A Course in Math Bio Sol

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Solutions Manual
to accompany
A Course in Mathematical Biology
Quantitative Modeling with Mathematical and Computational Methods
G. de Vries, T. Hillen, M. Lewis, J. M ulller, and B. Sch onsch,
Society for Industrial and Applied Mathematics, Philadephia, 2006
A. Beltaos G. de Vries T. Hillen
November 20, 2006 sol-gerda
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2 Solutions manual for de Vries et al, SIAM 2006 sol-gerda
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Contents
1.4 Exercises for Modeling . . . . . . . . . . . . . . . . . . . . . . . 6
2.4 Exercises for Discrete-Time Models . . . . . . . . . . . . . . . . 7
3.9 Exercises for ODEs . . . . . . . . . . . . . . . . . . . . . . . . . 29
4.5 Exercises for PDEs . . . . . . . . . . . . . . . . . . . . . . . . . 55
5.8 Exercises for Stochastic Models . . . . . . . . . . . . . . . . . . 61
6.6 Exercises for Cellular Automata . . . . . . . . . . . . . . . . . . 62
7.7 Exercises for Parameter Estimation . . . . . . . . . . . . . . . . 63
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4 Solutions manual for de Vries et al, SIAM 2006 sol-gerda
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Note to Readers
This document contains solutions to a selection of problems, primarily problems
from Chapters 2, 3, and 4.
We welcome submissions to ll in the missing solutions (preferably in latex format)
and gures (preferably in postscript or encapsulated postscript format). We will be
pleased to credit your authorship in the corresponding solution in updated versions
of this document.
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6 Solutions manual for de Vries et al, SIAM 2006
1.4 Exercises for Modeling
Exercise 1.4.1: Discrete-time versus continuous-time models
(a) t = 10 minutes, and the probability of one cell making one other cell in 10
minutes is p = 1, since we are told that each cell eectively makes one copy
of itself every 10 minutes.
(b) Letting N(t) be the population at time t, we have:
N(t + t) = N(t) + pN(t) (1.1)
= N(t) + N(t)
= 2N(t)
We can see that the whole population doubles every 10 minutes.
(c) We can dene :=
p
t
(in this case, =
1
10
min
1
), and we can change our
discrete model. Dividing (1.1) by t, we get:
N(t + t)
t
=
N(t)
t
+
p
t
N(t) =
N(t)
t
+ N(t)
N(t + t) N(t)
t
= N(t)
If we take the limit as t 0, we obtain the continuous model:
d
dt
N(t) = N(t)
Exercise 1.4.2: Comparison of discrete and continuous models
Solution not available.
Exercise 1.4.3: Structured populations
Solution not available. sol-gerda
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A. Beltaos, G. de Vries, T. Hillen, November 20, 2006 7
2.4 Exercises for Discrete-Time Models
Exercise 2.4.1: German population
Let x
n
= Germanys population at the end of year n. The simplest model incorpo-
rating the given information is
x
n+1
= x
n
x
n
+ x
n
= (1 + )x
n
(2.2)
= rx
n
,
where r = 1 + .
At the end of the year 1998, we have x
1998
= 82, 037, 000, and
x
1998
= 770, 744, and x
1998
= 846, 330,
which gives the values of and to be:
_
_
_
=
770,744
82,037,000
= 0.009395,
=
846,330
82,037,000
= 0.010316,
(2.3)
and hence r = 1 0.010316 + 0.009395 = 0.999079 < 1. That is, the population
will decay.
To make the model more realistic, we could add in terms for immigration and
emigration, and use a logistic equation.
Exercise 2.4.2: Drug prescriptions
(a) The dimensionless parameter k represents the fraction of the drugs dose which
is used up or broken down by the body. The amount of the drug in a dose is
represented by b. We know that b 0 (no negative doses), and k > 0 (the body
must use up some of the drug). Notice that if b = 0, then a
n+1
= a
n
ka
n
, so
we better have k 1 to avoid negative amounts. Summarizing the conditions
on k and b,
k (0, 1],
and
b 0.
(b) The xed point (we know there is only one, since it is a linear equation), a, of
this model can be found if we let f(x) := x kx + b, and solve for f( a) = a.
This gives us a = b/k. Further, f
(x) = 1 k. We found in part (a) that

k (0, 1], so that means that f
(x) [0, 1), which means that this xed point,

a = b/k, is stable. sol-gerda
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(c) In every case, we will start with the initial value of 0 (representing no dose
given yet). Looking at the cobweb diagrams below, we can see that the map
always settles down to the xed point. The drug will have a constant concen-
tration after sucient time has passed, unless the dose is changed. The xed
point is reached more quickly if k is larger. In the gure below on the left,
we have k = 0.1, and we can see many steps leading up to the xed point.
In the gure on the right, we set k = 0.6, and it takes fewer iterations of the
map to reach the xed point. Note that in these gures, we have set b = 1.
Also, a larger value of k gives a smaller actual value of the xed point. Since
the xed point is a = b/k, we see that b behaves like a scaling factor, simply
changing the value, but not the quality of the xed point.
0
2
4
6
8
10
12
y
2 4 6 8 10 12
x
0
0.5
1
1.5
2
y
0.5 1 1.5 2
x
(d) We assume that there is a minimum amount, E, for which the drug will be
eective, and a maximum amount, T, for which the drug becomes toxic. To
make sure the drug is eective but not toxic, we need the xed point to be
between the values E and T:
E a < T
Substituting in for a (found in part (b)), and multiplying through by k, we
need the dosage b to be between the values kE and kT:
kE b < kT.
Exercise 2.4.3: Improving the t of the logistic model to the data
Exercise 2.4.4: Fluctuations in the population of P. aurelia
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Exercise 2.4.5: Whale population
(a) The given equation,
a
n+1
= a
n
+ k(M a
n
)(a
n
m),
describes the population of whales next year in terms of the population this
year. The last term describes the change in the population, which is dependent
on the 3 parameters, k, M, and m, with m < M.
If 0 < a
n
< m, then k(Ma
n
)(a
n
m) < 0, so the population will decline
(specically, the population next year is smaller than the population this
year).
If m < a
n
< M, then k(M a
n
)(a
n
m) > 0, so the population will
grow.
If a
n
> M, then k(M a
n
)(a
n
m) < 0, so the population will decline.
If a
n
= m or a
n
= M, then k(Ma
n
)(a
n
m) = 0, so the population will
remain the same size (that is, m and M are xed points of the model).
(b) For xed points a, we require
a = a + 0.0001(5000 a)( a 100)
0.0001(5000 a)( a 100) = 0
a = M = 5000 or a = m = 100.
To determine their stability, we let
f(x) = x + 0.0001 (5000 x)(x 100)
= x + 0.0001 (5000x 500000 x
2
+ 100x)
= x + 0.0001 (5100x 500000 x
2
).
Then
f
(x) = 1 + 0.0001(5100 2x).

For the xed point a = m = 100, we have
f
(100) = 1 + 0.0001(5100 200)

= 1.49 > 1,
therefore a = m = 100 is unstable. For the xed point a = M = 5000 we have
f
(5000) = 1 + 0.0001(5100 10000)

