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Genere Badnavirus Classificazione

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Genere Badnavirus Classificazione Dott. Francesco Fiume Genere Badnavirus Classificazione Gruppo IV (+ssRNA) Ordine Ortervirales Genere Badnavirus. Il genere Badnavirus, la cui specie tipo è Commelina yellow mottle virus (CYMV, comprende le seguenti specie: 1. Aglaonema bacilliform virus (ABV) 2. Banana streak GF virus (BSGFV) 3. Banana streak IM virus (BSIMV) 4. Banana streak MY virus (BSMYV) 5. Banana streak OL virus (BSOLV) 6. Banana streak UA virus (BSUAV) 7. Banana streak UI virus (BSUIV) 8. Banana streak UL virus (BSULV) 9. Banana streak UM virus (BSUMV) 10. Banana streak VN virus (BSVNV) 11. Bougainvillea chlorotic vein banding virus (BCVBV) 12. Cacao mild mosaic virus (CMMV) 13. Cacao swollen shoot CD virus (CSSCDV) 14. Cacao swollen shoot Togo A virus (CSSTAV) 15. Cacao swollen shoot virus (CSSV) 16. Cacao yellow vein banding virus (CYVBV) 17. Canna yellow mottle virus (CaYMV) 18. Citrus yellow mosaic virus (CYMV) 19. Commelina yellow mottle virus (CoYMV) 20. Dioscorea bacilliform AL virus (DBALV) 21. Dioscorea bacilliform RT virus 1 (DBRTV1) 22. Dioscorea bacilliform RT virus 2 (DBRTV2) 23. Dioscorea bacilliform SN virus (DBSNV) 24. Dioscorea bacilliform TR virus (DBTRV) 25. Fig badnavirus 1 (FBaV-1) 26. Gooseberry vein banding associated virus (GVBaV) 27. Grapevine Roditis leaf discoloration-associated virus (GRLDaV) 28. Grapevine vein clearing virus (GVCV) 29. Kalanchoe top-spotting virus (KTSV) 30. Mulberry badnavirus 1 (MBV-1) 31. Pagoda yellow mosaic associated virus (PYMAV) 32. Pineapple bacilliform CO virus (PBCOV) 33. Pineapple bacilliform ER virus (PBERV) 34. Piper yellow mottle virus (PYMV) 35. Rubus yellow net virus (RYNV) 36. Schefflera ringspot virus (SRV) 37. Spiraea yellow leafspot virus (SYLSV) 38. Sugarcane bacilliform Guadeloupe A virus (SCBGAV) 39. Sugarcane bacilliform Guadeloupe D virus (SCBGDV) 40. Sugarcane bacilliform IM virus (SBIMV) 41. Sugarcane bacilliform MO virus (SBMOV) 42. Sweet potato pakakuy virus (SPPV) 43. Taro bacilliform CH virus (TBCHV) 44. Taro bacilliform virus (TBV) 45. Wisteria badnavirus 1 (WBV1) 46. Yacon necrotic mottle virus (YNMoV). Le piante sono gli ospiti naturali di questo virus. Ci sono attualmente 46 specie in questo genere le cui malattie associate includono i seguenti sintomi: clorosi fogliare, necrosi delle radici, benda delle vene rosse in foglie giovani, piccoli baccelli chiazzati, gonfiori nella radice e nel fusto cui può seguire la morte delle piante. L'infezione riduce la resa del 25% entro un anno, del 50% entro due anni e di solito le piante muoiono entro 3-4 anni. Distribuzione geografica Il virus è presente nella sottoregione dell'Africa occidentale nei paesi della Costa d’Avorio, Ghana, Togo e Nigeria (figura 1). Figura 1 – Distribuzione geografica dei Badnavirus. Struttura del virus e ciclo vitale I Badnavirus sono privi di pericapside, con geometrie bacilliformi. Il genoma è circolare con segmentazione monopartitica. Le particelle virali hanno una larghezza di circa 30 nm e una lunghezza di 90-900 nm (figura 2). 