劉樹(shù)芳, 董麗英, 李迅東, 張建洲, 楊勤忠*
1云南省農(nóng)業(yè)科學(xué)院農(nóng)業(yè)環(huán)境資源研究所,云南 昆明 650205; 2楚雄市種子管理站,云南 楚雄 657000
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水稻莖線蟲(chóng)病研究進(jìn)展
劉樹(shù)芳1, 董麗英1, 李迅東1, 張建洲2, 楊勤忠1*
1云南省農(nóng)業(yè)科學(xué)院農(nóng)業(yè)環(huán)境資源研究所,云南 昆明 650205;2楚雄市種子管理站,云南 楚雄 657000
水稻莖線蟲(chóng)病是水稻主要病害之一,能造成水稻嚴(yán)重減產(chǎn)。在我國(guó)水稻莖線蟲(chóng)屬于檢疫對(duì)象。該病害在孟加拉國(guó)、印度及緬甸等危害嚴(yán)重。本文結(jié)合國(guó)內(nèi)外研究現(xiàn)狀,從水稻莖線蟲(chóng)病原、形態(tài)、分布、癥狀等方面綜述了水稻莖線蟲(chóng)病的研究進(jìn)展,以期為我國(guó)開(kāi)展檢疫和防治提供參考。
水稻; 水稻莖線蟲(chóng); 發(fā)生; 分布
水稻莖線蟲(chóng)病又名稻窄莖線蟲(chóng)病、稻褐斑線蟲(chóng)病(洪劍鳴和董賢明,2006),該病于1913年在孟加拉國(guó)首先發(fā)現(xiàn)(Butler,1913),并命名為Tylenchusangustus,1936年更名為Ditylenchusangustus(Bultler) Filipjev并沿用至今。水稻莖線蟲(chóng)病在印度、孟加拉國(guó)、緬甸、巴基斯坦、泰國(guó)、越南、菲律賓、馬來(lái)西亞等亞洲國(guó)家均報(bào)道有發(fā)生。我國(guó)早在1986年就將水稻莖線蟲(chóng)列為禁止入境的危險(xiǎn)性有害生物,并對(duì)來(lái)自水稻莖線蟲(chóng)疫區(qū)的水稻及其附屬產(chǎn)品、寄主植物及其產(chǎn)品等采取了嚴(yán)格的檢疫措施,嚴(yán)防水稻莖線蟲(chóng)的傳入(黃可輝和郭瓊霞,2003; 李芳榮等,2015)。
隨著“一帶一路”國(guó)家戰(zhàn)略的實(shí)施,我國(guó)與世界各國(guó)的貿(mào)易日趨頻繁,因此在加強(qiáng)合作交流的基礎(chǔ)上,如何保障我國(guó)的糧食安全成為新的歷史使命。本文綜合國(guó)內(nèi)外相關(guān)研究,闡述了我國(guó)水稻檢疫性病害——水稻莖線蟲(chóng)病的國(guó)外發(fā)生危害和研究現(xiàn)狀,以期為我國(guó)的水稻生產(chǎn)和糧食安全提供參考。
1.1病原及形態(tài)學(xué)
水稻莖線蟲(chóng)病病原線蟲(chóng)(Butler,1913; Filipjev,1936),屬墊刃目粒線蟲(chóng)科莖線蟲(chóng)屬。由于物種內(nèi)的個(gè)體差異和環(huán)境因素的不同,導(dǎo)致D.angustus的體長(zhǎng)(L)、體長(zhǎng)/最大體寬(a)、體長(zhǎng)/體前端至食道與腸連接處的距離(b)、體長(zhǎng)/尾長(zhǎng)(c)等形態(tài)特征有所差別,雄蟲(chóng)L0.7~1.25 mm,a36.0~63.7,b5.42~9.0,c15.0~-24.0;雌蟲(chóng)L0.6~1.1 mm,a36.0~60.0,b6.0~8.0,c10.4~26.0(戚龍君等,2002; Bulter,1913; Das & Bajaj,2008; Goodey,1932; Mian & Latif,1994; Seshadri & Dasgupta,1975)。在分子鑒定方面,Ibrahimetal. (1994)依據(jù)秀麗隱桿線蟲(chóng)Caenorhabditiselegans的18S和26S核糖體RNA基因的保守序列設(shè)計(jì)PCR引物進(jìn)行擴(kuò)增,并利用限制性內(nèi)切酶酶切PCR產(chǎn)物后進(jìn)行電泳分析,可將D.angustus和Aphelenchoides劃分開(kāi)。
