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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.
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Leiner, L., & Fendt, M. (2011). Behavioural fear and heart rate responses of horses after exposure to novel objects: Effects of habituation. Appl. Anim. Behav. Sci., 131(3-4), 104–109.
Abstract: The emotion fear promotes the fitness of wild animals. In a farm environment, exaggerated fear, e.g., in horses, can cause several problems. Therefore, knowledge about fear in horses helps to prevent or to handle potential fear-inducing situations. The present study investigated which behavioural fear responses can be observed during exposure of horses to a novel stimulus, whether these behavioural responses are correlated with physiological changes, and whether and how specifically these changes are reduced after habituation training to one of the novel objects. Our data shows that behavioural and physiological fear responses in horses are correlated, are reliable to observe and to measure, and appear in a typical chronological order. Furthermore, after habituation-training to an object, the fear response to this object is specifically attenuated whereas the fear response to another object remains.
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Lee, J., Floyd, T., Erb, H., & Houpt, K. (2011). Preference and demand for exercise in stabled horses. Appl. Anim. Behav. Sci., 130(3-4), 91–100.
Abstract: Operant conditioning and two choice preference tests were used to assess the motivation of horses to be released from straight and from box stalls. The motivations for food, a companion, and release into a paddock were compared when the horses had to work for each commodity at increasing fixed ratios of responses (panel presses) to reward in an equine operant conditioning stall. The motivation for food (mean ± SEM = 258 ± 143) responses was much greater than that for either release (38 ± 32) from a straight stall into a large paddock alone or into a small paddock with another horse (95 ± 41) (P = 0.04). When given a two choice preference test between exercise on a treadmill for 20 min or returning to their box stalls, eight of nine horses chose to return to their stalls. In a two choice preference test six of eight horses in box stalls chose to be released into a paddock alone. Horses were given a series of two choice preference tests to determine how long they preferred to be in a paddock. After 15 min in the paddock the horses were re-tested, but all chose the paddock when released into a paddock with three other horses. They were retested every 15 min until they chose to return to their stalls. They chose to stay out for 35 ± 6 min when other horses were in the paddock but for only 17 ± 2 min when they would be alone. When deprived of stall release for 48 h the horses chose to remain in the paddock with other horses for 54 ± 6 min, but showed no compensatory behavior when they were alone (duration chosen = 16 ± 4 min). These findings indicate that horses are not strongly motivated to exercise alone and will choose not to endure forced exercise on a treadmill. The social context of voluntary exercise is important; horses are willing to stay out of their stalls longer if other horses are present and will show compensatory behavior only if other horses are present. These finding have implications for optimizing turnout time for stalled horses.
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Christensen, J. W., Zharkikh, T., & Chovaux, E. (2011). Object recognition and generalisation during habituation in horses. Appl. Anim. Behav. Sci., 129(2-4), 83–91.
Abstract: The ability of horses to habituate to frightening stimuli greatly increases safety in the horse-human relationship. A recent experiment suggested, however, that habituation to frightening visual stimuli is relatively stimulus-specific in horses and that shape and colour are important factors for object generalisation (Christensen et al., 2008). In a series of experiments, we aimed to further explore the ability of horses (n = 30, 1 and 2-year-old mares) to recognise and generalise between objects during habituation. TEST horses (n = 15) were habituated to a complex object, composed of five simple objects of varying shape and colour, whereas CONTROL horses (n = 15) were habituated to the test arena, but not to the complex object. In the first experiment, we investigated whether TEST horses subsequently reacted less to i) simple objects that were previously part of the complex object (i.e. testing for object recognition) and ii) a novel object (new shape and colour, i.e. testing for object generalisation), compared to CONTROLS. In the second experiment we investigated whether TEST horses reacted to a change in object order and object location. Behavioural reactions to the object, latency to eat, total eating time and heart rate were recorded. Compared to CONTROLS, TEST horses reacted significantly less towards objects, which were previously part of the complex object (e.g. mean heart rate; P = 0.006), indicating object recognition. In contrast to our expectations, TEST horses also reacted significantly less towards the novel object (e.g. mean heart rate; P = 0.018), suggesting that they were capable of object generalisation. We also found that TEST horses showed an increase in exploratory behaviour when objects within the complex object changed order and location (both P < 0.001), whereas there was no increase in heart rate, indicating that the horses were not frightened by the changes. The results demonstrate that it is possible to increase object generalisation in horses by habituating them to a range of colours and shapes simultaneously. This knowledge greatly affects the way in which horses may be trained to react calmly towards frightening objects.
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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.
