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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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Daniel, J. C., & Mikulka, P. J. (1998). Discrimination learning in the white rhinoceros. Appl. Anim. Behav. Sci., 58(1–2), 197–202.
Abstract: This study examined the ability of two adult white rhinoceroses (Ceratotherium simum simum) to develop a visual discrimination between an open circle and a triangle. These stimuli were presented as black symbols on large white cards. The cards were presented 4.6 m apart and a food reward was given if the subject approached the open circle. Ten discrimination choices were given daily until each subject reached the criterion of 80% correct responding over a block of 50 trials. The female reached the criterion over trials 151–200, while the male required considerably longer (trials 501–550). The male's discrimination was dramatically affected by a shift in the food reward. This study demonstrates that these rhinos were able to develop a successful discrimination and this protocol could be used to further examine their visual acuity.
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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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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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Fenner, K., Freire, R., McLean, A., & McGreevy, P. (2018). Behavioral, demographic and management influences on equine responses to negative reinforcement. Journal of Veterinary Behavior, .
Abstract: Understanding the factors that influence horse learning is critical to ensure horse welfare and rider safety. In this study, data were obtained from horses (n=96) training to step backwards through a corridor in response to bit pressure. Following training, learning ability was determined by the latency to step backwards through the corridor when handled on the left and right reins. Additionally, horse owners were questioned about each horse's management, training, behavior and signalment (such as horse breed, age and sex). Factors from these four broad domains were examined using a multiple logistic regression (MLR) model, following an Information Theoretic approach, for associations between horses' behavioral attributes and their ability to learn the task. The MLR also included estimates of the rider's ability and experience as well as owner's perceptions of their horse's trainability and temperament. Results revealed several variables including explanatory variables that correlated significantly with rate of learning. Horses were faster at backing, a behavioral trait, when handled on the right (t = 3.65, df = 94, P < 0.001) than the left side. Thoroughbred horses were slower at completing the tests than other breeds of horses when handled on the left side (LM, F1,48=4.5, P=0.04) and right side (LM, F1,45=6.0, P=0.02). Those in regular work, a training factor, did not learn faster than their unworked counterparts on the right rein but completed the task faster on the left rein (F1,44=5.47, P=0.02). This may reflect differences in laterality and habituation effects. In contrast, more anxious horses were faster at completing the test when handled from the right (Spearman, r=-0.22, P=0.04). It is possible that these horses have an increased arousal level when interacting with handlers, resulting in more engagement with the lesson, accounting for the improved performance results. The findings of this study will help clarify how horse behavior, training and management may influence learning and how their application may optimize learning outcomes. Future equine behavior assessment and research questionnaires should include items that assess these qualities.
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Hagen, K., & Broom, D. M. (2004). Emotional reactions to learning in cattle. Appl. Anim. Behav. Sci., 85(3), 203–213.
Abstract: It has been suggested that during instrumental learning, animals are likely to react emotionally to the reinforcer. They may in addition react emotionally to their own achievements. These reactions are of interest with regard to the animals' capacity for self-awareness. Therefore, we devised a yoked control experiment involving the acquisition of an operant task. We aimed to identify the emotional reactions of young cattle to their own learning and to separate these from reactions to a food reward. Twelve Holstein-Friesian heifers aged 7-12 months were divided into two groups. Heifers in the experimental group were conditioned over a 14-day period to press a panel in order to open a gate for access to a food reward. For heifers in the control group, the gate opened after a delay equal to their matched partner's latency to open it. To allow for observation of the heifers' movements during locomotion after the gate had opened, there was a 15m distance in the form of a race from the gate to the food trough. The heart rate of the heifers, and their behaviour when moving along the race towards the food reward were measured. When experimental heifers made clear improvements in learning, they were more likely than on other occasions to have higher heart rates and tended to move more vigorously along the race in comparison with their controls. This experiment found some, albeit inconclusive, indication that cattle may react emotionally to their own learning improvement.
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Pongrácz, P., Miklósi, Á., Vida, V., & Csányi, V. (2005). The pet dogs ability for learning from a human demonstrator in a detour task is independent from the breed and age. Appl. Anim. Behav. Sci., 90(3), 309–323.
