Holmes William R
Department of Mathematics, University of California Irvine, Irvine, CA, USA,
Bull Math Biol. 2014 Jan;76(1):157-83. doi: 10.1007/s11538-013-9914-6. Epub 2013 Oct 25.
Reaction diffusion systems are often used to study pattern formation in biological systems. However, most methods for understanding their behavior are challenging and can rarely be applied to complex systems common in biological applications. I present a relatively simple and efficient, nonlinear stability technique that greatly aids such analysis when rates of diffusion are substantially different. This technique reduces a system of reaction diffusion equations to a system of ordinary differential equations tracking the evolution of a large amplitude, spatially localized perturbation of a homogeneous steady state. Stability properties of this system, determined using standard bifurcation techniques and software, describe both linear and nonlinear patterning regimes of the reaction diffusion system. I describe the class of systems this method can be applied to and demonstrate its application. Analysis of Schnakenberg and substrate inhibition models is performed to demonstrate the methods capabilities in simplified settings and show that even these simple models have nonlinear patterning regimes not previously detected. The real power of this technique, however, is its simplicity and applicability to larger complex systems where other nonlinear methods become intractable. This is demonstrated through analysis of a chemotaxis regulatory network comprised of interacting proteins and phospholipids. In each case, predictions of this method are verified against results of numerical simulation, linear stability, asymptotic, and/or full PDE bifurcation analyses.
反应扩散系统常用于研究生物系统中的模式形成。然而,大多数理解其行为的方法具有挑战性,很少能应用于生物应用中常见的复杂系统。我提出了一种相对简单且高效的非线性稳定性技术,当扩散速率有显著差异时,该技术能极大地辅助此类分析。该技术将反应扩散方程组简化为常微分方程组,追踪均匀稳态的大振幅、空间局部扰动的演化。使用标准分岔技术和软件确定该系统的稳定性特性,描述了反应扩散系统的线性和非线性模式形成机制。我描述了可应用此方法的系统类别并展示了其应用。对施纳肯贝格模型和底物抑制模型进行分析,以证明该方法在简化设置中的能力,并表明即使是这些简单模型也存在先前未检测到的非线性模式形成机制。然而,该技术的真正优势在于其简单性以及对其他非线性方法难以处理的更大复杂系统的适用性。通过对由相互作用蛋白和磷脂组成的趋化调节网络的分析来证明这一点。在每种情况下,该方法的预测都与数值模拟、线性稳定性、渐近和/或全偏微分方程分岔分析的结果进行了验证。