Many mature bacterial colonies and biofilms are complex three-dimensional (3D) structures. One key step in their developmental program is a transition from a two-dimensional (2D) monolayer into a 3D architecture. Despite the importance of controlling the growth of microbial colonies and biofilms in a variety of medical and industrial settings, the underlying physical mechanisms behind single-cell dynamics, collective behaviors of densely-packed cells, and 3D complex colony expansion remain largely unknown. In this work, we explore the mechanisms behind the 2D-to-3D transition of motile Pseudomonas aeruginosa colonies; we provide a new motility-induced, rate-dependent buckling mechanism for their out-of-plane growth. We find that swarming of motile bacterial colonies generate sustained in-plane flows. We show that the viscous shear stresses and dynamic pressures arising from these flows allow cells to overcome cell-substrate adhesion, leading to buckling of bacterial monolayers and growth into the third dimension. Modeling bacterial monolayers as 2D fluid films, we identify universal relationships that elucidate the competition between in-plane viscous stresses, pressure and cell-substrate adhesion. Furthermore, we show that bacterial monolayers can exhibit crossover from swarming to kinetically-arrested, glassy-like states above an onset density, resulting in distinct 2D-to-3D transition mechanisms. Combining experimental observations of P. aeruginosa colonies at single-cell resolution, molecular dynamics simulations of active systems, and theories of glassy dynamics and 2D fluid films, we develop a dynamical state diagram that predicts the state of the colony, and the mechanisms governing their 2D-to-3D transitions.
Engineered bacteria in which motility is reduced by local cell density generate periodic stripes of high and low density when spotted on agar plates. We study theoretically the origin and mechanism of this process in a kinetic model that includes growth and density-suppressed motility of the cells. The spreading of a region of immotile cells into an initially cell-free region is analyzed. From the calculated front profile we provide an analytic ansatz to determine the phase boundary between the stripe and the no-stripe phases. The influence of various parameters on the phase boundary is discus…
Adhesion of bacteria to liquid-liquid interfaces can play a role in the biodegradation of dispersed hydrocarbons and in biochemical and bioprocess engineering. Whereas thermodynamic factors underpinning adhesion are well studied, the role of bacterial activity on adhesion is less explored.Here, we show that bacterial motility enhances adhesion to surfactant-decorated oil droplets dispersed in artificial sea water. Motile Halomonas titanicae adhered to hexadecane droplets stabilized with dioctyl sodium sulfosuccinate (DOSS) more rapidly and at greater surface densities compared to nonmotile H. …
Bacterial migration through confined spaces is critical for several phenomena like: biofilm formation, bacterial transport in soils, and bacterial therapy against cancer . In the present work, \textit{E. coli} (strain K12-MG1655 WT) motility was characterized by recording and analyzing individual bacterium trajectories in a simulated quasi-2-dimensional porous medium. The porous medium was simulated by enclosing, between slide and cover slip, a bacterial-culture sample mixed with uniform 2.98 $μm$ spherical latex particles. The porosity of the medium was controlled by changing the latex partic…
The motility mechanism of certain rod-shaped bacteria has long been a mystery, since no external appendages are involved in their motion which is known as gliding. However, the physical principles behind gliding motility still remain poorly understood. Using myxobacteria as a canonical example of such organisms, we identify here the physical principles behind gliding motility, and develop a theoretical model that predicts a two-regime behavior of the gliding speed as a function of the substrate stiffness. Our theory describes the elastic, viscous, and capillary interactions between the bacteri…
We demonstrate 'differential dynamic microscopy' (DDM) for the fast, high throughput characterization of the dynamics of active particles. Specifically, we characterize the swimming speed distribution and the fraction of motile cells in suspensions of Escherichia coli bacteria. By averaging over ~10^4 cells, our results are highly accurate compared to conventional tracking. The diffusivity of non-motile cells is enhanced by an amount proportional to the concentration of motile cells.
