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Psychology Review: Evidence from neuroeconomic studies delineating reasoned and emotional processes in decision-making in the ultimatum game

Psychology Review: Evidence from neuroeconomic studies delineating reasoned and emotional processes in decision-making in the ultimatum game

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An essay for the 2012 Undergraduate Awards (International Programme) Competition by Delia Fuhrmann. Originally submitted for PS4040 Psychology Review at University of St. Andrews, with lecturer Martin Campbell in the category of Psychology
An essay for the 2012 Undergraduate Awards (International Programme) Competition by Delia Fuhrmann. Originally submitted for PS4040 Psychology Review at University of St. Andrews, with lecturer Martin Campbell in the category of Psychology

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Psychology Review: Evidence from neuroeconomic studies delineating reasoned and emotional processes in decision-making in the ultimatum

game ABSTRACT The ultimatum game is a single-staged economic bargaining paradigm in which a proposer is given an endowment and makes an offer for how to split it with a recipient. If the recipient accepts, both players receive what the proposer offered. If the recipient rejects, both players get nothing. Contrary to predictions from traditional economic theory, participants reliably choose fairness over material utility (Güth, Schmittberger & Schwarze, 1982). Dual-process models of decision making predict that emotions are the proximate reason for favouring fairness norms and compete with reasoned processes favouring material utility (Harlé & Sanfey, 2007). In economic practise emotions appear to play an important role as well. Fenton-O’Creevy, Soane, Nicholson and Willman (2010), for example, showed that the economic decisions of traders were influenced by their emotion regulation strategy and had a direct impact on their performance. This essay reviews neuroeconomic studies that investigate the effect of reasoned and emotional processes on outcomes of economic decision-making in the ultimatum game. The neuroeconomic research designed to address the question of reason and emotion in the ultimatum game is diverse and creative. The convergent evidence from neuroimaging, electrophysiological, clinical and lesion studies powerfully demonstrates the importance of both emotional and reasoned processes and shows that the two interact dynamically in the decision-making in the ultimatum game. Lesion studies, for example, highlight the fact that a disruption of the reasoning system can lead to both decreased material utilitarianism (Moretti, Dragone and di Pellegrino, 2008) or increased material


utilitarianism (Knoch, Pascual-Leone, Meyer, Treyer & Fehr, 2006). Thus, both emotional and reasoned processes appear to be linked to materially utilitarian, as well as fair outcomes. This powerfully challenges dual-process models. A social utility of fairness perspective (Lowenstein, Thompson & Bazerman, 1989) is argued here, that may integrate the conflicting findings and emphasises the social dimension of processes in the ultimatum game for future research. From an evolutionary perspective Nowak, Page and Sigmund (2000), for example, argued that the short-term cost of rejecting a low offer might be outweighed by the long-term benefit of the reputation that one accepts only fair offers. Perhaps people therefore usually desire a good reputation over wealth and the choice to reject unfair offers may serve this ultimately selfish and utilitarian goal (Frith & Singer, 2008).

KEYWORDS Neuroeconomics; decision-making; emotion; rationality; ultimatum game


INTRODUCTION Rationalist philosophers such as Plato (427-347 BC), who described reason as the charioteer dominating unruly passions that drive the human soul, or Spinoza (1632-1677), who proposed that emotions are flawed thoughts that confound


superior rationality, disseminated the notion that rationality is the superior sentiment in the dichotomy of emotion and rationality (Blackburn, 2008). In tradition with this school of thought, economic models traditionally predict that the behaviour of homo economicus is largely ruled by rationality and material utility (Hausman, 2008). However, in economic practise emotions appear to play an important role, too. Fenton-O’Creevy, Soane, Nicholson and Willman (2010), for example, showed that the economic decisions of traders were influenced by their emotion regulation strategy and had a direct impact on their performance. Experimental findings, such as that of the ultimatum game, cast further doubt on the purely rational account of economy. The ultimatum game investigates economic bargaining behaviour. It involves two players - the proposer and the recipient. The proposer is given an endowment (e.g., 10 Pounds) and makes an offer for how to split it with the recipient. If the recipient accepts the proposal, both players receive what the proposer offered. If he rejects, both players get nothing. In any case, the single-staged game is over (Güth, Schmittberger & Schwarze, 1982). For an illustration of the computerised presentation of the game see Figure1.


