Abstract: Laterality is now known to be an ubiquitous phenomenon among the vertebrates. Particularly, laterality of auditory processing has been demonstrated in a variety of species, especially songbirds and primates. Such a hemispheric specialization has been shown to depend on factors such as sound structure, species specificity and types of stimuli. Much less is known on the possible influence of social familiarity although a few studies suggest such an influence. Here we tested the influence of the degree of familiarity on the laterality of the auditory response in the domestic horse. This species is known for its social system and shows visible reactions to sounds, with one or two ears moving towards a sound source. By comparing such responses to the playback of different conspecific whinnies (group member, neighbor and stranger), we could demonstrate a clear left hemisphere (LH) preference for familiar neighbor calls while no preference was found for group member and stranger calls. Yet, we found an opposite pattern of ear side preference for neighbor versus stranger calls. These results are, to our knowledge, the first to demonstrate auditory laterality in an ungulate species. They open further lines of thought on the influence of the social “value” of calls and the listener's arousal on auditory processing and laterality.

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: Domestic animals are highly capable of detecting human cues, while wild relatives tend to perform less well (e.g. responding to pointing gestures). It is suggested that domestication may have led to the development of such cognitive skills. Here, we hypothesized that because domestic animals are so attentive and dependant to humans' actions for resources, the counter effect may be a decline of self sufficiency, such as individual task solving. Here we show a negative correlation between the performance in a learning task (opening a chest) and the interest shown by horses towards humans, despite high motivation expressed by investigative behaviours directed at the chest. If human-directed attention reflects the development of particular skills in domestic animals, this is to our knowledge the first study highlighting a link between human-directed behaviours and impaired individual solving task skills (ability to solve a task by themselves) in horses.

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.