Onset Of Maternal Behavior Research Articles

Factors influencing ease of whelping and its relationship with maternal behaviour and puppy perinatal mortality in commercially bred dogs

For dog breeders, parturition is a critical stage in the reproductive cycle of the dam. Evidence in other mammals suggests that a difficult labour can influence maternal behaviour and offspring viability during the first hours postpartum. However, the effect of whelping difficulty on the onset of maternal behaviour has not yet been investigated in domestic dogs. Here we developed an ease of whelping (EoW) index in dams maintained within a Commercial dog Breeding Establishment (CBE) environment and investigated the relationship between intrinsic and extrinsic factors (breed group according to size/weight, litter size, parity, whelping season and origin of the dam), EoW, early maternal behaviour and puppy perinatal mortality. The behaviour of 30 dams was observed throughout the whelping process, starting 24 h before delivery of the first puppy until birth of the last puppy. Parturition duration, birth interval, and behaviours indicative of distress, restlessness, and general activity were scored and included in a Principal Component Analysis to construct the EoW index. Subsequently, mother–pup interactions and puppy perinatal mortality were recorded during the first 24 and 72 h postpartum respectively. Results showed that EoW was significantly affected by whelping season, litter size and origin of the dam (whether she was born and raised within the CBE or brought in). Furthermore, mothers that experienced more difficult parturitions (higher EoW score) spent more time lying in contact with their puppies during the first 24 h postpartum. Time in contact with puppies was also significantly affected by breed group. Nursing duration was significantly affected by breed group and origin of the dam. Additionally, medium-size breed (10–20 kg) puppies were significantly less likely to experience perinatal mortality than large breeds (> 20 kg). These findings are particularly relevant for the welfare of breeding dams maintained in large-scale CBEs where the staff-to-dog ratio might be insufficient to adequately manage multiple simultaneous parturitions.

The Onset of Maternal Behavior in Sheep and Goats: Endocrine, Sensory, Neural, and Experiential Mechanisms.

In sheep and goats, the onset of maternal behavior at parturition is characterized by a first phase called maternal responsiveness during which the mother is attracted to any newborn. In a second phase, called maternal selectivity, the mother establishes a selective bond with her young so that she only accepts it at suckling. After a description of the behavioral expression of both phases, this chapter reviews the physiological, sensory, and neural mechanisms involved. These two behavioral processes are synchronized with parturition by the vaginocervical stimulation induced by the expulsion of the newborn. Olfactory cues provided by the neonate are involved in maternal responsiveness and selectivity. Oxytocin supported by estrogens is the key factor for maternal responsiveness. The neural network involved in maternal responsiveness is mainly hypothalamic and is different from the circuitry involved in selectivity, which mainly concerns olfactory processing regions. Visual and auditory cues are necessary for offspring recognition at a distance. This multisensory recognition suggests that mothers form a mental image of their young. Maternal experience renders mothers more responsive to maternally relevant physiology and to young-related sensory inputs.

Innate and plastic mechanisms for maternal behaviour in auditory cortex.

Infant cries evoke powerful responses in parents1–4. To what extent are parental animals intrinsically sensitive to neonatal vocalizations, or might instead learn about vocal cues for parenting responses? In mice, pup-naive virgins do not recognize the meaning of pup distress calls, but retrieve isolated pups to the nest following cohousing with a mother and litter5–9. Distress calls are variable, requiring co-caring virgins to generalize across calls for reliable retrieval10,11. Here we show that the onset of maternal behavior in mice results from interactions between intrinsic mechanisms and experience-dependent plasticity in auditory cortex. In maternal females, calls with inter-syllable intervals (ISIs) from 75:375 ms elicited pup retrieval, and cortical responses generalized across these ISIs. In contrast, naive virgins were behaviorally sensitive only to the most common (‘prototypical’) ISIs. Inhibitory and excitatory neural responses were initially mismatched in naive cortex, with untuned inhibition and overly-narrow excitation. During cohousing, excitatory responses broadened to represent a wider range of ISIs, while inhibitory tuning sharpened to form a perceptual boundary. We presented synthetic calls during cohousing and observed that neurobehavioral responses adjusted to match these statistics, a process requiring cortical activity and the hypothalamic oxytocin system. Neuroplastic mechanisms therefore build on an intrinsic sensitivity in mouse auditory cortex, enabling rapid plasticity for reliable parenting behavior.

