Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor...
Transcript of Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor...
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Examples for Pattern Formation in Biological Systems
Nils GoerkeDepartment of Computer Science VIUniversity of Bonn
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Pattern Formation
• Cellular Automata (CA)
• Langton's Loop, Langton's Ant• Self Organising Criticality (SOC)
• Lindenmeyer systems
• Iterated functions
• Differential equations• Population dynamics
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Pattern Formation• Iterated functions
• Linear and exponential growth
• Fibonacci sequence
• Logistic growth
• Predator-prey system
• Lotka-Volterra equations
• Activator-inhibitor equations
• Reaction-diffusion systems
• Plant morphogenesis, phyllotaxis
• Golden section, Golden angle
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Iterated Functions
Iterated functions are closely related to sequences.
Sequence of values:
{ x(1), x(2), x(3), x(4), x(5), ... }
Sequences can be generated using recursive defined or iterated functions:
x(i+1) = f ( x(i) )
In iterated functions the next value x(i+1) is a function of the predecessing value x(i).
Iterated functions need an intitial value x(0) .
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Iterated Functions, examples
Examples for iterated functions:
x(i+1) = x(i) + 2 with x(0)=3{ 3, 5, 7, 9, 11, .... }
x(i+1) = a x(i) with x(0)=100.0, a=0.9
{ 100.00, 90.00, 81.00, 72.90, 65.61, 59.049, ... }
x(i+1) = (1+) x(i) with x(0)>0, and 0 1
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Iterated Functions, examples 2
The definition of iterated functions can contain logic functions as well: ½ x(i) if x(i) is even
x(i+1) =
3 x(i) +1 else{Can you predict the final value of x(i) when the starting value x(0) is given ?
Play this iterated function as a two person game: start with x(0) and iterate the function by turns:You lose, if you end up with x(i)=1.
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Linear Growth
Implementing linear growth of a population of individuals is easy with iterated functions:
x(i+1) = x(i) + c with x(0)>0, and c >0
time titeration i
x(t)
* * * * * * *
x(0)
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Linear Growth
Linear growth of a population of individuals is easily implemented using a constant increase c:
x(i+1) = x(i) + c with x(0)>0, and c >0
time titeration i
x(t)
* * * * * * *
x(0)
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Exponential Growth
To obtain exponential growth, the increase is proportional to the number of individuals (birthrate).
x(i+1) = x(i) + rr x(i) with x(0)>0 = ( 1 + r ) x(i) with b=1+r = b x(i)
Depending on b a different behaviour results: 0.0 < b < 1.0 exponential decay
b = 1.0 constant, stable population b > 1.0 exponential growth
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Exponential Growth
To obtain exponential growth, the increase is proportional to the number of individuals (birthrate).
x(i+1) = b x(i) with x(0)>0
x(i)
*x(0)
i
x(i)
*
x(i)
*x(0)
i i
x(0)
0.0 < b < 1.0 b = 1.0 b > 1.0
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci SequenceLeonardo of Pisa, also known as Fibonacci, has mentioned in 1202 AD a recursive sequence that has been inspired by a growing population (e.g. number of pairs of rabbits, unbounded reproduction).
x(i+1) = x(i) + x(i-1)
x(0)=0, x(1)=1
{ 0, 1, 1, 2, 3, 5, 8, 13, 21, 34, 55, 89, ... }
The sequence (although named after Fibonacci) has been reported to be known before that time,e.g. 200 BC, by Indian mathematicians. From Wikipedia, (9.6.2009)
http://en.wikipedia.org/wiki/Fibonacci_number
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Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci Sequence
From Ron Knott's web pages on Mathematics, Fibonacci Numbers and Nature , 9.6.2009http://www.mcs.surrey.ac.uk/Personal/R.Knott/Fibonacci/fibnat.html
Suppose a newly-born pair of rabbits, one male, one female, are put in a field. Rabbits are able to mate at the age of one month so that at the end of its second month a female can produce another pair of rabbits. Suppose that our rabbits never die and that the female always produces one new pair (one male, one female) every month from the second month on.
