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Versace, E.; Morgante, M.; Pulina, G.; Vallortigara, G. |
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Title |
Behavioural lateralization in sheep (Ovis aries) |
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Journal Article |
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Year |
2007 |
Publication |
Behavioural Brain Research |
Abbreviated Journal |
Behav. Brain. Res. |
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184 |
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1 |
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72-80 |
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Keywords |
Lateralization; Laterality; Brain asymmetry; Hemisphere; Sheep; Lamb; Strength of lateralization |
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This study investigates behavioural lateralization in sheep and lambs of different ages. A flock was tested in a task in which the animals were facing an obstacle and should avoid it on either the right or left side to rejoin flock-mates (adult sheep) or their mothers (lambs). A bias for avoiding the obstacle on the right side was observed, with lambs apparently being more lateralized than sheep. This right bias was tentatively associated with the left-hemifield laterality in familiar faces recognition which has been documented in this species. Differences between adult sheep and lambs were likely to be due to differences in social reinstatement motivation elicited by different stimuli (flock-mates or mothers) at different ages. Preferential use of the forelegs to step on a wood-board and direction of jaw movement during rumination was also tested in adult animals. No population bias nor individual-level lateralization was observed for use of the forelegs. At the same time, however, there was a large number of animals showing individual-level lateralization for the direction of jaw movement during rumination even though there was no population bias. These findings highlight that within the same species individual- and population-level lateralization can be observed in different tasks. Moreover, the results fit the general hypothesis that population-level asymmetries are more likely to occur in tasks that require social coordination among behaviourally asymmetric individuals. |
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0166-4328 |
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Equine Behaviour @ team @ |
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6701 |
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Krueger, K.; Schwarz, S.; Marr, I.; Farmer, K. |
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Title |
Laterality in Horse Training: Psychological and Physical Balance and Coordination and Strength Rather Than Straightness |
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Magazine Article |
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Year |
2022 |
Publication |
Animals |
Abbreviated Journal |
Animals |
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12 |
Issue |
8 |
Pages |
1042 |
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Keywords |
balance; body asymmetry; equitation; horse; motor laterality; sensory laterality; stress; welfare |
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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. |
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Animals |
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12 |
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8 |
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2076-2615 |
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Equine Behaviour @ team @ |
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6670 |
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Schwarz, S.; Marr, I.; Farmer, K.; Graf, K.; Stefanski, V.; Krueger, K. |
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Title |
Does Carrying a Rider Change Motor and Sensory Laterality in Horses? |
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Journal Article |
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Year |
2022 |
Publication |
Animals |
Abbreviated Journal |
Animals |
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12 |
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8 |
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992 |
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Keywords |
laterality; horse; rider; sensory laterality; motor laterality; novel object; side preference |
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Laterality in horses has been studied in recent decades. Although most horses are kept for riding purposes, there has been almost no research on how laterality may be affected by carrying a rider. In this study, 23 horses were tested for lateral preferences, both with and without a rider, in three different experiments. The rider gave minimal aids and rode on a long rein to allow the horse free choice. Firstly, motor laterality was assessed by observing forelimb preference when stepping over a pole. Secondly, sensory laterality was assessed by observing perceptual side preferences when the horse was confronted with (a) an unfamiliar person or (b) a novel object. After applying a generalised linear model, this preliminary study found that a rider increased the strength of motor laterality (p = 0.01) but did not affect sensory laterality (p = 0.8). This suggests that carrying a rider who is as passive as possible does not have an adverse effect on a horse�s stress levels and mental state. |
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Equine Behaviour @ team @ |
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6667 |
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Author |
Lucidi, P.; Bacco, G.; Sticco, M.; Mazzoleni, G.; Benvenuti, M.; Bernabò, N.; Trentini, R. |
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Title |
Assessment of motor laterality in foals and young horses (Equus caballus) through an analysis of derailment at trot |
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Journal Article |
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Year |
2013 |
Publication |
Physiology & Behavior |
Abbreviated Journal |
Physiol. Behav. |
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109 |
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8-13 |
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Keywords |
Horse; Laterality; Longeing; Sidedness |
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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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0031-9384 |
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Equine Behaviour @ team @ |
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6666 |
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Stomp, M.; d'Ingeo, S.; Henry, S.; Cousillas, H.; Hausberger, M. |
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Title |
Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model |
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Journal Article |
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Year |
2021 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
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Volume |
236 |
Issue |
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Pages |
105271 |
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Keywords |
Laterality; Electroencephalography; Theta wave; Welfare; Horses |
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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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0168-1591 |
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Equine Behaviour @ team @ |
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6629 |
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Author |
Stomp, M.; d'Ingeo, S.; Henry, S.; Cousillas, H.; Hausberger, M. |
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Title |
Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model |
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Journal Article |
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Year |
2021 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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236 |
Issue |
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Pages |
105271 |
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Keywords |
Laterality; Electroencephalography; Theta wave; Welfare; Horses |
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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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0168-1591 |
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Equine Behaviour @ team @ |
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6628 |
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Author |
Giljov, A.; Karenina, K. |
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Title |
Differential roles of the right and left brain hemispheres in the social interactions of a free-ranging ungulate |
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Journal Article |
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Year |
2019 |
Publication |
Behavioural Processes |
Abbreviated Journal |
Behav. Process. |
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168 |
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Pages |
103959 |
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Keywords |
Laterality; Hemispheric specialization; Brain asymmetry; Eye preference; Ungulate; Bovid |
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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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0376-6357 |
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Equine Behaviour @ team @ |
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6587 |
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Author |
Sankey, C.; Henry, S.; Clouard, C.; Richard-Yris, M.-A.; Hausberger, M. |
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Title |
Asymmetry of behavioral responses to a human approach in young naive vs. trained horses |
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Journal Article |
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Year |
2011 |
Publication |
Physiology & Behavior |
Abbreviated Journal |
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104 |
Issue |
3 |
Pages |
464-468 |
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Keywords |
Laterality; Emotionality; Human approach; Horse |
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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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0031-9384 |
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Equine Behaviour @ team @ |
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5688 |
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Author |
Komárková, M.; Bartošová, J. |
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Title |
Lateralized suckling in domestic horse foals (Equus caballus) |
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Conference Article |
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2012 |
Publication |
Proceedings of the 2. International Equine Science Meeting |
Abbreviated Journal |
Proc. 2. Int. Equine. Sci. Mtg |
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in press |
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laterality; suckling; horse; foal |
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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. |
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Xenophon Publishing |
Place of Publication |
Wald |
Editor |
Krueger, K. |
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978-3-9808134-26 |
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Equine Behaviour @ team @ |
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5505 |
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Author |
Krueger, K; Farmer, K. |
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Title |
Laterality in the Horse [Lateralität beim Pferd ] |
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Journal Article |
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Year |
2011 |
Publication |
mensch & pferd international |
Abbreviated Journal |
mup |
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4 |
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Pages |
160-167 |
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Keywords |
Laterality, horse, information processing, training, welfare, human-animal interaction |
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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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Equine Behaviour @ team @ |
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5444 |
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