JGSP 24 (2011) 45–75

GEOMETRY OF MEMBRANES

ZHANCHUN TU

Communicated by Boris Konopeltchenko

Abstract. This review reports some theoretical results on the geometry

of membranes. The governing equations to describe equilibrium configu-
rations of lipid vesicles, lipid membranes with free edges, and chiral lipid
membranes are derived from the variation of free energies of these struc-
tures. Some analytic solutions to these equations and their corresponding
configurations are also shown.

Contents
1 Introduction 46

2 Preliminaries in Mathematics and Physics 47

2.1 Surface Theory Based on Moving Frame . . . . . . . . . . . . . . 47
2.1.1. Moving Frame Method . . . . . . . . . . . . . . . . . . . 47
2.1.2. Stokes’ Theorem and Related Identities . . . . . . . . . . 49
2.2 Helfrich’s Model . . . . . . . . . . . . . . . . . . . . . . . . . . 50
2.3 Variational Method Based on Moving Frame . . . . . . . . . . . . 51

3 Lipid Vesicles 52
3.1 Shape Equation to Describe Equilibrium Configurations . . . . . . 52
3.2 Analytic Solutions and Corresponding Configurations . . . . . . . 54
3.2.1. Surfaces of Constant Mean Curvature . . . . . . . . . . . 54
3.2.2. Torus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
3.2.3. Biconcave Discoid . . . . . . . . . . . . . . . . . . . . . 56
3.2.4. Unduloid-Like and Cylinder-Like Surfaces . . . . . . . . 57

45
46 Zhanchun Tu

4 Lipid Membranes with Free Edges 58

4.1 Shape Equation and Boundary Conditions to Describe Equilibrium
Configurations . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
4.2 Theorems of Non-Existence . . . . . . . . . . . . . . . . . . . . 61
4.3 Quasi-Exact Solutions . . . . . . . . . . . . . . . . . . . . . . . 63

5 Chiral Lipid Membranes 64

5.1 Constructing the Free Energy . . . . . . . . . . . . . . . . . . . . 65
5.2 Governing Equations to Describe Equilibrium Configurations . . . 66
5.3 Solutions and Corresponding Configurations . . . . . . . . . . . . 67
5.3.1. Sphere . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
5.3.2. Cylinder . . . . . . . . . . . . . . . . . . . . . . . . . . 68
5.3.3. Torus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
5.3.4. Twisted Ribbons . . . . . . . . . . . . . . . . . . . . . . 69

6 Summary and Conclusions 71

1. Introduction

Membranes are very crucial to living organisms. They are the barriers of cells and
ensure cells to be relatively isolated individuals but still able to exchange some ma-
terials between the inner sides and outer surroundings through specific ways due
to the fancy properties of membranes. Membranes usually consist of lipid bilay-
ers mosaicks built of various kinds of proteins. They are also the key factors to
determine shapes of some kinds cells. In particular, the biconcave discoidal shape
of the red blood cells is regarded as a result of minimization of the free energy of
membranes under the area and volume constraints [4, 11] because red blood cells
have no complex inner structures. The equilibrium configurations of membranes
have attracted much attention of mathematicians and physicists [19, 32, 34, 38, 46].
A membrane is thought of as a two-dimensional (2D) smooth surface in the Eu-
clidean space E3 because its thickness is much smaller than its lateral dimension.
The first step to investigate configurations of membranes is the construction of a
free energy functional by consideration of symmetry. Then the governing equa-
tions to describe the equilibrium configurations can be derived by variation of the
free energy with some constraints. The next task is seeking for solutions to satisfy
the governing equations and comparing the results with the typical experiments.
 Geometry of Membranes 47

In this review, we will present some purely theoretical results on geometry of mem-
branes. For simplicity, we merely focus on structures of lipid membranes and
select only the theoretical problems that both physicists and mathematicians are
interested in. The governing equations to describe equilibrium configurations of
lipid structures are derived. Several solutions to these equations and their corre-
sponding geometries are also investigated. The rest of this review is organized as
follows.
In Section 2, we give a brief introduction to preliminary in mathematics and physics
including surface theory and variational method based on moving frame, and Hel-
frich’s model of lipid bilayer. In Section 3, we derive a shape equation that de-
scribes equilibrium configurations of lipid vesicles - closed lipid bilayers. Then
we discuss some analytic solutions and their corresponding configurations which
include surfaces of constant mean curvature, torus, biconcave discoid, cylinder-like
vesicles, and so on. In Section 4, we investigate a lipid membrane with free edge(s).
The shape equation and boundary conditions describing equilibrium configurations
of the membrane are derived. Then we discuss the compatibility between the shape
equation and boundary conditions, and verify five theorems of non-existence. In
Section 5, we construct the free energy functional of chiral lipid membranes in
terms of symmetric argument and then derive the governing equations to describe
their equilibrium configurations by variational method. Some analytic solutions
and their corresponding configurations are also shown. In the last section, we give
a brief summary and a list of related open questions.

2. Preliminaries in Mathematics and Physics

In order to continue our discussion, we first introduce several key mathematical

and physical concepts and tools that will be used in the following sections.

2.1. Surface Theory Based on Moving Frame

2.1.1. Moving Frame Method

A membrane can be regarded as a smooth orientable surface embedded in E 3 .

The properties of the surface such as its mean and Gaussian curvatures determine
the shape of the membrane. As shown in Fig. 1, each point on the surface M
can be represented by a position vector r. At that point we construct three unit
orthonormal vectors e1 , e2 , and e3 with e3 being the normal vector to the surface
M at the point r. The set of right-handed orthonormal triple-vectors {e 1 , e2 , e3 } is
48 Zhanchun Tu

called a frame at the point r. Different points on the surface have different vectors
r, e1 , e2 , and e3 , thus the set {r, e1 , e2 , e3 } is called a moving frame.

Figure 1. The moving frame on a surface.

