李 芳，張 峰，高 萍，段廷玉

(草地農(nóng)業(yè)生態(tài)系統(tǒng)國家重點實驗室，蘭州大學草地農(nóng)業(yè)科技學院，甘肅 蘭州 730020)

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農(nóng)業(yè)生態(tài)系統(tǒng)中AM真菌、禾草內(nèi)生真菌及病原菌互作

李 芳，張 峰，高 萍，段廷玉

(草地農(nóng)業(yè)生態(tài)系統(tǒng)國家重點實驗室，蘭州大學草地農(nóng)業(yè)科技學院，甘肅 蘭州 730020)

摘要：叢枝菌根真菌(Arbuscular Mycorrhizal Fungi，AMF)、禾草內(nèi)生真菌和植物病原菌廣泛存在于農(nóng)業(yè)生態(tài)系統(tǒng)中。AMF和禾草內(nèi)生真菌均能與宿主植物形成共生體，促進宿主植物對養(yǎng)分、水分的吸收，提高植物對病害、干旱等生物和非生物逆境的抗性。植物病原菌引致的植物病害可降低植物產(chǎn)量和品質(zhì)，造成生產(chǎn)上的重大損失。AMF可降低植物發(fā)病率和病情指數(shù)，其機理包括AMF與病原菌競爭侵染位點，養(yǎng)分和空間、消耗病原菌的能量、提高菌根植物對水分和養(yǎng)分的利用效率、調(diào)節(jié)病程相關(guān)蛋白。禾草內(nèi)生真菌可通過分泌抑菌活性物質(zhì)、誘導植物體產(chǎn)生抗性反應(yīng)及減少病害介體的傳播，提高植物地上生物量和抗病性。AMF和禾草內(nèi)生真菌與其它微生物的互作因植物、真菌而異，有相互促進的效應(yīng)，也有相互抑制的效應(yīng)。研究和明確農(nóng)業(yè)生態(tài)系統(tǒng)中三類微生物的互作機理，進而利用禾草內(nèi)生真菌和AMF提高抗逆性及產(chǎn)量、防治病害，對于促進和維持農(nóng)業(yè)生態(tài)系統(tǒng)可持續(xù)發(fā)展具有重要的意義。

關(guān)鍵詞：叢枝菌根真菌；禾草內(nèi)生真菌；植物病原菌；互作

微生物是農(nóng)業(yè)生態(tài)系統(tǒng)中的重要成員，有著不可替代的作用。作為生態(tài)系統(tǒng)中的重要分解者，參與C、N、P、S等養(yǎng)分循環(huán)[1]，或與植物建立共生關(guān)系，如叢枝菌根(Arbuscular Mycorrhiza，AM)真菌和禾草內(nèi)生真菌(Epichlo?)，兩類共生微生物均可提高植物對N、P、K等養(yǎng)分[2-4]、水分[5-6]的吸收和利用效率，提高植物對病[7-8]、蟲[8-9]等生物逆境以及干旱[8,10]、鹽堿[11-13]、低溫[14-15]和重金屬[16-17]等非生物逆境的抗性，從而促進植物的生長，提高植物在農(nóng)業(yè)生態(tài)系統(tǒng)中的競爭力，進而維持農(nóng)業(yè)生態(tài)系統(tǒng)的穩(wěn)定性和可持續(xù)性[18]。

同時，微生物亦可作為消費者，如，病原菌(病原真菌、細菌、病毒和線蟲等)可引致植物病害，降低植物產(chǎn)量，改變植被組成[19]，導致家畜中毒，造成重大經(jīng)濟損失。以天然草地為例， 2013年，我國天然草地鮮草總產(chǎn)量105 581.21萬t，折合干草約32 542.92萬t，以病害造成干草損失5%[19]計算，可造成干草產(chǎn)量損失1 627.15萬t，以每公斤干草0.2元計算，造成經(jīng)濟損失32.54億元。

