The reproductive cycle of yellowfin bream, Acanthopagms australis (Günther), with particular reference to protandrous sex inversion

Yellowfin bream, Acanthopagrus australis , of all age classes were collected from Moreton Bay, Australia. The species possessed typical sparid ovotestes in which the testis and ovary occur in separate zones. During the spawning period (June-August) juveniles, functional males and functional females could be distinguished by the macroscopic appearance of the gonad. The sex ratio of males to females decreases with age, indicating protandrous sex inversion.
Histological and structural study of the ovotestis showed all fish have previtellogenic cells in the ovarian zone but only juvenile and male fish have developing spermatogenic cells in the testis. Most juveniles become functional males by the age of two years but a small proportion of juveniles develop directly into functional females (primary females). Protandrous sex inversion commences after the spawning period when male fish appear with spermatozoa and no other spermatogenic cells in the testis. During the period November-January male fish with no spermatogenic cells are common and a reduction in size of the testis occurs so that by March-April the ovotestis becomes structurally and histologically similar to the female ovotestis. Some fish remain functional males during their whole life-history (primary males). In functional females vitellogenic cells are present in the ovary only during the spawning period and the testis remains very small in size.     

Reproductive Biology and Protandrous Hermaphroditism in Acanthopagrus latus

Detailed macroscopic and histological studies of the gonads of a full size and age range of Acanthopagrus latus from each season in Shark Bay, Western Australia, demonstrate that this species is a protandrous hermaphrodite in this large subtropical embayment. Although our scheme for the changes that occur in the ovotestes of A. latus during life is not consistent with some of the conclusions drawn for this species elsewhere, it is similar to that of Pollock (1985 J. Fish. Biol. 26: 301–311) for the congeneric Acanthopagrus australis. The ovotestes of males develop from gonads which contain substantial amounts of both testicular and ovarian tissue. The testicular component of the ovotestes of all males regresses markedly after spawning. During the next spawning season, the ovotestes either become gonads in which the testicular zone again predominates and contains spermatids and spermatozoa (functional males), or gonads in which the ovarian zone now predominates and contains mature oocytes (functional females). Once a fish has become a functional female, it remains a female throughout the rest of its life. In Shark Bay, A. latustypically spawns on a limited number of occasions during a short period in late winter and early spring and has determinate fecundity. The mean potential annual fecundity was ca. 2000000. The total length of 245mm, at which, during the spawning period, 50% of A. latus become identifiable as males, is very similar to the current minimum legal length (MLL) of 250mm, which corresponds to an age of ca. 2.5 years less than that at which 50% of males become females. Thus, although the spawning potential ratio suggests that the present fishing pressure is sustainable, the current MLL should be reviewed if recreational fishing pressure continues to increase.     

Experimentally induced true hermaphroditism in genetically female fowl

Summary Two types of hermaphroditism were experimentally induced in genetically female fowls by grafting of embryonic testes in embryos. Of the 27 hermaphrodites observed during the 8 months after hatching, 20 possessed a right testis and a left ovary and 7 a right testis and a left ovotestis. The testes and ovotestes contained seminiferous tubules with a more or less developed germ cell complement, attaining in many cases the early spermatid stage. The interstitial tissue was poorly functional, as shown by the absence of male secondary sex characters. The ovary or ovarian part of the ovotestes possessed numerous small ovarian follicles. The female arrangement of the plumage and the absence of spurs demonstrated the secretion of oestrogens. A mechanism is proposed for explaining this partial masculinization of genetically female gonads, a phenomenon which occurs during the period of embryonic sex differentiation, and is responsible for this experimental true hermaphroditism.     

