Mechanics of bundled semiflexible polymer networks

While actin bundles are used by living cells for structural fortification, the microscopic origin of the elasticity of bundled networks is not understood. Here, we show that above a critical concentration of the actin binding protein fascin, a solution of actin filaments organizes into a pure network of bundles. While the elasticity of weakly cross-linked networks is dominated by the affine deformation of tubes, the network of bundles can be fully understood in terms of nonaffine bending undulations.     

Deformation of cross-linked semiflexible polymer networks

Networks of filamentous proteins play a crucial role in cell mechanics. These cytoskeletal networks, together with various cross-linking and other associated proteins largely determine the (visco)elastic response of cells. In this Letter we study a model system of cross-linked, stiff filaments in order to explore the connection between the microstructure under strain and the macroscopic response of cytoskeletal networks. We find two distinct regimes as a function primarily of cross-link density and filament rigidity: one characterized by affine deformation and one by nonaffine deformation. We characterize the crossover between these two.     

Filamin cross-linked semiflexible networks: fragility under strain

The semiflexible F-actin network of the cytoskeleton is cross-linked by a variety of proteins including filamin, which contains Ig domains that unfold under applied tension. We examine a simple filament network model cross-linked by such unfolding linkers that captures the main mechanical features of F-actin networks cross-linked by filamin proteins and show that, under sufficient strain, the network spontaneously self-organizes so that an appreciable fraction of the filamin cross-linkers are at the threshold of domain unfolding. We propose and test a mean-field model to account for this effect. We also suggest a qualitative experimental signature of this type of network reorganization under applied strain that may be observable in intracellular microrheology experiments of Crocker et al.     

High-frequency stress relaxation in semiflexible polymer solutions and networks

We measure the linear viscoelasticity of sterically entangled and chemically cross-linked networks of actin filaments over more than five decades of frequency. The high-frequency response reveals rich dynamics unique to semiflexible polymers, including a previously unobserved relaxation due to rapid axial tension propagation. For high molecular weight, and for cross-linked gels, we obtain quantitative agreement with predicted shear moduli in both amplitude and frequency dependence.     

Dynamics of prestressed semiflexible polymer chains as a model of cell rheology

We report on a model of a prestressed nonlinear semiflexible polymer chain that links thermally driven dynamics to the creep behavior of living cells. Numerical simulations show that the chain's creep follows a power law with an exponent that decreases with increasing prestress. This is related to the propagation of free energy through the chain in response to stretching, where the propagation speed is regulated by the prestress via the chain's nonlinear elasticity. These results indicate that the main aspects of cell rheology are consistent with the dynamics of single polymer chains under tension.     

Surface rheology of two-dimensional percolating networks: Langmuir films of polymer pancakes

We report surface rheological measurements on Langmuir films of submicron-sized polymer coils (pancakes). The dynamics of the sol phase is found to be governed by free volume as in the simplest percolation problem. The observed rheology of the percolated phase (gel-like) is compatible with predictions for compressed arrangements of highly deformed soft particles interacting through their contact interfaces.     

The Bohm-Vigier subquantum fluctuations and nonlinear field theory

Utilising as starting point the double solution theory of Louis de Broglie and the Bohm-Vigier hypothesis of subquantum fluctuations, an attempt is made to give an example of the nonlinear field theory in which the guidance theorem of de Broglie can be realised. The simplest scalar model is considered.     

Elongation and fluctuations of semiflexible polymers in a nematic solvent

We directly visualize single polymers with persistence lengths l(p), ranging from 0.05 to 16 microm, dissolved in the nematic phase of rodlike fd virus. Polymers with a sufficiently large persistence length undergo a coil-rod transition at the isotropic-nematic transition of the background solvent. We quantitatively analyze the transverse fluctuations of the semiflexible polymers and show that at long wavelengths they are driven by the fluctuating nematic background. We extract the Odijk deflection length and the elastic constant of the background nematic phase from the data.     

Alternative explanation of stiffening in cross-linked semiflexible networks

Strain stiffening of filamentous protein networks is explored by means of a finite strain analysis of a two-dimensional network model of cross-linked semiflexible filaments. The results show that stiffening is caused by nonaffine network rearrangements that govern a transition from a bending-dominated response at small strains to a stretching-dominated response at large strains. Filament undulations, which are key in the existing explanation of stiffening, merely postpone the transition.     

The interplay of synchronization and fluctuations reveals connectivity levels in networks of nonlinear oscillators

We study spatiotemporal patterns produced by small-world networks of biologically motivated nonlinear oscillators from a data-analysis perspective. It is shown that the connectivity levels of such systems can be reconstructed by analyzing heterogeneity and fluctuation content of the patterns. These properties are determined by applying spatiotemporal filters described in [Physica A 289 (2001) 498] to pairs of oscillators in a network. Possible applications of our method to biological data (e.g., time-resolved cDNA microarray data), in order to distinguish densely connected systems from sparsely connected systems, are commented on.     

