Dafalias Yannis F, Pitouras Zacharias
Department of Mechanics, National Technical University of Athens, Athens, Greece.
Biomech Model Mechanobiol. 2009 Feb;8(1):9-24. doi: 10.1007/s10237-007-0113-y. Epub 2007 Dec 6.
Polymerization of actin to form an elastic gel is one of the main mechanisms responsible for cellular motility. The particular problem addressed here stems from the need to model theoretically the growth of actin gel under controlled conditions, as observed in experiments. A biomimetic in vitro system which consists of a spherical latex bead, coated by the enzymatic protein ActA, and a reconstituted cytoplasm within which such beads are placed, induces polymerization of actin on the surface of the bead in the form of successive elastic thin spherical layers. Each newly formed layer pushes outward, and is pushed inward by, the already formed spherical layers which altogether constitute an elastic spherical shell of thickness h varying with time. Thus, a stress field is created in the shell which in turn affects the rate of polymerization as well as that of dissociation of actin gel. Given this bio-chemo-mechanical coupling, the accurate determination of the stress field becomes a subject of great importance for the understanding of the process, and it is the main objective of this work. The problem is addressed by first assuming appropriate constitutive laws for the actin gel elastic material, and then solving the only non-trivial stress equilibrium differential equation along the radial direction assuming spherical symmetry. A linear and a non-linear constitutive model for isotropic elasticity is used, appropriate for small and finite strains, respectively, and the solution is found in closed analytical forms in both cases. Two important conclusions are reached. First, the stress field depends strongly on the compressibility of the actin gel medium via the value of the Poisson ratio, for both linear and non-linear analysis. Second, the linear and non-linear solutions are very close for small strains, but they diverge progressively as the strains increase from small to large. Guided by available experimental data on the observed strain levels, the analytical results are illustrated by selected graphs of stress variation along the radial direction. At the end some comments and suggestions on the bio-chemo-mechanical coupling of actin gel growth and resorption are presented, where the role of properly defined joint isotropic invariants of stress and a unit vector along the predominant direction of free ends of actin filaments at the polymerization site is introduced.
肌动蛋白聚合形成弹性凝胶是细胞运动的主要机制之一。这里所解决的特定问题源于需要在理论上对实验中观察到的在受控条件下肌动蛋白凝胶的生长进行建模。一种仿生体外系统,它由一个涂有酶蛋白ActA的球形乳胶珠以及放置有这种珠子的重构细胞质组成,能诱导肌动蛋白在珠子表面以连续弹性薄球层的形式聚合。每个新形成的层向外推,同时被已经形成的球层向内推,这些球层共同构成一个厚度h随时间变化的弹性球壳。因此,在壳中产生了一个应力场,该应力场反过来又影响肌动蛋白凝胶的聚合速率和解离速率。考虑到这种生物 - 化学 - 力学耦合,准确确定应力场对于理解该过程变得极为重要,这也是本工作的主要目标。通过首先为肌动蛋白凝胶弹性材料假设合适的本构定律,然后假设球对称性求解沿径向方向唯一的非平凡应力平衡微分方程来解决该问题。使用了分别适用于小应变和有限应变的各向同性弹性的线性和非线性本构模型,并且在两种情况下都以封闭解析形式找到了解。得出了两个重要结论。首先,对于线性和非线性分析,应力场都通过泊松比的值强烈依赖于肌动蛋白凝胶介质的可压缩性。其次,对于小应变,线性和非线性解非常接近,但随着应变从小变大,它们逐渐发散。根据关于观察到的应变水平的现有实验数据,通过沿径向方向应力变化的选定图表来说明分析结果。最后,对肌动蛋白凝胶生长和吸收的生物 - 化学 - 力学耦合提出了一些评论和建议,其中引入了适当定义的应力联合各向同性不变量以及沿聚合位点肌动蛋白丝自由端主要方向的单位向量的作用。