91:) usually use DNA primers, since they are more temperature stable. Primers can be designed in laboratory for specific reactions such as polymerase chain reaction (PCR). When designing PCR primers, there are specific measures that must be taken into consideration, like the melting temperature of the primers and the annealing temperature of the reaction itself. Moreover, the DNA binding sequence of the primer in vitro has to be specifically chosen, which is done using a method called basic local alignment search tool (BLAST) that scans the DNA and finds specific and unique regions for the primer to bind.
261:, in eukaryotes itβs known as the RNase H2. This enzyme degrades most of the annealed RNA primer, except the nucleotides close to the 5β end of the primer. Thus, the remaining nucleotides are displayed into a flap that is cleaved off using FEN-1. The last possible method of removing RNA primer is known as the long flap pathway. In this pathway several enzymes are recruited to elongate the RNA primer and then cleave it off. The flaps are elongated by a 5β to 3β
2110:
27:
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360:(PCR) uses a pair of custom primers to direct DNA elongation toward each other at opposite ends of the sequence being amplified. These primers are typically between 18 and 24 bases in length and must code for only the specific upstream and downstream sites of the sequence being amplified. A primer that can bind to multiple regions along the DNA will amplify them all, eliminating the purpose of PCR.
71:
using an enzyme called ligase. The removal process of the RNA primer requires several enzymes, such as Fen1, Lig1, and others that work in coordination with DNA polymerase, to ensure the removal of the RNA nucleotides and the addition of DNA nucleotides. Living organisms use solely RNA primers, while laboratory techniques in
413:
Selecting a specific region of DNA for primer binding requires some additional considerations. Regions high in mononucleotide and dinucleotide repeats should be avoided, as loop formation can occur and contribute to mishybridization. Primers should not easily anneal with other primers in the mixture;
363:
A few criteria must be brought into consideration when designing a pair of PCR primers. Pairs of primers should have similar melting temperatures since annealing during PCR occurs for both strands simultaneously, and this shared melting temperature must not be either too much higher or lower than the
252:
are formed, which are discontinuous strands of DNA. Then, when the DNA polymerase reaches to the 5β end of the RNA primer from the previous
Okazaki fragment, it displaces the 5β² end of the primer into a single-stranded RNA flap which is removed by nuclease cleavage. Cleavage of the RNA flaps involves
70:
before DNA polymerase can begin a complementary strand. DNA polymerase adds nucleotides after binding to the RNA primer and synthesizes the whole strand. Later, the RNA strands must be removed accurately and replace them with DNA nucleotides forming a gap region known as a nick that is filled in
405:
As of 2014, many online tools are freely available for primer design, some of which focus on specific applications of PCR. Primers with high specificity for a subset of DNA templates in the presence of many similar variants can be designed using by some software (e.g.
414:
this phenomenon can lead to the production of 'primer dimer' products contaminating the end solution. Primers should also not anneal strongly to themselves, as internal hairpins and loops could hinder the annealing with the template DNA.
277:, which has a helicase-nuclease activity, that cleaves the long flap of RNA primer, which then leaves behind a couple of nucleotides that are cleaved by FEN1. At the end, when all the RNA primers have been removed, nicks form between the
339:
before being extended by DNA polymerase. The ability to create and customize synthetic primers has proven an invaluable tool necessary to a variety of molecular biological approaches involving the analysis of DNA. Both the
417:
When designing primers, additional nucleotide bases can be added to the back ends of each primer, resulting in a customized cap sequence on each end of the amplified region. One application for this practice is for use in
177:. Reverse transcriptase is an enzyme that uses a template strand of RNA to synthesize a complementary strand of DNA. The DNA polymerase component of reverse transcriptase requires an existing 3' end to begin synthesis.
224:
direction, and polymerase I can do these activities simultaneously; this is known as βNick
Translationβ. Nick translation refers to the synchronized activity of polymerase I in removing the RNA primer and adding
385:
Additionally, primer sequences need to be chosen to uniquely select for a region of DNA, avoiding the possibility of hybridization to a similar sequence nearby. A commonly used method for selecting a primer site is
525:. Differences among sequences are accounted for by using IUPAC degeneracies for individual bases. PCR primers are then synthesized as a mixture of primers corresponding to all permutations of the codon sequence.
