<?xml version="1.0" ?>
<!DOCTYPE PubmedArticleSet PUBLIC "-//NLM//DTD PubMedArticle, 1st January 2025//EN" "https://dtd.nlm.nih.gov/ncbi/pubmed/out/pubmed_250101.dtd">
<PubmedArticleSet>
<PubmedArticle><MedlineCitation Status="MEDLINE" Owner="NLM" IndexingMethod="Automated"><PMID Version="1">39779974</PMID><DateCompleted><Year>2025</Year><Month>04</Month><Day>29</Day></DateCompleted><DateRevised><Year>2025</Year><Month>05</Month><Day>20</Day></DateRevised><Article PubModel="Print-Electronic"><Journal><ISSN IssnType="Electronic">1471-0048</ISSN><JournalIssue CitedMedium="Internet"><Volume>26</Volume><Issue>3</Issue><PubDate><Year>2025</Year><Month>Mar</Month></PubDate></JournalIssue><Title>Nature reviews. Neuroscience</Title><ISOAbbreviation>Nat Rev Neurosci</ISOAbbreviation></Journal><ArticleTitle>The curious case of dopaminergic prediction errors and learning associative information beyond value.</ArticleTitle><Pagination><StartPage>169</StartPage><EndPage>178</EndPage><MedlinePgn>169-178</MedlinePgn></Pagination><ELocationID EIdType="doi" ValidYN="Y">10.1038/s41583-024-00898-8</ELocationID><Abstract><AbstractText>Transient changes in the firing of midbrain dopamine neurons have been closely tied to the unidimensional value-based prediction error contained in temporal difference reinforcement learning models. However, whereas an abundance of work has now shown how well dopamine responses conform to the predictions of this hypothesis, far fewer studies have challenged its implicit assumption that dopamine is not involved in learning value-neutral features of reward. Here, we review studies in rats and humans that put this assumption to the test, and which suggest that dopamine transients provide a much richer signal that incorporates information that goes beyond integrated value.</AbstractText><CopyrightInformation>&#xa9; 2025. This is a U.S. Government work and not under copyright protection in the US; foreign copyright protection may apply.</CopyrightInformation></Abstract><AuthorList CompleteYN="Y"><Author ValidYN="Y"><LastName>Kahnt</LastName><ForeName>Thorsten</ForeName><Initials>T</Initials><Identifier Source="ORCID">0000-0002-3575-2670</Identifier><AffiliationInfo><Affiliation>Intramural Research Program, National Institute on Drug Abuse, Baltimore, MD, USA. thorsten.kahnt@nih.gov.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Schoenbaum</LastName><ForeName>Geoffrey</ForeName><Initials>G</Initials><Identifier Source="ORCID">0000-0001-8180-0701</Identifier><AffiliationInfo><Affiliation>Intramural Research Program, National Institute on Drug Abuse, Baltimore, MD, USA. schoenbg@schoenbaumlab.org.</Affiliation></AffiliationInfo></Author></AuthorList><Language>eng</Language><PublicationTypeList><PublicationType UI="D016428">Journal Article</PublicationType><PublicationType UI="D016454">Review</PublicationType></PublicationTypeList><ArticleDate DateType="Electronic"><Year>2025</Year><Month>01</Month><Day>08</Day></ArticleDate></Article><MedlineJournalInfo><Country>England</Country><MedlineTA>Nat Rev Neurosci</MedlineTA><NlmUniqueID>100962781</NlmUniqueID><ISSNLinking>1471-003X</ISSNLinking></MedlineJournalInfo><ChemicalList><Chemical><RegistryNumber>VTD58H1Z2X</RegistryNumber><NameOfSubstance UI="D004298">Dopamine</NameOfSubstance></Chemical></ChemicalList><CitationSubset>IM</CitationSubset><MeshHeadingList><MeshHeading><DescriptorName UI="D006801" MajorTopicYN="N">Humans</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D000818" MajorTopicYN="N">Animals</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D059290" MajorTopicYN="Y">Dopaminergic Neurons</DescriptorName><QualifierName UI="Q000502" MajorTopicYN="N">physiology</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D004298" MajorTopicYN="Y">Dopamine</DescriptorName><QualifierName UI="Q000378" MajorTopicYN="N">metabolism</QualifierName><QualifierName