Parameters fitted to the Eqs. (13)–(15) for the obtained patterns obtained using Eqs. (7)–(9).
Abstract
Turing demonstrated a coupled reaction-diffusion equation with two components produced steady-state heterogeneous spatial patterns, under certain conditions. The instability found by Turing is now called a diffusion-driven instability or Turing instability. Systems in two dimensions produce spot and stripe patterns, and these systems have been applied as models to explain patterns observed in biological and chemical fields and to develop image information processing tools. Previously, we developed a method that utilizes a reaction-diffusion system with anisotropic diffusion that exhibits triangular patterns, thereby introducing a certain anisotropic strength. In this chapter, we discuss the effects of anisotropic diffusion on the generation of triangular patterns. By defining the statistical index characterizing the spatial patterns, we investigated the parameter range over which the triangular patterns were obtained. We determined the explanatory variable based on the relative distance of the pitchfork bifurcation point between the maximum and minimum anisotropic diffusion functions. Its relevance to diffusion instability is also discussed.
Keywords
- Turing pattern
- reaction-diffusion system
- pitchfork bifurcation
- triangular pattern
- pattern statistics
1. Introduction
The formation of organisms and developmental phenomena are natural self-constructed phenomena. Mathematical and physical methods play an important role in the analysis of self-organizing phenomena, such as the collapse of sandy mountains [1], wind ripples on sand dunes, and snow crystal patterns [1, 2, 3].
For a number of pattern-formation phenomena, the so-called reaction-diffusion (RD) equation has been applied to model self-organizing patterns [1, 3]. In 1952, Alan Turing, a British mathematician well-known for proposing the basic principles of cryptanalysis and computers, proposed one of the most famous explanations for self-organizing phenomena by postulating a mechanism based on the reaction and diffusion of the materials [4]. This mechanism is now known as diffusion-driven instability or Turing instability [1, 2, 3].
Some researchers observed that Turing patterns can be realized in chemical reaction systems [1, 2, 3] and animal skin patterns [5, 6]. These patterns have two dimensions, and stripe, spot, and labyrinthine patterns can be realized in the RD equations [1].
Furthermore, several researchers have developed methods for applying the RD model to image information processing. Admatzky et al. [7] and Ito et al. [8] developed an algorithm for segmenting 2D periodic stripes or spotted images using an RD model. Applying such a mechanism has various advantages; for example, they do not require for noise removal mechanisms.
We applied the image-processing algorithm for pattern formation of outer hair cells in the ear [9, 10] that arranges periodic triangles on the surface of the organ of Corti in the inner ear [11, 12]. To quantify the arrangement of the periodic triangules, we developed an RD model with anisotropic diffusion. In Ref. [10], we explored the conditions for emerging periodic patterns. We found the following explanatory variables: on the heuristic argument of the relative difference in the bifurcation points between the maximum and minimum values of the anisotropic diffusion coefficient. This chapter presents the recent research on periodic triangles as Turing patterns.
2. Model
2.1 Turing instability
Turing [4] proposed the following concept as one of the mathematical mechanisms. The system is expressed as follows:
where
FitzHugh-Nagumo [13] chose the following reaction terms:
where the constants
There are several other choices for reaction terms, such as the Schnaclenberg model [14]. In the following sections, we study cases using the FitzHugh-Nagumo equation [13] given by Eq. (2). However, later, we discuss another choice of the reaction term.
First, we consider the case in which Eq. (1) has only one stable time-independent uniform solution
where the wavenumber
The condition for one of the eigenvalues,
This indicates that
At bifurcation, when
Therefore, in the case of
2.2 Turing patterns
The linear analysis performed in the previous section indicated that instability occured. However, an analysis that incorporates nonlinearities is necessary to investigate the time evolution in the region of
Starting from the random initial distributions shown in Figure 1(a) and (e), the time evolutions of the patterns in a density plot of
Various discussions have been conducted on pattern selection [1, 2, 3, 15, 16]. Some researchers have discussed this from the viewpoint of variational methods (energy) [2, 3, 16]. Others have discussed the transformation of the solution structure on the central manifold [2, 17].
