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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.
Keywords: Animals; Auditory Pathways/physiology; Auditory Perception/*physiology; Avoidance Learning/physiology; Brain Mapping; Electroshock; Female; Horses/*physiology; Male; Olivary Nucleus/anatomy & histology/*physiology; Orientation/physiology; Pitch Perception/physiology; Sound Localization/*physiology
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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.
Keywords: Cattle; Avoidance distance; Human-animal relationship (HAR); Dominance
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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.
Keywords: Anxiety; Avoidance; Behavioural test; Emotion; Fear; Flight; Habituation; Horse; Vocalization
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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.
Keywords: Avoidance task; Equus caballus; Fearfulness; Learning; Personality; Temperament
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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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Shier, D. M., & Owings, D. H. (2007). Effects of social learning on predator training and postrelease survival in juvenile black-tailed prairie dogs, Cynomys ludovicianus. Anim. Behav., 73(4), 567–577.
Abstract: We examined how social context and experience affected development of antipredator behaviour and subsequent postrelease survival in the black-tailed prairie dog. Captive-reared juveniles were initially exposed to four stimulus animals: a ferret, a rattlesnake, a hawk and a cottontail control (pretraining tests). Subjects were then trained with or without an adult female demonstrator. Training involved exposure to each stimulus animal two to three times over 5 weeks. After training, each juvenile was retested with each stimulus animal (post-training tests). During pretraining tests, juveniles responded differentially to the stimulus animals. They were least active with the snake, fled the most in tests with the hawk, and were less vigilant with the ferret than with the snake. Following training, juveniles trained with experienced adults were more wary with all three predators than juveniles trained without an experienced adult present. We then compared the antipredator behaviour of captive-reared juveniles trained with experienced adult females with that of wild-reared juveniles of the same age. For all behavioural measures except shelter use, wild-experienced animals differentiated more strongly among predator types than did captive-trained juveniles. One year after reintroduction, survivorship of juveniles trained with experienced adults was higher than that of juveniles trained without experienced adults, but did not differ from that of wild-reared juveniles. These findings provide the first evidence that social transmission of antipredator behaviour during training can enhance long-term survival following release and that as long as a social training regime is used, predator avoidance training can emulate experience acquired in the wild.
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McCall, C. A., Salters, M. A., & Simpson, S. M. (1993). Relationship between number of conditioning trials per training session and avoidance learning in horses. Appl. Anim. Behav. Sci., 36(4), 291–299.
Abstract: Sixteen horses were used to determine if number of trials given per training session (5, 10, 15 or 20) affected learning performance in an avoidance conditioning task. The horse had to move from one side of a test pen to the other during an auditory cue presentation to avoid aversive stimulation. A pen 8 mx3.6 m, divided into two equal sections by a 13-cm diameter plastic pipe lying on the ground, was used as the test pen. Painted plywood panels were fastened to the fence in half the pen to help horses distinguish visually between the two parts. A 10-s auditory cue was used as a signal for horses to move from one side of the test pen to the other. A 20-s intertrial interval was used. Training sessions were conducted every third day. Each trial was recorded as an avoidance (the horse completed the task during auditory cue presentation and avoided aversive stimulus) or an error (the horse received aversive stimulus). After completing ten consecutive avoidances (criterion), the horse was removed from the study. Numbers of training sessions, trials, avoidances and errors until reaching criterion were recorded for each horse. Horses varied greatly within these variables with ranges of 3-18 sessions, 37-121 trials, 20-68 avoidances and 17-53 errors to criterion. No differences were detected (P>0.05) in the number of conditioning trials per training session (treatment) for the mean number of trials, avoidances or errors to criterion. Number of training sessions to criterion differed (P<0.01) among treatments, indicating that an optimum number of learning trials per training session might exist. Mean sessions to criterion for horses receiving 5, 10, 15 and 20 trials per session were 15.1+/-1.3, 5.8+/-1.1, 5.3+/-1.1 and 4.6+/-1.1, respectively. Regression analysis indicated that 16.2 trials per training session would minimize number of sessions to criterion. Although it is widely assumed that learning efficiency in horses is decreased when intense activity is concentrated into a small number of sessions, these results indicate that moderate repetition of training activities is needed for efficient learning.
Keywords: Horse; Learning; Avoidance conditioning
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De Moraes Ferrari, E. A., & Todorov, J. C. (1980). Concurrent avoidance of shocks by pigeons pecking a key. J Exp Anal Behav., 30(3), 329–333.
Abstract: Three pigeons were studied on concurrent, unsignaled, avoidance schedules in a two-key procedure. Shock-shock intervals were two seconds in both schedules. The response-shock interval on one key was always 22 seconds, while the response-shock interval associated with the other key was varied from 7 to 52 seconds in different experimental conditions. Response rates on the key associated with the varied schedule tended to decrease when the response-shock interval length was increased. Responding on the key associated with the constant schedule was not systematically affected.
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Sachs, E. (1967). Dissociation of learning in rats and its similarities to dissociative states in man. Proc Annu Meet Am Psychopathol Assoc, 55, 249–304.
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Mitchell, D., Kirschbaum, E. H., & Perry, R. L. (1975). Effects of neophobia and habituation on the poison-induced avoidance of exteroceptive stimuli in the rat. J Exp Psychol Anim Behav Process, 1(1), 47–55.
Abstract: Two experiments on the role of neophobia in poison-induced aversions to exteroceptive stimuli are reported. In Experiment 1, rats were given either 10 or 25 days of habituation to the test situation prior to conditioning. Those animals with the longer habituation period avoided a complex of novel exteroceptive stimuli while those with the shorter habituation period did not. In Experiment 2 rats initially avoided the more novel of two containers, but gradually came to eat equal amounts from both. A single pairing of toxicosis with consumption from either the novel or the familiar container reinstated the avoidance of the novel container in both cases. The results were discussed in terms of an interaction between habituation and conditioning procedures. It was suggested that previously reported differences between interoceptive and exteroceptive conditioning effects may have been influenced by the differential novelty of the two classes of stimuli in the test situation. It was further suggested that non-contingently poisoned control groups should routinely be included in poison avoidance conditioning studies.
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