CHAPTER 2

Continuous Population Models for Single Species

2.1 Introduction
In this chapter, which treats only a single species, which are not intended to be very realis-
tic, various mathematical assumptions (a number of the difficulties which arise typically in
building mathematical models) on the growth rate are discussed. The birth rate of a human
population is usually given in terms of the number of births per thousand in one year. The
period of one year is also only a convention; the birth rate could just as well be given in terms
of a week, a second or any other unit of time. Similar remarks apply to the death rate and to
the growth rate. The dependence of the growth rate on food supply and the negative effects
of overcrowding.

Before describing a model, we explain some assumptions upon a single species population:

i. the ecological properties (e.g., age, sex, etc) of each individuals are identical

ii. the population growth rate is governed by the birth and death rates

iii. there is no inter specific competition

iv. the population size is large enough to be described by continuous variables

v. there is no time logging to respond to any change in their environment

vi. no migration is allowed.

2.2 Continuous Model of Single Species

Let t denote the time (non-negative real variable) and u(t) denote the population size of a
species at time t and the population changes at time t + ∂t is u(t + ∂t) of a given species
(the number of individuals) in the interval [t, t + ∂t]. The growth rate is thus the net change
in population per unit of time divided by the total population at the beginning of the time
period. Then the average growth rate is

u(t + ∂t) − u(t)

u(t)∂t

We assume that u(t) is continuously differentiable. We obtain the instantaneous per capita
growth rate at time t as
u(t + ∂t) − u(t) u′ (t)
lim =
∂t→0 u(t)∂t u(t)
This is function of t, the growth rate of the population at time t where the rate of change of
u(t) is
du
u′ (t) =
dt
The dynamics of a single species population can be described by

u′ (t) = u(t)f (t, u(t)) (2.2.1)

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Where, u(t) denotes the size of the population at time t. The simplest assumption is that of
constant growth rate which depends only on birth and death processes, that is,

f (t, u(t)) = b−d = intrinsic growth rate of the population

where b = intrinsic birth rate
d = intrinsic death rate.

Example: If the birth rate is 3.2 per hundred and the death rate is 1.8 per hundred,
then the growth rate is 3.2–1.8 = 1.4 per hundred = 1.4/100 = 0.014. Then the equation for
population growth
du
= 0.014u,
dt
where u is the density of population.

2.3 Malthusian Model

The mathematical model describe the growth of a single species population where u′ is pro-
portional to u with proportionality growth rate r as

u′ (t) = ru(t), u(t0 ) = u0 > 0. (2.3.1)

Thus u satisfies the differential equation

du
= ru.
dt
The simplest model we assume that growth rate r is a constant, that is, it does not change
with either time or population. We may solve the equation (2.3.1) by the method of separation
of variables as follows
du
= rdt.
u
Integrate in both sides from t0 to t, we obtain

ln(u/u0 ) = r(t − t0 ).

Thus the solution of mathematical model (2.3.1) is described by

u(t) = u0 er(t−t0 ) . (2.3.2)

The population will eventually either die out (r negative) or grow to fill the universe (r
positive) or constant (r = 0). The model is useful for short term predictions. This is known
as the exponential law of growth or Malthusian law of growth. It was first enunciated
by Thomas Robert Malthus (1766-1834) an English professor of history and political economy
in 1798, who concluded that, “Population, when unchecked, increases in a geometric ratio”
and so this is called Malthusian model.

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Behavior of the solution
Different values of r, we obtain three different solution of Malthusian model.
Case 1: If r < 0, we get,
u(∞) = lim u(t) = 0.
t→∞
This is called the case of extinction (decays exponentially).
Case 2: If r = 0, then we have
u(t) = u0 .
The population remains unchanged (constant).
Case 3: If r > 0, then we obtain
u(∞) = lim u(t) = ∞.
t→∞

The population grows exponentially (unlimited growth).

10

6
u(t)

4 r>0

1
r<0
0

0 1 2 3 4 5 6 7 8
t

Figure 2.3.1: Trajectories of Malthusian model.

