硬粒小麦品种<italic>Lpx-B1</italic>位点等位变异的分子鉴定及其脂肪氧化酶活性

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张福彦, 尚晓丽, 吴培培, 宋双, 陈锋, 崔党群. 硬粒小麦品种Lpx-B1位点等位变异的分子鉴定及其脂肪氧化酶活性 . 作物学报, 2014, 40(8): 1364-1370[ZHANG Fu-Yan, SHANG Xiao-Li, WU Pei-Pei, SONG Shuang, CHEN Feng, CUI Dang-Qun. Molecular Identification of Alleles onLpx-B1 Locus and Lipoxygenase Activity in Durum Wheat (Triticum turgidum L.) . ACTA AGRONOMICA SINICA, 2014, 40(8): 1364-1370] 复制到剪切板

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硬粒小麦品种Lpx-B1位点等位变异的分子鉴定及其脂肪氧化酶活性

张福彦^1,^2,^*, 尚晓丽^1,^*, 吴培培¹, 宋双¹, 陈锋^1,^*, 崔党群^1,^*

¹ 河南农业大学农学院 / 河南省粮食作物协同创新中心 / 小麦玉米作物学国家重点实验室, 河南郑州 450002

² 河南省核农学重点实验室 / 河南省科学院同位素研究所有限责任公司, 河南郑州 450015

**同等贡献(Contributed equally to this work)

收稿日期:2014-04-16

基金:本研究由国家重点基础研究计划(973计划)项目(2014CB138105), 教育部新世纪优秀人才项目(NCET-13-0776), 河南省高校青年骨干教师资助计划项目(2011GGJS-044)和河南省现代农业产业技术体系建设专项(Z2010-01-04)资助;

摘要

硬粒小麦籽粒中脂肪氧化酶(LOX)与硬粒小麦面制品的加工品质关系密切相关, 而Lpx-B1位点不同变异类型对LOX活性有重要影响。对来自不同国家和地区的167份硬粒小麦品种的LOX活性进行测定, 并对其Lpx-B1位点不同变异类型进行分子鉴定。不同品种间LOX活性差异明显, 变幅为0.20~7.98 AU min^-1 g^-1。在Lpx-B1.1位点鉴定出3种等位变异, 分别为Lpx-B1.1a、Lpx-B1.1b和Lpx-B1.1c, 以Lpx-B1.1a所占比例最高(55.1%), 其次为Lpx-B1.1c (37.1%), 而Lpx-B1.1b仅占7.8%;Lpx-B1.2和Lpx-B1.3二者总是互补出现在不同的品种中, 146份品种为Lpx-B1.2型, 其余21份品种均为Lpx-B1.3型, 表明二者可能互为一对等位因子。在Lpx-B1.1位点的3种等位变异类型中,Lpx-B1.1b类型品种的LOX活性显著高于Lpx-B1.1a和Lpx-B1.1c类型品种, 而Lpx-B1.1c类型品种的LOX活性最低。Lpx-B1.3类型品种的LOX活性显著高于Lpx-B1.2类型的品种。参试品种共有6种Lpx-B1基因型组合, 其中Lpx-B1.1b/Lpx-B1.3基因型的LOX活性显著高于其他基因型, 而Lpx-B1.1c/Lpx-B1.2和Lpx-B1.1c/Lpx-B1.3基因型的LOX活性最低。这些观测结果为硬粒小麦品质育种提供了重要信息。

关键词: 硬粒小麦; Lpx-B1位点; 等位变异; 脂肪氧化酶

Molecular Identification of Alleles onLpx-B1 Locus and Lipoxygenase Activity in Durum Wheat (Triticum turgidum L.)

ZHANG Fu-Yan^1,^2,^**, SHANG Xiao-Li^1,^**, WU Pei-Pei¹, SONG Shuang¹, CHEN Feng^1,^*, CUI Dang-Qun^1,^*

¹ College of Agronomy / Collaborative Innovation Center of Henan Grain Crops / National Key Laboratory of Wheat & Maize Crop Science, Henan Agricultural University, Zhengzhou 450002, China

² Henan Key Laboratory of Nuclear Agricultural Sciences / Isotope Institute Co., Ltd, Henan Academy of Sciences, Zhengzhou 450015, China

Fund:

Abstract

Lipoxygenase (LOX) has a close relationship to the processing quality of flour products in durum wheat (Triticum turgidum L.). Activity of LOX is greatly influenced by the locus ofLpx-B1. TheLpx-B1 alleles in 167 durum wheat varieties from different countries or regions were identified using gene-specific primers, and theLpx-B1 genotypes were analyzed based on LOX activity data. The LOX activity varied greatly in the 167 varieties, ranging from 0.20 to 7.98 AU min^-1 g^-1. OnLpx-B1.1 locus, three alleles (Lpx-B1.1a,Lpx-B1.1b, andLpx-B1.1c) were identified with frequencies of 55.1%, 7.8%, and 37.1%, respectively. LociLpx-B1.2andLpx-B1.3were always reciprocally present in varieties, in whichLpx-B1.2 was presented in 146 varieties andLpx-B1.3 in the remaining 21 varieties. This result indicates the allelic possibility ofLpx-B1.2andLpx-B1.3. In the threeLpx-B1.1 genotypes, theLpx-B1.1bvarieties had significantly higher LOX activity than theLpx-B1.1a andLpx-B1.1c varieties, and varieties carryingLpx-B1.1callele possessed the lowest LOX activity. The LOX activity inLpx-B1.3 genotypes was significantly higher than that inLpx-B1.2genotypes. A total of six genotypic combinations were found on the threeLpx-B1 loci in 167 varieties. Among them,Lpx-B1.1b/Lpx-B1.3combination had the highest LOX activity and combinations ofLpx-B1.1c/Lpx-B1.2 andLpx-B1.1c/Lpx-B1.3had the lowest LOX activity. These results are informative to quality breeding of durum wheat.

