Abstract: Some behaviours remain a mystery and their determinants are still uncertain. Here we propose a critical interpretation of two puzzling behaviours, i.e. yawning and adult play, recorded in large samples of domestic horses kept in sub-optimal conditions (e.g. time-restricted feeding practices, social isolation). Yawning is involved in behavioural state changes, especially in quiet contexts of motor relaxation, but seems also to be triggered by stress and emotional contexts. Thus, we investigated specific correlates of yawning and stereotypic behaviours in two large samples of working horses (n1 = 87, n2 = 59). A clear co-occurrences of yawning and stereotypic behaviours appeared: stereotypic horses yawned more than the non-stereotypic horses (Fisher test, p < 0.02), yawning increased at the same time periods as stereotypic behaviours did (Wilcoxon tests, p < 0.05 to 0.001) and yawning frequency was even sometimes positively correlated with stereotypic behaviour frequencies (Spearman correlations, p < 0.005). Play, more typical of young stages in healthy individuals, occurs rarely at adult stages but then more often in captive/domestic animals (which can be challenging to welfare) than in animals living in natural conditions. Thus, we tested the hypothesis that adult play may reflect altered welfare states in horses, where, as in several species, play rarely occurs at adult stages in natural conditions. We observed the behaviour (in particular social play) of 29 adult riding school horses during occasional outings in a paddock and measured several stress indicators (health-related, behavioural and physiological) when these horses were in their home individual boxes. Results revealed that the number of horses and rates of adult play appeared very high compared to field report data and that most stress indicators measured differed between “players” and “non-players”, revealing that most “playful” animals were more prone to suffer from chronic stress contrarily to “non-playful” horses (Fisher, Chi-square, Mann-Whitney and Spearman correlation tests, p < 0.05 to 0.001). Altogether, these results show that yawning and adult play can both reflect altered welfare states in horses, highlighting that these puzzling behaviours should not be considered, as currently thought, as reliable indicators of good welfare states and / or expression of positive emotions.

Abstract: <sec> <title>Background</title> <p>Adults play an important role in regulating the social behaviour of young individuals. However, a few pioneer studies suggest that, more than the mere presence of adults, their proportions in social groups affect the social development of young. Here, we hypothesized that aggression rates and social cohesion were correlated to adult-young ratios. Our biological model was naturally-formed groups of Przewalski horses, <italic>Equus f. przewalskii</italic>, varying in composition.</p> </sec><sec> <title>Methodology/Principal Findings</title> <p>We investigated the social interactions and spatial relationships of 12 one- and two-year-old Przewalski horses belonging to five families with adult-young ratios (AYR) ranging from 0.67 to 1.33. We found striking variations of aggression rates and spatial relationships related to the adult-young ratio: the lower this ratio, the more the young were aggressive, the more young and adults segregated and the tighter the young bonded to other young.</p> </sec><sec> <title>Conclusion/Significance</title> <p>This is the first study demonstrating a correlation between adult-young ratios and aggression rates and social cohesion of young individuals in a naturalistic setting. The increase of aggression and the emergence of social segregation in groups with lower proportions of adults could reflect a related decrease of the influence of adults as regulators of the behaviour of young. This social regulation has both theoretical and practical implications for understanding the modalities of the influence of adults during ontogeny and for recommending optimal settings, as for instance, for schooling or animal group management.</p> </sec>

Abstract: BACKGROUND: As visual attention is an intrinsic part of social relationships, and because relationships are built on a succession of interactions, their establishment involves learning and attention. The emotional, rewarding or punishing, content can modulate selective attention. In horses, the use of positive/negative reinforcement during training determines short and long-term human-horse relationships. In a recent study in horses, where either food or withers' grooming were used as a reward, it appeared that only the food-rewarded horses learned the task and show better relationship with humans. In the present study, we hypothesized that this differential effect of grooming/food rewards on learning performances could be due to attentional processes. Monitoring, gazes and behaviors directed towards the trainer revealed that the use of a food reward (FR) as positive reinforcement increased horses' selective attention towards their trainer. Conversely, horses trained with grooming reward (GR) expressed more inattentive responses and did not show a decrease of “agitated” behavior. However, individual plotting of attention vs. rate of learning performances revealed a complex pattern. Thus, while all FR horses showed a “window” of attention related to faster learning performances, GR horses' pattern followed an almost normal curve where the extreme animals (i.e., highest and lowest attention) had the slowest learning performances. On the other hand, learning was influenced by attention: at the end of training, the more attentive horses had also better learning performances. This study, based on horses, contributes to the general debate on the place of attentional processes at the interface of emotion and cognition and opens new lines of thought about individual sensitivities (only individuals can tell what an appropriate reward is), attentional processes and learning.

Abstract: Behavioural modifications, including modifications of emotional reactivity, can occur following early experience such as handling (manual rubbing). Here, we investigated the effects of unilateral tactile stimulation at an early stage on emotional reactions later on. We handled newborn foals intensively on one side of their body. This early unilateral tactile experience had medium-term effects: the reactions of foals to a human approach, when they were 10 days old, differed according to the side stimulated at birth. Fewer right-handled foals accepted contact with humans, they delayed first contact longer and they evaded approaching humans sooner than did non-handled and left-handled foals. These results raise questions concerning the organization of neonatal care in animals and humans.

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.