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Christensen, J. W., Zharkikh, T., & Ladewig, J. (2008). Do horses generalise between objects during habituation? Appl. Anim. Behav. Sci., 114(3-4), 509–520.
Abstract: Habituation to frightening stimuli plays an important role in horse training. To investigate the extent to which horses generalise between different visual objects, 2-year-old stallions were habituated to feeding from a container placed inside a test arena and assigned as TEST (n=12) or REFERENCE horses (n=12). In Experiment 1, TEST horses were habituated to six objects (ball, barrel, board, box, cone, cylinder) presented in sequence in a balanced order. The objects were of similar size but different colour. Each object was placed 0.5m in front of the feed container, forcing the horses to pass the object to get to the food. TEST horses received as many 2min exposures to each object as required to meet a habituation criterion. We recorded behavioural reactions to the object, latency to feed, total eating time, and heart rate (HR) during all exposures. There was no significant decrease in initial responses towards a novel object with increasing object number, indicating that habituation was stimulus-specific. REFERENCE horses were exposed to the test arena without objects in the same period, and reactions of TEST and REFERENCE horses were subsequently compared in a fear-test (2m2.5m rubber mat placed under the feed container, forcing the horses to step on the mat to get food). There were no significant differences between the treatment groups, i.e. previous habituation of TEST horses to six visual objects did not reduce responses in a fear-test involving visual and tactile stimulation. Due to the lack of generalisation in Experiment 1, we designed a supplemental experiment (Experiment 2), in which REFERENCE horses were exposed to the same six objects except that object colour was kept constant. We found a significant reduction in response (behaviour and HR) with increasing object number, indicating that horses generalise between similarly coloured objects of varying shape. We conclude that a high degree of object similarity, e.g. identical colouring, appears to be crucial for object generalisation in horses.
Keywords: Horse; Stimulus generalisation; Habituation; Fear
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Christensen, J. W., & Rundgren, M. (2008). Predator odour per se does not frighten domestic horses. Appl. Anim. Behav. Sci., 112(1-2), 136–145.
Abstract: Horses frequently react nervously when passing animal production farms and other places with distinctive smells, leading riders to believe that horses are innately frightened by certain odours. In three experiments, we investigated how horses respond to (1) urine from wolves and lions, (2) blood from slaughtered conspecifics and fur-derived wolf odour, and (3) a sudden auditory stimulus in either presence or absence of fur-derived wolf odour. The experiments were carried out under standardised conditions using a total of 45 naive, 2-year-old horses. In the first two experiments we found that horses showed significant changes in behaviour (Experiments 1 and 2: increased sniffing; Experiment 2 only: increased vigilance, decreased eating, and more behavioural shifts), but no increase in heart rate compared to controls when exposed to predator odours and conspecific blood in a known test environment. However, the third experiment showed that exposure to a combination of wolf odour and a sudden stimulus (sound of a moving plastic bag) caused significantly increased heart rate responses and a tendency to a longer latency to resume feeding, compared to control horses exposed to the sudden stimulus without the wolf odour. The results indicate that predator odour per se does not frighten horses but it may cause an increased level of vigilance. The presence of predator odour may, however, cause an increased heart rate response if horses are presented to an additional fear-eliciting stimulus. This strategy may be adaptive in the wild where equids share habitats with their predators, and have to trade-off time and energy spent on anti-predation responses against time allocated to essential non-defensive activities.
Keywords: Horse; Predator odour; Behaviour; Heart rate; Fear
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Christensen, J. W., Keeling, L. J., & Nielsen, B. L. (2005). Responses of horses to novel visual, olfactory and auditory stimuli. Appl. Anim. Behav. Sci., 93(1-2), 53–65.
Abstract: Responses of horses towards fear-eliciting stimuli can have important consequences for both human and horse safety. This experiment was designed to investigate behavioural and heart rate (HR) responses of horses to novel visual, auditory and olfactory stimuli. Twenty-four 2-year-old, previously unhandled, stallions were habituated to receive a food reward from a container in a test arena. Each horse was exposed to three 2 min tests in a balanced design where in addition to the feed container, either a traffic cone (visual test), white noise (auditory test) or eucalyptus oil applied to the inside of the container (olfactory test) were used as the novel stimuli. Compared to the control, less time was spent eating during all tests. There was no difference in locomotion activity in the different test situations, but presentation of the novel visual and auditory stimuli elicited significantly increased HR responses in the horses, compared to their response to the arena without novel stimuli (control), whereas there was no increase in HR response to the olfactory stimulus. However, during the olfactory test, the horses had an increased number of eating bouts and became more vigilant towards their surroundings, whereas during the visual and auditory tests, more time was spent alert towards the stimulus. The horses also took significantly more steps backwards in response to the auditory test. The heart rate responses correlated between tests and reflect a non-differentiated activation of the sympathetic nervous system, while the behavioural responses were linked to the type of stimulus.
