Mobile element type:  Integron
Name:  In2
Synonyms: 
Accession:  In2-AF071413
Family:  Tn402
Group:  Class 1
First isolate:  Yes
Partial:  ND
Evidence of transposition:  ND

Host Organism:  Shigella flexneri
Date of Isolation:  1950s
Country:  Japan
Molecular Source:  plasmid NR1 (R100)

Name Coordinates Direction Length Sequence
IRt 1-33 + 33 TGTCATTTTC AGAAGACGAC TGCACCAGTT GAT
repeat t1 9-27 + 19 TCAGAAGACG ACTGCACCA
repeat t2 49-67 - 19 TCAGGAGCTG GCTGCACAA
repeat t3 78-97 + 20 TCAGAAGTGA TCTGCACCAA
repeat t4 110-128 + 19 TCAATACTCG TGTGCACCA
repeat i4 10881-10899 - 19 TCAGCGGACG CAGGGAGGA
repeat i3 10909-10927 - 19 TCAGGCAACG ACGGGCTGC
repeat i2 10951-10969 - 19 TCAGAAGCCG ACTGCACTA
IRi 10968-11000 - 33 TGTCGTTTTC AGAAGACGGC TGCACTGAAC GTC
repeat i1 10974-10992 - 19 TCAGAAGACG GCTGCACTG

Name Associated TE Coordinates Orientation Length Sequence
attC qacH_sul1 In2 7473-7506 , 8784-8790 + 41 CCGCTAGCGG GCGGCCGGAA GGTGAATGCT AGGCATCTAA C
attC qacH_sul1 core In2 7473-7506 + 34 CCGCTAGCGG GCGGCCGGAA GGTGAATGCT AGGC
attC aadA1a core In2 8790-8843 + 54 CGCTTGAGTT AAGCCGCGCC GCGAAGCGGC GTCGGCTTGA ACGAATTGTT AGAC
attI In2 9646-9701 + 56 CTTTGTTTTA GGGCGACTGC CCTGCTGCGT AACATCGTTG CTGCTCCATA ACATCA

ORF Summary
Name Associated TE Coordinates Orientation Class Subclass
tniA In2 142-1821 + Transposase
tniB delta1 In2 1824-2684 + Accessory Gene
tnp IS1353 2735-4279 - Transposase
istA IS1326::IS1353 4402-5925 + Transposase None
istB IS1326::IS1353 5912-6697 + Accessory Gene ATPase Transposition Helper
GNAT_fam In2 6873-7373 - Passenger Gene Antibiotic Resistance
sul1 (ARO:3000410) In2 7501-8340 - Passenger Gene Antibiotic Resistance
qacEdelta1 (ARO:3005010) In2 8334-8681 - Passenger Gene Antibiotic Resistance
aadA (ARO:3002601) In2 8845-9636 - Passenger Gene Antibiotic Resistance
intI1 In2 9785-10798 + Integron Integrase Class 1
ORF Details
Gene Name Protein Name Associated TE Gene Length Coordinates Strand
tniA TniA In2 559 142-1821 +
Class: Transposase
Transposase Chemistry: DDE
Comment: can be extended upstream by 12 amino acids| identical to tniA (Tn1721 and In2)| 25% amino acid sequence identity to TnsB from Tn7
Protein Sequence: MATDTPRIPE QGVATLPDEA WERARRRAEI ISPLAQSETV GHEAADMAAQ ALGLSRRQVY VLIRRARQGS GLVTDLVPGQ SGGGKGKGRL PEPVERVIHE LLQKRFLTKQ KRSLAAFHRE VTQVCKAQKL RVPARNTVAL RIASLDPRKV IRRREGQDAA RDLQGVGGEP PAVTAPLEQV QIDHTVIDLI VVDDRDRQPI GRPYLTLAID VFTRCVLGMV VTLEAPSAVS VGLCLVHVAC DKRPWLEGLN VEMDWQMSGK PLLLYLDNAA EFKSEALRRG CEQHGIRLDY RPLGQPHYGG IVERIIGTAM QMIHDELPGT TFSNPDQRGD YDSENKAALT LRELERWLTL AVGTYHGSVH NGLLQPPAAR WAEAVARVGV PAVVTRATSF LVDFLPILRR TLTRTGFVID HIHYYADALK PWIARRERWP SFLIRRDPRD ISRIWVLEPE GQHYLEIPYR TLSHPAVTLW EQRQALAKLR QQGREQVDES ALFRMIGQMR EIVTSAQKAT RKARRDADRR QHLKTSARPD KPVPPDTDIA DPQADNLPPA KPFDQIEEW

