【現(xiàn)學(xué)現(xiàn)賣】review真菌病毒

Abstract

植物病原真菌多能被病毒侵染物独，并且常發(fā)生多種病毒共同侵染一個真菌寄主的共侵染現(xiàn)象紧帕，但是多數(shù)情況下，這種共侵染是無癥狀的锣夹。有癥狀的侵染分為有害和有益于真菌兩種，有害發(fā)癥狀包括：造成低毒力真菌或真菌衰弱現(xiàn)象苏潜，比如已知的板栗疫惨肌（chestnut blight），白紋羽残糇蟆（white root rot）贴唇，油菜腐爛膊笮濉（rapeseed rot）病原菌中的一些病毒造成此類癥狀；有益的影響目前研究發(fā)現(xiàn)存在于在一些植物內(nèi)生病原真菌中戳气，比如病毒的存在加強了真菌的耐熱性链患，提高了真菌的致病力，甚至植物的耐熱能力也提升了瓶您。

病毒共侵染給研究病毒之間互作提供了平臺麻捻。

1. Introduction

根據(jù)國際病毒分類委員會（International Committee onTaxonomy of Viruses (ICTV)）2016年發(fā)布的數(shù)據(jù)，真菌病毒大致有12個家族呀袱，一些家族與植物贸毕，動物病毒相似度較高，而另外一些自成一派夜赵。高通量實驗使我們在今年來獲得了大量的病毒基因數(shù)據(jù)明棍，dsRNA和（+）ssRNA病毒為真菌病毒主要遺傳物質(zhì)類型。后來人們發(fā)現(xiàn)了ssDNA病毒（Genomoviridae）在真菌和環(huán)境中的存在(Yu et al., 2010; Krupovic et al., 2016)寇僧。2013年Kondo等人在白粉菌中發(fā)現(xiàn)了單股負鏈病毒（-）ssRNA病毒（mononegaviruses）可能的存在證據(jù)(Kondo et al., 2013)摊腋。Liu等人在真菌中有與動物病毒Nyamiviridae和Bornaviridae親緣關(guān)系較近的（-）ssRNA病毒(Liu et al., 2014)。

由于真菌病毒獨特的生活史嘁傀，即沒有胞外過程兴蒸，所以造就了真菌病毒的一些特點，比如沒有衣殼也可以進行侵染(Dawe and Nuss, 2013; Kanhayuwa et al., 2015; Zhanget al., 2016)心包。

一般真菌病毒的傳播途徑是通過孢子的豎直傳播类咧，或者通過菌絲融合的水平傳播，沒有胞外傳播途徑蟹腾。目前只發(fā)現(xiàn)了一個例外痕惋，一種genomovirus-DNA病毒Sclerotiniagemycirculavirus 1 (SGCV1)可以在培養(yǎng)基上入侵真菌菌絲，并且食菌性昆蟲也可以作為它的載體(Liu et al., 2016)娃殖。真菌物種間的營養(yǎng)不親和是非特異性的抗病毒手段（【現(xiàn)學(xué)現(xiàn)賣】綜述-真菌與病毒的周旋）值戳，當然實驗室里也試圖打破這個屏障，完成真菌病毒的傳播(Liu et al., 2003)炉爆。

人們利用酵母及其病毒體系堕虹，研究真菌中影響dsRNA病毒復(fù)制的因子(Wickner et al., 2013)。然而在絲狀真菌中的研究依然不多(Xie and Jiang, 2014)芬首，有人研究鐮刀菌的HEX1基因?qū)gV1病毒在真菌中積累的影響(Son et al., 2013)赴捞，Sun等人研究板栗疫病病毒和其病毒體系，發(fā)現(xiàn)一種病毒對其他病毒復(fù)制的影響(Sun and Suzuki, 2008)郁稍。

2. Types of Virus/Host Interactions

病毒對寄主真菌的影響有很多赦政，比如影響真菌轉(zhuǎn)錄組，小RNA組，蛋白質(zhì)組恢着，代謝組桐愉，脂質(zhì)組和表觀基因組。多數(shù)呈現(xiàn)無癥狀感染掰派，不過有的無癥狀感染的真菌在不同環(huán)境壓力下也可能出現(xiàn)各種癥狀从诲。并且病毒的侵染有的時候有害，有的時候有益靡羡，根據(jù)寄主的不同系洛，環(huán)境因素的不同而變化(Vainio and Hantula, 2016)。

2.1 Beneficial Interactions With HostFungi

目前最有趣的一個例子略步，是植物碎罚，真菌和病毒與耐熱能力的發(fā)現(xiàn)(Márquez et al., 2007)。有的會增強真菌的致病力(Ahn and Lee, 2001)纳像。

2.2 Harmful Interactions With Host Fungi

有害影響最典型的例子就是板栗疫病的弱毒病毒，降低致病力常常伴隨著真菌色素分泌拯勉，營養(yǎng)生長受抑制竟趾，有性孢子產(chǎn)量降低。最有名的病毒是CHV1宫峦，其他影響板栗疫病真菌致病力的還有CHV2, CHV3,?mycoreovirus1(MyRV1), MyRV2岔帽。Rosellinia necatrix megabirnavirus?1(RnMBV1)和MyRV3可以降低白紋羽病真菌的致病力。核盤菌Sclerotinia sclerotiorum中也發(fā)現(xiàn)了許多引起弱毒力的病毒导绷，如SsHADV1, SsNSRV1, SsHV2犀勒，SsPV1。

3. Types of Virus/Virus Interactions

多種病毒共侵染一個真菌是很常見的妥曲，但是病毒之間的作用至今不明贾费。

3.1 Synergistic Interactions BetweenViruses

兩個病毒共侵染時，引起的癥狀比單獨侵染嚴重檐盟。

3.2 Mutualistic Interactions BetweenViruses

有的病毒依靠另一個病毒完成復(fù)制褂萧，但是它并不是另一個病毒的衛(wèi)星病毒或者遺傳片段(Zhang et al., 2016; Hisano et al., 2018)。

3.3 Antagonistic Interactions BetweenViruses

一種病毒的存在抑制另一種病毒的復(fù)制葵萎。

3.4 Genome Rearrangements DuringCoinfection

板栗疫病被MyRV1和CHV1共侵染時發(fā)現(xiàn)兩個病毒之間的影響导犹，MyRV1的segment10常發(fā)生重排(Sun and Suzuki, 2008)。同時寄主的RNA沉默機制也會造成CHV1病毒基因組的重排(Sun et al., 2009)羡忘。重排還發(fā)生在自然和實驗室條件下的partitiviruses之間(Chiba et al., 2013, 2016)谎痢。

3.5 Implication of Coinfections byMultiple Mito- and Mito-Like Viruses

線粒體病毒特點是使用與線粒體適配的一套密碼子。線粒體病毒共侵染在真菌中很常見卷雕，目前多數(shù)線粒體病毒存在于絲狀真菌中节猿，有趣的是一些植物里也有與線粒體病毒同源的片段。

4. Conclusions and Prospects

真菌病毒種類很多爽蝴，許多病毒與寄主之間的互作沐批，病毒共侵染之間的互作需要更深入的研究纫骑。

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