51:
990:
physical, chemical, and macromolecular properties. Viroids were shown to consist of short stretches (a few hundred nucleotides) of single-stranded RNA and, unlike viruses, did not have a protein coat. Viroids are extremely small, from 246 to 467 nucleotides, smaller than other infectious plant pathogens; they thus consist of fewer than 10,000 atoms. In comparison, the genomes of the smallest known viruses capable of causing an infection by themselves are around 2,000 nucleotides long.
861:. If so, viroids have assumed significance beyond plant virology for evolutionary theory, because their properties make them more plausible candidates than other RNAs to perform crucial steps in the evolution of life from inanimate matter (abiogenesis). Diener's hypothesis was mostly forgotten until 2014, when it was resurrected in a review article by Flores et al., in which the authors summarized Diener's evidence supporting his hypothesis as:
572:
4817:
994:
continuous loop, in which the 3' and 5' ends present in linear RNA molecules have been joined. Sanger et al. also provided evidence for the true circularity of viroids by finding that the RNA could not be phosphorylated at the 5' terminus. In other tests, they failed to find even one free 3' end, which ruled out the possibility of the molecule having two 3' ends. Viroids thus are true circular RNAs.
4841:
4829:
38:
993:
In 1976, Sanger et al. presented evidence that potato spindle tuber viroid is a "single-stranded, covalently closed, circular RNA molecule, existing as a highly base-paired rod-like structure"—believed to be the first such molecule described. Circular RNA, unlike linear RNA, forms a covalently closed
997:
The single-strandedness and circularity of viroids was confirmed by electron microscopy, The complete nucleotide sequence of potato spindle tuber viroid was determined in 1978. PSTVd was the first pathogen of a eukaryotic organism for which the complete molecular structure has been established. Over
985:
could not be found consistently associated with symptom-bearing plants, however, and therefore, it was assumed the disease was caused by a virus. Despite numerous attempts over the years to isolate and purify the assumed virus, using increasingly sophisticated methods, these were unsuccessful when
900:
The presence, in extant cells, of RNAs with molecular properties predicted for RNAs of the RNA world constitutes another powerful argument supporting the RNA world hypothesis. However, the origins of viroids themselves from this RNA world has been cast into doubt by several factors, including the
989:
In 1971, Theodor O. Diener showed that the agent was not a virus, but a totally unexpected novel type of pathogen, 1/80th the size of typical viruses, for which he proposed the term "viroid". Parallel to agriculture-directed studies, more basic scientific research elucidated many of viroids'
244:
973:
In the 1920s, symptoms of a previously unknown potato disease were noticed in New York and New Jersey fields. Because tubers on affected plants become elongated and misshapen, they named it the potato spindle tuber disease.
587:, by cross contamination following mechanical damage to plants as a result of horticultural or agricultural practices, or from plant to plant by leaf contact. Upon infection, viroids replicate in the nucleus (
686:. The viroid siRNAs contain sequences capable of complementary base pairing with the plant's own messenger RNAs, and induction of degradation or inhibition of translation causes the classic viroid symptoms.
4102:
694:"Viroid-like elements" refer to pieces of covalently closed circular (ccc) RNA molecules that do not share the viroid's lifecycle. The category encompasses satellite RNAs (including small plant satRNAs "
772:
in various combinations in both sense and antisense orientation – the detection of circular forms in both sense orientations suggest that "ambiviruses" use rolling circle replication for propagation.
925:). However, recent studies suggest that the diversity of viroids and others viroid-like elements is broader than previously thought and that it would not be limited to plants, encompassing even the
176:, plant pathologist at the U.S Department of Agriculture's Research Center in Beltsville, Maryland, in 1971. This viroid is now called potato spindle tuber viroid, abbreviated PSTVd. The
443:
435:
845:
in any other known life form, the researchers suggest that the obelisks are distinct from viruses, viroids and viroid-like entities, and thus form an entirely new class of organisms.
130:
study suggests that the host diversity of viroids and other viroid-like elements is broader than previously thought and that it would not be limited to plants, encompassing even the
2801:
4095:
282:
3272:
1293:
Lee BD, Neri U, Roux S, Wolf YI, Camargo AP, Krupovic M, et al. (RNA Virus
Discovery Consortium; Simmonds P, Kyrpides N, Gophna U, Dolja VV, Koonin EV) (2 Feb 2023).
459:
322:
314:
298:
4088:
427:
378:
977:
The symptoms appeared on plants onto which pieces from affected plants had been budded—indicating that the disease was caused by a transmissible pathogenic agent. A
451:
1156:
556:
500:
492:
484:
411:
338:
768:
has been established since the 2023 ICTV Virus
Taxonomy Release because of the unique features of encoding RNA-directed RNA polymerases but also having divergent
290:
1155:
Di Serio F, Owens RA, Li SF, Matoušek J, Pallás V, Randles JW, Sano T, Verhoeven JT, Vidalakis G, Flores R (November 2020). Zerbini FM, Sabanadzovic S (eds.).
3132:
Gross HJ, Domdey H, Lossow C, Jank P, Raba M, Alberty H, Sänger HL (May 1978). "Nucleotide sequence and secondary structure of potato spindle tuber viroid".
2595:"Microhomology-mediated End Joining and Homologous Recombination share the initial end resection step to repair DNA double-strand breaks in mammalian cells"
419:
394:
330:
3265:
784:
genome of the host. The only types found are closely related to the original "carnation small viroid-like RNA" (CarSV). These elements may act as a
780:"Retroviroids", more formally "retroviroid-like elements", are viroid-like circular RNA sequences that are also found with homologous copies in the
2791:
764:
154:
3097:
Sogo JM, Koller T, Diener TO (September 1973). "Potato spindle tuber viroid. X. Visualization and size determination by electron microscopy".
811:: there is no extracellular form of these elements; instead, they are spread only through pollen or egg-cells. They appear to co-occur with a
4787:
1729:"Retrozymes are a unique family of non-autonomous retrotransposons with hammerhead ribozymes that propagate in plants through circular RNAs"
3258:
2656:
1010:
science writer Carl Zimmer published a popularized piece that mistakenly credited Flores et al. with the hypothesis' original conception.
4658:
212:
Diener initially hypothesized in 1989 that viroids may represent "living relics" from the widely assumed, ancient, and non-cellular
2863:
674:
develops all the corresponding viroid-like symptoms. This indicates that when viroids replicate via a double stranded intermediate
2899:"Identification of viruses and viroids by next-generation sequencing and homology-dependent and homology-independent algorithms"
4225:
2973:
2720:
2257:
1361:
2327:
Sutela S, Forgia M, Vainio EJ, Chiapello M, Daghino S, Vallino M, Martino E, Girlanda M, Perotto S, Turina M (1 July 2020).
4591:
2991:"Viroids are single-stranded covalently closed circular RNA molecules existing as highly base-paired rod-like structures"
954:
853:
Diener's 1989 hypothesis had proposed that the unique properties of viroids make them more plausible macromolecules than
833:– the computer-aided search for RNA sequences and their analysis – biologists reported in January 2024 the discovery of "
3210:
1160:
1995:
231:
is a subviral agent similar in structure to a viroid, as it is a hybrid particle enclosed by surface proteins from the
182:(CEVd) was discovered soon thereafter, and together understanding of PSTVd and CEVd shaped the concept of the viroid.
2554:"The DNA form of a retroviroid-like element is involved in recombination events with itself and with the plant genome"
575:
The reproduction mechanism of a typical viroid. Leaf contact transmits the viroid. The viroid enters the cell via its
4648:
2329:"The virome from a collection of endomycorrhizal fungi reveals new viral taxa with unprecedented genome organization"
618:, viroids are entirely passive, relying entirely on the host. This is useful in the study of RNA kinetics in plants.
2097:
Flores R, Navarro B, Kovalskaya N, Hammond RW, Di Serio F (October 2017). "Engineering resistance against viroids".
4653:
248:
4792:
2230:
Pallas V, Martinez G, Gomez G (2012). "The
Interaction Between Plant Viroid-Induced Symptoms and RNA Silencing".
2134:"Analysis of the virulence modulating region of potato spindle tuber viroid (PSTVd) by site-directed mutagenesis"
1444:
Flores R, Hernández C, Martínez de Alba AE, Daròs JA, Di Serio F (2005). "Viroids and viroid-host interactions".
4576:
3926:
3521:
4564:
4062:
683:
220:, it has been proposed that viroids and other viroid-like elements may derive from this newly found class of
209:
properties that allow self-cleavage and ligation of unit-size genomes from larger replication intermediates.
2884:
Diener, T O. "Circular RNAs: relics of precellular evolution?."Proc.Natl.Acad.Sci.USA, 1989;86(23):9370-9374
3710:
3391:
3183:
Hammond RW, Owens RA (2006). "Viroids: New and
Continuing Risks for Horticultural and Agricultural Crops".
