Mammalian empathy: behavioural manifestations and neural basis

Journal name:
Nature Reviews Neuroscience
Year published:
Published online


Recent research on empathy in humans and other mammals seeks to dissociate emotional and cognitive empathy. These forms, however, remain interconnected in evolution, across species and at the level of neural mechanisms. New data have facilitated the development of empathy models such as the perception–action model (PAM) and mirror-neuron theories. According to the PAM, the emotional states of others are understood through personal, embodied representations that allow empathy and accuracy to increase based on the observer's past experiences. In this Review, we discuss the latest evidence from studies carried out across a wide range of species, including studies on yawn contagion, consolation, aid-giving and contagious physiological affect, and we summarize neuroscientific data on representations related to another's state.

At a glance


  1. The Russian-doll model of the evolution of empathy.
    Figure 1: The Russian-doll model of the evolution of empathy.

    Various components of the empathic response, which have been added layer upon layer during evolution, remain functionally integrated. At its core is the perception–action mechanism, which induces a similar emotional state in the observer as in the target. Its most basic expressions are motor mimicry and emotional contagion. The doll's outer layers, such as empathic concern and perspective-taking, build upon this core socio-affective basis while increasingly requiring emotion regulation, self–other distinction and cognition. Even though the doll's outer layers depend on prefrontal functioning, they remain fundamentally linked to the core perception–action mechanism. Adapted with permission from: de Waal, F. B. M. in Feelings & Emotions: The Amsterdam Symposium (eds Manstead T., Frijda, N. & Fischer A.) 379–399 (Cambridge Univ. Press, 2003).

  2. Behavioural manifestations of animal empathy.
    Figure 2: Behavioural manifestations of animal empathy.

    The table summarizes behavioural patterns that are considered to be expressions of empathy in non-human vertebrates, ranging from motor mirroring and yawn contagion to targeted helping. Most of these behaviours have been characterized by experimental research, whereas others have been documented observationally (see the references cited in the main text). The mechanisms and the non-human species in which these behaviours have been observed are shown. From top to bottom, the images show the following: two gelada baboon juveniles with playfaces (that is, they are showing rapid facial mimicry) (part a); a yawning wolf (part b); prairie vole mates matching each other's physiological stress level (part c); a juvenile bonobo wrapping her arms around another who has just lost a fight to provide consolation (part d); a rat that has learned to liberate another rat that was trapped in a container (part e); and an adult chimpanzee showing targeted helping by assisting a juvenile's descent from a tree (part f). The images in parts a–f are courtesy of Pier Francesco Ferrari, University of Parma, Italy; Teresa Romero, University of Lincoln, UK; Zack Johnson, Georgia Institute of Technology, USA; Zanna Clay, Durham University, UK; and Inbal Ben-Ami Bartal, University of California, Berkeley, USA.

  3. From affect transfer to altruism.
    Figure 3: From affect transfer to altruism.

    As shown in multiple animal studies, empathy may promote aid-giving behaviour between conspecifics6, 57, 58. First, the target's distress induces stress or distress in the observer through emotional transfer (step 1). The observer needs to downregulate its own distress in order to effectively attend to the target, such as through helping or consolation (step 2). The resulting reduction of the target's distress as a result of being helped is then transferred back to the observer, ameliorating the observer's caught distress (step 3); this reduction constitutes an intrinsic reward for performed altruism (step 4).

  4. Both cognitive and affective empathy access distributed, person-specific affective representations.
    Figure 4: Both cognitive and affective empathy access distributed, person-specific affective representations.

    Bottom-up, affective empathy (red box) occurs when an observer directly perceives the emotional state (such as a sad facial expression) of the target. This naturally activates distributed, personal representations of the target's state in the observer (purple box). These representations have developed over time with experience in the observer's life and include associated memories, semantic concepts, and bodily states and expressions. When empathy proceeds in a top-down, cognitive manner (blue box), the neural regions that support working memory, executive function, emotion regulation and visuospatial processes instead access the affective empathy representations from the top-down (indicated by the arrows). Thus, although the stimulation emerges from inside the mind rather than from the outside world, the affective regions and the associated representations are shared between cognitive and affective processes. Thus, subtractions of affective from cognitive forms of empathy reveal greater brain activity in the blue regions than in the red regions. However, the cognitive process must still access the affective regions and their shared associated representations (purple) to provide the imagination or simulation with content and meaning.

