Maternal behavior deficits in nulliparous oxytocin knockout mice

The first observations of postpartum oxytocin knockout (OTKO) mice found no maternal behavior deficits. However, it is unclear how detailed those observations were. In this study, we compared maternal behavior exhibited by OTKO and wild-type (WT) nullipara toward six 2-4-day-old foster pups during test sessions conducted on 3 successive days. Each day, subjects were placed in a clean cage 30 min prior to introduction of pups which were deposited in a clump adjacent to the middle of a long wall of each test cage. Behavior was measured for 3.5 h after which pups and test subjects were returned to their home cages. On test days 1 and 3, a significantly smaller proportion of OTKO females retrieved pups to a corner of their cage. Also, significantly fewer pups were retrieved to corners by OTKO females. In contrast to most WTs, most OTKO females mothered pups in the center of the cage where they were initially deposited. Pup-licking frequencies were significantly lower in OTKO females. Their self-grooming frequencies also trended toward being lower. Latencies to retrieve and lick pups, latencies to and frequencies of still crouching over pups and proportion of time in nest did not differ between groups. Our findings suggest that OT stimulates a significant proportion of pup-licking in nulliparous mice, a situation similar to lactating rat mothers. Our results also indicate that OT may play a role in the motivation to retrieve pups to a more secure location.     

Maternal behavior,milk ejection,and plasma hormones in Hatano high- and low-avoidance rats

Hatano high- and low-avoidance (in a two-way active avoidance task) animals (HAA and LAA, respectively) were successfully selected from a Sprague-Dawley rat population. Pup growth of both strains was dependent on the maternal strain in a cross-fostering study. To determine whether there are strain-specific differences between HAA and LAA in maternal care, both strains of dams were subjected to a test battery as measured by nest building, home cage behavior, pup retrieval, and milk ejection tests. In addition, changes in plasma concentrations of lactotropic and corticotropic hormones such as prolactin, ACTH, and corticosterone were examined during lactation. The test battery indicated that the dams of both strains built good nests and spent an identical amount of time with their offspring. However, LAA dams showed a prolonged latency time for pup retrieval and often left pups outside the nest until the end of the test period. LAA dams also showed a decreased amount of milk ejection, whereas no strain differences were observed in milk ejection after oxytocin treatment. During lactation, a lesser increase in plasma concentrations of prolactin and a greater increase in ACTH were found in LAA dams. There were no differences between the two strains in plasma concentrations of corticosterone. These results clearly demonstrated decreases in maternal behavior and milk ejection in LAA as compared to HAA dams. The present results also suggest that maternal motivation and mechanisms responsible for maternal hormones related to suckling are involved in the degree of pup growth.     

Genetic variation in oxytocin rs2740210 and early adversity associated with postpartum depression and breastfeeding duration

Mothers vary in duration of breastfeeding. These individual differences are related to a variety of demographic and individual maternal factors including maternal hormones, mood and early experiences. However, little is known about the role of genetic factors. We studied single‐nucleotide polymorphisms (SNPs) in the OXT peptide gene (rs2740210; rs4813627) and the OXT receptor gene (OXTR rs237885) in two samples of mothers from the Maternal adversity, Vulnerability and Neurodevelopment study (MAVAN), a multicenter (Hamilton and Montreal, Canada) study following mothers and their children from pregnancy until 7 years of age. Data from the Hamilton site was the primary sample (n = 201) and data from Montreal was the replication sample (n = 151). Breastfeeding duration, maternal mood (measured by the CES‐D scale) and early life adversity (measured by the CTQ scale) were established during 12 months postpartum. In our primary sample, polymorphisms in OXT rs2740210, but not the other SNPs, interacted with early life adversity to predict variation in breastfeeding duration (overall F_8,125 = 2.361, P = 0.021; interaction effect b = ?8.12, t = ?2.3, P = 0.023) and depression (overall F_8,118 = 5.751, P ≤ 0.001; interaction effect b = 6.06, t = 3.13, P = 0.002). A moderated mediation model showed that higher levels of depression mediated the inverse relation of high levels of early life adversity to breastfeeding duration, but only in women possessing the CC genotype [effect a′ = ?3.3401, 95% confidence interval (CI) = ?7.9466 to ?0.0015] of the OXT SNP and not in women with the AA/AC genotype (a′ = ?1.2942, ns). The latter findings (moderated mediation model) were replicated in our Montreal sample (a′ = ?0.277, 95% CI = ?0.7987 to ?0.0348 for CC; a′ = ?0.1820, ns for AA/AC) .     

