Power spectra methods for a stochastic description of diffusion on deterministically growing domains

A central challenge in developmental biology is understanding the creation of robust spatiotemporal heterogeneity. Generally, the mathematical treatments of biological systems have used continuum, mean-field hypotheses for their constituent parts, which ignores any sources of intrinsic stochastic effects. In this paper we consider a stochastic space-jump process as a description of diffusion, i.e., particles are able to undergo a random walk on a discretized domain. By developing analytical Fourier methods we are able to probe this probabilistic framework, which gives us insight into the patterning potential of diffusive systems. Further, an alternative description of domain growth is introduced, with which we are able to rigorously link the mean-field and stochastic descriptions. Finally, through combining these ideas, it is shown that such stochastic descriptions of diffusion on a deterministically growing domain are able to support the nucleation of states that are far removed from the deterministic mean-field steady state

Stochastic reaction & diffusion on growing domains: understanding the breakdown of robust pattern formation

Many biological patterns, from population densities to animal coat markings, can be thought of as heterogeneous spatiotemporal distributions of mobile agents. Many mathematical models have been proposed to account for the emergence of this complexity, but, in general, they have consisted of deterministic systems of differential equations, which do not take into account the stochastic nature of population interactions. One particular, pertinent criticism of these deterministic systems is that the exhibited patterns can often be highly sensitive to changes in initial conditions, domain geometry, parameter values, etc. Due to this sensitivity, we seek to understand the effects of stochasticity and growth on paradigm biological patterning models. In this paper, we extend spatial Fourier analysis and growing domain mapping techniques to encompass stochastic Turing systems. Through this we find that the stochastic systems are able to realize much richer dynamics than their deterministic counterparts, in that patterns are able to exist outside the standard Turing parameter range. Further, it is seen that the inherent stochasticity in the reactions appears to be more important than the noise generated by growth, when considering which wave modes are excited. Finally, although growth is able to generate robust pattern sequences in the deterministic case, we see that stochastic effects destroy this mechanism for conferring robustness. However, through Fourier analysis we are able to suggest a reason behind this lack of robustness and identify possible mechanisms by which to reclaim it

Influence of stochastic domain growth on pattern nucleation for diffusive systems with internal noise

Numerous mathematical models exploring the emergence of complexity within developmental biology incorporate diffusion as the dominant mechanism of transport. However, self-organizing paradigms can exhibit the biologically undesirable property of extensive sensitivity, as illustrated by the behavior of the French-flag model in response to intrinsic noise and Turing’s model when subjected to fluctuations in initial conditions. Domain growth is known to be a stabilizing factor for the latter, though the interaction of intrinsic noise and domain growth is underexplored, even in the simplest of biophysical settings. Previously, we developed analytical Fourier methods and a description of domain growth that allowed us to characterize the effects of deterministic domain growth on stochastically diffusing systems. In this paper we extend our analysis to encompass stochastically growing domains. This form of growth can be used only to link the meso- and macroscopic domains as the “box-splitting” form of growth on the microscopic scale has an ill-defined thermodynamic limit. The extension is achieved by allowing the simulated particles to undergo random walks on a discretized domain, while stochastically controlling the length of each discretized compartment. Due to the dependence of diffusion on the domain discretization, we find that the description of diffusion cannot be uniquely derived. We apply these analytical methods to two justified descriptions, where it is shown that, under certain conditions, diffusion is able to support a consistent inhomogeneous state that is far removed from the deterministic equilibrium, without additional kinetics. Finally, a logistically growing domain is considered. Not only does this show that we can deal with nonmonotonic descriptions of stochastic growth, but it is also seen that diffusion on a stationary domain produces different effects to diffusion on a domain that is stationary “on average.

Graph-Facilitated Resonant Mode Counting in Stochastic Interaction Networks

Oscillations in a stochastic dynamical system, whose deterministic counterpart has a stable steady state, are a widely reported phenomenon. Traditional methods of finding parameter regimes for stochastically-driven resonances are, however, cumbersome for any but the smallest networks. In this letter we show by example of the Brusselator how to use real root counting algorithms and graph theoretic tools to efficiently determine the number of resonant modes and parameter ranges for stochastic oscillations. We argue that stochastic resonance is a network property by showing that resonant modes only depend on the squared Jacobian matrix $J^2$ , unlike deterministic oscillations which are determined by $J$. By using graph theoretic tools, analysis of stochastic behaviour for larger networks is simplified and chemical reaction networks with multiple resonant modes can be identified easily.Comment: 5 pages, 4 figure

