External and internal controls of lunar-related reproductive rhythms in fishes.

Reproductive activities of many fish species are, to some extent, entrained to cues from the moon. During the spawning season, synchronous spawning is repeated at intervals of c. 1 month (lunar spawning cycle) and 2 weeks (semi-lunar spawning cycle) or daily according to tidal changes (tidal spawning cycle). In species showing lunar-related spawning cycles, oocytes in the ovary develop towards and mature around a specific moon phase for lunar spawners, around spring tides for semi-lunar spawners and at daytime high tides for tidal spawners. The production of sex steroid hormones also changes in accordance with synchronous oocyte development. Since the production of the steroid hormones with lunar-related reproductive periodicity is regulated by gonadotropins, it is considered that the higher parts of the hypothalamus-pituitary-gonad axis play important roles in the perception and regulation of lunar-related periodicity. It is likely that fishes perceive cues from the moon by sensory organs; however, it is still unknown how lunar cues are transduced as an endogenous rhythm exerting lunar-related spawning rhythmicity. Recent research has revealed that melatonin fluctuated according to the brightness at night, magnetic fields and the tidal cycle. In addition, cyclic changes in hydrostatic pressure had an effect on monoamine contents in the brain. These factors may be indirectly related to the exertion of lunar-related periodicity. Molecular approaches have revealed that mRNA expressions of light-sensitive clock genes change with moonlight, suggesting that brightness at night plays a role in phase-shifting or resetting of biological clocks. Some species may have evolved biological clocks in relation to lunar cycles, although it is still not known how lunar periodicities are endogenously regulated in fishes. This review demonstrates that lunar-related periodicity is utilized and incorporated by ecological and physiological mechanisms governing the reproductive success of fishes.