Sexual dimorphism in the vomeronasal pathway and sex differences in reproductive behaviors.

Several years ago we hypothesized that the vomeronasal system (VNS), a complex neural network involved in the control of reproductive behavior, might be sexually dimorphic. This hypothesis sprung from several facts; (a) the existence of steroid receptors in the VNS; (b) sexual dimorphism was already described in some structures that receive vomeronasal input, such as the medial preoptic area, the ventromedial hypothalamic nucleus, the ventral region of the premammillary nucleus and the medial amygdaloid nucleus; and (c) the vomeronasal organ, which is the receptor organ of the VNS, was also sexually dimorphic. After that point, the accessory olfactory bulb (AOB), the bed nucleus of the accessory olfactory tract (BAOT) and the bed nucleus of the stria terminalis were found to be sexually dimorphic. The aim of the present review is to show the experimental facts that confirm our earlier hypothesis and, consequently, to present the existence of a sexually dimorphic multisynaptic pathway for the first time in mammals. Sexual dimorphism in the VNS might provide a comprehensive approach to understanding the neural bases of sexually dimorphic reproductive behavior and it is suggested here that the greater number of neurons which male rats present in relation to females in most VNS structures might contribute to the inhibition of the expression of feminine copulatory behavior (lordosis) and maternal behavior in males. In addition, the mechanisms that control the development of sexual dimorphism in the VNS are discussed. The discussion takes into account the two patterns of sexual dimorphism found in the rat brain. Estrogens seem to promote the development of sexual dimorphism in both male and female rats. However, an inhibitory role of androgens might be necessary to hypothesize when males or females present a lower number of neurons and/or volume than the opposite sex. There are experimental data supporting this hypothesis in the female, since dihydrotestosterone seems to facilitate neuronal death in VNS structures, such as the AOB and the BAOT, in which females present a lower number of neurons and volume than male rats. Finally, since the lateral division of the bed nucleus of the stria terminalis, which belongs to the main olfactory system (MOS), is sexually dimorphic and presents anatomical relationships with some VNS structures the MOS might be sexually dimorphic.