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.

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.

Abstract: Despite attempts to generalise the left hemisphere-speech association of humans to animal communication, the debate remains open. More studies on primates are needed to explore the potential effects of sound specificity and familiarity. Familiar and non-familiar nonhuman primate contact calls, bird calls and non-biological sounds were broadcast to Japanese macaques. Macaques turned their heads preferentially towards the left (right hemisphere) when hearing conspecific or familiar primates supporting hemispheric specialisation. Our results support the role of experience in brain organisation and the importance of social factors to understand laterality evolution.

Abstract: 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.

Abstract: An animal that has stepped over an obstacle with its forelimbs uses a memory of the obstacle to guide the hind limbs so that they also clear the obstacle, even in situations in which long pauses are introduced between forelimb and hind limb stepping. To further clarify the features of hind limb obstacle clearance memory, the present study examined hind limb obstacle clearance in the horse. A rider guided horses over obstacles and paused the horse over obstacles in tests that examined the relationship between forelimb and hind limb stepping, with the following results. First, the horses displayed memory for an obstacle as measured by hind limb lifting over the obstacle for durations lasting as long as 15 min. The response was not dependent upon ongoing visualization of the obstacle, as limb lifting was unaffected by visual occlusion with blinders, a blindfold, or by removing the obstacle during the pause. Second, previous experience of stepping over an obstacle led to pause-related hind limb lifting at the object's previous location even on trials for which there was no obstacle and so no preceding forelimb lifting. Third, whereas a horse would lift its hind limbs to clear two successively presented obstacles, replacing an obstacle before the horse after the forelimbs had cleared the obstacle prevented subsequent hind limb lifting at the obstacle's previous location. Taken together the results show that hind limb obstacle clearance is guided by a place-object memory. The results are discussed in relation to the differential sensory and memonic control of forelimb and hind limb stepping with the suggestion that place-object memory can guide hind stepping as well as overshadow working memory from front leg stepping.