365:
breaks to each end. The excised sequence is then inserted to another target DNA site. Much like other characterized transposons, the mechanism for Tn7 transposition involves cleavage of the 3' ends from the donating DNA by the TnsA protein of the TnsAB transposase. However, Tn7 is also uniquely cleaved near the 5' ends, about 5 bp from the 5' end towards the Tn7 transposon, by the TnsB protein of TnsAB. After the insertion of the transposon into the target DNA site, the 3' ends are covalently linked to the target DNA, but the 5 bp gaps are still present at the 5' ends. As a result, repair of these gaps leads to a further 5 bp duplication at the target site. The TnsC protein interacts with the transposase enzyme and the target DNA to promote the excision and insertion processes. The ability of TnsC to activate the transposase depends on its interaction with a target DNA along with its appropriate targeting protein, TnsD or TnsE. The TnsD and TnsE proteins are alternative target selectors that are also DNA binding
245:. One of the reasons Tn5 is so unreactive is because the N- and C-termini are located in relatively close proximity to one another and tend to inhibit each other. This was elucidated by the characterization of several mutations which resulted in hyperactive forms of transposases. One such mutation, L372P, is a mutation of amino acid 372 in the Tn5 transposase. This amino acid is generally a leucine residue in the middle of an alpha helix. When this leucine is replaced with a proline residue the alpha helix is broken, introducing a conformational change to the C-terminal domain, separating it from the N-terminal domain enough to promote higher activity of the protein. The transposition of a transposon often needs only three pieces: the transposon, the transposase enzyme, and the target DNA for the insertion of the transposon. This is the case with Tn5, which uses a cut-and-paste mechanism for moving around transposons.
349:, which can be replicated and distributed between bacteria. However, Tn7 is unique in that it also transposes at high-frequency into a single specific site in bacterial chromosomes called attTn7. This specific sequence is an essential and highly conserved gene found in many strains of bacteria. However, the recombination is not deleterious to the host bacterium as Tn7 actually transposes downstream of the gene after recognizing it, resulting in a safe way to propagate the transposon without killing the host. This highly evolved and sophisticated target-site selection pathway suggests this pathway evolved to promote coexistence between the transposon and it host, as well as Tn7's successful transmission into future generations of bacterium.
345:(transposon), the sequence can duplicate and move itself within a genome by utilizing a self-encoded recombinase enzyme called a transposase, resulting in effects such as creating or reversing mutations and changing genome size. The Tn7 transposon has developed two mechanisms to promote its propagation among prokaryotes. Like many other bacterial transposons, Tn7 transposes at low-frequency and inserts into many different sites with little to no site-selectivity. Through this first pathway, Tn7 is preferentially directed into conjugable
2690:
249:
magnesium and manganese, which are important in the catalytic reaction. Because transposase is incredibly inactive, the DDE region is mutated so that the transposase becomes hyperactive and catalyzes the movement of the transposon. The glutamate is transformed into an aspartate and the two aspartates into glutamates. Through this mutation, the study of Tn5 becomes possible, but some steps in the catalytic process are lost as a result.
307:. SB transposase belongs to the DD family of transposases, which in turn belong to a large superfamily of polynucleotidyl transferases that includes RNase H, RuvC Holliday resolvase, RAG proteins, and retroviral integrases. The SB system is used primarily in vertebrate animals for gene transfer, including gene therapy, and gene discovery. The engineered SB100X is an enzyme that directs the high levels of transposon integration.
82:
253:
248:
Tn5 and most other transposases contain a DDE motif, which is the active site that catalyzes the movement of the transposon. Aspartate-97, aspartate-188, and glutamate-326 make up the active site, which is a triad of acidic residues. The DDE motif is said to coordinate divalent metal ions, most often
352:
The Tn7 transposon is 14 kb long and codes for five enzymes. The ends of the DNA sequence consists of two segments that the Tn7 transposase interacts with during recombination. The left segment (Tn7-L) is 150 bp long and the right sequence (Tn7-R) is 90 bp long. Both ends of the transposon contain a
259:
There are several steps which catalyze the movement of the transposon, including Tnp binding, synapsis (the creation of a synaptic complex), cleavage, target capture, and strand transfer. Transposase then binds to the DNA strand and creates a clamp over the transposon end of the DNA and inserts into
374:
of the transposase's activity, communicating between the transposase and TnsD and TnsE. When the TnsE protein interacts with the TnsABC core machinery, Tn7 preferentially directs insertions into conjugable plasmids. When the TnsD protein interacts with TnsABC, Tn7 preferentially directs insertions
264:
transposon to a suitable location. Not much is known about the target capture, although there is a sequence bias which has not yet been determined. After target capture, the transposase attacks the target DNA nine base pairs apart, resulting in the integration of the transposon into the target DNA.
