No Arabic abstract
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.
Experimental studies of protein-pattern formation have stimulated new interest in the dynamics of reaction-diffusion systems. However, a comprehensive theoretical understanding of the dynamics of such highly nonlinear, spatially extended systems is still missing. Here we show how a description in phase space, which has proven invaluable in shaping our intuition about the dynamics of nonlinear ordinary differential equations, can be generalized to mass-conserving reaction-diffusion (McRD) systems. We present a comprehensive analysis of two-component McRD systems, which serve as paradigmatic minimal systems that encapsulate the core principles and concepts of the local equilibria theory introduced in the paper. The key insight underlying this theory is that shifting local (reactive) equilibria -- controlled by the local total density -- give rise to concentration gradients that drive diffusive redistribution of total density. We show how this dynamic interplay can be embedded in the phase plane of the reaction kinetics in terms of simple geometric objects: the reactive nullcline and the diffusive flux-balance subspace. On this phase-space level, physical insight can be gained from geometric criteria and graphical constructions. The effects of nonlinearities on the global dynamics are simply encoded in the curved shape of the reactive nullcline. In particular, we show that the pattern-forming `Turing instability in McRD systems is a mass-redistribution instability, and that the features and bifurcations of patterns can be characterized based on regional dispersion relations, associated to distinct spatial regions (plateaus and interfaces) of the patterns. In an extensive outlook section, we detail concrete approaches to generalize local equilibria theory in several directions, including systems with more than two-components, weakly-broken mass conservation, and active matter systems.
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.
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.
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
Rippling patterns of myxobacteria appear in starving colonies before they aggregate to form fruiting bodies. These periodic traveling cell density waves arise from the coordination of individual cell reversals, resulting from an internal clock regulating them, and from contact signaling during bacterial collisions. Here we revisit a mathematical model of rippling in myxobacteria due to Igoshin et al. [Proc. Natl. Acad. Sci. USA {bf 98}, 14913 (2001) and Phys. Rev. E {bf 70}, 041911 (2004)]. Bacteria in this model are phase oscillators with an extra internal phase through which they are coupled to a mean-field of oppositely moving bacteria. Previously, patterns for this model were obtained only by numerical methods and it was not possible to find their wavenumber analytically. We derive an evolution equation for the reversal point density that selects the pattern wavenumber in the weak signaling limit, show the validity of the selection rule by solving numerically the model equations and describe other stable patterns in the strong signaling limit. The nonlocal mean-field coupling tends to decohere and confine patterns. Under appropriate circumstances, it can annihilate the patterns leaving a constant density state via a nonequilibrium phase transition reminiscent of destruction of synchronization in the Kuramoto model.