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Recent research on the circadian system of the zebrafish is reviewed. This teleost has become an attractive model system because of its advantages for genetic analyses. Circadian rhythms of zebrafish behavior, visual system function, and pineal melatonin synthesis have been described, and behavioral and pineal rhythms are being used to identify and characterize clock mutants. Zebrafish heart, kidney, and embryonic cell lines contain circadian oscillators and phototransduction mechanisms for entrainment, suggesting that circadian pacemaking functions may be distributed throughout the animal. Studies of circadian system development in zebrafish have found that a molecular circadian oscillation in unfertilized oocytes persists through embryonic development with its phase intact, but that the pacemakers that drive rhythms of melatonin synthesis and behavior require environmental entraining signals late in development for initial synchronization. Zebrafish homologs of several of the core clock genes identified in other animals have been cloned. Transcripts for most of these are rhythmically expressed in multiple tissues. The interactions of clock gene products are for the most part similar to their interactions in mammals, although there are some potentially interesting differences.

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Memory processes are modulated by the biological clock, although the mechanisms are unknown. Here, we report that in the diurnal zebrafish both learning and memory formation of an operant conditioning paradigm occur better during the day than during the night. Melatonin treatment during the day mimics the nighttime suppression of memory formation. Training in constant light improves nighttime memory formation while reducing endogenous melatonin concentrations. Treatment with melatonin receptor antagonists at night dramatically improves memory. Pinealectomy also significantly improves nighttime memory formation. We adduce that melatonin is both sufficient and necessary for poor memory formation during the night.

The signaling pathways that couple light photoreception to entrainment of the circadian clock have yet to be deciphered. Two prominent groups of candidates for the circadian photoreceptors are opsins (e.g., melanopsin) and blue-light photoreceptors (e.g., cryptochromes). We have previously showed that the zebrafish is an ideal model organism in which to study circadian regulation and light response in peripheral tissues. Here, we used the light-responsive zebrafish cell line Z3 to dissect the response of the clock gene zPer2 to light. We show that the MAPK (mitogen-activated protein kinase) pathway is essential for this response, although other signaling pathways may also play a role. Moreover, action spectrum analyses of zPer2 transcriptional response to monochromatic light demonstrate the involvement of a blue-light photoreceptor. The Cry1b and Cry3 cryptochromes constitute attractive candidates as photoreceptors in this setting. Our results establish a link between blue-light photoreceptors, probably cryptochromes, and the MAPK pathway to elicit light-induced transcriptional activation of clock genes.

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