Abstract: There are many indications and much practical knowledge about the different tasks which various breeds of dogs are selected for. Correspondingly these different breeds are known to possess different physical and mental abilities. We hypothesized that commonly kept breeds will show differences in their problem solving ability in a detour task around a V-shaped fence, and also, that breed differences will affect their learning ability from a human demonstrator, who demonstrates a detour around the fence. Subjects were recruited in Hungarian pet dog schools. We compared the results of the 10 most common breeds in our sample when they were tested in the detour task without human demonstration. There was no significant difference between the latencies of detour, however, there was a trend that German Shepherd dogs were the quickest and Giant Schnauzers were the slowest in this test. For testing the social learning ability of dogs we formed three breed groups (“utility”, “shepherd” and “hunting”). There were no significant differences between these, all the breed groups learned equally well from the human demonstrator. However, we found that dogs belonging to the “shepherd” group looked back more frequently to their owner than the dogs in the “hunting” group. Further, we have found that the age of pet dogs did not affect their social learning ability in the detour task. Our results showed that the pet status of a dog has probably a stronger effect on its cognitive performance and human related behaviour than its age or breed. These results emphasize that socialization and common activities with the dog might overcome the possible breed differences, if we give the dogs common problem solving, or social learning tasks.
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Ahrendt, L. P., Labouriau, R., Malmkvist, J., Nicol, C. J., & Christensen, J. W. (2015). Development of a standard test to assess negative reinforcement learning in horses. Appl. Anim. Behav. Sci., 169, 38–42.
Abstract: Most horses are trained by negative reinforcement. Currently, however, no standardised test for evaluating horses' negative reinforcement learning ability is available. The aim of this study was to develop an objective test to investigate negative reinforcement learning in horses. Twenty-four Icelandic horses (3 years old) were included in this study. The horses were tested in a pressure-release task on three separate days with 10, 7 and 5 trials on each side, respectively. Each trial consisted of pressure being applied on the hindquarter with an algometer. The force of the pressure was increased until the horse moved laterally away from the point of pressure. There was a significant decrease in required force over trials on the first test day (P<0.001), but not the second and third day. The intercepts on days 2 and 3 differed significantly from day 1 (P<0.001), but not each other. Significantly stronger force was required on the right side compared to the left (P<0.001), but there was no difference between first and second side tested (P=0.56). Individual performance was evaluated by median-force and the change in force over trials on the first test day. These two measures may explain different characteristics of negative reinforcement learning. In conclusion, this study presents a novel, standardised test for evaluating negative reinforcement learning ability in horses.
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Hartmann, E., Christensen, J. W., & McGreevy, P. D. (2017). Dominance and Leadership: Useful Concepts in Human-Horse Interactions? Proceedings of the 2017 Equine Science Symposium, 52, 1–9.
Abstract: Dominance hierarchies in horses primarily influence priority access to limited resources of any kind, resulting in predictable contest outcomes that potentially minimize aggressive encounters and associated risk of injury. Levels of aggression in group-kept horses under domestic conditions have been reported to be higher than in their feral counterparts but can often be attributed to suboptimal management. Horse owners often express concerns about the risk of injuries occurring in group-kept horses, but these concerns have not been substantiated by empirical investigations. What has not yet been sufficiently addressed are human safety aspects related to approaching and handling group-kept horses. Given horse's natural tendency to synchronize activity to promote group cohesion, questions remain about how group dynamics influence human-horse interactions. Group dynamics influence a variety of management scenarios, ranging from taking a horse out of its social group to the prospect of humans mimicking the horse's social system by taking a putative leadership role and seeking after an alpha position in the dominance hierarchy to achieve compliance. Yet, there is considerable debate about whether the roles horses attain in their social group are of any relevance in their reactions to humans. This article reviews the empirical data on social dynamics in horses, focusing on dominance and leadership theories and the merits of incorporating those concepts into the human-horse context. This will provide a constructive framework for informed debate and valuable guidance for owners managing group-kept horses and for optimizing human-horse interactions.
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Broad, K. D., Curley, J. P., & Keverne, E. B. (2006). Mother-infant bonding and the evolution of mammalian social relationships. Phil. Trans. Biol. Sci., 361(1476), 2199–2214.
Abstract: A wide variety of maternal, social and sexual bonding strategies have been described across mammalian species, including humans. Many of the neural and hormonal mechanisms that underpin the formation and maintenance of these bonds demonstrate a considerable degree of evolutionary conservation across a representative range of these species. However, there is also a considerable degree of diversity in both the way these mechanisms are activated and in the behavioural responses that result. In the majority of small-brained mammals (including rodents), the formation of a maternal or partner preference bond requires individual recognition by olfactory cues, activation of neural mechanisms concerned with social reward by these cues and gender-specific hormonal priming for behavioural output. With the evolutionary increase of neocortex seen in monkeys and apes, there has been a corresponding increase in the complexity of social relationships and bonding strategies together with a significant redundancy in hormonal priming for motivated behaviour. Olfactory recognition and olfactory inputs to areas of the brain concerned with social reward are downregulated and recognition is based on integration of multimodal sensory cues requiring an expanded neocortex, particularly the association cortex. This emancipation from olfactory and hormonal determinants of bonding has been succeeded by the increased importance of social learning that is necessitated by living in a complex social world and, especially in humans, a world that is dominated by cultural inheritance. © 2006 The Royal Society.
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