1. Introduction
Multicomponent lipid vesicles – composed of saturated lipids, unsaturated lipids and cholesterol – serve as minimal models for biological membranes. Their lateral phase separation into liquid-ordered (
$l_o$
) and liquid-disordered (
$l_d$
) domains (Baumgart, Hess & Webb Reference Baumgart, Hess and Webb2003; Veatch & Keller Reference Veatch and Keller2003, Reference Veatch and Keller2005; Lingwood & Simons Reference Lingwood and Simons2010; Wang et al. Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022) leads to spatial heterogeneity in membrane properties such as bending rigidity and surface tension (Evans & Rawicz Reference Evans and Rawicz1990; Baumgart et al. Reference Baumgart, Das, Webb and Jenkins2005; Yanagisawa, Imai & Taniguchi Reference Yanagisawa, Imai and Taniguchi2008). These heterogeneities underlie lipid rafts, which play critical roles in signal transduction (Simons & Ikonen Reference Simons and Ikonen1997; Simons & Toomre Reference Simons and Toomre2000; Sackmann & Smith Reference Sackmann and Smith2014), membrane trafficking (Simons & Vaz Reference Simons and Vaz2004) and molecular sorting (Mukherjee & Maxfield Reference Mukherjee and Maxfield2004). Understanding how such heterogeneity modulates membrane deformation under flow is crucial for the design of biomimetic materials (Elani, Law & Ces Reference Elani, Law and Ces2014) and drug delivery in cell biology (Allen & Cullis Reference Allen and Cullis2004; Herrmann, Wood & Fuhrmann Reference Herrmann, Wood and Fuhrmann2021; Kooijmans et al. Reference Kooijmans, de Jong and Schiffelers2021; Karaz & Senses Reference Karaz and Senses2023).
The dynamics of single-component vesicles in shear flow has been extensively investigated through experimental, theoretical and numerical methods, revealing a rich variety of behaviours such as tank treading, tumbling and swinging, primarily driven by viscosity contrast (Keller & Skalak Reference Keller and Skalak1982; Abkarian & Viallat Reference Abkarian and Viallat2005; Kantsler & Steinberg Reference Kantsler and Steinberg2006; Misbah Reference Misbah2006; Danker et al. Reference Danker, Biben, Podgorski, Verdier and Misbah2007; Noguchi & Gompper Reference Noguchi and Gompper2007; Vlahovska & Gracia Reference Vlahovska and Gracia2007; Lebedev, Turitsyn & Vergeles Reference Lebedev, Turitsyn and Vergeles2008; Deschamps, Kantsler & Steinberg Reference Deschamps, Kantsler and Steinberg2009). In contrast, the dynamics of multicomponent vesicles under shear flow remains poorly understood. Two-dimensional simulations have successfully reproduced phase treading and tumbling driven by bending heterogeneity and asymmetric phase distributions (Lowengrub, Xu & Voigt Reference Lowengrub, Xu and Voigt2007; Sohn et al. Reference Sohn, Tseng, Li, Voigt and Lowengrub2010; Liu et al. Reference Liu, Marple, Allard, Li, Veerapaneni and Lowengrub2017; Gera, Salac & Spagnolie Reference Gera, Salac and Spagnolie2022). However, these models are limited to planar dynamics and cannot account for three-dimensional (3-D) phenomena, such as vertical ring banding (Gera & Salac Reference Gera and Salac2018a , Reference Gera and Salacb ), neither can they incorporate line tension, which experimental studies have shown can induce tumbling (Tusch et al. Reference Tusch, Loiseau, Al-Halifa, Khelloufi, Helfer and Viallat2018). Recent 3-D simulations provide insights into the interplay of bending heterogeneity and flow (Venkatesh & Narsimhan Reference Venkatesh and Narsimhan2024), but are limited by assumptions of small deformations and neglect surface tension heterogeneity. Likewise, the two-phase fluid deformable surface sharp-interface model proposed in Sischka & Voigt (Reference Sischka and Voigt2025) incorporates bending heterogeneity but fails to capture topological transitions accounting for surface tension heterogeneity. To our knowledge, most numerical models enforce local membrane inextensibility by introducing a spatially varying Lagrange multiplier Arroyo & DeSimone (Reference Arroyo and DeSimone2009), Veerapaneni et al. (Reference Veerapaneni, Young, Vlahovska and Bławzdziewicz2011), Sohn et al. (Reference Sohn, Li, Li and Lowengrub2012), Aland et al. (Reference Aland, Egerer, Lowengrub and Voigt2014), Young, Veerapaneni & Miksis (Reference Young, Veerapaneni and Miksis2014), Lowengrub, Allard & Aland (Reference Lowengrub, Allard and Aland2016), often interpreted as an surface tension – a purely kinematic reaction field. Although composition-dependent tensions appear in several theoretical frameworks Lowengrub et al. (Reference Lowengrub, Xu and Voigt2007), Li, Lowengrub & Voigt (Reference Li, Lowengrub and Voigt2012), Destainville, Manghi & Cornet (Reference Destainville, Manghi and Cornet2018), Wu & Ou-Yang (Reference Wu and Ou-Yang2025), the dynamical consequences of the associated thermodynamic tension gradients have received relatively little attention.
To overcome these limitations, we developed a thermodynamically consistent phase-field model for multicomponent vesicles under shear flow. Our model effectively captures the coupling between background fluid and membrane dynamics, together with lateral phase separation, while incorporating line tension and component-dependent physical properties. Our 3-D simulations demonstrate excellent qualitative and quantitative agreement with experimental observations, accurately capturing phase separation and shear-induced dynamics. We further explore how shear-induced dynamics are governed by bending rigidity heterogeneity
$(\kappa _B=\kappa _{B_{l_d}}/\kappa _{B_{l_o}})$
, surface tension heterogeneity (
$\sigma _S=\sigma _{S_{l_d}}/\sigma _{S_{l_o}}$
) and bending capillary number
${\textit{Ca}}=\mu _{\textit{out}}UL^2/\kappa _{B_{l_o}}$
. Our results uncover a range of complex dynamic regimes, including three previously unreported ones. Notably, we identify a surface-tension-driven swinging and tumbling mechanism that arises without viscosity contrast or asymmetric phase distributions. These findings reveal previously unexplored physical pathways for regulating vesicle dynamics and offer guidance for biomimetic vesicle design in flow environments.
The outline of the paper is as follows. Section 2 presents a thermodynamically consistent phase-field model for multicomponent vesicles, and introduces key dimensionless parameters. Section 3 validates the model against experimental observations (Tusch et al. Reference Tusch, Loiseau, Al-Halifa, Khelloufi, Helfer and Viallat2018; Wang et al. Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022). Section 4 reports numerical results, elucidating the interplay between membrane heterogeneity and flow-induced forces. The appendices present additional methodological and numerical details.
2. Model
In this section, we develop a thermodynamically consistent phase-field model derived from the total energy of the system to investigate the dynamics of multicomponent vesicles with membrane heterogeneity under shear flow within a rectangular domain
$\varOmega$
(see figure 1).
(a) A multicomponent vesicle with lipid components (
$l_d$
(red) and
$l_o$
(blue)) exhibiting distinct properties, immersed in a background fluid, illustrating the lipid bilayer structure. For simplicity, the two components are distinguished only by colour. (b) Diffuse-interface representation of panel (a), showing the corresponding phase-field distribution.

