Enhanced maternal aggression and associated changes in neuropeptide gene expression in multiparous rats.

Although it has often been speculated that prior reproductive experience improves subsequent maternal care, few studies have examined specific changes in behavior during a 1st versus 2nd lactation. During lactation, mothers display heightened aggression toward male intruders, purportedly to protect vulnerable young. In the current study, maternal aggression was examined in primiparous and age-matched multiparous females on postpartum days 5 (PPD5) and PPD15. Expression of oxytocin, oxytocin receptor, arginine vasopressin, arginine vasopressin V1a receptors, and corticotrophin-releasing hormone mRNA was measured following aggression testing at both time points using real-time quantitative PCR in brain regions previously implicated in the regulation of maternal aggression. Multiparity significantly enhanced maternal aggression on PPD5 but not on PPD15. In addition, this increased aggression was associated with region- and gene-specific changes in mRNA expression. These findings indicate that reproductive experience enhances maternal aggression, an effect that may be mediated by region-specific alterations in neuropeptidergic activity. The adaptations observed in multiparous females provide an innate model for the study of neuroplasticity in the regulation of aggression. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Resource partitioning during reproduction in the Norway rat

Rat pups nursed by pregnant dams grow as fast as pups reared by dams that are not pregnant. Moreover, litters that were in utero during a lactation are as numerous at birth and grow as fast as pups developing in a nonlactating, pregnant mother. These litters continue to grow as fast as pups born to nonlactating dams whether or not the first litter remains after the birth of the second litter. When pregnant and lactating dams have a restricted food supply, some dams are capable of extending the duration of their pregnancies by over 2 weeks past that of nonlactating, pregnant dams. This facultative prolongation of pregnancy apparently allows females to carry normal litters to term.     

Plasticity in the maternal circuit: Effects of experience and partum condition on brain astroctye number in female rats.

Female rats that have received a maternal experience undergo enhanced c-fos expression in a number of brain sites when reexposed to pups. The present 2 studies examined changes in the expression of another brain protein, glial fibrillary acidic protein (GFAP), which is a major unit of the astrocytic cytoskeleton. In both experiments, primiparous and multiparous female rats were given varying amounts of postpartum contact with pups and overdosed after varying intervals, with no pups. Brains were prepared for GFAP immunohistochemical analysis. In both studies, Day 5 postpartum multiparous subjects given additional postpartum contact with pups, when compared with pup-exposed primiparous subjects, were found to have significantly higher numbers of GFAP positive cells in the medial preoptic area of the hypothalamus, an area critical for the expression of maternal behavior, but not in control sites. In Experiment 2, an opposite effect of parity was found in the medial amygdala and habenula. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The biparental care system of the California mouse, Peromyscus californicus..

Most knowledge of parent–offspring relations in mammals is derived from studies of mother–infant interactions. Male parental care has been less well studied. We explored maternal and paternal behavior of the California mouse, Peromyscus californicus. Six pairs of parents and their young were videotaped continuously for 12 hours/day, on alternate days from Days 1 to 31 postpartum. Males exhibit all parental activities and to the same extent as displayed by mothers, except lactation. Male parental behavior begins on the day of birth. Mothers and fathers spend substantial and equivalent amounts of time in the nest and in physical contact with pups throughout lactation. Males devote more time than females to licking pups, although females engage in more pup anogenital licking. Mothers nurse for at least 4 weeks, and fathers and mothers both build nests and carry young. The biparental care system of Peromyscus californicus affords an opportunity to develop a broader, more complete view of parent–offspring relations. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Endocrine mechanisms underlying reproductive toxicity in the developing rat chronically exposed to dietary lead

