- Open Access
Phase separation and coexistence in spatial coordination games between microbes
Phys. Rev. E 113, 014402 – Published 22 January, 2026
DOI: https://doi.org/10.1103/6fqz-fhjr
Abstract
Dense, microbial communities are shaped by local interactions between cells. Both the nature of interactions, spanning antagonistic to cooperative, and the strength of interactions vary between and across microbial species and strains. These local interactions can influence the emergence and maintenance of microbial diversity. However, it remains challenging to link features of local interactions with spatially mediated coexistence dynamics given the significant variation in the microscopic mechanisms involved in cell-to-cell feedback. Here, we explore how microbial interactions over a broad range of ecological contexts spanning antagonism to cooperation can enable coexistence as spatially explicit domains emerge. To do so, we introduce and analyze a family of stochastic coordination games, where individuals do better when playing (i.e., interacting) with individuals of the same type than when playing with individuals of a different type. Using this game-theoretic framework, we show that the population dynamics for coordination games is governed by a double-well shaped interaction potential. We find that in a spatial setting this double-well potential induces phase separation, facilitating coexistence. Moreover, we show that for microbes engaged in symmetric coordination games, phase separation takes on a universal scaling that follows “Model A” coarsening, consistent with prior experimental observations for Vibrio cholerae mutual killers. Finally, we derive a partial differential equation equivalent of the spatial stochastic game, confirming both the double-well nature of spatial coordination games and the universality of phase separation. Altogether, this work extends prior findings on the link between microbial interactions and population structure and suggests generic mechanisms embedded in local interactions that can enable coexistence.
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References (41)
- K. R. Foster and T. Bell, Competition, not cooperation, dominates interactions among culturable microbial species, Curr. Biol. 22, 1845 (2012).
- K. Z. Coyte, J. Schluter, and K. R. Foster, The ecology of the microbiome: Networks, competition, and stability, Science 350, 663 (2015).
- C. D. Nadell, K. Drescher, and K. R. Foster, Spatial structure, cooperation and competition in biofilms, Nat. Rev. Microbiol. 14, 589 (2016).
- S. Mukherjee and B. L. Bassler, Bacterial quorum sensing in complex and dynamically changing environments, Nat. Rev. Microbiol. 17, 371 (2019).
- L. Eigentler, F. A. Davidson, and N. R. Stanley-Wall, Mechanisms driving spatial distribution of residents in colony biofilms: An interdisciplinary perspective, Open Biol. 12, 220194 (2022).
- M. E. Hibbing, C. Fuqua, M. R. Parsek, and S. B. Peterson, Bacterial competition: Surviving and thriving in the microbial jungle, Nat. Rev. Microbiol. 8, 15 (2010).
- A. Dal Co, S. van Vliet, D. J. Kiviet, S. Schlegel, and M. Ackermann, Short-range interactions govern the dynamics and functions of microbial communities, Nat. Ecol. Evol. 4, 366 (2020).
- A. Lobanov, S. Dyckman, H. Kurkjian, and B. Momeni, Spatial structure favors microbial coexistence except when slower mediator diffusion weakens interactions, eLife 12, e82504 (2023).
- A. Swain, L. Fussell, and W. F. Fagan, Higher-order effects, continuous species interactions, and trait evolution shape microbial spatial dynamics, Proc. Natl. Acad. Sci. USA 119, e2020956119 (2022).
- J. A. Orr, D. W. Armitage, and A. D. Letten, Coexistence theory for microbial ecology, and vice versa, Environ. Microbiol. 27, e70072 (2025).
- M. O. Lavrentovich and D. R. Nelson, Asymmetric mutualism in two- and three-dimensional range expansions, Phys. Rev. Lett. 112, 138102 (2014).
- D. Yanni, P. Márquez-Zacarías, P. J. Yunker, and W. C. Ratcliff, Drivers of spatial structure in social microbial communities, Curr. Biol. 29, R545 (2019).
- M. K. Denk-Lobnig and K. B. Wood, Spatial population dynamics of bacterial colonies with social antibiotic resistance, Proc. Natl. Acad. Sci. USA 122, e2417065122 (2025).
- L. McNally, E. Bernardy, J. Thomas, A. Kalziqi, J. Pentz, S. P. Brown, B. K. Hammer, P. J. Yunker, and W. C. Ratcliff, Killing by type VI secretion drives genetic phase separation and correlates with increased cooperation, Nat. Commun. 8, 14371 (2017).
- G. Steinbach, C. Crisan, S. L. Ng, B. K. Hammer, and P. J. Yunker, Accumulation of dead cells from contact killing facilitates coexistence in bacterial biofilms, J. R. Soc. Interface. 17, 20200486 (2020).
