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Author |
Mendl, M. |
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Title |
Performing under pressure: stress and cognitive function |
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Journal Article |
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1999 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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65 |
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3 |
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221-244 |
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Stress; Cognition; Attention; Learning; Memory; Welfare |
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The way in which cognitive functioning is affected by stressors is an important area of research for applied ethologists because stress caused by captive conditions may disrupt cognitive processes and lead to welfare and husbandry problems. Such problems may be minimised through an understanding of the links between stress and cognition. The effects of stress on cognitive function have been studied in disciplines ranging from human perceptual psychology to animal neuroscience. The aim of this paper is to provide an introduction to this research, focusing on the effects of stressors on attention, memory formation and memory recall. Findings from such a diverse literature with little apparent inter-disciplinary communication are inevitably complex and often contradictory. Nevertheless, some generalities do emerge. The idea that an inverted U-shaped relationship exists between an individual's state of stress or arousal and its ability to perform a cognitive task effectively, the so-called Yerkes-Dodson law, is commonly encountered. The law has limited explanatory value because it is unlikely that different stressors act on cognitive function via the same intervening, non-specific state. Furthermore, the law only provides a very general description of the relationship between stress and cognitive function. Empirical research on attention and memory processes reveals more specific findings. Stressors appear to cause shifts, lapses and narrowing of attention, and can also influence decision speed. These processes may be viewed as serving an adaptive role helping the animal to search for and scrutinise a source of danger. There is conflicting evidence as to whether hormones involved in the hypothalamic-pituitary-adrenal stress response play a part in these processes. These hormones and those involved in the sympathetic-adrenomedullary stress response do appear to play an important role in memory formation. Low or moderate concentrations of circulating glucocorticoids and catecholamines can enhance memory formation, while excessively high or prolonged elevations of these hormones can lead to memory disruption. The effects of stressors on memory recall are less clear. There is evidence for disruptive effects, and for facilitatory effects indicating state-dependent memory recall; events experienced under conditions of high arousal may be best recalled under similar conditions. Applied ethologists have the opportunity to extend work in this area, which often involves studies of single stressors/stress hormones acting in isolation and limited measures of cognitive function, by focusing on real-life husbandry stressors encountered by captive animals. This will yield fundamental information which also has direct relevance to animal welfare and management issues. |
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Cooper, J.J.; Ashton, C.; Bishop, S.; West, R.; Mills, D.S.; Young, R.J. |
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Clever hounds: social cognition in the domestic dog (Canis familiaris) |
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Journal Article |
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2003 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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81 |
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3 |
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229-244 |
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Domestic dog; Canine; Social cognition; Counting; Theory of mind; Perspective taking |
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This paper reviews the reasons why domestic dogs make good models to investigate cognitive processes related to social living and describes experimental approaches that can be adopted to investigate such processes in dogs. Domestic dogs are suitable models for investigating social cognition skills for three broad reasons. First, dogs originated from wolves, social animals that engage in a number of co-operative behaviours, such as hunting and that may have evolved cognitive abilities that help them predict and interpret the actions of other animals. Second, during domestication dogs are likely to have been selected for mental adaptations for their roles in human society such as herding or companionship. Third, domestic dogs live in a human world and “enculturation” may facilitate the development of relevant mental skills in dogs. Studies of social cognition in animals commonly use experimental paradigms originally developed for pre-verbal human infants. Preferential gaze, for example, can be used as a measure of attention or “surprise” in studies using expectancy violation. This approach has been used to demonstrate simple numerical competence in dogs. Dogs also readily use both conspecific and human social signals (e.g. looking or pointing) as information sources to locate hidden rewards such as food or favourite toys. Such abilities make dogs particularly good models for investigating perspective-taking tasks, where animals are required to discriminate between apparently knowledgeable and apparently ignorant informants. |
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Hanggi, E.B. |
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Discrimination learning based on relative size concepts in horses (Equus caballus) |
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2003 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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83 |