Gene Expression Profiling during Pregnancy in Rat Brain Tissue.

The neurophysiological changes that occur during pregnancy in the female mammal have led to the coining of the phrases “expectant brain” and “maternal brain”. Although much is known of the hormonal changes during pregnancy, alterations in neurotransmitter gene expression have not been well-studied. We examined gene expression in the ventromedial nucleus of the hypothalamus (VMH) during pregnancy based on the fact that this nucleus not only modulates the physiological changes that occur during pregnancy but is also involved in the development of maternal behavior. This study was designed to identify genes that are differentially expressed between mid- and late-pregnancy in order to determine which genes may be associated with the onset and display of maternal behavior and the development of the maternal brain. A commercially available PCR array containing 84 neurotransmitter receptor and regulator genes (RT2 Profiler PCR array) was used. Brains were harvested from rats on days 12 and 21 of gestation, frozen, and micropunched to obtain the VMH. Total RNA was extracted, cDNA prepared, and SYBR Green qPCR was performed. In the VMH, expression of five genes were reduced on day 21 of gestation compared to day 12 (Chrna6, Drd5, Gabrr2, Prokr2, and Ppyr1) whereas Chat, Chrm5, Drd4, Gabra5, Gabrg2, LOC289606, Nmu5r2, and Npy5r expression was elevated. Five genes were chosen to be validated in an additional experiment based on their known involvement in maternal behavior onset. This experiment confirmed that gene expression for both the CCK-A receptor and the GABAAR γ2 receptor increases at the end of pregnancy. In general, these results identify genes possibly involved in the establishment of the maternal brain in rats and indicate possible new genes to be investigated.

Effect of a difficult calving on the vigour of the calf, the onset of maternal behaviour, and some behavioural indicators of pain in the dam

The neonate's development and survival is dependent upon being vigorous at birth and receiving appropriate maternal care. However, difficulty at delivery can result in less vigorous offspring and maternal care can be altered, probably as a consequence of exhaustion, pain and human intervention. The first 3h after expulsion of the calf were observed continuously from videos following twelve natural calvings and sixteen calvings assisted by farm staff (including four malpresentations) from Holstein cows. Calvings were balanced within groups for parity of the dam, genetic group, sex and birth weight of the calf, calving pen and calving season. Assisted calves were less vigorous with higher latencies to attempt to stand, achieve standing, walk and reach the udder than unassisted calves (P<0.05). Furthermore, assisted calves also tended to be less likely to stand and walk within the first 3h after birth (P<0.1), spent more time lying on their flank (P=0.019) and had more frequent bouts of this behaviour (P=0.033). Assisted dams did not take longer to lick the calf and performed as much licking as unassisted dams (P>0.05), indicating no delayed onset or impaired expression of maternal behaviour in dams given assistance at delivery. Study of potential pain-related behaviours revealed that assisted dams spent less time self-grooming (P=0.033) than dams delivering naturally, which could suggest greater pain. However, there were no significant differences in any of the other pain-related behaviours. Our results suggest that, although maternal behaviour was unaffected by a difficult delivery, dairy calves born following difficult calvings have lower vigour in the first 3h after birth than unassisted calves. This might have longer-term effects on the health and survival of the calves.