Pattern production through a chiral chasing mechanism

Recent experiments on zebrafish pigmentation suggests that their typical black and white striped skin pattern is made up of a number of interacting chromatophore families. Specifically, two of these cell families have been shown to interact through a nonlocal chasing mechanism, which has previously been modeled using integro-differential equations. We extend this framework to include the experimentally observed fact that the cells often exhibit chiral movement, in that the cells chase, and run away, at angles different to the line connecting their centers. This framework is simplified through the use of multiple small limits leading to a coupled set of partial differential equations which are amenable to Fourier analysis. This analysis results in the production of dispersion relations and necessary conditions for a patterning instability to occur. Beyond the theoretical development and the production of new pattern planiforms we are able to corroborate the experimental hypothesis that the global pigmentation patterns can be dependent on the chirality of the chromatophores

Likely equilibria of stochastic hyperelastic spherical shells and tubes

In large deformations, internally pressurised elastic spherical shells and tubes may undergo a limit-point, or inflation, instability manifested by a rapid transition in which their radii suddenly increase. The possible existence of such an instability depends on the material constitutive model. Here, we revisit this problem in the context of stochastic incompressible hyperelastic materials, and ask the question: what is the probability distribution of stable radially symmetric inflation, such that the internal pressure always increases as the radial stretch increases? For the classic elastic problem, involving isotropic incompressible materials, there is a critical parameter value that strictly separates the cases where inflation instability can occur or not. By contrast, for the stochastic problem, we show that the inherent variability of the probabilistic parameters implies that there is always competition between the two cases. To illustrate this, we draw on published experimental data for rubber, and derive the probability distribution of the corresponding random shear modulus to predict the inflation responses for a spherical shell and a cylindrical tube made of a material characterised by this parameter.Comment: arXiv admin note: text overlap with arXiv:1808.0126

Likely oscillatory motions of stochastic hyperelastic solids

Stochastic homogeneous hyperelastic solids are characterised by strain-energy densities where the parameters are random variables defined by probability density functions. These models allow for the propagation of uncertainties from input data to output quantities of interest. To investigate the effect of probabilistic parameters on predicted mechanical responses, we study radial oscillations of cylindrical and spherical shells of stochastic incompressible isotropic hyperelastic material, formulated as quasi-equilibrated motions where the system is in equilibrium at every time instant. Additionally, we study finite shear oscillations of a cuboid, which are not quasi-equilibrated. We find that, for hyperelastic bodies of stochastic neo-Hookean or Mooney-Rivlin material, the amplitude and period of the oscillations follow probability distributions that can be characterised. Further, for cylindrical tubes and spherical shells, when an impulse surface traction is applied, there is a parameter interval where the oscillatory and non-oscillatory motions compete, in the sense that both have a chance to occur with a given probability. We refer to the dynamic evolution of these elastic systems, which exhibit inherent uncertainties due to the material properties, as `likely oscillatory motions'

Effects of intrinsic stochasticity on delayed reaction-diffusion patterning systems

Cellular gene expression is a complex process involving many steps, including the transcription of DNA and translation of mRNA; hence the synthesis of proteins requires a considerable amount of time, from ten minutes to several hours. Since diffusion-driven instability has been observed to be sensitive to perturbations in kinetic delays, the application of Turing patterning mechanisms to the problem of producing spatially heterogeneous differential gene expression has been questioned. In deterministic systems a small delay in the reactions can cause a large increase in the time it takes a system to pattern. Recently, it has been observed that in undelayed systems intrinsic stochasticity can cause pattern initiation to occur earlier than in the analogous deterministic simulations. Here we are interested in adding both stochasticity and delays to Turing systems in order to assess whether stochasticity can reduce the patterning time scale in delayed Turing systems. As analytical insights to this problem are difficult to attain and often limited in their use, we focus on stochastically simulating delayed systems. We consider four different Turing systems and two different forms of delay. Our results are mixed and lead to the conclusion that, although the sensitivity to delays in the Turing mechanism is not completely removed by the addition of intrinsic noise, the effects of the delays are clearly ameliorated in certain specific cases

Non-linear effects on Turing patterns: time oscillations and chaos.

We show that a model reaction-diffusion system with two species in a monostable regime and over a large region of parameter space, produces Turing patterns coexisting with a limit cycle which cannot be discerned from the linear analysis. As a consequence, Turing patterns oscillate in time, a phenomenon which is expected to occur only in a three morphogen system. When varying a single parameter, a series of bifurcations lead to period doubling, quasi-periodic and chaotic oscillations without modifying the underlying Turing pattern. A Ruelle-Takens-Newhouse route to chaos is identified. We also examined the Turing conditions for obtaining a diffusion driven instability and discovered that the patterns obtained are not necessarily stationary for certain values of the diffusion coefficients. All this results demonstrates the limitations of the linear analysis for reaction-diffusion systems