Ontogeny of suckling in rats: Transitions toward adult ingestion.

Studied the suckling behavior of neonatal Sprague-Dawley rats on their own anesthetized mother from the day of birth until after weaning (35 days). Even newborn Ss were capable of nipple attachment without maternal assistance. Before 11–23 days of age, pups deprived of suckling for 22 hrs and nondeprived pups quickly attached to their mother's nipples, sucked, and remained attached to the nipple though no milk delivery occurred. Then behavior underwent at least 3 changes: (a) After 11–23 days of age, latency to attach became considerably elevated in nondeprived pups; (b) a 2nd change in latency occurred at 23–25 days of age, when nondeprived pups no longer even attached to the nipples of anesthetized mothers; and (c) about 14 days of age, deprived pups began to shift from 1 nipple to the next after initial attachment. These developmental changes were seen in other test situations in which pups were placed directly on a nipple and not required to search and when various periods of deprivation were utilized. These transitions were not critically dependent on the onset of visual function or on the pup's experience with food other than mother's milk. Suckling, therefore, in not an unmodified reflex but is an appetitive behavior that undergoes a series of changes during development. These transitions constitute major developmental events in the ontogeny of rat ingestive behavior. (48 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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.     

Methysergide-induced nipple attachment depends on suckling experience in juvenile rats.

These studies were designed to investigate the links between pharmacological and behavioral procedures that facilitate suckling in weanling rats by assessing the effects of methysergide on nipple attachment behavior following experiential manipulations known to either promote or attenuate suckling. In the first experiment, methysergide failed to stimulate suckling in 25-day-old rats separated from their dam on Day 20, although it facilitated suckling in rats kept with the dam until either Day 24 or Day 25. In the second experiment, methysergide did not facilitate suckling in 35-day-old rats separated on Day 25, although rats separated on Day 30 or 34 were induced to suckle. In the third experiment, rats were housed with preweanling litters until Day 35 and then separated, housed with 25-day-old litters, or housed with preweanling litters for another 10 days. Rats in a fourth group remained with their dams until Day 35 and were then separated for 10 days. When tested at Day 45, nipple attachment was facilitated by methysergide only in the three groups that had received extended suckling experience. These results demonstrate that serotonergic inhibitory mechanisms modulate nipple attachment only when suckling experience is recent or extensive. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Postnatal undernutrition: an alternative method

Rat pups were undernourished from birth by placing them for 12 hr/day with a normal lactating mother and 12 hr/day with a nipple-ligated mother each day for 25 days. The method resulted in a marked delay in the body growth of the undernourished pups, especially during the first 2 weeks of life. Observations of the behavior of the mothers towards the underfed pups were made at different times of the day and compared to the behavior of the mothers suckling well-fed pups. The results show that (1) nipple-ligated mothers are able to provide adequate maternal care for undernourished pups, and (2) both ligated and nonligated mothers caring for underfed pups spend more time with those pups than mothers caring for well-fed pups.     

Olfactory, thermal, and tactile controls of suckling in preauditory and previsual rats.

Determined exteroceptive control of nipple attachment in Sprague-Dawley rats and their litters by altering maternal sensory features. Preauditory and previsual pups, 2, 5, 8, and 11 days of age, were permitted to locate and attach to the nipples of their anesthetized mother after she had received one of the following treatments: (a) thermal--peripheral temperature was lowered to either 31 or 28°C; (b) tactile--the mother was shaved; (c) thermotactile--the mother was shaved and her skin temperature was lowered to 31°C; or (d) olfactory--all nipples were cleaned with methylene chloride and alcohol solvents. Only the olfactory treatment severely affected nipple attachment at all ages studied. Washing the nipples virtually eliminated suckling; painting them with a vacuum distillate of the nipple extract or of pup saliva reinstated suckling. The other manipulations either did not interfere with nipple attachment or did so at one age only. (42 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Light-dark variation and changes across the lactational period in the behaviors of undisturbed mother and infant guinea pigs (Cavia porcellus).

Lactating guinea pigs (Cavia porcellus) and their litters were observed by videophotography across the light/dark cycle at 1, 11, 21, and 31 days postpartum. The highest level of behavioral activity was seen in the dark, particularly in the hour after light offset. This circadian pattern was evident from Day 1 in mothers and from Day 11 in pups. Contact between mothers and pups was inversely related to activity, occurring more frequently during light. Maternal grooming of pups occurred on Day 1 and then declined; self-grooming by pups increased across days. Intake of solid food and water by pups occurred on Day 1 and increased thereafter. A nearly complete transition from nursing to independent ingestion was observed between 21–31 days of age. Overall, we document several ontogenetic changes in young guinea pigs and demonstrate that under laboratory conditions mother and infant guinea pigs exhibit a nocturnal activity pattern. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The role of experience in suckling maintenance in albino rats.

