Mathematical Modelling Group 11 Project 3

Continuous Population Modelling - Predator Prey

Contributors
Alexandra Hunt, Paul Martin, Lorna Gollop, Andrew Haspey and Jasmine Dhaliwal

1. Introduction .
In nature all organisms are mutually dependent on each other, so a change in the
population of one organism will have a knock on effect on the population of another
organism. For this reason, it is important that we model the interactions between
organisms. The interactions between organisms can either lead to co-existence,
extinction of one or both species, or symbiosis.
In the past a variety of modelling ideas have been used. The first population
modelling was carried out by Thomas Malthus in 1798 in a paper he wrote titled
Essay on the Principle of Population. As shown in Figure 1, Malthus argued that
populations grew logarithmically, but the populations that they depended on
remained constant or only increased arithmetically. Thus, the demand for resource
supply would cause the population to reach a natural level before dying off
(Berryman, 1992).

Figure 1: Urbano, 2011. A

graph showing Malthusian
Growth, showing that the
population will always over
take resources.

We decided to use the Lotka-Volterra model as it looks at multiple species

interactions. The Lotka-Volterra model was developed independently by Alfred Lotka
and Vito Volterra (Beals et al., 1999). The formula is the most successful attempt at
modelling the relationship between prey and predator (Lotka, 1922, Volterra, 1928;
cited in Bas, 2000). The Predator-Prey model plays a vital part in population
dynamics because it is able to determine the biomass transferred in an association
of different organisms forming a closely integrated community (Poggaile, 1998). The
Lotka-Volterra predator-prey equations were built around three main assumptions:
1. The Malthus Law is applied for both models
2. The principle of mass action so the responses were assumed to be
proportional to the product rather than the ratio of population densities
3. The predator density growth is proportional to the density of eaten prey.
(Berryman, 1992; Poggaile, 1998).
If the predator was absent, the population of prey would grow exponentially,
whereas in the absence of the prey, the predator population would decay
exponentially due to starvation.
We decided to use the Lotka-Volterra model to investigate the predator-prey
relationship between Brown Bears and Chum Salmon.
Our aim was to investigate whether the two populations could co-exist and if
so we aim to investigate what impact another factor would have.

2. Mathematical Model

2.1 Assumptions
No human influence.
Constant water level and flow a change increase in water level would make
it harder for the salmon to get up the stream, whilst a decrease in water level
would make it easier for the bears to catch the salmon.
The Bear only eats Salmon and they dont get their nutrients from anywhere
else this is to ensure the population can be accurately modelled.
The prey population has a constant source of food.
The rate of change of the populations are proportional to their size.
During the process, the environment does not change in favour of one
species and genetic adaptation is inconsequential.
Predators have limitless appetite.

Initially we decided to assume that there was a constant Birth and Death Rate
however, we then decided not to assume this as during the period of time, the
salmon do not reproduce.

2.2 Variables
Independent Variable: Salmon Population
Dependent Variable: Bear population
2.3 Model Construction and analysis
2.3.1 Sub-problem 1
This is our initial problem, with the two species interacting with each other, with no
other interactions made.
The rates of change of the populations , are denoted

!"
!"

and

!"
!"

respectively,

where () represents the Salmon population and () represents the Bear

population. The constant positive parameters , , and represent net growth rate
of , the interaction of with , the death rate of and the interaction of with .

=
= +

= 0 = 0

= 0,

+ = 0 + = 0

= 0,

! !

Therefore the Equilibrium Points are at (0,0) and (!, ! )

For (0,0)
:

det =

= ( )( )
! = ! =

!, !

This implies that we have a saddle.

= 0
0
0
=

+ 0 = 0 = 0
, = 0 = 0
! !

For (!, ! )
:

det =

= !

= ! +

This is a stable centre.

( , )

We have used Mathematica to plot the phase-plane diagram.

Figure 2

In Appendix 1 we used a=b=c=d=1 and plotted it on Mathematica. We can see from

Figure 2, that there is a saddle point at the origin and a stable centre at (1,1).

Figure 3

Figure 3 shows the oscillation of the population of bear and salmon against time.
() (red line) denotes the population of the salmon and () (blue line) denotes the
population of the bears.

