Morality is an evolved repertoire of cognitive and emotional mechanisms with distinct biological underpinnings, as modified by experience acquired throughout the human lifespan.*

Morality is derived from both the evolved biochemistry of the brain and sociocultural influences. Displayed below are the neuroanatomical structures associated with morality (Pascual et al., 2013).

Is human morality a product of nature? Of nurture? There has been a long dispute in the social sciences about which is responsible. The debate has largely been unproductive. It seems that morality results from both nature and nurture.[1][2]

Nature already supplies many elements of humanity’s moral psychology. Research in the evolutionary behavioral sciences has accumulated strong evidence: the hardware of our brains is pre-programmed with propensities for moral judgment. We are ready to feel shame or exhibit cooperation.[3] We are already outfitted for empathy, altruism, and compassion.[4]

So, too, do we come readily equipped for what most people regard as immoral behavior. Because of our brains, humans are capable of violence. We are prepared to sometimes inflict pain.[5][6]

Humans are geared by nature toward learning ethical behavior from their social environments.[7] So, as we live, nurture is also at work. Upbringing by family members counts. Socialization by peers and communities about “right” or “wrong” is critical to shaping conduct.[8][9]

Reconciling these disparate parts of ourselves requires an understanding of humankind’s social and cognitive complexity. One thing is clear from the research: religious imperative is not necessary for morality.[10] Research shows that such moral sentiments encourage our caring for and helping others evolved independently of religion.[11][12]

