以关联分析发掘小麦整穗发芽抗性基因分子标记

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朱玉磊, 王升星, 赵良侠, 张德新, 胡建帮, 曹雪连, 杨亚杰, 常成, 马传喜, 张海萍. 以关联分析发掘小麦整穗发芽抗性基因分子标记. 作物学报, 2014, 40(10): 1725-1732[ZHU Yu-Lei, WANG Sheng-Xing, ZHAO Liang-Xia, ZHANG De-Xin, HU Jian-Bang, CAO Xue-Lian, YANG Ya-Jie, CHANG Cheng, MA Chuan-Xi, ZHANG Hai-Ping. Exploring Molecular Markers of Preharvest Sprouting Resistance Gene Using Wheat Intact Spikes by Association Analysis. Acta Agronomica Sinica, 2014, 40(10): 1725-1732] 复制到剪切板

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以关联分析发掘小麦整穗发芽抗性基因分子标记

朱玉磊, 王升星, 赵良侠, 张德新, 胡建帮, 曹雪连, 杨亚杰, 常成, 马传喜, 张海萍^*

安徽农业大学农学院 / 农业部黄淮南部小麦生物学与遗传育种重点实验室, 安徽合肥230036

^*通讯作者(Corresponding author): 张海萍, E-mail:zhhp20@163.com

收稿日期:2014-07-06

基金:本研究由国家现代农业产业技术体系建设专项, 安徽省小麦产业技术体系建设专项和教育部高等学校博士学科点专项(20113418120004)资助;

摘要

利用分布于小麦全基因组的181对分子标记, 分析264份自然群体的基因型, 采用TASSLE软件的GLM和MLM模型检测与整穗发芽抗性紧密关联的标记位点, 发掘相关位点内的优异等位变异。在2012年和2013年室内整穗发芽率、2013年田间自然降雨整穗发芽率3个环境中, 共关联到20个显著位点(P<0.05), 分布于小麦染色体1AS、2DS、3AS、3BL、4AL、5AS、5BL、6BS、6DS、7AL和7BL上。分别位于2DS和7BL上的分子标记gwm102和barc340同时在3个环境下关联到, 属于稳定的抗性位点; 另有6个标记位点同时在2个环境下关联到; 其余12个标记位点仅在1个环境下关联到。位于7BL上的barc340标记位点为一新报道位点。从重复关联的8个标记位点内共检测出10种优异等位变异。barc28-229bp和barc28-217bp对提高整穗发芽抗性效应最显著, 主要分布在地方品种中(如遂宁坨坨麦等), 而gwm102-142bp和barc186-199bp效应虽然相对较小, 但多分布在推广品种中(如扬麦158等), 有利于穗发芽抗性分子育种的直接应用。

关键词: 小麦; 穗发芽; 休眠; 分子标记; 关联分析

Exploring Molecular Markers of Preharvest Sprouting Resistance Gene Using Wheat Intact Spikes by Association Analysis

ZHU Yu-Lei, WANG Sheng-Xing, ZHAO Liang-Xia, ZHANG De-Xin, HU Jian-Bang, CAO Xue-Lian, YANG Ya-Jie, CHANG Cheng, MA Chuan-Xi, ZHANG Hai-Ping^*

Key Laboratory of Wheat Biology and Genetic Improvement in South Yellow & Huai River Valley, Ministry of Agriculture / College of Agronomy, Anhui Agricultural University, Hefei 230036, China

Fund:

Abstract

To improve pre-harvest sprouting (PHS) resistance in wheat breeding, it is important to explore marker loci and elite alleles associated with PHS resistance using intact spikes. In this study, a total of 181 markers were used to 264 genotype materials. General and mixed linear models (GLM and MLM) were used to analyze PHS phenotypic data in three environments (2012-in house, 2013-in house and 2013-in field). The results showed that twenty markers were identified by association analysis, and located on chromosomes 1AS, 2DS, 3AS, 3BL, 4AL, 5AS, 5BL, 6BS, 6DS, 7AL, and 7BL. The markersgwm102 on 2DS andbarc340 on 7BL were detected stably in three environments, among whichbarc340 was likely to be novel and needs to be further studied through biparental linkage mapping analysis. Six markers were detected in two environments, and the other loci linked with 12 markers were detected only in one environment. A total of ten elite alleles were further explored among the eight loci with repeated associations. The allelesbarc28-229bp andbarc28-217bpfor high PHS resistance were all distributed in local cultivars (e.g. Suiningtuotuomai). However, the allelesgwm102-142bp andbarc186-199bp with intermediate PHS resistance were mainly detected in released cultivars (e.g. Yangmai 158), which could be beneficial to wheat molecular breeding.

Keyword: Triticum aestivumL.; Preharvest sprouting; Dormancy; Molecular marker; Association analysis

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小麦收获前遇连阴雨天气或处于湿热环境时间较长, 易发生穗发芽(pre-harvest sprouting, PHS)。穗发芽导致种子内部储藏物质消耗, 千粒重下降和减产, 同时营养和加工品质劣化^{[ 1]}。采用准确、可靠的评价方法筛选抗穗发芽资源, 从中挖掘抗性基因, 开发功能标记, 最终培育抗穗发芽品种是解决这一问题的根本途径。

小麦收获前穗发芽抗性受多种因素共同影响, 如种皮颜色、种子的休眠特性、小麦的穗部形态(有芒或无芒等)、颖壳内发芽抑制物质、籽粒吸水特性、α-淀粉酶活性、脱落酸(ABA)和赤霉素(GA)等因素, 以及外界环境因素如光照、温度和湿度等, 其中种子的休眠性被认为是穗发芽抗性的主要遗传因素^{[ 2, 3, 4, 5]}。目前, 对小麦穗发芽抗性的评价主要有籽粒发芽法、整穗发芽法和α-淀粉酶活性测定法等。籽粒发芽法仅能反映小麦种子本身的休眠状况, 却忽略了颖壳等外部抑制物质的作用。α-淀粉酶活性测定法操作相对较复杂, 不适于批量测定。相对于前两者, 整穗发芽法操作简单, 适合较多试验材料的批量测定, 且更能全面衡量小麦穗子的整体抗性水平, 主要有发芽纸保湿法、塑料袋保湿法、纱布保湿法、模拟降雨法等^{[ 6]}。

通过解析小麦穗发芽抗性分子机制, 发现小麦穗发芽抗性受多基因控制, 存在主效基因。利用DH或RIL等人工作图群体, 控制穗发芽抗性的主效基因被定位在2AL ( Xwmc1045和 Xgwm296), 3AS ( Xfbb370、 Xbarc310、 Xbcd907、 Xbarc57、 Xbarc321、 Xbarc12和 Xbarc310), 3AL ( Xgwm155和 Xwmc153), 4AL ( Xbarc170、 Xgwm397、 Xgwm269、 Xgwm937、 Xgwm894、 Xgwm637、 Xzxq118和 Xhbe03), 5D ( Xcfd40和 Xbarc1097), 6BS ( Xwmc104)及7DL ( XMST101)上^{[ 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21]}。而基于自然群体的关联分析是定位和挖掘数量性状位点的另一条有效途径。利用这一方法, Arif等^{[ 22]}检测出34个与小麦整穗发芽性状相关的位点, 分别位于1A、2A、2D、4A、5B、5D、6A、6B、7A和7D上, 其中2D、5B和7A是携带穗发芽抗性位点的密集区域; Kulwal等^{[ 23]}除发现位于1BS、2BS、2BL、2DS、4AL、6DL和7DS上已报道的QTL外, 还鉴定出位于7BS的新位点。

本试验采用发芽纸保湿法对264份小麦材料进行整穗发芽抗性测定, 采用6%变性聚丙烯酰胺凝胶电泳对其进行基因型分析, 利用TASSLE软件的一般线性模型(GLM)和混合线性模型(MLM)对基因型和表型值进行关联分析, 检测与整穗发芽抗性紧密关联的分子标记位点, 并进一步挖掘优异等位变异及其相应载体材料, 为小麦抗穗发芽分子育种提供参考。

1 材料与方法

1.1 试验材料

本课题组前期收集并保存410份育种亲本材料, 利用均匀分布于小麦21条染色体上的80对多态性丰富的SSR引物对其进行遗传多样性分析, 从中筛选出亲缘关系相对较远, 且穗发芽抗性不同的264份材料作为本试验材料, 包括推广品种204份、国外引进品种4份、地方品种18份、中间品系和高代材料38份。2011年和2012年, 所有材料均秋播于安徽农业大学合肥大杨店试验站, 每份材料种2行, 行长2 m, 行宽20 cm, 每行40粒, 常规田间管理。

