小麦慢白粉病QTL对条锈病和叶锈病的兼抗性

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刘金栋, 陈新民, 何中虎, 伍玲, 白斌, 李在峰, 夏先春. 小麦慢白粉病QTL对条锈病和叶锈病的兼抗性. 作物学报, 2014, 40(9): 1557-1564[LIU Jin-Dong, CHEN Xin-Min, HE Zhong-Hu, WU Ling, BAI Bin, LI Zai-Feng, XIA Xian-Chun. Resistance of Slow Mildewing Genes to Stripe Rust and Leaf Rust in Common Wheat. Acta Agronomica Sinica, 2014, 40(9): 1557-1564] 复制到剪切板

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作物学报编辑部

小麦慢白粉病QTL对条锈病和叶锈病的兼抗性

刘金栋¹, 陈新民¹, 何中虎^1,², 伍玲³, 白斌⁴, 李在峰⁵, 夏先春^1,^*

¹中国农业科学院作物科学研究所/国家小麦改良中心, 北京100081

²CIMMYT中国办事处, 北京100081

³四川省农业科学院作物研究所, 四川成都 610066

⁴甘肃省农业科学院小麦研究所, 甘肃兰州730070

⁵河北农业大学植物保护学院植物病理系, 河北保定071001

^*通讯作者(Corresponding author): 夏先春, E-mail:xiaxianchun@caas.cn, Tel: 010-82108610

收稿日期:2014-06-16

基金:本研究由国家重点基础研究发展计划项目(2013CB127700), 国家自然科学基金项目(31261140370), 国家高新技术研究发展计划(863计划)项目(2012AA101105)和国家现代农业产业技术体系建设专项(CARS-3-1-3)资助;

摘要

聚合兼抗白粉病、条锈病和叶锈病的慢病性基因, 是培育持久多抗小麦品种的重要措施。百农64和鲁麦21均为慢白粉病品种, 分别含有4个和3个慢白粉病抗性QTL。将百农64与鲁麦21杂交, 获得21个聚合2~5个慢白粉病抗性QTL的F₆株系, 于2012—2013年度分别在四川郫县和甘肃天水进行条锈病田间抗性鉴定,在河北保定和河南周口进行叶锈病田间抗性鉴定。分析21个株系条锈和叶锈病的最大严重度和病程曲线下面积, 检测单个QTL和QTL聚合体对条锈病和叶锈病的抗性效应。结果表明,QPm.caas-4DL、QPm.caas-6BS和QPm.caas-2BL对条锈病均有显著的抗性, 分别解释表型变异的16.9%、14.1%和17.3%;QPm.caas-4DL对叶锈病也有显著抗性, 可解释表型变异的35.3%;QPm.caas-1A/QPm.caas-4DL/QPm.caas-2DL/QPm.caas-2BS/QPm.caas-2BL和QPm.caas-1A/QPm.caas- 4DL/QPm.caas-2BS/QPm.caas-2BL聚合体对条锈病和叶锈病的抗性显著高于两亲本, 它们均含有来自百农64的QPm.caas-4DL以及来自鲁麦21的QPm.caas-2BL和QPm.caas-2BS, 表明这些QTL具有明显的兼抗性效应。在小麦抗病育种中, 聚合慢病性QTL越多, 慢病性越强, 聚合4~5个慢病性QTL时, 株系可达到高抗甚至接近免疫的水平, 是选育持久抗性小麦品种的重要手段。

关键词: 普通小麦; 慢病性; 持久抗性; 基因聚合; QTL

Resistance of Slow Mildewing Genes to Stripe Rust and Leaf Rust in Common Wheat

LIU Jin-Dong¹, CHEN Xin-Min¹, HE Zhong-Hu^1,², WU Ling³, BAI Bin⁴, LI Zai-Feng⁵, XIA Xian-Chun^1,^*

¹Institute of Crop Science / National Wheat Improvement Center, Chinese Academy of Agricultural Sciences (CAAS), Beijing 100081, China

²CIMMYT-China Office, c/o CAAS, Beijing 100081, China

³Crop Research Institute, Sichuan Academy of Agricultural Sciences, Chengdu 610066, China

⁴Wheat Research Institute, Gansu Academy of Agricultural Sciences, Lanzhou 730070, China

⁵Department of Plant Pathology, College of Plant Protection, Agricultural University of Hebei, Baoding 071001, China

Fund:

Abstract

Pyramiding quantitative trait loci (QTLs) is an effective method to improve resistance to powdery mildew, stripe rust, and leaf rust in common wheat. We have developed 21 lines (F₆) carrying 2-5 slow mildewing QTLs by crossing slow powdery mildew cultivars Bainong 64 and Lumai 21 possessing four and three slow mildewing QTLs, respectively. These F₆lines were evaluated in the field in Pianxian, Sichuan and Tianshui, Gansu for stripe rust resistance and in Baoding, Hebei and Zhoukou, Henan for leaf rust resistance during the 2012-2013 cropping season. According to the maximum disease severities (MDS) and the area under the disease progress curve (AUDPC), QTLsQPm.caas-4DL,QPm.caas-6BS, andQPm.caas-2BLwere highly resistant to stripe rust (P < 0.01), which explained 16.9%, 14.1%, and 17.3% of phenotypic variance, respectively. LocusQPm.caas-4DLalso showed high resistance to leaf rust (P < 0.01) with phenotypic contribution of 35.3%. Lines that pyramided five (QPm.caas-1A/QPm.caas-4DL/QPm.caas-2DL/QPm.caas-2BS/QPm.caas-2BL) and four (QPm.caas-1A/QPm.caas-4DL/QPm. caas-2BS/QPm.caas-2BL) QTLs exhibited higher resistance to both stripe and leaf rust compared with their parents. This result indicates that the combination ofQPm.caas-4DL (from Bainong 64),QPm.caas-2BS andQPm.caas-2BL(Lumai 21) has a marked effect on improving adult resistance to powdery mildew, stripe rust and leaf rust, and the more QTLs are pyramided, the stronger slow disease resistance can be achieved. In breeding practice, the combination of 4-5 slow mildewing or rusting QTLs can result in durable resistance to multiple diseases.

Keyword: Triticum aestivum L.; Slow mildewing and slow rusting resistance; Durable resistance; Gene pyramiding; QTL

Show Figures

小麦白粉病、条锈病和叶锈病是全球性重要病害, 分别由小麦白粉菌( Blumeria graminisf. sp. tritici)、小麦条锈菌( Puccinia striiformis f. sp. tritici)和小麦叶锈菌( P. recondita f. sp. tritici)引起, 具有发生频率高、流行范围广和暴发性强的特点, 可致3.0%~49.0%的产量损失^{[ 1, 2, 3]}。长期以来, 利用寄主抗性防治这些病害取得了显著成效^{[ 4]}。

寄主抗性分为垂直抗性和水平抗性。垂直抗性又称为生理小种专化性抗性、苗期抗性、全生育期抗性或主效基因抗性, 是由1个或少数几个主效基因控制, 对特定病原菌生理小种表现出高抗或免疫, 具有病原菌生理小种专化性, 常因病原菌生理小种变异而丧失抗性; 水平抗性亦称慢病性、成株抗性或非小种专化抗性, 由多个微效基因控制, 对病原菌无小种专化性或专化性弱, 苗期表现为感病, 成株期表现为中抗或高抗, 且抗性持久^{[ 5, 6]}。慢病性由多个微效基因的加性效应控制, 基因聚合是获得慢病性、选育兼抗多种病害品种的重要方式。Singh等^{[ 7]}发现, 聚合4~5个微效慢病基因的小麦材料对条锈和叶锈病呈高抗至免疫。意大利小麦品种Strampelli和Libellula及我国农家种平原50至今仍然高抗条锈病。抗性遗传分析表明, Strampelli和Libellula含有 Yr18/Lr34/Pm38/Sr57和2~4个其他慢病基因^{[ 8]}; 平原50含有3个慢条锈基因和3个慢白粉基因^{[ 9]}。 Yr18/Lr34/Pm38/Sr57对小麦条锈病、叶锈病、白粉病和秆锈病均具有抗性^{[ 10, 11]}。

