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Vol. 10, Issue 12, 4385-4402, December 1999
Department of Embryology, Carnegie Institution, Baltimore, Maryland
21210
We have examined the distribution of RNA transcription and
processing factors in the amphibian oocyte nucleus or germinal vesicle.
RNA polymerase I (pol I), pol II, and pol III occur in the Cajal
bodies (coiled bodies) along with various components required
for transcription and processing of the three classes of nuclear
transcripts: mRNA, rRNA, and pol III transcripts. Among these
components are transcription factor IIF (TFIIF), TFIIS, splicing
factors, the U7 small nuclear ribonucleoprotein particle, the
stem-loop binding protein, SR proteins, cleavage and polyadenylation factors, small nucleolar RNAs, nucleolar proteins that are probably involved in pre-rRNA processing, and TFIIIA. Earlier studies and data
presented here show that several of these components are first targeted
to Cajal bodies when injected into the oocyte and only subsequently
appear in the chromosomes or nucleoli, where transcription itself
occurs. We suggest that pol I, pol II, and pol III transcription and
processing components are preassembled in Cajal bodies before transport
to the chromosomes and nucleoli. Most components of the pol II
transcription and processing pathway that occur in Cajal bodies are
also found in the many hundreds of B-snurposomes in the germinal
vesicle. Electron microscopic images show that B-snurposomes consist
primarily, if not exclusively, of 20- to 30-nm particles, which closely
resemble the interchromatin granules described from sections of somatic
nuclei. We suggest the name pol II transcriptosome for
these particles to emphasize their content of factors involved in
synthesis and processing of mRNA transcripts. We present a model in
which pol I, pol II, and pol III transcriptosomes are assembled in the
Cajal bodies before export to the nucleolus (pol I), to the
B-snurposomes and eventually to the chromosomes (pol II), and directly
to the chromosomes (pol III). The key feature of this model is the
preassembly of the transcription and processing machinery into unitary
particles. An analogy can be made between ribosomes and
transcriptosomes, ribosomes being unitary particles involved in
translation and transcriptosomes being unitary particles for
transcription and processing of RNA.
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