微生物與植物在長期協(xié)同進化過程中形成了共生、共棲和寄生等多種復雜關(guān)系，如何利用農(nóng)業(yè)生態(tài)系統(tǒng)中的微生物與植物、微生物與微生物之間的關(guān)系，降低病害的發(fā)生和危害，提高農(nóng)業(yè)生態(tài)系統(tǒng)的生產(chǎn)力和可持續(xù)性是目前國際上研究的熱點[18]。

本文就近年來國內(nèi)外有關(guān)AM真菌、禾草內(nèi)生真菌與病原菌的互作進行綜述，以期為利用AM真菌和禾草內(nèi)生真菌來防治植物病害，提高和維持農(nóng)業(yè)生態(tài)系統(tǒng)生產(chǎn)力和可持續(xù)發(fā)展，為開展AM真菌、禾草內(nèi)生真菌和病原菌三者之間的互作研究提供理論依據(jù)。

1AM真菌與病原菌

AM真菌屬球囊菌門，可與約90%的維管植物形成共生體[20]，廣泛存在于農(nóng)業(yè)生態(tài)系統(tǒng)中。AM真菌與病原菌的互作關(guān)系較為復雜，有研究表明,AM真菌對植物具有一定的保護作用[21]、可減輕病害[22]，還有研究發(fā)現(xiàn)AM真菌與植物抗病性無關(guān)[23]，但也有AM真菌能加重植物病害的報道[24]。就目前的研究而言，有關(guān)抗病的報道居多[25-31]。

1.1AM真菌與病原真菌的互作

1968年Safir[32]首次報道了摩西球囊霉(Glomusmosseae)能減少洋蔥紅根腐菌(Pyrenochaetaterestris)對洋蔥根系的侵染并能減輕紅腐病(Red root)的危害，提高植物抗病性。此后，關(guān)于二者相互作用的關(guān)系被大量報道，但大部分研究僅限于球囊霉屬的幾個種，如摩西球囊霉、幼套球囊霉(Glomusetunicatum)、根內(nèi)球囊霉(Glomusintraradices)、地表球囊霉(Glomusversiforme)等。該屬AM真菌均可以不同程度地降低立枯絲核菌(Rhizoctoniasolani)、尖孢鐮刀菌(Fusairumoxysporum)、黃萎病原菌(Verticilliumdahliae)等引起的植物病害，發(fā)病率和病情指數(shù)下降[33-36]。隨著近年來栽培草地面積的不斷擴大，根腐病逐漸成為限制牧草生產(chǎn)的主要病害，僅在紫花苜蓿(Medicagosativa)上，就有鐮刀菌(Fusariumspp.)、絲核菌(Rhizoctoniaspp.)、腐霉(Pythiumspp.)和疫霉(Phytophthoraspp.)等病原菌可引致根腐病[37]，但之前AM真菌與病原真菌互作的研究多見于農(nóng)作物和蔬菜等(表1)，在牧草上的研究相對較少，以后應(yīng)加大在此方面的研究。

