Packing and delivery of a genetic message

Balbiani rings (BRs) 1 and 2 are two exceptionally large chromosomal puffs on chromosome IV in the salivary glands of the dipteran Chironomus tentans. The BR genes are 35-40 kb, contain four short introns, and encode salivary polypeptides of one million molecular weight. They have proven uniquely suited for visualization of the assembly and transport of a specific messenger ribonucleoprotein (RNP) particle. A BR transcript is packed with proteins into a thin RNP fibril, which is folded into a compact ring-like structure. The completed BR particle is released from the gene and moves randomly in the nucleoplasm before it becomes associated with the nuclear pore complex. The passage through the nuclear pore is a highly ordered process with a series of consecutive steps: initial binding, docking, unfolding, movement through the pore with the 5' end of the transcript in the lead, and exit into the cytoplasm. On the cytoplasmic side, the RNA becomes immediately engaged in protein synthesis. Recently, several major proteins in the BR particle have been identified and characterized. They are added to the BR RNA molecule concomitantly with transcription. During the ensuing RNA transport, the various proteins behave differently, some remaining in the nucleus, others entering the cytoplasm coupled to the RNA. The flow pattern of a given protein seems to be closely related to the specific function of the protein. The RNA-binding proteins are likely to play various active roles during gene expression rather than being solely packaging proteins. Finally, it is emphasized that the co-transcriptional loading of the transcript with proteins is probably a key process in gene expression that to a large extent determines the fate of an mRNA both in the nucleus and the cytoplasm.