警惕番茄褪綠病毒在我國的傳播和危害

周 濤, 楊普云, 趙汝娜, 師迎春, 原 鍇, 范在豐

(1. 中國農業(yè)大學植物病理學系,北京 100193；2. 全國農業(yè)技術推廣服務中心,北京 100125；3. 北京市植物保護站,北京 100029；4. 北京市房山區(qū)植物保護站,北京 102488)

有害生物動態(tài)

警惕番茄褪綠病毒在我國的傳播和危害

周 濤1*, 楊普云2, 趙汝娜1, 師迎春3, 原 鍇4, 范在豐1

(1. 中國農業(yè)大學植物病理學系,北京 100193；2. 全國農業(yè)技術推廣服務中心,北京 100125；3. 北京市植物保護站,北京 100029；4. 北京市房山區(qū)植物保護站,北京 102488)

番茄褪綠病毒(Tomatochlorosisvirus,ToCV)侵染番茄引起番茄褪綠病毒病。發(fā)病植株下部葉片黃化、脈間褪綠、邊緣輕微卷曲,葉片光合作用顯著降低,果實變小,產量和品質明顯下降。該病毒最早于1998年在美國佛羅里達州發(fā)現,隨后在世界多地陸續(xù)報道。我國首先于2004年在臺灣報道,2013年又在北京和山東發(fā)現并鑒定了該病毒,同時在辣椒上檢測到該病毒。ToCV屬于長線形病毒科(Closteroviridae)毛形病毒屬(Crinivirus)成員,基因組為二分體正義單鏈RNA,由粉虱傳播。經初步調查和檢測,ToCV在我國北京、山東、河北、天津等省市相繼發(fā)生,給當地番茄生產造成了嚴重危害。ToCV傳播迅速,成為我國番茄生產中又一重要病毒,防控形勢嚴峻?；谝陨显?建議有關部門立即采取相應預防和防治措施,組織開展相關研究和攻關,控制該病毒在我國的傳播和危害。

番茄褪綠病毒； 脈間褪綠； 粉虱； 防治措施

1989年美國佛羅里達州溫室栽培的番茄(Solanumlycopersicum)上出現“黃葉紊亂”(yellow leaf disorder)癥狀,懷疑為生理或營養(yǎng)失調、農藥藥害或者病毒侵染所致；1998年,該病原被鑒定為番茄褪綠病毒(Tomatochlorosisvirus, ToCV)[1]。ToCV侵染番茄導致番茄葉片脈間黃化,部分區(qū)域變紅,發(fā)育受阻,輕微卷曲,通常在老葉上表現明顯,新生枝條和葉片表現正常,隨著病害發(fā)展,發(fā)生脈間壞死,葉片變脆易碎、葉片變厚(圖1)。ToCV侵染番茄引起的癥狀極易與物理損傷、營養(yǎng)失調或農藥藥害混淆,但典型的癥狀特點是植株中下部葉片發(fā)病,而新生部分正常。

已知有兩種長線形病毒侵染番茄引起番茄褪綠病,一種是番茄褪綠病毒(ToCV),一種是番茄侵染性褪綠病毒(Tomatoinfectiouschlorosisvirus, TICV)。ToCV和TICV在番茄上引起癥狀相同,且均導致植株長勢變弱,果實產量降低[2]。我國目前僅發(fā)現ToCV。

1 世界分布和在我國的危害

目前,除大洋洲和南極洲未見報道外,其余各洲均報道有ToCV發(fā)生。在歐洲,葡萄牙[3]、意大利[4-5]、法國[6]、匈牙利[7]、西班牙[8]和希臘[9]報道有ToCV發(fā)生；在北美洲,美國[1]、墨西哥[10]、古巴[11]、哥斯達黎加[12]報道有ToCV的分布；南美洲的巴西[13]報道ToCV發(fā)生；在非洲,毛里求斯[14]和蘇丹[15]有ToCV的分布；在亞洲,以色列[16]、塞浦路斯[17]、黎巴嫩[18]、土耳其[19]、日本[20]和中國[21-23]均有ToCV的分布。

