Difference between revisions of "IS Families/ISAs1 family"

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====Original Identification and Distribution====
 
====Original Identification and Distribution====
The founder member of the family, IS1380, was identified in the early 1990s as responsible for the loss of ethanol-oxidizing ability in Acetobacter pasteurianus and is present in a high number of copies<ref><nowiki><pubmed>1657877</pubmed></nowiki></ref>. In the same year, a second member of the family, IS942, was identified upstream of a beta-lactamase, ccrA, gene in Bacteroides fragilis TAL3636<ref><nowiki><pubmed>1650006</pubmed></nowiki></ref> where it was proposed to activate gene expression.
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IS''As1'' was identified in the fish pathogen ''[[wikipedia:Aeromonas_salmonicida|Aeromonas salmonicida]]'' <ref name=":0"><pubmed>PMC40799</pubmed>
  
This family is represented by over 150 members from 100 bacterial species. Members of this family (Fig. IS1380.1) are between 1665 bp (IS1380<ref><nowiki><pubmed>1657877</pubmed></nowiki></ref><ref><nowiki><pubmed>8092854</pubmed></nowiki></ref><ref><nowiki><pubmed>8390818</pubmed></nowiki></ref> and 2071 bp (ISBj1) in length and each carries a single long open reading frame which includes a potential DDE motif. The terminal IRs are related and are approximately 15 bp long. They have conserved ends terminating with Cct/c (Fig. IS1380.1).
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</ref> as insertions into the ''[https://www.wikigenes.org/e/gene/e/1238265.html vapA]'' gene which is responsible for the formation of a virulence factor, the [https://pubmed.ncbi.nlm.nih.gov/1869553/ tetragonal paracrystalline surface protein array] (A-layer). Southern blot analysis showed it to be restricted to 2 [[wikipedia:Aeromonas_salmonicida|''A. salmonicida'']] strains and present in low copy number. The putative [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=ISAs1 IS''As1''] transposase was visualized as a 42,000 molecular weight (M(r)) protein <ref name=":0" />. 
[[Image:Fig. IS1380.1.png|thumb|center|850px|Fig. IS1380.1]]
 
  
====Organization====
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There are nearly 100 entries for this family in ISfinder from over 50 bacterial species. There are currently no archaeal members. [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=ISAs1 IS''As1''] family members are between 1200 and 1326 bp long and generally carry related terminal '''IRs''' of between 14 and 22 bp [[:File:ISAs1.1.png|(Fig.ISAs1.1]]). A single ''orf'' of between 294 and 376 amino acids occupies almost the entire length with between 26 and 50% identity. There are several conserved '''D''' and '''E''' residues but no clear C1 domain. The Tpases of this family include a β-strand insertion domain <ref><pubmed>PMC3107681</pubmed></ref>.
Insertion appears to generate a target duplication of 4 bp although several generate 5 bp DRs. Each member exhibits between 30% and 50% amino acid identity in the potential TpaseAlthough the majority of their Tpases often include the canonical DDE(6)K/R, several members exhibit other residues in place of the K/R. These include DDE(6)Q or DDE(6)I. A subgroup, IS942, composed of 13 members all restricted to Bacteriodetes, include DDE(6)N. None of these differences appear to affect the predicted secondary structure. In addition, to the host-restricted IS942 group, two other branches of the Tpase tree are restricted to the Actinobacteria. A single, poorly characterized NCBI database entry (WP_018034290) probably corresponds to an archaeal IS1380 member and intriguingly, an orf (XP_002337507) with a 100% match to ISLsp5 (Leptospirillum sp) has been identified in the genomic sequence of the black cottonwood tree, Populus trichocarpa. Tpases of this family include and insertion domain with a predominantly -strand secondary structure<ref><nowiki><pubmed>20067338</pubmed></nowiki></ref>.
 
