Math 155 Course Notes Exercises - Week 9 Solutions

7.11 Logistic equation. Assume that the size of a population evolves accord-
ing to the logistic equation with intrinsic rate of growth r = 1.5. Assume
that the carrying capacity K = 100.

(a) Find the differential equation that describes the rate of growth of this
population.
(b) Find all equilibria, and, using the graphical approach, discuss the
stability of the equilibria.
(c) Use the analytical approach to determine the stability of the equilibria.
Compare your answers with your results in (b).

Solution.
dN N
(a) dt = 1.5N (1 − 100 )
(b)
dN
dt

0 100 N

Equilibrium solutions: N̂ = 0, N̂ = 100. N̂ = 0 is a locally unstable

equilibrium, N̂ = 100 is a locally stable equilibrium.
N


(c) Let g(N ) = 1.5N 1 − 100 . Then after simplifications

3N
g0 (N ) = 1.5 − ,
100

g0 (0) = 1.5 > 0, g0 (100) = −1.5 < 0.

Answers in (b) and (c) agree.

7.12 Logistic equation. The logistic curve N (t ) is an S-shaped curve that

satisfies  
dN N
= rN 1 − with N (0) = N0
dt K
when N0 < K. Use the differential equation to show that the inflection point
of the logistic curve is at exactly half the saturation value of the curve.
[Hint: Differentiate the right-hand side with respect to t.]
Solution.
 2
To find the inflection point, we need to evaluate ddtN2 and find values of N
where it happens, and prove that there is change in the sign of the second
derivative when passing through this location. At first, differentiate the
right-hand side of the given equation:
        
d N N 1 2N
rN 1 − = rN 0 1 − + rN − N 0 = rN 0 1 − .
dt K K K K
2 0
Therefore, ddtN2 = 0 if 1 − 2N K
K = 0, N = 2 (N 6= 0 at this point). Also, note
dN dN
that dt > 0 if N < K and dt < 0 if N > K (from logistic growth equation).
Then
d2N
 
dN 2N
=r 1−
dt 2 dt K
2 2
and we see that ddtN2 > 0 if N < K2 , ddtN2 < 0 if K2 < N < r. Thus we
confirmed that the inflection point on the logistic curve occurs at N = K2 .
Note that if N (0) > K2 , the curve does not have an inflection point and does
not look like an S shape any more.

7.14 Levins model. Denote by p = p(t ) the fraction of occupied patches in a

metapopulation model, and assume that
dp
= 2p(1 − p) − p for t ≥ 0.
dt
(a) Set g( p) = 2p(1 − p) − p. Graph g( p) for p ∈ [0, 1].
(b) Find all equilibria of the differential equation that are in [0, 1]. Use your
graph in (a) to determine their stability.
(c) Use the analytical approach to analyze the stability of the equilibria that
you found in (b).

Solution.

(a) g( p) = 2p(1 − p) − p = p(2 − 2p − 1) = p(1 − 2p).

g(p)

0 0.5 p

(b) Equilibria: p̂ = 0 – locally unstable and p̂ = 1/2 – locally stable.

(c) g0 ( p) = −4p + 1, g0 (0) = 1 > 0, g0 (1/2) = −1 < 0.
Results in (b) and (c) agree well.
7.15 A metapopulation model with density-dependent extinction. Denote
by p = p(t ) the fraction of occupied patches in a metapopulation model,
and assume that
dp
= cp(1 − p) − p2 for t ≥ 0,
dt

where c > 0. The term p2 describes the density-dependent extinction

of patches; that is, the per-patch extinction rate is p, and a fraction p of
patches are occupied, resulting in an extinction rate of p2 . The coloniza-
tion of vacant patches is the same as in the Levins model.

(a) Set g( p) = cp(1 − p) − p2 and sketch the graph of g( p).

(b) Find all equilibria of the differential equation in [0, 1], and determine
their stability.
(c) Is there a nontrivial equilibrium when c > 0? Contrast your findings with
the corresponding results in the Levins model.

Solution.

(a)
g(p)

0 c p
c+1

(b) g( p) = cp(1 − p) − p2 = p(c − cp − p) = p[c − p(c + 1)].

Then p̂ = 0, p̂ = c+c 1 and from the graph we can see that p̂ = 0 is a
locally unstable and p̂ = c+c 1 is a locally stable equilibrium.
c
(c) Yes, there is a non-trivial equilibrium p̂ = c+1 for c > 0 which agrees
with the Levins model.

7.16 Allee effect. Denote the size of a population at time t by N (t ), and

assume that
 
dN N
= 2N (N − 10) 1 − for t ≥ 0.
dt 100

(a) Find all equilibria of the differential equation.

