小麦远缘杂交现状、抗病基因转移及利用研究进展

奥鹏网院作业

小麦远缘杂交现状、抗病基因转移及利用研究进展
刘成，韩冉，汪晓璐，宫文萍，程敦公，曹新有，刘爱峰，李豪圣，刘建军

（山东省农业科学院作物研究所/农业部黄淮北部小麦生物学与遗传育种重点实验室/小麦玉米国家工程实验室，济南 250100）

摘要：小麦近缘植物中含有丰富的抗病、抗逆和抗虫等基因，是小麦育种的优异基因源。通过远缘杂交可以将近缘植物优异基因转移给小麦，创制包括双二倍体或部分双二倍体、附加系、代换系和易位系等在内的小麦-近缘植物异染色体系。这些含小麦近缘植物血缘的异染色体系是研究物种染色体行为与进化、基因定位与作图的重要素材，也是拓宽小麦的遗传基础、抵御小麦重要病虫害、增加小麦产量和提升小麦品质的重要物质基础。为了更加清晰地了解小麦远缘杂交概况及小麦近缘植物抗病基因向小麦的转移，也为今后小麦远缘杂交研究和种质资源的开发利用提供参考，文中对小麦族物种分类、小麦远缘杂交的定义与意义、小麦族山羊草属、黑麦属、偃麦草属、簇毛麦属、冰草属、大麦属、披碱草属、赖草属、新麦草属以及旱麦草属物种与小麦远缘杂交现状和异染色体系创制情况进行了概括，并对来源于小麦近缘植物被正式命名的17个抗条锈病基因、35个抗叶锈病基因、30个抗秆锈病基因、41个抗白粉病基因、3个抗赤霉病基因、1个抗麦瘟病基因、1个抗叶枯病基因、1个抗颖枯病基因、4个抗褐斑病基因、2个抗眼斑病基因、1个抗梭条花叶病基因、2个抗线条花叶病基因和2个抗禾谷类黄矮病基因向小麦的转移情况及其所在染色体的位置信息进行了归纳。小麦-黑麦1RS·1BL易位系、1RS·1AL易位系和小麦-偏凸山羊草2NS/2AS易位系等抗病优良种质的育成与利用在世界小麦育种史上做出了突出贡献，然而，这仅仅得益于对少数抗病基因的利用。与目前已经被命名的基因数量相比，被利用到小麦育种中的抗病基因相对较少。文中分析了当前已命名抗病基因利用情况比例偏低的原因，并对今后如何利用这些抗病基因提出了建议。同时，还列举了已克隆的源自小麦近缘植物的抗病基因，并对克隆这些基因的方法以及今后可能的研究热点进行了分析，认为加强无遗传累赘的小麦-近缘植物易位系的创制与应用仍可能是今后小麦育种材料创新与新品种培育的一个重要发力点。

关键词：小麦；远缘杂交；异染色体系；抗病基因；衍生品种

1 小麦族分类
小麦族（Triticeae）有300多个物种，包含小麦属（Triticum）、山羊草属（Aegilops）、黑麦属（Secale）、偃麦草属（Elyrigia）、簇毛麦属（Dasypyrum）、冰草属（Agropyrum）、大麦属（Hordeum）、披碱草属（Elymus）、赖草属（Leymus）、新麦草属（Psathyrostachys）、旱麦草属（Eremopyrum）、类大麦属（Crithopsis）、无芒草属（Henrardia）、异形花属（Heteranthelium）、棱轴草属（Taeniatherum）、鹅观草属（Roegneria）、拟鹅观草属（Pseudoroegneria）和澳麦草属（Australopyrum）等，基本染色体组包含A—W和2个尚未确定的染色体组X和Y，表现出遗传变异的多样性[1-2]。小麦的近缘植物具有抗病[3-5]、抗虫[4-5]、抗旱[6]、抗寒[7-8]、耐盐[6,9-10]等优良性状，是小麦遗传改良的宝贵基因资源库[1-3]。

