Age-specific effects of density and weather on body condition and birth rates in a large herbivore, the Przewalski's horse.

Reproduction in young females can show a particularly sensitive response to environmental challenges, although empirical support from individual-based long-term studies is scarce. Based on a 20-year data set from a free-roaming Przewalski's horse population (Equus ferus przewalskii), we studied effects of large-herbivore density (horses + cattle) and weather conditions experienced during different life stages on females' annual birth rates. Foaling probability was very low in 2-year-olds, reaching maximum values in 5 to 10-year-olds, followed by a decrease in older females indicating reproductive senescence. Mother's previous reproductive investment affected her current reproduction; young and old mothers (as opposed to middle-aged ones), which had nursed a foal for at least 60 days during the previous year, reproduced with a lower probability. Foaling probability and body condition of young females were lower when large-herbivore density was high. Reproduction was also influenced by interactive weather effects during different life stages. Low late-summer precipitation during the females' year of birth was associated with a pronounced decrease in foaling probability in response to harsh late-winter temperatures prior to the mating season. In turn, increased amounts of late-summer rain during this early age together with more late-summer rain during the females' current pregnancy led to an increased reproductive probability in 2-3-year-olds. These results were corroborated by the ameliorating effects of late-summer rain on body condition in such females. In conclusion, our findings highlight the interactive importance of weather conditions experienced during early life, and of density and weather during current pregnancy on foaling probability, particularly in young females.

Patterns and predictors of inter-litter differences in rabbit pup locomotor activity, based on an automatized quantification method.

Individual-level sibling interactions in the litter huddle have been studied extensively, especially in the domestic rabbit (Oryctolagus cuniculus). However, little is known about inter-litter differences in pup activity patterns during early postnatal life, in particular regarding the drivers of such variation. In our study on 2-3-day-old rabbit pups, we predicted lower locomotor activity in litters with lower mean body masses on the day of birth (starting body mass) and with lower daily milk intake per pup, possibly constituting a behavioral strategy of pups to cope with associated energetic constraints. For an automatized assessment of pup locomotor activity in the litter huddle, we successfully developed and validated a method based on the quantification of dissimilarities between consecutive frames of video footage. Using this method, we could confirm a U-shaped time course of litter-level locomotor activity, with maximum values shortly before and after the once-daily nursing typical for the rabbit. As predicted, between-litter variation in mean starting body mass and in daily milk intake affected the degree of locomotor activity in the litter huddle, in an interactive way. That is, in litters with heavier starting body masses, pup locomotor activity was greater in pups with an initially higher milk intake, suggesting that only pups with better body condition and a higher energy intake could afford higher levels of activity. This interaction was exclusively apparent during the middle phase of the 24 h inter-nursing interval, when litter activity was low. Shortly before nursing, when pups show higher levels of locomotor behavior in anticipation of the mother's arrival, and shortly after nursing when the pups were more active possibly due to adjustments of their positions in the huddle, activity levels were decoupled from pups' starting body mass and previous milk intake. Our findings highlight the importance of pup body mass and daily energy intake, two parameters known to be related to maternal characteristics, in shaping inter-litter differences in pup locomotor activity.

Early life parameters and personality affect oxidative status during adulthood in an altricial rodent.

