花生葉斑病研究進(jìn)展

韓鎖義,　張新友*,　朱　軍,　許欣然,湯豐收,　董文召,　王忠于

(1. 河南省農(nóng)業(yè)科學(xué)院經(jīng)濟(jì)作物研究所,農(nóng)業(yè)部黃淮海油料作物重點(diǎn)實(shí)驗(yàn)室,國(guó)家油料作物改良中心河南花生分中心,河南省油料作物遺傳改良重點(diǎn)實(shí)驗(yàn)室,花生遺傳改良國(guó)家地方聯(lián)合工程實(shí)驗(yàn)室,鄭州　450002; 2. 海南大學(xué)農(nóng)學(xué)院,?？凇?70228)

?

花生葉斑病研究進(jìn)展

韓鎖義1,張新友1*,朱軍2,許欣然1,湯豐收1,董文召1,王忠于1

(1. 河南省農(nóng)業(yè)科學(xué)院經(jīng)濟(jì)作物研究所,農(nóng)業(yè)部黃淮海油料作物重點(diǎn)實(shí)驗(yàn)室,國(guó)家油料作物改良中心河南花生分中心,河南省油料作物遺傳改良重點(diǎn)實(shí)驗(yàn)室,花生遺傳改良國(guó)家地方聯(lián)合工程實(shí)驗(yàn)室,鄭州450002; 2. 海南大學(xué)農(nóng)學(xué)院,?？?70228)

花生葉斑病是一種世界性病害，其流行可以導(dǎo)致花生產(chǎn)量減少和品質(zhì)降低。本文對(duì)花生葉斑病的發(fā)生與危害、病原菌、防治措施、遺傳研究、分子標(biāo)記、轉(zhuǎn)基因及花生抗葉斑病種質(zhì)創(chuàng)制等多個(gè)方面的最新研究進(jìn)展進(jìn)行了綜述。為花生抗葉斑病研究提供參考。

花生;葉斑病;抗性

1　花生葉斑病的發(fā)生與危害

花生是世界上重要的經(jīng)濟(jì)作物和油料作物,在食用植物油消費(fèi)及休閑食品中占有舉足輕重的地位。世界范圍內(nèi),病害是花生減產(chǎn)的重要因素之一,其中,花生葉斑病在所有花生產(chǎn)區(qū)都發(fā)生或流行,可分為早斑病和晚斑病兩種,但發(fā)病程度因地區(qū)和季節(jié)不同而有差異[1]?；ㄉ~斑病主要危害葉片和莖稈,破壞葉綠素,造成光合作用效能下降,大量病斑會(huì)引起落葉,嚴(yán)重影響干物質(zhì)積累和莢果成熟。2種葉斑病發(fā)生初期在葉片上均產(chǎn)生黃褐色小斑點(diǎn),不易區(qū)分。癥狀因寄主基因型和環(huán)境因素的不同而有差異。早斑病在花期開始發(fā)病,隨著病情發(fā)展,產(chǎn)生圓形或不規(guī)則形病斑,葉正面病斑呈暗褐色,背面顏色較淺,呈淡褐色或褐色;病斑周圍有黃色暈圈。晚斑病病斑一般較小,圓形或近圓形。病斑呈黑褐色,在葉片正反兩面顏色相近。病斑邊緣通常沒(méi)有黃色暈圈,或有不明顯的淡黃色暈圈。在病斑背面,通常產(chǎn)生許多黑色小點(diǎn),即病菌分生孢子梗和分生孢子。病情嚴(yán)重時(shí),產(chǎn)生大量病斑,引起葉片干枯脫落[2-3]。這2種葉斑病可以在同一地區(qū)混合交叉發(fā)生,但通常只有1種葉斑病嚴(yán)重流行。