1.2 穗发芽抗性评价

采用整穗发芽率(germination percentage, GP)来评价穗发芽抗性水平, 分为抗穗发芽、中间型和感穗发芽3个抗性等级, 其整穗发芽率分别为0~30%、31%~60%和61%~100%^{[ 6]}。

2012年和2013年分别进行室内鉴定。于小麦蜡熟后期(茎秆、叶片和麦穗全部变黄)收获每份材料10个主茎穗, 5穗为一个重复, 室内风干1 d, 立即存放于-20℃冰箱以维持种子的休眠性, 待收获全部材料后一并进行整穗发芽试验。先把整穗在灭菌水中充分浸泡10 min, 用发芽纸卷裹, 保鲜袋保湿, 置人工气候培养箱中(温度20℃, 光周期14 h昼/10 h夜, 相对湿度80%)培养7 d。于第7天取出并除去保鲜袋, 但保留发芽纸, 于电热恒温干燥箱(150℃)快速烘干, 阻止其继续发芽; 再于阳光下暴晒至籽粒易剥取; 记录每个穗子穗发芽率后取两重复平均值, 以胚部出现中破裂迹象为鉴定标准^{[ 1]}。

2013年小麦收获期遇到连阴雨天气, 于田间进行了重复鉴定。每份材料留10个生理成熟一致主茎穗直接在田间自然发芽, 1周后收回室内, 置于电热恒温干燥箱中(150℃)快速烘干, 手工脱粒, 统计每份材料发芽种子数和总粒数, 计算平均整穗发芽率。

采用SPSS19.0软件分别对3个环境下小麦整穗发芽率进行统计描述和方差分析。

1.3 分子标记筛选及其扩增条件

选取15份亲缘关系较远的材料进行SSR引物筛选, 筛选标准为带型清晰、多态性丰富。引物信息来自Graingenes (http://wheat.pw.usda.gov/GG2/ index.shtm/), 由生工生物工程(上海)有限公司合成引物。最终筛选出180对SSR引物, 另外补充小麦穗发芽基因功能标记 Vp1B3^{[ 6]}, 总共181个分子标记用于关联分析。

采用SDS饱和酚法提取小麦基因组DNA。在NanoVue Plus微量分光光度计上检测DNA 浓度, 于-20℃冰箱保存。PCR扩增体系为10 μL, 包括10× buffer (含有2.0 mmol L^-1 Mg²⁺) 1.0 μL, 2.5 mmol L^-1 dNTPs 0.8 μL, 5 U μL^-1 Taq DNA polymerase 0.11 μL, 10 μmol L^-1引物各0.4 μL, 2.0 μL模板DNA (50~ 60 ng μL^-1), ddH₂O 5.29 μL。反应程序为94℃预变性5 min; 35个循环(95℃变性30 s, 48~62℃退火30 s, 72℃延伸30 s); 72℃延伸10 min; 4℃保存。用6%变性聚丙烯酰胺凝胶电泳检测扩增产物。利用软件Gelpro32 (http://www.mediacy.com/)分析带型。

1.4 关联分析

参考Mackay等^{[ 24]}的方法, 从180对SSR引物中选取随机分布于小麦全基因组的105对独立SSR引物对群体遗传结构进行评估。采用STRUCTURE软件^{[ 25]}的混合模型来确定群体结构, K值取1~11, 每个 K值单独运行6次。当 K=4时曲线出现最大拐点, 因此将264份材料分为4个类群。

利用TASSLE软件^{[ 26]}的一般线性模型(general linear model, GLM)和混合线性模型(mixed linear model, MLM), 将群体结构分析所得的 Q值和亲缘关系 K作为协变量, 进行标记与性状的关联分析, 选取两种模型共同关联到的显著( P<0.05)标记位点, 并计算这些标记位点对表型变异的解释率。

参考文自翔等^{[ 27]}描述的方法, 对整穗发芽抗性显著关联的标记位点进一步挖掘其优异等位变异, 明确各等位变异的表型效应及其载体材料。

a_i =

式中, a_i代表第 i个等位变异的表型效应值, x_ij为携带第 i个等位变异在第 j份材料中表型值, n_i为具有第 i等位变异的材料数, N_k为携带无效等位变异(某一位点中, 若其中一个等位变异被视为“有效”, 则其他等位变异相应地为“无效”)的第 k份料的表型值, n_k为具有无效等位变异的材料数。若 a_i > 0, 则认为该等位变异为增效等位变异, 反之为减效等位变异。

2 结果与分析

2.1 不同环境下整穗发芽抗性表型分析

264份材料在3个环境中均表现出丰富的表型变异, 整穗发芽率变幅分别为0~92%、0~100%和0~96%, 其中2012年室内和2013年田间自然环境下的变异系数均在0.80以上。2013年室内总体平均值最高, 为69%, 2012年室内和2013年田间自然环境下总体平均值较低, 分别为25%和16% (表1)。

表1 264份材料在3个环境下的整穗发芽率变异 Table 1 Variation of germination percentage of intact spike in 264 wheat genotypes detected in three environments

不同穗发芽抗性等级材料的分布在不同环境中表现不一, 2012室内和2013田间环境下分布趋势相似, 均有60%以上的材料分布在0~30%抗性等级中, 分布到61%~100%等级中的材料不满10%。2013室内频率分布不同于前两者, 其分布在0~30%和31%~60%等级中的材料均不满20%, 超过60%的材料分布在61%~100%等级中(图1)。究其原因可能是2013年小麦种子成熟期间的温度以及收获前降雨的影响, 造成整体供试材料穗发芽抗性水平下降。

方差分析(表2)显示, 3种环境下测得供试材料间整穗发芽率差异极显著( P=0.0001), 说明小麦穗发芽抗性既受基因型的控制, 也易受环境影响。供试材料的整穗发芽率在3种环境间的相关性均达极大, 为0.469; 2013室内和2013田间相关系数为0.334。综合分析上述结果说明, 整穗发芽抗性易受环境影响, 但在不同年份间, 以及同一年份内不同环境下, 不同供试材料整穗发芽率表现出相对稳定的趋势。

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图1 3个环境下不同级别穗发芽抗性材料的分布Resistant、Intermediate和Susceptible表示抗穗发芽、中间型和感穗发芽, 发芽率分别为0~30%、31%~60%和61%~100%。Fig. 1 Frequencies of materials with different resistances to preharvest spouting evaluated in three environmentsThe germination percentage of resistant, intermediate and susceptible cultivars separately was 0-30%, 31%-60%, and 61%-100%.

表2 供试264份材料整穗发芽抗性的方差分析 Table 2 ANOVA of sprouting resistance in 264 genotypes used in this study

显著水平( P<0.01)。其中, 2012室内和 2013室内相关系数最小, 为0.286; 2012室内与2013田间相关系数最

2.2 整穗发芽抗性基因定位与分子标记发掘

两种模型共同检测到20个标记位点与整穗发芽抗性显著相关( P<0.05)。表3为MLM模型的检测结果。其中与2012室内关联的分子标记位点有10个, 分别位于2DS ( gwm102和 barc1146)、3AS ( barc321和 gwm369)、4AL ( wmc617)、5AS ( barc186)、5BL ( barc156)、6BS ( barc14)和7BL ( barc1181和 barc340), 各标记位点对表型贡献为2.2%~10.2%, 其中6BS上的 barc14对表型贡献最大(10.2%)。与2013室内关联的分子标记位点有11个, 分别位于染色体1AS ( barc28)、2DS ( gwm102)、3BL ( barc344和 Vp1B3)、4AL ( gwm610和 barc78)、5BL ( barc109和 barc156)、6DS ( barc173)、7AL ( barc281)和7BL ( barc340)上, 各标记位点对表型贡献为3.1%~8.0%, 其中4AL上的 barc78对表型贡献最大(8.0%)。与2013田间关联的分子标记位点有9个, 分别位于染色体1AS ( barc28)、2DS ( gwm102和 barc1146)、3AS ( barc321和 barc57)、5AS ( barc186)、5DL ( barc347)和7BL ( barc1181和 barc340)上, 各标记位点对表型贡献为3.5%~15.0%, 其中 barc321和 barc57对表型贡献相对较大(10.6%和15.0%), 均位于3AS染色体区域, 且距离较近。

表3 混合线性模型检测到的与穗发芽抗性相关的标记位点( P<0.05)及其对表型贡献 Table 3 Marker loci associated with preharvest sprouting resistance ( P<0.05) detected by mixed linear model and their phenotypic contributions