  
For ISBj1 a simple phase change around co-ordinate 1365 results in the extension of the potential Tpase orf from 328 to 454 amino acids (similar to that of the other group members) and significantly increases the quality of the alignment. In addition, ISBj1 also carries a 112 bp duplication in its right end.
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A second, related IS, [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=IS1358 IS''1358''], was identified in ''[[wikipedia:Vibrio|Vibrio]]'' and is linked to [[wikipedia:Lipopolysaccharide#O-antigen|surface layer and O-antigen genes]] <ref name=":0" />. Its putative Tpase of has been visualized using a [[wikipedia:T7_phage|phage T7]] promoter-driven gene <ref name=":2"><pubmed>10708371</pubmed>
  
IS1380 itself<ref><nowiki><pubmed>1657877</pubmed></nowiki></ref> is present in high copy number in the Acetobacter pasteurianus NCI1380 genome and in several strains of acetic acid bacteria. The family contains at least two tIS present in a single copy: ISMsm12 (Mycobacterium smegmatis ; tetR + Methyltransferase) and ISRop1 (Rhodococcus opacus ; reverse transcriptase).  
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</ref> and that of [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=ISAs1 ISAs''1''] has been detected in ''[[wikipedia:Escherichia_coli|Escherichia coli]]'' [[wikipedia:Min_System|minicells]] <ref name=":0" />.  
  
This IS family is distantly related to the eukaryotic PiggyBac TE family (see <ref><nowiki><pubmed>26104701</pubmed></nowiki></ref>) which also includes an insertion domain largely in the form of β-strand<ref><nowiki><pubmed>18076328</pubmed></nowiki></ref>.
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The family also includes ‘‘H-repeats’’ which form part of several so-called RHS ('''R'''earrangement '''H'''ot '''S'''pot) elements containing another repeated sequence, the H-rpt element ('''H'''inc-'''r'''epeat) (see <ref><pubmed>7934896</pubmed></ref> <ref>&lt;nowiki&gt;<pubmed>PMC1206714</pubmed><br /></ref> <ref name=":3"><pubmed>PMC176749</pubmed>
  
====Activation of Neighboring Genes and Transposition ====
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</ref>). [[wikipedia:Escherichia_coli_in_molecular_biology|''Escherichia coli'' K-12]] contains 5 large repetitions of this type ([https://www.uniprot.org/uniprot/P16916 RhsA] to [https://www.uniprot.org/uniprot/P24211 RhsE], scattered around the chromosome with lengths of between 3.7 and 9.6 kb). They represent nearly 1 % of the chromosome and provide homology for [[wikipedia:RecA|RecA]]-dependent rearrangements. These elements are present in many but not all wild-type isolates of ''[[wikipedia:Escherichia_coli|E. coli]]''.
One member of this family, ISEcp1, has proved clinically important in the spread of extended spectrum resistance to beta-lactams. It was originally identified in plasmid plasmid pST01 from an E. coli isolate from the UK (P.D. Stapleton, AJ242809) upstream of a cephalosporinase gene, blaCMY and several lines of evidence suggested that it was often associated with bla genes. It was subsequently identified upstream of a gene encoding an extended spectrum CTX-M type beta-lactamase, blaCTX-M-1 [8], and its relationship to the bla gene was analysed in some detail: although there appeared to be no endogenous blaCTX-M-1 promoter, it was suggested that a promoter in ISEcp1 was capable of driving blaCTX-M-1<ref><nowiki><pubmed>11470367</pubmed></nowiki></ref> as had been proposed for IS942 previously<ref><nowiki><pubmed>1650006</pubmed></nowiki></ref>. Further study showed this to be true and primer extension indicated that a promoter was located within ISEcp1 towards its right end of the IS<ref><nowiki><pubmed>12936998</pubmed></nowiki></ref>. Other CTX-ML-beta-lactamases (including CTXM-1, -M-2, and -M-9)<ref><nowiki><pubmed>11470367</pubmed></nowiki></ref><ref><nowiki><pubmed>12205064</pubmed></nowiki></ref><ref><nowiki><pubmed>14693512</pubmed></nowiki></ref><ref><nowiki><pubmed>11959547</pubmed></nowiki></ref><ref><nowiki><pubmed>11850241</pubmed></nowiki></ref><ref><nowiki><pubmed>15135523</pubmed></nowiki></ref><ref><nowiki><pubmed>12007800</pubmed></nowiki></ref> driven by upstream insertions of ISEcp1 have been identified in various clinical isolates of E. coli and P. mirabilis isolated in Paris<ref><nowiki><pubmed>12007800</pubmed></nowiki></ref> and in Enterobacteriaceae from China<ref><nowiki><pubmed>11850241</pubmed></nowiki></ref> and Poland<ref><nowiki><pubmed>12205064</pubmed></nowiki></ref>.
 
