Deformation of crosslinked semiflexible polymer networks

Networks of filamentous proteins play a crucial role in cell mechanics. These cytoskeletal networks, together with various crosslinking and other associated proteins largely determine the (visco)elastic response of cells. In this letter we study a model system of crosslinked, 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 crosslink density and filament rigidity: one characterized by affine deformation and one by non-affine deformation. We characterize the crossover between these two.Comment: Typos fixed and some technical details clarified. To appear in Phys. Rev. Let

Elastic response of filamentous networks with compliant crosslinks

Experiments have shown that elasticity of disordered filamentous networks with compliant crosslinks is very different from networks with rigid crosslinks. Here, we model and analyze filamentous networks as a collection of randomly oriented rigid filaments connected to each other by flexible crosslinks that are modeled as worm-like chains. For relatively large extensions we allow for enthalpic stretching of crosslinks' backbones. We show that for sufficiently high crosslink density, the network linear elastic response is affine on the scale of the filaments' length. The nonlinear regime can become highly nonaffine and is characterized by a divergence of the elastic modulus at finite strain. In contrast to the prior predictions, we do not find an asymptotic regime in which the differential elastic modulus scales linearly with the stress, although an approximate linear dependence can be seen in a transition from entropic to enthalpic regimes. We discuss our results in light of the recent experiments.Comment: 10 pages, 11 figure

Non-equilibrium mechanics and dynamics of motor activated gels

The mechanics of cells is strongly affected by molecular motors that generate forces in the cellular cytoskeleton. We develop a model for cytoskeletal networks driven out of equilibrium by molecular motors exerting transient contractile stresses. Using this model we show how motor activity can dramatically increase the network's bulk elastic moduli. We also show how motor binding kinetics naturally leads to enhanced low-frequency stress fluctuations that result in non-equilibrium diffusive motion within an elastic network, as seen in recent \emph{in vitro} and \emph{in vivo} experiments.Comment: 21 pages, 8 figure

Driven diffusive systems with mutually interactive Langmuir kinetics

We investigate the simple one-dimensional driven model, the totally asymmetric exclusion process, coupled to mutually interactive Langmuir kinetics. This model is motivated by recent studies on clustering of motor proteins on microtubules. In the proposed model, the attachment and detachment rates of a particle are modified depending upon the occupancy of neighbouring sites. We first obtain continuum mean-field equations and in certain limiting cases obtain analytic solutions. We show how mutual interactions increase (decrease) the effects of boundaries on the phase behavior of the model. We perform Monte Carlo simulations and demonstrate that our analytical approximations are in good agreement with the numerics over a wide range of model parameters. We present phase diagrams over a selective range of parameters.Comment: 9 pages, 8 Figure

Active biopolymer networks generate scale-free but euclidean clusters

We report analytical and numerical modelling of active elastic networks, motivated by experiments on crosslinked actin networks contracted by myosin motors. Within a broad range of parameters, the motor-driven collapse of active elastic networks leads to a critical state. We show that this state is qualitatively different from that of the random percolation model. Intriguingly, it possesses both euclidean and scale-free structure with Fisher exponent smaller than $2$. Remarkably, an indistinguishable Fisher exponent and the same euclidean structure is obtained at the critical point of the random percolation model after absorbing all enclaves into their surrounding clusters. We propose that in the experiment the enclaves are absorbed due to steric interactions of network elements. We model the network collapse, taking into account the steric interactions. The model shows how the system robustly drives itself towards the critical point of the random percolation model with absorbed enclaves, in agreement with the experiment.Comment: 6 pages, 7 figure

Nonlinear elasticity of composite networks of stiff biopolymers with flexible linkers

Motivated by recent experiments showing nonlinear elasticity of in vitro networks of the biopolymer actin cross-linked with filamin, we present an effective medium theory of flexibly cross-linked stiff polymer networks. We model such networks by randomly oriented elastic rods connected by flexible connectors to a surrounding elastic continuum, which self-consistently represents the behavior of the rest of the network. This model yields a crossover from a linear elastic regime to a highly nonlinear elastic regime that stiffens in a way quantitatively consistent with experiment.Comment: 4 pages, 3 figure

