Rostislav Boltyanskiy

Membrane Tension Maintains Cell Polarity by Confining Signals to the Leading Edge during Neutrophil Migration

Little is known about how neutrophils and other cells establish a single zone of actin assembly during migration. A widespread assumption is that the leading edge prevents formation of additional fronts by generating long-range diffusible inhibitors or by sequestering essential polarity components. We use morphological perturbations, cell-severing experiments, and computational simulations to show that diffusion-based mechanisms are not sufficient for long-range inhibition by the pseudopod. Instead, plasma membrane tension could serve as a long-range inhibitor in neutrophils. We find that membrane tension doubles during leading-edge protrusion, and increasing tension is sufficient for long-range inhibition of actin assembly and Rac activation. Furthermore, reducing membrane tension causes uniform actin assembly. We suggest that tension, rather than diffusible molecules generated or sequestered at the leading edge, is the dominant source of long-range inhibition that constrains the spread of the existing front and prevents the formation of secondary fronts.

Universal deformation of soft substrates near a contact line and the direct measurement of solid surface stresses

Droplets deform soft substrates near their contact lines. Using confocal microscopy, we measure the deformation of silicone gel substrates due to glycerol and fluorinated-oil droplets for a range of droplet radii and substrate thicknesses. For all droplets, the substrate deformation takes a universal shape close to the contact line that depends on liquid composition, but is independent of droplet size and substrate thickness. This shape is determined by a balance of interfacial tensions at the contact line and provides a novel method for direct determination of the surface stresses of soft substrates. Moreover, we measure the change in contact angle with droplet radius and show that Youngs law fails for small droplets when their radii approach an elastocapillary length scale. For larger droplets the macroscopic contact angle is constant, consistent with Youngs law.

Surface tension and contact with soft elastic solids

The Johnson-Kendall-Roberts theory is the basis of modern contact mechanics. It describes how two deformable objects adhere together, driven by adhesion energy and opposed by elasticity. Here we characterize the indentation of glass particles into soft, silicone substrates using confocal microscopy. We show that, whereas the Johnson-Kendall-Roberts theory holds for particles larger than a critical, elastocapillary lengthscale, it fails for smaller particles. Instead, adhesion of small particles mimics the adsorption of particles at a fluid interface, with a size-independent contact angle between the undeformed surface and the particle given by a generalized version of the Youngs law. A simple theory quantitatively captures this behaviour and explains how solid surface tension dominates elasticity for small-scale indentation of soft materials.

Stiffening solids with liquid inclusions

Solids embedded with fluid inclusions are intuitively softer than their pure counterparts. But experiments show that when the droplets are small enough, material can become stiffer—highlighting a role for surface tension.

Wetting and Phase Separation in Soft Adhesion

Significance Modern contact mechanics was originally developed to describe adhesion to relatively stiff materials like rubber, but much softer sticky materials are ubiquitous in biology, medicine, engineering, and everyday consumer products. By studying adhesive contact between compliant gels and rigid objects, we demonstrate that soft materials adhere very differently than their stiffer counterparts. We find that the structure in the region of contact is governed by the same physics that sets the geometry of liquid droplets, even though the material is solid. Furthermore, adhesion can cause the local composition of a soft material to change, thus coupling to its thermodynamic properties. These findings may substantially change our understanding of the mechanics of soft contact. In the classic theory of solid adhesion, surface energy drives deformation to increase contact area whereas bulk elasticity opposes it. Recently, solid surface stress has been shown also to play an important role in opposing deformation of soft materials. This suggests that the contact line in soft adhesion should mimic that of a liquid droplet, with a contact angle determined by surface tensions. Consistent with this hypothesis, we observe a contact angle of a soft silicone substrate on rigid silica spheres that depends on the surface functionalization but not the sphere size. However, to satisfy this wetting condition without a divergent elastic stress, the gel phase separates from its solvent near the contact line. This creates a four-phase contact zone with two additional contact lines hidden below the surface of the substrate. Whereas the geometries of these contact lines are independent of the size of the sphere, the volume of the phase-separated region is not, but rather depends on the indentation volume. These results indicate that theories of adhesion of soft gels need to account for both the compressibility of the gel network and a nonzero surface stress between the gel and its solvent.

Direct measurement of strain-dependent solid surface stress

Surface stress, also known as surface tension, is a fundamental material property of any interface. However, measurements of solid surface stress in traditional engineering materials, such as metals and oxides, have proven to be very challenging. Consequently, our understanding relies heavily on untested theories, especially regarding the strain dependence of this property. Here, we take advantage of the high compliance and large elastic deformability of a soft polymer gel to directly measure solid surface stress as a function of strain. As anticipated by theoretical work for metals, we find that the surface stress depends on the strain via a surface modulus. Remarkably, the surface modulus of our soft gels is many times larger than the zero-strain surface tension. This suggests that surface stresses can play a dominant role in solid mechanics at larger length scales than previously anticipated.Solid surface stress is a fundamental property of solid interfaces. Here authors measure the solid surface stress of a gel, and show its dependence on surface strain through a surface modulus.

Erratum: Stiffening solids with liquid inclusions

Nature Physics 11, 82–87 (2015); published online 15 December 2014; corrected after print 8 January 2015. In the text following equation 7, the expression describing the limit where stiffening occurs was incorrect and should have read: surface tension dominates over elasticity (R ≪ ϒ/E). This has now been corrected in the online versions of the Article.