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Leliveld, L.M.C.; Düpjan, S.; Tuchscherer, A.; Puppe, B. |
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Title |
Hemispheric Specialization for Processing the Communicative and Emotional Content of Vocal Communication in a Social Mammal, the Domestic Pig |
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Journal Article |
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2020 |
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Frontiers in Behavioral Neuroscience |
Abbreviated Journal |
Front. Behav. Neurosci. |
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14 |
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596758 |
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In humans, speech perception is lateralized, with the left hemisphere of the brain dominant in processing the communicative content and the right hemisphere dominant in processing the emotional content. However, still little is known about such a division of tasks in other species. We therefore investigated lateralized processing of communicative and emotionally relevant calls in a social mammal, the pig (Sus scrofa). Based on the contralateral connection between ears and hemispheres, we compared the behavioural and cardiac responses of 36 young male pigs during binaural and monaural (left or right) playback to the same sounds. The playback stimuli were calls of social isolation and physical restraint, whose communicative and emotional relevance, respectively, were validated prior to the test by acoustic analyses and during binaural playbacks. There were indications of lateralized processing mainly in the initial detection (left head-turn bias, indicating right hemispheric dominance) of the more emotionally relevant restraint calls. Conversely, there were indications of lateralized processing only in the appraisal (increased attention during playback to the right ear) of the more communicative relevant isolation calls. This implies differential involvement of the hemispheres in the auditory processing of vocalizations in pigs and thereby hints at similarities in the auditory processing of vocal communication in non-human animals and speech in humans. Therefore, these findings provide interesting new insight in the evolution of human language and auditory lateralization. |
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Equine Behaviour @ team @ |
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6699 |
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Krueger, K.; Gruentjens, T.; Hempel, E. |
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Title |
Wolf contact in horses at permanent pasture in Germany |
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2023 |
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Plos One |
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Plos One |
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18 |
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8 |
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e0289767 |
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Wolves returned to Germany in 2000, leading to fear in German horse owners that their horses could be in danger of wolf attacks or panic-like escapes from pastures when sighting wolves. However, reports from southern European countries indicate that wolf predation on horses diminishes with increasing presence of wildlife. Therefore, we conducted a long-term, filed observation between January 2015 and July 2022 on 13 non breeding riding horses, mares and geldings, kept permanently on two pastures within the range of wildlife and a stable wolf pack with annual offspring. Wildlife cameras at the fences of the pastures made 984 times recordings of wolves and 3151 times recordings of wildlife in and around the pastures. Between 1 January 2022 and 23 March 2022 we observed two stable horse groups. Pasture 1 was grazed by five horses of mixed breed, four mares and one gelding, with the median age of 8 years (min. = 6y, max. = 29y). Pasture 2 was grazed by eight heavy warmbloods and draught horses, three mares and five geldings, with the median age of 16 years (min. = 13y, max. = 22y). During this period no wolf was recorded at pasture 2, but wild boar several times, whereas at pasture 1, wolves were recorded 89 times, and for the wildlife mostly hare. Wolves may have avoided pasture 2 because of the presence of wild boar or because the large group of older, heavy breed horses may have formed a stable, protective group. The latter needs to be confirmed in a follow-up field observation, which records anti-predator behavior and welfare indicators in horses. In conclusion, wolves did not attack the mature horses on pastures with plenty of wildlife and the horses did not respond to the presence of wolves with visible signs of reduced welfare or panic. This indicates that wolves may prefer to prey on easily accessible wildlife around and at horse pastures and that Central European horses become accustom to the presence of non-hunting wolves. |
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Equine Behaviour @ team @ |
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6708 |
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Krueger, K.; Esch, L.; Byrne, R. |
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Title |
Need or opportunity? A study of innovations in equids |
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2021 |
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Plos One |
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Plos One |
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16 |
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9 |
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e0257730 |
