211:. For example, in plasmid ColE1, the asRNA termed RNA I plays an important role in determining the plasmid copy number by controlling replication. The replication of ColE1 relies on the transcription of a primer RNA named RNA II. Once RNA II is transcribed, it hybridizes to its DNA template and later cleaved by RNase H. In the presence of the asRNA RNA I, RNA I and RNA II forms a duplex which introduces a conformational change of RNA II. Consequently, RNA II cannot hybridize with its DNA template which results in a low copy number of ColE1. In bacteriophage P22, the asRNA sar helps regulate between lytic and lysogenic cycle by control the expression of Ant. Besides being expressed in prokaryotes, asRNAs were also discovered in plants. The most well described example of asRNA regulation in plants is on
503:
locus-specific nature of the endogenous asRNAs, only 10–50% synthesized oligonucleotides showed expected targeting effect. One possible reason for this problem is the high requirement on the structure of the asRNAs to be recognized by the target sequence and RNase H. A single mismatch can result in distortion in the secondary structure and lead to off target effects. Lastly, artificial asRNAs have been shown to have limited intracellular uptake. Although neurons and glia have been shown to have the ability to freely uptake naked antisense oligonucleotides, a traceable carriers such as virus and lipid vesicles would still be ideal to control and monitor the intracellular concentration and metabolism.
31:
328:
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gene regulation. For example, in bacterial or eukaryotic cells where complex RNA polymerases are present, bidirectional transcription at the same locus can lead to polymerase collision and results in the termination of transcription. Even when polymerase collision is unlikely during weak transcription, polymerase pausing can also occur which blocks elongation and leads to gene repression. One of the examples is repression of
489:, it is always easier to have drugs functioning as downregulators or inhibitors. However, there is a need in developing drugs that can activate or upregulate gene expression such as tumor suppressor genes, neuroprotective growth factors and genes that are found silenced in certain Mendelian disorders. Currently, the approach to restore deficient gene expression or protein function include enzyme replacement therapies,
1875:
335::' a) AsRNAs can induce DNA methylation by recruiting a DNA methyltransferase (DNMT). b) AsRNAs can induce histone methylation by recruiting of a histone methyltransferase (HMT). 'Co-transcriptional regulation:' c) AsRNAs can cause RNA polymerase (Pol) collision and stop the transcription. d) AsRNAs can preference translation of a specific splice variant (mRNA V1) by blocking the other splice variant (mRNA V2). '
464:. With the asRNA of ZEB2 being expressed, it can mask the splicing site and maintain the IRES in the mRNA which results in an efficient synthesis of E-cadherin. Lastly, depending on the level of asRNA expression, different isoforms of the sense transcript can be produced. Therefore, asRNA dependent regulation is not limited to on/off mechanism; rather, it presents a fine tone control system.
473:
and its asRNA need to be present simultaneously in the same cell. As described in the cis-acting asRNAs, the mRNA-asRNA pairing can result in blockage of ribosome entry and RNase H dependent degradation. Overall, mRNA-targeting asRNAs can either activate or inhibit translation of the sense mRNAs with inhibitory effect being the most abundant.
482:
degree of complementarity with the targeting genes. Thirdly, the expression level of asRNAs is very small compared to that of the targeting mRNAs; therefore, only small amount of asRNAs is required to produce an effect. In terms of drug targets, this represents a huge advantage because only a low dosage is required for effectiveness.
494:
large financial burden for the patient. Because of the locus specific nature of asRNAs and evidences of changes in asRNA expression in many diseases, there have been attempts to design single stranded oligonucleotides, referred as antagoNATs, to inhibit asRNAs and ultimately to increase specific gene expression.
