The coarsening of biomimetic condensates in an active fluid is non-self-similar

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Nature Physics (2026)Cite this article Coarsening, the growth of larger structures at the expense of smaller ones, is a fundamental process in multiphase systems. The cell cytoplasm is an example of an out-of-equilibrium multiphase system in which molecular phase-separated condensates nucleate and grow within an active fluid composed of biopolymers and energy-consuming enzymes. Here we uncover the mechanisms that govern the growth of condensates in a self-stirring active fluid. We study the coarsening of synthetic DNA-based condensates embedded within a three-dimensional reconstituted cytoskeleton composed of microtubules and molecular motors. By combining experiments and modelling, we explain the absence of self-similarity in active coarsening and the origin of the continuously varying coarsening exponents for condensates within either active or passive fluids. The coarsening dynamics are set by the statistics of binary collisions among droplets, which depend on their size-dependent motility, irrespective of their active or passive origins. We find that the scaling exponent of the collision kernel is a unifying control parameter for the coarsening and the size distribution of motile condensates. Our results expand our understanding of phase separation in far-from-equilibrium systems, with potential implications in materials science and biology.This is a preview of subscription content, access via your institution Access Nature and 54 other Nature Portfolio journals Get Nature+, our best-value online-access subscription $32.99 / 30 days cancel any timeSubscribe to this journal Receive 12 print issues and online access $259.00 per yearonly $21.58 per issueBuy this articleUSD 39.95Prices may be subject to local taxes which are calculated during checkoutAll data generated or analysed during this study are included in the article and its Supplementary Information. Data are also available from the corresponding author on reasonable request. Source data are provided with this paper.The code used during this study is available via GitHub at http://github.com/Layne28/active-coarsening (ref. 45).Cugliandolo, L. F. Coarsening phenomena.
Comptes Rendus Phys. 16, 257–266 (2015).Article ADS Google Scholar Randrup, J. Phase transition dynamics for baryon-dense matter. Phys. Rev. C 79, 054911 (2009).Article ADS Google Scholar Riemer, N. & Wexler, A. S. Droplets to drops by turbulent coagulation. J. Atmos. Sci. 62, 1962–1975 (2005).Article ADS Google Scholar Baity-Jesi, M. et al. Memory and rejuvenation effects in spin glasses are governed by more than one length scale. Nat. Phys. 19, 978–985 (2023).Article Google Scholar Hyman, A. A., Weber, C. A. & Julicher, F. Liquid-liquid phase separation in biology. Annu. Rev. Cell Dev. Biol. 30, 39–58 (2014).Article Google Scholar Ratke, L. & Voorhees, P. W. Growth and Coarsening: Ostwald Ripening in Material Processing 1st edn (Springer, 2002).Stansell, P., Stratford, K., Desplat, J. C., Adhikari, R. & Cates, M. E. Nonequilibrium steady states in sheared binary fluids. Phys. Rev. Lett. 96, 085701 (2006).Article ADS Google Scholar Gonnella, G. & Lamura, A. Long-time behavior and different shear regimes in quenched binary mixtures. Phys. Rev. E 75, 011501 (2007).Article ADS Google Scholar Stratford, K., Desplat, J. C., Stansell, P. & Cates, M. E. Binary fluids under steady shear in three dimensions. Phys. Rev. E 76, 030501 (2007).Article ADS Google Scholar Brangwynne, C. P. et al. Germline P granules are liquid droplets that localize by controlled dissolution/condensation. Science 324, 1729–1732 (2009).Article ADS Google Scholar Feric, M. et al. Coexisting liquid phases underlie nucleolar subcompartments. Cell 165, 1686–1697 (2016).Article ADS Google Scholar Jambon-Puillet, E. et al. Phase-separated droplets swim to their dissolution. Nat. Commun. 15, 3919 (2024).Article ADS Google Scholar Lifshitz, I. M. & Slyozov, V. V. The kinetics of precipitation from supersaturated solid solutions. J. Phys. Chem. Solids 19, 35–50 (1961).Article ADS Google Scholar Weber, C. A., Zwicker, D., Jülicher, F. & Lee, C. F. Physics of active emulsions. Rep. Prog. Phys. 82, 064601 (2019).Article ADS MathSciNet Google Scholar Cates, M. E. & Nardini, C. Active phase separation: new phenomenology from non-equilibrium physics. Rep. Prog. Phys. 88, 056601 (2025).Article ADS Google Scholar Wagner, C. Theorie der Alterung von Niederschlägen durch Umlösen (Ostwald-Reifung) (theory of the aging of precipitates by dissolution). Z. Elektrochem. 65, 581–591 (1961).
