葡萄孢属植物病原菌真菌病毒研究
摘要
葡萄孢属真菌(Botrytis spp)是一种广泛分布的植物病原真菌,可引起多种作物的灰霉病,造成严重的经济损失。本研究对源于灰葡萄孢(B. cinerea)、大蒜盲种葡萄孢(B. porri)、葱鳞葡萄孢(B.squamosa)、中国葱葡萄孢(B. sinoallii)及拟蚕豆葡萄孢(B.fabiopsis)中的5类真菌病毒进行了研究,具体包括两种线粒体病毒(Mitovirus)、一种双分病毒(Partitivirus)、一种尚未分类的病毒及一个代表新病毒科的成员。在上述5种病毒中,有两种病毒已被证实与葡萄孢的致病力衰退有密切的关系,其中包括源于灰葡萄孢菌株CanBc-1的线粒体病毒Botrytis cinerea mitovirus1(BcMV1)和源于大蒜盲种葡萄孢菌株GarlicBc-72的真菌病毒Botrytis porri RNA virus1(BpRV1)。
BcMV1的全长约为2804nt,与OnuMV3b有极高的同源性,可以通过病菌的分生孢子进行垂直传染和通过菌丝的融合进行水平传染。成功传染BcMV1dsRNA的菌株均会在生长速度和致病力上出现明显的衰退。通过透射电镜对病毒感染后菌丝的超薄切片进行观察可以发现,病毒侵染后,菌丝细胞中线粒体会出现肿大、嵴退化和内部纤维化的症状,加之提纯的线粒体中也可检测到BcMV1的dsRNA,因此,BcMV1的攻击位点被认为是线粒体。BcMV1侵染线粒体后,造成线粒体的超微结构出现明显的异常,从而可能造成线粒体功能的紊乱,并最终导致灰葡萄孢致病力的严重衰退。此外,BcMV1在灰葡萄孢分生孢子形成过程中可以产生DIRNA,即BcMV1-S。BcMV1-S可以通过菌丝融合进行水平传染。虽然它可以抑制BcMV1的复制但不会影响BcMV1造成的弱毒特性。
BpRVl是本研究中另一种可引起葡萄孢致病力衰退的真菌病毒。BpRVl的基因组含有两条dsRNA片段,分别命名为dsRNA-1(6125bp)和dsRNA-2(5879bp)。这两条片段在5’和3’末端分别有95%和62%的相似度,且各编码一个大的开放阅读框(ORF),分别命名为ORFⅠ (dsRNA-1)和ORFⅡ (dsRNA-2)。ORFI所编码的蛋白在靠近N端的区域与Totiviridae、Chrysoviridae和Megabirnaviridae科病毒所编码的依赖于RNA的RNA聚合酶(RdRp)有一定的同源性,且同源性约为19%-23%。ORFⅠ所编码蛋白的其他区域及ORFⅡ所编码蛋白并未在数据库中搜索到与之有明显同源性的蛋白序列。系统进化分析表明,BpRV1属于一个单独的分支,与已报道真菌病毒的亲缘关系均较远。BpRV1的病毒粒体直径约为35nm,里面包含有两条dsRNA链(dsRNA-1和dsRNA-2)和大小分别为70kDa、80kDa及85kDa的三种病毒结构蛋白。蛋白质指纹图谱分析表明80kDa及85kDa的结构蛋白由dsRNA-1编码,而70kDa的结构蛋白由dsRNA-2编码。将BpRV1的病毒粒体导入强毒菌株GarlicBc-38后,会导致其病毒感染后的衍生菌株38T致病力和生长速度的衰退。以上结果都表明,BpRV1是一种新型的真菌病毒,并可能属于一个新的病毒科。此外,在葱鳞葡萄孢菌株LeekBc-10中也检测到了一种真菌病毒,Botrytis squamosa RNA virus1(BsRV1)。BsRV1与BpRV1的两条dsRNA基因组在核酸水平上的同源性为91%-97%,而在蛋白水平上的同源性则高达97%-98%,因此BpRV1与BsRV1被认为是同一种病毒的不同毒株。
剩余的3种真菌病毒对寄主的影响尚不十分明确,但病毒侵染的菌株都表现出了与正常菌株明显不同的培养形态。其中,已获得了来源于葱鳞葡萄孢菌株GarlicBc-21的Botrytis squamosa RNA virus2(BsRV2/Bc-21)和来源于拟蚕豆葡萄孢菌株BroadbeanBc-41的Botrytis fabiopsis RNA virus1(BfRV1/Bc-41)部分基因组序列。结果显示两者在蛋白水平上有91%以上的相似性,因而BsRV2/Bc-21与BfRV1/Bc-41可能也是同一种病毒。系统进化分析表明这两种病毒与FgV-DK21的亲缘关系最近,且与hypovirus有一定的亲缘关系,但具体的分类地位还尚待确定。另外在菌株BroadbeanBc-41中还发现了一种双分病毒Botrytis fabiopsis partitivirus1(BfPV1)与SsPVS有较近的亲缘关系。此外,BfPV1还与蚕豆上的一种双分病毒VfPV-1有较高的同源性。最后,在中国葱葡萄孢菌株OnionBc-59中发现了一种与SsMV1有较近的亲缘关系的线粒体病毒Botrytis sinoallii mitovirus1(BsMV1)。
Botrytis spp is a widely spread group of plant pathogenic fungi, which causes gray mold disease of many crops and leads serious economic losses. This research focused on the characterization of five mycoviruses originated from B. cinerea, B. porri, B. squamosa, B. sinoallii and B.fabiopsis, of which two belong to Mitovirus, one belongs to Partitivirus, one is from an unassigned viral group and one that may represent a novel viral family. Two mycoviruses from those five viruses were determined to be associated with the hypovirulence of Botrytis spp, including Botrytis cinerea mitovirus1(BcMV1) from strain CanBc-1of B. cinerea and Botrytis porri RNA virus1(BpRV1) from strain GarlicBc-72of B. porri.
