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A basic mechanism of domestication is the selection for fearlessness and acceptance of humans as social partners, which may affect risk-taking behavior and the ability to use humans as social support, both at the behavioural and physiological levels. We combined behavioural observations with heart rate parameters (i.e., HR and heart rate variability, HRV) in equally raised and housed wolves and dogs to assess the responses to food offered in the vicinity of a potential stressor (an unknown spinning object) with and without social support from a familiar human. Based on previous studies on neophobia in wolves and dogs, we expected dogs to be less scared of the object, approach more quickly, show less ambivalent behaviour, lower HR, and higher HRV, than wolves, especially at the presence of a human partner. However, we found that mainly age and the presence of a familiar human affected the behaviour of our subjects: older wolves and dogs were generally bolder and faster to approach the food and the familiar human's presence increased the likelihood of taking it. HR rate parameters were affected by age and the stage of the test. Wolves and dogs showed particularly high HRs at the beginning and end of the test sessions. We conclude that in our paradigm, wolves' and dogs' risk-proneness varied with age, rather than species. Additionally, the presence of a familiar human increased the motivation of both, dogs and wolves to take the food.

A basic mechanism of domestication is the selection for fearlessness and acceptance of humans as social partners, which may affect risk-taking behavior and the ability to use humans as social support, both at the behavioural and physiological levels. We combined behavioural observations with heart rate parameters (i.e., HR and heart rate variability, HRV) in equally raised and housed wolves and dogs to assess the responses to food offered in the vicinity of a potential stressor (an unknown spinning object) with and without social support from a familiar human. Based on previous studies on neophobia in wolves and dogs, we expected dogs to be less scared of the object, approach more quickly, show less ambivalent behaviour, lower HR, and higher HRV, than wolves, especially at the presence of a human partner. However, we found that mainly age and the presence of a familiar human affected the behaviour of our subjects: older wolves and dogs were generally bolder and faster to approach the food and the familiar human's presence increased the likelihood of taking it. HR rate parameters were affected by age and the stage of the test. Wolves and dogs showed particularly high HRs at the beginning and end of the test sessions. We conclude that in our paradigm, wolves' and dogs' risk-proneness varied with age, rather than species. Additionally, the presence of a familiar human increased the motivation of both, dogs and wolves to take the food. [ABSTRACT FROM AUTHOR]

Social learning is highly adaptive in transmitting essential information between individuals in many species. While several mechanisms have been observed, less is known about how much animals can remember. However, results on observational spatial memory among caching species, i.e. a form of social learning allowing individuals to remember and pilfer food caches made by others, suggest that this ability correlates with their social organization. Both wolves and their domesticated form, dogs, are social species known to make food caches, and previous studies have shown that they both can use observational spatial memory abilities to find hidden food. In order to test how much socially transmitted information wolves and dogs can remember, we tested both species in a task requiring them to find 4, 6 or 8 caches after they observed a human hiding food items, or after a control condition where they could not observe the hiding. We found that both wolves and dogs retrieved more caches and were more efficient for the first few caches if they observed the hiding than in the control condition, suggesting that they did not simply rely on scent to find the rewards. Interestingly, wolves outperformed dogs irrespective of whether the caching could be observed or not. We suggest that this result is due to a difference in motivation/persistence between wolves and dogs rather than observational spatial memory.

The reproductive season is energetically costly as revealed by elevated glucocorticoid concentrations, constrained immune functions and an increased risk of infections. Social allies and affiliative interactions may buffer physiological stress responses and thereby alleviate associated effects. In the present study, we investigated the seasonal differences of immune reactive corticosterone metabolite concentrations, endoparasite burden (nematode eggs and coccidian oocysts) and affiliative interactions in northern bald ibis (Geronticus eremita), a critically endangered bird. In total, 43 individually marked focal animals from a free-ranging colony were investigated. The analyses included a description of initiated and received affiliative interactions, pair bond status as well as seasonal patterns of hormone and endoparasite levels. During the reproductive season, droppings contained parasite eggs more often and corticosterone metabolite levels were higher as compared to the period after reproduction. The excretion rate of endoparasite products was lower in paired individuals than in unpaired ones, but paired animals exhibited higher corticosterone metabolite concentrations than unpaired individuals. Furthermore, paired individuals initiated affiliative behaviour more frequently than unpaired ones. This suggests that the reproductive season influences the excretion patterns of endoparasite products and corticosterone metabolites and that affiliative interactions between pair partners may positively affect endoparasite burden during periods of elevated glucocorticoid levels. Being embedded in a pair bond may have a positive impact on individual immune system and parasite resistance.

