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George, I., Cousillas, H., Richard, J. - P., & Hausberger, M. (2002). Song perception in the European starling: hemispheric specialisation and individual variations. Compt. Rend. Biol., 325(3), 197–204.
Abstract: Hemispheric specialisation for speech in humans has been well documented. The lateralisation for song production observed in songbirds is reminiscent of this hemispheric dominance. In order to investigate whether song perception is also lateralised, we made multiunit recordings of the neuronal activity in the field L of starlings during the presentation of species-specific and artificial non-specific sounds. We observed a systematic stronger activation in one hemisphere than in the other one during the playback of species-specific sounds, with inter-subject variability in the predominant hemisphere for song perception. Such an asymmetry was not observed for artificial non-specific sounds. Thus, our results suggest that, at least at the individual level, the two hemispheres of the starlings' brain perceive and process conspecific signals differently.
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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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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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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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Fureix, C., Bourjade, M., Henry, S., Sankey, C., & Hausberger, M. (2012). Exploring aggression regulation in managed groups of horses Equus caballus. Appl. Anim. Behav. Sci., 138(3–4), 216–228.
Abstract: Horses are highly social animals that have evolved to live in social groups. However, in modern husbandry systems, single housing prevails where horses experience social isolation, a challenge-to-welfare factor. One major reason for this single housing is the owners’ concerns that horses may injure each other during aggressive encounters. However, in natural conditions, serious injuries due to aggressive encounters are rare. What could therefore explain the claimed risks of group living for domestic horses? Basing our questioning on the current knowledge of the social life of horses in natural conditions, we review different practices that may lead to higher levels of aggression in horses and propose practical solutions. Observations of natural and feral horses mostly indicate a predominance of low frequencies and mild forms of aggression, based on subtle communication signals and ritualized displays and made possible by group stability (i.e. stable composition), dominance hierarchy and learning of appropriate social skills by young horses. Obviously, adults play a major role here in canalizing undesirable behaviours, and social experience during development, associated with a diversity of social partners, seems to be a prerequisite for the young horse to become socially skilled. Given the natural propensity of horses to have a regulation of aggression in groups, the tendency to display more aggression in groups of domestic horses under some management practices seems clearly related to the conditions offered. We therefore review the managing practices that could trigger aggressiveness in horses. Non social practices (space, resource availability) and social practices (group size, stability of membership, composition and opportunities for social experiences during development) in groups of domestic horses are discussed here. Finally, we propose simple practical solutions leading to more peaceful interactions in groups of domestic horses, based on the knowledge of horses’ natural social life which therefore should be enhanced (e.g. ensuring roughage availability, favouring group stability, introducing socially experienced adults in groups of young horses, etc.). The state of the art indicates that many questions still need to be answered. Given the importance of the associated welfare issues and the consequences on the use of horses, further research is required, which could benefit horses… and humans.
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Benhajali, H., Richard-Yris, M. - A., Ezzaouia, M., Charfi, F., & Hausberger, M. (2010). Reproductive status and stereotypies in breeding mares: A brief report. Appl. Anim. Behav. Sci., 128(1-4), 64–68.
Abstract: The aim of this study was to investigate the relationship between age, reproductive status and sire of the mare and the tendency to perform stereotypies. One hundred and fourteen purebred Arab mares, offspring of 39 sires ( offspring per sire) and aged 4-21 years () were observed from the 30th March to the 15th of May 2005 using instantaneous scan sampling. We used a multivariate logistic regression in order to study the relationship between age, reproductive status and sire of the mare and the tendency to perform stereotypies. 28% of the observed mares showed stereotypic behaviour, mostly weaving (22%). Neither age (χ2 Wald = 9.36, p = 0.89) nor sire (χ2 Wald = 4.34, p = 1.0) affected the occurrence of stereotypies whilst the reproductive status of the mare influenced significantly the occurrence (χ2 Wald = 10.75, p = 0.001) but also the type (χ2 = 12.1, p < 0.001) of stereotypic behaviour. Weaving was more frequent in non foaling mares (41.4%) than in foaling mares (1.8%) that performed mostly stall-walking.
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Rochais, C., Henry, S., Fureix, C., & Hausberger, M. (2016). Investigating attentional processes in depressive-like domestic horses (Equus caballus). Behavioural Processes, 124, 93–96.