A dose-response study was conducted in a rat model to examine the effects of lifetime lead exposure on the development of the reproductive system and the endocrine mechanisms underlying these effects. Time-impregnated female Sprague-Dawley rats (n = 10-15/group) were exposed to lead acetate in the drinking water at levels of 0.05%, 0. 15%, or 0.45% (w/v) initiated on gestational day 5. At birth, litters were culled to four male and four female pups. Exposure of dams to lead was continued until weaning, following which, the pups continued to be exposed to lead acetate in drinking water until sacrifice. One male and one female pup from each litter were sacrificed at age 21, 35, 55, and 85 d. A significant dose-responsive decrease in birth weight and crown-to-rump length was observed in all lead-exposed litters. However, no marked effects were observed on anogenital distance/crown-to-rump length ratios. Lead exposure resulted in a delay in sexual maturity as measured by prostate weight in male pups and time of vaginal opening in female pups, which increased with lead dose. These disruptions in reproductive physiology were accompanied by a significant decrease in neonatal sex steroid levels and suppression of the plasma concentrations of testosterone (male) and estradiol (female) during puberty. In male pups, this was accompanied by a significant decrease in plasma luteinizing hormone (LH), elevated pituitary LH content, and a decrease in plasma testosterone/LH ratios at the highest dose. In female pups, although no effects were observed on plasma LH concentration, a similar significant elevation in pituitary LH content was observed during early puberty. Postpuberty, plasma LH and sex steroid concentrations were unaffected at any dose in spite of continued lead exposure. No significant effects were observed on epididymal sperm count in male pups at 85 d of age. In female pups, estrus cycling was only significantly disrupted at the highest lead dose. These data suggest that the reproductive axis is particularly sensitive to lead during specific developmental periods, resulting in delayed sexual maturation produced by suppression by sex steroid biosynthesis. The mechanisms underlying this appear to involve lead actions on both LH release and gonadal function. At low, environmentally relevant blood lead concentrations, adaptation to the continuous presence of the metal ion occurs and surprisingly little effect is observed on adult reproductive endocrinology and physiology.     

Maternal behavior of rats is affected by hormonal condition of pups.

Previous research has found that maternal rats discriminate male from female offspring and provide more anogenital licking to males. In the present study, with 80 primiparous Long-Evans rats, 2 experiments determined whether this discrimination is based on hormonal condition of young. It was found that 500 μg of testosterone, estradiol, or dihydrotestosterone injected on the day of birth into female pups led to their receiving an equivalent amount of maternal anogenital licking as males and significantly more than either oil or untreated females. The different steroids had similar effects. Maternal nonanogenital licking was not affected by sex or hormonal condition of pups, but it was significantly increased by injection per se. Effects on maternal behavior of hormonal condition or neonatal injection of young were apparent 1 and 9 days after injection. (13 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Women's and men's personal goals during the transition to parenthood.

To investigate how women's and men's personal goals change during the transition to parenthood, the authors studied 348 women (152 primiparous and 196 multiparous) and 277 of their partners at 3 times: early in pregnancy, 1 month before the birth, and 3 months afterward. At each measurement, participants completed the Personal Project Analysis questionnaire (B. R. Little, 1983). The results showed that during pregnancy women became more interested in goals related to childbirth, the child's health, and motherhood and less interested in achievement-related goals. After the birth women were more interested in family and health-related issues. These changes were more substantial among the primiparous than among the multiparous mothers. Although the men's personal goals changed during the transition to parenthood, these changes were less substantial than those found among the women. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Mother's milk and adult presence: Two factors determining initial dietary selection by weanling rats.

Reports 3 experiments with (a) 10 female hooded rats and their litters (reduced to 6 pups/litter); (b) 1 male and 4 female wild rats (Rattus norvegicus) and 2 litters (6 pups/litter) born to colony members; and (c) 6 litters of wild and 4 litters of hooded rat pups. Results indicate that adult Ss were capable of influencing their pups' choice of diet during weaning in 2 ways: (a) cues transmitted from a mother to her pups during the nursing period were sufficient to determine the dietary preference of the young at weaning, and (b) the presence of adult Ss at a feeding site was sufficient to determine the choice of feeding site by the pups independent of cues transmitted during the nursing period. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Digestion and energy balance in lactating dairy cows fed varying ratios of alfalfa silage and grain

Forty-four multiparous and 43 primiparous Holstein cows were used to study the effect of dietary alfalfa silage: grain ratio on digestion, passage of digesta, and energy balance in a complete lactation experiment. Cows were placed on one of five treatments with forage contents from 38.2 to 98.2% (DM basis) during the first 12 wk of lactation. Forage content was increased during wk 13 to 26 of lactation to give diets containing 48.2 to 98.2% forage and again during wk 27 to 44 to give diets containing 68.2 to 98.2% forage. Both DMI and DM digestibilities decreased as proportion of dietary forage increased. The DM digestibilities for multiparous and primiparous cows ranged between 69.3 and 57.3% during early lactation and 64.9 and 55.5% during late lactation. Increased percentage of forage in the diet decreased and then increased (quadratic relationship) the ruminal retention time of La, a marker applied to the alfalfa silage in early and late lactation. Time cows spent eating and ruminating per kilogram of DMI increased as proportion of forage in the diet increased. Cows fed diets with a high proportion of alfalfa silage remained in negative energy balance longer than cows fed high grain diets. Intake of NEL (calculated by either of two methods) minus NEL output (milk, maintenance, and BW change) resulted in net balances of NEL after 36 wk of lactation within 5% of NEL intake and indicated that estimates of the NEL value of feedstuffs used in this experiment were reasonably accurate.     

