Introduction Experiment Conclusion Bayesian Learning under Three Kinds of Uncertainty: Risk, Estimation Uncertainty, and Unexpected Uncertainty Élise Payzan-LeNestour Australian School of Business, UNSW Sydney October 2010 Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation Natural sampling tasks: decision maker explores (“samples”) reward prospects and learns about their values Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation Natural sampling tasks: decision maker explores (“samples”) reward prospects and learns about their values Model-based (Bayesian) learning or model-free (RL) learning Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation Natural sampling tasks: decision maker explores (“samples”) reward prospects and learns about their values Model-based (Bayesian) learning or model-free (RL) learning ? Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation Natural sampling tasks: decision maker explores (“samples”) reward prospects and learns about their values Model-based (Bayesian) learning or model-free (RL) learning ? Both modes coexist in the brain (e.g., Balleine ea 2005, Gläscher ea 2010); the mode that influences behavior is the one that is more adapted to the current situation (Daw ea 2005) Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation Natural sampling tasks: decision maker explores (“samples”) reward prospects and learns about their values Model-based (Bayesian) learning or model-free (RL) learning ? Both modes coexist in the brain (e.g., Balleine ea 2005, Gläscher ea 2010); the mode that influences behavior is the one that is more adapted to the current situation (Daw ea 2005) → Question is irrelevant: when the two modes are algorithmically the same; see the Kalman Filter (Aoki 1987) when answer already known: Model-free RL does as well as model-based learning in the long run; model-based outperforms in the transient period (Balleine ea 2005, Daw ea 2005) Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation Natural sampling tasks: decision maker explores (“samples”) reward prospects and learns about their values Model-based (Bayesian) learning or model-free (RL) learning ? Both modes coexist in the brain (e.g., Balleine ea 2005, Gläscher ea 2010); the mode that influences behavior is the one that is more adapted to the current situation (Daw ea 2005) → Question is irrelevant: when the two modes are algorithmically the same; see the Kalman Filter (Aoki 1987) when answer already known: Model-free RL does as well as model-based learning in the long run; model-based outperforms in the transient period (Balleine ea 2005, Daw ea 2005) Here question asked in the context of unstable natural sampling Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation (cont.) Unstable sampling tasks: reward probabilities jump over time so decision maker is continuously experiencing transient periods Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation (cont.) Unstable sampling tasks: reward probabilities jump over time so decision maker is continuously experiencing transient periods In such contexts even most sophisticated kind of model-free RL can’t perform as well as Bayesian learning (Courville ea 2006, Choi ea 2009) Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation (cont.) Unstable sampling tasks: reward probabilities jump over time so decision maker is continuously experiencing transient periods In such contexts even most sophisticated kind of model-free RL can’t perform as well as Bayesian learning (Courville ea 2006, Choi ea 2009) So Bayesian shall control behavior Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation (cont.) Unstable sampling tasks: reward probabilities jump over time so decision maker is continuously experiencing transient periods In such contexts even most sophisticated kind of model-free RL can’t perform as well as Bayesian learning (Courville ea 2006, Choi ea 2009) So Bayesian shall control behavior ... to the extent that the brain can implement it! Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation (cont.) Unstable sampling tasks: reward probabilities jump over time so decision maker is continuously experiencing transient periods In such contexts even most sophisticated kind of model-free RL can’t perform as well as Bayesian learning (Courville ea 2006, Choi ea 2009) So Bayesian shall control behavior ... to the extent that the brain can implement it! Bayesian learning complex here: Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation (cont.) Unstable sampling tasks: reward probabilities jump over time so decision maker is continuously experiencing transient periods In such contexts even most sophisticated kind of model-free RL can’t perform as well as Bayesian