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The membrane tubulation induced by banana-shaped protein rods is investigatedby using coarse-grained meshless membrane simulations. It is found that thetubulation is promoted by laterally isotropic membrane inclusions that generatethe same sign of spontaneous curvature as the adsorbed protein rods. Theinclusions are concentrated in the tubules and reduce the bending energy of thetip of the tubules. On the other hand, the inclusions with an oppositecurvature suppress the tubulation by percolated-network formation at a highprotein-rod density while they induce a spherical membrane bud at a low roddensity. When equal amounts of the two types of inclusions (with positive andnegative curvatures) are added, their effects cancel each other for the firstshort period but later the tubulation is slowly accelerated. A positive surfacetension suppresses the tubulation. Out results suggest that the cooperation ofscaffolding of BAR (Bin/Amphiphysin/Rvs) domains and isotropic membraneinclusions is important for the tubulation.

Acceleration and suppression of banana-shaped-protein-induced tubulation by addition of small membrane inclusions of isotropic spontaneous curvatures