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CIMMYT小麦种质C615抗叶锈病QTL分析

本站小编 Free考研考试/2021-12-26

李玉玲1, 蒋正宁2, 胡文静2, 李东升2, 程婧晔3, 裔新1, 程晓明2, 吴荣林2, 程顺和,1,2,*1 南京农业大学, 江苏南京 210095
2 江苏里下河地区农业科学研究所 / 国家小麦改良中心扬州分中心, 江苏扬州 225007
3 扬州大学农学院, 江苏扬州 225009

Mapping QTLs against Leaf Rust in CIMMYT Wheat C615

LI Yu-Ling1, JIANG Zheng-Ning2, HU Wen-Jing2, LI Dong-Sheng2, CHENG Jing-Ye3, YI Xin1, CHENG Xiao-Ming2, WU Rong-Lin2, CHENG Shun-He,1,2,* 1 Nanjing Agricultural University, Nanjing 210095, Jiangsu, China
2 Lixiahe Institute of Agriculture Sciences of Jiangsu Province / Branch of National Wheat Improvement Center, Yangzhou 225007, Jiangsu, China
3 Agricultural College of Yangzhou University, Yangzhou 225009, Jiangsu, China

通讯作者: 程顺和, E-mail: yzcsh1939@126.com

第一联系人: E-mail: liyuling0817@126.com
收稿日期:2017-09-26接受日期:2018-03-25网络出版日期:2018-06-12
基金资助:本研究由国家现代农业产业技术体系建设专项.CARS-03-03B
本研究由国家现代农业产业技术体系建设专项.CARS-3-2-11
江苏省农业科技自主创新资金项目.CX(14)5080
江苏省自然科学基金项目资助.BK20171279


Received:2017-09-26Accepted:2018-03-25Online:2018-06-12
Fund supported: This study was supported by the China Agriculture Research System.CARS-03-03B
This study was supported by the China Agriculture Research System.CARS-3-2-11
the Independent Innovation Fund for Agricultural Science and Technology in Jiangsu Province.CX(14)5080
the Natural Science Foundation of Jiangsu Province.BK20171279


摘要
Puccinia triticina引起的叶锈病是小麦主要病害之一, 引进种质C615具有叶锈病成株期抗性, 但其抗病性遗传机制尚不清楚。本研究以抗病亲本C615与高感叶锈病亲本宁麦18构建的F2:7代重组自交系群体为材料, 利用337对多态性SSR标记构建遗传连锁图谱, 结合2016、2017连续两年的叶锈病鉴定结果进行复合区间作图, 结果在1BL、2DS、3BS、4DL和6BS染色体上共发现了5个抗性QTL, 暂命名为QLr.njau-1BLQLr.njau-2DSQLr.njau-3BSQLr.njau-4DLQLr.njau-6BS。其中, QLr.njau-1BLQLr.njau-3BSQLr.njau-4DL在两年均被检测到, 分别解释10.1%~15.7%、10.9%~13.5%和8.2%~9.0%的表型变异; 另2个QTL只在一年被检测到, 解释6.2%和9.2%的表型变异。除QLr.njau-2DS外的4个抗性QTL均来源于抗病亲本C615。QLr.njau-1BLQLr.njau-4DL分别与已报道的慢病性基因Lr46Lr67在同一区域, QLr.njau-3B可能为一个新的抗叶锈病QTL。此外, 本研究在C615/扬麦13 (轮回亲本)BC4F5回交群体中选出了15个农艺性状优良且抗叶锈病的株系, 利用与C615所含抗性QTL紧密连锁的7个SSR标记对其进行基因型检测, 结果显示所有这15个株系均含有来自C615的抗性QTL, 且有3个株系聚合了全部抗性位点, 表明C615可作为抗源亲本用于高产、抗病育种。本研究结果将为分子标记选育抗叶锈品种提供材料和技术支撑。
关键词: 小麦;叶锈病;QTL;标记辅助选择育种

Abstract
Leaf rust, caused by Puccinia triticina, is an important wheat disease. The introduced wheat line C615 shows adult resistance to leaf rust, but its mechanism is not clear. In this study, the quantitative trait loci (QTLs) for leaf rust resistance were identified and located in a linkage map constructed with 337 SSR markers using a recombinant inbred line (RIL) population (112 lines of the F2:7 generation) derived from CIMMYT wheat C615 (resistant) and Ningmai 18 (highly susceptible). Final disease severity (FDS) of leaf rust was evaluated in 2016 and 2017. Five QTLs were detected on chromosomes 1BL, 2DS, 3BS, 4DL, and 6BS, designated QLr.njau-1BL, QLr.njau-2DS, QLr.njau-3BS, QLr.njau-4DL, and QLr.njau-6BS, respectively. All the resistance alleles were contributed by the resistant parent C615, except for QLr.njau-2DS. Three QTLs, QLr.njau-1BL, QLr.njau-3BS, and QLr.njau-4DL, were detected in both years, which explained phenotypic variance by 10.1%-15.7%, 10.9%-13.5%, and 8.2%-9.0%, respectively. The remaining two QTLs were detected in one year with smaller contributions (6.2% and 9.2%, respectively). QLr.njau-1BL and QLr.njau-4DL were located in the same regions of the known resistance genes Lr46 and Lr67, respectively. QLr.njau-3BS in the marker interval Xbarc102-Xwmc623 is probably a novel QTL for leaf rust resistance. Fifteen BC4F5 lines derived from the cross of C615/Yangmai 13 (recurrent parent) were genotyped for leaf rust resistance using seven SSR markers closely linked to the QTLs from C615. All the 15 lines had QTLs donated by C615 and three of them pyramided four QTLs, indicating that C615 can serve as an elite donor parent in wheat breeding aiming at high-yield and high-resistance to leaf rust. These results provide a basis of marker-assisted selection and providing breeding materials.
Keywords:wheat (Triticum aestivum L.);leaf rust;QTL mapping;marker-assisted selection


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本文引用格式
李玉玲, 蒋正宁, 胡文静, 李东升, 程婧晔, 裔新, 程晓明, 吴荣林, 程顺和. CIMMYT小麦种质C615抗叶锈病QTL分析[J]. 作物学报, 2018, 44(6): 836-843. doi:10.3724/SP.J.1006.2018.00836
LI Yu-Ling, JIANG Zheng-Ning, HU Wen-Jing, LI Dong-Sheng, CHENG Jing-Ye, YI Xin, CHENG Xiao-Ming, WU Rong-Lin, CHENG Shun-He. Mapping QTLs against Leaf Rust in CIMMYT Wheat C615[J]. Acta Agronomica Sinica, 2018, 44(6): 836-843. doi:10.3724/SP.J.1006.2018.00836


小麦叶锈病是由叶锈菌(Puccinia triticina)引起的真菌性病害, 是世界范围内分布最广泛的小麦病害之一。该病害主要侵染小麦叶片, 进而影响光合作用, 导致感病品种灌浆不足、粒重下降, 严重时可使小麦减产40%以上[1]。近年来, 由于全球气候变暖, 气候条件更加适合小麦叶锈菌的生长与繁殖, 小麦叶锈病的发生日益严重[2]。选育和种植抗病品种是应对叶锈病危害的最经济、安全的途径。

小麦对叶锈病的抗性按照其生长发育阶段可分为苗期抗性和成株抗性。苗期抗性一般为主基因控制, 多表现为小种专化性抗性, 在全生育期表现抗病。与苗期抗性对应的是成株抗性, 一般受微效多基因控制, 在苗期表现感病, 但在成株期表现较好的耐病性[3]。与苗期抗性相比, 成株抗性减少了对病原小种的选择压力, 不易引起病原菌变异, 因而表现出更好的持久抗性[4]。目前国际上已发现了100多个小麦抗叶锈病基因, 其中正式命名的有76个, 具有成株抗性的仅有Lr34[5,6,7,8,9,10]、Lr46[11,12,13]Lr67[14,15]Lr68[16]等11个[17]。利用成株期抗性基因容易做到兼抗与持久抗性的结合, 已经定名的Lr34/Yr18/ Pm38/Sr57Lr46/Yr29/Pm39/Sr58Lr67/Yr46/ Pm46/Sr55等慢病基因都具有这一特点, 即表现出对叶锈、条锈、白粉及秆锈病的兼抗性。除此之外, Rosewarne等[18]、Crossa等[19]和Ren等[20]还在2BS、3BS、4BL和6BS染色体上发现了多个兼抗小麦叶锈、条锈和白粉病的基因簇。这些基因的发现为培育兼抗多种病害的持久抗病品种提供了可能。

