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Cortical Thickness of the Dorsolateral Prefrontal Cortex Predicts Strategic Choices in Economic Games

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Cortical thickness of the dorsolateral prefrontal cortex predicts strategic choices in economic games Toshio Yamagishia,b,1, Haruto Takagishib, Alan de Souza Rodrigues Ferminb, Ryota Kanaic, Yang Lib, and Yoshie Matsumotob aGraduate School of International Corporate Strategy, Hitotsubashi University, 2-1-2 Hitotsubashi, Chiyoda-ku, Tokyo 101-8439, Japan; bBrain Science Institute, Tamagawa University, 6-1-1 Tamagawagakuen, Machida, Tokyo 194-8610, Japan; and cDepartment of Neuroinformatics, Araya Brain Imaging, 3-16-16 Daizawa, Setagayaku, Tokyo 155-0032, Japan Edited by Susan T. Fiske, Princeton University, Princeton, NJ, and approved March 30, 2016 (received for review December 5, 2015) Human prosociality has been traditionally explained in the social cooperated more (23). Additionally, increased activity in the sciences in terms of internalized social norms. Recent neuroscien- lateral prefrontal cortex was negatively related to fairness-seeking tific studies extended this traditional view of human prosociality behavior in an economic game (24). According to the heuristic by providing evidence that prosocial choices in economic games prosociality model (14–17), humans are predisposed to cooperate require cognitive control of the impulsive pursuit of self-interest. in social exchange situations. People fail to behave in a prosocial However, this view is challenged by an intuitive prosociality view manner in social exchanges when this predisposition is overridden emphasizing the spontaneous and heuristic basis of prosocial by strategic reasoning to secure their self-interest. By comparing choices in economic games. We assessed the brain structure of participants’ behaviors in two economic games with brain structural 411 players of an ultimatum game (UG) and a dictator game (DG) differences and strategic reasoning abilities, we provide evidence and measured the strategic reasoning ability of 386. According to that strategic reasoning controls, and thus reduces rather than the reflective norm-enforcement view of prosociality, only those promotes, game players’ prosocial behavior. capable of strategically controlling their selfish impulses give a fair The contrast between two simple, two-person economic games— share in the UG, but cognitive control capability should not affect namely, the dictator game (DG) and the UG—is often used to behavior in the DG. Conversely, we support the intuitive proso- support the reflective prosocial model by demonstrating how strategic ciality view by showing for the first time, to our knowledge, that reasoning affects game players’ decisions. In both games, one player COGNITIVE SCIENCES strategic reasoning and cortical thickness of the dorsolateral freely decides how much of a fixed reward to take and how much to PSYCHOLOGICAL AND prefrontal cortex were not related to giving in the UG but were leave for the other player. The difference between the two games is negatively related to giving in the DG. This implies that the that the other player in the UG (termed “responder”)hastheoption uncontrolled choice in the DG is prosocial rather than selfish, and to reject the decision made by the first player (termed “proposer”), those who have a thicker dorsolateral prefrontal cortex and are causing both to earn nothing. This option is not provided to the capable of strategic reasoning (goal-directed use of the theory of second player in the DG, who plays the role of a “recipient”.The mind) control this intuitive drive for prosociality as a means to “ ” maximize reward when there are no future implications of choices. recipient simply receives whatever the first player ( dictator ) gives. The level of giving by the proposer in the UG is usually higher than ultimatum game | dictator game | strategic reasoning | DLPFC | that by the dictator in the DG (25). This is attributed to the pro- ’ ’ prosocial behavior poser s strategic reasoning, which requires inference of the recipient s internal state and prediction of the resulting response (e.g., anger on the basis of unfair giving and subsequent rejection) (8, 9, 13). Given umans are a cooperative species, and the question of why that neuroimaging and neuroendocrinological studies showed that humans are so cooperative has been a subject of consider- H negative emotions are associated with rejection of unfair offers able interest in social and biological sciences (1–4). The tradi- (24, 26, 27), UG proposers may anticipate negative responses to tional answer in the social sciences highlights critical roles of unfair offers. UG proposers anticipate norm-enforcing responses social norms and cultural values internalized as personal values (rejection of the offer) to