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PubMed2001

Decreases in calmodulin binding proteins and calmodulin dependent protein phosphorylation in the medial preoptic area at the onset of maternal behavior in the rat.

The onset of maternal behavior is characterized by the action of certain hormones, neuropeptides and neurotransmitters and a concomitant increase in the expression of c-Fos in the medial preoptic area (MPOA) but the signaling events that lie between have not been characterized. Because several of these hormones, neuropeptides and neurotransmitters function by activating Ca(2+)/calmodulin (CaM) mediated signaling pathways, many of which can lead to c-Fos expression, the goal of the current work was to identify calmodulin binding proteins (CaMBPs) or specific CaM-dependent phosphoproteins that might be involved. Probing of SDS-PAGE gels of extracts from the hippocampus, parietal cortex, basolateral amygdala and MPOA with recombinant (35)S-VU1-calmodulin (CaM) revealed 30 Ca(2+)-dependent and 4-6 Ca(2+)-independent CaMBPs. Statistically significant maternal behavior-related decreases in four Ca(2+)-dependent CaMBPs ( approximately 31 kDa, 50% decrease; approximately 33 kDa, 32%; approximately 50 kDa, 35%; approximately 60 kDa, 33%) were observed specifically in the MPOA. Numerous proteins were phosphorylated in a Ca(2+) CaM-dependent manner with two (MWs approximately 61 Da, approximately 58 kDa) showing a lack of phosphophorylation only in the MPOA. The selective decrease in CaMBPs coupled with the absence of CaM-dependent phosphoproteins implies that changes in Ca(2+)/CaM-mediated signaling may mediate some of the MPOA-specific processes during the onset of maternal behavior in the rat.

Amygdala↗

PubMed2001

Physiological regulation of maternal behavior in heifers: roles of genital stimulation, intracerebral oxytocin release, and ovarian steroids.

We tested the hypotheses that 1) epidural anesthesia at parturition would block both peripheral and central release of oxytocin and eliminate the development of maternal behavior in primiparous heifers and 2) estradiol priming, genital stimulation, and appropriate neonatal stimuli would induce maternal behavior in nulliparous heifers. In experiment 1, primiparous crossbred heifers (n = 13) with cannulas in the third cerebroventricle (IIIV) were assigned randomly to receive epidural treatments of saline (SAL; n = 6) or lidocaine HCl (EPI; n = 7) at the onset of labor induced between Days 270 and 280 of gestation. Epidural anesthesia blocked (P < 0.001) both central and peripheral release of oxytocin and markedly reduced (P < 0.05) or eliminated licking behaviors during a 3-h period following parturition as compared with SAL. Following approximately 1 wk of controlled daily suckling, during which calves were permitted access only to the inguinal region of their dams (three times daily for 10 min each time), a second maternal behavior test was performed. Although licking behavior remained markedly reduced (P < 0.001) in the EPI compared with the SAL groups, all heifers accepted their calf at the udder. In experiments 2-4, neither estradiol priming in ovariectomized heifers nor estradiol plus progesterone in intact heifers resulted in an induction of maternal behaviors following genital stimulation and presentation of a neonate wetted with amniotic fluid. Pelvic sensory deficits apparently block oxytocin release and disturb both short-latency and long-term maternal behaviors but do not result ultimately in rejection of the calf. Combinations of hormonal, sensory, olfactory, and visual cues observed previously to induce maternal behavior in nulliparous ewes do not appear adequate for induction of maternal behavior in nulliparous heifers.

Animals↗

PubMed1996

Opioid stimulation in the ventral tegmental area facilitates the onset of maternal behavior in rats.

This research investigated the effect of an increase or decrease in opioid activity in the ventral tegmental area (VTA) on the onset of maternal behavior in rats. In Experiment 1, the latency to show maternal behavior toward foster rat pups (sensitization latency) was determined in maternally naive female rats given either nothing or a unilateral intra-VTA injection of morphine sulfate (MS) (0.0, 0.01, 0.03, 0.1 or 0.3 microgram), on the first three days of a 10-day period of constant exposure to pups. Rats treated with 0.03 microgram MS had significantly shorter sensitization latencies than did rats treated with 0.0 microgram MS, 0.01 microgram MS, or receiving no treatment (higher doses of morphine produced intermediate results). The facilitating effect of intra-VTA MS on the onset of maternal behavior was blocked by pretreatment with naltrexone hydrochloride and was found to have a specific site of action in the VTA (MS injections dorsal to the VTA were ineffective). In Experiment 2, sensitization latencies were determined in periparturitional rats given a bilateral intra-VTA injection of either the opioid antagonist naltrexone methobromide (quaternary naltrexone), its vehicle, a sham injection, or left untreated 40 min after delivery of the last pup. The mothers' own pups were removed at delivery; mothers were nonmaternal at the time of testing. Quaternary naltrexone treatment produced significantly slower sensitization to foster pups than did control conditions. Total activity and pup-directed activity did not differ significantly with treatment. The results demonstrate that increased opioid activity in the VTA facilitates the onset of maternal behavior in inexperienced nonpregnant female rats, and decreased opioid activity in the VTA disrupts the rapid onset of maternal behavior at parturition.