Keywords: Horses; Novelty responses; Fear; Behaviour; Heart rate
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Christensen, J. W., Zharkikh, T., Ladewig, J., & Yasinetskaya, N. (2002). Social behaviour in stallion groups (Equus przewalskii and Equus caballus) kept under natural and domestic conditions. Appl. Anim. Behav. Sci., 76(1), 11–20.
Abstract: The aim of this study was to investigate social behaviour in differently reared stallions in their respective environments; one group of stallions was reared under typical domestic conditions whereas the other group was reared and lives under natural conditions. The domestic group consisted of 19, 2-year-old stallions (Equus caballus), which were all weaned at 4 months of age and experienced either individual or group housing facilities before being pastured with the other similarly aged stallions. The natural living and mixed age group of Przewalski stallions (E. przewalskii) consisted of 13 stallions, most of which were juveniles (n=11, <=4 years; n=2, >9 years). The domestic group was studied in a 4-ha enclosure at the Danish Institute of Agricultural Sciences and the Przewalski group under free-ranging conditions in a 75-ha enclosure in the Askania Nova Biosphere Reserve, Ukraine. Behavioural data was collected during 168 h of direct observation. The occurrence of 14 types of social interactions was recorded and group spacing behaviour was studied using nearest neighbour recordings. In spite of very different environments, reflecting domestic and natural rearing conditions, many similarities in behaviour was found. Play and play fight behaviour was very similar in the two stallion groups. Quantitative differences were found in social grooming since Przewalski stallions groomed more frequently (P=0.004), and in investigative behaviours, since domestic stallions showed more nasal (P=0.005) and body sniffing (P<0.001), whereas Przewalski stallions directed more sniffing towards the genital region (P<0.001). These differences may, however, be attributed to environmental factors and in the period of time the stallions were together prior to the study period. Quantitative differences appeared in some agonistic behaviours (kick threat, P<0.001; and kick, P<0.001), but data do not support earlier findings of Przewalski horses being significantly more aggressive than domestic horses. In general, Przewalski stallions engaged in more social interactions, and they showed less group spacing, i.e. maintained a significantly shorter distance between neighbours (P<0.001). The study indicates that also domestic horses, which have been reared under typical domestic conditions and allowed a period on pasture, show social behaviour, which is very similar to that shown by their non-domestic relatives.
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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.
Keywords: Horse; Individual stabling; Group stabling; Social behaviour
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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., Rundgren, M., & Olsson, K. (2006). Training methods for horses: habituation to a frightening stimulus. Equine Vet J, 38(5), 439–443.
Abstract: REASONS FOR PERFORMING STUDY: Responses of horses in frightening situations are important for both equine and human safety. Considerable scientific interest has been shown in development of reactivity tests, but little effort has been dedicated to the development of appropriate training methods for reducing fearfulness. OBJECTIVES: To investigate which of 3 different training methods (habituation, desensitisation and counter-conditioning) was most effective in teaching horses to react calmly in a potentially frightening situation. HYPOTHESES: 1) Horses are able to generalise about the test stimulus such that, once familiar with the test stimulus in one situation, it appears less frightening and elicits a reduced response even when the stimulus intensity is increased or the stimulus is presented differently; and 2) alternative methods such as desensitisation and counter-conditioning would be more efficient than a classic habituation approach. METHODS: Twenty-seven naive 2-year-old Danish Warmblood stallions were trained according to 3 different methods, based on classical learning theory: 1) horses (n = 9) were exposed to the full stimulus (a moving, white nylon bag, 1.2 x 0.75 m) in 5 daily training sessions until they met a predefined habituation criterion (habituation); 2) horses (n = 9) were introduced gradually to the stimulus and habituated to each step before the full stimulus was applied (desensitisation); 3) horses (n = 9) were trained to associate the stimulus with a positive reward before being exposed to the full stimulus (counter-conditioning). Each horse received 5 training sessions of 3 min per day. Heart rate and behavioural responses were recorded. RESULTS: Horses trained with the desensitisation method showed fewer flight responses in total and needed fewer training sessions to learn to react calmly to test stimuli. Variations in heart rate persisted even when behavioural responses had ceased. In addition, all horses on the desensitisation method eventually habituated to the test stimulus whereas some horses on the other methods did not. CONCLUSIONS AND POTENTIAL RELEVANCE: Desensitisation appeared to be the most effective training method for horses in frightening situations. Further research is needed in order to investigate the role of positive reinforcement, such as offering food, in the training of horses.
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Rørvang, M. V., Ahrendt, L. P., & Christensen, J. W. (2015). Horses fail to use social learning when solving spatial detour tasks. Anim.Cogn., 18(4), 847–854.
