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The red blood cell (RBC) membrane is composed of a lipid bilayer and acytoskeleton interconnected by protein junction complexes, allowing forpotential sliding between the lipid bilayer and the cytoskeleton. Despite thisbiological reality, it is most often modelled as a single-layer model, ahyperelastic capsule or a fluid vesicle. Another approach involvesincorporating the membrane's composite structure using double layers, where onelayer represents the lipid bilayer and the other represents the cytoskeleton.In this paper, we computationally assess the various modelling strategies byanalysing RBC behaviour in extensional flow and four distinct regimes thatsimulate RBC dynamics in shear flow. The proposed double-layer strategies, suchas the vesicle-capsule and capsule-capsule models, account for the fluidity andsurface incompressibility of the lipid bilayer in different ways. Our findingsdemonstrate that introducing sliding between the layers offers the cytoskeletona considerable degree of freedom to alleviate its elastic stresses, resultingin a significant increase in RBC elongation. Surprisingly, our study revealsthat the membrane modelling strategy for RBCs holds greater importance than thechoice of the cytoskeleton's reference shape. These results highlight theinadequacy of considering mechanical properties alone and emphasise the needfor careful integration of these properties. Furthermore, our findingsfortuitously uncover a novel indicator for determining the appropriatestress-free shape of the cytoskeleton.

Assessment of coupled bilayer-cytoskeleton modelling strategy for red blood cell dynamics in flow