  5. Neural regions that participate in human empathy.
    Figure 5: Neural regions that participate in human empathy.

    The figure shows a medial sagittal view (far left), a left lateral side view (centre) and a coronal view (far right) of the human brain, and indicates the relative locations and roles of brain areas that are involved in human empathy. Affective representations are required to imagine how another person feels. Depending on the task, however, the observed neural activity will emphasize bottom-up, affective brain areas or top-down, cognitive ones. Regions that are more associated with affective empathy tasks — such as direct perception of the emotion or pain of another person — are shown in red: the anterior cingulate cortex (ACC) including perigenual and subgenual regions of the ACC, amygdala, thalamus, hypothalamus, primary motor cortex (M1), premotor cortex, the primary and secondary somatosensory cortices (S1 and S2), and the temporal pole (TP). Brain regions that are more associated with higher-level, top-down forms of empathy — such as imagining how you would feel in the place of another or taking their perspective — are shown in blue: the dorsolateral prefrontal cortex (DLPFC), inferior parietal lobule (IPL), temporoparietal junction (TPJ), superior temporal gyrus (STG) and fusiform gyrus (FG). Two regions that are commonly activated in both affective and cognitive tasks are shown in green: the anterior insula (AI) and the anterior middle cingulate cortex (aMCC), which extends dorsally into the supplementary motor area (SMA) and cingulate motor area (CMA). Neural locations are approximated so that the regions can be viewed in just three images (for example, the FG is normally too medial to be seen in the lateral image, the amygdala is shown in the medial view as emerging from behind the visible midline slice in right temporal cortex, and differences in laterality by region and task are not represented). IFG, inferior frontal gyrus; PI, posterior insula; VMPFC, ventromedial PFC.


