On the Theory of Membrane Fusion. The Stalk Mechanism

Based on literary data, conditions necessary for membrane fusion are discussed. It is proposed that fusion mechanisms should be classified according to the primary act involving a change in the membrane structure. Two principal fusion mechanisms are identified: the stalk mechanism, starting with the appearance of a stalk between approaching membranes, and the adhesion mechanism which involves bilayer reorganization as a result of a tight junction of the membranes. The origin and evolution of the monolayer and bilayer stalks between membranes are analysed. Using the expression for the elastic energy of the stalk it was possible to find the value of the spontaneous curvature of its membrane, Ks, at which the existence of a stalk is in principle possible. It is shown that, within the framework of the stalk mechanism, there exists a possibility of either the formation of a stalk of a finite radius, or complete fusion. The Ks values have been determined at which one of the variants occur. The energy barrier of the hydrophobic interaction and the elastic energy barrier, which have to be overcome by the membranes to form the stalk are analysed. The theoretical analysis of the stalk formation mechanism is supported by experimental data. It has been shown by freeze-fracture electron microscopy that the addition of Ca+2, Mg+2, Mn+2 or Cd+2 to suspensions of egg phosphatidylcholine and cardiolipin (1:1 or 3:1) leads to the formation of numerous intramembrane particles (imp's) and crater-like (stalk) structures.