Reproduction in the dusky grouper from the southern Mediterranean

Demographic data and gonad histology confirmed that the dusky grouper Epinephelus marginatus is a protogynous hermaphrodite that follows a monandric pathway to sexual development. Females reached first sexual maturity at 36·7 cm L_s and estimated mean length at first maturity (L₅₀) was 43·8 cm L_s for females and 81·3 cm L_s for males. Adult sex ratios during the reproductive period were c. 3·5: 1 females to males. Females exhibited group-synchronous ovarian development and multiple ovulation occurred over the spawning period. Gonads were ripe from early May and spawning occurred from June until early September. The size of ripe testes (0·6% W )indicated strong oligospermy and suggested a mating system with no sperm competition. Sexual transition was protogynous involving regression of ovarian tissue and proliferation of testicular tissue in the gonads. Transitional individuals occurred from May through November and accounted for 9% of sampled adult population. Sex change occurred in fish 69–93 cm (L_s) long and the size distributions of males and females overlapped over 27% of the L_s range. Special zones were recognized as gathering areas for sexually mature dusky groupers during the reproductive period.     

Protogynous hermaphroditism in the half-banded sea perch, Hypoplectrodes maccullochi (Serranidae)

The half-banded sea perch, Hypoplectrodes maccullochi (Serranidae) exhibits protogynous hermaphroditism, based on the presence of transitional individuals collected from rocky reefs along the coast of New South Wales, Australia. Males dominated the larger size and age classes, whereas females predominated the smaller size and age groups. Histological preparations suggested that sex change occurs between 55–85 mm s.l. and 1 + to 4+ years of age. Three transitional phases were identified which varied according to the proportion of gametogenic tissue and remnant oocytes present within their gonads. All individuals greater than 85 mm s.l. were functional males containing well defined crypts of spermatids and spermatozoa. Remnant oocytes within these testes indicated prior female function. H. maccullochi were sexually monochromatic. The occurrence of some small males derived from pre-spawning females suggested pre-maturational sex change occurred in some individuals. Sex changing individuals were found in all the months of the year that gonads were examined in detail (January, February, April and June 1989). Fish were ripe in August and November. H. maccullochi conforms with the pattern of reproduction evident within the genus Hypopleclrodes .     

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.     

Reproductive biology and implications for management of the spangled emperor Lethrinus nebulosus in the southern Arabian Gulf

The reproductive biology of the spangled emperor Lethrinus nebulosus in the southern Arabian Gulf was determined from the examination of 781 individuals collected between September 2008 and August 2009. There was no histological evidence of adult sex change, and sex ratios did not differ significantly from unity across all size and age classes. Testes had an ovarian structure and a remnant lumen which was not used for sperm transport; furthermore, residual oocytes were observed in the testes of some immature, resting and developing males. A dramatic change in the sex ratio of the smallest and youngest size and age classes suggested that juvenile female to male sex change occurred between 21·0 cm fork length (L(F)) and 24·0 cm L(F) at 1 year of age. The combination of histological evidence with the sexual composition of the size and age structures suggests a non-functional protogynous hermaphroditic sexual pattern, which is analogous to functional gonochorism. The spawning season was well defined, occurring once a year during March, April and early May. Peaks in spawning occurred after full moons, there was a cessation in spawning activity after new moons and spawning was completed within three lunar cycles. The distribution of males over the entire size and age ranges and the absence of inactive mature females during the spawning season suggested that the population was not constrained by sperm limitation. Size-specific and age-specific reproductive potential indicated that conventional regulations that equate the mean size at first capture to sexual maturation are unsuitable for the management of L. nebulosus. The maximum recorded age (11 years), small mean size and young age at sexual maturation (L(m50) = 26·7 cm L(F), 2·1 years, for females and 19·4 cm L(F), 0·5 years, for males) may be a direct result of intensive demersal fishing in the southern Arabian Gulf.     

