Turing Patterns

Self-organization in physical systems: rhythms, patterns and chaos

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Turing Patterns by Miriam Daeubler

Existence of Turing Patterns and different types of patterns will be discussed

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2 3 4 5 6

Short Introduction to Turing Patterns

Linear Stability Analysis Existence of Turing Patterns Types of Turing Patterns in a System Summary and Outlook Literature

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Turing Patterns by Miriam Daeubler

Turing patterns can be used to describe evolution of cat pelts

Leopard (left) and jaguar (right) pelt patterns at different ages of the animal

Model not proven experimentally

Source:

R. T. Liu, S. S. Liaw, and P. K. Maini, Physical Review E 74, 011914 (2006)

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Turing Patterns by Miriam Daeubler

Turing patterns are topologically different, within one morphology different typical length scales exist
Experimentally observed patterns in chlorite-iodine-malonic acid reactions

Source:

Ouyang/Swinney, Transition from a uniform state hexagonal and striped Turing patterns, Nature, Vol. 352, 610-611, 1991

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Turing Patterns by Miriam Daeubler

The Turing model originally derived to explain morphogenesis in biological systems

Some facts about Turing patterns
Model systems of two reacting and diffusing substances with concentrations u, v = , + = , + The above system describes an activator-inhibitor system Activator: substance that stimulates the growth of the concentration of both chemicals Inhibitor: substance that leads to a decrease in the concentrations In 1952, Turing predicted that such a system can produce a stationary pattern, if the inhibitor diffuses much faster than the activator (local activation with lateral inhibition). Turing tried to explain morphogenesis, i.e. the development of shape or form in plants and animals.

24.11.2010

Turing Patterns by Miriam Daeubler

Existence of Turing Patterns and different types of patterns will be discussed

1
2 3 4 5 6

Short Introduction to Turing Patterns

Linear Stability Analysis Existence of Turing Patterns Types of Turing Patterns in a System Summary and Outlook Literature

2
6 11 20 26 29

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Turing Patterns by Miriam Daeubler

A diagonalizable coefficient matrix makes system of linear differential equations easy to handle
A homogeneous linear system of ODEs with constant coefficients
= (), () For further discussion, assume that is diagonalizable The eigenvalues are obtained by solving the secular equation det = 0 In order to construct fundamental system choose eigenbasis of , thus = ( 1 1 , . . , ), eigenvalues The solution of the homogeneous system is

= =

, =1

Usually, one is only interested in real solutions. Complex eigenvalues and eigenvectors always come in pairs, therefore, both can be combined to obtain only real solutions.

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Turing Patterns by Miriam Daeubler

Thanks to coordinate transform stability analysis only needs to be derived once

Stability analysis in the simplest linear system (1/2)
Consider the following system with constant coefficients = + , () For further discussion, assume that is diagonalizable Let be a special solution of interest, usually a fixed point of the system Transform coordinates such that = = ()

Thus, it is only necessary to assess the stability of = 0.

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Turing Patterns by Miriam Daeubler

Stability analysis reduces to an eigenvalue problem

Stability analysis in the simplest linear system (2/2)
The stability is described by the initial value problem = and 0 = with the solution = (0 )1 Theorem The solution = 0 is a) strictly stable, if and only if all eigenvalues of A have a real part smaller than zero: < 0 b) stable, if and only if all eigenvalues of A have a real part smaller or equal zero: 0 c) unstable, in every other case

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Turing Patterns by Miriam Daeubler

Under certain conditions even the stability analysis of a nonlinear system reduces to an eigenvalue problem
Stability analysis in the nonlinear case
Consider the following nonlinear system of differential equations with being a nonlinear function which can be continuously differentiated a sufficient number of times

= ( )
Assume = 0 is a fixed point with 0 = 0. Using Taylor expansion one obtains

= + ( ) = (0), 0 = 0 = ( )
It can be shown that if the shortened system of differential equations = is either strictly stable or unstable so is the nonlinear system.

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Turing Patterns by Miriam Daeubler

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Existence of Turing Patterns and different types of patterns will be discussed

1
2 3 4 5 6

Short Introduction to Turing Patterns

Linear Stability Analysis Existence of Turing Patterns Types of Turing Patterns in a System Summary and Outlook Literature

2
6 11 20 26 29

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Turing Patterns by Miriam Daeubler

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The possible existence of Turing patterns can be derived using stability analysis
Stability analysis of Turing model (1/7)
Let there be a steady, spatially homogeneous solution = 0 (fixed point)

= , + = , + Firstly, linearize the system of equations in the vicinity of the fixed point

+ =
With = c=
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0 0

Signs of a, b, c and d chooses in such way that u corresponds to the activator, v to the inhibitor

, = , =

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The instability shall be diffusion driven, the system must be stable in the absence of diffusion
Stability analysis of Turing model (2/7)
Neglecting the diffusion terms the linearized system becomes =

In order to test stability, one must solve the eigenvalue problem = 0 1 = + 2 det For the fixed point to be strictly stable < 0. Thus, < <

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Turing Patterns by Miriam Daeubler

