How do horses (Equus caballus) learn from observing human action?

  • Kira Bernauer
  • Hanna Kollross
  • Aurelia Schuetz
  • Kate Farmer
  • Konstanze KruegerEmail author
Original Paper


A previous study demonstrated that horses can learn socially from observing humans, but could not draw any conclusions about the social learning mechanisms. Here we develop this by showing horses four different human action sequences as demonstrations of how to press a button to open a feed box. We tested 68 horses aged between 3 and 12 years. 63 horses passed the habituation phase and were assigned either to the group Hand Demo (N = 13) for which a kneeling person used a hand to press the button, Head Demo (N = 13) for which a kneeling person used the head, Mixed Demo (N = 12) for which a squatting person used both head and hand, Foot Demo (N = 12) in which a standing person used a foot, or No Demo (N = 13) in which horses did not receive a demonstration. 44 horses reached the learning criterion of opening the feeder twenty times consecutively, 40 of these were 75% of the Demo group horses and four horses were 31% of the No Demo group horses. Horses not reaching the learning criterion approached the human experimenters more often than those who did. Significantly more horses used their head to press the button no matter which demonstration they received. However, in the Foot Demo group four horses consistently preferred to use a hoof and two switched between hoof and head use. After the Mixed Demo the horses’ actions were more diverse. The results indicate that only a few horses copy behaviours when learning socially from humans. A few may learn through observational conditioning, as some appeared to adapt to demonstrated actions in the course of reaching the learning criterion. Most horses learn socially through enhancement, using humans to learn where, and which aspect of a mechanism has to be manipulated, and by applying individual trial and error learning to reach their goal.


Copying Equus caballus Human demonstrator Interspecies specific social learning Social learning Social enhancement 



We thank Richard Byrne, Ádam Miklósi and many others for discussing the data, many persons for assisting in the experiments, several horse owners for supplying the experiment with their horses and three anonymous referees and the editor for helping to improve the manuscript.


This study was not funded.

Compliance with ethical standards

Ethical approval

All applicable international, national, and/or institutional guidelines for the care and use of animals were followed and approved by the animal welfare board of Nürtingen-Geislingen University. All procedures performed in the study involving human participants were in accordance with the ethical standards of the ethics committee of Nürtingen-Geislingen University and with the 1964 Helsinki declaration and its later amendments or comparable ethical standards.

Informed consent

Informed consent was obtained from all individual participants included in the study.

Supplementary material

10071_2019_1310_MOESM1_ESM.pdf (999 kb)
Source information for images all the images of the present manuscript (Fig. 1 and Fig. 2, main manuscript, and Fig. ESM 2, Fig ESM 3, Table ESM 1, Table ESM 4 and Table 5, supplementary material) were created from pictures, graphs and tables made by the authors of the present study themselves (PDF 999 kb)


