646:
phenotype with a second, non-overlapping
Morpholino targeting the same mRNA, by confirmation of the observed phenotypes by comparing with a mutant strain (though compensation will obscure a phenotype in some mutants), by testing the Morpholino in a null mutant background to detect additional phenotypic changes or by dominant-negative methods. As mentioned above, rescue of observed phenotypes by coinjecting a rescue mRNA is, when feasible, a reliable test of specificity of a Morpholino.
3634:
407:
458:
31:
339:
438:). This is useful experimentally when an investigator wishes to know the function of a particular protein; Morpholinos provide a convenient means of knocking down expression of the protein and learning how that knockdown changes the cells or organism. Some Morpholinos knock down expression so effectively that, after degradation of preexisting proteins, the targeted proteins become undetectable by
702:). Though they permeate through intercellular spaces in tissues effectively, unconjugated PMOs have limited distribution into the cytosol and nuclear spaces within healthy tissues following IV administration. Systemic delivery into many cells in adult organisms can be accomplished by using covalent conjugates of Morpholino oligos with
108:, but this usage is not consistent across scientific literature. Morpholino oligos are sometimes referred to as PMO (for phosphorodiamidate morpholino oligomer), especially in medical literature. Vivo-Morpholinos and PPMO are modified forms of Morpholinos with chemical groups covalently attached to facilitate entry into cells.
114:
is achieved by reducing the expression of a particular gene in a cell. In the case of protein-coding genes, this usually leads to a reduction in the quantity of the corresponding protein in the cell. Knocking down gene expression is a method for learning about the function of a particular protein; in
637:
using another antisense structural type, showing the p53-mediated apoptosis to be a consequence of the loss of the targeted protein and not a consequence of the knockdown oligo type. It appears that these effects are sequence-specific; as in most cases, if a
Morpholino is associated with non-target
581:
oligos with irrelevant sequences usually produce no change in embryonic phenotype, evidence of the
Morpholino oligo's sequence-specificity and lack of non-antisense effects. The dose required for a knockdown can be reduced by coinjection of several Morpholino oligos targeting the same mRNA, which is
598:
encodes the protein of interest. Translation of the rescue mRNA replaces production of the protein that was knocked down by the
Morpholino. Since the rescue mRNA would not affect phenotypic changes due to the Morpholino's off-target gene expression modulation, this return to wild-type phenotype is
658:
into the cytosol of a cell. Once in the cytosol, Morpholinos freely diffuse between the cytosol and nucleus, as demonstrated by the nuclear splice-modifying activity of
Morpholinos observed after microinjection into the cytosol of cells. Different methods are used for delivery into embryos, into
602:
In embryos, Morpholinos can be tested in null mutants to check for unexpected RNA interactions, then used in a wild-type embryo to reveal the acute knockdown phenotype. The knockdown phenotype is often more extreme than the mutant phenotype; in the mutant, effects of losing the null gene can be
645:
phenotype is due to the intended knockdown or an interaction with an off-target RNA can often be addressed in embryos by running another experiment to confirm that the observed morphant phenotype results from the knockdown of the expected target. This can be done by recapitulating the morphant
710:
for effective oligo delivery at doses below those causing observed toxicity. An octa-guanidinium dendrimer attached to the end of a
Morpholino can deliver the modified oligo (called a Vivo-Morpholino) from the blood to the cytosol. Delivery-enabled Morpholinos, such as peptide conjugates and
548:. Morpholinos can block RNA editing, poly(A) tailing and translocation sequences. Morpholino activities against this variety of targets suggest that Morpholinos can be used as a general-purpose tool for blocking interactions of proteins or nucleic acids with mRNA.
280:). Instead, Morpholinos act by "steric blocking", binding to a target sequence within an RNA, inhibiting molecules that might otherwise interact with the RNA. Morpholino oligos are often used to investigate the role of a specific mRNA transcript in an
2962:
Abes S, Moulton HM, Clair P, Prevot P, Youngblood DS, Wu RP, Iversen PL, Lebleu B (December 2006). "Vectorization of morpholino oligomers by the (R-Ahx-R)4 peptide allows efficient splicing correction in the absence of endosomolytic agents".
564:
and can be strongly affected by an off-target interaction. Following initial injections into frog or fish embryos at the single-cell or few-cell stages, Morpholino effects can be measured up to five days later, after most of the processes of
3047:
Amantana A, Moulton HM, Cate ML, Reddy MT, Whitehead T, Hassinger JN, Youngblood DS, Iversen PL (2007). "Pharmacokinetics, biodistribution, stability and toxicity of a cell-penetrating peptide-morpholino oligomer conjugate".
589:
phenotype to the embryos and provide evidence for the specificity of a
Morpholino. In an mRNA rescue, a Morpholino is co-injected with an mRNA that codes for the morphlino's protein. However, the rescue mRNA has a modified
2877:
Fletcher S, Honeyman K, Fall AM, Harding PL, Johnsen RD, Wilton SD (February 2006). "Dystrophin expression in the mdx mouse after localised and systemic administration of a morpholino antisense oligonucleotide".
2461:
Stainier DY, Raz E, Lawson ND, Ekker SC, Burdine RD, Eisen JS, Ingham PW, Schulte-Merker S, Yelon D, Weinstein BM, Mullins MC, Wilson SW, Ramakrishnan L, Amacher SL, Neuhauss SC, Meng A, Mochizuki N, Panula P,
1510:
Kos R, Reedy MV, Johnson RL, Erickson CA (April 2001). "The winged-helix transcription factor FoxD3 is important for establishing the neural crest lineage and repressing melanogenesis in avian embryos".
154:
or inhibit the maturation and activity of miRNA. Techniques for targeting
Morpholinos to RNAs and delivering Morpholinos into cells have recently been reviewed in a journal article and in book form.
667:, which can deliver oligos into tissues of later embryonic stages. Common techniques for delivery into cultured cells include the Endo-Porter peptide (which causes the Morpholino to be released from
2773:
Jubin R (2004). "Optimizing electroporation conditions for intracellular delivery of
Morpholino antisense oligonucleotides directed against the hepatitis C virus internal ribosome entry site".
2423:
Kamachi Y, Okuda Y, Kondoh H (January 2008). "Quantitative assessment of the knockdown efficiency of morpholino antisense oligonucleotides in zebrafish embryos using a luciferase assay".
2069:
Yen L, Svendsen J, Lee JS, Gray JT, Magnier M, Baba T, D'Amato RJ, Mulligan RC (September 2004). "Exogenous control of mammalian gene expression through modulation of RNA self-cleavage".
1475:
Howard EW, Newman LA, Oleksyn DW, Angerer RC, Angerer LM (February 2001). "SpKrl: a direct target of beta-catenin regulation required for endoderm differentiation in sea urchin embryos".
1135:
398:, is converted into the structure of a protein. A Morpholino can modify splicing, block translation, or block other functional sites on RNA depending on the Morpholino's base sequence.
633:
and can be suppressed by co-injection of an anti-p53 Morpholino along with the experimental
Morpholino. Moreover, the p53-mediated apoptotic effect of a Morpholino knockdown has been
663:
apparatus is usually used for delivery into an embryo, with injections most commonly performed at the single-cell or few-cell stage; an alternative method for embryonic delivery is
504:
from the mature mRNA. Targeting some splice targets results in intron inclusions, while activation of cryptic splice sites can lead to partial inclusions or exclusions. Targets of
544:
activity. U2 and U12 snRNP functions have been inhibited by Morpholinos. Morpholinos targeted to "slippery" mRNA sequences within protein coding regions can induce translational
2552:
Youngblood DS, Hatlevig SA, Hassinger JN, Iversen PL, Moulton HM (2007). "Stability of cell-penetrating peptide-morpholino oligomer conjugates in human serum and in cells".
