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Brust, V., & Guenther, A. (2015). Domestication effects on behavioural traits and learning performance: comparing wild cavies to guinea pigs. Anim. Cogn., 18(1), 99–109.
Abstract: The domestication process leads to a change in behavioural traits, usually towards individuals that are less attentive to changes in their environment and less aggressive. Empirical evidence for a difference in cognitive performance, however, is scarce. Recently, a functional linkage between an individual's behaviour and cognitive performance has been proposed in the framework of animal personalities via a shared risk-reward trade-off. Following this assumption, bolder and more aggressive animals (usually the wild form) should learn faster. Differences in behaviour may arise during ontogeny due to individual experiences or represent adaptations that occurred over the course of evolution. Both might singly or taken together account for differences in cognitive performance between wild and domestic lineages. To test for such possible linkages, we compared wild cavies and domestic guinea pigs, both kept in a university stock for more than 30 years under highly comparable conditions. Animals were tested in three behavioural tests as well as for initial and reversal learning performance. Guinea pigs were less bold and aggressive than their wild congeners, but learnt an association faster. Additionally, the personality structure was altered during the domestication process. The most likely explanation for these findings is that a shift in behavioural traits and their connectivity led to an altered cognitive performance. A functional linkage between behavioural and cognitive traits seems to exist in the proposed way only under natural selection, but not in animals that have been selected artificially over centuries.
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Gabor, V., & Gerken, M. (2014). Shetland ponies (Equus caballus) show quantity discrimination in a matching-to-sample design. Anim. Cogn., 17(6), 1233–1243.
Abstract: Numerical competence is one of the aspects of animal cognition with a long history of research interest, but few results are available for the horse. In the present study, we investigated the ability of three Shetland ponies to discriminate between different quantities of geometric symbols presented on a computer screen in a matching-to-sample arrangement. In Experiment 1, the ponies had to relate two similar quantities to another, paired in contrasts (1 vs. 2, 3 vs. 4 and 4 vs. 5) of the same stimulus (dot). Specific pairs of quantities (all differing by one) of up to five different geometrical symbols were displayed in Experiment 2. In each session, both quantities (more and less) were used as sample in such a way that each of the two quantities presented in one test served as positive and as negative stimulus, respectively. The three Shetland ponies were able to discriminate between the given quantities of dots by showing more than 80 % correct responses in two consecutive sessions. Only one of the ponies distinguished different shapes of geometric symbols at a level of 4 versus 5 items. The results show that all ponies were capable of visual quantity discrimination in the present matching-to-sample design, but task solving seemed more difficult when quantities were composed of heterogeneous stimuli. The present results confirm our hypothesis that the ponies based their decision on the matching concept of sameness and were not biased by a spontaneous preference for higher quantities.
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Miyata, H., Gajdon, G. K., Huber, L., & Fujita, K. (2011). How do keas (Nestor notabilis) solve artificial-fruit problems with multiple locks? Anim. Cogn., 14(1), 45–58.
Abstract: Keas, a species of parrots from New Zealand, are an interesting species for comparative studies of problem solving and cognition because they are known not only for efficient capacities for object manipulation but also for explorative and playful behaviors. To what extent are they efficient or explorative, and what cognitive abilities do they use? We examined how keas would solve several versions of artificial-fruit box problems having multiple locks. After training keas to remove a metal rod from over a Plexiglas lid that had to be opened, we exposed the birds to a variety of tasks having two or more locks. We also introduced a preview phase during which the keas had extended opportunity to look at the tasks before the experimenter allowed the birds to solve them, to examine whether the preview phase would facilitate the birds' performance on the tasks. In a large number of tests, the keas showed a strong trend to solve the tasks with no positive effect of previewing the tasks. When the tasks became complex, however, the keas corrected inappropriate responses more quickly when they had had chance to preview the problems than when they had not. The results suggest that the keas primarily used explorative strategies in solving the lock problems but might have obtained some information about the tasks before starting to solve them. This may reflect a good compromise of keas' trial-and-error tendency and their good cognitive ability that result from a selection pressure they have faced in their natural habitat.
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Basile, M., Boivin, S., Boutin, A., Blois-Heulin, C., Hausberger, M., & Lemasson, A. (2009). Socially dependent auditory laterality in domestic horses (Equus caballus). Anim. Cogn., 12, 611–619.
