PATTERN FORMATION IN A PMZC PLANKTON

MODEL
Tahani Al-Karkhi, Hadi Susanto, Edward A. Codling
Department of Mathematical Sciences, University of Essex, UK
Wivenhoe Park, Colchester, CO4 3SQ, United Kingdom

Abstract

We focus on the phenomena of pattern formation in a reaction diffusion

model of plankton dynamics that includes infochemical mediated trophic
interactions. We consider an extended four species model. Where the graz-
ing pressure of microzooplankton (M) on phytoplankton (P) is controlled
through external info-chemical (C) mediated predation by copepods (Z). We
undertake a stability analysis of both the two species and four species models
and compare the system dynamics. We also explore spatial pattern forma-
tion in both models and consider under what conditions a Turing pattern is
exhibited or when spatiotemporal chaos is observed. An attempts are made
to study the non-Turing patterns with special emphasis on spatiotemporal
chaos.
Keywords: , Plankton model, Asymptotic Expansion Analysis, Limit cycle,
Dynamical system,, Chaos, Systematic analysis.

1 1. Introduction

2 Turing and non-Turing Patterns ? was used by many researchers to un-

3 derstand the mechanism of pattern formation in various physical processes

Preprint submitted to MMNP October 2, 2017

 4 such as formation meandering of a river, mixing of chemical reagents or
5 chemical processes, ripple like structures in the sand. In this work we are
6 introducing a trophic web illustrated in Figure ??, where the direction of the
7 interaction is given by the arrows. There are four interacting components,
8 namely the Phytoplankton (P ), Microzooplankton (M ), Zooplankton (Z)
9 and (C) is the infochemical released as shown in Figure ??. The interactions
10 in the diagram are described by the system of four differential equations,
11 below is the fill description of the PMZC-model. Mostly we focus on spa-
12 tiotemporal system and provided an analytical and numerical explanation
13 for understanding the diffusion driven instability condition. attempts are
14 made to study the non-Turing patterns with special emphasis on spatiotem-
15 poral chaos, which is able to explain the irregular distribution of population
16 observed frequently in nature.

∂U
= Di ∇2 U + Fi (U ) (1)
∂t
17 Where U represents a vector of four components: P denotes to the pop-
18 ulation density of prey Phytoplankton, M denotes the predator Microzoo-
19 plankton, Z denotes the top predator copepod at time T , i = 1, 2, 3, 4 is the
20 number of species in the community, and finally C denotes the effect of prey
21 on the trophic interaction, and functions Fi take into account effects of birth
22 and mortality; in most biologically meaningful situations, the functions Fi
23 is nonlinear with respect to at least some of their arguments. The second
24 part of the above equation, in D∇2 U we have the simplest (but yet biolog-
25 ically meaningful) case is given by random isotropic motion, i.e., diffusion,
26 where Di is the diffusion coefficient of the ith species, and ∇2 is the Laplace

2
Figure 1: Schematic diagram of proposed model

3
27 operator:

∂2 ∂2
∇2 = + (2)
∂x2 ∂y 2
28 Keeping in view the above discussion, we will study a plankton model
29 for one prey and two predator system with Holling type II functional re-
30 sponse Fig.??. The main contribution in this paper is to expand the two
31 trophic model studied by (?) and then study the effect of diffusion on the
32 four species food web model with Holling type II functional response. We
33 have obtained analytically as well numerically the diffusion driven instability
34 condition for the spatial system. In addition, we have obtained the different
35 types of spatial patterns i.e. Turing and non-Turing spatial patterns. Finally,
36 the higher order stability analysis for the four species prey-predator system
37 has explored. The details of this paper is organized as follows. In Section 2,
38 we have proposed spatiotemporal PMZC model, Section 3 presents Asymp-
39 totic Expansion Analysis followed by section 4 the local stability analysis
40 and limit cycle condition for the temporal system. In Section 5, we have
41 derived the analytical conditions for diffusion driven instability using Ruth
42 Hurwitz criteria and the numerical simulation are performed. In Section
43 6, the systematic analysis is discussed. Finally, conclusion is given in last
44 Section 7.

45 2. The temporal PMZC-model

46 A developed mathematical model of one prey utilized by two predators

47 and chemical released has been studied in temporal domain with Holling
48 type II functional response (?). Motivated from the work (?). We derived

4
49 Fi , i = 1..4, which is the interaction function of the developed model and it
50 has the following format: Consider the following model:

dP
= F1 (P, M, Z, C), (3)
dt
dM
= F2 (P, M, Z, C), (4)
dt
dZ
= F3 (P, M, Z, C), (5)
dt
dC
= F4 (P, M, Z, C). (6)
dt
51 Here Fi , i = 1, 2, 3, 4, which is the interaction function of the developed
52 model, has the following format:
 
P aP M
F1 (P, M, Z, C) = rP 1− − , (7)
K 1 + b1 P
53

γ1 aP M βZM ζC
F2 (P, M, Z, C) = − m1 M − (1 + ), (8)
1 + b1 P 1 + b2 M 1 + ǫC
54

γ2 βZM ζC
F3 (P, M, Z, C) = (1 + ) − m2 Z, (9)
1 + b2 M 1 + ǫC
55

ηaP M
F4 (P, M, Z, C) = − m3 C + ωP. (10)
1 + b1 P
56 The model describes the interactions between small infochemical-producing
57 phytoplankton, microzooplankton and copepods in a system that is depleted
58 of nutrients. The parameter r represents phytoplankton intrinsic growth
59 rates, a is the clearance rate of microzooplankton at low food densities, bi
60 i = 1, 2 is half saturation constants, β is the copepod linear predation rate,
61 mi (i = 1, 2) is the predator’s death rate, m3 is the chemical evaporation

