No Arabic abstract
We discuss the electrostatic contribution to the elastic moduli of a cell or artificial membrane placed in an electrolyte and driven by a DC electric field. The field drives ion currents across the membrane, through specific channels, pumps or natural pores. In steady state, charges accumulate in the Debye layers close to the membrane, modifying the membrane elastic moduli. We first study a model of a membrane of zero thickness, later generalizing this treatment to allow for a finite thickness and finite dielectric constant. Our results clarify and extend the results presented in [D. Lacoste, M. Cosentino Lagomarsino, and J. F. Joanny, Europhys. Lett., {bf 77}, 18006 (2007)], by providing a physical explanation for a destabilizing term proportional to $kps^3$ in the fluctuation spectrum, which we relate to a nonlinear ($E^2$) electro-kinetic effect called induced-charge electro-osmosis (ICEO). Recent studies of ICEO have focused on electrodes and polarizable particles, where an applied bulk field is perturbed by capacitive charging of the double layer and drives flow along the field axis toward surface protrusions; in contrast, we predict reverse ICEO flows around driven membranes, due to curvature-induced tangential fields within a non-equilibrium double layer, which hydrodynamically enhance protrusions. We also consider the effect of incorporating the dynamics of a spatially dependent concentration field for the ion channels.
We analyse here the problem of large deformation of dielectric elastomeric membranes under coupled electromechanical loading. Extremely large deformations (enclosed volume changes of 100 times and greater) of a toroidal membrane are studied by the use of a variational formulation that accounts for the total energy due to mechanical and electrical fields. A modified shooting method is adopted to solve the resulting system of coupled and highly nonlinear ordinary differential equations. We demonstrate the occurrence of limit point, wrinkling, and symmetry-breaking buckling instabilities in the solution of this problem. Onset of each of these reversible instabilities depends significantly on the ratio of the mechanical load to the electric load, thereby providing a control mechanism for state switching.
We present two models for electron-driven uphill proton transport across lipid membranes, with the electron energy converted to the proton gradient via the electrostatic interaction. In the first model, associated with the cytochrome c oxidase complex in the inner mitochondria membranes, the electrostatic coupling to the site occupied by an electron lowers the energy level of the proton-binding site, making the proton transfer possible. In the second model, roughly describing the redox loop in a nitrate respiration of E. coli bacteria, an electron displaces a proton from the negative side of the membrane to a shuttle, which subsequently diffuses across the membrane and unloads the proton to its positive side. We show that both models can be described by the same approach, which can be significantly simplified if the system is separated into several clusters, with strong Coulomb interaction inside each cluster and weak transfer couplings between them. We derive and solve the equations of motion for the electron and proton creation/annihilation operators, taking into account the appropriate Coulomb terms, tunnel couplings, and the interaction with the environment. For the second model, these equations of motion are solved jointly with a Langevin-type equation for the shuttle position. We obtain expressions for the electron and proton currents and determine their dependence on the electron and proton voltage build-ups, on-site charging energies, reorganization energies, temperature, and other system parameters. We show that the quantum yield in our models can be up to 100% and the power-conversion efficiency can reach 35%.
Using numerical simulations, we characterized the behavior of an elastic membrane immersed in an active fluid. Our findings reveal a nontrivial folding and re-expansion of the membrane that is controlled by the interplay of its resistance to bending and the self-propulsion strength of the active components in solution. We show how flexible membranes tend to collapse into multi-folded states, whereas stiff membranes oscillates between an extended configuration and a singly folded state. This study provides a simple example of how to exploit the random motion of active particles to perform mechanical work at the micro-scale.
We discuss the directional motion of an elastic three-sphere micromachine in which the spheres are in equilibrium with independent heat baths having different temperatures. Even in the absence of prescribed motion of springs, such a micromachine can gain net motion purely because of thermal fluctuations. A relation connecting the average velocity and the temperatures of the spheres is analytically obtained. This velocity can also be expressed in terms of the average heat flows in the steady state. Our model suggests a new mechanism for the locomotion of micromachines in nonequilibrium biological systems.
Up-down asymmetric inclusions impose a local, spontaneous curvature to an elastic membrane. When several of them are inserted in a same membrane, they feel effective forces mediated by the membrane, both of elastic and entropic nature. Following an approach initiated by Dommersnes and Fournier in the vanishing tension case [Eur. Phys. J. B 12, 9 (1999)], and also using a pseudo-analytical micellization theory, we derive the statistical mechanics of asymmetric inclusion assemblies when they are also subject to an additional short-range, attractive interaction. Our main conclusion is that generically, when the membrane is under tension, these inclusions live in small clusters at equilibrium, leading to local membrane invaginations. We also propose a novel curvature-induced demixing mechanism: when inclusions imposing local curvatures of opposite sign coexist, they tend to demix in distinct clusters under realistic conditions. This work has potential implications in the context of the thermodynamics of proteins embedded in biological lipid bilayers.