117:, some of the ompC promoter clones observed were capable of repressing the expression of other membrane porin such as ompF. The region responsible for this repression function was found to be a 300 base-pair locus upstream of the ompC promoter. This 300 base-pair region is 70% homologous in sequence with the
502:
linkage to the backbones. However, the phosphrothioate modification can be proinflammatory. Adverse effects including fever, chills or nausea have been observed after local injection of phosphrothioate modified oligonucleotides. Secondly, off target toxicity also represents a big problem. Despite the
497:
Despite the promises of asRNAs as drug targets or drug candidates, there are some challenges remained to be addressed. First of all, asRNAs and antagoNATs can be easily degraded by RNase or other degrading enzymes. To prevent degradation of the therapeutic oliogoneucleotides, chemical modification is
252:, there are short asRNAs with length of less than 200 base pairs. Because the regulatory mechanism of asRNAs are found to be species specific, asRNAs can also be classified by species. One of the common ways of classifying asRNAs is by where the asRNAs are transcribe relatively to their target genes:
121:
of the ompF mRNA and thus the transcript of this 300 base pair locus was complementary to the ompF mRNA. Later on, this transcript, denoted micF, was found to be an asRNA of ompF and capable of downregulating the expression of ompF under stress by forming a duplex with the ompF mRNA. This induces the
493:
therapies and delivery of functional cDNA. However, each bears some drawbacks. For example, the synthesized protein used in the enzyme replacement therapies often cannot mimic the whole function of the endogenous protein. In addition, enzyme replacement therapies are life-long commitment and carry a
472:
The direct post transcriptional modulation by asRNAs refers to mRNAs being targeted by asRNAs directly; thus, the translation is affected. Some characteristics of this type of asRNAs are described in the cis- and trans- acting asRNAs. This mechanism is relatively fast because both the targeting mRNA
142:
can be used. In this method, one or both strands of encoding genes can be used as probes. In addition to computational searches and microarrays, some asRNAs were discovered by sequencing cDNA clones as well as mapping promoter elements. Although many findings from the approaches mentioned above gave
439:
Epigenetic regulations such as DNA methylation and histone methylation can repress gene expression by inhibiting initiation of transcription. Sometimes, however, gene repression can be achieved by prematurely terminating or slowing down transcription process. AsRNAs can be involved in this level of
398:
are context dependent. In general, histone methylation leads to gene repression but gene activation can also be achieved. Evidence has shown histone methylation can be induced by asRNAs. For instance, ANRIL, in addition to the ability to induce DNA methylation, can also repress the neighboring gene
219:
encodes for a transcription factor that prevent expression of a range of genes that induce floral transition. In cold environment, the asRNA of FLC gene, denoted COOLAIR, is expressed and inhibits the expression of FLC via chromatin modification which consequently allows for flowering. Another well
367:
gene (HBA1) is downregulated by an abnormal transcript of putative RNA-binding protein Luc7-like (LUC71) that serves as an asRNA to HBA1 and induces methylation of HBA1's promoter. Another example is silencing of a tumor suppressor gene p15INK4b, also called CDKN2B, in acute lymphoblastic leukemia
277:
asRNAs are transcribed from the opposite strand of the target gene at the target gene locus. They often show high degree or complete complementarity with the target gene. If the cis-acting asRNA regulates gene expression by targeting mRNA, it can only target individual mRNA. Upon interactions with
163:
in patients with AIDS. It works by targeting the transcribed mRNA of the virus and consequently inhibiting replication of cytomegalovirus. Despite fomivirsen being discontinued in 2004 due to the loss of the market, it served as a successful and inspiring example of using asRNAs as drug targets or
481:
As a regulatory element, asRNAs bear many advantages to be considered as a drug target. First of all, asRNAs regulate gene expression at multiple levels including transcription, post-transcription and epigenetic modification. Secondly, the cis-acting asRNAs are sequence specific and exhibits high
1094:
Kaur, Devinder; Agrahari, Mridula; Singh, Shashi
Shekhar; Mandal, Prabhat Kumar; Bhattacharya, Alok; Bhattacharya, Sudha (2021). "Transcriptomic analysis of Entamoeba histolytica reveals domain-specific sense strand expression of LINE-encoded ORFs with massive antisense expression of RT domain".
