Observations
Excitable media can support circular and spiral waves
Spiral formation can be triggered in a variety of ways
All seem to involve inhomogeneities (broken symmetries):
in space
in time
in activity
Amplification of random fluctuations
Circles & spirals are to be expected

StarLogo Simulation of Streaming Aggregation
 chemical diffuses
 if cell is refractory (yellow)
 then chemical degrades
 else (itÕs excitable, colored white)
 if chemical > movement threshold then
take step up chemical gradient
 else if chemical > relay threshold then
produce more chemical (red)
become refractory
 else wait

Demonstration of StarLogo Simulation of Streaming
Run SlimeStream.slogo

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

Zebra
Vermiculated Rabbit Fish
Activation & Inhibition
in Pattern Formation
Color patterns typically have a charac-teristic length scale
Independent of cell size and animal size
Achieved by:
short-range activation Þ local uniformity
long-range inhibition Þ separation

Emergent Hierarchical Structure
Characteristic length scale
Independent of cell size and space size
Structures created at intermediate level

Interaction Parameters
R1 and R2 are the interaction ranges
J1 and J2 are the interaction strengths

CA Activation/Inhibition Model
Let states si ë {Ð1, +1}
and h be a bias parameter
and rij be the distance between cells i and j
Then the state update rule is:

Example
(R1=1, R2=6, J1=1, J2=Ð0.1, h=0)
Effect of Bias
(h = Ð6, Ð3, Ð1; 1, 3, 6)
Effect of Interaction Ranges
Demonstration of StarLogo Program for Activation/Inhibition Pattern Formation
Run Pattern.slogo

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?

Emergent Regions of Acceptable Variation
A Key Element of
Self-Organization
Activation vs. Inhibition
Cooperation vs. Competition
Amplification vs. Stabilization
Growth vs. Limit
Positive Feedback vs. Negative Feedback
Positive feedback creates
Negative feedback shapes

Additional Bibliography
Kessin, R. H.  Dictyostelium: Evolution, Cell Biology, and the Development of Multicellularity.  Cambridge, 2001.
Gerhardt, M., Schuster, H., & Tyson, J. J.  ÒA Cellular Automaton Model of Excitable Media Including Curvature and Dispersion,Ó Science 247 (1990): 1563-6.
Tyson, J. J., & Keener, J. P.  ÒSingular Perturbation Theory of Traveling Waves in Excitable Media (A Review),Ó 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.
P‡lsson, E., & Cox, E. C.  ÒOrigin and Evolution of Circular Waves and Spiral in Dictyostelium discoideum Territories,Ó Proc. Natl. Acad. Sci. USA: 93 (1996): 1151-5.
SolŽ, R., & Goodwin, B.  Signs of Life: How Complexity Pervades Biology.  Basic Books, 2000.