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Fritz, J., Bisenberger, A., & Kotrschal, K. (2000). Stimulus enhancement in greylag geese: socially mediated learning of an operant task. Animal Behaviour, 59(6), 1119–1125.
Abstract: We recently observed the spreading of a novel tradition in a flock of semiferal greylag geese, Anser anser: an increasing number of individuals began to bite and chew the stems of butterbur, Petasites hybridus. Because this behaviour spread particularly fast within families, social learning seemed to be involved. We therefore designed an experiment with hand-reared goslings, which were socially imprinted on humans, to investigate whether and how the observation of an experienced tutor affects the acquisition of a novel skill. Goslings had to open the gliding lid of a box to get at a food reward. To each of seven hand-reared observers a human tutor demonstrated where and how to open the lid, whereas seven controls remained untutored. All observers learned to perform the task but only one of the controls succeeded. The observers explored more often at the position shown by the tutor than elsewhere and seemingly learned by trial and error. In contrast, control birds explored primarily at positions that did not allow them to open the box. These results indicate that in greylag goslings the observation of an experienced model facilitates the learning of an operant task. We conclude that stimulus enhancement followed by operant conditioning were the mechanisms involved, which may have accounted for the fast spread of the stem-chewing tradition between family members.
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Schmidt, J., Scheid, C., Kotrschal, K., Bugnyar, T., & Schloegl, C. (2011). Gaze direction – A cue for hidden food in rooks (Corvus frugilegus)? Behavioural Processes, 88(2), 88–93.
Abstract: Other individual's head- and eye-directions can be used as social cues indicating the presence of important events. Among birds, ravens and rooks have been shown to co-orient with conspecifics and with humans by following their gaze direction into distant space and behind visual screens. Both species use screens to cache food in private; also, it had been suggested that they may rely on gaze cues to detect hidden food. However, in an object-choice task, ravens failed to do so, and their competitive lifestyle may have prevented them from relying on these cues. Here we tested closely related and cooperative rooks. Food was hidden in one of two cups and the experimenter gazed at the baited cup. In a second experiment, we aimed to increase the birds’ motivation to choose correctly by increasing the investment needed to obtain the reward. To do so, the birds had to pull on a string to obtain the cup. Here, the birds as a group tended to rely on gaze cues. In addition, individual birds quickly learned to use the cue in both experiments. Although rooks may not use gaze cues to find hidden food spontaneously, they may quickly learn to do so.
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Schwab, C., Bugnyar, T., Schloegl, C., & Kotrschal, K. (2008). Enhanced social learning between siblings in common ravens, Corvus corax. Anim. Behav., 75(2), 501–508.
Abstract: It has been suggested that social dynamics affect social learning but empirical support for this idea is scarce. Here we show that affiliate relationships among kin indeed enhance the performance of common ravens, Corvus corax, in a social learning task. Via daily behavioural protocols we first monitored social dynamics in our group of captive young ravens. Siblings spent significantly more time in close proximity to each other than did nonsiblings. We subsequently tested birds on a stimulus enhancement task in model-observer dyads composed of both siblings and nonsiblings. During demonstration the observer could watch the model manipulating one particular object (target object) in an adjacent room. After removing the model, the observer was confronted with five different objects including the former target object. Observers from sibling dyads handled the target object for significantly longer periods of time as compared with the other four available objects, whereas observers from nonsibling dyads did not show a preference for the target object. Also, siblings matched the model's decision to cache or not to cache objects significantly more often than did nonsiblings. Hence, siblings were likely to attend to both, the behaviour of the model (caching or noncaching) and object-specific details. Our results support the hypothesis that affiliate relations between individuals affect the transmission of information and may lead to directed social learning even when spatial proximity has been experimentally controlled for.
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Wascher, C. A. F., Fraser, O. N., & Kotrschal, K. (2010). Heart Rate during Conflicts Predicts Post-Conflict Stress-Related Behavior in Greylag Geese. PLoS ONE, 5(12), e15751.
Abstract: Background
Social 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 Findings
We 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/Significance
To 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.
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Kotrschal, K., Schöberl, I., Bauer, B., Thibeaut, A. - M., & Wedl, M. (2009). Dyadic relationships and operational performance of male and female owners and their male dogs. Behav. Process., 81(3), 383–391.
Abstract: In the paper we investigate how owner personality, attitude and gender influence dog behavior, dyadic practical functionality and the level of dog salivary cortisol. In three meetings, 12 female and 10 male owners of male dogs answered questionnaires including the Neo-FFI human personality inventory. Their dyadic behavior was video-taped in a number of test situations, and saliva samples were collected. Owners who scored highly in neuroticism (Neo-FFI dimension one) viewed their dogs as social supporters and spent much time with them. Their dogs had low baseline cortisol levels, but such dyads were less successful in the operational task. Owners who scored highly in extroversion (Neo-FFI dimension two) appreciated shared activities with their dogs which had relatively high baseline cortisol values. Dogs that had female owners were less sociable-active (dog personality axis 1) than dogs that had male owners. Therefore, it appears that owner gender and personality influences dyadic interaction style, dog behavior and dyadic practical functionality.