Bacterial chemotaxis is one of the most extensively studied adaptive responses in cells. Many bacteria are able to bias their apparently random motion to produce a drift in the direction of the increasing chemoattractant concentration. It has been recognized that the particular motility pattern employed by moving bacteria has a direct impact on the efficiency of chemotaxis. The linear theory of chemotaxis pioneered by de Gennes allows for calculation of the drift velocity in small gradients for bacteria with basic motility patterns. However, recent experimental data on several bacterial specie…
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 poly…
Many types of bacteria swim by rotating a bundle of helical filaments also called flagella. Each filament is driven by a rotary motor and a very flexible hook transmits the motor torque to the filament. We model it by discretizing Kirchhoff's elastic-rod theory and develop a coarse-grained approach for driving the helical filament by a motor torque. A rotating flagellum generates a thrust force, which pushes the cell body forward and which increases with the motor torque. We fix the rotating flagellum in space and show that it buckles under the thrust force at a critical motor torque. Buckling…
Diverse processes--e.g. bioremediation, biofertilization, and microbial drug delivery--rely on bacterial migration in disordered, three-dimensional (3D) porous media. However, how pore-scale confinement alters bacterial motility is unknown due to the opacity of typical 3D media. As a result, models of migration are limited and often employ ad hoc assumptions. Here we reveal that the paradigm of run-and-tumble motility is dramatically altered in a porous medium. By directly visualizing individual Escherichia coli, we find that the cells are intermittently and transiently trapped as they navigat…
University of Pennsylvania（宾夕法尼亚大学）；National Institute for Mathematical Sciences（韩国数学科学国立研究所）；Chung-Ang University（中央大学）；University of Cincinnati（辛辛那提大学）
We study the transport of bacteria in a porous media modeled by a square channel containing one cylindrical obstacle via molecular dynamics simulations coupled to a lattice Boltzmann fluid. Our bacteria model is a rod-shaped rigid body which is propelled by a force-free mechanism. To account for the behavior of living bacteria, the model also incorporates a run-and-tumble process. The model bacteria are capable of hydrodynamically interacting with both of the channel walls and the obstacle. This enables the bacteria to get reoriented when experiencing a shear-flow. We demonstrate that this mod…
The presence of a nearby boundary is likely to be important in the life cycle and evolution of motile flagellate bacteria. This has led many authors to employ numerical simulations to model near-surface bacterial motion and compute hydrodynamic boundary effects. A common choice has been the method of images for regularized Stokeslets (MIRS); however, the method requires discretization sizes and regularization parameters that are not specified by any theory. To determine appropriate regularization parameters for given discretization choices in MIRS, we conducted dynamically similar macroscopic …
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, $Ω$, of motile Escherichia coli as a function of polymer concentration in polyvinylpyrrolidone…
Growing, Buckling, and Swirling: motility from polymerization
International Centre for Theoretical Sciences, Tata Institute of Fundamental Research（国际理论科学中心，塔塔基础研究所）；Center for Computational Biology, Flatiron Institute（计算生物学中心，平顿研究所）；Courant Institute of Mathematical Sciences, New York University（纽约大学库朗数学科学研究所）
Understanding how bacteria move in porous media is critical to applications in healthcare, agriculture, environmental remediation, and chemical sensing. Recent work has demonstrated that E. coli, which moves by run-and-tumble dynamics in a homogeneous medium, exhibits a new form of motility when confined in a disordered porous medium: hopping-and-trapping motility, in which cells perform rapid, directed hops punctuated by intervals of slow, undirected trapping. Here, we use direct visualization to shed light on how these processes depend on pore-scale confinement and cellular activity. We find…
Bacteria living on surfaces use different types of motility mechanisms to move on the surface in search of food or to form micro-colonies. Twitching is one such form of motility employed by bacteria such as N. gonorrhoeae, in which the polymeric extensions known as type IV pili mediate its movement. Pili extending from cell body adheres to the surface and pulls the bacteria by retraction. The bacterial movement is decided by the two-dimensional tug-of-war between the pili attached to the surface. Natural surfaces in which these micro-crawlers dwell are generally spatially inhomogeneous and hav…
Unraveling bacterial strategies for spatial exploration is crucial for understanding the complexity in the organization of life. Bacterial motility determines the spatio-temporal structure of microbial communities, controls infection spreading and the microbiota organization in guts or in soils. Most theoretical approaches for modeling bacterial transport rely on their run-and-tumble motion. For Escherichia coli, the run time distribution was reported to follow a Poisson process with a single characteristic time related to the rotational switching of the flagellar motors. However, direct measu…
As the constituent particles of a flock are polar and in a driven state, their interactions must, in general, be fore-aft asymmetric and non-reciprocal. Within a model that explicitly retains the classical spin angular momentum field of the particles we show that the resulting asymmetric contribution to interparticle torques, if large enough, leads to a buckling instability of the flock. Precisely this asymmetry also yields a natural mechanism for a difference between the speed of advection of information along the flock and the speed of the flock itself, concretely establishing that the absen…
