小麦面粉色泽(白度)与DArT标记的关联分析

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于海霞, 肖静, 田纪春. 小麦面粉色泽(白度)与DArT标记的关联分析. 作物学报, 40(12): 2198-2202
YU Hai-Xia, XIAO Jing, TIAN Ji-Chun. Genome-wide Association Analysis of Flour Color (Whiteness) Using DArT Markers in Common Wheat. Acta Agronomica Sinica, 40(12): 2198-2202 复制到剪切板

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作物学报编辑部

小麦面粉色泽(白度)与DArT标记的关联分析

于海霞¹, 肖静², 田纪春^1,^*

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

²山东省泰安市岱岳区农业局, 山东泰安 271018

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

收稿日期:2014-09-16

基金:本研究由国家自然科学基金项目(30971764)和国家转基因生物新品种培育重大专项(2008ZX08002-003)资助;

摘要

面粉色泽(白度)是评价小麦加工品质的重要指标之一。在4个环境中检测了109份矮孟牛姊妹系及其衍生系的面粉色泽(白度)参数, 并利用混合线性模型分析其与覆盖小麦基因组的971个DArT (Diversity Array Technology)标记的关联性。结果显示, 17个DArT标记与面粉色泽显著关联, 4个标记与白度显著关联(P≤0.001)。标记wPt-1196和wPt-669693在多个环境中与色泽a*和b*关联, 且效应值较大。该研究结果为小麦面粉色泽的分子标记辅助选择育种提供了参考信息。

关键词: 小麦; 面粉色泽(白度); 矮孟牛; 关联分析; DArT标记

Genome-wide Association Analysis of Flour Color (Whiteness) Using DArT Markers in Common Wheat

YU Hai-Xia¹, XIAO Jing², TIAN Ji-Chun^1,^*

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

²Daiyue District Agriculture Bureau of Tai#cod#x02019;an City, Shandong Province, Tai’an 271018, China

Abstract

Flour color (whiteness) is an important index to evaluate wheat quality. Parameters for flour color (L*, a*, and b*) and whiteness of 109 lines derived from wheat variety Aimengniu were evaluated in four environments, and association analysis was carried out using 971 DArT (Diversity Array Technology) markers under the mixed linear model. Twenty-seven DArT markers were found to be associated with flour color (P ≤ 0.001), while seven markers with whiteness. Markers wPt-1196 and wPt-669693 were associated with color a* and b* in multi-environments, and the percentages of phenotypic variation explained were high. These results provide important information for marker-assisted selection of wheat flour color (whiteness).

Keyword: Wheat; Flour color (whiteness); Aimengniu; Association analysis; DArT markers

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面粉色泽(白度)是小麦面粉品质的重要指标之一, 在很大程度上反映面粉的质量和制粉精度, 也是面粉分级的重要指标。在我国, 面粉等级标准对色泽(R457白度)的要求是一级大于76, 二级大于75, 三级大于72^{[ 1]}。馒头、面条等中国传统食品大多都需要较高的白度。

面粉色泽主要受遗传因素影响。Parker等^{[ 2]}用重组自交系, 将控制面粉黄色度(flour b*)基因位点定位在3A和7A, 分别解释13%和60%的表型变异, 确定遗传力为0.68, 表明面粉色泽的表达主要受遗传控制。Ma等^{[ 3]}在第1同源群和7B染色体上检测到亮度和黄色度(L*和b*)的QTL。Kuchel等^{[ 4]}在7B染色体上检测到一个主效QTL, 解释48%~60%的表型变异。Zhang等^{[ 5]}将控制黄色度b*的8个QTL定位于1DS、2DL、3A、4D、5D、6AL、6D和7AL。McCartney等^{[ 6]}将控制黄碱面条亮度(noodle L*)和黄色度(noodle b*)基因分别定位在5B、5D和7AL上。张晓和田纪春^{[ 7]}检测到控制面粉色泽的18个加性位点, 分布在19条染色体上, 其中一个主效QTL与位点Xbarc 372 (0.1 cM)邻近, 对红色度a*的贡献率高达25.64%。Zhang等^{[ 8]}在1A染色体上检测到关于色泽的QTL, 解释1.5%~4.1%的变异。大多数研究集中于色泽L*和b*, 而对同样重要的a*则研究较少。