Let us imagine a mass point that moves from position r to its neighbor position r 
on the surface. The length of the path is denoted by Δs. Then we can define the
differentiation of the frame as

dr = lim (r − r) = ω1 e1 + ω2 e2 (1)

Δs→0

and
dei = lim (ei − ei ) = ωij ej , i = 1, 2, 3 (2)
Δs→0

where ω1 , ω2 , and ωij (i, j = 1, 2, 3) are one forms, and ‘d’ is the exterior differ-
ential operator [7]. The one form ωij is anti-symmetric with respect to the indices
i and j, that is ωij = −ωji . Here and in the following contents without special
statements, the repeated subscripts represent summation from 1 to 3. Additionally,
the structure equations of the surface can be expressed as [7]

dω1 = ω12 ∧ ω2 , dω2 = ω21 ∧ ω1 , dωij = ωik ∧ ωkj , i, j = 1, 2, 3 (3)

and
ω13 ab ω1
= (4)
ω23 b c ω2
where ‘∧’ denotes the wedge product of the two differential forms. The matrix
ab
is the representation matrix of the curvature tensor. Its trace and determi-
b c
nant are two invariants under the coordinate rotation around e 3 which are denoted
 Geometry of Membranes 49

respectively by
2H = a + c and K = ac − b2 . (5)
H and K are called the mean curvature and Gaussian curvature, respectively. They
determine the shape of the surface and can be expressed as 2H = −(1/R 1 +1/R2 )
and K = R1−1 R2−1 via the two principal curvature radii R1 and R2 at each point
on the surface.
Now consider a curve on the surface M. Its tangent vector is denoted by t. Let φ
be the angle between t and e1 at the same point. Then the geodesic curvature kg ,
the geodesic torsion τg , and the normal curvature kn along the direction of t can
be expressed as [42]

kg = (dφ + ω12 )/ds

τg = b cos 2φ + (c − a) cos φ sin φ (6)
2 2
kn = a cos φ + 2b cos φ sin φ + c sin φ

where ds is the arc length along t. If t aligns with e1 , then φ = 0, kg = ω12 /ds,
τg = b, and kn = a. In the principal frame, the geodesic torsion and normal
curvature can be expressed as
cos2 φ sin2 φ
kn = − − , τg = (1/R1 − 1/R2 ) cos φ sin φ. (7)
R1 R2

2.1.2. Stokes’ Theorem and Related Identities

Stokes’ theorem is a crucial theorem in differential geometry, which reads

 
ω= dω (8)
∂D D

where D is a domain with boundary ∂D and ω is a differential form on ∂D. In

particular, D dω = 0 for a closed domain D.
From Stokes’ theorem, we can derive several identities listed as follows [42, 46].

i) For smooth functions f and h on 2D subdomain D ⊆ M

 
(f d∗dh − hd∗df ) = (f ∗dh − h∗df )
 D  ∂D

(f d∗d̃h − hd∗d̃f ) = (f ∗d̃h − h∗d̃f ) (9)

D ∂D
(f d˜∗d̃h − hd˜∗d̃f ) = (f ˜∗d̃h − h˜
∗d̃f )
D ∂D
50 Zhanchun Tu

where ∗ is Hodge star operator satisfying ∗ω1 = ω2 and ∗ω2 = −ω1 . Here d̃
and ˜∗ are generalized differential operator and generalized Hodge star which
satisfy d̃f = f1 ω13 + f2 ω23 and ˜∗d̃f = f1 ω23 − f2 ω13 if df = f1 ω1 + f2 ω2
[42, 43].

ii) If u and v are two vector fields defined on 2D subdomain D ⊆ M, then [44]
 
(u · d∗dv − v · d∗du) = (u · ∗dv − v · ∗du)
D
  ∂D

(u · d∗d̃v − v · d∗d̃u) = (u · ∗d̃v − v · ∗d̃u) (10)

D ∂D
(u · d˜∗d̃v − v · d˜∗d̃u) = (u · ˜∗d̃v − v · ˜
∗d̃u)
D ∂D

where the ‘dot’ represents the inner product of vectors. For simplicity, equa-
tions (9) and (10) are still called Stokes’ theorem in this review. They are
widely used in the variational process.

Additionally, we can also define the gradient, curl, divergent, Laplace operators,
etc. on the surface in terms of the differential operators and Hodge stars. They are
summarized as follows [45, 46]

(∇ × u)dA = d(u · dr), (∇ · u)dA = d(∗u · dr), ∇f · dr = df

˜ · u)dA = d(˜∗u · d̃r),
(∇ ¯ · u)dA = d(∗u · d̃r), ∇f
(∇ ˜ · dr = d̃f
(∇2 f )dA = d ∗ df, ¯ )dA = d ∗ d̃f,
(∇ · ∇f S · dr = −ω12
(u · ∇f )dA = u · dr ∧ ∗df, ˜ )dA = d˜∗d̃f
(∇ · ∇f (11)
˜ )dA = u · dr ∧ ˜∗d̃f,
(u · ∇f ¯ )dA = u · dr ∧ ∗d̃f
(u · ∇f
(∇2 u)dA = d ∗ du, (∇u)· dr = du
(∇u: ∇v)dA=du∧dv,
˙ (∇ · S)dA = −d ∗ ω12

where dA = ω1 ∧ω2 and S are the area element and spin connection of the surface,
respectively. And ∧˙ denotes the simultaneous calculations of the dot and wedge
products.

2.2. Helfrich’s Model

In 1973, Helfrich proposed a spontaneous curvature model to describe the free

energy of lipid membranes by analogy with a bent box of liquid crystal in Smectic
 Geometry of Membranes 51

A phase [11]. The free energy is in fact a functional defined in the space of shapes
of membranes, which reads
  
kc 2
FH = (2H + Ih) + k̄K dA (12)
M 2

where M, H, and K represent the membrane surface, mean curvature, and Gaus-
sian curvature, respectively. The constants kc and k̄ are two bending moduli. The
former should be positive, while the latter can be negative or positive for lipid
membranes. The real constant Ih is called spontaneous curvature, which reflects
the asymmetrical chemical or physical factors between two leaves of lipid bilay-
ers.
On the other hand, the spontaneous curvature model can also be obtained from
symmetric argument. A lipid membrane can be locally regarded as 2D isotropic
elastic entity. Thus the local free energy density f should be invariant under ro-
tational transformation around the normal direction of the membrane surface. In
other words, it should be a function of H and K because H and K are the funda-
mental invariants of the surface under rotational transformation. Up to the second
order terms of curvatures, it can be expanded as
fc = A 0 + A 1 H + A 2 H 2 + A 3 K (13)
which can be rewritten as
kc
fc = (2H + Ih)2 + k̄K (14)
2
by omitting an unimportant constant. This is nothing else than the integrand in
equation (12). Therefore, the spontaneous curvature model is of general signif-
icance not only for lipid membranes, but also for other membranes consisting
of isotropic materials. The following discussions are mainly based on Helfrich’s
spontaneous curvature model.