Abstract: Laterality is now known to be an ubiquitous phenomenon among the vertebrates. Particularly, laterality of auditory processing has been demonstrated in a variety of species, especially songbirds and primates. Such a hemispheric specialization has been shown to depend on factors such as sound structure, species specificity and types of stimuli. Much less is known on the possible influence of social familiarity although a few studies suggest such an influence. Here we tested the influence of the degree of familiarity on the laterality of the auditory response in the domestic horse. This species is known for its social system and shows visible reactions to sounds, with one or two ears moving towards a sound source. By comparing such responses to the playback of different conspecific whinnies (group member, neighbor and stranger), we could demonstrate a clear left hemisphere (LH) preference for familiar neighbor calls while no preference was found for group member and stranger calls. Yet, we found an opposite pattern of ear side preference for neighbor versus stranger calls. These results are, to our knowledge, the first to demonstrate auditory laterality in an ungulate species. They open further lines of thought on the influence of the social “value” of calls and the listener's arousal on auditory processing and laterality.
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Hanggi, E. B., & Ingersoll, J. F. (2009). Long-term memory for categories and concepts in horses (Equus caballus). Anim. Cogn., 13(3), 451–462.
Abstract: Three horses (Equus caballus) with a history of performing cognitive tasks including discrimination learning, categorization, and concept use were tested to evaluate their long-term memory (LTM) in three experiments. In addition, use of LCD multi-displays for stimulus presentation was incorporated into cognition testing protocol for the first time with horses. Experiment 1 tested LTM for discrimination learning that originally occurred 6 years earlier. Five sets of stimuli were used and the two horses tested showed no decrement in performance on four of the sets; however, both horses did score below chance on one set. Experiment 2 examined long-term categorization recall 10 years after horses had demonstrated the ability to make stimulus selections based on shared characteristics within a given category. The horse tested for LTM after the decade-long interval immediately and consistently applied the previously learned categorization rule to not only familiar but also novel sets of stimuli. Experiment 3 tested another horse for LTM for a relative size concept. This horse had originally demonstrated concept rule use in order to select stimuli based on their relative size to one another. More than 7 years later and without further training, this horse reliably applied the previously established size concept to both familiar and novel sets of stimuli. These findings are the first reports of long-term categorical and conceptual memory in horses and are consistent with observations of domestic and wild horses, which indicate that behavioral and ecological events may be remembered for long periods of time. These studies also demonstrate the adaptive nature of horses with regard to their ability to generalize over several different testing conditions.
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Gácsi, M., Kara, E., Belényi, B., Topál, J., & Miklósi, Á. (2009). The effect of development and individual differences in pointing comprehension of dogs. Anim. Cogn., 12(3), 471–479.
Abstract: In spite of the rather different procedures actually used in comparative studies to test the ability of different species to rely on the human pointing gesture, there is no debate on the high performance of dogs in such tasks. Very little is known, however, on the course through which they acquire this ability or the probable factors influencing the process. Important developmental questions have remained unsolved and also some methodological concerns should be addressed before we can convincingly argue for one interpretation or another. In this study we tested 180 dogs of different age (from 2 months to adults) to investigate their performance in the human distal momentary pointing gesture. The results, analyzed at both the group and the individual levels, showed no difference in the performance according to age, indicating that in dogs the comprehension of the human pointing may require only very limited and rapid early learning to fully develop. Interestingly, neither the keeping conditions nor the time spent in active interaction with the owner, and not even some special (agility) training for using human visual cues, had significant effect on the success and explained individual differences. The performance of the dogs was rather stable over time: during the 20 trials within a session and even when subsamples of different age were repeatedly tested. Considering that in spite of the general success at the group level, more than half of the dogs were not successful at the individual level, we revealed alternative “decision-making rules” other than following the pointing gesture of the experimenter.
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Fureix, C., Jego, P., Sankey, C., & Hausberger, M. (2009). How horses (Equus caballus) see the world: humans as significant “objects”. Anim. Cogn., 12(4), 643–654.
Abstract: Abstract  This study aimed to determine whether horses have a kind of memory of humans (based on previous interactions), leading to a general significance of humans revealed by their reactions to humans in subsequent interactions. Subjects were 59 adult horses used to interact daily with humans. Three types of behavioural tests involving an unknown experimenter evaluated three possibly different memorized types of human–animal interactions (not work-related, using work-related objects, unfamiliar working task). We also performed standardized observations of routine interactions between each horse and its familiar handler (caretaker). To get a broad overview of the horses’ reactions to humans, we recorded both investigative and aggressive behaviours during the tests, representing respectively a “positive” and a “negative” memory of the relationship. Whereas correlations between tests revealed a general perception of humans as either positive or negative, unusual tests, i.e. that are not usually performed, elicited more positive reactions. Moreover, some horses reacted positively to a motionless person in their box, but negatively when this same person approached them, for example for halter fitting. Overall, aggressive reactions were more reliable indicators of the relationship than positive reactions, both between tests and between familiar and unfamiliar humans. Our results also show generalization of the perception of humans. These results support our hypothesis that perception of humans by horses may be based on experience, i.e. repeated interactions. Altogether, our results support the hypothesis that horses can form a memory of humans that impacts their reactions in subsequent interactions.
