1077:
1138:
1109:
1271:
1245:
1219:
1193:
1167:
350:
411:(usually only 1) and read a completely different frame thereafter. In programmed −1 ribosomal frameshifting, the slippery sequence fits a X_XXY_YYH motif, where XXX is any three identical nucleotides (though some exceptions occur), YYY typically represents UUU or AAA, and H is A, C or U. In the case of +1 frameshifting, the slippery sequence contains codons for which the corresponding tRNA is more rare, and the frameshift is favored because the codon in the new frame has a more common associated tRNA. One example of a slippery sequence is the
308:
1063:
341:, has weaker tRNA anticodon binding specificity than the first and second nucleotides. In this model, the motif structure is explained by the fact that the first and second positions of the anticodons must be able to pair perfectly in both the 0 and −1 frames. Therefore, nucleotides 2 and 1 must be identical, and nucleotides 3 and 2 must also be identical, leading to a required sequence of 3 identical nucleotides for each tRNA that slips.
431:) is thought to pause the ribosome on the slippery site during translation, forcing it to relocate and continue replication from the −1 position. It is believed that this occurs because the structure physically blocks movement of the ribosome by becoming stuck in the ribosome mRNA tunnel. This model is supported by the fact that strength of the pseudoknot has been positively correlated with the level of frameshifting for associated mRNA.
384:
435:
and incomplete circles of mRNA represent linear regions. The secondary "stem-loop" structures, where "stems" are formed by a region of mRNA base pairing with another region on the same strand, are shown protruding from the linear DNA. The linear region of the HIV ribosomal frameshift signal contains a highly conserved UUU UUU A slippery sequence; many of the other predicted structures contain candidates for slippery sequences as well.
1007:
1021:
1035:
1049:
366:. Due to this lag, there exist in small sections of codons sequences that control the rate of ribosomal frameshifting. Specifically, the ribosome must pause to wait for the arrival of a rare tRNA, and this increases the kinetic favorability of the ribosome and its associated tRNA slipping into the new frame. In this model, the change in reading frame is caused by a single tRNA slip rather than two.
993:
337:
described by a tandem slippage model, in which the ribosomal P-site tRNA anticodon re-pairs from XXY to XXX and the A-site anticodon re-pairs from YYH to YYY simultaneously. These new pairings are identical to the 0-frame pairings except at their third positions. This difference does not significantly disfavor anticodon binding because the third nucleotide in a codon, known as the
357:
The slippery sequence for a +1 frameshift signal does not have the same motif, and instead appears to function by pausing the ribosome at a sequence encoding a rare amino acid. Ribosomes do not translate proteins at a steady rate, regardless of the sequence. Certain codons take longer to translate,
434:
Below are examples of predicted secondary structures for frameshift elements shown to stimulate frameshifting in a variety of organisms. The majority of the structures shown are stem-loops, with the exception of the ALIL (apical loop-internal loop) pseudoknot structure. In these images, the larger
438:
The mRNA sequences in the images can be read according to a set of guidelines. While A, T, C, and G represent a particular nucleotide at a position, there are also letters that represent ambiguity which are used when more than one kind of nucleotide could occur at that position. The rules of the
336:
The slippery sequence fits a X_XXY_YYH motif, where XXX is any three identical nucleotides (though some exceptions occur), YYY typically represents UUU or AAA, and H is A, C or U. Because the structure of this motif contains 2 adjacent 3-nucleotide repeats it is believed that −1 frameshifting is
1687:
Mulroney, Thomas E.; Pöyry, Tuija; Yam-Puc, Juan Carlos; Rust, Maria; Harvey, Robert F.; Kalmar, Lajos; Horner, Emily; Booth, Lucy; Ferreira, Alexander P.; Stoneley, Mark; Sawarkar, Ritwick; Mentzer, Alexander J.; Lilley, Kathryn S.; Smales, C. Mark; von der Haar, Tobias (6 December 2023).
323:
sites of the ribosome are indicated. Location of growing polypeptide chain is not indicated in image because there is not yet consensus on whether the −1 slip occurs before or after polypeptide is transferred from P-site tRNA to A-site tRNA (in this case from the Asn tRNA to the Leu tRNA).
374:
Ribosomal frameshifting may be controlled by mechanisms found in the mRNA sequence (cis-acting). This generally refers to a slippery sequence, an RNA secondary structure, or both. A −1 frameshift signal consists of both elements separated by a spacer region typically 5–9 nucleotides long.
275:
virus (flu), which all rely on frameshifting to create a proper ratio of 0-frame (normal translation) and "trans-frame" (encoded by frameshifted sequence) proteins. Its use in viruses is primarily for compacting more genetic information into a shorter amount of genetic material.
135:, meaning that a particular amino acid can be specified by more than one codon. However, a shift of any number of nucleotides that is not divisible by 3 in the reading frame will cause subsequent codons to be read differently. This effectively changes the ribosomal
1076:
353:+1 frameshift occurs as ribosome and P-site tRNA pause to wait for arrival of rare arginine tRNA. The A-site codon in the new frame pairs to anticodon of more common glycine tRNA, and translation continues.
