LTPJIT 8 SLNGLE SPECIES POPULATION

MODELS
Structure * . Page No.
8.1 Introduction
Objectives
8.2 Fundamental concepts 31
8.3 Exponential Growth Model
Formulation
Solution and Interpretation
Limitations
8.4 Logistic Growth Model
Formulation
Solution and Interpretation
Limitations
8.5 Extension Of The Logistic Model 40
8.6 Summary 42

8.1 INTRODUCTION
Ecology has attracted attention of scientists and philosophers from the early
ages of human civilisation. Some of the writings of great Greek philosophers
like Hippocrates, Aristotle, etc. dealt wit11 ecological materials although the
term "ecology" was not know to them. The word L'ecology" was first coined
by the german biologist Ernst Haeckel in 1869 to define "the science of the
interrelations between living organisms and their environment". The word
"ecology" owes its origin to the Greek: word "oikos" meaning L1house"or
"place to live". Because of growing environmental awarencss now-a-days,
ecology has become a branch of science that is most relevant to everyday life.
The fact that ecology is essentially a mathematical subject is becoming more
widely accepted. Population biology or mathematical ecology deals with the
increase and fluctuations of populations (e.g. plant popdlation, animal
population, or other organic population). The matllematical study of the
problems in ecology is not of recent origin. In fact, Lotka (1924) and
Volterra (1926) werk early pioneers developing foundation,work in this field.
They established their works on the expression of predator-prey and
competing species relations in terms of differential/integral equations.
In this unit we shdl first define some fundamental concepts used in
ecological studies and then develop mathematical models of some basic
principles in ecology dealing with the growth of single species biological
populations. We shall talk about two species biological population in Unit 9.
Objectives
After reading this unit, you should be able to
express population growth processes in a mathematical framework.
apply your knowledge of differential calculus, integral calculus and differen-
tial equations in building mathematical models of population dynamics.
@ to solve mathematical models or population aynamics. single species

e analyse inathematical relations obtained to understand how the change of

a population can be predicted

8.2 FUNDAMENTAL CONCEPTS

In naturc, an individual living organism of any species does not live in

isolation - the organis!ns live in groups which are called populations. The
term population ill(>:.fiIS n group of iridividuals of any one kind of living
organism. Ecological studies start at the population level.
The basic characteristic of a population is iiidicated by its density. The
density of a population is its size. in relation to some unit of space; it is
generally expressed as the n u m b e r of individupls or biomass per unit
area or volume. For example, 300 trees per acre of land or 2 quiritals of fish
per acre of water surface or 20,000 bacteria per cubic metre of a test tubc,
etc..
Since a population changes over time, we are interested in knowing how it is
changing or more precisely, what is its time-rate of cliange which we call the
growth-rate. The growth- rate of a population is tlie rate of change of its
density or size per unit time; it is determined by the birth-rate and the
death- rate. The birth-ralv .F a population is the ~naxiinumproduction of
new individuals per unit timc .~liderideal conditions (i.e. without any
ecological limiting fnctoi. : reprcduction being limited by physiological
factors only). Death-rase may .tc vxpressed as the nuinber of individuals
dying per unit time.
With these few definitic r L ::T r!ow j hrocecd to develop an exponential growth
population model.

B.3 EXPONENTIAL GROWTH MODEL

The principle of exponential growth for human populaiiior~swas first

propounded. by Thomas R. Malthus (1766-1834) an English clergyman and
political economist in the first edition of his famous book entitled A n Essay
o n t h e Principle of Populatioli published j11 1798. Malthus achieved
notoriety through this work for publishing that huma~ipopulation grows at a
(geometrical) rate that is faster than tlie (arithrnotical) rate of growth of the
supply of commod.itieu necessary for life. He predicted famine and wars as a
consequence.
Let us now discuss the exponential growth model propounded by Malthus.
8.3.1 Formulation of the Model
How does one predict the growth of a popnlation? If we are interested in a
single population, we car1 thinlc of species aa being contained in a
compartment (a Petrie dish, an island, a country etc.) and study the growth
process as one-compartment system. While the population say x(t) is always
an integer, it is usually assumed to be large enough so that very little error is
introduced in assuming that x(t) is a continuous function. In fact, to avoid
this problem, x(t) is often taken to be the population density or to be
biomass rather than the number of indivicluals.
Malthus, while formulating the population growth model ~nadethe following
three assumptions
 Biological i) The po:pulation is su%ciently large
El~vironment
ii) Popula,Lion is homogeneuims el-lal;is, it is evel~lj.spread over the living
spa,r:e.
iii) There are no lirnjtntions to growth, i.e., rw limitations of food, space and
so on. Popu'latio~~changes only by the occu~:r.(?nce of hirths and deaths.
Let us now discuss the model fcrmu1:ited iln.der thcse conditioils.

