小麦新品种“山农20”抗病基因的分子检测

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李继发, 邓志英, 孙福来, 关西贞, 王延训, 田纪春. 小麦新品种“山农20”抗病基因的分子检测. 作物学报, 40(4): 611-621
LI Ji-Fa, DENG Zhi-Ying, SUN Fu-Lai, GUAN Xi-Zhen, WANG Yan-Xun, TIAN Ji-Chun. Resistance Genes of Wheat Variety Shannong 20 Identified by Diagnostic Molecular Markers. 作物学报, 40(4): 611-621 复制到剪切板

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作物学报编辑部

小麦新品种“山农20”抗病基因的分子检测

李继发^1,^*, 邓志英^1,^*, 孙福来², 关西贞¹, 王延训¹, 田纪春^1,^*

¹山东农业大学作物生物学国家重点实验室 / 山东省作物生物学重点实验室, 山东泰安 271018

²山东省滨州市种子管理站, 山东滨州 256600

^*通讯作者(Corresponding author): 田纪春, E-mail:jctian@sdau.edu.cn, Tel: 0538-8242040

收稿日期:2014-01-12

基金:本研究由国家自然科学基金项目(31171154), 山东省农业良种工程项目(鲁农良种字[2011]7号和[2012]213号)和国家转基因生物新品种培育重大专项(2013ZX08002-003)资助。

摘要

山农20是2011年和2012年分别通过国家黄淮南、北片审定的小麦高产多抗新品种, 在国家区试抗病性鉴定和生产中都表现出良好的抗黄淮麦区主要病害的特性。本研究利用与小麦抗白粉病、条锈病、叶锈病、纹枯病基因和抗赤霉病主效QTL紧密连锁的SSR、SCAR、STS等标记对该品种进行了分子检测, 发现山农20含有6个抗白粉病基因(Pm12、Pm24、Pm30、Pm31、Pm35和Pm36), 6个抗条锈病基因(Yr5、Yr9、Yr15、Yr24、Yr26和YrTp1), 2个抗叶锈病基因(Lr21和Lr26), 1个抗纹枯病基因(Ses1), 但未检测到抗赤霉病主效QTL。分子检测结果部分解释了山农20的优良抗病性, 也为利用分子标记辅助选择培育抗病稳产小麦新品种提供参考。

关键词: 山农20; 抗病性相关基因; 分子标记

Resistance Genes of Wheat Variety Shannong 20 Identified by Diagnostic Molecular Markers

LI Ji-Fa^1,^**, DENG Zhi-Ying^1,^**, SUN Fu-Lai², GUAN Xi-Zhen¹, WANG Yan-Xun¹, TIAN Ji-Chun^1,^*

¹State Key Laboratory of Crop Biology / Shandong Provincial Key Laboratory of Crop Biology, Shandong Agricultural University, Tai#cod#x02019;an 271018, China

² Binzhou Seed Management Station of Shandong Province, Binzhou 256600, China

Fund:

Abstract

Shannong 20 is a newly released winter wheat variety in the National Regional Trials for North and South Yellow-Huai River Valleys, which exhibits excellent resistance to major diseases of wheat. In this study, the possible resistance genes in Shannong 20 were identified by using SSR, SCAR, and STS markers tightly linked to resistance genes/loci against powdery mildew, stripe rust, leaf rust, sheath blight, and scab. A total of 15 resistance genes were detected, including six (Pm12,Pm24,Pm30,Pm31,Pm35, andPm36) for powdery mildew resistance, six (Yr5,Yr9,Yr15,Yr24,Yr26, andYrTp1) for stripe rust resistance, two (Lr21 andLr26) for leaf rust resistance, and one (Ses1) for sheath blight resistance. However, no resistance locus to scab was found. The result may partially explain the multi- resistance in Shannong 20, and provide some references for cultivating new varieties with disease resistance and stable yield by using molecular marker-assisted selection.

Keyword: Shannong 20; Disease resistance genes; Molecular markers

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我国从20世纪50年代开始进行小麦抗病常规育种, 主要针对条锈病。近年来, 除条锈病外, 许多原来的次要病害逐渐成为限制小麦生产的主要病害, 如白粉病、赤霉病、纹枯病和全蚀病等, 其中最主要的病害是条锈病, 其次是白粉病。由于小麦条锈病和白粉病分布广泛、病原菌生理小种复杂多变等特点, 常常导致品种抗性频繁丧失, 因此, 这些病害的防治任务十分艰巨。除了利用现有抗病基因进行常规育种外, 挖掘新抗病基因和在同一品种中聚合多种抗病基因是重要的育种策略。另外, 随着分子标记的广泛应用和标记技术的发展, 获得了越来越多的抗病基因分子标记, 为分子标记辅助选择(MAS), 加快目标基因的聚合育种提供了便利。

目前, 已有60多个小麦抗白粉病基因被定位到47个位点( Pm1~ Pm47), 分布在除3A、4D以外的各条染色体上^{[ 1, 2]}。其中, Pm38和 Pm39为白粉病成株抗性基因^{[ 3]}, 其余抗性基因均为病原菌生理小种专化性的主效基因; 由于生理小种的不断变异, 大部分已丧失抗性^{[ 4]}。

国际上已命名51个抗条锈病主效基因, 涉及48个位点( Yr1~ Yr48), 除 Yr11~Yr14外均被定位于特定的染色体上^{[ 5]}。其中, Yr3位点有 Yr3a、 Yr3b和 Yr3c三个复等位基因, Yr4位点有 Yr4a和 Yr4b两个复等位基因, Yr11、 Yr12、 Yr13、 Yr14、 Yr16、 Yr18、 Yr29、 Yr30、 Yr36、 Yr39、 Yr46和 Yr48为成株抗性基因, 其他39个为苗期抗性基因^{[ 5]}。还有72个条锈病成株抗性QTL被定位在除1A、1D、3D和7A染色体外的其余各个染色体上^{[ 4]}。

目前已正式命名的小麦抗叶锈病基因有68个, 其中大多数为小种特异的主效抗叶锈病基因^{[ 6]}, 已报道39个分子标记与抗叶锈病基因连锁或共分离^{[ 7]}。

近年来, 小麦纹枯病在我国特别是黄淮麦区趋向严重, 已上升为我国小麦主要病害之一, 但关于抗纹枯病的遗传研究报道较少。张小村等^{[ 8]}利用2个重组自交系群体RIL-8和RIL-SES, 对抗纹枯病性QTL进行分析, 检测到1个SSR标记(Xgwm526)与纹枯病抗性具有关联性, 与纹枯病抗病基因 Ses1的遗传距离为27.9 cM。

小麦赤霉病主要发生在温暖潮湿和半潮湿地区, 在我国主要发生在长江中下游冬麦区和东北春麦区, 近年来随着全球气候变暖、灌溉条件的改善及耕作制度的变化, 有进一步蔓延到黄淮麦区的趋势^{[ 9]}。目前, 除7D染色体外的其他所有的染色体上都检测到抗赤霉病QTL的存在^{[ 10]}, 其中位于3B、5A和6B染色体上的QTL在苏麦3号、望水白、Wuhan、Nyubai、Frontana、CM82036和DH181中被检测到, 并且效应较大, 是小麦抗赤霉病的主效QTL。但到目前为止, 世界上还没有发现对赤霉病完全免疫的小麦品种。

尽管已获得一些主要抗病性的QTL及其分子标记, 但在实际生产中应用却很少。目前生产上应用的小麦品种基本上都是常规育种方法培育而成, 由于其抗病机制及其携带的抗病基因都不明确, 致使很难进一步研究和利用很多优良品种。因此, 利用分子标记分析重要品种的抗病基因, 不仅为优异抗病基因的鉴定和利用MAS技术进行多基因聚合提供技术基础, 而且有助于跟踪重要或骨干亲本所含抗病基因的来源和遗传, 为在育种中有效利用不同抗病基因及其合理布局奠定基础。

本课题组培育的山农20属于高产多抗小麦新品种, 其亲本为PH82-2-2和954072。2010年和2011年分别通过国家黄淮南片和黄淮北片试验程序, 是目前很少见的广适性“双国审”小麦新品种。该品种抗逆性好、抗病性强、品质优、高产、稳产、适应性强, 适宜在包括苏北、皖北、河南、山西、陕西、山东、河北7个省份和地区的黄淮麦区推广种植。在近几年的推广种植中, 通过田间接种鉴定, 该品种表现出对多种小麦主要病害的优良抗性, 但是其抗病机制尚不清楚。本研究针对小麦抗白粉病、抗条锈病、抗叶锈病、抗纹枯病和抗赤霉病等相关性状, 选取已开发的与有关抗性基因或主效QTL连锁距离相对较近和稳定的SSR、STS、SCAR等分子标记, 对山农20及其亲本进行抗病性和抗病基因的分子鉴定, 旨在明确该品种抗病基因及其抗性机制, 为对其进一步研究和利用奠定基础, 也为利用MAS方法培育具抗性和稳产小麦新品种提供参考。

1 材料与方法

1.1 供试材料及其抗病性评价

山农20及其亲本PH82-2-2 (母本)和954072 (父本)均由本课题组保存。

2008—2010年, 由中国农业科学院植物保护研究所根据全国农业技术推广服务中心《农技种函[2008]301号》文件要求, 分别对国家冬小麦品种区域试验品种进行人工接种抗病性鉴定, 其中供试条锈菌、叶锈菌、白粉菌均为混合优势小种, 赤霉菌和纹枯病菌以强致病力菌株接种。条锈病鉴定菌系包括CY29、CY30、CY31、CY32等生理小种; 叶锈病接种菌系以PHT、THT、PHK生理小种为主, 同时混有THP和FHT; 白粉病菌菌系包括V1、V3a、V3b、V3c、V3d、V3e、V3f、V4a、V4b、V5、V6、V7、V8、V17、V19、V25和V35; 赤霉菌菌株为F15、F34、F0301和F0609; 纹枯病菌菌株为R0301。各年度均按《小麦抗病虫性评价技术规范》(NY/T 1443-2007)进行接种和抗病性鉴定, 在4月上中旬分别进行田间和室内接种, 待对照品种充分发病后调查病情。

为了更准确地鉴定山农20在生产种植中的抗病性, 2010—2013年分别在山东的济宁、泰安、菏泽, 安徽的宿州、濉溪、阜阳, 河南的商丘、郑州、濮阳及江苏的徐州、新沂、连云港安排了0.67 hm²生产示范种植, 并按照《小麦白粉病测报调查规范》(NY/T 613-2002)、《小麦条锈病测报技术规范》 (GB/T 15795-2011)、《小麦叶锈病测报调查规范》(NY/T 617-2002)、《小麦纹枯病测报调查规范》(NY/T 614-2002)和《小麦赤霉病测报技术规范》(GB/T 15796-2011)调查田间病害, 反应型分为免疫(0)、高抗(1)、中抗(2)、中感(3)和高感(4)。每年每省选3点鉴定抗病性, 按抗性最弱点的抗病表现记载结果。

1.2 抗病基因的分子鉴定

采用改良CTAB法(http://www.triticarte.com.au/)提取山农20及其两个亲本的基因组DNA, 用1%琼脂糖凝胶电泳检测DNA的浓度和质量。选用与小麦抗白粉病、抗条锈病、抗叶锈病、抗纹枯病和抗赤霉病基因或QTL紧密连锁或共分离的47个分子标记(见附表)进行分子鉴定。所有标记引物均由英潍捷基(上海)贸易有限公司合成。

