pith. machine review for the scientific record. sign in

arxiv: 2602.16002 · v4 · submitted 2026-02-17 · ⚛️ physics.comp-ph · physics.bio-ph

Recognition: 2 theorem links

· Lean Theorem

Liquid Crystal Theory of Biomembranes

Authors on Pith no claims yet

Pith reviewed 2026-05-15 21:31 UTC · model grok-4.3

classification ⚛️ physics.comp-ph physics.bio-ph
keywords biomembranesliquid crystalsHelfrich modelmembrane shapesDelaunay surfacesred blood cellselastic theoryshape transitions
0
0 comments X

The pith

Biomembrane shapes such as spheres, tori and biconcave discs form a group due to shared intrinsic geometry.

A machine-rendered reading of the paper's core claim, the machinery that carries it, and where it could break.

The paper uses liquid crystal theory and the Helfrich elastic model to analyze biomembrane shapes. It establishes that shapes including cylinders, spheres, tori, biconcave discoids and Delaunay surfaces belong to a single group. This grouping is an intrinsic geometric property and does not depend on the specific form of the biomembrane equation. The shapes emerge when pressure, surface tension and bending moduli satisfy particular conditions. This provides a unified explanation for diverse membrane morphologies observed in biology and synthetic systems.

Core claim

The shapes such as cylinders, spheres, tori, biconcave discoids and Delaunay surfaces form a group. This result is merely an intrinsic geometric feature of these shapes and is independent of the biomembrane equation. When the pressure on the membrane, surface tension, and bending modules meet certain conditions, the biomembrane will take on these shapes.

What carries the argument

The group of membrane shapes unified by their intrinsic geometry under the Helfrich elastic model.

If this is right

  • Biomembranes will adopt these grouped shapes under tuned physical conditions without additional molecular constraints.
  • The biconcave shape of red blood cells follows directly from the elastic model matching the group properties.
  • Similar shape formations occur in multi-layer systems and peptide assemblies by analogy to smectic liquid crystals.
  • Reversible transitions in assemblies and icosahedral structures align with the geometric group.
  • The continuum theory unifies descriptions across biological and synthetic membrane systems.

Where Pith is reading between the lines

These are editorial extensions of the paper, not claims the author makes directly.

  • This geometric unification may extend to other curved surfaces in soft matter physics.
  • Testing by controlling pressure and tension in artificial membranes could confirm the shape selection.
  • The approach might connect to minimal surface mathematics through the Delaunay surfaces.
  • Limitations at nanoscale could require hybrid models incorporating molecular details.

Load-bearing premise

The observed shapes arise purely from the geometric properties of the surfaces under the elastic model without requiring additional molecular-level interactions or specific boundary conditions.

What would settle it

Observation of a biomembrane maintaining a shape outside the identified group, such as a highly irregular or asymmetric form, despite pressure, surface tension, and bending moduli satisfying the required conditions.

read the original abstract

Biomembranes, primarily composed of lipid bilayers, are not merely passive barriers but dynamic and complex materials whose shapes are governed by the principles of soft matter physics. This review explores the shape problem in biomembranes through the lens of material science and liquid crystal theory. Beginning with classical analogies to crystals and soap bubbles, it details the application of the Helfrich elastic model to explain the biconcave shape of red blood cells. The discussion extends to multi-layer systems, drawing parallels between the focal conic structures of smectic liquid crystals, the geometries of fullerenes and carbon nanotubes, and the reversible transitions in peptide assemblies. Furthermore, it examines icosahedral self-assemblies and shape formation in two-dimensional lipid monolayers at air/water interfaces. At the end of the paper, we find that the shapes such as cylinders, spheres, tori, biconcave discoids and Delaunay surfaces form a group. This result is merely an intrinsic geometric feature of these shapes and is independent of the biomembrane equation. When the pressure on the membrane, surface tension, and bending modules meet certain conditions, the biomembrane will take on these shapes. The review concludes by highlighting the unifying power of continuum elastic theories in describing a vast array of membrane morphologies across biological and synthetic systems.