314:
asRNAs are transcribed from loci that are distal from the targeting genes. In contrast to cis-acting asRNAs, they display low degree of complementarity with the target gene but can be longer than cis-acting asRNAs. They can also target multiple loci. Because of these properties of trans-acting
129:
analysis. Conventionally, the first step involves computational predictions based on some known characteristics of asRNAs. During computational searches, the encoding regions are excluded. The regions that are predicted to have conserved RNA structures and act as orphan promoters and
154:
and
Stephenson found an antisense oligonucleotide to the viral RNA of Rous scarcoma virus that was capable of inhibiting viral replication and protein synthesis. Since then, much effort has been devoted to developing asRNAs as drug candidates. In 1998, the first asRNA drug,
179:(HoFH), which is a rare autosomal dominant genetic condition. Because of the high level of total cholesterol (650–1000 mg/dL) and LDL receptor (above 600 mg/dL) in HoFH, patients with HoFH has a high risk for coronary heart disease. Because the protein
220:
studied example is DOG1 (Delay of
Germination 1) gene. Its expression level is negatively regulated by the antisense transcript (asDOG1 or 1GOD) acting in cis. In mammalian cells, a typical example of asRNA regulation is X chromosome inactivation.
248:). Antisense RNAs can be categorized by the type of the promoters that initiate expression of asRNAs: independent promoters, shared bidirectional promoters or cryptic promoters. In terms of length, although asRNA in general is classified as
143:
rise to a lot of possible asRNAs, only few were proven to be actual asRNAs via further functional tests. To minimize the number of false positive results, new approaches from recent years have been focusing on strand-specific transcription,
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where they are transcribed. Instead of targeting individual mRNAs, these cis-acting epigenetic regulators can recruit chromatin modifying enzymes which can exert effects on both the transcription loci and the neighboring genes.
282:
to degrade the targeting mRNAs. Consequently, the function of these cis-acting asRNAs is to repress translation of the targeting mRNAs. Besides cis-acting asRNAs that target mRNAs, there are cis-acting epigenetic
418:
ANRIL induced epigenetic modification is an example of cis acting epigenetic regulation. In addition, Antisense RNA-induced chromatin modification can be both trans-acting. For example, in mammals, the asRNA
339::' e) AsRNA-mRNA duplex can either block ribosome from binding to the mRNA or recruit RNase H to degrade the mRNA. By this mechanism, asRNAs directly inhibit translation of the mRNAs.
359:
can result in long term downregulation of specific genes. Repression of functional proteins via asRNA induced DNA methylation has been found in several human disease. In a class of
138:, the asRNAs that are transcribed from the opposite strand of an encoding gene are likely to be missed using this method. To detect asRNA transcribed from the encoding region,
236:
Antisense RNAs can be classified in different ways. In terms of regulatory mechanisms, some authors group asRNAs into RNA-DNA interactions, RNA-RNA interactions either in
1043:
Fedak, Halina; Palusinska, Malgorzata; Krzyczmonik, Katarzyna; Brzezniak, Lien; Yatusevich, Ruslan; Pietras, Zbigniew; Kaczanowski, Szymon; Swiezewski, Szymon (2016).
1180:
Weiss, B. (ed.): Antisense
Oligodeoxynucleotides and Antisense RNA : Novel Pharmacological and Therapeutic Agents, CRC Press, Boca Raton, FL, 1997.
485:
Recent years the idea of targeting asRNAs to increase gene expression in a locus specific manner has been drawing much attention. Due to the nature of
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such as Hfq to exert their functions. Due to the complexity of the trans-acting asRNAs, they are currently considered to be less druggable targets.
1470:
1208:
125:
Unlike micF RNA being discovered by accident, the majority of asRNAs were discovered by genome wide searches for small regulatory RNAs and by
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1008:
Ietswaart R, Wu Z, Dean C (September 2012). "Flowering time control: another window to the connection between antisense RNA and chromatin".
