劉成，韓冉，汪曉璐，宮文萍，程敦公，曹新有，劉愛峰，李豪圣，劉建軍

小麥遠緣雜交現(xiàn)狀、抗病基因轉(zhuǎn)移及利用研究進展

劉成，韓冉，汪曉璐，宮文萍，程敦公，曹新有，劉愛峰，李豪圣，劉建軍

（山東省農(nóng)業(yè)科學院作物研究所/農(nóng)業(yè)部黃淮北部小麥生物學與遺傳育種重點實驗室/小麥玉米國家工程實驗室，濟南 250100）

小麥近緣植物中含有豐富的抗病、抗逆和抗蟲等基因，是小麥育種的優(yōu)異基因源。通過遠緣雜交可以將近緣植物優(yōu)異基因轉(zhuǎn)移給小麥，創(chuàng)制包括雙二倍體或部分雙二倍體、附加系、代換系和易位系等在內(nèi)的小麥-近緣植物異染色體系。這些含小麥近緣植物血緣的異染色體系是研究物種染色體行為與進化、基因定位與作圖的重要素材，也是拓寬小麥的遺傳基礎(chǔ)、抵御小麥重要病蟲害、增加小麥產(chǎn)量和提升小麥品質(zhì)的重要物質(zhì)基礎(chǔ)。為了更加清晰地了解小麥遠緣雜交概況及小麥近緣植物抗病基因向小麥的轉(zhuǎn)移，也為今后小麥遠緣雜交研究和種質(zhì)資源的開發(fā)利用提供參考，文中對小麥族物種分類、小麥遠緣雜交的定義與意義、小麥族山羊草屬、黑麥屬、偃麥草屬、簇毛麥屬、冰草屬、大麥屬、披堿草屬、賴草屬、新麥草屬以及旱麥草屬物種與小麥遠緣雜交現(xiàn)狀和異染色體系創(chuàng)制情況進行了概括，并對來源于小麥近緣植物被正式命名的17個抗條銹病基因、35個抗葉銹病基因、30個抗稈銹病基因、41個抗白粉病基因、3個抗赤霉病基因、1個抗麥瘟病基因、1個抗葉枯病基因、1個抗穎枯病基因、4個抗褐斑病基因、2個抗眼斑病基因、1個抗梭條花葉病基因、2個抗線條花葉病基因和2個抗禾谷類黃矮病基因向小麥的轉(zhuǎn)移情況及其所在染色體的位置信息進行了歸納。小麥-黑麥1RS·1BL易位系、1RS·1AL易位系和小麥-偏凸山羊草2NS/2AS易位系等抗病優(yōu)良種質(zhì)的育成與利用在世界小麥育種史上做出了突出貢獻，然而，這僅僅得益于對少數(shù)抗病基因的利用。與目前已經(jīng)被命名的基因數(shù)量相比，被利用到小麥育種中的抗病基因相對較少。文中分析了當前已命名抗病基因利用情況比例偏低的原因，并對今后如何利用這些抗病基因提出了建議。同時，還列舉了已克隆的源自小麥近緣植物的抗病基因，并對克隆這些基因的方法以及今后可能的研究熱點進行了分析，認為加強無遺傳累贅的小麥-近緣植物易位系的創(chuàng)制與應用仍可能是今后小麥育種材料創(chuàng)新與新品種培育的一個重要發(fā)力點。

小麥；遠緣雜交；異染色體系；抗病基因；衍生品種

1 小麥族分類

小麥族（）有300多個物種，包含小麥屬（）、山羊草屬（）、黑麥屬（）、偃麥草屬（）、簇毛麥屬（）、冰草屬（）、大麥屬（）、披堿草屬（）、賴草屬（）、新麥草屬（）、旱麥草屬（）、類大麥屬（）、無芒草屬（）、異形花屬（）、棱軸草屬（）、鵝觀草屬（）、擬鵝觀草屬（）和澳麥草屬（）等，基本染色體組包含A—W和2個尚未確定的染色體組X和Y，表現(xiàn)出遺傳變異的多樣性[1-2]。小麥的近緣植物具有抗病[3-5]、抗蟲[4-5]、抗旱[6]、抗寒[7-8]、耐鹽[6,9-10]等優(yōu)良性狀，是小麥遺傳改良的寶貴基因資源庫[1-3]。

