CHAPTER-2

MATHEMATICAL STUDY OF AN ECO-SYSTEM CONSISTENT OF A

PREY-PREDATOR WITH HOLLING TYPE –II FUNCTIONAL RE-
SPONSE AND HOST COMMENSAL TO THE PREY: EFFECT OF
PREDATION OF HOST BY PREDATOR.

2.1 INTRODUCTION :

Ecology is the study of living beings such as animals and plants in relation
to their habits and habitats. It mainly deals with the evolutionary biology,
which explains us about how the living beings are regulated in nature.
Ecology can refer to any form of biodiversity.Hence ecologists are capable
of conducting research from the smallest bacteria to the global atmospheric
gases. Ecology is a modern discipline of science, which came into exist-
ence in 19th century. It blossomed from natural science. Research in the
area of theoretical Ecology was initiated in1925 by Lotka [1] and in 1931
by Volterra [2]. Since then many Mathematicians and Ecologists contrib-
uted to the growth of this area of knowledge reported in the treatises of
May [3], Smith [4], Kushing[5], Kapur [6] etc.
The ecological interactions can be broadly classified as Prey-Predator,
Commensalism, Competition, Neutralism, Mutualism and so on. In com-
mensalisms one organism benefits the other without getting effected due to
the interaction (i.e. it is neither benefited nor harmed). A common example
is an animal using a tree for shelter-tree (Host) does not get any benefit
from the animal (Commensal). Recently, Archana Reddy[1] and Bhaskara
Rama Sharma[2] investigated diverse problems related to two species
competitive systems with time delay, employing analytical and numerical
 techniques. Following this Ravindra Reddy [12],[13],[14] et al. Shanker et
al [15] and Papa Rao [16] have discussed different prey-predator models in
detail. Recently Upadhyay et al. [23] studied the model system under the
influence of environmental driving forces. Seshagiri Rao and Pattabhi Ra-
macharyuly [12] studied stability Three Species Ecosystem Consisting of a
Prey, Predator and a Host Commensal to the Prey (with mortaility rate to
the prey).

This chapter is organized as follows:

Section 2.2 discusses the basic equations and notations. Section 2.3
obtain boundedness of the solution .Section 2.4 shows investigation of equilib-
rium states. Section 2.5 discuss stability of the equilibrium states. Nonlinear
stability of co-existing state is in section 2.6. Section 2.7 include the numerical
solution and section 2.8 conclusion is given.

2.2 BASIC EQUATIONS:

In the present model we will study the dynamics of a three species ecosystem
consisting of a prey (N1), a predator (N2) surviving on commensal species with
holling type II and a host (N3) commensal to the prey (with mortality rate for the
prey).
The model equations for a three species multi-reactive ecosystem are given by
the following system of non-linear ordinary differential equations.

dN1 pN2
= a11 N1 [−e1 − N1 − + cN3 ]
dt h+N1

(2.1)
dN2 qN1
= a22 N2 [K 2 − aN2 + ] + cαN2 N3 (2.2)
dt h+N1
dN3
= a33 N3 [K 3 − N3 ] − αN2 N3
dt

(2.3)

where,
d1 = The natural death/decay of N1.
a2 = natural growth rate of N2.
a3 = natural growth rate of N3.
aii = the rate of decrease of Ni due to insufficient resource of Ni ; i=1,2,3.
a12 = the rate of decrease of N1 due to inhibition by predator N2.
a13 = the rate of increase of the prey commensal (N1) due to its successful pro-
motion by the host (N3).
a21 = the rate of increase of the predator N2 due to its successful attacks on the
prey (N1).
e1: d1/a11 extinction coefficient of the prey (N1).
K2: a2/a22 is the carrying capacity of the predator (N2).
K3: a3/a33 is the carrying capacity of the host (N3).
p: a12/a11 coefficient of commensal inhibition .
q : a21/a22 coefficient of the predator consumption of the prey.
c = a13/a11 coefficient of the prey-commensal .
a = intraspecific co-efficient.
α = the predation of host by predator.
t*= the dominance reversal time.

2.3 BOUNDEDNESS OF THE SOLUTION:

Lemma: - All the solution of the system (2.1) to (2.3) that initiate in R 3* are
eventually bounded and enter into a region R1, defined by
R1 = {(N1 , N2 , N3 )𝜖 𝑅+3 : N1L < N1 ≤ cK 3 , 0 < N2 ≤ N2u , N3L ≤ N3 ≤ K 3 }
 a22 (K2 +qck3 )+cαk3
where,N1L = (𝑐 N3L − 𝑝 N2u − e1 ), N2u = , N3L =
a22 a
(a33 K3 − αN2u )
a33

Proof: We have from equation (2.3)

dN3
= a33 N3 (K 3 − N3 ) − αN2 N3
dt

≤ a33 N3 (K 3 − N3 )
lim N3 ≤ K 3
t→∞

dN3
= a33 N3 (K 3 − N3 ) − αN2 N3
dt
≥ (a33 K 3 − αN2u ) N3 − a33 N3 2 where N2u =
upper bound of N2
as t → ∞ we get
(a33 K 3 − αN2u )
N3 ≥
a33
Now from equation (2.1), we have
dN1 pN2
= a11 N1 [−e1 − N1 − + cN3 ]
dt h+N1

