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Author McComb, K.; Moss, C.; Sayialel, S.; Baker, L. url  doi
openurl 
  Title Unusually extensive networks of vocal recognition in African elephants Type Journal Article
  Year 2000 Publication (up) Anim Behav Abbreviated Journal  
  Volume 59 Issue Pages  
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  Area Expedition Conference  
  Notes Approved no  
  Call Number Equine Behaviour @ team @ McComb2000 Serial 6281  
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Author Proops, L.; Walton, M.; McComb, K. url  doi
openurl 
  Title The use of human-given cues by domestic horses, Equus caballus, during an object choice task Type Journal Article
  Year 2010 Publication (up) Animal Behaviour. Abbreviated Journal Anim. Behav.  
  Volume 79 Issue 6 Pages 1205-1209  
  Keywords domestication; Equus caballus; horse; interspecific communication; marker cue; pointing; social cognition  
  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.  
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  Series Editor Series Title Abbreviated Series Title  
  Series Volume Series Issue Edition  
  ISSN 0003-3472 ISBN Medium  
  Area Expedition Conference  
  Notes Approved no  
  Call Number Equine Behaviour @ team @ Serial 5083  
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Author Proops, L.; McComb, K. url  doi
openurl 
  Title Attributing attention: the use of human-given cues by domestic horses (Equus caballus) Type Journal Article
  Year 2010 Publication (up) Animal Cognition Abbreviated Journal Anim. Cogn.  
  Volume 13 Issue 2 Pages 197-205  
  Keywords  
  Abstract Abstract  Recent research has shown that domestic dogs are particularly good at determining the focus of human attention, often outperforming chimpanzees and hand-reared wolves. 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 domestic species have been studied. We tested the ability of 36 domestic horses to discriminate between an attentive and inattentive person in determining whom to approach for food. The cues provided were body orientation, head orientation or whether the experimenters’ eyes were open or closed. 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, head cue, and eye cue but not the mixed cue. This result suggests that domestic horses are highly sensitive to human attentional cues, including gaze. The possible role of evolutionary and environmental factors in the development of this ability is discussed.  
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  Notes Approved no  
  Call Number Equine Behaviour @ team @ Serial 4984  
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Author Smith, A.V.; Proops, L.; Grounds, K.; Wathan, J.; McComb, K. url  doi
openurl 
  Title Functionally relevant responses to human facial expressions of emotion in the domestic horse (Equus caballus) Type Journal Article
  Year 2016 Publication (up) Biology Letters Abbreviated Journal Biol. Lett.  
  Volume 12 Issue 2 Pages  
  Keywords  
  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.  
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  Notes Approved no  
  Call Number Equine Behaviour @ team @ Serial 6010  
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Author Wathan, J.; McComb, K. url  doi
openurl 
  Title The eyes and ears are visual indicators of attention in domestic horses Type Journal Article
  Year 2014 Publication (up) Current Biology Abbreviated Journal  
  Volume 24 Issue 15 Pages R677-R679  
  Keywords  
  Abstract Summary Sensitivity to the attentional states of others has adaptive advantages [1], and in social animals, attending to others is important for predator detection, as well as a pre-requisite for normal social functioning and more complex socio-cognitive abilities [2]. Despite widespread interest in how social species perceive attention in others, studies of non-human animals have been inconclusive about the detailed cues involved [3]. Previous work has focused on head and eye direction, overlooking the fact that many mammals have obvious and mobile ears that could act as a visual cue to attention. Here we report that horses use the head orientation of a conspecific to locate food, but that this ability is disrupted when parts of the face (the eyes and ears) are covered up with naturalistic masks. The ability to correctly judge attention also interacted with the identity of the model horse, suggesting that individual differences in facial features may influence the salience of cues. Our results indicate that a combination of head orientation with facial expression, specifically involving both the eyes and ears, is necessary for communicating social attention. These findings emphasise that in order to understand how attention is communicated in non-human animals, it is essential to consider a broad range of cues.  
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  Series Editor Series Title Abbreviated Series Title  
  Series Volume Series Issue Edition  
  ISSN 0960-9822 ISBN Medium  
  Area Expedition Conference  
  Notes Approved no  
  Call Number Equine Behaviour @ team @ Serial 5929  
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Author Proops, L.; Grounds, K.; Smith, A.V.; McComb, K. url  doi
openurl 
  Title Animals Remember Previous Facial Expressions that Specific Humans Have Exhibited Type Journal Article
  Year 2018 Publication (up) Current Biology Abbreviated Journal  
  Volume 28 Issue 9 Pages 1428-1432.e4  
  Keywords affective processing; face processing; ; animal-human interaction; interspecific communication; animal memory  
  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.  
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  Series Volume Series Issue Edition  
  ISSN 0960-9822 ISBN Medium  
  Area Expedition Conference  
  Notes Approved no  
  Call Number Equine Behaviour @ team @ Serial 6394  
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Author Proops, L.; McComb, K.; Reby, D. pdf  openurl
  Title Cross-modal individual vocal recognition in the domestic horse Type Conference Article
  Year 2008 Publication (up) IESM 2008 Abbreviated Journal  
  Volume Issue Pages  
  Keywords social cognition, animal-human interaction, horses, attention  
  Abstract Horses fulfill many of the criteria for a species in which it would be adaptive to be capable of individual recognition: they are highly social, form strong and long lasting bonds, their affiliations are rarely kin based, they have a fission-fusion social structure and they possess inter and intra-group dominance hierarchies.

