小西葫芦黄花叶病毒山东南瓜和丝瓜分离物全基因组序列分析
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摘要
在山东泰安采集到两个表现黄花叶症状的丝瓜和南瓜样品中检测到小西葫芦黄花叶病毒(Zucchini yellow mosaic virus,ZYMV);通过RT-PCR和RACE方法获得了两个分离物的全基因组序列,分别命名为CN:Cm:17(丝瓜分离物)和CN:Lc:17(南瓜分离物)。结果表明,CN:Cm:17和CN:Lc:17基因组除Poly(A)尾巴外,分别含有9 593和9 591个核苷酸(Nucleotides,nt),通过多聚蛋白策略编码一个含3080个氨基酸的多聚蛋白。CN:Cm:17和CN:Lc:17在氨基酸水平上一致率为96.7%;CN:Cm:17与韩国分离物KR:RDA:07的氨基酸一致率最高(97.8%); CN:Lc:17与CN:CJLX30535:14和CN:spider131932:13的一致率最高(98.4%)。重组分析发现,CN:Lc:17具有显著重组事件,为KR:KR-PA:05和CN:Cm:17的重组体。基于多聚蛋白编码序列的系统进化聚类结果显示分组与分离物的地理起源存在密切相关性,所有中国分离物分别聚集在A-Ⅴ和A-Ⅵ亚组。
Zucchini yellow mosaic virus(ZYMV)is a major virus infecting cucurbit crops. ZYMV was detected in leaf samples of loofah and pumpkin displaying symptoms of yellowing and mosaic collected from Tai'an,Shandong Province. Their complete genome sequences were determined using RT-PCR and 5′ RACE,the resulting complete genomes were named as CN:Cm:17 and CN:Lc:17,respectively. Sequencing results showed that the complete genome excluding poly(A) tail of CN:Cm:17 and CN:Lc:17 was 9 593 and 9 591 nucleotides(nt),respectively,encoding a polyprotein of 3 080 amino acids. Amino acid sequence identity between CN:Cm:17 and CN:Lc:17 was 96.7%. Isolate CN:Cm:17 shared the highest sequence identity with KR:RDA:07 at amino acid level,CN:Lc:17 showed the highest sequence identity with CN:CJLX30535:14 and CN:spider131932:13. Recombinant analysis revealed that CN:Lc:17 was recombinant of isolates of KR:KR-PA:05 and CN:Cm:17. Phylogenetic analysis based on polyprotein-encoding sequences suggested a high correlation between phylo-groups and their geographical origins. All Chinese isolates were grouped in A-Ⅴ and A-Ⅵ,respectively.
Zucchini yellow mosaic virus(ZYMV)is a major virus infecting cucurbit crops. ZYMV was detected in leaf samples of loofah and pumpkin displaying symptoms of yellowing and mosaic collected from Tai'an,Shandong Province. Their complete genome sequences were determined using RT-PCR and 5′ RACE,the resulting complete genomes were named as CN:Cm:17 and CN:Lc:17,respectively. Sequencing results showed that the complete genome excluding poly(A) tail of CN:Cm:17 and CN:Lc:17 was 9 593 and 9 591 nucleotides(nt),respectively,encoding a polyprotein of 3 080 amino acids. Amino acid sequence identity between CN:Cm:17 and CN:Lc:17 was 96.7%. Isolate CN:Cm:17 shared the highest sequence identity with KR:RDA:07 at amino acid level,CN:Lc:17 showed the highest sequence identity with CN:CJLX30535:14 and CN:spider131932:13. Recombinant analysis revealed that CN:Lc:17 was recombinant of isolates of KR:KR-PA:05 and CN:Cm:17. Phylogenetic analysis based on polyprotein-encoding sequences suggested a high correlation between phylo-groups and their geographical origins. All Chinese isolates were grouped in A-Ⅴ and A-Ⅵ,respectively.
引文
Adams M J,Antoniw J F,Fauquet C M.2005.Molecular criteria for genus and species discrimination within the family Potyviridae.Archives of Virology,150(3):459-79.
