Pseudomonas aeruginosa distinguishes surfaces by stiffness using retraction of type IV pili.

The ability of eukaryotic cells to differentiate surface stiffness is fundamental for many processes like stem cell development. Bacteria were previously known to sense the presence of surfaces, but the extent to which they could differentiate stiffnesses remained unclear. Here we establish that the human pathogen Pseudomonas aeruginosa actively measures surface stiffness using type IV pili (TFP). Stiffness sensing is nonlinear, as induction of the virulence factor regulator is peaked with stiffness in a physiologically important range between 0.1 kPa (similar to mucus) and 1,000 kPa (similar to cartilage). Experiments on surfaces with distinct material properties establish that stiffness is the specific biophysical parameter important for this sensing. Traction force measurements reveal that the retraction of TFP is capable of deforming even stiff substrates. We show how slow diffusion of the pilin PilA in the inner membrane yields local concentration changes at the base of TFP during extension and retraction that change with substrate stiffness. We develop a quantitative biomechanical model that explains the transcriptional response to stiffness. A competition between PilA diffusion in the inner membrane and a loss/gain of monomers during TFP extension/retraction produces substrate stiffness-dependent dynamics of the local PilA concentration. We validated this model by manipulating the ATPase activity of the TFP motors to change TFP extension and retraction velocities and PilA concentration dynamics, altering the stiffness response in a predictable manner. Our results highlight stiffness sensing as a shared behavior across biological kingdoms, revealing generalizable principles of environmental sensing across small and large cells.

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.

Multiple strategies of improving photocatalytic water splitting efficiency in 2D arsenic sesquichalcogenides.

Improving the solar-to-hydrogen efficiency has always been a significant topic in the field of photocatalysis. Based on first-principles calculations, herein, we propose multiple strategies to improve the photocatalytic properties of 2D arsenic sesquichalcogenides for full water splitting. The new configurations As2STe2 and As2SeTe2 monolayers, derived from the As2Te3 monolayers by surface modification, are manifested to be typical infrared-light driven photocatalysts. Notably, under the built-in electric field, As2STe2 and As2SeTe2 monolayers can fulfil overall water splitting and the predicted solar-to-hydrogen efficiencies even reach up to 36.19% and 29.36%, respectively. The Gibbs free energy calculations indicate that the OER can be successfully driven under light irradiation. In addition, the overpotentials can provide most of the energy for HER when illuminated, especially for As2STe2 with the . In addition, both As2S3 and As2Se3 monolayers are capable of satisfying the conditions for photocatalytic water splitting. Furthermore, the band gaps of As2Se3 and As2S3 can dramatically be narrowed by increasing the number of layers and doping, respectively. These findings provide a theoretical basis for As2X3 monolayers to achieve efficient photocatalytic water splitting.

Tuning the mechanical impedance of disordered networks for impact mitigation.

Disordered-Network Mechanical Materials (DNMM), comprised of random arrangements of bonds and nodes, have emerged as mechanical metamaterials with the potential for achieving fine control over their mechanical properties. Recent computational studies have demonstrated this control whereby an extremely high degree of mechanical tunability can be achieved in disordered networks via a selective bond removal process called pruning. In this study, we experimentally demonstrate how pruning of a disordered network alters its macroscopic dynamic mechanical response and its capacity to mitigate impact. Impact studies with velocities ranging from 0.1 m s-1 to 1.5 m s-1 were performed, using a mechanical impactor and a drop tower, on 3D printed pruned and unpruned networks comprised of materials spanning a range of stiffness. High-speed videography was used to quantify the changes in Poisson's ratio for each of the network samples. Our results demonstrate that pruning is an efficient way to reduce the transmitted force and impulse from impact in the medium strain rate regime (101 s-1 to 102 s-1). This approach provides an interesting alternative route for designing materials with tailored impact mitigating properties compared to random material removal based on open cell foams.

Intrusion into Granular Media Beyond the Quasistatic Regime.

The drag force exerted on an object intruding into granular media is typically assumed to arise from additive velocity and depth dependent contributions. We test this with intrusion experiments and molecular dynamics simulations at constant speed over four orders of magnitude, well beyond the quasistatic regime. For a vertical cylindrical rod we find velocity dependence only right after impact, followed by a crossover to a common, purely depth-dependent behavior for all intrusion speeds. The crossover is set by the timescale for material, forced to well up at impact, to subsequently settle under gravity. These results challenge current models of granular drag.

Interparticle hydrogen bonding can elicit shear jamming in dense suspensions.

Dense suspensions of hard particles in a liquid can exhibit strikingly counter-intuitive behaviour, such as discontinuous shear thickening (DST)1-7 and reversible shear jamming (SJ) into a state where flow is arrested and the suspension is solid-like8-12. A stress-activated crossover from hydrodynamic interactions to frictional particle contacts is key for these behaviours2-4,6,7,9,13. However, in experiments, many suspensions show only DST, not SJ. Here we show that particle surface chemistry plays a central role in creating conditions that make SJ readily observable. We find the system's ability to form interparticle hydrogen bonds when sheared into contact elicits SJ. We demonstrate this with charge-stabilized polymer microspheres and non-spherical cornstarch particles, controlling hydrogen bond formation with solvents. The propensity for SJ is quantified by tensile tests12 and linked to an enhanced friction by atomic force microscopy. Our results extend the fundamental understanding of the SJ mechanism and open avenues for designing strongly non-Newtonian fluids.

