Professor Wolfram Schultz


Our group is interested to relate the mechanics of brain activity to measurable behaviour. We combine neurophysiological, imaging and behavioural techniques to investigate the neural correlates of goal-directed. We are interested in outcome value (in particular reward) signals in specific brain structures such as dopamine neurons, striatum, frontal cortex and amygdala. These rapid, global, evaluative and supervising neural signals may play a role in decision-making and choice behaviour. In investigating these outcome-coding mechanisms we try to establish a common biological basis for animal learning theory, microeconomic utility and game theories, and behavioural ecology. The larger background is to consider the brain in its capacity for processing reward information as an essential organ for assuring the fitness for survival in an evolutionary framework.

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Research Focus






neural circuit

Clinical conditions


Attention deficit hyperactivity disorder

Huntington's disease

Learning disbilities

Movement disorders

Obsessive compulsive disorder

Parkinson's disease



Behavioural analysis

Electrophysiological recording techniques

Magnetic resonance imaging (fMRI)



Anthony Dickinson

Paul Fletcher

Christopher Harris

United Kingdom

Ray Dolan Web:


Peter Bossaerts Web:

Masamichi Sakagami Web:

Masataka Watanabe Web:

Associated News Items

Key publications

O'Neill M, Schultz W (2010), “Coding of reward risk distinct from reward value by orbitofrontal neurons” Neuron 68: 789-800

Tobler PN, Fiorillo CD, Schultz W (2005), “Adaptive coding of reward value by dopamine neurons.” Science 307(5715):1642-5 Details

Fiorillo CD, Tobler PN, Schultz W (2003), “Discrete coding of reward probability and uncertainty by dopamine neurons” Science 299:1898-1902

Waelti P, Dickinson A, Schultz W (2001), “Dopamine responses comply with basic assumptions of formal learning theory.” Nature 412(6842):43-8 Details

Tremblay L, Schultz W (1999), “Relative reward preference in primate orbitofrontal cortex.” Nature 398(6729):704-8 Details

Schultz W, Dayan P, Montague PR (1997), “A neural substrate of prediction and reward.” Science 275(5306):1593-9 Details



Ferrari-Toniolo S, Bujold P, Schultz W (2020), “Compliance with the continuity axiom of Expected Utility Theory supports utility maximization in monkeys” BioRxiv


Al-Mohammad A, Schultz W (2019), “Let a monkey do your bidding: Reward valuation in an auction-like mechanism” BioRxiv

Ferrari-Toniolo S, Bujold P, Schultz W (2019), “Probability distortion depends on choice sequence in rhesus monkeys” J Neurosci 39: 2915-2929

Grabenhorst F, Báez-Mendoza R, Genest W, Deco G, Schultz W (2019), “Primate amygdala neurons simulate decision processes of social partners” Cell

Grabenhorst* F, Tsutsui* KI, Kobayashi S, Schultz W (2019), “Primate prefrontal neurons signal economic risk derived from the statistics of recent reward experience” eLife 8: e44838

Pastor-Bernier A, Stasiak A, Schultz W (2019), “Orbitofrontal signals for two-component choice options comply with indifference curves of Revealed Preference Theory” Nat Comm 10: 4885

Pastor-Bernier A, Volkmann K, Stasiak A, Grabenhorst F, Schultz W (2019), “Single-dimensional human revealed stochastic preferences for multi-dimensional economic choice options” BioRxiv

Schultz W (2019), “Recent advances in understanding the role of phasic dopamine activity” F1000


Brzosko Z, Zannone S, Schultz W, Clopath C, Paulsen O (2017), “Sequential neuromodulation of Hebbian plasticity offers mechanism for effective reward-based navigation” eLife 6:e27756

Diederen KMJ, Ziauddeen H, Vestergaard M, Spencer T, Schultz W, Fletcher P (2017), “Dopamine modulates adaptive prediction error coding in the human midbrain and striatum” J Neurosci 37: 1708-1720

Pastor-Bernier A, Plott CR, Schultz W (2017), “Monkeys choose as if maximizing utility compatible with basic principles of revealed preference theory” PNAS E114: 1766-E1775

Schultz W, Stauffer WR, Lak A (2017), “The phasic dopamine signal maturing: from reward via behavioural activation to formal economic utility” Curr Op Neurobiol 43: 139-148


Báez-Mendoza R, Schultz W (2016), “Performance error-related activity in monkey striatum during social interactions” Sci Reports 6: 37199

