This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
1. Introduction
Generally speaking, spatial or geographic heterogeneity plays an important role in the transmission process of many infectious diseases when species interact. This propagation of infectious diseases has been known to be an important regulating factor for human and animal population sizes. The study of the effect of infectious disease propagation in species in interaction has attracted some attention in ecology and epidemiology due to its seriousness threats all over the world. In most of the mathematical models, environment has been considered as homogeneous. However, in reality, environment is heterogeneous, and it can be considered as a set of different localities connected by migration [1, 2]. In particular, for prey–predator population, infectious diseases coupled with prey–predator model produce a complex dynamic given the multitude of species living in the environment and interacting with each other. This complexity is increased in the presence of migration in each species. We cannot ignore this factor because it is common in the ecological system and plays a major role in the natural regulation of populations.
After the pioneer work of Kermack and McKendricK [3], epidemiological models have drawn consideration of many biologists and ecologists. The infectious diseases in the community of prey–predator models has been studied extensively in literature [1, 4–14]. In [14], Tewa et al. proposed and investigated a prey–predator model with an SIS infectious disease affecting preys or predators or both. They showed that the disease can disappear from the community, persist in one or two populations of the community. Savadogo et al. [8] proposed and analyzed an eco-epidemiological model describing the effect of predation in the dynamic of propagation of disease. Results concerning the local and global stability have been analyzed according to Routh–Hurwitz criterion and Lyapunov principle, respectively. They also established the Hopf-bifurcation to highlight the periodic fluctuation with extinction or persistence of the disease in the preys and predators communities. In [5], Biswas et al. proposed and investigated a cannibalistic eco-epidemiological model with disease in the predator species. They conclude that cannibalism is process of regulation and govern the disease transmission in the predators community. Many works have been investigated in literature on ecological models with disease in prey only [15–18]. For instance, Greenhalgh et al. [17], studied an eco-epidemiological model with fatal disease in the prey population. In [16], Chattopadhyay and Arino studied a prey–predator model with disease in the prey population. They showed persistence and extinction conditions of the populations and also determined conditions for which the system enters a Hopf-bifurcation.
Furthermore, it is well-known that migration can be described as an important demographic process that occurs in all living beings. It can be occurred for many reasons such as education, employment, marriages, war for human beings. For animal species, migration is usually due to the climate change, for habitat, looking for food, predation, cannibalism, reproduction, urbanization, deforestation, etc. Many researches are mainly focused on the effect of infectious diseases in the community of prey–predator models under the influence of migration [4, 11, 13, 19–21]. Indeed, Kant and Kumar [11] proposed and studied a prey–predator model with disease in both species by taking into account the process of migration in only prey population. In their mathematical analysis, existence, positivity and stability of equilibria has been investigated. Moreover, the epidemiological thresholds are computed and used to determine the conditions of the disease propagation. In [4], Arora and Vivek proposed and investigated a prey–predator model when disease spread among prey populations with migration in both species. Holling type II functional response is used for interaction between prey and predator species. Their mathematical analysis has permitted to establish existence of equilibria and the local and global stability. Chowdhury [13] investigated an eco-epidemiological model describing the effect of fast migration on prey–predator model between two different patches. Mathematically, the author proved the asymptotic stability of the unique fast equilibrium point and the aggregated model.
It is in this line of thought that, in this paper, we propose and study an eco-epidemiological prey–predator model to study the effect of migration on the dynamic of prey and predator. Indeed, motivated by the works of Arora and Kumar [4], Kant and Kumar [11], and Tewa et al. [14], our main goal in this work, is to analyze the effect of migration on the dynamic of prey–predator model in the presence of an SIS infectious diseases. We found inspiration in the work of an eco-epidemiological model studied by Tewa et al. [14], by taking into account the migration. Besides, we established the conditions of existence when our model admits at least coexistence equilibrium. Based upon thorough of mathematical analysis of our model under consideration, all the equilibria point of the system are adequately characterized, and their stability analysis are investigated following the Routh–Hurwitz criteria and Lyapunov principle. Also, we established the existence of the limit cycles of the system studied arising from Hopf-bifurcation. Finally, in the numerical simulation, bifurcation diagrams and phase portraits are given, and some complex and rich dynamic behaviors, such as limit cycle, periodic solutions are found. The results show that variation of migration parameters may affect prey and predator density, thereby controlling species density can effectively maintain the ecological balance.
The remaining part of this paper is structured as follows after the statement of the problem. Section 2 is devoted to the formulation of the eco-epidemiological model. In Section 3, a mathematical analysis of the model is established, including well-posedness, stabilities analysis, and Hopf-bifurcation. We perform some numerical simulations to support our main results in Section 4. The paper ends with a conclusion and discussion in Section 5.
2. Mathematical Formulation of the Eco-Epidemiological Model
In this section, our goal is to establish an eco-epidemiological model in order to study the effect of migration on the dynamic of preys and predators population. Let’s denote by
We list the following key assumptions useful in the mathematical formulation of our prey–predator system.
