It is well-known that a “tether” may be drawn out from a pressurized liposome by means of a suitably applied radial-outward force applied locally to the lipid bilayer. The tether is a narrow, uniform cylindrical tube, which joins the main vesicle in a short “transition region.” A first-order energy analysis establishes the broad relationship between the force F needed to draw the tether, the radius R0 of the tether, the bending-stiffness constant B for the lipid bilayer and the membrane tension T in the pressurized liposome. The aim of the present paper is to study in detail the “transition region” between the tether and the main vesicle, by means of a careful application of the engineering theory of axisymmetric shell structures. It turns out that the well-known textbook “thin-shell” theory is inadequate for this purpose, because the tether is evidently an example of a thick-walled shell; and a novel ingredient of the present study is the introduction of elastic constitutive relations that are appropriate to the thick-shell situation. The governing equations are set up in dimensionless form, and are solved by means of a “shooting” technique, starting with a single disposable parameter at a point on the meridian in the tether, which can be adjusted until the boundary conditions at the far “equator” of the main vessel are satisfied. It turns out that the “transition region” between the tether and the main vessel is well characterized by only a few parameters, while the tether and main vessel themselves are described by very simple equations. Introduction of the thick-shell constitutive relation makes little difference to the conformation of, and stress-resultants in, the main vessel; but it makes a great deal of difference in the tether itself. Indeed, a kind of phase-change appears to take place in the “transition region” between these two zones of the liposome.

1.
Hochmuth
,
R. M.
, and
Evans
,
E. A.
,
1982
, “
Extensional Flow of Erythrocyte Membrane from Cell Body to Elastic Tether
,”
Biophys. J.
,
39
, pp.
71
81
.
2.
Waugh
,
R. E.
, and
Hochmuth
,
R. M.
,
1987
, “
Mechanical Equilibrium of Thick, Hollow, Liquid Membrane Cylinders
,”
Biophys. J.
,
52
, pp.
391
400
.
3.
Bo
,
L.
, and
Waugh
,
R. E.
,
1989
, “
Determination of Bilayer Membrane Bending Stiffness by Tether Formation from Giant, Thin-Walled Vesicles
,”
Biophys. J.
,
55
, pp.
509
517
.
4.
Bozic
,
B.
,
Sventina
,
S.
,
Zeks
,
B.
, and
Waugh
,
R. E.
,
1992
, “
Role of Lamellar Membrane Structure in Tether Formation from Bilayer Vesicles
,”
Biophys. J.
,
61
, pp.
963
973
.
5.
Waugh
,
R. E.
,
Song
,
J.
,
Svetina
,
S.
, and
Zeks
,
B.
,
1992
, “
Local and Nonlocal Curvature Elasticity in Bilayer Membranes by Tether Formation from Lecithin Vesicles
,”
Biophys. J.
,
61
, pp.
974
982
.
6.
Evans
,
E. A.
, and
Yeung
,
A.
,
1994
, “
Hidden Dynamics in Rapid Change of Bilayer Shape
,”
Chem. Phys. Lipids
,
73
, pp.
39
56
.
7.
Waugh
,
R. E.
, and
Bauserman
,
R. G.
,
1995
, “
Physical Measurements of Bilayer-Skeletal Separation Forces
,”
Ann. Biomed. Eng.
,
23
, pp.
308
321
.
8.
Hochmuth
,
R. M.
,
Shao
,
J.-Y.
,
Dai
,
J.
, and
Sheetz
,
M. P.
,
1996
, “
Deformation and Flow of Membrane into Tethers Extracted from Neuronal Growth Cones
,”
Biophys. J.
,
70
, pp.
358
369
.
9.
Bosic
,
B.
,
Svetina
,
S.
, and
Zeks
,
B.
,
1997
, “
Theoretical Analysis of the Formation of Membrane Microtubules on Axially strained Vesicles
,”
Phys. Rev. E
,
55
, pp.
5834
5842
.
10.
Dai
,
J.
, and
Sheetz
,
M. P.
,
1999
, “
Membrane Tether Formation from Blebbing cells
,”
Biophys. J.
,
77
, pp.
3363
3370
.
11.
Evans, E. A., and Skalak, R., 1980, Mechanics and Thermodynamics of Biomembranes, CRC Press, Boca Raton, FL.
