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But this also holds true for rather cultural activities such as dancing, singing, or playing board games, where the activity itself consti- tutes the goal. As a consequence, joint action research aims at studying how individuals bring about such tasks or playfully act in concert. This does not imply that those activities cannot be ana- lyzed by game theory or that decisions play no role in joint action. This focus rather motivates a research orientation that empha- sizes details different from those put forward by neuroeconomics, Frontiers in Human Neuroscience www.

We will suggest in the course of this review that both perspectives are not necessarily incompatible with each other. Despite marked differences regarding the proposed con- stituents of joint action, current research can be summarized by the following coarse taxonomy of joint action Bratman, ; Searle, ; Tuomela, , ; Tomasello et al, ; Knoblich and Sebanz, ; Pacherie, First, joint action implies at least two agents intentionally acting together, that is, consciously pursuing compatible goals.

In cases of mutual task dependence, this implicates the agents' ability to coordinate with each other, mostly controlled automatically and without con- scious awareness.

This mainly includes perceptual sensitivity and behavioral responsiveness to the other's actions and mental states. Second, joint action, in many cases, implicates rather explic- itly shared mindsets and motivations including specific beliefs, desires, goals, and intentions.

Of note, "shared" emphasizes that those mindsets and motives can be actively expressed by com- municative gestures and verbal behavior. This distinction, in particular, responds to the fact that the reasons for which indi- viduals act together often significantly differ and may include full-blown cooperative, but also selfish or simply socially com- patible but actually private motives. We will address this point in greater detail toward the end of this review in the section termed "modes of cooperation and we-thinking.

The next section will tap into the more explicitly processed facets of joint action by reviewing findings and concepts from developmental and comparative psychology. The subsequent three sections will review current neuroimaging research pursuing rather integrative approaches with regard to the behavioral and cognitive facets of joint action and cooperation.

A final section will examine recent theoretical contributions from social philosophy and economics with regard to their potential of bringing together together the different variants and aspects.

This can even be observed in experimental settings that are highly unlikely to elicit explicitreflec- tion of mutual actions. For example, Richardson and colleagues had subjects sit in front of each other on rocking-chairs.

Those chairs were shaped in a way to bias for different rock- ing frequencies. Nevertheless, synchronization of the subjects' rocking frequencies was observed in this scenario. Further stud- ies prompting individuals to coordinate their behavior explored cognitive mechanisms for mutual adjustments.

In a behavioral study by Knoblich and Jordan subjects shared control over a tracking-device that had to be kept aligned with a horizon- tally moving object. Crucially, this rather difficult task could not be achieved individually. The results suggested that individuals indeed solve coordination problems by anticipating the others' moves without relying on explicit communication. A joint tap- ping paradigm extends these findings Konvalinka et al, : subjects had to synchronize to external beats or to their partner.

The results suggested that anticipation and adjustment performed by each individual are necessary but not sufficient for high syn- chronization performance. Instead, anticipation and adjustment need to be bidirectional, i.

First insights into the neural basis of coordination in joint action were provided by Newman-Norlund and colleagues In their fMRI experiment, single and joint action versions of a virtual lifting task were compared, in which subjects had to adjust their actions in order to prevent a ball from slipping off a bar. The results include increased activity in the inferior frontal gyrus IFG; pars opercularis and the posterior superior temporal sulcus pSTS , which are both believed to be part of the putative human mirror neuron system Keysers and Perrett, ; Oberman et al, ; Schulte-Ruther et al, ; Schilbach et al, ; Wang et al, This has been taken to suggest that when human actors achieve mutual coordination they rely on a motor representation of their partner's ongoing action.

Taken together, these findings shed light on advanced coor- dination skills that go beyond mere temporal estimation and prediction as suggested by the individual-joint comparisons. However, some caution is warranted regarding the temptation to take these examples as instantiations of mental-state coordi- nation or full-blown cooperation, as these studies do not permit any principled conclusions about the nature of the entities coor- dinated nor the agents' attitudes toward their acting together.