172:
1556:
934:"Mipomersen (kynamro): a novel antisense oligonucleotide inhibitor for the management of homozygous familial hypercholesterolemia"
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and acute myeloid leukemia. The asRNA that is responsible for this silencing effect is antisense non-coding RNA in the INK locus (
1843:
1314:
512:
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In terms of epigenetic modification, cis-acting refers to the nature of these asRNAs that regulate epigenetic changes around the
269:
1848:
1242:
336:
327:
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1631:
176:
34:
AsRNA is transcribed from the lagging strand of a gene and is complementary to a specific mRNA or sense transcript.
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456:) which encodes E-cadherin, a transcriptional repressor. Efficient translation of ZEB2 mRNA requires the presence of an
184:
46:
448:. Another way of affecting transcription co-transcriptionally is by blocking splicing. One classic example in human is
30:
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131:
1201:
139:
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1427:
363:, a type of blood disorder that has reduced level of hemoglobin leading to insufficient oxygen in the tissues,
411:(PRC2) which leads to histone methylation (H3K27me). Another classic example is X chromosome inactivation by
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Many examples of asRNAs show the inhibitory effect on transcription initiation via epigenetic modifications.
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asRNAs, they form less stable complexes with their targeting transcripts and sometimes require aids from RNA
1904:
1899:
1878:
1571:
1456:
1252:
727:
Wahlestedt C (June 2013). "Targeting long non-coding RNA to therapeutically upregulate gene expression".
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332:
292:
62:
605:"Natural antisense RNAs as mRNA regulatory elements in bacteria: a review on function and applications"
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to HOXD which deposits H3K27 and silences HOXD. HOTAIR is highly expressed in primary breast tumors.
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Some of the earliest asRNAs were discovered while investigating functional proteins. An example was
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1509:
1324:
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364:
284:
249:
667:"Regulation of chromatin structure by long noncoding RNAs: focus on natural antisense transcripts"
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1549:
1354:
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752:
578:
85:. asRNAs may also be produced synthetically and have found wide spread use as research tools for
390:. Modification on histones can change interactions with DNA which can further induce changes in
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usually required. The most common chemical modification on the oligonucleotides is adding a
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159:, was approved by FDA. Fomivirsen, a 21 base-pair oligonucleotide, was developed to treat
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356:
108:
82:
50:
553:
Pelechano V, Steinmetz LM (December 2013). "Gene regulation by antisense transcription".
1045:"Control of seed dormancy in Arabidopsis by a cis-acting noncoding antisense transcript"
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852:
827:
77:, and can be classified into short (<200 nucleotides) and long (>200 nucleotides)
1805:
1708:
1525:
1157:
1071:
1044:
973:
Simons RW (December 1988). "Naturally occurring antisense RNA control—a brief review".
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171:, which was approved by FDA in 2013. Mipomersen was developed to manage the level of
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58:
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the targeting mRNAs, cis-acting asRNAs can either block ribosome binding or recruit
187:(VLDL) and LDL, mipomersen complements with the mRNA of apo-B-100 and target it for
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1437:
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869:
582:
311:
279:
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291:. Antisense RNA has been shown to repress the translation of LINE1-ORF2 domain of
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dependent degradation. Ultimately, mipomersen is able to reduce the level of LDL.
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134:
are preferenced during analysis. Because computational searches focuses on the
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828:"Antisense RNA gene therapy for studying and modulating biological processes"
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843:
779:"RNA therapeutics: beyond RNA interference and antisense oligonucleotides"
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1329:
603:
Saberi F, Kamali M, Najafi A, Yazdanparast A, Moghaddam MM (2016-07-28).
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885:"Bacterial antisense RNAs: how many are there, and what are they doing?"
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228:(PRC2) which results in heterochromatinization of the X chromosome.
372:), which is expressed in the same locus that encodes for p15INK4b.
295:. However it is not confirmed yet whether its cis-acting or trans.
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1293:
665:
Magistri M, Faghihi MA, St
Laurent G, Wahlestedt C (August 2012).
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69:. The asRNAs (which occur naturally) have been found in both
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The initial asRNAs discovered were in prokaryotes including
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Another example of using an asRNA as a therapeutic agent is
1137:. NIH U.S. National Library of Medicine. 14 November 2017.