PCR体系为20 μL, 含10×buffer (含Mg²⁺) 2 μL、dNTP 1.6 μL (10 mmol L^-1)、每条引物0.4 μL、模板DNA 50 ng、1 U Taq聚合酶。PCR扩增程序为 95℃预变性5 min; 95℃变性50 s; 退火1 min (退火温度因具体引物而定), 72℃延伸1 min, 35个循环; 72℃延伸

10 min, 8℃保存。采用8%非变性聚丙烯酰胺凝胶电泳(PAGE)分离扩增产物, 硝酸银染色, 显影后观察并照相。

2 结果与分析

2.1 国家区试和生产种植中的抗病性表现

国家区试中, 山农20对5种病害的抗病性在黄淮南片和北片表现不尽相同, 年度间也有差异, 但总体呈现对白粉、条锈和叶锈病有良好抗性, 多为免疫或高抗、或慢锈病, 而对赤霉病和纹枯病抗性较低, 甚至高感(表1)。

连续3个生长季在山东、安徽、河南和江苏田间自然感病条件下, 对山农20进行抗病性鉴定。总体来看, 白粉病由免疫到轻度发生; 条锈和叶锈病由免疫至高抗; 纹枯病由中抗至中感; 赤霉病发病较重, 表现为中抗至高感。2012年病害发生较重, 赤霉病表现最重, 其次是纹枯病。山东的抗病表现最好, 安徽、河南和江苏抗病性依次减弱(表2)。

表1 山农20在国家区试中的抗病性鉴定结果 Table 1 Resistance evaluation of Shannong 20 to main diseases of wheat in China national region trials

表2 自然感病条件下山农20对5种主要病害的抗病性鉴定 Table 2 Resistance of Shannong 20 to five major wheat diseases under natural disease fields

2.2 抗病基因的分子检测

2.2.1 抗白粉病基因利用与抗白粉病基因紧密连锁的分子标记, 对15个基因位点检测, 结果山农20在6个位点( Pm12、 Pm24、 Pm30、 Pm31、 Pm35和 Pm36)有扩增产物(图1), 且这些特异带在其一个亲本或双亲中也被检测到(表3), 说明山农20可能携带这6个抗白粉病基因。在 Pm16位点, 仅PH82-2-2有特异扩增, 而在山农20和954072中未检测到该片段(表3), 可能 Pm16基因未能从PH82-2-2传递到山农20。而 Pm36基因位点则是通过亲本954072传递给山农20的; Pm12、 Pm24、 Pm30、 Pm31、 Pm35基因位点在2个亲本和山农20中都含有; 其余8个基因位点在2个亲本及山农20中都不存在。

2.2.2 抗条锈病基因利用与抗条锈病基因紧密连锁的分子标记, 对12个基因位点检测, 结果在6个位点( Yr5、Yr9、Yr15、Yr24、Yr26和 YrTp1)有扩增产物(图1), 且这些特异带在其一个亲本或双亲中均被检测到(表4), 说明山农20可能携带这6个抗条锈病基因。其中, Yr5和 Yr15基因位点由亲本954072传给山农20; Yr9、Yr24、Yr26和 YrTp1基因位点在2个亲本和山农20都含有, 而在2个亲本和山农20中都未检测到其余6个基因位点的特异片段。

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图1 不同抗病基因分子标记的扩增结果各泳道自左向右依次为DL2000分子量标记、山农20、PH82-2-2和954072。A~F: 抗白粉病基因 Pm12(A)、 Pm16(B)、 Pm24(C)、 Pm30(D)、 Pm35(E)和 Pm36(F)的分子标记带型; G~L: 抗条锈病基因 Yr5(G)、 Yr9(H)、 Yr15(I)、 Yr24(J)、 Yr26(K)和 YrTp1(L)的分子标记带型; M~O: 抗叶锈病基因 Lr1(M)、 Lr21(N)和 Lr26(O)的分子标记带型。Fig. 1 PCR profiles of molecular markers for different resistance genesFrom left to right, the four lanes in each panel were loaded with DL2000 marker and genome DNA templates extracted from Shannong 20, PH82-2-2, and 954072, respectively. A-F: Banding patterns of markers for powdery mildew resistance genes Pm12(A), Pm16(B), Pm24(C), Pm30(D), Pm35(E), and Pm36(F); G-L: Banding patterns of markers for stripe rust resistance genes Yr5(G), Yr9(H), Yr15(I), Yr24(J), Yr26(K), and YrTp1(L); M-O: Banding patterns of markers for leaf rust resistance genes Lr1(M), Lr21(N), and Lr26(O).

表3 抗白粉病基因分子检测结果 Table 3 Molecular detection results of powdery mildew resistance genes

表4 抗条锈病基因分子检测结果 Table 4 Molecular detection results of stripe rust resistance genes

2.2.3 抗叶锈病基因利用与抗叶锈病基因紧密连锁的分子标记, 对12个基因位点检测, 结果在2个位点( Lr21和 Lr26)有扩增产物(图1), 且这些特异带在双亲中也被检测到(表5), 说明山农20可能携带这2个抗叶锈病基因。对于 Lr1基因位点只在PH82-2-2扩增出特异性片段(见附图), 但未传递给山农20。在山农20和2个亲本均未扩增出其余9个

基因位点的特异性片段。

2.2.4 抗纹枯病基因对于与抗纹枯病基因Ses1紧密连锁的SSR标记Xgwm526, 在山农20及其双亲中均扩增出约140 bp的特异性片段(图2), 推测山农20可能含有 Ses1基因。

表5 抗叶锈病基因分子检测结果 Table 5 Molecular detection results of leaf rust resistance genes

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	图2 Ses1基因的分子标记在山农20及其亲本中的扩增结果M: DL2000; 1: 山农20; 2: PH82-2-2; 3: 954072。Fig. 2 Banding patterns on Ses1locus in Shannong 20 and its parents amplified with specific primersM: DL2000; 1: Shannong 20; 2: PH82-2-2; 3: 954072.

2.2.5 抗赤霉病主效QTL 利用5个已开发SSR标记, 对山农20、PH82-2-2和954072进行抗赤霉病QTL检测, 结果均未扩增出相应标记的特异性片段, 说明山农20及其亲本没有这些主效QTL。

3 讨论

小麦品种抗病性鉴定最经典的方法是使用抗病菌株的田间鉴定后进行基因推导, 但这一方法易受外界环境条件、遗传背景及基因×环境和基因×基因互作的影响, 而且有些基因没有相对应的致病菌株, 因此, 这一方法在检测某些新的高抗品种时, 难以确定是否含有新的抗病基因。利用与抗病基因/QTL紧密连锁的分子标记, 可以弥补基因推导的不足, 而且随着标记开发, 可及时鉴定和检测新的抗病基因及其遗传传递。山农20是一个农艺性状优良的冬小麦新品种, 除产量和品质好以外, 其突出的特点是对多种小麦主要病害表现广谱抗病性, 在多年的国家区试鉴定和生产鉴定中, 表现对白粉病和条锈病免疫、抗叶锈病、中感纹枯病和赤霉病。利用抗病性鉴定和分子标记检测方法, 明确山农20可能携带的抗病基因, 将对该品种的推广应用和选育抗病性更好的新品种有重要意义。

目前, Pm12、 Pm13、 Pm16、 Pm20、 Pm21、 Pm30在中国对小麦白粉病表现高抗至免疫, 而 Pm1、 Pm2、 Pm3、 Pm4a、 Pm4b、 Pm5、 Pm6、 Pm7、 Pm8等已丧失抗性, 只是在聚合状态下有较好的抗病性^{[ 11]}。在已知的抗白粉病基因中, 来自簇毛麦的 Pm21抗性强, 其载体品种无明显的不良性状, 在不同小麦遗传背景下抗病性均表现稳定, 也是目前最有效的抗病基因之一^{[ 12]}。 Pm22、 Pm25、 Pm26、 Pm27、 Pm28、 Pm29、 Pm31、 Pm32、 Pm33、 Pm34、 Pm35、 Pm36、 Pm37、 Pm38、 Pm39、 Pm40、 Pm41、 Pm42和 Pm43在中国的利用价值有待进一步鉴定^{[ 11]}。本研究发现, 山农20可扩增出 Pm12、 Pm24、 Pm30、 Pm31、 Pm35和 Pm36等6个抗病基因的特异片段, 多个抗病基因的聚合可能是其对多个白粉病混合菌株免疫的重要原因。

从抗性基因在当前生产品种中出现的频率看, Yr9所占比例最大, Yr2和 Yr1次之^{[ 13]}。李在峰等^{[ 14]}发现 Yr5、 Yr10、 Yr15、 Yr26和 Yr31对当前流行的条锈病生理小种具有良好的抗性, 并认为 Yr16、 Yr18和 Yr29为条锈病持久抗性基因。殷学贵等^{[ 15]}在小麦与十倍体长穗偃麦草的杂交后代材料A-3中鉴定出抗条锈病新基因 YrTp1, 对条锈病有较好抗性。本研究检测结果显示, 山农20可能聚合了 Yr5、 Yr9、 Yr15、 Yr24、 Yr26和 YrTp1基因, 田间抗条锈病鉴定结果暗示这些抗病基因的聚合可使品种表现出对条锈病免疫或高抗。

我国小麦品种资源中主要利用的抗叶锈基因有 Lr1、 Lr10、 Lr26、 Lr28、 Lr34和 Lr35, 其中 Lr1、 Lr10和 Lr26因病原菌新致病小种的产生或者次要小种上升为流行小种而“丧失”抗叶锈性^{[ 7]}, 但Kolmer^{[ 16]}和German等^{[ 17]}发现, 将 Lr1、 Lr3、 Lr11、 Lr26等基因与 Lr13或 Lr34聚合后, 可提高小麦成株期的抗病性, 抗病性一般都大于基因单独存在时。 Lr34为慢锈基因, 在成株期表达, 表现持久抗性, 同时兼抗条锈、秆锈和白粉病^{[ 18]}。 Lr9、 Lr19、 Lr24和 Lr38具有全生育期抗病性, 但可能是典型的垂直抗病性基因, 应慎重应用^{[ 19]}。本研究检测结果显示, 山农20可能携带 Lr21和 Lr26。在国家黄淮南片区试鉴定中, 山农20连续两年表现慢叶锈病, 在黄淮北片区试鉴定中表现高抗和中抗叶锈病; 在多年多点的生产种植鉴定中, 抗病反应为免疫和高抗。这是抗性基因聚合的效应还是存在新的抗(慢)叶锈基因, 有待深入研究。

关于小麦纹枯病抗性基因定位和分子标记的报道较少。张小村等^{[ 8]}利用118个RIL-SES系, 发现1个与纹枯病抗性连锁的SSR标记, 即Xgwm526。该标记在山农20及其双亲中有相同的扩增带型, 结合田间纹枯病抗病性鉴定的结果, 推测山农20可能含有 Ses1基因。

小麦的赤霉病抗性是数量性状, 已在3B、4B和5A染色体上定位出抗赤霉病的主效QTL^{[ 20, 21]}。本研究利用这些QTL的双侧标记对山农20及其2个亲本进行检测, 均未扩增出相应标记的特异片段。田间的抗病性鉴定显示山农20对赤霉病表现中感, 个别环境下为高感。因此, 在对山农20进行抗病性遗传改良时, 应着重考虑改善赤霉病抗性。

