Abstract: Animals can indirectly gather meaningful information about other individuals by eavesdropping on their third-party interactions. In particular, eavesdropping can be used to indirectly attribute a negative or positive valence to an individual and to adjust one's future behavior towards that individual. Few studies have focused on this ability in nonhuman animals, especially in nonprimate species. Here, we investigated this ability for the first time in domestic horses (Equus caballus) by projecting videos of positive and negative interactions between an unknown human experimenter (a “positive” experimenter or a “negative” experimenter) and an actor horse. The horses reacted emotionally while watching the videos, expressing behavioral (facial expressions and contact-seeking behavior) and physiological (heart rate) cues of positive emotions while watching the positive video and of negative emotions while watching the negative video. This result shows that the horses perceived the content of the videos and suggests an emotional contagion between the actor horse and the subjects. After the videos were projected, the horses took a choice test, facing the positive and negative experimenters in real life. The horses successfully used the interactions seen in the videos to discriminate between the experimenters. They touched the negative experimenter significantly more, which seems counterintuitive but can be interpreted as an appeasement attempt, based on the existing literature. This result suggests that horses can indirectly attribute a valence to a human experimenter by eavesdropping on a previous third-party interaction with a conspecific.

Abstract: The effect of stabling for the first time on the behaviour and welfare of young and naïve horses has not yet been studied in detail. In this study we examined the effect of two typical housing systems on their subsequent behavioural and physiological responses upon first time stabling. Thirty-six 2-year-old Dutch warmbloods, 18 geldings and 18 mares were included in the study. Half of the horses were stabled in individual stables (10.5m2) and the other half in pair housing (48m2 for two horses). The study lasted 12 weeks. At the end of the study the physiological and temperamental responses of the horses on the different treatments was tested using a CRF challenge test (to test the HPA-axis function) and a Novel Object test (to test temperamental differences) respectively. Especially in the first week after stabling pair housed horses spent more time eating whereas individually housed horses spent more time either standing vigilant or sleeping. Stress-related behaviours like neighing, pawing, nibbling and snorting were all displayed significantly more frequently in the individually housed horses (P<0.01). At the end of the study 67% of the individually housed horses was seen performing one or more stereotypies (P<0.01). The cortisol response and ACTH response on the CRF challenge test were lower for horses in the individually housed boxes. It is suggested that this depression in socially isolated animals is caused by a desensitisation of the HPA axis in response to stress-induced elevations in ACTH and cortisol. In general there was no effect of the treatment on the reactivity of the horses during the Novel Object test. However, there were significant relations between the responses of horses in the Novel Object test and in the stable environment. It is concluded that sudden isolated stabling is stressful to young and naïve horses, resulting in a high prevalence of stereotypies and abnormal behaviours. This study also provided some support for the notion that social stress in horses may be associated with a blunted adrenocortical response to CRF challenge. The finding that responses of horses to a behavioural test are correlated with home environment behaviours suggests that individual horses exhibit consistent behavioural traits across different contexts, and opens the possibility of using behavioural tests in horses to predict more general underlying behavioural characteristics.

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.

Abstract: REASONS FOR PERFORMING STUDY: Little is known in quantitative terms about the influence of different head-neck positions (HNPs) on the loading pattern of the locomotor apparatus. Therefore it is difficult to predict whether a specific riding technique is beneficial for the horse or if it may increase the risk for injury. OBJECTIVE: To improve the understanding of forelimb-hindlimb balance and its underlying temporal changes in relation to different head and neck positions. METHODS: Vertical ground reaction force and time parameters of each limb were measured in 7 high level dressage horses while being ridden at walk and trot on an instrumented treadmill in 6 predetermined HNPs: HNP1 – free, unrestrained with loose reins; HNP2 – neck raised, bridge of the nose in front of the vertical; HNP3 – neck raised, bridge of the nose behind the vertical; HNP4 – neck lowered and flexed, bridge of the nose considerably behind the vertical; HNP5 – neck extremely elevated and bridge of the nose considerably in front of the vertical; HNP6 – neck and head extended forward and downward. Positions were judged by a qualified dressage judge. HNPs were assessed by comparing the data to a velocity-matched reference HNP (HNP2). Differences were tested using paired t test or Wilcoxon signed rank test (P<0.05). RESULTS: At the walk, stride duration and overreach distance increased in HNP1, but decreased in HNP3 and HNP5. Stride impulse was shifted to the forehand in HNP1 and HNP6, but shifted to the hindquarters in HNP5. At the trot, stride duration increased in HNP4 and HNP5. Overreach distance was shorter in HNP4. Stride impulse shifted to the hindquarters in HNP5. In HNP1 peak forces decreased in the forelimbs; in HNP5 peak forces increased in fore- and hindlimbs. CONCLUSIONS: HNP5 had the biggest impact on limb timing and load distribution and behaved inversely to HNP1 and HNP6. Shortening of forelimb stance duration in HNP5 increased peak forces although the percentage of stride impulse carried by the forelimbs decreased. POTENTIAL RELEVANCE: An extremely high HNP affects functionality much more than an extremely low neck.

