Multiflagellarity leads to the size-independent swimming speed of bacteria

Flagella are essential organelles of bacteria enabling their swimming motility. While monotrichous or uniflagellar bacteria possess a single flagellum at one pole of their body, peritrichous bacteria grow multiple flagella over the body surface, which form a rotating helical bundle propelling the bacteria forward. Although the adaptation of bacterial cellular features is under strong evolutionary pressure, existing evidence suggests that multiflagellarity confers no noticeable benefit to the swimming of peritrichous bacteria in bulk fluids compared with uniflagellar bacteria. This puzzling result poses a long-standing question: why does multiflagellarity emerge given the high metabolic cost of flagellar synthesis? Contrary to the prevailing wisdom that its benefit lies beyond the basic function of flagella in steady swimming, here we show that multiflagellarity provides a significant selective advantage to bacteria in terms of their swimming ability, allowing bacteria to maintain a constant swimming speed over a wide range of body size. By synergizing experiments of immense sample sizes with quantitative hydrodynamic modeling and simulations, we reveal how bacteria utilize the increasing number of flagella to regulate the flagellar motor load, which leads to faster flagellar rotation neutralizing the higher fluid drag on their larger bodies. Without such a precise balancing mechanism, the swimming speed of uniflagellar bacteria generically decreases with increasing body size. Our study sheds light on the origin of multiflagellarity, a ubiquitous cellular feature of bacteria. The uncovered difference between uniflagellar and multiflagellar swimming is important for understanding environmental influence on bacterial morphology and useful for designing artificial flagellated microswimmers.Comment: 23 pages, 4 figure

Physical mechanism reveals bacterial slowdown above a critical number of flagella

Numerous studies have explored the link between bacterial swimming and the number of flagella, a distinguishing feature of motile multiflagellated bacteria. We revisit this open question using augmented slender-body theory simulations, in which we resolve the full hydrodynamic interactions within a bundle of helical filaments rotating and translating in synchrony. Unlike previous studies, our model considers the full torque-speed relationship of the bacterial flagellar motor, revealing its significant impact on multiflagellated swimming. Because the viscous load per motor decreases with flagellar number, the bacterial flagellar motor (BFM) transitions from the high-load to the low-load regime at a critical number of filaments, leading to bacterial slowdown as further flagella are added to the bundle. We explain the physical mechanism behind the observed slowdown as an interplay between the load-dependent generation of torque by the motor, and the load-reducing cooperativity between flagella, which consists of both hydrodynamic and non-hydrodynamic components. The theoretically predicted critical number of flagella is remarkably close to the values reported for the model organism Escherichia coli. Our model further predicts that the critical number of flagella increases with viscosity, suggesting that bacteria can enhance their swimming capacity by growing more flagella in more viscous environments, consistent with empirical observations