小麦品种(系)抗叶锈病基因<italic>Lr10</italic> 位点基因型的多样性

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韩淑晓, 刘全兰, 董洁, 陈建省, 田纪春. 小麦品种(系)抗叶锈病基因Lr10 位点基因型的多样性 . 作物学报, 2013, 39(11): 1983-1991[HAN Shu-Xiao, LIU Quan-Lan, DONG Jie, CHEN Jian-Sheng, TIAN Ji-Chun. Genotypic Diversity atLr10 Locus for Leaf Rust Resistance in Various Hexaploid Wheat Varieties (Lines) . 作物学报, 2013, 39(11): 1983-1991] 复制到剪切板

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小麦品种(系)抗叶锈病基因Lr10 位点基因型的多样性

韩淑晓¹, 刘全兰², 董洁¹, 陈建省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青岛科技大学化工学院生物工程与技术系, 山东青岛266042

^* 通讯作者(Corresponding author): 田纪春, E-mail:jctian@sdau.edu.cn, Te1: 0538-8242040

收稿日期:2013-06-24

基金:本研究由国家自然科学基金项目(31171554)和国家转基因生物新品种培育科技重大专项(2011ZX08002-003)资助。

摘要

小麦抗叶锈病基因Lr10 的表达需与其紧密连锁的RGA2 基因调控, 按照这2个基因的连锁关系,Lr10 基因位点分成H1和H2两个古单倍型。为揭示我国小麦品种中Lr10 的遗传多样性, 对189个来自12省的小麦育成品种和58个品系的该基因位点变异进行了鉴定分析。绝大多数供试品种(系)为H2古单倍型, 其频率在育成品种和品系中分别为95.2% (180/189)和96.6% (56/58)。这2种古单倍型可进一步分为9种单倍型亚型, 其中H1-2、H2-4、H2-5、H2-6和H2-7亚型为首次报道。育成品种包含所有单倍型亚型, 以H2-1频率最高(69.3%), H2-3和H2-6频率最低(0.5%); 而在选育品系中仅检测到5种单倍型亚型, 其中H2-1频率最高(27.6%), H1-2频率最低(3.5%)。除H2-1与H2-2型外, 其他亚型与育成地显著相关(P< 0.05)。本研究结果支持Lr10 基因位点的古单倍型和单倍型亚型在育种中保持相对稳定的推断, 同时由于在育种过程中持续重组和变异而产生了新的单倍型亚型, 而且重组可以发生在H2和H1两种古单倍型之间。在现代育成品种和品系中, H1古单倍型所占比例已降至5%以下, 应加以保护。

关键词: 小麦; Lr10; 古单倍型; 单倍型亚型; 遗传多样性

Genotypic Diversity atLr10 Locus for Leaf Rust Resistance in Various Hexaploid Wheat Varieties (Lines)

HAN Shu-Xiao¹, LIU Quan-Lan², DONG Jie¹, CHEN Jian-Sheng¹, TIAN Ji-Chun^1,^*

¹State Key Laboratory of Crop Biology / Shandong Provincial Key Laboratory of Crop Biology, Shandong Agricultural University, Tai’an 271018, China

² Department of Bioengineering and Biotechnology, College of Chemical Engineering, Qingdao University of Science and Technology, Qingdao 266042, China

Abstract

Genotypes H1 and H2 are two ancient haplotypes onLr10 locus, which is a resistance gene to wheat leaf rust. They are identified based on the presence of bothLr10 andRGA2 in full length (H1), and the absence ofLr10 and chromatin rearrangement ofRGA2 (H2) due to chromosome 1AS reorganization during species evolution. Both haplotypes contain several subhaplotypes. To understand the genetic diversity onLr10 locus, we tested the frequencies of H1 and H2 haplotypes and their subhaplotypes in 189 wheat cultivars and 58 breeding lines from various provinces of China. The H2 haplotype was dominant in the developed cultivars and breeding lines with the frequencies of 95.2% (180/189) and 96.6% (56/58), respectively. Five novel subhaplotypes were detected in the developed cultivars, namely H1-2, H2-4, H2-5, H2-6, and H2-7. In cultivars, subhaplotype H2-1 was the most frequent (69.3%), whereas subhaplotypes H2-3 and H2-6 had the lowest frequencies (0.5%). Among the five subhaplotypes detected in breeding lines, subhaplotype H2-1 had the highest frequencies (27.6%), whereas subhaplotypes H1-2 was the least frequent (3.5%). Interestingly, the subhaplotypes except for H2-1 and H2-2 were highly breeding location-dependent (P< 0.05). These results support the hypothesis that both haplotypes have been maintained through a balanced polymorphism mechanism during wheat domestication. Furthermore, the five novel subhaplotypes suggest that genetic diversities resulting from recombination and other types of chromatin reorganizations (e.g., origin of the H2 haplotype and polymorphism for theLr10 locus) have been going on since the formation of the hexaploid wheat. Significantly, identification of the H1-2 subhaplotype is a clear evidence of occurrence of recombination between the two ancient haplotypes, which disagrees with the previous notion. Importantly, because the frequencies of H1 is lower than 5% in developed cultivars and breeding lines, actions for protecting germplasm with the H1 haplotype are highly recommended.

Keyword: Wheat; Lr10; Ancient haplotype; Subhaplotype; Genetic diversity

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引言

小麦是全球主要粮食作物之一, 种植面积仅次于玉米。叶锈病是小麦的主要病害之一, 发病严重时可减产50%^[1]。小麦叶锈病的病原菌为 Puccinia triticina , 具有小种专化性, 对不同小麦品种具有不同的致病性^[2]; 但病原小种易发生变异, 致使抗病品种丧失抗性, 特别是大面积推广品种都高抗当时流行的生理小种, 加速了病原菌的定向选择和新生理小种的形成。因此, 研究抗叶锈基因在小麦品种(系)中的多样性和分布规律, 对合理利用抗叶锈病基因资源, 延缓抗病品种的抗性丧失具有重要意义。

抗叶锈基因 Lr10 位于小麦1A染色体短臂上^[3,4],与另一抗病基因同源序列 RGA2 紧密连锁, 在该位点存在H1和H2两种古单倍型^[3]。H1古单倍型含有完整的 Lr10 基因及与其紧密连锁的 RGA2 基因, H2型仅含有 RGA2 的部分序列, 缺失 RGA2 的5′端部分序列及全部 Lr10 序列^[3,4,5,6]。群体遗传学研究表明, 这两种古单倍型原存在于一粒小麦( Triticum urartu )和野生二粒小麦( T. dicoccoides )中^[3,4,5,6]。在小麦进化和选育过程中, Lr10 基因在普通小麦中被保留下来, 并对叶锈菌菌株( AvrLr10 ) 89-201 CBTB(TX)表现抗性^[5]。Stein等^[5]检测了 Lr10 古单倍型在56份欧洲小麦品系中的分布规律, 发现H1古单倍型的比例为14.29% (8/56), H2古单倍型的比例为85.71% (48/56), 说明H2古单倍型是欧洲普通小麦的主要类型。Isidore等^[6]设计了4对H2单倍型的特异引物(A、B、C和B_3kb)用于研究不同倍性小麦中H2单倍型的保守性, 共检测出3种H2亚型, 在欧洲普通小麦中仅检测出2种H2亚型。尽管前人对 Lr10 基因位点的两种古单倍型及其亚型在小麦中的分布规律已有较多研究, 但 Lr10 基因位点在中国小麦品种中的分布规律未见报道。