  1. Blackmore TL, Foster TM, Sumpter CE, Temple W (2008) An investigation of colour discrimination with horses (Equus caballus). Behav Process 78:387–396. CrossRefGoogle Scholar
  2. Brubaker L, Udell MAR (2016) Cognition and learning in horses (Equus caballus): what we know and why we should ask more. Behav Process 126:121–131. CrossRefGoogle Scholar
  3. Byrne RW (1994) The evolution of intelligence. In: Slater PJB, Halliday TR (eds) Behaviour and evolution. Cambridge University Press, Cambridge, pp 223–265Google Scholar
  4. Byrne RW (2002) Imitation of novel complex actions: what does the evidence from animals mean? In: Snowdon CT, Roper TJ (eds) Advances in the study of behavior. Academic Press, San Diego, pp 77–105Google Scholar
  5. Byrne RW (2009) Animal imitation. Curr Biol 19:R111–R114. CrossRefGoogle Scholar
  6. Byrne R, Whiten A (1988) Machiavellian intelligence. Oxford Univ Press, OxfordGoogle Scholar
  7. Carroll J, Murphy CJ, Neitz M, Hoeve JN, Neitz J (2001) Photopigment basis for dichromatic color vision in the horse. J Vis 1:80–87. CrossRefGoogle Scholar
  8. Cook M, Mineka S, Wolkenstein B, Laitsch K (1985) Observational conditioning of snake fear in unrelated rhesus monkeys. J Abnorm Psychol 94:591–610CrossRefGoogle Scholar
  9. Custance D, Whiten A, Fredman T (1999) Social learning of an artificial fruit task in capuchin monkeys (Cebus apella). J Comp Psychol 113:13–23. CrossRefGoogle Scholar
  10. Deutsche Reiterliche Vereinigung e.V (2014) Pferde verstehen—Umgang und Bodenarbeit. FN Verlag der deutschen Reiterlichen Vereinigung GmbH, WarendorforeyGoogle Scholar
  11. d’Ingeo S, Quaranta A, Siniscalchi M, Stomp M, Coste C, Bagnard C et al (2019) Horses associate individual human voices with the valence of past interactions: a behavioural and electrophysiological study. Sci Rep 9:11568. CrossRefGoogle Scholar
  12. Dorey NR, Conover AM, Udell MAR (2014) Interspecific communication from people to horses (Equus ferus caballus) is influenced by different horsemanship training styles. J Comp Psychol 128(4):337–342. CrossRefGoogle Scholar
  13. Feh C, de Mazières J (1993) Grooming at a preferred site reduces heart rate in horses. Anim Behav 46:1191–1194. CrossRefGoogle Scholar
  14. Flannery B (1997) Relational discrimination learning in horses. Appl Anim Behav Sci 54:267–280. CrossRefGoogle Scholar
  15. Galef BG (1989) Enduring social enhancement of rats’ preferences for the palatable and the piquant. Appetite 13:81–92. CrossRefGoogle Scholar
  16. Greco BJ, Brown TK, Andrews JRM, Swaisgood RR, Caine NG (2013) Social learning in captive African elephants (Loxodonta africana africana). Anim Cogn 16:459–469. CrossRefGoogle Scholar
  17. Hall CA, Cassaday HJ, Derrington AM (2003) The effect of stimulus height on visual discrimination in horses. J Anim Sci 81(7):1715–1720CrossRefGoogle Scholar
  18. Healy SD, Jones CM (2002) Animal learning and memory: an integration of cognition and ecology. Zoology 105:321–327. CrossRefGoogle Scholar
  19. Heyes CM (1994) Social learning in animals: categories and mechanisms. Biol Rev 69:207–231. CrossRefGoogle Scholar
  20. Hopewell L, Leaver L, Lea S, Wills A (2010) Grey squirrels (Sciurus carolinensis) show a feature-negative effect specific to social learning. Anim Cogn 13:219–227. CrossRefGoogle Scholar
  21. Hoppitt W, Laland KN (2008) Social processes influencing learning in animals: a review of the evidence. Adv Study Behav 38:105–165. CrossRefGoogle Scholar
  22. Irving-Pease EK, Ryan H, Jamieson A, Dimopoulos EA, Larson G, Frantz LAF (2019) Paleogenomics of animal domestication. In: Lindqvist C, Rajora OP (eds) Paleogenomics: genome-scale analysis of ancient DNA. Springer International Publishing, Cham, pp 225–272Google Scholar
