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doi:10.1534/genetics.106.062133
A more recent version of this article appeared on February 1, 2007.
REGULAR RESEARCH PAPERS |
The SU(VAR)3-9/HP1 complex differentially regulates the compaction state and degree of underreplication of X chromosome pericentric heterochromatin in Drosophila melanogaster
Olga V. Demakova 1, Galina V. Pokholkova 1, Tatyana D. Kolesnikova 1, Sergey A. Demakov 1, Eugenia N. Andreyeva 1, Elena S. Belyaeva 1 and Igor F. Zhimulev 1*
1 Institute of Cytology and Genetics
* To whom correspondence should be addressed. E-mail: zhimulev{at}bionet.nsc.ru.
Submitted on June 18, 2006
Revised on August 7, 2006
Accepted on 14 November 2006
In polytene chromosomes of Drosophila melanogaster, regions of pericentric heterochromatin coalesce to form a compact chromocenter and are highly underreplicated. Focusing on study of X chromosome heterochromatin, we demonstrate that loss of either SU(VAR)3-9 HMTase activity or HP1 protein differentially affects the compaction of different pericentric regions. Using a set of inversions disrupting X chromosome heterochromatin at background of the Su(var)3-9 mutations, we show that distal heterochromatin (blocks h26 to h29) is the only one within the chromocenter to form a big "puff"-like structure. The "puffing" heterochromatin has not only unique morphology but very special protein composition as well: i) it does not bind proteins specific for active chromatin and should therefore be referred to as a pseudopuff; and ii) it strongly associates with heterochromatin-specific proteins SU(VAR)3-7 and SUUR, despite HP1 and HP2 are depleted particularly from this polytene structure. The pseudopuff completes replication earlier than when it is compacted as heterochromatin, and underreplication of some DNA sequences within the pseudopuff is strongly suppressed. So, we show that pericentric heterochromatin is heterogeneous in its requirement for SU(VAR)3-9 with respect to the establishment of the condensed state, time of replication and DNA polytenization.
Key Words: SU(VAR)3-9, heterochromatic proteins, heterochromatin, polytene chromosomes, replication timing
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