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The biogenesis of lipid droplets (LD) in the yeast Saccharomyces cerevisiaewas theoretically investigated on basis of a biophysical model. In accordancewith the prevailing model of LD formation, we assumed that neutral lipidsoil-out between the membrane leaflets of the endoplasmic reticulum (ER),resulting in LD that bud-off when a critical size is reached.  Mathematically, LD were modeled as spherical protuberances in an otherwiseplanar ER membrane. We estimated the local phospholipid composition, andcalculated the change in elastic free energy of the membrane caused by nascentLD. Based on this model calculation, we found a gradual demixing of lipids inthe membrane leaflet that goes along with an increase in surface curvature atthe site of LD formation. During demixing, the phospholipid monolayer was ableto gain energy during LD growth, which suggested that the formation of curvedinterfaces was supported by or even driven by lipid demixing. In addition, weshow that demixing is thermodynamically necessary as LD cannot bud-offotherwise.  In the case of Saccharomyces cerevisiae our model predicts a LD bud-offdiameter of about 13 nm. This diameter is far below the experimentallydetermined size of typical yeast LD. Thus, we concluded that if the standardmodel of LD formation is valid, LD biogenesis is a two step process. Small LDare produced from the ER, which subsequently ripe within the cytosol through aseries of fusions.

Phospholipid demixing and the birth of a lipid droplet