Aspects of reproductive biology of the Mediterranean amberjack (Seriola dumerilii Risso) during the spawning period

Wild Seriola dumerilii were collected in the South Mediterranean Sea during the 1990, 1991 and 1992 spawning seasons. Macroscopic and cytological characteristics of ovary and testis were analysed. Depending on the presence and the number of oocytes at different stages, a five point maturity scale was proposed for ovarian maturity. Nine maturity stages of the oocytes are described. Oocyte size-frequency distribution has shown a group synchronous ovarian development type. The testes have a typical lobular-type structure. Depending on lobules and sperm duct development and on the abundance of germ cells, testis maturity was classified in four stages. Maturity rate according to age and size was also determined. 100% of 1-year-old fish show immature gonads. The proportion of females with mature ovaries was 0, 12.5, 84.6 and 100% at the age of 2, 3, 4 and 5 years respectively. The proportion of males with mature testes was 14.3, 40, 80 and 100 at the same age. The median standard length at which 50% of the fish attained maturity is 109 and 113 cm SL in males and females respectively.     

Sexuality, sex change and maturation patterns in the yellowfin seabream, Acanthopagrus latus (Teleostei: Sparidae) (Houttuyn, 1782)

Macroscopic and histological studies were carried out to describe the reproductive styles and sex reversal and to follow gonadal changes in captive yellowfin seabream during the second year of life. Four reproductive styles are found in Acanthopagrus latus (Houttuyn, 1782): (i) males and females (gonochorism), (ii) functional males, (iii) transitionals and (iv) functional females. The species is a protandric hermaphrodite and begins life as a functional male with testicular zone undergoing active spermatogenesis, while the ovarian zone is arrested at the primary growth (perinucleolar) phase. Males and females were encountered in virtually all size‐classes. Functional males outnumbered the functional females in all size‐classes in which they were encountered. Sex reversal begins in the transitionals from July to August, after spawning in the functional males, at 14.9–20.2 cm standard length (SL) and, by November, maturation of the ovarian tissue begins. A. latus in cages in Kuwait waters spawns from January to April with a peak in February for males, and a peak in March for females and transitionals. Spawning begins in the 18.3–20.2 cm size‐range fish, peaking in the 20.3–22.2 cm size‐range in both males and females and also in the transitionals, although a few of the latter spawn from 14.9 cm SL. In relation to age, spawning begins at 20 months in males and peaks at 21 months. Females begin to spawn at 21 months with a peak at 22 months, while transitionals generally begin to spawn at 20 months, although a few 14–15‐month‐old sex‐changing individuals were encountered. Temperature, either alone or in combination with other unknown factors, triggers spawning in A. latus.     

Further studies on the sexuality of the hermaphroditic teleost Diplodus sargus, with particular reference to protandrous sex inversion

Histological events associated with the process of sex inversion are described in the protandrous teleost Diplodus sargus reared in captivity. This species possesses typical sparid ovotestes in which the testis and ovary occur in separate zones. After spawning, the testicular tissue within the bisexual gonad undergoes regressive changes such as spermatogonial degeneration, deposition of yellowish pigment and proliferation of connective cells and fibres. At the same time ovarian tissue begins to develop. Transitional individuals were 26–7% of examined fish, while the other 73–3% were functional males.     

Reproductive strategy of Leporinus piau (Fowler, 1941), a neotropical freshwater fish from the semi‐arid region of Brazil

This study concerns the reproductive strategy of the freshwater fish, Leporinus piau, native to the Brazilian semi‐arid region. The hypothesis of this work predicts that the species would spawn during the rainy season, presenting a seasonal reproductive strategy. The study investigated the structure of total body length and body mass, sex ratio, body size at first sexual maturity, developmental stages of gonads, fecundity, gonadosomatic index (GSI), condition factor (K) and the spawning season of L. piau. Twelve monthly samplings (mean n = 18) were carried out in 2009 and environmental parameters such as rainfall, temperature, pH, electrical conductivity and concentration of dissolved oxygen of water were registered. The sampled population of L. piau (n = 211) showed a slight predominance of males (55%), but with larger and heavier females. First sexual maturation in males occurred earlier (16.5 cm total length) than in females (20.5 cm total length). The species is a total spawner with an average fecundity of 55 000 mature oocytes. Rainfall (225 ± 7.2 mm) and concentration of dissolved oxygen (8.5 ± 0.2 mg L^?1) acted as influential factors during the spawning season. The monthly GSI shows that spawning starts when the rainy season is at its maximum and ends synchronously (females r = 0.91; males r = 0.89) with the end of the rainy season. L. piau presents a seasonal reproductive strategy with high reproductive output, providing a population increase and exploitation of the environmental resources.     