- S. M. Allen and J. W. Cahn, A microscopic theory for antiphase boundary motion and its application to antiphase domain coarsening, Acta Metall. 27, 1085 (1979).
- J. Hofbauer and K. Sigmund, Evolutionary game dynamics, Bull. Am. Math. Soc. 40, 479 (2003).
- M. A. Nowak and R. M. May, Evolutionary games and spatial chaos, Nature (London) 359, 826 (1992).
- C. Hauert and M. Doebeli, Spatial structure often inhibits the evolution of cooperation in the snowdrift game, Nature (London) 428, 643 (2004).
- M. Jusup, P. Holme, K. Kanazawa, M. Takayasu, I. Romić, Z. Wang, S. Geček, T. Lipić, B. Podobnik, L. Wang, et al., Social physics, Phys. Rep. 948, 1 (2022).
- M. Mobilia, I. T. Georgiev, and U. C. Täuber, Fluctuations and correlations in lattice models for predator-prey interaction, Phys. Rev. E 73, 040903(R) (2006).
- Y.-H. Lin and J. S. Weitz, Spatial interactions and oscillatory tragedies of the commons, Phys. Rev. Lett. 122, 148102 (2019).
- R. Durrett, Ten lectures on particle systems, Lectures on Probability Theory (Springer, Berlin, 1995), pp. 97–201.
- R. Durrett and S. Levin, The importance of being discrete (and spatial), Theor. Popul Biol. 46, 363 (1994).
- R. V. Kohn, F. Otto, M. G. Reznikoff, and E. Vanden-Eijnden, Action minimization and sharp-interface limits for the stochastic Allen-Cahn equation, Commun. Pure Appl. Math. 60, 393 (2007).
- M. A. Peletier, Variational modelling: Energies, gradient flows, and large deviations, arXiv:1402.1990.
- S. M. Kodera, P. Das, J. A. Gilbert, and H. L. Lutz, Conceptual strategies for characterizing interactions in microbial communities, iScience 25, 103775 (2022).
- J. D. Palmer and K. R. Foster, Bacterial species rarely work together, Science 376, 581 (2022).
- J. S. Weitz, C. Eksin, K. Paarporn, S. P. Brown, and W. C. Ratcliff, An oscillating tragedy of the commons in replicator dynamics with game-environment feedback, Proc. Natl. Acad. Sci. USA 113, E7518 (2016).
- A. R. Tilman, J. B. Plotkin, and E. Akçay, Evolutionary games with environmental feedbacks, Nat. Commun. 11, 915 (2020).
- T. Yao and D. B. Cooney, Spatial pattern formation in eco-evolutionary games with environment-driven motion, arXiv:2502.06723.
- F. A. Gorter, M. Manhart, and M. Ackermann, Understanding the evolution of interspecies interactions in microbial communities, Phil. Trans. R. Soc. B 375, 20190256 (2020).
- D. B. Borenstein, P. Ringel, M. Basler, and N. S. Wingreen, Established microbial colonies can survive type VI secretion assault, PLoS Comput. Biol. 11, e1004520 (2015).
- C.-Y. Chang, D. Bajić, J. C. C. Vila, S. Estrela, and A. Sanchez, Emergent coexistence in multispecies microbial communities, Science 381, 343 (2023).
- C. Ratzke, J. Barrere, and J. Gore, Strength of species interactions determines biodiversity and stability in microbial communities, Nat. Ecol. Evol. 4, 376 (2020).
- G. Li and G. Steinbach, Code repository for “Phase separation and coexistence in spatial coordination games between microbes,” GitHub (2025), https://github.com/Gabi-Steinbach/Spatial-microbial-games.
- G. Li and G. Steinbach, Code repository for “Phase separation and coexistence in spatial coordination games between microbes,” Zenodo (2025), https://zenodo.org/records/17039082.
- D. F. Anderson and T. G. Kurtz, Continuous time Markov chain models for chemical reaction networks, Design and Analysis of Biomolecular Circuits (Springer, Berlin, 2011), pp. 3–42.
- N. G. Van Kampen, Stochastic Processes in Physics and Chemistry (Elsevier, Amsterdam, 1992), Vol. 1.
- S. M. Krone, Spatial models: Stochastic and deterministic, Math. Comput. Modell. 40, 393 (2004).
- A. De Masi, P. A. Ferrari, and J. L. Lebowitz, Reaction-diffusion equations for interacting particle systems, J. Stat. Phys. 44, 589 (1986).