小麦育种过程中多选用少数综合性状好、配合力好的品种为亲本, 并以高产、优质、抗病虫和抗旱等为主要育种目标对后代进行更强有力的选择。这种强目的性的人工选择必将强化基因的瓶颈效应, 导致育成品种遗传基础变窄, 遗传多样性降低。与古老的地方品种和野生品种相比, 现代育成品种遗传多样性明显降低。为评价 Lr10 基因位点在中国小麦品种中的遗传多样性及可能经历的遗传瓶颈效应, 本研究选用247份中国小麦品种(系), 检测 Lr10 基因位点的古单倍型及其等位基因的遗传变异, 旨在阐明 Lr10 基因位点的H1和H2古单倍型在小麦驯化过程中的遗传瓶颈效应, Lr10 基因位点在中国小麦品种(系)中的等位变异及其遗传多样性, 及 Lr10 基因位点的多样性机制。

1 材料与方法

1.1 植物材料

189个小麦育成品种和58个高代品系(见附表), 均由国家小麦改良中心泰安分中心提供。189个育成品种包括山东103份、河北31份、河南22份、陕西10份、江苏7份、山西5份、北京4份、安徽2份、宁夏2份, 以及西藏、新疆和云南各1份; 58个高代品系均由国家小麦改良中心泰安分中心近年新育成。2011—2012年, 将所有品种(系)种植于山东农业大学试验站, 每个材料种植3行, 行长3.00 m, 行宽0.26 m, 四周各有1 m保护行。生育期间肥水按高产田管理, 未进行药剂防治病害, 小麦亦未发生倒伏。

1.2 Lr10 基因位点相关片段的PCR扩增

采用Doyle和Doyle^[7]的方法从小麦幼苗提取的基因组DNA。采用引物Control^[6]和rga1Pro^[8], 通过PCR扩增检测H1和H2单倍型, 其中引物Control (表1)用于扩增H1和H2两种单倍型的共同序列(图1), 引物rga1Pro (表1)特异性扩增 Lr10 基因的5′端非编码区及部分5′端编码区, 只在H1型个体中有扩增产物(图1)。引物ThLr10_T/P、ThLr10_E/H和ThLr10_G/J^[9]用于检测H1的亚型; 而引物A、B、C和B_3k^[6]用于检测H2的亚型。所有引物均由生工生物工程(上海)股份有限公司合成。

在T-gradient Thermal Cycler (Bio-metra)上, 按已报道的方法^[6,8,9]进行PCR扩增。扩增产物经琼脂糖电泳后, 1%~2%溴化乙锭染色, 用Bioshine GelX 1650凝胶成像系统(上海欧翔科学仪器有限公司)观察、照相、读带。

1.3 数据分析

Lr10 位点基因型按照PCR扩增片段的有或无命名。例如, 若两对引物Control和rga1Pro都有PCR产物, 则其古单倍型的类型为H1^[8]; 若仅引物Control有PCR产物, 引物rga1Pro无扩增产物, 则其古单倍型的类型为H2^[8]。H1和H2单倍型亚型的扩增带型如表2所示。采用SAS 9.0软件^[10]统计各古单倍型及其亚型的发生频率以及基因型。选用样品数较多的山东、河北、河南和陕西材料进行 Lr10 位点基因型与品种育成地的相关性分析。

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	图 1 引物扩增片段在 Lr10 位点的位置序列结构图引自Isidore 等^[6]的图2和图5, 有修改。Fig . 1 Positions of amplified fragments used different primers at locus Lr10The sequence structures were modified from the original Figs. 2 and 5 in the article by Isidore et al.^[6]

表 1 用于PCR扩增的引物序列 Tab le 1 Primers used in this study

表2 按PCR产物命名的基因型 Table 2 Genotypes classified by different amplicons produced by different primers

2 结果与分析

2.1 中国小麦品种(系)中 Lr10 基因型的多样性

小麦品种或品系中均检测到H1和H2两种古单倍型(图2), 但两种古单倍型的发生频率有显著差异(表3)。在189份小麦品种中, 有180份材料的古单倍

型为H2, 占供试品种的95.2%, 仅有9份材料含有H1单倍型, 占供试品种的4.8%; 在58份高代品系中, H2和H1单倍型的频率分别为96.6% (56/58)和3.5% (2/58)。

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图2 Lr10 基因位点的引物在小麦品种(系)中的扩增图谱M: 标准分子量DNA (Trans2K plus); 1: 濮兴02-16; 2: 红地963; 3: 石09-4276; 4: 高-W16; 5: 航麦0903; 6: LS6045; 7: 荷麦0302; 8: 西农9814; 9: 明麦1号; 10: 泰农984; 11: 22037; 12: 89100S。Fig. 2 Amplification profiles on locus Lr10 in wheat varieties (lines) and breeding linesM: tz DNA marker (Trans2K plus); 1: Puxing 02-16; 2: Hongdi 963;3: Shi 09-4276;4: Gao-W16;5: Hangmai 0903; 6: LS6045;7: Hemai 0302;8: Xinong 9814;9: Mingmai 1; 10: Tainong 984; 11: 22037; 12: 89100S.

表3 小麦品种(系)中 Lr10 位点各单倍型及其亚型发生频率 Table 3 Frequencies of genotypes at locus Lr10 from common wheat varieties (lines) and breeding lines

2.2 Lr10 基因型遗传多样性与品种育成地的相关性

在来自山东、河北、河南和陕西的育成品种中, H2-1型( P =0.2701)和H2-2型( P =0.51715)与育成地没有相关性, H2-3 ( P =0.04701)、H2-4 ( P = 0.0464)、H2-5 ( P =0.0397)和H2-7型( P =0.0100)与育成地有显著相关。H2-1和H2-2型广泛分布于这4个省, H2-3和H2-4型仅分布在山东, H2-5型分布在山东、河南和陕西, H2-7型仅分布在河北(图3); 而H2-6型在这4个省份中都没有分布, 仅在江苏的小麦品种中检测到(见附表)。总之, 除基因型H2-1和H2-2外, 其他基因型与其育成地存在显著相关性( P< 0.05)。

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	图3 4省小麦品种中基因型的分布品种(系)序号同附表中。Series numbers of wheat varieties (lines) correspond to the numbers for variety names given in the Appendix.Fig. 3 Distribution of genotypes across common wheat varieties (lines) from four provinces

3 讨论

3.1 Lr10 基因位点的遗传瓶颈效应

长期的自然选择和人工驯化最终将导致小麦基因的遗传多样性降低^[11]。尽管如此, 小麦基因组通过其可塑性可以弥补这种瓶颈效应。Dubcovsky和Dvorak^[12]指出, 小麦可以通过其A、B和D基因组的基因补偿效应来弥补基因瓶颈效应造成的不良后果, 杂交育种可以使新育成的小麦获得优于亲本的基因型, 杂交后代通过基因突变、重组、缺失等方式也可以形成全新的基因型。Stein等^[5]和Isidore等^[6]的研究也表明, 小麦的抗病基因同样经历了类似的遗传瓶颈效应, 但这些抗病基因经历瓶颈效应的程度和可能获得的新基因并没有得到深入研究。