Figure1. A typical computerised presentation of stimuli in the ultimatum game. Adapted from Sanfey, Rilling, Aronson, Nystrom and Cohen (2003). Economic utility predicts that the recipient should accept any offer greater than zero. Therefore, the proposer ought to always offer the smallest non-zero offer possible. With remarkable reliability however, a seemingly paradoxical behaviour is exhibited by both proposer and recipient. The proposer usually offers around 50% of the endowment and the recipient will reject offers lower than 30% (Camerer & Thaler, 1995). What motivates people to forgo material gain in such a one-shot interaction? What processes influence the outcome of decision-making in the ultimatum game? An explanation is offered by dual-process models of decision making, which describe decisions in terms of an affective, intuitive system (“System 1”) and a computational, deliberative component (“System 2”) and predict that emotions are the proximate reason for a normative decision favouring fairness and compete with reasoned processes favouring material utility (Harlé & Sanfey, 2007). The emerging field of Neuroeconomics has taken a keen interest in economic bargaining and used it to model the processes linking mind and action by 4

revealing the neurobiological mechanisms of decision making (Glimcher & Rustichini, 2004). This essay will review neuroeconomic studies that delineate rational and emotive processes to assess their contribution to decision-making in the ultimatum game. To this end convergent evidence from the following methodological approaches will be discussed: • • • • Neuroimaging studies Electrophysiological studies Clinical studies Lesion studies.

Science Direct, Web of Knowledge and PubMed were the primary databases used to review the available evidence. An initial review of studies was done by using “ultimatum game” and “emotion OR affect OR emotive” as a search term in topic, keywords, abstract or title. The results were then refined with the different methodological approaches discussed here, for example, “lesion” or “fMRI”. After key studies were identified, relevant references in these articles were interrogated. The search was concluded on the 13th of February 2012.

REVIEW Neuroimaging studies Neuroimaging, especially functional magnetic resonance imaging (fMRI), is becoming an increasingly popular and accessible methodological approach in Neuroscience and investigating task-related activity in the brain allows functional mapping with good spatial resolution and may yield insights into different processes involved in shaping behaviour (Gazzaniga, Ivry & Mangun, 2009). 5

Sanfey et al. (2003) laid the foundation for most of the Neuroecomic research on the ultimatum game by investigating activity in brain regions associated with emotion and cognition by using fMRI. Note that this study - like most studies reviewed here - only investigated the role of the recipient because the recipient’s decision between material utility and fairness is not hampered by tactical considerations (Camerer & Thaler, 1995). The first important finding of this study was that unfair offers were rejected significantly more often when the proposer was human, as compared to a computer (Figure2). This finding has been replicated many times (e.g., Knoch, Pascual-Leone, Meyer, Treyer & Fehr, 2006) and suggests that the attribution of unfair intent is associated with stronger emotional reactions in recipients. It also highlights an element of punishment in the rejection of unfair offers. This behaviour has been termed “altruistic punishment” because rejecting an unfair offer is costly for the individual but benefits everyone in the group since it may reinforce cooperation (Jensen, 2010). Sanfey et al.’s (2003) neuroimaging data revealed that unfair offers were related to activity in brain areas associated with both negative emotion (anterior insula) and reason (dorsolateral prefrontal cortex - dlPFC), as well as conflict mediation (anterior cingulate cortex – ACC). The authors found that rejected and unfair offers correlated with a greater anterior insula than dlPFC activation, whereas accepted unfair offers correlated with a greater dlPFC than anterior insula activity. Note that not the absolute activation, but the contrast of activation was found to be predictive of decisions. Subsequent studies largely replicated and elaborated on Sanfey et al.’s (2003) findings (see Chapman, Kim, Susskind & Anderson, 2009, for example). Some of Sanfey et al.’s (2003) conclusions are controversial, however. The authors related dlPFC activity to processing financial goals, for example, while other studies implicated the same region for processing fairness goals (e.g., Knoch et al., 2006). It remains for future research to devise a


functional model predictive of behaviour and for neuroimaging studies to specify the parameters of the relative activation of parts of the brain processing emotions and cognitions further. Perhaps optical imaging, a relatively new method which offers brilliant temporal and spatial resolution (Gazzaniga, et al., 2009) could be applied to the problem.