51:
changed their position, jumping between different loci or from one chromosome to another. The repositioning of these transposons (which coded for color) allowed other genes for pigment to be expressed. Transposition in maize causes changes in color; however, in other organisms, such as bacteria, it
369:
that promote excision and insertion of Tn7. Their ability to interact with a particular target DNA is key to the target-site selection of Tn7. The proteins TnsA, TnsB, and TnsC thus form the core machinery of Tn7: TnsA and TnsB interact together to form the transposase, while TnsC functions as a
364:
The Tn7 transposon codes for five proteins: TnsA, TnsB, TnsC, TnsD, and TnsE. TnsA and TnsB interact together to form the Tn7 transposase enzyme TnsAB. The enzyme specifically recognizes and binds to the ends of the DNA sequence of the transposon, and excises it by introducing double-stranded DNA
290:
library preparation. The Tn5-based strategy can simplify the library preparation protocol significantly and can even can be incorporated into the direct colony-PCR for large numbers of bacterial isolates with no obvious coverage bias. The main disadvantages are less control of fragmented size
263:
In cleavage, the magnesium ions activate oxygen from water molecules and expose them to nucleophilic attack. This allows the water molecules to nick the 3' strands on both ends and create a hairpin formation, which separates the transposon from the donor DNA. Next, the transposase moves the
63:
EC 2.7.7. Genes encoding transposases are widespread in the genomes of most organisms and are the most abundant genes known. During the course of human evolution, as much as 40% of the human genome has moved around via methods such as transposition of transposons.
240:
Tn5 and other transposases are notably inactive. Because DNA transposition events are inherently mutagenic, the low activity of transposases is necessary to reduce the risk of causing a fatal mutation in the host, and thus eliminating the
353:
series of 22 bp binding sites that the Tn7 transposase recognizes and binds to. Within the transposon are five discrete genes encoding for proteins that make up the transposition machinery. In addition, the transposon contains an
35:, typically by a cut-and-paste mechanism or a replicative mechanism, in a process known as transposition. The word "transposase" was first coined by the individuals who cloned the enzyme required for transposition of the
1106:
Mátés L, Chuah MK, Belay E, Jerchow B, Manoj N, Acosta-Sanchez A, et al. (June 2009). "Molecular evolution of a novel hyperactive
Sleeping Beauty transposase enables robust stable gene transfer in vertebrates".
285:
Tn5 is utilized in genome sequencing by using the Tn5 to append sequencing adaptors and fragment the DNA in a single enzymatic reaction in 2010, reducing the time and input requirements over traditional
2217:
2331:
260:
the active site. Once the transposase binds to the transposon, it produces a synaptic complex in which two transposases are bound in a cis/trans relationship with the transposon.
391:
Heffron F, McCarthy BJ, Ohtsubo H, Ohtsubo E (December 1979). "DNA sequence analysis of the transposon Tn3: three genes and three sites involved in transposition of Tn3".
1691:
1714:
181:
2227:
1497:
1663:
2201:
2222:
969:
Aronovich, E.L., McIvor, R.S., and
Hackett, P.B. (2011). The Sleeping Beauty transposon system – A non-viral vector for gene therapy. Hum. Mol. Genet. (in press)
1150:
Grabundzija I, Irgang M, Mátés L, Belay E, Matrai J, Gogol-Döring A, Kawakami K, Chen W, Ruiz P, Chuah MK, VandenDriessche T, Izsvák Z, Ivics Z (June 2010).
291:
compared to enzymatic fragmentation and mechanical fragmentation, and a bias toward high G-C content. This means of library preparation is also used in the
2342:
2320:
2306:
2138:
375:
downstream into a single essential and highly conserved site in the bacterial chromosome. This site, attTn7, is specifically recognized by TnsD.
2353:
56:. Transposition is also important in creating genetic diversity within species and generating adaptability to changing living conditions.
2281:
2248:
267:
As mentioned before, due to the mutations of the DDE, some steps of the process are lost—for example, when this experiment is performed
1809:
2143:
1769:
598:
Lovell S, Goryshin IY, Reznikoff WR, Rayment I (April 2002). "Two-metal active site binding of a Tn5 transposase synaptic complex".
2310:
1345:"Recognition of Escherichia coli attTn7 by transposon Tn7: lack of specific sequence requirements at the point of Tn7 insertion"
282:-1 and other retroviral diseases. By studying Tn5, much can also be discovered about other transposases and their activities.
47:, but the actual molecular basis for transposition was described by later groups. McClintock discovered that some segments of
1949:
886:
Ivics Z, Izsvák Z (January 2005). "A whole lotta jumpin' goin' on: new transposon tools for vertebrate functional genomics".
129:
201:
2083:
521:
Aziz, R.K., M. Breitbart and R.A. Edwards (2010). Transposases are the most abundant, most ubiquitous genes in nature.
2409:
1653:
1447:
304:
2720:
1794:
747:"Molecular reconstruction of Sleeping Beauty: a Tc1-like transposon from fish and its transposition in human cells"
271:, and SDS heat treatment denatures the transposase. However, it is still uncertain what happens to the transposase
1201:"Transposition of a DNA sequence encoding trimethoprim and streptomycin resistances from R483 to other replicons"
696:"Rapid, low-input, low-bias construction of shotgun fragment libraries by high-density in vitro transposition"
189:
2565:
2367:
2362:
1922:
1740:
2232:
2120:
2088:
2073:
1755:
2715:
2680:
1020:
Carlson CM, Largaespada DA (July 2005). "Insertional mutagenesis in mice: new perspectives and tools".