2.3.2 Sub-problem 2
!"

The rates of change of the populations , are denoted

!"

and

!"
!"

respectively,

where () represents the Salmon population and () represents the Bear

population. The constant positive parameters , , , and represent net growth
rate of , the interaction of with , the death rate of , the interaction of with
and carrying capacity constant in relation to .
This is the logistic model, where the prey could grow unboundedly.

= ! = 0
= +

! !!!!"

Therefore the Equilibrium Points are at (0,0), ( ! , 0) and (!,

2

det =

!, !

= ( )( )
! = ! =

This implies we have a saddle.

!

For ( ! , 0)
:

=
0

det =

= 0

= 0

We therefore have a stable node.

! !!!!"

! = ! = +

, !

For (!,

!"

!"

2
0

det =

! !!!"
,
!
!"

= 0
0

!
= + +

!
det =

4 det =

4 4 !

Since Tr(A)<0, both values are positive real eigenvalues.

Whenever 0<4det(B)<Tr(A) the eigenvalues are real and distinct and we have a
node that is stable if Tr(A)<0 and unstable if Tr(A)>0
Since our Tr(A) =

!!"
!

<0 it follows we have a stable node.

Appendix 2 shows a worked example where the constant positive parameters , , ,

and have been replaced with real values: = 2, = = = = 1. Figure 4
illustrates this model when plotted using Mathematica. The diagram displays a stable
spiral, unstable node and a stable node. Therefore the two populations can co-exist.

Figure 4

Figure 5 (below) shows a graph of each population plotted against time. As you can
see, the predator starts out high and the prey low, after the interaction both
populations plateau at a constant value.

Prey
Predator

Figure 5

2.3.3 Sub-problem 3
The rates of change of the populations , are denoted

!"

and

!"

!"
!"

respectively,

where () represents the Salmon population and () represents the Bear

population. The constant positive parameters , , , , and represent net growth
rate of , the interaction of with , the death rate of , the interaction of with ,
carrying capacity constant in relation to and the carrying capacity constant in
relation to .
For our third model we have a double logistic model.

= ! = 0
= + ! + = 0

2 = 0 = 0
2 + = 0 + = 0

Therefore, our Equilibrium points are at 0,0 , 0, ! ,

!
!

, 0

For 0,0
:

det =

2 +

!, !

= ( )( )
! = ! ! =

We therefore have an unstable node.

!

For 0, !
:

det =

= (

2 +

0

2

)( )

!,

!
!

!"!!" !"!!"

!"!!" !"!!"

 ! =

! =

This is a saddle.
For
:

!
!

,0

det =

2 +

2
0

= ( )

!
, !
!

! = ! =

Due to the complexity of the 4th equilibrium point, we decided just to apply numbers
straight to it, as shown in Appendix 3. We then used Mathematica to plot Figure 6.
illustrates this model when plotted using Mathematica. The diagram displays a
unstable node, two saddle points and a stable spiral. Therefore the two populations
can co-exist.

10

Figure 6

Figure 7 (below) shows a graph of each population plotted against time. As you can
see, the predator starts out high and the prey low, after the interaction both
populations plateau at a constant value. As the predator population decreases, the
prey population increases. This graph is almost identical to figure 5.

Prey
Predator

Figure 7

11

3. Discussion
As expected, all our equilibrium points are positive and in the first quadrant, since
populations cannot be negative. Appendix 1 -3 show some worked examples, and
the corresponding graphs also reflect what we expected the outcome to be.
Our basic model shows that the two species can co-exist at the trivial point
! !
(0,0) and then at (!, ! ). This is what we expected, that both species can have large

populations and can exist side by side. At (0,0) the equilibrium point is a saddle, this
means that it is an unstable point and a small deviation in the population will take
! !
you away from the equilibrium point. At (!, ! ) it is a stable centre, this means that coexistence is possible, which is what we expected.

For the logistic model, we have three equilibrium points. The trivial point of
!

! !!!!"

(0,0) and ( ! , 0) and (!,

!

!"

). The trivial point is a saddle which is the same as our

basic model. At ( ! , 0) we have two possible outcomes, ! will always be positive, !

can either be positive or negative. In the instant when ! is positive we will have a
saddle point, however, when ! is negative we have an asymptotically stable node.
! !"!!"