中國(guó)、美國(guó)和印度是世界上重要的花生生產(chǎn)國(guó)。花生主要產(chǎn)區(qū)受葉斑病的危害,每年均造成不同程度的損失。在美國(guó),早斑病流行較重, 僅在美國(guó)東南部每年就會(huì)造成數(shù)百萬(wàn)美元的損失[4]。在印度,晚斑病流行較重,早斑病則輕[2]。在我國(guó),葉斑病在花生產(chǎn)區(qū)普遍發(fā)生,危害重[3,5]。北方花生產(chǎn)區(qū)以晚斑病流行為主,在南方產(chǎn)區(qū)兩種葉斑病常交替發(fā)生,多數(shù)年份以晚斑病危害為主。晚斑病對(duì)全球花生生產(chǎn)造成的損失每年達(dá)6億美元[1,6-7],若與銹病同時(shí)發(fā)生,減產(chǎn)可超過(guò)70%[2,8]。在花生生育前期雨水多、光照嚴(yán)重不足的年份,早斑病可嚴(yán)重流行[3]。感染葉斑病的花生可以導(dǎo)致產(chǎn)量減少和品質(zhì)降低[9]。對(duì)于葉斑病的危害,如果不進(jìn)行有效的控制,可造成花生減產(chǎn)10%～20%,嚴(yán)重時(shí)達(dá)80%以上[1]。

2　花生葉斑病的病原菌

有2種真菌可引起花生葉斑病,其中落花生尾孢(CercosporaarachidicolaHori)引起早斑病,又稱褐斑病;落花生短胖孢[Cercosporidiumpersonatum(Berk. & Curt.) Deighton]引起晚斑病,又稱黑斑病。早斑病與晚斑病統(tǒng)稱花生葉斑病[1]。早斑病病菌無(wú)性階段為半知菌亞門尾孢屬落花生尾孢,有性階段為子囊菌綱球腔菌屬落花生球腔菌(Mycosphaerellaarachidicola)。晚斑病菌無(wú)性階段為半知菌類短胖孢屬落花生短胖孢,有性階段為子囊菌綱球腔菌屬伯克利球腔菌(Mycosphaerellaberkeleyi)。

落花生尾孢菌產(chǎn)生子座,深褐色,直徑為25～100 μm,多在葉片正面呈不規(guī)則分散排列。分生孢子梗叢生或散生,多數(shù)1個(gè),無(wú)分枝,黃褐色,基部顏色較暗,有1～2個(gè)隔膜或無(wú)隔膜。分生孢子頂生,淡橄欖色或無(wú)色,細(xì)長(zhǎng),一般5～7個(gè)隔膜[10]。病菌侵染寄主后,菌絲分布于寄主細(xì)胞間和細(xì)胞內(nèi),不產(chǎn)生吸器。病菌生長(zhǎng)的溫度范圍較廣,在5～35 ℃間均能生長(zhǎng),適宜溫度為15～30 ℃,最適為25 ℃;產(chǎn)孢適宜溫度為15～30 ℃,最適為25 ℃。低于5 ℃或者高于40 ℃,病菌則停止生長(zhǎng)和產(chǎn)孢。光照條件對(duì)病菌的生長(zhǎng)影響較大,短波長(zhǎng)的可見光能促進(jìn)孢子萌發(fā),近紫外照射對(duì)病菌的孢子萌發(fā)有明顯的促進(jìn)作用[10-11]。

落花生短胖孢子座多產(chǎn)生在葉片背面,半球形,包埋在寄主表皮下,直徑60～100 μm,在子座上叢生大量分生孢子梗。分生孢子梗呈褐色或暗褐色,有1～3個(gè)膝彎,多數(shù)無(wú)隔膜。分生孢子頂生,短而粗,圓筒形,多數(shù)3～5個(gè)隔膜[12]。病菌侵染寄主后,菌絲完全分布于細(xì)胞間隙,在海綿狀葉肉細(xì)胞內(nèi)產(chǎn)生分枝形吸器。病菌在10～35 ℃均可生長(zhǎng),20～30 ℃為適宜溫度,最適生長(zhǎng)溫度為28 ℃,最適pH為6,致死溫度是60 ℃,光暗交替有利于菌絲生長(zhǎng),病菌分生孢子萌發(fā)最適溫度為25 ℃,最適pH為6,在水滴中孢子有較高的萌發(fā)率,光/暗交替培養(yǎng)有利于孢子萌發(fā),分生孢子致死溫度為52 ℃[13]。該病菌喜高溫的習(xí)性導(dǎo)致田間發(fā)病高峰主要在花生生長(zhǎng)中后期,此時(shí)的溫度條件正適合病菌的生長(zhǎng)擴(kuò)展。