结合分子标记检测已知抗病基因/QTL和田间鉴定结果, 我们初步推测了山农20可能携带的抗病基因, 为解析该品种的优异抗性的遗传基础提供了初步资料。鉴于本试验田间抗病鉴定采用混合接种方法, 并且分子检测没有用原始载体品种作对照, 因而尚不能完全肯定候选基因的存在, 并且抗性鉴定结果是否完全是基因聚合的效应, 需作进一步验证或设计新的试验进行深入剖析。

4 结论

初步推测山农20可能聚合了 Pm12、 Pm24、 Pm30、 Pm31、 Pm35和 Pm36共6个抗白粉病基因, Yr5、 Yr9、 Yr15、 Yr24、 Yr26和 YrTp1共6个抗条锈病基因, Lr21和 Lr26两个抗叶锈病基因, 以及抗纹枯病基因 Ses1; 未检测到抗赤霉病的主效QTL。该结果从某种程度上部分解释了山农20的优良抗病性。

附表用于抗病性基因检测的分子标记 Supplementary Table Molecular markers used for the marker-assisted selection of disease resistance genes

基因/QTL Gene/QTL	标记类型 Marker type	引物 Primer	距离 Distance (cM)	引物序列 Primer sequence (5'-3')	退火温度 Annealing temp. (℃)	片段大小 Fragment size (bp)	参考文献 Reference
抗白粉病 Powdery mildew resistance
Pm2	STS	Whs350-F	—	AGCTGTTTGGGTACAAGGTG	60	498	[22]
Pm2	STS	Whs350-R	—	GCCATCGTTTTCTACTAG	60	498	[22]
Pm8	STS	IAG95-F	—	AGCAACCAAACACACCCATC	60	1050	[23]
Pm8	STS	IAG95-F	—	ATACTACGAACACACACCCC	60	1050	[23]
Pm12	SSR	Xcau108-F	co-segregating	CTGATCGTCCGTCTCTGTCC	62		[24]
Pm12	SSR	Xcau108-R	co-segregating	GGGTGAAACAGAATGTGGCG	62		[24]
Pm13	STS	Xutv14-F	—	CGCCAGCCAATTATCTCCATGA	62	564	[25]
Pm13	STS	Xutv14-R	—	AGCCATGCGCGGTGTCATGTGAA	62	564	[25]
Pm16	SSR	Xbarc004-F	2.2	GCGTGTTTGTGTCTGCGTTCTA	52	158	[26]
Pm16	SSR	Xbarc004-R	2.2	CACCACACATGCCACCTTCTTT	52	158	[26]
Pm21	SCAR	Pm21D	—	GTTTGTTCACGTTGAATGAATTC	55	1265	[27]
Pm21	SCAR	Pm21E	—	CACTCTCCTCCACTAACAGAGG	55	1265	[27]
Pm24	SSR	Xgwm337-F	2.4±1.2	CCTCTTCCTCCCTCACTTAGC	55	204	[28]
Pm24	SSR	Xgwm337-R	2.4±1.2	TGCTAACTGGCCTTTGCC	55	204	[28]
Pm30	SSR	Xgwm159-F	5.7	GCAGAAGCTTGTTGGTAGGC	60	187	[2]
Pm30	SSR	Xgwm159-R	5.7	GGGCCAACACTGGAACAC	60	187	[2]
Pm31	SSR	Xpsp3071-F	2.5	CGTGCCCTACACCTCCTTTTCTCTC	60	153	[3]
Pm31	SSR	Xpsp3071-R	2.5	TCCGTACATACTCCGGGAGACC	60	153	[3]
Pm32	SSR	Xgwm408-F	1.3	GTATAATTCGTTCACAGCACGC	56	187	[29]
Pm32	SSR	Xgwm408-R	1.3	TCGATTTATTTGGGCCACTG	56	187	[29]
Pm33	SSR	Xwmc317-F	1.1	TGCTAGCAATGCTCCGGGTAAC	60	132	[30]
Pm33	SSR	Xwmc317-R	1.1	TCACGAAACCTTTTCCTCCTCC	60	132	[30]
Pm34	SSR	Xbarc144-F	2.6	GCGTTTTAGGTGGACGACATAGATAGA	62	235	[31]
Pm34	SSR	Xbarc144-R	2.6	GCGCCACGGGCATTTCTCATAC	62	235	[31]
Pm35	SSR	Xcfd7-F	10.3	AGCTACCAGCCTAGCAGCAG	60	251	[32]
Pm35	SSR	Xcfd7-R	10.3	TCAGACACGTCTCCTGACAAA	60	251	[32]
Pm36	SSR	Xwmc75-F	10.0	GTCCGCCGCACACATCTTACTA	61	206	[33]
		Xwmc75-R		GTTTGATCCTGCGACTCCCTTG
Pm43	SSR	Xbarc11-F	4.2	GCGATGCGTGTAAAGTCTGAAGATGA	62	299	[34]
		Xbarc11-R		GCGTCCATGGAGCTCTGTTTTATCTGA

(续附表)

基因/QTL Gene/QTL	标记类型 Marker type	引物 Primer	距离 Distance (cM)	引物序列 Primer sequence (5'-3')	退火温度 Annealing temp. (℃)	片段大小 Fragment size (bp)	参考文献 Reference
抗条锈病 Stripe rust resistance
Yr2	SSR	Xwmc364-F	5.6	ATCACAATGCTGGCCCTAAAAC	58	207	[35]
		Xwmc364-R		CAGTGCCAAAATGTCGAAAGTC
Yr5	SSR	Xgwm501-F	10.5	GGCTATCTCTGGCGCTAAAA	57	166	[36]
		Xgwm501-R		TCCACAAACAAGTAGCGCC
Yr9	SSR	Xgwm582-F	3.7	AAGCACTACGAAAATATGAC	51	135	[37]
		Xgwm582-R		TCTTAAGGGGTGTTATCATA
Yr10	SCAR	SC-200-F	—	CTGCAGAGTGACATCATACA	60	200	[38]
		SC-200-R		TCGAACTAGTAGATGCTGGC
Yr15	SSR	Xgwm33-F	5.0	GGAGTCACACTTGTTTGTGCA	60	123	[39]
		Xgwm33-R		CACTGCACACCTAACTACCTGC
Yr17	SCAR	SC-372-F	0.8±0.7	ATGTCCGCCCTTCCACAACTC	60	372	[40]
		SC-372-R		CACTTGCCTATAAGCACAGAG
Yr18	STS	csLV34-F	—	GTTGGTTAAGACTGGTGATGG	57	150	[41]
		csLV34-R		TGCTTGCTATTGCTGAATAGT
Yr24	SSR	Xgwm273-F	6.1	AGCAGTGAGGAAGGGGATC	58	168	[42]
		Xgwm273-R		ATTGGACGGACAGATGCTTT
Yr26	SSR	Xgwm18-F	1.9	TGGCGCCATGATTGCATTATCTTC	59	186	[43]
		Xgwm18-R		GGTTGCTGAAGAACCTTATTTAGG
Yr36	SSR	Xbarc101-F	co-segregating	GCTCCTCTCACGATCACGCAAAG	62	125	[44]
		Xbarc101-R		GCGAGTCGATCACACTATGAGCCAATG
YrV23	SSR	Xwmc356-F	9.4	GCCGTTGCCCAATGTAGAAG	60	251	[45]
		Xwmc356-R		CCAGAGAAACTCGCCGTGTC
YrTp1	SSR	Xwmc477-F	0.4	CGTCGAAAACCGTACACTCTCC	61	167	[46]
		Xwmc477-R		GCGAAACAGAATAGCCCTGATG
抗叶锈病 Leaf rust resistance
Lr1	STS	WR003-F	—	GGGACAGAGACCTTGGTGGA	55	760	[47]
		WR003-R		GACGATGATGATTTGCTGCTGG
Lr9	STS	J13/1	—	TCCTTTTATTCCGCACGCCGG	66	1100	[48]
		J13/2		CCACACTACCCCAAAGAGACG
Lr10	STS	Lrk10D1	—	GAAGCCCTTCGTCTCATCTG	58	282	[49]
		Lrk10D2		TTGATTCATTGCAGATGAGATCACG
Lr19	STS	Lr19Gb-F	—	CATCCTTGGGGACCTC	56	130	[50]
		Lr19Gb-R		CCAGCTCGCATACATCCA
Lr20	STS	STS638-F	—	ACAGCGATGAAGCAATGAAA	60	542	[51]
		STS638-R		GTCCAGTTGGTTGATGGAAT
Lr21	STS	Lr21-F	—	CGCTTTTACCGAGATTGGTC	57	669	[52]
		Lr21-R		TCTGGTATCTCACGAAGCCTT
Lr24	STS	J09/1	—	TCTAGTCTGTACATGGGGGC	57	350	[53]
		J09/2		TGGCACATGAACTCCATACG
Lr26	STS	Lr26-F	—	GCAAGTAAGCAGCTTGATTTAGC	60	210	[54]
		Lr26-R		AATGGATGTCCCGGTGAGTGG
Lr29	SCAR	OPY10/1	—	GTGACCTCAGGCAATGCA	60	850	[55]
		OPY10/2		GTGACCTCAGAACCGATG
Lr34	STS	csLV34-F	—	GTTGGTTAAGACTGGTGATGG	57	150	[41]
		csLV34-R		TGCTTGCTATTGCTGAATAGT
Lr35	SCAR	Sr39 F2	—	AGAGAGAGTAGAAGAGCTGC	57	900	[55]
		Sr39 R3		AGAGAGAGAGCATCCACC
Lr38	SCAR	SCAR₉₈₂-F	—	GCTGAATCTGCGTATCGTCCC	68	982	[56]
		SCAR₉₈₂-R		GACTTGTTCTTCGGCGTGTTG

(续附表)

基因/QTL Gene/QTL	标记类型 Marker type	引物 Primer	距离 Distance (cM)	引物序列 Primer sequence (5'-3')	退火温度 Annealing temp. (℃)	片段大小 Fragment size (bp)	参考文献 Reference
抗纹枯病Sheath blight resistance
Ses1	SSR	Xgwm526-F	27.9	CAATAGTTCTGTGAGAGCTGCG	58	140	[8]
		Xgwm526-R		CCAACCCAAATACACATTCTCA
抗赤霉病 Scab resistance
Qfhs.ndsu -3BS	SSR	Xgwm533-F	4	AAGGCGAATCAAACGGAATA	60	316	[20]
Qfhs.ndsu -3BS		Xgwm533-R		GTTGCTTTAGGGGAAAAGCC
	SSR	Xgwm493-F	4	TTCCCATAACTAAAACCGCG	60	175
		Xgwm493-R		GGAACATCATTTCTGGACTTTG
Qfhs.ifa -5A	SSR	Xgwm293-F	2	TACTGGTTCACATTGGTGCG	55	205	[20]
Qfhs.ifa -5A		Xgwm293-R		TCGCCATCACTCGTTCAAG
	SSR	Xgwm304-F	2	AGGAAACAGAAATATCGCGG	55	202
		Xgwm304-R		AGGACTGTGGGGAATGAATG
Qfhi.nau -4B	SSR	Xgwm149-F	1	CATTGTTTTCTGCCTCTAGCC	55	152	[21]
Qfhi.nau -4B		Xgwm149-R		CTAGCATCGAACCTGAACAAG
Qfhi.nau -5A	SSR	Xwmc96-F	1	TAGCAGCCATGCTTAGCATCAA	61	280	[21]
Qfhi.nau -5A		Xwmc96-R		GTTTCAGTCTTTCACGAACACG