Bacteria predate plants and animals by billions of years. Today, they are the world's smallest cells yet they represent the bulk of the world's biomass, and the main reservoir of nutrients for higher organisms. Most bacteria can move on their own, and the majority of motile bacteria are able to swim in viscous fluids using slender helical appendages called flagella. Low-Reynolds-number hydrodynamics is at the heart of the ability of flagella to generate propulsion at the micron scale. In fact, fluid dynamic forces impact many aspects of bacteriology, ranging from the ability of cells to reorie…
The present article is an invited contribution to the Encyclopedia of Complexity and System Science, Robert A. Meyers Ed., Springer New York (2009). It is a review of the biophysical mechanisms that underly cell motility. It mainly focuses on the eukaryotic cytoskeleton and cell-motility mechanisms. Bacterial motility as well as the composition of the prokaryotic cytoskeleton is only briefly mentioned. The article is organized as follows. In Section III, I first present an overview of the diversity of cellular motility mechanisms, which might at first glance be categorized into two different t…
Active motility and wetting cooperatively regulate liquid-liquid phase separation
Advanced Materials Thrust, Function Hub, The Hong Kong University of Science and Technology (Guangzhou)（香港科技大学（广州）功能枢纽先进材料 thrust）；Institute of Chinese Medical Sciences & State Key Laboratory of Mechanism and Quality of Chinese Medicine, University of Macau（澳门大学中医药学院及中药质量研究国家重点实验室）；Department of Pharmaceutical Sciences, Faculty of Health Sciences, University of Macau（澳门大学健康科学学院药学系）；Zhuhai UM Science and Technology Research Institute, University of Macau（澳门大学珠海科学技术研究院）；Research Center for Advanced Science and Technology, The University of Tokyo（东京大学先进科学技术研究中心）；Department of Fundamental Engineering, Institute of Industrial Science, The University of Tokyo（东京大学工业科学研究所基础工程系）
How systems are endowed with migration capacity is a fascinating question with implications ranging from the design of novel active systems to the control of microbial populations. Bacteria, which can be found in a variety of environments, have developed among the richest set of locomotion mechanisms both at the microscopic and collective levels. Here, we uncover experimentally a new mode of collective bacterial motility in humid environment through the depinning of bacterial droplets. While capillary forces are notoriously enormous at the bacterial scale, even capable of pinning water droplet…
Bacteria on surfaces exhibit collective behaviors, such as active turbulence and active stresses, which result from their motility, growth, and interactions with their local surroundings. However, interfacial deformations on soft surfaces and liquid interfaces caused by active growth, particularly over long time scales, are not well understood. Here, we describe experimental observations on the emergence of tubular structures arising from the growth of rod-shaped bacteria at the interface of oil droplets in water. Using microfluidics and timelapse microscopy, the dimensions and extension rates…
Suspensions of motile cells are model systems for understanding the unique mechanical properties of living materials which often consist of ensembles of self-propelled particles. We present here a quantitative comparison of theory against experiment for the rheology of such suspensions. The influence of motility on viscosities of cell suspensions is studied using a novel acoustically-driven microfluidic capillary-breakup extensional rheometer. Motility increases the extensional viscosity of suspensions of algal pullers, but decreases it in the case of bacterial or sperm pushers. A recent model…
Dense active systems are widespread in nature, examples range from bacterial colonies to biological tissues. Dense clusters of active particles can be obtained by increasing the packing fraction of the system or taking advantage of a peculiar phenomenon named motility-induced phase separation (MIPS). In this work, we explore the phase diagram of a two-dimensional model of active glass and show that disordered active materials develop a rich collective behaviour encompassing both MIPS and glassiness. We find that, although the glassy state is almost indistinguishable from that of equilibrium gl…
Motile Bacteria Modify Salt Precipitation Patterns in Dried Sessile Droplet
Georgia Institute of Technology（佐治亚理工学院）；University of Nebraska - Lincoln（内布拉斯加大学林肯分校）；Illinois Institute of Technology（伊利诺伊理工学院）；Blue Valley North High School（蓝谷北高中）
We model an enclosed system of bacteria, whose motility-induced phase separation is coupled to slow population dynamics. Without noise, the system shows both static phase separation and a limit cycle, in which a rising global population causes a dense bacterial colony to form, which then declines by local cell death, before dispersing to re-initiate the cycle. Adding fluctuations, we find that static colonies are now metastable, moving between spatial locations via rare and strongly nonequilibrium pathways, whereas the limit cycle becomes almost periodic such that after each redispersion event…
Motile cells often explore natural environments characterized by a high degree of structural complexity. Moreover cell motility is also intrinsically noisy due to spontaneous random reorientation and speed fluctuations. This interplay of internal and external noise sources gives rise to a complex dynamical behavior that can be strongly sensitive to details and hard to model quantitatively. In striking contrast to this general picture we show that the mean residence time of swimming bacteria inside artificial complex microstructures, can be quantitatively predicted by a generalization of a rece…