370:
3396:
728:
are infectious circular RNA molecules that depend on a carrier virus to reproduce, being carried in their
3630:
3386:
748:
are circular, circa 5 kb in length. One of at least two open reading frames encodes a viral RNA-directed
671:
583:
Viroids are only known to infect plants, and infectious viroids can be transmitted to new plant hosts by
467:
274:
169:
146:
2284:
Balázs E, Hegedűs K, Divéki Z (15 April 2022). "The unique carnation stunt-associated pararetroviroid".
4807:
950:
541:
1380:
Diener TO (August 1971). "Potato spindle tuber "virus". IV. A replicating, low molecular weight RNA".
137:
The first discoveries of viroids in the 1970s triggered the historically third major extension of the
4080:
3931:
946:
998:
thirty plant diseases have since been identified as viroid-, not virus-caused, as had been assumed.
4866:
4635:
4581:
3778:
3679:
3596:
3239:
958:
857:, or other RNAs considered in the past as possible "living relics" of a hypothetical, pre-cellular
526:
50:
1184:"Next-Generation Sequencing and CRISPR/Cas13 Editing in Viroid Research and Molecular Diagnostics"
4670:
3805:
3581:
2173:"On the role of RNA silencing in the pathogenicity and evolution of viroids and viral satellites"
962:
662:
corresponding to sequences from viroid genomes have been isolated from infected plants. Finally,
153:(of the "submicroscopic" viruses). The unique properties of viroids have been recognized by the
142:
4714:
4033:
3349:
158:
123:
2827:
2708:
1017:", a new class of viroid-like elements, and "oblins", their related group of proteins, in the
909:
likely representing their ancestors) and their complete absence from organisms outside of the
837:", a new class of viroid-like elements, and "oblins", their related group of proteins, in the
3793:
3766:
3381:
2378:"Viroid-like RNA-dependent RNA polymerase-encoding ambiviruses are abundant in complex fungi"
800:
789:
3369:
1001:
Four additional viroids or viroid-like RNA particles were discovered between 2009 and 2015.
793:
4761:
4680:
3737:
3732:
3608:
3141:
3002:
2606:
2462:
2184:
1862:
1619:
1457:
1306:
1124:
1107:
1081:
1066:
45:
2510:"The DNA form of a retroviroid-like element characterized in cultivated carnation species"
1791:"Circular RNAs with hammerhead ribozymes encoded in eukaryotic genomes: The enemy at home"
882:
Existence of structural periodicity, which permits modular assembly into enlarged genomes.
8:
4255:
3973:
3815:
3771:
3727:
3465:
1071:
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834:
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173:
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127:
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2915:
2898:
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2764:
2737:
2736:
Breit TM, de Leeuw WC, van Olst M, Ensink WA, van
Leeuwen S, Dekker RJ (16 March 2023).
2675:
2610:
2466:
2404:
2377:
2188:
2048:"Viroids: from genotype to phenotype just relying on RNA sequence and structural motifs"
1866:
1623:
4549:
4471:
4023:
3165:
3074:
3049:
2629:
2594:
2353:
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1234:
1210:
1183:
842:
3250:
3110:
3025:
2990:
2207:
2172:
2011:
De Bokx JA, Piron PG (1981). "Transmission of potato spindle tuber viroid by aphids".
1642:
1607:
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4527:
4443:
4345:
4115:
4018:
3722:
3157:
3114:
3079:
3030:
2969:
2920:
2855:
2792:"'Obelisks': Entirely New Class of Life Has Been Found in The Human Digestive System"
2769:
2716:
2689:
2634:
2575:
2531:
2490:
2485:
2450:
2409:
2358:
2313:
2301:
2263:
2253:
2212:
2153:
2149:
2114:
2079:
1991:
1949:
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1880:
1851:"Viroids and Viroid-like Circular RNAs: Do They Descend from Primordial Replicators?"
1828:
1810:
1768:
1750:
1706:
1688:
1647:
1588:
1528:
Flores R, Di Serio F, Hernández C (February 1997). "Viroids: The
Noncoding Genomes".
1514:
1502:
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2389:
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2235:
2202:
2192:
2145:
2106:
2069:
2059:
2020:
1939:
1929:
1888:
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1818:
1802:
1758:
1740:
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1678:
1637:
1627:
1578:
1568:
1537:
1492:
1453:
1389:
1324:
1316:
1256:
1246:
1205:
1195:
1119:
1018:
838:
740:"Ambiviruses" are mobile genetic elements that were recently (2020s) discovered in
355:
306:
178:
2171:
Wang MB, Bian XY, Wu LM, Liu LX, Smith NA, Isenegger D, et al. (March 2004).
1806:
1418:
879:
Their circular structure, which assures complete replication without genomic tags.
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4387:
4011:
3705:
3684:
3613:
3416:
3065:
2239:
2110:
1051:
906:
251:
of the PSTVd viroid. The highlighted nucleotides are found in most other viroids.
221:
162:
65:
2570:
2553:
4845:
4821:
4436:
4401:
4394:
4373:
4352:
4329:
4209:
4001:
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3618:
3601:
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3336:
3215:
2995:
Proceedings of the
National Academy of Sciences of the United States of America
2947:
2599:
Proceedings of the
National Academy of Sciences of the United States of America
2526:
2509:
2455:
Proceedings of the
National Academy of Sciences of the United States of America
2394:
2177:
Proceedings of the
National Academy of Sciences of the United States of America
1612:
Proceedings of the National Academy of Sciences of the United States of America
1320:
1006:
812:
749:
608:
201:" synthesis of new RNA using the viroid's RNA as a template. Viroids are often
198:
2841:
1745:
1356:. Burlington, Massachusetts, US: Elsevier Academic Press. pp. 1221–1259.
1354:
Virus Taxonomy. Ninth Report of the International Committee for Virus Taxonomy
1311:
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4737:
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4603:
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4429:
4415:
4366:
4359:
4278:
4151:
4041:
4006:
3849:
3839:
3810:
3443:
3436:
2064:
1884:
1814:
1754:
1692:
1295:"Mining metatranscriptomes reveals a vast world of viroid-like circular RNAs"
1133:
1036:
808:
799:
These elements are dubbed retroviroids as the homologous DNA is generated by
758:
725:
655:
635:
604:
599:) of plant cells in three steps through an RNA-based mechanism. They require
576:
512:
260:
194:
97:
91:
3015:
2619:
2475:
2197:
1985:
1632:
1479:
Tsagris EM, Martínez de Alba AE, Gozmanova M, Kalantidis K (November 2008).
4692:
4625:
4489:
4457:
4422:
4408:
4380:
4306:
4285:
4028:
3871:
3831:
3798:
3570:
3556:
3454:
3083:
2924:
2859:
2796:
2773:
2693:
2638:
2579:
2535:
2413:
2362:
2305:
2267:
2216:
2118:
2083:
1953:
1902:
1832:
1772:
1710:
1592:
1541:
1506:
1465:
1338:
1270:
1219:
1141:
1031:
651:
592:
535:
111:
3118:
3034:
2989:
Sanger HL, Klotz G, Riesner D, Gross HJ, Kleinschmidt AK (November 1976).
2494:
2157:
1934:
1651:
1478:
1401:
4746:
4554:
4299:
4052:
3918:
3866:
3861:
3374:
3206:
3161:
2754:
2593:
Truong LN, Li Y, Shi LZ, Hwang PY, He J, Wang H, et al. (May 2013).
2344:
1875:
1683:
1573:
1251:
926:
708:
703:
667:
520:
405:
364:
349:
268:
131:
119:
2684:
2655:
Flores R, Gago-Zachert S, Serra P, Sanjuán R, Elena SF (June 18, 2014).
986:
applied to extracts from potato spindle tuber disease-afflicted plants.
185:
Although viroids are composed of nucleic acid, they do not code for any
141:—to include smaller lifelike entities —after the discoveries in 1675 by
4685:
4632:
4219:
4132:
3960:
3955:
3908:
3901:
3896:
3881:
3876:
3717:
3652:
3401:
3343:
2248:
2024:
914:
804:
663:
478:
216:, and others have followed this conjecture. Following the discovery of
4110:
1200:
579:. RNA polymerase II catalyzes rolling-circle synthesis of new viroids.
4774:
4292:
4271:
4173:
3965:
3950:
3937:
3689:
3477:
3308:
3153:
902:
893:
858:
753:
699:
695:
643:
571:
550:
217:
213:
206:
202:
138:
118:, they have no protein coating. All known viroids are inhabitants of
4708:
2508:
Hegedűs K, Palkovics L, Tóth EK, Dallmann G, Balázs E (March 2001).