2921:
Kipshidze NN, Kim HS, Iversen P, Yazdi HA, Bhargava B, New G, Mehran R, Tio F, Haudenschild C, Dangas G, Stone GW, Iyer S, Roubin GS, Leon MB, Moses JW (May 2002).
2834:
Partridge M, Vincent A, Matthews P, Puma J, Stein D, Summerton J (1996). "A simple method for delivering morpholino antisense oligos into the cytoplasm of cells".
1071:
McClorey G, Fall AM, Moulton HM, Iversen PL, Rasko JE, Ryan M, Fletcher S, Wilton SD (October 2006). "Induced dystrophin exon skipping in human muscle explants".
3083:
Li YF, Morcos PA (July 2008). "Design and synthesis of dendritic molecular transporter that achieves efficient in vivo delivery of morpholino antisense oligo".
480:
base and preventing it from forming the splice lariat structure, or by interfering with the binding of splice regulatory proteins such as splice silencers and
2517:
Hudziak RM, Barofsky E, Barofsky DF, Weller DL, Huang SB, Weller DD (1996). "Resistance of morpholino phosphorodiamidate oligomers to enzymatic degradation".
599:
further evidence of Morpholino specificity. In some cases, ectopic expression of the rescue RNA makes recovery of the wild-type phenotype impossible.
1709:"Peptide-conjugated phosphorodiamidate morpholino oligomer (PPMO) restores carbapenem susceptibility to NDM-1-positive pathogens in vitro and in vivo"
3179:
that consists of a series of peer-reviewed short papers using Morpholino knock downs of gene function in various animal and tissue culture systems.
264:
range, so Morpholinos in organisms or cells are uncharged molecules. The entire backbone of a Morpholino is made from these modified subunits.
2728:
Cerda GA, Thomas JE, Allende ML, Karlstrom RO, Palma V (July 2006). "Electroporation of DNA, RNA, and morpholinos into zebrafish embryos".
2923:"Intramural coronary delivery of advanced antisense oligonucleotides reduces neointimal formation in the porcine stent restenosis model"
104:
ring. To help avoid confusion with other morpholine-containing molecules, when describing oligos "Morpholino" is often capitalized as a
686:
is usually difficult, though there are a few systems allowing useful uptake of unmodified Morpholino oligos (including uptake into
236:
structure. Usually 25 bases in length, they bind to complementary sequences of RNA or single-stranded DNA by standard nucleic acid
1799:"Aberrant splicing in the ocular albinism type 1 gene (OA1/GPR143) is corrected in vitro by morpholino antisense oligonucleotides"
126:
by that exon or can sometimes knock down the protein activity altogether. These molecules have been applied to studies in several
3229:
256:. The figure compares the structures of the two strands depicted there, one of RNA and the other of a Morpholino. Replacement of
512:
snRNPs can also be blocked. Splice modification can be conveniently assayed by reverse-transcriptase polymerase chain reaction (
3197:
2790:
1707:
Sully EK, Geller BL, Li L, Moody CM, Bailey SM, Moore AL, Wong M, Nordmann P, Daly SM, Sturge CR, Greenberg DE (March 2017).
1286:
968:
1022:
Deas TS, Bennett CJ, Jones SA, Tilgner M, Ren P, Behr MJ, Stein DA, Iversen PL, Kramer LD, Bernard KA, Shi PY (July 2007).
81:
groups. Morpholinos block access of other molecules to small (~25 base) specific sequences of the base-pairing surfaces of
2220:"Steric antisense inhibition of AMPA receptor Q/R editing reveals tight coupling to intronic editing sites and splicing"
272:
Morpholinos do not trigger the degradation of their target RNA molecules, unlike many antisense structural types (e.g.,
446:
3315:
834:"Targeted inhibition of miRNA maturation with morpholinos reveals a role for miR-375 in pancreatic islet development"
1587:
Summerton JE (November 2005). "Endo-Porter: a novel reagent for safe, effective delivery of substances into cells".
1356:
Summerton J (December 1999). "Morpholino antisense oligomers: the case for an RNase H-independent structural type".
3602:
3182:"Peptide Nucleic Acids, Morpholinos and Related Antisense Biomolecules." eds. Janson & During (Springer, 2007)
3607:
3173:
Wiley-Liss, Inc. Special Issue: Morpholino Gene Knockdowns of genesis Volume 30, Issue 3 Pages 89-200 (July 2001)
1846:
Morcos PA (June 2007). "Achieving targeted and quantifiable alteration of mRNA splicing with Morpholino oligos".
790:"Inhibition of zebrafish fgf8 pre-mRNA splicing with morpholino oligos: a quantifiable method for gene knockdown"
2022:"Revealing details: whole mount microRNA in situ hybridization protocol for zebrafish embryos and adult tissues"
473:) complexes from binding to their targets at the borders of introns on a strand of pre-mRNA, or by blocking the
907:
899:
591:
390:
begins. This entire process is referred to as gene expression; it is the process by which the information in a
316:
embryos at later development stages. With appropriate cytosolic delivery systems, Morpholinos are effective in
641:
A cause for concern in the use of Morpholinos is the potential for "off-target" effects. Whether an observed
3504:
3390:
706:, and, while toxicity has been associated with moderate doses of the peptide conjugates, they have been used
119:
to be spliced out of the RNA transcript encoding a protein can help to determine the function of the protein
1682:
100:. The word "Morpholino" can occur in other chemical names, referring to chemicals containing a six-membered
3660:
1136:"FDA grants accelerated approval to first targeted treatment for rare Duchenne muscular dystrophy mutation"
1119:
1114:
375:
186:
1313:
Summerton J, Weller D (June 1997). "Morpholino antisense oligomers: design, preparation, and properties".
1269:
Summerton JE (2017). "Invention and Early History of Morpholinos: From Pipe Dream to Practical Products".
3670:
3617:
3612:
3597:
3222:
1758:"Correction of aberrant FGFR1 alternative RNA splicing through targeting of intronic regulatory elements"
691:
260:
phosphates with the uncharged phosphorodiamidate groups eliminates ionization in the usual physiological
190:
378:, forming the initiation complex. The initiation complex scans along the mRNA strand until it reaches a
594:(untranslated region) so that the rescue mRNA contains no target for the Morpholino. The rescue mRNA's
509:
217:) at AntiVirals Inc. (now Sarepta Therapeutics) and originally developed in collaboration with Weller.
2267:
Vorlová S, Rocco G, Lefave CV, Jodelka FM, Hess K, Hastings ML, Henke E, Cartegni L (September 2011).
1883:"Splicing segregation: the minor spliceosome acts outside the nucleus and controls cell proliferation"
1797:
Vetrini F, Tammaro R, Bondanza S, Surace EM, Auricchio A, De Luca M, Ballabio A, Marigo V (May 2006).
606:
Because of their completely unnatural backbones, Morpholinos are not recognized by cellular proteins.
3665:
1643:"Vivo-Morpholinos: a non-peptide transporter delivers Morpholinos into a wide array of mouse tissues"
1115:"Press Announcements - FDA grants accelerated approval to first drug for Duchenne muscular dystrophy"
720:
226:
2820:
1024:"In vitro resistance selection and in vivo efficacy of morpholino oligomers against West Nile virus"
3685:
3186:
Moulton J (2007). "Using Morpholinos to Control Gene Expression (Unit 4.30)". In Beaucage S (ed.).
