Equatorial deformation of homogeneous spherical fluid vesicles by a rigid ring

Pablo Vázquez-Montejo,1, ∗ Bojan Božič,2, † and Jemal Guven3, ‡

1
SECIHTI - Facultad de Matemáticas, Universidad Autónoma de Yucatán,
Periférico Norte, Tablaje 13615, 97110, Mérida, Yucatán, México
2
Institute of Biophysics, Faculty of Medicine, University of Ljubljana, Vrazov trg 2, SI-1000 Ljubljana, Slovenia
3
Instituto de Ciencias Nucleares, Universidad Nacional Autónoma de México,
Apdo. Postal 70-543, 04510, Coyoacán, Ciudad de México, México
We examine the deformation of homogeneous spherical fluid vesicles along their equator by a
circular rigid ring. We consider deformations preserving the axial and equatorial mirror symmetries
of the vesicles. The configurations of the vesicle are determined employing the spontaneous curvature
model subject to the constraints imposed by the ring as well as of having constant area or volume.
arXiv:2412.17940v1 [cond-mat.soft] 23 Dec 2024

We determine two expressions of the force exerted by the ring, one involving a discontinuity in
the derivative of the curvature of the membrane across the ring, and another one in terms of the
global quantities of the vesicle. For small enough values of the spontaneous curvature there is only
one sequence of configurations either for fixed area or volume. The behavior of constricted vesicles
is similar for both constraints, they follow a transition from prolate to dumbbell shapes, which
culminates in two quasispherical vesicles connected by a small catenoid-like neck. We analyze the
geometry and the force of the small neck employing a perturbative analysis about the catenoid. A
stretched vesicle initially adopts an oblate shape for either constraint. If the area is fixed the vesicle
increasingly flattens until it attains a disclike shape, which we examine using an asymptotic analysis.
If the volume is fixed the poles approach until they touch and the vesicle adopts a discocyte shape.
When the spontaneous curvature of the vesicle is close to the mean curvature of the constricted
quasi-spherical vesicles, the sequences of configurations of both constraints develop bifurcations,
and some of their configurations have the lowest energy.

I. INTRODUCTION In this work, as an approach to model the relevant phys-

ical aspects of this kind of cellular processes, we exam-
Many biophysical processes involve the interaction of ine the response of an homogeneous spherical fluid mem-
biopolymers with membranes [1]. Transmembrane pro- brane of radius RS and spontaneous curvature Cs under
teins as well as proteins docking on the membrane sur- the action of a rigid polymer loop of radius R0 binding
face will tend to deform the membrane geometry locally, along its equator. The deformed vesicles are character-
which in turn will mediate interactions between proteins. ized by the ratio r0 = R0 /RS : if r0 < 1 (r0 > 1) the
Dramatic large but still localized deformations may also ring constricts (stretches) the vesicle along its equator.
occur, as is strikingly illustrated by tethers formation Both deformations have biophysical relevance: the con-
[2–4]. Proteins can also bind to the membrane and as- striction of the membrane is an important step in the
semble into a filament or a protein complex, giving rise to fission process induced by a protein complex, whereas
stresses that induce large deformations, which may play a the stretching of the membrane might represent the situ-
role in shaping the global morphology of the membrane, ation when outward traction is applied to the membrane
as well as changing its topology. The prototypical exam- or the confinement of a loop within the vesicle, scenario
ple in the formation of vesicles is the endocytosis, driven that contrasts with the very different limit studied previ-
by the helical protein dynamin, which is also involved in ously in which a semi-flexible polymer is confined within
the process of cellular division in certain bacteria. The a rigid sphere [10–12]. Although this model greatly sim-
dynamin self-assembles into a helical elastic filament on plifies the biophysics, it provides several pointers to a
the narrow neck of a membrane, and by applying an in- better understanding of the different stages of such cellu-
ward traction promotes its fission [5, 6]. Another impor- lar processes, in particular, it permits us to quantify the
tant example is the cytokinesis, in which the ESCRT-III force deforming the vesicle.
protein polymerizes on the cell membrane forming a con- We determine the configurations of the membrane em-
tractile ring, which exerts a force that produces its divi- ploying the spontaneous curvature model subject to the
sion into two new cells [7, 8]. Also, by employing Monte local constraint that the equator conforms to the ring,
Carlo simulations it has been shown that the mechanical and to the global constraints that either the total area or
constriction of vesicles could be driven by synthetic con- the total volume of the membrane are fixed. The Euler-
tractile rings composed of nanoparticles [9]. Lagrange (EL) equation of a fluid membrane, whose so-
lutions describe the equilibrium configurations, can be
expressed in terms of the conservation law of the stress
tensor, which is also determined by the geometry of the
∗ pablo.vazquez@secihti.mx; pablo.vazquez@correo.uady.mx membrane [13–20]. In Sec. II we present how the con-
† bojan.bozic@mf.uni-lj.si straint imposed by the ring can be implemented in the
‡ jemal@nucleares.unam.mx
 2

variational principle by means of a vectorial Lagrange

multiplier, which modifies the boundary terms.
For the sake of simplicity, we consider deformations pre-
serving the axial symmetry, as well as the mirror sym-
metry with respect to the equatorial plane. The axial
symmetry implies the existence of a first integral of the
EL equation: the integration of the projection of conser-
vation law of the stress tensor along the axis of symmetry
provides a direct derivation [16], which we present in Sec.
III for the spontaneous curvature model. The presence
of the ring at the equator sets the boundary condition
for the radial coordinate of the membrane. Furthermore, (a) r0 = 0.95 (b) r0 = 0.845 (c) r0 = 0.5
the condition of finite energy demands the continuity of
the tangent planes across the equator and at the poles,
such that kinks do not occur. To examine the response
of the membrane to the force exerted by the ring, the
first integral is solved numerically with these boundary
conditions in the two source-free regions into which the
membrane is partitioned.
The force exerted by the ring induces a distribution of
stresses on the membrane. If the force is negative (pos-
itive) the ring exerts an inward (outward) radial force.
In Sec. IV we examine the local and global effects of the
force exerted by the ring on the surface geometry. First, (d) r0 = 10−1 (e) r0 = 10−2 (f) r0 = 10−3
from the balance of stresses at the equator we find that
the external force originates a discontinuity in the arc-
length derivative of the meridian curvature. Also, from
the scale invariance of the energy we obtain the external
force in terms of the global quantities of the membrane FIG. 1. (a)-(e) Sequence of constricted vesicles with fixed area
and the Lagrange multipliers fixing area and volume. and Cs = 0. (e) At the maximum constriction, the vesicle
The length scale is given by the radius of the spherical consists of two quasi-spherical surfaces connected by a small
vesicle RS . In Sec. V we define the geometric and phys- catenoid-like neck. (f) Magnification of the small neck. The
ical quantities rescaled with appropriate powers of RS bending energy density is color coded in these figures.
that we employ in the discussion of the deformed config-
urations of the membranes.
We examine the equilibrium shapes of membranes with apparently adopts a catenoidal shape in this limit (shown
different values of spontaneous curvature and with their in Fig. 1(f)). However, since the catenoid is a minimal
area or volume fixed in Secs. VI and VII respectively. surface it satisfies the source-free EL equation, so it can-
For membranes with spontaneous curvature lower than not represent a constricted small neck, for which a finite
certain values we obtain a unique configuration for each force persists. A more careful analysis employing pertur-
value of the equatorial radius. bation theory reveals that the geometry of the small neck
The behavior of constricted membranes (0 < r0 < 1), il- is well approximated by a first order perturbation of the
lustrated in Fig. 1, is similar for fixed area or volume, dif- catenoid, which modifies its mean curvature such that
fering only in their size. Slightly constricted membranes it alternates between the spontaneous curvature of the
adopt a prolate shape (Fig. 1(a)), followed by a transi- membrane and the mean curvature of the quasi-spherical
tion to a non-convex dumbbell geometry (Figs. 1(b)-(c)). vesicles, making it compatible with a finite equatorial
Upon further constriction, a second transition occurs as- force. In addition, the perturbative analysis provides the
sociated with the conversion to a pair of quasi-spherical force exerted on these small necks, which is proportional
vesicles connected by an infinitesimal neck (Figs. 1(d)- to the difference of between the mean curvature of the
(e)). The constriction involves a finite inward force, but quasispherical vesicles and the spontaneous curvature of
its dependence on the equatorial radius is not monotonic, the membrane.
it varies between a local minimum and local maximum
values before decreasing towards a finite value in the limit The sequences of stretched vesicles (r0 > 1) with fixed
of the very small neck. Usually, to analyze such necks it is area and fixed volume are illustrated in Figs. 2 and 3 re-
assumed that they can be described by a section of an un- spectively. Slightly stretched membranes adopt an oblate
duloid (a constant mean curvature surface) and the cor- shape for both constraints (Figs. 2(a)-(b) and 3(a)-(b)),
responding results are compared with numerical results but as the equatorial radius increases the shapes of vesi-
[21–24]. By looking at the numerical solutions the neck cles with fixed area and fixed volume become very dis-
similar. If the area is fixed, the membrane flattens more
 3

either constraint, the increase of the spontaneous curva-

ture to values close to the mean curvature of the quasi-
spherical vesicles, obtained in the limit of maximum con-
striction, gives rise to bifurcations in the solutions of the
EL equation. For both constraints the bifurcations be-
gin with different constricted vesicles and for the case
(a) r0 = 1.1 (b) r0 = 1.2 of fixed area they end with a common stretched vesicle,
whereas for the case of fixed volume they end with dif-
ferent stretched configurations. In both cases, below a
certain value of the equatorial radius the configurations
belonging to one of the bifurcations possess lower total
(c) r0 = 1.3 (d) r0 = 1.4 bending energy than the configurations of the original
sequence of constricted vesicles with the same equatorial
radius. The corresponding sequence of configurations is
illustrated in Fig. 4 for fixed area (the sequence for fixed
volume is similar). It begins with a dumbbell shape (Fig.
FIG. 2. (a)-(d) Sequence of stretched vesicles with fixed area 4(a)), as the vesicle is stretched its central region bulges,
and Cs = 0. (d) At the maximum stretching, the vesicle resembling an onduloid (Figs. 4(b)-(c)), which morphs
consists of two discs joined along their boundary. The bending into a central oblate shape connected by small necks to
energy density is color coded in these figures. two quasi-spherical vesicles above and below (Fig. 4(d)).
Upon further stretch and increase of spontaneous curva-
ture the necks become vanishingly small (Fig. 4(e)), but
which are free of external forces, similar to the remote
constriction of a vesicle [25].
The conclusions and discussion are presented in Sec. VIII.
The parametrization and geometric quantities of an ax-
isymmetric surface are reviewed in Appendix A. The
derivations of the first integral of the EL equation and
(a) r0 = 1.1 (b) r0 = 1.2 of the external force adapted to axisymmetric surfaces
are presented in Appendix B.

II. BENDING ENERGY AND

(c) r0 = 1.3 (d) r0 = 1.4 EULER-LAGRANGE EQUATION

The membrane is represented by its midsurface,

parametrized by coordinates ua , a = 1, 2, and embedded
in three-dimensional space by the mapping Σ : (u1 , u2 ) 7→
X(u1 , u2 ). The two tangent vectors adapted to this
(e) r0 = 1.5 (f) r0 = 1.63
parametrization are ea = ∂a X; n is the unit outward nor-
mal; gab = ea · eb is the metric tensor and g ab its inverse;
Kab = ea · ∂b n is the extrinsic curvature tensor. The
two invariants of the shape operator K ab = g ac Kcb are
FIG. 3. (a)-(f) Sequence of stretched vesicles with fixed vol- (twice) the mean curvature, K = trK ab = C1 + C2 and
ume and Cs = 0. (f) At the maximum stretching, the vesicle the Gaussian curvature, KG = detK ab = C1 C2 , where C1
adopts a discocyte shape. The bending energy density is color and C2 are the two principal curvatures [26]. The bound-
coded in these figures. ary of Σ, denoted by ∂Σ, is parametrized by its arc length
s, so its embedding functions are given by the composi-
tion of maps ∂Σ : s 7→ X0 (U a (s)), where U a are the
and more (Fig. 2(c)) until it consists of two flat discs surface coordinates along the boundary. The Darboux
joined along their perimeter (Fig. 2(d)). The force in- frame adapted to ∂Σ is given by the right handed basis
creases monotonically as the membrane is stretched and defined by the unit tangent vector T := dX0 /ds = Ta ea ,
diverges in the limit of the disc-like shape. If the volume where Ta = dU a /ds, the surface unit normal n and the
is fixed, the poles of the membrane get closer and closer unit outward conormal L := T × n = La ea .
(Figs. 3(c)-(e)) until they touch, whereas the exterior re- On mesoscopic scales, the free energy of a fluid mem-
mains round, so the membrane adopts a shape similar to brane is given by the Canham-Helfrich energy, given by
a discocyte (Fig. 3(f)), for which the force is finite. For the sum of the bending energy, quadratic in the mean
 4

multipliers σ and P. The former contributes to the tan-

gential stresses and the latter represents the difference be-
tween the internal and external pressure, P = Pint −Pext .
The ring will be represented by a space curve Y(s), which
is also parametrized by the boundary arc length s. The
constraint Y(s) = X0 (U a (s)), is implemented by intro-
ducing a vectorial Lagrange multiplier λ [12, 32]. Thus
the constrained functional to be minimized is