20世纪80年代中期以后我国长江下游麦区叶锈病较少发生, 即使发生范围也较小, 严重度较轻。但最近几年, 叶锈病危害呈急剧上升势头, 严重度显著高于常年, 对小麦产量造成重大损失[21,22]。由于本麦区小麦品种大多不具有叶锈病抗性, 发掘利用新的抗叶锈病基因资源具有重要意义。从国际玉米小麦改良中心(CIMMYT)引进的小麦种质C615成株期对叶锈病表现出较好的抗性, 但尚未见其抗病位点和染色体分布等报道。本研究旨在通过构建C615/宁麦18组合的RIL群体, 结合田间叶锈病严重度鉴定, 发掘C615中叶锈病抗性QTL及其紧密连锁标记, 为进一步利用该抗病资源及标记辅助抗叶锈病育种提供技术支撑。

1 材料与方法

1.1 试验材料及田间试验

以高感叶锈病品种宁麦18为母本, C615 (系谱为SABUF/3/BCN//CETA/AE.SQUARROSA(895))为父本进行杂交, F1代用单粒传法得到包含112个家系的RIL群体(F2:7代)。此外, 以C615为供体亲本, 综合性状优良但高感叶锈病品种扬麦13为轮回亲本构建了包含255个家系的BC4F5回交群体, 经过两年表型鉴定, 获得15个农艺性状较好的抗病株系(编号BL-1~BL-15)。2015—2016和2016— 2017连续2个生长季(简称为2016年和2017年), 将亲本、RIL群体及回交群体种植于江苏里下河地区农业科学研究所万福基地试验田, 采用随机区组设计, 3次重复, 行长1.3 m, 行距0.3 m, 每行点播40粒。

1.2 田间抗叶锈性鉴定

采用田间自然发病鉴定叶锈病, 在小麦成株期目测植株叶片上叶锈菌孢子堆占叶片总面积的百分比[23], 将每个家系叶锈病平均严重度分为1%、5%、10%、25%、40%、65%、80%和100%共8个等级。待感病亲本宁麦18的发病严重度达到80%时, 进行第一次严重度调查记录; 当宁麦18发病严重度达到100%时进行第二次调查, 此发病高峰时所达到的严重度称为最终严重度(final disease severity, FDS)。用SPSS 22软件进行统计分析和显著性比较。

1.3 SSR标记分析

苗期进行田间取样, 采用改良的CTAB法提取基因组DNA [24]。利用2423对SSR引物对两亲本基因组DNA进行扩增, 其中357对的扩增产物存在多态性, 从中筛选出条带清晰且均匀分布于21条染色体上的337个标记检测RIL群体112个家系。

为明确15个BC4F5回交株系的叶锈病抗性QTL组成, 利用与C615抗性QTL紧密连锁的7个SSR标记进行基因型检测。其中, QLr.njau-1BL的连锁标记有Xgwm259Xcfa2292Xwmc728, QLr.njau-3BS的连锁标记有Xbarc102Xwmc623, QLr.njau-4DL的连锁标记为Xgwm165, QLr.njau-6BS的连锁标记为Xmag1424

1.4 连锁图谱的构建及QTL定位

利用JoinMap 4.0软件构建连锁图谱, 按照Kosambi作图函数将标记间的重组交换率转化成遗传图距单位(cM) [25], 获得由238对SSR标记组成的48个连锁群, 总长1196.53 cM, 覆盖小麦21条染色体, 标记间平均距离为5.03 cM。结合田间叶锈病FDS表型数据, 使用Win QTLCart 2.5软件以复合区间作图法进行小麦成株期抗叶锈性QTL分析[26], LOD阈值设置为2.5。

2 结果与分析

2.1 亲本及RIL群体的抗病性表现

2016年总体发病较2017年重, 但发病趋势一致。抗病亲本C615成株期叶锈病FDS为5%, 而感病亲本宁麦18、扬麦13两年FDS均达到100%, 发病充分(图1)。两年的群体平均FDS分别为55.41%和46.72%, 家系FDS范围分别为5%~100%和3%~ 100%, 均呈连续性分布(表1图2), 符合数量遗传特征。

图1

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图1宁麦18 (A)、扬麦13 (B)和C615 (C)成株期叶锈病抗性表现

Fig.1Leaf rust resistance of Ningmai 18 (A), Yangmai 13 (B), and C615 (C) at adult stage



图2

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图2RIL群体2016年和2017年叶锈病最终严重度频次分布

亲本C615和宁麦18的最终病害严重度分别为5%和100%。
Fig. 2Frequency distributions of RILs in final leaf rust severity in 2016 and 2017

The final disease severities of parents were 5% for C615 and 100% for Ningmai 18.


Table 1
表1
表1亲本及其RIL群体叶锈病最终严重度表现
Table 1Final disease severity of leaf rust in parents and their RILs
年份
Year
亲本 ParentRIL群体 RIL population
C615
(%)
宁麦18
Ningmai 18 (%)
范围
Range (%)
均值
Mean (%)
标准差
SD (%)
变异系数
CV (%)
偏度
Skewness
峰度
Kurtosis
201651005-10055.4122.6738.52-0.61-0.53
201751003-10046.7222.4151.050.11-0.72

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2.2 叶锈病抗性QTL分析

两年共发现5个抗叶锈病QTL, 分布在5条染色体上(表2图3), 单个位点的表型贡献率为6.2%~15.7%, 除QLr.njau-2DS的抗性等位基因来自宁麦18 (加性效应为正)外, 其余4个QTL的抗性等位基因来自C615 (加性效应为负)。有3个QTL被两年重复检测到, 分别是QLr.njau-1BLQLr.njau-3BSQLr.njau-4DL, 效应较大且稳定; 另外2个QTL仅在一年被检测到, 并且表型贡献率较小(表2)。

Table 2
表2
表2叶锈病抗性QTL定位结果
Table 2QTL mapping result for leaf rust resistance
位点
QTL
标记区间
Marker interval
年份
Year
LOD加性效应
Additive effect
贡献率
Phenotypic variance explained (%)
QLr.njau-1BLXwmc728-Xbarc8020164.98-8.5515.7
Xgwm140-Xbarc8020174.34-9.6210.1
QLr.njau-2DSXgwm296-GPW408020172.546.586.2
QLr.njau-3BSXbarc102-Xwmc62320164.92-7.9513.5
20173.95-7.7810.9
QLr.njau-4DLXgwm165-Xcfd7120162.93-5.689.0
20173.52-6.878.2
QLr.njau-6BSXmag1424-Xmag1200.120164.18-5.309.2

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图3

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图3C615/宁麦18 RIL群体中检测到的叶锈病抗性QTL在染色体上的分布

Fig. 3Chromosomal locations of QTLs for leaf rust resistance in C615/Ningmai 18 RIL population



2.3 抗性QTL可靠性分析

根据QLr.njau-1BLQLr.njau-3BSQLr.njau- 4DL基因型, 将112个家系分为8种类型(表3), 分别用两年的FDS及其平均值进行类型间差异分析。有9个家系同时含有这3个QTL, 其发病严重度显著低于其他类型; 49个家系含有任意2个抗性QTL, 以同时含有QLr.njau-1BLQLr.njau- 3BS的家系抗病性较高; 含有1个抗性QTL的家系FDS普遍较高(50.6%~79.8%); 不含有抗性QTL的家系发病最重, 平均FDS在80%左右。说明聚合2个或2个以上抗性位点对降低叶锈病严重度效果明显。

