http://aerg.canberra.edu.au/library/sex_general/2002_Devlin_Nagahama_sex_determination_in_fish_review.pdf

Sex determination and sex differentiation in fish: an overview of genetic, physiological, and environmental influences

Spermatogenesis in fish.

Oogenesis in teleosts: How fish eggs are formed

Oocyte growth and development is an important issue in fish and fisheries biology. This paper reviews the information available on oocyte growth patterns and the rates and dynamics of oocyte growth in teleosts. In synchronous spawners, the weight of the gonad may represent as much as 40% of the overall body weight of the fish. In asynchronous spawners, the weight of the mature ovary is considerably less than in synchronous ovulators, but the ovary shows a more regular periodicity and may grow repeatedly many times during the breeding season. There is a huge variability in egg size in teleosts, with the largest known measuring up to 8 cm in diameter. Within the limits of variance set by genetic constraints, egg size may vary between populations of the same species. Oocytes in all teleosts undergo the same basic pattern of growth: oogenesis, primary oocyte growth, cortical alveolus stage, vitellogenesis, maturation and ovulation. The mechanisms that control oocyte growth are addressed in this review, albeit that the available information, as in all other vertebrates, is very limited. The main hormones that have been shown to affect ovarian growth are gonadotrophin, thyroid hormones, growth hormone, insulin and insulin-like growth factors. An overview of the determinants of fecundity, with particular reference to oocyte recruitment and atresia, is the focus of the second part of the paper. Genetics and nutrition have major effects on fecundity, and studies so far suggest that the determinants of fecundity usually operate during the early part of gametogenesis. The role of atresia in determining fecundity is less clear. The final part of this review highlights some areas of study that are priorities for research on ovarian development in fish.

Oocyte growth and development in teleosts

A period of oocyte growth is followed by a process called oocyte maturation (the resumption of meiosis) which occurs prior to ovulation and is a prerequisite for successful fertilization. Our studies using fish models have revealed that oocyte maturation is a three-step induction process involving gonadotropin (LH), maturation-inducing hormone (MIH), and maturation-promoting factor (MPF). LH acts on the ovarian follicle layer to produce MIH (17α, 20β-dihydroxy-4-pregnen-3-one, 17α, 20β-DP, in most fishes). The interaction of ovarian thecal and granulosa cell layers (two-cell type model), is required for the synthesis of 17α,20β-DP. The dramatic increase in the capacity of postvitellogenic follicles to produce 17α,20β-DP in response to LH is correlated with decreases in P450c17 (P450c17-I) and P450 aromatase (oP450arom) mRNA and increases in the novel form of P450c17 (P450c17-II) and 20β-hydroxysteroid dehydrogenase (20β-HSD) mRNA. Transcription factors such as Ad4BP/SF-1, Foxl2, and CREB may be involved in the regulation of expression of these steroidogenic enzymes. A distinct family of G-protein-coupled membrane-bound MIH receptors has been shown to mediate non-genomic actions of 17α, 20β-DP. The MIH signal induces the de novo synthesis of cyclin B from the stored mRNA, which activates a preexisting 35 kDa cdc2 kinase via phosphorylation of its threonine 161 by cyclin-dependent kinase activating kinase, thus producing the 34 kDa active cdc2 (active MPF). Upon egg activation, MPF is inactivated by degradation of cyclin B. This process is initiated by the 26S proteasome through the first cut in its NH2 terminus at lysine 57.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1440-169X.2008.01019.x

Regulation of oocyte maturation in fish.

The pituitary-gonadal axis plays an important role in regulating gametogenesis in vertebrates. In most cases, gonadotropins act through the biosynthesis of gonadal steroid hormones which in turn mediate various stages of gametogenesis. A series of studies in our laboratory using several species of teleost fishes as experimental animals has provided new information about the endocrine regulation of gametogenesis, including oocyte growth, oocyte maturation, spermatogenesis and sperm maturation. This article briefly reviews our findings on the identification of steroidal mediators involved in each process of gametogenesis, and the sites and mechanisms of action of the mediators. These observations collectively demonstrate the appropriateness of using teleost fishes as valid models for examining hormonal influences on gametogenesis. Such models could also have applications and validity for vertebrates in general.

/pdf/endocrine-regulation-of-gametogenesis-in-fish-c963fs25ox.pdf

Endocrine regulation of gametogenesis in fish.

/pdf/human-recombinant-activin-induces-proliferation-of-48ka74i2te.pdf

Human Recombinant Activin Induces Proliferation of Spermatogonia in vitro in the Japanese Eel Anguilla japonica

https://febs.onlinelibrary.wiley.com/doi/pdf/10.1016/S0014-5793%2896%2901187-8

Fish testicular 11β-hydroxylase : cDNA cloning and mRNA expression during spermatogenesis

Under cultivation conditions, male Japanese eels (Anguilla japonica) have immature testes containing only spermatogonia together with inactive testicular somatic cells, Leydig cells and Sertoli cells. Using a recently developed organ culture system for eel testes, we have shown that hormonal induction of spermatogenesis in eel testes involves gonadotropin stimulation of Leydig cells to produce 11-ketotestosterone, a potent androgen in fish. In turn, 11-ketotestosterone activates Sertoli cells to stimulate premitotic spermatogonia to complete spermatogenesis. Our current research focuses on the isolation and characterization of genes that show altered expression in eel testes during gonadotropin-induced spermatogenesis. One up-regulated and three down-regulated genes have been isolated. Northern blot analysis and in situ hybridization reveal that mRNA for activin B is absent in testes before gonadotropin injection and is abundant in Sertoli cells in testes injected with gonadotropin for one to six days after injection. This stimulation of activin B mRNA is accompanied by spermatogonial proliferation. Gonadotropin treatment also causes a rapid rise in the testicular concentrations of mRNA for 3 beta-hydroxysteroid dehydrogenase, the rate-limiting enzyme for gonadotropin-induced 11-ketotestosterone production. We have also obtained three down-regulated cDNAs which are abundant in testes before gonadotropin treatment and disappear almost completely in testes one day after gonadotropin injection.

The onset of spermatogenesis in fish.

The testis of Japanese eel (Anguilla japonica) consists of type A and early type B spermatogonia together with inactive Leydig and Sertoli cells. A single injection of human chorionic gonadotropin induced marked changes in the morphology of the testis and in the serum androgen levels within a period of 72 h. Morphological changes include spermatogonial proliferation, activation of Leydig and Sertoli cells, organization of seminiferous lobules and formation of lobular lumen in the testis. Leydig cells were enlarged, exhibiting characteristics of steroid-producing cells. Sertoli cells become elongated, show signs of high cellular activity and remain in close contact with spermatogonia. The lobular organization was achieved much earlier than the progression of spermatogenesis to late type B spermatogonia. Even 6 h after hCG injection, a significant increase in plasma levels of 11-ketotestosterone was observed, followed by a further time dependent increase. Plasma testosterone levels were also increased after injection, but the increase was much less than that of 11-ketotestosterone.

Yoshitaka Nagahama

Papers

Endocrine regulation of gametogenesis in fish.

Human Recombinant Activin Induces Proliferation of Spermatogonia in vitro in the Japanese Eel Anguilla japonica

Fish testicular 11β-hydroxylase : cDNA cloning and mRNA expression during spermatogenesis

The onset of spermatogenesis in fish.

A rapid activation of immature testis of Japanese eel (Anguilla japonica) by a single injection of human chorionic gonadotropin