1.2AM真菌與病原菌互作機理

AM真菌可以促進植物對養(yǎng)分、水分的吸收和利用，從而提高植物的生長，這是其提高植物抗病性最主要的功能，但亦存在其它抗病機制，主要包括：1)菌根表面本身分布著大量菌絲體，對病原物構(gòu)成天然的機械屏障，再加上AM真菌的侵染增強細胞壁木質(zhì)化程度，增加細胞壁富含羥基脯氨酸糖蛋白(Hydroxyproline-rich Glycoprotein)，使植物根尖表皮增厚，從而阻止病原物的侵入[44]；2)AM真菌能顯著改變植物根圍的微生物區(qū)系，刺激對土傳病原物(Soil-borne Pathogens)有拮抗作用的微生物活性，使促進植物生長的細菌[如熒光假單胞菌(Pseudomonasfluorescens)、芽孢桿菌(Bacillus)和其它拮抗菌木霉屬(Trichodermaspp.)、鏈霉菌(Streptomycesspp.)]數(shù)量增加[45]。這些有益菌的協(xié)同作用有利于降低植物土傳病害的危害[46]；3)AM真菌能誘導植物合成茉莉酸等物質(zhì)[47]產(chǎn)生或激活幾丁質(zhì)酶[48]、β-1，3葡聚糖酶[49]、苯丙氨酸裂解酶[50]等保護酶，從而明顯增強了植物抗病性；4)AM真菌可以和病原菌競爭侵染位點[29-30,51-53]、養(yǎng)分和空間[54]。在苗床上聚生球囊霉(G.fasciculatum)接種顯著限制了腐霉菌(Pythiumaphanidermatum)在沉香(Aquilariaagallocha)根系組織中的侵染，這正是由于叢枝菌根真菌先侵染根系位點所致[55]；5)AM真菌侵染感病寄主植物時，可以消耗病原菌的光合產(chǎn)物[52]，從而使病原菌的致病能力下降；6)AM真菌根外菌絲體相互交錯形成龐大的菌絲網(wǎng)，促進植物對水分和養(yǎng)分，特別是磷酸鹽和硝酸鹽的吸收利用[25,27]，通過對營養(yǎng)物質(zhì)和水分的吸收，補償了因病原菌侵染造成的根系生物量的損失，從而間接減輕病原微生物引起的危害，提高宿主植物的抗病能力[45]；7)AM真菌可以誘導病程相關(guān)蛋白，提高宿主對病原物的抵抗力[56]。研究發(fā)現(xiàn)，當病原菌侵染菌根化寄主植物時，AM真菌能誘導PAL5基因和幾丁質(zhì)酶基因Chib1等[57]，或通過調(diào)控各種抗病基因的表達量及特異性表達來增強宿主植物的抗病性[58]。此外，Gallou等[59]通過實時定量PCR分析發(fā)現(xiàn)，菌根化馬鈴薯(Solanumtuberosum)葉片的疫霉菌(P.infestans)病情指數(shù)和病害癥狀之所以減輕，可能和菌根化馬鈴薯中與植株系統(tǒng)抗性相關(guān)的PR1和PR2基因的誘導表達有關(guān)。

表1　AM真菌與病原真菌的互作效應(yīng)

續(xù)表1

AM真菌Arbuscularmycorrhizalfungi病原真菌Pathogenicfungi病害名稱Disease寄主植物Host互作結(jié)果Interactioneffect試驗條件Experimentalconditions參考文獻Reference幼套球囊霉G.etunicatum、根內(nèi)球囊霉G.intraradices立枯絲核菌R.solani菌核病Sclerotina甘薯Ipomoeabatatas病情指數(shù)下降Reduceddiseaseindex盆栽Pot[42]巨孢囊霉G.rosea、摩西球囊霉G.mosseae、地表球囊霉G.versiforme尖鐮孢菌F.oxysporum枯萎病Fusariumwilt西瓜Citrulluslanatus抑制病原菌的生長和侵染32.7%~53.7%和27.7%~35.1%Restrainpathogenicgrowthandinfection32.7%~53.7%and27.7%~35.1%,respectively大田Field[43]

2禾草內(nèi)生真菌與病原菌的互作

本文所指內(nèi)生真菌為寄生在植株體內(nèi)完成全部或大部分生活周期，而植物體不顯示外部癥狀的一類真菌，無性階段為Neotyphodium屬，有性階段為香柱菌屬(Epichlo?)真菌[60],目前眾多學者一致認為將Neotyphodium屬真菌并入Epichlo?屬，其菌絲體存在于植物的所有地上部分：莖桿、葉鞘、葉片、花序及種子，但未發(fā)現(xiàn)存在于根系中[8]。全世界已發(fā)現(xiàn)43種，我國已報道7種Epichlo?內(nèi)生真菌[61-62]。