3种环境下均检测到与整穗发芽抗性相关的分子标记位点有2个, 分别为 gwm102和 barc340; 2种环境下均检测到的分子标记位点有6个, 分别为 barc28、 barc1146、 barc321、 barc186、 barc156和 barc1181; 其余12个分子标记位点仅在一种环境下被检测到。说明小麦整穗发芽抗性虽然存在效应稳定的QTL, 但受基因型与环境互作的显著影响, 与方差分析结果吻合。

2.3 优异等位变异及载体材料的鉴定

对8个重复检测到的标记作等位变异分析, 共鉴定出10个降低整穗发芽率的优异等位变异(表4), 其载体主要为6份材料, 即茶淀红麦、遂宁坨坨麦、扬麦158、梓潼女儿麦、川麦42和扬麦20。从各等位变异的分布频率来看, barc186-199bp在供试材料中所占比例最大(62.1%, 164/264), 其次是 barc1181- 256bp (38.6%, 102/264), barc28-229bp最小(0.7%, 2/264)。从各等位变异的表型效应来看, barc28- 229bp和 barc28-217bp在3种环境下的穗发芽程度显著低于其他类型, 其来源主要是地方品种, 如遂宁坨坨麦、涪陵须须麦和梓潼女儿麦等, 在小麦育种中难以直接利用; 而广泛分布在推广品种中的 barc186-199bp和 gwm102-142bp虽然效应值相对较小, 但易于直接利用, 应该引起重视。

不同载体材料携带的等位变异及其组合不同, 如遂宁坨坨麦, 其同时携带 barc28-217bp、 barc321- 182/172bp和 barc1181-256bp三种等位变异组合; 扬麦20同时携带 barc186-199bp和 barc340-207bp两种组合; 梓潼女儿麦同时携带 barc1146-197bp和 barc340-194bp两种组合(表4)。这可能对表型起到一种聚合的效应, 从而导致其整穗发芽抗性水平较高。

进一步分析上述10种优异等位变异在不同抗性等级材料中的分布, 发现携带优异等位变异的材料大都表现抗穗发芽特性, 说明它们对整穗发芽抗性有不同程度的贡献(表4)。如携带 barc28-229bp和 barc28-217bp的材料全部表现抗穗发芽; 携带 gwm102-142bp、 barc1146-197bp和 barc321-182/ 172bp的材料中表现抗穗发芽的比率分别为65.2% (45/69)、73.3% (11/15)和66.7% (16/24)。

3 讨论

业已报道, 与小麦穗发芽相关的QTL涉及多条染色体和众多区段^{[ 12, 13, 15, 16, 19, 22, 23]}。本研究采用GLM和MLM模型对264份材料不同环境下的穗发芽表型性状及其基因型进行关联分析, 20个标记位点被共同鉴定出与小麦整穗发芽抗性关联, 分别位于1AS、2DS、3AS、3BL、4AL、5AS、5BL、5DL、6BS、6DS、7AL和7BL上, 这一结果支持上述结论。但本研究鉴定的部分标记位点与相同染色体上已报道的位点距离较远, 如 barc28(1AS)与Mohan等^{[ 5]}定位的QTL(连锁标记为 Xbarc24和 Xbarc119), barc340(7BL)与Liu等^{[ 17]}鉴定的2个QTL(标记区间分别为 Xbarc255- Xgwm43和 Xwmc517- Xbarc278)都较远。本研究是否发现了新位点, 有待进一步验证。今后将着手开发多种标记(SNP、EST-SSR、EST、SCAR和CAPS等)用于加密标记图谱, 同时构建适当的作图群体, 以准确定位这些关联位点。

另有一些已报道的位点在本研究中被重复鉴定, 但其表型的贡献率在不同试验中不尽相同。如3AS上的 barc321和 barc57, 在本研究中其对表型解释效应为5.7%~15.0%, 而在张海萍等^{[ 14]}的试验中为25.6%~48.3%。本研究检测到的 wmc617 (4AL)、 gwm610 (4AL)和 barc347(5DL)与Chen等^{[ 16]}、Torada等^{[ 21]}和Munkvold等^{[ 28]}定位的QTL具有相同标记, 但表型效应较小。说明小麦抗穗发芽性状受群体、遗传背景、环境条件等影响很大, 此外, 基因型与环境互作也很明显, 本研究多数位点(12个)只能在单一环境下被检测到, 方差分析也证实这一点。

本研究检测到若干整穗发芽抗性关联位点在不同环境中较为稳定, 如 gwm102和 barc340在3种环境下均显著关联目标性状, 且与之较近的标记 barc1146和 barc1181也能在2种环境下被检测到, 与Kulwal等^{[ 8, 23]}和Liu等^{[ 13]}的研究结果相似。说明生产上有可能通过分子标记辅助选择技术改良小麦穗发芽抗性。在8个重复检测位点上, barc28-229bp、 barc28-217bp、 gwm102-142bp和 barc340-207bp具有显著提高整穗发芽抗性的作用; 前二者可能是提高整穗发芽抗性的主要优异等位变异, 但其主要由地方品种携带, 无法直接用于小麦育种, 但应重视利用其创制中间材料。另外, 推广品种中存在一些优异等位变异, 便于直接利用, 如本研究发现扬麦158、扬麦20、川麦42均携带控制低穗发芽率(0~30%)等位变异。我们认为, 育种中可以尝试利用分布在育成品种中、效应适中且稳定的等位变异或变异组合(如 gwm102-142bp、 barc156-204bp、 barc340-207bp和 barc340-194bp)来提高穗发芽抗性水平; 在多种环境下表现稳定的QTL, 虽然其效应可能不是最大, 但比在特定环境下表达的QTL更有育种价值^{[ 29]}。

表4 与整穗发芽抗性显著关联的分子标记位点优异等位变异的表型效应和相应载体材料 Table 4 Phenotypic effects and carrier materials of elite alleles at loci strongly associated with PHS resistance

4 结论

鉴定出与小麦整穗发芽抗性显著关联的分子标记, 证实2DS、3AS和7BL染色体区域含有稳定的抗穗发芽位点, 1AS和7BL等染色体上可能存在新位点。小麦整穗发芽抗性受基因型和环境共同影响。

The authors have declared that no competing interests exist.

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

参考文献

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2002

3.658

0.0

Groos

, Gay

, Perretant

R M

, Bernard

G M

, Charmet

D G

. Study of the relationship between pre-harvest sprouting and grain color by quantitative trait loci analysis in a white red grain bread-wheat cross. Theor Appl Genet, 2002, 104: 39-47

In many wheat ( Triticum aestivum L.) growing areas, pre-harvest sprouting (PHS) may cause important damage, and in particular, it has deleterious effects on bread-making quality. The relationship between PHS and grain color is well known and could be due either to the pleiotropic effect of genes controlling red-testa pigmentation ( R ) or to linkage between these genes and other genes affecting PHS. In the present work, we have studied a population of 194 recombinant inbred lines from the cross between two cultivars, ’Renan’ and ’Récital’, in order to detect QTLs for both PHS resistance and grain color. The variety ’Renan’ has red kernels and is resistant to PHS, while ’Récital’ has white grain and is highly susceptible to PHS. A molecular-marker linkage map of this cross was constructed using SSRs, RFLPs and AFLPs. The population was evaluated over 2 years at Clermont-Ferrand (France). PHS was evaluated on mature spikes under controlled conditions and red-grain color was measured using a chromameter. Over the 2 years, we detected four QTLs for PHS, all of them being co-localized with QTLs for grain color. Three of them were located on the long arm of chromosomes 3 A, 3B and 3D, close to the loci where the genes R and taVp1 were previously mapped. For these three QTLs, the resistance to PHS is due to the allele of the variety ’Renan’. Another co-located QTL for PHS and grain color was detected on the short arm of chromosome 5 A. The resistance for PHS for this QTL is due to the allele of ’Récital’.

1.Institut National de la Recherche Agronomique (INRA), Domaine de Crouelle, 234 rue du Brézet, 63039 Clermont-Ferrand, France e-mail: charmet@clermont.inra.fr FR FR 2.Institut National de la Recherche Agronomique (INRA), Domaine du Rheu, BP 35327, 35650 Le Rheu Cedex, France FR FR

... 穗发芽导致种子内部储藏物质消耗, 千粒重下降和减产, 同时营养和加工品质劣化^[1] ...

... 记录每个穗子穗发芽率后取两重复平均值, 以胚部出现中破裂迹象为鉴定标准^[1] ...

2006

1.328

0.0

... -淀粉酶活性、脱落酸(ABA)和赤霉素(GA)等因素, 以及外界环境因素如光照、温度和湿度等, 其中种子的休眠性被认为是穗发芽抗性的主要遗传因素^[2,3,4,5] ...