随着QTL定位和分子标记研究工作的不断深入, 育种家可以利用分子标记聚合抗病QTL, 培育持久抗性品种^{[ 7, 12, 13]}。从20世纪70年代开始, 国际玉米小麦改良中心(CIMMYT)选育出一批抗性持久且兼抗多种病害的慢病性品种, 例如Amadina、Chapio、Cook、Kukuna、Parula、Pavon 76、Sonoita 81、Tonichi 81和Tukuru, 均含有 Yr18/Lr34/Pm38/Sr57或 Yr29/ Lr46/Pm39/Sr58位点及2~3个微效基因^{[ 14, 15]}。因此, 聚合慢病性基因是获得兼抗多种病害的持久抗性小麦品种的重要手段。

虽然小麦抗病性QTL定位研究很多, 但是由于不同遗传背景下的QTL聚合到同一遗传背景存在诸多困难^{[ 16]}, 不同来源的QTL可能存在互作, QTL聚合育种实践的报道很少。Miedaner等^{[ 17]}将3个来源不同的赤霉病抗性QTL聚合到同一种质, 发现单个QTL均可降低赤霉病发病率和脱氧雪腐镰刀菌烯醇(DON)含量, 但3A上的一个QTL单独存在或与3B和5A上QTL聚合时对赤霉病没有显著遗传效应。Lu等^{[ 18]}将5A上的QTL与 Fhb1聚合在一起, 发现可以降低矮秆基因 Rht-D1b对赤霉病引起的效应。可见, QTL聚合体遗传效应分析具有重要的应用价值。

百农64和鲁麦21均具有优良的农艺性状, 分别在20世纪90年代和21世纪初为我国黄淮麦区主推品种, 其苗期对白粉菌^{[ 19]}、条锈菌和叶锈菌^{[ 20]}流行小种均感病, 但田间成株期表现抗病, 具有典型的慢病性特征。在百农64中检测到4个慢白粉病抗性QTL ( QPm.caas-1A、 QPm.caas-4DL、 QPm.caas- 6BS和 QPm.caas-7A)^{[ 21]}, 在鲁麦21中检测到3个慢白粉病抗性QTL ( QPm.caas-2BS、 QPm.caas-2BL和 QPm.caas-2DL)^{[ 22]}。Bai等^{[ 23]}采用杂交和改良系谱法, 对百农64和鲁麦21所含慢白粉病抗性QTL进行聚合, 创制了21个F₆聚合株系, 并对这些材料的白粉病抗性进行了分析。在此基础上, 本研究将分析这21个聚合不同慢白粉病基因的株系对条锈病和叶锈病的遗传效应, 明确这些慢白粉病抗性QTL及QTL聚合体对条锈病和叶锈病的抗性, 发掘高抗条锈、叶锈和白粉病的慢病性材料和QTL组合, 为选育兼抗多种病害的持久抗性小麦品种提供材料和方法。

1 材料与方法

1.1 聚合株系基因型及其分子标记

百农64与鲁麦21杂交, 采用改良系谱法, 根据田间白粉病抗性和综合农艺性状选择, 在早代淘汰白粉病严重度高的株系, 选择低严重度株系继续种植, 经过连续多代鉴定和选择, 获得21个F₆抗病株系(编号: BFB5~BFB25)^{[ 23]}; 利用16个分子标记^{[ 21, 22]}进行检测, 明确这些株系携带2~5个慢白粉病QTL。这些慢白粉病QTL中, QPm.caas-1A的连锁标记为 Xbarc148和 Xwmc550, QPm.caas-4DL的连锁标记为 Xgwm165、 Xcfd23和 Xwmc331, QPm.caas-6BS的连锁标记为 Xbarc79和 Xgwm518, QPm.caas-7A的连锁标记为 Xbarc127和 Xbarc174, QPm.caas-2BS的连锁为 Xbarc98和 Xbarc1147, QPm.caas2BL的连锁标记为 Xbarc1139和 Xgwm47, QPm.caas-2DL的连锁标记为 Xbarc18、 Xgwm539和 Xcfd233。

1.2 田间设计

将21个F₆聚合基因株系及其亲本于2012—2013年度种植于四川郫县(30°05′N, 102°54′E)和甘肃天水(34°05′N, 104°35′E)进行条锈病抗性鉴定; 在河北保定(113°40′N, 38°10′E)和河南周口(114°38′N, 33°37′E)进行叶锈病抗性鉴定。采用完全随机区组设计, 3次重复, 单行区, 行长1.50 m, 行距0.25 m, 每行种植50粒。在郫县和天水点, 每10行种植1行高感条锈病品种对照辉县红, 小区周围种植高感品种作为诱发行; 在保定和周口, 每10行种植1行高感叶锈病对照郑州5389。以保证充分接种, 实际观察所有对照均发病完全。

1.3 抗病性评价

接种后6周左右, 当对照品种充分发病时开始调查发病严重度, 记录旗叶和倒二叶上条锈菌或叶锈菌孢子堆面积占总叶片面积的百分数。每隔7 d调查一次, 共调查2~3次, 直到叶片上孢子堆不再增加为止。

用最大严重度(maximum disease severity, MDS)和病程曲线下面积(area under the disease progress curve, AUDPC)作为抗病性评价指标。

式中, x_i表示第 i次调查的严重度, t_i表示第 i次调查距接种后的天数^{[ 24]}。用SAS 9.2软件进行统计分析和显著性比较。

2 结果与分析

2.1 21个基因聚合株系条锈病和叶锈病抗性鉴定

MDS和AUDPC在株系间和环境间的差异均达极显著水平( P<0.01)。百农64和鲁麦21在郫县和天水两个环境下条锈病平均MDS分别为36.8%和50.0%, 平均AUDPC分别为210.6和290.5; 12个聚合基因株系的条锈病平均MDS均低于百农64和鲁麦21。其中, 株系BFB14在两个环境下的条锈病平均MDS值(16.2%)和平均AUDPC值(92.1)最低; 在保定和周口的叶锈病平均MDS分别为15.8%和35.0%, 平均AUDPC分别为91.6和212.3。其他11个株系的叶锈病平均MDS低于百农64和鲁麦21。株系BFB9在两个环境下叶锈病的平均MDS (3.6%)和平均AUDPC (18.9)最低(表1)。

表1 21个聚合基因株系及其亲本3种病害的最大严重度(MDS)和病程曲线下面积(AUDPC) Table 1 Composition of slow mildewing QTL, and averaged MDS and AUDPC for powdery mildew, stripe rust and leaf rust response in 21 F₆ lines from the Bainong 64/Lumai 21 cross and their parents