5
62 rate, γi i = 1, 2, is a conversion parameter of the biomass of the prey to
63 predator. ζ is the chemical changing rate and ǫ is a key parameter that we
64 are going to use to reduce the general four species model to a special cased
65 model as in ?. η is the productivity rate of DMS-infochemical and ω is the
66 amount of chemical given off by each phytoplankton. In Eq. (??) we still
67 employ the logistic map to describe the growth rate of prey and Holling II
68 functional response to describe the effect of predator on prey. Eq. (??), we
69 define microzooplankton population growth using the Holling II functional
70 response with γ1 as a conversion parameter to convert prey biomass into
71 predator. The second term in Eq.(??) represents the normal mortality of
72 microzooplankton, while the third term represents the effect of zooplankton
73 on microzooplankton, which gives another reason for the microzooplankton
74 death. The third term also represents the increase of predation with β as
75 a linear predation rate 1 . Copepods saturate their behavior to handle prey
76 (microzooplankton) with b2 being the half saturation parameter. The re-
77 leased chemical could also be saturated by (1 + C) factor and ζ parameter is
78 used to measure the rate of the chemical increase that affects the increasing
79 of predation 2 . In Eq. (??) the first term we are trying to introduce is the
80 copepod population growth that connects both predators M and Z. This term
81 also describes how copepods consume microzooplankton following DM S re-

1
-β MZ represents the affect of copepod on microzooplankton, because any species
should saturate at some level. Therefor we changed the layout of this term to the Holling
II functional response.
2
ζ also can be defined as a maximum level of chemical released, especially if we modeled
 
this term by 1 + ζC max
1+C .

6
82 lease and how copepods saturate after the time it takes to handle prey. The
83 next term is copepods mortality due to the consuming of a higher trophic
84 predation. The last equation (??) has three terms. The first term is used
85 to describe the released infochemical following microzooplankton grazing on
86 phytoplankton with η being the productivity rate of DM S. The second term
87 in F4 stands for chemical evaporation. The third term represents chemical
88 (exuded) release by each cell. We can reduce the model in Eqs. (??)- (??)
89 into a special case model by setting ǫ = 0, b2 = 0, ω = 0, i.e:

 
P aP M
F1 (P, M, Z, C) = rP 1 − − (11)
K 1 + b1 P
γ1 aP M
F2 (P, M, Z, C) = − m1 M − βZM (1 + ζC) (12)
1 + b1 P
F3 (P, M, Z, C) = γ2 βZM (1 + ζC) − m2 Z (13)
ηaP M
F4 (P, M, Z, C) = − m3 C (14)
1 + b1 P

90 The main difference between the two set of nonlinearities and the model in
91 Eq (??) is the linear predation function, which describes the linear predation
92 of the copepod on microzooplankton. It can be assumed that the model
93 is valid over long time–scales, because we are adding more resources to the
94 basic food chain of the two species model by modeling the population density
95 of copepod and the chemical released. So, we are considering the time that
96 both predators M and Z take to handle the prey. The model in Eq. (??)
97 is considered as a special case model of Eqs. (??) - (??). Because when
98 ǫ = b2 = ω = 0 the model will be reduced to the two species model in (?) 3 .

3
not specific time scale is not convinced to use and need extra explanation

7
99 One goal of the model construction here is to predict of the predator–prey
100 kinetic and dynamic properties. Since our model is constructed from the two
101 species predator prey model P and M , a basic question to raise here is how
102 can the four species model provide better descriptions than the two species
103 model.

104 3. Qualitative analysis of the location of the equilibria

105 We now look for the steady-state solutions (P, M, Z, C) which satisfy
106 ( dp , dM , dZ
dt dt
, dC ) = 0. The system in Eqs. (??)-(??) possesses five possible
dt dt

107 nonnegative equilibria, namely the extinction equilibrium E0 , the microzoo-

108 plankton and copepod-eradication equilibrium E1 , phytoplankton and info-
109 chemical eradication equilibrium E2 , copepod eradication (free) equilibrium
110 E3 , and finally the coexistence equilibrium E4 . Table ?? shows the number
111 of equilibria and their type and definition.

Table 1: Biologically relevant and irrelevant possible equilibria of the system given by the
Eqs. (??)-(??)
Equilibrium Definition Value in Description Hyperbolic
parametrized Eigenvalues
system
E0 (Pe , Me , Ze , Ce ) (0, 0, 0, 0) Trivial (extinct) Eq. (??)
Kω
E1 (Pe , 0, 0, Ce ) (K, 0, 0, m3
) phytoplankton and infochemical equilibrium Eq. (??)
E2 (0, Me , Ze , 0) (0, Me , Ze , 0) Biologically irrelevant equilibrium given in Eq (??) Eq.(??)
E3 (Pe , Me , 0, Ce ) (P, M, 0, C) Copepod free equilibrium is given by Eq (??) Eqs. (??), (??), (??)
E4 (Pe , Me , Ze , Ce ) as in Eq Full Coexistence equilibrium given by Eq (??) Eqs.(??)
(??)

