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In a recent experiment, we showed that horses are sensitive to pet-directed speech (PDS), a kind of speech used to talk to companion animals that is characterized by high pitch and wide pitch variations. When talked to in PDS rather than adult-directed speech (ADS), horses reacted more favorably during grooming and in a pointing task. However, the mechanism behind their response remains unclear: does PDS draw horses’ attention and arouse them, or does it make their emotional state more positive? In this study, we used an innovative paradigm in which female horses watched videos of humans speaking in PDS or ADS to better understand this phenomenon. Horses reacted differently to the videos of PDS and ADS: they were significantly more attentive and their heart rates increased significantly more during PDS than during ADS. We found no difference in the expressions of negative or positive emotional states during PDS and ADS videos. Thus, we confirm that horses’ perception of humans can be studied by means of video projections, and we conclude that PDS attracts attention and has an arousing effect in horses, with consequences on the use of PDS in daily interactions with them.

Several studies have shown that horses have the ability to cross-modally recognize humans by associating their voice with their physical appearance. However, it remains unclear whether horses are able to differentiate humans according to different criteria, such as the fact that they are women or men. Horses might recognize some human characteristics, such as sex, and use these characteristics to classify them into different categories. The aim of this study was to explore whether domesticated horses are able to cross-modally recognize women and men according to visual and auditory cues, using a preferential looking paradigm. We simultaneously presented two videos of women and men’s faces, while playing a recording of a human voice belonging to one of these two categories through a loudspeaker. The results showed that the horses looked significantly more towards the congruent video than towards the incongruent video, suggesting that they are able to associate women’s voices with women’s faces and men’s voices with men’s faces. Further investigation is necessary to determine the mechanism underlying this recognition, as it might be interesting to determine which characteristics horses use to categorize humans. These results suggest a novel perspective that could allow us to better understand how horses perceive humans.

The welfare of an animal is closely linked to their emotional experiences, making it essential to identify reliable indicators of these emotions. This study aimed to identify behaviours and facial movements in horses experiencing contrasting emotional valence, triggered by the anticipation of a positive condition (going to pasture) or a negative condition (going alone to a novel environment). Twenty horses were daily trained to wait in a starting box before being exposed to these two conditions. After one week of positive training or negative training, we analysed horses’ behaviours, cortisol variations, and facial movements while they waited in their starting box. First, we confirmed that the two conditions induced contrasting emotional valence, as evidenced by the shorter time taken to approach in the positive condition compared to the negative, and by the higher maximal heart rate and cortisol variation in the negative condition. Then using the Equine Facial Action Coding System (EquiFACS) and network analysis (NetFACS) we revealed distinct behaviours and facial expression profiles. In positive anticipation, the horses exhibited a greater range of behaviours, including shaking their heads from side to side, stepping back, sniffing, and pawing at the ground. Additionally, two distinct facial expression profiles were identified as specific to positive and negative anticipation. In positive anticipation, the horses displayed a higher neck, accompanied by a greater frequency of half-blinks and mouth movements. Conversely, in negative anticipation, the horses exhibited a medium neck, with ears backward accompanied by more flattened ears and expressed more nostril movements. The findings highlight the importance of these indicators in characterising horses’ emotions and emphasise their significance for assessing equine welfare.

Social interactions shape both the physiological and behavioural development of offspring, and poor care/early caregiver loss is known to promote adverse outcomes during infancy in both animals and humans. How affiliative behaviours impact the future development of offspring remains an open question. Here, we used Equus caballus (domestic horse) as a model to investigate this question. By coupling magnetic resonance imaging, longitudinal biobehavioural assessments and advanced multivariate statistical modelling, we found that prolonged maternal presence during infancy promotes the maturation of brain regions involved in both social behaviour (anterior cingulate cortex and retrosplenial cortex) and physiological regulation (hypothalamus and amygdala). Additionally, offspring benefiting from a prolonged maternal presence showed higher default mode network connectivity, improved social competences and feeding behaviours, and higher concentrations of circulating lipids (triglyceride and cholesterol). The findings of the present study underscore the salient role of social interactions in the development of allostatic regulation in offspring. The present study shows that maternal presence beyond early life remains crucial for brain, behavioural and physiological development in young horses, highlighting the importance of the mother–offspring relationship during a childhood-like stage.