Brief communication: Self‐suckling in Barbary macaque (Macaca sylvanus) mothers before and after the death of their infant

We report here self‐suckling in four wild female Barbary macaques (Macaca sylvanus), living in two troops (i.e. “Flat face” and “Large” troop) in the middle‐Atlas Mountains, Morocco. The four females lost their infants due to predation or for unknown causes. Self‐suckling was observed before and after the infants died in the four females living in the “Flat face” troop. When the infants were still alive, self‐suckling was of short duration and it was probably a method to improve milk flow when the infant switched from one nipple to the other. After the infants died, self‐suckling lasted significantly longer and the females were apparently drinking their own milk. Self‐suckling was never observed among the four lactating females in the “Large” troop (including one monkey who lost her infant) and it could thus represent a cultural difference. Moreover, self‐suckling after the death of an infant may be explained by the energetic and immunological benefits that a monkey may gain from drinking their own milk. Finally, self‐suckling may have a stress‐releasing effect on the mothers who have lost their infants. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Effects of restricted nursing on milk production and collection, kid growth and plasma prolactin and growth hormone concentrations in dairy goats

The milk production of dairy goats under various regimes of mother-young contact from day 4 post partum were studied during the first 2 months of lactation, together with the prolactin (PRL) and growth hormone (GH) responses to udder stimulation. In the control group, 13 goats and their kids were left in permanent contact and did not undergo milking. In two additional groups, goats were machine milked once a day in the morning (at 0800 h) and kids were allowed 10 hours (from 1000 to 2000 h; 10H group, n = 11) or 5 h (from 1000 to 2000 h; 5H group, n = 11) of mother-young interaction per day. In the last group (MO, n = 10), mothers were permanently separated from their kids on day 4 post partum and milked once a day. Milk production during a 24-h period at 37 days post partum performed by controlled nursing and weighing of the kids (groups with kids) or by two machine milking 12 h apart (milking only group) revealed a higher production in the three groups with some mother-young contact than in the MO group. Total milk collected by milking over the 2 months of the study did not differ between the three groups that underwent milking. Kid weights at 2 months were 3.4 to 4.8 kg. lighter in the groups that underwent milking than in the control group. Hormonal profiles were significantly affected by restricted mother-young contact, with highest pre-stimulation concentrations of PRL and GH in the 5H group. Restricting mother-young contact from the first week postpartum can permit an early collection of milk without major effects on kid growth, when compared with one daily milking in goats totally separated from their young.     

Comparison of the "nursing" and other parental behaviors of nulliparous and lactating female rats

