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Environmental factors such as photoperiod play a regulatory role in the HPG axis via regulation of GnRH production (Porkka-Heiskanen et al., 1997). In view of its major role in the regulation of the GnRH neuron, kisspeptin neurons were hypothesised to relay photoperiod effects on GnRH neurons in seasonally breeding species. In seasonal breeders such as sheep, cattle and hamsters, activation of the reproductive axis is controlled by melatonin produced from the pineal gland, which is secreted at night only and therefore relays day length to the body (Bittman et al., 1985). Three animal models have been employed to look at these effects, the Siberian and Syrian hamsters (long day breeders), and sheep (short day breeders).

Female Siberian hamsters are sexually active during long days (LD), when melatonin secretion is reduced. During this time KiSS-1 levels in the ARC is decreased, while AVPV KiSS-1 level is increased, to give optimum conditions for conception (Mason et al., 2007). However, during short days (SD) when female Siberian hamsters are in anoestrus the opposite occurs, with high KiSS-1 levels in the ARC and low levels in the AVPV (Table 3) (Greives et al., 2006; Mason et al., 2007). In the Syrian Hamster, also a long day breeder, no staining was found in the AVPV, but an increase in KiSS-1 staining was seen in the ARC during LD and a decrease was seen during SD. This response to seasonal breeding appears to be a result of melatonin production as pineal gland ablation prevents the reduction in ARC KiSS-1 during SD (Revel et al., 2006). Thus melatonin is important for KiSS-1 regulation in both Siberian and Syrian hamsters but evidence of functional melatonin receptors on kisspeptin neuronal cells has not been demonstrated.

In the ewe, which is a short day breeder and so responds to a long melatonin secretion period, KiSS-1 mRNA in the ARC is increased during SD when estrus occurs, and ARC KiSS-1 expression is decreased during LDs for anoestrus (Table 3) (Smith et al., 2006b). This allows activation of the LH surge during estrus but inhibition during anoestrus (a mechanism which has evolved to co-ordinate reproduction with seasonal availability of food). However, seasonal endocrine physiology can be overcome with exogenous kisspeptin, which when given continuously for 30hrs can cause ovulation in 80% of ewes during anoestrus (Caraty et al., 2007). This shows that kisspeptin is a potent regulator of the HPG axis, even when the system is dampened.

Therefore, kisspeptin neurons represent a conserved mechanism for relaying photoperiodic cues in seasonal breeding animals. The neuronal networks and melatonin actions are yet to be delineated.

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