2133:
1443:
953:
has allowed for rapid and inexpensive detection of known viroids in
172:, was discovered, initially molecularly characterized, and named by
4816:
4779:
4731:
4665:
4610:
4197:
4181:
4146:
3996:
3942:
3844:
3788:
3783:
3674:
3635:
3591:
3493:
3298:
2738:"Genome Sequence of a New Carnation Small Viroid-Like RNA, CarSV-1"
1086:
982:
918:
886:
873:
769:
713:
22:
1352:
King AM, Adams MJ, Carstens EB, Lefkovitz EJ, et al. (2012).
933:
spacers suggest that some of them might replicate in prokaryotes.
4766:
4697:
4615:
4596:
4156:
4141:
3978:
3856:
3640:
3625:
3328:
3303:
2832:
2234:. Methods in Molecular Biology. Vol. 894. pp. 323–343.
1056:
922:
869:
732:. Like Avsunviroidae, however, they are capable of self-clevage.
631:
627:
186:
1986:
Brian W. J. Mahy, Marc H. V. Van Regenmortel, ed. (2009-10-29).
4643:
4586:
4571:
4521:
4069:
4047:
3744:
3667:
3662:
3318:
3313:
2427:
978:
930:
854:
745:
729:
639:
2654:
2096:
122:(flowering plants), and most cause diseases, whose respective
4620:
4247:
4191:
4186:
3551:
3543:
3361:
3323:
1076:
1061:
942:
910:
741:
712:) and "retroviroids". Most of them also carry some type of a
682:
enzyme and cleaved into siRNAs that are then loaded onto the
679:
659:
647:
584:
115:
4840:
2968:. Burlington, MA: Jones and Bartlett Learning. p. 482.
2507:
876:
content, which increases stability and replication fidelity.
243:
37:
4559:
4163:
4111:
4057:
3749:
3290:
2735:
2326:
2046:
Flores R, Serra P, Minoia S, Di Serio F, Navarro B (2012).
1232:
603:, a host cell enzyme normally associated with synthesis of
193:, a host cell enzyme normally associated with synthesis of
145:(of the "subvisible" microorganisms) and in 1892–1898 by
2451:"Identification of a retroviroid-like element from plants"
1351:
2709:"Chapter 5: Agents Resembling or Altering Virus Diseases"
2045:
781:
675:
638:
is involved in the process. First, changes to the viroid
168:
The first recognized viroid, the pathogenic agent of the
885:
Their lack of protein-coding ability, consistent with a
865:
Viroids' small size, imposed by error-prone replication.
634:
products within their sequences. Evidence suggests that
626:
There has long been uncertainty over how viroids induce
3280:
3244:
2988:
2938:
Owens RA, Verhoeven JT (2009). "Potato Spindle Tuber".
1726:
1527:
1154:
1013:
In January 2024, biologists reported the discovery of "
21:
For a variant of it which is dependent on viruses, see
4805:
3247:, a database of viroids and viroid-like circular RNAs
3131:
3050:"Current view and perspectives in viroid replication"
2551:
1419:"ARS Research Timeline – Tracking the Elusive Viroid"
1235:"Impact of Nucleic Acid Sequencing on Viroid Biology"
1106:
Navarro B, Flores R, Di Serio F (29 September 2021).
1105:
841:. Given that the RNA sequences recovered do not have
611:" synthesis of new RNA using the viroid as template.
423:; (CVd-III); accession AF184147, genome length 294nt
2283:
2229:
1233:Adkar-Purushothama CR, Perreault JP (August 2020).
2826:Zheludev, Ivan N., et al. (21 January 2024).
447:; (GVYSd-2); accession J04348, genome length 363nt
439:; (GVYSd-1); accession X06904, genome length 367nt
286:; (TCDVd); accession AF162131, genome length 360nt
2825:
1788:
1608:"Circular RNAs: relics of precellular evolution?"
504:; (CbVd-3); accession X95364, genome length 361nt
496:; (CbVd-2); accession X95365, genome length 301nt
398:; (CVd-IV); accession X14638, genome length 284nt
334:; (IrVd-1); accession X95734, genome length 370nt
4853:
3096:
2592:
1915:
1727:Cervera A, Urbina D, de la Peña M (2016-06-23).
1664:
1554:
807:. They are neither true viroids nor viroid-like
463:; (PBCVd); accession D12823, genome length 315nt
455:; (CBLVd); accession M74065, genome length 318nt
431:; (ADFVd); accession X99487, genome length 306nt
382:; (CTiVd); accession M20731, genome length 254nt
326:; (TASVd); accession K00818, genome length 360nt
302:; (TPMVd); accession K00817, genome length 360nt
2892:
2890:
2821:
2819:
2279:
2277:
1292:
566:
471:; (AGVd); accession X17101, genome length 369nt
390:; (HLVd); accession X07397, genome length 256nt
342:; (CLVd); accession X15663, genome length 370nt
318:; (CSVd); accession V01107, genome length 356nt
294:; (MPVd); accession L78454, genome length 360nt
2959:
2957:
2937:
2896:
2785:
2783:
2715:(Fifth ed.). London, UK: Academic Press.
2650:
2648:
1988:Desk Encyclopedia of Plant and Fungal Virology
1439:
1437:
1435:
1413:
1411:
155:International Committee on Taxonomy of Viruses
4096:
3266:
2789:
2170:
1375:
1373:
719:
126:importance to humans varies widely. A recent
16:Pathogenic small single-stranded circular RNA
3182:
3176:
3041:
2887:
2828:"Viroid-like colonists of human microbiomes"
2816:
2501:
2274:
2090:
2039:
2010:
1548:
4659:Reverse transcriptase-related cellular gene
3090:
2982:
2963:
2954:
2780:
2645:
2448:
2375:
2125:
1671:International Journal of Molecular Sciences
1599:
1561:International Journal of Molecular Sciences
1432:
1408:
1239:International Journal of Molecular Sciences
1226:
4103:
4089:
3273:
3259:
2547:
2545:
1370:
189:. The viroid's replication mechanism uses
36:
3073:
3024:
3014:
2914:
2849:
2763:
2753:
2683:
2628:
2618:
2569:
2525:
2484:
2474:
2403:
2393:
2352:
2247:
2206:
2196:
2073:
2063:
1943:
1933:
1918:"Hepatitis delta virus: a peculiar virus"
1892:
1874:
1848:
1822:
1762:
1744:
1700:
1682:
1641:
1631:
1582:
1572:
1496:
1328:
1310:
1260:
1250:
1209:
1199:
1123:
2657:"Viroids: survivors from the RNA world?"
2552:Hegedűs K, Dallmann G, Balázs E (2004).
913:(especially their complete absence from
752:, that firmly places "ambiviruses" into
570:
242:
4640:Retroelements not elsewhere classified
3211:"A Tiny Emissary From the Ancient Past"
2542:
2131:
848:
689:
650:produced would have less complementary
4854:
3205:
3199:
2897:Wu Q, Ding SW, Zhang Y, Zhu S (2015).
2586:
2013:Netherlands Journal of Plant Pathology
1789:de la Peña M, Cervera A (2017-08-03).
1605:
1458:10.1146/annurev.phyto.43.040204.140243
1379:
1181:
1125:10.1146/annurev-virology-091919-092331
1108:"Advances in Viroid-Host Interactions"
1099:
114:that are infectious pathogens. Unlike
4713:
4712:
4084:
3254:
2442:
1981:
1979:
1977:
1975:
1973:
1971:
1969:
1967:
1965:
1963:
1844:
1842:
1784:
1782:
1722:
1720:
1665:Moelling K, Broecker F (2021-03-28).
1555:Moelling K, Broecker F (March 2021).
1288:
1286:
1284:
1282:
1280:
929:. Matches between viroid cccRNAs and
3047:
2866:from the original on 29 January 2024
2804:from the original on 29 January 2024
2706:
4592:Integrative and conjugative element
2916:10.1146/annurev-phyto-080614-120030
2790:Koumoundouros T (29 January 2024).
2742:Microbiology Resource Announcements
2676:10.1146/annurev-micro-091313-103416
2428:"ICTV Virus Taxonomy: 2023 Release"
1916:Alves C, Branco C, Cunha C (2013).
614:Unlike plant viruses which produce
607:from DNA, which instead catalyzes "
197:from DNA, which instead catalyzes "
13:
2376:Chong LC, Lauber C (12 May 2023).
1990:. Academic Press. pp. 71–81.
1960:
1839:
1779:
1717:
1277:
941:The development of tests based on
896:—the fingerprint of the RNA world.
646:. This reflects the fact that any
14:
4878:
4649:Diversity-generating retroelement
3233:
444:Grapevine yellow speckle 2 viroid
436:Grapevine yellow speckle 1 viroid
4839:
4827:
4815:
4654:Telomerase reverse transcriptase
4226:Microbes with highly unusual DNA
1849:Lee BD, Koonin EV (2022-01-12).
