Rates of Gyrase Supercoiling and Transcription Elongation Control Supercoil Density in a Bacterial Chromosome 英文参考文献.docVIP

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Rates of Gyrase Supercoiling and Transcription Elongation Control Supercoil Density in a Bacterial Chromosome 英文参考文献.doc

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Rates of Gyrase Supercoiling and Transcription Elongation Control Supercoil Density in a Bacterial Chromosome 英文参考文献

RatesofGyraseSupercoilingandTranscription ElongationControlSupercoilDensityinaBacterial Chromosome NikolayRovinskiy1.,AndrewsAkwasiAgbleke1.,OlgaChesnokova1,ZhenhuaPang1,2 ,N. PatrickHiggins1* 1DepartmentofBiochemistryandMolecularGenetics,UniversityofAlabamaatBirmingham,Birmingham,Alabama,UnitedStatesofAmerica,2CathayIndustrialBiotech, Shanghai,China Abstract GyrasecatalyzesnegativesupercoilingofDNAinanATP-dependentreactionthathelpscondensebacterialchromosomes into a compact interwound ‘‘nucleoid.’’ The supercoil density (s) of prokaryotic DNA occurs in two forms. Diffusible supercoil density (sD) moves freely around the chromosome in 10kb domains, and constrained supercoil density (sC) resultsfrombindingabundantproteinsthatbend,loop,orunwindDNAatmanysites.Diffusibleandconstrainedsupercoils contribute roughly equally to the total in vivo negative supercoil density of WT cells, so s=sC+sD . Unexpectedly, Escherichia coli chromosomes have a 15% higher level of s compared to Salmonella enterica. To decipher critical mechanismsthatcanchangediffusiblesupercoildensityofchromosomes,weanalyzedstrainsofSalmonellausinga9kb ‘‘supercoil sensor’’ inserted attenpositions around thegenome.Thesensor contains acomplete Lacoperon flankedby directlyrepeatedresolvasebindingsites,andthesensorcanmonitorbothsupercoildensityandtranscriptionelongation ratesinWTandmutantstrains.RNAtranscriptioncaused(2)supercoilingtoincreaseupstreamanddecreasedownstream of highly expressed genes. Excess upstream supercoiling was relaxed by Topo I, and gyrase replenished downstream supercoil losses to maintain an equilibrium state. Strains with TS gyrase mutations growing at permissive temperature exhibitedsignificantsupercoillossesvaryingfrom30%ofWTlevelstoatotallossofsD atmostchromosomelocations. Supercoillosseswereinfluencedbytranscriptionbecauseadditionofrifampicin(Rif)causedsupercoildensitytorebound throughout the chromosome. Gyrase mutants that caused dramatic supercoil losses also reduced the