2.3. Variational Method Based on Moving Frame

To obtain governing equations that describe equilibrium configurations of lipid

membranes, we need to minimize the free energy. That is, we should calculate
the variation of functionals defined in the space of shapes of membranes. In this
space, each shape might be expressed as a function, but the defining domain is not
a fixed planar domain. It is difficult to deal with this case by using the traditional
variational calculus. Now we will introduce the variational method based on the
moving frame developed by the present author and Ou-Yang [41, 42, 46], which is
capable to deal with variational problems on surfaces.
52 Zhanchun Tu

Any infinitesimal deformation of a surface can be achieved by a displacement vec-

tor
δr ≡ v = Ωi ei (15)
at each point on the surface, where δ can be understood as a variational operator.
The frame is also changed because of the deformation of the surface, which is
denoted as
δei = Ωij ej , i = 1, 2, 3 (16)
where Ωij = −Ωji (i, j = 1, 2, 3) corresponds to the rotation of the frame due to
the deformation of the surface. From the general identities δdr = dδr, δde j =
dδej , and equations (1)–(4), we can derive
δω1 = dv · e1 − ω2 Ω21 = dΩ1 + Ω2 ω21 + Ω3 ω31 − ω2 Ω21
δω2 = dv · e2 − ω1 Ω12 = dΩ2 + Ω1 ω12 + Ω3 ω32 − ω1 Ω12
(17)
δωij = dΩij + Ωil ωlj − ωil Ωlj
dv · e3 = dΩ3 + Ω1 ω13 + Ω2 ω23 = Ω13 ω1 + Ω23 ω2 .
These equations are the essential ingredients of the variational method based on
the moving frame. With them as well as equations (4) and (5), we can easily derive
δdA = (div v − 2HΩ3 )dA
δ(2H) = [∇2 + (4H 2 − 2K)]Ω3 + ∇(2H) · v (18)
˜ 3 + 2KHΩ3 + ∇K · v.
δK = ∇ · ∇Ω
Using the above equations (3)-(6), (9)-(11), (17) and (18), we can deal with almost
all variational problems on surfaces.

3. Lipid Vesicles
Most of lipid molecules are amphiphiles with a hydrophilic head group and two
hydrophobic hydrocarbon tails. When a quantity of lipid molecules disperse in
water, they will assemble themselves into a bilayer vesicle as depicted in Fig. 2
due to hydrophobic forces. In this section, we will theoretically understand various
configurations of lipid vesicles.

3.1. Shape Equation to Describe Equilibrium Configurations

A lipid vesicle can be represented as a closed surface M. Its equilibrium shape is

determined by minimizing Helfrich’s free energy (12) under the constraints of con-
stant area and volume because experiments reveal that the area of lipid membranes
 Geometry of Membranes 53

Figure 2. A cartoon of lipid vesicle.

are almost incompressible and the membranes are impermeable for the solutions in
both sides of the membranes. Thus we can introduce two Lagrange multipliers λ
and p to replace these two constraints, and then minimize the following functional
  
kc 2
F = (2H + Ih) + k̄K dA + λA + pV (19)
M 2

where M, A and V represent the membrane surface, total area of the vesicle and
volume enclosed by the vesicle. The multipliers λ and p can also be understood as
the apparent surface tension and osmotic pressure of the lipid vesicle.
The Euler-Lagrange equation for the functional (19) can be derived by using the
variational method presented in Section 2, which reads [27, 28, 42]

p̃ − 2λ̃H + (2H + Ih)(2H 2 − IhH − 2K) + 2∇2 H = 0 (20)

with reduced parameters p̃ = p/kc and λ̃ = λ/kc . This formula is called the
shape equation because it describes the equilibrium shapes of lipid vesicles and
represents the force balance along the normal direction of the membrane surfaces.

Figure 3. The generation curve for an axisymmetric vesicle.

54 Zhanchun Tu

Now let us consider an axisymmetric vesicle generated by a planar curve shown in

Fig. 3. Rotate the curve about the z− axis and then mirror with respect to the hori-
zontal plane. Here ρ is the revolving radius, and ψ is the angle between the tangent
of the curve and the horizontal plane. That is, dz/dρ = tanψ. Through simple
calculations, we can obtain −2H = h ≡ sin ψ/ρ + (sin ψ) , K = sin ψ(sin ψ) /ρ,
and ∇2 (2H) = −(cos ψ/ρ)(ρ cos ψh ) , where the ‘prime’ represents the deriva-
tive with respect to ρ. Substituting these relations into equation (20), we derive
h2 c0 h cos ψ
(h − c0 ) + − 2K − p̃ − λ̃h + (ρ cos ψh ) = 0. (21)
2 2 ρ
The equivalent form of this equation is first derived by Hu and Ou-Yang [14]. The
shape equation (21) of axisymmetric vesicles is a third-order differential equation.
Following Zheng and Liu’s work [55], we can transform it into a second order
differential equation
2η0 − p̃ρ2 tanψ
cos ψh + (h − c0 ) sin ψψ  − λ̃tanψ + − (h − c0 )2 = 0 (22)
2ρ cos ψ 2
where η0 is the integration constant. It is found that the shape equation of axisym-
metric vesicles obtained by Hu and Ou-Yang reduces into that derived by Seifert et
al. [37] when η0 = 0 in equation (22) or for vesicles with spherical topology free
of singular points [33, 55].

3.2. Analytic Solutions and Corresponding Configurations

Now we will show several analytic solutions to the shape equations (20) or (22)
of lipid vesicles that we have known till now. They correspond to the surfaces
of constant mean curvature, torus, biconcave discoid, and so on. Most of these
solutions are found by Ou-Yang and his coworkers. We will see that only sphere,
torus, and biconcave discoid can correspond to lipid vesicles. We also recommend
the reader to note the classic paper by Konopelchenko [17] who expressed equation
(20) in terms of the inverse mean curvature and density of squared mean curvature,
and then found a broad variety of its solutions and corresponding possible shapes.

3.2.1. Surfaces of Constant Mean Curvature

Obviously, surfaces of constant mean curvature (including sphere, cylinder, and

unduloid shown in Fig. 4) can satisfy the shape equation. First, let us consider a
spherical surface with radius R. Then H = −1/R and K = 1/R 2 . Substituting
them into equation (20), we derive
p̃R2 + 2λ̃R − Ih(2 − IhR) = 0. (23)
 Geometry of Membranes 55

This equation gives the relation between the sphere radius R, spontaneous curva-
ture Ih, reduced osmotic pressure p̃, and reduced surface tension λ̃.