Reproduction, age and growth of Sillago maculata in south-eastern Australia

Sillago maculata is endemic to the east coast of Australia where it is harvested by recreational and commercial fishers; however, little is known of the important aspects of its biology and ecology to assist with fisheries management planning. This situation is redressed here by investigating aspects of the reproduction, age and growth of estuarine populations of S. maculata in south-eastern Australia. Gonadosomatic index (GSI) values indicated peak spawning occurred between September and February and that the estimated mean fork length at maturity (L₅₀) was 14.6 cm for males and 15.2 cm for females. Females displayed an asynchronous pattern of oocyte development, with individuals probably spawning multiple times in a spawning season. A validated aging protocol using thin sectioned otoliths was used to estimate the age of fish. The maximum ages for males and females were 9.5 and 12.0 years, respectively. Growth differed between sexes, with males ( L ∞ = 24.04, K  = 0.70, t ₀ = −0.09) attaining a smaller maximum length than females ( L ∞ = 25.01, K  = 0.72, t ₀ = −0.04). The age composition of gill-net and beach-seine samples mainly consisted of individuals aged 2–4 years, and there was evidence of variable recruitment. Management implications are also discussed.     

Reproductive biology of the silver pomfret, Pampus argenteus (Euphrasen), in Kuwait waters

The reproductive activities of the silver pomfret, Pampus argenteus (Euphrasen), in Kuwait waters were investigated from March 1996 to February 1998. Observations on the seasonal distribution of maturity stages and variations in seasonal fluctuations in the gonadosomatic index (GSI) confirmed recent findings that the spawning period begins in May. The species has a prolonged spawning period in the females extending from May to August, whereas the males mature in April and ripe specimens were encountered in monthly samples until September. The recruitment pattern confirmed the one breeding season. There are two spawning peaks, the first in May and the second in August. Variations in GSI relative to fish length indicated that females and males are most fecund at about 24.5–26.4 cm and 20.5–22.4 cm length classes, respectively. The males mature earlier than females, at a minimum size of 12.5–14.4 cm, while the females mature at 20.5–22.4 cm. The oocyte diameter‐frequency distribution suggests a serial rhythm of spawning. Fecundity ranged from 28 965 to 455 661 and correlated positively with: (a) standard length (P < 0.006); (b) ovary weight (P < 0.001); and (c) body weight (P < 0.001), and negatively with egg size (P < 0.003).     

Reproductive biology of the East African lungfish (Protopterus aethiopicus) in Mwanza Gulf, Lake Victoria

The population of lungfish (Protopterus aethiopicus (Heckel)) was sampled by bottom trawling for 12 months from June 1984 to May 1985. A total of 576 lungfish were investigated. The sex ratio in the population was 1 male : 1 -98 females. Average size at maturity was found to be 960cm total length (TL) but the smallest matured male measured 72-9 cm and the female 840 cm TL. It was not possible to establish spawning times and reproductive activity trends either from the variation in relative condition factor ‘kn’ or from gonadosomatic index (GSI). Whereas male lungfish seem to show no further increase in GSI after gonadal maturity stage III, females exhibit GSI values that vary directly with oocyte maturation and ovarian weights. Fat deposited along the gonads was found to be a better basis for assessing maturation and spawning in lungfish. The potential spawning period was found to be from September to May. Absence of lungfish with spent gonads over the 12-month period and the presence of different egg sizes in ovaries, suggest that the fish is a multiple spawner. Fecundity was found to range from 705 eggs (in 84-cm TL fish) to 14,922 eggs (in 130-cm TL Fish).     

