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Lim, M. M., & Young, L. J. (2006). Neuropeptidergic regulation of affiliative behavior and social bonding in animals. Hormon. Behav., 50(4), 506–517.
Abstract: Social relationships are essential for maintaining human mental health, yet little is known about the brain mechanisms involved in the development and maintenance of social bonds. Animal models are powerful tools for investigating the neurobiological mechanisms regulating the cognitive processes leading to the development of social relationships and for potentially extending our understanding of the human condition. In this review, we discuss the roles of the neuropeptides oxytocin and vasopressin in the regulation of social bonding as well as related social behaviors which culminate in the formation of social relationships in animal models. The formation of social bonds is a hierarchical process involving social motivation and approach, the processing of social stimuli and formation of social memories, and the social attachment itself. Oxytocin and vasopressin have been implicated in each of these processes. Specifically, these peptides facilitate social affiliation and parental nurturing behavior, are essential for social recognition in rodents, and are involved in the formation of selective mother-infant bonds in sheep and pair bonds in monogamous voles. The convergence of evidence from these animal studies makes oxytocin and vasopressin attractive candidates for the neural modulation of human social relationships as well as potential therapeutic targets for the treatment of psychiatric disorders associated with disruptions in social behavior, including autism.
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Beery, A. K., & Kaufer, D. (2015). Stress, social behavior, and resilience: Insights from rodents. Neurobiol. Stress, 1(Stress Resilience), 116–127.
Abstract: The neurobiology of stress and the neurobiology of social behavior are deeply intertwined. The social environment interacts with stress on almost every front: social interactions can be potent stressors; they can buffer the response to an external stressor; and social behavior often changes in response to stressful life experience. This review explores mechanistic and behavioral links between stress, anxiety, resilience, and social behavior in rodents, with particular attention to different social contexts. We consider variation between several different rodent species and make connections to research on humans and non-human primates.
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Sueur, C., Jacobs, A., Amblard, F., Petit, O., & King, A. J. (2010). How can social network analysis improve the study of primate behavior? Am. J. Primatol., 73(8), 703–719.
Abstract: Abstract When living in a group, individuals have to make trade-offs, and compromise, in order to balance the advantages and disadvantages of group life. Strategies that enable individuals to achieve this typically affect inter-individual interactions resulting in nonrandom associations. Studying the patterns of this assortativity using social network analyses can allow us to explore how individual behavior influences what happens at the group, or population level. Understanding the consequences of these interactions at multiple scales may allow us to better understand the fitness implications for individuals. Social network analyses offer the tools to achieve this. This special issue aims to highlight the benefits of social network analysis for the study of primate behaviour, assessing it's suitability for analyzing individual social characteristics as well as group/population patterns. In this introduction to the special issue, we first introduce social network theory, then demonstrate with examples how social networks can influence individual and collective behaviors, and finally conclude with some outstanding questions for future primatological research. Am. J. Primatol. 73:703?719, 2011. ? 2011 Wiley-Liss, Inc.
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Krueger., K., & Farmer, K. (2018). Social learning in Horses: Differs from individual learning only in the learning stimulus and not in the learning mechanisms. In 14th Meeting of the Internatinoal Society for Equitation Science.
Abstract: Equine welfare can be enhanced by applying species specific training. This may incorporate social learning, as horses are highly social and social stimuli are of primary importance. Social learning is comparable to individual learning in its learning mechanisms, differing primarily in the way it is stimulated. Our initial study showed that horses of different breeds (N = 38) follow humans after observing other horses doing so, but only if the observed horse was familiar to and higher ranking than the observer (Fisher's exact test: N = 12, P = 0.003). A second study showed that horses and ponies (N = 25) learned to pull a rope to open a feeding apparatus after observing demonstrations by conspecifics, again, only if the demonstrating horse was older and higher ranking than the observer (Fisher's combination test, N = 3, v2 = 27.71, p = 0.006). Our third approach showed that horses and ponies (N = 24) learned to press a switch to open a feeding apparatus after observing a familiar person (GzLM: N = 24, z = 2.33, P = 0.02). Most recently, we confronted horses and ponies (N = 50) with persons demonstrating different techniques for opening a feeding apparatus. In this study we investigated whether the horses would copy the demonstrators' techniques or apply their own. Here only some horses copied the technique, and most of the successful learners used their mouths irrespective of the demonstrators' postures (Chi Square Test: N = 40, df = 2, χ2 = 31.4, p < 0.001). In all the approaches social stimuli elicited learning processes in the test horses, while only a few individuals in the control groups mastered the tasks by individual learning. The following behaviour observed in the initial study may have been facilitated by a social stimuli (social facilitation), and the opening of the feed boxes in the subsequent studies appear to be mostly the result of enhancement (social enhancement). Some horses may have used the social stimuli at first and continued their learning process by individual trial and error. However, the horses were also selective in whom and some in how to copy. This may have been conditioned (socially conditioned) or the result of simple forms of reasoning on the reliability of the particular information provided by demonstrators of certain social ranks or social positions, as high ranking and familiar horses and familiar persons were copied and some imitated exactly.