Oxytocin changes primate paternal tolerance to offspring in food transfer

Oxytocin facilitates social recognition in rats and mice, onset of maternal behavior in virgin mice and formation of pair bonds without copulation in prairie voles. However, the relationship between this peptide and paternal behavior in primates remains largely unknown. We investigated whether oxytocin affects paternal behavior in common marmosets. In these primates, fathers as well as mothers take care of their infants, and transferring food to the infants is one of their more obvious caretaking behaviors. We tested whether oxytocin and an oxytocin receptor antagonist affect the transfer of food to offspirng by fathers. After intracerebroventricular infusion of the vehicle, oxytocin, or the oxytocin receptor antagonist, the fathers' behavior, including picking up food, transferring food to the offspring, and refusing to transfer food to the offspring, was analyzed. Compared with the vehicle, oxytocin reduced the frequency of refusal. This was not caused by reduction of appetite. Although the oxytocin receptor antagonist did not change the frequency of refusal behavior of the fathers statistically significant manner, these observations suggest that the tolerance of the adult male marmoset toward its offspring as shown by the transfer of food is increased by oxytocin administered into the central nervous system.

Maternal behaviour and peripartum levels of oestradiol and progesterone show little difference in Merino ewes selected for calm or nervous temperament under indoor housing conditions

Temperament influences the expression of maternal behaviour in sheep under experimental field conditions. We investigated whether maternal behaviour between ewes selected for a calm or nervous temperament is independent from environmental conditions. In addition, the level of maternal behaviour expressed by mothers is correlated with the concentration of hormones during the peripartum period. Therefore, we investigated whether the selection for temperament had resulted in hormonal differences between the two lines with regard to the hormones that could be involved in the onset of maternal behaviour. Oestradiol, progesterone and cortisol concentrations from 4 days before parturition to 24 h after parturition were determined from blood samples collected from 10 calm and 12 nervous ewes. Behavioural interactions between ewe and lamb were also recorded for 2 h starting at parturition. Mothers of both temperament lines showed adequate maternal behaviour under the controlled conditions of the study. Therefore, the results of the study do not suggest that selection for a calm or nervous temperament has profoundly affected the intrinsic ability of mothers to display adequate maternal behaviour. Hormonal differences between the two temperament lines were generally small and their possible influence on the display of maternal behaviour in the two temperament lines would have to be demonstrated.

Normal maternal behavior, but increased pup mortality, in conditional oxytocin receptor knockout females.

Oxytocin (Oxt) and the Oxt receptor (Oxtr) are implicated in the onset of maternal behavior in a variety of species. Recently, we developed two Oxtr knockout lines: a total body knockout (Oxtr-/-) and a conditional Oxtr knockout (OxtrFB/FB) in which the Oxtr is lacking only in regions of the forebrain, allowing knockout females to potentially nurse and care for their biological offspring. In the current study, we assessed maternal behavior of postpartum OxtrFB/FB females toward their own pups and maternal behavior of virgin Oxtr-/- females toward foster pups and compared knockouts of both lines to wildtype (Oxtr+/+) littermates. We found that both Oxtr-/- and OxtrFB/FB females appear to have largely normal maternal behaviors. However, with first litters, approximately 40% of the OxtrFB/FB knockout dams experienced high pup mortality, compared to fewer than 10% of the Oxtr+/+ dams. We then went on to test whether or not this phenotype occurred in subsequent litters or when the dams were exposed to an environmental disturbance. We found that regardless of the degree of external disturbance, OxtrFB/FB females lost more pups on their first and second litters compared to wildtype females. Possible reasons for higher pup mortality in OxtrFB/FB females are discussed.

Dopamine D 1 receptor activation of adenylyl cyclase, not phospholipase C, in the nucleus accumbens promotes maternal behavior onset in rats

