Presence of the father and parental experience have differentiated effects on pup development in Mongolian gerbils (Meriones unguiculatus)

Conflicting data exist on the importance of the father and parental experience during development in rodents. The purpose of this study was to investigate the effects of these two variables on development in Mongolian gerbils. Forty pairs of males and females with a litter size of between 4 and 7 pups were used as subjects. Twenty couples had no experience in raising young. After the birth of their pups, four experimental groups were formed: (I) inexperienced mother and father; (II) inexperienced mother; (III) experienced mother and father and (IV) experienced mother. When the pups reached 10 days of age, pup and parental behavior was recorded in experimental sessions of 15 min on 11 consecutive days. Through the statistical analysis it was found that the presence of the father significantly increased the physical contact between pups and parents and that pups opened their eyes earlier in comparison to the groups without the father. On the other hand, parental experience had a significant influence on the behavior of the pups (locomotion inside and outside the nest, and self-grooming). The results of this study suggest that parental experience and the presence of the father have differentiated effects on development in Mongolian gerbils.     

Birth and simultaneous rearing of two litters in a pack of captive African wild dogs (Lycaon pictus)

Two female African wild dogs (Lycaon pictus) in a pack of 10 animals at the Bronx Zoo gave birth to litters of nine and seven pups within a 2‐day period. Two pups from the litter of seven died before they were 2 weeks old, but the other 14 survived past weaning. Litter size, sex ratio, and pup developmental stages were all consistent with data obtained from other captive‐ and wild‐born litters. Both dams were very attentive to their litters, and during the pups' first 2 weeks spent >90% of their time in the dens with the pups. By 2 days after birth and during their first 2 weeks of life, the pups spent 86% of their time in a nursing position. Before parturition and during the first week post‐partum, one of the dams (DAL) was clearly dominant to the other (WHI). However, the females' dominance rank reversed 2 weeks post‐partum. On two occasions during the pups' first 2 weeks DAL stole and reared one of WHI's pups, but between Weeks 3 and 4 all of DAL's pups were stolen and WHI raised both litters as one until they were weaned. The pups began spending time out of their den at approximately 1 month of age. The timing of the births, the design of the wild dogs' management facility, and the presence of several dens in different enclosures within the facility all likely contributed to the successful rearing of the litters. The pups in both litters were very similar in size because of the short interval between births, so one litter did not have a competitive advantage over the other with respect to gaining access to the dam for milk. The facility's design helped mitigate aggression within the pack, and the presence of multiple dens enabled to dams to move the pups to different den sites and allowed the two females to stay visually and spatially apart from each other while remaining with the pack. Zoo Biol 0:1–17, 2006. © 2006 Wiley‐Liss, Inc.     

Mechanisms of Change in the Ontogeny of Black-tailed Prairie Dog Time Budgets

This study examines the behavioral mechanisms underlying changes in the time allocation patterns of black-tailed prairie dog (Sciuridae: Cynomys ludovicianus) pups. Previous analyses of time budgets had revealed substantial differences between pups (young of the year) and adults, and that pup time budgets changed significantly over time. This paper shows that these differences were due to two mechanisms: pups generally differed from adults in the frequency of performance of different behaviors; however, changes in pup behavior over time were due to changes in both bout frequencies and durations. Other influences on bout durations were also examined. Time of day dramatically affected the duration of most behaviors, with vigilance bouts longest in the morning and feeding bouts longest in the evening. The presence of a pup's mother above ground and the height of the vegetation a pup was in had much less influence on bout durations. Multiple regression analyses were used to identify other potential influences (e.g., weather and social conditions) on bout lengths. Of these, the distance of a pup from the nearest burrow produced the strongest relationships.     

Diet of lactating South American sea lions,as inferred from stable isotopes,influences pup growth

Serum and blood cell δ¹³C and δ¹⁵N signals from 26 suckling pups of the South American sea lion from northern Patagonia were used as proxies of the composition of their mothers' diet to test the hypothesis that the foraging habits of the mother influence pup growth. Samples of primary producers and the female potential prey were analyzed to establish baseline isotopic values and to determine energy density. Pups were weighed to determine specific growth rate. Individual variability in female diet was large, probably as a consequence of dissimilarities in the foraging performance that depends on the individual's age, body size, and/or foraging skills. Growth of a pup was influenced by its mother's diet, as pups of females mostly relying on pelagic offshore prey were found to grow faster than those of females basing their diet on benthic coastal prey.     

