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Register a new userMechanochemical induction of wrinkling morphogenesis on elastic shells
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Morphogenetic dynamics of tissue sheets require coordinated cell shape changes regulated by global patterning of mechanical forces. Inspired by such biological phenomena, we propose a minimal mechanochemical model based on the notion that cell shape changes are induced by diffusible biomolecules that influence tissue contractility in a concentration-dependent manner -- and whose concentration is in turn affected by the macroscopic tissue shape. We perform computational simulations of thin shell elastic dynamics to reveal propagating chemical and three-dimensional deformation patterns arising due to a sequence of buckling instabilities. Depending on the concentration threshold that actuates cell shape change, we find qualitatively different patterns. The mechanochemically coupled patterning dynamics are distinct from those driven by purely mechanical or purely chemical factors. Using numerical simulations and theoretical arguments, we analyze the elastic instabilities that result from our model and provide simple scaling laws to identify wrinkling morphologies.
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We systematically explore the self-assembly of semi-flexible polymers in deformable spherical confinement across a wide regime of chain stiffness, contour lengths and packing fractions by means of coarse-grained molecular dynamics simulations. Compliant, DNA-like filaments are found to undergo a continuous crossover from two distinct surface-ordered quadrupolar states, both characterized by tetrahedral patterns of topological defects, to either longitudinal or latitudinal bipolar structures with increasing polymer concentrations. These transitions, along with the intermediary arrangements that they involve, may be attributed to the combination of an orientational wetting phenomenon with subtle density- and contour-length-dependent variations in the elastic anisotropies of the corresponding liquid crystal phases. Conversely, the organization of rigid, microtubule-like polymers evidences a progressive breakdown of continuum elasticity theory as chain dimensions become comparable to the equilibrium radius of the encapsulating membrane. In this case, we observe a gradual shift from prolate, tactoid-like morphologies to oblate, erythrocyte-like structures with increasing contour lengths, which is shown to arise from the interplay between nematic ordering, polymer and membrane buckling. We further provide numerical evidence of a number of yet-unidentified, self-organized states in such confined systems of stiff achiral filaments, including spontaneous spiral smectic assemblies, faceted polyhedral and twisted bundle-like arrangements. Our results are quantified through the introduction of several order parameters and an unsupervised learning scheme for the localization of surface topological defects, and are in excellent agreement with field-theoretical predictions as well as classical elastic theories of thin rods and spherical shells.
Inspired by active shape morphing in developing tissues and biomaterials, we investigate two generic mechanochemical models where the deformations of a thin elastic sheet are driven by, and in turn affect, the concentration gradients of a chemical signal. We develop numerical methods to study the coupled elastic deformations and chemical concentration kinetics, and illustrate with computations the formation of different patterns depending on shell thickness, strength of mechanochemical coupling and diffusivity. In the first model, the sheet curvature governs the production of a contractility inhibitor and depending on the threshold in the coupling, qualitatively different patterns occur. The second model is based on the stress--dependent activity of myosin motors, and demonstrates how the concentration distribution patterns of molecular motors are affected by the long-range deformations generated by them. Since the propagation of mechanical deformations is typically faster than chemical kinetics (of molecular motors or signaling agents that affect motors), we describe in detail and implement a numerical method based on separation of timescales to effectively simulate such systems. We show that mechanochemical coupling leads to long-range propagation of patterns in disparate systems through elastic instabilities even without the diffusive or advective transport of the chemicals.
Experimentally measuring the elastic properties of thin biological surfaces is non-trivial, particularly when they are curved. One technique that may be used is the indentation of a thin sheet of material by a rigid indenter, whilst measuring the applied force and displacement. This gives immediate information on the fracture strength of the material (from the force required to puncture), but it is also theoretically possible to determine the elastic properties by comparing the resulting force-displacement curves with a mathematical model. Existing mathematical studies generally assume that the elastic surface is initially flat, which is often not the case for biological membranes. We previously outlined a theory for the indentation of curved isotropic, incompressible, hyperelastic membranes (with no bending stiffness) which breaks down for highly curved surfaces, as the entire membrane becomes wrinkled. Here we introduce the effect of bending stiffness, ensuring that energy is required to change the shell shape without stretching, and find that commonly neglected terms in the shell equilibrium equation must be included. The theory presented here allows for the estimation of shape- and size-independent elastic properties of highly curved surfaces via indentation experiments, and is particularly relevant for biological surfaces.
Thin elastic membranes form complex wrinkle patterns when put on substrates of different shapes. Such patterns continue to receive attention across science and engineering. This is due, in part, to the promise of lithography-free micropatterning, but also to the observation that similar patterns arise in biological systems from growth. The challenge is to explain the patterns in any given setup, even when they fail to be robust. Building on the theoretical foundation of [Tobasco, to appear in Arch. Ration. Mech. Anal., arXiv:1906.02153], we derive a complete and simple rule set for wrinkles in the model system of a curved shell on a liquid bath. Our rules apply to shells whose initial Gaussian curvatures are of one sign, such as cutouts of saddles and spheres. They predict the surprising coexistence of orderly wrinkles alongside disordered regions where the response appears stochastic, which we confirm in experiment and simulation. They also unveil the role of the shells medial axis, a distinguished locus of points that we show is a basic driver in pattern selection. Finally, they explain how the sign of the shells initial curvature dictates the presence or lack of disorder. Armed with our simple rules, and the methodology underlying them, one can anticipate the creation of designer wrinkle patterns.
Growing experimental evidence indicates that topological defects could serve as organizing centers in the morphogenesis of tissues. In this article we provide a quantitative explanation for this phenomenon, rooted in the buckling theory of deformable active polar liquid crystals. Using a combination of linear stability analysis and computational fluid dynamics, we demonstrate that confined cell layers are unstable to the formation of protrusions in the presence of disclinations. The instability originates from an interplay between the focusing of the elastic forces, mediated by defects, and the renormalization of the systems surface tension by the active flow. The post-transitional regime is also characterized by several complex morphodynamical processes, such as oscillatory deformations, droplet nucleation and active turbulence. Our findings offer an explanation of recent observations on tissue morphogenesis and shed light on the dynamics of active surfaces in general.
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