2.1. Governing equations for vesicle membrane interacting with background fluid
We consider a vesicle immersed in a background fluid within a rectangular domain
$\varOmega$
, whose membrane is represented by a closed surface
$\varGamma$
. The system is modelled as a two-phase flow, with
$\varGamma$
serving as the interface between the inner and outer fluids. To capture complex shape deformations, we adopt a phase-field formulation in which the sharp interface
$\varGamma$
is replaced by a diffuse interface characterized by a scalar field
$\phi (\boldsymbol{x}, t)$
. The phase-field variable
$\phi$
follows a hyperbolic tangent profile across the membrane interface, ensuring a smooth transition from the interior (
$\phi = 1$
) to the exterior (
$\phi = -1$
) of the vesicle. Specifically,
$\phi$
can be defined as
\begin{align} \phi (\boldsymbol{x}, t) = \tanh \left (\frac { r(\boldsymbol{x}, t)}{2\sqrt {2}\varepsilon _\phi }\right )\!, \quad \boldsymbol{x} \in \bar {\varOmega }, \end{align}
where
$r(\boldsymbol{x}, t)$
denotes the signed distance from
$\boldsymbol{x}$
to the nearest point on the membrane interface
$\varGamma$
, taken positive inside and negative outside, and
$\varepsilon _\phi$
controls the thickness of the diffuse interface.
Both the inner and outer fluids are assumed to be incompressible, with equal density (
$\rho _{{in}} = \rho _{\textit{out}}$
) but distinct viscosities, denoted by
$\mu _{{in}}$
and
$\mu _{\textit{out}}$
. The governing equations for the coupled two-phase flow and membrane evolution are formulated as follows:
where (2.2) and (2.3) represent the conservation of momentum and fluid incompressibility via the Navier–Stokes equations, and (2.4) describes the evolution of the vesicle membrane using a phase-field Cahn–Hilliard (CH) formulation. The membrane inextensibility condition is enforced through (2.5). Here,
$\boldsymbol{u}$
and
$p$
denote the bulk velocity and pressure, respectively. The rate of strain tensor is given by
$ \unicode{x1D63F}(\boldsymbol{u}) = (\boldsymbol{\nabla } \boldsymbol{u} + \boldsymbol{\nabla } \boldsymbol{u}^{{T}})/2$
. The spatially varying viscosity is interpolated as
$\mu (\phi ) = {(1-{\phi })\mu _{in}}/{2} + {(1+{\phi })\mu _{out}}/{2}$
. The vesicle–fluid interaction force
$\boldsymbol{F}$
and the interfacial flux
$\boldsymbol{q}_\phi$
will be defined later.
In addition, membrane inextensibility is modelled using the diffuse domain approach of Aland et al. (Reference Aland, Egerer, Lowengrub and Voigt2014), which extends the local surface constraint
$\boldsymbol{\nabla }_{\varGamma } \boldsymbol{\cdot }\boldsymbol{u} = \unicode{x1D64B} : \boldsymbol{\nabla } \boldsymbol{u} = 0$
from the vesicle membrane
$\varGamma$
to the entire computational domain
$\varOmega$
. Here, the surface gradient operator is defined as
where
$ \unicode{x1D64B} = \boldsymbol{I} - \boldsymbol{n} \otimes \boldsymbol{n}$
denotes the projection tensor onto the tangential plane of the membrane, with the unit normal vector
$\boldsymbol{n} = -\boldsymbol{\nabla } \phi / |\boldsymbol{\nabla } \phi |$
pointing from the interior to the exterior of the vesicle. The inextensibility constraint is enforced by introducing a local Lagrange multiplier
$\lambda _{local}$
together with a harmonic regularization term, leading to (2.5), where the parameter
$\xi \gt 0$
controls the strength of the regularization. The regularized surface delta function is then defined as
where
$f(\phi ) = {(1 - \phi ^2)^2}/{4}$
is a double-well potential,
$\varepsilon _\phi$
is the membrane thickness and
$\eta _\phi$
is a scaling parameter that relates the diffuse interface to the sharp-interface model. The resulting constraint term contributes additional membrane tension forces in the momentum equation, completing the formulation of (2.2).
Furthermore, the velocity
$\boldsymbol{u}$
is subject to Dirichlet boundary conditions, while the phase-field variable
$\phi$
and its associated flux
$\boldsymbol{q}_{\phi }$
satisfy no-flux boundary conditions,
where
$\boldsymbol{n_b}$
denotes the unit outward normal vector to the boundary
$\partial \varOmega$
of the rectangular computational domain
$\varOmega$
.
2.2. Governing equation for multicomponent on the vesicle membrane
We model the vesicle membrane
$\varGamma$
as a two-phase fluid consisting of liquid-ordered (
$l_o$
) and liquid-disordered (
$l_d$
) phases with distinct physical properties. The membrane composition is described by a second scalar phase-field variable
$c_{\varGamma }$
defined on the membrane
$\varGamma$
, where
$c_{\varGamma } = 1$
corresponds to the
$l_d$
phase, and
$c_{\varGamma } = -1$
to the
$l_o$
phase. The two components are separated by a diffuse interface of thickness
$\varepsilon _c$
, across which
$c_{\varGamma }$
transitions smoothly. The phase separation dynamics on
$\varGamma$
are governed by the surface CH equation (Zhiliakov et al. Reference Zhiliakov, Wang, Quaini, Olshanskii and Majd2021),
where the diffusive flux of the membrane component,
$\boldsymbol{q}_c$
, will be specified later.
To couple the surface phase dynamics (2.10) with the bulk phase-field system (2.2)–(2.5) in the rectangular domain
$\varOmega$
, we employ the diffuse domain method (Rätz & Voigt Reference Rätz and Voigt2006; Li et al. Reference Li, Lowengrub, Rätz and Voigt2009; Teigen et al. Reference Teigen, Li, Lowengrub, Wang and Voigt2009), which is an accurate and efficient numerical approach for solving partial differential equations (PDEs) on complex surfaces. The vesicle membrane
$\varGamma$
is embedded into the larger, rectangular domain
$\varOmega$
using a regularized surface delta function
$\zeta (\phi )$
derived from the phase-field variable
$\phi$
. This embedding eliminates the need for explicit interface tracking. Moreover, the method establishes a unified, energy-based framework by extending surface PDEs into the bulk domain
$\varOmega$
through the regularized delta function
$\zeta (\phi )$
(2.7). Accordingly, the surface CH (2.10) is reformulated in
$\varOmega$
as
Details of this extension are provided in Appendix A. Notably, previous studies have demonstrated that in the diffuse domain method, the extended PDE (2.11) asymptotically converges to the original surface PDE (2.10) as the diffuse interface thickness approaches zero (
$\varepsilon _{\phi } \to 0$
) (Li et al. Reference Li, Lowengrub, Rätz and Voigt2009; Yu, Chen & Thornton Reference Yu, Chen and Thornton2012; Lervåg & Lowengrub Reference Lervåg and Lowengrub2015; Chadwick et al. Reference Chadwick, Stewart, Enrique, Du and Thornton2018; Poulsen & Voorhees Reference Poulsen and Voorhees2018; Yu, Guo & Lowengrub Reference Yu, Guo and Lowengrub2020; Guo et al. Reference Guo, Yu, Lin, Wise and Lowengrub2021; Guo Reference Guo2023).
In addition, the membrane component
$c_\varGamma$
and its flux
$\boldsymbol{q_{c}}$
are subject to no-flux boundary conditions on the boundary of the rectangular domain
$\varOmega$
,
Thanks to the diffuse domain method, the membrane component equation for
$c_{\varGamma }$
(2.11) can be naturally coupled with the vesicle–fluid interaction system (2.2)–(2.5) and solved simultaneously within the same rectangular domain
$\varOmega$
. The unknown terms
$\boldsymbol{F}$
,
$\boldsymbol{q_{\phi }}$
and
$\boldsymbol{q_{c}}$
in (2.2)–(2.5) and (2.11) will be derived via an energy-variation approach based on the principle of free energy dissipation, thereby ensuring thermodynamic consistency.
2.3. Free energy of the multicomponent vesicle–fluid system
The free energy
$\mathcal{F}$
of a multicomponent vesicle immersed in a background fluid comprises contributions from both the fluid and the membrane. These include the kinetic energy of the background fluid
$ \mathcal{F}_{\textit{K}}$
, the membrane bending energy
$ \mathcal{F}_{\textit{B}}$
, the surface energy
$ \mathcal{F}_{\textit{S}}$
, the line energy
$ \mathcal{F}_{\textit{L}}$
and a penalty term
$ \mathcal{F}_{\textit{A}}$
enforcing surface area conservation. The free energy is expressed as
\begin{align} \mathcal{F} &= \mathcal{F}_{{K}} + \mathcal{F}_{{B}} + \mathcal{F}_{{S}} + \mathcal{F}_{{L}} + \mathcal{F}_{{A}} \nonumber\\ &= \int _{\varOmega } \frac {1}{2} \rho |\boldsymbol{u}|^2 \, \mathrm{d}V + \int _{\varOmega } \frac {1}{2 \varepsilon _\phi } \kappa _B(c_\varGamma ) \omega _\phi ^2 \, \mathrm{d}V + \int _{\varOmega } \sigma _S(c_\varGamma ) \zeta (\phi ) \, \mathrm{d}V \nonumber\\ &\quad + \int _{\varOmega } \sigma _L \zeta (c_\varGamma ) \zeta (\phi ) \, \mathrm{d}V \,\,+ \frac {M_s}{2} \frac {(S(\phi ) - S(\phi _0))^2}{S(\phi _0)}. \end{align}
In the bending energy
$\mathcal{F}_{{B}}$
, the scalar field
serves as a phase-field approximation of the membrane mean curvature. Here, zero spontaneous curvature is assumed and the contribution from Gaussian curvature is neglected. The bending rigidity
$\kappa _B(c_\varGamma )$
, associated with the bending energy
$\mathcal{F}_{{B}}$
, and the surface tension
$\sigma _S(c_\varGamma )$
, defined in the surface energy
$\mathcal{F}_{{S}}$
, are both modelled as functions of the membrane component field
$c_\varGamma$
,
In the line energy
$ \mathcal{F}_{{L}}$
, the function
serves as a regularized delta function that captures the diffuse interface between membrane components. Here, the double-well potential
$ f(c_\varGamma ) = (1 - c_\varGamma ^2)^2/4$
promotes phase separation,
$\varepsilon _c$
represents the interfacial thickness between the components,
$\eta _c$
is a scaling parameter and
$ \sigma _L$
denotes the line tension. In the penalty energy
$\mathcal{F}_{{A}}$
, the vesicle surface area is defined as
$S(\phi ) = \int _\varOmega \zeta (\phi ) \, \mathrm{d}V$
. The reference area
$S(\phi _0)$
is measured at the initial time
$t = 0$
and assumed constant during the simulation. Surface area conservation is enforced by this penalty term, with its strength controlled by the penalty coefficient
$M_S$
, which has been found to help improve the accuracy of the method (Aland et al. Reference Aland, Egerer, Lowengrub and Voigt2014).
By applying an energy-variation approach, we derive explicit expressions for the terms
$\boldsymbol{F}, \boldsymbol{q_\phi }$
and
$\boldsymbol{q_c}$
that were previously left unspecified in the governing equations. Detailed derivations are provided in Appendix B.
2.4. Non-dimensionalization
To non-dimensionalize the model, we choose the characteristic length, velocity and time scales as
$L$
,
$U$
and
$L/U$
, respectively. Fluid properties are normalized with respect to the outer fluid (where
$\phi = -1$
), using characteristic viscosity
$\mu _{\textit{out}}$
and density
$\rho _{\textit{out}}$
, while membrane properties are normalized by those of the
$l_o$
phase (
$c_\varGamma = -1$
). The non-dimensional form of the multicomponent vesicles–fluid model is then given by
where
with
The chemical potential associated with the membrane component field
$c_\varGamma$
is given by
\begin{align} \zeta (\phi )\omega _c &=\frac {1}{\textit{Ca}} \frac {{\varepsilon _\phi }^{-1}}{2}{\kappa _B}^{\prime }(c_{\varGamma }) {\omega _{\phi }}^{2} + Cs {\sigma _S}^{\prime }(c_{\varGamma }) {\zeta (\phi )} \nonumber \\ &\quad + {\textit{Cn}}_{\varGamma } \zeta (\phi )\left ({\eta _c} \frac {f^{\prime }(c_{\varGamma })}{\varepsilon _c}-{\eta _c} {\varepsilon _c} \varDelta c_{\varGamma }\right )- {\textit{Cn}}_{\varGamma } {\eta _c} {\varepsilon _c} \boldsymbol{\nabla } \zeta (\phi ) \boldsymbol{\cdot }\boldsymbol{\nabla }{c_{\varGamma }}. \end{align}
Equations (2.17) and (2.18) describe the dynamics of the incompressible background fluid, while the evolution of the vesicle membrane is governed by (2.19). The local inextensibility of the vesicle is enforced via (2.20), and the lateral phase separation of membrane components is governed by (2.21). The boundary conditions are applied as specified in (2.8), (2.9) and (2.12).
Notably, we emphasize that
$\sigma (c_\varGamma )$
and
$\lambda _{\textit{local}}$
play fundamentally different roles in our model. The field
$\lambda _{\textit{local}}$
arises solely from the geometric inextensibility constraint and therefore cannot represent composition-dependent surface tension. In contrast,
$\sigma (c_\varGamma )$
is a genuine material property that encodes the membrane’s intrinsic energetics, and its spatial gradients can actively drive Marangoni-type tangential flows. Thus, the two quantities operate at distinct physical levels:
$\lambda _{\textit{local}}$
enforces kinematic inextensibility, whereas
$\sigma (c_\varGamma )$
governs the membrane’s thermodynamic response to compositional heterogeneity. Although their associated tangential stresses may locally reinforce or partially counteract each other under certain flow conditions, the constraint-induced tension cannot generally replace or compensate for material surface tension since it is uniquely determined by the inextensibility requirement. A systematic investigation of how constraint-induced tension redistributes in the presence of heterogeneous material tension is beyond the scope of the present work and will be considered in future studies.
The vesicle dynamics described by (2.17)–(2.27) are governed by a set of dimensionless parameters that characterize the coupling between the background fluid and membrane dynamics, the lateral phase separation and the influence of line tension and component-dependent physical properties. These dimensionless parameters are summarized in table 1.
Physical and dimensionless parameters used in the simulations.

Our model integrates key physical features, including membrane component-dependent bending rigidity heterogeneity, surface tension heterogeneity, line tension, inextensibility and viscosity contrasts between the internal and external fluids. Its thermodynamically consistent formulation ensures free energy dissipation and physical consistency, enabling the simulation of complex dynamic processes under realistic biological conditions. This comprehensive framework provides a powerful tool for investigating vesicle behaviour under shear flow and for elucidating the role of membrane heterogeneity in governing non-equilibrium dynamics.
3. Numerical validations
To validate the model, we conducted systematic comparisons between our simulations and two sets of experimental observations: (i) phase separation dynamics on giant unilamellar vesicles (GUVs) reported in Wang et al. (Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022), and (ii) dynamic behaviours of prepositioned multicomponent vesicles under shear flow reported in Tusch et al. (Reference Tusch, Loiseau, Al-Halifa, Khelloufi, Helfer and Viallat2018). The numerical method is detailed in Appendix C, and additional numerical convergence tests are presented in Appendix D to verify the accuracy of the diffuse-interface formulation and to validate both the numerical method and the spatial–temporal resolutions used in all simulations. At the fully discrete level, we employ BSAM – a highly efficient block-structured adaptive full approximation scheme multigrid solver – for the simulations. The solver combines a staggered finite-difference method with a full approximation scheme multigrid algorithm and is specifically designed to address diffuse-interface problems (Guo et al. Reference Guo, Lin, Lowengrub and Wise2017). Within this framework, the velocity components
$\boldsymbol{u} = (u, v, w)$
are stored at edge centres, while all scalar quantities, including pressure and phase-field variables, are stored at cell centres. Details of the spatial discretization can be found in Guo et al. (Reference Guo, Lin, Lowengrub and Wise2017). Our simulations exhibit excellent qualitative and quantitative agreement with both experiments. These validations confirm the accuracy, effectiveness and predictive capability of the proposed model in capturing phase behaviours, vesicle–fluid coupling and dynamic transitions of multicomponent vesicles under shear flow.
3.1. Phase separation on stationary multicomponent vesicles
We first investigate phase separation dynamics on the membrane of multicomponent vesicles in comparison with the experimental results reported in Wang et al. (Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022). That study examined the phase behaviour of GUVs and provided detailed observations of domain evolution. To facilitate a direct comparison, we simplify our set-up by assuming a stationary vesicle with a fixed spherical shape, focusing on capturing the essential features of membrane phase separation.
Here
$\bar {\varOmega } = [-1.5, 1.5]^3$
and is discretized using a uniform Cartesian grid of
$128^3$
. The fixed spherical vesicle is represented by a phase-field with radius
$r_0=1$
, corresponding to a physical vesicle radius
$R = 10 \,{\unicode{x03BC}}$
m (Wang et al. Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022), and membrane thickness
$\varepsilon _\phi = 0.02$
. The initial phase-field variable
$\phi (\boldsymbol{x}, 0)$
is prescribed as
\begin{align} \phi (\boldsymbol{x}, 0) = \tanh \left (\frac {r_0 - r(\boldsymbol{x})}{2\sqrt {2}\varepsilon _\phi }\right )\!, \quad r(\boldsymbol{x}) = \sqrt {x^2 + y^2 + z^2}, \quad \boldsymbol{x} =(x,y,z) \in \bar {\varOmega }, \end{align}
representing a smooth diffuse interface centred around the vesicle surface. To model an initially homogeneous liposome, we first generate random numbers
$ c_{\textit{rand}}(\boldsymbol{x})$
with a uniform distribution between 0 and 1. Then, we apply the following cutoff for the initial surface fraction
$ c_\varGamma (\boldsymbol{x}, 0)$
:
\begin{align} c_{\varGamma }(\boldsymbol{x}, 0) = \begin{cases} \,\,1 \quad (l_{d}), & \text{if } c_{\textit{rand}}(\boldsymbol{x}) \in [0, a_{l_d}], \\ -1 \quad (l_{o}), & \text{if } c_{\textit{rand}}(\boldsymbol{x}) \in (a_{l_d}, 1], \end{cases} \end{align}
where
$a_{l_d}$
and
$a_{l_o} = 1 - a_{l_d}$
represent the prescribed area fractions of the
$l_d$
and
$l_o$
phases, respectively.
No-flux boundary conditions are applied to the phase-field variables
$\phi$
,
$c_{\varGamma }$
and
$\omega _c$
at the domain boundary
$\partial \varOmega$
.
The model parameters are chosen to match those reported in Wang et al. (Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022): line tension
$\sigma _L = 1.2 \,\text{pN}$
and diffusion coefficient
$D_c = 10^{-5}\,\text{cm}^2\,\text{s}^{-1}$
. The lipid mobility is adjusted such that
${\textit{Cn}}_\varGamma /{\textit{Pe}}_\varGamma = 0.12$
. Bending rigidity and surface tension are assumed to be uniform across the
$l_o$
and
$l_d$
phases, with
$\kappa _B = \sigma _S = 1$
. The interfacial thickness of membrane components is set to
$\varepsilon _c = 0.02$
, fitted to match experimental GUVs.
Two area fractions,
$a_{l_d} = 0.3$
and
$a_{l_d} = 0.7$
, are considered to match the experimental conditions in Wang et al. (Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022). To account for stochastic variability, we perform 10 independent simulations with different random seeds of
$c_{\varGamma }$
for each area fraction.
Figures 2 and 3 show the phase separation process on the membrane for area fractions
$ a_{l_d} = 0.3$
and
$ a_{l_d} = 0.7$
, respectively, compared with the experimental results from Wang et al. (Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022). During phase separation, membrane components coarsen, forming distinct, stable domains, as illustrated in the rightmost images of both figures 2 and 3 that closely match the experimental snapshots.
Phase separation on GUVs at
$a_{l_d} = 0.3$
: experimental snapshots (a) from Wang et al. (Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022) and numerical results (b), showing
$l_o$
domains (blue) forming within the
$l_d$
phase (red).