2 Figura 2 - Particelle del genere Badnavirus, senza pericapside, bacilliformi, di 30 nm di larghezza e 90-900 nm di lunghezza. I genomi sono circolari e non segmentati (figura 3). La replicazione virale è nucleare/citoplasmatica. L'ingresso nella cellula ospite si ottiene con l’attacco delle proteine virali ai recettori ospiti, che mediano l'endocitosi. La replica è quella del modello di replica del dsDNA (RT, reverse transcriptasi). La trascrizione basata sul DNA, in particolare la trascrizione dsdna (rt) è il metodo di trascrizione. Il virus esce dalla cellula ospite mediante l'esportazione di pori nucleari ed il movimento virale guidato dai tubuli. Le piante fungono da ospiti naturali. Il virus viene trasmesso tramite un vettore rappresentato da varie specie di cocciniglie, anche se, in generale, i percorsi della trasmissione sono vettoriali, meccanici ed attraverso i semi. Figura 3 – Genoma dei Badnavirus, monopartito, a DNA circolare aperto a doppio filamento, di circa 7200 paia di basi, con discontinuità in entrambi i filamenti: uno nel filamento trascritto e tre in quello non trascritto. La poliproteina ORF1 è presumibilmente tagliata in diverse catene da proteasi virali. La replicazione virale avviene secondo le seguenti fasi: 1. L’attacco delle proteine virali ai recettori ospiti media l’entrata virale nella cellula ospite. 3 2. Il dsDNA virale viene rilasciato nel nucleo dove viene trascritto dalla polimerasi II dell’RNA dell’ospite. 3. La traduzione dell'RNA produce proteine virali. 4. L'RNA genomico è retrotrascritto nel citoplasma in nuovi genomi di dsDNA. 5. I genomi sono incapsidati con la proteina capsidica, rilasciando nuovi virioni. Organizzazione del genoma e ciclo di replicazione Nella figura 4 è riportato il ciclo di Cauliflower mosaic virus (CaMV), un badnavirus il cui genoma è lungo da 7,2 a 9,2 kb volte quello do in DNA circolare con sovrapposizione del singolo filamento. Questi sono i segni dell'inizio della sintesi del DNA positivo e negativo per la trascrizione inversa (Hohn et Rothnie, 2013 ). La trascrizione inversa si riscontra all'interno del capside del virus con il risultato che il DNA viene trasferito al nucleo, dove le discontinuità vengono riparate ed i mini- cromosomi sono formati dall'associazione del DNA supercoltivato chiuso covalentemente con le proteine istoniche (Hull, 2002). Figura 4 - Ciclo di replicazione dei pararetrovirus delle piante (partenza in senso orario a 12 ore): una particella virale contenente DNA circolare aperto è entrato nella cellula. Il suo DNA è importato nel nucleo, dove gli spazi vuoti vengono riparati per produrre dsDNA superarrotolato. Viene prodotto RNA terminale, ridondante e trasportato nel citoplasma. Le copie dell'RNA sono tradotte in proteine virali e/o impaccate come due copie in particelle virali. Avviene la trascrizione inversa. Il dsDNA circolare aperto risultante rientra nel ciclo di replica. In basso a sinistra viene mostrato come la ricombinazione può avvenire per modello commutazione. A destra viene mostrato quanto raramente i frammenti del DNA del virus incorporino nella pianta genoma, che porta a sequenze virali endogene. 4 Una regione intergenomica lunga da 631 a 1177 pb (figura 5) include un promotore ed un segnale di poliadenilazione. Figura 5 - Organizzazione del genoma di badnavirus rappresentativi con un numero variabile di ORF. Banana streak MY virus (BSMYV); Bougainvillea spectabilis chlorotic vein banding virus (BsCVBV); Cacao swollen shoot virus (CSSV); Citrus yellow mosaic virus (CYMV), Rubus yellow net virus (RYNV). Il DNA mini-cromosomico viene quindi trascritto utilizzando l'RNA dipendente dalla polimerasi II del DNA codificato dall'ospite, creando un RNA terminale ridondante che funge da pregenoma e messaggero