Stability Analysis of a Prey-Predator

Model
Incorporating a Prey Refuge
Project report of
Mathematical Modeling (MA539)
Submitted by-
Prabhat Kumar
Research Scholar
2221MA04

under the supervision of

Dr. Prashant Kumar Srivastava
Associate Professor
IIT Patna

November, 2022

Department of Mathematics
Indian Institute of Technology, Patna
 ACKNOWLEDGEMENTS

I would like to extend my sincere thanks to the course instructor Dr. P. K. Srivastava,
Associate Professor, Department of Mathematics, IIT Patna for teaching the course on Math-
ematical Modelling (MA 539). His wonderful teaching skill makes him a visionary and in-
spiring professor.

Prabhat Kumar

I
Contents

ACKNOWLEDGEMENT I

1 Introduction 1
1.1 Objective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

2 DESCRIPTION AND FORMULATION OF THE MODEL 2

2.1 Basic Assumptions: . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2.2 Model Parameters: . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2.3 Model Equations: . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3 ANALYSIS OF THE MODEL 4

3.1 Equilibria : . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3.2 Boundedness of the System . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3.3 Dynamical Behaviour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

4 Numerical Simulation 8

5 Conclusion 12

II
Chapter 1

Introduction

In the dynamics of the ecosystems,the relationship between prey and predator has been
long and will continue to be one of the dominant themes due to its universal existence and
importance.Although these problems first seem to have easy mathematical solutions, they
are frequently quite difficult and complex.

Although the predator–prey theory has seen much progress in the last 50 to 60 years,
many long standing mathematical and ecological problems remain open. Differential equa-
tion models for interactions between prey-predator are one of the classical application of
mathematics to biology.

In this presentation we analyze a Lotka-Volterra type prey-predator model with Michaelis-

Menten type II functional response.

In addition,a spatial refuge protects a constant proportion of prey from predation.Due

to this spatial refuge, prey get some protection from predation and reduce the chance of ex-
tinction due to preda- tion.Maynard Smith shows that the presence of a constant proportion
refuge does not alter the dynamical stability of the neutrally stable Lotka-Volterra model,
while a constant number refuge of any size replaces the neutrally stable behaviour with a
stable equilibrium.

1.1 Objective
The main goal of this mini project is to study the mathematical modelling and stability
analysis of prey-predator model incorporating a prey refuge. A prey-predator model is used
to formulate a system of ordinary differential equations.The stability theory of differential
equation is used to investigate the qualitative behaviour of the system.
The purpose of the work is to offer mathematical analysis of the model and to discuss some
important results that are expected to arise from the interplay of biological forces.The nu-
merical simulation is also carried out.
Keywords : Prey–predator; Refuge; Stability; Persistent; Limit cycle.

1
Chapter 2

DESCRIPTION AND FORMULATION

OF THE MODEL

2.1 Basic Assumptions:

Stability Analysis of a Prey-Predator Model Incorporating a Prey Refuge is based on the
following assumptions:

• Let x(t) and y(t) be the population density of the prey and predator species at any time
t.

• The population density of the prey has limited food resource,so in the absence of preda-
tor the population density of the prey will grow logistically with intrinsic growth rate.

• The food supply of the predator population depends entirely on the size of the prey
population,so in the absence of prey the population density of predator could decrease
at the natural rate.

• There are no changes in the environment which would favor one species.

• Predators can consume infinite quantities of prey.

• This presentation extends the Lotk-Volterra model with Holling Type II functional
response by incorporating a refuge protecting mx of the prey,where 0 ≤ 𝑚 < 1 is
constant.This leaves (1-m)x of the prey available to the predator.

• All the parameters are positive.

2.2 Model Parameters:

The description of parameters used in this model is listed below :

• 𝛼 : Intrinsic growth rate of the Prey.

• 𝑘 : Carrying capacity of the Prey.

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 • 𝛽 : Consumption rate of the Predator.

• 𝑎 : Handling time of a Prey by Predator.

• 𝛾 : Death rate of the Predator.

• 𝑐 : Conversion factor denoting the number of newly born predators for each captured
prey.

• 𝑚 : Prey refuge.