= 0.51 < 1 (but greater than 1),
therefore a = M = 5000 is stable. sol-gerda
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(c)
100
a
0
5000 a
n
a
n+1
a
n+1
= a
n
As before, we conclude that a = m = 100 is unstable, and a = M = 5000 is
stable.
(d)
n
m = 100
M = 5000
a
n
n
m = 100
M = 5000
a
n
n
m = 100
M = 5000
a
n
n
m = 100
a
n
(e) When a
0
< m, the population declines, eventually becoming negative. When
a
0
M (i.e., when a
0
is greater than the biggest root of the map), then sol-gerda
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a
1
< 0, and the population continues to decline. Both cases are problematic,
since it is non-sensical to have a negative number of whales.
(f) If we plot a
n+1
versus a
n
for the given model, we obtain the following quali-
tative graph:
m
M a
n
a
n+1
stable unstable
a
n+1
= a
n
We see that if the whale population ever reaches a value located on the high-
lighted (thick) portions of the a
n
-axis, the population will be negative next
year, since the graph of a
n+1
versus a
n
lies below the a
n
-axis there.
To x the problem, we could ensure that the graph of a
n+1
versus a
n
always
lies above the a
n
-axis. For example:
m
M a
n
a
n+1
a
n+1
= a
n sol-gerda
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Exercise 2.4.6: Second-iterate map
(a)
f
2
(x) = r (rx(1 x)) (1 (rx(1 x)))
= r
2
x(1 (r + 1)x + 2rx
2
rx
3
)
(b) We already know the xed points of f. Since they are xed points of f, they
also are xed points of f
2
. These two trivial 2-cycles are x = 0 and x =
r1
r
.
Since we know they are xed points of f
2
, i.e., we know they satisfy f
2
(x) = x,
we know that they are solutions of f
2
(x) x = 0. This means we can rewrite
f
2
(x) x as x(x
r1
r
)q(x), where q(x) is an unknown quadratic polynomial.
We then set this new expression equal to what we know for f
2
(x) x:
x(x
r 1
r
)q(x) = r
2
x(1 (r + 1)x + 2rx
2
rx
3
) x.
Since we are interested in nonzero solutions, we can drop the common x factor
on each side. We dont want the (r 1)/r root either, so we can drop that,
by diving both sides by x
r1
r
. Hence,
q(x) =
r
2
(1 (r + 1)x + 2rx
2
rx
3
) 1
x
r1
r
.
Carrying out the long division, we end up with
q(x) = r
3
x
2
+ (r
3
+ r
2
)x r
2
r.
The roots of this polynomial are the last two roots of f
2
. Now we can nd
them by simply invoking the quadratic formula:
x =
(r
3
+ r
2
)
_
(r
3
+ r
2
)
2
4r
3
(r
2
r)
2r
3
.
This can be simplied to
x =
r + 1
_
(r 3)(r + 1)
2r
.
It is these two roots we are interested in, because the rst two are trivial
2-cycles. Recall the restriction of r [0, 4] which was discussed in section
(later), so the term (r + 1) in the radical poses no problem for us. It is the
(r 3) term which tells us when this 2-cycle exists. Obviously, this 2-cycle
does not exist if r < 3 (imaginary roots). If r = 3, this ceases to be a 2-cycle,
because the roots which make up the 2-cycle become the same (the radical
evaluates to zero). Hence, a nontrivial 2-cycle exists only for r > 3. sol-gerda
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(c) After a bit of simplication, we nd that
d
dx
f
2
(x) = 4r
3
x
3
+ 6r
3
x
2
2(r
2
+ r
3
)x + r
2
.
(d) This can be done with a few MAPLE commands. One is complicated. What
we want to do is turn the derivative, with the xed points substituted in
already, into a function of r:
> d:=q-> subs( r=q, abs( subs( x=sol[3], diff(f2(x), x)
) ) ):
The explanation of this command is as follows (from the inside out):
We substitute the 3rd solution that MAPLE gave for the xed points of f
2
(x)
into the derivative of f
2
(x), and then take the absolute value. We make it a
function by replacing the rs that appear with the dummy variable, q, which
we are using to dene the function in terms of. Now if we call d(r
0
), it will
give us the absolute value of the derivative of f
2
at the xed point with r = r
0
.
Next, we want to plot d(r), between 2.8 and 3.6 (to include the points 3 and
1 +
6 3.449). A horizontal line has been added to see where the graph
crosses d(r) = 1.
0
0.2
0.4
0.6
0.8
1
1.2
1.4
1.6
2.9 3 3.1 3.2 3.3 3.4 3.5 3.6
r
Looking at this, we can see that d is above 1 for r < 3 and below 1 for r > r
,
where r
3.5. This means that the 2-cycle is unstable and stable in those
regions, respectively. To nd the exact values at which the stability changes,
we use the solve command:
> solve(d(r)=1, r);
1 +
6, 1
6, 3, 1
Hence, the 2-cycle is stable for 3 < r < 1 +
6 and unstable for r > 1 +
6.
Note that 1 +
6 3.449, conrming our observation from the graph above. sol-gerda

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Exercise 2.4.7: Fourth-iterate map
Exercise 2.4.8: Exact solution for the Beverton-Holt model
We have
x
n+1
=
r
1 +
r1
K
x
n
x
n
Let u
n
=
1
xn
. Then
1
u
n+1
=
r
1 +
r1
K
1
un
1
u
n
u
n+1
=
1
r
_
1 +
r 1
K
1
u
n
_
u
n
=
1
r
u
n
+
r 1
rK
This equation is linear, of the form
u
n+1
= Au
n
+ B,
where A =
1
r
and B =
r1
rK
.
We can nd a solution to this equation as follows:
u
n+1
= Au
n
+ B
= A[Au
n1
+ B] + B
= A
2
u
n1
+ B(A + 1)
= A
2
[Au
n2
+ B] + B(A + 1)
= A
3
u
n2
+ B(A
2
+ A + 1)
.
.
.
= A
n+1
u
0
+ B(A
n
+ A
n1
+ + A+ 1)
= A
n+1
u
0
+ B
A
n+1
1
A1
Returning to the original variables, we have
1
x
n+1
=
1
r
n+1
1
x
0
+
r 1
rK
1
r
n+1
1
1
r
1
=
1
x
0
r
n+1
+
r
n+1
1
r
n+1
K
=
K + x
0
(r
n+1
1)
x
0
Kr
n+1
x
n+1
=
r
n+1
x
0
1 +
x0
K
(r
n+1
1 sol-gerda
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Exercise 2.4.9: Fitting the Beverton-Holt model to Gauses data
Exercise 2.4.10: The tent map
(a) We distinguish three cases: Case 1: < 1, Case 2: = 1, Case 3: > 1.
Graphs of f are:
1 1/2 0
x
/2
1/2
1/2 1 0
x
/2 = 1/2
1 1/2 0
x
/2
1/2
Case 1: < 1 Case 2: = 1 Case 3: > 1
(b) In case 1, there is one xed point, x = 0. In case 2, there are innitely many
xed points 0 x
1
2
. In case 3, there are two xed points, x
1
= 0 and
x
2
=

1+
.
To determine their stability, we look at the derivative of f(x) at each of the
xed points:
f
(x) =
_
for 0 x < 0.5
for 0.5 < x 1
In case 1, f
(0) = < 1, therefore x = 0 is stable. In case 2, f
( x) = = 1,
and the stability of these xed points is undetermined. In case 3, f
(0) =
> 1, therefore x
1
= 0 is unstable. Similarly, f
_

1+
_
= < 1, and so
x
2
=

1+
is unstable as well.
(c) Well dene two functions, f
1
(x) = x, and f
2
(x) = (1x), and loosely dene
the terms the rst region and the second region to mean {x : 0 x < 0.5},
and {x : 0.5 < x 1}, respectively. There are three possibilities for any 2-
cycle in this situation, namely that both iterates lie in the rst region, they
both lie in the second region, or one lies in the rst region and the second in
the second region (theres really one more with the opposite condition of the
last one, but this one can be found by simply applying f to the one found
in the last case, and those are the two iterates of the same 2-cycle). To nd
2-cycles, we set x = f(f( x)). sol-gerda
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Case (i): Both lie in the rst region. Then f(f(x)) = f
2
1
(x) = (x) =
2
x, and the xed point x must satisfy
x =
2
x,
which has only the trivial solution, x = 0, which we already know to be
1-cycle, so we are not interested in this case.
Case (ii): Both lie in the second region. Then f(f(x)) = f
2
2
(x) = (1
[(1 x)]) =
2
+
2
x. The xed point must satisfy
x =
2
+
2
x,
which has solution
x =
(1 )
(1 )(1 + )
=

1 +
.
Since x must lie in the second region, we require > 1. This is the same
solution as for the 1-cycle, so again, we are not interested in this case.
Case (iii): The rst lies in the rst region, and the second lies in the
second region. Here, applying f twice will give f(f(x)) = f
2
(f
1
(x)) =
(1 x) =
2
x. The xed point must satisfy
x =
2
x,
which has solution x = /(1 +
2
). Note that this point lies in the rst
region no matter what is. The proof lies in the fact that for any R,
we have ( 1)
2
0. With a little rearranging, the inequality turns
into /(
2
+ 1) 1/2, for all R. As described above, we can nd
the other orbit of this 2-cycle by applying f to x. Renaming x to p,
the 2-cycle will be p and q, where q = f(p) =
2
/(1 +
2
). Since we
found this 2-cycle by assuming that the rst iterate is in the rst region
(which it is) and the second is in the second region, we must restrict
such that q is in the second region. That is, we need to make sure
that 1/2 <
2
/(
2
+ 1) 1. The right-hand side of the inequality is
trivial, but the left-hand side forces us to put > 1 (keeping in mind
that > 0). Therefore, a nontrivial 2-cycle exists only for > 1, and it
is:
_
p =

