Abstract: Selection pressures during domestication are thought to lead to an enhanced ability to use human-given cues. Horses fulfil a wide variety of roles for humans and have been domesticated for at least 5000 years but their ability to read human cues has not been widely studied. We tested the ability of 28 horses to attend to human-given cues in an object choice task. We included five different cues: distal sustained pointing, momentary tapping, marker placement, body orientation and gaze (head) alternation. Horses were able to use the pointing and marker placement cues spontaneously but not the tapping, body orientation and gaze alternation cues. The overall pattern of responding suggests that horses may use cues that provide stimulus enhancement at the time of choice and do not have an understanding of the communicative nature of the cues given. As such, their proficiency at this task appears to be inferior to that of domestic dogs, Canis lupus familiaris, but similar to that of domestic goats, Caprus hircus.

Abstract: Whether non-human animals can recognize human signals, including emotions, has both scientific and applied importance, and is particularly relevant for domesticated species. This study presents the first evidence of horses' abilities to spontaneously discriminate between positive (happy) and negative (angry) human facial expressions in photographs. Our results showed that the angry faces induced responses indicative of a functional understanding of the stimuli: horses displayed a left-gaze bias (a lateralization generally associated with stimuli perceived as negative) and a quicker increase in heart rate (HR) towards these photographs. Such lateralized responses towards human emotion have previously only been documented in dogs, and effects of facial expressions on HR have not been shown in any heterospecific studies. Alongside the insights that these findings provide into interspecific communication, they raise interesting questions about the generality and adaptiveness of emotional expression and perception across species.

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.

Abstract: Recent research has shown that domestic dogs are particularly good at reading human attentional cues, often outperforming chimpanzees and hand reared wolves [1, 2]. It has been suggested that the close evolutionary relationship between humans and dogs has led to the development of this ability, however very few other species have been studied [3]. We tested the ability of 24 domestic horses to discriminate between an attentive and inattentive person when choosing whom to approach for food. While the attentive person faced forwards, the inattentive person either stood with their body turned 180° away from the subject (body orientation condition), stood with their body facing forwards but their head facing away (head orientation condition) or stood facing forwards but with their eyes closed (eyes closed condition). A fourth, mixed condition was included where the attentive person stood with their body facing away from the subjects but their head turned towards the subject while the inattentive person stood with their body facing the subject but their head turned away. Horses chose the attentive person significantly more often using the body cue (n = 24, k = 19, p = 0.003), the head cue (n = 24, k = 18, p = 0.011), and the eye cue (n = 24, k = 19, p = 0.003) but not the mixed cue (n = 24, k = 13, p = 0.42). In an additional pilot study, horses were tested in an object choice task. A human experimenter cued one of two buckets by either tapping the bucket (tap condition), orienting their body towards the bucket and pointing (body and point condition), pointing while facing forwards (point condition) or orienting their body towards the bucket (body condition). If the subjects chose the correct bucket they were rewarded. Subjects were able to use the tap cue (n = 31, k = 21, p = 0.035), body + point cue (n= 31, k = 21, p = 0.035) and the point cue (n = 30, k = 21, p = 0.021) but not the body cue (n = 31, k = 11, p = 0.076). These results taken together suggest that domestic horses are also very sensitive to human attentional cues, including gaze.
Keywords:
social cognition, animal-human interaction, horses, attention attribution, domestication
1. Hare, B., Brown, M., Williamson, C., and Tomasello, M. (2002). The domestication of social cognition in dogs. Science 298, 1634-1636.
2. Gacsi, M., Miklosi, A., Varga, O., Topal, J., and Csanyi, V. (2004). Are readers of our face readers of our minds` Dogs (Canis familiaris) show situation-dependent recognition of human’s attention. Animal Cognition 7, 144-153.
3. Hare, B., and Tomasello, M. (2005). Human-like social skills in dogs? Trends Cogn. Sci. 9, 439-444.

Abstract: <p>It has been suggested that the process of domestication, at least in some species, has led to an innate predisposition to be skilled at reading human communicative and attentional cues. Adult domestic horses (<italic>Equus caballus</italic>) are highly sensitive to subtle bodily cues when determining if a person is attending to them but they are less adept at using human communicative cues in object choice tasks. Here we provide the first study into the ontogeny of such skills in order to gain insights into the mechanisms underlying these abilities. Compared with adult horses, youngsters under the age of three could use body orientation but not more subtle cues such as head movement and open/closed eyes to correctly choose an attentive person to approach for food. Across two object choice experiments, the performance of young horses was comparable to that of adult horses – subjects were able to correctly choose a rewarded bucket using marker placement, pointing and touching cues but could not use body orientation, gaze, elbow pointing or tapping cues. Taken together these results do not support the theory that horses possess an innate predisposition to be particularly skilled at using human cues. Horses' ability to determine whether humans are attending to them using subtle body cues appears to require significant experience to fully develop and their perhaps less remarkable use of limited cues in object choice tasks, although present at a much earlier age, is likely to reflect a more general learning ability related to stimulus enhancement rather than a specific ‘human-reading’ skill.</p>