Abstract

Flagella, the primary means of motility in bacteria, are helical filaments that function as microscopic propellers composed of thousands of copies of the protein flagellin. Here, we show that many bacteria encode “giant” flagellins, greater than a thousand amino acids in length, and that two species that encode giant flagellins, the marine γ-proteobacteria Bermanella marisrubri and Oleibacter marinus, produce monopolar flagellar filaments considerably thicker than filaments composed of shorter flagellin monomers. We confirm that the flagellum from B. marisrubri is built from its giant flagellin. Phylogenetic analysis reveals that the mechanism of evolution of giant flagellins has followed a stepwise process involving an internal domain duplication followed by insertion of an additional novel insert. This work illustrates how “the” bacterial flagellum should not be seen as a single, idealised structure, but as a continuum of evolved machines adapted to a range of niches.

Highlights

  • Flagella are the organelles responsible for motility in diverse bacterial species

  • Flagellins are secreted by the flagellar type III secretion system and travel through the hollow core of the growing filament to assemble into a helical array at the distal tip [4]

  • We propose a mechanism for evolution of these giant flagellins based on phylogenetic analysis

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Summary

Introduction

Flagella are the organelles responsible for motility in diverse bacterial species. Flagella form helical filaments, several microns long, connected to a basal body that spans the cell envelope and functions as a rotary motor [1]. In contrast to the highly conserved N- and C-terminal flagellin domains, it has long been known that the central region of this protein is highly variable in both sequence and length [7]. Typhimurium and confirmed this experimentally by measuring filament thickness in two hydrocarbon-degrading marine γproteobacteria with predicted giant flagellins: Bermanella marisrubri Red65, originally isolated from the Red Sea [11] and Oleibacter marinus 2O1, originally isolated from Indonesian seawater [12].

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