Anatomical and pharmacological evidence suggests that the dorsal raphe serotonin system and the ventral tegmental and substantia nigra dopamine system may act as mutual opponents. In the light of the temporal difference model of the involvement of the dopamine system in reward learning, we consider three aspects of motivational opponency involving dopamine and serotonin. We suggest that a tonic serotonergic signal reports the long-run average reward rate as part of an average-case reinforcement learning model; that a tonic dopaminergic signal reports the long-run average punishment rate in a similar context; and finally speculate that a phasic serotonin signal might report an ongoing prediction error for future punishment.
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