Bananej K,Keshavarz T,Vahdat A,Salekdeh G H,Glasa M.2010.Biological and Molecular Variability of Zucchini yellow mosaic virus in Iran.Journal of Phytopathology,156:654-659.
Chare E R,Holmes E C.2006.A phylogenetic survey of recombination frequency in plant RNA viruses.Archives of Virology,151:933.
Chen Jie-yun,Chen Ji-shuang,Hong Jian.2003.Sequence diversity of 3′end genome for Zucchini yellow mosaic virus.Acta Microbiology Sinica,43(3):388-394.(in Chinese)陈洁云,陈集双,洪健.2003.小西葫芦黄化花叶病毒分离物的3′末端序列多态性研究.微生物学报,43(3):388-394.
Chen Y K,Hong Y H.2008.First report of Begonia chlorotic ringspot caused by Zucchini yellow mosaic virus in Taiwan.Plant Disease,92(8):1247.
Chung Y W,Miller W A,Atkins J F,Firth A E.2008.An overlapping essential gene in the Potyviridae.Proceedings of the National Academy of Sciences of the United States of America,105(15):5897.
Cordero T,Cerdán L,Carbonell A,Katsarou K,Kalantidis K,Daròs J A.2016.Dicer-like 4 is involved in restricting the systemic movement of Zucchini yellow mosaic virus in Nicotiana benthamiana.Molecular Plant-Microbe Interactions,MPMI:MPMI11160239R.
Coutts B A,Kehoe M A,Webster C G,Wylie S J,Jones R A.2011.Zucchini yellow mosaic virus:biological properties,detection procedures and comparison of coat protein gene sequences.Archives of Virology,156(12):2119-2131.
Davis R F.1986.Partial characterization of Zucchini yellow mosaic virus isolated from squash in Turkey.Plant Disease,70(8):735-738.
Davis R F,Mizuki M K.1987.Detection of cucurbit viruses in New Jersey.Plant Disease,22(71):199-202.
Desbiez C,Gal-On A,Girard M,Wipf-Scheibel C,Lecoq H.2003.Increase in Zucchini yellow mosaic virus symptom severity in tolerant zucchini cultivars is related to a point mutation in P3 protein and is associated with a loss of relative fitness on susceptible plants.Phytopathology,93:1478-1484.
Desbiez C,Wipf-Scheibel C,Lecoq H.2002.Biological and serological variability,evolution and molecular epidemiology of Zucchini yellow mosaic virus(ZYMV,Potyvirus)with special reference to Caribbean islands.Virus Research,85:5-16.
Dunham J P,Simmons H E,Holmes E C,Stephenson A G.2014.Analysis of viral(Zucchini yellow mosaic virus)genetic diversity during systemic movement through a C.pepo vine.Virus Research,191(1):172-179.
Gal-On A,Antignus Y,Rosner A,Raccah B.1990.Nucleotide sequence of the Zucchini yellow mosaic virus capsid-encoding gene and its expression in Escherichia coli.Gene,87:273-277.
Glasa M,PittnerováS.2010.Complete genome sequence of a slovak isolate of Zucchini yellow mosaic virus(ZYMV)provides further evidence of a close molecular relationship among central European ZYMV isolates.Journal of Phytopathology,154(7-8):436-440.
Green K J,Brown C J,Gray S M,Karasev A V.2017.Phylogenetic study of recombinant strains of Potato virus Y:Virology,507:40-52.
Green K J,Brown C J,Karasev A V.2018.Genetic diversity of Potato virus Y(PVY):sequence analyses reveal ten novel PVY recombinant structures.Arch Virol,163:23-32.
Gu Qin-sheng,Fan Zai-feng,Piero Rogger,Chen Hong-yun,Yu Zheng-wang,Li Huai-fang.2003.Cloning and sequence analysis of the coat protein gene of Zucchini yellow mosaic virus.Acta Phytopathologica Sinica,33(6):498-502.(in Chinese)古勤生,范在丰,Piero Rogger,陈红运,俞正旺,李怀方.2003.小西葫芦黄花叶病毒外壳蛋白基因的克隆及序列分析.植物病理学报,33(6):498-502.