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In order to reduce the complexity of the PDE system use Fourier integral transform
Stability analysis of Turing model (3/7)
Linearizing one obtained a system of PDEs + = +

Integral transforms are standard way to reduce complex PDEs to simpler PDEs or even ODEs, here use Fourier transform = = ( , ) 3 ( , ) 3
Fourier transforms describe a change in basis, new basis consists of eigenfunctions of Laplace operator

Fourier transforms of a function only exist if it is absolutely integrable ( , ) 3 <

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Properties of Fourier transforms needed to derive system of differential equations in Fourier space
Stability analysis of Turing model (4/7)
Fourier transform of gradient of a function

, 3 =
=

, 3

, 3

, 3 = (, )

Fourier transform of Laplace of a function , 3 = = , 3 , 3 = 2 (, )

, 3 = ,

Fourier transform of derivative of a function with respect to time , 3 1 = lim , + , 0

= lim
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1 0

, + (, ) =

(, )
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Turing Patterns by Miriam Daeubler

Fourier transforms reduce PDEs to ODEs and stability analysis becomes an eigenvalue problem
Stability analysis of Turing model (5/7)
Fourier transform the system of PDEs 3 = 3 = + + 3

3 +

2 = 2 2 = 2

Fourier transforms reduce PDEs to ODEs and stability analysis becomes an eigenvalue problem in Fourier space
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For a pattern to evolve some spatial modes must be unstable

Stability analysis of Turing model (6/7)
Fourier transforming the linearized system of differential equations yields 2 = 2

To find a criterion for unstable spatial modes find eigenvalues 2 = 0 2 2 + + + 2 + 2 + 2 2 + + = 0 Compare with = 0 2 + + = 0

and
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2 + + = 0 =
Turing Patterns by Miriam Daeubler

2 2

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Condition for instability to occur linked with typical length scales

Stability analysis of Turing model (7/7)
To sum up, the only possibility for an instability is for the product of the roots to be negative ( 2 ) 2 2 + + < 0

The Turing instability occurs as long as > and the minimum of 2 is

negative. First requirement can be rewritten

>

>

Source:

Hoyle, Pattern Formation, Cambridge University Press, 2006, p.20

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Nonlinear bifurcation theory has to be used to predict morphologies

Results obtained from stability analysis
Stability analysis the parameter range for which instabilities are present the characteristic length of the resulting patterns, which are independent of the initial conditions, therefore intrinsic to the system initial conditions only influence the phase of the pattern, i.e. position and alignment of pattern entities the different possible morphologies of the patterns the selection rules for multistable systems

Apply nonlinear bifurcation theory to approximately predict the stability of different Turing patterns

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Turing Patterns by Miriam Daeubler

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Existence of Turing Patterns and different types of patterns will be discussed

1
2 3 4 5 6

Short Introduction to Turing Patterns

Linear Stability Analysis Existence of Turing Patterns Types of Turing Patterns in a System Summary and Outlook Literature

2
6 11 20 26 29

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Turing Patterns by Miriam Daeubler

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Bifurcations describe sudden qualitative changes in the set of solutions of a system as parameters are varied
Bifurcation theory in a nutshell (1/2)
Consider the following nonlinear system of differential equations with being a nonlinear function which can be continuously differentiated a sufficient number of times

= ( )
The solutions of the system define a flow , with 0 , 0 = 0 . Definition bifurcation

At a bifurcation there is a sudden qualitative change in the flow in response to infinitesimal changes in one or more parameters of the system of differential equations. The phase portrait, the number and stability of fixed points or periodic orbits is usually affected.
Local vs. global bifurcation Local bifurcation: changes of the flow in the vicinity of a fixed point or periodic orbit Global bifurcation: changes that affect the large scale property of the flow
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For Turing patterns local bifurcations are of interest

Bifurcation theory in a nutshell (2/2)
Local bifurcations A local bifurcation occurs when a parameter change causes the stability of a fixed point or periodic orbit to change. This corresponds to the real part of an eigenvalue of a fixed point or PO passing through zero. Goal of bifurcation theory Bifurcation theory tries to produce parameter space maps or bifurcation diagrams that divide the parameter space into regions of topologically equivalent systems, in our case equivalent Turing pattern morphologies.

Source:

Ott, Chaos in Dynamical Systems, Second Edition, Cambridge University Press, 2002, p. 46

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Turing Patterns by Miriam Daeubler

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Stripes and hexagonal spotty pattern typical for 2D reaction-diffusion systems

Typical patterns of 2D reaction-diffusion systems

Source:

Leppnen, The theory of Turing pattern formation, Helsinki University of Technology, 2004

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Turing Patterns by Miriam Daeubler

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Bifurcation theory able to predict stable morphologies found in numerical simulations of 2D system
Bifurcation theory applied to Turing patterns: 2D hexagonal lattice

2D hexagonal lattice is a bistable system

Source:

Leppnen, The theory of Turing pattern formation, Helsinki University of Technology, 2004

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Turing Patterns by Miriam Daeubler

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Turing patterns discussed theoretically beforehand have been observed experimentally

