Abstract: The Eneolithic Botai culture of the Central Asian steppes provides the earliest archaeological evidence for horse husbandry, ~5,500 ya, but the exact nature of early horse domestication remains controversial. We generated 42 ancient horse genomes, including 20 from Botai. Compared to 46 published ancient and modern horse genomes, our data indicate that Przewalski's horses are the feral descendants of horses herded at Botai and not truly wild horses. All domestic horses dated from ~4,000 ya to present only show ~2.7% of Botai-related ancestry. This indicates that a massive genomic turnover underpins the expansion of the horse stock that gave rise to modern domesticates, which coincides with large-scale human population expansions during the Early Bronze Age.

Abstract: Evidence suggests that behavioural affective reactions to sweet and bitter substances are homologous in humans, nonhuman primates, and rodents. The sweet taste of sucrose elicits facial responses that include rhythmic tongue protrusions whereas the bitter taste of quinine elicits facial responses that include gapes, featuring an opening of the mouth and protrusion of the tongue. The present study using the horse (Equus caballus) was undertaken for three reasons: (1) there is debate about the presence of a sweet receptor gene in the horse, (2) there is a need to expand the examination of facial reactions to taste in lineages other than the closely related lineages of rodents and primates, and (3) the horse provides an opportunity to test the hypothesis that some social signals derive from movements related to taste reaction. The horses were given oral infusions of either sucrose or quinine and their behaviour was examined using frame-by-frame video analysis. Control groups were exposed received water or syringe insertion only. Amongst the many responses made to the infusions, the distinctive response to sucrose was a bob coupled with a slight tongue protrusion and forward movement of the ears; the distinctive response to quinine was a head extension and mouth gape accompanied by a large tongue protrusion and backward movement of the ears. Sucrose Bobs and Quinine Gapes are discussed with respect to: (1) the relevance of facial reactions to both sucrose and quinine to taste receptors in horses, (2) the similarity of features of taste expression in horses to those documented in rodents and primates, and (3) the dissimilarity between facial reactions to taste and other social signals displayed by 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.

Abstract: Many breeding organisations include a subjective scoring of rideability by a professional rider into their evaluation of sports horses, but the consistency and reliability of the scoring system is debateable. The aim of this study was to investigate (i) whether professional riders agree in their scoring of rideability, and (ii) whether rideability scores are affected by rein tension, horse conflict behaviour, heart rate, and salivary cortisol, and (iii) whether riders induce different levels of conflict behaviour and physiological responses in the horses. Ten professional, female riders each rode 10 dressage horses (level M German scale; n = 100 combinations) through a standardised dressage test (10 min warm-up followed by a 4-min test) and subsequently scored the horses for rideability on the official 1-10 scale (1 = poor to 10 = excellent) from the Danish Riding Federation. Rein tension, horse heart rate, saliva cortisol and conflict behaviour were measured for each rider-horse pair. The riders were inconsistent in their scoring of rideability to the individual horses, e.g. scores for one of the horses ranged from 1 to 8. There was a significant effect of rider (P = 0.003) and the frequency of conflict behaviour (undesired head movements: P < 0.001, breaking the gait: P = 0.013, and other evasive behaviour: P = 0.032) on rideability scores, i.e. the more conflict behaviour the lower the score. There was no significant effect of rein tension and the physiological measures on rideability scores. However, there was a significant effect of rider on rein tension, horses' heart rate and increases in saliva cortisol concentrations and a tendency for some types of conflict behaviour, suggesting that some riders induced more discomfort in the horses. Future studies could help shed light on which elements of riding style are particularly important for sports horse welfare. In conclusion, this study found a large variation in rideability scores assigned to ten sports horses by ten professional riders. Rideability scores were dependent on the level of horse conflict behaviour, but not rein tension and physiological measures. Further studies are needed to improve the objectivity, consistency and reliability of rideability assessment of sports horses.

Abstract: Growing evidence shows a variety of sensorial and motor asymmetries in social and non-social interactions in various species, indicating a lateralized processing of information by the brain. Using digital video cameras on tripods and drones, this study investigated lateralization in frequency and duration of social behavior patterns, in affiliative, agonistic, and resting contexts, in a feral population of horses (Equus ferus caballus) in Northern Portugal, consisting of 37 individuals organized in eight harem groups. Affiliative interactions (including grooming) were more often performed, and lasted longer, when recipients were positioned to the right side. In recumbent resting (animals lying down) episodes on the left side lasted longer. Our results of an affiliative behavior having a right side tendency, provide partial support to the valence-specific hypothesis of Ahern and Schwartz (1979) – left hemisphere dominance for positive affect, affiliative behaviors. Longer recumbent resting episodes on the left side may be due to synchronization. However, in both instances it is discussed how lateralization may be context dependent. Investigating the position asymmetries of social behaviors in feral equids will contribute to a better understanding of differential lateralization and hemispheric specialization from the ecological and evolutionary perspectives.