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Systems of Difference Equations-Host-parasitoid Models: Ap CN P Ap BN N
Systems of Difference Equations-Host-parasitoid Models: Ap CN P Ap BN N
N t 1 bN t exp( aPt )
Pt 1 cN t 1 exp( aPt )
for t=0,1,2,. Here, N and P denote the population sizes of susceptible hosts and
searching adult female parasitoids at time t and t+1, respectively. The parameter b is
interpreted as the net growth parameter. We see from the first equation that hosts grow
exponentially in the absence of parasitoids (P=0). The term exp(aPt ) is the fraction of
hosts that are not parasitized (and thus 1 exp(aPt ) is the fraction of hosts that are
parasitized) at generation t. Parasitized hosts produce parasitoids. The parameter c is
equal to the average number of female parasitoids produced per parasitized host. All
parameters are nonnegative.
The Nicholson-Bailey model assumes that hosts and parasitoids encounter each
other randomly. This means that the total number of encounters is proportional to the
product of host and parasitoid densities. That is,
N e #encounters per unit time between hosts and parasitoids aHP
Therefore, the average number of encounters per host is given by
Ne
aP
H
-1-
m
m!
It holds that E ( X ) var( X ) . The histograms for different values of the parameter
are shown below:
Poisson Distribution = 1
0.4
0.2
0
0.2
0
10
15
10
15
20
Poisson Distribution = 4
0.2
0.2
0.1
0
10
15
20
Poisson Distribution = 5
0.2
10
15
20
Poisson Distribution = 6
0.2
0.1
0
20
Poisson Distribution = 3
0.4
Poisson Distribution = 2
0.4
0.1
0
10
15
20
10
15
20
With aP , we find
-2-
(2)
x1 (t 1) a11 x1 (t ) a12 x 2 (t )
x 2 (t 1) a 21 x1 (t ) a 22 x 2 (t )
x (t 1)
x (t )
It shows that systems of linear difference equations are linear maps. (Recall that the
Leslie matrix is an example where linear systems are used.)
A point equilibrium
x* Ax *
It follows that the zero vector
0
0
0
0
, we proceed as
follows. Recall that if the eigenvalues of A are distinct, the corresponding eigenvectors
are distinct, and we can write any vector uniquely as a linear combination of the two
corresponding eigenvectors. Assume now that the eigenvalues are real and distinct. We
initial vector x (0) can be written as a linear combination of the two eigenvectors
x (0) c1u1 c 2 u 2
-3-
where c1 and c2 are real numbers. Using this representation of x (0) , we found
previously that
x (t ) c11t u1 c 2 t2 u 2 ,
t 0,1,2,...
x (t ) .
which immediately yields insights into the long-term behavior of x (t ) , tlim
Namely, if both 1
and 2
1,
0
in this case that is a stable equilibrium. If either 1 1 or 2 1 , then the
0
0
equilibrium is unstable. The stability condition holds more generally, not just for
0
x (t 1) Ax (t )
and 2
1.
If either 1
or 2
Task 1:
of the system
x1 (t 1)
0.4 0.2 x1 (t )
x (t 1) 0.3 0.1 x (t )
2
2
Are there any other equilibria? Find a geometric argument that explains your answer.
If one of the eigenvalues is equal to 1, then there are infinitely many equilibria, as
illustrated in the following example.
Example: Suppose
-4-
3 2.5
A
The eigenvalues of A are given by
1 1 and 2 0.5
r
r
To find equilibria of x (t 1) Ax (t ) , we solve
2 1.5 x1
3 2.5 x
x1
x
2
3 x1 2.5 x2 x2
3 x1 1.5 x2 0
3 x1 1.5 x2 0
The second set of equations consists of two identical equations, namely, after simplifying,
x1 0.5 x2 0 . This means that any point ( x1 , x2 ) that satisfies x1 0.5 x2 0 , is a point
equilibrium. These equilibria are neutrally stable. This means that they are neither
repellent nor attracting. After a small perturbation, the system will reach a different
equilibrium point on the line given by x1 0.5 x2 0 .
Equilibria and Stability of Nonlinear Systems of Difference Equations
The general form of a system of two nonlinear difference equations is
(3)
x1 (t 1) F x1 (t ), x 2 (t )
x 2 (t 1) G x1 (t ), x 2 (t )
where F and G are (nonlinear) functions. We say that x1* , x 2* is a point equilibrium if the
following two equalities hold simultaneously:
-5-
G x
x1* F x1* , x 2*
x 2*
* *
1 , x2
Task 2:
Find all equilibria of
x1 (t 1) 2 x1 (t )1 x1 (t )
x 2 (t 1) x1 (t )1 x 2 (t )
Task 3:
Find all biologically relevant equilibria of the Nicholson-Bailey model
N t 1 bN t exp( aPt )
Pt 1 cN t 1 exp( aPt )
x1 (t 1) F x1 (t ), x 2 (t )
x 2 (t 1) G x1 (t ), x 2 (t )
x1* F x1* , x 2*
and
x 2* G x1* , x 2*
and
x 2 (t ) x 2* z 2 (t )
linearization of F x1 (t ), x 2 (t ) is F x1* , x 2*
x1
-6-
F
x 2
z1 (t )
z 2 (t )
x1* , x 2*
x1
G
z1 (t )
x 2
z 2 (t )
where * means that we evaluate the expression in the parentheses at the equilibrium
x1* , x 2* .