It is increasingly recognized that alterations of the cellular oxidative status might be an important cost underlying challenging early life conditions. For example, an increased litter size can impose challenges as the offspring will face increased competition for maternal resources. Within a litter, individuals with relatively higher starting mass typically show higher growth rates, which can lead to increased oxidative damage. We investigated the long-term consequences of these early life parameters on the oxidative status in mature mound-building mice (Mus spicilegus). Individual differences in the animals' exploration tendency were assessed by repeated open field and novel object tests. We predicted less exploratory phenotypes, which typically show a higher stress responsiveness, to be particularly susceptible to possible effects of these early life parameters on oxidative status. We quantified oxidative damage of DNA (8-hydroxy-2'-deoxyguanosine levels, 8-OHdG) and proteins (protein carbonyl content, PCC), and activities of the antioxidants catalase (CAT), glutathione peroxidase (GPx), and superoxide dismutase (SOD) in liver and skeletal muscle tissue. 8-OHdG levels were positively associated with CAT and SOD in both tissues, indicating that increased oxidative DNA damage was associated with an upregulation of antioxidant production. Hepatic DNA damage after maturity was increased in animals from larger litters. In less exploratory animals, DNA damage and the activity of CAT and SOD in the muscle were increased, but only in individuals with higher relative starting mass (measured on postnatal day 9). This interaction may be explained by the typically higher adrenocortical activity in less exploratory phenotypes and by the higher growth in relatively heavier pups, two factors known to increase oxidative stress. These findings contribute to enlightening the complex interplay between early life conditions, personality, and oxidative status.

Is there a bias in spatial maze judgment bias tests? Individual differences in subjects' novelty response can affect test results.

Judgment bias tests have become an important tool in the assessment of animals' affective states. Subjects are first trained to discriminate between two cues associated with a positive and a less-positive outcome. After successful training, they are confronted with an ambiguous cue, and responses are used for judgment bias assessment. In spatial settings, ambiguous cue presentation is typically linked with novelty, i.e. to yet unexplored areas or areas to which the animal has a low degree of habituation. We hypothesized that in such settings, responses to ambiguity might be biased by the animals' perception of novelty. We conducted judgment bias tests in mound-building mice phenotyped for their exploration tendency. After subjects had learned to distinguish between the positively and less-positively rewarded arms of a maze, a new ambiguous middle-arm was introduced. During the first test trial, more exploratory, less neophobic individuals displayed higher bidirectional locomotion in the ambiguous arm, indicating intensive exploration. Although this resulted in longer latencies to the reward in more exploratory animals, we conclude that this did not reflect a 'more pessimistic judgment of ambiguity'. Indeed, during the following two trials, with increasing habituation to the ambiguous arm, the direction of the association was inversed compared to the first trial, as more exploratory individuals showed relatively shorter approach latencies. We suggest that in spatial test settings associating the ambiguous cue to novel areas, results can be confounded by subjects' personality-dependent motivational conflict between exploration and reaching the reward. Findings obtained under such conditions should be interpreted with care.

More exploratory house mice judge an ambiguous situation more negatively.

Exploration tendency, one of the most investigated animal personality traits, may be driven by either positive (when seeking interesting information) or negative (to reduce the uncertainty of the environment) affective/emotional profiles. To disentangle the valence of the affective state associated with exploration trait, we applied a judgment bias test to evaluate the animals' responses in an ambiguous situation, allowing an assessment of their affective state or mood. Experiments were carried out in male house mice (Mus musculus) of wild origin. Individual differences in exploration tendency were assessed by repeated open field and novel object tests. To evaluate the animals' judgment bias, we trained the subjects for 8 days in a 3-arm maze to discriminate between two extreme locations (outer arms: either positively reinforced with sugary water or less-positively reinforced with plain water), in terms of a shorter latency to approach the positively reinforced arm. After this learning criterion was reached, we repeatedly tested their responses to an ambiguous location (intermediate arm). The latencies to approach and consume the ambiguous reward were highly repeatable over the 3 days of testing; hence individuals expressed a stable judgment bias. Most importantly, more exploratory animals showed a more negative judgment bias, which supports the hypothesis that a higher exploration tendency was associated with a negative affective state. Further studies should investigate whether exploration in different situations might be due to distinct affective states.

Contexts of emission of non-signature whistles in bottlenose dolphins (Tursiops truncatus) under human care.

