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Visser, E. K., van Reenen, C. G., van der Werf, J. T. N., Schilder, M. B. H., Knaap, J. H., Barneveld, A., et al. (2002). Heart rate and heart rate variability during a novel object test and a handling test in young horses. Physiol. Behav., 76(2), 289–296.
Abstract: Forty-one Dutch Warmblood immature horses were used in a study to quantify temperamental traits on the basis of heart rate (HR) and heart rate variability (HRV) measures. Half of the horses received additional training from the age of 5 months onwards; the other half did not. Horses were tested at 9, 10, 21 and 22 months of age in a novel object and a handling test. During the tests, mean HR and two heart variability indices, e.g. standard deviation of beat-to-beat intervals (SDRR) and root mean square of successive beat-to-beat differences (rMSSD), were calculated and expressed as response values to baseline measures. In both tests, horses showed at all ages a significant increase in mean HR and decrease in HRV measures, which suggests a marked shift of the balance of the autonomic nervous system towards a sympathetic dominance. In the novel object test, this shift was more pronounced in horses that had not been trained. Furthermore, statistical analysis showed that the increase in mean HR could not be entirely explained by the physical activity. The additional increase in HR, the nonmotor HR, was more pronounced in the untrained horses compared to the trained. Hence, it is suggested that this nonmotor HR might be due to the level of emotionality. HR variables showed consistency between years, as well as within the second year. These tests bring about a HR response in horses, part of which may indicate a higher level of emotionality; and horses show individual consistency of these HR variables over ages. Therefore, it is concluded that mean HR and HRV measures used with these tests quantify certain aspects of a horse's temperament.
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Hausberger, M., & Muller, C. (2002). A brief note on some possible factors involved in the reactions of horses to humans. Appl. Anim. Behav. Sci., 76(4), 339–344.
Abstract: In order to investigate relationships of adult horses to humans, we developed a simple evaluation test and scores based on observations. The first reactions of 224 adult horses to the presence of an experimenter were observed and scored. All these horses belonged to the same riding school, had the same general housing conditions and were all geldings. The evaluation was based on the horse's posture. Individual differences that could be related to some extent to the breed but also to human factors emerged clearly. French saddlebreds showed more often friendly behaviour than Angloarabs, whereas thoroughbreds were more indifferent. Clear variations occurred between groups of horses that depended on different caretakers. In this school, one caretaker is responsible for the whole daily management of a group of horses and is probably a very important factor in their well-being. The effects of this daily relation to a human seemed to be involved in the reactions to a strange person. Further studies are required to investigate what, in practice, may be determinant.
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Jones, J. E., Antoniadis, E., Shettleworth, S. J., & Kamil, A. C. (2002). A comparative study of geometric rule learning by nutcrackers (Nucifraga columbiana), pigeons (Columba livia), and jackdaws (Corvus monedula). J Comp Psychol, 116(4), 350–356.
Abstract: Three avian species, a seed-caching corvid (Clark's nutcrackers; Nucifraga columbiana), a non-seed-caching corvid (jackdaws; Corvus monedula), and a non-seed-caching columbid (pigeons; Columba livia), were tested for ability to learn to find a goal halfway between 2 landmarks when distance between the landmarks varied during training. All 3 species learned, but jackdaws took much longer than either pigeons or nutcrackers. The nutcrackers searched more accurately than either pigeons or jackdaws. Both nutcrackers and pigeons showed good transfer to novel landmark arrays in which interlandmark distances were novel, but inconclusive results were obtained from jackdaws. Species differences in this spatial task appear quantitative rather than qualitative and are associated with differences in natural history rather than phylogeny.
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Shettleworth, S. J., & Westwood, R. P. (2002). Divided attention, memory, and spatial discrimination in food-storing and nonstoring birds, black-capped chickadees (Poecile atricapilla) and dark-eyed juncos (Junco hyemalis). J Exp Psychol Anim Behav Process, 28(3), 227–241.