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Debate persists over whether animals develop innovative solutions primarily in response to needs or conversely whether they innovate more when basic needs are covered and opportunity to develop novel behaviour is offered. We sourced 746 cases of “unusual” behaviour in equids by contacting equid owners and caretakers directly and via a website (https://innovative-behaviour.org), and by searching the internet platforms YouTube and Facebook for videos. The study investigated whether differences in need or opportunity for innovation were reflected in the numbers of different types of innovations and in the frequencies of repeating a once-innovative behaviour (i) with respect to the equids' sex, age, and breed type, (ii) across behavioural categories, and whether (iii) they were affected by the equids' management (single vs group housing, access to roughage feed, access to pasture, and social contact). We found that the numbers of different types of innovation and the frequency of displaying specific innovations were not affected by individual characteristics (sex, age, breed or equid species). Few types of innovation in escape and foraging contexts were observed, whilst the comfort, play, and social contexts elicited the greatest variety of innovations. We also found higher numbers of different types of innovations in horses kept in groups rather than in individual housing, and with unlimited rather than with restricted access to pasture and roughage. Equids in permanent social contact performed high rates of once-innovative behaviour. We suggest that equids produce goal-directed innovations and repeat the behaviour at high frequency in response to urgent needs for food and free movement or when kept in conditions with social conflict. However, equids devise the greatest variety of innovations when opportunity to play and to develop comfort behaviour arises and when kept in good conditions. |
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Equine Behaviour @ team @ |
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6653 |
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Marr, I.; Preisler, V.; Farmer, K.; Stefanski, V.; Krueger, K. |
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Non-invasive stress evaluation in domestic horses (Equus caballus): impact of housing conditions on sensory laterality and immunoglobulin A |
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2020 |
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Royal Society Open Science |
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Royal Society Open Science |
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7 |
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2 |
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191994 |
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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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Royal Society |
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doi: 10.1098/rsos.191994 |
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Equine Behaviour @ team @ |
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6608 |
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Goursot, C.; Düpjan, S.; Puppe, B.; Leliveld, L.M.C. |
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Title |
Affective styles and emotional lateralization: A promising framework for animal welfare research |
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Journal Article |
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2021 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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237 |
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105279 |
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Individuality; Motor lateralization; Hemispheric dominance; Motivational tendencies; Emotional reactivity; Emotional regulation |
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The growing recognition of animals as individuals has broader implications for farm animal welfare research. Even under highly standardized on-farm conditions, farm animals show heterogeneous but individually consistent behavioural patterns towards various stimuli, based on how they appraise these stimuli. As a result, animal welfare is likely to be highly individual as well, and studying the proximate mechanisms underlying distinct individual behaviour patterns and appraisal will improve animal welfare research. We propose to extend the framework of affective styles to bridge the gap between existing research fields on animal personality and affective states. Affective styles refer to consistent individual differences in emotional reactivity and regulation and can be predicted by baseline cerebral lateralization. Likewise, animals with consistent left or right motor biases--a proxy measure of individual patterns in cerebral lateralization--have been shown to differ in their personality, emotional reactivity, motivational tendencies or coping styles. In this paper, we present the current knowledge of the links between laterality and stable individual traits in behaviour and affect in light of hypotheses on emotional lateralization. Within our suggested framework, we make recommendations on how to investigate affective styles in non-human animals and give practical examples. This approach has the potential to promote a science of affective styles in nonhuman animals and significantly advance research on animal welfare. |
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0168-1591 |
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Equine Behaviour @ team @ |
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6698 |
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Stomp, M.; d'Ingeo, S.; Henry, S.; Cousillas, H.; Hausberger, M. |
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Title |
Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model |
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Journal Article |
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2021 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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236 |
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105271 |
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Laterality; Electroencephalography; Theta wave; Welfare; Horses |
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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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0168-1591 |
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Equine Behaviour @ team @ |
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6628 |
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Stomp, M.; d'Ingeo, S.; Henry, S.; Cousillas, H.; Hausberger, M. |
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Title |
Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model |
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Journal Article |
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Year |
2021 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
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Volume |
236 |
Issue |
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Pages |
105271 |
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Laterality; Electroencephalography; Theta wave; Welfare; Horses |
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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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Equine Behaviour @ team @ |
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6629 |
