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Schanz, L., Krueger, K., & Hintze, S. (2019). Sex and Age Don't Matter, but Breed Type Does--Factors Influencing Eye Wrinkle Expression in Horses. Front. Vet. Sci., 6, 154.
Abstract: Identifying valid indicators to assess animals' emotional states is a critical objective of animal welfare science. In horses, eye wrinkles above the eyeball have been shown to be affected by pain and other emotional states. From other species we know that individual characteristics, e.g. age in humans, affect facial wrinkles, but it has not yet been investigated whether eye wrinkle expression in horses is systematically affected by such characteristics. Therefore, the aim of this study was to assess how age, sex, breed type, body condition and coat colour affect the expression and/or the assessment of eye wrinkles in horses. To this end, we adapted the eye wrinkle assessment scale from Hintze et al. (2016) and assessed eye wrinkle expression in pictures taken from the left and the right eye of 181 horses in a presumably neutral situation, using five outcome measures: a qualitative first impression reflecting how worried the horse is perceived by humans, the extent to which the brow is raised, the number of wrinkles, their markedness and the angle between a line through both corners of the eye and the topmost wrinkle. All measures could be assessed highly reliable with respect to intra- and inter-observer agreement. Breed type affected the width of the angle (F2, 114 = 8.20, p < 0.001), with thoroughbreds having the narrowest angle (M = 23.80, SD = 1.60), followed by warmbloods (M = 28.00, SD = 0.60), and coldbloods (M = 31.00, SD = 0.90). None of the other characteristics affected any of the outcome measures, and eye wrinkle expression did not differ between the left and the right eye area (all p-values > 0.05). In conclusion, horses' eye wrinkle expression and its assessment in neutral situations was not systematically affected by the investigated characteristics, except for 'breed type', which accounted for some variation in 'angle'; how much eye wrinkle expression is affected by emotion or perhaps mood needs further investigation and validation.
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Rørvang, M. V., Christensen, J. W., Ladewig, J., & McLean, A. (2018). Social Learning in Horses--Fact or Fiction? Front. Vet. Sci., 5, 212.
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.
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Lesimple, C., Sankey, C., Richard, M. - A., & HAUSBERGER, M. (2012). Do Horses Expect Humans to Solve Their Problems? Front. Psychol., 3, 306.
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.
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McCoy, D. E., Schiestl, M., Neilands, P., Hassall, R., Gray, R. D., & Taylor, A. H. (2019). New Caledonian Crows Behave Optimistically after Using Tools. Current Biology, .
Abstract: Summary Are complex, species-specific behaviors in animals reinforced by material reward alone or do they also induce positive emotions? Many adaptive human behaviors are intrinsically motivated: they not only improve our material outcomes, but improve our affect as well [1, 2, 3, 4, 5, 6, 7, 8]. Work to date on animal optimism, as an indicator of positive affect, has generally focused on how animals react to change in their circumstances, such as when their environment is enriched [9, 10, 11, 12, 13, 14] or they are manipulated by humans [15, 16, 17, 18, 19, 20, 21, 22, 23], rather than whether complex actions improve emotional state. Here, we show that wild New Caledonian crows are optimistic after tool use, a complex, species-specific behavior. We further demonstrate that this finding cannot be explained by the crows needing to put more effort into gaining food. Our findings therefore raise the possibility that intrinsic motivation (enjoyment) may be a fundamental proximate cause in the evolution of tool use and other complex behaviors. Video Abstract
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Murphy, J., & Arkins, S. (2006). Laterality and visuo-spatial ability in the equine: Functional measures of sport horse selection? BSAP Occasional Publication, 35, 159–170.
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.
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Chaplin, S. J., & Gretgrix, L. (2010). Effect of housing conditions on activity and lying behaviour of horses. animal, 4(5), 792–795.
Abstract: Housing conditions for horses impose various levels of confinement, which may compromise welfare. Lying behaviour and activity can be used as welfare indicators for domestic animals and rebound behaviour suggests a build-up of motivation resulting from deprivation. The objective of this study was to determine if activity and lying behaviour of horses are affected by housing conditions and to investigate the occurrence of rebound behaviour after release from confinement. Eight horses were subjected, in pairs, to each of four experimental treatments; paddock (P), fully stabled (FS), partly stabled (PS) and yard (Y). Each horse received 6 days acclimatisation prior to the 24 h recording period. Time spent in lying and activity were electronically recorded using a tilt switch and motion sensor connected to a data logger worn on the horse's left foreleg. Time spent active during the first 5 min of release from stable to paddock in the PS treatment (days 1 and 5) and at the same time of day in the P treatment was used as a measure of rebound behaviour. Effect of housing conditions on total time spent active was highly significant (FS = 123 s, PS = 158 s, Y = 377 s, P = 779 s, P < 0.001). Housing conditions did not significantly affect total time spent lying (P = 0.646). Horses were significantly more active, compared with baseline paddock behaviour, on release from stabling on both days 1 (P = 0.006) and 5 (P = 0.025) of PS treatment. These results suggest that activity patterns of horses, but not lying behaviour, are affected by the housing conditions tested and that rebound activity occurs in horses after a period of confinement.
