1. The Physics of Decision-Making: Cognitive Control as the Optimization of Behavior Gary Aston-Jones ∞ Rafal Bogacz * † ª Eric Brown † Jonathan D. Cohen * ª Mark Gilzenrat * Philip Holmes † Patrick Simen ª *Department of Psychology ªCenter for the Study of Brain, Mind and Behavior † Program in Applied and Computational Mathematics Princeton University ∞ Department of Psychiatry, University of Pennsylvania Silvio Conte Center for Neuroscience Research, NIMH

2. Cognitive Control Definition: The ability to flexibly guide decision making and behavior in accord with internally represented goals or intentions

3. Time Scales of Decision Making & Control Single decisions: 100s of milleseconds - seconds “Should I swing the bat?” Real time dynamics of information integration and decision making Adaptive regulation of decision making: seconds - minutes “Was that last swing too fast? Should I wait longer this time?” Adaptive adjustment of decision making parameters Learning Long term decisions, prospective control: hours - years “Should I get some coaching help with my batting, or just retire?” Strategies

4. Time Scales of Decision Making & Control Single decisions: 100s of milleseconds - seconds “Should I swing the bat?” Real time dynamics of information integration and decision making Adaptive regulation of decision making: seconds - minutes “Was that last swing too fast? Should I wait longer this time?” Adaptive adjustment of decision making parameters Learning Long term decisions, prospective control: hours - years “Should I get some coaching help with my batting, or just retire?” Strategies ☞ ☞

5. Allocate Control Evaluate Outcomes Performance Control: Adaptive Regulation of Behavior Adjust Monitor Select / Bias

6. Allocate Control Prefrontal Areas Evaluate Outcomes Performance Poseterior Frontal, Parietal & Temporal Areas Control: Adaptive Regulation of Behavior Select / Bias Monitor Adjust

7. Control Response PFC Bias Studied in a variety of tasks – Selective attention tasks – Response inhibition tasks – Working memory tasks – Task switching Computational models have specified some of the mechanisms involved – Prefrontal cortex and attentional control (Cohen et al., 1990; Cohen & Servan-Schreiber, 1992) – Dopamine / basal ganglia and updating of task representations (Braver & Cohen, 2000; Frank, Loughry & O’Reilly, 2004) – Task switching (Gilbert & Shallice, 2002; Yeung & Monsell, 2003) Control: Adaptive Regulation of Behavior

8. Allocate Control Prefrontal Areas Evaluate Outcomes Oribitofrontal Anterior Cingulate Performance Poseterior Frontal, Parietal & Temporal Areas Adjust Monitor Control: Adaptive Regulation of Behavior

9. Studied in a variety of tasks – Simple reaction time tasks – Two alternative forced choice decision tasks – Attention tasks – Learning tasks Computational models have specified some of the mechanisms involved – Reinforcement learning (Montague et al., 1996) – Conflict monitoring (Botvinick et al., 2001) – Mismatch detection (Holroyd et al., 2002) Monitoring and Adjustments of Control Control Response Conflict Monitoring Anterior Cingulate Modulation of Control PFC Reinforcement Learning / Gating VTA

10. Lack of a single coherent framework for understanding control: – Multiple models; each addresses a different task How do their mechanisms relate to and interact with one another? –Parameterization problems –What are the fundamental principles of operation? –Relationship of neural implementation to behavior Need a more precise, formal definition of control… Shortcomings

11. Control as Optimization Refine definition of control: –adjustment of processing parameters to optimize task performance –assumes processing mechanisms are capable of (near) optimal function

12. Control as Optimization Refine definition of control: –adjustment of processing parameters to optimize task performance –assumes processing mechanisms are capable of (near) optimal function Precedence: This approach has been used productively in a variety of fields: -  Economics — utility maximization (standard economic model, homo economicus) -  Behavioral ecology — evolution of social behavior (e.g., Nowak, Boyd)  -  Psychology — rational analysis (John Anderson ) -  Neuroscience — perceptual, motor and learning systems (e.g., Barlow, Bialek, Gallistel)

13. Control as Optimization Refine definition of control: –adjustment of processing parameters to optimize task performance –assumes processing mechanisms are capable of (near) optimal function Precedence: This approach has been used productively in a variety of fields: -  Economics — utility maximization (standard economic model, homo economicus) -  Behavioral ecology — evolution of social behavior (e.g., Nowak, Boyd)  -  Psychology — rational analysis (John Anderson ) -  Neuroscience — perceptual, motor and learning systems (e.g., Barlow, Bialek, Gallistel) However: Scarcity of work that uses this approach to address cognitive control, and bridge between behavior and its neural mechanisms

14. Outline Simple behavioral task: – Two alternative forced choice (2AFC) decision task Current “state of play:” – Behavioral findings – Neurobiological findings – Neural network models Formal analysis: – Drift diffusion model (DDM) of decision making – Control as optimization of the DDM – Predictions and behavioral findings

15. Two Alternative Force Choice Task Are the dots moving left or right?

16. <> See dots moving left (<), press the left button See dots moving right (>), press the right button Measure reaction time (RT) and accuracy Two Alternative Force Choice Task Are the dots moving left or right?