Keyword: Triticum turgidum L.; Lpx-B1locus; Alleles; Lipoxygenase

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脂肪氧化酶(lipoxygenase, LOX)是一类不含血红素铁的蛋白质, 广泛存在于各种类型的植物中^{[ 1, 2]}。它在植物的生长、发育、衰老、脂质过氧化作用、光合作用及抗逆性等方面起重要作用^{[ 2]}。小麦籽粒中的LOX含量很低, 但其在酶促脂肪氧化过程中产生的活性氧, 不仅能通过降解色素类物质影响面粉及面条、馒头和面包等面制品颜色^{[ 3]}, 且能有效改善面制品的营养品质和加工品质^{[ 4, 5, 6]}。因而, 小麦脂肪氧化酶活性及其在品质改良中的重要作用已成为国内外学者研究的热点之一。

小麦籽粒LOX活性是一个受多基因控制的复杂数量性状, 受基因型和环境及其互作的显著影响, 且基因型的效应大于环境及互作效应^{[ 7, 8]}。Leenhardt等^{[ 9]}发现普通小麦籽粒中的LOX活性高于硬粒小麦和一粒小麦, 分别是其2.5倍和7.5倍, 表明小麦类型也是影响LOX活性的因素之一。Hart和Langston^{[ 10]}利用中国春缺体-四体系对LOX同工酶定位表明, Lpx-A1、 Lpx-B1和 Lpx-D1分别位于4AL、4BL和4DS染色体, 而 Lpx-A2、 Lpx-B2和 Lpx-D2分别位于5AL、5BL和5DL, 推测小麦LOX基因主要位于第4和第5同源群。Hsieh等^{[ 11]}从硬粒小麦胚乳中分离出3个与LOX活性有关的同工酶, 分别命名为lipoxygeanses-1 (L-1)、lipoxygeanses-2 (L-2)和lipoxygeanses-3 (L-3)。硬粒小麦中, 脂肪氧化酶由 Lpx编码, 其主效基因分别被定位在4A^{[ 12]}、4B^{[ 12, 13, 14, 15, 16, 17, 18]}、4D^{[ 18, 19]}和5D^{[ 19]}染色体上, Lpx-B1位点的等位变异对LOX活性有显著影响^{[ 3]}。Carrera等^{[ 14]}分析7个硬粒小麦品种的 Lpx-B1基因及其对LOX活性的影响发现, Lpx-B1位点部分片段重叠和 Lpx-B1.1基因部分片段缺失的现象, 并将这2个基因分别命名为 Lpx-B1.1(DQ474240)和 Lpx-B1.2(DQ474241), 且后者与LOX活性降低密切相关。此外, Verlotta等^{[ 15]}还报道了一个控制LOX活性的新基因, 将其命名为 Lpx-B1.3(HM126469), 同时认为 Lpx-B1.2和 Lpx-B1.3基因可能是一对等位基因; Lpx-B1.1位点存在3个等位变异, 分别命名为 Lpx-B1.1a、 Lpx-B1.1b和 L px-B1.1c, 并开发了相应的功能标记。

硬粒小麦是世界第二大麦类作物, 种植面积占世界小麦总面积的10%左右, 其蛋白质和面筋含量较高, 富含大量胡萝卜素, 具有较高的营养价值, 在欧洲, 拉丁美洲的许多国家很受欢迎。硬粒小麦在储藏和加工过程中脂肪氧化酶的氧化降解会导致胡萝卜素和类胡萝卜素大量损失, 降低部分营养价值^{[ 20, 21]}。郑文寅等^{[ 22]}分析了小麦全麦粉的色泽性状与LOX和类胡萝卜素含量之间的关系, 结果表明, LOX活性与类胡萝卜素含量呈极显著负相关, 而与全麦粉的白度、亮度和黄度无显著相关。本研究旨在明确硬粒小麦籽粒中 Lpx-B1基因的等位变异与LOX活性之间的关系, 为利用分子标记选择相应LOX活性的硬粒小麦品种提供理论依据。

1 材料与方法

1.1 试验材料

167份硬粒小麦(意大利博洛尼亚大学农业环境科技学院的Roberto Tuberosa教授提供)来自不同国家和地区, 为历史品种和当前主栽品种, 其中意大利47份、国际玉米小麦改良中心(CIMMYT) 12份、叙利亚80份、西班牙17份、摩洛哥11份。2010—2011年生长季种植于河南农业大学科教示范园区, 2次重复, 常规田间管理。所有材料在当地均能正常成熟, 且未发现有倒伏和穗发芽现象。

1.2 小麦籽粒LOX活性的测定

取完熟后籽粒, 用分光光度计法^{[ 16]}测定籽粒中LOX活性, 每份材料3次重复, 求平均值。操作步骤主要有LOX酶的提取、亚油酸底物的配制(现用现配)、分光光度计检测。所用试剂包括40 mmol L^-1 PBS缓冲液(pH 6.8)、亚油酸(Sigma, L1376)、Tween 20、1 mol L^-1 NaOH、0.1 mol L^-1 NaOH, 以及ddH₂O。使用UV-2600型紫外可见分光光度计(尤尼柯上海仪器有限公司)在234 nm处用1 cm光程石英比色皿测定共轭过氧化物的吸光度∆A, 以每毫克总蛋白每分钟吸光值增加0.001为一个酶活性单位(AU min^-1 g^-1), 用经过高温煮沸LOX粗酶液底物做对照。

1.3 Lpx-B1等位变异检测

从每份试验材料取3粒有代表性的种子, 用锤子砸碎后放入2.0 mL离心管, 参照单籽粒种子DNA快速提取方法^{[ 23]}提取DNA, 但用SLS (十二烷基肌氨酸钠)代替SDS (十二烷基硫酸钠)。利用Verlotta等^{[ 15]}在硬粒小麦开发的共显性标记 Lpx-B1.1a/b以及显性标记 Lpx-B1.1c、 Lpx-B1.2和 Lpx-B1.3 (表1)检测参试品种4B染色体上 Lpx-B1位点等位变异。所有引物均由生工生物工程(上海)有限公司合成。

在ABI 9700或MJ Research PTC-200中进行PCR扩增, 扩增体系为25 μL, 包含2 μL基因组DNA (50 ng μL^-1)、 0.5 μL dNTP (10 mmol L^-1)、各0.4 μL上、下游引物 (10 pmol)、2.5 μL 10× buffer (20 mmol L^-1 Tris-HCl pH 8.4, 25 mmol L^-1 KCl, 1.5 mmol L^-1 MgCl₂)、0.4 μL Taq DNA聚合酶(2.5 U μL^-1)。以上试剂均购自TIANGEN公司。PCR程序为95℃预变性5 min; 95℃变性30 s, 58~64℃退火30 s (不同引物退火温度见表1), 72℃延伸1 min, 35个循环; 72℃延伸10 min, 4℃保存。

表1 硬粒小麦中检测 Lpx-B1基因等位变异的引物 Table 1 Specific primers for identification of Lpx-B1 alleles in durum wheat

在1×TBE的缓冲液中, 用含有溴化乙锭(EB)的1.0% (w/v)琼脂糖凝胶电泳分离PCR扩增产物, 150 V电压下电泳15 min, 最后在凝胶成像系统中扫描观察, 并对电泳结果统计分析。 Lpx-B1.1a和 Lpx-B1.1b等位变异片段差别较小, 只相差74 bp, 故采用2.0% (w/v)琼脂糖凝胶电泳分离检测, 150 V电压下电泳30 min后扫描观察。