A body of evidence supports the idea that the mesolimbic dopamine (DA) system modulates the natural increase in responsiveness female rats show toward offspring (biological or foster) at birth. In the absence of the full hormonal changes associated with pregnancy and birth, female rats do not show immediate responsiveness toward foster offspring. Activation of the mesolimbic DA system can produce an immediate onset of maternal behavior in these females. For example, female rats that are hysterectomized and ovariectomized on day 15 of pregnancy (15HO) and presented with pups 48 hours later normally show maternal behavior after 2–3 days of pup exposure, but will show maternal behavior on day 0 of testing after microinjection of the DA D 1 receptor agonist, SKF 38393, into the nucleus accumbens (NA) at the time of pup presentation. DA D 1 receptor stimulation is known to activate cAMP intracellular signaling cascades via its stimulation of adenylyl cyclase (AC). However, some DA D 1 receptors are also linked to phospholipase C (PLC) and are capable of activating phosphatidylinositol signaling cascades. SKF 38393 stimulates both types of D 1 receptors. Here we provide evidence that the facilitatory effects of DA D 1 receptor stimulation in the NA on maternal behavior are mediated by AC-linked DA D 1 receptors. By examining the effects of intra-NA application of SKF 83822, a drug which selectively binds DA D 1–AC receptors, or SKF 83959, a drug which selectively activates D 1–PLC-linked receptors, we find that only SKF 83822 facilitates maternal behavior onset.

Steroid-induced alterations in mRNA expression of the long form of the prolactin receptor in the medial preoptic area of female rats: Effects of exposure to a pregnancy-like regimen of progesterone and estradiol

It is firmly established that the onset of maternal behavior in the female rat is stimulated by a combination of hormones that include prolactin (PRL), estradiol (E 2), and progesterone (P 4). Specifically, nulliparous rats display short latencies to respond to foster young when primed with Silastic capsules filled with P 4 and E 2 and then administered PRL centrally to the medial preoptic area (MPOA), an area integrally involved in the expression of maternal behavior in this species. PRL or P 4 treatments alone are ineffective in stimulating the expression of maternal care. Since the actions of PRL in the MPOA appear to be mediated by PRL receptors, it was of interest to determine whether and how treatment with P 4 and E 2 together or separately might alter mRNA expression of the long form of the PRL receptor (PRL-R L) in the MPOA. Using in situ hybridization histochemistry (ISHH), mRNA expression of the PRL-R L was measured in the MPOA of ovariectomized, nulliparous rats treated with various combinations of P 4 and E 2. Treatment of animals with P 4 alone for 10 days or with P 4 followed by E 2 for 1 or 4 days resulted in reductions in PRL receptor mRNA expression in the MPOA when compared with the expression in animals treated with E 2 alone or blank capsules. The actions of P 4 on mRNA expression of the PRL-R L were more pronounced in the dorsal MPOA. Circulating PRL levels collected at the time of sacrifice were elevated in all groups treated with E 2, but no association between PRL levels and receptor mRNA expression within the MPOA was evident. These findings indicate that the dorsal MPOA may be one site of progesterone's action in facilitating prolactin-mediated maternal behavior.

Prolactin receptor gene expression in the forebrain of pregnant and lactating rats

Prolactin plays a large role in the onset of maternal behavior at parturition. Knowledge of the change in expression of the prolactin receptor in the brain across pregnancy and lactation, however, is limited. Prolactin receptor gene expression was determined by in situ hybridization histochemistry during pregnancy and lactation in rats. Expression of the mRNA for the longform of the prolactin receptor (PRL-R-L) was measured in various forebrain structures in primigravid rats at different stages of pregnancy, in primiparous rats during early, mid-, and late lactation, and in age-matched, nulliparous females in diestrus. Hybridizations were performed using a [ 33P]-labeled riboprobe specific for the long form of the prolactin receptor mRNA complimentary to 290 bp of the prolactin receptor gene. The following areas of the forebrain were examined: medial preoptic area (MPOA), median preoptic nucleus both dorsal (MePOd) and ventral (MePOv) to the anterior commissure, ventral lateral septum (LSv), and the ventral and principal parts of the bed nucleus of the stria terminalis (BnSTv and BnSTpr, respectively). Overall, the number of cells expressing PRL-R-L mRNA was significantly higher at 2 h postpartum compared to diestrus in all areas examined except the LSv. In addition, there were lower numbers of PRL-R-L cells during all stages of lactation compared to pregnancy. The number of grains per cell in the MPOA and LSv did not change as dramatically as the number of cells expressing PRL-R-L mRNA in those brain regions. These data contribute to the growing body of evidence that the neural lactogenic system changes as a function of female reproductive state. Changes in PRL-R-L mRNA in terms of behavior and endocrine functions are discussed.