Bottlenose dolphins are social cetaceans that strongly rely on acoustic communication and signaling. The diversity of sounds emitted by the species has been structurally classified into whistles, clicks and burst-pulsed sounds. Although click sounds and individually-specific signature whistles have been largely studied, not much is known about non-signature whistles. Most studies that link behavior and whistle production conduct aerial behavioral observations and link the production of whistles to the general category of social interactions. The aim of this study was to determine if there was a correlation between the non-signature whistle production and the underwater behaviors of a group of bottlenose dolphins (Tursiops truncatus) under human care, during their free time in the absence of trainers. To do this we made audio-video recordings 15 min before and after 10 training sessions of eight dolphins in Boudewijn Seapark (Belgium). For the behavioral analysis we conducted focal follows on each individual based on six behavioral categories. For the acoustical analysis, carried out at the group level, we used the SIGID method to identify non-signature whistles (N = 661) and we classified them in six categories according to their frequency modulation. The occurrences of the six categories of whistles were highly collinear. Most importantly, non-signature whistle production was positively correlated with the time individuals spent slow swimming alone, and was negatively correlated with the time spent in affiliative body contact. This is the first analysis that links the production of non-signature whistles with particular underwater behaviors in this species.

Regulation of the rabbit's once-daily pattern of nursing: a circadian or hourglass-dependent process?

The European rabbit Oryctolagus cuniculus has an unusual pattern of nursing behavior. After giving birth in a nursery burrow (or laboratory nest box), the mother immediately leaves the young and only returns to nurse for a few minutes once approximately every 24 h. It has been assumed this schedule, like a variety of other functions in the rabbit, is under circadian control. This assumption has been largely based on findings from mothers only permitted restricted access to their young once every 24 h. However, in nature and in the laboratory, mothers with free access to young show nursing visits with a periodicity shorter than 24 h, that does not correspond to other behavioral and physiological rhythms entrained to the prevailing 24 h light/dark (LD) cycle. To investigate how this unusual, apparently non-circadian pattern might be regulated, we conducted two experiments using female Dutch-belted rabbits housed individually in cages designed to automatically register feeding activity and nest box visits. In Experiment 1 we recorded the behavior of 17 mothers with free access to their young under five different LD cycles with long photo and short scotoperiods, spanning the limits of entrainment of the rabbit's circadian system. Whereas feeding rhythms were entrained by LD cycles within the rabbit's circadian range of entrainment, nursing visits showed a consistently shorter periodicity regardless of the LD regimen, largely independent of the circadian system. In Experiment 2 we tested further 12 mothers under more conventional LD 16:8 cycles but "trained" by having access to the nest box restricted to 1 h at the same time each day for the first 7 d of nursing. Mothers were then allowed free access either when their young were left in the box (n = 6), or when the litter had been permanently removed (n = 6). Mothers with pups still present returned to nurse them on the following days according to a similarly advancing pattern to the mothers of Experiment 1 despite the previous 7 d of "training" to an experimentally enforced 24 h nursing schedule as commonly used in previous studies of rabbit maternal behavior. Mothers whose pups had been removed entered the box repeatedly several times on the first day of unrestricted access, but on subsequent days did so only rarely, and at times of day apparently unrelated to the previously scheduled access. We conclude that the pattern of the rabbit's once-daily nursing visits has a periodicity largely independent of the circadian system, and that this is reset at each nursing. When nursing fails to occur nest box visits cease abruptly, with mothers making few or no subsequent visits. Together, these findings suggest that the rabbit's once-daily pattern of nursing is regulated by an hourglass-type process with a period less than 24 h that is reset at each nursing, rather than by a circadian oscillator. Such a mechanism might be particularly adaptive for rhythms of short duration that should end abruptly with a sudden change in context such as death or weaning of the young.

Within-litter covariance of allele-specific MHC heterozygosity, coccidian endoparasite load and growth is modulated by sibling differences in starting mass.

