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Farmer, K., Krueger, K., & Byrne, R. (2010). Visual laterality in the domestic horse (Equus caballus) interacting with humans. Anim. Cogn., 13, 229–238.
Abstract: Most horses have a side on which they are easier to handle and a direction they favour when working on a circle, and recent studies have suggested a correlation between emotion and visual laterality when horses observe inanimate objects. As such lateralisation could provide important clues regarding the horse’s cognitive processes, we investigated whether horses also show laterality in association with people. We gave horses the choice of entering a chute to left or right, with and without the passive, non-interactive presence of a person unknown to them. The left eye was preferred for scanning under both conditions, but significantly more so when a person was present. Traditionally, riders handle horses only from the left, so we repeated the experiment with horses specifically trained on both sides. Again, there was a consistent preference for left eye scanning in the presence of a person, whether known to the horses or not. We also examined horses interacting with a person, using both traditionally and bilaterally trained horses. Both groups showed left eye preference for viewing the person, regardless of training and test procedure. For those horses tested under both passive and interactive conditions, the left eye was preferred significantly more during interaction. We suggest that most horses prefer to use their left eye for assessment and evaluation, and that there is an emotional aspect to the choice which may be positive or negative, depending on the circumstances. We believe these results have important practical implications and that emotional laterality should be taken into account in training methods.
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Schwarz, S., Marr, I., Farmer, K., Graf, K., Stefanski, V., & Krueger, K. (2022). Does Carrying a Rider Change Motor and Sensory Laterality in Horses? Animals, 12(8), 992.
Abstract: Laterality in horses has been studied in recent decades. Although most horses are kept for riding purposes, there has been almost no research on how laterality may be affected by carrying a rider. In this study, 23 horses were tested for lateral preferences, both with and without a rider, in three different experiments. The rider gave minimal aids and rode on a long rein to allow the horse free choice. Firstly, motor laterality was assessed by observing forelimb preference when stepping over a pole. Secondly, sensory laterality was assessed by observing perceptual side preferences when the horse was confronted with (a) an unfamiliar person or (b) a novel object. After applying a generalised linear model, this preliminary study found that a rider increased the strength of motor laterality (p = 0.01) but did not affect sensory laterality (p = 0.8). This suggests that carrying a rider who is as passive as possible does not have an adverse effect on a horse�s stress levels and mental state.
Keywords: laterality; horse; rider; sensory laterality; motor laterality; novel object; side preference
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Krueger, K., Schwarz, S., Marr, I., & Farmer, K. (2022). Laterality in Horse Training: Psychological and Physical Balance and Coordination and Strength Rather Than Straightness. Animals, 12(8), 1042.
Abstract: For centuries, a goal of training in many equestrian disciplines has been to straighten the horse, which is considered a key element in achieving its responsiveness and suppleness. However, laterality is a naturally occurring phenomenon in horses and encompasses body asymmetry, motor laterality and sensory laterality. Furthermore, forcibly counterbalancing motor laterality has been considered a cause of psychological imbalance in humans. Perhaps asymmetry and laterality should rather be accepted, with a focus on training psychological and physical balance, coordination and equal strength on both sides instead of enforcing “straightness”. To explore this, we conducted a review of the literature on the function and causes of motor and sensory laterality in horses, especially in horses when trained on the ground or under a rider. The literature reveals that body asymmetry is innate but does not prevent the horse from performing at a high level under a rider. Motor laterality is equally distributed in feral horses, while in domestic horses, age, breed, training and carrying a rider may cause left leg preferences. Most horses initially observe novel persons and potentially threatening objects or situations with their left sensory organs. Pronounced preferences for the use of left sensory organs or limbs indicate that the horse is experiencing increased emotionality or stress, and long-term insufficiencies in welfare, housing or training may result in left shifts in motor and sensory laterality and pessimistic mentalities. Therefore, increasing laterality can be regarded as an indicator for insufficiencies in housing, handling and training. We propose that laterality be recognized as a welfare indicator and that straightening the horse should be achieved by conducting training focused on balance, coordination and equal strength on both sides.
Keywords: balance; body asymmetry; equitation; horse; motor laterality; sensory laterality; stress; welfare
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Krueger, K., Trager, L., Farmer, K., & Byrne, R. (2022). Tool Use in Horses. Animals, 12(15), 1876.
Abstract: Tool use has not yet been confirmed in horses, mules or donkeys. As this subject is difficult to research with conventional methods, we used a crowdsourcing approach to gather data. We contacted equid owners and carers and asked them to report and video examples of �unusual� behaviour via a dedicated website. We also searched YouTube and Facebook for videos of equids showing tool use. From 635 reports, including 1014 behaviours, we found 20 cases of tool use, 13 of which were unambiguous in that it was clear that the behaviour was not trained, caused by reduced welfare, incidental or accidental. We then assessed (a) the effect of management conditions on tool use and (b) whether the animals used tools alone, or socially, involving other equids or humans. We found that management restrictions were associated with corresponding tool use in 12 of the 13 cases (p = 0.01), e.g., equids using sticks to scrape hay within reach when feed was restricted. Furthermore, 8 of the 13 cases involved other equids or humans, such as horses using brushes to groom others. The most frequent tool use was for foraging, with seven examples, tool use for social purposes was seen in four cases, and there was just one case of tool use for escape. There was just one case of tool use for comfort, and in this instance, there were no management restrictions. Equids therefore can develop tool use, especially when management conditions are restricted, but it is a rare occurrence.
Keywords: crowdsourcing; horse; innovation; mule; management; tool use
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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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Krueger, K., Esch, L., Farmer, K., & Marr, I. (2021). Basic Needs in Horses?--A Literature Review. Animals, 11(6), 1798.
