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2 Chapter 2 Solutions
100 ln (2) 69
t= < 72r.
r r
K (t) = k0 ek3 t ,
k1 k0
1 ek3 t ,
A (t) =
k3
k2 k0
1 ek3 t .
C (t) =
k3
(b) The sum of K, A and C is k0 , which represents the total number of atoms
in the system. This number is constant because the number of particles in the
system must be conserved. (This result also implies that both potassium-40
and argon-40 are stable isotopes.)
(f ) A/K 1.008 104 .
2
6 6
C (t) (106 parts/m3 )
4 4
3 3
2 2
1 1
0 2 4 6 8 0 2 4 6 8
t (years) t (years)
(a) Constant flow and concentration. (b) Constant flow and seasonal con-
centration.
10
12
C (t) (106 parts/m3 )
C (t) (106 parts/m3 )
10 8
8
6
4
4
2
2
0 2 4 6 8 0 2 4 6 8
t (years) t (years)
(c) Seasonal flow and constant con- (d) Seasonal flow and concentration.
centration.
3
(b)
dc1 F1 dc2 F2 F2
= c1 , = c1 c2 .
dt V1 dt V2 V2
(c)
F1 t F2 F1 t F2 t
c1 (t) = e V1
, c2 (t) = e V1 + e V2 ,
F2 F1
V2 V2
V1
4
2.10. Cold pills.
(b) Cannot divide by k1 k2
y (t) = k1 x0 tek1 t .
(c) Lowering the clearance coefficient does not change the GI-tract drug con-
centration since the differential equation describing the amount in the tract
is independent of k2 . It does, however, increase the time taken for the drug
to be removed from the blood stream. Accordingly, the peak concentration
in the blood stream is higher.
I
Z
e x(t) = ek1 t I dt = ek1 t + a1
k1 t
k1
I I
Z
e y = I ek2 t e(k2 k1 )t dt = ek2 t
k2 t
e(k2 k1 )t + a2
k2 k2 k1
I I
y(t) = (1 ek2 t ) + (ek2 t ek1 t ).
k2 k2 k1
Note: solution for y that was given in question and text is INCORRECT.
2.12. Antibiotics.
(a) ( )
rate of change n
rate drug
o
of drug in = leaves GI tract
GI tract
( )
rate of change n
rate drug
o n
rate drug
o
of drug = enters blood leaves blood
in blood
( )
rate of change n
rate drug enters
o
of drug in = urinary tract
urinary tract
5
(b) Let x (t) be the amount of the drug in the GI-tract, y (t) be the amount
of drug in the blood stream and z (t) be the amount of drug in the urinary
tract at time t. Then
x (t) = k1 x,
y (t) = k1 x k2 y,
z (t) = k2 y,
with
x(0) = x0 , y(0) = 0, z(0) = 0,
and where k1 , and k2 > 0 represent the rates at which the drug is removed
from or enters different parts of the system.
(c)
x(t) = 0.0001e0.72t ,
y(t) = 0.000126 e.15t e.72t ,
z(t) = 0.000026e.72t 0.000126e.15t + 0.0001.
x (t) = I k1 x, x (0) = 0,
y (t) = k1 x k2 y, y (0) = 0,
z (t) = k2 y, z (0) = 0,
6
2.14. Alcohol consumption.
(a) y (t) = k3 t + y0 , y (0) = y0 .
(b) y (t) = y0 e(k3/M )t , y (0) = y0 .
(c) See Figure 2
y
y0
7
2.16. Formulating DEs for alcohol case study.
Conservation of mass requires
d C 1 Vb
(Vb C1 ) = i F1 ,
dt Vg
d(Vb C2 ) C 1 Vb
= F1 F2 C2 ,
dt Vg
So
i F1 F1 F2
I= , k1 = , k2 = , k4 = .
Vb Vg Vg Vb
Since C1 is the concentration in the GI-tract measured in terms of the volume
in the blood, the mass of alcohol is therefore Vb C1 and since F1 measures flow
rate as volume of fluid in the GI tract we need to multiply by the fraction
(C1 Vb )/Vg as the appropriate concentration in the first equation.
dr s
= n r.
dt a
The solution, satisfying r(t) = r0 , is
r(t) = r0 e(s/an)t .