2種葉斑病病菌主要以分生孢子座和菌絲體在病株殘?bào)w上越冬?；蛞苑稚咦羽じ皆谇v果、莖稈表面越冬,翌年借風(fēng)雨傳播進(jìn)行初侵染和再侵染。分生孢子萌生芽管直接從花生葉片表皮或氣孔侵入。22～23 ℃時(shí),3～4 d顯病,再經(jīng)1周開始產(chǎn)孢。在有露水或水膜的情況下產(chǎn)孢量最大。清晨葉片上露水剛消失和下雨之前是分生孢子擴(kuò)散的高峰期,再侵染頻繁[13]。

3　花生葉斑病的防治途徑

3.1抗病品種的培育

花生不同種質(zhì)間抗、感病程度存在一定差異,尤其是野生花生中存在著優(yōu)異的抗病基因,因而培育抗病品種是減輕病害的有效途徑[14]。在同樣病原菌壓力的情況下,一般抗病品種比非抗病品種發(fā)病慢,感病輕,并能獲得較高的產(chǎn)量。此外,抗病品種的應(yīng)用可以減少殺菌劑的使用量、降低生產(chǎn)成本,減少風(fēng)險(xiǎn)。缺點(diǎn)是抗病新品種選育周期較長(zhǎng)。

3.2殺菌劑的推廣應(yīng)用

應(yīng)用殺菌劑是當(dāng)前田間防治葉斑病的重要措施。大規(guī)模推廣應(yīng)用殺菌劑對(duì)防治葉斑病起到了重要作用,化學(xué)藥劑對(duì)于花生葉部病害的防治效果一般在60%左右[15]。目前,對(duì)花生葉斑病防效較好的藥劑是多菌靈、代森錳鋅、百菌清、32.5%苯甲·嘧菌酯、60%唑醚·代森聯(lián)水分散粒劑和12.5%氟環(huán)唑懸浮劑[16-18]。但化學(xué)農(nóng)藥的不合理及過(guò)量使用會(huì)引起人畜中毒、生態(tài)環(huán)境破壞和植物藥害等一系列環(huán)境、社會(huì)問(wèn)題。

3.3生物防治措施

在探索生物防治在晚斑病上的應(yīng)用時(shí),Kishore等[19]采用甲殼素輔以葉面噴施黏質(zhì)沙雷氏菌(Serratiamarcescens) GPSS5 防治晚斑病,可減少該病的癥狀達(dá)64%。此外,鄢洪海等[20]研究了叢枝菌根真菌對(duì)黑斑病的生物防治作用,接種后可使花生株高、分枝、莢果數(shù)、莢果干重、地上部干重等生長(zhǎng)指標(biāo)顯著地增加,對(duì)花生黑斑病防治效果為5.4%。

3.4綜合管理措施

特殊的溫度與氣候條件對(duì)于葉斑病的發(fā)生有重要影響。因而,加強(qiáng)栽培管理,創(chuàng)造不利于發(fā)病的微生態(tài)環(huán)境,可減少初次侵染源及田間菌源。適期播種,合理密植,施足底肥,可以促進(jìn)花生健壯生長(zhǎng),來(lái)抵御病害的侵染。在發(fā)病前期盡早預(yù)報(bào)是一項(xiàng)重要預(yù)防措施,Olatinwo 等[4]建立的高分辨率天氣研究和預(yù)報(bào)(WRF)模型在花生早斑病的管理應(yīng)用中能發(fā)揮較大作用。該系統(tǒng)建立了精準(zhǔn)的空間-溫度-濕度模型,檢測(cè)病害發(fā)生的潛在威脅。在俄克拉荷馬州和佐治亞州的應(yīng)用結(jié)果表明,可以幫助種植者作出準(zhǔn)確的管理決策,降低病害的影響。