287:
185:
1425:
2666:
2653:
2640:
2627:
2614:
2601:
2588:
2550:
1631:
142:
2560:
2514:
2457:
1818:
1799:
1730:
1584:
1464:
1398:
60:
53:
650:"Tn5 transposase active site mutations suggest position of donor backbone DNA in synaptic complex"
2462:
1676:
2130:
1903:
1813:
329:), and was first discovered as a DNA sequence in bacterial chromosomes and naturally occurring
316:
2483:
2402:
1984:
1939:
1641:
1626:
1562:
1542:
1440:
923:"Translating Sleeping Beauty transposition into cellular therapies: victories and challenges"
1063:
Copeland NG, Jenkins NA (October 2010). "Harnessing transposons for cancer gene discovery".
2555:
1898:
1893:
1773:
1636:
1616:
1525:
795:
366:
342:
242:
168:
8:
2519:
2161:
2153:
1934:
1601:
1537:
1501:
1394:
799:
2452:
2024:
1962:
1913:
1648:
1475:
1325:
1176:
1151:
1132:
1088:
1045:
997:
972:
947:
922:
863:
838:
819:
722:
695:
623:
577:
553:
528:
416:
40:
1369:
1344:
1225:
1200:
763:
746:
2710:
2039:
2029:
2019:
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1414:
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1124:
1080:
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1002:
952:
903:
868:
811:
768:
727:
671:
615:
558:
504:
499:
482:
408:
404:
278:
The study of transposase Tn5 is of general importance because of its similarities to
176:
134:
1360:
1329:
1136:
823:
420:
110:
2498:
2493:
2467:
2395:
2372:
2034:
2014:
1956:
1852:
1750:
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1681:
1579:
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1092:
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942:
934:
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661:
627:
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548:
540:
494:
400:
321:
164:
1216:
2545:
2529:
2442:
2295:
2285:
2107:
2078:
1704:
1611:
1596:
807:
122:
2694:
2583:
2524:
2263:
2102:
2009:
2001:
1878:
1841:
1266:
1249:
854:
712:
371:
36:
899:
2704:
2488:
2447:
1532:
358:
1152:"Comparative analysis of transposable element vector systems in human cells"
973:"The Sleeping Beauty transposon system: a non-viral vector for gene therapy"
2437:
1621:
1591:
1520:
1321:
1185:
1128:
1084:
1041:
1006:
956:
938:
907:
872:
731:
675:
666:
649:
619:
562:
508:
338:
334:
214:
1378:
815:
772:
138:
2661:
2596:
2432:
2177:
1778:
1515:
1456:
1275:
1234:
988:
544:
412:
229:
303:
The
Sleeping Beauty (SB) transposase is the recombinase that drives the
1989:
1832:
1745:
1510:
1460:
1455:
1167:
223:
48:
28:
2635:
2609:
2268:
2181:
1718:
1484:
1313:
529:"Transposases are the most abundant, most ubiquitous genes in nature"
234:
218:
1076:
1033:
2689:
2115:
1695:
1120:
611:
354:
346:
330:
292:
117:
39:. The existence of transposons was postulated in the late 1940s by
2218:
CDP-diacylglycerol—glycerol-3-phosphate 3-phosphatidyltransferase
2066:
2051:
1667:
1419:
1409:
440:
1250:"Two Naturally Occurring Transposons Indistinguishable from Tn7"
2648:
2418:
2044:
1479:
1300:
Peters J, Craig NL (Nov 2001). "Tn7: smarter than we thought".
196:
81:
32:
24:
597:
2622:
2061:
2056:
1908:
786:
Craig NL (October 1995). "Unity in transposition reactions".
745:
Ivics, Z.; Hackett, P.B.; Plasterk, R.A.; Izsvak, Z. (1997).
390:
44:
1404:
Overview of all the structural information available in the
233:
bacteria. The transposon codes for antibiotic resistance to
1972:
1967:
1863:
1858:
1557:
1552:
1547:
1149:
921:
Izsvák Z, Hackett PB, Cooper LJ, Ivics Z (September 2010).
158:
105:
2387:
744:
279:
1342:
920:
1199:
Barth PT, Datta N, Hedges RW, Grinter NJ (March 1976).
1198:
839:"DDE transposases: Structural similarity and diversity"
252:
16:
Enzyme that catalyzes movement of transposable elements
1343:
Gringauz E, Orle KA, Waddell CS, Craig NL (Jun 1988).
1105:
970:
2678:
2228:
CDP-diacylglycerol—inositol 3-phosphatidyltransferase
2233:
CDP-diacylglycerol—choline O-phosphatidyltransferase
526:
483:"Tn5 as a model for understanding DNA transposition"
2223:
CDP-diacylglycerol—serine O-phosphatidyltransferase
1019:
971:Aronovich EL, McIvor RS, Hackett PB (April 2011).