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.

Chapter 18 Lactogenic hormone regulation of maternal behavior

Biological factors can profoundly affect a mother's response to her young. For example, it is well known that the hormones of pregnancy act on the maternal brain to stimulate the spontaneous onset of maternal behavior at parturition. Studies in the rat have provided an excellent model to investigate maternal behavior in mammals, since maternal behavior in rats is easily observable and readily quantifiable and it is well-documented that the endocrine state of gestation helps to bring about the onset of maternal behavior around the time of birth. The same response in virgin animals requires a number of days of constant exposure to pups before maternal-like behaviors emerge. To date, research has established that the steroid hormones, estradiol and progesterone, and the lactogenic hormones, prolactin and the placental lactogens, act in concert to stimulate maternal behavior in the pregnant female. Treatment of adult, virgin rats with these hormones can stimulate a rapid onset of maternal care. In the present chapter experiments are described that demonstrate key roles for prolactin and placental lactogens in the onset of maternal behavior. Central sites of action of prolactin and placental lactogens, including the medial preoptic area, appear to be involved in stimulating the onset of maternal care. Other studies are discussed which support the involvement of the prolactin receptor in the endocrine regulation of maternal behavior using prolactin receptor antagonist and 'knock-out' models in rats and mice, respectively. Overall, these studies indicate that during pregnancy the endocrine system primes the mother's brain so that the new mother displays appropriate and successful behaviors toward her newborn at parturition.

The effect of opioid antagonism and environmental restriction on plasma oxytocin and vasopressin concentrations in parturient gilts.

Oxytocin plays an important role at parturition due to its involvement in uterine contractions, foetal expulsion and the onset of maternal behaviour. The role of the related neurohypophysial hormone, vasopressin, is less clear; however, there is some evidence that it is also involved in maternal behaviour and its role in osmotic regulation is well established. The aim of this study was to investigate the inhibitory effects of endogenous opioids on these hormones during the expulsive phase of parturition in the pig, and to examine how opioid restraint interacts with environmental restriction. The subjects of this study were 31 Large Whitex Landrace primiparous sows (gilts). An indwelling jugular catheter was implanted under general anaesthesia at 12 days before the expected parturition day (EPD). From 5 days before the EPD 15 of the gilts were individually housed in a restrictive parturition crate without straw and 16 were individually housed in a straw-bedded pen. Blood samples were taken with increasing frequency towards and during parturition through a catheter extension to reduce disturbance. At 7.5 min after the birth of the first piglet half of the gilts in each environment received a dose of the opioid receptor antagonist naloxone (1 mg/kg, i.v.) with the remaining gilts receiving saline as a control. Overall, there was no effect of environment on either circulating oxytocin or vasopressin. However, both oxytocin and vasopressin were inhibited by endogenous opioids during the expulsive phase. The inhibitory effects of opioids on these hormones did not appear to have any adverse effects on the progress of parturition as judged by cumulative piglet birth intervals. The regulation of the opioid inhibition of oxytocin and vasopressin during parturition is discussed in relation to other neurotransmitters and whether opioid inhibition of these neurohypophysial hormones is part of the 'normal' physiological response to parturition or whether it is stress-induced.

Using c-Fos immunocytochemistry to identify forebrain regions that may inhibit maternal behavior in rats.

Evidence indicates there is a neural system that inhibits maternal behavior in virgin rats. It has been suggested that pregnancy hormones promote the onset of maternal behavior by reducing the behavioral influence of this system. The authors used c-Fos immunocytochemistry to identify brain regions more activated by pup exposure in nonmaternal rats than in maternal rats. Previous experiments indicated that some of these regions, such as the posterodorsal medial amygdala and several medial hypothalamic sites, inhibit maternal behavior. For others, such as the ventral lateral septum, dorsal premammillary nucleus, and principal bed nucleus of the stria terminalis, this is the first indication that they could also inhibit maternal responding. These regions have previously been implicated in promoting defensive behaviors, consistent with the finding that nonmaternal rats actively avoid pups. These findings suggest the existence of a neural circuit through which pup exposure could promote defensive responses in virgin rats, and how pregnancy hormones could reduce such activity to stimulate maternal behavior.

