An investigation of the effects of maternal separation and novelty on central mechanisms mediating pituitary-adrenal activity in infant guinea pigs (Cavia porcellus).

In mammalian species in which the young exhibit a strong filial attachment (e.g., monkeys, guinea pigs), numerous studies have shown that even brief separation from the attachment figure potently elevates circulating concentrations of glucocorticoids and adrenocorticotropic hormone (ACTH). However, effects of separation on central regulation of this stress response are not known. Therefore, we investigated central mechanisms mediating pituitary-adrenal activation during maternal separation and novelty exposure in guinea pig (Cavia porcellus) pups. Corticotropin-releasing factor (CRF) mRNA expression in the hypothalamic paraventricular nucleus (PVN), and plasma cortisol and ACTH levels, were elevated only during separation in a novel environment. C-Fos activity was elevated in the medial amygdala (MeA) and reduced in the bed nucleus of the stria terminalis (BNST) during novelty exposure, regardless of separation. On the other hand, c-Fos activity was elevated in the PVN during separation, regardless of novelty exposure. These results demonstrate independent and combined effects of separation and novelty in regions of the guinea pig CNS that regulate pituitary-adrenal activity. Moreover, they suggest that a pathway from MeA to BNST to PVN mediates responses to novelty in the guinea pig pup, as in the adult rat, though inputs from other cell populations appear required to fully account for the HPA activity observed here.

Development of selective social buffering of the plasma cortisol response in laboratory-reared male guinea pigs (Cavia porcellus).

The authors examined the effect of different classes of females on the plasma cortisol response of male guinea pigs (Cavia porcellus) from shortly after weaning through full adulthood. Males housed under standard laboratory conditions with artificial harems of 2 unrelated females of similar age were tested at 4 age ranges (approximately Days 40, 120, 180, and 240). At each age range, males were placed into a novel environment for 2 hr on 4 separate occasions: while alone, with 1 female cage mate, with the other female cage mate, or with an unfamiliar female. Relative preference for the 2 female cage mates was determined from observations in the home cage. Puberty was estimated by plasma testosterone levels. At the 3 youngest ages, the more preferred, less preferred, and unfamiliar females did not differentially reduce the plasma cortisol response. At 240 days of age, only the presence of the more preferred female was able to significantly reduce cortisol levels. These results demonstrate a very selective social buffering effect on the plasma cortisol response in a nonmonogamous species. This effect emerges well after sexual maturity and can be observed under standard laboratory housing.

Social buffering of the cortisol response of adult female guinea pigs.

When housed in spacious, mixed age/sex colonies, male guinea pigs form small harems of females with which attachment-like social bonds are established. A previous study indicated that the bonded male, but not an unfamiliar male from the same colony, could reduce the plasma cortisol response of a female when exposed to a novel environment. We examined the effect of several classes of social partners on the cortisol response of females maintained under standard laboratory housing conditions. Eight "artificial harems" comprised of one adult male and two adult females were formed in standard group-housing cages 2 weeks prior to the study. Behavior observed in these home cages indicated that interactions between males and females of the triads were overwhelmingly positive in nature. When exposed to a novel environment for 2 h, the presence of either the male or female cage mate reduced the subject female's plasma cortisol response, whereas the presence of an unfamiliar adult male did not. Changes in plasma cortisol levels were not predicted by the frequency of various social behaviors observed in the novel environment. These findings: (1) demonstrate that the buffering effect of male partners on the cortisol response of adult females can be observed in guinea pigs maintained in conventional laboratory housing; (2) indicate that unfamiliar adult males are ineffective at reducing cortisol responses of females under these conditions; and, (3) appear to be the first to show buffering effects by adult female guinea pigs on the cortisol responses of other adult females.

Anti-inflammatory agents attenuate the passive responses of guinea pig pups: evidence for stress-induced sickness behavior during maternal separation.

