Heiko G. Rödel

Behavioural and physiological responses of naïve European rabbits to predator odour

Animals show a variety of antipredator strategies in response to the presence of chemical cues from mammalian predators. Nevertheless, there is no general agreement as to whether recognition of predator odours is dependent upon experience. We conducted an experiment on European rabbits Oryctolagus cuniculus naive to any contact with predators and we investigated (1) whether they possessed a mechanism for the recognition of the odour of a predator, and (2) how they responded behaviourally and physiologically to that odour. We used fox Vulpes vulpes faeces as the source of the predator odour and sheep Ovis aries faeces as a nonpredator control odour. The experiments were conducted in small outdoor enclosures where the animals were kept singly. We recorded patterns of vigilance, activity and feeding, and changes in glucocorticoids and body mass. The rabbits showed a clear antipredator response to the presence of fox faeces, whereas they behaved neutrally in response to sheep odour. The response consisted of increased avoidance and vigilance while feeding and more investigation before feeding. Furthermore, the rabbits showed a physiological alarm response, that is, an increased responsiveness of their adrenocortical system and weight loss. However, the total activity budget, measured as time spent outside the burrow, the time spent feeding, and the amount of food ingested remained largely stable during the experiment. We conclude that rabbits recognised predator odours and that this recognition was independent of experience.

Female home range size is regulated by resource distribution and intraspecific competition: a long-term field study

The size of an individuals home range is an important feature, influencing reproduction and survival, but it can vary considerably among both populations and individuals. The factors accounting for such variation are still poorly understood, and comprehensive long-term field studies considering various environmental factors that influence home range size are rare. We investigated the effects of seasonality, availability of food, cover, number of direct neighbours and the relative individual body mass on home range sizes in 125 adult female striped mice, Rhabdomys pumilio, in South Africa from 2004 to 2008. We used radiotelemetry to estimate home range sizes, trapping to determine the number of direct neighbours, and plant surveys in every home range to determine availability of food and cover. Home ranges were smaller when food quantity was high, many territorial neighbours were present, females had a relatively small body mass and during the nonbreeding season. We conclude that the availability of food resources and intraspecific competition are the main factors influencing home range size in female striped mice. Females enlarged their home ranges when territorial neighbours were few, and there was a significant positive correlation between home range size and quantity of food plants. This indicates that home range size might not reflect the minimal trade-off between access to resources that allow for a females survival and lowest cost for defending and foraging in that area. Instead, we propose a hypothesis for future research that female striped mice occupy areas several times larger than needed to improve their fitness by providing resources for future offspring.

Non-invasive measurement of the physiological stress response of wild rabbits to the odour of a predator

Summary.Stress has been widely studied in different mammals, but the physiological stress reaction that the odour of a predator could induce in preys has not received much attention. Besides, not all the animals would respond to the same extent to a known stressor.We developed an experimental procedure with eleven naïve European rabbits (Oryctolagus cuniculus) in order to determine the individual physiological response to the olfactory detection of a potential predator. The rabbits were housed singly in small enclosures with a concrete burrow system and food and water were available ad libitum. The animals followed a control trial, without odour, and an experimental trial where we confronted the rabbits with fox (Vulpes vulpes) odour. Furthermore, another sample of eleven rabbits followed a control procedure subjected to the same housing and handling procedures but without facing the predator odour. In order to assess the physiological response we analysed the concentration of glucocorticoid metabolites in the faeces of the rabbits. Therefore, everyday faecal samples were collected and analysed with an enzyme immunoassay in order to measure the corticosterone metabolites (CM), particularly, those metabolites with a 5α-3β, 11β-diol structure.After validating the assay for wild rabbits, we found that the simulated presence of a predator (fox odour) in the enclosure resulted in an increase in faecal CM concentrations. However, the stressor did not affect all the animals in the same way. We found a general increase in the individual differences. In particular, males experienced a higher increase than females, though the overall response was similar for both sexes.To our knowledge this is one of the first attempts to analyse the assessment of the risk of predation by means of non-invasive methods.

Early development influences ontogeny of personality types in young laboratory rats

Features of an individuals early development are frequently reported to alter the postnatal ontogeny in litter-bearing mammals with respect to various physiological parameters. We hypothesized that such effects might also apply to the ontogeny of personality types. On the one hand, litter size effects by means of more contacts with littermates, might lead to the development of more offensive types. On the other hand, smaller and less developed young from larger litters might be less offensive due to their lower physical capabilities to deal with challenging situations. We studied these contrasting hypotheses in young rats, which we tested in a battery of emotionality tests. There were clear indications for the existence of distinct behavioral types by means of consistencies in behavioral responses within and across contexts. Based on these responses, we calculated three new variables by PCA, which we interpreted to mainly reflect boldness, exploration, and anxiety. Overall, our results strongly suggest that the early development alters the ontogeny of personality types, with heavier individuals being bolder and more explorative. Furthermore, body mass and litter size influenced the changes in the behavioral responses in successive tests, further supporting the importance of the litter size-dependent body mass for the ontogeny of personalities. Anxiety also depended on litter size, however, in a nonlinear way. Animals born to litters of small or large sizes had higher scores, whereas individuals from medium-sized litters were less anxious. This optimum curve indicates that opposing effects of litter size are involved in shaping personalities in young rats.

Development of behavior in the litter huddle in rat pups: Within- and between-litter differences

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 individuals long-term behavioral and physiological performance.