2 小麥遠緣雜交現(xiàn)狀

遠緣雜交是親緣關(guān)系較遠的（包括生物學規(guī)定的不同“種”間、“屬”間）以及親緣關(guān)系更遠的物種間雜交的統(tǒng)稱[11-12]。小麥族中，小麥與黑麥、小麥與偃麥草、小麥與山羊草以及不同小麥種間的雜交均屬遠緣雜交，而生物學上規(guī)定的“種”以內(nèi)的不同變種或品種間的雜交則統(tǒng)稱為近緣雜交[11]。將近緣植物與小麥雜交，不僅可以將其優(yōu)異基因?qū)胄←溸M行遺傳改良[12-15]，還可以用于基因及染色體作圖[16-17]、染色體行為及進化[18-19]等研究。自18世紀，科學家們就零星開始了小麥遠緣研究[20]。19世紀以來，國內(nèi)外科學家們在小麥遠緣雜交方面做了大量工作，不同小麥種間[21-22]、小麥與山羊草屬[2,23-26]、黑麥屬[2,20,26-29]、偃麥草屬[2,11,26,30-32]、簇毛麥屬[2,26,33-36]、冰草屬[26,37-39]、大麥屬[40-42]、披堿草屬[43-45]、賴草屬[26,46-48]、新麥草屬[49-51]、旱麥草屬[52-53]等屬物種遠緣雜交成功的結(jié)果陸續(xù)被報道出來。目前，除類大麥屬、擬鵝觀草屬、無芒草屬、異形花屬、棱軸草屬、澳麥草屬物種外，其余小麥族各屬物種均已有與小麥雜交成功的報道。

3 小麥-近緣植物異染色體系創(chuàng)制情況

包括小麥-近緣植物雙二倍體或部分雙二倍體、附加系、代換系和易位系等在內(nèi)的異染色體系，是向小麥轉(zhuǎn)移近緣植物優(yōu)異基因的橋梁和物質(zhì)基礎(chǔ)[5,54-55]。目前，小麥種間[21-22]、小麥與山羊草屬[2,55-57]、黑麥屬[2,58-60]、偃麥草屬[2,26,31,61-63]、簇毛麥屬[2,33-35,64]、冰草屬[2,65-67]、大麥屬[68-70]、披堿草屬[71-73]、賴草屬[2,74-76]、新麥草屬[2,77-80]等屬物種異染色體系創(chuàng)制成功的結(jié)果如雨后春筍般被報道。目前，雖然已有大量的小麥-近緣植物異染色體系被創(chuàng)制出來，然而，被直接用于小麥育種的小麥-近緣植物染色體易位系的比例還比較低。

4 近緣植物抗病基因向栽培小麥轉(zhuǎn)移情況

迄今為止，小麥與近緣植物雜交成功的報道已有數(shù)百個[5,11,19,26,31,81]，其中大部分與小麥五大主要病害抗病基因轉(zhuǎn)移有關(guān)[5,82]。目前，被國際小麥新基因命名委員會正式命名的抗小麥條銹病、葉銹病、稈銹病、白粉病和赤霉病的基因個數(shù)分別為82、79、60、65和7個，其中，來源于小麥近緣植物的基因個數(shù)分別有17（表1）、35（表2）、30（表3）、41（表4）和3個（表5），分別占被正式命名基因的20.7%、44.3%、50.0%、63.1%和42.9%。

此外，被正式命名的抗麥瘟病、葉枯病、穎枯病、褐斑病、眼斑病、梭條花葉病、線條花葉病和禾谷類黃矮病基因分別為8、18、3、7、3、1、3和3個，其中，來源于小麥近緣植物的基因個數(shù)分別有1、1、1、4、2、1、2和2個（表6），分別占被正式命名基因的12.5%、5.5%、33.3%、57.1%、66.7%、100%、66.7%和66.7%。

