13 Chromatin remodelling complexes of the SWI/SNF family function in the formation of 14 nucleosome-depleted regions and transcriptionally active promoters in the eukaryote 15 genome. The structure of the Saccharomyces cerevisiae SWI/SNF family member RSC in 16 complex with a nucleosome substrate reveals five protein modules and suggests key 17 features of the remodelling mechanism. A DNA-interacting module grasps extra-18 nucleosomal DNA and helps to recruit RSC to promoters. The ATPase and arm modules 19 sandwich the nucleosome disc with their 'SnAC' and 'finger' elements, respectively. The 20 translocase motor engages with the edge of the nucleosome at superhelical location +2 to 21 pump DNA along the nucleosome, resulting in a sliding of the histone octamer along DNA. 22

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