2
+ 1
, q =

2
2
+ 1
_
. sol-gerda
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(d)
0 1/2 1
x
1
y = f(x)
y = x
As in (c), we think of the dierent cases of where the iterates of a 3-cycle could
reside. There seem to be eight possibilities. We denote each possibility with
a 3-tuple, where 1 means the iterate lies in the rst region, and 2 means
the iterate lies in the second region. The eight possibilities thus are
(1, 1, 1),
(1, 1, 2),
(1, 2, 1),
(1, 2, 2),
(2, 1, 1),
(2, 1, 2),
(2, 2, 1),
(2, 2, 2).
Since a 3-cycle is literally a cycle, any cyclic permutation of the above 3-tuples
will be the same 3-cycle (it doesnt matter which is the rst iterate of the
three, the important thing is the order of the three). This means that we have
some repetition in the above list, for example (2, 1, 1) is the same as (1, 1, 2),
which is the same as (1, 2, 1). Eliminating the repeated ones, we obtain the
following four essential cases:
Case (i) : (1, 1, 1),
Case (ii) : (1, 1, 2),
Case (iii) : (1, 2, 2),
Case (iv) : (2, 2, 2).
To nd a rst point of a 3-cycle, we need x = f(f(f( x))).
Case (i): All of the iterates of our 3-cycle lie in the rst region. If this is
the case, then the equation we get for the xed point x of the third-iterate
function is (remember that = 2 in this problem)
x = f
3
( x) = f
3
1
( x) =
3
x = 8 x.
As in part (c), we have the 1-cycle x = 0 again, so this is a trivial 3-cycle. sol-gerda
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Case (ii): The rst two iterates are in the rst region and the third is in
the second region. Now we would have for x:
x = f
3
( x) = f
2
(f
1
(f
1
( x))) = f
2
(f
1
(2 x)) = f
2
(4 x) = 2(1 4 x) = 2 8 x.
Hence the xed point of the third-iterate function is x = 2/9. We can
nd the rest of this 3-cycle by applying f to this point, and applying
f again. Let p = 2/9, and then q and r will be the other two iterates
of the 3-cycle. q = f(p) = f(2/9) = f
1
(2/9) = 2 2/9 = 4/9, and
then r = f(q) = f(4/9) = f
1
(4/9) = 2 4/9 = 8/9. We can check that
p = f(r): f(r) = f(8/9) = f
2
(8/9) = 2(1 8/9) = 2/9 = p, as expected.
Hence, our 3-cycle is
_
p =
2
9
, q =
4
9
, r =
8
9
_
.
We could stop here, since the question asks to nd an orbit of period 3 (i.e.,
a 3-cycle), but well continue to check the cases, in case we can nd another
one.
Case (iii): The rst iterate lies in the rst region, and the other two are
in the second region. Our equation for the xed point of f
3
(x) is then
x = f
3
( x) = f
2
(f
2
(f
1
( x))) = f
2
(f
2
(2 x)) = f
2
(2(1 2 x))
= f
2
(2 4 x) = 2(1 (2 4 x)) = 2 4 + 8 x
= 2 + 8 x.
This time we get x = 2/7. Doing the same thing as for case (ii), we put
p = 2/7, and then q = f(p) = f(2/7) = f
1
(2/7) = 2 2/7 = 4/7, and
r = f(q) = f(4/7) = f
2
(4/7) = 2(1 4/7) = 2 3/7 = 6/7. We check the
solution by nding f(r) = f(6/7) = f
2
(6/7) = 2(16/7) = 2 1/7 = 2/7,
and we see that this is correct. Hence, another 3-cycle is
_
p =
2
7
, q =
4
7
, r =
6
7
_
.
Case (iv): All iterates of the 3-cycle are in the second region. Then the
equation for a xed point of f
3
(x) is
x = f
3
( x) = f
3
2
( x) = f
2
(f
2
(2(1 x))) = f
2
(f
2
(2 2 x)) = f
2
(2(1 (2 2 x)))
= f
2
(2 4 + 4 x) = f
2
(2 + 4 x) = 2(1 (2 + 4 x)) = 2(3 4 x)
= 6 8 x.
This gives us the solution x = 2/3. Again, we set p = 2/3, and let
q = f(2/3) = f
2
(2/3) = 2(1 2/3) = 2/3. We need not go any further,
because we can see that this is a 1-cycle, a trivial 3-cycle. This is the
1-cycle we found in part (b), with = 2 (we can verify: /(1 + ) =
2/(1 + 2) = 2/3). sol-gerda
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Exercise 2.4.11: Blood cell population
(a) First we graph the function on Maple for the three cases of m: m < 1, m = 1,
and m > 1, for a xed :
0
2
4
6
8
10
12
14
16
18
20
22
24
26
28
30
32
34
20 40 60 80 100
x
0
2
4
6
8
20 40 60 80 100
x
0
1
2
3
4
5
20 40 60 80 100
x
For m < 1, p
2
(x) as x . For m = 1, p
2
(x) approaches b for
x . If m > 1, then p
2
(x) will go to zero as x . It is actually this
case where p
2
(x) takes the shape of p
1
(x). The graph of p
1
(x) is shown below,
with a = 1 and b = 1.
0
0.02
0.04
0.06
0.08
0.1
0.12
0.14
0.16
0.18
0.2
0.22
0.24
0.26
0.28
0.3
0.32
0.34
0.36
2 4 6 8 10
x
If we look at the case m 1, we see that the peak gets sharper. In the limit
of m the graph looks linear with a sudden drop to zero, as shown below. sol-gerda
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0
2
4
6
8
20 40 60 80 100
x
Changing the value of causes the horizontal scale to change, but does not
aect the shape of the curve. Large gives larger p
2
(x) at larger x. Finally,
changing the value of b simply changes the vertical scale.
(b) Assume it is q days that red blood cell production is delayed. Then instead
of the original model,
x
n+1
= x
n
d(x
n
) + p(x
n
),
we take the number of cells gained q days earlier, so the model becomes
x
n+1
= x
n
d(x
n
) + p(x
nq
).
Exercise 2.4.12: Population genetics
Exercise 2.4.13: Competition
Exercise 2.4.14: Spread of infectious disease
(a) For xed points

I, we require
I =

I + k
I(N

I)
k
I(N

I) = 0

I = 0 or

I = N.
To determine the stability of the xed points, we let
f(I) = (1 + kN)I kI
2
. sol-gerda
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Dierentiating gives
f
(I) = 1 + kN 2kI.
Since
f
(0) = 1 + kN > 1,
the trivial xed point

I = 0 is unstable. The stability of the nontrivial xed
point

I = N is given by
f
(N) = 1 + kN 2kN = 1 kN.

Since kN < 2, we have 1 < 1 kN < 1, and therefore

I = N is stable.
Cobwebbing gives the following picture:
0 N
I
n
I
n+1
Note that iterates may
In that case, we may
it implies that everyone
I
n+1
= I
n
be greater than N.
stop the iteration, since
has caught the disease.
The model predicts that as time progresses, everyone catches the disease.
(b) We introduce a new class of individuals, namely those who have been ill, but
now have recovered with immunity. Let the number of such individuals on
day n be R
n
. Then the number of newly infected individuals on day n + 1 is
k
people who are sick
..
I
n
(N I
n
R
n
)
. .
people available to catch disease
.
The number of people who move from the infected to the recovered class on
day n are those that became newly infected d days ago. That is,
kI
nd
(N I
nd
R
nd
).
Putting things together, we obtain
_
I
n+1
= I
n
+ kI
n
(N I
n
R
n
) kI
nd
(N I
nd
R
nd
)
R
n+1
= R
n
+ kI
nd
(N I
nd
R
nd
). sol-gerda
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with
I
d
= = I
2
= I
1
= 0
I
0
= 1
R
d
= = R
1
= R
0
= R
1
= = R
d1
= 0
R
d
= 1
The additional assumption we made is that people remain sick for precisely d
days. That is, all people who become ill at the same time recover at the same
time.
Exercise 2.4.15: Jury conditions
We have
J =
_
a
R
p
R
p
J
a
J
_
.
(a) The characteristic polynomial of J is
det
_
a
R
p
R
p
J
a
J

_
= 0
( a
R
)( a
J
) p
R
p
J
= 0
2
(a
R
+ a
J
) + a
R
a
J
p
R
p
J
= 0
2
tr J + det J = 0,
as required.
(b) The eigenvalues of J are
=

_
2
4
2
,
where := tr J and := det J.
Thus, we need to prove
|| < 1 + < 2 || < 1. (2.4)
We will show both directions of the implication separately.
: Suppose the rst part of (2.4) is satised. We then need to show that both
eigenvalues will have magnitude less than 1. The form of the eigenvalues
tells us that they could be complex with nonzero imaginary part. If this
is the case, we know that the eigenvalues can be rewritten in the form
=
i
_
4
2
2
, (2.5) sol-gerda
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with 4 >
2
.
Thus, for complex eigenvalues,
||
2
=

2
4
+
4
2
4
= < 1,
since 1 + < 2 by assumption. Of course, once we have that ||
2
< 1,
we get automatically that || < 1.
Now that we have the complex case covered, we can assume
2
4, so
that the eigenvalues are real, and proceed the following way:
|| =
2
4
2
||
2
+
|
_
2
4|
2
(triangle inequality, regardless of the sign of the radical)
<
1 + +
_
(1 + )
2
4
2
(left-hand inequality relating and )
=
1 + +
_
(1 )
2
2
=
1 + +|1 |
2
=
1 + + 1
2
(since < 1, |1 | = 1 )
= 1.
Therefore, || < 1 for real or complex eigenvalues, and we have shown
this direction of the implication.
: Suppose now the right hand side of the implication in (2.4) is satised
(that is, || < 1 for both eigenvalues ). We then need to show that
|| < 1 + < 2.
Again we will break this up into the cases of complex with nonzero imag-
inary part, and purely real eigenvalues.
First, if
2
< 4, the eigenvalues will be complex and well get:
|| < 1 ||
2
< 1
2
4
+
4
2
4
< 1
< 1
1 + < 2,
which implies one of the Jury conditions. To show the other condition, sol-gerda
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we start with
2
2 + 1 > 0
2
+ 2 + 1 > 4
( + 1)
2
> 4
( + 1)
2
>
2
(4 >
2
, by assumption)
+ 1 > ||.
Note that the last step is allowed because 4 >
2
0 > 1 (if
1, we would need to put | + 1| > ||). We have now shown the
desired result in the case of complex eigenvalues when 4 >
2
.
In the real case, we substitute in for both of the eigenvalues in the in-
equality || < 1:
_
_
1 <
+
_
2
4
2
< 1,
1 <