In Exp I nipple-attachment behavior in Sprague-Dawley albino rats was disrupted by raising rat pups in isolation from their mother and siblings on the 3rd–5th day after birth. In Exp II nipple attachment was maintained, however, in isolation-reared pups that received, on Days 3 and 4 postnatally, either 6 or 12 opportunities to search for, locate, and attach to the nipples of an anesthetized mother. Suckling remained severely disrupted on Day 5 in pups whose experiences on Days 3 and 4 were restricted to either nipple search alone or nipple attachment without previous search. Findings focus attention on the role of experience in suckling maintenance and suggest that in rats the suckling system is not fully specified at birth. (35 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Postsuckling behavioral arousal in weanling rats (Rattus norvegicus).

Twenty-day-old litters and their dams were observed in seminatural habitats consisting of a nest compartment and adjacent open field that contained powdered rat chow. It was found that pups displayed marked bursts of activity after suckling. Independent feeding reliably followed nursing bout termination (Experiment 1). Nipple withdrawal, with or without milk transfer, induced behavioral arousal whereas withdrawal of thermotactile and conspecific odor cues did not (Experiments 2-3). Increased thermogenesis was observed following milk transfer (Experiment 4). Finally, preweanling pups (10- to 12-day-olds) also displayed postsuckling arousal within the confines of the nest; full locomotor expression of this arousal was not evident until weaning age (Experiment 5). It was concluded that postsuckling arousal in weanlings functions to stimulate activities performed away from the nest and suckling, propelling pups into the field where feeding begins. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effect of kitten vocalizations on maternal behavior.

Observation of 8 mother cats and their litters revealed that the previous occurrence of a kitten vocalization was associated with an increased probability that mothers would approach kittens, change to a different body position, shift their lactation position, or vocalize. Experimental study, in which the effects of kitten vocalizations were separated from other kitten-emitted cues, demonstrated that vocalizations reliably elicited approach and investigation of the sound source from mothers. Further, mothers exposed to kitten vocalizations removed kittens from a box and carried them to the home site. Repeated testing of mothers demonstrated that these effects of kitten vocalizations on maternal behavior remained strong across at least the 1st 30 days of life. It is concluded that vocalizations are a stimulus by which kittens influence their mother's behavior. (31 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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

Parental behavior, food consumption, and pup development in litters of CD-1 albino mice (Mus musculus) with single mothers or paired mothers in 3 different housing conditions were compared. Fewer pups survived when parents were required to run in a wheel to obtain food, and these pups had lower weights than pups in the wheel-noncontingent and no-wheel groups. The presence of the male facilitated pup survival in the wheel-contingent group but had no effect on pup growth. Mice in the wheel-contingent group ran more revolutions than those in the wheel-noncontingent group and spent less time in parental care. In the wheel-contingent group, males spent more time running and ate less food than females, suggesting that males may indirectly facilitate pup development by providing food for the mother. The results suggest that the presence of the male increases pup care and may facilitate maternal behavior under appropriate environmental conditions. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Translactational induction of CYP4A expression in 10.5-day neonatal rats by the hypolipidemic drug clofibrate

Lactating mothers of 7.5-day neonatal rats were injected intraperitoneally with 500 mg kg-1 clofibrate for 3 consecutive days at 24-hour intervals; 24 hours after the final injection, the maternal cytochrome P450 4A (CYP4A) mRNA levels had risen 14- and 2.5-fold above the constitutive levels of expression seen in the liver and kidney, respectively. Lactational transfer of clofibrate to the suckling 10.5-day litter was demonstrated by the 15- and 5-fold elevation observed in the neonatal hepatic and renal CYP4A mRNAs, respectively, following suckling from drug-induced mothers. A significant decrease in the relative liver weights of these neonatal pups was seen following clofibrate exposure via maternal milk, in total contrast to the normally observed increase in liver/body weight ratios of rats treated with clofibrate. Western blot analysis using a polyclonal goat anti-rat CYP4A1 antibody also demonstrated a rise in the CYP4A protein levels in both the mothers and their litters following maternal clofibrate treatment.     

