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Krueger, K., Schwarz, S., Marr, I., & Farmer, K. (2022). Laterality in Horse Training: Psychological and Physical Balance and Coordination and Strength Rather Than Straightness. Animals, 12(8), 1042.
Abstract: For centuries, a goal of training in many equestrian disciplines has been to straighten the horse, which is considered a key element in achieving its responsiveness and suppleness. However, laterality is a naturally occurring phenomenon in horses and encompasses body asymmetry, motor laterality and sensory laterality. Furthermore, forcibly counterbalancing motor laterality has been considered a cause of psychological imbalance in humans. Perhaps asymmetry and laterality should rather be accepted, with a focus on training psychological and physical balance, coordination and equal strength on both sides instead of enforcing “straightness”. To explore this, we conducted a review of the literature on the function and causes of motor and sensory laterality in horses, especially in horses when trained on the ground or under a rider. The literature reveals that body asymmetry is innate but does not prevent the horse from performing at a high level under a rider. Motor laterality is equally distributed in feral horses, while in domestic horses, age, breed, training and carrying a rider may cause left leg preferences. Most horses initially observe novel persons and potentially threatening objects or situations with their left sensory organs. Pronounced preferences for the use of left sensory organs or limbs indicate that the horse is experiencing increased emotionality or stress, and long-term insufficiencies in welfare, housing or training may result in left shifts in motor and sensory laterality and pessimistic mentalities. Therefore, increasing laterality can be regarded as an indicator for insufficiencies in housing, handling and training. We propose that laterality be recognized as a welfare indicator and that straightening the horse should be achieved by conducting training focused on balance, coordination and equal strength on both sides.
Keywords: balance; body asymmetry; equitation; horse; motor laterality; sensory laterality; stress; welfare
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Deuel, N. R., & Lawrence, L. M. (1987). Laterality in the gallop gait of horses. J Biomech, 20(6), 645–649.
Abstract: Bilateral asymmetry in gallop stride limb contact patterns of four Quarter Horse fillies was documented by high-speed cinematography. Horses were filmed with rider by two cameras simultaneously while galloping along a straightaway. Even though signaled for each gallop lead an equivalent number of times, horses frequently switched leads, selecting the left lead nearly twice as often as the right. Velocities and stride lengths were greater for the left lead than the right, but stride frequencies did not differ between leads. Velocity effects were partitioned out in limb contact data analysis to enable the determination of persistent gallop stride asymmetries. The contact duration for the trailing (right) fore limb on the left lead exceeded the contact duration for the trailing (left) fore limb on the right lead. Selecting the right fore limb as the trailing fore limb may have allowed horses to use it to withstand the greater stresses and caused them to preferentially gallop with the left fore limb leading. Laterality may have an important influence on equine gallop motion patterns and thereby influence athletic performance.
Keywords: Animals; *Functional Laterality; *Gait; Horses/*physiology; Kinesis
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Krueger, K., & Farmer, K. (2011). Laterality in the Horse [Lateralität beim Pferd ]. mup, 4, 160–167.
Abstract: Horses are one-sided, not only on a motor level, but they also prefer to use one eye, ear or nostril over the other under particular circumstances. Horses usually prefer using the left eye to observe novel objects and humans. This preference is more marked in emotional situations and when confronted with unknown persons. Thus the horse’s visual laterality provides a good option for assessing its mental state during training or in human-horse interactions. A strong preference for the left eye may signal that a horse cannot deal with certain training situations or is emotionally affected by a particular person.
Pferde benutzen für die Begutachtung von Objekten und Menschen bevorzugt eine bestimmte Nüster, ein Ohr oder ein Auge. So betrachten die meisten Pferde Objekte und Menschen mit dem linken Auge. Die Lateralitätsforschung erklärt diese sensorische Lateralität mit der Verarbeitung von Informationen unterschiedlicher Qualität in verschiedenen Gehirnhälften und zeigt auf, dass positive und negative emotionale Informationen sowie soziale Sachverhalte mit dem linken Auge aufgenommen und vorwiegend an die rechte Gehirnhälfte weitergegeben werden. In diesem Zusammenhang ermöglicht die visuelle Lateralität, den Gemütszustand des Pferdes im Training und im therapeutischen Fördereinsatz zu erkennen und zu berücksichtigen. |
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Sakai, M., Hishii, T., Takeda, S., & Kohshima, S. (2006). Laterality of flipper rubbing behaviour in wild bottlenose dolphins (Tursiops aduncus): Caused by asymmetry of eye use? Behav. Brain. Res., 170(2), 204–210.
