Abstract: Many aspects of an animal's early development might potentially contribute to long-term individual differences in physiology and behavior. Here we asked whether differences among littermates of the domestic rabbit in the position in the litter huddle that they occupy during the early nest period might contribute to the development of distinct behavioral and physiological phenotypes. In each of 12 litters we determined the pup occupying the most peripheral, the most central, and an intermediate position in the huddle during the first postnatal week. We then tested the responses of these same individuals as nestlings, juveniles and young adults when confronted by a range of age-appropriate environmental challenges. Two behavioral tests appeared particularly discriminatory in identifying differences associated with early position in the huddle; latency of pre-weaning pups to jump down from a shelf, and the response of young adults to the fearful screams of a conspecific. In both cases animals that had occupied the periphery of the huddle showed behavioral responses indicative of a more proactive behavioral style than their "intermediate" or "central" littermates. We conclude that while consistent long-term differences in behavioral style associated with early position in the litter huddle exist in rabbits, future work is needed to confirm the causal nature of this association, to identify underlying mechanisms, and to refine methods of behavioral and physiological testing across the life span.
Abstract: Although most mammals grow up in the company of same or different age sibs (or half sibs), surprisingly little attention has been given to how relations among them might influence the development of individual differences in morphology, physiology, and behavior. Here we review evidence from our work on domestic and wild European rabbits, and more recently on laboratory rats, mice, and domestic cats, supporting the proposition that in mammals early sibling relations contribute to the development of individual differences in these three domains and thereby to long-term behavioral differences of the kind we might consider part of an animal's behavioral style or personality. First we report a consistent and marked negative relation between litter size and individuals' body mass at birth and weaning, as well as marked within-litter differences in prenatal body mass and placental efficiency. We then report individual differences in preweaning behaviors associated with these morphological variables such as position occupied in the litter huddle and development of motor ability, as well as physiological differences in thermoregulation, immune parameters, and endocrine indicators of stress. Finally, we report first evidence from wild rabbits that early relations among littermates may have long-term consequences for individual differences in behavioral style. We conclude that in mammals, individual differences in early growth, physiology and behavior potentially important for the development of animal personality, are shaped to an appreciable extent by early sibling relations and that this little-researched field deserves closer attention.
Abstract: Interest has been growing in the influence siblings may have on individual development. While mammalian research has tended to emphasize competition among siblings for essential but often limited resources such as the mother's milk, there is also evidence of mutual benefits to be had from sibling presence, most notably for altricial young in enhanced thermoregulatory efficiency. In the present study we asked whether littermates of an altricial mammal, the domestic rabbit, might gain other developmental benefits from sibling presence. From postnatal days 1 to 25 we raised rabbit pups either together with their littermates or alone except for the brief, once daily nursing characteristic of this species, while controlling for litter size and ambient nest box temperature. At weaning on Day 25 the young were then transferred to individual cages. Before weaning, we found that pups raised separately from their littermates obtained less milk, and showed lower weight gain and slower development of the ability to maintain body equilibrium than their litter-raised sibs. This was the case even though the two groups did not differ in birth weight or in the ratio of converting milk into body mass in their temperature-controlled nest boxes. Postweaning, the isolation-raised animals were also less successful in competing for food and water when tested after deprivation than their litter-raised sibs. The present study adds to the growing evidence of the influence, in this case positive, that sibs (or half sibs) may have in shaping one another's development.
Abstract: Early postnatal growth in mammals can be considerably influenced by litter size and often differs among littermates in relation to birth mass. In a study of Long Evans laboratory rats we asked whether within- and between-litter differences in body mass and growth are related to behavioral development during early postnatal life. For this, we analyzed the amount of general motor activity and the display of directed, seemingly goal-oriented interactions within the litter huddle in previsual pups. During the study period from postnatal days 2 to 11, we found significant changes in pup behavior, showing a nonlinear, quadratic shape. General motor activity and, more specifically, the display of behaviors apparently directed to reaching central positions in the litter huddle increased during the first postnatal days and then decreased again. However, pups from small litters that grow more rapidly than pups from large litters, showed a faster increase in both behaviors, whereas the young from large litters reached a higher maximum. We also found striking within-litter differences in the amount of directed behavior performed by light and heavy pups, with higher levels in the former group, most probably because light pups that have a less favorable body mass-to-volume ratio and more often occupy peripheral positions in the litter huddle, make a greater effort to reach thermally favorable central positions. In conclusion, our study shows there to be consistent between-litter as well as within-litter differences in behavioral patterns during early life. These differences might have important implications for an individual's long-term behavioral and physiological performance.
