Principal Investigator/Program Director (Last, First, Middle): PI Name Roshan Cools

BIOGRAPHICAL SKETCH Provide the following information for the key personnel and other significant contributors in the order listed on Form Page 2. Follow this format for each person. DO NOT EXCEED FOUR PAGES.

NAME POSITION TITLE Cools, Roshan Professor of Cognitive Neuropsychiatry eRA COMMONS USER NAME Cools EDUCATION/TRAINING (Begin with baccalaureate or other initial professional education, such as nursing, and include postdoctoral training.) DEGREE INSTITUTION AND LOCATION YEAR(s) FIELD OF STUDY (if applicable) University of Groningen, The M.Sc.(equiv) 1993 Experimental Psychology University of Cambridge, UK M.Phil. 1999 Experimental Psychology University of Cambridge, UK Ph.D. 2003 Experimental Psychology

A. Personal Statement I have a broad background in psychology and cognitive neuroscience, with specific training and expertise in the neurochemistry of cognitive control and decision making, combining psychopharmacology, pharmacological fMRI and PET to advance our understanding of the role of the major ascending neuromodulators, such as and serotonin, in the cognitive and motivational control of behaviour. As PI or co-Investigator on several university- and NWO-funded grants, I laid the groundwork for the proposed research by establishing a key role for striatal dopamine in learning and cognitive control.

B. Positions and Honors. Appointments 2002-2007 Research Fellow, University of Cambridge UK 2003-2005 Visiting Research Fellow, University of California, Berkeley 2007-present Principal Investigator, Donders Institute for Brain, Cognition and Behaviour, Centre for Cognitive Neuroimaging, Radboud University 2011-present Professor of Cognitive Neuropsychiatry, Dept of Psychiatry, Radboud university medical centre 2014-present Member, (Government) Advisory Council for Science and Technology Policy, NL

Professional Memberships 2017 Member, VIDI award selection committee (Social Sciences), NWO, NL 2016 – present Member, Academia Europaea (Section Behavioural Sciences) 2012 – present Member, Board of the Rathenau Institute (research, debate on Science and Technology), NL 2010 & 2014 Member, Rubicon grant selection Committee (Social Sciences), NWO, NL 2011 – present Member, Advisory Council Attention and Performance, US 2012 – 2014 Contributor to Research Network “A Roadmap on Mental Health Research in Europe” 2011, 2014, 2017 Member, PI Selection Committee, Radboudumc, NL 2008 - 2013 Member, Steering Committee of 1 of 4 Themes (‘Perception, Action and Control’) at Donders Institute, Radboud University, NL 2008 – present Member, Education Committee, Radboud University, NL 2008 – 2012 Member, Research Infrastructure Committee, Radboud University, NL 2013 – present Fellow, Association for Psychological Science, UK 2003 – present Member, Society for Neuroscience, US 2003 – present Member, Cognitive Neuroscience Society, US

Commissions of Trust 2017 – now Reviewing Editor, Journal of Neuroscience 2010 - now Active Editor, Journal of Cognitive Neuroscience 2014 - now Member, Editorial Board Royal Society Open Science PHS 398/2590 (Rev. 05/01) Page Biographical Sketch Format Page Principal Investigator/Program Director (Last, First, Middle): PI Name

2014 - now Member, Editorial Board Current Opinion in Behavioral Sciences 2014 - now Member, Editorial Board Motivation Science 2012 – 2014 Associate Editor, special issue Frontiers Behavioral Neuroscience 2009 – 2015 Consulting Editor, Cognitive, Affective, & Behavioral Neuroscience

