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Department of Physiology, Development and Neuroscience



Reward processing in the brain

Our group is interested in identifying brain signals for reward and economic decisions. As information processing systems work with explicit signals, we like to identify and characterise such signals before investigating detailed neuronal mechanisms. We use concepts from animal learning theory and economic decision theory and combine behavioural, neurophysiological and neuroimaging (fMRI) methods. We search for neuronal responses that implement fundamental theoretical constructs underlying reward-seeking, learning and decision-making, such as reward prediction error, utility, probability, risk, object-action-chosen value, and revealed preference. Studied brain structures include dopamine neurons, striatum, frontal cortex and amygdala. Please find more information from a seminar lecture on this YouTube video, a short general article or update on dopamine reward prediction error coding, a brief overview or longer review on reward and economic decisions, or my CV, publication list, personal website and biography (SfN).

Laboratory members
Leo Chi U Seak (BSc Univ Hongkong)
Aled H. David (Lab manager) (BSc Univ Derby)
Simone Ferrari Toniolo (PhD Univ Rome)
Daniel F. Hill (PhD Univ Arizona)
Arkadiusz Stasiak (Computer Associate) (PhD Univ Lodz)

Main collaborations
Ralph Adolphs, Charles R. Plott, Ueli Rütishauser (Caltech)
Peter Bossaerts (Economics Univ Cambridge)
Anthony Dickinson (Psychology Univ Cambridge)
Paul Fletcher (Psychiatry Univ Cambridge)
Fabian Grabenhorst (Psychology Univ Oxford) with Fei-Yang Huang and Tavish Traut
Christopher Harris (Economics Univ Cambridge)
Masamichi Sakagami (Tamagawa Univ)

Main recent funding
Wellcome Trust, European Research Council (ERC), NIMH (Caltech Conte Center), Human Frontiers, Cambridge University studentships.

Prizes and honours
Lashley Award (Am Philos Soc) 2019, Gruber Prize 2018, Brain Prize 2017, Fellow EMBO 2014, Zülch Prize 2013, Thompson 99th % citations 2002-2012, EJN FENS Award 2010, Fellow Royal Society 2009, Ipsen Prize 2005, Golden Brain Award 2002, Theodore-Ott-Prize 1997, Ellermann Prize 1984.

Named Lectures
Hill Lecture Cambridge Univ 2021, FENS Eastern Europe Lecture (virtual) 2021, Skinner Lecture Assoc Behav Analysis Int (virtual) 2020, Waelsch Lecture Columbia Univ 2019, Swammerdam Lecture Utrecht 2018, Gruber Prize Lecture SfN San Diego 2018, Zülch Lecture Berlin 2018, Keynote Japanese Psychol Assoc Sendai 2018, Volker Henn Lecture Univ Zürich 2018, Brain Prize Lecture FENS Berlin 2018, Ruysch Lecture Univ Amsterdam 2018, Terry Lecture WashU St. Louis 2018, Erlanger Lecture Exp Biol San Diego 2018, Sprague Lecture UPenn Philadelphia 2018, Dean for Science Lecture NYU 2018, Caltech Chen Lecture 2017, Yale Endowed Lecture 2017, Colegio Nacional Mexico 2016, Plenary Lectures Int Congr Psychol and JNS Yokohama 2016, EBBS Prize Lecture FENS Copenhagen 2016, Distinguished Lecture Pharmacology Univ Toronto 2015, Zülch Lecture Göttingen 2015, Qi Zhen Global Lecture Hangzhou 2014, Brenda Milner Lecture McGill Univ Montreal 2014, Ting-Kai Li Lecture San Francisco 2012, Woolsey Lecture Madison 2012, Eli Lilly Lecture Univ Montreal 2011, Mars Lecture Florida 2011, Kavli Lecture Soc Neuroecon Evanston 2010, Roger Brown Louks Lecture Seattle 2009, Netherlands Cognition Lecture Amsterdam 2008, Paul Flechsig Lecture Univ Leipzig 2006, Presidential Lecture Biol Psych Atlanta 2005, Pfizer Lecture SfN San Diego 2004, Special Lecture JNS Osaka 2004, Teuber Lecture MIT 2001, NIH Director's Lecture Bethesda 1999, EBBS ENA Lecture Strassbourg 1996, Brooks Lecture Harvard 1993.