This implies that we would have a stable situation. At (!,

!"

) we have an unstable

node, this implies that the populations are unlikely to reach this point due to there
being an unstable system.
For the double logistic model, we have four equilibrium points, at
!

0,0 , 0, ! ,

!
!

, 0

!"!!" !"!!"

!"!!" !"!!"

. (0,0) is an unstable node which implies that

!

the population will always deviate away from this point. At 0, ! and

!
!

, 0 we have

saddles which, again, imply that a small deviation in the populations will take you
away from the equilibrium point. With regards to our 4th equilibrium point, with the
figure we put in, we resulted in a stable spiral. This implies that although the
populations are expected to fluctuate greatly, eventually they will reach a point where
they are able to co-exist together. However, we realise that this may not always be
the case for all values input.
For all our models we used Mathematica to plot the graphs. This was a very
time consuming task, as only one member of the group had any knowledge of the
complicated programme. The plots from Mathematica confirmed the points that we
had worked out.
The largest difficulty that we had was trying to classify the eigenvalues. We
got our selves stuck by doing unnecessary specific mathematics which over
complicated things.

12

4. Conclusion
We achieved our aim, and found that another factor impacts the model greatly. In
this instance a fisherman could be the influential factor.
Due to limited time and resources we had to make a number of assumptions in order
to simplify our phase-plane models. If we had more time we could have made other
refinements to make our model reflect the real world more accurately. These
refinements include:
-
-

Introducing another source of food for the bears and finding out what impact
they have on the salmon population.
Thinking about the weather and the climate. If there was a bad storm then
there could be a reduction in the number of salmon that make it back to the
stream. Over longer periods of time there could be an increase in surface sea
level temperatures which could cause a decrease in salmon numbers. Or, an
increase in temperature could cause rivers to have more water in due to a
greater rate of melting, this could make it harder for the bears to fish.
Introducing disease that could significantly reduce numbers of one or more
species.

Actual Contributors
Alexandra Hunt, Paul Martin, Lorna Gollop, Andrew Haspey and Jasmine Dhaliwal

5. References

Bas, Carles (2000). Fishery science: analysis and present situation. Contributions to
Science. Vol. 1, No. 4, pp. 489-510
Beals, M., Gross, L., Harrell, S. (1999). Predator-Prey Dynamics: Lotka-Volterra
http://www.tiem.utk.edu/~gross/bioed/bealsmodules/predator-prey.html Accessed
02/05/15
Berryman, Alan A. (1992). The Origins and Evolution of Predator-Prey Theory.
Ecology. Vol. 73, No. 5, pp. 1530-1535
Poggiale, J. C. (1998). Lotka-Volterras Model and Migrations: Breaking of the WellKnown Centre. Mathematical Computer Modelling. Vol. 27, No. 4, pp. 51-61
Urbano, L. (2011) Malthusian Growth.
http://montessorimuddle.org/2011/06/30/malthusian-growth/ Accessed 02/05/15

13

Appendix 1 A worked example of our basic scenario

In the instant where a=b=c=d=1:

=
= +

= 0 1 = 0
= 0,

= 1

+ = 0 1 + = 0
= 0,

= 1

Therefore the Equilibrium Points are at (0,0) and (1,1)

For (0,0)
:
=

1
0

1 +

!, !

0
=
1
1
0

det =

0

1

= 1 1 = 0
!,! = 1

This is a saddle
For (1,1)
:
=

0
1

1 +

1
=
0

det =

0
1

= ! + 1 = 0
!,! =

This is a stable centre.

!, !

14

1

0

Appendix 2- A worked example of our logistic equation

In the instant where:
= 2, = = = = 1

= 2 ! = 0
= + = 0

2 ! = 0 2 = 0
= 0 = 2,

= 1

+ = 0 1 + = 0
= 0,

= 1

Therefore the Equilibrium Points are at (0,0), (2,0) and (1,1)

For (0,0)
:

2 2

2
0

det =

1 +

!, !

0
=
1
2
0

0

1

= (2 )(1 ) = 0
! = 2,

! = 1

This implies a saddle

For (2,0)
:

2 2

2
0

det =

1 +

!, !

2
=
1
2
0

2

1

= (2 )(1 ) = 0
! = 2,

! = 1

This also implies a saddle

For (1,1)

15

2 2

1
1

1 +

!, !