1.2分布及危害
D.angustus主要分布在孟加拉國(guó)、印度、印度尼西亞、馬來(lái)西亞、菲律賓、緬甸、泰國(guó)南部以及越南的湄公河流域三角區(qū)(CABI/EPPO,2015; Kyndt,2014)。Cuc & Kinh (1981)報(bào)道了越南于1975—1980年在湄公河三角區(qū)發(fā)生水稻莖線蟲(chóng)病,水稻減產(chǎn)達(dá)50%~100%;Cox & Rahman(1980)報(bào)道,泰國(guó)南部水稻莖線蟲(chóng)病的發(fā)生造成水稻減產(chǎn)20%~90%;2014年,緬甸伊洛瓦底省因水稻莖線蟲(chóng)病暴發(fā),水稻產(chǎn)量較正常年份減產(chǎn)70%左右(IAPPS,2014)。Latifetal. (2013a)開(kāi)展了單一接種D.angustus或水稻干尖線蟲(chóng)A.besseyi和不同比例混合接種時(shí)對(duì)糧食產(chǎn)量影響的研究,試驗(yàn)表明,僅接種D.angustus時(shí),糧食產(chǎn)量損失達(dá)62%,比僅接種A.besseyi和不同比例混合接種的產(chǎn)量損失高。由于水稻莖線蟲(chóng)病危害的嚴(yán)重性,中國(guó)、老撾和日本等國(guó)家均將其列為檢疫對(duì)象(黃可輝和郭瓊霞,2003; 李芳榮等,2015; 李喜陽(yáng),1998)。
1.3癥狀
Dasetal. (2011)研究表明,D.angustus侵染水稻10~15 d后即表現(xiàn)病狀。其癥狀在水稻整個(gè)生育期不同部位均可表現(xiàn)。發(fā)病初期,幼嫩葉片褪綠、畸形;隨著病程的發(fā)展,葉片出現(xiàn)分散的暗斑,莖干節(jié)間區(qū)域變深褐色,葉基部和葉鞘扭曲或畸形,下部節(jié)間膨脹;發(fā)病末期,葉片褪綠,全株枯萎或死亡。若發(fā)生在孕穗期,受害嚴(yán)重的穗和穗軸變暗褐色,穗常被包裹在已發(fā)病的葉鞘內(nèi),造成水稻不能正常抽穗,發(fā)生較輕的雖能正常抽穗,但不能結(jié)實(shí)或僅穗的頂部少量結(jié)實(shí)(洪劍鳴和董賢明,2006; Rahman,2003)。
Butler (1913)按照是否抽穗將該病害的癥狀分為2種類型:“Thor”型,即葉片和穗扭曲,不能抽穗;“Pucca”型,即能抽穗但不能結(jié)實(shí)或少量結(jié)實(shí)。Cox & Rahman (1980)將該病害的癥狀分為3種類型:ufraⅠ,被葉鞘包裹不能正常抽穗;ufra Ⅱ,稻穗未完全抽穗,下部稻穗被葉鞘包裹,不能結(jié)實(shí);ufra Ⅲ,稻穗能正常抽穗但大部分不能結(jié)實(shí)。
1.4發(fā)病條件
在水稻生產(chǎn)季節(jié)特別是高溫高濕條件有利于病害發(fā)生,當(dāng)溫度為27~30 ℃、相對(duì)濕度在75%以上發(fā)病較嚴(yán)重(Bridgeetal.,2005; Bridge & Starr,2007; Cuc & Kinh,1981; Rahman & Evans,1987)。
1.5侵染危害
D.angustus在適宜的溫、濕度條件下,通過(guò)在植物體內(nèi)的取食和遷移,不僅造成水稻葉片褪綠、畸形,不能正常抽穗(Rahman & Evans,1987),而且降低了水稻的抵抗能力,易引起其他病害的發(fā)生,增加感染的風(fēng)險(xiǎn)。Alietal. (1997a)研究表明,由于D.angustus的侵染,水稻體內(nèi)氮含量明顯增加,加重了如稻瘟病MagnaportheoryzaeHebert、葉鞘腐敗病Sarocladiumoryzae(Sawada)和細(xì)菌性條斑病Xanthomonasoryzaepv.oryzicola等病害的發(fā)生。