Food preferences may be driven by a species' ecology. Closely related species such as dogs and wolves may have evolved preferences for different foods owing to their differing foraging styles. Wolves have been shown to be more persistent in problem-solving experiments and more risk-prone in a foraging task. A possible element affecting these (and other) results is a potential wolf-dog difference in food preferences. To address this possibility, we tested similarly raised and kept dogs and wolves in two different food choice tasks, a classic two-choice task and a multiple-choice paradigm. We predicted that if dogs have adapted to a more opportunistic, scavenging foraging style, they would show a weaker preference for meat over starch rich foods (such as kibble) and be less affected by hunger than wolves. Alternatively, given the recentness of the new niche dogs have created, we predicted no substantial differences between dogs' and wolves' food preferences. We found that our subjects did not differ in their preference for meat over kibble in either paradigm. However, wolves' (but not dogs') choice patterns were affected by satiation, with wolves being less "selective" when hungry. Furthermore, when fed before testing, wolves were more selective than dogs. These differences were more noticeable in the multiple-choice paradigm than the two-choice task, suggesting that the former, novel paradigm may be more sensitive and better capable of evaluating food preferences in a diverse range of species. Overall, we found that the distinct differences in wolves' and dogs' ecology and foraging styles do not appear to have affected their food preferences and thus, differences in food preferences are unlikely to have influenced results of previous experiments demonstrating wolf-dog differences in cognitive skills.

Stress responses within dyads are modulated by interactions such as mutual emotional support and conflict. We investigated dyadic psychobiological factors influencing intra-individual cortisol variability in response to different challenging situations by testing 132 owners and their dogs in a laboratory setting. Salivary cortisol was measured and questionnaires were used to assess owner and dog personality as well as owners' social attitudes towards the dog and towards other humans. We calculated the individual coefficient of variance of cortisol (iCV = sd/mean*100) over the different test situations as a parameter representing individual variability of cortisol concentration. We hypothesized that high cortisol variability indicates efficient and adaptive coping and a balanced individual and dyadic social performance. Female owners of male dogs had lower iCV than all other owner gender-dog sex combinations (F = 14.194, p

Social play is known as a cooperative interaction between individuals involving multiple mechanisms. However, the extent to which the equality of individuals' play styles affects the interaction has not been studied in many species. Dyadic play between wolf puppies, as well as between puppies and adults, was studied to investigate both self-handicapping and offensive behaviors to determine the extent to which wolves engage in play styles where one individual does not dominate the play. Our results did not support the hypothesized '50:50' rule, which suggests that more advantaged individuals should show higher rates of self-handicapping behaviors in order to facilitate play with others. Adult wolves performed significantly less self-handicapping behaviors than their puppy partners, and they performed significantly more offensive behaviors than their puppy partners. While the '50:50' rule was not supported at any time during our study period, dyads consisting of two puppies had significantly more equal play than dyads consisting of one puppy and one adult. These results suggest that wolf puppies are more likely to play on equal terms with similarly-aged play partners, while the dominance status of the partners dictates offensive and self-handicapping behaviors between animals of different ages.