Elevated Fos-like immunoreactivity in the brains of postpartum female prairie voles, Microtus ochrogaster.

This study investigated the expression of the protein product of the immediate early gene c-fos in the brains of female prairie voles (Microtus ochrogaster) in association with pregnancy and postparturient activities including maternal behavior, lactation and postpartum estrus. Fos expression was assessed in female voles that were late in pregnancy, nonpregnant or at one of three different times postpartum (0-8, 12-24, and 24-48 h, respectively). A significant increase in the number of cells displaying Fos immunoreactivity (Fos-ir) was observed during the 0-8 h and 12-24 h postpartum time periods in the accessory olfactory bulbs, medial preoptic area, hypothalamus (specifically, the supraoptic nucleus, ventro-medial hypothalamus, and paraventricular nucleus), lateral septum, bed nucleus of the stria terminalis, and primary somatosensory area of the brain. The number of Fos-ir cells decreased after 24 h postpartum. There were no significant changes in Fos-ir cell numbers in the primary olfactory bulbs, hippocampus, or caudate putamen. The neural activation of the medial preoptic area, accessory olfactory bulbs, hypothalamus, and bed nucleus is consistent with reports in rats of Fos induction associated with the onset of maternal behavior. In voles postpartum estrous behavior begins and ends 0-12 h after parturition. Maternal behavior, including lactation, is initiated at the same time but persists for several weeks. The highest Fos-ir cell numbers reported here coincide with the timing of postpartum estrous behavior in this species.

Physiological Correlates of Maternal–Offspring Behaviour in Sheep: A Factor Analysis

The onset of maternal behaviour in sheep is controlled by levels of oestrogen and progesterone in gestation and the release of oxytocin during delivery. Factor analysis was used to investigate the relationships between maternal behaviour, offspring behaviour, and maternal ovarian hormone levels during gestation in sheep. Ewes gave birth to lambs following embryo transfer between two breeds (Suffolk and Scottish Blackface), which differ in their expression of maternal behaviour. Plasma oestradiol-17β concentration was significantly higher in Blackface ewes in the last 6 weeks of pregnancy, whereas plasma progesterone was higher in Suffolk ewes in early and mid gestation. Factor analysis revealed three factors that accounted for 50% of the total variation between variables. Factor 1 described lamb activity and had positive loadings for lamb behavioural latencies immediately postpartum, and negative loadings for sucking frequency. Factor 2 described some aspects of ewe maternal behaviour, and had positive loadings for ewe–lamb separation and lamb vocalisation, and negative loadings for ewe grooming behaviour and plasma oestradiol concentration. Factor 3 described ewe and lamb-sucking interactions, and had positive loadings for ewe moving as the lamb made sucking attempts, and negative loadings for ewe and lamb vocalisation. Factor 1 scores were significantly affected by lamb breed: Suffolk lambs received positive scores, indicating a longer latency to right and stand, whereas Blackface lambs had negative scores. Maternal progesterone concentration was significantly correlated with Factor 1 scores. Factor 2 and 3 scores were significantly affected by ewe breed. This study has shown that there are two behavioural axes to maternal behaviour in sheep, and that maternal oestradiol concentration is related to affiliative behaviours (e.g., grooming) but only weakly to ewe responsiveness to lamb-sucking attempts.

Intact neurons of the lateral habenular nucleus are necessary for the nonhormonal, pup-mediated display of maternal behavior in sensitized virgin female rats.