Figure 4. Surfaces of constant mean curvature: sphere (left), cylinder (mid-

dle), and unduloid (right).

Next, H = −1/(2R) and K = 0 for a cylindrical surface with radius R. Then the
shape equation (20) requires

2p̃R3 + 2λ̃R2 − 1 + Ih2 R2 = 0. (24)

For other surfaces of constant mean curvature such as unduloid, H is a constant

but K is not a constant [20]. Via the shape equation (20), we obtain

H = −Ih/2 (25)

and p̃ = −λ̃Ih.
Note that cylindrical surface and unduloid are in fact not closed surfaces. Alexan-
drov proved that “an embedded surface (no self-intersection) with constant mean
curvature in E3 must be a spherical surface” [1]. Thus we can only observe one
kind of vesicles of constant mean curvature - spheres.

3.2.2. Torus

As shown in Fig. 5, a torus is a surface of revolution generated by a circle with

radius r rotated about an axis in the same plane of the circle. The revolving
radius R should be larger than r. The torus can be expressed in vector form
{(R + r cos ϕ) cos θ, (R + r cos ϕ) sin θ, r sin ϕ}. Through simple calculations,
we have 2H = −(R + 2r cos ϕ)/r(R + r cos ϕ) and K = cos ϕ/r(R + r cos ϕ).
56 Zhanchun Tu

Substituting them into equation (20), we derive

[(2Ih2 r2 − 4Ihr+4λ̃r2 + 2p̃r 3 )/ν 3 ] cos3 ϕ
+[(5Ih2 r2 − 8Ihr + 10λ̃r2 + 6p̃r 3 )/ν 2 ] cos2 ϕ
(26)
+[(4Ih2 r2 − 4Ihr + 8λ̃r2 + 6p̃r 3 )/ν] cos ϕ
+2/ν 2 + (Ih2 r2 − 1) + 2(p̃r + λ̃)r2 = 0
with ν = R/r. If ν is finite, then equation (26) holds if and only if the coefficients
of {1, cos ϕ, cos2 ϕ, cos3 ϕ} vanish. It follows that 2λ̃r = c0 (4−c0 r), p̃r 2 = −2c0
and √
ν = R/r = 2. (27)
That
√ is, there √ exists a lipid torus with the ratio of its two generation radii being
2 (called 2 torus by Ou-Yang [29]), which was confirmed in the experiment
[21]. It is also found
√ that nonaxisymmetric tori [36] constructed from conformal
transformations of 2 torus also satisfy the shape equation.

Figure 5. Torus (left) and its generation curve (right).

To check the consistency, we need also to verify that the above toroidal solution
indeed
√ satisfy shape equation (22) of axisymmetric vesicles. It is not hard to see
that 2 torus can be generated from a curve defined by
√
sin ψ = (p̃/r) ± 2. (28)
Substituting it into equation (22), we arrive at 2λ̃r = c0 (4 − c0 r), p̃r 2 = −2c0 and
η0 = −1/r = 0.

3.2.3. Biconcave Discoid

For 0 < IhρB < e, the parameter equations

sin ψ = −Ihρ ln(ρ/ρB )
ρ (29)
z = z0 + 0 tanψdρ
 Geometry of Membranes 57

corresponds to a planar curve shown in Fig. 6. Substituting it into equation (22),

we have p̃ = 0, λ̃ = 0, and η0 = 2Ih = 0. That is, a biconcave discoid generated by
revolving this planar curve about z-axis can satisfy the shape equation of vesicles.
This result can give a good explanation to the shape of human red blood cells under
normal physiological conditions [9, 22, 24].

Figure 6. Biconcave discoid (left) and its generation curve (right).

A small comment is that η0 = 2Ih = 0 reflects the singularity at two poles of

the biconcave discoid. Whether does this singularity exist in real red blood cells?
What is the biological meaning of this singularity? Or does there exist a normal
solution to the shape equation that can also explain the shape of red blood cells?
These open questions need further discussions.

3.2.4. Unduloid-Like and Cylinder-Like Surfaces

If we only concern solutions to the shape equation, two cases are also widely dis-
cussed. The first case [20,23] is an axisymmetric surface generated by planar curve
satisfying

1 ρ ρm 4
sin ψ = + − − 2, 0 < ρm Ih < 4/3. (30)
ρm Ih ρm ρ ρ2m Ih2

The generated surface abides by the shape equation with p̃ = −2Ihρ 4m , λ̃ = 2/ρ2m −
Ih2 /2, and η0 = 2Ih − 3/(Ihρ2m ). This surface has the unduloid-like shape but
nonconstant mean curvature.
The second one is a cylinder-like surface generated by a planar curve translating
along the normal of the plane. If we denote the curvature of the curve as κ, then
the geometric quantities of the generated surface can be expressed as 2H = −κ,
K = 0, and ∇2 (2H) = −κ̈, where the ‘dot’ above κ represents the derivative with
respect to the arc length of the curve. Thus the shape equation (20) degenerates
into [2, 49, 54]
p̃ + λ̄κ + κ3 /2 − κ̈ = 0 (31)
58 Zhanchun Tu

with λ̄ = λ̃ + Ih2 /2. The above equation is integrable, which results in

κ̇2 = ξ0 + 2p̃κ + λ̄κ2 − κ4 /4 (32)

with an integral constant ξ0 . This equation can be further solved in terms of elliptic
functions [2, 49, 54, 56].
It is necessary to note that these two cases do not correspond to real vesicles be-
cause they are not closed surfaces.

4. Lipid Membranes with Free Edges

The opening-up process of liposomal membranes by talin [35] was observed, which
gives rise to the study of equilibrium equation and boundary conditions of lipid
membranes with free exposed edges. This problem was theoretically investigated
by Capovilla et al. [5] and Tu et al. [41] in terms of different methods. In this
section, we will present these theoretical results and subsequent advancements.

Figure 7. An open smooth surface M with a boundary curve C. {e 1 , e2 , e3 }

forms the frame at some point on the surface. {t, b, e3 } also forms the right-
handed frame for the point in C such that t is the tangent of C and b points
to the side that the surface is located in.

