Mathematical Modelling

CHAPTER 03
FORMULATING INTERACTING POPULATION MODEL
Introduction
Interacting population models are relevant where two or more populations depend on each
other. We study, in detail, four examples of interacting population: (1) model of influenza
outbreak, (2) A model of a battle between two opposing groups, (3) a predator-prey
interaction, (4) a competing species interaction. (3) a competing species interaction.

3.1 Interacting population

Large number of one species may be unaffected by others, however, in some instances,
removing, introducing or modifying one resource or species through (for example) harvesting
or poisoning may have wide-ranging ramifications for the system. Evidence suggests that,
typically, communities with many interacting species have grater stability than those
comprising much simpler systems. That is, while rainforests are stable, cultivated land and
orchards are relatively unstable and the populations of species in laboratory controlled
predatory-prey system undergo large 0scillations.
The relationship between the species within a system are often highly nonlinear, such that it
is extremely difficult to establish, with certainly, a precise mathematical model describing the
processes involved. However, there is clearly a need to understand these systems, or aspects
of them. To this end we develop simple mathematical models where the independent variable
is time and the dependent variables are the numbers or densities (numbers per unit area) of
the various different interacting populations.
A different type of interaction, which occurs within a single species, is the interaction
between those who are infected with a disease and those who are not. We study, in detail, a
simple model for growth and then decline of the number of individuals in a population
infected with a disease.
The most examples come from nature, where different species interact with each other.
Example of population that interact include spread of a disease between population groups,
predator-prey interactions, battles between opposing groups and interactions between
different age or gender groups within a population.
There are many different types of interactions between populations in different trophic levels.
One such interaction, where some species use other species as their food supply, is the
predator-prey interaction. Another interaction process between populations on the same
trophic level is that of competing species, where two or more populations compete for a
limited resource, such as food or territory. We model this system in a manner very similar to
that developed for the predator-prey system.
Another example we study is that of a battle between two opposing groups, such as may
occur between two insect or human populations. The rate at which soldiers are wounded, or
killed, in a battle depends largely on the number of enemy soldiers.

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Case Study:
Nile Perch catastrophe We cite here a case where the ramifications of a disturbance within an
interacting population system were wide reaching and caused an ecological disaster for the
communities living in and around Lake Victoria. It emphasizes the responsibility which should
accompany any unsuitable modification of environmental processes, and the need for good
modelling and forecasting, incorporating a multidisciplinary approach.

Adapted from Murray (1990) and Quammen (1997)

Lake Victoria, which feeds into the Nile River, is the largest and most northern of a series of lakes
punctuating the Rift Valley in Africa. Round as a pond and nearly as big as Ireland it is bordered by
Kenya. Tanzania and Uganda, and until 1960 it supported many communities along its shores
providing them with a large and diverse population of fish as well as fresh water. In particular, it
supported an abundance of cichlids (tropical fresh water fish of the family Cichlidae) in both quantity
and species. The sort of fish known for their garish appearance and collected for aquaria.

Because of its scalloped shoreline, its irregular patterns of depth and shallows, and its subjection to
periods of intense drought and low water levels during the centuries long history of the lake, pockets
of cichlids were cut off from others for long periods. In this way they diverged and speciated. Later
reconnection allowed the species to mix spatially, but they had become incompatible repro ductively
and were now in competition. In order to survive they radiated: that is, they settled in different
niches within the environment. Thus the lake rep resented a kind of underwater Galapagos
archipelago with species for every niche: rock-scrapers, sand-digginginsect eaters, plant-scrapers,
scale-eaters, nibbler-of-other-fishes-fins, fish-eaters and the famed Haplochromis compres siceps
known for biting out eyeballs! Lake Victoria supported about 200 such cichlid species, all of which
are thought to have descended from a single an cestral species.

The ramifications spread far wider than the local ecological and economic disaster. The Nile Perch
are oily and cannot be sun dried, the traditional means by which fish had been preserved. Instead
they had to be smoked which resulted in major felling operations in the local environs to provide the
fuel.

All in all, the perch introduction was catastrophic. Many of the ramifications could have, and should
have, been foreseen and avoided. However, in spite of this, further introductions were being
planned in the late 1980s for Nile Perch into other African lakes such as Lake Malawi.

Modelling assumptions and approach

Different populations are expected to have different birth and death rates. For some
populations the size of another population will affect these parameters, and the populations
are then said to interact with each other.
Models can describe the number of individuals (population size), as will be the case for the
epidemic and battle models we develop. Alternatively, the population density, number per
unit area, can be modelled as in the predator-prey and competing species models that follow.
For both, approach is the same.