≤ a11 (cN3 N1 − N12)

≤ a11 (cK 3 N1 − N12 ) using N3 ≤ K 3
lim N1 ≤ cK 3
t→∞

dN1 pN2
= a11 N1 [−e1 − N1 − + cN3 ]
dt h + N1
≥ a11 N1 [ − e1 − N1 − 𝑝 N2u + 𝑐 N3L ]
= a11 N1 (𝑐 N3L − 𝑝 N2u − e1 ) − a11 N1 2
where N3L = lower bound of N3 , 𝑎𝑠 𝑡 → ∞ 𝑤𝑒 𝑔𝑒𝑡
N1 ≥ (𝑐 N3L − 𝑝 N2u − e1 )
Then from equation (2.2), we have
dN2 qN1
= a22 N2 [K 2 − aN2 + ] + cαN2 N3
dt h+N1

≤ a22 N2 [K 2 − aN2 + qN1 ] + cαN2 N3

 ≤ a22 N2 [K 2 − aN2 − qck 3 ] + cαN2 k 3
[using N3 ≤ K 3 and N1 ≤ cK 3 ]
= [(a22 (K 2 + qck 3 ) + cαk 3 ) N2 − a22 aN22 ]
a22 (K2 +qck3 )+cαk3
lim N2 ≤
t→∞ a22 a

2.4 EQUILIBRIUM STATES:

dNi
The equilibrium points are given by = 0, i = 1,2,3. the system under inves-
dt

tigation has six equilibrium points that can put in four categories A, B, C, D as
follows.
Fully washed out state.
E0 : ̅̅
N̅̅1 = 0; ̅̅̅̅
N2 = 0; ̅̅̅̅
N3 = 0 (2.4)
States in which two of three species are washed out and third is not.
E1 : ̅̅
N̅̅1 = 0; ̅̅̅̅
N2 = 0; ̅̅̅̅
N3 = k 3 (2.5)
K
E2 : ̅̅
N̅̅1 = 0; ̅̅̅̅
N2 = 2 ; ̅̅̅̅
N3 = 0 (2.6)
a

Only one of the three species is washed out while the other two are not.
𝜇1 𝜇2
E3 ∶ ̅̅
N̅̅1 = 0; ̅̅̅̅
N2 = ; ̅̅̅̅
N3 = (2.7)
𝛼`1 𝛼`1

Where 𝜇1 =a33 (a22 K 2 + cαK 3 ), 𝜇2 =a22 (aa33 K 3 − αK 2 )

𝛼`1 =(aa22 a33 + cα2 )
This would exists only when aa33 K 3 > αK 2 .
E4 ∶ ̅̅
N̅̅1 = cK 3 − e1 ; ̅̅̅̅
N2 = 0; ̅̅̅̅
N3 = K 3 (2.8)
This would exists only when cK 3 > e1
The Co-existence state or normal steady state.
E5 ∶ ̅̅
N̅̅1 = N ∗1 , ; ̅̅̅̅
N2 = N ∗ 2 ; ̅̅̅̅
N3 = N ∗ 3 (2.9)
N ∗1 𝟑 + N ∗1 𝟐 𝜎1 + N ∗1 σ2 – σ3 = 0

∗
a33 h(cK 3 − e1 ) − a33 N ∗1 2 − a33 N ∗1 (e1 + h − cK 3 )
N 2 =
a33 p + chα + cαN ∗1
 a33 K3 − αN∗ 2
N∗3 =
a33
𝑐𝛼(K2 a22 + cαK3 + a22 q)
where , 𝜎1 = [ (aa22 a33 +cα2 )
+ e1 + 2h − cK 3 ]

𝜎2 = h(2e1 + h - 2cK 3 )
(a22 K2 +cαK3 )(a33 p+chα)
𝜎3 = h [ h(ck 3 − e1 ) − (aa22 a33 +cα2 )
]

This would exists only when the following conditions are satisfied
a. cK 3 > e1
h
b. e1 + > 𝑐K 3
2
(a22 K2 +cαK3 )(a33 p+chα)
c. h(cK 3 − e1 ) > (aa22 a33 +cα2 )

d. a33 K 3 > 𝛼N ∗ 2

2.5 THE STABILITY OF THE EQUILIBRIUM STATES:

The basic equations (2.1), (2.2) and (2.3) are linearized to obtain the equations
for perturbed state.
du
= AU, (2.10)
dt

where,
̅̅̅2̅h
pN −a11 pN ̅̅̅1̅
̅̅̅1̅ −
a11 [−e1 − 2N ̅̅̅3̅]
+ cN ca11 ̅N̅̅1̅
(h + ̅N̅̅1̅)2 (h + N1̅)
̅̅̅
A= a 22̅N̅̅2̅qh qN̅̅̅1̅
̅̅̅2̅ +
a 22 [{K 2 − 2aN ̅̅̅3̅]
} + cαN ̅̅̅2̅
cαN
(h + ̅N̅̅1̅)2 (h + ̅N̅̅1̅)
[ 0 −αN̅̅̅3̅ a 33 (K 3 − 2N̅̅̅3̅) − αN
̅̅̅2̅]

2.5.1 FULLY WASHED OUT EQUILIBRIUM STATE:

In this state the corresponding linearized perturbed equations are
du1
= − a11 e1 u1
dt
du2
= a22 K 2 u2
dt
du3
= a33 K 3 u3 (2.10)
dt