Emotional contagion, the emotional state-matching of two individuals, has been documented in various species. Recent findings suggest emotional contagion could also take place between humans and domestic mammals. However, the range of targeted animal species and human emotions that have been studied is still limited, and the methodology to investigate emotional contagion in this context is not fully established. This study examined emotional contagion of fear and joy from humans to horses by measuring physiological (heart rate, infrared thermography) and behavioral responses (posture, laterality bias, facial expressions) to videos of human fear, joy, or neutral emotions. Horses (n = 45) exhibited higher heart rates and ear movements during the fear and joy videos compared to the neutral ones, suggesting heightened arousal. During fear videos, they showed a greater increase in eye temperature than during joy or neutral videos, and maintained an alert posture for longer than during neutral videos, expressing specific facial expressions including inner brow raising and blowing. During joy videos, they showed a higher right eye preference (indicating a higher left-hemisphere bias) than during neutral videos, suggesting a more positive emotional valence. These findings suggest that human facial and vocal cues of fear and joy trigger emotional contagion in horses, which may influence the human-horse relationship.

Recently, research on domestic mammals’ sociocognitive skills toward humans has been prolific, allowing us to better understand the human–animal relationship. For example, horses have been shown to distinguish human beings on the basis of photographs and voices and to have cross-modal mental representations of individual humans and human emotions. This leads to questions such as the extent to which horses can differentiate human attributes such as age. Here, we tested whether horses discriminate human adults from children. In a cross-modal paradigm, we presented 31 female horses with two simultaneous muted videos of a child and an adult saying the same neutral sentence, accompanied by the sound of an adult’s or child’s voice speaking the sentence. The horses looked significantly longer at the videos that were incongruent with the heard voice than at the congruent videos. We conclude that horses can match adults’ and children’s faces and voices cross-modally. Moreover, their heart rates increased during children’s vocalizations but not during adults’. This suggests that in addition to having mental representations of adults and children, horses have a stronger emotional response to children’s voices than adults’ voices.

The widespread failure of anthelmintic drugs against nematodes of veterinary interest requires novel control strategies. Selective treatment of the most susceptible individuals could reduce drug selection pressure but requires appropriate biomarkers of the intrinsic susceptibility potential. To date, this has been missing in livestock species. Here, we selected Welsh ponies with divergent intrinsic susceptibility (measured by their egg excretion levels) to cyathostomin infection and found that their divergence was sustained across a 10-year time window. Using this unique set of individuals, we monitored variations in their blood cell populations, plasma metabolites and faecal microbiota over a grazing season to isolate core differences between their respective responses under worm-free or natural infection conditions. Our analyses identified the concomitant rise in plasma phenylalanine level and faecal Prevotella abundance and the reduction in circulating monocyte counts as biomarkers of the need for drug treatment (egg excretion above 200 eggs/g). This biological signal was replicated in other independent populations. We also unravelled an immunometabolic network encompassing plasma beta-hydroxybutyrate level, short-chain fatty acid producing bacteria and circulating neutrophils that forms the discriminant baseline between susceptible and resistant individuals. Altogether our observations open new perspectives on the susceptibility of equids to strongylid infection and leave scope for both new biomarkers of infection and nutritional intervention.

