second-chance signal transduction explains cooperative flagellar switching二次信号转导解释合作鞭毛切换.pdfVIP

Second-Chance Signal Transduction Explains Cooperative Flagellar Switching 1 2 3 Henry G. Zot *, Javier E. Hasbun , Nguyen Van Minh 1 Department of Biology, University of West Georgia, Carrollton, Georgia, United States of America, 2 Department of Physics, University of West Georgia, Carrollton, Georgia, United States of America, 3 Department of Mathematics, University of West Georgia, Carrollton, Georgia, United States of America Abstract The reversal of flagellar motion (switching) results from the interaction between a switch complex of the flagellar rotor and a torque-generating stationary unit, or stator (motor unit). To explain the steeply cooperative ligand-induced switching, present models propose allosteric interactions between subunits of the rotor, but do not address the possibility of a reaction that stimulates a bidirectional motor unit to reverse direction of torque. During flagellar motion, the binding of a ligand-bound switch complex at the dwell site could excite a motor unit. The probability that another switch complex of the rotor, moving according to steady-state rotation, will reach the same dwell site before that motor unit returns to ground state will be determined by the independent decay rate of the excited-state motor unit. Here, we derive an analytical expression for the energy coupling between a switch complex and a motor unit of the stator complex of a flagellum, and demonstrate that this model accounts for the cooperative switching response without the need for allosteric interactions. The analytical result can be reproduced by simulation when (1) the motion of the rotor delivers a subsequent ligand-bound switch to the excited motor unit, thereby providing the excited motor unit with a second chance to remain ex