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Lampe, J. F., & Andre, J. (2012). Cross-modal recognition of human individuals in domestic horses (Equus caballus). Animal Cognition, 15(4), 623–630.
Abstract: This study has shown that domestic horses are capable of cross-modal recognition of familiar humans. It was demonstrated that horses are able to discriminate between the voices of a familiar and an unfamiliar human without seeing or smelling them at the same moment. Conversely, they were able to discriminate the same persons when only exposed to their visual and olfactory cues, without being stimulated by their voices. A cross-modal expectancy violation setup was employed; subjects were exposed both to trials with incongruent auditory and visual/olfactory identity cues and trials with congruent cues. It was found that subjects responded more quickly, longer and more often in incongruent trials, exhibiting heightened interest in unmatched cues of identity. This suggests that the equine brain is able to integrate multisensory identity cues from a familiar human into a person representation that allows the brain, when deprived of one or two senses, to maintain recognition of this person.
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Leliveld, L. M. C. (2019). From Science to Practice: A Review of Laterality Research on Ungulate Livestock. Symmetry, 11(9), 1157.
Abstract: In functional laterality research, most ungulate livestock species have until recently been mainly overlooked. However, there are many scientific and practical benefits of studying laterality in ungulate livestock. As social, precocial and domestic species, they may offer insight into the mechanisms involved in the ontogeny and phylogeny of functional laterality and help to better understand the role of laterality in animal welfare. Until now, most studies on ungulate livestock have focused on motor laterality, but interest in other lateralized functions, e.g., cognition and emotions, is growing. Increasingly more studies are also focused on associations with age, sex, personality, health, stress, production and performance. Although the full potential of research on laterality in ungulate livestock is not yet exploited, findings have already shed new light on central issues in cognitive and emotional processing and laid the basis for potentially useful applications in future practice, e.g., stress reduction during human-animal interactions and improved assessments of health, production and welfare. Future research would benefit from further integration of basic laterality methodology (e.g., testing for individual preferences) and applied ethological approaches (e.g., established emotionality tests), which would not only improve our understanding of functional laterality but also benefit the assessment of animal welfare.
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Maros, K., Boross, B., & Kubinyi, E. (2010). Approach and follow behaviour – possible indicators of the human-horse relationship. In Interaction Studies (Vol. 11, pp. 410–427).
Abstract: The aim of our study was to analyze the behavioural responses of horses (N = 51) to familiar humans and to find factors that may affect these responses in three tests: (1) approach to, (2) standing beside, and (3) following the familiar person. We investigated the impacts of horse-related factors (gender and age) and human-related factors (type of work, housing management, amount of handling, number of handlers and training to follow).<br xmlns=“http://pub2web.metastore.ingenta.com/ns/”></br> Horses with one handler needed less time to approach the human than horses with more handlers. Standing beside the human correlated positively with following. Following was mainly affected by training.<br xmlns=“http://pub2web.metastore.ingenta.com/ns/”></br> According to our results, the number of handlers has an important effect on horses' responses to familiar humans, especially regarding approach and follow behaviour. However, following behaviour is fundamentally determined by training.
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McGreevy, P., & Yeates, J. (2018). Horses (Equus caballus). In Companion Animal Care and Welfare. Companion Animal Care and Welfare.
Abstract: Summary Domestic horses are equid members of the class Mammalia, order Perissodactyla, and family Equidae. Horses are obligate herbivores, with nutritional requirements as listed in a table. Adequate space is necessary for exercise, exploration, flight, sharing resources, play, and rolling. Company is essential for all horses, including stallions. Company provides opportunities for mutual grooming and play and allows horses to stand head-to-tail to remove flies. Unhandled horses may respond to humans as they would to predators, whereas handled horses' responses depend on their previous interactions with humans. Horses can suffer from several diseases as listed in another table. The best method of euthanasia of horses is usually sedation followed by either cranial shooting or the injection of an overdose of pentobarbitone into the jugular vein. Behavioural signs of distress can include increased locomotory activity, vigilance behaviours, neighing, snorting, pawing, nibbling walls and buckets, defaecation, rearing, kicking stable walls or doors, and high-stepping 'prancing'.
