A minimal model of smoothly dividing disk-shaped cells

Lukas Hupe, Yoav G. Pollack, Jonas Isensee, Aboutaleb Amiri, Ramin Golestanian, Philip Bittihn
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Abstract

Replication through cell division is one of the most fundamental processes of life and a major driver of dynamics in systems ranging from bacterial colonies to embryogenesis, tissues and tumors. While regulation often plays a role in shaping self-organization, mounting evidence suggests that many biologically relevant behaviors exploit principles based on a limited number of physical ingredients, and particle-based models have become a popular platform to reconstitute and investigate these emergent dynamics. However, incorporating division into such models often leads to aberrant mechanical fluctuations that hamper physically meaningful analysis. Here, we present a minimal model focusing on mechanical consistency during division. Cells are comprised of two nodes, overlapping disks which separate from each other during cell division, resulting in transient dumbbell shapes. Internal degrees of freedom, cell-cell interactions and equations of motion are designed to ensure force continuity at all times, including through division, both for the dividing cell itself as well as interaction partners, while retaining the freedom to define arbitrary anisotropic mobilities. As a benchmark, we also translate an established model of proliferating spherocylinders with similar dynamics into our theoretical framework. Numerical simulations of both models demonstrate force continuity of the new disk cell model and quantify our improvements. We also investigate some basic collective behaviors related to alignment and orientational order and find consistency both between the models and with the literature. A reference implementation of the model is freely available as a package in the Julia programming language based on $\textit{InPartS.jl}$. Our model is ideally suited for the investigation of mechanical observables such as velocities and stresses, and is easily extensible with additional features.
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平滑分裂盘状细胞的最小模型
通过细胞分裂进行复制是生命最基本的过程之一,也是从细菌菌落到胚胎发生、组织和肿瘤等系统动态的主要驱动力。虽然调控通常在塑造自组织过程中发挥作用,但越来越多的证据表明,许多生物相关行为利用了基于有限物理成分的原理,而基于粒子的模型已成为重组和研究这些突发动力学的流行平台。然而,在这类模型中加入分裂往往会导致异常的机械波动,从而影响有物理意义的分析。在这里,我们提出了一个最小模型,重点研究分裂过程中的机械一致性。细胞由两节重叠的圆盘组成,在细胞分裂过程中相互分离,形成瞬时哑铃状。内部自由度、细胞-细胞相互作用和运动方程的设计确保了分裂细胞本身和相互作用伙伴在任何时候(包括分裂过程中)受力的连续性,同时保留了定义任意各向异性运动的自由度。作为一个基准,我们还将一个具有类似动力学的增殖球体模型转化为我们的理论框架。对这两个模型的数值模拟证明了新的盘状细胞模型的力连续性,并量化了我们的改进。我们还研究了一些与排列和定向顺序有关的基本集体行为,发现模型之间以及模型与文献之间的一致性。该模型的参考实现作为一个基于 $\textit{InPartS.jl}$ 的朱利编程语言包免费提供。我们的模型非常适合研究速度和应力等力学观测指标,而且很容易扩展附加功能。
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