464:. This allows different organisms to have a significantly different genetic sequence that code for a highly similar protein. For this reason, degenerate primers are also used when primer design is based on
257:(FEN-1), which cleaves the 5β overhanging flap. This method is known as the short flap pathway of RNA primer removal. The second way to cleave a RNA primer is by degrading the RNA strand using a
889:
Adenosine added on the primer 50 end improved TA cloning efficiency of polymerase chain reaction products, Ri-He Peng, Ai-Sheng Xiong, Jin-ge Liu, Fang Xu, Cai Bin, Hong Zhu, Quan-Hong Yao
2126:
162:
cannot add bases in the 3β²β5β² direction complementary to the template strand, DNA is synthesized βbackwardβ in short fragments moving away from the replication fork, known as
390:
search, whereby all the possible regions to which a primer may bind can be seen. Both the nucleotide sequence as well as the primer itself can be BLAST searched. The free
521:
or allow the recovery of genes from organisms where genomic information is not available. Usually, degenerate primers are designed by aligning gene sequencing found in
166:. Unlike in the leading strand, this method results in the repeated starting and stopping of DNA synthesis, requiring multiple RNA primers. Along the DNA template,
273:(RPA). The RPA-bound DNA inhibits the activity or recruitment of FEN1, as a result another nuclease must be recruited to cleave the flap. This second nuclease is
375:(melting temperature) too much higher than the reaction's annealing temperature may mishybridize and extend at an incorrect location along the DNA sequence. A
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In prokaryotes, DNA polymerase I synthesizes the
Okazaki fragment until it reaches the previous RNA primer. Then the enzyme simultaneously acts as a
663:
391:
507:. Degenerate primers may not perfectly hybridize with a target sequence, which can greatly reduce the specificity of the PCR amplification.
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394:
tool Primer-BLAST integrates primer design and BLAST search into one application, as do commercial software products such as ePrime and
142:. Starting from the free 3β-OH of the primer, known as the primer terminus, a DNA polymerase can extend a newly synthesized strand. The
934:
422:, a special subcloning technique similar to PCR, where efficiency can be increased by adding AG tails to the 5β² and the 3β² ends.
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three methods of primer removal. The first possibility of primer removal is by creating a short flap that is directly removed by
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849:"New Sets of Primers for DNA Identification of Non-Indigenous Fish Species in the Volga-Kama Basin (European Russia)"
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These are mixtures of primers that are similar, but not identical. These may be convenient when amplifying the same
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1969:
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847:
Karabanov, D.P.; Bekker, E.I.; Pavlov, D.D.; Borovikova, E.A.; Kodukhova, Y.V.; Kotov, A.A. (1 February 2022).
154:, requiring only an initial RNA primer to begin synthesis. In the lagging strand, the template DNA runs in the
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468:, as the specific sequence of codons are not known. Therefore, primer sequence corresponding to the
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402:) may be performed to assist in primer design by giving melting and annealing temperatures, etc.
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is essential for the completion of replication. Thus, as the lagging strand being synthesized by
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intersperses RNA primers that DNA polymerase uses to synthesize DNA from in the 5β²β3β² direction.
20:
593:"In vitro reconstitution of RNA primer removal in Archaea reveals the existence of two pathways"
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then joins the fragmented strands together, completing the synthesis of the lagging strand.
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behind. Both the activities of polymerization and excision of the RNA primer occur in the
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significantly lower than the annealing temperature may fail to anneal and extend at all.
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to a specific site on the template DNA. In solution, the primer spontaneously
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Diagrammatic representation of the forward and reverse primers for a standard
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Short strand of RNA or DNA that serves as a starting point for DNA synthesis
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Doudna; Cox; O'Donnell, Jennifer; Michael M.; Michael (December 21, 2016).
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570:. New York: W. H. Freeman and Company. pp. 221β238, 369β376, 592β593.