112 The biologically irrelevant equilibrium is given by:

m2 −γ2 m1
E2 = (0, , , 0). (15)
(βγ2 ) (βγ2 )

8
 The fourth equilibrium is given by:
m1 rγ1 (aKγ1 − Kb1 m1 − m1 )
E3 = ( , , 0,
(aγ1 − b1 m1 ) K(aγ1 − b1 m21 )
(aηKrγ1 − ηKrb1 m1 + aKωγ1 − Kωb1 m1 − ηrm1 )
). (16)
(K(aγ1 − b1 m1 )2 )
113 The full co-existence state satisfies a quartic polynomial
4
X
Ai Pe4−i = 0, (17)
i=0

where Ai , i = 0, .., 4 are cascading parameters given by:

A0 = γ2 b1 βηr2 + ζb1 βηr2 − b1 b2 ηm2 r2 ,

A1 = −2γ2 b1 βηkr2 − 2ζb1 βηkr2 + 2b1 b2 ηkm2 r2 − γ2 b1 βkωr − ζb1 βkωr

+ b1 b2 km2 ωr + γ2 βηr2 + ζβηr2 − b2 ηm2 r2 ,

A2 = −γ2 b1 βηk 2 r2 + ζb1 βηk 2 r2 − b1 b2 ηk 2 m2 r2 + γ2 b1 βk 2 ωr

+ ζb1 βk 2 ωr − b1 b2 k 2 m2 ωr − γ2 b1 βkm3 r − 2γ2 βηkr2 − 2ζβηkr2

+ b1 b2 km2 m3 r + 2b2 ηkm2 r2 − γ2 βkωr − ζβkωr + aηkm2 r + b2 km2 ωr,

A3 = γ2 b1 βk 2 rm3 + γ2 βηk 2 r2 + ζβηk 2 r2 − b1 b2 k 2 rm2 m3

− b2 ηk 2 r2 m2 + γ2 βk 2 ωr + ζβk 2 ωr − aηk 2 rm2 − b2 k 2 ωrm2

− γ2 βkrm3 − ak 2 ωm2 + b2 krm2 m3 ,

A4 = γ2 βk 2 m3 r − b2 k 2 m2 m3 r − ak 2 m2 m3 ,

while M , Z and C are given by

−r

M (Pe ) = b1 Pe2 − (Kb1 − 1)Pe − K ,
aK
9
114

Z(Pe ) = A5 (Pe )2 + B(Pe ) + G, (18)

(P 2 ηr − P ηKr − Kω)
C(Pe ) = .
(Km3 )
115 The full co-existence persistence state given by a quartic polynomial as
116 follows;
4
X
Ai Pe4−i (19)
i=0
where Ai , i = 0, .., 4 are all cascading parameters as follows:

A0 = γ2 b1 βηr2 + ζb1 βηr2 − b1 b2 ηm2 r2

A1 = −2γ2 b1 βηkr2 − 2ζb1 βηkr2 + 2b1 b2 ηkm2 r2 − γ2 b1 βkωr − ζb1 βkωr

+ b1 b2 km2 ωr + γ2 βηr2 + ζβηr2 − b2 ηm2 r2

A2 = −γ2 b1 βηk 2 r2 + ζb1 βηk 2 r2 − b1 b2 ηk 2 m2 r2 + γ2 b1 βk 2 ωr

+ ζb1 βk 2 ωr − b1 b2 k 2 m2 ωr − γ2 b1 βkm3 r − 2γ2 βηkr2 − 2ζβηkr2

+ b1 b2 km2 m3 r + 2b2 ηkm2 r2 − γ2 βkωr − ζβkωr + aηkm2 r + b2 km2 ωr

A3 = γ2 b1 βk 2 rm3 + γ2 βηk 2 r2 + ζβηk 2 r2 − b1 b2 k 2 rm2 m3

− b2 ηk 2 r2 m2 + γ2 βk 2 ωr + ζβk 2 ωr − aηk 2 rm2 − b2 k 2 ωrm2

− γ2 βkrm3 − ak 2 ωm2 + b2 krm2 m3

A4 = γ2 βk 2 m3 r − b2 k 2 m2 m3 r − ak 2 m2 m3

10
117 3.1. System behaviour near the origin E0

118 A straightforward calculation shows that, the hyperbolic equilibrium or

119 the first trivial (extinct) equilibria is an unstable saddle point, with the
120 unstable manifold in the direction orthogonal to the M − Z − C coordinate
121 plane.
λE0 = (r, −m1 , −m2 , −m3 ) (20)

122 3.2. System behaviour near Microzooplankton and copepod free equilibrium
123 E1

124 The equilibrium point E1 of the system with Microzooplankton and cope-
125 pod eradication is locally asymptotically stable if γ1 ak − b1 km1 − m1 ≤ 0 ,
126 and if γ1 ak < b1 km1 − m1 , 1 + b1 k > 0 then E1 is globally stable.

γ1 ak − b1 km1 − m1
λE1 = (−r, , −m2 , −m3 ) (21)
1 + b1 k

127 3.3. System behaviour near Phytoplankton and Infochemical free equilibrium
128 E2

129 The Jacobian matrix of the system ?? around the non feasible point,
130 Phytoplankton and Infochemical eradication equilibrium point E2 gives the
131 following eigenvalues.

γ2 βr − b2 m2 r − am2 1 p
λE 2 = ( , (α ∓ β), −m3 ) (22)
γ 2 β − b2 m 2 2A0

A0 = γ2 β (23)

α = −m1 m2 m3 b2 (24)

11
 q
β= (4γ2 β 2 m1 m2 − 4γ2 b2 βm1 m22 + b22 m21 m22 ) (25)

132 Now, this hyperbolic point is unstable saddle, since all the parameters
133 are positive. λ1 is unstable, and because λ2,3 has a negative real part then
134 they are stable focus and λ4 < 0 then we have saddle-focus point, as when
135 we have one real eigenvalue with the sign opposite to the sign of the real
136 part of a pair of complex-conjugate eigenvalues and the negative real fourth
137 eigenvalue; This type of equilibrium is always unstable.

