Abstract
Equilibrium self-assembly and conventional materials processing techniques fall far short of mimicking dynamic self-actuating processes that are commonplace throughout biology. Here, to bridge the gap between living and synthetic matter, we study passive adhesive non-thermal actin fibres immersed in an active microtubule-based fluid. We show that autonomous chaotic flows power non-equilibrium fibre dynamics, thus inducing collisions, generating connections and weaving a membrane-like elastic network. The ensuing active assembly generates a hierarchy of shapes, structures and dynamical processes spanning nanometres to centimetres. Ultimately, it generates an active membrane that exhibits global limit cycles induced by a non-reciprocal coupling between deformations of the elastic membrane and the alignment axis of the nematic active fluid. Our work merges self-assembly with active matter to demonstrate self-processing materials wherein hierarchical life-like structures and dynamics emerge from an initially structureless suspension.
This is a preview of subscription content, access via your institution
Access options
Access Nature and 54 other Nature Portfolio journals
Get Nature+, our best-value online-access subscription
$32.99 /Â 30Â days
cancel any time
Subscribe to this journal
Receive 12 print issues and online access
$259.00 per year
only $21.58 per issue
Buy this article
- Purchase on SpringerLink
- Instant access to the full article PDF.
USD 39.95
Prices may be subject to local taxes which are calculated during checkout





Data availability
Representative data from this study are available via Zenodo at https://doi.org/10.5281/zenodo.17215226 (ref. 67). Source data are provided with this paper.
Code availability
The code used for data analysis is available via Zenodo at https://doi.org/10.5281/zenodo.17215226 (ref. 67).
References
Cislo, D. J., Pavlopoulos, A. & Shraiman, B. I. ‘Morphogenetic action’ principle for 3D shape formation by the growth of thin sheets. Phys. Rev. X 15, 021068 (2025).
Mitchell, N. P. et al. Visceral organ morphogenesis via calcium-patterned muscle constrictions. eLife 11, e77355 (2022).
Nykypanchuk, D., Maye, M. M., van der Lelie, D. & Gang, O. DNA-guided crystallization of colloidal nanoparticles. Nature 451, 549–552 (2008).
Douglas, S. M. et al. Self-assembly of DNA into nanoscale three-dimensional shapes. Nature 459, 414–418 (2009).
Jacobs, W. M., Reinhardt, A. & Frenkel, D. Rational design of self-assembly pathways for complex multicomponent structures. Proc. Natl Acad. Sci. USA 112, 6313–6318 (2015).
He, M. et al. Colloidal diamond. Nature 585, 524–529 (2020).
Wei, W.-S. et al. Hierarchical assembly is more robust than egalitarian assembly in synthetic capsids. Proc. Natl Acad. Sci. USA 121, e2312775121 (2024).
Whitelam, S. & Jack, R. L. The statistical mechanics of dynamic pathways to self-assembly. Annu. Rev. Phys. Chem. 66, 143 (2015).
Keren, K. et al. Mechanism of shape determination in motile cells. Nature 453, 475 (2008).
Brugués, J. & Needleman, D. Physical basis of spindle self-organization. Proc. Natl Acad. Sci. USA 111, 18496 (2014).
Aditi Simha, R. & Ramaswamy, S. Hydrodynamic fluctuations and instabilities in ordered suspensions of self-propelled particles. Phys. Rev. Lett. 89, 058101 (2002).
Sanchez, T., Chen, D. T., DeCamp, S. J., Heymann, M. & Dogic, Z. Spontaneous motion in hierarchically assembled active matter. Nature 491, 431 (2012).
Wensink, H. H. et al. Meso-scale turbulence in living fluids. Proc. Natl Acad. Sci. USA 109, 14308 (2012).
Zhou, S., Sokolov, A., Lavrentovich, O. D. & Aranson, I. S. Living liquid crystals. Proc. Natl Acad. Sci. USA 111, 1265–1270 (2014).
Wu, X.-L. & Libchaber, A. Particle diffusion in a quasi-two-dimensional bacterial bath. Phys. Rev. Lett. 84, 3017–3020 (2000).
Di Leonardo, R. et al. Bacterial ratchet motors. Proc. Natl Acad. Sci. USA 107, 9541 (2010).
Sokolov, A., Apodaca, M. M., Grzybowski, B. A. & Aranson, I. S. Swimming bacteria power microscopic gears. Proc. Natl Acad. Sci. USA 107, 969 (2010).
