Photo-neuroendocrine control of seasonal cycles in body weight, pelage growth and reproduction: lessons from the HPD sheep model.

This chapter summarises the results of a recent study which investigated the role of the hypothalamo-pituitary relay system in mediating the effects of photoperiod on seasonal cycles in: (a) body weight; (b) pelage growth; and (c) reproduction in Soay rams. Hypothalamo-pituitary disconnected (HPD) and the control rams were housed indoors under an artificial lighting regimen of alternating 16-weekly periods. These periods consisted of long (16L:8D) and short days (8L:16D) and lasted for more than 2 years. The: (i) body weight; (ii) voluntary food intake; (iii) pelage and horn growth; and (iv) variations in testicular diameter were measured routinely every 2-4 weeks. Twice-weekly blood samples were collected to monitor long-term changes in the blood concentrations of: (1) pituitary; (2) metabolic; and (3) reproductive hormones (prolactin, GH, alpha-MSH, beta-endorphin, ACTH, TSH, LH, FSH, cortisol, insulin, IGF1 and testosterone). In control rams there were clearly defined photoperiod-induced cycles in blood concentrations of prolactin, alpha-MSH, beta-endorphin, LH, FSH, insulin and testosterone and associated morphological changes consistent with causal relationships (e.g. prolactin versus wool and horn growth, alpha-MSH, beta-endorphin and insulin versus body weight/food intake, LH and FSH versus testis size). In the HPD rams there were no photoperiod-induced cycles in the concentrations of any of the pituitary hormones with the exception of prolactin which varied as in controls (10-fold higher under long days). There was a permanent increase in blood concentrations of alpha-MSH, beta-endorphin and insulin in the HPD animals and a decrease in the concentrations of GH (loss of pulsatility) and IGF1. These changes were associated with the development of obesity. The reproductive axis was inactivated (basal LH, FSH and testosterone) although there was residual cyclicity in the size of the testis associated with the changes in prolactin secretion. Overall, the results support the view that the melatonin signal which encodes photoperiod, acts in the hypothalamus to regulate some photoperiodic responses (alpha-MSH and beta-endorphin-body weight axis, gonadotrophin-gonadal axis) but acts in the pituitary gland to regulate other responses (prolactin-pelage axis). However, a functional hypothalamus is required to generate normal seasonal cycles in: (a) body weight; (b) food intake; (c) growth; (d) fattening; and (e) reproduction, to provide the internal coordination between different systems and to facilitate the temporal entrainment to environmental cues.