Abstract: Social animals should have plenty of opportunities to learn from conspecifics, but most studies have failed to document social learning in horses. This study investigates whether young Icelandic horses can learn a spatial detour task through observation of a trained demonstrator horse of either the same age (Experiments 1 and 2, n = 22) or older (Experiment 3, n = 24). Observer horses were allowed to observe the demonstrator being led three times through the detour route immediately before being given the opportunity to solve the task themselves. Controls were allowed only to observe the demonstrator horse eating at the final position, but not the demonstration of the route. Although we found a tendency towards better performance by observer horses in the second experiment, we were unable to repeat this result in a similar set-up with a new group of horses and older, dominant demonstrator horses. We conclude that horses exposed to prior demonstration did not perform better than control horses in solving spatial detour tasks.
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Rørvang, M. V., Christensen, J. W., Ladewig, J., & McLean, A. (2018). Social Learning in Horses--Fact or Fiction? Front. Vet. Sci., 5, 212.
Abstract: Prima facie, the acquisition of novel behaviors in animals through observation of conspecifics seems straightforward. There are, however, various mechanisms through which the behavior of animals can be altered from observing others. These mechanisms range from simple hard-wired contagious processes to genuine learning by observation, which differ fundamentally in cognitive complexity. They range from social facilitation and local enhancement to true social learning. The different learning mechanisms are the subject of this review, largely because research on learning by observation can be confounded by difficulties in interpretation owing to the looming possibility of associative learning infecting experimental results. While it is often assumed that horses are capable of acquiring new behavior through intra-species observation, research on social learning in horses includes a variety of studies some of which may overestimate the possession of higher mental abilities. Assuming such abilities in their absence can have welfare implications, e.g. isolating stereotypical horses on the assumption that these behaviors can be learned though observation by neighboring horses. This review summarizes the definitions and criteria for the various types of social transmission and social learning and reviews the current documentation for each type in horses with the aim of clarifying whether horses possess the ability to learn through true social learning. As social ungulates, horses evolved in open landscapes, exposed to predators and grazing most of the day. Being in close proximity to conspecifics may theoretically offer an opportunity to learn socially, however anti-predator vigilance and locating forage may not require the neural complexity of social learning. Given the significant energetic expense of brain tissue, it is likely that social facilitation and local enhancement may have been sufficient in the adaptation of equids to their niche. As a consequence, social learning abilities may be maladaptive in horses. Collectively, the review proposes a novel differentiation between social transmission (social facilitation, local and stimulus enhancement) and social learning (goal emulation, imitation). Horses are undoubtedly sensitive to intra-species transfer of information but this transfer does not appear to satisfy the criteria for social learning, and thus there is no solid evidence for true social learning in horses.
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Christensen, J. W. (2012). Object habituation in horses: Voluntary vs. negatively reinforced approach to frightening stimuli. In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: The ability and ease of horses to habituate to frightening stimuli greatly increases safety in the horse-human relationship. Several different techniques have been suggested for habituation training of horses and under certain conditions, preventing animals from avoidance reactions during exposure to frightening stimuli is believed to facilitate habituation. Response prevention does, however, lead to a loss of control, which is a known stress inducer in both animals and humans. This experiment investigated whether horses show increased stress responses when negatively reinforced to approach a mildly frightening stimulus, compared to horses allowed to voluntarily explore the same stimulus. We further investigated whether the prevention of avoidance responses in horses that are negatively reinforced to approach the stimulus, facilitates habituation to the stimulus. Twenty-two 2-3 years old Danish warmblood geldings were included in the study. Half of the horses (NR group) were negatively reinforced (through halter and rope pressure) by a familiar human handler to approach a collection of frightening objects (six open and colourful umbrellas) placed in a semi-circle in a familiar test arena. The other half of the horses were released in the arena and were free to avoid or explore the objects (VOL group). On the next day, all horses were exposed to the objects again without a human to investigate the rate of habituation. Behavioural and heart rate responses were recorded on both days. Data were analysed in a two way repeated measures ANOVA and post hoc analysed via the Holm-Sidak method. In the VOL group, all horses initially chose to avoid the unknown objects, whereas the handler managed to get all horses in the NR group to approach and stand next to the objects within the first 2-min session. As expected, horses in the NR group had a significantly longer duration of alertness (sec, mean ± se: NR: 23 ± 4.1 vs. VOL: 16 ± 4.7, P=0.026) and a higher max HR in the first session (bpm, mean ± se: NR: 106 ± 5.2 vs. VOL: 88 ± 4.4, P=0.004). On the next day, however, the NR horses spent significantly less time investigating the objects (sec, mean ± se: NR: 13 ± 4.1 vs. VOL: 24 ± 6.0, P=0.005) and had a shorter latency to approach a feed container, placed next to the objects (sec, mean ± se: NR: 25 ± 3.9 vs. VOL: 47 ± 16.2, P=0.031), indicating increased habituation. In conclusion, negatively reinforced approach to mildly frightening objects appears to increase stress responses during the initial exposure, but also to facilitate habituation in young horses.
Keywords: Habituation; learning; fearfulness; stress; reinforcement
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