Google Scholar Smoluchowski, M. V. Versuch einer mathematischen Theorie der Koagulationskinetik kolloider Lösungen. Z. Phys. Chem. 92, 129–168 (1918).Article Google Scholar Siggia, E. D. Late stages of spinodal decomposition in binary mixtures. Phys. Rev. A 20, 595–605 (1979).Article ADS Google Scholar Mullins, W. W. The statistical self-similarity hypothesis in grain growth and particle coarsening. J. Appl. Phys. 59, 1341–1349 (1986).Article ADS Google Scholar Lee, D. S. W., Wingreen, N. S. & Brangwynne, C. P. Chromatin mechanics dictates subdiffusion and coarsening dynamics of embedded condensates. Nat. Phys. 17, 531–538 (2021).Article Google Scholar Caballero, F. & Marchetti, M. C. Activity-suppressed phase separation. Phys. Rev. Lett. 129, 268002 (2022).Article ADS Google Scholar Tayar, A. M. et al. Controlling liquid–liquid phase behaviour with an active fluid. Nat. Mater. 22, 1401–1408 (2023).Article ADS Google Scholar Arnold, D. P., Gubbala, A. & Takatori, S. C. Active surface flows accelerate the coarsening of lipid membrane domains. Phys. Rev. Lett. 131, 128402 (2023).Article ADS Google Scholar Tiribocchi, A., Wittkowski, R., Marenduzzo, D. & Cates, M. E. Active model H: scalar active matter in a momentum-conserving fluid. Phys. Rev. Lett. 115, 188302 (2015).Article ADS Google Scholar Singh, R. & Cates, M. E. Hydrodynamically interrupted droplet growth in scalar active matter. Phys. Rev. Lett. 123, 148005 (2019).Article ADS Google Scholar Adkins, R. et al. Dynamics of active liquid interfaces. Science 377, 768–772 (2022).Article ADS MathSciNet Google Scholar Zhao, L. et al. Asymmetric fluctuations and self-folding of active interfaces. Proc. Natl Acad. Sci. USA 121, e2410345121 (2024).Article Google Scholar Caballero, F., Maitra, A. & Nardini, C. Interface dynamics of wet active systems. Phys. Rev. Lett. 134, 087105 (2025).Article ADS MathSciNet Google Scholar Blow, M. L., Thampi, S. P. & Yeomans, J. M. Biphasic, lyotropic, active nematics. Phys. Rev. Lett. 113, 248303 (2014).Article ADS Google Scholar Biffi, S. et al. Phase behavior and critical activated dynamics of limited-valence DNA nanostars. Proc. Natl Acad. Sci. USA 110, 15633–15637 (2013).Article ADS Google Scholar Sanchez, T., Chen, D. T., DeCamp, S. J., Heymann, M. & Dogic, Z. Spontaneous motion in hierarchically assembled active matter. Nature 491, 431–434 (2012).Article ADS Google Scholar Jeon, B. J. et al. Salt-dependent properties of a coacervate-like, self-assembled DNA liquid. Soft Matter 14, 7009–7015 (2018).Article ADS Google Scholar Sato, Y., Sakamoto, T. & Takinoue, M. Sequence-based engineering of dynamic functions of micrometer-sized DNA droplets. Sci. Adv. 6, eaba3471 (2020).Article ADS Google Scholar Nguyen, D. T., Jeon, B. -j., Abraham, G. R. & Saleh, O. A. Length-dependence and spatial structure of DNA partitioning into a DNA liquid. Langmuir 35, 14849–14854 (2019).Article Google Scholar Berry, J., Weber, S. C., Vaidya, N., Haataja, M. & Brangwynne, C. P. RNA transcription modulates phase transition-driven nuclear body assembly. Proc. Natl Acad. Sci. USA 112, E5237–E5245 (2015).Article ADS Google Scholar Brangwynne, C. P., Mitchison, T. J. & Hyman, A. A. Active liquid-like behavior of nucleoli determines their size and shape in Xenopus laevis oocytes. Proc. Natl Acad. Sci. USA 108, 4334–4339 (2011).Article ADS Google Scholar Courchaine, E. M. et al. DMA-Tudor interaction modules control the specificity of in vivo condensates. Cell 184, 3612–3625 (2021).Article Google Scholar Derenzini, M., Montanaro, L. & Treré, D. What the nucleolus says to a tumour pathologist. Histopathology 54, 753–762 (2009).Article Google Scholar Buchwalter, A. & Hetzer, M. W. Nucleolar expansion and elevated protein translation in premature aging. Nat. Commun. 