Although littermates in altricial mammals usually experience highly similar environmental conditions during early life, considerable differences in growth and health can emerge among them. In a study on subadults of a European rabbit (Oryctolagus cuniculus) population with low MHC polymorphism, we tested whether litter-sibling differences in endoparasitic coccidia load and body mass at the end of the vegetation period were associated with within-litter differences in starting body mass (measured around 2 weeks prior to weaning) and in immune-genetic (MHC class II DRB) constitution. We hypothesized that siblings with a lighter starting mass might be more susceptible to endoparasite infections and thus, negative effects of a more unfavourable MHC constitution might be particularly pronounced in such individuals. Within-litter comparisons revealed that animals with a lighter starting mass reached a relatively lower body mass in autumn. Furthermore, there were indications for an allele-specific heterozygote advantage, as animals with heterozygous combinations of the allele Orcu-DRB*4 had relatively lower hepatic coccidia loads than their littermates with certain homozygous allele combinations. Consistent with our hypothesis, significantly higher hepatic coccidia loads and tendentially lower autumn body masses in homozygous compared to heterozygous individuals for the allele Orcu-DRB*4 were evident in initially lighter but not in heavier siblings, suggesting synergistic effects between an unfavourable MHC constitution and a light starting mass. Taken together, these effects might lead to notable differences in fitness among litter siblings, as a low body mass and a high endoparasite burden are key factors limiting young rabbits' survival during winter.

Dolphins' Willingness to Participate (WtP) in Positive Reinforcement Training as a Potential Welfare Indicator, Where WtP Predicts Early Changes in Health Status.

Welfare science has built its foundations on veterinary medicine and thus measures of health. Since bottlenose dolphins (Tursiops truncatus) tend to mask symptoms of poor health, management in captivity would benefit from advanced understanding on the links between health and behavioural parameters, and few studies exist on the topic. In this study, four representative behavioural and health measures were chosen: health status (as qualified by veterinarians), percentage of daily food eaten, occurrences of new rake marks (proxy measure of social activity), and willingness to participate (WtP) in Positive Reinforcement Training sessions as qualitatively measured by their caretakers. These data were collected multiple times a day, every day over the course of a year from a multi-facility, large sample size (n dolphins = 51), allowing powerful analyses of the relationships between measures. First, it was found that dolphins with a higher WtP score also had a significantly better health status, ate a higher percentage of their daily food, and a lower occurrence of new rake marks. In addition, the WtP score was significantly lower up to 3 days before the weekly veterinary diagnosis of a decrease in health state; the percentage of daily food eaten and new rake mark measures did not show any significant change before such a diagnosis. These results suggest that WtP in training sessions is a potential behavioural measure of dolphin welfare, and an indicator of early changes in the dolphins' health state. We therefore suggest measurement of WtP as a more practical and non-invasive tool to support veterinary care and general management. More work needs to be conducted to elucidate the influence of social behaviour on health, and to identify other potential welfare indicators, but this long-term study has shown that qualitative measures can be both practical and valid when assessing dolphin welfare.

Differences between fast and slow explorers in short-term tail temperature responses to handling in a rodent of wild origin.

Animals of different behavioral types typically show associated differences in their physiological stress response, including differential reactivity of the sympathetic nervous system. Infrared thermography offers the possibility to explore this link in a non-invasive way via the quantification of fine-scale changes in peripheral body temperature due to changes in cutaneous blood flow. We used this technique to investigate the association between exploration tendency, a behavioral trait frequently used to phenotype mammals and birds, and short-term thermal responses to challenge in a small rodent of wild origin, the mound-building mouse (Mus spicilegus). We applied a brief handling procedure consisting in the transfer of subjects into a small arena. This procedure led to a significant increase in subjects' maximum peripheral body temperature (mainly reflecting the temperature of the eyes) and to significant decreases in maximum temperatures at different positions on the tail. Maximum peripheral body and tail temperatures showed significant individual-level consistencies in response to repeated applications of the handling procedure, suggesting stable individual differences in the animals' sympathetic activity. We then compared the thermal responses to handling between 'fast' and 'slow' explorers, who were phenotyped through repeated open field and novel object tests. Fast explorers showed significantly lower tail temperatures than slow explorers shortly after handling, suggesting a stronger sympathetic reactivity in the former. Comparisons within sibling groups kept in different cages showed that the differences between explorer types were particularly pronounced during the first minute after handling, and increased in magnitude along the first millimeters distal to the tail base.