H = HCH [X] + σ(A[X] − A0 ) − P(V [X] − V0 )

Z
+ dsλ(s) · [X0 (U a (s)) − Y(s)] , (2)

(a) r0 = 0.313 (b) r0 = 0.325 (c) r0 = 0.5 where ds is the line element of the ring. The Lagrange
multiplier λ(s) quantifies the change of the bending en-
ergy under a deformation of the ring
δHB δHB
λ=− = , (3)
δX0 δY
so it can be identified as the linear force density exerted
by the ring on the membrane boundary.
From the variation of the coordinates U a we get that
λ · ea = 0, thus, the force density along the boundary is
normal to the membrane, [12]

λ = λn n . (4)
(d) r0 = 0.78 (e) r0 = 0.85 The total variation of the constrained functional, given
in Eq. (2), has two terms representing the changes in the
energies of the bulk and of the boundary [16, 17, 20]

δH = δHΣ + δH∂Σ , (5a)

FIG. 4. (a)-(d) Sequence of vesicles
√ corresponding to a bifur- Z
cation with fixed area and Cs = 2 2/RS and (e) Cs = 3/RS . δHΣ = dA (∇a f a − Pn) · δX , (5b)
The bending energy density is color coded in these figures. ZΣ

δH∂Σ = ds (La δQa − λ · δY) . (5c)

∂Σ
curvature, and the Gaussian energy [27–29]
In the bulk term δHΣ , ∇a is the covariant derivative com-
HCH [X] = HB [X] + HG [X] , (1a) patible with gab ; f a is the stress tensor, which expressed
in the adapted basis reads
Z
kB 2
HB [X] := dA KD , (1b)
2
Z f a = f ab eb + f a n , (6a)
HG [X] := kG dA KG , (1c)
 
1
f ab = kB KD K ab − KD g ab − σg ab , (6b)
2
where we have defined KD = K − Cs , the difference be- f a = −kB ∇a K . (6c)
tween the mean curvature and the spontaneous curvature
Cs ; dA is the area element; kB and kG are the bending The normal component f a vanishes for constant mean
and Gaussian moduli with units of energy [19, 30, 31]. An curvature surfaces, and in particular for minimal surfaces
important consequence of the fact that the energy HCH with vanishing mean curvature.
depends only on the geometrical degrees of freedom of In the boundary term δH∂Σ , La are the covariant com-
the membrane is that the stresses are also determined ponents of the outward conormal along the equator, L,
completely by the membrane geometry. and we have defined
The constraints of fixed area and volume, which are of
global character, as well as the restriction on the sur- δQa = −f a · δX0 + H ab eb · δn , (7a)
face to conform to the ring, which is enforced locally, are H ab = kB KD g ab + kG (Kg ab − K ab ) . (7b)
introduced explicitly in the variational principle. Fixing
the area A and volume V of the membrane to given values In equilibrium, the stresses within the membrane bulk,
A0 and V0 involves the introduction of constant Lagrange given by the divergence of the surface stress tensor, are
 5

balanced with the pressure difference, which acts in the where τg = Ta Lb Kab is the geodesic torsion; κn =
normal direction1 Ta Tb Kab and κn⊥ = La Lb Kab are the normal curvatures
in the directions along the boundary and orthogonal to
∇a f a = Pn . (8) it [26]; ∇⊥ = La ∇a is the projected covariant derivative
along L. The vector f⊥ is the force per unit length ex-
If P = 0, as in the case of open membranes, f a is con- erted by a region of the membrane on its neighbouring
served. Equation (8) captures the fluid character of the region separated by a line element ds [20]. In Sect. IV A,
bilayer lipid membranes: since ∇a f a · eb = 0, there is we specialize this equation to axisymmetric surfaces in
no cost associated with deformations along its tangen- order to analyze the external normal force deforming the
tial plane, only normal deformations are penalized. The equatorial radius of the membrane.
Euler-Lagrange (EL) equation is given by the normal pro-
jection of the divergence of the stress tensor, ∇a f a ·n = P,
which in full reads [16, 17, 20] III. AXIAL SYMMETRY AND FIRST
  INTEGRAL OF THE EL EQUATION
K
(−∆ + σ̄)K + KD 2KG − (K + Cs ) = P̄ , (9)
2
We assume that the membrane remains axially sym-
metric under a radial deformation along one of its paral-
where ∆ = g ab ∇a ∇b is the Laplace-Beltrami operator on
lels caused by a circular ring of radius R0 . Moreover, for
the surface and from now on parameters with an over-
simplicity we consider that the ring is located at equa-
bar will be scaled with the inverse of kB : σ̄ = σ/kB and
tor, so the deformed membrane also has mirror symmetry
P̄ = P/kB , which have dimensions of inverse area and
with respect to the equatorial plane (Fig. 5).
inverse volume respectively. These two unknown param-
eters are not independent, as shown below in Sec. IV B.
There is a scaling relation connecting them, so even if
both of them are non vanishing, there is actually only
one independent parameter, which is determined by im-
posing the area or volume constraint.
The Gauss-Bonnet theorem implies that the Gaussian en-
ergy is given by the sum of a topological invariant and the
line integrals of the geodesic curvature along the bound-
aries [26]. In consequence, neither the bulk membrane
stresses nor the EL equation depend on the Gaussian
rigidity kG , it only enters the change in the boundary en-
ergy. Stationarity of the energy also entails the vanishing
of δH∂Σ , given in Eq. (5), which by expressing the surface
tangent vectors ea in the tangent basis adapted to the
boundary ∂Σ and taking into account that δY = δX0 ,
can be recast as
Z
− ds (λ + f⊥ ) · δX0
Z
+ ds (kG τg T + (kB KD + kG κn ) L) · δn = 0 , (10)

where we have defined the projection of the stress tensor

along the outward conormal by FIG. 5. Axisymmetric membrane constrained by a ring at
the equator. The meridian is parametrized by its arclength l,
f⊥ := La f a = f⊥⊥ L + f⊥∥ T + f⊥ n , (11a) and Θ is the angle that the tangent vector el makes with the
ab radial direction ρ̂. The integration contour Γ on the cap S of
f⊥∥ := La Tb f = kB KD τg , (11b) the surface is shown with a dashed line.
kB
f⊥⊥ := La Lb f ab = KD (κn⊥ − κn + Cs ) − σ, (11c)
2 Therefore, the membrane can be parametrized by arc-
f⊥ := La f a = −kB ∇⊥ K , (11d) length l along the meridian (measured from the ring)
and the azimuthal angle φ. The arc-length l is measured
from the the ring, so l = 0 at the equator and at the pole
it will be denoted by lp . In these coordinates the line
1 In the presence of an external normal force per unit area across element assumes the form ds2 = dl2 + R(l)2 dφ2 , where
the surface, Φ(ua ), the right hand side of Eq. (8) is replaced by R is the radius of the parallel circles. As discussed in
(Φ + P)n. the Appendix A, the derivatives of the radial and height
 6

coordinates of the meridian can also be described using have n · ẑ = − cos Θ = −R′ . Thus, integrating the right
the angle Θ that its tangent vector makes with the radial hand side of Eq. (14) from a parallel with arc-length l to
direction ρ̂ (Fig. 5), the pole where l = lp and R(lp ) = 0, we get
Z lp Z R
R′ (l) = cos Θ , Z ′ (l) = sin Θ , R2
Z
(12) 1 ′
dA n · ẑ = − dlRR = dRR = . (18)
2π l 0 2
where the prime stands for differentiation with respect to Σ
l, ′ = d/dl. The principal curvatures of an axisymmetric
Substituting Eqs. (17) and (18) in Eq. (14) we obtain the
surface occur along the meridians and parallels, which in
first integral of the EL equation
terms of Θ are given by
P
sin Θ cos Θ f⊥ + sin Θ f⊥⊥ + R = 0, (19)
C⊥ = Θ′ , C∥ = . (13) 2
R
which estates the balance of normal and tangential
Thus, the mean and Gaussian curvatures are given by stresses with the pressure difference. By substituting
K = C⊥ + C∥ and KG = C⊥ C∥ . Eqs. (15), in full this equation reads
The rotational symmetry of the membrane geometry im-   
plies the existence of a first integral of the EL equation

′
 1
2 
cos Θ C⊥ + C∥′ + sin Θ C⊥2
− C∥ − Cs − σ̄
(9) [16]. Integrating Eq. (8) within the cap S bounded 2
by parallel Γ of radius R above the equator (see Fig. 1), P̄
and using the divergence theorem to recast the left hand + R = 0, (20)
2
side as a closed line integral along Γ, we get
I Z which is to be solved along with Eqs. (12). This axially
ds f⊥ = P dA n , (14) symmetric shape equation is a second order differential
equation for Θ. In practice, numerical precision and sta-
Γ Σ
bility are improved by casting it in the more familiar
where ds = Rdφ is the arc-length. Thus, the total force equivalent Hamiltonian form (as described in Appendix
exerted on S, given by the line integral of the projection B) in terms of Θ and its conjugate momenta PΘ = RKD
of stress tensor onto the unit outward conormal along Γ, (which replaces the curvature C⊥ = Θ′ ). Alternatively,
is proportional to the integrated normal vector over the instead of working with PΘ , it proves useful to work with
same region. The equator is not included in the region of KD as a dependent variable along with R and Θ, so we
integration, so Eq. (14) does not capture the force exerted solve the system of first order differential equations given
by the ring, which will be calculated in Sec. IV A. by Eqs. (12) and
In this case the normal curvatures along the parallel and Θ′ − KD + C∥ − CS = 0 , (21a)
orthogonal to it are κn = C∥ and κn⊥ = C⊥ , whereas    
τg = 0, (see Appendix A), so the projected stress tensor ′ KD
cos Θ KD + sin Θ KD − C∥ + Cs − σ̄
and its tangential and normal components, given in Eq. 2
(11), simplify to P̄
+ R = 0. (21b)
f⊥ = f⊥⊥ L + f⊥ n , (15a) 2
kB
To solve this system of four differential equations in or-
f⊥⊥ = KD C⊥ − C∥ + Cs − σ , (15b) der to determine the shape of the deformed equilibrium
2   states, it is required to identify the four boundary condi-
f⊥ = kB K ′ = kB C⊥
′
+ C∥′ , (15c) tions on the geometry along the equator that are consis-
tent with the constraint imposed by the ring.
where the derivatives of the curvatures are

′ cos Θ
C⊥ = Θ′′ , C∥′ = (C⊥ − C∥ ) . (16) A. Boundary Conditions
R
The only non-trivial projection of Eq. (14) is along the We consider deformed vesicles with mirror symmetry
axis of symmetry, ẑ, and it is independent of s. Using the with respect to the equatorial plane, so the system of
result (A12b) of Appendix A, we find that the vertical equations (12) and (21) are solved for one half of the
projection of the left hand side of Eq. (14) reads generating curve with total arc-length lp . Setting the
I equator at the plane Z = 0, the boundary conditions of
ds f⊥ · ẑ = −2πR (cos Θ f⊥ + sin Θ f⊥⊥ ) . (17) the radial and height coordinates are given by
Γ R0 := R(0) = Rring , Z0 := Z(0) = 0 . (22)
Using Eq. (A6) of Appendix A for the normal vector to Finite bending energy demands the surface to be con-
an axisymmetric surface along with Eq. 12 for R′ , we tinuous and smooth across the equator and at the poles,
 7

otherwise the curvatures (and the energy) would diverge. A. Equatorial force as a balance of stresses
The existence of tangent planes along the equator and at
the poles impose the boundary conditions (BCs) on the We can consider the vesicle as composed by two halves
angle Θ, bounded by the equator, where they are in contact with
π the ring. Taking into account the contributions of each
Θ0 := Θ(0) = , Θp := Θ(lp ) = π . (23) halve, the upper one denoted by a + and the bottom one
2 with a −, we have from Eq. (5)(c) that the change of the
On account of Eqs. (12), these BCs also imply that the energy along the equator is given by
equator and the poles correspond to extrema of the ra- Z
δH∂Σ = ds La+ δQa+ + La− δQa− − λ · δY . (26)


dial and the height coordinates respectively. It should be
noted that, since Eq. (21b) does not hold at the equator,
where the force is applied by the ring, it is not legitimate Stationarity of the energy implies the vanishing of δH∂Σ .
to evaluate the first integral at Θ0 = π/2. Doing so, the Taking into account that the deformations of the embed-
coefficient of the first term vanishes and it reduces to an ding functions and the normal δY and δn are the same
equation relating the Lagrange multiplier σ and P, which for both boundary parallels, and that the equator is also
in general is inconsistent with the constraints of fixed area a principal curve, so τg = 0, we have in this case Eq. (10)
and volume or with the condition of smoothness at the is given by
pole, so in practice we evaluate at a very close value, Z Z
such as Θ0 = (1 ± 10−6 )π/2. The second BC replaces the dsλ · δY = − ds (f⊥+ + f⊥− ) · δY
specification of KD (0), which involves Θ′ (0), value that Z
is not fixed by the requisite of continuity along the equa- +

ds kB KD+ + kG C∥+ L+


tor. Furthermore, since lp is not known a priori, this is
a free boundary problem. The shooting method is used


