Adding prediction risk to the theory of reward learning
Open accessAbstract
This article analyzes the simple Rescorla-Wagner learning rule from the vantage point of least squares learning theory. In particular, it suggests how measures of risk, such as prediction risk, can be used to adjust the learning constant in reinforcement learning. It argues that prediction risk is most effectively incorporated by scaling the prediction errors. This way, the learning rate needs adjusting only when the covariance between optimal predictions and past (scaled) prediction errors changes. Evidence is discussed that suggests that the dopaminergic system in the (human and nonhuman) primate brain encodes prediction risk, and that prediction errors are indeed scaled with prediction risk (adaptive encoding).
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When some amount of uncertainty is expected, learning rates should decrease.
“When some amount of uncertainty is expected (also referred to as outcome variance or risk), learning rates should decrease.10–14”
From Serotonin neurons modulate learning rate through uncertainty (2022) · cites this paper as [11]
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“In addition to the rapid glutamatergic (and GABAergic) signaling between layers in a cortical hierarchy, there are slower neuormodulatory effects, for example, dopamine and acetylcholine appear to code a confidence estimate or uncertainty in predictions and prediction errors (Preuschoff and Bossaerts, 2007; Yu and Day…”
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“But another important finding was that bilateral anterior insula also showed an ongoing hemodynamic response to the variance in payoff (i.e., risk) and to an error signal related to this variance (also see Preuschoff and Bossaerts, 2007; Knutson and Bossaerts, 2007).”
BackgroundSurprise! Neural correlates of Pearce–Hall and Rescorla–Wagner coexist within the brain
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“In fact, in order to determine the correct value of an uncertain outcome, it is necessary to evaluate the risk, which is then used to estimate the probability of a possible reward (Preuschoff and Bossaerts, 2007).”
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“…prediction parallels previous work demonstrating that humans (Behrens et al., 2007; Speekenbrink & Shanks, 2010) and animals (Dayan et al., 2000; Courville et al., 2006; Preuschoff & Bossaerts, 2007) modulate their learning rates in response to the volatility of changes in associational structure.”
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“The precision weight is also updated with every trial and can be regarded as equivalent to a dynamic learning rate in reward learning models (Preuschoff and Bossaerts, 2007).”
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“To avoid unstable learning, it is thus essential to compare PEs to the expected fluctuation in reward value (Preuschoff and Bossaerts, 2007), and update values more when PEs with higher precision are encountered.”
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“reward feedback: (1) to determine the optimal learning rates based on optimization or on the level of uncertainty in a given environment (RL models) (Doya, 2002; Schweighofer and Doya, 2003; Preuschoff and Bossaerts, 2007); (2) to identify the correct model of the world, which includes the amount of uncertainty to pro…”
Neural computations of threat in the aftermath of combat trauma
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“…rt and the estimated predicted reward ̂vt : (3) Empirical data and theoretical considerations suggest that reward prediction error is scaled by predicted risk in the brain (Preushoff and Bossaerts, 2007; Tobler et al., 2005), allowing the learning rate to be independent of the payoff variance.”
InfluentialNeural Correlates of Anticipation Risk Reflect Risk Preferences
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“, the expected outcome variance, also plays a significant role in reward learning because it represents a predictability measure (Preuschoff and Bossaerts, 2007).”
BackgroundScaling prediction errors to reward variability benefits error-driven learning in humans
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“Thus far, it is unclear whether scaling of prediction errors relative to the variability of reward distributions results in improved performance, as predicted by learning models (Preuschoff and Bossaerts 2007).”
BackgroundInfluentialComputational models of adaptive behavior and prefrontal cortex
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“…of research into the inner workings of the human brain and have subsequently made appeals to brain images common in contemporary discussions on just about everything from the causes of gambling addiction (Preuschoff and Bossaerts, 2007) to debates about why music stirs the emotions (Koelsch, 2005).”
BackgroundPerceptual Classification in a Rapidly Changing Environment
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“However, an observer who is using a statistical decision strategy will revise this estimate more when category variance is low than high (Preuschoff and Bossaerts, 2007).”
Balancing new against old information: the role of puzzlement surprise in learning
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“…errors (Pearce & Hall, 1980; Hayden, Heilbronner, Pearson, & Platt, 2011; Roesch, Esber, Li, Daw, & Schoenbaumn, 2012), risk prediction errors (Preuschoff & Bossaerts, 2007; Preuschoff, Quartz, & Bossaerts, 2008), uncertainty-based jump detection (Nassar, Wilson, Heasly, & Gold, 2010;…”
ResultDelusions and prediction error: clarifying the roles of behavioural and brain responses
Corlett, P. R., & Fletcher, P. C. (2015) · Cognitive Neuropsychiatry · doi:10.1080/13546805.2014.990625 · Cited by 63 · see citing works
“Over time, a haphazard error signal – even a strong one – will become discounted – it will not contribute to updating (Preuschoff & Bossaerts, 2007).”