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Krueger, K., Farmer, K., & Heinze, J. (2014). The effects of age, rank and neophobia on social learning in horses. Anim. Cogn., 17(3), 645–655.
Abstract: Social learning is said to meet the demands of complex environments in which individuals compete over resources and co-operate to share resources. Horses (Equus caballus) were thought to lack social learning skills because they feed on homogenously distributed resources with few reasons for conflict. However, the horse’s social environment is complex, which raises the possibility that its capacity for social transfer of feeding behaviour has been underestimated. We conducted a social learning experiment using 30 socially kept horses of different ages. Five horses, one from each group, were chosen as demonstrators, and the remaining 25 horses were designated observers. Observers from each group were allowed to watch their group demonstrator opening a feeding apparatus. We found that young, low ranking, and more exploratory horses learned by observing older members of their own group, and the older the horse, the more slowly it appeared to learn. Social learning may be an adaptive specialisation to the social environment. Older animals may avoid the potential costs of acquiring complex and potentially disadvantageous feeding behaviours from younger group members. We argue that horses show social learning in the context of their social ecology, and that research procedures must take such contexts into account. Misconceptions about the horse’s sociality may have hampered earlier studies.
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Farmer, K., Krueger, K., & Byrne, R. (2010). Visual laterality in the domestic horse (Equus caballus) interacting with humans. Anim. Cogn., 13, 229–238.
Abstract: Most horses have a side on which they are easier to handle and a direction they favour when working on a circle, and recent studies have suggested a correlation between emotion and visual laterality when horses observe inanimate objects. As such lateralisation could provide important clues regarding the horse’s cognitive processes, we investigated whether horses also show laterality in association with people. We gave horses the choice of entering a chute to left or right, with and without the passive, non-interactive presence of a person unknown to them. The left eye was preferred for scanning under both conditions, but significantly more so when a person was present. Traditionally, riders handle horses only from the left, so we repeated the experiment with horses specifically trained on both sides. Again, there was a consistent preference for left eye scanning in the presence of a person, whether known to the horses or not. We also examined horses interacting with a person, using both traditionally and bilaterally trained horses. Both groups showed left eye preference for viewing the person, regardless of training and test procedure. For those horses tested under both passive and interactive conditions, the left eye was preferred significantly more during interaction. We suggest that most horses prefer to use their left eye for assessment and evaluation, and that there is an emotional aspect to the choice which may be positive or negative, depending on the circumstances. We believe these results have important practical implications and that emotional laterality should be taken into account in training methods.
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Krueger, K., Flauger, B., Farmer, K., & Maros, K. (2011). Horses (Equus caballus) use human local enhancement cues and adjust to human attention. Anim. Cogn., 14(2), 187–201.
Abstract: This study evaluates the horse (Equus caballus) use of human local enhancement cues and reaction to human attention when making feeding decisions. The superior performance of dogs in observing human states of attention suggests this ability evolved with domestication. However, some species show an improved ability to read human cues through socialization and training. We observed 60 horses approach a bucket with feed in a three-way object-choice task when confronted with (a) an unfamiliar or (b) a familiar person in 4 different situations: (1) squatting behind the bucket, facing the horse (2) standing behind the bucket, facing the horse (3) standing behind the bucket in a back-turned position, gazing away from the horse and (4) standing a few meters from the bucket in a distant, back-turned position, again gazing away from the horse. Additionally, postures 1 and 2 were tested both with the person looking permanently at the horse and with the person alternating their gaze between the horse and the bucket. When the person remained behind the correct bucket, it was chosen significantly above chance. However, when the test person was turned and distant from the buckets, the horses’ performance deteriorated. In the turned person situations, the horses approached a familiar person and walked towards their focus of attention significantly more often than with an unfamiliar person. Additionally, in the squatting and standing person situations, some horses approached the person before approaching the correct bucket. This happened more with a familiar person. We therefore conclude that horses can use humans as a local enhancement cue independently of their body posture or gaze consistency when the persons remain close to the food source and that horses seem to orientate on the attention of familiar more than of unfamiliar persons. We suggest that socialization and training improve the ability of horses to read human cues.
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