norm-violating behavior (taking most of the and social preferences (5, 6). Recent neuroscientific studies of brain structure and activity extended this traditional view of human prosociality by showing that players of economic games act Significance prosocially when they cognitively control selfish impulses (7–13). Experimental evidence shows that prosocial choices in economic Is human prosociality a consequence of cognitive control of selfish games positively relate to local gray matter volume and thickness impulses? Alternatively, is it a default option that most people use and the activation of brain areas that control selfish impulsive unless they are cognitively persuaded that a given situation does drives, such as the dorsolateral prefrontal cortex (DLPFC) and not require them to behave prosocially? Our results support the temporoparietal junction (TPJ) (7–9). Furthermore, impairment of latter argument. Participants with weaker cognitive control fairly cognitive control by disruption of DLPFC function prevents re- shared a reward with another participant even when there was no jection of unfair offers in the ultimatum game (UG), which some chance of punishing unfair behavior, whereas those more capable authors considered prosocial and fairness-seeking behavior (10– of cognitive control behaved selfishly in the same situation. These findings demonstrate that participants’ intuitive choices in eco- 13). Recently, this reflective view of human prosociality has been nomic games are prosocial. challenged by an alternative view emphasizing the intuitive na- ture of prosocial behavior, subsumed under intuitive prosociality Author contributions: T.Y., Y.L., and Y.M. designed research; H.T., Y.L., and Y.M. per- (14) or heuristic cooperation (15–17). Support for the intuitive formed research; T.Y., H.T., A.d.S.R.F., and R.K. analyzed data; and T.Y., Y.L., and Y.M. and automatic nature of prosocial behavior is provided by find- wrote the paper. ings that prosocial choices are promoted under time pressure The authors declare no conflict of interest. (15, 16, 18), under cognitive load (19–21), or after priming This article is a PNAS Direct Submission. by successful experiences of intuitive decision making (15, 22). 1To whom correspondence should be addressed. Email: [email protected]. Also, participants who expressed more positive emotional words This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10. and less inhibitory words during and after an economic game 1073/pnas.1523940113/-/DCSupplemental. www.pnas.org/cgi/doi/10.1073/pnas.1523940113 PNAS Early Edition | 1of6 Downloaded by guest on October 1, 2021 thus make distinct predictions regarding the relationship between DLPFC thickness and behavior in the UG and DG. The reflective model predicts a positive relationship between DLPFC thickness and giving in the UG, whereas the intuitive model predicts a neg- ative relationship between DLPFC thickness and giving in the DG. These two alternative accounts of differences in giving in the UG and DG (8, 9) provide a way to test the intuitive selfishness assumption against the intuitive prosociality assumption. We first successfully replicated earlier findings that strategic behavior is more pronounced among those who had a thicker DLPFC than those who had a thinner DLPFC (8) in a study of 411 adult, nonstudent participants who played both the UG and DG and Fig. 1. Schematic representations of how strategic considerations generate from whom brain structural images were obtained. Then, we the difference between strategists (ST) and nonstrategists (Non-ST) in the UG found for the first time, to our knowledge, that local gray matter and DG. A shows the prediction that strategic considerations should improve thickness of the DLPFC negatively correlated with giving in the fair behavior in the UG. B shows the prediction that strategic considerations DG but was not correlated with giving in the UG (Fig. 2 C and should depress fair behavior in the DG. D). We further measured the strategic reasoning of 386 of these participants using a newly developed test of strategic reasoning, reward) and strategically adjust giving behavior to secure acceptance measured 411 participants’ Machiavellianism (34, 35) score, and by the responder. Thus, those capable of using strategic reasoning are found that those exhibiting better strategic reasoning behaved expected to make fair offers in the UG compared with those who more selfishly in the DG than those with poor strategic reasoning, struggle to control their selfish drive for immediate reward. but no relationship was found between task performance and In contrast, in the DG, which requires no strategic reasoning fairness in the UG. These striking findings provide strong evidence B to earn as much as possible, strategic control over selfish impulses
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