No Arabic abstract
Bundles of stiff filaments are ubiquitous in the living world, found both in the cytoskeleton and in the extracellular medium. These bundles are typically held together by smaller cross-linking molecules. We demonstrate analytically, numerically and experimentally that such bundles can be kinked, i.e., have localized regions of high curvature that are long-lived metastable states. We propose three possible mechanisms of kink stabilization: a difference in trapped length of the filament segments between two cross links; a dislocation where the endpoint of a filament occurs within the bundle, and the braiding of the filaments in the bundle. At a high concentration of cross links, the last two effects lead to the topologically protected kinked states. Finally, we explore numerically and analytically the transition of the metastable kinked state to the stable straight bundle.
We use continuum theory to show that chirality is a key thermodynamic control parameter for the aggregation of biopolymers: chirality produces a stable disperse phase of hexagonal bundles under moderately poor solvent conditions, as has been observed in {it in-vitro} studies of F-actin [O. Pelletier {it et al.}, Phys. Rev. Lett. {bf 91}, 148102 (2003)]. The large characteristic radius of these chiral bundles is not determined by a mysterious long-range molecular interaction but by in-plane shear elastic stresses generated by the interplay between a chiral torque and an unusual, but universal, non-linear gauge term in the strain tensor of ordered chains that is imposed by rotational invariance.
We discuss the response of biopolymer filament bundles bound by transient cross linkers to compressive loading. These systems admit a mechanical instability at stresses typically below that of traditional Euler buckling. In this instability, there is thermally-activated pair production of topological defects that generate localized regions of bending -- kinks. These kinks shorten the bundles effective length thereby reducing the elastic energy of the mechanically loaded structure. This effect is the thermal analog of the Schwinger effect, in which a sufficiently large electric field causes electron-positron pair production. We discuss this analogy and describe the implications of this analysis for the mechanics of biopolymer filament bundles of various types under compression.
We examine the nonequilibrium production of topological defects -- braids -- in semiflexible filament bundles under cycles of compression and tension. During these cycles, the period of compression facilitates the thermally activated pair production of braid/anti-braid pairs, which then may separate when the bundle is under tension. As result, appropriately tuned alternating periods of compression and extension should lead to the proliferation of braid defects in a bundle so that linear density of these pairs far exceeds that expected in thermal equilibrium. Secondly, we examine the slow extension of braided bundles under tension, showing that their end-to-end length creeps nonmonotonically under a fixed force due to braid deformation and the motion of the braid pair along the bundle. We conclude with a few speculations regarding experiments on semiflexible filament bundles and their networks.
Monolayers of anisotropic cells exhibit long-ranged orientational order and topological defects. During the development of organisms, orientational order often influences morphogenetic events. However, the linkage between the mechanics of cell monolayers and topological defects remains largely unexplored. This holds specifically at the time scales relevant for tissue morphogenesis. Here, we build on the physics of liquid crystals to determine material parameters of cell monolayers. In particular, we use a hydrodynamical description of an active polar fluid to study the steady-state mechanical patterns at integer topological defects. Our description includes three distinct sources of activity: traction forces accounting for cell-substrate interactions as well as anisotropic and isotropic active nematic stresses accounting for cell-cell interactions. We apply our approach to C2C12 cell monolayers in small circular confinements, which form isolated aster or spiral topological defects. By analyzing the velocity and orientational order fields in spirals as well as the forces and cell number density fields in asters, we determine mechanical parameters of C2C12 cell monolayers. Our work shows how topological defects can be used to fully characterize the mechanical properties of biological active matter.
We present a Landau type theory for the non-linear elasticity of biopolymer gels with a part of the order parameter describing induced nematic order of fibers in the gel. We attribute the non-linear elastic behavior of these materials to fiber alignment induced by strain. We suggest an application to contact guidance of cell motility in tissue. We compare our theory to simulation of a disordered lattice model for biopolymers. We treat homogeneous deformations such as simple shear, hydrostatic expansion, and simple extension, and obtain good agreement between theory and simulation. We also consider a localized perturbation which is a simple model for a contracting cell in a medium.