Abstract: In recent years, Przewalski’s horses have been increasingly kept in semi-reserves. However,there areonly few studies ontheir behaviour and their ability to adaptto management interventions.In the main part of my dissertation, I focus on investigatingthe animals’ behaviour in different semi-reserves with varyinghabitats and living spaces. In addition, I investigate the horses’ behaviour during various management interventionsand analysetheensuing changes instress levels. Another aspect of my dissertation is the studyof social behaviour inPrzewalski’s horses. I investigate theparameters that should be used to demonstrate social bonds between individualsandassess whichdata provide the most meaningful results.In the commentary tochapter 1,several studies investigatingsocial bonds in horsesare discussed. Comparing the various studies, it is strikingthat no homogeneous analyses orevaluation criteria exist. While some authors only considersocial grooming, others include data onthe spatial proximity of the individuals in their evaluations, and various definitionsof proximity can also be found in the literature. Additionally, someauthors use friendly approaches between individuals asa furtherparameter wheninvestigating the social bonds.Continuing with this theme, in chapter 2I investigate the social behaviour of the horses and compare various analysis methods. I show that proactive behaviour, such as friendly approaches, is a good alternative to spatial proximity when investigating social bonds between group members, andis also useful for expanding the often very small data sets of mutual grooming in horses. Comparing Przewalski’s horses with wild living horses, I found no significant differences in the social behavior and the frequency of social interactions, regardlessof group size, group composition, habitat, and individual parameters such as age and gender.Inchapter 3,I investigate the behaviour of a Przewalski’s horse group when exploring a new area of their enclosure. Their behaviour changed, showing less resting and more feeding. Furthermore, the animals maintained greater distances from each other, and the alpha male, instead of herding the group from behind, led the group around the new area and walked in front of the other group members. Moreover, he showed a substantial increase in stress level during the first day.A general comparison of the behaviour of the Przewalski’s horses in different semi-reserves is provided in chapter 4. In it, the habitat choice of the animals and their reactions to various management interventions are investigated. It is shown that Przewalski’s horses prefer open grassland to dense woods, although keeping Przewalski’s horses in a pine forest does not influence the animals’ stress level. In contrast to habitat, food range, and changes in the group composition, which do not appear to change stress levels, individual factors, such as the hierarchy, influence the glucocorticoid level of the animals significantly. The largest increases in stress hormones were demonstrated when the horses were temporarily confined in smaller areas.The importance of the available space is also discussed in chapter 5, where it is shown that horses show less aggressive behaviour when more space is provided. In contrast, the husbandry system does not influence the animals’ aggression, but the way of feeding can additionally reduce agonistic behaviour, for example if food is offered ad libitum.In summary, the results of this study provide indications for the optimization of keeping Przewalski’s horses in semi-reserves. The animals can adapt themselves to the environment and thrive in habitats which do not correspond to their original steppe-like home. Nevertheless, the semi-reserves should provide sufficient grassland, as the horses prefer this type of habitat. General speaking, any types of habitat can only offer a suitable living space if the food range is sufficient for the number of horses. Otherwise, and especially during could winter months, supplementary feeding is necessary according to the body condition of the animals. This is particularly important for older, weakened, or very young animals, which are still adapting to life in the semi-reserve. Without sufficient food, stress hormones can increase and negatively influence the well-being of the horses. The same is true for management interventions: restricting the animals to small enclosures, for example, can adversely affect the horses’ well-being and should be only done if absolutely necessary. Targetedbehaviour observations allow the animals that have a special meaning for the group to be identified, and these should not be taken out of the group unless it is unavoidable, as young and unexperienced horses orientate themselves on those animals. This is especially true for the alpha male in a bachelor group, as these groups are often composed of young horses and the alpha-male provides the necessary stability and experience. Social bonds between individuals can be investigated by observing friendly and proactive behaviour, and social grooming and friendly approaches yield suitable data for such analysis.