Nonlinear Poisson effect in affine semiflexible polymer networks.

Stretching an elastic material along one axis typically induces contraction along the transverse axes, a phenomenon known as the Poisson effect. From these strains, one can compute the specific volume, which generally either increases or, in the incompressible limit, remains constant as the material is stretched. However, in networks of semiflexible or stiff polymers, which are typically highly compressible yet stiffen significantly when stretched, one instead sees a significant reduction in specific volume under finite strains. This volume reduction is accompanied by increasing alignment of filaments along the strain axis and a nonlinear elastic response, with stiffening of the apparent Young's modulus. For semiflexible networks, in which entropic bending elasticity governs the linear elastic regime, the nonlinear Poisson effect is caused by the nonlinear force-extension relationship of the constituent filaments, which produces a highly asymmetric response of the constituent polymers to stretching and compression. The details of this relationship depend on the geometric and elastic properties of the underlying filaments, which can vary greatly in experimental systems. Here, we provide a comprehensive characterization of the nonlinear Poisson effect in an affine network model and explore the influence of filament properties on essential features of both microscopic and macroscopic response, including strain-driven alignment and volume reduction.

Field Theory for Mechanical Criticality in Disordered Fiber Networks.

Strain-controlled criticality governs the elasticity of jamming and fiber networks. While the upper critical dimension of jamming is believed to be d_{u}=2, non-mean-field exponents are observed in numerical studies of 2D and 3D fiber networks. The origins of this remains unclear. In this study we propose a minimal mean-field model for strain-controlled criticality of fiber networks. We then extend this to a phenomenological field theory, in which non-mean-field behavior emerges as a result of the disorder in the network structure. We predict that the upper critical dimension for such systems is d_{u}=4 using a Gaussian approximation. Moreover, we identify an order parameter for the phase transition, which has been lacking for fiber networks to date.

Effects of local incompressibility on the rheology of composite biopolymer networks.

Fibrous networks such as collagen are common in biological systems. Recent theoretical and experimental efforts have shed light on the mechanics of single component networks. Most real biopolymer networks, however, are composites made of elements with different rigidity. For instance, the extracellular matrix in mammalian tissues consists of stiff collagen fibers in a background matrix of flexible polymers such as hyaluronic acid (HA). The interplay between different biopolymer components in such composite networks remains unclear. In this work, we use 2D coarse-grained models to study the nonlinear strain-stiffening behavior of composites. We introduce a local volume constraint to model the incompressibility of HA. We also perform rheology experiments on composites of collagen with HA. Theoretically and experimentally, we demonstrate that the linear shear modulus of composite networks can be increased by approximately an order of magnitude above the corresponding moduli of the pure components. Our model shows that this synergistic effect can be understood in terms of the local incompressibility of HA, which acts to suppress density fluctuations of the collagen matrix with which it is entangled.

Mechanical criticality of fiber networks at a finite temperature.

At zero temperature, spring networks with connectivity below Maxwell's isostatic threshold undergo a mechanical phase transition from a floppy state at small strains to a rigid state for applied shear strain above a critical strain threshold. Disordered networks in the floppy mechanical regime can be stabilized by entropic effects at finite temperature. We develop a scaling theory for this mechanical phase transition at finite temperature, yielding relationships between various scaling exponents. Using Monte Carlo simulations, we verify these scaling relations and identify anomalous entropic elasticity with sublinear T dependence in the linear elastic regime. While our results are consistent with prior studies of phase behavior near the isostatic point, the present work also makes predictions relevant to the broad class of disordered thermal semiflexible polymer networks for which the connectivity generally lies far below the isostatic threshold.

Strain-Controlled Critical Slowing Down in the Rheology of Disordered Networks.

Networks and dense suspensions frequently reside near a boundary between soft (or fluidlike) and rigid (or solidlike) regimes. Transitions between these regimes can be driven by changes in structure, density, or applied stress or strain. In general, near the onset or loss of rigidity in these systems, dissipation-limiting heterogeneous nonaffine rearrangements dominate the macroscopic viscoelastic response, giving rise to diverging relaxation times and power-law rheology. Here, we describe a simple quantitative relationship between nonaffinity and the excess viscosity. We test this nonaffinity-viscosity relationship computationally and demonstrate its rheological consequences in simulations of strained filament networks and dense suspensions. We also predict critical signatures in the rheology of semiflexible and stiff biopolymer networks near the strain stiffening transition.

