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The Journal of Neuroscience, July 15, 2000, 20(14):5300-5311
Columnar Organization of Dendrites and Axons of Single and
Synaptically Coupled Excitatory Spiny Neurons in Layer 4 of the Rat
Barrel Cortex
Joachim
Lübke1,
Veronica
Egger2,
Bert
Sakmann2, and
Dirk
Feldmeyer2
1 Anatomisches Institut I,
Albert-Ludwigs-Universität Freiburg, D-79104 Freiburg, Germany,
and 2 Max-Planck-Institut für Medizinische Forschung,
Abteilung Zellphysiologie, D-69120 Heidelberg, Germany
Cortical columns are the functional units of the neocortex that are
particularly prominent in the "barrel" field of the somatosensory cortex. Here we describe the morphology of two classes of synaptically coupled excitatory neurons in layer 4 of the barrel cortex, spiny stellate, and star pyramidal cells, respectively. Within a single barrel, their somata tend to be organized in clusters. The dendritic arbors are largely confined to layer 4, except for the distal part of
the apical dendrite of star pyramidal neurons that extends into layer
2/3. In contrast, the axon of both types of neurons spans the cortex
from layer 1 to layer 6. The most prominent axonal projections are
those to layers 4 and 2/3 where they are largely restricted to a single
cortical column. In layers 5 and 6, a small fraction of axon
collaterals projects also across cortical columns. Consistent with the
dense axonal projection to layers 4 and 2/3, the total number and
density of boutons per unit axonal length was also highest there.
Electron microscopy combined with GABA postimmunogold labeling revealed
that most (>90%) of the synaptic contacts were established on
dendritic spines and shafts of excitatory neurons in layers 4 and
2/3.
The largely columnar organization of dendrites and axons of both cell
types, combined with the preferential and dense projections within
cortical layers 4 and 2/3, suggests that spiny stellate and star
pyramidal neurons of layer 4 serve to amplify thalamic input and relay
excitation vertically within a single cortical column.
Key words:
barrel cortex; layer 4; spiny stellate cell; star
pyramidal neuron; cortical column; axonal projection; paired
recordings; intracellular labeling
Copyright © 2000 Society for Neuroscience 0270-6474/00/20145300-12$05.00/0
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