Introduction 318Cycles in relation to the moon 318Moon-related reproductive activity 319Reproductive season and lunar-related spawning rhythm of rabbitfishes 320Ovarian development and the lunar cycle 321Testicular development and lunar periodicity 323Environmental factors regulating lunar-related reproduction 324Closing remarks 325 Abstract Cues from the moon influence synchrony in growth, feeding, migration, behaviour and reproduction of many reef fishes. Compared with comprehensive studies on the annual and daily activities of fish, few physiological studies have paid attention to the importance of lunar cues in reproductive activities. We review mutual and interesting relationships between fish reproduction and environmental changes induced by the moon, with particular emphasis on the reproductive activity of the rabbitfishes (Siganidae). Rabbitfish species exhibit, in nature, a definitive reproductive season, which differs among the tropical areas. During the reproductive season, synchronous spawning of rabbitfish is associated with a particular lunar phase. The lunar phase used by the respective species is similar in different regions on the earth. Histological observations revealed that gonads develop synchronously towards a peak around the spawning lunar phase, after which the gonads return to spent condition. Concomitant with gonadal development, sex steroid hormones were produced under the influence of gonadotropin (GtH). Injections of human chronic gonadotropin (hCG) to the fish that are undergoing active spermatogenesis accelerated testicular maturation. These results suggest that hormonal response in maturing the gonads in rabbitfish is under the regulation of GtH, and that pituitary secretion of GtH according to the lunar cycle accounts for the lunar rhythm in gonadal development. We speculate that the cues from the moon can be recognized by the higher parts of the hypothalamus-pituitary-gonadal axis. Possible relationships between exogenous environmental factors and the lunar-reproductive rhythm are also discussed.

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Rabbitfishes are known to spawn synchronously around the species-specific lunar phase. It is considered that they perceive and utilize cues from the moon in order to be synchronized gonadal development and spawning with the lunar cycle. Using the golden rabbitfish, Siganus guttatus, which spawns synchronously around the first quarter moon during the reproductive season, we measured the fluctuation of melatonin levels and examined the response of the fish to moonlight intensity. Daily fluctuation of melatonin concentration in the blood of golden rabbitfish showed low levels during daytime and high levels during night-time, suggesting that melatonin functions in the perception and utilization of photoperiod. Plasma melatonin concentration at the new moon was higher than that at the full moon. When the fish were exposed to moonlight at midnight of the both moon phases, the melatonin concentrations decreased to the control levels. These results show that the fish possibly perceive moonlight intensity and plasma melatonin fluctuates according to 'lightness' at a point of night. At the first spawning period (experiment was started one month before the spawning), the fish reared under natural conditions spawned at the expected spawning dates, whereas the fish reared under the constant darkness and lightness of night did not spawn. At the second spawning period (experiment was started 2 weeks before the spawning), the fish reared under the conditions of natural and constant darkness of night spawned but not that of constant lightness of night. It is possible that night conditions are related to synchronous gonadal development and spawning in the golden rabbitfish.

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Melatonin receptors are expressed in neural and peripheral tissues and mediate melatonin actions on the regulation of circadian rhythms in various species. For overall understanding of 'circa' rhythms in the golden rabbitfish, Siganus guttatus, which exhibits restricted lunar-related rhythms and spawns synchronously around the first quarter moon, the aim of the present study was to clone a melatonin receptor (Mel(lb)) cDNA and examine daily variations of Mel(lb) mRNA expression in certain tissues of the rabbitfish. The full-length Mel(lb) cDNA (1808 bp) contained an open reading frame to encode a protein with a length of 354 amino acids, which was highly homologous to a protein of nonmammalian species. Northern blot analysis showed transcripts of Mel(lb) in the brain and retina. Real-time quantitative polymerase chain reaction analysis also revealed expression of Mel(lb) in all tissues tested. Significantly high expression of the gene during daytime was evident in the liver and kidney. When the expression of Mel(lb) was examined in the brain and retina under conditions of light/dark cycles or constant darkness, daily and circadian variations of gene expression with two increases during daytime and nighttime for the brain and a single increase during nighttime for the retina were recognized. Moreover, daily variations in the expression of Mel(lb) were observed in the cultured pineal gland. These results suggest that the melatonin receptor plays a role in integration of melatonin actions in various tissues and that daily variations of Mel(lb) in the neural tissues may be related to regulation of circadian clock.