Duncan, I. J. H. (1995). D.G.M. Wood-Gush Memorial Lecture: An applied ethologist looks at the question “Why?”. Appl. Anim. Behav. Sci., 44(2-4), 205–217.
Abstract: The question “Why does an animal behave as it does?” can be answered in terms of ontogeny, function, phylogeny and causation. The achievements of applied ethology relative to those four approaches are reviewed, gaps in our knowledge are identified and predictions for fruitful avenues of future research are made. Ontogenic studies have been useful in the past and it is suggested that studies of the effects of early experience on the sexual behaviour of animals used in artificial breeding schemes might pay dividends. It is proposed that functional studies should be approached cautiously. More information is required on the process of domestication in order to increase the chances of success in the trend to farm exotic species. Studies on causation are likely to continue to be the mainstay of applied ethological research. It is suggested that within this category, studies on states of suffering, motivation and cognition are urgently required to answer the most pressing questions on animal welfare.
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Duncan, I. J. H., & Petherick, J. C. (1989). Proceeding (Paper presented at the Winter Meeting of the Society for Veterinary Ethology, London, Great Britain, 30 November 1988)Cognition: The implications for animal welfare. Appl. Anim. Behav. Sci., 24(1), 81–1010.
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Appleby M. (2002). Consciousness, Cognition and Animal Welfare – J.K. Kirkwood, R.C. Hubrecht, S. Wickens, H. O'Leary, S. Oakley (Eds.), Universities Federation for Animal Welfare, 2001, 251 pp., Paperback, Supplement to Volume 10 of Animal Welfare, 15/US$ 30, ISSN 0962-7286. Appl. Anim. Behav. Sci., 77, 239–241.
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Flannery, B. (1997). Relational discrimination learning in horses. Appl. Anim. Behav. Sci., 54(4), 267–280.
Abstract: This series of studies investigated horses' ability to learn the concept of sameness under several different conditions. Before experimentation began, three horses were shaped to touch individually presented stimuli with their muzzles, and then to make two responses to two matching cards from an array of three. A modified version of the identity matching-to-sample (IMTS) procedure was used to present stimuli in a variety of configural arrangements on a barn wall (Experiment 1 and Experiment 2), and on a flat panel mounted to a barn door (Experiment 3). The task in each experiment was to select the two stimulus cards that were the same (either circles or Xs) and to avoid the nonmatching stimulus card (either a star or a square). In Experiment 1, the mean accuracy rate for selecting the matching alternatives was 74%. The horses' accuracy levels reached a mean level of 83% during Experiment 2, in which they received additional trials and an intermittent secondary reinforcement schedule. In Experiment 3, when the stimuli were moved further apart from each other within arrangements and were presented on a novel background, the mean accuracy rate was 73%. These data demonstrate that horses can learn complex discrimination problems involving the concept of sameness, and that they are able to generalize this learning to a novel stimulus presentation situation. These results also suggest that a relational discrimination test may be useful for assessing horses' learning ability and the level of training appropriate for individual horses.
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Murphy, J., Waldmann, T., & Arkins, S. (2004). Sex differences in equine learning skills and visuo-spatial ability. Appl. Anim. Behav. Sci., 87(1-2), 119–130.
Abstract: There is evidence of superior visuo-spatial ability in males compared to females in most species investigated to-date. However, no known studies have addressed this issue in the equine. Equine visuo-spatial ability was investigated using a novel test apparatus with a sample of 62 horses (males=34 and females=28) during a series of six tests, where the horses were required to access a food source. The test apparatus consisted of a series of four adjacent stalls, each of which had a feed bin and a moveable barrier. The test apparatus was designed such that the breastplate barriers controlled and limited access by the horses to feed bins in all but one stall during each test. Male horses performed such that there were significant differences (P<0.05) in the ability of the subjects to complete all six tests in a mean time of 30 s or less per test. There were significant differences in mean completion times for male subjects between test 1 and test 2 (P<0.05), test 1 and test 3 (P<0.001), test 1 and test 4 (P<0.05) and test 1 and test 5 (P<0.05). There were no significant differences in mean completion times between any of the six tests for female subjects. Males had a lower mean total number of errors during all tests. Male horses also successfully completed significantly more tests than females (P<0.05). These results provide the first behavioural demonstration of superior visuo-spatial ability in male horses, similar to that reported in other species.