Abstract: To determine whether wild Indo-Pacific bottlenose dolphins (Tursiops aduncus) at Mikura Island, Japan, show asymmetry of eye or flipper use during a social behaviour, we investigated the laterality of flipper-to-body (F-B) rubbing, in which one dolphin (“rubber”) rubs the body of another (“rubbee”) with its flipper. We analysed 382 episodes of video-recorded F-B rubbings performed by identified individuals (N = 111 rubbers). F-B rubbing was conducted significantly more frequently with the left flipper than with the right flipper. The duration of F-B rubbings was also significantly longer with the left flipper than with the right flipper. Of 20 dolphins, nine individuals showed significant left-side bias as the rubber in this behaviour, whereas no dolphins showed significant right-side bias. The results indicate a population-level left-side bias of the rubber in F-B rubbing. An analysis of the swimming configurations during this behaviour suggests that the asymmetry in F-B rubbing was caused not only by the laterality of the rubber, but by a preference for use of the left eye in both dolphins during this behaviour. Dolphins used the left eye significantly more frequently than the right eye during the inquisitive behaviour, while they showed no significant bias in flipper use during the object-carrying behaviour. These facts also suggest that the asymmetry of F-B rubbing is caused by the preference for using the left eye. Significant left-side bias was observed only in F-B rubbings initiated by the rubbee, in which the rubbee determined its position during this behaviour. This suggests that this behavioural asymmetry was enhanced by the rubbees choosing the left side of the rubber to ensure better and longer rubs.
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Larose, C., Richard-Yris, M. - A., Hausberger, M., & Rogers, L. J. (2006). Laterality of horses associated with emotionality in novel situations. Laterality, 11(4), 355–367.
Abstract: We have established that lateral biases are characteristic of visual behaviour in 65 horses. Two breeds, Trotters and French Saddlebreds aged 2 to 3, were tested on a novel object test. The main finding was a significant correlation between emotionality index and the eye preferred to view the novel stimulus: the higher the emotionality, the more likely that the horse looked with its left eye. The less emotive French Saddlebreds, however, tended to glance at the object using the right eye, a tendency that was not found in the Trotters, although the emotive index was the same for both breeds. The youngest French Saddlebreds did not show this trend. These results are discussed in relation to the different training practices for the breeds and broader findings on lateralisation in different species.
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Chapelain, A., & Blois-Heulin, C. (2009). Lateralization for visual processes: eye preference in Campbell"s monkeys ( Cercopithecus c. campbelli ). Anim. Cogn., 12(1), 11–19.
Abstract: Abstract: Brain lateralization has been the matter of extensive research over the last centuries, but it remains an unsolved issue. While hand preferences have been extensively studied, very few studies have investigated laterality of eye use in non-human primates. We examined eye preference in 14 Campbell"s monkeys (Cercopithecus c. campbelli). We assessed eye preference to look at a seed placed inside a tube using monocular vision. Eye use was recorded for 100 independent and non-rewarded trials per individual. All of the 14 monkeys showed very strong preferences in the choice of the eye used to look inside the tube (mean preference: 97.6%). Eight subjects preferred the right eye and six subjects preferred the left eye. The results are discussed in light of previous data on eye preference in primates, and compared to data on hand preference from these subjects. Our findings would support the hypothesis for an early emergence of lateralization for perceptual processes compared to manual motor functions.
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Wells, D. L., & Millsopp, S. (2009). Lateralized behaviour in the domestic cat, Felis silvestris catus. Anim. Behav., 78(2), 537–541.
Abstract: Lateralized behaviour in the felids has been subject to little investigation. We examined the paw use of 42 domestic cats on three tasks designed to determine whether the animals performed asymmetrical motor behaviour. The influence of the cats' sex and age on their paw preferences was also explored. The distribution of the cats' paw preferences differed significantly between the three tasks. Task 1, the most complex exercise involving retrieval of a food treat from an empty jar, encouraged the most apparent display of lateralized behaviour, with all but one animal showing a strong preference to use either their left or right paw consistently. Tasks 2 (an exercise involving reaching for a toy suspended overhead) and 3 (a challenge involving reaching for a toy moving along the ground) encouraged ambilateral motor performance. Lateralized behaviour was strongly sex related. Male and female cats showed paw preferences at the level of the population, but in opposite directions. Females had a greater preference for using their right paw; males were more inclined to adopt their left paw. Feline age was unrelated to either strength or direction of preferred paw use. Overall, the findings suggest that there are two distinct populations of paw preference in the cat that cluster strongly around the animals' sex. The results also point to a relationship between lateralized behaviour and task complexity. More apparent patterns of lateralized behaviour were evident on more complex manipulatory tasks, hinting at functional brain specialization in this species.
