The Lipid A moiety of lipopolysaccharide forms the outer monolayer of the outer membrane in most Gram-negative bacteria. Escherichia coli lipid A is synthesized on the cytoplasmic side of the inner membrane by a conserved pathway of nine constitutive enzymes. Following attachment of the core oligosaccharide, nascent core-lipid A is flipped to the outer surface of the inner membrane by the ABC transporter MsbA, where the O-antigen polymer is attached. These complex glycoforms protect bacteria from antibiotics and complement-mediated lysis. The core and O-antigen domains are required for virulence and consequently are present in most clinical and environmental isolates (Raetz, 2007).
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