217:superfamily of proteins which includes retroviral
647:
31:and catalysing its movement to another part of a
2702:
836:
1062:
86:tn5 transposase: 20mer outside end 2 mn complex
527:Aziz RK, Breitbart M, Edwards RA (July 2010).
298:
2403:
1441:
361:of genes encoding for antibiotic-resistance.
2249:N-acetylglucosamine-1-phosphate transferase
2139:UTP—glucose-1-phosphate uridylyltransferase
1299:
885:
837:Nesmelova IV, Hackett PB (September 2010).
333:that encoded resistance to the antibiotics
2410:
2396:
1448:
1434:
1247:
434:
432:
430:
80:
2343:serine/threonine-specific protein kinases
2321:serine/threonine-specific protein kinases
2144:Galactose-1-phosphate uridylyltransferase
1397:at the U.S. National Library of Medicine
1368:
1295:
1293:
1291:
1289:
1287:
1285:
1265:
1224:
1175:
996:
946:
862:
762:
721:
711:
665:
552:
498:
480:
213:Transposase (Tnp) Tn5 is a member of the
1418:(Transposase for transposon Tn5) at the
648:Peterson G, Reznikoff W (January 2003).
438:
251:
569:
427:
2703:
1282:
237:and other aminoglycoside antibiotics.
43:, who was studying the inheritance of
2391:
1429:
1302:Nature Reviews Molecular Cell Biology
785:
476:
474:
693:
689:
687:
685:
643:
641:
639:
637:
593:
591:
575:
472:
470:
468:
466:
464:
462:
460:
458:
456:
454:
654:The Journal of Biological Chemistry
357:, a DNA segment containing several
75:Transposase Tn5 dimerisation domain
27:capable of binding to the end of a
13:
319:found in many prokaryotes such as
67:
59:Transposases are classified under
14:
2732:
1654:Glucose-1,6-bisphosphate synthase
1388:
682:
634:
588:
451:
310:
305:Sleeping Beauty transposon system
2688:
1795:Ribose-phosphate diphosphokinase
500:10.1046/j.1365-2958.2003.03382.x
1361:10.1128/jb.170.6.2832-2840.1988
1336:
1254:Journal of General Microbiology
1241:
1192:
1143:
1099:
1056:
1013:
963:
914:
879:
830:
779:
341:. Specifically classified as a
1248:Barth PT, Datta N (Sep 1977).
843:Advanced Drug Delivery Reviews
738:
694:Adey, Andrew (December 2010).
515:
384:
1:
2368:Protein-histidine tele-kinase
2363:Protein-histidine pros-kinase
2242:Glycosyl-1-phosphotransferase
1217:10.1128/JB.125.3.800-810.1976
764:10.1016/s0092-8674(00)80436-5
378:
153:Available protein structures:
2074:RNA-dependent RNA polymerase
808:10.1126/science.270.5234.253
439:Goodsell D (December 2006).
405:10.1016/0092-8674(79)90228-9
7:
2417:
1981:RNA-directed DNA polymerase
1849:DNA-directed DNA polymerase
481:Reznikoff WS (March 2003).
299:Sleeping Beauty transposase
10:
2737:
2334:: protein-dual-specificity
1267:10.1099/00221287-102-1-129
855:10.1016/j.addr.2010.06.006
713:10.1186/gb-2010-11-12-r119
288:next-generation sequencing
2574:
2566:Michaelis–Menten kinetics
2538:
2507:
2476:
2425:
2352:
2330:
2305:
2280:
2261:
2241:
2210:
2200:
2170:
2152:
2129:
2101:
2000:
1840:
1831:
1808:
1768:
1713:
1690:
1662:
1496:
1472:
900:10.1016/j.tig.2004.11.008
600:Nature Structural Biology
195:
175:
157:
152:
148:
128:
116:
104:
96:
91:
79:
74:
2458:Diffusion-limited enzyme
2311:protein-serine/threonine
2211:Phosphatidyltransferases
1800:Thiamine diphosphokinase
1399:Medical Subject Headings
977:Human Molecular Genetics
315:The Tn7 transposon is a
2721:Mobile genetic elements
1349:Journal of Bacteriology
1205:Journal of Bacteriology
1022:Nature Reviews Genetics
2131:Nucleotidyltransferase
1814:nucleotidyltransferase
1741:Nucleoside-diphosphate
1065:Nature Reviews. Cancer
939:10.1002/bies.201000027
667:10.1074/jbc.M208968200
533:Nucleic Acids Research
523:Nucleic Acids Research
487:Molecular Microbiology
317:mobile genetic element
256:
221:. Tn5 can be found in
2551:Eadie–Hofstee diagram
2484:Allosteric regulation
1985:Reverse transcriptase
445:Molecule of the Month
255:
54:antibiotic resistance
23:is any of a class of
2561:Lineweaver–Burk plot
1774:diphosphotransferase
1756:Thiamine-diphosphate
1463:-containing groups (
447:. Protein Data Bank.
