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Ahrendt, L. P., Labouriau, R., Malmkvist, J., Nicol, C. J., & Christensen, J. W. (2015). Development of a standard test to assess negative reinforcement learning in horses. Appl. Anim. Behav. Sci., 169, 38–42.
Abstract: Most horses are trained by negative reinforcement. Currently, however, no standardised test for evaluating horses' negative reinforcement learning ability is available. The aim of this study was to develop an objective test to investigate negative reinforcement learning in horses. Twenty-four Icelandic horses (3 years old) were included in this study. The horses were tested in a pressure-release task on three separate days with 10, 7 and 5 trials on each side, respectively. Each trial consisted of pressure being applied on the hindquarter with an algometer. The force of the pressure was increased until the horse moved laterally away from the point of pressure. There was a significant decrease in required force over trials on the first test day (P<0.001), but not the second and third day. The intercepts on days 2 and 3 differed significantly from day 1 (P<0.001), but not each other. Significantly stronger force was required on the right side compared to the left (P<0.001), but there was no difference between first and second side tested (P=0.56). Individual performance was evaluated by median-force and the change in force over trials on the first test day. These two measures may explain different characteristics of negative reinforcement learning. In conclusion, this study presents a novel, standardised test for evaluating negative reinforcement learning ability in horses.
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Trösch, M., Pellon, S., Cuzol, F., Parias, C., Nowak, R., Calandreau, L., et al. (2020). Horses feel emotions when they watch positive and negative horse-human interactions in a video and transpose what they saw to real life. Anim. Cogn., 23(4), 643–653.
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.
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Baumgartner, M., Boisson, T., Erhard, M. H., & Zeitler-Feicht, M. H. (2020). Common Feeding Practices Pose A Risk to the Welfare of Horses When Kept on Non-Edible Bedding. Animals, 10, 441.
Abstract: During the evolution of the horse, an extended period of feed intake, spread over the entire 24-h period, determined the horses� behaviour and physiology. Horses will not interrupt their feed intake for more than 4 h, if they have a choice. The aim of the present study was to investigate in what way restrictive feeding practices (non ad libitum) affect the horses� natural feed intake behaviour. We observed the feed intake behaviour of 104 horses on edible (n = 30) and non-edible bedding (n = 74) on ten different farms. We assessed the duration of the forced nocturnal feed intake interruption of horses housed on shavings when no additional roughage was available. Furthermore, we comparatively examined the feed intake behaviour of horses housed on edible versus non-edible bedding. The daily restrictive feeding of roughage (2 times a day: n = 8; 3 times a day: n = 2), as it is common in individual housing systems, resulted in a nocturnal feed intake interruption of more than 4 hours for the majority (74.32%, 55/74) of the horses on shavings (8:50 ± 1:25 h, median: 8:45 h, minimum: 6:45 h, maximum: 13:23 h). In comparison to horses on straw, horses on shavings paused their feed intake less frequently and at a later latency. Furthermore, they spent less time on consuming the evening meal than horses on straw. Our results of the comparison of the feed-intake behaviour of horses on edible and non-edible bedding show that the horses� ethological feeding needs are not satisfied on non-edible bedding. If the horses accelerate their feed intake (also defined as �rebound effect�), this might indicate that the horses� welfare is compromised. We conclude that in addition to the body condition score, the longest duration of feed intake interruption (usually in the night) is an important welfare indicator of horses that have limited access to roughage.
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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.
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Baragli, P., Scopa, C., Maglieri, V., & Palagi, E. (2021). If horses had toes: demonstrating mirror self recognition at group level in Equus caballus. Anim. Cogn., .