UI="Q000502" MajorTopicYN="N">physiology</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D012201" MajorTopicYN="Y">Reward</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D012054" MajorTopicYN="N">Reinforcement, Psychology</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D001245" MajorTopicYN="Y">Association Learning</DescriptorName><QualifierName UI="Q000502" MajorTopicYN="N">physiology</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D007858" MajorTopicYN="Y">Learning</DescriptorName><QualifierName UI="Q000502" MajorTopicYN="N">physiology</QualifierName></MeshHeading></MeshHeadingList><CoiStatement>Competing interests: The authors declare no competing interests.</CoiStatement></MedlineCitation><PubmedData><History><PubMedPubDate PubStatus="accepted"><Year>2024</Year><Month>12</Month><Day>11</Day></PubMedPubDate><PubMedPubDate PubStatus="medline"><Year>2025</Year><Month>2</Month><Day>22</Day><Hour>0</Hour><Minute>21</Minute></PubMedPubDate><PubMedPubDate PubStatus="pubmed"><Year>2025</Year><Month>1</Month><Day>9</Day><Hour>0</Hour><Minute>22</Minute></PubMedPubDate><PubMedPubDate PubStatus="entrez"><Year>2025</Year><Month>1</Month><Day>8</Day><Hour>23</Hour><Minute>35</Minute></PubMedPubDate></History><PublicationStatus>ppublish</PublicationStatus><ArticleIdList><ArticleId IdType="pubmed">39779974</ArticleId><ArticleId IdType="doi">10.1038/s41583-024-00898-8</ArticleId><ArticleId IdType="pii">10.1038/s41583-024-00898-8</ArticleId></ArticleIdList><ReferenceList><Reference><Citation>Mirenowicz, J. &amp; Schultz, W. Importance of unpredictability for reward responses in primate dopamine neurons. J. Neurophysiol. 72, 1024&#x2013;1027 (1994).</Citation><ArticleIdList><ArticleId IdType="pubmed">7983508</ArticleId></ArticleIdList></Reference><Reference><Citation>Schultz, W. Getting formal with dopamine and reward. Neuron 36, 241&#x2013;263 (2002).</Citation><ArticleIdList><ArticleId IdType="pubmed">12383780</ArticleId></ArticleIdList></Reference><Reference><Citation>Sutton, R. S. &amp; Barto, A. G. Reinforcement Learning: An Introduction (MIT Press, 2018).</Citation></Reference><Reference><Citation>Rescorla, R. A. &amp; Wagner, A. R. in Classical Conditioning II: Current Research and Theory (eds Black, A. H. &amp; Prokesy, W. F.) 64&#x2013;99 (Appleton-Century-Crofts, 1972).</Citation></Reference><Reference><Citation>Sutton, R. S. &amp; Barto, A. G. Toward a modern theory of adaptive networks: expectation and prediction. Psychol. Rev. 88, 135&#x2013;170 (1981).</Citation><ArticleIdList><ArticleId IdType="pubmed">7291377</ArticleId></ArticleIdList></Reference><Reference><Citation>Dayan, P. Improving generalization for temporal difference learning: the successor representation. Neural Comput. 5, 613&#x2013;624 (1993).</Citation></Reference><Reference><Citation>Lak, A., Stauffer, W. R. &amp; Schultz, W. Dopamine prediction error responses integrate subjective value from different reward dimensions. Proc. Natl Acad. Sci. USA 111, 2343&#x2013;2348 (2014).</Citation><ArticleIdList><ArticleId IdType="pubmed">24453218</ArticleId><ArticleId IdType="pmc">3926061</ArticleId></ArticleIdList></Reference><Reference><Citation>Tobler, P. N., Fiorillo, C. D. &amp; Schultz, W. Adaptive coding of reward value by dopamine neurons. Science 307, 1642&#x2013;1645 (2005).</Citation><ArticleIdList><ArticleId IdType="pubmed">15761155</ArticleId></ArticleIdList></Reference><Reference><Citation>Fiorillo, C. D., Tobler, P. N. &amp; Schultz, W. Discrete coding of reward probability and uncertainty by dopamine neurons. Science 299, 1898&#x2013;1902 (2003).</Citation><ArticleIdList><ArticleId IdType="pubmed">12649484</ArticleId></ArticleIdList></Reference><Reference><Citation>Roesch, M. R., Calu, D. J. &amp; Schoenbaum, G. Dopamine neurons encode the better option in rats deciding between differently delayed or sized rewards. Nat. Neurosci. 