The full length sequence of BcMV1is2084nt, which shows high sequence identity to OnuMV3b. BcMV1could be vertically and horizontally transmitted though conidia and by hyphal anastomosis. Infection of BcMV1dsRNA causes the serious debilitation of virulence and growth for virus-infected strains. TEM observation indicates the infection of BcMV1causes the swollen mitochondria, degeneration of cristae and presence of fibrous material in fungal mitochondria. In addition, BcMV1could be detected from purified mitochondria. Therefore, mitochondria are believed to be the target for BcMV1. The infection of mitochondria by BcMV1causes the unusual subcellular structure of mitochondria, which may lead the malfunction of mitochondria and causes the hypovirulence of B. cinerea. Moreover, BcMV1could generate defective interference (DI) RNA during the sporulation process of B. cinerea, designated as BcMV1-S. BcMV1-S could be horizontally transmitted. Although it inhibits the replication of BcMV1, but it does not affect the hypovirulence caused by BcMV1.
BpRV1is determined to be another mycovirus conferring hypovirulence to Botrytis spp in this study. The BpRV1genome comprises two dsRNAs, dsRNA-1(6,215bp) and dsRNA-2(5,879bp), which share sequence identities of62and95%at the3'-and5'-terminal regions, respectively. Two open reading frames (ORFs), ORF I (dsRNA-1) and ORF Ⅱ (dsRNA-2), were detected. The protein encoded by the3'-proximal coding region of ORF I shows sequence identities of19to23%with RNA-dependent RNA polymerases encoded by viruses in the families Totiviridae, Chrysoviridae, and Megabirnaviridae. However, the proteins encoded by the5'-proximal coding region of ORF I and by the entire ORF II lack sequence similarities to any reported virus proteins. Phylogenetic analysis showed that BpRV1belongs to a separate clade distinct from those of other known RNA mycoviruses. Purified virions of-35nm in diameter encompass dsRNA-1and dsRNA-2, and three structural proteins (SPs) of70,80, and85kDa, respectively. Peptide mass fingerprinting analysis revealed that the80-and85-kDa SPs are encoded by ORF I, while the70-kDa SP is encoded by ORF II. Introducing BpRV1purified virions into the virulent strain GarlicBc-38of B. porri caused derivative38T reduced mycelial growth and hypovirulence. These combined results suggest that BpRV1is a novel bipartite dsRNA virus that possibly belongs to a new virus family. Moreover, Botrytis squamosa RNA virus1(BsRV1) was isolated from strain LeekBc-10of B. squamosa. BsRV1and BpRV1shows highly sequence identity with91%-97%and97%-98%for nucleotide and protein sequence respectively, thus the two viruses is believed to be two strains of one viral species.
The effects to host fungi were unknown for the rest three mycoviruses, although the virus infected strains showed abnormal cultural morphology. Partial genome sequence of Botrytis squamosa RNA virus2(BsRV2/Bc-21) and Botrytis fabiopsis RNA virus1(BfRV1/Bc-41) were obtained. The two viruses were isolated from strain GarlicBc-21of B. sqaumosa and strain BroadbeanBc-41of B. fabiopsis respectively. BsRV2/Bc-21and BfRV1/Bc-41are believed to be one viral species, as they showed high protein sequence identity (91%) with each other. Phylogenetic analysis indicated the two viruses were close related to FgV-DK21, and grouped with hypo viruses. However, the classification of this virus remains unknown. Moreover, another partitivirus, Botrytis fabiopsis partitivirus1(BfPV1), was also identified from strain BroadbeanBc-41, which is close related to SsPVS. Moreover, BfPV1also showed homology to a plant partitivirus of Vicia faba, VfPV-1. In addition, a mitovirus with close relationship to SsMV1was isolated from strain OnionBc-59of B. sinoallii, and named as Botrytis sinoallii mitovirus1(BsMV1).