The characteristics roots of the system (2.10) are

λ1 = −a11 e1 , λ2 = a22 K 2 , λ3 = a33 K 3
Of these λ2 and λ3 are positive, so the state is unstable.
The equations (2.10) yield the solution curves
u1 = u10 e−a11e1t ; u2 = u20 ea22K2t ; u3 = u30 ea33K3t (2.11)
Case 1A: u10 > u20 > u30 ; a33 K 3 > a22 K 2
In spite of the low initial strength of host, it outnumbers the prey and predator at
the time.
1 u10 1 u20
t13 ∗ = log ( ) and t 23 ∗ = log ( )
a33 K 3 + a11 e1 u30 a33 K 3 − a22 K 2 u30
1 u10
Further the predator out-numbers the prey at time t12 ∗ = log ( ).
a11 e1 +a22 K2 u20

This happens due to a higher natural growth rate of host compared to that of
predator. This is illustrated in fig.1.

Case 1B: u10 > u30 > u20 ; a33 K 3 > a22 K 2
The host and predator both dominates the prey at times t13 ∗ and t12 ∗ as shown
in fig. 2.
Case 1C: u10 > u30 > u20 ; a22 K 2 > a33 K 3
The predator out numbers both the prey the host at time t12 ∗ and t 32 ∗ =
1 u30
(a22 K2 −a33 K3 )
log (
u20
). Further the host out numbers the prey at time t13 ∗ after

this illustrated in Fig. 3

2.5.2 PREY AND PREDATOR WASHED OUT STATE:
In this case, we get from (2.1), (2.2) and (2.3)
du1
= a11 (cK 3− e1 )u1
dt
du2
= (a22 K 2 + cαK 3 )u2
dt
du3
= −a33 K 3 u3 − αK 3 u2 (2.12)
dt

The characteristics roots of (3.12) are

λ1 = cK 3− e1 ; λ2 = a22 K 2 + cαK 3 ; λ3 = −a33 K 3
Of these, λ2 is always positive, so the state is unstable. The equations (2.12)
yield the solution curves.
u1 = u10 eλ1t ; u2 = u20 eλ2t ; u3 = (u30 + L1 )eλ3t − L1 eλ2t (2.13)
where L1 = αK 3 u20 /(λ2 − λ3 ).
Case A:- cK 3 − e1 > 0
Then first two roots are positive and third is negative.
Case 2A (i): u10 > u30 > u20 and λ1 > λ2 ;
The prey dominates the predator because of higher natural growth rate and ini-
tial take off, the predator out numbers the host at time t 32 ∗ as shown in fig.4.

Case 2A (ii):- u10 > u20 > u30 and λ1 < λ2 ;

The prey dominates the predator till the time t12 ∗ after that the predator domi-
nates, and the host asymptotic to the equilibrium point as shown in fig.5.
Case B: cK 3 − e1 < 0
In this case, λ2 is positive and the remaining two roots are negative.
Case 2B (i): u10 > u20 > u30
The commensal-prey out number the predator till the time t12 ∗ after that the
predator dominates , and the prey-commensal and host declines further as
shown in Fig.6.
Case 2B (ii): u30 > u10 > u20
Even with the low initial strength of the predator, it out numbers the commensal
prey and later host at time t12 ∗ and t 32 ∗ as shown in Fig.7.
2.5.3 PREY COMMENSAL AND HOST WASHED OUT STATE:
In this case, we get from (2.1), (2.2) and (2.3)
du1 pK2
= −a11 (e1 + ) u1
dt ah
du2 a22 K2 q cαK2
= u1 − a22 K 2 u2 + u3
dt ah a
du3 αK2
= [a33 K 3 − ] u3 (2.14)
dt a

The corresponding characteristics roots are

pK2 αK2
λ1 = −a11 (e1 + ) ; λ2 = −a22 K 2 , λ3 = [a33 K 3 − ]
ah a
𝛼𝐾2
Case 3A:- 𝑎33 𝐾3 >
𝑎
 αK2
When a33 K 3 > , third root is positive, so this state is unstable.
a

The equations (2.14) yield the solutions curves

𝑢1 = 𝑢10 𝑒 𝜆1𝑡 ; 𝑢3 = 𝑢30 𝑒 𝜆2𝑡
𝜆 𝑞𝑢10 𝑐𝛼𝐾 𝑢 𝜆 𝑞𝑢10 𝑐𝛼𝐾 𝑢
2
𝑢2 = [𝑢20 + (𝑎ℎ(𝜆 − 𝑎(𝜆 2−𝜆30))] 𝑒 𝜆2 𝑡 − (𝑎ℎ(𝜆
2
𝑒 𝜆1 𝑡 − 𝑎(𝜆 2−𝜆30) 𝑒 𝜆3 𝑡 ) (2.15)
1 −𝜆2 ) 3 2 1 −𝜆2 ) 3 2

𝜆2 𝑞𝑢10 𝑐𝛼𝐾2 𝑢30

When 𝑢20 = − + the equations (2.15) become
𝑎ℎ(𝜆1 −𝜆2 ) 𝑎(𝜆3 −𝜆2 )