Lr10 基因是小麦中4个已测序抗叶锈病基因之一^[4](http://www.ncbi.nlm.nih.gov/genbank/), 全长为3935 bp, 与其他抗叶锈病基因序列的相似度较低^[4]。该基因编码919个氨基酸长度的CC-NBS-LRR类型蛋白质^[9], 在蛋白CC结构域的N端经历多样性选择(diversifying selection)^[13]。另一抗病基因同源序列 RGA2 与 Lr10 基因紧密连锁, 是 Lr10 基因表现出抗性必需的调控基因^[13]。陈万权和王剑雄^[14]采用基因推导法在来自世界各地的76个小麦种质资源中鉴定出4个材料含 Lr10 基因; 丁艳红等^[15]采用STS分子标记在28个小麦微核心种质中鉴定出2个材料含 Lr10 基因。本研究在189份小麦品种中仅鉴定出9个同时含有 Lr10 基因和 RGA2 基因的品种, 其中2个品种含有 Lr10 基因和缺失的 RGA2 基因(表3); 在58份小麦品系中没有鉴定出同时含有完整 Lr10 基因和 RGA2 基因的材料, 仅有2个品系同时含有 Lr10 基因和缺失的 RGA2 基因(表3)。上述研究结果表明, Lr10 基因经历了强烈的瓶颈效应, 推测导致这种结果的原因有2个: 其一, 当前流行的叶锈病菌已经对 Lr10 基因产生抗性, 因而, 小麦抗病育种工作者已不选育含有此基因的材料^[2]; 其二, Lr10 基因可能对某些稀有致病小种或未知种有抗性, 但对当前流行的叶锈病病原菌没有效果, 且该基因的表达对产量或其他农艺性状有负面效应, 最终导致育种工作者不选育含有此基因的材料^[4]。我们认为这两种解释都有其合理性, 同时也认为国内外学者采用研究方法的不同也可能导致上述解释的差异。

Isidore等^[6]发现, 欧洲小麦中 Lr10 位点的H2-1型占优势。本研究检测结果表明, 该亚型在中国小麦中也有最高的发生频率, 其次是是H2-2 (表3), 同时也说明, H2-1基因型可能是世界性的优势基因型, 但导致H2-1基因型占优势的原因还需进一步探究。另外, Lr10 位点的瓶颈效应说明需要保护那些基因型发生频率低的材料, 如含H1-1基因型的材料, 这类材料含有较完整的 Lr10 基因和 RGA2 基因。

3.2 Lr10 基因位点的进化

致病菌的高度变异为植物抗病基因的进化提供了很强的自然选择压, 关于两者的相关性主要有2种假说, 即trench-warfare模型^[16]和arm-race模型^[17]。前者认为抗病基因进化形成的多样性依赖于致病菌的变异频率, 频率高时抗病基因的多样性高, 频率低时抗病基因的多样性低, 在此模型下, 抗病基因的多样性较高, 可以形成新基因和保留已有基因, 如拟南芥中的基因 Rpm1^[16]、 RPP8^[17]、 RPP13 等^[17]; 后者是birth-death模型^[18]的补充完善, 强调抗病基因的多样性有明显的方向性, 即选择更具抗性的基因, 在此模型下, 抗病基因的多样性很低, 有明显的方向性, 如番茄中对尖孢镰孢菌( Fusarium oxysporum )产生抗性的抗病基因^[19]。 Lr10 基因的进化模式为trench-warfare模型, 在野生群体中具有较高的遗传多样性^[4,20]。Sela等^[20]推测重组使 Lr10 基因具有较高的多样性; Isidore等^[6]推测逆转录子的插入、转座子的缺失以及 Lr10 基因与 RGA2 基因间元件的不等重组等事件形成H2古单倍型; Isidore等^[6]还推测H2和H1古单倍型之间非常稳定, 不会发生重组。本研究的结果支持Sela等^[20]的结论, 即H2和H1古单倍型确实存在于小麦品种中, 而且H2-1和H2-2基因型分布广泛并相对稳定, 同时又修正了Isidore等^[6]关于重组不能发生在H2和H1古单倍型之间的推断。

本研究鉴定出了5个新的单倍型亚型(H1-2、H2-4、H2-5、H2-6和H2-7, 见附表), 证明了重组机制在小麦育种过程中持续存在, 杂交育种可以创造出新的 Lr10 基因型。本研究结果证明, 在小麦育种过程中确实有些材料丢失了 Lr10 基因、 RGA2 基因及两基因间的元件, 但小麦通过重组机制可以产生新基因型, 以获得基因的可塑性来创造新的抗病性状。

4 结论

247份中国小麦育成品种和高代品系中H2型为优势基因型, 其中H2-1型出现频率最高。在育成品种中发现5个新单倍型亚型, 分别是H1-2、H2-4、H2-5、H2-6和H2-7, 其频率分别为1.1%、1.1%、2.1%、0.5%和1.6%。在现代小麦品种中, H1古单倍型所占比例低于5%, 亟需进行遗传资源保护。

附表材料信息及其 Lr10 位点基因型 Appendix Information of materials and their genotypes at locus Lr10

(续附表) 材料信息及其 Lr10 位点基因型 Appendix Information of materials and their genotypes at locus Lr10

序号No.	品种(系) Variety (line)	来源Origin	古单倍型 Ancient haplotype	单倍型亚型Subhaplotype	序号No.	品种(系) Variety (line)	来源Origin	古单倍型 Ancient haplotype	单倍型亚型Subhaplotype
29	滨轮77	山东	H2	H2-1	153	周麦22	河南	H2	H2-1
30	滨系10	山东	H2	H2-1	154	周麦25	河南	H2	H2-2
31	登海59	山东	H2	H2-2	155	驻0263-07142	河南	H2	H2-1
32	东方0516	山东	H2	H2-1	156	驻0324-06169	河南	H2	H2-1
33	东方09-8	山东	H1	H1-1	157	陕225-2	陕西	H2	H2-5^a
34	高-W16	山东	H2	H2-1	158	陕农84	陕西	H2	H2-1
35	航麦0903	山东	H2	H2-1	159	陕农131	陕西	H2	H2-1
36	菏麦0302	山东	H2	H2-4^a	160	陕农22SN055843	陕西	H2	H2-1
37	菏麦9946	山东	H2	H2-2	161	陕农33	陕西	H2	H2-1
38	红地963	山东	H2	H2-2	162	陕农342	陕西	H2	H2-1
39	济麦16	山东	H2	H2-1	163	陕农981	陕西	H2	H2-2
40	济麦22	山东	H2	H2-1	164	西农9814	陕西	H2	H2-1
41	济麦7195	山东	H2	H2-1	165	小偃166	陕西	H2	H2-2
42	济南17	山东	H2	H2-2	166	小偃6号	陕西	H2	H2-2
43	济宁045066	山东	H2	H2-1	167	华瑞1032	江苏	H2	H2-6^a
44	济宁07412	山东	H2	H2-1	168	连5036	江苏	H2	H2-2
45	金莱679	山东	H2	H2-1	169	连麦2	江苏	H2	H2-1
46	良星601	山东	H2	H2-1	170	明麦1号	江苏	H2	H2-2
47	良星99	山东	H2	H2-2	171	徐麦8093	江苏	H2	H2-1
48	聊0801	山东	H2	H2-1	172	扬06-164	江苏	H2	H2-1
49	聊9817	山东	H2	H2-1	173	扬06G5	江苏	H2	H2-1
50	临061	山东	H2	H2-3	174	西农157	山西	H1	H1-1
51	鲁麦205	山东	H2	H2-1	175	西农501	山西	H2	H2-1
52	鲁原185	山东	H2	H2-1	176	西农529	山西	H1	H1-2^a
53	鲁原208	山东	H2	H2-1	177	西农822	山西	H1	H1-1
54	鲁原262	山东	H2	H2-2	178	西农919	山西	H2	H2-2
55	鲁淄0318-1	山东	H2	H2-1	179	科遗4058	北京	H2	H2-2
56	鲁淄588	山东	H2	H2-1	180	科遗5214	北京	H2	H2-2
57	轮选165	山东	H2	H2-1	181	中麦349	北京	H2	H2-2
58	轮选799	山东	H2	H2-2	182	中麦895	北京	H2	H2-1
59	齐丰2号	山东	H2	H2-1	183	天禾3号	安徽	H2	H2-1
60	千禾麦17	山东	H2	H2-1	184	天禾6号	安徽	H2	H2-1
61	山农055849	山东	H2	H2-1	185	宁冬10号	宁夏	H2	H2-1
62	山农06-278	山东	H2	H2-1	186	宁冬11	宁夏	H2	H2-1
63	山农0905	山东	H2	H2-1	187	0901300988044	西藏	H2	H2-2
64	山农10-2	山东	H2	H2-1	188	0901051-新疆	新疆	H2	H2-2
65	山农1151	山东	H2	H2-1	189	南0901068M1	云南	H2	H2-1
66	山农17	山东	H2	H2-2	190	20668	NWIC-TA	H2	H2-2
67	山农19	山东	H2	H2-1	191	21916	NWIC-TA	H2	H2-5^a
68	山农20	山东	H2	H2-1	192	22037	NWIC-TA	H2	H2-4^a
69	农23SN032714	山东	H2	H2-1	193	22798	NWIC-TA	H2	H2-2
70	山农2478	山东	H2	H2-1	194	26115	NWIC-TA	H2	H2-2
71	山农2657	山东	H2	H2-1	195	26146	NWIC-TA	H1	H1-2^a
72	山农55843	山东	H2	H2-1	196	28642	NWIC-TA	H2	H2-2
73	山农711	山东	H2	H2-1	197	32752	NWIC-TA	H2	H2-4^a
74	山农优麦3号	山东	H2	H2-1	198	33117	NWIC-TA	H2	H2-5^a
75	圣丰2号	山东	H2	H2-1	199	33786	NWIC-TA	H2	H2-5^a
76	泰农 9571	山东	H2	H2-1	200	34455	NWIC-TA	H2	H2-1
77	泰农2987	山东	H2	H2-1	201	36039	NWIC-TA	H2	H2-1