Isolation calls in house mouse pups: Individual consistency across time and situations.

Isolation calls are emitted by the offspring of many mammalian species when separated from caregivers and siblings. Some studies indicate that isolation call rates constitute a consistent individual trait; others show that the young adjust their vocalization rate to the current situation. We studied this in the house mouse (Mus musculus domesticus) by exploring individual consistencies in pup isolation call rates and their potential modulation in different social situations. We carried out experiments including three treatments (repeated measurements) during consecutive days, all starting with an initial isolation of the pup, followed by (a) a reunion with mother and littermates and a second isolation hereafter, (b) the confrontation of isolated pup with cues of its own nest or (c) with cues of an unfamiliar adult male. The first treatment induced a significant increase, while the others induced significant decreases in pup isolation call rates. Pups showed consistent individual differences in initial call rates across the three days of testing (postnatal days 9-11), which were significantly associated with individual differences in call rates during the different treatments. We conclude that pup isolation calls represent a consistent, trait-like behaviour in the house mouse, which can also express flexibility in response to social cues.

Individual behavioural type and group performance in Formica fusca ants.

The link between individual and group-level behaviour may help understanding cooperation and division of labour in social animals. Despite the recent surge of studies, especially in social insects, the way individual differences translate into group performance remains debated. One hypothesis is that groups may simply differ in the average personality of their members and this would translate into inter-group differences in collective behaviour. We tested the hypothesis of a linear relationship between individual and group phenotype in the ant Formica fusca by using same-age groups of workers after measuring an individual behavioural trait. Individual exploratory activity in an open-field arena was significantly repeatable. Based on this trait, groups were composed, each consisting of 6 individuals with similar exploration tendency housed with 3 cocoons and a refuge. Individual exploratory activity was associated with the performance in cocoon recovery at the group level: groups composed of high exploratory individuals started transporting displaced cocoons significantly earlier and transported more cocoons into the refuge than groups with low exploratory workers. When in a group, more exploratory animals showed significantly more returns to the refuge than less exploratory ones and tended to transport more cocoons. These results show a direct linear link between individual and collective behaviour, suggesting that colony personality reflects the average personality of workers involved in a given task.

Individual differences in early body mass affect thermogenic performance and sibling interactions in litter huddles of the house mouse.

We asked whether within-litter differences in early body mass are associated with differences in house mouse pups' thermogenic performance and whether such variation predicts individual differences in competitive interactions for thermally more advantageous positions in the huddle. We explored pups' thermogenic performance in isolation by measuring changes in (maximal) peripheral body temperatures during a 5-min thermal challenge using infrared thermography. Changes in peripheral body temperature were significantly explained by individual differences in body mass within a litter; relatively lighter individuals showed an overall quicker temperature decrease leading to lower body temperatures toward the end of the thermal challenge compared to heavier littermates. Within the litter huddle, relatively lighter pups with a lower thermogenic performance showed consistently more rooting and climbing behavior, apparently to reach the thermally advantageous center of the huddle. This suggests that within-litter variation in starting body mass affects the pups' thermal and behavioral responses to environmental challenges.

Body mass modulates huddling dynamics and body temperature profiles in rabbit pups.