  1. Preston, S. D. & de Waal, F. B. M. Empathy: its ultimate and proximate bases. Behav. Brain Sci. 25, 171 (2002).
    This paper offers the first summary of empathy as a perception–action process and reviews evidence from multiple levels of analysis.
  2. Goldman, A. I. Simulating Minds: the Philosophy, Psychology, and Neuroscience of Mindreading (Oxford Univ. Press, 2006).
  3. Baron-Cohen, S. in The Lost Self: Pathologies of the Brain and Identity (eds Feinberg, T. E. & Keenan, J. P.) 166180 (Oxford Univ. Press, 2005).
  4. Zahavi, D. Simulation, projection and empathy. Conscious. Cogn. 17, 514522 (2008).
  5. Lipps, T. Einfühlung, innere nachahmung und organenempfindungen [German]. Arch. Gesamte Psychol. 1, 465519 (1903).
  6. de Waal, F. B. M. Putting the altruism back into altruism: the evolution of empathy. Annu. Rev. Psychol. 59, 279300 (2008).
  7. Preston, S. D. & Hofelich, A. J. The many faces of empathy: parsing empathic phenomena through a proximate, dynamic-systems view of representing the other in the self. Emot. Rev. 4, 2433 (2012).
  8. Zahn-Waxler, C. & Radke-Yarrow, M. The origins of empathic concern. Motiv. Emot. 14, 107130 (1990).
  9. Walter, H. Social cognitive neuroscience of empathy: concepts, circuits, and genes. Emot. Rev. 4, 917 (2012).
  10. Darwin, C. The Descent of Man, and Selection in Relation to Sex (Princeton Univ. Press, 1982).
  11. Plutchik, R. in Empathy and its Development (eds Eisenberg, N. & Strayer, J.) 346 (Cambridge Univ. Press, 1987).
  12. de Waal, F. B. M. Good Natured (Harvard Univ. Press, 1996).
  13. Krupenye, C., Kano, F., Hirata, S., Call, J. & Tomasello, M. Great apes anticipate that other individuals will act according to false beliefs. Science 354, 110114 (2016).
  14. de Waal, F. B. M. The Age of Empathy (Harmony Books, 2009).
  15. Yamamoto, S., Humle, T. & Tanaka, M. Chimpanzees' flexible targeted helping based on an understanding of conspecifics' goals. Proc. Natl Acad. Sci. USA 109, 35883592 (2012).
  16. Hattori, Y., Leimgruber, K., Fujita, K. & de Waal, F. B. M. Food-related tolerance in capuchin monkeys (Cebus apella) varies with knowledge of the partner's previous food-consumption. Behaviour 149, 171185 (2012).
  17. Bateson, P. & Laland, K. N. Tinbergen's four questions: an appreciation and an update. Trends Ecol. Evol. 28, 712718 (2013).
  18. Tinbergen, N. On aims and methods of ethology [German]. Z. Tierpsychol. 20, 410433 (1963).
  19. Church, R. M. Emotional reactions of rats to the pain of others. J. Comp. Physiol. Psychol. 52, 132134 (1959).
    This is the first rodent study to analyse emotional responses to the distress of others.
  20. Rice, G. E. & Gainer, P. “Altruism” in the albino rat. J. Comp. Physiol. Psychol. 55, 123125 (1962).
  21. Masserman, J. H., Wechkin, S. & Terris, W. “Altruistic” behavior in rhesus monkeys. Am. J. Psychiatry 121, 584585 (1964).
  22. Wechkin, S., Masserman, J. H. & Terris, W. Shock to a conspecific as an aversive stimulus. Psychon. Sci. 1, 4748 (1964).
  23. Miller, R. E., Murphy, J. V. & Mirsky, I. A. Relevance of facial expression and posture as cues in communication of affect between monkeys. AMA Arch. Gen. Psychiatry 1, 480488 (1959).
  24. Yerkes, R. M. Almost Human (Century, 1925).
  25. Ladygina-Kohts, N. N. Infant Chimpanzee and Human Child (ed. de Waal, F. B. M.) (Oxford Univ. Press, 2001)
  26. de Waal, F. B. M. & van Roosmalen, A. Reconciliation and consolation among chimpanzees. Behav. Ecol. Sociobiol. 5, 5566 (1979).
    This is the first non-human animal study to define and analyse consolation behaviour.
  27. Zahn-Waxler, C., Radke-Yarrow, M., Wagner, E. & Chapman, M. Development of concern for others. Dev. Psychol. 28, 126136 (1992).
    This is a pioneering study of the early development of human consolation behaviour.
  28. Liddle, M. J. E., Bradley, B. S. & Mcgrath, A. Baby empathy: infant distress and peer prosocial responses. Infant Ment. Health J. 36, 446458 (2015).
  29. Roth-Hanania, R., Davidov, M. & Zahn-Waxler, C. Empathy development from 8 to 16 months: early signs of concern for others. Infant Behav. Dev. 34, 447458 (2011).
  30. Dimberg, U., Thunberg, M. & Elmehed, K. Unconscious facial reactions to emotional facial expressions. Psychol. Sci. 11, 8689 (2000).
    This study is part of a series of studies on facial mimicry that first demonstrated the spontaneous nature of human empathic reactions.
  31. Mancini, G., Ferrari, P. F. & Palagi, E. Rapid facial mimicry in geladas. Sci. Rep. 3, 1527 (2013).
  32. Ross, M. D., Menzler, S. & Zimmermann, E. Rapid facial mimicry in orangutan play. Biol. Lett. 4, 2730 (2008).