Editorial analysis

A structured set of objections, weighed in public.

Desk editor's note, referee report, simulated authors' rebuttal, and a circularity audit. Tearing a paper down is the easy half of reading it; the pith above is the substance, this is the friction.

Referee Report

1 major / 1 minor

Summary. This review applies liquid crystal theory and the Helfrich elastic model to biomembrane shape formation, drawing analogies to soap bubbles, smectic focal conics, fullerenes, and peptide assemblies. It concludes that cylinders, spheres, tori, biconcave discoids, and Delaunay surfaces constitute a geometric group whose selection is an intrinsic surface property independent of the biomembrane equation, appearing only when pressure, surface tension, and bending moduli satisfy certain conditions.

Significance. If the geometric-independence claim can be rigorously separated from the Helfrich variational structure, the work would offer a unifying continuum framework for a wide range of observed membrane morphologies across biological and synthetic systems.

major comments (1)
  1. [Abstract/Conclusion] Abstract/Conclusion: The central claim that the listed shapes form a group 'merely as an intrinsic geometric feature ... independent of the biomembrane equation' is undercut by the immediately following statement that the shapes appear only when pressure, surface tension, and bending moduli meet conditions taken from the Helfrich functional. The manuscript must show explicitly (e.g., via constant-mean-curvature identities or group-theoretic properties of the surfaces) that the grouping follows from geometry alone, without invoking the Euler-Lagrange equation or its parameters.
minor comments (1)
  1. [Abstract] The phrase 'bending modules' should read 'bending moduli'.

Simulated Author's Rebuttal

1 responses · 0 unresolved

We thank the referee for the careful review and valuable feedback on our manuscript. We address the major comment below and propose revisions to strengthen the presentation of our central claim.

read point-by-point responses
  1. Referee: [Abstract/Conclusion] Abstract/Conclusion: The central claim that the listed shapes form a group 'merely as an intrinsic geometric feature ... independent of the biomembrane equation' is undercut by the immediately following statement that the shapes appear only when pressure, surface tension, and bending moduli meet conditions taken from the Helfrich functional. The manuscript must show explicitly (e.g., via constant-mean-curvature identities or group-theoretic properties of the surfaces) that the grouping follows from geometry alone, without invoking the Euler-Lagrange equation or its parameters.

    Authors: We agree that the current phrasing in the abstract and conclusion could be misinterpreted as contradictory. The intent of the claim is that the shapes constitute a geometric family (specifically, the constant-mean-curvature surfaces of revolution and related topologies) whose selection is dictated by intrinsic surface geometry once the parameters permit the energy minimization to yield constant mean curvature. To address this, we will revise the abstract and conclusion to explicitly demonstrate the grouping via constant-mean-curvature identities: these surfaces are classified by the Delaunay theorem as the only surfaces of revolution with constant mean curvature, and the biconcave discoid corresponds to the CMC surface with appropriate topology. This geometric classification holds independently of the specific values of the moduli or the precise form of the functional, provided the conditions reduce the problem to H = constant. We will add a brief paragraph in the conclusion citing the relevant differential geometry results to make this separation clear, without relying on the full Euler-Lagrange derivation in the revised text. revision: yes

Circularity Check

1 steps flagged

Independence claim for geometric shape group is qualified by model-specific parameter conditions from Helfrich equation

specific steps
  1. other [Abstract (final paragraph)]
    "At the end of the paper, we find that the shapes such as cylinders, spheres, tori, biconcave discoids and Delaunay surfaces form a group. This result is merely an intrinsic geometric feature of these shapes and is independent of the biomembrane equation. When the pressure on the membrane, surface tension, and bending modules meet certain conditions, the biomembrane will take on these shapes."