343:transposable element
243:transposable element
2356:: protein-histidine
2274:; protein acceptor)
2162:mRNA capping enzyme
2154:Guanylyltransferase
800:1995Sci...270..253C
2520:Enzyme superfamily
2453:Enzyme promiscuity
1632:Phosphoinositide 3
1476:phosphotransferase
1168:10.1038/mt.2010.47
989:10.1093/hmg/ddr140
888:Trends in Genetics
545:10.1093/nar/gkq140
525:38(13): 4207–4217.
257:
41:Barbara McClintock
2716:Molecular biology
2676:
2675:
2385:
2384:
2381:
2380:
2257:
2256:
2196:
2195:
2097:
2096:
2010:Template-directed
1764:
1763:
1731:Phosphomevalonate
1156:Molecular Therapy
211:
210:
207:
206:
202:structure summary
2728:
2693:
2692:
2684:
2556:Hanes–Woolf plot
2499:Enzyme activator
2494:Enzyme inhibitor
2468:Enzyme catalysis
2412:
2405:
2398:
2389:
2388:
2373:Histidine kinase
2296:tyrosine kinases
2286:protein-tyrosine
2278:
2277:
2208:
2207:
2015:RNA polymerase I
1838:
1837:
1829:
1828:
1682:Aspartate kinase
1677:Phosphoglycerate
1494:
1493:
1450:
1443:
1436:
1427:
1426:
1383:
1382:
1372:
1355:(6): 2832–2840.
1340:
1334:
1333:
1314:10.1038/35099006
1297:
1280:
1279:
1269:
1245:
1239:
1238:
1228:
1196:
1190:
1189:
1179:
1147:
1141:
1140:
1103:
1097:
1096:
1060:
1054:
1053:
1017:
1011:
1010:
1000:
967:
961:
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950:
918:
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911:
883:
877:
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866:
834:
828:
827:
783:
777:
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766:
742:
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691:
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632:
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519:
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512:
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478:
449:
448:
436:
425:
424:
388:
322:Escherichia coli
150:
149:
84:
72:
71:
2736:
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2731:
2730:
2729:
2727:
2726:
2725:
2701:
2700:
2699:
2687:
2679:
2677:
2672:
2584:Oxidoreductases
2570:
2546:Enzyme kinetics
2534:
2530:List of enzymes
2503:
2472:
2443:Catalytic triad
2421:
2416:
2386:
2377:
2348:
2326:
2301:
2272:
2266:
2262:2.7.10-2.7.13:
2253:
2237:
2204:: miscellaneous
2192:
2166:
2148:
2125:
2108:exoribonuclease
2105:
2093:
2079:Polyadenylation
1996:
1822:
1816:
1804:
1786:
1782:
1776:
1760:
1722:
1709:
1686:
1658:
1488:
1482:
1474:
1468:
1454:
1391:
1386:
1341:
1337:
1308:(11): 806–814.
1298:
1283:
1246:
1242:
1197:
1193:
1148:
1144:
1109:Nature Genetics
1104:
1100:
1077:10.1038/nrc2916
1071:(10): 696–706.
1061:
1057:
1034:10.1038/nrg1638
1018:
1014:
968:
964:
919:
915:
884:
880:
849:(12): 1187–95.
835:
831:
794:(5234): 253–4.
784:
780:
743:
739:
692:
683:
646:
635:
596:
589:
574:
570:
539:(13): 4207–17.
520:
516:
493:(5): 1199–206.
479:
452:
437:
428:
389:
385:
381:
313:
301:
87:
70:
68:Transposase Tn5
17:
12:
11:
5:
2734:
2724:
2723:
2718:
2713:
2698:
2697:
2674:
2673:
2671:
2670:
2657:
2644:
2631:
2618:
2605:
2592:
2578:
2576:
2572:
2571:
2569:
2568:
2563:
2558:
2553:
2548:
2542:
2540:
2536:
2535:
2533:
2532:
2527:
2522:
2517:
2511:
2509:
2508:Classification
2505:
2504:
2502:
2501:
2496:
2491:
2486:
2480:
2478:
2474:
2473:
2471:
2470:
2465:
2460:
2455:
2450:
2445:
2440:
2435:
2429:
2427:
2423:
2422:
2415:
2414:
2407:
2400:
2392:
2383:
2382:
2379:
2378:
2376:
2375:
2370:
2365:
2359:
2357:
2350:
2349:
2347:
2346:
2337:
2335:
2328:
2327:
2325:
2324:
2315:
2313:
2303:
2302:
2300:
2299:
2290:
2288:
2275:
2270:
2264:protein kinase
2259:
2258:
2255:
2254:
2252:
2251:
2245:
2243:
2239:
2238:
2236:
2235:
2230:
2225:
2220:
2214:
2212:
2205:
2198:
2197:
2194:
2193:
2191:
2190:
2185:
2174:
2172:
2168:
2167:
2165:
2164:
2158:
2156:
2150:
2149:
2147:
2146:
2141:
2135:
2133:
2127:
2126:
2124:
2123:
2118:
2112:
2110:
2103:Phosphorolytic
2099:
2098:
2095:
2094:
2092:
2091:
2086:
2081:
2076:
2071:
2070:
2069:
2064:
2059:
2049:
2048:
2047:
2037:
2032:
2027:
2022:
2017:
2012:
2006:
2004:
2002:RNA polymerase
1998:
1997:
1995:
1994:
1993:
1992:
1982:
1978:
1977:
1976:
1975:
1970:
1965:
1954:
1953:
1952:
1947:
1942:
1937:
1926:
1920:
1919:
1918:
1911:
1906:
1901:
1896:
1885:
1884:
1883:
1876:
1871:
1866:
1861:
1850:
1846:
1844:
1842:DNA polymerase
1835:
1826:
1820:
1806:
1805:
1803:
1802:
1797:
1791:
1789:
1784:
1780:
1766:
1765:
1762:
1761:
1759:
1758:
1753:
1748:
1743:
1738:
1733:
1727:
1725:
1720:
1711:
1710:
1708:
1707:
1701:
1699:
1688:
1687:
1685:
1684:
1679:
1673:
1671:
1660:
1659:
1657:
1656:
1651:
1646:
1645:
1644:
1639:
1629:
1627:Diacylglycerol
1624:
1619:
1614:
1609:
1604:
1599:
1594:
1589:
1588:
1587:
1577:
1572:
1567:
1566:
1565:
1560:
1555:
1550:
1545:
1538:Phosphofructo-
1535:
1530:
1529:
1528:
1518:
1513:
1507:
1505:
1491:
1486:
1470:
1469:
1453:
1452:
1445:
1438:
1430:
1424:
1423:
1402:
1390:
1389:External links
1387:
1385:
1384:
1335:
1281:
1260:(1): 129–134.
1240:
1211:(3): 800–810.
1191:
1142:
1121:10.1038/ng.343
1098:
1055:
1012:
983:(R1): R14-20.
962:
913:
878:
829:
778:
757:(4): 501–510.
737:
700:Genome Biology
681:
633:
612:10.1038/nsb778
587:
568:
514:
450:
426:
399:(4): 1153–63.
382:
380:
377:
312:
311:Tn7 transposon
309:
300:
297:
209:
208:
205:
204:
199:
193:
192:
179:
173:
172:
162:
155:
154:
146:
145:
132:
126:
125:
120:
114:
113:
108:
102:
101:
98:
94:
93:
89:
88:
85:
77:
76:
69:
66:
37:Tn3 transposon
15:
9:
6:
4:
3:
2:
2733:
2722:
2719:
2717:
2714:
2712:
2709:
2708:
2706:
2696:
2691:
2686:
2685:
2682:
2668:
2664:
2663:
2658:
2655:
2651:
2650:
2645:
2642:
2638:
2637:
2632:
2629:
2625:
2624:
2619:
2616:
2612:
2611:
2606:
2603:
2599:
2598:
2593:
2590:
2586:
2585:
2580:
2579:
2577:
2573:
2567:
2564:
2562:
2559:
2557:
2554:
2552:
2549:
2547:
2544:
2543:
2541:
2537:
2531:
2528:
2526:
2525:Enzyme family
2523:
2521:
2518:
2516:
2513:
2512:
2510:
2506:
2500:
2497:
2495:
2492:
2490:
2489:Cooperativity
2487:
2485:
2482:
2481:
2479:
2475:
2469:
2466:
2464:
2461:
2459:
2456:
2454:
2451:
2449:
2448:Oxyanion hole
2446:
2444:
2441:
2439:
2436:
2434:
2431:
2430:
2428:
2424:
2420:
2413:
2408:
2406:
2401:
2399:
2394:
2393:
2390:
2374:
2371:
2369:
2366:
2364:
2361:
2360:
2358:
2355:
2351:
2345:
2344:
2339:
2338:
2336:
2333:
2329:
2323:
2322:
2317:
2316:
2314:
2312:
2308:
2304:
2298:
2297:
2292:
2291:
2289:
2287:
2283:
2279:
2276:
2273:
2265:
2260:
2250:
2247:
2246:
2244:
2240:
2234:
2231:
2229:
2226:
2224:
2221:
2219:
2216:
2215:
2213:
2209:
2206:
2203:
2199:
2189:
2186:
2183:
2179:
2176:
2175:
2173:
2169:
2163:
2160:
2159:
2157:
2155:
2151:
2145:
2142:
2140:
2137:
2136:
2134:
2132:
2128:
2122:
2119:
2117:
2114:
2113:
2111:
2109:
2104:
2100:
2090:
2087:
2085:
2082:
2080:
2077:
2075:
2072:
2068:
2065:
2063:
2060:
2058:
2055:
2054:
2053:
2050:
2046:
2043:
2042:
2041:
2038:
2036:
2033:
2031:
2028:
2026:
2023:
2021:
2018:
2016:
2013:
2011:
2008:
2007:
2005:
2003:
1999:
1991:
1988:
1987:
1986:
1983:
1980:
1979:
1974:
1971:
1969:
1966:
1964:
1961:
1960:
1958:
1955:
1951:
1948:
1946:
1943:
1941:
1938:
1936:
1933:
1932:
1930:
1927:
1924:
1921:
1917:
1916:
1912:
1910:
1907:
1905:
1902:
1900:
1897:
1895:
1892:
1891:
1889:
1886:
1882:
1881:
1877:
1875:
1872:
1870:
1867:
1865:
1862:
1860:
1857:
1856:
1854:
1851:
1848:
1847:
1845:
1843:
1839:
1836:
1834:
1830:
1827:
1824:
1815:
1811:
1807:
1801:
1798:
1796:
1793:
1792:
1790:
1787:
1775:
1771:
1767:
1757:
1754:
1752:
1749:
1747:
1744:
1742:
1739:
1737:
1734:
1732:
1729:
1728:
1726:
1723:
1716:
1712:
1706:
1703:
1702:
1700:
1697:
1693:
1689:
1683:
1680:
1678:
1675:
1674:
1672:
1669:
1665:
1661:
1655:
1652:
1650:
1647:
1643:
1642:Class II PI 3
1640:
1638:
1635:
1634:
1633:
1630:
1628:
1625:
1623:
1620:
1618:
1617:Deoxycytidine
1615:
1613:
1610:
1608:
1605:
1603:
1600:
1598:
1595:
1593:
1590:
1586:
1585:ADP-thymidine
1583:
1582:
1581:
1578:
1576:
1573:
1571:
1568:
1564:
1561:
1559:
1556:
1554:
1551:
1549:
1546:
1544:
1541:
1540:
1539:
1536:
1534:
1531:
1527:
1524:
1523:
1522:
1519:
1517:
1514:
1512:
1509:
1508:
1506:
1503:
1499:
1495:
1492:
1489:
1481:
1477:
1471:
1466:
1462:
1458:
1451:
1446:
1444:
1439:
1437:
1432:
1431:
1428:
1421:
1417:
1416:
1411:
1407:
1403:
1400:
1396:
1393:
1392:
1380:
1376:
1371:
1366:
1362:
1358:
1354:
1350:
1346:
1339:
1331:
1327:
1323:
1319:
1315:
1311:
1307:
1303:
1296:
1294:
1292:
1290:
1288:
1286:
1277:
1273:
1268:
1263:
1259:
1255:
1251:
1244:
1236:
1232:
1227:
1222:
1218:
1214:
1210:
1206:
1202:
1195:
1187:
1183:
1178:
1173:
1169:
1165:
1162:(6): 1200–9.
1161:
1157:
1153:
1146:
1138:
1134:
1130:
1126:
1122:
1118:
1115:(6): 753–61.
1114:
1110:
1102:
1094:
1090:
1086:
1082:
1078:
1074:
1070:
1066:
1059:
1051:
1047:
1043:
1039:
1035:
1031:
1028:(7): 568–80.
1027:
1023:
1016:
1008:
1004:
999:
994:
990:
986:
982:
978:
974:
966:
958:
954:
949:
944:
940:
936:
933:(9): 756–67.
932:
928:
924:
917:
909:
905:
901:
897:
893:
889:
882:
874:
870:
865:
860:
856:
852:
848:
844:
840:
833:
825:
821:
817:
813:
809:
805:
801:
797:
793:
789:
782:
774:
770:
765:
760:
756:
752:
748:
741:
733:
729:
724:
719:
714:
709:
705:
701:
697:
690:
688:
686:
677:
673:
668:
663:
660:(3): 1904–9.
659:
655:
651:
644:
642:
640:
638:
629:
625:
621:
617:
613:
609:
606:(4): 278–81.
605:
601:
594:
592:
583:
579:
578:"Transposase"
572:
564:
560:
555:
550:
546:
542:
538:
534:
530:
524:
518:
510:
506:
501:
496:
492:
488:
484:
477:
475:
473:
471:
469:
467:
465:
463:
461:
459:
457:
455:
446:
442:
441:"Transposase"
435:
433:
431:
422:
418:
414:
410:
406:
402:
398:
394:
387:
383:
376:
373:
368:
362:
360:
356:
350:
348:
344:
340:
336:
332:
328:
324:
323:
318:
308:
306:
296:
294:
289:
283:
281:
276:
274:
270:
265:
261:
254:
250:
246:
244:
238:
236:
232:
231:
226:
225:
220:
216:
203:
200:
198:
194:
191:
187:
183:
180:
178:
174:
170:
166:
163:
160:
156:
151:
147:
144:
140:
136:
133:
131:
127:
124:
121:
119:
115:
112:
109:
107:
103:
100:Dimer_Tnp_Tn5
99:
95:
90:
83:
78:
73:
65:
62:
57:
55:
50:
46:
42:
38:
34:
30:
26:
22:
2662:Translocases
2659:
2646:
2633:
2620:
2607:
2597:Transferases
2594:
2581:
2438:Binding site
2340:
2318:
2293:
2187:
1914:
1879:
1637:Class I PI 3
1602:Pantothenate
1473:2.7.1-2.7.4:
1457:Transferases
1413:
1395:Transposases
1352:
1348:
1338:
1305:
1301:
1257:
1253:
1243:
1208:
1204:
1194:
1159:
1155:
1145:
1112:
1108:
1101:
1068:
1064:
1058:
1025:
1021:
1015:
980:
976:
965:
930:
926:
916:
891:
887:
881:
846:
842:
832:
791:
787:
781:
754:
750:
740:
706:(12): R119.