policistronico del RNA. La ridondanza viene creata ignorando il segnale di poliadenilazione sul DNA circolare, durante il primo incontro con la RNA polimerasi e riconoscendolo al secondo (Sanfaçon et Hohn, 1990). Nella fase finale della replicazione, l'RNA pregenomico viene riconvertito in dsDNA mediante l'azione di una trascrittasi inversa. La sintesi del DNA a singolo filamento, senso (-), è innescata da tRNAmet e di senso (+) del DNA da prodotti di clivaggio ricchi in purine rimasti dopo la digestione con RNasi H del pregenomico RNA (Medberry et al., 1990; Hohn, et Rothnie, 2013). L'ORF I dei badnavirus contiene da 399 a 927 bp a seconda della specie. Nel membro del tipo 5 ComYMV, ORF I è lungo 602 bp e si traduce in un polipeptide di 23 kD, che si è dimostrato essere (Cheng et al., 1996) associato al virione (virion-associated). L'ORF II è il più piccolo, che va da 312 a 561 bp. In CSSV, il suo prodotto è stato identificato come proteina legante l'acido nucleico (Jacquot et al 1996); ComYMV, è stato anche mostrato di legarsi al capside del virus (Medberry et al., 1990; Cheng et al., 1996). ORF III è il più grande ORF, di lunghezza compresa tra 5100 e 6000 bp. Codifica una poliproteina che viene scissa in quattro o cinque prodotti quali: 1. proteina del capside, 2. aspartato proteasi, 3. transcriptasi inversa, 4. RNasi H 5. una presunta proteina di movimento cellula-cellula (Medberry et al., 1990; Jacquot et al., 1996; Hohn et Fütterer, 1997). La scissione di questa poliproteina nelle sue subunità funzionali è raggiunta dalla proteasi aspartica (Hohn et Fütterer, 1997). La proteina del capside include un motivo Cys, come nella maggior parte degli elementi di trascrizione inversa, ed un secondo motivo Cys (CX2CX11CX2CX4CX2C) trovato solo nei membri dei generi. Alcuni badnavirus hanno ORF aggiuntivi: Piper yellow mottle virus (PYMoV), Bougainvillea spectabilis chlorotic vein-banding virus (BsCVBV), Dioscorea bacilliform SN virus (DBSNV), Grapevine Roditis leaf discoloration-associated virus (GRLDaV), Taro bacilliform virus (TaBV) e Yacon necrotic mottle virus (YNMoV) hanno un ORF aggiuntivo (Hany et al., 2014; Lee et al., 2015; Maliogka et al., 2015; Seal et al., 2007; Wang et al., 2016; Yang et al., 2003). Pagoda yellow mosaic associated virus (PYMAV) ne ha due (Hagen et al., 1993; Wang et al., 2014); Cymbidium mosaic virus (CYMV) e Taro bacilliform CH virus (TaBCHV) ne hanno tre (Borah et al., 2009; Huang et al., 2001; Kazmi et al., 2015); DrMV e RYNV hanno quattro ORF aggiuntivi. Due di questi ultimi sono posizionati sul filamento antisenso (Su et al., 2007; Kalischukng et al., 2013). Non è noto se questi ORF siano espressi, né quale potrebbe essere la loro funzione. Le trascrizioni dei pararetrovirus delle piante hanno una lunghezza di circa 600 nt, sono altamente strutturate, contengono diversi ORF corti, con la possibilità di formare una grande struttura, stem-loop (conformazione intramolecolare basata sull'accoppiamento delle basi azotate che può insorgere a livello di un DNA a singolo filamento) nota per essere la traduzione transitoria. La formazione di questa struttura porta il primo ORF lungo nella stretta vicinanza spaziale di un ORF prossimale 5' corto che termina a 5-10 nt a monte dell’elemento strutturale stabile. Questo elemento viene deviato dal ribosoma di scansione e porta alla traduzione del primo lungo ORF (Fütterer et al., 1993; Ryabova et Hohn, 2000).
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