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System of differential equations

Many processes are described by more than one differential equation. When these equations
need to be satisfied simultaneously the set of equation is known as a system of differential
equations. The system of equation is known as a dynamical system if it allows prediction of
future states given present and/or past states. We also develop system in which the equations
are nonlinear that is, they include products of the dependent variables or their derivatives.
The system of equation may be coupled, which implies that their solutions are
interdependent. For example, in the case of two equations x ' ( t )=F ( x , y , t )∧ y ' ( t )=G ( x , y , t ) ,
if the solution of x ' depends on the values of y and the solution of y ' depends on x , then the
equations are coupled.

3.2 Model for an influenza outbreak

We develop a model to describe the spread of a disease in a population overview and we use
it to describe the spread of in a boarding school.
To do so the population is divided into three groups:
i. Those susceptible to catching the disease.
ii. Those infected with the disease and capable of spreading it and
iii. Those who are immune from the disease.
Modelling these interacting groups leads to a system of two coupled differential equations.

Background
Over the centuries, there have been dramatic examples of how epidemics example of various
diseases has had a significant effect on the human population.one of the most well- known is
the Black Death in Europe in the fourteenth century. Today epidemic are still prevalent, the
most notable being possibly AIDS and the Ebola virus. If we can understand the nature of
how a disease spread through a population, then we are better equipped to contain it through
vaccination or quarantine. Or in the case of the biological control of pets we may wish to
determine how to increase the spread of the disease (for example, myxomatosis or calicivirus
in rabbits) so as to find an efficient way of reducing the population. unfortunately, humans
themselves have been subjected to this means of control. In colonial times, the spread of
European disease, such as measles and smallpox, had a disastrous impact on certain
indigenous populations who had no resistance to them.
Many diseases are spread by infected individuals in the population coming into close contact
with susceptible individuals. These include influenza, measles, chickenpox, glandular fever
and AIDS. On the other hand, malaria is transmitted through a host, a mosquito, which
carries the disease from individual to individual. Certain diseases are more contagious than
others. Measles and influenza are highly contagious, whereas glandular fever is much less so.
Many diseases, such as mumps and measles confer a lifelong immunity, however, influenza
and typhoid have short periods of immunity and can be contracted more than once.

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There are some naturals definitions that we require in order to proceed with our modelling.
The incubation period of a disease is the time between infection and the appearance of visible
symptoms. This should not be confused with the latent period, which is the period of time
between infection and the ability to infect someone else with the disease. The latent period is
shorter than the incubation period so that an individual can be spreading the disease, yet be
unaware of having it. For measles, the incubation period is approximately 2 weeks and the
latent period is approximately 1 week. We consider simple mathematical model for a flu
(influenza) epidemic at a boarding school over a period of about 15 days. For this period it is
reasonable to assume that reinfection does not occure.

Model assumption
When considering a disease, the population can be divided into distinct classes: susceptible
S(t ) and contagious infectives C ( t ), where t denotes time. The susceptible are those liable to
catch the disease, while the contagious infectives are those infected with the disease who are
capable of communicating it to a susceptible. There are also those who have recovered from
the disease and are no longer susceptible, who are now immune from further infection of the
disease.
Initially, we make some assumptions and then build the model based on them.
 We assume the populations of susceptible and contagious infective are large so that
random differences between individuals can be neglected.
 We ignore births and deaths in this model and assume the disease is spread by contact.
 We neglect the latent period for the disease, setting it equal to zero.
 We assume all those who recover from the disease are the immune (at least within the
time period considered).
 We also assume that, at any time, the population is homogeneously mixed, i.e., we
assume that the contagious infectives and susceptible are always randomly distributed
over the area in which the population lives.
Example 1: Develop appropriate word equations for the rates of change of susceptible and
contagious infectives.
Solution: The only way the number of susceptible can change is the loss of those who
become infected, as there are no births and none of those who become contagious infectives
can become susceptible again. The number of infectives changes due to the susceptible
becoming infected and decreases due to those infectives who die, become immune or are
quarantined. The latter cannot become susceptible again (from the assumption made).
The appropriate word equations are

{ } { }
rate of rate
change∈no . =− susceptible
susceptibles infected

{ }{ }{ }
rate of rate rate
change∈no . = susceptible − infectives (1)
infectives infected removed

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{ }{ }
rate of rate
change∈no . = infectives
recovered have recovered

