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Maternal behavior in F344/N and LEW/N rats. Effects on carrageenan-induced inflammatory reactivity and body weight.

Inbred Fischer (F344/N) and Lewis (LEW/N) rats differ on a myriad of behavioral and physiological endpoints, such as inflammatory, startle and drug responsivity. These differences point to underlying genetic differences between the strains. However, genetic models of hypertension have shown the importance of the maternal environment in the development of high blood pressure, suggesting that maternal influences might also play a role in adult phenotypes of the LEW/N and F344/N strains. This was tested in the present series of experiments in which the effects of crossfostering on carrageenan-induced inflammation and on body weight were examined in the two strains. Following the demonstration that the two strains differed in maternal behavior (Experiment 1), which was independent of the pup being reared (Experiment 2), crossfostered and in-fostered pups from the LEW/N and F344/N strains were injected with carrageenan (at 60 days of age) and subsequently assessed for the accumulation of exudate in response to the injection. Body weights were also monitored from birth through 60 days of age. Although crossfostering affected body weight of the two strains, specifically, reducing weights in LEW/N pups reared by F344/N dams and increasing weights of F344/N pups reared by LEW/N dams, crossfostering did not affect inflammatory reactivity to carrageenan. Specifically, LEW/N pups had a greater level of exudate than F344/N pups, independent of the conditions under which they were reared, suggesting that differences in the inflammatory response between these two strains are under a high degree of genetic control. These results were discussed in terms of genetic factors mediating the early form of immune reactivity induced by carrageenan.

Animals↗

A prospective study of the development of laterality: neonatal laterality in relation to perinatal factors and maternal behavior.

As part of an investigation of the development of laterality, 150 neonates were tested for bilateral asymmetries in behavior. Palmar and plantar grasping, and head-turning preference were measured, and observations were made of the way mothers held their babies and laid them in their cribs. Overall, right head-turning predominated. However, the side on which the baby had been lain pre-test markedly affected head-turning behavior: 120 of 147 babies turned predominantly to the right after lying on the right, whereas only 75 did so after lying on their left side (p < .001). There was no predominant asymmetry in the population in palmar or plantar grasping. These findings are explored in relation to perinatal factors and maternal behavior.

Female↗

Central nitric oxide synthase inhibition disrupts maternal behavior in the rat.

Blocking nitric oxide (NO) production, by 3rd ventricle administration of a nitric oxide synthase (NOS) inhibitor, N(G)-nitro-L-arginine methyl ester (L-NAME; 250 microg/5 microl, postpartum [pp]) decreased milk ejections in Day 10 pp rats. On Day 4 pp, L-NAME treatment eliminated pup retrieval and at both stages of lactation suppressed maternal aggression. Fewer rats treated with L-NAME on Day 10 pp retrieved 4-day-old pups than controls, although all nursed older litters. Following exposure to a mobile intruder, Fos expression was lower in the medial preoptic area and the bed nucleus of the stria terminalis in L-NAME-treated rats than in controls but was lower in the medial amygdala only following exposure to an anaesthetized intruder. Thus, the elevated levels of NO observed in lactation may contribute to the mechanism(s) that mediate maternal behavior and aggression.

Aggression↗

The influence of handling and isolation postweaning on open field, exploratory and maternal behavior of female rats.

Fifty-three black-hooded female rats, raised from the 28th to the 128th day of life under four types of environmental conditions differing in social and handling factors, had their behavioral performance evaluated in eight tests involving exploration of a complex environment (EX) in two open field (OF) tests and in three pup-retrieving tests. These tests were held before, during and after the period of lactation of their first brood. Females differed from (previously studied) males in that environmental raising conditions had a much smaller influence, especially during the lactation period. Nevertheless, females raised in isolation tended to remain for longer times inside a protected den in the exploration tests and to have greater locomotion in the open field. Previous handling allowed a greater exploration during the first contact with the novel complex environment as well as greater activity in the open field. The presence of small pups drastically reduced the exploratory motivation, but the presence of pups more than 20 days old tended to increase it. Maternal behavior evaluated in the pup retrieval test and in the test of time for the female to leave the nest side of the double box system was markedly refractory to previous environmental influences.

Analysis of Variance↗

Maternal behavior as a regulator of polyamine biosynthesis in brain and heart of the developing rat pup.