基于连锁不平衡(linkage disequilibrium, LD)的关联分析通过检测标记-性状关联已成为QTL分析的一个重要手段。近年来, 该技术已在小麦产量、品质、抗病性等遗传学研究上广泛采用^{[ 9, 10, 11, 12]}, 但是品质特性研究多集中于高分子量麦谷蛋白, 而对面粉色泽和白度关注较少, 尤其是使用DArT标记(Diversity Arrays Technology, http:// www.diversityarrays.com/)^{[ 13]}的研究还未见报道。本研究通过关联分析对小麦面粉色泽(白度)进行QTL定位, 希望找到与之紧密连锁的DArT标记, 为分子标记辅助选择和分子设计育种提供参考。

1 材料与方法

1.1 植物材料

选用矮孟牛姊妹系及其亲本和衍生系109份材料, 大部分来源于中国。使用DArT标记进行全基因组扫描, 标记的PIC平均值为0.408。使用STRUCTUE 2.0软件评估群体结构, 可分为4个子群体。整个群体LD的延伸距离为23.6 cM^{[ 14]}。

1.2 表型性状测定方法

分别在山东泰安的山东农业大学农场(2008—2010连续3年)和安徽宿州紫芦湖农场(2010年) 4个田间条件下种植, 采用完全随机区组设计, 2次重复, 行长2 m, 行距0.25 m。按当地小麦产量品种比较试验的要求进行田间管理。成熟期小区混收。

收获后室温储藏3个月, 用布勒实验磨(model-MLU 300 m/s Buhler, Uzwil, 瑞士)制粉。采用WSB-IV型智能白度测定仪测定面粉白度。用美能达CR-310型色差计(日本)测定面粉色泽(L*、a*、b*值), 以L*、a*和b*色空间

表示。L*为亮度(0表示黑色, 100表示白色), a*表示红-绿方向, b*表示黄-蓝方向。每样品重复5次, 每个环境下的观测值为样品平均值。

1.3 数据处理和关联分析

使用SAS 8.0 (SAS Institute Inc., Cary, NC, 美国)进行方差分析(ANOVA)和相关性分析, 用4个环境的面粉色泽(白度)平均值作相关分析。使用TASSEL 2.0.1 (http://www. maizegenetics.net/)计算DArT标记间的连锁不平衡( r²), 排除稀有等位基因( f < 0.10)后进行1000次排列计算。用Structure 2.0计算群体结构数据, 由TASSEL 2.0.1计算Kinship矩阵, 应用mixed linear model进行标记和性状之间的关联分析。当标记的 P≤0.001时认为与性状存在关联。

2 结果与分析

2.1 面粉色泽(白度)的表型变异及参数之间的相关性

分别对4个环境下材料的面粉色泽和白度参数考察, 显示这个群体的表型具有丰富的多样性, 在不同材料中变异幅度较大, 但在不同环境间差异较小, 尤其是2010年2个地点间差异更小。群体结构对面粉白度、a*和b*变异的解释率分别为12.9%、21.2%和14.6%, 而L*受遗传因素影响较小, 表型变异解释率仅为7.3% ( 表1)。除白度和色泽a*正相关外, 参数之间存在不同程度的负相关, 尤其是b*与a*和L*有显著负相关( 表2)。

表1 面粉色泽和白度的表型变异 Table 1 Phenotypic performance for flour color and whiteness

2.2 面粉色泽与白度的关联分析

利用各环境的数据独立寻找遗传关联, 在 P≤0.001水平上确定了34个标记-性状关联(marker-trait association, MTA), 包括20个分子标记, 分布在1A、1B、3B、5A、6A、6B、6D、7A和7B染色体上, 对表型变异的解释率( R²)为6.8%~18.1%。有8个标记关联2个性状, 其他12个标记关联单个性状; 除3个标记在3个环境、1个标记在2个环境中被重复检测到外, 其他均在单一环境下被检测到( 表3)。

表2 面粉色泽与白度参数之间的相关性系数 Table 2 Pairwise correlation between flour color and whiteness

表3 与面粉色泽(白度)关联标记的染色体定位及其对表型变异的解释 Table 3 Chromosomal location and phenotypic variation explained of marker loci associated with flour color (whiteness)

与面粉白度显著关联的标记共4个, 其中位于1A的wPt-731090在3个环境中被检测到, 位于6D的wPt- 666615在2个环境中被检测到。

与面粉色泽L*显著关联的标记有7个, 在1B和6B染色体上均有多个标记位点(2个和3个), 其中位于6B的wPt-730273的表型贡献率高达13.7%。

检测到6个与面粉色泽a*显著关联的标记位点, 其中位于7B的wPt-1196和wPt-669693都在3个环境下被重复检测到( P≤0.0001), 其表型贡献率平均值都超过10%。