𝜆2 𝑞𝑢10 𝑐𝛼𝐾2 𝑢30 𝜆 𝑡

𝑢1 = 𝑢10 𝑒 𝜆1 𝑡 , 𝑢2 = − ( 𝑒 𝜆1 𝑡 − 𝑒 3 )
𝑎ℎ(𝜆1 − 𝜆2 ) 𝑎(𝜆3 − 𝜆2 )
𝑢3 = u30 𝑒 𝜆3𝑡
𝜆2 𝑞𝑢10 𝑐∝𝑘2 𝑢30
Let y1 = − , y2 = −
𝑎ℎ(𝜆1 −𝜆2 ) 𝑎(𝜆3 −𝜆2 )

Then we have-
u1 =u10 eλ1t , u2 = u20 eλ1t +y2(eλ1t -eλ3t ) ;u3 =u30 eλ3t (2.16)
Case 3A (i) : u30>u20>u10
In this case the host exceeds both the predator and prey. Further, both the preda-
tor and the host species move towards equilibrium, which is shown in fig. 8.
Case 3A (ii): u10>u30>u20
∗ 1 u10
In this case, the prey commensal out number the host at time 𝑡13 = log
𝜆3 −𝜆1 u30

after that host dominates, and the prey commensal declines. Further as shown in
fig. 9.
∝k2
Case 3B : a33 k3 <
𝑎

In this case, all the three characteristics roots are negative. So this state is stable.
Case 3B (i) : u10>u30>u20
In this case all the three species move towards the equilibrium point asymptoti-
cally, as shown in fig. 10.

2.5.4 PREY WASHED OUT STATE:

In this case, we get from (2.1), (2.2) and (2.3)
𝑑𝑢1 𝑝µ1 𝑐µ2
= 𝑎11 (−𝑒1 − + )𝑢1
𝑑𝑡 ∝1 ℎ ∝1
 𝑑𝑢2 𝑎22 µ1 𝑞 2𝑎µ1 𝑐∝µ2 𝑐∝µ1
=( ) 𝑢1 + 𝑎22 (𝑘2 − + ) 𝑢2 + 𝑢3
𝑑𝑡 ∝1 ℎ ∝1 ∝2 ∝1
𝑑𝑢3 ∝µ2 2µ2 ∝µ1
=− 𝑢2 + (𝑎33 (𝑘3 − )− )𝑢3 (2.20)
𝑑𝑡 ∝1 ∝1 ∝1

Characteristic equation corresponding to the above equations is

𝑝µ1 𝑐µ2
(𝜆 − 𝑎11 (−𝑒1 − + ))(𝜆2
∝1 ℎ ∝1
2𝑎22 𝑎𝜇1 ∝ 𝜇1 2𝑎33 𝜇2 𝑎22 𝑐 ∝ 𝜇2
+( + + − 𝑎22 𝑘2 −
∝1 ∝1 ∝1 ∝1
4𝑎22 𝑎33 𝑎𝜇1 𝜇2 2𝑎22 𝑎𝜇12
− 𝑎33 𝑘3 ) 𝜆 + 𝑎22 𝑎33 𝑘2 𝑘3 + +
∝1 2 ∝12
2𝑎22 𝑎33 𝑘2 𝜇2 𝑎22 𝑘2 ∝ 𝜇1 2𝑎22 𝑘3 𝑎33 𝑎𝜇1
− − − − 2𝑐
∝1 ∝1 ∝1
𝑎22 𝑎33 𝑐 ∝ 𝑎22 𝑎33 𝑘3 µ2
∝ 𝜇22 + )=0
∝12 ∝1
𝑝µ1 𝑐µ2
Where 𝜆 = − 𝑎11 (𝑒1 + − )
∝1 ℎ ∝1

It is easy to find condition for 𝜆1 < 0 and by the following equation.

2𝑎22 𝑎𝜇1 ∝ 𝜇1 2𝑎33 𝜇1 𝑎22 𝑐 ∝ 𝜇2
𝜆2 + ( + + − 𝑎22 𝑘2 − − 𝑎33 𝑘3 ) 𝜆
∝1 ∝1 ∝1 ∝1
4𝑎22 𝑎33 𝑎𝜇1 𝜇2 2𝑎22 𝑎𝜇12 𝑐 ∝ 𝑎22 𝑎33 𝑘3 µ2
+ 𝑎22 𝑎33 𝑘2 𝑘3 + + + )
∝1 2 ∝12 ∝1
2𝑎22 𝑎33 𝑘2 𝜇2 𝑎22 𝑘2 ∝ 𝜇1 2𝑎22 𝑘3 𝑎33 𝑎𝜇1
− − − − 2𝑐
∝1 ∝1 ∝1
𝑎22 𝑎33
∝ 𝜇22 =0
∝12
Writing above equation as
𝜆2 + 𝑏1 𝜆 + 𝑏2 = 0
 𝑏1 ±√𝑏12 −4𝑏2
where 𝜆 = −
2

