, Volume 191, Issue 3, pp 335–354 | Cite as

‘Theory of mind’ in animals: ways to make progress

  • Elske van der VaartEmail author
  • Charlotte K. Hemelrijk
Open Access


Whether any non-human animal can attribute mental states to others remains the subject of extensive debate. This despite the fact that several species have behaved as if they have a ‘theory of mind’ in various behavioral tasks. In this paper, we review the reasons of skeptics for their doubts: That existing experimental setups cannot distinguish between ‘mind readers’ and ‘behavior readers’, that results that seem to indicate ‘theory of mind’ may come from studies that are insufficiently controlled, and that our own intuitive biases may lead us to interpret behavior more ‘cognitively’ than is necessary. The merits of each claim and suggested solution are weighed. The conclusion is that while it is true that existing setups cannot conclusively demonstrate ‘theory of mind’ in non-human animals, focusing on this fact is unlikely to be productive. Instead, the more interesting question is how sophisticated their social reasoning can be, whether it is about ‘unobservable inner experiences’ or not. Therefore, it is important to address concerns about the setup and interpretation of specific experiments. To alleviate the impact of intuitive biases, various strategies have been proposed in the literature. These include a deeper understanding of associative learning, a better knowledge of the limited ‘theory of mind’ humans actually use, and thinking of animal cognition in an embodied, embedded way; that is, being aware that constraints outside of the brain, and outside of the body, may naturally predispose individuals to produce behavior that looks smart without requiring complex cognition. To enable this kind of thinking, a powerful methodological tool is advocated: Computational modeling, namely agent-based modeling and, particularly, cognitive modeling. By explicitly simulating the rules and representations that underlie animal performance on specific tasks, it becomes much easier to look past one’s own biases and to see what cognitive processes might actually be occurring.


Theory of mind Animal cognition Computational modeling 



This research was supported by the Netherlands Organisation for Scientific Research, through TopTalent grant 021.001.089, Open Access grant 036.002.118 and Vici grant 277-80-001.

Open Access

This article is distributed under the terms of the Creative Commons Attribution License which permits any use, distribution, and reproduction in any medium, provided the original author(s) and the source are credited.


