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Almost nothing is known about dendritic activity in awake animals and even less about its relationship to behavior. The tuft dendrites of layer 5 (L5) pyramidal neurons lie in layer 1, where long-range axons from secondary thalamic nuclei and higher cortical areas arrive. This class of input is very dependent on active thalamo-cortical loops and activity in higher brain areas and so is likely to be heavily influenced by the conscious state of the animal. If, as has been suggested, the dendrites of pyramidal neurons actively participate in this process, dendritic activity should greatly increase in the awake state. Here, we measured calcium activity in L5 pyramidal neuron dendrites using the "periscope" fiberoptic system. Recordings were made in the sensorimotor cortex of awake and anesthetized rats following sensory stimulation of the hindlimb. Bi-phasic dendritic responses evoked by hindlimb stimulation were extremely dependent on brain state. In the awake state, there was a prominent slow, delayed response whose integral was on average 14-fold larger than in the anesthetized state. Moreover, the dramatic increases in dendritic activity closely correlated to the strength of subsequent hindlimb movement. These changes were confined to L5 pyramidal dendrites and were not reflected in the response of layer 2/3 (L2/3) neurons to air-puff stimuli in general (which actually decreased in the awake state). The results demonstrate that the activity of L5 pyramidal dendrites is a neural correlate of awake behavior.

Enhanced dendritic activity in awake rats