Virgin female rats display maternal behaviors after continuous exposure to pups (sensitization) that are in some respects similar to those of postpartum females. We herein provide a detailed comparison of the "nursing" and other parental behaviors of maternally sensitized virgin females and postpartum lactating dams. Ovariectomized and intact virgin females were exposed to pups until displaying maternal behavior. On the females' fourth day of maternal responsiveness, the pups were removed for 3 h and then returned, and subject-litter interactions were observed for 45 min. Behavior of maternal virgins was compared with that of lactating dams observed on day 4 postpartum interacting with either suckling pups or pups unable to suckle due to perioral anesthesia. Ovariectomy had no effect on behavior of virgins. Retrieval and licking of pups were deficient in virgins compared with lactating dams. Suckled dams showed prolonged kyphosis (upright crouched nursing), whereas nonsuckled dams displayed little kyphosis but rather were often in a hunched position over pups. Some aspects of quiescent "nursing" behaviors of virgins were surprisingly similar to those of suckled dams, including the latency to and duration of quiescence. Nonsuckling pup stimulation elicited more kyphosis in virgins than in lactating dams, which was still much less than in suckled dams. Virgins also "nursed" pups in hunched and prone postures. Differences between sensitized and postpartum females in their maternal behaviors likely reflect differences in motivation as well as sensory inputs they receive from pups. In particular, sensory regulation of "nursing" behaviors is influenced by reproductive state because nonsuckling pups elicit different postural responses in sensitized and lactating mothers.     

Increased aggression and lack of maternal behavior in Dio3‐deficient mice are associated with abnormalities in oxytocin and vasopressin systems

Thyroid hormones regulate many aspects of brain development and function, and alterations in the levels of thyroid hormone action lead to abnormal anxiety‐ and depression‐like behaviors. A complement of factors in the brain function independently of circulating levels of hormone to strictly controlled local thyroid hormone signaling. A critical factor is the type 3 deiodinase (DIO3), which is located in neurons and protects the brain from excessive thyroid hormone. Here, we examined whether a local increase in brain thyroid hormone action secondary to DIO3 deficiency is of consequence for social behaviors. Although we did not observe alterations in sociability, Dio3?/? mice of both sexes exhibited a significant increase in aggression‐related behaviors and mild deficits in olfactory function. In addition, 85% of Dio3?/? dams manifested no pup‐retrieval behavior and increased aggression toward the newborns. The abnormal social behaviors of Dio3?/? mice were associated with sexually dimorphic alterations in the physiology of oxytocin (OXT) and arginine vasopressin (AVP), 2 neuropeptides with important roles in determining social interactions. These alterations included low adult serum levels of OXT and AVP, and an abnormal expression of Oxt, Avp and their receptors in the neonatal and adult hypothalamus. Our results demonstrate that DIO3 is essential for normal aggression and maternal behaviors, and indicate that abnormal local regulation of thyroid hormone action in the brain may contribute to the social deficits associated with neurodevelopmental disorders.     

Parity-related differences in suckling behavior and nipple preference among free-ranging Japanese macaques

Suckling behavior of primiparae in free-ranging Japanese macaques (Macaca fuscata) was compared with that of multiparae at Jigokudani Monkey Park, the Shiga Heights, Nagano Prefecture, Japan, from April 1984 to June 1994. The estimated rates of milk secretion in a single preferred nipple among primiparae were lower than those among multiparae from birth to 5 months of age. Milk secretion capabilities of primiparous mothers, hence, appear to be inferior to those of multiparous mothers. In nutritive sucking, although Japanese macaque infants preferred one single nipple, nipple preferences in primiparae were weaker than those in multiparae. With supplementary two-nipple use during sucking, however, the infants of primiparae appear to overcome a suckling flaw of their primiparous mothers. After infants were 5 months of age and after a drop (from approximately 10% to approximately 5%) in the rates of milk secretion, however, milk secretion rates in a single preferred nipple among primiparae were similar to those in multiparae and the supplementary two-nipple use in primiparous mother-infant dyads disappeared. Am. J. Primatol. 42:331–339, 1997. © 1997 Wiley-Liss, Inc.     

