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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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Christensen, J. W., Ahrendt, L. P., Lintrup, R., Gaillard, C., Palme, R., & Malmkvist, J. (2012). Does learning performance in horses relate to fearfulness, baseline stress hormone, and social rank? In Applied Animal Behaviour Science (Vol. 140, pp. 44–52).
Abstract: The ability of horses to learn and remember new tasks is fundamentally important for their use by humans. Fearfulness may, however, interfere with learning, because stimuli in the environment can overshadow signals from the rider or handler. In addition, prolonged high levels of stress hormones can affect neurons within the hippocampus; a brain region central to learning and memory. In a series of experiments, we aimed to investigate the link between performance in two learning tests, the baseline level of stress hormones, measured as faecal cortisol metabolites (FCM), fearfulness, and social rank. Twenty-five geldings (2 or 3 years old) pastured in one group were included in the study. The learning tests were performed by professional trainers and included a number of predefined stages during which the horses were gradually trained to perform exercises, using either negative (NR) or positive reinforcement (PR). Each of the learning tests lasted 3 days; 7min/horse/day. The NR test was repeated in a novel environment. Performance, measured as final stage in the training programme, and heart rate (HR) were recorded. Faeces were collected on four separate days where the horses had been undisturbed at pasture for 48h. Social rank was determined through observations of social interactions during feeding. The fear test was a novel object test during which behaviour and HR were recorded. Performance in the NR and PR learning tests did not correlate. In the NR test, there was a significant, negative correlation between performance and HR in the novel environment (rS=-0.66, P<0.001, i.e. nervous horses had reduced performance), whereas there was no such correlation in the home environment (both NR and PR). Behavioural reactions in the fear test correlated significantly with performance in the NR test in the novel environment (e.g. object alertness and final stage: rS=-0.43, P=0.04), suggesting that performance under unfamiliar, stressful conditions may be predicted by behavioural responses in a fear test. There was a negative correlation between social rank and baseline stress hormones (rS=-0.43, P=0.04), i.e. high rank corresponded to low FCM concentrations, whereas neither rank nor FCM correlated with fearfulness or learning performance. We conclude that performance under stressful conditions is affected by activation of the sympathetic nervous system during training and related to behavioural responses in a standardised fear test. Learning performance in the home environment, however, appears unrelated to fearfulness, social rank and baseline FCM levels.
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Mormède, P., Andanson, S., Aupérin, B., Beerda, B., Guémené, D., Malmkvist, J., et al. (2007). Exploration of the hypothalamic-pituitary-adrenal function as a tool to evaluate animal welfare. Physiol. Behav., 92(3), 317–339.
Abstract: Measuring HPA axis activity is the standard approach to the study of stress and welfare in farm animals. Although the reference technique is the use of blood plasma to measure glucocorticoid hormones (cortisol or corticosterone), several alternative methods such as the measurement of corticosteroids in saliva, urine or faeces have been developed to overcome the stress induced by blood sampling itself. In chronic stress situations, as is frequently the case in studies about farm animal welfare, hormonal secretions are usually unchanged but dynamic testing allows the demonstration of functional changes at several levels of the system, including the sensitization of the adrenal cortex to ACTH and the resistance of the axis to feedback inhibition by corticosteroids (dexamethasone suppression test). Beyond these procedural aspects, the main pitfall in the use of HPA axis activity is in the interpretation of experimental data. The large variability of the system has to be taken into consideration, since corticosteroid hormone secretion is usually pulsatile, follows diurnal and seasonal rhythms, is influenced by feed intake and environmental factors such as temperature and humidity, age and physiological state, just to cite the main sources of variation. The corresponding changes reflect the important role of glucocorticoid hormones in a number of basic physiological processes such as energy metabolism and central nervous system functioning. Furthermore, large differences have been found across species, breeds and individuals, which reflect the contribution of genetic factors and environmental influences, especially during development, in HPA axis functioning. Usually, these results will be integrated with data from behavioral observation, production and pathology records in a comprehensive approach of farm animal welfare.
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Christensen, J. W., Malmkvist, J., Nielsen, B. L., & Keeling, L. J. (2008). Effects of a calm companion on fear reactions in naive test horses. Equine Vet J, 40(1), 46–50.
Abstract: REASON FOR PERFORMING STUDY: In fear-eliciting situations, horses tend to show flight reactions that can be dangerous for both horse and man. Finding appropriate methods for reducing fearfulness in horses has important practical implications. OBJECTIVES: To investigate whether the presence of a calm companion horse influences fear reactions in naive subject horses. HYPOTHESES: The presence of a habituated (calm) companion horse in a fear-eliciting situation can reduce fear reactions in naive subject horses, compared to subject horses with a nonhabituated companion (control). METHODS: Minimally handled (n = 36), 2-year-old stallions were used, 18 as subjects and 18 as companions. Companion horses (n = 9) were habituated to an otherwise frightening, standardised test stimulus (calm companions), whereas the rest (n = 9) of the companion horses remained nonhabituated (control companions). During the test, unique pairs of companion and subject horses were exposed to the test stimulus while heart rate and behavioural responses were registered. Subsequently, subject horses were exposed to the stimulus on their own (post test). RESULTS: Subject horses, paired with a calm companion horse, showed less fear-related behaviour and lower heart rate responses compared to subject horses with control companions. Results from the post test suggest that the difference between treatment groups remained in the subsequent absence of companion horses. CONCLUSIONS AND POTENTIAL RELEVANCE: It appears possible to reduce fear reactions in young, naive horses by allowing them to interact with a calm companion horse in fear-eliciting situations.
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Christensen, J. W., Ladewig, J., Sondergaard, E., & Malmkvist, J. (2002). Effects of individual versus group stabling on social behaviour in domestic stallions. Appl. Anim. Behav. Sci., 75(3), 233–248.
Abstract: Domestic horses (Equus caballus) are typically kept in individual housing systems, in which they are deprived of physical contact. In order to study the effects of social restrictions on behaviour in young horses, nineteen 2-year-old stallions were housed either singly (n=7), or in groups of three (n=12) for 9 months. Subsequently, the stallions were released into two separate 2 ha enclosures according to treatment, and recordings were made on social interactions and nearest neighbours during a 6-week-period, 28 h per week. Previously group stabled stallions frequently had a former group mate as their nearest neighbour (P=0.001), whereas previously singly stabled stallions did not associate more with their former box neighbours, to whom physical contact was limited by bars during the previous treatment. The nearest neighbour was more frequently recorded to be within one horselength of singly stabled than of group stabled stallions (P=0.005). More aggressive behaviour was recorded in the group of previously singly stabled stallions, i.e. bite threats (P=0.032), whereas group stabled stallions tended to make more use of subtle agonistic interactions (displacements, submissive behaviour). Singly stabled stallions also responded to the 9 months of social deprivation by significantly increasing the level of social grooming (P<0.001) and play behaviour (P<0.001), when subsequently interacting freely with other horses. The increased occurrence may relate to a build-up of motivation (a rebound effect), as well as to external factors, such as playful pasture companions and the increased space allowance of the pasture. It is concluded that 2-year-old domestic stallions are sensitive to social deprivation and that stabling has long-term effects, lasting 6 weeks at least, on the social behaviour in stallions.
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