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

参考文献

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[34]	He R L, Chang Z J, Yang Z J, Yuan Z Y, Zhan H X, Zhang X J, Liu J X. Inheritance and mapping of powdery mildew resistance gene Pm43 introgressed from Thinopyrum intermedium into wheat. Theor Appl Genet, 2009, 118: 1173-1180 [本文引用:1] [JCR: 3.658]
[35]	刘燕, 张增艳, 辛志勇, 林志珊, 杜丽璞, 徐惠君, 刘丽, 陈孝, 张改生. 利用分子标记解析小麦新种质YW243的抗锈病基因. 中国农业科学, 2006, 39: 295-299 Liu Y, Zhang Z Y, Xin Z Y, Lin Z S, Du L P, Xu H J, Liu L, Chen X, Zhang G S. Analysis on the rust resistance genes of a new wheat germplasm YW243 by molecular markers. Sci Agric Sin, 2006, 39: 295-299 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.889]
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[37]	翁东旭, 徐世昌, 万安民, 李景鹏, 吴立仁. 小麦条锈菌鉴别寄主抗条锈病基因Yr9的微卫星标记. 遗传学报, 2005, 32: 937-941 Weng D X, Xu S C, Wan A M, Li J P, Wu L R. Microsatellite marker linked with stripe rust resistant gene Yr9 in wheat. Acta Genet Sin, 2005, 32: 937--941 (in Chinese with English abstract) [本文引用:1]
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[39]	Sun G L, Fahima T, Korol A B, Turpeinen T, Grama A, Ronin Y I, Neve E. Identification of molecular markers linked to the Yr15 stripe rust resistance gene of wheat originated in wild emmer wheat, Triticum dicoccoides. Theor Appl Genet, 1997, 95: 622-628 [本文引用:1] [JCR: 3.658]
[40]	贾举庆, 雷孟平, 刘成, 李光蓉, 杨足君. 小麦抗条锈基因Yr17的新SCAR标记的建立与应用. 麦类作物学报, 2010, 30: 11-16 Jia J Q, Lei M P, Liu C, Li G R, Yang Z J. Exploitation and application of a new SCAR marker linked to strip rust resistance gene Yr17 in wheat. J Triticeae Crops, 2010, 30: 11-16 (in Chinese with English abstract) [本文引用:1]
[41]	Lagudah E S, McFadden H, Singh R P, Huerta-Espino J, Bariana H S, Spielmeyer W. Molecular genetic characterization of the Lr34/Yr18 slow rusting resistance gene region in wheat. Theor Appl Genet, 2006, 114: 21-30 [本文引用:1] [JCR: 3.658]
[42]	刘亚萍, 曹双河, 王献平, 徐志斌, 张相岐. 小麦抗条锈病基因Yr24的SSR标记. 植物病理学报, 2005, 35: 478-480 Liu Y P, Cao S H, Wang X P, Xu Z B, Zhang X Q. Molecular mapping of stripe rust resistance gene Yr24 in wheat. Acta Phytopathol Sin, 2005, 35: 478-480 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.088]
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2013

3.658

0.0

. 2013, 126:1397-1403 DOI:10.1007/s00122-013-2060-6

Identification of the gene Pm47 on chromosome 7BS conferring resistance to powdery mildew in the Chinese wheat landrace Hongyanglazi

<li>1.The National Key Facility for Crop Gene Resources and Genetic Improvement (NFCRI), Institute of Crop Science, Chinese Academy of Agricultural Sciences, Beijing, 100081, China</li><li>2.Forestry College of Henan University of Science and Technology, Luoyang, 471003, China</li><li>3.Center for Agricultural Resources Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Shijiazhuang, 050021, China</li>

Abstract

Powdery mildew, caused by Blumeria graminis f. sp. tritici (Bgt), is an important disease that causes substantial yield losses in wheat (Triticum aestivum) in China and other parts of the world. This foliar disease can be effectively managed by host resistance. The Chinese landrace Hongyanglazi from Shaanxi province is highly resistant to many Bgt isolates at the seedling stage. Genetic analysis using an F_2:3 population derived from a cross between Hongyanglazi and susceptible cultivar Zhongzuo 9504 indicated that Hongyanglazi carried a single recessive gene (tentatively designated PmHYLZ) conferring its resistance to Bgt isolate E09. PmHYLZ was flanked by EST marker BE606897 and microsatellite marker Xgwm46 on chromosome 7BS at genetic distances of 1.7 and 3.6 cM, respectively. This gene differed from Pm40, also located on 7BS, by origin, linked markers, and reactions to 13 Bgt isolates. Based on these findings, PmHYLZ was permanently designated as Pm47.

... 目前, 已有60多个小麦抗白粉病基因被定位到47个位点(Pm1~Pm47), 分布在除3A、4D以外的各条染色体上^[1,2] ...

2011

0.0

... 目前, 已有60多个小麦抗白粉病基因被定位到47个位点(Pm1~Pm47), 分布在除3A、4D以外的各条染色体上^[1,2] ...

2008

3.658

0.0

. 2008, 116:1155-1166 DOI:10.1007/s00122-008-0743-1

The adult plant rust resistance loci Lr34/Yr18 and Lr46/Yr29 are important determinants of partial resistance to powdery mildew in bread wheat line Saar

1.Department of Plant and Environmental Sciences, Norwegian University of Life Sciences, P.O. Box 5003, 1432 ?s, Norway 2.CIMMYT, Apdo. Postal 6-641, 06600 México, D.F. Mexico 3.Campo Experimental Valle de Mexico-INIFAP, Apdo. Postal 10, 56230 Chapingo, Edo. de Mexico, Mexico 4.Institute of Crop Sciences/National Wheat Improvement Center, Chinese Academy of Agricultural Sciences, Zhongguancun South Street 12, 100081 Beijing, People’s Republic of China 5.CIMMYT China Office, c/o Chinese Academy of Agricultural Sciences, Zhongguancun South Street 12, 100081 Beijing, People’s Republic of China

Powdery mildew, caused by Blumeria graminis f. sp. tritici is a major disease of wheat (Triticum aestivum L.) that can be controlled by resistance breeding. The CIMMYT bread wheat line Saar is known for its good level of partial and race non-specific resistance, and the aim of this study was to map QTLs for resistance to powdery mildew in a population of 113 recombinant inbred lines from a cross between Saar and the susceptible line Avocet. The population was tested over 2 years in field trials at two locations in southeastern Norway and once in Beijing, China. SSR markers were screened for association with powdery mildew resistance in a bulked segregant analysis, and linkage maps were created based on selected SSR markers and supplemented with DArT genotyping. The most important QTLs for powdery mildew resistance derived from Saar were located on chromosomes 7DS and 1BL and corresponded to the adult plant rust resistance loci Lr34/Yr18 and Lr46/Yr29. A major QTL was also located on 4BL with resistance contributed by Avocet. Additional QTLs were detected at 3AS and 5AL in the Norwegian testing environments and at 5BS in Beijing. The population was also tested for leaf rust (caused by Puccinia triticina) and stripe rust (caused by P. striiformis f. sp. tritici) resistance and leaf tip necrosis in Mexico. QTLs for these traits were detected on 7DS and 1BL at the same positions as the QTLs for powdery mildew resistance, and confirmed the presence of Lr34/Yr18 and Lr46/Yr29 in Saar. The powdery mildew resistance gene at the Lr34/Yr18 locus has recently been named Pm38. The powdery mildew resistance gene at the Lr46/Yr29 locus is designated as Pm39.

... 其中, Pm38和Pm39为白粉病成株抗性基因^[3], 其余抗性基因均为病原菌生理小种专化性的主效基因 ...

2010

0.0

... 由于生理小种的不断变异, 大部分已丧失抗性^[4] ...

... 还有72个条锈病成株抗性QTL被定位在除1A、1D、3D和7A染色体外的其余各个染色体上^[4] ...

2011

0.0

... 国际上已命名51个抗条锈病主效基因, 涉及48个位点(Yr1~Yr48), 除Yr11~Yr14外均被定位于特定的染色体上^[5] ...

... 其中, Yr3位点有Yr3a、Yr3b和Yr3c三个复等位基因, Yr4位点有Yr4a和Yr4b两个复等位基因, Yr11、Yr12、Yr13、Yr14、Yr16、Yr18、Yr29、Yr30、Yr36、Yr39、Yr46和Yr48为成株抗性基因, 其他39个为苗期抗性基因^[5] ...

2013

0.0

... 目前已正式命名的小麦抗叶锈病基因有68个, 其中大多数为小种特异的主效抗叶锈病基因^[6], 已报道39个分子标记与抗叶锈病基因连锁或共分离^[7] ...

2011

0.0

... 目前已正式命名的小麦抗叶锈病基因有68个, 其中大多数为小种特异的主效抗叶锈病基因^[6], 已报道39个分子标记与抗叶锈病基因连锁或共分离^[7] ...

... 抗叶锈性^[7], 但Kolmer^[16]和German等^[17]发现, 将Lr1、Lr3、Lr11、Lr26等基因与Lr13或Lr34聚合后, 可提高小麦成株期的抗病性, 抗病性一般都大于基因单独存在时 ...

2005

0.0

1.1628

. 2005, 6:276-279

QTL and molecular markers for resistance gene of wheat sharp eyespot

(小麦纹枯病抗性的QTL分析和抗病基因的分子标记) . (植物遗传资源学报) , 2005 , 6 : 276 - 279 Zhang X C , Li S S , Zhao X H , Fan Y D , Li R J . QTL and molecular markers for resistance gene of wheat sharp eyespot

Zhang X C

(张小村) , (李斯深) , (赵新华) , (范玉顶) , (李瑞军) . (小麦纹枯病抗性的QTL分析和抗病基因的分子标记) . (植物遗传资源学报) , 2005 , 6 : 276 - 279 Zhang X C

对RIL-8群体纹枯病抗性进行QTL检测,检测到一个加性QTL,位于1A染色体上,贡献率为21.57%;检测到4对QTL间互作位点,涉及7条染色体,互作贡献率分别为11.63%、6.54%、14.04%、20.01%,互作总贡献率为52.23%.通过对RIL-SES群体118个系检测,发现1个SSR标记Xgwm526,位于2B染色体上,与纹枯病抗病基因距离为27.9cM;一个ISSR标记IS840,与纹枯病抗病基因距离为16.9cM.

... 张小村等^[8]利用2个重组自交系群体RIL-8和RIL-SES, 对抗纹枯病性QTL进行分析, 检测到1个SSR标记(Xgwm526)与纹枯病抗性具有关联性, 与纹枯病抗病基因Ses1的遗传距离为27 ...

... 张小村等^[8]利用118个RIL-SES系, 发现1个与纹枯病抗性连锁的SSR标记, 即Xgwm526 ...

