汪 雷，劉 瑤，丁一娟，王 雨，萬(wàn)華方，梅家琴，錢 偉

(西南大學(xué) 農(nóng)學(xué)與生物科技學(xué)院，重慶市油菜工程技術(shù)研究中心，重慶 北碚400715)

油菜菌核病研究進(jìn)展

汪 雷，劉 瑤，丁一娟，王 雨，萬(wàn)華方，梅家琴，錢 偉

(西南大學(xué) 農(nóng)學(xué)與生物科技學(xué)院，重慶市油菜工程技術(shù)研究中心，重慶 北碚400715)

菌核病是危害油菜生產(chǎn)的主要病害之一。結(jié)合已有文獻(xiàn)與本研究室的研究結(jié)果，文章就核盤菌的致病過程和致病機(jī)理、油菜菌核病抗病資源篩選、抗病育種現(xiàn)狀、抗性遺傳規(guī)律、抗病QTL定位及抗性基因發(fā)掘等方面進(jìn)行了總結(jié)和展望。

油菜；菌核??；致病機(jī)理；分子機(jī)制；抗病育種

油菜是重要的油料作物，在食品、冶金、機(jī)械等多種行業(yè)應(yīng)用廣泛，經(jīng)濟(jì)價(jià)值高。核盤菌(Sclerotiniasclerotiorum)是一種廣譜性病原真菌，可寄生于70多科400多種植物，由其引起的菌核病是油菜的一大主要病害[1]。在中國(guó)，菌核病可引起油菜減產(chǎn)10%～80%[2]；在澳大利亞，菌核病導(dǎo)致油菜年均減產(chǎn)24%[3]；在北美雙低油菜主要種植區(qū)域[4]以及歐洲油菜種植區(qū)[5-7]該病的危害也日趨嚴(yán)重，加之全球氣候變化，更可能加速該病害的流行[8]?；瘜W(xué)殺菌劑的使用對(duì)該病害有一定防控效果，但效率卻依賴于施用時(shí)間、植株發(fā)育階段及環(huán)境等條件[9]，且提高了生產(chǎn)成本[4]，還會(huì)引發(fā)環(huán)境污染等問題。因此，選擇和培育抗病品種是控制油菜菌核病最經(jīng)濟(jì)有效的途徑[10-11]。本文參照國(guó)內(nèi)外及本課題組的研究成果，綜述了核盤菌的致病機(jī)理，并從抗源篩選、抗性遺傳規(guī)律探索、抗病分子機(jī)制解析等方面總結(jié)了油菜菌核病目前的研究狀況，以期為油菜菌核病的抗性改良提供參考。

1 核盤菌的致病過程及致病機(jī)理

1.1 致病過程

核盤菌經(jīng)典的侵染循環(huán)路徑主要是菌核在土壤、染病莖桿或種子中越冬、越夏，當(dāng)外界環(huán)境適宜時(shí)，菌核可萌發(fā)產(chǎn)生菌絲或子囊孢子。子囊孢子成熟后從子囊中彈出，借氣流傳播，飄落到植株的葉片或花瓣上，在營(yíng)養(yǎng)和環(huán)境條件適宜時(shí)萌發(fā)形成菌絲，侵入植物體，使植物受侵部位的細(xì)胞、組織壞死，形成水漬狀病斑并腐爛，再漫延到其他健康植株，最后在患病部位尤其是莖桿中形成菌核。對(duì)油菜而言，其菌核病的發(fā)病符合這個(gè)過程(圖1)。核盤菌在油菜花期來臨前萌發(fā)，通過產(chǎn)生子囊孢子附著在油菜花瓣，繼而以花瓣為營(yíng)養(yǎng)基質(zhì)生長(zhǎng)萌發(fā)產(chǎn)生菌絲，菌絲再隨著花瓣的凋零飄落到油菜的葉片或莖桿上，開始實(shí)現(xiàn)對(duì)油菜的侵染[12]。

圖1 油菜菌核病的病害循環(huán)