10, 1615&#x2013;1624 (2007).</Citation><ArticleIdList><ArticleId IdType="pubmed">18026098</ArticleId><ArticleId IdType="pmc">2562672</ArticleId></ArticleIdList></Reference><Reference><Citation>Schultz, W. Dopamine reward prediction-error signalling: a two-component response. Nat. Rev. Neurosci. 17, 183&#x2013;195 (2016).</Citation><ArticleIdList><ArticleId IdType="pubmed">26865020</ArticleId><ArticleId IdType="pmc">5549862</ArticleId></ArticleIdList></Reference><Reference><Citation>Watabe-Uchida, M., Eshel, N. &amp; Uchida, N. Neural circuitry of reward prediction error. Annu. Rev. Neurosci. 40, 373&#x2013;394 (2017).</Citation><ArticleIdList><ArticleId IdType="pubmed">28441114</ArticleId><ArticleId IdType="pmc">6721851</ArticleId></ArticleIdList></Reference><Reference><Citation>O&#x2019;Doherty, J. P., Dayan, P., Friston, K., Critchley, H. &amp; Dolan, R. J. Temporal difference models and reward-related learning in the human brain. Neuron 38, 329&#x2013;337 (2003).</Citation><ArticleIdList><ArticleId IdType="pubmed">12718865</ArticleId></ArticleIdList></Reference><Reference><Citation>D&#x2019;Ardenne, K., McClure, S. M., Nystrom, L. E. &amp; Cohen, J. D. BOLD responses reflecting dopaminergic signals in the human ventral tegmental area. Science 319, 1264&#x2013;1267 (2008).</Citation><ArticleIdList><ArticleId IdType="pubmed">18309087</ArticleId></ArticleIdList></Reference><Reference><Citation>Rutledge, R. B., Dean, M., Caplin, A. &amp; Glimcher, P. W. Testing the reward prediction error hypothesis with an axiomatic model. J. Neurosci. 30, 13525&#x2013;13536 (2010).</Citation><ArticleIdList><ArticleId IdType="pubmed">20926678</ArticleId><ArticleId IdType="pmc">2957369</ArticleId></ArticleIdList></Reference><Reference><Citation>Haber, S. N., Fudge, J. L. &amp; McFarland, N. R. Striatonigrostriatal pathways in primates form an ascending spiral from the shell to the dorsolateral striatum. J. Neurosci. 20, 2369&#x2013;2382 (2000).</Citation><ArticleIdList><ArticleId IdType="pubmed">10704511</ArticleId><ArticleId IdType="pmc">6772499</ArticleId></ArticleIdList></Reference><Reference><Citation>Fallon, J. H. &amp; Moore, R. Y. Catecholamine innervation of the basal forebrain. IV. Topography of the dopamine projection to the basal forebrain and neostriatum. J. Comp. Neurol. 180, 545&#x2013;580, (1978).</Citation><ArticleIdList><ArticleId IdType="pubmed">659674</ArticleId></ArticleIdList></Reference><Reference><Citation>Bjorklund, A. &amp; Dunnett, S. B. Dopamine neuron systems in the brain: an update. Trends Neurosci. 30, 194&#x2013;202 (2007).</Citation><ArticleIdList><ArticleId IdType="pubmed">17408759</ArticleId></ArticleIdList></Reference><Reference><Citation>Pessiglione, M., Seymour, B., Flandin, G., Dolan, R. J. &amp; Frith, C. D. Dopamine-dependent prediction errors underpin reward-seeking behaviour in humans. Nature 442, 1042&#x2013;1045 (2006).</Citation><ArticleIdList><ArticleId IdType="pubmed">16929307</ArticleId><ArticleId IdType="pmc">2636869</ArticleId></ArticleIdList></Reference><Reference><Citation>Knutson, B. et al. Amphetamine modulates human incentive processing. Neuron 43, 261&#x2013;269 (2004).</Citation><ArticleIdList><ArticleId IdType="pubmed">15260961</ArticleId></ArticleIdList></Reference><Reference><Citation>Schultz, W., Dayan, P. &amp; Montague, P. R. A neural substrate for prediction and reward. Science 275, 1593&#x2013;1599 (1997).</Citation><ArticleIdList><ArticleId IdType="pubmed">9054347</ArticleId></ArticleIdList></Reference><Reference><Citation>Glimcher, P. W. Understanding dopamine and reinforcement learning: the dopamine reward prediction error hypothesis. Proc. Natl Acad. Sci. USA 108, 15647&#x2013;15654 (2011).</Citation><ArticleIdList><ArticleId IdType="pubmed">21389268</ArticleId><ArticleId IdType="pmc">3176615</ArticleId></ArticleIdList></Reference><Reference><Citation>Kakade, S. &amp; Dayan, P. Dopamine: generalization and bonuses. Neural Netw. 15, 549&#x2013;559 (2002).