Formulating differential equations

The number of infectives removed in the time interval should not depends in any way upon
the number of susceptible, but only on the number infectives.
We assume that the rate at which infectives recover is directly proportional to the number of
infectives.
We write

{ }
rate
infective =γI (t )
recovered
(2)
Where γ is a positive constant of proportionality, called recovery rate.
To model the total rate of susceptibility infected, first consider the susceptibility by a single
infective. It is evident that the grater the number of susceptibles, the grater increases in the
number of infectives. Thus the rate of susceptibles infected by a single infective will be an
increasing function of the number of susceptibles.
Let us assume that this rate is directly proportional to the number of susceptibles.
We denote the number of susceptibles at time t by S(t ), and the number of infectives at timet
by I ( t ) , then the rate susceptibles are infected is λ (t)S ( t ).
The total rate that susceptibles are infected is

{ }
rate
susceptibles =λ (t) S ( t ) (3)
infected

The constant b is called the transmission coefficient or infection rate.

Putting together the assumption for rate of new infections and rate of recoveries gives the
system of differential equations,
dS dI dR
=− λ ( t ) S , =λ ( t ) S−γI , =γI (4 )
dt dt dt

Exercise:
(1)Continuous vaccination: Consider a model for the spread of a disease where life long
immunity is attained after catching the disease. The susceptibles are continuously vaccinated
against the disease at a rate proportional to their number. Write down suitable word equations
to describe the process, and hence obtain a pair of differential equations.
Solution: Suitable word equations are

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{ } { }
rate rate
of change ∈no . =− susceptibles
of susceptibles vaccinated

{ }{ }{ }
rate rate rate
of change ∈no . = susceptible − vaccinated (1)
vaccinated vaccinated removed

Let us assume that the rate of susceptibles vaccinated by a single infectives is directly
proportional to the number of susceptibles with a proportionality constant 𝑎.
At time 𝑡 the number of susceptibles is denoted by 𝑆(𝑡) and the number of infectives at time
𝑡 denoted by 𝑉(𝑡).

{ }
rate
susceptibles =¿ 𝑎𝑆(𝑡)𝑉(𝑡)
vaccinated
(2)
The constant 𝑎 is called vaccination rate.
We assume that the number of vaccinated removed is directly proportional to the number of
vaccinated.

{ }
rate
vaccinated =¿ 𝑟𝑉(𝑡)
removed
(3)
The constant 𝑟 is called removal rate.
dS
The rate of change in the number of susceptible is in interval and the rate of change in
dt
dV
the number of vaccinated is given by .
dt
By using (2) and (3), the suitable word equation (1) becomes,
dS dV
=−aSV , =aSV −rV (4 )
dt dt
Equation (4) is required paired differential equation.
2)Simple age-based model: Consider a population split into two groups: adults 𝐴(𝑡) and
juveniles 𝐽(𝑡). Starting with word equations formulate a pair of differential equations
describing the density of adults and the density of juveniles. Define all variables and
parameters used. You may assume constant per-capita birth and death rates for the
population, and also assume that the young mature into adults at a constant per-capita rate γ.

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Solution: suitable word equation is

{ }{ }{ }{ }
rate rate of rate at which rate of
of change of = juveniles − juveniles reach − juveniles
juveniles births maturity deaths

{ }{ }{ }
rate rat rate
of change of = of Juvenile − at which Juveniles (1)
Juveniles births reach maturity

Let us assume that rate of change of Juveniles is denoted by 𝐽(𝑡).

Let β 1 is the per-capita birth rate of Juvenile 𝑎𝑛𝑑 α 1and α 2 be the per-capita death rates of
Juveniles and adults.
Given that γ is the rate at which Juvenile mature into adults.

{ }
rate
of juvenile =β 1 A (2)
birth

{ }
rate
at which Juveniles =γJ (3)
reach maturity

{ }
rate
of Juveniles =α 1 J (4)
deathes

{ }
rate
of adults =α 2 A
deaths
(5)
By using equations (2), (3), (4) & (5)
The appropriate word equation (1) becomes,

dJ dA
=βA−γJ−α 1 J , =γJ−α 2 A (6)
dt dt

Equation (6) is paired differential equation.

