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

Euler-Helfrich problem for open vesicles bounded by elastic filaments

Chengyao Zhang and Xin Yi*

  • School of Mechanics and Engineering Science, Peking University, Beijing 100871, China

  • *Contact author: xyi@pku.edu.cn

Phys. Rev. Research 8, 013225 – Published 2 March, 2026

DOI: https://doi.org/10.1103/9byv-ndvs

Abstract

We develop a theoretical framework to elucidate the mechanics and morphogenesis of open vesicles coupled to boundary filaments, capturing their geometric, energetic, and mechanical interplay. Using a spherical harmonics parametrization and constrained energy minimization, we systematically explore equilibrium morphologies as functions of filament length, stiffness, and topology, as well as membrane spontaneous curvature, uncovering a rich spectrum of shapes and transitions. For open vesicles bounded by flexible closed filaments, increasing filament length or spontaneous curvature drives transitions from cuplike to stomatocyte and budded morphologies, both axisymmetric and nonaxisymmetric, with continuous and discontinuous regimes mapped in a phase diagram. For open vesicles partially bounded by open filaments, the coupled filament-membrane system exhibits more complex morphological evolution, including free edge stretching or contraction, localized filament bending, and opening expansion with increasing filament length. The derived force and moment balances at the vesicle edge connect local geometric quantities, such as filament and membrane curvatures, to filament internal forces and membrane tension, providing a direct mechanical interpretation of shape equilibria. Our results reveal how filament geometry, topology, and bending stiffness govern membrane shape transformations and establish a general mechanical framework for coupled filament-membrane systems, such as DNA-vesicle complexes and other filament-mediated membrane structures.

View figure in article

Physics Subject Headings (PhySH)

Article Text

References (48)

