Jonathan D. Cohen

Eugene Higgins Professor of Psychology Director, Center for the Study of Brain, Mind and Behavior Director, Program in Neuroscience Princeton University Professor of Psychiatry University of Pittsburgh Green Hall Princeton University Princeton, NJ 08544 (609) 258-2696 (609) 258-2574 (fax) jdc@princeton.edu Curriculum Vitae Representative Publications Related Links

Research in my laboratory focuses on the neurobiological mechanisms underlying cognitive control, and their disturbance in psychiatric disorders such as schizophrenia and depression. Cognitive control is the ability to guide attention, thought and action in accord with goals or intentions. One of the fundamental mysteries of neuroscience is how this capacity for coordinated, purposeful behavior arises from the distributed activity of many billions of neurons in the brain. Several decades of cognitive and neuroscientific research have focused on the mechanisms by which control influences processing (e.g., attentional effects in sensory processing, goal directed sequencing of motor output, etc.), and the brain structures upon which these functions depend, such as the prefrontal cortex, anterior cingulate cortex, basal ganglia and brainstem neuromodulatory systems. However, we still have a poor understanding of how these systems give rise to cognitive control. Our work seeks to develop formally explicit hypotheses about the functioning of these systems, and to test these hypotheses in empirical studies. An important motivation for this work is the development of a theoretically sound foundation for research on the relationship between disturbances of brain function and their manifestation as disorders of thought and behavior in psychiatric illness.

Theoretical work. Neural network models are developed as a way of articulating precise hypotheses about the function of particular brain systems, and their role in cognitive control. This work seeks to bridge between the traditionally disparate levels of analysis of neurophysiology, systems neuroscience, and cognitive psychology. Projects focus on the function of systems considered to be critical for cognitive control, including: a) the role of prefrontal cortex in biasing attention and response selection in posterior structures; b) the role of brainstem dopamine systems in regulating learning and updating of representations in prefrontal cortex; c) the role of the anterior cingulate cortex in monitoring performance, and its influence on adaptations in control; and d) the influence of locus coeruleus and norepinephrine on attentional state. In many cases, modeling work has led to novel predictions about neurophysiolgical mechanisms underlying systems-level function, such as: a) gain control as a mechanism for dopaminergic neuromodulation; b) the role of dopamine in coordinating reinforcement learning and the gating of information into prefrontal cortex; c) the influence of electrotonic coupling on population dynamics within the locus coeruleus; and d) the effects of changes in locus coeruleus physiological state on attentional mode. In other cases, this work has led to novel hypotheses about system level function, such as the response of anterior cinglulate cortex to conflict in processing and its influence on adaptive changes in cognitive control. This work has also predicted, and led to the discovery of new anatomic relationships, such as projections from the anterior cingulate cortex to locus coeruleus.

Empirical work. Experimental studies within the laboratory make use of behavioral testing, neuroimaging (using functional magnetic resonance imaging) and scalp electrical recordings in humans. Collaborations with neurophysiologists also involve direct neuronal recordings in awake behaving animals, and detailed anatomic studies. Experiments are designed to test predictions made by neural network models, and to provide data needed to guide their further development. An important focus of this work is the generation and testing of hypotheses about the neurobiological mechanisms underlying disturbances of behavior in psychiatric disorders. By manipulating variables of biological interest in our models, we are able to explore the effects that disturbances in these variables have on behavior, and then test these in empirical studies. Empirical findings emerging from this work include: a) the first demonstration in humans of sustained activity in prefrontal cortex associated with working memory performance; b) the correlation of prefrontal cortex activity with parametric manipulations of working memory load; c) the dissociation of frontal responses to working memory load from task difficulty; d) the effects of dopamine manipulation on selective attention and working memory tasks; e) selective deficits both in behavioral and prefrontal activity among patients with schizophrenia in these tasks; f) the response of the anterior cingulate to processing conflict in the absence of performance errors; g) the co-localization of event-related potentials associated with errors (ERN) and processing conflict (N2C).

