Cell Membranes Resist Flow.

The fluid-mosaic model posits a liquid-like plasma membrane, which can flow in response to tension gradients. It is widely assumed that membrane flow transmits local changes in membrane tension across the cell in milliseconds, mediating long-range signaling. Here, we show that propagation of membrane tension occurs quickly in cell-attached blebs but is largely suppressed in intact cells. The failure of tension to propagate in cells is explained by a fluid dynamical model that incorporates the flow resistance from cytoskeleton-bound transmembrane proteins. Perturbations to tension propagate diffusively, with a diffusion coefficient Dσ ∼0.024 µm2/s in HeLa cells. In primary endothelial cells, local increases in membrane tension lead only to local activation of mechanosensitive ion channels and to local vesicle fusion. Thus, membrane tension is not a mediator of long-range intracellular signaling, but local variations in tension mediate distinct processes in sub-cellular domains.

The mechanical world of bacteria.

In the wild, bacteria are predominantly associated with surfaces as opposed to existing as free-swimming, isolated organisms. They are thus subject to surface-specific mechanics, including hydrodynamic forces, adhesive forces, the rheology of their surroundings, and transport rules that define their encounters with nutrients and signaling molecules. Here, we highlight the effects of mechanics on bacterial behaviors on surfaces at multiple length scales, from single bacteria to the development of multicellular bacterial communities such as biofilms.

Hydration solids.

Hygroscopic biological matter in plants, fungi and bacteria make up a large fraction of Earth's biomass1. Although metabolically inert, these water-responsive materials exchange water with the environment and actuate movement2-5 and have inspired technological uses6,7. Despite the variety in chemical composition, hygroscopic biological materials across multiple kingdoms of life exhibit similar mechanical behaviours including changes in size and stiffness with relative humidity8-13. Here we report atomic force microscopy measurements on the hygroscopic spores14,15 of a common soil bacterium and develop a theory that captures the observed equilibrium, non-equilibrium and water-responsive mechanical behaviours, finding that these are controlled by the hydration force16-18. Our theory based on the hydration force explains an extreme slowdown of water transport and successfully predicts a strong nonlinear elasticity and a transition in mechanical properties that differs from glassy and poroelastic behaviours. These results indicate that water not only endows biological matter with fluidity but also can-through the hydration force-control macroscopic properties and give rise to a 'hydration solid' with unusual properties. A large fraction of biological matter could belong to this distinct class of solid matter.

Capillary forces generated by biomolecular condensates.

Liquid-liquid phase separation and related phase transitions have emerged as generic mechanisms in living cells for the formation of membraneless compartments or biomolecular condensates. The surface between two immiscible phases has an interfacial tension, generating capillary forces that can perform work on the surrounding environment. Here we present the physical principles of capillarity, including examples of how capillary forces structure multiphase condensates and remodel biological substrates. As with other mechanisms of intracellular force generation, for example, molecular motors, capillary forces can influence biological processes. Identifying the biomolecular determinants of condensate capillarity represents an exciting frontier, bridging soft matter physics and cell biology.

Building on-chip cytoskeletal circuits via branched microtubule networks.

Controllable platforms to engineer robust cytoskeletal scaffolds have the potential to create novel on-chip nanotechnologies. Inspired by axons, we combined the branching microtubule (MT) nucleation pathway with microfabrication to develop "cytoskeletal circuits." This active matter platform allows control over the adaptive self-organization of uniformly polarized MT arrays via geometric features of microstructures designed within a microfluidic confinement. We build and characterize basic elements, including turns and divisions, as well as complex regulatory elements, such as biased division and MT diodes, to construct various MT architectures on a chip. Our platform could be used in diverse applications, ranging from efficient on-chip molecular transport to mechanical nano-actuators. Further, cytoskeletal circuits can serve as a tool to study how the physical environment contributes to MT architecture in living cells.

Bacterial spores respond to humidity similarly to hydrogels.

Bacterial spores have outstanding properties from the materials science perspective, which allow them to survive extreme environmental conditions. Recent work by [S. G. Harrellson et al., Nature 619, 500-505 (2023)] studied the mechanical properties of Bacillus subtilis spores and the evolution of these properties with the change of humidity. The experimental measurements were interpreted assuming that the spores behave as water-filled porous solids, subjected to hydration forces. Here, we revisit their experimental data using literature data on vapor sorption on spores and ideas from polymer physics. We demonstrate that upon the change of humidity, the spores behave like rubber with respect to their swelling, elasticity, and relaxation times. This picture is consistent with the knowledge of the materials comprising the bacterial cell walls-cross-linked peptidoglycan. Our results provide an interpretation of the mechanics of bacterial spores and can help in developing synthetic materials mimicking the mechanical properties of the spores.