Postdoctoral scientists supervised
Alexandre Pastor-Bernier 2013-2021, Armin Lak 2013-15, Raymundo Báez-Mendoza 2013-15, Kelly M.J. Diederen 2012-15, Martin D. Vestergaard 2010-17, Fabian Grabenhorst 2009-17, William R. Stauffer 2009-15, Maria A. Bermudez 2005-09, Shunsuke Kobayashi 2005-10, Martin O’Neill 2005-14, Philippe Tobler 2004-09, Ofelia Carvalho 2002-07, Ken-ichiro Tsutsui 2002-05, Istvan Hernadi 2002-05, Christopher D. Fiorillo 1999-2003, Bertram Gerber 1999-2001, Oum K. Hassani 1997-99, Howard C. Cromwell 1996-99, Roland R. Suri 1995-97, Léon Tremblay 1992-96, Jeffrey R. Hollerman 1992-96, Paul Apicella 1987-1990, Tomas Ljungberg 1987-1990, Ranulfo Romo 1985-1987.

PhD students supervised
Mark Burrell 2020, Philipe Bujold 2020, Konstantin Volkmann 2019, Alaa Al-Mohammad 2018, Leopold Zangemeister 2017, Charlotte van Coeverden 2017, Wilfried Genest 2016, Armin Lak 2013, Raymundo Báez-Mendoza 2013, Ryan Haynes 2011, Christopher Burke 2010, Krishna Prasad Miyapuram 2009, George Christopoulos 2008, Philippe Tobler 2004, Pascale Waelti 2000, Jacques Mirenowicz 1995, Andreas Studer 1989, Din Lam Toan 1986, Patrick Aebischer 1983, André Ruffieux 1980.









Key publications: 

