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The Journal of Neuroscience, June 15, 2005, 25(24):5670-5679; doi:10.1523/JNEUROSCI.1173-05.2005

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Behavioral/Systems/Cognitive
Laminar and Columnar Organization of Ascending Excitatory Projections to Layer 2/3 Pyramidal Neurons in Rat Barrel Cortex

Gordon M. G. Shepherd and Karel Svoboda

Howard Hughes Medical Institute, Cold Spring Harbor Laboratory, Cold Spring Harbor, New York 11724

Excitatory synaptic projections to layer 2/3 (L2/3) pyramidal neurons in brain slices from the rat barrel cortex were measured using quantitative laser-scanning photostimulation (LSPS) mapping. In the barrel cortex, cytoarchitectonic "barrels" and "septa" in L4 define a stereotypical array of landmarks, allowing alignment and averaging of LSPS maps from multiple cells in different slices. We distinguished inputs to L2 and L3 neurons above barrels and septa. Average input maps revealed that barrel-related ascending projections (L42/3^barrel) interdigitated with a novel septum-related projection (L5A2^septum). We also explored the functional organization of these projections by comparing the input maps of multiple cells in individual slices. L2/3 cells sharing the same barrel-related column showed strong correlations in their input maps, independent of their precise locations within the column; otherwise, correlations fell rapidly as a function of intersomatic separation. Our data indicate that barrel-related and septum-related columns are associated with distinct functional circuits. These projections are likely to mediate parallel processing of somatosensory signals within the barrel cortex, with L42/3^barrel and L5A2^septum representing the intracortical continuations of, respectively, the subcortical lemniscal and paralemniscal systems conveying somatosensory information to the barrel cortex.

Key words: barrel; cortex; septum; paralemniscal; somatosensory; microcircuit; photostimulation; glutamate uncaging; connectivity

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Received Jan 25, 2005; revised April 20, 2005; accepted April 27, 2005.

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