1137:
1108:
1954:
Jagger BW, Wise HM, Kash JC, Walters KA, Wills NM, Xiao YL, Dunfee RL, Schwartzman LM, Ozinsky A, Bell GL, Dalton RM, Lo A, Efstathiou S, Atkins JF, Firth AE, Taubenberger JK, Digard P (July 2012).
295:. Other, rarer types of frameshifting include +1 and −2 frameshifting. −1 and +1 frameshifting are believed to be controlled by different mechanisms, which are discussed below. Both mechanisms are
1270:
1244:
1218:
1192:
1166:
311:
Tandem slippage of 2 tRNAs at rous sarcoma virus slippery sequence. After the frameshift, new base pairings are correct at the first and second nucleotides but incorrect at wobble position.
965:
423:
Efficient ribosomal frameshifting generally requires the presence of an RNA secondary structure to enhance the effects of the slippery sequence. The RNA structure (which can be a
328:
In −1 frameshifting, the ribosome slips back one nucleotide and continues translation in the −1 frame. There are typically three elements that comprise a −1 frameshift signal: a
263:
In viruses this phenomenon may be programmed to occur at particular sites and allows the virus to encode multiple types of proteins from the same mRNA. Notable examples include
211:
However, let's change the reading frame by starting one nucleotide downstream (effectively a "+1 frameshift" when considering the 0 position to be the initial position of
958:
1749:"Ribosomal frameshifting in decoding antizyme mRNAs from yeast and protists to humans: close to 300 cases reveal remarkable diversity despite underlying conservation"
2467:"Identification of a new antizyme mRNA +1 frameshifting stimulatory pseudoknot in a subset of diverse invertebrates and its apparent absence in intermediate species"
1478:
1392:
1068:
395:
protein-coding region, or open reading frame (ORF). Both gag and pol proteins are required for reverse transcriptase, which is essential to HIV1 replication.
1440:. Certain proteins which are needed for codon recognition or which bind directly to the mRNA sequence have also been shown to modulate frameshifting levels.
1851:
Jacks T, Power MD, Masiarz FR, Luciw PA, Barr PJ, Varmus HE (January 1988). "Characterization of ribosomal frameshifting in HIV-1 gag-pol expression".
2567:"Structural probing and mutagenic analysis of the stem-loop required for Escherichia coli dnaX ribosomal frameshifting: programmed efficiency of 50%"
279:
In eukaryotes it appears to play a role in regulating gene expression levels by generating premature stops and producing nonfunctional transcripts.
224:
Because of this +1 frameshifting, the DNA sequence is read differently. The different codon reading frame therefore yields different amino acids.
387:
This is a graphical representation of the HIV1 frameshift signal. A −1 frameshift in the slippery sequence region results in translation of the
240:
after the frameshift, or the creation of a completely new protein after the frameshift. In the case where a frameshift results in nonsense, the
1453:
1332:
1012:
87:
Small molecules, proteins, and nucleic acids have also been found to stimulate levels of frameshifting. In
December 2023, it was reported that
1458:
1346:
1026:
296:
1436:
synthesis pathway is based on polyamine levels stimulating an increase in +1 frameshifts, which results in production of an inhibitory
972:
1428:
Small molecules, proteins, and nucleic acids have been found to stimulate levels of frameshifting. For example, the mechanism of a
2183:"Expression levels influence ribosomal frameshifting at the tandem rare arginine codons AGG_AGG and AGA_AGA in Escherichia coli"
1062:
197:. When read from the beginning, these codons make sense to a ribosome and can be translated into amino acids (AA) under the
2717:
1463:
1360:
1040:
1515:"Ribosomal frameshifting and transcriptional slippage: From genetic steganography and cryptography to adventitious use"
2013:"Reprogramming the genetic code: The emerging role of ribosomal frameshifting in regulating cellular gene expression"
2426:"Apical loop-internal loop RNA pseudoknots: a new type of stimulator of −1 translational frameshifting in bacteria"
333:
161:
and H of the first word (effectively a +1 frameshift when considering the 0 position to be the initial position of
61:
2143:
375:
Frameshifting may also be induced by other molecules which interact with the ribosome or the mRNA (trans-acting).
1473:
1378:
1054:
1006:
221:|Start|ACG AAA ATC TGT TCG CTT CA... -|Start|123 123 123 123 123 123 12... | AA | T K I C S L ...
2648:
1020:
245:
208:
AC GAA AAT CTG TTC GCT TCA ... |Start|123 123 123 123 123 123 123 ... | AA | N E N L F A S ...
198:
2668:
2401:
147:
In this example, the following sentence of three-letter words makes sense when read from the beginning:
1034:
183:
17:
2617:
2712:
2621:
2340:"Correlation between mechanical strength of messenger RNA pseudoknots and ribosomal frameshifting"
1048:
2281:
Arthur L, Pavlovic-Djuranovic S, Smith-Koutmou K, Green R, Szczesny P, Djuranovic S (July 2015).