3) SEIR model: Many diseases have a latent period, which is when there is a period of time
between infection and when an infected individual becomes infectious. One example is
measles, where the latent period is approximately 5 days. Extend the SIR epidemic model to

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one with an additional population class 𝐸(𝑡), corresponding to individuals who have been
exposed to the disease, so they are no longer susceptibles, but are not yet infectious. You may
assume a latent period σ −1. Also, infectives recover in a mean time γ −1 and have lifelong
immunity.
Solution: Suitable word equation

{ } { }
rate rate
of change of =− of susceptible
no . of susceptibles infectious

{ }{ }{ }
rate rate rate
of change ∈no . = susceptible − infectious
of infectious infectious removed
(1)

Let us assume that rate of susceptible at time t be denoted by 𝑆(𝑡) & rate of susceptible
infectious is denoted by 𝐼(𝑡).

{ }
rate
of susceptible =βS ( t ) I ( t )
infected
(2)

{ }
rate
of infectious =rI (t)
removed
(3)
Using equation (2) & (3)
dS dI
=− βSI , =βSI −rI (4 )
dt dt

With an additional class of population 𝐸(𝑡)

{ }{ { }
rate rate
of susceptible
infectious
= rate
of additional population∧¿ latent period
−
}
of infectious∧recovery
meantime

E(t ) I (t)
I ( t )= − −1
σ −1 γ

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{ }
rate 1
of α
{latent period } { recovery meantime }
recovery

dR 1
= −1 −1 =σγ
dt σ γ

3.3 Simple model of a battle

We now consider a novel type of population interaction: a destructive competition or battle
between two opposing groups. These may be battles between two hostile insect groups,
athletic teams or human armies. While the models we derive here apply to the last case, the
principles can be generalised and would apply to many other examples. The model we
develop turns out to be a system of two coupled, linear differential equations.
Background:
Battles between armies have been fought since antiquity. In ancient times battles were
primarily hand -to-hand combat. With the development of archery and then gunpowder, a
crucial feature of battles has been aimed fire. Although many factors can affect the outcome
of a battle, experience has shown that numerical superiority and superior military training are
critical. The model we present was first developed in the 1920s by F.W. Lanchester who was
also well known for his contribution to the theory of flight.
Our aim is to develop a simple model that predicts the number of soldiers in each army at
any given time, provided we know the initial number of soldiers in each army.
Model assumptions
First we make some basic assumptions and then develop the model based on these.
 We assume the number of soldiers to be sufficiently large so that we can neglect random
differences between them.
 We also assume that there are no reinforcements and no operational losses (i.e. due to
desertion or disease).
These are assumptions that can easily be relaxed at a later stage if the model is inadequate.
Example 1: Determine the word equations for the number of soldiers in both the red and blue
armies.

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Solution: Since there are no reinforcements or operational losses, each population changes
by the number of soldiers who are wounded by the other army.
Thus, in any given instant of time,

{ } { }
rate rate red soldiers
of change of =− wounded by
red soldiers blue army

{ } { }
rate of rate blue soldiers
change of =− wounded by (1)
blue soldiers red army

In a real battle there will be a mixture of shots; those fired directly at an enemy soldier and
those fired into an area known to be occupied by an enemy, but where the enemy cannot be
seen. Some battles may be dominated by one or the other firing method. We consider these
two idealisations of shots fired as aimed fire and random fire. For the model we assume only
aimed fire for both armies.
In the aimed fire idealisation, we assume all targets are visible to those firing at them. If the
blue army uses aimed fire on the red army, then each time a blue soldier fires he/she takes
aim at an individual red soldier. The rate of wounding of the red army depends only on the
number of blue soldiers firing at them and not on the number of red soldiers. We see later that
this assumption is equivalent to assuming a constant probability of success (on average) for
each bullet fired.
For random fire a soldier firing a gun cannot see his/her target, but fires randomly into an
area where enemy soldiers are known to be. The more enemy soldiers in that given area then
the Formulating interaction population model Department of mathematics, GUK Page 10
greater the rate of wounding. For random fire we thus assume the rate of enemy soldiers
wounded is proportional to both the number firing and the number being fired at.
In summary then we make the following further assumptions:
 For aimed fire the rate of soldiers wounded is proportional to the number of enemy soldiers
only.
 For random fire the rate at which soldiers are wounded is proportional to both numbers of
soldiers.
Formulating the differential equations
Let 𝑅(𝑡) denote the number of soldiers of the red army and 𝐵(𝑡) the number of soldiers of the
blue army. We assume aimed fire for both armies. This information is expressed
mathematically by writing

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{ } { }
rate red soldiers rate blue soldiers
wounded by =a1 B ( t ) , wounded by =a2 R ( t ) (2)
blue army red army

