蓋云鵬， 潘汝謙， 關(guān)銘芳， 徐大高

(1.山西侯馬出入境檢驗(yàn)檢疫局，侯馬 043000；2.華南農(nóng)業(yè)大學(xué)資源環(huán)境學(xué)院植物病理學(xué)系，廣州 510642)

基礎(chǔ)知識(shí)

一種值得關(guān)注的大豆病害
——大豆紅冠腐病

蓋云鵬1，2*， 潘汝謙2， 關(guān)銘芳2， 徐大高2

(1.山西侯馬出入境檢驗(yàn)檢疫局，侯馬 043000；2.華南農(nóng)業(yè)大學(xué)資源環(huán)境學(xué)院植物病理學(xué)系，廣州 510642)

大豆紅冠腐病是一種為害大豆生產(chǎn)的重要病害，該病害的典型癥狀是大豆葉片變黃萎蔫，植株枯萎，根系變黑腐爛，莖基部及根系表面產(chǎn)生大量紅橙色小點(diǎn)，是病原菌的子囊殼。其致病菌為寄生柱枝孢菌(Cylindrocladiumparasiticum)，有性階段為冬青麗赤殼菌(Calonectriailicicola)。本文對(duì)大豆紅冠腐病及其致病菌的文獻(xiàn)進(jìn)行了簡(jiǎn)要的綜述。

大豆紅冠腐??； 寄生柱枝孢菌； 研究進(jìn)展

1 發(fā)生與分布

大豆紅冠腐病(red crown rot of soybean)是一種危害大豆(GlycinemaxLinn.)的重要病害，在環(huán)境有利于病害發(fā)生條件下可導(dǎo)致大豆減產(chǎn)。1968年在日本千葉縣首次報(bào)道了大豆紅冠腐病[1]。至2001年，包括北海道在內(nèi)，大豆紅冠腐病在全日本各地區(qū)廣泛發(fā)生，且危害嚴(yán)重，成為日本大豆的重要病害之一[2-3]。當(dāng)時(shí)該病害被定名為大豆黑根腐病(black root rot of soybean)，病原菌被描述為豬屎豆柱枝孢菌[Cylindrocladiumcrotalariae(Loos) Bell & Sobers]，有性階段為豬屎豆麗赤殼菌[Calonectriacrotalariae(Loos) Bell & Sobers][4]。1993年，Crous等將豬屎豆柱枝孢菌更名為寄生柱枝孢菌(C.parasiticumCrous, Wingfield & Alfenas)，隸屬于半知菌門(mén)、絲孢綱、絲孢目、絲孢科、柱枝孢屬；Crous等將豬屎豆柱枝孢菌有性態(tài)更改為冬青麗赤殼菌(C.ilicicolaBoedijn & Reitsma)，隸屬于子囊菌門(mén)、糞殼菌綱、肉座菌目、叢赤殼科、麗赤殼屬[5]。

目前，大豆紅冠腐病主要分布于美國(guó)、日本、中國(guó)、韓國(guó)[6]、喀麥隆[7]等多個(gè)國(guó)家和地區(qū)。在美國(guó)，大豆紅冠腐病主要分布在路易斯安那州和密西西比州，為該地區(qū)重要的大豆病害[8-9]。在我國(guó)江蘇省、云南省和廣東省的一些地區(qū)已報(bào)道大豆紅冠腐病的發(fā)生[10-12]。在廣東省，大豆紅冠腐病主要分布在博羅縣和梅州市等地。夏大豆結(jié)莢期最感病，且發(fā)病較普遍和嚴(yán)重，使大豆產(chǎn)量和品質(zhì)降低[13]。

2 典型癥狀

大豆紅冠腐病通常發(fā)生在大豆結(jié)莢期之后，在田間形成明顯的發(fā)病中心[6]。發(fā)病初期，罹病植株葉片葉脈間變黃，隨即萎蔫、落葉，植株枯萎。病株根系及近地面的莖基部變紅褐色，沿莖干向上擴(kuò)展，近地面莖皮層腐爛，嚴(yán)重的甚至木質(zhì)部組織也變褐色腐爛；拔起植株，可見(jiàn)根系變黑腐爛；植株莖基部的病部表面有大量紅橙色球狀子囊殼聚生。莖基部變紅褐色和紅橙色子囊殼為該病害診斷的典型特征；發(fā)病后期，整個(gè)植株根系腐爛，最終植株死亡[10-12]。