Table 3
表3
表3C615/宁麦18 RIL群体中聚合不同稳定QTL家系的抗性效应
Table 3Resistance of the C615/Ningmai 18 RILs in different QTL-composition genotypes
QTL组成
QTL composition
家系数
Number of lines
最终严重度 Final disease severity (%)
20162017平均 Mean
None1485.6 a78.6 a82.1 a
QLr.njau-4DL1979.8 a70.4 a75.1 a
QLr.njau-3BS1065.7 b51.0 b58.4 b
QLr.njau-1BL1162.3 bc50.6 b56.5 bc
QLr.njau-1BL+QLr.njau-4DL1550.0 cd45.0 b47.5 bc
QLr.njau-3BS+QLr.njau-4DL1642.2 d40.0 bc41.1 cd
QLr.njau-1BL+QLr.njau-3BS1836.5 d26.9 c31.7 d
QLr.njau-1BL+QLr.njau-3BS+QLr.njau-4DL921.1 e11.1 d16.1 e
Different letters after data indicate significant difference among genotypes (P < 0.05).
数据后不同字母表示不同基因型间差异显著(P < 0.05)。

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此外, 利用与QLr.njau-3BS紧密连锁的分子标记检测所有家系显示, 没有该位点的家系平均FDS为65.2%, 变化范围是15%~100%; 而具有该位点的家系, 其平均FDS为36.9%, 变化范围是1%~80% (图4)。可见, 该QTL在两年自然发病的情况下均能明显降低病害严重度, 推测是一个稳定有效的抗病QTL。

图4

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图4含有和不含QLr.njau-3BS家系的最终病害严重度比较

最终病害严重度为两年平均值。RIL群体中含有和不含QLr.njau-3BS家系数分别是53和59。
Fig. 4Comparison of final disease severity between RILs with or without the QLr.njau-3BS locus

Final disease severity was the mean value of two years. In the RIL population, 53 and 59 lines were QLr.njau-3BS and non-QLr.njau-3BS genotype, respectively.


2.4 C615中的抗性QTL在15个BC4F5回交株系中的聚合

利用与C615抗性QTL紧密连锁的7个SSR标记检测15个表现抗病的BC4F5株系, 结果所有株系均含有C615的抗性等位基因, 其中3个株系聚合了4个来自C615的抗性位点, 仅有1个株系含单个抗性位点。13个株系为QLr.njau-1BL基因型, 且其中9个株系为QLr.njau-1BL+QLr.njau-3BS基因型。表明QLr.njau-1BL对降低叶锈病严重度发挥了重要作用, 且其与QLr.njau-3BS的聚合能有效提高抗病性(表4), 与RIL群体检测结果相吻合。

Table 4
表4
表415个回交株系的抗性QTL组成及叶锈病抗性
Table 4QTL composition of 15 backcross lines and their genotypes of leaf rust resistance
抗性QTL组成
Composition of resistance QTL
轮回亲本或株系
Recurrent parent or line
最终病害严重度 Final disease severity (%)
20162017
None扬麦13 Yangmai 13100100
QLr.njau-1BL+QLr.njau-3BS+QLr.njau-4DL+QLr.njau-6BSBL-15.05.0
BL-210.05.0
BL-310.08.3
QLr.njau-1BL+QLr.njau-3BS+QLr.njau-4DLBL-410.08.3
BL-510.010.0
BL-615.010.0
QLr.njau-1BL+ QLr.njau-4DL+QLr.njau-6BSBL-715.015.0
QLr.njau-1BL+QLr.njau-3BSBL-810.020.0
BL-915.015.0
BL-1020.020.0
QLr.njau-1BL + QLr.njau-6BSBL-1120.025.0
QLr.njau-3BS+QLr.njau-4DLBL-1225.020.0
BL-1325.025.0
QLr.njau-1BL + QLr.njau-6BSBL-1425.025.0
QLr.njau-1BLBL-1531.725.0

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3 讨论

3.1 定位结果与已知抗病基因的比较

在C615和宁麦18中共检测到5个抗叶锈病QTL, 其中位于1BL、3BS和4DL染色体上的QTL两年环境下均能被检测到, 是稳定的QTL。其中, 来源于抗病亲本C615的主效QTL QLr.njau-1BL, 标记区间为Xgwm140-Xbarc80, 根据系谱推测该位点很可能来自硬粒小麦CETA, 并与墨西哥小麦Pavon76中发现的Lr46/Yr29 (贡献率为55.8%) [27,28]非常接近, 但是QLr.njau-1BL在本试验中的表型贡献率较小。另外, C615 (系谱为SABUF/3/BCN//CETA/AE. SQUARROSA(895))与Pavon76 (系谱为Vicams71// Cianof67/Siete Cerrost66/3/Kalyansona/Bluebird)没有亲缘关系, 二者是否存在等位关系还需进一步验证。本研究发现QLr.njau-1BL在降低叶锈病严重度上有重要作用, 可作为有效的抗病基因应用于本麦区抗叶锈病育种。

Herrera等[14]和Hiebert等[15]Lr67定位在4DL染色体上, 并发现与其紧密连锁的标记Xgwm165Xgwm192。本研究定位的QLr.njau-4DL, 其标记区间为Xgwm165-Xcfd71, 两标记间的遗传距离为3 cM, 包含与Lr67紧密连锁的Xgwm165, 推测这个位点的抗性可能由Lr67提供, 其关系有待进一步验证。

位于3BS染色体上的QLr.njau-3BS, 标记区间为Xbarc102-Xwmc623, 贡献率为10.9%~13.5%。该染色体上已定位了4个抗叶锈QTL, 分别是QLr.sfrs-3B [29]Lr27 [30]QLr.cim-3BS [31]QLr.hebau-3BS [3]。其中, QLr.sfrs-3B位于着丝粒附近; Lr27为小种专化性抗病基因, 与QLr.cim-3BS均位于3B染色体短臂末端; QLr.hebau-3BS位于标记barc147gwm493之间, 与QLr.njau-3BS的遗传距离超过20 cM。因此推测, QLr.njau-3BS可能是一个新的抗叶锈病位点。此外, 本研究发现QLr.njau-3BSQLr.njau-1BL的聚合能有效降低病害严重度, 说明QLr.njau-3BS在抗叶锈育种中可能具有较大的应用潜力。本试验并没有对C615进行苗期鉴定, 因而QLr.njau-3BS是否为成株抗性基因值得进一步研究。

位于2DS和6BS染色体上的QTL只能在一年环境下被检测出来, 受环境因子影响较大, 但QLr.njau-6BS效应较大, 能解释表型变异的9.2%, 其有效性还需进一步验证。

3.2 QTL聚合效应分析

本研究发现QLr.njau-1BL对降低病害严重度有重要作用。当该位点单独存在时, 多数家系的FDS在50%以上, 但当它与其他QTL聚合, 特别是与QLr.njau-3BS聚合时, 病害严重度明显降低, 并且聚合包含QLr.njau-1BL的3个稳定QTL的家系具有较好的叶锈病抗性, FDS变化范围为11.1%~21.1%, 显著低于含有2个或2个以下抗性QTL家系的FDS。利用15个回交系检测, 也得到相似结论, 说明多个抗病基因的聚合能表现出基因的互作与累加效应。Singh等[32]报道Lr34/Yr18/Pm38基因单独存在时可致产量损失31%~52%, 而与3~4个微效基因聚合时产量损失小于10%。因而, 利用与Lr46QLr.njau- 3BS等抗病基因紧密连锁的分子标记对抗病品种或品系进行辅助选择, 从而将多个抗病基因聚合到同一品种中, 对于持久抗病品种的选育具有重要意义。

