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Physical activity is generally considered as most relevant for modulating heart rate (HR). The authors show here that HR is not only modulated by physical activity but even more by social contexts. HR modulation in three free-ranging, socially embedded, male greylag geese fitted with implanted radiotransmitters was investigated. Measured HR ranged from 40 beats per minute (bpm) during rest to a maximum over 400 bpm during takeoff. Almost the same maximum HRs (400 bpm) were reached during social interactions, which however, generally require less bodily action. Mean HR during social interactions (agonistic interactions, vocalizations) was significantly higher than during behaviors with a less obvious social context (e.g., resting, comfort or feeding behavior), but with comparable physical activity involved. The authors also found significant and consistent differences in HR between the three focal individuals, probably because of individual behavioral phenotype. Our results show that social context has a strong modulatory effect on the sympathico-adrenergic activity in a social bird and conclude that particularly the latter may pose considerable energetic costs. Keywords: heart rate, social modulation of heart rate, physical activity, individual differences, greylag geese

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.

To link to full-text access for this article, visit this link: http://dx.doi.org/10.1016/j.anbehav.2006.08.017 Byline: Christian Schloegl, Kurt Kotrschal, Thomas Bugnyar Abstract: Co-orientation with others by using their gaze direction is considered to be adaptive for detecting food or predators or monitoring social interactions. Like the great apes, common ravens are capable of following human experimenters' gaze direction not only into distant space but also behind visual barriers. We investigated the ontogenetic development of these abilities by confronting birds with a human foster parent looking up (experiment 1) and behind visual barriers (experiment 3) and their modification by habituation (experiments 2 and 4). We tested a group of 12 hand-reared ravens during their first 10 months of life. Ravens responded to others' look-ups soon after fledging but could track their gaze behind a visual barrier only 4 months later, at the age they usually become independent from their parents. Furthermore, ravens quickly ceased responding to repeated look-ups by the model, but did not habituate to repeated gaze cues directed behind a barrier. Our findings support the idea that the two modes of gaze following reflect different cognitive levels in ravens and, possibly, have different functions. Author Affiliation: (a) Konrad Lorenz Forschungsstelle Grunau, Austria (a ) Department of Animal Physiology, University of Bayreuth, Germany (a ) Department of Behavioural Biology, University of Vienna, Austria Article History: Received 6 March 2006; Revised 25 April 2006; Accepted 16 August 2006 Article Note: (miscellaneous) MS. number: 8870R

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.

Flexible or innovative behavior is advantageous, especially when animals are exposed to frequent and unpredictable environmental perturbations. Improved cognitive abilities can help animals to respond quickly and adequately to environmental dynamics, and therefore changing environments may select for higher cognitive abilities. Increased cognitive abilities can be attained, for instance, if environmental change during ontogeny triggers plastic adaptive responses improving the learning capacity of exposed individuals. We tested the learning abilities of fishes in response to experimental variation of environmental quality during ontogeny. Individuals of the cichlid fish Simochromis pleurospilus that experienced a change in food ration early in life outperformed fish kept on constant rations in a learning task later in life--irrespective of the direction of the implemented change and the mean rations received. This difference in learning abilities between individuals remained constant between juvenile and adult stages of the same fish tested 1 y apart. Neither environmental enrichment nor training through repeated neural stimulation can explain our findings, as the sensory environment was kept constant and resource availability was changed only once. Instead, our results indicate a pathway by which a single change in resource availability early in life permanently enhances the learning abilities of animals. Early perturbations of environmental quality may signal the developing individual that it lives in a changing world, requiring increased cognitive abilities to construct adequate behavioral responses.

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.

The attribution of mental states (MS) to other species typically follows a scala naturae pattern. However, "simple" mental states, including emotions, sensing, and feelings are attributed to a wider range of animals as compared to the so-called "higher" cognitive abilities. We propose that such attributions are based on the perceptual quality (i.e. imageability) of mental representations related to MS concepts. We hypothesized that the attribution of highly imaginable MS is more dependent on the familiarity of participants with animals when compared to the attribution of MS low in imageability. In addition, we also assessed how animal agreeableness, familiarity with animals, and the type of human-animal interaction related to the judged similarity of animals to humans. Sixty-one participants (19 females, 42 males) with a rural (n = 20) and urban (n = 41) background rated twenty-six wild and domestic animals for their perceived similarity with humans and ability to experience a set of MS: (1) Highly imageable MS: joy, anger, and fear, and (2) MS low in imageability: capacity to plan and deceive. Results show that more agreeable and familiar animals were considered more human-like. Primates, followed by carnivores, suines, ungulates, and rodents were rated more human-like than xenarthrans, birds, arthropods, and reptiles. Higher MS ratings were given to more similar animals and more so if the MS attributed were high in imageability. Familiarity with animals was only relevant for the attribution of the MS high in imageability. [ABSTRACT FROM AUTHOR]

Dogs are becoming increasingly popular in pedagogical settings. Particularly children with special educational needs are believed to benefit from dog-assisted interventions. However, reliable evidence for supporting such claims is still scarce and reports on the effectiveness of this approach are often anecdotal. With our review we aim at evaluating the literature to answer the question, whether dog-assisted interventions in an educational setting can help children with special educational needs to improve and to develop their emotional, social and cognitive skills. Following the PRISMA Guidelines, the literature was systematically searched for experimental studies until February 2021. Eighteen studies were finally included, which varied greatly in type of intervention, outcomes measured, sample sizes, and scientific quality, which precluded a formal meta-analysis. Hence, we resorted to a narrative synthesis. Overall, the studies report mixed results in the different functional domains of stress reduction, motivation, social skills, cognitive abilities, reading abilities, social conduct, and mental wellbeing. No study reported any negative effects of the intervention. The most unequivocal evidence comes from studies on dogs' effects on physiological stress response in challenging situations and on motivation and adherence to instructions, reporting significantly lower levels of cortisol in both children and pedagogues in the presence of dogs, as well as increased motivation to learn and participate. Findings for other outcomes, academic or social, however, remain inconclusive. Data on long-term effects are lacking altogether. Still, this review indicates the potentials of dog-assisted interventions in special pedagogy, particularly towards supporting a calm and trustful social atmosphere. [ABSTRACT FROM AUTHOR]