  1. D. Bray, Cell Movements: From Molecules to Motility, 2nd ed. (Garland Science, New York, 2001).
  2. X.-Q. Feng, B. Li, S. Z. Lin, M. Y. Wang, X. D. Chen, H. X. Zhang, and W. Fang, Mechano-chemo-biological theory of cells and tissues: Review and perspectives, Acta Mech. Sinica 41, 625315 (2025).
  3. D. A. Fletcher and R. D. Mullins, Cell mechanics and the cytoskeleton, Nature (London) 463, 485 (2010).
  4. A. H. Bahrami, M. G. Lin, X. Ren, J. H. Hurley, and G. Hummer, Scaffolding the cup-shaped double membrane in autophagy, PLoS Comput. Biol. 13, e1005817 (2017).
  5. J. Agudo-Canalejo and R. Lipowsky, Domes and cones: Adhesion-induced fission of membranes by ESCRT proteins, PLoS Comput. Biol. 14, e1006422 (2018).
  6. B. Meadowcroft, I. Palaia, A. K. Pfitzner, A. Roux, B. Baum, and A. Šarić, Mechanochemical rules for shape-shifting filaments that remodel membranes, Phys. Rev. Lett. 129, 268101 (2022).
  7. C. M. Funkhouser, R. Sknepnek, T. Shimi, A. E. Goldman, R. D. Goldman, M. Olvera de la Cruz, Mechanical model of blebbing in nuclear lamin meshworks, Proc. Natl. Acad. Sci. USA 110, 3248 (2013).
  8. Y. Turgay, M. Eibauer, A. Goldman, T. Shimi, M. Khayat, K. Ben-Harush, A. Dubrovsky-Gaupp, K. T. Sapra, R. D. Goldman, and O. Medalia, The molecular architecture of lamins in somatic cells, Nature (London) 543, 261 (2017).
  9. D. Deviri, C. R. Pfeifer, L. J. Dooling, I. L. Ivanovska, D. E. Discher, and S. A. Safran, Scaling laws indicate distinct nucleation mechanisms of holes in the nuclear lamina, Nat. Phys. 15, 823 (2019).
  10. T. Litschel, C. F. Kelley, D. Holz, M. A. Koudehi, S. K. Vogel, L. Burbaum, N. Mizuno, D. Vavylonis, and P. Schwille, Reconstitution of contractile actomyosin rings in vesicles, Nat. Commun. 12, 2254 (2021).
  11. C. Shi, G. Zou, Z. Wu, M. Wang, X. Zhang, H. Gao, and X. Yi, Morphological transformations of vesicles with confined flexible filaments, Proc. Natl. Acad. Sci. USA 120, e2300380120 (2023).
  12. A. Sciortino, H. A. Faizi, D. A. Fedosov, L. Frechette, P. M. Vlahovska, G. Gompper, and A. R. Bausch, Active membrane deformations of a minimal synthetic cell, Nat. Phys. 21, 799 (2025).
  13. C. Shi, C. Zhang, Y. Fang, and X. Yi, Flexible filaments in vesicles with reduced volume: Anisotropic confinement and morphological response, J. Mech. Phys. Solids 206, 106383 (2026).
  14. C. Zhang, G. Zou, Y. Fang, H. Gao, and X. Yi, Interacting filaments drive vesicle morphogenesis, Nat. Commun. 17, 78 (2026).
  15. T. Savin, N. Kurpios, A. Shyer, P. Florescu, H. Liang, L. Mahadevan, and C. Tabin, The growth and form of the gut, Nature (London) 476, 57 (2011).
  16. A. Fragasso, N. De Franceschi, P. Stömmer, E. O. van der Sluis, H. Dietz, and C. Dekker, Reconstitution of ultrawide DNA origami pores in liposomes for transmembrane transport of macromolecules, ACS Nano 15, 12768 (2021).
  17. M. L. Daly, K. Nishi, S. J. Klawa, K. Y. Hinton, Y. Gao, and R. Freeman, Designer peptide–DNA cytoskeletons regulate the function of synthetic cells, Nat. Chem. 16, 1229 (2024).
  18. T. Umeda, Y. Suezaki, K. Takiguchi, and H. Hotani, Theoretical analysis of opening-up transformations within single and two holes, Phys. Rev. E 71, 011913 (2005).
  19. Z. Yao, R. Sknepnek, C. K. Thomas, M. Olvera de la Cruz, Shapes of pored membranes, Soft Matter 8, 11613 (2012).
  20. Y. Liu, G. Zou, and H. Gao, Domain aggregation and associated pore growth in lipid membranes, ACS Nano 15, 604 (2021).
  21. A. Saitoh, K. Takiguchi, Y. Tanaka, and H. Hotani, Opening-up of liposomal membranes by talin, Proc. Natl. Acad. Sci. USA 95, 1026 (1998).
  22. F. Nomura, M. Nagata, T. Inaba, H. Hiramatsu, and K. Takiguchi, Capabilities of liposomes for topological transformation, Proc. Natl. Acad. Sci. USA 98, 2340 (2001).
  23. R. J. C. Gilbert, M. Dalla Serra, C. J. Froelich, M. I. Wallace, and G. Anderluh, Membrane pore formation at protein–lipid interfaces, Trends Biochem. Sci. 39, 510 (2014).
  24. H. G. Franquelim, H. Dietz, and P. Schwille, Reversible membrane deformations by straight DNA origami filaments, Soft Matter 17, 276 (2021).
  25. J. Zhu, Z. A. McDargh, F. Li, S. S. Krishnakumar, J. E. Rothman, and B. O’Shaughnessy, Synaptotagmin rings as high-sensitivity regulators of synaptic vesicle docking and fusion, Proc. Natl. Acad. Sci. USA 119, e2208337119 (2022).
  26. I. G. Denisov and S. G. Sligar, Nanodiscs in membrane biochemistry and biophysics, Chem. Rev. 117, 4669 (2017).
  27. W. Helfrich, Elastic properties of lipid bilayers: Theory and possible experiments, Z. Naturforsch. C 28, 693 (1973).
  28. P. J. Flory, Statistical Mechanics of Chain Molecules (Hanser Publishers, Munich, 1988).
  29. L. Giomi and L. Mahadevan, Minimal surfaces bounded by elastic lines, Proc. R. Soc. A 468, 1851 (2012).
  30. A. Biria and E. Fried, Buckling of a soap film spanning a flexible loop resistant to bending and twisting, Proc. R. Soc. A 470, 20140368 (2014).
  31. A. Biria, M. Maleki, and E. Fried, Continuum theory for the edge of an open lipid bilayer, Adv. Appl. Mech. 46, 1 (2013).
  32. B. Palmer and Á. Pámpano, Minimizing configurations for elastic surface energies with elastic boundaries, J. Nonlinear Sci. 31, 23 (2021).
  33. B. Palmer and Á. Pámpano, The Euler–Helfrich functional, Calc. Var. 61, 79 (2022).
  34. L. D. Landau and E. M. Lifshitz, Theory of Elasticity (Butterworth-Heinemann, New York, 1986), 3rd ed.
  35. A. Nádai, Über das ausbeulen von kreisförmigen platten, Z. Ver. Dtsch. Ing. 59, 169 (1915).
  36. N. Yamaki, Buckling of a thin annular plate under uniform compression, J. Appl. Mech. 25, 267 (1958).
  37. M. Wang and X. Yi, Bulging and budding of lipid droplets from symmetric and asymmetric membranes: Competition between membrane elastic energy and interfacial energy, Soft Matter 17, 5319 (2021).
  38. H. A. Hameed, J. Paturej, and A. Erbaş, Shape spectra of elastic shells with surface-adsorbed semiflexible polymers, Soft Matter 22, 234 (2026).
  39. R. Capovilla and J. Guven, Stresses in lipid membranes, J. Phys. A: Math. Gen. 35, 6233 (2002).
  40. Z. C. Tu and Z.-C. Ou-Yang, A geometric theory on the elasticity of bio-membranes, J. Phys. A: Math. Gen. 37, 11407 (2004).
  41. M. Deserno, Fluid lipid membranes: From differential geometry to curvature stresses, Chem. Phys. Lipids 185, 11 (2015).
  42. P. Yang, Q. Du, and Z. C. Tu, General neck condition for the limit shape of budding vesicles, Phys. Rev. E 95, 042403 (2017).
  43. K. Khairy and J. Howard, Minimum-energy vesicle and cell shapes calculated using spherical harmonics, Soft Matter 7, 2138 (2011).
  44. C. Zhang, Y. Fang, C. Shi, H. Yuan, and X. Yi, Stretching transition of vesicles with confined filament loops: Morphological evolution with filament distortion and reorientation, Giant 17, 100233 (2024).
  45. H. W. Guggenheimer, Differential Geometry (Dover Publications, New York, 1977).
  46. R. Capovilla, J. Guven, and J. A. Santiago, Lipid membranes with an edge, Phys. Rev. E 66, 021607 (2002).
  47. Z. C. Tu and Z.-C. Ou-Yang, Lipid membranes with free edges, Phys. Rev. E 68, 061915 (2003).
  48. Z.-C. Ou-Yang and W. Helfrich, Bending energy of vesicle membranes: General expressions for the first, second, and third variation of the shape energy and applications to spheres and cylinders, Phys. Rev. A 39, 5280 (1989).

Outline

Information

Sign In to Your Journals Account

Filter

Filter

Article Lookup

Enter a citation