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Farmer, K., Krüger, K., Byrne, R. W., & Marr, I. (2018). Sensory laterality in affiliative interactions in domestic horses and ponies (Equus caballus). Anim. Cogn., 21(5), 631–637.
Abstract: Many studies have been carried out into both motor and sensory laterality of horses in agonistic and stressful situations. Here we examine sensory laterality in affiliative interactions within four groups of domestic horses and ponies (N = 31), living in stable social groups, housed at a single complex close to Vienna, Austria, and demonstrate for the first time a significant population preference for the left side in affiliative approaches and interactions. No effects were observed for gender, rank, sociability, phenotype, group, or age. Our results suggest that right hemisphere specialization in horses is not limited to the processing of stressful or agonistic situations, but rather appears to be the norm for processing in all social interactions, as has been demonstrated in other species including chicks and a range of vertebrates. In domestic horses, hemispheric specialization for sensory input appears not to be based on a designation of positive versus negative, but more on the perceived need to respond quickly and appropriately in any given situation.
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Schuetz, A., Farmer, K., & Krueger, K. (2017). Social learning across species: horses (Equus caballus) learn from humans by observation. Anim. Cogn., 20(3), 567–573.
Abstract: This study examines whether horses can learn by observing humans, given that they identify individual humans and orientate on the focus of human attention. We tested 24 horses aged between 3 and 12. Twelve horses were tested on whether they would learn to open a feeding apparatus by observing a familiar person. The other 12 were controls and received exactly the same experimental procedure, but without a demonstration of how to operate the apparatus. More horses from the group with demonstration (8/12) reached the learning criterion of opening the feeder twenty times consecutively than horses from the control group (2/12), and younger horses seemed to reach the criterion more quickly. Horses not reaching the learning criteria approached the human experimenters more often than those that did. The results demonstrate that horses learn socially across species, in this case from humans.
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Baragli, P., Scopa, C., Maglieri, V., & Palagi, E. (2021). If horses had toes: demonstrating mirror self recognition at group level in Equus caballus. Anim. Cogn., .
Abstract: Mirror self-recognition (MSR), investigated in primates and recently in non-primate species, is considered a measure of self-awareness. Nowadays, the only reliable test for investigating MSR potential skills consists in the untrained response to a visual body mark detected using a reflective surface. Here, we report the first evidence of MSR at group level in horses, by facing the weaknesses of methodology present in a previous pilot study. Fourteen horses were used in a 4-phases mirror test (covered mirror, open mirror, invisible mark, visible colored mark). After engaging in a series of contingency behaviors (looking behind the mirror, peek-a-boo, head and tongue movements), our horses used the mirror surface to guide their movements towards their colored cheeks, thus showing that they can recognize themselves in a mirror. The analysis at the group level, which 'marks' a turning point in the analytical technique of MSR exploration in non-primate species, showed that horses spent a longer time in scratching their faces when marked with the visible mark compared to the non-visible mark. This finding indicates that horses did not see the non-visible mark and that they did not touch their own face guided by the tactile sensation, suggesting the presence of MSR in horses. Although a heated debate on the binary versus gradualist model in the MSR interpretation exists, recent empirical pieces of evidence, including ours, indicate that MSR is not an all-or-nothing phenomenon that appeared once in phylogeny and that a convergent evolution mechanism can be at the basis of its presence in phylogenetically distant taxa.
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Trösch, M., Pellon, S., Cuzol, F., Parias, C., Nowak, R., Calandreau, L., et al. (2020). Horses feel emotions when they watch positive and negative horse-human interactions in a video and transpose what they saw to real life. Anim. Cogn., 23(4), 643–653.