4　花生葉斑病抗性遺傳及抗病基因分子標(biāo)記

花生葉斑病抗性的遺傳較復(fù)雜[21],至今還沒(méi)有完全闡明。Sharief等[22-23]研究認(rèn)為早斑病和晚斑病這兩種葉斑病抗性是獨(dú)立遺傳的,受2對(duì)以上的核基因控制,且抗性均是由隱性基因控制。Nevill[24]研究認(rèn)為晚斑病的抗性受5個(gè)隱性基因位點(diǎn)控制。Kornegay等[25]推測(cè)花生葉斑病抗性為數(shù)量性狀,且抗性以加性基因作用為主。Miller等[1]研究認(rèn)為葉斑病抗性通常與晚熟性相關(guān)。Soriano等[26]對(duì)花生突變體的研究結(jié)果表明,2個(gè)突變基因IS-1和IS-2控制對(duì)葉斑病的抗性。當(dāng)病原菌感染葉片后,表皮細(xì)胞質(zhì)膜中的突變體蛋白抑制病原菌菌絲生長(zhǎng)。突變蛋白能夠去除纖維素酶的多肽鐵輔因子。在雜合狀態(tài)中,基因IS-2控制著抑制菌絲分泌的多肽的合成。

目前,與葉斑病抗性相關(guān)的分子標(biāo)記有RAPD、AFLP、SSR。Stalker等[27]在對(duì)含有Arachiscardenasii親緣的遠(yuǎn)緣雜交種中發(fā)掘出與早斑病抗性連鎖的RAPD標(biāo)記。夏友霖[12]以抗/感晚斑病組合‘中花5號(hào)’בICGV 86699’的F2分離群體為材料,以AFLP分析結(jié)合BSA法篩選到與晚斑病抗性連鎖較緊密的AFLP標(biāo)記3個(gè),分別為E35/M51、E37/M48和E41/M47,其與抗性間的圖距分別為7.40、7.40和8.67 cM;所獲得的3個(gè)標(biāo)記間連鎖緊密,位于同一連鎖群上,其中與抗性連鎖最緊密的標(biāo)記E35/M551和E37/M48更具實(shí)用價(jià)值。Leal-Bertioli等[28]運(yùn)用含93個(gè)家系的F2群體發(fā)掘出花生抗病基因核苷酸結(jié)合位點(diǎn)(NBS)編碼區(qū)11個(gè),晚斑病相關(guān)的QTLs位點(diǎn)5個(gè),貢獻(xiàn)率介于4.2%～43.8%。Khedikar等[29]運(yùn)用含268個(gè)家系的重組自交系分離群體(TAG24×GPBD4)定位出花生抗晚斑病相關(guān)的QTLs位點(diǎn)11個(gè),貢獻(xiàn)率介于1.70%～6.50%。有2對(duì)SSR引物IPAHM103和PM384 可以用于標(biāo)記輔助選擇育種中[29-30]。

5　基因轉(zhuǎn)化研究

一些研究表明，轉(zhuǎn)基因是提高花生抗葉斑病的重要途徑之一。在花生中過(guò)表達(dá)其他物種抗病基因?qū)ㄉ戆卟】剐杂幸欢ㄐЧ??？寺～@得全長(zhǎng)的芥菜防御素基因(BjD),通過(guò)根癌農(nóng)桿菌介導(dǎo)轉(zhuǎn)化花生,無(wú)論離體葉片接菌還是田間植株接菌均表明,轉(zhuǎn)基因花生對(duì)晚斑病的抗性比對(duì)照明顯增加[31]?；ㄉ羞^(guò)表達(dá)細(xì)菌、水稻幾丁質(zhì)酶基因?qū)ν戆卟】剐运接幸欢ǖ奶岣遊32-36]。Vasavirama等[37]用龍葵中的病程相關(guān)蛋白編碼基因SniOLP和蘿卜中的病程相關(guān)蛋白編碼基因Rs-AFP2與35S啟動(dòng)子構(gòu)建載體進(jìn)行花生轉(zhuǎn)基因研究的結(jié)果表明,過(guò)表達(dá)SniOLP和Rs-AFP2 基因可以有效提高花生對(duì)晚斑病的抗性,病斑減少并延遲發(fā)病,轉(zhuǎn)基因植株病癥與對(duì)照相比減輕97%以上。劉風(fēng)珍等[38]利用花粉管通道技術(shù)將國(guó)槐DNA導(dǎo)入花生栽培品種可以提供植株對(duì)花生早斑病的抗性。Kumar等[39]從花生抗晚斑病的二倍體野生種Arachisdiogoi中克隆的花生親環(huán)素蛋白AdCyp基因轉(zhuǎn)化煙草也表明對(duì)黑斑病抗性增加。