We used a novel cross-modal, expectancy violation paradigm to provide the first systematic evidence that a non-human animal – the domestic horse- is capable of cross modal recognition. We believe this paradigm could provide an ideal way to study individual recognition across a wide range of species.

For full published details see: Proops L, McComb K, Reby D (2009) Cross-modal individual recognition in domestic horses (Equus caballus). Proc Natl Acad Sci U S A 106: 947-951.
 
  Address Centre for Mammal Vocal Communication Research, Psychology department,  
  Corporate Author Proops, L Thesis  
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  Area Expedition Conference IESM 2008  
  Notes Talk 15 min IESM 2008 Approved yes  
  Call Number Equine Behaviour @ team @ Serial 4469  
Permanent link to this record
 

 
Author Proops, L.; McComb, K.; Reby, D. pdf  openurl
  Title Horse-human interactions: Attention attribution and the use of human cues by domestic horses (Equus caballus). Type Conference Article
  Year 2008 Publication (up) IESM 2008 Abbreviated Journal  
  Volume Issue Pages  
  Keywords social cognition; animal-human interaction; horses; attention  
  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.
 
  Address  
  Corporate Author Proops, L. Thesis  
  Publisher Place of Publication Editor  
  Language Summary Language Original Title  
  Series Editor Series Title Abbreviated Series Title  
  Series Volume Series Issue Edition  
  ISSN ISBN Medium  
  Area Expedition Conference IESM 2008  
  Notes Poster IESM 2008 Approved no  
  Call Number Equine Behaviour @ team @ Serial 4502  
Permanent link to this record
 

 
Author Wathan, J.; Burrows, A.M.; Waller, B.M.; McComb, K. url  doi
openurl 
  Title EquiFACS: The Equine Facial Action Coding System Type Journal Article
  Year 2015 Publication (up) PLoS ONE Abbreviated Journal PLoS ONE  
  Volume 10 Issue 8 Pages e0131738  
  Keywords  
  Abstract <p>Although previous studies of horses have investigated their facial expressions in specific contexts, e.g. pain, until now there has been no methodology available that documents all the possible facial movements of the horse and provides a way to record all potential facial configurations. This is essential for an objective description of horse facial expressions across a range of contexts that reflect different emotional states. Facial Action Coding Systems (FACS) provide a systematic methodology of identifying and coding facial expressions on the basis of underlying facial musculature and muscle movement. FACS are anatomically based and document all possible facial movements rather than a configuration of movements associated with a particular situation. Consequently, FACS can be applied as a tool for a wide range of research questions. We developed FACS for the domestic horse (<italic>Equus caballus</italic>) through anatomical investigation of the underlying musculature and subsequent analysis of naturally occurring behaviour captured on high quality video. Discrete facial movements were identified and described in terms of the underlying muscle contractions, in correspondence with previous FACS systems. The reliability of others to be able to learn this system (EquiFACS) and consistently code behavioural sequences was high?and this included people with no previous experience of horses. A wide range of facial movements were identified, including many that are also seen in primates and other domestic animals (dogs and cats). EquiFACS provides a method that can now be used to document the facial movements associated with different social contexts and thus to address questions relevant to understanding social cognition and comparative psychology, as well as informing current veterinary and animal welfare practices.</p>  
  Address  
  Corporate Author Thesis  
  Publisher Public Library of Science Place of Publication Editor  
  Language Summary Language Original Title  
  Series Editor Series Title Abbreviated Series Title  
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  ISSN ISBN Medium  
  Area Expedition Conference  
  Notes Approved no  
  Call Number Equine Behaviour @ team @ Serial 5973  
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Author Proops, L.; Rayner, J.; Taylor, A.M.; McComb, K. url  doi
openurl 
  Title The Responses of Young Domestic Horses to Human-Given Cues Type Journal Article
  Year 2013 Publication (up) PLoS ONE Abbreviated Journal PLoS ONE  
  Volume 8 Issue 6 Pages e67000 EP -  
  Keywords  
  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>  
  Address  
  Corporate Author Thesis  
  Publisher Public Library of Science Place of Publication Editor  
  Language Summary Language Original Title  
  Series Editor Series Title Abbreviated Series Title  
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  ISSN ISBN Medium  
  Area Expedition Conference  
  Notes Approved no  
  Call Number Equine Behaviour @ team @ Serial 5676  
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