We propose a nonlinear extension of the standard tube model for semidilute solutions of freely-sliding semiflexible polymers. Non-affine filament deformations at the entanglement scale, the renormalisation of direct interactions by thermal fluctuations, and the geometry of large deformations are systematically taken into account. The stiffening response predicted for athermal solutions of stiff rods is found to be thermally suppressed. Instead, we obtain a broad linear response regime, supporting the interpretation of shear stiffening at finite frequencies in polymerised actin solutions as indicative of coupling to longitudinal modes. We observe a destabilizing effect of large strains (about 100%), suggesting shear banding as a plausible explanation for the widely observed catastrophic collapse of in-vitro biopolymer solutions, usually attributed to network damage. In combination with friction-type interactions, our analysis provides an analytically tractable framework to address the nonlinear viscoplasticity of biological tissue on a molecular basis.
Recent experimental investigations have revealed a non-Maxwellian absorption pattern in the rheological spectra of actin gels, which was interpreted in terms of transient bonds. Here we examine the consequences of reversible crosslinking on the apparent linear spectra of biopolymer solutions theoretically. For a schematic model consisting of a reversibly crosslinked power-law fluid we obtain a simple analytical prediction for the position of the absorption peak, which is backed up by a numerical evaluation of the inelastic glassy wormlike chain model. This establishes bond breaking as a nonlin…
Precise measuring the deforming force and observation of the deformation dynamics along with difficulties of micro sample preparation are the main problems which make the micro-scale experiments difficult. Here we present a method for direct observation and measurements of mechanical parameters of elastic microfibrils in air using the pressure of optical light. For DNA microfibers, observation of the behavior dynamics of mechanical characteristics and the method sensitivity to changing the structure are demonstrated.
We suggest a simple model for reversible cross-links, binding and unbinding to/from a network of semiflexible polymers. The resulting frequency dependent response of the network to an applied shear is calculated via Brownian dynamics simulations. It is shown to be rather complex with the timescale of the linkers competing with the excitations of the network. If the lifetime of the linkers is the longest timescale, as is indeed the case in most biological networks, then a distinct low frequency peak of the loss modulus develops. The storage modulus shows a corresponding decay from its plateau v…
We propose a physical model for the nonlinear inelastic mechanics of sticky biopolymer networks with potential applications to inelastic cell mechanics. It consists in a minimal extension of the glassy wormlike chain (GWLC) model, which has recently been highly successful as a quantitative mathematical description of the viscoelastic properties of biopolymer networks and cells. To extend its scope to nonequilibrium situations, where the thermodynamic state variables may evolve dynamically, the GWLC is furnished with an explicit representation of the kinetics of breaking and reforming sticky bo…
A long standing puzzle in the rheology of living cells is the origin of the experimentally observed long time stress relaxation. The mechanics of the cell is largely dictated by the cytoskeleton, which is a biopolymer network consisting of transient crosslinkers, allowing for stress relaxation over time. Moreover, these networks are internally stressed due to the presence of molecular motors. In this work we propose a theoretical model that uses a mode-dependent mobility to describe the stress relaxation of such prestressed transient networks. Our theoretical predictions agree favorably with e…
We investigate the delayed rupture of biopolymer gels under a constant shear load by simultaneous dynamic light scattering and rheology measurements. We unveil the crucial role of normal stresses built up during gelation: all samples that eventually fracture self-weaken during the gelation process, as revealed by a partial relaxation of the normal stress concomitant to a burst of microscopic plastic rearrangements. Upon applying a shear stress, weakened gels exhibit in the creep regime distinctive signatures in their microscopic dynamics, which anticipate macroscopic fracture by up to thousand…
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 p…
Live cells have ambiguous mechanical properties. They were often described as either elastic solids or viscoelastic fluids and have recently been classified as soft glassy materials characterized by weak power-law rheology. Nonlinear rheological measurements have moreover revealed a pronounced inelastic response indicative of a competition between stiffening and softening. It is an intriguing question whether these observations can be explained from the material properties of much simpler in-vitro reconstituted networks of biopolymers that serve as reduced model systems for the cytoskeleton. H…
Inulins are nano-meter size semi-crystalline particles, composed of oligomeric fructose units. It has been subjected to fine micro-structural analysis under temperature variations using mainly positron annihilation spectroscopy. The results show a non-monotonous temperature sensitive behaviour of the positron parameters, with considerable variation of its free volume size. The ortho-positronium pick-off component shows a major thermotropic transition at ~320K and a structure loss due to glass transition. Differential scanning calorimetry confirms the onset of the major molecular transition aro…
We present an exact analysis of the intrinsic dynamics of two types of biopolymers, described by the wormlike chain (WLC) and the wormlike rod chain (WLRC) models, respectively. We map each model to a classical field theory for an appropriate unit vector field defined on the space curve formed by the axis of the biopolymer. Conformation dynamics and energy transport are shown to be described, respectively, by the soliton-bearing Landau-Lifshitz equation and the nonlinear Schrödinger equation. This enables us to identify the soliton representing the ``conformon'' hypothesized in biopolymers.
