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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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Stachurska, A., Janczarek, I., Wilk, I., & Kedzierski, W. (2015). Does Music Influence Emotional State in Race Horses? Journal of Equine Veterinary Science, 35(8), 650–656.
Abstract: The aim of the study was to determine the effect of music featured in the barn, on the emotional state of race horses. Seventy 3-year-old Purebred Arabian horses in their first race season were divided into experimental group (EXP) of 40 horses and control group (CNT) of 30 horses and placed in separate barns. The EXP was subject to specifically composed music featured in the barn for 5 hours in the afternoon during the whole study. The emotional state in the horses was assessed at rest, saddling, and warm-up walk under rider. Measurements were taken six times, every 30 to 35 days, starting from the beginning of featuring the music. The horse's emotional state was assessed by cardiac activity variables. The music effect on the emotional state was also considered with regard to the horse's performance estimated by race records. The cardiac activity variables were compared with repeated measures design, whereas race records were analyzed with analysis of variance generalized linear model. The music positively affected the emotional state in race horses. The influence was noticeable already after the first month of featuring the music and increased in the second and third months. Despite the fact that later the variables began to return to initial levels, a positive effect of the music on prizes won by the horses in the EXP compared to the CNT was found (P < .05). The results suggest that the music may be featured in the barn, preferably for 2 to 3 months as a means of improving the welfare of race horses.
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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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Werhahn, H., Hessel, E. F., & Van den Weghe, H. F. A. (2012). Competition Horses Housed in Single Stalls (II): Effects of Free Exercise on the Behavior in the Stable, the Behavior during Training, and the Degree of Stress. Journal of Equine Veterinary Science, 32(1), 22–31.
Abstract: Although housing horses in single stalls limits their natural behavior to a great extent, this housing system is widespread in Germany, especially for competition horses. To improve the welfare of this system, free exercise on pastures or paddocks is deemed suitable, but it is also feared because of injuries and decreased willingness or motivation to perform. In the present study, three treatments were investigated with regard to their effect on the behavior of six competition horses in the stable, behavior during training, and on their degree of stress: daily training without free exercise (no turnout [NT]), solitary turnout for 2 hours after training, and 2-hour turnout in groups of two after training (group turnout). The horses' behavior in the stable was continuously analyzed through video recordings (2 pm to 6 am) on 3 days at the end of each treatment. The degree of stress was evaluated daily by heart rate variability at rest. The behavior during training was evaluated by a questionnaire answered by the riders, and the distance covered during training was measured by global positioning system. When NT was allowed, the horses showed less lying in the stable compared with the treatments with turnout. Heart rate variability measurements resulted in great individual differences, but generally, there was a higher degree of stress shown with the treatment NT according to the following parameters: standard deviation of inter-beat-intervals (SDNN), square root of the mean of the sum of the squares of differences between successive inter-beat-intervals (RMSSD), and ratio between low frequency and high frequency (LF/HF). The willingness to perform was evaluated as being slightly better in the treatments with turnout than in the treatment without turnout.
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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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Vitale, V., Balocchi, R., Varanini, M., Sgorbini, M., Macerata, A., Sighieri, C., et al. (2013). The effects of restriction of movement on the reliability of heart rate variability measurements in the horse (Equus caballus). J. Vet. Behav., 8(5), 400–403.
Abstract: Analysis of heart rate variability (HRV) is a noninvasive approach for investigating the sympathovagal balance of the autonomic nervous system. In recent years, HRV has been increasingly evaluated in animal research. In horses, it has been suggested that basal resting conditions can be achieved by restraining them. The aim of this study was to verify how restriction of movement influences HRV i2n horses. Ten healthy standardbred mares were used to measure the electrocardiographic signal under 2 conditions: free to move in the stall and restrained in the stock. Results indicate that the restriction of movement is associated with increased nervous system sympathetic activity not consistent with resting conditions.
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Morgan, K., Funkquist, P., & Nyman, G. (2002). The effect of coat clipping on thermoregulation during intense exercise in trotters. Equine Veterinary Journal, 34(S34), 564–567.
Abstract: Summary The aim of this study was to study the physiological, especially thermoregulatory, responses during intense exercise in the clipped horse compared to the horse with winter coat. Six Standardbred trotters were studied before and after clipping. They performed an inclined incremental high intensity treadmill exercise test and were monitored during recovery. The clipped horse differed significantly (ANOVA) during exercise as compare to coated: less increase in central venous blood temperature, higher skin surface temperature, greater difference skin to ambient temperature and higher rate of nonevaporative heat loss. The clipped horse had significantly lower total cutaneous evaporative heat loss from walk to end of peak exercise and a shorter time for recovery for the respiratory rate using a paired t test. The clipped horse showed a tendency (P = 0.059) to decreased oxygen uptake during the stepwise increase in workload. We concluded that the clipped horse experienced less strain on the thermoregulatory system due to an enhanced heat loss. Some clipped horses in the study showed a more efficient power output; future studies with emphasis on respiration and oxygen demand are needed to explain this.
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