Abstract
This paper investigates the impact of spatial heterogeneity on the interaction between similar strains in a dynamical system of coinfecting strains with spatial diffusion. The SIS model studied is a reaction–diffusion system with spatially heterogeneous coefficients. The study considers two limiting cases: asymptotically slow and fast diffusion coefficients. When the diffusion coefficient is small, the slow system is shown to be a semilinear system of “replicator equations,” describing the spatiotemporal evolution of the strains’ frequencies. This system is of the reaction–advection–diffusion type, with an additional advection term that explicitly involves the heterogeneity of the associated neutral system. In the case of fast diffusion, traditional methods of aggregating variables are used to reduce the spatialized SIS problem to a homogenized SIS system, on which the results of the non-spatial model can be applied directly.
Similar content being viewed by others
Availability of data and materials
Not applicable
Code availability
Not applicable
Notes
This is due to the fact that each equation reads \(\partial _t u_i -d\Delta u_i =f_i(u)\) with \(f_i(u_{|u_i=0})\ge 0\).
The name semi-neutral system comes from the fact that if \(\epsilon =0\), except the coefficients of diffusion terms, then the parameters do not depend on the strains as in the neutral theory.
We use the usual notation abuse. Rigorously speaking, we have to define \(\widetilde{X}(\tau )=X\left( \frac{\tau }{\epsilon }\right) \) and the same for each variables. Here we remove the \(\widetilde{}\) for simplicity.
References
Poggiale, J.C.: Lotka–Volterra’s model and migrations: breaking of the well-known center. Math. Comput. Model. 27(4), 51–61 (1998). https://doi.org/10.1016/S0895-7177(98)00005-3
Poggiale, J.C.: Predator–prey models in heterogeneous environment: emergence of functional response. Math. Comput. Model. 27(4), 63–71 (1998)
Wang, Z.-C., Wu, J.: Travelling waves of a diffusive Kermack–Mckendrick epidemic model with non-local delayed transmission. Proc R Soc A Math Phys Eng Sci 466, 237–261 (2009)
Wang, X.-S., Wang, H., Wu, J.: Traveling waves of diffusive predator–prey systems: disease outbreak propagation. Discrete Contin. Dynam. Syst. 32, 3303–3324 (2012)
Adler, F.R., Brunet, R.C.: The dynamics of simultaneous infections with altered susceptibilities. Theor. Popul. Biol. 40(3), 369–410 (1991). https://doi.org/10.1016/0040-5809(91)90061-J
Alizon, S.: Co-infection and super-infection models in evolutionary epidemiology. Interface Focus 3(6), 20130031 (2013)
Martcheva, M.: A non-autonomous multi-strain sis epidemic model. J. Biol. Dyn. 3(2–3), 235–251 (2009)
Marchaim, D., Perez, F., Lee, J., Bheemreddy, S., Hujer, A.M., Rudin, S., Hayakawa, K., Lephart, P.R., Blunden, C., Shango, M., Campbell, M.L., Varkey, J., Manickam, P., Patel, D., Pogue, J.M., Chopra, T., Martin, E.T., Dhar, S., Bonomo, R.A., Kaye, K.S.: “swimming in resistance’’: co-colonization with carbapenem-resistant enterobacteriaceae and Acinetobacter baumannii or Pseudomonas aeruginosa. Am. J. Infect. Control 40(9), 830–835 (2012). https://doi.org/10.1016/j.ajic.2011.10.013
Warren, D., Nitin, A., Hill, C., Fraser, V., Kollef, M.: Occurrence of co-colonization or co-infection with vancomycin-resistant enterococci and methicillin-resistant staphylococcus aureus in a medical intensive care unit. Infect. Control Hosp. Epidemiol. 25(2), 99–104 (2004)
Madec, S., Gjini, E.: Predicting n-strain coexistence from co-colonization interactions: Epidemiology meets ecology and the replicator equation. Bull. Math. Biol. 82(11), 142 (2020). https://doi.org/10.1007/s11538-020-00816-w
Le, T.M.T., Gjini, E., Madec, S.: Quasi-neutral dynamics in a coinfection system with N strains and asymmetries along multiple traits. J. Math. Biol. 87, 48 (2023)
Cantrell, R.S., Cosner, C.: Spatial Ecology Via Reaction–Diffusion Equations. Wiley, New York (2004)
Allen, L.J.S., Bolker, B.M., Lou, Y., Nevai, A.L.: Asymptotic profiles of the steady states for an sis epidemic reaction–diffusion model. Discrete Contin. Dynam. . 21(1), 1–20 (2008)
Castella, F., Hoffbeck, J.-P., Lagadeuc, Y.: A reduced model for spatially structured predator-prey systems with fast spatial migrations and slow demographic evolutions. Asymptot. Anal. 61(3–4), 125–175 (2009). https://doi.org/10.3233/asy-2008-0905