4.1. Shape Equation and Boundary Conditions to Describe Equilibrium Con-

figurations

As shown in Fig. 7, a lipid membrane with a free edge can be visualized as an

open smooth surface M with a boundary curve C in geometry. Because the free
exposed edge is energetically unfavorable, we assign the line tension (energy cost
 Geometry of Membranes 59

per length) to be γ > 0. Then the free energy functional that we need to minimize
can be expressed as
  
kc 2
F = (2H + Ih) + k̄K dA + λA + γL (33)
M 2
where L is the total length of the free edge.
By using the variational method introduced in Section 2, we can arrive at the shape
equation [41]
(2H + c0 )(2H 2 − c0 H − 2K) − 2λ̃H + ∇2 (2H) = 0 (34)
and three boundary conditions [41]


[(2H + c0 ) + k̃κn ] = 0 (35)
C

[−2∂H/∂b + γ̃κn + k̃ τ̇g ] = 0 (36)
C
2 
[(1/2)(2H + c0 ) + k̃K + λ̃ + γ̃κg ] = 0 (37)
C
where λ̃ ≡ λ/kc , k̃ ≡ k̄/kc , and γ̃ ≡ γ/kc are the reduced surface tension, reduced
bending modulus, and reduced line tension, respectively. κ n , κg , and τg are the
normal curvature, geodesic curvature, and geodesic torsion of the boundary curve,
respectively. The ‘dot’ represents the derivative with respect to the arc length of
the edge. Equation (34) expresses the normal force balance of the membrane.
Equations (35)–(37) represent the force and the moment balances at each point in
curve C. Thus, in general, the above four equations are independent of each other
and applicable for an open membrane with several edges.
Now we consider the axisymmetric membranes. As before when a planar curve AC
shown in Fig. 8 revolves about the z axis, an axisymmetric surface is generated.
Let ψ represent the angle between the tangent line and the horizontal plane. Each
point in the surface can be expressed as a vector r = {ρ cos φ, ρ sin φ, z(ρ)}, where
ρ and φ are the radius and azimuth angle that the point corresponds to. Introduce a
notation σ such that σ = 1 if t is parallel to ∂r/∂φ, and σ = −1 if t is antiparallel
to ∂r/∂φ on the boundary curve generated by point C. The above equations (34)–
(37) are transformed into
h2 c0 h cos ψ
(h − c0 ) + − 2K − λ̃h + (ρ cos ψh ) =0 (38)
2 2 ρ
 
h − c0 + k̃sin ψ/ρ =0 (39)
 C
−σ cos ψh + γ̃sin ψ/ρ C =0 (40)
 
1 2 cos ψ
(h − Ih) + k̃K + λ̃ − σγ̃ =0 (41)
2 ρ C
60 Zhanchun Tu

z
C

y
A

r
Figure 8. The profile curve of the axisymmetric surface. Such surface can
be generated by a planar curve AC rotating about z axis. The angle between
the tangent line and the horizontal plane is ψ.

with h ≡ sin ψ/ρ + (sin ψ) and K ≡ sin ψ(sin ψ) /ρ. The ‘prime’ represents the
derivative with respect to ρ.
The shape equation (38) is integrable, which reduces to a second order differential
equation

η0 tanψ
cos ψh + (h − c0 ) sin ψψ  − λ̃tanψ + − (h − c0 )2 = 0 (42)
ρ cos ψ 2
in which η0 is the integration constant [47]. This equation is equivalent to equa-
tion (22) with zero osmotic pressure. The configuration of an axisymmetric open
lipid membrane should satisfy shape equation (42) and boundary conditions (39)–
(41). In particular, the points on the boundary curve should satisfy not only the
boundary conditions, but also the shape equation (42) because they also locate in
the surface. That is, equations (39)-(41) and (42) should be compatible with each
other in the edge. Substituting equations (39)-(41) into (42), we derive the com-
patibility condition [47] to be
η0 = 0. (43)
It is a necessary (but not sufficient) condition for existence of axisymmetric open
membranes. Under this condition, the shape equation is reduced to

tanψ
cos ψh + (h − c0 ) sin ψψ  − λ̃tanψ − (h − c0 )2 = 0 (44)
2
while three boundary conditions are reduced to two equations, i.e., equations (39)
and (41).
 Geometry of Membranes 61

4.2. Theorems of Non-Existence

Now our task is to find analytic solutions that satisfy both the shape equation and
the boundary conditions. An obvious but trivial one is a circular disk with radius
R. In this case, equations (34)–(37) degenerate to
λ̃R + γ̃ = 0. (45)

Can we find nontrivial analytic solutions? We will prove several theorems of non-
existence in this subsection, which imply that it is almost hopeless to find nontrivial
analytic solutions.

Theorem 1. There is no open membrane being a part of a spherical vesicle.

Proof: For a sphere with radius R, we can calculate H = −1/R, κ n = −1/R and
τg = 0 in terms of equation (6) because a = c = −1/R and b = 0 for a sphere.
Boundary condition (36) cannot be abided by. Thus an open membrane cannot be
a part of a spherical vesicle. 

Theorem 2. There is no open membrane being a part of a cylindrical surface.

Proof: For any line element on the surface of a cylinder with radius R, we can
calculate κn = − cos2 θ/R from equation (7) where θ is the angle between the line
element and the circumferential direction. Additionally, H = −1/R is a constant.
If k̃ = 0, then boundary condition (36) results in κn = 0, that is θ = π/2. The
line along this direction is not a closed curve, and so cannot be as an edge of a
membrane. If k̃ = 0, then boundary condition (35) results in κn = (Ih − 1/R)/k̃,
which implies θ should be a constant. The unique closed curve is a circle, i.e.,
θ = 0 and κn = −1/R. But τg = 0 if θ = 0, then contradicts with boundary
condition (36). Thus an open membrane cannot be a part of a spherical surface. 

Theorem 3. There is no open membrane being a part of a curved surface with

constant mean curvature.

Proof: Two special surfaces (sphere and cylinder) with constant mean curvature
are discussed in the above theorems. Now we only need to investigate surfaces
with constant H but nonconstant K. From shape equation of open membranes, we
derive two possible cases: i) H = −Ih/2 = 0 and λ̃ = 0; ii) H = Ih = 0 and
λ̃ = 0.
In the former case, if k̃ = 0, then boundary conditions (36) and (37) result in
κn = κg = 0. Thus the curvature of the boundary curve is κ = κ2n + κ2g = 0.
62 Zhanchun Tu

That is, this curve is a straight line which is not closed curve. If k̃ = 0, then
boundary conditions (35) and (36) results in κn = 0 and τg = constant. Using
equation (7), we derive the principal curvatures for the points in the curve are
constant. Then equation (37) requires κg = constant. That is, the curvature
and torsion are constant in the curve. The unique closed curve is a circle. But
τg = 0 for a circle. Let c1 and c2 represent the two principal curvatures, θ is the
angle between the tangent of the curve and one principal direction at each point
in the curve. Then we have two equations: κn = c1 cos2 θ + c2 sin2 θ = 0 and
τg = (c2 − c1 ) sin θ cos θ = 0. Substituting these two equations and K = c1 c2
into equation (37), we obtain κg = 0. Then κ = κ2n + κ2g = 0, which contradicts
with the preassumption of a circle.
In the latter case, H = Ih = 0, similar to the proof in the former one, it also leads
to a contradiction. Thus there is no open membrane being a part of a curved surface
with constant mean curvature. 