  
Although there appears to be only a single copy of ISEcp1 associated with the blaCTX-M genes rather than two flanking copies constituting a compound transposon, several lines of evidence show that this single copy is involved in blaCTX-M mobilisation.
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The most prominent Rhs component is a giant core orf whose features are suggestive of a [https://www.ncbi.nlm.nih.gov/pmc/articles/PMC98962/#__sec23title cell wall surface ligand-binding protein]. Each Rhs element also contains another repeated sequence, the H-rpt element displaying features of typical insertion sequences (called [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=ISSe1 IS''Se1'']), although no transposition activity has yet been detected.  
  
In a first series of studies<ref><nowiki><pubmed>26104701</pubmed></nowiki></ref> detailed analysis of the DNA region surrounding the blaCTX-M gene a natural plasmid, pILT-3 from K. pneumoniae ILT-3 (Fig. IS1380.2) identified a single ISEcp1 copy upstream containing a potential promoter together with a sequence with some resemblance to an IRR (called IRR1) at some distance downstream. Moreover, the structure was flanked by a 5 bp direct repeat as expected for members of the IS1380 family suggesting that it could be a transposable unit. The transposition capacity of blaCTX-M was assessed<ref><nowiki><pubmed>15616333</pubmed></nowiki></ref> by cloning the region from pILT-3 into a high copy number plasmid in E. coli and using a mating out assay<ref><nowiki><pubmed>6281440</pubmed></nowiki></ref> with the conjugative plasmid pOX38<ref><nowiki><pubmed>6271456</pubmed></nowiki></ref> as a target. Surprisingly a number of different products were isolated, each carrying a the upstream ISEcp1 and its associated blaCTX-M gene but including different ends (Fig. IS1380.3 A and C) but these appear to be true transposition events since they generated 5 bp direct target repeats<ref><nowiki><pubmed>15616333</pubmed></nowiki></ref> (Fig. IS1380.3B). This unexpected result suggests that end recognition (at least at the IRR end) is somewhat relaxed because since the IR sequences show significant variation. 
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For the sake of clarity, the H-rpt DNA sequences B ([https://www.uniprot.org/uniprot/P16917 RhsB]), C1to C3 ([https://www.uniprot.org/uniprot/P16918 RhsC]), E ([https://www.uniprot.org/uniprot/P24211 RhsE]), and min.5 <ref><pubmed>PMC204594</pubmed></ref>, as well as H-rptF <ref name=":3" />, have been renamed [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=ISEc1 ISEc''1''] to [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=ISEc7 ISEc''7''], respectively ([[General Information/What Is an IS?#Characteristics of insertion sequence families|Table 1. Characteristics of insertion sequence families]]) <ref name=":6"><pubmed>PMC98933</pubmed>
[[Image:Fig. IS1380.2.png|thumb|center|850px|Fig. IS1380.2]]
 
[[Image:Fig. IS1380.3.png|thumb|center|850px|Fig. IS1380.3]]
 