3.3 抗叶锈资源的选育

近3年小麦叶锈病危害急剧上升, 长江下游麦区逐渐成为易感区, 本麦区品种大多不具有叶锈病抗性, 如果长期使用单基因抗性势必面临强大的寄主选择压力, 抗病基因会逐步丧失抗性。因而有针对性地培育多基因聚合的持久抗锈品种显得十分迫切且重要。多年来, C615对小麦叶锈病一直保持着较好的抗性水平, 是一个很好的抗病资源。为改良感叶锈病栽培品种扬麦13, 本研究以C615为供体亲本构建回交群体, 结合分子标记检测结果, 获得了15个聚合不同抗病QTL且农艺性状均优于扬麦13的株系, 可作为优质、抗叶锈材料用于进一步的育种研究。同时该工作也表明C615作为抗病性强的种质资源, 在抗病品种的选育中具有较大的应用价值, 可利用本研究获得的与抗病基因紧密连锁的分子标记辅助选择, 将其用于现有品种的改良或培育新的持久抗叶锈品种。

4 结论

利用CIMMYT抗叶锈病种质C615与感病品种构建分离群体, 鉴定出5个抗病QTL, 其中位于1BL、4DL和3BS上的QTL效应较强, 对标记辅助选择抗叶锈病育种价值较大。3BS上的QLr.njau-3BS可能是一个新的抗病位点, 其与QLr.njau-1BL的聚合能明显降低病害严重度, 可作为有效的抗病基因用于育种研究。以C615为供体, 对扬麦13进行抗锈病性状回交改良, 获得了15个农艺性状优于扬麦13的抗病株系, 说明C615在抗锈品种的选育中具有较大的应用潜力, 可结合本研究中获得的与抗病基因紧密连锁的分子标记, 将其用于选育具有持久抗性的小麦新品种。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。


参考文献 原文顺序
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被引期刊影响因子

Khan M H, Bukhari A, Dar Z A, Rizvi S M . Status and strategies in breeding for rust resistance in wheat
Agric Sci, 2013,4:292-301

DOI:10.4236/as.2013.46042URL [本文引用: 1]
Wheat along with riceand maize is fulfilling half of the calories demands of the world. Global Wheatproduction has increased tremendously since green revolution in 1960鈥檚 andhelped in minimizing hunger and malnutrition. Developing countries, whichconsume 60% of the global wheat production, have shown a higher yield increasethan the developed countries in the past [1]. It was driven by the hunger prevalencein these countries and was attributable to the introduction of high yieldingand rusted resistant semi dwarf varieties developed under the collaborativeefforts of International and National research systems during the last 50years. Whereas, climate change and the emergence of new pests and diseases arethreatening the food sustainability. The evolution of new races of disease pathogenslike stem rust (Ug 99) is of serious concern. In order to feed the ever increasingpopulation we have to increase wheat production at the rate 1.6% which can beachieved by developing high yielding varieties having a good tolerance levelfor biotic and abiotic stresses.

Singh D, Mohler V, Park R F . Discovery, characterization and mapping of wheat leaf rust resistance gene
Lr71. Euphytica, 2013,190:131-136

DOI:10.1007/s10681-012-0786-xURL [本文引用: 1]
A temporarily designated gene LrARK12c (identified from spelt wheat cv. Altgold Rotkorn) with an intermediate low infection type was found effective against prevalent Australian Puccinia triticina ( Pt ) pathotypes. The gene was mapped to chromosome 1B between markers Xgwm18 and Xbarc187 , with linkage distances of 1.0 and 1.3聽cM, respectively. While it was not possible to assign a definitive chromosomal arm location to LrARK12c , it maps close to the centromere based on physical mapping of SSR marker loci using deletion lines. Other genes conferring resistance to Pt in chromosome 1B include Lr33, Lr44 and Lr46 . Genetic analysis showed that LrARK12c and Lr44 are genetically independent. Comparisons of markers linked to LrARK12c and Lr46 indicate that Lr46 should be well distal to the centromere. Lr33 is not effective in the seedling stage with Australian Pt pathotypes, therefore question of possible allelism of LrARK12c and Lr33 cannot be resolved using Australian Pt pathotypes. Genetic studies, chromosome mapping and allelism tests indicated that LrARK12c is a new and genetically independent leaf rust resistance locus, and hence it was designated Lr71 in accordance with the rules of wheat gene nomenclature.

Zhang P P, Yin G H, Zhou Y, Qi A Y, Gao F M, Xia X C, He Z H, Li Z F, Liu D Q , QTL mapping of adult-plant resistance to leaf rust in the wheat cross Zhou 8425B/Chinese Spring using high-density SNP markers
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Spielmeyer W, Mcintosh R J, Lagudah E . Powdery mildew resistance and Lr34/Yr18 genes for durable resistance to leaf and stripe rust cosegregate at a locus on the short arm of chromosome 7D of wheat
Theor Appl Genet, 2005,111:731-735

DOI:10.1007/s00122-005-2058-9URLPMID:15965649 [本文引用: 1]
Abstract The incorporation of effective and durable disease resistance is an important breeding objective for wheat improvement. The leaf rust resistance gene Lr34 and stripe rust resistance gene Yr18 are effective at the adult plant stage and have provided moderate levels of durable resistance to leaf rust caused by Puccinia triticina Eriks. and to stripe rust caused by Puccinia striiformis Westend. f. sp. tritici. These genes have not been separated by recombination and map to chromosome 7DS in wheat. In a population of 110 F(7) lines derived from a Thatcher x Thatcher isogenic line with Lr34/Yr18, field resistance to leaf rust conferred by Lr34 and to stripe rust resistance conferred by Yr18 cosegregated with adult plant resistance to powdery mildew caused by Blumeria graminis (DC) EO Speer f. sp. tritici. Lr34 and Yr18 were previously shown to be associated with enhanced stem rust resistance and tolerance to barley yellow dwarf virus infection. This chromosomal region in wheat has now been linked with resistance to five different pathogens. The Lr34/Yr18 phenotypes and associated powdery mildew resistance were mapped to a single locus flanked by microsatellite loci Xgwm1220 and Xgwm295 on chromosome 7DS.

Bariana H S, Hayden M J, Ahmed N U , BellJ A, Sharp P J, McIntosh R A. Mapping of durable adult plant and seedling resistances to stripe rust and stem rust diseases in wheat
Aust J Agric Res, 2001,52:1247-1255

DOI:10.1071/ar01040URL [本文引用: 1]
Doubled haploid populations of CD87/Katepwa, Cranbrook/Halberd, and Sunco/Tasman were assessed for seedling response to stem rust and stripe rust. The CD87/Katepwa population was also screened as adult plants in the field against stripe rust. The respective parents differed in presence or absence of various stem rust and stripe rust resistance genes. At least 4 resistance loci controlled adult plant resistance to stripe rust in the CD87/Katepwa population, and based on quantitative trait loci mapping results, two of these were contributed by CD87. Pedigree information indicated that these regions correspond to durable adult plant stripe rust resistance genes Yr18 and Yr29. Yr29 was mapped to the distal region of chromosome 1BL. The third gene, contributed by Katepwa, YrKat, was located in chromosome arm 2DS. Sr30 mapped distal to markers abg3 and P36/M61-170 in chromosome arm 5DL. Genes Yr7 and Pbc (completely linked with durable stem rust resistance gene Sr2) showed close associations with markers in chromosome arms 2BL and 3BS, respectively. A distally located genomic region in chromosome 6AS also affected the expression of Pbc. The temperature-sensitive stripe rust resistance gene, YrCK, carried by Sunco showed monogenic inheritance and was located in chromosome arm 2DS. Several markers showed complete association with Triticum timopheevi derived stem rust resistance gene Sr36. Microsatellite markers stm773 and gwm271A were validated on a set of wheat genotypes and were found to be diagnostic for the detection of Sr36. TheSr36-linked Xstm773 allele showed better amplification than the Sr36-linked Xgwm271A allele. These markers could be used for marker assisted identification of Sr36 in breeding populations.