3 病原菌生物學(xué)特性

大豆紅冠腐病菌在PDA培養(yǎng)基上生長(zhǎng)迅速，菌落白色至淺灰色或紅褐色，絨毛狀。分生孢子梗直立，細(xì)長(zhǎng)，呈掃帚狀，簡(jiǎn)單分支，具有初生孢子梗和次生孢子梗，頂端為產(chǎn)孢細(xì)胞，瓶狀，分生孢子從產(chǎn)孢細(xì)胞內(nèi)生出。分生孢子圓柱形，無(wú)色，(54.1～76.3)μm×(4.9～7.4)μm，1～3個(gè)分隔。菌絲末端膨大，產(chǎn)生球形泡囊，直徑為(4.0～13.0)μm。厚垣孢子褐色，成串聚集，形成深褐色微菌核(microsclerotia)，(33.3～311.1)μm×(22.2～133.3)μm。病原菌為同宗配合，子囊殼紅色至橙紅色、橢圓形至倒卵形，表面粗糙，有孔口，單生， (212.1～454.5)μm×(111.1～333.3)μm。輕壓子囊殼后可見(jiàn)大量子囊噴出，子囊具長(zhǎng)柄，棍棒形，(121.0～200.8)μm×(11.5～25.6)μm，子囊內(nèi)有8個(gè)鐮刀形的具1～3個(gè)隔膜的子囊孢子，子囊孢子線形，大小為(29.5～73.8)μm×(4.9～9.8)μm[10-12]。菌絲的最適生長(zhǎng)溫度為26～28 ℃，最低為8 ℃，高于35 ℃時(shí)菌絲不生長(zhǎng)。微菌核形成的最適溫度為24～28 ℃，低于12 ℃和高于32 ℃均不形成微菌核[14]。分生孢子產(chǎn)生的最適溫度為28 ℃。28～30 ℃的較高溫度容易產(chǎn)生子囊殼。適合菌絲生長(zhǎng)的固體培養(yǎng)基有馬鈴薯葡萄糖瓊脂培養(yǎng)基、酵母蛋白胨培養(yǎng)基、合成真菌培養(yǎng)基、V8汁培養(yǎng)基、燕麥瓊脂培養(yǎng)基[15]。

4 寄主范圍

大豆紅冠腐病菌的寄主十分廣泛，可侵染經(jīng)濟(jì)作物、觀賞植物、林木及雜草在內(nèi)的20多種植物。除了侵染大豆引起大豆紅冠腐病外，也可侵染花生(ArachishypogaeaL.)引起花生黑腐病(Cylindrocladiumblack rot of peanut)。1965年Bell等在美國(guó)佐治亞州首次發(fā)現(xiàn)花生黑腐病，隨后，該病相繼在佐治亞州[16-17]、北卡羅來(lái)納州[18]、南卡羅來(lái)納州、佛羅里達(dá)州和德克薩斯州[19]發(fā)生，是美國(guó)境內(nèi)危害花生的一種重要病害[20]。該病原菌還可侵染高灌藍(lán)莓(VacciniumcorymbosumL.)引起藍(lán)莓莖腐和根腐病(stem and root rot of blueberry)[21]；侵染番木瓜(CaricapapayaL.)引起番木瓜莖腐病(collar rot of papaya)[22]；侵染夏威夷寇阿相思木(AcaciakoaGray)引起根莖腐病(collar root rot of koa)[23]；侵染北美楓香(LiquidambarstyracifluaL.)引起苗期根腐病(Cylindrocladiumroot rot of sweetgum seedling)[24]；侵染紅葉火筒樹(shù)(LeeacoccineaPlanch)引起莖腐和葉斑病(collar rot and leaf spot of leea)[25]；侵染夾竹桃(NeriumoleanderL.)引起夾竹桃基腐病(basal stem rot of oleander)[26]；侵染中華獼猴桃(ActinidiachinensisPlanch)引起獼猴桃根腐病(Cylindrocladiumroot rot of kiwifruit)[27]；侵染南美山螞蝗[Desmodiumtortuosum(Sw.) DC]引起南美山螞蝗黑腐病(Cylindrocladiumblack rot of beggarweed)[28]；侵染決明(SennaobtusifoliaL.)和鷓鴣豆(CassiafasciculateMichx)引起黑腐病(Cylindrocladiumblack rot)[29]；侵染西印度櫻桃(MalpighiaglabraL.)引起葉斑病(leaf spots of antilles cherry)[30]；侵染富貴椰子(HoweabelmoreanaBecc.)引起褐斑病(Cylindrocladiumbrown leaf spot)[31]；侵染紅掌(AnthuriumandraeanumLinden)引起紅掌根冠腐病(root and crown rot of anthurium)[32]；侵染月桂樹(shù)(LaurusnobilisLinn.)引起苗期的根冠腐病(crown and root rot)[33]；侵染鱷梨(PerseaamericanaMill.)引起黑根腐病(black root rot)[34]。