High-throughput measurement of elastic moduli of microfibers by rope coiling.

There are many fields where it is of interest to measure the elastic moduli of tiny fragile fibers, such as filamentous bacteria, actin filaments, DNA, carbon nanotubes, and functional microfibers. The elastic modulus is typically deduced from a sophisticated tensile test under a microscope, but the throughput is low and limited by the time-consuming and skill-intensive sample loading/unloading. Here, we demonstrate a simple microfluidic method enabling the high-throughput measurement of the elastic moduli of microfibers by rope coiling using a localized compression, where sample loading/unloading are not needed between consecutive measurements. The rope coiling phenomenon occurs spontaneously when a microfiber flows from a small channel into a wide channel. The elastic modulus is determined by measuring either the buckling length or the coiling radius. The throughput of this method, currently 3,300 fibers per hour, is a thousand times higher than that of a tensile tester. We demonstrate the feasibility of the method by testing a nonuniform fiber with axially varying elastic modulus. We also demonstrate its capability for in situ inline measurement in a microfluidic production line. We envisage that high-throughput measurements may facilitate potential applications such as screening or sorting by mechanical properties and real-time control during production of microfibers.

Evaporation of alcohol droplets on surfaces in moist air.

Droplets of alcohol-based formulations are common in applications from sanitizing sprays to printing inks. However, our understanding of the drying dynamics of these droplets on surfaces and the influence of ambient humidity is still very limited. Here, we report the drying dynamics of picoliter droplets of isopropyl alcohol deposited on a surface under controlled humidity. Condensation of water vapor in the ambient environment onto alcohol droplets leads to unexpectedly complex drying behavior. As relative humidity (RH) increases, we observed a variety of phenomena including enhanced spreading, nonmonotonic changes in the drying time, the formation of pancake-like shapes that suppress the coffee-ring effect, and the formation of water-rich films around an alcohol-rich drop. We developed a lubrication model that accounts for the coupling between the flow field within the drop, the shape of the drop, and the vapor concentration field. The model reproduces many of the experimentally observed morphological and dynamic features, revealing the presence of unusually large spatial compositional gradients within the evaporating droplet and surface-tension-gradient-driven flows arising from water condensation/evaporation at the surface of the droplet. One unexpected feature from the simulation is that water can evaporate and condense concurrently in different parts of the drop, providing fundamental insights that simpler models based on average fluxes lack. We further observed rim instabilities at higher RH that are well-described by a model based on the Rayleigh-Plateau instability. Our findings have implications for the testing and use of alcohol-based disinfectant sprays and printing inks.

Oil-on-water droplets faceted and stabilized by vortex halos in the subphase.

For almost 200 y, the dominant approach to understand oil-on-water droplet shape and stability has been the thermodynamic expectation of minimized energy, yet parallel literature shows the prominence of Marangoni flow, an adaptive gradient of interfacial tension that produces convection rolls in the water. Our experiments, scaling arguments, and linear stability analysis show that the resulting Marangoni-driven high-Reynolds-number flow in shallow water overcomes radial symmetry of droplet shape otherwise enforced by the Laplace pressure. As a consequence, oil-on-water droplets are sheared to become polygons with distinct edges and corners. Moreover, subphase flows beneath individual droplets can inhibit the coalescence of adjacent droplets, leading to rich many-body dynamics that makes them look alive. The phenomenon of a "vortex halo" in the liquid subphase emerges as a hidden variable.

Anomalous crystalline ordering of particles in a viscoelastic fluid under high shear.

Addition of particles to a viscoelastic suspension dramatically alters the properties of the mixture, particularly when it is sheared or otherwise processed. Shear-induced stretching of the polymers results in elastic stress that causes a substantial increase in measured viscosity with increasing shear, and an attractive interaction between particles, leading to their chaining. At even higher shear rates, the flow becomes unstable, even in the absence of particles. This instability makes it very difficult to determine the properties of a particle suspension. Here, we use a fully immersed parallel plate geometry to measure the high-shear-rate behavior of a suspension of particles in a viscoelastic fluid. We find an unexpected separation of the particles within the suspension resulting in the formation of a layer of particles in the center of the cell. Remarkably, monodisperse particles form a crystalline layer which dramatically alters the shear instability. By combining measurements of the velocity field and torque fluctuations, we show that this solid layer disrupts the flow instability and introduces a single-frequency component to the torque fluctuations that reflects a dominant velocity pattern in the flow. These results highlight the interplay between particles and a suspending viscoelastic fluid at very high shear rates.