New directions. A focus of increasing interest within the laboratory is the interaction between cognitive and emotional processing. This stems from an appreciation of the close interactions between executive (e.g. prefrontal) and evaluative (e.g., cingulate) functions evident in our work on cognitive control, and an appreciation of the fact that few, if any, aspects of real world behavior are governed exclusively by one form of processing or the other. Recent studies in the laboratory have begun to explore interactions between cognitive and emotional processes in a variety of behavioral domains, including economic choice (e.g., gambling tasks), social interaction (e.g., ultimatum and bargaining games), and moral decision making. Initial findings, using both behavioral and neuroimaging methods, have provided clear evidence for the prevalent engagement of emotional systems in tasks traditionally considered to be predominantly cognitive. This work offers the hope of producing a more accurate portrayal of real world behavior. It is also likely to have direct relevance to our understanding of psychiatric disorders, which invariably involve complex interactions between disturbances of thought and feeling.

REPRESENTATIVE PUBLICATIONS
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Prefrontal Cortex, Dopamine, and Cognitive Control

Computational Modeling and Theoretical Reviews

Cohen JD, Dunbar K, & McClelland JL (1990). On the control of automatic processes: A parallel distributed processing model of the Stroop effect. Psychological Review, 97(3):332-361.

Servan-Schreiber D, Printz H, & Cohen JD (1990). A network model of catecholamine effects: Gain, signal-to-noise ratio, and behavior. Science, 249, 892-895.

Cohen JD & Servan-Schreiber D (1992). Context, cortex and dopamine: A connectionist approach to behavior and biology in schizophrenia. Psychological Review, 99, 45-77.

Cohen JD & O'Reilly RC (1996). A preliminary theory of the interactions between prefrontal cortex and hippocampus that contribute to planning and prospective memory. In Brandimonte M, Einstein GO & McDaniel MA (Eds.), Prospective Memory: Theory and Applications. Hillsdale, NJ: Erlbaum.

O'Reilly RC, Braver TS, & Cohen JD (1999). A biologically-based neural network model of working memory. In Shah P & Miyake A (Eds.), Models of Working Memory. Cambridge University Press.

Braver TS & Cohen JD (2000). On the control of control: The role of dopamine in regulating prefrontal function and working memory. In Monsell S & Driver J (Eds.), Attention and Performance XVIII; Control of cognitive processes, pp.713-737.

Miller EK & Cohen JD (2001). An integrative theory of prefrontal cortex function. Annual Review of Neuroscience, 24:167-202.

Cohen JD, Braver TS & Brown JW (2002). Computational perspectives on dopamine function in prefrontal cortex, Current Opinion in Neurobiology, 12, 223-229.

O'Reilly RC, Noelle DC, Braver TS & Cohen JD (2002). Prefrontal cortex in dynamic categorization tasks: Representational organization and neuromodulatory control. Cerebral Cortex, 12, 246-257.

Cohen JD, Aston-Jones G & Gilzenrat MS (2004).  A systems-level perspective on attention and cognitive control:  Guided activation, adaptive gating, conflict monitoring, and exploitation vs. exploration. In Posner MI (Ed.), Cognitive Neuroscience of Attention. New York: Guilford Press, pp. 71-90.

Montague PR, Hyman SE & Cohen JD (2004). Computational roles for dopamine in behavioural control. Nature 431(7010), 760-767.

Rougier NP, Noelle DC, Braver TS, Cohen JD & O'Reilly (2005).  Prefrontal cortex and flexible cognitive control:  Rules without symbols.  Proceedings of the National Academy of Sciences, USA, 102(20), 7338-7343.

Bogacz R, Brown ET, Moehlis J, Hu P, Holmes P & Cohen JD (2006). The physics of optimal decision making:  A formal analysis of models of performance in two-alternative forced choice tasks.  Psychological Review, 113, 700-765.

Simen P, Cohen JD & Holmes P (2006).  Adaptation of decision making parameters by continuous performance monitoring.  Neural Networks, 19, 1013-1026.

Neuroimaging

Barch DM, Braver TS, Nystrom LE, Forman SD, Noll DC, & Cohen JD (1997). Dissociating working memory from task difficulty in human prefrontal cortex. Neuropsychologia, 35, 1373-1380.