1
=
0
1
1

det =

1

0

= ! + + 1 = 0
!,! =

1 3
2

This implies a stable spiral.

16

Appendix 3 A worked example of our double logistic model.

At the point a=100, r=1, b=1, c=30, q=1, d=1

= 100 ! = 0
= 30 + ! + = 0

For 0,0
:

100 2

100
0

det =

30 2 +

!, !

0
=
30
100
0

0

30

= (100 )(30 ) = 0
! = 100,

! = 30

For 0, !
:

100 2

70
30

30 2 +

0

30

70
30

det =

!, !"

0

30

= 70 30 = 0
! = 70,

! = 30

For
:

!
!

,0

100 2

30 2 +

100
0

det =

100

130

100
0

100

130

= 100 130 = 0
! = 100,

For

!"", !

! = 130

!"!!" !"!!"

!"!!" !"!!"

17

100 2

35
65

det =

30 2 +

!", !"

35

65

35
65

35

65

= 35 65 + 2275
! + 100 + 4550
!,! =

100 17200

2

Since ! and ! are complex conjugates with both real parts equalling less than zero, we
therefore have a stable spiral.

18

Appendix 4- Group Minutes

Meeting 1 2 hours 26/03/15
Attendees: Lorna, Alexandra, Paul, Jasmine, Andrew, Saranyan

Decided on population problem

Bears and Salmon
Will all investigate and do some reading over Easter

Meeting 2 1 hour 20/04/15

Attendees: Lorna, Alexandra, Paul, Jasmine, Andrew

Started working on the model

Meeting 3 1 hour 21/04/15

Attendees: Lorna, Alexandra, Paul, Andrew, Jasmine

Background research on both animals

Noted some factors which would affect the population rate of salmon, other
than bears. Weve made a start on constructing our simple interaction models
using Lotka-Volterra equations.

Meeting 4 2 hours 22/04/15

Attendees: Lorna, Alexandra, Paul, Andrew, Jasmine

We had started discussing the phase plane equations. Started to take into
consideration all the possible scenarios i.e. the bear population decreasing
and the effect this has on the salmon population and the effect a decrease in
the salmon population would have on the Bears population.

Note: Andrew had turned up 25 minutes before the session ended and had no input
with the discussion.
Meeting 5 1 hour 27/04/15
Attendees: Lorna, Alexandra, Paul, Andrew, Jasmine

We continued work on the phase-plane diagram.

We started a rough draft of the final report.

Meeting 6 1hour 28/04/15

Attendees: Lorna, Alexandra, Paul, Jasmine, Andrew

We continued work on the phase plane equations and started plotting them on
Mathematica. The report has been started and a plan made for it.

Note: Saranyan told us that he wasnt going to be part of the group anymore

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Meeting 7 2.5 hours 29/04/15

Attendees: Lorna, Alexandra, Paul, Jasmine, Andrew

We met with Dr Naire before the lecture to clear things up.

During the lecture Andrew and Jasmine worked on Mathematica, Alexandra

and Paul worked on the maths. Lorna typed up the report so far.

Meeting 8 3 hours 04/05/15

Attendees: Lorna, Alexandra, Paul, Jasmine, Andrew

Andrew worked on Mathematica

Alexandra and Paul worked on the maths

Lorna had written introduction over the weekend and wrote up the maths

Jasmine wrote about the maths and had to leave after 1 hour

Meeting 9 2 hours 05/05/15

Attendees: Lorna, Alexandra, Paul, Jasmine, Andrew

Andrew worked on Mathematica

Alexandra and Paul worked on the maths

Lorna worked on the report

We had a meeting with Dr Naire prior to the lecture. Paul was unable to attend due
to a lecture. Lorna, Alex and Jasmine attended, Andrew wasnt present.

Meeting 10 3 hours 06/05/15

Attendees: Lorna, Andrew, Paul, Alexandra, Jasmine

Paul and Alexandra finished up working on the third (more refined) model.
Lorna and Jasmine continued the type of the report.
Andrew created the phase-plane diagram for the third model on Mathematica.

Note: Jasmine left after 2 hours as she had another group project to work on.
Andrew arrived an hour late.

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