Abstract: We studied the development of suckling behavior and weight gain in 11 litters (52 kittens) of free-ranging domestic cats until postnatal day 28 just before the start of weaning. In six of these litters, we also recorded milk intake and contests for access to nipples. Already within 12 hr of birth kittens showed a preference for posterior nipples, and by postnatal day 3 each had developed a preference for particular nipples. In fact, 86% of kittens used one particular nipple most often, and even when the mother changed the side she lay on to nurse. Contests for access to nipples occurred throughout the study period at an average rate of one to two contests per kitten per hour of nursing. Contrary to suggestions in the literature that kittens compete for more productive nipples, we found no relation between kittens' use of particular nipples and their weight gain, milk intake, or involvement in contests during suckling. We suggest that kittens' preference for posterior nipples as well as their establishment of an individual "teat order" might function to optimize the number of nipples remaining productive across lactation, and to reduce energetically costly scrambles and potentially injurious contests among littermates.
Abstract: Interest is growing among psychobiologists and behavioral ecologists in the role of sibling relations in shaping individual development and life histories. In litters of domestic rabbits Oryctolagus cuniculus the heaviest pups at birth are more likely to survive the critical first postnatal week, they compete more effectively with littermates for milk and well-insulated positions in the litter huddle, and are the heaviest at weaning. Here we report that high birth weight pups are also better able to maintain body equilibrium. Testing pups' ability to maintain equilibrium when placed on a 15 degrees ramp for 2 min each day during the first postnatal week, we found that pups showed a continual daily improvement in their ability to maintain balance while moving on the ramp, rarely lost balance by postnatal day 8, and that heavier pups could maintain balance better and earlier than their lighter littermates. Better ability to maintain body equilibrium, however achieved, may help explain heavier pups' advantage in competing for vital resources such as milk and in gaining access to better-insulated positions in the litter huddle. It also provides further support for the usefulness of birth weight, not only as an absolute measure but also relative to the weight of other littermates, as a predictor of different developmental trajectories, behavioral and physiological, among same-age siblings in this mammal.
Abstract: The European rabbit, both in its wild and domesticated forms, has been a pioneer species in the study of mammalian chemical communication, and illustrates well the difficulty of understanding the functional significance of these often complex signals. Here we investigate the performance of one of the rabbit's most conspicuous chemical signaling behaviors, chin marking (chinning), and the hypothesis that this expresses social dominance. In tests of 21 chinchilla-strain sexually mature males we predicted 1) that animals would show marked and stable individual differences in the frequency of chinning, 2) that these differences would correlate with behaviors associated with dominance such as intrasexual mounting, and 3) that individual differences in the frequency of chinning and dominance-related behaviors would correlate with individual differences in a commonly used physiological indicator of dominance, concentration of serum testosterone. Supporting these predictions and consistent with previous reports, animals showed large and stable individual differences in the frequency of chinning which correlated with the behavioral indicators of dominance and less strongly, with serum testosterone. As our animals had been kept in single cages and without direct contact with other males since weaning, these findings raise the question as to how and when during development such differences among individuals arise. We are currently investigating the possible relation between pups' intrauterine position, postnatal competition among littermates for milk and thermally advantageous positions in the litter huddle, and later differences in indicators of dominance such as those reported here.
Abstract: Birth mass can vary considerably among mammalian littermates. Heavier pups often show higher growth rates than their lighter siblings, which might positively affect fitness-relevant parameters during later life. Such a correlation between birth mass and pre-weaning growth within litters was confirmed by our study of wild-type and domestic European rabbits (Oryctolagus cuniculus) living in a semi-natural environment and under laboratory housing conditions, respectively. Our study indicates that at least two main mechanisms account for this relationship in our study species: heavier pups had a higher milk intake and also showed a more efficient conversion of milk into body mass. Furthermore, our study suggests that the better milk conversion by heavy pups was driven by three synergistic mechanisms: heavier pups had comparatively more huddling partners in the nest, they did not need to perform large amounts of proactive behavior in order to reach and remain in a central position within the litter huddle, and they could maintain a comparatively higher body temperature most probably due to their more favorable surface area to volume ratio. In conclusion, our study of European rabbits provides strong evidence that both under natural conditions and in the laboratory, within-litter differences in birth mass are maintained and may even increase during pups' early postnatal development.
Abstract: Newborn rabbits compete vigorously for the mother's milk, and survivors benefit from littermate deaths. Here we report that rabbit pups also may benefit from littermate presence in terms of increased thermal efficiency. Pups nursed normally once a day by their mother but otherwise raised alone had a lower probability of survival, lower body temperature, and lower efficiency of converting milk into body mass than their siblings raised with littermates. The contribution of a more favorable thermal environment to the better growth and survival of group-raised pups was supported by the finding that single pups raised at higher ambient temperatures grew more rapidly than single pups raised at lower temperatures. These effects were most clearly seen across Days 2 to 5, after which time differences between treatment groups were no longer significant. Thus, the results of this study suggest that the benefits as well as costs of having siblings must be weighed against each other when considering the manner in which sibling presence influences individual development and survival.
