Behavioral and Neural Changes after Gains and Losses of Conditioned Reinforcers

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Abstract
Human behaviors can be more powerfully influenced by conditioned reinforcers, such as money, than by primary reinforcers. Moreover, people often change their behaviors to avoid monetary losses. However, the effect of removing conditioned reinforcers on choices has not been explored in animals, and the neural mechanisms mediating the behavioral effects of gains and losses are not well understood. To investigate the behavioral and neural effects of gaining and losing a conditioned reinforcer, we trained rhesus monkeys for a matching pennies task in which the positive and negative values of its payoff matrix were realized by the delivery and removal of a conditioned reinforcer. Consistent with the findings previously obtained with non-negative payoffs and primary rewards, the animal's choice behavior during this task was nearly optimal. Nevertheless, the gain and loss of a conditioned reinforcer significantly increased and decreased, respectively, the tendency for the animal to choose the same target in subsequent trials. We also found that the neurons in the dorsomedial frontal cortex, dorsal anterior cingulate cortex, and dorsolateral prefrontal cortex often changed their activity according to whether the animal earned or lost a conditioned reinforcer in the current or previous trial. Moreover, many neurons in the dorsomedial frontal cortex also signaled the gain or loss occurring as a result of choosing a particular action as well as changes in the animal's behaviors resulting from such gains or losses. Thus, primate medial frontal cortex might mediate the behavioral effects of conditioned reinforcers and their losses.