大豆籽粒维生素E含量的QTL分析

引用本文

张红梅, 李海朝, 文自翔, 顾和平, 袁星星, 陈华涛, 崔晓艳, 陈新, 卢为国. 大豆籽粒维生素E含量的QTL分析. 41(2): 187-196
ZHANG Hong-Mei, LI Hai-Chao, WEN Zi-Xiang, GU He-Ping, YUAN Xing-Xing, CHEN Hua-Tao, CUI Xiao-Yan, CHEN Xin, LU Wei-Guo. Identification of QTL Associated with Vitamin E Content in Soybean Seeds. Acta Agronomica Sinica, 41(2): 187-196 复制到剪切板

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作物学报编辑部

大豆籽粒维生素E含量的QTL分析

张红梅¹, 李海朝², 文自翔², 顾和平¹, 袁星星¹, 陈华涛¹, 崔晓艳¹, 陈新^1,^*, 卢为国^2,^*

¹江苏省农业科学院蔬菜研究所, 江苏南京 210014

²河南省农业科学院经济作物研究所 / 国家大豆改良中心郑州分中心 / 农业部黄淮海油料作物遗传育种重点实验室, 河南郑州 450002

^* 通讯作者(Corresponding authors): 陈新, E-mail: cx@jaas.ac.cn, Tel: 025-84391362; 卢为国, E-mail: 123bean@163.com, Tel: 0371-65733647

第一作者联系方式: E-mail: zhm@jaas.ac.cn

收稿日期:2014-12-19

基金:本研究由国家自然科学基金项目(30771360), 江苏省农业自主创新基金项目[cx(13)4070]和江苏省科技支撑计划项目(BE2013379)资助;

摘要

维生素E (VE)具有提高人体免疫力、抗癌、预防心血管疾病等保健作用, 从大豆中提取的VE安全性更高。本研究采用高效液相色谱技术(HPLC)检测大豆BIEX群体(Essex×ZDD2315)维生素E的α-生育酚、γ-生育酚和δ-生育酚含量。应用QTLNetwork 2.1软件分别检测到8个和12对控制大豆维生素E及组分含量的加性和互作QTL。α-生育酚含量加性和互作QTL累计贡献值分别为8.68% (2个)和15.57% (4对), γ-生育酚含量加性和互作QTL累计贡献值分别为8.59% (2个)和11.57% (2对), δ-生育酚含量加性和互作QTL累计贡献值分别为5.44% (1个)和17.61% (3对), 维生素E总含量的加性和互作QTL累计贡献值分别为11.39% (3个)和9.48% (3对)。未检测到维生素E及组分含量和环境互作的QTL。未定位到的微效QTL累计贡献值为66.16%~75.32%, 说明未定位到的微效基因的变异占2/3以上。各性状的遗传构成中, 未检测出的微效QTL份额最大, 加性QTL和互作QTL贡献相差不大。在育种中应考虑常规方法聚合微效QTL与标记辅助方法聚合主要QTL相结合。

关键词: 大豆; 籽粒; 维生素E含量; QTL定位

Identification of QTL Associated with Vitamin E Content in Soybean Seeds

ZHANG Hong-Mei¹, LI Hai-Chao², WEN Zi-Xiang², GU He-Ping¹, YUAN Xing-Xing¹, CHEN Hua-Tao¹, CUI Xiao-Yan¹, CHEN Xin^1,^*, LU Wei-Guo^2,^*

¹Institute of Vegetable Crops, Jiangsu Academy of Agricultural Sciences, Nanjing 210014, China

² Institute of Industrial Crops, Henan Academy of Agricultural Scienecs / Zhengzhou Subcenter of National Center for Soybean Improvement / Key laboratory of Oil Crops in Huanghuaihai Plains, Ministry of Agriculture, Zhengzhou 450002, China

