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Ahrendt, L. P., Labouriau, R., Malmkvist, J., Nicol, C. J., & Christensen, J. W. (2015). Development of a standard test to assess negative reinforcement learning in horses. Appl. Anim. Behav. Sci., 169, 38–42.
Abstract: Most horses are trained by negative reinforcement. Currently, however, no standardised test for evaluating horses' negative reinforcement learning ability is available. The aim of this study was to develop an objective test to investigate negative reinforcement learning in horses. Twenty-four Icelandic horses (3 years old) were included in this study. The horses were tested in a pressure-release task on three separate days with 10, 7 and 5 trials on each side, respectively. Each trial consisted of pressure being applied on the hindquarter with an algometer. The force of the pressure was increased until the horse moved laterally away from the point of pressure. There was a significant decrease in required force over trials on the first test day (P<0.001), but not the second and third day. The intercepts on days 2 and 3 differed significantly from day 1 (P<0.001), but not each other. Significantly stronger force was required on the right side compared to the left (P<0.001), but there was no difference between first and second side tested (P=0.56). Individual performance was evaluated by median-force and the change in force over trials on the first test day. These two measures may explain different characteristics of negative reinforcement learning. In conclusion, this study presents a novel, standardised test for evaluating negative reinforcement learning ability in horses.
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Giles, S. L., Nicol, C. J., Harris, P. A., & Rands, S. A. (2015). Dominance rank is associated with body condition in outdoor-living domestic horses (Equus caballus). Applied Animal Behaviour Science, .
Abstract: Abstract The aim of our study was to explore the association between dominance rank and body condition in outdoor group-living domestic horses, Equus caballus. Social interactions were recorded using a video camera during a feeding test, applied to 203 horses in 42 herds. Dominance rank was assigned to 194 individuals. The outcome variable body condition score (BCS) was recorded using a 9-point scale. The variables age and height were recorded and considered as potential confounders or effect modifiers. Results were analysed using multivariable linear and logistic regression techniques, controlling for herd group as a random effect. More dominant (p = 0.001) individuals generally had a higher body condition score (p = 0.001) and this association was entirely independent of age and height. In addition, a greater proportion of dominant individuals fell into the obese category (BCS >= 7/9, p = 0.005). There were more displacement encounters and a greater level of interactivity in herds that had less variation in age and height, lending strength to the hypothesis that phenotypic variation may aid cohesion in group-living species. In addition there was a strong quadratic relationship between age and dominance rank (p < 0.001), where middle-aged individuals were most likely to be dominant. These results are the first to link behavioural predictors to body condition and obesity status in horses and should prompt the future consideration of behavioural and social factors when evaluating clinical disease risk in group-living animals.
Keywords: Equine; fatness; obesity; social behaviour; displacement
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Søndergaard, E., Jensen, M. B., & Nicol, C. J. (2011). Motivation for social contact in horses measured by operant conditioning. Appl. Anim. Behav. Sci., 132(3-4), 131–137.
Abstract: Although horses are social animals they are often housed individually with limited social contact to other horses and this may compromise their welfare. The present study included eight young female horses and investigated the strength of motivation for access to full social contact, head contact and muzzle contact, respectively, to a familiar companion horse. Horses were housed individually next to their companion horse and separations between pens prevented physical contact. During daily test sessions horses were brought to a test area where they could access an arena allowing social contact. Arena access during 3 min was given after completion of a predetermined number of responses on a panel. Fixed ratios (FR) of 8, 16, 24, 32 and 40 responses per arena access were applied in a random order, one per daily test session, within each test week (Monday to Friday), and the number of rewards per daily test session was recorded. All horses could access all three types of social contact in a cross-over design, and an empty arena was used as control. Motivational strength was assessed using elasticity of demand functions, which were estimated based on the number of rewards earned and FR. Elasticities of demand for the three types of social contact were low (-0.20), and not significantly different, although increasing FR still resulted in a decrease in rewards obtained for all three types of social contact (P < 0.001). Across FR-levels horses earned more rewards for social contact than for an empty arena, as shown by much higher intercept values (2.51 vs. 0.99; P < 0.001). However, the elasticity of demand for infrequent access to an empty arena (-0.08) was lower than for social contact (P < 0.01) and not significantly different from zero (P = 0.07). Horses performed more social behaviour the lesser the restriction on social contact (full > head > muzzle). However, the finding that horses showed a similar and high motivation for all three types of social contact suggests that they are valued equally highly in a situation where the alternative is no social contact.
Keywords: Horse; Housing; Social behaviour; Operant conditioning; Motivation
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Hothersall, B., & Nicol, C. (2009). Role of Diet and Feeding in Normal and Stereotypic Behaviors in Horses. Clinical Nutrition, 25(1), 167–181.