Gene Name Protein Name Associated TE Gene Length Coordinates Strand
tniB delta1 TniB delta1 In2 286 1824-2684 +
Class: Accessory Gene
Function: probable ATP-binding protein.
Comment: probably truncated by insertion of IS1326::IS1353
Protein Sequence: MDEYPIIDLS HLLPAAQGLA RLPADERIQR LRADRWIGYP RAVEALNRLE ALYAWPNKQR MPNLLLVGPT NNGKSMIVEK FRRTHPASSD ADQEHIPVLV VQMPSEPSVI RFYVALLAAM GAPLRPRPRL PEMEQLALAL LRKVGVRMLV IDELHNVLAG NSVNRREFLN LLRFLGNELR IPLVGVGTRD AYLAIRSDDQ LENRFEPMML PVWEANDDCC SLLASFAASL PLRRPSPIAT LDMARYLLTR SEGTIGELAH LLMAAAIVAV ESGEEAINHR TLSMAC

Gene Name Protein Name Associated TE Gene Length Coordinates Strand
tnp Tnp IS1353 514 2735-4279 -
Class: Transposase
Function: transposase
Transposase Chemistry: DDE
Protein Sequence: MYSYEDRLRA VRLYLKLGRR MSATLRQLGY PTKNSLKAWL AEFERNQDLR RGYQRIKRQY TDEQKQRAVD HYIEQGYCLS HTIRSLGYPS REALRAWIRD LRPEFARTVV GSSAPTVARS RLEKQQAVIA LNLRVGSAKD VADTVGVSRP TLYNWQHRLL GKVPLKPMTK KKGDTSLEQR HEALLRELAE LESQNQRLRM ENAILEKASE LIKKDMGINP LELTSREKTK VVDALRVTFP LANLLCGLKL ARSTYFYQRL RQTRPDKYTQ VREVIRTIFE DNYRCYGYRR IDSALRLGGM RVSEKVVRRL MAQERLVVRT PRRRRFSAYA GDPTPAVPNL LNRDFHASAP NTKWLTDLTE IHIPAGKVYV SPIVDCFDGL VVAWNIGTSP DANLVNTMLD HAVRTLRPGE HPVIHSDRGS HYRWPAWIRR TENAQLTRSM SKKGCSPDNA ACEGFFGRLK TELIYPRNWQ HVTLKDLMTR IDAYIHWYNE RRIKVSLGGR SPIEYRHAVG LMSV

Gene Name Protein Name Associated TE Gene Length Coordinates Strand
istA IstA IS1326::IS1353 507 4402-5925 +
Class: Transposase
Transposase Chemistry: DDE
Protein Sequence: MINVAILSAI RRWHFRDGAS IREIARRSGL SRNTVRKYLQ SKVVEPQYPA RDSVGKLSPF EPKLRQWLST EHKKTKKLRR NLRSMYRDLV ALGFTGSYDR VCAFARQWKD SEQFKAQTSG KGCFIPLRFA CGEAFQFDWS EDFARIAGKQ VKLQIAQFKL AHSRAFVLRA YYQQKHEMLF DAHWHAFQIF GGIPKRGIYD NMKTAVDSVG RGKERRVNQR FTAMVSHYLF DAQFCNPASG WEKGQIEKNV QDSRQRLWQG APDFQSLADL NVWLEHRCKA LWSELRHPEL DQTVQEAFAD EQGELMALPN AFDAFVEQTK RVTSTCLVHH EGNRYSVPAS YANRAISLRI YADKLVMAAE GQHIAEHPRL FGSGHARRGH TQYDWHHYLS VLQKKPGALR NGAPFAELPP AFKKLQSILL QRPGGDRDMV EILALVLHHD EGAVLSAVEL ALECGKPSKE HVLNLLGRLT EEPPPKPIPI PKGLRLTLEP QANVNRYDSL RRAHDAA

