Part 2: Cellular Automata 9/13/04 StarLogo Simulation of Streaming Aggregation 1. 2. 3. 4. chemical diffuses if cell is refractory (yellow) then chemical degrades else (it’s excitable, colored white) 1. if chemical > movement threshold then 2. else if chemical > relay threshold then Demonstration of StarLogo Simulation of Streaming take step up chemical gradient Run SlimeStream.slogo produce more chemical (red) become refractory 3. else wait 9/13/04 1 9/13/04 Differentiation & Pattern Formation 2 Zebra • 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 9/13/04 photos ©2000, S. Cazamine 3 9/13/04 figs. from Cazamine & al.: Self-Org. Biol. Sys. 4 1 Part 2: Cellular Automata 9/13/04 Vermiculated Rabbit Fish Activation & Inhibition in Pattern Formation • 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 9/13/04 figs. from Cazamine & al.: Self-Org. Biol. Sys. 5 9/13/04 6 Interaction Parameters Emergent Hierarchical Structure • Characteristic length scale • Independent of cell size and space size • Structures created at intermediate level • R1 and R2 are the interaction ranges • J1 and J2 are the interaction strengths 9/13/04 7 9/13/04 8 2 Part 2: Cellular Automata 9/13/04 Example CA Activation/Inhibition Model • • • • (R1=1, R2=6, J1=1, J2=–0.1, h=0) 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: si ( t + 1) = signh + J1 s j ( t ) + J 2 s j ( t ) rij <R1 R1 <rij <R 2 9/13/04 9 figs. from Bar-Yam 9/13/04 Effect of Bias Effect of Interaction Ranges (h = –6, –3, –1; 1, 3, 6) R2 = 8 R1 = 1 h=0 R2 = 6 R1 = 1 h=0 R2 = 6 R1 = 1.5 h=0 9/13/04 figs. from Bar-Yam 10 11 9/13/04 R2 = 6 R1 = 1.5 h = –3 figs. from Bar-Yam 12 3 Part 2: Cellular Automata 9/13/04 Digression on Diffusion Differential Interaction Ranges • Simple 2-D diffusion equation: • 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 13 9/13/04 reaction 14 0 2 A f A ( A,I ) + dI 2 I f I ( A,I ) • Let be the logistic sigmoid function • Activator A and inhibitor I may diffuse at different rates in x and y directions • Cell is “on” if activator + bias exceeds inhibitor A 2A 2A = dAx 2 + dAy 2 + k A [ mA ( A + B I )] t x y A c˙ = D 2c + f (c), where c = I 9/13/04 2 A( x, y ) 2 A( x, y ) + x 2 y 2 Example: Activation-Inhibition System Reaction-Diffusion System A d A = t I 0 2 A( x, y ) = – positive in a local minimum – negative in a local maximum 9/13/04 diffusion • Recall the 2-D Laplacian: • The Laplacian (like 2nd derivative) is: – activator diffuses slowly (short range) – inhibitor diffuses rapidly (long range) A = dA 2 A + f A ( A,I ) t I = dI 2 I + f I ( A,I ) t A˙ ( x, y ) = c 2 A( x, y ) I 2I 2I = dIx 2 + dIy 2 + k I [ mI ( A + B I)] t x y 15 9/13/04 16 4 Part 2: Cellular Automata 9/13/04 Demonstration of StarLogo Program for Activation/Inhibition Pattern Formation Run Pattern.slogo 9/13/04 17 Abstract Activation/Inhibition Spaces • Consider two axes of cultural preference – E.g. hair length & interpersonal distance – Fictitious example! • Suppose there are no objective reasons for preferences • Suppose people approve/encourage those with similar preferences • Suppose people disapprove/discourage those with different preferences • What is the result? 9/13/04 18 A Key Element of Self-Organization Emergent Regions of Acceptable Variation • Activation vs. Inhibition • Cooperation vs. Competition • Amplification vs. Stabilization • Growth vs. Limit • Positive Feedback vs. Negative Feedback – Positive feedback creates – Negative feedback shapes 9/13/04 19 9/13/04 20 5 Part 2: Cellular Automata 9/13/04 Additional Bibliography 1. 2. 3. 4. 5. 6. 9/13/04 Kessin, R. H. Dictyostelium: Evolution, Cell Biology, and the Development of Multicellularity. Cambridge, 2001. Gerhardt, M., Schuster, H., & Tyson, J. J. “A Cellular Automaton Model of Excitable Media Including Curvature and Dispersion,” Science 247 (1990): 1563-6. Tyson, J. J., & Keener, J. P. “Singular Perturbation Theory of Traveling Waves in Excitable Media (A Review),” Physica D 32 (1988): 327-61. Camazine, S., Deneubourg, J.-L., Franks, N. R., Sneyd, J., Theraulaz, G.,& Bonabeau, E. Self-Organization in Biological Systems. Princeton, 2001. Pálsson, E., & Cox, E. C. “Origin and Evolution of Circular Waves and Spiral in Dictyostelium discoideum Territories,” Proc. Natl. Acad. Sci. USA: 93 (1996): 1151-5. Solé, R., & Goodwin, B. Signs of Life: How Complexity Pervades Biology. Basic Books, 2000. continue to “Autonomous Agents” 21 6