Part 2D: Pattern Formation 2013/2/1 Differentiation & Pattern Formation • A central problem in development: How do cells differentiate to fulfill different purposes? • How do complex systems generate spatial & temporal structure? • CAs are natural models of intercellular communication B. Pattern Formation 2013/2/1 1 2013/2/1 Zebra 2013/2/1 figs. from Camazine & al.: Self-Org. Biol. Sys. photos ©2000, S. Cazamine 2 Vermiculated Rabbit Fish 3 2013/2/1 Activation & Inhibition in Pattern Formation figs. from Camazine & al.: Self-Org. Biol. Sys. 4 Interaction Parameters • Color patterns typically have a characteristic length scale • Independent of cell size and animal size • Achieved by: – short-range activation ⇒ local uniformity – long-range inhibition ⇒ separation 2013/2/1 • R1 and R2 are the interaction ranges • J1 and J2 are the interaction strengths 5 2013/2/1 6 1 Part 2D: Pattern Formation 2013/2/1 CA Activation/Inhibition Model • • • • Let states si ∈ {–1, +1} and h be a bias parameter and rij be the distance between cells i and j Then the state update rule is: Demonstration of NetLogo Program for Activation/Inhibition Pattern Formation: Fur $ ' si ( t + 1) = sign&h + J1 ∑ s j ( t ) + J 2 ∑ s j ( t )) &% )( rij <R1 R1 ≤rij <R 2 2013/2/1 RunAICA.nlogo 7 2013/2/1 8 € Effect of Bias Example (h = –6, –3, –1; 1, 3, 6) (R1=1, R2=6, J1=1, J2=–0.1, h=0) 2013/2/1 figs. from Bar-Yam 9 Effect of Interaction Ranges R2 = 6 R1 = 1 h = 0 2013/2/1 10 • How can a system using strictly local interactions discriminate between states at long and short range? • E.g. cells in developing organism • Can use two different morphogens diffusing at two different rates R2 = 6 R1 = 1.5 h = –3 figs. from Bar-Yam figs. from Bar-Yam Differential Interaction Ranges R2 = 8 R1 = 1 h = 0 R2 = 6 R1 = 1.5 h = 0 2013/2/1 – activator diffuses slowly (short range) – inhibitor diffuses rapidly (long range) 11 2013/2/1 12 2 Part 2D: Pattern Formation 2013/2/1 Digression on Diffusion Reaction-Diffusion System • Simple 2-D diffusion equation: A ( x, y) = D∇ 2 A ( x, y) • Recall the 2-D Laplacian: ∂ 2 A( x, y ) ∂ 2 A( x, y ) ∇ 2 A( x, y ) = + ∂x 2 ∂y 2 • The Laplacian (like 2nd derivative) is: € diffusion ! ∂ ! A $ # DA # &= ∂ t " I % #" 0 0 $! ∇ 2 A &# DI &%#" ∇ 2 I $ ! fA ( A, I ) &+# & # f A, I % " I( ) reaction $ & & % # A& c˙ = D∇ 2c + f (c), where c = % ( $I' – positive in a local minimum – negative in a local maximum 2013/2/1 ∂A = DA ∇ 2 A + fA ( A, I ) ∂t ∂I = DI ∇ 2 I + fI ( A, I ) ∂t 13 2013/2/1 14 € Framework for Complexity General Reaction-Diffusion System # n & ∂ci = ∑ kαν iα % ∏ ckmkα ( − ∇ ⋅ ji ∂t $ k=1 ' α where ji = µi ci − div Di ci (flux) • change = source terms + transport terms • source terms = local coupling = interactions local to a small region • transport terms = spatial coupling = interactions with contiguous regions = advection + diffusion where kα = rate constant for reaction α and ν iα = stoichiometric coefficient and mkα = a non-negative integer and µi = drift vector – advection: non-dissipative, time-reversible – diffusion: dissipative, irreversible and Di = diffusivity matrix where div Dc = ∑ e j ∑ D jk ∂c 2013/2/1 j k ∂xk 15 2013/2/1 Continuous-time Activator-Inhibitor System NetLogo Simulation of Reaction-Diffusion System • Activator A and inhibitor I may diffuse at different rates in x and y directions • Cell becomes more active if activator + bias