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The idea of asRNAs as drug targets started in 1978 when
61:(mRNA) with which it hybridizes, and thereby blocks its
81:(ncRNAs). The primary function of asRNA is regulating
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147:binding noncoding RNAs and single cell studies.
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89:. They may also have therapeutic applications.
386:In eukaryotic cells, DNA is tightly packed by
1464:
1202:
777:Kole R, Krainer AR, Altman S (January 2012).
598:
596:
594:
592:
45:), also referred to as antisense transcript,
8:
1148:Whetstine JR (2010). "Histone Methylation".
826:Weiss B, Davidkova G, Zhou LW (March 1999).
772:
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768:
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107:. While characterizing the outer membrane
57:that is complementary to a protein coding
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183:has been found to be required to produce
93:Discovery and history in drug development
609:Cellular & Molecular Biology Letters
1152:(Second ed.). pp. 2389–2397.
524:
7:
932:Wong E, Goldberg T (February 2014).
832:Cellular and Molecular Life Sciences
460:(IRES) in intron of the mRNA at the
901:10.1146/annurev-genet-102209-163523
173:low-density lipoprotein cholesterol
1158:10.1016/b978-0-12-374145-5.00287-4
175:(LDL) in patients with homozygous
25:
1595:Micro
394:. The biological consequences of
1874:
1873:
1315:Cis-natural antisense transcript
513:Cis-natural antisense transcript
270:Cis-natural antisense transcript
1550:precursor, heterogenous nuclear
1243:Signal recognition particle RNA
468:Post-transcriptional regulation
452:E-box binding homeobox 2 gene (
427:C (HOXC) locus but it recruits
337:Post-transcriptional regulation
1680:Trans-acting small interfering
1644:Enhancer RNAs
1562:Transfer
783:Nature Reviews. Drug Discovery
729:Nature Reviews. Drug Discovery
244:and RNA-protein interactions (
122:degradation of the ompF mRNA.
1:
1567:Ribosomal
1545:Messenger
1109:10.1016/j.plasmid.2021.102560
883:Thomason MK, Storz G (2010).
435:Co-transcriptional regulation
409:polycomb repressive complex 2
226:polycomb repressive complex 2
177:familial hypercholesterolemia
987:10.1016/0378-1119(88)90125-4
458:internal ribosome entry site
185:very low-density lipoprotein
47:natural antisense transcript
140:oligonucleotide microarrays
132:Rho independent terminators
1921:
1746:Multicopy single-stranded
1590:Interferential
1402:Reverse transcribing virus
1150:Handbook of Cell Signaling
379:
267:
96:
1869:
1660:Guide
1022:10.1016/j.tig.2012.06.002
889:Annual Review of Genetics
683:10.1016/j.tig.2012.03.013
622:10.1186/s11658-016-0007-z
161:cytomegalovirus retinitis
1622:Small nuclear
555:Nature Reviews. Genetics
224:, an asRNA, can recruit
1736:Genomic
1369:Cis-regulatory elements
1340:Repeat-associated siRNA
1135:Genetics Home Reference
1062:10.1073/pnas.1608827113
213:Flowering Locus C (FLC)
195:Examples across species
53:, is a single stranded
1839:Artificial chromosomes
1627:Small nucleolar
1253:Transfer-messenger RNA
1049:Proc Natl Acad Sci USA
340:
35:
1632:Small Cajal Body RNAs
1345:Small interfering RNA
844:10.1007/s000180050296
477:Therapeutic potential
344:Epigenetic regulation
333:Epigenetic regulation
330:
293:Entamoeba histolytica
33:
1685:Subgenomic messenger
1600:Small interfering
1572:Transfer-messenger
1335:Piwi-interacting RNA
423:is transcribed from
382:Histone modification
376:Histone modification
217:Arabidopsis thaliana
1274:Small nucleolar RNA
1131:"alpha thalassemia"
1055:(48): E7846–E7855.