Analgesics, Opioid↗

PubMed1996

Early postnatal alcohol exposure in rats: maternal behavior and estradiol levels.

Long Evans male and female rats were exposed to alcohol or control conditions during the early postnatal period, and their maternal behavior was assessed between 25 and 29 days of age. Maternal behavior was induced by exposing experimental animals to pups, and behaviors included retrieval of pups and proximity to pups. After 5 days of continuous exposure to pups, experimental animals were decapitated, and trunk blood was measured for levels of estradiol and progesterone. Alcohol-exposed animals showed a deficit in maternal behavior as measured by a longer latency to retrieve pups than rats that were not treated with alcohol. In addition, alcohol-exposed animals had lower levels of circulating estradiol than controls. These effects may also be explained by undernutrition as a result of artificial rearing. No difference in progesterone levels was found. As estradiol is important for the onset of maternal behavior in adult rats, it may be that the alcohol-induced decrease in estradiol levels underlies the deficit in maternal behavior exhibited by alcohol-exposed rats.

Animals↗

PubMed2004

Bisphenol-A exposure during pregnancy and lactation affects maternal behavior in rats.

In mammals, endogenous estrogens are crucial for sexual differentiation during the perinatal period, and the modulation in adulthood of many neuroendocrine and behavioral functions involved in reproduction. In rats, the estrogenic environment during pregnancy and lactation affects directly maternal behavior. This experiment was aimed to test whether the exposure to the estrogenic compound bisphenol-A (BPA; 0.040 mg/kg/die, orally) of adult female rats, from mating to weaning of the pups, could alter maternal behavior. An appropriate methodology was applied to reveal differences in the behavior of dams directed to male and female pups, testing the dams on postnatal days 3-4 and 8-9. Results show different maternal behavioral patterns towards male and female pups of control mothers, with more ano-genital licking to males than to females. Exposure of mothers to BPA modified their behavior, reducing specific components of maternal behavior, both active and passive, irrespective of the sex of pups and the period of observation. This experiment shows that maternal behavior is affected by a prolonged exposure to a low dose of BPA during pregnancy and lactation, thus suggesting an effect on neural circuits in adulthood.

Animals↗

PubMed2002

Restraint stress impaired maternal behavior in female mice lacking the neuromedin B receptor (NMB-R) gene.

Neuromedin B (NMB) is a bombesin-like peptide that exerts its function via NMB receptor (NMB-R). The NMB/NMB-R pathway is involved in the regulation of a wide variety of behaviors, such as spontaneous activity, feeding, and anxiety-related behavior. In the current study, we assessed the effects of stress on maternal behavior in female NMB-R-deficient mice. Non-stressed NMB-R-deficient and wild-type mice showed normal maternal behavior. However, immediately after undergoing restraint-induced stress (30 min) both genotypes of mice exhibited severely decreased maternal behaviors. Furthermore, 30 min after stress induction, maternal behavior in wild-type mice recovered to near normal levels whereas that of NMB-R-deficient mice remained significantly lower. These results indicate that NMB-R-deficient mice suffer more severely from stress and suggest that dysfunction in the NMB/NMB-R pathway may constitute one of the risk factors of stress vulnerability.

Animals↗

PubMed1977

Uterine distention facilitates the onset of maternal behavior in pseudopregnant but not in cycling rats.

