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BIMM 130 Dr. Milton Saier Bacterial Flagellum Bacterial flagellum: 10 microns long (the helical period is ~2 microns); the size of the flagellum is about 1 billion Da! There are ~50 flagellum (flg, flh, fli) genes; about ~25 structural genes. + 2 mot + 6 che + 5 primary receptor genes encoding MCPs + many more secondary receptors E. coli: ~10 flagelli; 20,000 rpm; H+; 30 /sec (~ 2 mm/min) - other bacteria 1-1,000 flagelli (i.e., swarming), 101-105 rpm H+ or Na+; 10-200 /sec (~20 mm/min) If E. coli was the size of a car, how fast would it go? E. coli: 1 2 mm/min = 120 mm/hr = 0.12 meters/hr Car: 5 meters 65 mph ~100 km/hr; max, 250 km/hr; size difference 5 x 106 0.12 meters/hr (5 x 106) = 0.6 x 106 meters/hr = 600 km/hr But some bacteria swim 10x faster: 6,000 km/hr; faster than any jet airplane! Motor: a. Stator: MotAB is the ion selector (H+ or Na+) (Asai et al., 2003) b. Rotor: the flagellum (FliFGMN). The C-terminal domain of FliG interacts with MotAB to produce the torque. FliG, FliM and FliN form a complex that interacts with MotAB; only their genes when mutated (1) fla, (2) mot and (3) che phenotypes, i.e., phenotypes corresponding to (1) an absence of flagelli, (2) intact flagelli but no motor function, and (3) normal flagelli with an active motor but lacking chemotactic responses. MotAB complexes have been reconstituted in artificial membranes yielding channels specific for H+ or Na+. References: (Terashima et al. 2008) (Minamino et al. 2008) bushing OM PG IM L-ring rotor P-ring S-ring M-ring Stator (MotA + MotB) C-ring FliG Charged ridge Structure of the flagellar basal region and its association with the bacterial cell envelope. OM, outer membrane; PG, peptidoglycan cell wall; IM, inner membrane. References: Anderson JK, Smith TG, Hoover TR. Sense and sensibility: flagellum-mediated gene regulation. Trends Microbiol. 2010 Jan;18(1):30-7. Asai Y, Yakushi T, Kawagishi I, Homma M. Ion-coupling determinants of Na+-driven and H+driven flagellar motors. J Mol Biol. 2003 Mar 21;327(2):453-63. Egelman EH. Reducing irreducible complexity: divergence of quaternary structure and function in macromolecular assemblies. Curr Opin Cell Biol. [Review]. 2010 Feb;22(1):68-74. Minamino T, Imada K, Namba K (2008) Molecular motors of the bacterial flagella. Curr Opin Struct Biol 18:693-701 Smith TG, Hoover TR. Deciphering bacterial flagellar gene regulatory networks in the genomic era. Adv Appl Microbiol. [Review]. 2009;67:257-95. Terashima H, Kojima S, Homma M (2008) Flagellar motility in bacteria structure and function of flagellar motor. Int Rev Cell Mol Biol 270:39-85