  1. Anderson J.R. (2007) How can the human mind occur in the physical universe?. Oxford University Press, OxfordCrossRefGoogle Scholar
  2. Andrews K. (2005) Chimpanzee theory of mind: Looking in all the wrong places?. Mind and Language 20: 521–536CrossRefGoogle Scholar
  3. Andrews K. (2008) It’s in your nature: A pluralistic folk psychology. Synthese 165: 13–29CrossRefGoogle Scholar
  4. Apperly I.A., Butterfill S.A. (2009) Do humans have two systems to track beliefs and belief-like states?. Psychological Review 116: 953–970CrossRefGoogle Scholar
  5. Balda R.P., Kamil A.C., Grim K. (1986) Revisits to emptied cache sites by Clark’s nutcrackers (Nucifraga columbiana). Animal Behaviour 34: 1289–1298CrossRefGoogle Scholar
  6. Barrett L. (2010) Too much monkey business. In: Semin G., Echterhoff G. (Eds.) Grounding sociality. Psychology Press, London, pp 219–236Google Scholar
  7. Barrett L., Henzi P., Rendall D. (2007) Social brains, simple minds: Does social complexity really require cognitive complexity?. Philosophical Transactions of the Royal Society B 362: 561–575CrossRefGoogle Scholar
  8. Bolhuis J.J., Wynne C.D.L. (2009) Can evolution explain how minds work?. Nature 458: 832–833CrossRefGoogle Scholar
  9. Bräuer J., Call J., Tomasello M. (2007) Chimpanzees really know what others can see in a competitive situation. Animal Cognition 10: 439–448CrossRefGoogle Scholar
  10. Bryson J.J., Leong J.C.S. (2007) Primate errors in transitive ’inference’: A two-tier learning model. Animal Cognition 10: 1–15CrossRefGoogle Scholar
  11. Bulloch M.J., Boysen S.T., Furlong E.E. (2008) Visual attention and its relation to knowledge states in chimpanzees, Pan troglodytes. Animal Behaviour 76: 1147–1155CrossRefGoogle Scholar
  12. Byrne R.W., Bates L.A. (2010) Primate social cognition: Uniquely primate, uniquely social, or just unique?. Neuron 65: 815–830CrossRefGoogle Scholar
  13. Call J., Tomasello M. (2008) Does the chimpanzee have a theory of mind? 30 years later. Trends in Cognitive Sciences 12: 187–192CrossRefGoogle Scholar
  14. Camazine, S., Deneubourg, J.-L., Franks, N. R., Sneyd, J., Theraulaz, G., & Bonabeau, E. (2001). Self-organization in biological systems. Princeton: Princeton University Press.Google Scholar
  15. Clary D., Kelly D.M. (2011) Cache protection strategies of a non-social food-caching corvid, Clark’s nutcracker (Nucifrage columbiana). Animal Cognition 14: 735–744CrossRefGoogle Scholar
  16. Clayton N.S., Dally J.M., Emery N.J. (2007) Social cognition by food-caching corvids. The western scrub-jay as a natural psychologist. Philosophical Transactions of the Royal Society B 362: 505–522CrossRefGoogle Scholar
  17. Couzin I.D., Krause J. (2003) Self-organization and collective behavior in vertebrates. Advances in the Study of Behavior 32: 1–75CrossRefGoogle Scholar
  18. Dally J.M., Emery N.J., Clayton N.S. (2004) Cache protection strategies by western scrub-jays (Aphelocoma californica): Hiding food in the shade. Proceedings of the Royal Society B 271: 5387–5390CrossRefGoogle Scholar
  19. Dally J.M., Emery N.J., Clayton N.S. (2005) Cache protection strategies by western scrub-jays, Aphelocoma californica: Implications for social cognition. Animal Behaviour 70: 1251–1263CrossRefGoogle Scholar
  20. Dally J.M., Emery N.J., Clayton N.S. (2006) Food-caching western scrub jays keep track of who was watching when. Science 312: 1662–1666CrossRefGoogle Scholar
  21. de Kort S.R., Correia S.P.C., Alexis D.M., Dickinson A., Clayton N.S. (2007) The control of food-caching behavior by western scrub-jays (Aphelocoma californica). Journal of Experimental Psychology: Animal Behavior Processes 33: 361–370Google Scholar
  22. Dickinson A. (1980) Contemporary animal learning theory. Cambridge University Press, CambridgeGoogle Scholar
  23. Emery N.J. (2005) The evolution of social cognition. In: Easton A., Emery N.J. (Eds.) Cognitive neuroscience of social behaviour. Psychology Press, New YorkGoogle Scholar