Abstract

Vitamin E has effects on human immunity, anti-cancer and prevention of cardiovascular disease. Vitamin E from soybean has the advantages of higher security and higher human body absorption rate. The objective of the present study was to map the additive, additive × additive (epistasis), additive × year and epistasis × year QTLs for vitamin E and relative tocopherol contents with the RIL population BIEX (Essex×ZDD2315) using HPLC (high performance liquid chromatography) method and software QTLNetwork 2.1. Eight additive QTLs and twelve additive × additive (epistasis) QTLs were detected for vitamin E and relative tocopherol contents. The contributions to the phenotypic variances of additive QTL and epistatic QTL pairs were 8.68% (two QTLs) and 15.57% (four pairs) for α-tocopherol, 8.59% (two QTL) and 11.57% (two pairs) for γ-tocopherol, 5.44% (one QTL) and 17.61% (three pairs) for δ-tocopherol and 11.39% (three QTL) and 9.48% (three pairs) for total vitamin E contents, respectively. Those of additive and epistatic QTLs by year interaction were not found. The accumulated contribution of the unmapped minor QTLs was 66.16%-75.32%, indicating the variance of unmapped minor QTLs accounting for more than two thirds. In genetic composition, undetected minor QTLs accounted for a considerably large part additive QTLs and epistatic QTLs were nearly equal in α-tocopherol, γ-tocopherol, δ-tocopherol and total vitamin E contents. Accordingly, in breeding for vitamin E contents, the strategy of pyramiding multiple QTLs, both additive and epistatic, by using marker-assisted selection combined with accumulating minor effect QTLs through conversional procedures should be considered.

Keyword: Soybean [ Glycine max (L.) Merr.]; Seed; Vitamin E content; QTL mapping

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维生素E(VE)是脂溶性维生素, 主要由α 、β 、γ 、δ 型生育酚及其相应的三烯生育酚8种同系物组成。维生素E参与生物体内多种代谢途径, 不仅参与信号转导、基因表达调控、抗低温胁迫, 而且还具有提高机体免疫力、抗不育、抗癌及预防心血管疾病的作用^{[1, 2, 3, 4, 5]}。在天然存在的4种生育酚中, α -生育酚的生物活性最高且被人体优先吸收和利用, α -生育酚大多存在于植物绿色组织中, 而γ -生育酚大多在种子中发现, 它是大豆油中的一种天然抗氧化剂^[6], 可用来保持油脂的风味、延长油脂的储藏时间以及保证种子长期储藏后的种子活力。人体所需维生素E 70%来源于植物油, 它是良好天然维生素E来源。近年来, 天然维生素E年需求增长10%, 而实际年产量难以满足市场需求, 生产厂家主要集中在美国、日本、德国等发达国家, 中国国内产量有限。因此, 在有限的大豆种植面积下, 选育富含高维生素E的大豆品种, 尤其是富含高α -生育酚的大豆品种是满足大豆维生素E市场需求的必要途径^{[7, 8]}。本研究解析了大豆籽粒维生素E含量的遗传控制基础, 可为选育高维生素E的大豆品种提供参考。

目前国内外研究大豆维生素E组分含量的遗传机制比较少。李卫东等^[9]的研究表明, 大豆种子中VE含量受环境影响较大, 受多个基因控制。Dwiyanti等^[10]用SSR标记分析了F₂和F₃群体中α -生育酚含量, 发现K连锁群对大豆种子α -生育酚含量有重要影响。由于维生素E检测的费用比较高, 相关的QTL定位不是很多。Li等^[11]利用大豆RIL群体在3个环境发现在Dlb连锁群上同时控制α -、γ -、δ -生育酚的QTL, 在2个环境发现在N、A2和L连锁群上同时控制α -、γ -、δ -生育酚的QTL。定位大豆籽粒维生素E含量的QTL, 发掘紧密连锁的分子标记, 是解析大豆维生素E含量的遗传基础, 利用分子标记育种选育高维生素E含量大豆新品种的重要前提。本研究利用一个黄淮海产区夏大豆重组自交系群体(RIL)进行大豆种子维生素E含量的遗传分析, 检测α -、γ -、δ -生育酚和维生素E总含量的QTL, 以期为黄淮海产区大豆籽粒高维生素E或高α -生育酚含量的大豆品种选育奠定遗传学基础。