Báez-Mendoza R, van Coeverden C, Schultz W (2016), “A neuronal reward inequity signal in primate striatum” J Neurophysiol 115: 68-79

Diederen K, Spencer T, Vestergaard MD, Fletcher P, Schultz W (2016), “Adaptive prediction error coding in the human midbrain and striatum facilitates behavioral adaptation and learning efficiency” Neuron 90: 1127-1138

Genest W, Stauffer WR, Schultz W (2016), “Utility functions predict variance and skewness risk preferences in monkeys” Proc Natl Acad Sci (USA) 113: 8402-8407

Grabenhorst* F, Hernadi* I, Schultz W (2016), “Primate amygdala neurons evaluate the progress of self-defined economic choice sequences” eLife 5: e18731

Lak A, Stauffer WR, Schultz W (2016), “Dopamine neurons learn relative chosen value from probabilistic rewards” eLife 5: e18044

Schultz W (2016), “Dopamine reward prediction error signalling: a two-component response” Nat Rev Neurosci 17: 183-195

Stauffer WR, Lak A, Yang A, Borel M, Paulsen O, Boyden E, Schultz W (2016), “Dopamine neuron-specific optogenetic stimulation in Rhesus macaques” Cell 166: 1564-1571

Tsutsui* KI, Grabenhorst* F, Kobayashi S, Schultz W (2016), “A dynamic code for economic object valuation in prefrontal cortex neurons” Nat Comm 7: 12554

Zangemeister* L, Grabenhorst* F, Schultz W (2016), “Neural basis for economic saving strategies in human amygdala-prefrontal reward circuits” Curr Biol 26: 3004-3013


Brzosko Z, Schultz W, Paulsen O (2015), “Retroactive modulation of spike timing-dependent plasticity by dopamine” eLife 4: e09685

Diederen KMJ, Schultz W (2015), “Scaling prediction errors to reward variability benefits error-driven learning in humans” J Neurophysiol 114: 1628-1640

Hernadi* I, Grabenhorst* F, Schultz W (2015), “Planning activity for internally generated reward goals in monkey amygdala neurons” Nat Neurosci 18

Schultz W (2015), “Neuronal reward and decision signals: from theories to data” Physiol Rev 95: 853-951

Schultz W, Carelli RM, Wightman RM (2015), “Phasic dopamine signals: from subjective reward value to formal economic utility” Curr Op Behav Sci 5: 147-154

Stauffer WR, Lak A, Bossaerts P, Schultz W (2015), “Economic choices reveal probability distortion in macaque monkeys” J Neurosci 35: 3146-3154

Vestergaard MD, Schultz W (2015), “Choice mechanisms for past, temporally extended outcomes” Proc Roy Soc B 282: 20141766, 1810


Bermudez MA, Schultz W (2014), “Timing in reward and decision processes.” Philos Trans R Soc Lond B Biol Sci 369(1637):20120468 Details

Medic N, Ziauddeen H, Vestergaard MD, Henning E, Schultz W, Farooqi IS, Fletcher PC (2014), “Dopamine modulates the neural representation of subjective value of food in hungry subjects” J Neurosci 34: 16877-16889

O'Neill M, Schultz W (2014), “Economic risk coding by single neurons in the orbitofrontal cortex.” J Physiol Paris Details

Stauffer WR, Lak A, Schultz W (2014), “Dopamine reward prediction error responses reflect marginal utility” Curr Biol 2491-2500


Báez-Mendoza R, Schultz W (2013), “The role of the striatum in social behavior.” Front Neurosci 7:233 Details


Ariansen JL, Heien ML, Hermans A, Phillips PE, Hernadi I, Bermudez MA, Schultz W, Wightman RM (2012), “Monitoring extracellular pH, oxygen, and dopamine during reward delivery in the striatum of primates.” Front Behav Neurosci 6:36 Details

Miyapuram KP, Tobler PN, Gregorios-Pippas L, Schultz W (2012), “BOLD responses in reward regions to hypothetical and imaginary monetary rewards.” Neuroimage 59(2):1692-9 Details

Schultz W (2012), “Risky dopamine.” Biol Psychiatry 71(3):180-1 Details


Schultz W (2011), “Potential vulnerabilities of neuronal reward, risk, and decision mechanisms to addictive drugs.” Neuron 69(4):603-17 Details

Schultz W (2011), “Potential vulnerabilities of neuronal reward, risk, and decision mechanisms to addictive drugs” Neuron 69: 603-617

