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Collery, L. (1974). Observations of equine animals under farm and feral conditions. Equine Vet J, 6(4), 170–173.
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Kiley, M. (1972). The vocalizations of ungulates, their causation and function. Z. Tierpsychol., 31(2), 171–222.
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Devenport, J. A., Patterson, M. R., & Devenport, L. D. (2005). Dynamic averaging and foraging decisions in horses (Equus callabus). J. Comp. Psychol., 119(3), 352–358.
Abstract: The variability of most environments taxes foraging decisions by increasing the uncertainty of the information available. One solution to the problem is to use dynamic averaging, as do some granivores and carnivores. Arguably, the same strategy could be useful for grazing herbivores, even though their food renews and is more homogeneously distributed. Horses (Equus callabus) were given choices between variable patches after short or long delays. When patch information was current, horses returned to the patch that was recently best, whereas those without current information matched choices to the long-term average values of the patches. These results demonstrate that a grazing species uses dynamic averaging and indicate that, like granivores and carnivores, they can use temporal weighting to optimize foraging decisions.
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Jansen, T., Forster, P., Levine, M. A., Oelke, H., Hurles, M., Renfrew, C., et al. (2002). Mitochondrial DNA and the origins of the domestic horse. Proc. Natl. Acad. Sci. U.S.A., 99(16), 10905–10910.
Abstract: The place and date of the domestication of the horse has long been a matter for debate among archaeologists. To determine whether horses were domesticated from one or several ancestral horse populations, we sequenced the mitochondrial D-loop for 318 horses from 25 oriental and European breeds, including American mustangs. Adding these sequences to previously published data, the total comes to 652, the largest currently available database. From these sequences, a phylogenetic network was constructed that showed that most of the 93 different mitochondrial (mt)DNA types grouped into 17 distinct phylogenetic clusters. Several of the clusters correspond to breeds and/or geographic areas, notably cluster A2, which is specific to Przewalski's horses, cluster C1, which is distinctive for northern European ponies, and cluster D1, which is well represented in Iberian and northwest African breeds. A consideration of the horse mtDNA mutation rate together with the archaeological timeframe for domestication requires at least 77 successfully breeding mares recruited from the wild. The extensive genetic diversity of these 77 ancestral mares leads us to conclude that several distinct horse populations were involved in the domestication of the horse.
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Hall, C. A., Cassaday, H. J., & Derrington, A. M. (2003). The effect of stimulus height on visual discrimination in horses. J. Anim Sci., 81(7), 1715–1720.
Abstract: This study investigated the effect of stimulus height on the ability of horses to learn a simple visual discrimination task. Eight horses were trained to perform a two-choice, black/white discrimination with stimuli presented at one of two heights: ground level or at a height of 70 cm from the ground. The height at which the stimuli were presented was alternated from one session to the next. All trials within a single session were presented at the same height. The criterion for learning was four consecutive sessions of 70% correct responses. Performance was found to be better when stimuli were presented at ground level with respect to the number of trials taken to reach the criterion (P < 0.05), percentage of correct first choices (P < 0.01), and repeated errors made (P < 0.01). Thus, training horses to carry out tasks of visual discrimination could be enhanced by placing the stimuli on the ground. In addition, the results of the present study suggest that the visual appearance of ground surfaces is an important factor in both horse management and training.
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Harman, A. M., Moore, S., Hoskins, R., & Keller, P. (1999). Horse vision and an explanation for the visual behaviour originally explained by the 'ramp retina'. Equine Vet J, 31(5), 384–390.
Abstract: Here we provide confirmation that the 'ramp retina' of the horse, once thought to result in head rotating visual behaviour, does not exist. We found a 9% variation in axial length of the eye between the streak region and the dorsal periphery. However, the difference was in the opposite direction to that proposed for the 'ramp retina'. Furthermore, acuity in the narrow, intense visual streak in the inferior retina is 16.5 cycles per degree compared with 2.7 cycles per degree in the periphery. Therefore, it is improbable that the horse rotates its head to focus onto the peripheral retina. Rather, the horse rotates the nose up high to observe distant objects because binocular overlap is oriented down the nose, with a blind area directly in front of the forehead.