Winter browsing of brown hares: evidence for diet breadth expansion

Summary Foraging theory predicts that diet breadth should expand as food availability decreases. We tested this by looking at the winter browsing behaviour of the brown hare Lepus europaeus. We predicted selective feeding on different woody plant species, diet generalisation under increasingly severe winter conditions and a rank preference between the different food items. Following a period of severe winter conditions, we censused browsing marks of brown hares on woody plants at six sites situated at different altitudes (340 – 600 m a. s. l.) in Upper Franconia, Germany. We assumed that access to ground vegetation, which is the general diet of brown hares, was more restricted at sites with higher snow cover. The results provided support for all of our predictions: Feeding intensity varied strongly between the different plant species indicating selectivity of feeding. The number and also the percentage of browsed woody plant species was positively correlated with the altitude of the study site indicating a wider diet breadth in situations where less food was available. On the basis of a rank model on the food choice of brown hares, which was evaluated by an independent data set, we conclude that brown hares show a rank preference with regard to different groups of woody plant species. Our findings support the assumption that food restricted hares increasingly include unfavourable, low quality items into the diet.

Major histocompatibility complex variation and age‐specific endoparasite load in subadult European rabbits

Genes of the major histocompatibility complex (MHC) play a fundamental role in the vertebrate immune response and are amongst the most polymorphic genes in vertebrate genomes. It is generally agreed that the highly polymorphic nature of the MHC is maintained through host–parasite co‐evolution. Two nonexclusive mechanisms of selection are supposed to act on MHC genes: superiority of MHC heterozygous individuals (overdominance) and an advantage for rare MHC alleles. However, the precise mechanisms and their relative importance are still unknown. Here, we examined MHC dependent parasite load in European rabbits (Oryctolagus cuniculus) from a distinct population with low MHC diversity (three alleles, six genotypes). Using a multivariate approach, we tested for associations of individual MHC class II DRB constitution and the rabbits’ intestinal burden with nematodes and coccidia. Rabbits having a particular allele showed lower infestations with hepatic coccidia (E. stiedai). However, a comparison of all six genotypes in the population revealed that carriers of this allele only benefit when they are heterozygous, and furthermore, MHC heterozygosity in general did not affect individual parasite load. In conclusion, this study suggests an immunogenetic basis of European rabbit resistance to hepatic coccidiosis, which can strongly limit survival to maturity in this species. Our study gives a complex picture of MHC–parasite correlations, unveiling the limits of the classical hypotheses of how MHC polymorphism is maintained in natural systems.

The responsiveness of young rabbits to the mammary pheromone: developmental course in domestic and wild pups

Abstract.Newborn rabbits rely on odour cues to localize and seize maternal nipples. In particular, they orally respond to the mammary pheromone (MP) emitted in rabbit milk. The activity of the MP was so far mainly investigated in newborns, and a drop in its releasing effect was shown in 23-day old pups. Here, we evaluated the activity of the MP in a large sample (n=1588) of domestic rabbit pups tested between birth and weaning. We also assessed whether wild-type pups respond to the MP, and whether the developmental course of their responsiveness matches the one of domestic pups. During the first postnatal week, the MP revealed to be a strong releasing stimulus in all pups, regardless of their genetic and environmental background. However, the saliency of the MP decreased with approaching weaning, and this decrease was clearly steeper in wild-type than in domestic pups. These results are discussed with regard to the concept of ontogenetic adaptation and the effects of artificial selection.

Survival costs of fast exploration during juvenile life in a small mammal

Animal personality is considered to be subject to natural selection, and measuring its fitness consequences is an important step in the study of the evolution of this phenomenon. The evolutionary costs and benefits of certain personality traits are frequently assumed to fluctuate across different life history stages. However, little is still known about survival consequences of personality during juvenile life. We predicted survival costs of exploration tendency in juvenile European rabbits (Oryctolagus cuniculus), which we studied under quasi-natural conditions. In addition, consistencies between exploration and the animal’s behavioral responses in other contexts were tested. Early exploratory behavior was assessed in two annual cohorts, either observationally by quantifying the time course of exploring the area around the nursery burrow shortly after emergence above ground or experimentally by conducting open-field tests shortly before emergence. In both years, the study revealed consistent results: more exploratory individuals had a lower chance of survival until the end of the vegetation period, presumably due to a higher predation risk. Fast explorers, as assessed by their exploration of the burrow environment, were also less sociable and tended to be more aggressive toward conspecifics as subadults. Sociability was repeatable across time. Furthermore, fast explorers, as assessed by their open-field behavior, were bolder in a handling test before emergence. These consistencies across context and time suggest the existence of personality. In conclusion, the results provide evidence for survival costs of fast exploration during juvenile life. The findings are in line with predictions on personality-related life history variation.

Family matters: maternal and litter-size effects on immune parameters in young laboratory rats.

A functional immune system is important for the survival of mammalian young, particularly at weaning when they lose the immunological support provided by the mothers colostrum and milk. In altricial mammals, litter size and maternal characteristics are important components of an animals early environment, which affect postnatal growth and development. In a study of unculled litters of Long-Evans laboratory rats (Rattus norvegicus), we asked whether such parameters are also associated with the immune status of the young shortly before weaning. On postnatal day 17, we assessed numbers of several leukocyte and lymphocyte subsets, the activity of the complement system, and immunoglobulin G (IgG) concentrations in the serum. Averaging the values of all pups per litter, we found negative correlations between litter size and lymphocyte counts, complement system activity and IgG concentration. Maternal effects were seen in the positive correlation between maternal postpartum body mass and granulocyte and monocyte counts. In addition, lymphocyte and monocyte counts as well as complement activity were lower for the young of multiparous than of primiparous mothers. This suggests a trend towards a better developed immune system in such offspring, which may be relevant for their immediate and long-term survival. The effects described here have potential implications for the design and interpretation of biomedical studies of immune parameters in laboratory rats.