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Author Farmer, K.; Krueger, K.; Byrne, R.
Title Visual laterality in the domestic horse (Equus caballus) interacting with humans Type Journal Article
Year (up) 2010 Publication Animal Cognition Abbreviated Journal Anim. Cogn.
Volume 13 Issue Pages 229-238
Keywords Horse – Laterality – Eye preference – Emotion – Vision
Abstract Most horses have a side on which they are easier to handle and a direction they favour when working on a circle, and recent studies have suggested a correlation between emotion and visual laterality when horses observe inanimate objects. As such lateralisation could provide important clues regarding the horse’s cognitive processes, we investigated whether horses also show laterality in association with people. We gave horses the choice of entering a chute to left or right, with and without the passive, non-interactive presence of a person unknown to them. The left eye was preferred for scanning under both conditions, but significantly more so when a person was present. Traditionally, riders handle horses only from the left, so we repeated the experiment with horses specifically trained on both sides. Again, there was a consistent preference for left eye scanning in the presence of a person, whether known to the horses or not. We also examined horses interacting with a person, using both traditionally and bilaterally trained horses. Both groups showed left eye preference for viewing the person, regardless of training and test procedure. For those horses tested under both passive and interactive conditions, the left eye was preferred significantly more during interaction. We suggest that most horses prefer to use their left eye for assessment and evaluation, and that there is an emotional aspect to the choice which may be positive or negative, depending on the circumstances. We believe these results have important practical implications and that emotional laterality should be taken into account in training methods.
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Call Number Equine Behaviour @ team @ Serial 4953
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Author Siniscalchi, M.; Sasso, R.; Pepe, A.M.; Vallortigara, G.; Quaranta, A.
Title Dogs turn left to emotional stimuli Type Journal Article
Year (up) 2010 Publication Behavioural Brain Research Abbreviated Journal Behav. Brain. Res.
Volume 208 Issue 2 Pages 516-521
Keywords Dog; Laterality; Vision; Behaviour; Physiology; Cognition; Emotion; Animal welfare
Abstract During feeding behaviour, dogs were suddenly presented with 2D stimuli depicting the silhouette of a dog, a cat or a snake simultaneously into the left and right visual hemifields. A bias to turn the head towards the left rather than the right side was observed with the cat and snake stimulus but not with the dog stimulus. Latencies to react following stimulus presentation were lower for left than for right head turning, whereas times needed to resume feeding behaviour were higher after left rather than after right head turning. When stimuli were presented only to the left or right visual hemifields, dogs proved to be more responsive to left side presentation, irrespective of the type of stimulus. However, cat and snake stimuli produced shorter latencies to react and longer times to resume feeding following left rather than right monocular visual hemifield presentation. Results demonstrate striking lateralization in dogs, with the right side of the brain more responsive to threatening and alarming stimuli. Possible implications for animal welfare are discussed.
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ISSN 0166-4328 ISBN Medium
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Notes Approved no
Call Number Equine Behaviour @ team @ Serial 5080
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Author Stennett, C.R.; Strauss, R.E.
Title Behavioural lateralization in zebrafish and four related species of minnows (Osteichthyes: Cyprinidae) Type Journal Article
Year (up) 2010 Publication Animal Behaviour. Abbreviated Journal Anim. Behav.
Volume 79 Issue 6 Pages 1339-1342
Keywords binary data; Cyprinidae; Danio rerio; fish; laterality; monocular test; motor bias; score test; zebrafish
Abstract Behavioural lateralization has been observed in many species of fishes during stimulus-specific tasks. However, one area that has been overlooked is the study of naïve side bias in motor behaviour of fishes in the absence of direct visual stimulus. To this end, we examined naïve side biases in motor behaviour in five species of minnows (Osteichthyes: Cyprinidae). Fifteen individuals of each species were subjected to a T-shaped test arena, with 40 randomized replicates per individual. We took advantage of rheotaxis by running a slow current of water through each arm of the test apparatus. Of the 75 individuals tested, 55 showed a rightward turning preference. The overall right-biased behaviour observed in these fishes in the absence of systematic stimulus strongly suggests that a stimulus-free control condition be included in the experimental design whenever plausible for studies of laterality in fishes and presumably in other organisms.