Gene Name Protein Name Associated TE Gene Length Coordinates Strand
istB IstB IS1326::IS1353 261 5912-6697 +
Class: Accessory Gene
Subclass: ATPase Transposition Helper
Function: stimulates transposition
Protein Sequence: MMQHEGHVRI LKSLKLFGMA HAIEELGNQN SPAFNQALPM LDSLIKAEVA EREVRSVNYQ LRVAKFPVYR DLVGFDFSQS LVNEATVKQL HRCDFMEQAQ NVVLIGGPGT GKTHLATAIG TQAVMHLNRR VRFFSTVDLV NALEQEKSSG RQGQIANRLL YADLVILDEL GYLPFSQTGG ALLFHLLSKL YEKTSVILTT NLSFSEWSRV FGDEKMTTAL LDRLTHHCHI LETGNESYRF KHSSTQNKQE EKQTRKLKIE T

Gene Name Protein Name Associated TE Gene Length Coordinates Strand
GNAT_fam GNAT_fam In2 166 6873-7373 -
Class: Passenger Gene
Subclass: Antibiotic Resistance
Sequence Family: GNAT
Target: aminoglycoside antibiotic (ARO:0000016)
Protein Sequence: MDSEEPPNVR VACSGDIDEV VRLMHDAAAW MSAKGTPAWD VARIDRTFAE TFVLRSELLV ASCSDGIVGC CTLSAEDPEF WPDALKGEAA YLHKLAVRRT HAGRGVSSAL IEACRHAART QGCAKLRLDC HPNLRGLYER LGFTHVDTFN PGWDPTFIAE RLELEI

Gene Name Protein Name Associated TE Gene Length Coordinates Strand
sul1 (ARO:3000410) Sul1 In2 279 7501-8340 -
Class: Passenger Gene
Subclass: Antibiotic Resistance
Function: antibiotic target replacement (ARO:0001002)
Sequence Family: sulfonamide resistant sul (ARO:3004238)
Transposase Chemistry: dihydropteroate synthase
Target: sulfonamide antibiotic (ARO:3000282)||sulfone antibiotic (ARO:3003401)
Comment: perfect match to reference sequence for ARO:3000410
Protein Sequence: MVTVFGILNL TEDSFFDESR RLDPAGAVTA AIEMLRVGSD VVDVGPAASH PDARPVSPAD EIRRIAPLLD ALSDQMHRVS IDSFQPETQR YALKRGVGYL NDIQGFPDPA LYPDIAEADC RLVVMHSAQR DGIATRTGHL RPEDALDEIV RFFEARVSAL RRSGVAADRL ILDPGMGFFL SPAPETSLHV LSNLQKLKSA LGLPLLVSVS RKSFLGATVG LPVKDLGPAS LAAELHAIGN GADYVRTHAP GDLRSAITFS ETLAKFRSRD ARDRGLDHA

Gene Name Protein Name Associated TE Gene Length Coordinates Strand
qacEdelta1 (ARO:3005010) QacEdelta1 In2 115 8334-8681 -
Class: Passenger Gene
Subclass: Antibiotic Resistance
Function: antibiotic efflux (ARO:0010000)
Sequence Family: major facilitator superfamily (MFS) antibiotic efflux pump (ARO:0010002)
Target: disinfecting agents and antiseptics(ARO:3005386)
Comment: subunit of the qac multidrug efflux pump||perfect match to reference sequence for ARO:3005010 (bitscore:219)
Protein Sequence: MKGWLFLVIA IVGEVIATSA LKSSEGFTKL APSAVVIIGY GIAFYFLSLV LKSIPVGVAY AVWSGLGVVI ITAIAWLLHG QKLDAWGFVG MGLIIAAFLL ARSPSWKSLR RPTPW