2.1禾草內(nèi)生真菌與病原真菌的互作

從1983年Shimanuki和Sato[63]發(fā)現(xiàn)含有內(nèi)生真菌(Epichlo?typhina)的梯牧草(Phleumpratense)對引致葉斑病的病原真菌(Cladosporiumphlei)有抗性開始，相繼有很多試驗均證明禾草內(nèi)生真菌能夠提高植物的抗病性。田沛[64]發(fā)現(xiàn),內(nèi)生真菌對細交鏈孢、離蠕孢、燕麥鏈孢、新月彎孢、小孢殼二孢等10種病原真菌均有抗病性。高羊茅(Festucaarundinacea)、黑麥草(Loliumperenne)、羊草(Leymuschinensis)等植物的內(nèi)生真菌也能不同程度地降低寄主植物發(fā)病率、病情指數(shù)、病斑數(shù)和植株矮化程度，提高防御性保護酶活性、病葉損失率，增強抗病性，促進植物生長。在PDA培養(yǎng)基上，進行內(nèi)生真菌和病原真菌競爭試驗，發(fā)現(xiàn)內(nèi)生真菌對病原真菌的生長表現(xiàn)出明顯的抑制作用[65-68](表2)。

目前，美國和新西蘭已經(jīng)利用禾草內(nèi)生真菌培育出了對害蟲、干旱等具有抗性的品種，并被大量推廣、應(yīng)用[69]，但專門針對抗病性的品種，尚未有報道。

2.2禾草內(nèi)生真菌提高植物抗病性的機理

禾草內(nèi)生真菌與AMF一樣，可促進植物生長，提高植物抗逆性，這應(yīng)是該菌提高植物抗病性的機理之一。關(guān)于禾草內(nèi)生真菌的抗病機制，存在以下幾種可能：1)禾草內(nèi)生真菌與病原菌之間存在空間、養(yǎng)分和水分的競爭[8,70-74]。一方面，禾草內(nèi)生真菌菌絲體在植物葉片、葉鞘等部位的存在，形成菌絲防護網(wǎng)，占據(jù)了一定的空間和生態(tài)位，從而抵抗病原菌的侵入和定殖[8]；另一方面，禾草內(nèi)生真菌從寄主植物獲得營養(yǎng),使病害真菌侵染時可獲得的水分、養(yǎng)分減少，從而降低植株發(fā)病程度[70]；2)禾草內(nèi)生真菌能夠分泌抑制病原菌生長的活性物質(zhì)，從而降低病原真菌的繁殖和進一步擴展[71-73]。此外，禾草內(nèi)生真菌共生體產(chǎn)生的化學物質(zhì)可以作為殺菌劑抑制或殺死一些引起病害的病原真菌、細菌、病毒和原生動物[74]；3)當病原真菌入侵時，禾草內(nèi)生真菌能快速誘導宿主植物產(chǎn)生幾丁質(zhì)酶、β-1, 3-葡聚糖酶和過氧化物酶、病程相關(guān)蛋白、以及抗病相關(guān)物質(zhì)的生成，提高感染內(nèi)生真菌的禾草中酚類物質(zhì)和木質(zhì)素的含量等[75-76]；4)減少病害介體的傳播。有些病害會通過昆蟲介體傳播，而禾草內(nèi)生真菌能夠產(chǎn)生毒素，阻抑帶有病毒和真菌的昆蟲采食[77-78]，從而阻止了介體昆蟲傳播病毒，減輕了病毒病的發(fā)生。Mahmood等[79]發(fā)現(xiàn)E+高羊茅的大麥黃矮病毒(BYDVs)發(fā)病率低主要是由于其內(nèi)生真菌阻抑了大麥黃矮病毒介體的蚜蟲的采食率；5)禾草內(nèi)生真菌可以改變植物的營養(yǎng)，促進植物生長，從而強化寄主植物的能力，間接增加植物對病害的抗性[80]；6)禾草內(nèi)生真菌能誘導寄主植物本身產(chǎn)生系統(tǒng)抗性反應(yīng)，從而抑制病原菌的入侵，而植物本身的基因并未發(fā)生改變，仍然可以保持天然性狀[81]。