  33. Dimberg, U., Andréasson, P. & Thunberg, M. Emotional empathy and facial reactions to facial expressions. J. Psychophysiol. 25, 2631 (2011).
  34. Campbell, M. W. & de Waal, F. B. M. Ingroup-outgroup bias in contagious yawning by chimpanzees supports link to empathy. PLoS ONE 6, e18283 (2011).
  35. Norscia, I. & Palagi, E. Yawn contagion and empathy in Homo sapiens. PLoS ONE 6, e28472 (2011).
  36. Romero, T., Konno, A. & Hasegawa, T. Familiarity bias and physiological responses in contagious yawning by dogs support link to empathy. PLoS ONE 8, e71365 (2013).
  37. Gallup, A. C., Swartwood, L., Militello, J. & Sackett, S. Experimental evidence of contagious yawning in budgerigars (Melopsittacus undulatus). Anim. Cogn. 18, 10511058 (2015).
  38. Silva, K., Bessa, J. & de Sousa, L. Auditory contagious yawning in domestic dogs (Canis familiaris): first evidence for social modulation. Anim. Cogn. 15, 721724 (2012).
  39. Visalberghi, E. & Fragaszy, D. M. in 'Language' and Intelligence in Monkeys and Apes: Comparative Developmental Perspectives (eds Parker, S. T. & Gibson, K. R.) 247273 (Cambridge Univ. Press, 1990).
  40. Ferrari, P. F. et al. Neonatal imitation in rhesus macaques. PLoS Biol. 4, e302 (2006).
  41. Paukner, A., Suomi, S. J., Visalberghi, E. & Ferrari, P. F. Capuchin monkeys display affiliation toward humans who imitate them. Science 325, 880883 (2009).
  42. Perry, S. et al. Social conventions in wild white-faced capuchin monkeys: evidence for traditions in a neotropical primate. Curr. Anthropol. 44, 241268 (2003).
  43. Dindo, M., Whiten, A. & de Waal, F. B. M. In-group conformity sustains different foraging traditions in capuchin monkeys (Cebus apella). PLoS ONE 4, e7858 (2009).
  44. Horner, V. & Whiten, A. Causal knowledge and imitation/emulation switching in chimpanzees (Pan troglodytes) and children (Homo sapiens). Anim. Cogn. 8, 164181 (2005).
  45. Fuhrmann, D., Ravignani, A., Marshall-Pescini, S. & Whiten, A. Synchrony and motor mimicking in chimpanzee observational learning. Sci. Rep. 4, 5283 (2014).
  46. Hopper, L. M., Lambeth, S. P., Schapiro, S. J. & Whiten, A. Observational learning in chimpanzees and children studied through 'ghost' conditions. Proc. R. Soc. B Biol. Sci. 275, 835840 (2008).
  47. Langford, D. J. et al. Social modulation of pain as evidence for empathy in mice. Science 312, 19671970 (2006).
    This paper provides compelling evidence of emotional contagion in mice and set the stage for subsequent rodent research.
  48. Langford, D. J. et al. Social approach to pain in laboratory mice. Soc. Neurosci. 5, 163170 (2010).
  49. Martin, L. J. et al. Reducing social stress elicits emotional contagion of pain in mouse and human strangers. Curr. Biol. 25, 326332 (2015).
  50. Burkett, J. et al. Oxytocin-dependent consolation behavior in rodents. Science 351, 375378 (2016).
  51. Buchanan, T. W., Bagley, S. L., Stansfield, R. B. & Preston, S. D. The empathic, physiological resonance of stress. Soc. Neurosci. 7, 191201 (2012).
    This paper provides the first demonstration that stress — at the physiological level — spreads from one stressed human participant to empathic observers.
  52. Buchanan, T. W. & Preston, S. D. Stress leads to prosocial action in immediate need situations. Front. Behav. Neurosci. (2014).
  53. Clay, Z. & de Waal, F. B. M. Development of socio-emotional competence in bonobos. Proc. Natl Acad. Sci. USA 110, 1812118126 (2013).
  54. Fries, A. B. W. & Pollak, S. D. Emotion understanding in postinstitutionalized Eastern European children. Dev. Psychopathol. 16, 355369 (2004).
  55. Kagan, J. Human morality is distinctive. J. Conscious. Stud. 7, 4648 (2000).
  56. Fehr, E. & Fischbacher, U. The nature of human altruism. Nature 425, 785791 (2003).
  57. Bartal, I. B.-A., Decety, J. & Mason, P. Empathy and pro-social behavior in rats. Science 334, 14271430 (2011).
  58. Sato, N., Tan, L., Tate, K. & Okada, M. Rats demonstrate helping behavior toward a soaked conspecific. Anim. Cogn. 18, 10391047 (2015).
  59. Romero, T., Castellanos, M. A. & de Waal, F. B. M. Consolation as possible expression of sympathetic concern among chimpanzees. Proc. Natl Acad. Sci. USA 107, 1211012115 (2010).
  60. Clay, Z. & de Waal, F. B. M. Bonobos respond to distress in others: consolation across the age spectrum. PLoS ONE 8, e55206 (2013).
  61. Palagi, E., Dall'Olio, S., Demuru, E. & Stanyon, R. Exploring the evolutionary foundations of empathy: consolation in monkeys. Evol. Hum. Behav. 35, 341349 (2014).