1.4 数据分析

利用SPSS 19.0软件分析LOX活性平均值和标准差等参数, 用LSR方法比较不同基因型组合的LOX活性差异显著性。

2 结果与分析

2.1 Lpx-B1位点变异类型及分布

首先利用 Lpx-B1.1a/b标记检测, 扩增产物为1312 bp和1238 bp两种带型, 分别对应 Lpx-B1.1a和 Lpx-B1.1b等位变异(图1); 部分品种中没有检测到扩增产物, 可能为 Lpx-B1.1c等位变异, 进一步用 Lpx-B1.1c的特异引物检测, 结果Cannizzo、Claudio、Lesina和Norba等7个材料中扩增出1558 bp的条带(图2), 确认为 Lpx-B1.1c变异类型。

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	图1 Lpx-B1.1a/b引物在部分硬粒小麦品种中的扩增结果Fig. 1 Amplification profile of Lpx-B1.1a/b primers in partial varieties of durum wheatM: DL2000; 1: Meridiano; 2: Appio; 3: Cannizzo; 4: Trinakria; 5: Bolenga; 6: Appulo; 7: Cappelli; 8: Torrebianca; 9: Mongibello; 10: Roqueno; 11: Quadrato; 12: Colosseo.

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	图2 Lpx-B1.1c引物在部分硬粒小麦品种中的扩增结果Fig. 2 Amplification profile of Lpx-B1.1c primers in partial varieties of durum wheatM: DL2000; 1: Cannizzo; 2: Claudio; 3: Lesina; 4: West Bred Turbo; 5: Roqueno; 6: Torrebianca; 7: Bravadur; 8: Meridiano; 9: Norba; 10: Mongibello; 11: Colosseo; 12: Younes-1.

在 Lpx-B1位点共发现5种变异类型, 分别为 Lpx-B1.1a、 Lpx-B1.1b、 Lpx-B1.1c、 Lpx-B1.2和 Lpx- B1.3。 L px-B1.1位点有3种等位变异, 其中 Lpx-B1.1a类型品种有92份, 占55.1%, Lpx-B1.1b和 Lpx-B1.1c类型的品种分别为13份和62份, 占7.8%和37.1%。此外, 没有发现同一品种兼有 Lpx-B1.2和 Lpx-B1.32个等位变异, 即二者总是互补出现(表2), 146份品种为 Lpx-B1.2变异类型(占87.4%), 而另外21份品种则为 Lpx-B1.3类型(占12.6%)。

Lpx-B1位点基因型因品种来源不同而异。来自叙利亚和意大利的品种, Lpx-B1位点变异类型较为丰富, 拥有所有的5种变异类型; 而在来自CIMMYT、摩洛哥和西班牙的品种中, 没有发现 Lpx-B1.1b和 Lpx-B1.3两种类型。80份叙利亚材料中, Lpx-B1.1位点 Lpx-B1.1a、 Lpx-B1.1b和 Lpx-B1.1c类型分别有46、8和26份; 而在另一位点, 有65份为 Lpx-B1.2类型, 15份为 Lpx-B1.3类型。47份意大利材料中, Lpx-B1.1位点有19份为 Lpx-B1.1a类型, 5份为 Lpx-B1.1b类型, 23份为 Lpx-B1.1c类型; 另一位点 Lpx-B1.2类型有41份, Lpx-B1.3类型有6份(表3)。结果表明, Lpx-B1.1a、 Lpx-B1.1c和 Lpx-B1.2三种基因型在硬粒小麦中占主导地位。

2.2 Lpx-B1位点不同变异类型对LOX活性的影响

方差分析结果表明, 硬粒小麦籽粒中 Lpx-B1位点不同变异类型间LOX活性有显著差异(图3)。 Lpx-B1.1b和 Lpx-B1.3类型品种的LOX活性最高, 分别为5.74 AU min^-1 g^-1和5.04 AU min^-1 g^-1; Lpx-B1.1c变异类型的LOX活性最小, 仅为0.92 AU min^-1 g^-1; 其他2类品种的LOX活性分别为2.50 AU min^-1 g^-1( Lpx-B1 .1a)和1.75 AU min^-1 g^-1 ( Lpx-B1.2)。

表2 部分硬粒小麦品种的LOX活性和 Lpx-B1基因型鉴定 Table 2 Lpx-B1 genotypes and LOX activities in part of durum wheat varieties

表3 不同国家硬粒小麦种质的 Lpx-B1位点基因型分布 Table 3 Distribution of different Lpx-B1alleles in durum wheat varieties from various countries

167个品种在 Lpx-B1位点共有6种基因型组合, 分别为 Lpx-B1.1a/Lpx-B1.2、 Lpx-B1.1b/Lpx-B1.2、 Lpx-B1.1c/Lpx-B1.2、 Lpx-B1.1a/Lpx-B1.3、 Lpx-B1.1b/ Lpx-B1.3和 Lpx-B1.1c/Lpx-B1.3。其中, Lpx-B1.1b/ Lpx-B1.3基因型组合为高LOX活性类型, 有11个品种(6.6%), 其LOX活性平均值高达6.03 AU min^-1 g^-1; Lpx-B1.1a/Lpx-B1.3和 Lpx-B1.1b/Lpx-B1.2基因型组合为偏高类型, 分别有9个(5.4%)和2个(1.2%)品种, 其LOX活性分别为4.27 AU min^-1 g^-1和4.17 AU min^-1 g^-1; Lpx-B1.1a/Lpx-B1.2基因型组合为偏低类型, 是参试品种中的最主要的类型, 包括83个品种(49.7%), 其LOX活性为2.30 AU min^-1 g^-1; Lpx-B1.1c/Lpx-B1.2基因型组合属于低LOX活性类型, 共有61个品种(36.5%), 其平均LOX活性最低, 仅为0.92 AU min^-1 g^-1; 另外, 仅发现1份 Lpx-B1.1c/Lpx-B1.3组合类型, LOX活性为1.02AU min^-1 g^-1, 也属于低LOX活性类型(表4)。

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图3 硬粒小麦中 Lpx-B1位点不同变异类型的LOX活性LOX活性为同类品种的平均值及标准差; 柱形上面的不同字母表示不同变异类型间有显著性差异( P<0.05)。Fig. 3 LOX activities in different Lpx-B1 genotypes of durum wheatLOX activity is the mean of each variety group, and the error bar shows the standard error; Different letters above columns indicate significant difference among different types of variation at P < 0.05.