Awards and Honors 1998 MSc degree with Cum Laude 1998-1999 Dutch-British Partnership Award, Cambridge UK 1998-2001 Cambridge European Trust Award, UK 1999-2002 CD Marsden Studentship, Parkinson’s Disease Society of the UK 1999-2002 Mary Ann Eward Studentship, Newnham College, Cambridge UK 2002-2006 Junior Research Fellowship, St John’s College, Cambridge UK 2002-2006 Royal Society Dorothy Hodgkin Research Fellowship, University of Cambridge UK 2002 Human Frontiers Longterm Fellowship (declined) 2006-2007 Royal Society University Research Fellowship, University of Cambridge UK 2008 Vidi award from the Netherlands Organization for Scientific Research 2011 Radboud Science Award for best research at Radboud University 2012 Young Investigator Award of the Cognitive Neuroscience Society 2012 James McDonnell Scholar Award 2015 Vici award from the Netherlands Organization for Scientific Research 2017 KNAW-Ammodo Award

C. Contribution to Science I study the brain mechanisms of motivation and cognition by combining cognitive neuroscience with psychopharmacology, focusing on dopamine and serotonin. A strength of our group is the interventional nature of our work, which often involves manipulating the brain through pharmacological, electrical or psychological means in health and disease. The work is motivated by mechanistic questions about motivation and cognition. How do we motivate ourselves to exert cognitive control? How do we maximize reward? We leverage knowledge about dopamine and serotonin and combine the use of neurochemical PET, psychopharmacological fMRI, (computational) model-based analyses and trans-diagnostic patient work to resolve these questions. We hope to contribute to resolving a major problem in neurology and psychiatry: the huge variability in treatment efficacy.

Cognitive effects of Parkinson’s disease In my earliest work I studied effects of dopaminergic medication on learning and control in patients with Parkinson’s disease. These series of experiments provided empirical evidence for, among other things, contrasting effects of medication depending on task demands, which led us to advance the ‘dopamine overdose hypothesis’ (Cools et al., 2001 Cereb Cortex; Cools, 2006 NBR). This and subsequent evidence (Smittenaar et al., 2014; Cools et al., 2010) inspired novel theoretical work on dopamine’s role in behavior (Frank, 2005 J Cogn Neurosci; Frank et al., 2004 Science; Collin and Frank, 2014 Psych Rev) as well as novel recent work on the role of inhibitory control mechanisms in interrupting cognition (Wessel and Aron, 2017 Neuron). I continue to study Parkinson’s disease, in order to refine and revise the dopamine overdose hypothesis, clarifying why not every patients develops the severe psychiatric side effects, such as ICD, after starting medication.

Selected publications: 1. Cools R, Barker RA, Sahakian BJ, Robbins TW (2001). Enhanced or impaired cognitive function in Parkinson's disease as a function of dopaminergic medication and task demands. Cereb Cortex 11:1136-1143. 2. Cools R, Barker RA, Sahakian BJ, and Robbins TW (2001). Mechanisms of cognitive set flexibility in Parkinson’s disease. Brain 124:2503-2512. 3. Cools R, Stefanova E, Barker RA, Robbins TW, Owen AM (2002) Dopaminergic modulation of high-level cognition in Parkinson’s disease: The role of prefrontal cortex revealed by PET. Brain 125: 584-594. 4. Cools R, Barker R A, Sahakian B J, Robbins T W (2003). L-Dopa medication remediates cognitive inflexibility, but increases impulsivity in patients with Parkinson’s disease. Neuropsychologia 41:1431- 1441. PHS 398/2590 (Rev. 09/04, Reissued 4/2006) Page 6 Continuation Format Page Principal Investigator/Program Director (Last, First, Middle): PI Name