5 病害循環(huán)

大豆紅冠腐病菌的傳播擴(kuò)散靠子囊孢子、分生孢子和微菌核[35]。田間排水和雨水與病原菌傳播有密切關(guān)系，雨水沖刷和田間排水可以把子囊孢子、分生孢子和微菌核沖到以前未被侵染的土壤[1]。子囊殼產(chǎn)生與否和環(huán)境濕度有很大關(guān)系，高濕度有利于子囊殼的形成。子囊孢子和分生孢子對(duì)干燥的環(huán)境較敏感，子囊孢子暴露在33 ℃空氣中30 min后，存活率小于0.1%，田間也很少觀察到分生孢子[35]。當(dāng)攜帶有子囊孢子、分生孢子和微菌核的植物組織在土壤中存留8個(gè)月后，只有微菌核仍然存活，因此，子囊孢子和分生孢子在病害循環(huán)中只能短距離傳播[36]。病原菌主要以微菌核在土壤和植物病殘?bào)w上越冬和長(zhǎng)距離傳播。溫度20～30 ℃最適合病害發(fā)生。溫度在20 ℃以下或30 ℃以上時(shí)，病害發(fā)病程度會(huì)下降，當(dāng)溫度達(dá)到40 ℃時(shí)，基本上不發(fā)病[37]。因?yàn)楫?dāng)土壤溫度高于35 ℃時(shí)，病原菌所有營(yíng)養(yǎng)體和各種孢子都無(wú)法存活。適當(dāng)推遲大豆種植的時(shí)間，大豆紅冠腐病的發(fā)病率可顯著下降[38]。當(dāng)土壤溫度持續(xù)4～5周低于5 ℃或土壤水分結(jié)冰時(shí)，微菌核數(shù)量會(huì)減小[39]。大豆和花生輪作會(huì)增加田間微菌核的數(shù)量，加重病害的流行和發(fā)生[40]。

6 病原菌鑒定和分離方法

大豆紅冠腐病菌的形態(tài)鑒定主要采用康乃馨葉片瓊脂(carnation leaf agar, CLA)。25 ℃光暗交替培養(yǎng)7 d后觀察菌落培養(yǎng)形態(tài)和顯微特征，包括分生孢子形狀和隔膜數(shù)、子囊殼形態(tài)、子囊孢子和泡囊大小。病原菌泡囊的形狀和大小及分子孢子形態(tài)是病原菌鑒定最主要的依據(jù)[5]。分子生物學(xué)鑒定主要針對(duì)β-tubulin基因、His3基因、EF-1α基因、Cal基因進(jìn)行鑒定[41]。

