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Jankunis, E. S., & Whishaw, I. Q. (2013). Sucrose Bobs and Quinine Gapes: Horse (Equus caballus) responses to taste support phylogenetic similarity in taste reactivity. Behavioural Brain Research, 256, 284–290.
Abstract: Evidence suggests that behavioural affective reactions to sweet and bitter substances are homologous in humans, nonhuman primates, and rodents. The sweet taste of sucrose elicits facial responses that include rhythmic tongue protrusions whereas the bitter taste of quinine elicits facial responses that include gapes, featuring an opening of the mouth and protrusion of the tongue. The present study using the horse (Equus caballus) was undertaken for three reasons: (1) there is debate about the presence of a sweet receptor gene in the horse, (2) there is a need to expand the examination of facial reactions to taste in lineages other than the closely related lineages of rodents and primates, and (3) the horse provides an opportunity to test the hypothesis that some social signals derive from movements related to taste reaction. The horses were given oral infusions of either sucrose or quinine and their behaviour was examined using frame-by-frame video analysis. Control groups were exposed received water or syringe insertion only. Amongst the many responses made to the infusions, the distinctive response to sucrose was a bob coupled with a slight tongue protrusion and forward movement of the ears; the distinctive response to quinine was a head extension and mouth gape accompanied by a large tongue protrusion and backward movement of the ears. Sucrose Bobs and Quinine Gapes are discussed with respect to: (1) the relevance of facial reactions to both sucrose and quinine to taste receptors in horses, (2) the similarity of features of taste expression in horses to those documented in rodents and primates, and (3) the dissimilarity between facial reactions to taste and other social signals displayed by horses.
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Versace, E., Morgante, M., Pulina, G., & Vallortigara, G. (2007). Behavioural lateralization in sheep (Ovis aries). Behav. Brain. Res., 184(1), 72–80.
Abstract: This study investigates behavioural lateralization in sheep and lambs of different ages. A flock was tested in a task in which the animals were facing an obstacle and should avoid it on either the right or left side to rejoin flock-mates (adult sheep) or their mothers (lambs). A bias for avoiding the obstacle on the right side was observed, with lambs apparently being more lateralized than sheep. This right bias was tentatively associated with the left-hemifield laterality in familiar faces recognition which has been documented in this species. Differences between adult sheep and lambs were likely to be due to differences in social reinstatement motivation elicited by different stimuli (flock-mates or mothers) at different ages. Preferential use of the forelegs to step on a wood-board and direction of jaw movement during rumination was also tested in adult animals. No population bias nor individual-level lateralization was observed for use of the forelegs. At the same time, however, there was a large number of animals showing individual-level lateralization for the direction of jaw movement during rumination even though there was no population bias. These findings highlight that within the same species individual- and population-level lateralization can be observed in different tasks. Moreover, the results fit the general hypothesis that population-level asymmetries are more likely to occur in tasks that require social coordination among behaviourally asymmetric individuals.
Keywords: Lateralization; Laterality; Brain asymmetry; Hemisphere; Sheep; Lamb; Strength of lateralization
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Merkies, K., McKechnie, M. J., & Zakrajsek, E. (2018). Behavioural and physiological responses of therapy horses to mentally traumatized humans. Applied Animal Behaviour Science, .
Abstract: The benefits to humans of equine-assisted therapy (EAT) have been well-researched, however few studies have analyzed the effects on the horse. Understanding how differing mental states of humans affect the behaviour and response of the horse can assist in providing optimal outcomes for both horse and human. Four humans clinically diagnosed and under care of a psychotherapist for Post-Traumatic Stress Disorder (PTSD) were matched physically to four neurotypical control humans and individually subjected to each of 17 therapy horses loose in a round pen. A professional acting coach instructed the control humans in replicating the physical movements of their paired PTSD individual. Both horses and humans were equipped with a heart rate (HR) monitor recording HR every 5secs. Saliva samples were collected from each horse 30 min before and 30 min after each trial to analyze cortisol concentrations. Each trial consisted of 5 min of baseline observation of the horse alone in the round pen after which the human entered the round pen for 2 min, followed by an additional 5 min of the horse alone. Behavioural observations indicative of stress in the horse (gait, head height, ear orientation, body orientation, distance from the human, latency of approach to the human, vocalizations, and chewing) were retrospectively collected from video recordings of each trial and analyzed using a repeated measures GLIMMIX with Tukey's multiple comparisons for differences between treatments and time periods. Horses moved slower (p < 0.0001), carried their head lower (p < 0.0001), vocalized less (p < 0.0001), and chewed less (p < 0.0001) when any human was present with them in the round pen. Horse HR increased in the presence of the PTSD humans, even after the PTSD human left the pen (p < 0.0001). Since two of the PTSD/control human pairs were experienced with horses and two were not, a post-hoc analysis showed that horses approached quicker (p < 0.016) and stood closer (p < 0.0082) to humans who were experienced with horses. Horse HR was lower when with inexperienced humans (p < 0.0001) whereas inexperienced human HR was higher (p < 0.0001). Horse salivary cortisol did not differ between exposure to PTSD and control humans (p > 0.32). Overall, behavioural and physiological responses of horses to humans are more pronounced based on human experience with horses than whether the human is diagnosed with a mental disorder. This may be a reflection of a directness of movement associated with humans who are experienced with horses that makes the horse more attentive. It appears that horses respond more to physical cues from the human rather than emotional cues. This knowledge is important in tailoring therapy programs and justifying horse responses when interacting with a patient in a therapy setting.
