences in secretion of gonadal steroid hormones during critical periods of early development. This theory has received support primarily from many experiments conducted on mammals, but also from studies on other vertebrate classes, including birds. The only avian neural dimorphism that has allowed extensive tests of this hypothesis is the neural circuit for song in passerine birds, which is much larger in males than in females. Experiments in zebra finches have yielded contradictory results. Although it is relatively easy to induce masculine patterns of development in genetic females with estrogen, it has not been possible to induce feminine patterns of development in males with any treatments, including antiestrogens and inhibitors of estrogen synthesis. Moreover, genetic females that develop with large amounts of functional testicular tissue but with virtually no ovarian tissue nevertheless have a feminine song circuit. The latter studies fail to support the idea of steroid induction of sexual differentiation. An alternative to the steroidal control hypothesis is that nonhormonal gene products expressed in the brain early in development trigger sexually dimorphic patterns of development. Although current evidence in several neural and nonneural systems indicates that sexual differentiation of some somatic phenotypes cannot be explained by the actions of gonadal steroids, the idea of direct genetic (nonhormonal) induction of sexual differentiation has yet to be proved. © 1997 John Wiley & Sons, Inc. J Neurobiol 33: 572–584, 1997     

Testicular type Sox9 is not involved in sex determination but might be in the development of testicular structures in the medaka, Oryzias latipes

Testicular type Sox9 is the most upstream conserved gene in the sex determining cascade among vertebrate. However, in medaka, only one Sox9 gene was identified as expressed in the ovary; no other Sox9 gene was reported expressed in the testis. We explored the medaka genome and cloned a novel testicular type Sox9 cDNA. Phylogenetic analysis revealed that both our isolated Sox9 and the already reportedly cloned medaka Sox9 belongs zebrafish Sox9a branch. Therefore, we named our gene Sox9a2. Unexpectedly, Sox9a2 mRNA was expressed in somatic cells surrounding germ cells at similar high levels in both sexes during early gonadal sex differentiation. However, at the initial stage of testicular tubules development, the expression of Sox9a2 was maintained only in XY gonads, and was remarkably reduced in XX gonads. These results suggest that Sox9a2 is not involved in early sex determination and differentiation, but is involved in the later development of testicular tubules in medaka.     

Gonadal morphogenesis and sex differentiation in the viviparous fish Chapalichthys encaustus (Teleostei, Cyprinodontiformes, Goodeidae)

This study describes the structural and ultrastructural characteristics of gonadal sex differentiation and expression of Vasa, a germline marker, in different developmental stages of embryos and newborn fry of the barred splitfin Chapalichthys encaustus, a viviparous freshwater teleost endemic to Mexico. In stage 2 embryos, the gonadal crest was established; gonadal primordia were located on the coelomic epithelium, formed by scarce germ and somatic cells. At stage 3, the undifferentiated gonad appeared suspended from the mesentery of the developing swimbladder and contained a larger number of germ and somatic cells. At stages 4 and 5, the gonads had groups of meiotic and non-meiotic germ cells surrounded by somatic cells; meiosis was evident from the presence of synaptonemal complexes. These stages constituted a transition towards differentiation. At stage 6 and at birth, the gonad was morphologically differentiated into an ovary or a testis. Ovarian differentiation was revealed by the presence of follicles containing meiotic oocytes, and testicular differentiation by the development of testicular lobules containing spermatogonia in mitotic arrest, surrounded by Sertoli cells. Nuage, electron-dense material associated with mitochondria, was observed in germ cells at all gonadal stages. The Vasa protein was detected in all of the previously described stages within the germ-cell cytoplasm. This is the first report on morphological characteristics and expression of the Vasa gene during sexual differentiation in viviparous species of the Goodeidae family. Chapalichthys encaustus may serve as a model to study processes of sexual differentiation in viviparous fishes and teleosts.     

