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Chapter 41 : Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation

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Abstract:

Several gene products have been implicated in internalization, and the function and regulation of these and other pathogenicity factors in the intracellular environment are discussed in this chapter. Traditionally, bacterial pathogenicity or virulence factors are products whose role in the disease process is either clearly demonstrable, e.g., toxins, or more or less obvious on the basis of biological properties, e.g., enzymes that degrade tissue components. The chapter outlines current understanding of genetics and regulation of staphylococcal virulon. Staphylococcal pathogenesis is multifactorial, involving three classes of factors: secreted proteins, including superantigens (SAgs), cytotoxins, and tissue-degrading enzymes; cell surface-bound proteins, including fibrinogen-binding protein, fibronectin-binding protein, collagen-binding protein, other adhesins, and antiopsonins; and cell surface components, including the polysaccharide capsule and components of the cell wall peptidoglycan. Considering first the genetics of staphylococcal virulence factors, there would appear to be two classes—those encoded by constant chromosomal genes, present in most or all strains, and those encoded by variable genes, present in a minority of strains, and usually belonging to accessory genetic elements, including plasmids, transposons, prophages, and pathogenicity islands (SaPIs), some of which are mobile. The first evidence for virulence gene regulation was the isolation of pleiotropic staphylococcal mutants defective in the production of hemolysins and other virulence factors. A table in the chapter gives a summary of the known genes and environmental conditions that affect the expression of pathogenicity factors by .

Citation: Novick R. 2006. Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation, p 496-516. In Fischetti V, Novick R, Ferretti J, Portnoy D, Rood J (ed), Gram-Positive Pathogens, Second Edition. ASM Press, Washington, DC. doi: 10.1128/9781555816513.ch41

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Image of FIGURE 1
FIGURE 1

() Staphylococcal abscesses. A) Cutaneous furuncle (Nadir Goksugur, M.D., Dermatlas; http://www.dermatlas.org). (B) Stained section of pulmonary abscess. Kindly provided by Martin Nachbar.

Citation: Novick R. 2006. Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation, p 496-516. In Fischetti V, Novick R, Ferretti J, Portnoy D, Rood J (ed), Gram-Positive Pathogens, Second Edition. ASM Press, Washington, DC. doi: 10.1128/9781555816513.ch41
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Image of FIGURE 2
FIGURE 2

Temporal program of virulon expression in vitro.

Citation: Novick R. 2006. Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation, p 496-516. In Fischetti V, Novick R, Ferretti J, Portnoy D, Rood J (ed), Gram-Positive Pathogens, Second Edition. ASM Press, Washington, DC. doi: 10.1128/9781555816513.ch41
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Image of FIGURE 3
FIGURE 3

() , and TCS modules known to affect the virulon. (A) The system. The pro-AIP peptide is processed and secreted by AgrB, binds to an extracellular loop in the receptor-HPK, AgrC, activating autophosphorylation (or dephosphorylation), followed by phosphorylation or dephosphorylation of the response regulator, AgrA. AgrA, in conjunction with SarA, activates the two promoters, P2 and P3, leading to the production of RNAIII. RNAIII controls transcription of the target genes via one or more intracellular regulatory mediators, including a second two-component module, . (B) The locus encodes a receptor-HPK () and a response regulator (), driven by a single promoter and followed by a terminator stem-loop. (C) . The locus encodes a receptor-HPK () and a response regulator (), driven by a single promoter that generates two transcripts whose relative significance is unknown. (Reprinted from reference 94 with the kind permission of Blackwell Publishing, Ltd.)

Citation: Novick R. 2006. Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation, p 496-516. In Fischetti V, Novick R, Ferretti J, Portnoy D, Rood J (ed), Gram-Positive Pathogens, Second Edition. ASM Press, Washington, DC. doi: 10.1128/9781555816513.ch41
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Image of FIGURE 4
FIGURE 4

peptides from various staphylococcal species. Sequences were aligned visually. Predicted AIPs are in bold and are set between spaces. Saur, ; Sarl, ; Sarc, ; Scap, ; Scapr, ; Scarn, ; Sconc, ; Sconu, ; Sepi, ; Sint, ; Slug, ; Ssim, ; Sgal, ; Sxyl, Swar, . Sequence confirmed by in vitro synthesis or mass spectroscopy. Peptide sequence predicted from nucleotide sequence. (Reprinted from reference with the kind permission of Blackwell Publishing, Ltd.)

Citation: Novick R. 2006. Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation, p 496-516. In Fischetti V, Novick R, Ferretti J, Portnoy D, Rood J (ed), Gram-Positive Pathogens, Second Edition. ASM Press, Washington, DC. doi: 10.1128/9781555816513.ch41
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Image of FIGURE 5
FIGURE 5

The system. (A) The locus, about 3.5 kb, contains four open reading frames, P, Q, R, and S R and S form a classical TCS module. The functions of P and Q are unknown. is transcribed from two or three promoters, one of which is active in an -null strain and the other(s) is activated by RNAIII. All three major transcripts, A, B, and C, end at ter. D may be independently transcribed or derived from C by processing. PCR probes used to map the transcripts are shown. (B) Transcription pattern (see text). (Reprinted from reference .)