  
In an additional set of experiments, it was subsequently shown that ISEcp1B can mobilize a beta-lactamase gene from the chromosome of Klebsiella ascorbata to a plasmid in E.coli<ref><nowiki><pubmed>28416554</pubmed></nowiki></ref>. The study involved a number of steps. A plasmid carrying an ISEcp1 copy was first transformed into a K. ascorbata strain which carries a chromosomal blaCTX-M gene specifying low level resistance to the cephalosporin cefotaxime (MIC 0.06 micrograms/ml). High level cefotaxime resistance colonies (0.5 or 2 microgram/ml) were scored after growth in liquid culture and occurred at 10-7 per viable cell. They were assumed to represent ISEcp1 insertions. Mobilisation of blaCTX-M by ISEcp1 was then assessed by a direct mating out assay<ref><nowiki><pubmed>6281440</pubmed></nowiki></ref> from a Klebsiella ascorbata derivative to E.coli using the conjugative plasmid pOX38Gm<ref><nowiki><pubmed>28416554</pubmed></nowiki></ref>.
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</ref>.
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[[File:ISAs1.1.png|center|thumb|620x620px|'''Fig. ISAs1.1.''' '''General IS''As1'' characteristics, average length, and common ends. Top:''' Distribution of IS length (base pairs) of [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=ISAs1 IS''As1''] family members. The number of examples used in the sample is shown above each column. '''Bottom''': Left ('''IRL''') and right '''IRR''' inverted terminal repeats are shown in [http://weblogo.threeplusone.com/ WebLogo] format.|alt=]]
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====Mechanism====
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Little is known about the transposition properties of this family of elements. However, recent experiments with the ''[[wikipedia:Vibrio_cholerae|Vibrio cholera]]'' element [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=IS1358 IS''1358''] and [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=ISPG2 IS''PG2''] have demonstrated that insertion generates 10-bp <ref name=":6" /> or 8-bp <ref name=":0" /> '''DRs'''.
  
More recently, to study the increasing number of the relatively high copy number ColE1-type plasmids which are now beginning to be identified having acquired antibiotic resistance genes, ISEcp1-mediated transposition has been monitored from the chromosome of clinical E. coli isolates<ref><nowiki><pubmed>30087569</pubmed></nowiki></ref>. A number of these were observed to carry a collection of plasmids including ColE1 derivatives. None of the strains were able to transfer the resistances by conjugation but three could transfer cefoxitin resistance and the blaCMY-2 gene to a suitable E. coli recipient at very low frequencies (10-8 –10-9) when plasmid DNA was introduced into a suitable recipient by electroporation. In these cases, resistance levels were significantly higher than in the original host. A second round of electroporation of plasmid DNA isolated from resistant colonies gave high frequencies of resistance resulting from an insertion from the chromosome into into the endogenous plasmid pSC137. Sequence analysis of several plasmid derivatives revealed different insertion events using a the ISECP1 IRL and a surrogate downstream end all flanked by a 5 bp direct target duplication<ref><nowiki><pubmed>30087569</pubmed></nowiki></ref>. The association of ISEcp1 with beta-lactamases in clinical isolated appears to be a frequent events<ref><nowiki><pubmed>19596889</pubmed></nowiki></ref>. The association of ISEcp1 with beta-lactamases in clinical isolated appears to be a frequent events<ref><nowiki><pubmed>19596889</pubmed></nowiki></ref>. This is not limited to antibiotic resistance genes.
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It is interesting to note that several members of the family ([https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=IS1358 IS''1358''] ''<ref name=":1"><pubmed>PMC40799</pubmed>
  