4.1 抗條銹病基因向小麥轉(zhuǎn)移情況

來源于小麥近緣植物的抗條銹病基因有17個（表1），包括來自頂芒山羊草的、偏凸山羊草的、擬斯卑爾脫山羊草的、粗山羊草的、粘果山羊草的、沙融山羊草的、卵穗山羊草的、三芒山羊草的、小傘山羊草的、栽培黑麥的、中間偃麥草的、硬粒小麥的、和以及野生二粒小麥的、和。

表1 小麥近緣植物抗條銹病基因向小麥轉(zhuǎn)移情況

4.2 抗葉銹病基因向小麥轉(zhuǎn)移情況

來源于小麥近緣植物的抗葉銹病基因有35個（表2），包括來自小傘山羊草的和、粗山羊草的、、、、和、擬斯卑爾脫山羊草的、、、、和、偏凸山羊草的、粘果山羊草的、沙融山羊草的、卵穗山羊草的、鉤刺山羊草的、柱穗山羊草的、短穗山羊草的、栽培黑麥的、和、長穗偃麥草的和、彭提卡偃麥草的、中間偃麥草的、粗穗披堿草的、栽培二粒小麥的、野生二粒小麥的和、硬粒小麥的、一粒小麥的和提莫非維小麥的。

4.3 抗稈銹病基因向小麥轉(zhuǎn)移情況

來源于小麥近緣植物的抗稈銹病基因有30個（表3），包括來自頂芒山羊草的、偏凸山羊草的、擬斯卑爾脫山羊草的、和、希爾斯山羊草的、卵穗山羊草的、栽培黑麥的、、和、簇毛麥的、彭提卡偃麥草和、長穗偃麥草的和、中間偃麥草的、野生二粒小麥的、、、、和、一粒小麥的、、和、硬粒小麥的以及提莫非維小麥的和。

表2 小麥近緣植物抗葉銹病基因向小麥轉(zhuǎn)移情況

表3 小麥近緣植物抗桿銹病基因向小麥轉(zhuǎn)移情況

4.4 抗白粉病基因向小麥轉(zhuǎn)移情況

來源于小麥近緣植物的抗白粉病基因有41個（表4），包括來自擬斯卑爾脫山羊草的和、粗山羊草的、、和、高大山羊草的、希爾斯山羊草的卵穗山羊草的、栽培黑麥的、、、和、簇毛麥的、和、中間偃麥草的和、彭提卡偃麥草的、一粒小麥的和、野生一粒小麥的、波斯小麥的、栽培二粒小麥的、、和、野生二粒小麥的、、、、、和、硬粒小麥的、烏拉爾圖小麥的以及提莫非維小麥的、和。

表4 小麥近緣植物抗白粉病基因向小麥轉(zhuǎn)移情況

4.5 抗赤霉病基因向小麥轉(zhuǎn)移情況

來源于小麥近緣植物的抗赤霉病基因有3個（表5），包括來自大賴草的、柯孟披堿草（也有科學家稱其為鵝觀草）的和來自彭提卡偃麥草的。

4.6 其他抗病基因向小麥轉(zhuǎn)移情況

來源于小麥近緣植物的五大主要病害之外的抗病基因有14個（表6），包括來自栽培二粒小麥的抗麥瘟病基因、粗山羊草的抗葉枯病基因、抗穎枯病基因和抗褐斑病基因、野生二粒小麥的抗褐斑病基因和、圓錐小麥的抗褐斑病基因、偏凸山羊草的抗眼斑病基因、簇毛麥的抗眼斑病基因和抗梭條花葉病毒基因、中間偃麥草的抗線條花葉病基因、以及抗禾谷類黃矮病基因和。

表5 小麥近緣植物抗赤霉病基因向小麥轉(zhuǎn)移情況

表6 小麥近緣植物抗麥瘟病等基因向小麥轉(zhuǎn)移情況

—表示基因已命名但無文獻發(fā)表（McIntosh R A與Worland A K，私人通訊）

—indicates that the gene has been designated but no reference published (MCINTOSH R A and WORLAND A K, private communication)

