Abstract: Prima facie, the acquisition of novel behaviors in animals through observation of conspecifics seems straightforward. There are, however, various mechanisms through which the behavior of animals can be altered from observing others. These mechanisms range from simple hard-wired contagious processes to genuine learning by observation, which differ fundamentally in cognitive complexity. They range from social facilitation and local enhancement to true social learning. The different learning mechanisms are the subject of this review, largely because research on learning by observation can be confounded by difficulties in interpretation owing to the looming possibility of associative learning infecting experimental results. While it is often assumed that horses are capable of acquiring new behavior through intra-species observation, research on social learning in horses includes a variety of studies some of which may overestimate the possession of higher mental abilities. Assuming such abilities in their absence can have welfare implications, e.g. isolating stereotypical horses on the assumption that these behaviors can be learned though observation by neighboring horses. This review summarizes the definitions and criteria for the various types of social transmission and social learning and reviews the current documentation for each type in horses with the aim of clarifying whether horses possess the ability to learn through true social learning. As social ungulates, horses evolved in open landscapes, exposed to predators and grazing most of the day. Being in close proximity to conspecifics may theoretically offer an opportunity to learn socially, however anti-predator vigilance and locating forage may not require the neural complexity of social learning. Given the significant energetic expense of brain tissue, it is likely that social facilitation and local enhancement may have been sufficient in the adaptation of equids to their niche. As a consequence, social learning abilities may be maladaptive in horses. Collectively, the review proposes a novel differentiation between social transmission (social facilitation, local and stimulus enhancement) and social learning (goal emulation, imitation). Horses are undoubtedly sensitive to intra-species transfer of information but this transfer does not appear to satisfy the criteria for social learning, and thus there is no solid evidence for true social learning in horses.

Abstract: Domestic horses are faced with social challenges throughout their lives due to limitations in social contact, space restrictions and frequent changes in social companionship. This is in contrast to natural conditions where horses live in relatively stable harem bands. Currently, little is known about how repeated regrouping affect horse behaviour and welfare, and it is unknown whether horses may adapt to regrouping. In this study, we aimed to investigate the effects of an unstable group structure, caused by weekly regroupings, on behaviour and frequency of injuries in young horses. Forty-five horses were included in the study and were randomly assigned to the treatments; Stable (S; seven groups of three horses) or Unstable (U; eight groups of three horses). The experimental period lasted 7 weeks, during which horses in Stable groups remained in the same group, whereas one horse was exchanged between Unstable groups every week. The groups were kept in 80m×80m grass-covered enclosures and were fed additional roughage on the ground daily. Social interactions were recorded in Unstable groups immediately after each regrouping (30min), and in both Stable and Unstable groups on day 1, 3 and 6 after each regrouping (2×20min/group/day). Injuries were scored by the end of the experimental period. The level of aggression shown by horses in Unstable groups immediately after regrouping was not affected by week (F5,35=0.42, P=0.83), indicating that horses neither habituated, nor sensitized, to repeated regrouping. Compared to horses in Stable groups, more agonistic behaviour was shown by horses in Unstable groups (i.e. non-contact agonistic; F1,65=5.60, P=0.02), whereas there was no treatment effect on other variables. The level of play behaviour appeared, however, to be more variable in Unstable groups. There was a significant effect of week on the level of contact agonistic interactions as well as greeting behaviour, due to a high occurrence in weeks 4-6. Non-contact agonistic interactions constituted the major part of agonistic interactions (66%). Possibly as consequence, no serious injuries were registered and there was no treatment effect (U=184; P=0.11). We conclude that the behaviour of young horses is affected by group management, and that horses appear not to adapt to weekly regroupings.

Abstract: Summary Ten trotting Standardbred colts were recorded by high-speed cinematography at the ages of eight, 12 and 18 months. The horses were trotting on a treadmill operating at 4.0 m/secs. Five horses were subjected to a programme of intensified training from eight months of age, whereas the others were not trained and acted as controls. The films were analysed on a semi-automatic film-reading equipment and a number of variables used to demonstrate the gait symmetry were calculated and scaled by computer. Certain differences between left and right diagonal and contralateral pair of limbs, respectively, were noted, suggesting that laterality in horses may be inherited. The most pronounced systematic differences were found in 18-month old horses in the trained group. The results show the importance of careful gait examination and comprehensive coordination training at an early age.