2001

1.328

0.0

2006

1.643

0.0

Kottearachchi

N S

, Uchino

, Kato

, Miura

. Increased grain dormancy in white-grained wheat by introgression of preharvest sprouting tolerance QTLs. Euphytica, 2006, 152: 421-428

White-grained wheat cultivars have long been recognized to be less resistant to preharvest sprouting (PHS) than the red-grained ones. Previously two QTLs for grain dormancy, QPhs.ocs-3A.1 ( QPhs-3AS ) and QPhs.ocs-4A.1 ( QPhs-4AL ) were identified in a highly dormant Japanese red wheat, Zenkoujikomugi (Zen). Aiming at improvement of PHS tolerance in white-grained wheat, the introgression effect of these two QTLs in a white-grained population consisting of 40 recombinant inbred lines (RILs) developed from a cross between Zen and white-grained Spica was examined here. Random 20 RILs with red grains were also developed from the same cross and used as a control population. The RILs were grown in the field and in the glasshouse to evaluate the grain dormancy by germination test. Several SSR markers closely linked to the QPhs-3AS and QPhs-4AL were used to estimate the alleles at the QTLs. Dormancy variation in the RILs was significantly associated with the differences for grain color and the alleles at QPhs-3AS over several years. Although allelic variation was detected in a SSR marker closely linked to QPhs-4AL there was no difference in germination data between the Zen-allele and the Spica-allele groups. As expected, the red-grained RILs with the Zen allele at QPhs-3AS were the most dormant. Some white-grained RILs with the Zen allele at QPhs-3AS showed higher dormancy compared to the red-grained RILs with the alternative allele. These results demonstrated that introgression of the QPhs-3AS gene could contribute to the increased grain dormancy in white-grained wheat.

1.Department of Crop Science, Obihiro University of Agriculture & Veterinary Medicine, Obihiro, 080-8555 Japan

2002

2.471

0.0

... 但本研究鉴定的部分标记位点与相同染色体上已报道的位点距离较远, 如barc28 (1AS)与Mohan等^[5]定位的QTL(连锁标记为Xbarc24和Xbarc119), barc340 (7BL)与Liu等^[17]鉴定的2个QTL(标记区间分别为Xbarc255-Xgwm43和Xwmc517-Xbarc278)都较远 ...

2007

3.658

0.0

Yang

, Zhao

X L

, Xia

L Q

, Chen

X M

, Xia

X C

, Yu

, He

Z H

, Roder

. Development and validation of a Viviparous-1 STS marker for preharvest sprouting tolerance in Chinese wheats. Theor Appl Genet, 2007, 115: 971-980

Pre-harvest sprouting (PHS) of wheat reduces the quality of wheat grain, and improving PHS tolerance is a priority in certain wheat growing regions where conditions favorable for PHS exist. Two new Viviparous-1 allelic variants related to PHS tolerance were investigated on B genome of bread wheat, and designated as Vp-1Bb and Vp-1Bc , respectively. Sequence analysis showed that Vp-1Bb and Vp-1Bc had an insertion of 193-bp and a deletion of 83-bp fragment, respectively, located in the third intron region of the Vp-1B gene. The insertion and deletion affected the expression level of the Vp1 at mature seed stage, more correctly spliced transcripts were observed from the genotypes with either insertion or deletion than that of the wild type. Based on these insertions and deletions, a co-dominant STS marker of Vp-1B gene was developed and designated as Vp1B3 , which in most cases could amplify either 845 or 569-bp fragment from the tolerant cultivars, and 652-bp from the susceptible ones. This Vp1B3 marker was mapped to chromosome 3BL using a set of Chinese Spring nulli-tetrasomic and ditelosomic lines. A total of 89 white-grained Chinese wheat cultivars and advanced lines, were used to validate the relationship between the polymorphic fragments of Vp1B3 and PHS tolerance. Statistical analysis indicated that Vp1B3 was strongly associated with PHS tolerance in this set of Chinese germplasm, suggesting that Vp1B3 could be used as an efficient and reliable co-dominant marker in the evaluation of wheat germplasm for PHS tolerance and marker-assisted breeding for PHS tolerant cultivars.

1.Chinese Academy of Agricultural Sciences (CAAS) Institute of Crop Science, National Wheat Improvement Center, The National Key Facility for Crop Gene Resources and Genetic Improvement No 12 Zhongguancun South Street Beijing 100081 China 2.Inner Mongolia Agricultural University College of Agronomy No 306 Zhaowuda Road Hohhot 010018 Inner Mongolia China 3.Henan Academy of Agricultural Sciences Wheat Research Institute No 1 Nongye Road Zhengzhou 450002 Henan China 4.CIMMYT China Office, C/O CAAS Beijing 100081 China 5.Leibiniz Institute of Plant Genetics and Crop Plant Research (IPK) Corrensstrasse 3 06466 Gatersleben Germany

... 相对于前两者, 整穗发芽法操作简单, 适合较多试验材料的批量测定, 且更能全面衡量小麦穗子的整体抗性水平, 主要有发芽纸保湿法、塑料袋保湿法、纱布保湿法、模拟降雨法等^[6] ...

... 2 穗发芽抗性评价采用整穗发芽率(germination percentage, GP)来评价穗发芽抗性水平, 分为抗穗发芽、中间型和感穗发芽3个抗性等级, 其整穗发芽率分别为0~30%、31%~60%和61%~100% ^[6] ...

... 最终筛选出180对SSR引物, 另外补充小麦穗发芽基因功能标记Vp1B3^[6], 总共181个分子标记用于关联分析 ...

1999

3.658

0.0

... 利用DH或RIL等人工作图群体, 控制穗发芽抗性的主效基因被定位在2AL (Xwmc1045和Xgwm296), 3AS (Xfbb370、Xbarc310、Xbcd907、Xbarc57、Xbarc321、Xbarc12和Xbarc310), 3AL (Xgwm155和Xwmc153), 4AL (Xbarc170、Xgwm397、Xgwm269、Xgwm937、Xgwm894、Xgwm637、Xzxq118和Xhbe03), 5D (Xcfd40和Xbarc1097), 6BS (Xwmc104)及7DL (XMST101)上^{[7,8,9,10,11,12,13,14,15,16,17,18,19,20,21]} ...

2005

3.658

0.0

Kulwal

P L

, Kumar

, Gaur

, Khurana

J P

, Tyagi

A K

, Balyan

H S

, Gupta

P K

. Mapping of a major QTL for preharvest sprouting tolerance on chromosome 3A in bread wheat. Theor Appl Genet, 2005, 111: 1052-1059

Quantitative trait loci (QTL) analysis was conducted for pre-harvest sprouting tolerance (PHST) in bread wheat for a solitary chromosome 3A, which was shown to be important for this trait in earlier studies. An intervarietal mapping population in the form of recombinant inbred lines (RILs) developed from a cross between SPR8198 (a PHS tolerant genotype) and HD2329 (a PHS susceptible cultivar) was used for this purpose. The parents and the RIL population were grown in six different environments and the data on PHS were collected in each case. A framework linkage map of chromosome 3A with 13 markers was prepared and used for QTL analysis. A major QTL ( QPhs.ccsu-3A.1 ) was detected on 3AL at a genetic distance of ∼183 cM from centromere, the length of the map being 279.1 cM. The QTL explained 24.68% to 35.21% variation in individual environments and 78.03% of the variation across the environments (pooled data). The results of the present study are significant on two counts. Firstly, the detected QTL is a major QTL, explaining up to 78.03% of the variation and, secondly, the QTL showed up in all the six environments and also with the pooled data, which is rather rare in QTL analysis. The positive additive effects in the present study suggest that a superior allele of the QTL is available in the superior parent (SPR8198), which can be used for marker-aided selection for the transfer of this QTL allele to obtain PHS-tolerant progeny. It has also been shown that the red-coloured grain of PHS tolerant parent is not associated with the QTL for PHST identified during the present study, suggesting that PHS tolerant white-grained cultivars can be developed.

1.Molecular Biology Laboratory, Department of Genetics & Plant Breeding, Ch. Charan Singh University, Meerut, UP, 250004 India 2.Department of Plant Molecular Biology, University of Delhi South Campus, Benito Juarez Road, New Delhi, 110021 India

... 本研究检测到若干整穗发芽抗性关联位点在不同环境中较为稳定, 如gwm102和barc340在3种环境下均显著关联目标性状, 且与之较近的标记barc1146和barc1181也能在2种环境下被检测到, 与Kulwal等^[8,23]和Liu等^[13]的研究结果相似 ...