Cohen JD, Perlstein WM, Braver TS, Nystrom LE, Noll DC, Jonides J, & Smith EE (1997). Temporal dynamics of brain activation during a working memory task. Nature, 386, 604-608.

Berns GS, Cohen JD, & Mintun MA (1997). Brain regions responsive to novelty in the absence of awareness. Science, 276, 1272-1275.

Braver TS, Cohen JD, Jonides J, Smith EE, & Noll DC (1997). A parametric study of prefrontal cortex involvement in human working memory. NeuroImage, 5(1), 49-62.

Nystrom LE, Braver TS, Sabb FW, Delgado MR, Noll DC, & Cohen JD (2000). Working memory for letters, shapes and locations: fMRI evidence against stimulus-based regional organization of human prefrontal cortex. NeuroImage, 11, 424-446.

Kerns JG, Cohen JD, MacDonald III AW, Cho RY, Stenger VA, Carter CS (2004). Anterior cingulate conflict monitoring and adjustments in control. Science, 303, 1023-1026.

Yeung N, Nystrom LE, Aronson JA & Cohen JD (2006). Between-task competition and cognitive control in task switching.  Journal of Neuroscience, 26(5), 1429-1438.

Anterior Cingulate Cortex, Conflict Monitoring and Error Detection

Computational Modeling

Botvinick, MM, Braver TS, Carter CS, Barch DM & Cohen JD (2001). Conflict monitoring and cognitive control. Psychological Review, 108(3) 624-652.

Cho RY, Nystrom LE, Brown ET, Jones AD, Braver TS, Holmes PJ, & Cohen JD. (2002). Mechanisms underlying performance dependencies on stimulus history in a two-alternative forced choice task. Journal of Cognitive, Affective and Behavioral Neuroscience, 2(4), 283-299.

Jones, AD, Cho R, Nystrom LE, Cohen JD & Braver TS (2002). A computational model of anterior cingulate function in speeded response tasks: Effects of frequency, sequence and conflict. Journal of Cognitive, Affective and Behavioral Neuroscience, 2(4), 300-317.

Yeung N, Botvinic MM & Cohen JD (2004). The neural basis of error detection: conflict monitoring and the error-related negativity.  Psychological Review, 111(4), 931-959.

Yeung N & Cohen JD (2006).  The impact of cognitive deficits on conflict monitoring: Predictable dissociations between the ERN and N2.  Psychological Science, 17(2), 164-171.

fMRI

Carter CS, Braver TS, Barch DM, Botvinick MM, Noll DC, & Cohen JD (1998). Anterior cingulate cortex, error detection and the on-line monitoring of performance. Science, 280, 747-749.

Botvinick MM, Nystrom L, Fissell K, Carter CS, & Cohen JD (1999). Conflict monitoring vs. selection-for-action in anterior cingulate cortex. Nature, 402, 179-181.

MacDonald AW, Cohen JD, Stenger VA, & Carter CS (2000). Dissociating the role of dorsolateral prefrontal cortex and anterior cingulate cortex in cognitive control. Science, 288, 1835-1837.

van Veen V, Cohen JD, Botvinick MM, Stenger VA and Carter CS (2001). Anterior cingulate cortex, conflict monitoring, and levels of processing. Neuroimage, 14, 1302-1308.

Kerns JG, Cohen JD, MacDonald III AW, Cho RY, Stenger VA, Carter CS (2004). Anterior cingulate conflict monitoring and adjustments in control. Science, 303, 1023-1026.

Electrophysiology (ERP)

Holroyd CB, Nieuwenhuis S, Yeung N & Cohen JD (2003). Errors in reward prediction are reflected in the event-related brain potential.  NeuroReport, 14(18), 2481-2484.

Nieuwenhuis S, Yeung N, Cohen JD (2004). Stimulus modality, perceptual overlap, and the go/no-go N2. Psychophysiology, 41, 157-160.

Nieuwenhuis S, Yeung N, Holroyd CB, Schurger A & Cohen JD (2004).  Sensitivity of electrophysiological activity from medial frontal cortex to utilitarian and performance feedback. Cerebral Cortex, 14, 741-747

Holroyd CB, Larsen JT & Cohen JD (2004). Context dependence of the event-related brain potential associated with reward and punishment. Psychophysiology, 41(2), 245-53.