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Giraldeau, L. - A., Lefebvre, L., & Morand-Ferron, J. (2007). Can a restrictive definition lead to biases and tautologies? Behav. Brain Sci., 30(4), 411–412.
Abstract: We argue that the operational definition proposed by Ramsey et al. does not represent a significant improvement for students of innovation, because it is so restrictive that it might actually prevent the testing of hypotheses on the relationships between innovation, ecology, evolution, culture, and intelligence. To avoid tautological thinking, we need to use an operational definition that is taxonomically unbiased and neutral with respect to the hypotheses to be tested.
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Maloney, S. J. (2019). The Relationship Between Asymmetry and Athletic Performance: A Critical Review. The Journal of Strength & Conditioning Research, 33(9).
Abstract: Maloney, SJ. The relationship between asymmetry and athletic performance: A critical review. J Strength Cond Res 33(9): 2579-2593, 2019--Symmetry may be defined as the quality to demonstrate an exact correspondence of size, shape, and form when split along a given axis. Although it has been widely asserted that the bilateral asymmetries are detrimental to athletic performance, research does not wholly support such an association. Moreover, the research rarely seeks to distinguish between different types of bilateral asymmetry. Fluctuating asymmetries describe bilateral differences in anthropometric attributes, such as nostril width and ear size, and are thought to represent the developmental stability of an organism. There is evidence to suggest that fluctuating asymmetries may be related to impaired athletic performance, although contradictory findings have been reported. Sporting asymmetries is a term that may better describe bilateral differences in parameters, such as force output or jump height. These asymmetries are likely to be a function of limb dominance and magnified by long-standing participation within sport. Sporting asymmetries do not seem to carry a clear influence on athletic performance measures. Given the vast discrepancy in the methodologies used by different investigations, further research is warranted. Recent investigations have demonstrated that training interventions can reduce sporting asymmetries and improve performance. However, studies have not sought to determine whether the influence of sporting asymmetry is independent of improvements in neuromuscular parameters. It may be hypothesized that the deficient (weaker) limb has a greater potential for adaptation in comparison to the strong limb and may demonstrate greater responsiveness to training.
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Preiszner, B., Vincze, E., Seress, G., Papp, S., Bókony, V., Liker, A., et al. (2013). Necessity or capacity? Physiological state predicts problem-solving performance in house sparrows. Behav. Ecol., 25(1), 124–135.
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.
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Byström, A., Clayton, H. M., Hernlund, E., Rhodin, M., & Egenvall, A. (2020). Equestrian and biomechanical perspectives on laterality in the horse. Comp. Exerc. Physiol., 16(1), 35–45.
Abstract: It has been suggested that one of the underlying causes of asymmetrical performance and left/right bias in sound riding horses is laterality originating in the cerebral cortices described in many species. The aim of this paper is to review the published evidence for inherent biomechanical laterality in horses deemed to be clinically sound and relate these findings to descriptions of sidedness in equestrian texts. There are no established criteria to determine if a horse is left or right dominant but the preferred limb has been defined as the forelimb that is more frequently protracted during stance and when grazing. Findings on left-right differences in forelimb hoof shape and front hoof angles have been linked to asymmetric forelimb ground reaction forces. Asymmetries interpreted as motor laterality have been found among foals and unhandled youngsters, and the consistency or extent of asymmetries seems to increase with age. Expressions of laterality also vary with breed, sex, training and handling, stress, and body shape but there are no studies of the possible link between laterality and lameness. In a recent study of a group of seven dressage horses, a movement pattern in many ways similar to descriptions of sidedness in the equestrian literature, e.g. one hind limb being more protracted and placed more laterally than the other, has been documented. The role of innate laterality versus painful conditions, training, human handedness and simply habit remains to be determined. Understanding the biomechanical manifestations of laterality in healthy horses, including individual variation, would yield a potential basis for how laterality should be taken into account in relation to training/riding and rehabilitation of lameness.
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