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Introduction to the Modeling and Analysis of Complex Systems

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13.4. MODELING SPATIAL MOVEMENT 247• Cells nei<strong>the</strong>r die nor divide.Did you notice <strong>the</strong> similarity between <strong>the</strong>se assumptions <strong>and</strong> those we used for <strong>the</strong>population-economy model? Indeed, <strong>the</strong>y are identical, if you replace “cells” with “people”<strong>and</strong> “cAMP” with “money.” We could use <strong>the</strong> word “moneytaxis” for people’s migration<strong>to</strong>ward economically active areas!The actual Keller-Segel equations look like this:∂a∂t = µ∇2 a − χ∇ · (a∇c) (13.27)∂c∂t = D∇2 c + fa − kc (13.28)In fact, <strong>the</strong>se equations are simplified ones given in [49], as <strong>the</strong> original equations werera<strong>the</strong>r complicated with more biological details. a <strong>and</strong> c are <strong>the</strong> state variables for celldensity <strong>and</strong> cAMP concentration, respectively. µ is <strong>the</strong> parameter for cell mobility, χ is<strong>the</strong> parameter for cellular chemotaxis, D is <strong>the</strong> diffusion constant <strong>of</strong> cAMP, f is <strong>the</strong> rate <strong>of</strong>cAMP secretion by <strong>the</strong> cells, <strong>and</strong> k is <strong>the</strong> rate <strong>of</strong> cAMP decay. Compare <strong>the</strong>se equationswith Eqs. (13.25) <strong>and</strong> (13.26). They are <strong>the</strong> same! It is intriguing <strong>to</strong> see that completelydifferent phenomena at vastly distant spatio-temporal scales could be modeled in an identicalma<strong>the</strong>matical formulation.It is known that, for certain parameter settings (which will be discussed in <strong>the</strong> nextchapter), this model shows spontaneous pattern formation from almost homogeneousinitial conditions. A sample simulation result is shown in Fig. 13.12, where we see spots <strong>of</strong>aggregated cell clusters forming spontaneously. You will learn how <strong>to</strong> conduct simulations<strong>of</strong> continuous field models in <strong>the</strong> next section.In Fig. 13.12, we can also observe that <strong>the</strong>re is a characteristic distance betweennearby spots, which is determined by model parameters (especially diffusion constants).The same observation applies <strong>to</strong> <strong>the</strong> formation <strong>of</strong> cities <strong>and</strong> <strong>to</strong>wns at geographical scales.When you see a map, you will probably notice that <strong>the</strong>re is a typical distance betweenmajor cities, which was probably determined by <strong>the</strong> human mobility centuries ago, amongo<strong>the</strong>r fac<strong>to</strong>rs.Exercise 13.13 Consider introducing <strong>to</strong> <strong>the</strong> Keller-Segel model a new variable bthat represents <strong>the</strong> concentration <strong>of</strong> a <strong>to</strong>xic waste chemical. Make <strong>the</strong> followingassumptions:• cAMP gradually turns in<strong>to</strong> <strong>the</strong> waste chemical (in addition <strong>to</strong> natural decay).• The waste chemical diffuses over space.

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