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

Abstract

A great deal of information is known regarding the process of sex differentiation in fish, and the mechanisms involved in primary sex determination are now beginning to be defined. A range of gonadal differentiation types have been described for fish, including gonochoristic species possessing purely ovarian or testicular tissues, as well as hermaphroditic species that can initially mature either as males (protandrous) or females (protogynous). Sex determination in fish is a very flexible process with respect to evolutionary patterns observed among genera and families, and within individuals is subject to modification by external factors. These influences can affect the fate of both somatic and germ cells within the primordial gonad, and include the action of genetic, environmental (e.g. temperature), behavioural, and physiological factors. Exogenous sex steroids administered at the time of sex determination can strongly influence the course of sex differentiation in fish, suggesting that they play a critical role in assignment of gonad determination as well as subsequent differentiation. Detailed information is available from fish systems describing the production of sex steroids, as well as the enzymes involved in steroid production. Both estradiol and the maturation hormone 17α, 20β-dihydroxy-4-pregnen-3-one (17α, 20β-DP) are produced by a two-step process involving different cell layers in the gonad, and have effects on the differentiation of gonadal and nongonadal tissues. Gonadal development and differentiation in some fish is also controlled by hormones from the pituitary gland (gonadotropins) that are regulated by release hormones (GnRH) and other neuroendocrine and gonadal factors. Genetic determination of sex in fish can involve monogenic or polygenic systems, with factors located on the autosomes or on sex chromosomes. In the latter case, both male (XY) and female (ZW) heterogametic systems have been described, as well as many subtle variations on these themes. Sex chromosomes are found in approximately 10% of fish species examined, and sex-linked phenotypic traits, and protein and molecular genetic markers have been identified in several fish systems. Some species of fish reproduce gynogenetically, producing all-female populations. Several gene families known to be involved in sex determination in other vertebrates have recently been shown to be similarly involved in fish, suggesting conservation of sex determination pathways. The lability of sex-determination systems in fish makes some species sensitive to environmental pollutants capable of mimicking or disrupting sex hormone actions. Such observations provide important insight into potential impacts from endocrine disruptors, and can provide useful monitoring tools for impacts on aquatic environments.