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.     

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.     

Induction of meiosis in fetal mouse testis in vitro

Fetal mouse testes and ovaries with their urogenital connections were cultured singly or in pairs on Nuclepore filters. When a testis in which the sex was not yet morphologically detectable was cultured together with older ovaries containing germ cells which were progressing through the meiotic prophase, the male germ cells were triggered to enter meiosis. When older fetal testes in which the testicular cords have developed were cultured together with ovaries of the same age with germ cells in meiosis, the oocytes were prevented from reaching diplotene stage. It was concluded that the fetal male and female gonads secrete diffusable substances which influence germ cell differentiation. The male gonad secretes a "meiosis-preventing substance" (MPS) which can arrest the female germ cells within the meiotic prophase. The female gonad secretes a "meiosis-inducing substance" (MIS) which can trigger the nondifferentiated male germ cells to enter meiosis.     

Gonadal development and non-functional protogyny in a coral-reef damselfish, Dascyllus albisella Gill

All gonads of the Hawaiian dascyllus Dascyllus albisella , irrespective of the final sex of individuals, developed an ovarian lumen and primary-growth-stage oocytes after an initially undifferentiated state. From this ovarian state or from more differentiated ovaries, some gonads redifferentiated into testes. None of 117 individuals examined had a gonad containing degenerating vitellogenic oocytes and proliferating spermatogenic tissue. Eleven individuals had gonads containing degenerating cortical-alveolus-stage oocytes and developing spermatogenic tissue. The size of these individuals overlapped with the female size range in which the majority of the females were still in the middle of the maturation process. They were absent from the larger size range where the majority of females had vitellogenic oocytes. This indicated that the transition toward maleness is likely to have occurred after the onset of cortical-alveolus stage, but before final oocyte maturation and spawning as females. Therefore the protogynous pattern of gonadal development was non-functional. There was no dimorphism in the sperm duct configuration, and all the testes were secondary testes reported for diandric, protogynous species with undelimited gonads. Very early development of an ovarian lumen appeared to have resulted in a secondary-male configuration in all testes, although redifferentiation into males appeared to have occurred before sexual maturity and spawning as females.     

Gonadal development and differentiation in Alligator mississippiensis at male and female producing incubation temperatures

The development and differentiation of the gonads of embryonic alligators incubated at 30 °C (100% female producing) and 33 °C (100% male producing) was investigated histologically. The stage of development of the gonad and differentiation into an ovary or a testis occurred at essentially the same time at both temperatures. This contrasts with the overall development of the embryos which was slower at the lower temperature. A few days prior to differentiation, gonads grew more quickly at 33 °C than they did at 30 °C. However, once differentiated into a presumptive testis, gonads reduced in volume so that at hatching presumptive testes were smaller than presumptive ovaries. It is hypothesized that synchrony/asynchrony of development of the gonad and the rest of the embryo may account for temperature-dependent sex determination.     

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.     

Gonadal development in T16H/XSxr hermaphrodite mice

Sexual development was studied in 25 hermaphrodite mice with the genotype T16H/XSxr. The majority of the animals had a male phenotype similar to that seen in XXSxr males. A few, however, had feminized external genitalia and were classified as females. Examination of the gonads and reproductive tracts of the male hermaphrodites revealed a strong tendency for the left gonad to be more masculine than the right. Most of the gonads in male and female hermaphrodites appeared to be ovaries or testes rather than ovotestes.     

Cytomorphology of female and male gonads in Antarctic toothfish <Emphasis Type="Italic">Dissosctichus mawsoni</Emphasis> (Nototheniidae) from the Ross Sea in the summer period

Results of histological analysis of gonads of female and male Antarctic toothfish Dissostichus mawsoni caught in the Ross Sea of the Pacific sector in the period of Antarctic summer (December–February) 2004–2005 are presented. Morphological indices and index of gonad maturity are described and ecological criteria of assessment of maturity stages of ovaries, cytological indices of oocytes, and the type of toothfish oogenesis are determined. It was established that in the period of fishing Antarctic toothfish, females and males with gonads at maturity stage III dominate. Ovaries contain two groups of oocytes of the period of trophoplasmatic growth and large oocytes of the nearest spawning season that comprise the smallest proportion of total sex cells. In the testes of two male toothfish, the primary fusion of renal and generative tissues was revealed. It is suggested that the termination of gonad maturity of toothfish takes place from March to April, and spawning takes place from June to August.     