DEVELOPMENT OF DIVING BY HARBOR SEAL PUPS IN TWO REGIONS OF ALASKA: USE OF THE WATER COLUMN

Satellite-linked dive recorders were attached to 53 harbor seal pups in Prince William Sound (PWS) and at Tugidak Island, Alaska, during 1997–1999. We used generalized additive models and bootstrap techniques to describe pup diving behavior during their first year of life. Pups increased their ability to dive during the first 3–6 mo, as indicated by increases in proportion of time in the water (time wet) and maximum dive depth achieved by a pup each day (max-depth) values. Time wet and/or max-depth later decreased, suggesting a seasonal component to diving behavior. Monthly time wet varied from an overall minimum of 0.68 at tagging in July to a maximum of 0.89 in November. Pups spent half of their time wet swimming in water <25 m deep, the shallowest 30% of the available water column. They spent only 5% of their time swimming in the deepest 30% of the available water column, at depths >60–70 m. This strongly suggests they were not feeding on or near bottom during their first year. Average max-depths and deepest actual dives were similar for PWS and Tugidak pups. PWS pups dove deeper sooner and spent less time wet than Tugidak pups during the first few months after tagging, probably as a result of regional bathymetric differences. Diving behavior and body condition suggest that food availability was not likely a major factor in the population decline in PWS during the period of this study.     

Oxytocin mediates the acquisition of filial,odor-guided huddling for maternally-associated odor in preweanling rats

The present study was designed to examine possible roles of oxytocin (OT) in the acquisition of a filial huddling preference in preweanling rats. We used a procedure in which a scented, foster mother can induce an odor-guided huddling preference in preweanling pups, following a single, 2-h-long co-habitation (Kojima and Alberts, 2009, Kojima and Alberts, 2011). This single, discrete period for preference learning enables us to observe the mother–pup interactions that establish the pups' preferences and to intervene with experimental manipulations. Four, 14-day-old littermates interacted with a scented foster mother that provided maternal care during a 2-h session. Two of the pups were pretreated with an intracerebroventricular injection of OT or an oxytocin antagonist (OTA), and the others received a vehicle injection. Filial preference for a maternally-paired odor was measured in a huddling test the next day. OT is necessary for acquisition of the filial preference: The preference learning was blocked in the pups treated with OTA, but not in their vehicle-treated littermates who experienced the same mother at the same time. Injection with exogenous OT did not augment the pups' preference. Manipulating pups' central OT also altered the contact interactions of the mother and pups. When some pups received OT, mother–litter aggregations formed as frequently and with similar combinations of bodies, but contact aggregations were significantly more cohesive than when some pups in the litter received OTA. We discuss dual, behavioral and neuroendocrine roles of OT in social learning by preweanling rats.     

Effects of repeated pup exposure on behavioral,neural, and adrenocortical responses to pups in male California mice (Peromyscus californicus)

In biparental mammals, the factors facilitating the onset of male parental behavior are not well understood. While hormonal changes in fathers may play a role, prior experience with pups has also been implicated. We evaluated effects of prior exposure to pups on paternal responsiveness in the biparental California mouse (Peromyscus californicus). We analyzed behavioral, neural, and corticosterone responses to pups in adult virgin males that were interacting with a pup for the first time, adult virgin males that had been exposed to pups 3 times for 20 min each in the previous week, and new fathers. Control groups of virgins were similarly tested with a novel object (marble). Previous exposure to pups decreased virgins' latency to approach pups and initiate paternal care, and increased time spent in paternal care. Responses to pups did not differ between virgins with repeated exposure to pups and new fathers. In contrast, repeated exposure to a marble had no effects. Neither basal corticosterone levels nor corticosterone levels following acute pup or marble exposure differed among groups. Finally, Fos expression in the medial preoptic area, ventral and dorsal bed nucleus of the stria terminalis was higher following exposure to a pup than to a marble. Fos expression was not, however, affected by previous exposure to these stimuli. These results suggest that previous experience with pups can facilitate the onset of parental behavior in male California mice, similar to findings in female rodents, and that this effect is not associated with a general reduction in neophobia.     

