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Model 2

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Two Species Models

The models we have discussed so far (Malthus and Logistic) are single

species models. Many of the most interesting dynamics in the biological world

have to do with interactions between species. Mathematical models which

incorporate these interactions are required if we hope to simulate these dynamics.

One of the first models to incorporate interactions between predators and prey was

proposed in 1925 by the American biophysicist Alfred Lotka and the Italian

mathematician Vito Volterra. Unlike the Malthusian and Logistic models we have

previously seen, the Lotka-Volterra model is based on differential equations.

Differential Equations

For our purposes, the best way to understand differential equations is to contrast

them with the now familiar difference equations. When using difference equations

the population sizes are computed at discrete points in time. These results can be

likened to snapshots, we observe the situation at points in time but do not observe

what happens between snapshots. In the difference equation models we've

discussed so far, the snapshots have been taken at the beginning of each time

period, often years.

at every moment in time. They accomplish this by trying to find a formula X(t) for

the population where t can be any value, not necessary an integer. In our analogy,

the solution to a differential equation is like watching an event unfold, observing

every nuance of its evolution.

In practice, this lofty goal of differential equations is not fully met. This is because

of the difficulty of finding the formulas that actually solve differential equations.

What normally happens instead is that a numerical approximation of the

differential equation's solution is found. This is routinely done by converting the

differential equation into an "equivalent" difference equation, and solving that.

(For details, see any undergraduate differential equations text. Most modern

calculus text also discuss numerical solution of differential equations.) The

"equivalent" difference equation is usually solved using very small time steps. The

result is like a motion picture; the illusion of smooth motion is obtained while in

fact we are seeing a sequence of snapshots.

Instead of giving us information on how much something changes over the course

of one full time period, differential equations give us a formula for

the derivative (from calculus) of some interesting quantity like a population level.

Fortunately for us, the derivative is just a fancy notion for the change in a quantity

per unit time. So, as far as we're concerned, we can treat the formula for the

derivative in exactly the same manner as we treated the formula for change in the

difference equation case. The only change is that in the differential equation case it

is a good idea to use small time steps rather than the time step of 1 typically used

in difference equation models.

As those of you who have calculus in your backgrounds may remember, the

derivative has many different notations. If X(t) is a function, the derivative of

X(t) can be represented in a fraction-like form dX/dt or as X'(t). Another common

notation is to put a single dot over the function name. We will use the fraction-like

form.

symbol to the left of an equal sign and the formula for the derivative to the right.

For example, a differential equation form of the Logistic model would take the

form

dX/dt = r*X*(1-X/K).

To solve this model with Stella we note that both sides of this equation represent

the change in population size, just as X(i+1) - X(i) does in the difference equation

models. Consequently, the right hand side of the differential equation is precisely

the formula that should appear in the flow in the Stella model. Note that this gives

us exactly the same model as we derived in the Logistic difference equation case.

We can use either the bi-directional single flow version or the better birth/death

two flow version.

Just as with difference equation models, differential equation models require initial

conditions. These are indicated in precisely the same way as for the difference

equation models discussed earlier.

This works for any differential equation model, simply use the right hand side of

the differential equation as the formula for the flow setting. Let me emphasize

again that the only difference is that differential equation models should generally

be solved with small time steps, while difference equation models should be solved

with time steps of 1.

Exercise

1. Run the Logistic Stella model developed earlier under several different time

steps. Does the overall behavior of the model seem sensitive to the time

step?

differential equation case as it is in the difference equation case, and just as easy.

To find the equilibria simply set the derivative equal to zero (indicating no change)

and solve the resulting algebraic (no derivatives) equation for the equilibrium. In

the case of the logistic example given above, the equilibria are exactly the same for

the difference equation models and the differential equation models.

Lotka-Volterra

The Lotka-Volterra model describes interactions between two species in an

ecosystem, a predator and a prey. This represents our first multi-species model.

Since we are considering two species, the model will involve two equations, one

which describes how the prey population changes and the second which describes

how the predator population changes.

For concreteness let us assume that the prey in our model are rabbits, and that the

predators are foxes. If we let R(t) and F(t) represent the number of rabbits and

foxes, respectively, that are alive at time t, then the Lotka-Volterra model is:

dF/dt = e*b*R*F - c*F

The Stella model representing the Lotka-Volterra model will be slightly more

complex than the single species models we've dealt with before. The main

difference is that our model will have two stocks (reservoirs), one for each species.

Each species will have its own birth and death rates. In addition, the Lotka-Volterra

model involves four parameters rather than two. All told, the Stella representation

of the Lotka-Volterra model will use two stocks, four flows, four converters and

many connectors.

Exercises

1. Split the rabbit's differential equation into the births part and the deaths part.

3. Using the following parameter values, write down the differential equations

for the Lotka-Volterra model and find all equilibrium points. This will

involve solving two equations for two unknowns

(namely R(*)and F(*)). HINT: this model produces two steady states, one of

which should be unsurprising.

o a = 0.04

o b = 0.0005

o c = 0.2

o e = 0.1

terms of the parameters. In other words, try to find formulas

for R(*) and F(*) without plugging in specific values for the parameters.

5. Create a Stella model for the Lotka-Volterra model. Use the parameter

values given above as values for the four converters in your model. Try

various initial conditions for the rabbits and fox populations; choose some to

be near the equilibria you determined above, and have some be far away.

Use different time steps and different running times. Which equilibrium is

stable, unstable?

6. Try both the usual time series graph and the scatter graph to examine the

model output. A scatter plot of rabbit versus fox population is particularly

interesting. To produce such a graph pull down the graph icon, place it

somewhere in the model, double click on the graph when it appears and

select the scatter plot option. This will require you to choose two quantities

to plot. Pick the rabbit and fox populations. You should get some interesting

pictures if you let the model run long enough.

Analysis of Lotka-Volterra

The Lotka-Volterra model is one of the earliest predator-prey models to be based

on sound mathematical principles. It forms the basis of many models used today in

the analysis of population dynamics. Unfortunately, in its original form Lotka-

Volterra has some significant problems. As you may have noted in your

experiments, neither equilibrium point is stable. Instead the predator and prey

populations seem to cycle endlessly without settling down quickly. It can be shown

(see any undergraduate differential equations book for details) that this behavior

will be observed for any set of values of the model's four parameters. While this

cycling has been observed in nature, it is not overwhelmingly common. It appears

that Lotka-Volterra by itself is not sufficient to model many predator-prey systems.

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