与面粉色泽b*显著关联的标记位点最多, 共10个, 在7A和7B染色体上各有3个, 但所有标记仅在单一环境中被检测到。7A上的wPt-1958具有最高的表型解释率(13.3%); 7B上wPt-1196t也是a*的关联标记; 其余标记, 如wPt-731090 (1A)、wPt-6594 (6B)和wPt-5533 (7A)也与白度或L*关联。7A上遗传距离未知的wPt-1958与已定位标记wPt-5533 (221.8 cM)存在较强的LD ( r² > 0.8), 这2个标记均可视为在定位标记附近^{[ 17]}。

3 讨论

面粉色泽是小麦磨粉品质和商品品质的主要指标, 面粉白度直接影响中国馒头和面条的外观品质。早期已有许多面粉色泽的QTL研究, 本研究首次应用DArT标记和关联分析探究面粉色泽(白度)的遗传基础, 这将有助于分子标记辅助育种。我们发现关联标记大多集中在1A、1B、6B、7A和7B上, 与以往QTL定位结果吻合^{[ 2, 3, 4, 7, 13]}。

本研究结果进一步证实色泽b*与a*和L*有显著负相关^{[ 7]}。目前发现的面粉白度、色泽L*和a*相关QTL较少, 所以研究a*与L*的遗传基础对获得理想的面粉色泽很重要。本研究得到了a*和L*的大量MTA, 其中主效位点wPt-1196可解释色泽a*变异的18.1%, 且能在多个环境中被检测到, 是一个值得深入研究的位点。此外仅检测到1个与白度和b*关联的位点, 还需进一步研究。

关联结果与材料群体结构和试验环境高度相关^{[ 15]}, 使用具有较高遗传多样性的材料, 并在多个环境下重复检测, 可有效减少误差。Bordes等^{[ 13]}利用803个标记, 在来自世界各国的小麦群体中检测到与面粉色泽显著关联的位点, 其中wPt-6447为共同关联标记。本研究表明, 7B上的wPt-1196和wPt-669693都在3个环境下与a*和b*显著关联, 且具有高的贡献率(18.1%), 可用于分子标记辅助选择育种。

关联分析主要用于确定控制复杂性状且效应值较小的QTL^{[ 16]}。面粉色泽是受微效多基因控制的数量性状。Tsilo等^{[ 17]}在6条染色体(1A、1B、5A、5B、5D和7B)上发现了QTL簇, 单个标记的表型贡献率较小。本文确定的关联标记与以往研究相比, 大多位于相同染色体, 但位置不同。这可能与作图群体和试验环境、标记类型和数据、分析方法, 以及图谱基因组覆盖率和标记密度等有关。

小麦面粉色泽主要由胚乳颜色确定^{[ 18]}, 色泽组分包括黄色素和棕色素等。黄色素是决定面粉颜色的主要组分, 同时具有一定的抗氧化功能, 对慢性病有一定的抵抗功效^{[ 19, 20]}。加强对黄色素的研究有助于对面粉色泽的理解。另外, 谷物蛋白含量和硬度等同样影响面粉色泽。蛋白含量与色泽水平显著正相关^{[ 21]}, 硬度通常与白度负相关^{[ 22, 23]}。探讨相关性状的遗传基础可能有助于理解色泽的遗传机制。

面粉色泽(白度)的改良包括利用现有白度优良的品种资源, 可采用传统杂交育种和现代生物技术相结合, 选育白度优良的材料。本研究中, 携带优异位点或优异等位变异的材料主要有矮丰3号、鲁麦23、孟县201等^{[ 14]}, 广大育种者在材料选择时可加以参考。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。

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2000

0.0

... 在我国, 面粉等级标准对色泽(R457白度)的要求是一级大于76, 二级大于75, 三级大于72 ^[1] ...

1998

3.658

0.0

Parker

G D

, Chalmers

J K

, Rathjen

A J

, Langride

Mapping loci associated with flour color in wheat (Triticum aestivum L. ). Theor Appl Genet, 1998, 97: 238-245

A1.ARC Special Research Centre for Basic and Applied Plant Molecular Biology, Department of Plant Science, Waite Campus, University of Adelaide, South Australia 5064 Fax: +61 8 83037331, e-mail: gparker@waite.adelaide.edu.au, AU A2.Department of Plant Science, Waite Campus, University of Adelaide, South Australia 5064, AU

An RFLP map constructed using 150 single seed descent (SSD) lines from a cross between two hexaploid wheat varieties (‘Schomburgk’×‘Yarralinka’) was used to identify loci controlling flour colour. Flour colour data were obtained from field trials conducted over two seasons at different sites. The estimated heritability of this trait was calculated as 0.67. Two regions identified in the preliminary analysis on chromosomes 3A and 7A, accounted for 13% and 60% of the genetic variation respectively. A detailed analysis of the major locus on 7A was conducted through fine mapping of AFLP markers identified using bulked segregant analysis (BSA). Seven additional markers were identified by the BSA and mapped to the region of the 7A locus. The applicability of these markers to identify wheat lines with enhanced flour colour is discussed.

