Simple micromechanical methods provide direct measurements of surface cohesion, elasticity, rigidity, and mutual attraction properties of surfactant double-layer membranes in aqueous media. Temperature-dependent tests yield explicit data for thermal phase transitions. Properties of mixtures of phospholipids (neutral and charged), cholesterol, and polypeptides have been studied. The results are essential for evaluation of theories of surfactant mixing, lamellar phase transitions, colloidal stabilization, and flocculation.
IntroductionThe lamellar configuration of biological membrane structures is peculiar to the preferential assembly of amphiphilic molecules into molecular double layers. The strong preference for the lamellar configuration is evidenced by the negligible solubility of these molecules-and extremely slow rate of exchange from a membrane capsule-in aqueous media. In this condensed state, bilayer membranes exhibit solid-or liquidlike material behavior with the common feature of limited surface compressibility (Le. great resistance to change in surface density). Because of the thin structure (~( 3 -4 ) X lo-' cm), bilayer fragility and limited detectability by optical methods create significant obstacles to measurement of the physical-mechanical properties. Recently, micromechanical techniques have been developed to investigate thermal transitions, cohesion, elasticity, and surface rigidity of bilayer membranes in situ. '-3 Bilayers interact nonspecifically via long-range electrostatic, electrodynamic, and solvation These colloidal forces are commonly recognized as van der Waals attraction, electric double-layer repulsion, and hydration repulsion. Other steric and structural interactions exist that are not yet clearly understood. Even though the magnitude of each interaction can be very large, the free energy minimum at stable contact is extremely low. Again because of micromechanical technology, it is possible to directly measure the cumulated free energy potential for assembly (adhesion) of membrane surfaces to stable ~o n t a c t .~,~-' * In this article, we discuss how mixtures of lipids, cholesterol, and polypeptides affect the thermal-mechanical and colloidal interaction properties of bilayer membranes. We outline prominent