Jia, H. et al. 3D printed protein-based robotic structures actuated by molecular motor assemblies. Nat. Mater. 21, 703–709 (2022).
Grober, D. et al. Unconventional colloidal aggregation in chiral bacterial baths. Nat. Phys. 19, 1680–1688 (2023).
Pedersen, M. C., Mukherjee, S., Doostmohammadi, A., Mondal, C. & Thijssen, K. Active particles knead three-dimensional gels into porous structures. Phys. Rev. Lett. 133, 228301 (2024).
Frechette, L. B., Baskaran, A. & Hagan, M. F. Active-noise-induced dynamic clustering of passive colloidal particles. Newton 1, 100167 (2025).
Martinet, Q., Li, Y., Aubret, A., Hannezo, E. & Palacci, J. Emergent dynamics of active elastic microbeams. Phys. Rev. X 15, 041017 (2025).
Lieleg, O., Claessens, M. M. A. E., Heussinger, C., Frey, E. & Bausch, A. R. Mechanics of bundled semiflexible polymer networks. Phys. Rev. Lett. 99, 088102 (2007).
Kayser, J., Grabmayr, H., Harasim, M., Herrmann, H. & Bausch, A. R. Assembly kinetics determine the structure of keratin networks. Soft Matter 8, 8873 (2012).
Falzone, T. T., Lenz, M., Kovar, D. R. & Gardel, M. L. Assembly kinetics determine the architecture of α-actinin crosslinked F-actin networks. Nat. Commun. 3, 861 (2012).
Foffano, G., Levernier, N. & Lenz, M. The dynamics of filament assembly define cytoskeletal network morphology. Nat. Commun. 7, 13827 (2016).
Berezney, J., Goode, B. L., Fraden, S. & Dogic, Z. Extensile to contractile transition in active microtubule-actin composites generates layered asters with programmable lifetimes. Proc. Natl Acad. Sci. USA 119, e2115895119 (2022).
Chandrakar, P. et al. Engineering stability, longevity, and miscibility of microtubule-based active fluids. Soft Matter 18, 1825–1835 (2022).
Paret, J. & Tabeling, P. Experimental observation of the two-dimensional inverse energy cascade. Phys. Rev. Lett. 79, 4162 (1997).
Shimoyama, N., Sugawara, K., Mizuguchi, T., Hayakawa, Y. & Sano, M. Collective motion in a system of motile elements. Phys. Rev. Lett. 76, 3870 (1996).
Baconnier, P. et al. Selective and collective actuation in active solids. Nat. Phys. 18, 1234–1239 (2022).
Baconnier, P. et al. Self-aligning polar active matter. Rev. Mod. Phys. 97, 015007 (2025).
You, Z., Baskaran, A. & Marchetti, M. C. Nonreciprocity as a generic route to traveling states. Proc. Natl Acad. Sci. USA 117, 19767 (2020).
Scheibner, C. et al. Odd elasticity. Nat. Phys. 16, 475 (2020).
Saha, S., Agudo-Canalejo, J. & Golestanian, R. Scalar active mixtures: the nonreciprocal Cahn-Hilliard model. Phys. Rev. X 10, 041009 (2020).
Fruchart, M., Hanai, R., Littlewood, P. B. & Vitelli, V. Non-reciprocal phase transitions. Nature 592, 363 (2021).
Tan, T. H. et al. Odd dynamics of living chiral crystals. Nature 607, 287 (2022).
Gu, F., Guiselin, B., Bain, N., Zuriguel, I. & Bartolo, D. Emergence of collective oscillations in massive human crowds. Nature 638, 112 (2025).
Fructhart, M., Vitelli, V. Nonreciprocal many-body physics. preprint available at https://arxiv.org/abs/2602.11111
Oron, A., Davis, S. H. & Bankoff, S. G. Long-scale evolution of thin liquid films. Rev. Mod. Phys. 69, 931 (1997).
Köhler, S., Schaller, V. & Bausch, A. R. Structure formation in active networks. Nat. Mater. 10, 462 (2011).
Murrell, M. P. & Gardel, M. L. F-actin buckling coordinates contractility and severing in a biomimetic actomyosin cortex. Proc. Natl Acad. Sci. USA 109, 20820–20825 (2012).
Alvarado, J., Sheinman, M., Sharma, A., MacKintosh, F. C. & Koenderink, G. H. Molecular motors robustly drive active gels to a critically connected state. Nat. Phys. 9, 591–597 (2013).