Theorem 4. There is no open membrane being a part of a Willmore surface.

Proof: Let us consider the scaling transformation r → Λr, where the vector r
represents the position of each point in the membrane and Λ is a scaling parameter
[6]. Under this transformation, we have A → Λ2 A, L → ΛL, H → Λ−1 H, and
K → Λ−2 K. Thus, equation (33) is transformed into

F (Λ) = [(kc /2)(2H)2 + k̄K]dA
M  (46)
2 2
+2kc IhΛ HdA + (λ + kc Ih /2)Λ A + γΛL.
M

The equilibrium configuration should satisfy ∂F/∂Λ = 0 when Λ = 1. Thus we

obtain 
2Ih HdA + (2λ̃ + Ih2 )A + γ̃L = 0. (47)
M
This equation is an additional constraint for open membranes.
Willmore surfaces satisfy the special form of equation (20) with vanishing λ̃ and
Ih [51]. Because γ̃L > 0, thus the constraint (47) cannot be satisfied when λ̃ = 0
and Ih = 0. That is, there is no open membrane being a part of Willmore surface.

√
Corollary There is no open membrane being a part of 2 torus.
√
Proof: Substituting
√ 2 torus into equation (34), we obtain Ih = 0 and λ̃ = 0.
That is, 2 torus is a Willmore surface. In terms of Theorem 4, we arrive at this
corollary. 
 Geometry of Membranes 63

Figure 9. Schematics of several impossible open membranes with free

edges: parts of sphere, cylinder, unduloid, torus, biconcave discodal surface
(from left to right).

Theorem 5. There is no axisymmetric open membrane being a part of a biconcave

discodal surface generated by a planar curve defined by the Gauss map sin ψ =
−Ihρ ln(ρ/ρB ).

Proof: A biconcave discodal surface [22,24] generated by a planar curve specified

by sin ψ = −Ihρ ln(ρ/ρB ) with non-vanishing constants Ih and ρB . To avoid the
singularity at the two poles, we may dig two holes around the poles. Substituting
this equation into the shape equation (42), we obtain λ̃ = 0 and η0 = 2Ih. That
is, the biconcave discodal surface can be a solution to the shape equation. How-
ever, η0 = 2Ih = 0 contradicts to compatibility condition (43). Thus there is no
axisymmetric open membrane being a part of this biconcave discoidal surface. 
In Fig. 9, we show several impossible open membranes with free edges in terms
of the above theorems. These theorems suggest that it is hopeless to find exactly
analytic solutions to the shape equation and boundary conditions of open lipid
membranes. Quasi-exact solutions or numerical simulations are highly appreci-
ated.

4.3. Quasi-Exact Solutions

Here the quasi-exact solution is defined as a surface with free edge(s) such that
the points on that surface exactly satisfy the shape equation, and most of points in
the edge(s) abide by boundary conditions. In fact, the proofs of Theorem 2 and
Theorem 3 implies two possible solutions as shown in Fig. 10. One is a straight
stripe along the axial direction of cylinder, another is a twist ribbon which is a part
of a minimal surface (H=0).
Let we consider a long enough straight stripe along the axial direction of cylinder
that satisfies shape equation (34), that is, λ̃ = (1 − Ih2 R2 )/2R2 . The long enough
configuration ensure us to omit the boundary of two ends. The lateral edges are
64 Zhanchun Tu

Figure 10. Schematics of two quasi-exact solutions: straight stripe along

axial direction of cylinder (left) and twist ribbon which is a part of a minimal
surface (right).

straight lines which have κn = κg = τg = 0. Thus boundary conditions (35) and

(36) are trivial. The third boundary condition results in λ̃ = (1 − IhR)2 /2R2 . Thus
we arrive at λ̃ = 0 and R = 1/Ih. That is, a long enough straight stripe along the
axial direction of a cylinder with R = 1/Ih is a quasi-exact solution.
Next, a twisted ribbon with pitch T and width 2u0 can be expressed in the vector
form {u cos ϕ, u sin ϕ, αϕ} with |u| ≤ u0 , |ϕ| < ∞ and |α| = T /2π. By the
simple calculations, we have
H = 0, K = −α2 /(u2 + α2 )2 (48)
for the points on the surface, and
κn = 0, K = −α2 /(u20 + α2 )2
κg = u0 /(u20 + α2 ) (49)
τg = α/(u20 + α2 )
for the points on the edges.
It is easy to see that equation (48) can satisfy shape equation (34) when Ih = 0.
Then equation (49) naturally validates boundary conditions (35) and (36). The last
boundary condition (37) leads to λ̃ = [k̃α2 − γ̃u0 (u20 + α2 )]/(u20 + α2 )2 , which
can be satisfied by a proper choice of the parameters λ̃, k̃, γ̃, u0 and α. That is, the
twist ribbon is indeed a quasi-exact solution.

5. Chiral Lipid Membranes

In fact, our above discussions only concern lipid membranes where lipid molecules
are in Smectic A phase. In this phase, lipid molecules almost point to the normal
direction of the membrane surface. However, there are also many kinds of chiral
lipids in cell membranes. At body temperature, chiral lipids usually form Smec-
tic C∗ phase. They are tilting from the normal direction in a constant angle. It is
necessary to develop Helfrich’s spontaneous curvature model introduced in Sec-
tion 2.2 to cover the Smectic C∗ phase. Based on symmetric argument or Frank
 Geometry of Membranes 65

energy in the theory of liquid crystal, many theoretical models and results were
achieved [12,16,26,30,31,39,40]. These theoretical models contain much compli-
cated terms and many parameters, which make it is impossible to derive the exact
governing equations for describing equilibrium configurations of chiral lipid mem-
branes. Here we will discuss a simplified version proposed by the present author
and Seifert [45]. It is found that this concise theory can still explain most of the
experimental phenomena.

5.1. Constructing the Free Energy

The free energy density for a chiral lipid membrane are supposed to consist of the
following contributions.

i) The bending energy per area is still taken as Helfrich’s form (14). That is,
we neglect the anisotropic effect of lipid molecules’ tilting on the bending
moduli.

