In embryogenesis, epithelial cells, acting as individual entities or as coordinated aggregates in a tissue, exhibit strong coupling between chemical signalling and mechanical responses to internally or externally applied stresses. Intercellular communication in combination with such coordination of morphogenetic movements can lead to drastic modifications in the calcium distribution in the cells. In this paper we extend the recent mechanochemical model in [K. Kaouri, P.K. Maini, P.A. Skourides, N. Christodoulou, S.J. Chapman. J. Math. Biol., 78 (2019) 2059--2092], for an epithelial continuum in one dimension, to a more realistic multi-dimensional case. The resulting parametrised governing equations consist of an advection-diffusion-reaction system for calcium signalling coupled with active-stress linear viscoelasticity and equipped with pure Neumann boundary conditions. We implement a mixed finite element method for the simulation of this complex multiphysics problem. Special care is taken in the treatment of the stress-free boundary conditions for the viscoelasticity in order to eliminate rigid motions from the space of admissible displacements. The stability and solvability of the continuous weak formulation is shown using fixed-point theory. We investigate numerically the solutions of this system and show that solitary waves and periodic wavetrains of calcium propagate through the embryonic epithelial sheet. We analyse the bifurcations of the system guided by the bifurcation analysis of the one-dimensional model. We also demonstrate the nucleation of calcium sparks into synchronous calcium waves coupled with contraction. This coupled model can be employed to gain insights into recent experimental observations in the context of embryogenesis, but also in other biological systems such as cancer cells, wound healing, keratinocytes, or white blood cells.
翻译:在胚胎形成过程中,作为个体实体或作为组织中协调的聚合体,上皮细胞在化学信号和机械对内外部应用压力的反应之间表现出强烈的结合。细胞间交流,加上这种形态运动的协调,可以导致细胞内钙分布的急剧改变。在本文中,我们扩展了最近在[K. Kaouri,P.K.K.Miani,P.A.Skourides,N.Christodoul,S.J.Chapman.J.Math.Biol.,78(2019-2059-2092年),Biol.,78(2019-2059-2092年),用于一个层面的显性直向直向直向直向直径直径的直径直径的直径直径直径。由此形成的正对等方形平面的平衡化系统包括一个振动-振动-振荡-振动系统,以及电动线性直线性粘度和纯纽曼边界条件。我们在模拟这一复杂的多物理的直位直径直径直径直径直径分析中,我们从一个无压的直流的直径直径直位直的直的直的直系的直系的直的直向的直径直向的直向的直向的直向的直向的直的直的直的直向下运动的直向,通过直向运动向的直向的直向运动向运动向运动向的直向的直向运动向运动向运动向运动向运动向运动向运动向运动向运动向运动向运动向运动向运动向,展示了直向的直向运动。