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Schuetz, A., Farmer, K., & Krueger, K. (2017). Social learning across species: horses (Equus caballus) learn from humans by observation. Anim. Cogn., 20(3), 567–573.
Abstract: This study examines whether horses can learn by observing humans, given that they identify individual humans and orientate on the focus of human attention. We tested 24 horses aged between 3 and 12. Twelve horses were tested on whether they would learn to open a feeding apparatus by observing a familiar person. The other 12 were controls and received exactly the same experimental procedure, but without a demonstration of how to operate the apparatus. More horses from the group with demonstration (8/12) reached the learning criterion of opening the feeder twenty times consecutively than horses from the control group (2/12), and younger horses seemed to reach the criterion more quickly. Horses not reaching the learning criteria approached the human experimenters more often than those that did. The results demonstrate that horses learn socially across species, in this case from humans.
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Schneider, G., & Krueger, K. (2012). Third-party intervention. In Proceedings of the 2. International Equine Science Meeting (73). Wald: Xenophon Publishing.
Abstract: Third-party intervention is the interruption of a dyadic interaction by a third animal. We observed such interventions in affiliative interactions in free-ranging Esperia-ponies (Equus caballus). It is known that horses intervene in affiliative contexts especially when a preferred partner is involved, probably to protect their social bond to this preferred partner. To prove this hypothesis the present study investigated whether the preferred partner was targeted, i.e. challenged, or supported by the intervener or both randomly. Therefore we examined the social relationship between the intervener and both dyadic interacting individuals. We found that interveners usually supported individuals to which they have stronger social bonds than to other group mates, while they have no particular relationship to the targeted animals. This indicates that interveners in stable horse groups protect their social bonds to the supported animals by challenging their interaction partners. Of all observed horses only some mares showed intervention behaviour. Their social position, reflected by their position in the dominance hierarchy, social networks, and the spatial group structure were investigated. We found that interveners occupy no unique position, but they are involved in a high amount of affiliative interactions, high-ranking, and relatively aggressive. KW -
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Schneider, G., & Krueger, K. (2012). Third-party interventions keep social partners from exchanging affiliative interactions with others. Anim. Behav., 83(2), 377–387.
Abstract: Third-party interventions are defined as the interruption of dyadic interactions by third animals through direct physical contact, interposing or threats. Previous studies focused on the analysis of interventions against agonistic encounters. However, there have been no evaluations of interventions against affiliative behaviours, particularly in relation to the intervening animal�s social relationships and its social and spatial position. Horses, Equus caballus, are an interesting model species, as interventions against affiliative interactions occur more frequently than against agonistic interactions. In this study, 64 feral horses displayed 67 interventions in affiliative interactions and eight interventions in agonistic interactions within the observation period. We analysed the interventions in affiliative encounters, and found that it was mainly higher-ranking females that intervened in the affiliative interactions of group mates in the stable horse harems. The intervening animals took an active part in affiliative and agonistic encounters within the group, but did not occupy particular social roles or spatial positions. They intervened in affiliative interactions in which group mates with which they had social bonds interacted with other members of the group. They targeted the nonbonded animal and approached the one with which they were socially bonded. We suggest some species use third-party interventions in affiliative interactions to prevent competition for preferred social interaction partners from escalating into more costly agonistic encounters.
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Schanz, L., Krueger, K., & Hintze, S. (2019). Sex and Age Don't Matter, but Breed Type Does--Factors Influencing Eye Wrinkle Expression in Horses. Front. Vet. Sci., 6, 154.