(续附表) 材料信息及其 Lr10 位点基因型 Appendix Information of materials and their genotypes at locus Lr10

序号No.	品种(系) Variety (line)	来源Origin	古单倍型 Ancient haplotype	单倍型亚型Subhaplotype	序号No.	品种(系) Variety (line)	来源Origin	古单倍型 Ancient haplotype	单倍型亚型Subhaplotype
78	泰农7018	山东	H2	H2-2	202	41296	NWIC-TA	H2	H2-1
79	泰农8681	山东	H2	H2-1	203	41300	NWIC-TA	H2	H2-2
80	泰农9236	山东	H2	H2-1	204	41484	NWIC-TA	H2	H2-5^a
81	泰农984	山东	H2	H2-4^a	205	41569	NWIC-TA	H2	H2-4^a
82	泰农9862	山东	H2	H2-1	206	102-8	NWIC-TA	H2	H2-1
83	泰山 008-HF0901027	山东	H2	H2-2	207	106-1	NWIC-TA	H2	H2-4^a
84	泰山4173	山东	H2	H2-1	208	156-19	NWIC-TA	H2	H2-5^a
85	泰山4241	山东	H2	H2-1	209	171-1	NWIC-TA	H2	H2-1
86	潍64510	山东	H2	H2-1	210	185-15	NWIC-TA	H2	H2-5^a
87	潍麦8-0901010	山东	H2	H2-2	211	213-9	NWIC-TA	H2	H2-5^a
88	汶航1号	山东	H2	H2-1	212	217-1	NWIC-TA	H2	H2-1
89	汶农17	山东	H2	H2-1	213	220-7	NWIC-TA	H2	H2-2
90	汶农18	山东	H2	H2-1	214	232-5	NWIC-TA	H2	H2-2
91	汶农19	山东	H2	H2-1	215	238-10	NWIC-TA	H2	H2-4^a
92	新山农11	山东	H2	H2-1	216	247-25	NWIC-TA	H2	H2-4^a
93	鑫麦296	山东	H2	H2-1	217	258-7	NWIC-TA	H2	H2-4^a
94	烟0881	山东	H1	H1-1	218	305-6	NWIC-TA	H2	H2-1
95	烟09135	山东	H2	H2-1	219	315-419	NWIC-TA	H2	H2-1
96	烟99102	山东	H2	H2-1	220	315-461	NWIC-TA	H2	H2-2
97	烟农10854	山东	H2	H2-1	221	315-506	NWIC-TA	H2	H2-1
98	烟农578	山东	H2	H2-2	222	315-526	NWIC-TA	H2	H2-1
99	优麦2号	山东	H2	H2-2	223	318-418	NWIC-TA	H2	H2-5^a
100	早熟6061	山东	H1	H1-1	224	319-199	NWIC-TA	H2	H2-2
101	枣1802	山东	H2	H2-1	225	319-590	NWIC-TA	H2	H2-1
102	枣1864	山东	H2	H2-1	226	319-591	NWIC-TA	H2	H2-1
103	淄麦12	山东	H2	H2-5^a	227	321-134	NWIC-TA	H1	H1-2^a
104	藁9415	河北	H2	H2-1	228	321-170	NWIC-TA	H2	H2-1
105	藁城8901	河北	H2	H2-1	229	321-219	NWIC-TA	H2	H2-5^a
106	邯05-093	河北	H2	H2-1	230	321-244	NWIC-TA	H2	H2-4^a
107	衡08-6008	河北	H2	H2-1	231	321-270	NWIC-TA	H2	H2-5^a
108	衡4371	河北	H2	H2-1	232	321-47	NWIC-TA	H2	H2-5^a
109	衡4422	河北	H2	H2-1	233	321-56	NWIC-TA	H2	H2-4^a
110	衡4568	河北	H2	H2-1	234	321-89	NWIC-TA	H2	H2-2
111	衡5	河北	H2	H2-1	235	322-248	NWIC-TA	H2	H2-4^a
112	衡5364	河北	H2	H2-1	236	322-79	NWIC-TA	H2	H2-1
113	衡562	河北	H2	H2-1	237	323-11	NWIC-TA	H2	H2-1
114	衡6503	河北	H2	H2-1	238	324-315	NWIC-TA	H2	H2-5^a
115	衡6632	河北	H2	H2-2	239	324-316	NWIC-TA	H2	H2-5^a
116	衡矮1号	河北	H2	H2-1	240	324-344	NWIC-TA	H2	H2-4^a
117	衡观35	河北	H2	H2-1	241	324-361	NWIC-TA	H2	H2-4^a
118	衡观76	河北	H2	H2-2	242	324-95	NWIC-TA	H2	H2-5^a
119	冀麦112	河北	H2	H2-7^a	243	325-2	NWIC-TA	H2	H2-4^a
120	科农1006	河北	H2	H2-1	244	325-21	NWIC-TA	H2	H2-1
121	科农199	河北	H2	H2-1	245	325-55	NWIC-TA	H2	H2-4^a
122	科农2009	河北	H2	H2-1	246	329-100	NWIC-TA	H2	H2-2
123	科农3106	河北	H2	H2-2	247	329-20	NWIC-TA	H2	H2-4^a
124	师栾02	河北	H2	H2-2

^a新发现的基因型。NWIC-TA: 国家小麦改良中心泰安分中心。

^a Novel genotypes. NWIC-TA: Tai’an Subcenter of National Wheat Improvement Center.