12.
Canham
,
P. B.
,
1970
, “
The Minimum Energy of Bending as a Possible Explanation of the Biconcave Shape of the Human Red Blood Cell
,”
J. Theor. Biol.
,
26
, pp.
61
81
.
13.
Dueling
,
H. J.
, and
Helfrich
,
W.
,
1976
, “
Red Blood Cell Shapes as Explained on the Basis of Curvature Elasticity
,”
Biophys. J.
,
16
, pp.
861
868
.
14.
Sekimura
,
T.
, and
Hotani
,
H.
,
1991
, “
The Morphogenesis of Liposomes Viewed from the Aspect of Bending Energy
,”
J. Theor. Biol.
,
149
, pp.
325
337
.
15.
Timoshenko, S. P., and Woinowsky-Krieger, S., 1959, Theory of Plates and Shells, 2nd Edition, McGraw-Hill, New York.
16.
Flu¨gge, W., 1962, Stresses in Shells (2nd printing), Springer, Berlin.
17.
Novozhilov, V. V., 1964, The Theory of Thin Shells, Translation of 2nd Russian Edition by Lowe, P. G., ed. Radock, J. R. M., P. Noordhoff Ltd, Groningen.
18.
Calladine, C. R., 1983, Theory of Shell Structures, Cambridge University Press, Cambridge.
19.
Jenkins
,
J. T.
,
1977
, “
Static Equilibrium Configurations of a Model Red Blood Cell
,”
J. Math. Biol.
,
4
, pp.
149
169
.
20.
Pamplona
,
D. C.
, and
Calladine
,
C. R.
,
1993
, “
The Mechanics of Axially Symmetric Liposomes
,”
ASME J. Biomech. Eng.
115
, pp.
149
159
.
21.
Pamplona
,
D. C.
, and
Calladine
,
C. R.
,
1996
, “
Aspects of the Mechanics of Lobed Liposomes
,”
ASME J. Biomech. Eng.
118
, pp.
482
488
.
22.
Umeda
,
T.
,
Nakajima
,
H.
, and
Hotani
,
H.
,
1998
, “
Theoretical Analysis of Shape Transformations of Liposomes Caused by Microtubule Assembly
,”
Journal of the Physical Society of Japan
,
67
, pp.
682
688
.
23.
Bangham
,
A. D.
, and
Horne
,
R. W.
,
1964
, “
Negative Staining of Phospholipids and their Structural Modification by Surface-Active Agents as Observed in the Electron Microscope
,”
J. Mol. Biol.
,
8
, pp.
660
668
.
24.
Kornberg, R. D., and McConnell, H. M., 1971, “Lateral diffusion of phospholipids in a vesicle membrane,” Proceedings of the National Academy of Sciences of the USA, 68, pp. 2564–2568.
25.
Bangham, A. D., 1975, “Models of Cell Membranes,” Cell Membranes: Biochemistry, Cell Biology and Pathology, Weissmann, G., and Claiborne, R., eds. pp. 24–34. New York: Hospital Practice.
26.
Alberts, B., Bray, D., Lewis, L., Raff, M., Roberts, K., and Watson, J. D., 1983, Molecular Biology of the Cell, Garland, New York.
27.
Evans, E. A., Yeung, A., Waugh, R. E., and Song, J., 1983, “Dynamic Coupling and Nonlocal Curvature Elasticity in Bilayer Membranes,” Springer Proceedings in Physics, Vol. 66, The Structure and Conformation of Amphiphilic Membranes (Editors: Lipowski, R., Richter, D., and Kremer, K.), Springer-Verlag, Berlin Heidelberg.
28.
Mathivet
,
L.
,
Cribier
,
S.
, and
Devaux
,
P. F.
,
1996
, “
Shape Change and Physical Properties of Giant Phospholipid Vesicles Prepared in the Presence of an AC Electric Field
,”
Biophys. J.
,
70
, pp.
1112
1121
.
29.
Parker
,
K. H.
, and
Winlove
,
C. P.
,
1999
, “
The Deformation of Spherical Vesicles with Permeable, Constant-Area Membranes: Application to the Red Blood Cell
,”
Biophys. J.
,
77
, pp.
3096
3107
.
You do not currently have access to this content.