(2.21)
2𝑎22 𝑎𝜇1 ∝ 𝜇1 2𝑎33 𝜇2 𝑎22 𝑐 ∝ 𝜇2
𝑏1 = + + − 𝑎22 𝑘2 − − 𝑎33 𝑘3
∝1 ∝1 ∝1 ∝1
4𝑎22 𝑎33 𝑎𝜇1 𝜇2 2𝑎22 𝑎𝜇12 𝑐 ∝ 𝑎22 𝑎33 𝑘3 µ2
𝑏2 = 𝑎22 𝑎33 𝑘2 𝑘3 + + + )
∝12 ∝12 ∝1
2𝑎22 𝑎33 𝑘2 𝜇2 𝑎22 𝑘2 ∝ 𝜇1 2𝑘3 𝑎22 𝑎33 𝑎𝜇1
− − − − 2𝑐
∝1 ∝1 ∝1
𝑎22 𝑎33
∝ 𝜇22
∝12
using the Hurwitz criteria we derive that the equilibrium point E4 locally
asymptotically stable if
𝑝𝜇1 𝑐𝜇1
b1 > 0 and b2 >0 and (𝑒1 + )>
∝1 ℎ ∝1

Here the conditions are

𝑎22 𝑎𝜇1 ∝ 𝜇1 2𝑎33 𝜇2 𝑎22 𝑐 ∝ 𝜇2
2 + + > 𝑎22 𝑘2 + + 𝑎33 𝑘3
∝ ∝1 ∝1 ∝1
and
𝑐∝𝜇2 2𝑎𝑎22 𝜇1 𝑎22 𝑎33
𝑎22 𝑎33 𝑘3 (𝑘2 + )+ (2𝜇2 + 𝜇1 ) > (2𝑘2 𝜇2 +
∝1 ∝21 ∝1

2𝑐∝𝜇22 𝑎22 𝑘2 ∝𝜇1

2𝑎𝜇1 𝑘3 + )+
∝1 ∝1

The equations (2.20) yield the solution curves

𝑢1 = 𝑢10 𝑒 𝜆1𝑡
 𝜇1 𝑞𝑢10 (𝜆1 − 𝑐)
u2 = 𝑎22 ( ) 𝑒 𝜆1𝑡
∝1 ℎ(𝜆3 − 𝜆1 )(𝜆2 − 𝜆1 )
𝑎22 𝜇1 𝑞𝑢10 (𝜆2 − 𝑐) 𝑢20 (𝑐 − 𝜆2 )
+{ +
∝1 ℎ(𝜆3 − 𝜆1 ) (𝜆2 − 𝜆1 ) (𝜆3 − 𝜆1 )
𝑢30 𝑐 ∝ 𝜇1
+ } 𝑒 𝜆2𝑡
∝ (𝜆2 − 𝜆3 )
𝑎22 𝜇1 𝑞𝑢10 (𝜆3 − 𝑐) 𝑢20 (𝜆3 − 𝑐)
+{ +
∝1 ℎ(𝜆3 − 𝜆1 ) (𝜆2 − 𝜆1 ) (𝜆3 − 𝜆2 )
𝑢30 𝑐 ∝ 𝜇1
− } 𝑒 𝜆3𝑡
(𝜆
∝ 2 − 𝜆3 )
𝑎22 𝜇1 𝑞𝜇 𝜇30(𝜆2 −𝐵) ∝𝜇2 𝑢20
10 𝑒𝜆1 𝑡
𝑢3 (𝑡) = + { − −
∝1 ℎ(𝜆3 −𝜆1 )(𝜆2 −𝜆1 ) (𝜆2 −𝜆3 ) ∝1 (𝜆2 −𝜆3 )
∝𝑎22 𝜇1 𝑞𝑢10 𝜇2 (𝐵−𝜆3 )𝑢30 ∝𝜇2 𝑢20 𝑎22 𝜇1 𝑞𝑢10
2
∝1 ℎ(𝜆1 −𝜆2 )(𝜆3 −𝜆2 )
} 𝑒 𝜆2𝑡 + { (𝜆2 −𝜆3 )
−
∝1 (𝜆3 −𝜆2 )
−
∝1 ℎ(𝜆1 −𝜆3 )(𝜆2 −𝜆3 )
} 𝑒 𝜆3𝑡

where
2𝑎𝜇1 𝑐∝𝜇2
B = 𝑎22 (𝑘2 − +
∝1 ∝1
2𝜇2 ∝𝜇1
C= 𝑎33 (𝑘3 − )+ )
∝1 ∝1

2.5.5. PREDATOR WASHED OUT STATE:

In this case we get from (2.1), (2.2) and (2.3)
𝑑𝑢1 𝑎11 𝑝(𝑐𝑘3 −𝑒1 )
= − 𝑎11 (𝑐𝑘3 − 𝑒1 )𝑢1 − 𝑢2 + 𝑐𝑎11 (𝑐𝑘3 − 𝑒1 )𝑢3
𝑑𝑡 ℎ+(𝑐𝑘3 −𝑒1 )
𝑑𝑢2 𝑞(𝑐𝑘3 −𝑒1 )
= { 𝑎22 (𝑘2 + + 𝑐 ∝ 𝑘3 }𝑢2
𝑑𝑡 ℎ+(𝑐𝑘3 −𝑒1 )
𝑑𝑢3
= − ∝ 𝑘3 𝑢2 − 𝑎33 𝑘3 𝑢3
𝑑𝑡

(2.17)
The characteristic roots of (2.17) are
 𝑞(𝑐𝑘3 −𝑒1 )
𝜆1 = − 𝑎11 (𝑐𝑘3 − 𝑒1 ), 𝜆2 = 𝑎22 (𝑘2 + } + 𝑐 ∝ 𝑘3 , 𝜆3 =
(ℎ+(𝑐𝑘3 −𝑒1 )