Dynamic characteristics and processing of fillers in polyurethane elastomers for vibration damping applications

Polyurethane elastomers have the potential of being used to reduce vibrational noise in many engineering applications. The performance of the elastomer is directly related to matching the nature of the mechanical loss characteristics to the frequency and temperature dependence of the source of the vibration. Materials with a broad frequency response and good mechanical properties are desirable for situations were load bearing and isolation becomes an issue. Because automobile, and other related vehicles operate over a broad temperature range, it is desirable for the damping characteristics of the elastomer to ideally be independent of temperature and frequency. In practice, this is not possible and the creation of materials with a broad spectrum response is desirable. In this paper, the effects of various fillers on the breadth and temperature dependence of the vibration damping characteristics of a filled and crosslinked polyurethane elastomer are explored. The fillers studied are wollastonite, barium sulphate and talc. These materials have different shapes, sizes and surface chemistry and undergo different types of interaction with the matrix. The vibration damping characteristics were further varied by the use of a crosslinking agent. Data presented on the rheological characteristics indicate the strength of the filler-polyol interactions. Dielectric relaxation and dynamic mechanical thermal analysis demonstrate the way in which changes in the type of filler, concentration and amount of crosslinker lead to changes in the location and breadth of the energy dissipation process in these elastomers. The vibration damping characteristics of a selected material are presented to demonstrate the potential of these materials

Multi-scale strain-stiffening of semiflexible bundle networks

Bundles of polymer filaments are responsible for the rich and unique mechanical behaviors of many biomaterials, including cells and extracellular matrices. In fibrin biopolymers, whose nonlinear elastic properties are crucial for normal blood clotting, protofibrils self-assemble and bundle to form networks of semiflexible fibers. Here we show that the extraordinary strain-stiffening response of fibrin networks is a direct reflection of the hierarchical architecture of the fibrin fibers. We measure the rheology of networks of unbundled protofibrils and find excellent agreement with an affine model of extensible wormlike polymers. By direct comparison with these data, we show that physiological fibrin networks composed of thick fibers can be modeled as networks of tight protofibril bundles. We demonstrate that the tightness of coupling between protofibrils in the fibers can be tuned by the degree of enzymatic intermolecular crosslinking by the coagulation Factor XIII. Furthermore, at high stress, the protofibrils contribute independently to the network elasticity, which may reflect a decoupling of the tight bundle structure. The hierarchical architecture of fibrin fibers can thus account for the nonlinearity and enormous elastic resilience characteristic of blood clots.Comment: 27 pages including 8 figures and Supplementary Dat

Critical behaviour in the nonlinear elastic response of hydrogels

In this paper we study the elastic response of synthetic hydrogels to an applied shear stress. The hydrogels studied here have previously been shown to mimic the behaviour of biopolymer networks when they are sufficiently far above the gel point. We show that near the gel point they exhibit an elastic response that is consistent with the predicted critical behaviour of networks near or below the isostatic point of marginal stability. This point separates rigid and floppy states, distinguished by the presence or absence of finite linear elastic moduli. Recent theoretical work has also focused on the response of such networks to finite or large deformations, both near and below the isostatic point. Despite this interest, experimental evidence for the existence of criticality in such networks has been lacking. Using computer simulations, we identify critical signatures in the mechanical response of sub-isostatic networks as a function of applied shear stress. We also present experimental evidence consistent with these predictions. Furthermore, our results show the existence of two distinct critical regimes, one of which arises from the nonlinear stretch response of semi-flexible polymers.

Actively Contracting Bundles of Polar Filaments

We introduce a phenomenological model to study the properties of bundles of polar filaments which interact via active elements. The stability of the homogeneous state, the attractors of the dynamics in the unstable regime and the tensile stress generated in the bundle are discussed. We find that the interaction of parallel filaments can induce unstable behavior and is responsible for active contraction and tension in the bundle. Interaction between antiparallel filaments leads to filament sorting. Our model could apply to simple contractile structures in cells such as stress fibers.Comment: 4 pages, 4 figures, RevTex, to appear in Phys. Rev. Let