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517:. They allow for the amplification of genes from thus far uncultivated
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58:(responsible for DNA replication) enzymes are only capable of adding
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Another example of primers being used to enable DNA synthesis is
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682:"Primer removal during mammalian mitochondrial DNA replication"
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of an existing nucleic acid, requiring a primer be bound to
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add a complementary RNA primer to the reading template
806:. NCBI - National Center for Biotechnology Information
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that fill the gaps where the RNA primer was present.
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Balakrishnan, Lata; Bambara, Robert A. (2013-02-01).
537:– the methods by which primers are manufactured
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are widely used and extremely useful in the field of
30:
The DNA replication fork. RNA primer labeled at top.
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398:. Computer simulations of theoretical PCR results (
323:Synthetic primers, sometimes known as oligos, are
42:used by all living organisms in the initiation of
2162:
236:In eukaryotes the removal of RNA primers in the
112:RNA primers are used by living organisms in the
680:Uhler, Jay P.; Falkenberg, Maria (2015-10-01).
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316:. For possible methods involving primers, see
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327:, usually of DNA, which can be customized to
662:: CS1 maint: multiple names: authors list (
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738:Cold Spring Harbor Perspectives in Biology
640:Molecular Biology: Principles and practice
568:Molecular Biology: Principles and Practice
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436:Some situations may call for the use of
312:For the organic chemistry involved, see
300:
150:in one continuous piece moving with the
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325:chemically synthesized oligonucleotides
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255:flap structure-specific endonuclease 1
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50:primer may also be referred to as an
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475:might be "ATH", where A stands for
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1965:Post-transcriptional modification
1591:Control of chromosome duplication
1157:Autonomously replicating sequence
896:
828:. Wellcome Trust Sanger Institute
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670:
591:Henneke, Ghislaine (2012-09-26).
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1970:Post-translational modification
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342:Sanger chain termination method
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503:, using the IUPAC symbols for
1:
2092:Post-translational regulation
1314:DNA polymerase III holoenzyme
1164:Single-strand binding protein
734:"Okazaki fragment metabolism"
541:
54:, short for oligonucleotide.
2040:High-throughput technique ("
699:10.1016/j.dnarep.2015.07.003
456:, meaning several different
124:. A class of enzymes called
38:is a short, single-stranded
7:
1918:Functional biology/medicine
750:10.1101/cshperspect.a010173
528:
289:, through a process called
10:
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1410:Prokaryotic DNA polymerase
1111:Minichromosome maintenance
1058:Origin recognition complex
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335:with the template through
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2181:Polymerase chain reaction
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535:Oligonucleotide synthesis
358:polymerase chain reaction
337:Watson-Crick base pairing
314:Oligonucleotide synthesis
297:Uses of synthetic primers
285:using an enzyme known as
89:polymerase chain reaction
566:Cox, Michael M. (2015).
281:that are filled-in with
197:) and replaced with new
1051:Pre-replication complex
982:Pre-replication complex
185:After the insertion of
83:DNA synthesis (such as
460:can code for the same
309:
146:in DNA replication is
31:
1913:Developmental biology
1908:Computational biology
1474:Replication protein A
1243:Origin of replication
366:annealing temperature
304:
271:replication protein A
175:reverse transcription
102:Further information:
29:
2087:Post-transcriptional
1445:Replication factor C
318:Nucleic acid methods
283:deoxyribonucleotides
227:deoxyribonucleotides
218:deoxyribonucleotides
216:in front and adding
199:deoxyribonucleotides
19:For other uses, see
1882:Histone methylation
597:Biochemical Journal
495:, according to the
438:degenerate primers.
609:10.1042/BJ20120959
511:Degenerate primers
426:Degenerate primers
368:. A primer with a
310:
212:, removing primer
32:
2176:Molecular biology
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2115:Molecular biology
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2054:Mass spectrometry
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1698:Molecular biology
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1621:
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1457:Flap endonuclease
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1286:
1274:Okazaki fragments
868:10.3390/w14030437
642:. W. H. Freeman.
515:microbial ecology
352:PCR primer design
279:Okazaki fragments
250:Okazaki fragments
210:5β²β3β² exonuclease
187:Okazaki fragments
164:Okazaki fragments
77:molecular biology
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1150:Licensing factor
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826:"About DECIPHER"
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804:"Electronic PCR"
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242:DNA polymerase Ξ΄
152:replication fork
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790:"Primer-BLAST"
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744:(2): a010173.
