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.

Abstract: The present study aims to investigate whether the presence of unrelated adult horses at weaning would reduce the social stress of weaning and the emergence of undesirable behaviours. We tested this hypothesis in 32 domestic foals by comparing short and medium term behavioural and physiological responses to weaning in foals maintained in homogeneous groups of peers (PW) to those of foals grouped with both peers and unrelated adults (AW). In total, three trials were conducted, which each trial consisting of one AW group and one PW group. In all foals, weaning was followed by increased vocalization, increased locomotion and increased salivary cortisol concentration. However, signs of stress were less pronounced and shorter in duration in weanlings housed with unrelated adults (e.g. whinnies: p < 0.05; salivary cortisol: p < 0.05). Only foals without adults exhibited increased aggressiveness towards peers (p < 0.05) and abnormal behaviours (p < 0.05) such as excessive wood-chewing and redirected sucking towards peers. In conclusion, introducing adults to minimize weaning stress in foals and later on aggressiveness and abnormal behaviours appears as the most promising approach to date.

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.

Abstract: Brain lateralization is a phenomenon widely reported in the animal kingdom and sensory laterality has been shown to be an indicator of the appraisal of the stimulus valence by an individual. This can prove a useful tool to investigate how animals perceive intra- or hetero-specific signals. The human-animal relationship provides an interesting framework for testing the impact of the valence of interactions on emotional memories. In the present study, we tested whether horses could associate individual human voices with past positive or negative experiences. Both behavioural and electroencephalographic measures allowed examining laterality patterns in addition to the behavioural reactions. The results show that horses reacted to voices associated with past positive experiences with increased attention/arousal (gamma oscillations in the right hemisphere) and indicators of a positive emotional state (left hemisphere activation and ears held forward), and to those associated with past negative experiences with negative affective states (right hemisphere activation and ears held backwards). The responses were further influenced by the animals' management conditions (e.g. box or pasture). Overall, these results, associating brain and behaviour analysis, clearly demonstrate that horses' representation of human voices is modulated by the valence of prior horse-human interactions.

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.