IS and Gene Expression
Another important aspect of IS impact on their
bacterial hosts is their ability to modulate gene expression. In addition to acting as vectors for gene
transmission from one replicon to another in the form composite transposons
(two IS flanking any gene; Fig 1.2.3) and tIS (Fig 1.13.1) and their ability to interrupt
genes, it has been known for some time (Reif
& Saedler, 1974, Glansdorff, et al.,
1981) that IS can also activate gene expression. This capacity has
recently received much attention due to the increase in resistance to various
antibacterials (Aubert, et al., 2006, Soki, et al.,
2013), a worrying public health threat (Kieny,
2012, McKenna, 2013, Mole, 2013).
They
can accomplish this in two ways: either by providing internal promoters whose transcripts escape into neighbouring
DNA (Glansdorff, et al., 1981, Simons, et
al., 1983) or by hybrid promoter formation. Many IS carry -35
promoter components oriented towards the flanking DNA (Fig 1.24.1). In a number of cases this plays
an important part in their transposition since a significant number of IS
transpose using an excised transposon circle (Fig 1.24.1) with abutted left and right ends.
For these IS, the other end carries a -10 element oriented inwards towards the
Tpase gene. Together with the -35, this generates a strong promoter on
formation of the circle junction to drive Tpase expression required for
catalysis of integration (Fig 1.24.2) (Chandler, et al., 2015); (Ton-Hoang, et
al., 1997, Perkins-Balding, et al.,
1999, Duval-Valentin, et al., 2001).
Thus if integration occurs next to a resident -10 sequence, the IS -35 sequence
can contribute to a hybrid promoter to drive expression of neighboring genes
[see (Prentki,
et al., 1986)]. At present this phenomenon had been reported to
occur with over 30 different IS in at least 17 bacterial species (Depardieu et
al., 2007, Siguier et al., 2014) (Table 3: IS and Gene Expression). Indeed, specific vector
plasmids have been designed to identify activating insertions (e.g. (Szeverenyi et
al., 1996))
IS activity can affect efflux mechanisms
resulting in increased resistance: IS1 or IS10 insertion can up-regulate the AcrAB-TolC pump in Salmonella enterica (Olliver et al., 2005); IS1 or IS2 insertion upstream of AcrEF (Jellen-Ritter & Kern,
2001, Kobayashi, et al., 2001) and IS186 insertional inactivation of
the AcrAB repressor, AcrR, in Escherichia
coli (Jellen-Ritter
& Kern, 2001), all
lead to increased resistance
to fluoroquinolones. Insertional
inactivation of specific porins can also play a significant role (Wolter, et al.,
2004).
References :
- Aubert
D, Naas T, Heritier C, Poirel L & Nordmann P (2006) Functional
characterization of IS1999, an IS4 family element involved in mobilization and
expression of beta-lactam resistance genes. J
Bacteriol 188: 6506-6514.
- Chandler
M, Fayet O, Rousseau P, Ton Hoang B & Duval-Valentin G (2015)
Copy-out-Paste-in Transposition of IS911: A Major Transposition Pathway. Microbiol Spectr 3.
- Depardieu
F, Podglajen I, Leclercq R, Collatz E & Courvalin P (2007) Modes and
modulations of antibiotic resistance gene expression. Clin Microbiol Rev 20:
79-114.
- Duval-Valentin
G, Normand C, Khemici V, Marty B & Chandler M (2001) Transient promoter
formation: a new feedback mechanism for regulation of IS911 transposition. Embo J 20: 5802-5811.
- Glansdorff
N, Charlier D & Zafarullah M (1981) Activation of gene expression by IS2
and IS3. Cold Spring
Harb.Symp.Quant.Biol. 45 Pt 1:
153-156.
- Jellen-Ritter
AS & Kern WV (2001) Enhanced expression of the multidrug efflux pumps AcrAB
and AcrEF associated with insertion element transposition in Escherichia coli
mutants Selected with a fluoroquinolone. Antimicrob
Agents Chemother 45: 1467-1472.
- Kieny
M-P (2012) The evolving threat of antimicrobial resistance: Options for action.
ed.^eds.), p.^pp. World Health Organization.
- Kobayashi
K, Tsukagoshi N & Aono R (2001) Suppression of hypersensitivity of
Escherichia coli acrB mutant to organic solvents by integrational activation of
the acrEF operon with the IS1 or IS2 element. J Bacteriol 183:
2646-2653.
- McKenna
M (2013) The Last Resort. Nature 499: 394 - 396.
- Mole B
(2013) Farming up trouble. Nature 499: 398-400.
- Olliver
A, Valle M, Chaslus-Dancla E & Cloeckaert A (2005) Overexpression of the
multidrug efflux operon acrEF by insertional activation with IS1 or IS10
elements in Salmonella enterica serovar typhimurium DT204 acrB mutants selected
with fluoroquinolones. Antimicrob Agents
Chemother 49: 289-301.
- Perkins-Balding
D, Duval-Valentin G & Glasgow AC (1999) Excision of IS492 requires flanking
target sequences and results in circle formation in Pseudoalteromonas
atlantica. J Bacteriol 181: 4937-4948.
- Prentki
P, Teter B, Chandler M & Galas DJ (1986) Functional promoters created by
the insertion of transposable element IS1. J
Mol Biol 191: 383-393.
- Reif HJ
& Saedler H (1974) IS1 is Involved in Deletion Formation in the gal Region
of E.coli K12. Mol.Gen.Genet. 137: 17-28.
- Siguier
P, Gourbeyre E & Chandler M (2014) Bacterial insertion sequences: their
genomic impact and diversity. FEMS
Microbiol Rev.
- Simons
RW, Hoopes BC, McClure WR & Kleckner N (1983) Three promoters near the
termini of IS10: pIN, pOUT, and pIII. Cell 34: 673-682.
- Soki J,
Eitel Z, Urban E & Nagy E (2013) Molecular analysis of the carbapenem and
metronidazole resistance mechanisms of Bacteroides strains reported in a
Europe-wide antibiotic resistance survey. Int
J Antimicrob Agents 41: 122-125.
- Szeverenyi
I, Hodel A, Arber W & Olasz F (1996) Vector for IS element entrapment and
functional characterization based on turning on expression of distal
promoterless genes. Gene 174: 103-110.
- Ton-Hoang
B, Betermier M, Polard P & Chandler M (1997) Assembly of a strong promoter
following IS911 circularization and the role of circles in transposition. Embo J 16: 3357-3371.
- Wolter DJ, Smith-Moland E, Goering
RV, Hanson ND & Lister PD (2004) Multidrug resistance associated with mexXY
expression in clinical isolates of Pseudomonas aeruginosa from a Texas
hospital. Diagn Microbiol Infect Dis 50: 43-50.