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Functional Architecture of Biomembranes
Functional Architecture of Biomembranes
Functional Architecture of Biomembranes
Early microscopists who were contemplating the marvelous diversity of life forms
have noticed a slim line delimiting cells, but actually membranes are not directly
visible under a microscope ~ what we see is a difference in optical density or
refractive index between the interior and exterior of the cell.
In 1900, Ernst Overton, professor of pharmacology at Zrich, found that lipid
soluble compounds show the fastest penetration into cells, water penetrates
more slowly and ions even slower, concluding that the cell membrane contains
lipids. Irwing Langmuir, a renowned physical chemist, professor in New Jersey
and the founder of surface chemistry, describes the structure of lipids as bimodal
molecules ~ amphiphilic or amphipathic ~ composed of a hydrophilic head and a
hydrophobic tail.
In 1925, Grter and Grendel, professors at the University of Leyden, perform a
crucial experiment: they extract the lipids from erythrocyte ghosts, an easily
obtainable preparation, with acetone, and dispose them in a monolayer, following
Langmuirs method. Measuring the surface area of this monolayer, they inferred it
is double compared to the surface of erythrocytes, therefore the logical
conclusion is that the membrane should be a lipid bilayer. Their experiment
contained two mistakes, which fortunately compensated each other: first,
acetone does not extract all lipids, and second, they considered a smaller
erythrocyte surface (100 m2 instead of 140 m2).
In 1936, J.F. Danielli achieves at Buffalo another experiment: he obtained a
lecithin bilayer and measured its surface tension, finding it is approximately one
order of magnitude smaller than the surface tension of cell membranes. By
adding a protein to the bilayer (a solution of ovalbumin), the surface tension
raises and becomes equal to that of the membrane. Therefore, Danielli
developed the sandwich model of cell membrane, adding two protein layers,
one on each side of the lipid bilayer (Fig. 1).