6　花生葉斑病抗性種質(zhì)創(chuàng)制

長(zhǎng)久以來(lái),花生研究者致力于抗病種質(zhì)的創(chuàng)制。一般情況下,抗性品種比非抗病品種產(chǎn)量較低且晚熟,抗病性強(qiáng)的早熟花生品種還很欠缺[1]。美國(guó)20世紀(jì)90年代的品種‘Southern Runner’對(duì)晚斑病中等抗性,莢果產(chǎn)量與‘Florunner’ 相當(dāng),但比‘Florunner’ 晚熟10～14 d。后來(lái)還選育有‘Florida MDR98’、‘C-99R’和 ‘York’等抗葉斑病品種[40-41]。玻利維亞地方品種‘Bayo Grande’,以及其他一系列的育種系也有較好的葉斑病抗性[42]。澳大利亞新品種‘Sutherland’對(duì)晚斑病有明顯的高抗水平[43]。Dwivedi等[9]從 15份種間雜交材料中選出了ICGV 99006、99013、 99004、 9903、 99001等5個(gè)好的晚斑病抗性育種親本材料。董煒博等[44]引進(jìn)品種‘UF91108’對(duì)晚斑病有高抗水平,可以作為花生育種的抗源材料。袁虹霞等[45]篩選出‘開農(nóng)31’、‘濮花8030’、‘豫花15號(hào)’等對(duì)花生早斑病和晚斑病抗性較好的系列品種。路興濤等[46]鑒定出‘花育23’、‘日花1號(hào)’和‘四粒紅’為高抗早斑病品種。劉風(fēng)珍等[38]通過(guò)將國(guó)槐DNA導(dǎo)入花生栽培種‘79266’,篩選出抗花生早斑病的新材料‘05D1148’。

花生野生種質(zhì)中具有對(duì)葉斑病有高抗水平的材料。研究者正在努力將這些抗性基因轉(zhuǎn)入栽培種中,ICRISAT、中國(guó)農(nóng)業(yè)科學(xué)院油料研究所、河南省農(nóng)業(yè)科學(xué)院等單位也利用野生花生創(chuàng)制了大批的抗病中間材料[12,46-51]。目前,雖然抗性育種取得了一定的進(jìn)展,但要將高水平的抗性匯聚到綜合性狀優(yōu)良的高產(chǎn)栽培種中仍然有一定困難。

7　未來(lái)研究方向與展望

隨著花生葉斑病相關(guān)研究的不斷深入,病害田間防控將是以高效、低毒新型藥劑為主,結(jié)合多種農(nóng)藝措施進(jìn)行綜合防治。葉斑病的生物防治措施雖然已經(jīng)取得了一些可喜成績(jī),但從小面積試驗(yàn)階段到大面積生產(chǎn)實(shí)踐還有一段距離。如何全面闡明花生葉斑病抗性遺傳機(jī)制并根據(jù)生產(chǎn)需要定向培育綜合性狀優(yōu)良、高抗花生葉斑病的新品種是花生科研工作者面臨的重要課題之一。

[1]Miller I L, Norden A J, Knauft D A, et al. Influence of maturity and fruit yield on susceptibility of peanut toCercosporidiumpersonatum(late leafspot pathogen)[J]. Peanut Science, 1990, 17(2): 52-58.

[2]McDonald D, Subrahmanyam P, Gibbons R W, et al. Early and late leaf spots of groundnut[M]. India: International Crops Research Institute for the Semi-Arid Tropics,1985.

[3]廖伯壽,肖達(dá)人.花生的抗病育種[M]∥孫大容.花生育種學(xué).北京:中國(guó)農(nóng)業(yè)出版社,1998: 222-255.

[4]Olatinwo R O,Prabha T V,Paz J O, et al. Predicting favorable conditions for early leaf spot of peanut using output from the weather research and forecasting (WRF) model [J]. International Journal of Biometeorology, 2012, 56(2): 259-268.