1956:"An overlapping protein-coding region in influenza A virus segment 3 modulates the host response"
241:
2683:
415:
on mRNA, which is known to induce ribosome slippage even in the absence of any other elements.
2692:— Database of recoded genes, including those that require programmed Translational frameshift.
45:
2514:
Baranov PV, Henderson CM, Anderson CB, Gesteland RF, Atkins JF, Howard MT (February 2005).
2351:
2294:
1967:
1860:
1584:
244:(NMD) pathway may destroy the mRNA transcript, so frameshifting would serve as a method of
132:
349:
8:
1468:
255:
If a novel or off-target protein is produced, it can trigger other unknown consequences.
2355:
2298:
1971:
1864:
1724:
1689:
1588:
950:
2591:
2566:
2542:
2515:
2491:
2466:
2374:
2339:
2315:
2282:
2258:
2231:
2207:
2182:
2037:
2012:
1988:
1955:
1931:
1906:
1884:
1825:
1800:
1773:
1748:
1661:
1634:
1605:
1572:
1539:
1514:
1444:(miRNA) molecules may hybridize to an RNA secondary structure and affect its strength.
268:
2159:
2073:
2722:
2596:
2547:
2496:
2447:
2379:
2320:
2263:
2212:
2198:
2163:
2117:
2077:
2042:
1993:
1936:
1922:
1876:
1830:
1778:
1729:
1711:
1666:
1610:
1544:
1493:
1483:
1429:
1288:
1262:
1236:
1210:
1184:
1158:
1155:
1129:
1126:
1100:
1097:
1094:
404:
329:
233:
187:
107:
Proteins are translated by reading tri-nucleotides on the mRNA strand, also known as
948:
These symbols are also valid for RNA, except with U (uracil) replacing T (thymine).
2586:
2578:
2537:
2527:
2486:
2478:
2437:
2403:
2402:
Nomenclature
Committee of the International Union of Biochemistry (NC-IUB) (1984).
2369:
2359:
2310:
2302:
2253:
2243:
2202:
2194:
2155:
2107:
2069:
2032:
2024:
1983:
1975:
1926:
1918:
1888:
1868:
1820:
1812:
1768:
1760:
1719:
1701:
1656:
1646:
1600:
1592:
1534:
1526:
338:
73:
2232:"Programmed −1 frameshifting by kinetic partitioning during impeded translocation"
1571:
Napthine S, Ling R, Finch LK, Jones JD, Bell S, Brierley I, Firth AE (June 2017).
992:
2672:
1488:
1318:
998:
412:
249:
2532:
2112:
2099:
1816:
2344:
Proceedings of the
National Academy of Sciences of the United States of America
2248:
1706:
1573:"Protein-directed ribosomal frameshifting temporally regulates gene expression"
2482:
1907:"Signals for ribosomal frameshifting in the Rous sarcoma virus gag-pol region"
2701:
2627:
1715:
1651:
136:
127:
as the start (initiation) codon AUG. Each codon is translated into a single
112:
53:
2364:
1979:
2707:
2582:
2551:
2500:
2451:
2442:
2425:
2383:
2324:
2306:
2280:
2267:
2216:
2167:
2046:
2028:
1997:
1834:
1782:
1733:
1670:
1614:
1548:
1337:
171:|Start|HEC ATA NDT HEM ANA REF AT... -|Start|123 123 123 123 123 123 12...
69:
2600:
2121:
2081:
2060:
Crick FH (August 1966). "Codon—anticodon pairing: the wobble hypothesis".
1940:
1880:
1284:
1258:
1232:
1206:
1180:
1151:
1122:
1090:
27:
Phenomenon that occurs during translation of a messenger RNA into proteins
2404:"Nomenclature for Incompletely Specified Bases in Nucleic Acid Sequences"
2230:
Caliskan N, Katunin VI, Belardinelli R, Peske F, Rodnina MV (June 2014).
1764:
1530:
1351:
408:
232:
In the case of a translating ribosome, a frameshift can either result in
1596:
1513:
Atkins JF, Loughran G, Bhatt PR, Firth AE, Baranov PV (September 2016).
307:
99:(Pfizer–BioNTech) anti-COVID-19 vaccine caused ribosomal frameshifting.
2676:
2660:
2639:
747:
428:
237:
128:
124:
57:
1690:"N1-methylpseudouridylation of mRNA causes +1 ribosomal frameshifting"
407:
can potentially make the reading ribosome "slip" and skip a number of
157:
However, if the reading frame is shifted by one letter to between the
154:
HE CAT AND THE MAN ARE FAT ... |Start|123 123 123 123 123 123 123 ...