1667:"Viroids and the Origin of Life"
1557:"Viroids and the Origin of Life"
1498:10.1111/j.1462-5822.2008.01231.x
892:Replication mediated in some by
666:expression of the noninfectious
621:
49:
3193:10.1094/APSnetFeature-2006-1106
3125:
2931:
2903:Annual Review of Phytopathology
2878:
2729:
2700:
2514:The Journal of General Virology
2420:
2369:
2320:
2223:
2164:
2004:
1909:
1658:
1521:
1472:
1446:Annual Review of Phytopathology
775:
630:in plants without encoding any
4577:Defective interfering particle
3927:Last universal common ancestor
3522:Defective interfering particle
2298:10.1016/j.virusres.2022.198709
2232:Antiviral Resistance in Plants
1345:
1175:
1148:
735:
1:
4565:Clonally transmissible cancer
4063:Clonally transmissible cancer
3499:Satellite-like nucleic acids
3111:10.1016/s0042-6822(73)81009-8
2664:Annual Review of Microbiology
1807:10.1080/15476286.2017.1321730
1093:
684:RNA-induced silencing complex
283:Tomato chlorotic dwarf viroid
3066:10.1016/j.coviro.2020.12.004
2966:Fundamentals of Microbiology
2240:10.1007/978-1-61779-882-5_22
2150:10.1016/0042-6822(92)90468-5
2111:10.1016/j.coviro.2017.07.003
1394:10.1016/0042-6822(71)90342-4
794:double-stranded break repair
792:may occur and are linked to
567:Transmission and replication
371:Coconut cadang-cadang viroid
170:potato spindle tuber disease
7:
4118:, and comparable structures
3054:Current Opinion in Virology
2571:10.1016/j.virol.2004.04.035
2449:Daròs JA, Flores R (1995).
2099:Current Opinion in Virology
1606:Diener TO (December 1989).
1024:
818:
672:potato spindle tuber viroid
642:can dramatically alter its
468:Australian grapevine viroid
278:; 356–361 nucleotides(nt)
275:Potato spindle tuber viroid
263:: relies on host Rnase III
238:
147:Dmitri Iosifovich Ivanovsky
110:are small single-stranded,
10:
4883:
3619:Class II or DNA transposon
3614:Class I or retrotransposon
2948:10.1094/PHI-I-2009-0804-01
2527:10.1099/0022-1317-82-3-687
2395:10.3389/fmicb.2023.1144003
1321:10.1016/j.cell.2022.12.039
968:
951:nucleic acid hybridization
936:
822:
720:Viroid-like satellite RNAs
542:Peach latent mosaic viroid
460:Pear blister canker viroid
323:Tomato apical stunt viroid
315:Chrysanthemum stunt viroid
299:Tomato planta macho viroid
20:
4721:
4540:
4514:
4481:
4337:
4323:
4316:
4263:
4246:
4239:
4207:
4172:
4130:
4123:
3989:
3932:Earliest known life forms
3917:
3830:
3806:Repeated sequences in DNA
3698:
3580:
3569:
3542:
3514:
3464:
3453:
3426:
3415:
3360:
3289:
2842:10.1101/2024.01.20.576352
2382:Frontiers in Microbiology
2132:Hammond RW (April 1992).
2052:Frontiers in Microbiology
1746:10.1186/s13059-016-1002-4
1312:10.1101/2022.07.19.500677
1182:Hadidi A (January 2019).
1112:Annual Review of Virology
959:phytosanitary inspections
678:, they are targeted by a
428:Apple dimple fruit viroid
88:
83:
44:
35:
30:
4582:Endogenous viral element
3779:Endogenous viral element
3597:Horizontal gene transfer
2065:10.3389/fmicb.2012.00217
527:Avocado sunblotch viroid
515:: autocatalytic clevage
379:Coconut tinangaja viroid
3476:dsDNA satellite virus (
3185:APSnet Feature Articles
3016:10.1073/pnas.73.11.3852
2964:Pommerville JC (2014).
2940:Plant Health Instructor
2707:Hull R (October 2013).
2620:10.1073/pnas.1213431110
2476:10.1073/pnas.92.15.6856
2198:10.1073/pnas.0400104101
1633:10.1073/pnas.86.23.9370
955:biosecurity inspections
702:", and the much larger
452:Citrus bent leaf viroid
307:Citrus exocortis viroid
179:Citrus exocortis viroid
143:Antonie van Leeuwenhoek
4034:Helper dependent virus
3350:Biological dark matter
3209:(September 25, 2014).
1542:10.1006/smvy.1997.0107
580:
557:Eggplant latent viroid
501:Coleus blumei 3 viroid
493:Coleus blumei 2 viroid
485:Coleus blumei viroid 1
412:Apple scar skin viroid
339:Columnea latent viroid
252:
3794:Endogenous retrovirus
3767:Origin of replication
3483:ssDNA satellite virus
3473:ssRNA satellite virus
3048:Wang Y (April 2021).
1485:Cellular Microbiology
801:reverse transcriptase
788:substrate upon which
574:
291:Mexican papita viroid
246:
157:, in creating a new
4681:Transposable element
4671:Spiegelman's Monster
3738:Secondary chromosome
3733:Extrachromosomal DNA
3609:Transposable element
2755:10.1128/mra.01219-22
1922:Advances in Virology
1876:10.3390/life12010103
1684:10.3390/ijms22073476
1574:10.3390/ijms22073476
1530:Seminars in Virology
1252:10.3390/ijms21155532
1082:Virus classification
1067:RNA world hypothesis
849:RNA world hypothesis
762:; a separate phylum
690:Viroid-like elements
46:Virus classification
3974:Model lipid bilayer
3816:Interspersed repeat
3146:1978Natur.273..203G
3007:1976PNAS...73.3852S
2611:2013PNAS..110.7720T
2467:1995PNAS...92.6856D
2189:2004PNAS..101.3275W
1935:10.1155/2013/560105
1867:2022Life...12..103L
1624:1989PNAS...86.9370D
1163:on December 2, 2020
1072:Satellite (biology)
1047:Obelisk (life form)
831:metatranscriptomics
803:that is encoded by
249:secondary structure
227:The human pathogen
174:Theodor Otto Diener
151:Martinus Beijerinck
128:metatranscriptomics
4550:Bio-like structure
4472:Tolecusatellitidae
3284:organic structures
2345:10.1093/ve/veaa076
2025:10.1007/bf01976653
581:
253:
4803:
4802:
4715:Taxon identifiers
4706:
4705:
4536:
4535:
4510:
4509:
4506:
4505:
4444:Portogloboviridae
4346:Alphasatellitidae
4240:Non-cellular life
4235:
4234:
4116:non-cellular life
4078:
4077:
4019:Non-cellular life
3826:
3825:
3565:
3564:
3538:
3537:
3492:ssRNA satellite (
2975:978-1-284-03968-9
2722:978-0-12-384872-7
2520:(Pt 3): 687–691.
2461:(15): 6856–6860.
2259:978-1-61779-881-8
1363:978-0-12-384685-3
1201:10.3390/v11020120
1042:Non-cellular life
825:Obelisk (biology)
616:movement proteins
601:RNA polymerase II
420:Citrus III viroid
387:Hop latent viroid
233:hepatitis B virus
229:hepatitis D virus
191:RNA polymerase II
105:
104:
4874:
4844:
4843:
4832:
4831:
4830:
4820:
4819:
4811:
4796:
4795:
4783:
4782:
4770:
4769:
4757:
4756:
4755:
4742:
4741:
4740:
4710:
4709:
4321:
4320:
4261:
4260:
4244:
4243:
4128:
4127:
4105:
4098:
4091:
4082:
4081:
3755:Gene duplication
3578:
3577:
3574:self-replication
3462:
3461:
3424:
3423:
3282:Self-replicating
3275:
3268:
3261:
3252:
3251:
3228:
3227:
3225:
3223:
3203:
3197:
3196:
3180:
3174:
3173:
3154:10.1038/273203a0
3129:
3123:
3122:
3094:
3088:
3087:
3077:
3045:
3039:
3038:
3028:
3018:
2986:
2980:
2979:
2961:
2952:
2951:
2935:
2929:
2928:
2918:
2894:
2885:
2882:
2876:
2875:
2873:
2871:
2853:
2823:
2814:
2813:
2811:
2809:
2787:
2778:
2777:
2767:
2757:
2733:
2727:
2726:
2704:
2698:
2697:
2687:
2661:
2652:
2643:
2642:
2632:
2622:
2590:
2584:
2583:
2573:
2549:
2540:
2539:
2529:
2505:
2499:
2498:
2488:
2478:
2446:
2440:
2439:
2437:
2435:
2424:
2418:
2417:
2407:
2397:
2373:
2367:
2366:
2356:
2324:
2318:
2317:
2281:
2272:
2271:
2251:
2227:
2221:
2220:
2210:
2200:
2183:(9): 3275–3280.