Abstract: Mirror self-recognition (MSR), investigated in primates and recently in non-primate species, is considered a measure of self-awareness. Nowadays, the only reliable test for investigating MSR potential skills consists in the untrained response to a visual body mark detected using a reflective surface. Here, we report the first evidence of MSR at group level in horses, by facing the weaknesses of methodology present in a previous pilot study. Fourteen horses were used in a 4-phases mirror test (covered mirror, open mirror, invisible mark, visible colored mark). After engaging in a series of contingency behaviors (looking behind the mirror, peek-a-boo, head and tongue movements), our horses used the mirror surface to guide their movements towards their colored cheeks, thus showing that they can recognize themselves in a mirror. The analysis at the group level, which 'marks' a turning point in the analytical technique of MSR exploration in non-primate species, showed that horses spent a longer time in scratching their faces when marked with the visible mark compared to the non-visible mark. This finding indicates that horses did not see the non-visible mark and that they did not touch their own face guided by the tactile sensation, suggesting the presence of MSR in horses. Although a heated debate on the binary versus gradualist model in the MSR interpretation exists, recent empirical pieces of evidence, including ours, indicate that MSR is not an all-or-nothing phenomenon that appeared once in phylogeny and that a convergent evolution mechanism can be at the basis of its presence in phylogenetically distant taxa.
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Visser, E. K., Ellis, A. D., & Van Reenen, C. G. (2008). The effect of two different housing conditions on the welfare of young horses stabled for the first time. Applied Animal Behaviour Science, 114(3), 521–533.
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.
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Stomp, M., d'Ingeo, S., Henry, S., Cousillas, H., & Hausberger, M. (2021). Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model. Applied Animal Behaviour Science, 236, 105271.
Abstract: Assessing the animal welfare state is a challenge given the subjective individual cognitive and emotional processing involved. Electroencephalography (EEG) spectrum analysis has proved an ecologically valid recording situation to assess the link between brain processes and affective or cognitive states in humans: a higher slow wave/fast wave ratio has been associated with a positive internal state. In particular, a high production of theta power (3-8 Hz) has been related to positive emotions. On the other hand, it has been hypothesized that a left hemisphere (LH) dominance may be associated with a better welfare state. Here, we test the hypothesis that individual differences in the resting-state quantitative EEG power spectrum of adult horses (N = 18) and its lateralization pattern may reflect individual subjective perception of their conditions of life and welfare state. The results show clear individual differences in the proportions of the different waves and their inter-hemispheric distribution. Three different EEG power spectrum profiles were highlighted, from a bilateral predominance of theta waves in horses in a more positive welfare state to a bilateral predominance of beta waves in horses with clear expressions of compromised welfare. Interestingly, particular correlations were found between wave power activity and welfare parameters. We found a negative correlation between the number of stereotypic behaviours per hour and the median proportion of theta waves in the left hemisphere. and between the overall state (total chronic stress score) of welfare and gamma production in the right hemisphere (RH). These findings go along the hypothesis of a particular involvement of the left hemisphere for positive processing and of the right hemisphere for negative processing. However, the pattern of laterality did not appear as the most important feature here as both extreme clusters in terms of welfare showed bilateral predominance of one wave type. It is possible that hemispheric specialization makes more sense during acute emotion-inducing conditions rather than in this resting-state context (i.e. in absence of any high emotion-inducing stimulation), although the opposition gamma versus theta waves between both hemispheres in the horses with an intermediate welfare state is noticeable and intriguing. It seems that bilateral but also LH theta activity is a promising neurophysiological marker of good welfare in horses, while a bilateral or RH high production of gamma waves should alert about potential welfare alterations. Quantitative resting-state EEG power spectrum appears as a highly promising tool for exploring the brain processes involved in the subjective perception of chronic welfare, as a useful complementary tool for welfare assessment.
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Stomp, M., d'Ingeo, S., Henry, S., Cousillas, H., & Hausberger, M. (2021). Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model. Appl. Anim. Behav. Sci., 236, 105271.