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Christensen, J.W.; Munk, R.; Hawson, L.; Palme, R.; Larsen, T.; Egenvall, A.; König von Borstel, U.U.; Rørvang, M.V. |
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Rider effects on horses' conflict behaviour, rein tension, physiological measures and rideability scores |
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Journal Article |
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2021 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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234 |
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105184 |
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Equitation science; Heart rate; Horse riding; Sport horse; Temperament; Training cues |
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Many breeding organisations include a subjective scoring of rideability by a professional rider into their evaluation of sports horses, but the consistency and reliability of the scoring system is debateable. The aim of this study was to investigate (i) whether professional riders agree in their scoring of rideability, and (ii) whether rideability scores are affected by rein tension, horse conflict behaviour, heart rate, and salivary cortisol, and (iii) whether riders induce different levels of conflict behaviour and physiological responses in the horses. Ten professional, female riders each rode 10 dressage horses (level M German scale; n = 100 combinations) through a standardised dressage test (10 min warm-up followed by a 4-min test) and subsequently scored the horses for rideability on the official 1-10 scale (1 = poor to 10 = excellent) from the Danish Riding Federation. Rein tension, horse heart rate, saliva cortisol and conflict behaviour were measured for each rider-horse pair. The riders were inconsistent in their scoring of rideability to the individual horses, e.g. scores for one of the horses ranged from 1 to 8. There was a significant effect of rider (P = 0.003) and the frequency of conflict behaviour (undesired head movements: P < 0.001, breaking the gait: P = 0.013, and other evasive behaviour: P = 0.032) on rideability scores, i.e. the more conflict behaviour the lower the score. There was no significant effect of rein tension and the physiological measures on rideability scores. However, there was a significant effect of rider on rein tension, horses' heart rate and increases in saliva cortisol concentrations and a tendency for some types of conflict behaviour, suggesting that some riders induced more discomfort in the horses. Future studies could help shed light on which elements of riding style are particularly important for sports horse welfare. In conclusion, this study found a large variation in rideability scores assigned to ten sports horses by ten professional riders. Rideability scores were dependent on the level of horse conflict behaviour, but not rein tension and physiological measures. Further studies are needed to improve the objectivity, consistency and reliability of rideability assessment of sports horses. |
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0168-1591 |
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Equine Behaviour @ team @ |
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6696 |
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Lesimple, C.; Reverchon-Billot, L.; Galloux, P.; Stomp, M.; Boichot, L.; Coste, C.; Henry, S.; Hausberger, M. |
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Title |
Free movement: A key for welfare improvement in sport horses? |
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Journal Article |
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2020 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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Volume |
225 |
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104972 |
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Horse; Housing; Free movement; Horse welfare; Stereotypic behaviour; Oxytocin |
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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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Equine Behaviour @ team @ |
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6619 |
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da Cruz, A.B.; Hirata, S.; dos Santos, M.E.; Mendonça, R.S. |
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Show me your best side: Lateralization of social and resting behaviors in feral horses |
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Journal Article |
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2023 |
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Behavioural Processes |
Abbreviated Journal |
Behav. Process. |
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206 |
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104839 |
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Cerebral lateralization; Drone technology; ; Hemispheric specialization; Horses; Social interactions |
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Growing evidence shows a variety of sensorial and motor asymmetries in social and non-social interactions in various species, indicating a lateralized processing of information by the brain. Using digital video cameras on tripods and drones, this study investigated lateralization in frequency and duration of social behavior patterns, in affiliative, agonistic, and resting contexts, in a feral population of horses (Equus ferus caballus) in Northern Portugal, consisting of 37 individuals organized in eight harem groups. Affiliative interactions (including grooming) were more often performed, and lasted longer, when recipients were positioned to the right side. In recumbent resting (animals lying down) episodes on the left side lasted longer. Our results of an affiliative behavior having a right side tendency, provide partial support to the valence-specific hypothesis of Ahern and Schwartz (1979) – left hemisphere dominance for positive affect, affiliative behaviors. Longer recumbent resting episodes on the left side may be due to synchronization. However, in both instances it is discussed how lateralization may be context dependent. Investigating the position asymmetries of social behaviors in feral equids will contribute to a better understanding of differential lateralization and hemispheric specialization from the ecological and evolutionary perspectives. |
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0376-6357 |
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Equine Behaviour @ team @ |
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6697 |
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