Keywords: Equine-assisted therapy; Ptsd; Horse; Behaviour; Cortisol; Heart rate
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von Borstel, U. U., Duncan, I. J. H., Shoveller, A. K., Merkies, K., Keeling, L. J., & Millman, S. T. (2009). Impact of riding in a coercively obtained Rollkur posture on welfare and fear of performance horses. Appl. Anim. Behav. Sci., 116(2-4), 228–236.
Abstract: Rollkur, the usually coercively obtained hyperflexion of the horse's neck, is employed as a training method by some dressage riders; however, its use is controversial as it may cause discomfort and adversely affect the horse's welfare. The objectives of this study were to determine: (1) if horses showed differences in stress, discomfort and fear responses as measured by heart rate and behaviour when ridden in Rollkur (R) obtained by pressure on the reins compared to regular poll flexion (i.e. with the nose-line being at or just in front of the vertical; N), and (2) if they showed a preference between the two riding styles when given the choice. Fifteen riding horses were ridden 30 times through a Y-maze randomly alternating between sides. Riding through one arm of the Y-maze was always followed by a short round ridden in R, whereas riding through the other arm was followed by a short round ridden in N. Immediately after the conditioning phase, horses were again repeatedly ridden into the maze; however, riders left it to the horse to decide which arm of the maze to enter. During R, horses moved slower and showed more often behavioural signs of discomfort, such as tail-swishing, head-tossing or attempted bucks (P < 0.05), and 14 of the 15 horses chose significantly (P < 0.05) more often the maze-arm associated with N rather than R. Subsequently, eight of the horses were also subjected to two fear tests following a short ride in N as well as a ride in R. During R, horses tended to react stronger (P = 0.092) to the fear stimuli and to take longer (P = 0.087) to approach them. These findings indicate that a coercively obtained Rollkur position may be uncomfortable for horses and that it makes them more fearful and therefore potentially more dangerous to ride. Further studies need to assess horses' reaction to gradual training of Rollkur, as opposed to a coercively obtained hyperflexion, in order to decide whether the practice should be banned.
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Sato, S. (1984). Social licking pattern and its relationships to social dominance and live weight gain in weaned calves. Appl. Anim. Behav. Sci., 12(1), 25–32.
Abstract: Social licking patterns of heifer and steer herds were observed and recorded during periods of resting and intermittent feeding. The results revealed the following features: (1) heifers and steers had 15.0 and 15.2 social licking interactions per hour which lasted for 37.8 and 41.0 s on average, respectively. The average time an animal spent licking was about 25 s per hour; (2) all the animals in the herds were licked by others, but only 72.3% of the animals licked other animals; (3) the animals close in the social hierarchy tended to lick each other for a longer time than did remote animals; (4) the time receiving l licking and weight gain tended to be positively correlated. The observations suggest that (1) the motivation of giving licking may be individual-specific and may be influenced by genetic factors, while that of receiving licking appears to be general, and that (2) social licking may mean not only cleaning the skin and hair of a passive partner, but also leading it to psychological stability.
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Sato, S., Sako, S., & Maeda, A. (1991). Social licking patterns in cattle (<em>Bos taurus</em>): influence of environmental and social factors. Applied Animal Behaviour Science, 32(1), 3–12.
Abstract: To investigate the functions of social licking in cattle, four calves (one heifer and one steer in each of two herds), known to exhibit frequent social licking were observed continuously for 2 h before sunset for 13 days, using the focal animal sampling method. Calves were observed under various environmental conditions. Social licking significantly decreased on rainy days and tended to increase in a dirty barn and when food was restricted. Solicitation for social licking occurred not only from dominant animals of pairs but also from subordinates. Of the licking interactions, 31% occurred following solicitation, and these accounted for 39% of the total time spent licking. Following solicitation, 78% of social licking was oriented to the head and the neck regions that were inaccessible to self-licking animals. Unsolicited licking, however, was oriented not only to the head and the neck but also to the back and the rump regions, and these two latter regions were the major ones to receive licking. The effect of social relationships on social licking was investigated using least-squares analysis of variance. Social factors investigated were the difference of dominance values, the dominance-subordinance relationship, and kinship and familiarity; the sex of calves involved was also considered. Only familiarity had a significant effect on licking; exchanges of social licking increased with length of cohabitation. We suggest that social licking may have a cleaning effect, a tension-reducing effect and a bonding effect.
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Kusunose, R., & Yamanobe, A. (2002). The effect of training schedule on learned tasks in yearling horses. Appl. Anim. Behav. Sci., 78(2), 225–233.