2168:
2162:
2161:
2129:
2123:
2122:
2094:
2088:
2087:
2077:
2067:
2043:
2037:
2036:
2008:
2002:
2001:
1983:
1958:
1957:
1947:
1937:
1913:
1907:
1906:
1896:
1878:
1846:
1837:
1836:
1826:
1786:
1777:
1776:
1766:
1748:
1724:
1715:
1714:
1704:
1686:
1662:
1656:
1655:
1645:
1635:
1603:
1597:
1596:
1586:
1576:
1552:
1546:
1545:
1525:
1519:
1518:
1500:
1476:
1470:
1469:
1441:
1430:
1429:
1427:
1426:
1415:
1406:
1405:
1377:
1368:
1367:
1349:
1343:
1342:
1332:
1314:
1290:
1275:
1274:
1264:
1254:
1230:
1224:
1223:
1213:
1203:
1179:
1173:
1172:
1170:
1168:
1159:. Archived from
1152:
1146:
1145:
1127:
1103:
1019:human microbiome
839:human microbiome
554:; type species:
539:; type species:
524:; type species:
482:; type species:
409:; type species:
395:Citrus IV viroid
368:; type species:
356:Hop stunt viroid
353:; type species:
331:Iresine 1 viroid
272:; type species:
54:
53:
40:
28:
27:
4882:
4881:
4877:
4876:
4875:
4873:
4872:
4871:
4867:Subviral agents
4852:
4851:
4850:
4838:
4828:
4826:
4814:
4806:
4804:
4799:
4791:
4786:
4778:
4773:
4765:
4760:
4751:
4750:
4745:
4736:
4735:
4730:
4717:
4707:
4702:
4542:
4532:
4502:
4477:
4388:Finnlakeviridae
4333:
4312:
4254:
4250:
4231:
4203:
4168:
4119:
4109:
4079:
4074:
4024:Synthetic virus
4012:Artificial cell
3985:
3913:
3822:
3711:RNA replication
3706:DNA replication
3694:
3685:Group II intron
3583:
3573:
3561:
3552:Mammalian prion
3534:
3510:
3489:dsRNA satellite
3486:ssDNA satellite
3456:
3449:
3418:
3411:
3356:
3285:
3279:
3236:
3231:
3221:
3219:
3204:
3200:
3181:
3177:
3140:(5659): 203–8.
3130:
3126:
3095:
3091:
3046:
3042:
2987:
2983:
2976:
2962:
2955:
2936:
2932:
2895:
2888:
2883:
2879:
2869:
2867:
2824:
2817:
2807:
2805:
2788:
2781:
2748:(3): e0121922.
2734:
2730:
2723:
2705:
2701:
2659:
2653:
2646:
2605:(19): 7720–25.
2591:
2587:
2550:
2543:
2506:
2502:
2447:
2443:
2433:
2431:
2426:
2425:
2421:
2374:
2370:
2333:Virus Evolution
2325:
2321:
2282:
2275:
2260:
2228:
2224:
2169:
2165:
2130:
2126:
2095:
2091:
2044:
2040:
2009:
2005:
1998:
1984:
1961:
1914:
1910:
1847:
1840:
1787:
1780:
1725:
1718:
1663:
1659:
1604:
1600:
1553:
1549:
1526:
1522:
1491:(11): 2168–79.
1477:
1473:
1442:
1433:
1424:
1422:
1417:
1416:
1409:
1378:
1371:
1364:
1350:
1346:
1291:
1278:
1231:
1227:
1180:
1176:
1166:
1164:
1153:
1149:
1104:
1100:
1096:
1091:
1052:Plant pathology
1027:
971:
939:
907:retrotransposon
851:
829:After applying
827:
821:
778:
738:
722:
692:
624:
569:
241:
222:retrotransposon
163:subviral agents
95:
67:Subviral agents
48:
26:
17:
12:
11:
5:
4880:
4870:
4869:
4864:
4849:
4848:
4836:
4824:
4801:
4800:
4798:
4797:
4784:
4771:
4758:
4743:
4727:
4725:
4719:
4718:
4704:
4703:
4701:
4700:
4695:
4690:
4689:
4688:
4678:
4673:
4668:
4663:
4662:
4661:
4656:
4651:
4646:
4638:
4630:
4629:
4628:
4618:
4613:
4608:
4599:
4594:
4589:
4584:
4579:
4574:
4569:
4568:
4567:
4562:
4552:
4546:
4544:
4538:
4537:
4534:
4533:
4531:
4530:
4525:
4518:
4516:
4512:
4511:
4508:
4507:
4504:
4503:
4501:
4500:
4493:
4485:
4483:
4479:
4478:
4476:
4475:
4468:
4461:
4454:
4447:
4440:
4437:Polydnaviridae
4433:
4426:
4419:
4412:
4405:
4402:Globuloviridae
4398:
4395:Fuselloviridae
4391:
4384:
4377:
4374:Bicaudaviridae
4370:
4363:
4356:
4353:Ampullaviridae
4349:
4341:
4339:
4335:
4334:
4330:Naldaviricetes
4327:
4325:
4318:
4314:
4313:
4311:
4310:
4303:
4296:
4289:
4282:
4275:
4267:
4265:
4258:
4241:
4237:
4236:
4233:
4232:
4230:
4229:
4223:
4215:
4213:
4210:Incertae sedis
4205:
4204:
4202:
4201:
4194:
4189:
4184:
4178:
4176:
4170:
4169:
4167:
4166:
4161:
4160:
4159:
4154:
4144:
4138:
4136:
4125:
4121:
4120:
4108:
4107:
4100:
4093:
4085:
4076:
4075:
4073:
4072:
4067:
4066:
4065:
4060:
4050:
4044:
4038:
4037:
4036:
4031:
4021:
4016:
4015:
4014:
4009:
3999:
3993:
3991:
3987:
3986:
3984:
3983:
3982:
3981:
3976:
3968:
3963:
3958:
3953:
3947:
3946:
3945:
3934:
3929:
3923:
3921:
3915:
3914:
3912:
3911:
3906:
3905:
3904:
3899:
3891:
3889:Kappa organism
3886:
3885:
3884:
3879:
3874:
3869:
3864:
3854:
3853:
3852:
3847:
3836:
3834:
3828:
3827:
3824:
3823:
3821:
3820:
3819:
3818:
3813:
3803:
3802:
3801:
3796:
3791:
3786:
3776:
3775:
3774:
3764:
3763:
3762:
3760:Non-coding DNA
3757:
3752:
3742:
3741:
3740:
3735:
3730:
3725:
3715:
3714:
3713:
3702:
3700:
3696:
3695:
3693:
3692:
3687:
3682:
3680:Group I intron
3677:
3672:
3671:
3670:
3660:
3659:
3658:
3655:
3646:
3643:
3638:
3633:
3623:
3622:
3621:
3616:
3606:
3605:
3604:
3602:Genomic island
3599:
3588:
3586:
3582:Mobile genetic
3575:
3567:
3566:
3563:
3562:
3560:
3559:
3554:
3548:
3546:
3540:
3539:
3536:
3535:
3533:
3532:
3531:
3530:
3527:
3518:
3516:
3512:
3511:
3509:
3508:
3507:
3506:
3503:
3497:
3490:
3487:
3484:
3481:
3474:
3470:
3468:
3459:
3451:
3450:
3448:
3447:
3440:
3432:
3430:
3421:
3413:
3412:
3410:
3409:
3407:dsDNA-RT virus
3404:
3402:ssRNA-RT virus
3399:
3397:(−)ssRNA virus
3394:
3392:(+)ssRNA virus
3389:
3384:
3379:
3378:
3377:
3366:
3364:
3358:
3357:
3355:
3354:
3353:
3352:
3347:
3337:Incertae sedis
3333:
3332:
3331:
3326:
3321:
3316:
3306:
3301:
3295:
3293:
3287:
3286:
3278:
3277:
3270:
3263:
3255:
3249:
3248:
3242:
3235:
3234:External links
3232:
3230:
3229:
3216:New York Times
3198:
3175:
3124:
3089:
3040:
3001:(11): 3852–6.
2981:
2974:
2953:
2930:
2886:
2877:
2815:
2779:
2728:
2721:
2713:Plant virology
2699:
2644:
2585:
2564:(2): 277–286.
2541:
2500:
2441:
2419:
2368:
2339:(2): veaa076.
2319:
2286:Virus Research
2273:
2258:
2222:
2163:
2144:(2): 654–662.
2124:
2089:
2038:
2003:
1997:978-0123751485
1996:
1959:
1908:
1838:
1801:(8): 985–991.
1778:
1733:Genome Biology
1716:
1657:
1618:(23): 9370–4.
1598:
1547:
1520:
1471:
1431:
1407:
1369:
1362:
1344:
1305:(3): 646–661.
1276:
1225:
1174:
1147:
1118:(1): 305–325.
1097:
1095:
1092:
1090:
1089:
1084:
1079:
1074:
1069:
1064:
1059:
1054:
1049:
1044:
1039:
1034:
1028:
1026:
1023:
1007:New York Times
970:
967:
938:
935:
898:
897:
890:
889:-free habitat.