Phase separation on GUVs at
$a_{l_d} = 0.7$
: experimental snapshots (a) from Wang et al. (Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022) and numerical results (b), showing
$l_d$
domains (red) forming within the
$l_o$
phase (blue).

To further validate the accuracy of our model, we compare the time evolution of the number of lipid domains obtained from our numerical simulations with the experimental measurements reported in Wang et al. (Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022) (digitized using WebPlotDigitizer https://automeris.io/). In the simulations, lipid domains are identified directly on the membrane surface by extracting the isosurface
$\phi =0$
, interpolating
$c_\varGamma$
onto this surface, thresholding at
$c_\varGamma =0$
and applying MATLAB’s standard surface-connected-component labelling routine bwconncomp (https://www.mathworks.com/help/images/ref/bwconncomp.html.) As shown in figure 4, for both area fractions under investigation (
$a_{l_d} = 0.3$
and
$a_{l_d} = 0.7$
), the experimental data (markers) are generally bounded by the minimum (green) and maximum (red) values from the ensemble of our simulations. The average numerical result (blue) also lies in close agreement with the experimental trend.
Time evolution of number of lipid rafts for
$a_{l_d} = 0.3$
(a) and
$a_{l_d} = 0.7$
(b). Markers represent experimental data; solid lines represent numerical results (blue, average; green, minimum; red, maximum).

Time evolution of the total perimeter of lipid domains at
$a_{l_d} = 0.3$
(a) and
$a_{l_d} = 0.7$
(b). Markers represent experimental data; solid lines represent numerical results.

Experimental images (a) from Wang et al. (Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022) and numerical results (b) showing the time evolution of
$l_o$
domains (blue) within the
$l_d$
phase (red). The simulation is initialized with 12 symmetrically placed
$l_o$
domains and an area fraction
$a_{l_o} = 0.64$
.

In addition, we further compare the evolution of the total perimeter of lipid domains, quantified by
$\int _\varOmega \zeta (\phi ) \zeta (c_\varGamma )\, \mathrm{d}V$
in our simulations, where
$\zeta (\phi )$
localizes the membrane and
$\zeta (c_\varGamma )$
localizes the phase boundary, with the experimental data (digitized using WebPlotDigitizer), as shown in figure 5. The simulation results (solid lines), averaged over all numerical simulations, exhibit excellent agreement with the experimental measurements (markers) for both
$a_{l_d} = 0.3$
and
$a_{l_d} = 0.7$
.
As reported in Wang et al. (Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022), three distinct structural patterns were observed experimentally at an area fraction of
$a_{l_d} = 0.3$
prior to equilibrium: (i) the minority
$l_d$
phase domains (red) embedded within the majority
$l_o$
phase (blue), as shown in figure 2; (ii) majority
$l_o$
phase domains (blue) enclosed within the minority
$l_d$
phase (red), as illustrated in figure 6; and (iii) a mixed pattern featuring both types of domains, as depicted in figure 7.
Patterns (ii) and (iii) were attributed to the persistence of a metastable third phase, which influences domain coarsening and leads to fluctuations in domain area fractions before equilibrium (Van Santen Reference Van Santen1984; Anderson & Lekkerkerker Reference Anderson and Lekkerkerker2002). For instance, the measured
$l_o$
area fraction in case (ii) was approximately 6 % lower than its equilibrium value.
Following Wang et al. (Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022), we reproduce the experimentally observed patterns by adjusting the initial conditions: (i) prescribing prepositioned, symmetrically arranged
$l_o$
domains within an
$l_d$
background, and (ii) reducing the
$l_o$
area fraction by approximately
$6\,\%$
. Accordingly, the second pattern –
$l_o$
domains fully enclosed within the
$l_d$
phase (figure 6) – is reproduced by initializing 12 symmetrically placed
$l_o$
domains with an area fraction
$a_{l_o}=0.64$
. The third pattern, featuring coexisting
$l_o$
and
$l_d$
domains (figure 7), is obtained by initializing six symmetrically distributed
$l_o$
domains and setting
$a_{l_o}=0.65$
.
The quantitative and qualitative agreement between our reproduced simulations and the experimental observations demonstrates the effectiveness and accuracy of our model in capturing the phase dynamics associated on the vesicle membrane.
Experimental images (a) from Wang et al. (Reference Wang, Palzhanov, Quaini, Olshanskii and Majd2022) and numerical results (b) with front and rear views over time, showing a combination of
$l_o$
phase (blue) and
$l_d$
phase (red). The simulation is initialized with six symmetrically placed
$l_o$
domains and an area fraction
$a_{l_o} = 0.65$
.

3.2. Dynamics of a two-component vesicle under shear flow
To assess the capability of the model in capturing membrane–fluid interactions, we reproduce the experimental observations of multicomponent GUVs under shear flow reported in Tusch et al. (Reference Tusch, Loiseau, Al-Halifa, Khelloufi, Helfer and Viallat2018). These experiments revealed dynamic behaviours such as tank treading, swinging and tumbling, driven by asymmetric phase distributions and periodic elastic energy storage along the contact line.
In our simulations,
$\bar {\varOmega } = [-2, 2]^3$
, discretized on a uniform grid with
$128^3$
. To match the experimental set-up, the initial vesicle shape is modelled as a prolate ellipsoid with a specified reduced volume. The initial phase variable is given by
\begin{align} \phi (\boldsymbol{x}, 0) = \tanh \left (\frac {r_0 - r(\boldsymbol{x})}{2\sqrt {2}\varepsilon _\phi }\right )\!, \quad r (\boldsymbol{x})= \sqrt {\frac {x^2}{d_1^2} + \frac {y^2}{d_2^2} + \frac {z^2}{d_3^2}}, \quad \boldsymbol{x} = (x, y, z) \in \bar {\varOmega }, \end{align}
where
$d_1 \gt d_2 = d_3$
, resulting in elongation along the
$X$
-axis (see figure 8). The membrane component
$c_{\varGamma }$
is initially prepositioned at the tips of the vesicle along the
$X$
-axis, with a prescribed area fraction
$a_{l_d}$
representing the fraction of the
$l_d$
phase.
(a) Geometric parameters used to characterize vesicle dynamics under shear flow. The vesicle shape is described by its principal semiaxes
$(d_1,d_2,d_3)$
, and the inclination angle
$\theta$
is defined as the angle between the major axis
$d_1$
and the flow direction (X-axis). (b) Initial phase distribution used in the simulations of § 4, shown in the X–Z view.

Experimental images (a) from Tusch et al. (Reference Tusch, Loiseau, Al-Halifa, Khelloufi, Helfer and Viallat2018) and numerical results (b) showing tank treading and swinging at
$ {\textit{Cn}}_\varGamma = 8$
, for a vesicle with reduced volume
$v = 0.95$
, flattening
$f = 0.3$
and area fraction
$a_{l_d} = 0.7$
.

The shear flow is introduced by imposing boundary conditions. Specifically, Dirichlet boundary conditions
$\boldsymbol{u}_{bc} = (\gamma z, 0, 0)$
are imposed at the top and bottom boundaries in the
$z$
-direction, where
$\gamma$
denotes the dimensionless shear rate (see table 1). No-flux boundary conditions are enforced for all phase-field variables (
$\phi$
,
$\omega _\phi$
,
$c_\varGamma$
,
$\omega _c$
) in the
$z$
-direction, while periodic boundary conditions are applied in the
$x$
- and
$y$
-directions for all variables.
Physical parameters are mainly taken from Tusch et al. (Reference Tusch, Loiseau, Al-Halifa, Khelloufi, Helfer and Viallat2018): mean vesicle radius
$R = 10 \,{\unicode{x03BC}} \text{m}$
, fluid viscosity
$\mu _{in} = \mu _{out} = 1.2 \times 10^{-3} \,\text{Pa}\boldsymbol{\, }\text{s}$
, bending rigidity
$\kappa _{B_{l_o}} = \kappa _{B_{l_d}} = 10^{-19}\,\text{J}$
and shear rate
$\dot {\gamma } = 3\,\text{s}^{-1}$
. Fluid densities are taken as
$\rho _{in} = \rho _{out} = 1000 \,\text{kg}\,\text{m}^{-3}$
(Luby-Phelps Reference Luby-Phelps1999b
). Surface tensions are fixed at
$\sigma _{S_{l_d}} = \sigma _{S_{l_o}} = 10^{-6}\,\text{N}\,\text{m}^{-1}$
to ensure uniform membrane properties. The line tension
$\sigma _L$
is varied as
$4,1$
and
$0.1\,\text{pN}$
to reproduce the three dynamical regimes observed experimentally.
Using
$R$
as the characteristic length and
$\dot {\gamma }R$
as the velocity scale, the resulting dimensionless parameters are Reynolds number
${\textit{Re}} = 3 \times 10^{-4}$
, bending capillary number
${\textit{Ca}} = 36$
, Péclet numbers
${\textit{Pe}} = 1000$
and
${\textit{Pe}}_\varGamma = 10$
, dimensionless surface tension strength
$Cs_\varGamma = 27$
and surface area penalty coefficient
$\mathcal{M}_{{S}} = 2000$
. The Cahn number is set as
${\textit{Cn}}_\varGamma = 8,\,2,\,0.2$
for the three regimes (figures 9–11). Interfacial thickness parameters
$\varepsilon _c = 0.05$
for membrane components and
$\varepsilon _\phi = 0.05$
for the membrane are fitted to experimental GUVs.
Figures 9–11 illustrate vesicle dynamics under different reduced volumes, showing close agreement with the experimental results from Tusch et al. (Reference Tusch, Loiseau, Al-Halifa, Khelloufi, Helfer and Viallat2018). In figure 9, the initial ellipsoid with
$ d_1 = 1$
,
$ d_2 = d_3 = 0.7$
, yields
$ v \approx 0.95$
, flattening parameter
$ f = ({d_1 - d_2})/{d_1} = 0.3$
and area fraction
$ a_{l_d} = 0.7$
. At
${\textit{Cn}}_\varGamma =8$
, the vesicle exhibits tank treading and swinging characterized by a steady prolate shape with oscillatory inclination angle.
At a moderate reduced volume
$ v \approx 0.8$
, shown in figure 10, with
$ d_1=1, d_2 = d_3 = 0.35$
,
$ f = 0.65$
and
$ a_{l_d} = 0.6$
, the vesicle exhibits pronounced deformation at
${\textit{Cn}}_\varGamma =2$
. It appears elliptical when the
$ l_o$
–
$ l_d$
interface aligns with the short axis and resembles two adhered vesicles when aligned with the long axis, exhibiting tank treading and swinging accompanied by large deformations.
At lower volume
$v \approx 0.7$
(figure 11), with
$d_1=1, d_2 = d_3 = 0.25$
,
$f = 0.75$
and
$a_{l_d} = 0.5$
, the vesicle transitions to a tumbling regime at
${\textit{Cn}}_\varGamma =0.2$
. The interface length between
$l_o$
and
$l_d$
is markedly reduced along the long axis due to line tension, which drives the system to minimize line energy, facilitating large deformations and promoting tumbling under shear.
Experimental images (a) from Tusch et al. (Reference Tusch, Loiseau, Al-Halifa, Khelloufi, Helfer and Viallat2018) and numerical results (b) showing strong deformation during tank treading and swinging at
$ {\textit{Cn}}_\varGamma = 2$
, for a vesicle with reduced volume
$ v = 0.8$
, flattening parameter
$ f = 0.65$
and area fraction
$ a_{l_d} = 0.6$
.

Experimental images (a) from Tusch et al. (Reference Tusch, Loiseau, Al-Halifa, Khelloufi, Helfer and Viallat2018) and numerical results (b) showing the tumbling at
$ {\textit{Cn}}_\varGamma = 0.2$
, for a vesicle with reduced volume
$ v = 0.7$
, flattening parameter
$ f = 0.75$
and area fraction
$ a_{l_d} = 0.5$
.