Let x ( t ) ( >I]) be the size of the popnl;2tion at time t and x(0) -- xo. Suppose
.that thc.popu1d;ion changes only by the occurren.ce,of births and
dea,ths-thereis no immigration or emigratio~~. Let B(t) and l3I.b) tieizote,
rr?spectiv&l;y,the n~xnlbersof hirths and deaths 'that 11;lv.u occ.rnrred by time 1;.
Then the per capita birth rate b a,nd,the death rate 1.nare give11 by

1 dD
In = ----.
x(t) dl;

and the per capita grovrtb rate of tlie p~1puIati011

at a'wy t i ~ n et is given by

OF, --.-1 -dx(t)

-.-- =: b -- ni .= r (constant) say
x(t) dt
dx(t)
or, -- = m(t), where x(C1) = xo
dt

where consl;anl represents the net growth rate.

Eqn. (4) is thc niodel equation for the po~>-nliztisn
growth as given by
Maltl~hns.Let us now solve this equation aid see what it repl*eserll,:g.

The simple ordinary differential Xqrl.(Li) Carl be solved by t l ~ erl~ettlotlof

separation of variables,
We have I

Tntc:g~at;ingthis eqiiation, we obtain

+
In x(t) = C rt, C being a constant
To obtaivl C we use the illitid condition x(0)
In xo = c
- xo, arld get
(6)
Solution to Eqn.(5) then reduces to
X(t)
In- = rt -- ln ert
xo
or, ~ ( t )= xoert. , (7)
, Eqn. (7).gives the poprrlation size at any time t. If the net growth rate
I
r > 0, x(t) grows exponentially without any bound as shown in Fig: 1 For
Fig. 1 r '< O,x(t)-+ 0 as t -+ oo,implying that the populatioll is ultimately driven
t o extinction. Both these outcomes are extreme and are not found to occur
in the nature. In this sense the Maltllus model has severe limitations which
we shall discuss now, but before khat, let us solve this example.
32
Example 1: In a population of birds, the proportionate birth rate ,and the Single Species
death rate are both constant, being 0.45 per year and 0.65 per
year respectively. Formulate a model of the population and discuss its
long-term behavionr.
Solution: Let x(t) denotes the size of the population at any time t > 0. The
per capita birth-rate b = 0.45 and,per capita death-rate m = 0.65. Hence

Integrating Eqn. (8) we get

x(t) == xoe-0.2t
where xo is the initial size of the bird population.
Eqn.(B) gives tlie size of the bird population a;t any time t. Since e-0.2t ---+ 0
as t k- oo,x(t) -) 0 whatever (finite) value is assigned to xo . This shows
that the bird population goes to extinction in the long run.

8.3.3 Limitations
Under ideal conditions when the avhilability of space, food and other
resources do not inhibit growth, Inany biological populations are observed to
grow initially at an approximately exponeni;ialrate After some time, when
the population size becomes considerably lasge, there is lack of food, space
and other resources; also there is pollution due to overcrowding. All these
consequences are collectively called "crowding effects". The crowding
effects force the growth rate to decIine. These considerations make it clcar
Chat the growth rate r cannot. be conutant, but must depend on the size or
density of the populalion. This is where the !imitations of the Malthus
model precisely lie.
The above discussioii suggests that Eqr~(4)should be modified as

where r(x) after certain stage decreases as x increwes.

When r(x) is a decreasirlg function of x, the model is said to describe a
process of "feed back" or LLcompensation". The natural biological
population9 usually exhibit compensatory growth processes.
It is thus clear that Eqn.(4) does not provide a very accurate model for the
population growth when the population itself is very large. Therefore, there
is a need to improve this model. In the next section we shall develop a model -.
called Logistic model which takes care of the lwge population.
And now some exercises for y ~ u .