Morphological ontogeny of the gonad of three plectropomid species through sex differentiation and transition

The gonadal ontogeny through sex differentiation and transition of three protogynous coral trout species, Plectropomus leopardus , P. maculatus and P. laevis was described, based on anatomical and germinal differences along the length of the reproductive tract. Gonads of immature and mature females, sex changing individuals (transitionals) and males were examined. Specific anatomical features that were compared between sexual phases included the presence and structure of sperm sinuses, gonadal musculature and germinal cell types. All three coral trout species first differentiated as an immature female. The sexual pattern of P. leopardus and P. maculatus was concluded to be diandric protogynous hermaphroditism (males were derived from the juvenile phase as well as through sex change of mature females). Plectropomus laevis was found to be monandric as males were only derived through sex change in mature females. Structural changes did not occur concomitantly with the germinal changes associated with sex change in these Plectropomus species, which is atypical for protogynous species described to date. Precursory sperm sinuses in the dorso-medial region of the gonad were present, although non-functional, in a proportion of immature and mature females of all three species. These proportions, however, varied between species depending on the sexual pattern. The structural and germinal changes observed were hypothesized as anatomical adaptations that aid in minimizing time spent in the (non-reproductive) sexual transition phase and maximizing flexibility in male development in the diandric species.     

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.     

Aromatase inhibitor induces complete sex change in the protogynous honeycomb grouper (Epinephelus merra)

The protogynous hermaphrodite fish change sex from female to male at the certain stages of life cycle. The endocrine mechanisms involved in gonadal restructuring throughout protogynous sex change are not clearly understood. In the present study, we implanted maturing female honeycomb groupers with nonsteroidal aromatase inhibitor (AI), Fadrozole (0, 1, and 10 mg/fish) and examined changes in gonadal structures and serum levels of sex steroid hormones 2(1/2) months after implantation. The ovaries of control females had oocytes undergoing active vitellogenesis, whereas AI caused females to develop into functional males. These males had testes, which were indistinguishable in structure from those of normal males, but bigger in size, and completed all stages of spermatogenesis including accumulation of large amount of sperm in the seminiferous tubules. AI significantly reduced the serum levels of estradiol-17beta (E2) and increased levels of testosterone (T), 11-ketotestosterone (11-KT), and 17alpha, 20beta-dihydroxy-4-pregnen-3-one (DHP). Further, AI suppressed in vitro production of E2, and stimulated the production of T and 11-KT in the ovarian fragments of mature female. In the honeycomb grouper, suppression of both in vitro and in vivo production of E2 and degeneration of oocytes by AI suggests that AI induces complete sex change through inhibition of estrogen biosynthesis, and perhaps, subsequent induction of androgen function.     

Social system of an inshore stock of the red hind grouper,Epinephelus guttatus (Pisces: Serranidae)

Synopsis According to sex allocation theory, the decision by a female in protogynous fish species to change sex or not should be influenced by, among other things, the mating sex ratio during spawning periods and/or by factors that vary directly with the spawning sex ratio, such as relative rates of behavioral interaction with males and females outside of spawning periods. In groupers that only spawn during a few weeks of the year in large aggregations, individuals must assess the relative value of changing sex or not entirely within the aggregation unless the social system during the remainder of the year provides a behavioral equivalent of the mating sex ratio. Fifty-five individuals of the red hind,Epinephelus guttatus, were tagged and repeatedly located during a 152-day period within a 100 × 100 m grid on a shallow forereef off southwestern Puerto Rico. The home ranges of 22 tagged individuals sighted 10 or more times were 112–5636 m² in area. Individual home ranges overlapped with the home ranges of 1–18 other individuals. Home ranges of small fish were not clustered within the borders of the home range of larger fish, i.e. fish did not form spatially defined social units. At the end of the study, 31 tagged individuals remained on the grid together with five newly sighted fish. All 36 individuals proved on histological examination to be females similar in size to females in the spawning aggregation of the following year. The sex ratio of this all-female inshore stock differed significantly from the sex ratio of that spawning aggregation. Hence, information predicting the reproductive value of a sex change is not available to females in the inshore stock during nonspawning months.     