Burrell M, Pastor-Bernier A, Schultz W. Worth the work? Monkeys discount rewards by a subjective adapting effort cost. bioRxiv 523384, 2023. SSRN 4321740, 2023.
van Coeverden C, Schultz W. Formal cooperation in macaque monkeys obtained via single-payoff change from formal coordination games. bioRxiv 521899, 2022. SSRN 4321728, 2023. (pdf)
Pastor-Bernier A, Volkmann K, Seak LCU, Stasiak A, Plott CR, Schultz W. Concept-based neural investigations of multi-component economic choice in human and non-human primates. SSRN 4295148, 2022. (pdf)
• Ferrari-Toniolo S, Schultz W. Reliable population signal of subjective economic value from unreliable neurons in primate orbitofrontal cortex. bioRxiv 468353, 2022. (pdf)
Schultz W. My Biography. Society for Neuroscience (SfN) 2022. (pdf)
• Ferrari-Toniolo S, Seak LCU, Schultz W. Risky choice: probability weighting explains Independence Axiom violations in monkeys. J Risk Uncertain ( 2022. (pdf)
• Al-Mohammad A, Schultz W. Reward value revealed by auction. J Neurosci 42: 1510-1528, 2022. (pdf)
• Bujold PM, Ferrari-Toniolo S, Seak LCU, Schultz W. Comparing utility between risky and riskless choice in rhesus monkeys. Anim Cog 25: 385-399, 2022. (pdf)
• Pastor-Bernier AM, Stasiak A, Schultz W. Reward-specific satiety affects subjective value signals in orbitofrontal cortex during multi-component choice. Proc Nat Acad Sci (USA) 118: e2022650118, 2021. (pdf)
• Bujold PM, Ferrari-Toniolo S, Schultz W. Adaptation of utility functions functions to reward distribution in rhesus monkeys. Cognition 214: 104764, 2021. (pdf)
• Grabenhorst F, Schultz W. Functions of primate amygdala neurons in economic decisions and social decision simulation. Behav Brain Res 409: 113318, 2021. (pdf)
• Schultz W, Stauffer WR, Lak A. Pastor-Bernier A. Smarter than humans: rationality reflected in primate neuronal signals. Curr Op Behav Sci 41: 51-56, 2021. (pdf)
• Chi U Seak* L, Volkmann* K, Pastor-Bernier A, Grabenhorst F, Schultz W. Single-dimensional human brain signals for two-dimensional economic choice options. J Neurosci 41: 3000-3013, 2021. (pdf)
• Ferrari-Toniolo S, Bujold P, Schultz W. Non-human primates satisfy utility maximization in compliance with the continuity axiom of Expected Utility Theory. J Neurosci 41: 2964-2979, 2021. (pdf)
• Tremblay S, Acker L, Afraz A, Albaugh DL, Amita H, Andrei AR, Angelucci A, Aschner A, Balan PF, Basso MA, Benvenuti G, Bohlen MO, Caiola MJ, Calcedo R, Cavanaugh J, Chen Y, Chen S, MM, Clark AM, Dai J, Debes SR, Deisseroth K, Desimone R, Dragoi V, Egger SW, Eldridge MAG, El-Nahal HG, Fabbrini F, Federer F, Fetsch CR, Fortuna MG, Friedman RM, Fujii N, Gail A, Galvan A, Ghosh S, Gieselmann MA, Gulli RA, Hikosaka O, Hosseini EA, Hu X, Hüer J, Inoue K-I, Janz R, Jazayeri M, Jiang R, Ju N, Kar K, Klein C, Kohn A, Komatsu M, Maeda K, Martinez-Trujillo JC, Matsumoto M, Maunsell JHR, Mendoza-Halliday D, Monosov IE, Muers RS, Nurminen L, Ortiz-Rios M, O’Shea DJ, Palfi S, Petkov CI, Pojoga S, Rajalingham R, Ramakrishnan C, Remington ED, Revsine C, Roe AW, Sabes PN, Saunders RC, Scherberger H, Schmid MC, Schultz W, Seidemann E, Senova Y-S, Shadlen MN, Sheinberg DL, Siu C, Smith Y, Solomon SS, Sommer MA, Spudich JL, Stauffer WR, Takada M, Tang S, Thiele A, Treue S, Vanduffel W, Vogels R, Whitmire MP, Wichmann T, Wurtz RH, Xu H, Yazdan-Shahmorad A, Krishna V. Shenoy KV, DiCarlo JJ, Platt ML. An Open Resource for Non-human Primate Optogenetics. Neuron 108: 1075-1090, 2020. pdf
• Vestergaard MD, Schultz W. Retrospective valuation of experienced outcome encoded in distinct reward representations in the anterior insula and amygdala. J Neurosci 40: 8938-8950, 2020. pdf
• Pastor-Bernier A*, Volkmann K*, Stasiak A, Grabenhorst F, Schultz W. Experimentally revealed stochastic preferences for multicomponent choice options. J exp Psych: Anim Learn Cog 46: 367-384, 2020. pdf
• Grabenhorst F, Salzman D, Schultz W. The role of the primate amygdala in reward and decision-making. In: The Cognitive Neurosciences (Eds. Poeppel D, Mangun GR, Gazzaniga MS) pp 631-639, 2020.