    The asserted independence of the geometric grouping from the biomembrane equation is immediately followed by the statement that the membrane adopts precisely these shapes only when parameters of that equation (pressure, surface tension, bending moduli) satisfy model-specific conditions. The selection mechanism therefore remains tied to the variational structure of the Helfrich functional, undercutting the claimed separation without an explicit geometric derivation shown to be free of the elastic energy.

full rationale

The paper is a review synthesizing liquid-crystal and Helfrich-model results from prior literature. Its central closing claim asserts that the listed shapes form a geometric group independent of the biomembrane equation, yet immediately qualifies their adoption by conditions on pressure, tension and bending moduli drawn from that same model. This mixing does not reduce any derivation to a tautology by construction, nor does it rely on a load-bearing self-citation chain or fitted prediction; the geometric grouping is presented as an observation rather than a derived output. No self-definitional loop or ansatz smuggling is exhibited in the quoted text. The overall derivation chain therefore remains largely self-contained as a review, warranting only a modest circularity score.

Axiom & Free-Parameter Ledger

2 free parameters · 2 axioms · 0 invented entities

The review relies on established models from prior literature; no new free parameters or entities introduced in the abstract. The geometric feature is presented as independent but depends on the validity of the elastic theory assumptions.

free parameters (2)
  • bending modulus
    Central parameter in Helfrich model fitted to match observed shapes like biconcave discs.
  • surface tension
    Conditioned on meeting certain values along with pressure for shape selection.
axioms (2)
  • domain assumption Continuum elastic approximation for lipid bilayers
    Assumed in applying Helfrich model to biomembranes.
  • domain assumption Analogy between biomembranes and smectic liquid crystals
    Used to draw parallels with focal conic structures.

pith-pipeline@v0.9.0 · 5522 in / 1315 out tokens · 44069 ms · 2026-05-15T21:31:26.755925+00:00 · methodology

discussion (0)

Sign in with ORCID, Apple, or X to comment. Anyone can read and Pith papers without signing in.

Lean theorems connected to this paper

Citations machine-checked in the Pith Canon. Every link opens the source theorem in the public Lean library.

What do these tags mean?
matches
The paper's claim is directly supported by a theorem in the formal canon.
supports
The theorem supports part of the paper's argument, but the paper may add assumptions or extra steps.
extends
The paper goes beyond the formal theorem; the theorem is a base layer rather than the whole result.
uses
The paper appears to rely on the theorem as machinery.
contradicts
The paper's claim conflicts with a theorem or certificate in the canon.
unclear
Pith found a possible connection, but the passage is too broad, indirect, or ambiguous to say the theorem truly supports the claim.

Reference graph

Works this paper leans on

118 extracted references · 118 canonical work pages

  1. [1]

    = 0,(24) where λ=λ/k c +C 2 0 /2, ∆P= ∆P/k c. The shape of the red blood cell has been computed using the above elasticity model (Deuling & Helfrich, 1976 [33]; Jenkins, 1977 [34]; Peterson, 1985 [35]; Svetina & ˇZekˇ s, 1989 [36]; Miao et al., 1991 [37]). Another special variational method is the 1D plane curve variation δ ˆ F(ψ(s), dψ/ds, d 2ψ/ds2)ds= 0...

  2. [2]

    This solution is not a numerical approximation but a closed-form mathematical function: sinψ=C 0ρln(ρ/ρ B),(28) whereC 0 <0 for bioconcave shape of RBC

    was to demonstrate that under the condition of zero osmotic pressure difference, the axisymmetric shape equation admits a specific analytic solution that perfectly describes the classic biconcave disc profile. This solution is not a numerical approximation but a closed-form mathematical function: sinψ=C 0ρln(ρ/ρ B),(28) whereC 0 <0 for bioconcave shape of...

  3. [3]

    He then introduced infinitesimal shape perturbations, mathematically expressed as a series of spherical harmonicsY lm(θ, ϕ)

    They have treated the sphere as the ”ground state” of a closed vesicle under a reference pressure. He then introduced infinitesimal shape perturbations, mathematically expressed as a series of spherical harmonicsY lm(θ, ϕ). Each harmonicY lm(θ, ϕ) represents a distinct deformation mode: The indexl(degree) determines the wavelength or ”wiggliness” of the d...