Monogamous mating system and sexuality in the gobiid fish, <Emphasis Type="Italic">Trimma marinae</Emphasis> (Actinopterygii: Gobiidae)

The mating system and sexuality of the gobiid fish Trimma marinae were investigated in aquaria and by gonadal histological examination. The male to female sex ratio in the study aggregation was female biased (14:27), and females were larger than males. T. marinae were monogamous because they established continuous pairs and spawned repeatedly with the same individuals. Observations of aggressive behavior suggested that the monogamous mating system resulted from female mate guarding. We also performed a rearing experiment to test whether sex change occurs in this species. None of the males or females reared separately in aquaria for 63 days changed sex. Additionally, gonadal histology revealed that mature fish had unisexual gonads (testis or ovary). These results strongly suggest that T. marinae is gonochoristic. However, immature fish had a bisexual gonadal structure, indicating juvenile hermaphroditism.     

Steroid hormone profiles and correlative gonadal histological changes during natural sex reversal of sobaity kept in tanks and sea-cages

The gonadosomatic index of sobaity Sparidentex hasta fits well with the natural spawning season of February-March. At 12 and 24 months all the population matured as males. However, during this time, between April-September intersex gonads were seen. From October of the third year of life, females were seen for the first time. These changes in the histology of the gonads were correlated with the hormonal profile. Testosterone showed no significant correlation with sex but 11-ketotestosterone correlated very well with the mature male gonads and thus can be used for sexing. Oestradiol-17β higher in sex reversed individuals (females) in the spawning season than in males. The sequence of events in the protandrous sex change of this fish during the first two spawning seasons is male → intersex → male. However, during the third spawning season (age 36–37 months) this sequence is male →intersex → male or female (10–17%).     

Fertility of gynogenetic Atlantic cod (Gadus morhua L.)

In order to investigate whether meiotic gynogenetic Atlantic cod is fertile and able to produce viable offspring, meiotic gynogenetic females were produced in spring 2010 by activating cod eggs using irradiated sperm. The extrusion of the second polar body was prevented by the application of hydrostatic pressure (56.6 MPa) 36 min after fertilization. In February 2012, their mean round weight was 972 g, and 2580 g in March 2013. In 2012, when the fish were 2 years old, about 52% were mature, 33% were immature, and 13% had undifferentiated gonads. One year later, 77% were mature, 11% were immature, and 11% had undifferentiated gonads. Several of the mature females had malformed gonads, with only one developed ovary lobe or with the two lobes fused. The mean gonadosomic index (GSI) of the 2‐year‐old mature females was 5.2%, with an estimated relative fecundity of 581 000 eggs kg ovary‐free wet weight^?1. Females were stripped for eggs when 2 and 3 years old (2012 and 2013), and fertilized with sperm from normal males. Offspring were obtained from 12 of 17 and 12 of 15 egg batches incubated in 2012 and 2013, respectively, proving that the gynogenetic females are fertile. Furthermore, larvae in all but one of the hatched groups from 2013 had commenced feeding 2 h after being startfed using rotifers 4 days after hatch, indicating viable offspring.     

Biochemical content, energy composition and reproductive effort in the broadcasting sea star Asterias vulgaris over the spawning period

To assess sex differences in reproductive effort, we examined the biochemical composition and energetic content of the principal body components of the broadcast spawning sea star Asterias vulgaris in the Mingan Islands in the northern Gulf of St. Lawrence, eastern Canada. The body wall was the most stable body component, showing no variations in mass or in lipid and protein content (and total energetic content) between sexes or during spawning. Patterns in the gonads differed between sexes and with spawning. The lipid, protein and carbohydrate content of the ovary dropped during spawning, while only the protein content of the testis decreased significantly. Reproductive effort, expressed as loss of energy in the gonads during spawning for an individual weighing 10 g in underwater mass (8.2 cm in radius), was six times greater in females (49.5 kJ) than males (7.9 kJ). The energetic content of the pyloric caeca also decreased during spawning, by 17.7 kJ in females and 21.5 kJ in males, mainly due to a decrease in lipids. If this decrease is included as reproductive effort, it lessens the gender difference. The caecum decrease possibly represented expenditures due to formation of aggregations or the expulsion of gametes during spawning. Effectively, we observed aggregations during a massive spawning in this population. The sex ratio did not differ from 1:1 in all size classes sampled. This suggests that, unless males suffer higher mortality, females manage to allocate as much energy to somatic growth as males, possibly by feeding at higher rates to compensate for their higher reproductive effort. Stomach protein content tended to be higher in females than males and may indicate greater muscular development to facilitate digestion.     

