Levin et al. Neuropsychology of risky gains/losses Although a more detailed meta-analysis is clearly warranted, Table 1 shows that a wide variety of structures are involved in risky decision-making beyond those depicted in Figure 1. Never- theless, we feel that the relatively simple depiction of the model represents a good start in capturing the different neurological underpinnings of risk-taking for gains and losses. The comple- mentary roles of the VMPFC, amygdala, and insula depicted in the model are consistent with both the general hypothesis that separate processes underlie risk-taking for gains and losses, and the stricter hypothesis of separate neural structures coming together in different ways to guide risky decision-making in the gain and loss domains. In conclusion, we ﬁnd that evidence of differ- ent neural responses underlying risk-taking for gains and losses favors the hypothesis that decision makers react differently to risky gains and losses, both in terms of overt risk-taking and neural activation. REFERENCES Bechara, A., and Damasio, A. R. (2005). The somatic marker hypothesis: a neural theory of economic decision. Games Econ. Behav. 52, 336–372. Bechara, A., Damasio, A. R., Dama- sio, H., and Anderson, S. W. (1994). Insensitivity to future consequences following damage to human pre- frontal cortex. Cognition 50, 7–15. Bechara, A., Damasio, H., Damasio, A. R., and Lee, G. P. (1999). Differ- ent contributions of the amygdala and ventromedial prefrontal cortex to decision-making. J. Neurosci. 19, 5473–5481. Bechara, A., Damasio, H., Tranel, D., and Damasio, A. R. (1997). Deciding advantageously before knowing the advantageous strategy. Science 275, 1293–1295. Bechara, A., Tranel, D., Damasio, H., and Damasio, A. R. (1996). Failure to respond autonomically to antic- ipated future outcomes following damage to prefrontal cortex. Cereb. Cortex 6, 215–225. Breiter, H. C., Aharon, I., Kahne- man, D., Dale, A., and Shizgal, P. (2001). Functional imaging of neural responses to expectancy and experience of monetary gains and losses. Neuron 30, 619–639. Clark, L., Bechara, A., Damasio, H., Aitken, M. R. F., Sahakian, B. J., and Robbins, T. W. (2008). Distinct pat- terns of risky decision-making in patients with lesions to ventromedial prefrontal cortex and insular cortex. Brain 131, 1311–1322. Craig, A. D. (2009). How do you feel – now? The anterior insula and human awareness. Nat. Rev. Neurosci. 10, 59–70. Damasio, A. R. (1994). Descartes’ Error: Emotion, Reason, and the Human Brain. New York, NY: Putnam. Davidson, R., Ekman, P., Saron, C., Senulis, J., and Friesen, W. (1990). Approach-withdrawal and cerebral asymmetry: emotional expression and brain physiology I. J. Pers. Soc. Psychol. 58, 330–341. De Martino, B., Kumaran, D., Seymour, B., and Dolan, R. J. (2006). Frames, biases, and rational decision-making in the human brain. Science 313, 684–687. Fujiwara, J., Tobler, P. N., Taira, M., Iijima, T., and Tsutsui, K. I. (2009). Segregated and integrated coding of reward and punishment in the cin- gulate cortex. J. Neurophysiol. 101, 3284. Fukui, H., Murai, T., Fukuyama, H., Hayashi, T., and Hanakawa, T. (2005). Functional activity related to risk anticipation during perfor- mance of the Iowa gambling task. Neuroimage 24, 253–259. Gottfried, J. A., O’Doherty, J., and Dolan, R. J. (2002). Appetitive and aversive olfactory learning in humans studied using event- related functional magnetic reso- nance imaging. J. Neurosci. 22, 10829–10837. Hsu, M., Bhatt, M., Adolphs, R., Tranel, D., and Camerer, C. F. (2005). Neural systems responding to degrees of uncertainty in human decision- making. Science 310, 1680. Huettel, S. A., Song, A. W., and McCarthy, G. (2005). Decisions under uncertainty: probabilistic context inﬂuences activation of prefrontal and parietal cortices. J. Neurosci. 25, 3304–3311. Huettel, S. A., Stowe, C. J., Gor- don, E. M., Warner, B. T., and Platt, M. L. (2006). Neural signa- tures of economic preferences for risk and ambiguity. Neuron 49, 776–775. Kahneman, D. (2003). Perspective on judgment and choice: mapping bounded rationality. Am. Psychol. 