Gene Name Protein Name Associated TE Gene Length Coordinates Strand
aadA (ARO:3002601) AadA In2 263 8845-9636 -
Class: Passenger Gene
Subclass: Antibiotic Resistance
Function: antibiotic inactivation (ARO:0001004)
Sequence Family: ANT(3'')(ARO:3004275)
Transposase Chemistry: aminoglycoside nucleotidyltransferase
Target: aminoglycoside antibiotic (ARO:0000016)
Comment: perfect match to reference sequence for ARO:3002601||Synonyms: aadA1-pm, aadA, aadA1, aad(3'')(9)
Protein Sequence: MREAVIAEVS TQLSEVVGVI ERHLEPTLLA VHLYGSAVDG GLKPHSDIDL LVTVTVRLDE TTRRALINDL LETSASPGES EILRAVEVTI VVHDDIIPWR YPAKRELQFG EWQRNDILAG IFEPATIDID LAILLTKARE HSVALVGPAA EELFDPVPEQ DLFEALNETL TLWNSPPDWA GDERNVVLTL SRIWYSAVTG KIAPKDVAAD WAMERLPAQY QPVILEARQA YLGQEEDRLA SRADQLEEFV HYVKGEITKV VGK

Gene Name Protein Name Associated TE Gene Length Coordinates Strand
intI1 IntI1 In2 337 9785-10798 +
Class: Integron Integrase
Subclass: Class 1
Sequence Family: Class 1 Integron Tyrosine Integrase
Transposase Chemistry: Tyrosine
Protein Sequence: MKTATAPLPP LRSVKVLDQL RERIRYLHYS LRTEQAYVHW VRAFIRFHGV RHPATLGSSE VEAFLSWLAN ERKVSVSTHR QALAALLFFY GKVLCTDLPW LQEIGRPRPS RRLPVVLTPD EVVRILGFLE GEHRLFAQLL YGTGMRISEG LQLRVKDLDF DHGTIIVREG KGSKDRALML PESLAPSLRE QLSRARAWWL KDQAEGRSGV ALPDALERKY PRAGHSWPWF WVFAQHTHST DPRSGVVRRH HMYDQTFQRA FKRAVEQAGI TKPATPHTLR HSFATALLRS GYDIRTVQDL LGHSDVSTTM IYTHVLKVGG AGVRSPLDAL PPLTSER

TnCentral Accession TE Name Type Coordinates Strand Length
IS1326_IS1353-AF071413 IS1326::IS1353 insertion sequence 2679-6764 + 4086
IS1353-AF071413 IS1353 insertion sequence 2707-4320 + 1614

Name Associated TE Coordinates Direction Length Sequence
IRL IS1326::IS1353 2679-2704 + 26 TGTTGAGTTG CATCTAAAAT TGACCC
IRR IS1353 2707-2719 + 13 TGGGGGTGCG GAC
IRL IS1353 4309-4320 - 12 TGGGGTGCGG AC
IRR IS1326::IS1353 6739-6764 - 26 TGTTGATTTG CACCCAAATT TGACCC