Hydrodynamically induced helical particle drift due to patterned surfaces.

Advances in microfabrication enable the tailoring of surfaces to achieve optimal sorting, mixing, and focusing of complex particulate suspensions in microfluidic devices. Corrugated surfaces have proved to be a powerful tool to manipulate particle motion for a variety of applications, yet the fundamental physical mechanism underlying the hydrodynamic coupling of the suspended particles and surface topography has remained elusive. Here, we study the hydrodynamic interactions between sedimenting spherical particles and nearby corrugated surfaces, whose corrugations are tilted with respect to gravity. Our experiments show three-dimensional, helical particle trajectories with an overall drift along the corrugations, which agree quantitatively with our analytical perturbation theory. The theoretical predictions reveal that the interaction of the disturbance flows, induced by the particle motion, with the corrugations generates locally a transverse anisotropy of the pressure field, which explains the helical dynamics and particle drift. We demonstrate that this dynamical behavior is generic for various surface shapes, including rectangular, sinusoidal, and triangular corrugations, and we identify surface characteristics that produce an optimal particle drift. Our findings reveal a universal feature inherent to particle transport near patterned surfaces and provide fundamental insights for future microfluidic applications that aim to enhance the focusing or sorting of particulate suspensions.

Rayleigh-Taylor Instability in Soft Viscoelastic Solids.

We present an experimental characterization of the gravity-driven Rayleigh-Taylor instability in viscoelastic solids. The instability creates periodic patterns on the free surface of the soft solids that are distinct from the previously studied elastic Rayleigh-Taylor instability. The experimental results are supported by the linear stability analysis reported here. We identify the dependence of the steady-state pattern of deformations on the gel's geometry, complex shear modulus, and surface tension. This study provides quantitative measures applicable to the design of tunable surface textures, soft machines, and 3D structures.

Low-Reynolds-number, biflagellated Quincke swimmers with multiple forms of motion.

In the limit of zero Reynolds number (Re), swimmers propel themselves exploiting a series of nonreciprocal body motions. For an artificial swimmer, a proper selection of the power source is required to drive its motion, in cooperation with its geometric and mechanical properties. Although various external fields (magnetic, acoustic, optical, etc.) have been introduced, electric fields are rarely utilized to actuate such swimmers experimentally in unbounded space. Here we use uniform and static electric fields to demonstrate locomotion of a biflagellated sphere at low Re via Quincke rotation. These Quincke swimmers exhibit three different forms of motion, including a self-oscillatory state due to elastohydrodynamic-electrohydrodynamic interactions. Each form of motion follows a distinct trajectory in space. Our experiments and numerical results demonstrate a method to generate, and potentially control, the locomotion of artificial flagellated swimmers.

Evidence for biosurfactant-induced flow in corners and bacterial spreading in unsaturated porous media.

The spread of pathogenic bacteria in unsaturated porous media, where air and liquid coexist in pore spaces, is the major cause of soil contamination by pathogens, soft rot in plants, food spoilage, and many pulmonary diseases. However, visualization and fundamental understanding of bacterial transport in unsaturated porous media are currently lacking, limiting the ability to address the above contamination- and disease-related issues. Here, we demonstrate a previously unreported mechanism by which bacterial cells are transported in unsaturated porous media. We discover that surfactant-producing bacteria can generate flows along corners through surfactant production that changes the wettability of the solid surface. The corner flow velocity is on the order of several millimeters per hour, which is the same order of magnitude as bacterial swarming, one of the fastest known modes of bacterial surface translocation. We successfully predict the critical corner angle for bacterial corner flow to occur based on the biosurfactant-induced change in the contact angle of the bacterial solution on the solid surface. Furthermore, we demonstrate that bacteria can indeed spread by producing biosurfactants in a model soil, which consists of packed angular grains. In addition, we demonstrate that bacterial corner flow is controlled by quorum sensing, the cell-cell communication process that regulates biosurfactant production. Understanding this previously unappreciated bacterial transport mechanism will enable more accurate predictions of bacterial spreading in soil and other unsaturated porous media.

Hierarchical transitions and fractal wrinkling drive bacterial pellicle morphogenesis.

