Academic Appointments

Research & Scholarship

Current Research and Scholarly Interests

Reward learning, decision-making, dopamine function


2014-15 Courses

Graduate and Fellowship Programs


Journal Articles

  • Dissociating Motivation from Reward in Human Striatal Activity JOURNAL OF COGNITIVE NEUROSCIENCE Miller, E. M., Shankar, M. U., Knutson, B., McClure, S. M. 2014; 26 (5): 1075-1084


    Neural activity in the striatum has consistently been shown to scale with the value of anticipated rewards. As a result, it is common across a number of neuroscientific subdiscliplines to associate activation in the striatum with anticipation of a rewarding outcome or a positive emotional state. However, most studies have failed to dissociate expected value from the motivation associated with seeking a reward. Although motivation generally scales positively with increases in potential reward, there are circumstances in which this linkage does not apply. The current study dissociates value-related activation from that induced by motivation alone by employing a task in which motivation increased as anticipated reward decreased. This design reverses the typical relationship between motivation and reward, allowing us to differentially investigate fMRI BOLD responses that scale with each. We report that activity scaled differently with value and motivation across the striatum. Specifically, responses in the caudate and putamen increased with motivation, whereas nucleus accumbens activity increased with expected reward. Consistent with this, self-report ratings indicated a positive association between caudate and putamen activity and arousal, whereas activity in the nucleus accumbens was more associated with liking. We conclude that there exist regional limits on inferring reward expectation from striatal activation.

    View details for DOI 10.1162/jocn_a_00535

    View details for Web of Science ID 000333627800012

  • Intertemporal choice as discounted value accumulation. PloS one Rodriguez, C. A., Turner, B. M., McClure, S. M. 2014; 9 (2)


    Two separate cognitive processes are involved in choosing between rewards available at different points in time. The first is temporal discounting, which consists of combining information about the size and delay of prospective rewards to represent subjective values. The second involves a comparison of available rewards to enable an eventual choice on the basis of these subjective values. While several mathematical models of temporal discounting have been developed, the reward selection process has been largely unexplored. To address this limitation, we evaluated the applicability of the Linear Ballistic Accumulator (LBA) model as a theory of the selection process in intertemporal choice. The LBA model formalizes the selection process as a sequential sampling algorithm in which information about different choice options is integrated until a decision criterion is reached. We compared several versions of the LBA model to demonstrate that choice outcomes and response times in intertemporal choice are well captured by the LBA process. The relationship between choice outcomes and response times that derives from the LBA model cannot be explained by temporal discounting alone. Moreover, the drift rates that drive evidence accumulation in the best-fitting LBA model are related to independently estimated subjective values derived from various temporal discounting models. These findings provide a quantitative framework for predicting dynamics of choice-related activity during the reward selection process in intertemporal choice and link intertemporal choice to other classes of decisions in which the LBA model has been applied.

    View details for DOI 10.1371/journal.pone.0090138

    View details for PubMedID 24587243

  • Family, community and clinic collaboration to treat overweight and obese children: Stanford GOALS-A randomized controlled trial of a three-year, multi-component, multi-level, multi-setting intervention CONTEMPORARY CLINICAL TRIALS Robinson, T. N., Matheson, D., Desai, M., Wilson, D. M., Weintraub, D. L., Haskell, W. L., McClain, A., McClure, S., Banda, J. A., Sanders, L. M., Haydel, K. F., Killen, J. D. 2013; 36 (2): 421-435