Prey resources before spawning influence gonadal investment of female, but not male, white crappie

In this study, an outdoor pool experiment was used to evaluate the effect of prey resources during 4 months before spawning on the gonadal investments of male and female white crappie Pomoxis annularis , a popular freshwater sportfish that exhibits erratic recruitment. Fish were assigned one of three feeding treatments: starved, fed once every 5 days (intermediate) or fed daily (high). All measurements of male testes ( i.e. wet mass, energy density and spermatocrit) were similar across treatments. Conversely, high-fed females produced larger ovaries than those of intermediate-fed and starved fish, and invested more energy in their ovaries than starved fish. Compared to pre-experiment fish, starved and intermediate-fed females appeared to increase their ovary size by relying on liver energy stores ('capital' spawning). Conversely, high-fed females increased liver and gonad mass, implying an 'income'-spawning strategy (where gonads are built from recently acquired energy). Fecundity did not differ among treatments, but high-fed fish built larger eggs than those starved. Females rarely 'skipped' spawning opportunities when prey resources were low, as only 8% of starved females and 8% of intermediate-fed females lacked vitellogenic eggs. These results suggest that limited prey resources during the months before spawning can limit ovary production, which, in turn, can limit reproductive success of white crappies.     

Hermaphroditism and sexual function inCirrhitichthys aureus and the other Japanese hawkfishes (Cirrhitidae: Teleostei)

The present paper deals with gonadal formation, sex succession and sexual function in the Japanese hawkfishes,Cirrhitichthys aureus (Temminck et Schlegel),C. aprinus (Cuvier),C. falco Randall,Cirrhitops hubbardi (Schultz) andCyprinocirrhites polyactis (Bleeker). Detailed studies were made forC. aureus collected from Suruga Bay, central Japan. In this species the gonad of a young fish of 21.8 mm SL had begun to initially differentiate into an ovary forming an early ovarian cavity of the parovarian type. The gonadal structures of 63 further specimens ofC. aureus, ranging from 23.9 to 114.3 mm SL, could be separated into three categories: viz., ovaries (7), ambisexual gonads (54) and secondary testes (2). Reproduction ofC. aureus in Suruga Bay took place from June to November. Although ovaries which appeared throughout the year presented only in the immature stage and occurred only in small fish ranging from 25.1 to 46.6 mm SL, functional gonads occurred in 32 ambisexual fish ranging from 61.5 to 113.5 mm SL and in one secondary male of 101.5 mm SL, collected during the reproductive season. The smaller ambisexual fish, ranging from 61.5 to 92.0 mm SL, showed active oogenetic processes in the major ovarian zones with many ripened eggs ovulating into the ovarian cavity. These gynogenic gonads undoubtedly functioned as female; however, distinct meiosis proceeded partially in the minor testicular zones with a few spermatozoa occurring in the vas deferens. The larger ambisexual specimens, ranging from 71.5 to 113.5mm SL, had slender androgenic gonads, which showed active spermatogenesis in various testicular zones and which might function as males with reduced ovigerous lamellae. The secondary males had testes occupied by seminal lobules, but which retained the ovarian cavity within. According to aquarium rearing experiments, in a pair ofC. aureus twice crossed mutually after spawning, functional sex changes occurred in twice, opposite directions, in a single gonad, i.e., from female to male and from male to female. These gonadal aspects in captiveC. aureus seemed to show an intermediate phenomenon between consecutive and simultaneous hermaphroditism. This finding also suggested thatC. aureus had similar sex succession characteristics to those of the serranid,Serranus fasciatus, studied by Hasting and Petersen (1986) and the gobiid,Trimma okinawae, studied by Sunobe (1990). We also established that normal functional hermaphroditism occurred in the four other cirrhitid species, which showed almost identical gonadal characteristics toC. aureus.     

Morphological development of the gonads in zebrafish

Gonadogenesis of zebrafish Danio rerio was investigated by means of light microscopy to test the suitability of gonad histology as an endpoint in hazard assessment of endocrine‐active compounds. At age 2 weeks post‐fertilization (pf), primordial germ cells were found in a dorsocaudal position in the body cavity. At 4 weeks pf, the majority of the fish (86%) possessed paired gonads with meiotic germ cells; these gonads represented presumptive ovaries. At week 5 pf, 87% of the fish examined had ovaries with perinucleolar oocytes. Further development of the gonads in female zebrafish up to week 11 pf was characterized by an increase in gonad size as well as in the number and size of perinucleolar oocytes. Starting with week 5, some fish showed alterations of gonad morphology, including a decrease in the number and size of the oocytes, an enhanced basophilia and irregular shape of the oocytes, and finally their degeneration into residual bodies. With the decline in oocyte number, stromal cells became more numerous and they infiltrated the gonadal matrix. In several 7 week‐old zebrafish with altered gonadal morphology, enhanced numbers of gonial cells arranged in cyst‐like groups appeared. These gonads were interpreted as presumptive testes. In one fish out of 32 individuals examined, spermatocytes were detected, in addition to the gonial cells. During the subsequent weeks, the percentage of fish showing early testes with spermatogonia, spermatocytes and spermatids increased and reached 40% at 11 weeks pf. The sequence of gonadal alterations taking place in some of the individuals from week 5 pf onwards was interpreted to reflect the transition of protogynic ovaries into testes. The developmental pattern described identifies zebrafish to be a juvenile hermaphrodite. The results of this study are of relevance for the use of gonadal histopathology as endpoint in endocrine disruption testing, particularly in order to avoid false diagnoses of ‘intersex gonads’ in zebrafish.     