Although little is known about most other IS1380 family members, the ability to enhance expression of downstream genes may be a general characteristic. It was previously shown that IS1247 can activate gene expression and appears to participate in a similar type of transposition as ISEcp1<ref><nowiki><pubmed>7868610</pubmed></nowiki></ref>. In Xanthobacter autotrophicus GJ10 it can increase resistance to bromoacetate by insertion upstream and activation of the haloacetate dehalogenase gene and mobilizes the gene by transposition using a surrogate sequence resembling an IR downstream. More recently, another IS1380 family member, ISPme1 from Paracoccus methylutens<ref><nowiki><pubmed>18296518</pubmed></nowiki></ref>, which also uses a surrogate downstream IR-like sequence has been identified. Examples are shown in (Fig. IS1380.4A and B). These elements have been named TMO (ISX) for Transposition Modules where ISX is the name of the IS involved. Like ISEcp1, there is an internal promoter downstream of the transposase gene which is oriented downstream<ref><nowiki><pubmed>18296518</pubmed></nowiki></ref>. In addition, another Paracoccus IS1380 family member, an IS1247 derivative from Paracoccus marcusii was identified as part of a mosaic transposable element. It appears that a TMO(IS1247a) (Fig. IS1380.4C) had inserted into a member of the ISAs1 IS family, ISPmar4<ref><nowiki><pubmed>19187199</pubmed></nowiki></ref>. TMO(IS1247a) has flanking 5 bp DR in this transposition unit. It was suggested that the internal outwardly facing IS1247 promoter could activate the downstream putative alpha/beta hydrolase gene<ref><nowiki><pubmed>19187199</pubmed></nowiki></ref>.
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</ref>''<ref name=":2" /><ref name=":4"><pubmed>PMC98933</pubmed>
[[Image:Fig. IS1380.4.png|thumb|center|850px|Fig. IS1380.4]]
 
  
Although no details of the IS1380 transposition mechanism are known, these observations suggest that, like other IS such as IS91 (and potentially the ISCR although no formal proof has yet been presented) are capable of “capturing” and driving transposition of neighboring, downstream genes.
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</ref>, [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=ISAs1 ISAs''1''] <ref name=":0" /> and [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=ISSe1 IS''Se1''] <ref><pubmed>PMC205649</pubmed></ref>) are associated with cell surface component genes in their respective hosts, although this may reflect the interests of the researchers rather than reflecting a specific association.
<br/>
 
  
 +
In ''[[wikipedia:Streptococcus_agalactiae|Streptococcus agalactiae]]'', [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=IS1548 IS''1548''], is also associated with virulence modulation <ref><pubmed>24760965</pubmed></ref> and has been linked to 19 strains associated with [[wikipedia:Neonatal_meningitis|neonatal meningitis]] and [[wikipedia:Endocarditis|endocarditis]] <ref name=":5"><pubmed>31476407</pubmed>
  
<b>Bibliography</b>
+
</ref>. It has been reported to impact ''[[wikipedia:Streptococcus_agalactiae|S. agalactiae]]'' in two ways: by inactivating virulence genes such as ''[[wikipedia:Hyaluronidase|hylB]]'' (hyaluronidase ) or ''[https://www.uniprot.org/uniprot/P0C0T9 cpsD]'' by insertion and by activating the expression of downstream genes such as ''[https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2737970/ lmb]'' (laminin/Zn-binding protein gene) and ''murB'' ([[wikipedia:Peptidoglycan#Biosynthesis|peptidoglycan biosynthesis]]) <ref name=":5" /><ref><pubmed>PMC2875397</pubmed></ref>.
<references/>
+
 
<br/>
+
Another member, [https://tncentral.ncc.unesp.br/ISfinder/scripts/ficheIS.php?name=ISPmar4 IS''Pmar4''], has been implicated in driving transposition of a mosaic element in ''[[wikipedia:Paracoccus|Paracoccus]]''  <ref><pubmed>19187199</pubmed></ref>.
<br/>
+
<br />
 +
==Bibliography==
 +
<references />

Latest revision as of 00:02, 13 August 2021

Original Identification and Distribution

ISAs1 was identified in the fish pathogen Aeromonas salmonicida [1] as insertions into the vapA gene which is responsible for the formation of a virulence factor, the tetragonal paracrystalline surface protein array (A-layer). Southern blot analysis showed it to be restricted to 2 A. salmonicida strains and present in low copy number. The putative ISAs1 transposase was visualized as a 42,000 molecular weight (M(r)) protein [1].