The following two sections will address these nuances in greater detail. Accordingly, insights into the coordi- nation of mental states in joint action can be drawn from another set of studies that specifically investigated how people share repre- sentations of their partner's actions, that is, how they form action co-representations.

Studies on action co-representation charac- teristically exploit the so-called Simon-Effect Simon, This effect is elicited in subjects that are asked to respond spatially to non-spatial features of stimuli while ignoring the location of the stimulus presentation. Characteristically, in tasks that elicit the Simon-Effect actual stimulus location affects reaction times.

For instance, when subjects respond with left button presses to green and with right button presses to red stimuli, they tend to be sig- nificantly slower when the green stimulus appears on the right side incongruent response as compared to the left side con- gruent response. In a seminal behavioral study by Sebanz et al. That is, the task was to respond to certain stimuli, e. Importantly, a Simon-Effect increased reaction times during incompatible trials was observed in this Frontiers in Human Neuroscience www.

In a number of follow-up experiments Sebanz et al, it was demonstrated that individuals co-represent not only their partners' actions but also the rules guiding their actions.

These cognitive entities were teased apart by implementing different tasks for both players, which sometimes necessitated the same and sometimes necessitated different responses to the same stimuli. Concretely, one subject had to respond to the direction of a stim- ulus, whereas the other subject had to respond to its color.

The Social Simon-Effect, that can be regarded as a quantitative behav- ioral marker for joint action, was markedly stronger when both tasks required different responses by the subjects. This indicates a high sensitivity to the partner's task, even if it was irrelevant to the subject's task. Interestingly, the number of neuroimaging studies that make use of Social Simon-Tasks is still limited. As suggested by the authors, these findings might reflect intensified inhibitory pro- cesses in response to the challenge of disentangling one's own and the partner's representations during reciprocally dependent action.

In the same vein, Sebanz et al. Taken together, these studies suggest that individuals are highly sensitive to their partners' actions and mental states, even when mutual coordination is not relevant for achieving a given task.

More recent studies following this methodological track suggest that the set of mental entities that can be tracked using the inter- active Simon Task is to be extended to the personal relationship between the actors Hommel et al, and cooperative inten- tions Ruys and Aarts, ; Iani et al, On the other hand, the Social Simon-Effect does not involve conscious or explicit processing of the mental states governing ones partner's behavior.

Recent findings suggest this effect to be grounded on low-level saliency mechanisms rather than higher-level representational processing Vlainic et al, ; Dolk et al, Capturing the explicit dimension of cognitive processes subserving joint action, at least to some extent, requires to permit subjects to interact and express themselves in a less constrained fashion.

The subsequent section discusses findings from developmental and comparative psychology highlighting the role of shared intentions and reward in joint action and cooperation. In a study by Warneken et al. In the former acting together lead to material rewards, whereas the latter aim at maintaining and enjoying the shared activity per se.

For example, in a typical instrumental task, food or toys were hidden inside a long tube with two handles and could only be released by the subjects' and experimenters' combined efforts.

In contrast, a typical social game was constituted by a trampoline which the experimenter and subject could utilize in concert to make a ball jump up and down. Chimpanzees and children displayed substantial coordi- nation skills in all instrumental tasks. Contrarily, it was hardly possible to make chimpanzees engage in social games, which appeared to be intrinsically rewarding to and month-old human toddlers.

Finally, as activities got spontaneously inter- rupted by the experimenter, children but not chimpanzees, tried to reengage the experimenter rather than trying to complete the task individually or engage in other activities. This observed effect is noteworthy, as it cannot be attributed to a lack of cognitive or motor capacities in chimpanzees.

Rather, great apes are likely able to understand others' goals and even, to some extent, others' knowledge Hare et al, Moreover, the example demonstrates that apes are capable coordinators when food rewards are expected. Comparing instrumental activ- ities with social games, thus suggests that children, but not chimpanzees, exhibit an intrinsic motivation to collaborate. The findings eluded to above also highlight the importance of distin- guishing games from instrumental activities and demonstrate the limits of current neuroeconomic approaches relying on instru- mental payoff in investigating the putative reward mechanisms underlying cooperation.