Effect of crude fat and crude protein on reproduction and weaning growth in four strains of inbred mice

Diets made from natural ingredients were fed to four inbred strains of mice (BALB/cAnN, C3H/HeN, C57BL/6N and DBA/2N) to study the effects of different concentrations of dietary crude protein, 18% and 24% with crude fat concentrations of 4%, 8%, and 12% on reproduction and weanling growth. The parameters measured included the number of litters and pups born, the number of litters and pups weaned, weanling mortality and weanling weight. Neither crude protein nor crude fat concentrations had significant effects on any of the reproductive parameters tested. However, a significant fat x protein interaction was observed for reproduction. These results indicate that the absolute concentrations of crude protein and crude fat in diets for inbred mouse production are not as important as the ratio of these two nutrients. There was also a differential response in reproduction among the strains due to the level of fat which indicates different dietary requirements for fat among these four inbred strains for maximum reproduction. In contrast to the results for reproduction, crude protein and crude fat independently had significant effects on weanling weight, but there was no effect due to the ratio of the nutrients. There was a significant increase in the weanling growth rate for all four strains as the dietary fat level increased, but a decrease in growth rate as the protein level increased. The reduced growth rate due to the increased protein was not of the same magnitude for all four strains which indicates specific protein requirements among the strains for weanling growth.     

Early weaning in the rat: Effects on whole-brain RNA and free amino acids.

Each of 67 CAW:CFE(SD)spf rat litters was reared with its mother or with its mother and a virgin female ("aunt"). At pup age 15 days, some litters were deprived of their mothers, some remained with mothers whose nipples were cauterized to prevent suckling, and some remained intact until 21 days when all pups were housed individually or in peer groups until tested at 56 days. While increases in whole-brain free aspartic acid were attributed to loss of opportunity for suckling since they occurred in pups without mothers or with cauterized mothers, decreases in brain RNA caused by deprivation were prevented by the presence of mother or aunt. Rearing with aunts increased brain weight and emotional reactivity. (27 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Problems of undernutrition research with breeding mice

Rats undernourished throughout pregnancy and lactation lose few young during the suckling period and wean normal size litters of extremely growth-retarded offspring. Comparable treatment of maternal mice, on the other hand, resulted in considerable nestling mortality. Thus few young were reared to weaning and these were not nearly as growth-retarded. Detailed evidence is given of the effects of different schedules of undernutrition throughout pregnancy and lactation on the reproductive performance of mice. These findings are compared with those for rats and possible reasons for the difference in nestling mortality discussed.     

Complementary/alternative medicine for asthma: we do not know what we need to know

In a recent report, a series of studies is described showing that individual differences in rat maternal licking and grooming are correlated with their offsprings' later adrenal response to a stressor (Liu et al., 1997). Pups that received more maternal stimulation in infancy had lower ACTH, corticosterone, and CRH mRNA, while they had greater amounts of GR mRNA. Liu et al. also compared maternal behavior in litters where rat pups were handled daily to maternal behavior of nonhandled litters. They found that mothers of handled pups licked and groomed their young significantly more often than did mothers of control litters. In their discussion, Liu et al. proposed that their data support Levine's (1975) thesis that handling effects in infancy are mediated via the mother. Levine had proposed that handling of pups will modify the mother--pup interaction, thereby causing a change in maternal behavior. The purpose of this commentary is to summarize data obtained with mice and rabbits which, in part, support and, in part, challenge the conclusions of Liu et al.     

