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Ordine Tymovirales

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Ordine Tymovirales Ordine Tymovirales Classificazione Dominium/Dominio: Acytota o Aphanobionta Gruppo: IV (Virus a ssRNA+) Ordo/Ordine: Tymovirales Il nome deriva dal genere Tymovirus (e dalla famiglia Tymoviridae). Questo è stato scelto perché le altre famiglie costituenti hanno nomi che riflettono i loro virioni flessi (non una caratteristica di tutti i membri del’'ordine). Tymovirales è un ordine di virus proposto nel 2007 e ufficialmente approvato dall’International Committee on Taxonomy of Viruses nel 2009. Quest’ordine possiede un genoma ad RNA a singolo filamento a senso positivo, di conseguenza fanno parte del gruppo IV secondo la classificazione di Baltimore. I virus appartenenti a quest’ordine hanno, come ospite, le piante. I Tymovirales hanno capside senza pericapside, filamentoso e flessibile o isometrico a simmetria icosaedrica e possiedono tutti una poliproteina di replicazione alpha-like. I Tymovirales, hanno una singola molecola di ssRNA senso positivo e sono uniti dalle somiglianze nelle loro poliproteine associate alla replicazione. I virioni all’interno delle famiglie Alphaflexiviridae, Betaflexiviridae e Gammaflexiviridae sono filamenti flessuosi ed hanno solitamente un diametro di 12-13 nm e una lunghezza compresa tra circa 470 e 1000 nm, a seconda del genere. Hanno una simmetria elicoidale e in alcuni generi c’è un crossbanding ben visibile. Quasi tutti i membri hanno una singola proteina di rivestimento (CP) di 18-44 kDa e nel caso dei generi Lolavirus e alcuni Marafivirus, ci sono due proteine strutturali, che sono di forme diverse dallo stesso genere. La più grande proteina codificata è una poliproteina associata alla replicazione di circa 150-250 kDa vicino all'estremità 5' del genoma e che è tradotta direttamente dall’RNA genomico. La maggior parte dei membri dell’ordine infetta le piante, ma alcune specie provengono da funghi patogeni vegetali. La gamma e la trasmissione dell’ospite sono spesso caratteristiche dei singoli generi. Nell’analisi filogenetica della proteina di replicazione, ogni genere e famiglia forma un ramo distinto e ben supportato. I virus con virioni flessibili delle famiglie Alphaflexiviridae e Gammaflexiviridae sono più strettamente correlati a quelli con particelle icosaedriche nella famiglia Tymoviridae di quanto non lo siano per i membri dei Betaflexiviridae (figura 1). Figura 1 - Albero filogenetico basato sulle sequenze di amminoacidi dell’intera proteina di replicazione dei membri dell’ordine Tymovirales. È stato incluso un singolo isolato rappresentativo di ciascuna specie sequenziata nell’ordine. Generi e famiglie (che sono tutti monofiletici) sono stati collassati in un triangolo, la cui dimensione corrisponde alla variazione trovata all’interno del clade. I numeri sui rami indicano la percentuale del supporto di bootstrap su 1000 repliche di bootstrap (quando> 60%). La scala indica le distanze degli aminoacidi JTT (Jones- Taylor-Thornton matrix). Albero prodotto in MEGA4. L’ordine Tymovirales è diviso in quattro famiglie e sedici generi: 1. Famiglia Alphaflexiviridae, con i seguenti generi: 1. Allexivirus 2. Botrexvirus 3. Lolavirus 4. Mandarivirus 5. Protexvirus 6. Sclerodarnavirus 2. Famiglia Betaflexiviridae, con i seguenti generi: 7. Capillovirus 8. Carlavirus 9. Citrivirus 10. Foveavirus 11. Trichovirus 12. Vitivirus 3. Famiglia