The latency to onset of maternal behavior toward foster pups was examined in maternally-naive female rats treated either with uterine distention, a sham procedure, or no uterine manipulation. Uterine distention was achieved by intrauterine injection of hypertonic saline. The treatments were applied to either cycling, Day 10 pseudopregnant, or Day 11 pseudopregnant-decidualized virgins. The latency to onset of maternal behavior for both pseudopregnant groups was significantly shorter than that for the nonpseudopregnant group, when uterine distention was applied. The results suggest that uterine distention during pregnancy (during high progesterone level) may bring about both pregnancy termination (delivery) and the almost immediate maternal behavior seen at parturition, by the same hormonal mechanism.

Animals↗

PubMed1980

Hormonal and nonhormonal regulation of maternal behavior: a theoretical survey.

There is a general uniformity in the hormones that regulate pregnancy among a large variety of mammals. There is also a division of maternal behavior into two main patterns: that shown by mammals that build nests for the altricial young and those that only briefly establish a birth site for the precocial young. Apart from this difference, the patterns of maternal behavior are quite similar, centering around nursing, and weaning follows a similar course in different species. Is there a corresponding uniformity in the factors which regulate maternal behavior? Too few species have been studied to establish any general rule about the hormonal basis for the onset of maternal behavior. In the main, however, the ovarian hormones probably play a major role in most species and prolactin appears to play a role in at least one species. Post-partum maternal behavior appears to depend mainly upon stimulation from the young and hormones may not be required to maintain it or be involved in its decline. The transition period between hormonal and nonhormonal regulation of maternal behavior may emerge as an important phenomenon in many species.

Animals↗

PubMed2003

Long-term effects of prenatal morphine exposure on maternal behaviors differ from the effects of direct chronic morphine treatment.

Previous studies have demonstrated that chronic morphine treatment of pregnant rats alters maternal behavior. Other studies have shown long-term effects of prenatal exposure to morphine, including changes in reproductive behavior in adult females. The present study investigated the effects of prenatal morphine exposure on a variety of maternal behaviors such as nursing, maternal activities, nonmaternal activities, and pup retrieval. Prenatal morphine exposure increased active and decreased passive nursing. There were no differences in maternal activities such as presence in the nest, contact with pups, grooming of pups, and/or manipulation of nest shavings. In the retrieval test, prenatally morphine-exposed mothers were faster in carrying the first pup, retrieving the first pup back to the nest, and returning all pups to the nest than prenatally saline-exposed mothers. Maternal and nonmaternal activities also were affected by the light: dark cycle. All saline- and morphine-exposed mothers nursed more, were more often in the nest, and more often in contact with greater than half of their litter during the light than the dark sessions. On the other hand, nonmaternal activities increased during the dark sessions: Mothers cared for themselves (groomed, ate) more and displayed more rearing and sniffing. Mothers spent more time resting with their eyes closed during the light sessions regardless of prenatal drug exposure. The present study demonstrated that the effect of morphine on maternal behavior is different in adult exposed and prenatally exposed mothers. While direct morphine treatment impaired maternal behaviors, prenatal morphine exposure has the opposite effect.

Animals↗

PubMed1971

Olfactory bulb removal eliminates maternal behavior in the mouse.

Olfactory bulb removal eliminated maternal behavior in lactating and in virgin mice. The results are in contrast to the generally accepted concept of multisensory control of mammalian maternal behavior.

Animals↗

PubMed2002

Dopamine antagonists during parturition disrupt maternal care and the retention of maternal behavior in rats.

Brief contact with pups at parturition enables the female rat to establish and retain the full repertoire of maternal behaviors, allowing her to respond rapidly to pups in the future. To determine whether the dopamine system is involved in the retention of maternal behavior, females were continuously infused with dopamine antagonists during the periparturitional period and then allowed either a brief interaction period with pups (3 h) or no interaction with pups (pups removed as they were born). Females were exposed to either the D1-like antagonist SCH 23390 (0.1 or 1.0 mg/kg/day) or the D2-like antagonist clebopride (0.5 or 1.0 mg/kg/day). The high dose of either DA antagonist resulted in significant attenuation of maternal care immediately postpartum. When tested for the retention of maternal behavior 7 days later, however, only the females exposed to the D2 antagonist displayed a delayed response to shown full maternal behavior (FMB) towards donor pups. Thus, while both dopamine receptor subtypes appear necessary for the full and rapid expression of maternal behavior during the early postpartum period, only the D2 receptor subtype appears to be involved in the retention of this behavior.

Animals↗

PubMed1990

Maternal behavior of spontaneously hypertensive and Wistar-Kyoto normotensive rats: effects of reciprocal cross-fostering of litters.