Myeloperoxidase (MPO), as a marker of neutrophil activation, has been associated with equine endometritis. However, in absence of inflammation, MPO is constantly detected in the uterine lumen of estrous mares. The aim of this study was to characterize MPO in the uterus of mares under physiological conditions as a first step to better understand the role of this enzyme in equine reproduction. Total and active MPO concentrations were determined, by ELISA and SIEFED assay, respectively, in low-volume lavages from mares in estrus (n = 26), diestrus (n = 18) and anestrus (n = 8) in absence of endometritis. Immunohistochemical analysis was performed on 21 endometrial biopsies randomly selected: estrus (n = 11), diestrus (n = 6) and anestrus (n = 4). MPO, although mostly enzymatically inactive, was present in highly variable concentrations in uterine lavages in all studied phases, with elevated concentrations in estrus and anestrus, while in diestrus, concentrations were much lower. Intracytoplasmic immunoexpression of MPO was detected in the endometrial epithelial cells, neutrophils and glandular secretions. Maximal expression was observed during estrus in mid and basal glands with a predominant intracytoplasmic apical reinforcement. In diestrus, immunopositive glands were sporadic. In anestrus, only the luminal epithelium showed residual MPO immunostaining. These results confirm a constant presence of MPO in the uterine lumen of mares in absence of inflammation, probably as part of the uterine mucosal immune system, and suggest that endometrial cells are a source of uterine MPO under physiological cyclic conditions.

Precision livestock farming using omics approach to acquire precise and real-time data can help farmers in individual animal management and decision making. Since steroid hormones play a key role for the regulation of reproductive functions, reproduction management could be improved by characterizing the steroidome during reproductive stages. Moreover, saliva collection is a non-invasive, painless, inexpensive and easy sampling method. Thus, this prospective study proposes a steroidomic analysis in mare saliva during reproductive stages, that could help to identify potential biomarkers to accurately detect their reproductive stage in a welfare friendly production system, for real-time decision making at the individual animal level. Correlation between saliva and plasma steroidome was also investigated. Saliva and blood samples from 6 mares were collected in anestrus, in the follicular phase 3 days, 2 days and 1 day before ovulation and the day when ovulation was detected, in the luteal phase 6 days after ovulation and in gestation 18 days after ovulation and insemination. Steroidome analysis was performed by gas chromatography coupled to tandem mass spectrometry (GC-MS/MS). We characterized 25 and 36 steroids in saliva and plasma respectively. Pregnenolone concentrations in saliva during gestation were significantly higher than during anestrus or follicular phase and tended to be higher than during luteal phase. Most of the 5α-reduced metabolites of progesterone showed higher salivary concentrations during the luteal phase 6 days post-ovulation and during gestation 18 days post-ovulation compared to anestrus and follicular phase. These steroids could be potential salivary biomarkers of the reproductive stage of the mare. Further studies with a greater number of animals are in progress to confirm the reliability of these potential candidate biomarkers and to develop field-friendly assays.

Olfaction is the most widespread sensory modality animals use to communicate, yet much remains to be discovered about its role. While most studies focused on intraspecific interactions and reproduction, new evidence suggests chemosignals may influence interspecific interactions and emotional communication. This study explores this possibility, investigating the potential role of olfactory signals in human-horse interactions. Cotton pads carrying human odours from fear and joy contexts, or unused pads (control odour) were applied to 43 horses’ nostrils during fear tests (suddenness and novelty tests) and human interaction tests (grooming and approach tests). Principal component analysis showed that overall, when exposed to fear-related human odours, horses exhibited significantly heightened fear responses and reduced interaction with humans compared to joy-related and control odours. More precisely, when exposed to fear-related odours, horses touched the human less in the human approach test (effect size: Rate Ratio(RR)=0.60 ± 0.24), gazed more at the novel object (RR = 1.32 ± 0.14), and were more startled (startle intensity – Cohen’s d = −0.88 ± 0.39; and maximum heart rate – Cohen’s d = 1.16 ± 0.47) by a sudden event. These results highlight the significance of chemosignals in interspecific interactions and provide insights into questions about the impact of domestication on emotional communication. Moreover, these findings have practical implications regarding the significance of handlers’ emotional states and its transmission through odours during human-horse interactions.