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Neuromodulatory systems code for the reward-prediction errors, and feedback connections from the response-selection stage mediate top-down attention effects The combined influence of feedback connections and neuromodulatory substances on plasticity enables powerful learning rules for the training of 'deep', multilayered neuronal networks Feedback connections project to cortical layers that are distinct from feedforward input, where they impinge on distal dendritic segments, separate excitatory neuronal populations or inhibitory interneurons Feedback connections gate plasticity in cortical pyramidal neurons by promoting NMDA-receptor-driven calcium entry into dendrites and by disinhibiting the cortical column through activation of vasoactive-intestinal-peptide-positive interneurons (among others) Synaptic tags are biochemical processes that make synapses eligible for plasticity

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