  24. Emery N.J., Clayton N.S. (2001) Effects of experience and social context on prospective caching strategies by scrub jays. Nature 414: 443–446CrossRefGoogle Scholar
  25. Emery N.J., Clayton N.S. (2004) The mentality of crows: Convergent evolution of intelligence in corvids and apes. Science 306: 1903–1907CrossRefGoogle Scholar
  26. Emery N.J., Clayton N.S. (2008) How to build a scrub jay that reads minds. In: Itakura S., Fujita K. (Eds.) Origins of the social mind: Evolutionary and developmental perspectives. Springer, Tokyo, pp 65–98CrossRefGoogle Scholar
  27. Emery N.J., Clayton N.S. (2009) Comparative social cognition. Annual Review of Psychology 60: 87–113CrossRefGoogle Scholar
  28. Emery N.J., Dally J.M., Clayton N.S. (2004) Western scrub-jays (Aphelocoma californica) use cognitive strategies to protect their caches from thieving conspecifics. Animal Cognition 7: 37–43CrossRefGoogle Scholar
  29. Evers E., de Vries H., Spruijt B.M., Sterck E.H.M. (2011) Better safe than sorry - Socio-spatial group structure emerges from individual variation in fleeing, avoidance or velocity in an agent-based model. PLoS ONE 6: e26189. doi: 26110.21371/journal.pone.0026189 CrossRefGoogle Scholar
  30. Flombaum J.I., Santos L.R. (2005) Rhesus monkeys attribute perceptions to others. Current Biology 15: 447–452CrossRefGoogle Scholar
  31. Hare B. (2001) Can competitive paradigms increase the validity of experiments on primate social cognition?. Animal Cognition 4: 269–280CrossRefGoogle Scholar
  32. Hare B., Call J., Agnetta B., Tomasello M. (2000) Chimpanzees know what conspecifics do and do not see. Animal Behaviour 59: 771–785CrossRefGoogle Scholar
  33. Hare B., Call J., Tomasello M. (2001) Do chimpanzees know what conspecifics know?. Animal Behaviour 61: 139–151CrossRefGoogle Scholar
  34. Harrison, A., & Trafton, J.G. (2009). Gaze-following and awareness of visual perspective in chimpanzees. In: Howes A., Peebles D., Cooper R. (Eds.), Proceedings of the Ninth International Conference on Cognitive Modeling, (pp. 292–297), Manchester.Google Scholar
  35. Hemelrijk C. K. (1996) Reciprocation in great apes: From complex cognition to self-structuring. In: W.C. McGrew, M. L.F., & T. Nishida (Eds.), Great Ape societies. Cambridge: Cambridge University Press.Google Scholar
  36. Hemelrijk C.K. (2002) Understanding of social behaviour with the help of complexity science. Ethology 108: 655–671CrossRefGoogle Scholar
  37. Hemelrijk C.K., Bolhuis J.J. (2011) A minimalist approach to comparative psychology. Trends in Cognitive Science 15: 185–186CrossRefGoogle Scholar
  38. Hemelrijk C.K., Hildenbrandt H. (2011) Some causes of the variable shape of flocks of birds. PLoS ONE 6: e22479. doi: 22410.21371/journal.pone.0022479 CrossRefGoogle Scholar
  39. Heyes C.M. (1993) Anecdotes, training, trapping and triangulating: Do animals attribute mental states?. Animal Behaviour 46: 177–188CrossRefGoogle Scholar
  40. Heyes C.M. (1998) Theory of mind in nonhuman primates. Behavioral and Brain Sciences 21: 101–148Google Scholar
  41. Heyes C.M. (2012) A qualified defense of associative learning. Philosophical Transactions of the Royal Society B 367: 2695–2703CrossRefGoogle Scholar
  42. Hofstetter A.B., Russell J.L., Freeman H., Hopkins W.D. (2007) Now you see me, now you don’t: Evidence that chimpanzees understand the role of the eyes in attention. Animal Cognition 10: 55–62CrossRefGoogle Scholar
  43. Horner V., Carter J.D., Suchak M., de Waal F.B.M. (2011) Spontaneous prosocial choice by chimpanzees. Proceedings of the National Academy of Sciences of the United States of America 108: 13867–13851Google Scholar
  44. Hurley S., Nudds M. (2006) The question of animal rationality. In: Hurley S., Nudds M. (Eds.) Rational animals?. Oxford University Press, Oxford, pp 1–83CrossRefGoogle Scholar
  45. Kamil A.C., Balda R.P. (1990) Differential memory for different cache sites by Clark’s nutcrackers (Nucifraga columbiana). Journal of Experimental Psychology: Animal Behavior Processes 16: 162–168Google Scholar
