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.

Abstract: Lateralization in dogs, Canis familiaris, has been reported for paw usage and response to visual and acoustic stimuli. Surprisingly, however, no investigation of possible lateralization for the most relevant sensory domain of dogs, namely olfaction, has been carried out. Here we investigated left and right nostril use in dogs freely sniffing different emotive stimuli in unrestrained conditions. When sniffing novel nonaversive stimuli (food, lemon, vaginal secretion and cotton swab odours), dogs showed initial preferential use of the right nostril and then a shift towards use of the left nostril with repeated stimulus presentation. When sniffing arousal stimuli such as adrenaline and veterinary sweat odorants, dogs showed a consistent right nostril bias all over the series of stimulus presentations. Results suggest initial involvement of the right hemisphere in processing of novel stimuli followed by the left hemisphere taking charge of control of routine behaviour. Sustained right nostril response to arousal stimuli appears to be consistent with the idea that the sympathetic hypothalamic-pituitary-adrenal axis is mainly under the control of the right hemisphere. The implications of these findings for animal welfare are discussed.

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.

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.

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.