Taking t0 = 0, then the solution (2.3.2) reduces to

u(t) = u0 ert .
The populations tend to growth exponentially, so long as enough nutrients are available.
Remark 2.3.1. The growth rate of a population will depend on the size of population. If
the population becomes very large, we can expect the death rate to exceed the birth rate.
We replace the net reproduction rate r by r(u) which is strictly decreasing function of u for
large u and become negative when u is very large.

Malthusian model with reality

From the above observation it is seen that, r is positive and this makes the model unrealistic
for any long time prediction, so most population are constrained by limitation on resource
even in the short run. In reality, a population cannot increase indefinitely. However there is
evidence that population grow exponentially for a finite time period.
Proposition 2.3.1. Suppose the population u(t) is given by exponential growth law where
the growth rate r is positive for some time t0 . Let T be the amount of time required for the
starting population to double. Then
loge 2 0.693
T = ≈ . (T does not depend on u0 ) (2.3.3)
r r

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Proof. We know that
u(t) = u0 er(t−t0 ) .
By definition, u(t0 + T ) = 2u(t0 ). But the equation (2.3.2) implies that

u(t0 + T ) u0 erT loge 2

2= = ⇒T = , assume t0 = 0.
u(t0 ) u0 r

If r = 2%, that is, r = 0.02 then T ∼

= 0.69315
0.02 ≈ 35 years.
Proposition 2.3.2. Assume that u(t) is the size of population obeying an exponential growth
law. Let u1 and u2 the values of u(t) at distinct times t1 < t2 , respectively. Then the growth
rate r is given by the formula
1
r= loge (u2 /u1 ). (2.3.4)
t2 − t1
Proof. We know that
u(t) = u0 er(t−t0 ) ,
and assume that u(t1 ) = u1 , u(t2 ) = u2 .
Then u1 = u0 er(t1 −t0 ) ,
and u2 = u0 er(t2 −t0 ) .
We get,
u2
= er(t2 −t1 ) .
u1
1
Hence r= t2 −t1 loge (u2 /u1 ).

Example 2.3.1. Bacteria in a certain culture increase at a rate proportional to the number
present. If the number u increases from 1000 to 2000 in 1 hour, how many are present at the
end of 1.5 hours?
Solution. We have u(t) = u0 ert , we obtain

1000e1.r = 2000 ⇒ er = 2 ⇒ r = ln2.

Thus u(1.5) = 1000e1.5ln2 = 2828.4271.

Example 2.3.2. A population of a city increases at a rate proportional to the present number.
It has an initial population of 50,000 that increases by 15% in 10 years. What will be the
population in 30 years?
Solution. Let population at time t is u(t) and at t = 0, u0 = 50, 000. Since the rate of increase
is proportional to the number, then du rt
dt = ru, then u(t) = u0 e which yields u(t) = 50, 000e .
rt

Since the population increases by 15% in 10 years then

115
u(10) = 50, 000e10r ⇒ 50, 000 × = 50, 000e10r .
100
Thus e10r = 1.15 ⇒ r = ln(1.15)
10 ,
Thus population will be after 30 years,

u(30) = 50, 000e30r = 76043.74991.

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Example 2.3.3. If the population of a country doubles in 30 years, in how many years will
be triple under the assumption that the rate of increase is proportional to the number of
inhabitants?
Solution. Since the rate of increase is proportional to the number of inhabitants, so we get
du rt
dt = ru, then u(t) = u0 e .
Since the population is double in 30 years then u(30) = u0 e30r = 2u0 .
Thus e30r = 2 ⇒ r = ln302 .
Let the certain time t = T , the population is triple, that is, u(T ) = 3u0 that implies u(T ) =
u0 erT = 3u0 ⇒ erT = 3 which yields (for r = ln302 )