2009

1.643

0.0

Mohan

, Kulwal

, Singh

, Kumar

, Mir

R R

, Kumar

, Prasad

, Balyan

H S

, Gupta

P K

. Genome-wide QTL analysis for pre-harvest sprouting tolerance in bread wheat. Euphytica, 2009, 168: 319-329

A framework linkage map comprising 214 molecular marker (SSR, AFLP, SAMPL) loci was prepared using an intervarietal recombinant inbred line (RIL) mapping population of bread wheat. The RIL population that was developed from the cross SPR8198 (red-grained and PHS tolerant genotype) × HD2329 (white-grained and PHS susceptible genotype) following single seed descent segregated for pre-harvest sprouting (PHS). The RIL population and parental genotypes were evaluated in six different environments and the data on PHS were collected. Using the linkage map and PHS data, genome-wide single-locus and two-locus QTL analyses were conducted for PHS tolerance (PHST). Single-locus analysis following composite interval mapping (CIM) detected a total of seven QTL, located on specific arms of five different chromosome (1AS, 2AL, 2DL, 3AL and 3BL). These seven QTL included two major QTL one each on 2AL and 3AL. Two of these seven QTL were also detected following two-locus analysis, which resolved a total of four main-effect QTL (M-QTL), and 12 epistatic QTL (E-QTL), the latter involved in 7 QTL × QTL interactions. Interestingly, none of these M-QTL and E-QTL detected by two-locus analysis was involved in Q × E and Q × Q × E interactions, supporting the results of ANOVA, where genotype × environment interaction were non-significant. The QTL for PHS detected in the present study may be efficiently utilized for marker-aided selection for enhancing PHST in bread wheat.

1.Molecular Biology Lab, Department of Genetics & Plant Breeding, C.C.S. University, Meerut, 250004 India 2.Biotechnology Centre, Department of Agriculture Botany, Dr. Panjabrao Deshmukh Agricultural University, Akola, 444104 India 3.National Institute for Plant Genome Research, JNU Campus, Aruna Asaf Ali Marg, P.O. Box 10531, New Delhi, 110067 India

2003

3.658

0.0

Osa

, Kato

, Mori

, Shindo

, Torada

, Miura

. Mapping QTLs for seed dormancy and the Vp1 homologue on chromosome 3A in wheat. Theor Appl Genet, 2003, 106: 1491-1496

1., Department of Crop Science, Obihiro University of Agriculture and Veterinary Medicine, Obihiro, 080-8555, Japan 2., Kihara Institute for Biological Research, Yokohama City University, Maioka, Totsuka-ku, Yokohama, 244-0813, Japan 3., Hokkaido Green-Bio Institute, Naganuma, Hokkaido, 069-1301, Japan

2005

3.658

0.0

Mori

, Uchino

, Chono

, Kato

, Miura

. Mapping QTLs for grain dormancy on wheat chromosome 3A and group 4 chromosomes, and their combined effect. Theor Appl Genet, 2005, 110: 1315-1323

A major QTL for grain dormancy, QPhs.ocs-3A.1 , derived from the highly dormant wheat Zenkoujikomugi (Zen), has been identified in a study made under a controlled environment. Further investigations were needed to dissect the precise position and expression of QPhs.ocs-3A.1 under different field conditions because the ability to detect genetic loci for grain dormancy traits is compromised by environmental effects and genotype/environment interactions. Group 4 chromosomes have also been shown to be possible sites of QTLs for grain dormancy. The objectives of this study were (1) to locate additional molecular markers in the QPhs.ocs-3A.1 region, (2) to identify QTLs on the group 4 chromosomes and (3) to elucidate their combined effects. We examined the recombinant inbred lines (RILs) from a cross between Chinese Spring (CS) and Zen over a 3-year period in one location and 1 year in a different location. In an interval mapping study QPhs.ocs-3A.1 was mapped to within the 4.6 cM region flanked by Xbarc310 and Xbcd907 at the proximal end of the short arm of chromosome 3A. QPhs.ocs-3A.1 was confirmed to be the predominant dormancy QTL since it explained a large portion (11.6–44.8%) of the phenotypic variation, and was strongly displayed under dormancy-breaking conditions or at low germination temperatures. For QPhs.ocs-4A.1 , identified on the long arm of chromosome 4A, and QPhs.ocs-4B.1 , on the centromeric region of the long arm of Chr 4B, the LOD peak positions and the desirable allele were consistent between the trials, while the LOD scores and contribution to the phenotypic variation varied. Transgressive segregants were observed among the 125 RILs and most of them had a combination of the three alleles conferring a higher dormancy: the Zen alleles at QPhs.ocs-3A.1 and QPhs.ocs-4A.1 and the CS allele at QPhs.ocs-4B1 . This demonstrated a combined effect of the desirable alleles on accelerating grain dormancy, with their total effect being superior to that of Zen.

1.Department of Crop Science Obihiro University of Agriculture and Veterinary Medicine Obihiro 080-8555 Hokkaido Japan 2.Department of Wheat and Barley Research National Institute of Crop Science, National Agriculture and Bio-oriented Research Organization 2-1-18 Kannondai Tsukuba 305-8505 Ibaraki Japan

2008

3.658

0.0

Liu

S B

, Cai

S B

, Robert

, Chen

C X

, Bai

G H

. Quantitative trait loci for resistance to pre-harvest sprouting in US hard white winter wheat Rio Blanco. Theor Appl Genet, 2008, 117: 691-699

Pre-harvest sprouting (PHS) of wheat is a major problem that severely limits the end-use quality of flour in many wheat-growing areas worldwide. To identify quantitative trait loci (QTLs) for PHS resistance, a population of 171 recombinant inbred lines (RILs) was developed from the cross between PHS-resistant white wheat cultivar Rio Blanco and PHS-susceptible white wheat breeding line NW97S186. The population was evaluated for PHS in three greenhouse experiments and one field experiment. After 1,430 pairs of simple sequence repeat (SSR) primers were screened between the two parents and two bulks, 112 polymorphic markers between two bulks were used to screen the RILs. One major QTL, QPhs.pseru-3AS , was identified in the distal region of chromosome 3AS and explained up to 41.0% of the total phenotypic variation in three greenhouse experiments. One minor QTL, QPhs.pseru-2B.1, was detected in the 2005 and 2006 experiments and for the means over the greenhouse experiments, and explained 5.0–6.4% of phenotypic variation. Another minor QTL, QPhs.pseru-2B . 2 , was detected in only one greenhouse experiment and explained 4.5% of phenotypic variation for PHS resistance. In another RIL population developed from the cross of Rio Blanco/NW97S078, QPhs.pseru-3AS was significant for all three greenhouse experiments and the means over all greenhouse experiments and explained up to 58.0% of phenotypic variation. Because Rio Blanco is a popular parent used in many hard winter wheat breeding programs, SSR markers linked to the QTLs have potential for use in high-throughput marker-assisted selection of wheat cultivars with improved PHS resistance as well as fine mapping and map-based cloning of the major QTL QPhs.pseru-3AS .

1.Kansas State University Department of Agronomy Manhattan KS 66506 USA 2.Jiangsu Academy of Agricultural Science Nanjing 210014 China 3.University of Nebraska USDA-ARS, 314 Biochem Hall Lincoln NE 68583 USA 4.USDA-ARS, Plant Science and Entomology Research Unit 4008 Throckmorton Hall Manhattan KS 66506 USA

... 3 讨论业已报道, 与小麦穗发芽相关的QTL涉及多条染色体和众多区段^{[12,13,15,16,19,22,23]} ...

2010

3.658

0.0

Liu

S B

, Bai

G H

. Dissection and fine mapping of a major QTL for preharvest sprouting resistance in white wheat Rio Blanco. Theor Appl Genet, 2010, 121: 1395-1404

Preharvest sprouting (PHS) is a major constraint to white wheat production. Previously, we mapped quantitative trait loci (QTL) for PHS resistance in white wheat by using a recombinant inbred line (RIL) population derived from the cross Rio Blanco/NW97S186. One QTL, QPhs.pseru - 3A , showed a major effect on PHS resistance, and three simple sequence repeat (SSR) markers were mapped in the QTL region. To determine the flanking markers for the QTL and narrow down the QTL to a smaller chromosome region, we developed a new fine mapping population of 1,874 secondary segregating F 2 plants by selfing an F6 RIL (RIL25) that was heterozygous in the three SSR marker loci. Segregation of PHS resistance in the population fitted monogenic inheritance. An additive effect of the QTL played a major role on PHS resistance, but a dominant effect was also observed. Fifty-six recombinants among the three SSR markers were identified in the population and selfed to produce homozygous recombinants or QTL near-isogenic lines (NIL). PHS evaluation of the recombinants delineated the QTL in the region close to Xbarc57 flanked by Xbarc321 and Xbarc12 . To saturate the QTL region, 11 amplified fragment length polymorphism (AFLP) markers were mapped in the QTL region with 7 AFLP co-segregated with Xbarc57 by using the NIL population. Dissection of the QTL as a Mendelian factor and saturation of the QTL region with additional markers created a solid foundation for positional cloning of the major QTL.