[5]湯豐收, 盧為國(guó), 張玉聚, 等, 中國(guó)植保技術(shù)原色圖解 4. 花生大豆病蟲草害原色圖解[M].北京: 中國(guó)農(nóng)業(yè)科學(xué)技術(shù)出版社, 2010.

[6]Dwivedi S L, Crouch J H, Nigam S N, et al. Molecular breeding of groundnut for enhanced productivity and food security in the Semi-Arid Tropics: opportunities and challenges [J]. Advances in Agronomy, 2003, 80: 153-221.

[7]Ogwulumba S I, Ugwuoke K I, Iloba C. Prophylactic effect of paw-paw leaf and bitter leaf extracts on the incidence of foliar myco-pathogens of groundnut (ArachishypogaeaL.) in Ishiagu, Nigeria [J]. African Journal of Biotechnology, 2008, 7(16): 2878-2880.

[8]Shokes F M, Culbreath A K. Early and late leaf spots [M]∥Kokalis-Burelle N,Porter D M,Rodriguez-kbana R, et al.Compendium of peanut disease, 2nd ed. APS Press, 1997.

[9]Dwivedi S L, Pande S, Rao J N, et al. Components of resistance to late leaf spot and rust among interspecific derivatives and their significance in a foliar disease resistance breeding in groundnut (ArachishypogaeaL.) [J]. Euphytica, 2002, 125(1): 81-88.

[10]薛其勤, 萬(wàn)勇善, 劉風(fēng)珍. 花生褐斑病病菌的分離培養(yǎng)及致病性研究[J]. 中國(guó)農(nóng)學(xué)通報(bào), 2007, 23(3): 343-346.

[11]黎穗臨, 葉維霖．花生褐斑病菌培養(yǎng)基篩選及生理性狀和致病性研究[J]．植物病理學(xué)報(bào), 1991, 21(1): 31-35．

[12]夏友霖. 花生晚斑病抗性AFLP 標(biāo)記篩選和川渝地區(qū)花生遺傳多樣性分析[D].重慶: 西南大學(xué)， 2003.

[13]鄢洪海, 張茹琴. 環(huán)境因子對(duì)花生黑斑病菌生長(zhǎng)發(fā)育的影響[J]. 湖北農(nóng)業(yè)科學(xué), 2009, 48(9): 2146-2149.

[14]Mallikarjuna N, Jadhav D R, Reddy K, et al. Screening new arachis amphidiploids, and autotetraploids for resistance to late leaf spot by detached leaf technique [J]. European Journal of Plant Pathology, 2012, 132(1): 17-21.

[15]周蓉, 段乃雄, 談?dòng)羁? 花生葉部病害抗病性種質(zhì)鑒定[J]. 花生科技, 1990 (2): 33-34.

[16]潘一展, 王繼臣. 花生葉斑病的發(fā)生規(guī)律和防治技術(shù)[J]. 河南農(nóng)業(yè)科學(xué), 1999(6): 21-22.

[17]范君龍, 殷君華, 姚潛, 等. 五種殺菌劑對(duì)花生葉斑病防治效果的試驗(yàn)[J]. 農(nóng)業(yè)科技通訊, 2014 (5): 96-98.

[18]鄒宗峰, 田明英, 繆玉剛,等. 新型農(nóng)藥防治花生葉斑病試驗(yàn)研究[J]. 現(xiàn)代農(nóng)業(yè)科技, 2012(2):146-147.

[19]Kishore G K, Pande S, Podile A R. Chitin-supplemented foliar application of Serratia marcescens GPS5 improves control of late leaf spot disease of groundnut by activating defense-related enzymes [J].Journal of Phytopathology,2005,153:169-173.

[20]鄢洪海,張茹琴,安佰國(guó).AM真菌摩西球囊霉對(duì)2種花生葉斑病的生防及促生作用[J].中國(guó)農(nóng)學(xué)通報(bào)，2011,27(30):209-213.

[21]Holbrook C C, Stalker H T. Peanut breeding and genetic resources [J]. Plant Breeding Reviews, 2003, 22: 297-356.

[22]Sharief Y. The inheritance ofCercosporaleafspot resistance inArachisspecies [D].USA:North Carolina State University,1972.