</Citation><ArticleIdList><ArticleId IdType="pubmed">12371511</ArticleId></ArticleIdList></Reference><Reference><Citation>Starkweather, C. K. &amp; Uchida, N. Dopamine signals as temporal difference errors: recent advances. Curr. Opin. Neurobiol. 67, 95&#x2013;105 (2021).</Citation><ArticleIdList><ArticleId IdType="pubmed">33186815</ArticleId></ArticleIdList></Reference><Reference><Citation>Dabney, W. et al. A distributional code for value in dopamine-based reinforcement learning. Nature 577, 671&#x2013;675 (2020).</Citation><ArticleIdList><ArticleId IdType="pubmed">31942076</ArticleId><ArticleId IdType="pmc">7476215</ArticleId></ArticleIdList></Reference><Reference><Citation>Jeong, H. et al. Mesolimbic dopamine release conveys causal associations. Science 378, eabq6740 (2022).</Citation><ArticleIdList><ArticleId IdType="pubmed">36480599</ArticleId><ArticleId IdType="pmc">9910357</ArticleId></ArticleIdList></Reference><Reference><Citation>Coddington, L. T., Lindo, S. E. &amp; Dudman, J. T. Mesolimbic dopamine adapts the rate of learning from action. Nature 614, 294&#x2013;302 (2023).</Citation><ArticleIdList><ArticleId IdType="pubmed">36653450</ArticleId><ArticleId IdType="pmc">9908546</ArticleId></ArticleIdList></Reference><Reference><Citation>Kutlu, M. G. et al. Dopamine release in the nucleus accumbens core signals perceived saliency. Curr. Biol. 31, 4748&#x2013;4761.e8 (2021).</Citation><ArticleIdList><ArticleId IdType="pubmed">34529938</ArticleId><ArticleId IdType="pmc">9084920</ArticleId></ArticleIdList></Reference><Reference><Citation>Lee, R. S., Sagiv, Y., Engelhard, B., Witten, I. B. &amp; Daw, N. D. A feature-specific prediction error model explains dopaminergic heterogeneity. Nat. Neurosci. 27, 1574&#x2013;1586 (2024).</Citation><ArticleIdList><ArticleId IdType="pubmed">38961229</ArticleId></ArticleIdList></Reference><Reference><Citation>Takahashi, Y. K. et al. Dopamine neurons respond to errors in the prediction of sensory features of expected rewards. Neuron 95, 1395&#x2013;1405.e3 (2017).</Citation><ArticleIdList><ArticleId IdType="pubmed">28910622</ArticleId><ArticleId IdType="pmc">5658021</ArticleId></ArticleIdList></Reference><Reference><Citation>Howard, J. D. &amp; Kahnt, T. Identity prediction errors in the human midbrain update reward-identity expectations in the orbitofrontal cortex. Nat. Commun. 9, 1611 (2018).</Citation><ArticleIdList><ArticleId IdType="pubmed">29686225</ArticleId><ArticleId IdType="pmc">5913228</ArticleId></ArticleIdList></Reference><Reference><Citation>Boorman, E. D., Rajendran, V. G., O&#x2019;Reilly, J. X. &amp; Behrens, T. E. Two anatomically and computationally distinct learning signals predict changes to stimulus-outcome associations in hippocampus. Neuron 89, 1343&#x2013;1354 (2016).</Citation><ArticleIdList><ArticleId IdType="pubmed">26948895</ArticleId><ArticleId IdType="pmc">4819449</ArticleId></ArticleIdList></Reference><Reference><Citation>Suarez, J. A., Howard, J. D., Schoenbaum, G. &amp; Kahnt, T. Sensory prediction errors in the human midbrain signal identity violations independent of perceptual distance. eLife 8, e43962 (2019).</Citation><ArticleIdList><ArticleId IdType="pubmed">30950792</ArticleId><ArticleId IdType="pmc">6450666</ArticleId></ArticleIdList></Reference><Reference><Citation>Witkowski, P. P., Park, S. A. &amp; Boorman, E. D. Neural mechanisms of credit assignment for inferred relationships in a structured world. Neuron 110, 2680&#x2013;2690.e9 (2022).</Citation><ArticleIdList><ArticleId IdType="pubmed">35714610</ArticleId></ArticleIdList></Reference><Reference><Citation>Liu, Q. et al. Midbrain signaling of identity prediction errors depends on orbitofrontal cortex networks. Nat. Commun. 15, 1704 (2024).