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The mammalian brain can form associations between behaviorally relevant stimuli in an animal's environment. While such learning is thought to primarily involve high-order association cortex, even primary sensory areas receive long-range connections carrying information that could contribute to high-level representations. Here, we imaged layer 1 apical dendrites in the barrel cortex of mice performing a whisker-based operant behavior. In addition to sensory-motor events, calcium signals in apical dendrites of layers 2/3 and 5 neurons and in layer 2/3 somata track the delivery of rewards, both choice related and randomly administered. Reward-related tuft-wide dendritic spikes emerge gradually with training and are task specific. Learning recruits cells whose intrinsic activity coincides with the time of reinforcement. Layer 4 largely lacked reward-related signals, suggesting a source other than the primary thalamus. Our results demonstrate that a sensory cortex can acquire a set of associations outside its immediate sensory modality and linked to salient behavioral events.

Original publication




Journal article


Cell Rep

Publication Date





2000 - 2008.e2


GCaMP, apical dendrites, barrel cortex, detection, reward, two-photon, vibrissa, voltage-gated calcium, Animals, Calcium Signaling, Dendrites, Female, Male, Mice, Mice, Inbred C57BL, Reinforcement, Psychology, Sensory Receptor Cells, Somatosensory Cortex, Vibrissae