The occurrence of serological H-Y antigen (Sxs antigen) in the diandric protogynous wrasse, Coris julis (L.) (Labridae, Teleostei)

The serological sex-specific (Sxs) antigen (previously called 'H-Y antigen') has been shown, in various vertebrate species ranging from fish to mammals, to be characteristic of the heterogametic sex. We studied a protogynous hermaphrodite, Coris julis, in order to examine whether the change of a female to a secondary male also involves a change in the Sxs-antigen phenotype. The (homogametic) females of this species were found to be Sxs negative, while both primary and secondary males were Sxs positive. This was true not only for gonads but also for nongonadal tissues. The administration of androgen to females is known to cause sex inversion in this species; we were able to demonstrate this again at the histological level, and found that androgen results in a Sxs positive phenotype in all tissues studied (gonads, spleen, muscle). We propose that androgen is responsible, directly or indirectly, for the occurrence of the Sxs antigen.     

Seasonal changes in sex ratio and size‐related sex reversal of the protogynous hermaphroditic bluespotted seabass Cephalopholis taeniops (Valenciennes, 1828)

Seasonal changes in sex ratio and size‐related sex reversal of the protogynous hermaphroditic Cephalopholis taeniops were studied from histological and population data of 218 individuals captured by hook and line, July 2009–November 2012, in Cape Verde archipelago. This study showed that C. taeniops have a diandric protogynous hermaphrodite sexual model, with young individuals undergoing bisexual development and hermaphrodites above 28 cm. All gonads had a bisexual immature stage with primary and secondary males. Primary males possibly originate from immature bisexual individuals, whereas secondary males likely result from females that have already reproduced and changed sex.     

Aspects of the reproductive biology of the grouper, Epinephelus tauvina (Forskål), in Kuwaiti waters

Gonad maturation in Epinephelus tauvina is examined over one annual reproductive cycle. The species has one major spawning period from April to May and this is associated with increasing water temperatures and relatively low salinity. The histological changes in the gonads in the female, male and transitional phase are examined, and oogenesis and spermatogenesis are described in five developmental stages. That E. tauvina is a protogynous hermaphrodite is confirmed, and sexual transition is found to occur in individuals 55–75 cm in length, and is related to spawning activity. Fecundity estimates for E. tauvina of length 35.1–62.3 cm ranged from 850 186 to 2 904 921.     

Evidence that estrogen regulates the sex change of honeycomb grouper (Epinephelus merra), a protogynous hermaphrodite fish

Circulating estradiol-17beta (E2) levels decrease precipitously during female to male (protogynous) sex change in fish. Whether this drop in E2 levels is a cause or consequence of sex change is still largely unknown. The present study treated adult female honeycomb groupers (Epinephelus merra) with aromatase inhibitor (AI, Fadrozole), either alone or in combination with E2, to investigate the role of estrogen in protogynous sex change. Control fish had ovaries undergoing active vitellogenesis; the gonads of AI-treated fish had already developed into testes, which produced sperm capable of fertilization. In contrast, co-treatment of fish with E2 completely blocked AI-induced sex reversal. AI treatment significantly reduced circulating levels of E2, whereas the addition of E2 to AI prevented the loss. The plasma androgen (testosterone and 11-ketotestosterone) levels were increased in the AI-treated fish, while the levels in the E2-supplemented fish were low compared to controls. Present results show that E2 plays an important role in maintaining female sex of hermaphrodite fishes, and that the inhibition of E2 synthesis causes oocyte degeneration leading to testicular differentiation in the ovary.     

Sex-inversion of the hermaphroditic, protogynous teleost Coris julis L. (Labridae)

Sex-inversion of Coris julis , a protogynous hermaphrodite, was studied using histological and cytological criteria. Four stages were recognized: oocyte and oogonial atresia; occurrence of spermatogonia with primordial germ cells (PGC) in the ovarian wall; onset of spermatogonial proliferation; considerable proliferation of spermatogonia and PGC with build-up of seminiferous tubules. Spermatogonia arise from PGC, undifferentiated and bipotential cells, in which mitotic activity was detected.     