30 ln 3
T = = 47.55 Years.
ln 2
Example 2.3.4. A population of a town increases from 15,000 to 25,000 in 10 years. As-
suming that the population is growing exponentially, how long does it take the population to
increase by 40%.
Solution. We have u(t), since u0 = 15, 000, and u(t) = 25, 000 when t = 10 then 15000e10r =
25000.
Hence e10r = 5/3, so r = (1/10) ln(5/3).
Thus we obtain,
 t/10
1
( 10 ln 53 )t 5
u(t) = 15, 000e = 15, 000 .
3
When the population has increased by 40%, it will be equal to

15, 000 + (0.40)15, 000 = (1.40)15, 000, so u = 1.4(15, 000),

t/10
then 15, 000 53 = 1.4(15, 000).
Thus we obtain
10 ln(1.4)
t= years.
ln(5/3)
Example 2.3.5. A bank account earns interest at an annual rate of 5% compounded con-
tinuously. How long will it take the amount of money in the account to triple?
Solution. We have u(t) = u0 ert , since r = 0.05, then u = u0 e0.05t , such that 3u0 = u0 e0.05t .
ln 3
So we obtain 0.05t = ln 3, thus we obtain t = 0.05 = 20 ln 3 years.

Estimating the U.S. population by exponential law of growth

We need two base years for and we consider 1790 and 1910 and the corresponding U.S popula-
tion are 3.929 million and 91.972 million, respectively. The growth rate r is obtained by using
1
the formula (2.3.4) which was noted as r = t2 −t 1
loge (u2 /u1 ). The growth rate r = 0.0262758.
We obtain

u(t) = 3.929e−47.0337+0.0262758t , regarding formula u(t) = u0 er(t−t0 ) (2.3.5)

For defining the U.S population we plot the U.S population predicated by the equation (2.3.5)
[Derive from the points (t0 , u0 ) = (1790, 3.929) and (t1 , u1 ) = (1910, 91.9272)].

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 600

500

400

u(t)
300

200

100

0
1800 1850 1900 1950 2000
t

Figure 2.3.2: Exponential model of the U.S population.

600

500

400
u(t)

300

200

100

0
1800 1850 1900 1950 2000
t

Figure 2.3.3: U.S actual population with U.S predicated population.

Estimating the Bangladesh population by exponential law of growth

We choose 1901 and 1991 as base year, the corresponding Bangladesh population are 28.92
and 111.45 million, respectively.
1
The growth rate, r = 0.0149894, the noted formula was r = t2 −t1
loge (u2 /u1 ).
Thus we have for Bangladesh population,

u(t) = 28.92e−26.4948+0.0149894t . (2.3.6)

140

120

100
u(t)

80

60

40

20
1900 1920 1940 1960 1980 2000
t

Figure 2.3.4: Exponential growth curve for Bangladesh population.

6
 140

120

100

u(t)
80

60

40

20
1900 1920 1940 1960 1980 2000
t

Figure 2.3.5: Actual population of Bangladesh with estimated population of Bangladesh.

Estimating the Global population by exponential law of growth

Let us examine whether Malthusian model has any relationship at all with reality for the
earth’s human population. Let u(t) denote the human population of the earth at time t. It
was estimated that the earth’s human population was increasing at an average rate of 2% per
year during the period 1971 – 1991. Let us start in the middle of this decade on March 5,
1981 at which time the Bangladesh estimated the earth’s population to be 95.93 million. We
have
t0 = 1981, u0 = 95.93.
So that we have from (2.3.2) as established u(t) = u0 er(t−t0 ) such that

u(t) = 95.93e0.02(t−1981) .

Now, we can check this formula for past population. It reflects with surprisingly accuracy the
population estimated for the period 1971 – 1991. Again, using the Proposition 2.3.1, we get
the doubling time T of earth’s population. The doubling time, T = 34.6574.

2.4 Population Growth with Power Law Model

The generalization of Malthusian model can be described by the power law model. It was
first introduced by Mendelsohn in 1963. This model has the following form:
du
= rub (2.4.1)
dt
where r is the intrinsic growth rate. If b = 1, we get Malthusian model which we have
already discussed. This equation has the solution of the following form:
1
u(t) = [u01−b + r(1 − b)(t − t0 )] b−1 .