Abstract: Assessing the animal welfare state is a challenge given the subjective individual cognitive and emotional processing involved. Electroencephalography (EEG) spectrum analysis has proved an ecologically valid recording situation to assess the link between brain processes and affective or cognitive states in humans: a higher slow wave/fast wave ratio has been associated with a positive internal state. In particular, a high production of theta power (3-8 Hz) has been related to positive emotions. On the other hand, it has been hypothesized that a left hemisphere (LH) dominance may be associated with a better welfare state. Here, we test the hypothesis that individual differences in the resting-state quantitative EEG power spectrum of adult horses (N = 18) and its lateralization pattern may reflect individual subjective perception of their conditions of life and welfare state. The results show clear individual differences in the proportions of the different waves and their inter-hemispheric distribution. Three different EEG power spectrum profiles were highlighted, from a bilateral predominance of theta waves in horses in a more positive welfare state to a bilateral predominance of beta waves in horses with clear expressions of compromised welfare. Interestingly, particular correlations were found between wave power activity and welfare parameters. We found a negative correlation between the number of stereotypic behaviours per hour and the median proportion of theta waves in the left hemisphere. and between the overall state (total chronic stress score) of welfare and gamma production in the right hemisphere (RH). These findings go along the hypothesis of a particular involvement of the left hemisphere for positive processing and of the right hemisphere for negative processing. However, the pattern of laterality did not appear as the most important feature here as both extreme clusters in terms of welfare showed bilateral predominance of one wave type. It is possible that hemispheric specialization makes more sense during acute emotion-inducing conditions rather than in this resting-state context (i.e. in absence of any high emotion-inducing stimulation), although the opposition gamma versus theta waves between both hemispheres in the horses with an intermediate welfare state is noticeable and intriguing. It seems that bilateral but also LH theta activity is a promising neurophysiological marker of good welfare in horses, while a bilateral or RH high production of gamma waves should alert about potential welfare alterations. Quantitative resting-state EEG power spectrum appears as a highly promising tool for exploring the brain processes involved in the subjective perception of chronic welfare, as a useful complementary tool for welfare assessment.
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McBride, S. D., Parker, M. O., Roberts, K., & Hemmings, A. (2017). Applied neurophysiology of the horse; implications for training, husbandry and welfare. Appl. Anim. Behav. Sci., 190, 90–101.
Abstract: Understanding the neural circuits underlying equine behaviour has the potential to help optimise strategies of husbandry and training. This review discusses two areas of neurophysiological research in a range of species and relates this information to the horse. The first discussion focuses on mechanisms of learning and motivation and assesses how this information can be applied to improve the training of the horse. The second concerns the identification of the equine neurophysiological phenotype, through behavioural and genetic probes, as a way of improving strategies for optimal equine husbandry and training success. The review finishes by identifying directions for future research with an emphasis on how neurophysiological systems (and thus behaviour) can be modified through strategic husbandry. This review highlights how a neurophysioloigical understanding of horse behaviour can play an important role in attaining the primary objectives of equitation science as well as improving the welfare of the horse.
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Lesimple, C., Reverchon-Billot, L., Galloux, P., Stomp, M., Boichot, L., Coste, C., et al. (2020). Free movement: A key for welfare improvement in sport horses? Appl. Anim. Behav. Sci., 225, 104972.
Abstract: Horses, and in particular sport horses, remain housed predominantly in single stalls. One of the main reported reasons is the fear that they will become agitated and injure themselves and thereby impair their performance if released in paddocks. The hour spent daily at work is also assumed to be sufficient to satisfy the horses' needs for locomotion. Growing scientific evidence shows that single stall housing has negative consequences on horses' welfare and that time for free movement is necessary. Our aim was to assess the feasibility of allowing sport horses used to staying permanently in their stall (except for 1 h riding/day) daily free time in a paddock and to evaluate its potential impact on their welfare using two behavioural reliable indicators (stereotypic behaviours and ear position), as well as selected blood parameters (blood cell count, oxytocin and serotonin concentrations). Two experiments were conducted on the same site. The first experiment evaluated sport horses' habituation to the novel situation of daily sessions in a paddock, and recorded welfare indicators in their stall before and during the experiment, comparing horses that were taken outdoors (experimental) and those that stayed in their stall (control). The second experiment evaluated the impact of this daily free time in a paddock on horses' welfare and its durability, focusing on positive indicators. Behavioural observations in paddocks showed that even horses that had never experienced free movement outside their stall habituated rapidly to this situation. The presence of hay in the paddock, may have speeded up habituation. Their restricted living conditions were associated with abnormalities in blood cell count that were not overcome during the time of daily paddock sessions but behavioural indicators showed that their welfare improved. In the second study, the experimental horses' welfare improved during the paddock release period, in particular their stereotypic behaviours decreased and oxytocin levels increased. No effects on serotonin concentrations could be evidenced. These effects were directly associated with being in paddock, as the indicators returned rapidly to their previous levels indicating compromised welfare when the paddock release sessions stopped. In conclusion, it can be recommended to release sport horses for free movement in paddocks as welfare is improved and subjective assessment by caretakers indicated minimal risks.
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