Figure 11. Right-handed orthonormal frame {e1 , e2 , e3 } at any point in

a surface where e3 is the normal vector of the surface. a) Surface without
boundary curve. b) Surface with boundary curve where t is the tangent
vector of the boundary curve, and b, in the tangent plane of the surface, is
perpendicular to t.

ii) The energy per area originating from the chirality of tilting molecules has
the form [30]
fch = −hτm (50)
where h reflects the strength of molecular chirality. Without losing the gen-
erality, here we only discuss the case of h > 0. τm is the geodesic torsion
along the unit vector m at each point. Here m represents the projected
direction of the lipid molecules on the membrane surface. If we take a
right-handed orthonormal frame {e1 , e2 , e3 } as shown in Fig. 11, m can
66 Zhanchun Tu

be expressed as m = cos φe1 + sin φe2 , where φ is the angle between m

and e1 . then geodesic torsion τm and normal curvature κm along m can be
expressed in the similar form of equation (6).

(iii) The energy per area due to the orientational variation is taken as

fov = (kf /2)[(∇ × m)2 + (∇ · m)2 ] (51)

where kf is a constant in the dimension of energy. This is the simplest term

of energy cost due to tilting order invariant under the coordinate rotation
around the normal of the membrane surface. By defining a spin connection
field S such that ∇.S = K, one can derive (∇×m)2 +(∇·m)2 = (∇φ−S)2
through simple calculations [25].

The total free energy density adopted in the present paper, G = f c + fch + fov , has
the following concise form
kc kf
G= (2H + c0 )2 + k̄K − hτm + v2 (52)
2 2
with v ≡ ∇φ − S. This special form might arguably be the most natural and
concise construction including the bending, chirality and tilting order, for the given
vector field m and normal vector field e3 .

5.2. Governing Equations to Describe Equilibrium Configurations

The free energy for a closed chiral lipid vesicle may be expressed as

F = GdA + λA + pV (53)
M

where A is the area of the membrane and V the volume enclosed by the vesicle.
Here again λ and p are two multipliers to implement area and volume constraints.
Using the variational method mentioned in Section 2, we can obtain two governing
equations to describe equilibrium configurations [45] as

2h̃(κm − H) − k̃f ∇2 φ = 0 (54)

and

2∇2 H + (2H + c0 )(2H 2 − c0 H − 2K) − 2λ̃H + p̃ + h̃[∇ · (m∇ × m)

(55)
+ ∇ × (m∇ · m)] + k̃f [(κv − H)v2 − ∇v : ∇e3 ] = 0
 Geometry of Membranes 67

with reduced parameters h̃ = h/kc , k̃f = kf /kc , p̃ = p/kc , and λ̃ = λ/kc . κm and
κv are the normal curvature along the directions of m and v, respectively. When
writing equation (54), we have selected the proper gauge such that ∇ · S = 0, or
else ∇2 φ should be replaced with ∇2 φ − ∇ · S. Additionally, we do not consider
singular points for closed vesicles different from toroidal topology.
Consider a chiral lipid membrane with a free edge as shown in Fig. 11b. Its free
energy can be expressed as

F = GdA + λA + γL (56)
M
where A is the area of the membrane and L the total length of the edge and γ
represents the line tension of the edge.
Using the variational method mentioned in Section 2, we can obtain the governing
equations to describe equilibrium configurations of membrane surfaces as
2h̃(κm − H) − k̃f ∇2 φ = 0 (57)
and
2∇2 H + (2H + c0 )(2H 2 − c0 H − 2K) − 2λ̃H + p̃ + h̃[∇ · (m∇ × m)
(58)
+ ∇ × (m∇ · m)] + k̃f [(κv − H)v2 − ∇v : ∇e3 ] = 0.
Simultaneously, the boundary conditions obeyed by the free edge are derived as
[45]
vb = 0 (59)
2 2
(1/2)(2H + c0 ) + k̃K − h̃τm + (k̃f /2)v + λ̃ + γ̃κg = 0 (60)
(2H + c0 ) + k̃κn − (h̃/2) sin 2φ̄ = 0 (61)
˙ sin 2φ̄ + k̃ κ v = 0
γ̃κn + k̃ τ̇g − 2∂H/∂b − h̃(vt + φ̄) (62)
f n t

where κn , τg and κg are the normal curvature, geodesic torsion, and geodesic cur-
vature of the boundary curve (i.e., the edge), respectively. Here v b and vt are the
components of v in the directions of b and t. The ‘dot’ denotes the derivative
with respect to arc length parameter s and φ̄ denotes the angle between m and t at
the boundary curve. Equations (59)–(62) describe the force and moment balance
relations in the edge. Thus they are also available for a chiral lipid membrane with
several edges.

5.3. Solutions and Corresponding Configurations

Now we will present some analytic solutions to the governing equations of chiral
lipid membranes.
68 Zhanchun Tu

5.3.1. Sphere

For spherical vesicles of chiral lipid molecules with radius R, τm is always van-
ishing because a = c = 1/R and b = 0. Thus the free energy (53) is independent
of the molecular chirality and permits the same existence probability of left- and
right-handed spherical vesicles. This is uninteresting case in practice.

5.3.2. Cylinder

Here we consider a long enough cylinder with radius R such that its two ends
can be neglected. The cylinder can be parameterized by two variables s and z
which are the arc length along the circumferential direction and coordinate along
axial direction, respectively. Let φ be the angle between m and the circumferential
direction. Then equations (54) and (55) are transformed into [45]

k̃f (φss + φzz ) + (h̃/R) cos 2φ = 0 (63)

and

h̃[2(φ2z − φ2s + φsz ) sin 2φ + (φss − φzz + 4φz φs ) cos 2φ] + λ̃/R
(64)
+ p̃ + (c20 − 1/R2 )/2R + k̃f [(φ2z − φ2s )/2R + φsz /R] = 0

where the subscripts s and z denote the partial derivatives respect to s and z, re-
spectively.
It is not hard to see that φ = π/4 and 2p̃R3 + 2λ̃R2 − 1 + Ih2 R2 = 0 can satisfy
the above two equations. Thus a cylinder shown in Fig. 12a with uniform tilting
state (tilting angle φ = π/4) is a solution.