Maternal behavior of spontaneously hypertensive (SHR) and Wistar-Kyoto (WKY) normotensive dams rearing either their natural pups or pups of the opposite strain was assessed when pups were 4-6 days of age. Consistent with previous reports, SHR dams caring for their natural pups were observed nursing and licking their pups more often than WKY dams and were away from their pups less often. The behavior of dams rearing cross-fostered pups demonstrated that this strain difference in patterns of maternal behavior is dependent on pup type. Mothers of both strains shifted their behavior in the direction of the opposite strain's maternal behavior pattern when caring for cross-fostered pups. This shift was most pronounced in SHR dams caring for WKY pups. These data suggest that alterations in patterns of maternal behavior produced by cross-fostering may play an important role in cross-fostering effects on the physiological development of SHRs previously reported.

Animals↗

PubMed1986

Effect of varying the duration of pre-test cage habituation on oxytocin induction of short-latency maternal behavior.

Intracerebroventricular (ICV) infusion of the peptide hormone oxytocin has been previously reported to induce the performance of short-latency maternal behavior (less than one hour of exposure to foster pups required) in estrogen-treated, ovariectomized virgin rats. Tests for the effect of ICV oxytocin in maternal behavior latency have included transfer of animals from their home cage to a larger test cage one to two hours before oxytocin infusion. The importance of this test feature on peptide-induced short-latency maternal behavior was evaluated by varying the duration of the pre-test cage habituation. The responses of ovariectomized, estrogen-primed Zivic-Miller Sprague-Dawley rats housed in the test cages one week, two hours, or 0 hours before oxytocin or saline infusion were compared. It was found that only the rats given two hours of pre-test cage habituation responded to ICV oxytocin treatment with short-latency maternal behavior. This result is discussed with regard to the failure of other investigators to elicit short-latency maternal behavior with oxytocin. Possible neuroendocrine mechanisms for the interaction of degree of environment novelty with oxytocin-induced behavior are considered.

Animals↗

PubMed1998

Opiate disruption of maternal behavior: morphine reduces, and naloxone restores, c-fos activity in the medial preoptic area of lactating rats.

Morphine significantly impairs maternal behavior; naloxone, an opiate antagonist, restores it. Maternal behavior is associated with c-fos expression, an immediate early gene product, in the medial preoptic area (mPOA) of females. In two experiments, the effects of morphine-alone and morphine plus naloxone on the expression of c-fos were examined. On postpartum day 5, females were injected with morphine or saline (experiment 1), and morphine + naloxone or morphine + saline (experiment 2) and placed back in the home-cage, separated from their pups by a wire-mesh partition. A separate group in experiment 1 was injected but not exposed to pups. Processing for c-fos immunohistochemistry commenced, and c-fos positive cells in a proscribed portion of mPOA were counted. Morphine-treated females had fewer c-fos cells in mPOA compared to saline-treated females, and the presence of pups accounted for a significant increase in c-fos-expressing neurons, whereas in females not exposed to pups, morphine treatment did not significantly reduce baseline c-fos expression (experiment 1). Furthermore, naloxone mitigated the effect as morphine + naloxone-treated females expressed more c-fos cells compared to morphine + saline females (experiment 2). Morphine-treated females, therefore, may exhibit reductions in maternal behavior because of relative opiate-induced inactivation of areas of the brain devoted to the regulation of maternal behavior.

Analgesics, Opioid↗

PubMed1977

Alterations in the maternal behavior of rats rearing malnourished offspring.

Investigations of maternal behavior of mothers fed a low-protein diet indicated deficits in retrieval and in the rate of nest-building. In addition, they indicated a concomitant increase in time spent with young when assessed during periods not associated with the retrieval/nest-building test session. The adrenalectomized mother, another case that produces growth-stunted progeny, was compared with both low-protein and control mothers for maternal behavior. Unlike the low-protein mother, the adrenalectomized mother did not exhibit retrieval or nest-building deficits; however, the adrenalectomized mother did display an increase in time spent with young. These data suggest that although deficits in retrieval and nest-building can be attributed to the nutritional condition of the mother, the stimulus characteristics of the malnourished pup are important in eliciting the increased time spent with the litter.

Adrenalectomy↗

PubMed1995

Human placental lactogen infusions into the medial preoptic area stimulate maternal behavior in steroid-primed, nulliparous female rats.