表4 硬粒小麦中 Lpx-B1位点不同基因型组合LOX活性比较 Table 4 Comparison of LOX activities of durum wheat varieties with different genotype combinations

3 讨论

面粉或面制品的表观色泽是评价小麦品质的重要指标之一。LOX对面粉色泽有很大影响, 且对小麦品质性状具有双重效应。籽粒中LOX会偶联氧化小麦中的类胡萝卜素, 取代化学漂白剂使小麦粉变白, 提高面粉或面制品的白度; 但是过高的LOX活性则会破坏小麦籽粒中的黄色素, 使小麦粉过白而丧失许多营养成分, 降低麦类食品的营养价值^{[ 3, 6, 22]}。研究表明, LOX与小麦籽粒、面粉及其面制品的储藏特性的关系密切, 降低LOX活性有利于延长硬粒小麦籽粒、通心粉等食品的保存期, 从而提高产品的附加值^{[ 24]}, 降低LOX活性也是长期保存种子的重要方法之一。可见, 低LOX活性可以有效地减轻脂质的氧化反应, 减少籽粒的氧化变质, 延长其储藏期, 从而减少粮食的损失^{[ 6, 25]}。本研究在分析硬粒小麦品种籽粒LOX活性及其基因型分布特点的同时, 也试图筛选出低LOX活性遗传材料, 为今后综合性状评价和硬粒小麦品质育种提供基础信息和种质资源储备。

硬粒小麦中, LOX为受多基因控制的数量性状, Lpx-B1位点变异类型是导致LOX活性变化的重要原因。我们在167份硬粒小麦材料中发现LOX活性变异范围为0.20~7.98 AU min^-1 g^-1, 品种间差异较大。Verlotta等^{[ 15]}把 Lpx-B1不同基因型组合分成3个单体型, 即单体型I ( Lpx-B1.1b/Lpx-B1.3)、单体型II ( Lpx- B1.1a/Lpx-B1.2)和单体型III ( Lpx-B1.1c/Lpx-B1.2), 其LOX活性平均值分别为4.60、2.59和0.12 AU min^-1 g^-1。本研究结果与此类似, LOX活性最高和最低的也分别是 Lpx-B1.1b/Lpx-B1.3和 Lpx-B1.1c/Lpx-B1.2组合类型, 平均LOX活性依次是6.03 AU min^-1 g^-1和0.92 AU min^-1 g^-1, 同时本研究结果也支持 Lpx-B1位点等位变异能够引起LOX活性的显著变化的观点。在硬粒小麦主要消费国家, 低LOX活性的硬粒小麦更受欢迎。研究表明, LOX活性与蛋白质、淀粉、湿面筋等大多数品质参数呈负相关, 且能有效增加面筋强度, 导致面团延展性降低^{[ 26, 27]}。与普通小麦相比, 硬粒小麦中LOX活性更低, 同时由于硬粒小麦中的蛋白质和抗氧化剂化合物也显著高于普通小麦, 因此认为硬粒小麦比普通小麦具有更高的营养价值^{[ 28]}。

在对小麦籽粒LOX活性相关基因的定位研究中, 发现一些与目标性状紧密连锁的SSR标记, 如 Xbcd1262、 Xksud2a、 Xwmc312和 Xgwm251^{[ 13, 29]}, 但这些标记不在基因内部, 不能与目的基因共分离, 因而预测基因型的准确性较低, 在实际应用中存在局限性。功能标记是根据基因内部引起表型性状变异的多态性序列而开发出的一种分子标记, 利用其进行辅助选择育种具有高效、准确、适用性广等特点^{[ 30, 31]}。目前, 功能性标记已广泛用于小麦品质和农艺性状以及抗病性鉴定等领域, 是小麦育种的重要辅助手段之一^{[ 31]}。本试验用功能标记对来自不同国家和地区的167份硬粒小麦品种进行 Lpx-B1位点的分子鉴定, 发现主要为 Lpx-B1.1a、 Lpx-B1.1c和 Lpx-B1.2基因型, 与Verlotta等^{[ 15]}利用85份硬粒小麦品种的检测结果基本一致。结合LOX活性测定, 我们发现 Lpx-B1位点上的2个基因是决定硬粒小麦LOX活性的最为主要因素之一。Verlotta等^{[ 15]}筛选出低LOX活性、综合性状优异的硬粒小麦品种, 可直接用作硬粒小麦育种亲本。与LOX活性相关的功能性标记具有很好的应用前景, 尤其适合快速、准确、简便地判定小麦种质资源中LOX基因型, 并能有效预测表型, 在一定程度上加快硬粒小麦品质改良进程。

4 结论

来自不同国家和地区的167份硬粒小麦品种中 Lpx-B1基因位点有 Lpx-B1.1a、 Lpx-B1.1b、 Lpx-B1.1c、 Lpx-B1.2和 Lpx-B1.3共5种变异类型, 且在同一个品种中 Lpx-B1.1a、 Lpx-B1.1b和 Lpx-B1.1c基因交替出现, 不存在2个或2个以上基因共存现象, 同样 Lpx- B1.2和 Lpx-B1.3基因也表现互补, 似为一对等位基因。根据LOX活性, 将不同等位变异分成高或低LOX活性的变异类型, 前者包括 Lpx-B1.1b和 Lpx-B1.3基因型, 后者包括 Lpx-B1.1c和 Lpx-B1.2基因型。硬粒小麦品种在2个位点上聚合高LOX活性等位变异, 则表现为高LOX活性特征, 反之则表现为低LOX活性特征; 而当2个位点分别有高和低LOX活性等位变异时, 其表现型多属于中间类型。

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

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2001

1.931

0.0

... 脂肪氧化酶(lipoxygenase, LOX)是一类不含血红素铁的蛋白质, 广泛存在于各种类型的植物中^[1,2] ...

2005

0.0

... 脂肪氧化酶(lipoxygenase, LOX)是一类不含血红素铁的蛋白质, 广泛存在于各种类型的植物中^[1,2] ...

... 它在植物的生长、发育、衰老、脂质过氧化作用、光合作用及抗逆性等方面起重要作用^[2] ...

2002

1.513

0.0

... 小麦籽粒中的LOX含量很低, 但其在酶促脂肪氧化过程中产生的活性氧, 不仅能通过降解色素类物质影响面粉及面条、馒头和面包等面制品颜色^[3], 且能有效改善面制品的营养品质和加工品质^[4,5,6] ...