2004

0.0

1.667

. 2004, 30:739-744

Analysis of QTL for Fusarium head blight resistance in wheat

(小麦赤霉病抗性QTL分析) . (作物学报) , 2004 , 30 : 739 - 744 Zhou M P , Ren L J , Zhang X , Olga-E Scholten , Huang Y H , Ma H X , Lu W Z . Analysis of QTL for Fusarium head blight resistance in wheat

Olga-E Scholten

(周淼平) , (任丽娟) , (张旭) , Olga-E Scholten

National Wheat Improvement Center/Institute of Crop Breeding and Cultivation, Chinese Academy of Agricultural Sciences, Beijing 100081

Improving the processing quality of pan bread and dry white Chinese noodle (DWCN) are the major breeding objectives for North China Winter Wheat Region, and understanding the association in wheat quality parameters between pan bread and DWCN will contribute to the efficiency of breeding program. Eig

改良面包和面条品质是北方冬麦区的主要育种目标，了解二者的品质异同有助于提高育种效率。本研究以81份冬小麦品种品系为材料，比较了小麦品质性状与面包和面条品质表现的关系。结果表明，籽粒硬度、蛋白质含量、SDS沉降值、形成时间、稳定时间、拉伸面积和抗延阻力是影响面包和面条品质的共同性状，但它们对面包、面条品质

... 小麦赤霉病主要发生在温暖潮湿和半潮湿地区, 在我国主要发生在长江中下游冬麦区和东北春麦区, 近年来随着全球气候变暖、灌溉条件的改善及耕作制度的变化, 有进一步蔓延到黄淮麦区的趋势^[9] ...

0.0

... 目前, 除7D染色体外的其他所有的染色体上都检测到抗赤霉病QTL的存在^[10], 其中位于3B、5A和6B染色体上的QTL在苏麦3号、望水白、Wuhan、Nyubai、Frontana、CM82036和DH181中被检测到, 并且效应较大, 是小麦抗赤霉病的主效QTL ...

2009

0.0

. 2009, 25(10):53-58

Researches and application on molecular markers of powdery mildew resistant genes in wheat

(小麦白粉病抗病基因分子标记开发及应用研究进展) . (中国农学通报) , 2009 , 25 ( 10 ): 53 - 58 Li C-X , Xu W G . Researches and application on molecular markers of powdery mildew resistant genes in wheat

Li C-X

(李春鑫) , (许为钢) . (小麦白粉病抗病基因分子标记开发及应用研究进展) . (中国农学通报) , 2009 , 25 ( 10 ): 53 - 58 Li C-X

1. Henan academy of agricultural sciences wheat research center 2.

The use of resistant cultivars is an effective, economical, and environmentally safe approach that eliminates the use of fungicides and reduces production losses due to powdery mildew caused by Blumeria graminis f. sp. tritici (syn. Erysiphe graminis f. sp. tritici). So far, 55 genes/alleles for resistance to powdery mildew at 39 loci have been identified and located on 16 different chromosomes,of which 30 resistance genes/alleles have been tagged by different molecular marker systems. Several quantitative trait loci (QTLs) for adult plant resistance to powdery mildew have been associated with molecular markers. The detailed information on chromosomal location,the sources, available molecular markers and cloning of these genes has been reviewed. The fresh researches, existed problem and the resolvent in marker-assisted selection using these genes are also discussed.

选育和利用抗病品种是防治白粉病最经济、有效、安全的措施，优异抗白粉病基因资源的挖掘及其紧密连锁分子标记的开发是开展抗白粉病分子标记育种的基础。目前，国内外已命名了分布于16条染色体上的39个位点共55个白粉病抗病基因，其中30个已开发出分子标记,另外还发现数个与成株期抗性相关的数量性状位点。本文综述了国内外小麦白粉病抗病基因的定位、来源、分子标记开发、克隆，以及白粉病抗病基因的分子标记育种的最新研究进展,并分析了当前小麦抗白粉病育种存在的主要问题及解决建议。

... 目前, Pm12、Pm13、Pm16、Pm20、Pm21、Pm30在中国对小麦白粉病表现高抗至免疫, 而Pm1、Pm2、Pm3、Pm4a、Pm4b、Pm5、Pm6、Pm7、Pm8等已丧失抗性, 只是在聚合状态下有较好的抗病性^[11] ...

... Pm22、Pm25、Pm26、Pm27、Pm28、Pm29、Pm31、Pm32、Pm33、Pm34、Pm35、Pm36、Pm37、Pm38、Pm39、Pm40、Pm41、Pm42和Pm43在中国的利用价值有待进一步鉴定^[11] ...

2004

0.0

. 2004, 24:127-132

Researches on powdery mildew resistant genes and their molecular markers in wheat

(小麦抗白粉病基因及其分子标记研究进展) . (麦类作物学报) , 2004 , 24 : 127 - 132 Qiu Y C , Zhang S S . Researches on powdery mildew resistant genes and their molecular markers in wheat

Qiu Y C

(邱永春) , (张书绅) . (小麦抗白粉病基因及其分子标记研究进展) . (麦类作物学报) , 2004 , 24 : 127 - 132 Qiu Y C

小麦白粉病是小麦生产的主要病害之一,应用抗病品种是防治该病的十分经济、有效的措施.近年来分子标记技术的发展为小麦抗白粉病基因的研究提供了极大的方便.目前为止,小麦基因组中已定名抗白粉病基因31个,其中对20个基因位点的25个抗白粉病基因进行了标记和作图.本文对小麦抗白粉病基因染色体定位、来源、评价利用及其分子标记研究现状进行了综述,并对分子标记在小麦抗病遗传育种中的应用进行了探讨.

... 在已知的抗白粉病基因中, 来自簇毛麦的Pm21抗性强, 其载体品种无明显的不良性状, 在不同小麦遗传背景下抗病性均表现稳定, 也是目前最有效的抗病基因之一^[12] ...

2009

0.0

. 2009, 25(13):190-196

Progress on stripe rust resistance genes and resistant breeding in wheat

(小麦条锈病抗性基因研究进展及分子标记辅助聚合育种) . (中国农学通报) , 2009 , 25 ( 13 ): 190 - 196 Dong S J , Xu W G . Progress on stripe rust resistance genes and resistant breeding in wheat

Dong S J

(董淑静) , (许为钢) . (小麦条锈病抗性基因研究进展及分子标记辅助聚合育种) . (中国农学通报) , 2009 , 25 ( 13 ): 190 - 196 Dong S J

The origin, chromosomal location and molecular markers of stripe rust resistance genes in wheat are summarized .The evolvement of resistance varieties and physiological race are reviewed.The stripe rust resistance genes in wheat are valued.And the applications of molecular markers in wheat of resistance breeding are discussed.

概述了已知小麦条锈病抗性基因的来源、染色体定位和分子标记研究现状，回顾了抗性品种与生理小种的演变，评价了小麦抗条锈基因，并浅谈了小麦抗条锈基因的分子标记在育种中的应用

... 从抗性基因在当前生产品种中出现的频率看, Yr9所占比例最大, Yr2和Yr1次之^[13] ...

2006

3.658

0.0

. 2006, 112:1098-1103 DOI:10.1007/s00122-006-0211-8

Molecular tagging of stripe rust resistance gene YrZH84 in Chinese wheat line Zhou 8425B

1.Institute of Crop Science/National Wheat Improvement Center, Chinese Academy of Agricultural Sciences (CAAS), Zhongguancun South Street 12, 100081 Beijing, China 2.Zhoukou Academy of Agricultural Sciences, Jianshe East Road, 466001 Zhoukou, Henan Province China 3.International Maize and Wheat Improvement Center (CIMMYT), CIMMYT China Office, c/o CAAS, Zhongguancun South Street 12, 100081 Beijing, China 4.Institute of Plant Protection, Chinese Academy of Agricultural Sciences (CAAS), Yuanmingyuan West Road 2, 100094 Beijing, China 5.International Maize and Wheat Improvement Center (CIMMYT), Apdo. Postal 6-641, 06600 Mexico D.F, Mexico

Stripe rust, caused by Puccinia striiformis f. sp. tritici (PST), is one of the most damaging diseases in common wheat (Triticum aestivum L.). With the objective of identifying and tagging new genes for resistance to stripe rust, F₁, F₂and F₃ populations from the cross Zhou 8425B/Chinese Spring were inoculated with Chinese PST isolate CYR32 in the greenhouse. A total of 790 SSR primers were used to test the parents and resistant and susceptible bulks. The resulting seven polymorphic markers on chromosome 7BL were used for genotyping F₂and F₃ populations. Results indicated that Zhou 8425B carries a single dominant resistance gene, temporarily designated YrZH84, closely linked to SSR markers Xcfa2040-7B and Xbarc32-7B with genetic distances of 1.4 and 4.8 cM, respectively. In a seedling test with 25 PST isolates, the reaction patterns of YrZH84 were different from those of lines carrying Yr2 and Yr6. It was concluded that YrZH84 is probably a new stripe rust resistance gene.

... 李在峰等^[14]发现Yr5、Yr10、Yr15、Yr26和Yr31对当前流行的条锈病生理小种具有良好的抗性, 并认为Yr16、Yr18和Yr29为条锈病持久抗性基因 ...

2005

0.0

... 殷学贵等^[15]在小麦与十倍体长穗偃麦草的杂交后代材料A-3中鉴定出抗条锈病新基因YrTp1, 对条锈病有较好抗性 ...

1992

1.643

0.0

1992

3.658

0.0

1996

10.229

0.0

... Lr34为慢锈基因, 在成株期表达, 表现持久抗性, 同时兼抗条锈、秆锈和白粉病^[18] ...

2011

0.0

... Lr9、Lr19、Lr24和Lr38具有全生育期抗病性, 但可能是典型的垂直抗病性基因, 应慎重应用^[19] ...

2002

3.658

0.0

. 2002, 104:84-91 DOI:10.1007/s001220200009

Ruckenbauer P. Molecular mapping of QTLs for Fusarium head blight resistance in spring wheat: I. Resistance to fungal spread (Type II resistance)

1.IFA-Tulln, Institute for Agrobiotechnology, Department of Biotechnology in Plant Produktion, Konrad Lorenz Strasse 20, A-3430 Tulln, Austria e-mail: buerst@ifa-tulln.ac.at Fax: +43-2272-66280-77 AT 2.Bayerische Landesanstalt für Bodenkultur und Pflanzenbau, V?ttingerstrasse 38, D-85354 Freising, Germany DE

Fusarium head blight (FHB, scab) is a fungal disease of wheat and other small cereals that is found in both temperate and semi-tropical regions. FHB causes severe yield and quality losses, but the most-serious concern is the possible mycotoxin contamination of cereal food and feed. Breeding for FHB resistance by conventional selection is feasible, but tedious and expensive. This study was conducted to identify and map DNA markers associated with FHB resistance genes in wheat. A population of 364 F₁-derived doubled-haploid (DH) lines from the cross ’CM-82036’ (resistant)/’Remus’ (susceptible) was evaluated for Type II resistance (spread within the spike) during 2 years under field conditions. Marker analysis was performed on 239 randomly chosen DH lines. Different marker types were applied, with an emphasis on AFLP and SSR markers. Analysis of variance, as well as simple and composite interval mapping, were applied. Three genomic regions were found significantly associated with FHB resistance. The most-prominent effect was detected on the short arm of chromosome 3B, explaining up to 60% of the phenotypic variance for Type II FHB resistance. A further QTL was located on chromosome 5A and a third one on 1B. The QTL regions on 3B and 5A were tagged with flanking SSR markers, the 1B QTL was found associated with the high-molecular-weight glutenin locus. These results indicate that FHB resistance is under control of a few major QTLs operating together with unknown numbers of minor genes. Marker-assisted selection for these major QTLs involved in FHB resistance appears feasible and should accelerate the development of resistant and agronomically improved wheat cultivars.