Stress Controlled Rheology of Dense Suspensions Using Transient Flows.

Dense suspensions of hard particles in a Newtonian liquid can be jammed by shear when the applied stress exceeds a certain threshold. However, this jamming transition from a fluid into a solidified state cannot be probed with conventional steady-state rheology because the stress distribution inside the material cannot be controlled with sufficient precision. Here we introduce and validate a method that overcomes this obstacle. Rapidly propagating shear fronts are generated and used to establish well-controlled local stress conditions that sweep across the material. Exploiting such transient flows, we can track how a dense suspension approaches its shear-jammed state dynamically and quantitatively map out the onset stress for solidification in a state diagram.

Measuring the porosity and compressibility of liquid-suspended porous particles using ultrasound.

A key parameter describing the behavior of suspensions is the volume fraction ϕ of the solid particles that are dispersed in the liquid. Obtaining accurate values for ϕ becomes difficult for porous particles, because they can absorb some of the liquid. A prime example are the widely used cornstarch suspensions, for which ϕ usually is only estimated from the mass fraction of particles. Here we present a method to measure the effective porosity and compressibility of porous particles with ultrasound. We obtain the speed of sound in dilute cornstarch suspensions at multiple particle concentrations and with different solvent compressibilities. With the measured particle porosity of 0.31 we are able to calculate the volume fraction of the saturated particles reliably.

Local polar order controls mechanical stress and triggers layer formation in developing Myxococcus xanthus colonies.

Colonies of the social bacterium Myxococcus xanthus go through a morphological transition from a thin colony of cells to three-dimensional droplet-like fruiting bodies as a strategy to survive starvation. The biological pathways that control the decision to form a fruiting body have been studied extensively. However, the mechanical events that trigger the creation of multiple cell layers and give rise to droplet formation remain poorly understood. By measuring cell orientation, velocity, polarity, and force with cell-scale resolution, we reveal a stochastic local polar order in addition to the more obvious nematic order. Average cell velocity and active force at topological defects agree with predictions from active nematic theory, but their fluctuations are anomalously large due to polar active forces generated by the self-propelled rod-shaped cells. We find that M. xanthus cells adjust their reversal frequency to tune the magnitude of this local polar order, which in turn controls the mechanical stresses and triggers layer formation in the colonies.

Effect of two parallel intruders on total work during granular penetrations.

The intrusion of single passive intruders into granular particles has been studied in detail. However, the intrusion force produced by multiple intruders separated at a distance from one another, and hence the effect of their presence in close proximity to one another, is less explored. Here, we used numerical simulations and laboratory experiments to study the force response of two parallel rods intruding vertically into granular media while varying the gap spacing between them. We also explored the effect of variations in friction, intruder size, and particle size on the force response. The total work (W) of the two rods over the depth of intrusion was measured, and the instantaneous velocities of particles over the duration of intrusion were calculated by simulations. We found that the total work done by the intruders changes with distance between them. We observed a peak in W at a gap spacing of ∼3 particle diameters, which was up to 25% greater than W at large separation (>11 particle diameters), beyond which the total work plateaued. This peak was likely due to reduced particle flow between intruders as we found a larger number of strong forces-identified as force chains-in the particle domain at gaps surrounding the peak force. Although higher friction caused greater force generation during intrusion, the gap spacing between the intruders at which the peak total work was generated remained unchanged. Larger intruder sizes resulted in greater total work with the peak in W occurring at slightly larger intruder separations. Taken together, our results show that peak total work done by two parallel intruders remained within a narrow range, remaining robust to most other tested parameters.

Dynamic shear jamming in dense granular suspensions under extension.

Unlike dry granular materials, a dense granular suspension like cornstarch in water can strongly resist extensional flows. At low extension rates, such a suspension behaves like a viscous fluid, but rapid extension results in a response where stresses far exceed the predictions of lubrication hydrodynamics and capillarity. To understand this remarkable mechanical response, we experimentally measure the normal force imparted by a large bulk of the suspension on a plate moving vertically upward at a controlled velocity. We observe that, above a velocity threshold, the peak force increases by orders of magnitude. Using fast ultrasound imaging we map out the local velocity profiles inside the suspension, which reveal the formation of a growing jammed region under rapid extension. This region interacts with the rigid boundaries of the container through strong velocity gradients, suggesting a direct connection to the recently proposed shear-jamming mechanism.

High-speed ultrasound imaging in dense suspensions reveals impact-activated solidification due to dynamic shear jamming.

A remarkable property of dense suspensions is that they can transform from liquid-like at rest to solid-like under sudden impact. Previous work showed that this impact-induced solidification involves rapidly moving jamming fronts; however, details of this process have remained unresolved. Here we use high-speed ultrasound imaging to probe non-invasively how the interior of a dense suspension responds to impact. Measuring the speed of sound we demonstrate that the solidification proceeds without a detectable increase in packing fraction, and imaging the evolving flow field we find that the shear intensity is maximized right at the jamming front. Taken together, this provides direct experimental evidence for jamming by shear, rather than densification, as driving the transformation to solid-like behaviour. On the basis of these findings we propose a new model to explain the anisotropy in the propagation speed of the fronts and delineate the onset conditions for dynamic shear jamming in suspensions.