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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. (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.
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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.
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Lindberg, A. C., Kelland, A., & Nicol, C. J. (1999). Effects of observational learning on acquisition of an operant response in horses. Appl. Anim. Behav. Sci., 61(3), 187–199.
Abstract: The effect of observational learning on the acquisition of an operant response was examined in eighteen riding horses and ponies. The test horses were randomly divided into three groups of six and individually exposed to one of three treatments. An additional horse was trained as a demonstrator, to perform the operant response. The observer horses watched either the demonstrator performing the bin-opening response (Group D+B); the demonstrator standing passively (Group D); or the operant bin in the absence of the demonstrator (Group B). Observers had access to and were free to interact with an identical bin during testing. Observers in Groups D+B and D were socially familiar with the demonstrator. Each test horse was tested once a day for 10 days. An ANOVA revealed no significant differences between treatment groups in the number of responses or the time taken to reach the learning criterion. However, there were highly significant differences between breed types, with non-warmbloods performing more bouts of opening the bin and feeding (p=0.02), feeding from the bin sooner (p=0.01) and reaching the criterion for learning sooner than warmbloods (p=0.05). There was also a significant negative linear relationship between horses' ages and time spent investigating the bin, with younger horses performing more investigative behaviour (y=-3.08x+106.86; p=0.02).
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Murphy, J., Sutherland, A., & Arkins, S. (2005). Idiosyncratic motor laterality in the horse. Appl. Anim. Behav. Sci., 91(3-4), 297–310.
Abstract: Idiosyncratic motor behaviour was investigated during four experimental procedures in 40 horses (males = 20, females = 20) to establish if horses revealed evidence of significant right or left laterality. The experimental procedures included (1) detection of the preferred foreleg to initiate movement (walk or trot), (2) obstacle avoidance within a passageway (right or left), (3) obstacle avoidance when ridden and (4) idiosyncratic motor bias when rolling. The influence of the horses' sex on both the direction and the degree of the laterality was explored within and between experimental procedures. The findings showed that the direction, but not the degree of idiosyncratic motor preference in the horses was strongly sex-related. Male horses exhibited significantly more (t = 3.74, d.f. = 79, P < 0.001) left lateralised responses and female horses exhibited significantly more (t = -6.35, d.f. = 79, P < 0.01) right lateralised responses. There was also significant positive correlation (P < 0.05) between four of six possible inter-experimental relationships. The results suggest two discrete trends of laterality associated with the sex of the horse. The primary cause of idiosyncratic motor laterality may be genetically predetermined, influenced by environmental factors or a combination of these two and the current findings may support the development of sex-specific training schedules for the horse. Further, work in this area might assist in defining the mechanisms of brain hemisphere lateralisation and allocation of cognitive function in the horse.
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Seaman, S. C., Davidson, H. P. B., & Waran, N. K. (2002). How reliable is temperament assessment in the domestic horse (Equus caballus)? Appl. Anim. Behav. Sci., 78(2-4), 175–191.
Abstract: Differences in behavioural characteristics between individuals of the same species are often described as being due to the temperament of the individuals. These differences can have enormous implications for welfare with some individuals apparently being able to adapt to environmental challenge more easily than others. Such differences have resulted in animals often being described as either `active' copers, which try to escape from or remove an aversive stimulus, or `passive' copers, which show no outward signs of a situation being aversive, thus, appearing to be unaffected. Tests previously developed to assess the temperament of animals have been criticised for several reasons. Behaviour is often recorded and categorised using methods that are not objective and tests are generally carried out once with no consideration of whether or not behavioural responses are consistent over time. This study takes these factors into account. The behaviour of 33 horses was recorded in three types of test--an arena test, response to a person and response to an object. In order to test whether or not responses were consistent over time, the tests were repeated three times with an average of 9 days between trials. Test results were validated using responses from questionnaires completed by the farm team leader. The data were analysed using an initial principal component analysis (PCA) and factor analysis. The horses were found to behave consistently over the three trials in their responses in the arena test. The responses to the person test and the object test were similar to each other; however, these responses were not consistent over trials. The behaviour in the arena test was unable to be used to make a prediction of behaviour in the person and object tests and vice versa. The responses shown by the horses did not enable them to be categorised as either active or passive copers. Behavioural responses in the tests were not predictive of the response to a startle test (water spray), nor could they be used to predict status or response to being reintroduced to the group after testing. There was no relationship between the responses in the tests and the ratings given by the farm team leader. It was concluded that horses vary widely in their responses to artificial behavioural tests, with only the responses to an open-field arena test being consistent over time, and therefore, the only type of test which can indicate some core factor of temperament.