表2　禾草內(nèi)生真菌與病原真菌的互作結(jié)果

3AM真菌與禾草內(nèi)生真菌的互作

農(nóng)業(yè)生態(tài)系統(tǒng)中存在著豐富的微生物類群，因此在植物組織中，這些微生物也必然存在復雜的雙共生，甚至三共生的相互作用。Porras-Alfaro等[82]利用免培(Culture Independent)環(huán)境PCR方法，通過真菌特異性引物擴增發(fā)現(xiàn)，植株體內(nèi)存在豐富多樣的真菌類群，包括菌根真菌和內(nèi)生真菌，因此,植物同時形成兩種共生現(xiàn)象在農(nóng)業(yè)生態(tài)系統(tǒng)中十分普遍，研究菌根真菌、內(nèi)生真菌與植物的雙共生可以真實反映植物的微生態(tài)系統(tǒng)[75-84]。

禾草內(nèi)生真菌和土壤中的AM真菌可以相互影響。研究發(fā)現(xiàn),禾草內(nèi)生真菌和AM真菌存在相互競爭的關(guān)系，且?guī)?nèi)生真菌的禾草植物AM真菌侵染率會降低[85-87]，而Novas等[88]卻發(fā)現(xiàn),禾草內(nèi)生真菌可以增加AM真菌的侵染率，而且在一定濃度內(nèi)生真菌浸提液作用下可以增加AM真菌菌絲長度。Liu等[89]報道禾草內(nèi)生真菌降低AM真菌侵染率的同時，AM真菌反過來又減少了內(nèi)生真菌的菌絲密度和寄主植物葉片中生物堿的含量(生物堿是禾草內(nèi)生真菌提高宿主抗性的主要原因)；Vicari等[84]和Barker[90]的研究發(fā)現(xiàn)，帶內(nèi)生真菌Epichlo?festucae的黑麥草被摩西球囊霉(G.mosseae)和聚生球囊霉(G.fasciculatum)侵染后抗蟲性降低。也有研究發(fā)現(xiàn)，內(nèi)生真菌Epichlo?elymi可以促進摩西球囊霉侵染，抑制近明球囊霉(G.claroideum)的侵染[91]，即摩西球囊霉和內(nèi)生真菌E.elymi共同作用于寄主植物時，對植物的生長效應(yīng)是疊加的。研究表明，在無脅迫條件下，禾草內(nèi)生真菌和菌根真菌均可顯著增加野大麥(Hordeumbrevisubulatum)的地上生物量和總生物量，而菌根真菌可顯著增加野大麥的地下生物量，禾草內(nèi)生真菌的存在可顯著抑制叢枝菌根真菌的侵染，抑制率為41.67%；高磷、低磷和無磷水平下，禾草內(nèi)生真菌處理顯著提高了野大麥的株高和分蘗，兩種共生體共存時，地上、地下生物量顯著大于其它水平的處理。在無磷條件下，有AM真菌處理的地下生物量顯著高于其它處理。在低磷水平下，禾草內(nèi)生真菌的處理和AM真菌的處理對野大麥氮、磷累積有顯著的促進作用[92-93]。

綜上所述，關(guān)于AM真菌與禾草內(nèi)生真菌的互作，目前主要有兩種觀點：一種觀點認為，禾草內(nèi)生真菌與AM真菌互作對植物體具有正效應(yīng)，禾草內(nèi)生真菌感染的牧草能提高叢枝菌根真菌的侵染率[88]。另一種觀點則認為，菌根真菌和禾草內(nèi)生真菌有相互抑制效應(yīng)：內(nèi)生真菌的存在顯著減少菌根真菌的侵染[85-87]，而AM真菌侵染可以減少禾草內(nèi)生真菌菌絲密度和寄主植物葉片中生物堿含量，二者的互作結(jié)果可能與土壤營養(yǎng)水平、內(nèi)生真菌菌株類型和AM真菌類型等因素有關(guān)[89]。探究二者互作的內(nèi)在機制，明確植物是如何同時與不同的微生物建立共生關(guān)系，并從中找到平衡點將對未來的實際應(yīng)用有指導意義。