Alietal.(1997b)研究表明,D.angustus能在1~30 ℃下存活,發(fā)育最適宜溫度為25 ℃。雌蟲(chóng)成熟后,當(dāng)溫度達(dá)到10.6 ℃即可產(chǎn)卵;隨著溫度升高,孵化時(shí)間縮短,在20、25、30 ℃下分別需4、3、2 d即可孵化;溫度高于35 ℃時(shí),雌蟲(chóng)不產(chǎn)卵且卵也不能孵化。在相對(duì)濕度75%、溫度為30 ℃條件下水稻莖線蟲(chóng)10~20 d可完成一個(gè)世代(Kyndt,2014)。在東南亞和南亞,由于地理位置鄰近赤道和印度洋,受海洋季風(fēng)的影響常年高溫高濕,為D.angustus的侵染提供了有利的自然條件(Plowright & Gill,1994)。在植物收獲后,寄生于水稻殘株、土壤、寄主植物和種子里越冬(Cox & Rahman,1979; Cuc,1982, Ibrahim & Perry,1993; Kinh,1981; Prasad & Varaprasad,2002)。當(dāng)溫度和濕度適宜時(shí),D.angustus再次侵染水稻植株或寄主。
研究表明,當(dāng)濕度達(dá)75%以上時(shí),D.angustus能完成從土壤到水稻植株、從病殘?bào)w到健康植株的侵染過(guò)程,當(dāng)濕度大于85%時(shí)也可通過(guò)水稻葉片的接觸從帶病植株侵染到健康植株(Bridgeetal.,2005; Cuc & Kinh,1981; Perry,1995; Rahman & Evans,1987)。
D.angustus的生命力極強(qiáng),在花和葉鞘內(nèi)至少能存活6個(gè)月(Bridgeetal.,2005; Latifetal.,2011);在干燥的土壤里能存活1.5個(gè)月,在淹水的土壤里至少也存活4個(gè)月(Butler,1913);在含水量高于14%的飽滿谷粒和空癟的谷粒中也能存活(Butler,1919; Dasetal.,2011; Ibrahim & Perry,1993)。研究還表明,將種子在太陽(yáng)下暴曬1 d(32 ℃),并存放在(22±5) ℃保存3個(gè)月后仍有27%的水稻感染莖線蟲(chóng)病(Prasad & Varaprasad,2002)。因此,灌溉水、地表徑流、水稻殘株、穗、種子等為D.angustus的傳播創(chuàng)造了條件。
D.angustus對(duì)水稻專性寄生,寄主包括栽培稻和野生稻以及部分雜草,如亞洲栽培水稻OryzasativaL.、普通野生稻O.rufipogon、非洲栽培稻O.glaberrima、闊葉稻O.latifolia、疣粒稻O.meyeriana、印度野生稻O.nivara、藥用稻O.officinalis、野生紅稻O.perennis和光頭稗Echinochloacolona(L.)、水禾Hygroryzaaristata(Retz.) Nees、間序囊穎草Sacciolepisinterrupta(Wild.) Stapf等雜草(Prasadetal.,2011; USDA,2011)。
清潔田園,清除病殘?bào)w和田間雜草,采取水旱輪作模式以及種植早熟品種,能降低水稻莖線蟲(chóng)病發(fā)病(Chakraborti,2000; Rahman,2003)。利用抗性品種也能較好地控制水稻莖線蟲(chóng)的發(fā)生。Khanametal. (2016)從85個(gè)水稻品種中,篩選到1個(gè)具有高抗水稻莖線蟲(chóng)病的地方品種(Manikpukha)及6個(gè)抗病的品種。呋喃丹、滅線磷、異狄氏劑和久效磷等化學(xué)藥劑對(duì)D.angustus的防治效果較好,其中以呋喃丹的防治效果最好(Latifetal.,2011、2013b)。由于化學(xué)藥劑易造成環(huán)境污染的問(wèn)題,相關(guān)國(guó)家正開(kāi)展植物源殺蟲(chóng)劑如印楝素防治水稻莖線蟲(chóng)病的研究工作(Chakraborti,2000; Latifetal.,2006)。