Pasture Quality Affects Juvenile Survival through Reduced Maternal Care in a Mountain‐Dwelling Ungulate

Lactation exerts heavy energetic and physiological costs to mothers, whilst determining early growth and survival of offspring. To mountain ungulates, access to high‐quality forage during nursing and weaning is crucial for reproductive success. We have evaluated the effects of pasture quality on suckling behaviour and winter survival of Apennine chamois Rupicapra pyrenaica ornata kids, across three areas. Areas A‐B (‘poor’ areas) were characterised by a reduced availability of nutritious forage, thus a lower diet quality for female chamois and kids; Area C (a ‘rich’ area) included a much greater availability of nutritious forage. In poor areas, pasture quality has been reduced by climatic and plant composition changes, as well as the presence of a herbivore competitor (red deer Cervus elaphus). In poor areas, we recorded a significantly (1) lower suckling success of chamois kids (number of suckling bouts/number of suck attempts); (2) lower frequency of suckling bouts (n. suckling bouts/kid/h); and (3) lower suckling intensity (suck duration/kid/h) in respect to the rich area. Conversely, frequencies of suckling rejections and those of suckling attempts (n. events/kid/h) were the lowest in the rich area. Winter survival of chamois kids was c. 2 times greater in the rich area (45%) than in poor areas (20–26%). In the poor areas, resource scarcity induced adult female chamois to decrease maternal cares and favour their own maintenance, ultimately affecting population dynamics through kid winter mortality.     

Litter Size Determines Circadian Periodicity of Nursing in Rabbits

Nursing in rabbits occurs inside the nest with circadian periodicity. To determine the contribution of suckling stimulation in regulating such periodicity, we varied the size of the litters provided (1, 2, 4, or 6–8 pups). Nursing does, kept under a 14:10 (L:D) photoperiod, were continuously videotaped from parturition into lactation day 15. Although parturitions occurred throughout the day, a significant negative linear correlation (p?<?0.0001; r?=??0.68) was evident between time of delivery and time of nursing on lactation day 1, regardless of newborn number: longer intervals between these two events were seen in does delivering in the early morning than in those that gave birth late in the day. In rabbits suckling 6–8 pups, a Rayleigh analysis revealed that the population vector best describing their nursing pattern (across lactation days 1–15) had a phase angle?=?58° (corresponding to solar time 0352?h and rho?=?0.78; p?<?0.001). In contrast, the nursing pattern of does nursing litters smaller than 6 pups did not show circadian periodicity; rather, mothers showed multiple entrances into the nest box throughout the day. Cluster analysis revealed that the main equilibrium point of intervals between suckling bouts shifted from 24?h (6–8 pups) to 6?h (4 and 2 pups) and to as low as 4?h with 1 pup. In the groups nursing 2, 4, or 6–8 pups, most nursing episodes were followed by food and water intake. Those mothers also showed self-grooming of the ventrum and nipples after nursing. The incidence of these behaviors was lower in does nursing 1 pup. In conclusion, nursing in rabbits spontaneously occurs with circadian periodicity, but it is largely modulated by a threshold of suckling stimulation. (Author correspondence: gabygmm@gmail.com)     

The effect of a mixed-management system on the release of oxytocin, prolactin, and cortisol in ewes during suckling and machine milking

Milk yield and plasma oxytocin (OT), prolactin (PRL), and cortisol (CORT) during suckling and machine milking were measured in multiparous ewes subjected to a mixed management system of 3 sucklings and two daily milkings. Peak hormones were significantly increased and were similar during suckling and milking for PRL (181 vs. 163.3 ng x mL(-1)) and CORT (12.5 vs. 11.5 ng x mL(-1)). During the period of exclusive suckling, OT was always significantly released (90.3 pg x mL(-1)); however, during the period of mixed management, OT concentrations only increased during suckling compared to milking (91.7 vs. 13.1 pg x mL(-1)). The mean volume of milk obtained during suckling (632 mL) was significantly higher than during milking (255 mL). Thus, during a mixed management system, oxytocin and prolactin releases are not under similar central regulation. A mixed system, without OT release during milking, does not contribute to accelerate the conditioning of ewes for machine milking.     