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430:Main article:
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400:Electronic PCR
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238:lagging strand
182:
181:Primer removal
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160:DNA polymerase
144:leading strand
104:DNA polymerase
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85:DNA sequencing
56:DNA polymerase
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2066:
2060:
2059:Lab-on-a-chip
2057:
2055:
2052:
2050:
2047:
2043:
2038:
2037:
2034:
2033:Radioactivity
2030:
2026:
2023:
2019:
2016:
2014:
2011:
2010:
2008:
2005:
2001:
1998:
1996:
1993:
1992:
1990:
1986:
1980:
1976:
1973:
1971:
1968:
1966:
1963:
1961:
1958:
1956:
1953:
1951:
1950:Cultured meat
1948:
1947:
1945:
1941:
1938:
1934:
1924:
1921:
1919:
1916:
1914:
1911:
1909:
1906:
1904:
1901:
1899:
1896:
1895:
1893:
1889:
1883:
1880:
1878:
1875:
1871:
1870:trp repressor
1868:
1866:
1865:lac repressor
1863:
1862:
1861:
1858:
1856:
1853:
1851:
1848:
1846:
1843:
1841:
1838:
1834:
1831:
1829:
1826:
1824:
1821:
1820:
1819:
1816:
1812:
1809:
1807:
1804:
1803:
1802:
1799:
1798:
1790:
1785:
1784:
1783:
1782:
1780:
1776:
1769:
1765:
1762:
1759:
1755:
1754:Transcription
1752:
1749:
1745:
1742:
1741:
1739:
1737:
1736:Central dogma
1733:
1730:
1726:
1720:
1717:
1715:
1712:
1710:
1707:
1706:
1703:
1699:
1692:
1687:
1685:
1680:
1678:
1673:
1672:
1669:
1655:
1652:
1650:
1647:
1645:
1642:
1641:
1640:
1636:
1633:
1632:
1630:
1628:
1624:
1614:
1611:
1609:
1605:
1604:
1602:
1598:
1592:
1589:
1588:
1583:
1580:
1579:
1578:
1575:
1574:
1569:
1566:
1564:
1561:
1559:
1556:
1554:
1551:
1550:
1549:
1546:
1542:
1539:
1537:
1534:
1532:
1529:
1527:
1524:
1523:
1522:
1519:
1515:
1512:
1510:
1507:
1505:
1502:
1500:
1497:
1496:
1495:
1492:
1489:
1486:
1485:
1480:
1477:
1476:
1475:
1472:
1470:
1469:Topoisomerase
1467:
1463:
1460:
1459:
1458:
1455:
1451:
1448:
1447:
1446:
1443:
1442:
1440:
1437:
1428:
1420:
1417:
1416:
1415:
1411:
1408:
1407:
1402:
1399:
1398:
1397:
1396:Topoisomerase
1394:
1392:
1389:
1387:
1384:
1382:
1379:
1375:
1372:
1370:
1367:
1365:
1362:
1360:
1357:
1355:
1352:
1350:
1347:
1345:
1342:
1340:
1337:
1335:
1332:
1330:
1327:
1325:
1322:
1320:
1317:
1316:
1315:
1312:
1311:
1309:
1306:
1299:
1296:
1294:
1290:
1280:
1277:
1275:
1272:
1268:
1264:
1261:
1260:
1259:
1256:
1255:
1252:
1248:
1244:
1241:
1240:
1238:
1234:
1228:
1225:
1223:
1219:
1216:
1215:
1212:
1208:
1205:
1204:
1199:
1196:
1194:
1191:
1190:
1189:
1186:
1185:
1180:
1177:
1175:
1172:
1170:
1167:
1166:
1165:
1162:
1161:
1158:
1155:
1154:
1151:
1148:
1147:
1142:
1139:
1137:
1134:
1132:
1129:
1127:
1124:
1122:
1119:
1117:
1114:
1113:
1112:
1109:
1108:
1105:
1102:
1101:
1098:
1095:
1094:
1089:
1086:
1084:
1081:
1079:
1076:
1074:
1071:
