Bacterial attachment to a fluid interface can lead to the formation of a film with physicochemical properties that evolve with time. We study the time evolution of interface (micro)mechanics for interfaces between oil and bacterial suspensions by following the motion of colloidal probes trapped by capillarity to determine the interface microrheology. Initially, active bacteria at and near the interface drive superdiffusive motion of the colloidal probes. Over timescales of minutes, the bacteria form a viscoelastic film which we discuss as a quasi-two-dimensional, active, glassy system. To study late stage mechanics of the film, we use pendant drop elastometry. The films, grown over tens of hours on oil drops, are expanded and compressed by changing the drop volume. For small strains, by modeling the films as 2D Hookean solids, we estimate the film elastic moduli, finding values similar to those reported in the literature for the bacteria themselves. For large strains, the films are highly hysteretic. Finally, from wrinkles formed on highly compressed drops, we estimate film bending energies. The dramatic restructuring of the interface by such robust films has broad implications, e.g. in the study of active colloids, in understanding the community dynamics of bacteria, and in applied settings including bioremediation.
Bacteria at fluid interfaces endure physical and chemical stresses unique to these highly asymmetric environments. The responses of Pseudomonas aeruginosa PAO1 and PA14 to a hexadecane-water interface are compared. PAO1 cells form elastic films of bacteria, excreted polysaccharides and proteins, whereas PA14 cells move actively without forming an elastic film. Studies of PAO1 mutants show that, unlike solid-supported biofilms, elastic interfacial film formation occurs in the absence of flagella, pili, or certain polysaccharides. Highly induced genes identified in transcriptional profiling incl…
Bacterial cellulose biofilms are complex networks of strong interwoven nanofibers that control transport and protect bacterial colonies in the film. Design of diverse applications of bacterial cellulose films also relies on understanding and controlling transport through the fiber mesh, and transport simulations of the films are most accurate when guided by experimental characterization of the structures and the resultant diffusion inside. Diffusion through such films is a function of their key microstructural length scales, determining how molecules, as well as particles and microorganisms, p…
The population growth of a Staphylococcus aureus culture was followed by rheological measurements, under steady-state and dynamic shear flows. We observed a rich viscoelastic behaviour as a consequence of the bacteria activity. First, the viscosity increased ~10 times due to cell multiplication and aggregation. This viscosity increase presented several drops and full recoveries, which are reproducible, allowing us to evoke the existence of a percolation phenomenon. Eventually, as the bacteria population reached a final stage of development, fulfilling the sample volume, the viscosity returned …
The activity of growing living bacteria was investigated using real-time and in situ rheology -- in stationary and oscillatory shear. Two different strains of the human pathogen Staphylococcus aureus -- strain COL and its isogenic cell wall autolysis mutant -- were considered in this work. For low bacteria density, strain COL forms small clusters, while the mutant, presenting deficient cell separation, forms irregular larger aggregates. In the early stages of growth, when subjected to a stationary shear, the viscosity of both strains increases with the population of cells. As the bacteria reac…
Improving building energy performance requires the development of new highly insulative materials. An affordable retrofitting solution comprising a thin film could improve the resistance to heat flow in both residential and commercial buildings and reduce overall energy consumption. Here we propose cellulose aerogel films formed from pellicles produced by the bacteria Gluconacetobacter hansenii as insulation materials. We studied the impact of density and nanostructure on the aerogels' thermal properties. Thermal conductivity as low as 13 mW/(K*m) was measured for native pellicle-based aerogel…
There is a deep analogy between the physics of crystalline solids and the behavior of superfluids, dating back to pioneering work of Phillip Anderson, Paul Martin and others. The stiffness to shear deformations in a periodic crystal resembles the superfluid density that controls the behavior of supercurrents in neutral superfluids such as He^4. Dislocations in solids have a close analogy with quantized vortices in superfluids. Remarkable recent experiments on the way rod-shaped bacteria elongate their cell walls have focused attention on the dynamics and interactions of point-like dislocation …
Hydrodynamic flows in biological systems are often generated by active chiral processes near or on surfaces. Important examples are beating cilia, force generation in actomyosin networks, and motile bacteria interacting with surfaces. Here we develop a coarse grained description of active chiral films that captures generic features of flow and rotation patterns driven by chiral motors. We discuss force and torque balances within the film and on the surface and highlight the role of the intrinsic rotation field. We arrive at a two dimensional effective theory and discuss our results in the cont…