Foster, P. J., Fürthauer, S., Shelley, M. J. & Needleman, D. J. Active contraction of microtubule networks. eLife 4, e10837 (2015).
Lee, G. et al. Myosin-driven actin-microtubule networks exhibit self-organized contractile dynamics. Sci. Adv. 7, eabe4334 (2021).
Livne, G., Gat, S., Armon, S. & Bernheim-Groswasser, A. Self-assembled active actomyosin gels spontaneously curve and wrinkle similar to biological cells and tissues. Proc. Natl Acad. Sci. USA 121, e2309125121 (2024).
Hemingway, E. et al. Active viscoelastic matter: from bacterial drag reduction to turbulent solids. Phys. Rev. Lett. 114, 098302 (2015).
Liu, S., Shankar, S., Marchetti, M. C. & Wu, Y. Viscoelastic control of spatiotemporal order in bacterial active matter. Nature 590, 80–84 (2021).
Levanon, M. et al. Active flow-driven DNA remodeling generates millimeter-scale mechanical oscillations. Preprint at http://arxiv.org/abs/2511.22589 (2025).
Turlier, H. et al. Equilibrium physics breakdown reveals the active nature of red blood cell flickering. Nat. Phys. 12, 513–519 (2016).
Vutukuri, H. R. et al. Active particles induce large shape deformations in giant lipid vesicles. Nature 586, 52–56 (2020).
Sciortino, A. et al. Active membrane deformations of a minimal synthetic cell. Nat. Phys. 21, 799–807 (2025).
Nelson, D. & Peliti, L. Fluctuations in membranes with crystalline and hexatic order. J. Phys. 48, 1085 (1987).
Blees, M. K. et al. Graphene kirigami. Nature 524, 204–207 (2015).
Zhang, J., Childress, S., Libchaber, A. & Shelley, M. Flexible filaments in a flowing soap film as a model for one-dimensional flags in a two-dimensional wind. Nature 408, 835 (2000).
Saffman, P. & Turner, J. On the collision of drops in turbulent clouds. J. Fluid Mech. 1, 16 (1956).
Arguedas-Leiva, J.-A., Słomka, J., Lalescu, C. C., Stocker, R. & Wilczek, M. Elongation enhances encounter rates between phytoplankton in turbulence. Proc. Natl Acad. Sci. USA 119, e2203191119 (2022).
Broedersz, C. P., Mao, X., Lubensky, T. C. & MacKintosh, F. C. Criticality and isostaticity in fibre networks. Nat. Phys. 7, 983–988 (2011).
Sharma, A. et al. Strain-controlled criticality governs the nonlinear mechanics of fibre networks. Nat. Phys. 12, 584–587 (2016).
Bantawa, M. et al. The hidden hierarchical nature of soft particulate gels. Nat. Phys. 19, 1178–1184 (2023).
Ross, T. D. et al. Controlling organization and forces in active matter through optically defined boundaries. Nature 572, 224 (2019).
Klein, Y., Efrati, E. & Sharon, E. Shaping of elastic sheets by prescription of non-Euclidean metrics. Science 315, 1116–1120 (2007).
Kim, J., Hanna, J. A., Byun, M., Santangelo, C. D. & Hayward, R. C. Designing responsive buckled surfaces by halftone gel lithography. Science 335, 1201–1205 (2012).
Bearce, E. A. et al. Motile cilia spin the Reissner fiber, a tensioned and anchored extracellular thread essential for body morphogenesis. Preprint at bioRxiv https://doi.org/10.1101/2025.09.25.678623 (2025).
Bellegarda, C. et al. The Reissner fiber under tension in vivo shows dynamic interaction with ciliated cells contacting the cerebrospinal fluid. eLife 12, e86175 (2023).
Loiseau, E. et al. Active mucus–cilia hydrodynamic coupling drives self-organization of human bronchial epithelium. Nat. Phys. 16, 1158 (2020).
Ray, S., Berezney, J. & Kolvin, I. Data for: Active assembly and non-reciprocal dynamics of elastic membranes. Zenodo https://doi.org/10.5281/zenodo.17215226 (2025).
Acknowledgements
We thank L. Frechette, B. Chakraborty and X. Mao for insightful discussions. This work was primarily supported by the US Department of Energy (Grant No. DE-SC0022291 to J.B., S.R., I.K., S.F. and Z.D.). F.B. acknowledges support from a Gordon and Betty Moore post-doctoral fellowship (Award No. 2919). The theoretical work was supported in part by the National Science Foundation through Grant No. PHY-2309135 to the Kavli Institute for Theoretical Physics (M.B.). V.V. and S.C. acknowledge partial support from the Army Research Office (Grant Nos. W911NF-22-2-0109 and W911NF-23-1-0212), the National Science Foundation through the Center for Living Systems (Grant No. 2317138), the National Institute for Theory and Mathematics in Biology, the Simons Foundation and the Chan Zuckerberg Foundation.