... Parker等^[2]用重组自交系, 将控制面粉黄色度(flour b#cod#x0002A ...

1999

0.0

... Ma等^[3]在第1同源群和7B染色体上检测到亮度和黄色度(L#cod#x0002A ...

2006

3.658

0.0

Kuchel

, Langride

, Mosionek

, Williams

, Jefferies

S P.

The genetic control of milling yield, dough rheology and baking quality of wheat. Theor Appl Genet, 2006, 112: 1487-1495

Improving the end-use quality of wheat is a key target for many breeding programmes. With the exception of the relationship between glutenin alleles and some dough rheological characters, knowledge concerning the genetic control of wheat quality traits is somewhat limited. A doubled haploid population produced from a cross between two Australian cultivars ‘Trident’ and ‘Molineux’ has been used to construct a linkage map based largely on microsatellite molecular makers. ‘Molineux’ is superior to ‘Trident’ for a number of milling, dough rheology and baking quality characteristics, although by international standards ‘Trident’ would still be regarded as possessing moderately good end-use quality. This population was therefore deemed useful for investigation of wheat end-use quality. A number of significant QTL identified for dough rheological traits mapped to HMW and LMW glutenin loci on chromosomes 1A and 1B. However, QTL associated with dough strength and loaf volume were also identified on chromosome 2A and a significant QTL associated with loaf volume and crumb quality was identified on chromosome 3A. A QTL for flour protein content and milling yield was identified on chromosome 6A and a QTL associated with flour colour reported previously on chromosome 7B was confirmed in this population. The detection of loci affecting dough strength, loaf volume and flour protein content may provide fresh opportunities for the application of marker-assisted selection to improve bread-making quality.

1.University of Adelaide Australian Grain Technologies Pty Ltd Roseworthy Campus 5371 Roseworthy SA Australia 2.University of Adelaide School of Agriculture and Wine Waite Campus 5064 Glen Osmond SA Australia 3.University of Adelaide Molecular Plant Breeding Cooperative Research Centre Waite Campus 5064 Glen Osmond SA Australia 4.Plant Genomics Building Australian Centre for Plant Functional Genomics Waite Campus 5064 Glen Osmond SA Australia 5.South Australian Research and Development Institute Waite Campus 5064 Glen Osmond SA Australia

... Kuchel等^[4]在7B染色体上检测到一个主效QTL, 解释48%~60%的表型变异 ...

2006

1.643

0.0

Zhang

, He

Z H

, Zhang

A M

, Ginkel

, Ye

G Y.

Pattern analysis on grain yield performance of Chinese and CIMMYT spring wheat cultivars sown in China and CIMMYT. Euphytica, 2006, 147: 409-420

Understanding the relationships among testing environments is essential for better targeting cultivars to production environments. To identify patterns of cultivar, environment, cultivar-by-environment interactions, and opportunities for indirect selection for grain yield, a set of 25 spring wheat cultivars from China and the International Maize and Wheat Improvement Center (CIMMYT) was evaluated in nine environments in China and four management environments at CIMMYT in Cd. Obregon, Mexico, during two wheat seasons. Genetic background and original environment were the main factors influencing grain yield performance of the cultivars. Baviacora M 92, Xinchun 2 and Xinchun 6 showed relatively more stable and higher grain yields, whereas highly photoperiod sensitive cultivars Xinkehan 9, Kefeng 6 and Longmai 19 proved consistently inferior across environments, except in Harbin and Keshan, the two high latitude environments. Longmai 26, also from high latitude environments in the north-eastern Heilongjiang province, was however probably not as photoperiodicly sensitive as other cultivars from that region, and produced much higher grain yield and expressed a broader adaptation. None of the environments reported major diseases. Pattern analyses revealed that photoperiod response and planting option on beds were the two main factors underlying the observed interactions for grain yield. The production environment of planting on the flat in Mexico grouped together with Huhhot and Urumqi in both wheat seasons, indicating an indirect response to selection for grain yield in this CIMMYT managed environment could benefit the two Chinese environments. Both the environment of planting on the flat with Chinese Hejin and Yongning, and the three CIMMYT environments planting on raised beds with Chinese Yongning grouped together only in one season, showing that repeatability may not be stable in this case.

