Second variation of the Helfrich-Canham Hamiltonian and reparametrization invariance

A covariant approach towards a theory of deformations is developed to examine both the first and second variation of the Helfrich-Canham Hamiltonian -- quadratic in the extrinsic curvature -- which describes fluid vesicles at mesoscopic scales. Deformations are decomposed into tangential and normal components; At first order, tangential deformations may always be identified with a reparametrization; at second order, they differ. The relationship between tangential deformations and reparametrizations, as well as the coupling between tangential and normal deformations, is examined at this order for both the metric and the extrinsic curvature tensors. Expressions for the expansion to second order in deformations of geometrical invariants constructed with these tensors are obtained; in particular, the expansion of the Hamiltonian to this order about an equilibrium is considered. Our approach applies as well to any geometrical model for membranes.Comment: 20 page

How to determine local elastic properties of lipid bilayer membranes from atomic-force-microscope measurements: A theoretical analysis

Measurements with an atomic force microscope (AFM) offer a direct way to probe elastic properties of lipid bilayer membranes locally: provided the underlying stress-strain relation is known, material parameters such as surface tension or bending rigidity may be deduced. In a recent experiment a pore-spanning membrane was poked with an AFM tip, yielding a linear behavior of the force-indentation curves. A theoretical model for this case is presented here which describes these curves in the framework of Helfrich theory. The linear behavior of the measurements is reproduced if one neglects the influence of adhesion between tip and membrane. Including it via an adhesion balance changes the situation significantly: force-distance curves cease to be linear, hysteresis and nonzero detachment forces can show up. The characteristics of this rich scenario are discussed in detail in this article.Comment: 14 pages, 9 figures, REVTeX4 style. New version corresponds to the one accepted by PRE. The result section is restructured: a comparison to experimental findings is included; the discussion on the influence of adhesion between AFM tip and membrane is extende

Stresses in lipid membranes

The stresses in a closed lipid membrane described by the Helfrich hamiltonian, quadratic in the extrinsic curvature, are identified using Noether's theorem. Three equations describe the conservation of the stress tensor: the normal projection is identified as the shape equation describing equilibrium configurations; the tangential projections are consistency conditions on the stresses which capture the fluid character of such membranes. The corresponding torque tensor is also identified. The use of the stress tensor as a basis for perturbation theory is discussed. The conservation laws are cast in terms of the forces and torques on closed curves. As an application, the first integral of the shape equation for axially symmetric configurations is derived by examining the forces which are balanced along circles of constant latitude.Comment: 16 pages, introduction rewritten, other minor changes, new references added, version to appear in Journal of Physics

Two-colour generation in a chirped seeded Free-Electron Laser

We present the experimental demonstration of a method for generating two spectrally and temporally separated pulses by an externally seeded, single-pass free-electron laser operating in the extreme-ultraviolet spectral range. Our results, collected on the FERMI@Elettra facility and confirmed by numerical simulations, demonstrate the possibility of controlling both the spectral and temporal features of the generated pulses. A free-electron laser operated in this mode becomes a suitable light source for jitter-free, two-colour pump-probe experiments

The prolate-to-oblate shape transition of phospholipid vesicles in response to frequency variation of an AC electric field can be explained by the dielectric anisotropy of a phospholipid bilayer

The external electric field deforms flaccid phospholipid vesicles into spheroidal bodies, with the rotational axis aligned with its direction. Deformation is frequency dependent: in the low frequency range (~ 1 kHz), the deformation is typically prolate, while increasing the frequency to the 10 kHz range changes the deformation to oblate. We attempt to explain this behaviour with a theoretical model, based on the minimization of the total free energy of the vesicle. The energy terms taken into account include the membrane bending energy and the energy of the electric field. The latter is calculated from the electric field via the Maxwell stress tensor, where the membrane is modelled as anisotropic lossy dielectric. Vesicle deformation in response to varying frequency is calculated numerically. Using a series expansion, we also derive a simplified expression for the deformation, which retains the frequency dependence of the exact expression and may provide a better substitute for the series expansion used by Winterhalter and Helfrich, which was found to be valid only in the limit of low frequencies. The model with the anisotropic membrane permittivity imposes two constraints on the values of material constants: tangential component of dielectric permittivity tensor of the phospholipid membrane must exceed its radial component by approximately a factor of 3; and the membrane conductivity has to be relatively high, approximately one tenth of the conductivity of the external aqueous medium.Comment: 17 pages, 6 figures; accepted for publication in J. Phys.: Condens. Matte

Hamilton's equations for a fluid membrane: axial symmetry

Consider a homogenous fluid membrane, or vesicle, described by the Helfrich-Canham energy, quadratic in the mean curvature. When the membrane is axially symmetric, this energy can be viewed as an `action' describing the motion of a particle; the contours of equilibrium geometries are identified with particle trajectories. A novel Hamiltonian formulation of the problem is presented which exhibits the following two features: {\it (i)} the second derivatives appearing in the action through the mean curvature are accommodated in a natural phase space; {\it (ii)} the intrinsic freedom associated with the choice of evolution parameter along the contour is preserved. As a result, the phase space involves momenta conjugate not only to the particle position but also to its velocity, and there are constraints on the phase space variables. This formulation provides the groundwork for a field theoretical generalization to arbitrary configurations, with the particle replaced by a loop in space.Comment: 11 page

Efficient and accurate simulations of deformable particles immersed in a fluid using a combined immersed boundary lattice Boltzmann finite element method

The deformation of an initially spherical capsule, freely suspended in simple shear flow, can be computed analytically in the limit of small deformations [D. Barthes-Biesel, J. M. Rallison, The Time-Dependent Deformation of a Capsule Freely Suspended in a Linear Shear Flow, J. Fluid Mech. 113 (1981) 251-267]. Those analytic approximations are used to study the influence of the mesh tessellation method, the spatial resolution, and the discrete delta function of the immersed boundary method on the numerical results obtained by a coupled immersed boundary lattice Boltzmann finite element method. For the description of the capsule membrane, a finite element method and the Skalak constitutive model [R. Skalak et al., Strain Energy Function of Red Blood Cell Membranes, Biophys. J. 13 (1973) 245-264] have been employed. Our primary goal is the investigation of the presented model for small resolutions to provide a sound basis for efficient but accurate simulations of multiple deformable particles immersed in a fluid. We come to the conclusion that details of the membrane mesh, as tessellation method and resolution, play only a minor role. The hydrodynamic resolution, i.e., the width of the discrete delta function, can significantly influence the accuracy of the simulations. The discretization of the delta function introduces an artificial length scale, which effectively changes the radius and the deformability of the capsule. We discuss possibilities of reducing the computing time of simulations of deformable objects immersed in a fluid while maintaining high accuracy.Comment: 23 pages, 14 figures, 3 table