2 小麦远缘杂交现状
远缘杂交是亲缘关系较远的（包括生物学规定的不同“种”间、“属”间）以及亲缘关系更远的物种间杂交的统称[11-12]。小麦族中，小麦与黑麦、小麦与偃麦草、小麦与山羊草以及不同小麦种间的杂交均属远缘杂交，而生物学上规定的“种”以内的不同变种或品种间的杂交则统称为近缘杂交[11]。将近缘植物与小麦杂交，不仅可以将其优异基因导入小麦进行遗传改良[12-15]，还可以用于基因及染色体作图[16-17]、染色体行为及进化[18-19]等研究。自18世纪，科学家们就零星开始了小麦远缘研究[20]。19世纪以来，国内外科学家们在小麦远缘杂交方面做了大量工作，不同小麦种间[21-22]、小麦与山羊草属[2,23-26]、黑麦属[2,20,26-29]、偃麦草属[2,11,26,30-32]、簇毛麦属[2,26,33-36]、冰草属[26,37-39]、大麦属[40-42]、披碱草属[43-45]、赖草属[26,46-48]、新麦草属[49-51]、旱麦草属[52-53]等属物种远缘杂交成功的结果陆续被报道出来。目前，除类大麦属、拟鹅观草属、无芒草属、异形花属、棱轴草属、澳麦草属物种外，其余小麦族各属物种均已有与小麦杂交成功的报道。

3 小麦-近缘植物异染色体系创制情况
包括小麦-近缘植物双二倍体或部分双二倍体、附加系、代换系和易位系等在内的异染色体系，是向小麦转移近缘植物优异基因的桥梁和物质基础[5,54-55]。目前，小麦种间[21-22]、小麦与山羊草属[2,55-57]、黑麦属[2,58-60]、偃麦草属[2,26,31,61-63]、簇毛麦属[2,33-35,64]、冰草属[2,65-67]、大麦属[68-70]、披碱草属[71-73]、赖草属[2,74-76]、新麦草属[2,77-80]等属物种异染色体系创制成功的结果如雨后春笋般被报道。目前，虽然已有大量的小麦-近缘植物异染色体系被创制出来，然而，被直接用于小麦育种的小麦-近缘植物染色体易位系的比例还比较低。

4 近缘植物抗病基因向栽培小麦转移情况
迄今为止，小麦与近缘植物杂交成功的报道已有数百个[5,11,19,26,31,81]，其中大部分与小麦五大主要病害抗病基因转移有关[5,82]。目前，被国际小麦新基因命名委员会正式命名的抗小麦条锈病、叶锈病、秆锈病、白粉病和赤霉病的基因个数分别为82、79、60、65和7个，其中，来源于小麦近缘植物的基因个数分别有17（表1）、35（表2）、30（表3）、41（表4）和3个（表5），分别占被正式命名基因的20.7%、44.3%、50.0%、63.1%和42.9%。

此外，被正式命名的抗麦瘟病、叶枯病、颖枯病、褐斑病、眼斑病、梭条花叶病、线条花叶病和禾谷类黄矮病基因分别为8、18、3、7、3、1、3和3个，其中，来源于小麦近缘植物的基因个数分别有1、1、1、4、2、1、2和2个（表6），分别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 抗条锈病基因向小麦转移情况
来源于小麦近缘植物的抗条锈病基因有17个（表1），包括来自顶芒山羊草的Yr8、偏凸山羊草的Yr17、拟斯卑尔脱山羊草的Yr19、粗山羊草的Yr28、粘果山羊草的Yr37、沙融山羊草的Yr38、卵穗山羊草的Yr40、三芒山羊草的Yr42、小伞山羊草的Yr70、栽培黑麦的Yr9、中间偃麦草的Yr50、硬粒小麦的Yr7、Yr24和Yr53以及野生二粒小麦的Yr15、Yr35和Yr36。

表1 小麦近缘植物抗条锈病基因向小麦转移情况

Table 1 Stripe rust resistance genes transferred from wild relatives to wheat


4.2 抗叶锈病基因向小麦转移情况
来源于小麦近缘植物的抗叶锈病基因有35个（表2），包括来自小伞山羊草的Lr9和Lr76、粗山羊草的Lr21、Lr22a、Lr22b、Lr32、Lr39和Lr42、拟斯卑尔脱山羊草的Lr28、Lr35、Lr36、Lr47、Lr51和Lr66、偏凸山羊草的Lr37、粘果山羊草的Lr54、沙融山羊草的Lr56、卵穗山羊草的Lr57、钩刺山羊草的Lr58、柱穗山羊草的Lr59、短穗山羊草的Lr62、栽培黑麦的Lr25、Lr26和Lr45、长穗偃麦草的Lr19和Lr29、彭提卡偃麦草的Lr24、中间偃麦草的Lr38、粗穗披碱草的Lr55、栽培二粒小麦的Lr14a、野生二粒小麦的Lr53和Lr64、硬粒小麦的Lr23、一粒小麦的Lr63和提莫非维小麦的Lr50。