1.Institute of Crop Sciences/National Wheat Improvement Center, Chinese Academy of Agriculture Sciences (CAAS), Beijing, 100081 China 2.CIMMYT-China Office, c/o CAAS, Beijing, 100081 China 3.Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing, 100101 China 4.CIMMYT, Apdo. Postal 6-641, 06600 Mexico, D.F. Mexico 5.School of Land and Food, The University of Queensland, Brisbane, Qld 4072, Australia

... Zhang等^[5]将控制黄色度b#cod#x0002A ...

2006

1.175

0.0

McCartney

C A

, Somers

D J

, Lukow

, Ames

, Noll

, Cloutier

, Humphreys

D G

, McCallum

B D

. QTL analysis of quality traits in the spring wheat cross RL4452 × “AC Domain”Plant Breed, 2006, 125: 565-575

1 Agriculture and Agri-Food Canada, Cereal Research Centre, 195 Dafoe Rd., Winnipeg, MB R3T 2M9, Canada 2 Corresponding author, E-mail: <a href="mailto:somersd@agr.gc.ca" title="Link to email address" shape="rect">somersd@agr.gc.ca</a>

Wheat grain quality is a complex group of traits of tremendous importance to wheat producers, end-users and breeders. Quantitative trait locus (QTL) analysis studied the genetics of milling, mixograph, farinograph, baking, starch and noodle colour traits in the spring wheat population RL4452/‘AC Domain’. Forty-seven traits were measured on the population and 99 QTLs were detected over 18 chromosomes for 41 quality traits. Forty-four of these QTLs mapped to three major QTL clusters on chromosomes 1B, 4D, and 7D. Fourteen QTLs mapped near Glu-B1, 20 QTLs mapped near a major plant height QTL on chromosome 4D, and 10 QTLs mapped near a major time to maturity QTL on chromosome 7D. Large QTLs were detected for grain and flour protein content, farinograph absorption, mixograph parameters, and dietary fibre on chromosome 2BS. QTLs for yellow alkaline noodle colour parameter L* mapped to chromosomes 5B and 5D, while the largest QTL for the b* parameter mapped to 7AL.

... McCartney等^[6]将控制黄碱面条亮度(noodle L#cod#x0002A ...

2008

0.0

1.889

张晓, 田纪春. 若干高白度小麦的色泽优势及形成因素分析. 中国农业科学, 2008, 41: 347-353

Zhang

, Tian

J C.

The color advantage of Chinese wheat with high whiteness and analysis of factors affecting color formation. Sci Agric Sin, 2008, 41: 347-353 (in Chinese with English abstract)

【objective】In this test, we chose Aobaimai imported from foreign country and Jimai 21 in course of spreading in Shandong province as contrasts. Firstly, we studied the color advantage of Chinese wheat cultivars(line) Youmai3hao, Shannong12, Shannong8355 and 62008 which have good appearance to investigate , and followedly analyzed the main factors influencing the formation of the color, to offer theoretic evidences for the breeding of high whiteness traits of different wheat genotypes. 【Method】We mainly used intelligent brightnessmeter and color and color difference meter to evaluate the color of flour, fresh sheet and steamed bread. 【Result】Results showed that the brightness of the four wheat cultivars(line) was lower than Aobaimai, but the yellowness was also lower than Aobaimai, especially the appearance of steamed bread of 62008 and Shannong12 was better than Aobaimai. The variation of flour color after added with BPO in the four wheat genotypes were smaller than Aobaimai. So we can satisfy the request of high white wheat-based products without BPO by the breeding of wheat varieties. Through the analysis of the affecting factors for the color formation, we found that the main affecting factors of color formation were low PPO activity and yellow pigment content for high-protein content varieties and low protein content and high starch content for low-protein content varieties.【Conclusion】In the breeding selection of wheat color, we can choose wheat lines with low PPO activity and yellow pigment content in wheat materials with high protein content, thus we can acquire wheat cultivars with both high quality and good color to make steamed bread and white noodle. We can also choose wheat lines with low PPO activity and yellow pigment content in wheat materials with low protein content and high starch content. Moreover, we can choose wheat lines with low PPO activity and high yellow pigment content in wheat lines with low protein content and high starch content to make high brightness and yellowness foods.