El) In a population of birds, the proportionate birth rate and death rate
are both constant, being 0.48 per year and 0.65 per year respectively.
Immigration occurs at a conatant rate of 2000 birds and emigration at
a con'ktant rate of 1000 birds per year, Use these assuinptions to
formulate a model of the hopulation. Solve the model and describe the
long-term behaviour of the population in the two cases when the initial
?.\populationis 3000 or 8000.
E2) The population of fish in a reservoir is affected by both fishing and
restocking. The proportionate birth rate is constant at 0.6 per year and
the proportionate death rate is constant at 0.65 per year. The reservoir
 Biological is restocked at a constant rate of 4000 fish per year and fishermen are
Environment allo-ived to catch 3500 fish per year.
Use these assum2tions to derive a model for the population. Solve the
rnodel and describe the loqg~terrnbehaviour of the fish population in
the two cases when the initial population is 5000 or 15000.

We shall now discuss the Ilogistic Model based on a density dependent,

comper~atorygrowth process.

8.4 LOGISTIC MODEL

--.-------

-;f:
A
0
B
1
When a ~opulationis growing in a limited space, the density gractually rises
until eventually the presence of other organisms reduce the fertility and
longevity of the population. This reduce the rate of increase of the
population until eventually the population ceases to grow. The growth curve
defi~iedby such a population follows sigmoid, or S-shaped pattern when
density is plotted against tiale (see Fig. 2). This curve was first suggested to
Fig.2 describe the growth of human populatioiis by PF. Verhulst in 1838. The
sigmoid curve arises due to greater and greater action of detrimental factors
(environplental resistance) as the density of populaltion increases. The
simplest, case that can be conceived is the one in which the detrimental
factors are linearly proportional to the density. Such sirriple or ideal growth
form is call(?d 'Llogistic" and the corresponding growth equation is called
the L L l o g i s tequation".
i~
If you, look at the shape of the curve in Fig.2 you will notice that the curve
consists of three different patterns AB, BC and CD. From A to I3 the curve
gradually rises, from B to.C it is almost; an.exponentia1 increase and from C
to D it gets flattened. This curve is found t,o represent adequately t l ~
popula,tion growth which has steady growth initially until the growth ratc. is
reduced due to various factors like mowding effects, epidemics etc. and
ultimately tending almost to zero. In other words, we can say tliat
ultimately the population gets stabilized/reaches an equilibrium value
without any appreciable increase or decrease. We now take up mathematical
formulation of the logistic model.

8.4.1 Formulation
Assumilig r(x) to be positive and putting r(x) = rr (1 - $) in Eqn.(9) where,
constants rl > 0 and K > 0 we get the Verhulst's f i ~ ~ o '?ogistic
us equation".

Since rf(x)= --% < 0 for all x > 0, the per capita growth rate r(x) declines
as the density x increases. This decrewe in r(x) is broughl; about by
environlnent a1 resistance term $ which is linearly proportional to the
density. [Since r(x) FZ r l for sniall x, r is called the "intrinsic growth rate"
i.e,, growth rate free from environmental constraints.]
Note that Eqn.(lO) is non-linear, first order equation. It is easy to solve it by
the method of separation of variables. Before we do that let us discuss the
qualitative behaviour of the solution by using geometric reasoning.
, ,
The graph of $f against x, where 3
is given by Eqn.(lO) gives the graph of 1I
the logistic mowth function as show11in. Fig.3. The graph i~ a parabola wit11
I . 34

i 1
-
 intercepts at (0,O) and (K, 0) and with vertex at T ,r~ Single Species
"1
I

&/dl ("

Fig.3: T h e logistic growth function

hhen 0< x < K, > 0 so that x increases towards K. When
/x > K, < 0 so that x decreases towards K.
This shows that the populatiun level x(t) always approaches K.
We can infact, express this by writing
lim x(t) = I< provided xo*> 0.
t+ca 9 (I1) A constant solution of
If x = 0 or x = K, then 3= 0 and x(t) does not change. The constant a differential equation is
called an equilibrium
solutions x = 0 and x = I(: axe called equilibrium solutions. solution.
Corresponding to them the points x = 0 and x = K are called equilibrium
points or critical points. You would notice in Unit 9 when we discuss the
stability of the critical points in detail that the constant I< defined by
Eqn.(ll) is an asymptotically stable equilibrium. It is a "saturation
level" or 'Lupperlimit" of the population. It is called the 'carryifig
capacity' of the population- the maximum number of individuals that can
persist under the conditions specified.
In many situations it is sufficient to have the qualitative information about
the solution x(t) of Eqn.(lO). However, if we wish to have a more detailed
description of logistic growth - for example, if we wish to know the
population at some particular time, then we must solve Eqn.(lO). Let; us now
do that.