1436:"Beta-catenin signaling activity dissected in the early Xenopus embryo: a novel antisense approach"
926:
703:
570:
481:
427:
351:
2998:
Burrer R, Neuman BW, Ting JP, Stein DA, Moulton HM, Iversen PL, Kuhn P, Buchmeier MJ (June 2007).
3172:
2269:"Induction of antagonistic soluble decoy receptor tyrosine kinases by intronic polyA activation"
3690:
3675:
3637:
3330:
3215:
618:
244:
is that, while Morpholinos have standard nucleic acid bases, those bases are bound to methylene
97:
1549:
Morcos PA (July 2001). "Achieving efficient delivery of morpholino oligos in cultured cells".
582:
an effective strategy for reducing or eliminating dose-dependent off-target RNA interactions.
3514:
3358:
2171:"Efficient stimulation of site-specific ribosome frameshifting by antisense oligonucleotides"
876:
725:
537:
423:
383:
3000:"Antiviral effects of antisense morpholino oligomers in murine coronavirus infection models"
536:-tagged Morpholinos combined with fluorescein-specific antibodies can be used as probes for
3680:
3549:
3532:
3363:
2078:
1596:
1391:
Nasevicius A, Ekker SC (October 2000). "Effective targeted gene 'knockdown' in zebrafish".
578:
505:
182:
2318:"Participation of Xenopus Elr-type proteins in vegetal mRNA localization during oogenesis"
8:
3544:
3527:
3385:
3268:
2628:
Robu ME, Larson JD, Nasevicius A, Beiraghi S, Brenner C, Farber SA, Ekker SC (May 2007).
545:
517:
497:
493:
489:
485:
469:
processing steps either by preventing splice-directing small nuclear ribonucleoproteins (
445:
In 2016 a synthetic peptide-conjugated PMO (PPMO) was found to inhibit the expression of
374:
usually starts by binding at the 5' end of the mRNA and is joined there by various other
359:
147:
74:
2082:
1600:
1273:. Methods in Molecular Biology. Vol. 1565. Humana Press (Springer). pp. 1–15.
671:), the Special Delivery system (no longer commercially available, used a Morpholino-DNA
3438:
3433:
3308:
3146:
3119:
3024:
2999:
2903:
2859:
2808:
2705:
2680:
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2400:
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2219:
2195:
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2102:
2046:
2021:
1997:
1972:
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1708:
1620:
1416:
1338:
1251:
1196:
1096:
1048:
1023:
974:
860:
833:
193:, although the approval process was mired in controversy. Other Morpholino-based drugs
120:
78:
2939:
2922:
2146:
2121:
1948:
1923:
1369:
955:. Methods in Molecular Biology. Vol. 1565. Humana Press (Springer). p. 284.
77:
contains DNA bases attached to a backbone of methylenemorpholine rings linked through
3522:
3484:
3477:
3428:
3380:
3193:
3187:
3151:
3100:
3065:
3029:
2980:
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2534:
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2440:
2405:
2349:
2298:
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2200:
2151:
2094:
2051:
2002:
1953:
1904:
1863:
1820:
1779:
1738:
1664:
1612:
1566:
1528:
1492:
1457:
1408:
1373:
1330:
1292:
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1255:
1243:
1235:
1200:
1188:
1180:
1088:
1053:
1001:
964:
888:
865:
811:
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324:, enter cells when administered systemically in adult animals or in tissue cultures.
66:
2907:
1832:
1624:
1420:
1342:
1100:
978:
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3141:
3131:
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1992:
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2284:
884:
850:
832:
Kloosterman WP, Lagendijk AK, Ketting RF, Moulton JD, Plasterk RH (August 2007).
664:
613:
Up to 18% of Morpholinos appear to induce nontarget-related phenotypes including
435:
333:
309:
249:
70:
2782:
2741:
1278:
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1899:
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1231:
711:
Vivo-Morpholinos, show promise as therapeutics for viral and genetic diseases.
660:
561:
431:
382:, and then the large subunit of the ribosome attaches to the small subunit and
127:
111:
90:
3136:
1924:"Substrate requirements for let-7 function in the developing zebrafish embryo"
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960:
625:
tissues of zebrafish embryos. Most of these effects are due to activation of
205:(also for Duchenne muscular dystrophy) were approved by the FDA in 2019–2021.
3654:
3499:
3489:
3370:
3325:
3303:
2847:
2530:
2463:
2316:
Arthur PK, Claussen M, Koch S, Tarbashevich K, Jahn O, Pieler T (July 2009).
1326:
1239:
1184:
655:
595:
566:
273:
2334:
1973:"Zebrafish miR-214 modulates Hedgehog signaling to specify muscle cell fate"
1608:
1215:
1160:
284:. Developmental biologists inject Morpholino oligos into eggs or embryos of
3582:
3402:
3375:
3320:
3239:
3155:
3104:
3069:
3033:
2984:
2948:
2899:
2800:
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2714:
2665:
2614:
2573:
2503:
2444:
2409:
2353:
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2155:
2098:
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2006:
1957:
1908:
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1092:
1057:
1005:
892:
869:
815:
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of messenger RNA (mRNA), Morpholinos can interfere with progression of the
363:
317:
233:
158:
123:
2855:
2538:
2391:
2235:
1971:
Flynt AS, Li N, Thatcher EJ, Solnica-Krezel L, Patton JG (February 2007).
1488:
1334:
449:, an enzyme that many drug-resistant bacteria use to destroy carbapenems.
320:. Vivo-Morpholinos, in which the oligo is covalently linked to a delivery
96:
This article discusses only the Morpholino antisense oligomers, which are
3494:
3340:
3207:
3015:
2137:
1939:
1774:
1757:
1724:
1039:
699:
695:
533:
474:
379:
285:
198:
30:
3295:
2750:
2186:
2090:
2037:
638:
effects, the 4-base mismatch Morpholino will not trigger these effects.
610:
do not degrade Morpholinos, nor are they degraded in serum or in cells.
3263:
3258:
3253:
1815:
1798:
896:. If this is an intentional citation to a such a paper, please replace
614:
406:
295:
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143:
105:
101:
20:
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2436:
1659:
1642:
1562:
806:
789:
3418:
2374:
Bill BR, Petzold AM, Clark KJ, Schimmenti LA, Ekker SC (March 2009).
634:
630:
586:
574:
422:
initiation complex from the 5' cap to the start codon. This prevents
415:
367:
347:
321:
300:
237:
135:
2891:
496:(at the polypyrimidine moiety and acceptor site) can cause modified
457:
3572:
3423:
3353:
1988:
1921:
1358:
Biochimica et Biophysica Acta (BBA) - Gene Structure and Expression
668:
642:
607:
551:
541:
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419:
371:
305:
229:
166:
162:
151:
58:
55:
24:
2551:
1404:
831:
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systems, which have a broader range of gene expression than adult
338:
3577:
2696:
2122:"Targeted 'knockdown' of spliceosome function in mammalian cells"
477:
387:
313:
290:
139:
1881:
König H, Matter N, Bader R, Thiele W, Müller F (November 2007).
