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: This paper describes a method in which horses learn to communicate by touching different neutral visual symbols, in order to tell the handler whether they want to have a blanket on or not. Horses were trained for 10-15min per day, following a training program comprising ten steps in a strategic order. Reward based operant conditioning was used to teach horses to approach and touch a board, and to understand the meaning of three different symbols. Heat and cold challenges were performed to help learning and to check level of understanding. At certain stages, a learning criterion of correct responses for 8-14 successive trials had to be achieved before proceeding. After introducing the free choice situation, on average at training day 11, the horse could choose between a “no change” symbol and the symbol for either “blanket on” or “blanket off” depending on whether the horse already wore a blanket or not. A cut off point for performance or non-performance was set to day 14, and 23/23 horses successfully learned the task within this limit. Horses of warm-blood type needed fewer training days to reach criterion than cold-bloods (P<0.05). Horses were then tested under differing weather conditions. Results show that choices made, i.e. the symbol touched, was not random but dependent on weather. Horses chose to stay without a blanket in nice weather, and they chose to have a blanket on when the weather was wet, windy and cold (χ2=36.67, P<0.005). This indicates that horses both had an understanding of the consequence of their choice on own thermal comfort, and that they successfully had learned to communicate their preference by using the symbols. The method represents a novel tool for studying preferences in horses.

Abstract: The growing recognition of animals as individuals has broader implications for farm animal welfare research. Even under highly standardized on-farm conditions, farm animals show heterogeneous but individually consistent behavioural patterns towards various stimuli, based on how they appraise these stimuli. As a result, animal welfare is likely to be highly individual as well, and studying the proximate mechanisms underlying distinct individual behaviour patterns and appraisal will improve animal welfare research. We propose to extend the framework of affective styles to bridge the gap between existing research fields on animal personality and affective states. Affective styles refer to consistent individual differences in emotional reactivity and regulation and can be predicted by baseline cerebral lateralization. Likewise, animals with consistent left or right motor biases--a proxy measure of individual patterns in cerebral lateralization--have been shown to differ in their personality, emotional reactivity, motivational tendencies or coping styles. In this paper, we present the current knowledge of the links between laterality and stable individual traits in behaviour and affect in light of hypotheses on emotional lateralization. Within our suggested framework, we make recommendations on how to investigate affective styles in non-human animals and give practical examples. This approach has the potential to promote a science of affective styles in nonhuman animals and significantly advance research on animal welfare.

Abstract: This study investigates behavioural lateralization in sheep and lambs of different ages. A flock was tested in a task in which the animals were facing an obstacle and should avoid it on either the right or left side to rejoin flock-mates (adult sheep) or their mothers (lambs). A bias for avoiding the obstacle on the right side was observed, with lambs apparently being more lateralized than sheep. This right bias was tentatively associated with the left-hemifield laterality in familiar faces recognition which has been documented in this species. Differences between adult sheep and lambs were likely to be due to differences in social reinstatement motivation elicited by different stimuli (flock-mates or mothers) at different ages. Preferential use of the forelegs to step on a wood-board and direction of jaw movement during rumination was also tested in adult animals. No population bias nor individual-level lateralization was observed for use of the forelegs. At the same time, however, there was a large number of animals showing individual-level lateralization for the direction of jaw movement during rumination even though there was no population bias. These findings highlight that within the same species individual- and population-level lateralization can be observed in different tasks. Moreover, the results fit the general hypothesis that population-level asymmetries are more likely to occur in tasks that require social coordination among behaviourally asymmetric individuals.

Abstract: Despite the abundant empirical evidence on lateralized social behaviours, a clear understanding of the relative roles of two brain hemispheres in social processing is still lacking. This study investigated visual lateralization in social interactions of free-ranging European bison (Bison bonasus). The bison were more likely to display aggressive responses (such as fight and side hit), when they viewed the conspecific with the right visual field, implicating the left brain hemisphere. In contrast, the responses associated with positive social interactions (female-to-calf bonding, calf-to-female approach, suckling) or aggression inhibition (fight termination) occurred more likely when the left visual field was in use, indicating the right hemisphere advantage. The results do not support either assumptions of right-hemisphere dominance for control of various social functions or hypotheses about simple positive (approach) versus negative (withdrawal) distinction between the hemispheric roles. The discrepancy between the studies suggests that in animals, the relative roles of the hemispheres in social processing may be determined by a fine balance of emotions and motivations associated with the particular social reaction difficult to categorize for a human investigator. Our findings highlight the involvement of both brain hemispheres in the control of social behaviour.