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Clarke, J. V., Nicol, C. J., Jones, R., & McGreevy, P. D. (1996). Effects of observational learning on food selection in horses. Appl. Anim. Behav. Sci., 50(2), 177–184.
Abstract: Fourteen riding horses of mixed age and breed were randomly allocated to observer and control treatments. An additional horse was pre-trained as a demonstrator to walk the 13.8 m length of the test arena and select one of two food buckets using colour and pattern cues. Observer horses were exposed to correct performances of the task by the trained demonstrator, for 20 trials held over 2 days. Control horses were subjected to the same handling and placement procedures as the observer horses but without exposure to the behaviour of the demonstrator. The third day for all subjects was designated as a test day. Each subject was released individually in a predetermined place in the arena, and the latency to walk the length of the test arena to the food buckets, the latency to feed, the identity of the bucket approached and the identity of the bucket selected were recorded on ten consecutive trials. During tests both food buckets contained food to minimize the possibility of individual trial and error learning. On the first trial the mean latency to approach the goal area was 18 s for observer horses, compared with 119 s for control horses (t = 2.8, d.f. = 12, P < 0.01) and the mean latency to eat was 35 s for observer horses, compared with 181 s for control horses (t = 4.86, d.f. = 11, P < 0.001). However, observer horses were no more likely to choose the demonstrated bucket than control horses on the first trial. Twelve of the 14 horses decreased their latency to approach the goal area during the series of ten trials, but there were no significant changes in the buckets selected.
Keywords: Horse; Observational learning; Food discrimination
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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. (1995). The social transmission of information and behaviour. Appl. Anim. Behav. Sci., 44(2-4), 79–98.
Abstract: Social influences on established behaviour and on the acquisition of new information and behaviour are reviewed. Distinctions between social facilitation and contagious behaviour are drawn and suggestions for further research on contagious behaviour are made. Socially derived visual, olfactory and auditory cues are considered as important influences on behaviour and subsequent learning. The evidence supporting two potential mechanisms of social learning, i.e. stimulus enhancement followed by individual learning, and imitation, is reviewed in detail. It is argued that the functions of social learning are similarly heterogeneous and include motor skill acquisition, gathering of environmental information, and social conformity. Factors affecting the spread of socially acquired skills, including the social relationship between demonstrator and observer, are highlighted. Lastly, the few studies of social learning that have been conducted with domestic species are reviewed and potential applied goals that could stimulate further research in this area are suggested.
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Nicol, C. J., Badnell-Waters, A. J., Bice, R., Kelland, A., Wilson, A. D., & Harris, P. A. (2005). The effects of diet and weaning method on the behaviour of young horses. Appl. Anim. Behav. Sci., 95(3-4), 205–221.
Abstract: The effects of diet on horse behaviour have not previously been quantified in detail. In this study, we examined the behaviour of 17 foals from the age of 2 to 40 weeks. Each foal received either a starch and sugar (SS) diet or a fat and fibre (FF) diet. The two diets contained similar digestible energy, crude protein and micronutrients, but differed in the fat and non-structural carbohydrate balance. The baseline behaviour of the foals was observed every 2 weeks by focal animal sampling. Additional behavioural observations were conducted when the foals were weaned by one of two methods. Approximately 2 months after weaning, the temperament and tractability of the young horses was assessed using standardised tests. Responses to a novel object, to a novel person, and during a handling test were observed and quantified. Horses grew well on both diets with no apparent effects of diet on growth rate or baseline behaviour. Immediately after weaning, horses receiving the FF diet cantered less frequently (F = 5.10; p < 0.05), for a shorter duration (F = 7.23; p < 0.05) and appeared to be more settled. Foals that were barn-weaned appeared more stressed than foals that were paddock-weaned. In the temperament tests, horses receiving the FF diet spent significantly more time investigating (F = 6.78; p < 0.05), and less time looking at (F = 7.93; p < 0.05), the novel object than horses receiving the SS diet. They also spent less time walking away from the novel person (F = 5.16; p < 0.05) and their time taken to complete the handling test was significantly lower (F = 8.72; p = 0.01). Overall, the horses that received the FF diet appeared less distressed immediately after weaning and seemed calmer and more inquisitive during a range of temperament tests.
Keywords: Horse; Diet; Weaning; Temperament test
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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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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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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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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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Abeyesinghe, S. M., Nicol, C. J., Wathes. C.M., & Randall, J. M. (2001). Development of a raceway method to assess aversion of domestic fowl to concurrent stressors. Behav. Process., 56(3), 175–194.
Abstract: The requirement for assessing the effects of stressor combinations in improving the welfare of animals has not been widely recognised. Knowledge of the effects of concurrent stressors is needed to improve environments such as transport, where animals are presented with many simultaneous challenges. However, no method for measuring the effects of different stressors with a common unit is currently available. A locomotor passive avoidance method was developed as a common currency measure of the aversion of domestic fowl to concurrent stressors, using vibrational and thermal stressors as an exemplar. Juvenile fowl, fasted overnight, were trained to run a raceway into a goal-box for small food rewards (FR1). When running consistently, the reinforcement schedule was superimposed with a FR5 treatment schedule (60 min confinement in the goal-box with either a control of no other stressors [N] or concurrent vibration and thermal stressors [VT]). Subsequent latency to return to the goal-box was recorded as a measure of aversion. The factors affecting bird response were addressed in a series of experiments to optimise the method and clarify interpretation of results. Pre-feeding (20% ration 2 h prior to testing) did not affect response, but increasing the number of treatment presentations facilitated learning and increased method sensitivity. Treatment responses were consistent across experiments; overall VT was avoided (P<0.001), but N was not. However, there was large individual variation in response to VT. A final experiment indicated that, given a visual discriminatory cue, birds were capable of learning the required association between entering the goal-box and receiving the treatment, suggesting that the delay responses were due to aversion rather than the immediate impact of treatment on ability to respond. Further work is required to test the singular stressors, but the method retains common currency potential for assessing aversion to multiple stressors.
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Hothersall, B., & Nicol, C. (2007). Equine learning behaviour: accounting for ecological constraints and relationships with humans in experimental design. Behav. Process., 76(1), 45–48.
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