Fu Jing-jing,Yang Cai-xia,Han Tong,Liao Yi-ming,Li Liang,Li Meng-xin,Sun Wei,Wang Shuo.2017.Identification and sequence analysis of Zucchini yellow mosaic virus in Liaoning.Journal of Fujian Agriculture and Forestry University(Natural Science Edition),46(5):488-494.(in Chinese)付晶晶,杨彩霞,韩彤,廖一鸣,李梁,李梦新,孙伟,王蒴.2017.小西葫芦黄花叶病毒辽宁分离物的鉴定与序列分析.福建农林大学学报(自然科学版),46(5):488-494.
Kimalov B,Gal-On A,Stav R,Belausov E,Arazi T.2004.Maintenance of coat protein N-terminal net charge and not primary sequence is essential for Zucchini yellow mosaic virus systemic infectivity.Journal of General Virology,85:3421.
Kwon S W,Kim M S,Choi H S,Kim K H.2005.Biological characteristics and nucleotide sequences of three Korean isolates of Zucchini yellow mosaic virus.Journal of General Plant Pathology,71:80-85.
Lesemann D E,Makkouk K M,Koenig R,Samman E N.1983.Natural infection of cucumbers by Zucchini yellow mosaic virus in lebanon.Journal of Phytopathology,108(3-4):304-313.
Lin S S,Hou R F,Yeh S D.2001.Complete genome sequence and genetic organization of a Taiwan isolate of Zucchini yellow mosaic virus.Botanical Bulletin of Academia Sinica,42:243-250.
Lisa V,Boccardo G,D’Agostino G,Dellavalle G,D’Aquilio M.1981.Characterization of potyvirus that causes Zucchini yellow mosaic.Phytopathology,71(7):667.
Liu Jin-liang,Shao Yun-hua,Zhang Gaung-min,Zhu Xiao-ping,Yang Guang-ling,Shi Cheng-kui,Li Xiang-dong.2009.Molecular characterization of Zucchini yellow mosaic virus isolate infecting pumpkin in Shandong.Acta Phytopathologica Sinica,39(5):544-548.(in Chinese)刘金亮,邵云华,张广民,竺晓平,杨广玲,时呈奎,李向东.2009.小西葫芦黄花叶病毒山东南瓜分离物的分子特性.植物病理学报,39(5):544-548.
Maina S,Coutts B A,Edwards O,Almeida L D,Kehoe M,Ximenes A.2017.Zucchini yellow mosaic virus populations from east timorese and northern australian cucurbit crops:molecular properties,genetic connectivity and biosecurity implications.Plant Disease,101(7):1236-1245.
Monger W A,Seal S,Isaac A M,Foster G D.2001.Molecular characterization of the Cassava brown streak virus coat protein.Plant Pathology,50(4):527-534.
NovákováS,Svoboda J,Glasa M.2014.Analysis of the complete sequences of two biologically distinct Zucchini yellow mosaic virus isolates further evidences the involvement of a single amino acid in the virus pathogenicity.Acta Virologica,58(4):364-367.
Pan Ya-nan,Han Jian,Wu Hai-bo,Ji Yan-ling,Wang Chun-li,Liu Fang,Luo Ming,Zhang Xiang-lin.2016.Detection of viruses infecting Hami melon and their molecular identification in Xinjiang.Acta Horticulturae Sinica,43(6):1107-1116.(in Chinese)潘亚南,韩剑,吴海波,吉艳玲,王纯利,刘芳,罗明,张祥林.2016.新疆哈密瓜病毒的DAS-ELISA检测和分子鉴定.园艺学报,43(6):1107-1116.
Provvidenti R,Gonsalves D,Humaydan H S.1984.Occurrence of Zucchini yellow mosaic virus in cucurbits from Connecticut,New York,Florida and California.Plant Disease,68(1):443-446.