品系 Line	慢白粉病QTL Slow mildewing resistance QTL	QTL数 No. of QTLs	白粉病Powdery mildew		条锈病Stripe rust		叶锈病Leaf rust
品系 Line	慢白粉病QTL Slow mildewing resistance QTL	QTL数 No. of QTLs	MDS	AUDPC	MDS	AUDPC	MDS	AUDPC
BN64	1A /4DL/6BS/7A	4	7.0 cde	45.0 defg	36.8 cdefgh	210.6 efgh	15.8 ef	91.6 ef
LM21	2BS /2BL /2DL	3	12.0 a	75.3 a	50.0 abcd	290.5 bcde	35.0 a	212.9 a
BFB5	1A /4DL /2BS	3	3.5 f	22.8 hig	33.7 fghi	195.5 fghi	9.4 fgh	52.0 fgh
BFB6	1A /4DL /2DL	3	4.5 ef	33.3 fghi	35.3 fghi	198.6 fghi	9.0 fgh	50.1 fgh
BFB7	1A /4DL /2DL	3	4.0 ef	31.5 fghi	48.0 abcd	288.3 cde	8.7 fgh	48.0 fgh
BFB8	1A /4DL /2DL /2BS /2BL	5	4.3 ef	39 efghi	27.5 ghijk	160.5 ghijk	4.6 h	25.4 gh
BFB9	1A/4DL /2BS /2BL	4	9.5 abc	63.5 abcd	24.7 hijk	143.3 hijk	3.6 h	18.9 h
BFB10	1A /4DL /2DL /2BS /2BL	5	2.8 f	20.8 hi	21.0 hijk	123.7 jk	8.9 gh	48.5 fgh
BFB11	1A /4DL/2DL/2BS	4	2.8 f	17.3 i	18.2 jk	101.6 jk	5.7 gh	31.6 gh
BFB12	4DL/2BS	2	11.3 ab	72.8 ab	36.5 efgh	186.1 fghij	35.7 a	200.7 a
BFB13	1A/4DL/2DL	3	4.0 ef	23.5 hig	31.5 ghij	172.5 ghijk	15.4 ef	87.2 ef
BFB14	1A/4DL/2DL	3	3.8 ef	22.8 hig	16.2 k	91.7 k	10.0 fgh	54.8 fgh
BFB15	1A/4DL/2DL/2BS	4	8.5 bcd	59.3 abcde	54.3 ab	324.2 abcd	26.7 cd	145.8 cd
BFB16	1A/4DL/2BL/7A	4	5.8 def	51.0 abcdef	26.8 ghijk	168.0 ghijk	15.5 ef	83.3 ef
BFB17	1A/4DL/2BL/7A	4	5.0 ef	35.5 fghi	29.7 ghijk	177.7 fghij	14.3 efg	72.9 efg
BFB18	1A/4DL/2DL/2BL/7A	5	5.8 def	43.8 defgh	27.3 ghijk	162.7 ghijk	32.9 bc	182.1 bc
BFB19	1A/4DL/2DL/2BL/7A	5	5.8 def	32.5 fghi	39.5 cdefg	241.6 defg	21.3 de	114.3 de
BFB20	1A/4DL/2BL/7A	4	4.0 ef	32.8 fghi	39.2 cdefgh	227.1 efgh	21.0 de	113.2 de
BFB21	1A/4DL/2DL/2BL/7A	5	5.3 def	47.8 cdef	45.3 cdef	257.9 cdef	27.9 cd	155.8 cd
BFB22	1A/4DL/7A	3	9.8 abc	68.8 abc	53.5 ab	327.1 abc	31.9 bc	175.9 bc
BFB23	1A/4DL/7A	3	11.3 ab	76.5 a	61.7 a	374.5 a	29.2 bcd	158.8 cd
BFB24	1A/4DL/7A	3	10.0 abc	59.0 abcde	57.7 ab	349.7 ab	32.8 bc	179.0 bc
BFB25	1A/4DL/7A	3	7.0 cde	45.0 defg	50.8 abc	316.2 abcd	27.9 cd	157.0 cd

Slow powdery mildew QTL were mapped by Lan et al.^{[ 21, 22]} QTL 4DL is short for QPm.caas-4DLand by analogy for other QTLs. QPm.caas-4DL and QPm.caas-6BShave significant resistance to powdery mildew, stripe rust and leaf rust; QPm.caas-2BSand QPm.caas- 2BL have significant resistance to powdery mildew and stripe rust^{[ 20, 25]}. Powdery mildew resistance was the means of evaluation values in Beijing and Anyang of Henan province in 2009-2010 and 2010-2011 cropping seasons^{[ 23]}. Stripe rust resistance was the means of evaluation values in Pixian of Sichuan province and Tianshui of Gansu province in 2012-2013 cropping season; Leaf rust resistance was the means of evaluation values in Baoding of Hebei province and Zhoukou of Henan province in 2012-2013 cropping season. Values followed by different letters are significantly different at P < 0.05. BN64: Bainong 64; LM21: Lumai 21.

慢白粉病QTL由Lan等^{[ 21, 22]}鉴定, 其中 QPm.caas-4DL缩写为4DL, 余此类推。 QPm.caas-4DL和 QPm.caas-6BS兼抗白粉病、条锈病和叶锈病, QPm.caas-2BS和 QPm.caas-2BL兼抗白粉病和条锈病^{[ 20, 25]}。白粉病数据为 2009-2010和2010-2011年度北京和河南安阳两点的平均值^{[ 23]}。条锈病数据为2012-2013年度四川郫县和甘肃天水两点的平均值。叶锈病数据为2012-2013年度河北保定和河南周口2点的平均值。平均值后不同字母表示品系间差异显著( P < 0.05)。BN64: 百农64; LM21: 鲁麦21。

2.2 慢白粉病QTL对条锈和叶锈病的抗性效应与互作分析

QPm.caas-4DL、 QPm.caas-2BS和 QPm.caas-2BL对条锈病具有显著的抗性遗传效应( P<0.01), 分别解释条锈病平均表型变异的16.9%、14.1%和17.3%, QPm.caas-6BS可解释1.8%的表型变异。 QPm.caas- 2BS和 QPm.caas-2BL之间存在互作, 但效应较低, 环境与QTL间也存在互作。 QPm.caas-4DL对叶锈病也具有显著抗性, 可解释表型变异的35.3%, 而 QPm.caas-6BS可解释8.3%的表型变异(表2)。

表2 4个环境下慢白粉病抗性QTL对条锈病和叶锈病的效应及其互作 Table 2 Effects of slow mildewing QTL on stripe rust and leaf rust, and their interactions from four environments

2.3 慢白粉病QTL聚合体对条锈和叶锈病的抗性效应

百农64和鲁麦21以及21个慢病性QTL聚合株系中共有11种QTL组合(表3)。 QPm.caas-1A/ QPm.caas-4DL/ QPm.caas-2DL/ QPm.caas-2BS/ QPm.caas-2BL和 QPm.caas-1A/ QPm.caas-4DL/ QPm.caas- 2BS/ QPm.caas-2BL聚合体的条锈病MDS及AUDPC值最低, 远低于两亲本及其他7种QTL聚合体, 且差异显著( P<0.05), 对条锈和叶锈病有较好的抗性。 QPm.caas-1A/ QPm.caas-4DL/ QPm.caas-7A组合的条锈病MDS和AUDPC最高, 与鲁麦21差异不显著, 但显著高于百农64及其他8种QTL聚合体( P<0.05), 21个聚合株系中其余6种QTL聚合体抗性与百农64差异不显著。

表3 不同慢白粉病抗性QTL聚合体对白粉病、条锈病和叶锈病抗性效应 Table 3 Effects of different slow mildewing QTL combinations on powdery mildew, stripe rust, and leaf rust response

QPm.caas-1A/ QPm.caas-4DL/ QPm.caas-2DL/ QPm.caas-2BS/ QPm.caas-2BL、 QPm.caas-1A/ QPm.caas- 4DL/ QPm.caas-2BS/ QPm.caas-2BL和 QPm.caas-1A/ QPm.Caas-4DL/ QPm.caas-2BS组合的叶锈病MDS及AUDPC值较低, 显著低于双亲及其他6种QTL聚合体, 且差异显著( P<0.05), 对叶锈病有较好抗性。 QPm.caas-1A/ QPm.caas-4DL/ QPm.caas-7A和 QPm.caas-4DL/ QPm.caas-2BS组合的叶锈病MDS和AUDPC较高, 与鲁麦21差异不显著, 但显著高于21个聚合株系中其他7种QTL聚合体( P<0.05), 其余4种QTL聚合体抗性与百农64无显著差异。

除株系BFB12外, 随着含有白粉病抗性QTL数量的增多, 聚合株系的条锈病和叶锈病MDS和AUDPC呈现逐渐降低趋势(图1)。

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	图1 不同数量慢白粉病抗性QTL对条锈病和叶锈病的抗性效应Fig. 1 Effects of slow mildewing resistance QTL number on reducing stripe rust and leaf rust