作物生物学重点实验室/山东农业大学小麦品质育种室

【目的】以进口的高白度品种澳白麦和山东省现在主要推广的济麦21作对照，研究山东省最新育成的几个高自然白度的小麦品种优麦3号、山农12、山农8355及品系62008的色泽优势，对影响面粉及面制品色泽形成的各种因素进行研究，分析其色泽优势形成的主要影响因素，为面粉及其制品色泽性状的改良和高自然白度优质专用小麦品种的选育提供理论依据。【方法】运用智能白度仪和色彩色差计对面粉、面片和馒头的色泽进行评价。【结果】中国的高白度品种（系）具有很大的色泽优势，在面制品色泽上，和对照澳白麦相比，L*值差异不显著，但b*值极显著降低，特别是62008和山农12加工成馒头后L*值比澳白麦高，b*值比对照澳白麦低。加入增白剂后，中国的高白度品种（系）色泽变化的幅度比澳白麦小。对面粉色泽形成影响因素的研究结果表明，蛋白质含量高的品种，其色泽优势形成的主要原因是PPO活性和黄色素含量低；蛋白质含量低的品种，淀粉含量高是其形成高亮度的主要原因；PPO活性和黄色素含量越高，面粉及面制品黄度越大。【结论】在不用增白剂的情况下，通过选育高白度品种，可以满足人们对面制食品高白度的要求。对蛋白质含量高的材料，不过分追求面粉的白度和亮度，但可通过对低PPO活性和低黄色素含量的选择，培育食品色泽优良的高蛋白小麦新品种。对淀粉含量高的材料，既要重视面粉的白度和亮度，又要重视对低PPO活性和低黄色素含量的选择，培育用于加工中国传统食品的高白度小麦新品种。考虑到面食品的多样化，也可以选择蛋白质含量高、PPO活性低、黄色素含量高的特殊材料，培育加工亮黄色食品的品种，丰富中国的小麦品种类型。

... 张晓和田纪春^[7]检测到控制面粉色泽的18个加性位点, 分布在19条染色体上, 其中一个主效QTL与位点Xbarc 372 (0 ...

... 我们发现关联标记大多集中在1A、1B、6B、7A和7B上, 与以往QTL定位结果吻合^[2,3,4,7,13] ...

... 有显著负相关^[7] ...

2009

3.658

0.0

Zhang

L Y

, March

, Tinker

N A

, Belzile

Population structure and linkage disequilibrium in barley assessed by DArT markers. Theor Appl Genet, 2009, 119: 43-52

Diversity Array Technology (DArT) markers were used to investigate the genetic diversity, population structure, and extent of linkage disequilibrium (LD) on a genome-wide level in Canadian barley (Hordeum vulgare L.). Approximately 1,000 DArT markers were polymorphic and scored with high confidence among a collection of 170 barley lines composed mostly of Canadian cultivars and breeding lines. The reproducibility of DArT markers proved very high, as 99.9% of allele calls were identical among seven replicated samples. The polymorphism information content (PIC) of DArT markers ranged between 0.04 and 0.50 with an average of 0.38. Using principal coordinate analysis (PCoA), most lines fell into one of two major groups reflecting inflorescence type (two-row versus six-row). Within these two large groups, evidence of geographic clustering of genotypes was also observed. A cluster analysis Unweighted Pair Group Method with Algorithmic Mean suggested the existence of three subgroups within the two-row group and four subgroups within the six-row group. An analysis of molecular variance (AMOVA) revealed highly significant (P < 0.001) genetic variance within subgroups, among subgroups, and among groups. Values of LD, expressed as r ², declined with increasing genetic distance, and mean values of r ² fell below 0.2 for markers located 2.6 cM apart. Approximately 8% of marker pairs located on the same chromosome and 3.4% of pairs located on different chromosomes were in LD (r ² > 0.2). Within both the subsets of two-row and six-row lines, LD extended slightly further (3.5 cM) than for the entire set, while 7.5% of intra-chromosomal locus pairs and <2% of inter-chromosomal pairs were in LD. We discuss the implications of these findings with regard to the prospects of association mapping of complex traits in barley.

1.Département de Phytologie, Université Laval, 1243 Marchand Bldg, Québec, QC G1K 7P4, Canada 2.Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Central Experimental Farm, 960 Carling Ave, Ottawa, ON K1A 0C6, Canada

... Zhang等^[8]在1A染色体上检测到关于色泽的QTL, 解释1 ...

2001

35.209

0.0

Thornsberry

J M

, Goodman

M M

, Doebley

, Kresovich

, Nielsen

, Buckler

E S.