(H1): When there is no predator, the prey population growth logistically
(H2): The Holling function response of type II is used to represent the process of predation and is defined by
(H3): The disease is transmitted by contact between an infected and susceptible prey by standard incidence
(H4): The disease is not genetically inherited. The infected population can recover or become susceptible to diseases;
(H5): Only susceptible prey is capable of reproducing and contributing to their carrying capacity.
According to the above assumptions and the interaction diagram of Figure 1, the dynamics of the global eco-epidemiological model is given by the following set of differential equations:
(1)
(2)
(3)
(4)
(5)
(6)
(7)
By setting
[figure(s) omitted; refer to PDF]
3. Mathematical Investigation of Model
This section deals with mathematical analysis of system (2) [1].
3.1. Existence, Positivity, and Boundedness Properties
For system (2) to be ecologically and epidemiologically meaningful, it is important to prove that all its state variables are nonnegative for all time. Then, we rewrite model (2) in the following form:
The following results hold for model (2) [1, 8].
Theorem 1.
The nonnegative orthant
Theorem 2.
System (2) admits a unique global solution
Proof.
Indeed,
(1) The theorem of Cauchy–Lipschitz assures the existence and uniqueness of local solution of system (2) on
(2) Now let us show that
The boundedness of system (2) is given by the following theorem.
Theorem 3.
The closed set defined by:
Proof.
Indeed, adding the three equations of system (2), we obtain:
The time derivatives
From Equation (8), we get:
By integrating the differential inequality Equation (7) and by using the theory of Birkoff and Rota [23] yields:
Then,
Thus, the system (2) are bounded. Particularly,
3.2. Stability Analysis of Trivial Equilibria
In this subsection, we discuss the existence and the local stability of each trivial equilibrium point. Let’s consider the following epidemiological and ecological thresholds, with clear and distinct biological meaning [14, 24]:
(i)
(ii)
(iii)
The trivial equilibria is obtained by setting the right-hand sides of system (2) to zero. The explicit expressions of the trivial equilibria are given by the following proposition:
Proposition 1.
(i)
(ii)
(iii)
This equilibrium is ecologically admissible if:
Now, we are in position to investigate the local stability of trivial equilibria.
Proposition 2.
(i)
(ii)
(iii)
Proof.
Let us consider the variational matrix of system (2):
(i) The Jacobian matrix of system (2) at
The eigenvalues are
(ii) The Jacobian matrix of system (2) at
The characteristic polynomial is given by:
Thus, the eigenvalues of
In the light of the Routh–Hurwitz criteria,
(iii) The Jacobian matrix of system (2) at
The characteristic polynomial associated is as follows:
Then, the eigenvalue of
3.3. Existence of Coexistence Equilibria
To compute the equilibrium solutions, we set the right-hand-side of system (2) to zero. Thus, we get:
Assume that
By plugging
By substituting
Since
Table 1
Various possibilities for positive real roots of
Cases | Number of sign changes | Number of positive roots | ||||
1 | + | + | + | + | 0 | 0 |
2 | + | + | + | — | 1 | 1 |
3 | + | + | — | — | 1 | 1 |
4 | + | — | — | — | 1 | 1 |
5 | + | — | + | + | 2 | 0,2 |
6 | + | + | — | + | 2 | 0,2 |
7 | + | — | — | + | 2 | 0,2 |
8 | + | — | + | — | 3 | 1,3 |
Then, the following result gives the existence of the coexistence equilibrium point [25].
Theorem 4.
System (2):
(i) has no feasible coexistence equilibria points if case 1 is satisfied;
(ii) has a unique feasible coexistence equilibrium points if the conditions (40) and (42) and cases 2, 3, and 4 are satisfied;
(iii) has two feasible coexistence equilibria or no feasible coexistence equilibria points if the conditions (40) and (42) and cases 5, 6, and 7 are satisfied;
(iv) has three feasible coexistence equilibria points or a unique feasible coexistence equilibrium point if the conditions (40) and (42) and case 8 are satisfied.
3.4. Local Stability Analysis of the Coexistence Equilibrium Point
Let us define
Proposition 3.
If the conditions (ii) of Theorem 4 and the following conditions are satisfied:
Proof.
Indeed, by linearizing model (2) around the coexistence equilibrium point
Therefore, the characteristic polynomial is given by:
From expressions (48) and (50), we get, respectively,
According to Equations (49) and (51), we get, respectively,
3.5. Bifurcation Analysis of the Coexistence Equilibrium
Here, we establish the conditions when Hopf-bifurcation occurs at
The following theorem gives the conditions that Hopf-bifurcation occurs [15, 26–28].
Theorem 5.
If the conditions (ii) of Theorem 4 and the following conditions are satisfied:
Proof.
Indeed, assuming that
According to expressions (49), (65), and (66), we get
At
According to
These roots are functions of
We have:
Differentiating Equation (69) with respect
By separating real and imaginary parts, one has:
Verifying that
At
Therefore, the transversality condition hold. Thus, Hopf-bifurcation occurs at
Now, let us investigate the global stability of the coexistence equilibrium point [12, 26, 29–32].