Ontogeny of sexual development in the roach (Rutilus rutilus) and its interrelationships with growth and age

The roach (Rutilus rutilus) has become a sentinel species for the study of sexual disruption in wild fish populations as a consequence of exposure to endocrine disrupting chemicals (EDCs). Little is known, however, about the normal ontogeny of sexual development in this species. Here, we analyzed the ontogeny of sexual development in captive‐bred roach and assessed how growth rate and fish size affected the timing of both sexual differentiation and sexual development over a 2‐year period. Ovarian differentiation was first recorded at 68 days post‐fertilization (dpf) and this preceded testicular differentiation (first recorded at 98 dpf). In contrast, sexual maturation occurred at an earlier age in males (300 dpf) compared with females (728 dpf). No differences in body size (length or weight) were recorded between male and female roach until the fish were 415 dpf. Studies on three populations of roach which grew at different rates showed that the timing of sexual differentiation was highly variable and more related to fish size than to fish age. Time to sexual maturation was also variable among populations but, subsequent to their first year of life, gonadal status was less well associated with fish size. Interestingly, the sex ratio of the population was biased towards females in populations that grew more rapidly during early life. The findings presented here provide a valuable foundation of work to support both field‐ and laboratory‐based assessments on the effects of EDCs, and other stressors, on sexual differentiation and development in the roach. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Sexual differentiation in the spiny softshell turtle (Apalone spinifera), a species with genetic sex determination

It is hypothesized on the basis of sex determination theory that species exhibiting genetic sex determination (GSD) may undergo sexual differentiation earlier in development than species with environmental sex determination (ESD). Most turtle species exhibit a form of ESD known as temperature-dependent sex determination (TSD), and in such species the chronology of sex differentiation is well studied. Apalone spinifera is a species of softshell turtle (Trionychidae) that exhibits GSD. We studied sexual differentiation in this species in order to facilitate comparison to TSD species. Eggs were incubated at two different temperatures and embryos were harvested at various stages of mid to late development. Gonad length was measured with image analysis software, then prepared histologically. Indifferent gonads have differentiated in stage 19 embryos. Histological details of gonadogenesis follow the same pattern as described for other reptiles. Regression of the male paramesonephric duct closely follows testicular differentiation. Gonad lengths are longer at the warmer incubation temperature, and ovaries are generally longer than testes at each stage and for each temperature. Although sexual differentiation takes place at about the same stage as in other turtles with TSD (18-20), in A. spinifera this differentiation is irreversible at this stage, while in some of the TSD species sex is reversible until about stage 22. This immutable, definitive sexual differentiation may support the hypothesis of an accelerated chronology of sex differentiation for this species. We also note that sexual dichromatism at hatching is known in this species and may provide additional evidence of early differentiation. J. Exp. Zool. 290:190-200, 2001.     

PROCESS OF FUNCTIONAL SEX REVERSAL OF THE GONAD IN THE FEMALE GUPPY, POECILIA RETICULATA, TREATED WITH ANDROGEN BEFORE BIRTH

In the guppy, Poecilia reticulata , ovarian differentiation occurs during the embryonal life by 14 days after the preceding parturition. Testicular differentiation begins with the appearance of prominent aggregations of stroma cells in the gonadal hilus occurring by 18 days following the last parturition.
Oral administration of methyltestosterone (400 μ/g diet) to gravid guppies, begun 13–15 days after the preceding parturition and continued until the end of gestation, induced a male-type aggregation of somatic cells in the hilus of ovaries of female embryos. Gonads of newly born, androgenized females still had developing oocytes but were always provided with atypical clusters of stroma cells in their hilus. The gonads of affected female offspring developed successively into definite testes within 20 days after birth, displaying a precocious differentiation of the hilar stroma into sperm ducts and testicular interstitium, a concomitant initiation of spermatogenesis, and a conspicuous degeneration of oocytes. A successful masculinization of the somatic element, which may occur prior to that of the germ cells, in androgen-affected embryonic ovaries seems to be essential for the functional sex reversal of genetic females in the guppy.     

