Synaptic responses to whisker deflections in rat barrel cortex as a function of cortical layer and stimulus intensity

To study the synaptic and spike responses of barrel cortex neurons as a function of cortical layer and stimulus intensity, we recorded intracellularly in vivo from barbiturate anesthetized rats while increasing the velocity-acceleration of the whisker deflection. Granular (Gr; layer 4) cells had the EPSP with the shortest peak and onset latency, whereas supragranular (SGr; layers 2-3) cells had the EPSP with longest duration and slowest rate of rise. Infragranular (Igr; layers 5-6) cells had intermediate values, and thus each layer was unique. The spike response peak of Gr cells was followed by IGr and then by SGr cells. In all cells, depolarization reduced the duration and amplitude of the response, but only in Gr cells did it reveal an early IPSP that cut short the EPSP. This early IPSP was associated with a large decrease in input resistance and an apparent reversal potential below spike threshold; consequently, synaptic integration in Gr cells was limited to the initial 5-7 msec of the response. In contrast, in SGr and IGr cells, results suggest an overlap in time of the EPSP and IPSP, with a small drop in input resistance and an apparent reversal potential above spike threshold, facilitating input integration for up to 20 msec. Decreasing stimulus intensity (velocity-acceleration) reduced the amplitude and increased the peak latency of the response without altering its synaptic composition. We propose that layer 4 circuits are better suited to perform coincidence detection, whereas supra and infragranular circuits are better designed for input integration.