Maturation and sexual ontogeny in the spangled emperor Lethrinus nebulosus

The reproductive development and sexual ontogeny of spangled emperor Lethrinus nebulosus populations in the Ningaloo Marine Park (NMP) were investigated to obtain an improved understanding of its evolved reproductive strategy and data for fisheries management. Evidence derived from (1) analyses of histological data and sampled sex ratios with size and age, (2) the identification of residual previtellogenic oocytes in immature and mature testes sampled during the spawning season and (3) observed changes in testis internal structure with increasing fish size and age, demonstrated a non‐functional protogynous hermaphroditic strategy (or functional gonochorism). All the smallest and youngest fish sampled were female until they either changed sex to male at a mean 277·5 mm total length (L_T) and 2·3 years old or remained female and matured at a larger mean L_T (392·1 mm) and older age (3·5 years). Gonad masses were similar for males and females over the size range sampled and throughout long reproductive lives (up to a maximum estimated age of c. 31 years), which was another correlate of functional gonochorism. That the mean L_T at sex change and female maturity were below the current minimum legal size (MLS) limit (410 mm) demonstrated that the current MLS limit is effective for preventing recreational fishers in the NMP retaining at least half of the juvenile males and females in their landed catches.     

Reproductive biology and implications for management of the orange-spotted grouper Epinephelus coioides in the southern Arabian Gulf

The reproductive biology of Epinephelus coioides was determined from the examination of 1455 individuals collected between July 2005 and June 2007 in the southern Arabian Gulf. Histological preparations of gonads indicated that males were either derived from a juvenile phase or the transition of postspawning females, confirming a diandric protogynous sexual pattern. The spawning season was well defined, occurring once a year during April and early May. Peaks in spawning occurred after the full and new moons and was completed within a single lunar cycle. The presence of mature males over the entire size and age range and the absence of inactive mature females during the spawning season suggested that the population was not constrained by sperm limitation. While specimens undergoing sexual transition were only observed in size and age ranges of 335–685 mm total length ( L _T) and 5–6 years, patterns in the proportion of males in size and age classes suggested that sex change occurred at a relatively constant rate after female maturation up to the maximum size (1002 mm L _T) and age (11 years). Relationships between reproductive output and capacity with size and age indicated that conventional regulations that equate the mean size at first capture to sexual maturation are unsuitable for the management of E. coioides . The maximum age, small size and young age at sexual maturation ( L _min= 320 mm L _T, 2 years, for females and 242 mm L _T, 1 year, for males) conflict with the general pattern for large epinepheline groupers and may be a direct result of the intensive demersal fishery in the southern Arabian Gulf.     

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.     

A social basis for the development of primary males in a sex-changing fish

An example of alternative male strategies is seen in diandric protogynous (female first) hermaphrodites, where individuals either mature directly as male (primary males) or first reproduce as female and then change sex to male (secondary males). In some sex-changing fishes, the testes of primary males appear anatomically similar to those of non-sex-changing species, whereas the testes of secondary males have anatomical evidence of their former ovarian function. Here, we provide evidence that in the bluehead wrasse, Thalassoma bifasciatum, these strikingly different male phenotypes arise from differences in the ontogenetic timing of environmental sex determination, timing that can be experimentally altered through changes in the social circumstances. Juveniles differentiated almost exclusively as females when reared in isolation, regardless of whether they were collected from a reef with a high proportion of primary males or from a reef with a low proportion of primary males. In contrast, one individual usually differentiated as a primary male when reared in groups of three. Our results indicate that primary males of the bluehead wrasse are an environmentally sensitive developmental strategy that has probably evolved in response to variation in the reproductive success of primary males in populations of different sizes.     

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.     