Suenaga K, Singh R P, Huertaespino J, William H M . Microsatellite markers for genes Lr34/Yr18 and other quantitative trait loci for leaf rust and stripe rust resistance in bread wheat
. Phytopathology, 2003,93:881-890

DOI:10.1094/PHYTO.2003.93.7.881URLPMID:18943170 [本文引用: 1]
ABSTRACT Leaf rust and stripe rust, caused by Puccinia triticina and P. striiformis, respectively, are important diseases of wheat in many countries. In this study we sought to identify molecular markers for adult plant resistance genes that could aid in incorporating such durable resistance into wheat. We used a doubled haploid population from a Japanese cv. Fukuho-komugi x Israeli wheat Oligoculm cross that had segregated for resistance to leaf rust and stripe rust in field trials. Joint and/or single-year analyses by composite interval mapping identified two quantitative trait loci (QTL) that reduced leaf rust severity and up to 11 and 7 QTLs that might have influenced stripe rust severity and infection type, respectively. Four common QTLs reduced stripe rust severity and infection type. Except for a QTL on chromosome 7DS, no common QTL for leaf rust and stripe rust was detected. QTL-7DS derived from 'Fukuho-komugi' had the largest effect on both leaf rust and stripe rust severities, possibly due to linked resistance genes Lr34/Yr18. The microsatellite locus Xgwm295.1, located almost at the peak of the likelihood ratio contours for both leaf and stripe rust severity, was closest to Lr34/Yr18. QTLs located on 1BL for leaf rust severity and 3BS for stripe rust infection type were derived from 'Oligoculm' and considered to be due to genes Lr46 and Yr30, respectively. Most of the remaining QTLs for stripe rust severity or infection type had smaller effects. Our results indicate there is significant diversity for genes that have minor effects on stripe rust resistance, and that successful detection of these QTLs by molecular markers should be helpful both for characterizing wheat genotypes effectively and combining such resistance genes.

Schnurbusch T, Paillard S, Schori A, Messmer M, Schachermayr G, Winzeler M, Keller B . Dissection of quantitative and durable leaf rust resistance in Swiss winter wheat reveals a major resistance QTL in the Lr34 chromosomal region
Theor Appl Genet, 2004,108:477-484

[本文引用: 1]

Schnurbusch T, Bossolini E, Messmer M, Keller B . Tagging and validation of a major quantitative trait locus for leaf rust resistance and leaf tip necrosis in winter wheat cultivar Forno
Phytopathology, 2004,94:1036-1041

DOI:10.1094/PHYTO.2004.94.10.1036URLPMID:18943790 [本文引用: 1]
Abstract ABSTRACT A major leaf rust (Puccinia triticina) resistance quantitative trait locus (QTL) (QLrP.sfr-7DS) previously has been described on chromosome 7DS in the winter wheat (Triticum aestivum) cv. Forno. It was detected in a population of single-seed descent (SSD) lines derived from the cross Arina x Forno. QLrP.sfr-7DS conferred a durable and slow-rusting resistance phenotype, co-segregated with a QTL for leaf tip necrosis (LTN) and was mapped close to Xgwm295 at a very similar location as the adult plant leaf rust resistance gene Lr34 found in some spring wheat lines. Here, we describe the validation of this QTL by mapping it to the same chromosomal region close to Xgwm295 on chromosome 7DS in a population of SSD lines from the winter wheat x spelt (T. spelta) cross Forno x Oberkulmer. In both populations, the log of the likelihood ratio curves for leaf rust resistance and LTN peaked at identical or very similar locations, indicating that both traits are due to the same gene. We have improved the genetic map in the target region of QLrP.sfr-7DS using microsatellite and expressed sequence tag (EST) markers. Two EST loci (Xsfr.BF473324 and Xsfr.BE493812) define a genetic interval of 7.6 centimorgans containing QLrP.sfr-7DS, a considerably more precise genetic location for this QTL than previously described both in spring and winter wheat. The identified genetic interval is physically located in the distal 39% of chromosome 7DS. Single-marker analysis identified Xsfr.BF473324 and Xgwm1220 as the most informative loci for QLrP.sfr-7DS and QLtn.sfr-7DS. In the rice genome, the two ESTs flanking the QLrP.sfr-7DS/QLtn.sfr-7DS chromosomal segment in wheat are conserved on chromosome 6S in a region colinear with wheat chromosome 7DS. There, they define a physical region of three rice bacterial artificial chromosomes spanning approximately 300 kb.

Zhang P P, Zhang Y Q, Han B, Gao P, Liu L, Xing L, Li Z F, Liu D Q . QTL mapping of adult-plant resistance genes to leaf rust and stripe rust in CIMMYT wheat Kingbird and PBW343
J Henan Agric Sci, 2014,54:1907-1925

URL [本文引用: 1]
In order to detect and map the QTL to leaf rust and stripe rust in CIMMYT wheat lines Kingbird and PBW343,a total of 181 recombinant inbred lines(RILs)derived from the cross Kingbird and PBW343 were planted in Mexico to score disease severity of leaf rust and stripe rust to obtain the phenotypic data.A total of 406 DArT markers with polymorphism between parents were used to test the whole 181 RILs to obtain the genotypic data.Phenotypic data and genotypic data were used to map QTL by softwares Map manager QTXb20 and QTL IciMapping 3.2with composite interval mapping strategy.The results showed that a total of three QTLs for leaf rust resistance and one QTL for stripe rust resistance were detected on chromosome 1A,7DS,5BL and3 BS,respectively,and explaining 5.57%,11.94%,5.11%and 9.03%of the phenotypic variance.The QTLs on 7DS and 3BS were from Kingbird,and the other two QTLs were detected in PBW343.These QTLs enrich the wheat slow rusting gene pool and will give theory and technology support for wheat resistance breeding.

William M, Singh R P, Huerta-Espino J, Islas S O, Hoisington D . Molecular marker mapping of leaf rust resistance gene Lr46 and its association with stripe rust resistance gene Yr29 in wheat
Phytopathology, 2003,93:153-159

DOI:10.1094/PHYTO.2003.93.2.153URLPMID:18943129 [本文引用: 1]
ABSTRACT Leaf and stripe rusts, caused by Puccinia triticina and P. striiformis, respectively, are globally important fungal diseases of wheat that cause significant annual yield losses. A gene that confers slow rusting resistance to leaf rust, designated as Lr46, has recently been located on wheat chromosome 1B. The objectives of our study were to establish the precise genomic location of gene Lr46 using molecular approaches and to determine if there was an association of this locus with adult plant resistance to stripe rust. A population of 146 F(5) and F(6) lines produced from the cross of susceptible 'Avocet S' with resistant 'Pavon 76' was developed and classified for leaf rust and stripe rust severity for three seasons. Using patterns of segregation for the two diseases, we estimated that at least two genes with additive effects conferred resistance to leaf rust and three to four genes conferred resistance to stripe rust. Bulked segregant analysis and linkage mapping using amplified fragment length polymorphisms with the 'Avocet' x 'Pavon 76' population, F(3) progeny lines of a single chromosome recombinant line population from the cross 'Lalbahadur' x 'Lalbahadur (Pavon 1B)', and the International Triticeae Mapping Initiative population established the genomic location of Lr46 at the distal end of the long arm of wheat chromosome 1B. A gene that is closely linked to Lr46 and confers moderate levels of adult plant resistance to stripe rust is identified and designated as Yr29.

Singh R P, Mujeebkazi A, Huertaespino J . Lr46: a gene conferring slow-rusting resistance to leaf rust in wheat. Phytopathology, 1998,88:890-894
[本文引用: 1]

Lan C, Basnet B R, Singh R P, Huerta-Espino J , Herrera-Foessel S A, Ren Y, Randhawa M S. Genetic analysis and mapping of adult plant resistance loci to leaf rust in durum wheat cultivar Bairds
Theor Appl Genet, 2017,130:609-619

DOI:10.1007/s00122-016-2839-3URLPMID:28004134 [本文引用: 1]
New leaf rust adult plant resistance (APR) QTLQLr.cim-6BLwas mapped and confirmed the known pleotropic APR geneLr46effect on leaf rust in durum wheat line Bairds. CIMMYT-derived durum wheat line Baird

Herrera-Foessel S A, Lagudah E S, Huertaespino J, Hayden M J, Bariana H S . New slow-rusting leaf rust and stripe rust resistance genes Lr67 and Yr46 in wheat are pleiotropic or closely linked
Theor Appl Genet, 2011,122:239-249

DOI:10.1007/s00122-010-1439-xURLPMID:20848270 [本文引用: 2]
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, F and F populations from photoperiod-insensitive F 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 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 脙鈥 RL6077 F 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.