396:histone methylation
49:(NAT) or antisense
27:Single stranded RNA
1714:Chloroplast
1557:modified Messenger
1520:Ribonucleic acids
1355:Trans-acting siRNA
1350:Small temporal RNA
1325:Long noncoding RNA
1010:Trends in Genetics
671:Trends in Genetics
444:gene by its asRNA
341:
215:gene. FLC gene in
36:
1887:
1886:
1764:Xeno
1726:Complementary
1699:Deoxyribonucleic
1693:
1692:
1670:Small hairpin
1446:
1445:
1360:Short hairpin RNA
1269:Small nuclear RNA
1226:Protein synthesis
1167:978-0-12-374148-6
365:hemoglobin alpha1
361:alpha-thalassemia
317:chaperone protein
164:drug candidates.
136:intergenic region
99:Antisense therapy
16:(Redirected from
1912:
1877:
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1854:Yeast
1675:Small temporal
1605:Piwi-interacting
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1510:Deoxynucleotides
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500:phosphorothioate
487:drug development
407:, by recruiting
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1806:Cloning vectors
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1397:Retrotransposon
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1303:Gene regulation
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567:10.1038/nrg3594
561:(12): 880–893.
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392:gene expression
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357:DNA methylation
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352:DNA methylation
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79:non-coding RNAs
51:oligonucleotide
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1634:
1629:
1624:
1614:
1609:
1608:
1607:
1602:
1597:
1586:
1584:
1578:
1577:
1575:
1574:
1569:
1564:
1559:
1554:
1553:
1552:
1541:
1539:
1530:
1516:
1515:
1513:
1512:
1507:
1502:
1497:
1491:
1489:
1485:
1484:
1481:nucleic acids
1478:
1476:
1475:
1468:
1461:
1453:
1444:
1443:
1441:
1440:
1435:
1430:
1428:Telomerase RNA
1424:
1422:
1418:
1417:
1415:
1414:
1409:
1404:
1399:
1393:
1391:
1387:
1386:
1384:
1383:
1378:
1372:
1370:
1366:
1365:
1363:
1362:
1357:
1352:
1347:
1342:
1337:
1332:
1327:
1322:
1317:
1312:
1306:
1304:
1300:
1299:
1297:
1296:
1291:
1286:
1281:
1276:
1271:
1265:
1263:
1262:RNA processing
1259:
1258:
1256:
1255:
1250:
1245:
1240:
1235:
1229:
1227:
1223:
1222:
1216:
1214:
1213:
1206:
1199:
1191:
1183:
1182:
1173:
1166:
1140:
1122:
1086:
1035:
1016:(9): 445–453.
1000:
981:(1–2): 35–44.
965:
944:(2): 119–122.
924:
895:(1): 167–188.
875:
838:(3): 334–358.
818:
789:(2): 125–140.
762:
735:(6): 433–446.
706:
677:(8): 389–396.