Quasistatic rheology and the origins of strain

Features of rheological laws applied to solid-like granular materials are recalled and confronted to microscopic approaches via discrete numerical simulations. We give examples of model systems with very similar equilibrium stress transport properties—the much-studied force chains and force distribution—but qualitatively different strain responses to stress increments. Results on the stability of elastoplastic contact networks lead to the definition of two different rheological regimes, according to whether a macroscopic fragility property (propensity to rearrange under arbitrary small stress increments in the thermodynamic limit) applies. Possible consequences are discussed. To cite this article: J.-N. Roux, G. Combe, C. R. Physique 3 (2002) 131–140.     

Universal elasticity and fluctuations of nematic gels

We study elasticity of spontaneously orientationally ordered amorphous solids, characterized by a vanishing transverse shear modulus, as realized by nematic elastomers and gels. We show that local heterogeneities and elastic nonlinearities conspire to lead to anomalous nonlocal universal elasticity controlled by a nontrivial infrared fixed point. Namely, such solids are characterized by universal shear and bending moduli that, respectively, vanish and diverge at long scales, are universally incompressible, and exhibit a universal negative Poisson ratio and a non-Hookean elasticity down to arbitrarily low strains. Based on expansion about five dimensions, we argue that the nematic order is stable to thermal fluctuation and local heterogeneities down to d(lc)<3.     

Thermal and non-thermal fluctuations in active polar gels

We discuss general features of noise and fluctuations in active polar gels close to and away from equilibrium. We use the single-component hydrodynamic theory of active polar gels built by Kruse and coworkers to describe the cytoskeleton in cells. Close to equilibrium, we calculate the response function of the gel to external fields and introduce Langevin forces in the constitutive equations with correlation functions respecting the fluctuation-dissipation theorem. We then discuss the breakage of the fluctuation-dissipation theorem due to an external field such as the activity of the motors. Active gels away from equilibrium are considered at the scaling level. As an example of application of the theory, we calculate the density correlation function (the dynamic structure factor) of a compressible active polar gel and discuss possible instabilities.     

Generalized nonlinear master equations for local fluctuations: Dimensionality expansions and augmented mean field theory

By application of a projection operator technique we derive a formally exact generalization of the nonlinear mean field master equation introduced recently for the study of local fluctuations in a reacting medium. Our starting point is a phenomenological cell master equation. The results of our theory are applicable to the theory of a fluctuating hydrodynamic reacting system. The mean field equation is placed on a firm theoretical foundation by showing it to be the lowest order approximation in an expansion in the dimensionality of the physical space keeping the product of the number of nearest neighbors (an increasing function of dimensionality) and the typical diffusion coefficient constant. A more accurate nonlinear master equation that allows for the correlation and fluctuations in the environment of a given volume element is derived in the form of an augmented mean field equation.Work supported in part by a grant from the National Science Foundation.     

Scaling of F-actin network rheology to probe single filament elasticity and dynamics

The linear and nonlinear viscoelastic response of networks of cross-linked and bundled cytoskeletal filaments demonstrates remarkable scaling with both frequency and applied prestress, which helps elucidate the origins of the viscoelasticity. The frequency dependence of the shear modulus reflects the underlying single-filament relaxation dynamics for 0.1-10 rad/sec. Moreover, the nonlinear strain stiffening of such networks exhibits a universal form as a function of prestress; this is quantitatively explained by the full force-extension relation of single semiflexible filaments.     

Linear and nonlinear rheology of wormlike micelles

Several surfactant molecules self-assemble in solution to form long, cylindrical, flexible wormlike micelles. These micelles can be entangled with each other leading to viscoelastic phases. The rheological properties of such phases are very interesting and have been the subject of a large number of experimental and theoretical studies in recent years. We shall report our recent work on the macrorheology, microrheology and nonlinear flow behaviour of dilute aqueous solutions of a surfactant CTAT (Cetyltrimethylammonium Tosilate). This system forms elongated micelles and exhibits strong viscoelasticity at low concentrations (∼0.9 wt%) without the addition of electrolytes. Microrheology measurements of G(θ) have been done using diffusing wave spectroscopy which will be compared with the conventional frequency sweep measurements done using a cone and plate rheometer. The second part of the paper deals with the nonlinear rheology where the measured shear stress σ is a nonmonotonic function of the shear rate . In stress-controlled experiments, the shear stress shows a plateau for larger than some critical strain rate, similar to the earlier reports on CPyCl/NaSal system. Cates et al have proposed that the plateau is a signature of mechanical instability in the form of shear bands. We have carried out extensive experiments under controlled strain rate conditions, to study the time-dependence of shear stress. The measured time series of shear stress has been analysed in terms of correlation integral and Lyapunov exponent to show unambiguously that the behaviour is typical of low dimensional dynamical systems.     

半刚性高分子链螺旋结构诱导纳米棒的有序结构

在自然界中, 螺旋结构广泛存在. 在熵的驱动下, 高分子链能在某些特殊情形下形成螺旋结构. 采用分子动力学方法研究了高分子链诱导纳米棒的自组装行为, 发现纳米棒/高分子链体系的构象与纳米棒的数量、高分子链的刚性等密切相关. 当纳米棒与高分子链之间存在适度吸附能时, 纳米棒能够形成三种完全不同的构象, 特别是在半刚性高分子链诱导下纳米棒能够形成线型排列. 研究结果对新型材料制备具有一定指导意义.