Interestingly, when looking at specific social motives that might explain the observed differences, these seem to go beyond altruistic helping helping irrespective of external rewards , which can be reliably elicited in chimpanzees Warneken et al, and, therefore, is not specific to humans. However, another series of studies by Grafenhain et al. This proposal is further motivated by philosophical analyses by Gilbert , Bratman , and Tuomela , on shared intentions.

The basic idea is that people form joint commitments as they act together. Hence, they strongly expect each other to fulfill their respective roles and try to help each other when problems arise during cooperation. Consequently, if one of the agents interrupts his participation in the shared task, her partner should not only be surprised, resulting in updated beliefs about the world, but should also exhibit disappointment, resentment, and other indi- cators for normative charge Rakoczy et al, The latter may then encourage attempts to reengage the cooperator, given the two individuals sharing social commitments to the joint activity.

This line of thought led to the experiments by Grafenhain et al. In one characteristic game, child and experimenter sat alongside each other in front Frontiers in Human Neuroscience www. The box was equipped with one handle for each of the two players that sets a rabbit free.

That is, the other's partic- ipation was not necessary to play the game, and moreover, the experimental setting allowed taking the other's role. This experi- mental detail constitutes a decisive variation of the games used by Warneken and colleagues that ultimately allowed the researchers to modulate explicitly the children's joint commitments while playing social games.

Assuming that joint commitments arise out of, within and by social interaction, they were varied by either positively engaging the child before and during the play contingent acting, mutual gaze, smiling at the child versus neu- trally acting in a rather parallel, unrelated fashion no contingent actions, smile not directed at child.

In subsequent interrup- tion periods, around 2-year-old children more often attempted to reengage the experimenter and less often continued the activ- ity on their own when playing in the commitment-facilitating condition compared to the neutral condition.

Furthermore, in a subsequent experiment Grafenhain and colleagues nicely demonstrated that young children anticipate the experimenter's expectations toward the child depending on their commitments.

As a second experimenter tried to engage them into a more inter- esting game, the children displayed leave-taking behavior e. What makes these findings particularly intriguing is, that the partner, i. Besides again highlighting the importance of social games as a means to access the intrinsic nature of human motivations underlying cooperation, these find- ings also emphasize the importance of social gaze and contingent social interaction for establishing joint commitments.

In conclusion, developmental and cross-species research strongly suggests a unique cognitive and motivational infrastruc- ture in humans which relies on sharing intentions and forming joint commitments in order to support cooperative joint actions. Consequently, specific reward-related and emotional neural cir- cuits might be expected to be involved. It is to the discussion of this topic that we now turn.

In this and the following section we will briefly introduce and discuss findings related to those two concepts in probably more elaborate forms of joint action. Neuroscientific research recently started to investigate the role of shared intentions in joint action, despite the methodological difficulties that almost naturally arise from inves- tigating complex notions of cooperation using neuroscientific methods Decety et al, ; de Bruijn et al. One viable approximation to testing shared intentions in cooperation includes biasing the participant's interpretation of a shared activity either toward a cooperative or toward a competitive setting.

This manipulation can be achieved when modifying the structure of the task, e. For example, in one of Decety's et al. Further, both conditions were compared to individual game performance. The contrast comparing joint and individual action revealed increased activation in the superior frontal gyrus, the superior parietal lobe and the anterior insula.

This pattern of neural activity might be attributed to higher executive demands of joint activ- ities for which coordination matters as compared to individual activities.

Additionally, this interpretation is consistent with find- ings from research on shared action and task representations. Yet, networks specifically more active during cooperative versus competitive conditions have also been identified.

