- Physical Review Journals
All JournalsPhysics Magazine

Physical Review Letters

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

Exchange Controls Coarsening of Surface Condensates

Riccardo Rossetto1,2, Marcel Ernst1,2, and David Zwicker1,*

  • *Contact author: david.zwicker@ds.mpg.de
PDF

Phys. Rev. Lett. 136, 198401 – Published 15 May, 2026

DOI: https://doi.org/10.1103/ndvb-yklf

Abstract

Biological membranes often exhibit heterogeneous protein patterns, which cells control. Strong patterns, like the polarity spot in budding yeast, can be described as surface condensates, formed by physical interactions between constituents. However, it is unclear how these interactions affect the material exchange with the bulk. To study this, we analyze a thermodynamically consistent model, which reveals that passive exchange generally accelerates the coarsening of surface condensates. Active exchange can further accelerate coarsening, although it can also fully arrest it and induce complex patterns involving various length scales. We reveal how these behaviors are related to nonlocal transport via diffusion through the bulk, rationalizing the various scaling laws that we observe and allowing us to interpret biologically relevant scenarios.

View figure in article

Physics Subject Headings (PhySH)

Article Text

Supplemental Material

References (64)

  1. S. Banjade and M. K. Rosen, eLife 3, e04123 (2014).
  2. D. B. McAffee, M. K. O’Dair, J. J. Lin, S. T. Low-Nam, K. B. Wilhelm, S. Kim, S. Morita, and J. T. Groves, Nat. Commun. 13, 7446 (2022).
  3. T. Litschel, C. F. Kelley, X. Cheng, L. Babl, N. Mizuno, L. B. Case, and P. Schwille, Nat. Commun. 15, 4986 (2024).
  4. A. Kamatar, J. P. K. Bravo, F. Yuan, L. Wang, E. M. Lafer, D. W. Taylor, J. C. Stachowiak, and S. H. Parekh, Biophys. J. 123, 1494 (2024).
  5. J.-g. Chiou, M. K. Balasubramanian, and D. J. Lew, Annu. Rev. Cell Dev. Biol. 33, 77 (2017).
  6. M.-P. Gulli, M. Jaquenoud, Y. Shimada, G. Niederhäuser, P. Wiget, and M. Peter, Mol. Cell 6, 1155 (2000).
  7. M. S. Turner, P. Sens, and N. D. Socci, Phys. Rev. Lett. 95, 168301 (2005).
  8. C. F. Lang and E. M. Munro, Biophys. J. 121, 4543 (2022).
  9. K. John and M. Bär, Phys. Rev. Lett. 95, 198101 (2005).
  10. K. John and M. Bär, Phys. Biol. 2, 123 (2005).
  11. R. Geßele, J. Halatek, L. Würthner, and E. Frey, Nat. Commun. 11, 539 (2020).
  12. F. Brauns, G. Pawlik, J. Halatek, J. Kerssemakers, E. Frey, and C. Dekker, Nat. Commun. 12, 3312 (2021).
  13. X. Zhao, G. Bartolucci, A. Honigmann, F. Jülicher, and C. A. Weber, New J. Phys. 23, 123003 (2021).
  14. X. Zhao, S. Liese, A. Honigmann, F. Jülicher, and C. A. Weber, New J. Phys. 26, 103025 (2024).
  15. X. Zhao, D. Sun, G. Bartolucci, A. A. Hyman, A. Honigmann, and C. A. Weber, eLife (2025).
  16. L. Foret, Europhys. Lett. 71, 508 (2005).
  17. N. Caballero, K. Kruse, and T. Giamarchi, Phys. Rev. E 108, L012801 (2023).
  18. D. Zwicker, O. Paulin, and C. ter Burg, Rep. Prog. Phys. 88, 116601 (2025).
  19. D. Zwicker, Curr. Opin. Colloid Interface Sci. 61, 101606 (2022).
  20. J. Kirschbaum and D. Zwicker, J. R. Soc. Interface 18, 20210255 (2021).
  21. R. Rossetto, G. Wellecke, and D. Zwicker, Phys. Rev. Res. 7, 023145 (2025).
  22. D. Qian, T. C. T. Michaels, and T. P. J. Knowles, J. Phys. Chem. Lett. 13, 7853 (2022).
  23. See Supplementl Material at http://link.aps.org/supplemental/10.1103/ndvb-yklf for the derivation of Eq. (4), linear stability analysis, details on the scaling behavior of the internal and diffusive fluxes and on the estimation of timescales, and evidence for multistability related to Fig. 3(a), which includes Refs. [18,21,24–28].
  24. D. Zwicker, J. Open Source Software 5, 2158 (2020).
  25. I. Lifshitz and V. Slyozov, J. Phys. Chem. Solids 19, 35 (1961).
  26. A. Kulkarni, E. Vidal-Henriquez, and D. Zwicker, Sci. Rep. 13, 733 (2023).
  27. C. Wagner, Z. Elektrochem. Ber. Bunsenges. Phys. Chem. 65, 581 (1961).
