Abstract: Using heart rate (HR) measurements we investigated whether potential calming effects of social licking were evident for both active (performers) and passive (receivers) licking partners. A HR decline was assumed to indicate relaxation and thus the experience of positive emotions. Effects of the licking category (spontaneous, solicited), the animals' basic activity (standing, lying) and the licked body region (head, neck, rest) were also considered. Two studies (A, B) were carried out in the same loose housed Austrian Simmental dairy herd. HR was recorded in up to 20 focal animals on 16 and 18 days, respectively. Using either direct observations (A) or video recordings (B), social licking interactions were continuously observed. The cow's basic activity was recorded using scan sampling at 5 min intervals. Linear mixed effects models were applied separately for Study A and B in order to compare the mean HR of the licking bouts with the mean of the respective 5 min pre- and post-licking periods. In receivers we found a significant calming effect in terms of a HR decline during allogrooming in both studies (A: -1.3 beats per minute, B: -1.1 bpm). This effect was more pronounced when animals were standing (A/B: -2.4 bpm/-3.8 bpm). However, it was not affected by the licked body region. In dairy cows performing social licking, we did not find an overall calming effect. On the contrary, in Study B, HR significantly increased during licking in lying performers (+2.5 bpm). This reaction was even stronger, when licking was directed to the receivers' head (+3.5 bpm) or neck (+3.0 bpm) as compared to the rest of the body (+1.4 bpm). The licking category had no effect on HR changes during the licking events. Our findings suggest that relaxation effects induced by social licking differ between performers and receivers and are affected by the cows' basic activity. In receivers, there were clear indications of a calming effect implying the experience of positive affective states. In performers, such calming effects during social licking were not identified.

Abstract: Summary For humans, facial expressions are important social signals, and how we perceive specific individuals may be influenced by subtle emotional cues that they have given us in past encounters. A wide range of animal species are also capable of discriminating the emotions of others through facial expressions [1, 2, 3, 4, 5], and it is clear that remembering emotional experiences with specific individuals could have clear benefits for social bonding and aggression avoidance when these individuals are encountered again. Although there is evidence that non-human animals are capable of remembering the identity of individuals who have directly harmed them [6, 7], it is not known whether animals can form lasting memories of specific individuals simply by observing subtle emotional expressions that they exhibit on their faces. Here we conducted controlled experiments in which domestic horses were presented with a photograph of an angry or happy human face and several hours later saw the person who had given the expression in a neutral state. Short-term exposure to the facial expression was enough to generate clear differences in subsequent responses to that individual (but not to a different mismatched person), consistent with the past angry expression having been perceived negatively and the happy expression positively. Both humans were blind to the photograph that the horses had seen. Our results provide clear evidence that some non-human animals can effectively eavesdrop on the emotional state cues that humans reveal on a moment-to-moment basis, using their memory of these to guide future interactions with particular individuals.