The welfare of animals in captivity is of considerable societal concern. A major source of stress, especially for wild animals, is the lack of control over their environment, which includes not being able to avoid contact with human beings. Paradoxically, some studies have shown that interactions with human beings may improve the welfare of wild animals in captivity. Here, we investigated the behavioural (behaviours indicative of cooperation or stress) and physiological (variations in salivary cortisol concentrations) effects of the increasingly used practice of training wild animals as a way to facilitate handling and/or as behavioural enrichment. We evaluated the effects of indoor training sessions with familiar caretakers on nine human-socialised individuals of a wild species, the wolf (Canis lupus), in comparison to nine individuals of its domesticated form, the dog (Canis lupus familiaris). All animals were raised and kept in intraspecific packs under identical conditions-in accordance with the social structure of the species-in order to control for socialisation with human beings and familiarity with training. We also collected saliva samples of trainers to measure GC and testosterone concentrations, to control for the effects of trainers' stress levels on the responses of the animals. During the training sessions, separated from pack members, the animals stayed voluntarily close to the trainers and mostly adequately performed requested behaviours, indicating concentration to the task. Similarly to dogs, the salivary cortisol level of wolves-used as an index of stress-dropped during these sessions, pointing to a similar stress-reducing effect of the training interaction in both subspecies. The responses to the requested behaviours and the reduction in salivary cortisol level of wolves and dogs varied across trainers, which indicates that the relaxing effect of training has a social component. This points to another factor affecting the welfare of animals during the sessions, beside the rewarding effect of getting food and control over the situation by successfully completing a task. As all responses performed by the animals corresponded to cues already familiar to them, the reported effects were likely due to the above cited factors rather than to a learning process. Our results support previous findings suggesting that training is a potentially powerful tool for improving welfare in some wild social canids by creating structured and positive interactions between these animals and their human caretakers.

Northern Bald Ibis are socially monogamous and year-round colonial birds with a moderate repertoire of calls. Their 'croop', for example, is used during greeting of mates, but also during agonistic encounters, and provides an ideal case to study whether calls are revealing with respect to motivational states. We recorded croop calls in a semi-tame and free-roaming flock of Northern Bald Ibis in Austria, and analysed the vocal structure to identify parameters (e.g. call duration, fundamental frequency) potentially differing between social contexts, sexes and individuals. Additionally, we conducted playback experiments to test whether mated pairs would discriminate each other by their greeting croops. Acoustic features showed highly variable temporal and structural parameters. Almost all calls could be classified correctly and assigned to the different social contexts and sexes. Classification results of greeting croops were less clear for individuality. However, incubating individuals looked up more often and longer in response to playbacks of the greeting calls of their mate than to other colony members, indicating mate recognition. We show that acoustic parameters of agonistic and greeting croops contain features that may indicate the expression of affective states, and that greeting croops encode individual differences that are sufficient for individual recognition.

Selection pressures that act differently on males and females produce numerous differences between the sexes in morphology and behaviour. However, apart from the controversial report that males have slightly heavier brains than females in humans, evidence for substantial sexual dimorphism in brain size is scarce. This apparent sexual uniformity is surprising given that sexually distinct selection pressures are ubiquitous and that brains are one of the most plastic vertebrate organs. Here we demonstrate the highest level of sexual brain size dimorphism ever reported in any vertebrate: male three-spined stickleback of two morphs in an Icelandic lake have 23% heavier brains than females. We suggest that this dramatic sexual size dimorphism is generated by the many cognitively demanding challenges that males are faced in this species, such as an elaborate courtship display, the construction of an ornate nest and a male-only parental care system. However, we consider also alternative explanations for smaller brains in females, such as life-history trade-offs. Our demonstration of unprecedented levels of sexual dimorphism in brain size in the three-spined stickleback implies that behavioural and life-history differences among the sexes can have strong effects also on neural development and proposes new fields of research for understanding brain evolution.

Other-regarding preferences are a critical feature of human cooperation but to what extent non-human animals exhibit these preferences is a matter of intense discussion. We tested whether jackdaws show prosocial behaviour (providing benefits to others at no cost to themselves) and altruism (providing benefits to others while incurring costs) with both sibling and non-sibling recipients. In the prosocial condition, a box was baited on both the actor's and the recipient's side (1/1 option), whereas another box provided food only for the actor (1/0 option). In the altruistic condition, the boxes contained food for either the actor (1/0 option) or the recipient (0/1 option). The proportion of selfish (1/0 option) and cooperative (1/1 and 0/1 option, respectively) actors' choices was significantly affected by the recipients' behaviour. If recipients approached the boxes first and positioned themselves next to the box baited on their side, trying to access the food reward (recipient-first trials), actors were significantly more cooperative than when the actors approached the boxes first and made their choice prior to the recipients' arrival (actor-first trials). Further, in recipient-first trials actors were more cooperative towards recipients of the opposite sex, an effect that was even more pronounced in the altruistic condition. Hence, at no cost to the actors, all recipients could significantly influence the actors' behaviour, whereas at high costs this could be achieved even more so by recipients of different sex. Local/stimulus enhancement is discussed as the most likely cognitive mechanism to account for these effects.