• Stauffer WR, Schultz W. Dopamine prediction error responses reflect economic utility. In: The Cognitive Neurosciences (Eds. Poeppel D, Mangun GR, Gazzaniga MS) pp 587-595, 2020.
• Pastor-Bernier A, Stasiak A, Schultz W. Orbitofrontal signals for two-component choice options comply with indifference curves of Revealed Preference Theory. Nat Comm 10: 4885, 2019 (57 pages). pdf
• Schultz W. Recent advances in understanding the role of phasic dopamine activity. F1000, 2019. pdf
• Grabenhorst* F, Tsutsui* KI, Kobayashi S, Schultz W. Primate prefrontal neurons signal economic risk derived from the statistics of recent reward experience. eLife 8: e44838, 2019 (30 pages). pdf
• Grabenhorst F, Báez-Mendoza R, Genest W, Deco G, Schultz W. Primate amygdala neurons simulate decision processes of social partners. Cell 177: 986-998, 2019. pdf
• Ferrari-Toniolo S, Bujold P, Schultz W. Probability distortion depends on choice sequence in rhesus monkeys. J Neurosci 39: 2915-2929, 2019. pdf
• Brzosko Z, Zannone S, Schultz W, Clopath C, Paulsen O. Sequential neuromodulation of Hebbian plasticity offers mechanism for effective reward-based navigation. eLife 6:e27756, 2017 (18 pages). pdf
• Schultz W, Stauffer WR, Lak A. The phasic dopamine signal maturing: from reward via behavioural activation to formal economic utility. Curr Op Neurobiol 43: 139-148, 2017.  pdf
• Pastor Bernier A, Plott CR, Schultz W. Monkeys choose as if maximizing utility compatible with basic principles of revealed preference theory. Proc Natl Acad Sci (USA) 114: E1766-E1775, 2017. pdf
• Báez-Mendoza R, Schultz W. Performance error-related activity in monkey striatum during social interactions. Sci Reports 6: 37199, 2016 (8 pages).
• Lak A, Stauffer WR, Schultz W. Dopamine neurons learn relative chosen value from probabilistic rewards. eLife 5: e18044, 2016 (19 pages). pdf
• Zangemeister* L, Grabenhorst* F, Schultz W. Neural basis for economic saving strategies in human amygdala-prefrontal reward circuits. Curr Biol 26: 3004-3013, 2016.  pdf
• Grabenhorst* F, Hernadi* I, Schultz W. Primate amygdala neurons evaluate the progress of self-defined economic choice sequences. eLife 5: e18731, 2016 (24 pages). pdf
• Tsutsui* KI, Grabenhorst* F, Kobayashi S, Schultz W. A dynamic code for economic object valuation in prefrontal cortex neurons. Nat Comm 7: 12554, 2016 (14 pages). pdf
• Stauffer, Lak A, Yang A, Borel M, Paulsen O, Boyden E, Schultz W. Dopamine neuron-specific optogenetic stimulation. Cell 166, 1564-1571, 2016. pdf
• Genest W, Stauffer WR, Schultz W. Utility functions predict variance and skewness risk preferences. Proc Natl Acad Sci (USA) 113: 8402-8407, 2016. pdf
• Diederen K, Spencer T, Vestergaard MD, Fletcher P, Schultz W. Adaptive prediction error coding in the human midbrain and striatum facilitates behavioral adaptation and learning efficiency. Neuron 90: 1127-1138, 2016. pdf
• Schultz W. Dopamine reward prediction error signalling: a two-component response. Nature Rev Neurosci 17: 183-195, 2016. pdf
• Schultz W. Neuronal reward and decision signals: from theories to data. Physiol Rev 95: 853-951, 2015. pdf (or with its content list)
• Schultz W, Carelli RM, Wightman RM. Phasic dopamine signals: from subjective reward value to formal economic utility. Curr Op Behav Sci 5: 147-154, 2015. pdf
• Hernadi* I, Grabenhorst* F, Schultz W. Planning activity for internally generated reward goals in amygdala neurons. Nat Neurosci 18: 461-469, 2015. pdf
• Stauffer WR, Lak A, Bossaerts P, Schultz W. Economic choices reveal probability distortion. J Neurosci 35: 3146-3154, 2015. pdf
• Stauffer WR, Lak A, Schultz W. Dopamine reward prediction error responses reflect marginal utility. Curr Biol 24: 2491-2500, 2014. pdf
• Lak A, Stauffer WR, Schultz W. Dopamine prediction error responses integrate subjective value from different reward dimensions. Proc Natl Acad Sci (USA) 111: 2343-2348, 2014. pdf
• Kobayashi S, Schultz W. Reward contexts extend dopamine signals to unrewarded stimuli. Curr Biol 24: 56-62, 2014.