The equilibrium structure and dynamics of a single polymer chain in a thermal solvent is by now well-understood in terms of scaling laws. Here we consider a polymer in a bacterial bath, i.e. in a solvent consisting of active particles which bring in nonequilibrium fluctuations. Using computer simulations of a self-avoiding polymer chain in two dimensions which is exposed to a dilute bath of active particles, we show that the Flory-scaling exponent is unaffected by the bath activity provided the chain is very long. Conversely, for shorter chains, there is a nontrivial coupling between the bacte…
In the present work we simulate the basic two-dimensional dynamics of swarming E. coli bacteria on the surface of a moderately soft agar plate. Individual bacteria are modelled by self-propelled ridged bodies (agents), which interact with each other only through inelastic collision and with the highly viscous environment through damping forces. The motion of single agents is modelled closely corresponding to the behaviour of swimming bacteria. The dynamics of the model were adjusted to reproduce the experimental measurements of swimming E. coli K 12. Accordingly, simulations with loosely packe…
We study theoretically the phase diagram of compressible active polar gels such as the actin network of eukaryotic cells. Using generalized hydrodynamics equations, we perform a linear stability analysis of the uniform states in the case of an infinite bidimensional active gel to obtain the dynamic phase diagram of active polar films. We predict in particular modulated flowing phases, and a macroscopic phase separation at high activity. This qualitatively accounts for experimental observations of various active systems, such as acto-myosin gels, microtubules and kinesins in vitro solutions, or…
Motile bacteria are known to accumulate at surfaces, eventually leading to changes in bacterial motility and bio-film formation. We use a novel two-colour, three-dimensional Lagrangian tracking technique, to follow simultaneously the body and the flagella of a wild-type ${\it Escherichia~coli}$. We observe long surface residence times and surface escape corresponding mostly to immediately antecedent tumbling. A motility model accounting for a large behavioural variability in run-time duration, reproduces all experimental findings and gives new insights into surface trapping efficiency.
Dense suspensions of swimming bacteria are known to exhibit collective behaviour arising from the interplay of steric and hydrodynamic interactions. Unconfined suspensions exhibit transient, recurring vortices and jets, whereas those confinedin circular domains may exhibit order in the form of a spiral vortex. Here we show that confinement into a long and narrow macroscopic `racetrack' geometry stabilises bacterial motion to form a steady unidirectional circulation. This motion is reproduced in simulations of discrete swimmers that reveal the crucial role that bacteria-driven fluid flows play …
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 Pseudom…
We confine a dense suspension of motile \textit{Escherichia coli} inside a spherical droplet in a water-in-oil emulsion, creating a "bacterially" propelled droplet. We show that droplets move in a persistent random walk, with a persistence time $τ\sim 0.3\, {\rm s}$, a long-time diffusion coefficient $D\sim 0.5\, μ{\rm m}^2/{\rm s}$, and an average instantaneous speed $V\sim 1.5\, μ{\rm m/s}$ when the bacterial suspension is at the maximum studied concentration. Several droplets are analyzed, varying the drop radius and bacterial concentration. We show that the persistence time, diffusion coef…
Swimming bacteria create long-range velocity fields that stir a large volume of fluid and move around passive particles dispersed in the fluid. Recent experiments and simulations have shown that long-time mean-squared displacement of passive particles in a bath of swimming bacteria exhibits diffusive behaviour with the effective diffusion coefficient significantly larger than its thermal counterpart. Comprehensive theoretical prediction of this effective diffusion coefficient and understanding of the enhancement mechanism remain a challenge. Here, we adapt the kinetic theory by Lin et al., J. …
Spontaneous oscillations and geometric cutoff in confined bacterial swarms
Center of Materials Science and Optoelectronics Engineering, College of Materials Science and Opto-Electronic Technology, University of Chinese Academy of Sciences (UCAS)（中国科学院大学材料科学与光电技术学院物质科学与光电工程中心）；Department of Physics, Hong Kong Baptist University（香港浸会大学物理系）；Center for Interdisciplinary Studies, Westlake University（西湖大学交叉研究中心）
We present a joint experimental and computational study of the effect of bacterial motion on micron-scale colloids contained in a two-dimensional suspension of Bacillus subtilis. With respect to previous work using E. coli, here we introduce a novel experimental set-up that allows us to realise a two-dimensional bacterial suspension insensitive to either evaporation or fluid flow. By analysing the mean square displacements of both bacteria and colloids, we confirm the existence of a crossover from super-diffusive behaviour at short time scales to normal diffusion at longer times. We also study…
The overdamped Brownian motion of a self-propelled particle which is driven by a projected internal force is studied by solving the Langevin equation analytically. The "active" particle under study is restricted to move along a linear channel. The direction of its internal force is orientationally diffusing on a unit circle in a plane perpendicular to the substrate. An additional time-dependent torque is acting on the internal force orientation. The model is relevant for active particles like catalytically driven Janus particles and bacteria moving on a substrate. Analytical results for the fi…
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…