5 小麥近緣植物抗病基因的利用

在小麥遠緣雜交種質(zhì)應用方面，對世界小麥育種做出突出貢獻的當屬小麥-黑麥1RS·1BL易位系。1RS染色體上由于含和等基因，受到了廣大育種工作者的普遍青睞[14,203-206]，國外育種家們利用該易位系及其衍生系作親本，育成了山前麥、高加索、無芒一號和洛夫林13等高產(chǎn)抗病小麥，被全世界幾十個國家作為骨干親本應用，育成了一大批優(yōu)異小麥新品種[207-210]，在推動小麥品種的更新?lián)Q代中發(fā)揮了重要作用[203,211-212]。除了1RS·1BL易位系，國外育種家們還培育出了含1RS·1AL易位系的Amigo等品種，并以此為骨干親本，培育出了含該易位系的Zhytnytsa、Nota和Duma[203]、Columbia、Etude和Rastavitsa[213]、TAM107、TAM303、TAM305、AG Robust、Fannin、N96L9970[214-216]和Helami-105等小麥新品種/系[217]，在美國、墨西哥和歐洲等國家推廣應用。近年來，國際玉米小麥改良中心（CYMMYT）以該易位系為親本育成了CM409和CM451等一批小麥新品種/系（劉彩云，私人通訊）。

據(jù)報道，19世紀后期中國約70%小麥品種含1RS·1BL易位系[204-205]，其中，為中國小麥育種做出突出貢獻的矮孟牛（Ⅱ型、Ⅳ—Ⅶ型）、周麥22、周8425B和石4185等骨干親本材料均含有1RS染色體。近年來，由于新的致病生理小種的產(chǎn)生與流行，使得和等基因的抗性迅速喪失[177,218-219]，加上育種家們在育種過程中注意雜交親本的遺傳多樣性，因此，該易位系在中國小麥中的比例明顯下降[220]。雖然等基因的抗性已經(jīng)喪失[177,218-219]，但近期的研究發(fā)現(xiàn)，不同黑麥來源的1RS·1BL易位系可能含有不同的抗病等位基因[14,221]，即表明不同黑麥來源的該易位系仍能在小麥育種中發(fā)揮重要作用。尤其是近年來，不含黑麥堿但仍具有良好抗病性的1RS·1BL易位系的創(chuàng)制[222-224]，為小麥育種提供了新的育種資源。

除含1RS染色質(zhì)的育種材料外，含、和的小麥-偏凸山羊草2NS/2AS易位系對世界小麥育種也做出了突出貢獻。以該易位系為抗源育成的Mace（還含和）[225-226]、Jagger、Madsen、Overley、SY Gold、Trident、EGA Eaglehawk和Espada等小麥品種在美國、澳大利亞和歐洲等國家推廣應用[225-229]。研究發(fā)現(xiàn)，源自中國10余個省份69個小麥品種中的49%含該易位系[230]。此外，據(jù)報道，川育18、川麥25和川麥39等[231]、新麥19、濟麥20、濟麥21和師欒02-1[232]、蘭考906、西農(nóng)739、陜872和小偃216等品種/系[233]含有該易位系。近期研究發(fā)現(xiàn)，濟麥20和濟麥21中不含但中麥175中含有等基因，然而已對中國當前葉銹生理小種表現(xiàn)為感病[234]。此外，在澳大利亞、歐洲和中國的條銹抗性已經(jīng)完全或部分喪失[228,235-236]。因此，今后在育種中應減少對該易位系的利用。

自19世紀以來，中國在小麥遠緣雜交領(lǐng)域研究一直處在世界前列。中國科學家先后將偃麥草[31,61-63,81,237-244]、黑麥[58-60,245-252]、簇毛麥[35,253]和冰草[254-255]等種質(zhì)轉(zhuǎn)移給了小麥，育成了一大批遠緣雜交新材料。在對這些小麥遠緣雜交種質(zhì)利用方面，取得了舉世矚目的研究成果，培育出的抗條銹病的小偃系列品種及其衍生品種[256-258]、普冰系列及其衍生品種（張錦鵬，私人通訊）和陜麥號及西農(nóng)號小麥[259-260]、抗黃矮病的張春號、臨抗號、晉麥號小麥[261-262]和黑小麥品種[263-264]、抗白粉病的南農(nóng)號小麥及其衍生品種[265-267]、抗條銹和白粉等病害的川農(nóng)系列小麥及其衍生品種[268-269]和遠豐號小麥[270]等在中國大面積推廣應用。上述品種抗病性來源主要為、、、、、和尚未被證明命名的少數(shù)幾個基因。