... 小麦的赤霉病抗性是数量性状, 已在3B、4B和5A染色体上定位出抗赤霉病的主效QTL ^[20,21] ...

2008

0.0

1.667

. 2008, 34:539-544 DOI:10.3724/SP.J.1006.2008.00539

Mapping QTLs associated with resistance to Fusarium head blight using an #cod#x0201c;immortalized F₂#cod#x0201d; population

(利用#cod#x0201c;永久F 2) #cod#x0201d;群体定位抗赤霉病QTL

Tian D G

(田大刚) , (林峰) , (张彩琴) , (张政值) , (薛树林) , (曹勇) , (李春军) , (马正强) . (利用#cod#x0201c;永久F 2) #cod#x0201d;群体定位抗赤霉病QTL . (作物学报) , 2008 , 34 : 539 - 544 Tian D G

Laboratory of Applied Plant Genomics and the National Key Laboratory of Crop Genetics and Germplasm Enhancement, Nanjing Agricultural University, Nanjing 210095, Jiangsu, China

Fusarium head blight (FHB) or scab disease, caused by Gibberrella zeae (Schw.) Petch, is a serious disease in many wheat (Triticum aestivum L.) growing regions worldwide. To study the inheritance of scab resistance against fungal penetration (type I resistance), an 198 lines “immortalized F₂”population was constructed by crossing between recombinant inbred lines chosen by random permutation of 132 RILs from the RIL population derived from the cross between scab-susceptible cultivar Nanda 2419 and scab-resistant cultivar Wangshuibai. The population were then evaluated for percentage of infected spikes (PIS) across two years and six chromosome regions were detected for their association with type I resistance through interval mapping, among which Qfhi.nau-4B and Qfhi.nau-5A with the resistance alleles originated from Wangshuibai and Qfhi.nau-2B with the resistance allele from Nanda 2419 were consistently detected. Qfhi.nau-4B and Qfhi.nau-5A had the largest effects among the detected QTLs and both showed mainly the additive allelic effects, the former also had partial dominance. In addition, four pairs of significant interaction loci were identified. These results further demonstrate that wheat scab resistance are under complex genetic control and also imply that early generation selection for type I resistance in scab resistance breeding is feasible with Wangshuibai as the parent.

为了研究抗赤霉病侵染性的遗传, 利用感赤霉病品种南大2419和抗赤霉病品种望水白杂交单粒传获得的重组自交系群体132个株系间的随机配对组合, 构建了一个包含198个株系的“永久F2”群体。通过两年抗侵染田间试验和QTL作图, 定位了6个抗侵染QTL, 其中抗性等位位点源于望水白的Qfhi.nau-4B和Qfhi.nau-5A以及源于南大2419的Qfhi.nau-2B的效应较为稳定。Qfhi.nau-4B和Qfhi.nau-5A的效应较大且以加性效应为主, 前者存在部分显性基因效应。此外, 还检测到4对显著的互作位点。这些结果进一步说明赤霉病抗性遗传的复杂性, 同时也表明在利用望水白进行抗赤霉病育种时早代选择抗侵染性是可行的。

... 小麦的赤霉病抗性是数量性状, 已在3B、4B和5A染色体上定位出抗赤霉病的主效QTL ^[20,21] ...

1996

3.658

0.0

2001

1.175

0.0

2004

0.0

1999

3.658

0.0

2005

1.175

0.0

1999

1.175

0.0

2000

3.658

0.0

. 2000, 101:407-414 DOI:10.1007/s001220051497

Molecular mapping of the wheat powdery mildew resistance gene Pm24 and marker validation for molecular breeding

1.Technische Universit?t München, Lehrstuhl für Pflanzenbau und Pflanzenzüchtung, Alte Akademie 12, D-85350 Freising-Weihenstephan, Germany e-mail: mohler@weihenstephan.de Fax: +49 8161 715173 Present address: X.Q. Huang, Institut für Pflanzengenetik und Kulturpflanzenforschung, Corrensstr. 3, D-06466 Gatersleben, Germany DE DE

Molecular markers were identified in common wheat for the Pm24 locus conferring resistance to different isolates of the powdery mildew pathogen, Erysiphe graminis DM f. sp. tritici (Em. Marchal). Bulked segregant analysis was used to identify amplified fragment length polymorphism (AFLP) markers and microsatellite markers linked to the gene Pm24 in an F₂ progeny from the cross Chinese Spring (susceptible)× Chiyacao (resistant). Two AFLP markers XACA/CTA-407 and XACA/CCG-420, and three microsatellite markers Xgwm106, Xgwm337 and Xgwm458, were mapped in coupling phase to the Pm24 locus. The AFLP marker locus XACA/CTA-407 co-segregated with the Pm24 gene, and XACA/CCG-420 mapped 4.5 cM from this gene. Another AFLP marker locus XAAT/CCA-346 co- segregated in repulsion phase with the Pm24 locus. Pm24 was mapped close to the centromere on the short arm of chromosome 1D, contrary to the previously reported location on chromosome 6D. Pm24 segregated independently of gene Pm22, also located on chromosome 1D. An allele of microsatellite locus Xgwm337 located 2.4±1.2 cM from Pm24 was shown to be diagnostic and therefore potentially useful for pyramiding two or more genes for powdery mildew resistance in a single genotype.

2003

1.643

0.0

. 2003, 133:367-370 DOI:10.1023/A:1025738513638

Chromosomal location of genes for resistance to powdery mildew in common wheat (Triticum aestivum L. em Thell.): 8. Gene Pm32 in a wheat- Aegilops speltoides translocation line

1.Technical University of Munich Division of Plant Breeding and Applied Genetics D–85350 Freising-Weihenstephan Germany D–85350 Freising-Weihenstephan Germany 2.Research Institute of Agriculture in Central Regions of Non-chernozem Zone Nemchinovka 1 Moscow oblast 143013 Russia Nemchinovka 1 Moscow oblast 143013 Russia

The wheat-Aegilops speltoides translocation line L501 exhibits a disease response pattern distinctive from that of documented powdery mildew genes after inoculation with differential Blumeria graminis tritici isolates. Results based on cytological C-bandings and monosomic analyses reveal that a dominant resistance gene derived from Ae. speltoides is located on a T1BL·1SS chromosome translocation in this line. The new gene is designated Pm32.

2005

1.668

0.0

2006

3.658

0.0

. 2006, 113:1497-1504 DOI:10.1007/s00122-006-0397-9

Pm34: a new powdery mildew resistance gene transferred from Aegilops tauschii Coss. to common wheat (Triticum aestivum L.)

1.Department of Crop Science, North Carolina State University, Box 7629, Raleigh, NC 27695, USA 2.United States Department of Agriculture—Agricultural Research Service, North Carolina State University, Box 7619, Raleigh, NC 27695, USA 3.Department of Plant Pathology, North Carolina State University, Box 7619, Raleigh, NC 27695, USA

Powdery mildew is a major fungal disease in wheat growing areas worldwide. A novel source of resistance to wheat powdery mildew present in the germplasm line NC97BGTD7 was genetically characterized as a monogenic trait in greenhouse and field trials using F₂ derived lines from a NC97BGTD7 X Saluda cross. Microsatellite markers were used to map and tag this resistance gene, now designated Pm34. Three co-dominant microsatellite markers linked to Pm34 were identified and their most likely order was established as: Xbarc177-5D, 5.4cM, Pm34, 2.6cM, Xbarc144-5D, 14cM, Xgwm272-5D. These microsatellite markers were previously mapped to the long arm of the 5D chromosome and their positions were confirmed using Chinese Spring nullitetrasomic Nulli5D-tetra5A and ditelosomic Dt5DL lines. Pm2, the only other known Pm gene on chromosome 5D, has been mapped to the short arm and its specificity is different from that of Pm34.

2007

3.658

0.0

. 2007, 114:1451-1456 DOI:10.1007/s00122-007-0530-4

Pm35: a novel Aegilops tauschii derived powdery mildew resistance gene introgressed into common wheat (Triticum aestivum L.)

1.Department of Crop Science, North Carolina State University, Box 7629, Raleigh, NC 27695, USA 2.Department of Crop and Soil Sciences, University of Georgia, 1109 Experiment street Griffin, Georgia, 30223 USA 3.United States Department of Agriculture, Agricultural Research Service, Department of Plant Pathology, North Carolina State University, Raleigh, NC 27695, USA 4.Department of Plant Pathology, North Carolina State University, Box 7619, Raleigh, NC 27695, USA

A single gene controlling powdery mildew resistance was identified in the North Carolina germplasm line NC96BGTD3 (NCD3) using genetic analysis of F₂ derived lines from a NCD3 X Saluda cross. Microsatellite markers linked to this Pm gene were identified and their most likely order was Xcfd7, 10.3 cM, Xgdm43, 8.6 cM, Xcfd26, 11.9 cM, Pm gene. These markers and the Pm gene were assigned to chromosome 5DL by means of Chinese Spring Nullitetrasomic (Nulli5D-tetra5A) and ditelosomic (Dt5DL) lines. A detached leaf test showed a distinctive disease reaction to six pathogen isolates among the NCD3 Pm gene, Pm2 (5DS) and Pm34 (5DL). An allelism test showed independence between Pm34 and the NCD3 Pm gene. Together, the tests provided strong evidence for the presence of a novel Pm gene in NCD3, and this gene was designated Pm35.

2008

3.658

0.0

. 2008, 117:135-142 DOI:10.1007/s00122-008-0760-0

Molecular mapping of the novel powdery mildew resistance gene Pm36 introgressed from Triticum turgidum var. dicoccoides in durum wheat

1.University of Bari Department of Agro-Forestry and Environmental Biology and Chemistry via Amendola, 165/A 70126 Bari Italy

Powdery mildew, caused by Blumeria graminis f.sp. tritici, is one of the most important wheat diseases in many regions of the world. Triticum turgidum var. dicoccoides (2n = 4x = AABB), the progenitor of cultivated wheats, shows particular promises as a donor of useful genetic variation for several traits, including disease resistances. The wild emmer accession MG29896, resistant to powdery mildew, was backcrossed to the susceptible durum wheat cultivar Latino, and a set of backcross inbred lines (BC₅F₅) was produced. Genetic analysis of F₃ populations from two resistant introgression lines (5BIL-29 × Latino and 5BIL-42 × Latino) indicated that the powdery mildew resistance is controlled by a single dominant gene. Molecular markers and the bulked segregant analysis were used to characterize and map the powdery mildew resistance. Five AFLP markers (XP43M32 ₍₂₅₀₎, XP46M31 ₍₄₁₀₎, XP41M37 ₍₁₀₀₎, XP41M39 ₍₂₅₀₎, XP39M32 ₍₁₂₀₎), three genomic SSR markers (Xcfd07, Xwmc75, Xgwm408) and one EST-derived SSR marker (BJ261635) were found to be linked to the resistance gene in 5BIL-29 and only the BJ261635 marker in 5BIL-42. By means of Chinese Spring nullisomic–tetrasomic, ditelosomic and deletion lines, the polymorphic markers and the resistance gene were assigned to chromosome bin 5BL6-0.29-0.76. These results indicated that the two lines had the same resistance gene and that the introgressed dicoccoides chromosome segment was longer (35.5 cM) in 5BIL-29 than that introgressed in 5BIL-42 (less than 1.5 cM). As no powdery mildew resistance gene has been reported on chromosome arm 5BL, the novel resistance gene derived from var. dicoccoides was designated Pm36. The 244 bp allele of BJ261635 in 5BIL-42 can be used for marker-assisted selection during the wheat resistance breeding process for facilitating gene pyramiding.

