No Arabic abstract
It is widely believed that the swimming speed, $v$, of many flagellated bacteria is a non-monotonic function of the concentration, $c$, of high-molecular-weight linear polymers in aqueous solution, showing peaked $v(c)$ curves. Pores in the polymer solution were suggested as the explanation. Quantifying this picture led to a theory that predicted peaked $v(c)$ curves. Using new, high-throughput methods for characterising motility, we have measured $v$, and the angular frequency of cell-body rotation, $Omega$, of motile Escherichia coli as a function of polymer concentration in polyvinylpyrrolidone (PVP) and Ficoll solutions of different molecular weights. We find that non-monotonic $v(c)$ curves are typically due to low-molecular weight impurities. After purification by dialysis, the measured $v(c)$ and $Omega(c)$ relations for all but the highest molecular weight PVP can be described in detail by Newtonian hydrodynamics. There is clear evidence for non-Newtonian effects in the highest molecular weight PVP solution. Calculations suggest that this is due to the fast-rotating flagella `seeing a lower viscosity than the cell body, so that flagella can be seen as nano-rheometers for probing the non-Newtonian behavior of high polymer solutions on a molecular scale.
The ability to propel against flows, i.e., to perform positive rheotaxis, can provide exciting opportunities for applications in targeted therapeutics and non-invasive surgery. To date, no biocompatible technologies exist for navigating microparticles upstream when they are in a background fluid flow. Inspired by many naturally occurring microswimmers such as bacteria, spermatozoa, and plankton that utilize the non-slip boundary conditions of the wall to exhibit upstream propulsion, here, we report on the design and characterization of self-assembled microswarms that can execute upstream motility in a combination of external acoustic and magnetic fields. Both acoustic and magnetic fields are safe to humans, non-invasive, can penetrate deeply into the human body, and are well-developed in clinical settings. The combination of both fields can overcome the limitations encountered by single actuation methods. The design criteria of the acoustically-induced reaction force of the microswarms, which is needed to perform rolling-type motion, are discussed. We show quantitative agreement between experimental data and our model that captures the rolling behaviour. The upstream capability provides a design strategy for delivering small drug molecules to hard-to-reach sites and represents a fundamental step toward the realization of micro- and nanosystem-navigation against the blood flow.
Peritrichous bacteria such as Escherichia coli swim in viscous fluids by forming a helical bundle of flagellar filaments. The filaments are spatially distributed around the cell body to which they are connected via a flexible hook. To understand how the swimming direction of the cell is determined, we theoretically investigate the elastohydrodynamic motility problem of a multi-flagellated bacterium. Specifically, we consider a spherical cell body with a number N of flagella which are initially symmetrically arranged in a plane in order to provide an equilibrium state. We analytically solve the linear stability problem and find that at most 6 modes can be unstable and that these correspond to the degrees of freedom for the rigid-body motion of the cell body. Although there exists a rotation-dominated mode that generates negligible locomotion, we show that for the typical morphological parameters of bacteria the most unstable mode results in linear swimming in one direction accompanied by rotation around the same axis, as observed experimentally.
The natural habitats of microorganisms in the human microbiome and ocean and soil ecosystems are full of colloids and macromolecules, which impart non-Newtonian flow properties drastically affecting the locomotion of swimming microorganisms. Although the low-Reynolds-number hydrodynamics of the swimming of flagellated bacteria in simple Newtonian fluids has been well developed, our understanding of bacterial motility in complex non-Newtonian fluids is still primitive. Even after six decades of research, fundamental questions about the nature and origin of bacterial motility enhancement in polymer solutions are still under debate. Here, we study the motility of flagellated bacteria in colloidal suspensions of varying sizes and volume fractions. We find that bacteria in dilute colloidal suspensions display quantitatively the same motile behaviors as those in dilute polymer solutions, where a universal particle-size-dependent motility enhancement up to 80% is uncovered, accompanied by strong suppression of bacterial wobbling. By virtue of the well-controlled size and the hard-sphere nature of colloids, the finding not only resolves the long-standing controversy over bacterial motility enhancement in complex fluids but also challenges all the existing theories using polymer dynamics to address the swimming of flagellated bacteria in dilute polymer solutions. We further develop a simple physical model incorporating the colloidal nature of complex fluids, which quantitatively explains bacterial wobbling dynamics and mobility enhancement in both colloidal and polymeric fluids. Our study sheds light on the puzzling motile behaviors of bacteria in complex fluids relevant to a wide range of microbiological processes and provides a cornerstone in engineering bacterial swimming in complex environments.
Self-sustained turbulent structures have been observed in a wide range of living fluids, yet no quantitative theory exists to explain their properties. We report experiments on active turbulence in highly concentrated 3D suspensions of Bacillus subtilis and compare them with a minimal fourth-order vector-field theory for incompressible bacterial dynamics. Velocimetry of bacteria and surrounding fluid, determined by imaging cells and tracking colloidal tracers, yields consistent results for velocity statistics and correlations over two orders of magnitude in kinetic energy, revealing a decrease of fluid memory with increasing swimming activity and linear scaling between energy and enstrophy. The best-fit model parameters allow for quantitative agreement with experimental data.
Colonies of bacterial cells endowed with a pili-based self-propulsion machinery represent an ideal model system for studying how active adhesion forces affect structure and dynamics of many-particle systems. As a novel computational tool, we describe here a highly parallel molecular dynamics simulation package for modeling of textit{Neisseria gonorrhoeae} colonies. Simulations are employed to investigate growth of bacterial colonies and the dependence of the colony structure on cell-cell interactions. In agreement with experimental data, active pilus retraction is found to enhance local ordering. For mixed colonies consisting of different types of cell types, the simulations show a segregation of cell types depending on the pili-mediated interactions, as seen in experiments. Using a simulated experimental setup, we study the power-spectral density of colony-shape fluctuations and the associated fluctuation-response relation. The simulations predict a strong violation of the equilibrium fluctuation-response relation across the measurable frequency range. Lastly, we illustrate the essential role of active force generation for colony dynamics by showing that pilus-mediated activity drives the spreading of colonies on surfaces and the invasion of narrow channels.