Dopamine-melatonin neurons in the avian hypothalamus controlling seasonal reproduction

Day length cues are used by temperate zone birds to time seasonal changes in reproductive physiology and behavior. However, the neuronal and neurochemical circuits used to measure day length (photoperiodic time measurement; PTM), transduce light information and activate the reproductive neuroendocrine system have not been definitely established. Recent findings from our laboratory provide data showing dopamine (DA) neurons within the premammillary nucleus (PMM) of the caudal turkey hypothalamus are putative photoreceptive neurons. These neurons reach threshold activation when a brief pulse of light is provided during the photo-inducible phase for photosexual stimulation. To further clarify the role of PMM neurons in coding daylight information, we showed that by using double-label immunocytochemistry (ICC) these neurons are immunoreactive (ir) to both tyrosine hydroxylase (TH; the rate limiting enzyme in DA biosynthesis) and melatonin (MEL). Moreover, we found these neurons to express tryptophan hydroxylase 1 (TPH1; the first enzyme in MEL biosynthesis) and 5-HT N-acetyltransferase (AANAT; a key regulatory enzyme in MEL synthesis) mRNAs but not neuronal tryptophan hydroxylase 2 mRNA (TPH 2; the rate limiting enzyme in 5-HT pathway). Both TH and TPH1 mRNAs were shown to cycle rhythmically, and with opposite phases, in PMM neurons of birds kept under a diurnal illumination cycle (12-h light/dark; LD). These neurons could also generate 24 h TH and TPH1 mRNA expression rhythms with the same phase relationship in constant light (LL) and constant dark (DD). In addition, the expression patterns and amplitudes of TH and TPH1 mRNAs were different between long and short photoperiods. These findings may form the basis for an endogenous dual-oscillator circadian system within PMM DA-MEL co-localized neurons controlling reproductive seasonality in birds.