Abstract: Every animal species has particular environmental requirements that are essential for its welfare, and when these so-called “basic needs” are not fulfilled, the animals suffer. The basic needs of horses have been claimed to be social contact, social companionship, free movement and access to roughage. To assess whether horses suffer when one or more of the four proposed basic needs are restricted, we examined several studies (n = 38) that reported behavioural and physiological reactions to these restrictions. We assigned the studies according to the four types of responses investigated: (a) Stress, (b) Active, (c) Passive, and (d) Abnormal Behaviour. Furthermore, the number of studies indicating that horses reacted to the restrictions were compared with the number of studies reporting no reaction. The limited number of studies available on single management restrictions did not allow conclusions to be drawn on the effect of each restriction separately, especially in the case of social companionship. However, when combinations of social contact, free movement and access to roughage were restricted, many of the horses had developed responses consistent with suffering. Passive Responses, indicating acute suffering, and Abnormal Behaviour, indicating suffering currently or at some time in the past, were especially clearly demonstrated. This provides further evidence of the usefulness of assessing behavioural parameters in combination with physiological measurements when evaluating horse welfare. This meta-analysis of the literature confirms that it is justified to claim that social contact, free movement and access to roughage are basic needs in horses.
Keywords: abnormal behaviour; active responses; horse; movement; passive responses; roughage; stress; social contact
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Esch, L., Wöhr, C., Erhard, M., & Krueger, K. (2019). Horses� (Equus Caballus) Laterality, Stress Hormones, and Task Related Behavior in Innovative Problem-Solving. Animals, 9(5), 265.
Abstract: Domesticated horses are constantly confronted with novel tasks. A recent study on anecdotal data indicates that some are innovative in dealing with such tasks. However, innovative behavior in horses has not previously been investigated under experimental conditions. In this study, we investigated whether 16 horses found an innovative solution when confronted with a novel feeder. Moreover, we investigated whether innovative behavior in horses may be affected by individual aspects such as: age, sex, size, motor and sensory laterality, fecal stress hormone concentrations (GCMs), and task-related behavior. Our study revealed evidence for 25% of the horses being capable of innovative problem solving for operating a novel feeder. Innovative horses of the present study were active, tenacious, and may be considered to have a higher inhibitory control, which was revealed by their task related behavior. Furthermore, they appeared to be emotional, reflected by high baseline GCM concentrations and a left sensory and motor laterality. These findings may contribute to the understanding of horses� cognitive capacities to deal with their environment and calls for enriched environments in sports and leisure horse management.
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Wolter, R., Stefanski, V., & Krueger, K. (2018). Parameters for the Analysis of Social Bonds in Horses. Animals, 8(11), 191.
Abstract: Social bond analysis is of major importance for the evaluation of social relationships in group housed horses. However, in equine behaviour literature, studies on social bond analysis are inconsistent. Mutual grooming (horses standing side by side and gently nipping, nuzzling, or rubbing each other), affiliative approaches (horses approaching each other and staying within one body length), and measurements of spatial proximity (horses standing with body contact or within two horse-lengths) are commonly used. In the present study, we assessed which of the three parameters is most suitable for social bond analysis in horses, and whether social bonds are affected by individual and group factors. We observed social behaviour and spatial proximity in 145 feral horses, five groups of Przewalski�s horses (N = 36), and six groups of feral horses (N = 109) for 15 h per group, on three days within one week. We found grooming, friendly approaches, and spatial proximity to be robust parameters, as their correlation was affected only by the animals� sex (GLMM: N = 145, SE = 0.001, t = �2.7, p = 0.008) and the group size (GLMM: N = 145, SE < 0.001, t = 4.255, p < 0.001), but not by the horse breed, the aggression ratio, the social rank, the group, the group composition, and the individuals themselves. Our results show a trend for a correspondence between all three parameters (GLMM: N = 145, SE = 0.004, t = 1.95, p = 0.053), a strong correspondence between mutual grooming and friendly approaches (GLMM: N = 145, SE = 0.021, t = 3.922, p < 0.001), and a weak correspondence between mutual grooming and spatial proximity (GLMM: N = 145, SE = 0.04, t = 1.15, p = 0.25). We therefore suggest either using a combination of the proactive behaviour counts mutual grooming and friendly approaches, or using measurements of close spatial proximity, for the analysis of social bonds in horses within a limited time frame.
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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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Krueger, K., Marr, I., Dobler, A., & Palme, R. (2019). Preservation of fecal glucocorticoid metabolites and immunoglobulin A through silica gel drying for field studies in horses. conphys, 7(1).
Abstract: Non-invasive methods enable stress evaluation through measuring fecal glucocorticoid metabolites (FGMs), and immunoglobulin A (IgA) in the feces avoiding stressful blood drawing or stressful restraining of animals in the field. However, FGMs and IgA are mostly analysed in freshly frozen samples, which is difficult when fresh samples cannot be frozen immediately or frozen samples cannot be stored or transported. Good results were also derived from air-dried fecal samples, which are hampered by unstable air humidity in the field. These difficulties may be overcome, when drying of samples could be induced with colorless silica gel (SiO2) granules in a secure set-up, such as an air tight tube. We determined the speed of drying 1.5 g of a fresh fecal sample from six horses on air and on silica gel. Furthermore, FGMs and IgA were analysed in differently stored subsamples from 12 horses: in frozen fecal samples, in air- or silica gel-dried samples stored for 1 day and for 7 days, and in wet fecal samples kept in a tube at room temperature for 7 days. FGM levels remained stable in feces dried on air or on silica gel for 7 days, whereas IgA quantities showed a significant loss. Under field conditions, when freezing or transporting the frozen samples is not possible and humidity hampers air drying, drying samples on silica gel in air tight tubes appears to be very helpful and reliable for analysing FGMs.
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