Abstract: Classically, biologists have considered adaptation of behavioural characteristics in terms of long-term functional benefits to the individual, such as survival or reproductive fitness. In captive species, including the domestic horse, this level of explanation is limited, as for the most part, horses are housed in conditions that differ markedly from those in which they evolved. In addition, an individual horse's reproductive fitness is largely determined by man rather than its own behavioural strategies. Perhaps for reasons of this kind, explanations of behavioural adaptation to environmental challenges by domestic animals, including the capacity to learn new responses to these challenges, tend to concentrate on the proximate causes of behaviour. However, understanding the original function of these adaptive responses can help us explain why animals perform apparently novel or functionless activities in certain housing conditions and may help us to appreciate what the animal welfare implications might be. This paper reviews the behavioural adaptation of the domestic horse to captivity and discusses how apparently abnormal behaviour may not only provide a useful practical indicator of specific environmental deficiencies but may also serve the animal as an adaptive response to these deficiencies in an “abnormal” environment.

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.

Abstract: Innovative behaviors such as exploiting novel food sources can grant significant fitness benefits for animals, yet little is known about the mechanisms driving such phenomena, and the role of physiology is virtually unexplored in wild species. Two hypotheses predict opposing effects of physiological state on innovation success. On one hand, poor physiological condition may promote innovations by forcing individuals with poor competitive abilities to invent alternative solutions. On the other hand, superior physiological condition may ensure greater cognitive capacity and thereby better problem-solving and learning performance. To test these hypotheses, we studied the behavior of wild-caught house sparrows (Passer domesticus) in 4 novel tasks of food acquisition, one of which was presented to the birds in repeated trials, and we investigated the relationships of individual performance with relevant physiological traits. We found that problem-solving performance across the 4 tasks was moderately consistent within individuals. Birds with lower integrated levels of corticosterone, the main avian stress hormone, solved the most difficult task faster and were more efficient learners in the repeated task than birds with higher corticosterone levels. Birds with higher concentration of total glutathione, a key antioxidant, solved 2 relatively easy tasks faster, whereas birds with fewer coccidian parasites tended to solve the difficult task more quickly. Our results, thus, indicate that aspects of physiological state influence problem-solving performance in a context-dependent manner, and these effects on problem-solving capacity, probably including cognitive abilities, are more likely to drive individual innovation success than necessity due to poor condition.