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Acuna, B. D., Sanes, J. N., & Donoghue, J. P. (2002). Cognitive mechanisms of transitive inference. Exp Brain Res, 146(1), 1–10.
Abstract: We examined how the brain organizes interrelated facts during learning and how the facts are subsequently manipulated in a transitive inference (TI) paradigm (e.g., if A<B and B<C, then A<C). This task determined features such as learned facts and behavioral goals, but the learned facts could be organized in any of several ways. For example, if one learns a list by operating on paired items, the pairs may be stored individually as separate facts and reaction time (RT) should decrease with learning. Alternatively, the pairs may be stored as a single, unified list, which may yield a different RT pattern. We characterized RT patterns that occurred as participants learned, by trial and error, the predetermined order of 11 shapes. The task goal was to choose the shape occurring closer to the end of the list, and feedback about correctness was provided during this phase. RT increased even as its variance decreased during learning, suggesting that the learnt knowledge became progressively unified into a single representation, requiring more time to manipulate as participants acquired relational knowledge. After learning, non-adjacent (NA) list items were presented to examine how participants reasoned in a TI task. The task goal also required choosing from each presented pair the item occurring closer to the list end, but without feedback. Participants could solve the TI problems by applying formal logic to the previously learnt pairs of adjacent items; alternatively, they could manipulate a single, unified representation of the list. Shorter RT occurred for NA pairs having more intervening items, supporting the hypothesis that humans employ unified mental representations during TI. The response pattern does not support mental logic solutions of applying inference rules sequentially, which would predict longer RT with more intervening items. We conclude that the brain organizes information in such a way that reflects the relations among the items, even if the facts were learned in an arbitrary order, and that this representation is subsequently used to make inferences.
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Christensen, J. W., Ahrendt, L. P., Lintrup, R., Gaillard, C., Palme, R., & Malmkvist, J. (2012). Does learning performance in horses relate to fearfulness, baseline stress hormone, and social rank? In Applied Animal Behaviour Science (Vol. 140, pp. 44–52).
Abstract: The ability of horses to learn and remember new tasks is fundamentally important for their use by humans. Fearfulness may, however, interfere with learning, because stimuli in the environment can overshadow signals from the rider or handler. In addition, prolonged high levels of stress hormones can affect neurons within the hippocampus; a brain region central to learning and memory. In a series of experiments, we aimed to investigate the link between performance in two learning tests, the baseline level of stress hormones, measured as faecal cortisol metabolites (FCM), fearfulness, and social rank. Twenty-five geldings (2 or 3 years old) pastured in one group were included in the study. The learning tests were performed by professional trainers and included a number of predefined stages during which the horses were gradually trained to perform exercises, using either negative (NR) or positive reinforcement (PR). Each of the learning tests lasted 3 days; 7min/horse/day. The NR test was repeated in a novel environment. Performance, measured as final stage in the training programme, and heart rate (HR) were recorded. Faeces were collected on four separate days where the horses had been undisturbed at pasture for 48h. Social rank was determined through observations of social interactions during feeding. The fear test was a novel object test during which behaviour and HR were recorded. Performance in the NR and PR learning tests did not correlate. In the NR test, there was a significant, negative correlation between performance and HR in the novel environment (rS=-0.66, P<0.001, i.e. nervous horses had reduced performance), whereas there was no such correlation in the home environment (both NR and PR). Behavioural reactions in the fear test correlated significantly with performance in the NR test in the novel environment (e.g. object alertness and final stage: rS=-0.43, P=0.04), suggesting that performance under unfamiliar, stressful conditions may be predicted by behavioural responses in a fear test. There was a negative correlation between social rank and baseline stress hormones (rS=-0.43, P=0.04), i.e. high rank corresponded to low FCM concentrations, whereas neither rank nor FCM correlated with fearfulness or learning performance. We conclude that performance under stressful conditions is affected by activation of the sympathetic nervous system during training and related to behavioural responses in a standardised fear test. Learning performance in the home environment, however, appears unrelated to fearfulness, social rank and baseline FCM levels.
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Ahrendt, L. P., Christensen, J. W., & Ladewig, J. (2012). The ability of horses to learn an instrumental task through social observation. In Applied Animal Behaviour Science (Vol. 139, pp. 105–113).