4.3 抗秆锈病基因向小麦转移情况
来源于小麦近缘植物的抗秆锈病基因有30个（表3），包括来自顶芒山羊草的Sr34、偏凸山羊草的Sr38、拟斯卑尔脱山羊草的Sr32、Sr39和Sr47、希尔斯山羊草的Sr51、卵穗山羊草的Sr53、栽培黑麦的Sr27、Sr31、Sr50和Sr59、簇毛麦的Sr52、彭提卡偃麦草Sr24和Sr25、长穗偃麦草的Sr26和Sr43、中间偃麦草的Sr44、野生二粒小麦的Sr2、Sr9d、Sr9e、Sr13、Sr14和Sr17、一粒小麦的Sr21、Sr22、Sr35和Sr60、硬粒小麦的Sr12以及提莫非维小麦的Sr36和Sr37。

表2 小麦近缘植物抗叶锈病基因向小麦转移情况

Table 2 Leaf rust resistance genes transferred from wild relatives to wheat


表3 小麦近缘植物抗杆锈病基因向小麦转移情况

Table 3 Stem rust resistance genes transferred from wild relatives to wheat


4.4 抗白粉病基因向小麦转移情况
来源于小麦近缘植物的抗白粉病基因有41个（表4），包括来自拟斯卑尔脱山羊草的Pm12、Pm32和Pm53、粗山羊草的Pm19、Pm34、Pm35和Pm58、高大山羊草的Pm13、希尔斯山羊草的Pm57、卵穗山羊草的Pm29、栽培黑麦的Pm7、Pm8、Pm17、Pm20和Pm56、簇毛麦的Pm21、Pm55和Pm62、中间偃麦草的Pm40和Pm43、彭提卡偃麦草的Pm51、一粒小麦的Pm1b和Pm4d、野生一粒小麦的Pm25、波斯小麦的Pm4b、栽培二粒小麦的Pm4a、Pm5a、Pm49和Pm50、野生二粒小麦的Pm16、Pm26、Pm30、Pm36、Pm41、Pm42和Pm64、硬粒小麦的Pm3h、乌拉尔图小麦的Pm60以及提莫非维小麦的Pm6、Pm27和Pm37。

表4 小麦近缘植物抗白粉病基因向小麦转移情况

Table 4 Powdery mildew resistance genes transferred from wild relatives to wheat


4.5 抗赤霉病基因向小麦转移情况
来源于小麦近缘植物的抗赤霉病基因有3个（表5），包括来自大赖草的Fhb3、柯孟披碱草（也有科学家称其为鹅观草）的Fhb6和来自彭提卡偃麦草的Fhb7。

4.6 其他抗病基因向小麦转移情况
来源于小麦近缘植物的五大主要病害之外的抗病基因有14个（表6），包括来自栽培二粒小麦的抗麦瘟病基因Rmg7、粗山羊草的抗叶枯病基因Stb5、抗颖枯病基因Snb3和抗褐斑病基因Tsr3、野生二粒小麦的抗褐斑病基因Tsr2和Tsr7、圆锥小麦的抗褐斑病基因Tsr5、偏凸山羊草的抗眼斑病基因Pch1、簇毛麦的抗眼斑病基因Pch3和抗梭条花叶病毒基因Wss1、中间偃麦草的抗线条花叶病基因Wsm1、Wsm3以及抗禾谷类黄矮病基因Bdv2和Bdv3。

表5 小麦近缘植物抗赤霉病基因向小麦转移情况

Table 5 Fusarium head blight resistance genes transferred from wild relatives to wheat


表6 小麦近缘植物抗麦瘟病等基因向小麦转移情况

Table 6 Wheat blast resistance gene and other disease resistance genes transferred from wild relatives to wheat