5. Cools R, Robbins TW (2004). Chemistry of the Adaptive Mind. Philos Transact Ser A Math Phys Eng Sci 362 (1825): 2871-2888. 6. Cools R (2006) Dopaminergic modulation of cognitive function – Implication for L-DOPA therapy in Parkinson’s disease. Neurosci Biobehav Rev 30 (1):1-34. 7. Cools R, Altamirano L, D’Esposito M (2006). Reversal learning in Parkinson’s disease depends on medication status and outcome valence. Neuropsychologia 44 (10):1663-1673 8. Cools R, Lewis SGJ, Clark L, Barker RA, Robbins TW (2007). L-DOPA disrupts activity in the nucleus accumbens during reversal learning in Parkinson’s disease. Neuropsychopharmacology 32 (1): 180- 189. 9. Cools R, Miyakawa A, Sheridan M, D'Esposito M (2010). Enhanced frontal function in Parkinson's disease. Brain 133:225-33. 10. De Wit S, Barker RA, Dickinson A, Cools R (2011). Habitual versus goal-directed action control in Parkinson disease. J Cogn Neurosci. 23(5):1218-1229 [5.7] 11. Smittenaar P, Chase HW, Aarts E, Nusselein B, Bloem BR, Cools R (2012). Decomposing effects of dopaminergic medication in Parkinson’s disease on probabilistic action selection: learning or performance? Eur J Neurosci 35(7):1144-51 [3.9] 12. Aarts E, Nusselein AA, Smittenaar P, Helmich RC, Bloem BR, Cools R (2014). Greater striatal s disease are associated with better task-switching but worse reward׳responses to medication in Parkinson performance. Neuropsychologia 62:390-7 13. Robbins TW, Cools R (2014). Cognitive Deficits in Parkinson’s Disease: A Cognitive Neuroscience Perspective. Movement Dis 29:597:607 14. Timmer MHM, Sescousse G, Van der Schaaf ME, Esselink RAJ, Cools R (in press). Reward learning deficits in Parkinson’s disease depend on depression. Psychol Med

Reversal learning In a second line of work, I have introduced a paradigm for assessing the neural mechanisms of probabilistic reversal learning in healthy volunteers. Many other research groups have subsequently used this paradigm to study reversal learning.

Selected publications on reversal learning: 1. Cools R, Clark L, Owen AM, Robbins TW (2002). Defining the neural mechanisms of probabilistic reversal learning using event-related functional MRI. J Neurosci 22: 4563-4567. 2. Chase HW, Swainson R, Durham L, Benham L, Cools R (2011). Feedback-related negativity codes prediction error, but not behavioural adjustment during probabilistic reversal learning. J Cogn Neurosci. 23(4):936-946 3. Van der Schaaf ME, Van Schouwenburg MR, Geurts D, Schellekens AFA, Buitelaar J, Verkes RJ, Cools R (2014). Establishing the dopamine-dependency of human striatal signals during reward and punishment reversal learning. Cereb Cortex 24(3):633-42

Role of serotonin in learning and control Inspired by the observation that detrimental overdose effects of dopaminergic medication in PD resemble effets of serotonin depletion, I developed a parallel line of work on the effects of central serotonin manipulation, using the dietary tryptophan depletion procedure, on learning and control. This work has demonstrated a selective role for serotonin in punishment learning and led to recent work providing empirical foundations for current theoretical ideas about setononin’s role in coupling aversive motivation to behavioural inhibition (Boureau et al., 2011 Neuropsychopharmacology; Cools et al., 2001 Neuropsychopharmacology).

Selected publications: 1. Cools R, Roberts AC, Robbins TW (2008). Serotoninergic regulation of emotional and behavioural control processes. Trends Cogn Sci 12(1):31-40. 2. Cools R, Blackwell A, Clark L, Menzies L, Cox S, Robbins TW (2005). Tryptophan depletion disrupts the motivational guidance of goal-directed behavior as a function of trait impulsivity. Neuropsychopharmacology 30 (7): 1362-1373.