Altricial mammals typically lack the physiological capacity to thermoregulate independently during the early postnatal period, and in litter-bearing species the young benefit strongly from huddling together with their litter siblings. Such litter huddles are highly dynamic systems, often characterized by competition for energetically favorable, central positions. In the present study, carried out in domestic rabbits Oryctolagus cuniculus, we asked whether individual differences in body mass affect changes in body temperature during changes in the position within the huddle. We predicted that pups with relatively lower body mass should be more affected by such changes arising from huddle dynamics in comparison to heavier ones. Changes in pups' maximum body surface temperature (determined by infrared thermography) were significantly affected by changes in the number of their neighbors in the litter huddle, and indeed these temperature changes largely depended on the pups' body mass relative to their litter siblings. Lighter pups showed significant increases in their maximum body surface temperature when their number of huddling partners increased by one or two siblings whereas pups with intermediate or heavier body mass did not show such significant increases in maximum body temperature when experiencing such changes. A similar pattern was found with respect to average body surface temperature. This strong link between changes in the number of huddling partners and body surface temperature in lighter pups might, on the one hand, arise from a higher vulnerability of such pups due to their less favorable body surface area-to-volume ratio. On the other hand, as lighter pups generally had fewer neighbors than heavier ones and thus typically a comparatively smaller body surface in contact with siblings, they potentially had more to gain from increasing their number of neighbors. The present findings might help to understand how individual differences in body mass within a litter lead to the emergence of individual differences in sibling interactions during early postnatal life in different species of altricial and litter-bearing mammals.

Schedule of human-controlled periods structures bottlenose dolphin (Tursiops truncatus) behavior in their free-time.

Behavioral patterns are established in response to predictable environmental cues. Animals under human care frequently experience predictable, human-controlled events each day, but very few studies have questioned exactly how behavioral patterns are affected by such activities. Bottlenose dolphins (Tursiops truncatus) maintained for public display are good models to study such patterns since they experience multiple daily human-controlled periods (e.g., shows, training for shows, medical training). Thus, we investigated the effect of training session schedule on their "free-time" behavior, studying 29 individuals within 4 groups from 3 European facilities. Our initial time budget analyses revealed that among the behaviors studied, dolphins spent the most time engaged in synchronous swimming, and within this category swam most at slow speeds and in close proximity to each other. "Slow-close" synchronous swimming peaked shortly after training sessions and was low shortly before the next session. Play behavior had significantly higher frequencies in juveniles than in adults, but the effect was only seen during the in-between session period (interval neither shortly before nor after sessions). Anticipatory behavior toward sessions was significantly higher shortly before sessions and lower afterward. We conclude that dolphin behaviors unconnected to the human-controlled periods were modulated by them: slow-close synchronous swimming and age-dependent play, which have important social dimensions and links to welfare. We discuss potential parallels to human-controlled periods in other species, including humans themselves. Our findings could be taken into account when designing welfare assessments, and aid in the provision of enrichment and maintaining effective schedules beneficial to animals themselves. (PsycINFO Database Record

Highly stable individual differences in the emission of separation calls during early development in the domestic cat.

Study of the development of individuality is often hampered by rapidly changing behavioral repertoires and the need for minimally intrusive tests. We individually tested 33 kittens from eight litters of the domestic cat in an arena for 3 min once a week for the first 3 postnatal weeks, recording the number of separation calls and the duration of locomotor activity. Kittens showed consistent and stable individual differences on both measures across and within trials. Stable individual differences in the emission of separation calls across trials emerged already within the first 10 s of testing, and in locomotor activity within the first 30 s. Furthermore, individual kittens' emission of separation calls, but not their locomotor activity, was highly stable within trials. We conclude that separation calls provide an efficient, minimally intrusive and reliable measure of individual differences in behavior during development in the cat, and possibly in other species emitting such calls.

Early development and the emergence of individual differences in behavior among littermates of wild rabbit pups.