Molecular and cellular mechanisms of the first social relationship: A conserved role of 5-HT from mice to monkeys,upstream of oxytocin

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Effect of prolactin and glucocorticoids on P-enolpyruvate carboxykinase activity in liver and mammary gland from diabetic and lactating rats

Summary The administration of 2 bromo--ergocryptine, to reduce serum prolactin decreased the activity of cytosolic P-enolpyruvatc carboxykinase (GTP) (EC4.1.1.32) about 50% in both liver and mammary gland of lactating animals. Adrenalectomy had similar effects to those of bromo-a-ergocryptine. In contrast, there was a 50% increase in enzyme activity in the mammary gland of diabetic, lactating rats and a 10-fold increase in liver as compared with normal rats. P-enolpyruvate carboxykinase activity in mammary gland as liver is coordinately regulated by prolactin, glucocorticoids and insulin.     

(d-Met2,Pro5)Enkephalinamide causes a decrease in plasma prolactin levels of lactating rats continuously suckled and an increase in those deprived of their litter

The effect of an enkephalin analogue, (d-Met²,Pro⁵)enkephalinamide (EKNH₂) on prolactin (PRL) secretion of lactating rats continuously suckled or separated from their pups was investigated. In rats together with their pups 0.5 mg/kg EKNH₂ caused a dramatic decrease, 0.25 mg/kg a mild and short-lasting reduction in plasma PRL levels. In contrast, in lactating rats separated for 4 h from their pups 0.5 mg/kg of the drug induced a slight and 1.0 mg/kg a considerable increase in plasma PRL levels. The data indicate that in lactating rats depending on the circumstances the enkephalin analogue causes opposite effects on PRL secretion.     

Intermittent suckling enables estrus and pregnancy during lactation in sows: effects of stage of lactation and lactation during early pregnancy

Previously we demonstrated that pre-ovulatory LH and post-ovulatory progesterone (P4) concentrations in plasma were low and embryo development was retarded when sows were induced to ovulate during lactation by submitting them to intermittent suckling (IS). The present study investigated whether this was due to: (1) stage of lactation when IS was initiated, and (2) continuation of IS post-ovulation. Multiparous Topigs40 sows were studied under three conditions: conventional weaning at Day 21 of lactation (C21; n = 30), intermittent suckling from Day 14 of lactation (IS14; n = 32), and intermittent suckling from Day 21 of lactation (IS21; n = 33). Sows were separated from piglets for 12 h daily during IS. IS sows were either weaned at ovulation or 20 d following ovulation. One-third (21/63) of the IS21 and C21 sows had already ovulated or had large pre-ovulatory follicles at Day 21 and were excluded from further study. Initiation of IS at Day 14 instead of Day 21 of lactation tended to reduce P4 at 7 d post-ovulation (P = 0.07), did not affect pregnancy rate, and tended to reduce embryo survival (P = 0.06). Continuation of IS during pregnancy resulted in lower P4 at 7 and 12 d post-ovulation, tended to reduce embryo weight and pregnancy rate (P < 0.10), whereas embryo survival was not affected. This study presents data for a population of sows in which follicle growth and ovulation are easily triggered under suckling conditions. Further, when these sows are bred during lactation, initiation of IS at 21 rather than 14 d of lactation with weaning at ovulation yields the most desirable reproductive performance.     

Maternal protectiveness and response to the unfamiliar in vervet monkeys

Vervet monkeys (Cercopithecus aethiops sabaeus) show individual differences in approach-avoidance behavior when faced with an unfamiliar and potentially threatening situation. Prior research from our colony demonstrated that juveniles who had experienced high levels of early maternal protectiveness were more cautious in response to novelty, compared to juveniles who had had less protective mothers. The research reported here was designed to verify this result in a paradigm that experimentally varied maternal protectiveness through the introduction of new breeding adult males. Mothers responded to the presence of new males by increased maternal protectiveness toward infants born in the year following the introductions. Individual differences in response to the unfamiliar were later evaluated by measuring the latency to approach within 1 m of novel food containers placed into the home enclosure of four naturally composed social groups. Infants approached with the same latency and in the same order as their mothers. Juveniles approached sooner than, and independent of, their mother's current behavior, but their latency to approach could be predicted by the experimentally induced variation in maternal protectiveness they had experienced as infants. Immatures who had been born in New Male years, when maternal protectiveness was high, were more cautious and had significantly longer latencies to approach the novel stimulus compared to immatures who had been born in Resident Male years. © 1993 Wiley-Liss, Inc.     