    To test the effects of a three-year, community-based, multi-component, multi-level, multi-setting (MMM) approach for treating overweight and obese children.Two-arm, parallel group, randomized controlled trial with measures at baseline, 12, 24, and 36months after randomization.Seven through eleven year old, overweight and obese children (BMI≥85th percentile) and their parents/caregivers recruited from community locations in low-income, primarily Latino neighborhoods in Northern California.Families are randomized to the MMM intervention versus a community health education active-placebo comparison intervention. Interventions last for three years for each participant. The MMM intervention includes a community-based after school team sports program designed specifically for overweight and obese children, a home-based family intervention to reduce screen time, alter the home food/eating environment, and promote self-regulatory skills for eating and activity behavior change, and a primary care behavioral counseling intervention linked to the community and home interventions. The active-placebo comparison intervention includes semi-annual health education home visits, monthly health education newsletters for children and for parents/guardians, and a series of community-based health education events for families.Body mass index trajectory over the three-year study. Secondary outcome measures include waist circumference, triceps skinfold thickness, accelerometer-measured physical activity, 24-hour dietary recalls, screen time and other sedentary behaviors, blood pressure, fasting lipids, glucose, insulin, hemoglobin A1c, C-reactive protein, alanine aminotransferase, and psychosocial measures.The Stanford GOALS trial is testing the efficacy of a novel community-based multi-component, multi-level, multi-setting treatment for childhood overweight and obesity in low-income, Latino families.

    View details for DOI 10.1016/j.cct.2013.09.001

    View details for Web of Science ID 000329265300012

  • Neural Correlates of Reinforcement Learning and Social Preferences in Competitive Bidding JOURNAL OF NEUROSCIENCE van den Bos, W., Talwar, A., McClure, S. M. 2013; 33 (5): 2137-2146


    In competitive social environments, people often deviate from what rational choice theory prescribes, resulting in losses or suboptimal monetary gains. We investigate how competition affects learning and decision-making in a common value auction task. During the experiment, groups of five human participants were simultaneously scanned using MRI while playing the auction task. We first demonstrate that bidding is well characterized by reinforcement learning with biased reward representations dependent on social preferences. Indicative of reinforcement learning, we found that estimated trial-by-trial prediction errors correlated with activity in the striatum and ventromedial prefrontal cortex. Additionally, we found that individual differences in social preferences were related to activity in the temporal-parietal junction and anterior insula. Connectivity analyses suggest that monetary and social value signals are integrated in the ventromedial prefrontal cortex and striatum. Based on these results, we argue for a novel mechanistic account for the integration of reinforcement history and social preferences in competitive decision-making.

    View details for DOI 10.1523/JNEUROSCI.3095-12.2013

    View details for Web of Science ID 000314351300036

    View details for PubMedID 23365249



    Psychological models of temporal discounting have now successfully displaced classical economic theory due to the simple fact that many common behavior patterns, such as impulsivity, were unexplainable with classic models. However, the now dominant hyperbolic model of discounting is itself becoming increasingly strained. Numerous factors have arisen that alter discount rates with no means to incorporate the different influences into standard hyperbolic models. Furthermore, disparate literatures are emerging that propose theoretical constructs that are seemingly independent of hyperbolic discounting. We argue that, although hyperbolic discounting provides an eminently useful quantitative measure of discounting, it fails as a descriptive psychological model of the cognitive processes that produce intertemporal preferences. Instead, we propose that recent contributions from cognitive neuroscience indicate a path for developing a general model of time discounting. New data suggest a means by which neuroscience-based theory may both integrate the diverse empirical data on time preferences and merge seemingly disparate theoretical models that impinge on time preferences.

    View details for DOI 10.1002/jeab.6

    View details for Web of Science ID 000319862600005

    View details for PubMedID 23344988

  • Pyrrhic victories: the need for social status drives costly competitive behavior. Frontiers in neuroscience van den Bos, W., Golka, P. J., Effelsberg, D., McClure, S. M. 2013; 7: 189-?


    Competitive behavior is commonly defined as the decision to maximize one's payoffs relative to others. We argue instead that competitive drive derives from a desire for social status. We make use of a multi-player auction task in which subjects knowingly incur financial losses for the sake of winning auctions. First, we show that overbidding is increased when the task includes members of a rival out-group, suggesting that social identity is an important mediator of competitiveness. In addition, we show that the extent that individuals are willing to incur losses is related to affective responses to social comparisons but not to monetary outcomes. Second, we show that basal levels of testosterone predict overbidding, and that this effect of testosterone is mediated by affective responses to social comparisons. Based on these findings, we argue that competitive behavior should be conceptualized in terms of social motivations as opposed to just relative monetary payoffs.