  28. C. F. Lee, Phys. Rev. Res. 3, 043081 (2021).
  29. J. W. Cahn and J. E. Hilliard, J. Chem. Phys. 28, 258 (1958).
  30. J. A. Marqusee, J. Chem. Phys. 81, 976 (1984).
  31. O. Bénichou, C. Loverdo, M. Moreau, and R. Voituriez, Rev. Mod. Phys. 83, 81 (2011).
  32. J. Rubinstein and P. Sternberg, IMA J. Appl. Math. 48, 249 (1992).
  33. A. Bray, Adv. Phys. 43, 357 (1994).
  34. C. A. Weber, D. Zwicker, F. Jülicher, and C. F. Lee, Rep. Prog. Phys. 82, 064601 (2019).
  35. S. C. Glotzer and A. Coniglio, Phys. Rev. E 50, 4241 (1994).
  36. J. J. Christensen, K. Elder, and H. C. Fogedby, Phys. Rev. E 54, R2212 (1996).
  37. C. B. Muratov, Phys. Rev. E 66, 066108 (2002).
  38. D. Zwicker, A. A. Hyman, and F. Jülicher, Phys. Rev. E 92, 012317 (2015).
  39. S. Köstler, Y. Qiang, G. Kusters, and D. Zwicker, Phys. Rev. Res. 8, 013107 (2026).
  40. Y. Qiang, C. Luo, and D. Zwicker, Phys. Rev. Lett. 135, 268301 (2025).
  41. Y. Cho and W. M. Jacobs, Proc. Natl. Acad. Sci. U.S.A. 122, e2501145122 (2025).
  42. J. Bauermann, G. Bartolucci, J. Boekhoven, F. Jülicher, and C. A. Weber, Phys. Rev. X 15, 041027 (2025).
  43. U. Thiele, A. J. Archer, M. J. Robbins, H. Gomez, and E. Knobloch, Phys. Rev. E 87, 042915 (2013).
  44. Y. Qiang, C. Luo, and D. Zwicker, Phys. Rev. X 14, 021009 (2024).
  45. G. Bartolucci, O. Adame-Arana, X. Zhao, and C. A. Weber, Biophys. J. 120, 4682 (2021).
  46. F. C. Thewes, Y. Qiang, O. W. Paulin, and D. Zwicker, arXiv:2511.05214.
  47. T. Ohta and K. Kawasaki, Macromolecules 19, 2621 (1986).
  48. F. Liu and N. Goldenfeld, Phys. Rev. A 39, 4805 (1989).
  49. J. W. Gibbs, Trans. Conn. Acad. Arts Sci. 3, 1 (1876).
  50. C. F. Lang and E. Munro, Development 144, 3405 (2017).
  51. M. Zeng, Y. Shang, Y. Araki, T. Guo, R. L. Huganir, and M. Zhang, Cell 166, 1163 (2016).
  52. G. Benn, I. V. Mikheyeva, P. G. Inns, J. C. Forster, N. Ojkic, C. Bortolini, M. G. Ryadnov, C. Kleanthous, T. J. Silhavy, and B. W. Hoogenboom, Proc. Natl. Acad. Sci. U.S.A. 118, e2112237118 (2021).
  53. M. Loose, K. Kruse, and P. Schwille, Annu. Rev. Biophys. 40, 315 (2011).
  54. L. Würthner, F. Brauns, G. Pawlik, J. Halatek, J. Kerssemakers, C. Dekker, and E. Frey, Proc. Natl. Acad. Sci. U.S.A. 119, e2206888119 (2022).
  55. L. Babl, A. Merino-Salomón, N. Kanwa, and P. Schwille, Sci. Rep. 12, 17949 (2022).
  56. T. Bland, N. Hirani, D. C. Briggs, R. Rossetto, K. Ng, I. A. Taylor, N. Q. McDonald, D. Zwicker, and N. W. Goehring, EMBO J. 43, 3214 (2024).
  57. N. W. Goehring, P. K. Trong, J. S. Bois, D. Chowdhury, E. M. Nicola, A. A. Hyman, and S. W. Grill, Science 334, 1137 (2011).
  58. M. Ernst, R. Rossetto, and D. Zwicker, PRX Life 4, 013033 (2026).
  59. J. A. Morin, S. Wittmann, S. Choubey, A. Klosin, S. Golfier, A. A. Hyman, F. Jülicher, and S. W. Grill, Nat. Phys. 18, 271 (2022).
  60. Q. Yu and A. Košmrlj, Soft Matter 21, 4288 (2025).
  61. A. Winter, Y. Liu, A. Ziepke, G. Dadunashvili, and E. Frey, Phys. Rev. E 111, 044405 (2025).
  62. N. Ziethen, J. Kirschbaum, and D. Zwicker, Phys. Rev. Lett. 130, 248201 (2023).
  63. R. Rossetto, M. Ernst, and D. Zwicker, Python code for the article “Exchange controls coarsening of surface condensates,” Zenodo (2026), 10.5281/zenodo.19610241.
  64. D. M. Rutkowski, V. Vincenzetti, D. Vavylonis, and S. G. Martin, Nat. Commun. 15, 8363 (2024).

Outline

Information

Phys. Rev. Lett. 136, 198401– Published 15 May, 2026
  • Received 5 November 2025
  • Accepted 13 April 2026
Crossmark - Check for updates button
Creative Commons Logo

Published by the American Physical Society under the terms of the Creative Commons Attribution 4.0 International license. Further distribution of this work must maintain attribution to the author(s) and the published article’s title, journal citation, and DOI. Open access publication funded by the Max Planck Society.

Published by the American Physical Society

Sign In to Your Journals Account

Filter

Filter

Article Lookup

Enter a citation