_
2
4
2
< 1.
We can then isolate /2:
_
_
1
_
2
4
2
<

2
< 1
_
2
4
2
,
1 +
_
2
4
2
<

2
< 1 +
_
2
4
2
.
Now we can mix these together,
_
_
1 +
_
2
4
2
_
<

2
< 1 +
_
2
4
2
,
_
1
_
2
4
2
_
<

2
< 1
_
2
4
2
.
so as to put it into the following more compact form:
||
2
< 1
_
2
4
2
.
It follows that
1
||
2
>
_
2
4
2
, (2.6)
and since
_
2
4 0, we get
||
2
< 1. (2.7) sol-gerda
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In addition, we can square both sides of (2.6), since they are both posi-
tive, and obtain
1 +

2
4
|| >

2
4

1 || >
|| < 1 + ,
(2.8)
which is one of the Jury conditions. To obtain the other Jury condition,
note that since
2
4, we can write
_
||
2
_
2
. (2.9)
Combining (2.7) and (2.9) gives
_
||
2
_
2
< 1
2
= 1, and hence +1 <
2, which is the second Jury condition. The two Jury conditions together
give
|| < 1 + < 2,
which is what we were trying to prove.
Now putting together the results from and , we have shown (2.4):
|| < 1 + < 2 || < 1.
Exercise 2.4.16: Romeo and Juliet in love/hate-preserving mode
(a)
det A = det
_
a
R
1 p
R
p
J
a
J
1
_
= (a
R
1)(a
J
1) p
R
p
J
= (p
J
)(p
R
) p
R
p
J
= 0, as required
(b)
J =
_
a
R
p
R
p
J
a
J
_
The eigenvalues are given by
det
_
a
R
p
R
p
J
a
J

_
= ( a
R
)( a
J
) p
R
p
J
=
2
(a
R
+ a
J
) + a
R
a
J
p
R
p
J
=
2
(a
R
+ a
J
) + a
R
+ a
J
1 (since det A = (a
R
1)(a
J
1) p
R
p
J
= 0)
= [ 1][ (a
R
+ a
J
1)] = 0, sol-gerda
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That is, = 1 or = a
R
+ a
J
1, as required.
Exercise 2.4.17: Host-parasitoid systems: the Poisson distribution
i=0
P(i) =
i=0
i
e
i!
= e
i=0
i
i!
= e
(Taylor expansion)
= 1
Exercise 2.4.18: Host-parasitoid systems: the Nicholson-Bailey model
(a) Fixed points (H, P) must satisfy
H = kHe
aP
P = cH[1 e
aP
]
The rst equationgives H = 0 or P =
ln k
a
.
Substituting H = 0 into the second equation gives P = 0, thus yielding the
trivial xed point,
(H
1
, P
1
) = (0, 0).
Substituting P =
ln k
a
into the second equation gives H =
k ln k
ac(k1)
, yielding
the nontrivial xed point,
(H
2
, P
2
) =
_
k ln k
ac(k 1)
,
ln k
a
_
.
To ensure P
2
> 0, we require k > 1.
(b)
J(H, P) =
_
ke
aP
akHe
aP
c[1 e
aP
] acHe
aP
_
,
so
J(H
1
, P
1
) = J(0, 0) =
_
k 0
0 0
_
.
The eigenvalues are
1
= k and
2
= 0. Thus, the trivial xed point is stable
when 0 < k < 1, and unstable when k > 1. sol-gerda
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(c)
ln k +
ln k
k 1
> 1
ln k
_
k
k 1
_
> 1
k ln k > k 1
k ln k k + 1 > 0.
Let f(k) = k ln k k + 1. Since f
(k) = 1 + ln k 1 = ln k > 0, we see that

f(k) is a monotonically increasing function for k > 1. Further, f(1) = 0.
thus
k ln k k + 1 > 0,
or
ln k +
ln k
k 1
> 1,
as required.
Exercise 2.4.19: Host-parasitoid systems: the Beddington model
Note that the problem as stated in the text is very challenging. As such, this solution
is not complete, but reects approximately what we would expect a student to be able
to do.
(a) We can nd two trivial xed points by inspection, namely
(

H
1
,

P
1
) = (0, 0)
and
(

H
2
,

P
2
) = (K, 0).
The rst of these xed points represents the situation that both hosts and par-
asitoids are extinct. The second represents the situation that the parasitoids
are extinct, and the host population is at its carrying capacity.
There is a third xed point, representing co-existence. This xed point cannot
be solved for explicitly (our apologies for the misleading wording in the prob-
lem statement). Please consult the paper by Beddington, Free and Lawton
for details on how they handled this case (see text for the complete reference
to the paper).
(b) We can determine the stability of the two trivial xed points found above.
The Jacobian matrix of the Beddington model is
J(

H,

P) =
_
_
_
exp
_
r
_
1

H
K
_
a

P
_ _
1

Hr
K
_
a

H exp
_
r
_
1

H
K
_
a
P
_
c(1 e
a
P
) ac

He
a
P
_
_
_. sol-gerda
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The stability of (

H
1
,

P
1
) = (0, 0) is determined by
J(0, 0) =
_
e
r
0
0 0
_
.
Here, tr J = e
r
and det J = 0. Since r > 0 (why?), tr J > 1, and hence the
xed point (

H
1
,

P
1
) = (0, 0) is unstable.
The stability of (

H
2
,

P
2
) = (K, 0) is determined by
J(K, 0) =
_
1 r aK
0 acK
_
.
Here, tr J = 1 r +acK and det J = acK(1 r). Depending on the value of
the model parameters, this xed point may be stable or unstable. Stability is
guaranteed by the Jury conditions, which are
|1 r + acK| < 1 + acK(1 r) < 2.
Determining the conditions that guarantee stability of the steady state rep-
resenting co-existence is beyond the scope of this book and solution manual.
The reader is encouraged to consult the paper by Beddington, Free and Law-
ton for details. It turns out that the co-existence xed point is stable in a
large region of parameter space, especially for small values of r. For large
values of r, the xed point usually is unstable (depending on the values of the
other model parameters).
(c) The numerical experiment is left to the reader. Lots of interesting patterns
are possible depending on the choice of parameters. It is especially revealing
to plot the orbits in the (H, P) phase plane. We refer the reader to the book
Mathematical Models in Biology by Leah Edelstein-Keshet for a nice discussion
of a variety of numerical solutions. sol-gerda
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3.9 Exercises for ODEs
Exercise 3.9.1: The C
14
-method
Let C(t) be the concentration of carbon at any time t. We use the equation described
in the chapter on ODEs,
C
(t) = k C(t),
where k > 0 is a constant. This equation has the solution
C(t) = Ae
kt
,
where A is a constant of integration. From the initial condition, C(0) = c
0
, we nd
that A = c
0
, so the equation we get for C is
C(t) = c
0
e
kt
.
We know by denition that C(T
1/2
) =
1
2
c
0
= c
0
e
kT
1/2
. Solving for k, we nd
k =
log(2)
T
1/2
.
The wood now has 75% of its original concentration. Call the time T
3/4
. Currently,
we have C(T
3/4
) =
3
4
c
0
. We set this value for the function and solve for T
3/4
:
3
4
c
0
= c
0
e
kT
3/4
log(4/3) = kT
3/4
T
3/4
= log(4/3)
1
k
= T
1/2
log(4/3)
log(2)
= 2390 a
The model tells us that this piece of wood is about 2390 years old. This is about
1000 years after Tutankhamens time (about 3300 years ago), so Tutankhamen could
not have sat in a boat made of wood from the same tree that this piece came from.
Exercise 3.9.2: Learning curves
(a) dP/dt represents the rate of change of the performance over time, or how fast
someone picks up a skill.
(b) When M P, dP/dt 0, so P(t) is increasing or staying constant in time.
If M < P, then dP/dt < 0, which means that P(t) is decreasing in time.
We expect that with more and more training, a person will never have a sol-gerda
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decrease in performance. Notice that if a we start with P below M, P can
never get larger than M. If P = M, P will remain constant. This model is
reasonable. We interpret M as the level when someone has mastered the skill
(M for master). A reasonable initial condition could be P(0) = 0; no previous
knowledge.
0
0
graph of dP/dt versus P
P
d
P
/
d
t

M
kM
(c)
0
0
graph of P versus t
t
P
(
t
)
P(0) = 0
P(0) > M
M
Exercise 3.9.3: Harvesting
The dierential equation we have is
u
(t) = g(u), sol-gerda