Figure2. Sanfey et al.’s (2003) behavioural results in the ultimatum game comparing the recipient’s acceptance of offers made by a human proposer or a computer. In another fMRI study, Tabinibia, Satpute and Lieberman’s (2008) used a modified ultimatum game paradigm that varied the size of the offer and thus very elegantly isolated fairness from monetary gain (Figure3). They found that accepting fair, compared to unfair, but more monetarily attractive offers, led to higher happiness ratings and correlated with an activity in reward regions, such as the ventral striatum and the amygdala. The preference for unfair but desirable offers, in contrast, correlated with an increased activity in the right ventrolateral prefrontal cortex (rvlPFC), a region associated with emotion regulation, as well as a decrease in anterior insula activity, a region associated with disgust. First of all, 7

this study illustrates how powerful self-report data may be in strengthening a neuroeconomic argument. The authors’ eloquent conclusion that there is hedonic value in fairness is of great interest to our current discussion as well, since it illustrates the intriguing possibility that the reward associated with supporting fairness and punishing unfairness could be greater than the monetary gain of not doing so. Rejections in the ultimatum game might not equal non-utilitarian behaviour. Note however, that the inherent correlational nature of neuroimaging findings cannot rule out alternative explanations and thus is not sufficient to establish causality here. For example, a reduction in insula activity measured by fMRI could be due to emotion regulation or a simply a lack of negative emotions.

Figure3. The isolation of material utility and fitness via the variation of the offer size devised by Tabibnia et al. (2008). All in all, the neuroimaging data is able to make invaluable contributions to the current discussion by illustrating dynamic patterns of processing, as seen in Tabibnia et al. (2008). Indeed, one recent study even showed that it was possible to predict decisions in the ultimatum game using real-time fMRI (Hollmann et al., 2011). Further, fMRI can be tremendously useful for the generation of 8

hypotheses, as demonstrated by Sanfey et al. (2003). However, the inverse inference from fMRI employed by these studies, that is, the inference of cognitive processes from patterns of activation, is not strictly deductively valid (Poldrack, 2006). Corroborating these neuroimaging findings using methods other than neuroimaging is thus absolutely necessary. Future research could employ neuroimaging methods to investigate developmental change in economic decision-making, an area that has received little attention in the literature so far (Kogut, 2012).

Electrophysiological studies Electrophysiological studies may yield information about the brain’s activity as well as the bodies’ response to certain tasks by recording electrical signals. For instance, sizable populations of firing neurons in the brain emit electrical signals that can be recorded on the scalp with electroencephalography (EEG). The changes in activity in response to an event are then recorded with very high temporal resolution by averaging signals to compute event-related potentials (ERPs). Other measurements commonly used are skin-conductance recordings (SCRs) that record changes in skin moisture related to sympathetic arousal, or electromyograms (EMs), which reflect muscle activity (Gazzaniga et al., 2009). Chapman et al. (2009) devised an ingenious study to investigate the role of disgust in the ultimatum game. They recorded EMs of the levator labii muscle region of the face, which is characteristic of the typical oralnasal rejection response of disgust (Figure4). They compared moral disgust elicited by unfair treatment in an economic game to gustatory distaste and basic disgust and found that all evoked the same muscle activity. The authors conclude that these results suggest that immorality elicits the same disgust as disease vectors and


distastes. If moral disgust is indeed closely related to simple disgust, one might expect that moral disgust leads to later avoidance of transgressors just like basic disgust leads to an avoidance of contamination. This intriguing possibility could be investigated neuroeconomically with multi-staged ultimatum game paradigms, for example. The reputation of the proposer could influence the bargaining behaviour of recipients in that they might be less willing to accept offers from individuals who are known to have made unfair offers in the past (Sigmund, Hauert & Nowak, 2001).

