Steroids as local regulators of ovarian activity in domestic animals.

The presented overview gives clear evidence for steroids as local regulators of follicular and luteal activity. In the follicle, estrogen receptor-alpha (ERalpha) and ERbeta expression are demonstrated in cow, ewe and pig. Besides species specific effects in general, there is evidence that estradiol-17beta (E(2)) exerts a dose-dependent inhibition on the secretion of progesterone (P(4)) by both theca interna cells (TI) and granulosa cells (GC). GC enhance the ability of the TI to produce androstendione by supplying them with progestin precursor. Androgen produced by TI enhances the ability of the GC to make E(2), and high concentrations of E(2) in the preovulatory follicle inhibit 3beta-HSD in both TI and GC and thus, may promote the use of the pathway Delta(5) for TI androgen production. The authors suggest that E(2) acts within the follicle to exert positive feedback on androgen and E(2) production, and exerts mitotic and anti-atretic or anti-apoptotic effects on follicular cells. Parts of the E(2)-mediated local action are regulated by stimulating effects on hormone receptors (LH, FSH, oxytocin). Gap junctions permit transfer of nutrients and cytokines to and from the avascular GC and oocyte, and formation is stimulated by estrogens. In bovine corpus luteum (CL) there is evidence that P(4) may directly regulate the production of P(4), oxytocin and prostaglandins (PGs) in a cycle dependent fashion. In most of domestic animal species, there is clear evidence for CL production of E(2) with clear stimulatory and luteotropic effects on P(4), and an intraluteal circuit that involves paracrine effects of E(2), oxytocin and PGF(2alpha) (especially in pigs). In contrast, there are species (ruminants, mares) in which the evidence for important local effects of E(2) is less clear, although expression of ERalpha, ERbeta and progesterone receptor (PR) is documented. Progesterone is very important for the regulation of CL lifetime by effects on the endometrium and release of the luteolytic signal PGF(2alpha). In conclusion, steroids as local regulators of ovarian activity are now documented and may stimulate further research in this field.