(续附表) 材料信息及其 Lr10 位点基因型 Appendix Information of materials and their genotypes at locus Lr10

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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[1]	Komer J A. Genetics of resistance to wheat leaf rust. Annu Rev Phytopathol, 1996, 34: 435-455 [本文引用:1] [JCR: 9.875]
[2]	Hu Y-Y(胡亚亚), Zhang N(张娜), Li L-M(李林懋), Yang W-X(杨文香), Liu D-Q(刘大群). Analysis of wheat leaf rust resistance genes in 14 wheat cultivars or lines. Acta Agron Sin(作物学报), 2011, 37(12): 2158-2166 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.8267]
[3]	Feuillet C, Gabriele S, Keller B. Molecular cloning of a new receptor-like kinase gene encoded at the Lr10 disease resistance locus of wheat. Plant J, 1997, 11: 45-52 [本文引用:4] [JCR: 6.16]
[4]	Sela H, SpiridonL N, Petrescu A J, Akerman M, Mand el-Gutfreund Y, Nevo E, Loutre C, Keller B, Schulman A H, Fahima T. Ancient diversity of splicing motifs and protein surfaces in the wild emmer wheat (Triticum dicoccoides) Lr10 coiled coil (CC) and leucine-rich repeat (LRR) domains. Mol Plant Pathol, 2012, 13: 276-287 [本文引用:7] [JCR: 3.899]
[5]	Stein N, Feuillet C, Wicker T, Schlagenhauf E, Keller B. Subgenome chromosome walking in wheat: a 450-kb physical contig in Triticum monococcum L. spans the Lr10 resistance locus in hexaploids wheat (Triticum aestivum L. ). Proc Natl Acad Sci USA, 2000, 97: 13436-13441 [本文引用:5] [JCR: 9.737]
[6]	Isidore E, Scherrer B, Chalhoub B, Feuillet C, Keller B. Ancient haplotypes resulting from extensive molecular rearrangements in the wheat A genome have been maintained in species of three different ploidy levels. Genome Res, 2005, 15: 526-536 [本文引用:11] [JCR: 13.608]
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[8]	Scherrer B, Keller B, Feuillet C. Two haplotypes of resistance gene analogs have been conserved during evolution at the leaf rust resistance locus Lr10 in wild and cultivated wheat. Funct Integrat Genomic, 2002, 2: 40-50 [本文引用:4] [JCR: 2.842]
[9]	Feuillet C, Travella S, Stein N, Albar L, Nublat A, Keller B. Map-based isolation of the leaf rust disease resistance gene Lr10 from the hexaploid wheat (Triticum aestivum L. ) genome. Proc Natl Acad Sci USA, 2003, 100: 15253-15258 [本文引用:3] [JCR: 9.737]
[10]	Cody R. Learning SAS by Example: A Programmer’s Guide, SAS Institute Inc. , Cary, NC, USA, 2007 [本文引用:1]
[11]	Nalam V J, Vales M I, Watson C J W, Kianian S F, Riera-Lizarazu O. Map-based analysis of genes affecting the brittle rachis character in tetraploid wheat (Triticum turgidum L. ). Theor Appl Genet, 2006, 112: 373-381 [本文引用:1] [JCR: 3.297]
[12]	Dubcovsky J, Dvorak J. Genome plasticity a key factor in the success of polyploid wheat under domestication. Science, 2007, 316: 1862-1866 [本文引用:1]
[13]	Loutre C, Wicker T, Travella S, Galli P, Scofield S, Fahima T, Feuillt C, Keller B. Two different CC-NBS-LRR genes are required for Lr10-mediated leaf rust resistance in tetrsploid and hexaploid wheat. Plant J, 2009, 60: 1043-1054 [本文引用:2] [JCR: 6.16]
[14]	Chen W-Q(陈万权), Wang J-X(王剑雄). Genes for leaf and stem rust resistance in 76 wheat genetic resources. Acta Agron Sin(作物学报), 1997, 23(6): 655-663 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.8267]
[15]	Stahl E A, Dwyer G, Mauricio R, Kreitman M, Bergelson J. Dynamics of disease resistance polymorphism at the Rpm1 locus of Arabidopsis. Nature, 1999, 400: 667-671 [本文引用:1] [JCR: 36.28]
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1996

9.875

0.0

Annu Rev Phytopathol. 1996, 34:435-55

Genetics of resistance to wheat leaf rust.

Kolmer JA

Agriculture and Agri-Food Canada, Cereal Research Centre, 195 Dafoe Road, Winnipeg, Manitoba R3T 2M9, Canada.

Leaf rust (caused by Puccinia recondita f. sp. tritici) is the most widespread and regularly occurring rust on wheat. Genetic resistance is the most economical method of reducing yield losses due to leaf rust. To date, 46 leaf rust resistance genes have been designated and mapped in wheat. Resistance gene expression is dependent on the genetics of host-parasite interaction, temperature conditions, plant developmental stage, and interaction between resistance genes with suppressors or other resistance genes in the wheat genomes. Genes expressed in seedling plants have not provided long-lasting effective leaf rust resistance. Adult-plant resistance genes Lr13 and Lr34 singly and together have provided the most durable resistance to leaf rust in wheat throughout the world. Continued efforts to isolate, characterize, and map leaf rust resistance genes is essential given the ability of the leaf rust fungus to overcome deployed resistance genes.

PMID: 15012551

... 叶锈病是小麦的主要病害之一, 发病严重时可减产50%^[1] ...

2011

0.0

1.8267

Acta Agron Sin. 2011, 37(12):2158 - 2166 DOI:10.3724/SP.J.1006.2011.02158

Analysis of Wheat Leaf Rust Resistance Genes in 14 Wheat Cultivars or Lines

14个小麦品种(系)抗叶锈性分析

HU Ya-Ya, ZHANG Na, LI Lin-Mao, YANG Wen-Xiang^*,LIU Da-Qun^*

胡亚亚,张娜,李林懋,杨文香^*,刘大群^*

Department of Plant Pathology, Agricultural University of Hebei / Biological Control Center of Plant Diseases and Plant Pests of Hebei Province / National Engineering Research Center for Agriculture in Northern Mountainous Areas, Baoding 071001, China

The objective of this study was todetect resistance genes to leaf rust in 14 wheat cultivars or lines. The resistance of the 14 wheat cultivars or lines wasinvestigated at seedling and adult stages, and the resistance genes at seedling stage were postulated byinoculating 16 races ofPuccinia triticina. Furthervalidation was conducted by using 21 molecular markers cosegregated with or closely linked to the knownLr genes. According to the results of bothphenotypic identification and molecular marker data, the information of the resistance genes in the 14 cultivars or lines was obtained. GenesLr3a, Lr28 andLr50 were postulated in s98351-2-2-2-1. GeneLr37 was possibly carried by 9629-03A-4-1-1. Lr26 was present in 97167-1-2-1-1-2-1, 919-20-2c2, 9589, Mianzhong 438, 9916-8-6, and 9916-8-18.Lr28 was possibly present in 96104-1-5-1c2.Lr1 was contained in 00-55-3-1-1.Lr24,Lr37, andLr38were possibly in1R13.Lr24andLr38 were possibly carried by 1R17. GeneLr10 andLr34 were detected in 1R35, andLr3a andLr50 were also postulated in 1R35. Besides, 9524-1-2-2-1 might contain unknown or untested resistance genes againstP. triticina pathotypes. GenesLr9,Lr19,Lr20, Lr21, Lr29, Lr35, andLr42 were not present in all cultivars or lines. The results indicate that these cultivars and lines carry diverse resistance genes to wheat leaf rust, and can be used as resistance resource for resistance breeding.