17. Characteristically skewed reaction time distribution: Speed / accuracy tradeoff: fast responding → less accurate more accurate → slower responding Behavioral Findings Probability Reaction Time

18. Neural Findings Area MT (temporal cortex) motion sensitive visual cortex Areas LIP (parietal cortex) and SEF (supplementary eye fields): control of eye movements

19. Simple Neural Network Model of Two Alternative Decision Task Information Competition Processing –Flow of activity from stimulus inputs to a pair decision units –Each decision unit accumulates / integrates input –Decision units compete Decision –Occurs when activity of one decision unit exceeds a specified threshold >< Integration Usher & McClelland, 2001

20. Some Problems with Neural Network Models Large parameter space: difficult to parameterize –How to set stimulus strength, connection weights, thresholds, etc. Complex dynamics: hard to characterize and compare Theoretical degeneracy: proliferation of models…

21. Some Problems with Neural Network Models Large parameter space: difficult to parameterize –How to set stimulus strength, connection weights, thresholds, etc. Complex dynamics: hard to characterize and compare Theoretical degeneracy: proliferation of models…

22. Simplification & Analysis Activity of R 2 Attractors (stable fixed points, R 1 correct) Step 1: Construct geometric representation of model’s behavior Activity of R 1 Threshold for R 2 Threshold for R 1 >< R1R1 R2R2

23. Step 2: Examine Dynamics Activity of R 1 Activity of R 2 Threshold for R 2 Threshold for R 1 Simplification & Analysis

24. Step 2: Examine Dynamics Activity of R 1 Activity of R 2 Threshold for R 2 Threshold for R 1 Simplification & Analysis

25. Threshold for R 2 Step 3: Note that there are two components of the trajectory… Activity of R 1 Activity of R 2 Threshold for R 1 Decision (difference in activity) Co-activation (sum of activity) Simplification & Analysis

26. Threshold for R 2 and that they have different dynamics Activity of R 1 Activity of R 2 Threshold for R 1 Decision ➠ slower Co-activation ➠ faster

27. Focus only on one dimension Brown & Holmes (Stochastics and Dynamics, 2001) –assume that most of the “action” is along the decision line –therefore, decision process can be approximated by a one-dimensional process (difference in activity) Simplification & Analysis Activity of R 1 Activity of R 2 Decision Co-activation (faster) Step 4: Dimensional Reduction

28. Focus only on one dimension Brown & Holmes (Stochastics and Dynamics, 2001) –assume that most of the “action” is in along the decision line –therefore, decision process can be approximated by a one-dimensional process (difference in activity) Focus on linear range of function –assume that units in the “focus of attention” are on the linear part of their activation function (i.e., most sensitive part of their dynamic range) Cohen et al (Psychological Review, 1990) Simplification & Analysis Activity of R 1 Activity of R 2 Decision Co-activation (faster) Step 5: Linearization

29. Simplification Drift Diffusion Model (DDM): x = A + c A = drift rate c = noise P(x,t) = N(At, c√t) Process ends when x exceeds ± z Activity of R 1 Activity of R 2 x -Z +Z

30. Simplification Drift Diffusion Model (DDM): –This process is described by the Fokker-Planck equation for the evolution of a Gaussian probability distribution toward a pair of boundaries = mathematical description of diffusion of an ideal gas x = A + c A = drift rate c = noise P(x,t) = N(At, c√t) Process ends when x exceeds ± z Drift (A) Diffusion (c) Response 2Response 1 Thresholds Activity of R 1 Activity of R 2 x -Z +Z +z+z -z-z Early Middle Late

31. Simplification Drift Diffusion Model (DDM): x = A + c A = drift rate c = noise P(x,t) = N(At, c√t) Process ends when x exceeds ± z Drift (A) Diffusion (c) Response 2Response 1 Thresholds Activity of R 1 Activity of R 2 x -Z +Z +z+z -z-z Early Middle Late