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Mills, D. S. (1998). Applying learning theory to the management of the horse: the difference between getting it right and getting it wrong. Equine Vet J Suppl, (27), 44–48.
Abstract: Horses constantly modify their behaviour as a result of experience. This involves the creation of an association between events or stimuli. The influence of people on the modification and generation of certain behaviour patterns extends beyond the intentional training of the horse. The impact of any action depends on how it is perceived by the horse, rather than the motive of the handler. Negative and positive reinforcement increase the probability of specific behaviours recurring i.e. strengthen the association between events, whereas punishment reduces the probable recurrence of a behaviour without providing specific information about the desired alternative. In this paper the term 'punishers' is used to refer to the physical aids, such as a whip or crop, which may be used to bring about the process of punishment. However, if their application ceases when a specific behaviour occurs they may negatively reinforce that action. Intended 'punishers' may also be rewarding (e.g. for attention seeking behaviour). Therefore, contingency factors (which define the relationship between stimuli, such as the level of reinforcement), contiguity factors (which describe the proximity of events in space or time) and choice of reinforcing stimuli are critical in determining the rate of learning. The many problems associated with the application of punishment in practice lead to confusion by both horse and handler and, possibly, abuse of the former. Most behaviour problems relate to handling and management of the horse and can be avoided or treated with a proper analysis of the factors influencing the behaviour.
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Cooper, J. J. (1998). Comparative learning theory and its application in the training of horses. Equine Vet J Suppl, (27), 39–43.
Abstract: Training can best be explained as a process that occurs through stimulus-response-reinforcement chains, whereby animals are conditioned to associate cues in their environment, with specific behavioural responses and their rewarding consequences. Research into learning in horses has concentrated on their powers of discrimination and on primary positive reinforcement schedules, where the correct response is paired with a desirable consequence such as food. In contrast, a number of other learning processes that are used in training have been widely studied in other species, but have received little scientific investigation in the horse. These include: negative reinforcement, where performance of the correct response is followed by removal of, or decrease in, intensity of a unpleasant stimulus; punishment, where an incorrect response is paired with an undesirable consequence, but without consistent prior warning; secondary conditioning, where a natural primary reinforcer such as food is closely associated with an arbitrary secondary reinforcer such as vocal praise; and variable or partial conditioning, where once the correct response has been learnt, reinforcement is presented according to an intermittent schedule to increase resistance to extinction outside of training.
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Ishida, N., Hirano, T., & Mukoyama, H. (1994). Detection of aberrant alleles in the D-loop region of equine mitochondrial DNA by single-strand conformation polymorphism (SSCP) analysis. Anim Genet, 25(4), 287.
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Dougherty, D. M., & Lewis, P. (1991). Stimulus generalization, discrimination learning, and peak shift in horses. J Exp Anal Behav, 56(1), 97–104.
Abstract: Using horses, we investigated three aspects of the stimulus control of lever-pressing behavior: stimulus generalization, discrimination learning, and peak shift. Nine solid black circles, ranging in size from 0.5 in. to 4.5 in. (1.3 cm to 11.4 cm) served as stimuli. Each horse was shaped, using successive approximations, to press a rat lever with its lip in the presence of a positive stimulus, the 2.5-in. (6.4-cm) circle. Shaping proceeded quickly and was comparable to that of other laboratory organisms. After responding was maintained on a variable-interval 30-s schedule, stimulus generalization gradients were collected from 2 horses prior to discrimination training. During discrimination training, grain followed lever presses in the presence of a positive stimulus (a 2.5-in circle) and never followed lever presses in the presence of a negative stimulus (a 1.5-in. [3.8-cm] circle). Three horses met a criterion of zero responses to the negative stimulus in fewer than 15 sessions. Horses given stimulus generalization testing prior to discrimination training produced symmetrical gradients; horses given discrimination training prior to generalization testing produced asymmetrical gradients. The peak of these gradients shifted away from the negative stimulus. These results are consistent with discrimination, stimulus generalization, and peak-shift phenomena observed in other organisms.
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