Heffner, R. S., & Heffner, H. E. (1986). Localization of tones by horses: use of binaural cues and the role of the superior olivary complex. Behav Neurosci, 100(1), 93–103.
Abstract: The ability of horses to use binaural time and intensity difference cues to localize sound was assessed in free-field localization tests by using pure tones. The animals were required to discriminate the locus of a single tone pip ranging in frequency from 250 Hz to 25 kHz emitted by loudspeakers located 30 degrees to the left and right of the animals' midline (60 degrees total separation). Three animals were tested with a two-choice procedure; 2 additional animals were tested with a conditioned avoidance procedure. All 5 animals were able to localize 250 Hz, 500 Hz, and 1 kHz but were completely unable to localize 2 kHz and above. Because the frequency of ambiguity for the binaural phase cue delta phi for horses in this test was calculated to be 1.5 kHz, these results indicate that horses can use binaural time differences in the form of delta phi but are unable to use binaural intensity differences. This finding was supported by an unconditioned orientation test involving 4 additional horses, which showed that horses correctly orient to a 500-Hz tone pip but not to an 8-kHz tone pip. Analysis of the superior olivary complex, the brain stem nucleus at which binaural interactions first take place, reveals that the lateral superior olive (LSO) is relatively small in the horse and lacks the laminar arrangement of bipolar cells characteristic of the LSO of most mammals that can use binaural delta I.
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Hedberg, Y., Dalin, A. - M., Ohagen, P., Holm, K. R., & Kindahl, H. (2005). Effect of oestrous-cycle stage on the response of mares in a novel object test and isolation test. Reprod Domest Anim, 40(5), 480–488.
Abstract: In various species, sex, hormonal treatments and oestrous-cycle stage have been shown to affect the animal's response in behavioural tests. Few such studies have been performed in the horse. The main aim of the present study was to investigate whether oestrous-cycle stage affects mares' response to a novel object test and isolation test and, in part, to study whether mares, assumed to suffer from oestrous-related behavioural problems, respond differently in these tests when compared with controls. Twelve mares were tested twice, in oestrus and dioestrus, in a crossover design. Seven behavioural and two heart rate variables were measured for the novel object test and two heart rate variables for the isolation test. Oestrous-cycle stage and whether a mare was classified as a 'problem' mare did not affect the mare's response. However, test order, i.e. the cycle stage a mare was tested in first, affected its reaction. This effect could partly be explained by significant differences between test occasions 1 and 2 in three behavioural variables and one heart rate variable (p < 0.05) in the novel object test. The mares explored the novel object more and had a higher mean heart rate in the first test. Exploring the novel object more could largely be attributed to those mares tested in dioestrus first, perhaps indicating that the mares in oestrus were less receptive to the novel object. The reason for the differences between test occasions could be an effect of learning or habituation.
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De Boyer Des Roches, A., Richard-Yris, M. - A., Henry, S., Ezzaouia, M., & Hausberger, M. (2008). Laterality and emotions: visual laterality in the domestic horse (Equus caballus) differs with objects' emotional value. Physiol. Behav., 94(3), 487–490.
Abstract: Lateralization of emotions has received great attention in the last decades, both in humans and animals, but little interest has been given to side bias in perceptual processing. Here, we investigated the influence of the emotional valence of stimuli on visual and olfactory explorations by horses, a large mammalian species with two large monocular visual fields and almost complete decussation of optic fibres. We confronted 38 Arab mares to three objects with either a positive, negative or neutral emotional valence (novel object). The results revealed a gradient of exploration of the 3 objects according to their emotional value and a clear asymmetry in visual exploration. When exploring the novel object, mares used preferentially their right eyes, while they showed a slight tendency to use their left eyes for the negative object. No asymmetry was evidenced for the object with the positive valence. A trend for an asymmetry in olfactory investigation was also observed. Our data confirm the role of the left hemisphere in assessing novelty in horses like in many vertebrate species and the possible role of the right hemisphere in processing negative emotional responses. Our findings also suggest the importance of both hemispheres in the processing positive emotions. This study is, to our knowledge, the first to demonstrate clearly that the emotional valence of a stimulus induces a specific visual lateralization pattern.
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Kaseda, Y., Ogawa, H., & Khalil, A. M. (1997). Causes of natal dispersal and emigration and their effects on harem formation in Misaki feral horses. Equine Vet J, 29(4), 262–266.
Abstract: Misaki feral horses were separated into 2 herds and the difference between dispersal from natal group (natal dispersal) and dispersal from natal area (natal emigration) was studied. The causes of dispersal and emigration and their effects on harem formation were studied 1979-1994. The number of horses ranged from 73 (mature males: 8, mature females: 26, young males: 8, young females: 3, colt foals: 6, filly foals: 10 and geldings: 12) in 1979 and 86 (mature males: 14, mature females: 37, young males: 12, young females: 7, colt foals: 5, filly foals: 7 and geldings: 4) in 1994 when the present study ended. All 29 males which survived to age 4 years and 58 females which survived to age 3 years left their natal or mother groups at age one to 3. Seventeen of 22 dispersing males and 29 of 39 dispersing females left their natal groups around the birth of their siblings and significant correlations were found between natal dispersal and birth of a sibling. The number of emigrating young males correlated negatively and significantly with the total number of young males in another herd and the number of emigrating young females correlated positively and significantly with the total number of young females in the natal herd. All 13 emigrating stallions which survived to age 5 years formed stable harem groups and a significant correlation was found between natal emigration and harem formation. Twenty-three of 35 resident mares formed stable consort relations with harem stallions and a significant correlation was found between residence and formation of stable consort relations.
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Berger, J. (1983). Induced abortion and social factors in wild horses. Nature, 303(5912), 59–61.
Abstract: Much evidence now suggests that the postnatal killing of young in primates and carnivores, and induced abortions in some rodents, are evolved traits exerting strong selective pressures on adult male and female behaviour. Among ungulates it is perplexing that either no species have developed convergent tactics or that these behaviours are not reported, especially as ungulates have social systems similar to those of members of the above groups. Only in captive horses (Equus caballus) has infant killing been reported. It has been estimated that 40,000 wild horses live in remote areas of the Great Basin Desert of North America (US Department of Interior (Bureau of Land Management), unpublished report), where they occur in harems (females and young) defended by males. Here I present evidence that, rather than killing infants directly, invading males induce abortions in females unprotected by their resident stallions and these females are then inseminated by the new males.
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