During cooper- ation, the medial orbitofrontal cortex, the posterior cingulate, as well as bilateral anterior insula increased in activity. Consistent with above mentioned ideas, cooperation might thus be intrinsi- cally rewarding and might automatically raise expectations about ones partner. During competition, however, a network includ- ing the inferior parietal cortex, the medial prefrontal cortex and the superior frontal gyrus exhibited increased activity.

This pos- sibly reflects that not only cooperation but also competition encourages mental-state ascription reflecting the strategic aspects of competition, being a potential link to the game theoretic paradigms discussed above. Moreover, the aspect of processing unexpected events during cooperation, e. In two stud- ies by de Bruijn and colleagues de Bruijn et al, ; Radke et al, subjects played the so-called cannon-shooting game on a computer, either alone, in cooperation or in competition with their partner.

This game requires precisely aligning a can- non to hit a given target. Importantly, it was played in such a way that participants were aware of their own and the other's error. Cooperation versus competition was established by cou- pling versus decoupling participants' overall outcome based on their respective performance. The functional analyses revealed increased neural activity in the MPFC during cooperation as well as in competition when focusing on the observation of errors that only affected the other compared to errors that only affected oneself.

This result is interesting in the context of Decety's et al. Thus, during joint action the MPFC might subserve cognition associated with scanning potential threats to one's own plans and predicting the other's behavior, irrespectively of whether he or she is a competitor or rather an incompetent collaborator.

Frontiers in Human Neuroscience www. Therefore, the neural sub- strates of commitment-disclosing behavior during joint action introduced by Warneken, Tomasello and colleagues as impor- tant indicators of cooperation Tomasello et al, ; Warneken et al, ; Grafenhain et al, still remain unknown see Figure 2 for an illustration of the current neuroscientific coverage of psychological constructs in the context of cooperation. The phenomenon of joint attention has been intensely discussed by Tomasello and col- leagues Tomasello, ; Tomasello et al, ; Tomasello and Carpenter, as one important example for shared intention- ality, a bundle of abilities and motivations subserving the coor- dination and sharing of mental states.

These authors proposed that the evolution of human cooperation and culture critically depended on the emergence of joint attention. In a large num- ber of studies, they were able to show that children around their 9th month of age are able to make others follow their own gaze.

Moreover, most children learn to exactly discern what others want them to attend to when looking at various objects or at compound objects, which emphasizes the intentional nature of joint atten- tion see Carpenter et al, for an exhaustive treatment of the topic. Finally, recent research indicates that early joint atten- tion is even predictive of speech and theory of mind performance Aschersleben et al, Despite its importance, the neurosci- entific understanding of joint gaze and attention is still limited, which can be attributed to the methodological challenge of inves- tigating eye-movements and naturalistic social interaction in an fMRI environment.

In order to establish visual joint attention, in the experiment by Schilbach et al. This algorithm was then used to control the gaze interaction between the participant and a fictive confederate rep- resented by a virtual face, in a way that allowed for capturing gaze following and, ultimately, joint attention.

The neuroimag- ing analyses suggest an increased BOLD response in the ventral striatum during joint attention initiated by the participant, as compared to joint attention initiated by the other suggestive of an inherently rewarding experience associated with establishing a shared experience with another person.

This was corrobo- rated by making use of correlation analyses with a postscan pleasantness rating, which indicated that participants actually preferred looking at objects together with the virtual other, rather than alone.

Purely illustrative depiction that demonstrates to what extent different psychological facets underlying cooperative phenomena have up to now been subject to neuroimaging-based investigations The measures are based on the authors' subjective impression rather on any objective measure given general inconsistencies in nomenclature and diverging experimental settings throughout existing research on cooperation Frontiers in Human Neuroscience www.

First, the task involves externally unrewarded activity and thus the results are in favor of intrinsic rewards driving joint action. Second, joint attention is likely to involve shared intentions and can be understood as a very basic form of joint action Fiebich and Gallagher, ; Pfeiffer et al, Consequently, and in line with Schilbach et al.