To further validate our model, we track the time evolution of the inclination angle
$\theta$
of a vesicle undergoing tank treading and swinging behaviours at reduced volume
$v = 0.95$
and area fraction
$a_{l_d} = 0.7$
(see figure 9). The inclination angle
$\theta$
is defined as the angle between the vesicle’s long axis and the flow direction, constrained within
$\theta \in [-{\pi }/{2}, {\pi }/{2} ]$
(see figure 8). As shown in figure 12, the simulated results (solid line) closely match the experimental measurements (circles) reported in Tusch et al. (Reference Tusch, Loiseau, Al-Halifa, Khelloufi, Helfer and Viallat2018) (digitized using WebPlotDigitizer), capturing the characteristic oscillations of the tank treading and swinging regime with a mean angle of approximately
$30^\circ$
. The numerical results remain consistently within the experimental variability, thereby confirming not only the accuracy and robustness of the model but also its predictive capability in reproducing flow-induced vesicle dynamics.
Time evolution of inclination angle
$ \theta$
for a vesicle with reduced volume
$ v = 0.95$
, flattening parameter
$ f = 0.3$
and area fraction
$ a_{l_d} = 0.7$
at
${\textit{Cn}}_\varGamma =8$
, comparing numerical results (solid line) with experimental data (circles) from Tusch et al. (Reference Tusch, Loiseau, Al-Halifa, Khelloufi, Helfer and Viallat2018).

4. Numerical experiments
In this section, we investigate the flow-induced dynamics of multicomponent vesicles, with a particular focus on the coupling between membrane heterogeneity and hydrodynamic forces. Through comprehensive numerical simulations, we demonstrate that the competition among shear flow, bending rigidity heterogeneity and surface tension heterogeneity governs complex vesicle behaviours, including shape deformation, domain evolution and transitions between distinct dynamic regimes.
4.1. Bending heterogeneity-driven dynamics
We systematically explore the coupling between shear flow and bending heterogeneity in multicomponent vesicles by varying the bending rigidity contrast
$\kappa _B = \kappa _{B_{l_d}} / \kappa _{B_{l_o}}$
and the bending capillary number
${\textit{Ca}} = \mu _{\textit{out}} U L^2 / \kappa _{B_{l_o}}$
, both readily tuneable in experiments.
The numerical set-up closely follows § 3.2. The vesicle is initialized as a prolate ellipsoid with principal axes
$d_1 = 1$
and
$d_2 = d_3 = 0.565$
, as defined in (3.3), corresponding to a reduced volume
$v \approx 0.91$
. Two symmetric
$l_d$
-phase domains (red) are prepositioned at the vesicle tips along the
$X$
-axis (see figure 8), with an initial area fraction
$a_{l_d} = 0.4$
for the membrane component field
$c_{\varGamma }$
. The dimensionless shear rate is set to
$\gamma = 1$
.
Moreover, the dimensionless parameters are specified as follows: Reynolds number
${\textit{Re}} = 10^{-3}$
; Cahn number
${\textit{Cn}}_{\varGamma } = 0.05$
; Péclet numbers
${\textit{Pe}} = 1000$
and
${\textit{Pe}}_\varGamma = 1$
; dimensionless surface-tension strength
$Cs_{\varGamma } = 10$
; surface-area penalty coefficient
$\mathcal{M}_{\mathrm{S}} = 2000$
; surface tension contrast
$\sigma _S = 1$
. In all cases, the
$l_d$
phase is assumed to be softer than the
$l_o$
phase, i.e.
$\kappa _{B_{l_d}} \lt \kappa _{B_{l_o}}$
.
Figure 13 presents a phase diagram spanned by
$(\kappa _B, Ca)$
, capturing diverse dynamics of multicomponent vesicles and identifying six distinct dynamical regimes.
-
(i) Lateral ring banding. The vesicle adopts an oblate, tilted shape, with
$ l_d$
phase domains stretching along the edges to form a lateral ring band composed of a single slender domain. -
(ii) Lateral broken-ring banding. Similar to (I), but the
$ l_d$
phase domains form a broken lateral ring composed of two slender domains. -
(iii) Tank treading. The vesicle assumes a prolate, tilted shape, with
$ l_d$
phase domains stably located at the high-curvature tips (reported in Liu et al. (Reference Liu, Marple, Allard, Li, Veerapaneni and Lowengrub2017) and Venkatesh & Narsimhan (Reference Venkatesh and Narsimhan2024)). -
(iv) Vertical broken-ring banding. The
$l_d$
phase domains stretch vertically along the flow direction on the membrane of a prolate, tilted vesicle. This results in a vertical broken ring composed of two slender domains. -
(v) Vertical ring banding. Similar to (IV), but the
$ l_d$
phase domains merge into a complete vertical ring band composed of a single slender domain (reported in Gera & Salac (Reference Gera and Salac2018b
)). -
(vi) Phase treading. The
$ l_d$
phase domains migrate periodically along the membrane of a prolate, tilted vesicle from one tip to the other tip in the flow direction, while maintaining the initial domain symmetry (reported in Liu et al. (Reference Liu, Marple, Allard, Li, Veerapaneni and Lowengrub2017), Gera & Salac (Reference Gera and Salac2018b
) and Venkatesh & Narsimhan (Reference Venkatesh and Narsimhan2024)).
Note that, regimes (I), (II) and (IV) have not been previously reported and represent novel dynamical regimes driven by membrane bending heterogeneity under shear flow.
Phase diagram of vesicle dynamics under shear flow as a function of bending rigidity contrast
$\kappa _B$
and capillary number
${\textit{Ca}}$
. Regimes (I)–(VI) are distinguished by colours and markers.

The phase diagram also reveals two prominent ‘dynamical belts’: (i) a ‘low-rigidity full-dynamics belt’ (
$ 0.25 \leq \kappa _B \leq 0.65$
), where the membrane rigidity contrast is small enough for all six regimes to appear across a wide range of
${\textit{Ca}}$
; and (ii) a ‘low-capillary full-dynamics belt’ (
$0.027 \lesssim Ca \lesssim 0.043$
), where relatively weak shear allows all six regimes to emerge over a range of
$\kappa _B$
.
Within the ‘low-rigidity full-dynamics belt’ (
$0.25 \leq \kappa _B \leq 0.65$
), increasing
${\textit{Ca}}$
drives a flow-mediated transition pathway through the dynamical regimes: at low
${\textit{Ca}}$
, weak shear and dominant bending energy lead the vesicle to adopt an oblate shape, with low-rigidity
$l_d$
domains accumulating along high-curvature lateral edges, forming (I) lateral ring banding. As
${\textit{Ca}}$
increases moderately, enhanced shear stresses partially rupture the
$l_d$
domains at the lateral edges, producing (II) lateral broken-ring banding. With further increase in
${\textit{Ca}}$
, the vesicle elongates into a prolate shape, and the
$l_d$
domains stabilize at the high-curvature tips, resulting in (III) tank treading. At higher
${\textit{Ca}}$
, the
$l_d$
domains stretch vertically along the membrane, but insufficient membrane fluidity prevents full closure into a ring, resulting in (IV) vertical broken-ring banding. At sufficiently high
${\textit{Ca}}$
, the
$l_d$
domains are further stretched and coalesce into (V) vertical ring banding. Under even stronger
${\textit{Ca}}$
, persistent and periodic domain migration emerges, characterized by symmetric phase migration, known as (VI) phase treading.
Outside the ‘low-rigidity full-dynamics belt’ (
$\kappa _B \gt 0.65$
), the diversity of accessible dynamical regimes progressively narrows. For
$0.65 \leq \kappa _B \lt 0.75$
, only regimes (III)–(VI) remain, while for
$\kappa _B \gt 0.75$
, regime (III) also vanishes. This reduction in dynamical regimes results from the diminishing bending-rigidity contrast between the
$l_d$
and
$l_o$
phases, which weakens the tendency of low rigidity
$l_d$
domains to localize at high-curvature regions and suppresses bending-driven behaviours such as (I) lateral ring banding.
Notably, within the ‘low-capillary full-dynamics belt’ (
$0.027 \leq Ca \lesssim 0.0435$
) shown in figure 13, we observe a complete sequence of transitions from regime (I) to regime (VI) as
$\kappa _B$
increases. This highlights the strong sensitivity of both domain and shape evolution to bending rigidity contrast under moderate shear, and identifies this belt as a critical parameter space for capturing diverse vesicle dynamical regimes.
We further observe that the critical capillary numbers separating successive regimes-for example, the transition from (IV) to (V) – generally decrease with increasing
$\kappa _B$
, except for the (V) to (VI) transition at
$\kappa _B \leq 0.45$
. This trend indicates that stronger flow is typically required to induce domain migration and evolution under pronounced bending heterogeneity, reflecting a competition between shear-induced forces and membrane bending rigidity contrast.
Moreover, within the (V) vertical ring banding regime in figure 13, we identify three subregimes based on ring dynamics.
-
(V a ) Temporary ring. The
$l_d$
domains briefly merge into a vertical ring that ruptures and stabilizes into a broken-ring configuration. -
(V b ) Cyclic broken and closing ring. The
$l_d$
domains repeatedly merge into a vertical ring that undergoes persistent cycles of rupture and reconnection. -
(V c ) Stable ring. The
$l_d$
domains experiences several rupture-reconnect cycles but eventually stabilizes and remains intact over long times.
These subregimes emerge from transitions driven by the bending heterogeneity
$\kappa _B$
and capillary number
${\textit{Ca}}$
. At
$\kappa _B \geq 0.65$
, a weak bending heterogeneity reduces the preference of
$l_d$
domains for high-curvature tips, restricting dynamics to (V
$_{{a}}$
). For
$0.45 \lt \kappa _B \lt 0.65$
, increasing
${\textit{Ca}}$
triggers sequential transitions from (V
$_{{a}}$
) to (V
$_{{b}}$
) and ultimately to (V
${c}$
). At
$\kappa _B \leq 0.45$
, a strong heterogeneity destabilizes the ring, suppressing (V
$_{{c}}$
). Notably, (V
$_{{a}}$
) is distinct from regime (IV): in (V
$_{{a}}$
), a transient ring forms and then ruptures, whereas in (IV), the
$l_d$
domains never coalesce into a ring.
Figure 14 illustrates six distinct vesicle dynamic regimes (I)–(VI) at
$\kappa _B = 0.65$
within the ‘low-rigidity full-dynamics belt’. Figure 14(a) shows their temporal evolution (see supplementary Movie-1, Movie-2, Movie-3, Movie-4, Movie-5 and Movie-6, corresponding to regimes (I)–(VI), respectively), are available at https://doi.org/10.1017/jfm.2026.11276, while figure 14(b) presents the corresponding bending and line energy profiles from
$t = 2$
to
$t = 38$
. To capture the key dynamical processes, we mark the critical turning points (
$t_{i}$
for
$i=1,\ldots ,9$
, see the caption of figure 14) on the energy curves. These turning points of bending and line energies largely coincide.
(a) Time evolution of six vesicle dynamic regimes (I)–(VI) at
$ \kappa _B = 0.65$
with varying
${\textit{Ca}}$
. (b) Corresponding evolution of bending and line energies. Key time points:
$t_1 = 2.6$
,
$t_2 = 9$
,
$t_3 = 11.5$
,
$t_4 = 15$
,
$t_5 = 17.5$
,
$t_6 = 18.7$
,
$t_7 = 25$
,
$t_8 = 26.7$
,
$t_9 = 32$
.

(a) Time evolution of subregimes (V
$_a$
)–(V
$_c$
) within regime (V) at
$ \kappa _B = 0.45$
with varying
${\textit{Ca}}$
. (b) Corresponding evolution of bending and line energies. Key time points:
$t_1 = 8.2$
,
$t_2 = 11.6$
,
$t_3 = 14.3$
,
$t_4 = 20.5$
,
$t_5 = 25.7$
,
$t_6 = 28.9$
,
$t_7 = 43.6$
,
$t_8 = 62$
,
$t_9 = 66$
.