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McCall, C. A. (1989). The effect of body condition of horses on discrimination learning abilities. Appl. Anim. Behav. Sci., 22(3-4), 327–334.
Abstract: Discriminative learning abilities were studied in 12 mature, malnourished horses. All horses initially received a condition score (CS) between 2 and 4 on a scale of 1 (poor) to 9 (extremely fat). Then horses were assigned to one of 3 treatments based on their eventual, rehabilitated CS during discrimination testing: thin, CS 1-3; moderate, CS 4-6; and fat, CS 7-9. The discrimination learning task was performed for 14 days with a maximum of 20 trials per day. Daily criterion was set at eight consecutively correct trails. Total trials to first criteria and total errors during testing were recorded. Analysis of variance showed that treatments did not differ (P>0.05) in total trials to first criterion, however horses on the fat treatment did have higher total error scores (P<0.05) than horses on the thin or moderate treatments. This difference was probably owing to lack of motivation in the fat treatment horses, rather than to true learning ability differences. The sex of the horse did not significantly affect either learning score.
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Marinier, S. L., & Alexander, A. J. (1994). The use of a maze in testing learning and memory in horses. Appl. Anim. Behav. Sci., 39(2), 177–182.
Abstract: Two mazes were used to test the learning ability and memory of horses, and changes in these abilities. Testing was done on four occasions. On Occasion 1, the horses were run through Maze A until they had reached the criterion of three consecutive correct runs. A week later (Occasion 2), they were retested in Maze A to the same criterion as a measure of memory. On Occasion 3,2 months later, the horses were run through Mazes A and B until they reached the criterion. Occasion 4 took place 1 week later when they were run through Mazes A and B. An estimation of changes in ability to learn came from a comparison of results from Occasions 1 and 3. Similarly, changes in ability to remember came from a comparison of results from Occasions 2 and 4. Nine horses with a variable amount of riding training were the subjects. All horses were able to learn the maze, but the ability varied among horses. There was no obvious correlation between quality of handling of the horses and learning ability. Once the horses had learned the maze, they remembered it perfectly on subsequent occasions. There were changes in the memory and learning ability of the horses, but no clear explanation for this could be found.
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McCall, C. A., Hall, S., McElhenney, W. H., & Cummins, K. A. (2006). Evaluation and comparison of four methods of ranking horses based on reactivity. Appl. Anim. Behav. Sci., 96(1-2), 115–127.
Abstract: Four methods of ranking horses on reactivity were evaluated and compared: isolation from conspecifics, presentation of a static novel stimulus, traversing a novel stimulus in a runway (isolation, novel stimulus and runway tests, respectively) and assigning subjective emotionality scores. In all tests, horses' heart rates were recorded and behaviour was videotaped. To be considered a valid test of reactivity, at least one heart rate and one behavioural measurement in the test had to change significantly between treatments (tranquilizer administation versus sham tranquilizer administration), and behavioural measures had to be displayed in at least 75% of the trials. Forty horses performed each of the three tests daily on three different days in a switchback design. Horses were assigned randomly to a daily test sequence, which was maintained throughout the study. In the runway test, no significant difference in heart rate values in tranquilized and non-tranquilized horses was found, and no behavioural attribute was displayed in more than 52% of the trials; therefore it was rejected as a valid test of reactivity. Both isolation and novel stimulus tests produced valid measurements. Mean heart rate was the most precise physiological measure for these tests, and walking and defecation frequency were the most precise behavioural measures for novel stimulus and isolation tests, respectively. Mean heart rates on the novel stimulus and isolation tests were correlated (rs = 0.79, P < 0.01) indicating that these tests produced similar rankings based on physiological responses. However, behavioural measures ranked horses differently (rs = 0.27, P < 0.10) on the tests. Rank correlations between mean heart rates and behavioural measures were higher in the novel stimulus (rs = 0.66, P < 0.01) than the isolation test (rs = 0.55, P < 0.01), indicating that the novel stimulus test ranked horses based on either physiological or behavioural responses more similarly than did the isolation test. Therefore, the novel stimulus test was considered the more accurate evaluation of reactivity. Subjective emotionality scores were correlated moderately with mean heart rates (rs > 0.33, P < 0.01) from the novel stimulus and isolation tests and with walking scores (rs = 0.47, P < 0.01) from the novel stimulus test. Assignment of subjective emotionality scores was not as accurate as the novel stimulus or isolation tests in ranking horses for reactivity. Using physiological data alone, combining physiological and behavioural measurements or using more than one behavioural measurement in reactivity tests may reflect the reactivity of the horse better than a single behavioural measurement.