Abstract: Abstract Some captive/domestic animals respond to confinement by becoming inactive and unresponsive to external stimuli. Human inactivity is one of the behavioural markers of clinical depression, a mental disorder diagnosed by the co-occurrence of symptoms including deficit in selective attention. Some riding horses display ‘withdrawn’ states of inactivity and low responsiveness to stimuli that resemble the reduced engagement with their environment of some depressed patients. We hypothesized that ‘withdrawn’ horses experience a depressive-like state and evaluated their level of attention by confronting them with auditory stimuli. Five novel auditory stimuli were broadcasted to 27 horses, including 12 ‘withdrawn’ horses, for 5 days. The horses’ reactions and durations of attention were recorded. Non-withdrawn horses reacted more and their attention lasted longer than that of withdrawn horses on the first day, but their durations of attention decreased over days, but those of withdrawn horses remained stable. These results suggest that the withdrawn horses’ selective attention is altered, adding to already evidenced common features between this horses’ state and human depression.
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Hausberger, M., Bruderer, C., Le Scolan, N., & Pierre, J. - S. (2004). Interplay between environmental and genetic factors in temperament/personality traits in horses (Equus caballus). J Comp Psychol, 118(4), 434–446.
Abstract: The aim of the present study was to broach the question of the relative influence of different genetic and environmental factors on different temperament/personality traits of horses (Equus caballus). The researchers submitted 702 horses to standardized experimental tests and investigated 9 factors, either genetic or environmental. Genetic factors, such as sire or breed, seemed to influence more neophobic reactions, whereas environmental factors, such as the type of work, seemed to play a more dominant role in reactions to social separation or learning abilities. Additive effects were evident, showing how environmental factors may modulate behavioral traits. This study constitutes a first step toward understanding the relative weights of genetic factors and how the environment may intervene in determining individual behavioral characteristics.
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Benhajali, H., Richard-Yris, M. - A., Leroux, M., Ezzaouia, M., Charfi, F., & Hausberger, M. (2008). A note on the time budget and social behaviour of densely housed horses: A case study in Arab breeding mares. Appl. Anim. Behav. Sci., 112(1-2), 196–200.
Abstract: We observed a high-density herd (200 mares/ha) of 44 Arab breeding mares, while in a bare paddock in Tunisia. Twenty-minute animal focal samples and scan sampling were used to determine the time budget of the mares during the period from 9 a.m. to 3 p.m. and study their social behaviour. The data obtained reveal restricted behavioural repertoires with missing behaviour like rolling, allogrooming and lying down; unusual time budgets with a high frequency of locomotion that constitutes the most frequent activity (27.9 ± 19.47%) of the mares. Social interactions were restricted to agonistic interactions but despite the high stocking density, aggressions were not that frequent among mares.
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Hausberger, M., Roche, H., Henry, S., & Visser, E. K. (2008). A review of the human-horse relationship. Appl. Anim. Behav. Sci., 109(1), 1–24.
Abstract: Despite a long history of human-horse relationship, horse-related incidents and accidents do occur amongst professional and non professional horse handlers. Recent studies show that their occurrence depend more on the frequency and amount of interactions with horses than on the level of competency, suggesting a strong need for specific research and training of individuals working with horses. In the present study, we review the current scientific knowledge on human-horse relationships. We distinguish here short occasional interactions with familiar or unfamiliar horses (e.g. veterinary inspection) and long-term bonds (e.g. horse-owner). An important aspect of the horse-human relationship is to try and improve the development and maintenance of a strong positive relationship. Studies show that deficits in the management conditions (housing, feeding, possibilities for social contact, and training methods) may lead to relational problems between horses and humans. Different methods have been used to assess and improve the human-horse relation, especially at the young age. They reveal that the time and type of contact all play a role, while recent studies suggest that the use of familiarized social models might be a great help through social facilitation. We argue that an important theoretical framework could be Hinde's [Hinde, R., 1979. Towards Understanding Relationships. Academic Press, Londres] definition of a relationship as an emerging bond from a series of interactions: partners have expectations on the next interaction on the basis of the previous ones. Understanding that a relationship is built up on the basis of a succession of interactions is an important step as it suggests that attention is being paid to the “positive” or “negative” valence of each interaction as a step for the next one. A better knowledge of learning rules is certainly necessary in this context not only to train the horse but also to counterbalance the unavoidable negative inputs that exist in routine procedures and reduce their impact on the relationship. It appears clearly that research is needed in order to assess how to better and safely approach the horse (e.g. research in position, posture, gaze, etc.), what type of approaches and timing may help in developing a positive bond, what influence human management and care have on the relationship, and how this can be adapted to have a positive influence on the relationship. Also the interaction between rider and horse, the search for the optimal match between two individuals, is an aspect of the horse-human relationship that requires attention in order to decrease the number of horse-riding accidents and reduced states of welfare. On the other hand, adequate knowledge is readily available that may improve the present situation rapidly. Developing awareness and attention to behavioural cues given by horses would certainly help decreasing accidents among professionals when interacting. Scientists therefore should play a major role in transmitting not only elements of the current knowledge of the ethology of the horse but also by helping developing observational skills.
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