−𝑎33 𝑘3 .
Of these, , 𝜆2 𝑖𝑠 always positive, so the state is unstable. The equations (2.17)
yield the solution curves.
1 𝜆1 𝑝𝑢20
𝑢1 = (𝑢10 − 𝐿3 )𝑒 𝜆1𝑡 + (𝑐𝜆1 𝐿2 − )𝑒 𝜆2𝑡 −
𝜆2 −𝜆1 (ℎ+(𝑐𝑘3 −𝑒1 )

(𝑢30 + 𝐿2 )𝑐𝜆1 𝑒 𝜆3𝑡

𝜆3 −𝜆1

𝑢2 = 𝑢20 𝑒 𝜆2𝑡
𝑢3 = (𝑢30 + 𝐿2 𝑒 𝜆3𝑡 − 𝐿2 𝑒 𝜆3𝑡 ) (2.18)
∝𝐾3 𝑢20
where L2 =
𝜆2 −𝜆3
𝐿2 𝑐 𝑝𝑢20 𝑐(𝑢30 +𝐿2 )
L3 = 𝜆1 ( − + ]
𝜆2 −𝜆1 (ℎ+(𝑐𝑘3 −𝑒1 ))(𝜆2 −𝜆1 ) 𝜆2 −𝜆3

When 𝑢10 = 𝐿3, then equation (2.18) become

(𝑢30 + 𝐿2 )𝑐𝜆1
𝑢1 = 𝑢10 e λ2t + (e λ2t − eλ3t )
𝜆3 −𝜆1

u2 = u20 e λ2t ; 𝑢3 = (u30 + L2 )eλ3t − L2 e λ2t

Case 4A (i) u10>u20>u30

In this case, the prey species and predator species go away from the equilibrium
point while, the third species move towards the equilibrium point as shown in
Fig. 11.
Case 4A (ii): u30>u20>u10
In this case, the host species dominates the predator species and prey commen-
∗ ∗
sal species till the time instant 𝑡32 and 𝑡31 respectively, which is shown in Fig.
12
2.5.6 CO-EXISTENT STATE:-
In this case from (2.9) we have
𝐴

∗ 𝑝𝑁2 ∗ ℎ ∗ ∗ −𝑎11 𝑝 ∗
𝑎11 [−𝑒1 − 2𝑁1 − 2 + 𝑐𝑁3 ] ∗ 𝑁1
(ℎ + 𝑁1 ∗ )
∗ (ℎ + 𝑁1 )
= 𝑎22 𝑞ℎ𝑁2 ∗ 𝑞 ∗ ∗∗
𝑎22 [𝐾2 + ∗ 𝑁1 − 2𝑎𝑁2 ]
(ℎ + 𝑁1 ∗ )2 (ℎ + 𝑁1 )
[ 0 −𝛼𝑁3 ∗ 𝑎33 (𝐾3
Using (2.1)-(2.3.), above Jacobean Matrix converts to
𝐵11 −𝐵12 𝐶𝑎11 𝑁1 ∗
𝐴 = [𝐵21 −𝑎𝑎22 𝑁2 ∗ 𝐶𝛼𝑁2 ∗ ]
0 −𝛼𝑁3 ∗ −𝑎33 𝑁3 ∗

𝑝𝑁 ∗ 𝑁2 ∗
1 ∗ 𝑎11 𝑝𝑁1 ∗ 𝑎22 𝑞ℎ𝑁2 ∗
where 𝐵11 = 𝑎11 ((ℎ+𝑁 ∗ )2 − 𝑁1 ), 𝐵12 = (ℎ+𝑁1 ∗ ) , 𝐵21 = (ℎ+𝑁1 ∗ )2
1

Characteristics equation corresponding to the above Jacobean is –

𝜆3 + 𝜆2 (𝑎𝑎22 𝑁2 ∗ + 𝑎33 𝑁3 ∗ − 𝐵11 ) + 𝜆{(𝑎22 𝑎𝑎33 + 𝐶𝛼 2 )𝑁2 ∗ 𝑁3 ∗ −
𝐵11 (𝑎𝑎22 + 𝑎33 )𝑁3 ∗ + 𝐵12 𝐵21 } + 𝐵21 𝑁3 ∗ (𝑎𝑎22 + 𝑎33 )𝑁3 ∗ −
𝐵11 (𝑎𝑎22 𝑎33 + 𝐶𝛼 2 )𝑁2 ∗ 𝑁3 ∗ = 0
Writing above equation as
𝜆3 + 𝑏1 𝜆2 + 𝑏2 𝜆 + 𝑏3 = 0 (2.22)
Where,
𝑏1 = 𝑎𝑎22 𝑁2 ∗ + 𝑎33 𝑁3 ∗ − 𝐵11
𝑏2 = (𝑎𝑎22 𝑎33 + 𝐶𝛼 2 )𝑁2 ∗ 𝑁3 ∗ − 𝐵11 (𝑎𝑎22 + 𝑎33 )𝑁3 ∗ + 𝐵12 𝐵21
𝑏3 = 𝐵21 𝑁3 ∗ (𝐵12 𝑎33 + 𝑐𝑎11 𝛼𝑁1 ∗ ) − 𝐵11 (𝑎𝑎22 𝑎33 + 𝐶𝛼 2 )𝑁2 ∗ 𝑁3 ∗
 Using the Routh-Hurwitz criteria, the condition for the stationary point
to be locally asymptotically stable are b1>0, b2>0, b3>0 and b1b2-b3>0.