8.4.2 Solution and Interpretation

Consider Eqn. (lo), viz.,

It can be easily solved by the method. of separation of variables, by writing it

in the form
Kdx
= rldt,
x(K - x)

[:+ &]
We can write Eqn.(l2) in the form
dx = rldt: (13)
If we assume that x < K,then Eqn. (13) on integqation yields
+
I

lnx - ln(K - x) = rlt C, C being a constant (14)

X
or, ln- = rlt +C. (15)
K-x
Biological Using the initial condition x(0) = XQ, we obtain C = In&.
Environment Therefore,
xo
In-
X
K-x
+
= 1nerlt ln------
K -x0
xOerl
= In-
K - xo
X xO erl
o r , = -
K-x K - xo

Therefore, x(t) = K
1 + Cle-rt
' K-xo
where C1 = --- is a constant.
xo
You may observe here that we made the assumption that x < K in order to
derive Eqn.(l9). But this restriction is unnecessary, because you can easily
verify that Eq11. (19) gives the solution of the logistic equation whether x < K
or x 2 K. We are leaving it for you to verify

E3) Verify that solution of Eqn.(lO) for x 2 K is given by Eqn.(l9).

Thus the solution of Eqn.(lO) as given by Eqn.(l!J)represents t h e size of I
t h e population at any time t. It is evident from Eqn.(lS) that x(t) + K I

as t + m. Thus a population that satisfies the logistic equation is not like a ,

naturally growing population; it does not grow without bound: but instead
approaches the finite limiting population K as t + m. But because $ > 0 in
I

this case, we see that population is steadily increasiiig. 1

I
Moreover, differentiating Eqn.(lO) with respect to t, we have II
d 2x - r - dx ---2~ dx
dt2 - [dt K d t l

(ili)
If (K - 2x) > 0, we have K - x > x > 0. Then a dx > 0. SO that the
r a t e of increase increases with time. This shows that there is an
accelerated growth of the population in the range 0 < x < $.
0ntheotherhand,if~<x<~,thcn~-2~<0and~-x>0,sothat~
is a decreasing function of time. Thus there is a retarded growth of the
5
population in range < x < K.
We have shown the two typical solution curves x(t) of the loghtic Eqn.(lO)
in Fig.4.
The graphs1of solution of Eqn.(lO) must have the general shape shown in
Fig.4, regardless of the values of r and K. The horizontal lines are the
equilibrium solutions x(t) = 0 and x(t) = K. If the initial population level i
XQ > K, x(t) monotonically decreases towards K; the upper curve depicts this 1
situation. The lower curve, with its characteristic "sigmoid" or "ogive"
shape, is usuaUy referred to as the "logistic growth curve". 1i
1 I

i '
 Single Species

Fig.4: The solution curves of t h e logistic growth equation

The logistic curve rises at an increasing rate to start with, like an
exponential curve, and then gradually slows down and finally flattens out to
approach the horizontal line x = K as t becomes very large, The time-period
before the population reaches half its equilibrium value (K/2) is a period of
"accelerated growth". Thereafter, the rate of growth diminishes and
gradually becomes zero.
If you compare Fig.1 and Fig.4, you would notice that solutions of the
non-linear Eqn.(lO) are strilrillgly different from those of the linear Eqn.(5),
at least for large values of t. Regardless of the value as K, that is, no matter
how small the non-linear term i11 Eqn.(lO), solution of the equation approach
a finite value of t + rn, whereas solution of Eqn,(5) grow (exponentially)
without bound as t - ; 10-. Thus, even a tiny izan-linear term in the
differential equation las a decisive effect on the sohtinn for large t .
Let us now consider the followirlg examples,
Example 2: the logistic model has beell applied to the nat,ural growth of
the halibut population in certaiu areas of the Pacific Ocean. Let x(t),
measured in kilograms he the total mass, or biomass, of the halibut
population at time t. The parameters in the logistic equation are estimated
to have the values rl = 0.71 / year and K = 80.5 x lowKg. If the initial
biomass is xo = 0.25K, find the biomass two years latter. Also find the time
T for which X(T)= 0.75K.

Solution: We can rewrite Eqn.(l9) in the form

Using the data given in toheproblem we find that

Hence x(2) 2 46.7 x 1 0 6 ~ g .