Hiebert C W, Thomas J B , McCallum B D, Humphreys D G, De Pauw R M, Hayden M J, Mago R, Schnippenkoetter W, Spielmeyer W. An introgression on wheat chromosome 4DL in RL6077 (Thatcher*6/PI 250413) confers adult plant resistance to stripe rust and leaf rust ( Lr67)
Theor Appl Genet, 2010,121:1083-1091

DOI:10.1007/s00122-010-1373-yURLPMID:20552325 [本文引用: 2]
Adult plant resistance (APR) to leaf rust and stripe rust derived from the wheat (Triticum aestivum L.) line PI250413 was previously identified in RL6077 (=Thatcher*6/PI250413). The leaf rust resistance gene in RL6077 is phenotypically similar to Lr34 which is located on chromosome 7D. It was previously hypothesized that the gene in RL6077 could be Lr34 translocated to another chromosome. Hybrids between RL6077 and Thatcher and between RL6077 and 7DS and 7DL ditelocentric stocks were examined for first meiotic metaphase pairing. RL6077 formed chain quadrivalents and trivalents relative to Thatcher and Chinese Spring; however both 7D telocentrics paired only as heteromorphic bivalents and never with the multivalents. Thus, chromosome 7D is not involved in any translocation carried by RL6077. A genome-wide scan of SSR markers detected an introgression from chromosome 4D of PI250413 transferred to RL6077 through five cycles of backcrossing to Thatcher. Haplotype analysis of lines from crosses of Thatcher x RL6077 and RL6058 (Thatcher*6/PI58548) x RL6077 showed highly significant associations between introgressed markers (including SSR marker cfd71) and leaf rust resistance. In a separate RL6077-derived population, APR to stripe rust was also tightly linked with cfd71 on chromosome 4DL. An allele survey of linked SSR markers cfd71 and cfd23 on a set of 247 wheat lines from diverse origins indicated that these markers can be used to select for the donor segment in most wheat backgrounds. Comparison of RL6077 with Thatcher in field trials showed no effect of the APR gene on important agronomic or quality traits. Since no other known Lr genes exist on chromosome 4DL, the APR gene in RL6077 has been assigned the name Lr67.Digital Object Identifier http://dx.doi.org/10.1007/s00122-010-1373-y

Herrera-Foessel S A, Singh R P, Huerta-Espino J, Rosewarne G M, Periyannan S K, Viccars L, Calvo-Salazar V, Lan C, Lagudah E S . Lr68: a new gene conferring slow rusting resistance to leaf rust in wheat
Theor Appl Genet, 2012,124:1475-1486

DOI:10.1007/s00122-012-1802-1URLPMID:22297565 [本文引用: 1]
The common wheat cultivar Parula possesses a high level of slow rusting, adult plant resistance (APR) to all three rust diseases of wheat. Previous mapping studies using an Avocet- YrA /Parula recombinant inbred line (RIL) population showed that APR to leaf rust ( Puccinia triticina ) in Parula is governed by at least three independent slow rusting resistance genes: Lr34 on 7DS, Lr46 on 1BL, and a previously unknown gene on 7BL. The use of field rust reaction and flanking markers identified two F 6 RILs, Arula1 and Arula2, from the above population that lacked Lr34 and Lr46 but carried the leaf rust resistance gene in 7BL, hereby designated Lr68 . Arula1 and Arula2 were crossed with Apav, a highly susceptible line from the cross Avocet- YrA /Pavon 76, and 396 F 4 -derived F 5 RILs were developed for mapping Lr68 . The RILs were phenotyped for leaf rust resistance for over 2 years in Ciudad Obregon, Mexico, with a mixture of P. triticina races MBJ/SP and MCJ/SP. Close genetic linkages with several DNA markers on 7BL were established using 367 RILs; Psy1 - 1 and gwm146 flanked Lr68 and were estimated at 0.5 and 0.6 cM, respectively. The relationship between Lr68 and the race-specific seedling resistance gene Lr14b , located in the same region and present in Parula, Arula1 and Arula2, was investigated by evaluating the RILs with Lr14b -avirulent P. triticina race TCT/QB in the greenhouse. Although Lr14b and Lr68 homozygous recombinants in repulsion were not identified in RILs, -irradiation-induced deletion stocks that lacked Lr68 but possessed Lr14b showed that Lr68 and Lr14b are different loci. Flanking DNA markers that are tightly linked to Lr68 in a wide array of genotypes can be utilized for selection of APR to leaf rust.

Li Z F, Lan C X, He Z H, Singh R P, Rosewarne M R, Chen X M, Xia X C . Overview and application of QTL for adult plant resistance to leaf rust and powdery mildew in wheat
Crop Sci, 2014,54:1907-1925

DOI:10.2135/cropsci2014.02.0162URL [本文引用: 1]
ABSTRACT Leaf rust and powdery mildew, caused by Puccinia triticina and Blumeria graminis f. sp. tritici, respectively, are widespread fungal diseases of wheat (Triticum aestivum L.). Development of cultivars with durable resistance is crucially important for global wheat production. This paper reviews the progress of genetic study and application of adult plant resistance (APR) to wheat leaf rust and powdery mildew. Eighty leaf rust and 119 powdery mildew APR quantitative trait loci (QTL) have been reported on 16 and 21 chromosomes, respectively, in over 50 publications during the last 15 yr. More important, we found 11 loci located on chromosomes 1BS, 1BL, 2AL, 2BS (2), 2DL, 4DL, 5BL, 6AL, 7BL, and 7DS showing pleiotropic effects on resistance to leaf rust, stripe rust, and powdery mildew. Among these, QTL on chromosomes 1BL, 4DL, and 7DS also correlate with leaf tip necrosis. Fine mapping and cloning of these QTL will be achieved with the advent of cheaper high-throughput genotyping technologies. Germplasm carrying these potential resistance genes will be useful for developing cultivars with durable multidisease resistance. In addition to its non-NBS-LRR (nucleotide binding site-leucine rich repeat) structure, the senescence-like processes induced by Lr34 could be the reason for durability of resistance; however, more information is needed for a full understanding of the molecular mechanism related to durability. Adult plant resistance genes have been used by CIMMYT for more than 30 yr and have also been transferred to many Chinese wheat varieties through shuttle breeding.

Rosewarne G M, Singh R P, Huerta-Espino J, Rebetzke G J . Quantitative trait loci for slow-rusting resistance in wheat to leaf rust and stripe rust identified with multi-environment analysis
Theor Appl Genet, 2008,116:1027-1034

DOI:10.1007/s00122-008-0736-0URLPMID:18335201 [本文引用: 1]
Rust diseases are a major cause of yield loss in wheat worldwide, and are often controlled through the incorporation of resistance genes using conventional phenotypic selection methods. Slow-rusting resistance genes are expressed quantitatively and are typically small in genetic effect thereby requiring multiple genes to provide adequate protection against pathogens. These effects are valuable and are generally considered to confer durable resistance. Therefore an understanding of the chromosomal locations of such genes and their biological effects are important in order to ensure they are suitably deployed in elite germplasm. Attila is an important wheat grown throughout the world and is used as a slow-rusting donor in international spring wheat breeding programs. This study identified chromosomal regions associated with leaf rust and stripe rust resistances in a cross between Attila and a susceptible parent, Avocet-S, evaluated over 3 years in the field. Genotypic variation for both rusts was large and repeatable with line-mean heritabilities of 94% for leaf rust resistance and 87% for stripe rust. Three loci, including Lr46/Yr29 on chromosome 1BL, were shown to provide resistance to leaf rust whereas six loci with small effects conferred stripe rust resistance, with a seventh locus having an effect only by epistasis. Disease scoring over three different years enabled inferences to be made relating to stripe rust pathogen strains that predominated in different years.

