Studying vocal correlates of emotions is important to provide a better understanding of the evolution of emotion expression through cross-species comparisons. Emotions are composed of two main dimensions: emotional arousal (calm versus excited) and valence (negative versus positive). These two dimensions could be encoded in different vocal parameters (segregation of information) or in the same parameters, inducing a trade-off between cues indicating emotional arousal and valence. We investigated these two hypotheses in horses. We placed horses in five situations eliciting several arousal levels and positive as well as negative valence. Physiological and behavioral measures collected during the tests suggested the presence of different underlying emotions. First, using detailed vocal analyses, we discovered that all whinnies contained two fundamental frequencies (“F0” and “G0”), which were not harmonically related, suggesting biphonation. Second, we found that F0 and the energy spectrum encoded arousal, while G0 and whinny duration encoded valence. Our results show that cues to emotional arousal and valence are segregated in different, relatively independent parameters of horse whinnies. Most of the emotion-related changes to vocalizations that we observed are similar to those observed in humans and other species, suggesting that vocal expression of emotions has been conserved throughout evolution.

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.

BackgroundNon-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.ResultsWe 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.ConclusionsHorses 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.Electronic supplementary materialThe online version of this article (doi:10.1186/s12983-017-0193-1) contains supplementary material, which is available to authorized users.