Where a 1and a 2 are positive constants of proportionality. The constants a 1and a 2 measure the
effectiveness of the blue army and red army respectively, and are called attrition coefficients.
We thus assume that attrition rates are dependent only on the firing rates and are a measure
of the success of each firing.
We now substitute (2) into the basic word equation (1), where the rate of change in the
number of red soldiers is 𝑑𝑅/𝑑𝑡 and for the blue soldiers it is 𝑑𝐵/𝑑𝑡.
The two simultaneous differential equations are thus,
dR dB
=−a1 B , =−a2 R(3)
dt dt

Exercise:
(1)Battle loss due to disease: Develop a model (a pair of differential equations) for a battle
between two armies where both groups use aimed fire. Assume that the red army has a
significant loss due to disease, where the associated death rate (from disease) is proportional
to the number of soldiers in that army.
Solution: Let 𝑅(𝑡) 𝑎𝑛𝑑 𝐵(𝑡) denotes the number of red soldiers and blue soldiers
respectively at any time 𝑡.
Assume that the number of soldiers to the sufficiently large do that we are neglecting random
differences between them.
Let the per-capita death rate of red army due to disease is α and the rate of death of red army
due to disease is 𝛼𝑅(𝑡)
Word equations are

{ } { }{ }
rate rate of change of rate of deaths
of change of =− red soldiers wounded by − of red soldiersdue
red soldiers blue soldiers ¿ disease

{ } { }
rate of rate of blue
change of =− soldiers wounded
blue soldiers by red soldiers

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Since both the armies use aimed fire is the difference equation describing this model is

dR dB
=−a1 B ( t ) −αR (t ) , =−a2 R(t)
dt dt

Where a 1∧a2 are positive constants of proportionality and a 1∧a2 are known as coefficients.
(2)Jungle warfare. We developed a simple model for a battle between two armies. We
assumed that the probability of a single bullet hitting its target is constant. This is not a good
assumption in jungle warfare where one, or both, of the armies may be hidden from view.
Suppose that soldiers from the red army are visible to the blue army, but soldiers from the
blue army are hidden. Thus, all the red army can do is fire randomly into an area and hope
they hit something. The blue army uses aimed fire.

(a) Write down appropriate word equations describing the rate of change of the number
of soldiers in each of the armies.

(b) By making appropriate assumptions, obtain two coupled differential equations

describing this system.

(c) Extend the model to include reinforcements if both of the armies receive
reinforcements at constant rates.
Solution: Suppose that the soldiers from the red army are visible but the Soldiers from the
blue army are hidden.
Thus, all the army of red will do random firing and blue army will do aim firing.
For random fire a Soldier firing a gun cannot see his target but fires randomly into an area
where enemy soldiers are known to be
For random fire we assume the rate of enemy soldiers wounded is proportional to both are
number of firing and the number being fired
For aimed fire, we assume the rate of enemy soldiers wounded is proportional to the number
of soldiers firing at them.
(a)Word equations are

{ } { }
rate of rate of red
change of =− soldiers wounded
red soldiers by blue soldiers

{ } { }
rate of rate of blue
change of =− soldiers wounded by
blue soldiers red sodiers

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We make some assumptions:

Assume that number of soldiers to be sufficiently large do that we can neglect random
difference between them.
(b) Assume that there are no reinforcements and operational issues
Let R(t) and B(t) denote the number of soldiers of red and blue army respectively then the
differential equation describing the systems are

dR dB
=−a1 ( t ) , =−a2 R ( t ) B (t)
dt dt

Where a 1∧a2 are positive constants of proportionality. These are called coefficients.
(c) If both the armies receive reinforcements at constant rates with this differential equation
are,
dR dB
=−a1 B ( t ) +r 1 , =−a 2 R ( t ) B ( t ) +r 2
dt dt
Where r 1∧r 2 are its constants rates of reinforcements of the red and blue armies respectively.

3.4 Predators and prey

In this section, we develop a predator-prey model for carnivores using the growth of a
population of small insect pests which interact with another population of beetle predators.

Background
There are several types of predator-prey interaction: that of several herbivores which eat plant
species, that of carnivores which eat animal species, that of parasites which lives on or in
another species(the host), and that of cannibals which eat their own species, often an
interaction between the adults and youngs. One interesting example of a predator-prey
interaction occurred in the late nineteenth century, when the American citrus industry was
almost destroyed by the accidental introduction from Australia of the cottony scale insect.To
combat this pest its natural predator, the Australian ladybird beetle, was also imported, but
this did not solve the problem and finally DDT was used to kill both predator and prey in a
bid to eradicate the pest.
Model assumption
We make a few preliminary assumptions on which to build the model.
 Initially we assume the populations are large, sufficiently large to neglect random
differences.