  4. [4]

    The nanotube to spherical vesicle transition is linked through joined necklace-like struc- tures

    find beautiful way to construct such a surface with rotationally symmetry: By rolling a given conic section on a straight line in a plane, and then rotating the trace of a focus about the line, one obtains the surface. The nanotube to spherical vesicle transition is linked through joined necklace-like struc- tures. The metastable necklace-like structure c...

  5. [5]

    Contrary to this prediction, high-resolution structural studies revealed that these viruses are constructed exclusively from pentagonal morphological units

    and Simian Virus 40 [99], the Caspar-Klug model predicts a capsid built from a combi- 33 nation of hexagonal and pentagonal capsomeres. Contrary to this prediction, high-resolution structural studies revealed that these viruses are constructed exclusively from pentagonal morphological units. This discrepancy highlights a significant limitation in the prev...

  6. [6]

    Les etats mesomorphes de la matiere,

    G. Friedel, “Les etats mesomorphes de la matiere,” Annales de Physique, vol. 18, pp. 273–474, (1922)

  7. [8]

    Helfrich, Elastic properties of lipid bilayers: theory and possible experiments, Z

    W. Helfrich, Elastic properties of lipid bilayers: theory and possible experiments, Z. Natur- forsch. C28, 693 (1973)

  8. [9]

    Helfrich, Instability and deformation of a spherical vesicle by pressure, Vol

    Ou-Yang Zhong-Can and W. Helfrich, Instability and deformation of a spherical vesicle by pressure, Vol. 59, No. 21, 2486-2488, (1987). DOI: 10.1103/PhysRevLett.59.2486

  9. [10]

    Bending energy of vesicle membranes: General expressions for the first, second, and third variation of the shape energy and applications to spheres and cylinders,

    Ou-Yang Zhong-Can and Wolfgang 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, vol. 39, pp. 5280–5288, (1989)

  10. [11]

    Naito, M

    H. Naito, M. Okuda & Z.C. Ou-Yang, Analytic solutions for the shape of the human red blood cell. Physical Review E, 48(3), 2304–2309 (1993)

  11. [12]

    Podgornik, R.; Parsegian, V. A. Thermal-mechanical fluctuations of fluid membranes in confined geometries: The case of soft confinement. Langmuir 1992, 8 (2), 557–562. DOI: 10.1021/la00038a041

  12. [13]

    S., Parsegian, V

    Dean, D. S., Parsegian, V. A., and Podgornik, R. (2015). Fluctuation mediated interactions 47 due to rigidity mismatch, Journal of Physics: Condensed Matter, 27(21), 214004

  13. [14]

    Majee, A., Bier, M., Blossey, R., and Podgornik, R. (2019). Charge regulation radically modifies electrostatics in membrane stacks. Physical Review E, 100(5), 050601

  14. [15]

    Khunpetch, P., Majee, A., and Podgornik, R. (2022). Curvature effects in charge-regulated lipid bilayers. Soft Matter, 18(13), 2597-2610

  15. [16]

    Markovich, T., Andelman, D., and Podgornik, R. (2016). Charge regulation: A generalized boundary condition?. Europhysics Letters, 113(2), 26004

  16. [17]

    Wu, Zhong-Can Ou-Yang, & R

    H. Wu, Zhong-Can Ou-Yang, & R. Podgornik. Electrostatic-elastic coupling in colloidal crystals, EPL, 148, 47001 (2024)

  17. [18]

    Wu, Zhong-Can Ou-Yang, & R

    H. Wu, Zhong-Can Ou-Yang, & R. Podgornik. Continuum theory of electrostatic-elastic coupling interactions in colloidal crystals, Commun. Theor. Phys. 77, 055602(2025)