—表示基因已命名但无文献发表（McIntosh R A与Worland A K，私人通讯）

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

5 小麦近缘植物抗病基因的利用
在小麦远缘杂交种质应用方面，对世界小麦育种做出突出贡献的当属小麦-黑麦1RS·1BL易位系。1RS染色体上由于含Yr9、Pm8、Lr26和Sr31等基因，受到了广大育种工作者的普遍青睐[14,203-206]，国外育种家们利用该易位系及其衍生系作亲本，育成了山前麦、高加索、无芒一号和洛夫林13等高产抗病小麦，被全世界几十个国家作为骨干亲本应用，育成了一大批优异小麦新品种[207-210]，在推动小麦品种的更新换代中发挥了重要作用[203,211-212]。除了1RS·1BL易位系，国外育种家们还培育出了含1RS·1AL易位系的Amigo等品种，并以此为骨干亲本，培育出了含该易位系的Zhytnytsa、Nota和Duma[203]、Columbia、Etude和Rastavitsa[213]、TAM107、TAM303、TAM305、AG Robust、Fannin、N96L9970[214-216]和Helami-105等小麦新品种/系[217]，在美国、墨西哥和欧洲等国家推广应用。近年来，国际玉米小麦改良中心（CYMMYT）以该易位系为亲本育成了CM409和CM451等一批小麦新品种/系（刘彩云，私人通讯）。

据报道，19世纪后期中国约70%小麦品种含1RS·1BL易位系[204-205]，其中，为中国小麦育种做出突出贡献的矮孟牛（Ⅱ型、Ⅳ—Ⅶ型）、周麦22、周8425B和石4185等骨干亲本材料均含有1RS染色体。近年来，由于新的致病生理小种的产生与流行，使得Yr9和Pm8等基因的抗性迅速丧失[177,218-219]，加上育种家们在育种过程中注意杂交亲本的遗传多样性，因此，该易位系在中国小麦中的比例明显下降[220]。虽然Yr9等基因的抗性已经丧失[177,218-219]，但近期的研究发现，不同黑麦来源的1RS·1BL易位系可能含有不同的抗病等位基因[14,221]，即表明不同黑麦来源的该易位系仍能在小麦育种中发挥重要作用。尤其是近年来，不含黑麦碱但仍具有良好抗病性的1RS·1BL易位系的创制[222-224]，为小麦育种提供了新的育种资源。

除含1RS染色质的育种材料外，含Yr17、Lr37和Sr38的小麦-偏凸山羊草2NS/2AS易位系对世界小麦育种也做出了突出贡献。以该易位系为抗源育成的Mace（还含Lr14a和Wsm1）[225-226]、Jagger、Madsen、Overley、SY Gold、Trident、EGA Eaglehawk和Espada等小麦品种在美国、澳大利亚和欧洲等国家推广应用[225-229]。研究发现，源自中国10余个省份69个小麦品种中的49%含该易位系[230]。此外，据报道，川育18、川麦25和川麦39等[231]、新麦19、济麦20、济麦21和师栾02-1[232]、兰考906、西农739、陕872和小偃216等品种/系[233]含有该易位系。近期研究发现，济麦20和济麦21中不含但中麦175中含有Lr37等基因，然而Lr37已对中国当前叶锈生理小种表现为感病[234]。此外，Yr17在澳大利亚、欧洲和中国的条锈抗性已经完全或部分丧失[228,235-236]。因此，今后在育种中应减少对该易位系的利用。

自19世纪以来，中国在小麦远缘杂交领域研究一直处在世界前列。中国科学家先后将偃麦草[31,61-63,81,237-244]、黑麦[58-60,245-252]、簇毛麦[35,253]和冰草[254-255]等种质转移给了小麦，育成了一大批远缘杂交新材料。在对这些小麦远缘杂交种质利用方面，取得了举世瞩目的研究成果，培育出的抗条锈病的小偃系列品种及其衍生品种[256-258]、普冰系列及其衍生品种（张锦鹏，私人通讯）和陕麦号及西农号小麦[259-260]、抗黄矮病的张春号、临抗号、晋麦号小麦[261-262]和黑小麦品种[263-264]、抗白粉病的南农号小麦及其衍生品种[265-267]、抗条锈和白粉等病害的川农系列小麦及其衍生品种[268-269]和远丰号小麦[270]等在中国大面积推广应用。上述品种抗病性来源主要为Yr9、Yr41、Yr50、Pm21、Bdv2、Bdv3和尚未被证明命名的少数几个基因。

除此之外，值得一提的是，中国科学家分别将来自彭提卡偃麦草抗赤霉病基因Fhb7[189,271]和长穗偃麦草的尚未被命名的赤霉病基因[272-273]分别转移到小麦，创制了一批小麦-偃麦草染色体易位系，并将其导入中国主栽小麦，培育出了一批赤霉抗性达到中抗水平正在参加区域试验的小麦新品系，有望对小麦抗赤霉病育种发挥重要作用。