Gammaflexiviridae, con il seguente genere: 13. Mycovirus 4. Famiglia Deltaflexiviridae, con il seguente genere: 14. Deltaflexivirus 5. Famiglia Tymoviridae, con i seguenti generi: 14. Maculavirus 15. Maravirus 16. Tymovirus Le specie dell’ordine Tymovirales sono di seguito elencate: A - Alphaflexiviridae 1. Actinidia virus X 2. Alfalfa virus S 3. Allium virus X 4. Alstroemeria virus x 5. Alternanthera mosaic virus 6. Arachis pintoi virus 7. Asparagus virus 3 8. Babaco mosaic virus 9. Bamboo mosaic virus 10. Blackberry virus E 11. Botrytis virus X 12. Cactus virus X 13. Cassava common mosaic virus 14. Cassava virus X 15. Citrus yellow vein clearing virus 16. Clover yellow mosaic virus 17. Cymbidium mosaic virus 18. Donkey orchid symptomless virus 19. Euonymus yellow vein virus 20. Foxtail mosaic virus 21. Garlic virus A 22. Garlic virus B 23. Garlic virus C 24. Garlic virus D 25. Garlic virus E 26. Garlic virus X 27. Hosta virus X 28. Hydrangea ringspot virus 29. Indian citrus ringspot virus 30. Lettuce virus X 31. Lily virus X 32. Lolium latent virus 33. Malva mosaic virus 34. Mint virus X 35. Narcissus mosaic virus 36. Nerine virus X 37. Opuntia virus X 38. Papaya mosaic virus 39. Pepino mosaic virus 40. Phaius virus X 41. Pitaya virus X 42. Plantago asiatica mosaic virus 43. Potato aucuba mosaic virus 44. Potato virus X 45. Schlumbergera virus X 46. Sclerotinia sclerotiorum 47. Senna mosaic virus 48. Shallot virus X 49. Strawberry mild yellow edge virus 50. Tamus red mosaic virus 51. Tulip virus X 52. Vanilla latent virus 53. Vanilla virus X 54. White clover mosaic virus 55. Yam virus X 56. Zygocactus virus X B - Betaflexiviridae 57. Aconitum latent virus 58. Actinidia virus B 59. African oil palm ringspot virus 60. Agave tequilana leaf virus 61. Pea streak virus: a. Alfalfa latent virus b. Pea streak virus 62. American hop latent virus 63. Apple chlorotic leaf spot virus 64. Apple green crinkle associated virus 65. Apple stem grooving virus 66. Apple stem pitting virus 67. Apricot latent virus 68. Apricot pseudo-chlorotic leaf spot virus 69. Apricot vein clearing associated virus 70. Arracacha virus V 71. Asian prunus virus 1 72. Asian prunus virus 73. Asian prunus virus 3 74. Banana mild mosaic virus 75. Blueberry scorch virus 76. Butterbur mosaic virus 77. Carrot Ch virus 1 78. Carrot Ch virus 2 79. Cherry green ring mottle virus 80. Cherry mottle leaf virus 81. Cherry necrotic rusty mottle virus 82. Cherry rusty mottle associated virus 83. Cherry twisted leaf associated virus 84. Cherry virus A 85. Chrysanthemum virus B 86. Citrus leaf blotch virus 87. Coleus vein necrosis virus 88. Cowpea mild mottle virus 89. Currant virus A 90. Daphne virus S 91. Diuris virus A 92. Diuris virus B 93. Elderberry carlavirus A 94. Elderberry carlavirus B 95. Elderberry carlavirus C 96. Elderberry carlavirus D 97. Elderberry carlavirus E 98. Gaillardia latent virus 99. Garlic common latent virus 100. Garlic latent virus 101. Grapevine Pinot gris virus 102. Grapevine berry inner necrosis virus 103. Grapevine rupestris stem pitting-associated virus 104. Grapevine virus A 105. Grapevine virus B 106. Grapevine virus E 107. Grapevine virus F 108. Grapevine virus I 109. Grapevine virus K 110. Grapevine virus T 111. Hardenbergia virus A 112. Helleborus net necrosis virus 113. Hippeastrum latent virus 2 114. Hop latent virus 115. Hop mosaic virus 116. Hydrangea chlorotic mottle virus 117. Jasmine virus C 118. Kalanchoe latent virus 