  62. Cools, A. K., Van Hout, A. J. M. & Nelissen, M. H. Canine reconciliation and third-party-initiated postconflict affiliation: do peacemaking social mechanisms in dogs rival those of higher primates? Ethology 114, 5363 (2008).
  63. Plotnik, J. M. & de Waal, F. B. M. Asian elephants (Elephas maximus) reassure others in distress. PeerJ 2, e278 (2014).
  64. Seed, A. M., Clayton, N. S. & Emery, N. J. Postconflict third-party affiliation in rooks, Corvus frugilegus. Curr. Biol. 17, 152158 (2007).
  65. Fraser, O. N., Stahl, D. & Aureli, F. Stress reduction through consolation in chimpanzees. Proc. Natl Acad. Sci. USA 105, 85578562 (2008).
  66. Aureli, F., Preston, S. D. & de Waal, F. B. M. Heart rate responses to social interactions in free-moving rhesus macaques (Macaca mulatta): a pilot study. J. Comp. Psychol. 113, 5965 (1999).
  67. Ben-Ami Bartal, I. et al. Anxiolytic treatment impairs helping behavior in rats. Front. Psychol. 7, 850 (2016).
  68. Warneken, F., Hare, B., Melis, A. P., Hanus, D. & Tomasello, M. Spontaneous altruism by chimpanzees and young children. PLoS Biol. 5, e184 (2007).
  69. Warneken, F. & Tomasello, M. Extrinsic rewards undermine altruistic tendencies in 20-month-olds. Motiv. Sci. 1, 4348 (2014).
  70. Andreoni, J. Giving with impure altruism: applications to charity and Ricardian equivalence. J. Polit. Econ. 97, 14471458 (1989).
  71. Wager, T. D. et al. Pain in the ACC? Proc. Natl Acad. Sci. USA 113, E2474E2475 (2016).
  72. Ballesta, S. & Duhamel, J.-R. Rudimentary empathy in macaques' social decision-making. Proc. Natl Acad. Sci. USA 112, 1551615521 (2015).
  73. Christov-Moore, L. & Iacoboni, M. Self–other resonance, its control and prosocial inclinations: brain–behavior relationships. Hum. Brain Mapp. 37, 15441558 (2016).
  74. Hoffman, M. L. Is altruism part of human nature? J. Pers. Soc. Psychol. 40, 121137 (1981).
  75. Nowbahari, E., Scohier, A., Durand, J.-L. & Hollis, K. L. Ants, Cataglyphis cursor, use precisely directed rescue behavior to free entrapped relatives. PLoS ONE 4, e6573 (2009).
  76. Vasconcelos, M., Hollis, K., Nowbahari, E. & Kacelnik, A. Pro-sociality without empathy. Biol. Lett. 8, 910912 (2012).
  77. Wilson, E. O. A chemical releaser of alarm and digging behavior in the ant Pogonomyrmex badius (Latreille). Psyche 65, 4151 (1958).
  78. Preston, S. D. The origins of altruism in offspring care. Psychol. Bull. 139, 13051341 (2013).
    This is a comprehensive review of the offspring-care perspective on how altruism and heroic rescue evolved and are encoded in the brain.
  79. MacLean, P. D. Brain evolution relating to family, play, and the separation call. Arch. Gen. Psychiatry 42, 405417 (1985).
  80. Sivaselvachandran, S., Acland, E. L., Abdallah, S. & Martin, L. J. Behavioral and mechanistic insight into rodent empathy. Neurosci. Biobehav. Rev. (2016).
  81. Edgar, J., Lowe, J., Paul, E. & Nicol, C. Avian maternal response to chick distress. Proc. R. Soc. B Biol. Sci. 278, 31293134 (2011).
  82. Doody, J. S., Burghardt, G. M. & Dinets, V. Breaking the social–non-social dichotomy: a role for reptiles in vertebrate social behavior research? Ethology 119, 95103 (2013).
  83. Donaldson, Z. R. & Young, L. J. Oxytocin, vasopressin, and the neurogenetics of sociality. Science 322, 900904 (2008).
  84. Chen, Q., Panksepp, J. B. & Lahvis, G. P. Empathy is moderated by genetic background in mice. PLoS ONE 4, e4387 (2009).
  85. Dittus, W. P. & Ratnayeke, S. M. Individual and social behavioral responses to injury in wild toque macaques (Macaca sinica). Int. J. Primatol. 10, 215234 (1989).
  86. De Vignemont, F. & Singer, T. The empathic brain: how, when and why? Trends Cogn. Sci. 10, 435441 (2006).
  87. de Waal, F. B. M. Chimpanzee Politics (John's Hopkins Univ. Press, 1998).
  88. Wilson, M. L. et al. Lethal aggression in Pan is better explained by adaptive strategies than human impacts. Nature 513, 414417 (2014).
  89. Lanzetta, J. T. & Englis, B. G. Expectations of cooperation and competition and their effects on observers' vicarious emotional responses. J. Pers. Soc. Psychol. 56, 543554 (1989).
  90. Prinz, W. Perception and action planning. Eur. J. Cogn. Psychol. 9, 129154 (1997).
  91. di Pellegrino, G., Fadiga, L., Fogassi, L., Gallese, V. & Rizzolatti, G. Understanding motor events: a neurophysiological study. Exp. Brain Res. 91, 176180 (1992).
    This study reports the discovery of mirror neurons, which are a subclass of cells that activate similarly when a macaque or the experimenter performs a goal-directed motor action.
  92. Grafton, S. T., Fadiga, L., Arbib, M. A. & Rizzolatti, G. Premotor cortex activation during observation and naming of familiar tools. Neuroimage 6, 231236 (1997).