1069:
1066:
1064:
1061:
1060:
1059:
1056:
1055:
1052:
1049:
1048:
1046:
1043:
1034:
1026:
1023:
1022:
1021:
1018:
1017:
1012:
1009:
1007:
1004:
1002:
999:
998:
997:
994:
993:
990:
987:
986:
983:
980:
979:
977:
974:
967:
964:
962:
958:
953:
949:
945:
938:
933:
931:
926:
924:
919:
918:
915:
909:
906:
904:
901:
900:
886:
878:
874:
869:
864:
860:
856:
855:
850:
843:
827:
821:
805:
799:
791:
785:
777:
773:
768:
763:
759:
755:
751:
747:
743:
739:
735:
728:
726:
717:
713:
709:
705:
700:
695:
691:
687:
683:
676:
674:
665:
659:
651:
649:9781319116378
645:
641:
634:
626:
622:
618:
614:
610:
606:
602:
598:
594:
587:
579:
577:9781464126147
573:
569:
562:
560:
558:
556:
554:
552:
547:
536:
533:
532:
526:
524:
520:
516:
512:
508:
506:
502:
498:
494:
490:
486:
482:
478:
474:
471:
467:
463:
459:
455:
451:
447:
443:
439:
433:
423:
421:
415:
411:
409:
403:
401:
397:
393:
389:
383:
378:
371:
367:
361:
359:
349:
347:
343:
338:
334:
330:
326:
319:
315:
308:
303:
294:
292:
288:
284:
280:
276:
275:DNA2 nuclease
272:
268:
264:
260:
256:
251:
247:
243:
239:
234:
232:
228:
223:
219:
215:
211:
206:
204:
200:
196:
192:
188:
178:
176:
171:
169:
165:
161:
157:
153:
149:
145:
141:
137:
133:
132:
127:
123:
119:
115:
109:
105:
98:
92:
90:
86:
82:
79:that require
78:
74:
69:
65:
61:
57:
53:
49:
45:
44:DNA synthesis
41:
37:
28:
22:
2125:
2113:
2025:Fluorescence
2013:Nucleic acid
2004:C57BL/6 mice
1995:Cell culture
1903:Biochemistry
1898:Cell biology
1608:Processivity
1434:synthesis in
1278:
908:Primer-BLAST
885:
858:
852:
842:
830:. Retrieved
820:
808:. Retrieved
798:
784:
741:
737:
689:
685:
639:
633:
600:
596:
586:
567:
510:
509:
497:genetic code
483:, and H for
450:genetic code
437:
435:
416:
412:
404:
384:
376:
369:
362:
355:
322:
235:
207:
184:
172:
134:on both the
129:
120:a strand of
118:synthesizing
111:
96:
95:RNA primers
73:biochemistry
68:the template
51:
40:nucleic acid
35:
33:
2127:WikiProject
1936:Engineering
1891:Linked life
1806:Pribnow box
1764:Translation
1627:Termination
1301:Prokaryotic
1293:Replication
969:Prokaryotic
948:prokaryotic
946:(comparing
832:12 February
364:reaction's
265:, known as
248:direction,
191:prokaryotes
148:synthesized
60:nucleotides
2165:Categories
2077:Epigenetic
1988:Techniques
1850:Terminator
1833:trp operon
1828:lac operon
1823:gal operon
1639:Telomerase
1613:DNA ligase
1606:Movement:
1430:Eukaryotic
1401:DNA gyrase
1386:DNA ligase
1305:elongation
1036:Eukaryotic
973:initiation
961:Initiation
952:eukaryotic
861:(3): 437.
686:DNA Repair
542:References
473:isoleucine
470:amino acid
462:amino acid
454:degenerate
452:itself is
420:TA cloning
333:hybridizes
231:DNA ligase
203:DNA ligase
195:eukaryotes
114:initiation
2002:(such as
1860:Repressor
1577:DNA clamp
1391:DNA clamp
1381:Replisome
877:2073-4441
758:1943-0264
708:1568-7864
692:: 28β38.