选用16个小麦叶锈菌菌系对14个小麦品种(系)进行抗叶锈性鉴定和苗期抗叶锈基因推导，初步分析这些品种(系)的抗性和携带的抗病基因；进一步利用21个与Lr基因紧密连锁或共分离的分子标记，对这14个品种(系)中可能含的抗叶锈基因进行鉴定。结果表明，s98351-2-2-2-1可能含Lr3a、Lr28和Lr50；9629-03A-4-1-1可能含Lr37；97167-1-2-1-1-2-1、919-20-2c2、9589、免中438、9916-8-6和9916-8-18含Lr26；96104-1-5-1c2可能含Lr28；00-55-3-1-1含Lr1；1R13可能含Lr24、Lr37和Lr38；1R17可能含Lr24和Lr38；1R35含Lr10和Lr34，还可能含Lr3a和Lr50；9524-1-2-2-1含未知抗叶锈基因或本试验使用的已知抗病基因以外的抗叶锈基因。所有品种(系)均不含Lr9、Lr19、Lr20、Lr21、Lr29、Lr35、Lr42和Lr47基因。测试的14个品种(系)中有比较丰富的抗叶锈病基因，可为育种提供丰富的抗源。

... 小麦叶锈病的病原菌为Puccinia triticina , 具有小种专化性, 对不同小麦品种具有不同的致病性^[2] ...

... 上述研究结果表明, Lr10 基因经历了强烈的瓶颈效应, 推测导致这种结果的原因有2个: 其一, 当前流行的叶锈病菌已经对Lr10 基因产生抗性, 因而, 小麦抗病育种工作者已不选育含有此基因的材料^[2] ...

1997

6.16

0.0

... 抗叶锈基因Lr10 位于小麦1A染色体短臂上^[3,4],与另一抗病基因同源序列RGA2 紧密连锁, 在该位点存在H1和H2两种古单倍型^[3] ...

... H1古单倍型含有完整的Lr10 基因及与其紧密连锁的RGA2 基因, H2型仅含有RGA2 的部分序列, 缺失RGA2 的5′端部分序列及全部Lr10 序列^[3,4,5,6] ...

... dicoccoides )中^[3,4,5,6] ...

2012

3.899

0.0

Mol. Plant Pathol.. 2012, 13(3):276-87 DOI:10.1111/j.1364-3703.2011.00744.x

Ancient diversity of splicing motifs and protein surfaces in the wild emmer wheat (Triticum dicoccoides) LR10 coiled coil (CC) and leucine-rich repeat (LRR) domains.

Sela H, Spiridon LN, Petrescu AJ, Akerman M, Mandel-Gutfreund Y, Nevo E, Loutre C, Keller B, Schulman AH, Fahima T

Department of Evolutionary and Environmental Biology, Institute of Evolution, University of Haifa, Mt. Carmel, Haifa 31905, Israel.

In this study, we explore the diversity and its distribution along the wheat leaf rust resistance protein LR10 three-dimensional structure. Lr10 is a leaf rust resistance gene encoding a coiled coil-nucleotide-binding site-leucine-rich repeat (CC-NBS-LRR) class of protein. Lr10 was cloned and sequenced from 58 accessions representing diverse habitats of wild emmer wheat in Israel. Nucleotide diversity was very high relative to other wild emmer wheat genes (π= 0.029). The CC domain was found to be the most diverse domain and subject to positive selection. Superimposition of the diversity on the CC three-dimensional structure showed that some of the variable and positively selected residues were solvent exposed and may interact with other proteins. The LRR domain was relatively conserved, but showed a hotspot of amino acid variation between two haplotypes in the ninth repeat. This repeat was longer than the other LRRs, and three-dimensional modelling suggested that an extensive α helix structure was formed in this region. The two haplotypes also differed in splicing regulation motifs. In genotypes with one haplotype, an intron was alternatively spliced in this region, whereas, in genotypes with the other haplotype, this intron did not splice at all. The two haplotypes are proposed to be ancient and maintained by balancing selection.

PMID: 21952112

... 抗叶锈基因Lr10 位于小麦1A染色体短臂上^[3,4],与另一抗病基因同源序列RGA2 紧密连锁, 在该位点存在H1和H2两种古单倍型^[3] ...

... H1古单倍型含有完整的Lr10 基因及与其紧密连锁的RGA2 基因, H2型仅含有RGA2 的部分序列, 缺失RGA2 的5′端部分序列及全部Lr10 序列^[3,4,5,6] ...

... dicoccoides )中^[3,4,5,6] ...

... Lr10 基因是小麦中4个已测序抗叶锈病基因之一^[4](http://www ...

... gov/genbank/), 全长为3935 bp, 与其他抗叶锈病基因序列的相似度较低^[4] ...

... 其二, Lr10 基因可能对某些稀有致病小种或未知种有抗性, 但对当前流行的叶锈病病原菌没有效果, 且该基因的表达对产量或其他农艺性状有负面效应, 最终导致育种工作者不选育含有此基因的材料^[4] ...

... Lr10 基因的进化模式为trench-warfare模型, 在野生群体中具有较高的遗传多样性^[4,20] ...

2000

9.737

0.0

... H1古单倍型含有完整的Lr10 基因及与其紧密连锁的RGA2 基因, H2型仅含有RGA2 的部分序列, 缺失RGA2 的5′端部分序列及全部Lr10 序列^[3,4,5,6] ...

... dicoccoides )中^[3,4,5,6] ...

... 在小麦进化和选育过程中, Lr10 基因在普通小麦中被保留下来, 并对叶锈菌菌株(AvrLr10 ) 89-201 CBTB(TX)表现抗性^[5] ...

... Stein等^[5]检测了Lr10 古单倍型在56份欧洲小麦品系中的分布规律, 发现H1古单倍型的比例为14 ...

... Stein等^[5]和Isidore等^[6]的研究也表明, 小麦的抗病基因同样经历了类似的遗传瓶颈效应, 但这些抗病基因经历瓶颈效应的程度和可能获得的新基因并没有得到深入研究 ...

2005

13.608

0.0

Genome Res.. 2005, 15(4):526-36

Ancient haplotypes resulting from extensive molecular rearrangements in the wheat A genome have been maintained in species of three different ploidy levels.

Isidore E, Scherrer B, Chalhoub B, Feuillet C, Keller B

Institute of Plant Biology, University of Zürich, 8008 Zürich, Switzerland.

Plant genomes, in particular grass genomes, evolve very rapidly. The closely related A genomes of diploid, tetraploid, and hexaploid wheat are derived from a common ancestor that lived<3 million years ago and represent a good model to study molecular mechanisms involved in such rapid evolution. We have sequenced and compared physical contigs at the Lr10 locus on chromosome 1AS from diploid (211 kb), tetraploid (187 kb), and hexaploid wheat (154 kb). A maximum of 33% of the sequences were conserved between two species. The sequences from diploid and tetraploid wheat shared all of the genes, including Lr10 and RGA2 and define a first haplotype (H1). The 130-kb intergenic region between Lr10 and RGA2 was conserved in size despite its activity as a hot spot for transposon insertion, which resulted in >70% of sequence divergence. The hexaploid wheat sequence lacks both Lr10 and RGA2 genes and defines a second haplotype, H2, which originated from ancient and extensive rearrangements. These rearrangements included insertions of retroelements and transposons deletions, as well as unequal recombination within elements. Gene disruption in haplotype H2 was caused by a deletion and subsequent large inversion. Gene conservation between H1 haplotypes, as well as conservation of rearrangements at the origin of the H2 haplotype at three different ploidy levels indicate that the two haplotypes are ancient and had a stable gene content during evolution, whereas the intergenic regions evolved rapidly. Polyploidization during wheat evolution had no detectable consequences on the structure and evolution of the two haplotypes.

