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Vol. 13, Issue 7, 2207-2222, July 2002
Department of Molecular and Cell Biology, University of California,
Berkeley, Berkeley, California 94720
In Saccharomyces cerevisiae, silencing at the
HM loci depends on Sir proteins, which are structural
components of silenced chromatin. To explore the structure and assembly
of silenced chromatin, the associations of Sir proteins with sequences
across the HMR locus were examined by chromatin
immunoprecipitation. In wild-type cells, Sir2p, Sir3p, and Sir4p were
spread throughout and coincident with the silenced region at
HMR. Sir1p, in contrast, associated only with the
HMR-E silencer, consistent with its role in
establishment but not maintenance of silencing. Sir4p was required for
the association of other Sir proteins with silencers. In contrast, in
the absence of Sir2p or Sir3p, partial assemblies of Sir proteins could
form at silencers, where Sir protein assembly began. Spreading across HMR required Sir2p and Sir3p, as well as the deacetylase
activity of Sir2p. These data support a model for the spreading of
silenced chromatin involving cycles of nucleosome deacetylation by
Sir2p followed by recruitment of additional Sir2p, Sir3p, and Sir4p to
the newly deacetylated nucleosome. This model suggests mechanisms for
boundary formation, and for maintenance and inheritance of silenced
chromatin. The principles are generalizable to other types of heritable
chromatin states.
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