exceeds inhibitor • Otherwise, less active ∂A ∂2A ∂2A = dAx 2 + dAy 2 + kA ( A + B − I ) (1− A) ∂t ∂x ∂y ∂I ∂ 2I ∂ 2I = dIx 2 + dIy 2 + kI ( A + B − I ) (1− I ) ∂t ∂x ∂y 2013/2/1 16 17 1. Diffuse activator in X and Y directions 2. Diffuse inhibitor in X and Y directions 3. Each patch performs: stimulation = bias + activator – inhibitor + noise if stimulation > 0 then set activator and inhibitor to 100 else set activator and inhibitor to 0 2013/2/1 18 3 Part 2D: Pattern Formation 2013/2/1 Demonstration of NetLogo Program for Activator/Inhibitor Pattern Formation Demonstration of NetLogo Program for Activator/Inhibitor Pattern Formation with Continuous State Change Run Pattern.nlogo Run Activator-Inhibitor.nlogo 2013/2/1 19 2013/2/1 20 A Key Element of Self-Organization Turing Patterns • Alan Turing studied the mathematics of reaction-diffusion systems • Activation vs. Inhibition • Turing, A. (1952). The chemical basis of morphogenesis. Philosophical Transactions of the Royal Society B 237: 37–72. • Amplification vs. Stabilization • Cooperation vs. Competition • Positive Feedback vs. Negative Feedback – Positive feedback creates • The resulting patterns are known as Turing patterns 2013/2/1 • Growth vs. Limit – Negative feedback shapes 21 Reaction-Diffusion Computing 2013/2/1 22 Image Processing in BZ Medium • Has been used for image processing – diffusion ⇒ noise filtering – reaction ⇒ contrast enhancement • Depending on parameters, RD computing can: – restore broken contours – detect edges – improve contrast 2013/2/1 • (A) boundary detection, (B) contour enhancement, (C) shape enhancement, (D) feature enhancement 23 2013/2/1 Image < Adamatzky, Comp. in Nonlinear Media & Autom. Coll. 24 4 Part 2D: Pattern Formation 2013/2/1 Voronoi Diagrams Some Uses of Voronoi Diagrams • Given a set of generating points: • Construct a polygon around each generating point of set, so all points in a polygon are closer to its generating point than to any other generating points. 2013/2/1 Image < Adamatzky & al., Reaction-Diffusion Computers 25 Computation of Voronoi Diagram by Reaction-Diffusion Processor 2013/2/1 Image < Adamatzky & al., Reaction-Diffusion Computers 27 Path Planning via BZ medium: No Obstacles 2013/2/1 Image < Adamatzky & al., Reaction-Diffusion Computers • Collision-free path planning • Determination of service areas for power substations • Nearest-neighbor pattern classification • Determination of largest empty figure 2013/2/1 26 Mixed Cell Voronoi Diagram 2013/2/1 Image < Adamatzky & al., Reaction-Diffusion Computers 28 Path Planning via BZ medium: Circular Obstacles 29 2013/2/1 Image < Adamatzky & al., Reaction-Diffusion Computers 30 5 Part 2D: Pattern Formation 2013/2/1 Mobile Robot with Onboard Chemical Reactor 2013/2/1 Image < Adamatzky & al., Reaction-Diffusion Computers Actual Path: Pd Processor 31 2013/2/1 Actual Path: Pd Processor 2013/2/1 Image < Adamatzky & al., Reaction-Diffusion Computers Image < Adamatzky & al., Reaction-Diffusion Computers 32 Actual Path: BZ Processor 33 2013/2/1 Image < Adamatzky & al., Reaction-Diffusion Computers 34 Bibliography for Reaction-Diffusion Computing 1. Adamatzky, Adam. Computing in Nonlinear Media and Automata Collectives. Bristol: Inst. of Physics Publ., 2001. 2. Adamatzky, Adam, De Lacy Costello, Ben, & Asai, Tetsuya. Reaction Diffusion Computers. Amsterdam: Elsevier, 2005. 2013/2/1 35 6