3 讨论

利用分子标记辅助选择聚合慢病性基因是培育兼抗白粉病、条锈病和叶锈病品种的重要途径。迄今为止, 已证实 Yr18/Lr34/Pm38/Sr57、 Yr29/Lr46/ Pm39/Sr58和 Yr46/Lr67/Pm46/Sr55等多个小麦慢病性基因兼抗条锈病、叶锈病、白粉病和秆锈病^{[ 26, 27, 28, 29]}。同时, 还在1BL、2BS、2BL、3BS、6BS和7DS上发现了多个兼抗条锈病、叶锈病和白粉病的基因簇^{[ 30]}。这些基因的发现为培育兼抗多种病害的小麦品种提供了可能。CIMMYT约60%的小麦品种聚合多个慢病性基因, 并已建立通过聚合 Yr18/Lr34/ Pm38/Sr57、 Yr29/Lr46/ Pm39/Sr58和 Yr46/Lr67/Pm46/ Sr55等几个慢病性基因, 选育兼抗几种病害的持久抗性品种的主要育种策略, 运用该策略成功选育出一批兼抗型小麦品种^{[ 4]}。携带 Yr18/Lr34/Pm38/Sr57基因的材料在CIMMYT小麦种质资源中广泛存在, 已保持70多年的抗性, 发展中国家含 Yr18/Lr34/ Pm38/Sr57的小麦品种种植面积约有2600万公顷^{[ 31]}, 在病害流行年份发挥着重要作用。美国、澳大利亚和欧洲的研究重点近年也从垂直抗性逐步转向慢病性^{[ 32, 33, 34]}。

百农64含有的抗白粉病 QPm.caas-4DL与 Yr46/ Lr67/Pm46/Sr55很可能为同一基因, 虽然 Yr46/Lr67/ Pm46/Sr55抗性效应不及 Yr18/Lr34/Pm38/Sr57, 但百农64的农艺性状良好, 该位点将是小麦持久抗病育种的另一重要基因资源。Ren等^{[ 25]}在 QPm. caas-6BS同一位置分别发现条锈病和叶锈病抗性QTL, 该位点是一个新发现的兼抗多种病害的位点。任妍^{[ 20]}在鲁麦21中定位了2个条锈病抗性QTL QYr.caas-2BS和 QYr.caas-2BL, 分别与抗白粉病 QPm.caas-2BS和 QPm.caas-2BL位置一致, 这2个位点也是小麦持久兼抗育种的重要基因。利用慢病性基因容易做到兼抗与持久抗性的结合, 兼抗几种病害的慢病基因都具有持久抗性(Ravi Singh, 个人交流)。已经定名的几个慢病性基因(如 Yr18/Lr34/ Pm38/Sr57、 Yr29/Lr46/Pm39/Sr58和 Yr46/Lr67/Pm46/ Sr55等)都具有这种特点。我国小麦白粉病、条锈病和叶锈病发生严重, 选育兼抗多种病害小麦品种对于我国小麦生产具有重要意义。来自百农64的 QPm.caas-4DL、 QPm.caas-6BS和来自鲁麦21的 QPm.caas-2BL和 QPm.caas-2BS等对条锈病、叶锈病和白粉病皆表现慢病性, 利用这些基因选育兼抗多种病害的小麦品种可取得事半功倍的效果。虽然过去国内也有育成兼抗且具有持久抗性的小麦品种, 但有目的地进行这种育种的尚不多。

在21个聚合株系中, 含有抗病QTL数量较多的株系的MDS和AUDPC值均较低, 表明QTL加性效应起了重要作用。Singh等^{[ 7]}发现聚合4~5个微效基因的小麦材料对条锈和叶锈病呈高抗至免疫。但是, QTL聚合体的遗传效应并不总是与所含抗病QTL的数量成正比。如株系BFB12和BFB14分别含有2个和3个抗性QTL, 但条锈病MDS和AUDPC值均较低。其原因可能是不同的QTL聚合体效应不同, 部分QTL间互作会产生加性效应, 或者某些株系含有未知的抗性QTL, 导致条锈病和叶锈病MDS和AUDPC值较低, 抗性增强。

任妍^{[ 20]}在鲁麦21中还发现另外2个条锈病抗性QTL ( QYr.caas-4DL.2和 QPm.caas-2DS.2), 我们推测本研究的部分株系可能含有这2个QTL, 导致BFB12和BFB14等含有较少兼抗型QTL的株系抗性较强。相同QTL聚合体的株系抗性并不完全一致, 如BFB11和BFB15均含有 QPm.caas-1A/ QPm. caas-4DL/ QPm.caas-2DL/ QPm.caas-2BS基因组合, 但BFB11抗性远大于BFB15, 估计BFB11含有其他条锈病抗性QTL或存在QTL间互作。但综合21个株系数据来看, 株系含有QTL越多, 对多种病害的抗性越强。

结合Bai等^{[ 23]}对21个聚合株系的白粉病鉴定结果进行分析, 株系BFB5、BFB7、BFB8、BFB10、BFB11和BFB14对白粉病、条锈病和叶锈病均具有较高的抗性, 可作为育种亲本材料用于小麦抗锈病和白粉病育种。 QPm.caas-1A/ QPm.caas-4DL/ QPm. caas-2DL/ QPm.caas-2BS/QPm.caas-2BL、QPm.caas-1A/ QPm.caas-4DL/ QPm.caas-2DL和 QPm.caas-1A/ QPm. caas-4DL/ QPm.caas-2BS组合对白粉病、条锈病和叶锈病均具有较好的抗性, 是选育兼抗多种病害小麦慢病性品种的良好材料。 QPm.caas-1A/ QPm.caas- 4DL/ QPm.caas-2BS/QPm.caas-2BL对条锈病和叶锈病抗性较好, 可用于条锈病和叶锈病多发区的抗病育种。

4 结论

6个聚合百农64和鲁麦21慢白粉病抗性QTL的F₆株系对白粉病、条锈病和叶锈病均具有较好的慢病性, 且农艺性状优良, 是选育抗病高产品种的优良材料。证实了利用分子标记进行慢病性QTL聚合的可行性及其有效性, 进一步说明聚合4~5个慢病性QTL足以在田间表现高水平抗性, 为小麦抗病育种提供了材料和育种思路。

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.

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1988

1.175

0.0

... 0%的产量损失^[1,2,3] ...

1996

10.229

0.0

... 0%的产量损失^[1,2,3] ...

2003

0.716

0.0

... 0%的产量损失^[1,2,3] ...

2005

0.731

0.0

... 长期以来, 利用寄主抗性防治这些病害取得了显著成效^[4] ...

... CIMMYT约60%的小麦品种聚合多个慢病性基因, 并已建立通过聚合Yr18/Lr34/ Pm38/Sr57、Yr29/Lr46/ Pm39/Sr58和Yr46/Lr67/Pm46/ Sr55等几个慢病性基因, 选育兼抗几种病害的持久抗性品种的主要育种策略, 运用该策略成功选育出一批兼抗型小麦品种^[4] ...

1970

0.0

... 水平抗性亦称慢病性、成株抗性或非小种专化抗性, 由多个微效基因控制, 对病原菌无小种专化性或专化性弱, 苗期表现为感病, 成株期表现为中抗或高抗, 且抗性持久^[5,6] ...