138 3.4. System behaviour near copepod free equilibrium E3

139 The Jacobian matrix of the system ?? for the fourth equilibrium with
140 copepod eradication has the following four eigenvalues:

A
λ1 = (26)
B
141 where

12
A = −γ14 γ2 a3 βk 2 m3 r + γ14 a3 b2 k 2 m2 m3 r + 3γ13 γ2 a2 b1 βk 2 m1 m3 r

− γ13 γ2 a2 βηk 2 m1 r2 − γ13 ζa2 βηk 2 m1 r2

− 3γ13 a2 b1 b2 k 2 m1 m2 m3 r + γ13 a2 b2 ηk 2 m1 m2 r2 − 3γ12 γ2 ab21 βk 2 m21 m3 r

+ 2γ12 γ2 ab1 βηk 2 m21 r2 + 2γ12 ζab1 βηk 2 m21 r2

+ 3γ12 ab21 b2 k 2 m21 m2 m3 r − 2γ12 ab1 b2 ηk 2 m21 m2 r2 + γ1 γ2 b31 βk 2 m31 m3 r

+ γ1 γ2 b21 βηk 2 m31 r2 − γ1 ζb21 βηk 2 m31 r2 + γ1 b31 b2 k 2 m31 m2 m3 r

+ γ1 b21 b2 ηk 2 m31 m2 r2 + γ14 a4 k 2 m2 m3 − γ13 γ2 a2 βk 2 m1 ωr

− γ13 ζa2 βk 2 m1 ωr + 4γ13 a3 b1 k 2 m1 m2 m3 − γ13 a3 ηk 2 m1 m2 r

+ γ13 a2 b2 k 2 m1 m2 ωr − 2γ12 γ2 ab1 βk 2 m21 ωr − 2γ12 ζab1 βk 2 m21 ωr

+ 6γ12 a2 b21 k 2 m21 m2 m3 + 3γ12 a2 b1 ηk 2 m21 m2 r + 2γ12 ab1 b2 k 2 m21 m2 ωr

− γ1 γ2 b21 βk 2 m31 ωr + γ1 ζb21 βk 2 m31 ωr + 4γ1 ab31 k 2 m31 m2 m3

+ 3γ1 ab21 ηk 2 m31 m2 r − γ1 b21 b2 k 2 m31 m2 ωr − b41 k 2 m41 m2 m3

− b31 ηk 2 m41 m2 r − γ13 γ2 a2 βkm1 m3 r − γ13 a3 k 2 m1 m2 ω

− γ13 a2 b2 km1 m2 m3 r + 2γ12 γ2 ab1 βkm21 m3 r − 2γ12 γ2 aβηkm21 r2

+ 2γ12 ζaβηkm21 r2 − 3γ12 a2 b1 k 2 m21 m2 ω − 2γ12 ab1 b2 km21 m2 m3 r

− 2γ12 ab2 ηkm21 m2 r2 + γ1 γ2 b21 βkm31 m3 r − 2γ1 γ2 b1 βηkm31 r2

− 2γ1 ζb1 βηkm31 r2 + 3γ1 ab21 k 2 m31 m2 ω − γ1 b21 b2 km31 m2 m3 r

+ 2γ1 b1 b2 ηkm31 m2 r2 − b31 k 2 m41 m2 ω − γ12 γ2 aβkm21 ωr

+ γ12 ζaβkm21 ωr − γ12 a2 ηkm21 m2 r − γ12 ab2 km21 m2 ωr

− γ1 γ2 b1 βkm31 ωr − γ1 ζb1 βkm31 ωr + 2γ1 ab1 ηkm31 m2 r

+ γ1 b1 b2 km31 m2 ωr − b21 ηkm41 m2 r − γ1 γ2 βηm1 3 r2 − γ1 ζβηm1 3 r2

− γ1 b2 ηm31 m2 r2 .
13
B = γ14 a3 b2 k 2 m3 r − 3γ13 a2 b1 b2 k 2 m1 m3 r + γ13 a2 b2 ηk 2 m1 r2

+ 3γ12 ab21 b2 k 2 m21 m3 r − 2γ12 ab1 b2 ηk 2 m21 r2 − γ1 b31 b2 k 2 m31 m3 r

+ γ1 b21 b2 ηk 2 m31 r2 + γ14 a4 k 2 m3 − 4γ13 a3 b1 k 2 m1 m3 + γ13 a3 ηk 2 m1 r

+ γ13 a2 b2 k 2 m1 ωr + 6γ12 a2 b21 k 2 m21 m3 − 3γ12 a2 b1 ηk 2 m21 r

− 2γ12 ab1 b2 k 2 m21 ωr − 4γ1 ab31 k 2 m31 m3 + 3γ1 ab21 ηk 2 m31 r

+ γ1 b21 b2 k 2 m31 ωr + b41 k 2 m41 m3 − b31 ηk 2 m41 r + γ13 a3 k 2 m1 ω

− γ13 a2 b2 km1 m3 r − 3γ12 a2 b1 k 2 m21 ω + 2γ12 ab1 b2 km21 m3 r

− 2γ12 ab2 ηkm21 r2 + 3γ1 ab21 k 2 m31 ω − γ1 b21 b2 km31 m3 r

+ 2γ1 b1 b2 ηkm31 r2 − b31 k 2 m41 ω − γ12 a2 ηkm21 r − γ12 ab2 km21 ωr

+ 2γ1 ab1 ηkm31 r + γ1 b1 b2 km31 ωr − b21 ηkm41 r + γ1 b2 ηm31 r2

λ2 = −m3

1 p
λ3,4 = (α ∓ β). (27)
2A0

A0 = γ1 ak(γ1 a − b1 m1 ) (28)