2.3 Model Equations:

Considering all the variables, assumptions, and parameters the general form of this model
is given by the following system of ordinary differential equations :

𝑑𝑥 𝑥 𝛽(1 − 𝑚)𝑦𝑥
= 𝛼𝑥(1 − ) −
𝑑𝑡 𝑘 1 + 𝑎(1 − 𝑚)𝑥
𝑑𝑦 𝑐𝛽(1 − 𝑚)𝑥𝑦
= −𝛾𝑦 +
𝑑𝑡 1 + 𝑎(1 − 𝑚)𝑥

3
Chapter 3

ANALYSIS OF THE MODEL

To ensure the existence and uniqueness of above system,we seek the solution in 𝑅+2 = {𝑥, 𝑦 ∶
𝑥 > 0, 𝑦 > 0} so that all the standard results of existence,uniqueness and continuous depen-
dence on initial conditions are evidently satisfied.

3.1 Equilibria :
We now study of equilibrium point of the model.Particularly we interested in the interior or
positive equilibrium point.So for equilibrium point we solve the following equations for x
and y :
𝑥 𝛽(1 − 𝑚)𝑦𝑥
𝛼𝑥(1 − ) − =0 (1)
𝑘 1 + 𝑎(1 − 𝑚)𝑥
𝑐𝛽(1 − 𝑚)𝑥𝑦
−𝛾𝑦 + =0 (2)
1 + 𝑎(1 − 𝑚)𝑥
Case I

When x=0,y=0

We get trivial equilibrium point 𝐸0 = (0, 0)

Case II

When y=0,𝑥 ≠ 0

That is,equilibrium point in the absence of predator 𝐸1 = (𝑘, 0)

Case III

When 𝑥 ≠ 0,𝑦 ≠ 0

4
 The interior(positive) equilibrium point 𝐸2 = (𝑥 ∗ , 𝑦 ∗ ),where
𝛾
𝑥∗ = (3)
(𝑐𝛽 − 𝛾𝑎)(1 − 𝑚)
,
𝛼𝑐 𝑘(𝑐𝛽 − 𝛾𝑎)(1 − 𝑚) − 𝛾
[ 𝑦∗ = ] (4)
𝑘 (𝑐𝛽 − 𝛾𝑎)2 (1 − 𝑚)2
For the equilibrium point 𝐸2 to be positive we first need

𝑐𝛽 − 𝛾𝑎 > 0

For 𝑦 ∗ to be positive, the condition on m to be

𝛾
0≤𝑚<1− (5)
𝑘(𝑐𝛽 − 𝛾𝑎)

Thus,for the existence of the positive equilibrium both 𝑐𝛽 > 𝛾𝑎 and

0≤𝑚<1− 𝛾
𝑘(𝑐𝛽−𝛾𝑎)
must be hold.

Also we see that 𝑥 ∗ increases as m.

3.2 Boundedness of the System

The boundedness of solutions of the system is proved in the following theorem.

Theorem 1. All the solutions of system which start in 𝑅+2 are uniformly bounded.

Proof. We define the function w = c x + y. Therefore, time derivative

𝑑𝑤 𝜕𝑤 𝑑𝑥 𝑑𝑦
𝑑𝑡
= + 𝜕𝑤
𝜕𝑥 𝑑𝑡𝜕𝑦 𝑑𝑡

or, 𝑑𝑤
𝑑𝑡
= 𝑐 𝑑𝑥
𝑑𝑡
+ 𝑑𝑦
𝑑𝑡
= 𝑐[𝛼𝑥(1 − 𝑥𝑘 ) − 𝛽(1−𝑚)𝑦𝑥
1+𝑎(1−𝑚)𝑥
] + −𝛾𝑦 + 𝑐𝛽(1−𝑚)𝑥𝑦
1+𝑎(1−𝑚)𝑥

so, 𝑑𝑤
𝑑𝑡
= 𝛼𝑐𝑥(1 − 𝑥𝑘 ) − 𝛾𝑦
For 𝛾 > 0, we have,
𝑑𝑤 𝑐𝑘(𝛼 + 𝛾)(𝛼 − 𝛾) 𝛾𝑐𝑘(𝛼 + 𝛾)
+ 𝛾𝑤 ≤ +
𝑑𝑡 4𝛼 2𝛼
Now if we choose 𝛼 > 𝛾, then right hand side is bounded for all (𝑥, 𝑦)𝜖𝑅+2 .So we choose a L
> 0, such that
𝑑𝑤
+ 𝛾𝑤 < 𝐿.
𝑑𝑡
Now we apply the theory of 1st order differential linear differential equation to solve
𝑑𝑤
+ 𝛾𝑤 = 𝐿
𝑑𝑡
which is linear in w,after that taking some initial value we get bounded solution in 𝑅+2 .