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3 |
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201-213 |
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Horse; Concept; Size transposition; Generalization; Learning; Training |
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This study explored whether or not horses (Equus caballus) could respond to stimuli using a concept based on relative size. In Experiment 1, after learning to respond to the larger of the two stimuli for six sets of two-dimensional (2D) training exemplars, one horse was tested for size transposition that used novel larger and smaller stimuli as well as three-dimensional (3D) objects (5 two-dimensional sets and 5 three-dimensional sets with large, medium, small, and tiny sizes). The horse correctly chose (significantly above chance) the larger of two stimuli regardless of novelty or dimension or combination. In Experiment 2, two additional horses were tested using a subset of the stimuli from Experiment 1. One horse was required to select the larger stimulus--as in Experiment 1--and the other the smaller stimulus. After learning the task, both horses responded correctly to new stimuli and showed size transposition. These results suggest that at least some horses are capable of solving problems based on relative size concepts. Moreover, they are able to generalize across situations that vary from flat, black shapes to objects of different materials and colors including balls, flower pots, and PVC connectors. These findings support earlier research that showed that horses could categorize certain stimuli, and provide new evidence that they are capable of using some form of concept for problem solving. Understanding that horses have more advanced learning abilities than was previously believed should help improve training methods and management. |
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398 |
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Saslow, C.A. |
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Title |
Understanding the perceptual world of horses |
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2002 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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78 |
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2-4 |
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209-224 |
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Horse; Perception; Vision; Olfaction; Touch; Hearing; Pain; Training; Psychophysics; Umwelt |
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From the viewpoint of experimental psychology, there are two problems with our current knowledge of equine perception. The first is that the behavioral and neurophysiological research in this area has enormous gaps, reflecting that this animal is not a convenient laboratory subject. The second is that the horse, having been a close companion to humans for many millennia, entrenched anecdotal wisdom is often hard to separate from scientific fact. Therefore, any summary at present of equine perception has to be provisional. The horse appears to have developed a visual system particularly sensitive to dim light and movement, it may or may not have a weak form of color vision in part of the retina, it has little binocular overlap, and its best acuity is limited to a restricted horizontal band which is aimed primarily by head/neck movements. However, the total field of view is very large. Overall, as would be expected for a prey animal, horse vision appears to have evolved more for detection of predator approach from any angle than for accurate visual identification of stationary objects, especially those seen at a distance. It is likely that, as for most mammals except the primates, horses rely more heavily on their other senses for forming a view of their world. Equine high-frequency hearing extends far above that of humans, but horses may be less able to localize the point of origin of brief sounds. The horse's capacity for chemoreception and its reliance on chemical information for identification may more closely resemble that of the dog than of the human. Its tactile sensitivity is high, and the ability of its brain and body to regulate pain perception appears to be similar to that found in other mammals. There is room for a great deal of future research in both the area of equine perception and sensory-based cognition, but for the present time persons interacting with this animal should be made aware of the importance of the sounds they make, the movements of their bodies, the way they touch the animal, and the odors they emit or carry on their clothing. |
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McLean, A.N. |
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Title |
Short-term spatial memory in the domestic horse |
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2004 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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85 |
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1-2 |
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93-105 |
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Animal cognition; Delayed response; Equine memory; Horse; Object permanence; Temporal cognition |
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This study investigates the ability of horses to recall a feeding event in a two-point choice apparatus. Twelve horses were individually tested whereby they were maintained immobile in a test arena and visually and aurally experienced the delivery of food into one of two feed goals. The horses were then released to make their choice in two experimental contexts: immediate release after experiencing the delivery of food, and release 10 s after food delivery. Each horse performed 40 immediate-release (IR) trials, followed by forty 10-s release trials over a 3-day period. In addition, the same horses were tested 3 months later in the spring with the same number and sequence of trials. Results were analysed by log-linear analysis of frequencies. Results showed that while horses were able to achieve the correct feed goal choice in the immediate-release trials, they were unsuccessful with the 10-s release trials. This suggests that there are limitations in recall abilities in horses, in that they may not possess a prospective type of memory. There are welfare and training implications in these findings concerning the effects of overestimating the mental abilities of horses during training and the effects of delays in reinforcements. |