α = −m1 (γ1 ab1 kr + b21 km1 r + rγ1 a + b1 m1 r)

14
 β = −4γ14 a4 k 2 m1 r + 12γ13 a3 b1 k 2 m21 r − 12γ12 a2 b21 k 2 m31 r

+ γ12 a2 b21 k 2 m21 r2 + 4γ1 ab31 k 2 m41 r − 2γ1 ab31 k 2 m31 r2

+ b41 k 2 m41 r2 + 4γ13 a3 km21 r − 8γ12 a2 b1 km31 r

− 2γ12 a2 b1 km21 r2 + 4γ1 ab21 km41 r + 2b31 km41 r2

+ γ12 a2 m21 r2 + 2γ1 ab1 m31 r2

+ b21 m41 r2

142 Now, this hyperbolic point is unstable, since all the parameters are posi-
143 tive. λ1 is unstable if A > B > 0, and because λ2,3 has a negative real part
144 then they are stable focus and λ4 < 0 then we have saddle-focus point, as
145 when we have one real eigenvalue with the opposite sign to the sign of the
146 real part of a pair of complex-conjugate eigenvalues and the negative real
147 fourth eigenvalue; This type of equilibrium is always unstable.

148 3.5. System behaviour around the coexistence equilibrium point E4

149 The Jacobian matrix J4 = (aij )4×4 . Let λi , i = 1, 2, 3, 4 be the roots of

150 the characteristic polynomially of J4 which is given by:

4
X
Ai λ4−i = 0, (29)
i=0

151 where Ai are cascading parameters and A0 = 1;

A1 = −a11 − a22 − a33 − a44 . (30)

15
 A2 = a11 a22 +a11 a33 +a11 a44 −a12 a21 +a22 a33 +a22 a44 −a23 a32 −a24 a42 +33 a44 .
(31)

A3 = −a11 a22 a33 − a11 a22 a44 + a11 a23 a32 + a11 a24 a42 − a11 a33 a44 + a12 a21 a33 + a12 a21 a44

− a12 a24 a41 − a22 a33 a44 + a23 a32 a44 − a23 a34 a42 +24 a33 a42 . (32)

A4 = a11 a22 a33 a44 − a11 a23 a32 a44 +11 a23 a34 a42 − a11 a24 a33 a42 − a12 a21 a33 a44

− a12 a23 a34 a41 + a12 a24 a33 a41 . (33)

152 From Routh-Hurwitz criterion, all the roots of Jacobian matrix have neg-
153 ative real part iff the determinant of all Harwitz matrix are positive ? then
154 any E, is locally asymptotically stable if and only if A1 > 0, A3 > 0 and
p
155 A1 A2 > A3 and A3 > A1 (A1 A4 − A2 A3 ) or A1 A2 A3 > A23 + A21 A4 .
156 Obviously, we have A1 < 0 and A3 < 0 and by depending on the Ja-
157 cobian element matrix, when a12 < 0, a21 > 0, a23 < 0, a32 > 0, a33 < 0
158 and a44 < 0. It is readily seen that A1 A2 A3 > A23 + A21 A4 . Therefore, we
159 formulate necessary and sufficient conditions for the positive equilibrium to
160 be locally asymptotically stable, follows from the RouthHurwitz criteria. For
161 this purpose, we use the following notation
aMe aPe Me b1 k
162 a11 > 0 i.e if r > (1+b1 Pe )
+ (1+b 1 Pe )
2 ( (1−2P ) ) and a12 < 0 i.e if
e
−aPe
(1+b1 Pe )
<0
Pe Ce −1
163 and a21 > 0 i.e. b1 < (1+b1 Pe )
and a23 < 0 if ζ < Ce
and a44 < 0. Hence E4
164 is locally asymptotically stable Equilibria.

16
165 4. Asymptotic Expansion Analysis

166 Using an asymptotic approach it is possible to make some limited analyt-

167 ical progress with the general system given in Equations??. In this analysis,
168 we study how to scale our parameters in order to determine the general
169 stability for PMZC-models’ roots. We can start performing the method by
a = aǫ , bb1 = bb2 =
b
k b1 b2 m1
170 setting the following assumption: k = ǫ2
, b ǫ2
, ǫ2
, c1 =
m ǫ
,
m2 m3 eta γ1 γ2
171 c2 =
m ǫ
, c3 =
m ǫ
, η= ǫ
, γ1 = ǫ
, γ2 = ǫ
. and by substitute our scaled
172 parameters and the following expansion into the full co-existence persistence
173 state, which is given by ??:
P0 P1
P = + + P2 + ... (34)
ǫ2 ǫ
174 we will obtain an expanded polynomial, by collecting the coefficient of its
175 leading order we could obtain the appropriate value of prey density Pe . Which
176 is:

b
k
PE4 = (35)
ǫ2
and by substituting the expanded (scaled) value of P into the quadratic
polynomial of M ZC, we could have the following values for the predators
M Zand chemical release C.
1
ME 4 =
b
aǫ
−ζη 2 rb
k γb2 b1 m3 (b
kγ2 b1 + b
kb1 + γ2 + 1)
ZE4 =
ǫ
b
ηbkr
CE4 = .
m3

17
177 Now, after determine the values of scaled root E4 , we could study the gen-
178 eral stability analysis by following the same procedures method we presented
179 on this section i.e
λ0 λ1
λE 4 = + ... (36)
ǫ +
180 An expanded characteristic polynomial could be obtained following the
181 same procedures and by substituting ?? and collecting the coefficient of the
182 leading order and then solve for λ, we could obtain the following eigenvalues:

γb1ba ζb + γb2
λE4 = (0, 0, , ). (37)
bb1 bb2

183 5. Parameter Values investigation

184 A major reason for modelling the dynamics of a population is to under-

185 stand the principle controlling features and to be able to predict the likely
186 pattern of development consequent upon a change of environmental param-
187 eters ?. In PMZC model of ?? we assumed that the parameters within the
188 elementary analysis are similar to their values in the two previous chapters
189 ?.We denote their values as default values. Apparently, oscillatory solutions
190 were present in the two models, when κ = b2 = ω = 0 and when κ = 1,
191 b2 = 0.05, ω = 0.1 which makes the two systems consistent with the result
192 of the two previous chapters, i.e the model is still relaxing to stable steady
193 state. However, in this chapter we are studying 4 species system. Therefore,
194 we need to consider carefully the effect of each parameter on PMZC food
195 chain, that might help us to gain a valid solution especially we are introduc-
196 ing the effect of Zooplankton on this system. Following ?, parameter values

18
197 used to model zooplankton mortality can have a major influence on the dy-
198 namics of simple models ?, ?. They found that, for their particular parameter
199 values, limit cycle behaviour (unforced oscillations) which occurred when us-
200 ing a linear zooplankton mortality term did not occur when using a quadratic
201 term. Depending on the ? and ? we will keep the maximum growth rate
202 of logistic growth models, and belongs to the range 0.1 < r < 2d−1 . As
203 ? considered the phytoplankton carrying capacity of K = 50µgCI −1 and ?
204 considered a much wider range K = ∞. Therefor, we will use 50 < K < ∞
205 ?.
206 ?, ? estimated the half saturation constant of phytoplankton to belong
207 to the range 20 < b1 < 150µgCI −1 , zooplankton (copepod) half saturation
208 constant 20 < b2 < 100µgCI −1 to reflect that copepod dynamics evolve
209 on slower time scale than microzooplankton dynamics. However, because
210 we are introducing zooplankton (copepod) and we are going to study it’s
211 effect on this food chain. the accurate value for b1 and b2 to be used in
212 this model is much smaller than the literature suggested. We will have a
213 very small value for zooplankton(copepod) population density in contrast
214 with microzooplankton, as we are not introducing any higher trophic, and
215 because of the non-nutrient limited. We have chosen b1 P and b2 M terms,
216 because this term may be regarded as reflection the time it takes for predator
γ1 a
217 to handle ? and if we choose b1
− m1 > 0 the predator density tend to zero
γ1 a γ2 β
218 over time. Therefore, we postulate that b1
> m1 similarly for b2
> m2 .
219 Also, we can’t choose b2 > b1 in this model for same reasons.
220 In ? microzooplankton conversion efficiency is estimated as 0.15 < γ1 <
221 0.64, ? states that the conversion efficiency may be higher when considering

19
222 zooplankton feeding on microplankton, hence a higher value of γ2 = 0.7 is
223 chosen for the copepod assimilation efficiency (nicola2014). Also, a value
224 of the maximum copepod predation rate were chosen as β = 1d−1 . In our
225 model, copepods are specified with a normal mortality value as we didn’t
226 account for predation from higher predators as well as natural mortality.
227 Moreover, if we want to check the effect of choosing these specific values we
228 can do so numerically by substituting the default values from table ?? into
229 the functional responses to check the corresponded population density.

20
 Table 2: Outlines, default values and ranges of the parameters. The ranges cover values used by a variety of authors in different
models.?, ? and ?.
Parameters Definition Value Unit Range
r Phytoplankton intrinsic growth rate 1.5 days−1
K Phytoplankton Carrying capacity 120 µg C I −1
a clearance rate of microzooplankton at low food densities 0.3 µg CI −1 days−1
b1 Half saturation density 0.05 µg C I −1 0.01 −
0.05
b2 Half saturation density 0.02 µg C I −1 0.01 −
21

0.05
γ1 Microzooplankton grazing efficiently 0.3 days−1 0.3 −
0.64
γ2 Mesozooplankton grazing efficiently 0.7 - 0.3 − 0.7
m1 Microzooplankton mortality in the absence of DMS 0.1 days−1 0.015 −
0.15
m2 zooplankton mortality in the absence of DMS 0.15 days−1 0.015 −
0.15
m3 Chemical evaporate or flux to the outomosphere 0.03 days−1
η DMS production rate 0.1 -
β Mesozooplankton Linear Predation Term 1 -
ζ Chemical increasing rate CP -
ω exotic rate for each phytoplankton cell 0.01 -
230 Here we report the parameter values that have been used in the current
231 study.

232 6. Numerical simulation Results

233 6.1. Time Series and Phase Portraits

234 The main objective of this section is to support the analytical finding
235 with the help of experimented parameter values from published literature,
236 reported in table ??. The table is of the 14 parameters of the model ??
237 taken in the same range of the corresponding range of parameters shown in
238 the analysis of the previous chapters. Also they are closely related to the
239 value of the main control parameter ν or m3 that helped us to set initial
240 predict (IC) in the numerical analysis to obtain consistent results. One of
241 the integral purpose of this section is to verify our analytical finding in Table
242 ??, these numerical simulation show important features of the system from
243 practical point of view.
244 Figure ?? exhibits the local stability of the model around a proposed
245 initial condition that we set to test the consistency in between the two species
246 and the expanded model with the parameter values given in table ??.
247 In figure ??, time evolution shows the plot of the special case model
248 and the trajectory in P M ZC space of the system from the proposed initial
249 condition (Pe , Me , Ze , Ce ) = (3.942, 5.789, 0.0481, 20.379) and ζ = 0.01 and
250 all parameters fixed at their default values. In ?? The trajectory is attracted
251 onto a limit cycle, with a specific period of roughly 500 days. The trajectory
252 exhibits large-amplitude fluctuations in P .