654:
For a Morpholino to be effective, it must be delivered past the
240:. In terms of structure, the difference between Morpholinos and
3592:
3587:
3537:
687:
622:
557:
524:
Other applications: blocking other mRNA sites and use as probes
513:
355:
342:
Eukaryotic gene expression without intervention by a Morpholino
281:
3120:"Gene knockdowns in adult animals: PPMOs and vivo-morpholinos"
2681:"Microinjection of zebrafish embryos to analyze gene function"
2315:
1161:"Viltolarsen for the treatment of Duchenne muscular dystrophy"
189:
in September 2016 for the treatment of some mutations causing
3395:
2516:
1970:
1922:
Kloosterman WP, Wienholds E, Ketting RF, Plasterk RH (2004).
529:
470:
277:
170:
131:
2833:
2627:
1796:
1683:"New molecule knocks out superbugs' immunity to antibiotics"
556:
Morpholinos have become a standard knockdown tool in animal
2876:
2727:
501:
391:
116:
2373:
1474:
1070:
3457:
3278:
2266:
1262:
626:
395:
257:
241:
82:
3046:
2920:
2019:
1509:
1021:
679:
delivery reagent), electroporation, or scrape loading.
327:
261:
2961:
2460:
1880:
2997:
2168:
34:
Segment of a Morpholino-RNA heteroduplex, 8-mer shown
2589:"Morphant technology in model developmental systems"
2169:Howard MT, Gesteland RF, Atkins JF (October 2004).
1848:
Biochemical and Biophysical Research Communications
992:Geller BL (April 2005). "Antibacterial antisense".
944:
85:(RNA). Morpholinos are used as research tools for
2422:
2369:
2367:
2365:
2363:
2068:
1706:
1433:
1308:
1306:
787:
585:mRNA rescue experiments can sometimes restore the
2678:
2020:Lagendijk AK, Moulton JD, Bakkers J (June 2012).
927:"Guide for Morpholino Users: Toward Therapeutics"
3652:
552:Specificity, stability and non-antisense effects
2456:
2454:
2360:
2217:
1390:
1312:
1303:
746:"Morpholino oligos: making sense of antisense?"
362:out, then the mature mRNA is exported from the
213:Morpholino oligos were conceived by Summerton (
950:
452:
232:that are the product of a redesign of natural
3223:
2927:Journal of the American College of Cardiology
2836:Antisense & Nucleic Acid Drug Development
2679:Rosen JN, Sweeney MF, Mably JD (March 2009).
2519:Antisense & Nucleic Acid Drug Development
1640:
1315:Antisense & Nucleic Acid Drug Development
827:
825:
788:Draper BW, Morcos PA, Kimmel CB (July 2001).
3117:
2468:"Guidelines for morpholino use in zebrafish"
2451:
2218:Penn AC, Balik A, Greger IH (January 2013).
1755:
1636:
1634:
1582:
1580:
1158:
1133:
3189:Current Protocols in Nucleic Acid Chemistry
2586:
2119:
3237:
3230:
3216:
3192:. New Jersey: John Wiley & Sons, Inc.
2630:"p53 activation by knockdown technologies"
2376:"A primer for morpholino use in zebrafish"
1641:Morcos PA, Li Y, Jiang S (December 2008).
1589:Annals of the New York Academy of Sciences
1434:Heasman J, Kofron M, Wylie C (June 2000).
1159:Roshmi, R. R.; Yokota, T. (October 2019).
1134:Commissioner, Office of the (2019-12-12).
910:|...|intentional=yes}}
822:
3145:
3135:
3023:
2938:
2749:
2704:
2655:
2645:
2604:
2493:
2483:
2399:
2343:
2333:
2292:
2243:
2194:
2145:
2045:
1996:
1947:
1898:
1814:
1773:
1756:Bruno IG, Jin W, Cote GJ (October 2004).
1732:
1713:The Journal of Antimicrobial Chemotherapy
1658:
1631:
1586:
1577:
1544:
1542:
1451:
1355:
1268:
1047:
994:Current Opinion in Molecular Therapeutics
859:
849:
805:
761:
410:Translation blocked by a Morpholino oligo
3082:
783:
781:
659:cultured cells or into adult animals. A
456:
405:
337:
29:
3185:
1213:
1064:
924:
743:
577:consistent with target-gene knockdown.
401:
185:received accelerated approval from the
3653:
1845:
1548:
1539:
1349:
1017:
1015:
991:
461:Splicing blocked by a Morpholino oligo
48:phosphorodiamidate Morpholino oligomer
3211:
2772:
1028:Antimicrobial Agents and Chemotherapy
951:Moulton HM, Moulton JD, eds. (2017).
778:
426:of the coding region of the targeted
115:a similar manner, causing a specific
918:
528:Morpholinos have been used to block
516:) and is seen as a band shift after
394:, encoded as a sequence of bases in
328:Normal gene expression in eukaryotes
2322:The Journal of Biological Chemistry
2120:Matter N, König H (February 2005).
1012:
603:concealed by genetic compensation.
13:
3165:
3118:Moulton JD, Jiang S (March 2009).
2777:. Vol. 106. pp. 309–22.
484:. Preventing the binding of snRNP
157:Morpholinos are in development as
146:. Morpholinos can also modify the
14:
3702:
3354:Micro
2685:Journal of Visualized Experiments
2587:Ekker SC, Larson JD (July 2001).
540:to miRNAs. Morpholinos can block
3633:
3632:
447:New Delhi Metallo-beta-lactamase
3309:precursor, heterogenous nuclear
3111:
3076:
3040:
2991:
2955:
2914:
2870:
2827:
2766:
2721:
2672:
2621:
2580:
2545:
2510:
2416:
2309:
2260:
2211:
2162:
2113:
2062:
2013:
1964:
1915:
1874:
1839:
1790:
1749:
1700:
1675:
1503:
1468:
1427:
1384:
1207:
1152:
1127:
934:J Drug Discov Develop and Deliv
465:Morpholinos can interfere with
187:US Food and Drug Administration
3439:Trans-acting small interfering
3403:Enhancer RNAs
3321:Transfer
1177:10.1358/dot.2019.55.10.3045038
1107:
985:
737:
698:cells stressed during balloon
161:therapeutics targeted against
1:
3326:Ribosomal
3304:Messenger
3175:. This is a special issue of
2977:10.1016/j.jconrel.2006.09.011
2965:Journal of Controlled Release
2940:10.1016/S0735-1097(02)01830-2
1653:(6): 613–4, 616, 618 passim.
1370:10.1016/S0167-4781(99)00150-5
875:(This paper currently has an
731:
376:eukaryotic initiation factors
2880:The Journal of Gene Medicine
2647:10.1371/journal.pgen.0030078
2485:10.1371/journal.pgen.1007000
2285:10.1016/j.molcel.2011.08.009
1216:"Casimersen: First Approval"
1120:Food and Drug Administration
885:10.1371/journal.pbio.3001631
851:10.1371/journal.pbio.0050203
220:
7:
2742:10.1016/j.ymeth.2005.12.009
1279:10.1007/978-1-4939-6817-6_1
714:
692:Duchenne muscular dystrophy
649:
453:Modifying pre-mRNA splicing
370:. The small subunit of the
267:
191:Duchenne muscular dystrophy
10:
3707:
3505:Multicopy single-stranded
3349:Interferential
1900:10.1016/j.cell.2007.09.043
1860:10.1016/j.bbrc.2007.04.172
1232:10.1007/s40265-021-01512-2
1214:Shirley, Matt (May 2021).
532:activity and maturation.