1.  NAKAYA R, NAKAMURA A, MURATA Y.  Resistance transfer agents in Shigella..  Biochem Biophys Res Commun. 1960 Dec;3:654-9. doi: 10.1016/0006-291x(60)90081-4.  PubMed ID: 13727669 .
2.  Kopecko DJ, Brevet J, Cohen SN.  Involvement of multiple translocating DNA segments and recombinational hotspots in the structural evolution of bacterial plasmids..  J Mol Biol. 1976 Dec;108(2):333-60. doi: 10.1016/s0022-2836(76)80124-6.  PubMed ID: 796463 .
3.  Diver WP, Grinsted J, Fritzinger DC, Brown NL, Altenbuchner J, Rogowsky P, Schmitt R.  DNA sequences of and complementation by the tnpR genes of Tn21, Tn501 and Tn1721..  Mol Gen Genet. 1983;191(2):189-93. doi: 10.1007/BF00334812.  PubMed ID: 6312271 .
4.  Zheng ZX, Chandler M, Hipskind R, Clerget M, Caro L.  Dissection of the r-determinant of the plasmid R100.1: the sequence at the extremities of Tn21..  Nucleic Acids Res. 1981 Dec 11;9(23):6265-78. doi: 10.1093/nar/9.23.6265.  PubMed ID: 6275355 .
5.  Barrineau P, Gilbert P, Jackson WJ, Jones CS, Summers AO, Wisdom S.  The DNA sequence of the mercury resistance operon of the IncFII plasmid NR1..  J Mol Appl Genet. 1984;2(6):601-19.  PubMed ID: 6530603 .
6.  Misra TK, Brown NL, Fritzinger DC, Pridmore RD, Barnes WM, Haberstroh L, Silver S.  Mercuric ion-resistance operons of plasmid R100 and transposon Tn501: the beginning of the operon including the regulatory region and the first two structural genes..  Proc Natl Acad Sci U S A. 1984 Oct;81(19):5975-9. doi: 10.1073/pnas.81.19.5975.  PubMed ID: 6091128 .
7.  Misra TK, Brown NL, Haberstroh L, Schmidt A, Goddette D, Silver S.  Mercuric reductase structural genes from plasmid R100 and transposon Tn501: functional domains of the enzyme..  Gene. 1985;34(2-3):253-62. doi: 10.1016/0378-1119(85)90134-9.  PubMed ID: 2989109 .
8.  Rogowsky P, Halford SE, Schmitt R.  Definition of three resolvase binding sites at the res loci of Tn21 and Tn1721..  EMBO J. 1985 Aug;4(8):2135-41. doi: 10.1002/j.1460-2075.1985.tb03904.x.  PubMed ID: 2998784 .
9.  Hyde DR, Tu CP.  tnpM: a novel regulatory gene that enhances Tn21 transposition and suppresses cointegrate resolution..  Cell. 1985 Sep;42(2):629-38. doi: 10.1016/0092-8674(85)90120-5.  PubMed ID: 2992807 .
10.  Brown NL, Misra TK, Winnie JN, Schmidt A, Seiff M, Silver S.  The nucleotide sequence of the mercuric resistance operons of plasmid R100 and transposon Tn501: further evidence for mer genes which enhance the activity of the mercuric ion detoxification system..  Mol Gen Genet. 1986 Jan;202(1):143-51. doi: 10.1007/BF00330531.  PubMed ID: 3007931 .
11.  Ward E, Grinsted J.  The nucleotide sequence of the tnpA gene of Tn21..  Nucleic Acids Res. 1987 Feb 25;15(4):1799-806. doi: 10.1093/nar/15.4.1799.  PubMed ID: 3029727 .
12.  Sundstrom L, Radstrom P, Swedberg G, Skold O.  Site-specific recombination promotes linkage between trimethoprim- and sulfonamide resistance genes. Sequence characterization of dhfrV and sulI and a recombination active locus of Tn21..  Mol Gen Genet. 1988 Aug;213(2-3):191-201. doi: 10.1007/BF00339581.  PubMed ID: 3054482 .
13.  Schmidt FR, Nucken EJ, Henschke RB.  Structure and function of hot spots providing signals for site-directed specific recombination and gene expression in Tn21 transposons..  Mol Microbiol. 1989 Nov;3(11):1545-55. doi: 10.1111/j.1365-2958.1989.tb00140.x.  PubMed ID: 2559298 .
14.  Stokes HW, Hall RM.  A novel family of potentially mobile DNA elements encoding site-specific gene-integration functions: integrons..  Mol Microbiol. 1989 Dec;3(12):1669-83. doi: 10.1111/j.1365-2958.1989.tb00153.x.  PubMed ID: 2560119 .