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 We ignore the effect of DDT initially, but modify the model later to incorporate its
impact on the system.
 We also assume there are only two populations the predator and prey which affect
the ecosystem.
 We assume that the prey population grows exponentially in the absence of a
predators.
Example
Develop appropriate word equation for each of two populations, the predator and the prey.
Solution: The only inputs for each populations are births and the only outputs are deaths.
However the prey deaths occure due to the predators capturing and eating them.
The appropriate word equations are

{ }{ }{ }
rate of rate of rate of
change of = prey − prey killed
prey births by predators

{ }{ }{ }
rate of rate of rate of
change of = predator − predator (1)
predators births death

Formulating the differential equations

Let us assume that the per-capita birth rate for the prey is a constant β . The per-capita birth
rates give the rate of births from an individual prey. The rate of births from each individual is
not depends on the predator density. Alternatively the death of the prey are due to being eaten
by the predators.We assume a constant per-capita death rate for the predators independent of
the prey density. At any time the per-capita birth rate will increase with more food available
and depend heavily on the amount of prey available.
To model this we assume that the overall birth rate for the predators is directly proportional to
the food supply. i.e., number of prey killed by the predators.
 Prey per-capita birth rate is constant.
 Predator per-capita death rate is constant.
 Prey per-capita death rate is proportional to the predator density.
 Predator per-capita birth rate is proportional to the rate at which prey are eaten.
Example
Using the assumption and word equations for the prey and predator densities.

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Solution: let X ( t ) denotes the number of prey per unit area and Y (t ) the number of predator
per unit area.
Let the prey per-capita birth rate is β 1. Predator per-capita death rate be α 2.

{ } { }
rate of rate of
prey =β 1 X ( t ) , predator =α 2 Y (t) (2)
birth deaths

Prey deaths we denote the per-capita death rate as c 1 Y ( t ) .c 1is positive constant of
proportionality. Thus the rate at which prey are eaten is given by c 1 Y ( t ) X (t). The predator
birth rate is proportional to this rate of prey eaten.

{ } { }
rate of rate of
prey killed =c 1 Y ( t ) X ( t ) , predators =f c1 Y ( t ) X ( t) (3)
by predators deaths

Where f is a positive constant of proportionality. X ( t ) Y (t) is that it is proportional to the rate

of encounters between prey and predators.
Now substitute equation (2) and (3) into word equation (1)
dX dY
=β 1 X −c 1 XY , =c 2 XY −α 2 Y (4)
dt dt
Where c 2=f c 1

Exercise
(1) Check the Lotka -Volterra model in the limiting cases of prey with no predators or
predators with no prey.
Solution : Suppose there are no predators Y =0.
The equations then reduces to
dX
=β 1 X
dt
Which is the equation for exponential growth. The prey grows exponentially.
If there are no prey, then X =0 ,
dY
∴ =−α 2 Y
dt

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That is exponential decay. Which means that the predator population decreases exponentially
and dies out.
(2) Two prey and one predator, Develop a model with three differential equations
decreasing a predator-prey interaction, where there are two different species of prey and one
species of predator.
Solution: let X ( t ) denotes the number of prey per unit area. Y (t ) denote the number of
predator per unit area.

{ } { }
rate of rate of
prey 1 =β 1 X ( t ) , predator =α 2 Y (t);
birth deaths

{ } { }
rate of rate of
prey 2 =β 3 X ( t ) ; predator =α 4 Y ( t ) ;
births deaths

For prey1 death we denote the per-capita death rate C 1 Y ( t ). For prey2 death we denote the
per-capita death rate D1 Y (t).
C 1∧D1 as the positive constants.

{ }
rate of prey 1
killed by =C1 Y ( t ) X (t )
predators

{ }
ratr of
predator =f C 1 Y ( t ) X (t)
birth

{ }
rate of prey 2
killled by =D1 Y ( t ) X (t)
predator

{ }
rate of
predator =f D1 Y ( t ) X (t)
birth

dX dY
=β 1 X −C 1 XY ; =C2 XY −α 2 Y
dt dt

dX dY
=β 2 X−D 1 XY ; =D1 XY −α 4 Y
dt dt
3.5 Competing species

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Another simple ecosystem to model is that of competing species, where two (or more)
populations compete for limited resources such as food or territory. There are two aspects of
competition : exploitation, when the competitor uses the resource itself and
interference ,where the population behaves in such a manner as to prevent the competitor
from utilizing the resource.
This system is very similar to the predator – prey model.