We present a generic theory for the dynamics of a stiff filament under tension, in an active medium with orientational correlations, such as a microtubule in contractile actin. In sharp contrast to the case of a passive medium, we find the filament can stiffen, and possibly oscillate, \textit{or} buckle, depending on the contractile or tensile nature of the activity \textit{and} the filament-medium anchoring interaction. We present experiments on the behaviour of microtubules in the growth cone of a neuron, which provide evidence for these apparently opposing behaviours. We also demonstrate a …
Active systems, or active matter, are self-driven systems which live, or function, far from equilibrium - a paradigmatic example which we focus on here is provided by a suspension of self-motile particles. Active systems are far from equilibrium because their microscopic constituents constantly consume energy from the environment in order to do work, for instance to propel themselves. The nonequilibrium nature of active matter leads to a variety of non-trivial intriguing phenomena. An important one which has recently been the subject of intense interest among biological and soft matter physici…
Recent studies have shown that packings of cells, both eukaryotic cellular tissues and growing or swarming bacterial colonies, can often be understood as active nematic fluids. A key property of volume-conserving active nematic model systems is chaotic self-mixing characterized by motile topological defects. However, for active nematics driven by growth rather than motility, less is understood about mixing and defect motion. Mixing could affect evolutionary outcomes in bacterial colonies by counteracting the tendency to spatially segregate into monoclonal sectors, which reduces the local genet…
Bacterial processes ranging from gene expression to motility and biofilm formation are constantly challenged by internal and external noise. While the importance of stochastic fluctuations has been appreciated for chemotaxis, it is currently believed that deterministic long-range fluid dynamical effects govern cell-cell and cell-surface scattering - the elementary events that lead to swarming and collective swimming in active suspensions and to the formation of biofilms. Here, we report the first direct measurements of the bacterial flow field generated by individual swimming Escherichia coli …
Bacterial turbulence drives interfacial waves and shape dynamics in phase-separated droplets
School of Physics and Astronomy, Institute of Natural Sciences and MOE-LSC, Shanghai Jiao Tong University（上海交通大学物理与天文学院、自然科学研究院和MOE-LSC）；Fujian Provincial Key Laboratory for Soft Functional Materials Research, Research Institute for Biomimetics and Soft Matter, Department of Physics, Xiamen University（厦门大学物理系仿生与软物质研究院福建省软功能材料研究重点实验室）；Universal Biology Institute, Graduate School of Science, The University of Tokyo（东京大学理学研究科全球生物学研究所）
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 …
Meso-scale turbulence was originally observed experimentally in various suspensions of swimming bacteria, as well as in the collective motion of active colloids. The corresponding large-scale dynamical patterns were reproduced in a simple model of a polar fluid, assuming a constant density of active particles. Recent, more detailed experimental studies revealed additional interesting aspects, such as anomalous velocity statistics and clustering phenomena. Those phenomena cannot be explained by currently available models for active polar fluids. Herein, we extend the continuum model suggested b…
We use moving light patterns to control the motion of {\it Escherichia coli} bacteria whose motility is photo-activated. Varying the pattern speed controls the magnitude and direction of the bacterial flux, and therefore the accumulation of cells in up- and down-stream reservoirs. We validate our results with two-dimensional simulations and a 1-dimensional analytic model, and use these to explore parameter space. We find that cell accumulation is controlled by a competition between directed flux and undirected, stochastic transport. We articulate design principles for using moving light patter…
Using a fluid-particle dynamics approach, we numerically study the effects of hydrodynamic interactions on the collective dynamics of active suspensions within a simple model for bacterial motility: each microorganism is modeled as a stroke-averaged dumbbell swimmer with prescribed dipolar force pairs. Using both simulations and qualitative arguments, we show that, when the separation between swimmers is comparable to their size, the swimmers' motions are strongly affected by activity-induced hydrodynamic forces. To further understand these effects, we investigate semidilute suspensions of swi…
The soil bacterium Myxococcus xanthus lives in densely packed groups that form dynamic three-dimensional patterns in response to environmental changes, such as droplet-like fruiting bodies during starvation. The development of these multicellular structures begins with the sequential formation of cell layers in a process that is poorly understood. Using confocal three-dimensional imaging, we find that motile rod-shaped M. xanthus cells are densely packed and aligned in each layer, forming an active nematic liquid crystal. Cell alignment is nearly perfect throughout the population except at poi…
How particles are deposited at the edge of evaporating droplets, i.e. the {\em coffee ring} effect, plays a crucial role in phenomena as diverse as thin-film deposition, self-assembly, and biofilm formation. Recently, microorganisms have been shown to passively exploit and alter these deposition dynamics to increase their survival chances under harshening conditions. Here, we show that, as the droplet evaporation rate slows down, bacterial mobility starts playing a major role in determining the growth dynamics of the edge of drying droplets. Such motility-induced dynamics can influence several…