1872:
1433:
1397:
1365:
424:
272:
120:
116:
2516:"Programmed ribosomal frameshifting in decoding the SARS-CoV genome"
383:
2679:
1441:
1401:
1369:
1323:
498:
96:
77:
31:
1635:"On programmed ribosomal frameshifting: the alternative proteomes"
2656:
2631:
2513:
1413:
1383:
552:
525:
468:
363:
194:
190:
65:
49:
2424:
Mazauric MH, Licznar P, Prère MF, Canal I, Fayet O (July 2008).
2338:
Hansen TM, Reihani SN, Oddershede LB, Sørensen MA (April 2007).
2229:
2665:
2643:
2337:
1437:
718:
690:
579:
320:
316:
312:
108:
2465:
Ivanov IP, Anderson CB, Gesteland RF, Atkins JF (June 2004).
2464:
2181:
Gurvich OL, Baranov PV, Gesteland RF, Atkins JF (June 2005).
2180:
2144:"An "integrated model" of programmed ribosomal frameshifting"
661:
440:
264:
1905:
Jacks T, Madhani HD, Masiarz FR, Varmus HE (November 1988).
2283:"Translational control by lysine-encoding A-rich sequences"
1280:
1254:
1228:
1202:
1176:
1147:
1118:
1086:
359:
182:
In this example, the following sequence is a region of the
92:
81:
2652:
2635:
2423:
1904:
1479:
Insertion sequence IS1222 ribosomal frameshifting element
1393:
Insertion sequence IS1222 ribosomal frameshifting element
1069:
Insertion sequence IS1222 ribosomal frameshifting element
980:
2689:
1686:
1512:
302:
2141:
1850:
1570:
174:
then the sentence reads differently, making no sense.
60:
sequence of the mRNA and is sometimes affected by the
2564:
2142:
Harger JW, Meskauskas A, Dinman JD (September 2002).
1953:
923:
258:
1801:"Translation Elongation and Recoding in Eukaryotes"
439:
International Union of Pure and
Applied Chemistry (
48:that results in the production of multiple, unique
2565:Larsen B, Gesteland RF, Atkins JF (August 1997).
2699:
1798:
44:, is a biological phenomenon that occurs during
1454:Antizyme RNA frameshifting stimulation element
1333:Antizyme RNA frameshifting stimulation element
1013:Antizyme RNA frameshifting stimulation element
293:programmed −1 ribosomal frameshifting (−1 PRF)
80:insertion elements, and also in some cellular
1459:Coronavirus frameshifting stimulation element
1347:Coronavirus frameshifting stimulation element
1027:Coronavirus frameshifting stimulation element
966:
378:
344:
2417:
2010:
1799:Dever TE, Dinman JD, Green R (August 2018).
1746:
1805:Cold Spring Harbor Perspectives in Biology
973:
959:
418:
2620:at the U.S. National Library of Medicine
2590:
2541:
2531:
2490:
2441:
2373:
2363:
2314:
2257:
2247:
2206:
2111:
2036:
1987:
1930:
1824:
1772:
1723:
1705:
1660:
1650:
1604:
1538:
369:
287:The most common type of frameshifting is
282:
2397:
2395:
2393:
2097:
1682:
1680:
1632:
1423:
382:
348:
306:
358:because there are not equal amounts of
62:secondary, 3-dimensional mRNA structure
56:. The process can be programmed by the
14:
2700:
1279:: Secondary structure taken from the
1253:: Secondary structure taken from the
1227:: Secondary structure taken from the
1201:: Secondary structure taken from the
1175:: Secondary structure taken from the
1146:: Secondary structure taken from the
1117:: Secondary structure taken from the
1085:: Secondary structure taken from the
2659:sequence database, allowing gaps and
2390:
2137:
2135:
2133:
2131:
2093:
2091:
2059:
2011:Advani VM, Dinman JD (January 2016).
1677:
1566:
1564:
1562:
1560:
1558:
981:Gallery of secondary structure images
954:
303:Programmed −1 ribosomal frameshifting
267:(human immunodeficiency virus), RSV (
2100:"Ribosomal frameshifting viral RNAs"
1900:
1898:
1846:
1844:
1794:
1792:
1628:
1626:
1624:
1464:DnaX ribosomal frameshifting element
1361:DnaX ribosomal frameshifting element
1041:DnaX ribosomal frameshifting element
399:
2430:The Journal of Biological Chemistry
142:
102:
24:
2128:
2088:
2004:
1555:
259:Function in viruses and eukaryotes
64:. It has been described mainly in
25:
2734:
2611:
1895:
1841:
1789:
1621:
2199:10.1128/JB.187.12.4023-4032.2005
1269:
1243:
1217:
1191:
1165:
1136:
1107:
1075:
1061:
1047:
1033:
1019:
1005:
991:
362:of that particular codon in the
131:. The code itself is considered
2558:
2507:
2458:
2331:
2274:
2223:
2174:
2104:The Journal of General Virology
2053:
1474:HIV ribosomal frameshift signal
1379:HIV ribosomal frameshift signal
1055:HIV ribosomal frameshift signal
884:
740:
711:
683:
654:
628:
599:
123:) starting with the amino acid
2148:Trends in Biochemical Sciences
1947:
1740:
1506:
252:level of the associated gene.