Qin Bi-xia,Zheng Hong-ying,Cai Jian-he,Chen-Jiong,Liu Zhi-ming,Chen Jian-ping.2005.Identification of a Zucchini yellow mosaic virus isolate from Siraitia grosvenori(Swingle)C.Jeffrey in Guangxi Province and preparation of coat protein antiserum.Chinese Journal of Virology,21(6):468-472.(in Chinese)秦碧霞,郑红英,蔡健和,陈炯,刘志明,陈剑平.2005.侵染广西罗汉果的小西葫芦黄花叶病毒的鉴定及抗血清制备.病毒学报,21(6):468-473.
Reyes A,Wu M,Mcnulty N P,Rohwer F L,Gordon J I.2013.Gnotobiotic mouse model of phage-bacterial host dynamics in the human gut.PNAS,110(50):20236-20241.
Romay G,Lecoq H,Desbiez C.2013.First Report of Watermelon mosaic virus naturally infecting Cucumis anguria.Plant Disease,97(11):1515.
Rybicki E P,Shukla D D.1992.Coat protein phylogeny and systematics of potyviruses.Archives of Virology Supplementum,5:139.
Scotto-Lavino E,Du G,Frohman M A.2006.5′end cDNA amplification using classic RACE.Nature Protocols,1:2555-2562.
Shi M,Lin X D,Tian J H,Chen L J,Chen X,Li C X.2016a.Redefining the invertebrate RNA virosphere.Nature,540(7634):539.
Shi Y,Sun X,Wang Z,Li H,Yan Z.2016b.First report of Zucchini yellow mosaic virus infecting sesame in China.Plant Disease,100(12):2545.
Simmons H E,Holmes E C,Gildow F E,Bothegoralczyk M A,Stephenson A G.2011.Experimental verification of seed transmission of Zucchini yellow mosaic virus.Plant Disease,95:751-754.
Simmons H E,Dunham J P,Zinn K E,Munkvold G P,Holmes E C,Stephenson A G.2013.Zucchini yellow mosaic virus(ZYMV,Potyvirus):vertical transmission,seed infection and cryptic infections.Virus Research,176(1-2):259-264.
Tang Chao.2016.Phylogenetic analysis of Zucchini yellow mosaic virus in Hunan Province,China[M.D.Dissertation].Changsha:Hunan Agricultural University.(in Chinese)唐超.2016.湖南省小西葫芦黄花叶病毒进化分析[硕士论文].长沙:湖南农业大学.
Wang W Q,Natsuaki T,Kosaka Y,Okuda S.2006.Comparison of the nucleotide and amino acid sequences of parental and attenuated isolates of Zucchini yellow mosaic virus.Journal of General Plant Pathology,72(1):52-56.
Wen Zhao-hui,Liu Ya-li.2010.Identification of Zucchini yellow mosaic virus infecting squash and Zucchini in Gansu Province.Plant Protection,36(4):120-122.(in Chinese)文朝慧,刘雅莉.2010.甘肃省南瓜及西葫芦小西葫芦黄花叶病毒病鉴定.植物保护,36(4):120-122.
Wisler G C,Purcifull D E,Hiebert E.1995.Characterization of the P1 protein and coding region of the Zucchini yellow mosaic virus.J Gen Virol,76:37-45.
Zhao M F,Chen J,Zheng H Y,Adams M J,Chen J P.2003.Molecular analysis of Zucchini yellow mosaic virus isolates from Hangzhou,China.Journal of Phytopathology,151:307-311.
Zhao Qin,Xie Da-sen,He Xiao-ming,Li Hua-ping,You Yi,Luo Shao-bo,Peng Qing-wu.2016.Detection of Zucchini yellow mosaic virus infecting wax gourd in Guangdong Province and genetic variation of its coat protein gene.Acta Horticulturae Sinica,43:151-160.(in Chinese)赵芹,谢大森,何晓明,李华平,尤毅,罗少波,彭庆务.2016.广东地区冬瓜感染的小西葫芦黄花叶病毒检测及其外壳蛋白基因多样性分析.园艺学报,43:151-160.