[23]Sharief Y, Rawlings J O, Gregory W C. Estimates of leafspot resistance in three interspecific hybrids ofArachis[J]. Euphytica, 1978, 27(3): 741-751.

[24]Nevill D J. Inheritance of resistance to Cercosporidiumpersonatumin groundnuts: a genetic model and its implications for selection [J]. Oleagineux, 1982, 37(7): 355-366.

[25]Kornegay J L, Beute M K, Wynne J C. Inheritance of resistance toCercosporaarachidicolaandCercosporidiumpersonatumin six virginia-type peanut lines [J]. Peanut Science, 1980, 7(1): 4-9.

[26]Soriano J D, Villanueva A C, Calaguas A M, et al. Gene expression in peanut mutants resistant to leaf spot disease [J]. Cell Biology International Reports, 1990, 14: 181.

[27]Stalker H T, Mozingo L G. Molecular markers ofArachisand marker-assisted selection [J]. Peanut Science, 2001, 28(2): 117-123.

[28]Leal-Bertioli S C M, José V F A C, Alves-Freitas D M T, et al. Identification of candidate genome regions controlling disease resistance inArachis[J]. BMC Plant Biology, 2009, 9: 112-123.

[29]Khedikar Y P, Gowda M V C, Sarvamangala C, et al. A QTL study on late leaf spot and rust revealed one major QTL for molecular breeding for rust resistance in groundnut (ArachishypogaeaL.)[J]. Theoretical and Applied Genetics, 2010, 121(5): 971-984.

[30]Shoba D, Manivannan N, Vindhiyavarman P, et al. SSR markers associated for late leaf spot disease resistance by bulked segregant analysis in groundnut (ArachishypogaeaL.)[J]. Euphytica, 2012, 188(2): 265-272.

[31]Anuradha T S, Divya K, Jami S K, et al. Transgenic tobacco and peanut plants expressing a mustard defensin show resistance to fungal pathogens[J]. Plant Cell Reports, 2008, 27(11): 1777-1786.

[32]Rohini V K, Rao K S. Transformation of peanut (ArachishypogaeaL.) with tobacco chitinase gene: variable response of transformants to leaf spot disease[J]. Plant Science, 2001, 160 (5): 889-898.

[33]許澤永, 廖伯壽, 晏立英, 等. 轉(zhuǎn)基因花生研究進(jìn)展[J]. 中國(guó)油料作物學(xué)報(bào), 2007, 29 (4): 489-496.

[34]Iqbal M M, Zafar Y, Nazir F, et al. Over expression of bacterial chitinase gene in Pakistani peanut (ArachishypogaeaL.) cultivar GOLDEN[J]. African Journal of Biotechnology, 2011, 10(31): 5838-5844.

[35]Iqbal M M, Nazir F, Ali S, et al. Over expression of rice chitinase gene in transgenic peanut (ArachishypogaeaL.) improves resistance against leaf spot[J]. Molecular Biotechnology, 2012, 50(2): 129-136.

[36]Prasad K, Bhatnagar M P, Waliyar F, et al. Overexpression of a chitinase gene in transgenic peanut confers enhanced resistance to major soil borne and foliar fungal pathogens [J]. Journal of Plant Biochemistry and Biotechnology,2013, 22(2): 222-233.

[37]Vasavirama K, Kirti P B. Increased resistance to late leaf spot disease in transgenic peanut using a combination of PR genes[J]. Functional & Integrative Genomics, 2012, 12(4): 625-634.

[38]劉風(fēng)珍, 萬(wàn)勇善, 薛其勤. 國(guó)槐DNA導(dǎo)入花生栽培品種選育抗葉斑病新種質(zhì)的鑒定[J]. 華北農(nóng)學(xué)報(bào), 2010, 25(6): 113-117.

[39]Kumar K R R, Kirti P B. Differential gene expression inArachisdiogoiupon interaction with peanut late leaf spot pathogen,Phaeoisariopsispersonataand characterization of a pathogen induced cyclophilin [J]. Plant Molecular Biology, 2011, 75(4/5): 497-513.