To find T , we solve Eqn.(23) for t and obtain
Biological Using r l = 0.71, 9 = 0.25 and 2 = 0.75, we find
Environment 1 (0.25)(0.25) 1
7 = --In = ----ln9 E 3.095 years.
0.71 (0.75)(0.75) 0.71
E x a m p l e 3: T h e poplation of fish in a large lake has been stable for some
time. Prior to this situation the population was decreasing from an initially
relatively high level. When the population was 4000, the proportionate birth
rate was 10% and tbe proportionate death rate was 70% . When the
population was 3000, the proportionate birth rate was 30% and the
proportionate death rate was 60%.
A model of the population is based on the follo~l~lng
assumptions:

(i) there is no exploitation and no restocking;

(ii) the proportionate birth rate is a decreasing linear function of the popu-
lation;
(iii) the proportio~~ate
death rate is an increasing linear function of the pop-
ulation.

Show that the rnodel based on these assumptions and the above data
predicts that population falls according to the logistic model; find the
equilibrium population size.
Restocking of the lake now takes place at a rate of 20%)of the population per
year. Find the equilibrium population in this case.
So1ution:Let x(t) denotes the size of the fish population a t any time t > 0.
By the given conditions, the proportionate birth rate is
b(x) = A1 - pix (24)
and the proportionate death rate is
m(x) = A2 p2x+ (25)
where Xi, pi(i =.I, 2) are all positive constants.
Then the net proportionate growth rate is
b(x) - m(x) = A1 - A2 - (PI + PZ)x (26)
= A- px (27)
where X = XI - X2 and p = p1+ pp are constants. Here X may have ally sign
but p is always positive. i
Using the given conditions,

3 6 3
and X - 3000p = - -- -
10 l o - - - 10
Subtracting, -1000p = -&
Therefore,
= 3 10-~
.3
and X = 3000 x p - 10
- = 0.6.

Hence, b(x) - m(x) = X - ux = 0.6 - 3 x 1 0 - ~ x

This implies that
This is the 1ogi.sticgrowth equation with carrying capacity AIJ for the Single Species
population. As we hare seen in the 'logistic grov~thmodel', this carrying
capacity is the stable equilibrium population.
Hence the equilibrium popnla!tion level is "a--_
IJ = 2000.
When restocking of the population is allowed, the governing Eqn. (31) is
rnod.ified to the form

This also represents the logistic law of growth with l;fie new carrying
capacity .= 3,";:-F 2227 approximately.
Hence the new equilibrium level of the fish population afi;er ~restocltingis,
2667.
And now a few exercises for you.

E4) A colony of birds has a stable population. Prior t;o this situation tlie
population iucreasod from an initially low I.evel. When the population
was 10:000 the proportionate birth rate was 50% per year and .the
proportionate death rate was 10% per year. When the population was
20,000 the proportionate birth rate was 30% a i d the prbporticlrlate
death rate was 20%.
A model of the population is based on the following assumptions:
(i)there is no migration and no exp1,oitation (sudl as shooting);
(ii)the proportionate birth rate is a decreasing linear func1,ion of the
population;
(iii)the proportionate death rate is an increasiilg liiiear funcl;ion of pop-
ulation.

Show that a lriodel hasecl on thcse assurnptionu and above ciatn preclicks
that the population grows according to the logistic niodel and iind the
stable population size.
Shooting of the birds is now allowed at a rate of 20% of tihe population
per yeas. Find the new equilibrilinl populat'1011.

E5) For the model

where rl , E and 'Kare constaiit , determine x(t) explicitly. Verify froni

the form of the solution that for x > lc(1 - f ) if E 5 rl, then
x(t) + K ( l - E/rl) as t -+ oo whereas if E > rl, then s(k) 3 O
exponentially as t + oo.

We shall now discuss several Jimitations of the logistic model.