331:
208:
177:. A Morpholino-based drug
18:
3628:
3563:
3513:
3456:
3419:Guide
3411:
3339:
3294:
3277:
3246:
3137:10.3390/molecules14031304
2783:10.1385/1-59259-854-4:309
1085:10.1016/j.nmd.2006.05.017
961:10.1007/978-1-4939-6817-6
721:Oligonucleotide synthesis
704:cell-penetrating peptides
3381:Small nuclear
2848:10.1089/oli.1.1996.6.169
2531:10.1089/oli.1.1996.6.267
1762:Human Molecular Genetics
1327:10.1089/oli.1.1997.7.187
744:Heasman J (March 2002).
573:are past, with observed
19:Not to be confused with
3495:Genomic
2335:10.1074/jbc.M109.009928
1609:10.1196/annals.1359.012
1073:Neuromuscular Disorders
488:(at the donor site) or
350:organisms, pre-mRNA is
288:, African clawed frog (
3598:Artificial chromosomes
3386:Small nucleolar
3085:Bioconjugate Chemistry
3050:Bioconjugate Chemistry
2775:Antisense Therapeutics
2554:Bioconjugate Chemistry
2224:Nucleic Acids Research
2126:Nucleic Acids Research
1928:Nucleic Acids Research
1525:10.1242/dev.128.8.1467
1453:10.1006/dbio.2000.9720
763:10.1006/dbio.2001.0565
619:central nervous system
462:
416:5'-untranslated region
411:
343:
35:
3391:Small Cajal Body RNAs
2392:10.1089/zeb.2008.0555
1489:10.1242/dev.128.3.365
1440:Developmental Biology
908:expression of concern
902:|...}}
900:expression of concern
877:expression of concern
750:Developmental Biology
726:Nucleic acid analogue
538:in-situ hybridization
500:, commonly excluding
460:
409:
341:
248:rings linked through
33:
3444:Subgenomic messenger
3359:Small interfering
3331:Transfer-messenger
3016:10.1128/JVI.02360-06
1271:Morpholino Oligomers
1040:10.1128/AAC.00069-07
953:Morpholino Oligomers
696:vascular endothelial
682:Delivery into adult
520:of RT-PCR products.
402:Blocking translation
183:Sarepta Therapeutics
98:nucleic acid analogs
3661:Genetics techniques
3004:Journal of Virology
2236:10.1093/nar/gks1044
2187:10.1261/rna.7810204
2091:10.1038/nature02844
2083:2004Natur.431..471Y
2038:10.1242/bio.2012810
1601:2005NYASA1058...62S
925:Moulton JD (2016).
675:and an ethoxylated
518:gel electrophoresis
75:molecular structure
44:Morpholino oligomer
3671:Molecular genetics
3473:Chloroplast
3316:modified Messenger
3279:Ribonucleic acids
2138:10.1093/nar/gni041
1940:10.1093/nar/gkh968
1816:10.1002/humu.20303
1775:10.1093/hmg/ddh272
1725:10.1093/jac/dkw476
463:
412:
344:
252:groups instead of
250:phosphorodiamidate
165:organisms such as
79:phosphorodiamidate
61:(colloquially, an
42:, also known as a
36:
3646:
3645:
3523:Xeno
3485:Complementary
3458:Deoxyribonucleic
3452:
3451:
3429:Small hairpin
3199:978-0-471-24662-6
3097:10.1021/bc8001437
3062:10.1021/bc070060v
2792:978-1-59259-854-0
2606:10.1002/gene.1038
2566:10.1021/bc060138s
2437:10.1002/dvg.20361
1660:10.2144/000113005
1563:10.1002/gene.1039
1288:978-1-4939-6817-6
970:978-1-4939-6815-2
807:10.1002/gene.1053
312:Morpholinos into
274:phosphorothioates
67:molecular biology
16:Chemical compound
3698:
3666:Phosphoramidates
3636:
3635:
3613:Yeast
3434:Small temporal
3364:Piwi-interacting
3292:
3291:
3288:
3269:Deoxynucleotides
3232:
3225:
3218:
3209:
3208:
3203:
3160:
3159:
3149:
3139:
3115:
3109:
3108:
3080:
3074:
3073:
3044:
3038:
3037:
3027:
2995:
2989:
2988:
2959:
2953:
2952:
2942:
2918:
2912:
2911:
2874:
2868:
2867:
2831:
2825:
2824:
2818:
2814:
2812:
2804:
2770:
2764:
2763:
2753:
2725:
2719:
2718:
2708:
2676:
2670:
2669:
2659:
2649:
2625:
2619:
2618:
2608:
2584:
2578:
2577:
2549:
2543:
2542:
2514:
2508:
2507:
2497:
2487:
2478:(10): e1007000.
2466:(October 2017).
2458:
2449:
2448:
2420:
2414:
2413:
2403:
2371:
2358:
2357:
2347:
2337:
2328:(30): 19982–92.
2313:
2307:
2306:
2296:
2264:
2258:
2257:
2247:
2215:
2209:
2208:
2198:
2166:
2160:
2159:
2149:
2117:
2111:
2110:
2066:
2060:
2059:
2049:
2017:
2011:
2010:
2000:
1968:
1962:
1961:
1951:
1919:
1913:
1912:
1902:
1878:
1872:
1871:
1843:
1837:
1836:
1818:
1794:
1788:
1787:
1777:
1753:
1747:
1746:
1736:
1704:
1698:
1697:
1695:
1694:
1679:
1673:
1672:
1662:
1638:
1629:
1628:
1584:
1575:
1574:
1546:
1537:
1536:
1507:
1501:
1500:
1472:
1466:
1465:
1455:
1431:
1425:
1424:
1388:
1382:
1381:
1353:
1347:
1346:
1310:
1301:
1300:
1266:
1260:
1259:
1211:
1205:
1204:
1156:
1150:
1149:
1147:
1146:
1131:
1125:
1124:
1111:
1105:
1104:
1079:(9–10): 583–90.
1068:
1062:
1061:
1051:
1019:
1010:
1009:
989:
983:
982:
948:
942:
941:
931:
922:
916:
915:
913:
911:
903:
873:
863:
853:
829:
820:
819:
809:
785:
776:
775:
765:
741:
677:polyethylenimine
482:splice enhancers
354:in the nucleus,
225:Morpholinos are
175:genetic diseases
87:reverse genetics
83:ribonucleic acid
54:), is a type of
3706:
3705:
3701:
3700:
3699:
3697:
3696:
3695:
3686:Gene expression
3651:
3650:
3647:
3642:
3624:
3565:Cloning vectors
3559:
3545:Locked
3509:
3459:
3448:
3407:
3335:
3282:
3281:
3273:
3242:
3236:
3206:
3200:
3168:
3166:Further reading
3163:
3116:
3112:
3081:
3077:
3045:
3041:
3010:(11): 5637–48.
2996:
2992:
2960:
2956:
2933:(10): 1686–91.
2919:
2915:
2892:10.1002/jgm.838
2875:
2871:
2832:
2828:
2816:
2815:
2806:
2805:
2793:
2771:
2767:
2726:
2722:
2677:
2673:
2626:
2622:
2585:
2581:
2550:
2546:
2515:
2511:
2459:
2452:
2421:
2417:
2372:
2361:
2314:
2310:
2265:
2261:
2216:
2212:
2181:(10): 1653–61.
2167:
2163:
2118:
2114:
2077:(7007): 471–6.
2067:
2063:
2018:
2014:
1977:Nature Genetics
1969:
1965:
1934:(21): 6284–91.