Abstract: The ability of horses to learn through social observation may ease the implementation of new management systems, because the use of automatic feeders etc. by naive horses could be facilitated by observation of experienced horses. However, previous studies found no documentation for observational learning abilities in horses. This study aimed to investigate the ability of horses to learn an instrumental task from a familiar conspecific when social interaction was allowed during the demonstration. Two similar experiments were performed. In the first experiment, Observer horses (n=11) participated in ten successive demonstrations, where a trained Demonstrator opened an operant device by pushing a sliding lid aside with the muzzle in order to obtain a food reward. Immediately after the demonstrations the Observer horses were given the opportunity to operate the device alone. Control horses (n=11) were aware that the device contained food but were presented to the operant device without demonstration of the task. The learning criterion was at least two openings. Accomplishment of and latency to accomplish the learning criterion, and investigative behaviour towards the operant device were recorded. Five Observers and one Control, out of the eleven horses in each treatment group, accomplished the learning criterion. Even though this presents a high odds ratio (OR) in favour of the Observer treatment (OR=7.6), there was no significant difference between the treatment groups (P=0.15). Analysis of investigative behaviour showed, however, that the demonstrations increased the motivation of the Observer horses to investigate the device. Subsequently, a similar experiment was performed in a practical setting with 44 test horses (mixed age, gender and breed). We used the same operant device and the same number and type of demonstrations, although the horses were held on a loose rope to minimise aggression. In this second experiment, six of 23 Observer horses and five of 21 Control horses learned the instrumental task, representing no influence of the demonstration. Thus, this study did not demonstrate an ability of horses to learn an instrumental task through observation.
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Beckers, T., Miller, R. R., De Houwer, J., & Urushihara, K. (2006). Reasoning rats: forward blocking in Pavlovian animal conditioning is sensitive to constraints of causal inference. J Exp Psychol Gen, 135(1), 92–102.
Abstract: Forward blocking is one of the best-documented phenomena in Pavlovian animal conditioning. According to contemporary associative learning theories, forward blocking arises directly from the hardwired basic learning rules that govern the acquisition or expression of associations. Contrary to this view, here the authors demonstrate that blocking in rats is flexible and sensitive to constraints of causal inference, such as violation of additivity and ceiling considerations. This suggests that complex cognitive processes akin to causal inferential reasoning are involved in a well-established Pavlovian animal conditioning phenomenon commonly attributed to the operation of basic associative processes.
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Zentall, T. R. (2002). A cognitive behaviorist approach to the study of animal behavior. J Gen Psychol, 129(4), 328–363.
Abstract: Traditional psychological approaches to animal learning and behavior have involved either the atheoretical behaviorist approach proposed by B. F. Skinner (1938), in which input-output relations are described in response to environmental manipulations, or the theoretical behaviorist approach offered by C. L Hull (1943), in which associations mediated by several hypothetical constructs and intervening variables are formed between stimuli and responses. Recently, the application of a cognitive behaviorist approach to animal learning and behavior has been found to have considerable value as a research tool. This perspective has grown out of E. C. Tolman's cognitive approach to learning in which behavior is mediated by mechanisms that are not directly observable but can be inferred from the results of critical experiments. In the present article, the author presents several examples of the successful application of the cognitive behaviorist approach. In each case, the experiments have been designed to distinguish between more traditional mechanisms and those mediated by hypothesized internal representations. These examples were selected because the evidence suggests that some form of active cognitive organization is needed to account for the behavioral results.
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Zentall, T. R., & Riley, D. A. (2000). Selective attention in animal discrimination learning. J Gen Psychol, 127(1), 45–66.
Abstract: The traditional approach to the study of selective attention in animal discrimination learning has been to ask if animals are capable of the central selective processing of stimuli, such that certain aspects of the discriminative stimuli are partially or wholly ignored while their relationships to each other, or other relevant stimuli, are processed. A notable characteristic of this research has been that procedures involve the acquisition of discriminations, and the issue of concern is whether learning is selectively determined by the stimulus dimension defined by the discriminative stimuli. Although there is support for this kind of selective attention, in many cases, simpler nonattentional accounts are sufficient to explain the results. An alternative approach involves procedures more similar to those used in human information-processing research. When selective attention is studied in humans, it generally involves the steady state performance of tasks for which there is limited time allowed for stimulus input and a relatively large amount of relevant information to be processed; thus, attention must be selective or divided. When this approach is applied to animals and alternative accounts have been ruled out, stronger evidence for selective or divided attention in animals has been found. Similar processes are thought to be involved when animals search more natural environments for targets. Finally, an attempt is made to distinguish these top-down attentional processes from more automatic preattentional processes that have been studied in humans and other animals.
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Gabor, V., & Gerken, M. (2010). Horses use procedural learning rather than conceptual learning to solve matching to sample. Appl. Anim. Behav. Sci., 126(3-4), 119–124.