2009

3.658

0.0

. 2009, 118:1173-1180 DOI:10.1007/s00122-009-0971-z

Inheritance and mapping of powdery mildew resistance gene Pm43 introgressed from Thinopyrum intermedium into wheat

1.Shanxi University School of Life Science and Technology 030006 Taiyuan Shanxi China 2.Shanxi Academy of Agricultural Sciences Institute of Crop Genetics 030031 Taiyuan Shanxi China 3.University of Electronic Science and Technology of China School of Life Science and Technology 610054 Chengdu Sichuan China 4.Shanxi Academy of Agricultural Sciences Institute of Plant Protection 030031 Taiyuan Shanxi China

Powdery mildew resistance from Thinopyrum intermedium was introgressed into common wheat (Triticum aestivum L.). Genetic analysis of the F₁, F₂, F₃ and BC₁ populations from powdery mildew resistant line CH5025 revealed that resistance was controlled by a single dominant allele. The gene responsible for powdery mildew resistance was mapped by the linkage analysis of a segregating F₂ population. The resistance gene was linked to five co-dominant genomic SSR markers (Xcfd233, Xwmc41, Xbarc11, Xgwm539 and Xwmc175) and their most likely order was Xcfd233–Xwmc41–Pm43–Xbarc11–Xgwm539–Xwmc175 at 2.6, 2.3, 4.2, 3.5 and 7.0 cM, respectively. Using the Chinese Spring nullisomic-tetrasomic and ditelosomic lines, the polymorphic markers and the resistance gene were assigned to chromosome 2DL. As no powdery mildew resistance gene was previously assigned to chromosome 2DL, this new resistance gene was designated Pm43. Pm43, together with the identified closely linked markers, could be useful in marker-assisted selection for pyramiding powdery mildew resistance genes.

2006

0.0

1.889

. 2006, 39:295-299

Analysis on the rust resistance genes of a new wheat germplasm YW243 by molecular markers

(利用分子标记解析小麦新种质YW243的抗锈病基因) . (中国农业科学) , 2006 , 39 : 295 - 299 Liu Y , Zhang Z Y , Xin Z Y , Lin Z S , Du L P , Xu H J , Liu L , Chen X , Zhang G S . Analysis on the rust resistance genes of a new wheat germplasm YW243 by molecular markers

Liu Y

(刘燕) , (张增艳) , (辛志勇) , (林志珊) , (杜丽璞) , (徐惠君) , (刘丽) , (陈孝) , (张改生) . (利用分子标记解析小麦新种质YW243的抗锈病基因) . (中国农业科学) , 2006 , 39 : 295 - 299 Liu Y

【Objective】 A new wheat line YW243 is a recently developed new germplasm with multi-resistance to wheat stripe rust, stem rust, leaf rust, wheat yellow dwarf and powdery mildew. In this study, the gene composition of the stripe and stem rust resistance of YW243 was analyzed. In order to clarify the sources of the rust resistance genes of YW243, YW243 and its parents, such as Fengkang 8, Fengkang 13, Shan 7859, Suwon 11 and Heine Hvede, were analyzed by the molecular markers of the resistance genes . 【Method】 The composition and the sources of the stripe and stem rust resistance genes of YW243 were analyzed by using PCR markers of Yr9, YrX, Yr2, Yr1, Sr31 and SDS-PAGE. 【Result】 YW243 possessed at least four stripe rust resistance genes of Yr1, Yr2, Yr9 and YrX, one stem rust resistance gene Sr31. Shan 7859 and Fengkang13 are the sources of Yr1 in YW243, Shan7859, Fengkang 8 and Fengkang13 are the sources of Yr2 in YW243, Fengkang 13 is the source of YrX of YW243. Fengkang 8 and Shan 7859 are the sources of Yr9 and Sr31 of YW243. 【Conclusion】YW243 is a good resistance germplasm for wheat breeding in the future.

【目的】对兼抗条锈病、秆锈病、叶锈病、黄矮病、白粉病等5种病害的小麦新种质YW243中抗锈病基因的组成和来源进行解析。【方法】通过系谱推断，利用了抗条锈病基因Yr9、YrX、Yr2、Yr1以及抗秆锈病基因Sr31的特异PCR标记和醇溶蛋白蛋白质电泳图谱，分析YW243及其部分相关亲本材料丰抗8号、丰抗13、陕7859中相关的抗条锈病基因和抗秆锈病基因的有无，解析YW243中抗锈病基因的组成。【结果】YW243含有抗条锈病基因Yr1、Yr2、Yr9、YrX，抗秆锈病基因Sr31、抗叶锈病基因Lr26。Yr1源于亲本陕7859或丰抗13，Yr2源于陕7859或丰抗8号或丰抗13；YrX源于丰抗13；Yr9 、Sr31和Lr26源于陕7859或丰抗8号中。【结论】解析了YW243中抗条锈病基因和抗秆锈病基因的组成，证明YW243是一个含有Yr1、Yr2、Yr9、YrX、Sr31等抗锈病基因的多基因聚合体，这些基因特异的SSR，STS特异标记可用于检测多基因聚合体YW243中目标基因，为在抗锈育种中利用、选择YW243的抗锈基因提供了快捷方法。

2002

1.175

0.0

2005

0.0

2001

0.0

0.95

. 2001, 46:669-672

Identification of an AFLP marker linked to the stripe rust resistance gene Yrl0 in wheat

以抗条锈病基因Yr10的供体亲本Moro作参照,用6个PstⅠ端引物和10个TaqⅠ端引物对抗条锈病基因Yr10的近等基因系和轮回亲本Avocet S进行了AFLP分析,共扩增出约4200条可分辨的带,其中5条为稳定的差异带.用Yr10基因的供体亲本Moro和感病品种铭贤169杂交产生的F₂代分离群体,对5个特异条带与目的基因的遗传连锁性进行了初步分析,发现特异条带PT0502与Yr10基因连锁.将该片段进行克隆、测序,根据其序列设计了PCR特异扩增用专化引物.经用195株F₂代分离株进行遗传连锁性分析表明,用所设计的引物进行PCR,其主要产物SC₂₀₀片段与抗条锈病基因Yr10的连锁距离为0.5 cM,可用于该基因的快速、有效、准确检测.

1997

3.658

0.0

2010

0.0

. 2010, 30:11-16

Exploitation and application of a new SCAR marker linked to strip rust resistance gene Yr17 in wheat

(小麦抗条锈基因Yr17的新SCAR标记的建立与应用) . (麦类作物学报) , 2010 , 30 : 11 - 16 Jia J Q , Lei M P , Liu C , Li G R , Yang Z J . Exploitation and application of a new SCAR marker linked to strip rust resistance gene Yr17 in wheat

Jia J Q

(贾举庆) , (雷孟平) , (刘成) , (李光蓉) , (杨足君) . (小麦抗条锈基因Yr17的新SCAR标记的建立与应用) . (麦类作物学报) , 2010 , 30 : 11 - 16 Jia J Q

为深入了解四川小麦新品种(系)中抗条锈病基因的组成,并为开展分子标记辅助育种提供依据,以小麦抗条锈病基因Yr17的近等基因系以及抗条锈病品系L15与感条锈病材料SY95-71的杂交F2代群体为材料,采用集群分离分析法(BSA)结合SSR分子标记进行分析,发现1条由引物Xgwm415扩增的与抗条锈基因Yr17连锁的特异带.对该特异带克隆测序发现,该序列长390 bp,与栽培大麦中的一个序列表达标签(EST)有90%的相似性,与水稻基因组中的一个"核苷酸结合位点(NBS)-富亮氨酸重复(LRR)"型抗病基因的编码区有70%的相似性.根据该序列设计特异引物,建立SCAR标记,命名为SC-372.利用SC-372对54个四川省近年育成的小麦品种(系)进行检测,发现在15个抗病品种(系)中能特异扩增.进一步利用SC-372及前人报道的Yr17连锁标记VENTRUP/LN2对Yr17近等基因系及偏凸山羊草进行扩增比较发现,本实验建立的SCAR标记对小麦背景中的Yr17基因有更好的检测效率.因此可将SC-372用于分子标记聚合小麦抗条锈病基因的育种实践中.

Abstract：

To further clarify the composition of stripe rust resistance gene in the newly developed wheat cultivars (lines) of Sichuan and establish molecular marker-assisted breeding system for rust resistance, the Thatcher Yr17 near isogeneic lines (NILs) and a F2 population derived from a cross between developed resistant line L15 and susceptible line SY95-71 were used to screen the SSR markers by bulked segregation analysis (BSA). A PCR product amplified by Xgwm415 was linked to resistance gene to strip rust resistance gene Yr17. The 390bp sequence of the PCR fragment had 90% identity to one EST of barley and 70% identity to the CDS of a NBS-LRR disease resistance protein in rice. Basing on this sequence, a pair of sequence-characterized amplified region (SCAR) primers were designed, and the SCAR marker, named SC-372, was able to target 15 resistant cultivars (lines) in all 54 wheat cultivars of newly developed in Sichuan Province. Compared with the amplification of marker VENTRUP/LN2 linked to Yr17, the SC-372 marker showed a higher detection efficiency to Yr17 when a set of NILs and Aegilops ventricosa were amplified. It was therefore to conclude that the new SCAR marker for Yr17 gene and the identified resistant lines could be used in wheat breeding for stripe rust resistance.

2006

3.658

0.0

. 2006, 114:21-30 DOI:10.1007/s00122-006-0406-z

Molecular genetic characterization of the Lr34/Yr18 slow rusting resistance gene region in wheat

1.CSIRO Plant Industry, GPO Box 1600, Canberra, ACT 2601, Australia 2.International Maize and Wheat Improvement Center (CIMMYT), Apartado Postal 6-641, 06600 Mexico, D.F., Mexico 3.Campo Experimental Valle de Mexico-INIFAP, Apdo. Postal 10, 56230 Chapingo, Edo. de Mexico, Mexico 4.Plant Breeding Institute, University of Sydney, Cobbitty, NSW 2570, Australia

Wheat expressed sequence tags (wESTs) were identified in a genomic interval predicted to span the Lr34/Yr18 slow rusting region on chromosome 7DS and that corresponded to genes located in the syntenic region of rice chromosome 6 (between 2.02 and 2.38 Mb). A subset of the wESTs was also used to identify corresponding bacterial artificial chromosome (BAC) clones from the diploid D genome of wheat (Aegilops tauschii). Conservation and deviation of micro-colinearity within blocks of genes were found in the D genome BACs relative to the orthologous sequences in rice. Extensive RFLP analysis using the wEST derived clones as probes on a panel of wheat genetic stocks with or without Lr34/Yr18 revealed monomorphic patterns as the norm in this region of the wheat genome. A similar pattern was observed with single nucleotide polymorphism analysis on a subset of the wEST derived clones and subclones from corresponding D genome BACs. One exception was a wEST derived clone that produced a consistent RFLP pattern that distinguished the Lr34/Yr18 genetic stocks and well-established cultivars known either to possess or lack Lr34/Yr18. Conversion of the RFLP to a codominant sequence tagged site (csLV34) revealed a bi-allelic locus, where a variant size of 79 bp insertion in an intron sequence was associated with lines or cultivars that lacked Lr34/Yr18. This association with Lr34/Yr18 was validated in wheat cultivars from diverse backgrounds. Genetic linkage between csLV34 and Lr34/Yr18 was estimated at 0.4 cM

2005

0.0

1.088

. 2005, 35:478-480

Molecular mapping of stripe rust resistance gene Yr24 in wheat

(小麦抗条锈病基因Yr24的SSR标记) . (植物病理学报) , 2005 , 35 : 478 - 480 Liu Y P , Cao S H , Wang X P , Xu Z B , Zhang X Q . Molecular mapping of stripe rust resistance gene Yr24 in wheat

Liu Y P

(刘亚萍) , (曹双河) , (王献平) , (徐志斌) , (张相岐) . (小麦抗条锈病基因Yr24的SSR标记) . (植物病理学报) , 2005 , 35 : 478 - 480 Liu Y P

抗性鉴定表明,含有抗小麦条锈病基因Yr24的近等基因系Yr24/3*Avocet S对我国流行的条锈菌小种CY30、CY31和CY32均具有良好的抗性.遗传学分析证明,Yr24/3*Avocet S的抗条锈病性状为显性遗传.利用Yr24/3*Avocet S×感病品种铭贤169的F2群体进行SSR分析,筛选到2个位于目的基因两侧的标记Xgwm273和Xgwm11,遗传距离分别为6.1和7.1 cM.双侧分子标记的建立可为标记辅助选择育种提供更有力的分子选择工具.