4問題與展望

目前，國內(nèi)外有關(guān)AM真菌、禾草內(nèi)生真菌和植物病原菌兩者之間互作關(guān)系的研究已取得一定的成果，且在生產(chǎn)上開始應(yīng)用，如美國、澳大利亞、日本等國家，已開始利用AM真菌菌劑，提高作物生產(chǎn)；美國和新西蘭已培育出含內(nèi)生真菌的黑麥草品種，且已投放市場[68]。而關(guān)于這3類微生物共同作用關(guān)系的研究尚未見報道。

隨著社會的不斷發(fā)展，人民生活水平逐漸提高，對食品安全的要求也不斷增長。但在城市化和工業(yè)化不斷推進的今天，生態(tài)環(huán)境在不斷惡化，食品安全也存在較多隱患，而且大面積種植同一品種植物，可能導致植物病害的爆發(fā)和流行，因此必須密切關(guān)注植物病害的監(jiān)測和防控。農(nóng)業(yè)生態(tài)系統(tǒng)中蘊藏著豐富的植物、微生物資源。鑒于AM真菌與禾草內(nèi)生真菌對于植物病原菌的顯著防效，有必要研究農(nóng)業(yè)生態(tài)系統(tǒng)中這3類重要微生物的互作機理，并在此基礎(chǔ)上，利用禾草內(nèi)生真菌和AM真菌提高植物抗逆性及產(chǎn)量、防治植物病害、促進農(nóng)業(yè)生態(tài)系統(tǒng)可持續(xù)發(fā)展。

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(責任編輯張瑾)

Progress of research on the interactions of arbuscular mycorrhizal fungi,grass endophyte and plant pathogens in agro-ecosystem

Li Fang, Zhang Feng, Gao Ping, Duan Ting-yu

(State Key Laboratory of Grassland Agro-ecosystems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou 730020, China)

Abstract:Arbuscular myucorrhizal fungi (AMF), grass endophyte and plant pathogens are widely distributed in agro-ecosystems. Both AMF and grass endophyte could establish symbiosis with plants which could improve plant nutrient uptake, water absorption and enhance plant stress resistance, such as disease and drought stress. AMF could reduce plant disease incidence and disease index by a series of mechanisms including competing invading sites, nutrition and space with plant pathogens, consuming plant pathogens energy, improving plant water and nutrition efficiency and adjusting plant disease related proteins. Grass endophyte also could decrease plant disease incidence and the possible mechanism including excreting antimicrobial metabolites, inducing plant resistance and inhibiting medium of disease spread. The interactions of AMF and grass endophyte depended on plants and fungi which are mutual or antagonism. Research and utilization of AMF and grass endophyte are very important for improving and maintaining agro-ecosystem productivity and sustainability.

Key words:arbuscular mycorrhizal fungi; grass endophyte; plant pathogen; interaction

Corresponding author:Duan Ting-yuE-mail:duanty@lzu.edu.cn

中圖分類號：S181.6;S432.4+4

文獻標識碼：A

文章編號：1001-0629(2016)2-0219-11

通信作者：段廷玉(1976-)，男，甘肅靖遠人，副教授，博士，研究方向為菌根生態(tài)學。E-mail:duanty@lzu.edu.cn

基金項目：甘肅省自然科學基金項目(1208RJZA270)；國家自然科學基金項目(31100368/C0309)；教育部博士點基金新青年教師項目(20110211120032)

*收稿日期：2015-03-18接受日期：2015-07-24

DOI:10.11829/j.issn.1001-0629.2015-0151

李芳，張峰，高萍，段廷玉.農(nóng)業(yè)生態(tài)系統(tǒng)中AM真菌、禾草內(nèi)生真菌及病原菌互作.草業(yè)科學,2016,33(2)：219-229.

Li F,Zhang F,Gao P,Duan T Y.Progress of research on the interactions of arbuscular mycorrhizal fungi, grass endophyte and plant pathogens in agro-ecosystem.Pratacultural Science，2016,33(2)：219-229.

第一作者：李芳(1990-)，女，青海西寧人，在讀碩士生，研究方向為菌根生態(tài)學。E-mail:lif2013@lzu.edu.cn