Transcription by RNA polymerase I on nucleosomal templates requires binding
of the transcription termination factor TTF-I to a cognate site 160 bp ups
tream of the transcription start site. Binding of TTF-I is accompanied by c
hanges in the chromatin architecture which suggests that TTF-I recruits a r
emodeling activity to the rDNA promoter. We have cloned a cDNA that encodes
TIP5 (TTF-I-interacting protein 5), a 205 kDa protein that shares a number
of important protein domains with WSTF (Williams syndrome transcription fa
ctor) and hAcf1/WCRF180, the largest subunits of human chromatin remodeling
complexes hCHRAC and WCRF. TIP5 co-localizes with the basal RNA polymerase
I transcription factor UBF in the nucleolus and is associated with SNF2h.
The cellular TIP5-SNF2h complex, termed NoRC (nucleolar remodeling complex)
, induces nucleosome sliding in an ATP- and histone H4 tail-dependent fashi
on. The results suggest that NoRC is a novel nucleolar chromatin remodeling
machine that may serve a role in the regulation of the rDNA locus.