(b) For s/a n < 0 then r is always decreasing. With r0 > a/b, then r
decreases towards s/nb < a/b. When r(t) reaches a/b it decreases further.
Note that at r = a/b, labour supply and capital stock are in balance. There-
after, the capitallabour ratio decreases with labour becoming redundant,
and this redundancy growing.
8
r Value of sf (r) Sign of dr/dt
(1)
r< re sf (r) > nr +
(1) (2)
re < r < re sf (r) < nr -
(2) (3)
re < r < re sf (r) > nr +
(1)
r > re sf (r) < nr -
9
3 Chapter 3 Solutions
dX
= 0.7X 0.2X 300, X (0) = X0 .
dt
(b) 1 176 000
(c) Xe = 600.
X X X
X = ( ) + ( ) (1 ) X = ( ) X 1 ,
K K K
which is in the form of the standard logistic equation with r = .
and with the parameter values in Figure 3.3.2 the larger equilibrium popu-
lation is r !
10 49
Xe = 1+ 1 = 9.
2 10 1 10
(b) Non-zero equilibrium values do not exist when 1 4h/ (rK) < 0. The
largest value of h for which non-zero equilibrium values exist is hc = rK/4.
10
(c) Factoring the right-hand side of the differential equation gives
" r !# " r !#
r K 4h K 4h
X = X 1 1 X 1+ 1 .
K 2 rK 2 rK
Using this factored form, the behaviour of the function for different initial
conditions can be determined and is shown in Table 2. If x0 is lower than
the critical value, r !
K 4h
xc = 1 1 ,
2 rK
the population becomes extinct.
x0 Sign of X Behaviour of X
X tends
q
K 4h
x0 < 11 rK X < 0
2 to zero
X tends to the
q q
K 4h
1 1 rK < x0 < K2 1 + 1 4h
X > 0
2 rK larger steady state
X tends to the
q
x0 > K2 1 + 1 rK 4h
X < 0
larger steady state
dx
= x (xf x) , x (0) = x0 .
dt
(b) The analytic solution is
x0 xf
x (t) = .
x0 (1 e f t) +
x xf exf t
11
Algebraic simplification leads us to the equation
exf t (xf x0 ) = 0.
We are not interested in the case where xf = x0 because then x = 0 for all
time. However, exf t 0 as t . The plant biomass cannot reach xf in
a finite time.
Let the population at time t be x (t), the per-capita birth rate be and the
per-capita death rate be . Let the immigration rate be i and the emigration
rate be e. Then
dx
= ( ) x + i e.
dt
(b) ( ) ( ) ( ) ( )
rate of change rate rate net rate of
of population = of of + movement to
in country X births deaths country X
( ) ( ) ( ) ( )
rate of change rate rate net rate of
of population = of of + movement to
in country Y births deaths country Y
where the net movement into each country incorporates immigration to and
emigration from the country.
Let x (t) be the population in country X at time t and y (t) be the population
in the neighbouring country Y at time t. Let the per-capita birth rate be
and the per-capita death rate be . Subscripts x and y denote the rates
in countries X and Y , respectively. Let be the constant of proportionality
associated with movement of people between the two countries. Then
dx
= x x x x + (x y) ,
dt
dy
= y y y y (x y) .
dt
12
3.11. Investigating parameter change.
Similar behaviour to the logistic model is exhibited. With small r the pop-
ulation grows exponentially and then levels off as overcrowding becomes sig-
nificant. With higher r values, the steady state is again reached, although
damped oscillations are exhibited. With r > 7 and r > 12 we see 2-cycle and
4-cycle solutions, respectively. With r = 15 chaotic behaviour is observed.
A 2-cycle appears when r 7 and 4-cycles occur from r 12 to r 13.
medium + tank
nutrient mixture mixture
tank
(b) S (t) = (F/V ) cin (F/V ) S (t) where cin is the concentration of nutrient
in the inflowing mixture and V is the volume of the tank.
(c) S (t) = (F/V ) cin (F/V ) S (t) p (S) x (t)
(d) x (t) = p (S) x (t) (F/V ) x (t)
(e)
ma
p (S) = > 0 S R.