1.Department of Agronomy, Kansas State University, Manhattan, KS 66506, USA 2.Hard Winter Wheat Genetics Research Unit, USDA-ARS, 4008 Throckmorton Hall, Manhattan, KS 66506, USA

... 3 讨论业已报道, 与小麦穗发芽相关的QTL涉及多条染色体和众多区段^{[12,13,15,16,19,22,23]} ...

2011

0.0

张海萍, 冯继明, 常成, 马传喜, 张秀英, 闫长生, 游光霞, 肖世和. 中国小麦地方品种籽粒强休眠特性的主效基因鉴定. 农业生物技术学报, 2011, 19: 270-277

Zhang

H P

, Feng

J M

, Chang

, Ma

C X

, Zhang

X Y

, Yan

C S

, You

G X

, Xiao

S H

. Investigation of main loci contributing to strong seed dormancy of Chinese wheat land race. J Agric Biotechnol, 2011, 19: 270-277 (in Chinese with English abstract)

前人研究表明,我国一些小麦地方品种籽粒休眠性强,穗发芽抗性好,可能受1～2对主效基因控制.本文利用小麦(Triticum aestivum)2个重组自交系群体(RILs,万县白麦子/京411,万县白麦子/中优9507)进行2点4年的田间试验,以期发掘控制我国小麦地方品种(万县白麦子)籽粒休眠的主效QTL位点.通过复合区间作图进行分析,分别在3AS和3BL染色体上鉴定出2个主效QTL,前者分布在Xbarc57和Xbarc294标记区间(在2个RILs群体中遗传距离分别为5.8和8.5 cM),在多年多点的实验中可解释25.6%～48.3%的表型变异;后者分布在Vp1和Xwmc446标记区间,在万县白麦子/中优9507群体中的遗传距离为8.1 cM,可解释23.5%～37.8%的表型变异.上述研究表明,我国小麦地方品种万县白麦子籽粒强休眠特性主要受Osd.ahau-3A、Qsd.ahau-3B这两个主效基因控制,在不同环境中表现出较好的遗传稳定性.可通过聚合育种的方法获得籽粒休眠性强、穗发芽抗性好的小麦新品种.

... 0%, 而在张海萍等^[14]的试验中为25 ...

2010

0.0

朱占玲, 田宾, 刘宾, 谢全刚, 田纪春. 小麦整穗发芽的QTL定位分析. 山东农业科学, 2010, (6): 19-23

Zhu

Z L

, Tian

, Liu

, Xie

Q G

, Tian

J C

. Quantitative trait loci analysis for pre-harvest sprouting using intact spikes in wheat (Triticum aestivum L. ). Shand ong Agric Sci, 2010, (6): 19-23 (in Chinese with English abstract)

为了研究小麦穗发芽的抗性水平,揭示其遗传机理,并筛选抗性较强的家系用于育种实践,利用黄淮海地区主要推广的两个小麦品种花培3号/豫麦57构建的DH群体和基于混合线性模型的QTLNetwork 2.0软件,对3种环境下的小麦整穗发芽进行了QTL定位分析.3种环境条件下分别检测到3、3、2个与整穗发芽相关的加性QTL位点,这些位点分别位于1B、2B、4A和5D染色体上,总共可解释23.28%、21.83%和11.55%的表型变异.在所有检测到的QTL位点中,只有qPhs5D.1位点在3个环境中均能检测到,剩余位点只能在单独一个环境中检测到.3种环境条件下分别检测到2对、1对和2对上位性位点,总共可解释8.01%、9.50%和21.67%的表型变异,不同环境条件下,检测到的上位性位点均不相同.结果表明,小麦整穗发芽的遗传同时受加性效应和上位性效应控制,且易受环境条件的影响.

... 3 讨论业已报道, 与小麦穗发芽相关的QTL涉及多条染色体和众多区段^{[12,13,15,16,19,22,23]} ...

2008

3.251

0.0

Chen

C X

, Cai

S B

, Bai

G H

. A major QTL controlling seed dormancy and pre-harvest sprouting resistance on chromosome 4A in a Chinese wheat land race. Mol Breed, 2008, 21: 351-358

Wheat pre-harvest sprouting (PHS) can cause significant reduction in yield and end-use quality of wheat grains in many wheat-growing areas worldwide. To identify a quantitative trait locus (QTL) for PHS resistance in wheat, seed dormancy and sprouting of matured spikes were investigated in a population of 162 recombinant inbred lines (RILs) derived from a cross between the white PHS-resistant Chinese landrace Totoumai A and the white PHS-susceptible cultivar Siyang 936. Following screening of 1,125 SSR primers, 236 were found to be polymorphic between parents, and were used to screen the mapping population. Both seed dormancy and PHS of matured spikes were evaluated by the percentage of germinated kernels under controlled moist conditions. Twelve SSR markers associated with both PHS and seed dormancy were located on the long arm of chromosome 4A. One QTL for both seed dormancy and PHS resistance was detected on chromosome 4AL. Two SSR markers, Xbarc 170 and Xgwm 397 , are 9.14 cM apart, and flanked the QTL that explained 28.3% of the phenotypic variation for seed dormancy and 30.6% for PHS resistance. This QTL most likely contributed to both long seed dormancy period and enhanced PHS resistance. Therefore, this QTL is most likely responsible for both seed dormancy and PHS resistance. The SSR markers linked to the QTL can be used for marker-assisted selection of PHS-resistant white wheat cultivars.

1.Department of Plant Pathology, Kansas State University, Manhattan, KS 66506, USA 2.Food Crop Institute, Jiangsu Academy of Agricultural Science, Nanjing, 210014 China 3.Department of Agronomy, Kansas State University, Manhattan, KS 66506, USA 4.Plant Science and Entomology Research Unit, USDA-ARS, Kansas State University, 4008 Throckmorton Hall, Manhattan, KS 66506, USA

... 3 讨论业已报道, 与小麦穗发芽相关的QTL涉及多条染色体和众多区段^{[12,13,15,16,19,22,23]} ...

... 本研究检测到的wmc617 (4AL)、gwm610 (4AL)和barc347 (5DL)与Chen等^[16]、Torada等^[21]和Munkvold等^[28]定位的QTL具有相同标记, 但表型效应较小 ...

2011

3.251

0.0

Liu

S B

, Bai

G H

, Cai

S B

, Chen

C X

. Dissection of genetic components of preharvest sprouting resistance in white wheat. Mol Breed, 2011, 27: 511-523

Preharvest sprouting (PHS) in rain-affected wheat ( Triticum aestivum ) is a major constraint to the production of high-quality wheat, especially in regions where white grain wheat cultivars are preferred. To characterize quantitative trait loci (QTLs) for PHS resistance and seed dormancy (SD), we evaluated 162 recombinant inbred lines developed from the cross between PHS-resistant white wheat landrace Tutoumai A and PHS-susceptible white wheat cultivar ‘Siyang 936’ for PHS resistance and SD in field and greenhouse experiments. Composite interval mapping (CIM) identified four QTLs for PHS resistance and long SD that explained up to 45 and 40.8% of the phenotypic variation in five PHS and four SD experiments, respectively. Qphs.pseru - 4A.1 was detected in three of the five PHS experiments, and Qphs.pseru - 5B.1 , Qphs.pseru - 5B.2 , and Qphs.pseru - 4B.1 were detected in two of the five PHS experiments, respectively. All four QTLs for PHS resistance also affected SD. Qphs.pseru - 4A.1 was significant in all four SD experiments; the other three QTLs were detected only in one experiment. Additive and epistatic effects were observed for PHS resistance and SD. Besides three additive QTLs for PHS resistance and two for long SD, an additional 11 and 10 QTLs were detected with epistatic effects on PHS resistance and SD, respectively. The major genetic component of PHS resistance was SD, and other genetic factors may also contribute to PHS resistance in this population.