8, 328 (2017).Article ADS Google Scholar Fogelson, A. L. & Neeves, K. B. Fluid mechanics of blood clot formation. Annu. Rev. Fluid Mech. 47, 377–403 (2015).Article ADS MathSciNet Google Scholar Pradel, A., Catrouillet, C. & Gigault, J. The environmental fate of nanoplastics: what we know and what we need to know about aggregation. NanoImpact 29, 100453 (2023).Article Google Scholar Edelstein, A. D. et al. Advanced methods of microscope control using μManager software. J. Biol. Methods 1, e10 (2014).Article Google Scholar Ershov, D. et al. TrackMate 7: integrating state-of-the-art segmentation algorithms into tracking pipelines. Nat. Methods 19, 829–832 (2022).Article Google Scholar Varghese, M., Baskaran, A., Hagan, M. F. & Baskaran, A. Confinement-Induced self-pumping in 3D active fluids. Phys. Rev. Lett. 125, 268003 (2020).Article ADS MathSciNet Google Scholar Frechette, L. Layne28/active-coarsening. GitHub https://github.com/Layne28/active-coarsening (2024).Download referencesThis work was supported primarily by the NSF CAREER award DMR-2047119 (J.L., A.T.C. and G.D.) and by the NSF-funded Brandeis Bioinspired MRSEC award DMR-2011846 (L.B.F., C.A. and A.B.). We thank S. Dalal, director of the Brandeis Biomaterial Facility, for help with protein purification. We also acknowledge the use of the optical, microfluidics and biomaterial facilities supported by the NSF MRSEC award DMR-2011846. W.B.R. acknowledges support from the Human Frontier Science Program (RGP0029).Department of Physics, Brandeis University, Waltham, MA, USAJeremy Laprade, Layne B. Frechette, Christopher Amey, Adrielle T. Cusi, Aparna Baskaran, W. Benjamin Rogers & Guillaume DuclosSearch author on:PubMed Google ScholarSearch author on:PubMed Google ScholarSearch author on:PubMed Google ScholarSearch author on:PubMed Google ScholarSearch author on:PubMed Google ScholarSearch author on:PubMed Google ScholarSearch author on:PubMed Google ScholarJ.L. performed the experiments and the simulations and analysed the data. L.B.F. and C.A. developed the computer codes for the simulations. A.T.C. performed the experiments for the coarsening in a passive fluid. A.B. and W.B.R. contributed materials and analysis tools. G.D. and W.B.R. performed experiments in the early stage of the project. G.D. supervised the project and wrote the paper. All co-authors provided feedback on the paper.Correspondence to Guillaume Duclos.The authors declare no competing interests.Nature Physics thanks Atul Parikh, David Zwicker and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.Discussion about the biological relevance and Supplementary Figs. 1–19.Coarsening of DNA droplets embedded in an active fluid.Tracking droplet trajectories advected by the active flows.Simulations of droplet coarsening in an active fluid.Simulations of coarsening for diffusive droplets with m = 1, 0 or −1.Tables with data points.Tables with data points.Tables with data points.Tables with data points.Tables with data points.Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.Reprints and permissionsLaprade, J., Frechette, L.B., Amey, C. et al. The coarsening of biomimetic condensates in an active fluid is non-self-similar. Nat. Phys. (2026). https://doi.org/10.1038/s41567-026-03191-wDownload citationReceived: 17 December 2024Accepted: 23 January 2026Published: 06 March 2026Version of record: 06 March 2026DOI: https://doi.org/10.1038/s41567-026-03191-wAnyone you share the following link with will be able to read this content:Sorry, a shareable link is not currently available for this article. Provided by the Springer Nature SharedIt content-sharing initiative
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