Effective medium approach for stiff polymer networks with flexible cross-links

Recent experiments have demonstrated that the nonlinear elasticity of in vitro networks of the biopolymer actin is dramatically altered in the presence of a flexible cross-linker such as the abundant cytoskeletal protein filamin. The basic principles of such networks remain poorly understood. Here we describe an effective medium theory of flexibly cross-linked stiff polymer networks. We argue that the response of the cross-links can be fully attributed to entropic stiffening, while softening due to domain unfolding can be ignored. The network is modeled as a collection of randomly oriented rods connected by flexible cross-links to an elastic continuum. This effective medium is treated in a linear elastic limit as well as in a more general framework, in which the medium self-consistently represents the nonlinear network behavior. This model predicts that the nonlinear elastic response sets in at strains proportional to cross-linker length and inversely proportional to filament length. Furthermore, we find that the differential modulus scales linearly with the stress in the stiffening regime. These results are in excellent agreement with bulk rheology data.Comment: 12 pages, 8 figure

Instability and front propagation in laser-tweezed lipid bilayer tubules

We study the mechanism of the `pearling' instability seen recently in experiments on lipid tubules under a local applied laser intensity. We argue that the correct boundary conditions are fixed chemical potentials, or surface tensions \Sigma, at the laser spot and the reservoir in contact with the tubule. We support this with a microscopic picture which includes the intensity profile of the laser beam, and show how this leads to a steady-state flow of lipid along the surface and gradients in the local lipid concentration and surface tension (or chemical potential). This leads to a natural explanation for front propagation and makes several predictions based on the tubule length. While most of the qualitative conclusions of previous studies remain the same, the `ramped' control parameter (surface tension) implies several new qualitative results. We also explore some of the consequences of front propagation into a noisy (due to pre-existing thermal fluctuations) unstable medium.Comment: 12 page latex + figures using epsf.sty to be published in Journal de Physique II, January 199

Breakup coupling effects on near-barrier ⁶Li, ⁷Be and ⁸B + ⁵⁸Ni elastic scattering compared

New data for near-barrier 6Li, 7Be and 8B + 58Ni elastic scattering enable a comparison of breakup coupling effects for these loosely-bound projectiles. Coupled Discretised Continuum Channels (CDCC) calculations suggest that the large total reaction cross sections for 8B + 58Ni are dominated by breakup at near-barrier energies, unlike 6Li and 7Be where breakup makes a small contribution. In spite of this, the CDCC calculations show a small coupling influence due to breakup for 8B, in contrast to the situation for 6Li and 7Be. An examination of the S matrices gives a clue to this counter-intuitive behaviour

Determining Microscopic Viscoelasticity in Flexible and Semiflexible Polymer Networks from Thermal Fluctuations

We have developed a new technique to measure viscoelasticity in soft materials such as polymer solutions, by monitoring thermal fluctuations of embedded probe particles using laser interferometry in a microscope. Interferometry allows us to obtain power spectra of fluctuating beads from 0.1 Hz to 20 kHz, and with sub-nanometer spatial resolution. Using linear response theory, we determined the frequency-dependent loss and storage shear moduli up to frequencies on the order of a kHz. Our technique measures local values of the viscoelastic response, without actively straining the system, and is especially suited to soft biopolymer networks. We studied semiflexible F-actin solutions and, as a control, flexible polyacrylamide (PAAm) gels, the latter close to their gelation threshold. With small particles, we could probe the transition from macroscopic viscoelasticity to more complex microscopic dynamics. In the macroscopic limit we find shear moduli at 0.1 Hz of G'=0.11 +/- 0.03 Pa and 0.17 +/- 0.07 Pa for 1 and 2 mg/ml actin solutions, close to the onset of the elastic plateau, and scaling behavior consistent with G(omega) as omega^(3/4) at higher frequencies. For polyacrylamide we measured plateau moduli of 2.0, 24, 100 and 280 Pa for crosslinked gels of 2, 2.5, 3 and 5% concentration (weight/volume) respectively, in agreement to within a factor of two with values obtained from conventional rheology. We also found evidence for scaling of G(omega) as \omega^(1/2), consistent with the predictions of the Rouse model for flexible polymers.Comment: 16 pages, with 15 PostScript figures (to be published in Macromolecules

Origin of slow stress relaxation in the cytoskeleton

Dynamically crosslinked semiflexible biopolymers such as the actin cytoskeleton govern the mechanical behavior of living cells. Semiflexible biopolymers nonlinearly stiffen in response to mechanical loads, whereas the crosslinker dynamics allow for stress relaxation over time. Here we show, through rheology and theoretical modeling, that the combined nonlinearity in time and stress leads to an unexpectedly slow stress relaxation, similar to the dynamics of disordered systems close to the glass transition. Our work suggests that transient crosslinking combined with internal stress can explain prior reports of soft glassy rheology of cells, in which the shear modulus increases weakly with frequency.Comment: 6 pages, 4 figure

Normal stress anisotropy and marginal stability in athermal elastic networks

Hydrogels of semiflexible biopolymers such as collagen have been shown to contract axially under shear strain, in contrast to the axial dilation observed for most elastic materials. Recent work has shown that this behavior can be understood in terms of the porous, two-component nature and consequent time-dependent compressibility of hydrogels. The apparent normal stress measured by a torsional rheometer reflects only the tensile contribution of the axial component $\sigma_{zz}$ on long (compressible) timescales, crossing over to the first normal stress difference, $N_1 = \sigma_{xx}-\sigma_{zz}$ at short (incompressible) times. While the behavior of $N_1$ is well understood for isotropic viscoelastic materials undergoing affine shear deformation, biopolymer networks are often anisotropic and deform nonaffinely. Here, we numerically study the normal stresses that arise under shear in subisostatic, athermal semiflexible polymer networks. We show that such systems exhibit strong deviations from affine behavior and that these anomalies are controlled by a rigidity transition as a function of strain

Budding and Domain Shape Transformations in Mixed Lipid Films and Bilayer Membranes

We study the stability and shapes of domains with spontaneous curvature in fluid films and membranes, embedded in a surrounding membrane with zero spontaneous curvature. These domains can result from the inclusion of an impurity in a fluid membrane, or from phase separation within the membrane. We show that for small but finite line and surface tensions and for finite spontaneous curvatures, an equilibrium phase of protruding circular domains is obtained at low impurity concentrations. At higher concentrations, we predict a transition from circular domains, or "caplets", to stripes. In both cases, we calculate the shapes of these domains within the Monge representation for the membrane shape. With increasing line tension, we show numerically that there is a budding transformation from stable protruding circular domains to spherical buds. We calculate the full phase diagram, and demonstrate a two triple points, of respectively bud-flat-caplet and flat-stripe-caplet coexistence.Comment: 14 pages, to appear in Phys Rev