learning (Courville ea 2006, Choi ea 2009) So Bayesian shall control behavior ... to the extent that the brain can implement it! Bayesian learning complex here: requires assessment of Risk (Expected Uncertainty), Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation (cont.) Unstable sampling tasks: reward probabilities jump over time so decision maker is continuously experiencing transient periods In such contexts even most sophisticated kind of model-free RL can’t perform as well as Bayesian learning (Courville ea 2006, Choi ea 2009) So Bayesian shall control behavior ... to the extent that the brain can implement it! Bayesian learning complex here: requires assessment of Risk (Expected Uncertainty), jump likelihood (Unexpected Uncertainty) (Yu&Dayan 2005), Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation (cont.) Unstable sampling tasks: reward probabilities jump over time so decision maker is continuously experiencing transient periods In such contexts even most sophisticated kind of model-free RL can’t perform as well as Bayesian learning (Courville ea 2006, Choi ea 2009) So Bayesian shall control behavior ... to the extent that the brain can implement it! Bayesian learning complex here: requires assessment of Risk (Expected Uncertainty), jump likelihood (Unexpected Uncertainty) (Yu&Dayan 2005), and Estimation Uncertainty (Ambiguity) Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation (cont.) Unstable sampling tasks: reward probabilities jump over time so decision maker is continuously experiencing transient periods In such contexts even most sophisticated kind of model-free RL can’t perform as well as Bayesian learning (Courville ea 2006, Choi ea 2009) So Bayesian shall control behavior ... to the extent that the brain can implement it! Bayesian learning complex here: requires assessment of Risk (Expected Uncertainty), jump likelihood (Unexpected Uncertainty) (Yu&Dayan 2005), and Estimation Uncertainty (Ambiguity) combined Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Motivation (cont.) Unstable sampling tasks: reward probabilities jump over time so decision maker is continuously experiencing transient periods In such contexts even most sophisticated kind of model-free RL can’t perform as well as Bayesian learning (Courville ea 2006, Choi ea 2009) So Bayesian shall control behavior ... to the extent that the brain can implement it! Bayesian learning complex here: requires assessment of Risk (Expected Uncertainty), jump likelihood (Unexpected Uncertainty) (Yu&Dayan 2005), and Estimation Uncertainty (Ambiguity) combined May the human brain approximate it? Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Evidence for Bayesian Learning Behavioral evidence exists but with rat subjects (Gallistel ea 2001) Neural evidence exists for separate encoding of the three levels of uncertainty (e.g., Preuschoff ea 2006-2008, Hsu ea 2005, Yoshida ea 2006, Huettel ea 2006, Rutishauser ea 2006, Behrens ea 2007, Den Ouden ea, 2010, Watson ea 2007) But levels studied separately or without independent control of the three levels (Behrens ea 2007); unclear whether human brain can tease apart representations of Risk and Unexpected Uncertainty which are antagonistic (Yu&Dayan 2005) Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Evidence for Bayesian Learning Behavioral evidence exists but with rat subjects (Gallistel ea 2001) Neural evidence exists for separate encoding of the three levels of uncertainty (e.g., Preuschoff ea 2006-2008, Hsu ea 2005, Yoshida ea 2006, Huettel ea 2006, Rutishauser ea 2006, Behrens ea 2007, Den Ouden ea, 2010, Watson ea 2007) But levels studied separately or without independent control of the three levels (Behrens ea 2007); unclear whether human brain can tease apart representations of Risk and Unexpected Uncertainty which are antagonistic (Yu&Dayan 2005) This study: provides behavioral and neural evidence in a six-armed bandit task where the arms jump over time Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Restless Bandit Task Six-armed bandit in which reward processes jump over time Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Two ways to learn option values in this task Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Two ways to learn option values in this task Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Two ways to learn option values in this task 1 Model-based (Bayesian): tracks hidden outcome contingencies by detecting jumps on the spot → quick to adapt Requires to tease apart Risk, Estimation Uncertainty and Unexpected Uncertainty 2 Model-free RL: predicts next outcome from observation of past outcome; purely correlational → slower to adapt (“backward looking”) Representation of uncertainty either absent (Rescorla-Wagner) or monolithic (Pearce-Hall) Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Evidence for Bayesian Learning Sought at Two Levels Behavior: Bayesian and RL learning produce different behaviors in the task ⇒ Did subjects act more like Bayesians? Note: Bayesian and RL use the same exploration policy – softmax choice rule (Ishii ea 2005) Imaging: Truly Bayesian metrics that RL would ignore ⇒ Do we see neural activation correlating with these metrics? Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Behavioral Evidence for Bayesian Learning Bayesian model better predicts behavior than does RL (62 subjects, 500 choices per subject) (Payzan 2010) Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Behavioral Evidence for Bayesian Learning Bayesian model better predicts behavior than does RL (62 subjects, 500 choices per subject) (Payzan 2010) Subjects directed exploration towards best known options i.e., were ambiguity-averse more Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Behavioral Evidence for Bayesian Learning Bayesian model better predicts behavior than does RL (62 subjects, 500 choices per subject) (Payzan 2010) Subjects directed exploration towards best known options i.e., were ambiguity-averse more Behavioral marker of Bayesian learning (no representation of ambiguity under RL) Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Behavioral Evidence for Bayesian Learning Bayesian model better predicts behavior than does RL (62 subjects, 500 choices per subject) (Payzan 2010) Subjects directed exploration towards best known options i.e., were ambiguity-averse more Behavioral marker of Bayesian learning (no representation of ambiguity under RL) Neural markers? ⇒ Examine whether activation correlating with unexpected uncertainty, estimation uncertainty, and risk Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Imaging Study Adapted the task: same stochastic structure, same information, but only 2 options proposed for choice on each trial design Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Imaging Study Adapted the task: same stochastic structure, same information, but only 2 options proposed for choice on each trial design Replicated previous behavioral results (17 subjects, 260 choices per subject) Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Imaging Study Adapted the task: same stochastic structure, same information, but only 2 options proposed for choice on each trial design Replicated previous behavioral results (17 subjects, 260 choices per subject) GLM with 4 onset regressors: cue (phasic), cue (tonic), outcome (phasic), outcome (tonic) Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Imaging Study Adapted the task: same stochastic structure, same information, but only 2 options proposed for choice on each trial design Replicated previous behavioral results (17 subjects, 260 choices per subject) GLM with 4 onset regressors: cue (phasic), cue (tonic), outcome (phasic), outcome (tonic) Included (model-derived) Unexpected Uncertainty, Estimation Uncertainty, and Risk signals as parametric modulators at cue and at outcome Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Imaging Study Adapted the task: same stochastic structure, same information, but only 2 options proposed for choice on each trial design Replicated previous behavioral results (17 subjects, 260 choices per subject) GLM with 4 onset regressors: cue (phasic), cue (tonic), outcome (phasic), outcome (tonic) Included (model-derived) Unexpected Uncertainty, Estimation Uncertainty, and Risk signals as parametric modulators at cue and at outcome GLM Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Imaging Study Adapted the task: same stochastic structure, same information, but only 2 options proposed for choice on each trial design Replicated previous behavioral results (17 subjects, 260 choices per subject) GLM with 4 onset regressors: cue (phasic), cue (tonic), outcome (phasic), outcome (tonic) Included (model-derived) Unexpected Uncertainty, Estimation Uncertainty, and Risk signals as parametric modulators at cue and at outcome GLM Look at activations across the group Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Main Results (preliminary) Risk signals in anterior insula (Huettel ea 2006, Preuschoff ea 2006, Christopoulos ea 2009,...) Ambiguity signals in ACC (Behrens ea 2007) , right superior temporal lobule, bilateral MFG (Huettel ea 2006, Gläscher ea 2010) Unexpected Uncertainty signals in vmPFC (Hampton ea 2006, Den Ouden ea 2010), parahippocampal gyrus (Rutishauser ea 2006), post cingulate and ACC (anterior to Behrens ea 2007; Cf. Den Ouden ea 2010), anterior insula (Watson ea, 