Normal prenatal but arrested postnatal sexual development of luteinizing hormone receptor knockout (LuRKO) mice

To study further the role of gonadotropins in reproductive functions, we generated mice with LH receptor (LHR) knockout (LuRKO) by inactivating, through homologous recombination, exon 11 on the LHR gene. LuRKO males and females were born phenotypically normal, with testes, ovaries, and genital structures indistinguishable from their wild-type (WT) littermates. Postnatally, testicular growth and descent, and external genital and accessory sex organ maturation, were blocked in LuRKO males, and their spermatogenesis was arrested at the round spermatid stage. The number and size of Leydig cells were dramatically reduced. LuRKO females also displayed underdeveloped external genitalia and uteri postnatally, and their age of vaginal opening was delayed by 5-7 days. The (-/-) ovaries were smaller, and histological analysis revealed follicles up to the early antral stage, but no preovulatory follicles or corpora lutea. Reduced gonadal sex hormone production was found in each sex, as was also reflected by the suppressed accessory sex organ weights and elevated gonadotropin levels. Completion of meiosis of testicular germ cells in the LuRKO males differs from other hypogonadotropic/cryptorchid mouse models, suggesting a role for FSH in this process. In females, FSH appears to stimulate developing follicles from the preantral to early antral stage, and LH is the stimulus beyond this stage. Hence, in each sex, the intrauterine sex differentiation is independent of LH action, but it has a crucial role postnatally for attaining sexual maturity. The LuRKO mouse is a close phenocopy of recently characterized human patients with inactivating LHR mutations, although the lack of pseudohermaphroditism in LuRKO males suggests that the intrauterine sex differentiation in this species is not dependent on LH action.     

Gonadal sex differentiation in Alligator mississippiensis,a species with temperature-dependent sex determination

Temperature of egg incubation determines sex in Alligator mississippiensis hatchlings. To define the timing and morphology of sexual differentiation, alligator gonads were examined histologically and ultrastructurally throughout embryogenesis. At the male-producing temperature (33° C), the onset of testis differentiation occurred in most embryos during developmental stages 21–22, when a number of somatic cells in the medulla of the gonad became enlarged, forming presumptive Sertoli cells. Some enlarged somatic cells were also observed at the female-producing temperature (30° C) during gonadogenesis, but they were less widespread than at 33° C. Ovarian differentiation at 30° C began slighlty later, during stage 22–23, and was characterised by proliferation of germs cells in the cortex of the gonad. Testis formation in alligators may depend upon presumptive Sertoli cells differentiating prior to a critical event in embryogenesis, such as germ cell proliferation and meiosis. If follows that ovary formation occurs if this requirement is not met, as at lower incubation temperatures.     

Ultrastructural aspects of gonadal morphogenesis in Bufo bufo (Amphibia Anura) 1. Sex differentiation

The morphogenesis of gonads in Bufo bufo tadpoles was studied, and ultrastructural differences between sexes were identified. All specimens analyzed initially developed gonads made up of a peripheral fertile layer (cortex) surrounding a small primary cavity. Subsequently a central layer of somatic cells (medulla) developed. Both layers were separated by two uninterrupted basal laminae between which a vestige of the primary cavity persisted. During female differentiation, the peripheral layer continued to be the fertile layer. In males, the central layer blended into the peripheral layer and the basal laminae disappeared. The somatic cells of the central layer came into direct contact with the germ cells; this did not occur in females. Testicular differentiation continued with the migration of germ cells towards the center of the gonad. The somatic elements surrounding the germ cells appeared to play an active role in their transfer to the center of the gonad. The peripheral layer shrank and became sterile. Two basal laminae then re-formed to separate the fertile central layer from the peripheral sterile one. Germ cells have always been thought to perform a passive role in sex differentiation in amphibians. Following the generally accepted "symmetric model", the mechanism of gonad development is symmetrical, with cortical somatic cells determining ovarian differentiation and medullary somatic cells determining testicular differentiation. In contrast, we found that sex differentiation follows an "asymmetric" pattern in which germ cells tend primarily toward a female differentiation and male differentiation depends on a secondary interaction between germ cells and medullary somatic cells.     

