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Topographic reorganization of the hand representation in cortical area 3b owl monkeys trained in a frequency-discrimination task. Cerebellar Purkinje cell activity related to the classically conditioned nictitating membrane response. While outcome-mediated feedback likely still provides basic boundaries for the behavior to ensure that it is shaped based on outcome, the existing motor-related ensemble at this stage of learning likely operates mostly within these bounds. Basal ganglia and cerebellar circuits provide major inputs to the motor cortex through the thalamus, so the feedback-based learning in these circuits likely assist in selecting effective circuits in the motor cortex from the broadened population initially explored. Rule learning enhances structural plasticity of long-range axons in frontal cortex. When this tool was activated in the motor cortex following training, impairment in motor skill learning was observed.