Zheng Guang-yu,Dong Tao.1991.Occurrence of Zucchini yellow mosaic virus in Xinjiang.Acta Phytopathologica Sinica,21(1):72.(in Chinese)郑光宇,董涛.1991.在新疆发生的小西葫芦黄化花叶病毒的研究初报.植物病理学报,21(1):72.
Bananej K,Keshavarz T,Vahdat A,Salekdeh G H,Glasa M.2010.Biological and Molecular Variability of Zucchini yellow mosaic virus in Iran.Journal of Phytopathology,156:654-659.
Chare E R,Holmes E C.2006.A phylogenetic survey of recombination frequency in plant RNA viruses.Archives of Virology,151:933.
Chen Jie-yun,Chen Ji-shuang,Hong Jian.2003.Sequence diversity of 3′end genome for Zucchini yellow mosaic virus.Acta Microbiology Sinica,43(3):388-394.(in Chinese)陈洁云,陈集双,洪健.2003.小西葫芦黄化花叶病毒分离物的3′末端序列多态性研究.微生物学报,43(3):388-394.
Chen Y K,Hong Y H.2008.First report of Begonia chlorotic ringspot caused by Zucchini yellow mosaic virus in Taiwan.Plant Disease,92(8):1247.
Chung Y W,Miller W A,Atkins J F,Firth A E.2008.An overlapping essential gene in the Potyviridae.Proceedings of the National Academy of Sciences of the United States of America,105(15):5897.
Cordero T,Cerdán L,Carbonell A,Katsarou K,Kalantidis K,Daròs J A.2016.Dicer-like 4 is involved in restricting the systemic movement of Zucchini yellow mosaic virus in Nicotiana benthamiana.Molecular Plant-Microbe Interactions,MPMI:MPMI11160239R.
Coutts B A,Kehoe M A,Webster C G,Wylie S J,Jones R A.2011.Zucchini yellow mosaic virus:biological properties,detection procedures and comparison of coat protein gene sequences.Archives of Virology,156(12):2119-2131.
Davis R F.1986.Partial characterization of Zucchini yellow mosaic virus isolated from squash in Turkey.Plant Disease,70(8):735-738.
Davis R F,Mizuki M K.1987.Detection of cucurbit viruses in New Jersey.Plant Disease,22(71):199-202.
Desbiez C,Gal-On A,Girard M,Wipf-Scheibel C,Lecoq H.2003.Increase in Zucchini yellow mosaic virus symptom severity in tolerant zucchini cultivars is related to a point mutation in P3 protein and is associated with a loss of relative fitness on susceptible plants.Phytopathology,93:1478-1484.
Desbiez C,Wipf-Scheibel C,Lecoq H.2002.Biological and serological variability,evolution and molecular epidemiology of Zucchini yellow mosaic virus(ZYMV,Potyvirus)with special reference to Caribbean islands.Virus Research,85:5-16.
Dunham J P,Simmons H E,Holmes E C,Stephenson A G.2014.Analysis of viral(Zucchini yellow mosaic virus)genetic diversity during systemic movement through a C.pepo vine.Virus Research,191(1):172-179.
Gal-On A,Antignus Y,Rosner A,Raccah B.1990.Nucleotide sequence of the Zucchini yellow mosaic virus capsid-encoding gene and its expression in Escherichia coli.Gene,87:273-277.
Glasa M,PittnerováS.2010.Complete genome sequence of a slovak isolate of Zucchini yellow mosaic virus(ZYMV)provides further evidence of a close molecular relationship among central European ZYMV isolates.Journal of Phytopathology,154(7-8):436-440.
Green K J,Brown C J,Gray S M,Karasev A V.2017.Phylogenetic study of recombinant strains of Potato virus Y:Virology,507:40-52.