使用選擇性培養(yǎng)基可以從土壤中分離大豆紅冠腐病菌，例如在酵母粉葡萄糖瓊脂培養(yǎng)基中加入抗生素如噻菌靈、氯霉素和金霉素等[42]；在蔗糖蛋白胨瓊脂培養(yǎng)基中加入鏈霉素、氯霉素、牛膽汁、五氯硝基苯和噻苯咪唑等，可以從土壤中定量分離病原菌[43]。在普通的葡萄糖蛋白胨培養(yǎng)基中添加乳酸，將pH調(diào)至4，制成酸性培養(yǎng)基，也能從土壤中分離病原菌[44]。Ochi等將山梨糖、酵母膏、氟酰胺、噻菌靈、金霉素鹽酸鹽、氯霉素、表面活性劑、瓊脂和蒸餾水混合滅菌制成選擇性培養(yǎng)基，將土壤樣品過(guò)篩，用次氯酸鈉溶液表面消毒后接種到選擇性培養(yǎng)基上，該方法成功地分離到該病原菌。與其他方法相比，這種方法能提高病原菌的分離率，降低污染[45]。

7 防治措施

大豆紅冠腐病的防治措施主要包括農(nóng)業(yè)防治和化學(xué)防治。大豆不能與花生輪作，否則會(huì)增加田間微菌核數(shù)量和加重病害的發(fā)生[40]。在大豆紅冠腐病發(fā)生的地區(qū)，大豆種植時(shí)間應(yīng)參考當(dāng)?shù)氐耐寥罍囟萚8]。如果適當(dāng)推遲播種時(shí)間，土壤溫度將隨之升高，這樣可以減輕大豆紅冠腐病的發(fā)生[38]。在大豆播種前，使用草甘膦除草，可以減輕和控制大豆紅冠腐病[46]。在大豆播種前兩個(gè)星期，使用威百畝、異硫氰酸甲酯、疊氮化鈉和氯化苦等殺菌兼殺線蟲(chóng)劑進(jìn)行土壤熏蒸，可以減少土壤微菌核數(shù)量，降低病害的發(fā)生[47]。

我國(guó)是世界大豆的主產(chǎn)區(qū)，氣候條件適合大豆紅冠腐病的發(fā)生，對(duì)該病原菌的風(fēng)險(xiǎn)性評(píng)估分析結(jié)果表明：該菌是高度危險(xiǎn)的入侵性植物病原菌，山東、河南和河北等我國(guó)廣大的大豆和花生種植區(qū)都是該菌的適生分布區(qū)，該菌的入侵將對(duì)我國(guó)大豆和花生產(chǎn)業(yè)的可持續(xù)發(fā)展和生態(tài)安全構(gòu)成嚴(yán)重威脅[48]。為此，廣東省農(nóng)業(yè)廳已將該病原菌增列為《廣東省農(nóng)業(yè)植物檢疫性有害生物補(bǔ)充名單》。

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Anotablesoybeandisease,redcrownrotofsoybean

Gai Yunpeng1，2, Pan Ruqian2, Guan Mingfang2, Xu Dagao2

(1.HoumaEntry-ExitInspectionandQuarantineBureau,Shanxi043000,China; 2.DepartmentofPlantPathology，CollegeofNaturalResourcesandEnvironment,SouthChinaAgriculturalUniversity,Guangzhou510642,China)

Red crown rot (RCR) is an important disease in soybean. Typical symptoms of RCR are as follows: chlorotic, yellowish and blighted leaves, and wilting of the plants. Taproots were blackened and rotted. Numerous of reddish-orange spherical perithecia appeared on basal stems and roots of soybean. The causal pathogen of RCR isCylindrocladiumparasiticum(anamorph) andCalonectriailicicola(teleomorph). The research progress of RCR was also summarized.

red crown rot of soybean;Cylindrocladiumparasiticum; research progress

2013-08-27

：2013-09-18

廣東省科技計(jì)劃項(xiàng)目(2009B020310014)

S 435.65

：BDOI：10.3969/j.issn.0529-1542.2014.04.024

* 通信作者 E-mail: gaiyunpeng@gmail.com