1.2 致病機(jī)理

目前，對(duì)核盤菌的致病機(jī)理尚缺乏系統(tǒng)全面的闡述，但研究表明細(xì)胞壁降解酶和草酸的分泌對(duì)核盤菌的致病有十分重要的作用[13-18]。核盤菌能分泌多種作用于寄主表皮的水解酶，如多聚半乳糖醛酸酶(Polygalacturonase,PG)、纖維素酶(Cellulase)、半纖維素酶(Hemicellulase)、果膠酶(Pectinase)、蛋白酶(Protease)、角質(zhì)酶(Cutinase)、葡聚糖酶(Dextranase)等酶類，攻破寄主表層防護(hù)，降解細(xì)胞壁，侵染組織，為病菌提供營(yíng)養(yǎng)，便于菌絲入侵和病菌繁殖[13-16,19]。核盤菌分泌的草酸是其關(guān)鍵致病因子之一[17]。研究發(fā)現(xiàn)，草酸通過以下方式在核盤菌的致病過程中發(fā)揮作用：①酸化、腐敗寄主組織[20]；②與細(xì)胞壁中膠層的Ca2+結(jié)合形成草酸鈣結(jié)晶損壞細(xì)胞壁結(jié)構(gòu)[21-22]，與Mg2+螯合使葉綠素合成受阻和核糖體功能退化[23]，與二價(jià)金屬離子結(jié)合改變膜透性，導(dǎo)致電解質(zhì)外滲，使細(xì)胞生理機(jī)能退化[24-25]；③影響寄主氧化還原環(huán)境，促進(jìn)寄主的氧化爆發(fā)，破壞寄主體內(nèi)平衡，誘發(fā)細(xì)胞程序性死亡[26-31]；④為病菌分泌水解酶營(yíng)造適宜的pH環(huán)境[15,24-25]，抑制與寄主抗病性相關(guān)的酶[15,32]；⑤刺激保衛(wèi)細(xì)胞對(duì)K+的吸收及對(duì)淀粉的降解，影響其細(xì)胞膜的滲透性，使保衛(wèi)細(xì)胞功能失調(diào)，促使氣孔張開，引發(fā)葉片發(fā)生脫水等癥狀，利于病菌入侵[33]。然而，有學(xué)者發(fā)現(xiàn)了分泌草酸毒素能力強(qiáng)，但致病力卻極弱的核盤菌菌株[34-35]，表明草酸并非核盤菌的惟一致病因子，核盤菌存在更復(fù)雜的致病機(jī)理，尚待進(jìn)行更深入和系統(tǒng)的研究。

2 油菜菌核病抗性資源的篩選及抗病育種

2.1 抗病資源的篩選與鑒定

優(yōu)良的抗源對(duì)作物抗病育種有至關(guān)重要的作用，因此學(xué)者們對(duì)甘藍(lán)型油菜資源進(jìn)行了系統(tǒng)而全面的菌核病抗性鑒定，發(fā)現(xiàn)甘藍(lán)型油菜對(duì)菌核病的抗性存在差異，但并未找到免疫或者高抗材料[36-39]，因此學(xué)者們嘗試在甘藍(lán)型油菜的近緣物種中發(fā)掘優(yōu)良抗源。早在20世紀(jì)末，中國(guó)農(nóng)業(yè)科學(xué)院油料作物研究所等對(duì)2 000余份蕓薹屬材料進(jìn)行了鑒定，發(fā)現(xiàn)芥菜型油菜抗性最強(qiáng)，并選出約6%的高抗種質(zhì)[37]； Li等[38]和Barbetti等[39]報(bào)道了在芥菜型油菜中存在較好的抗病資源，如澳大利亞的BrassicajunceaJM06018、BrassicajunceaJM06006和中國(guó)的Brassicajuncea2等材料。也有學(xué)者報(bào)道蕓薹屬中的埃芥對(duì)菌核病具有很強(qiáng)的抗性[40]。蕓薹屬中的甘藍(lán)，尤其是野生甘藍(lán)擁有很強(qiáng)的菌核病抗性[41-42]；此外，還有學(xué)者在十字花科植物中尋找抗病資源，發(fā)現(xiàn)Erucastrumcardaminoides、Diplotaxistenuisiliqua和Erucastrumabyssinicum等野生型十字花科植物具有高抗菌核病的性能[43]。這些研究盡管均未找到完全免疫的材料，但明確了不同蕓薹屬物種對(duì)菌核病的抗性存在明顯差異，甘藍(lán)型油菜的近緣物種中存在抗性較強(qiáng)的資源，這對(duì)甘藍(lán)型油菜的抗菌核病育種具有較大的應(yīng)用價(jià)值。