... 硬粒小麦中, 脂肪氧化酶由Lpx编码, 其主效基因分别被定位在4A^[12]、4B^{[12,13,14,15,16,17,18]}、4D^[18,19]和5D^[19]染色体上, Lpx-B1位点的等位变异对LOX活性有显著影响^[3] ...

... 但是过高的LOX活性则会破坏小麦籽粒中的黄色素, 使小麦粉过白而丧失许多营养成分, 降低麦类食品的营养价值^[3,6,22] ...

1991

1.251

0.0

2000

2.088

0.0

2009

0.0

. 2009, (8):5-6

Research and application of lipoxygenase for improvement of wheat quality

小麦脂肪氧化酶活性及其对品质改良的作用是当前国内外小麦育种者关注的课题之一.综述了脂肪氧化酶类型及性质、其影响因素和测定方法、小麦等作物脂肪氧化酶的遗传、脂肪氧化酶对小麦加工品质和储藏品质的影响等.讨论了脂肪氧化酶在小麦品质改良中的应用等问题.

... 但是过高的LOX活性则会破坏小麦籽粒中的黄色素, 使小麦粉过白而丧失许多营养成分, 降低麦类食品的营养价值^[3,6,22] ...

... 可见, 低LOX活性可以有效地减轻脂质的氧化反应, 减少籽粒的氧化变质, 延长其储藏期, 从而减少粮食的损失^[6,25] ...

2008

2.088

0.0

. 2008, 48:494-502 DOI:10.1016/j.jcs.2007.11.007

Distribution along durum wheat kernel of the components involved in semolina color

Abstract A bright yellow colour of pasta is an important qualitative trait for the durum wheat industry. Final colour is the result of the balance between yellow and brown components in semolina. Carotenoid pigments and lipoxygenase (LOX) enzyme are mainly involved in yellowness, whereas peroxidase (POD) and ash affect brown hue. All these components have a different distribution across the kernel, with varietal differences too. This study aimed to evaluate the distribution pattern of carotenoid pigments, α-tocopherol, linoleic acid, and ash content as well as of LOX and POD activities within the kernel of six durum wheat cultivars characterised by different pigment content and hydroperoxidation activity of LOX in semolina. The results confirmed differences in the distribution of these components across the kernel and among varieties. Additionally, this study identified for some components (POD, pigments and bleaching activity of LOX) a higher effect of genotype whereas for others (ash, α-tocopherol, hydroperoxidation activity) a marked effect of the debranning process. These results suggest that improvement of the final semolina colour could be reached both through breeding activity, enabling an early selection of better lines, and through an appropriate debranning process.

... 小麦籽粒LOX活性是一个受多基因控制的复杂数量性状, 受基因型和环境及其互作的显著影响, 且基因型的效应大于环境及互作效应^[7,8] ...

2011

0.0

. 2011, 26(1):11-14

Lipoxygenase activity and its genotype and environment interactions for different wheat varietier

选用了安徽省淮南片小麦区试5个生态点的10个小麦品种(系)为试验材料,测定其全麦粉脂肪氧化酶(LOX)活性.研究了基因型、环境及基因型×环境互作对小麦LOX活性的影响.结果表明,全麦粉LOX活性的变幅为4.27～8.94 nkat·g-1,平均值为6.61 nkat·g-1,LOX活性的基因型和环境差异均达到极显著,基因型效应大于环境及基因型与环境互作效应.同时,选用皖麦48等10个小麦品种(系)为试材,测定其LOX活性等9个品质性状,分析了小麦籽粒LOX活性与若干品质性状的关系.结果表明,全麦粉LOX活性与小麦粉LOX活性呈极显著正相关,与水溶性戊聚糖含量呈显著正相关,小麦粉LOX活性与水溶性戊聚糖含量和总戊聚糖含量呈显著正相关.讨论了脂肪氧化酶在小麦品质改良中的利用等问题.

... 小麦籽粒LOX活性是一个受多基因控制的复杂数量性状, 受基因型和环境及其互作的显著影响, 且基因型的效应大于环境及互作效应^[7,8] ...

2006

2.8

0.0

. 2006, 25:170-176 DOI:10.1016/j.eja.2006.04.010

Genetic variability of carotenoid concentration, and lipoxygenase and peroxidase activities among cultivated wheat species and bread wheat varieties

Abstract Lutein and its stereo-isomer zeaxanthin, are the major carotenoids of wheat grain and function as antioxidants in plants as well as in humans. Dietary intake and serum levels of these xanthophylls have been correlated with a beneficial impact on the development of cataracts, age-related maculopathy and other degenerative diseases. The objective of the present study was to explore the genetic variability of carotenoid concentration and oxidase activity among cultivated diploid (einkorn, T. monococcum L.), tetraploid (durum wheat, T. turgidum L. subsp. durum (Desf.) Husn.) and hexaploid (bread wheat, Triticum aestivum L.) wheat species and bread wheat varieties. Varietal mix of the three cultivated wheat species and 48 varieties of bread wheat adapted to north-western European conditions were analysed. The carotenoid concentration of the wholemeal flour of the three wheat species was determined by high performance liquid chromatography. The extractable activities of lipoxygenase (LOX) and peroxidase, involved in the degradation of carotenoid pigments during the kneading step of bread making, were also determined for the varietal mix of the three cultivated wheat species. For the three cultivated wheat species, lutein was the major carotenoid and only traces of zeaxanthin were detected, confirming previous results obtained for durum and bread wheat. Lutein concentration was significantly different among the three wheat species, and averaged 5.75 ± 0.17, 3.32 ± 0.18, and 1.24 ± 0.17 μg g −1 DM for einkorn, durum and bread wheat, respectively. The LOX activity of the three wheat species was negatively correlated to the total carotenoid concentration, and was 3-times lower in einkorn (11.91 ± 1.56 nkat g −1 ) than in durum wheat (33.61 ± 1.59 nkat g −1 ), which was 2.5-times lower than in bread wheat (84.7 ± 7.72 nkat g −1 ). The peroxidase activity was not significantly different among the three wheat species. Einkorn appears to be a parent of interest for breeding, since it presents a high carotenoid concentration to LOX activity ratio, a selection criteria that predicts an optimal conservation of carotenoid pigments in bread. Total carotenoid concentration of the 48 bread wheat varieties was assessed spectrophotometrically after calibration against HPLC determination. The results show significant varietal variation of the total carotenoid concentration, ranging from 0.43 ± 0.01 μg g −1 DM (cv. Panifor) to 1.74 ± 0.07 μg g −1 DM (cv. Apache). Therefore, grain selection could be used to increase the nutritional value of wheat products.

