Jean-François Joanny

A model for contact angle hysteresis

We discuss the behavior of a liquid partially wetting a solid surface, when the contact angle at equilibrium θ0 is small, but finite. The solid is assumed to be either flat, but chemically heterogeneous (this in turn modulating the interfacial tensions), or rough. For weak heterogeneities, we expect no hysteresis, but the contact line becomes wiggly. For stronger heterogeneities, we first discuss the behavior of the contact line in the presence of a single, localized defect, and show that there may exist two stable positions for the line, obtained by a simple graphic construction. Hysteresis shows up when the strength of the defect is above a certain threshold. Extending this to a dilute system of defects, we obtain formulas for the ‘‘advancing’’ and ‘‘receding’’ contact angles θa, θr, in terms of the distribution of defect strength and defect sharpness. These formulas might be tested by controlled contamination of a solid surface.

Fast DNA Translocation through a Solid-State Nanopore

We report experiments and modeling of translocation of double-strand DNA through a siliconoxide nanopore. Long DNA molecules with different lengths ranging from 6500 to 97000 base pairs have been electrophoretically driven through a 10 nm pore. We observe a power-law caling of the translocation time with the length, with an exponent of 1.27. This nonlinear scaling is strikingly different from the well-studied linear behavior observed in similar experiments performed on protein pores. We present a theoretical model where hydrodynamic drag on the ection of the polymer outside the pore is the dominant force counteracting the electrical driving force. We show that this applies to our experiments, and we derive a power-law scaling with an exponent of 1.22, in good agreement with the data.

Role of cortical tension in bleb growth

Blebs are spherical membrane protrusions often observed during cell migration, cell spreading, cytokinesis, and apoptosis, both in cultured cells and in vivo. Bleb expansion is thought to be driven by the contractile actomyosin cortex, which generates hydrostatic pressure in the cytoplasm and can thus drive herniations of the plasma membrane. However, the role of cortical tension in bleb formation has not been directly tested, and despite the importance of blebbing, little is known about the mechanisms of bleb growth. In order to explore the link between cortical tension and bleb expansion, we induced bleb formation on cells with different tensions. Blebs were nucleated in a controlled manner by laser ablation of the cortex, mimicking endogenous bleb nucleation. Cortical tension was modified by treatments affecting the level of myosin activity or proteins regulating actin turnover. We show that there is a critical tension below which blebs cannot expand. Above this threshold, the maximal size of a bleb strongly depends on tension, and this dependence can be fitted with a model of the cortex as an active elastic material. Together, our observations and model allow us to relate bleb shape parameters to the underlying cellular mechanics and provide insights as to how bleb formation can be biochemically regulated during cell motility.

Curvature-driven lipid sorting needs proximity to a demixing point and is aided by proteins

Sorting of lipids and proteins is a key process allowing eukaryotic cells to execute efficient and accurate intracellular transport and to maintain membrane homeostasis. It occurs during the formation of highly curved transport intermediates that shuttle between cell compartments. Protein sorting is reasonably well described, but lipid sorting is much less understood. Lipid sorting has been proposed to be mediated by a physical mechanism based on the coupling between membrane composition and high curvature of the transport intermediates. To test this hypothesis, we have performed a combination of fluorescence and force measurements on membrane tubes of controlled diameters pulled from giant unilamellar vesicles. A model based on membrane elasticity and nonideal solution theory has also been developed to explain our results. We quantitatively show, using 2 independent approaches, that a difference in lipid composition can build up between a curved and a noncurved membrane. Importantly, and consistent with our theory, lipid sorting occurs only if the system is close to a demixing point. Remarkably, this process is amplified when even a low fraction of lipids is clustered upon cholera toxin binding. This can be explained by the reduction of the entropic penalty of lipid sorting when some lipids are bound together by the toxin. Our results show that curvature-induced lipid sorting results from the collective behavior of lipids and is even amplified in the presence of lipid-clustering proteins. In addition, they suggest a generic mechanism by which proteins can facilitate lipid segregation in vivo.

Generic theory of active polar gels: a paradigm for cytoskeletal dynamics

Abstract.We develop a general theory for active viscoelastic materials made of polar filaments. This theory is motivated by the dynamics of the cytoskeleton. The continuous consumption of a fuel leads to a non equilibrium state characterized by the generation of flows and stresses. Our theory can be applied to experiments in which cytoskeletal patterns are set in motion by active processes such as those which are at work in cells.

