Non-human animals often produce different types of vocalisations in negative and positive contexts (i.e. different valence), similar to humans, in which crying is associated with negative emotions and laughter is associated with positive ones. However, some types of vocalisations (e.g. contact calls, human speech) can be produced in both negative and positive contexts, and changes in valence are only accompanied by slight structural differences. Although such acoustically graded signals associated with opposite valence have been highlighted in some species, it is not known if conspecifics discriminate them, and if contagion of emotional valence occurs as a result. We tested whether domestic horses perceive, and are affected by, the emotional valence of whinnies produced by both familiar and unfamiliar conspecifics. We measured physiological and behavioural reactions to whinnies recorded during emotionally negative (social separation) and positive (social reunion) situations. We show that horses perceive acoustic cues to both valence and familiarity present in whinnies. They reacted differently (respiration rate, head movements, height of the head and latency to respond) to separation and reunion whinnies when produced by familiar, but not unfamiliar individuals. They were also more emotionally aroused (shorter inter-pulse intervals and higher locomotion) when hearing unfamiliar compared to familiar whinnies. In addition, the acoustic parameters of separation and reunion whinnies affected the physiology and behaviour of conspecifics in a continuous way. However, we did not find clear evidence for contagion of emotional valence. Horses are thus able to perceive changes linked to emotional valence within a given vocalisation type, similar to perception of affective prosody in humans. Whinnies produced in either separation or reunion situations seem to constitute acoustically graded variants with distinct functions, enabling horses to increase their apparent vocal repertoire size.

Measuring emotions in nonhuman mammals is challenging. As animals are not able to verbally report how they feel, we need to find reliable indicators to assess their emotional state. Emotions can be described using two key dimensions: valence (negative or positive) and arousal (bodily activation or excitation). In this study, we investigated vocal expression of emotional valence in wild boars (Sus scrofa). The animals were observed in three naturally occurring situations: anticipation of a food reward (positive), affiliative interactions (positive), and agonistic interactions (negative). Body movement was used as an indicator of emotional arousal to control for the effect of this dimension. We found that screams and squeals were mostly produced during negative situations, and grunts during positive situations. Additionally, the energy quartiles, duration, formants, and harmonicity indicated valence across call types and situations. The mean of the first and second formants also indicated valence, but varied according to the call type. Our results suggest that wild boars can vocally express their emotional states. Some of these indicators could allow us to identify the emotional valence that wild boars are experiencing during vocal production and thus inform us about their welfare.

Vocal expression of emotions has been suggested to be conserved throughout evolution. However, since vocal indicators of emotions have never been compared between closely related species using similar methods, it remains unclear whether this is the case. Here, we investigated vocal indicators of emotional valence (negative versus positive) in Przewalski’s horses, in order to find out if expression of valence is similar between species and notably among Equidae through a comparison with previous results obtained in domestic horse whinnies. We observed Przewalski’s horses in naturally occurring contexts characterised by positive or negative valence. As emotional arousal (bodily activation) can act as a confounding factor in the search for indicators of valence, we controlled for its effect on vocal parameters using a behavioural indicator (movement). We found that positive and negative situations were associated with specific types of calls. Additionally, the acoustic structure of calls differed according to the valence. There were some similarities but also striking differences in expression of valence between Przewalski’s and domestic horses, suggesting that vocal expression of emotional valence, unlike emotional arousal, could be species specific rather than conserved throughout evolution.