</Citation><ArticleIdList><ArticleId IdType="pubmed">38402210</ArticleId><ArticleId IdType="pmc">10894191</ArticleId></ArticleIdList></Reference><Reference><Citation>Millidge, B., Song, Y., Lak, A., Walton, M. E. &amp; Bogacz, R. Reward bases: a simple mechanism for adaptive acquisition of multiple reward types. PLoS Comput. Biol. 20, e1012580 (2024).</Citation><ArticleIdList><ArticleId IdType="pubmed">39561186</ArticleId><ArticleId IdType="pmc">11614280</ArticleId></ArticleIdList></Reference><Reference><Citation>Papageorgiou, G. K., Baudonnat, M., Cucca, F. &amp; Walton, M. E. Mesolimbic dopamine encodes prediction errors in a state-dependent manner. Cell Rep. 15, 221&#x2013;228 (2016).</Citation><ArticleIdList><ArticleId IdType="pubmed">27050518</ArticleId><ArticleId IdType="pmc">4835543</ArticleId></ArticleIdList></Reference><Reference><Citation>Kim, H. R. et al. A unified framework for dopamine signals across timescales. Cell 183, 1600&#x2013;1616 (2020).</Citation><ArticleIdList><ArticleId IdType="pubmed">33248024</ArticleId><ArticleId IdType="pmc">7736562</ArticleId></ArticleIdList></Reference><Reference><Citation>Ogasawara, T. et al. A primate temporal cortex &#x2014; zona incerta pathway for novelty seeking. Nat. Neurosci. 25, 50&#x2013;60 (2022).</Citation><ArticleIdList><ArticleId IdType="pubmed">34903880</ArticleId></ArticleIdList></Reference><Reference><Citation>Akam, T. &amp; Walton, M. E. What is dopamine doing in model-based reinforcement learning? Curr. Opin. Behav. Sci. 38, 74&#x2013;82 (2021).</Citation><ArticleIdList><ArticleId IdType="pubmed">37082448</ArticleId><ArticleId IdType="pmc">7614453</ArticleId></ArticleIdList></Reference><Reference><Citation>Bromberg-Martin, E. S., Matsumoto, M. &amp; Hikosaka, O. Dopamine in motivational control: rewarding, aversive, and alerting. Neuron 68, 815&#x2013;834 (2010).</Citation><ArticleIdList><ArticleId IdType="pubmed">21144997</ArticleId><ArticleId IdType="pmc">3032992</ArticleId></ArticleIdList></Reference><Reference><Citation>Pearce, J. M. &amp; Hall, G. A model for Pavlovian learning: variations in the effectiveness of conditioned but not of unconditioned stimuli. Psychol. Rev. 87, 532&#x2013;552 (1980).</Citation><ArticleIdList><ArticleId IdType="pubmed">7443916</ArticleId></ArticleIdList></Reference><Reference><Citation>Pearce et al. in Quantitative Analyses of Behavior Vol. 3 (eds Commons, M. L., Herrnstein, R. J. &amp; Wagner, A. R.) 241&#x2013;255 (Ballinger, 1982).</Citation></Reference><Reference><Citation>Stalnaker, T. A. et al. Dopamine neuron ensembles signal the content of sensory prediction errors. eLife 8, e49315 (2019).</Citation><ArticleIdList><ArticleId IdType="pubmed">31674910</ArticleId><ArticleId IdType="pmc">6839916</ArticleId></ArticleIdList></Reference><Reference><Citation>Howard, J. D., Edmonds, D., Schoenbaum, G. &amp; Kahnt, T. Distributed midbrain responses signal the content of positive identity prediction errors. Curr. Biol. 34, 241&#x2013;4240.e4 (2024).</Citation></Reference><Reference><Citation>Garr, E. et al. Mesostriatal dopamine is sensitive to specific cue-reward contingencies. Sci. Adv. https://doi.org/10.1126/sciadv.adn4203 (2023).</Citation></Reference><Reference><Citation>Steinberg, E. E. et al. A causal link between prediction errors, dopamine neurons and learning. Nat. Neurosci. 16, 966&#x2013;973 (2013).</Citation><ArticleIdList><ArticleId IdType="pubmed">23708143</ArticleId><ArticleId IdType="pmc">3705924</ArticleId></ArticleIdList></Reference><Reference><Citation>Kamin, L. J. "Attention-like" processes in classical conditioning. In Miami Symposium on the Prediction of Behavior, 1967: Aversive Stimulation (ed. Jones, M. R.) 9&#x2013;31 (Univ. of Miami Press, 1968).</Citation></Reference><Reference><Citation>Keiflin, R., Pribut, H. J., Shah, N. B. &amp; Janak, P. H. Ventral tegmental dopamine neurons participate in reward identity predictions. Curr. Biol. 29, 93&#x2013;103.e3 (2019).