How the concentric organization of the nucleolus and chromatin ensures accuracy of ribosome biogenesis and drives transport.

The biogenetic transport of ribosomal subunit precursors must be conducted with precision to ensure production of functional ribosomes. With a focus on ribosome biogenesis in higher eukaryotic cells, we here discuss the following: (1) the concentric organization of the phases/subcompartments of the nucleus-including chromatin, (2) why the nucleolus reorganizes when ribosomal RNA synthesis is inhibited, and (3) the mechanism responsible for vectorial transport of particulate subunit intermediates between subcompartments. We call attention to evidence that (1) nucleolar proteins can access the entire volume of the nucleus, (2) that the packaging of rDNA is a key determinant of topology, (3) the constancy of contacts between subcompartments, and the likely importance of a Brownian ratchet for imparting both directionality and quality control upon transport. Transport appears to depend on "self-immersion," whereby the surfaces of particulate intermediates successively interact with components of the surrounding milieux, each of which may be thought of as a distinct solvent. The result is a vectorial and ordered process.