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Register a new userThe colloidal nature of complex fluids leads to enhanced motility of flagellated bacteria
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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.
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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.
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.
We have developed a lab work module where we teach undergraduate students how to quantify the dynamics of a suspension of microscopic particles, measuring and analyzing the motion of those particles at the individual level or as a group. Differential Dynamic Microscopy (DDM) is a relatively recent technique that precisely does that and constitutes an alternative method to more classical techniques such as dynamics light scattering (DLS) or video particle tracking (VPT). DDM consists in imaging a particle dispersion with a standard light microscope and a camera. The image analysis requires the students to code and relies on digital Fourier transform to obtain the intermediate scattering function, an autocorrelation function that characterizes the dynamics of the dispersion. We first illustrate DDM on the textbook case of colloids where we measure the diffusion coefficient. Then we show that DDM is a pertinent tool to characterize biologic systems such as motile bacteria i.e.bacteria that can self propel, where we not only determine the diffusion coefficient but also the velocity and the fraction of motile bacteria. Finally, so that our paper can be used as a tutorial to the DDM technique, we have joined to this article movies of the colloidal and bacterial suspensions and the DDM algorithm in both Matlab and Python to analyze the movies.
Colloidal particles hold promise for mobilizing and removing trapped immiscible fluids from porous media, with implications for key energy and water applications. Most studies focus on accomplishing this goal using particles that can localize at the immiscible fluid interface. Therefore, researchers typically seek to optimize the surface activity of particles, as well as their ability to freely move through a pore space with minimal deposition onto the surrounding solid matrix. Here, we demonstrate that deposition can, surprisingly, promote mobilization of a trapped fluid from a porous medium without requiring any surface activity. Using confocal microscopy, we directly visualize both colloidal particles and trapped immiscible fluid within a transparent, three-dimensional (3D) porous medium. We find that as non-surface active particles deposit on the solid matrix, increasing amounts of trapped fluid become mobilized. We unravel the underlying physics by analyzing the extent of deposition, as well as the geometry of trapped fluid droplets, at the pore scale: deposition increases the viscous stresses on trapped droplets, overcoming the influence of capillarity that keeps them trapped. Given an initial distribution of trapped fluid, this analysis enables us to predict the extent of fluid mobilized through colloidal deposition. Taken together, our work reveals a new way by which colloids can be harnessed to mobilize trapped fluid from a porous medium.
The study of how photosynthetic organisms convert light offers insight not only into natures evolutionary process, but may also give clues as to how best to design and manipulate artificial photosynthetic systems -- and also how far we can drive natural photosynthetic systems beyond normal operating conditions, so that they can harvest energy for us under otherwise extreme conditions. In addition to its interest from a basic scientific perspective, therefore, the goal to develop a deep quantitative understanding of photosynthesis offers the potential payoff of enhancing our current arsenal of alternative energy sources for the future. In the following Chapter, we consider the trade-off between dynamics, structure and function of light harvesting membranes in Rps. Photometricum purple bacteria, as a model to highlight the priorities that arise when photosynthetic organisms adapt to deal with the ever-changing natural environment conditions.
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