... Leenhardt等^[9]发现普通小麦籽粒中的LOX活性高于硬粒小麦和一粒小麦, 分别是其2 ...

1977

4.11

0.0

... Hart和Langston ^[10]利用中国春缺体-四体系对LOX同工酶定位表明, Lpx-A1、Lpx-B1和Lpx-D1分别位于4AL、4BL和4DS染色体, 而Lpx-A2、Lpx-B2和Lpx-D2分别位于5AL、5BL和5DL, 推测小麦LOX基因主要位于第4和第5同源群 ...

1984

1.251

0.0

... Hsieh等^[11]从硬粒小麦胚乳中分离出3个与LOX活性有关的同工酶, 分别命名为lipoxygeanses-1 (L-1)、lipoxygeanses-2 (L-2)和lipoxygeanses-3 (L-3) ...

2009

2.088

0.0

. 2009, 50:67-73 DOI:10.1016/j.jcs.2009.02.008

Physical mapping of durum wheat lipoxygenase genes

Abstract A bright yellow color is an important quality criterion for pasta making. Yellow color depends on the amount of carotenoid pigments in grain, which is the result of the balance between pigment synthesis and degradation by lipoxygenases (LPX). The organization of genes coding for lipoxygenases in the tetraploid wheat genome is not completely understood. Here, we report the screening of a durum wheat BAC library with barley probes to characterize the physical distribution of Lpx genes. PCR characterization and BAC fingerprinting of the positive clones suggests that Lpx-B1.1 and Lpx-B3 are less than 103-kb apart, whereas Lpx-B1.2 is further apart from them. In the A genome a partially deleted copy of Lpx-1 ( Lpx-A1_like ) was found, colocalizing within a 42 kbp region with Lpx-A3 , confirming that in both genomes these two genes are close to each other. The knowledge of the physical location of these two genes is important to understand the evolution of this family but also has practical implications since closely linked genes are difficult to separate by recombination. This may limit the number of Lpx allele combinations that can be obtained and affect the selection of optimal Lpx allele combinations for pasta quality improvement.

2008

0.427

0.0

. 2008, 44:567-574 DOI:10.1134/S1022795408050098

Mapping of quantitative trait loci (QTL) associated with activity of disulfide reductase and lipoxygenase in grain of bread wheat

1.Russian Academy of Sciences Institute of Cytology and Genetics Novosibirsk 630090 Russia 2.Russian Academy of Sciences Institute of Plant Physiology and Biochemistry Irkutsk 664033 Russia 3.Leibniz-Institut für Pflanzengenetik und Kulturpflanzenforschung (IPK) Gatersleben D-06466 Germany

Activity of two enzymes of thiol-disulfide cell metabolism, lipoxygenase (LOX, EC 1.13.11.12) and disulfide-reductase (TPDO, EC 1.8.4.2) was studied in recombinant inbred lines of bread wheat ITMI. Their activity in the caryopsis may be connected with the gluten quality, one of the most important traits significant for breeding. The activity of lipoxygenase under favorable and droughty environmental conditions was shown to be associated with the quantitative trait locus (QTL) located on chromosome 4BS near the structural gene of a subunit of this enzyme. However, no QTL common to this enzyme and any characteristic of gluten quality have been found. Four loci responsible for the activity of disulfide reductase were identified on chromosomes 4A, 5D, 6A, and 7D. Previously, indicators of grain and flour properties, such as elasticity, flour strenght, and grain hardiness were mapped at the same loci. This indicates that the given enzyme participates in the formation of the protein complex upon maturation of wheat grain. The detected QTL can be involved in further genetic studies designed to establish the regularities of gluten formation.

... 在对小麦籽粒LOX活性相关基因的定位研究中, 发现一些与目标性状紧密连锁的SSR标记, 如Xbcd1262、Xksud2a、Xwmc312和Xgwm251^[13,29], 但这些标记不在基因内部, 不能与目的基因共分离, 因而预测基因型的准确性较低, 在实际应用中存在局限性 ...

2007

2.088

0.0

. 2007, 45:67-77 DOI:10.1016/j.jcs.2006.07.001

A deletion at the Lpx-B1 locus is associated with low lipoxygenase activity and improves pasta color in durum wheat (Triticum turgidum ssp. durum)

Abstract The concentration of yellow carotenoid pigments in durum wheat grain is an important quality criterion and is determined both by their accumulation and by their degradation by lipoxygenase enzymes ( Lpx loci). The existence of a duplication at the Lpx-B1 locus and the allelic variation for a deletion of the Lpx-B1.1 copy is reported. This deletion was associated with a 4.5-fold reduction in lipoxygenase activity and improved pasta color ( P Lpx-A3 , was mapped on the homoeologous region on chromosome 4A and was associated with semolina and pasta color ( P Lpx-A3 and the Kofa Lpx-B1.1 deletion resulted in a 10% increase in yellow scores for dry pasta relative to the opposite allele combination. This result indicates that the markers and the new allelic variants reported here will be useful tools to manipulate the wheat Lpx loci and to improve pasta color.

... Carrera等^[14]分析7个硬粒小麦品种的Lpx-B1基因及其对LOX活性的影响发现, Lpx-B1位点部分片段重叠和Lpx-B1 ...

2010

4.354

0.0

... 此外, Verlotta等^[15]还报道了一个控制LOX活性的新基因, 将其命名为Lpx-B1 ...

... 利用Verlotta等^[15]在硬粒小麦开发的共显性标记Lpx-B1 ...

... Verlotta等^[15]把Lpx-B1不同基因型组合分成3个单体型, 即单体型I (Lpx-B1 ...

... 2基因型, 与Verlotta等^[15]利用85份硬粒小麦品种的检测结果基本一致 ...

... Verlotta等^[15]筛选出低LOX活性、综合性状优异的硬粒小麦品种, 可直接用作硬粒小麦育种亲本 ...

2012

3.251

0.0

. 2012, 30:113-124 DOI:10.1007/s11032-011-9603-9

Molecular characterization of a novel type of lipoxygenase (LOX) gene from common wheat (Triticum aestivum L.)

1. Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu, 610041, China 3. ECORES Laboratory, Chinese Academy of Sciences, Chengdu, 610041, China 2. The State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing, 100101, China

Abstract Lipoxygenases (LOXs) are key enzymes catalyzing the oxidation of polyunsaturated fatty acids, and play an important role in grain storage, seed vigor and bread-making. In this study, we report cloning and characterization of TaLOX3 , the homolog gene of barley LoxB , from common wheat ( Triticum aestivum L.). The full-length cDNA sequence of TaLOX3 had an open reading frame of 2,634 bp encoding a deduced polypeptide of 878 amino acid residues with a predicted molecular weight of 99,549.79 Da and an estimated isoelectric point of 6.26. TaLOX3 was assigned to chromosome 4A. The multiple alignment result showed that TaLOX3 protein had an identity of 91.3% to barley LOX3 (encoded by LoxB ). Profile analyses revealed that the three TaLOX genes are expressed in unique patterns in various organs. Furthermore, the expression profiles of the TaLOX3 gene were determined by q-PCR using the high and low seed LOX activity of different wheat varieties. The result showed that TaLOX3 transcript levels increased significantly during the development of wheat grain. Further analysis indicated genotypes with high LOX activity were associated with higher expression levels than the low LOX activity ones. This indicates that TaLOX3 may contribute to the total LOX activity in common wheat seeds. Phylogenetic analysis conducted with LOXs from rice, maize, Brachypodium distachyon genome, barley and wheat seeds (TaLOX1, TaLOX2 and TaLOX3) revealed that LOXs could be well resolved into two clades (G1 and G2). Moreover, G1 can be divided to three subgroups, with each TaLOX in one subgroup. Collectively, these results can extent our knowledge of the molecular characterization of LOXs, and will provide a valuable resource for wheat quality improvement and breeding.

... 2 小麦籽粒LOX活性的测定取完熟后籽粒, 用分光光度计法^[16]测定籽粒中LOX活性, 每份材料3次重复, 求平均值 ...

2012

1.513

0.0

2013

2.088

0.0

2010

2.088

0.0

. 2010, 52:387-394 DOI:10.1016/j.jcs.2010.06.019

Molecular analysis of lipoxygenase (LOX) genes in common wheat and phylogenetic investigation of LOX proteins from model and crop plants

Abstract Lipoxygenase (LOX) proteins constitute an important class of lipid-hydrolyzing enzymes in cellular organisms. Here, we report a molecular analysis of LOX genes in common wheat ( Triticum aestivum L.) and a more comprehensive phylogenetic investigation of the LOX proteins from higher plants. The full-length nucleotide sequences of two LOX genes ( TaLOX1 and TaLOX2 ) from common wheat were isolated. TaLOX1 and TaLOX2 were assigned to the short arm of chromosome 4D and the long arm of chromosome 5D, respectively. TaLOX1 and TaLOX2 were both expressed in the developing grains of two common wheat varieties, indicating that they may contribute to the total LOX activity in common wheat seeds. A more extensive phylogenetic analysis conducted with 143 unique LOXs from model and crop plants revealed that plant LOXs could be classified into two subfamilies, which correlated with the absence (subfamily I) or presence (subfamily II) of a predicted chloroplast targeting peptide in the deduced LOX proteins. Our work provides new information on the LOX genes in common wheat and the phylogenetic relationships of higher plant LOXs, which may aid further functional and evolutionary studies of plant LOXs.

2006

2.906

0.0

... 硬粒小麦在储藏和加工过程中脂肪氧化酶的氧化降解会导致胡萝卜素和类胡萝卜素大量损失, 降低部分营养价值^[20,21] ...

2006

1.24

0.0

... 硬粒小麦在储藏和加工过程中脂肪氧化酶的氧化降解会导致胡萝卜素和类胡萝卜素大量损失, 降低部分营养价值^[20,21] ...

2013

0.0

1.889

. 2013, 46:1087-1094 DOI:10.3864/j.issn.0578-1752.2013.06.001

Variations of LOX and PPO activities and carotenoid content as well as their influence on whole flour color in common wheat

College of Agronomy, Anhui Agricultural University/Key Laboratory of Wheat Biology and Genetic Breeding in Southern Huang-huai Wheat Region, Ministry of Agriculture, Hefei 230036

【Objective】The aim of this study is to provide a theoretical basis for improving flour color and related traits in breeding practice. LOX and PPO activities and carotenoids concerntration of fresh and stored whole flours were determined to study their variation, analyze the effects of these traits on the whole flour color by correlation coefficient, and evaluate the influence of storage time on flour color.【Method】The LOX and PPO activities and carotenoids concerntration were determined spectrophotometrically in 63 wheat varieties. The whole flour color was evaluated by Color difference meter.【Result】There were significant or very significant variations of LOX and PPO activities and carotenoids concerntration as well as other color index in both fresh and stored whole flours. During the storage, the LOX and PPO activities reduced quickly, but the carotenoids concentration remained steadily. For fresh whole flour, the whiteness and brightness had a significant negative correlation with PPO activity, while yellowness showed a significant positive correlation with carotenoids concerntration. LOX activity was only significantly negatively correlated with carotenoid content whereas no significant relationships with the whole flour whiteness, brightness and yellowness were found. After storage for 3 months, the whole flour whiteness and yellowness were only significantly negatively correlated with PPO activity.【Conclusion】To improve the flour color, the genetic control of PPO activity and carotenoids concerntration is essential. Storing whole flour is an impracticable method as it cannot result in the increase of whole flour whiteness and brightness, and the reduction of yellowness in wide range. The genetic breeding would be an important way to improve flour whiteness or yellowness with high nutritional value.

【目的】研究新鲜全麦粉及储存3个月后全麦粉中脂肪氧化酶（LOX）和多酚氧化酶（PPO）的活性及类胡萝卜素含量的变化情况，分析其色泽性状与LOX、PPO活性和类胡萝卜素含量之间的关系，了解储存时间对全麦粉色泽的影响及其品种间的差异，为全麦粉或面粉色泽的改良和相关的育种工作提供参考依据。【方法】选用63个普通小麦品种（系）为试验材料，用分光光度计法测定其LOX、PPO活性和类胡萝卜素含量，用色差计法测定全麦粉色泽性状。【结果】品种间LOX、PPO活性、类胡萝卜素含量及其它色泽性状差异显著或极显著；新鲜全麦粉和存储3个月的全麦粉的LOX、PPO活性变化较大，储存期间LOX、PPO活性丧失较快，类胡萝卜素含量相对稳定；储存3个月后的白度和亮度等色泽性状存在一定程度的变化，但变化不大。新鲜全麦粉的白度和亮度均与PPO活性呈极显著负相关，黄度与类胡萝卜素含量呈极显著正相关，LOX活性与类胡萝卜素含量呈极显著负相关，而LOX活性与白度、亮度和黄度间相关均不显著；储存3个月的全麦粉的白度和亮度仅与PPO活性呈极显著负相关。【结论】全麦粉色泽改良中，PPO活性和类胡萝卜素的遗传控制是关键；通过储存全麦粉或面粉的方式并不能大幅度的提高其亮度、白度和降低黄度。所以，遗传育种方法是改良小麦粉白度性状和类胡萝卜素含量等品质性状的重要途径。

... 郑文寅等^[22]分析了小麦全麦粉的色泽性状与LOX和类胡萝卜素含量之间的关系, 结果表明, LOX活性与类胡萝卜素含量呈极显著负相关, 而与全麦粉的白度、亮度和黄度无显著相关 ...