22
 (a) (b)

(c) (d)

Figure 2: Time series and phase-space trajectory for the two cases of P M ZC systems
around the proposed initial condition and all of the other parameters are fixed as in table
?? and ζ = 0.01. The trajectories in the two cases are attracting to the equilibrium point
generating a stable limit cycle.

23
 (a) (b)

Figure 3: Time Series and Phase Portraits near Microzooplankton and copepod free equi-
librium point E1 and ζ = 0.01.It is readily seen that the trajectories are attracted onto a
stable limit cycle

(a) (b)

Figure 4: Time Series and Phase Portraits free equilibrium point E3 with all of the other
parameters fixed as in ?? and ζ = 0.01. The trajectories are attracted onto a stable limit
cycle

24
 (a) (b)

Figure 5: Time Series and Phase Portraits around the coexistence equilibrium point E4
with all of the other parameters fixed at their default values and ζ = 0.01. In ?? shows
the limit-cycle component, in red, blue, dashed blue and in black for PMZC-model. The
limit-cycle shows the overall movement in time series, ignoring the seasonality and any
small random fluctuations. In??.shows that trajectories are attracted onto a stable limit
cycle.

25
253 The dynamics of the prey-dependent model are either stable coexistence,
254 unstable coexistence, or extinction of the predator.

255 6.2. One Parameter Bifurcation Behavior

256 we proposed numerical simulations of the proposed model ??. Figure

257 ?? exhibits the local stability diagram around E4 with the parameter values
258 given in table ??. It is readily seen that ?? shows that, the coexistence equi-
259 libria E4 < K then the population of phytoplankton will bloom. Especially,
260 if E4 is a function of ζ which can reflect the consistency in between our pre-
261 vious work and the current one. However, if E4 > K then the population
262 of prey will decrease significantly. Also, if E4 = K then the population will
263 remain constant (?).
264 These graphs illustrate P M ZC biological model: For a specific choice of
265 parameters, in each cycle, the prey population is increased to extremely high
266 numbers, yet recovers (while the predator population remains sizeable at the
267 highest prey density). In real-life situations, however, chance fluctuations
268 of the discrete numbers of individual structure and life-cycle of prey might
269 cause the prey to actually go extinct, and, by consequence, the predators as
270 well (?).

26
 (a) (b)

(c) (d)

Figure 6: The coexistent equilibrium point w.r.t ζ as infochemical measurement, shows

that the range of our conrol parameter should be 0 < ζ < 0.1 to avoid having negative
real root for Z.

27
271 7. Turing analysis in P M ZC model

272 Several studies have been devoted to the various modes of spatiotemporal
273 organization generated by different models: limit cycles, Turing patterns, and
274 traveling waves in one-dimensional systems .The interest here is to provide a
275 fundamental understanding of how a steady state that is locally asymptot-
276 ically stable in a nonspatial system can become unstable in corresponding
277 diffusive system. Considering the system in ??, where Ui and i = 1, 2, 3, 4 is
278 vector of the system state variables P, M, Z, C at the position X and time t.
279 It is readily seen that the corresponding linearized system in the vicinity of
280 a spatial homogeneous steady state (Pe , Me , Ze , Ce ) has the following form :

4
dUi (t) X
= aij Uj (38)
dt j=1

281 where Ui = P, M, Z, C, and i = 1, .., 4 this is in the case of a spatially

282 homogeneous perturbation, and

4
dUi (t, k) X

= aij − k 2 Di δij Ui (t, k) (39)
dt j=1

283 In the case of spatially inhomogeneous e perturbation with k as a wave

∂fi
284 number. Here aij = ∂Uj
and δij is the Kronecker symbol, and Ui (t) and
285 Ui (t, k) are the amplitudes of the perturbation and its Fourier transform,
286 respectively ?. It is readily seen that when k = 0 system ?? coincides with

28
287 ??. The matrix of the linearised system is:

  
a11 − DP k 2 a12 a13 a14 P1
  
 2  
 a21 a22 − DM k a23 a24   M1 
A=





 a31 a32 a33 − DZ k 2 a34   Z1 
  
2
a41 a42 a43 a44 − DC k C1
(40)
288 We obtained the dispersion relation as a quartic polynomial and by solv-
289 ing this polynomial for δ we can study Turing instability conditions

4
X
Pi δ 4−i = 0, (41)
i=0

290 where Pi are all cascading parameters.

291 According to Ruth-Hurwitz criterion, all the eigenvalues have negative
292 real part if and only if the following conditions hold:

293 • P1 > 0,

294 • P3 > 0,

295 • P1 P2 > P3 ,
p
296 • P3 < P1 (P1 P4 − P2 P3 ) or P1 P2 P3 > P32 + P12 P4

297 Apparently, depending on the analytical analysis in table ?? E1 is steady

298 state of the system ??, called phytoplankton and infochemical equilibrium.
299 This state is stable under the common two conditions trace(A) < 0 i.e a11 +
300 a22 + a33 + a44 < 0.

29
301 Spatiotemporal chaos arising from the diffusive coupling of local limit
302 cycle oscillators (?) patterns in two- and three-dimensional systems including
303 patterns arising from the interference of different instability mechanisms.

304 8. Spatial distribution of the species

305 In this section we are considering P M ZC model in ?? of time and space

306 continuous, which is of four interacting species as an example of population
307 community with oscillatory kinetics. Where U is a vector of four components
308 (P, M, Z, C). In this section we are interested in the possibility of non Turing
309 patterns, we assume that the diffusivity is the same for all species. However,
310 It appears that the type of the choice of the initial condition given by equation
311 ?? could effect the type of patterns (?).