1 材料与方法

1.1 试验材料与设计

供试材料为河南省农业科学院经济作物研究所配制的Essex× ZDD2315杂交组合。包含208个家系的重组自交家系群体BIEX (F_2:7-12), 即每一F₂单株衍生成F₇家系, 每一F₇家系再选株建成家系, 至2008年是F₁₃代。亲本Essex和ZDD2315分别来自美国南方和我国黄淮海地区。于2008年和2009年夏季在江苏省农业科学院六合基地种植试验群体及亲本, 采用随机区组设计, 3次重复, 2行区, 行长4 m。田间管理同一般大田生产。

1.2 大豆籽粒维生素E提取和检测

维生素E测定方法参照GB/T5009.82-2003, 并略作改进, 改进的关键步骤是以正己烷代替乙醚, 提高了提取效果; 提取次数增加一次。从每个家系选取10.00~15.00 g籽粒饱满大小一致的大豆籽粒, 经样品研磨(FOSS, Knifetec 1095) 60 s粉碎, 称取2.5 g粉碎后的豆粉样品, 加入30 mL无水乙醇混匀, 再加10%抗坏血酸溶液5 mL和氢氧化钾溶液10 mL恒温水浴皂化30 min; 然后, 用100 mL正己烷分3次提取皂化液, 弃去水层; 最后, 将正己烷提取液通过无水硫酸钠, 过滤到旋转蒸发瓶中旋转近干, 用氮气吹干残留液后, 用2.5 mL甲醇溶解, 过0.45 µ m滤膜, 4℃冰箱保存备用。

利用高效液相色谱技术(HPLC), 采用外标法对生育酚各异构体进行定量分析。色谱柱为DIKMA公司产品, 色谱柱填料为symmetry, 钻石C18, 5 μ m, 柱规格为250.0 mm × 4.6 mm; 荧光检测器激发波长290 nm, 发射波长330 nm; 流动相为甲醇, 流速1.5 mL min^-1; 柱温40℃; 进样量20 μ L; 检测时间10 min。以α -生育酚、γ -生育酚、δ -生育酚峰面积代入回归方程进行定量分析。由于β -生育酚在大豆籽粒中的含量很低, 忽略不计。维生素E总含量为α -、γ -、δ -生育酚值之和。

1.3 数据分析

用SAS 8.01描述统计及方差分析。遗传率h²=σ _g²/[σ _g²+(σ _ge²/n)+(σ ²/rn)], 其中σ _g²为遗传方差, σ _ge²为基因型× 年份(环境)方差, σ ²为误差方差, r为重复数, n为年份数; 遗传变异系数GCV(%)=σ _g/μ × 100。其中σ _g为遗传方差的标准差, μ 为群体平均数, 均由试验数据估计。

1.4 遗传图谱构建和QTL定位

利用Mapmaker/Exp 3.0构建连锁图谱, 24个连锁群包括198个SSR标记, 覆盖2179 cM, 平均图距为11.0 cM。经与Song等^[12]整合的图谱相比对, 标记排序相对一致。但O、D1b、N和F这4个连锁群由于大的gap分出小连锁群, 标记引物信息、标记试验过程及作图过程见参考文献[13]。