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3. Cools R, Robinson OJ, Sahakian BJ (2008). Acute tryptophan depletion in healthy volunteers enhances punishment prediction but does not affect reward prediction. Neuropsychopharmacology 33(9):2291-9. 4. Cools R, Nakamura K, Daw ND (2011): Serotonin and Dopamine: Unifying Affective, Activational, and Decision Functions. Neuropsychopharmacology Reviews 36(1):98-113 5. Den Ouden H, Fernandez G, Elshout J, Rijpkema M, Hoogman M, Franke B, Daw ND, Cools R (2013). Dissociable effects of dopamine and serotonin on reversal learning. Neuron 80(4):1090-100 [16.5] 6. Geurts D, Huys Q, den Ouden H, Cools R (2013). Serotonin and aversive Pavlovian control of instrumental behavior in humans. J Neurosci 33(48):18932-9 [7.6] 7. den Ouden HEM, Swart JC, Schmidt K, Fekkes D, Geurts DEM, Cools R (2015). Acute serotonin depletion releases motivated inhibition of response vigour. Psychopharm 232(7):1303-12 [4.1] 8. Crockett M and Cools R (2015). Serotonin and aversive processing in affective and social decision- making. Current Opinion in Behavioral Sciences 5:64-70

Paradoxical effects of dopamine on healthy human cognition In a fourth line of work I aim to identify the mechanistic factors that contribute to the large variation in dopaminergic drug effects on cognitive control in the healthy population. This work provided key empirical foundations for the proposal that there is an Inverted-U shaped relationship between dopamine and human cognition. Moreover pharmacological fMRI work with healthy volunteers demonstrated that dopaminergic drugs have opposite effects on the stabilizing and flexing aspects of cognitive control.

Selected publications: 1. Cools R, Sheridan M, Jacobs EJ, D’Esposito MD (2007). Impulsive personality predicts dopamine- dependent changes in fronto-striatal activity during component processes of working memory. J Neurosci 27(20): 5506-5514. 2. Cools R, Gibbs SE, Miyakawa A, Jagust W, D'Esposito M (2008).Working memory capacity predicts dopamine synthesis capacity in the human striatum. J Neurosci 28(5):1208-12. 3. Cools R, Frank MF, Gibbs SE, Miyakawa A, Jagust W, D'Esposito M (2009). Striatal dopamine predicts outcome-specific reversal learning and its sensitivity to dopaminergic drug administration. J Neurosci 29: 1538-1543. 4. Cools R and D’Esposito M (2011). Inverted-U-Shaped dopamine actions on human working memory and cognitive control. Biol Psychiatry 69(12):e113-25. 5. Van Holstein M, Aarts E, van der Schaaf ME, Geurts DE, Verkes RJ, Franke B, van Schouwenburg MR, Cools R (2011). Human cognitive flexibility depends on dopamine D2 receptor signaling. Psychopharm 218(3):567-78. [4.1] 6. Van der Schaaf ME, Fallon SJ, ter Huurne N, Buitelaar J, Cools R (2013). Working memory capacity predicts effects of methylphenidate on reversal learning. Neuropsychopharmacology 38(10):2011- 8. 7. Van Schouwenburg MR, Zwiers MP, van der Schaaf ME, Geurts DE, Schellekens AF, Buitelaar JK, Verkes RJ, Cools R (2013). Anatomical connection strength predicts dopaminergic drug effects on fronto-striatal function. Psychopharm 227(3):521-31 8. Fallon SJ, Van der Schaaf ME, ter Huurne N, Cools R (2016). The neurocognitive cost of enhancing cognition with methylphenidate: improved distractor resistance but impaired updating. J Cogn Neurosci 25:1-12.

Cognitive role of the basal ganglia: A fifth line of work has contributed by revising the classic view that dopamine’s effects on cognition reflect modulation of exclusively the prefrontal cortex and has highlighted a key role for the basal ganglia in cognitive control and learning.