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Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 0003-3472 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 5358
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Author Tomkins, L.M.; McGreevy, P.D.; Branson, N.J.
Title Lack of standardization in reporting motor laterality in the domestic dog (Canis familiaris) Type Journal Article
Year (up) 2010 Publication Journal of Veterinary Behaviour Abbreviated Journal
Volume 5 Issue 5 Pages 235-239
Keywords dog; motor laterality; lateralization; paw preference; standardization
Abstract Over the past 2 decades, numerous studies have been undertaken to assess motor laterality in the domestic dog. In anticipation of growth in this area of enquiry, we decided to review the literature on canine motor biases to identify any shortcomings, reflect on the lessons to be learned from and offer ways forward for future research into canine laterality. The aim of this review is to (i) summarize motor laterality findings in the dog, (ii) highlight areas lacking in standardization, and (iii) propose necessary criteria for future tests and global reporting protocols. Our review of the literature highlighted the lack of standardization between studies in task selection, sample size, number of behavior scores recorded, and the methods by which motor laterality were classified and reported. This review illustrates the benefits of standardizing methods of motor laterality assessment so that comparisons can be made between the populations sampled. By adopting such an approach, researchers should mutually benefit as motor laterality data could then be compared and subjected to meta-analysis.
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ISSN 1558-7878 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ S1558-7878(10)00038-9 Serial 5378
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Author Krueger, K; Farmer, K.
Title Laterality in the Horse [Lateralität beim Pferd ] Type Journal Article
Year (up) 2011 Publication mensch & pferd international Abbreviated Journal mup
Volume 4 Issue Pages 160-167
Keywords Laterality, horse, information processing, training, welfare, human-animal interaction
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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Notes Approved no
Call Number Equine Behaviour @ team @ Serial 5444
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Author Sankey, C.; Henry, S.; Clouard, C.; Richard-Yris, M.-A.; Hausberger, M.
Title Asymmetry of behavioral responses to a human approach in young naive vs. trained horses Type Journal Article
Year (up) 2011 Publication Physiology & Behavior Abbreviated Journal
Volume 104 Issue 3 Pages 464-468
Keywords Laterality; Emotionality; Human approach; Horse
Abstract The aim of this study was to investigate the impact of training experience on young horses (Equus caballus)’ lateralized responses to an approaching human. The results show that the one year old untrained horses display asymmetrical responses to an approaching human, with more negative reactions (escapes, threats) when approached from the left side, while approaches towards the right shoulder elicited more positive behaviors. On the contrary, two years old trained horses reacted equally positively to approaches and contact on both sides. Our findings support those of previous studies investigating a link between emotionality and laterality and confirm the role of the left hemisphere in the processing of novel or negative stimuli. Moreover, the data underline the impact work and training can have on this laterality in horses.
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ISSN 0031-9384 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 5688
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Author Komárková, M.; Bartošová, J.
Title Lateralized suckling in domestic horse foals (Equus caballus) Type Conference Article
Year (up) 2012 Publication Proceedings of the 2. International Equine Science Meeting Abbreviated Journal Proc. 2. Int. Equine. Sci. Mtg
Volume in press Issue Pages
Keywords laterality; suckling; horse; foal
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.
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Corporate Author Komárková, M. Thesis
Publisher Xenophon Publishing Place of Publication Wald Editor Krueger, K.
Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 978-3-9808134-26 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 5505
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Author Lucidi, P.; Bacco, G.; Sticco, M.; Mazzoleni, G.; Benvenuti, M.; Bernabò, N.; Trentini, R.
Title Assessment of motor laterality in foals and young horses (Equus caballus) through an analysis of derailment at trot Type Journal Article
Year (up) 2013 Publication Physiology & Behavior Abbreviated Journal Physiol. Behav.
