Review Article

Regulation of Ribosomal RNA Production by RNA Polymerase I: Does Elongation Come First?

Figure 3

Miller spreading of nontranscribed rDNA. Single-gene analysis of nontranscribed rRNA genes reveals nucleosomal (a) and nonnucleosomal (b) organization. Chromatin spreading (upper panel) and a schematic representation of rDNA spreading (lower panel) are shown from strain NOY1071, bearing 25 rDNA copies. Transcribed rRNA genes (green) are identified by high Pol I density, and nascent rRNAs are shown in black. Non-transcribed regions are depicted in red. Intergenic spaces (IGSs) are short (600 nm) and can be easily distinguished from inactive rRNA genes. Inactive genes are flanked by two IGSs. Due to the DNA wrapping around nucleosomes, nontranscribed genes associated with nucleosomes appear shorter (2,100 nm − (2 × 600 nm) = 900 nm) than genes devoid of nucleosomes (3,100 nm − (2 × 600) = 1,900 nm). With 15 nucleosomes detected and each wrapped around 146 bp, we observed a length reduction of approximately 1,000 nm.