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where g(u) = au(1 u/K) cu. Rewriting, we get
g(u) = u
_
(a c)
a
K
u
_
.
We can see that the graph of g(u) vs u is a parabola. Since a, K > 0, the parabola
opens downwards. It has intersections with the u axis at u = 0 and at u =
K
a
(ac).
We refer to u = 0 as the trivial steady state, and u =
K
a
(a c) as the nontrivial
steady state.
There are 3 dierent scenarios which will give three dierent qualitative sketches.
Case (i): a < c. This means that the nontrivial steady state is negative. There
is only one biologically realistic steady state, namely the trivial steady state
u = 0.
0
0
case(i): a < c
u
d
u
/
d
t

The phase portrait shows that the trivial steady state u = 0 is stable.
Case (ii): a = c. Now there is just one intersection, at the origin. Hence there
is only one steady state, namely the trivial steady state u = 0. sol-gerda
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0
0
case(ii): a = c
u
d
u
/
d
t

The phase portrait shows that u = 0 is stable (for u > 0).
Case (iii): a > c. Both steady states are biologically realistic.
0
0
case(iii): a > c
u
d
u
/
d
t

The phase portrait shows that the nontrivial steady state at u =
K
a
(a c) is
stable, and the trivial one at u = 0 is unstable.
In summary, the nontrivial steady state exists only for a > c, and it is stable. If
a c, there is only the trivial steady state, and it is stable.
Biologically, this means that when the population is not reproducing faster than it
is being harvested, it will die out. If it is reproducing fast enough, it settles down sol-gerda
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to a steady state, determined by how much faster it is reproducing than it is being
harvested (the larger the dierence, the larger the population at the steady state).
Exercise 3.9.4: Fishing
(a) 1: The shing term says that there is a constant number of sh per unit
time being removed due to shing. This number is H
1
, in the units of
sh per unit time.
2: Here the shing term says that the number of sh which are shed out
per unit time is proportional to the current population. H
2
, which is the
fraction of sh caught per unit time, must be between 0 and 1, since you
cannot catch more than all of the sh, or fewer than none of them. This
model reects the fact that sh must be found to be caught. A certain
fraction of them can be found.
3: In the third model, the limits of small and large values for N reduce to
models 2 and 1, respecively. As N 0, the shing term, H
3
N/(A+N),
goes to H
3
N/A. This is the same situation as model 2, but with H
2
= H
3
/A. That is, for small amounts of sh, the number of sh caught
per unit time it proportional to the population. More sh implies more
shing, and fewer sh implies less shing. As N , the shing term
will go to H
3
, a constant, which works the same way as H
1
in model 1.
That is, when there are more sh than can be caught, or more sh than
are wanted, there is just a constant amount of sh being caught per unit
time. A determines how fast this limit of too many sh is eectively
reached. When A is higher, model 3 stays at model 2 for a longer time.
When A is lower, model 3 turns into model 1 sooner. We can see that
A is actually in the units of sh. When N = A, we see that the shing
term becomes H
3
/2, giving an idea of how fast we are approaching the
limit of H
3
, in terms of N.
(b) Model 1 is not biologically realistic because a constant number of sh caught
per unit time regardless of how many sh there are makes no sense. For exam-
ple, suppose there are 10 sh in the pond, and H
1
= 3 sh/hour. Neglecting
the rst term in the model equation, there is no way that this model can work
after 3 hours, because there will be only 1 sh left. Model 1 would work only
if there was an abundance of sh. When the abundance becomes less, and the
sh population becomes closer to the value H
1
, the model breaks down.
(c) Model 3 is a better model, because, as explained in part (a), it takes the
best parts of both models. We see that model 3 is precisely the modication
to model 1 desired in part (b). The problem with model 2 is that there is
a limited number of sh which can actually be caught. Model 3 says what
happens when this limit is reached. For example, say you have a boat which
can catch 50 sh per hour. We use model 2. Say that H
2
= 0.5, and that
initially, N = 100. After an hour, you will have caught 50 sh. After 2 hours, sol-gerda
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you will have caught 25 more sh, etc. If, on the other hand, you start shing
at N = 1000, then model 2 says you will have caought 500 sh after an hour,
even though your boat can catch only 50 sh per hour. Model 3 corrects
this and says that there is a constant number of sh that will be caught,
when the population is high. To a close approximation, H
3
represents the
combined eects of people who sh with limited boat capacities, or limited
desired quantities of sh, and puts it into a single term; one big boat which
can catch at the most ||H
3
|| sh per hour, where ||H
3
|| means the magnitude
of H
3
, without the units.
Exercise 3.9.5: A metapopulation model
Thanks to Pandora Lam, University of Alberta, for providing this solution.
(a) We rst rewrite P
:
P
= cP(h P) P
= chP cP
2
P
= (ch )P cP
2
= P((ch ) cP)
and then set P
= 0 to nd the steady states, namely P = 0 and P =

ch
c
=
h

c
.
Note that the steady state P = 0 always exists. The steady state P = h

c
exists only if h

c
> 0 or h >

c
.
(b) Let f(P) = cP(h P) P. Then f
(P) = ch 2cP.
The stability of the steady state P = 0 is determined by f
(0) = ch .
If h >

c
, f
(0) > 0, and P = 0 is a stable steady state.

If h <

c
, f
(0) < 0, and P = 0 is an unstable steady state.

The stability of the steady state P = h

c
is determined by
f
(h

c
) = ch 2c(h

c
) = (ch ).
Recall that this steady state exists only if h >

c
, so that f
(h
c
) < 0, which
implies that the steady state is stable.
The bifurcation diagram follows: sol-gerda
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0
0
bifurcation diagram
h
P
P = 0
P = h /c
/c
In conclusion, the population dies out as soon as the number of habitable
patches falls below

c
.
Note that the horizontal line in the bifurcation diagram should be dotted for
h > /c.
Exercise 3.9.6: Gene activation
(a) The rst term is a constant growth term. The growth of g depends linearly
on the concentration of S, with a growth rate k
1
. In this case, it is a constant
growth, because s
0
is constant.
The second term is the genes natural decay term. With nothing else, the
gene would decay exponentially at a rate k
2
.
The third term is a self-production term, with a limited rate of reproduction
of k
3
. When g gets large, this term becomes approximately constant, keeping
a the k
3
factor (lim
g
g
2
k
2
4
+g
2
= 1). The parameter k
4
determines how large
g has to be before this term starts behaving as in the large g limit.
(b) We start with the rst equation,
dg
dt
= k
1
s
0
k
2
g +
k
3
g
2
k
2
4
+ g
2
.
In order to get the last term to look like x
2
/(1+x
2
), we must divide everything
by k
3
, and then cleverly mulitply the last term by 1, as in
1/k
2
4
1/k
2
4
= 1. We then
get:
1
k
3
dg
dt
=
k
1
s
0
k
3
k
2
k
3
g +
(g/k
4
)
2
1 + (g/k
4
)
2 sol-gerda
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It seems logical to dene x := g/k
4
, so that the last term is correct. We now
must think about how we can dene in order to make the dx/d =
1
k3
dg/dt.
To get just a single
1
k3
as the dierence, well need =
k3
k4
t, to cancel out the
k
4
s and to bring in the
1
k3
. Since x and are just scalar mulitples of g and
t, respectively, we can pull the scalars out of the dierential operator, so that
dx
d
=
d(g/k
4
)
d(k
3
t/k
4
)
=
1/k
4
k
3
/k
4
dg
dt
=
1
k
3
dg
dt
.
Hence,
dx
d
=
1
k
3
dg
dt
=
k
1
s
0
k
3
k
2
k
3
g +
(g/k
4
)
2
1 + (g/k
4
)
2
=
k
1
s
0
k
3
k
2
k
4
k
3
x +
x
2
1 + x
2
= s rx +
x
2
1 + x
2
,
with s := k
1
s
0
/k
3
, and r := k
2
k
4
/k
3
.
(c) We really need to graph only two more cases; when s is such that there are
two intersections of the horizontal axis, and when s is high enough that there
are no intersections of the horizontal axis.
0 0.5 1 1.5 2 2.5
0.15
0.1
0.05
0
0.05
0.1
0.15
0.2
0.25
graph of dx/d versus x
x
d
x
/
d
s = 0
s = 0.02
s = 0.1
s = 0.04
(d) We note that a bifurcation in the number of steady states occurs at s* 0.04.
For s < s*, there are three steady states; the outer two steady states are
stable, and the inner steady state is unstable.
For s > s*, there is one stable steady state.
The bifurcation diagram follows: sol-gerda
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0 0.05 0.1 0.15 0.2 0.25 0.3
0
0.5
1
1.5
2
2.5
3
qualitative sketch of bifurcation diagram
x
s
X: 0.04183
Y: 0.2198
Exercise 3.9.7: Linear systems
(a)
A =
_
1 1
3 1
_
tr A = a + d = 1 + (1) = 0
det A = ad bc = 1 (1) 1 3 = 4 < 0
1
,
2
=
tr A
2