Castella, F., Madec, S., Lagadeuc, Y.: Global behavior of n competing species with strong diffusion: diffusion leads to exclusion. Appl. Anal. 95(2), 341–372 (2016). https://doi.org/10.1080/00036811.2015.1004320
Bratus, A.S., Posvyanskii, V.P., Novozhilov, A.S.: Replicator equations and space. Math. Model. Nat. Phenom. 9(3), 47–67 (2014)
van Baalen, M., Sabelis, M.W.: The dynamics of multiple infection and the evolution of virulence. Am. Nat. 146, 881–910 (1995)
Hollis, S.L., Martin, R.H., Jr., Pierre, M.: Global existence and boundedness in reaction–diffusion systems. SIAM J. Math. Anal. 18(3), 744–761 (1987). https://doi.org/10.1137/0518057
Morgan, J.: Global existence for semilinear parabolic systems. SIAM J. Math. Anal. 20(5), 1128–1144 (1989). https://doi.org/10.1137/0520075
Pao, C.V.: Nonlinear Parabolic and Elliptic Equations. Springer, Berlin (1993)
Biegert, M.: The Neumann Laplacian on spaces of continuous functions. Note di Matematica 22(1), 65–74 (2003)
Pazy, A.: Semigroups of Linear Operators and Applications to Partial Differential Equations. Applied Mathematical Sciences, vol. 44. Springer, Berlin (1983)
Diekmann, O., Heesterbeek, J., Metz, J.: On the definition and the computation of the basic reproduction ratio r0 in models for infectious diseases in heterogeneous populations. J. Math. Biol. 28, 365–382 (1990)
Smoller, J.: Shock Waves and Reaction–Diffusion Equations. Grundlehren der mathematischen Wissenschaften, vol. 258. Springer, Berlin (1994)
Anderson, R.M., May, R.: Infectious Diseases of Humans: Dynamics and Control. Oxford University Press, Oxford (1991)
Dushoff, J., Levin, S.: The effects of population heterogeneity on disease invasion. Math. Biosci. 128(1–2), 25–40 (1995)
Lajmanovich, A., Yorke, J.: A deterministic model for gonorrhea in a nonhomogeneous population. Bellman Prize Math. Biosci. 28, 221–236 (1976)
Lloyd, A., May, R.: Spatial heterogeneity in epidemic models. J. Theor. Biol. 179(1), 1–11 (1996)
Mottoni, P., Orlandi, E., Tesei, A.: Asymptotic behavior for a system describing epidemics with migration and spatial spread of infection. Nonlinear Anal. Theory Methods Appl. 3, 663–675 (1979)
Pang, D., Xiao, Y.: The sis model with diffusion of virus in the environment. Math. Biosci. Eng. 16, 2852–2874 (2019)
Fitzgibbon, W., Langlais, M., Morgan, J.: A reaction–diffusion system modeling direct and indirect transmission of diseases. Discrete Contin. Dyn. Syst. Ser. B 4, 893–910 (2004)
Fitzgibbon, W., Langlais, M.: Simple models for the transmission of microparasites between host populations living on noncoincident spatial domains. In: Structured Population Models in Biology and Epidemiology (2008)
Ruan, S.: Spatial-temporal dynamics in nonlocal epidemiological models. In: Mathematics for Life Science and Medicine (2007)
Hofbauer, J., Sigmund, K.: Evolutionary Games and Population Dynamics. Cambridge University Press, London (1998)
Wu, Y., Zou, X.: Asymptotic profiles of steady states for a diffusive sis epidemic model with mass action infection mechanism. J. Differ. Equ. 261, 4424–4447 (2016)
Megretski, A.: Singular Perturbations and Averaging. Lecture Notes, Massachusetts Institute of Technology (2003)
Duan, G.-R., Patton, R.J.: A note on Hurwitz stability of matrices. Automatica 34, 509–511 (1998)
Acknowledgements
We would like to thank Professor Boris ANDREIANOV, Laboratory of Mathematics and Theoretical Physics, University of Tours. Professor Andreianov helped us with several techniques in the proof of Theorem 7.
Funding
We have no funding received for this paper.
Author information
Authors and Affiliations
Contributions
Two authors have the same contributions, including writing and revising this manuscript.
Corresponding author
Ethics declarations
Conflict of interest
We declare that the authors have no competing interests as defined by Springer, or other interests that might be perceived to influence the results and/or discussion reported in this paper.
Ethics approval
Not applicable
Consent to participate
Two authors consent to participate.
Consent for publication
Two authors consent to participate.
Additional information
Publisher's Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Appendices
Appendix A Proof for Theorem 7
The idea of our proof bases on the technique mentioned in [36].