Experimentally observed patterns in chlorite-iodine-malonic acid reactions

Source:

Ouyang/Swinney, Transition from a uniform state hexagonal and striped Turing patterns, Nature, Vol. 352, 610-611, 1991

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Turing Patterns by Miriam Daeubler

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Existence of Turing Patterns and different types of patterns will be discussed

1
2 3 4 5 6

Short Introduction to Turing Patterns

Linear Stability Analysis Existence of Turing Patterns Types of Turing Patterns in a System Summary and Outlook Literature

2
6 11 20 26 29

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Turing Patterns by Miriam Daeubler

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There is no general way to determine which state a morphologically multistable system will choose
Results obtained from stability analysis and nonlinear bifurcation theory
Stability analysis and nonlinear bifurcation theory the parameter range for which instabilities are present the characteristic length of the resulting patterns, which are independent of the initial conditions, therefore intrinsic to the system initial conditions only influence the phase of the pattern, i.e. position and alignment of pattern entities the different possible morphologies of the patterns the selection rules for multistable systems

Formal theory of self-organization based on non-equilibrium thermodynamics, however, this theory is incomplete There is no general way to figure out the selection rules for multistable systems

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Besides missing selection rules there are many more topics concerning Turing patterns currently analyzed
Further topics concerning Turing patterns
Turing patterns also appear in 3D systems Turing patterns can be generalized to allow for transport mechanisms other than diffusion Interaction between Turing instability (constant in time) and other instabilities such as Hopf instability (constant in space) Spatially correlated noise can induce Turing patterns in systems that under normal conditions would not exhibit any

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Turing Patterns by Miriam Daeubler

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Existence of Turing Patterns and different types of patterns will be discussed

1
2 3 4 5 6

Short Introduction to Turing Patterns

Linear Stability Analysis Existence of Turing Patterns Types of Turing Patterns in a System Summary and Outlook Literature

2
6 11 20 26 29

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Turing Patterns by Miriam Daeubler

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Literature used
1) Hoyle, Pattern Formation: An Introduction to methods, Cambridge University Press, 2006 2) Ott, Chaos in Dynamical Systems, Second Edition, Cambridge University Press, 2002 3) Leppnen et al., Spatio-temporal dynamics in a Turing model, submitted to InterJournal 2004 4) Sanz-Anchelergues et al., Turing pattern formation induced by spatially correlated noise, Phys. Rev. E, vol. 63, 056124 5) Guckenheimer, Bifurcation, doi: 10.4249/scholarpedia.1517 6) Leppnen et al., A new dimension to Turing patterns, Physica D, 168-169 (2002) 35-44 7) Leppnen et al., Dimensionality effects in Turing pattern formation, arXiv:condmat/0306121v1, 2003 8) Nakao/Mikhailov, Turing patterns in network-organized activator-inhibitor systems, doi: 10.1038/NPHYS1651 9) Gierer/Meinhardt, A Theory of Biological Pattern Formation, Kybernetik 12, 30-39 (1972) 10) Leppnen et al., Morphological transitions and bistability in Turing systems, Phys. Rev. E, vol. 70, 066202 (2004) 11) Leppnen, The theory of Turing pattern formation, Helsinki University of Technology, 2004

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Literature used
12) Ouyang/Swinney, Transition from a uniform state hexagonal and striped Turing patterns, Nature, Vol. 352, 610-611, 1991 13) Leppnen et al., Turing Systems as Models of Complex Pattern Formation, Brazil Journal of Physics, vol. 34, No. 2A, June 2004 14) Movie: Turing-Hopf bifurcation in monostable system downloaded from http://www.apmaths.uwo.ca/~mkarttu/turing.shtml 15) Movie: Self-organization of spherical structures under Gaussian noise in 3D downloaded from http://www.apmaths.uwo.ca/~mkarttu/turing.shtml

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Backup

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In order to solve a system of linear differential equations one needs to find the homogeneous solution first
General remarks on systems of linear differential equations
Definition of an explicit system of linear first order differential equations = + with

Systems of higher order can always be reduced to a system of first order

and () continuous functions of time .

General solution of an explicit system of linear first order differential equations

= +
being a particular solution of the inhomogeneous differential equation being any solution of the homogeneous differential equation Study solutions of homogenous system of differential equations
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One needs to construct a basis of the vector space spanned by the solutions
Solutions of a system of homogeneous linear differential equations
The solutions of the homogeneous linear system of differential equations form a subspace of 1 , , the real vector space of all continuously differentiable functions , with finite dimension.

Receipe to construct a basis of this subspace for an initial value problem

Choose 0 and a basis , 1, . . , of Solve = (), 0 = ( 1 , . . , )

The matrix () is called a fundamental system.

Then the general solution of the homogeneous equation is = =

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=1 (),

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Many methods enable one to obtain a particular solution of the inhomogeneous system
Solutions of a system of inhomogeneous linear differential equations
First a construct a basis of the solutions of the homogeneous differential equations () Use one of the abundant methods to obtain a solution for the inhomogeneous system of differential equations for initial value problems, such as Variation of constants Greens functions

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