646:
588:
523:
522:
520:
517:
516:
515:
508:
505:
478:
475:
469:
466:
436:
433:
380:Main article:
377:
374:
353:
350:
345:
342:
324:
321:
308:
305:
265:
262:
233:
232:Classification
230:
196:
193:
97:Main article:
94:
91:
87:gene knockdown
26:
24:
18:Rna, antisense
14:
13:
10:
9:
6:
4:
3:
2:
1917:
1906:
1903:
1901:
1900:Antisense RNA
1898:
1897:
1895:
1880:
1872:
1871:
1868:
1860:
1857:
1855:
1852:
1850:
1847:
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1837:
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1827:
1825:
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1807:
1803:
1797:
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1777:
1775:
1774:Threose
1772:
1770:
1767:
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1765:
1762:
1761:
1759:
1757:
1753:
1747:
1744:
1742:
1739:
1737:
1734:
1732:
1731:Deoxyribozyme
1729:
1727:
1724:
1720:
1719:Mitochondrial
1717:
1715:
1712:
1711:
1710:
1707:
1706:
1704:
1702:
1696:
1686:
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1560:
1558:
1555:
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1548:
1547:
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1540:
1538:
1537:Translational
1534:
1531:
1527:
1521:
1517:
1511:
1508:
1506:
1503:
1501:
1498:
1496:
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1410:
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1405:
1403:
1400:
1398:
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1392:
1388:
1382:
1381:SECIS element
1379:
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1367:
1361:
1358:
1356:
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1338:
1336:
1333:
1331:
1328:
1326:
1323:
1321:
1318:
1316:
1313:
1311:
1310:Antisense RNA
1308:
1307:
1305:
1301:
1295:
1292:
1290:
1287:
1285:
1282:
1280:
1277:
1275:
1272:
1270:
1267:
1266:
1264:
1260:
1254:
1251:
1249:
1246:
1244:
1241:
1239:
1238:Ribosomal RNA
1236:
1234:
1233:Messenger RNA
1231:
1230:
1228:
1224:
1220:
1212:
1207:
1205:
1200:
1198:
1193:
1192:
1189:
1177:
1174:
1169:
1163:
1159:
1155:
1151:
1144:
1141:
1136:
1132:
1126:
1123:
1118:
1114:
1110:
1106:
1102:
1098:
1090:
1087:
1082:
1078:
1073:
1068:
1063:
1058:
1054:
1050:
1046:
1039:
1036:
1031:
1027:
1023:
1019:
1015:
1011:
1004:
1001:
996:
992:
988:
984:
980:
976:
969:
966:
961:
957:
952:
947:
943:
939:
935:
928:
925:
920:
916:
911:
906:
902:
898:
894:
890:
886:
879:
876:
871:
867:
863:
859:
854:
849:
845:
841:
837:
833:
829:
822:
819:
814:
810:
805:
800:
796:
792:
788:
784:
780:
773:
771:
769:
767:
763:
758:
754:
750:
746:
742:
738:
734:
730:
723:
721:
719:
717:
715:
713:
711:
707:
702:
698:
693:
688:
684:
680:
676:
672:
668:
661:
659:
657:
655:
653:
651:
647:
642:
638:
633:
628:
623:
618:
614:
610:
606:
599:
597:
595:
593:
589:
584:
580:
576:
572:
568:
564:
560:
556:
549:
547:
545:
543:
541:
539:
537:
535:
533:
531:
529:
525:
518:
514:
511:
510:
506:
504:
501:
495:
492:
488:
483:
476:
474:
467:
465:
463:
459:
455:
451:
447:
443:
434:
432:
430:
426:
422:
416:
414:
410:
406:
402:
397:
393:
389:
383:
375:
373:
371:
366:
362:
358:
351:
349:
343:
338:
334:
329:
322:
320:
318:
313:
306:
304:
301:
296:
294:
290:
286:
281:
276:
271:
263:
261:
259:
255:
251:
247:
243:
239:
231:
229:
227:
223:
218:
214:
210:
206:
205:bacteriophage
202:
194:
192:
190:
186:
182:
178:
174:
170:
165:
162:
158:
153:
148:
146:
141:
137:
133:
128:
127:transcriptome
123:
120:
116:
115:
110:
106:
100:
92:
90:
88:
84:
80:
76:
72:
68:
64:
60:
59:messenger RNA
56:
52:
48:
44:
40:
39:Antisense RNA
32:
19:
1849:Bacterial
1824:Lambda phage
1611:
1488:Constituents
1438:List of RNAs
1309:
1248:Transfer RNA
1176:
1149:
1143:
1134:
1125:
1100:
1096:
1089:
1052:
1048:
1038:
1013:
1009:
1003:
978:
974:
968:
941:
937:
927:
892:
888:
878:
835:
831:
821:
786:
782:
732:
728:
674:
670:
612:
608:
558:
554:
496:
484:
480:
471:
438:
417:
385:
355:
347:
312:Trans-acting
310:
307:Trans-acting
297:
273:
258:trans-acting
235:
198:
166:
149:
124:
113:
102:
42:
38:
37:
1844:P1-derived
1612:Antisense
1505:Nucleotides
1500:Nucleosides
1495:Nucleobases
450:zinc-finger
71:prokaryotes
63:translation
1894:Categories
1796:Morpholino
1709:Organellar
1617:Processual
1582:Regulatory
1526:non-coding
1376:Riboswitch
1320:CRISPR RNA
1103:: 102560.
