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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Mazurek, M., McGee, M., Minchin, W., Crowe, M. A., & Earley, B. (2011). Is the avoidance distance test for the assessment of animals' responsiveness to humans influenced by either the dominant or flightiest animal in the group? Appl. Anim. Behav. Sci., 132(3-4), 107–113.
Abstract: A previously described (Windschnurer et al., 2009) avoidance distance test was used to assess animals’ fear of humans in order to quantify the human–animal relationship (HAR). This study investigated the influence of the dominant and flightiest animals within a group on the responsiveness of animals during the avoidance distance test. Eighty-eight pregnant heifers comprised of four different genotypes were used (22 animals per genotype): Limousin × Holstein-Friesian, Limousin × Simmental, Charolais × Limousin, and Charolais × Simmental. Sixty of the 88 heifers were group housed (n = 5) into 12 pens with 3 pens per breed, while 28 heifers were singly housed (seven heifers per breed). A reactivity test was performed on days 10, 18, 25 and 30 post-housing on the singly housed heifers, and then on the group housed heifers, on the same days, to calculate a reactivity score. On days 33 and 37 flight and dominance tests, respectively, were performed to identify the flightiest and the dominant animal within each group. On day 41, an avoidance test, measuring both the avoidance distance towards a familiar and an unfamiliar human, was performed on all heifers. No difference (P > 0.05) in reactivity scores was found between the genotypes, between pens for the group housed heifers or between singly housed and group housed heifers (P = 0.28). The avoidance distance (AD) of singly (S) housed heifers towards a familiar (F) (ADSF) human was shorter (P < 0.001) than the avoidance distance of group (G) housed heifers towards an unfamiliar human (ADSU). The ADSF and ADGF were correlated with the ADSU and ADGU (R = 0.87 for singly housed heifers; R = 0.61 for group housed heifers, P < 0.001). For the singly housed heifers, no correlation was observed between reactivity score and ADSF (R = 0.36, P = 0.18), whereas the reactivity score and ADSU were correlated (R = 0.68, P = 0.004). For the group housed heifers no significant correlation was detected between the reactivity score and ADGF (R = 0.18, P = 0.22) or ADGU (R = −0.11, P = 0.39). No influence of the most dominant animal and the flightiest animals was found on the behaviour of the group in term of avoidance distance and reactivity (P > 0.05). It is concluded that the assessment of the fear of the animals towards humans using the avoidance test at the feed bunk may be useful for singly and group housed heifers and that the leaders of a group such as the flightiest animal or the dominant animal did not influence the avoidance distance test.
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Leiner, L., & Fendt, M. (2011). Behavioural fear and heart rate responses of horses after exposure to novel objects: Effects of habituation. Appl. Anim. Behav. Sci., 131(3-4), 104–109.
Abstract: The emotion fear promotes the fitness of wild animals. In a farm environment, exaggerated fear, e.g., in horses, can cause several problems. Therefore, knowledge about fear in horses helps to prevent or to handle potential fear-inducing situations. The present study investigated which behavioural fear responses can be observed during exposure of horses to a novel stimulus, whether these behavioural responses are correlated with physiological changes, and whether and how specifically these changes are reduced after habituation training to one of the novel objects. Our data shows that behavioural and physiological fear responses in horses are correlated, are reliable to observe and to measure, and appear in a typical chronological order. Furthermore, after habituation-training to an object, the fear response to this object is specifically attenuated whereas the fear response to another object remains.
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Lansade, L., & Simon, F. (2010). Horses' learning performances are under the influence of several temperamental dimensions. Appl. Anim. Behav. Sci., 125(1-2), 30–37.
Abstract: Learning performances are influenced by many factors, not only breed, age and sex, but also temperament. The purpose of this study was to understand how different temperamental dimensions affect the learning performance of horses, Equus caballus. First, we carried out a series of behavioural tests on 36 Welsh ponies aged 5-7 years to measure five temperamental dimensions: fearfulness (novel area test and surprise test), gregariousness (social isolation test), reactivity to humans (passive human test), tactile sensitivity (von Frey filament test) and activity level (evaluation of locomotor activity during all the tests). We then presented them with two learning tasks (avoidance and backwards-forwards tasks). In the avoidance task they had to learn to jump over a fence when they heard a sound associated with an aversive stimulus (puff of air). In the backwards-forwards task they had to walk forwards or move backwards in response to a tactile or vocal command to obtain a food reward. There was no correlation between performances on the two learning tasks, indicating that learning ability is task-dependent. However, correlations were found between temperamental data and learning performance (Spearman correlations). The ponies that performed the avoidance task best were the most fearful and the most active ones. For instance, the number of trials required to perform 5 consecutive correct responses (learning criterion) was correlated with the variables aimed at measuring fearfulness (way of crossing a novel area: rs = -0.41, P = 0.01 and time to start eating again after a surprise effect: rs = -0.33, P = 0.05) and activity level (frequency of trotting during all the tests: rs = -0.40, P = 0.02). The animals that performed the backwards-forwards task best were the ones that were the least fearful and the most sensitive. For instance, the learning criterion (corresponding to the number of trials taken to achieve five consecutive correct responses) was correlated with the variables aimed at measuring fearfulness (latency to put one foot on the area: rs = 0.43, P = 0.01; way of crossing a novel area: rs = 0.31, P = 0.06; and time to start eating again after a surprise effect: rs = 0.43, P = 0.009) and tactile sensitivity (response to von Frey filaments: rs = -0.44, P = 0.008). This study revealed significant links between temperament and learning abilities that are highly task-dependent.
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Griffin, A. S. (2008). Social learning in Indian mynahs, Acridotheres tristis: the role of distress calls. Anim. Behav., 75(1), 79–89.
Abstract: Socially acquired predator avoidance is a phenomenon in which individuals acquire an avoidance response towards an initially neutral stimulus after they have experienced it together with the antipredator signals of social companions. Earlier research has established that alarm calls used for intraspecific communication are effective stimuli for triggering acquisition. However, animals produce a large range of other antipredator responses that might engage antipredator learning. Here, I examine the effects of conspecific distress calls, a signal that is produced by birds when restrained by a predator, and that appears to be directed towards predators, rather than conspecifics, on predator avoidance learning in Indian mynahs, Acridotheres tristis. Distress calls reflect high levels of alarm in the caller and should, therefore, mediate robust learning. Experiment 1 revealed that subjects performed higher rates of head movements in response to a previously unfamiliar avian mount after it had been presented simultaneously with playbacks of conspecific distress vocalizations. Experiment 2 revealed that increased rates of head saccades resembled the spontaneous response evoked by a novel stimulus more closely than it resembled the response evoked by a perched raptor, suggesting that distress calls inculcated a visual exploratory response, rather than an antipredator response. While it is usually thought that the level of acquisition in learners follows a simple relationship with the level of alarm shown by demonstrators, the present results suggest that this relationship may be more complex. Antipredator signals with different functions may have differential effects on learners.
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