However, further experiments are needed that clearly establish hypotheses about the intrinsic reward of joint attention in ruling out mere contingency or efficacy-experiences as potential confounds. To sum up, joint action research elucidates several aspects and preconditions of cooperation in the sense of acting together; some of them even tap into mutual helping and support.

It demon- strates individuals' sensitivity to coordinate, to establish common representations of their joint activity in cognitive and motiva- tional terms.

Neuroimaging studies complement these findings and corroborate psychological assumptions regarding the role of executive functions, reward processing, action mirroring and mentalizing in joint action. At the same time, it has to be stated that neuroimaging studies have not yet been able to fully capture the details revealed by developmental studies based on interaction-based methodologies.

In order to fill this gap, future neuroimaging paradigms should employ externally unrewarded social games and include naturalistic interactions that allow for mutual interventions reengagement attempts, criticism, teach- ing, reassuring as exceptions interruptions, problems, errors arise during joint action.

The latter point is of considerable sig- nificance in unveiling the agent's motives and experiences that govern their mutual cooperation, in other words telling to which extent agents cooperate when acting together.

Importantly, this is not just cosmetic in nature, as comparative studies reveal distinctions in ape and human cooperation only at this level.

The studies discussed up to this point capture many important facets of cooperation: striving for mutual benefit, acting together and supporting each other. Nevertheless, it becomes clear that the individual pieces of evidence are difficult to integrate with each other, as the different trends of cooperation research signif- icantly differ in their underlying key concepts decision making, joint action, shared intentionality and employed methods game theory, neuroimaging, action-based approaches.

In the following sections, we will discuss conceptual advances that might help to remedy this shortcoming. Concretely, the findings suggest mechanisms that invoke "social" utility func- tions, thus coupling personal with social welfare and encouraging cooperative choices by punishment of non-cooperative behavior.

In fact, research on social preferences taps into the motivational contingencies in human cooperation. Conceptually, social pref- erences fulfill a similar role as the joint commitments unveiled by comparative and developmental studies. Both approaches argue for the intrinsic nature of the mechanisms captured by their paradigms.

Yet, both perspectives suffer from their own limitations. Also, findings from comparative and developmental research are difficult to compare to findings from neuroeco- nomics. Developmental and comparative approaches convincingly rely on naturalistic joint action scenarios in which social interactions during instrumental and non-instrumental activities are com- pared.

Yet, research on social preferences relies on one-shot games in which the social interaction is reduced to making choices known to have consequences for the other's payoff. While the former seem to deemphasize formal analysis, the latter hardly seems to capture ecologically valid scenarios Schonberg et al, Studying decision-making using material payoffs thus seems more comparable to instrumental activities but not to the games employed by developmental studies.

Moreover, in the neu- roeconomic paradigms used to test social preferences selfish and social preferences usually form a potential conflict, at least in the light of the classical game theoretic framework. For example in the PD, uncooperative choices constitute the only Nash-equilibrium but are not Pareto efficient, indicating rational options that, if chosen by all players, would not yield the best possible outcome for all Myerson, Therefore, one may also call these games mixed-motive games.

This, however, does not hold for most of the joint action paradigms which may be more appropriately analyzed as com- mon interest games in which one Pareto superior Nash equi- librium exists Bacharach et al, implicating congruence between selfish and social preferences, hence, facilitating the for- mation of shard intentions.

One such game is the stag hunt game that can be traced back to a parable by Jean-Jacques Rousseau, in which two hunters can either conjointly hunt a stag together or hunt a hare separately.

Stags possess considerable strength and, therefore, hunting them requires successfully combining efforts, but also is most rewarding. Conversely, hunting a hare does not necessitate assistance of the other but yields only limited reward, whereas hunting a stag alone, constitutes the least efficient option see Figures 3A,B for a schematic depiction of the strategic inter- action captured by the stag hunt game.

From a cooperative stance, thus, the essence of this game is to coordinate each other's action toward hunting a stag together and only to choose hunting hares if it is indicated that hunting a stag is unlikely to succeed.

