Rigid body rotation and chiral reorientation combine in filamentous E. coli swimming in low-Re flows

초록

When treated with doses of antibiotics below the minimum inhibitory concentration, bacterial cell division turns off, but cell growth does not. As a result, rod-like bacteria, including E. coli, can elongate many times their original length without increasing their width. The swimming behavior of these filamentous bacteria through small channels may provide insights into how bacteria that survive antibiotic treatment can reach channel walls. Such swimming behaviors in settings like hospital tubing may signal precursors to adhesion, biofilm formation, and infection. Despite the importance of understanding the behavior of bacteria not killed by antibiotics, the hydrodynamics of filamentous bacteria swimming in external flows has not received much attention. We study the swimming behavior of stressed, filamentous E. coli. In quiescence, highly elongated E. coli swim with a sinusoidal undulating motion, suggesting rigid body rotation of long, rigid, buckled cell bodies. In low Reynolds number pressure-driven flows through a microchannel, the undulating motion becomes irregular; it may even stop and start within a particular bacterial trajectory. We refer to this behavior in flow as "wiggling." Rigid body rotation persists in flow, appearing as a high-frequency change in body orientation on top of a slower one. Chiral reorientation can explain the slower reorientation frequency. We quantify swimming behaviors in two different flow rates and observe rheotaxis in addition to the preferential orientation of bacterial bodies. Faster flow constrains wiggling bacteria trajectories and orientations compared to those observed in slower flow, with rheotaxis taking bacteria toward the wall. Interestingly, not all bacteria in flow wiggle. Populations of non-motile "non-wiggling" filamentous E. coli follow streamlines, without preferential alignment of their orientation. Non-motile bacteria do not behave like chiral rods propelled and rotated by flagellar bundles, but like rigid rods. Motility slows the swimmers in comparison. Differentiating these two populations may have important implications for understanding consequences of motility loss that inevitably occurs as bacteria die.

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