1
2
_
(tr A)
2
4 det A = 0
1
2
_
0 4 (4) =
1
2
4 = 2
Hence, (0, 0) is a saddle point. sol-gerda
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5000
10000
15000
20000
25000
30000
35000
y2
5000 10000 15000 20000 25000 30000 35000
y1
(b)
A =
_
2 1
2 3
_
tr A = 2 + 3 = 5 > 0
det A = ad bc = 2 (3) 1 2 = 4 > 0
1
,
2
=
tr A
2

1
2
_
(tr A)
2
4 det A =
5
2

1
2
_
25 4 (4) =
5
2

3
2
= 4, 1
Hence, (0, 0) is an unstable node.
0
1000
2000
3000
4000
5000
y2
500 1000 1500 2000 2500
y1 sol-gerda
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(c)
A =
_
1 2
2 1
_
tr A = 1 + (1) = 2 < 0
det A = ad bc = 1 (1) (2) 2 = 5 > 0
1
,
2
=
tr A
2

1
2
_
(tr A)
2
4 det A =
2
2

1
2
_
4 4 (5) = 1
1
2
16 = 1 2i
Hence, (0, 0) is a stable spiral.
0.2
0
0.2
0.4
0.6
0.8
1
1.2
y2
0.5 0.5 1 1.5 2
y1
(d)
A =
_
1 2
2 1
_
tr A = 1 + 1 = 2 > 0
det A = ad bc = 1 1 2 (2) = 5 > 0
1
,
2
=
tr A
2

1
2
_
(tr A)
2
4 det A =
2
2

1
2
_
4 4 (5) = 1
1
2
16 = 1 2i
Hence, (0, 0) is an unstable spiral. sol-gerda
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100
50
0
50
100
150
y2
300 250 200 150 100 50 50
y1
(e)
A =
_
0 2
2 0
_
tr A = 0 + 0 = 0
det A = ad bc = 0 0 (2) 2 = 4 > 0
1
,
2
=
tr A
2

1
2
_
(tr A)
2
4 det A = 0
1
2
_
0 4 (4) =
1
2
16 = 2i
Hence, (0, 0) is a center.
2
1
1
2
y2
2 1 1 2
y1 sol-gerda
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Exercise 3.9.8: A linear system with complex eigenvalues
We need to show that both x
(1)
(t) and x
(2)
(t) satisfy the dierential equation
d
dt
_
x
1
x
2
_
=
_

_ _
x
1
x
2
_
.
If we let
_
x
1
x
2
_
= x
(1)
(t) =
_
e
t
cos t
e
t
sint
_
,
then
d
dt
_
x
1
x
2
_
=
_
e
t
cos t e
t
sin t
e
t
sint e
t
cos t
_
=
_
(e
t
cos t) + (e
t
sin t)
(e
t
cos t) + (e
t
sin t)
_
=
_

_ _
e
t
cos t
e
t
sin t
_
=
_

_ _
x
1
x
2
_
,
as required.
Similarly, if we let
_
x
1
x
2
_
= x
(2)
(t) =
_
e
t
sin t
e
t
cos t
_
,
then
d
dt
_
x
1
x
2
_
=
_
e
t
sin t + e
t
cos t
e
t
cos t e
t
sin t
_
=
_
(e
t
sin t) + (e
t
cos t)
(e
t
sin t) + (e
t
cos t)
_
=
_

_ _
e
t
sin t
e
t
cos t
_
=
_

_ _
x
1
x
2
_
,
as required.
We now let x(t) = c
1
x
(1)
(t) + c
2
x
(2)
(t), and rewrite x(t) in the required form as sol-gerda
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follows:
x(t) = c
1
x
(1)
(t) + c
2
x
(2)
(t)
= c
1
_
e
t
cos t
e
t
sint
_
+ c
2
_
e
t
sin t
e
t
cos t
_
= e
t
_
c
1
cos t + c
2
sin t
c
1
sint + c
2
cos t
_
.
Introducing a and such that c
1
= a cos() and c
2
= a sin(), we get
x(t) = ae
t
_
cos t cos() + sin t sin()
(sin t cos() cos t sin())
_
= ae
t
_
cos(t + )
sin(t + )
_
.
Note that c
1
= a cos() and c
2
= a sin() imply
c
2
1
+ c
2
2
= a
2
cos
2
() + a
2
sin
2
() = a
2
,
or
a =
_
c
2
1
+ c
2
2
and
a sin()
a cos()
= tan() = tan() =
c
2
c
1
,
or
= arctan(
c
2
c
1
).
Exercise 3.9.9: The trace-determinant formula
Given a matrix,
A =
_
a b
c d
_
,
The eigenvalues of A are the satisfying |I A| = 0, where I is the 2 2 identity
matrix. Notice that tr(A) = a + d, and det(A) = ad bc. Hence,
0 = |I A|
= ( a)( d) bc
=
2
+ (a d) + ad bc
=
2
tr(A) + det(A).
From the quadratic formula, we nd
1/2
=
tr(A)
_
(tr(A))
2
4 det(A)
2
. sol-gerda
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Q.E.D.
Exercise 3.9.10: Using the trace-determinant formula
(a)
A =
_
1 5
3 2
_
tr A = a + d = 1 + 2 = 3
det A = ad bc = 1 2 5 3 = 13 < 0
1
,
2
=
tr A
2

1
2
_
(tr A)
2
4 det A =
3
2

1
2
_
9 4 (13) =
3
2

1
2
61 5.41, 2.41
Hence, (0, 0) is a saddle point.
(b)
A =
_
0 2
1 3
_
tr A = a + d = 0 + (3) = 3 < 0
det A = ad bc = 0 (3) 1 (2) = 2 > 0
1
,
2
=
tr A
2

1
2
_
(tr A)
2
4 det A =
3
2

1
2
_
9 4 (2) =
3
2

1
2
= 1, 2
Hence, (0, 0) is a stable node.
(c)
A =
_
2 4
3 4
_
tr A = a + d = 2 + 4 = 2 > 0
det A = ad bc = 2 4 4 (3) = 4 > 0
1
,
2
=
tr A
2

1
2
_
(tr A)
2
4 det A =
2
2

1
2
_
4 4 (4) = 1
1
2
12 = 1
3i
Hence, (0, 0) is an unstable spiral.
(d)
A =
_
2 1
1 3
_
tr A = a + d = 2 + 3 = 5 > 0
det A = ad bc = 2 3 1 1 = 5 > 0
1
,
2
=
tr A
2

1
2
_
(tr A)
2
4 det A =
5
2

1
2
_
25 4 (5) =
5
2

1
2
5 3.62, 1.38
Hence, (0, 0) is an unstable node. sol-gerda
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(e)
A =
_
2 1
1 2
_
tr A = a + d = 2 + 2 = 0
det A = ad bc = 2 2 (1) 1 = 3 < 0
1
,
2
=
tr A
2

1
2
_
(tr A)
2
4 det A = 0
1
2
_
0 4 (3) =
1
2
12 =
3
Hence, (0, 0) is a saddle point.
(f)
A =
_
1 2
2 1
_
tr A = a + d = 1 + 1 = 0
det A = ad bc = 1 1 (2) 2 = 3 > 0
1
,
2
=
tr A
2

1
2
_
(tr A)
2
4 det A = 0
1
2
_
0 4 (3) =
1
2
12 =
3i
Hence, (0, 0) is a center.
Exercise 3.9.11: Two-population model
Thanks to Pandora Lam, University of Alberta, for providing the outline of this
solution.
The two-population model, (3.8), is
x = x + xy,
y = y + xy.
There are two steady states, namely P
1
= (0, 0) and P
2
= (
).
In the solutions shown below, we determine the stability of any biologically relevant
steady states. Note that P
1
always is biologically relevant. However, P
2
only is
biologically relevant if and as well as and have opposite signs.
Knowing the stability of the steady states will be helpful in sketching the phase
portraits, not (yet) provided here.
The Jacobian matrix for the system is
J =
_
f1
x
f1
y
f2
x
f2
y
_
=
_
+ y x
y + x
_
. sol-gerda
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In general then, the stability of P
1
= (0, 0) is determined by
J(0, 0) =
_
0
0
_
,
with eigenvalues
1
= and
2
= .
Similarly, the stability of P
2
= (
) is determined by
J(
) =
_
0