15.  Guerineau F, Brooks L, Mullineaux P.  Expression of the sulfonamide resistance gene from plasmid R46..  Plasmid. 1990 Jan;23(1):35-41. doi: 10.1016/0147-619x(90)90042-b.  PubMed ID: 2190244 .
16.  Mercier J, Lachapelle J, Couture F, Lafond M, Vezina G, Boissinot M, Levesque RC.  Structural and functional characterization of tnpI, a recombinase locus in Tn21 and related beta-lactamase transposons..  J Bacteriol. 1990 Jul;172(7):3745-57. doi: 10.1128/jb.172.7.3745-3757.1990.  PubMed ID: 2163386 .
17.  Allmeier H, Cresnar B, Greck M, Schmitt R.  Complete nucleotide sequence of Tn1721: gene organization and a novel gene product with features of a chemotaxis protein..  Gene. 1992 Feb 1;111(1):11-20. doi: 10.1016/0378-1119(92)90597-i.  PubMed ID: 1312499 .
18.  Summers AO.  Untwist and shout: a heavy metal-responsive transcriptional regulator..  J Bacteriol. 1992 May;174(10):3097-101. doi: 10.1128/jb.174.10.3097-3101.1992.  PubMed ID: 1577681 .
19.  Levesque C, Brassard S, Lapointe J, Roy PH.  Diversity and relative strength of tandem promoters for the antibiotic-resistance genes of several integrons..  Gene. 1994 May 3;142(1):49-54. doi: 10.1016/0378-1119(94)90353-0.  PubMed ID: 8181756 .
20.  Radstrom P, Skold O, Swedberg G, Flensburg J, Roy PH, Sundstrom L.  Transposon Tn5090 of plasmid R751, which carries an integron, is related to Tn7, Mu, and the retroelements..  J Bacteriol. 1994 Jun;176(11):3257-68. doi: 10.1128/jb.176.11.3257-3268.1994.  PubMed ID: 8195081 .
21.  Recchia GD, Stokes HW, Hall RM.  Characterisation of specific and secondary recombination sites recognised by the integron DNA integrase..  Nucleic Acids Res. 1994 Jun 11;22(11):2071-8. doi: 10.1093/nar/22.11.2071.  PubMed ID: 8029014 .
22.  Hall RM, Brown HJ, Brookes DE, Stokes HW.  Integrons found in different locations have identical 5' ends but variable 3' ends..  J Bacteriol. 1994 Oct;176(20):6286-94. doi: 10.1128/jb.176.20.6286-6294.1994.  PubMed ID: 7929000 .
23.  Recchia GD, Hall RM.  Gene cassettes: a new class of mobile element..  Microbiology (Reading). 1995 Dec;141 ( Pt 12):3015-27. doi: 10.1099/13500872-141-12-3015.  PubMed ID: 8574395 .
24.  Brown HJ, Stokes HW, Hall RM.  The integrons In0, In2, and In5 are defective transposon derivatives..  J Bacteriol. 1996 Aug;178(15):4429-37. doi: 10.1128/jb.178.15.4429-4437.1996.  PubMed ID: 8755869 .
25.  Collis CM, Kim MJ, Stokes HW, Hall RM.  Binding of the purified integron DNA integrase Intl1 to integron- and cassette-associated recombination sites..  Mol Microbiol. 1998 Jul;29(2):477-90. doi: 10.1046/j.1365-2958.1998.00936.x.  PubMed ID: 9720866 .
26.  Gravel A, Fournier B, Roy PH.  DNA complexes obtained with the integron integrase IntI1 at the attI1 site..  Nucleic Acids Res. 1998 Oct 1;26(19):4347-55. doi: 10.1093/nar/26.19.4347.  PubMed ID: 9742234 .
27.  Liebert CA, Hall RM, Summers AO.  Transposon Tn21, flagship of the floating genome..  Microbiol Mol Biol Rev. 1999 Sep;63(3):507-22. doi: 10.1128/MMBR.63.3.507-522.1999.  PubMed ID: 10477306 .
28.  Partridge SR, Hall RM.  In34, a complex In5 family class 1 integron containing orf513 and dfrA10..  Antimicrob Agents Chemother. 2003 Jan;47(1):342-9. doi: 10.1128/AAC.47.1.342-349.2003.  PubMed ID: 12499211 .
29.  Kholodii G, Mindlin S, Petrova M, Minakhina S.  Tn5060 from the Siberian permafrost is most closely related to the ancestor of Tn21 prior to integron acquisition..  FEMS Microbiol Lett. 2003 Sep 26;226(2):251-5. doi: 10.1016/S0378-1097(03)00559-7.  PubMed ID: 14553919 .