Bacterial cells can self-organize into structured communities at fluid-fluid interfaces. These soft, living materials composed of cells and extracellular matrix are called pellicles. Cells residing in pellicles garner group-level survival advantages such as increased antibiotic resistance. The dynamics of pellicle formation and, more generally, how complex morphologies arise from active biomaterials confined at interfaces are not well understood. Here, using Vibrio cholerae as our model organism, a custom-built adaptive stereo microscope, fluorescence imaging, mechanical theory, and simulations, we report a fractal wrinkling morphogenesis program that differs radically from the well-known coalescence of wrinkles into folds that occurs in passive thin films at fluid-fluid interfaces. Four stages occur: growth of founding colonies, onset of primary wrinkles, development of secondary curved ridge instabilities, and finally the emergence of a cascade of finer structures with fractal-like scaling in wavelength. The time evolution of pellicle formation depends on the initial heterogeneity of the film microstructure. Changing the starting bacterial seeding density produces three variations in the sequence of morphogenic stages, which we term the bypass, crystalline, and incomplete modes. Despite these global architectural transitions, individual microcolonies remain spatially segregated, and thus, the community maintains spatial and genetic heterogeneity. Our results suggest that the memory of the original microstructure is critical in setting the morphogenic dynamics of a pellicle as an active biomaterial.

A calibration-free model of micropipette aspiration for measuring properties of protein condensates.

There is growing evidence that biological condensates, which are also referred to as membraneless organelles, and liquid-liquid phase separation play critical roles regulating many important cellular processes. Understanding the roles these condensates play in biology is predicated on understanding the material properties of these complex substances. Recently, micropipette aspiration (MPA) has been proposed as a tool to assay the viscosity and surface tension of condensates. This tool allows the measurement of both material properties in one relatively simple experiment, in contrast to many other techniques that only provide one or a ratio of parameters. While this technique has been commonly used in the literature to determine the material properties of membrane-bound objects dating back decades, the model describing the dynamics of MPA for objects with an external membrane does not correctly capture the hydrodynamics of unbounded fluids, leading to a calibration parameter several orders of magnitude larger than predicted. In this work we derive a new model for MPA of biological condensates that does not require any calibration and is consistent with the hydrodynamics of the MPA geometry. We validate the predictions of this model by conducting MPA experiments on a standard silicone oil of known material properties and are able to predict the viscosity and surface tension using MPA. Finally, we reanalyze with this new model the MPA data presented in previous works for condensates formed from LAF-1 RGG domains.

Dynamics of Droplet Pinch-Off at Emulsified Oil-Water Interfaces: Interplay between Interfacial Viscoelasticity and Capillary Forces.

The presence of submicrometer structures at liquid-fluid interfaces modifies the properties of many science and technological systems by lowering the interfacial tension, creating tangential Marangoni stresses, and/or inducing surface viscoelasticity. Here we experimentally study the break-up of a liquid filament of a silica nanoparticle dispersion in a background oil phase that contains surfactant assemblies. Although self-similar power-law pinch-off is well documented for threads of Newtonian fluids, we report that when a viscoelastic layer is formed in situ at the interface, the pinch-off dynamics follows an exponential decay. Recently, such exponential neck thinning was found theoretically when surface viscous effects were taken into account. We introduce a simple approach to calculate the effective relaxation time of viscoelastic interfaces and estimate the thickness of the interfacial layer and the viscoelastic properties of liquid-fluid interfaces, where the direct measurement of interfacial rheology is not possible.

Volatile Droplets on Water are Sculpted by Vigorous Marangoni-Driven Subphase Flow.

The shapes of highly volatile oil-on-water droplets become strongly asymmetric when they are out of equilibrium. The unsaturated organic vapor atmosphere causes evaporation and leads to a strong Marangoni flow in the bath, unlike that previously seen in the literature. Inspecting these shapes experimentally on millisecond and submillimeter time and length scales and theoretically by scaling arguments, we confirm that Marangoni-driven convection in the subphase mechanically stresses the droplet edges to an extent that increases for organic droplets of smaller contact angle and accordingly smaller thickness. The viscous stress generated by the subphase overcomes the thermodynamic Laplace pressure. The oil droplets develop copious regularly spaced fingers, and these fingers develop spike-shaped and branched treelike structures. Unlike this behavior for single-component (surfactant-free) oil droplets, droplets composed of two miscible (surfactant-free) organic liquids develop a rim of the less volatile component along the droplet perimeter, from which jets of monodisperse smaller droplets eject periodically due to the Rayleigh-Plateau instability. When evaporation shrinks droplets to µm size, their shapes fluctuate chaotically, and ellipsoidal shapes rupture into smaller daughter droplets when subphase convection flow pulls them in opposite directions. The shape of the evaporating oil droplets is kneaded and sculpted by vigorous flow in the water subphase.

A note about convected time derivatives for flows of complex fluids.

We present a direct derivation of the typical time derivatives used in a continuum description of complex fluid flows, harnessing the principles of the kinematics of line elements. The evolution of the microstructural conformation tensor in a flow and the physical interpretation of different derivatives then follow naturally.