0
_
,
with tr J = 0 and det J = .
(a) Case > 0, > 0, > 0, < 0
For P
1
= (0, 0):
The eigenvalues are
1,2
> 0, therefore P
1
= (0, 0) is an unstable node.
For P
2
= (
):
Since and have the same sign, P
2
is not biologically relevant.
INSERT PHASE PORTRAIT HERE
Biological interpretation: We have a predator-prey model . . .
(b) Case > 0, > 0, < 0, < 0
For P
1
= (0, 0):
The eigenvalues are
1
= > 0 and
2
= < 0, therefore P
1
= (0, 0) is a
saddle point.
For P
2
= (
):
P
2
(c) Case < 0, > 0, < 0, < 0
For P
1
= (0, 0):
The eigenvalues are
1
= < 0 and
2
= < 0, therefore (0, 0) is a stable
node.
For P
2
= (
):
P
2
INSERT PHASE PORTRAIT HERE sol-gerda
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(d) Case > 0, > 0, > 0, > 0
For P
1
= (0, 0):
The eigenvalues are
1
= > 0 and
2
= > 0, therefore (0, 0) is an unstable
node.
For P
2
= (
):
P
2
Biological interpretation: We have a mutualism or symbiosis model . . .
(e) Case > 0, > 0, < 0, > 0
For P
1
= (0, 0):
The eigenvalues are
1
= > 0 and
2
= < 0, therefore (0, 0) is a saddle
point.
For P
2
= (
):
P
2
Biological interpretation: We have a mutualism or symbiosis model . . .
(f) Case > 0, < 0, > 0, < 0
For P
1
= (0, 0):
The eigenvalues are
1
= > 0 and
2
= > 0, therefore (0, 0) is an unstable
node.
For P
2
= (
):
P
2
IS biologically relevant! Since tr J = 0 and det J = < 0, P
2
=
(
) is a saddle point.
Biological interpretation: We have a competition model . . .
(g) Case < 0, < 0, < 0, < 0
For P
1
= (0, 0):
The eigenvalues are
1
= < 0 and
2
= < 0, therefore (0, 0) is a stable
node.
For P
2
= (
):
P
2
INSERT PHASE PORTRAIT HERE sol-gerda
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Biological interpretation: We have a competition model . . .
Exercise 3.9.12: Predator-prey model
(a) Let x(t) be the prey population, and y(t) be the natural predator population.
Assuming exponential growth for the prey population in the absense of the
predator, and exponential decay for the predator population in the absense of
prey, the 2-species interaction model reads
dx
dt
= x xy,
dy
dt
= y + xy.
(b) Let r
1
be the rate that the poison kills the prey population, and r
2
be the
rate that the poison kills the predator population.
The new model then reads
dx
dt
= x xy r
1
x,
dy
dt
= y + xy r
2
y.
Exercise 3.9.13: Inhibited enzymatic reaction
Let s = [S], e = [E], b
1
= [B
1
], q = [Q], b
2
= [B
2
], and i = [I].
The rst reaction gives the following dierential equations:
ds
dt
= k
1
se + K
1
b
1
,
de
dt
= k
1
se + K
1
b
1
+ k
2
b
1
,
db
1
dt
= k
1
se K
1
b
1
k
2
b
1
,
dq
dt
= k
2
b
1
. sol-gerda
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The second gives the following three equations:
db
2
dt
= k
1
ei k
1
b
2
,
de
dt
= k
1
ei + k
1
b
2
,
di
dt
= k
1
ei + k
1
b
2
.
Exercise 3.9.14: A feedback mechanism for oscillatory reactions
We are given the following pathway:
A
k
1
k
1
B
k
2
k
2
C
k
3
k
3
A.
Let a = [A], b = [B], and c = [C].
A dierential equation model for the above pathway then is
da
dt
= k
1
b + k
3
c k
1
a k
3
a,
db
dt
= k
1
a + k
2
c k
1
b k
2
b,
dc
dt
= k
2
b + k
3
a k
2
c k
3
c.
Exercise 3.9.15: Enzymatic reaction with two intermediate steps
We are given the following reaction:
S + E
k
1
k
1
C
1
k
2
k
2
C
2
k
3
k
3
E + P.
Let s = [S], e = [E], c
1
= [C
1
], c
2
= [C
2
], and p = [P]. sol-gerda
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A dierential equation model for the above reaction then is
ds
dt
= k
1
c
1
k
1
se,
de
dt
= k
1
c
1
+ k
3
c
2
k
1
se k
3
ep,
dc
1
dt
= k
1
se + k
2
c
2
k
1
c
1
k
2
c
1
,
dc
2
dt
= k
2
c
1
+ k
3
ep k
2
c
2
k
3
c
2
,
dp
dt
= k
3
c
2
k
3
ep.
Exercise 3.9.16: Self-intoxicating population
We are working with the following system:
n = ( Ky)n,
y = n y.
To avoid having to consider all sorts of special cases in the solution below, we assume
, , , , K > 0 instead of , , , , K 0.
(a) The term n represents birth, increasing the population.
The term n represents natural death, decreasing the population.
The term Kyn represents death due to a toxic environment, decreasing the
population.
The term n represents the production of waste products, proportional to the
size of the population.
The term y represents natural degradation of the waste products.
(b) We begin with the nullclines.
There are two n-nullclines, given by n = 0, namely the vertical line
n = 0
and the horizontal line
y =

K
.
Similarly, there is one y-nullcline, given by y = 0, namely the straight line
passing through the origin (with positive, nite slope /)
y =

n. sol-gerda
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We now nd steady states by looking for all the intersections of an n-nullcline
with a y-nullcline.
The intersection of the rst n-nullcline, n = 0, and the y-nullcline is given by
the solution of n = 0 and y = n/, that is, at
P
1
:= (n, y) = (0, 0).
The intersection of the second n-nullcline, y = ()/K, and the y-nullcline
is given by the solution of y = ( )/K and y = n/, that is, at
P
2
:= (n, y) =
_

K
,

K
_
.
We will refer to P
1
as the trivial steady state and P
2
as the nontrivial (co-
existence) steady state. Note that P
2
is biologically relevant only provided
> .
We think it doesnt make sense to sketch a phase portrait here (since there are
too many cases, and not all information has been determined yet). It should
come later, in part (e).
(c) We think it doesnt make sense to sketch a vector eld here (since there are
too many cases, and not all information has been determined yet). It should
come later, in part (e).
(d) The Jacobian matrix of the system is
J(n, y) =
_
f1
n
f1
y
f2
n
f2
y
_
=
_
Ky Kn

_
The stability of P
1
is determined by
J(0, 0) =
_
0

_
.
The eigenvalues of J(0, 0) are
1
= and
2
= < 0.
If < , then P
1
is the only biologically relevant steady state. In this case,
1
< 0, and P
1
is a stable node.
If > , then both steady states are relevant. In this case,
1
> 0, and P
1
is
a saddle point.
Similarly, the stability of P
2
is determined by
J
_

K
,

K
_
=
_
0
( )

_
.
We have tr J = < 0 and det J = ( ). sol-gerda
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If < , then det J < 0, and P
2
is a saddle point (but in this case, P
2
is not
biologically relevant).
If > , then det J > 0, and P
2
is either a stable node or a stable spiral.
To summarize what we have so far:
If < , then P
1
is the only relevant steady state, and it is a stable node.
If > , then both P
1
and P
2
are biologically relevant. In this case, P
1
is a
saddle point, and P
2
is a stable node or a stable spiral.
For P
2
to be a stable node, we need (tr J)
2
4 det J > 0, that is
2
4() >
0, or > 4( ) > 0.
Similarly, for P
2
to be a stable spiral, we need (tr J)
2
4 det J < 0, or
< 4( ).
(e) Here we look at one of the cases determined above, namely when < 4().
In this case, P
1
is a saddle point, and P
2
is a stable spiral.
INSERT VECTOR FIELD AND PHASE PORTRAIT HERE
Interpretation in terms of the biology: Starting from any initial population
(other than zero), the population and amount of toxicity eventually reach a
steady state. That is, under ideal conditions (no stochasticity), the population
persists, no matter how much waste it produces. The steady state is reached
in a damped oscillatory fashion. However, depending on the initial conditions,
trajectories may pass close to the rst n-nullcline, n = 0. When this happens,
n is very small. That is, in the presence of stochastic events, the population
could become extinct.
(f) Solution not available.
Exercise 3.9.17: Fish populations in a pond
(a) Exponential growth:
dT
dt
= r
T
T
(b) Growth with competition:
dT
dt
= (mB + r
T
)T
(c) Exponential growth:
dB
dt
= r
B
b
Growth with competition:
dB
dt
= (nT + r
B
)B sol-gerda
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(d) Solution not available.
(e) We get the system
dT
dt
= r
T
T mBT,
dB
dt
= r
B
B nBT.
The steady states are determined by dT/dt = dB/dt = 0. This means that
any steady state (

T,

B) must satisfy
r
T

T = m

B

T,
r
B

B = n

B

T.
Therefore, we get the trivial steady state,
(
T,

B) = (0, 0),
and the nontrivial steady state,
(

T,

B) = (
r
B
n
,
r
T
m
).
The jacobian matrix of this system, evaluated at the nontrivial steady state,
is
J
_
r
B
n
,
r
T
m
_
=
_
_
r
T
m

B m
T
n

B r
B
n
T
_
_
=
_
_
0
mrB
n
nrT
m
0
_
_
.
Exercise 3.9.18: Exact solution for the logistic equation
(a) We have
N
= N
_
1
N
K
_
, N(0) = N
0
.
Solution method 1: We recognize the dierential equation as a separable
equation, so that we can write
_
N(t)
N0
d

N
N
_
1

N
K
_ =
_
t
0
d
t.
Using partial fractions, we can rewrite the left hand side:
_
N(t)
N0
_
1
N
+
1
K
1