Influence of the social environment on parental behavior and pup development of meadow voles (Microtus pennsylvanicus) and prairie voles (M. Ochrogaster).

Parental behavior and pup development in meadow and prairie voles were examined. Social units were manipulated for the presence of fathers and of juveniles. Meadow vole mothers alone with pups showed less maternal care and spent more time resting than did mothers with juveniles or with fathers and juveniles. Pups reared only with mothers developed faster than did pups under other conditions. Number of animals in the social unit was negatively correlated with the rate of pup development. Meadow vole fathers and juveniles showed no parental care. In contrast, prairie vole mothers spent less time in the nest when both fathers and juveniles were present. Fathers assisted in rearing pups, and litters developed more rapidly when fathers were present. Juveniles remained in the natal nest and engaged in parentlike behavior. The findings relate to differences in the life history strategy for the 2 vole species. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Mother as shield: Differential effects of contact and nursing on pain responsivity in infant rats: Evidence for nonopioid mediation.

To determine how rat mothers protect their pups against pain, we applied focal heat (34–51°C) to the ear or shoulder of 10-day-old rats that were isolated in contact among themselves or with their mother, suckling nonnutritively, or in the hyperextension position normally caused by milk letdown. Relative to isolated rats, contact doubled withdrawal latencies from heat (43 or 45°C) applied to the ear. Suckling quadrupled heat-escape latencies. During hyperextension, rats essentially did not escape from thermal stimulation of up to 48°C. Protection provided by maternal contact, especially suckling, was not mediated by either mu or kappa opioid receptors: Neither systemic injections of naltrexone nor norbinaltorphimine reduced heat-escape latencies. Morphine (0.125 and 0.250 mg/kg) added to the effects of contact but multiplied the effects of suckling to produce heat-escape latencies that were upward of 2 min. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Combination therapy of daily intramuscular injection of interferon-alpha and oral administration of fluorouracil for advanced renal cell carcinoma]

The present study, using proton nuclear magnetic resonance relaxation (H1 NMR) measurements, was undertaken to quantitate water fractions with different mobility in the brain tissue obtained form New Zealand White rabbit pups. Serial studies were carried out at the postnatal age of 0-1, 24, 48, 72, and 96 h in pups nursed with their mothers and suckling ad libitum (group I) and in those pups separated from their mothers and completely withheld from suckling (group II). Tissue water content (desiccation method) and T1 and T2 relaxation times (H1 NMR method) were measured. Free, loosely bound, and tightly bound water fractions were calculated by applying multicomponent fits of the T2 relaxation curves. It was demonstrated that brain water content and T1 and T2 relaxation times did not change with age in the suckling pups. In pups withheld from suckling brain water decreased from 89.4 +/- 0.5% at birth to 87.7 +/- 0.1% at the age of 96 h (p < 0.05), T1 remained unchanged, and there was a significant fall in T2 by the age of 72 h (188 +/- 12 versus 178 +/- 4 ms, p < 0.05) and 96 h (171 +/- 6 ms, p < 0.01). Partition of brain water into bound and free fractions as derived from biexponential fits of T2 decay curve showed that the percent contribution of bound water fraction in pups of group I fell progressively from 61% at birth to 3% at the age of 72-96 h (p < 0.05). This fall was accelerated by the complete deprival of fluid intake, and the level of about 4% could be attained as early as the age of 24 h. Triexponential analysis of T2 relaxation curves revealed that the loosely bound fraction (middle component) predominated over the free (slow component) and the tightly bound (fast component) water fractions. In response to withholding fluid intake, the free water fraction increased 4-fold at the expense of tightly bound brain water. It is concluded that the majority of neonatal brain water is motion-constrained. The free, the loosely bound, and the tightly bound water fractions appear to be interrelated; from the brain water store water can be released to supply free water for volume regulation.     

Suckling behavior and intake control in the developing rat pup.