Yeung N, Holroyd CB & Cohen JD (2005). ERP correlates of feedback and reward processing in the presence and absence of response choice. Cerebral Cortex, 15, 535-544..

Usher M, Cohen JD, Rajkowsky J, & Aston-Jones G (1999). The role of locus coeruleus in the regulation of cognitive performance. Science, 283, 549-554.

Gilzenrat MS, Holmes BD, Holmes PJ, Rajkowski J, Aston-Jones G & Cohen JD (2002). A modified Fitzhugh-Nagumo system simulates locus coeruleus-mediated regulation of cognitive performance. Neural Networks, 15, 647-663.

Clayton EC, Rajkowski J, Cohen JD & Aston-Jones G (2004). Phasic activation of monkey locus coeruleus neurons by simple decisions in a forced choice task.  Journal of Neuroscience, 24(45).

Aston-Jones G & Cohen JD (2005).  An integrative theory of locus coeruleus-norepinephrine function:  Adaptive gain and optimal performance.  Annual Review of Neuroscience, 28, 403-450.

Nieuwenhuis S, Aston-Jones G & Cohen JD (2005). Decision making, the P3, and the locus coeruleus-norepinephrine system.  Psychological Bulletin, 131(4), 510–532.

Nieuwenhuis S, Gilzenrat MS, Holmes BD, Cohen JD (2005). The role of the locus coeruleus in mediating the attentional blink: A neurocomputational theory.  Journal of Experimental Psychology:  General, 134, 291-307.

Greene JD, Sommerville RB, Nystrom LE, Darley JM, & Cohen JD. (2001). An fMRI investigation of emotional engagement in moral judgment. Science, 293, 2105-2108.

Greene JDC, Nystrom LE, Engell AD, Darley JM & Cohen JD (2004). The neural bases of cognitive conflict and control in moral judgment.   Neuron, 44(2), 389-400.

Cohen JD and Blum KI (2002). Overview: Reward and decision. Introduction to special i ssue. Neuron, 36(2), 193-198.

Sanfey AG, Rilling JK, Aronson JA, Nystrom LE & Cohen JD (2003). The neural basis of economic decision making in the ultimatum game. Science, 300, 1755-1757.

McClure SM, Laibson DI, Loewenstein G & Cohen JD (2004). Separate neural systems value immediate and delayed monetary rewards.  Science, 306, 503-507.

Rilling JK, Sanfey AG, Aronson JA, Nystrom LE & Cohen JD (2004). The neural correlates of theory of mind within interpersonal interactions. Neuroimage 22(4), 1694-703.

Rilling JK, Sanfey AG, Aronson JA, Nystrom LE & Cohen JD (2004). Opposing BOLD responses to reciprocated and unreciprocated altruism in putative reward pathways.  NeuroReport, 15(16), 2539-2243.

Wager TD, Rilling JK, Smith EE, Sokolik A, Casey KL, Davidson RJ, Kosslyn SK, Rose RM &  Cohen JD (2004). Placebo-induced changes in fMRI in the anticipation and experience of pain.  Science, 303, 1162-1167.

Cohen JD, Romero RD, Servan-Schreiber D, & Farah MJ (1994). Mechanisms of spatial attention: The relation of macrostructure to microstructure in parietal neglect. Journal of Cognitive Neuroscience, 6(4), 377-387.

Armony JL, Servan-Schreiber D, Cohen JD, & LeDoux JE (1995). An anatomically-constrained neural network model of fear conditioning. Behavioral Neuroscience, 109(2), 246-256.

Cohen JD and Servan-Schreiber D (1992). Context, cortex and dopamine: A connectionist approach to behavior and biology in schizophrenia. Psychological Review, 99, 45-77.

Cohen JD & Servan-Schreiber D (1993). A theory of dopamine function and cognitive deficits in schizophrenia. Schizophrenia Bulletin, 19(1), 85-104.

Cohen JD, Barch DM, Carter CS, & Servan-Schreiber D (1999). Schizophrenic deficits in the processing of context: Converging evidence from three theoretically motivated cognitive tasks. Journal of Abnormal Psychology, 108, 120-133.