1920:
1916:
1879:
1875:
1844:
1840:
1795:
1791:
1768:(20): 2409–20.
1754:
1750:
1705:
1701:
1692:
1690:
1681:
1680:
1676:
1639:
1632:
1585:
1578:
1547:
1540:
1508:
1504:
1473:
1469:
1432:
1428:
1393:Nature Genetics
1389:
1385:
1354:
1350:
1311:
1304:
1289:
1267:
1263:
1212:
1208:
1171:(10): 627–639.
1157:
1153:
1144:
1142:
1132:
1128:
1113:
1112:
1108:
1069:
1065:
1020:
1013:
990:
986:
971:
949:
945:
929:
923:
919:
905:
897:
895:
874:
830:
823:
786:
779:
742:
738:
734:
717:
665:electroporation
652:
571:differentiation
554:
526:
455:
436:gene expression
404:
336:
334:Gene expression
330:
301:F. heteroclitus
298:and killifish (
270:
223:
211:
128:model organisms
93:gene function.
71:gene expression
28:
17:
12:
11:
5:
3704:
3694:
3693:
3688:
3683:
3678:
3673:
3668:
3663:
3644:
3643:
3641:
3640:
3629:
3626:
3625:
3623:
3622:
3621:
3620:
3615:
3610:
3605:
3595:
3590:
3585:
3580:
3575:
3569:
3567:
3561:
3560:
3558:
3557:
3552:
3550:Peptide
3547:
3542:
3541:
3540:
3535:
3530:
3528:Glycol
3519:
3517:
3511:
3510:
3508:
3507:
3502:
3497:
3492:
3487:
3482:
3481:
3480:
3475:
3464:
3462:
3454:
3453:
3450:
3449:
3447:
3446:
3441:
3436:
3431:
3426:
3421:
3415:
3413:
3409:
3408:
3406:
3405:
3400:
3399:
3398:
3393:
3388:
3383:
3373:
3368:
3367:
3366:
3361:
3356:
3345:
3343:
3337:
3336:
3334:
3333:
3328:
3323:
3318:
3313:
3312:
3311:
3300:
3298:
3289:
3275:
3274:
3272:
3271:
3266:
3261:
3256:
3250:
3248:
3244:
3243:
3240:nucleic acids
3235:
3234:
3227:
3220:
3212:
3205:
3204:
3198:
3183:
3180:
3169:
3167:
3164:
3162:
3161:
3130:(3): 1304–23.
3110:
3091:(7): 1464–70.
3075:
3056:(4): 1325–31.
3039:
2990:
2954:
2913:
2869:
2826:
2817:|journal=
2791:
2765:
2720:
2671:
2620:
2579:
2544:
2509:
2450:
2415:
2359:
2308:
2273:Molecular Cell
2259:
2230:(2): 1113–23.
2210:
2161:
2112:
2061:
2012:
1989:10.1038/ng1953
1963:
1914:
1873:
1838:
1803:Human Mutation
1789:
1748:
1719:(3): 782–790.
1699:
1674:
1630:
1576:
1538:
1519:(8): 1467–79.
1502:
1467:
1426:
1383:
1348:
1302:
1287:
1261:
1226:(7): 875–879.
1206:
1165:Drugs of Today
1151:
1126:
1106:
1063:
1034:(7): 2470–82.
1011:
984:
969:
943:
917:
821:
777:
735:
733:
730:
729:
728:
723:
716:
713:
661:microinjection
651:
648:
553:
550:
525:
522:
454:
451:
403:
400:
329:
326:
269:
266:
222:
219:
210:
207:
159:pharmaceutical
112:Gene knockdown
15:
9:
6:
4:
3:
2:
3703:
3692:
3691:Biotechnology
3689:
3687:
3684:
3682:
3679:
3677:
3676:Nucleic acids
3674:
3672:
3669:
3667:
3664:
3662:
3659:
3658:
3656:
3649:
3639:
3631:
3630:
3627:
3619:
3616:
3614:
3611:
3609:
3606:
3604:
3601:
3600:
3599:
3596:
3594:
3591:
3589:
3586:
3584:
3581:
3579:
3576:
3574:
3571:
3570:
3568:
3566:
3562:
3556:
3553:
3551:
3548:
3546:
3543:
3539:
3536:
3534:
3533:Threose
3531:
3529:
3526:
3525:
3524:
3521:
3520:
3518:
3516:
3512:
3506:
3503:
3501:
3498:
3496:
3493:
3491:
3490:Deoxyribozyme
3488:
3486:
3483:
3479:
3478:Mitochondrial
3476:
3474:
3471:
3470:
3469:
3466:
3465:
3463:
3461:
3455:
3445:
3442:
3440:
3437:
3435:
3432:
3430:
3427:
3425:
3422:
3420:
3417:
3416:
3414:
3410:
3404:
3401:
3397:
3394:
3392:
3389:
3387:
3384:
3382:
3379:
3378:
3377:
3374:
3372:
3369:
3365:
3362:
3360:
3357:
3355:
3352:
3351:
3350:
3347:
3346:
3344:
3342:
3338:
3332:
3329:
3327:
3324:
3322:
3319:
3317:
3314:
3310:
3307:
3306:
3305:
3302:
3301:
3299:
3297:
3296:Translational
3293:
3290:
3286:
3280:
3276:
3270:
3267:
3265:
3262:
3260:
3257:
3255:
3252:
3251:
3249:
3245:
3241:
3233:
3228:
3226:
3221:
3219:
3214:
3213:
3210:
3201:
3195:
3191:
3190:
3184:
3181:
3178:
3174:
3171:
3170:
3157:
3153:
3148:
3143:
3138:
3133:
3129:
3125:
3121:
3114:
3106:
3102:
3098:
3094:
3090:
3086:
3079:
3071:
3067:
3063:
3059:
3055:
3051:
3043:
3035:
3031:
3026:
3021:
3017:
3013:
3009:
3005:
3001:
2994:
2986:
2982:
2978:
2974:
2971:(3): 304–13.
2970:
2966:
2958:
2950:
2946:
2941:
2936:
2932:
2928:
2924:
2917:
2909:
2905:
2901:
2897:
2893:
2889:
2886:(2): 207–16.
2885:
2881:
2873:
2865:
2861:
2857:
2853:
2849:
2845:
2842:(3): 169–75.
2841:
2837:
2830:
2822:
2810:
2802:
2798:
2794:
2788:
2784:
2780:
2776:
2769:
2761:
2757:
2752:
2747:
2743:
2739:
2736:(3): 207–11.
2735:
2731:
2724:
2716:
2712:
2707:
2702:
2698:
2694:
2690:
2686:
2682:
2675:
2667:
2663:
2658:
2653:
2648:
2643:
2639:
2635:
2634:PLOS Genetics
2631:
2624:
2616:
2612:
2607:
2602:
2598:
2594:
2590:
2583:
2575:
2571:
2567:
2563:
2559:
2555:
2548:
2540:
2536:
2532:
2528:
2525:(4): 267–72.
2524:
2520:
2513:
2505:
2501:
2496:
2491:
2486:
2481:
2477:
2473:
2472:PLOS Genetics
2469:
2465:
2457:
2455:
2446:
2442:
2438:
2434:
2430:
2426:
2419:
2411:
2407:
2402:
2397:
2393:
2389:
2385:
2381:
2377:
2370:
2368:
2366:
2364:
2355:
2351:
2346:
2341:
2336:
2331:
2327:
2323:
2319:
2312:
2304:
2300:
2295:
2290:
2286:
2282:
2279:(6): 927–39.