883:
880:
877:
866:
850:
847:
823:Main article:
820:
817:
813:pararetrovirus
809:satellite RNAs
777:
774:
750:RNA polymerase
737:
734:
726:satellite RNAs
721:
718:
691:
688:
623:
620:
609:rolling circle
568:
565:
564:
563:
562:
561:
546:
531:
509:
508:
507:
506:
505:
497:
474:
473:
472:
464:
456:
448:
440:
432:
424:
401:
400:
399:
391:
383:
374:; 246–247 nt
360:
345:
344:
343:
335:
327:
319:
311:
303:
295:
287:
240:
237:
199:rolling circle
103:
102:
86:
85:
81:
80:
75:
71:
70:
63:
60:Informal group
56:
55:
42:
41:
33:
32:
15:
9:
6:
4:
3:
2:
4879:
4868:
4865:
4863:
4860:
4859:
4857:
4847:
4842:
4837:
4835:
4825:
4823:
4818:
4813:
4812:
4809:
4794:
4789:
4785:
4781:
4776:
4772:
4768:
4763:
4759:
4754:
4748:
4744:
4739:
4733:
4729:
4728:
4726:
4724:
4720:
4716:
4711:
4699:
4696:
4694:
4691:
4687:
4684:
4683:
4682:
4679:
4677:
4676:Tandem repeat
4674:
4672:
4669:
4667:
4664:
4660:
4657:
4655:
4652:
4650:
4647:
4645:
4642:
4641:
4639:
4637:
4634:
4631:
4627:
4624:
4623:
4622:
4619:
4617:
4614:
4612:
4609:
4606:
4605:
4604:Nanobacterium
4600:
4598:
4595:
4593:
4590:
4588:
4585:
4583:
4580:
4578:
4575:
4573:
4570:
4566:
4563:
4561:
4558:
4557:
4556:
4553:
4551:
4548:
4547:
4545:
4539:
4529:
4526:
4523:
4520:
4519:
4517:
4513:
4499:
4498:
4497:Rhizidiovirus
4494:
4492:
4491:
4487:
4486:
4484:
4480:
4474:
4473:
4469:
4467:
4466:
4465:Thaspiviridae
4462:
4460:
4459:
4455:
4453:
4452:
4451:Pospiviroidae
4448:
4446:
4445:
4441:
4439:
4438:
4434:
4432:
4431:
4430:Plasmaviridae
4427:
4425:
4424:
4420:
4418:
4417:
4416:Halspiviridae
4413:
4411:
4410:
4406:
4404:
4403:
4399:
4397:
4396:
4392:
4390:
4389:
4385:
4383:
4382:
4378:
4376:
4375:
4371:
4369:
4368:
4367:Avsunviroidae
4364:
4362:
4361:
4360:Anelloviridae
4357:
4355:
4354:
4350:
4348:
4347:
4343:
4342:
4340:
4336:
4332:
4331:
4326:
4322:
4319:
4315:
4309:
4308:
4304:
4302:
4301:
4297:
4295:
4294:
4290:
4288:
4287:
4283:
4281:
4280:
4279:Duplodnaviria
4276:
4274:
4273:
4269:
4268:
4266:
4262:
4259:
4257:
4253:
4249:
4245:
4242:
4238:
4227:
4224:
4222:
4221:
4217:
4216:
4214:
4212:
4211:
4206:
4199:
4195:
4193:
4190:
4188:
4185:
4183:
4180:
4179:
4177:
4175:
4171:
4165:
4162:
4158:
4155:
4153:
4152:Mitochondrion
4150:
4149:
4148:
4145:
4143:
4140:
4139:
4137:
4134:
4129:
4126:
4124:Cellular life
4122:
4117:
4113:
4106:
4101:
4099:
4094:
4092:
4087:
4086:
4083:
4071:
4068:
4064:
4061:
4059:
4056:
4055:
4054:
4051:
4049:
4045:
4043:
4042:Nanobacterium
4039:
4035:
4032:
4030:
4027:
4026:
4025:
4022:
4020:
4017:
4013:
4010:
4008:
4007:Cell division
4005:
4004:
4003:
4000:
3998:
3995:
3994:
3992:
3988:
3980:
3977:
3975:
3972:
3971:
3969:
3967:
3964:
3962:
3959:
3957:
3954:
3952:
3948:
3944:
3941:
3940:
3939:
3935:
3933:
3930:
3928:
3925:
3924:
3922:
3920:
3916:
3910:
3907:
3903:
3900:
3898:
3895:
3894:
3892:
3890:
3887:
3883:
3880:
3878:
3875:
3873:
3870:
3868:
3865:
3863:
3860:
3859:
3858:
3855:
3851:
3850:Hydrogenosome
3848:
3846:
3843:
3842:
3841:
3840:Mitochondrion
3838:
3837:
3835:
3833:
3832:Endosymbiosis
3829:
3817:
3814:
3812:
3811:Tandem repeat
3809:
3808:
3807:
3804:
3800:
3797:
3795:
3792:
3790:
3787:
3785:
3782:
3781:
3780:
3777:
3773:
3770:
3769:
3768:
3765:
3761:
3758:
3756:
3753:
3751:
3748:
3747:
3746:
3743:
3739:
3736:
3734:
3731:
3729:
3726:
3724:
3721:
3720:
3719:
3716:
3712:
3709:
3708:
3707:
3704:
3703:
3701:
3699:Other aspects
3697:
3691:
3688:
3686:
3683:
3681:
3678:
3676:
3673:
3669:
3666:
3665:
3664:
3661:
3656:
3654:
3650:
3647:
3644:
3642:
3639:
3637:
3634:
3632:
3629:
3628:
3627:
3624:
3620:
3617:
3615:
3612:
3611:
3610:
3607:
3603:
3600:
3598:
3595:
3594:
3593:
3590:
3589:
3587:
3585:
3579:
3576:
3572:
3568:
3558:
3555:
3553:
3550:
3549:
3547:
3545:
3541:
3528:
3525:
3524:
3523:
3520:
3519:
3517:
3513:
3504:
3501:
3500:
3498:
3495:
3491:
3488:
3485:
3482:
3479:
3475:
3472:
3471:
3469:
3467:
3463:
3460:
3458:
3452:
3446:
3445:
3444:Avsunviroidae
3441:
3439:
3438:
3437:Pospiviroidae
3434:
3433:
3431:
3429:
3425:
3422:
3420:
3414:
3408:
3405:
3403:
3400:
3398:
3395:
3393:
3390:
3388:
3385:
3383:
3380:
3376:
3373:
3372:
3371:
3368:
3367:
3365:
3363:
3359:
3351:
3348:
3346:
3345:
3341:
3340:
3339:
3338:
3334:
3330:
3327:
3325:
3322:
3320:
3317:
3315:
3312:
3311:
3310:
3307:
3305:
3302:
3300:
3297:
3296:
3294:
3292:
3291:Cellular life
3288:
3283:
3276:
3271:
3269:
3264:
3262:
3257:
3256:
3253:
3246:
3243:
3241:
3238:
3237:
3218:
3217:
3212:
3208:
3202:
3194:
3190:
3186:
3179:
3171:
3167:
3163:
3159:
3155:
3151:
3147:
3143:
3139:
3135:
3128:
3120:
3116:
3112:
3108:
3104:
3100:
3093:
3085:
3081:
3076:
3071:
3067:
3063:
3059:
3055:
3051:
3044:
3036:
3032:
3027:
3022:
3017:
3012:
3008:
3004:
3000:
2996:
2992:
2985:
2977:
2971:
2967:
2960:
2958:
2949:
2945:
2941:
2934:
2926:
2922:
2917:
2912:
2908:
2904:
2900:
2893:
2891:
2881:
2865:
2861:
2857:
2852:
2847:
2843:
2839:
2835:
2834:
2829:
2822:
2820:
2803:
2799:
2798:
2793:
2786:
2784:
2775:
2771:
2766:
2761:
2756:
2751:
2747:
2743:
2739:
2732:
2724:
2718:
2714:
2710:
2703:
2695:
2691:
2686:
2681:
2677:
2673:
2669:
2665:
2658:
2651:
2649:
2640:
2636:
2631:
2626:
2621:
2616:
2612:
2608:
2604:
2600:
2596:
2589:
2581:
2577:
2572:
2567:
2563:
2559:
2555:
2548:
2546:
2537:
2533:
2528:
2523:
2519:
2515:
2511:
2504:
2496:
2492:
2487:
2482:
2477:
2472:
2468:
2464:
2460:
2456:
2452:
2445:
2429:
2423:
2415:
2411:
2406:
2401:
2396:
2391:
2387:
2383:
2379:
2372:
2364:
2360:
2355:
2350:
2346:
2342:
2338:
2334:
2330:
2323:
2315:
2311:
2307:
2303:
2299:
2295:
2291:
2287:
2280:
2278:
2269:
2265:
2261:
2255:
2250:
2245:
2241:
2237:
2233:
2226:
2218:
2214:
2209:
2204:
2199:
2194:
2190:
2186:
2182:
2178:
2174:
2167:
2159:
2155:
2151:
2147:
2143:
2139:
2135:
2128:
2120:
2116:
2112:
2108:
2104:
2100:
2093:
2085:
2081:
2076:
2071:
2066:
2061:
2057:
2053:
2049:
2042:
2034:
2030:
2026:
2022:
2018:
2014:
2007:
1999:
1993:
1989:
1982:
1980:
1978:
1976:
1974:
1972:
1970:
1968:
1966:
1964:
1955:
1951:
1946:
1941:
1936:
1931:
1927:
1923:
1919:
1912:
1904:
1900:
1895:
1890:
1886:
1882:
1877:
1872:
1868:
1864:
1860:
1856:
1852:
1845:
1843:
1834:
1830:
1825:
1820:
1816:
1812:
1808:
1804:
1800:
1796:
1792:
1785:
1783:
1774:
1770:
1765:
1760:
1756:
1752:
1747:
1742:
1738:
1734:
1730:
1723:
1721:
1712:
1708:
1703:
1698:
1694:
1690:
1685:
1680:
1676:
1672:
1668:
1661:
1653:
1649:
1644:
1639:
1634:
1629:
1625:
1621:
1617:
1613:
1609:
1602:
1594:
1590:
1585:
1580:
1575:
1570:
1566:
1562:
1558:
1551:
1543:
1539:
1535:
1531:
1524:
1516:
1512:
1508:
1504:
1499:
1494:
1490:
1486:
1482:
1475:
1467:
1463:
1459:
1455:
1451:
1447:
1440:
1438:
1436:
1420:
1414:
1412:
1403:
1399:
1395:
1391:
1388:(2): 411–28.
