Cooperation based on reciprocal altruism has evolved in only a small number of species, yet it constitutes the core behavioral principle of human social life. The iterated Prisoner's Dilemma Game has been used to model this form of cooperation. We used fMRI to scan 36 women as they played an iterated Prisoner's Dilemma Game with another woman to investigate the neurobiological basis of cooperative social behavior. Mutual cooperation was associated with consistent activation in brain areas that have been linked with reward processing: nucleus accumbens, the caudate nucleus, ventromedial frontal/orbitofrontal cortex, and rostral anterior cingulate cortex. We propose that activation of this neural network positively reinforces reciprocal altruism, thereby motivating subjects to resist the temptation to selfishly accept but not reciprocate favors.

Functional MRI experiments in human subjects strongly suggest that the striatum participates in processing information about the predictability of rewarding stimuli. However, stimuli can be unpredictable in character (what stimulus arrives next), unpredictable in time (when the stimulus arrives), and unpredictable in amount (how much arrives). These variables have not been dissociated in previous imaging work in humans, thus conflating possible interpretations of the kinds of expectation errors driving the measured brain responses. Using a passive conditioning task and fMRI in human subjects, we show that positive and negative prediction errors in reward delivery time correlate with BOLD changes in human striatum, with the strongest activation lateralized to the left putamen. For the negative prediction error, the brain response was elicited by expectations only and not by stimuli presented directly; that is, we measured the brain response to nothing delivered (juice expected but not delivered) contrasted with nothing delivered (nothing expected).

A recent flurry of neuroimaging and decision-making experiments in humans, when combined with single-unit data from orbitofrontal cortex, suggests major additions to current models of reward processing. We review these data and models and use them to develop a specific computational relationship between the value of a predictor and the future rewards or punishments that it promises. The resulting computational model, the predictor-valuation model (PVM), is shown to anticipate a class of single-unit neural responses in orbitofrontal and striatal neurons. The model also suggests how neural responses in the orbitofrontal-striatal circuit may support the conversion of disparate types of future rewards into a kind of internal currency, that is, a common scale used to compare the valuation of future behavioral acts or stimuli.

The application of neuroimaging methods to product marketing -neuromarketing -has recently gained considerable popularity. We propose that there are two main reasons for this trend. First, the possibility that neuroimaging will become cheaper and faster than other marketing methods; and second, the hope that neuroimaging will provide marketers with information that is not obtainable through conventional marketing methods. Although neuroimaging is unlikely to be cheaper than other tools in the near future, there is growing evidence that it may provide hidden information about the consumer experience. The most promising application of neuroimaging methods to marketing may come before a product is even released -when it is just an idea being developed.Despite many common beliefs about the inherently evil nature of marketing, the main objective of marketing is to help match products with people. Marketing serves the dual goals of guiding the design and presentation of products such that they are more compatible with consumer preferences and facilitating the choice process for the consumer. Marketers achieve these goals by providing product designers with information about what consumers value and want before a product is created. After a product emerges on the marketplace, marketers attempt to maximize sales by guiding the menu of offerings, choices, pricing, advertising and promotions.In their attempts to provide these types of inputs, marketers use a range of market research techniques, from focus groups and individual surveys to actual market tests -with many approaches in between (see Supplementary information S1 (box)). In general, the simpler approaches (focus groups and surveys) are easy and cheap to implement but they provide data that can include biases, and are therefore seen as not very accurate 1-4. The approaches that are more complex and therefore harder to implement, such as market tests, provide more accurate data but incur a higher cost, and the product, production and distribution systems have to be in place for market tests to be conducted. There are some compromise approaches between

Certain classes of stimuli, such as food and drugs, are highly effective in activating reward regions. We show in humans that activity in these regions can be modulated by the predictability of the sequenced delivery of two mildly pleasurable stimuli, orally delivered fruit juice and water. Using functional magnetic resonance imaging, the activity for rewarding stimuli in both the nucleus accumbens and medial orbitofrontal cortex was greatest when the stimuli were unpredictable. Moreover, the subjects' stated preference for either juice or water was not directly correlated with activity in reward regions but instead was correlated with activity in sensorimotor cortex. For pleasurable stimuli, these findings suggest that predictability modulates the response of human reward regions, and subjective preference can be dissociated from this response.

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Intertemporal choices are decisions with consequences that play out over time. These choices range from the prosaichow much food to eat at a meal -to life-changing decisions about education, marriage, fertility, health behaviors and savings. Intertemporal preferences also affect policy debates about long-run challenges, such as global warming. Historically, it was assumed that delayed rewards were discounted at a constant rate over time. Recent theoretical and empirical advances from economic, psychological and neuroscience perspectives, however, have revealed a more complex account of how individuals make intertemporal decisions. We review and integrate these advances. We emphasize three different, occasionally competing, mechanisms that are implemented in the brain: representation, anticipation and self-control . 55 56 57 58 59 60 61 62 63 64 65 66 67 68 Economic, psychological and neuroscientific perspectives on intertemporal choice 70Intertemporal choices -decisions with consequences that play out over time -are important and ubiquitous. Decisions about spending, investments, diet, relationships, fertility, crime and education all contain intertemporal tradeoffs. In this paper, we discuss interrelated perspectives on intertemporal choice from the fields of economics, psychology and neuroscience. 73 74 75 76 77Until recently, the main contribution of economics to the study of intertemporal decisions was modeling. For nearly 80 years, economists have analyzed intertemporal decisions using the discounted utility (DU) model, which assumes that people evaluate the pleasures and pains resulting from a decision in much the same way that financial markets evaluate losses and gains, exponentially 'discounting' the value of outcomes according to how delayed they are in time. DU has been used to describe how people actually make intertemporal choices and it has been used as a tool for public policy. Policy decisions about how much to spend on research and development, health and education all depend on the discount rate used to analyze the decision. Indeed, recently the discount rate has proven to be a key parameter in the policy debate about global warming [1]. 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 The main contribution of psychology has been to identify, through empirical research, psychological mechanisms underlying intertemporal choice. For example, George Ainslie's research on the structure of time discounting posed the first serious challenge to the DU model -specifically to the assumption that people discount the future exponentially [2,3]. The concept of 'hyperbolic time discounting' (explained below) can be considered the first observed pattern of behavior that is inconsistent with DU -a DU 'anomaly'. Subsequent research by both psychologists and economists has identified a wide range of additional anomalies [4][5][6][7][8][9][10][11][12]. Economists have responded to these findings by constructing new models of intertemporal choice, which incorporate psychological insights,...

The mesolimbic dopaminergic system has long been known to be involved in the processing of rewarding stimuli, although recent evidence from animal research has suggested a more specific role of signaling errors in the prediction of rewards. We tested this hypothesis in humans, using functional magnetic resonance imaging (fMRI) and an operant conditioning paradigm for the discrete delivery of small quantities of fruit juice, along with a control experiment in which juice was substituted with a neutral visual stimulus. A local estimation of the activity in the ventral striatum showed a significant differentiation when the juice was withheld at the expected time of delivery; this finding was not replicated in the case of visual stimulation, providing evidence for time-locked processing of reward prediction errors in human ventral striatum.