Photoperiodism and circadian rhythms have been studied intensively in birds because Aves are typical seasonal breeders and diurnal animals. Light is the most important environmental factor involved in entrainment of circadian rhythms and photoperiodism. The eyes and the extraocular photoreceptors, such as the pineal organ and hypothalamus, are reported to have an important function not only for photoreception but also for circadian organization in nonmammalian vertebrates, including birds. In this report, we review the roles of the eyes, pineal organ, and deep brain as the components of the multiphotoreceptor and multioscillator system in avian circadian organization.

We investigated the photoperiodic response of serotonin- and galanin (GA)- immunoreactive (ir) cells in the paraventricular organ (PVO) and infundibular nucleus (IF) of the Japanese quail and the interaction of these cells with gonadotropin-releasing hormone (GnRH)-ir neurons in the hypothalamus. Serotonin-ir cells were located in series from the PVO to the IF, and were connected with each other. The number of serotonin-ir cells differed significantly between light and dark phases on the short days (SD), but did not differ between light and dark phases on long days (LD). GA-ir cells were also found in the PVO and IF. The number of GA-ir cells under SD conditions was significantly greater than under LD conditions but did not change diurnally. Both serotonin-ir and GA-ir fibers ran along the GnRH-ir cells in the nucleus commissurae pallii. Serotonin-ir and GA-ir fibers were connected with the GnRH-ir fibers in the external layer of the median eminence (ME). We confirmed that GA-ir fibers were closely associated with serotonin-ir neurons in the PVO and IF. GA-ir neurons have at least 2 routes of regulating GnRH neurons directly, and indirectly via the serotonin-ir cells in the PVO and IF.

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