  18. [19]

    Ou-Yang, Z.B

    Z.C. Ou-Yang, Z.B. Su & C.L. Wang, Coil Formation in Multishell Carbon Nanotubes: Competition between Curvature Elasticity and Interlayer Adhesion. Physical Review Letters, 78(21), 4055–4058 (1997)

  19. [20]

    Ou-Yang, The study of complex shapes of fluid membrane, the Helfrich variation model and new application, conference of soft matter in Xiamen city, (2025)

    Z.C. Ou-Yang, The study of complex shapes of fluid membrane, the Helfrich variation model and new application, conference of soft matter in Xiamen city, (2025)

  20. [21]

    M. Yan, X. Yao, Y. Zhang, Z.C. Ou-Yang & W. Huang, Chemistry, Manipulating Carbon Nanotubes with Optical Tweezers, Cemistry-A European Journal, Vol. 14, 5699–6007 (2008)

  21. [22]

    Wulff, Zur Frage der Geschwindigkeit des Wachsthums und der Aufloung der Krystallfla- gen

    G. Wulff, Zur Frage der Geschwindigkeit des Wachsthums und der Aufloung der Krystallfla- gen. Zeitschrift fur Kristallographie-Crystalline Materials, 34(5/6), 449–530, (1901)

  22. [23]

    Plateau, Statique experimentale et theorique des liquides soumis aux seules forces moleculaires (Vol

    J.A.F. Plateau, Statique experimentale et theorique des liquides soumis aux seules forces moleculaires (Vol. 2). Gauthier-Villars (1873)

  23. [24]

    Young, An Essay on the Cohesion of Fluids

    T. Young, An Essay on the Cohesion of Fluids. Philosophical Transactions of the Royal Society of London, 95, 65–87. (Introduces the concepts of surface tension, the contact angle, and a qualitative link between curvature and pressure), (1805)

  24. [25]

    Laplace, Traite de mecanique celeste, Supplement to Book 10, ”On Capillary Action.” Courcier, Paris

    P.S. Laplace, Traite de mecanique celeste, Supplement to Book 10, ”On Capillary Action.” Courcier, Paris. (Presents the first rigorous mechanical and mathematical derivation of the pressure equation), (1806)

  25. [26]

    Young, Miscellaneous Works of the Late Thomas Young, Vol

    T. Young, Miscellaneous Works of the Late Thomas Young, Vol. I (edited by G. Peacock). John Murray, London. (Contains a later commentary and consolidation of his ideas on cap- illarity), (1855). 48

  26. [27]

    A. D. Aleksandrov, Uniqueness theorems for surfaces in the large. I. Vestnik Leningrad University, 11(19), 5–17, (1956)

  27. [28]

    E. A. Evans, & P.F. Leblond, Image Holograms of Single Red Blood Cell Discocyte- Spheroechinocyte Transformations. In Red Cell Shape (pp. 138-145). Springer, (1973)

  28. [29]

    Korpman, D

    R.A. Korpman, D. Dorrough, J.D. Brailsford,and B.S. Bull, The Red Cell Shape as an Indicator of Membrane Structure: Ponder’s Rule Reexamined. In Red Cell Rheology (pp. 199-209). Springer. (1978)

  29. [30]

    Bartosz, E

    G. Bartosz, E. Grzeli´ nska & J. Wagner, Aging of the erythrocyte. XIV. ATP content does decrease. Cellular and Molecular Life Sciences, 38(5), 575, (1982)

  30. [31]

    Lopez, S

    R.A. Lopez, S. Schoetz, K. DeAngelis, D. O’Neill, & A. Bank, Multiple hematopoietic defects and delayed globin switching in Ikaros null mice. Proceedings of the National Academy of Sciences of the United States of America, 99(1), 602-607, (2002)

  31. [32]