Figure4. The typical oralnasal rejection response in disgust associated with gustatory distaste, basic disgust and moral disgust. Adapted from Chapman et al. (2009). Van’t Wout, Kahn, Sanfey and Aleman (2006) recorded SCRs prior to decisionmaking in the ultimatum game and found that these were higher for unfair offers as well as predictive of rejections for offers made by a human proposer, but not by a computer. This they took as evidence for the importance of emotions in economic decision-making. Arguably however, the possibility that rational processes are the origin of physiological arousal and emotion just a by-product cannot be ruled out. While this is good evidence that emotions play some role in decision-making in the ultimatum game, it remains unclear if they are the only 10

determinant of rejections in the game. One study by Hewig et al., (2010) combined ERPs, SCRs and subjective recordings. Stronger SCRs, as well as negative emotions in self-report, were predictive of rejections in the ultimatum game. Importantly, they related SCRs to the amygdala, a well established generator of negative emotions. The feedback negativity (FN), a brain potential generated by the anterior cingulate cortex (ACC), was measured with ERP and found to be predictive as well. This they took as evidence for a role of the ACC in the integration of reinforcement and punishment. Critically however, they found that rejections were related to more positive amplitudes with fair offers but no systematic relationship for unfair offers was found, which contradicts their hypothesis. While the findings for SCRs and self-report data were quite robust and explained about 40% of the variance, the FN explained only 10%, which casts doubt on the proposed importance of the ACC. One ERP component, perhaps more predictive of behaviour in the ultimatum game, is a group of waves related to conflicts and learning – the medial frontal negativities (MFNs; Boksem & De Cremer, 2010). In order to investigate if physiological measurements indeed reflect reasoned and emotional components differentially, one highly relevant study (van der Veen & Sahibdin, 2011) disentangled the size of the offer and fairness, similar to Tabibnia et al. (2008), and investigated cardiac deceleration as an affective measure, as well as MFNs as a reasoning measure. Unfair and high offers were more readily accepted than unfair and low offers, an effect not described by Tabibnia et al. (2008) who simply looked at the offer as a percentage of the endowment. This indicates that utilitarian motives can influence decision-making in the ultimatum game to some extent at least. They further found that the amplitude of the MFN correlated positively with the willingness to accept offers, whereas cardiac deceleration correlated negatively. Overall, cardiac and 11

electrocortical measures were found to only modestly correlate with each other. This dissociation they interpreted as evidence for the independence between affective, cardiac responses and brain potentials reflecting deliberative processes. While this finding is highly relevant, the authors unfortunately present the electrophysiological results inconsistently and imprecisely. It remains unclear how exactly fairness and offer size selectively influence cardiac and electrocortical measures. Therefore no final conclusions can be drawn from this research. Knoch, Gianotti, Baumgartner and Fehr (2010) applied ERPs in an unusual way. They looked at the overall strength of resting state activation instead of specific negativities or positivities. They found that lower cortical activity in the right prefrontal cortex (rPFC), as part of the reasoning system, was correlated with fewer rejections of unfair offers. They concluded that the energisation provided by the rPFC is a prerequisite for fairness. This highlights the importance of motivation for effortful behaviour and shows that a compromised reasoning process can lead to more material utilitarianism. This finding is of pivotal importance since it contradicts the dual process models of decision-making. What remains unclear, however, is how this decreased cortical activation might interact with personal goals. If someone’s goal was to maximise material utility we might expect a preference for material gain to be compromised by decreased rPFC activation due to a lack of energy for effortful behaviour. Overall, electrophysiological research makes valuable contributions to the discussion of the current matter in that it consistently highlights the importance of affective arousal as measured by SCRs (van t’Wout et al., 2006 and Hewig et al., 2010). It is able to provide good support for the importance of disgust for punishing unfairness (Chapman et al., 2009) and challenge the dual process


model of decision-making (Knoch et al., 2010). While frontal negativities generated by the ACC were implicated in decision-making in the ultimatum game repeatedly by ERP studies, the findings are inconsistent. Future research should investigate the specific processes associated with different ERP measurements and specify the function of the ACC in decision-making.