(a + S)2
The function is monotonically increasing and its maximum value is given by
limx p (S) = m. Therefore, the function is bounded from above. The
value of p (a) = m/2 is half this maximum, m, which is why it is called the
half-saturation constant.
Xn+2 Xn = 2r + r 2 S 2r + 3r 2 + r 3 S 2 + 2r 2 + 2r 3 S 3 r 3 S 4 = 0.
13
The solutions to the quadratic equation (r 2 S 2 (2r + r 2 )S + 2 + r) = 0 are
(2 + r) r 2 4
S= .
2r
mCemt = Cem(t1)
m = em .
14
4 Chapter 4 Solutions
50
40
y(t)
30
20
10
0 1 2 3 4
t
Figure 3: Analytical solution (black solid line) and solutions computed using
a forward Euler method (black dashed line), Huens method (grey solid line)
and RK4 (grey dashed line) for the system in Question 2. All numerical
solutions use a stepsize of 0.1.
50
40
y(t)
30
20
10
0
1 2 3 4
t
Figure 4: Analytical solution (black solid line) and solutions computed using
a forward Euler method (black dashed line), Huens method (grey solid line)
and RK4 (grey dashed line) for the system in Question 2. All numerical
solutions use a stepsize of 0.5.
15
10
y(x)
4
0
0 2 4 6 8 10
x
Figure 5: The difference between the solutions obtained for the system in
Question 3 by different numerical methods. The black line shows the dif-
ference between the forward Euler method and RK4 solutions and the grey
line shows the difference between the Huens method and RK4 solutions. All
numerical solutions use use a stepsize of 0.1.
10
6
y(x)
0
0 2 4 6 8 10
x
Figure 6: The difference between the analytical solution and the solution
obtained using the forward Euler method (black line), Heuns method (dark
grey line) and RK4 (light grey line) for the system in Question 3. All nu-
merical solutions use a stepsize of 0.1.
16
4.4. Discretisation and round-off errors.
When Digits is set to 20, the error decreases as the stepsize increases. With
Digits set to 10, however, it appears that the error is larger when a stepsize
of 0.005 is used than when as stepsize of 0.01 is used.
17
5 Chapter 5 Solutions
Let S (t) and I (t) be the number of susceptibles and infectives at time t,
respectively, and let be the transmission coefficient. Then
S = SI, I = SI.
18
5.5. SEIR model, disease with latent period.
( ) ( )
rate of rate
change in no. = susceptibles
susceptibles exposed
( ) ( ) ( )
rate of rate rate exposed
change in no. = susceptibles become
exposed exposed infected
( ) ( ) ( )
rate of rate exposed rate
change in no. = become infectives
infectives infected removed
S = SI, E = SI E, I = E I,
where is the transmission coefficient, 1 is the latent period and 1 is
the infectious period.
( ) ( ) ( )
rate of rate of rate prey
change of = prey X X killed
prey X births by predators
( ) ( ) ( )
rate of rate of rate prey
change of = prey Y Y killed
prey Y births by predators
( ) ( ) ( )
rate of rate of rate of
change of = predator predator
predators births deaths
Let X (t) and Y (t) be the two prey species and Z (t) be the predator species.
We assume that the populations are sufficiently large to neglect random dif-
ferences between individuals, that only the two prey species and one predator
species affect the ecosystem, that the prey populations grow exponentially in
the absence of a predator and that the prey do not compete with each other.
Then
X = b1 X c1 XZ Y = b2 Y c2 Y Z, Z = d1 XZ + d2 Y Z a3 Z,
where b1 and b2 are the per-capita birth rates of the prey species, a3 is the
per-capita death rate of the predators and c1 , c2 , d1 and d2 are positive
constants representing the interaction terms.
19
5.8. Predator-prey with density dependence.
(c) The populations oscillate around the equilibrium populations and settle
down to these values over time.
5.11. Symbiosis.
births deaths
species X
births deaths
species Y
Let X (t) and Y (t) be the density of species X and Y , respectively, at time
t. Then
X = c1 XY 1 X, Y = c2 XY 2 Y,
where 1 and 2 are per-capita death rates and c1 and c2 are interaction
coefficients.
J = 1 A J 1 J, A = J 2 A,
where 1 is the per-capita birth rate of juveniles, 1 and 2 are the per-capita
death rates of juveniles and adults, respectively, and is the rate at which
juveniles mature into adults.