Protogynous hermaphroditism in the parrotfish,Calotomus japonicus

Sexual patterns of the parrotfish,Calotomus japonicus, were studied using fifteen specimens collected in February and April, 1990, near Makurazaki, Kagoshima Prefecture. The males ranged from 265 to 345 mm SL, and were larger than the females on average. Most females ranged from 202 to 260 mm SL, but a female of 305 mm SL was obtained. Sexual dichromatism was conspicuous. Histological examination of the gonads and low values of gonado-somatic indices of both males and females indicated that they were not in spawning season. It was concluded that C.japonicus is a protogynous hermaphrodite, because all male specimens had secondary testes. It was pointed out that the present species has a weak size-sex relationship. Because the specimens examined did not include primary males, it is suggested that primary males are relatively rare, if they occur at all.     

Induction of female-to-male sex change in the honeycomb grouper (Epinephelus merra) by 11-ketotestosterone treatments

The honeycomb grouper, Epinephelus merra, is a protogynous hermaphrodite fish. Sex steroid hormones play key roles in sex change of this species. A significant drop in endogenous estradiol-17beta (E2) levels alone triggers female-to-male sex change, and the subsequent elevation of 11-ketotestosterone (11KT) levels correlates with the progression of spermatogenesis. To elucidate the role of an androgen in sex change, we attempted to induce female-to-male sex change by exogenous 11KT treatments. The 75-day 11KT treatment caused 100% masculinization of pre-spawning females. Ovaries of the control (vehicle-treated) fish had oocytes at various stages of oogenesis, while the gonads of the 11KT-treated fish had transformed into testes; these contained spermatogenic germ cells at various stages, including an accumulation of spermatozoa in the sperm duct. In the sex-changed fish, plasma levels of E2 were significantly low, while both testosterone (T) and 11KT were significantly increased. Our results suggest that 11KT plays an important role in sex change in the honeycomb grouper. Whether the mechanism of 11KT-induced female-to-male sex change acts through direct stimulation of spermatogenesis in the ovary or via the inhibition of estrogen synthesis remains to be clarified.     

Gonadal development and an indication of functional protogyny in the Indian damselfish (Dascyllus carneus)

The objective of this study was to determine the sexual pattern of the Indian dascyllus Dascyllus carneus . After an initially undifferentiated state, gonads of D. carneus developed an ovarian lumen and primary growth stage oocytes, and subsequently cortical-alveolus stage oocytes. From ovaries with cortical-alveolus stage oocytes and from more developed ovaries, some gonads redifferentiated into testes. From a sample of 163 individuals, two had a gonad containing degenerating vitellogenic oocytes and proliferating spermatogenic tissue, nine had a gonad containing degenerating cortical-alveolus stage oocytes and spermatogenic tissue, and five had a gonad with degenerating primary growth stage oocytes and spermatogenic tissue. The size of these individuals overlapped greatly with the size range of mature females, suggesting that at least in some individuals, redifferentiation toward a testis occurred after spawning as females. This indicates that D. carneus is a functional, diandric protogynous hermaphrodite. Removal of a dominant male(s) did not induce a sex change in any of the ranking females in the laboratory and field groups. There was no difference in the number of chases and signal jumps performed by the ranking female between control and experimental field groups, or before and after removal of the male. However, the sizes of the ranking females were at or beyond the size range of individuals with a mixed-stage gonad, suggesting that the developmental window for female-to-male sex change may not be open ended. In 41 of 43 field groups, in which sex of fish was determined histologically or by the shape of the urogenital papilla, one to several highest size ranks were occupied by males, followed by one to several females. Mature males, however, were not limited to the highest ranks and occurred at various lower size ranks within groups. Individuals with a mixed-stage gonad also occupied various size ranks within groups.     

Socially controlled male-to-female sex reversal in the protogynous orange-spotted grouper,Epinephelus coioides

Socially controlled sex change in teleosts is a dramatic example of adaptive reproductive plasticity. In many cases, the occurrence of sex change is triggered by a change in the social context, such as the disappearance of the dominant individual. The orange-spotted grouper Epinephelus coioides is a typical protogynous hermaphrodite fish that changes sex from female to male and remains male throughout its life span. In this study, male-to-female sex reversal in male Epinephelus coioides was successfully induced by social isolation. The body length and mass, gonadal change, serum sex steroid hormone levels and sex-related gene expression patterns during the process of socially controlled male-to-female sex reversal in E. coioides were systematically examined. This report investigates the physiological mechanisms of the socially controlled male-to-female sex reversal process in a protogynous hermaphrodite grouper species. The results enable us to study the physiological control of sex change, not only from female to male, but also from male to female.     