Abstract: This article reviews the effects of diet on equine feeding behavior and feeding patterns, before considering the evidence that diet affects reactivity in horses. A growing body of work suggests that fat- and fiber-based diets may result in calmer patterns of behavior, and possible mechanisms that may underpin these effects are discussed. In contrast, there is little evidence that herbal- or tryptophan-containing supplements influence equine behavior in any measurable way. The role of diet in the development of abnormal oral behaviors, particularly the oral stereotypy crib-biting, is also reviewed, and suggestions for future work are presented.
Keywords: Equine behavior; Diet; Crib-biting; Stereotypy; Weaning; Tryptophan; Insulin
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Abeyesinghe, S. M., Nicol, C. J., Hartnell, S. J., & Wathes, C. M. (2005). Can domestic fowl, Gallus gallus domesticus, show self-control? Anim. Behav., 70(1), 1–11.
Abstract: An important aspect of cognition is whether animals live exclusively in the present or can anticipate the future. Defined as self-control, the ability to choose a large, remote reinforcer over a small, proximate reinforcer available at the same frequency has been examined in a number of species, often proving difficult to demonstrate. We investigated self-control for food in domestic fowl using a standard two-key operant task and an equivalent two-choice return maze (TCRM) task. When hens chose between a 2-s delay to a 3-s feed access (impulsive) and a 6-s delay to a 7-s feed access (self-control), they appeared unable to discriminate in the TCRM but were impulsive in the operant task. We explored reasons for not choosing self-control in the operant task, first by examining the relation between feed access time and actual feed intake. A second operant experiment examined whether failure to show self-control could be attributed to an inability to combine the delay and access (quantity) reward information associated with choices to reach overall predictions of value. New hens chose between a 2-s delay to a 3-s feed access (impulsive) and either a 22-s delay to a 22-s feed access (standard self-control) or a 6-s delay to a 22-s feed access (jackpot self-control). While hens were impulsive in the standard condition, they showed significant and pronounced self-control in the jackpot condition, eliminating the possibility of an absolute cognitive constraint. Impulsive behaviour can instead be explained by temporal discounting: perceived depreciation of reward value as a function of the uncertainty associated with delay. Implications for welfare are discussed.
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Nicol, C. J., & Pope, S. J. (1999). The effects of demonstrator social status and prior foraging success on social learning in laying hens. Anim. Behav., 57(1), 163–171.
Abstract: Opportunities for social learning within a group of animals are likely to be influenced by the social dynamics of that group. Some individuals may be more influential demonstrators than others even when there are no differences in their skill level or performance. In this study of domestic hens,Gallus gallus domesticus, differences in demonstrator salience were examined. From 24 separate flocks we selected as demonstrators a dominant cockerel, a dominant hen, a mid-ranking hen or a subordinate hen. Demonstrators were pretrained to perform an operant discrimination task to obtain food. Six observers from each flock individually watched the demonstrator perform the task for four 5-min sessions held on consecutive days. On the fifth day observers were tested individually in the operant chamber. We analysed data from 19 flocks, where there were no quantitative differences in demonstrator performance. Observer hens of relatively high social status performed more correct operant pecks than observer hens of relatively low social status. Demonstrator category also had a significant effect on subsequent observer behaviour. Hens that had observed cockerels performed very few general pecks or operant pecks. Hens that had observed dominant hens performed more operant pecks, but hens that had observed sub-ordinate hens performed more general pecks in the chamber. The results suggested either that there was an interaction between dominance and gender in demonstrator salience or that dominant hens might have been influential because of some factor imperfectly associated with their dominance status. A possible candidate was the foraging ability of the dominant hens. In a second experiment using the same protocol, we manipulated the prior foraging success of dominant hens from four additional flocks but this had no significant effect on their subsequent influence as demonstrators.
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Søndergaard, E., Jensen, M. B., & Nicol, C. J. (). Motivation for social contact in horses measured by operant conditioning. Appl. Anim. Behav. Sci., In Press, Corrected Proof.
Abstract: Although horses are social animals they are often housed individually with limited social contact to other horses and this may compromise their welfare. The present study included eight young female horses and investigated the strength of motivation for access to full social contact, head contact and muzzle contact, respectively, to a familiar companion horse. Horses were housed individually next to their companion horse and separations between pens prevented physical contact. During daily test sessions horses were brought to a test area where they could access an arena allowing social contact. Arena access during 3 min was given after completion of a predetermined number of responses on a panel. Fixed ratios (FR) of 8, 16, 24, 32 and 40 responses per arena access were applied in a random order, one per daily test session, within each test week (Monday to Friday), and the number of rewards per daily test session was recorded. All horses could access all three types of social contact in a cross-over design, and an empty arena was used as control. Motivational strength was assessed using elasticity of demand functions, which were estimated based on the number of rewards earned and FR. Elasticities of demand for the three types of social contact were low (-0.20), and not significantly different, although increasing FR still resulted in a decrease in rewards obtained for all three types of social contact (P < 0.001). Across FR-levels horses earned more rewards for social contact than for an empty arena, as shown by much higher intercept values (2.51 vs. 0.99; P < 0.001). However, the elasticity of demand for infrequent access to an empty arena (-0.08) was lower than for social contact (P < 0.01) and not significantly different from zero (P = 0.07). Horses performed more social behaviour the lesser the restriction on social contact (full > head > muzzle). However, the finding that horses showed a similar and high motivation for all three types of social contact suggests that they are valued equally highly in a situation where the alternative is no social contact.