PMID: 15805493

... H1古单倍型含有完整的Lr10 基因及与其紧密连锁的RGA2 基因, H2型仅含有RGA2 的部分序列, 缺失RGA2 的5′端部分序列及全部Lr10 序列^[3,4,5,6] ...

... dicoccoides )中^[3,4,5,6] ...

... Isidore等^[6]设计了4对H2单倍型的特异引物(A、B、C和B_3kb)用于研究不同倍性小麦中H2单倍型的保守性, 共检测出3种H2亚型, 在欧洲普通小麦中仅检测出2种H2亚型 ...

... 采用引物Control^[6]和rga1Pro^[8], 通过PCR扩增检测H1和H2单倍型, 其中引物Control (表1)用于扩增H1和H2两种单倍型的共同序列(图1), 引物rga1Pro (表1)特异性扩增Lr10 基因的5′端非编码区及部分5′端编码区, 只在H1型个体中有扩增产物(图1) ...

... 而引物A、B、C和B_3k^[6]用于检测H2的亚型 ...

... 在T-gradient Thermal Cycler (Bio-metra)上, 按已报道的方法^[6,8,9]进行PCR扩增 ...

... Isidore等^[6]发现, 欧洲小麦中Lr10 位点的H2-1型占优势 ...

... Isidore等^[6]推测逆转录子的插入、转座子的缺失以及Lr10 基因与RGA2 基因间元件的不等重组等事件形成H2古单倍型 ...

... Isidore等^[6]还推测H2和H1古单倍型之间非常稳定, 不会发生重组 ...

... 本研究的结果支持Sela等^[20]的结论, 即H2和H1古单倍型确实存在于小麦品种中, 而且H2-1和H2-2基因型分布广泛并相对稳定, 同时又修正了Isidore等^[6]关于重组不能发生在H2和H1古单倍型之间的推断 ...

1990

0.0

... 采用Doyle和Doyle^[7]的方法从小麦幼苗提取的基因组DNA ...

2002

2.842

0.0

Funct Integr Genomic. 2002, 2(1-2):40 - 50 DOI:10.1007/s10142-002-0051-9

Two haplotypes of resistance gene analogs have been conserved during evolution at the leaf rust resistance locusLr10 in wild and cultivated wheat

B. Scherrer(1) B. Keller(1) C. Feuillet(1)

1.Institute of Plant Biology, University of Zürich, Zollikerstrasse 107, 8008 Zürich, Switzerland

The isolation of genes of agronomic interest such as disease resistance genes is a central issue in wheat research. A good knowledge of the organization and evolution of the genome can greatly help in defining the best strategies for efficient gene isolation. So far, very few wheat disease resistance loci have been studied at the molecular level and little is known about their evolution during polyploidization and domestication. In this study, we have analyzed the haplotype structure at loci orthologous to the leaf rust resistance locusLr10in hexaploid wheat which spans 350 kb in diploid wheat. Two haplotypes (H1, H2) were defined by the presence (H1) or the absence (H2) of two different resistance gene analogs (rga1, rga2) at this locus on chromosome 1AS. Both haplotypes were found in a collection of 113 wild and cultivated diploid and polyploid wheat lines and they do not reflect phylogenetic relationships. This indicates an ancient origin for this disease resistance locus and the independent conservation of the two haplotypes throughout the evolution of the wheat genome. Finally, the coding regions of the H1 haplotype RGAs are extremely conserved in all the species. This suggests a selective pressure for maintaining the structural and functional configuration of this haplotype in wheat. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s10142-002-0051-9.

... 在T-gradient Thermal Cycler (Bio-metra)上, 按已报道的方法^[6,8,9]进行PCR扩增 ...

... 例如, 若两对引物Control和rga1Pro都有PCR产物, 则其古单倍型的类型为H1^[8] ...

... 若仅引物Control有PCR产物, 引物rga1Pro无扩增产物, 则其古单倍型的类型为H2^[8] ...

2003

9.737

0.0

Proc. Natl. Acad. Sci. U.S.A.. 2003, 100(25):15253-8

Map-based isolation of the leaf rust disease resistance gene Lr10 from the hexaploid wheat (Triticum aestivum L.) genome.

Feuillet C, Travella S, Stein N, Albar L, Nublat A, Keller B

Institute of Plant Biology, University of Zürich, Zollikerstrasse 107, CH-8008 Zürich, Switzerland.

More than 50 leaf rust resistance (Lr) genes against the fungal pathogen Puccinia triticina have been identified in the wheat gene pool, and a large number of them have been extensively used in breeding. Of the 50 Lr genes, all are known only from their phenotype and/or map position except for Lr21, which was cloned recently. For many years, the problems of molecular work in the large (1.6 x 10(10) bp), highly repetitive (80%), and hexaploid bread wheat (Triticum aestivum L.) genome have hampered map-based cloning. Here, we report the isolation of the Lr gene Lr10 from hexaploid wheat by using a combination of subgenome map-based cloning and haplotype studies in the genus Triticum. Lr10 is a single-copy gene on chromosome 1AS. It encodes a CC-NBS-LRR type of protein with an N-terminal domain, which is under diversifying selection. When overexpressed in transgenic wheat plants, Lr10 confers enhanced resistance to leaf rust. Lr10 has similarities to RPM1 in Arabidopsis thaliana and to resistance gene analogs in rice and barley, but is not closely related to other wheat Lr genes based on Southern analysis. We conclude that map-based cloning of genes of agronomic importance in hexaploid wheat is now feasible, opening perspectives for molecular bread wheat improvement trough transgenic strategies and diagnostic allele detection.

PMID: 14645721

... 引物ThLr10_T/P、ThLr10_E/H和ThLr10_G/J^[9]用于检测H1的亚型 ...

... 在T-gradient Thermal Cycler (Bio-metra)上, 按已报道的方法^[6,8,9]进行PCR扩增 ...

... 该基因编码919个氨基酸长度的CC-NBS-LRR类型蛋白质^[9], 在蛋白CC结构域的N端经历多样性选择(diversifying selection)^[13] ...

2007

0.0

... 0软件^[10]统计各古单倍型及其亚型的发生频率以及基因型 ...

2006

3.297

0.0

Theor Appl Genet. 2006, 112(2):373 - 381 DOI:10.1007/s00122-005-0140-y

Map-based analysis of genes affecting the brittle rachis character in tetraploid wheat (Triticum turgidum L.)

Vamsi J. Nalam(13) M. Isabel Vales(1) Christy J. W. Watson(1) Shahryar F. Kianian(2) Oscar Riera-Lizarazu(1)

1.Department of Crop and Soil Science, Oregon State University, 107 Crop Science Bldg, Corvallis, OR 97331, USA
2.Department of Plant Sciences, North Dakota State University, Fargo, ND 58105, USA
3.Department of Plant Pathology, Kansas State University, Manhattan, KS 66506, USA

The mature spike rachis of wild emmer [Triticum turgidum L. ssp.dicoccoides (K?rn. ex Asch. and Graebner) Thell.] disarticulates spontaneously between each spikelet leading to the dispersion of wedge-type diaspores. By contrast, the spike rachis of domesticated emmer (Triticum turgidum L. ssp.turgidum) fails to disarticulate and remains intact until it is harvested. This major distinguishing feature between wild and domesticated emmer is controlled by two major genes,brittle rachis 2 (Br-A2) andbrittle rachis 3 (Br-A3) on the short arms of chromosomes 3A and 3B, respectively. Because of their biological and agricultural importance, a map-based analysis of these genes was undertaken. Using two recombinant inbred chromosome line (RICL) populations,Br-A2, on chromosome 3A, was localized to a ~11-cM region betweenXgwm2 and a cluster of linked loci (Xgwm666.1, Xbarc19, Xcfa2164, Xbarc356, andXgwm674), whereasBr-A3, on chromosome 3B, was localized to a ~24-cM interval betweenXbarc218 andXwmc777. Comparative mapping analyses suggested that bothBr-A2 andBr-A3 were present in homoeologous regions on chromosomes 3A and 3B, respectively. Furthermore,Br-A2 andBr-A3 from wheat andBtr1/Btr2 on chromosome 3H of barley (Hordeum vulgare L.) also were homoeologous suggesting that the location of major determinants of the brittle rachis trait in these species has been conserved. On the other hand, brittle rachis loci of wheat and barley, and a shattering locus on rice chromosome 1 did not appear to be orthologous. Linkage and deletion-based bin mapping comparisons suggested thatBr-A2 andBr-A3 may reside in chromosomal areas where the estimated frequency of recombination was ~ 4.3 Mb/cM. These estimates indicated that the cloning ofBr-A2 andBr-A3 using map-based methods would be extremely challenging.

