| H. Davson and J. Danielli, in 1935, proposed a membrane structure 
              that was refined by Robertson into the "unit-membrane" 
              model, which has served as the basis for membrane structure theories 
              for many years. The unit-membrane model is diagrammed above: a lipid 
              bilayer core coated on either side with proteins. The inside of 
              the membrane in this model is strongly hydrophobic and the outside 
              is polar. Lipid molecules are held together only by hydrophobic interactions, 
              and lipid and protein molecules are held by hydrophobic forces, 
              plus possible charge attractions between lipid heads and side groups 
              on the proteins. No covalent bonds are assumed to exist between 
              molecules, since membranes can be taken apart gently by solvents 
              and then reconstituted in an apparently intact form. The dimensions 
              of the Davson-Danielli unit membrane correspond to those observed 
              in electron micrographs.  |