2007) Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Main Results (preliminary) Risk signals in anterior insula (Huettel ea 2006, Preuschoff ea 2006, Christopoulos ea 2009,...) Ambiguity signals in ACC (Behrens ea 2007) , right superior temporal lobule, bilateral MFG (Huettel ea 2006, Gläscher ea 2010) Unexpected Uncertainty signals in vmPFC (Hampton ea 2006, Den Ouden ea 2010), parahippocampal gyrus (Rutishauser ea 2006), post cingulate and ACC (anterior to Behrens ea 2007; Cf. Den Ouden ea 2010), anterior insula (Watson ea, 2007) images Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Results (cont.) Attentional (salience and executive control) processes in amygdala, IFG (Corbetta ea 2000, MacDonald ea 2000, Gläscher ea 2010) , superior temporal lobule (Yantis ea 2002, Gläscher ea 2010) learning rate Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Bringing the Three Kinds of Uncertainty Together Behavioral Evidence for Bayesian Learning Neural Evidence for Bayesian Learning Results (cont.) Attentional (salience and executive control) processes in amygdala, IFG (Corbetta ea 2000, MacDonald ea 2000, Gläscher ea 2010) , superior temporal lobule (Yantis ea 2002, Gläscher ea 2010) learning rate Part of vmPFC covaries with Bayesian expected value (Tricomi ea) expected value Ventral striatum responds to value of realized outcome (O’Doherty ea 2002, McClure ea 2004-2007,...) Élise Payzan-LeNestour reward Society for Neuroeconomics Introduction Experiment Conclusion Summary Discussion Conclusion Found neural correlates of Risk, Unexpected Uncertainty, and Estimation Uncertainty in ant insula, ACC, lat PFC and superior parietal Since 1 Correlation of neural activity with Bayesian uncertainty signals: not parasitic on general attentional mechanisms (orienting or executive control) 2 Encoding of the three categories of uncertainty is quintessentially Bayesian (RL ignores them) ⇒ Neural evidence for Bayesian learning in the task Strengthens the behavioral evidence Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Summary Discussion Boundaries of Bayesian Learning? Follow-up experiment: same boardgame but with a fourth level of uncertainty added: Model (or Knigtian) Uncertainty (Knight 1921, Keynes 1921, Basili ea 2009,...) Parameter uncertainty (Ambiguity) vs. Model Uncertainty Bayesian learning becomes inefficient in this case (Draper 1995, Diaconis&Freedman 1985) According to Daw ea 2005, people should fall back to RL Confirmed by our data Élise Payzan-LeNestour Society for Neuroeconomics Introduction Experiment Conclusion Summary Discussion Main Message of This Study We found that Bayesian learning controlled behavior when Estimation Uncertainty reduced to Ambiguity Evaporated when Estimation Uncertainty further meant Model Uncertainty ⇒ Decision making under Model Uncertainty does not represent a special more complex case of decision making under Ambiguity Élise Payzan-LeNestour Society for Neuroeconomics Acknowledgments Collaborators Peter Bossaerts Simon Dunne John O’Doherty Funding Science Foundation Ireland, Wellcome Trust NCCR Finrisk and Swiss Finance Institute. Behavioral Evidence for Bayesian Learning 900 850 800 750 -LL of FB 700 650 600 550 500 450 400 460 510 560 610 660 -LL of RL 710 760 810 860 Legend: Comparative fits for each subject (point below 45 degree line indicates that Bayesian model fits better) Evidence for ambiguity aversion 900 -LL of Ambiguity-Averse FB 800 700 600 500 400 300 200 400 450 500 550 600 650 700 750 800 850 -LL of standard FB With “Ambiguity-Averse” model, option value = expected value penalty proportional to level of estimation uncertainty back 900 fMRI Design Cue presentation and choice (2s) Waiting stage (4s) Outcome presentation (1.5s) back "# GLM $%&' ()*+,-$. $%&' (478-$. 7%4$7:&' 7%4$7:&' ()*+,-$. (478-$. 7%4$7:&' ;+5%& "/ 0-,1 "/ "/ 2(9. 23 23 40-+5'46)& 40-+5'46)&' 33 Note: Uncertainty signals orthogonalized relative to learning rate (LR): LR meant to pick up general attentional activation (salience and executive control) back Expected value p<.01, SVC at (9, 45, -13): vmPFC back Outcome delivery outcome (phasic) p<.001 (uncorrected): ventral striatum, mPFC back Neural Correlates of Learning rate left ht p<.001 (uncorrected): right cerebellum, left MFG, left amygdala back Risk signal sic) p<.001 (uncorrected): left anterior insula back Ambiguity signals p<.001 (uncorrected): anterior cingulate, right superior parietal lobule, bilateral MFG, bilateral occipital lobe, left precuneus Unexpected Uncertainty signals %49" 8&)9" 5%49" &" 38 p<.001: posterior left cingulate, vmPFC, bilateral insula, caudate, bilateral parahippocampal gyrus, anterior STG, right inferior parietal lobule
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