BackgroundNegative and interactive effects of sex, aging, and alcohol abuse on gray matter morphometry
Thayer, R. E., Hagerty, S. L., Sabbineni, A., Claus, E. D., Hutchison, K. E., & Weiland, B. J. (2016) · Human Brain Mapping · doi:10.1002/hbm.23172 · Cited by 61 · see citing works
“Given the role of the insula in reward learning [Preuschoff and Bossaerts, 2007] and interoception, our results suggest that with increased age, heavy alcohol exposure might lead to deficits in self awareness and increase sensitivity to physical sensations of craving [Goldstein et al., 2009].”
ResultIigaya, K., Ahmadian, Y., Sugrue, L. P., Corrado, G. S., Loewenstein, Y., Newsome, W. T., & Fusi, S. (2019) · Nature Communications · doi:10.1038/s41467-019-09388-3 · Cited by 58 · see citing works
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Supplementary Eye Field Encodes Reward Prediction Error
So, N., & Stuphorn, V. (2012) · Journal of Neuroscience · doi:10.1523/jneurosci.4419-11.2012 · Cited by 56 · see citing works
“Theoretical models have suggested that it would be useful to track prediction risk (i.e., the expected size of a prediction error) (Yu and Dayan, 2005; Preuschoff and Bossaerts, 2007), as it could be used to optimize the reaction to the prediction error.”
BackgroundMethodSchultz, W. (2009) · Elsevier eBooks · doi:10.1016/b978-0-12-374176-9.00021-x · Cited by 54 · see citing works
Liraglutide restores impaired associative learning in individuals with obesity
Hanssen, R., Rigoux, L., Kuzmanovic, B., Iglesias, S., Kretschmer, A. C., Schlamann, M., Albus, K., Edwin Thanarajah, S., Sitnikow, T., Melzer, C., et al. (2023) · Nature Metabolism · doi:10.1038/s42255-023-00859-y · Cited by 52 · see citing works
“4Faculty of Medicine and University Hospital Cologne, Department I of Internal Medicine, Center for Integrated Oncology Aachen Bonn Cologne Duesseldorf (CIO ABCD) and Excellence Center for Medical Mycology (ECMM), University of Cologne, Cologne, Germany.”
MethodInfluentialAdaptive coding of reward prediction errors is gated by striatal coupling
Park, S. Q., Kahnt, T., Talmi, D., Rieskamp, J., Dolan, R. J., & Heekeren, H. R. (2012) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1119969109 · Cited by 48 · see citing works
“Adaptive coding of PEs enables an enhanced discriminability to remain sensitive for changes of all sizes, which optimizes efficient coding in neural circuits with given limitations, namely, its firing ranges (28).”
BackgroundAdvanced Reinforcement Learning
Daw, N. D. (2013) · Elsevier eBooks · doi:10.1016/b978-0-12-416008-8.00016-4 · Cited by 44 · see citing works
“One way to understand this effect (due to Preuschoff and Bossaerts, 2007) is to assume that the neurons report not the raw prediction error δk but instead the prediction error scaled by the learning rate, κkUδk, which is, anyway, the quantity that should ultimately control the amount of plasticity…”
BackgroundA new predictive coding model for a more comprehensive account of delusions
Harding, J. N., Wolpe, N., Brugger, S. P., Navarro, V., Teufel, C., & Fletcher, P. C. (2024) · The Lancet Psychiatry · doi:10.1016/s2215-0366(23)00411-x · Cited by 45 · see citing works
Wang, Y., Liu, Y., Yang, L., Gu, F., Li, X., Zha, R., Wei, Z., Pei, Y., Zhang, P., Zhou, Y., et al. (2015) · Scientific Reports · doi:10.1038/srep10534 · Cited by 42 · see citing works
Dopamine, inference, and uncertainty
Gershman, S. J. (2017) · Neural Computation · doi:10.1162/neco_a_01023 · Cited by 40 · see citing works
Nassar, M. R., & Gold, J. I. (2013) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1003015 · Cited by 40 · see citing works
Risk and Reward Preferences under Time Pressure
Nursimulu, A., & Bossaerts, P. L. (2013) · European Finance Review · doi:10.1093/rof/rft013 · Cited by 40 · see citing works
Stolyarova, A., & Izquierdo, A. (2017) · eLife · doi:10.7554/elife.27483 · Cited by 39 · see citing works
Mice infer probabilistic models for timing
Li, Y., & Dudman, J. T. (2013) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1310666110 · Cited by 36 · see citing works
“By contrast, strong sensitivity to recent trials is suboptimal in tasks where there is substantial uncertainty about reward delivery (16).”
