Murein structure

Bacitracin contains a cyclic peptide as part of a complex structure of 12 amino acid residues (if one considers the thiazole ring as cysteine derived). Bacitracin-treated bacterial cultures accumulate precursor peptidoglycan chains, as do the penicillins. However, the mechanism is very different. Its prevention of lysine inclusion into the murein structure is probably of lesser significance. Later demonstration show that drug binding to the membrane-bound bactoprene phosphate and the metal ions (Zn2+ are particularly involved) is more likely to be important to the antibacterial mechanism, namely, inhibition of dephosphorylation of the phospholipid carrier. 

Braun, V. and Rehn, K. (1969). Chemical characterization, spatial distribution and function of a lipoprotein (murein-lipoprotein) of the E. coli cell wall. The specific effect of trypsin on the membrane structure, Eur. J. Biochem., 10, 426-438. 

The (3-lactam antibiotics structurally resemble the terminal D-alanyl-D-alanine (o-Ala-o-Ala) in the pen-tapeptides on peptidoglycan (murein) (Fig. 45.1). Bacterial transpeptidases covalently bind the (3-lactam antibiotics at the enzyme active site, and the resultant acyl enzyme molecule is stable and inactive. The intact (3-lactam ring is required for antibiotic action. The (3-lactam ring modifies the active serine site on transpeptidases and blocks further enzyme function. 

The complex structure of bacterial cell walls is discussed in Chapter 8. However, it is appropriate to mention a few bacterial polysaccharides here. The innermost layer of bacterial cell walls is a porous network of a highly crosslinked material known as pepti-doglycan or murein (see Fig. 8-29). The backbone of the peptidoglycan is a P-l,4-linked... 

Figure 8-29 (A) Repeating unit of structure of a bacterial peptidoglycan (murein). Some connecting bridges are pentaglycine (Staphylococcus aureus), trialanylthreonine (Micrococcus roseum), and polyserine (S. epidermis).

Principles to stabilize lipid bilayers by polymerization have been outlined schematically in Fig. 4a-d. Mother Nature — unfamiliar with the radically initiated polymerization of unsaturated compounds — uses other methods to-stabilize biomembranes. Polypeptides and polysaccharide derivatives act as a type of net which supports the biomembrane. Typical examples are spectrin, located at the inner surface of the erythrocyte membrane, clathrin, which is the major constituent of the coat structure in coated vesicles, and murein (peptidoglycan) a macromolecule coating the bacterial membrane as a component of the cell wall. Is it possible to mimic Nature and stabilize synthetic lipid bilayers by coating the liposome with a polymeric network without any covalent linkage between the vesicle and the polymer One can imagine different ways for the coating of liposomes with a polymer. This is illustrated below in Fig. 53. 

Polysaccharides and molecules whose structures contain polysaccharide residues have a wide variety of biochemical roles. They occur as integral parts of the structures of specific tissues the mureins, for example, (Figure 1.21(a)) are... 

Cell envelopes of archaea diifer distinctly from those of bacteria and show remarkable structural and chemical diversity. Murein, the typical sacculus-forming polymer of bacteria, and lipopolysaccharide-containing outer membranes, characteristic of gramnegative bacteria, are not found in archaea. Crystalline surface layers (S-layers) are common in both prokaryotic domains and they consist of protein or glycoprotein subunits (Table 1). However, S-layers in archaea have a form-stabilizing function especially when they are the only envelope layer outside the cytoplasmic membrane, while in bacteria S-layers have no distinct form-stabilizing function. 

Fig. 2. Primary structure of dimers of murein and pseudomurein (compounds in parentheses may be missing in some cases). From ref. [46].

X-ray diffraction measurements and structural calculations on murein [22-25] and pseudomurein [26-28] have revealed several common structural features in both polymers. Murein and pseudomurein sacculi possess a density of p= 1. 39-1.46g/cm which is characteristic of highly ordered material. A much lower density, in the range of p= 1.24-1.32g/cm is to be expected for amorphous polymers [26]. X-ray diffraction showed diffuse Debye-Scherer rings with Bragg periodicities of about 0.45 nm and 0.94 nm in the planes and of 4.3-4.5nm vertically to the planes of both types of cell walls. These data have been interpreted in two different ways ... 

The first steps of murein biosynthesis take place in the cytoplasm, whereas the two final steps occur at the inner and outer face of the cytoplasmic membrane, respectively [38]. This may also be true for the pseudomurein, according to a tentative scheme of the biosynthesis of pseudomurein (Fig. 4) [39-43], proposed on the basis of the structure of putative precursors isolated from cell extracts. 

The lack of a murein cell-wall sacculus and the discovery of different cell-envelope polymers and structures in some physiologically unusual prokaryotes, was one of the first biochemical and cytological evidences in favour of Carl Woese s archaebacteria concept [46,149,150]. Since then, increasingly more unique cell-envelope polymers and new types of biosynthetic pathways have become known. These findings corroborate the proposal that the archaea represent a third lineage of organisms [150] in addition to bacteria and eucarya, and that the common ancestor or ancestral population of the archaea did not evolve any cell-wall polymer before it radiated into the various sublineages known today [46,151]. 

A recent study [30] has shown that the murein consists of a mac-romolecular bag-shaped network of covalently linked repeating units. Metal ions do not play a role in conserving its structure and the layer is covalently linked to protein. 

Haworth structures of linked segments of cellulose, chitin, chitosan, murein, xylo-/ -glucan, and... 

A large number of bacterial polysaccharides are known [104]. The major structural component of the bacterial cell wall is a polysaccharide, known as murein and composed of a repeating unit of one A-acetyl-D-glucosamine and an O-lactyl substituted IV-acetyl-D-glucosamine (A-acetyl-D-muramic acid) see Sect. 7.3. 

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