8.4.3 Limitations
(i) The logistic model is not suitable for a population of small size. The
reason is obvious; for small x, rlx. (1- $) NN rlx neglecting the
shcond-order small quantity x2. The logistic equation reduces Lo that of
Malthus for'srnall x.
Biological (ii) It has been observed in both laboratory and natural population that the
Environment growth of mani populations (of microorganisms, plants and anirnals)
exhibit a sigmoid pattern, although such populations do not increase
according t o t h e logistic equation. Almost any equation in which
the negative factors increase in some hsnner with density will yield
sigmoid curves. The S-shaped logistic curve isIan adequate description
for the laboratory growth of paramecium, yeast and other organisms
wiih simple life cycles. Population growth in organisms with more
complex life cycles seldom follows the logistticvery closely.
(iii) The basic assumption in the logistic model that,"the environmental
resistance increases linearly with demlsity" is violated in many growing
populations when tested through direct experirneilts. 'This holds for
populations with siniple life histories, as for example, yeast growing in a
limited space.
(iv) Some populations, fluctuate periodically between, two values. These
fluctuations occcr when certain populations reach a sufficiently high
.density, they become susceptible to epidemics. The epidemic brings the
population down to a lower value where it again begins to increase, u$il
when it is large enough, the epidemic strikes again. E u t any llriild of
fluct~a~tion is ruled out in Ez logistic curve.
In addition to the above, the following limitations pertain to both the
population models considered in this unit.
The models of population growth operato in a closed system, without
input or output. Only self-crowding or other internal factors are
modeled. The real world consists of open systems i11 which the input
and output enviroilmeiits play major roles in the behaviour of the
component considered. This short coming is especially apparent, when it
comes to modelling the growth form of human populations. Clearly, the
technological developmenis, pollution camlsideratiorl and sociological
trends have significant influence on the coefficients r and K.
(2) We have considered the population as made up of one homogeneous
group of individuals. Wc should subdivide it into different age groups,
1
iiii;o males and females since tlie reproduction rate in a population I
usualy depends more on the number of females than ou the number of I
males.

8.5 EXTENSION OF THE LOGISTIC MODEL

- I

In the logistic model just discussed the fuilction r(x) is positive a,nd linear.
We shall now consider a simple extension of this model with an assumption I

of r(x) being a non-linear function of x. Three types pf deiisily dependent

r(x) are depicted in Fig.5.
Fig.5 (a) shows that logistic growth decremes linearly with density i.e.
r(x) = rl (1- t) which corresponds to the model discussed in Sec.8.4.1 (ref.
Eqn.(lC))
Fig.S(b) corresponds to the function which has a maximum a t a n
intermediate point. The function r(x) corresponding to this case is of the
. form
r(x) = a1 + a2x + a3x2 (34)
with a2 > 0 and a3 < 0. This represents a situation in which a population
has a maximal intrinsic growth rate a t intermediate density. This is known
.asthe Allee effect. Single Species
The general character of this density dependent function r(x) can be
summarised by the inequalities r(x) >'O.(x < 71) and r(x) < O(x > rl) (where
q is the density for optimal reproduction) 0 < q < k, where k is the carrying
capacity.
Fig.5 (c) corresponds to r(x) = -1nx. You may observe that this is a
non-lineax curve vhich becomes uegative for x > 1 and not defined at x = 0.
This represents a situation in which there is a, negative logarithmic
dependence of growth rate on population size. This relation is not valid for
very small populations since function is not defined at x = 0. This modei is
known as the Gompertz law, which is used mainly for depicting the growth
of solid tumors. Let us consider a, simple example of an Allee effect.

. .
Fig.5
'
Example 4 Discuss the Allee effect .givcnq'that

where ro and q axc positive constaxlts. Can you relate r(x) corresponding to
this situation with Fig.Fi(b).
Solution: Here r(x) = ro - a ( x ;-,rl)2.Comparing this with Eqn.(34) we
get a1 = ro - c q 2 , a2 = 2aq > 0, as = -a < 0. As in Fig.5 (b) this would be
-
an inverted parabola which intersects the axis at r(x) = a1 = ro aq2 it has
a maximum of ro when x = q and drops below zero when x > xo = q t @.-
--
Thus for densities above xo, the population begins to decline. x xo is a
.-"-
stable equilibiium for the population.
I
You may now try this exercise.

E6) A solid turner usually grows at a declining fate because its interiar has
no access to oxygen and other necessary substancls that the circulation
supplies. This has been modeled empirically by the Gornpetz growth -
law

dN ' dy
- = ryN where - =
dt dt l a y

. y is the effective tumor growth rate, which will decrease exponentially

 Biological by this assumption. Show that equivalent ways of writing this are
Environment
I =.
-dN N = (-a1nN)N
dt

We now end this unit by giving a summary of what we have covered in it.