2.2 抗菌核病育種

受限于油菜中抗病資源的缺乏，國(guó)內(nèi)育種工作者通過傳統(tǒng)育種方式選育出了一批具有部分抗性的甘藍(lán)型油菜品種，如中雙2號(hào)[44]、中油821[45]、中雙4號(hào)[46]，中雙9號(hào)[47]、湘油11號(hào)和湘油15號(hào)[48]、中雙11號(hào)[49]等；針對(duì)北美油菜種植，先鋒育種公司也培育出了對(duì)菌核病抗性顯著提高的雜交春油菜品種Pioneer Brand?45S51和Pioneer Brand?45S52[50]。雖然有學(xué)者曾采用無(wú)花瓣育種[51]、誘變育種[52]和基因工程育種[53-55]等方式來培育抗菌核病的油菜，但目前尚沒有通過此類途徑獲得抗菌核病品種的報(bào)道。由此看來，要培育高抗菌核病的油菜必須依賴于優(yōu)良抗源的發(fā)掘和先進(jìn)育種技術(shù)的應(yīng)用。

3 油菜菌核病的抗性遺傳規(guī)律

由于沒有找到高抗菌核病的油菜資源，人們只能采用具有部分抗性的油菜來研究菌核病的抗性遺傳規(guī)律。早期研究發(fā)現(xiàn)，甘藍(lán)型油菜BC1F1與BC2F1代具有明顯偏親本較強(qiáng)抗性的特性，其抗性表現(xiàn)為部分顯性[56]。隨后有學(xué)者發(fā)現(xiàn)，油菜菌核病抗性主要受核基因控制，由多個(gè)基因共同作用，抗性遺傳由基因的加性效應(yīng)和非加性效應(yīng)共同控制，且加性遺傳方差大于顯性遺傳方差[57-59]。何昆燕等[60]利用六世代分析模型研究了甘藍(lán)型油菜的菌核病抗性遺傳規(guī)律，發(fā)現(xiàn)抗性受2對(duì)主基因的控制，并受多基因修飾，基因效應(yīng)及其互作隨病情發(fā)展呈加性至顯性的動(dòng)態(tài)變化。近年來對(duì)油菜抗菌核病QTL定位的研究結(jié)果表明，油菜的菌核病抗性為數(shù)量性狀，受多基因共同作用，各抗病位點(diǎn)間以加性效應(yīng)為主，存在一定的上位性，且可能存在與環(huán)境的互作[61]。本研究小組在對(duì)甘藍(lán)的抗病位點(diǎn)定位研究中也發(fā)現(xiàn)了同樣的規(guī)律[62-63]。綜上可知，各研究得出的結(jié)論不盡相同，這預(yù)示著油菜菌核病抗性遺傳機(jī)制的復(fù)雜性，但可以確定的是，油菜菌核病抗性為數(shù)量性狀，其抗性遺傳受多基因控制，并傾向于認(rèn)為加性效應(yīng)在油菜菌核病的抗性遺傳中起著主要作用。

除了缺乏完全免疫或高抗資源外，在分子水平上對(duì)油菜菌核病抗性的不完全闡述也是限制相關(guān)研究的一大因素。為此，學(xué)者們從DNA、RNA以及蛋白質(zhì)水平上對(duì)油菜的抗菌核病基因進(jìn)行了發(fā)掘，以期解析油菜抗菌核病的分子機(jī)制。