+ kB KD− + kG C∥− L− · δn . (27)
to solve this boundary value problem: the initial value
KD (0) (or Θ′ (0)) is fine-tuned so that the second BC in The avoidance of kinks at the equator implies that the an-
Eq. (23) is fulfilled and the geometry closes smoothly at gle Θ ought to be continuous and smooth, rendering the
the pole. parallel curvature C∥ to be also continuous and smooth
The poles with Θ = 0, π and R = 0 are regular singular across the equator. However its counterpart along the
points where the curvature C∥ seems to be undetermined. meridian, C⊥ = Θ′ also should be continuous, but not
However, taking the limit l → lp and using the expres- necessarily smooth, for the presence of external forces will
sion of R′ , given in Eq. (12), we find that the poles are be reflected in its derivatives as explained below. Thus,
umbilical points, i. e. both curvatures are equal, [33] C∥+ = C∥− and C⊥+ = C⊥− . Furthermore, the tangent
vectors along the two boundaries have opposite directions
cos ΘΘ′ T+ = −T− = T = φ̂, and since the normal is the same
C∥p := lim C∥ = = C⊥p . (24) along both of them, the conormals also have opposite
l→lp R′ lp
directions L+ = −L− = L = −el . Taking this into ac-
Thus, at the poles we have Kp = 2C⊥p = 2Θ′p . count, we have that the vector multiplying δn vanishes,2
From Eq. (16), we have that C∥ ∝ (C⊥ − C∥ )/R, so on so the Gaussian rigidity does not enter. Moreover, since
account of Eq. (24) and also that R(lp ) = 0, apparently δY is arbitrary, we have that the stress transmitted to
also this derivative is undefined at the poles, but by tak- the membrane boundary is
ing the limit l → lp , we get
λ = −f⊥+ − f⊥− . (28)
′ ′
!
C⊥ − C∥ Thus, λ is identified as the external force exerted by the
lim C∥′ = cos Θ ′
= (C⊥ − C∥′ ) . (25) ring on the membrane boundary (recall f⊥ is the force per
l→lp R′ lp
lp unit length exerted by the membrane). Decomposing the
′ ′
force vector in the basis adapted to the boundary
Therefore at the poles we have C⊥p = 2C∥p . Further-
more, from Eq. (20) we have that at the poles Kp′ = f⊥± = f⊥∥± T± + f⊥⊥± L± + f⊥± n . (29)
′ ′
C⊥p + C∥p = 0, so the derivatives of the principal curva-
′ ′
Since the parallels are principal lines of curvature, the
tures vanish at the poles, C∥p = 0 and C⊥p = Θ′′p = 0. tangential projections vanish f⊥∥± = 0. The projection
of Eq. (28) onto the conormal reads

IV. FORCE EXERTED BY THE RING λL := λ · L = −f⊥⊥+ + f⊥⊥− . (30)

In this section we determine the external force in two

2
ways, using the balance of forces established at the equa- Besides, we are are interested in the particular case of equatorial
tor and by means of the scale invariance of the energy. deformation in which the normal does not change, so δn = 0.
 8

Since the curvatures are continuous, we have f⊥⊥+ = If Cs = 0 the two Lagrange multipliers are related,
f⊥⊥− and in consequence the tangential projection van- 2σA = 3PV .
ishes, so λL = 0, consistent with Eq. (4). The projection Let us apply the scaling argument for a surface con-
onto the normal reads strained by a ring to see how this relation generalizes.
Let us now consider the effect of a membrane rescaling
λn := λ · n = −f⊥+ − f⊥− . (31) X → ΛX on the constrained energy H, given by Eq.
(2). Under a rescaling the line element of the boundary
From Eq. (11d) we have that the normal components of changes as s → Λs, whereas the area, volume and mean
the stress tensor on each side are given by the derivatives curvature of the membrane scale as A → Λ2 A, V → Λ3 V
of the mean curvature along the conormal (L+ = −1 and K → Λ−1 K, respectively, so the energy changes as
′ ′
and L− = 1), so f⊥+ = kB K+ and f⊥− = −kB K− . Z
Substituting in Eq. (31) it reads kB
H[ΛX] = dA(K − ΛCs )2
2
′ ′
λ̄n = −K+ + K− , (32) +σ (Λ2 A − A0 ) − P (Λ3 V − V0 )
Z
where K ′ = C⊥ ′
+ C∥′ , and C⊥
′
and C∥′ were defined in + dsΛ λ(s) · (ΛX0 (s) − Y(s)) . (38)
Eq. (16). Due to the continuity of Θ and Θ′ , we have
′ ′
C∥+ = C∥− , so Eq. (32) reduces to While the bending energy itself is scale invariant, the
spontaneous curvature breaks this invariance. Stationar-
′ ′
λ̄n = −C⊥+ + C⊥− = −Θ′′+ + Θ′′− . (33) ity of the energy under a rescaling imposes the condition
dH
Thus, the linear force density on the equator involves = 0, (39)
only the difference of the derivatives of the meridian cur- dΛ Λ=1

vatures. This contrasts with the case of lipid membrane which implies that
domains where the force is due to the line tension and is Z
given in terms of the difference of the curvatures on both ds λ · X0 = −(2σ + kB Cs2 )A0 + 3PV0 + kB Cs M , (40)
sides, rather than of their derivatives, [34]. In particular,
if the vesicle has mirror symmetry with respect to the where we have used that A = A0 and V = V0 . This
equatorial plane, Θ(−l) = π − Θ(l), so the derivative of quantity is proportional to the change of the energy of
Θ is an odd function of arc length, Θ′′+ = −Θ′′− = Θ′′0 ,
R
the membrane boundary, δH∂Σ = dsλ · δX0 , under an
and the linear force density is proportional to the second infinitesimal dilation, δX0 = δΛX0 . Since the change of
derivative of Θ at the equator the energy of the membrane boundary is the negative of
the change of the energy of the ring, δHring = −δHb ,
λ̄n = −2Θ′′0 . (34) we get that under an infinitesimal dilation the change in
the energy of the ring is proportional to minus the the
The total normal force on the equator is obtained by right hand side of Eq. (40). In particular, for a circular
integrating this linear force density ring acting on an axially symmetric vesicle, we have on
Z account of the rotational symmetry Y(s) = R0 ρ̂ and
F̄ := dsλ̄n = −4πR0 Θ′′0 . (35) λ = λn ρ̂, so the magnitude of the total force exerted by
the ring is
These expressions for the linear force density and the to- 1
(2σ + kB Cs2 )A0 − 3PV0 − kB Cs M .


tal force can be derived also by integrating the EL equa- F = (41)
R0
tion in a region centered at the equator [20].
If Cs = 0 the constraining force is proportional to σ for
fixed area (P = 0), F̄ = 2σA0 /R0 , whereas it is pro-
B. Equatorial force from the scale invariance of the portional to P for fixed volume (σ = 0) F̄ = −3PV0 /R0 ,
bending energy which is less than evident at the level of the EL equation.
In each case, in the limit of maximum constriction, with
In the absence of external forces, the scale invariance R0 → 0, the force is proportional to the derivatives of
of bending energy, Eq. (1), provides a useful relation be- σ and P with respect to R0 , F̄ = 2A0 ∂σ/∂R0 for fixed
tween the spontaneous curvature Cs , the Lagrange multi- area and F̄ = −3V0 ∂P/∂R0 for fixed volume. The in-
pliers σ and P and the global quantities of the membrane troduction of a new scale associated with the constraint
on the ring is consistent with a non-vanishing value for
(Cs2 + 2σ̄)A − 3P̄V − Cs M = 0 , (36) σ or P. Only in the limit R0 → 0, the scale invariance is
restored.
where we have defined the total mean curvature Combining Eqs. (35) and (41) we can express the second
Z derivative of the angle Θ as
M = dAK . (37) 4πR02 Θ′′0 = −(2σ̄ + Cs2 )A0 + 3P̄V0 + Cs M . (42)
 9

Since this equation involves the total mean curvature M , The bending energy of a spherical vesicle with sponta-
which is a global quantity, it is not useful in the deter- neous curvature is HBS (Cs ) = 2πkB (2 − cs )2 . We rescale
mination of σ, P or Θ′′0 . However, as shown below it the total bending energy of the vesicles with the energy
is useful in the limit of maximum constriction, where it of a spherical vesicle with null spontaneous curvature,
permits us to determine σ if the area is fixed (P = 0) or HBS (0) = 8πkB . Thus, for a spherical vesicle we have
to determine P if the volume is fixed (σ = 0).
HB  cs 2
hB := = 1− (49)
HBS (0) 2
V. NONDIMENSIONAL QUANTITIES
which vanishes if cs = 2. The scaled total enegy of sym-
The relevant length scale is set by the radius of the metric membranes is given by
spherical vesicle. In the following we consider nondimen- Z ℓp
1
sional quantities obtained by a rescaling with appropriate hB = dℓκ2 r . (50)
powers of RS . The scaled arc-length along the meridian, 4 0
measured from the equator is
Since σ̄ and P̄ have dimensions of inverse area and in-
l verse volume, the nondimensional intrinsic tension and
ℓ := . (43) nondimensional pressure are defined by
RS
The scaled arclength at the poles is ℓp = lp /RS . The µ := RS2 σ̄ , p := RS3 P̄ . (51)
derivative with respect to the nondimensional arc-length
will be denoted by a dot, ˙ = d/dℓ = RS d/dl. The reduced We define the scaled linear force density and the scaled
coordinates are defined by total force on the equator (given by Eqs. (41))
R Z
r(ℓ) := , z(ℓ) := . (44) ϕ0 := RS2 λ̄n = −2Θ̈0 , (52a)
RS RS
f0 := RS F̄ = −4πr0 Θ̈0
Thus, the nondimensional embedding functions are 4π
(c2s + 2µ)a0 − p v0 − 2cs m .


x(ℓ) := X/RS = r(ℓ)ρ̂ + z(ℓ)ẑ. The radius of the ring = (52b)
r0
located at the equator with ℓ = 0 is r0 := R0 /Rs .
The principal curvatures and the spontaneous curvature So the analogue of Eq. (42) reads
are scaled with RS ,
r02 Θ̈0 = − 2µ + c2s a0 + p v0 + 2cs m .


