We develop an effective medium approach to the mechanics of disordered, semiflexible polymer networks and study the response of such networks to uniform and nonuniform strain. We identify distinct elastic regimes in which the contributions of either filament bending or stretching to the macroscopic modulus vanish. We also show that our effective medium theory predicts a crossover between affine and nonaffine strain, consistent with both prior numerical studies and scaling theory.
The cytoskeleton of living cells contains many types of crosslinkers. Some crosslinkers allow energy-free rotations between filaments and others do not. The mechanical interplay between these different crosslinkers is an open issue in cytoskeletal mechanics. Therefore, we develop a theoretical framework based on rigidity percolation to study a generic filamentous system containing both stretching and bond-bending forces to address this issue. The framework involves both analytical calculations via effective medium theory and numerical simulations on a percolating triangular lattice with very good agreement between both. We find that the introduction of angle-constraining crosslinkers to a semiflexible filamentous network with freely rotating crosslinks can cooperatively lower the onset of rigidity to the connectivity percolation threshold—a result argued for years but never before obtained via effective medium theory. This allows the system to ultimately attain rigidity at the lowest concentration of material possible. We further demonstrate that introducing angle-constraining crosslinks results in mechanical behaviour similar to just freely rotating crosslinked semflexible filaments, indicating redundancy and universality. Our results also impact upon collagen and fibrin networks in biological and bio-engineered tissues.
Among mammalian soft tissues, articular cartilage is particularly interesting because it can endure a lifetime of daily mechanical loading despite having minimal regenerative capacity. This remarkable resilience may be due to the depth-dependent mechanical properties, which have been shown to localize strain and energy dissipation. This paradigm proposes that these properties arise from the depth-dependent collagen fiber orientation. Nevertheless, this structure-function relationship has not yet been quantified. Here, we use confocal elastography, quantitative polarized light microscopy, and Fourier-transform infrared imaging to make same-sample measurements of the depth-dependent shear modulus, collagen fiber organization, and extracellular matrix concentration in neonatal bovine articular cartilage. We find weak correlations between the shear modulus |G(∗)| and both the collagen fiber orientation and polarization. We find a much stronger correlation between |G(∗)| and the concentration of collagen fibers. Interestingly, very small changes in collagen volume fraction vc lead to orders-of-magnitude changes in the modulus with |G(∗)| scaling as (vc - v0)(ξ). Such dependencies are observed in the rheology of other biopolymer networks whose structure exhibits rigidity percolation phase transitions. Along these lines, we propose that the collagen network in articular cartilage is near a percolation threshold that gives rise to these large mechanical variations and localization of strain at the tissue's surface.
Motivated by recent observations of rheochaos in sheared wormlike micelles, we study the coupled nonlinear partial differential equations for the hydrodynamic velocity and order-parameter fields in a sheared nematogenic fluid. In a suitable parameter range, we find irregular, dynamic shear banding and establish by decisive numerical tests that the chaos we observe in the model is spatiotemporal in nature.
The cytoskeleton precisely tunes its mechanics by altering interactions between semiflexible actin filaments, rigid microtubules, and crosslinking proteins. We use optical tweezers microrheology and confocal microscopy to characterize how varying crosslinking motifs impact the mesoscale mechanics and mobility of actin-microtubule composites. We show that, upon subtle changes in crosslinking patterns, composites can exhibit two distinct classes of force response – primarily elastic versus more viscous. For example, a composite in which actin and microtubules are crosslinked to each other but not to themselves is markedly more elastic than one in which both filaments are independently crosslinked. Notably, this distinction only emerges at mesoscopic scales in response to nonlinear forcing, whereas varying crosslinking motifs have little impact on the microscale mechanics and mobility. Our unexpected scale-dependent results not only inform the physics underlying key cytoskeleton processes and structures, but, more generally, provide valuable perspective to materials engineering endeavors focused on polymer composites.
Motivated by recent experiments showing the compressive buckling of microtubules in cells, we study theoretically the mechanical response of, and force propagation along elastic filaments embedded in a non-linear elastic medium. We find that embedded microtubules buckle when their compressive load exceeds a critical value fc, and that the resulting deformation is restricted to a penetration depth that depends on both the non-linear material properties of the surrounding cytoskeleton, as well as the direct coupling of the microtubule to the cytoskeleton. The deformation amplitude depends on the applied load f > fc as (f − fc) 1/2 . This work shows how the range of compressive force transmission by microtubules can be tens of microns and is governed by the mechanical coupling to the surrounding cytoskeleton.
The cytoskeleton is a dynamic network of proteins, including actin, microtubules, and their associated motor proteins, that enables essential cellular processes such as motility, division, and growth. While actomyosin networks are extensively studied, how interactions between actin and microtubules, ubiquitous in the cytoskeleton, influence actomyosin activity remains an open question. Here, we create a network of co-entangled actin and microtubules driven by myosin II. We combine dynamic differential microscopy, particle image velocimetry, and particle tracking to show that both actin and microtubules undergo ballistic contraction with unexpectedly indistinguishable characteristics. This contractility is distinct from faster disordered motion and rupturing that active actin networks exhibit. Our results suggest that microtubules enable self-organized myosin-driven contraction by providing flexural rigidity and enhanced connectivity to actin networks. Beyond the immediate relevance to cytoskeletal dynamics, our results shed light on the design of active materials that can be precisely tuned by the network composition.
With a view to understanding the "rheochaos" observed in recent experiments in a variety of orientable fluids, we study numerically the equations of motion of the spatiotemporal evolution of the traceless symmetric order parameter of a sheared nematogenic fluid. In particular we establish, by decisive numerical tests, that the irregular oscillatory behavior seen in a region of parameter space where the nematic is not stably flow-aligning is in fact spatiotemporal chaos. We outline the dynamical phase diagram of the model and study the route to the chaotic state. We find that spatiotemporal chaos in this system sets in via a regime of spatiotemporal intermittency, with a power-law distribution of the widths of laminar regions, as in H. Chaté and P. Manneville, Phys. Rev. Lett. 58, 112 (1987). Further, the evolution of the histogram of band sizes shows a growing length scale as one moves from the chaotic towards the flow-aligned phase. Finally we suggest possible experiments in which one can observe the intriguing behaviors discussed here.
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