3.6. Global Stability Analysis of the Coexistence Equilibrium Point
Let us define:
Setting
The following theorem give the global stability of coexistence equilibrium.
Theorem 6.
Setting
Proof.
Indeed, let us consider the function
It is straightforward to see that
The time derivative of
Using the fact that
By substituting
It is obvious to see that:
Plugging the above expressions into Equation (86) and rearranging gives:
Noticing that,
Applying the following classical relation:
By using the above relations, one has
However, by grouping term by term, we obtain:
Consequently,
4. Numerical Simulation
In this section, numerical simulation has been performed to illustrate our theoretical findings by using the parameters given in Table 2 [1, 14]. Our goal here is to illustrate numerically the effect of migration on the dynamics of system (2). Through Figures 2–4 we present the evolution of the population of the prey and predators. Figure 2 shows the convergence of the solution of system (2) toward the coexistence equilibrium
Table 2
Numerical values of the parameters of system (2) used in the simulation.
Parameters | Values | References |
1.5 | [8, 14] | |
366 | Estimated | |
0.2 | Estimated | |
0.03 | Estimated | |
0.02 | Estimated | |
0.1 | Estimated | |
0.001 | Estimated | |
0.015 | [8, 14] | |
0.5 | Estimated | |
2.5 | Estimated | |
1 | [14] | |
0.15 | [14] | |
0.02 | Estimated | |
0.5 | Estimated |
[figure(s) omitted; refer to PDF]
We continue our numerical analysis in order to study the effect of migration in the community of prey and predators. From Figure 5(e), we observe that the trajectories of system (2) approach asymptotically toward
Remark 7.
The dynamics of system (2) present variations when the migration parameters are varied while keeping the other parameters fixed. Thus, when the migration rate of the susceptible prey population
[figure(s) omitted; refer to PDF]
5. Conclusion
The process of migration in the dynamics of a prey–predator model in the presence of infectious disease play a major role in the natural mechanisms of regulation of species. It is in this line of thought that we are interested in this paper, to the study of the dynamics of prey–predator populations with infectious disease to describe the effects of migration in the dynamics of species. The formulation of the model derives from an ODE system by considering Holling function response of type II to represent the strategy of predation between the prey and the predator. The mathematical analysis allowed us first to establish that the model is ecologically and epidemiologically well-posedness. Thus, the existence, the positivity and the boundedness of the solutions are proved. Moreover, we established the conditions of existence of the coexistence equilibria. Under certain thresholds
We used different scenarios in the model numerical simulation in order to show the effect of migration on the dynamics of the prey and predator populations. Thus, for
In last, in this present work, the model is formulated by using the ordinary differential equations. However, it has been proven that the use of fractional derivatives gives a more realistic description of most biological issues. Therefore, for our future study, it will be interesting to consider fractional derivatives while formulating an eco-epidemiological model with migration that would give a better description of the ecological process [33, 34].
Disclosure
A preprint of this paper presenting the main findings is available online: https://papers.ssrn.com/sol3/papers.cfm?abstract_id=4347564 [31].
Authors’ Contributions
All the authors read and approved the final manuscript.
Acknowledgments
The authors are also very grateful to professor Pierre-Alexandre Bliman of the Sorbonne Université for his insightful and constructive comments and suggestions which helped to improve the original manuscript. This work was supported by the PDI-MSC program. The authors are very much thankful to Prof. Christophe Cambier of the Sorbonne Université (France), the coordinator of the program.
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Abstract
In this paper, we propose an eco-epidemiological mathematical model in order to describe the effect of migration on the dynamics of a prey–predator population. The functional response of the predator is governed by the Holling type II function. First, from the perspective of mathematical results, we develop results concerning the existence, uniqueness, positivity, boundedness, and dissipativity of solutions. Besides, many thresholds have been computed and used to investigate the local and global stability results by using the Routh–Hurwitz criterion and Lyapunov principle, respectively. We have also established the appearance of limit cycles resulting from the Hopf bifurcation. Numerical simulations are performed to explore the effect of migration on the dynamic of prey and predator populations.
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1 Jacques-Louis Lions Laboratory (LJLL), Sorbonne University, Paris UMR 7598, France; Laboratory of Mathematics, Computer Sciences and Applications (LaMIA), Nazi BONI University, 01 BP 1091 Bobo Dsso 01, Bobo-Dioulasso, Burkina Faso
2 Laboratory of Mathematics, Computer Sciences and Applications (LaMIA), Nazi BONI University, 01 BP 1091 Bobo Dsso 01, Bobo-Dioulasso, Burkina Faso
3 Laboratory of Mathematics, Computer Sciences and Applications (LaMIA), Gaoua University Center, Nazi BONI University, 01 BP 1091 Bobo Dsso 01, Bobo-Dioulasso, Burkina Faso