Brain vasopressin signaling modulates aspects of maternal behavior in lactating rats

The brain vasopressin system mediates various social behaviors as has been studied mostly in males. Only recently, advances in social neuroscience revealed that central vasopressin signaling via its V1a and V1b receptors also facilitates female social behavior, including maternal behavior. In this review, we show how maternal care, maternal motivation and maternal aggression of lactating rat mothers are modulated in a V1 receptor subtype‐ and brain region‐specific manner. Measuring local release pattern of vasopressin via intracerebral microdialysis in the behaving rat mother as well as using pharmacological approaches to activate or block vasopressin receptors with subsequent behavioral observation provide detailed insight into the functional role of the vasopressin system in maternal behavior. In this context, the complementary rat animal model of high (HAB) and low anxiety‐related behavior (LAB) is particularly helpful due to the genetically determined high activity of the vasopressin gene in HAB rats, which also underlies their high levels of maternal behavior. Furthermore, first studies in humans indicate that the vasopressin system in general and the V1a receptor in more particular might mediate mothering.     

Maternal chitosan oligosaccharide supplementation affecting expression of circadian clock genes,and possible association with hepatic cholesterol accumulation in suckling piglets

Chitosan oligosaccharide (COS) has been shown to reduce lipid accumulation in liver in mice and rats. The purpose of this study was to investigate whether maternal COS feeding affects hepatic lipid metabolism via influencing the expression of circadian clock genes in piglets. From day (d) 85 of gestation to d 14 of lactation, sixteen pregnant sows were divided into a control group (basal diet without COS supplementation) and a COS group (30 mg COS/kg basal diet). After farrowing, one piglet per litter in each group was selected for the collection of plasma and liver samples on d 0 and d 14 of age, respectively. Interestingly and significantly, we found that maternal COS supplementation promoted plasma and hepatic cholesterol accumulation and up-regulated the mRNA level of negative-regulated element period 1 (Per1), and reduced the abundance of the positive elements, circadian locomotor output cycles kaput (CLOCK), and brain muscle Arnt-like 1 (BMAL1) in the suckling piglets on d 14. These alterations may promote the hepatic cholesterol accumulation, which, in turn, activates hepatic bile acid metabolism and attenuates the relative expression levels of lipid metabolism-associated genes in the liver. However, the expression of CLOCK and BMAL1 and the lipid profile in the plasma and liver were not affected by COS supplementation on d 0. Collectively, our results indicate that maternal supplementation with COS postpartum up-regulates cholesterol accumulation in suckling piglets at age d 14, in part, by the regulation of circadian clock genes.     

Dopaminergic mediation of Y-aminobutyric acid in the control of prolactin release: Plasma prolactin and brain tyrosine hydroxylase levels in ovariectomized conscious rats

The present experiments were carried out to further elucidate the mechanism by which dopamine mediates the actions of Y-aminobutyric acid on prolactin release from anterior pituitary following its intraventricular injection in overiectomized conscious rats, Y-Aminobutyric acid significantly suppressed the prolactin levels at 0.1 Μmol concentration while at 4 Μmol dose, the level was elevated. The activity of tyrosine hydroxylase was increased significantly in the anterior pituitary at the lower dose while the higher concentration of Y-aminobutyric acid did not bring about any change in the activity both in the hypothalamus and the anterior pituitary. The results presented suggest that intracellular dopamine in the anterior pituitary may directly inhibit prolactin release; the plasma prolactin level is elevated by Y-aminobutyric acid, by way of either inhibiting dopaminergic tone or possible stimulation of a physiological prolactin releasin g hormone.