This work presents the study of some new anomalous electromagnetic effects in graphite-like thin carbon films. These are: The fast switching (1nanosecond) of electrical conductivity The detection of microwave radiation and its temperature dependence The oscillations of film stack magnetization in the magnetic field of 1-5 T. The optical radiation under process of spasmodic switching of conductivity Results of magnetic force microscopy (MFM), DC SQUID magnetization, reversed Josephson effect (RJE), and resistance measurements in thin carbon arc (CA) films are presented. The observation of a RJE…
Flocks of birds, schools of fish, mixtures of motors and cytoskeletal filaments, swimming bacteria and driven granular media are systems of interacting motile units that exhibit collective behaviour. These can all be described as active matter systems, since each individual unit takes energy from an internal energy depot and transduces it into work performed on the environment. We review a model for cyanobacterial phototaxis, emphasising the differences from other models for collective behaviour in active matter systems. The interactions between individual cells during phototaxis are dominated…
The inter-connection between the elasticity of a dielectric film and the wetting of a sessile drop on the same, under an applied electrical voltage, remains unaddressed. Here, we report the electrowetting-on-dielectric (EWOD) behaviour of sessile drops on dielectric elastomer films of varying elasticities- from an apparently rigid dielectric film to a soft, deformable dielectric film. Our results reveal that the elasticity of the underlying dielectric film provides an additional control over the droplet electrowetting behaviour, which may be best addressed from free energy based consideration,…
In this work we introduce a stochastic model to describe directional changes in the movement of swimming bacteria. We use the probability density function (PDF) of turn angles, measured on tumbling wild-type {\it E. coli}, to build a Langevin equation for the deflection of the bacterial body swimming in isotropic media. We have solved this equation analytically by means of the Green function method and shown that three parameters are sufficient to describe the movement: the characteristic time, the steady-state solution and the control parameter. We conclude that the tumble motion, which is ma…
Motivated by the need to conceive freely-precessing gyroscopes for detecting acoustic frame-dragging predicted recently in rotating acoustic analogue black holes, we report an incipient investigation on the hydrodynamics of nematic active fluids. With a specific assumption on barotropicity of a nematic fluid, we discern acoustic analogue black hole spacetimes experienced by linear perturbations of the velocity potential. For vanishingly small diffusivity of the active particles, linear perturbations of the active particle concentration reveal a profile with an enhancement close to the acoustic…
The spatial organization of bacteria in dense biofilms is key to their collective behaviour, and understanding it will be important for medical and technological applications. Here we study the morphology of a compact biofilm that undergoes unidirectional growth, and determine the condition for the stability of the growing interface as a function of the nutrient concentration and mechanical tension. Our study suggests that transient behaviour may play an important role in shaping the structure of a biofilm.
We formulate a simple Landau type model describing macroscopic behaviour recently discovered new smectic phases composed of achiral bent-shaped molecules. Films of such smectics exhibit three types of ordering related to dipole polarization, molecular tilt, and chirality. However due to specific third order coupling of the order parameters these three types of symmetry-breaking are not independent ones, and this fact leads to specific domain structures really observed in experiments.
We study the adsorption of a fluid in the grand canonical ensemble occurring at a planar heterogeneous wall which is decorated with a chemical stripe of width $L$. We suppose that the material of the stripe strongly preferentially adsorbs the liquid in contrast to the outer material which is only partially wet. This competition leads to the nucleation of a droplet of liquid on the stripe, the height $h_m$ and shape of which (at bulk two-phase coexistence) has been predicted previously using mesoscopic interfacial Hamiltonian theory. We test these predictions using a microscopic Fundamental Mea…
Self-propelled colloidal objects, such as motile bacteria or synthetic microswimmers, have microscopically irreversible individual dynamics - a feature they share with all living systems. The incoherent behaviour of individual swimmers can then be harnessed (or "rectified") by microfluidic devices that create systematic motions impossible in equilibrium. Examples include flow of rotor particles round a circuit, steady rotation of a gear wheel in a bacterial bath, and pumping of bacteria between chambers by "funnel gates". Here we present a computational proof-of-concept study, showing that suc…
We investigate the critical behaviour of the $N$-component Euclidean $λϕ^4$ model at leading order in $\frac{1}{N}$-expansion. We consider it in three situations: confined between two parallel planes a distance $L$ apart from one another, confined to an infinitely long cylinder having a square cross-section of area $A$ and to a cubic box of volume $V$. Taking the mass term in the form $m_{0}^2=α(T - T_{0})$, we retrieve Ginzburg-Landau models which are supposed to describe samples of a material undergoing a phase transition, respectively in the form of a film, a wire and of a grain, whose bulk…
