LAUSR

Significance In order to understand the evolution of complex biological machines like the bacterial flagellar motor, it is crucial to know what each component does and when it arose. Here, we show that a subcomplex of the motor thought to act as a bushing for the spinning motor likely also serves another function—it plugs the hole in the outer membrane left when the flagellum disassembles. Moreover, this component and function is ancient, since it appears in diverse phyla without evidence of recent gene transfer. The bacterial flagellum is an amazing nanomachine. Understanding how such complex structures arose is crucial to our understanding of cellular evolution. We and others recently reported that in several Gammaproteobacterial species, a relic subcomplex comprising the decorated P and L rings persists in the outer membrane after flagellum disassembly. Imaging nine additional species with cryo-electron tomography, here, we show that this subcomplex persists after flagellum disassembly in other phyla as well. Bioinformatic analyses fail to show evidence of any recent horizontal transfers of the P- and L-ring genes, suggesting that this subcomplex and its persistence is an ancient and conserved feature of the flagellar motor. We hypothesize that one function of the P and L rings is to seal the outer membrane after motor disassembly.