BackgroundResultRisks and rewards in adolescent decision-making
van Duijvenvoorde, A. C. K., van Hoorn, J., & Blankenstein, N. E. (2022) · Current Opinion in Psychology · doi:10.1016/j.copsyc.2022.101457 · Cited by 41 · see citing works
“…past functional neuroimaging studies have identified multiple brain regions associated with making decisions involving risk [43,44]: The (posterior) parietal cortex is thought to code probability [45] and its gray matter density has been related to individual differences in risk preference [46,47].”
BackgroundDynamic Estimation of Task-Relevant Variance in Movement under Risk
Landy, M. S., Trommershäuser, J., & Daw, N. D. (2012) · Journal of Neuroscience · doi:10.1523/jneurosci.6160-11.2012 · Cited by 35 · see citing works
“) Note that this will be the qualitative direction of update even in a variance-sensitive version of the mean-tracking approach in which the variance estimate controls the step size for the update (Dayan and Long, 1998; Dayan et al., 2000; Courville et al., 2006; Preuschoff and Bossaerts, 2007).”
BackgroundInfluentialAltered cingulo-striatal function underlies reward drive deficits in schizophrenia
Park, I. H., Chun, J., Park, H., Koo, M., Park, S., Kim, S.-H., & Kim, J. (2014) · Schizophrenia Research · doi:10.1016/j.schres.2014.11.005 · Cited by 34 · see citing works
“Likewise, risk prediction is also incorporated in the reinforcement learning process in which the trade-off between expected reward and risk determines behavior (Preuschoff and Bossaerts, 2007).”
BackgroundMultiple Mechanisms for Processing Reward Uncertainty in the Primate Basal Forebrain
Ledbetter, N. M., Chen, C. D., & Monosov, I. E. (2016) · Journal of Neuroscience · doi:10.1523/jneurosci.1123-16.2016 · Cited by 33 · see citing works
“…interact with other brain areas that are modulated by reward uncertainty (Hsu et al., 2005; McCoy and Platt, 2005; Preuschoff et al., 2006, 2008; Preuschoff and Bossaerts, 2007; O’Neill and Schultz, 2010; Burke and Tobler, 2011a; Schultz et al., 2011; Monosov and Hikosaka, 2012, 2013; Monosov…”
BackgroundThe Neurobiological Foundations of Valuation in Human Decision Making Under Uncertainty
Bossaerts, P. L., Preuschoff, K., & Hsu, M. (2009) · Elsevier eBooks · doi:10.1016/b978-0-12-374176-9.00023-3 · Cited by 32 · see citing works
“The intuition has a rigorous underpinning [see (Preuschoff and Bossaerts 2007)].”
BackgroundBai, Y., Katahira, K., & Ohira, H. (2014) · Frontiers in Psychology · doi:10.3389/fpsyg.2014.00871 · Cited by 29 · see citing works
“Several Bayesian approaches (Kakade and Dayan, 2002; Courville et al., 2006; Preuschoff and Bossaerts, 2007) suggested that after a surprising event, animals should pay more attention to stimuli and that these events provoke the animals into learning faster.”
BackgroundO’Connell, L. A., Fontenot, M. R., & Hofmann, H. A. (2013) · Journal of Chemical Neuroanatomy · doi:10.1016/j.jchemneu.2012.12.007 · Cited by 29 · see citing works
“On the other hand, the VTA is involved in risk-taking and reward evaluation (Schultz, 1998; Tobler et al., 2005; Preuschoff and Bossaerts, 2007) as well as the processing of nociception (Sotres-Bayón et al.”
Relative salience signaling within a thalamo-orbitofrontal circuit governs learning rate
Namboodiri, V. M. K., Hobbs, T. G., Trujillo‐Pisanty, I., Simon, R. C., Hjort, M. M., & Stuber, G. D. (2021) · Current Biology · doi:10.1016/j.cub.2021.09.037 · Cited by 25 · see citing works
Neural substrates of updating the prediction through prediction error during decision making
Wang, Y., Ma, N., He, X., Li, N., Wei, Z., Yang, L., Zha, R., Han, L., Li, X., Zhang, D., et al. (2017) · NeuroImage · doi:10.1016/j.neuroimage.2017.05.041 · Cited by 27 · see citing works
“This model included prediction errors for both reward and risk, following the proposition of Preushoff and Bossaerts (Preuschoff and Bossaerts, 2007).”
