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Membrane heterogeneity-driven dynamics of multicomponent vesicles in shear flow

Published online by Cambridge University Press:  05 March 2026

Shuqi Tang
Affiliation:
Beijing Computational Science Research Center , Beijing 100193, PR China
Steven M. Wise
Affiliation:
Department of Mathematics, The University of Tennessee, Knoxville, TN 37996, USA
John S. Lowengrub
Affiliation:
Department of Mathematics, University of California Irvine, Irvine, CA 92697, USA
Zhenlin Guo*
Affiliation:
Beijing Computational Science Research Center , Beijing 100193, PR China
*
Corresponding author: Zhenlin Guo, zguo@csrc.ac.cn

Abstract

Despite its significance in biology and materials science, the dynamics of multicomponent vesicles under shear flow remains poorly understood because of its nonlinear and strongly coupled nature, especially regarding the role of membrane heterogeneity in driving non-equilibrium behaviour. Here we present a thermodynamically consistent phase-field model, which is validated against experiments, for the quantitative investigation of the dynamics. While prior research has primarily focused on viscosity or bending rigidity contrasts, we demonstrate that surface tension heterogeneity can also trigger swinging and tumbling in vesicles under shear. Additionally, our systematic phase diagram reveals three previously unreported dynamical regimes arising from the interplay between bending rigidity heterogeneity and shear flow. Overall, our model provides a robust framework for understanding multicomponent vesicle dynamics, with findings offering new physical insights and design principles for tuneable vesicle-based carriers.

Information

Type
JFM Papers
Copyright
© The Author(s), 2026. Published by Cambridge University Press
Figure 0

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.

Figure 1

Table 1. Physical and dimensionless parameters used in the simulations.

Figure 2

Figure 2. Phase separation on GUVs at $a_{l_d} = 0.3$: experimental snapshots (a) from Wang et al. (2022) and numerical results (b), showing $l_o$ domains (blue) forming within the $l_d$ phase (red).

Figure 3

Figure 3. Phase separation on GUVs at $a_{l_d} = 0.7$: experimental snapshots (a) from Wang et al. (2022) and numerical results (b), showing $l_d$ domains (red) forming within the $l_o$ phase (blue).

Figure 4

Figure 4. 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).

Figure 5

Figure 5. 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.

Figure 6

Figure 6. Experimental images (a) from Wang et al. (2022) 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 $a_{l_o} = 0.64$.

Figure 7

Figure 7. Experimental images (a) from Wang et al. (2022) 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 $a_{l_o} = 0.65$.

Figure 8

Figure 8. (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.

Figure 9

Figure 9. Experimental images (a) from Tusch et al. (2018) 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 $a_{l_d} = 0.7$.

Figure 10

Figure 10. Experimental images (a) from Tusch et al. (2018) 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$.

Figure 11

Figure 11. Experimental images (a) from Tusch et al. (2018) 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$.

Figure 12

Figure 12. 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. (2018).

Figure 13

Figure 13. 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.

Figure 14

Figure 14. (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: $t_1 = 2.6$, $t_2 = 9$, $t_3 = 11.5$, $t_4 = 15$, $t_5 = 17.5$, $t_6 = 18.7$, $t_7 = 25$, $t_8 = 26.7$, $t_9 = 32$.

Figure 15

Figure 15. (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: $t_1 = 8.2$, $t_2 = 11.6$, $t_3 = 14.3$, $t_4 = 20.5$, $t_5 = 25.7$, $t_6 = 28.9$, $t_7 = 43.6$, $t_8 = 62$, $t_9 = 66$.

Figure 16

Figure 16. (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 $\sigma _S$.

Figure 17

Figure 17. (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: $t_1 = 5.4$, $t_2 = 10$, $t_3 = 15.5$, $t_4 = 17.5$, $t_5 = 23.6$, $t_6 = 27.4$, $t_7 = 30$, $t_8 = 37$, $t_9 = 39.5$.

Figure 18

Table 2. Convergence of numerical solutions towards the analytical interface component at $t=1$ for surface diffusion on a stationary vesicle.

Figure 19

Figure 18. 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 ${\textit{Pe}}_\varGamma =1,\,2,\,4,\,8,\,16$.

Figure 20

Figure 19. Diffusion patterns of the surface component $c_\varGamma$ on the membrane corresponding to the temporal evolution shown in figure 18(a).

Figure 21

Figure 20. Grid-refinement study for an initially presegregated vesicle (figure 8b) 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.

Figure 22

Figure 21. 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).

Figure 23

Figure 22. 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).

Supplementary material: File

Tang et al. supplementary movie 1

(I) Lateral ring banding
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Tang et al. supplementary movie 2

(II) Lateral broken-ring banding
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Tang et al. supplementary movie 3

(III) Tank treading
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Tang et al. supplementary movie 4

(VI) Vertical broken-ring banding
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Tang et al. supplementary movie 5

(V) Vertical ring banding
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Tang et al. supplementary movie 6

(VI) Phase treading
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Tang et al. supplementary movie 7

(V_a) Temporary ring
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Tang et al. supplementary movie 8

(V_b) Cyclic broken and closing ring
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Tang et al. supplementary movie 9

(V_c) Stable ring
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Tang et al. supplementary movie 10

(VII) Swinging
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Tang et al. supplementary movie 11

(VIII) Tumbling
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