Figure 12. Two possible chiral lipid membranes: a) Cylinder with uniform
tilting state; b) Torus with uniform tilting state.
 Geometry of Membranes 69

5.3.3. Torus

A torus is a surface of revolution generated by a circle with radius r rotated about

an axis in the same plane as shown in Fig. 5. It can be presented in the vector
formes {(R + r cos ϕ) cos θ, (R + r cos ϕ) sin θ, r sin ϕ}. Equation (54) is trans-
formed into [45]
 
1 ∂2φ ∂ ∂φ ν h̃r
2
+ (ν + cos ϕ) − cos 2φ = 0 (65)
ν + cos ϕ ∂θ ∂ϕ ∂ϕ k̃f

where φ is the angle between m and the latitude of the torus, while ν ≡ R/r is the
ratio between two generated radii of the torus.

The uniform tilting state (φ = −π/4) satisfies equation (65) and makes − hτm dA
to take the minimum. With φ = −π/4, equation (55) is transformed [45] into

(2 − k̃f )/ν 2 + (Ih2 r2 − 1) + 2(p̃r + λ̃)r2

+ [(4Ih2 r2 − 4Ihr − 2h̃r + 8λ̃r2 + 6p̃r 3 )/ν] cos ϕ
(66)
+ [(5Ih2 r2 − 8Ihr − 4h̃r + 10λ̃r2 + 3k̃f + 6p̃r 3 )/ν 2 ] cos2 ϕ
+ [(2Ih2 r2 − 4Ihr − 2h̃r + 4λ̃r2 + 2k̃f + 2p̃r 3 )/ν 3 ] cos3 ϕ = 0

Because ν is finite for a torus, then the above equation holds if and only if the
coefficients in front of {1, cos ϕ, cos2 ϕ, cos3 ϕ} vanish. It follows that 2λ̃r2 =
(4rc0 − r2 c20 ) − 3k̃f + 2h̃r, p̃r 3 = 2k̃f − 2rc0 − h̃r and

ν= (2 − k̃f )/(1 − k̃f ). (67)

Thus a torus with uniform tilting state as shown in Fig. 12b is an exact solution
to governing equations of chiral lipid vesicles. The ratio of two generation
√ radii
satisfies equation (67),
√ which increases with k̃ f . Especially, ν = 2 for k̃ f = 0,
which leads to the 2 torus of non-tilting lipid molecules √[29]. Since this kind of
torus was observed in the experiment [21], tori with ν > 2 for 0 < k̃f < 1 might
also be observed in some experiments on chiral lipid membranes.

5.3.4. Twisted Ribbons

Here we consider a quasi-exact solution for the governing equations to describing

equilibrium configurations of chiral lipid membranes with free edges. Two long
enough twisted ribbons with lipid molecules in different tilting states are shown
in Fig. 13. Similar to Section 4.3, a twist ribbon can be expressed as vector form
70 Zhanchun Tu

{u cos ϕ, u sin ϕ, αϕ} with |u| ≤ W/2, |ϕ| < ∞ and |α| = T /2π. Equation (57)
is transformed into [45]

uφu + φϕϕ 2h̃α sin 2φ

k̃f φuu + + =0 (68)
u2 + α 2 u2 + α 2

where φ is the angle between m and the horizontal.

Figure 13. Long enough twisted ribbons with lipid molecules in different
tilting states: a) m is perpendicular to the edges; b) m parallels the edges.
Arrows represent the projected directions {m} of the tilting molecules on
the ribbons’ surface.

If we only consider the uniform tilting state, theabove equation requires φ = 0 or

 see that φ = 0 minimizes −h τm dA for α < 0 while
π/2. It is easy to φ = π/2
minimizes −h τm dA for α > 0 because τm = −α cos 2φ/(u + α2 ) [45]. 2

Thus we should take φ = 0 for α < 0 and φ = π/2 for α > 0. The former
case corresponds to Fig. 13a where m is perpendicular to the edges and the latter
corresponds to Fig. 13b where m is parallel to the edges. Both for φ = 0 and π/2,
equation (58) leads to Ih = 0 for non-vanishing α. Among the boundary conditions
(59)–(62), only equation (60) is nontrivial, which reduces to

k̃f x2 − 2k̃
λ̃(1 + x2 )α2 − (h̃ − γ̃x)|α| + =0 (69)
2(1 + x2 )

with x ≡ W/2|α|. Solutions to this equation exists for proper parameters. Thus,
there are indeed twisted ribbons in two states as shown in Fig. 13, they have differ-
ent chirality and tilting angles.
 Geometry of Membranes 71

6. Summary and Conclusions

In the above discussions, we have presented some theoretical results on the geom-
etry of membranes, which include the surface theory and variational method based
on moving frame, the governing equations to describe equilibrium configurations
of various lipid structures derived from the variation of free energy functionals,
some analytic solutions to these equations and their corresponding configurations.
We only focus on the pure theoretical researches and miss all experimental and
numerical results related to our topic on which gentle readers may consult the Ref-
erences [3, 8, 9, 13, 18, 48, 50, 52].
Although many theoretical advancements have been achieved, there are still a lot
of challenges waiting for further investigations. Several key open questions among
them are listed below:

i) Lipid vesicles of multi-components. Cell membranes contains many kinds

of lipids. At body temperature, different kinds of lipids usually separate
into several lipid domains. Lipid vesicles with two or several domains have
been investigated from experimental and theoretical points of view [3, 10,
15, 42, 46, 50, 53]. However, there is still lack of strictly exact solutions to
the governing equations [42,46] describing the vesicles with multi-domains.

ii) Other solutions on the shape equations of lipid membranes. We have found
only a few analytic solutions to the governing equations of lipid structures.
Whether are there other solutions, in particular to the simplest equations
(20) and (22)? Or can we prove that there is no other analytic solutions
except the ones that we have mentioned?

iii) Generalized boundary conditions for open lipid membranes. Although we

have investigated the boundary conditions of lipid membranes with free
edges, there are still other cases, such as confined edges, contact lines, and
so on. Can we develop a generalized variational principle covering such
cases?

iv) Non-orientable membranes. All membranes that we have considered are ori-
entable membranes. How can we deal with the non-orientable membranes,
such as Möbius band [57]?
72 Zhanchun Tu

Acknowledgements
The author is grateful to the financial supports from the Nature Science Founda-
tion of China (grant # 10704009), the Foundation of National Excellent Doctoral
Dissertation of China (grant # 2007B17) and the Fundamental Research Funds for
the Central Universities.

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Zhanchun Tu
Department of Physics
Beijing Normal University
Beijing 100875
PR CHINA
E-mail address: tuzc@bnu.edu.cn