</Citation><ArticleIdList><ArticleId IdType="pubmed">30581025</ArticleId></ArticleIdList></Reference><Reference><Citation>Holland, P. C. &amp; Rescorla, R. A. The effects of two ways of devaluing the unconditioned stimulus after first and second-order appetitive conditioning. J. Exp. Psychol. Anim. Behav. Process. 1, 355&#x2013;363 (1975).</Citation></Reference><Reference><Citation>Howard, J. D., Gottfried, J. A., Tobler, P. N. &amp; Kahnt, T. Identity-specific coding of future rewards in the human orbitofrontal cortex. Proc. Natl Acad. Sci. USA 112, 5195&#x2013;5200 (2015).</Citation><ArticleIdList><ArticleId IdType="pubmed">25848032</ArticleId><ArticleId IdType="pmc">4413264</ArticleId></ArticleIdList></Reference><Reference><Citation>Stalnaker, T. A. et al. Orbitofrontal neurons infer the value and identity of predicted outcomes. Nat. Commun. 5, 3926 (2014).</Citation><ArticleIdList><ArticleId IdType="pubmed">24894805</ArticleId></ArticleIdList></Reference><Reference><Citation>Stoll, F. M. &amp; Rudebeck, P. H. Preferences reveal dissociable encoding across prefrontal-limbic circuits. Neuron 112, 2241&#x2013;2256.e8 (2024).</Citation><ArticleIdList><ArticleId IdType="pubmed">38640933</ArticleId></ArticleIdList></Reference><Reference><Citation>Burke, K. A., Franz, T. M., Miller, D. N. &amp; Schoenbaum, G. The role of the orbitofrontal cortex in the pursuit of happiness and more specific rewards. Nature 454, 340&#x2013;344 (2008).</Citation><ArticleIdList><ArticleId IdType="pubmed">18563088</ArticleId><ArticleId IdType="pmc">2727745</ArticleId></ArticleIdList></Reference><Reference><Citation>Howard, J. D. et al. Targeted stimulation of human orbitofrontal networks disrupts outcome-guided behavior. Curr. Biol. 30, 490&#x2013;498.e4 (2020).</Citation><ArticleIdList><ArticleId IdType="pubmed">31956033</ArticleId><ArticleId IdType="pmc">7291849</ArticleId></ArticleIdList></Reference><Reference><Citation>Rudebeck, P. H., Saunders, R. C., Prescott, A. T., Chau, L. S. &amp; Murray, E. A. Prefrontal mechanisms of behavioral flexibility, emotion regulation and value updating. Nat. Neurosci. 16, 1140&#x2013;1145 (2013).</Citation><ArticleIdList><ArticleId IdType="pubmed">23792944</ArticleId><ArticleId IdType="pmc">3733248</ArticleId></ArticleIdList></Reference><Reference><Citation>Sias, A. C. et al. A bidirectional corticoamygdala circuit for the encoding and retrieval of detailed reward memories. eLife 10, e68617 (2021).</Citation><ArticleIdList><ArticleId IdType="pubmed">34142660</ArticleId><ArticleId IdType="pmc">8266390</ArticleId></ArticleIdList></Reference><Reference><Citation>Ostlund, S. B. &amp; Balleine, B. W. Orbitofrontal cortex mediates outcome encoding in Pavlovian but not instrumental learning. J. Neurosci. 27, 4819&#x2013;4825 (2007).</Citation><ArticleIdList><ArticleId IdType="pubmed">17475789</ArticleId><ArticleId IdType="pmc">6672090</ArticleId></ArticleIdList></Reference><Reference><Citation>McDannald, M. A., Saddoris, M. P., Gallagher, M. &amp; Holland, P. C. Lesions of orbitofrontal cortex impair rats&#x2019; differential outcome expectancy learning but not conditioned stimulus-potentiated feeding. J. Neurosci. 25, 4626&#x2013;4632 (2005).</Citation><ArticleIdList><ArticleId IdType="pubmed">15872110</ArticleId><ArticleId IdType="pmc">1201522</ArticleId></ArticleIdList></Reference><Reference><Citation>Sias, A. C. et al. Dopamine projections to the basolateral amygdala drive the encoding of identity-specific reward memories. Nat. Neurosci. 27, 728&#x2013;736 (2024).</Citation><ArticleIdList><ArticleId IdType="pubmed">38396258</ArticleId><ArticleId IdType="pmc">11110430</ArticleId></ArticleIdList></Reference><Reference><Citation>Brogden, W. J. Sensory pre-conditioning. J. Exp. Psychol. 25, 323&#x2013;332 (1939).</Citation></Reference><Reference><Citation>Jones, J. L. et al. Orbitofrontal cortex supports behavior and learning using inferred but not cached values. Science 338, 953&#x2013;956 (2012).