  93. Adolphs, R., Damasio, H., Tranel, D., Cooper, G. & Damasio, A. R. A role for somatosensory cortices in the visual recognition of emotion as revealed by three-dimensional lesion mapping. J. Neurosci. 20, 26832690 (2000).
    This study offers early support for the PAM by showing that damage to the somatosensory cortex (where the body is represented) impairs an individual's ability to recognize the facial emotions of others.
  94. Jackson, P. L., Brunet, E., Meltzoff, A. N. & Decety, J. Empathy examined through the neural mechanisms involved in imagining how I feel versus how you feel pain. Neuropsychologia 44, 752761 (2006).
  95. Singer, T. & Lamm, C. The social neuroscience of empathy. Ann. NY Acad. Sci. 1156, 8196 (2009).
  96. Zaki, J. & Ochsner, K. N. The neuroscience of empathy: progress, pitfalls and promise. Nat. Neurosci. 15, 675680 (2012).
  97. Wicker, B. et al. Both of us disgusted in my insula: the common neural basis of seeing and feeling disgust. Neuron 40, 655664 (2003).
  98. Preston, S. D. et al. The neural substrates of cognitive empathy. Soc. Neurosci. 2, 254275 (2007).
  99. Morelli, S. A. & Lieberman, M. D. The role of automaticity and attention in neural processes underlying empathy for happiness, sadness, and anxiety. Front. Hum. Neurosci. 7, 160 (2013).
  100. Zaki, J., Wager, T. D., Singer, T., Keysers, C. & Gazzola, V. The anatomy of suffering: understanding the relationship between nociceptive and empathic pain. Trends Cogn. Sci. 20, 249259 (2016).
  101. Mohr, A. H., Kross, E. & Preston, S. D. Devil in the details: effects of depression on the prosocial response depend on timing and similarity. Adapt. Hum. Behav. Physiol. 2, 281297 (2016).
  102. Hodges, S. D., Kiel, K. J., Kramer, A. D., Veach, D. & Villanueva, B. R. Giving birth to empathy: the effects of similar experience on empathic accuracy, empathic concern, and perceived empathy. Pers. Soc. Psychol. Bull. 36, 398409 (2010).
  103. Völlm, B. A. et al. Neuronal correlates of theory of mind and empathy: a functional magnetic resonance imaging study in a nonverbal task. Neuroimage 29, 9098 (2006).
  104. Lamm, C., Nusbaum, H. C., Meltzoff, A. N. & Decety, J. What are you feeling? Using functional magnetic resonance imaging to assess the modulation of sensory and affective responses during empathy for pain. PLoS ONE 2, e1292 (2007).
  105. Nummenmaa, L., Hirvonen, J., Parkkola, R. & Hietanen, J. K. Is emotional contagion special? An fMRI study on neural systems for affective and cognitive empathy. Neuroimage 43, 571580 (2008).
  106. Schnell, K., Bluschke, S., Konradt, B. & Walter, H. Functional relations of empathy and mentalizing: an fMRI study on the neural basis of cognitive empathy. Neuroimage 54, 17431754 (2011).
  107. Blair, R. J. R. Responding to the emotions of others: dissociating forms of empathy through the study of typical and psychiatric populations. Conscious. Cogn. 14, 698718 (2005).
  108. Derntl, B., Seidel, E.-M., Schneider, F. & Habel, U. How specific are emotional deficits? A comparison of empathic abilities in schizophrenia, bipolar and depressed patients. Schizophr. Res. 142, 5864 (2012).
  109. Samson, D., Apperly, I. A., Chiavarino, C. & Humphreys, G. W. Left temporoparietal junction is necessary for representing someone else's belief. Nat. Neurosci. 7, 499500 (2004).
  110. Saxe, R. & Wexler, A. Making sense of another mind: the role of the right temporo-parietal junction. Neuropsychologia 43, 13911399 (2005).
  111. Adolphs, R., Baron-Cohen, S. & Tranel, D. Impaired recognition of social emotions following amygdala damage. J. Cogn. Neurosci. 14, 12641274 (2002).
  112. Blair, R. J. R. Fine cuts of empathy and the amygdala: dissociable deficits in psychopathy and autism. Q. J. Exp. Psychol. (Hove) 61, 157170 (2008).
  113. Marsh, A. A. Neural, cognitive, and evolutionary foundations of human altruism. Wiley Interdiscip. Rev. Cogn. Sci. 7, 5971 (2016).
  114. Singer, T. et al. Empathy for pain involves the affective but not sensory components of pain. Science 303, 11571162 (2004).
  115. Morrison, I., Lloyd, D., di Pellegrino, G. & Roberts, N. Vicarious responses to pain in anterior cingulate cortex: is empathy a multisensory issue? Cogn. Affect. Behav. Neurosci. 4, 270278 (2004).
  116. Jackson, P. L., Meltzoff, A. N. & Decety, J. How do we perceive the pain of others? A window into the neural processes involved in empathy. Neuroimage 24, 771779 (2005).
  117. Morrison, I. & Downing, P. E. Organization of felt and seen pain responses in anterior cingulate cortex. Neuroimage 37, 642651 (2007).