水稻莖線蟲(chóng)病是水稻重要病害,該病具有發(fā)病快、防治難且病原線蟲(chóng)在水稻植株、種子中存活時(shí)間長(zhǎng)等特點(diǎn)。病原線蟲(chóng)一旦侵入水稻,會(huì)造成水稻減產(chǎn)甚至絕收。大湄公河次區(qū)域是亞洲主要的水稻種植區(qū)和出口區(qū),而該病在泰國(guó)、緬甸、越南均有報(bào)道,這些國(guó)家又與中國(guó)云南省比鄰?!耙粠б宦贰眹?guó)家戰(zhàn)略的實(shí)施和國(guó)際間貿(mào)易的日趨頻繁,增加了水稻莖線蟲(chóng)病的傳入風(fēng)險(xiǎn)。為保障我國(guó)糧食安全,在加強(qiáng)進(jìn)口種質(zhì)材料及其附屬物嚴(yán)格檢疫的同時(shí),需進(jìn)一步開(kāi)展風(fēng)險(xiǎn)評(píng)估分析,并通過(guò)國(guó)際合作,監(jiān)測(cè)水稻莖線蟲(chóng)病在周邊國(guó)家的發(fā)生,以防傳入我國(guó)(王峰等,2007)。
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(責(zé)任編輯:郭瑩)
Research progress of rice ufra disease
Shu-fang LIU1, Li-ying DONG1, Xun-dong LI1, Jian-zhou ZHANG2, Qin-zhong YANG1*
1InstituteofAgriculturalEnvironmentandResources,YunnanAcademyofAgriculturalSciences,Kunming,Yunnan650205,China;2ChuxiongSeedServiceCenter,Chuxiong,Yunnan657000,China
The rice ufra disease, caused byDitylenchusangustus, is one of the important rice diseases that can cause severe yield loss.D.angustusis the quarantined pest in China. Ufra disease has seriously damaged rice production in countries such as Bangladesh, India and Myanmar. Here, we summarized the research progress of rice ufra disease in terms of its biology, morphology, distribution ofD.angustus, as well as symptoms on rice in the field. This information is of significance for carrying out plant quarantine and management in China.
Oryzasativa;Ditylenchusangustus; occurrence; distribution
2016-05-21接受日期(Accepted): 2016-07-04
科技部國(guó)家國(guó)際科技合作項(xiàng)目(2011DFB30040); 云南省對(duì)外科技合作計(jì)劃(國(guó)際科技合作)項(xiàng)目(2014IA009)
劉樹(shù)芳, 男, 碩士研究生。 研究方向: 植物病理。 E-mail: lshufang80@163.com
Author for correspondence), E-mail: qzhyang@163.com
10. 3969/j.issn.2095-1787.2016.03.014