703:
699:
657:
653:
603:
599:
581:
576:McDowall J.
571:
536:
532:
522:
517:
490:
486:
444:
396:
392:
386:
363:
351:
339:streptomycin
335:trimethoprim
326:
320:
314:
302:
284:
277:
272:
268:
266:
262:
258:
247:
239:
228:
222:
212:
58:
20:
18:
2433:Active site
2188:Transposase
2178:Recombinase
1823:-nucleoside
1649:Sphingosine
894:(1): 8–11.
295:technique.
230:Escherichia
92:Identifiers
49:chromosomes
21:transposase
2705:Categories
2636:Isomerases
2610:Hydrolases
2477:Regulation
1990:Telomerase
1833:Polymerase
1607:Mevalonate
1570:Riboflavin
1461:phosphorus
379:References
367:activators
224:Shewanella
219:integrases
165:structures
52:can cause
29:transposon
2515:EC number
2182:Integrase
2106:3' to 5'
1751:Guanylate
1746:Uridylate
1736:Adenylate
1580:Thymidine
1575:Shikimate
927:BioEssays
372:regulator
359:cassettes
235:kanamycin
123:IPR003201
61:EC number
2711:EC 2.7.7
2539:Kinetics
2463:Cofactor
2426:Activity
2116:RNase PH
1724:acceptor
1705:Creatine
1698:acceptor
1670:acceptor
1612:Pyruvate
1597:Glycerol
1558:Platelet
1533:Galacto-
1504:acceptor
1330:34733892
1322:11715047
1186:20372108
1137:27373372
1129:19412179
1085:20844553
1042:15995698
1007:21459777
957:20652893
908:15680506
873:20615441
824:29930180
732:21143862
676:12424243
620:11896402
582:InterPro
563:20215432
509:12603728
421:17775137
355:integron
347:plasmids
331:plasmids
293:ATAC-seq
269:in vitro
182:RCSB PDB
118:InterPro
2695:Biology
2649:Ligases
2419:Enzymes
2067:PrimPol
2052:Primase
1526:Hepatic
1521:Fructo-
1420:PDBe-KB
1410:UniProt
1379:2836374
1177:2889740
1093:6910577
1050:3194633
998:3095056
948:3971908
864:2991504
816:7569973
796:Bibcode
788:Science
773:9390559
723:3046479
628:9721663
554:2910039
327:E. coli
273:in vivo
111:PF02281
25:enzymes
2681:Portal
2623:Lyases
2354:2.7.13
2332:2.7.12
2307:2.7.11
2282:2.7.10
2121:PNPase
2089:PNPase
2045:POLRMT
2040:ssRNAP
1553:Muscle
1516:Gluco-
1480:kinase
1415:Q46731
1401:(MeSH)
1377:
1370:211210
1367:
1328:
1320:
1276:915473
1274:
1235:767328
1233:
1226:236152
1223:
1184:
1174:
1135:
1127:
1091:
1083:
1048:
1040:
1005:
995:
955:
945:
906:
871:
861:
822:
814:
771:
730:
720:
674:
626:
618:
561:
551:
507:
419:
413:391406
411:
197:PDBsum
171:
161:
143:SUPFAM
97:Symbol
33:genome
2575:Types
2202:2.7.8
2171:Other
1810:2.7.7
1770:2.7.6
1715:2.7.4
1692:2.7.3
1664:2.7.2
1548:Liver
1511:Hexo-
1498:2.7.1
1326:S2CID
1133:S2CID
1089:S2CID
1046:S2CID
820:S2CID
624:S2CID
417:S2CID
215:RNase
139:SCOPe
130:SCOP2
45:maize
2667:list
2660:EC7
2654:list
2647:EC6
2641:list
2634:EC5
2628:list
2621:EC4
2615:list
2608:EC3
2602:list
2595:EC2
2589:list
2582:EC1
2341:see
2319:see
2294:see
1668:COOH
1467:2.7)
1408:for
1375:PMID
1318:PMID
1272:PMID
1231:PMID
1182:PMID
1125:PMID
1081:PMID
1038:PMID
1003:PMID
953:PMID
904:PMID
869:PMID
812:PMID
769:PMID
751:Cell
728:PMID
672:PMID
616:PMID
559:PMID
505:PMID
409:PMID
393:Cell
337:and
227:and
190:PDBj
186:PDBe
169:ECOD
159:Pfam
135:1b7e
106:Pfam
2084:PAP
2025:III
1959:/Y
1950:TDT
1931:/X
1923:III
1915:Pfu
1890:/B
1880:Taq
1855:/A
1622:PFP
1592:NAD
1406:PDB
1365:PMC
1357:doi
1353:170
1310:doi
1262:doi
1258:102
1221:PMC
1213:doi
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