A universal role of the ventral striatum in reward-based learning: evidence from human studies

Reinforcement learning enables organisms to adjust their behavior in order to maximize rewards. Electrophysiological recordings of dopaminergic midbrain neurons have shown that they code the difference between actual and predicted rewards, i.e., the reward prediction error, in many species. This error signal is conveyed to both the striatum and cortical areas and is thought to play a central role in learning to optimize behavior. However, in human daily life rewards are diverse and often only indirect feedback is available. Here we explore the range of rewards that are processed by the dopaminergic system in human participants, and examine whether it is also involved in learning in the absence of explicit rewards. While results from electrophysiological recordings in humans are sparse, evidence linking dopaminergic activity to the metabolic signal recorded from the midbrain and striatum with functional magnetic resonance imaging (fMRI) is available. Results from fMRI studies suggest that the human ventral striatum (VS) receives valuation information for a diverse set of rewarding stimuli. These range from simple primary reinforcers such as juice rewards over abstract social rewards to internally generated signals on perceived correctness, suggesting that the VS is involved in learning from trial-and-error irrespective of the specific nature of provided rewards. In addition, we summarize evidence that the VS can also be implicated when learning from observing others, and in tasks that go beyond simple stimulus-action-outcome learning, indicating that the reward system is also recruited in more complex learning tasks.