Reproduction, age, growth and mortality of axillary seabream, Pagellus acarne (Sparidae), from the Canarian archipelago

Axillary seabream Pagellus acarne (Risso 1826) caught off the Canary Islands from January 1991 to December 1994 were studied. The length range of the catches was between 11 and 31 cm, with a modal distribution between 17 and 21 cm. The overall ratio of males to females was 1:1.74. Males were observed up to a length of 24 cm. Hermaphrodites were recorded at lengths between 15 and 23 cm. The species was characterized by protandric hermaphroditism. The reproductive season extended from October to March, with a peak in spawning activity in December–January. The size at sexual maturity was 15.8 cm total length (2 years old) for males and 19.4 cm total length (3 years old) for females. The total length–total weight relationship for the entire population is described by the parameters a = 0.0068, and b = 3.2401. Otolith age readings showed that the population exploited consisted of 10 age groups (1–10 years), including a very high proportion of individuals between 1 and 4 years old. The von Bertalanffy growth parameters for all individuals were L_∞ = 32.98 cm, k = 0.22 years⁻¹, and t₀ = −0.87 years. Males grew comparatively slower than females. The instantaneous rates of mortality for all fish were Z = 0.96 years⁻¹, M = 0.30 years⁻¹, and F = 0.66 years⁻¹. The exploitation ratio and the length at first capture were, respectively, E = 0.69 and LC₅₀ = 16.1 cm. The stock is overexploited, therefore measures such as closed seasons or changes in fishing patterns would be desirable to safeguard the spawning stock and recruits.     

Sex Change in Both Directions by Alteration of Social Dominance in Trimma okinawae (Pisces: Gobiidae)

The mating system of the gobiid fish Trimma okinawae is one of polygynous hermaphroditism, in which the largest female of a social unit changes sex following the removal of the dominant male. Histological observations of the gonads however, revealed that males have an ovarian tissue within a functional testis. The occurrence of ovarian tissue in the functional male suggests that T. okinawae males should be able to revert back into being functional females. To test this prediction, we placed females in an aquarium and allowed them to change sex. After confirming sex change from female to male, we individually placed new males into another aquarium and added a larger male to each. Our experiments revealed that females change sex and become males upon the removal of dominant males, and that those males changed sex again and became females in the presence of larger males. Sex change in both directions may be advantageous when a male is forced to become subordinate following the take over of the social unit by a larger male.     

Sex reversal and aromatase in chicken.