Social species that maintain individualised relationships with certain others despite continuous changes in age, reproductive status and dominance rank between group members ought to be capable of individual recognition. Tests of "true" individual recognition, where an individual recognises unique features of another, are rare, however. Often kinship and/or familiarity suffice to explain dyadic interactions. The complex relationships within a greylag goose flock suggest that they should be able to recognise individuals irrespective of familiarity or kinship. We tested whether six-week-old hand-raised greylags can discriminate between two of their siblings. We developed a new experimental protocol, in which geese were trained to associate social siblings with geometrical symbols. Subsequently, focals were presented with two geometrical symbols in the presence of a sibling associated with one of the symbols. Significant choice of the geometrical symbol associated with the target present indicated that focals were able to distinguish between individual targets. Greylag goslings successfully learned this association-discrimination task, regardless of genetic relatedness or sex of the sibling targets. Social relationships within a goose flock thus may indeed be based on recognition of unique features of individual conspecifics.

An animal's emotional responses are the result of its cognitive appraisal of a situation. This appraisal is notably influenced by the possibility of an individual to exert control over an aversive event. Although the fact that environment controllability decreases emotional responses in animals is well established, far less is known about its potential trans-generational effects. As the levels of avian yolk hormones can vary according to the mother's environment, we hypothesized that housing environment of mothers would modulate the quality of her eggs and in turn her offspring's behaviour. Two groups of female Japanese quail were constituted: a group that had access to a place to hide in their home-cage (Hd, n = 20) and a group that had nowhere to hide (NoHd, n = 20) when stressed. Both groups were submitted to daily human disturbances for a twenty-day-period. Hd females produced eggs with both less testosterone and androstenedione than did NoHd females. The emotional and social reactivity of Hd females' offspring were lower and their growth was slower than those of NoHd females' offspring. Our results show that a minor difference in housing environment had substantial effects on eggs and offspring. The presence of a shelter probably helped quail to cope with daily human disturbances, producing less reactive offspring. This transgenerational effect caused by an opportunity to hide could lead to applications in care of laboratory animals, conservation biology and animal welfare.

BackgroundSocial stressors are known to be among the most potent stressors in group-living animals. This is not only manifested in individual physiology (heart rate, glucocorticoids), but also in how individuals behave directly after a conflict. Certain 'stress-related behaviors' such as autopreening, body shaking, scratching and vigilance have been suggested to indicate an individual's emotional state. Such behaviors may also alleviate stress, but the behavioral context and physiological basis of those behaviors is still poorly understood.Methodology/principal findingsWe recorded beat-to-beat heart rates (HR) of 22 greylag geese in response to agonistic encounters using fully implanted sensor-transmitter packages. Additionally, for 143 major events we analyzed the behavior shown by our focal animals in the first two minutes after an interaction. Our results show that the HR during encounters and characteristics of the interaction predicted the frequency and duration of behaviors shown after a conflict.Conclusions/significanceTo our knowledge this is the first study to quantify the physiological and behavioral responses to single agonistic encounters and to link this to post conflict behavior. Our results demonstrate that 'stress-related behaviors' are flexibly modulated by the characteristics of the preceding aggressive interaction and reflect the individual's emotional strain, which is linked to autonomic arousal. We found no support for the stress-alleviating hypothesis, but we propose that stress-related behaviors may play a role in communication with other group members, particularly with pair-partners.

Individual phenotypic characteristics of many species are influenced by non-genetic maternal effects. Female birds can influence the development of their offspring before birth via the yolk steroid content of their eggs. We investigated this prenatal maternal effect by analysing the influence of laying females' social environment on their eggs' hormonal content and on their offspring's development. Social instability was applied to groups of laying Japanese quail females. We evaluated the impact of this procedure on laying females, on yolk steroid levels and on the general development of chicks. Agonistic interactions were more frequent between females kept in an unstable social environment (unstable females) than between females kept in a stable social environment (stable females). Testosterone concentrations were higher in unstable females' eggs than in those of stable females. Unstable females' chicks hatched later and developed more slowly during their first weeks of life than those of stable females. The emotional reactivity of unstable females' chicks was higher than that of stable females' chicks. In conclusion, our study showed that social instability applied to laying females affected, in a non-genetic way, their offspring's development, thus stressing the fact that females' living conditions during laying can have transgenerational effects.