    View details for DOI 10.3389/fnins.2013.00189

    View details for PubMedID 24167468

  • Training Cognition in ADHD: Current Findings, Borrowed Concepts, and Future Directions NEUROTHERAPEUTICS Rutledge, K. J., van den Bos, W., McClure, S. M., Schweitzer, J. B. 2012; 9 (3): 542-558


    With both its high prevalence and myriad of negative outcomes, Attention-Deficit/Hyperactivity Disorder (ADHD) demands a careful consideration of the efficacy of its treatment options. Although the benefits of medication have a robust empirical background, nonpharmaceutical interventions evoke particular interest, as they are often viewed more favorably by parents. This review pays special attention to the use of working memory and recent cognitive training attempts in ADHD, describing its cognitive, behavioral, and biological effects in relation to current neurological theory of the disorder. While these treatments have demonstrated positive effects on some measures, there are limitations, as studies have failed to demonstrate generalization to critical measures, such as teacher-rated classroom behaviors, and have provided limited but growing evidence of functionally significant improvements in behavior. There is also a clear lack of research on the effects of training on reward systems and self-control. These limitations may be addressed by broadening the scope and procedures of the training and incorporating research concepts from other fields of study. First, it is important to consider the developmental trajectories of brain regions in individuals with the disorder, as they may relate to the effectiveness of cognitive training. Notions from behavioral economics, including delay discounting and framing (i.e., context) manipulations that influence present orientation, also have applications in the study of cognitive training in ADHD. In considering these other domains, we may find new ways to conceptualize and enhance cognitive training in ADHD and, in turn, address current limitations of interventions that fall in this category.

    View details for DOI 10.1007/s13311-012-0134-9

    View details for Web of Science ID 000308826800007

    View details for PubMedID 22911054

  • Theories of Willpower Affect Sustained Learning PLOS ONE Miller, E. M., Walton, G. M., Dweck, C. S., Job, V., Trzesniewski, K. H., McClure, S. M. 2012; 7 (6)


    Building cognitive abilities often requires sustained engagement with effortful tasks. We demonstrate that beliefs about willpower-whether willpower is viewed as a limited or non-limited resource-impact sustained learning on a strenuous mental task. As predicted, beliefs about willpower did not affect accuracy or improvement during the initial phases of learning; however, participants who were led to view willpower as non-limited showed greater sustained learning over the full duration of the task. These findings highlight the interactive nature of motivational and cognitive processes: motivational factors can substantially affect people's ability to recruit their cognitive resources to sustain learning over time.

    View details for DOI 10.1371/journal.pone.0038680

    View details for Web of Science ID 000305730900016

    View details for PubMedID 22745675

  • Are executive function and impulsivity antipodes? A conceptual reconstruction with special reference to addiction PSYCHOPHARMACOLOGY Bickel, W. K., Jarmolowicz, D. P., Mueller, E. T., Gatchalian, K. M., McClure, S. M. 2012; 221 (3): 361-387


    Although there is considerable interest in how either executive function (EF) or impulsivity relate to addiction, there is little apparent overlap between these research areas.The present paper aims to determine if components of these two constructs are conceptual antipodes--widely separated on a shared continuum.EFs and impulsivities were compared and contrasted. Specifically, the definitions of the components of EF and impulsivity, the methods used to measure the various components, the populations of drug users that show deficits in these components, and the neural substrates of these components were compared and contrasted.Each component of impulsivity had an antipode in EF. EF, however, covered a wider range of phenomena, including compulsivity.Impulsivity functions as an antipode of certain components of EF. Recognition of the relationship between EF and impulsivity may inform the scientific inquiry of behavioral problems such as addiction. Other theoretical implications are discussed.