BcMV1的全长约为2804nt,与OnuMV3b有极高的同源性,可以通过病菌的分生孢子进行垂直传染和通过菌丝的融合进行水平传染。成功传染BcMV1dsRNA的菌株均会在生长速度和致病力上出现明显的衰退。通过透射电镜对病毒感染后菌丝的超薄切片进行观察可以发现,病毒侵染后,菌丝细胞中线粒体会出现肿大、嵴退化和内部纤维化的症状,加之提纯的线粒体中也可检测到BcMV1的dsRNA,因此,BcMV1的攻击位点被认为是线粒体。BcMV1侵染线粒体后,造成线粒体的超微结构出现明显的异常,从而可能造成线粒体功能的紊乱,并最终导致灰葡萄孢致病力的严重衰退。此外,BcMV1在灰葡萄孢分生孢子形成过程中可以产生DIRNA,即BcMV1-S。BcMV1-S可以通过菌丝融合进行水平传染。虽然它可以抑制BcMV1的复制但不会影响BcMV1造成的弱毒特性。
BpRVl是本研究中另一种可引起葡萄孢致病力衰退的真菌病毒。BpRVl的基因组含有两条dsRNA片段,分别命名为dsRNA-1(6125bp)和dsRNA-2(5879bp)。这两条片段在5’和3’末端分别有95%和62%的相似度,且各编码一个大的开放阅读框(ORF),分别命名为ORFⅠ (dsRNA-1)和ORFⅡ (dsRNA-2)。ORFI所编码的蛋白在靠近N端的区域与Totiviridae、Chrysoviridae和Megabirnaviridae科病毒所编码的依赖于RNA的RNA聚合酶(RdRp)有一定的同源性,且同源性约为19%-23%。ORFⅠ所编码蛋白的其他区域及ORFⅡ所编码蛋白并未在数据库中搜索到与之有明显同源性的蛋白序列。系统进化分析表明,BpRV1属于一个单独的分支,与已报道真菌病毒的亲缘关系均较远。BpRV1的病毒粒体直径约为35nm,里面包含有两条dsRNA链(dsRNA-1和dsRNA-2)和大小分别为70kDa、80kDa及85kDa的三种病毒结构蛋白。蛋白质指纹图谱分析表明80kDa及85kDa的结构蛋白由dsRNA-1编码,而70kDa的结构蛋白由dsRNA-2编码。将BpRV1的病毒粒体导入强毒菌株GarlicBc-38后,会导致其病毒感染后的衍生菌株38T致病力和生长速度的衰退。以上结果都表明,BpRV1是一种新型的真菌病毒,并可能属于一个新的病毒科。此外,在葱鳞葡萄孢菌株LeekBc-10中也检测到了一种真菌病毒,Botrytis squamosa RNA virus1(BsRV1)。BsRV1与BpRV1的两条dsRNA基因组在核酸水平上的同源性为91%-97%,而在蛋白水平上的同源性则高达97%-98%,因此BpRV1与BsRV1被认为是同一种病毒的不同毒株。
剩余的3种真菌病毒对寄主的影响尚不十分明确,但病毒侵染的菌株都表现出了与正常菌株明显不同的培养形态。其中,已获得了来源于葱鳞葡萄孢菌株GarlicBc-21的Botrytis squamosa RNA virus2(BsRV2/Bc-21)和来源于拟蚕豆葡萄孢菌株BroadbeanBc-41的Botrytis fabiopsis RNA virus1(BfRV1/Bc-41)部分基因组序列。结果显示两者在蛋白水平上有91%以上的相似性,因而BsRV2/Bc-21与BfRV1/Bc-41可能也是同一种病毒。系统进化分析表明这两种病毒与FgV-DK21的亲缘关系最近,且与hypovirus有一定的亲缘关系,但具体的分类地位还尚待确定。另外在菌株BroadbeanBc-41中还发现了一种双分病毒Botrytis fabiopsis partitivirus1(BfPV1)与SsPVS有较近的亲缘关系。此外,BfPV1还与蚕豆上的一种双分病毒VfPV-1有较高的同源性。最后,在中国葱葡萄孢菌株OnionBc-59中发现了一种与SsMV1有较近的亲缘关系的线粒体病毒Botrytis sinoallii mitovirus1(BsMV1)。
Botrytis spp is a widely spread group of plant pathogenic fungi, which causes gray mold disease of many crops and leads serious economic losses. This research focused on the characterization of five mycoviruses originated from B. cinerea, B. porri, B. squamosa, B. sinoallii and B.fabiopsis, of which two belong to Mitovirus, one belongs to Partitivirus, one is from an unassigned viral group and one that may represent a novel viral family. Two mycoviruses from those five viruses were determined to be associated with the hypovirulence of Botrytis spp, including Botrytis cinerea mitovirus1(BcMV1) from strain CanBc-1of B. cinerea and Botrytis porri RNA virus1(BpRV1) from strain GarlicBc-72of B. porri.