Clinical studies The investigation of the effect of relevant psychopathologies on behaviour in the ultimatum game is of great significance to the present discussion since it may illustrate the broader implications of selective impairments of rational or emotional processing. Csukly, Polgár, Tombor, Réthelyi and Kéri (2011), for example, found deviations in the behaviour of patients with schizophrenia. This patient group accepted unfair offers at a higher rate and fair offers at a lower rate. The authors also investigated the effect of emotion recognition and found that unlike controls who show an increased likelihood of accepting fairer offers when the proposer exhibits a positive facial expression, patients with schizophrenia did not show a similar pattern. However, the study failed to record behaviour for a neutral expression which could serve as a baseline comparator, which lessens the validity of conclusions about the impact of facial expressions. Additionally, most patients studied here were medicated, which is not unusual, but limits the power of the study considerably. The anti-social behaviour exhibited in psychopathy has been linked to an impairment of affect regulation (Davey, 2008) and is therefore highly interesting for the present discussion. Koenigs, Kruepke and Newman (2010) looked at the behaviour of imprisoned criminals as the recipients in the ultimatum game. They compared non-psychopaths, primary and secondary psychopaths. Primary, low13

anxious, psychopathy is a consequence of an intrinsic, perhaps biological, deficit present throughout development, whereas secondary, high-anxious, psychopathy reflects acquired behaviours that are the consequence of other psychopathologies or the membership in a deviant group (Newman, MacCoon, Vaughn & Sadeh, 2005). Koeigs et al. (2010) found that primary psychopathy was associated with significantly lower acceptance of unfair offers, while secondary psychopathy was not. Interestingly, ventromedial prefrontal cortex (vmPFC) patients have been described as “pseudo-psychopathic” since they exhibit symptoms of exaggerated anger with diminished shame and guilt. Based on this, the authors related primary psychopathy to vmPFC dysfunction and a defect in affect regulation. Osumi and Ohira (2010) found contradicting results however. In their study, high primary psychopathic participants more often chose economic utility than low psychopathic individuals by accepting unfair offers. They recorded SCRs as well and found that high psychopathic participants less often exhibited an electrodermal response to unfair offers. The authors concluded that a lack of affect in psychopathy leads to insensitivity to unfairness that may even be advantageous in economic settings. Since this aspect of psychopathy resembled Machiavellism, exhibited for example by successful businessman, it may be adaptive, they write. Due to the fact that the authors only investigated behaviour in the role of the recipient in a single-staged game, their study arguably does not allow for such a generalisation, however. For substantial supports of their claims, it would be necessary to investigate if the anti-social tendencies are indeed advantageous in a more naturalistic, multi-staged game (Sigmund, et al., 2001). Why the studies discussed above produced conflicting results is not clear. Perhaps psychopathy manifests differently in different cultures. Japanese participants generally exhibit more self- control than American ones (Weisz, 14

Rothbaum & Blackburn, 1984) and are perhaps not as prone to punishing unfairness. Future research will have to explore this and other possibilities and clarify if psychopathy is associated with a problem in affect regulation or affect per se. In summary, clinical studies may uniquely illustrate how a clinical impairment of affect or rationality may manifest in economic decision making. Moreover, it shows how clinical disorders may influence economic behaviour, furthering our understanding of their possible socioeconomic consequences. However, impairments in clinical populations are usually not restricted to single domains of processing, which limits the generalisability of these findings from a neuroeconomic perspective. Surprisingly anxiety, a pathology associated not with a lack, but with a detrimental intrusion of emotion, has received little attention so far. Future research could investigate social phobia, for example. This pathology with its insecurity and fear of judgement has a strong theoretical link to the present matter. Another promising approach for future research might be to integrate neuroimaging data into clinical studies to relate differential patterns of brain activation to behavioural deviations.