Abstract: Food-storing birds, black-capped chickadees (Poecile atricapilla), and nonstoring birds, dark-eyed juncos (Junco hyemalis), matched color or location on a touch screen. Both species showed a divided attention effect for color but not for location (Experiment 1). Chickadees performed better on location than on color with retention intervals up to 40 s, but juncos did not (Experiment 2). Increasing sample-distractor distance improved performance similarly in both species. Multidimensional scaling revealed that both use a Euclidean metric of spatial similarity (Experiment 3). When choosing between the location and color of a remembered item, food storers choose location more than do nonstorers. These results explain this effect by differences in memory for location relative to color, not division of attention or spatial discrimination ability.
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Hampton, R. R., Healy, S. D., Shettleworth, S. J., & Kamil, A. C. (2002). Neuroecologists' are not made of straw. Trends. Cognit. Sci., 6(1), 6–7.
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Saslow, C. A. (2002). Understanding the perceptual world of horses. Appl. Anim. Behav. Sci., 78(2-4), 209–224.
Abstract: 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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Nicol, C. J. (2002). Equine learning: progress and suggestions for future research. Appl. Anim. Behav. Sci., 78(2-4), 193–208.
Abstract: 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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Broom, M. (2002). A unified model of dominance hierarchy formation and maintenance. J. Theor. Biol., 219(1), 63–72.
Abstract: In many different species it is common for animals to spend large portions of their lives in groups. Such groups need to divide available resources amongst the individuals they contain and this is often achieved by means of a dominance hierarchy. Sometimes hierarchies are stable over a long period of time and new individuals slot into pre-determined positions, but there are many situations where this is not so and a hierarchy is formed out of a group of individuals meeting for the first time. There are several different models both of the formation of such dominance hierarchies and of already existing hierarchies. These models often treat the two phases as entirely separate, whereas in reality, if there is a genuine formation phase to the hierarchy, behaviour in this phase will be governed by the rewards available, which in turn depends upon how the hierarchy operates once it has been formed. This paper describes a method of unifying models of these two distinct phases, assuming that the hierarchy formed is stable. In particular a framework is introduced which allows a variety of different models of each of the two parts to be used in conjunction with each other, thus enabling a wide range of situations to be modelled. Some examples are given to show how this works in practice.
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Chase, I. D., Tovey, C., Spangler-Martin, D., & Manfredonia, M. (2002). Individual differences versus social dynamics in the formation of animal dominance hierarchies. Proc. Natl. Acad. Sci. U.S.A., 99(8), 5744–5749.
Abstract: Linear hierarchies, the classical pecking-order structures, are formed readily in both nature and the laboratory in a great range of species including humans. However, the probability of getting linear structures by chance alone is quite low. In this paper we investigate the two hypotheses that are proposed most often to explain linear hierarchies: they are predetermined by differences in the attributes of animals, or they are produced by the dynamics of social interaction, i.e., they are self-organizing. We evaluate these hypotheses using cichlid fish as model animals, and although differences in attributes play a significant part, we find that social interaction is necessary for high proportions of groups with linear hierarchies. Our results suggest that dominance hierarchy formation is a much richer and more complex phenomenon than previously thought, and we explore the implications of these results for evolutionary biology, the social sciences, and the use of animal models in understanding human social organization.
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Broom, M., & Cannings, C. (2002). Modelling Dominance Hierarchy formation as a Multi-player game. J. Theor. Biol., 219(3), 397–413.
Abstract: Animals who live in groups need to divide available resources amongst themselves. This is often achieved by means of a dominance hierarchy, where dominant individuals obtain a larger share of the resources than subordinate individuals. This paper introduces a model of dominance hierarchy formation using a multi-player extension of the classical Hawk-Dove game. Animals play non-independent pairwise games in a Swiss tournament which pairs opponents against those which have performed equally well in the conflict so far, for a fixed number of rounds. Resources are divided according to the number of contests won. The model, and its emergent properties, are discussed in the context of experimental observations.
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