1. The brain basis of cognitive control Todd Braver Cognitive Control and Psychopathology Lab, Washington University St. Louis

2. I do cognitive neuroscience Cognitive Psychology+Neuroscience = Cognitive Neuroscience What is that?

3. Cognitive Psychology+Neuroscience = Cognitive Neuroscience Study of knowledge and thinking What is cognitive neuroscience?

4. Cognitive Psychology+Neuroscience = Cognitive Neuroscience What is cognitive neuroscience? Study of how the brain works

5. Cognitive Psychology+Neuroscience = Cognitive Neuroscience What is cognitive neuroscience? Study of how the brain thinks

6. The mystery of control William James on will (1890) “...wherever movement follows unhesitatingly and immediately the notion of it in the mind, we have ideo-motor action. We are then aware of nothing between the conception and the execution. In contrast, some acts require will, such that an additional conscious element in the shape of a fiat, mandate, or expressed consent is involved” The mind/body and homunculus problems –Is there a “controller” in our mind/brain –What is it? Is it made out of the same neural “stuff” of the rest of the brain –That’s what I want to find out: Why I am a cognitive neuroscientist »A bit on my background… This is the domain of “cognitive control” (also called self-control, executive control) –Some subtopics: behavioral inhibition, sustained attention, working memory (goal- maintenance), planning, multi-tasking »Real world examples –How can these be studied in the lab?

7. Time for some demos! Tap your desk if the letter you see is anything but X If it is an X, DO NOT RESPOND Tap as fast as you can (it’s rapid paced)….

8. P CXK T FMBQXLH

9. Now let’s do some Stroop! Name the color of the Ink, ignore word name Answer as fast as you can….

10. YELLOW

11. BLUE

12. RED

13. YELLOW

14. RED

15. BLUE

16. RED

17. YELLOW

18. Now some working memory Study word names, hold on to them over delay Check if the next word matches one of them –If so, say YES (quickly) –If not, say NO (quickly)

19. + NAIL TRAIN GRAPE SHIRTMOUSE NAIL

20. + GRATE ROCK STEEL FANGBREAD TRAIN

21. What do these demos illustrate? First one: go-nogo –Behavioral inhibition: How can we stop ourselves from the impulse to respond to the X? –What causes us to fail (make an error)? How do we monitor & adjust performance based on failure? Second one: Stroop –Selective attention: How do we override the automatic tendency to attend to word name, even when irrelevant? –How do we respond and adjust to interference between word & color? Again, performance monitoring Third one: Short-term item recognition –Working memory: How do we maintain information in an accessible form during delay? –How do we decide which trial the target (probe) item belongs to? Current or previous? Using working memory to prevent sources of interference (i.e., familiarity)

22. Brain Mechanisms of Cognitive Control Studies of performance monitoring –What are brain circuits and computations that determine when control is needed? –i.e., to detect likelihood of errors, compensate for errors, or deal with sources of interference Studies of motivation –Why don’t people always exert optimal cognitive control? Does it have a cost? –How do increases in the (reward) value of cognitive control modify brain activity and behavior?

23. Gehring et al., 1993 The ERN -- an error-specific brainwave –ERP: event-related (brain electrical) potential

24. Gehring et al., 1993 Compensatory Behaviors Error Correction Subsequent Performance ERN magnitude -> What is the inference from these findings?

25. Where is the ERN located in the brain? The anterior cingulate cortex Dorsal/Caudal Rostral/Ventral

26. Carter et al., 1998 Conflict Monitoring theory The key points of the theory: Errors are a special case of high conflict ACC detects conflicts rather than errors per se High Conflict Low Conflict What does conflict mean? But also this was found in the same ACC region

27. The Conflict-Control Loop What is the point of performance monitoring in the ACC? –To provide signals that indicate when control processes need to be adjusted –Control state needs to adapt to environmental demands & contingencies »Low interference = low control (unbiased or bottom-up attention) »High interference = high control (focused attention) Dorsal ACC & PFC form a feedback loop Dynamically adjusts control in response to experienced conflict (Botvinick et al., 2001 Psych Review)

28. How does PFC implement control? Dorsolateral Ventrolateral Active maintenance (Store words over delay) Of cognitive goals (Name color; nogo for X) To bias attention & action (Filter out words; slow down responding)

29. Computational Modeling of Conflict-Control Loop (Flanker task) H H S H H S S S S S Compatible Incompatible

30. Evidence for ACC-PFC loop (Kerns et al., 2003 ) ACC activity increases following conflict (incongruent) or error trials Increased PFC activity and decreased interference on subsequent trial

31. Take-Home Points Performance monitoring & cognitive control might be implemented in the brain through division of labor –ACC: When are control demands high? –PFC: Modify control state (attention, goals, active maintenance) based on ACC demand signal ACC might respond to conflict rather than errors per se –Conflict could be a good signal of control demand –Doesn’t require knowledge of error This is a very active area of research! –Lots of new ideas, findings & theoretical accounts

32. Motivation & Cognitive Control How come people seem to perform better when there is a lot at stake? –Offering rewards seems to optimize behavior (e.g., gold stars, contests) –Does it work by modulating cognitive control state in the brain (e.g., PFC)? People seem to vary in their motivational levels –Some people seem to be very motivated by the prospect of rewards, others not »Intrinsic motivation vs. extrinsic motivation –Do ‘reward-driven’ people do better on cognitive tasks, when rewards are present? »Is it because they are more likely to utilize motivation to modify cognitive control? We can answer these questions using brain imaging methods Functional Magnetic Resonance Imaging (fMRI)

33. fMRI Scanning

34. FMRI Principles Increased Neural Response Increased Local Cerebral Blood Flow Decreased deoxyhemoglobin Change in Magnetic Susceptibility Change in fMRI Signal

35. Neuroimaging Activation Maps

36. Event-related fMRI -0.1 0 0.1 0.2 0.3 0.4 012345678910111213141516171819 PERCENT MR SIGNAL TIME (SEC) Blood flow responses can be detected (and isolated) for individual events of brief duration

37. Motivation Study design Verbal WM task

38. Performance & Reward- sensitivity Group Average Individual Differences These are effects on non-incentive trials! A motivational context effect

39. Neural effects of motivation Jimura, Locke, and Braver (2010) Right DLPFC No Reward Reward Sustained activity Trial-related activity Motivational context shifts PFC activity dynamics from transient (late-trial) to sustained (+ early- trial). What about individual differences?

40. Reward Sensitivity Behavioral Effects Of Motivational Context No Reward Reward Shift in PFC activity dynamics statistically explains Motivation-Personality-Behavior association Right DLPFC Individual Differences Jimura, Locke, & Braver (2010) PNAS

41. A causal model Jimura, Locke, and Braver (in preparation) High Reward Sensitivity -> Proactive Control = Better Performance Low Reward Sensitivity -> Reactive Control = Poorer Performance

42. Take-Home Points Motivation seems to modulate cognitive control mode –Proactive (sustained) mode might be optimized for obtaining rewards –Sustained control helps maintain task goals and attention Individual differences –People that seem to care more about rewards are more likely to shift modes (PFC activity) Cognitive control might be costly –Might be penalties for using sustained control (effort biases) »But Why? –The way that control is implemented could be dependent upon estimates of task reward value »Incentives might increase task reward value more for highly reward sensitive people –ACC might also be involved here (determining cost vs. benefit of control)