Growth of Flagellar Filaments of Escherichia coli is Independent of Filament Length

Abstract

Bacterial flagellar filaments grow at their distal ends, from flagellin that travels through a central channel (\sim 2 nm) in diameter. The flagellin is extruded from the cytoplasm by a pump powered by a proton motive force (PMF). We measured filament growth in cells near the mid-exponential-phase with flagellin bearing a specific cysteine-for-serine substitution, allowing filaments to be labeled with sulfhydryl-specific fluorescent dyes. We labeled filaments first with a green maleimide dye and then, following an additional period of growth, with a red maleimide dye. The contour lengths of the green and red segments were measured. The average lengths of red segments ((2.3 \mu m)) were the same regardless of the lengths of the green segments from which they grew (ranging from less than 1 to more than (9 \mu m) in length). Thus, flagellar filaments do not grow at a rate that decreases exponentially with length, as formerly supposed. If flagellar filaments were broken by viscous shear, the broken filaments continued to grow. Identical results were obtained whether flagellin was expressed from fliC on the chromosome under the control of its native promoter or on a plasmid under the control of the arabinose promoter.