  46. Kaminski J., Call J., Tomasello M. (2004) Body orientation and face orientation: Two factors controlling apes’ begging behavior from humans. Animal Cognition 7: 216–223CrossRefGoogle Scholar
  47. Kaminski J., Call J., Tomasello M. (2008) Chimpanzees know what others know, but not what they believe. Cognition 109: 224–234CrossRefGoogle Scholar
  48. Karin-D’Arcy M.R., Povinelli D.J. (2002) Do chimpanzees know what each other see? A closer look. International Journal of Comparative Psychology 110: 21–54Google Scholar
  49. Keysar B., Barr D.J., Balin J.A., Brauner J.S. (2000) Taking perspective in conversation: The role of mutual knowledge in comprehension. Pyschological Sciences 11: 32–38CrossRefGoogle Scholar
  50. King A.J., Sueur C., Huchard E., Cowlishaw G. (2011) A rule-of-thumb based on social affiliation explains collective movement in desert baboons. Animal Behaviour 82: 1337–1345CrossRefGoogle Scholar
  51. Krachun C., Call J., Tomasello M. (2009) Can chimpanzees (Pan troglodytes) discriminate appearance from reality?. Cognition 112: 435–450CrossRefGoogle Scholar
  52. Kummer H., Dasser V., Hoyningen-Huene P. (1990) Exploring primate social cognition: Some critical remarks. Behaviour 112: 84–98CrossRefGoogle Scholar
  53. Lucas J.R., Freeberg T.M., Egbert J., Schwabl H. (2006) Fecal corticosterone, body mass, and caching rates of Carolina chickadees (Poecili carolinensis) from disturbed and undisturbed sites. Hormones and Behavior 49: 634–643CrossRefGoogle Scholar
  54. Lurz R. (2009) If chimpanzees are mindreaders, could behavioral science tell? Towards a solution of the logical problem. Philosophical Psychology 22: 305–328CrossRefGoogle Scholar
  55. Lurz R. (2011) Belief attribution in animals: On how to move forward conceptually and empirically. Review of Philosophy and Psychology 22: 19–59CrossRefGoogle Scholar
  56. Lurz R., Krachun C. (2011) How could we know whether nonhuman primates understand other’s internal goals and intentions? Solving Povinelli’s problem. Review of Philosophy and Psychology 2: 449–481CrossRefGoogle Scholar
  57. Lyons D.E., Santos L.R. (2006) Ecology, domain specificity, and the origins of theory of mind: Is competition the catalyst?. Philosophy Compass 1: 481–492CrossRefGoogle Scholar
  58. Onishi K.H., Baillargeon R. (2005) Do 15-month-old infants understand false beliefs?. Science 308: 255–258CrossRefGoogle Scholar
  59. Pearce J.M. (2008) Animal learning and cognition: An introduction. Psychology Press, LondonGoogle Scholar
  60. Penn D.C., Holyoak K.J., Povinelli D.J. (2008) Darwin’s mistake: Explaining the discontinuity between human and nonhuman minds. Behavioral and Brain Sciences 31: 109–178Google Scholar
  61. Penn D.C., Povinelli D.J. (2007) On the lack of evidence that non-human animals possess anything remotely resembling a ’theory of mind’. Philosophical Transactions of the Royal Society B 362: 731–744CrossRefGoogle Scholar
  62. Pfeifer R., Scheier C. (1999) Understanding intelligence. MIT Press, CambridgeGoogle Scholar
  63. Povinelli D.J., & Eddy T.J. (1996) What young chimpanzees know about seeing (Vol. 61, No. 3, Monographs of the Society for Research in Child Development). Chicago: University of Chicago Press.Google Scholar
  64. Povinelli D.J., Vonk J. (2003) Chimpanzee minds: Suspiciously human?. Trends in Cognitive Sciences 7: 157–160CrossRefGoogle Scholar
  65. Povinelli D.J., Vonk J. (2004) We don’t need a microscope to explore the chimpanzee’s mind. Mind and Language 19: 1–28CrossRefGoogle Scholar
  66. Pravosudov V.V. (2003) Long-term moderate elevation of corticosterone facilitates avian food-caching behaviour and enhances spatial memory. Proceedings of the Royal Society B 270: 2599–2604CrossRefGoogle Scholar
  67. Premack D. (2007) Human and animal cognition: Continuity and discontinuity. Proceedings of the National Academy of Sciences of the USA 104: 13861–13867CrossRefGoogle Scholar
