Wellcome Principal Research Fellow, Professor of Neuroscience
Email: ws234@cam.ac.uk, Tel: +44 (0)1223 333 779, Fax: +44 (0)1223 333 840
Visiting Research Associate, California Institute of Technology, Pasadena
Visiting Professor, Tamagawa University, Tokyo
Fellow, Churchill College, Cambridge

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 first like to detect and characterise such signals before investigating more detailed neuronal mechanisms. We investigate fundamental reward and decision variables, including reward prediction errors, subjective reward value, utility, probability, risk and object-action-chosen value. We search for their signals in various brain structures, including dopamine neurons, striatum, frontal cortex and amygdala. We use concepts from animal learning theory and economic decision theory and combine behavioural, neurophysiological and neuroimaging (fMRI) methods. Please find a recent seminar lecture on this YouTube video, recent texts on reward and economic decisions in this brief overview or this longer review with its content list, or my CV or publication list.

Joining our laboratory: Please contact us for postdoctoral positions if you are enthusiastic about neuroscience research and interested in behaviour (experimental economics, experimental psychology) and neurophysiology or neuroimaging. Familiarity with a computer language (Matlab, C, Pascal or similar) and excellent writing skills are essential. We promise to provide the rest necessary for successful work, plus a vibrant lab and university atmosphere in a gorgeous city. Please email CV with list of experience and names of two references to ws234@cam.ac.uk.

Laboratory members
Alaa Al-Mohammad (BSc Univ Cambridge)
Alexandra Battaglia-Mayer (Prof Univ Rome)
Philipe Bujold (BA, BSc McGill Univ Montreal)
Mark Burrell (BSc Univ Auckland)
Charlotte R. van Coeverden (MSc Univ Amsterdam)
Helen Cousins (Computer Associate) (MSc Univ Cambridge)
Aled H. David (Lab manager) (BSc Univ Derby)
Kelly M J Diederen (PhD Univ Utrecht)
Simone Ferrari Toniolo (PhD Univ Rome)
Fabian Grabenhorst (PhD Univ Oxford)
Alexandre Pastor-Bernier (PhD Univ Montreal, MSc Univ Oslo)
Martin Vestergaard (PhD Tech Univ Copenhagen)
Konstantin Volkmann (MSc Univ Durham)
Leopold Zangemeister (BSc Univ Edinburgh)

Main collaborations
Ralph Adolphs, Charles R. Plott, Ueli Rütishauser (Caltech)
Christopher Harris (Economics Univ Cambridge)
Anthony Dickinson (Psychology Univ Cambridge)
Paul Fletcher (Psychiatry Univ Cambridge)
Florian Mormann (Univ Bonn)
Peter Bossaerts (Univ Melbourne)
Aldo Rustichini (Univ Minnesota)
Masamichi Sakagami (Tamagawa Univ)

Main funding
Wellcome Trust, European Research Council, NIH Conte Center Caltech, Cambridge University scholarships.

Prizes and honours
Thompson 99th percentile citations 2002-2012, Fellow EMBO 2014, Zülch Prize 2013, EJN FENS Award 2010, Fellow Royal Society 2009, Ipsen Prize 2005, Golden Brain Award 2002, Theodore-Ott-Prize 1997, Ellermann Prize 1984.

Named Lectures
EBBS Prize Lecture FENS Copenhagen 2016, Distinguished Lecture in Pharmacology Toronto 2015, Zülch Lecture Göttingen 2015, Qi Zhen Global Lecture Hangzhou 2014, Brenda Milner Lecture Montreal 2014, Woolsey Lecture Madison 2012, Eli Lilly Lecture 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 Leipzig 2006, Presidential Lecture Biol Psych Atlanta 2005, Pfizer Lecture SfN San Diego 2004, Special Lecture JNS Osaka 2004, Hans-Lukas-Teuber Lecture MIT 2001, NIH Director’s Lecture Bethesda 1999, EBBS ENA Lecture Strassbourg 1996, Brooks Lecture Harvard 1993.