    View details for DOI 10.1007/s00213-012-2689-x

    View details for Web of Science ID 000304615200001

    View details for PubMedID 22441659

  • The neural basis of cultural differences in delay discounting PHILOSOPHICAL TRANSACTIONS OF THE ROYAL SOCIETY B-BIOLOGICAL SCIENCES Kim, B., Sung, Y. S., McClure, S. M. 2012; 367 (1589): 650-656


    People generally prefer to receive rewarding outcomes sooner rather than later. Such preferences result from delay discounting, or the process by which outcomes are devalued for the expected delay until their receipt. We investigated cultural differences in delay discounting by contrasting behaviour and brain activity in separate cohorts of Western (American) and Eastern (Korean) subjects. Consistent with previous reports, we find a dramatic difference in discounting behaviour, with Americans displaying much greater present bias and elevated discount rates. Recent neuroimaging findings suggest that differences in discounting may arise from differential involvement of either brain reward areas or regions in the prefrontal and parietal cortices associated with cognitive control. We find that the ventral striatum is more greatly recruited in Americans relative to Koreans when discounting future rewards, but there is no difference in prefrontal or parietal activity. This suggests that a cultural difference in emotional responsivity underlies the observed behavioural effect. We discuss the implications of this research for strategic interrelations between Easterners and Westerners.

    View details for DOI 10.1098/rstb.2011.0292

    View details for Web of Science ID 000299897700003

    View details for PubMedID 22271781



    Rewards that are not immediately available are discounted compared to rewards that are immediately available. The more a person discounts a delayed reward, the more likely that person is to have a range of behavioral problems, including clinical disorders. This latter observation has motivated the search for interventions that reduce discounting. One surprisingly simple method to reduce discounting is an "explicit-zero" reframing that states default or null outcomes. Reframing a classical discounting choice as "something now but nothing later" versus "nothing now but more later" decreases discount rates. However, it is not clear how this "explicit-zero" framing intervention works. The present studies delineate and test two possible mechanisms to explain the phenomenon. One mechanism proposes that the explicit-zero framing creates the impression of an improving sequence, thereby enhancing the present value of the delayed reward. A second possible mechanism posits an increase in attention allocation to temporally distant reward representations. In four experiments, we distinguish between these two hypothesized mechanisms and conclude that the temporal attention hypothesis is superior for explaining our results. We propose a model of temporal attention whereby framing affects intertemporal preferences by modifying present bias.

    View details for DOI 10.1901/jeab.2011.96-363

    View details for Web of Science ID 000297609500005

    View details for PubMedID 22084496

  • Social Anxiety Modulates Risk Sensitivity through Activity in the Anterior Insula. Frontiers in neuroscience Tang, G. S., van den Bos, W., Andrade, E. B., McClure, S. M. 2011; 5: 142-?


    Decision neuroscience offers the potential for decomposing differences in behavior across individuals into components of valuation intimately tied to brain function. One application of this approach lies in novel conceptualizations of behavioral attributes that are aberrant in psychiatric disorders. We investigated the relationship between social anxiety and behavior in a novel socially determined risk task. Behaviorally, higher scores on a social phobia inventory (SPIN) among healthy participants were associated with an increase in risky responses. Furthermore, activity in a region of the dorsal anterior insula (dAI) scaled in proportion to SPIN score in risky versus non-risky choices. This region of the insula was functionally connected to areas in the intraparietal sulcus and anterior cingulate cortex that were related to decision-making across all participants. Overall, social anxiety was associated with decreased risk aversion in our task, consistent with previous results investigating risk taking in many everyday behaviors. Moreover, this difference was linked to the anterior insula, a region commonly implicated in risk attitudes and socio-emotional processes.

    View details for DOI 10.3389/fnins.2011.00142

    View details for PubMedID 22319462

  • An Expectation-Based Approach to Explaining the Crossmodal Influence of Color on Orthonasal Odor Identification: The Influence of Expertise CHEMOSENSORY PERCEPTION Shankar, M., Simons, C., Shiv, B., McClure, S., Spence, C. 2010; 3 (3-4): 167-173
  • An expectations-based approach to explaining the cross-modal influence of color on orthonasal olfactory identification: The influence of the degree of discrepancy ATTENTION PERCEPTION & PSYCHOPHYSICS Shankar, M., Simons, C., Shiv, B., McClure, S., Levitan, C. A., Spence, C. 2010; 72 (7): 1981-1993