  68. Puga-Gonzalez I., Hildenbrandt H., Hemelrijk C.K. (2009) Emergent patterns of social affiliation in primates: A model. PLoS Computational Biology 5: e1000630. doi: 1000610.1001371/journal.pcbi.1000630 CrossRefGoogle Scholar
  69. Reaux J.E., Theall L.A., Povinelli D.J. (1999) A longitudinal investigation of chimpanzees’ understanding of visual perception. Child Development 70: 275–290CrossRefGoogle Scholar
  70. Salvucci D.D. (2006) Modeling driver behavior in a cognitive architecture. Human Factors 48: 362–380CrossRefGoogle Scholar
  71. Seed A.M., Tomasello M. (2010) Primate cognition. Topics in Cognitive Science 2: 407–419CrossRefGoogle Scholar
  72. Shettleworth S.J. (2010) Clever animals and killjoy explanations in comparative pyschology. Trends in Cognitive Sciences 14: 477–481CrossRefGoogle Scholar
  73. Shettleworth S.J. (2010) Cognition, evolution and behavior. Oxford University Press, New YorkGoogle Scholar
  74. Singh D., Dixson B.J., Jessop T.S., Morgan B., Dixson A.F. (2010) Cross-cultural consensus for waist-hip ratio and women’s attractiveness. Evolution and Human Behavior 31: 176–181CrossRefGoogle Scholar
  75. Sueur C., Deneubourg J.-L. (2011) Self-organization in primates: Understanding the rules underlying collective movements. International Journal of Primatology 32: 1413–1432CrossRefGoogle Scholar
  76. Sun, R. (Ed.) (2008) The Cambridge handbook of computational psychology. Cambridge University Press, CambridgeGoogle Scholar
  77. Taatgen N.A., Anderson J.R. (2010) The past, present, and future of cognitive architectures. Topics in Cognitive Science 2: 693–704CrossRefGoogle Scholar
  78. Tempelmann S., Kaminski J., Liebal K. (2011) Focus on the essential: All great apes know when others are being attentive. Animal Cognition 14: 433–439CrossRefGoogle Scholar
  79. Tomasello M., Call J. (2006) Do chimpanzees know what others see—or only what they are looking at?. In: Hurley S., Nudds M. (Eds.) Rational animals. Oxford University Press, Oxford, pp 371–384CrossRefGoogle Scholar
  80. Tomasello M., Call J., Hare B. (2003) Chimpanzees understand psychological states—the question is which ones and to what extent. Trends in Cognitive Sciences 7: 153–156CrossRefGoogle Scholar
  81. Tomasello M., Call J., Hare B. (2003) Chimpanzees versus humans: It’s not that simple. Trends in Cognitive Sciences 7: 239–240CrossRefGoogle Scholar
  82. van der Vaart, Verbrugge R., Hemelrijk C.K. (2011) Corvid caching: Insights from a cognitive model. Journal of Experimental Psychology: Animal Behavior Processes 37: 330–340Google Scholar
  83. van der Vaart E., Verbrugge R., Hemelrijk C.K. (2012) Corvid re-caching without ’theory of mind’. PLoS ONE 7: e32904. doi: 32910.31371/journal.pone.0032904 CrossRefGoogle Scholar
  84. van Maanen L., van Rijn H., Taatgen N.A. (2012) RACE/A: An architectural account of the interactions between learning, task control, and retrieval dynamics. Cognitive Science 26: 62–101CrossRefGoogle Scholar
  85. Vonk J., Povinelli D.J. (2011) Preliminary investigations of cognitive plasticity: Social and physical causality in home-reared chimpanzees. In: Eilan N., Lerman H., Roessler J. (Eds.) Perception, causation and objectivity. Oxford University Press, Oxford, pp 342–367CrossRefGoogle Scholar
  86. Vonk J., Shackelford T.K. (2012) Toward bridging gaps: Finding commonality between evolutionary and comparative psychology. In: Vonk J., Shackelford T.K. (Eds.) Oxford handbook of comparative evolutionary psychology. Oxford University Press, New York, pp 3–16Google Scholar
  87. Wein J.M., Stephens D.W. (2011) Caching economics: Jays cache more when handling times are short and habitats are poor. Animal Behaviour 82: 579–585CrossRefGoogle Scholar

Copyright information

© The Author(s) 2012

Authors and Affiliations

  • Elske van der Vaart
    • 1
    • 2
    Email author
  • Charlotte K. Hemelrijk
    • 1
  1. 1.Behavioural Ecology and Self-Organisation, University of GroningenGroningenThe Netherlands
  2. 2.Institute of Artificial IntelligenceUniversity of GroningenGroningenThe Netherlands

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