20
Expressed in words the equations describing the inputs and outputs are
( ) ( ) ( ) ( )
rate of rate rate rate larvae
change of = larvae larvae age to
larvae born die pupae
( ) ( ) ( ) ( )
rate of rate larvae rate rate pupae
change of = age to pupae age to
pupae pupae die adults
( ) ( ) ( )
rate of rate pupae rate
change of = age to adults .
adults adults die
Y = 1 SY 1 AY, A = 2 Y A, S = 3 Y S,
where 1 S (t) is the per-capita birth rate of yeast, 2 Y (t) is the production
rate of alcohol, 1 A (t) is the per-capita death rate of yeast and 3 Y (t) is
the consumption rate of the sugar.
S = aS SI, I = SI I,
where a is the per-capita birth rate of susceptibles.
21
5.16. Density dependent contact rate.
With = 0.5, the number of infectives is at a maximum after approximately
8 days. As increases, the maximum number of infectives is higher and is
reached after only 7 days. Conversely, with = 0, the maximum number of
infectives is lower and is reached after 9 days.
As decreases from 1 to 0, the spread of infection among the susceptibles
takes longer to occur and the number of susceptibles who never become
infected increases.
R = a1 B R, B = a2 R,
(b)
R = a1 B, B = a2 RB.
(c)
R = a1 B + r1 , B = a2 RB + r2 ,
where r1 and r2 are the rates of reinforcements of the red and blue armies,
respectively.
22
where pr = 1/100.
(c) The differential equations are
B = a1 RB, R = a2 R
d2 R
= a1 a2 R.
dt2
(b) Assume R (t) = Cet . Substitution into the differential equation in (a)
gives
C 2 a1 a2 et = 0
and
B (t) = b0 cosh (t) r0 1 sinh (t) ,
p
where = a1 /a2 .
23
from a single bite may be different for humans than mosquitoes. Hence we
obtain the differential equations
dSh
= 1 Sh Im ,
dt
dSm
= 2 Sm Ih ,
dt
dIh
= 1 Sh Im ,
dt
dIm
= 2 Sm Ih .
dt
converted ordained
unconverted converted missionary
24
5.24. Diseases with carriers.
dS
= (SI + SC),
dt
dI
= (SI + SC) I
dt
dC
= qI
dt
dR
= (1 q)I.
dt
S = SI S, E = SI + S, I = S I.
(c) The revised SEIR model model, including natural deaths, is now
S = SI S aS, E = SI + S aE, I = S I aI
s0
R0 = .
+d
The value of R0 is smaller than for the SIR model because this model accounts
for some people dying of natural causes before they recover for the disease.
25
6 Chapter 6 Solutions
(c) Yes.
26
I
/b S
p1 2 + p2 1 2 + p2 p1
(X, Y ) = (0, 0) , K 1 ,0 , , 1
1 c2 c1 Kc2 c1
27
Y
1 /c1
2 /c2 X
c/b X
Figure 12: A typical trajectory for Question 11. The dashed line is the
non-zero nullcline.
(c) Yes. From Figure 12 it is clear that intersection with the Y axis is possible.
This corresponds to the rabbits completely dying out.
28
The number of microorganisms initially increases but then decreases as more
toxin is produced.
p0
r/ T
Figure 13: Regions in the phase-plane illustrating the trajectory directions
for Question 12. The dashed line is the non-zero nullcline.
29
R
K>0
K<0
1K 1 B
Figure 14: Regions in the phase-plane illustrating the trajectory directions
for Question 14.
B
Figure 15: Regions in the phase-plane illustrating the trajectory directions
for Question 15.
30
6.16. Battle with long range weapons.
(a)
c1 c1
R= B + K, K = r0 b0 .
c2 c2
(b) The numbers of red and blue soldiers can only decrease so the direction
of trajectories is as shown in Figure 16.
31
6.18. Bacteria in gut.
Easiest to use Maple to find the equilibrium points.
This gives
(F/V )Xin
X=
x + x + F/V
and
x y + y F/V + y x Yin (x F/V + (F/V )2 + x F/V )
Y =
F/V (F/V x + x + c2 Xin )
6.19. .
dR
Correction: last equation should be = I aR.
dt
(a) It is easiest to use Maple to find all the equilibrium points. This gives
+ a a(N ( + a))
(S, I) = (N, 0), (S, I) = , .