1.Department of Agronomy, Kansas State University, Manhattan, KS 66506, USA 2.USDA-ARS, Hard Winter Wheat Genetics Research Unit, 4008 Throckmorton Hall, Manhattan, KS 66506, USA

2005

3.658

0.0

Mares

D J

, Mrva

, Cheong

, Williams

, Watson

, Storlie

, Sutherland

, Zou

. A QTL located on chromosome 4A associated with dormancy in white- and red grained wheats of diverse origin. Theor Appl Genet, 2005, 111: 1357-1364

Improved resistance to preharvest sprouting in modern bread wheat ( Triticum aestivum . L.) can be achieved via the introgression of grain dormancy and would reduce both the incidence and severity of damage due to unfavourable weather at harvest. The dormancy phenotype is strongly influenced by environmental factors making selection difficult and time consuming and this trait an obvious candidate for marker assisted selection. A highly significant Quantitative Trait Locus (QTL) associated with grain dormancy and located on chromosome 4A was identified in three bread wheat genotypes, two white- and one red-grained, of diverse origin. Flanking SSR markers on either side of the putative dormancy gene were identified and validated in an additional population involving one of the dormant genotypes. Genotypes containing the 4A QTL varied in dormancy phenotype from dormant to intermediate dormant. Based on a comparison between dormant red- and white-grained genotypes, together with a white-grained mutant derived from the red-grained genotype, it is concluded that the 4A QTL is a critical component of dormancy; associated with at least an intermediate dormancy on its own and a dormant phenotype when combined with the R gene in the red-grained genotype and as yet unidentified gene(s) in the white-grained genotypes. These additional genes appeared to be different in AUS1408 and SW95-50213.

1.School of Agriculture and Wine, University of Adelaide, Waite Campus, Glen Osmond, SA 5064, Australia 2.SARDI, GPO Box 397, Adelaide, SA 5001, Australia 3.Faculty of Sciences, University of Southern Queensland, Toowoomba, QLD 4350, Australia 4.Sichuan Academy of Agricultural Science, Chengdu City, Sichuan 610066, China

2008

3.658

0.0

Ogbonnaya

F C

, Imtiaz

, Ye

, Hearnden

P R

, Hernand ez

, Eastwood

R F

, Van

Ginkel M

, Shorter

S C

, Winchester

J M

. Genetic and QTL analysis of seed dormancy and preharvest sprouting resistance in the wheat germplasm CN10955. Theor Appl Genet, 2008, 116: 891-902

The inheritance and genetic linkage analysis for seed dormancy and preharvest sprouting (PHS) resistance were carried out in an F 8 recombinant inbred lines (RILs) derived from the cross between “CN19055” (white-grained, PHS-resistant) with locally adapted Australian cultivar “Annuello” (white-grained, PHS-susceptible). Seed dormancy was assessed as germination index (GI7) while assessment for preharvest sprouting resistance was based on whole head assay (sprouting index, SI) and visibly sprouted seeds (VI). Segregation analysis of the F 2 , F 3 data from the glasshouse and the RIL population in 2004 and 2005 field data sets indicated that seed dormancy and PHS resistance in CN19055 is controlled by at least two genes. Heritabilities for GI7 and VI were high and moderate for SI. The most accurate method for assessing PHS resistance was achieved using VI and GI7 while SI exhibited large genotype by environment interaction. Two quantitative trait loci (QTLs) QPhs.dpivic.4A.1 and QPhs.dpivic.4A.2 were identified. On pooled data across four environments, the major QTL, QPhs.dpivic.4A.2 , explained 45% of phenotypic variation for GI7, 43% for VI and 20% for SI, respectively. On the other hand, QPhs.dpivic.4A.1 which accounted for 31% of the phenotypic variation in GI7 in 2004 Horsham field trial, was not stable across environments. Physical mapping of two SSR markers, Xgwm937 and Xgwm894 linked to the major QTL for PHS resistance, using Chinese Spring deletions lines for chromosome 4AS and 4AL revealed that the markers were located in the deletion bins 4AL-12 and 4AL-13. The newly identified SSR markers ( Xgwm937 / Xgwm894 ) showed strong association with seed dormancy and PHS resistance in a range of wheat lines reputed to possess PHS resistance. The results suggest that Xgwm937 / Xgwm894 could be used in marker-assisted selection (MAS) for incorporating preharvest sprouting resistance into elite wheat cultivars susceptible to PHS.

1.Department of Primary Industries, Primary Industries Research Victoria (PIRVic), Private Bag 260, Horsham, VIC 3401, Australia 2.Molecular Plant Breeding CRC, Latrobe University, Suite 21, 2 Park Drive, Bundoora, VIC 3080, Australia 3.International Centre for Agricultural Research in the Dry Areas (ICARDA), P.O. Box 5466, Aleppo, Syria 4.School of Agriculture and Wine, University of Adelaide, Waite Campus, Glen Osmond, SA 5064, Australia 5.International Maize and Wheat Improvement Center (CIMMYT), El Batan, Mexico 6.Australian Grain Technology, Private Bag 260, Horsham, VIC 3401, Australia 7.NZ Institute for Crop and Food Research Limited, Private Bag 4704, Christchurch, New Zealand

... 3 讨论业已报道, 与小麦穗发芽相关的QTL涉及多条染色体和众多区段^{[12,13,15,16,19,22,23]} ...

2008

3.251

0.0

Zhang

X Q

, Li

C D

, Tay

, Lance

, Mares

, Cheong

, Cakir

, Ma

J H

, Appels

. A new PCR-based marker on chromosome 4AL for resistance to pre-harvest sprouting in wheat (Triticum aestivum L. ). Mol Breed, 2008, 22: 227-236

Pre-harvest sprouting (PHS) is a complex trait controlled by multiple genes with strong interaction between environment and genotype that makes it difficult to select breeding materials by phenotypic assessment. One of the most important genes for pre-harvest sprouting resistance is consistently identified on the long arm of chromosome 4A. The 4AL PHS tolerance gene has therefore been targeted by Australian white-grained wheat breeders. A new robust PCR marker for the PHS QTL on wheat chromosome 4AL based on candidate genes search was developed in this study. The new marker was mapped on 4AL deletion bin 13-0.59-0.66 using 4AL deletion lines derived from Chinese Spring. This marker is located on 4AL between molecular markers Xbarc170 and Xwg622 in the doubled-haploid wheat population Cranbrook × Halberd. It was mapped between molecular markers Xbarc170 and Xgwm269 that have been previously shown to be closely linked to grain dormancy in the doubled haploid wheat population SW95-50213 × Cunningham and was co-located with Xgwm269 in population Janz × AUS1408. This marker offers an additional efficient tool for marker-assisted selection of dormancy for white-grained wheat breeding. Comparative analysis indicated that the wheat chromosome 4AL QTL for seed dormancy and PHS resistance is homologous with the barley QTL on chromosome 5HL controlling seed dormancy and PHS resistance. This marker will facilitate identification of the gene associated with the 4A QTL that controls a major component of grain dormancy and PHS resistance.

1.CRC Molecular Plant Breeding/DAFWA, State Agricultural Biotechnology Center, Murdoch University, Perth, WA 6150, Australia 2.Department of Agriculture and Food, Western Australia, 3 Baron-Hay Court, South Perth, WA 6151, Australia 3.School of Agriculture, Food and Wine, University of Adelaide, Waite Campus, Glen Osmond, SA 5064, Australia 4.SARDI, GPO Box 397, Adelaide, SA 5001, Australia

2005

1.643

0.0

Torada

, Ikeguchi

, Koike

. Mapping and validation of PCR-based markers associated with a major QTL for seed dormancy in wheat. Euphytica, 2005, 143: 251-255

Seed dormancy is one of the important factors controlling pre-harvest sprouting (PHS) resistance in wheat. We identified a major quantitative trait locus (QTL) for seed dormancy on the long arm of wheat chromosome 4A (4AL) via simple sequence repeat (SSR)-based genetic mapping using doubled haploid lines from a cross between Japanese PHS resistant variety ‘Kitamoe’ and the Alpine non-resistant variety “Münstertaler” (K/M). The QTL explained 43.3% of total phenotypic variation for seed dormancy under greenhouse conditions. SSR markers flanking the QTL were assigned to the chromosome long arm fraction length 0.59–0.66 on the basis of chromosome deletion analysis, suggesting that the gene(s) controlling seed dormancy are probably located within this region. Under greenhouse conditions, the QTL explained 28.5 and 39.0% of total phenotypic variation for seed dormancy in Haruyutaka/Leader (HT/L) and OS21-5/Haruyokoi (O/HK) populations, respectively. However, in field conditions, the effect was relatively low or not significant in both the K/M and HT/L populations. These markers were considered to be widely useful in common with various genetic backgrounds for improvement of seed dormancy through the use of marker-assisted selection. Further detailed research using near isogenic lines will be needed to define how this major QTL interacts with environmental conditions in our area.

1.Hokkaido Green-Bio Institute Naganuma Hokkaido 069-1317 Japan 2.Hokuren Agricultural Research Institute Naganuma Hokkaido 069-1316 Japan

... 本研究检测到的wmc617 (4AL)、gwm610 (4AL)和barc347 (5DL)与Chen等^[16]、Torada等^[21]和Munkvold等^[28]定位的QTL具有相同标记, 但表型效应较小 ...