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Logistic GrowthUnbounded reproduction is not realistic.Several circumstances exist (e.g. restricted resources) that yield to a maximal, bounded growth rate.
x(i+1) = x(i) + g x(i) with a growth rate g
The growth rate g shall now reflect restricted resources.The available ressources are those ressources that have been left over by the population x(i). Thus g is modeled as a growing rate a that is bounded by
g = a ( M – x(i) ) with M maximum of the ressources
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Artificial Life & Computational Intelligence, Wed. 10. June 2009
Logistic GrowthThus a modified growth formula can be used:
x(i+1) = x(i) + g x(i) with g=a( M-x(i) ) = x(i) + a ( M – x(i) ) x(i)
This iterated function implements the differential equation published by P.-F. Verhulst (1838) to model realistic growth of populations:
From Wikipedia, (9.6.2009)http://en.wikipedia.org/wiki/Logistic_function
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Artificial Life & Computational Intelligence, Wed. 10. June 2009
Logistic GrowthIterating the equation with a small initial value of x(0)
x(i+1) = x(i) + a( M-x(i) ) x(i)
The population starts to increase with an almost exponential growth of x(i). In this phase the increase is getting larger from step to step. This continues until x(i) reaches a value of x(j)=½ M. After that, the increase is getting smaller and smaller; until the value of x(i) converges to the maximal possible value of x(i) = M.
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Artificial Life & Computational Intelligence, Wed. 10. June 2009
Logistic GrowthSolving the Verhulst differential equation yields the logistic function with it`s typical sigmoid shape.
From Wikipedia, (9.6.2009)http://en.wikipedia.org/wiki/Logistic_function
In the field of neural networks one can find this function denoted as:Sigmoid function or Fermi function.
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Predator-Prey SystemConsider a situation with more than one species that interact with each other:
Predator and Preyone species (prey) is chased by the other species (predator).
Both, predator and prey are growing, and both are interacting. The dynamics can be modeled by the following equations:
N(i+1) = N(i) + aN(i) – b N(i)P(i) )
P(i+1) = P(i) + cP(i)N(i) – d P(i)
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Predator-Prey System
N(i+1) = N(i) + aN(i) – b N(i)P(i) ) prey
P(i+1) = P(i) + cP(i)N(i) – d P(i) predator
The number N(i) of prey individuals: N(i) is increasing with the growth factor of a (birth)and is decreasing due to limited ressources bN(i)and due to the number of predators P(i) (hunt).
The number P(i) of predators: P(i) is increasing proportional to the number of predators P(i) (birth) and the number of prey N(i) (food), and is decreasing due to limited ressources defined by d.
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Lotka-Volterra EquationsThe prey-predator equations written as system of two coupled differential equations: Lotka-Volterra (1925, 1926)
∂N/∂t = aN – bPN = N (a – bP) (prey)∂P/∂t = cPN – d P = P (cN – d) (predator)
Several observations from real animal data indicate that the Lotka-Volterra model is describing population dynamics sufficiently:Hudson-Bay Company, 1845 – 1935, lynx, hares.
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Lotka-Volterra Equations
From Wikipedia, (9.6.2009)http://en.wikipedia.org/wiki/Lotka-Volterra_equation
The prey-predator equations from Lotka and Volterra establish a periodic oscillation over time.
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Activator-Inhibitor EquationsThe prey-predator equations are an example of a complete family of differential equations called:
Activator – Inhibitor Systems
Typical Activator – Inhibitor systems have two components that interact with each other. One component is called the Activator, making the system grow.The other component is called the Inhibitor, which is the component that restricts the systems and prevents the growth from getting exponential..
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Activator-Inhibitor Equations
The Activator X enforces it's own growth, and enforces the growth of the inhibitor.
X(i+1) = ... + aX(i) ... Y(i+1) = ... +cX(i) ...