Models of neuromodulation for computational psychiatry
Iglesias, S., Tomiello, S., Schneebeli, M., & Stephan, K. E. (2016) · Wiley Interdisciplinary Reviews Cognitive Science · doi:10.1002/wcs.1420 · Cited by 26 · see citing works
Flannery, J. S., Riedel, M. C., Bottenhorn, K. L., Poudel, R., Salo, T., Hill-Bowen, L. D., Laird, A. R., & Sutherland, M. T. (2019) · Cognitive Affective & Behavioral Neuroscience · doi:10.3758/s13415-019-00763-7 · Cited by 25 · see citing works
“…influenced by external contextual factors (Behrens et al., 2007; Fiorillo, Tobler & Schultz, 2007; Hariri et al., 2006; Porcelli & Delgado, 2009; Preuschoff & Bossaerts, 2007; Rangel, 2008; Seaman et al., 2018), and/or by subjective internal factors (Hetherington, Pirie & Nabb, 2002; Juechems…”
Does reward unpredictability reflect risk?
Anselme, P. (2014) · Behavioural Brain Research · doi:10.1016/j.bbr.2014.12.003 · Cited by 25 · see citing works
Surprise beyond prediction error
Chumbley, J. R., Burke, C. J., Stephan, K. E., Friston, K. J., Tobler, P. N., & Fehr, E. (2014) · Human Brain Mapping · doi:10.1002/hbm.22513 · Cited by 23 · see citing works
“In this context, conventional model-free RPE learning algorithms (Rescorla and Wagner, 1972) can only learn the average or “expected” reward that is constant over cues, while more recent theories permit subjects to discriminate based on reward variance, risk or precision (Preuschoff and Bossaerts, 2007; Schultz et al.…”
BackgroundInfluentialLearning and Generalization under Ambiguity: An fMRI Study
Chumbley, J. R., Flandin, G., Bach, D. R., Jean Daunizeau, A., Fehr, E., Dolan, R. J., & Friston, K. J. (2012) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1002346 · Cited by 20 · see citing works
“While recent RL developments in neuroscience incorporate some notion of uncertainty [21], learning and generalization are typically non-probabilistic.”
BackgroundToward a Mechanistic Understanding of Human Decision Making
O’Doherty, J. P., & Bossaerts, P. L. (2008) · Current Directions in Psychological Science · doi:10.1111/j.1467-8721.2008.00560.x · Cited by 19 · see citing works
“This parameter should change with the uncertainty in the environment (Behrens, Woolrich, Walton, & Rushworth, 2007; Preuschoff & Bossaerts, 2007; Yu & Dayan, 2003).”
BackgroundElectrophysiological Signatures of Hierarchical Learning
Liu, M., Dong, W., Qin, S., Verguts, T., & Chen, Q. (2021) · Cerebral Cortex · doi:10.1093/cercor/bhab245 · Cited by 15 · see citing works
Synthetic surprise as the foundation of the psychedelic experience
Filippo, R. D., & Schmitz, D. (2024) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2024.105538 · Cited by 15 · see citing works
“…corresponding to precision, modulates the influence of prediction errors on updating priors, thereby adjusting the learning rate from current sensory information (Preuschoff et al., 2006; Preuschoff and Bossaerts, 2007; Nassar et al., 2012; Mcguire et al., 2014; Diederen and Schultz, 2015).”
BackgroundAnswering some phenomenal challenges to the prediction error model of delusions
Corlett, P. R. (2015) · World Psychiatry · doi:10.1002/wps.20211 · Cited by 16 · see citing works
Gender Differences in Behavioral and Neural Responses to Unfairness Under Social Pressure
Zheng, L., Ning, R., Li, L., Wei, C., Cheng, X. M., Zhou, C., & Guo, X. (2017) · Scientific Reports · doi:10.1038/s41598-017-13790-6 · Cited by 15 · see citing works
Formalizing the Function of Anterior Insula in Rapid Adaptation
Bossaerts, P. L. (2018) · Frontiers in Integrative Neuroscience · doi:10.3389/fnint.2018.00061 · Cited by 15 · see citing works
“There, the learning rate increases, and hence, learning accelerates, when an outcome is dissimilar to outcomes experienced in the past; it decreases if surprise is low (Pearce and Hall, 1980; Behrens et al., 2007; Preuschoff and Bossaerts, 2007; Payzan-LeNestour et al., 2013; McGuire et al., 2014).”