Lesion studies The study of patients with brain damage has been critical for the development of cognitive neuroscience since it may illustrate the impact of selective damage of certain regions on behaviour. Some relatively recent technical advances have made it possible to induce virtual lesions non-invasively with transcranial magnetic stimulation (TMS). This provides non-correlational data and may therefore indicate necessity of certain brain regions for certain functions (Gazzaniga et al., 2009). Van’t Wout, Kahn, Sanfey and Aleman (2005) found a 15

trend to accept unfair offers, as well as longer reaction times for accepting unfair offers, when the right dlPFC was disrupted with TMS. Knoch et al. (2006) replicated these results. They found an increased willingness to accept unfair offers, while fairness judgements were similar to those of control groups (Figure5). They conclude that the right dlPFC is necessary for the implementation of fairness-related goals, independent of the abstract knowledge about fairness. These conclusions differ from Sanfey et al. (2003), who related the dlPFC to carrying out materially utilitarian behaviour. Perhaps there is a more generalised role for the dlPFC in the computation of goals, irrespectively of fairness or material utility (Rangel & Hare, 2010). Again this shows that the dual process model might be too simplistic since the disruption of a reasoning process can lead to both decreased fairness and material utility depending on individual goals.

Figure5. Acceptance rates (A) and fairness rating (B) of unfair offers when the left dlPFC was disrupted with TMS (Left TMS), the right dlPFC was disrupted with TMS (Right TMS) or participants received a placebo treatment (Sham). Adapted from Knoch et al. (2006).


The role of the vmPFC, an area implicated by the electrophysiological measurements (van t’Wout et al., 2006), was extensively researched in lesion studies. Patients with damage to the vmPFC typically exhibit reduced ability to regulate emotions, in particular frustration and anger. Koenings and Tranel (2007) found that they also opted for rejecting unfair offers more often than controls and described them as “hyper-irrational”. The authors conclude that emotion regulation is a key component of economic decision-making. Interestingly, the behaviour of healthy participants was similar when serotonin depletion diminished vmPFC activity (Crockett, Clark, Tabibnia, Lieberman & Robbins, 2008). However, the fact that emotions other than anger are often found to be flattened, not heightened, in vmPFC patients casts doubt on this account of the vmPFC (Koenings et al., 2007). Indeed, Moretti, Dragone and di Pellegrino (2008) conducted a very insightful study which found that acceptance rates of unfair offers were not reduced in vmPFC patients if the financial gains were visible and readily available. What is more, they did not differ between offers from computers or human proposers. On the basis of this finding they convincingly argued that emotions related to self-interest, such as anger about financial loss are intact, or even heightened in vmPFC patients, and propose a role of the vmPFC in social emotions, such as compassion or shame. This is yet another indication that a dual process model of decision making might need revision. One important limitation of lesion studies is that the network properties of the brain are often not well understood. Indeed, lesions to the vmPFC may lead to effects in brain regions distant from the lesion and future research needs to establish how this region interacts with others, such as the ACC or dlPFC. To summarise, lesion studies have proven a powerful tool in the investigation of reason and emotion in the ultimatum game. They show that the disruption of a reasoning system can lead to both decreased material utilitarianism (Moretti et 17

al., 2008) or increased material utilitarianism (Knoch et al., 2006), which powerfully challenges dual process models. Apparently, the impairment of different types of emotions can lead to differential outcomes in the ultimatum game. The distinction found in the social vs. individual axis is intriguing because it raises the question how social vs. individualistic reasoning may affect outcomes in the ultimatum game. Future studies can elaborate on these findings and perhaps the controversial role of the ACC for economic bargaining could be clarified with lesion studies as well.

CONCLUSION This review discussed neuroimaging, electrophysiological, clinical and lesion studies delineating reasoned and emotional processes underlying decisionmaking in the ultimatum game. Overall, there is strong convergent evidence for the importance of both emotional and reasoned processes for decision-making in the ultimatum game, revealing a complex interdependence. Crucially and in opposition to traditional dual process models, such as that of Harlé and Sanfey (2007), the contribution of neither process was limited to one outcome – material utilitarian or fairness. Instead, the two processes appear to interact dynamically and dependant on individual goals. In order to make sense of these findings, perhaps the problem needs to be reassessed from a different angle: Rejecting unfair offers in the ultimatum game can be described as utilitarian if the object of choice is not financial gain but fairness (Hausman, 2008). Indeed, Lowenstein, Thompson and Bazerman (1989) argued the existence of a social utility, that is people feel greater satisfaction with an equal division of resources, even if that is not in favour of themselves. Therefore rejecting unfair offers and making fair offers may be reinforced by reward processes and serve goals that are ultimately