Gonad development and evidence of protogyny in the red-throat emperor on the Great Barrier Reef

The gonad development in the red-throat emperor Lethrinus miniatus is described and the first detailed evidence for protogyny in this species provided. The identification of transitional individuals, bimodal sex-specific size-frequency distributions and female biased sex ratios suggest that L. miniatus is most likely a protogynous hermaphrodite. Transitional L. miniatus gonads were characterized by the concurrent degeneration of all oocytes and the proliferation of spermatocysts near the edge of the lamellae, an increase in blood vessels along strands of stromal tissue within the lamellae and the formation of multiple sperm sinuses. The sites of oocyte degeneration and proliferation of spermatocysts were spatially segregated. An increase in blood vessels along strands of stromal tissue within the lamellae of transitional phase gonads is likely to assist in the breakdown of oocytes and the proliferation of spermatocysts. Most mature resting females containing spermatocysts occurred within the transitional size-frequency distribution, suggesting that the presence of spermatocysts in these females may be an early sign of sex change. Oocytes within female gonads were interrupted by filamentous strands of stromal tissue within the lamellae. The testis contained a remanent ovarian lumen but no residual oocytes. Three characteristics of transitional L. miniatus gonads were found to be unusual and described for few other species of coral reef fishes. These included the absence of oocytes within testes, increased numbers of blood vessels, and the presence of strands of stromal tissue within the lamellae.     

Ultrastructural study of sex inversion in a protogynous hermaphrodite, Epinephelus microdon (Teleostei, Serranidae).

Sex inversion of Epinephelus microdon, a protogynous hermaphrodite was studied using light and electron microscopic criteria. The sex changes takes place within ex-ovarian lamellae in which, in addition to spermatogonia, numerous primordial germ cells (PGCs) were detected. These undifferentiated and bipotential early germ cells are involved, as well as spermatogonia, in the building up of the testis.     

Growth and reproductive biology of the foxfish Bodianus frenchii,a very long‐lived and monandric protogynous hermaphroditic labrid

Samples of the foxfish Bodianus frenchii, collected over reefs on the lower west and south coasts of Western Australia, contained individuals ranging up to 78 years old. Although B. frenchii is far smaller than many other species within the Labridae, its maximum age is the greatest yet recorded for this highly speciose family and, together with Achoerodus gouldii, provides an example of a temperate hypsigenyine with exceptional longevity. Length and age compositions of females and males and the histological characteristics of gonads of a wide length range of individuals demonstrated that B. frenchii is a protogynous hermaphrodite. Furthermore, as, on both coasts, the length of the smallest male was greater than that at which all females had become mature, B. frenchii is a monandric protogynous hermaphrodite, i.e. all of its males are derived from functional females. Attainment of maturity by females is related more to length than age, whereas the reverse is true for sex change. On the basis of Schnute growth equations and length‐to‐body mass regression equations, the predicted length at age and body mass at length of fish on the south coast were greater than those on the west coast throughout life. Although B. frenchii spawns daily during the main spawning season, which extends from October to February on both coasts, its fecundity at any given length is substantially greater on the south than on the west coast. The more rapid growth of juveniles and earlier attainment of maturity by B. frenchii on the south coast than on the warmer west coast, together with maturation at a similar size on both coasts, run counter to the trends observed in many species and certain ecological theories regarding the relationships between life‐cycle traits and latitude and temperature. The attainment by B. frenchii of a larger body length at age, of greater body mass at length and of greater fecundity at both length and body mass in fish on the south than on the west coast strongly suggests that conditions on the former, cooler coast are more favourable for this labrid, which belongs to a sub‐genus whose other species typically live in cool, deep, temperate waters.     

Habitat Patch Size,Facultative Monogamy and Sex Change in a Coral-dwelling Fish, <Emphasis Type="Italic">Caracanthus unipinna</Emphasis>

Synopsis We investigated the inter-relationships between coral colony size, social group size, mating system, and patterns of sex allocation in the pygmy coral croucher, Caracanthus unipinna (Caracanthidae), an obligate coral-dwelling fish. Histological examination of the gonads from all individuals in social groups revealed that the predominant mating system was harem polygyny. However, both group size and mating system co-varied with coral colony size, with pair forming and monogamy occurring on small corals and group forming and harem polygyny on large corals. This species therefore displays mating system plasticity in response to varying habitat patch size. Within-group sexual size dimorphism and individual gonad structure indicate that C. unipinna is also likely to be a protogynous hermaphrodite. These social and reproductive features of C. unipinna contrast with some other coral-dwelling fishes, which display a lack of social and mating system plasticity in response to habitat patch size, and either bi-directional or protandrous sex change. Possible reasons for this dichotomy include differences in spawning mode, parental care and levels of intrasexual aggression.     