Comparing the
$ l_d$
phase domain positions in figure 14(a) with the energy evolution in figure 14(b) provides valuable insights into vesicle dynamics. In regimes (I)–(III),
$l_d$
domains (with lower rigidity
$\kappa _{B_{l_d}}$
) localize at high-curvature regions, minimizing bending energy and resulting in similar energy levels. However, due to the relatively low
${\textit{Ca}}$
, bending forces dominate over viscous forces, preventing further reduction in line energy. In regime (IV), the
$l_d$
domains stretch from the high-curvature tips towards low-curvature regions away from the tips, leading to a temporary increase in bending energy from
$t_1$
to
$t_3$
. Unable to overcome the energy barrier required to form a vertical ring, both the bending and line energies stabilize thereafter at
$t_7$
. In regime (V), the
$l_d$
domains initially stretch towards low-curvature membrane regions away from the tips, increasing bending and line energies from
$t_1$
to
$t_2$
. At
$t_4$
, the domains merge into a vertical ring band, minimizing both energies. After
$t_9$
, the domains stabilize into a broken-ring configuration. Strictly, this case at
$\kappa _B = 0.65$
corresponds to subregime (V
$_{{a}}$
), with detailed subregime dynamics presented in figure 15. In (VI),
$l_d$
domains periodically migrate along the membrane. Bending energy peaks when domains align along flatter regions (e.g.
$t_2$
,
$t_7$
) and decreases when they reach high-curvature tips (e.g.
$t_4$
,
$t_9$
).
To further investigate the (V) vertical ring banding regime, figure 15 presents three subregimes: (V
$_{{a}}$
), (V
$_{{b}}$
) and (V
$_{{c}}$
). Figure 15(a) shows the temporal evolution of each dynamics subregime, while figure 15(b) shows the corresponding bending and line energies from
$t=2$
to
$t = 70$
, with key transition points (
$t_i$
,
$i = 1, \ldots , 9$
, see the caption of figure 15) marked. Consistent with figure 14, the
$l_{d}$
phase domain positions correlate with energy evolution. We find that the three subregimes exhibit similar energy evolution before
$t_4$
, as all initially form a vertical ring. Subsequent divergence arises from differences in
$l_d$
domain evolution. In (V
$_{{a}}$
), the ring ruptures and stabilizes as a broken-ring after
$t_7$
, with both bending and line energies reaching stability. In (V
$_{{b}}$
), the ring undergoes periodic rupture–reconnection, leading to cyclic oscillations of the energy: bending energy increases during rupture (e.g.
$t_3$
to
$t_4$
) as more low-rigidity
$l_d$
domains migrate to low-curvature regions away from the tips, and decreases during reconnection (e.g.
$t_5$
to
$t_6$
). In (V
$_{{c}}$
), after two rupture–reconnection cycles (from
$t_3$
to
$t_8$
), the
$l_d$
domains form a stable vertical ring at
$t_8$
, with both bending and line energies stabilizing thereafter. Notably, the equilibrium energies in (V
$_{{c}}$
) are higher than in (V
$_{{a}}$
) due to (i) a larger fraction of low-rigidity
$l_{d}$
domains remaining in low-curvature regions, increasing bending energy and (ii) a longer
$l_d$
-
$l_o$
interface, elevating line energy.
4.2. Surface tension heterogeneity-driven swinging and tumbling
We next investigate the role of membrane surface tension heterogeneity
$\sigma _S = \sigma _{S_{l_d}} / \sigma _{S_{l_o}}$
in modulating vesicle dynamics under shear flow. Within the ‘low-rigidity full-dynamics belt’ at a fixed bending rigidity contrast
$ \kappa _B = 0.65$
, we systematically vary
$ \sigma _S \leq 1$
and the bending capillary number
${\textit{Ca}}$
.
The numerical set-up follows § 4.1. In all cases, the
$l_d$
phase is assumed to possess lower surface tension than the
$l_o$
phase, i.e.
$\sigma _{S_{l_d}} \lt \sigma _{S_{l_o}}$
.
The phase diagram (figure 16) reveals eight dynamical regimes: the six previously identified (I–VI) and two additional regimes.
-
(vii) Swinging. Small-amplitude oscillations of the vesicle inclination angle, coupled with cyclic rupture and reconnection of the
$l_d$
-formed vertical ring, similar to (V
$_{{b}}$
). -
(viii) Tumbling. Rigid-body-like vesicle rotation, with
$l_d$
domains stably anchored at high-curvature tips.
Swinging and tumbling dynamics have been previously reported in single-component vesicles with viscosity contrast, and in multicomponent vesicles with small-deformation bending heterogeneity (Gera et al. Reference Gera, Salac and Spagnolie2022; Venkatesh & Narsimhan Reference Venkatesh and Narsimhan2024) or with shape asymmetry arising from initial phase distributions and bending heterogeneity (Liu et al. Reference Liu, Marple, Allard, Li, Veerapaneni and Lowengrub2017). Here, we identify a novel mechanism: surface tension heterogeneity, without viscosity contrast or asymmetric phase distributions, can also induce swinging and tumbling. As shown in figure 16, regimes (VII) and (VIII) occur only under heterogeneous surface tension (
$\sigma _S \lt 1$
).
(a) Phase diagram of vesicle dynamics under shear flow as a function of surface tension contrast
$\sigma _S$
and capillary number
${\textit{Ca}}$
at fixed bending rigidity contrast
$\kappa _B = 0.65$
. Eight dynamic regimes (I)–(VIII) are identified and distinguished by colours and markers. (b) Inclination angle evolution in regime (VII) at
$\sigma _S = 0.85$
,
${\textit{Ca}} = 0.02$
. (c) Inclination angle evolution in regime (VIII) at
${\textit{Ca}} = 1$
for varying
$\sigma _S$
.

Under uniform surface tension (
$\sigma _S = 1$
), moderate capillary numbers (
$0.0189 \lesssim {\textit{Ca}} \lesssim 0.037$
) drive the vesicle into regime (IV), characterized by vertical broken-ring banding. As
$\sigma _S$
decreases to 0.85, enhanced membrane fluidity enables
$l_d$
domains to overcome the energy barrier, forming a complete ring. Periodic rupture and reconnection of this vertical ring dynamically alter the inclination angle, resulting in regime (VIII). At high capillary numbers (
$ 0.037 \lesssim Ca \leq 1$
), uniform surface tension (
$\sigma _S = 1$
) induces regimes (V), characterized by vertical ring banding, and (VI), defined by phase treading. As
$\sigma _S$
decreases to 0.85, regime (VIII) tumbling emerges. This transition occurs because the lower-tension
$l_d$
domains become strongly anchored at the high-curvature tips, suppressing their migration and introducing hydrodynamic asymmetry that promotes rigid-body-like rotation.
Overall, decreasing
$\sigma _S$
lowers the energetic cost of deforming
$l_d$
domains, enhancing their mobility and reducing the critical capillary number
${\textit{Ca}}$
required for successive dynamic regime transitions. As a result, at
$0.85 \leq \sigma \lt 1$
only regimes (II)–(IV), (VII) and (VIII) persist, regime (I) becomes unsustainable under shear, while swinging and tumbling emerge as the dominant dynamic responses. In addition, for
$0.35 \leq \sigma _S \lt 0.85$
, only regimes (II), (III) and (VIII) persist. In this range, strong surface tension contrast firmly anchors
$l_d$
domains at the tips, promoting vesicle rotation and lowering the threshold for tumbling, while suppressing other regimes as domain evolution rapidly stabilizes. When
$\sigma _S \lt 0.35$
, only (VIII) is observed across all tested
${\textit{Ca}}$
, indicating that extreme surface tension asymmetry robustly favours rigid-body-like rotation.
The distinct signatures of (VII) swinging and (VIII) tumbling are further quantified in figure 16(b), which show the time evolution of the vesicle inclination angle for regimes (VII) and (VIII). In (VII) Swinging, the angle exhibits step-like segments linked to the rupture and reconnection of the
$l_d$
ring. As
$l_d$
domains stretch towards low-curvature regions, surface tension gradients reduce the inclination angle; after rupture, domains shift back to the tips, increasing the angle. These non-smooth variations reflect the interplay between domain evolution and shear force. In (VIII) tumbling, further decreasing
$\sigma _S$
at a fixed
${\textit{Ca}}$
accelerates the angular rotation, driven by stronger surface tension gradients.
In addition, figure 17 provides a comprehensive depiction of the (VII) swinging and (VIII) tumbling regimes. Figure 17(a) illustrates their temporal evolution (see supplementary Movie-7 and Movie-8, corresponding to regimes (VII)–(VIII), respectively), while figure 17(b) presents the corresponding bending, line and surface energies starting from
$t = 2$
to
$t = 40$
. The critical time points (
$t_i$
, for
$i = 1, \ldots , 9$
; see caption of figure 17) are marked on the energy curves. The evolution of bending and line energies, which reflects the migration of
$l_d$
domains, is consistent with the discussion in § 4.1 and is not repeated here. The following analysis focuses on the variations in surface energy.
In (VII) swinging, as shown in figure 17(a), periodic rupture and reconnection of the ring cause surface energy to oscillate. When the ring reconnects,
$l_d$
domains stretch into low-curvature regions, increasing both their area and the surface energy (e.g. around
$t_4$
). When the ring ruptures,
$l_d$
domains cluster near the high-curvature tips, reducing their area and lowering surface energy (e.g. around
$t_5$
).
In (VIII) tumbling, the
$l_d$
domains remain anchored at the tips throughout the vesicle’s rotation. The vesicle alternates between elongated shapes, tilted along the flow direction (with inclination angle
$\theta \in (0, \pi /2)$
), and flattened shapes, tilted against the flow (
$\theta \in (-\pi /2, 0)$
). When elongated (e.g. at
$t_3$
), the vesicle adopts a thin, prolate shape, where the
$l_o$
domains are located in relatively low-curvature regions. When flattened (e.g. at
$t_5$
), the vesicle deforms into a more oblate shape under shear flow, causing the curvature at the
$l_o$
locations to decrease further. This curvature reduction lowers the surface energy. Such cyclic shape evolution, driven by the interplay between shear forces and surface tension heterogeneity, accounts for the periodic surface energy variations observed in the tumbling regime.
(a) Time evolution of vesicle dynamic regimes (VII) and (VIII) at
$ \sigma _S = 0.85$
and
$ \kappa _B = 0.65$
. (b) Corresponding evolution of bending, line and surface energies. Key time points:
$t_1 = 5.4$
,
$t_2 = 10$
,
$t_3 = 15.5$
,
$t_4 = 17.5$
,
$t_5 = 23.6$
,
$t_6 = 27.4$
,
$t_7 = 30$
,
$t_8 = 37$
,
$t_9 = 39.5$
.