1982

2.968

0.0

2011

1.643

0.0

. 2011, 179:175-186 DOI:10.1007/s10681-010-0322-9

Race non-specific resistance to rust diseases in CIMMYT spring wheats

1. CIMMYT, Apdo. Postal 6-641, 06600, Mexico, DF, Mexico 2. INIFAP-CEVAMEX, Apdo. Postal 10, 56230, Chapingo, Mexico 3. CIMMYT, Nairobi, Kenya 4. Cereal Disease Laboratory, USDA-ARS, St. Paul, MN, 55108, USA 5. Njoro Plant Breeding Research Center-Kenya Agricultural Research Institute (KARI-NPBRC), P.O. Njoro, Kenya

Abstract Rust diseases continue to cause significant losses to wheat production worldwide. Although the life of effective race-specific resistance genes can be prolonged by using gene combinations, an alternative approach is to deploy varieties that posses adult plant resistance (APR) based on combinations of minor, slow rusting genes. When present alone, APR genes do not confer adequate resistance especially under high disease pressure; however, combinations of 4–5 such genes usually result in “near-immunity” or a high level of resistance. Although high diversity for APR occurs for all three rusts in improved germplasm, relatively few genes are characterized in detail. Breeding for APR to leaf rust and stripe rust in CIMMYT spring wheats was initiated in the early 1970s by crossing slow rusting parents that lacked effective race-specific resistance genes to prevalent pathogen populations and selecting plants in segregating populations under high disease pressure in field nurseries. Consequently most of the wheat germplasm distributed worldwide now possesses near-immunity or adequate levels of resistance. Some semidwarf wheats such as Kingbird, Pavon 76, Kiritati and Parula show high levels of APR to stem rust race Ug99 and its derivatives based on the Sr2 -complex, or a combination of Sr2 with other uncharacterized slow rusting genes. These parents are being utilized in our crossing program and a Mexico-Kenya shuttle breeding scheme is used for selecting resistance to Ug99. High frequencies of lines with near-immunity to moderate levels of resistance are now emerging from these activities. After further yield trials and quality assessments these lines will be distributed internationally through the CIMMYT nursery system.

... Singh等^[7]发现, 聚合4~5个微效慢病基因的小麦材料对条锈和叶锈病呈高抗至免疫 ...

... 随着QTL定位和分子标记研究工作的不断深入, 育种家可以利用分子标记聚合抗病QTL, 培育持久抗性品种^[7,12,13] ...

... Singh等^[7]发现聚合4~5个微效基因的小麦材料对条锈和叶锈病呈高抗至免疫 ...

2009

3.658

0.0

. 2009, 119:1349-1359 DOI:10.1007/s00122-009-1139-6

QTL mapping for adult-plant resistance to stripe rust in Italian common wheat cultivars Libellula and Strampelli

1.Institute of Crop Science, National Wheat Improvement Centre/The National Key Facility for Crop Gene Resources and Genetic Improvement, Chinese Academy of Agricultural Sciences (CAAS), 12 Zhongguancun South Street, 100081 Beijing, China 2.College of Plant Protection, Northwest A&F University, 712100 Yangling, Shaanxi China 3.Gansu Wheat Research Institute, Gansu Academy of Agricultural Sciences, 730070 Lanzhou, Gansu Province China 4.International Maize and Wheat Improvement Centre (CIMMYT), China Office, c/o CAAS, 12 Zhongguancun South Street, 100081 Beijing, China

Italian common wheat cultivars Libellula and Strampelli, grown for over three decades in Gansu province of China, have shown effective resistance to stripe rust. To elucidate the genetic basis of the resistance, F 3 populations were developed from crosses between the two cultivars and susceptible Chinese wheat cultivar Huixianhong. The F 3 lines were evaluated for disease severity in Beijing, Gansu and Sichuan from 2005 to 2008. Joint- and single-environment analyses by composite interval mapping identified five quantitative trait loci (QTLs) in Libellula for reduced stripe rust severity, designated QYr.caas-2DS , QYr.caas-4BL , QYr.caas-5BL.1 , QYr.caas-5BL.2 and QYr.caas-7DS , and explained 8.1–12.4, 3.6–5.1, 3.4–8.6, 2.6 and 14.6–35.0%, respectively, of the phenotypic variance across four environments. Six interactions between different pairs of QTLs explained 3.2–7.1% of the phenotypic variance. The QTLs QYr.caas-4BL , QYr.caas-5BL.1 and QYr.caas-7DS were also detected in Strampelli, explaining 4.5, 2.9–5.5 and 17.1–39.1% of phenotypic variance, respectively, across five environments. Three interactions between different pairs of QTLs accounted for 6.1–35.0% of the phenotypic variance. The QTL QYr.caas-7DS flanked by markers csLV34 and Xgwm295 showed the largest effect for resistance to stripe rust. Sequence analyses confirmed that the lines with the QYr.caas-7DS allele for resistance carried the resistance allele of the Yr18/Lr34 gene. Our results indicated that the adult-plant resistance gene Yr18 and several minor genes confer effective durable resistance to stripe rust in Libellula and Strampelli.

... 抗性遗传分析表明, Strampelli和Libellula含有Yr18/Lr34/Pm38/Sr57和2~4个其他慢病基因^[8] ...

2010

2.968

0.0

... 平原50含有3个慢条锈基因和3个慢白粉基因^[9] ...

2009

0.0

Science. 2009, 323(5919):1360-1363

A Putative ABC Transporter Confers Durable Resistance to Multiple Fungal Pathogens in Wheat

Simon G. Krattinger 1 , * , Evans S. Lagudah 2 , * , † , Wolfgang Spielmeyer 2 , Ravi P. Singh 3 , Julio Huerta-Espino 4 , Helen McFadden 2 , ‡ , Eligio Bossolini 1 , § , Liselotte L. Selter 1 , Beat Keller 1 , †

1 Institute of Plant Biology, University of Zurich, Zollikerstrasse 107, 8008 Zurich, Switzerland. 2 Commonwealth Scientific and Industrial Research Organisation (CSIRO) Plant Industry, GPO Box 1600, Canberra, ACT, 2601, Australia. 3 International Maize and Wheat Improvement Center (CIMMYT), Apdo. Postal 6-641, 06600 Mexico DF, Mexico. 4 Campo Experimental Valle de Mexico INIFAP, Apdo. Postal 10, 56230 Chapingo, Edo de Mexico, Mexico. ↵ † To whom correspondence should be addressed. E-mail: evans.lagudah@csiro.au (E.S.L.), bkeller@botinst.uzh.ch (B.K.)

... Yr18/Lr34/Pm38/Sr57对小麦条锈病、叶锈病、白粉病和秆锈病均具有抗性^[10,11] ...

2011

0.0

... Yr18/Lr34/Pm38/Sr57对小麦条锈病、叶锈病、白粉病和秆锈病均具有抗性^[10,11] ...

2000

0.0

... 随着QTL定位和分子标记研究工作的不断深入, 育种家可以利用分子标记聚合抗病QTL, 培育持久抗性品种^[7,12,13] ...

2003

1.513

0.0

... 随着QTL定位和分子标记研究工作的不断深入, 育种家可以利用分子标记聚合抗病QTL, 培育持久抗性品种^[7,12,13] ...

2002

0.0

... 从20世纪70年代开始, 国际玉米小麦改良中心(CIMMYT)选育出一批抗性持久且兼抗多种病害的慢病性品种, 例如Amadina、Chapio、Cook、Kukuna、Parula、Pavon 76、Sonoita 81、Tonichi 81和Tukuru, 均含有Yr18/Lr34/Pm38/Sr57或Yr29/ Lr46/Pm39/Sr58位点及2~3个微效基因^[14,15] ...

2004

0.0

2003

41.063

0.0

... 虽然小麦抗病性QTL定位研究很多, 但是由于不同遗传背景下的QTL聚合到同一遗传背景存在诸多困难^[16], 不同来源的QTL可能存在互作, QTL聚合育种实践的报道很少 ...