2012

1.643

0.0

Rehman Arif M A, Neumann

, Nagel

, Kobiljski

, Lohwasser

, Börner

. An association mapping analysis of dormancy and pre-harvest sprouting in wheat. Euphytica, 2012, 188: 409-417

Abstract Seed dormancy and pre-harvest sprouting are important traits in bread wheat. Bi-parental populations have permitted the identification of several genes/quantitative trait loci controlling these traits, mapping to various bread wheat chromosomes. Here, we report the use of association mapping to uncover the genetic basis of both traits in a panel of 96 diverse winter wheat cultivars to establish the presence of marker-trait associations on many chromosomes. Potential candidate genes were identified by studying the gene content of the chromosome bins into which the major marker trait associations mapped.

1. Leibniz Institute of Plant Genetics and Crop Plant Research (IPK), Corrensstrasse 3, 06466, Gatersleben, Germany 2. Institute of Field and Vegetable Crops, Maksima Gorkog 30, 21000, Novi Sad, Serbia

... 利用这一方法, Arif等^[22]检测出34个与小麦整穗发芽性状相关的位点, 分别位于1A、2A、2D、4A、5B、5D、6A、6B、7A和7D上, 其中2D、5B和7A是携带穗发芽抗性位点的密集区域 ...

... 3 讨论业已报道, 与小麦穗发芽相关的QTL涉及多条染色体和众多区段^{[12,13,15,16,19,22,23]} ...

2012

3.658

0.0

... Kulwal等^[23]除发现位于1BS、2BS、2BL、2DS、4AL、6DL和7DS上已报道的QTL外, 还鉴定出位于7BS的新位点 ...

... 3 讨论业已报道, 与小麦穗发芽相关的QTL涉及多条染色体和众多区段^{[12,13,15,16,19,22,23]} ...

2007

11.808

0.0

Mackay

, Powell

. Methods for linkage disequilibrium mapping in crops. Trends Plant Sci, 2007, 12: 57-63

Linkage disequilibrium (LD) mapping in plants detects and locates quantitative trait loci (QTL) by the strength of the correlation between a trait and a marker. It offers greater precision in QTL location than family-based linkage analysis and should therefore lead to more efficient marker-assisted selection, facilitate gene discovery and help to meet the challenge of connecting sequence diversity with heritable phenotypic differences. Unlike family-based linkage analysis, LD mapping does not require family or pedigree information and can be applied to a range of experimental and non-experimental populations. However, care must be taken during analysis to control for the increased rate of false positive results arising from population structure and variety interrelationships. In this review, we discuss how suitable the recently developed alternative methods of LD mapping are for crops.

... 4 关联分析参考Mackay等^[24]的方法, 从180对SSR引物中选取随机分布于小麦全基因组的105对独立SSR引物对群体遗传结构进行评估 ...

2005

6.275

0.0

... 采用STRUCTURE软件^[25]的混合模型来确定群体结构, K值取1~11, 每个K值单独运行6次 ...

2007

5.323

0.0

... 利用TASSLE软件^[26]的一般线性模型(general linear model, GLM)和混合线性模型(mixed linear model, MLM), 将群体结构分析所得的Q值和亲缘关系K作为协变量, 进行标记与性状的关联分析, 选取两种模型共同关联到的显著(P参考文自翔等^[27]描述的方法, 对整穗发芽抗性显著关联的标记位点进一步挖掘其优异等位变异, 明确各等位变异的表型效应及其载体材料 ...

2008

0.0

1.667

文自翔, 赵团结, 郑永战, 刘顺湖, 王春娥, 王芳, 盖钧镒. 中国栽培和野生大豆农艺及品质性状与SSR标记的关联分析: II. 优异等位变异的发掘. 作物学报, 2008, 34: 1339-1349

Wen

Z X

, Zhao

T J

, Zheng

Y Z

, Liu

S H

, Wang

C E

, Wang

, Gai

J Y

. Association analysis of a agronomic and quality traits with SSR markers in Glycine max and Glycine soja in China: II. Exploration of elite alleles. Acta Agron Sin, 2008, 34: 1339-1449 (in Chinese with English abstract)

On the basis of our previous results of SSR loci associated with the 16 traits, alleles of loci significantly associated with the traits were analyzed further in the present paper. The phenotypic allele effect was estimated through comparison between the average phenotypic value over accessions with the specific allele and that of accessions with “ allele”, and then the average positive (negative) allele effect (AAE) of a locus was calculated over the estimated phenotypic effects of all positive (negative) alleles. The results showed that a set of elite alleles, loci and their carrier materials were screened out. Among the elite alleles existed in Glycine max and Glycine soja , some were consistent, some inconsistent and some complementary. The phenotypic effects of posi-tive (negative) alleles were different from each other, and therefore, the average positive (negative) effects of a locus were also different from each other, indicating the potential of genetic recombination for breeding purposes. A same marker locus could associate with multiple traits with its alleles performed in their own way in direction and size, and the covariation of a same allele performed in two related traits might be the genetic basis of their phenotypic correlation. In summary, the results implied that association mapping could offer further genetic information complementary to the family-based linkage mapping for the im-provement of breeding procedures. In addition, further detailed information on elite alleles and phenotypic effects of the 16 traits were shown in the text.

Soybean Research Institute of Nanjing Agricultural University / National Center for Soybean Improvement / National Key Laboratory for Crop Ge-netics and Germplasm Enhancement, Nanjing 210095, Jiangsu, China

在前文研究已检出与农艺品质性状显著关联的SSR位点的基础上, 本文进一步对与性状关联位点的等位变异作解析, 通过将携带某等位变异的所有材料表型均值与携带无效等位基因( allele)材料表型均值做比较, 估计等位变异的潜在表型效应增量(减量), 进一步利用该信息估计位点增效(减效)等位变异的平均效应, 鉴别出一批农艺品质性状优异位点、等位变异及携带优异等位变异的载体材料。发现在栽培及野生种质中检出的优异等位变异有同、有异、有互补性。发现关联位点正、负效应等位变异均值间有差异, 可根据育种目标性状选择要求, 选取适合的位点及相应等位变异。同一标记位点可与多性状关联, 其等位变异在不同性状间各有其表型效应的方向和大小; 等位变异在相关性状效应上方向、大小的异同解释了性状间正、负相关的遗传原因。关联作图得到的信息可以弥补家系连锁法QTL定位信息的不足, 并直接利用等位变异信息进行亲本选拔、组合选配及后代等位条带辅助选择以提高育种成效。

2009

3.658

0.0

Munkvold

J D

, Tanaka

, Benscher

, Sorrells

M E

. Mapping quantitative trait loci for preharvest sprouting resistance in white wheat. Theor Appl Genet, 2009, 119: 1223-1235

The premature germination of seeds before harvest, known as preharvest sprouting (PHS), is a serious problem in all wheat growing regions of the world. In order to determine genetic control of PHS resistance in white wheat from the relatively uncharacterized North American germplasm, a doubled haploid population consisting of 209 lines from a cross between the PHS resistant variety Cayuga and the PHS susceptible variety Caledonia was used for QTL mapping. A total of 16 environments were used to detect 15 different PHS QTL including a major QTL, QPhs.cnl-2B.1, that was significant in all environments tested and explained from 5 to 31% of the trait variation in a given environment. Three other QTL QPhs.cnl-2D.1, QPhs.cnl-3D.1, and QPhs.cnl-6D.1 were detected in six, four, and ten environments, respectively. The potentially related traits of heading date (HD), plant height (HT), seed dormancy (DOR), and rate of germination (ROG) were also recorded in a limited number of environments. HD was found to be significantly negatively correlated with PHS score in most environments, likely due to a major HD QTL, QHd.cnl-2B.1, found to be tightly linked to the PHS QTL QPhs.cnl-2B.1. Using greenhouse grown material no overlap was found between seed dormancy and the four most consistent PHS QTL, suggesting that greenhouse environments are not representative of field environments. This study provides valuable information for marker-assisted breeding for PHS resistance, future haplotyping studies, and research into seed dormancy.

1.Department of Plant Breeding and Genetics, Cornell University, 240 Emerson Hall, Ithaca, NY 14853, USA 2.Boyce Thompson Institute for Plant Research, Tower Road, Ithaca, NY 14853, USA 3.Department of Plant Breeding and Genetics, Cornell University, 240 Emerson Hall, Ithaca, NY 14853, USA

... 本研究检测到的wmc617 (4AL)、gwm610 (4AL)和barc347 (5DL)与Chen等^[16]、Torada等^[21]和Munkvold等^[28]定位的QTL具有相同标记, 但表型效应较小 ...

1997

3.658

0.0

... 但比在特定环境下表达的QTL更有育种价值^[29] ...