Selected publications: 1. Cools R, Clark L, Robbins TW (2004). Differential responses in human striatum and prefrontal cortex to changes in object- and rule-relevance. J Neurosci 24:1129-35. 2. Cools R, Ivry RB, D’Esposito M (2006). The human striatum is necessary for responding to changes in stimulus relevance. J Cogn Neurosci 18: 1973-1983 3. Van Schouwenburg M, Den Ouden H, Cools R (2010). The human basal ganglia modulate fronto-

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posterior connectivity during attention shifting. J Neurosci 30:9910-9918. 4. Cools R (2011). Dopaminergic control of the striatum for high-level cognition. Curr Opin Neurobiol 21:402-407 5. Van Schouwenburg M, O'Shea J, Mars R, Rushworth R, and Cools R (2012). Controlling human striatal cognitive function via the frontal cortex. J Neurosci 32(16):5631-7 6. Van Schouwenburg MR, Onnink AMH; ter Huurne N; Kan CC, Zwiers MP, Hoogman M, Franke B, Buitelaar JK, Cools R (2014). Cognitive flexibility depends on white matter microstructure of the basal ganglia. Neuropsychologia 53:171-7 7. Van Schouwenburg MR, den Ouden H, Cools R (2015). Selective attentional enhancement and inhibition of fronto-posterior connectivity by the basal ganglia during attention switching. Cerebral Cortex 25(6):1527-34 8. Piray P, Toni I, Cools R (2016). Human choice strategy varies with anatomical projections from ventromedial prefrontal cortex to medial striatum. J Neurosci 36(10):2857-67 9. Piray P, den Ouden HEM, Van der Schaaf ME, Toni E, Cools R (in press). Dopaminergic modulation of the functional ventrodorsal architecture of the human striatum. Cerebral Cortex

Cognitive neuropsychiatry of learning, motivation and control I have begun to leverage these various lines of mechanistic work to advance our understanding, in a transdiagnostic RDoCs-like manner, of a greater variety of neuropsychiatric disorders that implicate learning and control deficits.

Selected publications: 1. Robinson OJ, Cools R, Carlisi CO, Sahakian BJ, Drevets WC (2012). Ventral striatum response during reward and punishment reversal learning in unmedicated major depressive disorder. Am J Psychiatry 169(2):152-9 2. Ly V, Cools R, Roelofs K (2014). Aversive disinhibition of behavior and striatal signaling in social avoidance. Social Cogn Aff Neurosci 9(10):1530-6 [6.4] 3. Von Rhein D, Cools R, Zwiers MP, van der Schaaf M, Franke B, Luman M, Oosterlaan J, Heslenfeld DJ, Hoekstra PJ, Hartman CA, Faraone SV, van Rooij D, van Dongen EV, Lojowska M, Mennes M, Buitelaar J. (2015). Increased neural responses to reward in adolescents and young adults with Attention- deficit/Hyperactiviy Disorder and their unaffected siblings. J Am Acad Child Adolesc Psychiatry 54(5):394-402 4. Janssen L, Sescousse G; Hashemi M; Timmer M, ter Huurne N, Geurts D, Cools R (2015). Abnormal modulation of reward versus punishment learning by a dopamine D2-receptor antagonist in pathological gamblers. Psychopharmacology 232(18):3345-53. 5. Sescousse G, Janssen L, Hashemi M, Timmer M, Geurts D, ter Huurne N, Clark L, Cools R (2016). Amplified striatal responses to near-miss outcomes in pathological gamblers. Neuropsychopharmacology 41(10):2614-23 6. Ly V, Von Borries AKL, Brazil IA, Bulten BH, Cools R*, Roelofs K* (2016). Reduced transfer of affective value to instrumental behavior in violent offenders. Journal of Abnormal Psychology 125(5):657-63 7. Geurts D, von Borries K, Volman I, Bulten B, Cools R*, Verkes R*. (2016). Neural connectivity during reward expectation dissociates psychopathic criminals from noncriminal individuals with high impulsive/antisocial psychopathic traits. Social Cognitive and Affective Neuroscience 11(8):1326- 34

Our drive towards cognition Finally, the empirical evidence that we have provided for a key role of striatal dopamine in motivating cognitive control has provided the basis for my most recent work, which I leverage in the ongoing Vici project, within which Dr Westbrook will be embedded.

