Reuse & Permissions

It is not necessary to obtain permission to reuse this article or its components as it is available under the terms of the Creative Commons Attribution 4.0 International license. This license permits unrestricted use, distribution, and reproduction in any medium, provided attribution to the author(s) and the published article's title, journal citation, and DOI are maintained. Please note that some figures may have been included with permission from other third parties. It is your responsibility to obtain the proper permission from the rights holder directly for these figures.

Export citation

Export citation

Choose format for download:

Download Citation
  • Open Access

Spatiotemporal Organization of Chemical Oscillators via Phase Separation

Jonathan Bauermann1,*, Giacomo Bartolucci2,3, and Artemy Kolchinsky4,5,6

  • 1Department of Physics, Harvard University, Cambridge, Massachusetts 02138, USA
  • 2Department of Condensed Matter Physics, Universitat de Barcelona, 08007 Barcelona, Spain
  • 3Universitat de Barcelona Institute of Complex Systems (UBICS), Universitat de Barcelona, 08028 Barcelona, Spain
  • 4ICREA-Complex Systems Lab, Universitat Pompeu Fabra, 08003 Barcelona, Spain
  • 5Barcelona Collaboratorium for Modelling and Predictive Biology, Wellington 30, 08005 Barcelona, Spain
  • 6Universal Biology Institute, University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan

  • *Contact author: jbauermann@fas.harvard.edu

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.

View figure in article

Physics Subject Headings (PhySH)

Article Text

Supplemental Material

References (76)