Sxl in the germline of Drosophila: A target for somatic late induction

In Drosophila, the sex of germ cells is determined by autonomous and inductive signals. Somatic inductive signals can drive XX germ cells into oogenesis or into spermatogenesis. An autonomous signal makes XY germ cells male and unresponsive to sex determination by induction. The elements forming the X:A ratio in the soma and the genes tra, tra2, dsx, and ix that determine the sex of somatic cells have no similar role in the germline. The gene Sxl, however, is required for female differentiation of somatic and germ cells. Inductive signals that are dependent on somatic tra and dsx expression already affect the sex-specific development of germ cells of first instar larvae. At this early stage, however, germline expression of Sxl does not appear to affect the sexual characteristics of germ cells. Since inductive signals dependent on tra and dsx nevertheless influence the choice of sex-specific splicing of Sxl, it can be concluded that Sxl is a target of the inductive signal, but that its product is required late for oogenesis. Other genes must therefore control the early sexual dimorphism of larval germ cells. © 1994 Wiley-Liss, Inc.     

Gonadal development and sex differentiation in the cichlid fish Cichlasoma dimerus (Teleostei, Perciformes): a light- and electron-microscopic study

Although the overall pattern and timing of gonadal sex differentiation have been established in a considerable number of teleosts, the ultrastructure of early stages of gonadal development is not well documented. In this study, gonads from larval and juvenile stages of laboratory-reared Cichlasoma dimerus were examined at the light-microscopic and ultrastructural levels. This freshwater species adapts easily to captivity and spawns with high frequency during 8 months of the year, providing an appropriate model for developmental studies. Larvae and juveniles were kept at a water temperature of 26.5 +/- 1 degrees C and a 12:12 hour photoperiod. Gonadal development was documented from 14-100 days postfertilization, covering the period of histologically discernible sex differentiation. Gonadal tissue was processed according to standard techniques for light and electron microscopy. C. dimerus, a perciform teleost, is classified as a differentiated gonochorist, in which an indifferent gonad develops directly into a testis or ovary. On day 14, the gonadal primordium consists of a few germ cells surrounded by enveloping somatic cells. Ovarian differentiation precedes testicular differentiation, as usual in teleost fishes. The earliest signs of differentiation, detected from day 42 onward, include the onset of meiotic activity in newly formed oocytes, which is soon accompanied by increased oogonial mitotic proliferation and the somatic reorganization of the presumptive ovary. The ovarian cavity is completely formed by day 65. Numerous follicles containing perinucleolar oocytes are observed by day 100. In contrast, signs of morphological differentiation in the presumptive testis are not observed until day 72. By day 100, the unrestricted lobular organization of the testis is evident. The latest stage of spermatogenesis observed by this time of testicular development is spermatocyte II.     

Sex-related genes expression in juveniles of red abalone,Haliotis rufescens (Swanson, 1822)

The red abalone, Haliotis rufescens (Swanson 1822), is an important species for commercial aquaculture in Mexico, with annual production levels around 68 t. Not surprisingly, there is great interest in increasing production and its cultivation success. In order to have a better understanding of the genes involved in the sexual differentiation, this study analyzed the early differential expression of eight sex-related genes (VCP 2.2, VERL, VTGI, DMRT1, FP, LYS, SARIP, TEKT A1) by RT-qPCR in abalones from 5 to 45 mm of shell length. Sex identification was evaluated using VERL and LYS genes expression levels. These genes differentiated females and males from 16 mm of shell length and above. All eight sex-specific genes showed expression in all organisms. However, their level was higher in the sex group to which they correspond. In contrast, differential expression levels occurred in individuals with a shell length of 26–35 mm and 36–45 mm. These results suggest that sex gene expression is not entirely sex specific, and sexual differentiation in red abalone occurs between 16 and 25 mm of shell length. More studies are needed to determine the function of these genes in the sex that they are not associated with.     

Fibroblast growth factor-2 stimulates cell proliferation and decreases sexually dimorphic cell death in an avian song control nucleus

The neural system controlling song in birds has proven a useful model for investigating how neuronal growth and survival are regulated by sexual differentiation. The present study focused on one song control area, the robust nucleus of the archistriatum (RA), and explored how sex differences in the proliferation of putative glia cells in this region influence sexually dimorphic cell survival. In zebra finches (Poephila guttata), RA neuron death is much greater in young females than in males, resulting in marked sex differences in RA neuron number. An earlier study indicated that just prior to this sexually dimorphic neuron death the proliferation of putative glia cells within the RA is significantly lower in females than in males and remains so throughout the peak of neuron death. This suggests that sex differences in glia (or glia-derived molecules) might regulate neuron survival during sexual differentiation of the RA. To determine whether increased cell proliferation within the RA favors increased cell survival, we infused the potent glia mitogen fibroblast growth factor-2 (FGF-2) into the RA unilaterally in young females. We find that FGF-2 infusions increase RA cell proliferation and concurrently decrease the incidence of degenerating RA cells, results consistent with the hypothesis that glia exert neurotrophic effects on RA neurons during sexual differentiation. © 1998 John Wiley & Sons, Inc. J Neurobiol 37: 573–581, 1998     