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403 |
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Author |
Nicol, C. J. |
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Title |
Equine learning: progress and suggestions for future research |
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Journal Article |
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2002 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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78 |
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2-4 |
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193-208 |
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Learning; Horse; Equine; Discrimination; Training |
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Horses are well able to form classical and instrumental associations and so the focus of much recent research has been on the stimulus control of instrumental learning. Horses appear to discriminate using spatial cues more easily than other stimulus features, as indicated both by the speed of initial task acquisition and by the extent to which acquired discriminations can be reversed. Phenomena associated with discrimination learning in laboratory animals, including generalisation and peak shift, have been demonstrated in horses. However, the ability of horses to classify stimuli into categories is more controversial. Although there is some evidence that horses may be able to form categories based on similarities in the physical appearance of different stimuli, there is currently no evidence that they are able to develop abstract concepts. Their performance on social learning tasks has also been poor. Few correlations are observed between the learning ability of individual horses on different tasks, suggesting that it may not be possible to classify individual horses as `good' or `poor' learners. Better learning performance by horses that are naturally calm is probably due to reduced interference in the learning process. Correct handling procedures can lower reactivity levels in horses, and may facilitate learning in some circumstances. Future research on equine learning needs to take into account the complex nature of equine social interaction. Studies on the effects of stress on learning, and on social and spatial cognition, are also particularly needed. |
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405 |
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Marinier, S.L.; Alexander, A.J.; Waring, G.H. |
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Flehmen behaviour in the domestic horse: Discrimination of conspecific odours |
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1988 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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19 |
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3-4 |
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227-237 |
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American Saddlebred horses were used to test the responses of domestic horses to the odours of conspecifics. In all cases the odours were tested in the absence of the donor animal. Thus the test animal's behavioural responses were concentrated on the olfactory stimuli, and possible interference from donor behaviour was eliminated. Stallions were significantly more responsive than mares and geldings. This was shown in both flehmen and sniffing behaviour to urine/vaginal secretions and in sniffing behaviour to faecal samples. Only stallions were used for subsequent tests. Stallions showed no significant differences in response to the odour of urine/vaginal secretions of an oestrus mare from that when she was not in season. Parameters used for analysis of data were frequency, latency and duration of flehmen as well as duration of responsiveness to samples. In testing for differences in odours between individual mares, two methods were used. The stallions differentiated between samples from individual mares. In some cases this differentiation was exhibited when the stallions were merely presented with the two samples in sequence. In other cases statistically significant differences in response to the odours were shown only by simultaneous presentation of the two samples to the test stallion. Parameters used for data analysis were frequency and duration of flehmen and duration of responsiveness. |
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507 |
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Seaman, S.C.; Davidson, H.P.B.; Waran, N.K. |
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How reliable is temperament assessment in the domestic horse (Equus caballus)? |
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Journal Article |
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2002 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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78 |
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2-4 |
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175-191 |
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Temperament assessment; Behavioural tests; Horses; Active and passive copers; Factor analysis |