  1. 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).
  2. S. F. Banani, H. O. Lee, A. A. Hyman, and M. K. Rosen, Nat. Rev. Mol. Cell Biol. 18, 285 (2017).
  3. 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).
  4. A. A. Hyman, C. A. Weber, and F. Jülicher, Annu. Rev. Cell Dev. Biol. 30, 39 (2014).
  5. D. M. Mitrea and R. W. Kriwacki, Cell Commun. Signaling 14, 1 (2016).
  6. C. A. Strulson, R. C. Molden, C. D. Keating, and P. C. Bevilacqua, Nat. Chem. 4, 941 (2012).
  7. K. K. Nakashima, M. A. Vibhute, and E. Spruijt, Front. Mol. Biosci. 6, 21 (2019).
  8. C. A. Weber, D. Zwicker, F. Jülicher, and C. F. Lee, Rep. Prog. Phys. 82, 064601 (2019).
  9. A. Oparin, The Origin of Life (Dover Publications, New York, 1953).
  10. J. B. S. Haldane, Rationalist Annu. 148, 3 (1929).
  11. 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).
  12. 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).
  13. T. Aslyamov, F. Avanzini, É. Fodor, and M. Esposito, Phys. Rev. Lett. 131, 138301 (2023).
  14. F. Avanzini, T. Aslyamov, É. Fodor, and M. Esposito, J. Chem. Phys. 161, 174108 (2024).
  15. J. Kirschbaum and D. Zwicker, J. R. Soc. Interface 18, 20210255 (2021).
  16. Y. Cho and W. M. Jacobs, J. Chem. Phys. 159, 154101 (2023).
  17. S. Laha, J. Bauermann, F. Jülicher, T. C. T. Michaels, and C. A. Weber, Phys. Rev. Res. 6, 043092 (2024).
  18. S. C. Glotzer, E. A. Di Marzio, and M. Muthukumar, Phys. Rev. Lett. 74, 2034 (1995).
  19. D. Zwicker, A. A. Hyman, and F. Jülicher, Phys. Rev. E 92, 012317 (2015).
  20. J. D. Wurtz and C. F. Lee, Phys. Rev. Lett. 120, 078102 (2018).
  21. A. Kumar and S. A. Safran, Phys. Rev. Lett. 131, 258401 (2023).
  22. J. Bauermann, G. Bartolucci, C. A. Weber, and F. Jülicher, Phys. Rev. Lett. 135, 148201 (2025).
  23. N. Ziethen, J. Kirschbaum, and D. Zwicker, Phys. Rev. Lett. 130, 248201 (2023).
  24. Y. Cho and W. M. Jacobs, Phys. Rev. Lett. 130, 128203 (2023).
  25. D. Zwicker, R. Seyboldt, C. A. Weber, A. A. Hyman, and F. Jülicher, Nat. Phys. 13, 408 (2017).
  26. J. Bauermann, C. A. Weber, and F. Jülicher, Ann. Phys. (Amsterdam) 534, 2200132 (2022).
  27. L. Demarchi, A. Goychuk, I. Maryshev, and E. Frey, Phys. Rev. Lett. 130, 128401 (2023).
  28. G. Häfner and M. Müller, ACS Nano 18, 16530 (2024).
  29. A. Goychuk, arXiv:2506.07753.
  30. 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).
  31. D. Tariq, N. Maurici, B. M. Bartholomai, S. Chandrasekaran, J. C. Dunlap, A. Bah, and B. R. Crane, eLife 12, RP90259 (2024).
  32. M. S. Heltberg, A. Lucchetti, F.-S. Hsieh, D. P. M. Nguyen, S.-h. Chen, and M. H. Jensen, Cell 185, 4394 (2022).
  33. I. S. Haugerud, H. D. Vuijk, J. Boekhoven, and C. A. Weber, arXiv:2503.11604.
  34. J. Sastre, A. Thatte, A. M. Bergmann, M. Stasi, M. Tena-Solsona, C. A. Weber, and J. Boekhoven, Nat. Commun. 16, 2003 (2025).
  35. C. Luo and D. Zwicker, Phys. Rev. E 108, 034206 (2023).
  36. I. B. A. Smokers, B. S. Visser, W. P. Lipiński, K. K. Nakashima, and E. Spruijt, ChemSystemsChem 7, e202400056 (2024).
  37. R. M. May and W. J. Leonard, SIAM J. Appl. Math. 29, 243 (1975).
  38. J. Hofbauer and K. Sigmund, Evolutionary Games and Population Dynamics (Cambridge University Press, Cambridge, England, 1998).
  39. J. M. Smith, Evolution and the Theory of Games (Cambridge University Press, Cambridge, England, 2012).
  40. T. Reichenbach, M. Mobilia, and E. Frey, Nature (London) 448, 1046 (2007).
  41. M. Mobilia, J. Theor. Biol. 264, 11 (2010).
  42. G. Szabó and G. Fáth, Phys. Rep. 446, 97 (2007).
  43. B. Sinervo and C. M. Lively, Nature (London) 380, 240 (1996).
  44. B. Kerr, M. A. Riley, M. W. Feldman, and B. J. M. Bohannan, Nature (London) 418, 171 (2002).
  45. M. J. Liao, M. O. Din, L. Tsimring, and J. Hasty, Science 365, 1045 (2019).
  46. S. C. Takatori and J. F. Brady, Phys. Rev. E 91, 032117 (2015).
  47. M. E. Cates and J. Tailleur, Annu. Rev. Condens. Matter Phys. 6, 219 (2015).
  48. A. P. Solon, J. Stenhammar, M. E. Cates, Y. Kafri, and J. Tailleur, Phys. Rev. E 97, 020602(R) (2018).
  49. N. Srinivas, J. Parkin, G. Seelig, E. Winfree, and D. Soloveichik, Science 358, eaal2052 (2017).
  50. See Supplemental Material at http://link.aps.org/supplemental/10.1103/shz6-7fj9 for details.
  51. J. W. Cahn and J. E. Hilliard, J. Chem. Phys. 28, 258 (1958).
  52. G. I. Tóth, T. Pusztai, and L. Gránásy, Phys. Rev. B 92, 184105 (2015).
  53. G. I. Tóth, M. Zarifi, and B. Kvamme, Phys. Rev. E 93, 013126 (2016).
  54. L. Onsager, Phys. Rev. 37, 405 (1931).
  55. S. R. De Groot and P. Mazur, Non-Equilibrium Thermodynamics, Dover Books on Physics (Dover Publications, Mineola, NY, 2003).
  56. E. J. Kramer, P. Green, and C. J. Palmstrøm, Polymer 25, 473 (1984).
  57. S. Bo, L. Hubatsch, J. Bauermann, C. A. Weber, and F. Jülicher, Phys. Rev. Res. 3, 043150 (2021).
  58. T. Reichenbach, M. Mobilia, and E. Frey, J. Theor. Biol., 254 368 (2008).
  59. Q. He, M. Mobilia, and U. C. Täuber, Phys. Rev. E 82, 051909 (2010).
  60. S. Safran, Statistical Thermodynamics of Surfaces, Interfaces, and Membranes (CRC Press, London, 2019).
  61. M. Kardar, Statistical Physics of Fields (Cambridge University Press, Cambridge, England, 2007).
  62. J. Bauermann, S. Laha, P. M. McCall, F. Jülicher, and C. A. Weber, J. Am. Chem. Soc. 144, 19294 (2022).
  63. M. Peltomäki and M. Alava, Phys. Rev. E 78, 031906 (2008).
  64. B. Szczesny, M. Mobilia, and A. M. Rucklidge, Phys. Rev. E 90, 032704 (2014).
  65. 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).
  66. J. Halatek and E. Frey, Nat. Phys. 14, 507 (2018).
  67. J. Halatek, F. Brauns, and E. Frey, Phil. Trans. R. Soc. B 373, 20170107 (2018).
  68. 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).
  69. J. F. Robinson, T. Machon, and T. Speck, Phys. Rev. E 111, 065417 (2025).
  70. M. Tateno and O. A. Saleh, Phys. Rev. Lett. 136, 068403 (2026).
  71. 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.
  72. S. H. Strogatz, Nonlinear Dynamics and Chaos (CRC Press, London, 2018).
  73. D. Deviri and S. A. Safran, Proc. Natl. Acad. Sci. U.S.A. 118, e2100099118 (2021).
  74. C. Zechner and F. Jülicher, Cell Syst. 16, 101168 (2025).
  75. A. Klosin, F. Oltsch, T. Harmon, A. Honigmann, F. Jülicher, A. A. Hyman, and C. Zechner, Science 367, 464 (2020).
  76. A. Bray, Adv. Phys. 43, 357 (1994).

Outline

Information

Sign In to Your Journals Account

Filter

Filter

Article Lookup

Enter a citation