除此之外，值得一提的是，中國科學家分別將來自彭提卡偃麥草抗赤霉病基因[189,271]和長穗偃麥草的尚未被命名的赤霉病基因[272-273]分別轉(zhuǎn)移到小麥，創(chuàng)制了一批小麥-偃麥草染色體易位系，并將其導入中國主栽小麥，培育出了一批赤霉抗性達到中抗水平正在參加區(qū)域試驗的小麥新品系，有望對小麥抗赤霉病育種發(fā)揮重要作用。

6 結(jié)論與展望

目前，眾多個有明顯育種價值的小麥-近緣植物染色體易位系/漸滲系被成功創(chuàng)制出來[4-5]，并且有近140個抗病新基因被正式命名（表1—表6），但就基因利用狀況來看，被利用到小麥抗病育種上的基因的比例還比較低。其原因可能是：（1）部分基因的抗性已經(jīng)/正在喪失，例如、和等[274]，、和等[234]，、和等[275]，、和等[276]；（2）部分易位染色體具有遺傳累贅，例如[277]、[278]、和[179]等基因所在近緣植物染色體臂。因此，在今后的研究中，應該做到：（1）加強二倍體和四倍體小麥抗病基因向栽培小麥的導入與利用；（2）加強對有遺傳累贅效應易位系的染色體工程誘導。通過抗病基因轉(zhuǎn)育，創(chuàng)制出更多的抗病種質(zhì)資源并對其進行育種學評價。目前，已有多個研究團隊在開展這項工作[267,279]；（3）加強對無遺傳累贅且具優(yōu)異抗性易位系[267,279-280]的利用工作。

克隆抗病基因是研究其抗病機理的基礎(chǔ)。目前，從小麥近緣植物中克隆出的抗病基因主要包括[281]、[282]、[283]、[284]、[285]、[286]、[287]、[288]、[289-290]等。其中，除和[85,162]外，其他幾個基因均源自二倍體或四倍體小麥[96,98,100,141,143-145]。由于缺乏參考基因組信息，目前，從小麥-近緣植物（這里指非小麥屬物種）染色體易位系中克隆抗病基因還有一定困難。當前，克隆這些基因可以利用不同居群的抗病性不同的同一小麥近緣種進行雜交（或利用誘變技術(shù)創(chuàng)制突變體），配置抗感分離群體進行基因定位與克隆，例如[282]和[289]的克??；還可以創(chuàng)制更多的小麥-近緣植物染色體結(jié)構(gòu)變異體，將抗病基因定位到近緣植物某一染色體小片段上，進而利用已克隆的模式植物抗病基因所在染色體區(qū)間與上述小片段區(qū)間進行基因共線性分析與確證，同源克隆近緣植物的抗病基因。近期筆者及其合作者們利用該方法克隆了（待發(fā)表）。

從理論研究上講，隨著小麥族物種基因組測序工作的陸續(xù)開展與完成，能夠用于抗病基因克隆的參考基因組信息越來越多，今后克隆小麥近緣植物抗病基因?qū)兊迷絹碓饺菀祝虼?，這些基因的抗病調(diào)控機制以及不同物種共線性抗病基因的進化可能將成為新的研究熱點。從應用研究上講，小麥-黑麥1RS·1BL易位系、1RS·1AL易位系和小麥-偏凸山羊草2NS/2AS易位系等抗病優(yōu)良種質(zhì)的育成與利用在世界小麥育種史上做出了突出貢獻，然而，這僅僅得益于對少數(shù)抗病基因的利用。雖然目前被利用到小麥育種中的抗病基因相對較少，但加強無遺傳累贅的小麥-近緣植物易位系的創(chuàng)制與應用仍可能是今后小麥育種材料創(chuàng)新與新品種培育的一個重要發(fā)力點。