Citation: Novick R. 2006. Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation, p 496-516. In Fischetti V, Novick R, Ferretti J, Portnoy D, Rood J (ed), Gram-Positive Pathogens, Second Edition. ASM Press, Washington, DC. doi: 10.1128/9781555816513.ch41
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Image of FIGURE 6
FIGURE 6

Sar homologs. Amino acid sequence alignment of Sar homologs, including TcaR and MgaA, against MarR from and other homologous protein sequences identified from the N315 genome by BLASTP analysis. The sequences of SarS (SarH1) and SarH2, which both contain two domains with homology to SarA, were truncated so that only the N-terminal domain of each was included in the alignment. The region containing the predicted helix-turn-helix motif of TcaR (Network Protein Sequence analysis [ ]) and several of the other homologs is highlighted in gray. Strongly conserved residues are indicated in the consensus line and universally conserved residues are in bold type. The arrowhead represents the position (amino acid 79) at which the TcaR protein in NCTC8325-4 is truncated. (Reprinted from reference .)

Citation: Novick R. 2006. Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation, p 496-516. In Fischetti V, Novick R, Ferretti J, Portnoy D, Rood J (ed), Gram-Positive Pathogens, Second Edition. ASM Press, Washington, DC. doi: 10.1128/9781555816513.ch41
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Image of FIGURE 7
FIGURE 7

Regulatory interactions involving SarA and its homologs. Arrows represent up-regulation, bars represent down-regulation. The two outermost curved lines represent translation; the other lines represent interactions that are probably, but not always certainly, transcriptional. The interactions illustrated are based on reviews by Arvidson and Tegmark ( ) and Cheung and Zhang ( ) and on recent papers by Manna and Cheung ( ) and Said-Salim et al. ( ). Although the abbreviations are mostly in italics, on the assumption that the interactions are likely to be at the transcriptional level, there is actually very little evidence to indicate whether they are direct or indirect or at what level they occur. Question marks represent the most speculative. σ is shown entering the system via and , which have σ-dependent promoters and are likely to represent important intermediates in the pathways by which environmental signals are handled. (Reprinted from reference .)

Citation: Novick R. 2006. Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation, p 496-516. In Fischetti V, Novick R, Ferretti J, Portnoy D, Rood J (ed), Gram-Positive Pathogens, Second Edition. ASM Press, Washington, DC. doi: 10.1128/9781555816513.ch41
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Image of FIGURE 8
FIGURE 8

Temporal expression of in vivo. Agr bacteria with plasmid-carried -P3:: fusion. 1.5 × 10 organisms in early exponential phase injected subcutaneously with cytodex beads at time T1, three mice. Imaged with IVIS (Xenogen) system at times indicated. Images are in false color with increasing intensity from dark gray to light gray to white to medium gray (grayscale representations of blue to green to yellow to red). Signal intensity is plotted below (■) along with bacterial counts obtained by sacrificing infected mice, excising and homogenizing the lesion, and plating for viable bacteria.

Citation: Novick R. 2006. Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation, p 496-516. In Fischetti V, Novick R, Ferretti J, Portnoy D, Rood J (ed), Gram-Positive Pathogens, Second Edition. ASM Press, Washington, DC. doi: 10.1128/9781555816513.ch41
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Tables

Generic image for table
TABLE 1

Staphylococcal extracellular accessory proteins

Designation in n315 genome.

Not identified in n315 sequence or absent from n315.

Xp, throughout exponential phase; exp, early exponential phase only; pxp, postexponential phase.

Citation: Novick R. 2006. Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation, p 496-516. In Fischetti V, Novick R, Ferretti J, Portnoy D, Rood J (ed), Gram-Positive Pathogens, Second Edition. ASM Press, Washington, DC. doi: 10.1128/9781555816513.ch41
Generic image for table
TABLE 2

Regulatory genes and their roles

“None” indicates that the gene is conserved among the seven sequenced genomes.

Citation: Novick R. 2006. Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation, p 496-516. In Fischetti V, Novick R, Ferretti J, Portnoy D, Rood J (ed), Gram-Positive Pathogens, Second Edition. ASM Press, Washington, DC. doi: 10.1128/9781555816513.ch41
Generic image for table
TABLE 3

Virulon expression in vivo

Scored by flow cytometry as percent -positive bacteria.

Derivative of NCTC 8325.

Correlated with down-regulation of by CO in vitro ( ).

Enzyme-linked immunosorbent assay.

Citation: Novick R. 2006. Staphylococcal Pathogenesis and Pathogenicity Factors: Genetics and Regulation, p 496-516. In Fischetti V, Novick R, Ferretti J, Portnoy D, Rood J (ed), Gram-Positive Pathogens, Second Edition. ASM Press, Washington, DC. doi: 10.1128/9781555816513.ch41

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