177:
13:
1:
2160:10.1016/S0968-0004(02)02149-7
2074:10.1016/S0022-2836(66)80022-0
1747:Ivanov IP, Atkins JF (2007).
1499:
199:vertebrate mitochondrial code
2571:Journal of Molecular Biology
2471:Journal of Molecular Biology
2106:. 76 (Pt 8) (8): 1885–1892.
2062:Journal of Molecular Biology
1923:10.1016/0092-8674(88)90031-1
242:nonsense-mediated mRNA decay
7:
2718:Cis-regulatory RNA elements
2533:10.1016/j.virol.2004.11.038
2113:10.1099/0022-1317-76-8-1885
1817:10.1101/cshperspect.a032649
1447:
38:translational frameshifting
10:
2739:
2249:10.1016/j.cell.2014.04.041
2098:Brierley I (August 1995).
1707:10.1038/s41586-023-06800-3
1287:. Derived from Pseudobase
1261:. Derived from Pseudobase
1235:. Derived from Pseudobase
1209:. Derived from Pseudobase
1183:. Derived from Pseudobase
1154:. Derived from Pseudobase
1125:. Derived from Pseudobase
1093:. Derived from Pseudobase
922:
890:
862:
834:
806:
775:
746:
717:
689:
660:
634:
605:
578:
551:
524:
497:
467:
379:Frameshift signal elements
345:+1 ribosomal frameshifting
332:, a spacer region, and an
184:human mitochondrial genome
2675:— tool that compares two
2483:10.1016/j.jmb.2004.03.082
986:
793:
621:
484:
454:
227:
2622:Medical Subject Headings
2618:Frameshifting,+Ribosomal
1652:10.3389/fgene.2012.00242
334:RNA secondary structure.
2365:10.1073/pnas.0608668104
2187:Journal of Bacteriology
1980:10.1126/science.1222213
419:RNA secondary structure
115:to the other (from the
2583:10.1006/jmbi.1997.1162
2443:10.1074/jbc.M802829200
2307:10.1126/sciadv.1500154
2029:10.1002/bies.201500131
1753:Nucleic Acids Research
1519:Nucleic Acids Research
895:ucleotide (not a gap)
396:
370:Controlling mechanisms
354:
325:
283:Types of frameshifting
111:, from one end of the
42:translational recoding
1639:Frontiers in Genetics
1577:Nature Communications
1424:Trans-acting elements
1410:Ribosomal frameshift
867:comes after T and U)
386:
352:
310:
2671:19 July 2011 at the
1302:Frameshift elements
2436:(29): 20421–20432.
2356:2007PNAS..104.5830H
2299:2015SciA....1E0154A
1972:2012Sci...337..199J
1865:1988Natur.331..280J
1633:Ketteler R (2012).
1597:10.1038/ncomms15582
1589:2017NatCo...815582N
1469:Frameshift mutation
1303:
91:-transcribed (IVT)
2638:sequence allowing
1765:10.1093/nar/gkm035
1531:10.1093/nar/gkw530
1301:
455:Bases represented
443:) are as follows:
405:Slippery sequences
397:
355:
326:
297:kinetically driven
269:Rous sarcoma virus
2350:(14): 5830–5835.
2193:(12): 4023–4032.
1966:(6091): 199–204.
1859:(6153): 280–283.
1700:(7993): 189–194.
1525:(15): 7007–7078.
1494:Slippery sequence
1430:negative feedback
1421:
1420:
1298:
1297:
1283:database. Family
1257:database. Family
1231:database. Family
1205:database. Family
1179:database. Family
1150:database. Family
1121:database. Family
1089:database. Family
946:
945:
400:Slippery sequence
330:slippery sequence
234:nonsense mutation
188:overlapping genes
16:(Redirected from
2730:
2605:
2604:
2594:
2562:
2556:
2555:
2545:
2535:
2511:
2505:
2504:
2494:
2462:
2456:
2455:
2445:
2421:
2415:
2414:
2412:
2410:
2399:
2388:
2387:
2377:
2367:
2335:
2329:
2328:
2318:
2287:Science Advances
2278:
2272:
2271:
2261:
2251:
2242:(7): 1619–1631.
2227:
2221:
2220:
2210:
2178:
2172:
2171:
2139:
2126:
2125:
2115:
2095:
2086:
2085:
2057:
2051:
2050:
2040:
2008:
2002:
2001:
1991:
1951:
1945:
1944:
1934:
1902:
1893:
1892:
1873:10.1038/331280a0
1848:
1839:
1838:
1828:
1796:
1787:
1786:
1776:
1759:(6): 1842–1858.