(53)
c⊥ := RS C⊥ = Θ̇ , (45a)
sin Θ Expressing Eqs. (12) and (21) in terms of these nondi-
c∥ := RS C∥ = , (45b)
r mensional coordinates and parameters, we have (˙ :=
cs := RS Cs . (45c) d/dℓ)
The nondimensional mean curvature and the mean cur- ṙ = cos Θ , (54a)
vature difference are ż = sin Θ , (54b)
k := RS K = c⊥ + c∥ , (46a) sin Θ
Θ̇ = κ − + cs , (54c)
κ := RS KD = k − cs . (46b) r   
κ sin Θ
We define the reduced area, volume and total mean cur- cos Θ κ̇ + sin Θ κ − + cs − µ
2 r
vature by p
+ r = 0. (54d)
A V M 2
a := , v := , m := , (47)
AS VS MS The second derivative of Θ is given by
where AS = 4πRS2 , VS = 4πRS3 /3 and MS = 8πRS are  
the area, volume and total mean curvature of a sphere of cos Θ 2 sin Θ
Θ̈ = κ̇ − κ− + cs . (55)
radius RS . For surfaces with equatorial mirror symmetry r r
we have
Z ℓp The initial values of the reduced mean curvature differ-
ence and its derivative can expressed in terms of the ini-
a = dℓr , (48a) tial values of the angle and its derivatives as
0
Z ℓp
3 sin Θ0
v = dℓ sin Θ r2 , (48b) κ0 = Θ̇0 + − cs , (56a)
2 0 r0
ℓp  
cos Θ0 sin Θ0
Z
1
m = dℓ k r . (48c) κ̇0 = Θ̈0 + Θ̇0 − . (56b)
2 0 r0 r0
 10

√
The reduced coordinates, angle and mean curvature dif- cs 2.75 2 2 3
ference of a sphere of radius Rs , whose equator is at a a
r01 0.44 0.49 0.56
height Ze and with arc length le (so the reduced height a
and arc length of the equator are ze = Ze /RS and r02 1.14 1.15 1.17
a
ℓe = le /RS ), are given by r03 0.35 0.31 0.28
a
r04 0.74 0.78 0.85
r(ℓ) = rs cos θ , z(ℓ) = rs sin θ + ze , (57a)
π 2 TABLE I. Numerical values of r0 for vesicles with a = 1
Θ(ℓ) = θ + , κ(ℓ) = − cs , (57b) belonging to the bifurcations.
2 rs
where we have defined the reduced radius and the angle
θ by radius r0 , which are illustrated in Fig. 6 for constriction
and in Fig. 7 for dilation. However, for higher values of
Rs ℓ − ℓe
rs = , θ := . (58) cs two bifurcations in the solutions arise, which comprise
RS rs constricted and dilated vesicles, shown in Figs. 8 - 10.
The sphere is a solution of the first integral (54d) if the The equatorial radius of a constricted vesicle is in the in-
parameters satisfy the equation [35] terval 0 < r0 < 1. As r0 is decreased, the vesicle adopts a
prolate shape (Fig. 6(a)) and elongates along the symme-
p  c
s

try axis. Then the equatorial region becomes cylindrical
rs µ − rs2 + cs rs − 1 = 0 . (59)
2 2 (Θ̇0 = 0), setting the beginning of the transition from
a prolate shape (Fig. 6(b)) to a dumbbell shape with a
This agrees with Eq. (53) for a0 = 1, v0 = 1, m = 1, and waistline, (Fig 6(c)). As r0 is further decreased the waist
Θ̈0 = 0, so the sphere is a solution free of external forces. decreases rapidly and the vesicle is mainly composed of
If cs = 0, 2/rs the Lagrange multipliers enforcing the area two increasing quasi spherical vesicles (Fig. 6(d)). In the
and volume are proportional, prs = 2µ. For the initial limit of maximum constriction, r0 → 0, the vesicle con-
spherical configuration Rs = RS , so the scaled radius is √
sists of two√quasi-spherical√ vesicles of radius ra = 1/ 2
unit, rs = 1, and we set the equator on the plane Z = 0, (so ℓp = π/ 2 and ka = 2 2) connected by a vanishingly
so ℓe = 0 and ze = 0. small catenoid-like neck (Figs. 6(e)-(f)).
The equatorial radius √ of a stretched vesicle is in the
interval 1 < r0 < 2. As r0 is increased the vesi-
VI. EQUILIBRIUM SHAPES WITH FIXED
AREA cle becomes oblate and the distance between the poles
decreases (Figs. 7(a)-(c)), then the polar regions flatten
more and more, (Fig. √ 7(d)), up to the limit of maximum
We first consider deformed vesicles whose area is equal
stretching with r0 → 2, where the vesicle is composed
to the area of the spherical vesicle, A = AS , so their re-
of two √discs joined along their boundary, (Fig. 7(e)), so
duced area is unit, a = 1. The reduced volume of the
ℓp → 2. As the spontaneous curvature is increased
vesicle v is variable, so p = 0. This case could repre-
above cs = 2.7 the sequence of configurations is not
sent the situation in which the control parameter is the
simple, for two bifurcations occurs in the solutions of
pressure difference across the membrane and with a fixed
the system of differential equations. Some relevant val-
temperature, so that the enclosed volume varies. In this a
ues pertaining to these bifurcations, referred to as r0i ,
case Eq. (54d) reduces to
i = 1, 2, 3, 4 in the following discussion, are presented
 
κ sin Θ
  in Table I. In the sequence of constricted vesicles, as
cos Θκ̇ + sin Θ κ − + cs − µ = 0 . (60) r0 is reduced there are no further configurations below
2 r a
r0 = r01 . However, the sequence of the configurations
Evaluating Eq. (60) at a parallel with Θ0 and r0 , and continues for higher values of r0 along a bifurcation in
a a a
using Eqs. (56), we can determine µ in terms of the initial the interval r01 < r0 < r02 , illustrated in Fig. 8. At r01
values of the angle and its derivatives the vesicle is in the transition from prolate to dumbbell
   shapes (Fig. 8(a)). As r0 is increased the equatorial re-
cos Θ0 sin Θ0 gion begins to bulge (Figs. 8(b)-(c)), and as r0 is further
µ = cot Θ0 Θ̈0 + Θ̇0 − increased it swells while the regions close to the poles be-
r0 r0
a
 2 ! come rounded and develop necks (Figs. 8(d)-(e)). At r02
1 sin Θ0 the central region reaches a maximum stretching (Fig.
+ Θ̇20 − − cs . (61)
2 r0 8(f)), beyond which there are no further solutions with
higher r0 . Afterwards, the sequence of configurations
We solved the system of differential equations (54) for dif- continues for lower values of r0 along another bifurcation
a a
ferent values of cs . Starting with values close to cs = 0 in the interval r03 < r0 < r02 , illustrated in Fig. 9. As r0
a
and up to cs = 2.7 we found that there is a single se- is reduced below r02 the central region deflates and the
quence of configurations characterized by the equatorial necks shrink, while the top and bottom round regions
 11

(a) r0 = 0.95 (b) r0 = 0.845 (c) r0 = 0.5 (d) r0 = 10−1 (e) r0 = 10−2 (f) r0 = 10−3

FIG. 6. (a)-(e) Sequence of constricted vesicles with a = 1 and cs = 0 as the radius of the ring is reduced. (e) For a very
small radius, the vesicle is constituted by two quasi-spherical vesicles connected by a small catenoid-like neck. (f) Magnification
of the profile curve of the small neck (the solid line corresponds to the numerical solution and the black dashed line to the
catenoid). The scaled bending energy density is color coded in these figures.

(a) r0 = 1.05 (b) r0 = 1.1 (c) r0 = 1.2 (d) r0 = 1.3

(e) r0 = 1.375

FIG. 7. (a)-(e) Sequence of stretched vesicles with a = 1 and cs = 0 as the radius of the ring is increased. The scaled bending
energy density is color coded in these figures.

become quasi-spherical, Figs. 9(a)-(b). Then, the cen- rounded (Fig. 9(d)), until the vesicle adopts a dumbbell
tral region keeps deflating, but the radius of the necks shape Fig. 9(e), after which the constriction proceeds just
increases and the vesicle adopts an onduloid-like shape as in the case of the single sequence of configurations dis-
Fig. 9(c). Upon further reduction of r0 the necks con- cussed above.
flate with the central region while the extremes are kept The radius of the necks of the configurations of the second
 12

(a) r0 = 0.49 (b) r0 = 0.6 (c) r0 = 0.75 (d) r0 = 1 (e) r0 = 1.1 (f) r0 = 1.151

√ a a
FIG. 8. (a)-(f) Sequence of vesicles with a = 1 and cs = 2 2 belonging to the first bifurcation as r0 is increased (r01 < r0 < r02 ).
The scaled bending energy density is color coded in these figures.

√
bifurcation can be rendered very small by increasing cs , cs -1 0 1 2 2.7 2.75 2 2 3
for instance for cs = 3 in the interval 0.77 < r0 < 0.95 the
µamc -1.91 0 0.91 0.83 0.17 0.11 0 -0.26
configurations consist of oblate membranes connected to
quasispherical vesicles above and below by an infinitesi- haBmc 3.66 2 0.84 0.17 0.004 0.0015 0 0.0074
1 a
mal necks (see Fig. 10). This situation could be regarded f
4π 0mc
-3.83 -2.83 -1.83 -0.83 -0.13 -0.08 0 0.17
as the remote constriction of the vesicle, similar to the
process presented in Ref. [25]. The quasispherical vesi- TABLE II. Numerical values of quantities of vesicles with
cles satisfy Eq. (59) with p = 0, so they are not subject a = 1 and different values of cs in the limit of maximum
to a external forces and both of them have a radius given constriction.
by
2cs plotted in Fig 12. For the spherical membrane with r0 =
rs = . (62)
c2s + 2µ 1 it has a value µ = cs (1 − cs /2). For values close to cs =
0, as r0 is decreased µ first decreases reaching a global
The plots of the parameters of the vesicle are shown in minimum and then increases up to the value µamc (given
Figs. 11-15, where the values corresponding to these con- in Table II) in the limit of maximum constriction. For
figurations with small necks are shown with a dashed line. stretched membranes µ increases with r0 , diverging in√ the
The values of Θ̇0 and Θ̈0 as functions of r0 are plotted in limit of maximum stretching, i.e. µ → +∞ as r0 → 2.
Figs. 11. For the initial spherical state with r0 = 1 their For cs > 2.7, µ increases along the first bifurcation and
values are Θ̇0 = 1 and Θ̈0 = 0. For values close to cs = 0 decreases along the second one.
and constriction, Θ̇0 decreases (Θ̈0 increases) monotoni- The scaled total length of the profile curve 2ℓp and the
cally, tending to −∞ (+∞) as r0 → 0. As cs is increased reduced volume v are plotted in Fig. 13. The scaled to-
Θ̇0 (Θ̈0 ) takes lower (higher) values until
√ it develops a tal length begins at the value π and for values close to
maximum (minimum) and for cs > 2 2, Θ̈0 → −∞ as cs = 0 it increases as r0 is decreased, reaching √ a max-
r0 → 0, which has the consequence that the total force imum and then decreases reaching the value 2 in the
changes of sign. Above cs = 2.7, Θ̇0 (Θ̈0 ) increases (de- limit of maximum constriction. For√dilation it decreases
creases) along the first bifurcation, whereas along the sec- monotonically reaching the value 2 2 in the maximum
ond one it decreases (first increases and then decreases). stretching limit. For cs > 2.7, along both bifurcations
For dilation Θ̇0 > 1 (Θ̈0 < 0) and it increases (decreases)
√ 2ℓp first increases and then decreases.
monotonically, tending to +∞ (−∞) as r0 → 2. The behavior of the reduced volume is practically the
The dependence of the Lagrange multiplier µ on r0 is same for cs < 2.7, it begins with the value v = 1 at r0 = 1
 13

(a) r0 = 1 (b) r0 = 0.78 (c) r0 = 0.5 (d) r0 = 0.325 (e) r0 = 0.313

√ a
FIG. 9. (a)-(e) Sequence of vesicles with a = 1 and cs = 2 2 belonging to the second bifurcation as r0 is decreased (r03 < r0 <
a
r02 ). The scaled bending energy density is color coded in these figures.

and for constriction it decreases with r0 , after √

it reaches tend to decay to the configurations of the second bifur-
a minimum it increases up to the value v = 1/ 2 at the cation.
limit of maximum constriction. For dilation v increases The scaled total force f0 is plotted in Fig. 15. The data of
with r0 reaching a maximum and then it decreases to- the numerical solutions confirm that the equatorial force
ward 0 in the limit of maximum stretching. For cs > 2.7, is given by the derivative of the bending energy with re-
along the first bifurcation v increases reaching a maxi- spect to the equatorial radius, F = ∂HB /∂R0 or in terms
mum, then it decreases and along the second bifurcation of scaled quantities f0 = 8π∂hb /∂r0 . The initial spherical
it keeps decreasing, it reaches a minimum and then it membrane is free of external forces, f0 = 0. For values
increases. close to cs = 0 the total force on the constricted mem-
The scaled total energy hB is plotted in Fig. 14. The total branes is negative (so it is constrictive), and displays an
energy of the initial spherical configuration with r0 = 1 is oscillating behavior: as r0 is decreased f0 also decreases
given in Eq. (49). For values close to cs = 0, as the equa- to a minimum, then it increases to a maximum, after
tor is reduced, hB increases monotonically, reaching the which it decreases again towards the limit of maximum
value haBmc (given in Table II) for the totally constricted constriction, where it reaches a finite value f0amc (given in
membranes, which agrees with the exact value Table II). Let us examine the behavior of the total force in
√ the limit of maximum constriction a little more closely.
haBmc = ( 2 − cs /2)2 (63) Although the linear force density √ diverges as r0 → 0,
corresponding to the reduced total energy of two quasi- ϕ0 = 2Θ̈0 → ∞ (−∞ for cs > 2 2), their product r0 Θ̈0
spherical vesicles joined by an infinitesimal neck, which converges to the finite value given above, so f0 is finite.
√
vanishes if cs = 2 2. For stretched membranes As we will see below, this is possible because in this limit
√ hB in-
creases with r0 , diverging to +∞ as r0 → 2. For the geometry of the neck does not correspond to a sur-
cs > 2.7, hB changes twice in a non-smooth way, first at face of constant mean curvature, in particular it is not
the transition from the original sequence of constricted the central segment of a catenoid, which could not sus-
configurations to the first bifurcation, where it changes tain an equatorial force. For stretched vesicles the force
from decreasing to increasing and also at the transition increases monotonically as r0 is increased, diverging in
to the second bifurcation, where it becomes decreasing the limit of two flat
√ discs connected along their rim, i.e
again and intersects the original sequence at r0 = r04 a
, f0 → ∞ as r0 → 2. Since it is not possible to stretch
corresponding to two different configurations with the the membrane into such disc-like geometry by applying a
same total energy. For r0 < r04 a
the configurations of finite force, one would expect the membrane to rupture
the second bifurcation have the lowest energy, so it is beyond some critical force, which would occur through
conceivable that the original constricted configurations pore formation [36, 37]. As cs is increased f0 takes more
 14