Studied the behavioral development of suckling and intake control in 2 experiments with Charles River CD strain rat pups. Ss were observed at the initiation, during the course, and at the termination of suckling from their anesthetized mothers. Diet was delivered intraorally through a fine tongue cannula which enabled control of timing and volume. The control of diet intake and the behavior at termination of suckling showed correlated changes from 5 to 20 days of age. When deprived of suckling (and food and water) for 8 hr, 5- and 10-day-old Ss consumed large volumes of diet (10% of body weight or greater) and terminated suckling only in the presence of extreme gastrointestinal filling. These Ss were immediately lethargic and slept after intake termination. Five-day-old Ss persisted in reattaching to the nipple when manually stimulated; 10-day-old Ss eventually refused to reattach. In contrast, 20-day-old Ss consumed more moderate volumes of diet (5% of body weight). These Ss also remained awake for a period after feeding and engaged in the exploratory and grooming activities characteristic of adult rats at the termination of feeding. These observations demonstrate major changes in suckling behavior during development. They suggest that intake control processes shift from indirect to direct and become more effective and specifically food intake related in older pups. (20 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Decline of maternal behavior in the virgin and lactating rat.

Gave to sensitized virgins and postpartum lactating mothers, both exhibiting maternal behavior, donor litters that increased in age by 1 day, for 28 days, starting at the onset of maternal behavior. Ss were 18 Charles River rats. Each day Ss were tested for maternal behavior with 4-8 day old pups. Maternal care (i.e., nursing-crouching, retrieving, nest building, and licking and maternal withdrawal, rejection, and prevention of nursing were recorded. After the 9th day, Ss were also tested with the progressively older pups 10-28 days of age with which they were living. Virgins and lactating mothers showed generally similar patterns of maternal care although some differences were found, and they declined in maternal behavior toward the older pups in a similar manner. Maternal behavior did not decline in tests with younger pups. Results are interpreted as supporting the hypothesis that the decline as well as the maintenance of maternal behavior postpartum is nonhormonally mediated. (24 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Intragastric intubation: important aspects of the model for administration of ethanol to rat pups during the postnatal period

One technique for the controlled delivery of ethanol to neonatal rat pups is intragastric intubation. Often, the vehicle used for delivery of ethanol is composed of a nutrient mixture to compensate for decreased suckling or other possible nutritional compromise. This study analyzed the selection of nutrient vehicle, the combination of experimental treatment groups within a litter, and the overall litter size on the growth rate of ethanol-intubated and intubated-control pups, compared with mother-raised control pups. Sprague-Dawley rat pups were raised in litters of 8 or 10, and administered ethanol by intragastric intubation with 20% (v/v) Sustacal or 80% (v/v) Intralipid-II nutrient vehicle. Pups were treated between postnatal days 2 and 10, and body weight was analyzed on day 10. Pups were assigned to a treatment group as either intubated ethanol, intubated control, or nonintubated mother-raised controls. Experimental comparison by statistical analyses was performed to identify the optimal treatment design (mixed treatment groups in a single litter or a single treatment group per litter), the optimal vehicle (Sustacal or Intralipid-II), and the optimal number of pups per litter (8 vs. 10). The analyses demonstrate that the mixing of intubated control, intubated ethanol, and nonintubated mother-raised control treatment groups within a single litter introduced an uncontrolled variable that confounded measurement of ethanol-specific alterations. The sensitivity of treatment groups to inclusion in mixed litters was dependent on the nutrient vehicle and thus nutritional adequacy. Our results suggest that an optimal design was achieved with eight pups per litter. Furthermore, ethanol intubated and intubated control pups grow at a rate identical to parallel litters of eight mother-raised control pups when Intralipid-II is used as nutrient vehicle, and a single treatment group is present in a litter. Optimization of these experimental parameters has provided an excellent neonatal rat model for analysis of specific ethanol effects on brain development during the third trimester.     

Evidence for a transitional period for development of persistence in infant rats.

Conducted 4 experiments with a total of 188 rat pups to test the hypothesis that rat pups can learn persistence at age 11 days but cannot express this learning in their behavior until they are a few days older. Exp I used a space-trial procedure to investigate the relative reinforcing effects of milk suckling, dry suckling, and contact without suckling at ages 11 and 14 days. Exp II used the Exp I procedure, manipulating schedule of reward at 2 ages and under 2 of the reward conditions. Exp III manipulated the dry suckling reward schedule in 11- and 14-day-old pups. Exp IV manipulated partial and continuous reinforcement on Day 11 but tested for persistence on Day 14. Results suggest the existence of a traditional period in neonatal rats for the learning of persistence. Partial reinforcement training with suckling on an anesthetized dam as reward induced greater persistence in extinction of the approach response than did continuous reinforcement in rat pups 14 days old but not in 11-day-old pups. Other aspects of this period of development are discussed. (26 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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.     

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)