    In the present study, we explored the conditions under which color-generated expectations influence participants' identification of flavored drinks. Four experiments were conducted in which the degree of discrepancy between the expected identity of a flavor (derived from the color of a drink) and the actual identity of the flavor (derived from orthonasal olfactory cues) was examined. Using a novel experimental approach that controlled for individual differences in color-flavor associations, we first measured the flavor expectations held by each individual and only then examined whether the same individual's identification responses were influenced by his or her own expectations. Under conditions of low discrepancy, the perceived disparity between the expected and the actual flavor identities was small. When a particular color--identified by participants as one that generated a strong flavor expectation--was added to these drinks (as compared with when no such color was added), a significantly greater proportion of identification responses were consistent with this expectation. This held true even when participants were explicitly told that color would be an uninformative cue and were given as much time as desired to complete the task. By contrast, under conditions of high discrepancy, adding the same colors to the drinks no longer had the same effect on participants' identification responses. Critically, there was a significant difference in the proportion of responses that were consistent with participants' color-based expectations in conditions of low as compared with high discrepancy, indicating that the degree of discrepancy between an individual's actual and expected experience can significantly affect the extent to which color influences judgments of flavor identity.

    View details for DOI 10.3758/APP.72.7.1981

    View details for Web of Science ID 000284449900024

    View details for PubMedID 20952794

  • Protocol for the Examination of Specimens From Patients With Non-Hodgkin Lymphoma/Lymphoid Neoplasms ARCHIVES OF PATHOLOGY & LABORATORY MEDICINE Hussong, J. W., Arber, D. A., Bradley, K. T., Brown, M. S., Chang, C. (., de Baca, M. E., Ellis, D. W., Foucar, K., Hsi, E. D., Jaffe, E. S., Lill, M., McClure, S. P., Medeiros, L. J., Perkins, S. L. 2010; 134 (6): E40-E47

    View details for Web of Science ID 000278642000001

    View details for PubMedID 20524855

  • Protocol for the Examination of Specimens From Patients With Hematopoietic Neoplasms of the Ocular Adnexa ARCHIVES OF PATHOLOGY & LABORATORY MEDICINE Bradley, K. T., Arber, D. A., Brown, M. S., Chang, C., Coupland, S. E., de Baca, M. E., Ellis, D. W., Foucar, K., Hsi, E. D., Jaffe, E. S., Lill, M. C., McClure, S. P., Medeiros, L. J., Perkins, S. L., Hussong, J. W. 2010; 134 (3): 336-340

    View details for Web of Science ID 000275529200003

    View details for PubMedID 20196660

  • The Wick in the Candle of Learning: Epistemic Curiosity Activates Reward Circuitry and Enhances Memory PSYCHOLOGICAL SCIENCE Kang, M. J., Hsu, M., Krajbich, I. M., Loewenstein, G., McClure, S. M., Wang, J. T., Camerer, C. F. 2009; 20 (8): 963-973


    Curiosity has been described as a desire for learning and knowledge, but its underlying mechanisms are not well understood. We scanned subjects with functional magnetic resonance imaging while they read trivia questions. The level of curiosity when reading questions was correlated with activity in caudate regions previously suggested to be involved in anticipated reward. This finding led to a behavioral study, which showed that subjects spent more scarce resources (either limited tokens or waiting time) to find out answers when they were more curious. The functional imaging also showed that curiosity increased activity in memory areas when subjects guessed incorrectly, which suggests that curiosity may enhance memory for surprising new information. This prediction about memory enhancement was confirmed in a behavioral study: Higher curiosity in an initial session was correlated with better recall of surprising answers 1 to 2 weeks later.

