Viroids are small single-stranded, circular RNAs that are infectious pathogens. Unlike viruses, they have no protein coating. A 2023 metatranscriptomics study suggests that viroids and viroid-like elements can be found in all domains of life. The first discoveries of viroids in the 1970s triggered the historically third major extension of the biosphere—to include smaller entities—after the discoveries in 1675 by Antonie van Leeuwenhoek (of the "subvisible" protozoa) and in 1892–1898 by Dmitri Iosifovich Ivanovsky and Martinus Beijerinck (of the "submicroscopic" viruses). The unique properties of viroids have been recognized by the International Committee on Taxonomy of Viruses, in creating a new order of subviral agents. The first recognized viroid, the pathogenic agent of the potato spindle tuber disease, was discovered, initially molecularly characterized, and named by Theodor Otto Diener, a 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 Citrus exocortis viroid (CEVd) was discovered soon thereafter, and together understanding of PSTVd and CEVd shaped the concept of the viroid. Although viroids are composed of nucleic acid, they do not code for any protein. The viroid's replication mechanism uses RNA polymerase II, a host cell enzyme normally associated with synthesis of messenger RNA from DNA, which instead catalyzes "rolling circle" synthesis of new RNA using the viroid's RNA as a template. Viroids are often ribozymes, having catalytic properties that allow self-cleavage and ligation of unit-size genomes from larger replication intermediates. Diener initially hypothesized in 1989 that viroids may represent "living relics" from the widely assumed, ancient, and non-cellular RNA world, and others have followed this conjecture. Following the discovery of retrozymes, it has been proposed that viroids and other viroid-like elements may derive from this newly found class of retrotransposon.
Taxonomy
As of 2024:
Family Pospiviroidae: relies on host Rnase III Genus Pospiviroid; type species: Pospiviroid fusituberis (former name Potato spindle tuber viroid); 356–361 nucleotides(nt) Pospiviroid chloronani (former name Tomato chlorotic dwarf viroid); (TCDVd); accession AF162131, genome length 360nt Mexican papita viroid; (MPVd); accession L78454, genome length 360nt Pospiviroid machoplantae (former name Tomato planta macho viroid); (TPMVd); accession K00817, genome length 360nt Pospiviroid exocortiscitri (former name Citrus exocortis viroid); 368–467 nt Pospiviroid impedichrysanthemi (former name Chrysanthemum stunt viroid); (CSVd); accession V01107, genome length 356nt Pospiviroid apicimpeditum (former name Tomato apical stunt viroid); (TASVd); accession K00818, genome length 360nt Pospiviroid alphairesinis (former name Iresine 1 viroid); (IrVd-1); accession X95734, genome length 370nt Pospiviroid latenscolumneae (former name Columnea latent viroid); (CLVd); accession X15663, genome length 370nt Pospiviroid latensportulacae (former name Pospiviroid plvd) Pospiviroid parvicapsici (former name Pepper chat fruit viroid) Genus Hostuviroid; type species: Hostuviroid impedihumuli (former name Hop stunt viroid); 294–303 nt Hostuviroid latensdahliae (former name Dahlia latent viroid) Genus Cocadviroid; type species: Cocadviroid cadangi (former name Coconut cadang-cadang viroid); 246–247 nt Cocadviroid tinangajae (former name Coconut tinangaja viroid); (CTiVd); accession M20731, genome length 254nt Cocadviroid latenshumuli (former name Hop latent viroid); (HLVd); accession X07397, genome length 256nt Cocadviroid rimocitri (former names Citrus bark cracking viroid, Citrus IV viroid); (CVd-IV); accession X14638, genome length 284nt Genus Apscaviroid; type species: Apscaviroid cicatricimali (former name Apple scar skin viroid); 329–334 nt Citrus III viroid; (CVd-III); accession AF184147, genome length 294nt Apscaviroid fossulamali (former name Apple dimple fruit viroid); (ADFVd); accession X99487, genome length 306nt Apscaviroid alphaflavivitis (former name Grapevine yellow speckle viroid 1); (GVYSd-1); accession X06904, genome length 367nt Apscaviroid betaflavivitis (former name Grapevine yellow speckle viroid 2); (GVYSd-2); accession J04348, genome length 363nt Apscaviroid curvifoliumcitri (former name Citrus bent leaf viroid); (CBLVd); accession M74065, genome length 318nt Apscaviroid pustulapyri (former name Pear blister canker viroid); (PBCVd); accession D12823, genome length 315nt Apscaviroid austravitis (former name Australian grapevine viroid); (AGVd); accession X17101, genome length 369nt Apscaviroid maculamali (former name Apscaviroid aclsvd) Apscaviroid etacitri (former name Apscaviroid cvd-VII) Apscaviroid dendrobii (former name Apscaviroid dvd) Apscaviroid latensvitis (former name Apscaviroid glvd) Apscaviroid litchis (former name Apscaviroid lvd) Apscaviroid latenspruni (former name Apscaviroid plvd-I) Apscaviroid diospyri (former name Apscaviroid pvd) Apscaviroid betadiospyri (former name Apscaviroid pvd-2) Apscaviroid nanocitri (former name Citrus dwarfing viroid) Apscaviroid epsiloncitri (former name Citrus viroid V) Apscaviroid zetacitri (former name Citrus viroid VI) Apscaviroid japanvitis Genus Coleviroid; type species: Coleviroid alphacolei (former name Coleus blumei viroid 1); (CbVd-1); 248–251 nt Coleviroid betacolei (former name Coleus blumei viroid 2); (CbVd-2); accession X95365, genome length 301nt Coleviroid gammacolei (former name Coleus blumei viroid 3); (CbVd-3); accession X95364, genome length 361nt Coleviroid epsiloncolei (former name Coleviroid cbvd-5) Coleviroid zetacolei (former name Coleviroid cbvd-6) Family Avsunviroidae: autocatalytic clevage Genus Avsunviroid; type species: Avsunviroid albamaculaperseae (former name Avocado sunblotch viroid); 246–251 nt Genus Pelamoviroid; type species: Pelamoviroid latenspruni (former name Peach latent mosaic viroid); 335–351 nt Pelamoviroid maculachrysanthemi (former name Chrysanthemum chlorotic mottle viroid) Pelamoviroid malleusmali (former name Apple hammerhead viroid) Genus Elaviroid; type species: Elaviroid latensmelongenae (former name Eggplant latent viroid); 332–335 nt
Transmission and replication
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