Volume 109 Issue Pages 8-13
Keywords Horse; Laterality; Longeing; Sidedness
Abstract The conflicting results regarding the study of motor laterality in horses may indicate that there does not exist a proper method to assess the degree and the direction of motor bias in these animals. Unfortunately, even less is known about the development of laterality in horses, and to what extent early manipulations can still exert their effects in adulthood. We propose a new method that can be easily applied at a very early age thus avoiding testing adult horses eventually biased by human handling and/or training. Forty-six horses (29 nine-month-old foals and 17 two-year old horses) were handled since birth bilaterally and housed in groups in wide areas. At the time of the analysis, in order to minimize environmental and sensorial disturbances, each horse was tested in a round pen individually or as dyad mother-foal. The ability/inability to properly execute a circle at trot was then recorded, assuming the direction of derailment, i.e. the cutting of the circle, as an indicator of motor bias. From the results of the study it is arguable that motor laterality in horses is acquired over time: in fact foals tested while their mothers were being subjected to longeing showed a higher percentage of ambidextrous animals, while two-year-old horses appeared biased toward the right (p<0.05). Results are discussed in the light of the scientific knowledge about equine biomechanics, taking into account horses' locomotion that leads to the advancement of the body mass through the activation of a kinetic chain that originates from the hindquarters.
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Publisher Place of Publication Editor
Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 0031-9384 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 6666
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Author Giljov, A.; Karenina, K.
Title Differential roles of the right and left brain hemispheres in the social interactions of a free-ranging ungulate Type Journal Article
Year (up) 2019 Publication Behavioural Processes Abbreviated Journal Behav. Process.
Volume 168 Issue Pages 103959
Keywords Laterality; Hemispheric specialization; Brain asymmetry; Eye preference; Ungulate; Bovid
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.
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Corporate Author Thesis
Publisher Place of Publication Editor
Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 0376-6357 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 6587
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Author Stomp, M.; d'Ingeo, S.; Henry, S.; Cousillas, H.; Hausberger, M.
Title Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model Type Journal Article
Year (up) 2021 Publication Applied Animal Behaviour Science Abbreviated Journal Appl. Anim. Behav. Sci.
Volume 236 Issue Pages 105271
Keywords Laterality; Electroencephalography; Theta wave; Welfare; Horses
Abstract Assessing the animal welfare state is a challenge given the subjective individual cognitive and emotional processing involved. Electroencephalography (EEG) spectrum analysis has proved an ecologically valid recording situation to assess the link between brain processes and affective or cognitive states in humans: a higher slow wave/fast wave ratio has been associated with a positive internal state. In particular, a high production of theta power (3-8 Hz) has been related to positive emotions. On the other hand, it has been hypothesized that a left hemisphere (LH) dominance may be associated with a better welfare state. Here, we test the hypothesis that individual differences in the resting-state quantitative EEG power spectrum of adult horses (N = 18) and its lateralization pattern may reflect individual subjective perception of their conditions of life and welfare state. The results show clear individual differences in the proportions of the different waves and their inter-hemispheric distribution. Three different EEG power spectrum profiles were highlighted, from a bilateral predominance of theta waves in horses in a more positive welfare state to a bilateral predominance of beta waves in horses with clear expressions of compromised welfare. Interestingly, particular correlations were found between wave power activity and welfare parameters. We found a negative correlation between the number of stereotypic behaviours per hour and the median proportion of theta waves in the left hemisphere. and between the overall state (total chronic stress score) of welfare and gamma production in the right hemisphere (RH). These findings go along the hypothesis of a particular involvement of the left hemisphere for positive processing and of the right hemisphere for negative processing. However, the pattern of laterality did not appear as the most important feature here as both extreme clusters in terms of welfare showed bilateral predominance of one wave type. It is possible that hemispheric specialization makes more sense during acute emotion-inducing conditions rather than in this resting-state context (i.e. in absence of any high emotion-inducing stimulation), although the opposition gamma versus theta waves between both hemispheres in the horses with an intermediate welfare state is noticeable and intriguing. It seems that bilateral but also LH theta activity is a promising neurophysiological marker of good welfare in horses, while a bilateral or RH high production of gamma waves should alert about potential welfare alterations. Quantitative resting-state EEG power spectrum appears as a highly promising tool for exploring the brain processes involved in the subjective perception of chronic welfare, as a useful complementary tool for welfare assessment.
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Publisher Place of Publication Editor
Language Summary Language Original Title
Series Editor Series Title Abbreviated Series Title
Series Volume Series Issue Edition
ISSN 0168-1591 ISBN Medium
Area Expedition Conference
Notes Approved no
Call Number Equine Behaviour @ team @ Serial 6628
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