Author information
Authors and Affiliations
Contributions
J.B., S.R., I.K. and Z.D. conceptualized the work. J.B., S.R. and I.K. performed the experiments. J.B., S.R., I.K. and F.B. analysed the data. F.B., S.C., M.B. and V.V. developed the theoretical analysis. All authors contributed to the writing of the paper.
Corresponding author
Ethics declarations
Competing interests
The authors declare no competing interests.
Peer review
Peer review information
Nature Physics thanks Jennifer Ross and Ishant Tiwari for their contribution to the peer review of this work. Peer reviewer reports are available.
Additional information
Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Supplementary information
Supplementary Information (download PDF )
Theoretical model, experimental methods, Figs. 1–5, Videos 1–9 and video captions.
Supplementary Video 1 (download MP4 )
Active MT fluid rearranges the passive actin network in the quasi-2D chamber. Preformed actin–fascin bundles (green) form anetwork and contract into a membrane under the influence of an active MT fluid (magenta). Rendering from a 3D imageacquired with a spinning-disc confocal microscope. Same conditions as in Fig 1a–e.
Supplementary Video 2 (download MP4 )
Active assembly of actin–fascin bundles. Maximum intensity of the z-projection of actin–fascin bundles forming a networkunder the influence of active MT flows (1.5 μM actin, 0.5 μM fascin, active buffer 1). Same sample as in Fig. 2. The averagetranslation within the displayed region was removed to highlight network reconfiguration during periods of large in-planemotion.
Supplementary Video 3 (download MP4 )
Actin assembly from very dilute preformed bundles. Active assembly generates a percolated actin network with a very largemesh size. Large deformations driven by the active fluid indicate that the percolated network lacks a finite elastic modulus andis floppy (1.0 μM actin, 0.33 μM fascin, active buffer 1).
Supplementary Video 4 (download MP4 )
Active assembly of actin–fascin network from unbundled actin. The active fluid induces both bundling and network assembly. Inthe initial state, the sample consists of unbundled actin (1.5 μM actin,  μM fascin, active buffer 2). Same sample as in Fig. 3.
Supplementary Video 5 (download MP4 )
Height fluctuations of actin membrane. Top left: 3D rendering of segmented actin network. Colour indicates local height (samecolour bar as Supplementary Fig. 1a). Top right: reconstruction of the surface based on actin segmentation. Bottom: sideview of the actin membrane (x – z slice).
Supplementary Video 6 (download MP4 )
Height fluctuations of actin membrane. Top left: 3D rendering of segmented actin network. Colour indicates local height (samecolour bar as Supplementary Fig. 1a). Top right: reconstruction of the surface based on actin segmentation. Bottom: sideview of the actin membrane (x – z slice).
Supplementary Video 7 (download MOV )
Macroscopic shear oscillations. System-sized shear oscillations emerge in the actin membrane (3.0 μM actin, 2.2 μM fascin,active buffer 2). Same sample as in Fig. 5a–c.
Supplementary Video 8 (download MOV )
Simulation of actin membrane oscillations. Velocity field (arrows) and vorticity (colour) showing self-excited waves in asimulation with w / ℓv−el = A = 1.6 and channel length L = 4w.
Supplementary Video 9 (download MOV )
Shear oscillations emerge in membranes above the critical width. Width dependence of system-sized shear oscillations (6.0 μMactin, 2.0 μM fascin, active buffer 1). Same sample as in Fig. 5e,f.
Source data
Source Data Fig. 2 (download XLSX )
Numerical source data for figure.
Source Data Fig. 3 (download XLSX )
Numerical source data for figure.
Source Data Fig. 4 (download XLSX )
Numerical source data for figure.
Source Data Fig. 5 (download XLSX )
Numerical source data for figure.
Rights and permissions
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.
About this article
Cite this article
Berezney, J., Ray, S., Kolvin, I. et al. Active assembly and non-reciprocal dynamics of elastic membranes. Nat. Phys. (2026). https://doi.org/10.1038/s41567-026-03215-5
Received:
Accepted:
Published:
Version of record:
DOI: https://doi.org/10.1038/s41567-026-03215-5