    Greer, & R.F

    W.J.N. Greer, & R.F. Baker, The minimum energy of bending as a possible explanation of the biconcave shape of the human red blood cell. Journal of Theoretical Biology, 26(1), 61-81. (Note: Reference details beyond title/year are illustrative based on common citation format), (1970)

  32. [33]

    Frank, On the theory of liquid crystals

    F.C. Frank, On the theory of liquid crystals. Discussions of the Faraday Society, 25, 19-28, (1958)

  33. [34]

    Eriksson, S

    J.C. Eriksson, S. Ljunggren, The multiple chemical equilibrium approach to the theory of droplet microemulsions. In: Zulauf, M., Lindner, P., Terech, P. (eds) Trends in Col- loid and Interface Science IV. Progress in Colloid & Polymer Science, vol 81. Steinkopff. https://doi.org/10.1007/BFb0115521

  34. [35]

    Kazuhiko Seki, Shigeyuki Komura, Brownian dynamics in a thin sheet with momentum decay, Physical Review E, Vol. 47, No. 4, P2377-2383, (1993). DOI: 10.1103/physreve.47.2377

  35. [36]

    FEBS Letters, 333(1-2), 169–174, (1993)

    Tsukada, M., & Ohsumi, Y., Isolation and characterization of autophagy-defective mutants of Saccharomyces cerevisiae. FEBS Letters, 333(1-2), 169–174, (1993)

  36. [38]

    Deuling, & W

    H.J. Deuling, & W. Helfrich, The curvature elasticity of fluid membranes: A catalogue of vesicle shapes. Journal De Physique, 37, 1335–1345, (1976) 49

  37. [39]

    Jenkins, J. T. (1977). The equations of mechanical equilibrium of a model membrane. SIAM Journal on Applied Mathematics, 32(4), 755–764

  38. [40]

    Peterson, M. A. (1985). Geometrical methods for the elasticity theory of membranes. Journal of Mathematical Physics, 26(4), 711–717

  39. [41]

    Svetina, S., & ˇZekˇ s, B. (1989). Membrane bending energy and shape determination of phos- pholipid vesicles and red blood cells. European Biophysics Journal, 17(2), 101–111

  40. [42]

    Miao, L., Fourcade, B., Rao, M., Wortis, M., & Zia, R. K. P. (1991). Equilibrium budding and vesiculation in the curvature model of fluid lipid vesicles. Physical Review A, 43(12), 6843–6856

  41. [43]

    Berndl, J

    J. Berndl, J. Kas, R. Lipowsky, E. Sackmann, & U. Seifert, Shape Transformations of Giant Vesicles: Extreme Sensitivity to Bilayer Asymmetry, Europhysics Letters, Vol. 13, No 7, 659 (1990), DOI 10.1209/0295-5075/13/7/015

  42. [44]

    Seifert, Vesicles of toroidal topology, Physical Review Letters, Vol 66, No.18, 2404-2407 (1991)

    U. Seifert, Vesicles of toroidal topology, Physical Review Letters, Vol 66, No.18, 2404-2407 (1991)

  43. [45]

    Seifert, K

    U. Seifert, K. Berndl, & R. Lipowsky, Shape transformations of vesicles: Phase diagram for spontaneous-curvature and bilayer-coupling models, Physical Review A, Vol 44, No.2, 1182-1202, (1991)

  44. [46]

    Shape equations for axisymmetric vesicles: A clarification

    Julicher F, Seifert U. Shape equations for axisymmetric vesicles: A clarification. Phys Rev E Stat Phys Plasmas Fluids Relat Interdiscip Topics. May;49(5):4728-4731, (1994). doi: 10.1103/physreve.49.4728. PMID: 9961774

  45. [47]

    Derenyi, F

    I. Derenyi, F. Julicher, J. Prost, Formation and Interaction of Membrane Tubes, Phys. Rev. Lett. 88, 238101 (2002)

  46. [48]

    M., Liu, J

    Zheng, W. M., Liu, J. X. (1993). Relation between shape equations for axisymmetric vesicles. Physical Review E, 48(4), 2856–2859. doi: 10.1103/PhysRevE.48.2856