We note that the phase diagram presented in this section is constructed using a symmetric initial distribution of the
$l_d$
phase (see figure 8
b). Although the detailed dynamical trajectories may vary when different initial conditions are used, the qualitative dynamical regimes and their underlying physical mechanisms remain robust. Additional simulations starting from random initial conditions, provided in Appendix E, confirm that the vesicle consistently evolves towards dynamical states that are identifiable as the same regime classes reported here. These results demonstrate that the phase behaviour captured by our model is governed primarily by the interplay between membrane heterogeneity and shear flow, rather than strictly by the particular choice of initial phase distribution.
5. Conclusion
We have developed a comprehensive, thermodynamically consistent phase-field model that accurately captures the intricate interplay among vesicle dynamics, membrane heterogeneity and shear flow. The model successfully reproduces experimental observations of lateral phase separation and shear-induced morphological transitions. This underscores the accuracy, versatility and predictive capability of the model and numerical methods.
Systematic variation of the bending rigidity contrast
$\kappa _B$
and capillary number
${\textit{Ca}}$
reveals six dynamic regimes, including three previously unreported: (I), (II) and (IV). These regimes organize into two critical regions – the ‘low-rigidity full-dynamics belt’ and the ‘low-capillary full-dynamics belt’ – where all six dynamical regimes coexist. Within regime (V), we further identify three subregimes that reflect the rich ring structure of domain redistribution under shear. Beyond bending effects, we uncover surface tension heterogeneity as a previously unknown mechanism that suffices to induce (VII) and (VIII), revealing a new pathway for vesicle dynamics under shear. This unveils a distinct physical mechanism for tuning vesicle dynamics.
These findings offer a comprehensive understanding of the role of membrane heterogeneity in governing vesicle dynamics under shear flow, providing new insights and design principles for tuneable vesicle-based carriers.
Supplementary movies
Supplementary movies are available at https://doi.org/10.1017/jfm.2026.11276.
Funding
S.T. and Z.G. acknowledge support by the National Natural Science Foundation of China (grant no. 12371387). S.M. and J.L. acknowledge partial support by the National Science Foundation of the USA through grants NSF-DMS 2309547 (S.M.) and NSF-DMS 2309800 (J.L.).
Declaration of interests
The authors report no conflict of interest.
Appendix A. Diffuse domain method
We describe the diffuse domain method used to extend membrane dynamics from the sharp interface
$\varGamma$
to the rectangular domain
$\varOmega$
. This extension is achieved through a surface delta function
$\zeta (\phi )$
(2.7), constructed from the phase-field variable
$\phi$
, which is initialized with a hyperbolic tangent profile (2.1) to characterize the diffuse vesicle. The surface delta function
$\zeta (\phi )$
satisfies the following approximation membrane (Teigen et al. Reference Teigen, Song, Lowengrub and Voigt2011):
for any scalar or vector field
$w$
(e.g.
$c_{\varGamma }$
or
$\boldsymbol{q}_c$
). This relation enables the approximation of surface integrals and surface differential operators by volume integrals over
$\varOmega$
, facilitating the extension of surface quantities, equations and energy functionals into
$\varOmega$
.
We now describe the detailed procedure for coupling the surface phase separation dynamics, governed by the surface CH equation on
$\varGamma$
(2.10) to
$\varOmega$
. Using the diffuse domain method (Rätz & Voigt Reference Rätz and Voigt2006; Li et al. Reference Li, Lowengrub, Rätz and Voigt2009; Teigen et al. Reference Teigen, Li, Lowengrub, Wang and Voigt2009), we assume that the membrane component
$c_{\varGamma }$
is smoothly extended off
$ \varGamma$
, remaining constant in the normal direction within a narrow band around the membrane. This yields the normal extension condition:
Additionally, since the diffusive flux
$\boldsymbol{q}_c$
is tangential to
$\varGamma$
, its extension satisfies
Using the incompressibility of the background fluid (2.3) and the surface gradient operator defined in (2.6), we derive the following identity near the vesicle membrane
$\varGamma$
:
Applying the identity (A4) to the surface CH (2.10) yields
Moreover, we further employ the identity from James & Lowengrub (Reference James and Lowengrub2004):
Multiplying (A5) by
$ \zeta (\phi )$
, and (A6) by
$ c_{\varGamma }$
, with the help of the assumptions (A2) and (A3), we obtain
For the flux term, noting that
$\zeta (\phi )$
varies predominantly in the normal direction near
$\varGamma$
with negligible tangential variation (especially for thin interfaces) and applying (A3), we obtain
\begin{align} \boldsymbol{\nabla } \boldsymbol{\cdot }(\zeta (\phi ) \boldsymbol{q_c}) &= \zeta (\phi ) \boldsymbol{\nabla } \boldsymbol{\cdot }\boldsymbol{q_c} + \boldsymbol{\nabla } \zeta (\phi ) \boldsymbol{\cdot }\boldsymbol{q_c} \nonumber\\ &= \zeta (\phi ) \boldsymbol{\nabla } \boldsymbol{\cdot }\boldsymbol{q_c} + \boldsymbol{{\boldsymbol{\nabla} }_{\varGamma }} \zeta (\phi ) \boldsymbol{\cdot }\boldsymbol{q_c} + (\boldsymbol{n} \boldsymbol{\cdot }\boldsymbol{\nabla } \zeta (\phi )) (\boldsymbol{n} \boldsymbol{\cdot }\boldsymbol{q_c}) \nonumber\\ &= \zeta (\phi ) \boldsymbol{\nabla } \boldsymbol{\cdot }\boldsymbol{q_c}. \end{align}
Thus, the resulting surface CH equation in diffuse domain formulation within
$ \varOmega$
can be written as
Notably, previous studies have demonstrated that in the diffuse domain method, the extended PDE (A9) asymptotically converges to the original surface PDE (2.10) as the diffuse interface thickness approaches zero,
$\varepsilon _{\phi } \to 0$
(Li et al. Reference Li, Lowengrub, Rätz and Voigt2009; Yu et al. Reference Yu, Chen and Thornton2012; Lervåg & Lowengrub Reference Lervåg and Lowengrub2015; Chadwick et al. Reference Chadwick, Stewart, Enrique, Du and Thornton2018; Poulsen & Voorhees Reference Poulsen and Voorhees2018; Yu et al. Reference Yu, Guo and Lowengrub2020; Guo et al. Reference Guo, Yu, Lin, Wise and Lowengrub2021; Guo Reference Guo2023).
Appendix B. Model derivation
Next, we derive the free energy dissipation law and the governing equations for the coupled multicomponent vesicle–fluid system. In accordance with the second law of thermodynamics, the free energy
$\mathcal{F}$
(2.13) must satisfy the following dissipation law:
where
${\delta \mathcal{F}}/{\delta \boldsymbol{u}}$
,
${\delta \mathcal{F}}/{\delta \phi }$
and
${\delta \mathcal{F}}/{\delta c_{\varGamma }}$
denote the variational derivatives of the free energy
$\mathcal{F}$
with respect to
$\boldsymbol{u}$
,
$\phi$
and
$c_{\varGamma }$
, respectively.
For the fluid velocity field
$\boldsymbol{u}$
, the variational derivative of the free energy
$\mathcal{F}$
(2.13) with respect to
$\boldsymbol{u}$
is combined with the momentum conservation (2.2), applying integration by parts, we obtain
\begin{align} \int _{\varOmega } \frac {\delta \mathcal{F}}{\delta \boldsymbol{u}} \boldsymbol{u}_t \, \mathrm{d} V &= \int _{\varOmega } \frac {\delta \mathcal{F}_{\textit{K}}}{\delta \boldsymbol{u}}\boldsymbol{u}_t \, \mathrm{d} V = \int _{\varOmega }\rho \boldsymbol{u} \boldsymbol{\cdot }\boldsymbol{u}_t\, \mathrm{d} V \notag\\&= \int _{\varOmega } -\mu | \unicode{x1D63F}(\boldsymbol{u})|^2 + \boldsymbol{F} \boldsymbol{\cdot }\boldsymbol{u} -\xi \varepsilon _\phi ^{2} \phi ^{2}\left |\boldsymbol{\nabla } \lambda _{local}\right |^{2} \, \mathrm{d} V. \end{align}
Similarly, for the phase-field variable
$\phi$
, the variational derivative of
$\mathcal{F}$
with respect to
$\phi$
yields
\begin{align} \frac {\delta \mathcal{F}}{\delta \phi } & = \frac {\delta \mathcal{F}_{\textit{L}}}{\delta \phi } + \frac {\delta \mathcal{F}_{\textit{S}}}{\delta \phi }+\frac {\delta \mathcal{F}_{\textit{B}}}{\delta \phi } + \frac {\delta \mathcal{F}_{\textit{A}}}{\delta \phi } \nonumber \\ &=\left ( \sigma _L \delta (c_{\varGamma }) + {\sigma _S(c_{\varGamma })} + M_{s} \frac {S(\phi )-S(\phi _0)}{S(\phi _0)} \right ) \omega _{\phi } \nonumber \\ &\quad - {\eta _\phi } {\varepsilon _\phi }\boldsymbol{\nabla }\left ( \sigma _L \delta (c_{\varGamma }) + {\sigma _S(c_{\varGamma }) + M_{s} \frac {S(\phi )-S(\phi _0)}{S(\phi _0)}} \right ) \boldsymbol{\cdot }\boldsymbol{\nabla } \phi \nonumber \\ &\quad +{\kappa _B(c_{\varGamma })}{\eta _\phi } \frac {f^{\prime \prime }(\phi )}{{\varepsilon _\phi }^2} \omega _{\phi } - {\eta _\phi } \varDelta \left ( {\kappa _B(c_{\varGamma })} \omega _{\phi }\right ) , \end{align}
where
Combining (B3) with the CH evolution equation for the vesicle membrane (2.4) and applying integration by parts, we obtain
\begin{align} \int _{\varOmega } \frac {\delta \mathcal{F}}{\delta \phi } {\phi }_t \, \mathrm{d} V &=\int _{\varOmega } - \boldsymbol{q_{\phi }} \boldsymbol{\cdot }\boldsymbol{\nabla } \left [\left ( b + \omega _{c} c_{\varGamma } \right ) \omega _{\phi } - {\eta _\phi } {\varepsilon _\phi } \boldsymbol{\nabla }\left ( b + \omega _{c} c_{\varGamma } \right ) \boldsymbol{\cdot }\boldsymbol{\nabla } \phi \vphantom{\frac {f^{\prime \prime }(\phi )}{{\varepsilon _\phi }^2}} \right . \nonumber \\ &\quad \left. +{\kappa _B(c_{\varGamma })}{\eta _\phi } \frac {f^{\prime \prime }(\phi )}{{\varepsilon _\phi }^2} \omega _{\phi } - {\eta _\phi } \varDelta \left ( {\kappa _B(c_{\varGamma })} \omega _{\phi }\right ) \right ] \nonumber \\ &\quad -\boldsymbol{u} \boldsymbol{\cdot }\boldsymbol{\nabla } \phi \left [\left ( b + \omega _{c} c_{\varGamma } \right ) \omega _{\phi } - {\eta _\phi } {\varepsilon _\phi } \boldsymbol{\nabla }\left ( b + \omega _{c} c_{\varGamma } \right ) \boldsymbol{\cdot }\boldsymbol{\nabla } \phi \vphantom{\frac {f^{\prime \prime }(\phi )}{{\varepsilon _\phi }^2}} \right . \nonumber \\ &\quad \left. +{\kappa _B(c_{\varGamma })}{\eta _\phi } \frac {f^{\prime \prime }(\phi )}{{\varepsilon _\phi }^2} \omega _{\phi } - {\eta _\phi } \varDelta \left ( {\kappa _B(c_{\varGamma })} \omega _{\phi }\right ) \right ] \, \mathrm{d} V , \end{align}
where
For the membrane component field
$c_\varGamma$
, multiplying both sides of (2.11) by the chemical potential
$\omega _c$
, defined as the variational derivative of
$\mathcal{F}$
with respect to
$c_\varGamma$
,
\begin{align} \zeta (\phi )\omega _c= \frac {\delta \mathcal{F}}{\delta c_{\varGamma }} & = \frac {\delta \mathcal{F}_{\textit{L}}}{\delta c_{\varGamma }}+\frac {\delta \mathcal{F}_{\textit{S}}}{\delta c_{\varGamma }} + \frac {\delta \mathcal{F}_{\textit{B}}}{\delta c_{\varGamma }} \nonumber\\ &= \frac {{\varepsilon _\phi }^{-1}}{2}{\kappa _B}^{\prime }(c_{\varGamma }) {\omega _{\phi }}^{2} + {\sigma _S}^{\prime }(c_{\varGamma }) {\zeta (\phi )} + \sigma _L \zeta (\phi )\left ({\eta _c} \frac {f^{\prime }(c_{\varGamma })}{\varepsilon _c}-{\eta _c} {\varepsilon _c} \varDelta c_{\varGamma }\right ) \nonumber\\ &\quad - \sigma _L {\eta _c} {\varepsilon _c} \boldsymbol{\nabla } \zeta (\phi ) \boldsymbol{\cdot }\boldsymbol{\nabla }{c_{\varGamma }}, \end{align}
we obtain
Using the definition of the chemical potential
$\omega _c$
(B7), (2.4) and (2.11), which describe membrane evolution and the component dynamics of phase separation, respectively, we derive
\begin{align} \int _{\varOmega } \frac {\delta \mathcal{F}}{\delta c_{\varGamma }} (c_{\varGamma })_t \, \mathrm{d} V & = \int _{\varOmega } {\omega _{c}} \boldsymbol{\nabla } \boldsymbol{\cdot }\boldsymbol{j}-{\omega _{c}} c_{\varGamma } ({ \zeta (\phi )})_t \, \mathrm{d} V \nonumber\\ & = \int _{\varOmega } {\omega _{c}} \boldsymbol{\nabla } \boldsymbol{\cdot }\boldsymbol{j}- {{\phi }_t} \left ({\omega _{c}} c_{\varGamma } {\omega _{\phi }}-\eta _{\phi } {\varepsilon _{\phi }} \boldsymbol{\nabla }\left ({\omega _{c}} c_{\varGamma }\right ) \boldsymbol{\cdot }\boldsymbol{\nabla } \phi \right ) \, \mathrm{d} V \nonumber\\ & = \int _{\varOmega } {{\phi }_t} \left (-{\omega _{c}} c_{\varGamma } {\omega _{\phi }}+\eta _{\phi } {\varepsilon _{\phi }} \boldsymbol{\nabla }\left ({\omega _{c}} c_{\varGamma }\right ) \boldsymbol{\cdot }\boldsymbol{\nabla } \phi \right ) -\zeta (\phi ) \boldsymbol{\nabla } \omega _{c} \boldsymbol{\cdot }\boldsymbol{q_c} \nonumber\\ &\quad - \omega _{c} \boldsymbol{\nabla } (\zeta (\phi ) c_{\varGamma }) \boldsymbol{\cdot }\boldsymbol{u} \, \mathrm{d} V \nonumber\\ & = \int _{\varOmega } - \boldsymbol{q_{\phi }} \boldsymbol{\cdot }\boldsymbol{\nabla } \left (-{\omega _{c}} c_{\varGamma } {\omega _{\phi }}+\eta _{\phi } {\varepsilon _{\phi }} \boldsymbol{\nabla }\left ({\omega _{c}} c_{\varGamma }\right ) \boldsymbol{\cdot }\boldsymbol{\nabla } \phi \right ) -\zeta (\phi ) \boldsymbol{\nabla } \omega _{c} \boldsymbol{\cdot }\boldsymbol{q_c} \nonumber\\ &\quad -\boldsymbol{u} \boldsymbol{\cdot }\left [ \boldsymbol{\nabla } \phi \left (-{\omega _{c}} c_{\varGamma } {\omega _{\phi }}+\eta _{\phi } {\varepsilon _{\phi }} \boldsymbol{\nabla }\left ({\omega _{c}} c_{\varGamma }\right ) \boldsymbol{\cdot }\boldsymbol{\nabla } \phi \right ) { + \omega _{c} \boldsymbol{\nabla } (\zeta (\phi ) c_{\varGamma }) } \right ] \, \mathrm{d} V, \end{align}
where
$\boldsymbol{j}= \zeta (\phi )\boldsymbol{{q_c}} - \zeta (\phi ) c_{\varGamma } \boldsymbol{u}$
.
Finally, by substituting (B2), (B5) and (B9) into the free energy dissipation law (B1), we obtain
\begin{align} \frac {{\rm d} \mathcal{F}}{{\rm d} t}&= \int _{\varOmega } \frac {\delta \mathcal{F}}{\delta \boldsymbol{u}} { \boldsymbol{u}}_{t} + \frac {\delta \mathcal{F}}{\delta \phi } {\phi }_{t}+ \frac {\delta \mathcal{F}}{\delta c_{\varGamma }} (c_{\varGamma })_{t} \, \mathrm{d} V \nonumber\\ &= \int _{\varOmega }-\boldsymbol{q}_{\boldsymbol{\phi }} \boldsymbol{\cdot }\boldsymbol{\nabla }\left [ b\omega _{\phi } + {\kappa _B(c_{\varGamma })}{\eta _\phi } \frac {f^{\prime \prime }(\phi )}{{\varepsilon _\phi }^2} \omega _{\phi } - {\eta _\phi } \varDelta ({\kappa _B(c_{\varGamma })} \omega _{\phi } ) - {\eta _\phi } {\varepsilon _\phi } \boldsymbol{\nabla } b \boldsymbol{\cdot }\boldsymbol{\nabla } \phi \right ] \nonumber\\ &\quad -\boldsymbol{u} \boldsymbol{\cdot }\left [\boldsymbol{ \boldsymbol{\nabla }} \phi \left [ b\omega _{\phi } + {\kappa _B(c_{\varGamma })}{\eta _\phi } \frac {f^{\prime \prime }(\phi )}{{\varepsilon _\phi }^2} \omega _{\phi } \right . \right . \nonumber\\ &\quad - {\eta _\phi } \left . \left . \varDelta ({\kappa _B(c_{\varGamma })} \omega _{\phi } ) - {\eta _\phi } {\varepsilon _\phi } \boldsymbol{\nabla } b \boldsymbol{\cdot }\boldsymbol{\nabla } \phi { + \omega _{c} \boldsymbol{\nabla } (\zeta (\phi ) c_{\varGamma }) }\vphantom{\frac {f^{\prime \prime }(\phi )}{{\varepsilon _\phi }^2}}\right ]\right ] \nonumber\\ &\quad - \mu \left |\boldsymbol{\nabla }\boldsymbol{u}\right |^{2} + \boldsymbol{F} \boldsymbol{\cdot }\boldsymbol{u}-\xi \varepsilon _{\phi }^{2} \phi ^{2}\left |\boldsymbol{\nabla }\lambda _{local}\right |^{2} -\zeta (\phi ) \boldsymbol{\nabla } \omega _{c} \boldsymbol{\cdot }\boldsymbol{q}_{\boldsymbol{c}} \, \mathrm{d} V \leq 0. \end{align}
From these relations, we derive explicit expressions for the previously undetermined terms
$\boldsymbol{F}$
,
$\boldsymbol{q}_\phi$
and
$\boldsymbol{q}_c$
in (2.2)–(2.5) and (2.11),
where
Here, the functions
$m(\phi )=D_\phi (1 - \phi ^2 )^2$
and
$m(c_\varGamma ) = D_c (1 - c_\varGamma ^2 )^2$
denote the mobilities, which are phenomenological parameters with
$D_\phi$
and
$D_c$
being the diffusion coefficients.
By substituting (B11)–(B15) into (B10), we obtain the following free energy dissipation identity:
\begin{align} \frac {{\rm d} \mathcal{F}}{{\rm d} t}= & \int _{\varOmega } \frac {\delta \mathcal{F}}{\delta \boldsymbol{u}} { \boldsymbol{u}}_{t} + \frac {\delta \mathcal{F}}{\delta \phi } {\phi }_{t}+ \frac {\delta \mathcal{F}}{\delta c_{\varGamma }} (c_{\varGamma })_{t} \, \mathrm{d} V \nonumber\\ = & \int _{\varOmega } -\mu \left |\boldsymbol{\nabla }\boldsymbol{u}\right |^{2} -m(\phi )|\boldsymbol{q}_{\boldsymbol{\phi }}|^2 -\zeta (\phi ) m(c_\varGamma )|\boldsymbol{\nabla }\omega _c|^2 -\xi \varepsilon _{\phi }^{2} \phi ^{2}\left |\boldsymbol{\nabla }\lambda _{local}\right |^{2} \leq 0. \end{align}
This expression confirms the thermodynamic consistency of the model, with free energy monotonically decreasing due to viscous dissipation, diffusive fluxes and membrane tension relaxation.
Appendix C. Numerical method
To solve the coupled system numerically, we discretize the time interval
$I = [0, T]$
into uniform time steps
$\varDelta t = t_{n+1} - t_n$
. Time-dependent variables at the
$n$
th and
$(n+1)$
th time steps are denoted as
$(\boldsymbol{\cdot })^n$
and
$(\boldsymbol{\cdot })^{n+1}$
, respectively.
At each time step, the solution procedure consists of the following sequential steps.
-
(i) We first solve the CH (2.19) governing the membrane evolution to update
$\phi ^{n+1}$
,
$g^{n+1}$
and
$\omega _{\phi }^{n+1}$
by the following method: (C1)
\begin{align} &\qquad\qquad\qquad\qquad\quad\frac {\phi ^{n+1} - \phi ^{n}}{\varDelta t} + \boldsymbol{\nabla } \boldsymbol{\cdot }(\phi ^{n} \boldsymbol{u}^{n}) = \frac {1}{\textit{Pe}}\boldsymbol{\nabla } \boldsymbol{\cdot }\boldsymbol{q_{\phi }}^{n+1}, \end{align}
(C2)
\begin{align} &\qquad\qquad\qquad\qquad\qquad\qquad\boldsymbol{q_{\phi }}^{n+1} = m(\phi ^{n}) \boldsymbol{\nabla } g^{n+1}, \end{align}
(C3)
\begin{align} &g^{n+1}= b^{n} \omega _{\phi }^{n+1} + \frac {1}{\textit{Ca}} \!\left (\! {\kappa _B(c_{\varGamma }^{n})} {\eta _\phi } \frac {f^{\prime \prime }(\phi ^{n})}{{\varepsilon _\phi }^{2}} \omega _{\phi }^{n+1}\! - {\eta _\phi } \varDelta \left ({\kappa _B(c_{\varGamma }^{n})} \omega _{\phi }^{n+1} \right ) \!\right ) - {\eta _{\phi }} {\varepsilon _{\phi }} \boldsymbol{\nabla } b^{n}\! \boldsymbol{\cdot }\!\boldsymbol{\nabla } \phi ^{n} , \end{align}
(C4)
\begin{align} &\qquad\qquad b^{n} = -\omega _{c}^{n} c_{\varGamma }^{n} + {{\textit{Cn}}_{\varGamma }} \delta (c_{\varGamma }^{n}) + \mathcal{M}_{\text{s}} \frac {S(\phi ^{n})-S(\phi _0)}{S(\phi _0)} + {Cs_{\varGamma }}{\sigma _S(c_{\varGamma }^{n})} , \end{align}
(C5)
\begin{align} &\qquad\qquad\qquad\qquad\omega _{\phi }^{n+1} = {\eta _{\phi }} \left ( \frac {f^{\prime }({\phi }^{n+1})}{\varepsilon _{\phi }}- {\varepsilon _{\phi }} \varDelta \phi ^{n+1} \right ). \end{align}
The nonlinear terms, such as
$f^{\prime }(\phi ^{n+1})$
and
$f^{\prime \prime }(\phi ^{n+1})$
, are linearized via a first-order Taylor expansion, as described in Lowengrub et al. (Reference Lowengrub, Allard and Aland2016). -
(ii) Next, we solve the CH (2.21) governing the diffusion of membrane components to update
$c_{\varGamma }^{n+1}$
and
$\omega _c^{n+1}$
by the following method: (C6)
\begin{align} \frac { \zeta (\phi ^{n+1}) c_{\varGamma }^{n+1} - \zeta (\phi ^{n}) c_{\varGamma }^{n} }{ \varDelta t } + \boldsymbol{\nabla } \boldsymbol{\cdot }( \zeta (\phi ^{n}) c_{\varGamma }^{n} \boldsymbol{u}^{n} ) = \frac {1}{{\textit{Pe}}_{\varGamma }} \boldsymbol{\nabla } \boldsymbol{\cdot }\left ( \zeta (\phi ^{n+1}) \boldsymbol{q_{c}}^{n+1} \right )\!, \\[-28pt] \nonumber \end{align}
(C7)
\begin{align} \boldsymbol{q_{c}}^{n+1} = m\bigl(c_{\varGamma }^{n}\bigr) \boldsymbol{\nabla } \omega _{c}^{n+1}, \\[-28pt] \nonumber \end{align}
(C8)
\begin{align} \zeta (\phi ^{n+1})\omega _c^{n+1} & =\frac {1}{\textit{Ca}}\frac {{\varepsilon _\phi }^{-1}}{2}{\kappa _B}^{\prime }\bigl(c_{\varGamma }^{n}\bigr) \left ({\omega _{\phi }^{n+1}}\right )^{2} + {Cs_{\varGamma }}{\sigma _S}^{\prime }\bigl(c_{\varGamma }^{n}\bigr) {\zeta \bigl(\phi ^{n+1}\bigr)} \nonumber\\& \quad +{{\textit{Cn}}_{\varGamma }} \zeta \bigl(\phi ^{n+1}\bigr)\!\left (\!{\eta _c} \frac {f^{\prime }\bigl(c_{\varGamma }^{n+1}\bigr)}{\varepsilon _c}-{\eta _c} {\varepsilon _c} \varDelta c_{\varGamma }^{n+1}\!\right ) \nonumber \\ & \quad - {{\textit{Cn}}_{\varGamma }} {\eta _c} {\varepsilon _c} \boldsymbol{\nabla } \zeta \bigl(\phi ^{n+1}\bigr) \!\boldsymbol{\cdot }\!\boldsymbol{\nabla }{c_{\varGamma }^{n}}. \\[0pt] \nonumber \end{align}
Here, the nonlinear term
$f^{\prime }(c_{\varGamma }^{n+1})$
is likewise linearized using a first-order Taylor expansion. -
(iii) Finally, we solve the Navier–Stokes equations (2.17)–(2.18) for the background fluid together with the local inextensibility constraint (2.20) using a projection method, which includes the following substeps.
Step 1. (Temporary velocity prediction) We first solve for the temporary velocity
$\boldsymbol{\tilde {u}}^{n+1}$
,(C9)
\begin{align} &\rho \left ( \frac { \boldsymbol{\tilde {u}}^{n+1} - \boldsymbol{u}^{n} }{ \varDelta t } \right ) + \rho \boldsymbol{\nabla } \boldsymbol{\cdot }\left ( \boldsymbol{u}^{n} \otimes \boldsymbol{u}^{n} \right ) = \frac {1}{\textit{Re}}\boldsymbol{\nabla } \boldsymbol{\cdot }\left ( \mu \bigr(\phi ^{n+1}\bigr) \unicode{x1D63F}\bigl(\boldsymbol{\tilde {u}}^{n+1}\bigr) \right ) - \boldsymbol{\nabla }\! p^{n}\end{align}
(C10)
\begin{align} &\qquad \qquad \qquad \qquad \qquad \qquad \qquad \quad + \frac {1}{\textit{Re}} \boldsymbol{F}^{n} + \boldsymbol{\nabla } \boldsymbol{\cdot }\left ( \zeta \bigl(\phi ^{n+1}\bigr) \unicode{x1D64B}^{n} \lambda _{local}^{n} \right )\!,\end{align}
(C11)
\begin{align} &\boldsymbol{F}^{n} = g^{n+1} \boldsymbol{\nabla } \phi ^{n+1} + {\omega _{c}^{n+1} \boldsymbol{\nabla } \left (\zeta \bigl(\phi ^{n+1}\bigr) c_{\varGamma }^{n+1}\right )} ,\end{align}
where
$\mu (\phi ^{n+1})=(1+\phi ^{n+1})\mu _{{in}}/2+(1-\phi ^{n+1})\mu _{\textit{out}}/2$
. The fields
$\phi ^{n+1}$
,
$g^{n+1}$
,
$c_\varGamma ^{n+1}$
and
$\omega _c^{n+1}$
are obtained from (C1)–(C8).Step 2. (Pressure correction) We then solve for the pressure and then update the velocity field. Having in mind of the incompressibility constraint,
(C12)the velocity correction is given by
\begin{align} \boldsymbol{\nabla } \boldsymbol{\cdot }\boldsymbol{u}^{n+1} = 0, \end{align}
(C13)
\begin{align} \boldsymbol{u}^{n+1} = \boldsymbol{\tilde {u}}^{n+1} - \frac { \varDelta t }{ \rho } \boldsymbol{\nabla } \tilde {p}^{n+1}. \end{align}
By combining (C12) and (C13), the pressure
$\tilde {p}^{n+1}$
is obtained by solving the following Poisson equation:(C14)
\begin{align} {\nabla} ^{2} \tilde {p}^{n+1} = \frac { \rho }{ \varDelta t } \boldsymbol{\nabla } \boldsymbol{\cdot }\boldsymbol{\tilde {u}}^{n+1}. \end{align}
Step 3. (Velocity and pressure update) The velocity
$\boldsymbol{u}^{n+1}$
and pressure
${p}^{n+1}$
are then updated as (C15)
\begin{align} &\boldsymbol{u}^{n+1} = \boldsymbol{\tilde {u}}^{n+1} - \frac { \varDelta t }{ \rho } \boldsymbol{\nabla } \tilde {p}^{n+1}, \end{align}
(C16)
\begin{align} &p^{n+1} = p^{n} + \tilde {p}^{n+1}. \end{align}
Step 4. (Local inextensibility correction) Finally, the Lagrange multiplier
$\lambda _{local}^{n+1}$
is then updated by the following method:(C17)where
\begin{align} \xi \varepsilon _{\phi }^{2} \boldsymbol{\nabla } \boldsymbol{\cdot }\left ( \bigl(\phi ^{n+1}\bigr)^{2} \boldsymbol{\nabla } \lambda _{local}^{n+1} \right ) + \zeta \bigl(\phi ^{n+1}\bigr) \unicode{x1D64B}^{n+1} : \boldsymbol{\nabla } \boldsymbol{u}^{n} = 0; \end{align}
$ \unicode{x1D64B}^{n+1}=\boldsymbol{I} - (\boldsymbol{\nabla }\phi ^{n+1}/|\boldsymbol{\nabla }\phi ^{n+1}|)\otimes (\boldsymbol{\nabla }\phi ^{n+1}/|\boldsymbol{\nabla }\phi ^{n+1}|)$
.
In addition, Dirichlet boundary conditions can imposed on
$\boldsymbol{\tilde {u}}$
, consistent with
$\boldsymbol{u}$
in (2.8), while no-flux boundary conditions are applied to
$\phi$
,
$\boldsymbol{q}_\phi$
,
$c_\varGamma$
and
$\boldsymbol{q}_c$
as in (2.9)–(2.12).
Appendix D. Numerical convergence tests
This appendix presents two numerical convergence studies designed to validate the diffuse-interface formulation and the numerical method described in Appendix C. The first test examines surface diffusion on a stationary spherical vesicle and provides direct quantitative comparison with an analytical solution. The second test evaluates grid-refinement behaviour for fully coupled vesicle dynamics in shear flow. Together, these tests demonstrate that the proposed formulation accurately captures interfacial transport and vesicle hydrodynamics, and that the computational parameters adopted in the main text ensure both accuracy and efficiency.
D.1. Surface-diffusion convergence on a stationary spherical vesicle
To validate the diffuse-interface formulation, we examine the diffusion of a membrane component on a stationary spherical vesicle, following Guo et al. (Reference Guo, Yu, Lin, Wise and Lowengrub2021). The governing equation is
which evaluates the diffuse-domain representation of the membrane and the numerical implementation of the surface diffusion operator.
Under the axisymmetric setting, the surface concentration depends only on the polar angle, i.e.
$c_\varGamma =c_\varGamma (\beta ,t)$
, and is independent of the azimuthal angle. We consider a spherical vesicle of radius
$r_0=1$
. The initial distribution of the membrane component is prescribed as
where
$\beta$
denotes the polar angle measured from the negative
$Z$
-axis, so that
$\cos \beta =-z/r_0$
.
Since the vesicle is stationary (so that
$\phi _t=0$
), (D1) reduces on the spherical surface to the standard axisymmetric surface-diffusion equation
for
$\beta \in [0,\pi ]$
, subject to the initial condition (D2) and the natural symmetry conditions
$\partial _\beta c_{\varGamma }(0,t)=\partial _\beta c_{\varGamma }(\pi ,t)=0$
at the poles. This yields the analytical solution
\begin{equation} c_{\varGamma }(\beta ,t) =\frac {1}{2}\left [1-\exp \!\left (-\frac {2t}{{\textit{Pe}}_{\varGamma }r_0^2}\right )\cos \beta \right ]\!. \end{equation}
To compare the numerical solution with the analytical solution (D4), we solve (D1) on the computational domain
$\bar {\varOmega }=[-1.5,1.5]^3$
. The stationary spherical interface is represented by the phase field
\begin{align} \phi (\boldsymbol{x},0) =\frac {1}{2}\tanh \!\left (\frac {r_0-r(\boldsymbol{x})}{2\sqrt {2}\varepsilon _\phi }\right ) +\frac {1}{2}, \quad r(\boldsymbol{x})=\sqrt {x^2+y^2+z^2}, \quad \boldsymbol{x} =(x,y,z) \in \bar {\varOmega }. \end{align}
Both
$\phi$
and
$c_\varGamma$
are stored at cell centres in the finite-difference discretization, while the membrane interface is defined as the isosurface
$\phi =0.5$
, which does not in general coincide with grid points. To evaluate
$c_\varGamma$
on the membrane, the interface geometry is first extracted using a marching–cubes procedure Teigen et al. (Reference Teigen, Song, Lowengrub and Voigt2011), and the values of
$c_\varGamma$
at the interface vertices are then obtained by trilinear interpolation from the surrounding cell-centred data. The resulting interpolated interface values are compared with the analytical solution at
$t=1$
to compute the
$L_\infty$
errors for various grid sizes and interface thicknesses
$\varepsilon _\phi$
, as reported in table 2. For each refinement level, the time step
$\varDelta t$
is reduced consistently with the mesh size so that temporal errors remain negligible compared with spatial discretization errors. Although the interpolation introduces a small auxiliary error, the results still exhibit clear second-order convergence with mesh refinement and decreasing interface thickness
$\varepsilon _\phi$
.
Convergence of numerical solutions towards the analytical interface component at
$t=1$
for surface diffusion on a stationary vesicle.