Crossa J, Burgue?o J, Dreisigacker S, Vargas M , Herrera-Foessel S A, Lillemo M, Singh R P, Trethowan R, Warburton M, Franco J, Reynolds M, Crouch J H, Ortiz R. Association analysis of historicalbread wheat germplasm using additive genetic covariance of relatives and population structure
Genetics, 2007,177:1889-1913

DOI:10.1534/genetics.107.078659URLPMID:17947425 [本文引用: 1]
Linkage disequilibrium can be used for identifying associations between traits of interest and genetic markers. This study used mapped diversity array technology (DArT) markers to find associations with resistance to stem rust, leaf rust, yellow rust, and powdery mildew, plus grain yield in five historical wheat international multienvironment trials from the International Maize and Wheat Improvement Center (CIMMYT). Two linear mixed models were used to assess marker-trait associations incorporating information on population structure and covariance between relatives. An integrated map containing 813 DArT markers and 831 other markers was constructed. Several linkage disequilibrium clusters bearing multiple host plant resistance genes were found. Most of the associated markers were found in genomic regions where previous reports had found genes or quantitative trait loci (QTL) influencing the same traits, providing an independent validation of this approach. In addition, many new chromosome regions for disease resistance and grain yield were identified in the wheat genome. Phenotyping across up to 60 environments and years allowed modeling of genotype x environment interaction, thereby making possible the identification of markers contributing to both additive and additive x additive interaction effects of traits.

Ren Y, Li Z, He Z H , W Ling, Bai B, Lan C X, Wang C F, Zhou G, Zhu H Z, Xia X C. QTL mapping of adult-plant resistances to stripe rust and leaf rust in Chinese wheat cultivar Bainong 64
Theor Appl Genet, 2012,125:1253-1262

DOI:10.1007/s00122-012-1910-yURLPMID:22806327 [本文引用: 1]
Abstract 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.

Zhou H X, Xia X C, He Z H, LI X, Wang C F, Li Z F, Liu D Q . Molecular mapping of leaf rust resistance gene LrNJ97 in Chinese wheat line Neijiang 977671
Theor Appl Genet, 2013,126:2141-2147

DOI:10.1007/s00122-013-2124-7URLPMID:23689746 [本文引用: 1]
Neijiang 977671 and 19 near-isogenic lines with known leaf rust resistance genes were inoculated with 12 pathotypes of Puccinia triticina for postulation of leaf rust resistance genes effective at the seedling stage. The reaction pattern of Neijiang 977671 differed from those of the lines with known leaf rust resistance genes used in the test, indicating that Neijiang 977671 may carry a new leaf rust resistance gene(s). With the objective of identifying and mapping the new gene for resistance to leaf rust, F 1 and F 2 plants, and F 2:3 families, from Neijiang 977671脳Zhengzhou 5389 (susceptible) were inoculated with Chinese P. triticina pathotype FHNQ in the greenhouse. Results from the F 2 and F 2:3 populations indicated that a single dominant gene, temporarily designated LrNJ97 , conferred resistance. In order to identify other possible genes in Neijiang 977671 other eight P . triticina pathotypes avirulent on Neijiang 977671 were used to inoculate 25 F 2:3 families. The results showed that at least three leaf rust resistance genes were deduced in Neijiang 977671. Bulked segregant analysis was performed on equal amounts of genomic DNA from 20 resistant and 20 susceptible F 2 plants. SSR markers polymorphic between the resistant and susceptible bulks were used to analyze the F 2:3 families. LrNJ97 was linked to five SSR loci on chromosome 2BL. The two closest flanking SSR loci were Xwmc317 and Xbarc159 at genetic distances of 4.2 and 2.2cM, respectively. At present two designated genes ( Lr50 and Lr58 ) are located on chromosome 2BL. In the seedling tests, the reaction pattern of LrNJ97 was different from that of Lr50 . Lr50 and Lr58 were derived from T. armeniacum and Ae. triuncialis , respectively, whereas according to the pedigree of Neijiang 977671 LrNJ97 is from common wheat. Although seeds of lines with Lr58 were not available, it was concluded that LrNJ97 is likely to be a new leaf rust resistance gene.

彭红, 吕国强, 王江蓉 . 河南省2015 年小麦主要病害发生特点及原因分析
中国植保导刊, 2016,36(4):29-33

[本文引用: 1]

Peng H, Lyu G Q, Wang J R . Analysis on the characteristics and causes of main diseases of wheat in Henan province in 2015
China Plant Prot, 2016,36(4):29-33 (in Chinese)

[本文引用: 1]

Peterson R F, Campbell A B, Hannah A E . A diagrammatic scale for estimating rust intensity on leaves and stems of cereals
Can J Res, 1948,26:496-500

[本文引用: 1]

张晓祥, 王玲, 寿路路 . 一种快速提取小麦基因组DNA的改良CTAB方法
中国农学通报, 2012,28(36):46-49

[本文引用: 1]

Zhang X X, Wang L, Shou L L . A rapid modified CTAB method of extracting genomic DNA from wheat leaf
Chin Agric Sci Bull, 2012,28(36):46-49 (in Chinese with English abstract)

[本文引用: 1]

张利军, 李在峰 , Morten L, 夏先春, 刘大群, 杨文香, 罗家传, 王海燕. CIMMYT小麦品种Saar的叶锈成株抗性QTL分析
中国农业科学, 2009,42:388-397

DOI:10.3864/j.issn.0578-1752.2009.02.002URL [本文引用: 1]
[目的]小麦品种Saar由CIMMYT育成,在欧洲、亚洲和南美洲对小麦叶锈、条锈和白粉病均表现出很高的成株抗性,发掘其成株抗叶锈QTL对于选育持久抗锈品种有重要作用.[方法]以Avocet与Saar杂交的109个F6代重组自交系为材料,利用142个SSR标记和209 DArT(Diversity Arrays Technology)标记构建连锁图,对Saar和Avocet的成株抗性进行QTL分析.试验材料于2006-2007年度种植在河北保定和河南新乡两个试验点,调查各个家系对叶锈病的成株抗性.[结果]由351个位点组成的遗传连锁图,覆盖小麦21个连锁群,全长3 083 cM.采用复合区间作图法进行叶锈成株抗性的QTL分析,在1BL、2DS、5BL、6AL和7DS染色体上发现了5个抗叶锈病QTL,分别解释4.5%~6.4%、12.2%~12.5%,4.9%~11.2%、4.9%~7.8%和14.0%~67.6%的表型变异.[结论]叶锈成株抗性基因及其紧密连锁分子标记的发掘,将为小麦抗叶锈病育种的分子标记辅助选择(MAS)提供理论和技术支持.
Zhang L J, Li Z F, Morten L, Xia X C, Liu D Q, Yang W X, Luo J C, Wang H Y . QTL Mapping for adult-plant resistance to leaf rust in CIMMYT wheat cultivar Saar
Sci Agric Sin, 2009,42:388-397 (in Chinese with English abstract)

DOI:10.3864/j.issn.0578-1752.2009.02.002URL [本文引用: 1]
[目的]小麦品种Saar由CIMMYT育成,在欧洲、亚洲和南美洲对小麦叶锈、条锈和白粉病均表现出很高的成株抗性,发掘其成株抗叶锈QTL对于选育持久抗锈品种有重要作用.[方法]以Avocet与Saar杂交的109个F6代重组自交系为材料,利用142个SSR标记和209 DArT(Diversity Arrays Technology)标记构建连锁图,对Saar和Avocet的成株抗性进行QTL分析.试验材料于2006-2007年度种植在河北保定和河南新乡两个试验点,调查各个家系对叶锈病的成株抗性.[结果]由351个位点组成的遗传连锁图,覆盖小麦21个连锁群,全长3 083 cM.采用复合区间作图法进行叶锈成株抗性的QTL分析,在1BL、2DS、5BL、6AL和7DS染色体上发现了5个抗叶锈病QTL,分别解释4.5%~6.4%、12.2%~12.5%,4.9%~11.2%、4.9%~7.8%和14.0%~67.6%的表型变异.[结论]叶锈成株抗性基因及其紧密连锁分子标记的发掘,将为小麦抗叶锈病育种的分子标记辅助选择(MAS)提供理论和技术支持.