我國首先在臺灣地區(qū)報道了ToCV的發(fā)生[21],隨后在北京[22]、江蘇[23]、山東[24]三省市溫室栽培的番茄植株上檢出ToCV,并且其發(fā)病率呈逐漸增長之勢。2013年我們在常規(guī)蔬菜病毒病調查過程中發(fā)現,番茄褪綠病毒病已在山東(壽光、莘縣)、河北(唐山、保定)、北京、天津等省市發(fā)生,一般田塊的發(fā)病率高于50%(圖2)。采集樣品帶回實驗室進行分子檢測,這些樣品全部被ToCV侵染(未發(fā)表結果)。ToCV已給當地番茄生產造成了嚴重危害,受害番茄產量降低60%,單價降低50%,成為我國番茄生產中又一重要病毒。

2 分類、基因組和粒子特性

根據國際病毒分類委員會(International Committee on Taxonomy of Viruses,ICTV)第9次分類報告所述,ToCV屬于長線形病毒科(Closteroviridae)毛形病毒屬(Crinivirus)[25]。ToCV的基因組為二分體正義單鏈RNA(+ssRNA),RNA的5′端可能有一個甲基化帽子結構,3′端既無Ploy(A),也不形成tRNA樣結構,可能形成發(fā)夾結構[25]。目前世界上報道了7個分離物的基因組全序列,包括美國分離物[26],西班牙的2個分離物[8,27],希臘分離物[9]、巴西分離物[28],以及中國北京分離物[29]和山東分離物[24]。病毒粒子彎曲長線形,長800～850 nm,直徑約為12 nm,呈螺旋對稱結構,螺距3.4～3.8 nm[30]。ToCV基因組RNA1和RNA2分別包裝在兩種不同的病毒粒子中,兩者對成功侵染寄主都是必需的。

3 寄主范圍

ToCV可以侵染茄科(Solanaceae)、菊科(Compositae)、藜科(Chenopodiaceae)、莧科(Amaranthaceae)、番杏科(Aizoaceae)、夾竹桃科(Apocynaceae)及白花丹科(Plumbaginaceae)等7科25種植物。其中,茄科寄主數目最多,如：番茄(S.lycopersicum)、辣椒(Capsicumannuum)、普通煙(Nicotianatabacum)、本生煙(N.benthamiana)等多種煙草[2]。一些常見雜草和觀賞植物也是ToCV的寄主,如苦苣菜(Sonchusoleraceus)、百日菊(Zinniaelegans)、矮牽牛(Petuniahybrida)等。

4 傳播方式

ToCV不能通過汁液摩擦傳播,其自然傳毒介體為粉虱。有4種粉虱能夠傳播ToCV,但傳毒效率差異較大。紋翅粉虱(Trialeurodesabutilonea)和B型煙粉虱(BemisiatabaciB-biotype)的傳毒效率較高,A型煙粉虱(B.tabaciA-biotype)和溫室白粉虱(T.vaporariorum)的傳毒效率則較低。紋翅粉虱帶毒長達5 d,B型煙粉虱帶毒為2 d,而A型煙粉虱和溫室白粉虱只能帶毒1 d,雖然傳毒效率有差異,但都可以進行有效的傳毒[2]。

5 防控措施

目前國際上還未發(fā)現對ToCV表現抗性的番茄材料,無抗病品種可以使用。因為ToCV的傳播介體為粉虱,使用殺蟲劑殺滅粉虱是控制損失的首選方法。因目前我國番茄產區(qū)粉虱種群主要為煙粉虱,煙粉虱同時又是番茄黃化曲葉病毒(Tomatoyellowleafcurlvirus,TYLCV)的傳播介體,所以可以借鑒番茄黃化曲葉病毒病的防治方法[31]來防治番茄褪綠病毒病。但要注意的是,雖然殺蟲劑能夠有效控制粉虱群體,但不能有效控制病毒,因為粉虱在被殺死前可能已經傳播了病毒。另外,番茄被ToCV侵染后3～4周才表現癥狀。因此,在觀察到癥狀時病毒已經侵入植株體內,這時再通過控制煙粉虱來防控,效果就不明顯了。

清除雜草等ToCV的寄主是控制ToCV的另一項重要措施。ToCV寄主范圍中等,其中許多能感染ToCV的雜草在番茄田間和園藝植物附近非常常見。因此清除番茄生產棚室內及周邊的其他寄主植物,同時控制傳播介體能夠降低病毒的擴散和損失。