Green K J,Brown C J,Karasev A V.2018.Genetic diversity of Potato virus Y(PVY):sequence analyses reveal ten novel PVY recombinant structures.Arch Virol,163:23-32.
Gu Qin-sheng,Fan Zai-feng,Piero Rogger,Chen Hong-yun,Yu Zheng-wang,Li Huai-fang.2003.Cloning and sequence analysis of the coat protein gene of Zucchini yellow mosaic virus.Acta Phytopathologica Sinica,33(6):498-502.(in Chinese)古勤生,范在丰,Piero Rogger,陈红运,俞正旺,李怀方.2003.小西葫芦黄花叶病毒外壳蛋白基因的克隆及序列分析.植物病理学报,33(6):498-502.
Fu Jing-jing,Yang Cai-xia,Han Tong,Liao Yi-ming,Li Liang,Li Meng-xin,Sun Wei,Wang Shuo.2017.Identification and sequence analysis of Zucchini yellow mosaic virus in Liaoning.Journal of Fujian Agriculture and Forestry University(Natural Science Edition),46(5):488-494.(in Chinese)付晶晶,杨彩霞,韩彤,廖一鸣,李梁,李梦新,孙伟,王蒴.2017.小西葫芦黄花叶病毒辽宁分离物的鉴定与序列分析.福建农林大学学报(自然科学版),46(5):488-494.
Kimalov B,Gal-On A,Stav R,Belausov E,Arazi T.2004.Maintenance of coat protein N-terminal net charge and not primary sequence is essential for Zucchini yellow mosaic virus systemic infectivity.Journal of General Virology,85:3421.
Kwon S W,Kim M S,Choi H S,Kim K H.2005.Biological characteristics and nucleotide sequences of three Korean isolates of Zucchini yellow mosaic virus.Journal of General Plant Pathology,71:80-85.
Lesemann D E,Makkouk K M,Koenig R,Samman E N.1983.Natural infection of cucumbers by Zucchini yellow mosaic virus in lebanon.Journal of Phytopathology,108(3-4):304-313.
Lin S S,Hou R F,Yeh S D.2001.Complete genome sequence and genetic organization of a Taiwan isolate of Zucchini yellow mosaic virus.Botanical Bulletin of Academia Sinica,42:243-250.
Lisa V,Boccardo G,D’Agostino G,Dellavalle G,D’Aquilio M.1981.Characterization of potyvirus that causes Zucchini yellow mosaic.Phytopathology,71(7):667.
Liu Jin-liang,Shao Yun-hua,Zhang Gaung-min,Zhu Xiao-ping,Yang Guang-ling,Shi Cheng-kui,Li Xiang-dong.2009.Molecular characterization of Zucchini yellow mosaic virus isolate infecting pumpkin in Shandong.Acta Phytopathologica Sinica,39(5):544-548.(in Chinese)刘金亮,邵云华,张广民,竺晓平,杨广玲,时呈奎,李向东.2009.小西葫芦黄花叶病毒山东南瓜分离物的分子特性.植物病理学报,39(5):544-548.
Maina S,Coutts B A,Edwards O,Almeida L D,Kehoe M,Ximenes A.2017.Zucchini yellow mosaic virus populations from east timorese and northern australian cucurbit crops:molecular properties,genetic connectivity and biosecurity implications.Plant Disease,101(7):1236-1245.
Monger W A,Seal S,Isaac A M,Foster G D.2001.Molecular characterization of the Cassava brown streak virus coat protein.Plant Pathology,50(4):527-534.
NovákováS,Svoboda J,Glasa M.2014.Analysis of the complete sequences of two biologically distinct Zucchini yellow mosaic virus isolates further evidences the involvement of a single amino acid in the virus pathogenicity.Acta Virologica,58(4):364-367.
Pan Ya-nan,Han Jian,Wu Hai-bo,Ji Yan-ling,Wang Chun-li,Liu Fang,Luo Ming,Zhang Xiang-lin.2016.Detection of viruses infecting Hami melon and their molecular identification in Xinjiang.Acta Horticulturae Sinica,43(6):1107-1116.(in Chinese)潘亚南,韩剑,吴海波,吉艳玲,王纯利,刘芳,罗明,张祥林.2016.新疆哈密瓜病毒的DAS-ELISA检测和分子鉴定.园艺学报,43(6):1107-1116.