Potential of the Newborn Rabbit for Circadian Rhythms Research

Developmental studies of circadian function in mammals are generally difficult because of the close interaction between mother and young. The European rabbit presents an exception, providing developmental chronobiologists with an unusual opportunity to study the early development of circadian function. Doe rabbits only visit their newborn young once a day to nurse for a few minutes, and pups anticipate this regular event with heightened arousal and by uncovering of the nest. Both the mother's nursing visit and pups' anticipatory arousal represent well synchronized circadian rhythms. They also represent discrete, quantifiable events that can be readily manipulated by controlling does' access to the pups, by cross-fostering or by eliminating a nursing either before, during or after the development of visual function. The doe's long absence makes it possible to carry out surgical or other interventions without disrupting the normal pattern of maternal care, and the correspondence between pups' anticipatory arousal and the expression of c-Fos in hypothalamic nuclei demonstrates the suitability of this model for investigating the neural basis of early circadian function.     

Self-Grooming as a Tactic Used by Prairie Voles Microtus ochrogaster to Enhance Sexual Communication

Observations of numerous mammals suggest males self-groom more than females in response to the odours of opposite-sex conspecifics. Two experiments tested the hypothesis that self-grooming may be a tactic used by males to attract mates in prairie voles Microtus ochrogaster . In the first experiment, we measured the amounts of time voles self-groomed during exposure either to male-scented cotton bedding, female-scented cotton bedding, or clean cotton bedding. Results from this experiment support the hypothesis and also show that female prairie voles self-groom in response to odours of males. In addition, male prairie voles groom more in response to male odours than to female odours, suggesting that self-grooming also serves a role in male–male competition. In the second experiment, male and female voles spent more time investigating scent marks of opposite-sex conspecifics that recently self-groomed at a higher rate than those of opposite-sex conspecifics that self-groomed at a lower rate. Female, but not male prairie voles, spent more time investigating scent marks of opposite-sex conspecifics that self-groomed at a high rate than those of same-sex conspecifics that self-groomed at a high rate. For prairie voles, self-grooming may increase the detection of their scent marks by conspecifics. By self-grooming, prairie voles may be attempting to attract opposite-sex conspecifics, and males may also be attempting to deter encounters with male conspecifics.     

Behaviour of female and pup grey seals Halichoerus grypus during the breeding period at Froan, Norway

We studied the behaviour of grey seals, Halichoerus grypus , during the breeding period at Froan, Norway, and compared our findings with existing studies on grey seals at breeding sites in Britain and Canada. The pups at Froan spent more time in water than pups at other breeding sites. While on-shore, the pups at Froan spent most of their time resting, behaviour similar to pups at other breeding sites. Lactating females at Froan spent most of their time in the sea, thus differing from females at most other breeding sites which spend most of their time on-shore. While in the sea, the females at Froan spent most time diving, typically interrupted by regular periods of surface swimming.     

The importance of paternal care on pup survival and pup growth in Peromyscus californicus when required to work for food

We examined paternal care in the monogamous California mouse when wheel running was necessary to obtain food. Paternal and maternal care, pup survival and feeding were compared in litters with single females (Male Absent) and paired females (Male Present) under three different foraging requirements. When wheel running was required for obtaining food (Wheel Contingent), the male's presence significantly increased pup survival. There were no significant effects of the male's presence on pup survival in the No Wheel (standard laboratory housing with ad lib food) or Wheel Noncontingent (running wheel with ad lib food) groups. Males may have indirectly increased pup survival by helping generate food through wheel running and then subsequently spending less time eating than females. While females reduced the time spent in parental care as pups aged, males increased the time spent in parental care as pups aged. Thus as pups became larger and more expensive to rear, the male increased parental care. These results indicate that paternal care benefits pup survival more under conditions where the parents must forage to obtain food than when food is provided ad lib in the standard laboratory environment.     

Prolactin levels and maternal behavior induced by ultrasonic vocalizations of the rat pup.

The relationship among ultrasonic vocalization (USV), prolactin and maternal behavior was investigated in lactating rat mothers and their pups. The lactating mother had a cannula inserted into the external jugular vein, and was exposed to USVs emitted from a pup immediately. Changes of prolactin and maternal behavior were determined. Prolactin increased dramatically during exposure to USVs, when maternal search, retrieving and nest building behavior appeared significantly. These results suggested that the relationship among USV, prolactin and maternal behavior was included in communication between lactating mother and pup.     