Motor-free contractility of active biopolymer networks.

Contractility in animal cells is often generated by molecular motors such as myosin, which require polar substrates for their function. Motivated by recent experimental evidence of motor-independent contractility, we propose a robust motor-free mechanism that can generate contraction in biopolymer networks without the need for substrate polarity. We show that contractility is a natural consequence of active binding-unbinding of crosslinkers that breaks the principle of detailed balance, together with the asymmetric force-extension response of semiflexible biopolymers. We have extended our earlier work to discuss the motor-free contraction of viscoelastic biopolymer networks. We calculate the resulting contractile velocity using a microscopic model and show that it can be reduced to a simple coarse-grained model under certain limits. Our model may provide an explanation of recent reports of motor-independent contractility in cells. Our results also suggest a mechanism for generating contractile forces in synthetic active materials.

Effective medium theory for mechanical phase transitions of fiber networks.

Networks of stiff fibers govern the elasticity of biological structures such as the extracellular matrix of collagen. These networks are known to stiffen nonlinearly under shear or extensional strain. Recently, it has been shown that such stiffening is governed by a strain-controlled athermal but critical phase transition, from a floppy phase below the critical strain to a rigid phase above the critical strain. While this phase transition has been extensively studied numerically and experimentally, a complete analytical theory for this transition remains elusive. Here, we present an effective medium theory (EMT) for this mechanical phase transition of fiber networks. We extend a previous EMT appropriate for linear elasticity to incorporate nonlinear effects via an anharmonic Hamiltonian. The mean-field predictions of this theory, including the critical exponents, scaling relations and non-affine fluctuations qualitatively agree with previous experimental and numerical results.

Nonaffine Deformation of Semiflexible Polymer and Fiber Networks.

Networks of semiflexible or stiff polymers such as most biopolymers are known to deform inhomogeneously when sheared. The effects of such nonaffine deformation have been shown to be much stronger than for flexible polymers. To date, our understanding of nonaffinity in such systems is limited to simulations or specific 2D models of athermal fibers. Here, we present an effective medium theory for nonaffine deformation of semiflexible polymer and fiber networks, which is general to both 2D and 3D and in both thermal and athermal limits. The predictions of this model are in good agreement with both prior computational and experimental results for linear elasticity. Moreover, the framework we introduce can be extended to address nonlinear elasticity and network dynamics.

Structural Features and Nonlinear Rheology of Self-Assembled Networks of Cross-Linked Semiflexible Polymers.

Disordered networks of semiflexible filaments are common support structures in biology. Familiar examples include fibrous matrices in blood clots, bacterial biofilms, and essential components of cells and tissues of plants, animals, and fungi. Despite the ubiquity of these networks in biomaterials, we have only a limited understanding of the relationship between their structural features and their highly strain-sensitive mechanical properties. In this work, we perform simulations of three-dimensional networks produced by the irreversible formation of cross-links between linker-decorated semiflexible filaments. We characterize the structure of networks formed by a simple diffusion-dependent assembly process and measure their associated steady-state rheological features at finite temperature over a range of applied prestrains that encompass the strain-stiffening transition. We quantify the dependence of network connectivity on cross-linker availability and detail the associated connectivity dependence of both linear elasticity and nonlinear strain-stiffening behavior, drawing comparisons with prior experimental measurements of the cross-linker concentration-dependent elasticity of actin gels.

Unique Role of Vimentin Networks in Compression Stiffening of Cells and Protection of Nuclei from Compressive Stress.

In this work, we investigate whether stiffening in compression is a feature of single cells and whether the intracellular polymer networks that comprise the cytoskeleton (all of which stiffen with increasing shear strain) stiffen or soften when subjected to compressive strains. We find that individual cells, such as fibroblasts, stiffen at physiologically relevant compressive strains, but genetic ablation of vimentin diminishes this effect. Further, we show that unlike networks of purified F-actin or microtubules, which soften in compression, vimentin intermediate filament networks stiffen in both compression and extension, and we present a theoretical model to explain this response based on the flexibility of vimentin filaments and their surface charge, which resists volume changes of the network under compression. These results provide a new framework by which to understand the mechanical responses of cells and point to a central role of intermediate filaments in response to compression.