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Annu Rev Neurosci. 2017 Jul 25;40:373-394. doi: 10.1146/annurev-neuro-072116-031109. Epub 2017 Apr 24.

Neural Circuitry of Reward Prediction Error.

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Department of Molecular and Cellular Biology, Center for Brain Science, Harvard University, Cambridge, Massachusetts 02138; email: ,
Department of Psychiatry and Behavioral Sciences, Stanford University School of Medicine, Stanford, California 94305; email:


Dopamine neurons facilitate learning by calculating reward prediction error, or the difference between expected and actual reward. Despite two decades of research, it remains unclear how dopamine neurons make this calculation. Here we review studies that tackle this problem from a diverse set of approaches, from anatomy to electrophysiology to computational modeling and behavior. Several patterns emerge from this synthesis: that dopamine neurons themselves calculate reward prediction error, rather than inherit it passively from upstream regions; that they combine multiple separate and redundant inputs, which are themselves interconnected in a dense recurrent network; and that despite the complexity of inputs, the output from dopamine neurons is remarkably homogeneous and robust. The more we study this simple arithmetic computation, the knottier it appears to be, suggesting a daunting (but stimulating) path ahead for neuroscience more generally.


arithmetic; circuitry; dopamine; learning; prediction error; reward

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