N
K
_
d

N =
_
t
0
d
t. sol-gerda
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We integrate to obtain
_
ln

N ln
_
1
N
K
__
N(t)
N0
= t
ln
_

N
1

N
K
_
N(t)
N0
= t
ln
_
N(t)
1
N(t)
K
_
ln
_
N
0
1
N0
K
_
= t
ln
_
_
_
1
N0
K
_
N(t)
_
1
N(t)
K
_
N
0
_
_
= t.
Exponentiating both sides and rearranging gives
K N
0
K N(t)
N(t) = N
0
e
t
(K N
0
)N(t) = N
0
e
t
(K N(t))
(K N
0
+ N
0
e
t
)N(t) = N
0
Ke
t
N(t) =
N
0
Ke
t
K N
0
+ N
0
e
t
=
e
t
N
0
1 +
N0
K
(e
t
1)
.
Solution method 2: Let u =
1
N
. Then N =
1
u
and
dN
dt
=
1
u
2
du
dt
.
Substitution into the logistic equation gives
1
u
2
du
dt
=
1
u
_
1
1
K
1
u
_
du
dt
=
_
1
K
u
_
. sol-gerda
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We separate variables and integrate, as follows:
_
u(t)
u0
d u
1
K
u
=
_
t
0
d
t
ln
_
1
K
u
_
u(t)
u0
= t
ln
_
1
K
u(t)
_
+ ln
_
1
K
u
0
_
= t
ln
_
1
K
u
0
1
K
u(t)
_
= t
1 Ku
0
1 Ku(t)
= e
t
1 Ku(t) = (1 Ku
0
)e
t
u(t) =
1
K
_
1 (1 Ku
0
)e
t
.
Now we return to original variables, as follows:
1
N(t)
=
1
K
_
1
_
1 K
1
N
0
_
e
t
_
N(t) =
K
1
_
1
K
N0
_
e
t
=
Ke
t
e
t
1 +
K
N0
=
e
t
N
0
N0
K
e
t
N0
K
+ 1
=
e
t
N
0
1 +
N0
K
(e
t
1)
.
(b) This solution is of the same form as that of the Beverton-Holt model, except
we have e
t
in place of r
n+1
. sol-gerda
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4.5 Exercises for PDEs
Exercise 4.5.1: Diusion through a membrane
u
t
= Du
xx
, u
t
= 0
(a) u
xx
= 0 u
x
= const = c u(x) = cx + d
Boundary conditions:
u(0) = c
1
d = c
1
u(L) = c
2
cL + c
1
= c
2
c =
c
2
c
1
L
Solution:
u(x) =
c
2
c
1
L
x + c
1
For c
2
> c
1
:
0 L
x
u(x)
outside inside
u(0) = c
1
u(L) = c
2
(b) J(x) = D

x
u(x) = D
c2c1
L
=
D
L
(c
2
c
1
). The ux is proportional to the
concentration dierence. The proportionality factor
D
L
is called permeability.
Exercise 4.5.2: Fundamental solution
Exercise 4.5.3: Signalling in ant populations
u
t
= Du
xx
, u(0) =
0
(x), D = 1 (4.10)
(a) Fundamental solution of {u
t
= Du
xx
, u(0) =
0
(x)} is g(x) =
1
2t
e
x
2
4t
.
Hence u(x) = g(x) solves (4.10). sol-gerda
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At x(t): u(x(t)) = 0.1 = g(x)
g(x) =
1
10
, e
x
2
4t
=
2t
10
, e
x
2
4t
=
10
2t
, x
2
= 4t ln
_
10
2t
_
x(t) =
4t ln
_
10
2t
_
(b)
0 5 10 15
t
0
1
2
3
x(t)
Range of Influence
(c) x(t) dened only for ln
_
10
2t
_
> 0, hence
10
2t
> 1. So
10 >
2t, 100 > 2t,

50
> t
t
=
50
15.9
Exercise 4.5.4: Dingos in Australia
(Thanks to Dr. Markus Owen (Nottingham), who used this problem in one of his
Math-bio classes).
u
t
= Du
xx
+ ku(1 u), k = 1
(a) D
1
= 100, wave speed of a travelling wave,
c
= 2
_
D
1
f
(0), f
(0) = k = 1
= 2
_
D
1
= 20
_
miles
month
_
distance = 100 miles T =
100 miles
c
=
100
20
= 5 months. sol-gerda
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The decay rate of this wave front is:
1
=
c
2D
1
=
10
100
=
1
10
,
The wave looks like e
1
10
x
near farm A.
e
(x/10)
c*
(b) Between A and B: D
2
= 50
Decay rate
1
=
1
10
=
c
2D2
wave speed = c =
1
2D
2
=
1
10
2 50 = 10
T
2
= 10 months from farm A to B.
c
A B
Exercise 4.5.5: Signal transport in the axon
u
t
= u
xx
+ u(1 u)(u
1
2
)
u
x
(t, 0) = 0, u
x
(t, l) = 0
0 l
(a) Steady states: u
t
= 0. Introduce v := u
x
.
u
x
= v
v
x
= u(1 u)(u
1
2
) = u
3
3
2
u
2
+
1
2
u
(b) equlibria of (a): v = 0, u = 0, 1,
1
2
. sol-gerda
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Jacobian:
Df(u, v) =
_
_
0 1
3u
2
3u +
1
2
0
_
_
Df(0, 0) =
_
_
0 1
1
2
0
_
_
, tr(Df(0, 0)) = 0, det(Df(0, 0)) < 0 saddle
Df(
1
2
, 0) =
_
_
0 1
3
4

3
2
+
1
2
0
_
_
=
_
_
0 1
1
4
0
_
_
tr(Df(
1
2
, 0)) = 0, det(Df(
1
2
, 0)) > 0 center
Df(1, 0) =
_
_
0 1
1
2
0
_
_
, tr(Df(0, 0)) = 0, det(Df(0, 0)) < 0 saddle
(c) Hamilton function if
d
dx
H(u, v) = 0 and u
x
=
H
v
, v
x
=
H
u
,
Here H(u, v) =
1
2
(v)
2
1
4
u
4
+
1
2
u
3
1
4
u
2
.
Lets check:
H
v
= v = u
x
H
u
= u
3
3
2
u
2
1
2
u = v
x
d
dx
H(u, v) =
H
u
du
dx
+
H
v
dv
dx
= v
x
u
x
+ u
x
v
x
= 0.
(d) sol-gerda
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0 1/2 1
u
v
(e) Neumann boundary conditions:
v(0) = 0 v(l) = 0
Following candidates in the phase-portrait of (d):
1/2
u
v
I
1/2
u
v
II
1/2
u
v
III
etc.
As functions of x:
x
u(x)
0 l
I
x
u(x)
0 l
II
x
u(x)
0 l
III
etc. sol-gerda
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x
u(x)
l 0
a = 9
a = 8 + 1
(4.11)
(f) Solution not available.
Exercise 4.5.6: Separation
Exercise 4.5.7: Linear transport
Exercise 4.5.8: Correlated random walk
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5.8 Exercises for Stochastic Models
Exercise 5.8.1: Forest ecosystem succession
Exercise 5.8.2: Princeton forest
Exercise 5.8.3: Mean and variance for a sum of random variables
Exercise 5.8.4: Mean and variance for a negative binomial distribution
Exercise 5.8.5: Random walk derivation of a diusion-advection equation
Exercise 5.8.6: Spatially varying diusion model
Exercise 5.8.7: Variance for a branching process
Exercise 5.8.8: The survival of right whales
Exercise 5.8.9: An explicit solution for the pure birth process
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6.6 Exercises for Cellular Automata
Exercise 6.6.1: Wolfram rule 108
Exercise 6.6.2: Monotonous automaton
Exercise 6.6.3: Greenberg-Hastings automata
Exercise 6.6.4: Game of life
Exercise 6.6.5: Boundaries of nite grids
Exercise 6.6.6: Stochastic epidemic automaton
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7.7 Exercises for Parameter Estimation
Exercise 7.7.1: Maximum likelihood estimation for binomial distribution
Exercise 7.7.2: Maximum likelihood estimation for exponential distri-
bution
Exercise 7.7.3: Hill-climbing algorithm

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sol-gerda

(x) = 1 k. We found in part (a) that

(x) [0, 1), which means that this xed point,

(x) = 1 + 0.0001(5100 2x).

(100) = 1 + 0.0001(5100 200)

(5000) = 1 + 0.0001(5100 10000)

6 and unstable for r > 1 +

6 3.449, conrming our observation from the graph above. sol-gerda

(0) = < 1, therefore x = 0 is stable. In case 2, f

(N) = 1 + kN 2kN = 1 kN.

(k) = 1 + ln k 1 = ln k > 0, we see that

(t) = g(u), sol-gerda

= 0 to nd the steady states, namely P = 0 and P =

(0) > 0, and P = 0 is a stable steady state.

(0) < 0, and P = 0 is an unstable steady state.

2t, 100 > 2t,

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