相比于感染后2～3周發(fā)病的番茄黃化曲葉病毒病,ToCV引起的番茄褪綠病毒病潛育期更長,隱蔽性強,容易與缺素癥混淆,被發(fā)現或鑒定時,已大面積嚴重發(fā)生,造成防治不及時。近幾年來,菜農逐漸對番茄黃化曲葉病毒病熟悉,在番茄田間發(fā)現病株能即刻拔除,而對番茄褪綠病毒病還沒有認識,這也是番茄褪綠病毒病嚴重發(fā)生的重要原因。基于以上原因,建議有關部門立即采取相應預防和防治措施,組織研究單位開展相關研究和攻關,控制ToCV在我國的傳播和危害。

[1]Wisler G C, Li R H, Liu H, et al.Tomatochlorosisvirus: a new whitefly-transmitted, phloem-limited, bipartite closterovirus of tomato[J]. Phytopathology，1998,88(5):402-409.

[2]Wintermantel W M, Wisler G C. Vector specificity, host range, and genetic diversity ofTomatochlorosisvirus[J]. Plant Disease. 2006,90(6):814-819.

[3]Louro D, Accotto G P, Vaira A M. Occurrence and diagnosis ofTomatochlorosisvirusin Portugal[J]. European Journal of Plant Pathology， 2000,106(6):589-592.

[4]Accotto G P, Vaira A M, Vecchiati M, et al. First report ofTomatochlorosisvirusin Italy[J]. Plant Disease， 2001,85(11):1208.

[5]Parrella G, Scassillo L. First report ofTomatochlorosisvirus(ToCV) in Campania and ofTomatoinfectiouschlorosisvirus(TICV) in Calabria (Southern Italy)[J]. Informatore Fitopatologico, 2006,56(6):33-34.

[6]Dalmon A, Bouyer S, Cailly M, et al. First report ofTomatochlorosisvirus andTomatoinfectiouschlorosisvirusin tomato crops in France[J]. Plant Disease，2005,89(11):1243.

[7]Bese G, Bóka K, Krizbai L, et al. First report ofTomatochlorosisvirusin tomato from Hungary[J]. Plant Disease，2011,95(3):363.

[8]Lozano G, Moriones E, Navas-Castillo J. Complete nucleotide sequence of the RNA2 of the crinivirusTomatochlorosisvirus[J]. Archives of Virology， 2006,151(3):581-587.

[9]Kataya A, Stavridou E, Farhan K, et al. Nucleotide sequence analysis and detection of a Greek isolate ofTomatochlorosisvirus[J]. Plant Pathology， 2008,57(5):819-824.

[10]Alvarez Ruiz P, Jimenez C G, Leyva López N E, et al. First report ofTomatochlorosisvirusinfecting tomato crops in Sinaloa, Mexico[J]. Plant Pathology,2007,56(6):1043.

[11]Martínez-Zubiaur Y, Fiallo-Olivé E, Carrillo-Tripp J, et al. First report ofTomatochlorosisvirusinfecting tomato in single and mixed infections withTomatoyellowleafcurlvirusin Cuba[J]. Plant Disease,2008,92(5):836.

[12]Vargas J A, Hammond R, Hernandez E, et al. First report ofTomatochlorosisvirusinfecting sweet pepper in Costa Rica[J]. Plant Disease, 2011,95(11):1482.

[13]Barbosa J C, Teixeira A, Moreira A G, et al, First report ofTomatochlorosisvirusinfecting tomato crops in Brazil[J]. Plant Disease, 2008,92(12):1709.

[14]Lett J M, Hoareau M, Reynaud B, et al. First report ofTomatochlorosisvirusin tomato on Mauritius Island[J]. Plant Disease,2009,93(1):111.

[15]Fiallo-Olivé E, Hamed A A, Moriones E, et al. First report ofTomatochlorosisvirusinfecting tomato in Sudan[J]. Plant Disease,2011,95(12):1592.

[16]Segev L, Wintermantel W M, Polston J E, et al. First report ofTomatochlorosisvirusin Israel[J]. Plant Disease,2004,88(10):1160.

[17]Papayiannis L C, Ioannou N, Dovas C I, et al. First report ofTomatochlorosisviruson tomato crops in Cyprus[J]. Plant Pathology,2006,55(4):567.

[18]Abou-Jawdah Y, El Mohtar C, Atamian H, et al. First report ofTomatochlorosisvirusin Lebanon[J]. Plant Disease，2006,90(3):378.

[19]Cevik B, Erkis G. First report ofTomatochlorosisvirusin Turkey[J]. Plant Pathology，2008,57(4):767.