58, 697–720. Kahneman, D., and Tversky, A. (1979). Prospect theory – analysis of deci- sion under risk. Econometrica 47, 263–291. Knutson, B., Adams, C. M., Fong, G. W., and Hommer, D. (2001). Anticipation of increasing mon- etary reward selectively recruits nucleus accumbens. J. Neurosci. 21, 159–164. Knutson, B., Rick, S., Wimmer, G. E., Prelec, D., and Loewenstein, G. (2007). Neural predictors of pur- chases. Neuron 53, 147–156. Knutson, B., Wimmer, G. E., Kuh- nen, C. M., and Winkielman, P. (2008a). Nucleus accumbens acti- vation mediates the inﬂuence of reward cues on ﬁnancial risk taking. Neuroreport 19, 509–513. Knutson, B., Wimmer, G. E., Rick, S., Hollon, N. G., Prelec, D., and Loewenstein, G. (2008b). Neural antecedents of the endowment effect. Neuron 58, 814–822. Kuhnen, C. M., and Knutson, B. (2005). The neural basis of ﬁnancial risk taking. Neuron 47, 763–770. LeDoux, J. E. (2000). Emotion circuits in the brain. Annu. Rev. Neurosci. 2, 155–184. Levin, I. P., Gaeth, G. J., Schreiber, J., and Lauriola, M. (2002). A new look at framing effects: Distribution of effect sizes, individual differences, and independence of types of effects. Organ. Behav. Hum. Decis. Process. 88, 411–429. Levin, I. P., Weller, J. A., Pederson, A., and Harshman, L. A. (2007). Agere- lated differences in adaptive decision making: sensitivity to expected value in risky choice. Judgm. Decis. Mak. 2, 225–233. Loewenstein, G. F., Weber, E. U., Hsee, C. K., and Welch, N. (2001). Risk as feelings. Psychol. Bull. 127, 267–286. Matthews, S. C., Simmons, A. N., Lane, S. D., and Paulus, M. P. (2004). Selective activation of the nucleus accumbens during risk-taking decision making. Neuroreport 15, 2123–2127. Mellers, B. A., Schwartz, A., and Ritov, I. (1999). Emotion-based choice. J. Exp. Psychol. 128, 1–14. Mohr, P. N. C., Biele, G., and Heek- eren, H. R. (2010). Neural pro- cessing of risk. J. Neurosci. 30, 6613–6619. O’Doherty, J. P., Kringelbach, M. L., Rolls, E. T., Hornak, J., and Andrews, C. (2001). Abstract reward and punishment representations in the human orbitofrontal cortex. Nat. Neurosci. 4, 95–102. Paton, J. J., Belova, M. A., Morrison, S. E., and Salzman, C. D. (2006). The primate amygdala represents the positive and negative value of visual stimuli during learning. Nature 439, 865–870. Paulus, M. P., and Frank, L. R. (2006). Anterior cingulate activity mod- ulates nonlinear decision weight function of uncertain prospects. Neuroimage 30, 668–677. Paulus, M. P., Rogalsky, C., Sim- mons, A., Feinstein, J. S., and Stein, M. B. (2003). Increased activation in the right insula during risk-taking decision mak- ing is related to harm avoidance and neuroticism. Neuroimage 19, 1439–1448. Paulus, M. P., and Stein, M. B. (2006). An insular view of anxiety. Biol. Psychiatry 60, 383–387. Phelps, E. A. (2006). Emotion and cog- nition: insights from studies of the human amygdala. Annu. Rev. Psy- chol. 57, 27–53. Pinker, S., and Ullman, M. T. (2002). The past and future of past tense. Trends Cogn. Sci. (Regul. Ed.) 6, 456–463. Plassmann, H., O’Doherty, J. P., and Rangel, A. (2010). Appetitive and aversive goal values are encoded in the medial orbitofrontal cortex at the time of decision making. J. Neurosci. 30, 10799. Preuschoff, K., Quartz, S. R., and Bossaerts, P. (2008). Human insula activation reﬂects risk prediction errors as well as risk. J. Neurosci. 28, 2745–2752. Reimann, M., and Bechara, A. (2010). The somatic marker framework as a neurological theory of decision- making: review, conceptual com- parisons, and future neuroeco- nomic research. J. Econ. Psychol. 31, 767–776. Sanfey, A. G., Rilling, J. K., Aron- son, J. A., Nystrom, L. E., and Cohen, J. D. (2003). The neural basis of economic decision-making in the ultimatum game. Science 300, 1755–1758. Seymour, B., Daw, N., Dayan, P., Singer, T., and Dolan, R. (2007). Differen- tial encoding of losses and gains in the human striatum. J. Neurosci. 27, 4826–4831. Slovic, P., Finucane, M., Peters, E., and MacGregor, D. (2002). “The affect heuristic,” in Heuristic and Biases: The Psychology of Intuitive Judgment, eds T. Gilovich, D. Grif- ﬁn, and D. Kahneman (New York, NY: Cambridge University Press), 397–420. Frontiers in Neuroscience | Decision Neuroscience February 2012 | Volume 6 | Article 15 | 10