Aromatase inhibitors administered before sexual differentiation of the gonads can induce sex reversal in female chickens. To analyze the process of sex reversal, we have followed for several months the changes induced by Fadrozole, a nonsteroidal aromatase inhibitor, in gonadal aromatase activity and in morphology and structure of the female genital system. Fadrozole was injected into eggs on day four of incubation, and its effects were examined during the embryonic development and for eight months after hatching. In control females, aromatase activity in the right and the left gonad was high in the middle third of embryonic development, and then decreased up to hatching. After hatching, aromatase activity increased in the left ovary, in particular during folliculogenesis, whereas in the right regressing gonad, it continued to decrease to reach testicular levels at one month. In treated females, masculinization of the genital system was characterized by the maintenance of the right gonad and its differentiation into a testis, and by the differentiation of the left gonad into an ovotestis or a testis; however, in all individuals, the left Müllerian duct and the posterior part of the right Müllerian duct were maintained. In testes and ovotestes, aromatase activity was lower than in gonads of control females (except in the right gonad as of one month after hatching) but remained higher than in testes of control and treated males. Moreover, in ovotestes, aromatase activity was higher in parts displaying follicles than in parts devoid of follicles. The main structural changes in the gonads during sex reversal were partial (in ovotestes) or complete (in testes) degeneration of the cortex in the left gonad, and formation of an albuginea and differentiation of testicular cords/tubes in the two gonads. Testicular cords/tubes transdifferentiated from ovarian medullary cords and lacunae whose epithelium thickened and became Sertolian. Transdifferentiation occurred all along embryonic and postnatal development; thus, new testicular cords/tubes were continuously formed while others degenerated. The sex reversed gonads were also characterized by an abundant fibrous interstitial tissue and abnormal medullary condensations of lymphoid-like cells; in the persisting testicular cords/tubes, spermatogenesis was delayed and impaired. Related to aromatase activity, persistence of too high levels of estrogens can explain the presence of oviducts, gonadal abnormalities and infertility in sex reversed females.     

Life history of the orange-striped emperor <Emphasis Type="Italic">Lethrinus obsoletus</Emphasis> from the Mariana Islands

The present study investigated the age-based demography of the orange-striped emperor, Lethrinus obsoletus, from commercial samples in the Commonwealth of the Northern Mariana Islands (CNMI), spanning a 24-month market sampling period. Information on growth, life span, mortality, and reproduction was derived through analysis of sectioned sagittal otoliths and gonad material. The species had a moderate longevity of 13 years while females and males reached 50% sexual maturity at 3.8 years (22.9 cm fork length) and 2.8 years (19.9 cm fork length), respectively. Histological examination of gonads and sex-specific age frequency distributions suggest the sexual ontogeny of the species conforms to juvenile hermaphroditism, whereby sexual transition from female to male occurred prior to maturation. No annual spawning periods were identified, but the ubiquitous presence of post-ovulatory follicles in females and spermatogenic material in males coupled with a consistent lunar trend in gonadosomatic index patterns suggests the species spawns every lunar cycle throughout the year with spawning activity potentially increasing around the new moon. Ratios of natural to fishing mortality indicate a moderate level of exploitation (0.37); however, a large portion of harvested females had not reached reproductive maturity, suggesting that formal assessments of stock status are warranted to ensure the sustainable harvest of the species in CNMI.     

Gonadal development and spawning cycle in the digynic hermaphrodite sharpsnout seabream Diplodus puntazzo (Sparidae) off the Canary Islands, northwest of Africa

Diplodus puntazzo is a species characterized by a rudimentary hermaphroditism, with a low level of protandry. The study found that the overall sex ratio was one male to 2.03 females, diverging significantly from 1 : 1, and that there was a ratio of one male to 0.11 transitional fish. The sex ratio varied with season: during the reproductive period the ratio was 1 : 1.86 (male : female), significantly lower than in the non‐reproductive period in which the sex ratio was approximately 1 : 2.26. Both sexes had similar size distributions; however, females were predominant in all size intervals, especially in the larger ones. No significant difference in mean total length (TL) between females (330 mm), males (319 mm) or transitionals (321 mm) was found. The smallest functional female and male analysed were 159 and 157 mm TL, and the largest were 520 and 510 mm TL, respectively. No significant difference in length at first level of maturity, attained at the second year of life, was found between males (281 mm) and females (292 mm), respectively. A protracted spawning season was identified for the species: from September to February, with a peak in November. GSI values followed the same pattern for males and females, with values of females being similar to those of males; the highest values occurred between September and December, with a maximum in November. Occurrence of transitional fish was significantly higher during the month with highest values of GSI than in the resting period. The study found that vitellogenesis did not begin synchronously for all females, and the simultaneous presence of oocytes in all stages of development indicated an asynchronous mode of ovarian development. The presence of postovulatory, together with tertiary yolk vesicle oocytes, indicated that the species is a multiple spawner.