  23. Krueger K, Heinze J (2008) Horse sense: social status of horses (Equus caballus) affects their likelihood of copying other horses` behavior. Anim Cogn 11:431–439. CrossRefGoogle Scholar
  24. Krueger K, Flauger B, Farmer K, Maros K (2011) Horses (Equus caballus) use human local enhancement cues and adjust to human attention. Anim Cogn 14:187–201. CrossRefGoogle Scholar
  25. Krueger K, Farmer K, Heinze J (2014) The effects of age, rank and neophobia on social learning in horses. Anim Cogn 17:645–655. CrossRefGoogle Scholar
  26. Kubinyi E, Topál J, Miklósi Á, Csányi V (2003) Dogs (Canis familiaris) learn their owners via observation in a manipulation task. J Comp Psychol 117:156–165. CrossRefGoogle Scholar
  27. Lamp JF, Andre J (2012) Cross-modal recognition of human individuals in domestic horses (Equus caballus). Anim Cogn 15:623–630. CrossRefGoogle Scholar
  28. Leadbeater E, Dawson EH (2017) A social insect perspective on the evolution of social learning mechanisms. Proc Natl Acad Sci USA 114:7838–7845CrossRefGoogle Scholar
  29. Lesimple C, Sankey C, Richard M-A, Hausberger M (2012) Do horses expect humans to solve their problems? Front Psychol 3:306. CrossRefGoogle Scholar
  30. Lovrovich P, Sighieri C, Baragli P (2015) Following human-given cues or not? Horses (Equus caballus) get smarter and change strategy in a delayed three choice task. Appl Anim Behav Sci 166:80–88. CrossRefGoogle Scholar
  31. Malavasi R, Huber L (2016) Evidence of heterospecific referential communication from domestic horses (Equus caballus) to humans. Anim Cogn 19:899–909. CrossRefGoogle Scholar
  32. Maros K, Gácsi M, Miklósi Á (2008) Comprehension of human pointing gestures in horses (Equus caballus). Anim Cogn 11:457–466. CrossRefGoogle Scholar
  33. McDonnell SM, Haviland JCS (1995) Agonistic ethogram of the equid bachelor band. Appl Anim Behav Sci 43:147–188. CrossRefGoogle Scholar
  34. Mejdell CM, Buvik T, Jørgensen GHM, Bøe KE (2016) Horses can learn to use symbols to communicate their preferences. Appl Anim Behav Sci 184:66–73. CrossRefGoogle Scholar
  35. Mersmann D, Tomasello M, Call J, Kaminski J, Taborsky M (2011) Simple mechanisms can explain social learning in domestic dogs (Canis familiaris). Ethology 117:675–690. CrossRefGoogle Scholar
  36. Miklósi Á, Soproni K (2006) A comparative analysis of animals’ understanding of the human pointing gesture. Anim Cogn 9:81–93. CrossRefGoogle Scholar
  37. Mills DS, McDonnell SM (2005) The domestic horse. Cambridge University Press, CamebridgeGoogle Scholar
  38. Nicol CJ (2002) Equine learning: progress and suggestions for future research. Appl Anim Behav Sci 78:193–208. CrossRefGoogle Scholar
  39. Pfungst O (1907) Der Kluge Hans. Ein Beitrag zur nichtverbalen Kommunikation. Frankfurter Fachbuchhandlung für Psychologie, Frankfurt am MainGoogle Scholar
  40. Pongrácz P, Miklósi Á, Kubinyi E, Gurobi K, Topál J, Csányi V (2001) Social learning in dogs: the effect of a human demonstrator on the performance of dogs in a detour task. Anim Behav 62:1109–1117. CrossRefGoogle Scholar
  41. Pongrácz P, Miklósi Á, Timár-Geng K, Csányi V (2004) Verbal attention getting as a key factor in social learning between dog and human. J Comp Psychol 118:375–383CrossRefGoogle Scholar
  42. Pongrácz P, Miklósi Á, Vida V, Csányi V (2005) The pet dogs ability for learning from a human demonstrator in a detour task is independent from the breed and age. Appl Anim Behav Sci 90:309–323. CrossRefGoogle Scholar
  43. Proops L, McComb K (2010) Attributing attention: the use of human-given cues by domestic horses (Equus caballus). Anim Cogn 13:197–205. CrossRefGoogle Scholar