Rat pups removed from the mother and placed in a warm incubator for 1 hour or more show a 50 percent reduction in ornithine decarboxylase activity in the brain and heart. The decline is not caused by lack of nutrition. Instead, these studies suggest that active maternal behavior is necessary to maintain normal polyamine metabolism in brain and heart of the pup during development.

Adrenalectomy↗

Maternal behavior, paternal behavior, and pup survival in CD-1 albino mice (Mus musculus) in three different housing conditions.

Parental behavior, food consumption, and pup development in litters of CD-1 albino mice (Mus musculus) with single mothers or paired mothers in 3 different housing conditions were compared. Fewer pups survived when parents were required to run in a wheel to obtain food, and these pups had lower weights than pups in the wheel-noncontingent and no-wheel groups. The presence of the male facilitated pup survival in the wheel-contingent group but had no effect on pup growth. Mice in the wheel-contingent group ran more revolutions than those in the wheel-noncontingent group and spent less time in parental care. In the wheel-contingent group, males spent more time running and ate less food than females, suggesting that males may indirectly facilitate pup development by providing food for the mother. The results suggest that the presence of the male increases pup care and may facilitate maternal behavior under appropriate environmental conditions.

Animals↗

The ability of oxytocin to induce short latency maternal behavior is dependent on peripheral anosmia.

Nulliparous female Sprague-Dawley rats, cannulated in the left lateral ventricle, were ovariectomized and estrogen primed, then either rendered anosmic via intranasal irrigation with zinc sulfate or left with intact olfaction. Forty-eight hr later, after a 2-hr habituation to the test cage, these animals were injected with either intracerebroventricular oxytocin (400 ng in 2 microliter saline) or saline (2 microliter). Only the group receiving both zinc sulfate and oxytocin became maternal. Additionally, approximately one third of the olfaction-intact rats and none of the anosmic rats cannibalized the rat pups. These results are discussed in regard to discrepancies in the literature regarding oxytocin's role in inducing maternal behavior, as well as the functional connection of the olfactory and oxytocin systems.

Animals↗

Inhibitory effect of postpartum lesions or cuts in median raphe nucleus on maternal behavior in female rats.

In order to clarify the role of the median (MRN) and dorsal (DRN) raphe nuclei in regulating maternal care (retrieving and licking behavior), radiofrequency lesions or microknife cuts were made in postpartum rats on the day after delivery. Animals were housed individually without pups after the operation. One week after the surgery, maternal behavioral test was carried out daily for 3 days using pups of 2-6 days age. The results demonstrated that rats with MRN lesions or ventral horizontal cuts of the MRN showed extremely low incidence of the maternal behavior, as compared to those in control and sham-operated groups. DRN-lesions or dorsal cuts of the MRN had no effect. In locomotor activities measured by the infrared sensor system, there was no difference between the groups. This suggest that the MRN but not DRN plays a critical role in regulating retrieving and licking behaviors and ventral outputs are involved in this function in postpartum rats.

Animals↗

The differential effect of the anxiolytic agent 8-OH-DPAT during lactation is independent of pup withdrawal and maternal behavior.

Injection of the serotonergic agonist, 8-hydroxy-2-(di-n-propylamino-tetralin (8-OH-DPAT) (0.5 mg/kg ip) produced a clear anxiolytic-like effect (as measured in the burying behavior test), after parturition, which remains until day 6 of lactation. Thereafter 8-OH-DPAT completely lacked action. In order to analyze whether lactation prevented the action of 8-OH-DPAT, dams were separated from their pups for five consecutive days. The blockade of the anxiolytic effect of 8-OH-DPAT does not disappear by isolation of the mothers from their offspring or from neighboring pups. Finally, to investigate the possible role of maternal behavior in the blockade of the anxiolytic effect of 8-OH-DPAT a third experiment was made in which ovariectomized females were rendered maternal by the sensitization procedure. These females respond normally to the antianxiety actions of 8-OH-DPAT. Results suggest that a long-term clue triggered by lactation, possibly related to prolactin secretion, interferes with the anxiolytic effect of 8-OH-DPAT.

8-Hydroxy-2-(di-n-propylamino)tetralin↗

Nitric oxide prolongs parturition and inhibits maternal behavior in rats.