Daytime rest behavior of the Welsh pony (Equus caballus) mare and foal

Upright and recumbent rest of 15 Welsh pony foals and their mothers was studied over a 2 year period. During their first week of life, the foals spent 32% of the time in recumbent rest. Subsequently, the percentage of time spent in recumbent rest decreased, but was still greater than for the foal's mother by Week 21, when the foals spent 6.5% of their time in recumbent rest. Adults spent little time in recumbent rest. Foals rested upright only 3.5% of the time during their first week of life. Mares rested upright more than foals did to Week 13, at which time peak values for time spent in upright rest occurred for both mares (32.5%) and foals (23%). Subsequently, mares and foals spent equal, but decreasing, amounts of time resting upright. The total time spent resting by the foals decreased gradually, and was characterized by a transition from recumbent rest to upright rest. Foals were more likely to be resting, either recumbent or upright, if their mother was resting upright. During the late spring, summer, and early autumn, mares and foals were most likely to be resting upright between 09:00 and 17:00 h.     

Bioenergetics of reproduction and pup development in a subterranean rodent (Ctenomys talarum)

The objective of this study was to evaluate the maternal costs of reproduction and pup development in the subterranean rodent Ctenomys talarum (Thomas 1898). Statistical differences were detected in whole-animal metabolic rates between nonreproductive and pregnant or lactating females. Whole-animal metabolic rates during pregnancy and lactation were 128% and 151% of the resting metabolic rate (RMR) observed in nonreproductive females. The total additional energy cost of reproduction (above the nonreproductive level) was similar for both the gestation and lactation periods. Mass-specific RMR revealed an upregulation of cell or tissue metabolism during lactation but not during gestation. The mass-specific metabolic rate of pups was 237% of the adults' metabolic rates. No differences were observed in body temperature among nonreproductive, pregnant, or lactating females. No differences were detected in body mass at birth among pups from litters with different numbers of nestlings. Pups increased their body temperature, reaching adult temperature at 30 d of age, when they were near weaning. Milk constituted the exclusive food for pups until they started eating solid food at 10 d old. Suckling time decreased with age of pups, and at the same time, mother chases directed toward their pups increased. These reproductive characteristics may contribute to successful existence in a subterranean habitat.     

Maternal behaviour and early behavioural ontogeny of grey seals (Halichoerus grypus) on the Isle of May, UK

Female grey seals allocated more resources to individual sons than to individual daughters. Mothers of male pups spent significantly larger proportions of their time defending and feeding their young than did mothers of female pups. The amount of time females spent ashore and the proportion of time spent in various behavioural states was influenced by the habitat of the birth site and the age of the pup.
Grey seal neonates spent the majority of their time prone and immobile, perhaps to allow maximum energy retention (maximum mass gain) and to facilitate mothers relocating their pups. The proportion of time pups spent in the various behavioural states was influenced by the whelping habitat. Pups became increasingly active with age. Males were significantly more aggressive than females and, at one site they performed nosing behaviour with their mothers significantly more often than did female neonates.     

Weaning and Parent-Offspring Conflict in the Domestic Dog

Weaning and signs of a parent-offspring conflict were studied in four females of the Swedish Dachsbracken breed of domestic dog and their pups. The animals were observed from the second to the seventh week of age of the pups. In addition to regular weighing, measurements of milk and solid food intake per pup and meal were also made, and samples of milk from the mothers were collected and analysed. The most important mechanism for weaning seemed to be on the behavioural level. The time that the mother spent with her pups decreased continuously from week 2 to week 7, as did the number of sucklings per hour. Furthermore, both duration of suckling and the number of sucklings initiated by the mother decreased during the period, while the proportion of sucklings where the mother was standing increased steadily. The weights of mothers stayed rather constant during the period and there was no difference in the amount of milk given per suckling or in the composition of milk between the early and late weeks of lactation. Consequently, costs for the mother, in terms of loss of weight, were negligible as she was able to compensate for the increased energy demand of lactation with an increased food intake. There was a tendency for care-giving behaviour to decrease and aggression from the mother to increase at the same time as there was a tendency for care-seeking and contact-seeking behaviour from the pups to increase. These changes, together with the less frequent initiation of suckling by the mother, could perhaps be seen as signs of conflict. Conflict was defined according to TRIVERS' theory (1974) and referred to the disagreement between the female and her pups about the amount of care given. However, although the animals were kept in a way that allowed them to perform as much as possible of their natural behaviour, the good nutritional conditions, one of the characteristics of captive life, may have reduced overt parent-offspring conflict.     