(b) Use the analytical approach to determine the stability of the equilibria
you found in (a).
 (c) Set  
N
g(N ) = 2N (N − 10) 1 − ,
100
for N ≥ 0, and graph g(N ). Identify the equilibria of the differential
equation on your graph, and use the graph to determine the stability of
the equilibria. Compare your results with your findings in (b).

Solution.

(a) N̂ = 0, N̂ = 10, N̂ = 100.

(b)
 
N
g(N ) = 2N (N − 10) 1 − ,
100
   
0 N N 1
g (N ) = 2(N − 10) 1 − + 2N 1 − − 2N (N − 10) ,
100 100 100
g(0) = −20, thus N̂ = 0 is a locally stable equilibrium,
9
g(10) = 2 · 10 · = 18 > 0, so that N̂ = 10 is a locally unstable equilibrium,
10
1
g(100) = −2 · 100 · 90 · = −180 < 0, thus N̂ = 100 is a locally stable equilibrium.
100

(c)
g(N )

0 10 100 N
This graph confirms our findings in (b) regarding local stability of
steady states.

7.17 Kermack–McKendrick model. We will investigate the classical

Kermack–McKendrick model for the spread of an infectious disease in
a population of fixed size N. If S(t ) denotes the number of susceptibles
at time t, I (t ) the number of infectives at time t, and R(t ) the number of
immune individuals at time t, then
dS
= −bSI,
dt
dI
= bSI − aI,
dt
and R(t ) = N − S(t ) − I (t ).
(a) Divide second differential equation by first one to show that when I > 0,
dI a1
= − 1.
dS bS
Also, show that when R(0) = 0, I (0) = I0 , and S(0) = S0 , then solution
of this new differential equation satisfies
a S(t )
I (t ) = N − S(t ) + ln .
b S0
(b) Since I (t ) gives the number of infectives at time t and dI/dt = bSI − aI,
if S(0) > a/b, then dI/dt > 0 at time t = 0. Also, since limt→∞ I (t ) = 0,
there is a time t > 0 at which I (t ) is maximal. Show that the number of
susceptibles when I (t ) is maximal is given by S = a/b. [Hint: When
I (t ) attains a maximum, the derivative of I (t ) with respect to t, dI/dt,
is equal to 0.]
(c) In (a), you expressed I (t ) as a function of S(t ). Use your result in (b) to
show that the maximal number of infectives is given by
 
a a a/b
Imax = N − + ln .
b b S0
(d) Use your result in (c) to show that Imax is a decreasing function of the
parameter a/b for a/b < S0 (i.e., in the case in which the infection can
spread). Use the latter statement to explain how a and b determine the
severity (as measured by Imax ) of the disease. Does this make sense?

Solution.

(a) The classical Kermack–McKendrick model equations


dS
= −bSI,





 dt
dI

= bSI − aI,

 dt
 dR



 = aI.
dt
Divide second equation by first equation
dI
dt bSI − aI
dS
= .
dt
−bSI
On the left-hand side, multiply and divide by dt
dI
dt dt bSI − aI
dS
= .
dt dt
−bSI
dI dS
Recall differential notation: dI = dt dt and dS = dt dt. Then
dI bSI − aI
= , or
dS −bSI
dI a1
= −1 + .
dS bS
 Multiply both sides of the equation by dS and again use the fact that
dI
dI = dS dS
 
dI a1
dS = −1 + dS,
dS bS
 
a1
dI = −1 + dS.
bS
Integrate
Z  
a1
Z
dI = −1 + dS,
bS
a
I (t ) = −S + ln S + C.
b
Given the initial conditions R(0) = 0, I (0) = 0, S(0) = S0 , we get
a
I0 = −S0 + ln S0 + C.
b
Solving for the constant of integration
a a
C = I0 + S0 − ln S0 = N − ln S0 .
b b
Bringing this constant back into the solution I (t ) leads to
a a
I (t ) = −S + ln S + N − ln S0 .
b b
Finally, the general solution
a S(t )
I (t ) = ln + N − S(t ). (1)
b S0
(b) In order to find the number of susceptibles when I (t ) is maximal, we
need to find when I 0 (t ) = 0:
dI dI a
= bSI − aI, − 0 ⇒ bSI = aI, S= .
dt dt b
a
(c) After we plug in S = b into Eqn. (1), the maximal number of infectives
then becomes a
a a
Imax = ln b + N − . (2)
b S0 b
(d) Let the parameter λ = a/b. Then from Eqn. (2)
λ
Imax (λ ) = λ ln +N −λ.
S0
Differentiate with respect to λ :

0 λ 1 S0 λ
Imax (λ ) = ln +λ · − 1 = ln .
S0 S0 λ S0
0 (λ ) < 0 and I
If Sλ0 < 1 (that is, λ < S0 ), then Imax max (λ ) is a decreasing
function.
Maximum number of infectives is lower for lower values of constant a
and higher values of constant b.