Schultz W, O'Neill M, Tobler PN, Kobayashi S (2011), “Neuronal signals for reward risk in frontal cortex.” Ann N Y Acad Sci 1239:109-17 Details


Burke CJ, Tobler PN, Schultz W, Baddeley M (2010), “Striatal BOLD Response Reflects the Impact of Herd Information on Financial Decisions.” Front Hum Neurosci 4:48 Details

Kobayashi S, Pinto de Carvalho O, Schultz W (2010), “Adaptation of reward sensitivity in orbitofrontal neurons” J Neurosci 30: 534-544

Kobayashi S, Pinto de Carvalho O, Schultz W (2010), “Adaptation of reward sensitivity in orbitofrontal neurons.” J Neurosci 30(2):534-44 Details

Kobayashi S, Schultz W, Sakagami M (2010), “Operant conditioning of primate prefrontal neurons.” J Neurophysiol 103(4):1843-55 Details

Nomoto K, Schultz W, Watanabe T, Sakagami M (2010), “Temporally extended dopamine responses to perceptually demanding reward-predictive stimuli.” J Neurosci 30(32):10692-702 Details

O'Neill M, Schultz W (2010), “Coding of reward risk by orbitofrontal neurons is mostly distinct from coding of reward value.” Neuron 68(4):789-800 Details

Schultz W (2010), “Subjective neuronal coding of reward: temporal value discounting and risk.” Eur J Neurosci 31(12):2124-35 Details

Schultz W (2010), “Dopamine signals for reward value and risk: basic and recent data.” Behav Brain Funct 6:24 Details


Christopoulos GI, Tobler PN, Bossaerts P, Dolan RJ, Schultz W (2009), “Neural correlates of value, risk, and risk aversion contributing to decision making under risk.” J Neurosci 29(40):12574-83 Details


Fiorillo CD, Newsome WT, Schultz W (2008), “The temporal precision of reward prediction in dopamine neurons.” Nat Neurosci Details

Hare TA, O'Doherty J, Camerer CF, Schultz W, Rangel A (2008), “Dissociating the role of the orbitofrontal cortex and the striatum in the computation of goal values and prediction errors.” J Neurosci 28(22):5623-30 Details

Kobayashi S, Schultz W (2008), “Influence of reward delays on responses of dopamine neurons” J Neurosci 28: 7837-7846

Kobayashi S, Schultz W (2008), “Influence of reward delays on responses of dopamine neurons.” J Neurosci 28(31):7837-46 Details


Tobler PN, Fletcher PC, Bullmore ET, Schultz W (2007), “Learning-related human brain activations reflecting individual finances.” Neuron 54(1):167-75 Details


Blatter K, Schultz W (2006), “Rewarding properties of visual stimuli.” Exp Brain Res 168(4):541-6 Details

Kobayashi S, Nomoto K, Watanabe M, Hikosaka O, Schultz W, Sakagami M (2006), “Influences of rewarding and aversive outcomes on activity in macaque lateral prefrontal cortex.” Neuron 51(6):861-70 Details

Schultz W (2006), “Behavioral theories and the neurophysiology of reward” Ann Rev Psychol 57:87-115

Schultz W (2006), “Behavioral theories and the neurophysiology of reward.” Annu Rev Psychol 57:87-115 Details


Cromwell HC, Hassani OK, Schultz W (2005), “Relative reward processing in primate striatum.” Exp Brain Res 162(4):520-5 Details

Fiorillo CD, Tobler PN, Schultz W (2005), “Evidence that the delay-period activity of dopamine neurons corresponds to reward uncertainty rather than backpropagating TD errors.” Behav Brain Funct 1(1):7 Details


Schultz W (2004), “Neural coding of basic reward terms of animal learning theory, game theory, microeconomics and behavioural ecology.” Curr Opin Neurobiol 14(2):139-47 Details


Fiorillo CD, Tobler PN, Schultz W (2003), “Discrete coding of reward probability and uncertainty by dopamine neurons.” Science 299(5614):1898-902 Details

Martin-Soelch C, Missimer J, Leenders KL, Schultz W (2003), “Neural activity related to the processing of increasing monetary reward in smokers and nonsmokers.” Eur J Neurosci 18(3):680-8 Details

Schultz W, Tremblay L, Hollerman JR (2003), “Changes in behavior-related neuronal activity in the striatum during learning.” Trends Neurosci 26(6):321-8 Details


