Chapter 26 : Genetics of Peptidoglycan Biosynthesis

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The central core of the mycobacterial cell envelope consists of the mycolyl-arabinogalactan-peptidoglycan complex, also known as the MAPc. Anchoring the entire MAPc is the peptidoglycan (PG), which is composed of a glycan chain with alternating -acylated glucosamine (GlcNAc) and muramic acid (MurNAcyl) residues bearing peptide chains, which may be cross-linked ( ). The PG of mycobacteria belongs to the A1γ chemotype along with that of and a number of other organisms, but it has modifications to the monomeric units and other structural aspects that are likely related to a role for the PG in stabilizing the mycobacterial MAPc ( ). In this article, we will review the genetics of several aspects of PG biosynthesis in mycobacteria, including the production of monomeric precursors in the cytoplasm, assembly of the monomers into the mature wall, wall turnover, and cell division. Finally, we will touch upon the resistance of mycobacteria to β-lactam antibiotics, an important class of drugs that, until recently, have not been extensively investigated as potential antimycobacterial agents.

Citation: Pavelka M, Mahapatra S, Crick D. 2014. Genetics of Peptidoglycan Biosynthesis, p 513-533. In Hatfull G, Jacobs W (ed), Molecular Genetics of Mycobacteria, Second Edition. ASM Press, Washington, DC. doi: 10.1128/microbiolspec.MGM2-0034-2013
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Image of Figure 1
Figure 1

PG nucleotide precursor (Park's nucleotide). Basic structure of the PG monomer precursor with the muropeptide -alanyl--glutaminyl--DAP--alanyl--alanine. R denotes the presence of either an -acetyl or -glycolyl modification of the muramic acid moiety. -Ala, -Glu, -DAP, and -Ala are depicted in gold, blue, green, and red, respectively.

Citation: Pavelka M, Mahapatra S, Crick D. 2014. Genetics of Peptidoglycan Biosynthesis, p 513-533. In Hatfull G, Jacobs W (ed), Molecular Genetics of Mycobacteria, Second Edition. ASM Press, Washington, DC. doi: 10.1128/microbiolspec.MGM2-0034-2013
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Figure 2

PG cross-links. The direct 4-3 cross-link between -Ala and -DAP. The direct 3-3 cross-link between two -DAP residues. Also shown are various modifications of the PG: R = H or disaccharide linker connecting the PG to the arabinan of the arabinogalactan; R = -acetyl or -glycolyl on the muramic acid residue; R = OH, NH or glycine; R = OH or NH. -Ala, -Glu, -DAP, and -Ala are depicted in gold, blue, green, and red, respectively.

Citation: Pavelka M, Mahapatra S, Crick D. 2014. Genetics of Peptidoglycan Biosynthesis, p 513-533. In Hatfull G, Jacobs W (ed), Molecular Genetics of Mycobacteria, Second Edition. ASM Press, Washington, DC. doi: 10.1128/microbiolspec.MGM2-0034-2013
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Figure 3

Pathways for cytoplasmic steps of PG precursor synthesis.

Citation: Pavelka M, Mahapatra S, Crick D. 2014. Genetics of Peptidoglycan Biosynthesis, p 513-533. In Hatfull G, Jacobs W (ed), Molecular Genetics of Mycobacteria, Second Edition. ASM Press, Washington, DC. doi: 10.1128/microbiolspec.MGM2-0034-2013
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Figure 4

Structure of the mycobacterial Lipid II PG precursor. R = -acetyl or -glycolyl on the muramic acid residue.

Citation: Pavelka M, Mahapatra S, Crick D. 2014. Genetics of Peptidoglycan Biosynthesis, p 513-533. In Hatfull G, Jacobs W (ed), Molecular Genetics of Mycobacteria, Second Edition. ASM Press, Washington, DC. doi: 10.1128/microbiolspec.MGM2-0034-2013
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Figure 5

PG assembly proteins. The various PBPs and ,-transpeptidases are shown as they exist in the genome of . Gene designations from the H37Rv genome: (), (), (), (PBP3, , ), (PBP4, , ), (), (), (, ), (), (), (), (). These genes are also present in , with the exception of and , the latter of which is a pseudogene. and other soil organisms have the novel gene, an extra variant of , and an additional copy of the gene as described in the text. The various domains in each protein are also indicated. Note that PonA2 is unique because it bears a single PASTA domain, which likely binds unlinked PG precursors, and that PonA1, PonA2, LdtC, and LdtE bear extensive proline-rich regions. The class B PBP encoded by Rv2864c and the ,-transpeptidase encoded by are putative lipoproteins.

Citation: Pavelka M, Mahapatra S, Crick D. 2014. Genetics of Peptidoglycan Biosynthesis, p 513-533. In Hatfull G, Jacobs W (ed), Molecular Genetics of Mycobacteria, Second Edition. ASM Press, Washington, DC. doi: 10.1128/microbiolspec.MGM2-0034-2013
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Generic image for table
Table 1

Genes involved with PG turnover

Citation: Pavelka M, Mahapatra S, Crick D. 2014. Genetics of Peptidoglycan Biosynthesis, p 513-533. In Hatfull G, Jacobs W (ed), Molecular Genetics of Mycobacteria, Second Edition. ASM Press, Washington, DC. doi: 10.1128/microbiolspec.MGM2-0034-2013
Generic image for table
Table 2

Genes involved with cell division

Citation: Pavelka M, Mahapatra S, Crick D. 2014. Genetics of Peptidoglycan Biosynthesis, p 513-533. In Hatfull G, Jacobs W (ed), Molecular Genetics of Mycobacteria, Second Edition. ASM Press, Washington, DC. doi: 10.1128/microbiolspec.MGM2-0034-2013
Generic image for table
Table 3

Genes involved with β-lactam antibiotic resistance

Citation: Pavelka M, Mahapatra S, Crick D. 2014. Genetics of Peptidoglycan Biosynthesis, p 513-533. In Hatfull G, Jacobs W (ed), Molecular Genetics of Mycobacteria, Second Edition. ASM Press, Washington, DC. doi: 10.1128/microbiolspec.MGM2-0034-2013

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