119. Ligustrum necrotic ringspot virus 120. Ligustrum virus A 121. Lily symptomless virus 122. Mirabilis jalapa mottle virus 123. Narcissus common latent virus 124. Narcissus symptomless virus 125. Nerine latent virus 126. Ocimum basilicum RNA virus 1 127. Passiflora latent virus 128. Peach chlorotic mottle virus 129. Peach mosaic virus 130. Pepper virus A 131. Phlox virus B 132. Phlox virus S 133. Poplar mosaic virus 134. Potato latent virus 135. Potato virus H 136. Potato virus M 137. Potato virus P 138. Potato virus S 139. Potato virus T 140. Prunus virus T 141. Red clover vein mosaic virus 142. Rubus canadensis virus 1 143. Sugarcane striate mosaic-associated virus 144. Sweet potato C6 virus 145. Sweet potato chlorotic fleck virus 146. Watermelon virus A 147. Yacon virus A 148. Yam latent virus C - Gammaflexiviridae 149. Botrytis virus F D - Deltaflexiviridae 150. Fusarium graminearum deltaflexivirus 1 E - Tymoviridae 151. Anagyris vein yellowing virus 152. Andean potato latent virus 153. Andean potato mild mosaic virus 154. Asclepias asymptomatic virus 155. Bee Macula-like virus 156. Bombyx mori Macula-like virus 157. Chayote mosaic virus 158. Chiltepin yellow mosaic virus 159. Citrus sudden death-associated virus 160. Culex originated Tymoviridae-like virus 161. Diascia yellow mottle virus 162. Dulcamara mottle virus 163. Eggplant mosaic virus 164. Erysimum latent virus 165. Fig fleck-associated virus 166. Grapevine Red Globe virus 167. Grapevine Syrah virus 1 168. Grapevine asteroid mosaic-associated virus 169. Grapevine fleck virus 170. Grapevine rupestris vein feathering virus 171. Kennedya yellow mosaic virus 172. Maize rayado fino virus 173. Nectarine marafivirus M 174. Nemesia ring necrosis virus 175. Oat blue dwarf virus 176. Okra mosaic virus 177. Olive latent virus 3 178. Ononis yellow mosaic virus 179. Peach virus D 180. Physalis mottle virus 181. Plantago mottle virus 182. Poinsettia mosaic virus 183. Scrophularia mottle virus 184. Switchgrass mosaic virus 185. Tomato blistering mosaic virus 186. Turnip yellow mosaic virus 187. Varroa Tymo-like virus F – Flexiviridae non classificati 188. Rhizoctonia solani flexivirus 1 189. Sclerotinia sclerotiorum debilitation-associated RNA virus 2 G - Tymovirales non classificati 190. Bat tymo-like virus Organizzazione del genoma e replicazione La proteina più grande codificata è una poliproteina associata alla replicazione di circa 150-250 kDa vicino al terminale 5’ del genoma e che è tradotta direttamente dall’RNA genomico. La proteina contiene un insieme di domini funzionali, le cui sequenze e ordine di amminoacidi sono conservati in tutti i virus della superfamiglia simile agli alfavirus dei virus a RNA a filamento positivo. Un dominio di tipo 1 methyltransferase (Mtr) si trova vicino al terminale N e un dominio RNA- dipendente RNA polimerasi (RdRp) con un motivo centrale caratteristico S / TGx3Tx3NS / Tx22GDD si riscontra vicino al terminale C. Un dominio dell’elicasi RNA di superfamily 1 (Hel) è localizzato a monte del dominio RdRp. Mtr è coinvolto nell’incapsulamento dell’RNA, l’Hel nello svolgimento dell’RNA e RdRp nella sintesi dell’RNA. A monte dell’Hel in alcuni generi, esiste un dominio della proteasi della cisteina simile alla papaina (P-Pro) che elabora la poliproteina di replicazione. Immediatamente a monte della P-Pro, alcuni membri hanno una proteasi della famiglia del tumore ovarico (OTU).
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