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Merkies, K., Isensee, A., MacGregor, H., Koenig von Borstel, U., Tucker, A., Carson. J., et al. (2012). Influence of psychological and physiological arousal in humans on horse heart rate and behaviour. In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: The interaction of horses with humans is a dynamic state, but it is not clearly understood how horses perceive humans. Nervousness is transmissible from humans to horses indicated by increased horse heart rate (HR), however no studies have investigated whether horses can differentiate between humans who are physiologicallystressed (eg. after exercising) as opposed to psychologically-stressed (eg. feeling nervous/afraid). Horses (N=10) were randomly subjected to each of four treatments: 1) no human [control], 2) a calm human comfortable around horses [CALM; N=2 humans], 3) a physically-stressed human [PHYS; human exercised to reach 70% of maximum HR; N=2 humans], and 4) a psychologically-stressed human [PSYCH; human who was nervous around horses; N=14 humans]. Humans ranked themselves on a scale of 1-10 for their nervousness around horses. Both humans and horses were equipped with a HR monitor. Behavioural observations of the horses [gait, head position relative to the withers, distance from human, orientation toward human] were recorded live. Horses were allowed to wander loose in a round pen for 5 minutes of baseline recordings, at which time the human subject entered the round pen, stood in the centre and placed a blindfold over his/her eyes. The human remained in the centre of the round pen for an additional 5 minutes. Horse HR during control did not differ from when the human was present in the CALM and PSYCH treatment, and was lower during the PHYS treatment (51a vs 54a vs 55a vs 45b bpm for control, CALM, PSYCH and PHYS respectively; a,b differ p<0.0001). Over the 5 minute test period, horse HR decreased in PHYS and PSYCH (p<0.01) whereas it increased in CALM (p<0.0001). Horse HR decreased with increasing human rank of nervousness around horses (p=0.0156), and horses stood nearer to the human when they faced the human (p<0.0001) regardless of treatment. Horses moved at a faster gait in the control treatment, and their gait was slowest in the PSYCH treatment (p<0.0001), and the horse’s head position was lower in the PHYS and PSYCH treatments compared to CALM or baseline (p< 0.0001). A lower horse head position was positively correlated to a lower horse HR (p<0.0001) and negatively correlated to horse age (p<0.0001). Human HR was affected by treatment, with PHYS having the highest HR (p<0.0001). Human HR increased when the horse was facing away from the human, even though the human was blindfolded (p=0.0395). Overall, horses appear to be influenced by the physiological and psychological state of a human without any direct contact. Horses’ posture does reflect their physiological state. Understanding how horses react to human physiological and psychological states is especially important in equine-assisted activities, where the response of the horse has specific implications for the human participant.
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Munsters, C. C. B. M., Visser, K. E. K., van den Broek, J., & Sloet van Oldruitenborgh-Oosterbaan, M. M. (2012). The influence of challenging objects and horse-rider matching on heart rate, heart rate variability and behavioural score in riding horses. The Veterinary Journal, 192(1), 75–80.
Abstract: A good horse-rider ‘match’ is important in the context of equine welfare. To quantify the influence of repetition and horse-rider matching on the stress of horses encountering challenging objects, 16 Warmblood horses were ridden in a test-setting on three occasions. On each occasion the horse was ridden by a different rider and was challenged by three objects (A–C). Heart rate (HR), heart rate variability (HRV) of horse and rider, and behaviour score (BS) of the horse were obtained for each object and as a total for each test. The horse-rider interaction was evaluated with each combination and assessed as ‘matching’ or ‘mismatching’, and the horses were categorised as ‘compliant’, ‘partly-compliant’ or ‘non-compliant’. Horses exhibited a decreased HR (P = 0.015) and a decreased BS (P = 0.004) within and across different tests. ‘Matching’ horse-rider combinations exhibited less stress as indicated by reduced HR (‘match’ 69 ± 10 vs. ‘mismatch’ 72 ± 9, P = 0.001) and BS (‘match’ 1.9 ± 1.1 vs. ‘mismatch’ 3.8 ± 1.4, P = 0.017) of the horse. ‘Compliant’ (68 ± 8, P < 0.001) and ‘partly-compliant’ (71 ± 9, P = 0.002) horses had significantly lower HR than ‘non-compliant’ (75 ± 9) animals. The findings of the study indicate that HR and BS measurements support a subjective ‘match’ diagnosis and HR measurement may be a valuable tool in assessing horse compliance.