Abstract: Identifying valid indicators to assess animals' emotional states is a critical objective of animal welfare science. In horses, eye wrinkles above the eyeball have been shown to be affected by pain and other emotional states. From other species we know that individual characteristics, e.g. age in humans, affect facial wrinkles, but it has not yet been investigated whether eye wrinkle expression in horses is systematically affected by such characteristics. Therefore, the aim of this study was to assess how age, sex, breed type, body condition and coat colour affect the expression and/or the assessment of eye wrinkles in horses. To this end, we adapted the eye wrinkle assessment scale from Hintze et al. (2016) and assessed eye wrinkle expression in pictures taken from the left and the right eye of 181 horses in a presumably neutral situation, using five outcome measures: a qualitative first impression reflecting how worried the horse is perceived by humans, the extent to which the brow is raised, the number of wrinkles, their markedness and the angle between a line through both corners of the eye and the topmost wrinkle. All measures could be assessed highly reliable with respect to intra- and inter-observer agreement. Breed type affected the width of the angle (F2, 114 = 8.20, p < 0.001), with thoroughbreds having the narrowest angle (M = 23.80, SD = 1.60), followed by warmbloods (M = 28.00, SD = 0.60), and coldbloods (M = 31.00, SD = 0.90). None of the other characteristics affected any of the outcome measures, and eye wrinkle expression did not differ between the left and the right eye area (all p-values > 0.05). In conclusion, horses' eye wrinkle expression and its assessment in neutral situations was not systematically affected by the investigated characteristics, except for 'breed type', which accounted for some variation in 'angle'; how much eye wrinkle expression is affected by emotion or perhaps mood needs further investigation and validation.
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Ruess, M., Schmelz, A., & Krueger, K. (2013). Einfluss vitomechanischer Schwingungen auf das Muskuloskeletalsystem der Pferde. In Göttinger Pferdetage’13 (111).
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Pick, D., & Krueger, K. (2012). Heuristics and complex decisions in man and horses. In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: Darwin’s claim of differences in mental abilities between man and other animals is a matter of degree rather than of kind and appears to apply nicely to a comparison of choice behaviour in humans and equine. Humans and horses make two kinds of decisions, fast reflexive decisions based on heuristics, and slower more considered decisions that require more complex cognitive processing. Heuristic use is adaptive in that decisions based on them are most often useful in helping an organism to survive while requiring little time and cognitive effort. There is considerable evidence that there is an innate basis to many heuristics in humans as well as horses. A case will be made that the investigation of heuristics specific to horses will lead to an understanding of equine behaviour that has not been possible using traditional learning theories alone. Traditional learning theories are restricted in explaining behaviour to appeals to reinforcement regimes or to the formation of associations between novel stimuli and stimuli that cause innate reflexes, but they are of no use when behaviour is controlled by stimuli that have never been reinforced or been systematically paired with other stimuli that cause reflexive responses. For example, a horse in a two-choice discrimination task may choose a familiar stimulus which has never been reinforced over a new stimulus added to the discrimination task, simply due to the use of an heuristic that usually works -- when in doubt, choose a more familiar object over an unfamiliar one. Conflict resolution strategies provide another good example of heuristic vs. slow decision making. In both humans and horses, conflict resolution strategies are used within, but not typically between social groups. Such groups can be defined as community of interest. Depending on the resource that has to be defended, groups may well be small identities, such as human families or horse harems, or large aggregations such as herds in horses, or religious groups or nations in humans. Fast and simple resolution is possible with stable social identities in simple environments, but more complicated and time consuming deliberative processes are required to resolve conflict over long-term resource acquisition, for decisions in complex conflicts situations and complex social settings.
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Marr, I., Preisler, V., Farmer, K., Stefanski, V., & Krueger, K. (2020). Non-invasive stress evaluation in domestic horses (Equus caballus): impact of housing conditions on sensory laterality and immunoglobulin A. Royal Society Open Science, 7(2), 191994.
Abstract: The study aimed to evaluate sensory laterality and concentration of faecal immunoglobulin A (IgA) as non-invasive measures of stress in horses by comparing them with the already established measures of motor laterality and faecal glucocorticoid metabolites (FGMs). Eleven three-year-old horses were exposed to known stressful situations (change of housing, initial training) to assess the two new parameters. Sensory laterality initially shifted significantly to the left and faecal FGMs were significantly increased on the change from group to individual housing and remained high through initial training. Motor laterality shifted significantly to the left after one week of individual stabling. Faecal IgA remained unchanged throughout the experiment. We therefore suggest that sensory laterality may be helpful in assessing acute stress in horses, especially on an individual level, as it proved to be an objective behavioural parameter that is easy to observe. Comparably, motor laterality may be helpful in assessing long-lasting stress. The results indicate that stress changes sensory laterality in horses, but further research is needed on a larger sample to evaluate elevated chronic stress, as it was not clear whether the horses of the present study experienced compromised welfare, which it has been proposed may affect faecal IgA.
