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Vol. 11, Issue 8, 2705-2717, August 2000
and§
This report examines the distribution of an RNA polymerase I
transcription factor (upstream binding factor; UBF), pre-rRNA processing factors (nucleolin and fibrillarin), and pre-rRNAs throughout mitosis and postmitotic nucleologenesis in HeLa cells. The
results demonstrate that nucleolin, fibrillarin, and pre-rRNAs synthesized at G2/M phase of the previous cell cycle are directly recruited to UBF-associated nucleolar organizer regions (NORs) early in telophase before chromosome decondensation. Unlike the fusion
of prenucleolar bodies to the nucleoli, this early recruitment of
processing factors and pre-rRNAs is independent of RNA polymerase I
transcription. In the absence of polymerase I transcription, the
initial localization of nucleolin, fibrillarin, and pre-rRNAs to
UBF-associated NORs generates segregated mininucleoli that are similar
to the larger ones observed in interphase cells grown under the same
conditions. Pre-rRNAs are juxtaposed to UBF-nucleolin-fibrillarin caps
that may represent the segregated nucleoli observed by electron microscopy. These findings lead to a revised model of nucleologenesis. We propose that nucleolar formation at the end of mitosis results from
direct recruitment of processing factors and pre-rRNAs to UBF-associated NORs before or at the onset of rDNA transcription. This
is followed by fusion of prepackaged prenucleolar bodies into the
nucleolus. Pre-ribosomal ribonucleoproteins synthesized in the
previous cell cycle may contribute to postmitotic nucleologenesis.
Institut Jacques Monod, 75251 Paris, France; and
Department of Cell and Molecular Biology, Northwestern
University Medical School, Chicago, Illinois 60611
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