( + a)
32
7 Chapter 7 Solutions
1
y
4
2 1 0 1 2 3 4
x
7.3. Linearisation.
(a) (x, y) = (1, 1) , (1, 1) .
(b) At an equilibrium point (x, y) = (xe , ye ) we have the linearised system
x 1 1 x xe
= .
y ye xe y ye
1.5
0.5
0
y
0.5
1.5
2
2 1.5 1 0.5 0 0.5 1 1.5 2
x
33
7.4. Linearisation.
(a) At an equilibrium point (x, y) = (xe , ye ) the linearised system is
x 1 1 x xe
= .
y ye xe y ye
(b) (x, y) = (4, 1) is an unstable node and (x, y) = (1, 4) is a saddle point.
See Figure 19 for a typical phase-plane diagram.
2
y
2
4 3 2 1 0 1 2
x
(c)
q q
J= .
y x
The two eigenvalues of J must have different signs at one or more equilibrium
points for a saddle point to exist. The determinant of J, evaluated at the
equilibrium point, is the product of the eigenvalues and so we must show
that the determinant is negative. Now det (J) = q (xe + ye ), where (xe , ye )
is the equilibrium point. We must have, therefore that q > 0 and xe + ye > 0
or q < 0 and xe + ye < 0.
The two equilibrium points are
s s
2 2
3 1 3 3 1 3
(xe , ye ) = + 16, + 16
2q 2 q 2q 2 q
and so s
2
3
xe + ye = + 16.
q
Regardless of the value of q the determinant will be negative at one of the
equilibrium points and so one of the equilibrium points will be a saddle point.
34
(b) The eigenvalues of the Jacobian at each steady state are shown in Table
3. (0, 0) is an unstable node, (0, 1.25) and (1.071, 0) are saddle points and
(1.111, 0.278) is a stable node. The species will coexist. This is verified by
the phase-plane in Figure 20.
Equilibrium point 1 2
(0, 0) 3 2.5
(0, 1.25) 0.5 2.5
(1.071, 0) 3 1.429
(1.111, 0.278) 0.433 2.567
1
y
4
2 1 0 1 2 3 4
x
(c) It can be shown, by examining the eigenvalues of the general system, that
the steady state (0, 2 /d2 ) will be a stable node provided
2 c1
<1+ .
1 d1
This requirement is met if 1 = 2.5 and 2 = 3 with the other parameter
values unchanged. From Table 4, we see that the only stable steady state
is (0, 2 /d2 ) and so X will die out. Note that the steady state representing
coexistence is not feasible.
Equilibrium point 1 2
(0, 0) 2.5 3
(0, 1.5) 3 0.5
(0.893, 0) 2.5 2.107
35
7.6. Comparison of nonlinear and linearised models.
(a) The steady states are
2 1
(0, 0) , ,
c2 c1
and, at an equilibrium point (X, Y ) = (xe , ye ), the linearised system is
X 1 c1 ye c1 xe X xe
= .
Y c2 y e 2 + c2 xe Y ye
Time-dependent plots for the nonlinear and linearised systems are shown in
Figure 21. For initial values close to the non-zero equilibrium solution of
(100, 130), the solution obtained using the linearised system is similar to the
full solution.
145
110
140
105
135
x 100 y 130
125
95
120
90
115
0 5 10 15 20 0 5 10 15 20
t t
(a) (b)
Figure 21: Prey (a) and predator (b) populations over time computed using
the linearised system (grey line) and the full system (black line) for Ques-
tion 6. The initial prey and predator populations are (110, 120) and (90,
140).
(b) A number of trajectories in the phase plane are compared in Figure 22.
36
150
140
y 130
120
110
85 90 95 100 105 110 115
x
Figure 22: Phase portrait for Question 6 using the nonlinear (black) and
linearised (grey) systems.
37
8 Chapter 8 Solutions
= 23 1 13 2 ,
= 3 (13 22 12 23 ) + 32 ,
= 3 (11 23 13 21 ) 31 ,
= 1 (22 31 21 32 ) + 2 (11 32 12 31 ) + 3 (12 21 11 22 ) ,
= 11 23 32 + 13 22 31 12 23 31 13 21 32 .