Martin-Soelch C, Chevalley AF, Künig G, Missimer J, Magyar S, Mino A, Schultz W, Leenders KL (2001), “Changes in reward-induced brain activation in opiate addicts.” Eur J Neurosci 14(8):1360-8 Details

Martin-Soelch C, Leenders KL, Chevalley AF, Missimer J, Künig G, Magyar S, Mino A, Schultz W (2001), “Reward mechanisms in the brain and their role in dependence: evidence from neurophysiological and neuroimaging studies.” Brain Res Brain Res Rev 36(2-3):139-49 Details

Martin-Sölch C, Magyar S, Künig G, Missimer J, Schultz W, Leenders KL (2001), “Changes in brain activation associated with reward processing in smokers and nonsmokers. A positron emission tomography study.” Exp Brain Res 139(3):278-86 Details

Schultz W (2001), “Reward signaling by dopamine neurons.” Neuroscientist 7(4):293-302 Details


Cavada C, Schultz W (2000), “The mysterious orbitofrontal cortex. foreword” Cereb Cortex 10(3):205 Details

Hollerman JR, Tremblay L, Schultz W (2000), “Involvement of basal ganglia and orbitofrontal cortex in goal-directed behavior.” Prog Brain Res 126:193-215 Details

Künig G, Leenders KL, Martin-Sölch C, Missimer J, Magyar S, Schultz W (2000), “Reduced reward processing in the brains of Parkinsonian patients.” Neuroreport 11(17):3681-7 Details


Contreras-Vidal JL, Schultz W (1999), “A predictive reinforcement model of dopamine neurons for learning approach behavior.” J Comput Neurosci 6(3):191-214 Details

Schultz W (1999), “The Reward Signal of Midbrain Dopamine Neurons.” News Physiol Sci 14:249-255 Details


Schultz W (1998), “The phasic reward signal of primate dopamine neurons.” Adv Pharmacol 42:686-90 Details

Schultz W, Tremblay L, Hollerman JR (1998), “Reward prediction in primate basal ganglia and frontal cortex.” Neuropharmacology 37(4-5):421-9 Details

Suri RE, Schultz W (1998), “Learning of sequential movements by neural network model with dopamine-like reinforcement signal.” Exp Brain Res 121(3):350-4 Details


Schultz W (1997), “Dopamine neurons and their role in reward mechanisms.” Curr Opin Neurobiol 7(2):191-7 Details


Schultz W (1994), “Behavior-related activity of primate dopamine neurons.” Rev Neurol (Paris) 150(8-9):634-9 Details


Galbraith GC, Schultz W (1993), “Perceptual adaptation in the vestibulo-ocular system: EEG correlates of spatial and temporal rearrangement.” Percept Mot Skills 77(2):595-608 Details

Schultz W, Apicella P, Ljungberg T, Romo R, Scarnati E (1993), “Reward-related activity in the monkey striatum and substantia nigra.” Prog Brain Res 99:227-35 Details


Apicella P, Scarnati E, Schultz W (1991), “Tonically discharging neurons of monkey striatum respond to preparatory and rewarding stimuli.” Exp Brain Res 84(3):672-5 Details

Ljungberg T, Apicella P, Schultz W (1991), “Responses of monkey midbrain dopamine neurons during delayed alternation performance.” Brain Res 567(2):337-41 Details


Romo R, Schultz W (1989), “Somatosensory input to dopamine neurones of the monkey midbrain: responses to pain pinch under anaesthesia and to active touch in behavioural context.” Prog Brain Res 80:473-8; discussion 465-6 Details

Schultz W, Romo R, Scarnati E, Sundström E, Jonsson G, Studer A (1989), “Saccadic reaction times, eye-arm coordination and spontaneous eye movements in normal and MPTP-treated monkeys.” Exp Brain Res 78(2):253-67 Details


Schultz W (1988), “MPTP-induced parkinsonism in monkeys: mechanism of action, selectivity and pathophysiology.” Gen Pharmacol 19(2):153-61 Details

Studer A, Sundström E, Jonsson G, Schultz W (1988), “Acute electrophysiological and neurochemical effects of administration of MPTP in mice.” Neuropharmacology 27(9):923-31 Details


Schultz W, Romo R (1987), “Responses of nigrostriatal dopamine neurons to high-intensity somatosensory stimulation in the anesthetized monkey.” J Neurophysiol 57(1):201-17 Details

Studer A, Schultz W (1987), “The catecholamine uptake inhibitor nomifensine depresses impulse activity of dopamine neurons in mouse substantia nigra.” Neurosci Lett 80(2):207-12 Details