  118. Lamm, C., Decety, J. & Singer, T. Meta-analytic evidence for common and distinct neural networks associated with directly experienced pain and empathy for pain. Neuroimage 54, 24922502 (2011).
  119. Mischkowski, D., Crocker, J. & Way, B. M. From painkiller to empathy killer: acetaminophen (paracetamol) reduces empathy for pain. Soc. Cogn. Affect. Neurosci. 11, 13451353 (2016).
  120. Rütgen, M. et al. Placebo analgesia and its opioidergic regulation suggest that empathy for pain is grounded in self pain. Proc. Natl Acad. Sci. USA 112, E5638E5646 (2015).
  121. Rutgen, M., Seidel, E. M., Riecansky, I. & Lamm, C. Reduction of empathy for pain by placebo analgesia suggests functional equivalence of empathy and first-hand emotion experience. J. Neurosci. 35, 89388947 (2015).
  122. Danziger, N., Prkachin, K. M. & Willer, J.-C. Is pain the price of empathy? The perception of others' pain in patients with congenital insensitivity to pain. Brain 129, 24942507 (2006).
  123. Lieberman, M. D. & Eisenberger, N. I. The dorsal anterior cingulate cortex is selective for pain: results from large-scale reverse inference. Proc. Natl Acad. Sci. USA 112, 1525015255 (2015).
  124. Krishnan, A. et al. Somatic and vicarious pain are represented by dissociable multivariate brain patterns. eLife 5, e15166 (2016).
  125. Fan, Y., Duncan, N. W., de Greck, M. & Northoff, G. Is there a core neural network in empathy? An fMRI based quantitative meta-analysis. Neurosci. Biobehav. Rev. 35, 903911 (2011).
    This paper offers a meta-analysis of the role of the dACC–aMCC–SMA and the bilateral anterior insula across empathy tasks, demonstrating a domain-general role with similar recruitment for self and other experiences.
  126. Corradi-Dell'Acqua, C., Tusche, A., Vuilleumier, P. & Singer, T. Cross-modal representations of first-hand and vicarious pain, disgust and fairness in insular and cingulate cortex. Nat. Commun. 7, 10904 (2016).
    In this study, the anterior insula and aMCC are engaged across pain, disgust and unfairness, showing that these regions are not pain specific but participate in various negative self and other experiences.
  127. Danziger, N., Faillenot, I. & Peyron, R. Can we share a pain we never felt? Neural correlates of empathy in patients with congenital insensitivity to pain. Neuron 61, 203212 (2008).
  128. Craig, A. D. How do you feel? Interoception: the sense of the physiological condition of the body. Nat. Rev. Neurosci. 3, 655666 (2002).
  129. de la Vega, A., Chang, L. J., Banich, M. T., Wager, T. D. & Yarkoni, T. Large-scale meta-analysis of human medial frontal cortex reveals tripartite functional organization. J. Neurosci. 36, 65536562 (2016).
  130. Shackman, A. J. et al. The integration of negative affect, pain and cognitive control in the cingulate cortex. Nat. Rev. Neurosci. 12, 154167 (2011).
  131. Morrison, I., Peelen, M. V. & Downing, P. E. The sight of others' pain modulates motor processing in human cingulate cortex. Cereb. Cortex 17, 22142222 (2007).
  132. Carr, L., Iacoboni, M., Dubeau, M. C., Mazziotta, J. C. & Lenzi, G. L. Neural mechanisms of empathy in humans: a relay from neural systems for imitation to limbic areas. Proc. Natl Acad. Sci. USA 100, 54975502 (2003).
  133. Calder, A. J., Keane, J., Manes, F., Antoun, N. & Young, A. W. Impaired recognition and experience of disgust following brain injury. Nat. Neurosci. 3, 10771078 (2000).
  134. Bechara, A., Damasio, H. & Damasio, A. R. Emotion, decision making and the orbitofrontal cortex. Cereb. Cortex 10, 295307 (2000).
    This paper shows how the orbitofrontal cortex informs decisions from one's own simulated affective state, allowing humans to imagine how another person feels and how they would feel as a result of possible decision outcomes.
  135. Gallese, V., Keysers, C. & Rizzolatti, G. A unifying view of the basis of social cognition. Trends Cogn. Sci. 8, 396403 (2004).
  136. Iacoboni, M. Imitation, empathy, and mirror neurons. Annu. Rev. Psychol. 60, 653670 (2008).
  137. Mukamel, R., Ekstrom, A. D., Kaplan, J., Iacoboni, M. & Fried, I. Single-neuron responses in humans during execution and observation of actions. Curr. Biol. 20, 750756 (2010).
  138. Keysers, C. & Gazzola, V. Social neuroscience: mirror neurons recorded in humans. Curr. Biol. 20, R353R354 (2010).
  139. Shamay-Tsoory, S. G., Aharon-Peretz, J. & Perry, D. Two systems for empathy: a double dissociation between emotional and cognitive empathy in inferior frontal gyrus versus ventromedial prefrontal lesions. Brain 132, 617627 (2009).