    View details for Web of Science ID 000268713200008

    View details for PubMedID 19619181

  • Patients with Schizophrenia have a Reduced Neural Response to Both Unpredictable and Predictable Primary Reinforcers NEUROPSYCHOPHARMACOLOGY Waltz, J. A., Schweitzer, J. B., Gold, J. M., Kurup, P. K., Ross, T. J., Salmeron, B. J., Rose, E. J., McClure, S. M., Stein, E. A. 2009; 34 (6): 1567-1577


    One prevalent theory of learning states that dopamine neurons signal mismatches between expected and actual outcomes, called temporal difference errors (TDEs). Evidence indicates that dopamine system dysfunction is involved in negative symptoms of schizophrenia (SZ), including avolition and anhedonia. As such, we predicted that brain responses to TDEs in dopamine midbrain nuclei and target areas would be abnormal in SZ. A total of 18 clinically stable patients with chronic SZ and 18 controls participated in an fMRI study, which used a passive conditioning task. In the task, the delivery of a small amount of juice followed a light stimulus by exactly 6 s on approximately 75% of 78 total trials, and was further delayed by 4-7 s on the remaining trials. The delayed juice delivery was designed to elicit the two types of TDE signals, associated with the recognition that a reward was omitted at the expected time, and delivered at an unexpected time. Main effects of TDE valence and group differences in the positive-negative TDE contrast (unexpected juice deliveries-juice omissions) were assessed through whole-brain and regions of interest (ROI) analyses. Main effects of TDE valence were observed for the entire sample in the midbrain, left putamen, left cerebellum, and primary gustatory cortex, bilaterally. Whole-brain analyses revealed group differences in the positive-negative TDE contrast in the right putamen and left precentral gyrus, whereas ROI analyses revealed additional group differences in the midbrain, insula, and parietal operculum, on the right, the putamen and cerebellum, on the left, and the frontal operculum, bilaterally. Further, these group differences were generally driven by attenuated responses in patients to positive TDEs (unexpected juice deliveries), whereas responses to negative TDEs (unexpected juice omissions) were largely intact. Patients also showed reductions in responses to juice deliveries on standard trials, and more blunted reinforcer responses in the left putamen corresponded to higher ratings of avolition. These results provide evidence that SZ patients show abnormal brain responses associated with the processing of a primary reinforcer, which may be a source of motivational deficits.

    View details for DOI 10.1038/npp.2008.214

    View details for Web of Science ID 000265221000019

    View details for PubMedID 19052540

  • The value of victory: social origins of the winner's curse in common value auctions JUDGMENT AND DECISION MAKING van den Bos, W., Li, J., Lau, T., Maskin, E., Cohen, J. D., Montague, P. R., McClure, S. M. 2008; 3 (7): 483-492


    Auctions, normally considered as devices facilitating trade, also provide a way to probe mechanisms governing one's valuation of some good or action. One of the most intriguing phenomena in auction behavior is the winner's curse - the strong tendency of participants to bid more than rational agent theory prescribes, often at a significant loss. The prevailing explanation suggests that humans have limited cognitive abilities that make estimating the correct bid difficult, if not impossible. Using a series of auction structures, we found that bidding approaches rational agent predictions when participants compete against a computer. However, the winner's curse appears when participants compete against other humans, even when cognitive demands for the correct bidding strategy are removed. These results suggest the humans assign significant future value to victories over human but not over computer opponents even though such victories may incur immediate losses, and that this valuation anomaly is the origin of apparently irrational behavior.

    View details for Web of Science ID 000260422900001

    View details for PubMedID 20305741

  • Anchors, scales and the relative coding of value in the brain CURRENT OPINION IN NEUROBIOLOGY Seymour, B., McClure, S. M. 2008; 18 (2): 173-178


    People are alarmingly susceptible to manipulations that change both their expectations and experience of the value of goods. Recent studies in behavioral economics suggest such variability reflects more than mere caprice. People commonly judge options and prices in relative terms, rather than absolutely, and display strong sensitivity to exemplar and price anchors. We propose that these findings elucidate important principles about reward processing in the brain. In particular, relative valuation may be a natural consequence of adaptive coding of neuronal firing to optimise sensitivity across large ranges of value. Furthermore, the initial apparent arbitrariness of value may reflect the brains' attempts to optimally integrate diverse sources of value-relevant information in the face of perceived uncertainty. Recent findings in neuroscience support both accounts, and implicate regions in the orbitofrontal cortex, striatum, and ventromedial prefrontal cortex in the construction of value.

    View details for DOI 10.1016/j.conb.2008.07.010

    View details for Web of Science ID 000260279400010

    View details for PubMedID 18692572

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