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Williams, J. L., Friend, T. H., Nevill, C. H., & Archer, G. (2004). The efficacy of a secondary reinforcer (clicker) during acquisition and extinction of an operant task in horses. Appl. Anim. Behav. Sci., 88(3-4), 331–341.
Abstract: “Clicker training” is a popularly promoted training method based on operant conditioning with the use of a secondary reinforcer (the clicker). While this method draws from theories of learning and is used widely, there has been little scientific investigation of its efficacy. We used 60 horses, Equus callabus, and assigned each horse to one of six reinforcement protocols. The reinforcement protocols involved combinations of reinforcers administered (primary versus secondary plus primary), schedule of reinforcement (continuous versus variable ratio), and reinforcers applied during extinction (none or secondary). There were no differences (P>=0.11) between horses which received a secondary reinforcer (click) followed by the primary reinforcer (food) and those which received only the primary reinforcer (food) in the number of trials required to train the horses to touch their noses to a plastic cone (operant response). There also were no differences (P>=0.12) between horses which received the secondary reinforcer plus primary reinforcer and those which received only the primary reinforcer in regards to the number of trials to extinction. We conclude that there is no difference in the amount of training required to learn the operant task or in the task's resistance to extinction between horses that received a secondary reinforcer followed by a primary reinforcer versus horses which received only a primary reinforcer.
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Langbein, J., Siebert, K., Nuernberg, G., & Manteuffel, G. (2007). The impact of acoustical secondary reinforcement during shape discrimination learning of dwarf goats (Capra hircus). Appl. Anim. Behav. Sci., 103(1-2), 35–44.
Abstract: The use of secondary reinforcement is widely accepted to support operant learning in animals. In farm animals, however, the efficacy of secondary reinforcement has up to now been studied systematically only in horses (“clicker training”), and the results are controversial. We investigated the impact of acoustical secondary reinforcement on voluntary, self-controlled visual discrimination learning of two-dimensional shapes in group-housed dwarf goats (Capra hircus). Learning tests were conducted applying a computer-controlled learning device that was integrated in the animals' home pen. Shapes were presented on a TFT-screen using a four-choice design. Drinking water was used as primary reinforcement. In the control group (Gcontrol, n = 5) animals received only primary reinforcement, whereas in the sound group (Gsound, n = 6) animals got additional acoustical secondary reinforcement. Testing recall of shapes which had been successfully learned by the goats 6 weeks earlier (T1), we found a weak impact of secondary reinforcement on daily learning success (P = 0.07), but not on the number of trials the animals needed to reach the learning criterion (trials to criterion, n.s.). Results in T1 indicated that dwarf goats did not instantly recall previously learned shapes, but, re-learned within 250-450 trials. When learning a set of new shapes (T2), there was a strong influence of secondary reinforcement on daily learning success and on trials to criterion. Animals in Gsound reached the learning criterion earlier (P < 0.05) and needed fewer trials (1320 versus 3700; P < 0.01), compared to animals in Gcontrol. Results suggest that acoustical secondary reinforcement supports visual discrimination learning of dwarf goats, especially when the task is new and the salience of S+ is low.
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