3.1 抗病QTL的定位

在DNA水平上，學(xué)者們主要采取QTL定位途徑來尋找抗病相關(guān)基因(表1)。劉春林等[64]采用RAPD標(biāo)記構(gòu)建圖譜與F2單株苗期葉片抗性結(jié)合的方式檢測(cè)到3個(gè)抗病主效QTL，但未能提供該圖譜與甘藍(lán)型油菜各連鎖群的對(duì)應(yīng)關(guān)系，導(dǎo)致其所鑒定的抗病QTL難以與其他類似研究進(jìn)行比較。何昆燕[65]和Yin等[66]各自利用3種不同的鑒定方法對(duì)同一個(gè)甘藍(lán)型油菜DH群體進(jìn)行多年、多環(huán)境下的抗病性鑒定，分別檢測(cè)到18和21個(gè)抗病QTL，但可能由于鑒定方法的不完全相同，兩者鑒定出的QTL重疊較少。Zhao等[2]在甘藍(lán)型油菜F2∶3家系中鑒定出3個(gè)苗期和3個(gè)成株期抗病QTL，但苗期與成株期抗病QTL之間未檢測(cè)到重疊，隨后其又利用葉柄接種法在2個(gè)DH群體中分別獲得11和2個(gè)抗病位點(diǎn)，單個(gè)QTL能解釋的最高表型變異度為22%，并在N12和N02連鎖群上的QTL區(qū)域檢測(cè)到同源區(qū)段[67]。本研究室利用甘藍(lán)型油菜DH群體在N12和N02上鑒定出抗病QTL，并發(fā)現(xiàn)2個(gè)QTL區(qū)域之間存在同源性[68]。近來，一些研究者通過不同的抗性鑒定方法在不同的群體中也鑒定出了一些抗病QTL，這些QTL多數(shù)分布在甘藍(lán)型油菜的A亞基因組上，少數(shù)存在于C亞基因組，其中單個(gè)QTL能解釋的最大變異度達(dá)到42%[67,69-72]。在最近的一篇報(bào)道中，研究者將重要的QTL位點(diǎn)鎖定在了甘藍(lán)型油菜的A9和C6染色體上，通過生物信息學(xué)手段并結(jié)合RT-PCR驗(yàn)證的方法，確定了一個(gè)吲哚硫代甲基轉(zhuǎn)移酶(Indole glucosinolate methyltransferase,IGMT)編碼基因(BnIGMT5.a)，該基因?yàn)镃6上QTL區(qū)間中的候選抗病基因[61]。

表1 甘藍(lán)型油菜的抗菌核病QTLTable 1 Resistant QTLs against Sclerotinia sclerotiorum of Brassica napuse

注：R.抗病，S.感??；A.苗期離體葉片接種法，B.成株期田間牙簽接種法，C.成株期田間菌絲瓊脂塊貼附法，D.活體葉柄接種法，E.花瓣接種法，F(xiàn).草酸浸根法；括號(hào)中的編號(hào)代表該連鎖群在Yin等[66]圖譜中所對(duì)應(yīng)的染色體情況；“-”表示數(shù)據(jù)缺失。

Note:R.Represents resistant,S.Represents susceptible;A.Represents detached leaf inoculation,B.Represents toothpick inoculation,C.Represents mycelial plug inoculation,D.Represents petiole inoculation,E.Represents petal inoculation,F.Represents oxalic-acid soak root inoculation;Numbers in parentheses represent the linkage group in the map of the corresponding chromosomal condition in Yin et al[66];“-” indicates missing data.