2278:
2274:
2270:
2263:
2255:
2251:
2246:
2241:
2237:
2233:
2229:
2225:
2221:
2214:
2206:
2202:
2197:
2192:
2188:
2184:
2180:
2176:
2172:
2165:
2157:
2153:
2148:
2143:
2139:
2135:
2131:
2127:
2123:
2116:
2108:
2104:
2100:
2096:
2092:
2088:
2084:
2080:
2076:
2072:
2065:
2057:
2053:
2048:
2043:
2039:
2035:
2031:
2027:
2023:
2016:
2008:
2004:
1999:
1994:
1990:
1986:
1983:(2): 259–63.
1982:
1978:
1974:
1967:
1959:
1955:
1950:
1945:
1941:
1937:
1933:
1929:
1925:
1918:
1910:
1906:
1901:
1896:
1893:(4): 718–29.
1892:
1888:
1884:
1877:
1869:
1865:
1861:
1857:
1853:
1849:
1842:
1834:
1830:
1826:
1822:
1817:
1812:
1808:
1804:
1800:
1793:
1785:
1781:
1776:
1771:
1767:
1763:
1759:
1752:
1744:
1740:
1735:
1730:
1726:
1722:
1718:
1714:
1710:
1703:
1688:
1684:
1678:
1670:
1666:
1661:
1656:
1652:
1648:
1647:BioTechniques
1644:
1637:
1635:
1626:
1622:
1618:
1614:
1610:
1606:
1602:
1598:
1594:
1590:
1583:
1581:
1572:
1568:
1564:
1560:
1557:(3): 94–102.
1556:
1552:
1545:
1543:
1534:
1530:
1526:
1522:
1518:
1514:
1506:
1498:
1494:
1490:
1486:
1483:(3): 365–75.
1482:
1478:
1471:
1463:
1459:
1454:
1449:
1446:(1): 124–34.
1445:
1441:
1437:
1430:
1422:
1418:
1414:
1410:
1406:
1405:10.1038/79951
1402:
1399:(2): 216–20.
1398:
1394:
1387:
1379:
1375:
1371:
1367:
1364:(1): 141–58.
1363:
1359:
1352:
1344:
1340:
1336:
1332:
1328:
1324:
1321:(3): 187–95.
1320:
1316:
1309:
1307:
1298:
1294:
1290:
1284:
1280:
1276:
1272:
1265:
1257:
1253:
1249:
1245:
1241:
1237:
1233:
1229:
1225:
1221:
1217:
1210:
1202:
1198:
1194:
1190:
1186:
1182:
1178:
1174:
1170:
1166:
1162:
1155:
1141:
1137:
1130:
1123:. 2019-09-10.
1122:
1121:
1116:
1110:
1102:
1098:
1094:
1090:
1086:
1082:
1078:
1074:
1067:
1059:
1055:
1050:
1045:
1041:
1037:
1033:
1029:
1025:
1018:
1016:
1007:
1003:
1000:(2): 109–13.
999:
995:
988:
980:
976:
972:
966:
962:
958:
954:
947:
939:
935:
928:
921:
909:
901:
894:
890:
886:
882:
878:
871:
867:
862:
857:
852:
847:
843:
839:
835:
828:
826:
817:
813:
808:
803:
799:
795:
791:
784:
782:
773:
769:
764:
759:
756:(2): 209–14.
755:
751:
747:
740:
736:
727:
724:
722:
719:
718:
712:
709:
705:
701:
697:
693:
689:
685:
680:
678:
674:
670:
666:
662:
657:
656:cell membrane
647:
644:
639:
636:
632:
628:
624:
620:
616:
611:
609:
604:
600:
597:
596:coding region
593:
588:
583:
580:
576:
572:
568:
567:organogenesis
563:
559:
549:
547:
543:
539:
535:
531:
521:
519:
515:
511:
507:
503:
499:
495:
491:
487:
483:
479:
476:
472:
468:
459:
450:
448:
443:
441:
437:
433:
432:knocking down
429:
425:
421:
417:
414:Bound to the
408:
399:
397:
393:
389:
385:
381:
377:
373:
369:
365:
361:
357:
353:
349:
340:
335:
325:
323:
319:
315:
311:
310:electroporate
307:
303:
302:
297:
293:
292:
287:
283:
279:
275:
265:
263:
259:
255:
251:
247:
243:
239:
235:
231:
228:
218:
216:
206:
204:
200:
196:
192:
188:
184:
180:
176:
172:
168:
164:
160:
155:
153:
149:
145:
141:
137:
133:
129:
125:
122:
118:
113:
109:
107:
103:
99:
94:
92:
91:knocking down
88:
84:
80:
76:
72:
68:
64:
60:
57:
53:
49:
45:
41:
32:
26:
22:
3648:
3608:Bacterial
3583:Lambda phage
3554:
3247:Constituents
3188:
3176:
3127:
3123:
3113:
3088:
3084:
3078:
3053:
3049:
3042:
3007:
3003:
2993:
2968:
2964:
2957:
2930:
2926:
2916:
2883:
2879:
2872:
2839:
2835:
2829:
2774:
2768:
2751:10533/178089
2733:
2729:
2723:
2697:10.3791/1115
2688:
2684:
2674:
2637:
2633:
2623:
2599:(3): 89–93.
2596:
2592:
2582:
2560:(1): 50–60.
2557:
2553:
2547:
2522:
2518:
2512:
2475:
2471:
2428:
2424:
2418:
2386:(1): 69–77.
2383:
2379:
2325:
2321:
2311:
2276:
2272:
2262:
2227:
2223:
2213:
2178:
2174:
2164:
2129:
2125:
2115:
2074:
2070:
2064:
2032:(6): 566–9.
2029:
2026:Biology Open
2025:
2015:
1980:
1976:
1966:
1931:
1927:
1917:
1890:
1886:
1876:
1854:(2): 521–7.
1851:
1847:
1841:
1809:(5): 420–6.
1806:
1802:
1792:
1765:
1761:
1751:
1716:
1712:
1702:
1691:. Retrieved
1689:. 2017-01-20
1687:newatlas.com
1686:
1677:
1650:
1646:
1595:(1): 62–75.
1592:
1588:
1554:
1550:
1516:
1512:
1505:
1480:
1476:
1470:
1443:
1439:
1429:
1396:
1392:
1386:
1361:
1357:
1351:
1318:
1314:
1270:
1264:
1223:
1219:
1209:
1168:
1164:
1154:
1143:. Retrieved
1139:
1129:
1118:
1109:
1076:
1072:
1066:
1031:
1027:
997:
993:
987:
952:
946:
937:
933:
920:
906:{{
898:{{
841:
838:PLOS Biology
837:
800:(3): 154–6.
797:
793:
753:
749:
739:
707:
681:
673:heteroduplex
653:
640:
612:
605:
601:
584:
555:
527:
475:nucleophilic
464:
444:
440:Western blot
413:
345:
318:cell culture
308:embryos, or
304:) producing
299:
289:
271:
238:base-pairing
234:nucleic acid
224:
212:
156:
130:, including
110:
95:
62:
51:
47:
43:
39:
37:
3681:Morpholines
3603:P1-derived
3371:Antisense
3264:Nucleotides
3259:Nucleosides
3254:Nucleobases
1513:Development
1477:Development
879:, see
844:(8): e203.