2001

1.643

0.0

. 2001, 120:219-226 DOI:10.1023/A:1017510331721

Molecular mapping and detection of the yellow rust resistance gene Yr26 in wheat transferred from Triticum turgidum L1 using microsatellite markers

1.Chinese Academy of Agricultural Sciences Institute of Crop Germplasm Resources Beijing 100081 P.R. China Beijing 100081 P.R. China

Yellow rust (stripe rust), caused by Puccinia striiformis Westend f. sp. tritici, is one of the most devastating diseases of wheat throughout the world. Wheat-Haynaldia villosa 6AL.6VS translocation lines R43, R55, R64 and R77, derived from the cross of three species, carry resistance to both yellow rust and powdery mildew. An F₂ population was established by crossing R55 with the susceptible cultivar Yumai 18. The yellow rust resistance in R55 was controlled by a single dominant gene, which segregated independently of the powdery mildew resistance gene Pm21 located in the chromosome 6VS segment, indicating that the yellow rust resistance gene and Pm21 are unlikely to be carried by the same alien segment. This yellow rust resistance gene was considered to beYr26, originally thought to be also located in chromosome arm 6VS. Bulked Segregation Analysis and microsatellite primer screens of the population F₂ of Yumai 18 × R55 identified three chromosome 1B microsatellite locus markers, Xgwm11, Xgwm18 and Xgwm413, closely linked to Yr26. Yr26 was placed 1.9 cM distal of Xgwm11/Xgwml8, which in turn were 3.2 cM from Xgwm413. The respective LOD values were 21 and 36.5. Therefore, Yr26 was located in the short arm of chromosome 1B. The origin and distribution of Yr26 was investigated by pedigree, inheritance of resistance and molecular marker analysis. The results indicated that Yr26 came from Triticum turgidum L. Three other 6AL.6VS translocation lines, R43, R64 and R77, also carried Yr26. These PCR-based microsatellite markers were shown to be very effective for the detection of the Yr26 gene in segregating populations and therefore can be applied in wheat breeding.

2006

1.513

0.0

2006

0.0

2007

3.658

0.0

. 2007, 115:159-168 DOI:10.1007/s00122-007-0551-z

Physical mapping and identification of a candidate for the leaf rust resistance gene Lr1 of wheat

1.Chinese Academy of Sciences State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology Datun Road Chaoyang District, Beijing 100101 China Datun Road Chaoyang District, Beijing 100101 China 2.Graduate University of the Chinese Academy of Sciences 19 Yuquan Road Beijing 100049 China 19 Yuquan Road Beijing 100049 China 3.University of Zurich Institute of Plant Biology Zollikerstrasse 107 8008 Zürich Switzerland Zollikerstrasse 107 8008 Zürich Switzerland

Lr1 is a dominant leaf rust resistance gene located on chromosome 5DL of bread wheat and the wild species Aegilops tauschii. In this study, three polymorphic markers (WR001, WR002, and WR003) were developed from resistance gene analogs (RGAs) clustering around the Lr1 locus. Using these and other markers, Lr1 was mapped to a genetic interval of 0.79 cM in Ae. tauschii and 0.075 cM in wheat. The CAPS marker WR003, derived from LR1RGA1, co-segregated with Lr1 in both mapping populations of wheat and Ae. tauschii. For isolation of Lr1, two genomic BAC libraries (from Ae. tauschii and hexaploid wheat) were screened using the tightly flanking marker PSR567F and a set of nested primers derived from the conserved region of the RGA sequences. Approximately 400 kb BAC contig spanning the Lr1 locus was constructed. The LR1RGA1 encoding a CC-NBS-leucine-rich repeat (LRR) type of protein was the only one of the four RGAs at the Lr1 locus, which co-segregated with leaf rust resistance. Therefore, it represents a very good candidate for Lr1. The allelic sequences of LR1RGA1 from resistant and susceptible lines revealed a divergent DNA sequence block of ∼605 bp encoding the LRR repeats 9–15, whereas the rest of the sequences were mostly identical. Within this sequence block, the 48 non-synonymous changes resulted in 44 amino acid differences. This indicates that LR1RGA1 likely evolved through one or more recombination or gene conversion events with unknown genes.

1994

3.658

0.0

1997

3.251

0.0

2001

3.658

0.0

. 2001, 103:618-624 DOI:10.1007/PL00002918

AFLP and STS tagging of Lr19, a gene conferring resistance to leaf rust in wheat

1.ARC-Small Grain Institute, P.O. Box 3507, Matieland, South Africa, 7602 e-mail: rprins@maties.sun.ac.za Fax: +27-21-8874171 ZA ZA 2.Department of Genetics, Private Bag X1, Matieland, South Africa, 7602 ZA ZA 3.Cereals Research Department, John Innes Centre, Norwich Research Park, Colney Lane, Norwich NR4 7UH, UK GB GB

Amplified fragment length polymorphism (AFLP) markers were used to enrich the map of the wheat chromosomal region containing the Thinopyrum-derived Lr19 leaf rust resistance gene. The region closest to Lr19 was targeted through the use of deletion and recombinant lines of the translocated segment. One of the AFLP bands thus identified was converted into a sequence-tagged-site (STS) marker. This assay generated a 130-bp PCR fragment in all Lr19-carrying lines tested, except for one deletion mutant, while non-carrier template failed to amplify any product. This sequence represents the first marker to map on the distal side of Lr19 on chromosome 7el₁. The conversion process of AFLP fragments to STS markers was technically difficult, mainly because of the presence of contaminating fragments. Various approaches were taken to reduce the frequency of false positives and to identify the correct clone. We were able to formulate a general verification strategy prior to clone sequencing. Various other factors causing problems with converting AFLP bands to an STS assays are also discussed.

2002

1.668

0.0

2001

3.658

0.0

. 2001, 103:1007-1013 DOI:10.1007/s001220100701

An RGA-like marker detects all known Lr21 leaf rust resistance gene family members in Aegilops tauschii and wheat

1.Wheat Genetics Resources Center, Department of Plant Pathology, Throckmorton Plant Science Center, Kansas State University, Manhattan, KS 66506-5502, USA e-mail: bsg@ksu.edu US US

Leaf rust is one of the most important diseases of wheat worldwide, particularly in the Great Plains region of the USA. One long-term strategy for the control of this disease may be through durable genetic resistance by gene pyramiding. An important step in this strategy is identifying molecular markers linked to different leaf rust-resistance genes. Here we report the molecular tagging of a leaf rust-resistance gene that may have the potential for durable resistance through further genetic manipulation and gene pyramiding. Lr39 was previously designated for a leaf rust-resistance gene introgressed from Aegilops tauschii accession TA1675 into the common wheat germplasm WGRC2. Lr40 was designated for a gene derived from Ae. tauschii accession TA1649 and is present in germplasm WGRC7. These genes are now believed to be allelic to Lr21, which was transferred to wheat from a different accession of Ae. tauschii. Molecular mapping of Lr39 and Lr40 indicates that both genes come from TA1649. WGRC2 and WRGC7 also have a similar infection type against rust culture PRTUS6. We suggest the designation of the gene in WGRC2 should be changed to Lr40. RFLP marker KSUD14 (locus Xksud14) was found 0.2-cM proximal to Lr40 in a WGRC2/Wichita F₂ population (218 individuals), and co-segregated with the gene in a WGRC7/ Wichita F₂ population (165 individuals). A PCR-based molecular marker developed from the sequence-tagged-site (STS) of Xksud14 was mapped to the same locus as the RFLP marker KSUD14 in both populations. KSUD14 has the structure of a resistance gene analog (RGA) including kinase2a and kinase3 domains similar to the Cre3 gene of wheat and the rust resistance gene Rp1-D of maize. When the PCR products amplified from KSU14 STS were cleaved with restriction enzyme MspI, an 885-bp fragment was found in WGRC2, WGRC7, the Lr21 near-isogenic line, and eight accessions of Ae. tauschii shown to have resistance gene alleles at the Lr21 locus. The KSUD14 PCR-based assay provides an excellent marker for Lr40 and Lr21 in diverse wheat breeding and wild Ae. tauschii populations.

1995

3.658

0.0

2002

3.658

0.0

. 2002, 104:1317-1324 DOI:10.1007/s00122-002-0879-3

Identification and mapping of molecular markers linked to rust resistance genes located on chromosome 1RS of rye using wheat-rye translocation lines

1.CSIRO Plant Industry, GPO Box 1600, Canberra, ACT 2601, Australia

2002

0.0

1999

0.0

2008

0.0

1.889

. 2008, 41:3604-3609

A new marker tagged to the leaf rust resistance gene Lr38

(小麦抗叶锈基因Lr38的一个新标记) . (中国农业科学) , 2008 , 41 : 3604 - 3609 Yan H F , Yang W X , Chu D , Liu D Q . A new marker tagged to the leaf rust resistance gene Lr38

Yan H F

(闫红飞) , (杨文香) , (褚栋) , (刘大群) . (小麦抗叶锈基因Lr38的一个新标记) . (中国农业科学) , 2008 , 41 : 3604 - 3609 Yan H F

河北农业大学植物病理学系/河北省农作物病虫害生物防治工程技术研究中心

【目的】建立小麦抗叶锈近等基因系材料TcLr38特异的PCR标记。【方法】以Thatcher为对照，以TcLr38和TcLr38×Thatcher F2代单株构建的分离群体为材料，利用TcLr38远缘亲本中间偃麦草特异的SCAR标记引物进行PCR扩增获得单一条带，并以50个小麦抗叶锈近等基因系材料对该标记特异性进行检测，Mapmaker 3.0软件计算该标记与Lr38的遗传连锁距离。【结果】中间偃麦草特异的SCAR标记引物在TcLr38中获得单一扩增片段大小为982 bp，50个小麦抗叶锈近等基因系检测表明为TcLr38的特有条带，F2代分离群体验证表明为与Lr38共分离的分子标记，命名为Y38SCAR982。【结论】Y38SCAR982可以作为Lr38的分子标记，并证实该基因由中间偃麦草向普通小麦转移的事实。