FoxLipopolysaccharide (LPS) is a major component of the outer membrane of gram-negative bacteria. The diverse O antigen portion is an important determinant of the surface character of the cells, including their outer membrane permeability (38), electrical charge (44), reactivity to antibodies, and susceptibility to phages (32), and so affects the adaptation of bacteria to their environments. LPSs of many bacteria also show potent endotoxin activity (22). In cyanobacteria, LPS may adsorb cyanophage (42), and its endotoxin activity may affect aquatic habitats (25).LPS consists of three covalently linked domains: lipid A, oligosaccharide core, and O antigen. The O antigen portion is comprised of serially repeated, strain-specific oligosaccharide units. This polymer of oligosaccharide units is the immunodominant portion of the LPS and often serves as a bacteriophage receptor. Assembly of the lipid A-core portion and synthesis of the O antigen units are directed by the rfa and rfb genes, respectively. In a "block" mechanism of O antigen synthesis, repeating subunits of the O antigen, synthesized by sequential transfer of monosaccharide constituents to a lipid carrier at the cytosolic side of the cytoplasmic membrane, are transported to the periplasmic side of the cell membrane and polymerized. The polymerization, directed by rfc genes, entails transfer of an already synthesized unit or polymer of units to the reducing terminus of a single lipid-linked unit (34,45). Assembly of LPS is completed by ligation of an entire polysaccharide chain to oligosaccharide core-lipid A. Finally, LPS is translocated to the outer membrane. All of the enzymes, lipid carriers, intermediates, and LPS itself are membrane bound. In this mechanism for LPS synthesis, the first monosaccharide residue transferred to a lipid carrier is galactose-1-phosphate. The gene, rfbP, responsible for the transfer of that galactosyl residue is indicative of the block mechanism of LPS synthesis. An alternative, monomeric mechanism of O antigen synthesis proceeds at the cytoplasmic membrane by sequential addition of mannosyl residues to the nonreducing terminus of the forming chain and leads to the synthesis of an LPS in which the repeating subunit of O antigen contains three or five interlinked mannosyl residues (22,45).Cyanobacteria are a group of oxygenic, photosynthetic gram-negative bacteria. Although the wall of cyanobacterial vegetative cells has been incompletely defined, chemically, LPS is typically a constituent and contains domains that are similar to those in other bacteria (14). The LPS of the unicellular species Anacystis nidulans KM has been studied in detail (24). Its O antigen, reminiscent of the O antigen of Escherichia coli O8 and O9, contains 1,4-interlinked trisaccharides of 1,3-linked mannosyl residues. As in other phototrophic bacteria, some of the mannosyl residues are methylated. The O antigen of a filamentous species, Anabaena variabilis, contains L-acofriose, L-rhamnose, D-mannose, D-glucose, and D-galactose (50), suggestive of a b...