Abstract: Research into higher cognitive abilities of the horse may be limited by developing the adequate experimental design. In this study four pony mares between 8 and 19 years old were included. Three of them reached the criterion to be tested in a new design of matching to sample using a black circle and a cross as visual cues attached to an apparatus. The attention was directed to the question of whether the animals are able to concept formation in a given time period or if their decisions depend on other cues or strategies. After familiarization to the testing area and the test procedure, the animals were given 27 sessions of 20 trials each during 14 weeks. While there was no preference for one of the stimuli used, horses showed a significant left sidedness. None of the mares reached the learning criterion of 80% correct answers in one session. However, the ponies showed procedural learning based on correction runs that were given between incorrect decisions, by then selecting the correct stimulus on the other side of the apparatus. This learning type arose in three individuals in session four, six and eleven, respectively. It is concluded that discrimination tasks may be biased by the involvement of unexpected learning strategies, which complicates the interpretation of such tests and may even mask possible conceptualization capabilities.
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Brubaker, L., & Udell, M. A. R. (2016). Cognition and learning in horses (Equus caballus): What we know and why we should ask more. Behavioural Processes, 126, 121–131.
Abstract: Abstract Horses (Equus caballus) have a rich history in their relationship with humans. Across different cultures and eras they have been utilized for work, show, cultural rituals, consumption, therapy, and companionship and continue to serve in many of these roles today. As one of the most commonly trained domestic animals, understanding how horses learn and how their relationship with humans and other horses impacts their ability to learn has implications for horse welfare, training, husbandry and management. Given that unlike dogs and cats, domesticated horses have evolved from prey animals, the horse-human relationship poses interesting and unique scientific questions of theoretical value. There is still much to be learned about the cognition and behaviour of horses from a scientific perspective. This review explores current research within three related areas of horse cognition: human-horse interactions, social learning and independent learning in horses. Research on these topics is summarized and suggestions for future research are provided.
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Lansade, L., & Simon, F. (2010). Horses' learning performances are under the influence of several temperamental dimensions. Appl. Anim. Behav. Sci., 125(1-2), 30–37.
Abstract: Learning performances are influenced by many factors, not only breed, age and sex, but also temperament. The purpose of this study was to understand how different temperamental dimensions affect the learning performance of horses, Equus caballus. First, we carried out a series of behavioural tests on 36 Welsh ponies aged 5-7 years to measure five temperamental dimensions: fearfulness (novel area test and surprise test), gregariousness (social isolation test), reactivity to humans (passive human test), tactile sensitivity (von Frey filament test) and activity level (evaluation of locomotor activity during all the tests). We then presented them with two learning tasks (avoidance and backwards-forwards tasks). In the avoidance task they had to learn to jump over a fence when they heard a sound associated with an aversive stimulus (puff of air). In the backwards-forwards task they had to walk forwards or move backwards in response to a tactile or vocal command to obtain a food reward. There was no correlation between performances on the two learning tasks, indicating that learning ability is task-dependent. However, correlations were found between temperamental data and learning performance (Spearman correlations). The ponies that performed the avoidance task best were the most fearful and the most active ones. For instance, the number of trials required to perform 5 consecutive correct responses (learning criterion) was correlated with the variables aimed at measuring fearfulness (way of crossing a novel area: rs = -0.41, P = 0.01 and time to start eating again after a surprise effect: rs = -0.33, P = 0.05) and activity level (frequency of trotting during all the tests: rs = -0.40, P = 0.02). The animals that performed the backwards-forwards task best were the ones that were the least fearful and the most sensitive. For instance, the learning criterion (corresponding to the number of trials taken to achieve five consecutive correct responses) was correlated with the variables aimed at measuring fearfulness (latency to put one foot on the area: rs = 0.43, P = 0.01; way of crossing a novel area: rs = 0.31, P = 0.06; and time to start eating again after a surprise effect: rs = 0.43, P = 0.009) and tactile sensitivity (response to von Frey filaments: rs = -0.44, P = 0.008). This study revealed significant links between temperament and learning abilities that are highly task-dependent.
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Gibson, B. M., Shettleworth, S. J., & McDonald, R. J. (2001). Finding a goal on dry land and in the water: differential effects of disorientation on spatial learning. Behav. Brain. Res., 123(1), 103–111.
Abstract: Two previous studies, Martin et al. (J. Exp. Psychol. Anim. Behav. Process. 23 (1997) 183) and Dudchenko et al. (J. Exp. Psychol. Anim. Behav. Process. 23 (1997) 194), report that, compared to non-disoriented controls, rats disoriented before testing were disrupted in their ability to learn the location of a goal on a dry radial-arm maze task, but that both groups learned at the same rate in the Morris water maze. However, the radial-arm maze task was much more difficult than the water maze. In the current set of experiments, we examined the performance of control and disoriented rats on more comparable dry land and water maze tasks. Compared to non-disoriented rats, rats that were disoriented before testing were significantly impaired in locating a goal in a circular dry arena, but not a water tank. The results constrain theoretical explanations for the differential effects of disorientation on different spatial tasks.
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