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Ransom, J. I., Cade, B. S., & Hobbs, N. T. (2010). Influences of immunocontraception on time budgets, social behavior, and body condition in feral horses. Appl. Anim. Behav. Sci., 124(1-2), 51–60.
Abstract: Managers concerned with shrinking habitats and limited resources for wildlife seek effective tools for limiting population growth in some species. Fertility control is one such tool, yet little is known about its impacts on the behavioral ecology of wild, free-roaming animals. We investigated influences of the immunocontraceptive porcine zona pellucida (PZP) on individual and social behavior in bands of feral horses (Equus caballus) in three discrete populations and used 14 hierarchical mixed effect models to gain insight into the influences of PZP treatment on feral horse behavior. A model of body condition was the strongest predictor of feeding, resting, maintenance, and social behaviors, with treated females allocating their time similarly to control females. Time spent feeding declined 11.4% from low condition to high condition females (F1,154 = 26.427, P < 0.001) and was partially reciprocated by a 6.0% increase in resting (F1,154 = 7.629, P = 0.006), 0.9% increase in maintenance (F1,154 = 7.028, P = 0.009), and 1.8% increase in social behavior (F1,154 = 15.064, P < 0.001). There was no difference detected in body condition of treated versus control females (F1,154 = 0.033, P = 0.856), but females with a dependent foal had lower body condition than those without a foal (F1,154 = 4.512, P = 0.038). Herding behavior was best explained by a model of treatment and the interaction of band fidelity and foal presence (AICc weight = 0.660) which estimated no difference in rate of herding behavior directed toward control versus treated females (F1,102 = 0.196, P = 0.659), but resident females without a dependent foal were herded 50.9% more than resident females with a foal (F3,102 = 8.269, P < 0.001). Treated females received 54.5% more reproductive behaviors from stallions than control mares (F1,105 = 5.155, P = 0.025), with the model containing only treatment being the most-supported (AICc weight = 0.530). Treated and control females received harem-tending behaviors from stallions equally (F1,105 = 0.001, P = 0.969) and agonistic behaviors from stallions equally (F1,105 < 0.001, P = 0.986). Direct effects of PZP treatment on the behavior of feral horses appear to be limited primarily to reproductive behaviors and most other differences detected were attributed to the effects of body condition, band fidelity, or foal presence. PZP is a promising alternative to traditional hormone-based contraceptives and appears to contribute few short-term behavioral modifications in feral horses.
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Stachurska, A., Pieta, M., Ussing, A. P., Kapron, A., & Kwiecinska, N. (2010). Difficulty of cross-country obstacles for horses competing in Three Day Events. Appl. Anim. Behav. Sci., 123(3-4), 101–107.
Abstract: The objective of the study was to determine which cross-country obstacles are more difficult for eventing horses. Jumping scores were considered in terms of the horse's reaction to novelty and to the fearfulness of novel objects which are the fences situated in novel terrain. The data concerned 11 classes of One to Four Star level (stars showing the difficulty of the class) held at the Olympic Games and three international Three Day Events. A total of 400 entries, in which 259 horses jumped 372 obstacles were considered. Scores of 11,341 jumps at particular fences were categorized either as faulty jumps or non-faulty jumps. Factors describing the fences versus the jumping scores were studied with least square analysis of variance, with respect to the interaction between the star level and the fence traits. The overall frequency of faults at the cross-country amounted to 4.33 ± 0.57%. Among the effects analyzed, the difficulty of cross-country fences for the horses depends upon whether an obstacle is single or is an element of a combination, whether it is straight or requiring an effort in both height and spread, is broad or narrow, has a solid top or a brush, has an alternative or not and whether it is a water crossing or not. At One Star level, the less experienced horses react differently to certain fence traits compared to horses participating in Three or Four Star levels. It is concluded that the equine visionary system, being less developed towards identification of stationary objects than the human visionary system may be a key towards explaining the horse's behaviour while jumping the obstacle. The height-spread obstacles, those of the narrow front, with the brush and with the alternative seem to involve more faults because of the equine low-acuity vision. The same reason may justify the similar frequency of faults at the single obstacles and the first elements in combinations. Accurate methods of measuring equine vision would complement behavioural tests and should both be introduced into the selection of eventing horses.
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Jørgensen, G. H. M., Borsheim, L., Mejdell, C. M., Søndergaard, E., & Bøe, K. E. (2009). Grouping horses according to gender--Effects on aggression, spacing and injuries. Appl. Anim. Behav. Sci., 120(1-2), 94–99.