There are nearly 100 entries for this family in ISfinder from over 50 bacterial species. There are currently no archaeal members. ISAs1 family members are between 1200 and 1326 bp long and generally carry related terminal IRs of between 14 and 22 bp (Fig.ISAs1.1). A single orf of between 294 and 376 amino acids occupies almost the entire length with between 26 and 50% identity. There are several conserved D and E residues but no clear C1 domain. The Tpases of this family include a β-strand insertion domain [2].

A second, related IS, IS1358, was identified in Vibrio and is linked to surface layer and O-antigen genes [1]. Its putative Tpase of has been visualized using a phage T7 promoter-driven gene [3] and that of ISAs1 has been detected in Escherichia coli minicells [1].

The family also includes ‘‘H-repeats’’ which form part of several so-called RHS (Rearrangement Hot Spot) elements containing another repeated sequence, the H-rpt element (Hinc-repeat) (see [4] [5] [6]). Escherichia coli K-12 contains 5 large repetitions of this type (RhsA to RhsE, scattered around the chromosome with lengths of between 3.7 and 9.6 kb). They represent nearly 1 % of the chromosome and provide homology for RecA-dependent rearrangements. These elements are present in many but not all wild-type isolates of E. coli.

The most prominent Rhs component is a giant core orf whose features are suggestive of a cell wall surface ligand-binding protein. Each Rhs element also contains another repeated sequence, the H-rpt element displaying features of typical insertion sequences (called ISSe1), although no transposition activity has yet been detected.

For the sake of clarity, the H-rpt DNA sequences B (RhsB), C1to C3 (RhsC), E (RhsE), and min.5 [7], as well as H-rptF [6], have been renamed ISEc1 to ISEc7, respectively (Table 1. Characteristics of insertion sequence families) [8].

Fig. ISAs1.1. General ISAs1 characteristics, average length, and common ends. Top: Distribution of IS length (base pairs) of ISAs1 family members. The number of examples used in the sample is shown above each column. Bottom: Left (IRL) and right IRR inverted terminal repeats are shown in WebLogo format.

Mechanism

Little is known about the transposition properties of this family of elements. However, recent experiments with the Vibrio cholera element IS1358 and ISPG2 have demonstrated that insertion generates 10-bp [8] or 8-bp [1] DRs.

It is interesting to note that several members of the family (IS1358 [9][3][10], ISAs1 [1] and ISSe1 [11]) are associated with cell surface component genes in their respective hosts, although this may reflect the interests of the researchers rather than reflecting a specific association.

In Streptococcus agalactiae, IS1548, is also associated with virulence modulation [12] and has been linked to 19 strains associated with neonatal meningitis and endocarditis [13]. It has been reported to impact S. agalactiae in two ways: by inactivating virulence genes such as hylB (hyaluronidase ) or cpsD by insertion and by activating the expression of downstream genes such as lmb (laminin/Zn-binding protein gene) and murB (peptidoglycan biosynthesis) [13][14].

Another member, ISPmar4, has been implicated in driving transposition of a mosaic element in Paracoccus [15].