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Stöwe, M. & K., K. (2007). Behavioural phenotypes may determine whether social context facilitates or delays novel object exploration in ravens. International Journal of Ornithology, 148(Suppl 2), 179–184.
Abstract: Individuals consistently differ in behavioural phenotypes. Here we examine the interaction between behavioural phenotype and response to social context during novel object exploration in a neophobic corvid species, the raven (Corvus corax). The presence of conspecifics tends to encourage object exploration and learning but may also delay or even inhibit exploratory behaviour. Factors such as individual differences in response to social context may determine whether the presence of a conspecific facilitates or inhibits approach to novel objects. We confronted eleven six-month-old hand-raised ravens with novel objects, both individually and in dyadic combinations. We defined individuals as “fast” and “slow” explorers on the basis of their approach latency to novel objects when tested individually. The presence of a conspecific delayed the approach of fast birds to novel objects. Slow birds, in contrast, approached the novel objects with lower latencies and spent more time close to them when in dyads with fast siblings than when alone. The individuals" approach behaviour seemed to determine whether social context facilitated or delayed exploratory behaviour. This may contribute to explaining ambiguous results concerning the effects of social context in previous studies.
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Schloegl, C., Kotrschal, K., & Bugnyar, T. (2008). Modifying the object-choice task: Is the way you look important for ravens? Behav. Process., 77(1), 61–65.
Abstract: Most animals seem to have difficulties in using gaze cues to find hidden food in object-choice tasks. For instance, chimpanzees usually fail in these tests, even though they are capable of following other's gaze geometrically behind barriers. Similar to chimpanzees, common ravens are skilled in tracking other's gaze but fail in object-choice tasks. We here explored whether procedural modifications, which had been used successfully in chimpanzees, would also yield positive results in ravens. In our modifications (a) the experimenter approached the cup while gazing at it, (b) the gaze cue was accompanied by a sound and (c) the experimenter could actually see the food while giving the gaze cue. Two out of seven birds performed above chance level in some of these conditions. However, we ascribe this improvement to the individuals' learning ability rather than to an understanding of the communicative nature of the task. This interpretation is further supported by results of a follow-up experiment suggesting that ravens may not rely on conspecifics' gaze cues for finding food caches in a natural foraging context. In sum, our results suggest that ravens may not transfer their gaze follow abilities to foraging situations involving hidden food.
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Scheiber, I. B. R., WeiB, B. M., Hirschenhauser, K., Wascher, C. A. F., Nedelcu, J. T., & Kotrschal, K. (2008). Does 'Relationship Intelligence' Make Big Brains in Birds? Open Biol J, 1, 6–8 (3).
Abstract: Lately, Emery et al. developed a bird-specific modification of the *social brain hypothesis*, termed *relationship intelligence hypothesis*. Although the idea may be valuable, we doubt that it is supported by sufficient evidence and critically discuss some of the arguments raised by the authors in favour of their new idea.
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Schloegl, C., Kotrschal, K., & Bugnyar, T. (2007). Gaze following in common ravens, Corvus corax: ontogeny and habituation. Anim. Behav., 74(4), 769–778.
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.
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Kralj-Fiser, S., Scheiber, I. B. R., Blejec, A., Moestl, E., & Kotrschal, K. (2007). Individualities in a flock of free-roaming greylag geese: behavioral and physiological consistency over time and across situations. Horm Behav, 51(2), 239–248.
Abstract: The concept of personality implies individual differences in behavior and physiology that show some degree of repeatability/consistency over time and across contexts. Most studies of animal personality, particularly studies of individuals' variation in physiological mechanisms, have been conducted on selected individuals in controlled conditions. We attempted to detect consistent behaviors as well as physiological patterns in greylag ganders (Anser anser) from a free-roaming flock living in semi-natural conditions. We tested 10 individuals repeatedly, in a handling trial, resembling tests for characterization of “temperaments” in captive animals. We recorded the behavior of the same 10 individuals during four situations in the socially intact flock: (1) a “low density feeding condition”, (2) a “high density feeding condition”, (3) a “low density post-feeding situation” and (4) while the geese rested. We collected fecal samples for determination of excreted immuno-reactive corticosterone (BM) and testosterone metabolites (TM) after handling trials, as well as the “low density feeding” and the “high density feeding” conditions. BM levels were very highly consistent over the repeats of handling trials, and the “low density feeding condition” and tended to be consistent over the first two repeats of the “high density feeding condition”. Also, BM responses tended to be consistent across contexts. Despite seasonal variation, there tended to be inter-test consistency of TM, which pointed to some individual differences in TM as well. Aggressiveness turned out to be a highly repeatable trait, which was consistent across social situations, and tended to correlate with an individual's resistance during handling trials. Also, “proximity to the female partner” and “sociability” – the average number of neighboring geese in a close distance while resting – were consistent. We conclude that aggressiveness, “affiliative tendencies” and levels of excreted corticosterone and testosterone metabolites may be crucial factors of personality in geese.
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