Endocrine manipulations of spawning in cultured fish: from hormones to genes

Almost all fish reared in captivity exhibit some form of reproductive dysfunction. In females, there is often failure to undergo final oocyte maturation, ovulation and spawning; while in males milt production may be reduced and of low quality. These dysfunctions are due to the fact that fish in captivity do not experience the conditions of the spawning grounds, and as a result there is a failure of the pituitary to release the maturational gonadotropin, luteinizing hormone (LH). Reproductive hormones have been utilized since the 1930s to stimulate reproductive processes and induce ovulation/spermiation and spawning. The first methods employed freshly ground pituitaries collected from reproductively mature fish, which contained gonadotropins (mainly LH) and induced steroidogenesis and gonadal maturation. Eventually, purified gonadotropins became available, both of piscine and mammalian origin, e.g., carp or salmon gonadotropin, and human chorionic gonadotropin. In the 1970s, spawning induction methods begun employing the newly discovered gonadotropin-releasing hormone (GnRH), which induces the secretion of the fish's own gonadotropin from the pituitary, thereby overcoming the endocrine failure observed in captive broodstocks. Development of highly potent, synthetic agonists of GnRH (GnRHa) constituted the next generation of hormonal manipulation therapies, and created a surge in the use of hormones to control reproductive processes in aquaculture. The most recent development is the incorporation of GnRHa into polymeric sustained-release delivery systems, which release the hormone over a period of days to weeks. These delivery systems alleviate the need for multiple treatments and induce (a) long-term elevation in sperm production and (b) multiple spawning in fish with asynchronous or multiple-batch group-synchronous ovarian physiology. Based on the recent discovery of GnRH multiplicity in fish and the increasing understanding of its functional significance, new GnRH agonists can be designed for more potent, affordable and physiologically-compatible spawning induction therapies. Future strategies for improved spawning manipulations will be based on understanding the captivity-induced alterations in the GnRH system, and on new approaches for their repair at the level of GnRH gene expression and release.