32. Simplification Drift Diffusion Model (DDM): Can analytically solve for Error Rate and Decision Time: x = A + c A = drift rate c = noise P(x,t) = N(At, c√t) Process ends when x exceeds ± z Error Rate (ER) = Decision Time (DT) = Tanh( ) Threshold Drift DriftThreshold Noise 2 1 + e 2DriftThreshold Noise 2 1 Drift (A) Diffusion (c) Response 2Response 1 Thresholds Activity of R 1 Activity of R 2 x -Z +Z +z+z -z-z Early Middle Late

33. Evidence Threshold +Z Threshold -Z Probability Reaction Time Accurately describes reaction time distributions and error rates in simple decision making tasks (Ratcliff, 1978, 1999) Drift Diffusion Model Drift (A)

34. Accurately describes reaction time distributions and error rates in simple decision making tasks (Ratcliff, 1978, 1999) Accurately describes dynamics of firing among stimulus- and response-selective neurons in simple decision making tasks (Schall, 1994; Gold & Schadlen, 2002) Drift Diffusion Model

35. Theoretical Traction Formal reduction of neural network models: (Bogacz et al., 2006)

36. Theoretical Traction Formal reduction of neural network models (Bogacz et al., 2006) Optimal decision making process –Formally equivalent to sequential probability ratio test (SPRT) (used by Turing to crack German Enigma code in WWII) –Fastest to reach a decision for given threshold and error rate and most accurate for a given decision time (Wald, 1948; Turing [Good, 1979]; Rouder, 1996) –Guarantees arbitrarily low error rate as threshold is increased (Bogacz et al., 2006)

37. Theoretical Traction Formal reduction of neural network models (Bogacz et al., 2006) Optimal decision making process –Formally equivalent to sequential probability ratio test (SPRT) (used by Turing to crack German Enigma code in WWII) –Fastest to reach a decision for given threshold and error rate and most accurate for a given decision time (Wald, 1948; Turing [Good, 1979]; Rouder, 1996) –Guarantees arbitrarily low error rate as threshold is increased (Bogacz et al., 2006) However, presents optimization problem of its own: –How to set parameters (e.g., threshold and starting point)? –Here is where control comes in…

38. DDM and Control DDM specifies psychologically relevant control parameters –Starting point: expectations (priors) –Drift rate: signal strength / attention –Threshold: speed-accuracy trade-off

39. DDM and Control DDM specifies psychologically relevant control parameters –Starting point: expectations (priors) –Drift rate: signal strength / attention –Threshold: speed-accuracy trade-off Empirical question: –Do people in fact adjust these parameters to optimize performance?

40. DDM and Control DDM specifies psychologically relevant control parameters –Starting point: expectations (priors) –Drift rate: signal strength / attention –Threshold: speed-accuracy trade-off Empirical question: –Do people adjust these parameters to optimize performance? –We can analyze the DDM to determine optimal parameters under various experimental conditions, and use this to generate testable predictions. For example, what is the optimal threshold, and do people use this? –First, however, must assume an “objective (utility) function” — that is, the function that control seeks to optimize

41. Reward rate: 1-Error Rate Reaction Time + Delay Reward Rate Optimization RR =

42. Reward rate: Re-express RT and ER in terms of DDM parameters: 1-Error Rate Reaction Time + Delay Reward Rate Optimization RR = Error Rate (ER) = Decision Time (DT) = Tanh( ) Threshold Drift DriftThreshold Noise 2 1 + e 2DriftThreshold Noise 2 1

43. Reward rate: Re-express RT and ER in terms of DDM parameters: RR = 1-Error Rate Reaction Time + Delay Reward Rate Optimization - Delay - Delay- Drift Threshold Drift Threshold e 2DriftThreshold Noise 2 RR = Error Rate (ER) = Decision Time (DT) = Tanh( ) Threshold Drift DriftThreshold Noise 2 1 + e 2DriftThreshold Noise 2 1

44. Reward rate: Re-express RT and ER in terms of DDM parameters: RR = Solve for threshold that maximizes RR: 1-Error Rate Reaction Time + Delay Reward Rate Optimization - Delay - Delay- Drift Threshold Drift Threshold e 2DriftThreshold Noise 2 Drift * Delay 2 Delay 2 Thresh*Drift = Noise 2 Drift 2 Drift NoiseDelay RR =

45. Reward rate: Re-express RT and ER in terms of DDM parameters: RR = Solve for threshold that maximizes RR: Predict changes in speed-accuracy tradeoff (threshold) as a function of task parameters (delay, drift, and noise) (Bogacz et al., 2006) Reward Rate Optimization Drift * Delay 2 Delay 2 Thresh*Drift = Noise 2 Drift 2 Drift NoiseDelay 1-Error Rate Reaction Time + Delay - Delay - Delay- Drift Threshold Drift Threshold e 2DriftThreshold Noise 2 RR =