</Citation><ArticleIdList><ArticleId IdType="pubmed">23162000</ArticleId><ArticleId IdType="pmc">3592380</ArticleId></ArticleIdList></Reference><Reference><Citation>Wang, F., Schoenbaum, G. &amp; Kahnt, T. Interactions between human orbitofrontal cortex and hippocampus support model-based inference. PLoS Biol. 18, e3000578 (2020).</Citation><ArticleIdList><ArticleId IdType="pubmed">31961854</ArticleId><ArticleId IdType="pmc">6974031</ArticleId></ArticleIdList></Reference><Reference><Citation>Sharpe, M. J. et al. Dopamine transients are sufficient and necessary for acquisition of model-based associations. Nat. Neurosci. 20, 735&#x2013;742 (2017).</Citation><ArticleIdList><ArticleId IdType="pubmed">28368385</ArticleId><ArticleId IdType="pmc">5413864</ArticleId></ArticleIdList></Reference><Reference><Citation>Esmoris-Arranz, F. J., Miller, R. R. &amp; Matute, H. Blocking of subsequent and antecedent events. J. Exp. Psychol. Anim. Behav. Process. 23, 145&#x2013;156 (1997).</Citation><ArticleIdList><ArticleId IdType="pubmed">9095539</ArticleId></ArticleIdList></Reference><Reference><Citation>Kamin, L. J. in Punishment and Aversive Behavior (eds Campbell, B. A. &amp; Church, R. M.) 242&#x2013;259 (Appleton-Century-Crofts, 1969).</Citation></Reference><Reference><Citation>Mackintosh, N. J. A theory of attention: variations in the associability of stimuli with reinforcement. Psychol. Rev. 82, 276&#x2013;298 (1975).</Citation></Reference><Reference><Citation>Hart, E. E., Sharpe, M. J., Gardner, M. P. &amp; Schoenbaum, G. Responding to preconditioned cues is devaluation sensitive and requires orbitofrontal cortex during cue-cue learning. eLife 9, e59998 (2020).</Citation><ArticleIdList><ArticleId IdType="pubmed">32831173</ArticleId><ArticleId IdType="pmc">7481003</ArticleId></ArticleIdList></Reference><Reference><Citation>Sharpe, M. J., Batchelor, H. M. &amp; Schoenbaum, G. Preconditioned cues have no value. eLife 6, e28362 (2017).</Citation><ArticleIdList><ArticleId IdType="pubmed">28925358</ArticleId><ArticleId IdType="pmc">5619948</ArticleId></ArticleIdList></Reference><Reference><Citation>Wong, F. S., Westbrook, R. F. &amp; Holmes, N. M. &#x2018;Online&#x2019; integration of sensory and fear memories in the rat medial temporal lobe. eLife 8, e47085 (2019).</Citation><ArticleIdList><ArticleId IdType="pubmed">31180324</ArticleId><ArticleId IdType="pmc">6592679</ArticleId></ArticleIdList></Reference><Reference><Citation>Costa, K. M., Raheja, N., Mirani, J., Sercander, C. &amp; Schoenbaum, G. Striatal dopamine release reflects a domain-general prediction error. Preprint at bioRxiv https://doi.org/10.1101/2023.08.19.553959 (2023).</Citation></Reference><Reference><Citation>Moser, E. I., Kropff, E. &amp; Moser, M. B. Place cells, grid cells, and the brain&#x2019;s spatial representation system. Annu. Rev. Neurosci. 31, 69&#x2013;89 (2008).</Citation><ArticleIdList><ArticleId IdType="pubmed">18284371</ArticleId></ArticleIdList></Reference><Reference><Citation>Witten, I. B. et al. Recombinase-driver rat lines: tools, techniques, and optogenetic application to dopamine-mediated reinforcement. Neuron 72, 721&#x2013;733 (2011).</Citation><ArticleIdList><ArticleId IdType="pubmed">22153370</ArticleId><ArticleId IdType="pmc">3282061</ArticleId></ArticleIdList></Reference><Reference><Citation>Ilango, S. et al. Similar roles of substantia nigra and ventral tegmental dopamine neurons in reward and aversion. J. Neurosci. 34, 817&#x2013;822 (2014).</Citation><ArticleIdList><ArticleId IdType="pubmed">24431440</ArticleId><ArticleId IdType="pmc">3891961</ArticleId></ArticleIdList></Reference><Reference><Citation>Covey, D. P. &amp; Cheer, J. F. Accumbal dopamine release tracks the expectation of dopamine neuron-mediated reinforcement. Cell Rep. 27, 481&#x2013;490 (2019).