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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 |
Appl. Anim. Behav. Sci. |
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Volume |
236 |
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Pages |
105271 |
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Keywords |
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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Equine Behaviour @ team @ |
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6628 |
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Author |
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 |
Type |
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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Keywords |
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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0168-1591 |
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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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Title |
Rider effects on horses' conflict behaviour, rein tension, physiological measures and rideability scores |
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Journal Article |
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Year |
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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Equine Behaviour @ team @ |
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6696 |
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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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Year |
2021 |
Publication |
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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Lesimple, C.; Reverchon-Billot, L.; Galloux, P.; Stomp, M.; Boichot, L.; Coste, C.; Henry, S.; Hausberger, M. |
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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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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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0168-1591 |
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Equine Behaviour @ team @ |
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6619 |
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Amici, F.; Widdig, A.; Lehmann, J.; Majolo, B. |
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A meta-analysis of interindividual differences in innovation |
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2019 |
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Animal Behaviour |
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Anim. Behav. |
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155 |
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257-268 |
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age; bad competitor hypothesis; excess of energy hypothesis; innovation; interindividual differences; intraspecific variation; personality; rank; sex |
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The ability to innovate and the social transmission of innovations have played a central role in human evolution. However, innovation is also crucial for other animals, by allowing them to cope with novel socioecological challenges. Although innovation plays such a central role in animals' lives, we still do not know the conditions required for innovative behaviour to emerge. Here, we focused on interindividual differences in innovation by (1) extensively reviewing existing literature on innovative behaviour in animals and (2) quantitatively testing the different evolutionary hypotheses that have been proposed to explain interindividual variation in innovation propensity during foraging tasks. We ran a series of phylogenetically controlled mixed-effects meta-regression models to determine which hypotheses (if any) are supported by currently available empirical studies. Our analyses show that innovation is more common in individuals that are older and belong to the larger sex, but also in more neophilic and/or explorative individuals. Moreover, these effects change depending on the study setting (i.e. wild versus captive). Our results provide no clear support to the excess of energy or the bad competitor hypotheses and suggest that study setting and interindividual differences in traits related to personality are also important predictors of innovation. |
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0003-3472 |
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Equine Behaviour @ team @ |
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6589 |
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Merkies, K.; McKechnie, M.J.; Zakrajsek, E. |
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Behavioural and physiological responses of therapy horses to mentally traumatized humans |
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2018 |
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Applied Animal Behaviour Science |
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Equine-assisted therapy; Ptsd; Horse; Behaviour; Cortisol; Heart rate |
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The benefits to humans of equine-assisted therapy (EAT) have been well-researched, however few studies have analyzed the effects on the horse. Understanding how differing mental states of humans affect the behaviour and response of the horse can assist in providing optimal outcomes for both horse and human. Four humans clinically diagnosed and under care of a psychotherapist for Post-Traumatic Stress Disorder (PTSD) were matched physically to four neurotypical control humans and individually subjected to each of 17 therapy horses loose in a round pen. A professional acting coach instructed the control humans in replicating the physical movements of their paired PTSD individual. Both horses and humans were equipped with a heart rate (HR) monitor recording HR every 5secs. Saliva samples were collected from each horse 30 min before and 30 min after each trial to analyze cortisol concentrations. Each trial consisted of 5 min of baseline observation of the horse alone in the round pen after which the human entered the round pen for 2 min, followed by an additional 5 min of the horse alone. Behavioural observations indicative of stress in the horse (gait, head height, ear orientation, body orientation, distance from the human, latency of approach to the human, vocalizations, and chewing) were retrospectively collected from video recordings of each trial and analyzed using a repeated measures GLIMMIX with Tukey's multiple comparisons for differences between treatments and time periods. Horses moved slower (p < 0.0001), carried their head lower (p < 0.0001), vocalized less (p < 0.0001), and chewed less (p < 0.0001) when any human was present with them in the round pen. Horse HR increased in the presence of the PTSD humans, even after the PTSD human left the pen (p < 0.0001). Since two of the PTSD/control human pairs were experienced with horses and two were not, a post-hoc analysis showed that horses approached quicker (p < 0.016) and stood closer (p < 0.0082) to humans who were experienced with horses. Horse HR was lower when with inexperienced humans (p < 0.0001) whereas inexperienced human HR was higher (p < 0.0001). Horse salivary cortisol did not differ between exposure to PTSD and control humans (p > 0.32). Overall, behavioural and physiological responses of horses to humans are more pronounced based on human experience with horses than whether the human is diagnosed with a mental disorder. This may be a reflection of a directness of movement associated with humans who are experienced with horses that makes the horse more attentive. It appears that horses respond more to physical cues from the human rather than emotional cues. This knowledge is important in tailoring therapy programs and justifying horse responses when interacting with a patient in a therapy setting. |
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0168-1591 |
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Equine Behaviour @ team @ |
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6385 |
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McVey, A.; Wilkinson, A.; Mills, D.S. |
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Social learning in horses: the effect of using a group leader demonstrator on the performance of familiar conspecifics in a detour task |
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2018 |
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Applied Animal Behaviour Science |
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Equine; Imitation; Leader; Social facilitation; Social learning |