A previous study found that intracerebroventricular (ICV) infusion of 25 microg of alpha-MSH reduced the passive responses (crouched stance, eye-closing, piloerection) of guinea pig pups during a 3-h isolation in a novel environment. Because alpha-MSH has broad anti-inflammatory properties, the results suggested that proinflammatory factors play a role in mediating the behavior of isolated infants. The present study further investigated this possibility. In Experiment 1, injection of lipopolysacchride (LPS) increased the number of 60-s intervals in which pups expressed the same three responses during a 1-h test, and ICV infusion of alpha-MSH significantly reduced the effect of LPS on crouching and piloerection. In Experiment 2, the prostaglandin synthesis inhibitor indomethacin (10 mg/kg) reduced the number of 60-s intervals in which pups exhibited both crouching and the full suite of passive responses during a 3-h isolation in a novel environment. Together these results provide further support for the hypothesis that the passive behaviors exhibited during prolonged isolation are "stress-induced sickness behaviors" mediated by proinflammatory factors.

Increases in the circulating testosterone of maturing male guinea pigs appear neither necessary nor sufficient for heightened maternally directed sexual and social/courtship behavior.

Periadolescent male guinea pigs housed continuously with their mother since birth exhibit little maternally directed sexual behavior. However, if rehoused apart from the mother for 24 h, they show elevations in circulating testosterone concentrations and display frequent sexual responses and increased social/courtship behavior upon reunion with her. We investigated the role of testosterone in the disinhibition of maternally directed sexual and social/courtship behavior. Subcutaneous implants of testosterone (Experiment 1) did not trigger maternally directed sexual behavior or an increase in social/courtship behavior among males housed continuously with their mothers. Further, neither blocking androgen receptors (Experiment 2) nor preventing the surge in testosterone (Experiment 3) prevented males housed without the mother from exhibiting increased maternally directed sexual and social/courtship behavior upon reunion. These findings indicate that the increase in testosterone that males exhibit when rehoused apart from the mother is neither sufficient nor necessary for the disinhibition of maternally directed sexual and social/courtship behavior observed when mother and son are reunited.

Social preferences of developing guinea pigs (Cavia porcellus) from the preweaning to the periadolescent periods.

Preference tests in a novel environment (Experiment 1) and unobtrusive observations in a specialized living environment (Experiment 2) examined the attractiveness of various classes of conspecifics for maturing guinea pigs (Cavia porcellus). It was found that (a). the young continued to remain near the mother well beyond weaning; (b). there was increased time spent with unrelated adult females, but not males, after weaning; (c). male and female offspring behaved similarly; and (d). littermates spent considerable time with each other. These results provide no evidence that guinea pigs approaching sexual maturity begin to associate preferentially with novel animals or potential breeding partners. Choices were largely predictable from earlier findings of the ability of various classes of conspecifics to reduce hypothalamic-pituitary-adrenal activity of the young.

Presence of mother and unfamiliar female alters levels of testosterone, progesterone, cortisol, adrenocorticotropin, and behavior in maturing Guinea pigs.

Although the guinea pig is characterized by precocial physical development and minimal active maternal care, studies suggest the presence of the mother can influence neuroendocrine and behavioral activity of offspring even well beyond weaning. Previous results may have been influenced by the procedure of housing weaned subjects with the mother to within 2 days of testing. The present study examined approximately 40-day-old guinea pigs housed apart from the mother for 0 (not rehoused), 2, or 10 days. Rehousing without the mother led to elevations in plasma testosterone (measured in males), progesterone (measured in females), cortisol, and adrenocorticotropin (ACTH) (both measured in males and females). Offspring housed without the mother for 10 days had the highest progesterone, cortisol, and ACTH levels. Testosterone elevations were observed in 2-day-, but not 10-day-, rehoused animals. Regardless of rehousing condition, 60 min isolation in a novel test cage elevated progesterone, cortisol, and ACTH, and reduced testosterone. These effects were all moderated if the subject was tested with the mother or another female. Sexual behavior toward the mother was observed frequently, but only in males housed apart from her prior to testing. Overall, males and females that had been housed apart from the mother interacted with her as they would an unfamiliar female. Our results corroborate previous findings, suggest the effect of housing apart from the mother on male testosterone is transitory, and indicate that continuous housing with the mother past weaning suppresses circulating progesterone in females and cortisol and ACTH in both sexes.