EEG correlates of physical effort and reward processing during reinforcement learning.

Effort-based decision making is often modeled using subjective value, a function of reward discounted by effort. We asked whether EEG event-related potential (ERP) correlates of reward processing are also modulated by physical effort. Human participants performed a task in which they were required to accurately produce target levels of muscle activation to receive rewards. Quadriceps muscle activation was recorded with electromyography (EMG) during isometric knee extension. On a given trial, the target muscle activation required either low or high effort. The effort was determined probabilistically according to a binary choice, such that the responses were associated with 20% and 80% probability of high effort. This contingency could only be learned through experience, and it reversed periodically. Binary reinforcement feedback depended on accurately producing the target muscle activity. Participants adaptively avoided effort by switching responses more frequently after choices that resulted in hard effort. Feedback after participants' choices that revealed the resulting effort requirement did not elicit modulation of the feedback-related negativity/reward positivity (FRN/RP). However, the neural response to reinforcement outcome after effort production was increased by preceding physical effort. Source decomposition revealed separable early and late positive deflections contributing to the ERP. The main effect of reward outcome, characteristic of the FRN/RP, loaded onto the earlier component, whereas the reward × effort interaction was observed only in the later positivity, which resembled the P300. Thus, retrospective effort modulates reward processing. This may explain paradoxical behavioral findings whereby rewards requiring more effort to obtain can become more powerful reinforcers. Choices probabilistically determined the physical effort requirements for a subsequent task, and reward depended on task performance. Feedback revealing whether choices resulted in easy or hard effort did not elicit reinforcement learning signals. However, the neural responses to reinforcement were modulated by preceding effort. Thus, effort itself was not treated as loss or punishment, but it affected the responses to subsequent reinforcement outcomes. This may explain how effort can enhance the motivational effect of reward.