Abstract: Persistence of individual differences in animal behavior in reactions to various environmental challenges could reflect basic divergences in temperament, which might be used to predict details of adaptive response. Although studies have been carried out on fear and anxiety in various species, including laboratory, domestic and wild animals, no consistent definition of fearfulness as a basic trait of temperament has emerged. After a classification of the events that may produce a state of fear, this article describes the great variability in behavior and in physiological patterns generally associated with emotional reactivity. The difficulties of proposing fearfulness-the general capacity to react to a variety of potentially threatening situations-as a valid basic internal variable are then discussed. Although there are many studies showing covariation among the psychobiological responses to different environmental challenges, other studies find no such correlations and raise doubts about the interpretation of fearfulness as a basic personality trait. After a critical assessment of methodologies used in fear and anxiety studies, it is suggested that discrepancies among results are mainly due to the modulation of emotional responses in animals, which depend on numerous genetic and epigenetic factors. It is difficult to compare results obtained by different methods from animals reared under various conditions and with different genetic origins. The concept of fearfulness as an inner trait is best supported by two kinds of investigations. First, an experimental approach combining ethology and experimental psychology produces undeniable indicators of emotional reactivity. Second, genetic lines selected for psychobiological traits prove useful in establishing between behavioral and neuroendocrine aspects of emotional reactivity. It is suggested that fearfulness could be considered a basic feature of the temperament of each individual, one that predisposes it to respond similarly to a variety of potentially alarming challenges, but is nevertheless continually modulated during development by the interaction of genetic traits of reactivity with environmental factors, particularly in the juvenile period. Such interaction may explain much of the interindividual variability observed in adaptive responses.

Abstract: Laterality in any organism or species can be manifest as morphological, sensory and functional degrees of asymmetry such as hemispheric dominance, handedness or sidedness and other motor functional behaviours and as such is equally important in equitation. The influence of the horses' sex on both the direction and the degree of the laterality was explored within and between 4 experimental procedures in the 1st study. The findings showed that the direction, but not the degree of idiosyncratic motor preference in the horses was strongly sex-related. Male horses exhibited significantly more left lateralized responses and female horses exhibited significantly more right lateralized responses. Visuo-spatial ability is also likely to be important in the performance horse. In many species, moderate to large differences in visuo-spatial ability have been reported between the sexes, with superior visuo-spatial ability being reported in males of all species investigated to date. As no known studies had addressed visuo-spatial ability in the equine, the objective of the 2nd study, was to determine if visuo-spatial ability differed between male and female horses. The results produced the first behavioural demonstration of superior visuo-spatial ability in male horses, similar to that reported in other species. There is evidence to suggest that visuospatial ability and motor laterality are associated with cerebral hemispheric asymmetry and may be intrinsically linked. Brain development and laterality have also been associated with hair patterning, and, in a 3rd study we attempted to identify predictors of lateral bias in motor behaviour in horses. We investigated the relationship between the direction of facial hair whorl rotation and the incidence/direction of laterality in the horse. The findings suggest that direction of facial hair whorl rotation may be a useful indicator of lateralised motor behavioural preferences in the horse. We then attempted to establish if laterality was evident at birth in a 4th study, where we explored if neonatal foals exhibited lateralised patterns during and immediately post the birthing process that were correlated with their facial hair whorl patterns. The results showed a significant association between the sex of the foal and the choice of foreleg presented initially during 2nd stage parturition. Significantly more colt foals led with the left foreleg and significantly more filly foals led with the right foreleg than expected purely by random and the behaviour was correlated with facial hair whorl patterns. The findings also suggest that lateralisation in the horse is determined in utero as has also been shown in humans. Comparisons of wholly intact male and female horses are warranted as they might elucidate additional linkages between motor behaviour, visuo-spatial ability and brain organisation and development in the horse. Further research in this area could lead to more appropriate competition conditions (better fence design/construction on cross-country tracks) and so eliminate unnecessary levels of risk associated with many equestrian sports.