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Learning through the observation of others allows the transfer of information without the costs incurred during individual trial and error learning. Horses (Equus caballus) are a highly social species, which might be expected to be capable of learning from others, but experimental findings are inconsistent, and potentially confounded by social facilitation effects not related directly to the learning of the task. We refined the methods used in previous equine social learning studies, to examine and distinguish specific social influences on learning of a task: we used predefined group leaders rather than agonistically dominant individuals to demonstrate a detour task to familiar conspecific observers; in addition we had two control groups: a non-observer (true control) and a group with the demonstrator simply present at the goal (social facilitation control). 44 socially kept horses were allocated to one of the three test conditions and took part in five trials each. Success rate, latency and detour direction were recorded. There was no significant difference between the three groups in the likelihood of them succeeding in the task nor latency to succeed; however there was a significant difference in the route chosen by the groups, with the true control choosing the side with the entrance gate significantly more than either the observer group or social facilitation group. Both of the latter two groups chose to go in the same direction relative to themselves, regardless of which side the gate was. Seven out of nine horses in the observer group chose the same direction as their demonstrator every time. Our results show a significant role of social facilitation on detour behaviour and highlight the importance of including adequate controls for simpler cognitive influences on behaviour before claims can be made about the specific learning of motor actions or goal directed behaviour. Social cues may be important to horses if the task is sufficiently challenging and motivationally important, so future work should consider more demanding, but ecologically relevant situations, in order to maximise the potential revelation of social learning effects which do not depend on simple local or stimulus enhancement effects. |
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Equine Behaviour @ team @ |
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6395 |
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Rocha, A.D. de L.; Menescal-de-Oliveira, L.; da Silva, L.F.S. |
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Effects of human contact and intra-specific social learning on tonic immobility in guinea pigs, Cavia porcellus |
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Journal Article |
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2017 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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Cohabitation; Fear; Motor response; Defensive behaviour; Predator-prey |
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Abstract Social learning is the capacity of animals to acquire adaptive information from others. In the case of fear responses, animals can learn fearful or non-fearful responses by observing the behavior of conspecifics. Tonic immobility (TI) is an anti-predatory behavior elicited during intense fear situations. Studies have revealed that regular contact with humans can reduce TI responses in animals. In our study, we evaluated the effect of human contact on the TI responses in guinea pigs. We also evaluated the effect of cohabitation (non-fearful animals with fearful animals) on their TI responses. To achieve this, we measured the TI responses induced by postural inversion and restraint in guinea pigs as a result of different treatments. In our first experiment, we determined the effect of human contact on TI responses by establishing 3 treatment groups: no contact, handled, and habituated. In our second experiment, we addressed the effect of social learning on TI response by testing TI response in habituated, and unhabituated animals that had cohabitated for 10 days. In the first experiment, 10 days of either handling or habituation did not prevent TI in guinea pigs, but habituation did increase latency [F(2,119) = 14.19; p < 0.0001] and handling or habituation decrease duration [F(2,119) = 15.01; p < 0.0001] of the TI behavior in the guinea pigs. In the second experiment, the cohabitation of unhabituated and habituated animals reduced TI duration [F(2,93) = 5.058; p < 0.008]. These data suggest that both forms of human interaction can reduce experimenter fear in guinea pigs. It therefore seems that unhabituated guinea pigs learn not to fear the experimenter by cohabitating with habituated guinea pigs. |
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Equine Behaviour @ team @ |
Serial |
6133 |
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Author |
König v. Borstel, U.; Visser, E.K.; Hall, C. |
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Title |
Indicators of stress in equitation |
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Journal Article |
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2017 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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190 |
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43-56 |
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Keywords |
Stress; Horse; Riding; Heart rate variability; Cortisol; Behaviour |
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Abstract |
Abstract Stress is a generic concept describing the body's reaction to external stimuli, including both physiological and psychological factors. Therefore, by definition, the assessment of psychological stress in the exercising horse encompasses the problem of teasing apart the psychological and physiological factors both of which result in stress responses. The present study reviews the existing literature on various measures of stress taken specifically in the context of equitation science. Particular attention has been paid to short-term effects, and commonly used measurements of short-term stress include heart rate, a number of heart rate variability parameters, blood or saliva cortisol levels, eye temperature, and various behaviour parameters including in particular behaviour patterns presumably indicative of conflict with the rider's/trainer's aids. Inspection of the individual studies' results revealed that disagreement between these different measures of stress is commonplace. For physiological parameters, the largest proportion of agreement (i.e. both parameters simultaneously indicated either higher, insignificant or lower stress compared to a control treatment) was found for heart rate and heart rate variability parameters, while generally limited agreement was found for cortisol. It appears that cortisol levels may not be particularly useful for assessing/assessment of the valence of a situation in the exercising horse as cortisol levels are predominantly linked to activation and exercise levels. Although heart rate variability parameters reflect in theory more closely sympathovagal balance compared to cortisol levels, great care has to be taken regarding the use of appropriate time-frames, appropriate raw data correction methods as well as the use of appropriate equipment. In spite of its wide-spread and apparently successful use, popular equipment may in fact not be accurate enough under field conditions. Eye temperature is another promising parameter for assessment of psychological stress, but the technique is likewise susceptible to application errors. Given the high susceptibility of physiological parameters to errors at various experimental stages, behavioural rather than physiological parameters may in fact provide more accurate measures of valence when conducting experiments in the exercising horse. Behavioural parameters that appear to be particularly practical in assessing stress in ridden horses' behaviour are associated with frequencies of behaviour indicative of conflict. However, while increased frequencies of are a good indicator of stress, the absence of conflict behaviour does not provide proof of the absence of stress due to the possible occurrence of conditions such as Learned Helplessness. In future studies, the above issues should be taken into consideration when designing experiments to assess psychological stress in ridden horses. |
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0168-1591 |
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Equine Behaviour @ team @ |
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6160 |
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