Dwarg 8 polymorphisms associate with variation in flowering time. Nat Genet, 2001, 28: 286-289

2006

3.658

0.0

Ravel

, Praud

, Murigneux

, Linossier

, Darderet

, Balfourier

, Dufourier

, Dufour

, Brunet

, Charmet

Identification of Glu-B1-1 as a cand idate gene for the quantity of high molecular-weight glutenin in bread wheat (Triticum aestivum L. ) by means of an association study. Theor Appl Genet, 2006, 112: 738-743

A previous study in wheat (Triticum aestivum L.) identified two candidate genes controlling a quantitative trait locus (QTL) for high-molecular-weight glutenin subunit (HMW-GS) GluBx. These candidates were Glu-B1-1, the structural gene coding for Glu1Bx, and the B homoeologous gene coding for SPA (spa-B), a seed storage protein activator. The goal of this study was to identify the best candidate gene for this QTL. Single nucleotide polymorphisms (SNPs) are an abundant source of DNA polymorphisms that have been successfully used to identify loci associated with particular phenotypes. As no linkage disequilibrium was detected between Glu-B1-1 and spa-B, we performed an association study to identify the individual gene responsible for the QTL. Six SNPs, three located in Glu-B1-1 and three in spa-B, were genotyped by mass spectrometry in a collection of 113 bread wheat lines. These lines were also evaluated for protein content as well as the total quantity of HMW-GSs and of each HMW-GS in seed samples from two harvest years. Significant associations were detected only between Glu-B1-1 polymorphism and most of the traits evaluated. Spa-B was unambiguously discarded as a candidate. To our knowledge, this is the first report on an association study that was successfully used to discriminate between two candidate genes.

1.Amélioration et Santé des Plantes (UMR 1095) 63100 Clermont-Ferrand France 2.Biogemma, Les Cézeaux 63170 Aubière France 3.ULICE, Zac des portes de Riom 63200 Riom France 4.INRA-Centre National de Génotypage CP 5708 91057 Evry France

2007

3.658

0.0

Tommasini

, Schnurbusch

, Fossati

, Mascher

, Keller

Association mapping of Stagonospora nodorum blotch resistance in modern European winter wheat varieties. Theor Appl Genet, 2007, 115: 697-708

Association mapping in populations relevant for wheat breeding has a large potential for validating and fine-mapping QTLs identified in F2- or DH (double haploid)-derived populations. In this study, associations between markers in the region of QSng.sfr-3BS, a major QTL for resistance to Stagonospora nodorum glume blotch (SNG), and SNG resistance were investigated by linkage and association analyses. After increasing marker density in 240 F_5:7 recombinant inbred lines (RILs), QSng.sfr-3BS explained 43% of the genetic variance and peaked 0.6 cM proximal from the marker SUN2-3B. Association between SNG resistance and markers mapped in the region of QSng.sfr-3BS was investigated in a population of 44 modern European winter wheat varieties. Two genetically distinct subpopulations were identified within these lines. In agreement with linkage analyses, association mapping by a least squares general linear model (GLM) at marker loci in the region of QSng.sfr-3BS revealed the highest association with SNG resistance for SUN2-3B (p < 0.05). Association mapping can provide an effective mean of relating genotypes to complex quantitative phenotypes in hexaploid wheat. Linkage disequilibrium (r ²) in chromosome 3B extended less than 0.5 cM in 44 varieties, while it extended about 30 cM in 240 RILs, based on 91 SSR and STS marker-pair comparisons. This indicated that the association mapping population had a marker resolution potential at least 390-fold higher compared to the RIL population.

1.Department of Plant Molecular Biology, Institute of Plant Biology, University of Zürich, Zürich, 8008 Switzerland 2.Department of Botany and Plant Sciences, University of California Riverside, 2150 Batchelor Hall, Riverside, CA 92521-0124, USA 3.University of Adelaide, Waite Campus, Australian Centre for Plant Functional Genomics, PMB1, Glen Osmond, SA 5064, Australia 4.Agroscope Changins, Station fédérale de recherches agronomiques, Nyon1, 1260 Switzerland

2009

1.681

0.0

Yao

, Wang

, Liu

, Zhao

, Zheng

Association mapping of agronomic traits on chromosome 2A of wheat. Genetica, 2009, 137: 67-75

Association mapping is a method to test the association between molecular markers and quantitative trait loci (QTL) based on linkage disequilibrium (LD). In this study, the collection of 108 wheat germplasm accessions form China were evaluated for their plant heights, spike length, spikelets per spike, grains per spike, thousand kernel weight and spikelets density in 3 years at three locations. And they were genotyped with 85 SSR markers and 40 EST-SSR markers. The population structure was inferred on the basis of unlinked 48 SSR markers and 40 EST-SSR markers. The extent of LD on chromosome 2A was 2.3 cM. Association of 37 SSR loci on chromosomes 2A with six agronomic traits was analysed with a mixed linear model. A total of 14 SSR loci were significantly associated with agronomic traits. Some of the associated markers were located in the QTL region detected in previous linkage mapping analysis. Our results demonstrated that association mapping can enhance QTL information and achieves higher resolution with short LD extent.