With x1 (t ) x1* z1 (t ) and x 2 (t ) x2* z 2 (t ) , we find that approximately
x1* z1 (t 1) F x1* , x 2*
x1
F
x 2
z1 (t )
x1
x 2* z 2 (t 1) G x1* , x 2*
x1
G
x 2
z1 (t )
x2
z 2 (t )
z 2 (t )
Canceling x1* from the first equation and x 2* from the second equation, and writing the
resulting approximation in matrix form, we obtain
(4)
x1
z1 (t 1)
z (t 1)
2
G
x1
*
F
x 2 z1 (t )
*
G z 2 (t )
x 2
stability. The right-hand side of Equation (4) is a linear map and we can thus use the
results from linear systems of difference equations to draw conclusions about the stability
of the equilibrium. We summarize the results in the following box.
The point equilibrium x1* , x 2* of the nonlinear system (3) is locally stable if the two
eigenvalues 1 and 2 of the Jacobian matrix evaluated at x1* , x 2* ,
-7-
x1
G
x1
*
F
x 2
*
x 2
satisfy
1 1
If 1
or 2
1,
2 1
and
Task 4:
Discuss the stability of the equilibria of the system in Task 2.
To calculate the Jacobian for the Nicholson Bailey model, we can use Matlab:
>>
>>
>>
>>
syms a b c x y
f = [b*x*exp(-a*y);c*x*(1-exp(-a*y))];
v=[x,y];
R = jacobian(f,v);
be ay
bxae ay
R
ay
ay
c(1 e ) cxae
The following Matlab code calculates eigenvalues for different values of the parameter b:
%Nicholson-Bailey Stability Analysis
syms x y
a=0.023;c=2;k=1;
for b=0.5:0.1:1.5
-8-
end
abslambda=abs(lambda);
-9-
A
B
C
Nicholson-Bailey Model
a
0.023
Time
1.5
N_t
N_0
P_0
25
10
P_t
0
25
10
1 29.79501 10.27332
2 35.28718 12.54045
3 39.66838 17.68318
3 2.5
Open the EXCEL spreadsheet (Tab: Step 3) to investigate the dynamics of the system
r
r
of difference equations given by x (t 1) Ax (t ) . Recall from the worksheet that the
eigenvalues of A are 1 and -0.5 and that all point equilibria are on the line given by
x1 0.5 x2 0 . These point equilibria are neutrally stable.
(a) Run the dynamics for the first 40 time steps for the following three initial conditions:
(i) ( x1 (0), x2 (0)) (0.1, 0.2) , (ii) ( x1 (0), x2 (0)) (3, 1) , and (iii) ( x1 (0), x2 (0)) (5, 2) . In
your write-up, include the values at time 1 and time 40. (You do not have to include
graphs.)
(b) Calculate the ratios x2 (t ) / x1 (t ) for the three runs in (a). What happens for t large?
List the ratios at time 40 in your homework and explain your result in light of the
statement that all point equilibria are on the line given by x1 0.5 x2 0 . (You do not
have to include graphs.)
Step 4
(a) Over the past several weeks, you have gained significant experience with
mathematical modeling in the context of population models. Population models
have the advantage of allowing us to introduce the mathematics without having to
deal with complex biology. We will soon leave the topic of population models and
discuss other biological systems, continuing to broaden our toolbox and learning
about other systems. Write a short essay on which areas of biology you have
encountered (or hope to encounter) mathematical models in.
(b) Reflecting over the past several weeks and looking back at mathematics courses you
have taken in the past, write a short essay on what works best for you when you
learn mathematics. Address both in-class learning styles and how you learn at
home.
- 11 -
N t 1 bN t 1 t
k
aP
Pt 1 cN t 1 1 t
k
The parameter k is positive and measures aggregation of parasitoid attacks. The model is
locally stable when k<1. For very small values of k, the approach is monotonic, whereas
larger values of k result in oscillations.
Use MATLAB to code up the negative binomial model for the parameters a=0.023,
b=1.5, and c=2 with initial values N0=25 and P0=10. Investigate the temporal dynamics
for t=0,1,2,,100 for k between 0.1 and 1.3 by varying k in increments of 0.4. What do
you observe? Include graphs for each value of k. Include your MATLAB code in your
homework.
Figure 2: The parasitoid Cyzenis albicans (tachnid fly) parasitizes the winter moth, Operophtera brumata,
by laying eggs on the leaves of the winter moths food plant. The black bars are the observed values, the
hatched are fitted from a negative binomial distribution with parameter k=0.28, and the hollow bars are
from a Poisson distribution. The figure is from Hassell (2000), page 181.
Hassell, M.P. 2000. The Spatial and Temporal Dynamics of Host-Parasitoid Interactions. Oxford
University Press.
- 12 -