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Differences in behavioural characteristics between individuals of the same species are often described as being due to the temperament of the individuals. These differences can have enormous implications for welfare with some individuals apparently being able to adapt to environmental challenge more easily than others. Such differences have resulted in animals often being described as either `active' copers, which try to escape from or remove an aversive stimulus, or `passive' copers, which show no outward signs of a situation being aversive, thus, appearing to be unaffected. Tests previously developed to assess the temperament of animals have been criticised for several reasons. Behaviour is often recorded and categorised using methods that are not objective and tests are generally carried out once with no consideration of whether or not behavioural responses are consistent over time. This study takes these factors into account. The behaviour of 33 horses was recorded in three types of test--an arena test, response to a person and response to an object. In order to test whether or not responses were consistent over time, the tests were repeated three times with an average of 9 days between trials. Test results were validated using responses from questionnaires completed by the farm team leader. The data were analysed using an initial principal component analysis (PCA) and factor analysis. The horses were found to behave consistently over the three trials in their responses in the arena test. The responses to the person test and the object test were similar to each other; however, these responses were not consistent over trials. The behaviour in the arena test was unable to be used to make a prediction of behaviour in the person and object tests and vice versa. The responses shown by the horses did not enable them to be categorised as either active or passive copers. Behavioural responses in the tests were not predictive of the response to a startle test (water spray), nor could they be used to predict status or response to being reintroduced to the group after testing. There was no relationship between the responses in the tests and the ratings given by the farm team leader. It was concluded that horses vary widely in their responses to artificial behavioural tests, with only the responses to an open-field arena test being consistent over time, and therefore, the only type of test which can indicate some core factor of temperament. |
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Murphy, J.; Sutherland, A.; Arkins, S. |
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Idiosyncratic motor laterality in the horse |
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2005 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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91 |
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3-4 |
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297-310 |
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Horse; Idiosyncratic motor behaviour; Laterality; Sidedness |
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Idiosyncratic motor behaviour was investigated during four experimental procedures in 40 horses (males = 20, females = 20) to establish if horses revealed evidence of significant right or left laterality. The experimental procedures included (1) detection of the preferred foreleg to initiate movement (walk or trot), (2) obstacle avoidance within a passageway (right or left), (3) obstacle avoidance when ridden and (4) idiosyncratic motor bias when rolling. The influence of the horses' sex on both the direction and the degree of the laterality was explored within and between experimental procedures. The findings showed that the direction, but not the degree of idiosyncratic motor preference in the horses was strongly sex-related. Male horses exhibited significantly more (t = 3.74, d.f. = 79, P < 0.001) left lateralised responses and female horses exhibited significantly more (t = -6.35, d.f. = 79, P < 0.01) right lateralised responses. There was also significant positive correlation (P < 0.05) between four of six possible inter-experimental relationships. The results suggest two discrete trends of laterality associated with the sex of the horse. The primary cause of idiosyncratic motor laterality may be genetically predetermined, influenced by environmental factors or a combination of these two and the current findings may support the development of sex-specific training schedules for the horse. Further, work in this area might assist in defining the mechanisms of brain hemisphere lateralisation and allocation of cognitive function in the horse. |
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Lindberg, A.C.; Kelland, A.; Nicol, C.J. |
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Effects of observational learning on acquisition of an operant response in horses |
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1999 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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61 |
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3 |
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187-199 |
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Horse; Observational learning; Stereotyped behaviour; Operant behaviour; Breed influence; Age influence |
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The effect of observational learning on the acquisition of an operant response was examined in eighteen riding horses and ponies. The test horses were randomly divided into three groups of six and individually exposed to one of three treatments. An additional horse was trained as a demonstrator, to perform the operant response. The observer horses watched either the demonstrator performing the bin-opening response (Group D+B); the demonstrator standing passively (Group D); or the operant bin in the absence of the demonstrator (Group B). Observers had access to and were free to interact with an identical bin during testing. Observers in Groups D+B and D were socially familiar with the demonstrator. Each test horse was tested once a day for 10 days. An ANOVA revealed no significant differences between treatment groups in the number of responses or the time taken to reach the learning criterion. However, there were highly significant differences between breed types, with non-warmbloods performing more bouts of opening the bin and feeding (p=0.02), feeding from the bin sooner (p=0.01) and reaching the criterion for learning sooner than warmbloods (p=0.05). There was also a significant negative linear relationship between horses' ages and time spent investigating the bin, with younger horses performing more investigative behaviour (y=-3.08x+106.86; p=0.02). |
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