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rRNA Synthesis and Processing
The genes coding for rRNA (except 5S rRNA) are
located in the nucleolar part of the nucleus. The
rRNA genes are highly repetitious and mammalian cells
contain 100 to 2000 copies of the rRNA genes per
cell. The genes are organised in transcription units
separated by non-transcribed spacers. Each
transcription unit contains sequences coding for 18S,
5.8S and 28S rRNA.
The transcription units are transcribed by RNA
polymerase I into giant RNA molecules, primary
transcripts, that in addition to the sequences
corresponding to 18S, 5.8S and 28S rRNA contains
external and internal transcribed spacer sequences.
The rate of nucleolar transcription is very high and
many polymerases operate on the same transcription
unit. The transciptionally active DNA therefore has a
Christmas tree-like appearance on electron
microscopic pictures.
The primary transcript is
processed into the mature 18S, 5.8S and 28S rRNAs.
The processing involves exo- and endo-nucleolytic
cleavages guided by snoRNA (small nucleolar RNAs) in
complex with proteins. The mature rRNAs contain
modified nucleotides which are added after
transcription by a snoRNA-dependent
mechanism.
5S ribosomal RNA is transcribed by RNA polymerase III
in the nucleoplasm. Each eukaryotic cell contains a
high number of copies of the 5S coding gene (up to 20
000 copies per cell). 5S rRNA contains overlapping
binding sites for two different proteins, ribosomal
protein L5 and transcription factor TFIIIA. The
mutual exclusive binding of these two proteins to 5S
rRNA is important for coordinating the expression of
5S rRNA to the production of the other
rRNAs.
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