1744:
1738:
1737:
1727:
1709:
1684:
1675:
1674:
1664:
1654:
1630:
1619:
1618:
1608:
1568:
1553:
1552:
1542:
1510:
1304:
1300:
1273:
1247:
1221:
1195:
1169:
1140:
1111:
1079:
1065:
1051:
1037:
1023:
1009:
995:
975:
968:
961:
952:
951:
446:
445:
289:−1 frameshifting
143:Sentence example
103:Process overview
74:retrotransposons
36:, also known as
21:
2738:
2737:
2733:
2732:
2731:
2729:
2728:
2727:
2713:Gene expression
2698:
2697:
2673:Wayback Machine
2614:
2609:
2608:
2563:
2559:
2512:
2508:
2463:
2459:
2422:
2418:
2408:
2406:
2400:
2391:
2336:
2332:
2293:(6): e1500154.
2279:
2275:
2228:
2224:
2179:
2175:
2140:
2129:
2096:
2089:
2058:
2054:
2009:
2005:
1952:
1948:
1903:
1896:
1849:
1842:
1797:
1790:
1745:
1741:
1685:
1678:
1631:
1622:
1569:
1556:
1511:
1507:
1502:
1489:Ribosomal pause
1484:Recode database
1450:
1426:
1319:ALIL pseudoknot
1299:
1294:
1291:
1274:
1265:
1248:
1239:
1222:
1213:
1196:
1187:
1170:
1161:
1141:
1132:
1112:
1103:
1080:
1071:
1066:
1057:
1052:
1043:
1038:
1029:
1024:
1015:
1010:
1001:
999:ALIL pseudoknot
996:
982:
979:
839:comes after G)
811:comes after C)
780:comes after A)
421:
402:
391:instead of the
381:
372:
347:
339:wobble position
305:
285:
261:
230:
222:
209:
180:
172:
155:
145:
105:
95:in response to
28:
23:
22:
15:
12:
11:
5:
2736:
2726:
2725:
2720:
2715:
2710:
2694:
2693:
2687:
2663:
2655:sequence to a
2646:
2625:
2613:
2612:External links
2610:
2607:
2606:
2557:
2526:(2): 498–510.
2506:
2477:(3): 495–504.
2457:
2416:
2389:
2330:
2273:
2222:
2173:
2154:(9): 448–454.
2127:
2087:
2068:(2): 548–555.
2052:
2003:
1946:
1917:(3): 447–458.
1894:
1840:
1811:(8): a032649.
1788:
1739:
1676:
1620:
1554:
1504:
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1501:
1498:
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935:
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931:
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854:
851:
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843:
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833:
827:
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823:
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716:
710:
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696:
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682:
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678:
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659:
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652:
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647:
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623:
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591:
589:
587:
585:
577:
571:
570:
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558:
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540:
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531:
523:
517:
516:
513:
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509:
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496:
490:
489:
486:
483:
481:
479:
477:
474:
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459:
456:
453:
450:
420:
417:
401:
398:
380:
377:
371:
368:
346:
343:
304:
301:
284:
281:
260:
257:
236:, a premature
229:
226:
217:
203:
179:
176:
167:
149:
144:
141:
104:
101:
52:from a single
26:
9:
6:
4:
3:
2:
2735:
2724:
2721:
2719:
2716:
2714:
2711:
2709:
2706:
2705:
2703:
2696:
2691:
2688:
2685:
2681:
2678:
2674:
2670:
2667:
2664:
2662:
2658:
2654:
2650:
2647:
2645:
2641:
2637:
2633:
2629:
2626:
2623:
2619:
2616:
2615:
2602:
2598:
2593:
2588:
2584:
2580:
2576:
2572:
2568:
2561:
2553:
2549:
2544:
2539:
2534:
2529:
2525:
2521:
2517:
2510:
2502:
2498:
2493:
2488:
2484:
2480:
2476:
2472:
2468:
2461:
2453:
2449:
2444:
2439:
2435:
2431:
2427:
2420:
2405:
2398:
2396:
2394:
2385:
2381:
2376:
2371:
2366:
2361:
2357:
2353:
2349:
2345:
2341:
2334:
2326:
2322:
2317:
2312:
2308:
2304:
2300:
2296:
2292:
2288:
2284:
2277:
2269:
2265:
2260:
2255:
2250:
2245:
2241:
2237:
2233:
2226:
2218:
2214:
2209:
2204:
2200:
2196:
2192:
2188:
2184:
2177:
2169:
2165:
2161:
2157:
2153:
2149:
2145:
2138:
2136:
2134:
2132:
2123:
2119:
2114:
2109:
2105:
2101:
2094:
2092:
2083:
2079:
2075:
2071:
2067:
2063:
2056:
2048:
2044:
2039:
2034:
2030:
2026:
2022:
2018:
2014:
2007:
1999:
1995:
1990:
1985:
1981:
1977:
1973:
1969:
1965:
1961:
1957:
1950:
1942:
1938:
1933:
1928:
1924:
1920:
1916:
1912:
1908:
1901:
1899:
1890:
1886:
1882:
1878:
1874:
1870:
1866:
1862:
1858:
1854:
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1818:
1814:
1810:
1806:
1802:
1795:
1793:
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1762:
1758:
1754:
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1743:
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1717:
1713:
1708:
1703:
1699:
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1691:
1683:
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1672:
1668:
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1653:
1648:
1644:
1640:
1636:
1629:
1627:
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1598:
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1574:
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1565:
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1546:
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1536:
1532:
1528:
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1509:
1505:
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1409:
1408:
1405:
1403:
1399:
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1390:
1387:
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1382:
1380:
1377:
1376:
1373:
1371:
1367:
1364:
1362:
1359:
1358:
1355:
1353:
1350:
1348:
1345:
1344:
1341:
1339:
1338:Invertebrates
1336:
1334:
1331:
1330:
1327:
1325:
1322:
1320:
1317:
1316:
1312:
1310:Distribution
1309:
1306:
1305:
1290:
1286:
1282:
1278:
1272:
1267:
1264:
1260:
1256:
1252:
1246:
1241:
1238:
1234:
1230:
1226:
1220:
1215:
1212:
1208:
1204:
1200:
1194:
1189:
1186:
1182:
1178:
1174:
1168:
1163:
1160:
1157:
1153:
1149:
1145:
1139:
1134:
1131:
1128:
1124:
1120:
1116:
1110:
1105:
1102:
1099:
1096:
1092:
1088:
1084:
1078:
1073:
1070:
1064:
1059:
1056:
1050:
1045:
1042:
1036:
1031:
1028:
1022:
1017:
1014:
1008:
1003:
1000:
994:
989:
988:
985:
976:
971:
969:
964:
962:
957:
956:
953:
949:
941:
938:
936:
934:
932:
930:
928:
926:
920:
917:
916:
912:
909:
906:
903:
900:
897:
894:
888:
885:
880:
878:
875:
872:
869:
866:
860:
857:
856:
852:
849:
847:
844:
841:
838:
832:
829:
828:
824:
821:
818:
816:
813:
810:
804:
801:
800:
796:
790:
787:
784:
782:
779:
773:
770:
769:
765:
762:
760:
757:
755:
753:
751:
744:
741:
736:
734:
731:
729:
726:
724:
722:
715:
712:
707:
704:
701:
699:
697:
695:
693:
687:
684:
679:
677:
675:
672:
669:
667:
665:
658:
655:
650:
648:
645:
642:
640:
637:
632:
629:
624:
618:
616:
614:
611:
608:
603:
600:
595:
592:
590:
588:
586:
584:
582:
576:
573:
572:
568:
565:
563:
561:
559:
557:
555:
549:
546:
545:
541:
539:
536:
534:
532:
530:
528:
522:
519:
518:
514:
512:
510:
507:
505:
503:
501:
495:
492:
491:
487:
482:
480:
478:
475:
473:
471:
465:
462:
461:
457:
451:
448:
447:
444:
442:
436:
432:
430:
426:
416:
414:
410:
406:
394:
390:
385:
376:
367:
365:
361:
351:
342:
340:
335:
331:
322:
318:
314:
309:
300:
298:
294:
290:
280:
277:
274:
270:
266:
256:
253:
251:
247:
243:
239:
235:
225:
220:
216:
214:
207:
202:
200:
196:
192:
189:
186:with the two
185:
175:
170:
166:
164:
160:
153:
148:
140:
138:
137:reading frame
134:
130:
126:
122:
118:
114:
110:
100:
98:
94:
90:
85:
83:
79:
75:
71:
67:
63:
59:
55:
51:
47:
43:
39:
35:
34:frameshifting
33:
19:
2695:
2651:— compare a
2577:(1): 47–60.
2574:
2570:
2560:
2523:
2519:
2509:
2474:
2470:
2460:
2433:
2429:
2419:
2407:. Retrieved
2347:
2343:
2333:
2290:
2286:
2276:
2239:
2235:
2225:
2190:
2186:
2176:
2151:
2147:
2103:
2065:
2061:
2055:
2023:(1): 21–26.
2020:
2016:
2006:
1963:
1959:
1949:
1914:
1910:
1856:
1852:
1808:
1804:
1756:
1752:
1742:
1697:
1693:
1642:
1638:
1580:
1576:
1522:
1518:
1508:
1432:loop in the
1427:
1276:
1250:
1224:
1198:
1172:
1143:
1114:
1082:
947:
924:
918:
892:
886:
864:
858:
836:
830:
808:
802:
777:
771:
749:
742:
720:
713:
691:
685:
663:
656:
635:
630:
606:
601:
580:
574:
553:
547:
526:
520:
499:
493:
469:
463:
452:Description
437:
433:
422:
403:
392:
388:
373:
356:
327:
292:
288:
286:
278:
262:
254:
231:
223:
218:
212:
210:
205:
181:
173:
168:
162:
158:
156:
151:
146:
106:
88:
86:
70:retroviruses
68:(especially
41:
37:
30:
29:
2684:translation
2661:frameshifts
2640:frameshifts
2630:— aligns a
1352:Coronavirus
458:Complement
409:nucleotides
178:DNA example
46:translation
2702:Categories
2686:principle)
2677:frameshift
2634:against a
2409:4 February
1500:References
429:pseudoknot
271:) and the
250:expression
246:regulating
238:stop codon
133:degenerate
129:amino acid
125:methionine
58:nucleotide
18:Frameshift
2017:BioEssays
1716:1476-4687
1583:: 15582.