Proof
Firstly, we make a convention for the norm using in this proof. For each \(t \in \mathbb {R}^+\), for every \(f_1, f_2 \in L^2\left( \Omega \times \mathbb {R},\mathbb {R}^n\right) \) we denote
where the \(f_1\cdot f_2\) representing for the usual scalar product \(\sum _{i=1}^{n}f^i_1 f^i_2\) in \(\mathbb {R}^n\). This scalar product \(\langle \cdot ,\cdot \rangle \) induces the norm
For the sake of convenience in this proof, we only write \(\left\| \cdot \right\| \) instead of \(\left\| \cdot \right\| _2\).
We do the same convention for \(\langle g_1,g_2 \rangle \) and \(\left\| g\left( \cdot ,t\right) \right\| \) for all \(g_1,g_2,g \in C^1\left( \Omega \times \mathbb {R},\mathbb {R}^m\right) \).
Because in the finite dimensional space, all norms are equivalent, we then denote \(|\cdot |\) to be the usual 2-Euclidean norm. Moreover, we recall the notation \(A \prec 0\) for a symmetric matrix A if A is definitely negative, and \(A \succ 0\) for definitely positive symmetric matrix.
First, let us show that the interval \([t_0,t_1]\) can be subdivided into subinterval \(\Delta _k = [\tau _{k-1},\tau _k]\), where \(k \in \{1,2,\dots ,N\}\) and \(t_0 = \tau _0< \tau _1< \dots <\tau _N = t_1\) in such a way that for every k, there exists a symmetric matrix \(P_k = P^T_k \succ 0\) for which
Indeed, since A(x, t) is a Hurwitz matrix for every \(t \in [t_0,t_1]\), according to [37], there exists \(P(x,t) = P^T(x,t) \succ 0\) such that
Since A depends continuously on t, there exists an open interval \(\Delta (t)\) such that \(t \in \Delta (t)\) and
Now the open intervals \(\Delta (t)\) with \(t \in \left[ t_0,t_1\right] \) cover the whole closed bounded interval \(\left[ t_0,t_1\right] \) and taking a finite number of \(\tau _k\), \(k=1,\dots ,N\) such that \(\left[ t_0,t_1\right] \) is completely covered by \(\Delta (\tau _k)\) yields the desired partition subdivision.
We can note that a strictly negative upper bound is not required on the real parts’ eigenvalues uniformly in space because the spatial domain is supposed to be compact.
Note that, from (A1), for all \(y \in \mathbb {R}^m\) we have that
Second, because F, G are continuously differential in x and t, then for every \(\mu >0\) there exists \(C,r >0\) such that
for all \(t \in \mathbb {R}\), \(\bar{\delta }_f\left( x,t\right) \in \mathbb {R}^n\), \(\bar{\delta }_g\left( x,t\right) \in \mathbb {R}^m\) satisfying
For the sake of simplicity, we write \(\bar{\delta }_f\) and \(\bar{\delta }_g\) instead of \(\bar{\delta }_f\left( x,t\right) \) and \(\bar{\delta }_g\left( x,t\right) \). We now have the Taylor expansion as follows, noting that \(G\left( f_0(x,t),g_0(x,t),x,t\right) = 0\),
with B(x, t) is the Jacobian matrix of \(G\left( \cdot ,\cdot ,t\right) \) with respect to the first variable.
For each \(k = 1,\dots , N\), and \(u \in \mathbb {R}^m\), set \(|u|_k = \left( u^T P_k u\right) ^{1/2}\), then \(|\cdot |_k\) is a norm in \(\mathbb {R}^m\). Indeed, because \(P_k \succ 0\) then \(|\cdot |_k\) is well-defined, it suffices to check the condition \(|u + v|_k \le |u|_k + |v|_k\), which is equivalent to
It now becomes
thanks to the Cholesky’s factorization, which states that, if \(P_k \succ 0\), there exist a square matrix such that \(P_k = L^T_kL_k\). Note that, (A5) holds because of the inequality Cauchy-Schwarz. Hence, \(|\cdot |_k\) is a norm in \(\mathbb {R}^m\) and it is equivalent to an arbitrary norm in \(\mathbb {R}^m\).
Then, for \(\delta _f(x,t) = f(x,t) - f_0(x,t)\), \(\delta _g(x,t) = g(x,t) - g_0(x,t)\), we have that
as long as \(\delta _f\), \(\delta _g\) are sufficiently small, where \(C_1\), q are positive constants which do not depend on k.
Initially, for the sake of simplicity, in the following arguments, we write f, g instead of \(f\left( x,t\right) \) and \(g\left( x,t\right) \), respectively. Then, we have the equation for \(\delta _f\left( x,t\right) \) as follows
By the convention of \(\left\| \cdot \right\| \), we have that
On the other hand, recalling that \(K_f = a_f\left( x\right) \nabla + \Delta \) implies
which leads to, when we apply the Young inequality for the term \(\int _{\Omega }a\left( x\right) \nabla \delta _f\cdot \delta _f dx\),
where \(\left| a_f\left( x\right) \right| \) is the matrix in which entries are absolute values of corresponding coordinates of \(a_f\left( x\right) \).