Another way of looking at this game is to regard the stag choice as payoff dominant, as compared to the risk dominant choice for hunting hares: players can gather sufficient knowledge about each other, e.

Contrastingly, a lack of such knowledge would make hunting hares more advisable. Therefore, proficient stag hunters should be equipped with dis- tinct abilities to assess the other's mental states or even establish shared intentions which should be easier in this strategic inter- action due to the absence of principled conflict between social and selfish motives. Interestingly, according to Tomasello and col- leagues Tomasello, ; Hamann et al, ;Rekers et al, intensified collaboration, is an essential feature of human cooper- ation that requires exactly theses capacities.

Therefore, typical stag Frontiers in Human Neuroscience www. Panel B gives an example of the stag hunt's payoff matrix. Analogous to the original description byпїЅJean-Jacques Rousseau, the stag yields a higher reward but can only be caught by both players'combined efforts, that is, by conjointly blocking the stag big gray square from two sides to prevent its escape. In this implementation of the game, the hares are realized as stationary objects small gray squares , referred to as "rabbits. Additionally, the round is over when a hunter caught a rabbit.

Similar to a board game players take turns moving their token between fields. Consequently, paradigms incorporating versions of the stag hunt game potentially with different pay-off schedules might help to combine the interactive features of Tomasello's and col- leagues' joint action paradigms with the analytic power provided by game theory. Neural evidence regarding stag hunt interactions, however, is currently rather sparse.

In a recent fMRI study comparing high- payoff choices in a stag hunt game and a differently framed but payoff identical lottery game, Ekins et al. These findings are in support the "social knowledge" account depicted above and implicate social-intentional processing to support cooperative decisions in stag hunt interactions.

However, the evi- dence is somewhat limited by the lack of whole-brain analyses. Moreover, the employed experimental setup largely resembled typical neuroeconomic paradigms in de-emphasizing the fine- grained details and interactive aspects common to joint action tasks. A novel approach combining the advantages of neuroeco- nomic and joint action methods has recently been developed by Yoshida and colleagues in a series of studies Yoshida et al, , a,b.

Here, the stag hunt parable was implemented almost literally as a hunting task, in which players move their respective hunter figure on a labyrinth-like grid to hunt stags or rabbits see Figure 3C. Interestingly this implementation of the game, entails continuous joint action moving one figure toward a target as well as strategic choice moving toward the rabbit versus moving toward the stag together with the other.

At the same time, the authors drew on the game theoretic basis of the paradigm, which lends itself to quantitative modeling. In their computational model of theory of mind, Yoshida et al. The model assumed that agents employ higher order belief inference in predicting whether their partner will cooperate, given their own behavior.

Moreover, the model issues a cooperation param- eter which estimates the probability at which an agent chooses to hunt a stag. This model was then compared to a fixed-strategy model which assumes a constant cooperation rate Yoshida et al, Applied to experimental data from the stag hunt paradigm, in which subjects played the hunting task together with a com- puter agent, the theory of mind model was significantly more predictive than the fixed-strategy model.

Interestingly, this only holds for healthy controls but not for subjects suffering from ASD, whose behavior is better characterized by the fixed-strategy model Yoshida et al, a. These results nicely illustrate the synergic potential of combining joint action paradigms and game theoretic modeling. This becomes even more evident, when this strategy is com- bined with neuroscientific methods. In their event-related fMRI study Yoshida et al. This approach not only allows inference about the brain areas involved in a given experimental task, but moreover allows to address hypotheses about the computational operations imple- mented in those areas.

The analyses revealed increased activity in the rostral mPFC during movements of the computer-agent as well as the activity in the bilateral ventral striatum.

This entices to speculate that mentalizing and reward processes, respectively, might be involved in performing the stag hunt game. Put into practice, the model-based approach allowed for understanding the activity in the rostral mPFC as a function of uncertainty of belief inferences and the activity in the ventral striatum as a func- tion of the outcome.