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  • Abe, N., & Greene, J. D. (2014). Response to anticipated reward in the nucleus accumbens predicts behavior in an independent test of honesty. The Journal of Neuroscience, 34(32), 10564-10572.PDF
  • Bernhard, R. M., Chaponis, J., Siburian, R., Gallagher, P., Ransohoff, K., Wikler, D., ... & Greene, J. D. (2016). Variation in the oxytocin receptor gene (OXTR) is associated with differences in moral judgment. Social Cognitive and Affective Neuroscience, 11(12), 1872-1881.PDF
  • Burkart, J. M., Allon, O., Amici, F., Fichtel, C., Finkenwirth, C., Heschl, A., ... & Meulman, E. J. (2014). The evolutionary origin of human hyper-cooperation. Nature Communications, 5.PDF
  • Cesarini, D., Dawes, C. T., Fowler, J. H., Johannesson, M., Lichtenstein, P., & Wallace, B. (2008). Heritability of cooperative behavior in the trust game. Proceedings of the National Academy of Sciences USA, 105, 3721-3726. PDF
  • Chakroff, A., & Young, L. (2015). How the Mind Matters for Morality. AJOB Neuroscience, 6(3), 43-48.PDF
  • Cosmides, L. & Tooby, J. (2002). Knowing thyself: The evolutionary psychology of moral reasoning and moral sentiments.  Business, Science and Ethics, 91-127.
  • Cosmides, L., & Tooby, J. (2005). Neurocognitive adaptations designed for social exchange. In D. M. Buss (Ed.), Evolutionary Psychology Handbook. NY: Wiley. PDF
  • Curry, O. S. (2016). Morality as Cooperation - A Problem-Centred Approach. In The Evolution of Morality (pp. 27-51). Springer International Publishing.PDF
  • Curry, O. S., Rowland, L. A., Van Lissa, C. J., Zlotowitz, S., McAlaney, J., & Whitehouse, H. (2018). Happy to Help?: A systematic review and meta-analysis of othe effects of performing acts of kindness on the well-being of the actor. Journal of Experimental Social Psychology, 76, 320-329.PDF
  • Cushman, F., Kumar, V., & Railton, P. (2017). Moral learning: Current and future directions. Cognition, 167, 1-10.PDF
  • Cushman, F. (2015). Deconstructing intent to reconstruct morality. Current Opinion in Psychology, 6, 97-103.PDF
  • Cushman, F. A., & Greene, J. D. (2012). Finding faults: How moral dilemmas reveal cognitive structure. Social Neuroscience, 7(3), 269-279. PDF
  • Cushman, F. A., Young, L., & Greene, J. (2010). Our multi-system moral psychology: Towards a consensus view. In J. Doris et al. (Eds.), The Oxford Handbook of Moral Psychology. Oxford University Press. PDF
  • Darby, R. R., Horn, A., Cushman, F., & Fox, M. D. (2017). Lesion network localization of criminal behavior. Proceedings of the National Academy of Sciences, 201706587. PDF
  • DeQuervain, DJ-F., Fischbacher U., Treyer V., Schellhammer M., Schnyder U., Buck A., & Fehr E. (2004). The neural basis of altruistic punishment. Science, 305, 1254-1258.
  • DeScioli, P., & Kurzban, R. (2009). Mysteries of morality. Cognition, 112, 281-299. PDF
  • DeScioli, P., & Kurzban, R. (2013). A solution to the mysteries of morality. Psychological Bulletin, 139, 477-496. PDF
  • DeScioli, P. (2016). The side-taking hypothesis for moral judgment. Current Opinion in Psychology, 7, 23-27. PDF
  • Decety, J., & Chaminade, T. (2003). Neural correlates of feeling sympathy. Neuropsychologia - Special Issue on Social Cognition, 41(2), 127-138. PDF
  • Efferson, C., & Fehr, E. (2018). Simple moral code supports cooperation. Nature, 555 (7695), 169. PDF
  • Everett, J.A.C., Pizarro, D. A. & Crockett, M.J., (in press). Inference of trustworthiness from intuitive moral judgments. Journal of Experimental Psychology: General. PDF
  • Fehr, E. & Rockenbach, B. (2004). Human altruism: economic, neural, and evolutionary perspectives. Current Opinion in Neurobiology, 14, 784–790. PDF
  • Gervais, W. M., & Norenzayan, A. (2012) Like a camera in the sky? Thinking about God increases public self-awareness and socially desirable responding. Journal of Experimental Social Psychology, 48, 298-302. PDF
  • Graham, J., Nosek, B. A., Haidt, J., Iyer, R., Koleva, S., & Ditto, P. H. (2011). Mapping the moral domain. Journal of Personality and Social Psychology, 101(2), 366-385. PDF
  • Greene, J. D. (2017). The rat-a-gorical imperative: Moral intuition and the limits of affective learning. Cognition, 167, 66-77.PDF
  • Greene, J. D. (2015). The rise of moral cognition. Cognition, 135, 39-42.PDF
  • Greene, J. D., Nystrom, L. E., Engell, A. D., Darley, J. M. & Cohen, J. D. (2004). The neural bases of cognitive conflict and control in moral judgment. Neuron, 44, 389−400. PDF
  • Greene, J.D., Sommerville, R.B., Nystrom, L.E., Darley, J.M. & Cohen, J.D. (2001). An fMRI investigation of emotional engagement in moral judgment. Science, 293, 2105-2108. PDF