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Author information

  1. Both authors contributed equally to this work.

    • Frans B. M. de Waal &
    • Stephanie D. Preston


  1. Psychology Department and Living Links, Yerkes National Primate Research Center, Emory University, 36 Eagle Row, Atlanta, Georgia 30322, USA.

    • Frans B. M. de Waal
  2. Department of Psychology, University of Michigan, 530 Church Street, Ann Arbor, Michigan 48109, USA.

    • Stephanie D. Preston

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The authors declare no competing interests.

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  • Frans B. M. de Waal

    Frans B. M. de Waal is a Dutch-American ethologist and biologist who has published widely about primate behaviour and social cognition. He received his Ph.D. in 1977 from the University of Utrecht, The Netherlands, and is presently C. H. Candler Professor at Emory University and the Director of Living Links at the Yerkes National Primate Research Center, Georgia, USA. He is also a distinguished professor at the University of Utrecht and is a member of the National Academy of Sciences.

  • Stephanie D. Preston

    Stephanie D. Preston is an interdisciplinary psychologist with a B.A. in cognitive science from the University of Virginia, USA, an M.A. and a Ph.D. in behavioural neuroscience from the University of California, Berkeley, USA, and a postdoctoral fellowship in neurology from the University of Iowa, USA. She is a professor at the University of Michigan, USA, where she studies the impact of implicit emotional processes on prosocial behaviour, hoarding, consumer behaviour and pro-environmentalism.

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