1.Huazhong Agricultural University National Key Laboratory of Crop Genetic Improvement 430070 Wuhan China 2.Beijing Academy of Agricultural and Forestry Sciences Beijing Hybrid Wheat Research Centre 100097 Beijing China

2001

2.088

0.0

... com/) ^[13]的研究还未见报道 ...

... 我们发现关联标记大多集中在1A、1B、6B、7A和7B上, 与以往QTL定位结果吻合^[2,3,4,7,13] ...

... Bordes等^[13]利用803个标记, 在来自世界各国的小麦群体中检测到与面粉色泽显著关联的位点, 其中wPt-6447为共同关联标记 ...

2012

0.0

1.667

... 6 cM ^[14] ...

... 本研究中, 携带优异位点或优异等位变异的材料主要有矮丰3号、鲁麦23、孟县201等^[14], 广大育种者在材料选择时可加以参考 ...

2006

4.389

0.0

... 关联结果与材料群体结构和试验环境高度相关^[15], 使用具有较高遗传多样性的材料, 并在多个环境下重复检测, 可有效减少误差 ...

1996

0.0

Risch

, Merikangas

The future of genetic studies of complex human disease. Science, 1996, 273: 1516-1517

N. Risch is in the Department of Genetics, Stanford University School of Medicine, Stanford, CA 94305-5120, USA. E-mail: risch{at}lahmed.stanford.edu. K. Merikangas is in the Departments of Epidemiology and Psychiatry Unit, Yale University School of Medicine, New Haven, CT06510, USA. E-mail: kath{at}zeus.psych.yale.edu

The identification of the genetic basis of complex human diseases such as schizophrenia and diabetes has proven difficult.In their Perspective, Risch and Merikangas propose that we can best accomplish this goal by combining the power of the humangenome project with association studies, a method for determining the basis of a genetic disease.

... 关联分析主要用于确定控制复杂性状且效应值较小的QTL^[16] ...

2011

1.513

0.0

... 8), 这2个标记均可视为在定位标记附近^[17] ...

... Tsilo等^[17]在6条染色体(1A、1B、5A、5B、5D和7B)上发现了QTL簇, 单个标记的表型贡献率较小 ...

1962

1.251

0.0

... 小麦面粉色泽主要由胚乳颜色确定^[18], 色泽组分包括黄色素和棕色素等 ...

1992

1.251

0.0

2002

4.519

0.0

Meydani

Antioxidants in the prevention of chronic diseases. Nutr Clin Care, 2002, 5: 47-49

Director, Vascular Biology Laboratory. USDA Human Nutrition. Research Centre of Aging, Tufts University, Boston, Massachusetts.

1984

0.0

Miskelly

D M.

Flour components affecting paste and noodle colour. J Sci Food Agric, 1984, 35: 463-471

Bread Research Institute of Australia, North Ryde, N.S.W., 2113, Australia

The influence of components contributing to the colour and brightness of flour, flour paste and Chinese and Japanese style noodles has been investigated. Differences in brightness and yellowness were attributable to wheat cultivar, milling extraction rate, protein content, starch damage and brown and yellow pigments.

... 蛋白含量与色泽水平显著正相关^[21], 硬度通常与白度负相关^[22,23] ...

1985

1.251

0.0

2001

0.0

1.007

周艳华, 何中虎. 小麦品种磨粉品质研究概况. 麦类作物学报, 2001, 21(4): 91-95

Zhou

Y H

, He

Z H.

Review on wheat milling quality. J Triticeae Crops, 2001, 21(4): 91-95 (in Chinese with English abstract)

Wheat milling quality traits including flour yield, flour color and ash content and its major affecting factors were reviewed. Flour yield was controlled by multi genes located on 3A, 7D and 5A chromosome. Flour color was controlled by one or two major g

本文对出粉率，粉色和灰分三个主要磨粉品质性状及其影响因素进行了述。出粉率受多基因控制，分别位于3A，7D和5A染色体上，粉色的遗传较简单，受1个或2个基因控制，灰分含量主要受蛋白质含量及环境影响；因此有可能通过遗传改良提高小麦出粉率，改善面粉色泽。在影响磨粉品质的诸多因素中，主要讨论了籽粒形状与磨粉品质之间的关系。一般而言，籽度大，容粒大，容重高，圆形或椭圆形且籽粒硬的小麦出粉率高。粒长，粒宽，粒重等性状受多基因控制，籽籽硬度是影响磨粉品质和烘焙品质最重要的籽粒性状，由1对或2对主效基因和修饰基因控制。