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: The objective of this study was to investigate the ability of horses (Equus caballus) to detour around symmetric and asymmetric obstacles. Ten female Italian saddle horses were each used in three detour tasks. In the first task, the ability to detour around a symmetrical obstacle was evaluated; in the second and third tasks subjects were required to perform a detour around an asymmetrical obstacle with two different degrees of asymmetry. The direction chosen to move around the obstacle and time required to make the detour were recorded. The results suggest that horses have the spatial abilities required to perform detour tasks with both symmetric and asymmetric obstacles. The strategy used to perform the task varied between subjects. For five horses, lateralized behaviour was observed when detouring the obstacle; this was consistently in one direction (three on the left and two on the right). For these horses, no evidence of spatial learning or reasoning was found. The other five horses did not solve this task in a lateralized manner, and a trend towards decreasing lateralization was observed as asymmetry, and hence task difficulty, increased. These non-lateralized horses may have higher spatial reasoning abilities.

Abstract: In most cases, the social organisation of each of the seven species of Equidae existing today outside captivity is either territorial or non-territorial. The striking differences found between these two types of organisation in the social grouping and bonds, mating behaviour, leadership and dominance hierarchies of the animals are examined. It is thought that the non-territorial species show a less primitive type of organisation than the territorial animals. Infant Equidae are precocious animals and are able to follow their dams soon after birth. They stay close by their dams and travel with the herd from an early age and are therefore classified as “followers”, in contrast to the species which have a period of hiding after birth. Dams recognise their foals immediately after birth, whereas it takes 2 or 3 days for a foal to form an attachment to its dam. Being in close proximity to their dams, foals are able to nurse frequently and, unless artificially weaned, a foal will nurse until its dam foals again. Foals start to graze during their first week and as they grow older they spend more time grazing and less time nursing and resting. It is normal for foals to be corprophagic until one month old, and this provides them with bacteria essential for the digestion of fibre. Play behaviour is solitary in very young foals, but after 4 weeks of age, foals play together, with male foals playing more than females and showing more aggressive, fighting movements in play.

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.

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.