3.2 基因表達(dá)層面對(duì)抗病基因的挖掘

在RNA水平上，多數(shù)研究者采用cDNA芯片技術(shù)來研究油菜受核盤菌侵染后誘導(dǎo)表達(dá)的基因，或不同抗性材料在核盤菌侵染后基因的表達(dá)差異。研究發(fā)現(xiàn)，受核盤菌誘導(dǎo)的基因主要包括：①幾丁質(zhì)酶(Chitinase)、葡聚糖酶、滲調(diào)蛋白(Osmotins)和凝集素(Agglutinin)等病程相關(guān)蛋白(Pathogenesis-related proteins,PRs)編碼基因；②細(xì)胞色素P450(Cytochrome P450,CYP450)相關(guān)蛋白基因以及谷胱甘肽轉(zhuǎn)移酶(Glutathione transferase)基因；③茉莉酸(Jasmonic acid,JA)生物合成和信號(hào)傳遞、活性氧代謝(Reactive oxygen metabolism)及氧爆反應(yīng)(Oxidative burst)相關(guān)蛋白基因；④細(xì)胞壁結(jié)構(gòu)與功能相關(guān)基因；⑤蛋白激酶(Protein kinases)、分子轉(zhuǎn)運(yùn)蛋白和細(xì)胞發(fā)育相關(guān)基因等；⑥轉(zhuǎn)錄因子(鋅指、WRKY、AP2和MYB家族)、次生代謝物合成基因等[73-75]。其中，初步確定的候選抗病相關(guān)基因包括：Bn-29、Bn-30和Bn-10(三者一起正調(diào)控油菜菌核病的抗性)[76]，促分裂原活化蛋白激酶(Mitogen-activated protein kinase,MAPK)基因[77-78]、BnMAPK7-1、BnMAPK7-2和BnMAPK7-3[79]，正調(diào)控抗性的乙烯信號(hào)轉(zhuǎn)導(dǎo)途徑(Ethylene-signaling pathway)關(guān)鍵調(diào)節(jié)因子BnEIN3[80]，乙烯應(yīng)答轉(zhuǎn)錄因子(Ethylene-responsive transcriptional factors,ERFs)BnERF104和BnERF50基因[81-82]及多聚半乳糖醛酸酶抑制蛋白(Polygalacturonase-inhibiting protein,PGIP)基因Pgip1和Pgip2[83-84]等。盡管眾多學(xué)者都致力于揭示油菜抗病分子機(jī)制，但受cDNA芯片技術(shù)本身的限制，這些研究無(wú)法在整個(gè)轉(zhuǎn)錄組水平上對(duì)油菜接種核盤菌后基因的表達(dá)變化進(jìn)行系統(tǒng)而全面地跟蹤，因而這些研究類似于一幅拼圖中的少數(shù)幾個(gè)小單元，只有用更多的單元來補(bǔ)充和完善整幅圖片后，才能明確油菜抗核盤菌的分子機(jī)制。

3.3 蛋白質(zhì)層面對(duì)抗病基因的挖掘

在蛋白質(zhì)水平上，早期學(xué)者們主要是通過檢測(cè)少量與植物抗逆/抗病相關(guān)蛋白在接種核盤菌前后的變化情況來判斷其是否與菌核病抗性相關(guān)，如王漢中等[85]研究了苯丙氨酸解氨酶(Phenylalanine ammonia-lyase,PAL)、幾丁質(zhì)酶、β-1,3-葡聚糖酶(β-1,3-dextranase)、過氧化物酶(Peroxidase,POD)和多酚氧化酶(Polyphenol oxidase,PPO)等油菜中重要的防御酶類在抗、感材料接種前后的活性變化，發(fā)現(xiàn)PAL、幾丁質(zhì)酶和β-1,3-葡聚糖酶、POD、PPO與油菜對(duì)菌核病的抗性密切相關(guān)；張志元等[86]發(fā)現(xiàn)，超氧物歧化酶(Superoxide dismutase,SOD)與油菜對(duì)菌核病的抗性有關(guān)。后來學(xué)者們開始采用蛋白質(zhì)組學(xué)來研究油菜抗菌核病的機(jī)制和抗病基因?；矢Ｑ嗟萚84]對(duì)油菜抗菌核病近等基因系及其感病親本在核盤菌侵染后的蛋白質(zhì)組進(jìn)行了分析，發(fā)現(xiàn)組成線粒體ATP合酶F1的重要亞基F1β、糖酵解關(guān)鍵酶磷酸甘油酸激酶(Phosphoglycerate kinase,PGK)、參與合成維生素B1的輔酶噻唑(thiazole)生物合成酶THI1和1,5-二磷酸核酮糖羧化酶/加氧酶(Ribulose-1,5-bisphosphate carboxylase/oxygenas,RuBisCO)在油菜對(duì)菌核病的抗性反應(yīng)中差異表達(dá)。Wen等[87]對(duì)甘藍(lán)型油菜抗、感品種接種核盤菌前后進(jìn)行了比較蛋白組學(xué)分析，發(fā)現(xiàn)與抗性相關(guān)的蛋白主要在病原抗性、抗氧化作用和轉(zhuǎn)錄調(diào)節(jié)這幾個(gè)部分發(fā)生作用，并發(fā)現(xiàn)一些作用于活性氧清除的胰蛋白酶抑制蛋白(Trypsin inhibitor protein,TIP)、富含甘氨酸的細(xì)胞壁蛋白(Glycine rich protein,GRP)、熱激蛋白(Heat shock protein,HSP)和硫醇甲基轉(zhuǎn)移酶(Thiol methyltransferase,TMT)在抗性品種中呈上調(diào)或特異表達(dá)。