Lipoprotein lipase activity and its relationship to high milk fat transfer during lactation in grey seals

Lipoprotein lipase regulates the hydrolysis of circulating triglyceride and the uptake of fatty acids by most tissues, including the mammary gland and adipose tissue. Thus, lipoprotein lipase is critical for the uptake and secretion of the long-chain fatty acids in milk and for the assimilation of a high-fat milk diet by suckling young. In the lactating female, lipoprotein lipase appears to be regulated such that levels in adipose tissue are almost completely depressed while those in the mammary gland are high. Thus, circulating fatty acids are directed to the mammary gland for milk fat production. Phocid seals serve as excellent models in the study of lipoprotein lipase and fat transfer during lactation because mothers may fast completely while secreting large quantities of high fat milks and pups deposit large amounts of fat as blubber. We measured pup body composition and milk fat intake by isotope (deuterium oxide) dilution and plasma post-heparin lipoprotein lipase activity in six grey seal (Halichoerus grypus) mother-pup pairs at birth and again late in the 16-day lactation period. Maternal post-heparin lipoprotein lipase activity increased by an average of four-fold by late lactation (P = 0.027), which paralleled an increase in milk fat concentration (from 38 to 56%; P = 0.043). Increasing lipoprotein lipase activity was correlated with increasing milk fat output (1.3-2.1 kg fat per day) over lactation (P = 0.019). Maternal plasma triglyceride (during fasting) was inversely correlated to lipoprotein lipase activity (P = 0.027) and may be associated with the direct incorporation of long-chain fatty acids from blubber into milk. In pups, post-heparin lipoprotein lipase activity was already high at birth and increased as total body fat content (P = 0.028) and the ratio of body fat: protein increased (P = 0.036) during lactation. Although pup plasma triglyceride increased with increasing daily milk fat intake (P = 0.023), pups effectively cleared lipid from the circulation and deposited 70% of milk fat consumed throughout lactation. Lipoprotein lipase may play an important role in the mechanisms involved with the extraordinary rates of fat transfer in phocid seals.     

Reproductive experience and activation of maternal memory.

The maternal and neurobiological responses of biological mothers and pup-induced maternal virgin rats were compared 55 and 80 days after an initial 2-day maternal experience. When tested for home cage responsiveness after prolonged isolation from young, the biological, primiparous rats displayed shorter maternal latencies. Primiparous females tested in the presence of pups on the elevated plus-maze displayed increased exploration of the open arms and increased c-Fos expression in the cortical nucleus of the amygdala. Pup exposure and parity also enhanced activation of the nucleus accumbens shell and medial nucleus of the amygdala, respectively. Therefore, although both nulliparous and primiparous rats retain a maternal memory for a prolonged time, the memory and neurochemical response appear stronger in primiparous mothers. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Influence of litter size and breed on variation in length of gestation in the dog

The variation in the length of gestation, the period from mating until parturition, was studied in 77 dogs of different breeds; the time for mating was determined by measuring peripheral blood progesterone levels. The mean length of gestation was 62.1 +/- 0.2 (S.E.M.) days, with a variation of 11 days. The number of pups appeared to influence the length of gestation. Length of gestation was negatively correlated (r = -0.96, P < 0.001, n = 44) with litter size in litters with 7 or fewer pups. The intra-breed variation in length of gestation in the five breeds represented by five or more bitches was 3-6 days. The mean gestation of Alsatians (60.1 +/- 0.5, n = 9) was shorter (P < 0.005) than that of the other breeds combined (62.3 +/- 0.3, n = 68). The primiparous/multiparous status of the bitch did not influence the length of gestation.     

Maternal entrainment of the developing circadian system in the Siberian hamster (Phodopus sungorus)

The aim of these studies was to investigate maternal entrainment of developing circadian locomotor activity rhythms in the Siberian hamster. In Experiment 1, mothers were transferred from a 16:8 LD cycle into constant dim red light (DD) from the day of parturition, and wheel-running activity of the mother and pups was individually monitored from the time of weaning. The phases of the individual pups' rhythms were found to be synchronized both to the phase of the mother and to the phase of lights off (ZT 12) of the photo cycle that the mother was exposed to until the day of parturition. To investigate whether this synchrony might reflect direct effects of light acting upon the fetal circadian system in late gestation, the experiment was repeated but with mothers placed into DD early in pregnancy (< or = day 7 of gestation). The results were similar to the first study, suggesting that the mother rather than the photo cycle during the latter part of gestation entrains the developing circadian system. The third experiment investigated whether this entrainment occurred during the postnatal period. Breeding pairs were maintained on alternative light-dark cycles, LD and DL, that were 12 h out of phase. Litters born to mothers on one light-dark cycle were exchanged on the day of birth with foster mothers from the reversed light-dark cycle, then raised in DD. Control litters exchanged between mothers from the same light-dark cycle had similar litter synchrony as shown by nonfostered litters of Experiment 1. However, pups cross-fostered with mothers on reversed LD cycles showed a very different distribution of pup phases. Pups were not synchronized to their natural mother but to their foster mother. Moreover, pups were more scattered over the 24-h period and were found to be significantly synchronized to the phase of the reversed LD cycle. These results demonstrate the occurrence of postnatal entrainment in the Siberian hamster. The increased scatter produced by the cross-fostering paradigm results from some litters being completely entrained to the phase of the foster mother, some with an intermediate distribution between the phase of the natural and foster mothers, and a minority being associated with the phase of the natural mother. These results suggest that Siberian hamster pups are initially synchronized either prenatally or at birth but that the mother continues to provide entrainment signals during the postnatal period.     