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Marinier, S. L., Alexander, A. J., & Waring, G. H. (1988). Flehmen behaviour in the domestic horse: Discrimination of conspecific odours. Appl. Anim. Behav. Sci., 19(3-4), 227–237.
Abstract: American Saddlebred horses were used to test the responses of domestic horses to the odours of conspecifics. In all cases the odours were tested in the absence of the donor animal. Thus the test animal's behavioural responses were concentrated on the olfactory stimuli, and possible interference from donor behaviour was eliminated. Stallions were significantly more responsive than mares and geldings. This was shown in both flehmen and sniffing behaviour to urine/vaginal secretions and in sniffing behaviour to faecal samples. Only stallions were used for subsequent tests. Stallions showed no significant differences in response to the odour of urine/vaginal secretions of an oestrus mare from that when she was not in season. Parameters used for analysis of data were frequency, latency and duration of flehmen as well as duration of responsiveness to samples. In testing for differences in odours between individual mares, two methods were used. The stallions differentiated between samples from individual mares. In some cases this differentiation was exhibited when the stallions were merely presented with the two samples in sequence. In other cases statistically significant differences in response to the odours were shown only by simultaneous presentation of the two samples to the test stallion. Parameters used for data analysis were frequency and duration of flehmen and duration of responsiveness.
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Gorecka, A., Golonka, M., Chruszczewski, M., & Jezierski, T. (2007). A note on behaviour and heart rate in horses differing in facial hair whorl. Appl. Anim. Behav. Sci., 105(1-3), 244–248.
Abstract: The relationship between facial hair whorl position and reactivity, as assessed by behavioural measures (handling score = HS; startle reaction to a suddenly appearing novel object = SR; latency to touch a novel object = LNO) and heart rate measures (mean HR; increase in heart rate = IHR) were studied using 55 Konik horses reared either under conventional stable conditions or in the forest reserve. Horses were classified into four groups according to the whorl position and/or shape: (1) high, single whorl above the top eye line, n = 9; (2) medium, single whorl between the top and the bottom eye line, n = 30; (3) low, single whorl below the bottom eye line, n = 10; and (4) elongated or double whorl, n = 6. Horses with a high whorl position demonstrated a lesser degree of manageability as expressed by a lower HS compared to individuals with medium (P = 0.002) or low whorl positions (P = 0.016). Horses with different whorl positions did not differ significantly in their startle response to a suddenly appearing novel object (P = 0.685). The horses with an elongated or double whorl, which appeared only in the forest group, took significantly longer to approach the novel object than horses with medium (P = 0.006) or low (P = 0.005) whorl positions. No significant differences in mean HR and IHR between groups (HR: P = 0.629 and IHR: P = 0.214) were found. In conclusion, this study supports the relationship between the position of the hair whorl on the horses' head and their manageability during handling, as well as the latency to approach an unknown object.
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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.
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McLean, A. N. (2004). Short-term spatial memory in the domestic horse. Appl. Anim. Behav. Sci., 85(1-2), 93–105.
Abstract: This study investigates the ability of horses to recall a feeding event in a two-point choice apparatus. Twelve horses were individually tested whereby they were maintained immobile in a test arena and visually and aurally experienced the delivery of food into one of two feed goals. The horses were then released to make their choice in two experimental contexts: immediate release after experiencing the delivery of food, and release 10 s after food delivery. Each horse performed 40 immediate-release (IR) trials, followed by forty 10-s release trials over a 3-day period. In addition, the same horses were tested 3 months later in the spring with the same number and sequence of trials. Results were analysed by log-linear analysis of frequencies. Results showed that while horses were able to achieve the correct feed goal choice in the immediate-release trials, they were unsuccessful with the 10-s release trials. This suggests that there are limitations in recall abilities in horses, in that they may not possess a prospective type of memory. There are welfare and training implications in these findings concerning the effects of overestimating the mental abilities of horses during training and the effects of delays in reinforcements.
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