2.6 NON LINEAR STABILITY OF THE EQUILIBRIUM POINT E5:-

∗
̅̅̅2 ∗ = 𝑁2 ∗ ; ̅𝑁
𝑁1 = 𝑁1 ∗ ; ̅𝑁
𝐸5 ∶ ̅̅̅ ̅̅̅3 ∗ = 𝑁3 ∗ (2.23)
We define a Liapunov’s Function
𝑁1
V = (𝑁1 , 𝑁2 , 𝑁3 ) = 𝑁1 − ̅̅̅
𝑁1 − ̅̅̅
𝑁1 log ( )
̅̅̅
𝑁1
𝑁 𝑁
+ 𝑙1 {𝑁2 − ̅𝑁
̅̅̅2 − ̅𝑁
̅̅̅2 log ( 2 )} + 𝑙2 {𝑁3 − ̅𝑁
̅̅̅3 − ̅𝑁
̅̅̅3 log ( 3 )}
̅𝑁
̅̅̅2 ̅̅̅̅3
𝑁
Where l1 and l2 are suitable constants to be determined in subsequent steps.
Its time derivative is,
𝑑𝑉 𝑁1 − ̅̅̅
𝑁1 𝑑𝑁1 𝑁2 − ̅𝑁
̅̅̅2 𝑑𝑁2 𝑁3 − ̅𝑁
̅̅̅3 𝑑𝑁3
=( ) + 𝑙1 ( ) + 𝑙2 ( )
𝑑𝑡 𝑁1 𝑑𝑡 𝑁2 𝑑𝑡 𝑁3 𝑑𝑡
Put 𝑁1 − ̅̅̅
𝑁1 = 𝑧1 ; 𝑁2 − ̅𝑁
̅̅̅2 = 𝑧2 ; 𝑁3 − ̅𝑁
̅̅̅3 = 𝑧3 and using the equa-
tion (2.1), (2.2), (2.3).
 𝑝𝑁2
= 𝑧1 {𝑎11 (−𝑒1 − 𝑁1 − + 𝐶𝑁3 )}
ℎ + 𝑁1
𝑞𝑁1
+ 𝑙1 𝑧2 {𝑎22 (𝐾2 − 𝑎𝑁2 + ) + 𝐶𝛼𝑁3 }
(ℎ + 𝑁1 )
+ 𝑙2 𝑧3 {𝑎33 (𝐾3 − 𝑁3 ) − 𝛼𝑁2 }
𝑑𝑁̅1 𝑝𝑁̅2
𝑆𝑖𝑛𝑐𝑒 = 0 ⇒ −𝑒1 − ̅̅̅
𝑁1 − + 𝐶𝑁 ̅3 = 0
𝑑𝑡 (ℎ + 𝑁1 )
𝑝𝑁̅2
̅̅̅
⇒ −𝑒1 = 𝑁1 + − 𝐶𝑁̅3 ,
(ℎ + 𝑁1 )
𝑑𝑁̅2 𝑑𝑁̅3
[ 𝑠𝑎𝑚𝑒 𝑎𝑠 𝑎𝑏𝑜𝑣𝑒 𝑢𝑠𝑖𝑛𝑔 = 0 𝑎𝑛𝑑 =0 ]
𝑑𝑡 𝑑𝑡
Then,
d𝑉 pN̅2 pN2
̅1 +
= z1 {a11 (N ̅ 3 − N1 −
− CN + CN3 )}
dt ̅1 )
(h + N (h + N1 )
qN̅1 CαN̅3 qN1
̅2 −
+ l1 z2 {a22 (aN − − aN2 + )
̅1)
(h + N a22 (h + N1 )
αN̅2
+ CαN3 } + l2 z3 {a33 ( ̅ 3 − N3 ) − αN2 }
+N
a33
a11 pN̅ 2 z1 2 a11 p
2
= −a11 z1 + a11 cz1 z3 + − z z
(h + N1 )(h + N ̅ 1 ) (h + N1 ) 1 2
l1 a22 qhz1 z2
− l1 a22 az2 2 + l1 cαz2 z3 + − l αz z
(h + N1 )(h + N ̅1 ) 2 2 3
− l2 a33 az3 2
a11 p(h + N̅1 )
choosing l1 =
a22 qh
a11 pN̅ 2 z1 2
2
= −a11 z1 + a11 cz1 z3 + − l a az 2 + l1 cαz2 z3
(h + N1 )(h + N ̅ 1 ) 1 22 2
− l2 cαz2 z3 − l2 a33 az3 2
Now choosing l1 = l2 , we have
a11 pN̅ 2 z1 2
2
= −a11 z1 + a11 cz1 z3 + − l1 a22 az2 2 − l1 a33 az3 2
̅
(h + N1 )(h + N1 )
 ̅ 2 z1 2
a11 pN
2
≤ − a11 z1 + a11 cz1 z3 + − l1 a22 az2 2 − l1 a33 az3 2
̅
(h + N1L )(h + N1 )
dV
= −( 𝑃11 z1 2 − 𝑃13 z1 z3 + 𝑃33 z3 2 + 𝑃22 z2 2 )
dt
̅̅̅̅
pN 2
Where, P11 = a11 (1 − (h+N ̅ 1)
) , P22 = l1 aa22
1L )(h+N

