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Journal of Neuroscience, Vol 12, 1010-1023, Copyright © 1992 by Society for Neuroscience
Molecular cloning and development analysis of a new glutamate receptor subunit isoform in cerebellum
V Gallo, LM Upson, WP Hayes, L Vyklicky Jr, CA Winters and A Buonanno
Unit on Molecular Neurobiology, National Institute of Child Health and Human Development, National Institutes of Health, Bethesda, Maryland 20892.
The glutamate receptor gene GluR-4 is proposed to generate two spliced
isoforms (Sommer et al., 1990). Screening a rat cerebellar cDNA library, we
have now identified a third type of transcript derived from GluR-4 gene by
differential RNA processing. This transcript encodes a protein with a
"flop" module between transmembrane regions 3 and 4, but with a C-terminus
segment of 36 amino acids different from the previously described GluR-4
flip/flop cDNAs. This subunit was therefore designated as GluR-4c flop.
Transcripts synthesized in vitro from GluR- 4c cDNA form
kainate/AMPA-activated channels when expressed in Xenopus oocytes. The
current-voltage relationship for kainate-evoked responses in oocytes
injected with GluR-4c showed strong inward rectification. The different
transcripts derived from the GluR-4 gene were studied on Northern blots
hybridized with either a cDNA probe or oligonucleotides specific for the
GluR-4 flip/flop and C-terminal domains. Three transcripts of 6.2, 4.2, and
3.0 kilobases (kb) derived from the GluR-4 gene were identified on Northern
blots containing total RNA prepared from different brain regions, using a
cDNA probe or an oligonucleotide corresponding to the N-terminal region
common to all transcripts. These transcripts were much more abundant in the
cerebellum than in other brain areas, and their levels increased during
cerebellar development. The maximal increase was observed between postnatal
days 1 and 20, an age corresponding to the division and maturation of
granule neurons. The flip/flop and the C-terminal oligonucleotides
hybridized to the two higher molecular weight transcripts but did not
hybridize to the small RNA. Interestingly, using cerebellar cells that were
cultured for up to 12 d, we observed that the three transcripts are present
in granule neurons, but that astrocytes only express the 6.2 and the 4.2 kb
transcripts. The 3.0 kb transcript accumulates in cerebellar granule cells
during development in vitro. Furthermore, in situ hybridization
histochemistry revealed that the GluR-4c transcripts are preferentially
expressed in cerebellar granule cells and Bergmann glial cells, whereas the
expression of GluR-4 flip mRNAs is restricted to Bergmann glial cells.
Interestingly, we also show that granule cells already express GluR-4c in
the premigratory zone of the external granular layer, indicating that
intrinsic or highly localized cues induce GluR-4c expression before these
cells reach their final position.
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