Abstract: Animals can indirectly gather meaningful information about other individuals by eavesdropping on their third-party interactions. In particular, eavesdropping can be used to indirectly attribute a negative or positive valence to an individual and to adjust one's future behavior towards that individual. Few studies have focused on this ability in nonhuman animals, especially in nonprimate species. Here, we investigated this ability for the first time in domestic horses (Equus caballus) by projecting videos of positive and negative interactions between an unknown human experimenter (a “positive” experimenter or a “negative” experimenter) and an actor horse. The horses reacted emotionally while watching the videos, expressing behavioral (facial expressions and contact-seeking behavior) and physiological (heart rate) cues of positive emotions while watching the positive video and of negative emotions while watching the negative video. This result shows that the horses perceived the content of the videos and suggests an emotional contagion between the actor horse and the subjects. After the videos were projected, the horses took a choice test, facing the positive and negative experimenters in real life. The horses successfully used the interactions seen in the videos to discriminate between the experimenters. They touched the negative experimenter significantly more, which seems counterintuitive but can be interpreted as an appeasement attempt, based on the existing literature. This result suggests that horses can indirectly attribute a valence to a human experimenter by eavesdropping on a previous third-party interaction with a conspecific.
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Bernauer, K., Kollross, H., Schuetz, A., Farmer, K., & Krueger, K. (2020). How do horses (Equus caballus) learn from observing human action? Anim. Cogn., 23, 1–9.
Abstract: A previous study demonstrated that horses can learn socially from observing humans, but could not draw any conclusions about the social learning mechanisms. Here we develop this by showing horses four different human action sequences as demonstrations of how to press a button to open a feed box. We tested 68 horses aged between 3 and 12 years. 63 horses passed the habituation phase and were assigned either to the group Hand Demo (N = 13) for which a kneeling person used a hand to press the button, Head Demo (N = 13) for which a kneeling person used the head, Mixed Demo (N = 12) for which a squatting person used both head and hand, Foot Demo (N = 12) in which a standing person used a foot, or No Demo (N = 13) in which horses did not receive a demonstration. 44 horses reached the learning criterion of opening the feeder twenty times consecutively, 40 of these were 75% of the Demo group horses and four horses were 31% of the No Demo group horses. Horses not reaching the learning criterion approached the human experimenters more often than those who did. Significantly more horses used their head to press the button no matter which demonstration they received. However, in the Foot Demo group four horses consistently preferred to use a hoof and two switched between hoof and head use. After the Mixed Demo the horses' actions were more diverse. The results indicate that only a few horses copy behaviours when learning socially from humans. A few may learn through observational conditioning, as some appeared to adapt to demonstrated actions in the course of reaching the learning criterion. Most horses learn socially through enhancement, using humans to learn where, and which aspect of a mechanism has to be manipulated, and by applying individual trial and error learning to reach their goal.
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Giljov, A., Malashichev, Y., & Karenina, K. (2019). What do wild saiga antelopes tell us about the relative roles of the two brain hemispheres in social interactions? Anim. Cogn., .
Abstract: Two brain hemispheres are unequally involved in the processing of social stimuli, as demonstrated in a wide range of vertebrates. A considerable number of studies have shown the right hemisphere advantage for social processing. At the same time, an approach-withdrawal hypothesis, mainly based on experimental evidence, proposes the involvement of both brain hemispheres according to approach and withdrawal motivation. The present study aimed to test the relative roles of the two hemispheres in social responses displayed in a natural context. Visual biases, implicating hemispheric lateralization, were estimated in the social interactions of saiga antelope in the wild. In individually identified males, the left/right visual field use during approach and withdrawal responses was recorded based on the lateral head/body position, relative to the conspecific. Lateralized approach responses were investigated in three types of interactions, with left visual field bias found for chasing a rival, no bias--for attacking a rival, and right visual field bias--for pursuing a female. In two types of withdrawal responses, left visual field bias was found for retreating after fighting, while no bias was evident in fight rejecting. These findings demonstrate that neither the right hemisphere advantage nor the approach-withdrawal distinction can fully explain the patterns of lateralization observed in social behaviour. It is clear that both brain hemispheres play significant roles in social responses, while their relative contribution is likely determined by a complex set of motivational and emotional factors rather than a simple dichotomous distinction such as, for example, approach versus withdrawal motivation.
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Liedtke, J., & Schneider, J. M. (2017). Social makes smart: rearing conditions affect learning and social behaviour in jumping spiders. Anim. Cogn., 20(6), 1093–1106.