(a) r0 = 0.77 (b) r0 = 0.85 (c) r0 = 0.95

FIG. 10. (a)-(c) Vesicles with a = 1 and cs = 3 belonging to the second bifurcation and with vanishingly small necks. The
scaled bending energy density is color coded in these figures.

positive values and for cs > 2 it becomes positive (so it is borhood of the neck we have ℓ ≪ 1 (l ≪ RS ), we consider
dilative) for some configurations. For cs > 2.7, f0 is pos- the rescaled arclength
itive and increasing along the first bifurcation, whereas
along the second one it decreases towards negative values, ℓ l
ℓ̃ = = . (64)
reaches a minimum and then increases to positive values r0 R0
where it merges with the original sequence of constricted
vesicles. Thus the rescaled arc length derivative is given by d/dℓ̃ =
r0 d/dℓ. Likewise, since r ≪ 1 (R ≪ RS ) and |z| ≪ 1
(Z ≪ RS ) we consider the rescaled coordinates
A. Limit of maximum constriction:
quasi-catenoidal neck
r R z Z
r̃(ℓ̃) = = , z̃(ℓ̃) = = . (65)
r0 R0 r0 R0
At first glance, from the numerical solutions it appears Besides, the magnitudes of the curvatures are large
that, regardless of the value of cs , the shape of the neck
is well approximated by the central section of a catenoid 1
c∥ = −c⊥ =   ≫ 1, (66)
(its generating curve, a catenary, is shown with a dashed r0 1 + ℓ̃2
line in Figs. 1(f) and 6(f)). However, such comparison
is deceptive, although the catenoid is a solution of Eq.
so we consider the rescaled curvatures c̃∥ = r0 c∥ and c̃⊥ =
(54d) with vanishing µ and p, as well as null sponta-
r0 c⊥ , as well as the rescaled mean curvature difference
neous curvature, since it is a minimal surface it cannot
κ̃ = r0 κ = c̃⊥ + c̃∥ − c̃s , where c̃s = r0 cs . We expand the
withstand a force, in stark contrast with the fact that a
coordinates employing r0 as the small parameter
finite force is required to hold the membrane at the max-
imum constriction. Furthermore, we see in Figs. 1(f) and r̃(ℓ̃) = r̃(0) (ℓ̃) + r̃(1) (ℓ̃) + . . . , (67a)
6(f) that the mean curvature of the neck varies along it,
so it cannot be part of constant mean curvature surface z̃(ℓ̃) = z̃(0) (ℓ̃) + z(1) (ℓ̃) + . . . , (67b)
(in particular of a minimal surface). To resolve this ap- Θ(ℓ̃) = Θ(0) (ℓ̃) + Θ(1) (ℓ̃) . . . , (67c)
parent contradiction we employ a perturbative analysis
about the catenoid. To this end, we consider a catenoidal κ̃(ℓ̃) = κ̃(0) (ℓ̃) + κ̃(1) (ℓ̃) + . . . , (67d)
neck whose radius R0 is much smaller than the radius of
the vesicle RS , so the scaled neck radius r0 = R0 /RS ≪ 1 where in the mean curvature difference κ̃(0) (ℓ̃) = c̃⊥(0) +
sets the length scale in this regime. Since in the neigh- c̃∥(0) and κ̃(1) (ℓ̃) = c̃⊥(1) + c̃∥(1) − c˜s . The deformation
 15

FIG. 11. Dependence of Θ̇0 and Θ̈0 on r0 for membranes with

a = 1. The dashed line represents configurations with very FIG. 12. Lagrange multiplier µ as a function of r0 for mem-
small necks. branes with a = 1. The insets show values close to the limit
of maximum constriction. The dashed line represents config-
urations with very small necks.
should keep the equator fixed, so the following BCs are
to be fulfilled by the first order perturbations
The curvatures are given by
r̃(1) (0) = 0 , z̃(1) (0) = 0 , Θ(1) (0) = 0 . (68)
sin Θ(0) 1
c̃∥(0) = = 2 , (71a)
The zeroth order terms are given by the functions corre- r̃(0) r̃(0)
sponding to the catenoid3
dΘ(0) 1
c̃⊥(0) = =− 2 . (71b)
r̃(0)
q
dℓ̃
r̃(0) (ℓ̃) = 1 + ℓ̃2 , (69a)
z̃(0) (ℓ̃) = arcsinh ℓ̃ , (69b) Thus, the scaled mean curvature k̃(0) = c̃∥(0) + c̃⊥(0) van-
Θ(0) (ℓ̃) = arccot ℓ̃ , (69c) ishes as it is a minimal surface. Moreover, the derivative
of the meridian curvature is proportional to the scaled
κ̃(0) (ℓ̃) = 0 . (69d) arc-length
The derivatives of the radial and height coordinates are dc̃⊥(0) d2 Θ(0) 2ℓ̃
= = 4 , (72)
dr̃(0) ℓ̃ dℓ̃ dℓ̃2 r̃(0)
= cos Θ(0) = , (70a)
dℓ̃ r̃(0)
so it vanishes at the equator where ℓ̃ = 0. In consequence
dz̃(0) 1
= sin Θ(0) = . (70b) the force vanishes as well
dℓ̃ r̃(0)
4π d2 Θ(0)
f0 = −4πr0 Θ̈(0) (0) = − (0) = 0 , (73)
r0 dℓ̃2
3 κ̃(0) = 0 because the mean curvature of the catenoid is zero and consistent with the fact that the catenoid is a minimal
it is only a solution of the EL for null spontaneous curvature. surface, which satisfies the source-free EL equation when
 16

FIG. 13. Total length of the generating curve 2ℓp and volume
v as functions of r0 for membranes with a = 1. The dashed FIG. 14. Scaled total energy hB as a function of r0 for mem-
lines represent configurations with very small necks. branes with a = 1. The dashed line represents configurations
with very small necks.

the parameters vanish, i.e.

µ(0) = 0 , p(0) = 0 , cs(0) = 0 . (74)

The system of differential equations, given in Eqs. (12)

and (21), read
Substituting Eqs. (67) we get that at first order the sys-
dr̃ tem of Eqs. (75) are given by (like for the sphere we
= cos Θ , (75a) consider the deformation of just one halve, say the one
dℓ̃
dz̃ with ℓ̃ > 0)
= sin Θ , (75b)
dℓ̃
dΘ sin Θ
= κ̃ − + c̃s , (75c)
dℓ̃ r̃
   
dκ̃ κ̃ sin Θ
cos Θ = − sin Θ κ̃ − + r0 cs − µ̃
dℓ̃ 2 r̃
dr̃(1) Θ(1)
p̃ + = 0, (77a)
− r̃ , (75d) dℓ̃ r̃(0)
2
dz̃(1) ℓ̃
where − Θ(1) = 0 , (77b)
dℓ̃ r̃(0)
µ̃ = r02 µ , p̃ = r03 p . (76) dΘ(1) ℓ̃ 1
+ 2 Θ(1) − r̃ 3 r̃(1) − κ̃(1) = 0 ,
r̃(0)
(77c)
dℓ̃ (0)
Since the scaled spontaneous curvature and the Lagrange
multipliers are proportional to powers of r0 , they only en- dκ̃(1) 1
ℓ̃ − 2 κ̃(1) = 0. (77d)
ter at orders of the expansion higher than one. dℓ̃ r̃(0)
 17

a
the mean curvature of the neck is given by kmc = k̃(1) /r0 ,
which reads

!
a
√ ℓ̃ ℓ̃
kmc =2 2 + cs 1− . (80)
r̃(0) r̃(0)

Therefore, the mean curvature of the neck starts at the

value of cs at the equator, feature known as kissing con-
dition [7, 21, 34], and far from it reaches the value√ ka .
Thus, up to first order, we see that only for cs = 2 2 the
neck is represented by a constant mean curvature sur-
face. The mean curvature of the neck of the numerical
solutions and this first order correction are plotted in Fig.
16, showing a good agreement.

FIG. 15. Scaled total force f0 as a function of r0 for mem-

branes with a = 1. The dashed line represents configurations
with very small necks.

The solution to Eq. (77d) is4

FIG. 16. Mean curvature as a function of the arc-length of
ℓ̃ configurations with a = 1 in the region close to the neck. At
κ̃(1) (ℓ̃) = κ̃(1)A , (78)
r̃(0) the equator its value is given by spontaneous curvature and
away from the equator its value tends to √the mean curvature
where κ̃(1)A is a constant of integration. This first order of the quasi-spherical membrane, ka = 2 2. The solid lines
correction to the mean curvature difference vanishes at correspond to the numerical data and the dashed lines to plots
the equator and tends asymptotically to κ̃(1)A for ℓ̃ ≫ 1. of the first order correction k̃(1) /r0 .
The asymptotic value κ̃(1)A ought to match the value of
the mean curvature difference of the touching spherical
√
vesicles (scaled by r0 ), κa = ka − cs , where ka = 2 2 is
the mean curvature of the quasi-spherical vesicles, so we Using Eq. (77a) in Eq. (77c) to replace Θ(1) in favor of
get r̃(1) , substituting Eq. (78) and integrating with the BCs
 √  (68) we determine Θ(1) in terms of r(1)
κ̃(1)A = r0 κa = r0 2 2 − cs . (79)

The first order correction to the mean curvature is k̃(1) = ℓ̃

κ̃(1) +c̃s . Since the lowest order mean curvature vanishes, Θ(1) = r̃(1) + κ̃(1)A (r̃(0) − 1) . (81)
r̃(0)

4 In general, the first order

√ correction to the mean curvature dif- Substituting this result in Eq. (77a) we obtain an ODE
ference is κ(1) = C(1) r2 − 1/r where C(1) is a constant [7, 21]. for r̃(1) , whose solution also determines Θ(1) on account
√
So by substituting the catenoidal radial coordinate r = 1 + ℓ2 of Eq. (81), and which in turn allows for the integration
we obtain the same result. of (77b) to determine z̃(1) . These solutions with the BCs
 18

(68) are given by tor (ℓ̃ = 0), the second order derivative is non-vanishing
  d2 Θ(1)  √ 
r̃(1) 1 1 Υ(ℓ̃) (0) = κ̃(1)A = r0 2 2 − cs . (84)
= − ℓ̃ − , (82a) dℓ̃2
κ̃(1)A r̃(0) 2 2r̃(0)
Since r02 Θ̈(1) = d2 Θ(1) /dℓ̃2 , we have
!
z̃(1) 1 ℓ̃Υ(ℓ̃)
= 1+ √
κ̃(1)A r̃(0) 2 2 2 − cs
Θ̈(1) (0) = . (85)
1 2  r0
+ ℓ̃ − Υ(ℓ̃)2 − 1 , (82b)
4 Thus, although the scaled equatorial force density ϕ0 =
Θ(1) 1 1   2Θ̈0 diverges as r0 → 0, the total scaled force converges
2 = r̃(0) + − 2 ℓ̃Υ(ℓ̃) + 2 , (82c) to a finite value
κ̃(1)A r̃(0) r̃(0)
! a
f0mc √
ℓ̃ = −r0 Θ̈0 = −(2 2 − cs ) . (86)
Υ(ℓ̃) := arctanh . (82d) 4π
r̃(0) This result agrees with the numerical results of the scaled
force, see Table II and the insets of Fig. (15).
√ In particu-
These first order deformations make the generating
curve of the neck slightly wider in comparison with the lar f0 is negative
√ (constrictive) for cs < 2 2, it vanishes
√
catenoid, as shown in Fig. 17. for cs = 2 2 and becomes positive for cs > 2 2, so
a stretching force is required to maintain the vesicle in
equilibrium.
The value of µ at the maximum constriction is deter-
mined from the scaling relation, Eq. (53), relating the
Θ̈0 and the global quantities of the surface, which in this
case reads
−r02 Θ̈0 = 2µ + cs (cs − 2m) . (87)
√
In the limit r0 → 0, r02 Θ̈0
→ 0 and m → ka aa = 2, so
we have that µ is quadratic in cs
√ cs 
µamc (cs ) = cs 2− . (88)
2
√
Thus, µamc is√positive for 0 < cs < 2 2 and it vanishes
for cs = 0, 2 2, which agrees with the numerical values
presented in Table II.