utilitarian. The true distinction may then lie in a social vs. an individualistic choice - not an emotional vs. reasoned one. See Figure6 for a proposed model of a feedback loop designed to integrate evidence from the studies discussed here.

Reason Goals (dlPFC) Energisation (rPFC) Conflict mediatio n and learning (ACC) Emotion regulation (rvlPFC) Accounting for social motives (vmPFC)

Emotion Disgust (insula) Arousal (amygdala) Reward (ventral striatum, amygdala)


Figure6. A proposed feedback loop of reasoned and emotional processes driving decision-making in the ultimatum game, that builds on Sanfey et al’s (2003) notion of the relative engagement of cognitive and emotional regions mediated by the ACC and integrates findings from neuroimaging, physiological, lesion and clinical studies. Abbreviations used: anterior cingulate cortex - ACC, dorsolateral prefrontal cortex – dlPFC,, right prefrontal cortex – rPFC, right ventrolateral prefrontal cortex –rvlPFC, ventromedial prefrontal cortex – vmPFC.


The neuroeconomic research designed to address the question of reason and emotion in the ultimatum game is very diverse and remarkably creative. It powerfully demonstrates a multitude of processes likely to be involved in many situations of decision-making. However the limitations of different neuroecomic methods are certainly not negligible, likely contributing to conflicting findings. Therefore the integration of evidence from several approaches is vital and has proven valuable here. Further, the research on this topic suffers from a noticeable value-leadenness. Osumi and Ohira (2010), as much of the Neuroscientific literature, for example, make the implicit assumption that emotionality leads to a less than optimal choice (Frith & Singer, 2008). For other studies, such as Sanfey et al. (2003), the opposite seems to be true and research appears to be strongly motivated by a wish to promote the positive role of emotionality. As argued here from a social utility perspective, both processes could be important and may contribute to differential outcomes of the game. In order to investigate the role of social and individualistic motivations in the ultimatum game further, revisiting the evolutionary perspective put forward by Chapman et al. (2009) may be beneficial. Nowak, Page and Sigmund (2000), for example, argued that fairness is evolutionarily stable if the proposer has information on what offers the responder has accepted in the past and highlighted a role for reputation. The short-term cost of rejecting a low offer might be outweighed by the long-term benefit of the reputation that one accepts only fair offers. This could then foster co-operation. This climate of cooperation that is beneficial to each individual in the group may be reinforced by altruistic punishment mechanisms, similar to those observed in the ultimatum game. The punishment of free-riders may then be crucial to maintain co-operation in a society (Jensen, 2010). Indeed, this reinforcement of egalitarianism might have played a decisive role in human evolution – our closest primate relative, the 20

chimpanzee (pan troglodytes), does not show such an other-regarding preference in the ultimatum game (Jensen, Call & Tomasello, 2007). Perhaps people therefore usually desire a good reputation over wealth and the choice to reject unfair offers may serve this ultimately selfish and utilitarian goal (Frith & Singer, 2008).


In order to investigate this possibility in future research, neuroeconomic methods could be used to illustrate social emotions and cognitions in relation to fairness or material utilitarian motivations guiding behaviour in the ultimatum game. Neuroeconomic research might be especially valuable for investigating how the ACC could mediate social and individualistic motivations and thus inform our understanding of the computation of conflicting interests in the brain. Further, social and individualistic motivations and their processing could be investigated in the more naturalistic multi-staged games (Sigmund, et al., 2001). Another promising line of research for this field could be developmental change, as suggested by Kogut (2012). We might then be able to begin to disentangle nature from nurture and gain novel insights into the role reason and emotion, material utility and fairness, and social and individualistic choice play in decision-making in the ultimatum game.


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