致謝：堪薩斯州立大學Friebe B教授、德克薩斯農(nóng)工生命研究和推廣中心Liu SY教授、北達科他州立大學Cai XW教授、悉尼大學Zhang P博士和李建波博士、阿德萊德大學Dundas I博士、John Innes Centre的Griffiths S研究員、CYMMYT劉彩云博士后、烏克蘭國家種子與品種調(diào)查中心Motsnyi I研究員、電子科技大學楊足君教授、中國農(nóng)業(yè)科學院張錦鵬研究員、西北農(nóng)林科技大學王長有教授、魯東大學崔法教授、山東農(nóng)業(yè)大學鮑印廣教授在不同國家小麥品種所含外源染色質(zhì)信息搜集中給予的大力幫助，在此表示感謝。

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Research Progress of Wheat Wild Hybridization, Disease Resistance Genes Transfer and Utilization

LIU Cheng, HAN Ran, WANG XiaoLu, GONG WenPing, CHENG DunGong, CAO XinYou, LIU AiFeng, LI HaoSheng, LIU JianJun

(Crop Research Institute, Shandong Academy of Agricultural Sciences/Key Laboratory of Wheat Biology and Genetic Improvement in the North Huang-Huai River Valley, Ministry of Agriculture/National Engineering Laboratory for Wheat and Maize, Jinan 250100)

Wheat alien species are vast reservoir of diversity for disease and pest resistance as well as stress tolerance, which are excellent gene sources for wheat breeding. Through wide hybridization, the genes of alien species could be transferred to wheat to create wheat-alien chromosome lines such as amphiploids or partial amphiploids, additions, substitutions and translocation lines. These genetic stocks could be utilized to study chromosome behavior and genome evolution, mapping genes, and diversifying the genetic basis of wheat for diseases and pest resistance, as well as yield and quality improvement. In order to understand the progress of wheat wide hybridization and useful gene transfer from alien species to wheat, in this paper, the classification of the tribe Triticeae, the definition and significance of wheat wide hybridization, alien transfers progress from species belonging to genera,,,,,,,,andto wheat have been summarized and discussed. To date, the official designated genes originated from wheat alien species include 17 stripe rust resistance genes, 35 leaf rust resistance gens, 30 stem rust resistance genes, 41 powdery mildew resistance genes, 3 Fusarium head blight-resistance genes, one wheat blast resistance gene, one Septoria tritici blotch resistance genes, one Septoria nodorum blotch resistance gene, 4 tan spot resistance genes, 2 eyespot resistance genes, one wheat spindle streak mosaic virus resistance gene, 2 wheat streak mosaic virus resistance genes and 2 cereal yellow dwarf resistance genes. Names and the chromosomal locations of these disease resistance genes were inducted. Moreover, the utilization of these genes in wheat breeding has also been reviewed and summarized. In the history of world wheat breeding, disease resistant germplasms such as wheat-rye 1RS·1BL translocation, 1RS·1AL translocation and wheat-2NS/2AS translocation have made outstanding contributions. However, this only benefited from the utilization of a few disease resistant genes. Compared to the number of the designated genes, relatively few disease-resistant genes have been used in wheat breeding. In this paper, the limiting factors for the underutilization are discussed. Suggestions on how to use these disease-resistant genes in the future are put forward. Meanwhile, the cloned disease-resistant genes from wheat alien species are listed. The methods of cloning these genes and the possible research hotspots in the future are also analyzed. It is believed that the development and application of wheat-wild species translocation lines without genetic drag may be an important driving force for material innovation and variety breeding in the future.

wheat; wild hybridization; chromosome line; disease resistance gene; derived varieties

10.3864/j.issn.0578-1752.2020.07.001

2019-07-31；

2019-11-14

泰山學者工程專項經(jīng)費（tsqn201812123）、山東省良種工程（2019LZGC016）、山東省自然科學基金（ZR2017MC004）

劉成，E-mail：lch6688407@163.com

（責任編輯 李莉）