5
3.3 Dynamical Behaviour
In this subsection we will discuss the stability property of equilibrium point 𝐸0 , 𝐸1 𝑎𝑛𝑑𝐸2 .

The Jacobian of the system about the equilibrium point 𝐸0 = (0, 0) is given by

𝛼 0
𝐽𝐸0 (0,0) =
[ 0 −𝛾 ]

Hence the eigenvalues of the system are the roots of the equation (𝛼 − 𝜆)(−𝛾 − 𝜆) = 0.

Here one eigenvalue is positive and another is negative.

Therefore 𝐸0 is a saddle point.

Jacobian matrix for 𝐸1 is given by

−𝛽(1−𝑚)𝑘
−𝛼 1+𝑎(1−𝑚)𝑘
𝐽𝐸1 (𝑘,0) = 𝑐𝛽(1−𝑚)𝑘
[ 0 −𝛾 + 1+𝑎(1−𝑚)𝑘 ]

The eigenvalues of the matrix are −𝛼 and −𝛾 + 1+𝑎(1−𝑚)𝑘

𝑐𝛽(1−𝑚)𝑘
. Hence 𝐸1 is locally asymptoti-
cally stable when 𝑚 > 1 − 𝑘(𝑐𝛽−𝛾𝑎) and unstable (saddle) when 𝑚 ≤ 1 − 𝑘(𝑐𝛽−𝛾𝑎)
𝛾 𝛾
.

Here we observed that, if 𝐸1 is unstable (saddle) then 𝐸2 exists.

Now, Jacobian matrix for 𝐸2 = (𝑥 ∗ , 𝑦 ∗ ) is given by

𝑎1 −𝑎2
𝐽𝐸2 =
[ 𝑎3 0 ]

where
𝑎1 = 𝛼 − 2𝛼𝛾
𝑘(𝑐𝛽−𝛾𝑎)(1−𝑚)
− 𝛼[𝑘(𝑐𝛽−𝛾𝑎)(1−𝑚)−𝛾]
𝑘𝑐𝛽(1−𝑚)
,

𝑎2 = −𝛾
𝑐
,

𝑎3 = 𝛼[𝑘(𝑐𝛽−𝛾𝑎)(1−𝑚)−𝛾]
𝑘𝛽(1−𝑚)
.

The characteristic equation is

𝜆2 − 𝑎1 𝜆 + 𝑎2 𝑎3 = 0.
The sum of the roots is equal to 𝑎1 and the product of the roots is equal to 𝑎2 𝑎3 , is always
positive by condition 5.

6
Now 𝑎1 will be negative if
𝛾 𝑐𝛽
𝑚>1− − (6)
𝑘(𝑐𝛽 − 𝛾𝑎) 𝑘𝑎(𝑐𝛽 − 𝛾𝑎)
From (6) it is clear that if 𝑚 > 1 − 𝛾
𝑘(𝑐𝛽−𝛾𝑎)
− 𝑐𝛽
𝑘𝑎(𝑐𝛽−𝛾𝑎)
, then 𝐸2 is locally asymptotically stable.
Now if 𝑚 < 1 − 𝑘(𝑐𝛽−𝛾𝑎)
𝛾 𝑐𝛽
− 𝑘𝑎(𝑐𝛽−𝛾𝑎) , then 𝐸2 is locally unstable.
If 𝑚 = 1 − 𝑘(𝑐𝛽−𝛾𝑎)
𝛾 𝑐𝛽
− 𝑘𝑎(𝑐𝛽−𝛾𝑎) , then our system enters into Hopf type small amplitude periodic
solution near 𝐸2 .

