This study establishes that ovulated female goldfish release F type prostaglandins (PGFs) to the water where they stimulate male spawning behavior and comprise the goldfish postovulatory pheromone. We first demonstrated that ovulated and prostaglandin-injected female goldfish release immunoreactive PGFs to the water. Next, using electro-olfactogram recording (EOG), we determined that waterborne prostaglandins function as potent olfactory stimulants for mature male goldfish. Prostaglandin F2 alpha (PGF2 alpha) and its metabolite 15-keto-prostaglandin F2 alpha (15K-PGF2 alpha) were the most potent prostaglandins; the former had a detection threshold of 10(-10) M and the latter a detection threshold of 10(-12) M. Studies of prostaglandin-injected fish indicated that PGF metabolites are an important component of the pheromone. Cross-adaptation experiments using the EOG demonstrated that goldfish have separate olfactory receptor sites for PGF2 alpha and 15K-PGF2 alpha that are independent from those that detect other olfactory stimulants. Finally, we established that male goldfish exposed to low concentrations of waterborne PGFs exhibit reproductive behaviors similar to those elicited by exposure to the odor of ovulated fish. Together with our recent discovery that a steroidal maturational hormone functions as a preovulatory "priming" pheromone for goldfish, these findings suggest that hormones and their metabolites may commonly serve as reproductive pheromones in fish.

The major progestin in teleosts is not progesterone, as in tetrapods, but 17,20beta-dihydroxypregn-4-en-3-one (17,20beta-P) or, in certain species, 17,20beta,21-trihydroxy-pregn-4-en-3-one (17,20beta,21-P). Several functions for 17,20beta-P and 17,20beta,21-P have been proposed (and in some cases proved). These include induction of oocyte final maturation and spermiation (milt production), enhancement of sperm motility (by alteration of the pH and fluidity of the seminal fluid) and acting as a pheromone in male cyprinids. Another important function, initiation of meiosis (the first step in both spermatogenesis and oogenesis), has only very recently been proposed. This is a process that takes place at puberty in all fishes and once a year in repeat spawners. The present review critically examines the evidence to support the proposed functions of 17,20beta-P in males, including listing of the evidence for the presence of 17,20beta-P in the blood plasma of male fishes and discussion of why, in many species, it appears to be absent (or present at low and, in some cases, unvarying concentrations); consideration of the evidence, obtained mainly from in vitro studies, for this steroid being predominantly produced by the testis, for its production being under the control of luteinizing hormone (gonadotrophin II) and, at least in salmonids, for two cell types (Leydig cells and sperm cells) being involved in its synthesis; discussion of the factors involved in the regulation of the switch from androgen to 17,20beta-P production that seems to occur in many species just at the time of spermiation; discussion of the effects of in vivo injection and application of 17,20beta-P (and closely related compounds) in males; a listing of previously published evidence that supports the proposed new function of 17,20beta-P as an initiator of meiosis; finally, discussion of the evidence for environmental endocrine disruption by progestins in fishes.

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Prostaglandins (PGs) have been identified in gonads, semen, ovarian fluid, blood, and in vitro ovarian incubates from a variety of teleosts. In teleosts, PGs appear to be involved in ovulation (follicular rupture) and female sexual behavior, and possibly in gonadotropin (GtH) secretion. An increase in prostaglandin F (PGF) levels associated with GtH-induced ovulation occurs in vivo in the pond loach (Misgurnus anguillicaudatus) and goldfish (Carassius auratus). Indomethacin (PG synthesis inhibitor) blocks ovulation in these species and, in goldfish, PG injection reverses this blockade. PGF2α stimulates in vitro ovulation in rainbow trout (Salmo gairdneri), brook trout (Salvelinus fontinalis), and yellow perch (Perca flavescens); however, in perch, PGE2 is the most potent prostaglandin. Addition of melatonin to incubation medium both inhibits ovulation and decreases PGE and PGF synthesis in yellow perch, while addition of epinephrine and theophylline both enhances ovulation and increases PGE and PGF synthesis. Several studies indicate that PG, released from the ovaries or oviduct in response to the presence of ovulated oocytes, acts on the brain to stimulate female spawning behavior in the goldfish. Other externally fertilizing teleosts may use similar mechanisms to synchronize female sexual behavior with ovulation.Key words: prostaglandins, fish reproduction, ovulation, sexual behavior