700:angioplasty
690:cells with
635:phenocopied
546:frameshifts
534:Fluorescein
424:translation
384:translation
380:start codon
352:transcribed
199:viltolarsen
144:sea urchins
3655:Categories
3555:Morpholino
3468:Organellar
3376:Processual
3341:Regulatory
3285:non-coding
2640:(5): e78.
2431:(1): 1–7.
2132:(4): e41.
1693:2017-01-25
1145:2019-12-14
940:(2): 1023.
732:References
629:-mediated
615:cell death
575:phenotypes
428:transcript
348:eukaryotic
332:See also:
296:sea urchin
254:phosphates
246:morpholine
215:Gene Tools
203:casimersen
195:golodirsen
179:eteplirsen
163:pathogenic
106:trade name
102:morpholine
69:to modify
65:) used in
40:Morpholino
21:morpholine
3515:Analogues
3500:Hachimoji
3283:(coding,
3238:Types of
3124:Molecules
2819:ignored (
2809:cite book
2380:Zebrafish
1256:233248050
1240:1179-1950
1201:207935659
1185:1699-3993
669:endosomes
631:apoptosis
608:Nucleases
587:wild-type
558:embryonic
430:(called "
420:ribosomal
368:cytoplasm
322:dendrimer
286:zebrafish
230:molecules
227:synthetic
221:Structure
136:zebrafish
46:and as a
3638:Category
3573:Phagemid
3424:Ribozyme
3156:19325525
3105:18564870
3070:17583927
3034:17344287
2985:17097177
2949:12020498
2908:42463260
2900:16285002
2801:15375324
2760:16837210
2715:19274045
2666:17530925
2615:11477681
2574:17226957
2504:29049395
2464:Moens CB
2445:18196596
2410:19374550
2354:19458392
2303:21925381
2254:23172291
2205:15383681
2156:15731334
2099:15386015
2056:23213449
2007:17220889
1958:15585662
1909:18022366
1868:17493584
1833:21092974
1825:16550551
1784:15333583
1743:27999041
1669:19238792
1625:28590429
1617:16394126
1571:11477682
1533:11262245
1497:11152635
1462:10885751
1421:21451111
1413:11017081
1378:10807004
1343:19372403
1297:28364229
1248:33861387
1193:31720560
1101:21338534
1093:16919955
1058:17485503
1006:15844617
979:28338321
893:35486652
870:17676975
816:11477696
772:11884031
715:See also
650:Delivery
643:morphant
542:ribozyme
498:splicing
467:pre-mRNA
372:ribosome
306:morphant
268:Function
167:bacteria
152:pre-mRNA
148:splicing
59:molecule
56:oligomer
25:morphine
3578:Plasmid
3177:Genesis
3147:6253989
3025:1900280
2864:8161222
2856:8915501
2730:Methods
2706:2762901
2657:1877875
2593:Genesis
2539:9012862
2495:5648102
2425:Genesis
2401:2776066
2345:2740424
2294:3781938
2245:3553965
2196:1370650
2107:4425795
2079:Bibcode
2047:3509442
1998:3982799
1734:5890718
1597:Bibcode
1551:Genesis
1335:9212909
1049:1913242
861:1925136
794:Genesis
708:in vivo
694:or the
684:tissues
617:in the
579:Control
478:adenine
388:protein
366:to the
364:nucleus
360:spliced
356:introns
291:Xenopus
258:anionic
209:History
171:viruses
124:encoded
3593:Fosmid
3588:Cosmid
3538:Hexose
3460:acids
3412:Others
3196:
3154:
3144:
3103:
3068:
3032:
3022:
2983:
2947:
2906:
2898:
2862:
2854:
2799:
2789:
2758:
2713:
2703:
2691:(25).
2664:
2654:
2613:
2572:
2537:
2502:
2492:
2443:
2408:
2398:
2352:
2342:
2301:
2291:
2252:
2242:
2203:
2193:
2154:
2147:549580
2144:
2105:
2097:
2071:Nature
2054:
2044:
2005:
1995:
1956:
1949:535676
1946:
1907:
1866:
1831:
1823:
1782:
1741:
1731:
1667:
1623:
1615:
1569:
1531:
1495:
1460:
1419:
1411:
1376:
1341:
1333:
1295:
1285:
1254:
1246:
1238:
1199:
1191:
1183:
1099:
1091:
1056:
1046:
1004:
977:
967:
891:
887:,
868:
858:
814:
770:
688:muscle
623:somite
592:5'-UTR
514:RT-PCR
282:embryo
201:, and
121:moiety
73:. Its
3618:Human
3396:Y RNA
2904:S2CID
2860:S2CID
2103:S2CID
1829:S2CID
1621:S2CID
1417:S2CID
1339:S2CID
1252:S2CID
1220:Drugs
1197:S2CID
1097:S2CID
975:S2CID
930:(PDF)
904:with
562:cells
530:miRNA
502:exons
471:snRNP
386:of a
314:chick
278:siRNA
181:from
140:frogs
63:oligo
3194:ISBN
3152:PMID
3101:PMID
3066:PMID
3030:PMID
2981:PMID
2945:PMID
2896:PMID
2852:PMID
2821:help
2797:PMID
2787:ISBN
2756:PMID
2711:PMID
2662:PMID
2611:PMID
2570:PMID
2535:PMID
2500:PMID
2441:PMID
2406:PMID
2350:PMID
2299:PMID
2250:PMID
2201:PMID
2152:PMID
2095:PMID
2052:PMID
2003:PMID
1954:PMID
1905:PMID
1887:Cell
1864:PMID
1821:PMID
1780:PMID
1739:PMID
1665:PMID
1613:PMID
1593:1058
1567:PMID
1529:PMID
1493:PMID
1458:PMID
1409:PMID
1374:PMID
1362:1489
1331:PMID
1293:PMID
1283:ISBN
1244:PMID
1236:ISSN
1189:PMID
1181:ISSN
1089:PMID
1054:PMID
1002:PMID
965:ISBN
889:PMID
866:PMID
812:PMID
768:PMID
621:and
569:and
392:gene
358:are
173:and
142:and
132:mice
117:exon
3142:PMC
3132:doi
3093:doi
3058:doi
3020:PMC
3012:doi
2973:doi
2969:116
2935:doi
2888:doi
2844:doi
2779:doi
2746:hdl
2738:doi
2701:PMC
2693:doi
2652:PMC
2642:doi
2601:doi
2562:doi
2527:doi
2490:PMC
2480:doi
2433:doi
2396:PMC
2388:doi
2340:PMC
2330:doi
2326:284
2289:PMC
2281:doi
2240:PMC
2232:doi
2191:PMC
2183:doi
2175:RNA
2142:PMC
2134:doi
2087:doi
2075:431
2042:PMC
2034:doi
1993:PMC
1985:doi
1944:PMC
1936:doi
1895:doi
1891:131
1856:doi
1852:358
1811:doi
1770:doi
1729:PMC
1721:doi
1655:doi
1605:doi
1559:doi
1521:doi
1517:128
1485:doi
1481:128
1448:doi
1444:222
1401:doi
1366:doi
1323:doi
1275:doi
1228:doi
1173:doi
1140:FDA
1081:doi
1044:PMC
1036:doi
957:doi
881:doi
856:PMC
846:doi
802:doi
758:doi
754:243
627:p53
510:U12
506:U11
396:DNA
346:In
294:),
242:DNA
169:or
150:of
89:by
52:PMO
23:or
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