U = Ue + ǫcos(Lx .w). (42)

312 The initial condition in ?? depends on the value of ǫ and the value of the w
313 the wave number. Figure ?? shows how the initial condition evolve to smooth
314 spatial distribution of predators only. The spatial distribution gradually vary
315 in time : the local temporal behaviour of the dynamics variables P, M, Z, C
316 are strictly periodic depending on the limit cycle of the nonspatial system.
317 Figure ?? shows the population density of the dynamics variables in time
318 t = 150 using Ue = E1 and ?? shows the population density of the dynam-
319 ics variables in time t = 135 using Ue = E3 . Both figures show irregular
320 spatiotemporal oscillations invade over the whole domain.
321 In Fig.??, the spatial distributions of prey species at different time is
322 presented to show the continuous change in the distribution of species. Pat-

30
 (a) (b)

Figure 7: Population distribution over space at t = 150 and t = 135 for different steady
state obtained for the initial condition ?? with ǫ = 10−3 ; other parameters are the same
as in ??.

323 terns are presented here for the latest time t = 1500 , but the existence of
324 similar patterns are verified with long time simulations. This type of pattern
325 is classified as interacting spiral pattern and it is a non-Turing pattern.
326 In Fig.??, We find Turing patterns for PMZC model only for ζ > 0.01
327 and suitable choice of the parameter, we have observed that the stationary
328 Turing patterns are cold spot patterns which shows the existence of circular
329 patches with lower concentration of prey and predators. The stationary
330 Turing patterns observed for the classical Holling-functional response are of
331 two types, hot spot pattern and cold spot pattern. Hot spots are generated
332 through the localized circular patches with high concentration of population
333 density. Our stationary cold spot pattern changed to chaos pattern due to
334 coalesce of nearby circular patches with low concentration of population. The
335 stationary patterns obtained for the PMZC model are independent of initial
336 condition, we have checked this independence numerically, by considering E4

31
(a) (b)

(c) (d)

(e) (f)

32

(g) (h)
 (a) (b)

(c) (d)

(e) (f)

Figure 9: Turing Patterns predator-prey distribution over two dimensional space for t =
150, 350 and t = 1500; other parameters are the same as in ??.

33
337 as initial guess without perturbing it and we obtained a flat state.
338 This non-stationary property is illustrated in Fig.??, where spatial aver-
339 age of population densities are plotted against time as in ??. It is important
340 to note here that the temporal steady-state is unstable and started oscillating
341 for ζ > 0.001.

34
 (a) (b)

(c) (d)

(e)

Figure 10: Simulation analysis for Turing patterns for ζ = 0.001; other parameters are the
same as in ??.

35
342 9. Discussion

343 Many classical prey dependent food chain models, were studied by many
344 authors since the innovative work of Freedman ? and ? to obtain a better
345 insight of the behaviour of the solution trajectory. In the present investiga-
346 tion, an attempt has been made to discuss the occurrences of the trajectories.
347 To summaries the analytical finding we used table ?? and we obtained some
348 numerical results by using the parameter values in table ?? generated us-
349 ing ode45 in MATLAB. The model in ?? exhibits four biological feasible
350 equilibria and one non feasible equilibria E2 . The first one, is E0 where all
351 populations are extinct which is always unstable. The second one is E1 ,
352 where the prey is at the highest population while the predator Microzoo-
353 plankton and the top predator Zooplankton are extinct and the release of
354 infochemical is limited. The third one is E3 , where the prey and predator
355 Microzooplankton and infochemical release are persist while the top predator
356 Zooplankton is extinct. The fourth equilibria is the coexistence E4 where all
357 species are exist ? .
358 The set of parameters in table ?? allows a wide range of behaviour to
359 be observed in a relatively small ζ parameter space. The stability of the
360 system falls into four categories, based on the predator-prey population, or
361 limit-cycle, graph behaviour. An unstable spiral spirals away from the initial
362 condition, yielding population oscillations that grow without bound. A neu-
363 tral center indicates a closed loop, where population oscillations stay steady
364 without damping. A stable spiral spirals in to reach a point of stability, and
365 population oscillations are damped until both populations remain steady.
366 Also, it should be noted in here that the value of ζ should not exceed 0.01,

36
367 this is to obtain a feasible solution and set well posed model, because if we
368 increased this value we will have a negative value for Z and lead to a non
369 biological relevant root. Studying Turing instability in P M ZC model using
370 Routh Hurwitz criterion helped to understand the effect of the space on the
371 homogeneous steady state i.e. temporal model. Furthermore, based on the
372 numerical experiments results, we obtained that depending on the parameter
373 range given in table ?? the spatial homogeneity in Equations ?? has gave
374 way to regular spatiotemporal pattern that emerged to give chance to the
375 chaotic pattern appears. It is important to clarify that the spatiotemporal
376 chaos is typically for two dimensional reaction-diffusion system with oscilla-
377 tory dynamics. Different patterns may obtained by seeking different initial
378 conditions and using different numerical methods.
379 Finally, we have addressed the issue of Turing and non-Turing Patterns
380 in spatio-temporal PMZC model. Spatiotemporal chaotic pattern exists for
381 parameter values given in Table ??.These observations confirm the fact that
382 the interaction between the temporal and spatial instability are unable to
383 drive the system towards spatial and temporal irregularity under any circum-
384 stances. Relatively, the existence of the irregular distribution of population
385 over space and its continuous change with time depend on the complex inter-
386 action is taking place over spatial and temporal scale. There are no unique
387 criteria for the existence of spatio-temporal chaos ?.

37
388 10. References

38