519:References
289:activators
275:Cis-acting
268:See also:
264:Cis-acting
254:cis-acting
246:epigenetic
169:mipomersen
157:fomivirsen
105:micF asRNA
75:eukaryotes
1756:Analogues
1741:Hachimoji
1524:(coding,
1479:Types of
1433:Vault RNA
1407:RNA virus
1390:Parasites
1289:RNase MRP
1279:Guide RNA
1217:Types of
938:P & T
285:silencers
242:cytoplasm
181:apo-B-100
145:chromatin
1879:Category
1814:Phagemid
1665:Ribozyme
1330:MicroRNA
1117:33482228
1081:27856735
1030:22785023
960:24669178
919:20707673
862:10228554
853:11146801
813:22262036
749:23722346
701:22541732
641:28536609
575:24217315
507:See also
491:microRNA
425:homeobox
388:histones
323:Function
209:bacteria
201:plasmids
152:Zamecnik
111:ompC in
1819:Plasmid
1284:RNase P
1097:Plasmid
1072:5137729
995:2468573
951:3956393
910:3030471
870:9448271
804:4743652
692:3768148
632:5415839
583:2152962
250:lncRNAs
238:nucleus
189:RNAse H
67:protein
1834:Fosmid
1829:Cosmid
1779:Hexose
1701:acids
1653:Others
1412:Viroid
1164:
1115:
1079:
1069:
1028:
993:
958:
948:
917:
907:
868:
860:
850:
811:
801:
757:288163
755:
747:
699:
689:
639:
629:
581:
573:
462:5' end
421:HOTAIR
405:CDKN2A
401:CDKN2B
280:RNAase
119:5' end
114:E.coli
1859:Human
1637:Y RNA
1421:Other
1294:Y RNA
866:S2CID
753:S2CID
615:: 6.
579:S2CID
370:ANRIL
109:porin
65:into
43:asRNA
1162:ISBN
1113:PMID
1077:PMID
1026:PMID
991:PMID
975:Gene
956:PMID
915:PMID
858:PMID
809:PMID
745:PMID
697:PMID
637:PMID
571:PMID
454:ZEB2
446:RME2
442:IME4
429:PRC2
413:XIST
300:loci
287:and
256:and
222:Xist
207:and
73:and
1219:RNA
1154:doi
1105:doi
1101:114
1067:PMC
1057:doi
1053:113
1018:doi
983:doi
946:PMC
905:PMC
897:doi
848:PMC
840:doi
799:PMC
791:doi
737:doi
687:PMC
679:doi
627:PMC
617:doi
563:doi
399:of
240:or
55:RNA
1896::
1160:.
1133:.
1111:.
1099:.
1075:.
1065:.
1051:.
1047:.
1024:.
1014:28
1012:.
989:.
979:72
977:.
954:.
942:39
940:.
936:.
913:.
903:.
893:44
891:.
887:.
864:.
856:.
846:.
836:55
834:.
830:.
807:.
797:.
787:11
785:.
781:.
765:^
751:.
743:.
733:12
731:.
709:^
695:.
685:.
675:28
673:.
669:.
649:^
635:.
625:.
613:21
611:.
607:.
591:^
577:.
569:.
559:14
557:.
527:^
415:.
403:,
260:.
203:,
1528:)
1472:e
1465:t
1458:v
1210:e
1203:t
1196:v
1170:.
1156::
1119:.
1107::
1083:.
1059::
1032:.
1020::
997:.
985::
962:.
921:.
899::
872:.
842::
815:.
793::
759:.
739::
703:.
681::
643:.
619::
585:.
565::
331:'
41:(
20:)
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