Gonadal development and diandric protogyny in two populations of Dascyllus reticulatus from Madang, Papua New Guinea

Two Dascyllus reticulatus populations from Madang, Papua New Guinea exhibited diandric protogyny. In both populations, gonads began as undifferentiated, and then developed oocytes in the primary growth stage and an ovarian lumen. From this ovarian state or from more developed ovaries containing oocytes beyond the primary‐growth stage, some gonads developed into testes. The first sign of testicular development was degeneration of oocytes, degeneration of oocytes in the primary growth stage in ovarian gonads and degeneration of oocytes of all growth stages present including the primary growth stage in ovaries, which was then followed by development of spermatogenic tissue. In both populations, most of the fish that had gonads with degenerating oocytes were smaller than the smallest mature females, indicating that development towards testes was mostly initiated in immature gonads containing only pre‐vitellogenic oocytes. On some occasions, however, females as large as other mature females also had gonads with degenerating oocytes, suggesting that development towards testes may have occurred in mature ovaries as well. This latter notion is further strengthened by the discovery of a fish having a gonad that contained both degenerating vitellogenic oocytes and developing spermatogenic tissue. Taken together, these results suggest that D. reticulatus can exhibit diandric protogyny, because testes in D. reticulatus developed from juvenile gonads as well as from mature ovaries.     

Gonadal maturation is dependent on body size in the sea urchin,Echinometra tsumajiroi

Summary

This study examined gonadal maturation in the sea urchin, Echinometra tsumajiroi, in relationship to body size. In the breeding season, E. tsumajiroi were collected and divided into six groups based on the length of the major axis (LM) as a measure of body size: Group 1 (6–10 mm in LM), Group 2 (11–15 mm in LM), Group 3 (16–20 mm in LM), Group 4 (21–25 mm in LM), Group 5 (26–30 mm in LM), and Group 6 (> 31 mm in LM). Although gonads were present in individuals larger than 8 mm in LM, the germ cells were undifferentiated in Group 1. Gonadal sex differentiation was first seen in Group 2: a few young oocytes appeared at the periphery of the ovary, and both spermatogonia and spermatocytes were also observed at the periphery of the testis. As size increased from Group 3 to Group 4, more mature germ cells such as ripe ova or spermatozoa were observed in ovaries and testes. In Group 5, the ovaries and testes were filled with a large number of mature eggs and spermatozoa, respectively. Spawning could be induced in this group by the injection of 0.5 M KCl into the coelomic cavity. In contrast, gonads of some of the biggest individuals, belonging to Group 6, appeared to be spent due to spawning, because few eggs were present in the ovary and relict spermatozoa were present in the testis. These results suggest that reproductive activity in the sea urchin, E. tsumajiroi, depends on body size.     

Gonadal sexual differentiation of European sea bass (Dicentrarchus labrax,L. 1758) of fingerlings in different size classes

This report describes a study of the gonad differentiation in fingerlings of European sea bass (Dicentrarchus labrax) belonging to different size groups. Fishes were divided into four groups according to their weight: Group 1 (4.5 ± 0.7 g), Group 2 (9.2 ± 0.8 g), Group 3 (14.8 ± 1.8 g), and Group 4 (21.8 ± 2.3 g). In all groups, low percentages of undifferentiated or early sexually differentiated fish were found. Higher percentages of fully differentiated ovaries were found in Groups 1 and 3. Fully differentiated testes occurred in 90% of Group 4. Ninety percent of fish in Group 2 had gonads with intratesticular oocytes. This highlights the occurrence of intersexual cases and suggests that at this size, the fish is susceptible to phenotypic plasticity. The results of the present study show that sexual differentiation of the gonads is found in fish at 4.5 g weight. The ambiguous relationship between the observed gonad morpho-functional characteristics and fish size suggests a subject-dependent sexual differentiation.     