The Inhibitor Y constrains it's own growth, and constrains the growth of the activator.
X(i+1) = ... – bY(i) ... Y(i+1) = ... – dY(i) ...
X(i+1) = X(i) +aX(i) – bY(i) activator
Y(i+1) = Y(i) +cX(i) – dY(i) inhibitor
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Activator-Inhibitor EquationsActivator-Inhibitor characteristics can be found in a lot of systems from biology, physics, medicine and technics.
Very often the Activator-Inhibitor principle is combined with a spatial component.Not only the conditions at one spatial position are considered, but also the conditions from the direct heighborhood are taken into account via a diffusion process.
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Reaction-Diffusion SystemThe differential equations of a Reaction-Diffusion system try to model the local influence from the neighborhood via a diffusion processes and the activity within one compartment using the reaction terms.
dV/dt = μ ∂2V/∂x2 + fV(V,W)
dW/dt = σ ∂2W/∂x2 + fW
(V,W)
Example for a one-dimensional Reaction-Diffusion systemwith the diffusion terms: μ ∂2V/∂x2 and σ ∂2W/∂x2 and the reaction terms: f
V(V,W) and f
W(V,W)
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Reaction-Diffusion SystemCellular automata can be seen as discretised versions of Reaction-Diffusion systems.
The rule of a cellular automaton takes the state of the direct neighborhood into account, thus the diffusion part of a Reaction-Diffusion system can be easily implemented.
0 1 000 111 90D
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Reaction-Diffusion System
http://www.uni-muenster.de/Physik.AP/Purwins/DE/ElNw-de.html
An real implementation of an Activator-Inhibitor Reaction-Diffusion system is a field (1-dim, or 2-dim) of nonlinear electrical oscillators.
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Reaction-Diffusion System
http://www.uni-muenster.de/Physik.AP/Purwins/DE/ElNw-de.html
Spatial structures emerge as result of the coupling.
128 oscillators are connected to a one- dimensional field.
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Reaction-Diffusion System
http://www.uni-muenster.de/Physik.AP/Purwins/DE/ElNw-de.html
Spatial structures are the result of the coupling.
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Reaction-Diffusion System
Spatial structures are the result of the coupling.http://www.uni-muenster.de/Physik.AP/Purwins/DE/ElNw-de.html
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Belousov-Zhabotinsky-Reaction
http://www.scholarpedia.org/article/Belousov-Zhabotinsky_reaction
Example for an Activator-Inhibitor Reaction-Diffusion System from chemistry.
The Belousov-Zhabotinsky reaction yields a spatio-temporal moving pattern.
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Belousov-Zhabotinsky-Reaction
http://www.uni-muenster.de/Physik.AP/Purwins/DE/BZ-Reaktion-de.tml
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Patterns in Mixtures of Granulated Material
http://www.uni-muenster.de/Physik.AP/Purwins/DE/Streifen_im_Salz-Mohn-Gemisch-de.html
A mixture of granulated material with different density consisting of Salt (white) and Poppy Seed (black) can establish a spatial structure.
The nonlinear combination of different densities, the movement and the friction cause this effect.
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Growth PatternsGrowth patterns of plants are often determined by activator-inhibitor reaction-diffusion systems.
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Growth PatternsThe growing plant produces ressources (activator)
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Growth PatternsAs soon as enough ressources (activator) are available, a leaf is growing in that position.
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Growth PatternsThe growing leaf is producing an inhibitor, to prevent that a second leaf will grow near by.
Inhibitor distribution
Activator distribution
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Growth PatternsWith the growing plant, the concentration of activator and inhibitor is decaying.
Inhibitor distribution
Activator distribution
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Growth PatternsAfter a while, enough ressources are available again to produce a new leaf.
Inhibitor distribution
Activator distribution
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Growth PatternsBut the inhibitor is still present; thus the position of the next leaf will be a far avay from the fist one.