1387:
1383:
1376:
1374:
1365:
1359:
1355:
1348:
1340:
1336:
1331:
1326:
1322:
1318:
1313:
1308:
1304:
1300:
1296:
1289:
1287:
1285:
1283:
1281:
1272:
1268:
1263:
1258:
1253:
1248:
1244:
1240:
1236:
1229:
1221:
1217:
1212:
1207:
1202:
1197:
1193:
1189:
1185:
1178:
1162:
1158:
1151:
1143:
1139:
1135:
1131:
1126:
1121:
1117:
1113:
1109:
1102:
1098:
1088:
1085:
1083:
1080:
1078:
1075:
1073:
1070:
1068:
1065:
1063:
1060:
1058:
1055:
1053:
1050:
1048:
1045:
1043:
1040:
1038:
1037:Microparasite
1035:
1033:
1030:
1029:
1022:
1020:
1016:
1011:
1009:
1008:
1002:
999:
995:
991:
987:
984:
980:
975:
966:
964:
960:
956:
952:
948:
944:
934:
932:
928:
924:
920:
916:
912:
908:
905:(a family of
904:
901:discovery of
895:
891:
888:
884:
881:
878:
875:
871:
867:
864:
863:
862:
860:
856:
846:
844:
840:
836:
832:
826:
816:
814:
810:
806:
802:
797:
795:
791:
790:recombination
787:
783:
773:
771:
767:
766:
761:
760:
759:Orthornavirae
755:
751:
747:
743:
733:
731:
727:
717:
715:
711:
710:
705:
701:
697:
687:
685:
681:
677:
673:
669:
665:
661:
657:
656:messenger RNA
653:
649:
645:
641:
637:
636:RNA silencing
633:
629:
622:RNA silencing
619:
617:
612:
610:
606:
605:messenger RNA
602:
598:
597:Avsunviroidae
594:
590:
589:Pospiviroidae
586:
578:
577:plasmodesmata
573:
559:
558:
553:
552:
547:
544:
543:
538:
537:
532:
529:
528:
523:
522:
517:
516:
514:
513:Avsunviroidae
510:
503:
502:
498:
495:
494:
490:
489:
488:; 248–251 nt
487:
486:
481:
480:
475:
470:
469:
465:
462:
461:
457:
454:
453:
449:
446:
445:
441:
438:
437:
433:
430:
429:
425:
422:
421:
417:
416:
415:; 329–334 nt
414:
413:
408:
407:
402:
397:
396:
392:
389:
388:
384:
381:
380:
376:
375:
373:
372:
367:
366:
361:
358:
357:
352:
351:
346:
341:
340:
336:
333:
332:
328:
325:
324:
320:
317:
316:
312:
309:
308:
304:
301:
300:
296:
293:
292:
288:
285:
284:
280:
279:
277:
276:
271:
270:
265:
264:
262:
261:Pospiviroidae
258:
257:
256:
250:
245:
236:
234:
230:
225:
223:
219:
215:
210:
208:
204:
200:
196:
195:messenger RNA
192:
188:
183:
181:
180:
175:
171:
166:
164:
160:
156:
152:
148:
144:
140:
135:
133:
129:
125:
121:
117:
113:
112:circular RNAs
109:
101:
100:
99:
98:Avsunviroidae
94:
93:
92:Pospiviroidae
87:
82:
79:
76:
73:
72:
69:
68:
64:
61:
58:
57:
52:
47:
43:
39:
34:
29:
24:
19:
4722:
4693:Transpoviron
4626:Fungal prion
4602:
4495:
4490:Dinodnavirus
4488:
4470:
4463:
4458:Spiraviridae
4456:
4449:
4442:
4435:
4428:
4423:Ovaliviridae
4421:
4414:
4409:Guttaviridae
4407:
4400:
4393:
4386:
4381:Clavaviridae
4379:
4372:
4365:
4358:
4351:
4344:
4328:
4307:Varidnaviria
4305:
4298:
4291:
4286:Monodnaviria
4284:
4277:
4270:
4251:
4218:
4208:
4029:Viral vector
3872:Gerontoplast
3799:Transpoviron
3571:Nucleic acid
3557:Fungal prion
3455:Helper-virus
3442:
3435:
3427:
3342:
3335:
3240:Viroids/ATSU
3222:November 22,
3220:. Retrieved
3214:
3201:
3184:
3178:
3137:
3133:
3127:
3105:(1): 70–80.
3102:
3098:
3092:
3057:
3053:
3043:
2998:
2994:
2984:
2965:
2939:
2933:
2906:
2902:
2880:
2868:. Retrieved
2831:
2806:. Retrieved
2797:ScienceAlert
2795:
2745:
2741:
2731:
2712:
2702:
2685:10261/107724
2667:
2663:
2602:
2598:
2588:
2561:
2557:
2517:
2513:
2503:
2458:
2454:
2444:
2432:. Retrieved
2422:
2385:
2381:
2371:
2336:
2332:
2322:
2289:
2285:
2231:
2225:
2180:
2176:
2166:
2141:
2137:
2127:
2102:
2098:
2092:
2055:
2051:
2041:
2019:(2): 31–34.
2016:
2012:
2006:
1987:
1925:
1921:
1911:
1858:
1854:
1798:
1794:
1736:
1732:
1674:
1670:
1660:
1615:
1611:
1601:
1564:
1560:
1550:
1536:(1): 65–73.
1533:
1529:
1523:
1488:
1484:
1474:
1449:
1445:
1423:. Retrieved
1421:. 2006-03-02
1385:
1381:
1353:
1347:
1302:
1298:
1245:(15): 5532.
1242:
1238:
1228:
1191:
1187:
1177:
1165:. Retrieved
1161:the original
1150:
1115:
1111:
1101:
1032:Circular RNA
1012:
1005:
1003:
1000:
996:
992:
988:
976:
972:
940:
899:
852:
828:
805:retroviruses
798:
779:
776:Retroviroids
765:Ambiviricota
763:
757:
744:. Their RNA
739:
724:Viroid-like
723:
707:
693:
658:. Secondly,
654:with target
652:base pairing
625:
613:
596:
593:chloroplasts
588:
582:
560:; 332–335 nt
555:
549:
545:; 335–351 nt
540:
536:Pelamoviroid
534:
530:; 246–251 nt
525:
519:
499:
491:
483:
477:
466:
458:
450:
442:
434:
426:
418:
410:
404:
393:
385:
377:
369:
363:
359:; 294–303 nt
354:
348:
337:
329:
321:
313:
310:; 368–467 nt
305:
297:
289:
281:
273:
267:
255:As of 2005:
254:
226:
211:
184:
177:
167:
136:
107:
106:
96:
90:
89:
77:
74:(unranked):
66:
59:
18:
4747:Wikispecies
4636:microsphere
4555:Cancer cell
4300:Ribozyviria
4053:Cancer cell
3919:Abiogenesis
3867:Chromoplast
3862:Chloroplast
3645:Degradative
3387:dsRNA virus
3382:ssDNA virus
3375:Giant virus
3370:dsDNA virus
2670:: 395–414.