Further publications
• Báez-Mendoza R, Harris C, Schultz W. Activity of striatal neurons reflects social action and own reward. Proc Natl Acad Sci (USA) 110: 16634-16639, 2013. pdf
• O'Neill, M, Schultz W. Risk prediction error coding in orbitofrontal neurons. J Neurosci 33: 15810-15814, 2013. pdf
• d'Acremont M, Schultz W, Bossaerts P. The human brain encodes event frequencies while forming subjective beliefs. J Neurosci 33: 10887-10897, 2013.
• Schultz W. Updating dopamine reward signals. Curr Op Neurobiol 23: 229-238, 2013. pdf
• Grabenhorst* F, Hernadi* I, Schultz W. Prediction of economic choice by amygdala neurons. Proc Natl Acad Sci (USA) 109: 18950-18955, 2012. pdf
• Bermudez M, Schultz W. Sensitivity to temporal reward structure in amygdala neurons. Curr Biol 22: 1839-1844, 2012. pdf
• Hare TA, Schultz W, Camerer CF, O'Doherty JP, Rangel A. Transformation of stimulus value signals into motor commands during simple choice. Proc Natl Acad Sci (USA) 108: 18120-18125, 2011. pdf
• Schultz W. Potential vulnerabilities of neuronal reward, risk, and decision mechanisms to addictive drugs. Neuron 69: 603-617, 2011. pdf
• O'Neill M, Schultz W. Coding of reward risk by orbitofrontal neurons is mostly distinct from coding of reward value. Neuron 68: 789-800, 2010. pdf
• Burke CJ, Tobler PN, Baddeley M, Schultz W. Neuronal mechanisms of observational learning. Proc Natl Acad Sci (USA) 107: 14431-14436, 2010. pdf
• Schultz W. Dopamine signals for reward value and risk: basic and recent data. Behav Brain Funct 6:24, 2010. pdf
• Bermudez MA, Schultz W. Responses of amygdala neurons to positive reward predicting stimuli depend on background reward (contingency) rather than stimulus-reward pairing (contiguity). J Neurophysiol 103: 1158-1170, 2010. pdf
• Kobayashi S, Pinto de Carvalho O, Schultz W. Adaptation of reward sensitivity in orbitofrontal neurons. J Neurosci 30: 534-544, 2010. pdf
• Tobler PN, Christopoulos GI, O'Doherty JP, Dolan RJ, Schultz W. Risk-dependent reward value signal in human prefrontal cortex. Proc Natl Acad Sci (USA) 106: 7185-7190, 2009. pdf
• Gregorios-Pippas L, Tobler PN, Schultz W. Short term temporal discounting of reward value in human ventral striatum. J Neurophysiol 101: 1507-1523, 2009. pdf
• Christopoulos GI, Tobler PN, Bossaerts P, Dolan RJ, Schultz W. Neural correlates of value, risk, and risk aversion contributing to decision making under risk. J Neurosci. 29: 12574-12583, 2009. pdf
• Kobayashi S, Schultz W. Influence of reward delays on responses of dopamine neurons. J Neurosci 28: 7837-7846, 2008. pdf
• Hare TA, O’Doherty J, Camerer CF, Schultz W, Rangel A. Dissociating the role of the orbitofrontal cortex and the striatum in the computation of goal values and prediction errors. J Neurosci 28: 5623-5630, 2008. pdf
• Tobler PN, O'Doherty JP, Dolan R, Schultz W. Reward value coding distinct from risk attitude-related uncertainty coding in human reward systems. J Neurophysiol 97: 1621-1632, 2007. pdf
• Schultz W. Multiple dopamine functions at different time courses. Ann Rev Neurosci 30: 259-288, 2007. pdf
• Schultz W. Behavioral theories and the neurophysiology of reward. Ann Rev Psychol 57: 87-115, 2006. pdf, Podcast
• Tobler PN, Fiorillo CD, Schultz W. Adaptive coding of reward value by dopamine neurons. Science 307: 1642-1645, 2005. pdf
• Fiorillo CD, Tobler PN, Schultz W. Discrete coding of reward probability and uncertainty by dopamine neurons. Science 299: 1898-1902, 2003. pdf
• Waelti P, Dickinson A, Schultz W. Dopamine responses comply with basic assumptions of formal learning theory. Nature 412: 43-48, 2001. pdf
• Tremblay L, Schultz W. Relative reward preference in primate orbitofrontal cortex. Nature 398: 704-708, 1999. pdf
• Schultz W. Predictive reward signal of dopamine neurons. J Neurophysiol. 80: 1-27, 1998. pdf
• Hollerman JR, Tremblay L, Schultz W. Influence of reward expectation on behavior-related neuronal activity in primate striatum. J Neurophysiol 80: 947-963, 1998. pdf
• Hollerman JR, Schultz W. Dopamine neurons report an error in the temporal prediction of reward during learning. Nature Neurosci 1: 304-309, 1998. pdf
• Schultz W, Dayan P, Montague RR. A neural substrate of prediction and reward. Science 275: 1593-1599, 1997. pdf
• Mirenowicz J, Schultz W. Preferential activation of midbrain dopamine neurons by appetitive rather than aversive stimuli. Nature 379: 449-451, 1996. pdf
• Schultz W, Apicella P, Ljungberg T. Responses of monkey dopamine neurons to reward and conditioned stimuli during successive steps of learning a delayed response task. J Neurosci 13: 900-913, 1993. pdf
• Schultz W, Romo R. Role of primate basal ganglia and frontal cortex in the internal generation of movements: I. Preparatory activity in the anterior striatum. Exp Brain Res 91: 363-384, 1992. pdf
• Schultz W, Romo R. Dopamine neurons of the monkey midbrain: Contingencies of responses to stimuli eliciting immediate behavioral reactions. J Neurophysiol 63: 607-624, 1990. pdf
• Romo R, Schultz W. Dopamine neurons of the monkey midbrain: Contingencies of responses to active touch during self-initiated arm movements. J Neurophysiol 63: 592-606, 1990. pdf
• Schultz W. Responses of midbrain dopamine neurons to behavioral trigger stimuli in the monkey. J Neurophysiol 56: 1439-1462, 1986. pdf


Teaching and Supervisions


Neurobiology of Reward, Dept PDN Univ Cambridge (part II undergraduate)

Experimental Approaches in Brain Research, Dept PDN Univ Cambridge (part II undergraduate)

Neuroeconomics of Reward, Faculty of Economics Univ Cambridge (graduate)

Neuronal Reinforcement Learning, University College London (graduate)

Medical Sciences supervisions Churchill College 2005-2012 (part IB undergraduate)

Wellcome Principal Research Fellow
Professor of Neuroscience
Fellow of Churchill College Cambridge
Visiting Research Associate Caltech
Picture of Prof Wolfram   Schultz

Contact Details

Department of Physiology, Development & Neuroscience
University of Cambridge
Downing Street
Phone +44 (0)1223 333 779
Fax +44 (0)1223 333 840
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