Behaviour of the Solution

Now we can draw the graph from this solution using different value of b. For b = 1, we get
the Malthusian model. We consider here two cases for b > 1 and b < 1 and analyze the effect
of the exponent in the model.
In Figure 2.4.1, we can see that the graphs are different for b = 1.05 and b = 0.9, therefore
the model behaves differently.

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 1000 1000

900 900

800 800
a>0 a>0
700 700

600 600

u(t)

u(t)
500 500
a=0 a=0
400 400

300 300

200 a<0 200 a<0

100 100

0 0
0 2 4 6 8 10 0 2 4 6 8 10
t t

Figure 2.4.1: Power law model (2.4.1) with initial volume u0 = 5 × 10−4 and the exponent, b = 1.05
(left) and b = 0.9 (right).

1. For b = 1.05 we can observe that the volume increases faster for r > 0 (r = a), and it
decays at faster rate for r < 0.

2. For b = 0.9, we see that the growth rate of population is slower than the case of the
other graph. For r > 0 the population still increases exponentially but at a slower rate
and for r < 0 the population reduces slowly. So, we can conclude that the growth rate
gets faster with increase in the value b.

2.5 Population Growth with Migration

We can also describe a population that obeys an exponential law of growth with migration
du
= ru + k, (2.5.1)
dt
where the migration rate is k. Assume that, when k is positive we have immigration, and
when k is negative we have emigration.
Let u(t) = u + kr , r ̸= 0.
Thus u′ = u′ and u = u − kr . Observe that u(t0 ) = u0 + kr .
Thus we obtain from (2.5.1), u′ = r u − kr + k = ru, then u(t) = u(t0 )er(t−t0 ) .

The solution of the model is

 
k r(t−t0 )
r(t−t0 )
u(t) = u0 e + e −1 where u(t0 ) = u0 .
r
 
r(t−t0 ) k k
=e u0 + − .
r r
du
Equilibrium point, dt = 0, implies u(t) = − kr .

8
 26 6
x 10 x 10
12 5

4.5
10
4

3.5
8
3

2.5

u(t)

u(t)
6
2

4 r<0 1.5

1 r>0
2 0.5

u0= − (K/r) 0
0
−0.5
0 2 4 6 8 10 0 2 4 6 8 10
t t

Figure 2.5.1: Graph of migration model (2.5.1) with k = 3 × 10−5 and (left) r = −5 × 10−7 , (right)
r = 2.5 × 10−6 .

Limiting behavior
ˆ If r < 0, then u(t) → − kr as t → ∞, a stable fixed point of the system.

ˆ However, if u0 = − kr , r < 0 then the system is stuck at − kr . The value u(t) = − kr , is

an unstable fixed point of the system.
ˆ If r > 0 then u(t) explodes too large and uncountable as t → ∞. It is unstable because
even if we start very near u, the system moves far away from u. So, the equilibrium
points is unstable as t → ∞.

2.6 Population Growth with Gompertz Model

The Gompertz model exhibits an exponential decay of the growth rate. It has been
successfully used to model breast and lung cancer growth. It is a sigmoid function (function
that has S shaped curve) which describes growth as being slowest at start and end. It has
been modified suitably for use in biology, with regard to detailing populations. The model
can be described by the following form: For t0 = 0 the graph of this model (2.6.1) is shown
below:
800

700

600

500
u(t)

400

300 a>0

200

100
a=0
a<0
0
0 20 40 60 80 100
t

Figure 2.6.1: Graph of Gompertz model (2.6.1) with β = 1 × 10−7 and u0 = 10−4 .

du
= re−βt u (2.6.1)
dt
where r is the intrinsic growth parameter and β is the parameter of growth deceleration.
Integrating both sides by taking limits from u0 to u and t0 to t, we get:
r
(e−βt0 −e−βt )
u(t) = u0 e β .

9
 The population increases exponentially over time when the growth rate r > 0, (r = a) as
seen in Figure 2.6.1. Also we can see that the population decreases exponentially over time
if r < 0 and population growth is constant if the growth rate is 0.

