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Bacillus anthracis , a spore-forming pathogen, replicates as chains of vegetative cells by regulating the separation of septal peptidoglycan. Surface (S)-layer proteins andB. anthracis S-layer-associated proteins (BSLs) function as chain length determinants and are assembled in the envelope by binding to the secondary cell wall polysaccharide (SCWP).B. anthracis expresses six different genes encoding LytR-CpsA-Psr (LCP) enzymes (lcpB1 to -4 ,lcpC , andlcpD ), which when expressed inStaphylococcus aureus promote attachment of wall teichoic acid to peptidoglycan. Mutations inB. anthracis lcpB3 andlcpD cause aberrations in cell size and chain length that can be explained as discrete defects in SCWP assembly; however, the function of the otherlcp genes is not known. By deleting combinations oflcp genes from theB. anthracis genome, we generated variants with singlelcp genes.B. anthracis expressinglcpB3 alone displayed physiological cell size, vegetative growth, spore formation, and S-layer assembly. Strains expressinglcpB1 orlcpB4 displayed defects in cell size and shape, S-layer assembly, and spore formation yet sustained vegetative growth. In contrast, thelcpB2 strain was unable to grow unless the gene was expressed from a multicopy plasmid (lcpB2 ++ ), and variants expressinglcpC orlcpD displayed severe defects in growth and cell shape. ThelcpB2 ++ ,lcpC , orlcpD strains supported neither S-layer assembly nor spore formation. We propose a model whereby LCP enzymes fulfill partially overlapping functions in transferring SCWP molecules to discrete sites within the bacterial envelope.IMPORTANCE Products of genes essential for bacterial envelope assembly represent targets for antibiotic development. The LytR-CpsA-Psr (LCP) enzymes tether bactoprenol-linked intermediates of secondary cell wall polymers to the C6 hydroxyl ofN -acetylmuramic acid in peptidoglycan; however, the role of LCPs as a target for antibiotic therapy is not defined. We show here that LCP enzymes are essential for the cell cycle, vegetative growth, and spore formation ofBacillus anthracis , the causative agent of anthrax disease. Furthermore, we assign functions for each of the six LCP enzymes, including cell size and shape, vegetative growth and sporulation, and S-layer and S-layer-associated protein assembly.

Bacillus anthracis lcp Genes Support Vegetative Growth, Envelope Assembly, and Spore Formation