38
0.22
Stable point
0.20
0.18
0.16
3
0.14
Periodic behaviour
0.12
0.10
0.08
2
4 5 6 7 8 9 10
8.5. Lemmings.
(a) Over the first 4-8 years it appears that a four-year cycle is followed. Af-
ter this, oscillations with a yearly period are observed. The phase plane, in
which the trajectory is a closed loop, is further evidence of the simple oscil-
latory behaviour observed over long time periods.
(b)
a1 a2 a1 b2 a2 c1
(0, 0) , ,0 , ,
c1 b2 b1 b2
(c) At an equilibrium point (n1 , n2 ) = (n1e , n2e ) the linearised system is
n1 a1 b1 n2e 2c1 n1e b1 n1e n1 n1e
=
n2 b2 n2e a2 + b2 n1e n2 n2e
The (0, 0) steady state is a saddle point and ac11 , 0 is a stable node if =
a1 b2 a2 c1 < 0 and a saddle point otherwise. The point (a2 /b2 , / (b1 b2 )) is
a stable node if 4b 2
a2 c2
< 1 and is otherwise a stable spiral.
1
8.6. Prickly-pears.
(a) The growth in plant biomass and growth in the moth population are
logistic, while the reduction in plant biomass is approximately exponential
for small V . The functional response is
V
F (V ) = c1 ,
V +D
39
and the numerical response is
JH
N (V, H) = r2 1 .
V
(b) If the plant density V is large compared with D, this term is approxi-
mately c1 H, corresponding to an exponential decline of herbivores indepen-
dent of the plant density V . Alternatively, if V is small compared with D,
the rate of decline follows a different rate of decay that is dependent on V,
proportional to the product HV .
40
8.9. Bovine tuberculosis and a vaccination program.
(a)
deaths
vaccinated
vaccination
births infection
susceptibles infectives
deaths deaths
41
9 Chapter 9 Solutions
( ) ( ) ( ) ( )
rate of rate of flow rate of flow rate heat
change of = of heat of heat + generated
heat in section in at x out at x + x in section
2h 2h
= , = ua .
ak ak
d2 U q (x)
2
+ = 0.
dx k
(b)
V = (r + r)2 r 2 ,
42
( )
rate heat
produced in = Q0 r (2r + r) t.
shell in t
(c)
d dU Q0 r
r + = 0.
dr dr k
43
10 Chapter 10 Solutions
( ) ( ) ( ) ( )
rate of change rate heat rate heat rate heat
of temperature = produced by lost to lost to
in room 1 heater surroundings room 2
( ) ( ) ( )
rate of change rate heat rate heat
of temperature = lost to gained from
in room 2 surroundings room 1
44
dU1 S10 S12
M1 = q0 + (us U1 ) + (U2 U1 )
dt R10 R12
dU2 S20 S12
M2 = (us U2 ) + (U1 U2 )
dt R20 R12
45
11 Chapter 11 Solutions
k2 (uo ui ) k1 (uo ui )
U1 (x) = ui + (x + d1 ) , U2 (x) = uo + (x d2 )
k1 d2 + k2 d1 k1 d2 + k2 d1
(c) In general, J = (ui uo ) /R and so R = d1 /k1 + d2 /k2 . The total
resistance is the sum of the thermal resistances of each material.
q
U (x) = x (l x) + u0
2k
46
q (x)
q0
x
Figure 25: Internal heat generation function for Question 7.
(c)
q0 ax x a
u2 u1
U (x) = 1 e 1 e + x + u1 .
a2 k
47
and from this we find that
(b)
qr 2
U (r) = + C1 ln (r) + C2
4k
(c) As r 0, ln (r) and so C1 = 0.
(d)
q 2 k
U (r) = u0 + a + r2
4k h
48
12 Chapter 12 Solutions
/(2)x
r
= q
|U| e
k
49
0.0012
0.0011
0.0010
U1
0.0009
0.0008
0.0007
Figure 26: Amplitude of the soil surface temperature versus depth of land
mine for Question 8. The temperature is plotted at times t = 0 (black),
t = /4 (thin black) and t = (grey) radians, with = 1.
50