  47. [49]

    C., Hu, J

    Ou-Yang, Z. C., Hu, J. G., & Liu, J. X. (1992). Spontaneous curvature of fluid vesicles and the structure of biomembranes. Science in China Series A-Mathematics, Physics, Astronomy & Technological Sciences, 35(11), 1301–1309

  48. [50]

    A., Fung, Y.C

    Evans, E. A., Fung, Y.C. (1972). Improved measurements of the erythrocyte geometry. Mi- crovascular Research, 4(4), 335–347

  49. [51]

    Meyer, R. B. (1969). Piezoelectric Effects in Liquid Crystals. Physical Review Letters, 22(18), 918-921. 50

  50. [52]

    Schmidt, M

    D. Schmidt, M. Schadt and W. Helfrich, Z. Naturforsch, A 27, 277 (1972)

  51. [53]

    Dozov, Ph

    L. Dozov, Ph. Martinot-Lagarde and G. Durand, J. Phys. Lett. (paris) 43, L365, (1982)

  52. [54]

    Lassen, O

    U.V. Lassen, O. Sten-Knudsen, J. Physiol, 195, 681, (1968)

  53. [55]

    J., Lecar, H

    Nossal, R. J., Lecar, H. (1991). Molecular and cell biophysics. Addison-Wesley Pub. Co., the Advanced Book Program

  54. [56]

    C., Liu, J

    Ou-Yang, Z. C., Liu, J. X., & Xie, Y. Z. (1990). Torus vesicle. Europhysics Letters (EPL), 13(5), 421–426

  55. [57]

    M. Mutz, D. Bensimon, Observation of toroidal vesicles, Physical Review A, 43, 4525 (1991). DOI: 10.1103/PhysRevA.43.4525

  56. [58]

    A. S. Rudolph, B. R. Ratna, B. Kahn, Self-assembling phospholipid filaments, Vol. 352, No. 6330, 52-55, (1991). DOI: https://doi.org/10.1038/352052a0

  57. [59]

    Z. LinJ. J. CaiL. E. ScrivenH. T. Davis, J. Phys. Chem. 1994, 98, 23, 5984–5993, (1994). https://doi.org/10.1021/j100074a027

  58. [60]

    Hotani, Transformation pathways of liposomes, J

    H. Hotani, Transformation pathways of liposomes, J. Mol. Biol. 178, 113 (1984)

  59. [61]

    Pond, R.V

    E. Pond, R.V. Pond, Observations quantitatives sur la production de formes myeliniques par les stromas de globules rouges. Nouv. Rev. fr. Hemat.3, 553 (1963)

  60. [62]

    Bessis, Living blood cells and their ultrastructure, (Springer-Verlag, Berlin, 1973)

    M. Bessis, Living blood cells and their ultrastructure, (Springer-Verlag, Berlin, 1973)

  61. [63]

    Evans and W

    E. Evans and W. Rawicz, Entropy-Driven Tension and Bending Elasticity in Condensed-Fluid Membranes, Phys. Rev. Lett.64, 2094 (1990)

  62. [64]

    Zhou J.J, Y

    J.J. Zhou J.J, Y. Zhang, X. Zhou, Z.C. Ouyang, Large deformation of spherical vesicle studied by perturbation theory and surface evolver, IJMPB 15, 2977 (2001)

  63. [65]

    T. Xu, Z.C. Ouyang, Formation of multispheres and myelin based on multiple solutions of membrane shape equation. Membranes 15, 319, (2025)

  64. [66]

    Lipowsky, Understanding giant vesicles: a theoretical perspective, (CRC press) (2020)

    R. Lipowsky, Understanding giant vesicles: a theoretical perspective, (CRC press) (2020)

  65. [67]

    Bhatia, S

    T. Bhatia, S. Christ, J. Steinkuhler, R. Dimova, and R. Lipowsky, Simple sugars shape giant vesicles into multispheres with many membrane necks, Soft Matter,16, 1246 (2020)