Growth and sexual strategies in the fishThalassoma duperrey (Labridae), a protogynous hermaphrodite

Synopsis Adults of the fishThalassoma duperrey, a protogynous hermaphrodite, were collected and growth observed in captivity to study the relationship between growth and reproduction among primary males, females, and secondary males. Sexual maturity is reached at about 60 mm standard length, probably less than 1 year after fertilization. Gonosomatic index in both males and females peaks at about 120 mm, nearly 2 years later. Shortly thereafter females typically change sex, and both primary and secondary males undergo color change. At the same time gonosomatic index falls abruptly and remains low in large fish. The above changes appear to reflect differences in reproductive effort over a lifetime and are interpreted as the optimum strategy given the social and mating system of this fish. Unless they cannot acquire enough food to develop large gonads, small individuals put a much greater proportion of energy into growth than reproduction apparently to minimize the period of low fitness. Intermediate-sized males and females generally invest heavily in gametes, though some retain small gonads. Large individuals (both primary and secondary males) greatly reduce their investment in gametes, probably trading the energy required to maintain reproductive territories for it. This kind of gonad ontogeny involving gonad regression, as inT. duperrey and other labrid fishes, is unique among vertebrates.     

Reproduction of the inshore coral trout Plectropomus maculates (Perciformes: Serranidae) from the Central Great Barrier Reef, Australia

The reproductive biology of the serranid fish Plectropomus maculatus sampled from inshore waters of the Central Great Barrier Reef was studied based on histological analyses of gonad material. This species was shown to be a monandric protogynous hermaphrodite. The process of sex change foliowed the spawning period observed during September through November. Plectropomus: maculatus showed multiple spawning during this period. Sex change followed the usual protogynous mode with degeneration of ovarian germinal tissue accompanied by proliferation of male germinal tissue in the gonad. The sex structure of the sampled population was analysed based on age and size information. The size and age of first reproduction for females was 30.0cm s. L. and 2 years of age. The size and age of sex-transition was 35.4 cm s. L. and 4.4 years of age. The sex/size and sex/age relationships indicated that sex-change can occur over a broad range of sizes and ages. The sizes and age distributions of males and females P. maculates overlapped over 38% of the length range and over 42% of the maximum age observed.     

Preliminary observations on the life history of the white-streaked grouper, <Emphasis Type="Italic">Epinephelus ongus</Emphasis>, from Okinawa,Japan

A preliminary analysis of 175 specimens of the white-streaked grouper, Epinephelus ongus (Serranidae), was undertaken to determine life history characteristics of the species. Sagittal otoliths, stomachs, and a subsample of gonads were removed to determine age at length, diet, and reproductive strategy. The von Bertalanffy growth equation was used to describe growth in this species and yielded the growth parameters L^∞ = 438.3, K = 0.04334, and t₀ = −8.752. Fish ranged in age from 1 to 20 years. Diet was consistent with other serranid species and included crabs, shrimps, octopi, and fishes. Based on a very limited number of specimens (n = 12), the larger size and older age of males compared to females suggests that E. ongus may be a protogynous hermaphrodite.     

Increased H-Y antigen levels associated with behaviorally induced, female-to-male sex reversal in a coral-reef fish

It has been proposed that H-Y antigen is the synthetic product of sex-determining genes, and that H-Y antigen controls ontogenetic differentiation of the heterogametic sex throughout vertebrates. The coral-reef fish Anthias squamipinnis is a protogynous hermaphrodite in which all individuals mature initially as females. Males result when adult females change sex as a consequence of alterations in behavioral interactions within social groups. Three assay methods were used to measure H-Y antigen levels in the spleens, gonads, and epidermal tissue of 16 adult females and in 16 males that had been induced to change sex from a prior female phase by the removal of a pre-existing male from each of 16 social groups. In 15 male-female pairs, the H-Y antigen levels were higher in male than in female spleen, gonad, and epidermis tissues. The precise temporal relationship between the onset of sex change and the increase in the H-Y antigen level was not examined. If, as we strongly suspect, the temporal relationship proves to be close, the inference will be that the behavioral cues inducing sex change also influence the synthetic activity of genes controlling H-Y antigen production.