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小麦抗条锈新基因YrTp1和YrTp2的发现和分子标记定位
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摘要
条锈病是小麦的主要病害之一,在我国曾多次流行造成小麦严重减产。应用抗病品种是防治小麦条锈病最为经济、有效和对环境安全的途径。小麦条锈菌新小种的不断产生和发展,使小麦品种的抗锈性屡屡“丧失”,导致我国小麦条锈病周期性大流行。目前只有Yr5、Yr10等极少数抗条锈病基因对我国小麦条锈病优势生理小种条中30、31、32号仍表现免疫或高抗。其中强优势小种条中32号毒性谱更宽,致病范围更广,相对寄生适合度更高,已使我国大部分麦区的主栽品种沦为感病,发病面积占播种面积的90%,它对Yr1、Yr2、Yr3、Yr4、Yr6、Yr9、Yr15、Yr27、YrA、Yralba、YrCle、YrCV、YrGaby、YrRes、YrSD、YrSpP、YrSO等抗病基因均有毒性,对生产品种和抗源均有致病性。因此,我国的小麦生产和育种面临抗源危机。
     利用外源抗性基因是小麦抗条锈育种的有效措施。继来自黑麦的Yr9在生产上发挥巨大作用之后,抗性来自中间偃麦草(Thinopyrum intermedium Barkworth & Dewey)的无芒中4已成为国内外小麦抗条锈育种的主要抗源。鉴于利用野生近缘种(属)抗源已经成为当前抗病育种的主攻方向之一,而国内育种上利用的主要抗源只有无芒中4、92R系列和贵农21等少数几个,为避免抗源利用单一化而重蹈1BL/1RS及繁6的覆辙,寻找新的抗源、尤其从小麦近缘种(属)中发掘新的有效抗病基因,开展一些前瞻性的工作迫在眉睫。
     分子标记技术为新基因的发现和定位研究提供了新的工具。R?der et al (1998)构建了包含279个标记位点的小麦微卫星(Simple Sequence Repeats, SSR)图谱,Somers et al(2004)将来自不同研究小组的四张遗传图谱整合为一张包含1235个微卫星标记的高密度遗传图谱,为小麦目的基因的标记、定位提供了极大的方便。目前在已建立分子标记的15个抗条锈基因中,10个找到了SSR标记。
     从普通小麦(Triticum aestivum L.)与十倍体长穗偃麦草(Thinopyrum ponticum (Host) Liu & Wang)杂交创造的125个农艺性状较好的后代株系中,筛选出了基本稳定的抗条锈株系20个,其中有免疫的、也有其它低反应型类型和迟慢锈类型。2001年,在甘肃天水田间鉴定发现已经稳定的后代材料A-3对条锈病免疫;2001-2005年连续观察,结果相同;在苗期鉴定,对混合菌免疫;成株期分小种鉴定,对水源11致病类型-3、水源11致病类型-7、水源11致病类型-14、条中29号、31号、32号等生理小种和混合菌免疫。A-3系谱为“晋2148///Fuhuko/R431//北京837/2”,其普通小麦亲本晋2148、Fuhuko、北京837对条中31、32号小种和
Stripe rust (Puccinia striiformis Westend f. sp. tritici), caused by Puccinia strriformis Westend f. sp. tritici Eriks is one of the most important disease of wheat (Triticum aestivum L.), which resulted in massive wheat yield loss in several epidemics in China. Loss of variety resistance to stripe rust pathogene is an important factor causing massive periodical epidemic of stripe rust in wheat production. Using resistant varieties is the most economic, effective and environment-friend method. Emergency and development of new races of stripe rust pathogene have led widely planted varieties into serious crisis of resistance losing. Currently, only very few genes, such as Yr5、Yr10,keep immune or highly resistant to high frequency virulent races CY30, CY31 and CY32, among which the main preponderant race CY32 has led major varieties into susceptible in China because of its wider virulence array and higher autoecious fitness. It has affected 90% of planting area. CY32 was virulent to Yr1、Yr2、Yr3、Yr4、Yr6、Yr9、Yr15、Yr27、YrA、Yralba、YrCle、YrCV、YrGaby、YrRes、YrSD、YrSpP、YrSO as well as most varieties and resistance resources used in production. Utilizing alien resistant resources was a powerful breeding mean for stripe rust resistance. After Yr9 derived from Secale cereale L. had taken a very important role in production for nearly 20 years, Zhong 4 derived from Triticum aestivum×Thinopyrum intermedium has become the main resistant resource for stripe rust resistance breeding all over the world. As there were only very few resistant resources in use currently, in order to avoid disaster loss of variety resistance like 1BL/1RS translocation and Fan 6 because of simplification of resistance resources it is in urgent need to search new resistant resources, especially to unlocking effective novel resistant genes from wheat relative species and genus.
     Molecular markers could facilitate the identification and location of novel genes. As so far, among 15 Yr genes, 10 Yr genes were linked with SSR markers.
     Twenty lines with resistance and good agronomic traits were screened out in 125 lines derived from T. aestivum×Th. Ponticum, most of which were immune and 2-3 were high resistant including one late-slow-rusting line. A line A-3 immune to current epidemic yellow rust races (CY29, CY30, CY31, CY32) was screened out in offspring of T. aestivum×Th. ponticum. Previous GISH analysis did not detect any
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