Molecular Organization and Assembly of the Export Apparatus of Flagellar Type III Secretion Systems

The bacterial flagellum is a supramolecular motility machine consisting of the basal body, the hook, and the filament. For construction of the flagellum beyond the cellular membranes, a type III protein export apparatus uses ATP and proton-motive force (P

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Contents 1 2

Introduction.......................................................................................................................... Structure of the Transmembrane Export Gate Complex .................................................... 2.1 FlhA Ring Structure ................................................................................................... 2.2 FlhB ............................................................................................................................ 2.3 Core Structure of the Export Gate Complex ............................................................. 3 Cytoplasmic ATPase Ring Complex .................................................................................. 4 Sorting Platform .................................................................................................................. 5 Assembly of the Type III Protein Export Apparatus ......................................................... 6 Conclusion ........................................................................................................................... References ..................................................................................................................................

Abstract The bacterial flagellum is a supramolecular motility machine consisting of the basal body, the hook, and the filament. For construction of the flagellum beyond the cellular membranes, a type III protein export apparatus uses ATP and proton-motive force (PMF) across the cytoplasmic membrane as the energy sources to transport flagellar component proteins from the cytoplasm to the distal end of the growing flagellar structure. The protein export apparatus consists of a PMF-driven transmembrane export gate complex and a cytoplasmic ATPase complex. In T. Minamino (&)  A. Kawamoto  M. Kinoshita  K. Namba Graduate School of Frontier Biosciences, Osaka University, 1-3 Yamadaoka, Suita, Osaka 565-0871, Japan e-mail: [email protected] A. Kawamoto Institute for Protein Research, Osaka University, 3-2 Yamadaoka, Suita, Osaka 565-0871, Japan K. Namba Center for Biosystems Dynamics Research & SPring-8 Center, RIKEN, 1-3 Yamadaoka, Suita, Osaka 565-0871, Japan Current Topics in Microbiology and Immunology https://doi.org/10.1007/82_2019_170 © Springer Nature Switzerland AG 2019

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addition, the basal body C ring acts as a sorting platform for the cytoplasmic ATPase complex that efficiently brings export substrates and type III export chaperone–substrate complexes from the cytoplasm to the export gate complex. In this book chapter, we will summarize our current understanding of molecular organization and assembly of the flagellar type III protein export apparatus.

1 Introduction Bacteria swim in liquid environments and move on solid surfaces by rotating a very long filamentous assembly called the bacterial flagellum. The flagellum consists of at least three parts: the basal body as a bidirectional rotary motor, the hook as a universal joint, and the filament