The reproductive stage and experience of sexually receptive mothers alter their preference for pups or males.

Female rats show postpartum estrus, a unique stage in their reproductive cycle in which they are able to display maternal and sexual responses at the same time. To assess the relative value of pups or males for sexually receptive mothers with different hormonal profiles and reproductive experiences, we employed a 3-point star maze with 3 choice compartments containing: pups, a sexually active male, or no stimulus (neutral). Cycling maternal and nonmaternal females in late proestrus, independently of their previous reproductive experience, strongly preferred the male to the pups, although most postpartum estrous dams did not exhibit preference for the male. The majority of the postpartum primiparous females did not prefer the litter's chamber either, but a previous reproductive experience strongly determined their preference for the pups. These results suggest that the hormonal changes of the proestrus, in contrast to those of the postpartum estrus, promote a strong preference for the male that is not diminished by the maternal condition. Conversely, the endocrine changes of the postpartum facilitate the effect of previous reproductive experience in strengthening the incentive value of the pups. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Energy intake and utilisation by nursing bearded seal (Erignathus barbatus) pups from Svalbard, Norway

In this study we measure energy intake via milk in nursing bearded seal (Erignathus barbatus) pups and determine how this energy is allocated into metabolism and storage of new tissues. This was accomplished using longitudinal mass gain records and the doubly labelled water technique on nursing pups in combination with cross-sectional data on changes in milk composition from bearded seal mothers. The pups (n = 3) were all less than a week old at the start of the experiments. Pups gained 3.3 +/- 0.4 kg.day-1 of which 50% was fat, 14% protein and 36% water. Average daily water influx for the pups was 69.5 +/- 9.0 ml.kg-1. day-1. Average CO2 production during the study period was 0.99 +/- 0.10 ml.g-1.h-1, which corresponds to a field metabolic rate of 642 +/- 67 kJ.kg-1. day-1, or 6.0 +/- 0.5 times the predicted basal metabolic rate according to Kleiber (1975). The pups drank an average of 7.6 +/- 0.5 kg of milk daily. This corresponds to a daily energy intake of 154 +/- 8 MJ, 47 +/- 14% of which was stored as new body tissue. Despite this high energy intake bearded seal pups do not get as fat as do other nursing phocids. This is in part due to their larger body size but also due to their very active aquatic lifestyle and the lower and more consistent fat content of the milk compared to other phocid species. Bearded seal mothers forage during lactation and may also be involved in teaching their pups to feed independently. All these data suggest that the lactation strategy of bearded seals differs from the phocid norm.     

Retention of maternal behavior in nulliparous and primiparous rats: Effects of duration of previous maternal experience.

Lactating primiparous and nonlactating pup-induced maternal nulliparous female CD rats were given 1, 4, or 9 days of maternal experience with rat pups before they were isolated from the young. 25 days later, Ss were reexposed to 3–8 day old foster pups, and latencies to show maternal behavior in the home cage and a T-maze test were scored. In the home cage, the latencies of all nulliparous groups were shorter upon reinduction. Comparisons of primiparous and nulliparous groups revealed that primiparous females (1, 4, and 9 days combined) carried a pup, crouched, retrieved, and grouped the pups and built a good nest faster than did nulliparous females (combined groups). The number of behavioral differences between specific primiparous and nulliparous groups decreased as the length of prior maternal experience increased. In the T-maze, latencies to retrieve a pup were shorter in primiparous females. Results indicate that the processes underlying establishment of the long-term retention of short-latency maternal behavior in these groups may be comparable. (20 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)