Nonlinear integration of sensory responses in the rat barrel cortex: an intracellular study in vivo

To study integration of converging sensory inputs on single cortical neurons, we performed intracellular recordings in vivo in the barrel cortex of the barbiturate-anesthetized rat. We deflected the principal whisker (PW) for each cell either alone or preceded (at 20, 50, and 100 msec) by the deflection of a small number of remote whiskers (RWs) far from the PW. The synaptic responses to both the PW and the RW were similar qualitatively and consisted of excitation followed by inhibition that comprised an early and a late component. The RW response was of smaller amplitude and more often subthreshold for action potential generation. The main effect of the RW deflection was a suppression of the subsequent response to the PW that was most pronounced at the 20 msec interval and decreased progressively at the 50 and 100 msec intervals. Suppression of the spike output of the cell was not caused by hyperpolarization (subtractive inhibition) but by a reduction in the EPSP amplitude (divisive inhibition), resulting in a highly sublinear summation of the two responses. The small decrease in input resistance caused by the RW responses is not consistent with synaptic shunting as the main cause of the reduction of the EPSP amplitude. Instead, our results suggest that suppression results from a decrease in the amount of synaptic input triggered by the PW, particularly the early excitation. We suggest that this process involves a reduction in reverberant granular cell excitation that is induced by PW deflection.