Abstract
A number of Gram-negative bacteria have a membrane surrounding their flagella, referred to as the flagellar sheath, which is continuous with the outer membrane. The flagellar sheath was initially described in Vibrio metschnikovii in the early 1950s as an extension of the outer cell wall layer that completely surrounded the flagellar filament. Subsequent studies identified other bacteria that possess flagellar sheaths, most of which are restricted to a few genera of the phylum Proteobacteria. Biochemical analysis of the flagellar sheaths from a few bacterial species revealed the presence of lipopolysaccharide, phospholipids, and outer membrane proteins in the sheath. Some proteins localize preferentially to the flagellar sheath, indicating mechanisms exist for protein partitioning to the sheath. Recent cryo-electron tomography studies have yielded high resolution images of the flagellar sheath and other structures closely associated with the sheath, which has generated insights and new hypotheses for how the flagellar sheath is synthesized. Various functions have been proposed for the flagellar sheath, including preventing disassociation of the flagellin subunits in the presence of gastric acid, avoiding activation of the host innate immune response by flagellin, activating the host immune response, adherence to host cells, and protecting the bacterium from bacteriophages.
Highlights
The bacterial flagellum is a complex organelle used for motility and is organized into three basic structures referred to as the basal body, hook and filament
The 1983 review of bacterial flagellar sheaths by Sjoblad and colleagues begins with the statement, “ bacterial flagellar sheaths were observed over 30 years ago, they may still be characterized as structures in search of a function” [3]
Additional roles have been postulated for bacterial flagellar sheaths over the last 37 years, limitations in our knowledge of flagellar sheath biosynthesis and the lack of mutants that synthesize sheath-less flagella thwart efforts to confirm proposed functions for flagellar sheaths
Summary
The bacterial flagellum is a complex organelle used for motility and is organized into three basic structures referred to as the basal body, hook and filament. The filament in several genera of Gram-negative bacteria, is surrounded by a membranous sheath that is contiguous with the outer membrane. Flagella in these bacteria are located almost exclusively at the cell pole, and occur as a single flagellum at one cell pole (polar flagellum), as a single flagellum at each cell pole (amphitrichous or bipolar flagella) or as multiple flagella at one cell pole (lophotrichous flagella). It has been nearly four decades since the last comprehensive review of the bacterial flagellar sheath [3], which was a major impetus for this review
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