... 但是过高的LOX活性则会破坏小麦籽粒中的黄色素, 使小麦粉过白而丧失许多营养成分, 降低麦类食品的营养价值^[3,6,22] ...

2005

2.088

0.0

... 0 mL离心管, 参照单籽粒种子DNA快速提取方法^[23]提取DNA, 但用SLS (十二烷基肌氨酸钠)代替SDS (十二烷基硫酸钠) ...

1999

1.251

0.0

... 研究表明, LOX与小麦籽粒、面粉及其面制品的储藏特性的关系密切, 降低LOX活性有利于延长硬粒小麦籽粒、通心粉等食品的保存期, 从而提高产品的附加值^[24], 降低LOX活性也是长期保存种子的重要方法之一 ...

2012

0.0

... 可见, 低LOX活性可以有效地减轻脂质的氧化反应, 减少籽粒的氧化变质, 延长其储藏期, 从而减少粮食的损失^[6,25] ...

2007

0.689

0.0

. 2007, 43:91-97 DOI:10.1134/S0003683807010164

The effect of intercultivar substitution of wheat Triticum aestivum L. chromosomes on lipoxygenase activity and its correlation with the technological properties of flour

1.Siberian Institute of Plant Physiology and Biochemistry, Siberian Branch of the Russian Academy of Sciences, Irkutsk, 664033 Russia 2.Institute of Cytology and Genetics, Siberian Branch of the Russian Academy of Sciences, Novosibirsk, 630090 Russia

The effects of intercultivar substitution of individual chromosome pairs (except for 1B, 6D, and 7A) in the wheat cultivars Saratovskaya 29 and Janetzkis Probat , differing in quality, on specific lipoxygenase activity, the grain yield per spike, and the main technological properties of flour and dough were studied. It was demonstrated that the substitution of individual chromosomes of the recipient cultivar Saratovskaya 29 with the homologous chromosomes of the donor cultivar Janetzkis Probat caused significant changes in lipoxygenase activity and several other quality characteristics. The correlations between the lipoxygenase activity and the parameters of physical dough properties were determined. Three molecular forms of lipoxygenase (Lpx-1, Lpx-2, and Lpx-3), differing in the value of surface charge and enzymatic activity, were detected by native PAGE.

... 研究表明, LOX活性与蛋白质、淀粉、湿面筋等大多数品质参数呈负相关, 且能有效增加面筋强度, 导致面团延展性降低^[26,27] ...

2010

0.0

... 研究表明, LOX活性与蛋白质、淀粉、湿面筋等大多数品质参数呈负相关, 且能有效增加面筋强度, 导致面团延展性降低^[26,27] ...

2010

1.24

0.0

... 与普通小麦相比, 硬粒小麦中LOX活性更低, 同时由于硬粒小麦中的蛋白质和抗氧化剂化合物也显著高于普通小麦, 因此认为硬粒小麦比普通小麦具有更高的营养价值^[28] ...

2011

3.251

0.0

. 2011, 28:117-126 DOI:10.1007/s11032-010-9466-5

Molecular markers for tracking variation in lipoxygenase activity in wheat breeding

1. College of Agronomy, Xingjiang Agricultural University, 42 Nanchang Road, 830052, Urumqi, Xinjiang, China 2. Institute of Crop Science, National Wheat Improvement Center/The National Key Facility for Crop Gene Resources and Genetic Improvement, Chinese Academy of Agricultural Sciences (CAAS), 12 Zhongguancun South Street, 100081, Beijing, China 3. International Maize and Wheat Improvement Center (CIMMYT) China Office, c/o CAAS, 12 Zhongguancun South Street, 100081, Beijing, China 4. Beijing Engineering and Technique Research Center of Hybrid Wheat, Beijing Academy of Agricultural and Forestry Sciences, 100097, Beijing, China 5. Centre for Comparative Genomics, Murdoch University, Perth, WA, 6150, Australia

Abstract Lipoxygenase (LOX) activity is an important factor determining the color of flour and end-use products of wheat. In the present study, quantitative trait loci (QTL) for LOX activity in common wheat were mapped using 71 doubled haploid (DH) lines derived from a Zhongyou 9507 × CA9632 cross, and SSR markers. Two QTL, QLpx.caas.1AL and QLpx.caas - 4B , were identified on chromosomes 1AL and 4B, closely associated with LOX activity. The SSR loci Xwmc312 and Xgwm251 proved to be diagnostic and explained 13.4–25.2% of the phenotypic variance for the 1AL locus and 14.3–27.0% for the 4B locus across four environments. The SSR markers Xgwm251 and Xwmc312 were validated across 198 Chinese wheat cultivars and advanced lines and showed highly significant ( P Xwmc312/Xgwm251 to test these wheat cultivars and advanced lines. The results suggested that the marker combination Xwmc312/Xgwm251 is efficient and reliable for evaluating LOX activity and can be used in marker-assisted selection (MAS) for targeting flour color attributes to noodle and other wheat-based products.

2007

8.455

0.0

. 2007, 10:211-216 DOI:10.1016/j.pbi.2007.01.009

Functional markers in wheat

Wheat ( Triticum aestivum L.) is one of the most important staple crops world-wide. Gene-derived ‘functional’ markers (FMs, also called perfect or diagnostic markers elsewhere) would be the ideal tools for marker-assisted breeding of wheat but, at present, their utility is restricted by the limited availability of genes that control agronomic characters. This bottleneck will be overcome in the next decade by ongoing genomics projects. Another obstacle for assigning sequence to trait variation is the high level of linkage disequilibrium (LD) found in elite materials. Thus, although laborious, TILLING seems to be the most promising approach for targeting sites in genes of interest for FM development. Once larger numbers of FMs become available for wheat breeding, they might be useful in exploiting the fixed genetic variation that is present in regions of high LD.

... 功能标记是根据基因内部引起表型性状变异的多态性序列而开发出的一种分子标记, 利用其进行辅助选择育种具有高效、准确、适用性广等特点^[30,31] ...

2012

3.658

0.0

... 目前, 功能性标记已广泛用于小麦品质和农艺性状以及抗病性鉴定等领域, 是小麦育种的重要辅助手段之一^[31] ...