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Jankunis, E. S., & Whishaw, I. Q. (2013). Sucrose Bobs and Quinine Gapes: Horse (Equus caballus) responses to taste support phylogenetic similarity in taste reactivity. Behavioural Brain Research, 256, 284–290.
Abstract: Evidence suggests that behavioural affective reactions to sweet and bitter substances are homologous in humans, nonhuman primates, and rodents. The sweet taste of sucrose elicits facial responses that include rhythmic tongue protrusions whereas the bitter taste of quinine elicits facial responses that include gapes, featuring an opening of the mouth and protrusion of the tongue. The present study using the horse (Equus caballus) was undertaken for three reasons: (1) there is debate about the presence of a sweet receptor gene in the horse, (2) there is a need to expand the examination of facial reactions to taste in lineages other than the closely related lineages of rodents and primates, and (3) the horse provides an opportunity to test the hypothesis that some social signals derive from movements related to taste reaction. The horses were given oral infusions of either sucrose or quinine and their behaviour was examined using frame-by-frame video analysis. Control groups were exposed received water or syringe insertion only. Amongst the many responses made to the infusions, the distinctive response to sucrose was a bob coupled with a slight tongue protrusion and forward movement of the ears; the distinctive response to quinine was a head extension and mouth gape accompanied by a large tongue protrusion and backward movement of the ears. Sucrose Bobs and Quinine Gapes are discussed with respect to: (1) the relevance of facial reactions to both sucrose and quinine to taste receptors in horses, (2) the similarity of features of taste expression in horses to those documented in rodents and primates, and (3) the dissimilarity between facial reactions to taste and other social signals displayed by horses.
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Stahlbaum, C. C., & Houpt, K. A. (1989). The role of the Flehmen response in the behavioral repertoire of the stallion. Physiol. Behav., 45(6), 1207–1214.
Abstract: The role of the Flehmen response in equine behavior was investigated under field and laboratory conditions. In Experiment 1, a field study made of five stallions on pasture with between three and eighteen mares each during the season indicated the following: 1) The Flehmen response was most frequently preceded by nasal, rather than oral, investigation of substances; 2) The stallions' rate of Flehmen varied with the estrous cycles of the mares; 3) The rate of Flehmen response did not show a variation with time of day; and 4) The Flehmen response was most frequently followed by marking behaviors rather than courtship behaviors. The results suggest that the Flehmen response is not an immediate component of sexual behavior, e.g., courtship of the stallion but may be involved in the overall monitoring of the mare's estrous cycle. Therefore the Flehmen response may contribute to the chemosensory priming of the stallion for reproduction. In Experiment 2 stallions were presented with urine or feces of mares in various stages of the reproductive cycle as well as with their own or other males' urine or feces. The occurrence of sniffing and Flehmen was used to determine the discriminatory ability of the stallions. Stallions can differentiate the sex of a horse on the basis of its feces alone, but cannot differentiate on the basis of urine. This ability may explain the function of fecal marking behavior of stallions.
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Ralston, S. L. (1984). Controls of feeding in horses. J. Anim Sci., 59(5), 1354–1361.
Abstract: Members of the genus Equus are large, nonruminant herbivores. These animals utilize the products of both enzymatic digestion in the small intestine and bacterial fermentation (volatile fatty acids) in the cecum and large colon as sources of metabolizable energy. Equine animals rely primarily upon oropharyngeal and external stimuli to control the size and duration of an isolated meal. Meal frequency, however, is regulated by stimuli generated by the presence and (or) absorption of nutrients (sugars, fatty acids, protein) in both the large and small intestine plus metabolic cues reflecting body energy stores. The control of feeding in this species reflects its evolutionary development in an environment which selected for consumption of small, frequent meals of a variety of forages.
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Houpt, K. A., Zahorik, D. M., & Swartzman-Andert, J. A. (1990). Taste aversion learning in horses. J. Anim Sci., 68(8), 2340–2344.
Abstract: The ability of ponies to learn to avoid a relatively novel food associated with illness was tested in three situations: when illness occurred immediately after consuming a feed; when illness occurred 30 min after consuming a feed; and when illness was contingent upon eating one of three feeds offered simultaneously. Apomorphine was used to produce illness. The feeds associated with illness were corn, alfalfa pellets, sweet feed and a complete pelleted feed. The ponies learned to avoid all the fees except the complete feed when apomorphine injection immediately followed consumption of the feed. However, the ponies did not learn to avoid a feed if apomorphine was delayed 30 min after feed consumption. They could learn to avoid alfalfa pellets, but not corn, when these feeds were presented with the familiar “safe foods,” oats and soybean meal. Ponies apparently are able to learn a taste aversion, but there were constraints on this learning ability. Under the conditions of this study, they did not learn to avoid a food that made them sick long after consumption of the food, and they had more difficulty learning to avoid highly palatable feeds.
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