Sex determination and evolution of unisexuality in the Conchostraca

Field collected or laboratory-reared samples of 60 species of conchostracans (representing all extant genera) indicate that males and females are equally common in most species. Deviations from this pattern occur in four lineages.Cyzicus andLeptestheria each include at least one unisexual species; many species of Limnadiinae are either unisexual or characterized by female-biased sex ratios; and Cyclestheriidae are either unisexual or express males in the later generations of their life cycles. Laboratory studies indicate that species with sex ratios near unity are gonochoric (obligately sexual), whereas females in species with female-biased sex ratios are capable of both outcrossing and selfing modes of reproduction. Phylogenetic analysis of patterns of reproduction suggest that sexual reproduction is the primitive condition. Genetic analysis of sexual species indicate that gender is determined by one or a few genetic factors and that the male-determining allele is recessive. The inheritance of gender in androdioecious species (where females are capable of self-fertilization) is similar to that in sexual species. Androdioecy is likely to be the intermediate stage between obligately sexual reproduction and unisexuality in the Limnadiinae. The phylogenetic distribution of sex ratio variation suggests that unisexuality in Cyzicidae, Leptestheriidae, and Cyclestheriidae has arisen independently of that in the Limnadiinae and that these cases have evolved by different evolutionary pathways.     

Sexual differentiation in three unconventional mammals: spotted hyenas, elephants and tammar wallabies

The present review explores sexual differentiation in three non-conventional species: the spotted hyena, the elephant and the tammar wallaby, selected because of the natural challenges they present for contemporary understanding of sexual differentiation. According to the prevailing view of mammalian sexual differentiation, originally proposed by Alfred Jost, secretion of androgen and anti-Mullerian hormone (AMH) by the fetal testes during critical stages of development accounts for the full range of sexually dimorphic urogenital traits observed at birth. Jost's concept was subsequently expanded to encompass sexual differentiation of the brain and behavior. Although the central focus of this review involves urogenital development, we assume that the novel mechanisms described in this article have potentially significant implications for sexual differentiation of brain and behavior, a transposition with precedent in the history of this field. Contrary to the "specific" requirements of Jost's formulation, female spotted hyenas and elephants initially develop male-type external genitalia prior to gonadal differentiation. In addition, the administration of anti-androgens to pregnant female spotted hyenas does not prevent the formation of a scrotum, pseudoscrotum, penis or penile clitoris in the offspring of treated females, although it is not yet clear whether the creation of masculine genitalia involves other steroids or whether there is a genetic mechanism bypassing a hormonal mediator. Wallabies, where sexual differentiation occurs in the pouch after birth, provide the most conclusive evidence for direct genetic control of sexual dimorphism, with the scrotum developing only in males and the pouch and mammary glands only in females, before differentiation of the gonads. The development of the pouch and mammary gland in females and the scrotum in males is controlled by genes on the X chromosome. In keeping with the "expanded" version of Jost's formulation, secretion of androgens by the fetal testes provides the best current account of a broad array of sex differences in reproductive morphology and endocrinology of the spotted hyena, and androgens are essential for development of the prostate and penis of the wallaby. But the essential circulating androgen in the male wallaby is 5alpha androstanediol, locally converted in target tissues to DHT, while in the pregnant female hyena, androstenedione, secreted by the maternal ovary, is converted by the placenta to testosterone (and estradiol) and transferred to the developing fetus. Testicular testosterone certainly seems to be responsible for the behavioral phenomenon of musth in male elephants. Both spotted hyenas and elephants display matrilineal social organization, and, in both species, female genital morphology requires feminine cooperation for successful copulation. We conclude that not all aspects of sexual differentiation have been delegated to testicular hormones in these mammals. In addition, we suggest that research on urogenital development in these non-traditional species directs attention to processes that may well be operating during the sexual differentiation of morphology and behavior in more common laboratory mammals, albeit in less dramatic fashion.     