658:cite book
617:0264-6021
499:for each
446:organisms
344:and the β
48:synthetic
2009:Methods
1943:Concepts
1923:Genetics
1877:Silencer
1855:Enhancer
1811:TATA box
1801:Promoter
1792:Heredity
1728:Overview
1719:Glossary
1635:Telomere
1251:Replicon
1207:Helicase
1198:RNASEH2A
1042:G1 phase
996:Helicase
810:13 March
776:23378587
716:26303841
625:22849643
529:See also
493:cytosine
479:, T for
408:DECIPHER
346:Next-Gen
291:ligation
263:helicase
158:. Since
126:primases
81:in vitro
2155:Biology
2082:Genetic
2029:Pigment
2018:Protein
1979:Wet lab
1975:Dry lab
1955:Mitosis
1787:Genetic
1778:Element
1768:protein
1709:History
1548:epsilon
1436:S phase
1263:Lagging
1218:Primase
1193:RNASEH1
1188:RNase H
1020:Primase
903:Primer3
767:3552508
523:GenBank
489:thymine
485:adenine
481:thymine
477:adenine
287:ligase1
168:primase
136:leading
131:de novo
97:in vivo
62:to the
2141:Portal
2042:-omics
2031:&
1840:Intron
1818:Operon
1279:Primer
875:
774:
764:
756:
714:
706:
646:
623:
615:
574:
458:codons
329:anneal
64:3β-end
36:primer
21:Primer
1714:Index
1568:POLE4
1563:POLE3
1558:POLE2
1541:POLD4
1536:POLD3
1531:POLD2
1526:POLD1
1521:delta
1514:PRIM2
1509:PRIM1
1504:POLA2
1499:POLA1
1494:alpha
1227:PRIM2
1222:PRIM1
1179:SSBP4
1174:SSBP3
1169:SSBP2
854:Water
501:codon
491:, or
388:BLAST
259:RNase
246:5β²β3β²
222:5β²β3β²
52:oligo
1845:Exon
1654:DKC1
1649:TERC
1644:TERT
1600:Both
1582:PCNA
1553:POLE
1479:RPA1
1462:FEN1
1450:RFC1
1374:holE
1369:holD
1364:holC
1359:holB
1354:holA
1349:dnaX
1344:dnaT
1339:dnaQ
1334:dnaN
1329:dnaH
1324:dnaE
1319:dnaC
1265:and
1236:Both
1211:HFM1
1141:MCM7
1136:MCM6
1131:MCM5
1126:MCM4
1121:MCM3
1116:MCM2
1104:Cdt1
1097:Cdc6
1088:ORC6
1083:ORC5
1078:ORC4
1073:ORC3
1068:ORC2
1063:ORC1
1025:dnaG
1006:dnaB
1001:dnaA
989:dnaC
873:ISSN
834:2014
812:2012
772:PMID
754:ISSN
712:PMID
704:ISSN
664:link
644:ISBN
621:PMID
613:ISSN
572:ISBN
442:gene
392:NCBI
356:The
267:Pif1
193:and
138:and
106:and
87:and
75:and
46:. A
1758:RNA
1748:DNA
1247:Ori
950:to
863:doi
762:PMC
746:doi
694:doi
605:doi
601:447
307:PCR
244:in
122:DNA
116:of
2167::
2044:")
2027:,
1977:/
1637::
1412::
1220::
1209::
1011:T7
871:.
859:14
857:.
851:.
770:.
760:.
752:.
740:.
736:.
724:^
710:.
702:.
690:34
688:.
684:.
672:^
660:}}
656:{{
619:.
611:.
599:.
595:.
550:^
487:,
293:.
233:.
34:A
2143::
2006:)
1770:)
1766:(
1760:)
1756:(
1750:)
1746:(
1690:e
1683:t
1676:v
1490::
1438:)
1432:(
1307:)
1303:(
1249:/
1245:/
1044:)
1038:(
975:)
971:(
954:)
936:e
929:t
922:v
879:.
865::
836:.
814:.
792:.
778:.
748::
742:5
718:.
696::
666:)
652:.
627:.
607::
580:.
380:m
377:T
373:m
370:T
320:.
23:.
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