Chapter 26 : Enterococcal Genetics

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The majority of interest in enterococcal genetics has been generated in response to three landmark discoveries: (i) identification of the first conjugative plasmids whose transfer systems are induced by an identifiable signal, (ii) identification of the first "transposons" capable of intercellular (conjugative) transposition, and (iii) the acquisition of vancomycin resistance. Because the most prevalent vancomycin resistance genes are located on plasmids and transposons, most work on enterococcal genetics has focused on mobile genetic elements. Examination of the complete sequence of a vancomycin-resistant clinical isolate of reaffirmed the importance of such elements in the evolution of this species, revealing that over a quarter of the genome consists of mobile and/or exogenously acquired DNA. However, understanding of the basic mechanisms of DNA replication and repair, chromosomal segregation, cell division, and transcription in this genus remains limited. This chapter provides a review of what is known or can be discerned from the genome sequence about these basic genetic mechanisms. Next, it focuses on the known mobile genetic elements, which seem to play such a significant role in the evolution of the enterococci.

Citation: Weaver K. 2006. Enterococcal Genetics, p 312-331. 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.ch26
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Image of FIGURE 1

() Organization of the VanA and VanB vancomycin resistance systems. Gene functions are color-coded: green, sensory and regulatory; red, essential resistance genes; blue, auxiliary resistance genes. Detailed functions of each gene are described in the text. Green arrows show the position of the VanR-inducible promoters. The percent identity of the VanB genes to their VanA homologs is given below the individual VanB genes. Homologs have identical names in both systems except for the and ligases.

Citation: Weaver K. 2006. Enterococcal Genetics, p 312-331. 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.ch26
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Image of FIGURE 2

() Regulatory circuitry of the pheromone response. (A) Events occurring at the cell surface. Important chromosomally encoded determinants are colored yellow, with the exception of the pheromone itself, which is shown as blue circles. Plasmid-encoded determinants are color-coordinated with their genes shown in B. The inhibitor peptide (ip) is shown as red circles and competitively inhibits pheromone binding to TraC. TraB inhibition of pheromone secretion is depicted here as sequestration, but this is only one possible mechanism and has not been proven. (B) Events occurring at the DNA level. Binding of pheromone by TraA links events at the cell surface with events at the DNA level. A conformational change in TraA due to pheromone binding is indicated by the change in shape of the molecule and change in dimerization state, although the precise changes are still unknown. Pheromone-free TraA binds P0 and inhibits transcription. Direction of transcription from P0 and Pa is indicated by the arrows. The antisense RNA, generalized to aR in this figure, stimulates termination at t1, indicated by the green arrow. Positive regulatory elements vary between different pheromone-responsive plasmids, and their mechanisms of inducing downstream transcription of the conjugation structural genes may also vary. For simplicity, the more common gene names and order are used. It should be noted that the gene order of the and genes is reversed in pAD1. The RepA gene is shown to orient the reader relative to Fig. 3 . (C) Events at the RNA level. Relative levels of RNA produced from the P0 and Pa promoters under uninduced (red) and induced (green) conditions are depicted by the size of the arrows.

Citation: Weaver K. 2006. Enterococcal Genetics, p 312-331. 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.ch26
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Image of FIGURE 3

Genetic organization of a general pheromone-responsive plasmid replicon. Gene names are given as pAD1/pCF10. The stippled box within represents the origin of replication () to which the product binds. The lined boxes at each end of the operon represent the likely centromere-like sites to which the products bind and, with the product, direct plasmid partition. The organization of the region is enlarged below the replicon. Arrowheads at each end of the locus marked P represent the promoters for the RNA I and RNA II transcripts. The extent and direction of transcription of the transcripts are shown above and below the genes for RNA II and RNA I, respectively. The gene is designated by the diagonally lined box and the sequence of the peptide is shown at the bottom. DRa and DRb are direct repeats in the DNA sequence that provide complementarity between RNA I and RNA II when transcribed in opposite directions. Overlapping transcription at the bidirectional transcriptional terminator also provides complementarity.

Citation: Weaver K. 2006. Enterococcal Genetics, p 312-331. 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.ch26
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Image of FIGURE 4

() Model of RNA I-RNA II interaction. RNA I is the larger black structure and RNA II is the smaller blue structure. Stems, loops, and bulges are depicted in their approximate locations and sizes as determined by experimentation. The red stem-loop structure within RNA I sequesters the ribosomebinding site and prevents translation until a complex is formed. The initial interaction occurs at a U-turn motif in the terminator loop of RNA I (A), followed by interaction between the complementary repeats in the 5′ end of each RNA (B and C). Because RNA II-mediated protection from the RNA I-encoded toxin occurs in vivo even when one of the repeats is mutated, the structure shown in panel C is apparently sufficient to revent translation in vivo. Once complex formation is complete (D), the structure is extremely stable in vivo and in vitro, perhaps due to the gap between the interacting repeats.

Citation: Weaver K. 2006. Enterococcal Genetics, p 312-331. 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.ch26
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Image of FIGURE 5

() Genetic organization of Tn and the relative positions of Int and Xis binding sites. (A) Functions of Tn genes are color-coded: blue, recombination; green and red, positive and negative regulation, respectively; yellow, tetracycline resistance; magenta, conjugation. Gene names and open reading frame numbers are shown above the individual genes. The origin of transfer is designated by the line between s and labeled . Promoters and the direction of transcription are designated by labeled arrows below the gene line. (B) Binding sites for the Int-C and Int-N DNA-binding domains and Xis are designated by marked circles and triangles. Int and Xis binding sites are color-coordinated with the genes shown in panel A.

Citation: Weaver K. 2006. Enterococcal Genetics, p 312-331. 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.ch26
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