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Pierard, M. (2012). Agonistic and affiliative interactions in group housed riding horses (Equus caballus). In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: Group housed horses at a stud farm/riding stable in Belgium were observed on 17 days between 21 February and 25 April 2008, totalling 54hr25min of detailed data. The original group consisted of 8 Irish Cob mares, 1 Warmblood mare, 1 Arabian gelding and 2 Arabian mares. The group had been established in December 2007. During the course of the study 5 horses were removed from the group and 2 foals were born. 3 highly pregnant mares were housed adjacent to the group for part of the period. Horses were regularly used for lessons. Available surface area differed with the group on pasture at the end. Continuous all occurrence sampling of 10 agonistic and 2 affiliative behaviours was carried out for all group members present. Overall the group showed a frequency of 44.75 agonistic interactions per hour and 11.25 affiliative per hour. Of those agonistic interactions 46.3% were threats while 47% were less active interactions (displacement, being avoided), leaving only 6.7% more aggressive interactions ( mainly biting, some kicking and chasing). The effect on acting agonistically was not significant for age (p=0.1591) and borderline significant for density (p=0.0627). The analysis of the frequency of affiliative interactions showed there is no significant effect of age (p=0.1865) or density (p=0.7923). Agonistic and affiliative interactions were not significantly correlated (p=0.72). Affiliative behaviour a horse received showed a borderline effect (p=0.0787) on agonistic behaviour, as did the interaction between received agonistic and affiliative interactions (p=0.0725). Received agonistic interactions had a borderline negative effect (p=0.0656) on affiliative behaviour. A dominance hierarchy was calculated based on agonistic interactions using Empirical Bayes’ estimates based on Poisson regression with random effects. Agonistic behaviour expressed to other horses was significantly effected by relative rank (p=0.0243). Overall horses tended to be 3.7 times more aggressive towards lower ranking horses compared to higher ranking horses. Affiliative behaviour shown to other horses was not significantly influenced by the rank of the social partner (p=0.7915). Some individuals did show a significant effect whereby they showed more affiliative behaviour towards lower ranking individuals. This study was a small project to look at a practical situation of riding horses being kept in group housing. The frequent changes in group composition and available surface made it possible to look at agonistic and affiliative interactions in such circumstances. This is useful as instability in group composition is often used as main reason not to keep horses in social groups. The results from this study showed a borderline effect of density on agonistic behaviour. In reality it was also influenced by practical details, like a narrow paddock with only 2 hay crates on the smallest surface. Rank in a dominance hierarchy, based on agonistic behaviour, had a significant effect on the agonistic behaviour expressed towards higher or lower ranking horses. No injuries or escalating fights were observed. This study shows it is possible to keep a group of riding horses in a social context without excessive aggression.
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Proops, L., Grounds, K., Smith, A. V., & McComb, K. (2018). Animals Remember Previous Facial Expressions that Specific Humans Have Exhibited. Current Biology, 28(9), 1428–1432.e4.
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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Proops, L., McComb, K., & Reby, D. (2008). Cross-modal individual vocal recognition in the domestic horse. In IESM 2008.
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.
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Proops, L., McComb, K., & Reby, D. (2008). Horse-human interactions: Attention attribution and the use of human cues by domestic horses (Equus caballus). In IESM 2008.
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.
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