Additionally, increased activity found in the dorsolateral prefrontal cortex was shown to follow a model parameter that described how many levels of recursion where involved when thinking about another agent's. The modeling strategy employed in this study convincingly allows addressing specific hypotheses regarding the nature of the cognitive processes underlying cooperation and social interac- tion.

Complementary to Yoshida's ToM model, Braun et al. Further success in closing the loop between decision making, motor activity, and joint action is to be expected from future studies com- bining and exploiting related methodologies in a neuroimaging environment. Interestingly, the neural correlates underlying the payoff and agent-movement events closely matched those reported for the self-initiated joint attention episodes from Schilbach et al.

Although only speculation, this might indicate a common neural basis subserving stag hunt cooperation and basic social interac- tions, such as joint attention. This would support Tomasello's Frontiers in Human Neuroscience www. Concretely, one could investigate the neural responses to violations of expectation, that is, when subjects recognize the other's choice to hunt a hare individually versus to assume commitment to hunt a stag.

Manipulation of the others' reputation, sympathy or interpersonal responsiveness might help creating joint commitments and social expectations. Although more difficult to control, one might alternatively vary to what degree the players can communicate e.

Verbal responses to exceptions might then serve as additional indicators of the under- lying joint commitments. Thus, if implemented in an fMRI or a neurophysiological paradigm that elicits, e. Figure 4 illustrates the neuroimaging findings from the studies discussed in the course of this work.

The analysis of cooperation by Tuomela , and the theory of team-reasoning by Bacharach Bacharach, ; Bacharach et al. Further, both theoretical approaches are designed in a comprehensive fashion that allows for a wide range of applications.

In particular, Tuomela proposed that cooperation comprises all activities in which agents share and jointly pursue goals, whether specified as concrete ends involv- ing high levels of behavioral coordination or as group norms and ideals that do not exactly specify how to bring about the implicit common goal. Another important aspect of Tuomela's theory is the distinction between "i-mode" and "we-mode" coop- eration, which refers to the mindset involved in joint actions.

According to this idea, agents might construe their shared activ- ity as either involving commitment and giving rise to strong mutual expectationsпїЅa stance toward the joint action he terms we-mode. The color of the symbol indicates the publication from which the coordinates were drawn.

The shape of the symbol describes the study category. In the i-mode, agents might, for example, stop cooperating as soon as costs increase. Concretely, Tuomela argues that both aspects constitute inde- pendent dimensions yielding a taxonomy that allows describing the structure of cooperation on a task dimension dependence versus independence and a motivational dimension individual [i-mode] versus social [we-mode] commitments. Applied to our purposes, Tuomela's theoretical framework allows separating the different facets of cooperation discussed so far.

For example, the rabbit-game from Grafenhain's and colleagues , see above for discussion experiments constitute an independent task while at the same time involving a pronounced we-mode in the joint commitment condition1.

In contrast, in Newman-Norlund's and colleagues virtual-lifting task, which also was discussed above, the subjects' tasks are dependent while it remains unclear whether the subjects operate in a we-mode or in the i-mode. Consequently, "we-mode" and "i-mode" can be conceived as mindsets or schemes between which individuals can undulate, reflecting the ubiquitous fact that humans tend to cooperation selectively.

Based on a comprehensive review of the relevant lit- erature, Tuomela further contends that the we-mode is a uniquely human phenomenon that emerges early in ontogeny as indicated by developmental research reviewed above. However, while models have recently been proposed to incorporate these ideas into neuroscientific research Becchio and Bertone, ; Adenzato et al, ; Becchio et al, , this has so far hardly been put into practice. We propose that the we-mode theory might be a useful concept when it comes to systemiz- ing cooperation research.

Moreover, the specific assumptions of the we-mode theory are interesting by themselves and might help to design experiments that systematically tease apart dif- ferent psychological dimensions constitutive for the wide range of cooperative phenomena. Whether such a thing as the we- mode really exists is an empirical question. However, gath- ering systematic evidence for or against this concept might help to better understand whether different facets of cooper- ation discussed in this paper share a common psychological and neural ground.