Keywords: cat; Felis silvestris catus; handedness; laterality; paw preference
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Komárková, M., & Bartošová, J. (2012). Lateralized suckling in domestic horse foals (Equus caballus). In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: Animals with eyes placed laterally on their head would respond differently to stimuli detected on their different sides. Visual cues from the left eye are processed in the right hemisphere which controls the intensive emotion expression and fast response to the stimuli, while the left hemisphere receiving the information from the contralateral eye is responsible for the routine behaviour and concentration. Different use of left/right eye for exploring environment and evaluation of different stimuli was recently described in domestic horses. They use left eye predominantly for watching novel object or frightening stimulus which indicates specialization of right hemisphere (RH) for appraisal of these types of stimuli. We presumed such lateralisation also in foals during suckling when only monocular vision can be employed for observing surroundings (foals suck mostly in antiparallel body position). Two mutually exclusive hypotheses about the origin of probable foal’s preference for particular suckling side were postulated: 1) visual lateralization, then the foals should suck more often from the mother’s right side (‘from right’), so as potential danger would be detected by the better adapted right hemisphere (i.e. left eye); 2) motor lateralization (the analogous concept as human “handness”), then a foal should suck predominantly from one side, either left or right, more often. Within two seasons, suckling behaviour of 59 Kladruby mares giving birth to 79 foals was observed from deliveries to abrupt weaning (4-7 months of age). From 10 607 recorded suckling solicitations, 50.2% were performed from right, thus there was no preference of the suckling side on a population level. However, we found large variability in the probability of suckling from right among individual foals (p<0.0001); the probability ranged from 0.22-0.98. One third of the foals (35.4%) showed strong, either right (N=13) or left (N=15) side preference which further strengthened with age (interaction suckling side preference*foal’s age: p<0.0001). Other tested variables, i.e. sex of the foal, age of the mother and identity of the herd were not significant. The probability that the mother rejected foal’s suckling solicitation (N=108) did not differ according to the suckling side (right/left) or suckling side preference (right/left/no preference). Suckling bout duration was not significantly influenced neither by the suckling side nor suckling side preference regardless the mother or the foal terminated the suckling bout. In conclusion, only 16 % of the foals revealed significant right side preference for suckling while 19 % left side and 65 % no preference. Thus, we can reject the visual hypothesis because a general trait to suck with left eye open for better danger detection and recognition is unlikely to occur, at least in up to 7 months old foals. Such a young foal is probably fully focused on suckling and relies on mother’s vigilance. Observed individual side preferences indicated more likely a kind of motor lateralization in part of the foals. Further research is needed to clear up the effects of age and individual experience of the foal. Supported by AWIN, EU FP7 project No. 266213.
Keywords: laterality; suckling; horse; foal
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Corballis, M. C. (2008). Of mice and men – and lopsided birds. Cortex, 44(1), 3–7.
Abstract: The article by Zucca and Sovrano (2008, this issue) represents part of a new wave of studies of lateralization in nonhuman species. This work is often in conflict with earlier studies of human cerebral asymmetry and handedness, and the associated claim that these asymmetries are uniquely human, and perhaps even a result of the “speciation event” that led to modern humans. It is now apparent that there are close parallels between human and nonhuman asymmetries, suggesting that they have ancient roots. I argue that asymmetries must be seen in the context of a bilaterally symmetrical body plan, and that there is a balance to be struck between the adaptive advantages of symmetry and asymmetry. In human evolution, systematic asymmetries were incorporated into activities that probably are unique to our species, but the precursors of these asymmetries are increasingly evident in other species, including frogs, fish, birds, and mammals – especially primates.
Keywords: Cerebral asymmetry; Handedness; Evolution; Laterality
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Chappell, J., & Kacelnik, A. (2004). Selection of tool diameter by New Caledonian crows Corvus moneduloides. Anim. Cogn., 7(2), 121–127.
Abstract: One important element of complex and flexible tool use, particularly where tool manufacture is involved, is the ability to select or manufacture appropriate tools anticipating the needs of any given task-an ability that has been rarely tested in non-primates. We examine aspects of this ability in New Caledonian crows-a species known to be extraordinary tool users and manufacturers. In a 2002 study, Chappell and Kacelnik showed that these crows were able to select a tool of the appropriate length for a task among a set of different lengths, and in 2002, Weir, Chappell and Kacelnik showed that New Caledonian crows were able to shape unfamiliar materials to create a usable tool for a specific task. Here we examine their handling of tool diameter. In experiment 1, we show that when facing three loose sticks that were usable as tools, they preferred the thinnest one. When the three sticks were presented so that one was loose and the other two in a bundle, they only disassembled the bundle when their preferred tool was tied. In experiment 2, we show that they manufacture, and modify during use, a tool of a suitable diameter from a tree branch, according to the diameter of the hole through which the tool will have to be inserted. These results add to the developing picture of New Caledonian crows as sophisticated tool users and manufacturers, having an advanced level of folk physics.
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