is, by ligand-or steroid-independent activation. More importantly, ligand-independent activation of steroid receptors (that is, progestin receptors) within the brain can influence reproductive behaviour in rodents. This review focuses on recent findings which indicate that some somatosensory cues normally experienced by females, such as those associated with sexual contact with males, can activate progestin receptors to influence both neuronal response and oestrous behaviour in a ligand-independent manner.
Hormonal requirements for the induction of sexual behaviourDuring the oestrous cycle in rats, sexual behaviour is dependent upon an increase in serum oestradiol followed by an increase in progesterone concentrations. Sexual behaviour is abolished by ovariectomy, and reinstated by the injection of oestradiol followed 1 or 2 days later by progesterone. Treatment with oestradiol alone can induce some aspects of sexual behaviour, such as receptivity (for example, lordosis); however, oestradiol followed by progesterone administration induces the full complement of proceptive behaviours that occur during normal oestrus, such as ear wiggling, hopping and darting behaviour (Blaustein and Olster, 1989). This finding indicates that both oestradiol and progesterone are critical for the occurrence of normal oestrous behaviour. Oestradiol and progesterone are secreted from both the ovaries and the adrenal glands (Shaikh and Shaikh, 1975). Steroid hormone secretion from the adrenal glands appears to be important for the normal timing of sexual behaviour in rats, as stress resulting from sham ovariectomy advances, and stress resulting from adrenalectomy delays, the onset of oestrous behaviour. In addition, the small increase in solicitational behaviour induced by oestradiol treatment alone is blocked by adrenalectomy. Therefore, sexual behaviour is influenced by secretion of steroid hormones not only from the ovary, but also from the adrenal glands.
Behavioural and physiological changes associated with mating stimulationWhen mating occurs, the female rat experiences a variety of somatosensory and olfactory stimuli from the male that further influences female sexual behaviour (Pfaff et al., 1994). For example, ovariectomized rats primed with low doses of oestradiol exhibit reduced lordosis in the presence of a sexually active male; however, repeated exposure of ovariectomized oestradiol-primed female rats to males will induce increased lordosis (Hardy and Debold, 1973). The increase in lordosis owing to repeated mating of oestradiolprimed female rats occurs within an hour, and this behavioural phenomenon is not dependent upon the adrenal glands (Foreman and Moss, 1977). Therefore, mating-related stimuli increase sexual receptivity in the absence of circulating progesterone. It is not clear which somatosensory and olfactory cues are important for mediat-