We examined the effect of nitric oxide (NO) on the process of parturition in rats. Subcutaneous injection of the NO donor sodium nitroprusside (SNP) in late pregnancy prolonged the total parturition time. The effect of N omega-nitro-L-arginine was not significantly different from that of saline. Intracerebroventricular injection of SNP in parturient rats delayed the progress of parturition. Both modes of SNP treatment also inhibited expression of maternal behavior. Central injection of oxytocin (OXT) with SNP failed to reduce parturition time significantly, but intrapartum, not postpartum, maternal behaviour was restored. These observations suggest that NO interferes with the release of OXT within the brain, hence affecting the initiation of maternal behaviour, and may also impair oxytocin secretion from the neurohypophysis.

Animals↗

Artificial rearing causes changes in maternal behavior and c-fos expression in juvenile female rats.

This study investigated the effects of early-rearing experiences on responsiveness to pups and on the pattern of c-fos activation in the brain of juvenile female rats. From Days 4 to 20, littermate females were reared with their mothers (MR) or artificially (AR). AR rats received minimal licking-like tactile stimulation (AR-min) or maximal stimulation (AR-max). On Day 20, rats were exposed to pups for 4 or 8 days, exposed to a playmate for 4 or 8 days, or left in isolation for 4 or 8 days. Compared with MR rats, pup-exposed AR rats engaged in less pup licking, and all AR rats showed significant reductions in c-fos immunoreactivity in the medial preoptic area and the parietal and piriform cortices. The AR-min group showed the greatest difference in Fos-lir compared with the MR groups. Possible mechanisms that mediate the effects of rearing on the development of neural circuits underlying maternal behavior are discussed.

Animals↗

Genetic and ethanol-related differences in maternal behavior and offspring viability in mice.

Primiparous female mice of 2 lines genetically selected for different narcotic responses to ethanol were administered 10% (v/v) ethanol in their drinking water during Days 2-14 of lactation. Ethanol treatment resulted in a marked decrease in growth and viability of pups in both lines. Fewer offspring of the ethanol-sensitive line survived to weaning, apparently because of the failure of a large number of dams to exhibit maternal care. The combination of ethanol treatment and an additional environment stressor (mouse hepatic virus) had a devastating effect on maternal behavior and offspring survival in the ethanol-sensitive line, but far less effect on the ethanol-insensitive mice. Thus, ethanol-sensitive animals appear to be more susceptible to a variety of stressors, including ethanol.

Alcoholic Intoxication↗

Dose-dependent effects of multiple acute cocaine injections on maternal behavior and aggression in Sprague-Dawley rats.

Rat dams, which had no prior drug treatment, were either nontreated controls or were injected subcutaneously 4 times during a 10-day period with a single dose of 30, 15 or 7.5 mg/kg of cocaine hydrochloride HCl, or normal saline. Injections were given immediately postpartum following delivery of their final pup (PPD 1), and again on postpartum day 3 (PPD 3), postpartum day 6 (PPD 6) and postpartum day 10 (PPD 10). Dams were observed 30 min following injections for maternal behavior (MB) towards 8 surrogate male pups on PPD 1 and PPD 3 and for aggression towards a male or female intruder in the presence of their litter on PPD 6 and PPD 10. Compared to saline and untreated controls, cocaine-treated dams exhibited more disruptions in MB on both PPD 1 and PPD 3 and were less aggressive towards an intruder, regardless of intruder sex, on PPD 6 and PPD 10. In most cases MB was altered in a dose-dependent manner with the higher doses of cocaine resulting in a greater disruption of behavior.

Aggression↗

Oxytocin activates the postpartum onset of rat maternal behavior in the ventral tegmental and medial preoptic areas.

Oxytocin binding (Bmax) was found to be higher in the ventral tegmental area (VTA) and the medial preoptic area (MPOA) at midparturition compared with Pregnancy Days 15-17 or Postpartum Days 5-7 in rat dams. Pup retrieval and assuming a nursing posture over pups were blocked in parturient dams by infusions of an oxytocin antagonist into the VTA or MPOA and by infusions of a vasopressin (V1) antagonist into the MPOA. These results implicate oxytocin in the VTA and MPOA and vasopressin in the MPOA, as well as a parturition-associated rise in oxytocin binding in these sites in the postpartum activation of maternal behavior.

Animals↗

Infant and maternal behaviors regulate infant reactivity to novelty at 6 months.