Abstract: Twelve yearlings were divided into two groups and subjected to two different training schedules: (a) 30min of training daily (the daily trained group); and (b) 30min of training for 4 days, followed by a 3-day rest (the intermittently trained group), in order to compare the effect of two training methods on the ability of the horses to learn to be driven and ridden and to respond to the handlers? cues. The length of this experimental training was 17 days. The first step of training was surcingling and proceeded to lunging, to driving from the ground, and finally to being ridden at a trot on a track. Both groups were tested four times during the experimental period when they were at the same stage of training. They were driven and then ridden at a walk by a rider on a specified course and evaluated. The time to complete the course, accuracy of traveling the course, and heart rate during the test were used as the indicators of success in training. In three out of the four tests, the daily trained group tended to move faster and with more accuracy than the intermittently trained group. It would appear that daily training without a long interruption is more effective for yearlings.
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McVey, A., Wilkinson, A., & Mills, D. S. (2018). Social learning in horses: the effect of using a group leader demonstrator on the performance of familiar conspecifics in a detour task. Applied Animal Behaviour Science, .
Abstract: Learning through the observation of others allows the transfer of information without the costs incurred during individual trial and error learning. Horses (Equus caballus) are a highly social species, which might be expected to be capable of learning from others, but experimental findings are inconsistent, and potentially confounded by social facilitation effects not related directly to the learning of the task. We refined the methods used in previous equine social learning studies, to examine and distinguish specific social influences on learning of a task: we used predefined group leaders rather than agonistically dominant individuals to demonstrate a detour task to familiar conspecific observers; in addition we had two control groups: a non-observer (true control) and a group with the demonstrator simply present at the goal (social facilitation control). 44 socially kept horses were allocated to one of the three test conditions and took part in five trials each. Success rate, latency and detour direction were recorded. There was no significant difference between the three groups in the likelihood of them succeeding in the task nor latency to succeed; however there was a significant difference in the route chosen by the groups, with the true control choosing the side with the entrance gate significantly more than either the observer group or social facilitation group. Both of the latter two groups chose to go in the same direction relative to themselves, regardless of which side the gate was. Seven out of nine horses in the observer group chose the same direction as their demonstrator every time. Our results show a significant role of social facilitation on detour behaviour and highlight the importance of including adequate controls for simpler cognitive influences on behaviour before claims can be made about the specific learning of motor actions or goal directed behaviour. Social cues may be important to horses if the task is sufficiently challenging and motivationally important, so future work should consider more demanding, but ecologically relevant situations, in order to maximise the potential revelation of social learning effects which do not depend on simple local or stimulus enhancement effects.
Keywords: Equine; Imitation; Leader; Social facilitation; Social learning
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Crowell-Davis, S. L. (1985). Nursing behaviour and maternal aggression among Welsh ponies (Equus caballus). Appl Anim Behav Sci, 14(1), 11–25.
Abstract: Nursing behaviour and related aggression of mare-foal pairs was studied from birth (n = 21) to 24 weeks of age (n = 15) of the foal. Foals exhibited a decreasing length and frequency of nursing as they grew older. Mares rarely aggressed against their foals during nursing in the foal's first 4 weeks of life, but did so increasingly through Weeks 13-16, after which the rate of aggression during nursing decreased. Mares terminated nursing primarily by moving away, and were most likely to do so during the foal's first 4 weeks of life. They became gradually less likely to do so as the foal grew older. It was concluded that mares sometimes flex their hind limb on the side opposite the foal during nursing in order to conserve energy in a situation in which they would be remaining still anyway. There was no difference between colts and fillies in the frequency or duration of nursing or in the frequency with which their mothers aggressed against them or terminated nursing.
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Hagen, K., & Broom, D. M. (2004). Emotional reactions to learning in cattle. Appl. Anim. Behav. Sci., 85(3), 203–213.
Abstract: It has been suggested that during instrumental learning, animals are likely to react emotionally to the reinforcer. They may in addition react emotionally to their own achievements. These reactions are of interest with regard to the animals' capacity for self-awareness. Therefore, we devised a yoked control experiment involving the acquisition of an operant task. We aimed to identify the emotional reactions of young cattle to their own learning and to separate these from reactions to a food reward. Twelve Holstein-Friesian heifers aged 7-12 months were divided into two groups. Heifers in the experimental group were conditioned over a 14-day period to press a panel in order to open a gate for access to a food reward. For heifers in the control group, the gate opened after a delay equal to their matched partner's latency to open it. To allow for observation of the heifers' movements during locomotion after the gate had opened, there was a 15m distance in the form of a race from the gate to the food trough. The heart rate of the heifers, and their behaviour when moving along the race towards the food reward were measured. When experimental heifers made clear improvements in learning, they were more likely than on other occasions to have higher heart rates and tended to move more vigorously along the race in comparison with their controls. This experiment found some, albeit inconclusive, indication that cattle may react emotionally to their own learning improvement.
Keywords: Cattle; Expressive behaviour; Operant learning; Reinforcer
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