Food Supply Model: The growth rate can depend on many things. Let us assume that
it depends only on the per capita food supply σ and that σ ≥ 0 is constant. There will be a
minimum σ0 necessary to sustain the population. For σ > σ0 , the growth rate is positive, for
σ < σ0 it is negative, while for σ = σ0 the growth rate is 0. The simplest model is the growth
rate a linear function of σ − σ0 . Then

du
= a(σ − σ0 )u, a > 0 (2.6.2)
dt
where a and σ0 are constants, dependent only on the species, and σ is a parameter, dependent
on the particular environment. The solution of the model is

u(t) = u0 eta(σ−σ0 ) , u(0) = u0

Thus the population must increase without limit, remain constant or approach 0, depending
on whether σ > σ0 , σ = σ0 or σ < σ0 .

Another Linear Model: Consider the equation

du
= r(a − u). (2.6.3)
dt
Then the solution of the equation (2.6.3) is u(t) = a − ce−rt .
Thus u → a as t → ∞ for a > u0 and r > 0.

a 4.5

4
u(t)

3.5

u0 2.5
0 0.5 1 1.5 2 2.5 3
t

Figure 2.6.2: Graph of the model (2.6.3) with r = .9, u0 = 2 and a = 4.5.

2.7 Dynamics of Tumor Growth Model Using Malthusian Model

If free-living dividing cells, such as bacteria cells grow at a rate proportional to the volume of
dividing cells at that moment. Let u(t) denote the volume of dividing cells at time t. Then

du
= ku (2.7.1)
dt
for some positive constant k. The solution of equation (2.7.2) is

u(t) = u0 ek(t−t0 )

10
where u0 is the volume of dividing cells at time t0 (initial time). Thus, free-living dividing
cells growing exponentially with time, whereas solid tumors do not grow exponentially with
time. As the tumor becomes larger, the doubling time of the total tumor volume continuously
increases.

Remark 2.7.1. To derive more accurate model of population growth, it should not consider
the population as one homogeneous group of individuals. Rather it should also subdivide
the population into males and females. Since the reproduction rate in a population usually
depends on the number of females rather than on the number of males.

Modification of Malthusian Growth Model: We conclude that Malthusian model is

satisfactory as long as the population is not too large. When the population gets large, the
model cannot be realistic. The fact that individual members are competing with each other
for the limited living space, natural resources and food available. Thus we must add a
competition term to the linear differential equation. A suitable choice of a competition term
is −bu2 where b is a constant. We consider therefore, the modified equation

du
= au − bu2 . (2.7.2)
dt
The constant b, in general will be very small compared to a, so that if u is not too large then
the term −bu2 will be negligible compared to au and the population will grow exponentially.
However, when u becomes large, the term −bu2 is no longer negligible and to slow down the
rapid growth rate of the population.

2.8 Problems
1. Describe the population growth model dx/dt = ax and comment on the effects of
immigration and emigration.

2. Formulate a “reasonable model” describing the growth of a single species population,

the growth rate depending on the population size. Discuss the stability of equilibrium
states, if any, in your model.

3. Describe the Malthusian model for the dynamics of a single species population. Com-
ment on its plausibility. Determine how long it takes the population to exactly double
in size under this model.

4. Discuss Malthusian model for single species population growth. Discuss the limiting
behavior of the model as t → ∞. Comment on the appropriateness of the model and
suggest some improvement.

5. The population of a city increases at a rate proportional to the present number. It has
an initial population of 50,000 that increases by 15% in 10 years. What will be the
population in 30 years?

6. The population of a certain community is increase at a rate proportional to the present

number. If the population has doubled in 15 years, how long will it take to triple or
quadruple?

11
7. Describe a model for population growth. Draw graph and show that when does the
population grow most rapidly.

8. Explain a single species population model together with its stability and equilibrium
point.

9. Discuss the single species population growth model with migration. Explain the lim-
iting behavior of the model. Also discuss the points of equilibrium of the model and
investigate their stability.

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