For the profile comparisons shown in figure 18,
$c_\varGamma$
is additionally interpolated along a great circle cross-section on the plane
$y=0$
, parametrized by the polar angle
$\beta$
. For visualization, the axisymmetric solution
$c_{\varGamma }(\beta ,t)$
is plotted over
$\beta \in [0,2\pi ]$
by using its
$2\pi$
-periodic extension. Figure 18(a) presents the temporal evolution of
$c_\varGamma$
at
${\textit{Pe}}_{\varGamma }=1$
, demonstrating excellent agreement with the analytical solution. Figure 18(b) compares results at
$t=1$
for
${\textit{Pe}}_{\varGamma }=1,2,4,8,16$
, confirming that the numerical method accurately captures the dependence on
${\textit{Pe}}_{\varGamma }$
. The corresponding diffusion patterns of
$c_\varGamma$
on the membrane for the temporal evolution in figure 18(a) are illustrated in figure 19.
Comparison between numerical (dotted lines) and analytical (solid coloured lines) solutions for surface component diffusion on a stationary spherical vesicle. The grid spacing is
$h=3/128$
with
$\varepsilon _\phi =0.02$
. (a) Temporal evolution at
${\textit{Pe}}_\varGamma =1$
for
$t=0,\,0.2,\,0.4,\,0.6,\,0.8,\,1$
. (b) Profiles at
$t=1$
for
${\textit{Pe}}_\varGamma =1,\,2,\,4,\,8,\,16$
.

