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To swim through a viscous fluid, a flagellated bacterium must overcome the fluid drag on its body by rotating a flagellum or a bundle of multiple flagella. Because the drag increases with the size of bacteria, it is expected theoretically that the swimming speed of a bacterium inversely correlates with its body length. Nevertheless, despite extensive research, the fundamental size–speed relation of flagellated bacteria remains unclear with different experiments reporting conflicting results. Here, by critically reviewing the existing evidence and synergizing our own experiments of large sample sizes, hydrodynamic modeling, and simulations, we demonstrate that the average swimming speed ofEscherichia coli, a premier model of peritrichous bacteria, is independent of their body length. Our quantitative analysis shows that such a counterintuitive relation is the consequence of the collective flagellar dynamics dictated by the linear correlation between the body length and the number of flagella of bacteria. Notably, our study reveals how bacteria utilize the increasing number of flagella to regulate the flagellar motor load. The collective load sharing among multiple flagella results in a lower load on each flagellar motor and therefore faster flagellar rotation, which compensates for the higher fluid drag on the longer bodies of bacteria. Without this balancing mechanism, the swimming speed of monotrichous bacteria generically decreases with increasing body length, a feature limiting the size variation of the bacteria. Altogether, our study resolves a long-standing controversy over the size–speed relation of flagellated bacteria and provides insights into the functional benefit of multiflagellarity in bacteria.
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Effects of swimming environment on bacterial motility
Swimming trajectories of bacteria can be altered by environmental conditions, such as background flow and physical barriers, that limit the free swimming of bacteria. We present a comprehensive model of a bacterium that consists of a rod-shaped cell body and a flagellum which is composed of a motor, a hook, and a filament. The elastic flagellum is modeled based on the Kirchhoff rod theory, the cell body is considered to be a rigid body, and the hydrodynamic interaction of a bacterium near a wall is described by regularized Stokeslet formulation combined with the image system. We consider three environmental conditions: (1) a rigid surface is placed horizontally and there is no shear flow, (2) a shear fluid flow is present and the bacterium is near the rigid surface, and (3) while the bacterium is near the rigid surface and is under shear flow, an additional sidewall which is perpendicular to the rigid surface is placed. Each environmental state modifies the swimming behavior. For the first condition, there are two modes of motility, trap and escape, whether the bacterium stays near the surface or moves away from the surface as we vary the physical and geometrical properties of the model bacterium. For the second condition, there exists a threshold of shear rate that classifies the motion into two types of paths in which the bacterium takes either a periodic coil trajectory or a linear trajectory. For the last condition, the bacterium takes upstream motility along the sidewall for lower shear rates and downstream motility for larger shear flow rates.
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- Award ID(s):
- 1853591
- PAR ID:
- 10337822
- Date Published:
- Journal Name:
- Physics of Fluids
- Volume:
- 34
- Issue:
- 3
- ISSN:
- 1070-6631
- Page Range / eLocation ID:
- 031907
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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- Free Publicly Accessible Full Text
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Accepted Manuscript1.0
- Journal Article:
- https://doi.org/10.1063/5.0082768