Dynamics of sex reversal in the marbled swamp eel (Synbranchus marmoratus Bloch, 1795), a diandric hermaphrodite from Marechal Dutra Reservoir,northeastern Brazil

This study characterizes the dynamics of sex reversal in the marbled swamp eel, Synbranchus marmoratus (Osteichthyes: Synbranchidae), a diandric hermaphrodite, within the context of managing species with complex sex allocations. Monthly sampling in Marechal Dutra Reservoir, northeastern Brazil, was conducted using metal eel traps from July, 2013, to June, 2014, during which a total of 288 individuals were captured. Morphological and histological comparisons of gonads identified four sex types: primary males (n = 18), females (n = 197), transitional individuals (n = 30), and secondary males (n = 43). Primary males were smallest, ranging 18–32 cm total length. Females were numerically dominant throughout the 1‐year sampling period, and ranged 20–60 cm. Transitional individuals ranged 32–60 cm, and secondary males ranged 46–74 cm. The otolith‐based age of 52 specimens ranged 0.5 to 5+ year. Primary males were only observed at ages 0.5 and 1, and transitional individuals were only observed at ages 3 and 4 during the female‐to‐secondary‐male transition, supporting the existence of two types of individuals: gonochoristic males and protogynous hermaphrodites. This observation was further supported by histological observations of deteriorating ovarian tissue in transitional individuals. Given the length of time required for individuals to attain secondary male status, this species appears to be particularly vulnerable to over‐exploitation. Comparisons with results from other studies suggest sex allocations and adult size distributions vary substantially within this species’ range, adding complexity to management efforts.     

Long-term study of the ecology of wild Atlantic salmon smolts in a small Norwegian river

Backcalculated lengths at the end of the first growth season in wild Atlantic salmon Salmo salar differed significantly between parr smolting at age 1, 2 and 3 years over a period of 11 years (i.e. 1983–1993). Mean body lengths of the respective age groups at the end of the first growth period were 11·1, 6·2 and 4·7 cm, respectively. The mean percentage distribution of fish smolting at age 1, 2 and 3 was 14, 78 and 7%, and the mean smolt age was 1·95 years. Mean lengths at smolting of age groups 1, 2 and 3 were 13·6, 15·8 and 17·5 cm, respectively. Females outnumbered males among the downstream migrating smolts with a mean sex ratio (females/ males) estimated at 1·61, with a significant female surplus in 7 of the 11 years sampled. Of the smolts sampled, 14% exhibited enlarged gonads indicative of parr maturation, and all were males (37% of the parr males sampled). Mean annual smolt density from 1975 to 1996 was 13·4 individuals 100 m⁻² ranging between 0·3–31 smolts 100 m⁻². Mean densities (100 m⁻²) of the smolts aged 1, 2 and 3 years were 1·5, 9·3 and 0·9 fish, respectively. Mean annual biomass for the 22-year period (1975–1996) was estimated at 437 g 100 m⁻², with a range of variation from 136 to 683 g 100 m⁻². Smolt age 2 made up 81% of the mean annual biomass (355 g 100 m⁻²) and smolt age 1 and 3, 8% (35 g 100 m⁻²) and 11% (47 g 100 m⁻²), respectively.     

Seasonal characteristics of the functioning of the hypophysis-gonad system in the suslik Citellus parryi

In experiments on the arctic ground squirrel C. parryi, studies have been made on seasonal changes in the weight of testes, follicular diameter in the ovaries and the content of sex and gonadotropic hormones in the peripheral blood. Testicular involution and arrest of follicular development were observed in prehibernation period. During hibernation, follicular growth and the increase in the weight of testes take place. The level of LH decreases during hibernation. In sleeping animals, its level is higher as compared to that in active animals during short periods of arousal. The increase in LH level takes place both in males and females in April. FSH can not be detected in males during the first half of hibernation, appearing in the peripheral blood only in March and April. In females, FSH was found in the blood in October, being absent from November to January; beginning from February, it may be found both in sleeping and active animals. Testosterone was found in hibernating males and females, its level significantly increased in March in males, being approximately constant in hibernating females. Estradiol secretion was noted in hibernating females, whereas progesterone was found in the blood only in May.     

