Abstract
We propose a model of chemostat where the bacterial population is individually-based, each bacterium is explicitly represented and has a mass evolving continuously over time. The substrate concentration is represented as a conventional ordinary differential equation. These two components are coupled with the bacterial consumption. Mechanisms acting on the bacteria are explicitly described (growth, division and washout). Bacteria interact via consumption. We set the exact Monte Carlo simulation algorithm of this model and its mathematical representation as a stochastic process. We prove the convergence of this process to the solution of an integro-differential equation when the population size tends to infinity. Finally, we propose several numerical simulations.
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Notes
Note that our situation is simpler than that studied by Roelly-Coppoletta [26] and Mard and Roelly[19] since in our case \({\mathcal {X}}\) is compact: in fact in our case the weak topology—the smallest topology which makes the applications \(\nu \rightarrow \langle \nu ,f \rangle \) continuous for any \(f\) continuous and bounded—and the vague topology—the smallest topology which makes the applications \(\nu \rightarrow \langle \nu ,f \rangle \) continuous for all \(f\) continuous with compact support—are identical.
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Acknowledgments
The authors are grateful to Jme Harmand and Claude Lobry for discussions on the model, to Pierre Pudlo and Pascal Neveu for their help concerning the programming of the IBM. This work is partially supported by the project “Modles Numriques pour les cosystmes Microbiens” of the French National Network of Complex Systems (RNSC call 2012). The work of Coralie Fritsch is partially supported by the Meta-omics of Microbial Ecosystems (MEM) metaprogram of INRA.
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Fabien Campillo and Coralie Fritsch are members of the MODEMIC joint INRA and INRIA project-team.
Appendix: Skorohod Topology
Appendix: Skorohod Topology
The space of finite measures \({\mathcal {M}}_{F}(\mathcal {X})\) on \({\mathcal {X}}\) is equipped with the topology of the weak convergence, that is the smallest topology for which the applications \(\zeta \rightarrow \langle \zeta ,f \rangle =\int _{{\mathcal {X}}}f(x)\,\zeta ({\mathrm{d }}x)\) are continuous for any \(f\in \mathcal {C}(\mathcal {X})\). This topology is metrized by the Prokhorov metric:
where \(F^\epsilon \mathop {=}\limits ^\mathrm{\tiny def }\{x\in {\mathcal {X}};\inf _{y\in F}|x-y|<\epsilon \}\) (see [7, Appendix A2.5]). The Prokhorov distance is bounded by the distance of the total variation \(d_\mathrm{\tiny \mathrm TV }(\zeta ,\zeta ')=\Vert \zeta -\zeta ' \Vert _\mathrm{\tiny TV }\) associated with the norm defined by:
for any finite and signed measure \(\zeta \) where \(\zeta =\zeta _{+}-\zeta _{-}\) is the Hahn–Jordan decomposition of \(\zeta \).
The space \(\mathcal D([0,T],{\mathcal {M}}_{F}(\mathcal {X}))\) is equipped with the Skorohod metric \(d_\mathrm{\tiny \mathrm S }\). Instead of giving the definition of this metric [12, Eq. (5.2) p. 117] we recall a characterization of the convergence for this metric given in [12, Proposition 5.3, p. 119].
A sequence \((\zeta ^{n})_{n\in \mathbb N}\) converges to \(\zeta \) in \({\mathcal {D}}([0,T],{\mathcal {M}}_{F}(\mathcal {X}))\), i.e. \(d_\mathrm{\tiny \mathrm S }(\zeta ^{n},\zeta )\rightarrow 0\), if and only if there exists a sequence \(\lambda _{n}(t)\) of time change functions (i.e. strictly increasing bijective functions on \([0,T]\), with \(\lambda _{n}(0)=0\) and \(\lambda _{n}(T)=T\)) satisfying:
and
If \((\zeta ^{n})_{n\in \mathbb N}\) converges to \(\zeta \) in \({\mathcal {D}}([0,T],{\mathcal {M}}_{F}(\mathcal {X}))\) and if \(\zeta \in {\mathcal {C}}([0,T],{\mathcal {M}}_{F}({\mathcal {X}}))\) then in:
the first term of the right-hand side tends to 0 because of (25); the second one tends to 0 because of (26) and the uniform continuity of \(\zeta \) in \([0,T]\). This proves that \(\zeta ^{n}\) converges to \(\zeta \) in \({\mathcal {D}}([0,T],{\mathcal {M}}_{F}(\mathcal {X}))\) also for the uniform metric.
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Campillo, F., Fritsch, C. Weak Convergence of a Mass-Structured Individual-Based Model. Appl Math Optim 72, 37–73 (2015). https://doi.org/10.1007/s00245-014-9271-3
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DOI: https://doi.org/10.1007/s00245-014-9271-3
Keywords
- Individually-based model
- Integro-differential equation
- Weak convergence of Markov processes
- Mass-structured chemostat model
- Monte Carlo