  • Heekeren, H. R., Wartenburger, I., Schmidt, H., Schwintowski, H. P. & Villringer, A. (2003). An fMRI study of simple ethical decision-making. Neuroreport, 14, 1215−1219.
  • Heiphetz, L., & Young, L. (2014). A social cognitive developmental perspective on moral judgment. Behaviour, 151(2-3). PDF
  • Holbrook, C., Fessler, D. M., & Pollack, J. (2016). With God on our side- Religious primes reduce the envisioned physical formidability of a menacing adversary. Cognition, 146, 387-392.PDF
  • Johnson, D. D., & MacKay, N. J. (2015). Fight the power- Lanchester's laws of combat in human evolution. Evolution and Human Behavior. 36(2), 152-163.PDF
  • Kiehl, K. A., Smith, A. M., Hare, R. D., Mendrek, A., Forster, B. B., Brink, J., & Liddle, P. F. (2001). Limbic abnormalities in affective processing by criminal psychopaths as revealed by functional magnetic resonance imaging. Biological Psychiatry50(9), 677-684. PDF
  • Koenigs, M., Young, L., Adolphs, R., Tranel, D., Hauser, M., Cushman, F., & Damasio A. (2007). Damage to the prefrontal cortex increases utilitarian moral judgments. Nature, 446, 908-911. PDF
  • Kosfeld, M., Heinrichs, M., Zak, P. J., Fischbacher, U., & Fehr, E. (2005). Oxytocin increases trust in humans. Nature, 435, 673-676. PDF
  • Margoni, F., & Surian, L. (2015). Explaining the U-shaped development of intent-based moral judgments. Frontiers in Psychology, 7.PDF
  • Martin, J. W., & Cushman, F. (2016). Why we forgive what can't be controlled. Cognition, 147, 133-143.PDF
  • McCabe K., Houser D., Ryan L., Smith V., & Trouard T. (2001). A functional imaging study of cooperation in two-person reciprocal exchange. Proceedings of the National Academy of Sciences USA, 98, 11832-11835. PDF
  • Moll, J., Eslinger, P. J. & Oliveira-Souza, R. (2001). Frontopolar and anterior temporal cortex activation in a moral judgment task: preliminary functional MRI results in normal subjects. Arq. Neuropsiquiatry, 59, 657−664. PDF
  • Moll, J., de Oliveira-Souza, R., Bramati, I. E. & Grafman, J. (2002). Functional networks in emotional moral and nonmoral social judgments. Neuroimage, 16, 696−703. PDF
  • Moll, J., de Oliveira-Souza, R., Eslinger, P. J., Bramati, I. E., Mourão-Miranda, J., Andreiuolo, P. A., & Pessoa, L. (2002). The neural correlates of moral sensitivity: a functional magnetic resonance imaging investigation of basic and moral emotions. The Journal of Neuroscience22(7), 2730-2736. PDF
  • Morris, A., MacGlashan, J., Littman, M. L., & Cushman, F. (2017). Evolution of flexibility and rigidity in retaliatory punishment. Proceedings of the National Academy of Sciences, 201704032. PDF
  • Müller, J. L., Sommer, M., Wagner, V., Lange, K., Taschler, H., Röder, C. H., Schuierer, G., Klein, H.E., & Hajak, G. (2003). Abnormalities in emotion processing within cortical and subcortical regions in criminal psychopaths: Evidence from a functional magnetic resonance imaging study using pictures with emotional content. Biological Psychiatry54(2), 152-162. PDF
  • Niemi, L., Wasserman, E., & Young, L. (2018). The behavioral and neural signatures of distinct conceptions of fairness. Social Neuroscience, 13(4), 399-415. PDF
  • Norenzayan, A. (2014). Does religion make people moral? Behaviour, 151, 365-384. PDF
  • Norenzayan, A., Henrich, J., & E. Slingerland (2013) Religious Prosociality: A Synthesis. In P. J. Richerson & M. H. Christiansen (Eds.), Cultural Evolution: Society, Technology, Language and Religion. Cambridge, Massachusetts: MIT Press. PDF
  • Park, S. Q., Kahnt, T., Dogan, A., Strang, S., Fehr, E., & Tobler, P. N. (2017). A neural link between generosity and happiness. Nature Communications, 8, 15964. PDF
  • Pascual, L., Rodrigues, P., & Gallardo-Pujol, D. (2013). How does morality work in the brain?: A functional and structural perspective of moral behaviors. Frontiers in Integrative Neuroscience, 7 (65). PDF
  • Patil, I., Calò, M., Fornasier, F., Young, L., & Silani, G. (2017). Neuroanatomical correlates of forgiving unintentional harms. Scientific Reports, 7, 45967. PDF
  • Pizarro, D.A., Inbar, Y., & Helion, C. (2011). On disgust and moral judgment. Emotion Review, 3, 267–268. PDF
  • Pizarro, D.A., Tannenbaum, D., & Uhlmann, E.L. (2012). Mindless, harmless, and blameworthy. Psychological Inquiry, 23, 185-188. PDF
  • Pollack, J., Holbrook, C., Fessler, D. M., Sparks, A. M., & Zerbe, J. G. (2018). May God guide our guns. Human Nature, 1-17. PDF
  • Raine, A., Lencz, T., Bihrle, S., LaCasse, L. & Colletti, P. (2000). Reduced prefrontal gray matter volume and reduced autonomic activity in antisocial personality disorder. Arch. Gen. Psychiatry 57, 119−127; discussion 128−129. PDF
  • Rilling, J.K., Gutman, D.A., Zeh, T.R., Pagnoni, G., Berns, G.S., & Kilts, C.D. (2002.) A neural basis for social cooperation. Neuron, 35, 395-405. PDF