Phase transitions in Langmuir monolayers of polar molecules

Insoluble Langmuir monolayers are investigated in the presence of dipolar forces which can have two origins: permanent dipoles in neutral monolayers and induced dipoles in charged monolayers. The main effect of the additional long‐range repulsive interactions is to stabilize undulating phases at thermodynamic equilibrium. Phase diagrams are obtained in two limits: close to the liquid–gas critical point via a Ginzburg–Landau expansion of the free energy (mainly within a mean‐field approximation), and at low temperatures by free energy minimization. Possible applications of this theory to experiments at the liquid–gas, liquid expanded–liquid condensed, and solid–liquid transitions are discussed.

Fluidization of tissues by cell division and apoptosis

During the formation of tissues, cells organize collectively by cell division and apoptosis. The multicellular dynamics of such systems is influenced by mechanical conditions and can give rise to cell rearrangements and movements. We develop a continuum description of tissue dynamics, which describes the stress distribution and the cell flow field on large scales. In the absence of division and apoptosis, we consider the tissue to behave as an elastic solid. Cell division and apoptosis introduce stress sources that, in general, are anisotropic. By combining cell number balance with dynamic equations for the stress source, we show that the tissue effectively behaves as a viscoelastic fluid with a relaxation time set by the rates of division and apoptosis. If the system is confined in a fixed volume, it reaches a homeostatic state in which division and apoptosis balance. In this state, cells undergo a diffusive random motion driven by the stochasticity of division and apoptosis. We calculate the expression for the effective diffusion coefficient as a function of the tissue parameters and compare our results concerning both diffusion and viscosity to simulations of multicellular systems using dissipative particle dynamics.

ATP-dependent mechanics of red blood cells

Red blood cells are amazingly deformable structures able to recover their initial shape even after large deformations as when passing through tight blood capillaries. The reason for this exceptional property is found in the composition of the membrane and the membrane-cytoskeleton interaction. We investigate the mechanics and the dynamics of RBCs by a unique noninvasive technique, using weak optical tweezers to measure membrane fluctuation amplitudes with μs temporal and sub nm spatial resolution. This enhanced edge detection method allows to span over >4 orders of magnitude in frequency. Hence, we can simultaneously measure red blood cell membrane mechanical properties such as bending modulus κ = 2.8 ± 0.3 × 10−19J = 67.6 ± 7.2 kBT, tension σ = 6.5 ± 2.1 × 10−7N/m, and an effective viscosity ηeff = 81 ± 3.7 × 10−3 Pa s that suggests unknown dissipative processes. We furthermore show that cell mechanics highly depends on the membrane-spectrin interaction mediated by the phosphorylation of the interconnection protein 4.1R. Inhibition and activation of this phosphorylation significantly affects tension and effective viscosity. Our results show that on short time scales (slower than 100 ms) the membrane fluctuates as in thermodynamic equilibrium. At time scales longer than 100 ms, the equilibrium description breaks down and fluctuation amplitudes are higher by 40% than predicted by the membrane equilibrium theory. Possible explanations for this discrepancy are influences of the spectrin that is not included in the membrane theory or nonequilibrium fluctuations that can be accounted for by defining a nonthermal effective energy of up to Eeff = 1.4 ± 0.1 kBT, that corresponds to an actively increased effective temperature.

Spontaneous flow transition in active polar gels

We study theoretically the effects of confinement on active polar gels such as the actin network of eukaryotic cells. Using generalized hydrodynamics equations derived for active gels, we predict, in the case of quasi–one-dimensional geometry, a spontaneous flow transition from a homogeneously polarized immobile state for small thicknesses, to a perturbed flowing state for larger thicknesses. The transition is not driven by an external field but by the activity of the system. We suggest several possible experimental realizations.

Contractility and retrograde flow in lamellipodium motion

We present a phenomenological description of cell locomotion on a solid substrate. The material properties of the actin cytoskeleton in the lamellipodium are described by the constitutive equations of a viscous polar gel with intrinsic activity. The polymerization of the gel takes place in a localized region near the leading edge. Using a simple two-dimensional description, we calculate in the steady state the thickness profile of the lamellipodium which at the rear connects to the cell body; we also calculate the flow profiles and the forces exerted on the substrate. The cell velocity is estimated as a function of externally applied forces. Our description is consistent with experimentally observed properties of motile cells such as the existence of a retrograde flow in the lamellipodium and a dipolar force distribution exerted by the cell on the substrate.