̅ 1)
a11 p(h+N
P33 = l1 a33 , P13 = ca11 , l1 =
a22 qh

Now, if 𝑃11 > 0, 𝑃22 > 0, 𝑃33 > 0, 𝑃13 > 0,. Then the sufficient condi-
𝑑𝑉
tions for to be negative definite 𝑃13 2 < 4𝑃11 . 𝑃33 .
𝑑𝑡

Hence, V is a Liapunov’s Function with respect to equilibrium point E5.

Thus, it concludes that the equilibrium point E5 is non-linearly asymp-
totically stable.

2.7 NUMERICAL SIMULATION:

 Regarding equilibrium point E6(N1*, N2*,𝐍𝟑 ∗ ) :

Consider the following sets of parameters :

a1 =1, e1= 0.5, p= 0.5, h= 5, c = 2, a2 = 2, K2 = 1, a = 2, q = 0.8, alpha = 0 .3,
a3 = 3, K3 = 1.
For this numerical example, we get that the conditions for existence of interior
equilibrium point E6(N1*, N2*,N3 ∗ ) are satisfied and E6 is given by
E6 = {1.37211,0.45926 ,0.95407}
and the following region of attraction:
R1 = {(N1 , N2 , N3 ) ϵ R3+ : 0.485 ≤ N1 ≤ 2, 0 ≤ N2 ≤ 1.45, 0.855 ≤ N3
≤ 1}
With the above set of parameters we have observed that this numerical
example satisfies the Liapunov’s non-linear stability condition for global
stability.
In figure (3.1) ,we can also observed that interior equilibrium point E6(N1*,
N2*,N3 ∗ ) is asymptotically stable.

Now for α1 = 0.1 we obtained that

E6 = {1.44169,0.40424 ,0.98653}
and the following region of attraction:
R1 = {(N1 , N2 , N3 ) ϵ R3+ : 0.735 ≤ N1 ≤ 2, 0 ≤ N2 ≤ 1.35, 0.955 ≤ N3
≤ 1}

Now for α1 = 0.2 we obtained that

E6 = {1.40895,0.4308 ,0.97128}
and the following region of attraction:
R1 = {(N1 , N2 , N3 ) ϵ R3+ : 0.61334 ≤ N1 ≤ 2, 0 ≤ N2
≤ 1.4, 0.90667 ≤ N3 ≤ 1}

Now for α1 = 0.4 we obtained that

E6 = {1.33145,0.48765 ,0.93498}
and the following region of attraction:
R1 = {(N1 , N2 , N3 ) ϵ R3+ : 0.35 ≤ N1 ≤ 2, 0 ≤ N2 ≤ 1.5, 0.8 ≤ N3 ≤ 1}

𝐊𝟐
Regarding equilibrium point E2(0, , 𝟎):
𝐚

a1 =.1, e1= 0.5, p= 0.5, h= 5, c = 2, a2 = .2, K2 = 3, a = 2, q = 0.8, 𝛼1 = 0 .9,

a3 = 2.5, K3 = .5.

𝝁𝟏 𝝁𝟏
Regarding equilibrium point E3(𝟎, , ):
𝜶`𝟏 𝜶`𝟏
a1 =1, e1= 0.5, p= 1, h= 5, c = 2, a2 = 2, K2 = 2, a = 2, q = 0.8, alpha = 0 .9,
a3 = 2.5, K3 = .5.

3
N1
N2
N3
2.5

1.5

0.5
0 10 20 30 40 50 60 70 80 90 100
Time
fig.2.1
 3

2.8

2.6

2.4

2.2

1.8

1.6

1.4

1.2

1
0 10 20 30 40 50 60 70 80 90 100
Time
fig.2.2
 3

2.5

1.5

0.5

0
0 10 20 30 40 50 60 70 80 90 100
Time
fig.2.3

1.8
alpha=.1
1.7 alpha=.2
alpha =.3
alpha=.4
1.6

1.5
N1

1.4

1.3

1.2

1.1

1
0 10 20 30 40 50 60 70 80 90 100
Time
fig.2.4
 2
alpha=.1
alpha=.2
1.8
alpha =.3
alpha=.4
1.6

1.4
N2

1.2

0.8

0 10 20 30 40 50 60 70 80 90 100
Time
fig.2.5
 3
alpha=.4
alpha=.3
alpha =.2
2.5
alpha=.1

N3

1.5

0.5
0 10 20 30 40 50 60 70 80 90 100
Time
fig.2.6
2.8 CONCLUSION
In this chapter, a three species ecosystem is considered. The interactions that
are considered are Prey-Predator with Holling type II functional response,
Commensalism. In all, six equilibrium points are identified out of which one is
trivial.
The trivial equilibrium state and prey and predator-commensal washed out state
are found locally asymptotically unstable while other four found to be locally
asymptotically stable under some conditions involving parameters. The co-exist
state is identified nonlinearly asymptotically stable by using the Liapunov’s
function method.
Numerical example is given to support the theoretical results.