  44. Proops L, McComb K (2012) Cross-modal individual recognition in domestic horses (Equus caballus) extends to familiar humans. Proc R Soc B 279:3131–3138. CrossRefGoogle Scholar
  45. Proops L, McComb K, Reby D (2009) Cross-modal individual recognition in domestic horses (Equus caballus). Proc Natl Acad Sci USA 106:947–951. CrossRefGoogle Scholar
  46. Proops L, Walton M, McComb K (2010) The use of human-given cues by domestic horses, Equus caballus, during an object choice task. Anim Behav 79:1205–1209. CrossRefGoogle Scholar
  47. Proops L, Rayner J, Taylor AM, McComb K (2013) The responses of young domestic horses to human-given cues. PLoS One. Google Scholar
  48. R Development Core Team (2019) R: a language and environment for statistical computing.
  49. Rescorla RA, Holland PC (1982) Behavioral Studies of Associative Learning in Animals. Ann Rev Psychol 33:265–308CrossRefGoogle Scholar
  50. Ringhofer M, Yamamoto S (2017) Domestic horses send signals to humans when they face with an unsolvable task. Anim Cogn 20:397–405. CrossRefGoogle Scholar
  51. Rørvang MV, Ahrendt LP, Christensen JW (2015) A trained demonstrator has a calming effect on naïve horses when crossing a novel surface. Appl Anim Behav Sci 171:117–120. CrossRefGoogle Scholar
  52. Sankey C, Henry S, André N, Richard-Yris M-A, Hausberger M (2011) Do horses have a concept of person? PLoS One. Google Scholar
  53. Schuetz A, Farmer K, Krueger K (2017) Social learning across species: horses (Equus caballus) learn from humans by observation. Anim Cogn 20:567–573. CrossRefGoogle Scholar
  54. Shettleworth SJ (1998) Cognition, evolution and behaviour. Oxford University Press, OxfordGoogle Scholar
  55. Smith AV, Proops L, Grounds K, Wathan J, McComb K (2016) Functionally relevant responses to human facial expressions of emotion in the domestic horse (Equus caballus). Biol Lett. Google Scholar
  56. Thorpe WH (1963) Learning and instinct in animals. Methuen, LondonGoogle Scholar
  57. Tomasello M (1990) Cultural transmission in the tool use and communicatory signalling of chimpanzees? In: Parker ST, Gibson KR (eds) Language and intelligence in monkeys and apes. Cambridge University Press, Cambridge, pp 274–311CrossRefGoogle Scholar
  58. Topál J, Miklósi A, Csányi V (1997) Dog-human relationship affects problem solving be-havior in the dog. Anthrozoos 10:214–224CrossRefGoogle Scholar
  59. Whalen A, Cownden D, Laland K (2015) The learning of action sequences through social transmission. Anim Cogn 18:1093–1103. CrossRefGoogle Scholar
  60. Whiten A, Ham R (1992) On the nature and evolution of imitation in the animal kingdom: reappraisal of a century of research. Adv Study Behav 21:239–283CrossRefGoogle Scholar
  61. Whiten A, Horner V, Litchfield CA, Marshall-Pescini S (2004) How do apes ape? Learn Behav 32:36–52. CrossRefGoogle Scholar
  62. Zajonc RB (1965) Social facilitation. Science 149:269–274. CrossRefGoogle Scholar
  63. Zentall TR (2006) Imitation: definitions, evidence, and mechanisms. Anim Cogn 9:335–353. CrossRefGoogle Scholar

Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2019

Authors and Affiliations

  1. 1.Department Equine Economics, Faculty Agriculture, Economics and ManagementNuertingen-Geislingen UniversityNürtingenGermany
  2. 2.Department of Agricultural Economics and Rural Development, Faculty of Agricultural SciencesGeorg-August-University GoettingenGoettingenGermany
  3. 3.St Andrews University, School of Psychology and NeuroscienceSt Andrews, FifeUK
  4. 4.University of Regensburg, Zoology/Evolutionary BiologyRegensburgGermany

Personalised recommendations