B. Equatorial stretching: disc limit

FIG. 17. Profile curve of the narrow neck, the solid line cor- In the limit of maximum stretching the curvature is
responds to the first order deformed catenoid and the black
concentrated near the equator, where the meridian cur-
dashed line to the original catenoid. The magnitude of the
perturbation has been augmented for illustration purposes.
vature is large, c⊥ ≫ 1. Since in this region r0 ≈ 1 and
The bending energy density is color coded in this figure. | sin Θ| ≤ 1, we have c∥ ≈ 1, so c⊥ ≫ c∥ . We also assume
that the spontaneous curvature is finite, c⊥ ≫ cs . Thus,
to analyze this limit of a disc-like surface, we can con-
The derivatives of Θ(1) are
sider as a small parameter the radius of curvature given
! ! by the inverse of the value of c⊥ at the equator,
1 dΘ(1) 1 1 ℓ̃ Υ(ℓ̃)
= − 2 + 2 1 1
κ̃(1)A dℓ̃ 2 r̃(0) r̃(0) r̃(0) ρD = = ≪ 1. (89)
c⊥0 Θ̇0
2ℓ̃
+ 4 , (83a) Taking into account these considerations, the first inte-
r̃(0) gral, Eq. (54d), simplifies to5
!
d2 Θ(1)
   2 
1 2 3 4 c
= 1− 3 − r̃ 5 cos Θċ⊥ + sin Θ ⊥ − µ = 0 . (90)
κ̃(1)A dℓ̃2 r̃(0) r̃(0) (0) 2
!
4 ℓ̃
+ 2 −1
r̃(0) 4 Υ(s)
r̃(0)
(83b) 5 Even if we considered the term proportional to p it would also be
small compared with the other quantities, because in this limit
Although the first order derivative vanishes at the equa- V → 0 so the pressure difference vanishes, P → 0.
 19

Evaluating at the equator we determine the constant 2 ρD

c2⊥0 1
µ= = 2 . (91)
2 2ρD
ρD
√ 0

z
Thus, µ diverges with 1/ρ2D in the limit r0 → 2. Inte-
grating Eq. (90) and imposing the asymptotic boundary
condition that far from the equator the surface becomes
planar, Θ → π and Θ̇ → 0, yields
√ -2 ρD
2 Θ
c⊥ = Θ̇ = cos . (92) r0
ρD 2
r
Integrating once more, with the boundary condition
Θ(0) = π/2, we get

Θ ℓ − ℓ0
sin = tanh Ξ(ℓ) , Ξ(ℓ) := √ , (93) FIG. 18. Profile curve of the disc limit near the equatorial
2 2ρD
region. The osculating circle of radius ρD is shown with a
√ √ dashed line.
where ℓ0 = − 2ρD arctanh(1/ 2). Since ρD ≪ 1, Θ ≈ π
away from the equator, so in this planar region c∥ ≈ 0.
From this result, we can express the first order differential glued together along their perimeters (see Fig. 7(e)). The
for the coordinates as profile of the equatorial region, which agrees very well
with the numerical solution, is plotted in Fig. (18).
ṙ = sech2 Ξ(ℓ) − tanh2 Ξ(ℓ) , (94a)
Differentiating Eq. (92) and using it to substitute Θ̇ in
ż = 2 sech Ξ(ℓ) tanh Ξ(ℓ) . (94b) favor of Θ, we obtain the second order derivative
Integrating these equations with the boundary conditions 1
Θ̈ = − sin Θ . (99)
r(0) = r0 and z(0) = 0, we get 2ρ2D
√ 
r = r0 − ℓ + 2ρD 2 tanh Ξ(ℓ) − 1 , (95a) Evaluating at the equator we have that √ the2 scaled force
is given by f0 /(4π) = −r0 Θ̈0 = r√ 0 /( 2ρD ). Thus, it
 √ 
z = 2ρD 1 − 2 sech Ξ(ℓ) . (95b) diverges as 1/ρ2D in the limit r0 → 2.
Since in this limit c⊥ ≫ c∥ , the scaled mean curvature is
The scaled arclength at which the pole is reached (r = 0) given only by the curvature along the meridian,
√ √
is ℓp ≈ r0 + 2( 2 − 1)ρD . Calculating the scaled area, 2
we have κ ≈ c⊥ = sech Ξ(ℓ), (100)
ρD
Zℓp
ℓ2p so the total scaled bending energy reads
a= dℓr = − + ℓp (r0 − 2ρD ) √
2 Z2
0

1
 hB ≈ dℓc2⊥ r
2
+4ρD ln √ cosh Ξ(ℓp ) . (96)
2 0
"
Using the approximation cosh x ≈ ex /2 for x ≫ 1 and = 4 ln (cosh Ξ(ℓ)) − sech2 Ξ(ℓ)
imposing the condition of unit scaled area, to first order
√
in ρD we have a quadratic equation for r0 √
! # 2
2ℓ − 2
√ − + 2 tanh Ξ(ℓ) . (101)
r02 
2ρD
+2 2 − 1 r0 ρD − 1 = 0 . (97) 0
2
Taking into account that ρD ≪ 1, we get that the scaled
Solving for r0 and taking the positive solution we get total energy diverges with the inverse of ρD ,
√ √ √
r0 = 2 − 2( 2 − 1)ρD . (98) 2(2 − 2)
hB ≈ . (102)
ρD
Therefore, the small parameter is proportional to the dif-
ference√of the maximum radius and the equatorial radius,
√ These results agree with the behavior of the total force
ρD ∝ 2−r0 , whereas
√ the total arc-length is ℓp ≈ 2. In and enegy of the vesicles in the limit of maximum stretch-
the limit r0 → 2, the geometry consists of two flat discs ing obtained from the numerical results.
 20

VII. EQUILIBRIUM SHAPES WITH FIXED cs -1 0 1 2 2.25 2.4 24/3 2.55

VOLUME
r0v sc 1.68 1.66 1.68 1.74 1.75 1.76 1.77 1.78
Θ̇v0sc 3.83 3.62 3.63 3.93 4.04 4.11 4.17 4.19
In this section we consider membranes whose enclosed
volume is fixed, so V = VS or v = 1. This case could Θ̈v0sc -4.17 -2.74 -2.38 -3.08 -3.45 -3.69 -3.91 -3.96
represent the physical situation in which the control pa- pvmc 4.44 0 -1.91 -1.3 -0.76 -0.36 0 0.1
rameter is the temperature, whereas the difference pres- pvsc -7.25 -7.56 -7.75 -7.7 -7.69 -7.67 -7.66 -7.64
sure across the membrane is fixed. In this setting the 2 v
ℓ
π p sc
1.21 1.20 1.22 1.23 1.24 1.25 1.25 1.26
membrane area varies freely on account that its ther- avsc 1.73 1.71 1.74 1.83 1.84 1.86 1.87 1.88
mal expansivity is large compared to that of the aqueous hvBmc 3.9 2 0.73 0.08 0.02 0.004 0 0.0003
medium, so that the area changes faster than the volume
hvBsc 4.25 2.4 1.51 1.58 1.74 1.89 2.03 2.06
as the temperature is varied [38, 39]. Thus, µ = 0 and 1 v
Eq. (54d) reduces to f
4π 0mc
-3.52 -2.52 -1.52 -0.52 -0.27 -0.12 0 0.03
1 v
  f
4π 0sc
6.98 4.56 4 5.36 6.05 6.52 6.93 7.04
κ sin Θ p
cos Θ κ̇ + sin Θκ − + cs + r = 0 . (103) TABLE III. Numerical values of quantities of vesicles with
2 r 2
v = 1 and different values of cs in the limits of maximum
Evaluating at a parallel with Θ0 and r0 to determine p, constriction and self contact.
we get
 
p cos Θ0 sin Θ0 κ0 sin Θ0 that of a vesicle with fixed area, it adopts an oblate shape
=− κ̇0 − κ0 − + cs . (104)
2 r0 r0 2 r0 (Figs. 20(a)-(c)), and as r0 is increased the vesicle be-
comes flat at the poles, (Fig. 20(d)). However, unlike
Similar to the case of vesicles with fixed area, for values the case of fixed area, in order to satisfy the volume con-
close to cs = 0 and up to cs = 2.25 we obtained a sin- straint it does not flatten more and more as the radius is
gle sequence of configurations. Constricted and stretched increased. Instead, the neighborhood of the pole lowers
vesicles with fixed volume are plotted in Figs. 19 and 20, (where ż ≤ 0) and as r0 is increased the poles approach
respectively. For values of the spontaneous curvature in each other, (Figs. 20(e)-(f)), until they get in contact at
the interval 2.25 < cs < 2.55 two bifurcations of the so- v
r0sc (given in Table III), while the equatorial region re-
lutions arise, plotted in Figs. 21 and 22. These two bifur- mains round (similar to the outer region of a torus), such
cations do not meet, so the sequence becomes discontin- that in this limit the generating curve has a drop-like
uous, unlike the case of fixed area where the bifurcations shape and the vesicle resembles a discocyte (Fig. 20(g)).
were connected. Although there are solutions for r0 greater than the equa-
For values of the spontaneous curvature close to cs = 0 torial radius of the discocyte, we do not consider them,
a constricted vesicle exhibits a behavior similar to the because the poles cross each other and the generating
case of fixed area. As r0 is reduced the vesicle initially curve self-intersects, which would be not only unphysi-
adopts a prolate shape (Fig. 19(a)); after the equato- cal, for it does not comply with the self-avoidance of the
rial region becomes cylindrical, there is a transition to membrane, but it would also change the topology of the
a dumbbell shape (Figs. 19(b)-(d)), and in the limit of corresponding axisymmetric membrane. It is conceivable
maximum constriction, r0 → 0, the membrane consists that for slightly larger values of r0 the membrane adopts
of two quasi-spherical vesicles connected by an infinitesi- the shape of a stretched discocyte, such that the flat cen-
mal neck, (Figs. 19(e)). However, in this case the radius tral region (where the membrane is in self-contact) be-
of the quasi-spherical vesicles is rv = 2−1/3 , so their to- comes larger as r0 is increased, while the profile curve of
tal length, area and mean curvature are ℓvp = 2−1/3 π, the outer region maintains its drop-like shape. This kind
av = 2−2/3 and k v = 24/3 . Thus, in this case the asymp- of self-contact configurations could be studied by con-
totic value of the first order correction of the mean cur- sidering a linear repulsive force [11], however it is likely
vature difference is that the self-exerted force breaks the up-down or the ax-
ial symmetries [40–42], so such analysis lies beyond the
κ(1)A = 24/3 − cs . (105) scope of this paper.
v As the spontaneous curvature is increased two bifurca-
The mean curvature of the neck, kmc := k̃(1) /r0 , is
tions appear in the sequence of constricted configura-
! tions. Relevant values of the equatorial radius of con-
v 4/3 ℓ̃ ℓ̃ figurations belonging to these bifurcations, denoted by
kmc =2 + cs 1− , (106) v
r̃(0) r̃(0) r0i , i = 1, 2, . . . , 6, are presented in Table IV. The first
bifurcation, shown in Fig. 21, occurs in the interval
v v
plotted in Fig. (23) along with the numerical data, show- r01 < r0 < r02 . At r01 it begins with a dumbbell shape
ing a good agreement, like the case of vesicles with a = 1. whose central region is almost flat (Fig. 21(a)), but as r0
The initial behavior of a stretched vesicle is similar to is increased it develops a central protrusion (Fig. 21(b))
 21

(a) r0 = 0.95 b) r0 = 0.8625 (c) r0 = 0.5 d) r0 = 10−1 e) r0 = 10−2

FIG. 19. (a)-(e) Sequence of vesicle configurations with v = 1 and cs = 0 as the radius of the ring is reduced. The scaled
bending energy density is color coded in these figures.

(a) r0 = 1.05 (b) r0 = 1.1 (c) r0 = 1.2 (d) r0 = 1.3

(e) r0 = 1.4 (f) r0 = 1.5 (g) r0 = 1.633

FIG. 20. (a)-(e) Sequence of vesicle configurations with v = 1 and cs = 0 as the radius of the ring is increased. The scaled
bending energy density is color coded in these figures.
 22

(a) r0 = 0.5 b) r0 = 0.6 (c) r0 = 0.8 (d) r0 = 1 (e) r0 = 1.45 (f) r0 = 1.8 (g) r0 = 1.9

FIG. 21. (a)-(e) Sequence of vesicle configurations with v = 1 and cs = 24/3 of the first bifurcation as the radius of the ring is
increased. The scaled bending energy density is color coded in these figures.