Parameters 𝐸0 (0, 0) 𝐸1 (𝑘, 0) 𝐸2 (𝑥 ∗ , 𝑦 ∗ ) Phase portrait

𝑚 ∈ [0, 𝐴) Unstable Unstable Unstable, limit cycle exists Figs. 1 and 2
𝑚=𝐴 Unstable Unstable Unstable, small amplitude periodic solution occurs Fig. 3
𝑚 ∈ (𝐴, 𝐵) Unstable Unstable Asymptotically stable Fig. 4
𝑚=𝐵 Unstable Unstable Does not exist
𝑚 ∈ (𝐵, 1) Unstable Stable Does not exist Fig. 7

Table 3.1: Nature of equilibria of our model when 𝑐𝛽 > 𝛾𝑎.

In the above table,

𝛾 𝑐𝛽
𝐴=1− 𝑘(𝑐𝛽−𝛾𝑎)
− 𝑘𝑎(𝑐𝛽−𝛾𝑎)

𝐵 =1− 𝛾
𝑘(𝑐𝛽−𝛾𝑎)
.

7
Chapter 4

Numerical Simulation

Numerical Example
Let 𝛼 = 10, 𝑘 = 100, 𝑎 = 0.02, 𝛾 = 0.09, 𝛽 = 0.6, 𝑐 = 0.02 in appropriate units.For these
values of parameters, we verify the existence and stability properties of the equilibrium for
the system.
Here m as the bifurcation parameter, so we take different values of m.

Figure 4.1: The phase portrait of our model for m = 0.1.Here the equilibrium point (9.8,19.65)
is unstable.

8
Figure 4.2: The phase portrait of our model for m = 0.3.Here corresponding equilibrium point
(12.6,24.5) is also unstable.

Figure 4.3: Solution curves for m = 0.32.There is a periodic solution around the equilibrium
point 𝐸2 (13.0, 25.09).

9
Figure 4.4: Solution curves for m = 0.4. Both the populations converge to their equilibrium
values.

Figure 4.5: The phase portrait of our model for m = 0.5.𝐸2 (17.65, 32.3) is a global attractor.

10
Figure 4.6: Solution curves for m = 0.85. Both the populations converge to their equilibrium
values (58.82,53.82) .

Figure 4.7: The phase portrait of our model for m = 0.95.It is seen that (100,0) is globally
asymptotically stable.

11
Chapter 5

Conclusion

In this project first we have studied a prey-predator system incorporating a prey refuge. We
have assumed that the predator response function is of Holling type II. Here we have studied
impact of prey refuge in the dynamics of prey-predator model.

To study the dynamical behaviour of the model first we have calculated equilibrium point
of the model and we have observed that there are two boundary equilibrium points which
always exist and one interior (positive) equilibrium point exists with some constraint.

After that we have shown that all solutions of our model in 𝑅+2 are uniformly bounded.
Then we have studied the dynamical behaviour of the equilibrium points. Here we have
observed that 𝐸0 is always unstable (saddle). Equilibrium point 𝐸1 is conditionally asymp-
totically stable.

Since 𝐸1 is conditionally stable so when 𝐸1 is unstable then 𝐸2 will exist (by condition 5).
𝐸2 is also conditionally locally asymptotically stable.

Since all the dynamical behaviour of the equilibrium points depend on parameter m,
therefore m is the bifurcation parameter of our proposed model.

To see the impact of prey refuge in the consider model we have plotted phase portrait for
different value of prey refuge. At low level of prey refuge we have observed both population
coexist at oscillatory state that is stable limit cycle has observed.

Now we gradually increase our refuge parameter then system become stable at coex-
istence steady state. But further increasing refuge parameter we have seen that predator
population die out.

Thus, we have seen that high level of prey refuge is not beneficial for predator population.

Since many prey populations do have some type of refuge accessible, adding a refuge to
our system makes it more realistic.So it can be an important factor in the biological control
of a pest.For example, Hoy mentions that "hotspots" of high spider mite densities in almond

12
orchards can trigger orchard-wide outbreaks.These hotspots are places where the predator
is having trouble keeping the prey under control, and they can thus be regarded as refuges.

13
Bibliography

[1] Kar, T.K., 2005. Stability analysis of a prey–predator model incorporating a prey refuge.
Communications in Nonlinear Science and Numerical Simulation, 10(6), pp.681-691.

[2] Berryman, A.A., 1992. The orgins and evolution of predator-prey theory. Ecology, 73(5),
pp.1530-1535.

[3] Helle W, Sabelis MW, editors. Spider mites: their biology, natural enemies and control.
Amsterdam: Elsevier; 1985.

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