Abstract: Many horse owners tend to group horses according to gender, in an attempt to reduce aggressive interactions and the risk of injuries. The aim of our experiment was to test the effects of such gender separation on injuries, social interactions and individual distance in domestic horses. A total of 66 horses were recruited from 4 different farms in Norway and Denmark and divided into six batches. Within each batch, horses were allotted into one mare group, one gelding group and one mixed gender group, with most groups consisting of three or four animals. After 4-6 weeks of acclimatisation, a trained observer recorded all social interactions using direct, continuous observation 1 h in the morning and 1 h in the afternoon for three consecutive days. Recordings of the nearest neighbour of each horse were performed using instantaneous sampling every 10 min. The horses were inspected for injuries before grouping, day 1 after grouping and after 4-6 weeks. No significant effect of gender composition was found on social interactions (P > 0.05), spacing (P > 0.07) or injuries (P > 0.23). Eighty percent of all aggressive interactions recorded were threats, not involving physical contact. Horses with the smallest space allowance showed the highest mean number of aggressive interactions (28.6 ± 6.1 interactions per 6 h) compared to the mean of all the other batches (8.3 ± 1.0 interactions per 6 h). Very few injuries were found and most were superficial. In conclusion, gender composition does not seem to have any effect on aggression level, spacing or injuries. However, the early social experience of horses, management of feeding and space allowance probably represents more important factors for successful group housing of domestic horses.
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Edouard, N., Fleurance, G., Dumont, B., Baumont, R., & Duncan, P. (2009). Does sward height affect feeding patch choice and voluntary intake in horses? Appl. Anim. Behav. Sci., 119(3-4), 219–228.
Abstract: The numbers of horses grazing at pasture are increasing in developed countries, so a proper understanding of their feeding selectivity and of the tactics they use for extracting nutrients from swards is essential for the management of horses and grasslands. Resource acquisition in herbivores can be optimised through the modulation of their intake and patch selection, both being strongly dependent on the characteristics of swards. However, the principles by which horses adjust their grazing behaviour in response to variations in sward features are not completely understood. The aim of this study was to determine whether the behaviour of horses conforms to optimal foraging models. We hypothesized that, faced with binary choices between vegetative swards of a good and similar quality at three different heights, horses would select the taller sward, i.e. that allowing a higher reward in terms of dry matter intake rate. Three groups of three 2-year-old saddle horses were grazed on a semi-natural pasture that was managed to produce three contrasting sward heights at 6, 11 and 17 cm, in a Latin-square design. The instantaneous intake rate was determined from bite rate measured at pasture on the three sward heights, and bite mass estimated from measurements using swards offered indoors in experimental trays. Daily dry matter intake was estimated individually by total faecal collection and an estimation of digestibility from faecal nitrogen. Short-term (first 30 min) and daily preferences were assessed from the time spent grazing each sward offered in pair-wise tests at pasture. The results show that daily voluntary intake (an average of 21 g DM kg LW-1 day-1) and total grazing time (an average of 14 h day-1) were independent of sward height and of the choice of patches offered. In choice situations, the animals spent more time grazing on the taller sward, both during the first 30 min and at the daily scale. These results show that horses choose between vegetative patches of a good and similar quality according to the predictions from optimal foraging models, and select the one that they can ingest faster. Further research will now have to explore how the horses will adapt their feeding behaviour when they face a trade-off between sward height and quality.
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Nuñez, C. M. V., Adelman, J. S., Mason, C., & Rubenstein, D. I. (2009). Immunocontraception decreases group fidelity in a feral horse population during the non-breeding season. Appl. Anim. Behav. Sci., 117(1-2), 74–83.
Abstract: The behavioral effects of the immunocontraceptive agent porcine zona pellucida (PZP) have not been adequately studied. Important managerial decisions for several species, including the wild horse (Equus caballus), have been based on this limited research. We studied 30 horses on Shackleford Banks, North Carolina, USA to determine the effects of PZP contraception on female fidelity to the harem male. We examined two classes of females: contracepts, recipients of the PZP vaccine (n = 22); and controls, females that have never received PZP (n = 8). We conducted the study during the non-breeding season from December 2005 to February 2006, totaling 102.2 h of observation. Contracepted mares changed groups more often than control mares (P = 0.04). Contracepts also visited more harem groups than did control mares (P = 0.02) and exhibited more reproductive interest (P = 0.05). For both contracepted and control females, the number of group changes (P = 0.01) and number of groups visited (P = 0.003) decreased with the proportion of years mares were pregnant. Our study shows that the application of PZP has significant consequences for the social behavior of Shackleford Banks horses. In gregarious species such as the horse, PZP application may disrupt social ties among individuals and inhibit normal social functioning at the population level.
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