4 油菜菌核病的研究展望

盡管科學(xué)家們對(duì)油菜抗菌核病育種及相關(guān)研究投入了較大精力，也在取得了一定成效，但還存在一些重要問題需要解決：一是油菜中缺乏可利用的優(yōu)良抗源，導(dǎo)致抗性位點(diǎn)定位的重復(fù)性差，抗病基因鑒定工作進(jìn)展緩慢，其研究結(jié)果難以應(yīng)用于育種實(shí)踐，油菜菌核病抗性的改良難以獲得突破性進(jìn)展；二是寄主與核盤菌的互作機(jī)制復(fù)雜，病原菌的致病機(jī)理、寄主的抗病機(jī)制至今仍不清晰，從而限制了抗病基因的鑒定和油菜抗菌核病的分子設(shè)計(jì)育種；三是研究手段尚需改進(jìn)，當(dāng)前對(duì)抗病基因的鑒定研究多采用QTL定位或通過檢驗(yàn)少數(shù)基因?qū)吮P菌的響應(yīng)情況來推測(cè)其與抗性的關(guān)系，難以準(zhǔn)確鑒定出抗病基因，更無(wú)法了解寄主的抗病機(jī)制。

鑒于上述問題，筆者認(rèn)為油菜菌核病研究的首要內(nèi)容仍然是廣泛搜尋可利用的優(yōu)良抗病資源，包括油菜的近緣物種乃至十字花科其他植物。在獲得優(yōu)良抗源的基礎(chǔ)上，通過遠(yuǎn)緣雜交或傳統(tǒng)育種方式將其抗病成分轉(zhuǎn)入油菜，利用其進(jìn)行抗病位點(diǎn)鑒定的準(zhǔn)確性必將得到有效提高，抗病機(jī)制的分析難度也將降低。隨著生物學(xué)和生物信息學(xué)研究技術(shù)的快速發(fā)展，以及甘藍(lán)型油菜、甘藍(lán)和白菜全基因組測(cè)序的進(jìn)行和完成，結(jié)合全基因組水平、轉(zhuǎn)錄組水平和代謝組水平的分析手段，可快速、全面、準(zhǔn)確地解析油菜的抗菌核病機(jī)制并發(fā)掘抗病基因，這對(duì)于油菜抗菌核病的分子設(shè)計(jì)育種具有重要的實(shí)踐意義。

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Advance inSclerotiniastem rot of rapeseed

WANG Lei,LIU Yao,DING Yi-juan,WANG Yu,WAN Hua-fang,MEI Jia-qin,QIAN Wei

(SchoolofAgronomyandBiotechnology,SouthwestUniversity,ChongqingRapeseedEngineering&TechnologyResearchCenter,Beibei,Chongqing400715,China)

Sclerotiniastem rot caused by fungal pathogenSclerotiniasclerotiorumis one of the most devastating diseases in rapeseed production.Combining with the existing research literature and the findings of our lab,the review summarized and prospected the researches in infection process and pathogenesis mechanism ofS.sclerotiorum,disease-resistant resources,breeding progress,inheritance of resistance,resistant QTL loci,and resistance genes in rapeseed.

rapeseed;Sclerotiniastem rot; pathogenesis;molecular mechanism;resistance breeding

時(shí)間:2015-09-09 15:41

10.13207/j.cnki.jnwafu.2015.10.012

2014-03-05

中央高?；緲I(yè)務(wù)費(fèi)項(xiàng)目(SWU113106)；國(guó)家自然科學(xué)基金項(xiàng)目(31171585)

汪 雷(1991-)，女，重慶銅梁人 ，碩士，主要從事作物遺傳育種研究。E-mail:wangleicqtl@163.com

梅家琴(1982-)，女，四川彭州人，副教授，主要從事植物分子育種及蕓薹屬物種菌核病抗性研究。 E-mail:jiaqinmay@163.com

S332.2

A

1671-9387(2015)10-0085-09

網(wǎng)絡(luò)出版地址:http://www.cnki.net/kcms/detail/61.1390.S.20150909.1541.024.html