Absentee Mothering – Not So Absent? Responses of European Rabbit (Oryctolagus cuniculus) Mothers to Pup Distress Calls

Longer‐range acoustic parent‐offspring communication is widespread, but might be absent in species in which young are hidden in burrows during the mother's absence. The European rabbit (Oryctolagus cuniculus) is such a species, with unusually limited maternal care largely restricted to a short daily nursing bout. Based, however, on evidence of frequent infanticide in this species, we hypothesize that rabbits possess a mechanism promoting a maternal response to pup distress calls. We conducted playback experiments with distress calls of pre‐weaning pups played next to the breeding burrows of mothers in a field enclosure (i.e. next to the burrows where mothers give birth and raise their young). Calls were played shortly after pups were born (T1) when infanticide risk is maximal, and shortly before the pups start dispersing from the breeding burrow (T2). A high proportion of mothers (60.6%) responded to pup calls by rapidly returning to their breeding burrow and 40% of them investigated the area around the entrance. Return responses to the playback of pup calls did not differ between mothers during T1 and T2. Thus, our results confirm that rabbit mothers respond rapidly to pup distress calls and that this responsiveness may adaptively serve to repel potentially infanticidal females.     

Infant development in two aquarium bottlenose dolphins

The behavioral development of two aquarium bottlenose dolphin calves, one male (born to a primiparous mother) and one female (born to a multiparous mother), was examined. From birth to 9 weeks of age, a total of 558.7 h of nursing and behavioral data were collected using focal sampling methods. Data from both calves revealed a pattern of increasing independence as they aged, but there were significant differences between the two calves. Suckling rates, mean suckles per bout, suckling bout rates, and bout durations for the male calf were higher than those for the female calf throughout the study. These frequencies declined significantly over time in the male's case but not in the female's case. Her suckling durations (lock-on time) were longer than his. The male spent more time in infant position (under the mother) than the female calf and less time swimming in echelon position (alongside the mother). Over time, the male spent less time in infant position, and the female calf spent more time socializing and less time echelon swimming. Both calves decreased the amount of time spent close to their mothers over the 9 week period, and the mothers decreased their role in proximity maintenance over time. We suggest that factors related to maternal primiparity caused the male calf to develop an effective compensatory response of frequent nursing and spending more time in infant position. Both calves survived and were healthy at the end of the observation period. © 1995 Wiley-Liss, Inc.     

Effect of weaning time on the growth rate and food intake of the spiny mouse pup

The spiny mouse Acomys cahirinus produces well-developed pups although it is a relatively small mammal (45 g). We envisioned two opposing hypotheses on the effect of early weaning on the growth rate of pups. The first predicts little effect since the increase in energy intake of dams above non-reproducing females is relatively low, suggesting that pups consume a large portion of their energy as solid food, and the pups are very well developed at birth. The second predicts a substantial effect since the 'index of precociality', that is the energy intake for maintenance of a pup as a proportion of that predicted for a rodent of its body mass, falls within values for altricial rodents, suggesting an extended maternal dependence of the young. To test these hypotheses, we measured the growth rate and food intake of pups weaned after either 7, 14, 21 or 28 days. Only three of 12 pups weaned after 7 days survived and, consequently, the latter hypothesis was supported. All pups weaned at 14–28 days survived. There was a significant decrease in growth rate during the first day after weaning in pups weaned at 7, 14 and 21 days but not after 28 days, suggesting that pups did not require parental care by day 28. Peak growth rate in pups weaned at 14 days occurred in the second week but occurred in the third week in pups weaned at 21 and 28 days. In spite of these differences, pups in all treatments had similar body mass at 64 days, indicating compensatory growth. We concluded that pups of A. cahirinus are precocial from a morphological aspect in that they are well developed at birth but altricial from a nutritional aspect in that they require extended maternal support.