Reproductive life history of the introduced peacock grouper Cephalopholis argus in Hawaii

This research investigated the reproductive biology (sex ratio, hermaphroditic pattern, size and age at maturity) of Cephalopholis argus, known locally in Hawaii by its Tahitian name roi. The results suggest that C. argus exhibits monandric protogyny (female gonad differentiation with female to male sex change) with females reaching sexual maturity at 1.2 years (95% c.i .: 0.6, 1.6) and 20.0 cm total length (L_T; 95% c.i .: 19.6, 21.2). The female to male sex ratio was 3.9:1. The average age and L_T at sex change was 11.5 years (95% c.i .: 11.1, 12.9) and 39.9 cm (95% c.i .: 39.5, 41.2), respectively. Current information on spawning seasonality of this species is incomplete, but based on the occurrence of spawning capable and actively spawning females, spawning probably takes place from May to October. Evidence of lunar spawning periodicity was found, with an increased proportion of spawning capable and actively spawning females, and an increased female gonado‐somatic index during first quarter and full‐moon phases. This information fills a valuable information gap in Hawaii and across the species' native range.     

Reproductive traits of the “ocellated killifish” Floridichthys polyommus Hubbs, 1936 (Pisces: Cyprinodontidae) inhabiting estuary of the Champotón River (Campeche,Mexico)

This study explores the reproductive biology and the influence of environmental factors on the reproductive traits of Floridichthys polyommus inhabiting the estuary of the Champoton River. The analysis included population structure, gonadal developmental stages, reproductive period, size at first sexual maturity, fecundity, sex ratio, and somatic indexes. A spatio‐temporal pattern in the distribution of the seven size classes was detected. The species showed a positive allometric growth. Sex ratio was 1.21:1 (females:males). Size at first maturity was 42.23 mm (females) and 47.8 mm (males). A higher absolute and relative fecundity was detected in Puente Champotón estuary (PCH) with lower salinities than those detected in the Delta (DE). The adaptive significance of this trait could be in response to the heavy osmotic constraint imposed by extreme salinities in the DE. Floridichthys polyommus prefers PCH for spawning, and as a result a higher number of ripe individuals was observed in this site, characterized by the highest nutrient levels, which can boost productivity and food availability for fish. Hepatosomatic index was negatively correlated with gonadosomatic index, suggesting energy transfer from the liver towards gamete production. Floridichthys polyommus exhibits an opportunistic strategy (early maturity, small oocytes, small clutches) suggesting that energy is allocated towards reproduction to compensate for the unpredictable hydrological conditions imposed by the estuary (particularly during the hurricane season). Floridichthys polyommus shows a reproductive strategy exhibited by fish living in unstable systems. A seasonal divergence was observed in the reproductive traits of F. polyommus, while differences between study sites were for fecundity.     

Juvenile bisexuality in the red sea bream,Pagrus major

The histology of the gonad of the red sea bream,Pagrus major, was examined in order to study the early gonadal development, sexual maturation and sex ratio in a natural population. A total of 1,117 fish between the ages of 4 months and 8 years were examined. Gonads of 4-month-old fish were either sexually undifferentiated with a central cavity, or ovarian in form. Gonads of 12- and 18-month-old fish were ovaries or bisexual gonads, while those of 2-year-old fish were ovaries, bisexual gonads or testes. Fish aged between 3 and 8 years had ovaries or testes, except for a few bisexual gonads found in 3- and 4-year-old fish. The chronological appearance of females, hermaphrodites and males in that order, and histological evidence, suggested that the testis originates from the ovary via a bisexual gonad in the juvenile stage. The sex ratio of females to males at the age of 2 years and over was about 1:1, suggesting that hermaphroditic red sea bream appear in about 50% of the juvenile population. The sexual pattern in this species, therefore, is concluded to be gonochorism with a bisexual juvenile stage.