Lay person message: Traditional riding instructions suggest that horses learn by observing other horses. For example, older, more experienced driving horses are used for initial training of young driving horses. We have shown that horses indeed use learning stimuli provided by other horse, as well as by humans. Horses readily accept stimuli observed in high ranking and familiar horses, and familiar persons. Such stimuli elicit learning processes which are comparable to individual learning. We suggest applying social learning whenever possible, as it is much faster and less stressful than individual learning, where learners experience negative outcomes in trial and error learning.
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Quaresmini, C., Forrester, G. S., Spiezio, C., & Vallortigara, G. (2014). Social environment elicits lateralized behaviors in gorillas (Gorilla gorilla gorilla) and chimpanzees (Pan troglodytes). Journal of Comparative Psychology, 128(3), 276–284.
Abstract: The influence of the social environment on lateralized behaviors has now been investigated across a wide variety of animal species. New evidence suggests that the social environment can modulate behavior. Currently, there is a paucity of data relating to how primates navigate their environmental space, and investigations that consider the naturalistic context of the individual are few and fragmented. Moreover, there are competing theories about whether only the right or rather both cerebral hemispheres are involved in the processing of social stimuli, especially in emotion processing. Here we provide the first report of lateralized social behaviors elicited by great apes. We employed a continuous focal animal sampling method to record the spontaneous interactions of a captive zoo-living colony of chimpanzees (Pan troglodytes) and a biological family group of peer-reared western lowland gorillas (Gorilla gorilla gorilla). We specifically focused on which side of the body (i.e., front, rear, left, right) the focal individual preferred to keep conspecifics. Utilizing a newly developed quantitative corpus-coding scheme, analysis revealed both chimpanzees and gorillas demonstrated a significant group-level preference for focal individuals to keep conspecifics positioned to the front of them compared with behind them. More interestingly, both groups also manifested a population-level bias to keep conspecifics on their left side compared with their right side. Our findings suggest a social processing dominance of the right hemisphere for context-specific social environments. Results are discussed in light of the evolutionary adaptive value of social stimulus as a triggering factor for the manifestation of group-level lateralized behaviors. (PsycINFO Database Record (c) 2016 APA, all rights reserved)
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McVey, A., Wilkinson, A., & Mills, D. S. (2018). Social learning in horses: the effect of using a group leader demonstrator on the performance of familiar conspecifics in a detour task. Applied Animal Behaviour Science, .
Abstract: Learning through the observation of others allows the transfer of information without the costs incurred during individual trial and error learning. Horses (Equus caballus) are a highly social species, which might be expected to be capable of learning from others, but experimental findings are inconsistent, and potentially confounded by social facilitation effects not related directly to the learning of the task. We refined the methods used in previous equine social learning studies, to examine and distinguish specific social influences on learning of a task: we used predefined group leaders rather than agonistically dominant individuals to demonstrate a detour task to familiar conspecific observers; in addition we had two control groups: a non-observer (true control) and a group with the demonstrator simply present at the goal (social facilitation control). 44 socially kept horses were allocated to one of the three test conditions and took part in five trials each. Success rate, latency and detour direction were recorded. There was no significant difference between the three groups in the likelihood of them succeeding in the task nor latency to succeed; however there was a significant difference in the route chosen by the groups, with the true control choosing the side with the entrance gate significantly more than either the observer group or social facilitation group. Both of the latter two groups chose to go in the same direction relative to themselves, regardless of which side the gate was. Seven out of nine horses in the observer group chose the same direction as their demonstrator every time. Our results show a significant role of social facilitation on detour behaviour and highlight the importance of including adequate controls for simpler cognitive influences on behaviour before claims can be made about the specific learning of motor actions or goal directed behaviour. Social cues may be important to horses if the task is sufficiently challenging and motivationally important, so future work should consider more demanding, but ecologically relevant situations, in order to maximise the potential revelation of social learning effects which do not depend on simple local or stimulus enhancement effects.
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Burla, J. - B., Siegwart, J., & Nawroth, C. (2018). Human Demonstration Does Not Facilitate the Performance of Horses (Equus caballus) in a Spatial Problem-Solving Task. Animal, 8(6), 96.