46. Reward rate (and ∴ optimal threshold) varies with total delay: where “ ” (D total )= + (ER*D pen ) Predictions: Effects of Delay 1-Error Rate + Reward Rate =

47. Reward rate (and ∴ optimal threshold) varies with total delay: where “ ” (D total )= + (ER*D pen ) ⇒ Optimal threshold is same for the following two conditions: ITI = 0.5 secITI = 2 sec D pen = 1.5 secNo D pen D total = 2 sec and 1-Error Rate + Reward Rate = Effects of Delay

48. Empirical Results Reaction Time Error Rates Thresholds Effects of Delay

49. If one stimulus is more frequent than the other, it is optimal to move the starting point, not the threshold assuming constant drift (SNR) Predictions: Effects of Stimulus Frequency threshold B threshold A Stimulus A = 50%Stimulus B = 50% Stimulus A = 25% Stimulus B = 75% Starting Point

50. Effects of Stimulus Frequency threshold B threshold A Stimulus A = 50%Stimulus B = 50% Stimulus A = 25% Stimulus B = 75% Stimulus A = 10% Stimulus B = 90% For sufficiently extreme frequencies: –the optimal starting point exceeds the optimal threshold –the model predicts a switch from integration to stereotyped responding –the stimulus frequency at which this occurs varies according to delay and drift Starting Point

51. Effects of Stimulus Frequency Empirical Data

52. The fact that there is a single optimal threshold for a given set of task parameters means that the DDM equations: – Error Rate (ER) = – Decision Time (DT) = – Reward Rate (RR) = can be solved for DT as a function of ER: Optimal Performance Curve - Delay -Delay- Drift Threshold Drift Threshold e 2DriftThreshold Noise 2 Tanh( ) Threshold Drift DriftThreshold Noise 2 1 + e 2DriftThreshold Noise 2 1

53. The fact that there is a single optimal threshold for a given set of task parameters means that the DDM equations: – Error Rate (ER) = – Decision Time (DT) = – Reward Rate (RR) = can be solved for DT as a function of ER: In other words, there is a single, optimal speed-accuracy curve that should quantitatively define performance… Optimal Performance Curve - Delay -Delay- Drift Threshold Drift Threshold e 2DriftThreshold Noise 2 Tanh( ) Threshold Drift DriftThreshold Noise 2 1 + e 2DriftThreshold Noise 2 1

54. Optimal Performance Curve ER DT (mean normalized) Theoretical Prediction

55. Optimal Performance Curve ER DT (mean normalized) Empirical Data

56. Optimal Performance Curve ER DT (mean normalized) Empirical Data accuracy weight increasing

57. Optimal Performance Curve ER DT (mean normalized) Empirical Data accuracy weight increasing

58. Summary Drift Diffusion Model: –explains human RT distributions and accuracy in simple decision tasks –explains dynamics of neural firing in simple decision tasks –formally equivalent to neural network models of simple decision tasks –describes parameters of optimal performance (maximizing reward rate) –predicts influence of task parameters on speed-accuracy tradeoff that approximate those observed empirically Defines, in formal and principled terms, the mechanisms underlying decision making and its interaction with evaluation in simple two alternative forced choice tasks Defines, in formal and principled terms, variables that are subject to regulation by control mechanisms to optimize outcomes

59. Current Directions Probe neural mechanisms of control (fMRI/EEG): –Information integration process (posterior and frontal mechanisms) –Outcome monitoring (OFC? ACC?) –Threshold adjustment and starting point biases (BG? Supplementary Motor Areas?) –Drift control / attention (PFC?) Extend to more interesting behavioral domains –Competition / selection tasks — time varying drift (attentional control) –Multi-choice decisions Contact with other normative approaches –Bayesian theory: optimal computation –Information theoretic approaches: distribution free analyses

60. Acknowledgements Investigators: Rafal Bogacz - Bristol University Eric Brown - NYU Patrick Simen - Princeton Phil Holmes - Princeton Jeff Moehlis - UC Santa Barbara Tyler McMillen - Princeton Phil Eckhoff - Princeton Angela Yu - Princeton Neuroscience of Cognitive Control Laboratory Department of Psychology Program in Applied and Computational Mathematics Center for the Study of Brain, Mind and Behavior Princeton University Funding Support: NIMH NSF Center for the Study of Brain, Mind and Behavior