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Journal of Neuroscience, Vol 10, 1134-1153, Copyright © 1990 by Society for Neuroscience
Emergence and refinement of clustered horizontal connections in cat striate cortex
EM Callaway and LC Katz
Laboratory of Neurobiology, Rockefeller University, New York, New York 10021.
Pyramidal cells in layer 2/3 of adult cat striate cortex have long,
intrinsic horizontal axon collaterals within both layer 2/3 and layer 5.
These collaterals form periodic "clusters" of finer axon branches that link
columns of similar orientation selectivity. We have investigated the
sequence of events and possible mechanisms underlying the development of
these clustered intrinsic horizontal connections using a combination of
neuronal tracers and intracellular staining. Small injections of
fluorescent latex microspheres made during the first postnatal week (at
P4-6), when examined in tangential sections, produced an even, unclustered
distribution of retrogradely labeled cells up to 2 mm from the injection
site. At P8, retrograde labeling extended over a larger area and clustering
was discernible, primarily among the most distant labeled cells. At both P6
and P8, labeling was similar in layers 2/3 and 5, indicating that the
transition from clustered to unclustered connections occurred
simultaneously for cells in superficial and deep laminae. By the end of the
second postnatal week (P12-15), retrogradely labeled cells were far more
clustered both within and beyond the extent of P6 label; the density of
labeled cells was high throughout the labeled region, but much higher
within clusters. The periodicity of these nascent clusters was similar to
that in the adult. Despite obvious clustering, the pattern of retrograde
label observed following injections at 2-3 weeks (P12-21) differed markedly
from the adult, in that the regions between clusters contained many labeled
cells. Over the next 3 weeks, the connections were refined, so that by the
sixth postnatal week (P36-38), regions between clusters contained very few
retrogradely labeled cells and the overall pattern of retrograde label was
indistinguishable from that in adults. Despite differences in postmigratory
ages of neurons from the superficial and deep laminae, clustering of
retrogradely labeled cells from these 2 populations was similar at all
ages. Experiments in which 2-3 weeks elapsed between the time microsphere
injections were made and animals were killed demonstrated that neither the
initial formation of crude clusters nor their refinement was due to cell
death. Instead, cluster refinement resulted from specific process
elimination. When a red microsphere injection at P15 was followed by a
green microsphere injection at exactly the same location on P29, the
earlier injection resulted in crude clustering, as expected. Virtually all
of the cells double-labeled by the later injection were within the densest
clusters of label from the early injection.(ABSTRACT TRUNCATED AT 400
WORDS)
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