Latest publications
• Tsutsui KI, Grabenhorst F, Kobayashi S, Schultz W. A dynamic code for economic object valuation in prefrontal cortex neurons. Nat Comm (in press) 2016
• Stauffer, Lak A, Yang A, Borel M, Paulsen O, Boyden E, Schultz W. Dopamine neuron-specific optogenetic stimulation. Cell 166, 1564-1571, 2016
• Genest W, Stauffer WR, Schultz W. Utility functions predict variance and skewness risk preferences. Proc Natl Acad Sci (USA) 113: 8402-8407, 2016
• 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 download pdf
• Schultz W. Dopamine reward prediction error signalling: a two-component response. Nature Rev Neurosci 17: 183-195, 2016 download pdf
• Schultz W. Neuronal reward and decision signals: from theories to data. Physiol Rev 95: 853-951, 2015 download 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 download pdf
• Diederen KMJ, Schultz W. Scaling prediction errors to reward variability benefits error-driven learning in humans. J Neurophysiol 114: 1628-1640, 2015 download pdf
• Vestergaard MD, Schultz W. Choice mechanisms for past, temporally extended outcomes. Proc Roy Soc B 282: 20141766, 1810 (10 pages), 2015 download pdf
• Hernadi I, Grabenhorst F, Schultz W. Planning activity for internally generated reward goals in amygdala neurons. Nat Neurosci 18: 461-469, 2015
• Stauffer WR, Lak A, Schultz W. Economic choices reveal probability distortion. J Neurosci 35: 3146-3154, 2015
• Stauffer WR, Lak A, Schultz W. Dopamine reward prediction error responses reflect marginal utility. Curr Biol 24: 2491-2500, 2014
• 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
• Kobayashi S, Schultz W. Reward contexts extend dopamine signals to unrewarded stimuli. Curr Biol 24: 56-62, 2014

Selected 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
• O'Neill, M, Schultz W. Risk prediction error coding in orbitofrontal neurons. J Neurosci 33: 15810-15814, 2013
• 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 download pdf
• Grabenhorst F, Hernadi I, Schultz W. Prediction of economic choice by amygdala neurons. Proc Natl Acad Sci (USA) 109: 18950-18955, 2012
• Bermudez M, Schultz W. Sensitivity to temporal reward structure in amygdala neurons. Curr Biol 22: 1839-1844, 2012
• 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 download pdf
• Schultz W. Potential vulnerabilities of neuronal reward, risk, and decision mechanisms to addictive drugs. Neuron 69: 603-617, 2011
• 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
• Burke CJ, Tobler PN, Baddeley M, Schultz W. Neuronal mechanisms of observational learning. Proc Natl Acad Sci (USA) 107: 14431-14436, 2010 download pdf, suppl mat pdf
• Schultz W. Dopamine signals for reward value and risk: basic and recent data. Behav Brain Funct 6:24, 2010 download 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
• Kobayashi S, Pinto de Carvalho O, Schultz W. Adaptation of reward sensitivity in orbitofrontal neurons. J Neurosci 30: 534-544, 2010
• 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 download pdf, suppl mat 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 download 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 download pdf
• Kobayashi S, Schultz W. Influence of reward delays on responses of dopamine neurons. J Neurosci 28: 7837-7846, 2008
• 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 download 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 download pdf
• Schultz W. Multiple dopamine functions at different time courses. Ann Rev Neurosci 30: 259-288, 2007 download pdf
• Schultz W. Behavioral theories and the neurophysiology of reward. Ann Rev Psychol 57: 87-115, 2006 download pdf, Podcast
• Tobler PN, Fiorillo CD, Schultz W. Adaptive coding of reward value by dopamine neurons. Science 307: 1642-1645, 2005
• Fiorillo CD, Tobler PN, Schultz W. Discrete coding of reward probability and uncertainty by dopamine neurons. Science 299: 1898-1902, 2003
• Waelti P, Dickinson A, Schultz W. Dopamine responses comply with basic assumptions of formal learning theory. Nature 412: 43-48, 2001
• Tremblay L, Schultz W. Relative reward preference in primate orbitofrontal cortex. Nature 398: 704-708, 1999
• Schultz W. Predictive reward signal of dopamine neurons. J Neurophysiol. 80: 1-27, 1998
• Hollerman JR, Schultz W. Dopamine neurons report an error in the temporal prediction of reward during learning. Nature Neurosci 1: 304-309, 1998
• Schultz W, Dayan P, Montague RR. A neural substrate of prediction and reward. Science 275: 1593-1599, 1997
• Mirenowicz J, Schultz W. Preferential activation of midbrain dopamine neurons by appetitive rather than aversive stimuli. Nature 379: 449-451, 1996
• 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
• 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
• 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
• Schultz W. Responses of midbrain dopamine neurons to behavioral trigger stimuli in the monkey. J Neurophysiol 56: 1439-1462, 1986