Schultz W (1986), “Responses of midbrain dopamine neurons to behavioral trigger stimuli in the monkey.” J Neurophysiol 56(5):1439-61 Details

Schultz W (1986), “Activity of pars reticulata neurons of monkey substantia nigra in relation to motor, sensory, and complex events.” J Neurophysiol 55(4):660-77 Details

Schultz W, Scarnati E, Sundström E, Tsutsumi T, Jonsson G (1986), “The catecholamine uptake blocker nomifensine protects against MPTP-induced parkinsonism in monkeys.” Exp Brain Res 63(1):216-20 Details

Schultz W, Scarnati E, Sundström E, Tsutsumi T, Jonsson G (1986), ““The catecholamine uptake blocker nomifensine protects against MPTP-induced parkinsonism in monkeys.” ” Exp Brain Res 63(1):216-20


Romo R, Schultz W (1985), “Prolonged changes in dopaminergic terminal excitability and short changes in dopaminergic neuron discharge rate after short peripheral stimulation in monkey.” Neurosci Lett 62(3):335-40 Details

Schultz W, Studer A, Jonsson G, Sundström E, Mefford I (1985), “Deficits in behavioral initiation and execution processes in monkeys with 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine-induced parkinsonism.” Neurosci Lett 59(2):225-32 Details

Toan DL, Schultz W (1985), “Responses of rat pallidum cells to cortex stimulation and effects of altered dopaminergic activity.” Neuroscience 15(3):683-94 Details


Aebischer P, Schultz W (1984), “The activity of pars compacta neurons of the monkey substantia nigra is depressed by apomorphine.” Neurosci Lett 50(1-3):25-9 Details

Schultz W (1984), “Recent physiological and pathophysiological aspects of Parkinsonian movement disorders.” Life Sci 34(23):2213-23 Details


Schultz W (1982), ““Depletion of dopamine in the striatum as an experimental model of Parkinsonism: direct effects and adaptive mechanisms.” ” Prog Neurobiol 18(2-3):121-66


Ruffieux A, Schultz W (1981), “[Influence of dopamine on pars reticulata neurones of substantia nigra (author's transl)].” J Physiol (Paris) 77(1):63-9 Details

Schultz W, Wiesendanger R, Hess B, Ruffieux A, Wiesendanger M (1981), “The somatotopy of the gracile nucleus in cats with agenesis of a hindfoot.” Exp Brain Res 43(3-4):413-8 Details


Ruffieux A, Schultz W (1980), ““Dopaminergic activation of reticulata neurones in the substantia nigra.” ” Nature 285(5762):240-1


Schultz W, Montgomery EB Jr, Marini R (1979), “"Proximal limb movements in response to microstimulation of primate dentate and interpositus nuclei mediated by brain-stem structures.” ” Brain 102(1):127-46


Schultz W, Ungerstedt U (1978), “Striatal cell supersensitivity to apomorphine in dopamine-lesioned rats correlated to behaviour.” Neuropharmacology 17(6):349-53 Details

Schultz W, Ungerstedt U (1978), “A method to detect and record from striatal cells of low spontaneous activity by stimulating the corticostriatal pathway.” Brain Res 142(2):357-62 Details

Ungerstedt U, Ljungberg T, Schultz W (1978), “Dopamine receptor mechanisms: behavioral and electrophysiological studies.” Adv Biochem Psychopharmacol 19:311-21 Details

Zangger P, Schultz W (1978), “The activity of cells of nucleus reticularis tegmenti pontis during spontaneous locomotion in the decorticate cat.” Neurosci Lett 7(2-3):95-9 Details


Allen GI, Gilbert PF, Marini R, Schultz W, Yin TC (1977), “Integration of cerebral and peripheral inputs by interpositus neurons in monkey.” Exp Brain Res 27(1):81-99 Details

Hellweg FC, Schultz W, Creutzfeldt OD (1977), ““Extracellular and intracellular recordings from cat's cortical whisker projection area: thalamocortical response transformation.”” J Neurophysiol 40(3):463-79


Schultz W, Galbraith GC, Gottschaldt KM, Creutzfeldt OD (1976), “A comparison of primary afferent and cortical neurone activity coding sinus hair movements in the cat.” Exp Brain Res 24(4):365-81 Details

Schultz W, Montgomery EB Jr, Marini R (1976), “Stereotyped flexion of forelimb and hindlimb to microstimulation of dentate nucleus in cebus monkeys.” Brain Res 107(1):151-1 Details