Our research has demonstrated that the lateral habenular nucleus (Lhb) is necessary for the hormonal onset but not the postpartum maintenance of maternal behavior in the rat (K. P. Corodimas, J. S. Rosenblatt, & J. I. Morrell, 1992; K. P. Corodimas, J. S. Rosenblatt, M. E. Canfield, & J. I. Morell, 1993; T. Matthews-Felton, K. P. Corodimas, J. S. Rosenblatt, & J. I. Morell, 1995). To test the role of the Lhb in the nonhormonal onset of maternal behavior, we used the sensitization model in which the continual exposure of females to pups induces maternal behavior. Ovariectomized females received bilateral cytotoxic lesions of neurons of either the Lhb or the dorsal medial cingulate cortex-hippocampus, or they were unoperated. Maternal behavior, activity, and oromotor carrying capability were tested. Complete lesions of the neurons of the Lhb induced significant deficits in pup retrieval and nest building. Sniffing, licking, and crouching behaviors were unaltered. Activity and carrying ability were normal. These results indicate a role for the Lhb that extends to the nonhormonally dependent onset of maternal behavior, but they also indicate a more limited role than in the mediation of the hormonal onset of the behavior.

The duration of estradiol and progesterone exposure prior to progesterone withdrawal regulates oxytocin mrna levels in the paraventricular nucleus of the rat

The nonapeptide oxytocin (OT) is important for milk ejection during lactation, uterine contractility at parturition, and the onset of maternal behavior. Sequential exposure to estradiol (E2) and progesterone (P) followed by P withdrawal increases OT mRNA in the paraventricular nucleus (PVN), and to a lesser degree the supraoptic nucleus (SON), of the rat 48 hours after the P is removed. Although increases in PVN OT mRNA are not accompanied by changes in posterior pituitary OT peptide content, the PVN contains OT neurons that project to both the posterior pituitary (magnocellular group) and extra pituitary sites (parvocellular groups). Steroid-induced increases in OT mRNA occur in both the magnocellular and the parvocellular regions of the PVN. The latter are believed to contribute to CNS release of OT which may be important for certain behaviors including the onset of maternal behavior. The same steroid sequence that increases PVN OT mRNA also induces maternal behavior in virgin ovariectomized rats. Exposure of animals to E2 and P for 2 weeks resulted in the shortest latency to the onset of maternal behavior in ovariectomized rats, whereas exposure for 6 days was associated with a longer latency. In this study we questioned if the duration of E2 and P exposure prior to P withdrawal is an important regulator of PVN OT mRNA levels. We compared OT mRNA levels in the PVN of virgin ovariectomized rats administered no steroid or sequential E2 and P for 2 weeks versus 6 days. On day 1 animals received steroid-filled or empty capsules followed by P-filled or empty capsules on day 3. In one steroid-treated group, E2 and P were continued for 6 days and in the other group for 14 days prior to P removal. Animals were sacrificed 48 hours after P removal. Levels of OT mRNA were compared among 6 day and 2 week steroid-treated animals and sham-treated animals. The relative abundance of OT mRNA was significantly increased, P < 0.05, in animals receiving the 2-week, but not the 6-day, steroid treatment compared to sham-treated animals. Pituitary OT peptide content was not significantly different among the three groups. We conclude that the duration of steroid exposure may be an important regulator of the level of OT mRNA in the PVN of the rat.

Effects of short- and long-term withdrawal from gestational cocaine treatment on maternal behavior and aggression in Sprague-Dawley rats.

Pregnant rats were treated with 30 mg/kg per day cocaine or normal saline either throughout gestation (GD 1-20, cocaine and saline withdrawal) or throughout the gestation and continuing into lactation for 10 days postpartum (cocaine and saline nonwithdrawal). All cocaine-treated dams exhibited more disruptions in the onset of maternal behavior (retrieval, licking, crouching) and were more aggressive (threats and attacks) towards an intruder on postpartum day 6 than saline-treated dams. There were no significant differences in these behaviors between withdrawn and nonwithdrawn cocaine-treated dams. These findings indicate that changes in maternal behavior following chronic moderate cocaine treatment are not simply the result of withdrawal from cocaine treatment following gestation and that other possible mechanisms should be examined.