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Marr, I., Farmer, K., & Krueger, K. (2018). Evidence for Right-Sided Horses Being More Optimistic than Left-Sided Horses. Animals, 8(12), 219.
Abstract: An individual's positive or negative perspective when judging an ambiguous stimulus (cognitive bias) can be helpful when assessing animal welfare. Emotionality, as expressed in approach or withdrawal behaviour, is linked to brain asymmetry. The predisposition to process information in the left or right brain hemisphere is displayed in motor laterality. The quality of the information being processed is indicated by the sensory laterality. Consequently, it would be quicker and more repeatable to use motor or sensory laterality to evaluate cognitive bias than to perform the conventional judgment bias test. Therefore, the relationship between cognitive bias and motor or sensory laterality was tested. The horses (n = 17) were trained in a discrimination task involving a box that was placed in either a “positive” or “negative” location. To test for cognitive bias, the box was then placed in the middle, between the trained positive and negative location, in an ambiguous location, and the latency to approach the box was evaluated. Results indicated that horses that were more likely to use the right forelimb when moving off from a standing position were more likely to approach the ambiguous box with a shorter latency (generalized linear mixed model, p < 0.01), and therefore displayed a positive cognitive bias (optimistic).
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Marr, I., Bauer, T., Farmer, K., & Krueger, K. (2016). Gibt die sensorische Lateralität im Objekttest Aufschluss über das Interieur, den aktuellen Gemütszustand, oder den Trainingszustand der Pferde? In 33. FFP-Jahrestagung.
Abstract: Vor dem obersten Ziel der klassischen Ausbildungsskala für Pferde, die Versammlung, steht das Geraderichten. Ein jedes Pferd ist jedoch von Geburt an asymmetrisch, also schief. Hinter dieser Schiefe verbirgt sich nicht nur die natürliche Schiefe (asymmetrische muskuläre Entwicklung der beiden Körperhälften), die von Geburt an zu beobachten ist, sondern auch die sensorische und motorische Lateralität, also dem ungleichmäßigen Gebrauch der rechten oder linken Sinnesorgane sowie Gliedmaßen, der sich mit der Reifung des Gehirns entwickelt. Alle drei müssen als eigenständige Faktoren, die sich gegenseitig beeinflussen, gesehen werden (Krüger 2014). Um den Weg des Geraderichtens zu erleichtern, sollte in der Ausbildung eines Pferdes nicht nur an der natürlichen Schiefe gearbeitet werden, sondern auch die sensorische und motorische Lateralität beachtet werden um den Prozess des Geraderichtens für das Pferd zu erleichtern. Die sensorische und motorische Lateralität resultiert aus der Aufgabenteilung/Spezialisierung beider Gehirnhälften (Hemisphären) (Rogers 2010). Die rechte Hemisphäre ist dabei für die Verarbeitung von Emotionen (z.B. Angst, Aggression, Freude, Zufriedenheit) sowie für lebenserhaltende Spontanreaktionen zuständig. Die linke Hemisphäre ist für die rationale Verarbeitung von Informationen essentiell (Adolphs et al. 1996, Rogers 2010, Austin und Rogers 2012, De Boyer Des Roches et al. 2008, Demaree et al. 2005, Austin und Rogers 2014). Rückschlüsse auf die Informationsverarbeitung lassen sich über die Beobachtung der verwendeten Sinnesorgane (Ohren und Augen) ziehen, die kontralateral mit den Großhirnhemisphären verbunden sind (Brooks et al. 1999). Es wird vermutet, dass Stress zu einer verstärkten Informationsverarbeitung durch die rechte Großhirnhemisphäre führt (Rogers 2010, Schultheiss et al. 2009). Diese konnte auch in ersten Untersuchungen am Pferd bestätigt werden (unveröffentlichte Daten). Forscher, die den einseitigen Gebrauch von rechten und linken Gliedmaßen (motorische Lateralität) bei Menschen und Tieren untersuchten, zeigten weiterhin Zusammenhänge