We employ a matrix-based solver for the linear rheology of fluid-immersed disordered spring networks to reveal four distinct dynamic response regimes. One regime - completely absent in the known vacuum response - exhibits coupled fluid flow and network deformation, with both components responding non-affinely. This regime contains an additional plateau (peak) in the frequency-dependent storage (loss) modulus - features which vanish without full hydrodynamic interactions. The mechanical response of immersed networks such as biopolymers and hydrogels is thus richer than previously established, a…
Nonlinear stiffening is a ubiquitous property of major types of biopolymers that make up the extracellular matrices (ECM) including collagen, fibrin and basement membrane. Within the ECM, many types of cells such as fibroblasts and cancer cells are known to mechanically stretch their surroundings that locally stiffens the matrix. Although the bulk nonlinear elastic behaviors of these biopolymer networks are well studied, their local mechanical responses remain poorly characterized. Here, to understand how a living cell feels the nonlinear mechanical resistance from the ECM, we mimic the cell-a…
The response of cells during spreading and motility is dictated by several multi-physics events, which are triggered by extracellular cues and occur at different time-scales. For this sake, it is not completely appropriate to provide a cell with classical notions of the mechanics of materials, as for "rheology" or "mechanical response". Rather, a cell is an alive system with constituents that show a reproducible response, as for the {\em{contractility}} for single stress fibers or for the mechanical response of a biopolymer actin network, but that reorganize in response to external cues in a n…
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 rod…
The flow of colloidal suspensions is ubiquitous in nature and industry. Colloidal suspensions exhibit a wide range of rheological behavior, which should be closely related to the microscopic structure of the systems. With in-situ small-angle neutron scattering complemented by rheological measurements, we investigated the deformation behavior of a charge-stabilized colloidal glass at particle level undergoing steady shear. A short-lived, localized elastic response at particle level, termed as transient elasticity zone (TEZ), was identified from the neutron spectra. The existence of the TEZ is a…
We employ a first principles, force-level approach to self-consistently construct the anharmonic tube confinement field for entangled fluids of rigid needles and for primitive-path (PP) level chains in two limiting situations where chain stretching is assumed to either completely relax or remain unrelaxed. The influence of shear and extensional deformation and polymer orientation is determined in a nonlinear elastic limit where dissipative relaxation processes are intentionally neglected. For needles and PP-level chains, a Gaussian analysis of transverse polymer dynamical fluctuations predicts…
The activity of growing living bacteria was investigated using real-time and in situ rheology -- in stationary and oscillatory shear. Two different strains of the human pathogen Staphylococcus aureus -- strain COL and its isogenic cell wall autolysis mutant -- were considered in this work. For low bacteria density, strain COL forms small clusters, while the mutant, presenting deficient cell separation, forms irregular larger aggregates. In the early stages of growth, when subjected to a stationary shear, the viscosity of both strains increases with the population of cells. As the bacteria reac…
We use numerical simulations to study the flow of a bidisperse mixture of athermal, frictionless, soft-core two dimensional spherocylinders driven in uniform steady state shear. Energy dissipation is via a viscous drag with respect to a uniformly sheared host fluid, giving a model for a non-Brownian suspension with a Newtonian rheology. We study pressure $p$ and deviatoric shear stress $σ$ as a function of packing fraction $ϕ$, strain rate $\dotγ$, and a parameter $α$ that measures the asphericity of the particles. We consider the anisotropy of the stress tensor, the macroscopic friction $μ=σ/…
We solve exactly and describe in detail a simplified scalar model for the low frequency shear rheology of foams, emulsions, slurries, etc. [P. Sollich, F. Lequeux, P. Hebraud, M.E. Cates, Phys. Rev. Lett. 78, 2020 (1997)]. The model attributes similarities in the rheology of such ``soft glassy materials'' to the shared features of structural disorder and metastability. By focusing on the dynamics of mesoscopic elements, it retains a generic character. Interactions are represented by a mean-field noise temperature x, with a glass transition occurring at x=1 (in appropriate units). The exact sol…