Abstract: Horses’ ability to adapt to new environments and to acquire new information plays an important role in handling and training. Social learning in particular would be very adaptive for horses as it enables them to flexibly adjust to new environments. In the context of horse handling, social learning from humans has been rarely investigated but could help to facilitate management practices. We assessed the impact of human demonstration on the spatial problem-solving abilities of horses during a detour task. In this task, a bucket with a food reward was placed behind a double-detour barrier and 16 horses were allocated to two test groups of 8 horses each. One group received a human demonstration of how to solve the spatial task while the other group received no demonstration. We found that horses did not solve the detour task more often or faster with human demonstration. However, both test groups improved rapidly over trials. Our results suggest that horses prefer to use individual rather than social information when solving a spatial problem-solving task
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KOIZUMI, R., MITANI, T., UEDA, K., & KONDO, S. (2017). Skill reading of human social cues by horses (Equus caballus) reared under year-round grazing conditions. Animal Behaviour and Management, 53(2), 69–78.
Abstract: Animals use communicative signals, such as gesture or gaze, to communicate to someone the intention or expression of the sender, which is called social cue. In the previous studies, it was suggested the skill of reading human social cue in domestic animals are influenced to the domestication, the experience contacting with human and training to obey human. In this present study, we tested the skill for horses (Equus caballus) kept in year-round grazing conditions using 33 horses differed from breed and the degree of the experience with human by object-choice task subjects choosing either of bait boxes located at the end of experimenter. As results, non-socialized horses hardly responded to human social cues. Habituated horses that were both of trained and untrained responded to human social cues, but their accuracy rates were not more than 50% except for two trained subjects. For the skill of reading human social cues, there was high individual variation in responding to human social cues in horses kept in year-round grazing conditions. The individual characteristics influenced to it more than domestication, the experience with human, and training to obey human.
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Briard, L., Dorn, C., & Petit, O. (2015). Personality and Affinities Play a Key Role in the Organisation of Collective Movements in a Group of Domestic Horses. Ethology, 121(9), 888–902.
Abstract: Understanding how groups of individuals with different motives come to daily decisions about the exploitation of their environment is a key question in animal behaviour. While interindividual differences are often seen only as a threat to group cohesion, growing evidence shows that they may to some extent facilitate effective collective action. Recent studies suggest that personality differences influence how individuals are attracted to conspecifics and affect their behaviour as an initiator or a follower. However, most of the existing studies are limited to a few taxa, mainly social fish and arthropods. Horses are social herbivores that live in long-lasting groups and show identifiable personality differences between individuals. We studied a group of 38 individuals living in a 30-ha hilly pasture. Over 200 h, we sought to identify how far individual differences such as personality and affinity distribution affect the dynamic of their collective movements. First, we report that individuals distribute their relationships according to similar personality and hierarchical rank. This is the first study that demonstrates a positive assortment between unrelated individuals according to personality in a mammal species. Second, we measured individual propensity to initiate and found that bold individuals initiated more often than shy individuals. However, their success in terms of number of followers and joining duration did not depend on their individual characteristics. Moreover, joining process is influenced by social network, with preferred partners following each other and bolder individuals being located more often at the front of the movement. Our results illustrate the importance of taking into account interindividual behavioural differences in studies of social behaviours.
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Boogert, N. J., Reader, S. M., Hoppitt, W., & Laland, K. N. (2008). The origin and spread of innovations in starlings. Anim. Behav., 75(4), 1509–1518.
Abstract: There are numerous reports of novel learned behaviour patterns in animal populations, yet the factors influencing the invention and spread of these innovations remain poorly understood. Here we investigated to what extent the pattern of spread of innovations in captive groups of starlings, Sturnus vulgaris, could be predicted by knowledge of individual and social group variables, including association patterns, social rank orders, measures of neophobia and asocial learning performance. We presented small groups of starlings with a series of novel extractive foraging tasks and recorded the latency for each bird to contact and solve each task, as well as the orders of contacting and solving. We then explored which variables best predicted the observed diffusion patterns. Object neophobia and social rank measures characterized who was the first of the group to contact the novel foraging tasks, and the subsequent spread of contacting tasks was associated with latency to feed in a novel environment. Asocial learning performance, measured in isolation, predicted who was the first solver of the novel foraging tasks in each group. Association patterns did not predict the spread of solving. Contact latency and solving duration were negatively correlated, consistent with social learning underlying the spread of solving. Our findings indicate that we can improve our understanding of the diffusion dynamics of innovations in animal groups by investigating group-dependent and individual variables in combination. We introduce novel methods for exploring predictors of the origin and spread of behavioural innovations that could be widely applied.
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