</Citation><ArticleIdList><ArticleId IdType="pubmed">30970251</ArticleId><ArticleId IdType="pmc">6481661</ArticleId></ArticleIdList></Reference><Reference><Citation>Wolff, A. R. &amp; Saunders, B. T. Sensory cues potentiate VTA dopamine mediated reinforcement. eNeuro 11, ENEURO.0421-0423.2024 (2024).</Citation></Reference><Reference><Citation>Chang, C. Y. et al. Brief optogenetic inhibition of VTA dopamine neurons mimics the effects of endogenous negative prediction errors during Pavlovian over-expectation. Nat. Neurosci. 19, 111&#x2013;116 (2016).</Citation><ArticleIdList><ArticleId IdType="pubmed">26642092</ArticleId></ArticleIdList></Reference><Reference><Citation>Chang, C. Y., Gardner, M., Di Tillio, M. G. &amp; Schoenbaum, G. Optogenetic blockade of dopamine transients prevents learning induced by changes in reward features. Curr. Biol. 27, 3480&#x2013;3486 (2017).</Citation><ArticleIdList><ArticleId IdType="pubmed">29103933</ArticleId><ArticleId IdType="pmc">5698141</ArticleId></ArticleIdList></Reference><Reference><Citation>Chang, C. Y., Gardner, M. P. H., Conroy, J. S., Whitaker, L. R. &amp; Schoenbaum, G. Brief, but not prolonged, pauses in the firing of midbrain dopamine neurons are sufficient to produce a conditioned inhibitor. J. Neurosci. 38, 8822&#x2013;8830 (2018).</Citation><ArticleIdList><ArticleId IdType="pubmed">30181136</ArticleId><ArticleId IdType="pmc">6181314</ArticleId></ArticleIdList></Reference><Reference><Citation>Millard, S. J. et al. Cognitive representations of intracranial self-stimulation of midbrain dopamine neurons depend on stimulation frequency. Nat. Neurosci. 27, 1253&#x2013;1259 (2024).</Citation><ArticleIdList><ArticleId IdType="pubmed">38741021</ArticleId><ArticleId IdType="pmc">11239488</ArticleId></ArticleIdList></Reference><Reference><Citation>Takahashi, Y. K. et al. Dopaminergic prediction errors in the ventral tegmental area reflect a multithreaded predictive model. Nat. Neurosci. 26, 830&#x2013;839 (2023).</Citation><ArticleIdList><ArticleId IdType="pubmed">37081296</ArticleId><ArticleId IdType="pmc">10646487</ArticleId></ArticleIdList></Reference><Reference><Citation>Gardner, M. P. H., Schoenbaum, G. &amp; Gershman, S. J. Rethinking dopamine as generalized prediction error. Proc. Biol. Sci. 285, 20181645 (2018).</Citation><ArticleIdList><ArticleId IdType="pubmed">30464063</ArticleId><ArticleId IdType="pmc">6253385</ArticleId></ArticleIdList></Reference><Reference><Citation>Gershman, S. J. The successor representation: its computational logic and neural substrates. J. Neurosci. 38, 7193&#x2013;7200 (2018).</Citation><ArticleIdList><ArticleId IdType="pubmed">30006364</ArticleId><ArticleId IdType="pmc">6096039</ArticleId></ArticleIdList></Reference><Reference><Citation>Langdon, A. J., Sharpe, M. J., Schoenbaum, G. &amp; Niv, Y. Model-based predictions for dopamine. Curr. Opin. Neurobiol. 49, 1&#x2013;7 (2018).</Citation><ArticleIdList><ArticleId IdType="pubmed">29096115</ArticleId></ArticleIdList></Reference><Reference><Citation>German, D. C., Schlusselberg, D. S. &amp; Woodward, D. J. Three-dimensional computer reconstruction of midbrain dopaminergic neuronal populations: from mouse to man. J. Neural Transm. 57, 243&#x2013;254 (1983).</Citation><ArticleIdList><ArticleId IdType="pubmed">6140298</ArticleId></ArticleIdList></Reference><Reference><Citation>Kahnt, T. in Encyclopedia of the Human Brain 2nd edn (ed. Grafman, J. H.) 387&#x2013;400 (Elsevier, 2025).</Citation></Reference><Reference><Citation>Tegelbeckers, J., Porter, D. B., Voss, J. L., Schoenbaum, G. &amp; Kahnt, T. Lateral orbitofrontal cortex integrates predictive information across multiple cues to guide behavior. Curr. Biol. 33, 4496&#x2013;4504.e5 (2023).</Citation><ArticleIdList><ArticleId IdType="pubmed">37804827</ArticleId><ArticleId IdType="pmc">10622115</ArticleId></ArticleIdList></Reference></ReferenceList></PubmedData></PubmedArticle></PubmedArticleSet>