(a) r0 = 0.348 b) r0 = 0.4 (c) r0 = 0.6 (d) r0 = 0.857 (e) r0 = 1.2 (f) r0 = 1.5 (g) r0 = 1.63

FIG. 22. (a)-(e) Sequence of vesicle configurations with v = 1 and cs = 24/3 of the second bifurcation as the radius of the ring
is increased. The scaled bending energy density is color coded in these figures.
 23

by one of its solutions [43], namely

p
c2 − c4s − 8pcs
rs = s . (107)
2p
The values of the parameters corresponding to configu-
rations with vanishingly small necks are shown with a
dashed line in Figs. 24-28. Also, some values of the pa-
rameters for configurations in the limits of maximum con-
striction and self contact mentioned below are given in
Table III.
The plots of the initial values of the first and second
derivatives of Θ are shown in Fig. 24. For values close to
cs = 0, as the membrane is constricted, Θ̇0 decreases
monotonically and it diverges to −∞ in the limit of
FIG. 23. Mean curvature along the neck as a function of the maximum constriction. For stretched membranes Θ̇0 in-
scaled arc-length of configurations with v = 1. At the equator creases, reaching a value Θ̇0 sc for the discocytes. For
its value is given by cs and away from the equator its value cs = −1, 0 as r0 is decreased Θ̈0 increases monotonically,
tends to the mean curvature of the quasi-spherical membrane,
but as cs is increased it oscillates between a maximum
kv = 24/3 . The solid lines correspond to the numerical data
and the dashed lines to scaled first order correction k̃(1) /r0 . and a minimum after which it diverges to +∞ (−∞ for
cs > 24/3 ) in the limit of maximum constriction r0 → 0.
As the vesicle is stretched Θ̈0 decreases, reaches a mini-
mum and then it increases up to the value Θ̈0 sc for the
cs 2.4 24/3 3 discocytes. For cs > 2.25, as r0 is increased Θ̇0 increases
v along both bifurcations, ending at their corresponding
r01 0.461 0.505 0.512
v
self touching configurations. Along the first bifurcation
r02 1.894 1.904 1.907 as r0 is increased Θ̈0 first decreases toward a minimum
v
r03 0.376 0.348 0.34 and then it increases, whereas along the second bifur-
v
r04 1.623 1.632 1.635 cation it first increases rapidly, then it decreases to a
v
r05 0.925 0.855 0.839 minimum, after which it increases.
v
r06 0.804 0.859 0.872 The pressure difference p is plotted in Fig. 25. For the
initial spherical configuration p = cs (cs − 2). For values
TABLE IV. Numerical values of r0 for vesicles with v = 1 close to cs = 0, as the vesicle is constricted p increases
belonging to the bifurcations. reaching a maximum, after which it decreases reaching
a value pvmc in the limit of maximum constriction. This
limit value of p can be determined analytically from the
scaling relation, which in this case reads
which swells while remaining connected to the rounded r02 Θ̈0 = −c2s a + 2cs m + p . (108)
extremes by almost cylindrical regions (Fig. 21(c)-(d)).
By further increasing r0 the central protrusion flattens, In the limit r0 → 0, we have r02 Θ̈0 → 0, a → 2av = 21/3
so it adopts a disc-like shape (Fig. 21(e)), then its cen- and m → kv av = 22/3 , thus the pressure difference is
tral region lowers (Fig. 21(f)), until it gets in self con- given by
v
tact, such that the final configuration at r02 resembles
a discocyte connected to a dumbbell (Fig. 21(g)). The pvmc = 21/3 cs (cs − 24/3 ) , (109)
second bifurcation, shown in Fig. 22, occurs in the inter-
v v v
val r03 < r0 < r04 . At r03 it also starts with a dumb- which vanishes for cs = 0, 24/3 . This result is in good
bell (Fig. 22(a)) and as r0 is increased the central region agreement with the numerical data presented in Table
bulges (Fig. 22(b)) and swells, similar to the sequence of III. For stretched vesicles p decreases to a minimum and
configurations of the first bifurcation. However, in this then it increases reaching the value pvsc for the discocytes.
case the vesicle develops two necks, above and below of For cs > 2.25 as r0 is increased p has a similar behavior
the central region, whose radii decrease as r0 increases, along both bifurcations, it first decreases to a minimum
v
becoming vanishingly small at r05 . As a result, the vesicle and then increases.
adopts a unduloid-like shape (Fig. 22(c)), after which its The scaled total length of the generating curve 2ℓp and
central region follows a transition from a disc-like shape the total area a, plotted in Fig. 26, exhibit a similar
v
to a discocyte (in the limit of maximum stretching at r04 ), behavior. For constricted vesicles with values close to
joined to quasi-spherical vesicles above and below by in- cs = 0, as r0 is decreased 2ℓp and a increase, reaching
finitesimal necks (Figs. 22(d)-(g)). These quasi-spherical a maximum and then they decrease towards the limit of
vesicles satisfy Eq. (59) with µ = 0, so their radii is given maximum constriction, where they reach the values 22/3 π
 24

FIG. 24. Dependence of Θ̇0 and Θ̈0 on r0 for membranes with FIG. 25. Scaled pressure difference p as a function of r0 for
v = 1. The dashed lines represent configurations with very membranes with v = 1. The dashed lines represent configu-
small necks. rations with very small necks.

and 2av = 21/3 respectively. For stretching, as r0 is in- hB increases monotonically along the first bifurcation,
creased, 2ℓp decreases, reaches a minimum, after which whereas it first decreases to a minimum and then in-
it increases to the value 2ℓvp sc for the discocytes, whereas creases along the second bifurcation. The configurations
a increases to the value avsc for the same configurations. of the second bifurcation have the same energy as the con-
For cs > 2.25, as r0 is increased the total length and figurations of the original sequence of constricted vesicles
v v
the total area exhibit a similar oscillatory behavior along at r06 , so they have lower energy for r0 < r06 . Further-
4/3 v v
both bifurcations, they first increase up to a maximum more, for cs = 2 , 2.55 in the interval r0 ∈ [r05 , r06 ]
(more rapidly for the second bifurcation), then decrease these configurations of minimum total energy have van-
towards a minimum after which they increase. ishingly small necks.
The scaled total energy hB and the total force f0 are The force vanishes for the initial spherical configurations.
plotted in Figs. 27 and 28. For values close to cs = 0, the For values close to cs = 0 the force on constricted mem-
initial spherical states, whose energy is given in Eq. (49), branes exhibits an oscillatory behavior, it decreases to a
correspond to a local minimum of hB . Thus, either for minimum, then it increases to a maximum after which it
constricted or stretched membranes hB increases, reach- finally achieves the value f0vmc at the limit of maximum
ing the values hvBmc and hvBsc at the limit of maximum constriction. The exact value of the total force in the
constriction and for the discocytes, respectively. The ex- limit of maximum constriction is given by
act value in the limit of maximum constriction corre- v
f0mc
sponds to the bending energy of two spherical vesicles of = −(24/3 − cs ) . (111)
radius rv , given by 4π
Both, hvB mc and f0vmc vanish for cs = 24/3 and agree with
(24/3 − cs )2
hvBmc = . (110) the numerical data presented in Table III. For stretched
25/3 membranes, the total force increases toward a maximum
As cs is increased hB develops a maximum for constricted and then it decreases reaching the value f0vsc for the dis-
membranes and for cs > 2.25 it changes non smoothly at cocytes. For cs > 2.25, as r0 is increased f0 increases
the beginning of both bifurcations. As r0 is increased to a maximum and then it decreases along the first bi-
 25

FIG. 26. Total length of the generating curve 2ℓp and scaled
area a as functions of r0 for membranes with v = 1. The FIG. 27. Scaled total energy hB as a function of r0 for mem-
dashed lines represent configurations with very small necks. branes with v = 1. The dashed lines represent configurations
with very small necks.

furcation, whereas it first decreases to a minimum, then

increases to a maximum, after which it decreases again Constricted vesicles have a similar behavior for both con-
along the second bifurcation. straints, differing only by their size. As the equatorial ra-
dius is reduced, they exhibit a transition from prolate to
dumbbell shapes, which in turn terminates in two quasi-
VIII. CONCLUDING REMARKS AND spherical vesicles connected by a small catenoid-like neck.
DISCUSSION Although we employed a perturbative analysis to exam-
ine the geometry of such small necks, as well as the ex-
We examined the reaction of homogeneous axisymmet- ternal force required to hold them, remarkably the first
ric membranes to the force exerted by a rigid ring located order corrections suffice to determine them accurately in
at its equator. We considered membranes with different comparison with the numerical results. Unlike previous
values of spontaneous curvature and with their area or treatments where a small neck is assumed to have con-
volume fixed. For either constraint, starting with val- stant mean curvature, we found that it is described with
ues of the spontaneous curvature about zero we obtained a good degree of precision by a deformed catenoid whose
a single sequence of configurations, but as the sponta- mean curvature varies. Together, the external force and
neous curvature increases close to the mean curvature the area or volume constraints, compel the mean curva-
of the quasi-spherical vesicles in the limit of maximum ture of the neck to interpolate between the spontaneous
constriction, value for which the total force vanishes, we curvature at the equator (kissing condition) and the mean
found bifurcations in the sequence of configurations. We curvature of the quasi-spherical vesicles. A similar situ-
demonstrated that locally the effect of the external force ation occurs in the formation of a neck in the spherical
is to introduce a discontinuity in the derivatives of the confinement of a vesicle, which seems to be described by
meridional curvature of the membrane across the ring, a catenoidal shape, but by analyzing its mean curvature
whereas globally it introduces a source in the scaling re- it is found to be non-vanishing [41].
lation between the total area, volume and mean curvature Stretched vesicles with fixed area behave differently to
of the vesicle. vesicles with fixed volume as the equatorial radius is in-
 26

very small necks connecting the lobes with the central

region, akin to the remote constriction of a membrane
[25]. We found for both constraints that below a critical
value of the equatorial radius the lowest energy configu-
rations belong to one of the bifurcations, some of which
have very small necks.
There are several possible extensions and refinements of
our framework. Here we determined the configurations
of the membranes employing the spontaneous curvature
model, but it would be interesting to generalize our re-
sults using the bilayer couple model [33] or the area dif-
ference elasticity model [47, 48]. Also, we considered con-
figurations satisfying the area or the volume constraint
separately, so one of the two Lagrange multipliers van-
ishes, but by considering both of them non-vanishing one
could have access to a broader class of configurations.
We focused on relatively small values of the spontaneous
curvature, for which the vesicle follows a single sequence
of configurations and up to values where bifurcations oc-
cur for the first time. However, for higher values more
bifurcations arise, some of them showing a complex be-
havior, which complicates the identification of the con-
formations with the lowest energy. In this regard, though
laborious, it would be useful to classify such lowest en-
ergy configurations using a phase diagram. Furthermore,
to assess their stability one could analyze the spectrum
of the second variation of the energy. It might be inter-
esting to analyze the instabilities of the configurations
FIG. 28. Scaled total force f0 as function of r0 for membranes belonging to the bifurcations and to confirm that they
with v = 1. The dashed lines represent configurations with indeed decay to the configurations with lowest energy.
very small necks. Here we assumed that the deformed membrane remains
axisymmetric as well as symmetric with respect to the
equatorial plane. However, one cannot discount the pos-
sibility of new asymmetrical states, possibly stable, which
creased. In the former case, the vesicle flattens progres-
do not possess equatorial mirror symmetry, such as the
sively until it becomes disc-like, limit that we examined
shapes observed in the exobudding and endobudding pro-
using an asymptotic analysis. These stretched configu-
cesses of a vesicle [24, 25, 31], or like the configurations
rations require an increasing total force, which diverges
observed in the transition from symmetric to asymmetric
for the disc. For the latter case, the vesicle also starts
dumbbells due to a periodic change of spontaneous cur-
to flatten, but after a critical stretching the poles be-
vature [49]. For instance it is possible that constricted
gin to approach until they get in contact, so the vesi-
membranes with prolate shapes exhibit an instability by
cle achieves a discocyte shape. Along such sequence the
slipping through the ring and adopting an asymmetric
force is bounded, it increases reaching a maximum and shape. However, once the membrane adopts a dumbbell
then decreases to a finite value for the discocyte shape. shape and a neck forms, it should become stable. To relax
A similar configuration with a discocyte shape can be ob- the axial symmetry one may look at the configurations of
tained from the inversion of the catenoid with respect to a spherical membrane deformed by a non-circular loop,
a sphere centered at the origin. However in such case the say an elliptic one, which would constrict and stretch the
force is not equatorial, but applied at the poles, where membrane along orthogonal directions, producing a force
it gives rise to logarithmic singularities in the curvature dipole on the equator.
[44–46]. We considered a ring of infinite bending rigidity, so it re-
The increase of the spontaneous curvature elicits the ap- mains circular regardless of the force exerted by the mem-
pearance of bifurcations in the solutions corresponding brane, which for instance diverges in the unphysical limit
to constricted membranes. These bifurcations are con- of the disc-like configuration. More realistically, the ring
nected for vesicles with fixed area but disconnected for will have a finite bending rigidity, and under confinement
vesicles with fixed volume. In general, the vesicles be- it will not remain circular, but adjust to its confinement
longing to these bifurcations tend to develop lobes in the [50]. Even if the ring rigidity is large but finite, since
regions near the poles, while the central region bulges. the total force ought to remain finite, one would expect
For large enough values of the spontaneous curvature, that axial symmetry breaking occurs before the disc-like
as the equatorial radius is increased the vesicle develops
 27

