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Register a new userIsolating Patterns in Open Reaction-Diffusion Systems
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Realistic examples of reaction-diffusion phenomena governing spatial and spatiotemporal pattern formation are rarely isolated systems, either chemically or thermodynamically. However, even formulations of `open reaction-diffusion systems often neglect the role of domain boundaries. Most idealizations of closed reaction-diffusion systems employ no-flux boundary conditions, and often patterns will form up to, or along, these boundaries. Motivated by boundaries of patterning fields related to the emergence of spatial form in embryonic development, we propose a set of mixed boundary conditions for a two-species reaction-diffusion system which forms inhomogeneous solutions away from the boundary of the domain for a variety of different reaction kinetics, with a prescribed uniform state near the boundary. We show that these boundary conditions can be derived from a larger heterogeneous field, indicating that these conditions can arise naturally if cell signalling or other properties of the medium vary in space. We explain the basic mechanisms behind this pattern localization, and demonstrate that it can capture a large range of localized patterning in one, two, and three dimensions, and that this framework can be applied to systems involving more than two species. Furthermore, the boundary conditions proposed lead to more symmetrical patterns on the interior of the domain, and plausibly capture more realistic boundaries in developmental systems. Finally, we show that these isolated patterns are more robust to fluctuations in initial conditions, and that they allow intriguing possibilities of pattern selection via geometry, distinct from known selection mechanisms.
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Certain two-component reaction-diffusion systems on a finite interval are known to possess mesa (box-like) steadystate patterns in the singularly perturbed limit of small diffusivity for one of the two solution components. As the diffusivity D of the second component is decreased below some critical value Dc, with Dc = O(1), the existence of a steady-state mesa pattern is lost, triggering the onset of a mesa self-replication event that ultimately leads to the creation of additional mesas. The initiation of this phenomena is studied in detail for a particular scaling limit of the Brusselator model. Near the existence threshold Dc of a single steady-state mesa, it is shown that an internal layer forms in the center of the mesa. The structure of the solution within this internal layer is shown to be governed by a certain core problem, comprised of a single non-autonomous second-order ODE. By analyzing this core problem using rigorous and formal asymptotic methods, and by using the Singular Limit Eigenvalue Problem (SLEP) method to asymptotically calculate small eigenvalues, an analytical verification of the conditions of Nishiura and Ueyema [Physica D, 130, No. 1, (1999), pp. 73-104], believed to be responsible for self-replication, is given. These conditions include: (1) The existence of a saddle-node threshold at which the steady-state mesa pattern disappears; (2) the dimple-shaped eigenfunction at the threshold, believed to be responsible for the initiation of the replication process; and (3) the stability of the mesa pattern above the existence threshold. Finally, we show that the core problem is universal in the sense that it pertains to a class of reaction-diffusion systems, including the Gierer-Meinhardt model with saturation, where mesa self-replication also occurs.
Robustness of spatial pattern against perturbations is an indispensable property of developmental processes for organisms, which need to adapt to changing environments. Although specific mechanisms for this robustness have been extensively investigated, little is known about a general mechanism for achieving robustness in reaction-diffusion systems. Here, we propose a buffered reaction-diffusion system, in which active states of chemicals mediated by buffer molecules contribute to reactions, and demonstrate that robustness of the pattern wavelength is achieved by the dynamics of the buffer molecule. This robustness is analytically explained as a result of the scaling properties of the buffered system, which also lead to a reciprocal relationship between the wavelengths robustness and the plasticity of the spatial phase upon external perturbations. Finally, we explore the relevance of this reciprocity to biological systems.
Wavelength selection in reaction--diffusion systems can be understood as a coarsening process that is interrupted by counteracting processes at certain wavelengths. We first show that coarsening in mass-conserving systems is driven by self-amplifying mass transport between neighboring high-density domains. We derive a general coarsening criterion and show that coarsening is generically uninterrupted in two-component systems that conserve mass. The theory is then generalized to study interrupted coarsening and anti-coarsening due to weakly-broken mass conservation, providing a general path to analyze wavelength selection in pattern formation far from equilibrium.
In the nearly seven decades since the publication of Alan Turings work on morphogenesis, enormous progress has been made in understanding both the mathematical and biological aspects of his proposed reaction-diffusion theory. Some of these developments were nascent in Turings paper, and others have been due to new insights from modern mathematical techniques, advances in numerical simulations, and extensive biological experiments. Despite such progress, there are still important gaps between theory and experiment, with many examples of biological patterning where the underlying mechanisms are still unclear. Here we review modern developments in the mathematical theory pioneered by Turing, showing how his approach has been generalized to a range of settings beyond the classical two-species reaction-diffusion framework, including evolving and complex manifolds, systems heterogeneous in space and time, and more general reaction-transport equations. While substantial progress has been made in understanding these more complicated models, there are many remaining challenges that we highlight throughout. We focus on the mathematical theory, and in particular linear stability analysis of `trivial base states. We emphasise important open questions in developing this theory further, and discuss obstacles in using these techniques to understand biological reality.
This paper proposes the Ricci-flow equation from Riemannian geometry as a general geometric framework for various nonlinear reaction-diffusion systems (and related dissipative solitons) in mathematical biology. More precisely, we propose a conjecture that any kind of reaction-diffusion processes in biology, chemistry and physics can be modelled by the combined geometric-diffusion system. In order to demonstrate the validity of this hypothesis, we review a number of popular nonlinear reaction-diffusion systems and try to show that they can all be subsumed by the presented geometric framework of the Ricci flow. Keywords: geometrical Ricci flow, nonlinear reaction-diffusion, dissipative solitons and breathers
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