RIO

RIO kinases RIOK1, RIOK2, and RIOK3 are ancient, highly conserved atypical kinases dedicated to ribosome biogenesis. RIOK1 and RIOK2 are required for the final cytoplasmic maturation of the 40S subunit, acting at pre-rRNA processing and nuclear-export checkpoints, while RIOK3 has additional reported roles in signaling and immune regulation. Structurally they adopt a distinctive fold that lacks several canonical eukaryotic protein kinase motifs, and they exploit ATP binding and hydrolysis as a conformational switch during pre-40S particle remodeling in addition to their catalytic activity. Because rapidly proliferating cells and many tumors depend on elevated ribosome production, RIOK activity is being explored as an anticancer target. Recombinant RIOKs are core reagents for ribosome-assembly reconstitution and maturation studies.