BACKGROUND:Among birds, corvids and parrots are prime candidates for advanced cognitive abilities. Still, hardly anything is known about cognitive similarities and dissimilarities between them. Recently, exclusion has gained increasing interest in comparative cognition. To select the correct option in an exclusion task, one option has to be rejected (or excluded) and the correct option may be inferred, which raises the possibility that causal understanding is involved. However, little is yet known about its evolutionary history, as only few species, and mainly mammals, have been studied. METHODOLOGY/PRINCIPAL FINDINGS:We tested ravens and keas in a choice task requiring the search for food in two differently shaped tubes. We provided the birds with partial information about the content of one of the two tubes and asked whether they could use this information to infer the location of the hidden food and adjust their searching behaviour accordingly. Additionally, this setup allowed us to investigate whether the birds would appreciate the impact of the shape of the tubes on the visibility of food. The keas chose the baited tube more often than the ravens. However, the ravens applied the more efficient strategy, choosing by exclusion more frequently than the keas. An additional experiment confirmed this, indicating that ravens and keas either differ in their cognitive skills or that they apply them differently. CONCLUSION:To our knowledge, this is the first study to demonstrate that corvids and parrots may perform differently in cognitive tasks, highlighting the potential impact of different selection pressures on the cognitive evolution of these large-brained birds.

BackgroundTelomeres--the terminal caps of chromosomes--become shorter as individuals age, and there is much interest in determining what causes telomere attrition since this process may play a role in biological aging. The leading hypothesis is that telomere attrition is due to inflammation, exposure to infectious agents, and other types of oxidative stress, which damage telomeres and impair their repair mechanisms. Several lines of evidence support this hypothesis, including observational findings that people exposed to infectious diseases have shorter telomeres. Experimental tests are still needed, however, to distinguish whether infectious diseases actually cause telomere attrition or whether telomere attrition increases susceptibility to infection. Experiments are also needed to determine whether telomere erosion reduces longevity.Methodology/principal findingsWe experimentally tested whether repeated exposure to an infectious agent, Salmonella enterica, causes telomere attrition in wild-derived house mice (Mus musculus musculus). We repeatedly infected mice with a genetically diverse cocktail of five different S. enterica strains over seven months, and compared changes in telomere length with sham-infected sibling controls. We measured changes in telomere length of white blood cells (WBC) after five infections using a real-time PCR method. Our results show that repeated Salmonella infections cause telomere attrition in WBCs, and particularly for males, which appeared less disease resistant than females. Interestingly, we also found that individuals having long WBC telomeres at early age were relatively disease resistant during later life. Finally, we found evidence that more rapid telomere attrition increases mortality risk, although this trend was not significant.Conclusions/significanceOur results indicate that infectious diseases can cause telomere attrition, and support the idea that telomere length could provide a molecular biomarker for assessing exposure and ability to cope with infectious diseases.

Northern Bald Ibis are socially monogamous and year-round colonial birds with a moderate repertoire of calls. Their ‘croop’, for example, is used during greeting of mates, but also during agonistic encounters, and provides an ideal case to study whether calls are revealing with respect to motivational states. We recorded croop calls in a semi-tame and free-roaming flock of Northern Bald Ibis in Austria, and analysed the vocal structure to identify parameters (e.g. call duration, fundamental frequency) potentially differing between social contexts, sexes and individuals. Additionally, we conducted playback experiments to test whether mated pairs would discriminate each other by their greeting croops. Acoustic features showed highly variable temporal and structural parameters. Almost all calls could be classified correctly and assigned to the different social contexts and sexes. Classification results of greeting croops were less clear for individuality. However, incubating individuals looked up more often and longer in response to playbacks of the greeting calls of their mate than to other colony members, indicating mate recognition. We show that acoustic parameters of agonistic and greeting croops contain features that may indicate the expression of affective states, and that greeting croops encode individual differences that are sufficient for individual recognition. [ABSTRACT FROM AUTHOR]