shape is attained. Likewise, it is conceivable that if the by

equator of the discocyte shape is stretched, the planar
central region enlarges and the force increases up to a ρ̂ = cos φ x̂ + sin φ ŷ , (A1a)
point that, in order to minimize its bending energy, the φ̂ = − sin φ x̂ + cos φ ŷ , (A1b)
membrane adopts a non axi-symmetric configuration.
The opposing limit corresponding to an elastic loop of the embedding functions and the two tangent vectors
fixed length confined within a rigid sphere was consid- adapted to this parametrization are spanned as
ered in Ref. [12]. The ground state of the loop was found
X(l, φ) = R(l) ρ̂ + Z(l) ẑ , (A2a)
to assume a saddle shape. In general both the loop and
the membrane will deform in response to the constraint el = R′ ρ̂ + Z ′ ẑ , (A2b)
imposed by the confinement, which involves playing two eφ = Rφ̂ , (A2c)
competing tendencies against each other. Technically,
the determination of the shape will involve the non-trivial where R(l) and Z(l) are the radial and height coordi-
task of minimizing the sum of the membrane and loop nates; the prime indicates derivation with respect to l.
bending energies. Solving the coupled EL equations of Since, R(l) and Z(l) are parametrized by arc-length, their
the membrane-loop complex is not likely to be tractable derivatives are related by R′ (l)2 + Z ′ (l)2 = 1. This con-
analytically. The ground state of a confined loop would dition permits us to define
again be expected to assume a saddle shape in a deformed
geometry that resembles an ellipsoid with its long axis R′ = cos Θ , Z ′ = sin Θ , (A3)
aligned along the direction that the transmitted force is where Θ is the angle that the tangent along the meridian
maximum. However, less obvious possibilities should not el makes with the radial direction ρ̂.
be discounted. In contrast, in a membrane with a con- In this parametrization the metric tensor is diagonal
stricted equator, the circular loop will minimize the total
bending energy so that the behavior should not change. gll = 1 , gφl = glφ = 0 , gφφ = R2 . (A4)
One would also still expect that a finite force will be re-
quired to constrict the membrane completely. The line element on the surface is given by ds2 = dl2 +
Instead of considering the deformation of the membrane R(l)2 dφ2 , whereas the area element is dA = Rdφdl, so
due to a ring, one could also study its deformation by the area of a region with φ ∈ [0, 2π] and l ∈ [li , lf ] is
an open polymer, which also plays a role in shaping the
morphology of membranes. In particular, the protein Zlf
reticulin is suspected to be implicated in determining the A = 2π dlR . (A5)
morphology of the endoplasmic reticulum. To examine li
the equilibrium shapes of a polymer-membrane complex,
one could consider a semi-flexible rigid polymer segment The outward unit normal vector, defined by n =
of fixed length, be it straight, a hairpin, or the arc of a g −1/2 eφ × el , reads
circle, bounded to a fluid membrane.
n = sin Θ ρ̂ − cos Θ ẑ . (A6)

The enclosed volume is determined by the integrating

R of
the support function over the surface, V = 1/3 dAX·n.
ACKNOWLEDGEMENTS
For an axisymmetric surface

We have benefited from conversations with Saša Zlf

2π
Svetina. B. B. acknowledges partial support by the Slove- V = dlR (sin ΘR − cos ΘZ) . (A7)
nian research agency through the research program P1- 3
l0
0055. P. V.-M. acknowledges support by SECIHTI un-
der programs Investigadores por México (grant 439-2018) In the case of a closed surface symmetric with respect to
and Sistema Nacional de Investigadores (131142) and is the plane Z = 0 and satisfying Z(0) = 0 and R(lf ) = 0,
grateful for the hospitality of the Josef Stefan Institute this equation can be simplified by integrating by parts,
and the Institute of Biophysics of the University of Ljubl-
jana. Zlf
V = 2π dl sin ΘR2 . (A8)
0

Appendix A: Axisymmetric surfaces The extrinsic curvature tensor is diagonal

Kll = Θ′ , Kφl = Klφ = 0 Kφφ = R sin Θ . (A9)

An axially symmetric membrane can be parametrized
by the arc length l along its meridian and the azimuthal The shape operator K ab = g ac Kcb is also diagonal, so
angle φ. In terms of the cylindrical basis {ρ̂, φ̂, ẑ}, given its eigenvectors lie along meridians and parallels and the
 28

principal curvatures are the corresponding eigenvalues. stresses on the membrane. Thus, omitting the constant
These curvatures also correspond to the normal curva- terms −σA0 + P V0 , the Lagrangian is
tures along meridians and parallels, obtained from the Z lp
L
projections of the curvature tensor onto the outward =2 dlL , (B3)
conormal and the tangent vectors to a parallel, which 2π 0
are given by where the constrained Lagrangian density is defined by
1 P̄ 2
L = −el , T = φ̂ = eφ , (A10) L = LB + σ̄ R − R sin Θ
R 2
+ Rλ̄R (R − cos Θ) + Rλ̄Z (Z ′ − sin Θ) . (B4)
′
so the components are Ll = −1 and Tφ = 1/R. Thus,
the projections of Kab onto L and T are The conjugate momenta to Θ and to the coordinates R
and Z are
C⊥ = La Lb Kab = Θ′ , (A11a) ∂L
sin Θ PΘ := = RKD , (B5a)
C∥ = Ta Tb Kab = . (A11b) ∂Θ′
R ∂L
PR := = RλR , (B5b)
The geodesic torsion vanishes along a parallel τg = ∂R′
Ta Lb Kab = 0, because it is a principal curve. Thus the ∂L
PZ := = RλZ . (B5c)
tangential component f∥⊥ of the stress tensor projected ∂Z ′
along the parallel outward, given in Eq. (11b) vanishes. The corresponding Hamiltonian density is
Thus, the projected stress tensor is orthogonal to the  
PΘ sin Θ P̄
parallel, f⊥ = f⊥⊥ L + f⊥ n. Using the expressions of the H = PΘ − + Cs − σ̄R + R2 sin Θ
vectors L and T, Eqs. (A10), along with the expressions 2R R 2
of the adapted basis (A2b), (A2c) and (A6), we get that + PR cos Θ + PZ sin Θ . (B6)
the projections onto the unit vectors ρ̂ and ẑ are Since H does not depend explicitly on l, it is conserved,
H ′ = 0. Moreover, in order to make L stationary with
f⊥ · ρ̂ = sin Θf⊥ − cos Θf⊥⊥ , (A12a)
respect to total length variations, the Hamiltonian must
f⊥ · ẑ = − cos Θf⊥ − sin Θf⊥⊥ . (A12b) vanish
H = 0, (B7)
Appendix B: Hamiltonian formulation
which avoids the introduction of an unphysical constraint
fixing the interval of l, (the arc length at the pole, lp , is
The bending energy (scaled by 1/kB ), given in Eq. (1), not fixed) [39, 44, 45].
of an axially symmetric geometry with mirror symmetry The Hamilton equations for Θ, R and Z are identified as
with respect to the equatorial plane is
Z lp PΘ − sin Θ
LB Θ′ = + Cs , (B8a)
=2 dlLB , (B1) R
2π 0 R′ = cos Θ , (B8b)
where the energy density is given by LB = 2
RKD /26 , Z ′ = sin Θ , (B8c)
with KD the mean curvature difference, defined by whereas those for the corresponding momenta are
sin Θ
 
KD = Θ′ + − Cs . (B2) ′ PΘ P̄ 2
R PΘ = − R − PZ cos Θ + PR sin Θ , (B9a)
R 2
 
In order to implement in the variational principle the ′ PΘ PΘ
PR = − sin Θ − P̄R sin Θ + σ̄ , (B9b)
constraints of fixed area and volume, given by Eqs. (A5) R2 2
and (A8), we introduce two global Lagrange multipliers PZ′ = 0 . (B9c)
σ and P. Another two local Lagrange multipliers λR and
λZ are also introduced to enforce the relations between The momentum PZ is constant because Z is a cyclic co-
derivatives of R and Z and the angle Θ, given by Eq. ordinate.
(A3), which together imply that l is arc-length. As shown Multipliying Eqs. (B9a) and (B6) by cos Θ and sin Θ re-
below, these two Lagrange multipliers are related to the spectively and taking their difference in order to elimi-
nate PR , we get
 ′    
PΘ PΘ sin Θ
R cos Θ + sin Θ PΘ − + Cs − σ̄ R
6 The Gaussian R 2R R
R energy densityR is not included because it is a total
derivative, dAKG = −2π dl(cos Θ)′ , so it does not contribute P̄
to the energy of the bulk. + R 2 + PZ = 0 . (B10)
2
 29

Dividing by R, using Eq. (B5) to replace PΘ and PZ in An important example of a surface with a net verti-
favor of KD and λZ , we obtain a first order differential cal force is provided by the Clifford Torus, whose ra-
equation for the mean curvature difference dial, height and functions are R = RT + rc cos θ and
    Z = rc sin θ, where θ = l/rc (Θ = θ√+ π/2) and the two
′ KD sin Θ constant radii are related by RT = 2 rc , which satisfies
cos ΘKD + sin Θ KD − + Cs − σ̄
2 R Eq. (B14) for the following values of the parameters
P̄  
+ R + λ̄Z = 0 . (B11) 2 Cs
2 σ̄ = Cs − , (B16a)
rc 2
Thus, in order to determine the equilibrium configura- 2Cs
tions, we can solve the equivalent system of first order P̄ = , (B16b)
rc2
differential equations, given by Eqs. (B8) and (B11) for 1
R, Z, Θ, and KD , where the equations for the last two Rλ̄Z = − − 2Cs . (B16c)
rc
variables can be recast as We see that although σ and P vanish on a toroidal vesi-
sin Θ cle with null spontaneous curvature, it requires a non-
Θ′ = KD − + Cs , (B12a) vanishing vertical force.
R
    The other Lagrange multiplier λR can be determined
′ KD sin Θ
KD = − tan Θ KD − + Cs − σ̄ from Eq. (B9a), which divided by R and using Eqs. (B5)
2 R for the momenta, can be recast as
 
P̄
− sec Θ R + λ̄Z . (B12b) 
P̄

2 ′
λR sin Θ = KD + cos Θ R + λZ . (B17)
2
Substituting Eqs. (15) for the projections of the stress
tensor in Eq. (B11), the first integral of Eq. (9) is ex- Recall that f¯⊥ = K ′ , using Eq. (B13) to substitute λZ
pressed as a balance of stresses and using Eq. (A12a) for the radial projection of f⊥ , we
identify λR as the radial force density on the membrane
P
cos Θf⊥ + sin Θf⊥⊥ + R + λZ = 0 . (B13) λR = sin Θf⊥ − cos Θf⊥⊥ = f⊥ · ρ̂ . (B18)
2
Substituting Eq. (A12b), in Eq. (B13), we determine the By taking into account that at the poles R(lp ) = 0, the
Lagrange multiplier λZ constraint of fixed radius at equator, R(0) = R0 , can be
Rl
expressed as 0 p dlR(l)′ = −R0 , which is implemented
P with a constant Lagrange multiplier λn , so L → L +
λZ = f⊥ · ẑ − R. (B14)
2 (R0 /2)λ̄n R′ . Since the added term is a total derivative, it
does not modify either the Hamilton or the EL equations
The momentum PZ can be determined from the bound- for R. However it modifies the momentum conjugate to
ary termsR in the variation of L in Eq. (B3), which are R in the boundary term in Eq. (B15) corresponding to
given by dsδQ′ , where R, PR → PR +(R0 /2)λ̄n , so at the equator where δΘ = 0
and δZ = 0, we have R0 λ̄n = −2PR (0), or by using Eq.
δQ = PΘ δΘ + PR δR + PZ δZ . (B15) (B18) as well as that Θ0 = π/2, we obtain
For a surface with spherical topology, the angle Θ and λ̄n = −2λ̄R (0) = −2f¯⊥ (0) = −2Θ′′0 , (B19)
the radial coordinate R are fixed at the poles, thus δΘ
and δR vanish, whereas δZ is arbitrary at the poles so PZ which reproduces the force density, given in Eq. (34),
must vanish [39], so the first integral of the EL equation, and whose integration along the ring provides the total
given by Eq. (20), is reproduced. constraining force, F̄ = −4πR0 Θ′′0 .

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