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Stomp, M., d'Ingeo, S., Henry, S., Cousillas, H., & Hausberger, M. (2021). Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model. Appl. Anim. Behav. Sci., 236, 105271.
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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Stomp, M., d'Ingeo, S., Henry, S., Cousillas, H., & Hausberger, M. (2021). Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model. Applied Animal Behaviour Science, 236, 105271.
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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Giljov, A., & Karenina, K. (2019). Differential roles of the right and left brain hemispheres in the social interactions of a free-ranging ungulate. Behav. Process., 168, 103959.
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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Stennett, C. R., & Strauss, R. E. (2010). Behavioural lateralization in zebrafish and four related species of minnows (Osteichthyes: Cyprinidae). Anim. Behav., 79(6), 1339–1342.
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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Ruggieri, V. (1999). The running horse stops: the hypothetical role of the eyes in imagery of movement. Percept Mot Skills, 89(3 Pt 2), 1088–1092.
Abstract: To examine the hypothetical role of the eyes in visual mental imagery of movement 72 undergraduate women students in psychology were asked to imagine a running horse and then to produce the same mental image without moving the eyes and the head. In 59% of the subjects interesting modifications of the imagined movement appeared: 37% observed an inhibition of the movement and 19% an evident slowing up of the moving figure. The interpretation of this result was made by hypothesizing that the eyes are concretely involved in visual imagery processes.
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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.
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Schwarz, S., Marr, I., Farmer, K., Graf, K., Stefanski, V., & Krueger, K. (2022). Does Carrying a Rider Change Motor and Sensory Laterality in Horses? Animals, 12(8), 992.
Abstract: 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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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.
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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.
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Siniscalchi, M., Sasso, R., Pepe, A. M., Vallortigara, G., & Quaranta, A. (2010). Dogs turn left to emotional stimuli. Behav. Brain. Res., 208(2), 516–521.
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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