Deconstructing early life experiences: distinguishing the contributions of prenatal and postnatal factors to adult male sexual behavior in the rat.

In rodents, a pup's experiences in utero and during postnatal development shape its sexual behavior as an adult and how it is perceived by potential mates. We show that the male rat's sexuality is primarily influenced by the postnatal sex ratio of its litter, but not by the litter's prenatal intrauterine sex ratio or the behavior of its mother. Pups from litters with differing prenatal sex ratios were divided into litters with differing postnatal sex ratios. We found that males raised in a female-biased litter exhibited less mounting than males raised in either a male-biased litter or one with an equal sex ratio, and were less attractive to sexually receptive females, eliciting fewer soliciting behaviors, such as hop-darts, and fewer lordosis behaviors. However, the number of intromissions and ejaculations did not differ across groups, which suggests that males from female-biased litters mate as efficiently as males raised in other sex ratios, but do not require as many mounts to do so. The reported differences in sexual behavior did not vary with the quality of maternal behavior or with sexual experience in adulthood.

Effects of early deprivation and maternal separation on pup-directed behavior and HPA axis measures in the juvenile female rat.

Juvenile female rats show maternal-like behavior toward pups. The purpose of the following experiment was to investigate whether the HPA axis, through the use of early separation manipulations that alter HPA functioning in rats, plays a role in the juvenile response to foster pups. Female rats were early deprived or maternally separated for 5 hours daily from PND 2 to 14 and compared to animal facility-reared rats. Deprivation or separation increased CRH-R1 IR in the juvenile PVN, but had no other effects on other HPA measures or on maternal behavior. Pup-exposure during the juvenile period blunted corticosterone levels after acute and repeated pup-exposures when compared to exposure to novelty and conspecifics respectively. Repeated exposures to pups also increased CRH-R1 IR relative to isolation during the juvenile period. Overall, the data suggest that although pup-exposure affects corticosterone levels, the HPA axis does not relate to juvenile maternal behavior in the present experiments.

Early deprivation, but not maternal separation, attenuates rise in corticosterone levels after exposure to a novel environment in both juvenile and adult female rats.

Separation from the maternal nest alters the hypothalamic-pituitary-adrenal (HPA) axis stress response in adult male rats, but little research has addressed how separation affects female rats. The following experiments investigated how early maternal separation from postnatal day (PND) 2 to 14 affected stress-induced corticosterone and ACTH after exposure to an open field in juvenile and adult female rats. Female rats were separated for 5 h daily from mother and littermates (early deprivation: ED), separated from mother but not littermates (maternal separation: MS), or animal facility reared (AFR). Male siblings were left with the mother rat during separation. Female rats were exposed to an open field arena either during the juvenile period (PND 30) or during adulthood (PND 80-100). Results show that ED juvenile female rats showed a lower corticosterone stress response than MS and AFR female rats when measured at 5 min post-stress, but no difference at 20 or 60 min post-stress. In adulthood, ED female rats showed comparable elevations of corticosterone as MS and AFR rats at 5 min post-stress but lower elevations at 20 min. In terms of behavior, there were no significant effects of early experience. However, in adulthood, ED and MS rats tended to show a decreased proportion of inner grid crossings of the open field compared to AFR rats, suggesting a tendency for increased anxiety in these two separation groups.

The effects of adrenalectomy and corticosterone replacement on induction of maternal behavior in the virgin female rat.

Maternal behavior of the sensitized virgin rat is affected by approach-avoidance systems as well as by hypothalamic-pituitary-adrenal (HPA) axis, which is also activated during stress. The present experiments investigated the effects of adrenalectomy and of varying corticosterone concentrations on the onset and expression of maternal behavior in sensitized virgin rats. In the first experiment, latency to onset of maternal behavior and time spent licking once maternal were positively related to endogenous levels of corticosterone. However, few rats showed licking. In the second experiment, virgin rats were adrenalectomized or given sham surgeries before being sensitized and being given 0, 25, 100, 300, or 500 microg/mL of corticosterone in their drinking water. In the third experiment, virgin rats were adrenalectomized or given sham surgeries and given either control or corticosterone time-release pellets after being sensitized. Maternal behavior was then tested. Adrenalectomy increased licking in the second experiment and time over pups in the third experiment. Corticosterone replacement reduced licking in the second experiment and both licking and time over pups in the third experiment. In conclusion, exogenous corticosterone had an inhibitory effect on the expression of maternal behavior in the sensitized virgin rat, unlike the facilitatory effect previously found in the postpartum rat.

The effects of adrenalectomy and corticosterone replacement on maternal memory in postpartum rats.

Hormones associated with parturition prime rats to behave maternally, although hormonal changes are not necessary for these behaviors to occur. Experience with pups after birth enhances maternal responsiveness after a period of isolation, creating a maternal memory. The purpose of this study was to determine the role of corticosterone in the formation of maternal memory. Adrenalectomy or sham surgeries were performed in late gestation with corticosterone or vehicle pellets being given to adrenalectomized rats. Pups were removed immediately following parturition, and half of the rats received 4 h of pup experience, while the other half received only brief pup experience associated with parturition. Ten days following pup experience, foster pups were given to all rats. Latency to become maternal and maternal behaviors on the first 2 days of re-exposure and the first two maternal days were recorded. Among adrenalectomized rats given corticosterone, 4-h experience with pups decreased maternal latency when compared to brief experience with pups. This maternal experience effect was not found in comparisons between adrenalectomized rats not given corticosterone. In addition, corticosterone decreased latencies regardless of pup experience. Corticosterone also increased maternal behavior upon initial exposure to foster pups. In conclusion, corticosterone enhanced maternal memory and initial maternal behavior in postpartum rats.

The effects of adrenalectomy and corticosterone replacement on maternal behavior in the postpartum rat.

It is well known that the hypothalamic-pituitary-adrenal (HPA) axis is activated during stress. Recent work suggests it is also implicated in the regulation of "normal" behaviors. The present studies investigated the effects of adrenalectomy and of varying glucocorticoid concentrations on adult maternal behavior in primiparous rats. In two studies, rats in late pregnancy were adrenalectomized or given sham surgeries and were tested for maternal behavior. In the first study, primiparous rats were given 0, 25, 100, 300, or 500 microg/ml of corticosterone in their drinking water. In the second study, primiparous rats were given either control or corticosterone time-release pellets. Blood samples were taken to ensure that rats demonstrated levels of corticosterone in blood that were relative to doses received. In studies one and two, primiparous adrenalectomized rats showed slightly, but significantly, lower levels of some maternal behaviors, including licking and time in nest, than primiparous sham rats. Primiparous rats given higher doses of corticosterone replacement showed higher levels of these maternal behaviors than primiparous rats given lower doses of corticosterone. In conclusion, adrenalectomy decreases, but does not abolish, maternal behavior. Corticosterone replacement reverses these effects. Corticosterone is not necessary for the initiation or maintenance of maternal behavior but plays a role in the modulation of ongoing maternal behavior.