Inhibitor distribution
Activator distribution
![Page 40: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/40.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Growth PatternsAgain, the growing leaf will produce inhibitor.
Inhibitor distribution
Activator distribution
![Page 41: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/41.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Growth PatternsThe next leaf to grow is influenced by the inhibitor of both existing leaves, leading to a position in between.
![Page 42: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/42.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Growth PatternsThe next leaf to grow is influenced by the inhibitor of all the existing leaves, leading to a positions in between.
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Growth PatternsAfter a while the angle between subsequent leafs has become almost the golden angle of 137.51 deg..
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Growth Patterns
Growth patterns of plants are often determined by activator-inhibitor reaction-diffusion systems.
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci Spirals in Plant Morphogenesis
http://www.mcs.surrey.ac.uk/Personal/R.Knott/Fibonacci/fibnat.html#pinecones
![Page 46: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/46.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci Spirals in Plant Morphogenesis
http://www.mcs.surrey.ac.uk/Personal/R.Knott/Fibonacci/fibnat.html#pinecones
![Page 47: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/47.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci Spirals in Plant Morphogenesis
http://www.mcs.surrey.ac.uk/Personal/R.Knott/Fibonacci/fibnat.html#pinecones
![Page 48: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/48.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci Spirals in Plant Morphogenesis
http://www.mcs.surrey.ac.uk/Personal/R.Knott/Fibonacci/fibnat.html#pinecones
![Page 49: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/49.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci Spirals in Plant Morphogenesis
http://www.mcs.surrey.ac.uk/Personal/R.Knott/Fibonacci/fibnat.html#pinecones
![Page 50: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/50.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci Spirals in Plant Morphogenesis
From Ron Knott's web pages on Mathematics, Fibonacci Numbers and Nature , 9.6.2009http://www.mcs.surrey.ac.uk/Personal/R.Knott/Fibonacci/fibnat.html#pinecones
The number of spirals that can be found in either direction are typically consequitive Fibonacci numbers.
![Page 51: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/51.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci Spirals in Plant Morphogenesis
http://www.mcs.surrey.ac.uk/Personal/R.Knott/Fibonacci/fibnat.html#pinecones
![Page 52: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/52.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci Spirals in Plant Morphogenesis
http://www.mcs.surrey.ac.uk/Personal/R.Knott/Fibonacci/fibnat.html#pinecones
![Page 53: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/53.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci Spirals in Plant Morphogenesis
http://www.mcs.surrey.ac.uk/Personal/R.Knott/Fibonacci/fibnat.html#pinecones
![Page 54: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/54.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci Spirals in Plant Morphogenesis
![Page 55: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/55.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Fibonacci Spirals in Plant Morphogenesis
![Page 56: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/56.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Self Similarity in Plant Morphogenesis
Brassica oleracea (Romanescu)© Biopix.dk: N Sloth
![Page 57: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/57.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Self Similarity in Plant Morphogenesis
Brassica oleracea (Romanescu)© Biopix.dk: N Sloth
![Page 58: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/58.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Self Similarity in Plant Morphogenesis
Brassica oleracea (Romanescu)© Biopix.dk: N Sloth
![Page 59: Examples for Pattern Formation in Biological Systems · 2009-06-10 · Activator-Inhibitor Equations The prey-predator equations are an example of a complete family of differential](https://reader033.fdocuments.in/reader033/viewer/2022060506/5f1ee37bec59172317353341/html5/thumbnails/59.jpg)
© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Pattern Formation• Iterated functions
• Linear and exponential growth
• Fibonacci sequence
• Logistic growth
• Predator-prey system
• Lotka-Volterra equations
• Activator-inhibitor equations
• Reaction-diffusion systems
• Plant morphogenesis, phyllotaxis
• Golden section, Golden angle
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© Nils Goerke, University Bonn, 6/2009
Artificial Life & Computational Intelligence, Wed. 10. June 2009
Vielen Dank für Ihre die Aufmerksamkeit
Thank you for your attention.
und