6 结论与展望
目前，众多个有明显育种价值的小麦-近缘植物染色体易位系/渐渗系被成功创制出来[4-5]，并且有近140个抗病新基因被正式命名（表1—表6），但就基因利用状况来看，被利用到小麦抗病育种上的基因的比例还比较低。其原因可能是：（1）部分基因的抗性已经/正在丧失，例如Yr7、Yr9和Yr17等[274]，Lr21、Lr26和Lr37等[234]，Sr31、Sr34和Sr38等[275]，Pm8、Pm17和Pm19等[276]；（2）部分易位染色体具有遗传累赘，例如Yr8[277]、Yr38/Lr56[278]、Pm12和Pm20[179]等基因所在近缘植物染色体臂。因此，在今后的研究中，应该做到：（1）加强二倍体和四倍体小麦抗病基因向栽培小麦的导入与利用；（2）加强对有遗传累赘效应易位系的染色体工程诱导。通过抗病基因转育，创制出更多的抗病种质资源并对其进行育种学评价。目前，已有多个研究团队在开展这项工作[267,279]；（3）加强对无遗传累赘且具优异抗性易位系[267,279-280]的利用工作。

克隆抗病基因是研究其抗病机理的基础。目前，从小麦近缘植物中克隆出的抗病基因主要包括Yr15[281]、Yr28[282]、Yr36[283]、Lr21[284]、Sr13[285]、Sr21[286]、Sr35[287]、Sr60[288]、Pm21[289-290]等。其中，除Yr28和Pm21[85,162]外，其他几个基因均源自二倍体或四倍体小麦[96,98,100,141,143-145]。由于缺乏参考基因组信息，目前，从小麦-近缘植物（这里指非小麦属物种）染色体易位系中克隆抗病基因还有一定困难。当前，克隆这些基因可以利用不同居群的抗病性不同的同一小麦近缘种进行杂交（或利用诱变技术创制突变体），配置抗感分离群体进行基因定位与克隆，例如Yr28[282]和Pm21[289]的克隆；还可以创制更多的小麦-近缘植物染色体结构变异体，将抗病基因定位到近缘植物某一染色体小片段上，进而利用已克隆的模式植物抗病基因所在染色体区间与上述小片段区间进行基因共线性分析与确证，同源克隆近缘植物的抗病基因。近期笔者及其合作者们利用该方法克隆了Pm12（待发表）。

从理论研究上讲，随着小麦族物种基因组测序工作的陆续开展与完成，能够用于抗病基因克隆的参考基因组信息越来越多，今后克隆小麦近缘植物抗病基因将会变得越来越容易，因此，这些基因的抗病调控机制以及不同物种共线性抗病基因的进化可能将成为新的研究热点。从应用研究上讲，小麦-黑麦1RS·1BL易位系、1RS·1AL易位系和小麦-偏凸山羊草2NS/2AS易位系等抗病优良种质的育成与利用在世界小麦育种史上做出了突出贡献，然而，这仅仅得益于对少数抗病基因的利用。虽然目前被利用到小麦育种中的抗病基因相对较少，但加强无遗传累赘的小麦-近缘植物易位系的创制与应用仍可能是今后小麦育种材料创新与新品种培育的一个重要发力点。

致谢：堪萨斯州立大学Friebe B教授、德克萨斯农工生命研究和推广中心Liu SY教授、北达科他州立大学Cai XW教授、悉尼大学Zhang P博士和李建波博士、阿德莱德大学Dundas I博士、John Innes Centre的Griffiths S研究员、CYMMYT刘彩云博士后、乌克兰国家种子与品种调查中心Motsnyi I研究员、电子科技大学杨足君教授、中国农业科学院张锦鹏研究员、西北农林科技大学王长有教授、鲁东大学崔法教授、山东农业大学鲍印广教授在不同国家小麦品种所含外源染色质信息搜集中给予的大力帮助，在此表示感谢。

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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)

Abstract: 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 Aegilops, Secale, Thinopyrum, Dasypyrum, Agropyron, Hordeum, Elymus, Leymus, Psathyrostachys and Eremopyrum to 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-Aegilops ventricosa 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.

Key words: wheat; wild hybridization; chromosome line; disease resistance gene; derived varieties

开放科学（资源服务）标识码（OSID）：width=42.5,height=42.5

doi: 10.3864/j.issn.0578-1752.2020.07.001

收稿日期：2019-07-31；

接受日期：2019-11-14

基金项目：泰山学者工程专项经费（tsqn201812123）、山东省良种工程（2019LZGC016）、山东省自然科学基金（ZR2017MC004）

联系方式：刘成，E-mail：lch6688407@163.com

（责任编辑 李莉）