Conversely, they might also constitute dis- tinct phenomena that are only associated by conventional use of language. The innovative potential of this direction of thought is fur- ther illustrated by Bacharach's Bacharach, ; Bacharach et al, theory of team-reasoning, which is highly related to Tuomela's we-mode theory. Accordingly, thinking can either operate individually, as analyzed by classical economic thought, or socially, as described by his alternative game theoretic model of team-reasoning.

When thinking as a team, individuals may overcome social dilemmas by modifying their frame of reasoning This at least holds for the child, whereas, the experimenter's motivation underlying his or her play might be due to his individual commitments to playing a larger paper-writing and publishing game, as kindly pointed out by one of the reviewers. It has to be added that this larger game can of course be played in a we-mode, as in the best case the authors constitute a team.

Even competition in a denned game implicates acknowledging the constitutive rules denning that game Tomasello, ; Searle, and hence implicates a minimum of cooperation. Moreover, Bacharach argues, that a skill to team-reason developed during human evo- lution and constitutes the key capacity underlying cooperative solutions of mixed-motive games such as the PD and common interest games as the stag hunt, again, suggesting a common biological basis for different kinds of cooperation.

Moreover, Bacharach's theory of we-thinking is ontologically more parsi- monious as team-reasoning is basically a consequence of framing which neither requires sophisticated mentalizing nor complex normative entities Bacharach et al, ; Pacherie, Such a theoretical framework thus more easily allows for including developmental and certain clinical populations lacking full-blown mentalizing capacities into the family of cooperators and joint actors.

Taken together, the benefit of a theoretical perspective in neuroscience acknowledging the notion of we-thinking, as high- lighted by Tuomela and Bacharach, would clearly provide a rich framework for cooperation research, whether focusing on decision making or joint action.

Paradigms capitalizing on this framework may thus constitute a promising direction to assess behavioral, goal-related, and motivational aspects of cooperation. Hence, it can and should be viewed from various angles and dis- sected by diverse scientific disciplines. Anthropological research emphasizes that social selection pressures have shaped human evolution and have led to the emergence of cooperative social systems that appear to be without parallel in the animal king- dom.

Experimental psychology has proposed the concept of joint action as one paramount aspect of human cooperation, which refers to the automatic synchronization of behavior during coordinated action execution. Comparative investigations stress that sharing mental states during cooperation is more prevalent in human children than in great apes and altruistic punish- ment is probably characteristic of human but not non-human primates. Cross-cultural studies likewise suggest that children's capacity and propensity for interpersonal cooperation is an inter- ethnically stable human trait.

Finally, the advances in imaging neuroscience have begun to allow mapping the neural corre- lates and brain networks that subserve decision-making during cooperation tasks.

However, neuroeconomic research on cooperation has so far been mostly based on a small number of paradigms that emphasize material payoffs and decision-making, hereby often disregarding other aspects of naturalistic cooperation. We con- tend, however, that the stag hunt game, describing a highly under-researched strategic interaction, lends itself to the inte- gration of game theory with findings from joint action research representing the interactive and embedded nature of coopera- tion.

Further progress in this area of research, we hold, will be made by employing stag hunt paradigms to link decision- making with other socio-cognitive momentums, such as joint Frontiers in Human Neuroscience www.

Needless to say, this move does not imply ignoring the computational and mathematical advances in neuroeconomic cooperation research. On the contrary, we believe that a model- based approach which potentially allows for regional mapping of computational mechanisms, will play an important role in the development of this field.

In this way, social- philosophical and economic proposals for taxonomically categorizing cooper- ative phonomena might be efficiently tested for neurobiological pertinence.

Taken together, employing more paradigms based on games and mutual benefits in neuroeconomics might help to link up with psychological research on cooperation, to go beyond mere decision-making aspects during cooperation and to pro- mote computational modeling in the context of ecologically valid cooperation-scenarios.

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