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Spatiotemporal Organization of Chemical Oscillators via Phase Separation
Phys. Rev. Lett. 136, 178001 – Published 28 April, 2026
DOI: https://doi.org/10.1103/shz6-7fj9
Abstract
We develop a method for studying chemical oscillators in the presence of phase separation. Specifically, we define a dynamics at phase equilibrium by imposing timescale separation between slow reactions and fast diffusion. We show that colocalization of components can alter oscillator frequency and amplitude, and that it determines the stability of oscillations and fixed points. Although our method applies to general reaction networks with phase separation, we illustrate it on a concrete example of a three-component oscillator (“rock-paper-scissors” model) with two-phase coexistence. The analysis is validated with a spatial model, where relaxing the timescale separation between reactions and diffusion leads to waves of phase equilibria at mesoscopic scales.
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References (76)
- C. P. Brangwynne, C. R. Eckmann, D. S. Courson, A. Rybarska, C. Hoege, J. Gharakhani, F. Jülicher, and A. A. Hyman, Science 324, 1729 (2009).
- S. F. Banani, H. O. Lee, A. A. Hyman, and M. K. Rosen, Nat. Rev. Mol. Cell Biol. 18, 285 (2017).
- S. Boeynaems, S. Alberti, N. L. Fawzi, T. Mittag, M. Polymenidou, F. Rousseau, J. Schymkowitz, J. Shorter, B. Wolozin, L. Van Den Bosch, P. Tompa, and M. Fuxreiter, Trends Cell Biol. 28, 420 (2018).
- A. A. Hyman, C. A. Weber, and F. Jülicher, Annu. Rev. Cell Dev. Biol. 30, 39 (2014).
- D. M. Mitrea and R. W. Kriwacki, Cell Commun. Signaling 14, 1 (2016).
- C. A. Strulson, R. C. Molden, C. D. Keating, and P. C. Bevilacqua, Nat. Chem. 4, 941 (2012).
- K. K. Nakashima, M. A. Vibhute, and E. Spruijt, Front. Mol. Biosci. 6, 21 (2019).
- C. A. Weber, D. Zwicker, F. Jülicher, and C. F. Lee, Rep. Prog. Phys. 82, 064601 (2019).
- A. Oparin, The Origin of Life (Dover Publications, New York, 1953).
- J. B. S. Haldane, Rationalist Annu. 148, 3 (1929).
- Q.-X. Liu, A. Doelman, V. Rottschäfer, M. de Jager, P. M. Herman, M. Rietkerk, and J. van de Koppel, Proc. Natl. Acad. Sci. U.S.A. 110, 11905 (2013).
- K. Siteur, Q.-X. Liu, V. Rottschäfer, T. van der Heide, M. Rietkerk, A. Doelman, C. Boström, and J. van de Koppel, Proc. Natl. Acad. Sci. U.S.A. 120, e2202683120 (2023).
- T. Aslyamov, F. Avanzini, É. Fodor, and M. Esposito, Phys. Rev. Lett. 131, 138301 (2023).
- F. Avanzini, T. Aslyamov, É. Fodor, and M. Esposito, J. Chem. Phys. 161, 174108 (2024).
- J. Kirschbaum and D. Zwicker, J. R. Soc. Interface 18, 20210255 (2021).
- Y. Cho and W. M. Jacobs, J. Chem. Phys. 159, 154101 (2023).
- S. Laha, J. Bauermann, F. Jülicher, T. C. T. Michaels, and C. A. Weber, Phys. Rev. Res. 6, 043092 (2024).
- S. C. Glotzer, E. A. Di Marzio, and M. Muthukumar, Phys. Rev. Lett. 74, 2034 (1995).
- D. Zwicker, A. A. Hyman, and F. Jülicher, Phys. Rev. E 92, 012317 (2015).
- J. D. Wurtz and C. F. Lee, Phys. Rev. Lett. 120, 078102 (2018).
- A. Kumar and S. A. Safran, Phys. Rev. Lett. 131, 258401 (2023).
- J. Bauermann, G. Bartolucci, C. A. Weber, and F. Jülicher, Phys. Rev. Lett. 135, 148201 (2025).
- N. Ziethen, J. Kirschbaum, and D. Zwicker, Phys. Rev. Lett. 130, 248201 (2023).
- Y. Cho and W. M. Jacobs, Phys. Rev. Lett. 130, 128203 (2023).
- D. Zwicker, R. Seyboldt, C. A. Weber, A. A. Hyman, and F. Jülicher, Nat. Phys. 13, 408 (2017).
- J. Bauermann, C. A. Weber, and F. Jülicher, Ann. Phys. (Amsterdam) 534, 2200132 (2022).
- L. Demarchi, A. Goychuk, I. Maryshev, and E. Frey, Phys. Rev. Lett. 130, 128401 (2023).
- G. Häfner and M. Müller, ACS Nano 18, 16530 (2024).
- A. Goychuk, arXiv:2506.07753.
- Y. Zhuang, Z. Li, S. Xiong, C. Sun, B. Li, S. A. Wu, J. Lyu, X. Shi, L. Yang, Y. Chen et al., Cell 186, 3245 (2023).
- D. Tariq, N. Maurici, B. M. Bartholomai, S. Chandrasekaran, J. C. Dunlap, A. Bah, and B. R. Crane, eLife 12, RP90259 (2024).
- M. S. Heltberg, A. Lucchetti, F.-S. Hsieh, D. P. M. Nguyen, S.-h. Chen, and M. H. Jensen, Cell 185, 4394 (2022).
- I. S. Haugerud, H. D. Vuijk, J. Boekhoven, and C. A. Weber, arXiv:2503.11604.
