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sandwich. A slight improvement on this model was proposed by Robertson in his 1957 Unit
Membrane model. His model was based on electron micrographs, the first direct observa-
tions of membrane structure, and featured structures that resembled 'railroad tracks'. In
the 1960s, Benson (for the photosynthetic thylakoid) and Green (for the mitochondrial inner
membrane) proposed lipid bilayer-free models, called the Lipoprotein Subunit model. These
early membrane models did not appreciate membrane asymmetry or dynamics. In 1972
Singer and Nicholson proposed their Fluid Mosaic model that retained the lipid bilayer,
but accounted for previous model shortcomings, recognizing membrane fluidity and lateral
mobility. An extension of this model was provided by Kai Simons' 1997 Lipid Raft model that
proposed sphingolipid- and cholesterol-rich, liquid ordered, signaling microdomains.
The next three chapters will discuss various membrane physical properties. These prop-
erties are essential to understanding how membranes work. Chapter 9 will discuss
membrane thickness, bilayer stability, membrane protein, carbohydrate and lipid asymme-
try, lipid trans-membrane movement (flip-flop), membrane lateral diffusion, membrane-
cytoskeleton interaction, lipid melting behavior, and membrane fluidity.
References
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