Three issues were investigated: (a) the regulatory effects of presumed infant and maternal regulation behaviors on infant distress to novelty at 6 months, (b) stability of infant regulatory effects across contexts that vary in maternal involvement, and (c) associations and temporal dynamics between infant and maternal regulation behaviors. Participants were 87 low-risk infants and their mothers, observed at 6 months postpartum during infant exposure to novel toys. Contingencies derived from sequential analyses demonstrate that, by 6 months, some infants reduce their own distress to novelty by looking away from the novel toy or self-soothing, maternal engagement and support have comparable effects, and certain infant and maternal behaviors co-occur. Moreover, infants whose mothers engaged contingently when they looked away from the novel toy expressed less distress than comparable infants whose mothers did not. These findings implicate both infants and mothers in the development of emotion regulation during the infant's first year.

Adult↗

Trigeminal lesions and maternal behavior in Norway rats: I. Effects of cutaneous rostral snout denervation on maintenance of nurturance and maternal aggression.

Cutaneous desensitization of the upper, rostral snout by bilateral section of the infraorbital nerves in lactating Norway rat dams markedly, but transiently, disrupted retrieval and licking of pups, nursing behavior, and fighting with a strange male intruder. Removal of the mystacial vibrissae, which provide major inputs to the infraorbital nerves, or cutaneous desensitization of the chin by bilateral sectioning of the mental nerves did not disrupt these behaviors. However, cutaneous desensitization of both the upper and lower rostral snout exacerbated and prolonged the effects produced by infraorbital denervation alone. The control of maternal behavior and aggression by somatosensory reflexes and the possible bases for rapid behavioral recovery after partial trigeminal sensory denervations are discussed.

Aggression↗

Maternal behavior and neuroendocrine regulation of suckling-mediated anovulation in cows.

Although it is clear that the initial endocrine deficit contributing to the length of the postpartum anovulatory period in suckled cows is of pituitary origin, this limitation is only operative for about the first 2 weeks of the puerperium. Thereafter, the pattern of LH secretion required for the final stages of follicular development and ovulation is blocked or attenuated through a more centrally regulated phenomenon. Hence, much of the recently published work in this area has focused on regulatory processes within the hypothalamus. This focus ignores the critical issue of exteroceptive signalling. Recent observations in this laboratory suggest that exteroceptive cues responsible for suckling-mediated anovulation are specifically attributable to the dams' own calf, but are not teat-specific. In fact, suckling does not produce an extended anovulatory state in the absence of a maternal-offspring bond. This review provides an historical perspective of investigations leading to this conclusion, and proposes a conceptual working model that links the special senses, maternal behavior and neuroendocrine centers that drive gonadotropin secretion during the puerperium.

Animals↗

Variations in nucleus accumbens dopamine associated with individual differences in maternal behavior in the rat.

Lactating rats exhibit stable individual differences in pup licking/grooming. We used in vivo voltammetry to monitor changes in extracellular dopamine (DA) in the nucleus accumbens (n. Acc) shell of lactating rats interacting with pups and found that (1) the DA signal increased significantly with pup licking/grooming; (2) the onset of such increases preceded pup licking/grooming; and (3) the magnitude and duration of the increase in the DA signal were significantly correlated with the duration of the licking/grooming bout. In females characterized on the basis of behavioral observations as high-licking/grooming mothers, the magnitude of the increase in the DA signal associated with licking/grooming was significantly greater than in low-licking/grooming dams. Dopamine transporter binding in the n. Acc was increased in low-compared with high-licking/grooming mothers. Injection of the selective DA uptake inhibitor GBR 12909 [1-(2-(Bis-(4-fluorophenyl)methoxy)ethyl)-4-(3 phenypropyl)piperazine dihydrochloride] (5 mg/kg, s.c.) increased the DA signal in the n. Acc and pup licking/grooming in low-licking/grooming mothers to levels comparable with those observed in high-licking/grooming dams. Receptor autoradiographic studies showed elevated levels of D1 and D3 receptors in the n. Acc shell region in high-licking/grooming dams. These results suggest that high- and low-licking/grooming dams differ in mesolimbic dopaminergic activity associated with mother-pup interactions. Such differences may serve as neural substrates for individual differences in the motivational component of maternal behavior.

Animals↗