... 1 Lr10 基因位点的遗传瓶颈效应长期的自然选择和人工驯化最终将导致小麦基因的遗传多样性降低^[11] ...

2007

0.0

... Dubcovsky和Dvorak^[12]指出, 小麦可以通过其A、B和D基因组的基因补偿效应来弥补基因瓶颈效应造成的不良后果, 杂交育种可以使新育成的小麦获得优于亲本的基因型, 杂交后代通过基因突变、重组、缺失等方式也可以形成全新的基因型 ...

2009

6.16

0.0

Plant J.. 2009, 60(6):1043-54 DOI:10.1111/j.1365-313X.2009.04024.x

Two different CC-NBS-LRR genes are required for Lr10-mediated leaf rust resistance in tetraploid and hexaploid wheat.

Loutre C, Wicker T, Travella S, Galli P, Scofield S, Fahima T, Feuillet C, Keller B

Institute of Plant Biology, University of Zurich, Zollikerstrasse 107, 8008 Zürich, Switzerland.

Comparative study of disease resistance genes in crop plants and their relatives provides insight on resistance gene function, evolution and diversity. Here, we studied the allelic diversity of the Lr10 leaf rust resistance gene, a CC-NBS-LRR coding gene originally isolated from hexaploid wheat, in 20 diploid and tetraploid wheat lines. Besides a gene in the tetraploid wheat variety 'Altar' that is identical to the hexaploid wheat Lr10, two additional, functional resistance alleles showing sequence diversity were identified by virus-induced gene silencing in tetraploid wheat lines. In contrast to most described NBS-LRR proteins, the N-terminal CC domain of LR10 was found to be under strong diversifying selection. A second NBS-LRR gene at the Lr10 locus, RGA2, was shown through silencing to be essential for Lr10 function. Interestingly, RGA2 showed much less sequence diversity than Lr10. These data demonstrate allelic diversity of functional genes at the Lr10 locus in tetraploid wheat, and these new genes can now be analyzed for agronomic relevance. Lr10-based resistance is highly unusual both in its dependence on two, only distantly, related CC-NBS-LRR proteins, as well as in the pattern of diversifying selection in the N-terminal domain. This indicates a new and complex molecular mechanism of pathogen detection and signal transduction.

PMID: 19769576

... 该基因编码919个氨基酸长度的CC-NBS-LRR类型蛋白质^[9], 在蛋白CC结构域的N端经历多样性选择(diversifying selection)^[13] ...

... 另一抗病基因同源序列RGA2 与Lr10 基因紧密连锁, 是Lr10 基因表现出抗性必需的调控基因^[13] ...

1997

0.0

1.8267

Acta Agron Sin. 1997, 23(6):655 - 663

Genes for Leaf and Stem Rust Resistance in 76 Wheat Genetic Resources

76个小麦种质资源抗叶锈及秆锈基因初步分析

Chen Wanquan; Wang Jianxiong

陈万权;王剑雄

Institute of Plant Protection, CAAS, Beijing, 100094

... 陈万权和王剑雄^[14]采用基因推导法在来自世界各地的76个小麦种质资源中鉴定出4个材料含Lr10 基因 ...

1999

36.28

0.0

... 丁艳红等^[15]采用STS分子标记在28个小麦微核心种质中鉴定出2个材料含Lr10 基因 ...

2001

38.075

0.0

... 2 Lr10 基因位点的进化致病菌的高度变异为植物抗病基因的进化提供了很强的自然选择压, 关于两者的相关性主要有2种假说, 即trench-warfare模型^[16]和arm-race模型^[17] ...

... 前者认为抗病基因进化形成的多样性依赖于致病菌的变异频率, 频率高时抗病基因的多样性高, 频率低时抗病基因的多样性低, 在此模型下, 抗病基因的多样性较高, 可以形成新基因和保留已有基因, 如拟南芥中的基因Rpm1^[16]、RPP8^[17]、RPP13 等^[17] ...

1998

13.608

0.0

2010

3.899

0.0

Mol. Plant Pathol.. 2010, 11(2):309-14 DOI:10.1111/j.1364-3703.2009.00605.x

The arms race between tomato and Fusarium oxysporum.

Takken F, Rep M

Plant Pathology, Swammerdam Institute for Life Sciences, University of Amsterdam, P.O. Box 94215, 1090 GE Amsterdam, the Netherlands. f.l.w.takken@uva.nl

The interaction between tomato and Fusarium oxysporum f. sp. lycopersici has become a model system for the study of the molecular basis of disease resistance and susceptibility. Gene-for-gene interactions in this system have provided the basis for the development of tomato cultivars resistant to Fusarium wilt disease. Over the last 6 years, new insights into the molecular basis of these gene-for-gene interactions have been obtained. Highlights are the identification of three avirulence genes in F. oxysporum f. sp. lycopersici and the development of a molecular switch model for I-2, a nucleotide-binding and leucine-rich repeat-type resistance protein which mediates the recognition of the Avr2 protein. We summarize these findings here and present possible scenarios for the ongoing molecular arms race between tomato and F. oxysporum f. sp. lycopersici in both nature and agriculture.

PMID: 20447279

... 后者是birth-death模型^[18]的补充完善, 强调抗病基因的多样性有明显的方向性, 即选择更具抗性的基因, 在此模型下, 抗病基因的多样性很低, 有明显的方向性, 如番茄中对尖孢镰孢菌(Fusarium oxysporum )产生抗性的抗病基因^[19] ...

2011

3.297

0.0

Theor. Appl. Genet.. 2011, 122(1):175-87 DOI:10.1007/s00122-010-1434-2

Rapid linkage disequilibrium decay in the Lr10 gene in wild emmer wheat (Triticum dicoccoides) populations.

Sela H, Loutre C, Keller B, Schulman A, Nevo E, Korol A, Fahima T

Department of Evolutionary and Environmental Biology, Institute of Evolution, Faculty of Natural Sciences, University of Haifa, Mt. Carmel, Haifa, 31905, Israel.

Recombination is a key evolutionary factor enhancing diversity. However, the effect of recombination on diversity in inbreeding species is expected to be low. To estimate this effect, recombination and diversity patterns of Lr10 gene were studied in natural populations of the inbreeder species, wild emmer wheat (Triticum dicoccoides). Wild emmer wheat is the progenitor of most cultivated wheats and it harbors rich genetic resources for disease resistance. Lr10 is a leaf rust resistance gene encoding three domains: a coiled-coil, nucleotide-binding site, and leucine-rich repeat (CC-NBS-LRR).

PMID: 20859611