Bibliography

  1. 1.0 1.1 1.2 1.3 1.4 1.5 Stroeher UH, Jedani KE, Dredge BK, Morona R, Brown MH, Karageorgos LE, Albert MJ, Manning PA . Genetic rearrangements in the rfb regions of Vibrio cholerae O1 and O139. - Proc Natl Acad Sci U S A: 1995 Oct 24, 92(22);10374-8 [PubMed:7479787] [DOI]
  2. Hickman AB, Chandler M, Dyda F . Integrating prokaryotes and eukaryotes: DNA transposases in light of structure. - Crit Rev Biochem Mol Biol: 2010 Feb, 45(1);50-69 [PubMed:20067338] [DOI]
  3. 3.0 3.1 Jedani KE, Stroeher UH, Manning PA . Distribution of IS1358 and linkage to rfb-related genes in Vibrio anguillarum. - Microbiology: 2000 Feb, 146 ( Pt 2);323-331 [PubMed:10708371] [DOI]
  4. Hill CW, Sandt CH, Vlazny DA . Rhs elements of Escherichia coli: a family of genetic composites each encoding a large mosaic protein. - Mol Microbiol: 1994 Jun, 12(6);865-71 [PubMed:7934896] [DOI]
  5. <nowiki> Hill CW, Feulner G, Brody MS, Zhao S, Sadosky AB, Sandt CH . Correlation of Rhs elements with Escherichia coli population structure. - Genetics: 1995 Sep, 141(1);15-24 [PubMed:8536964]
  6. 6.0 6.1 Zhao S, Hill CW . Reshuffling of Rhs components to create a new element. - J Bacteriol: 1995 Mar, 177(5);1393-8 [PubMed:7868617] [DOI]
  7. Zhao S, Sandt CH, Feulner G, Vlazny DA, Gray JA, Hill CW . Rhs elements of Escherichia coli K-12: complex composites of shared and unique components that have different evolutionary histories. - J Bacteriol: 1993 May, 175(10);2799-808 [PubMed:8387990] [DOI]
  8. 8.0 8.1 Mahillon J, Chandler M . Insertion sequences. - Microbiol Mol Biol Rev: 1998 Sep, 62(3);725-74 [PubMed:9729608]
  9. Stroeher UH, Jedani KE, Dredge BK, Morona R, Brown MH, Karageorgos LE, Albert MJ, Manning PA . Genetic rearrangements in the rfb regions of Vibrio cholerae O1 and O139. - Proc Natl Acad Sci U S A: 1995 Oct 24, 92(22);10374-8 [PubMed:7479787] [DOI]
  10. Mahillon J, Chandler M . Insertion sequences. - Microbiol Mol Biol Rev: 1998 Sep, 62(3);725-74 [PubMed:9729608]
  11. Xiang SH, Hobbs M, Reeves PR . Molecular analysis of the rfb gene cluster of a group D2 Salmonella enterica strain: evidence for its origin from an insertion sequence-mediated recombination event between group E and D1 strains. - J Bacteriol: 1994 Jul, 176(14);4357-65 [PubMed:8021222] [DOI]
  12. Fléchard M, Gilot P . Physiological impact of transposable elements encoding DDE transposases in the environmental adaptation of Streptococcus agalactiae. - Microbiology: 2014 Jul, 160(Pt 7);1298-1315 [PubMed:24760965] [DOI]
  13. 13.0 13.1 Khazaal S, Al Safadi R, Osman D, Hiron A, Gilot P . Dual and divergent transcriptional impact of IS1548 insertion upstream of the peptidoglycan biosynthesis gene murB of Streptococcus agalactiae. - Gene: 2019 Dec 15, 720;144094 [PubMed:31476407] [DOI]
  14. Al Safadi R, Amor S, Hery-Arnaud G, Spellerberg B, Lanotte P, Mereghetti L, Gannier F, Quentin R, Rosenau A . Enhanced expression of lmb gene encoding laminin-binding protein in Streptococcus agalactiae strains harboring IS1548 in scpB-lmb intergenic region. - PLoS One: 2010 May 24, 5(5);e10794 [PubMed:20520730] [DOI]
  15. Szuplewska M, Bartosik D . Identification of a mosaic transposable element of Paracoccus marcusii composed of insertion sequence ISPmar4 (ISAs1 family) and an IS1247a-driven transposable module (TMo). - FEMS Microbiol Lett: 2009 Mar, 292(2);216-21 [PubMed:19187199] [DOI]