The ontogeny of associated individual differences in behavior and physiology during early postnatal life, and in particular the emergence of such differences among litter siblings, has been hardly explored in mammals under natural conditions. We studied such within-litter differences in behavior in European rabbit pups Oryctolagus cuniculus prior to weaning, and whether and how these differences co-varied with other individual characteristics such as postnatal body temperature and early growth. The study was conducted under semi-natural conditions in a colony of rabbits of wild origin, where the young were born and developed in nursery burrows. We equipped two siblings per litter with interscapular skin temperature loggers on postnatal day 2 and recorded temperature profiles for 48h. Individual body (skin) temperatures of pups within litters were repeatable across time, indicating the existence of consistent individual differences. Such differences within litters were associated with relative differences in pre-weaning growth, revealing that relatively warmer pups showed a greater increase in body mass during the nest period. Between postnatal days 12 and 17, after the pups had reached a developmental stage of greater mobility, we carried out different behavioral tests: a handling-restraint test, an open field test and a jump-down test from a platform. Individual responses in the former two tests were associated, as those pups showing a quicker struggling response to restraint during handling also exhibited greater exploratory activity in the open field. This correlation across contexts suggests the existence of personality types in wild rabbits at an early developmental stage. Furthermore, pups' behavioral responses were strongly associated with their relative within-litter body mass at testing. Animals with a lower body mass compared to their siblings showed a relatively quicker struggle response to handling restraint and covered a relatively larger distance in the open field, suggesting greater reactivity and responsiveness of relatively lighter pups in these tests. In contrast, relatively heavier pups jumped sooner from the platform, which may have been due to their greater physical maturation. In conclusion, our study shows that individual differences in behavior and associated differences in body temperature and growth are already present during early postnatal life, although such relationships can be easily overlooked, as they predominantly emerge as relative differences among littermates.

Bottlenose dolphins engaging in more social affiliative behaviour judge ambiguous cues more optimistically.

Cognitive bias tests measure variation in emotional appraisal and are validated methods to assess animals' affective states. However, the link between social behaviours and cognitive bias has not yet been investigated. Bottlenose dolphins are a gregarious species for whom welfare research is increasing in importance, and thus are a good model to test such an association. We adapted a spatial location judgement bias test for eight captive bottlenose dolphins to investigate the link between cognitive bias and social behaviour, where we conducted behavioural observations outside of training sessions and did not experimentally induce an affective state. Subjects showed stable individual differences in cognitive biases across the three test days. Furthermore, dolphins showing more synchronous swimming, a fundamental affiliative behaviour, judged ambiguous cues significantly more optimistically. Our longer-term data showed cognitive bias and synchronous swimming frequency were significantly associated for up to two months preceding the test, but disappeared prior to that, suggesting that here cognitive bias differences were reflected by transitory affective states rather than longer-term traits. We hypothesise that the frequency of synchronous swimming may induce affective states and/or be induced by them; either way, it has strong potential as an indicator of affective state in this species and beyond.

Ambient temperature affects postnatal litter size reduction in golden hamsters.

BACKGROUND: To better understand how different ambient temperatures during lactation affect survival of young, we studied patterns of losses of pups in golden hamsters (Mesocricetus auratus) at different ambient temperatures in the laboratory, mimicking temperature conditions in natural habitats. Golden hamsters produce large litters of more than 10 young but are also known to wean fewer pups at the end of lactation than they give birth to. We wanted to know whether temperature affects litter size reductions and whether the underlying causes of pup loss were related to maternal food (gross energy) intake and reproductive performance, such as litter growth. For that, we exposed lactating females to three different ambient temperatures and investigated associations with losses of offspring between birth and weaning. RESULTS: Overall, around one third of pups per litter disappeared, obviously consumed by the mother. Such litter size reductions were greatest at 30 °C, in particular during the intermediate postnatal period around peak lactation. Furthermore, litter size reductions were generally higher in larger litters. Maternal gross energy intake was highest at 5 °C suggesting that mothers were not limited by milk production and might have been able to raise a higher number of pups until weaning. This was further supported by the fact that the daily increases in litter mass as well as in the individual pup body masses, a proxy of mother's lactational performance, were lower at higher ambient temperatures. CONCLUSIONS: We suggest that ambient temperatures around the thermoneutral zone and beyond are preventing golden hamster females from producing milk at sufficient rates. Around two thirds of the pups per litter disappeared at high temperature conditions, and their early growth rates were significantly lower than at lower ambient temperatures. It is possible that these losses are due to an intrinsic physiological limitation (imposed by heat dissipation) compromising maternal energy intake and milk production.