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Received August 8, 2003
Revised September 15, 2003
Accepted after revision January 7, 2004
1 Max-Planck Institut für medizinische Forschung
* To whom correspondence should be addressed. E-mail: ian.manns{at}mail.mcgill.ca.
Layer 5 pyramidal neurons constitute a major sub- and intracortical output of the somatosensory cortex. This layer 5 is segregated into layers 5A and 5B which receive and distribute relatively independent afferent and efferent pathways. We performed in vivo whole-cell recordings from layer 5 neurons of the somatosensory (barrel) cortex of urethane anesthetized rats (P27-31). By delivering 6° single whisker deflections, whisker pad receptive fields were mapped for 16 layer 5A and 11 layer 5B neurons located below the layer 4 whisker-barrels. Average resting membrane potentials were -75.6 ± 1.1 mV, and spontaneous action potential (AP) rates were 0.54 ± 0.14 APs per second. Principal whisker (PW) evoked responses were similar in layers 5A and 5B neurons with an average 5.0 ± 0.6 mV post-synaptic potential (PSP) and 0.12 ± 0.03 APs per stimulus. The layer 5A sub- and suprathreshold receptive fields (RFs) were more confined to the principle whisker than those of layer 5B. The basal dendritic arbors of layer 5A and 5B cells were located below both layer 4 barrels and septa, but biased towards the barrel walls. Responses in both layers 5A and 5B developed slowly with onset latencies of 10.1 ± 0.5 ms and peak latencies of 33.9 ± 3.3 ms. Contralateral multi-whisker stimulation evoked PSPs similar in amplitude to those of PW deflections; whereas, ipsilateral stimulation evoked smaller and longer latency PSPs. We conclude that in L5 a whisker deflection is represented 2-fold: focally by L5A pyramids and more diffusely by L5B pyramids as a result of combining different inputs from lemniscal and paralemniscal pathways. The relevant output may be by the ensemble activity of the anatomically defined cortical modules associated with a single or a few barrel-columns.
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