One critic, two actors:Evidence for covert learning in the basal ganglia

This paper introduces a new hypothesis concerning the dissociated role of the basal ganglia in the selection and the evaluation of action that has been formulated using a theoretical model and confirmed experimentally in monkeys. To do so, prior to learning, we inactivated the internal part of the Globus Pallidus (GPi, the main output structure of the BG) with injections of muscimol and we tested monkeys on a variant of a two-armed bandit task where two stimuli are associated with two distinct reward probabilities (0.25 and 0.75 respectively). Unsurprisingly, performance in such condition are at the chance level because the output of basal ganglia is suppressed and they cannot influence behaviour. However, the theoretical model predicts that in the meantime, values of the stimuli are nonetheless covertly evaluated and learned. This has been tested and confirmed on the next day, when muscimol has been replaced by a saline solution: monkeys instantly showed significantly improved performances (above chance level), hence demonstrating they have covertly learned the relative value of the two stimuli. This tends to suggest a competition takes place in the Cortex-BG loop between two actors, one of whom being sensitive to criticism and the other not. Ultimately, the actual choice is valuated, independently of the origin of the decision.