Identification of key nutrients for gonadal development by comparative analysis of proximate composition and fatty/amino acid profile in tissues and eggs of Chinese sturgeon (Acipenser sinensis Gray, 1835)

The present study was conducted to characterize the proximate composition and fatty/amino acid profile in muscle, liver and gonads obtained from farmed male and female Chinese sturgeon (Acipenser sinensis) with gonad development at stage II, and to discern the key nutrients for gonads by comparing the nutrients between tissues and between fish at different gonad stages. Chinese sturgeon were reared in an outdoor circular cement pool at temperatures ranging from 8°C to 26°C, and fed with 70% artificial diets and 30% frozen forage fish at a feeding rate of 0.2%–1.5% body weight according to the water temperature. Three male and three female A. sinensis were sampled for analyses. Results showed that the lipid content in gonads was significantly higher than in other tissues in both males and females (p < .05). The protein content in male and female gonads was significantly lower than that in the muscle (p < .05). Total content of mono‐unsaturated fatty acids (MUFA) in females was significantly higher than in males (p < .05), while the total content of n‐6 PUFA in females was significantly lower than in males (p < .05). Palmitic acid (C16:0) and oleic acid (C18:1n9) were the major saturated fatty acid (SFA) and MUFA, respectively, in both females and males. The contents of C18:2n6, C20:4n6 (ARA), C18:3n3 and C22:6n3 (DHA) in male gonads were significantly higher than in females (p < .05). The contents of ARA, EPA and DHA in gonads were lower, possibly playing a pivotal role in gonad development. Comparing the nutrients of ovaries in broodstocks at stages III and IV and the eggs at stage IV, the present study suggests that the lipid and fatty acid profile in fish is vital to the gonad development and that proper improvement of lipid nutrition in the broodstock diet to provide sufficient energy and essential fatty acids would be beneficial for the gonad development of Acipenser sinensis.     

Accumulation of the major yolk protein and zinc in the agametogenic sea urchin gonad

Sea urchins of both sexes store the nutrients necessary for gametogenesis in nutritive phagocytes of the agametogenic gonad. A zinc-binding protein termed the major yolk protein (MYP) is stored here as two isoforms: the egg-type (predominant in egg yolk granules) and the coelomic fluid-type (a precursor with greater zinc-binding capacity). MYP is used during gametogenesis as material for synthesizing gametic proteins and other components. We investigated its accumulation and relationship to zinc contents in gonads during the non-reproductive season in Pseudocentrotus depressus. MYP constituted most of the protein in coelomic fluid and gonads. Both ovaries and testes grew gradually, accumulating MYP and zinc during the year. Total zinc contents and the ratio of coelomic fluid-type to egg-type protein were higher in ovaries than in testes as gametogenesis approached. Most of the zinc in the coelomic fluid was bound to MYP, and the concentrations of MYP and zinc were elevated toward the onset of oogenesis in the female coelomic fluid. Thus, MYP accumulates in the agametogenic ovaries and testes during the non-reproductive season, playing a role as a carrier to transport zinc to the gonad. Transportation of zinc by MYP is more active in females than in males.     

Histological and ultrastructural investigation of early gonad development and sex differentiation in Adriatic sturgeon (Acipenser naccarii,Acipenseriformes, Chondrostei)

Gonad development and sex differentiation from embryos to 594‐day‐old individuals were investigated in farmed Acipenser naccarii using light and transmission electron microscopy. The migrating primordial germ cells first appear along the dorsal wall of the body cavity in embryos 1.5 days before hatching. The gonadal ridge, containing a few primary primordial germ cells (PGC‐1) surrounded by enveloping cells, appears in 16‐day‐old larvae. At 60 days, the undifferentiated gonad is lamellar and PGC‐1 multiply, producing PGC‐2. In 105‐day‐old juveniles, a distinct germinal area with advanced PGC‐2 appears on the lateral side near the mesogonium and the first blood vessels are visible. At 180 days, putative ovaries with a notched gonadal epithelium and putative testes with a smooth one appear, together with adipose tissue on the distal side. In 210‐day‐old juveniles, active proliferation of germ cells begins in the putative ovaries, whereas putative testes still contain only a few germ cells. The onset of meiosis and reorganization of stromal tissue occurs in ovaries of 292‐day‐old individuals. Ovaries with developed lamellae enclosing early oocyte clusters and follicles with perinucleolar oocytes occur at 594 days. Meiotic stages are never found, even in anastomozing tubular testes of 594‐day‐old individuals. Steroid producing cells are detected in the undifferentiated gonad and in the differentiated ones of both sexes. Anatomical differentiation of the gonad precedes cytological differentiation and female differentiation largely precedes that of the male. Gonad development and differentiation are also associated with structural changes of connective tissue, viz. collagen‐rich areas are massive in developing testes and reduced in ovaries. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.