Accordingly, we have the estimation for \(\dfrac{d}{d t}\left\| \delta _f\right\| ^2\) as follows
Next, we come to control the growth of \(\left\| \delta _g\right\| _k\). We first observe that
We denote \(\epsilon \left[ K_g g_0\left( x,t\right) + \dfrac{\partial }{\partial t}g_0\left( x,t\right) + G_1(x)\cdot \nabla f_0\right] \) as \(O\left( \epsilon \right) \), then
Using the Taylor expansion for G in (A4) and the equation (A8), we obtain the following computations
Using the Young inequality, we have the estimation for \(\langle o\left( \left| \delta _f\right| \right) + o\left( \left| \delta _g\right| \right) + O\left( \epsilon \right) ,P\delta _g + \delta _g\rangle \) as follows
Alternatively, applying the Young inequality, we have that
since \(\nabla \delta _f\) is bounded in \(\Omega \).
For the term \(\langle K_g\delta _g, P\delta _g + \delta _g\rangle \), we get that
Note that \(P \succ 0\) then \(\int _{\Omega }\nabla \delta _gP \nabla \delta _gdx \ge \lambda \left\| \nabla \delta _g\right\| ^2\). Applying the Young inequality once more for the terms
we have that
which implies
with \(C\left( 1 + \lambda \right) \) denoting a constant depending on \(1 + \lambda \).
Combining these equations (A9), (A10), (A11), and (A12), and noting that two norms \(\left\| \cdot \right\| _k\) and \(\left\| \cdot \right\| _k\) are equivalent, we observe that
which implies when \(\epsilon \) small enough
Thus, combine (A7) and (A13) and we obtain that
for some constant \(C_1\) independent of k.
By the Gronwall’s inequality for \(\left( \left\| \delta _f\right\| ^2 + \frac{\epsilon C_1}{q}\left\| \delta _g\right\| _k^2\right) \), for each \(k \ge 1\), we can regard \(\tau _{k-1}\) as the initial value, and then deduce that
for \(\tau \in [0,\tau _k - \tau _{k-1}]\). With the aid of this bound for the growth of \(|\delta _f|\), the second inequality of (A6) implies a bound for \(\left\| \delta _g\right\| _k\) as following
We already have that \(\delta _f\left( x,t_0\right) = \delta _f\left( x,\tau _0\right) \le \epsilon \) and \(\delta _g\left( x,t_0\right) = \delta _g\left( x,\tau _0\right) \le \epsilon _0\) for \(\epsilon _0\) small enough. Then, by the compactness of \(\bar{\Omega }\), for \(\tau \in \left[ 0,\tau _1 - \tau _0\right] \), \(\left\| \delta _f\left( \tau \right) \right\| ^2 \le O\left( \epsilon \right) \), for all \(x \in \Omega \). Make a process similarly and successively for \(k = 1,2,\dots \), we have that \(\left\| \delta _f\right\| ^2 \le O\left( \epsilon \right) \) for all \(x\in \Omega \). Analogously, we can also prove that \(\left\| \delta _g\right\| ^2 \le O\left( \epsilon \right) \).
Therefore, \(\int _{\Omega }\left| f(x,t) - f_0(x,t)\right| ^2dx \le C\epsilon \) and \(\int _{\Omega }|g(x,t) - g_0(x,t)|^2dx \le C\epsilon \), and we have the conclusion of the theorem. \(\square \)
Appendix B Proof for theorem 8
Proof
Note that \(\left\| F\left( u_1,x\right) - F\left( u_2,x\right) \right\| \le C\left\| u_1 - u_2\right\| ,\forall u_1,u_2 \in D\left( F\right) \) and \(|G\left( u,x\right) v|\) is bounded, \(\forall u,v\) bounded due to the continuous differentiability of G in a bounded domain. Consider
Taking the integral of (B1) over \(\Omega \) and using the Neumann boundary condition implies that
which leads to
Applying Gronwall’s inequality, and we have that
which implies \(\int _{\Omega } |u -v|^2dx = O\left( \epsilon \right) \) for all \(t < \mathcal {T}\) with given \(\mathcal {T} > 0\), by the compactness of \(\bar{\Omega }\). \(\square \)
Rights and permissions
Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.
About this article
Cite this article
Le, T.M.T., Madec, S. Spatiotemporal Evolution of Coinfection Dynamics: A Reaction–Diffusion Model. J Dyn Diff Equat (2023). https://doi.org/10.1007/s10884-023-10285-z
Received:
Revised:
Accepted:
Published:
DOI: https://doi.org/10.1007/s10884-023-10285-z