1434:polyamine
1398:Eukaryota
1366:Eukaryota
425:stem-loop
273:influenza
78:bacterial
32:Ribosomal
2723:Genetics
2680:proteins
2669:Archived
2552:15680415
2520:Virology
2501:15147837
2452:18474594
2384:17389398
2325:26322332
2268:24949973
2217:15937165
2168:12217519
2047:26661048
1998:22745253
1835:29610120
1783:17332016
1734:38057663
1725:10764286
1671:23181069
1615:28593994
1549:27436286
1448:See also
1442:MicroRNA
1402:bacteria
1370:bacteria
1324:Bacteria
1289:PKB00080
1277:RF_site9
1263:PKB00107
1251:RF_site8
1237:PKB00128
1225:RF_site6
1211:PKB00258
1199:RF_site5
1185:PKB00257
1173:RF_site4
1159:PKB00043
1156:PKB00042
1144:RF_site3
1130:PKB00233
1127:PKB00218
1115:RF_site2
1101:PKB00240
1098:PKB00044
1095:PKB00046
1083:RF_site1
752:rimidine
97:BNT162b2
89:in vitro
50:proteins
2690:Recode2
2657:protein
2644:introns
2632:protein
2601:9300054
2592:7126992
2543:7111862
2492:7125782
2375:1838403
2352:Bibcode
2316:4552401
2295:Bibcode
2259:7112342
2208:1151738
2122:7636469
2082:5969078
2038:4749135
1989:3552242
1968:Bibcode
1960:Science
1941:2846182
1932:7133365
1889:4242582
1881:2447506
1861:Bibcode
1826:6071482
1774:1874602
1662:3500957
1645:: 242.
1606:5472766
1585:Bibcode
1540:5009743
1414:Viruses
1384:Viruses
1285:RF01098
1259:RF01097
1233:RF01094
1207:RF01093
1181:RF01090
1152:RF01079
1123:RF01076
1091:RF01074
863:not T (
835:not G (
807:not C (
776:not A (
502:ytosine
449:Symbol
364:cytosol
204:|Start|
195:MT-ATP6
191:MT-ATP8
150:|Start|
119:to the
66:viruses
2682:(back-
2624:(MeSH)
2599:
2589:
2550:
2540:
2499:
2489:
2450:
2382:
2372:
2323:
2313:
2266:
2256:
2215:
2205:
2166:
2120:
2080:
2045:
2035:
1996:
1986:
1939:
1929:
1887:
1879:
1853:Nature
1833:
1823:
1781:
1771:
1732:
1722:
1714:
1694:Nature
1669:
1659:
1613:
1603:
1547:
1537:
1438:enzyme
638:trong
556:hymine
529:uanine
472:denine
319:, and
228:Effect
121:3' end
109:codons
2649:FastY
2628:Wise2
1885:S2CID
1313:Ref.
1307:Type
583:racil
441:IUPAC
413:polyA
265:HIV-1
93:mRNAs
82:genes
2666:Path
2642:and
2597:PMID
2548:PMID
2497:PMID
2448:PMID
2411:2008
2380:PMID
2321:PMID
2264:PMID
2236:Cell
2213:PMID
2164:PMID
2118:PMID
2078:PMID
2043:PMID
1994:PMID
1937:PMID
1911:Cell
1877:PMID
1831:PMID
1779:PMID
1730:PMID
1712:ISSN
1667:PMID
1611:PMID
1545:PMID
1281:Rfam
1255:Rfam
1229:Rfam
1203:Rfam
1177:Rfam
1148:Rfam
1119:Rfam
1087:Rfam
891:any
609:eak
360:tRNA
248:the
193:and
113:mRNA
76:and
54:mRNA
2708:RNA
2653:DNA
2636:DNA
2587:PMC
2579:doi
2575:271
2538:PMC
2528:doi
2524:332
2487:PMC
2479:doi
2475:339
2438:doi
2434:283
2370:PMC
2360:doi
2348:104
2311:PMC
2303:doi
2254:PMC
2244:doi
2240:157
2203:PMC
2195:doi
2191:187
2156:doi
2108:doi
2070:doi
2033:PMC
2025:doi
1984:PMC
1976:doi
1964:337
1927:PMC
1919:doi
1869:doi
1857:331
1821:PMC
1813:doi
1769:PMC
1761:doi
1720:PMC
1702:doi
1698:625
1657:PMC
1647:doi
1601:PMC
1593:doi
1535:PMC
1527:doi
927:ero
723:ine
694:eto
666:ino
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