Provvidenti R,Gonsalves D,Humaydan H S.1984.Occurrence of Zucchini yellow mosaic virus in cucurbits from Connecticut,New York,Florida and California.Plant Disease,68(1):443-446.
Qin Bi-xia,Zheng Hong-ying,Cai Jian-he,Chen-Jiong,Liu Zhi-ming,Chen Jian-ping.2005.Identification of a Zucchini yellow mosaic virus isolate from Siraitia grosvenori(Swingle)C.Jeffrey in Guangxi Province and preparation of coat protein antiserum.Chinese Journal of Virology,21(6):468-472.(in Chinese)秦碧霞,郑红英,蔡健和,陈炯,刘志明,陈剑平.2005.侵染广西罗汉果的小西葫芦黄花叶病毒的鉴定及抗血清制备.病毒学报,21(6):468-473.
Reyes A,Wu M,Mcnulty N P,Rohwer F L,Gordon J I.2013.Gnotobiotic mouse model of phage-bacterial host dynamics in the human gut.PNAS,110(50):20236-20241.
Romay G,Lecoq H,Desbiez C.2013.First Report of Watermelon mosaic virus naturally infecting Cucumis anguria.Plant Disease,97(11):1515.
Rybicki E P,Shukla D D.1992.Coat protein phylogeny and systematics of potyviruses.Archives of Virology Supplementum,5:139.
Scotto-Lavino E,Du G,Frohman M A.2006.5′end cDNA amplification using classic RACE.Nature Protocols,1:2555-2562.
Shi M,Lin X D,Tian J H,Chen L J,Chen X,Li C X.2016a.Redefining the invertebrate RNA virosphere.Nature,540(7634):539.
Shi Y,Sun X,Wang Z,Li H,Yan Z.2016b.First report of Zucchini yellow mosaic virus infecting sesame in China.Plant Disease,100(12):2545.
Simmons H E,Holmes E C,Gildow F E,Bothegoralczyk M A,Stephenson A G.2011.Experimental verification of seed transmission of Zucchini yellow mosaic virus.Plant Disease,95:751-754.
Simmons H E,Dunham J P,Zinn K E,Munkvold G P,Holmes E C,Stephenson A G.2013.Zucchini yellow mosaic virus(ZYMV,Potyvirus):vertical transmission,seed infection and cryptic infections.Virus Research,176(1-2):259-264.
Tang Chao.2016.Phylogenetic analysis of Zucchini yellow mosaic virus in Hunan Province,China[M.D.Dissertation].Changsha:Hunan Agricultural University.(in Chinese)唐超.2016.湖南省小西葫芦黄花叶病毒进化分析[硕士论文].长沙:湖南农业大学.
Wang W Q,Natsuaki T,Kosaka Y,Okuda S.2006.Comparison of the nucleotide and amino acid sequences of parental and attenuated isolates of Zucchini yellow mosaic virus.Journal of General Plant Pathology,72(1):52-56.
Wen Zhao-hui,Liu Ya-li.2010.Identification of Zucchini yellow mosaic virus infecting squash and Zucchini in Gansu Province.Plant Protection,36(4):120-122.(in Chinese)文朝慧,刘雅莉.2010.甘肃省南瓜及西葫芦小西葫芦黄花叶病毒病鉴定.植物保护,36(4):120-122.
Wisler G C,Purcifull D E,Hiebert E.1995.Characterization of the P1 protein and coding region of the Zucchini yellow mosaic virus.J Gen Virol,76:37-45.
Zhao M F,Chen J,Zheng H Y,Adams M J,Chen J P.2003.Molecular analysis of Zucchini yellow mosaic virus isolates from Hangzhou,China.Journal of Phytopathology,151:307-311.
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