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Register a new userQuantifying material properties of cell monolayers by analyzing integer topological defects
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In developing organisms, internal cellular processes generate mechanical stresses at the tissue scale. The resulting deformations depend on the material properties of the tissue, which can exhibit long-ranged orientational order and topological defects. It remains a challenge to determine these properties on the time scales relevant for developmental processes. Here, we build on the physics of liquid crystals to determine material parameters of cell monolayers. Specifically, we use a hydrodynamic description to characterize the stationary states of compressible active polar fluids around defects. We illustrate our approach by analyzing monolayers of C2C12 cells in small circular confinements, where they form a single topological defect with integer charge. We find that such monolayers exert compressive stresses at the defect centers, where localized cell differentiation and formation of three-dimensional shapes is observed.
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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.
Monolayers of transition-metal dichalcogenides such as WSe2 have become increasingly attractive due to their potential in electrical and optical applications. Because the properties of these 2D systems are known to be affected by their surroundings, we report how the choice of the substrate material affects the optical properties of monolayer WSe2. To accomplish this study, pump-density-dependent micro-photoluminescence measurements are performed with time-integrating and time-resolving acquisition techniques. Spectral information and power-dependent mode intensities are compared at 290K and 10K for exfoliated WSe2 on SiO2/Si, sapphire (Al2O3), hBN/Si3N4/Si, and MgF2, indicating substrate-dependent appearance and strength of exciton, trion, and biexciton modes. Additionally, one CVD-grown WSe2 monolayer on sapphire is included in this study for direct comparison with its exfoliated counterpart. Time-resolved micro-photoluminescence shows how radiative decay times strongly differ for different substrate materials. Our data indicates exciton-exciton annihilation as a shortening mechanism at room temperature, and subtle trends in the decay rates in correlation to the dielectric environment at cryogenic temperatures. On the measureable time scales, trends are also related to the extent of the respective 2D-excitonic modes appearance. This result highlights the importance of further detailed characterization of exciton features in 2D materials, particularly with respect to the choice of substrate.
Defects, and in particular topological defects, are architectural motifs that play a crucial role in natural materials. Here we provide a systematic strategy to introduce such defects in mechanical metamaterials. We first present metamaterials that are a mechanical analogue of spin systems with tunable ferromagnetic and antiferromagnetic interactions, then design an exponential number of frustration-free metamaterials, and finally introduce topological defects by rotating a string of building blocks in these metamaterials. We uncover the distinct mechanical signature of topological defects by experiments and simulations, and leverage this to design complex metamaterials in which we can steer deformations and stresses towards parts of the system. Our work presents a new avenue to systematically include spatial complexity, frustration, and topology in mechanical metamaterials.
It is known that mechanical interactions couple a cell to its neighbors, enabling a feedback loop to regulate tissue growth. However, the interplay between cell-cell adhesion strength, local cell density and force fluctuations in regulating cell proliferation is poorly understood. Here, we show that spatial variations in the tumor growth rates, which depend on the location of cells within tissue spheroids, are strongly influenced by cell-cell adhesion. As the strength of the cell-cell adhesion increases, intercellular pressure initially decreases, enabling dormant cells to more readily enter into a proliferative state. We identify an optimal cell-cell adhesion regime where pressure on a cell is a minimum, allowing for maximum proliferation. We use a theoretical model to validate this novel collective feedback mechanism coupling adhesion strength, local stress fluctuations and proliferation.Our results, predicting the existence of a non-monotonic proliferation behavior as a function of adhesion strength, are consistent with experimental results. Several experimental implications of the proposed role of cell-cell adhesion in proliferation are quantified, making our model predictions amenable to further experimental scrutiny. We show that the mechanism of contact inhibition of proliferation, based on a pressure-adhesion feedback loop, serves as a unifying mechanism to understand the role of cell-cell adhesion in proliferation.
Recent experiments on monolayers of spindle-like cells plated on adhesive stripe-shaped domains have provided a convincing demonstration that certain types of collective phenomena in epithelia are well described by active nematic hydrodynamics. While recovering some of the hallmark predictions of this framework, however, these experiments have also revealed a number of unexpected features that could be ascribed to the existence of chirality over length scales larger than the typical size of a cell. In this article we elaborate on the microscopic origin of chiral stresses in nematic cell monolayers and investigate how chirality affects the motion of topological defects, as well as the collective motion in stripe-shaped domains. We find that chirality introduces a characteristic asymmetry in the collective cellular flow, from which the ratio between chiral and non-chiral active stresses can be inferred by particle-image-velocimetry measurements. Furthermore, we find that chirality changes the nature of the spontaneous flow transition under confinement and that, for specific anchoring conditions, the latter has the structure of an imperfect pitchfork bifurcation.
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