[40]Gorbet D W, Shokes F M, Culbreath A K, et al. C-99R——A new multiple disease resistant peanut cultivar[R]. Marianna NFREC Resarchi Report 99-2,1999.

[41]Singh M P, Erichson J E, Boote K J, et al. Late leaf spot effects on growth, photosynthesis, and yield in peanut cultivars of differing resistance [J]. Agronomy Journal, 2011, 103(1): 85-91.

[42]Gremillion S, Culbreath A, Gorbet D, et al. Response of progeny bred from Bolivian and North American cultivars in integrated management systems for leaf spot of peanut (Arachishypogaea)[J]. Crop Protection, 2011, 30(6): 698-704.

[43]Kelly L A, Ryley M J, Trevorrow P R, et al. Reduced fungicide use on a new Australian peanut cultivar, highly resistant to the late leaf spot and rust pathogens [J]. Australasian Plant Pathology, 2012, 41(4): 359-373.

[44]董煒博, 石延茂, 趙志強(qiáng), 等. 花生品種(系)葉部病害綜合抗性鑒定[J]. 中國(guó)油料作物學(xué)報(bào), 2000, 22(3): 71-74.

[45]袁虹霞, 孫炳劍, 李洪連, 等. 花生品種(系) 對(duì)葉斑病的抗性鑒定[J]. 河南農(nóng)業(yè)科學(xué), 2004 (12): 35-38.

[46]路興濤, 張成玲,張?zhí)锾?等. 不同花生品種對(duì)花生褐斑病和網(wǎng)斑病抗病性鑒定[J].花生學(xué)報(bào), 2013, 42(3):52-55.

[47]Anderson W F, Holbrook C C, Brenneman T B. Resistance toCercosporidiumpersonatumwithin peanut germplasm[J]. Peanut Science, 1993, 20(1): 53-57.

[48]Waliyar F, McDonald D, Subba R P V, et al. Components of resistance to an Indian source ofCercosporaarachidicolain selected peanut lines [J]. Peanut Science,1993, 20(2): 93-96.

[49]Ouedraogo M, Smith O D, Simpson C E, et al. Early and late leaf spot resistance and agronomic performance of nineteen interspecific derived peanut lines [J]. Peanut Science, 1994, 21(2): 99-104.

[50]Mehan V K, Reddy P M, Subrahmanyam P, et al. Identification of new sources of resistance to rust and late leaf spot in peanut [J]. International Journal of Pest Management, 1996, 42(4): 267-271.

[51]Singh A K, Mehan V K, Nigam S N. Sources of resistance to groundnut fungal and bacterial disease: an update and appraisal [M]. India: International Crops Research Institute for the Semi-Arid Tropics, 1997.

(責(zé)任編輯：田喆)

Progresses in the research on peanut leaf spot disease

Han Suoyi1,Zhang Xinyou1,Zhu Jun2,Xu Xinran1,Tang Fengshou1,Dong Wenzhao1,Wang Zhongyu1

(1. Industrial Crops Research Institute, Henan Academy of Agricultural Sciences / Key Laboratory of Oil Crops in Huanghuaihai Plains, Ministry of Agriculture /National Center for Peanut Improvement,Division of Peanut in Henan / Henan Provincial Key Laboratory for Oil Crops Improvement/National and Local Joint Engineering Laboratory for Peanut Genetic Improvement, P R China;Zhengzhou450002, China; 2. College of Agriculture, Hainan University, Haikou570228, China)

Leaf spot disease is one of the most important leaf diseases of the peanutArachishypogaea. The disease could cause substantial yield losses and reduce the fodder and seed quality in peanut. In this paper, we review the research status of leaf spot disease in peanut, including the occurrence and damage, pathogens, control measures, resistance genetics, molecular markers, transgenic research and germplasm creation on leaf spot disease resistance. The article provides a reference for the research of leaf spot disease resistance in peanut.

peanut;leaf spot disease;resistance

2015-01-26

2015-03-23

河南省農(nóng)業(yè)科學(xué)院優(yōu)秀青年科技基金(2013YQ08); 國(guó)家花生產(chǎn)業(yè)技術(shù)體系(CARS-14)

E-mail:Haasz@126.com

S 435.652

A

10.3969/j.issn.0529-1542.2016.02.003