  66. [68]

    Iwamoto, Z.C

    M. Iwamoto, Z.C. Ouyang, Anharmonic magnetic deformation of spherical vesicle: Field- induced tension and swelling effects, Chem. Phys. Lett. 590, 183-187 (2013)

  67. [69]

    Saitoh A, Takiguchi K, Tanaka Ya, et al., Opening-up of liposomal membranes by talin, PNAS 95, 1026 (1998). 51

  68. [70]

    C., & Ou-Yang, Z

    Tu, Z. C., & Ou-Yang, Z. C. (2003). A geometric theory on the elasticity of bio-membranes. Phys. Rev. E, 68(6), 061915

  69. [71]

    C., & Ou-Yang, Z

    Tu, Z. C., & Ou-Yang, Z. C. (2004). Lipid membranes with free edges. J. Phys. A: Math. Gen., 37(47), 11407

  70. [72]

    Tu, Z. C. (2010). A molecular theory for modeling the stability of cell membranes. J. Chem. Phys., 132(8), 084111

  71. [73]

    Z.C. Tu, Z.C. Ouyang, J.X. Liu, Y.Z. Xie, Geometric methods in elastic theory of membranes in liquid crystal phases, (Peking university press, Beijing, 2014)

  72. [74]

    Happel, H

    J. Happel, H. Brenner, Low Reynolds number hydrodynamics with special applications to particulate media, (Martinus Nijhoff Publishes, Boston), (1983)

  73. [75]

    & Liu, J.-X

    Ouyang, Z.-C. & Liu, J.-X. Vesicle shapes with varying spontaneous curvature and area- difference elasticity. Phys. Rev. Lett. 65, 1679–1682 (1990)

  74. [76]

    & Liu, J.-X

    Ouyang, Z.-C. & Liu, J.-X. Phase diagrams for vesicles with axisymmetric shapes. Phys. Rev. A 43, 6826–6836 (1991)

  75. [77]

    Ito, Stochastic thermodynamic interpretation of informa- tion geometry, Physical Review Letters121, 10.1103/phys- revlett.121.030605 (2018)

    Helfirch and J. Prost, Intrinsic bending force in anisotropic membranes made of chi- ral molecules. Phys Rev A Gen Phys. 1988 Sep 15;38(6):3065-3068. doi: 10.1103/phys- reva.38.3065. PMID: 9900723

  76. [78]

    Schnur, J. M. Lipid Tubules: A Paradigm for Molecularly Engineered Structures. Science 262, 1669–1676 (1993)

  77. [79]

    Diacetylenic lipid tubules: experimental evidence for a chiral molecular architecture

    Schnur JM, Ratna BR, Selinger JV, Singh A, Jyothi G, Easwaran KR. Diacetylenic lipid tubules: experimental evidence for a chiral molecular architecture. Science. 1994 May 13;264(5161):945-7. doi: 10.1126/science.264.5161.945. PMID: 17830081

  78. [80]

    Chiral molecular self-assembly of phospholipid tubules: a circular dichroism study

    Spector MS, Easwaran KR, Jyothi G, Selinger JV, Singh A, Schnur JM. Chiral molecular self-assembly of phospholipid tubules: a circular dichroism study. Proc Natl Acad Sci U S A. 1996 Nov 12;93(23):12943-6. doi: 10.1073/pnas.93.23.12943. PMID: 8917523; PMCID: PMC24025

  79. [81]

    G., The Physics of Liquid Crystals, Second Edition, (Clarendon Press, 1993)

    de Gennes, P. G., The Physics of Liquid Crystals, Second Edition, (Clarendon Press, 1993)

  80. [82]

    Elasticity of chiral membranes

    Lubensky, T. C. & Prost, J. “Elasticity of chiral membranes.” J. Phys. II France 2, 371 (1992)

Showing first 80 references.