Carter CS, Perlstein P, Ganguli R, Brar J, Mintun M, & Cohen JD (1998). Functional hypofrontality and working memory dysfunction in schizophrenia. American Journal of Psychiatry, 155(9), 1285-1287.

Perlstein WM, Carter CS, Noll DC, Cohen JD. (2001). Relation of prefrontal cortex dysfunction to working memory and symptoms in schizophrenia. American Journal of Psychiatry, 158(7), 1105-1113.

Barch DM, Carter CS, Braver TS, Sabb FW, MacDonald A, Noll DC & Cohen JD (2001). Selective deficits in prefrontal cortex function in medication naïve patients with schizophrenia. Archives of General Psychiatry, 58, 280-8.

Perlstein WM, Dixit NK, Carter CS, Noll DC & Cohen JD (2003). Prefrontal cortex dysfunction mediates deficits in working memory and prepotent responding in schizophrenia. Biological Psychiatry, 58, 25-38.

Kerns JG, Cohen JD, MacDonald AW, 3rd, Johnson MK, Stenger VA, Aizenstein H & Carter CS (2005). Decreased conflict- and error-related activity in the anterior cingulate cortex in subjects with schizophrenia. American Journal of Psychiatry, 162(10), 1833-9.

MacDonald III A, Carter CS, Kerns J, Ursu S, Barch DM, Holmes A, Stenger VA & Cohen JD (2005). Specificity of prefrontal dysfunction and context processing deficits to schizophrenia in a never medicated first-episode psychotic sample.  American Journal of Psychiatry, 162:475–484.

Methods (Behavioral, fMRI, EEG and Neural Network Modeling)

Cohen JD, Noll DC & Schneider W (1993). Functional Magnetic Resonance Imaging: Overview and methods for psychological research. Behavioral Research Methods, Instruments & Computers, 25(2), 101-113.

Cohen JD, MacWhinney B, Flatt M & Provost J (1993). PsyScope: A new graphic interactive environment for designing psychology experiments. Behavioral Research Methods, Instruments & Computers, 25(2), 257-271.

Noll DC, Cohen JD, Meyer CH, Schneider W (1995). Spiral k-space MR imaging of cortical activation. Journal of Magnetic Resonance Imaging, 45, 49-56.

Goddard N, Hood G, Cohen JD, Eddy WF, Genovese CR, Noll DC, & Nystrom LE. (1997). Online analysis of functional MRI datasets on parallel platforms. Journal of Supercomputing, 11, 295-318.

Genovese CR, Noll DC, Cohen JD & Eddy WF (1997). Estimating test-retest reliability in functional MR imaging I: Statistical methodology. Magnetic Resonance in Medicine, 38, 497-507.

Noll DC, Genovese CR, Nystrom L, Vazquez A, Forman SD, Eddy WF & Cohen JD (1997). Estimating test-retest reliability in functional MR imaging II: Application to motor and cognitive activation studies. Magnetic Resonance in Medicine, 38, 508-517.

Fissell K, Tseytlin E, Cunningham D, Karthickeyan I, Carter CS, Schneider W & Cohen JD (2002). Fiswidgets: A graphical computing environment for neuroimaging analysis. Neuroinformatics, 1, 111-125.

Montague PR, Berns GS, Cohen JD, McClure SM, Pagnoni G, Dhamala M, Wiest MC, Karpov I, King RD, Apple N, Fisher RE (2002). Hyperscanning: Simultaneous fMRI during linked social interactions. NeuroImage, 16(4), 1159-64.

 Yeung N, Bogacz R, Holroyd C & Cohen JD (2004). Detection of synchronized oscillations in the electroencephalogram:  An evaluation of methods.  Psychophysiology, 41, 822-832.

Bogacz R & Cohen JD (2004).  Parameterization of connectionist models.  Behavioral Research Methods, Instruments & Computers,  36 (4), 732-741.

Yeung N, Bogacz R, Holroyd CB, Nieuwenhuis S & Cohen JD (2007).  Theta phase resetting and the error-related negativity.   Psychophysiology, 44, 39–49.