2006

3.658

0.0

. 2006, 112:562-569 DOI:10.1007/s00122-005-0163-4

Stacking quantitative trait loci (QTL) for Fusarium head blight resistance from non-adapted sources in an European elite spring wheat background and assessing their effects on deoxynivalenol (DON) content and disease severity

1.State Plant Breeding Institute, University of Hohenheim (720), 70593 Stuttgart, Germany 2.Department for Agrobiotechnology Tulln, Institute of Plant Production Biotechnology, BOKU-University of Natural Resources and Applied Life Sciences Vienna, Konrad Lorenz Str. 20, 3430 Tulln, Austria 3.Lochow-Petkus GmbH, 29296 Bergen, Germany

Fusarium head blight (FHB) is a devastating disease in wheat that reduces grain yield, grain quality and contaminates the harvest with deoxynivalenol (DON). As potent resistance sources Sumai 3 and its descendants from China and Frontana from Brazil had been analysed by quantitative trait loci (QTL) mapping. We introgressed and stacked two donor QTL from CM82036 (Sumai 3/Thornbird) located on chromosomes 3B and 5A and one donor QTL from Frontana on chromosome 3A in elite European spring wheat and estimated the effects of the three individual donor QTL and their four combinations on DON, Fusarium exoantigen content, and FHB rating adjusted to heading date. One class with the susceptible QTL alleles served as control. Each of the eight QTL classes was represented by 12–15 F 3 -derived lines tested in F 5 generation as bulked progeny possessing the respective marker alleles homozygously. Traits were evaluated in a field experiment across four locations with spray inoculation of Fusarium culmorum. All three individual donor-QTL alleles significantly reduced DON content and FHB severity compared to the marker class with no donor QTL. The only exception was the donor-QTL allele 3A that had a low, but non-significant effect on FHB severity. The highest effect had the stacked donor-QTL alleles 3B and 5A for both traits. They jointly reduced DON content by 78% and FHB rating by 55% compared to the susceptible QTL class. Analysis of Fusarium exoantigen content illustrates that lower disease severity is associated with less mycelium content in the grain. In conclusion, QTL from non-adapted sources could be verified in a genetic background of German elite spring wheat. Within the QTL classes significant ( P <0.05) genotypic differences were found among the individual genotypes. An additional phenotypic selection would, therefore, be advantageous after performing a marker-based selection.

... Miedaner等^[17]将3个来源不同的赤霉病抗性QTL聚合到同一种质, 发现单个QTL均可降低赤霉病发病率和脱氧雪腐镰刀菌烯醇(DON)含量, 但3A上的一个QTL单独存在或与3B和5A上QTL聚合时对赤霉病没有显著遗传效应 ...

2011

1.513

0.0

... Lu等^[18]将5A上的QTL与Fhb1聚合在一起, 发现可以降低矮秆基因Rht-D1b对赤霉病引起的效应 ...

2005

2.455

0.0

... 百农64和鲁麦21均具有优良的农艺性状, 分别在20世纪90年代和21世纪初为我国黄淮麦区主推品种, 其苗期对白粉菌^[19]、条锈菌和叶锈菌^[20]流行小种均感病, 但田间成株期表现抗病, 具有典型的慢病性特征 ...

2012

0.0

... 任妍^[20]在鲁麦21中定位了2个条锈病抗性QTL QYr ...

... 任妍^[20]在鲁麦21中还发现另外2个条锈病抗性QTL (QYr ...

2009

2.968

0.0

... caas-7A) ^[21], 在鲁麦21中检测到3个慢白粉病抗性QTL (QPm ...

... 利用16个分子标记^[21,22]进行检测, 明确这些株系携带2~5个慢白粉病QTL ...

2010

3.251

0.0

. 2010, 25:615-622 DOI:10.1007/s11032-009-9358-8

Quantitative trait loci mapping of adult-plant resistance to powdery mildew in Chinese wheat cultivar Lumai 21

1.Institute of Crop Science, National Wheat Improvement Center/The National Key Facility for Crop Gene Resources and Genetic Improvement, Chinese Academy of Agricultural Sciences (CAAS), 12 Zhongguancun South Street, 100081 Beijing, China 2.Cotton Research Institute, Chinese Academy of Agricultural Sciences (CAAS), Huanghedadao, 455000 Anyang, Henan China 3.International Maize and Wheat Improvement Center (CIMMYT) China Office, c/o CAAS, 12 Zhongguancun South Street, 100081 Beijing, China

Powdery mildew, caused by Blumeria graminis f. sp. tritici , is a major fungal disease in common wheat ( Triticum aestivum L.) worldwide. The Chinese winter wheat cultivar Lumai 21 has shown good and stable adult plant resistance for 19 years. The aim of this study was to map quantitative trait loci (QTLs) for resistance to powdery mildew in a population of 200 F 3 lines from the cross Lumai 21/Jingshuang 16. The population was tested for powdery mildew reaction in Beijing and Anyang in the 2005–2006 and 2006–2007 cropping seasons, providing data for 4 environments. A total of 1,375 simple sequence repeat (SSR) markers were screened for associations with powdery mildew reactions, initially in bulked segregant analysis. Based on the mean disease values averaged across environments, broad-sense heritabilities of maximum disease severity and area under the disease progress curve were 0.96 and 0.77, respectively. Three QTLs for adult plant resistance were detected by inclusive composite interval mapping. These were designated QPm.caas-2BS , QPm.caas-2BL and QPm.caas-2DL , respectively, and explained from 5.4 to 20.6% of the phenotypic variance across environments. QPm.caas-2BS and QPm.caas-2DL were likely new adult plant resistance QTLs flanked by SSR markers Xbarc98 – Xbarc1147 and Xwmc18 – Xcfd233 , respectively. These markers could be useful for improving wheat powdery mildew resistance in breeding programs.

... caas-2DL) ^[22] ...

... 利用16个分子标记^[21,22]进行检测, 明确这些株系携带2~5个慢白粉病QTL ...

2011

1.133

0.0

... Bai等^[23]采用杂交和改良系谱法, 对百农64和鲁麦21所含慢白粉病抗性QTL进行聚合, 创制了21个F₆聚合株系, 并对这些材料的白粉病抗性进行了分析 ...

... 1 聚合株系基因型及其分子标记百农64与鲁麦21杂交, 采用改良系谱法, 根据田间白粉病抗性和综合农艺性状选择, 在早代淘汰白粉病严重度高的株系, 选择低严重度株系继续种植, 经过连续多代鉴定和选择, 获得21个F₆抗病株系(编号: BFB5~BFB25)^[23] ...

... 结合Bai等^[23]对21个聚合株系的白粉病鉴定结果进行分析, 株系BFB5、BFB7、BFB8、BFB10、BFB11和BFB14对白粉病、条锈病和叶锈病均具有较高的抗性, 可作为育种亲本材料用于小麦抗锈病和白粉病育种 ...

2009

3.658

0.0

. 2009, 118:631-642 DOI:10.1007/s00122-008-0894-0

Quantitative trait loci for non-race-speci#cod#x0fb01;c, high-temperature adult-plant resistance to stripe rust in wheat cultivar Express

1.Washington State University Department of Plant Pathology Pullman WA 99164-6430 USA 2.Shenyang Agricultural University Bioscience and Technology College Liaoning 110161 People’s Republic of China 3.US Department of Agriculture Agricultural Research Service, Wheat Genetics, Quality, Physiology, and Disease Research Unit Pullman WA 99164-6430 USA

Wheat cultivar Express has durable, high-temperature adult-plant (HTAP) resistance to stripe rust ( Puccinia striiformis f. sp. tritici ). To elucidate the genetic basis of the resistance, Express was crossed with ‘Avocet Susceptible’ (AVS). A mapping population of 146 F 5 recombinant inbred lines (RILs) was developed using single-seed descent. The RILs were evaluated at two sites near Pullman in eastern Washington and one site near Mount Vernon in western Washington in 2005, and were evaluated near Pullman in 2006 under natural stripe rust infection of predominant races virulent on seedlings of Express. Infection type (IT) and disease severity (DS) were recorded three times for each line during each growing season. The DS data were used to calculate relative area under the disease progress curve (rAUDPC) values. Both IT and rAUDPC data showed continuous distributions, indicating that the Express HTAP resistance was controlled by quantitative trait loci (QTL). Resistance gene analog polymorphism (RGAP) and simple sequence repeat (SSR) techniques were used to map the HTAP resistance QTL. Three QTL were detected with significant additive effects, explaining 49.5–69.6% of the phenotypic variation for rAUDPC. Two of the QTL explained 30.8–42.7% of the phenotypic variation for IT. The three QTL were mapped to wheat chromosomes 6AS, 3BL and 1BL, and were designated as QYrex.wgp-6AS , QYrex.wgp-3BL and QYrex.wgp-1BL , respectively. QYrex.wgp-6AS and QYrex.wgp-3BL , which had higher effects than QYrex.wgp-1BL , were different from previously reported QTL/genes for adult-plant resistance. Markers Xgwm334–Xwgp56 and Xgwm299–Xwgp66 flanking the two major QTL were highly polymorphic in various wheat genotypes, suggesting that these markers are useful in marker-assisted selection.