- J. Sastre, A. Thatte, A. M. Bergmann, M. Stasi, M. Tena-Solsona, C. A. Weber, and J. Boekhoven, Nat. Commun. 16, 2003 (2025).
- C. Luo and D. Zwicker, Phys. Rev. E 108, 034206 (2023).
- I. B. A. Smokers, B. S. Visser, W. P. Lipiński, K. K. Nakashima, and E. Spruijt, ChemSystemsChem 7, e202400056 (2024).
- R. M. May and W. J. Leonard, SIAM J. Appl. Math. 29, 243 (1975).
- J. Hofbauer and K. Sigmund, Evolutionary Games and Population Dynamics (Cambridge University Press, Cambridge, England, 1998).
- J. M. Smith, Evolution and the Theory of Games (Cambridge University Press, Cambridge, England, 2012).
- T. Reichenbach, M. Mobilia, and E. Frey, Nature (London) 448, 1046 (2007).
- M. Mobilia, J. Theor. Biol. 264, 11 (2010).
- G. Szabó and G. Fáth, Phys. Rep. 446, 97 (2007).
- B. Sinervo and C. M. Lively, Nature (London) 380, 240 (1996).
- B. Kerr, M. A. Riley, M. W. Feldman, and B. J. M. Bohannan, Nature (London) 418, 171 (2002).
- M. J. Liao, M. O. Din, L. Tsimring, and J. Hasty, Science 365, 1045 (2019).
- S. C. Takatori and J. F. Brady, Phys. Rev. E 91, 032117 (2015).
- M. E. Cates and J. Tailleur, Annu. Rev. Condens. Matter Phys. 6, 219 (2015).
- A. P. Solon, J. Stenhammar, M. E. Cates, Y. Kafri, and J. Tailleur, Phys. Rev. E 97, 020602(R) (2018).
- N. Srinivas, J. Parkin, G. Seelig, E. Winfree, and D. Soloveichik, Science 358, eaal2052 (2017).
- See Supplemental Material at http://link.aps.org/supplemental/10.1103/shz6-7fj9 for details.
- J. W. Cahn and J. E. Hilliard, J. Chem. Phys. 28, 258 (1958).
- G. I. Tóth, T. Pusztai, and L. Gránásy, Phys. Rev. B 92, 184105 (2015).
- G. I. Tóth, M. Zarifi, and B. Kvamme, Phys. Rev. E 93, 013126 (2016).
- L. Onsager, Phys. Rev. 37, 405 (1931).
- S. R. De Groot and P. Mazur, Non-Equilibrium Thermodynamics, Dover Books on Physics (Dover Publications, Mineola, NY, 2003).
- E. J. Kramer, P. Green, and C. J. Palmstrøm, Polymer 25, 473 (1984).
- S. Bo, L. Hubatsch, J. Bauermann, C. A. Weber, and F. Jülicher, Phys. Rev. Res. 3, 043150 (2021).
- T. Reichenbach, M. Mobilia, and E. Frey, J. Theor. Biol., 254 368 (2008).
- Q. He, M. Mobilia, and U. C. Täuber, Phys. Rev. E 82, 051909 (2010).
- S. Safran, Statistical Thermodynamics of Surfaces, Interfaces, and Membranes (CRC Press, London, 2019).
- M. Kardar, Statistical Physics of Fields (Cambridge University Press, Cambridge, England, 2007).
- J. Bauermann, S. Laha, P. M. McCall, F. Jülicher, and C. A. Weber, J. Am. Chem. Soc. 144, 19294 (2022).
- M. Peltomäki and M. Alava, Phys. Rev. E 78, 031906 (2008).
- B. Szczesny, M. Mobilia, and A. M. Rucklidge, Phys. Rev. E 90, 032704 (2014).
- F. Pedregosa, G. Varoquaux, A. Gramfort, V. Michel, B. Thirion, O. Grisel, M. Blondel, P. Prettenhofer, R. Weiss, V. Dubourg, J. Vanderplas, A. Passos, D. Cournapeau, M. Brucher, M. Perrot, and E. Duchesnay, J. Mach. Learn. Res. 12, 2825 (2011).
- J. Halatek and E. Frey, Nat. Phys. 14, 507 (2018).
- J. Halatek, F. Brauns, and E. Frey, Phil. Trans. R. Soc. B 373, 20170107 (2018).
- A. W. Fritsch, A. F. Diaz-Delgadillo, O. Adame-Arana, C. Hoege, M. Mittasch, M. Kreysing, M. Leaver, A. A. Hyman, F. Jülicher, and C. A. Weber, Proc. Natl. Acad. Sci. U.S.A. 118, e2102772118 (2021).
- J. F. Robinson, T. Machon, and T. Speck, Phys. Rev. E 111, 065417 (2025).
- M. Tateno and O. A. Saleh, Phys. Rev. Lett. 136, 068403 (2026).
- J. Bauermann, G. Bartolucci, and A. Kolchinsky, Data for reproducing the figures of the paper: “Spatiotemporal organization of chemical oscillators via phase separation” (2026), 10.5281/zenodo.18990195.
- S. H. Strogatz, Nonlinear Dynamics and Chaos (CRC Press, London, 2018).
- D. Deviri and S. A. Safran, Proc. Natl. Acad. Sci. U.S.A. 118, e2100099118 (2021).
- C. Zechner and F. Jülicher, Cell Syst. 16, 101168 (2025).
- A. Klosin, F. Oltsch, T. Harmon, A. Honigmann, F. Jülicher, A. A. Hyman, and C. Zechner, Science 367, 464 (2020).
- A. Bray, Adv. Phys. 43, 357 (1994).