Model assumption
We assume the population to be sufficiently large so that random fluctuations can be ignored
without consequence. we assume that two species model reflects the ecosystem sufficiently
accurately. we assume each population grows exponentially in the absence of the other
competitor(s), although we later incorporate density dependent growth for each.
Let x(t) and y(t) be the two population densities (number per unit area) where t is again
time.as before we have birth and death rates associated with each population. However , in
this case an increase in the number of deaths in one population causes a decrease in the
number of deaths in other population.
In words, we have for each population in the system

{ }{ }{ }
rate of rate of rate of
change of = population − population (1)
population births death

Formulating the differential equations

Since neither population is dependent on the other as for as growth rates (birth rates) are
concerned (unlike in the predictor – prey β 1and β 2describe the per – capita birth rates for
species x and y respectively. Since the two populations are competing for the same resource ,
the density of each population has a restraining effect, proportional to this density , on the
other.
So the pre-capita death rate for Y is proportional to X and that for X is proportional to Y ,
which in symbols is

{ }
rate of
species X =( c 1 Y ) X ,
deaths

{ }
rate of
species Y = (c2 X ) Y
deaths

Here c 1and c 2 are the constants of proportionality for this restraining effect
Our model becomes,

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dX dY
=β 1 X −c 1 XY ; =β 2 Y −c 2 XY (2)
dt dt
Where β 1∧β 2 were per-capita birth rate, we may also consider them as overall per-capita
growth rates which incorporate deaths (independent of the other species) as well as births.
Thus they are per-capita growth rates, or per-capita reproduction rate. While the parameters
c 1∧c 2 are the intersection parameters. These equations are known as Gauses equation and are
a coupled pair of first order non-linear differential equations.

Interpretation of parameters:
The system of equations developed so for the model of competitive species has four
parameters: the two per-capita growth rates β 1∧β 2 which are independent of any other
species, and the two constants of proportionality c 1∧c 2 which describes the interaction
between the species. The competition intersection can be interpreted as stating that the more
deaths there are in species Y, the more resource will be available to species x and thus the
fewer deaths there will be for X.
To explain interaction we consider a small time interval ∆ t ,sufficiently small so that X and Y
do not change significantly.
Let a ' be the rate of area used by one individual in species Y (for food), hence making it
unavailable to X. Assume that no such area can overlap. Then one individual ‘uses’ an area of
' '
a ∆ t ,∧Y individual will ‘use’ a Y (t)∆ t .
Assuming that each area of resource ‘used’ by Y implies it is unavailable to X, the rate at
which area is made unavailable is the density (number per unit area) of X in an area divided
by ∆ t , which is a ' X ( t ). And hence the total effect of the removal of resources from X by Y
becomes a ' X ( t ) Y (t). So a ' ,∨c 1 in our model can be interpreted as the rate at which X dies
off due to the removal of resources by Y.
c2
If we let f = ,then f is a measure of the efficiency of the competitive interaction. It can be
c1
considered as the number of units of species Y required to reduce species X by one unit.

Limitations of the model:

 One immediate and obvious limitation of this model is that each population grows
exponentially in the absence of the other.
 Next we improve on this by including density dependent growth.
 Early this century, research tended to accept and support Gauses principle (i.e., the
result suggest that in the case of two species competing for the same resource, in the
long run one species will survive and the other become extinct).

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Scenario: Aggressive protection of lerps and nymphs