  • Rottman, J., & Young, L. (2015). Mechanisms of Moral Development. The Moral Brain: A Multidisciplinary Perspective, 123.PDF
  • Saslow, L. R., Willer, R., Feinberg, M., Piff, P. K., Clark, K., Keltner, D., & Saturn, S. R. (2012). My brother’s keeper? Compassion predicts generosity more among less religious individuals. Social Psychological and Personality Science, 4, 31–38. PDF
  • Schaefer, M., Haun, D., & Tomasello, M. (2015). Fair is not fair everywhere. Psychological Science, 26, 1252-1260. PDF
  • Sell, A., Sznycer, D., Al-Shawaf, L., Lim, J., Krauss, A., Feldman, A., ... & Tooby, J. (2017). The grammar of anger: Mapping the computational architecture of a recalibrational emotion. Cognition, 168, 110-128. PDF
  • Shariff, A. F. (2015). Does religion increase moral behavior?. Current Opinion in Psychology, 6, 108-113.PDF
  • Shariff, A. F., Willard, A. K., Andersen, T., & Norenzayan, A. (2016). Religious Priming- A metanalysis with a focus on prosociality. Personality and Social Psychology Review, 20, 27-48. PDF
  • Shariff, A.F. & Norenzayan, A. (2007). God is watching you: Priming God concepts increases prosocial behavior in an anonymous economic game. Psychological Science, 18, 803-809. PDF
  • Singer, T., Kiebel, S.J., Winston, J.S., Kaube, H., Dolan, R.J., & Frith, C.D. (2004). Brain responses to the acquired moral status of faces. Neuron, 41,653-662. PDF
  • Sivan, J., Curry, O. S., & Van Lissa, C. J. (2018). Excavating the foundations: Cognitive adaptations for multiple moral domains. Evolutionary Psychological Science, 1-12. PDF
  • Sznycer, D., Delton, A. W., Robertson, T. E., Cosmides, L., & Tooby, J. (2018). The ecological rationality of helping others: Potential helpers integrate cues of recipients' need and willingness to sacrifice. Evolution and Human Behavior. 40(1), 34-45. PDF
  • Soderstrom, H., Hultin, L., Tullberg, M., Wikkelso, C., Ekholm, S., & Forsman, A. (2002). Reduced frontotemporal perfusion in psychopathic personality. Psychiatry Research: Neuroimaging114(2), 81-94. PDF
  • Stone, V., Cosmides, L., Tooby, J., Kroll, N. & Knight, R.T. (2002.) Selective impairment of reasoning about social exchange in a patient with bilateral limbic system damage. Proceedings of the National Academy of Sciences USA, 99, 11531-11536. PDF
  • Takahashi, H., Yahata, N., Koeda, M., Matsuda, T., Asai, K., & Okubo, Y. (2004). Brain activation associated with evaluative processes of guilt and embarrassment: An fMRI study. Neuroimage23(3), 967-974.
  • Tomasello, M.  & Vaish, A. (2013). Origins of human cooperation and morality.  Annual Review of Psychology, 64, 231–55. PDF
  • Tsoi, L., Dungan, J., Waytz, A., & Young, L. (2016). Distinct neural patterns of social cognition for cooperation versus competition. Neuroimage, 137, 86-96. PDF
  • Uhlmann, E. L., Pizarro, D. A., & Diermeier, D. (2015). A person-centered approach to moral judgment. Perspectives on Psychological Science, 10(1), 72-81.PDF
  • Van Elk, M., Matzke, D., Gronau, Q. F., Guan, M., Vandekerckhove, J., & Wagenmakers, E. J. (2015). Meta-analyses are no substitute for registered replications: A skeptical perspective on religious priming. Frontiers in psychology, 6.PDF
  • Wallace, B., Cesarini, D., Lichtenstein, P., & Johannesson, M. (2007). Heritability of ultimatum game responder behavior. Proceedings of the National Academy of Sciences, 10(1073). PDF
  • Wallach, W., Franklin, S., & Allen, C. (2010). A conceptual and computational model of moral decision making in human and artificial agents. In W. Wallach & S. Franklin (Eds.), Topics in Cognitive Science, special issue on Cognitive Based Theories of Moral Decision Making (pp. 454-485): Cognitive Science Society. PDF
  • Willard, A. K., Shariff, A. F., & Norenzayan, A. (2016). Religious priming as a research tool for studying religion- Evidentiary value, current issues, and future directions. Current Opinion in Psychology, 12, 71-75. PDF
  • Young, L., Bechara, A., Tranel, D., Damasio, H., Hauser, M., & Damasio, A. (2010). Damage to ventromedial prefrontal cortex impairs judgment of harmful intent. Neuron, 65, 845-851. PDF
  • Young, L., Camprodon, J., Hauser, M., Pascual-Leone, A., & Saxe, R. (2010). Disruption of the right temporoparietal junction with transcranial magnetic stimulation reduces the role of beliefs in moral judgments. Proceedings of the National Academy of Sciences, 107, 6753-6758. PDF
  • Young, L., Cushman, F., Adolphs, R., Tranel, D., & Hauser, M. D. (2006). Does emotion mediate the relationship between an action’s moral status and its intentional status? Neuropsychological evidence. Journal of Cognition and Culture6(1-2), 265-278. PDF
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