QTL定位主软件为QTLNetwork 2.1^[14]。该软件是基于混合线性模型的复合区间作图方法, 遗传模型中包括加性、显性、上位性和环境互作等各种效应。利用回归分析对基因组进行一维扫描探测加性QTL以及二维扫描探测上位性, 采用基于Henderson方法III的F检验。用基于Gibbs抽样的Bayesian方法估计遗传效应。采用Piepho和Gauch^[15]提出的标记成对选择方法(marker pair selection)选择复合区间作图中背景标记。本研究中将全基因组扫描的显著性水准设置为0.05, QTL检测的阈值以排列组合1000次结果为准^[16], 将检测到的所有QTL以及它们之间的上位性互作整合到一个全QTL模型中估计遗传效应。

为核对定位效果, 本研究同时还利用WinQTLCart 2.5的复合区间作图(CIM)^[17]和QTL IciMapping 2.0的复合区间作图(ICIM)^[18]定位软件比较QTL定位结果。前者的遗传模型中包含加性和显性效应。后者的遗传模型中包括加性、显性和上位性效应。CIM法扫描时, 采用模型6 (标准模型), 在每个被测标记区间两侧设置10 cM的窗口(Window), 同时设置该窗口之外的5个标记作为余因子, 步移速度为2 cM, 向前回归法。对每个性状分别进行1000次排列测验(permutation test), 以确定每个性状的LOD阈值(显著水准0.05)。利用QTL IciMapping 2.0软件MET模块中ICIM法进行上位性QTL定位, 常用LOD值为2.0, 由于本研究包含2个环境, 再结合定位结果, 故选取LOD值为4.0, 超过阈值就认为存在QTL。

参考Chantret等的方法^{[19, 20, 21, 22]}判断相同QTL, 并采取更为严格的条件, 即在置信区间相互重叠的基础上, 增加2个QTL位置不超过5 cM, 两侧标记至少有一个相同的条件, 这样的QTL看作同一个QTL。

2 结果与分析

2.1 大豆维生素E及组分含量的表型分析

表1列出了BIEX群体中维生素E组分的表型变异情况。α -生育酚亲本平均差为10.41 μ g g^-1; γ -生育酚的亲本平均差为80.55 μ g g^-1; δ -生育酚亲本平均差为13.98; 维生素E总含量亲本平均差为104.94 μ g g^-1。维生素E及组分含量分布都呈数量性状的连续分布特点, 遗传变异系数从γ -生育酚的15.72%到δ -生育酚的27.08%, 同一性状不同年份间遗传变异系数接近。维生素E总含量两年遗传率96.18%, 3种组分2年综合的遗传率在89.21%~ 96.67%之间。所有性状均有较大幅度超亲分离, 表明亲本间位点互补。表2列出了2年联合方差分析结果。除了α -生育酚和δ -生育酚含量间表现差异不显著外, 其他生育酚在组分间均存在极显著差异 (P< 0.01), 年份和家系× 年份的互作方差未达显著水平, 即维生素E含量的表达受环境影响的程度较低。

表1 BIEX群体大豆种子维生素E含量表型变异 Table 1 Phenotypic variation of vitamin E contents of soybean seed in BIEX population

表2 BIEX群体大豆种子维生素E含量方差分析 Table 2 Analysis of variance of vitamin E contents in two years in BIEX population

2.2 大豆维生素E及组分含量的相关性分析

如表3所示, 不同环境间α -生育酚与γ -生育酚和VE总含量之间都呈极显著相关性, 且相关系数均小于0.50, α -生育酚与δ -生育酚含量之间不显著相关, 这可能与它们在大豆籽粒中含量都比较少有关; γ -、δ -生育酚和VE总含量的相关系数均表现为γ -生育酚较高(0.98和0.83)、δ -生育酚较低(0.60和0.49), 这与它们在大豆籽粒中含量规律表现一致。

表3 大豆种子维生素E及相关生育酚含量的相关性分析 Table 3 Correlation analysis of vitamin E and correlated tocopherol contents in soybean seeds