Abstract: There is a long-standing debate as to whether social or physical environmental aspects drive the evolution and development of cognitive abilities. Surprisingly few studies make use of developmental plasticity to compare the effects of these two domains during development on behaviour later in life. Here, we present rearing effects on the development of learning abilities and social behaviour in the jumping spider Marpissa muscosa. These spiders are ideally suited for this purpose because they possess the ability to learn and can be reared in groups but also in isolation without added stress. This is a critical but rarely met requirement for experimentally varying the social environment to test its impact on cognition. We split broods of spiders and reared them either in a physically or in a socially enriched environment. A third group kept under completely deprived conditions served as a 'no-enrichment' control. We tested the spiders' learning abilities by using a modified T-maze. Social behaviour was investigated by confronting spiders with their own mirror image. Results show that spiders reared in groups outperform their conspecifics from the control, i.e. 'no-enrichment', group in both tasks. Physical enrichment did not lead to such an increased performance. We therefore tentatively suggest that growing up in contact with conspecifics induces the development of cognitive abilities in this species.
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Riley, J. L., Noble, D. W. A., Byrne, R. W., & Whiting, M. J. (2017). Does social environment influence learning ability in a family-living lizard? Anim. Cogn., 20(3), 449–458.
Abstract: Early developmental environment can have profound effects on individual physiology, behaviour, and learning. In birds and mammals, social isolation during development is known to negatively affect learning ability; yet in other taxa, like reptiles, the effect of social isolation during development on learning ability is unknown. We investigated how social environment affects learning ability in the family-living tree skink (Egernia striolata). We hypothesized that early social environment shapes cognitive development in skinks and predicted that skinks raised in social isolation would have reduced learning ability compared to skinks raised socially. Offspring were separated at birth into two rearing treatments: (1) raised alone or (2) in a pair. After 1 year, we quantified spatial learning ability of skinks in these rearing treatments (N = 14 solitary, 14 social). We found no effect of rearing treatment on learning ability. The number of skinks to successfully learn the task, the number of trials taken to learn the task, the latency to perform the task, and the number of errors in each trial did not differ between isolated and socially reared skinks. Our results were unexpected, yet the facultative nature of this species' social system may result in a reduced effect of social isolation on behaviour when compared to species with obligate sociality. Overall, our findings do not provide evidence that social environment affects development of spatial learning ability in this family-living lizard.
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Schwartz, L. P., Silberberg, A., Casey, A. H., Kearns, D. N., & Slotnick, B. (2017). Does a rat release a soaked conspecific due to empathy? Anim. Cogn., 20(2), 299–308.
Abstract: In Experiment 1, rats choosing in an E maze preferred to release a rat standing in a pool of water to dry ground over a rat already standing on dry ground. Five additional experiments showed that the choosing rat's preference for releasing the wet rat was maintained by two separable outcomes: (1) the social contact offered by the released rat and (2) the reinforcing value of proximity to a pool of water. These results call into question Sato et al.'s (Anim Cogn 18:1039-1047, 2015) claim to have demonstrated that a rat's releasing of a wet rat to dry ground is empathically motivated.
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Whalen, A., Cownden, D., & Laland, K. (2015). The learning of action sequences through social transmission. Anim. Cogn., 18(5), 1093–1103.
Abstract: Previous empirical work on animal social learning has found that many species lack the ability to learn entire action sequences solely through reliance on social information. Conversely, acquiring action sequences through asocial learning can be difficult due to the large number of potential sequences arising from even a small number of base actions. In spite of this, several studies report that some primates use action sequences in the wild. We investigate how social information can be integrated with asocial learning to facilitate the learning of action sequences. We formalize this problem by examining how learners using temporal difference learning, a widely applicable model of reinforcement learning, can combine social cues with their own experiences to acquire action sequences. The learning problem is modeled as a Markov decision process. The learning of nettle processing by mountain gorillas serves as a focal example. Through simulations, we find that the social facilitation of component actions can combine with individual learning to facilitate the acquisition of action sequences. Our analysis illustrates that how even simple forms of social learning, combined with asocial learning, generate substantially faster learning of action sequences compared to asocial processes alone, and that the benefits of social information increase with the length of the action sequence and the number of base actions.
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