The following scenario does not incorporate the above theory specifically, but describe the
occurrence of species, but describes the occurrence of species competing for territory with a
colony of birds protecting its territory in order to maintain an adequate food supply to support
itself. It is included an example of the value of interspecific territoriality, a system for which
the above theory would have relevance.
In regions of southeastern Australia, infestation of psyllids have inflicted severe damage on
the eucalypt forests causing deterioration of the foliage, and in extremes cases, completely
destroying certain trees. This has prompted an examination of the conditions under which it
may occure many species of birds live in the canopy of these eucalypts and complete to feed
on these insects; the Bell Miners are one such species.
Bell Miner ( Manorina melanophrys ) is a honeyeater living in large colonies that they define
aggressively against other competing birds often subsequently larger than themselves. They
feed on the nymphs, sweet secretions and lerps (protective carbohydrate covers) of psyllids;
however where their colonies occur, the canopy foliage appears unhealthy and infested with
these insects. The Miners aggressively protected this abundance of prey.
In an experiment a colony of Bell Miners was removed to join another colony 45km away.
They settled into this new location and did not return.
Other bird flocks soon moved into the site of the old colony and rapidly the foliage improved.
This insect infestation declined, with the numbers of psyllids remaining very small thereafter.
It appears that the insect infestations can be controlled by the removal of the Bell Miner
colonies. The interspecific territorial behavior of the Miners prevents this control and
introduces a control of its own. Miners effectively maintain an abundant and exclusive food
supply, albeit detrimental to the eucalypts, enabling a colony to remain within the same
territory for up to 40 years.
3.6 Analysis of an epidemic model:
Here we use the chain rule and some analysis to prove that the disease, described by this
model, can never infect the entire population.
Review of the model
In the epidemic model, assumed the population 𝑁 was divided into susceptibles, denoted by
𝑆(𝑡), infectives, denoted by 𝐶(𝑡) and removals 𝑁 − 𝑆(𝑡) − 𝐶(𝑡). We assumed the disease to
confer life-long immunity and we neglected to include natural births and deaths. The model
pair of differential equations we obtained was
dS dC
= −𝑏𝑆𝐶, = 𝑏𝑆𝐶 − 𝑟𝐶. − − − − − (1)
dt dt
The parameter 𝑏 is the transmission coefficient and 𝑟 is the recovery rate.
Example: Use the chain rule to find 𝐶 in terms of 𝑆, given that the initial number of
susceptibles is s0 and the initial number of contagious infectives is c 0
Solution: Using the chain rule

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dC
dC dt
=
dS dS
dt
Substituting from the differential equations (1) we eliminate the time variable, obtaining the
single differential equation
dC bSC−rC
=
dS −bSC
Which simplifies to
dC r
=−1+ (2)
dS bS

This differential equation relates the number of infectives 𝐶 to the number of susceptibles 𝑆
but does not retain explicit information about time.
The differential equation (2) is a first-order separable differential equation of a trivial kind
and to solve it we have only to integrate both sides with respect to 𝑆.
The solution is
r
C=−S+ ln(𝑆) + 𝐾 − − − −(3)
b
Where 𝐾 is an arbitrary constant of integration.
The initial numbers of susceptibles and infectives are 𝑠଴ & 𝑐଴, respectively, and hence the
initial condition for the differential equation (2) is
C ( s 0 )=c 0

Applying this to the general solution (3) gives an equation for 𝐾 from which we deduce that
r
𝐾 = c 0 + s0 − ln( s0 )
b

Exercise:
(1) Rabbits and foxes. A population of sterile rabbits 𝑋(𝑡) is preyed upon by a population of
foxes 𝑌(𝑡). A model for this population interaction is the pair of differential equations

dX dY
=−aXY , =bXY −cY − − − −(1)
dt dt

Where 𝑎, 𝑏 𝑎𝑛𝑑 𝑐 are positive constants. Use the chain rule to obtain a relationship
between the density of foxes and the density of rabbits.

Solution:

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Using chain rule

dY
dY dt
=
dX dX
dt

The required differential equation

dY bXY −cY
=
dX −aXY
Which simplifies to

dY −b c
= + (2)
dX a aX

This differential equation relates the number of sterile rabbits X to the number of
susceptible S but does not retain explicit information about time.
Solve and integrate both sides of equation (2) with respect to X .
This solution is,
−b c
Y= X + ln ( X ) + K −−−−( 3)
a a

Putting initial conditions

X ( y 0 ) =x 0 Y ( x 0 )= y 0

Applying this to the general solution equation (3) gives an equation for K from which we
deduce that
b c
K= y 0+ x 0− ln ⁡(x 0 )
a a

(2) Microorganism and toxins: the pair of differential equations

dP dT
=rP−γPT , =qP
dt dt

Where r , γ ∧q are positive constants, is a model for a population of microorganism P,

which produces toxins T which kill the microorganism.

Given that initially there are no toxins and P0 microorganisms, obtain an expression
relating the population density and the amount of toxins.(hint use the chain rule)

Solution:
dP dT
=rP−γPT , =qP−−−−(1)
dt dt

Where r , γ ∧q are positive constants.

Use the chain rule,

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dP
dP dt
=
dT dT
dt
Which simplified to,

dP r −γT
=
dT q

The differential equation (2) is a first order seperable differential equation of trivial kind
and to solve it we have only to integrate both sides with respect to P.

The solution is,

rT γT
P= − + c−−−−(3)
q 2q

With initial condition c ( P0 )=P 0

Applying the general solution equation (3) gives an equation for P(t) from which we
deduce that,
rT γT
P ( T ) = − + P0
q 2q

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