... 式中, x_i表示第i次调查的严重度, t_i表示第i次调查距接种后的天数^[24] ...

2012

3.658

0.0

Theor Appl Genet. 2012, 125(6):1253-1262

QTL mapping of adult-plant resistances to stripe rust and leaf rust in Chinese wheat cultivar Bainong 64

Yan Ren (1) , Zaifeng Li (2) , Zhonghu He (1) (5) , Ling Wu (3) , Bin Bai (1) (4) , Caixia Lan (1) , Cuifen Wang (2) , Gang Zhou (4) , Huazhong Zhu (3) , Xianchun Xia (1)

1. Institute of Crop Science, National Wheat Improvement Center/The National Key Facility for Crop Gene Resources and Genetic Improvement, Chinese Academy of Agricultural Sciences (CAAS), 12 Zhongguancun South Street, Beijing, 100081, China 2. Department of Plant Pathology, College of Plant Protection, Agricultural University of Hebei, Biological Control Center for Plant Diseases and Plant Pests of Hebei, 289 Lingyusi Street, Baoding, 071001, Hebei, China 5. International Maize and Wheat Improvement Center (CIMMYT) China Office, c/o CAAS, 12 Zhongguancun South Street, Beijing, 100081, China 3. Crop Research Institute, Sichuan Academy of Agricultural Sciences, Chengdu, 610066, Sichuan, China 4. Wheat Research Institute, Gansu Academy of Agricultural Sciences, Lanzhou, 730070, Gansu, China

Stripe rust and leaf rust, caused by Puccinia striiformis Westend. f. sp. tritici Erikss. and P . triticina , respectively, are devastating fungal diseases of common wheat ( Triticum aestivum L.). Chinese wheat cultivar Bainong 64 has maintained acceptable adult-plant resistance (APR) to stripe rust, leaf rust and powdery mildew for more than 10 years. The aim of this study was to identify quantitative trait loci/locus (QTL) for resistance to the two rusts in a population of 179 doubled haploid (DH) lines derived from Bainong 64 × Jingshuang 16. The DH lines were planted in randomized complete blocks with three replicates at four locations. Stripe rust tests were conducted using a mixture of currently prevalent P . striiformis races, and leaf rust tests were performed with P . triticina race THTT. Leaf rust severities were scored two or three times, whereas maximum disease severities (MDS) were recorded for stripe rust. Using bulked segregant analysis (BSA) and simple sequence repeat (SSR) markers, five independent loci for APR to two rusts were detected. The QTL on chromosomes 1BL and 6BS contributed by Bainong 64 conferred resistance to both diseases. The loci identified on chromosomes 7AS and 4DL had minor effects on stripe rust response, whereas another locus, close to the centromere on chromosome 6BS, had a significant effect only on leaf rust response. The loci located on chromosomes 1BL and 4DL also had significant effects on powdery mildew response. These were located at the same positions as the Yr29/Lr46 and Yr46/Lr67 genes, respectively. The multiple disease resistance locus for APR on chromosome 6BS appears to be new. All three genes and their closely linked molecular markers could be used in breeding wheat cultivars with durable resistance to multiple diseases.

... Ren等^[25]在QPm ...

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 .

... 迄今为止, 已证实Yr18/Lr34/Pm38/Sr57、Yr29/Lr46/ Pm39/Sr58和Yr46/Lr67/Pm46/Sr55等多个小麦慢病性基因兼抗条锈病、叶锈病、白粉病和秆锈病^{[26,27,28,29]} ...

1992

2.968

0.0

... 迄今为止, 已证实Yr18/Lr34/Pm38/Sr57、Yr29/Lr46/ Pm39/Sr58和Yr46/Lr67/Pm46/Sr55等多个小麦慢病性基因兼抗条锈病、叶锈病、白粉病和秆锈病^{[26,27,28,29]} ...

1994

1.668

0.0

... 迄今为止, 已证实Yr18/Lr34/Pm38/Sr57、Yr29/Lr46/ Pm39/Sr58和Yr46/Lr67/Pm46/Sr55等多个小麦慢病性基因兼抗条锈病、叶锈病、白粉病和秆锈病^{[26,27,28,29]} ...

2011

3.658

0.0

Theor Appl Genet. 2011, 122(1):239-249

New slow-rusting leaf rust and stripe rust resistance genes Lr67 and Yr46 in wheat are pleiotropic or closely linked

Sybil A. Herrera-Foessel (1) , Evans S. Lagudah (2) , Julio Huerta-Espino (3) , Matthew J. Hayden (4) , Harbans S. Bariana (5) , Davinder Singh (6) , Ravi P. Singh (1)

1. International Maize and Wheat Improvement Center (CIMMYT), Apdo. Postal 6-641, 06600, Mexico, D.F., Mexico 2. CSIRO Plant Industry, GPO Box 1600, Canberra, ACT, 2601, Australia 3. Campo Experimental Valle de México INIFAP, Apdo. Postal 10, 56230, Chapingo, Edo. de Mexico, Mexico 4. Department of Primary Industries, Victorian AgriBiosciences Center, Bundoora, VIC, 3083, Australia 5. The University of Sydney Plant Breeding Institute-Cobbitty, Private Bag 4011, Narellan, NSW, 2567, Australia 6. CIMMYT, ICRAF House, United Nations Avenue, Gigiri, Village Market, PO Box 1041, Nairobi, 00621, Kenya

The common wheat genotype ‘RL6077’ was believed to carry the gene Lr34 / Yr18 that confers slow-rusting adult plant resistance (APR) to leaf rust and stripe rust but located to a different chromosome through inter-chromosomal reciprocal translocation. However, haplotyping using the cloned Lr34/Yr18 diagnostic marker and the complete sequencing of the gene indicated Lr34/Yr18 is absent in RL6077. We crossed RL6077 with the susceptible parent ‘Avocet’ and developed F 3 , F 4 and F 6 populations from photoperiod-insensitive F 3 lines that were segregating for resistance to leaf rust and stripe rust. The populations were characterized for leaf rust resistance at two Mexican sites, Cd. Obregon during the 2008–2009 and 2009–2010 crop seasons, and El Batan during 2009, and for stripe rust resistance at Toluca, a third Mexican site, during 2009. The F 3 population was also evaluated for stripe rust resistance at Cobbitty, Australia, during 2009. Most lines had correlated responses to leaf rust and stripe rust, indicating that either the same gene, or closely linked genes, confers resistance to both diseases. Molecular mapping using microsatellites led to the identification of five markers ( Xgwm165 , Xgwm192 , Xcfd71 , Xbarc98 and Xcfd23 ) on chromosome 4DL that are associated with this gene(s), with the closest markers being located at 0.4 cM. In a parallel study in Canada using a Thatcher × RL6077 F 3 population, the same leaf rust resistance gene was designated as Lr67 and mapped to the same chromosomal region. The pleiotropic, or closely linked, gene derived from RL6077 that conferred stripe rust resistance in this study was designated as Yr46 . The slow-rusting gene(s) Lr67 / Yr46 can be utilized in combination with other slow-rusting genes to develop high levels of durable APR to leaf rust and stripe rust in wheat.