zur Emotionalität und Reaktivität (McGreevy und Thomson 2006, Rogers 2009, Austin und Rogers 2012, Deesing und Grandin 2014). Die Tendenz zum einseitigen Gebrauch der Gliedmaßen gibt Hinweise auf den „Cognitive Bias“ (= individuelle, kognitive Verzerrung der Wahrnehmung und Verarbeitung von Informationen ins Positive oder Negative) und steht im Zusammenhang mit der persönlichen Neigung auf Stressfaktoren zu reagieren (zusammengefasst von Rogers 2010). Die sensorische Lateralität ändert sich jedoch schneller und situationsgebundener als die motorische Lateralität. Sie wird mittels Objekttests bestimmt, die ebenfalls verwendet werden können um die Reaktivität und Emotionalität zu untersuchen. Für eine objektivere Beurteilung des Interieurs eines Pferdes ist daher zu überlegen, ob die sensorische Lateralität als objektiver Parameter integriert werden kann, welchen Einflüssen diese unterliegt und mit welchen Persönlichkeitsmerkmalen sie korreliert. Wie in der Studie von Farmer et al. (2010) dargestellt werden konnte, zeigten bilateral trainierte Pferde eine weniger stark ausgeprägte Präferenz für die linken Sinnesorgane als traditionell trainierte Pferde in Tests mit Personen (ohne Interaktion). Es stellt sich daher die Frage, ob diese Beobachtung ein Resultat von langjährigem Training ist oder ob es sich bereits nach wenigen Wochen Training einstellt sowie ob solche Entwicklungen auch bei Objekten beobachtet werden können.
Für die erste Untersuchung ergaben sich daher in dieser Studie folgende Fragstellungen: Sind die Ergebnisse eines Objekttests mit Evaluierung der sensorischen Lateralität hinsichtlich der Lateralität wiederholbar? Denn, sollte es möglich sein mittels der sensorische Lateralität auf bestimmte
Persönlichkeitsmerkmale rückschließen zu können, so muss diese genauso stabil und reproduzierbar sein, wie die betreffenden Persönlichkeitsmerkmale. Unterscheiden sich Pferde hinsichtlich ihrer Lateralität in Objekttests, die bereits intensiv gleichmäßig beidseitig trainiert wurden, von Pferden, bei denen weniger Augenmerk auf gleichmäßiges beidseitiges Training gelegt wurde? Kann die Lateralität in Objekttests mit einer definierten gleichmäßigen beidseitigen Trainingsmethode beeinflusst werden? In welche Richtung verschiebt sich gegebenenfalls die Lateralität? Beeinflusst das Alter gegebenenfalls das Ausmaß von Veränderungen, da sich die sensorische Lateralität mit der Reifung des Gehirns entwickelt?
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Krueger, K., & Heinze, J. (2008). Horse sense: social status of horses (Equus caballus) affects their likelihood of copying other horses` behavior. Anim. Cogn., 11(3), 431–439.
Abstract: Animals that live in stable social groups need to gather information on their own relative position in the group`s social hierarchy, either by directly threatening or challenging others, or in a less costly manner, by observing interactions among others. Such indirect inference of dominance relationships has previously been reported from primates, rats, and birds and fish. Here, we show that domestic horses, Equus caballus, are similarly capable of social cognition. Taking advantage of a specific “following behavior” that horses show towards humans in a riding arena, we investigated whether bystander horses adjust their response to an experimenter according to the observed interaction and their own dominance relationship with the horse whose reaction to the experimenter they had observed before. Horses copied the following behavior towards an experimenter after watching a dominant horse following, but did not follow after observing a subordinate horse or a horse from another social group doing so. The following behavior which horses show towards an experimenter therefore appears to be affected by the demonstrator`s behavior and social status relative to the observer.
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