Selected publications 1. Cools R (2008). Role of dopamine in the motivational and cognitive control of behavior. Neuroscientist 14(4):381-95. 2. Aarts E, Roelofs A, Franke B, Rijpkema M, Fernández G, Helmich RC, Cools R (2010). Striatal dopamine mediates the interface between motivational and cognitive control in humans: Evidence from

PHS 398/2590 (Rev. 09/04, Reissued 4/2006) Page 9 Continuation Format Page Principal Investigator/Program Director (Last, First, Middle): PI Name genetic imaging. Neuropsychopharmacology 35:1943-1951. 3. Aarts E, van Holstein M, Cools R (2011) Striatal dopamine and the interface between motivation and cognition. Frontiers in Cognition 2:article 163 (doi: 10.3389/fpsyg.2011.00163) 4. Aarts E, Helmich RC, Janssen MJ, Oyen WJ, Bloem BR, Cools R (2012). Aberrant reward processing in Parkinson's disease is associated with dopamine cell loss. Neuroimage 59(4):3339-46. [7.0] 5. Aarts E, Wallace DL, Dang LC, Jagust W, Cools R, D'Esposito M (2014). Dopamine and the cognitive downside of a promised bonus. Psych Sci 25(4):1003-9 6. Fallon SJ, Cools R (2014). Reward Acts on the pFC to Enhance Distractor Resistance of Working Memory Representations. J Cogn Neurosci 26(12):2812-26 [5.7] 7. Cools R (2016). The costs and benefits of brain dopamine for cognitive control. Wiley Interdiscip Rev Cogn Sci 7(5):317-329

D. Additional information: Research Support and/or Scholastic Performance

Ongoing Research Support

Ammodo KNAW Award 2017 Cools (PI) 9/1/17-8/31/2022 An award from the Ammodo Foundation and The Royal Netherlands Academy of Arts and Sciences to encourage unfettered fundamental scientific research. The Ammodo KNAW Award is granted once every two years to eight awardees covering all scientific disciplines. The funds will be used to study dopamine’s role in the incentivisation and valuation of mentally effortful control processes. Role: PI

JSMF #2200202328 Cools (PI) 1/1/13-12/31/2018 Understanding motivation and cognition: Combining psychology and neuropharmacology. The goal of the work funded by this James McDonnell Scholar Award is to advance our understanding of dopamine’s role in the contrasting effects of motivation on cognitive control Role: PI

#453-14-015 NWO Vici Innovational Research Incentives Scheme; Cools (PI) 2/1/16-1/31/2021 Netherlands Organization for Scientific Research The cost of cognitive doping: From dopamine to cognitive control via dynamic neural coding The goal of this project is to build a proxy model of dopamine in order to predict the cognitive effects of dopaminergic drugs Role: PI

2014/22011/MaGW Research Talent grant 406-14-028 Social Sciences; PI: Cools 2014-2018 Netherlands Organization for Scientific Research Title: Overcoming affective biases of decision-making: Combining neurophysiology, neurostimulation and computational psychology Role: PI

2013.WP4.PhD.RUN.002 PhD grant; PI: Cools 2014-2018 Language in Interaction Gravitation project Title: Unity and Diversity of the Neural Substrates of Cognitive Control Functions in Speaking The goal of this project is to assess the neural mechanisms underlying interactions between motor and cognitive control, assessing effects of (vocal vs manual) effector systems on cognitive control Role: PI

Internal competition grant Junior Researcher position; PI: Schene 2014-2018 Donders Centre for Neuroscience Title: Comorbidity of Autism Spectrum Disorders (ASD) and Major Depressive Disorder (MDD): a multi-level study of affective processing and executive control. The goal of this project is to assess the neurocognitive mechanisms underlying comorbidity between depression and autism