Keywords: Horse; Housing; Social behaviour; Operant conditioning; Motivation
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Hothersall, B., Harris, P., Sörtoft, L., & Nicol, C. J. (2010). Discrimination between conspecific odour samples in the horse (Equus caballus). Appl. Anim. Behav. Sci., 126(1-2), 37–44.
Abstract: Behavioural observations suggest that smell is important in social discriminations between horses but balanced studies of this capacity are lacking. We used a habituation-discrimination procedure to investigate the ability of horses to distinguish between pairs of odour samples from different individuals. In Study 1, separate tests were conducted for urine, faeces or fleece fabric previously rubbed on the coat (to pick up body odour samples (BOS)) and donor pairs differed in sex, and age. 10 pregnant mares each underwent three tests, one per sample type. A test consisted of three successive 2-min presentations of a sample from Individual A with a simultaneous presentation of a sample from Individual B during the final presentation. Doubly repeated measures ANOVA indicated a main effect of sample type on investigative response (df = 2, f = 7.98, P = 0.004): durations were longer for BOS than for urine or faeces but habituation across trials was most consistent for urine. In the final presentation, mares demonstrated discrimination by investigating the novel urine sample (B) more than the repeated sample (novel: median 8.0 s, IQR = 10; repeated: median 2.5 s, IQR = 6; z = -2.558, P = 0.008). In Study 2, urine samples from castrated male donors were used and neither mares nor their 4-month-old foals discriminated between samples from different individuals in the final presentation. The findings suggest that urine odour may contain some information that horses can use to discriminate between conspecifics. This may be limited to the level of broad categories such as sex or reproductive status; further investigation is needed to reveal what functional information can be transmitted and what compounds are involved.
Keywords: Horse; Olfaction; Social discrimination; Urine; Faeces
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Nicol, C. J. (2006). How animals learn from each other. Appl. Anim. Behav. Sci., 100(1-2), 58–63.
Abstract: This paper explores ways by which animals may learn from one another, using examples drawn mostly from the chicken, an animal for which social learning is likely to be less dangerous than individual learning. In early life, the behaviour of the hen is important in encouraging chicks to peck at edible items. Maternal display not only attracts chicks to profitable food items, but also redirects their attention away from harmful or non-profitable items. Older chicks can enhance their foraging success by observing the behaviour of conspecifics within their own social group. Hens have been trained to perform a novel behaviour (key-pecking for food) by observation of a trained demonstrator bird. Moreover, observers learnt most from watching dominant demonstrators. Thus the ability to learn from others is not `fixed', but depends on the context and the social identity of both the observer and the demonstrator.
Keywords: Social learning; Chickens; Demonstrators; Dominance
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Nicol, C. J. (2002). Equine learning: progress and suggestions for future research. Appl. Anim. Behav. Sci., 78(2-4), 193–208.
Abstract: Horses are well able to form classical and instrumental associations and so the focus of much recent research has been on the stimulus control of instrumental learning. Horses appear to discriminate using spatial cues more easily than other stimulus features, as indicated both by the speed of initial task acquisition and by the extent to which acquired discriminations can be reversed. Phenomena associated with discrimination learning in laboratory animals, including generalisation and peak shift, have been demonstrated in horses. However, the ability of horses to classify stimuli into categories is more controversial. Although there is some evidence that horses may be able to form categories based on similarities in the physical appearance of different stimuli, there is currently no evidence that they are able to develop abstract concepts. Their performance on social learning tasks has also been poor. Few correlations are observed between the learning ability of individual horses on different tasks, suggesting that it may not be possible to classify individual horses as `good' or `poor' learners. Better learning performance by horses that are naturally calm is probably due to reduced interference in the learning process. Correct handling procedures can lower reactivity levels in horses, and may facilitate learning in some circumstances. Future research on equine learning needs to take into account the complex nature of equine social interaction. Studies on the effects of stress on learning, and on social and spatial cognition, are also particularly needed.
Keywords: Learning; Horse; Equine; Discrimination; Training
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