... 迄今为止, 已证实Yr18/Lr34/Pm38/Sr57、Yr29/Lr46/ Pm39/Sr58和Yr46/Lr67/Pm46/Sr55等多个小麦慢病性基因兼抗条锈病、叶锈病、白粉病和秆锈病^{[26,27,28,29]} ...

2011

0.0

1.889

. 2011, 44:2193-2215 DOI:10.3864/j.issn.0578-1752.2011.11.001

Progress and perspective in research of adult-plant resistance to stripe rust and powdery mildew in wheat

中国农业科学院作物科学研究所/国家小麦改良中心

Stripe rust, caused by Puccinia striiformis f. sp. tritici, and powdery mildew, caused by Blumeria graminis f. sp. tritici, are the devastating diseases in common wheat (Triticum aestivum L.) worldwide. Use of adult-plant resistance (APR) genes is an important method for the development of durable resistant cultivars. A total of 72 quantitative trait loci (QTLs) for APR to stripe rust and 82 QTLs for APR to powdery mildew were integrated into a linkage map based on the information of DNA markers linked to individual QTL. Eight gene clusters (≥5 QTLs) conferred resistance to both stripe rust and powdery powdery, among them, Yr18/Lr34/Pm38, Yr29/Lr46/Pm39, and Yr46/Lr67 showed resistance to stripe rust, leaf rust and powdery mildew. Yr18/Lr34/Pm38 and Yr36 have been cloned. Xiannong 4 and Xiaoyan 6 were very important resistant germplasm for APR to stripe rust and powdery midew. The application of APR to wheat breeding in China was summarized. The use of APR genes will be a major method for improving stripe rust and powdery mildew resistance in wheat breeding. The strategies for APR on wheat breeding were also discussed.

利用成株抗性是小麦抗病育种的重要方向，本文综述了小麦条锈病和白粉病成株抗性鉴定方法、基因定位和克隆及其在育种中的应用。将报道的72个条锈病成株抗性数量性状遗传位点（quantitative trait loci，QTL）和82个白粉病成株抗性QTL整合到一张连锁图谱上，控制2种病害的基因簇（≥5个QTL）有8个，其中位于7DS的Yr18/Lr34/Pm38和1BS的Yr29/Lr46/Pm39对条锈病、叶锈病和白粉病均表现成株抗性，位于4DL的Yr46/Lr67位点可能也对白粉病表现成株抗性，Yr18/Lr34/Pm38和Yr36已被克隆，Yr29/Lr46/Pm39的克隆已取得良好进展，为培育兼抗和成株抗性相结合的品种提供了可用基因。总结了成株抗性在中国小麦育种中的应用现状，并用实例证实了培育成株抗性品种的可行性，建议对兼抗条锈病和白粉病成株抗性的咸农4号和小偃6号等进行遗传分析，育种工作者和品种审定部门需要转变观念，将成株抗性利用作为国内条锈病和白粉病抗性育种的重要内容。

... 同时, 还在1BL、2BS、2BL、3BS、6BS和7DS上发现了多个兼抗条锈病、叶锈病和白粉病的基因簇^[30] ...

2003

0.0

. 2003, 29(3):253-263

The economic impact of productivity maintenance research: breeding for leaf rust resistance in modern wheat

C.N. Marasas 1,‡,* ,M. Smale 2 andR.P. Singh 3

1 Agriculturul Research Council, Pretoria, South Africa 2 International Food Policy Research Institute (IFPRI) and International Plant Genetic Resources Institute (IPGRI), Washington, DC, USA 3 International Maize and Wheat Improvement Center (CIMMYT), Mexico, DF, Mexico * *Corresponding author. E-mail address: carissa.marasas@usda.gov (C.N. Marasas).

> This paper reports the results of a study undertaken to estimate the economic impact in developing countries of efforts by the International Maize and Wheat Improvement Center (CIMMYT) to breed leaf rust resistant spring bread wheat varieties since 1973. The challenge in estimating these benefits lies in the pathogen's ability to mutate to new races, which may infect previously resistant varieties. Genetic resistance, rather than fungicide application, is the principal means of controlling leaf rust in developing countries. Whereas productivity enhancement is often estimated in terms of yield gains and increased supply, productivity maintenance is measured in terms of the yield losses avoided by the research investment. The internal rate of return on CIMMYT's research investment was estimated at 41%. When discounted by 5%, the net present value was US$ 5.36 billion in 1990 dollars, and the benefit-cost ratio was 27:l. These findings emphasise the economic importance of maintenance research in crop breeding programs.

... 携带Yr18/Lr34/Pm38/Sr57基因的材料在CIMMYT小麦种质资源中广泛存在, 已保持70多年的抗性, 发展中国家含Yr18/Lr34/Pm38/Sr57的小麦品种种植面积约有2600万公顷^[31], 在病害流行年份发挥着重要作用 ...

1995

2.968

0.0

... 美国、澳大利亚和欧洲的研究重点近年也从垂直抗性逐步转向慢病性^[32,33,34] ...

1998

0.0

... 美国、澳大利亚和欧洲的研究重点近年也从垂直抗性逐步转向慢病性^[32,33,34] ...

1999

3.658

0.0

Theor Appl Genet. 1999, 98(6-7):903-912

Quantitative trait loci for resistance against powdery mildew in a segregating wheat×spelt population

M. Keller (A1) , B. Keller (A3) , G. Schachermayr (A1) , M. Winzeler (A1) , J. E. Schmid (A2) , P. Stamp (A2) , M. M. Messmer (A1)

A1. Swiss Federal Research Station for Agroecology and Agriculture (FAL-Reckenholz), Reckenholzstrasse 191, CH-8046 Zurich, Switzerland Tel.: 41-1-3777 445 Fax: 41-1-3777 201 E-mail: monika.messmer@fal.admin.ch, CH A3. Institute of Plant Biology, University of Zurich, Zollikerstrasse 107, CH-8008 Zurich, Switzerland, CH A2. Institute of Plant Sciences, Swiss Federal Institute of Technology (ETH), LFW, ETH-Zentrum, Universitätsrasse 2, CH-8092 Zurich, Switzerland, CH

Powdery mildew is one of the major diseases of wheat in regions with a maritime or semi-continental climate and can strongly affect grain yield. The attempt to control powdery mildew with major resistance genes ( Pm genes) has not provided a durable resistance. Breeding for quantitative resistance to powdery mildew is more promising, but is difficult to select on a phenotypic basis. In this study, we mapped and characterised quantitative trait loci (QTLs) for adult-plant powdery mildew resistance in a segregating population of 226 recombinant inbred lines derived from the cross of the Swiss wheat variety Forno with the Swiss spelt variety Oberkulmer. Forno possibly contains the Pm5 gene and showed good adult-plant resistance in the field. Oberkulmer does not have any known Pm gene and showed a moderate susceptible reaction. Powdery mildew resistance was assessed in field trials at two locations in 1995 and at three locations in 1996. The high heritability (h 2 =0.97) for powdery mildew resistance suggests that the environmental influence did not affect the resistance phenotype to a great extent. QTL analysis was based on a genetic map containing 182 loci with 23 linkage groups (2469 cM). With the method of composite interval mapping 18 QTLs for powdery mildew resistance were detected, explaining 77% of the phenotypic variance in a simultaneous fit. Two QTLs with major effects were consistent over all five environments. One of them corresponds to the Pm5 locus derived from Forno on chromosome 7B. The other QTL on 5A, was derived from the spelt variety Oberkulmer and did not correspond to any known Pm gene. In addition, five QTLs were consistent over three environments, and six QTLs over two environments. The QTL at the Pm5 locus showed a large effect, although virulent races for Pm5 were present in the mixture of isolates. Molecular markers linked with QTLs for adult-plant resistance offer the possibility of simultaneous marker-assisted selection for major and minor genes.

... 美国、澳大利亚和欧洲的研究重点近年也从垂直抗性逐步转向慢病性^[32,33,34] ...