Notably, Eq. (1) is one of the partial differential equations. This system is not immune to the dependence of the obtained pattern on the initial distribution. In other words, a slight modulation in the initial distribution can affect the directionality of the stripes in the resulting stripe pattern or the arrangement of the spots [1, 2, 3].
3. Triangular patterns obtained by RD model with anisotropic diffusion
3.1 RD model with anisotropic diffusion
Kondo and Miura [6] revealed that some of developmental processes are affected by RD systems, as in Eq. (1). It is also important to determine the direction in which organs and tissues self-organize during development [18, 19].
The development of V-shaped bundles for sensorineural hearing in the inner ear periodic triangular pattern area has self-organized in a specific direction based on the surrounding conditions [11, 12]. For the morphometric measurements of V-shaped bundles, we developed a system that applied a self-organizing periodic triangular system using an RD model with the anisotropic diffusion [9]. As the simple diffusion function model, we applied the periodic function depending on the diffusive directions with adjacent reaction cells [9].
In the study by Shoji and Iwamoto [9], we focused on the case of strong enough anisotropy to obtain periodic triangular patterns. Here, we considered the general case as follows
The diffusion coefficient of
where
We would like to note that the flux is proportional to the gradient vector, however the multiplication factor is determined by the angle of the vector. Eq. (8) implies that the diffusivity of
This form of anisotropic diffusion was applied by Kobayashi [20] for self-organizing dendritic crystal. In previous studies [19, 21], we also applied this form of anisotropy to explain the directionality of stripes on fish skin.
Figure 2 shows the time evolution of the two-dimensional distribution of
We start with the random initial distribution shown in Figure 2(a)
3.2 Image process
To examine the structure of the triangular pattern shown in Figure 3(a), the numerical data of
By displaying the intensity
A Fourier analysis are performed for the obtained distribution of
Figure 3(e) shows the Bragg peak intensities for
Using these features, we introduced the triangular clearness
where
4. Conditions for triangular patterns obtained
4.1 Comparison between TC and δ
Using the index
As
4.2 Heuristic index for triangular
Based on a heuristic mathematical argument, we developed the mathematical relations of the obtained periodic triangular patterns. The relations was provided by focusing on the range of the values of the diffusion coefficient function
We show the results when the parameters
When diffusion anisotropy is not introduced
Note that
Figure 4(d-f) summarizes the results for
4.3 Regression analysis
Figure 4(d-f) shows the changes in
Regression analysis was performed using the following function:
where
Figure | Parameters in (13) | ||||
---|---|---|---|---|---|
a | b | r | |||
Figure 4(d) | 7.92 | −1.45 | |||
Figure 4(e) | 11.0 | −1.48 | |||
Figure 4(f) | 2.05 | −0.93 |
The coordinates of the inflection point of the function (13),
The
Therefore, these points were subjected to changes until they reached approximately 21% and 79% for the final
We also performed a trial in which the initial distribution was given in a different manner; however, as mentioned above, we obtained the same values for the parameters
4.4 Range of δ
As seen in the previous section,
Transforming this into the
where
In the case of
5. Discussion
In this section, we explain how the triangular patterns were obtained by numerical simulation of Eqs. (7)-(9) as a two-dimensional Turing pattern. As shown in Figure 3, a triangular pattern was obtained using a complex combination of Fourier modes, including high wavelengths.
In this section, we discuss when the reaction term in (2). We also examined whether the same conclusion holds for other choice of reaction terms. We examined another type of reaction term introduced by Schnackenberg [13]. The model described a simple chemical reaction for glycolysis given by the following reaction terms
where
Moreover, we examined the substance independence of the diffusion anisotropy. We also examined not only the case in which diffusion anisotropy is introduced only into the inhibitor
Thus far, our understanding has not progressed to the point where we can fully describe the mechanism of triangular pattern formation for Turing patterns. However, the conditions under which the triangular pattern is formed and maintained as a motionless fixed pattern, as represented in this chapter, were derived mainly through numerical calculations. We hope that mathematical progress will be achieved in this area.
In contrast, this model was also created to automatically extract triangle images. Therefore, we also hope that this model will be relevant and developed for applications in image analysis application.
Acknowledgments
We thank Prof. M. Mimura and Prof. K. Osaki for helpful comments, and K. Yamada for technical helps.
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