8.6 SUMMARY

In this unit, we have covered the following:

(1) Mathematical.study of the problems in ecology deals with the increase

and fluctuations of populations. (e.g. plant: populal;ion, animal popula-
tion or other organic population).
(2) Maithus model and Logistic model deal with the growth of a single
species biological populations.
(3) F G a~ population of size x(t)(> 0) at any time t, the Malthus model is
given by the equation

where r > 0 is a constant and is the growth rate of the population.

(4) Malthus model works well only for small populations. For large popula-
tions the growth rate r cannot be constant, but'depends on the size or
density of the population.

I
(5) For large populations the logistic model give11 by the equation

(the constant K > 0 being the saturation level of the population) gives
a type of growth which follows an S;shaped or sigmoid pattern when
density is plotted against time.
(6) In nature, growth of many populations of plants and animals exhibit a
sigmoid pattern though they do not increase according to the logistic
equation.

E l ) The differential equation describing the growth of the population is

x(0) = xo
This can be written as

~ 0 . 2 ~I )= -0%
Integrating, l n l ( 1 0 - + lnlC1I
Therefore, 1000 - 0 . 2 = ~ cJe-Oa2"
 Using the initial condition and finding the value of C, we obtain Single Species
x(t) = 5000 - xoe-0q2t
As t -+ oo, e-0*2t -+ 0 and hence x(t) + 5000 = R (say). Thus the
stable population level is 5000 whatever (finite) value the initial
population level xo may have.
Hence, if the initial population be 3000, it rises upto 5000; if the initid
population be 8000, it ultimately drops to 5000.

E2) Proceeding exactly as above obtain the stable population level as

10,000.
E3) Eqn.(13) can be written as

Using initial condition x(0) k xo, we obtain

x xOerlt
or, - = - -
X- K xo - K

E4) Using the same notations as in Example-2, we have

and X - 20, OOOp = -3 2 -- '

- -- -
Subtracting, 10,000p =
Therefore, p = 3 x lo-'
10 10
-4 1
-- -1
10 10 10
-
10

4
Therefore, X = - 10,000
10
+

Therefore X - px = (0.7 - 3 x 1 0 - ~ x )
IJence the equation governing, the growth of the population is

This represents logistic law of growth with carrying capacity

Hence the equilibrium population level is equal to 23333. When

shooting of the birds is allowed, the Eqn. (26) is modified into the form

.I
This also represents a logistic law of growth with carrying capacity
- -s = 16667. Hence the new equilibrium poplilation level after
- ,,?,0.5
shooting is allowed is 16667.
Biological .
Environment

Integrating and using the boundary condition x(0) = k, we get for

. x>k(l-:)
k (1 - H)
x(t) =. 1.- $e-(n-E)t
rl-

a E6) Hint: Use the fact, a = $-( l n ~ )

UNIT 9 TWO-SPECIES POPULATION
MODELS
Structure Page No.

9.1 Introduction.
Objectives
9.2 Types of Interactions Between Two Species
9.3 Prey-Predator Model
Formulation
Solution and Interpretation
Limitations
9.4 Competing Species
Formulation
Solution and Interprekation
Limitations
9.5 Summary
9.6 Solution/Answers
Appendix

9.1 INTRODUCTION

In Unit 8, we discussed two mathelnatical models on the growth of a sing10

species biological population. In reality, any ecosystenl consists of several
species which are interrelated amongst themselves. It is, therefore, necessary
to study multi-species population models to understand the nature and
diversity of natural e'cosystem. For the sake of simplicity, we shall confine
our discussions in this unit to two species only. We shall develop simple
mathematical models for the growth of two populatioils having interrelations
in the form of pre-predator or competition. But, $0 start with in Sec 9.2, we
have discussed different types of interactions between two different species
living in the same ecosystem. The prey-predator model developed by
Vito-Volterra is discussed in Sec 9.3. Tlle population growth model for two
competing species is discussed in Sec 9.4. For understanding the discussion
in this unit the knowledge of the critical points of a system of differential
equations and their stability is essential. For those who are not familiar with
the stability of the system of equations we &regiving the details in the
appendix. You must go through the appendix carefully while reading this
unit this will provide you with the necessary background.
Objectives
After reading this unit, you should be able to
o identify different types of interactions between the populations of two species.

0 get acquainted with the fundamental mathematical model of a pre-predator

system developed by LothVolterra.
0 identify some of the major limitations of prey-predator model.
0 learn the different features of the basic model for two competing species.