The effects of central administration of human placental lactogen (hPL) on the onset of maternal behavior were measured in steroid-primed, adult ovariectomized, nulliparous rats. Rats were fitted with bilateral cannulas directed at the medial preoptic area (MPOA) and gonadectomized 1 week before being implanted sc with progesterone (P)-filled Silastic implants (treatment Day 1). On Day 11 P capsules were removed, and each female was implanted sc with a single estradiol (E2) capsule. On Days 11 to 13 animals were infused bilaterally with 40 ng of hPL/infusion or given 0.4 microliter vehicle. Subjects were given hormone or vehicle infusions five times during this period, twice each on Days 11 and 12 (1000 and 1600 hr) and once on Day 13 (1000 hr). Behavioral testing began on day 12 after the 1000 hr infusions and continued daily for 6 days. All females were injected sc twice daily with bromocriptine (2 mg/ke) to suppress endogenous PRL secretion from Day 11 to the completion of testing. The results showed that central infusions of hPL stimulated a fast onset of maternal behavior relative to controls. Latencies to display specific components as well as complete maternal behavior toward foster young were about 1 day in the hPL-treated group and 4 days in vehicle-infused controls. Infusions of hPL into the MPOA between 7.8 and 8.0 AP resulted in the fastest rate of onset of maternal behavior. These findings demonstrate that central infusion of a heterologous placental lactogen, hPL, is capable of stimulating maternal behavior may normally be brought about by exposure to placental lactogens as well as prolactin.

Animals↗

PubMed2005

A cross-fostering analysis of the effects of PCB 77 on the maternal behavior of rats.

Polychlorinated biphenyls (PCBs) are environmental contaminants known to cause multiple behavioral and developmental problems in humans and animals. In rats, gestational exposure to the PCB congener 3, 4, 3', 4'-tetrachlorobiphenyl (PCB 77) affects the brain and behavior of the offspring as well as the maternal behavior of the dams. Whether the behavior of dams is affected by direct effects of the contaminant or indirectly by actions of the PCB on the developing offspring is not known. We investigated this question using a cross-fostering paradigm in which pregnant rats were exposed to either oil vehicle or 2 mg/kg of PCB 77 on gestational days 6 through 18, and then raised pups that had been exposed to either oil or PCB 77 during gestation. Maternal behavior was observed on postnatal days 1, 2, 4 and 6. Some of the effects on maternal behavior, including an increase in the frequency of nursing bouts and in the amount of maternal autogrooming, can be ascribed to prenatal exposure of the litters to the PCB. Other behavioral effects, including an increase in time on the nest and in the amount of pup grooming as well as a reduction in high-crouch nursing, appear to be due to both direct effects of the PCB on the dams and effects mediated by changes in the offspring. Our results show that exposure to PCB 77 can have complex effects on the behavioral interactions between the dams and their litters with a potential impact on the development of the offspring.

Analysis of Variance↗

PubMed1990

Central prolactin infusions stimulate maternal behavior in steroid-treated, nulliparous female rats.

A series of experiments were conducted to determine whether and under what conditions central prolactin (PRL) administration would stimulate the onset of maternal behavior in female rats and to identify possible neural sites of PRL action. In each experiment ovariectomized, nulliparous rats whose endogenous PRL levels were suppressed with bromocriptine were tested for maternal behavior toward foster young. In experiments 1, 2, and 4, females were also exposed to pregnancy-like levels of progesterone (days 1-11) followed by estradiol (days 11-17). In experiment 1 infusions (days 11-13) of four doses of ovine PRL (400 ng, 2 micrograms, 10 micrograms, or 50 micrograms, but not 80 ng) into the lateral ventricle resulted in a rapid onset of maternal behavior (behavioral testing, days 12-17). The stimulatory action of these doses of PRL appears to be central, since subcutaneous injections of 50 micrograms of ovine PRL failed to affect maternal responsiveness (experiment 2). Experiment 3 indicated that the stimulatory effect of intracerebroventricularly administered PRL is steroid dependent. Infusions of either 10 micrograms of ovine PRL or 10 micrograms of rat PRL failed to induce maternal behavior in nonsteroid-treated animals. In the final experiment (no. 4) bilateral infusions of 40 ng of ovine PRL into the medial preoptic area of steroid-treated rats resulted in a pronounced stimulation of maternal behavior. These findings demonstrate a central site of PRL action in the stimulation of maternal responsiveness and point to the medial preoptic area as a key neural site for PRL regulation of maternal behavior.

Animals↗