I n eukaryotic cells containing tandem repeated ribosomal RNA genes, there appears a specialized region of chromatin, carrying out gene transcription, rRNA processing, and nascent ribosomal subunit assembly—the nucleolus. One of the first intracellular structures to be described (Montgomery, 1898; Franke, 1988), the nucleolus was established in the 1960s as the center of ribosome synthesis (Sch...