2.3 大豆维生素E及组分含量的QTL定位总体情况

以QTLNetwork 2.1的定位结果为主, 共在6个连锁群上检测到维生素E及组分含量的8个加性QTL, 解释率2.40%~6.18%, QTL贡献值均小于10%, 这些检测到的QTL均在WinQTLCart 2.5中检测到(表4)。4种性状累计表型变异解释率依次为8.58%、7.98%、5.49%和11.49%。其中2个QTL在2年同时被WinQTL Cart 2.5和QTL IciMapping 2.0检测到(表4和图1)。

表4 BIEX群体中以 MCIM 法检测出的大豆籽粒维生素E含量的加性QTL Table 4 Additive QTL of vitamin E contents of soybean seed detected by MCIM in the BIEX population

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图1 以MCIM法检测出大豆维生素E的QTL在连锁图上的位置椭圆: α -生育酚QTL; 方框: γ -生育酚QTL; 三角: δ -生育酚QTL; 菱形: VE含量QTL; 直线: 相连的2个加性QTL间有互作。Fig. 1 Locations of QTL for vitamin E contents detected by MCIM in soybean linkage mapOval: QTL for α -Toc; Square: QTL for γ -Toc; Triangle: QTL for δ -Toc; Rhombus: QTL for total vitamin E contents. Line: joining two additive QTLs represent epistatic interactions between them.

QTLNetwork2.1检测到了12对上位性QTL, 它们所占份额较小。同时各个性状还有许多未检测出来的微效QTL, 表型变异解释率从1.34%到7.79%。4种性状累计表型变异解释率依次为16.11%、12.14%、18.24%和8.02% (表5)。

表5 BIEX群体中以MCIM法检测出的大豆籽粒维生素E含量的互作QTL对 Table 5 Epistatic QTL pairs of VE contents of soybean seed detected by MCIM in the BIEX population

3类QTL中, 各性状上位性互作QTL具有与加性QTL相近的贡献份额, 而未检测到与环境互作的QTL (表6和图2)。以下将分别对维生素E及组分含量各个性状的QTL作分析。

2.3.1 α -生育酚含量在O-1和C2连锁群上各发现1个控制α -生育酚含量的加性QTL (表4和图1), 表型变异解释率分别为2.40%和6.18%, 累计8.58%。其中, 位于O-1连锁群的QTL加性效应为-1.08, 增加α -生育酚含量的等位基因来自ZDD2315; 而位于C2连锁群的QTL加性效应为1.78, 增加α -生育酚含量的等位基因来自Essex。qα 17-1可以同时被WinQTLCart 2.5和QTL IciMapping 2.0连续2年检测到。检测到4对显著互作的上位性QTL, 累计有16.11%的贡献值, 略大于加性QTL的累计贡献值。这说明, 在α -生育酚的遗传体系中, 加性和上位性QTL起重要作用。鉴于加性和上位性QTL贡献的总和只占24.69%, 而总遗传率为96.67%, 说明遗传变异中有约2/3以上为微效基因的变异(表6和图2)。

2.3.2 γ -生育酚含量在K和D1a连锁群上发现2个控制γ -生育酚含量的加性QTL, 表型变异解释率为3.64%和4.34%, 总计解释7.98%的表型变异, 单个QTL效应都不大。2个QTL的加性效应均为负值, 增效基因均来自父本ZDD2315, qγ 20-1和qγ 24-1可以被WinQTLCart 2.5检测到。检测到了2对显著互作的上位性QTL, 累计表型变异解释率12.14%。说明在γ -生育酚的遗传体系中, 加性QTL和上位性QTL占有同样重要的地位。鉴于加性和上位性QTL贡献的总和只占20.12%, 而总遗传率为92.59%, 说明该性状遗传变异中有3/4以上为微效基因的变异(表6和图2)。