Li H, Ribaut J M, Li Z, Wang J . Inclusive composite interval mapping (ICIM) for digenic epistasis of quantitative traits in biparental populations
Theor Appl Genet, 2008,116:243-260

DOI:10.1007/s00122-007-0663-5URL [本文引用: 1]

William H M, Singh R P, Huertaespino J, Palacios G, Suenaga K . Characterization of genetic loci conferring adult plant resistance to leaf rust and stripe rust in spring wheat
Genome, 2006,49:977-990

DOI:10.1139/g06-052URL [本文引用: 1]

Rosewarne G M, Singh R P, Huerta-Espino J , Herrera-Foessel S A, Forrest K L, Hayden M J, Rebetzke G J. Theor Appl Genet, 2012,124:1283-1294
DOI:10.1007/s00122-012-1786-xURLPMID:22274764 [本文引用: 1]
Leaf rust and stripe rust are important diseases of wheat world-wide and deployment of cultivars with genetic resistance is an effective and environmentally sound control method. The use of minor, additive genes conferring adult plant resistance (APR) has been shown to provide resistance that is durable. The wheat cultivar ‘Pastor’ originated from the CIMMYT breeding program that focuses on minor gene-based APR to both diseases by selecting and advancing generations alternately under leaf rust and stripe rust pressures. As a consequence, Pastor has good resistance to both rusts and was used as the resistant parent to develop a mapping population by crossing with the susceptible ‘Avocet’. All 148 F 5 recombinant inbred lines were evaluated under artificially inoculated epidemic environments for leaf rust (3 environments) and stripe rust (4 environments, 2 of which represent two evaluation dates in final year due to the late build-up of a new race virulent to Yr31 ) in Mexico. Map construction and QTL analysis were completed with 223 polymorphic markers on 84 randomly selected lines in the population. Pastor contributed Yr31 , a moderately effective race-specific gene for stripe rust resistance, which was overcome during this study, and this was clearly shown in the statistical analysis. Linked or pleiotropic chromosomal regions contributing to resistance against both pathogens included Lr46/Yr29 on 1BL, the Yr31 region on 2BS, and additional minor genes on 5A, 6B and 7BL. Other minor genes for leaf rust resistance were located on 1B, 2A and 2D and for stripe rust on 1AL, 1B, 3A, 3B, 4D, 6A, 7AS and 7AL. The 1AL, 1BS and 7AL QTLs are in regions that were not identified previously as having QTLs for stripe rust resistance. The development of uniform and severe epidemics facilitated excellent phenotyping, and when combined with multi-environment analysis, resulted in the relatively large number of QTLs identified in this study.

Messmer M M, Seyfarth R, Keller M, Schachermayr G, Winzeler M, Zanetti S, Feuillet C, Keller B . Genetic analysis of durable leaf rust resistance in winter wheat
Theor Appl Genet, 2000,100:419-431

DOI:10.1007/s001220050055URL [本文引用: 1]
Quantitative resistance that delays the epidemic development of leaf rust in wheat is an important source for durable resistance breeding. The Swiss winter wheat variety ’Forno’ shows a high level of quantitative resistance against leaf rust. This resistance has been effective for more than 10 years and can therefore be considered to be durable. In order to map quantitative trait loci (QTL) for durable leaf rust resistance we analysed 204 F 5 recombinant inbred lines (RILs) of the cross between the winter wheat ’Forno’ and the winter spelt ’Oberkulmer’ for their level of leaf rust resistance (LR) and leaf tip necrosis (LTN) in four different environments. Both traits showed a continuous distribution and were significantly correlated ( r =610.5). Across environments we detected 8 QTL for leaf rust resistance (6 inherited from ’Forno’) and 10 QTL for the quantitative expression of LTN (6 inherited from ’Forno’). Of the 6 QTL responsible for the durable leaf rust resistance of ’Forno’, 1 major QTL coincided with a thaumatin locus on 7BL explaining 35% of the phenotypic variance. Four QTL for LR coincided with QTL for LTN. At these loci the alleles of ’Forno’ increased the level of resistance as well as the extent of LTN, indicating pleiotropy.

Singh R P, Mcintosh R A . Complementary genes for reaction to Puccinia recondita tritici in Triticum aestivum: I. Genetic and linkage studies
Genome, 2011,26:723-735

[本文引用: 1]

Calvo-Salazar, SinghR P, Huerta-Espino J, Cruz-Izquierdo S, Lobato-Ortiz, Sandoval-Islas S, Vargas-Hernandez M, German S, Silva P, Basnet B R, Lan C X, Herrera-Foessel S A. Genetic analysis of resistance to leaf rust and yellow rust in spring wheat cultivar Kenya Kongoni
Plant Dis, 2015,99:1153-1159

DOI:10.1094/PDIS-07-14-0718-REURL [本文引用: 1]
Abstract The Kenyan wheat (Triticum aestivum L.) ‘Kenya Kongoni’ exhibits high levels of adult plant resistance (APR) to leaf rust (LR) and yellow rust (YR). We determined the genomic regions associated with LR and YR resistance in a population of 148 recombinant inbred lines generated from a cross between ‘Avocet-YrA’ and Kenya Kongoni. Field experiments to characterize APR to LR and YR were conducted in four and two Mexican or Uruguayan environments, respectively. A linkage map was constructed with 438 diversity arrays technology and 16 simplesequence repeat markers by JoinMap 4.1 software. Genetic analyses showed that resistance to both rusts was determined by four to five APR genes, including Lr46/Yr29 and Sr2/Lr27/Yr30. Quantitative trait loci (QTL) analysis indicated that pleiotropic APR loci QYLr.cim-1BL corresponding to Lr46/Yr29 and QYLr.cim-7BL that is a putative novel QTL accounted for 5 to 57% and 12 to 35% of the phenotypic variation for resistance to LR and YR, respectively. These loci, in combination with another three LR QTL and two YR QTL, respectively, conferred high levels of resistance to both LR and YR in wheat under Mexican and Uruguayan environments. Among other detected QTL, QLr.cim-1DS, QLr.cim-2BL, and QYLr.icm-7BL may be new loci for APR to both rusts in common wheat.

Singh R P, Huerta-Espino J, William H M . Genetics and breeding for durable resistance to leaf and stripe rusts in wheat
Turkish J Agric For, 2005,29:121-127

URL [本文引用: 1]
ABSTRACT Singh, R.P., J. Huerta-Espino and H.M. William. 2004. Genetics and breeding for durable resistance to leaf and stripe rusts in wheat. Turk. J. Agric. For. 28: xxx-xxx. Yellow (or stripe) and leaf (or brown) rusts, caused by Puccinia striiformis and P. triticina, respectively, are important diseases of wheat worldwide. Growing resistant cultivars is the most economical and environmentally safe control measure and has no cost to growers. Wheat (Triticum aestivum) cultivars that have remained resistant for a long time, or in other words carry durable or race-nonspecific resistance, are known to occur. Inheritance of resistance indicates that these cultivars often carry a few slow rusting genes that have small-to-intermediate, but additive, effects. Our genetic studies show that a high level of resistance (approaching immunity) to both rusts could be achieved by accumulating from 4 to 5 such genes. We recommend that a group of winter and spring wheat cultivars known to carry adequate levels of durable resistance to yellow and/or leaf rusts are assembled and further evaluated in the region to identify those cultivars that show resistance stability. Resistance from these cultivars should then be transferred in a planned manner to the susceptible but locally adapted cultivars through a 'Single Backcross Breeding Approach', that allows the simultaneous accumulation of desired number of slow rusting genes with increased grain yield potential and other traits.
相关话题/基因 材料 遗传 叶锈病 鉴定