BackgroundWang, B., & Pleger, B. (2020) · Cerebral Cortex · doi:10.1093/cercor/bhaa073 · Cited by 14 · see citing works
Emotion dynamics as hierarchical Bayesian inference in time
Majumdar, G., Yazin, F., Banerjee, A., & Roy, D. (2022) · Cerebral Cortex · doi:10.1093/cercor/bhac305 · Cited by 13 · see citing works
Wei, Z., Han, L., Zhong, X., Liu, Y., Zha, R., Wang, Y., Yang, L., Bu, J., Song, H., Wang, W., et al. (2017) · NeuroImage · doi:10.1016/j.neuroimage.2017.11.048 · Cited by 12 · see citing works
“RPEs were included in this model, following the suggestion by Preushoff and Bossaert (Preuschoff and Bossaerts, 2007).”
Wang, B., Schlaffke, L., & Pleger, B. (2020) · Journal of Neuroscience · doi:10.1523/jneurosci.2904-19.2020 · Cited by 11 · see citing works
“At the second level of the HGF, the precision weight (c 2) is updated with every trial, so that it also can be considered as the equivalence of a dynamic learning rate in reward-learning models (Preuschoff and Bossaerts, 2007; Mathys et al., 2011).”
BackgroundModulation of Dopamine for Adaptive Learning: a Neurocomputational Model
Inglis, J. B., Valentin, V. V., & Ashby, F. G. (2020) · Computational Brain & Behavior · doi:10.1007/s42113-020-00083-x · Cited by 9 · see citing works
The Experimental Manipulation of Uncertainty
Bach, D. R., Pryce, C. R., & Seifritz, E. (2010) · Neuromethods · doi:10.1007/978-1-60761-883-6_8 · Cited by 8 · see citing works
Organization of neuropeptide systems in the human brain
Ceballos, E. G., Farahani, A., Liu, Z.-Q., Milisav, F., Hansen, J. Y., Dagher, A., & Mišić, B. (2026) · Nature Neuroscience · doi:10.1038/s41593-026-02236-w · Cited by 7 · see citing works
Liu, M., Dong, W., Wu, Y., Verbeke, P., Verguts, T., & Chen, Q. (2022) · Cerebral Cortex · doi:10.1093/cercor/bhac352 · Cited by 6 · see citing works
Predictive coding and hallucinations: a question of balance
Fletcher, P. C. (2017) · Cognitive Neuropsychiatry · doi:10.1080/13546805.2017.1391083 · Cited by 7 · see citing works
“Or should we learn to ignore our errors in prediction, as would be needful in a setting that was simply noisy and unpredictable (Preuschoff & Bossaerts, 2007)?”
Background
- Authors
- PREUSCHOFF, K., & BOSSAERTS, P.
- Affiliations
- California Institute of Technology (US)
- Corresponding author
- BOSSAERTS, PETER
- Venue
- Annals of the New York Academy of Sciences
- Publisher
- Wiley
- Published
- 2007-05
- Volume
- 1104
- Issue
- 1
- Pages
- 135-146
- ISSN
- 0077-8923 (ISSN-L)
- 1749-6632
- OpenAlex
- W2136500228
- Cited by
- 133
- References
- 22
- Citations by year
- 2026: 4 · 2025: 2 · 2024: 7 · 2023: 3 · 2022: 5 · 2021: 5 · 2020: 4 · 2019: 5 · 2018: 4 · 2017: 13 · 2016: 7 · 2015: 8 · 2014: 9 · 2013: 10 · 2012: 10
- FWCI
- 4.66
- Type
- journal article
- Language
- en
- Open access
- green
- infoscience.epfl.ch
- Licence
- Locations
- Annals of the New York Academy of Sciences (publishedVersion)
- PubMed (publishedVersion)
- CaltechAUTHORS (California Institute of Technology) (submittedVersion)
- Infoscience (Ecole Polytechnique Fédérale de Lausanne) (submittedVersion, cc-by-nc-nd)
- Infoscience (Ecole Polytechnique Fédérale de Lausanne) (submittedVersion)
- Funders
- Swiss Finance Institute
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural and Behavioral Psychology Studies
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurotransmitter Receptor Influence on Behavior
- Physical Sciences › Computer Science › Artificial Intelligence › Reinforcement Learning in Robotics
- Keywords
- prediction risk, reward learning, adaptive learning rate, dopaminergic system, adaptive coding, reinforcement learning, primate brain
- MeSH terms
- Animals, Association Learning, Brain, Brain Mapping, Conditioning, Classical, Dopamine, Humans, Learning, Neurons, Probability, Reinforcement, Psychology, Reward, Risk, Least-Squares Analysis
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Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
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