A 2 2 Matrix Model of Population Growth

Author(s): D. Cooke
Source: The Mathematical Gazette, Vol. 61, No. 416 (Jun., 1977), pp. 120-123
Published by: The Mathematical Association
Stable URL: http://www.jstor.org/stable/3616410
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120

THE MATHEMATICAL GAZETTE

A 2 x 2 matrix model of population growth

D. COOKE

Matrix models are used extensively in demography and pop

The example discussed here shows how a simple 2 x 2 matr
use in the study of bird populations. Teachers may find it
illustration of the application of matrices. The knowledge of

required to appreciate the model and investigate it num

summarised as (1) the expression of two linear equatio

equation, (2) the multiplication of matrices. A deeper unde

behaviour of the model comes from the use of eigenvalues
I shall begin by recalling the familiar simple discrete mod
growth. Suppose that generations are distinct and that eac
duces 2 offspring and then dies or disappears. An example
divide into two at regular intervals, hence A = 2. If the num
in the tth generation is n,, it is related to the number in th
tion by the equation
nt= -nt-,.

It follows that if the number in the initial generation is no

n,
The

An.

(1)

population

in

2
=
1.
The
model
m
producing
offspri
present
in
Genera
tion
(t
1),
and
as

where

the

model

offspring
produce
the
multiplication

human

populat

very
different
mu
were
between
20
a
age
groups
with
d

Model
We

of

bird

consider

spring

in

the

Year

(t

equals
that
of
fem
male
birds
and
th
die
during
the
yea
spring
in
Year
t.

which

hatch

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to

121

A 2 X 2 MATRIX MODEL OF POPULATION GROWTH

spring. Adults also die, and the proportion of adults that survive from
spring to the next isp,. It is a striking fact that this constant survival rat
adults appears to be the case with most of the natural bird populations
have been studied. Expressed in other words, this means that the surviv
rate of adult birds is independent of age; it may be that no bird in the
survives long enough to exhibit the effects of old age. Also, for many spe
the number of offspring appears to be uninfluenced by the age of the mot
If the numbers of juvenile and adult female birds in Year t are no, and
respectively, we have the equations:
no, t f nl, t-l
nl, t =tpo0no. t-l +Pl n1 t-1.

These equations may be written in matrix form,

( to = 0 i) (O :t-i:)

nl, \Po Pi[ n, t-l

or, using an obvious notation,

nt = MInt-I.
It can be seen that

nt = Mn-i_ = MMlnt-2 = M2 nt-2,

and that if the vector no specifies the numbers of birds initially, then

nt

Mtno.

(2)

We have derived an equ

replace scalars and, in p
matrix M instead of th
implications of the mod

matrix

(? 2 5)

\0-3 0-5

Each adult female produces 2 juvenile females in the following year. Therefore, on average, each adult female would produce at least 4 eggs, allowing
for male and female offspring, and probably many more since losses among
fledgelings are likely to be high. The survival rates po, P1 can take values
only in the range 0 to 1. Juvenile birds are not likely to be able to survive as
well as adult birds, and therefore usually Po < P.
If the initial population is of 10 adult females (and 10 adult males), the
numbers of females in successive generations are given in the table below.
The table also includes the total number of female birds, N, = n, t + n1, ,
the ratio of juvenile to adult birds no. tnl, t, and the ratio of the total numbers

in successive generations. (The calculations were done by computer. The

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122

THE MATHEMATICAL GAZETI E

Generation, t 01 2 3 4 5 ... 10 11 12 ... 20

No.ofjuv., no,t 0 20 10 17 14 17 22 24 25 42
No.ofadults,nl,t 10 5 8 7 8 8 12 12 13 22

Total no., N, 10 25 18 24 22 25 34 36 38 64

no,t/nl,t 0 4-00 1-18 2-34 1.66 2-00 1-87 1-88 1-88 1.88

N/INt-i 2-50 0.74 1.31 0.96 1-13 1.06 1-07 1-06 1.06

numbers of birds have been rounded to the nearest integer in

were maintained to many places of decimals in the calculatio

this table reveals that the ratio of juvenile to adult bird

constant ratio 1-88, and that when this is attained the total n
increases by a factor of 1-06 each year.
It would make an interesting project for a pupil to investi
by varying the birth and survival rates. The calculations ma
a desk or hand-held calculator, or by means of a simple com
The results may be illustrated by plots of no. t against nl t, an
quantities in the table against time.
Analytic investigation of the model

The eigenvalues of the matrix M are given by the roots of

teristic equation IM - I1 = 0. The two roots A;, 22, which

show are distinct in our model, have corresponding eige

which may be derived by substituting Ai, 22 respectively for A

Me = Ae. Since 2A ? 22, el and e2 are linearly independent. We
express the initial population no in terms of e1 and e2, i.e.

no = al el + a2e2,

where a1 and a2 are constants. We have

nt = Mt no = Mt (a el + a2 e2) = al e + a2 e2.

Knowledge of 2l and 22 will enable us to predict the behavio

The matrix M has the characteristic equation

-2 fi =0-- 0,
Po Pi -

which is 22 - Ai -po fi- 0. The roots are given

4pofl)}. The restrictions on the parameters are th
0 < Pi < 1. It follows that the roots are real and dist
eigenvalue A2 is positive, that 22 is negative and th

write

n,= {ai el + (2/2)t a2 e2},

from which it will be seen that, for large t, nt ~ 2 1 a1 el. Thus in the long

term the age distribution stabilises and is proportional to e,; each age

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123

A 2 X 2 MATRIX MODEL OF POPULATION GROWTH

group then changes by a factor of A each year. When stability is reache

number in each age group, and the total number, obey an equation of
same form as (1). Before stability is attained the numbers oscillate; the m

nitude of the oscillation depends on the magnitude of Ii/AiJ. Fo

numerical example above, Ai = 1-06 and 2 = -0-56.

It is of great interest to know whether a population will ultimately

crease or decrease. It will increase if;Al > 1, and the condition for this is

fi > (1 -pi)lpo; for the numerical example we requiref, > 1-67. The s
age distribution is proportional to el which, in turn, is proportion

(fi )JT

Limitations of the model

It is always important to appreciate the limitations of any mathemat

model. The three major limitations of this model are:

(i) Theparameters change. Birth and death rates change from year to ye

a natural population, and are particularly dependent on the weather

model assumes a constant environment.

(ii) The parameters may be functions of nt. Ecologists believe that in many
populations the birth and death rates vary with the size of the population,
and in this way a population regulates itself. The necessity for this is most
obvious when it is seen that a population cannot increase indefinitely.
(iii) A stochastic model may be more realistic. It would be more realistic to
specify probabilities of survival and of offspring being produced, rather
than assuming constant proportions. The discrepancies introduced by using
proportions are likely to be largest with small populations. The numerical
example above is likely to be more realistic if we took the values to be in
units of 100 birds, so that the initial population was 2000 and not 20.
D. COOKE

School of Mathematical and Physical Sciences, University of Sussex, Fa

Brighton BNI 9QH

Encore!

"In all, 789 performances were given by Covent Garden companies, said Mr John Tooley,

general administrator, 'infinitely more than those promoted by any other European
opera house.' This was at a cost to the British public of about 001p per performance."
From the Daily Telegraph for 22 October 1976 (per John Hersee).
At least they removed the net

"On the inaugural flight to Morocco in 1919, they found out too late that the man who
had agreed to fix up an airfield at Alicante had confused metres with square metres, producing a landing strip the size and shape of a tennis court." From the Daily Telegraph for
19 July 1976 (per A. K. Austin).

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