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Role: Co-PI

Completed Research Support FOCOM Consortium Grant 2012-2015 European Fonds voor Regionale Ontwikkeling) The goal of FOCOM (Food and Cognition Model systems) is to design novel test setups that can help us to understand how food and brain influence each other. Role: Co-PI

International Parkinson’s Disease Foundation Research Grant; PI: Esselink 2012-2015 Title: Neurobiological mechanisms of depression and cognition in Parkinson’s disease The goal of this project is to assess effects of depression in Parkinson’s disease on learning, choice and motivation and associated fronto-striatal mechanisms Role: co-PI

AGIKO 40-00703-098-11621 grant Medical Sciences; PI: Cools 2011-2014 Netherlands Organization for Scientific Research Title: Cognitive neuropsychiatry: Understanding and treating emotional dysregulation using functional magnetic resonance imaging Role: PI

2011/00477/MaGW Open Competition grant Social Sciences; PI: Ivan Toni 2010-2014 Netherlands Organization for Scientific Research Title: Integrating motivation, cognition and action – Frontal control of dopamine-dependent striatal processing Role: Co-investigator

2010/16453/MaGW Open Competition grant Social Sciences; PI: Cools 2010-2014 Netherlands Organization for Scientific Research Title: Pharmacological modulation of alpha and gamma oscillations: implications for working memory maintenance Role: PI

RGP0036/2009-C Human Frontiers Science Program Research grant; PI: Kae Nakamura 2009–2012 Title: Serotonin and decision-making: integrating interspecies experimental and computational approaches. Role: Co-investigator

ALW2PJ/08042 Open Competition grant Life Sciences; PI: Forstmann 2009-2013 Netherlands Organization for Scientific Research Title: The Anatomical and Neurochemical Foundations of Decision-Making under Time Pressure Role: Co-investigator.

016.095.340 VIDI Innovational Research Incentives Scheme; PI: Cools 09/01/2008 – 09/01/2013 Netherlands Organization for Scientific Research Title: Dopaminergic modulation of fronto-striatal activity during cognitive control Purpose: Examine the effects of dopaminergic drugs on set-shifting in healthy volunteers as a function of genetic predisposition and in patients with focal lesions. Role: PI

F2008(1)-01 Dutch Brain foundation fellowship; PI: Cools 04/01/2009-04/01/2011 Title: Compensatie van cognitieve achteruitgang door motivatie in de ziekte van Parkinson - een fMRI studie

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NIH R01 DA020600; PI: D’Esposito 04/10/06 – 04/10/11 National Institute of Neurological Disorders and Stroke Title: Dopaminergic modulation of fronto-striatal function Purpose: Examine the effects of dopaminergic drugs on component processes of working memory and cognitive control in healthy volunteers and patients with Parkinson’s disease. Role: Co-investigator

RG 47229; PI: Cools 03/01/07-10/30/07 Fast track grant, Parkinson’s Disease Society of the UK Title: Dopaminergic modulation of habit formation in Parkinson’s disease Purpose: Examine the beneficial and detrimental effects of dopaminergic medication on different forms of learning in Parkinson’s disease. Role: PI

APDA SPO018682; PI: D’Esposito 09/01/04 – 09/30/05 American Parkinson’s Disease Association Title: Defining the cognitive enhancing and impairing effects of dopaminergic medication in Parkinson’s Disease; Purpose: Examine the beneficial and detrimental effects of dopaminergic medication in Parkinson’s disease on reward- and punishment-based reversal learning Role: Co-Investigator

Royal Society University Research Fellowship; PI: Cools 2006-2011 (cut short in 2007) Topic: Role of the basal ganglia in cognitive flexibility

Royal Society Dorothy Hodgkin Research Fellowship; PI: Cools 2002-2006 Topic: Cognitive and reward functions of fronto-striatal circuitry

Junior Research Title A Fellowship; PI: Cools 2002-2005 St John's College Cambridge

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