The full length sequence of BcMV1is2084nt, which shows high sequence identity to OnuMV3b. BcMV1could be vertically and horizontally transmitted though conidia and by hyphal anastomosis. Infection of BcMV1dsRNA causes the serious debilitation of virulence and growth for virus-infected strains. TEM observation indicates the infection of BcMV1causes the swollen mitochondria, degeneration of cristae and presence of fibrous material in fungal mitochondria. In addition, BcMV1could be detected from purified mitochondria. Therefore, mitochondria are believed to be the target for BcMV1. The infection of mitochondria by BcMV1causes the unusual subcellular structure of mitochondria, which may lead the malfunction of mitochondria and causes the hypovirulence of B. cinerea. Moreover, BcMV1could generate defective interference (DI) RNA during the sporulation process of B. cinerea, designated as BcMV1-S. BcMV1-S could be horizontally transmitted. Although it inhibits the replication of BcMV1, but it does not affect the hypovirulence caused by BcMV1.
BpRV1is determined to be another mycovirus conferring hypovirulence to Botrytis spp in this study. The BpRV1genome comprises two dsRNAs, dsRNA-1(6,215bp) and dsRNA-2(5,879bp), which share sequence identities of62and95%at the3'-and5'-terminal regions, respectively. Two open reading frames (ORFs), ORF I (dsRNA-1) and ORF Ⅱ (dsRNA-2), were detected. The protein encoded by the3'-proximal coding region of ORF I shows sequence identities of19to23%with RNA-dependent RNA polymerases encoded by viruses in the families Totiviridae, Chrysoviridae, and Megabirnaviridae. However, the proteins encoded by the5'-proximal coding region of ORF I and by the entire ORF II lack sequence similarities to any reported virus proteins. Phylogenetic analysis showed that BpRV1belongs to a separate clade distinct from those of other known RNA mycoviruses. Purified virions of-35nm in diameter encompass dsRNA-1and dsRNA-2, and three structural proteins (SPs) of70,80, and85kDa, respectively. Peptide mass fingerprinting analysis revealed that the80-and85-kDa SPs are encoded by ORF I, while the70-kDa SP is encoded by ORF II. Introducing BpRV1purified virions into the virulent strain GarlicBc-38of B. porri caused derivative38T reduced mycelial growth and hypovirulence. These combined results suggest that BpRV1is a novel bipartite dsRNA virus that possibly belongs to a new virus family. Moreover, Botrytis squamosa RNA virus1(BsRV1) was isolated from strain LeekBc-10of B. squamosa. BsRV1and BpRV1shows highly sequence identity with91%-97%and97%-98%for nucleotide and protein sequence respectively, thus the two viruses is believed to be two strains of one viral species.
The effects to host fungi were unknown for the rest three mycoviruses, although the virus infected strains showed abnormal cultural morphology. Partial genome sequence of Botrytis squamosa RNA virus2(BsRV2/Bc-21) and Botrytis fabiopsis RNA virus1(BfRV1/Bc-41) were obtained. The two viruses were isolated from strain GarlicBc-21of B. sqaumosa and strain BroadbeanBc-41of B. fabiopsis respectively. BsRV2/Bc-21and BfRV1/Bc-41are believed to be one viral species, as they showed high protein sequence identity (91%) with each other. Phylogenetic analysis indicated the two viruses were close related to FgV-DK21, and grouped with hypo viruses. However, the classification of this virus remains unknown. Moreover, another partitivirus, Botrytis fabiopsis partitivirus1(BfPV1), was also identified from strain BroadbeanBc-41, which is close related to SsPVS. Moreover, BfPV1also showed homology to a plant partitivirus of Vicia faba, VfPV-1. In addition, a mitovirus with close relationship to SsMV1was isolated from strain OnionBc-59of B. sinoallii, and named as Botrytis sinoallii mitovirus1(BsMV1).
引文
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