Diffusion patterns of the surface component
$c_\varGamma$
on the membrane corresponding to the temporal evolution shown in figure 18(a).

D.2. Convergence of the method and verification of domain parameters
To further validate our numerical method, and to confirm that the computational domain parameters – in particular the grid resolution and time step size – are sufficient to accurately resolve the vesicle dynamics, we consider the evolution of a prepositioned multicomponent vesicle subjected to a shear flow of strength
$\gamma = 1$
. The computational domain is defined as
$\bar {\varOmega } = [-2,2]^3$
. The initial vesicle shape is prescribed using
$d_{1}=1$
and
$d_{2}=d_{3}=0.565$
, corresponding to a reduced volume of
$v \approx 0.91$
. The
$l_d$
phase is presegregated such that the area fraction is
$a_{l_d} = 0.4$
, as illustrated in figure 8(b).
To assess spatial convergence, simulations are performed on four uniform grids of size
$64^3$
,
$96^3$
,
$128^3$
and
$192^3$
. For each grid, the time step sizes
$\varDelta t = 2\times 10^{-4},\;1.5\times 10^{-4},\;1\times 10^{-4},\;7.5\times 10^{-5}$
, respectively, are used. These choices maintain a comparable ratio between spatial and temporal resolutions, ensuring that differences observed in the results primarily reflect spatial accuracy. The dimensionless parameters used in all simulations are:
${\textit{Re}} = 10^{-3}$
,
${\textit{Ca}}=1$
,
${\textit{Cn}}_{\varGamma }=0.05$
,
${\textit{Pe}} = 1000$
,
${\textit{Pe}}_{\varGamma }=1$
,
$Cs_{\varGamma }=10$
,
$\mathcal{M}_S=2000$
,
$\sigma _S = 1$
and
$k_B = 0.5$
.
We monitor three quantities that characterize vesicle dynamics – namely (figure 20
a) the bending energy, (figure 20
b) the line energy, and (figure 20
c) the inclination angle – and compare their evolution up to time
$t=18$
. The simulations on the coarsest grid,
$64^3$
, exhibit noticeable deviations from the finer-grid results across all three diagnostics, indicating insufficient spatial resolution. The
$96^3$
grid reduces but does not fully remove these discrepancies. In contrast, the
$128^3$
and
$192^3$
simulations produce nearly indistinguishable results for all three quantities, demonstrating convergence with respect to spatial refinement.
Since the
$128^3$
and
$192^3$
grids yield quantitatively and qualitatively identical vesicle dynamics, while the coarser grids (
$64^3$
and
$96^3$
) show clear under-resolution effects, we conclude that a grid of size
$128^3$
provides a reliable balance between computational accuracy and efficiency. Accordingly, a resolution of
$128^3$
is adopted for all simulations in this study.
Grid-refinement study for an initially presegregated vesicle (figure 8
b) in shear flow. Shown are the time evolutions of (a) bending energy, (b) line energy and (c) inclination angle for grids of size
$64^3$
,
$96^3$
,
$128^3$
and
$192^3$
. The corresponding time step sizes are
$\varDelta t = 2\times 10^{-4}$
,
$1.5\times 10^{-4}$
,
$1\times 10^{-4}$
and
$7.5\times 10^{-5}$
, respectively.

Appendix E. Dynamical regimes using random initial conditions
To examine the dependence of the dynamical regimes on the initial distribution of membrane components, we performed simulations starting from a random initial distribution of the
$l_d$
phase with the same area fraction
$a_{l_d}=0.4$
, for several parameter combinations spanning regimes (I)–(VIII) in the phase diagrams, including those in figures 14(a) and 17(a). For each parameter set, we compare the time evolution obtained from the symmetric initial condition with that obtained from the random initial condition, using the same time instances as in the corresponding figures. The results for random initial conditions are shown in figures 21 and 22, where we denote the dynamical states by (I’)–(VIII’) to distinguish them from their symmetric counterparts.
Comparison of vesicle dynamical regimes at
$k_{B}=0.65$
for varying
${\textit{Ca}}$
. (a) Dynamical regimes (I)–(VI) obtained from a symmetric initial condition (shown above the panel), corresponding to the classification presented in figure 14 of the original manuscript. (b) Time evolution of the corresponding regimes (I’)–(VI’) obtained under the same
$(k_{B},Ca)$
values but starting from a random initial condition (also shown above the panel).

Comparison of vesicle dynamical regimes at
$k_{B}=0.65$
and
$\sigma _{S}=0.85$
for varying
${\textit{Ca}}$
. (a) Dynamical regimes (VII)–(VIII) obtained from a symmetric initial condition (shown above the panel), corresponding to the classification presented in figure 17(a). (b) Time evolution of the corresponding regimes (VII’)–(VIII’) obtained under the same
$(\sigma _{S},Ca)$
values but starting from a random initial condition (also shown above the panel).

For regimes (I)–(VI) in figure 14(a), the random initial condition generally leads to dynamical behaviours that are qualitatively similar – but not strictly identical – to those in the phase diagram. In particular, (III’) and (IV’) recover the same tank-treading and vertical broken-ring banding behaviours observed in (III) and (IV): both the vesicle shape and the organization of the
$l_d$
domains are very close to those obtained from the symmetric initial condition. By contrast, (I’), (II’), (V’) and (VI’) exhibit behaviours that remain clearly related to (I), (II), (V) and (VI), but with noticeable differences.
For (I’) and (II’), the vesicle no longer adopts a strongly oblate shape, yet the softer
$l_d$
phase still migrates robustly towards the tips and accumulates there. We attribute this to the random initial distribution: because the
$l_d$
phase is initially scattered over the membrane, the domains can reorganize and migrate towards the tips while the vesicle shape adjusts more gradually, without necessarily passing through a strongly oblate configuration to minimize bending energy.
For (V’) and (VI’), the stronger shear accelerates phase separation from the random initial condition, but the
$l_d$
domains do not evolve into two perfectly symmetric caps as in (V) and (VI). As a result, the dynamics do not form an ideal single, slender vertical band (V’), nor a strictly periodic phase-treading motion between two symmetric tips (VI’). Nevertheless, the essential features persist: in (V’), the two
$l_d$
-rich regions still migrate towards the tips and elongate along the flow direction, although they do not merge into a continuous vertical ring; in (VI’), the
$l_d$
phase still travels back and forth between the two tip regions, but with more irregularity due to the asymmetry introduced by the random initial distribution.
For regimes (VII)–(VIII) in figure 17(a), which probe the effect of composition-dependent surface tension under shear, the simulations with random initial conditions likewise exhibit the same categories of dynamical responses. In (VII’), we again observe swinging-like behaviour, though with less regular oscillations due to the asymmetric initial
$l_d$
distribution. In (VIII’), the vesicle displays a tumbling-like motion, but the
$l_d$
phase separates into multiple distinct patches. The lower surface tension in the
$l_d$
regions allows local outward protrusions, creating shape asymmetries that, together with shear, yield a more irregular tumbling cycle. For the particular parameter shown in figure 22(b) (VIII’), the relatively strong shear amplifies the influence of the asymmetric domain pattern. However, within the same parameter region as (VIII) in figure 16(a), we also tested cases with slightly weaker shear, for which more regular tumbling dynamics are recovered even under random initial conditions.
Taken together, these additional simulations demonstrate that the phase diagram in § 4 is not strictly independent of the initial condition, in the sense that different initial distributions may lead to differences in domain rearrangements or symmetry at the same parameter point. However, the qualitative dynamical regimes and the underlying physical mechanisms are robust: starting from a random initial condition primarily affects the detailed morphology of the domains, while the vesicle still evolves towards dynamical states that are clearly recognizable as the same classes reported in the phase diagram (figures 13 a and 16 a). In particular, the key signatures of the interplay among membrane heterogeneity, shear flow and phase separation – such as preferential accumulation of the softer phase in high-curvature regions and shear-induced reorganization of domains – remain unchanged.


















































































































