Hydrodynamics and direction change of tumbling bacteria.

The bacterium Escherichia coli (E. coli) swims in viscous fluids by rotating several helical flagellar filaments, which are gathered in a bundle behind the cell during 'runs' wherein the cell moves steadily forward. In between runs, the cell undergoes quick 'tumble' events, during which at least one flagellum reverses its rotation direction and separates from the bundle, resulting in erratic motion in place and a random reorientation of the cell. Alternating between runs and tumbles allows cells to sample space by stochastically changing their propulsion direction after each tumble. The change of direction during a tumble is not uniformly distributed but is skewed towards smaller angles with an average of about 62°-68°, as first measured by Berg and Brown (1972). Here we develop a theoretical approach to model the angular distribution of swimming E. coli cells during tumbles. We first use past experimental imaging results to construct a kinematic description of the dynamics of the flagellar filaments during a tumble. We then employ low-Reynolds number hydrodynamics to compute the consequences of the kinematic model on the force and torque balance of the cell and to deduce the overall change in orientation. The results of our model are in good agreement with experimental observations. We find that the main change of direction occurs during the 'bundling' part of the process wherein, at the end of a tumble, the dispersed flagellar filaments are brought back together in the helical bundle, which we confirm using a simplified forced-sphere model.