2249:10261/74632
1795:RNA Biology
1677:(7): 3476.
1567:(7): 3476.
1167:February 3,
927:prokaryotes
915:prokaryotes
868:Their high
736:Ambiviruses
709:Ribozyviria
698:", fungal "
521:Avsunviroid
406:Apscaviroid
365:Cocadviroid
350:Hostuviroid
269:Pospiviroid
132:prokaryotes
120:angiosperms
4856:Categories
4686:Retroposon
4633:Proteinoid
4543:structures
4541:Comparable
4317:Unassigned
4220:Parakaryon
4133:Prokaryota
3961:Proteinoid
3956:Coacervate
3909:Nitroplast
3902:Trophosome
3897:Bacteriome
3882:Apicoplast
3877:Leucoplast
3718:Chromosome
3636:Resistance
3344:Parakaryon
2909:: 425–44.
2870:29 January
2808:29 January
2292:: 198709.
1928:: 560105.
1861:(1): 103.
1739:(1): 135.
1452:: 117–39.
1425:2007-07-18
1194:(2): 120.
1094:References
963:quarantine
917:including
903:retrozymes
843:homologies
786:homologous
754:ribovirian
664:transgenic
479:Coleviroid
218:retrozymes
4293:Riboviria
4272:Adnaviria
4256:Satellite
4174:Eukaryota
3970:Research
3951:Protocell
3690:Retrozyme
3649:Virulence
3631:Fertility
3478:Virophage
3466:Satellite
3457:dependent
3309:Eukaryota
3060:: 32–37.
2314:246987005
1885:2075-1729
1815:1547-6286
1755:1474-760X
1693:1422-0067
1515:221581424
1481:"Viroids"
1157:"Viroids"
1134:2327-056X
1004:In 2014,
983:bacterium
894:ribozymes
859:RNA world
770:ribozymes
700:ambivirus
696:virusoids
644:virulence
551:Elaviroid
247:Putative
214:RNA world
207:catalytic
205:, having
203:ribozymes
139:biosphere
84:Families
4834:Medicine
4767:60605243
4732:Wikidata
4666:Ribozyme
4611:Phagemid
4338:Families
4198:Protista
4182:Animalia
4147:Bacteria
3997:Organism
3990:See also
3966:Sulphobe
3943:Ribozyme
3938:RNA life
3845:Mitosome
3789:Prophage
3784:Provirus
3772:Replicon
3728:Circular
3675:Phagemid
3592:Mobilome
3584:elements
3494:Virusoid
3417:Subviral
3329:Protista
3314:Animalia
3299:Bacteria
3245:ViroidDB
3207:Zimmer C
3170:19398777
3099:Virology
3084:33460914
2925:26047558
2864:Archived
2860:38293115
2851:10827157
2802:Archived
2774:36840552
2765:10019309
2694:25002087
2639:23610439
2580:15246267
2558:Virology
2536:11172112
2414:37275138
2405:10237039
2363:33324490
2306:35183574
2268:22678590
2217:14978267
2138:Virology
2119:28738223
2084:22719735
2033:44660564
1954:24198831
1903:35054497
1833:28448743
1773:27339130
1711:33800543
1593:33800543
1507:18764915
1466:16078879
1382:Virology
1339:36696902
1271:32752288
1220:30699972
1142:34255541
1087:Virusoid
1025:See also
1015:obelisks
919:bacteria
887:ribosome
874:cytosine
835:obelisks
819:Obelisks
756:kingdom
714:ribozyme
628:symptoms
239:Taxonomy
124:economic
23:Virusoid
4862:Viroids
4846:Viruses
4822:Biology
4808:Portals
4753:Viroids
4738:Q209917
4723:Viroids
4698:Xenobot
4616:Plasmid
4597:Jeewanu
4528:Obelisk
4324:Classes
4192:Plantae
4157:Plastid
4142:Archaea
3979:Jeewanu
3893:Organs
3857:Plastid
3657:Cryptic
3626:Plasmid
3324:Plantae
3304:Archaea
3142:Bibcode
3119:4728831
3075:8068583
3035:1069269
3003:Bibcode
2833:bioRxiv
2630:3651503
2607:Bibcode
2495:7542779
2463:Bibcode
2354:7724248
2185:Bibcode
2158:1546460
2105:: 1–7.
2075:3376415
2058:: 217.
1945:3807834
1894:8781251
1863:Bibcode
1824:5680766
1764:4918200
1702:8036462
1652:2480600
1620:Bibcode
1584:8036462
1402:5095900
1330:9911046
1307:bioRxiv
1262:7432327
1211:6409718
1188:Viruses
1057:Plasmid
969:History
937:Control
923:archaea
870:guanine
855:introns
746:genomes
730:capsids
632:protein
511:Family
259:Family
187:protein
116:viruses
108:Viroids
31:Viroid
4780:1VIRDD
4644:Retron
4587:Fosmid
4572:Cosmid
4522:Nanobe
4482:Genera
4264:Realms
4252:Viroid
4070:Virome
4048:Nanobe
3745:Genome
3723:Linear
3668:Fosmid
3663:Cosmid
3428:Viroid
3419:agents
3168:
3162:643081
3160:
3134:Nature
3117:
3082:
3072:
3033:
3026:431239
3023:
2972:
2923:
2858:
2848:
2772:
2762:
2719:
2692:
2637:
2627:
2578:
2534:
2493:
2483:
2430:. ICTV
2412:
2402:
2361:
2351:
2312:
2304:
2266:
2256:
2215:
2208:365780
2205:
2156:
2117:
2082:
2072:
2031:
1994:
1952:
1942:
1901:
1891:
1883:
1831:
1821:
1813:
1771:
1761:
1753:
1709:
1699:
1691:
1650:
1643:298497
1640:
1591:
1581:
1513:
1505:
1464:
1400:
1360:
1337:
1327:
1309:
1269:
1259:
1218:
1208:
1140:
1132:
979:fungus
961:, and
949:, and
931:CRISPR
911:plants
706:-like
660:siRNAs
648:siRNAs
640:genome
585:aphids
548:Genus
533:Genus
518:Genus
476:Genus
403:Genus
362:Genus
347:Genus
266:Genus
78:Viroid
4793:12884
4621:Prion
4515:Other
4248:Virus
4187:Fungi
3544:Prion
3515:Other
3362:Virus
3319:Fungi
3166:S2CID
2660:(PDF)
2486:41428
2434:2 May
2310:S2CID
2029:S2CID
1511:S2CID
1077:Virus
1062:Prion
943:ELISA
742:fungi
680:dicer
668:hpRNA
591:) or
159:order
4788:NCBI
4775:EPPO
4560:HeLa
4164:LUCA
4112:Life
4058:HeLa
4002:Cell
3750:Gene
3224:2014
3158:PMID
3115:PMID
3080:PMID
3031:PMID
2970:ISBN
2921:PMID
2872:2024
2856:PMID
2810:2024
2770:PMID
2717:ISBN
2690:PMID
2635:PMID
2576:PMID
2532:PMID
2491:PMID
2436:2024
2410:PMID
2359:PMID
2302:PMID
2264:PMID
2254:ISBN
2213:PMID
2154:PMID
2115:PMID
2080:PMID
1992:ISBN
1950:PMID
1926:2013
1899:PMID
1881:ISSN
1855:Life
1829:PMID
1811:ISSN
1769:PMID
1751:ISSN
1707:PMID
1689:ISSN
1648:PMID
1589:PMID
1503:PMID
1462:PMID
1398:PMID
1358:ISBN
1335:PMID
1299:Cell
1267:PMID
1216:PMID
1169:2021
1138:PMID
1130:ISSN
921:and
872:and
149:and
4762:EoL
4524:(?)
4228:(?)
3641:Col
3529:DNA
3526:RNA
3505:DNA
3502:RNA
3189:doi
3150:doi
3138:273
3107:doi
3070:PMC
3062:doi
3021:PMC
3011:doi
2944:doi
2911:doi
2846:PMC
2838:doi
2760:PMC
2750:doi
2680:hdl
2672:doi
2625:PMC
2615:doi
2603:110
2566:doi
2562:325
2522:doi
2481:PMC
2471:doi
2400:PMC
2390:doi
2349:PMC
2341:doi
2294:doi
2290:312
2244:hdl
2236:doi
2203:PMC
2193:doi
2181:101
2146:doi
2142:187
2107:doi
2070:PMC
2060:doi
2021:doi
1940:PMC
1930:doi
1889:PMC
1871:doi
1819:PMC
1803:doi
1759:PMC
1741:doi
1697:PMC
1679:doi
1638:PMC
1628:doi
1579:PMC
1569:doi
1538:doi
1493:doi
1454:doi
1390:doi
1325:PMC
1317:doi
1303:186
1257:PMC
1247:doi
1206:PMC
1196:doi
1120:doi
981:or
947:PCR
782:DNA
704:HDV
676:RNA
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