[20]Hirota T, Natsuaki T, Murai T, et al. Yellowing disease of tomato caused byTomatochlorosisvirusnewly recognized in Japan[J]. Journal of General Plant Pathology，2010,76(2):168-171.

[21]Tsai W S, Shih S L, Green S K, et al. First report of the occurrence ofTomatochlorosisvirusandTomatoinfectiouschlorosisvirusin Taiwan[J]. Plant Disease，2004,88(3):311.

[22]Zhao R, Wang R, Wang N, et al. First report ofTomatochlorosisvirusin China[J]. Plant Disease，2013,97(8):1123.

[23]Karwitha M, Feng Z, Yao M, et al. The complete nucleotide sequence of the RNA 1 of a chinese isolate ofTomatochlorosisvirus[J]. Journal of Phytopathology,2014,162(4):411-415.

[24]Zhao L, Li G, Gao Y, et al. Molecular detection and complete genome sequences ofTomatochlorosisvirusisolates from infectious outbreaks in China[J/OL]. Journal of Phytopathology, 2014-02-27. DOI:10.1111/jph.12236.

[25]King A M Q, Adams M J, Carstens E B, et al. Virus taxonomy:ninth report of the international committee on taxonomy of viruses[M]. Amsterdam:Elsevier Academic Press,2011.

[26]Wintermantel W M, Wisler G C, Anchieta A G, et al. The complete nucleotide sequence and genome organization ofTomatochlorosisvirus[J]. Archives of Virology,2005,150(11):2287-2298.

[27]Lozano G, Moriones E, Navas-Castillo J. Complete sequence of the RNA1 of a European isolate ofTomatochlorosisvirus[J]. Archives of Virology,2007,152(4):839-841.

[28]Albuquerque L C, Villanueva F, Resende R O, et al. Molecular characterization reveals BrazilianTomatochlorosisvirusto be closely related to a Greek isolate[J]. Tropical Plant Pathology,2013,38(4):332-336.

[29]Zhao R, Wang N, Wong R, et al. Characterization and full genome sequence analysis of a Chinese isolate ofTomatochlorosisvirus[J]. Acta Virologica,2014,58(1):92-94.

[30]Liu H, Wisler G C, Duffus J E. Particle lengths of whitefly-transmitted criniviruses[J]. Plant Disease,2000,84(7):803-805.

[31]周濤,師迎春,陳笑瑜,等. 北京地區(qū)番茄黃化曲葉病毒病的鑒定及防治對策討論[J]. 植物保護，2010,36(2):116-118.

AlarmingthetransmissionanddamageofTomatochlorosisvirusinChina

Zhou Tao1, Yang Puyun2, Zhao Runa1, Shi Yingchun3, Yuan Kai4, Fan Zaifeng1

(1.DepartmentofPlantPathology,ChinaAgriculturalUniversity,Beijing100193,China; 2.NationalAgro-TechnicalExtensionandServiceCentre,Beijing100125,China; 3.BeijingPlantProtectionStation,Beijing100029,China; 4.FangshanStationofPlantProtection,Beijing102488,China)

Tomatochlorosisvirus(ToCV) causes leaf chlorosis on infected tomato plants. The lower leaves become yellow, interveinal chlorosis, slightly curling on edges, and photosynthesis decreases obviously. The fruits on infected plants turn smaller, and the production and quality are reduced significantly. ToCV was firstly identified in Florida, USA, and then was reported in many countries. In China, ToCV was reported firstly from Taiwan in 2004. In 2013, ToCV was found on tomato plants in Beijing and Shandong Province, and was detected on pepper plants at the same time. Being a member ofCrinivirusin family Closteroviridae, ToCV has a bipartite positive single-stranded RNA, and is transmitted by whitefly. Preliminary survey indicated ToCV had spread in Beijing, Shandong, Hebei and Tianjin, and caused severe damage on local tomato production. As its fast transmission, ToCV has become one of the most important viruses for China’s tomato production. It is suggested that measures must be carried out to perform research on ToCV, and to control its transmission and damage on tomato plants.

Tomatochlorosisvirus； interveinal chlorosis； whitefly； control measures

2014-05-03

：2014-06-03

國家“863”計劃項目(2012AA100103)

S 432.41

：BDOI：10.3969/j.issn.0529-1542.2014.05.038

* 通信作者 E-mail: taozhoucau@cau.edu.cn