Sex ratio, survival and territorial behaviour of polygynous Hen Harriers Circus c. cyaneus in Orkney

Polygyny is widespread among Orkney's Hen Harriers Circus cyaneus . From 1975–81, it was associated with a sex ratio estimated from resightings of colour-marked birds up to six years of age to be 29 females:ten males. The sex ratio of fledglings changed significantly from a greater proportion of females in the 1950s and early 1960s to a greater proportion of males subsequently. The mean estimated 'survival' rates (birds colour-marked in Orkney which were seen there in later years) of males and of females 0–2 years old were 14 and 29%. The mean estimated annual survival rates of males and females from 2–6 years old were 72 and 90%. It is suggested that the uneven sex ratio resulted in more frequent intra-sexual encounters and displays by females. The results of temporary removals of two females (immediately replaced) and of two males (not replaced) in spring indicates that there was a shortage of males in the population.     

The light and the electron microscopic studies of Peyer's patches in non germ-free adult mice

The sexual apparatus was studied in 100 adult axolotls (Siredon mexicanum) for 13 different spawnings. The ages of the animals varied between two and six years. Additional material from Indiana University was also studied. Altogether there were 55 female and 52 male adult axolotls represented. The purpose of the study was to investigate the limits of the variations occurring in normal axolotls and to compare the incidences of variations and developmental abnormalities in adult animals of both sexes at various ages and belonging to different strains. Among the 13 spawnings examined, five strains were completely normal in 100% of the animals, but the remaining eight strains all included abnormal animals. The incidence of abnormal animals in some of these latter strains was 40% or even 50%. Since all of the animals were under the same conditions, the variability and the occurrence of developmental abnormalities most likely depended upon hereditary factors. Among 55 females, only seven (12.7%) were abnormal; only four of these had developmental abnormalities, and only one was hermaphrodite. Among 45 males from the author's axolotl colony, 16 (28%) were abnormal. Of these latter, six had no sex cells or very few; this variation must be regarded as a developmental abnormality. All of these malformations resulted from major degeneration processes and abnormal morphogenesis. Arrested development was also observed in many males. Spermatozoa were completely absent from the testes of eight animals. In the additional material from Indiana University (testes from 7 males), there was also one completely abnormal testis with major degeneration processes and complete absence of sex cells. It is evident that variability and the incidence of developmental abnormalities in the sexual apparatus in adult axolotls of some strains are very great.     

The Effect of Aluminum Exposure on Reproductive Ability in the Bank Vole (<Emphasis Type="Italic">Myodes glareolus</Emphasis>)

Human impact on the environment is steadily increasing the amounts of aluminum in the ecosystems. This element accumulates in plants and water, potentially exposing herbivores to its harmful effect. In heavily polluted sites, a decrease in the density of small rodent populations has been observed. This decline may be caused by many factors, including decreased fertility. The aim of the presented research was to determine how aluminum, administered at concentrations similar to those recorded in industrial districts (Al I = 3 mg/l, Al II = 200 mg/l), affects the reproductive abilities of small rodents. As the indicators of reproductive abilities, body weight, weight of the testes and accessory sex glands of males, and uterus weight of females were estimated. In females, the number of matured follicles (types 6, 7, and 8) was analyzed, while in males, the quantity and quality (matured, viable, swollen, motile, head abnormalities) of epididymal sperm cells were assessed. Moreover, the development of testes, measured by spermatogenic index, was determined. The model species was the bank vole. Our results have proven that aluminum impairs adult individuals’ reproductive abilities by decreasing the quality and quantity of sperm cells and by causing morphologically abnormal development of the gonads. However, no difference in male organometric parameters was found, and only in females treated with 3 mg/l Al, the uterus weight was higher than control. No differences were found in the total number of matured follicles. These results suggest that the decline in rodent numbers in industrial districts is due, at least in part, to poorer males’ reproductive abilities, resulting from exposure to aluminum contamination.