2.3.3 δ -生育酚含量发现1个控制δ -生育酚含量的加性QTL和3对上位性的QTL, 加性QTL解释5.49%的表型变异, 3对上位性QTL对表型变异解释率为6.19%、4.85%和7.20%, 累计表型变异解释率18.24%。这说明加性和上位性效应对δ -生育酚的遗传起重要作用。加性QTL效应为负值, 说明增效基因来自ZDD2315, 可以同时被WinQTLCart 2.5和QTL IciMapping 2.0连续2年检测到。鉴于加性和上位性QTL贡献总和只占23.73%, 而总遗传率为89.21%, 说明该性状遗传变异中有2/3以上为微效基因的变异(表6和图2)。

2.3.4 维生素E总含量的QTL定位结果在3个连锁群(G、K和D1a)上检测到了3个控制维生素E总含量的加性QTL, 总共解释了11.49%的表型变异, 范围从2.80%到5.63%。qTVE20-1和qTVE24-1的加性效应是负的, 增效基因来自ZDD2315; qTVE10-1加性效应是正的, 增效基因来自Essex。qTVE24-1、qTVE10-1和qTVE20-1可以同时被WinQTLCart 2.5检测到。检测到了3对显著互作的上位性QTL, 总计表型变异解释率8.02%, 略小于加性QTL的累计贡献值。这表明, 在维生素E总含量的遗传中, 加性QTL和上位性QTL占有同样重要的地位。维生素E总含量的加性和上位性QTL贡献总和占19.51%, 总遗传率为88.87%, 说明该性状遗传变异中有3/4以上为微效基因的变异(表6和图2)。

表6 QTL 及其互作对大豆种子维生素E含量表型变异的贡献 Table 6 Contributions of QTLs and their interactions to phenotypic variation for VE contents of soybean seed (%)

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	图2 维生素E含量遗传体系中各种效应方差比例Fig. 2 The percentage of different effects of vitamin E contents

2.3.5 维生素E及组分含量遗传体系的特点和性状间共享的标记以上QTL结果表明, 维生素E及组分含量遗传体系中加性QTL和上位性QTL同样重要, 并且单个加性和上位性QTL的贡献均不大, γ -生育酚和δ -生育酚含量加性增效QTL均来自亲本ZDD2315, α -生育酚和维生素E总含量均有双亲增效位点互补, 微效基因贡献均在2/3以上。

本研究检测到的共享标记有: (1)控制δ -生育酚含量的加性和上位性位点qδ 14-1和δ -Toc-e1共享标记Sat_320和Satt313; (2)控制γ -生育酚和维生素E总含量的位点qγ 20-1和qTVE20-1共享标记Satt617和Satt001, 位点qγ 24-1和qTVE24-1共享标记Sat_346和Satt436。相关分析表明, γ -生育酚和维生素E总含量不同年份的相关系数为0.83和0.98, 上述共享的位点和标记可能是两性状相关的遗传基础。

3 讨论

3.1 控制大豆维生素E及组分含量的主要QTL

大豆BIEX群体维生素E及α -、γ -、δ -生育酚含量定位的QTL中, 有4个标记在2年3种方法中均被检测到(表4), 包括标记Sat_402、Satt277、Sat_320和Satt313。如与α -生育酚含量相关的位于第6染色体上的标记Sat_402和Satt277, Li等^[23]定位到标记Sat_402与叶绿素的合成有关, Satt277被认为与大豆花发育、种子蛋白质含量、油分、荚宽、荚成熟度、种子重量有关^{[24, 25]}, 说明在大豆营养生长和种子发育各个阶段, 可以通过有效措施调控α -生育酚的合成与积累。另外, Li等^[11]检测到与α -生育酚含量相关的标记Satt376也位于第6染色体, Satt376 (GmComposite 2003_M, 97.83 cM)与Sat_402 (GmComposite 2003_M, 103.33 cM)相距5.50 cM。标记Satt376还被Han等^[26]定位到与大豆单粒重相关, 并且定位区间为Satt376~Satt277, 与本文定位到控制α -生育酚含量的QTL区间Sat_402~Satt277有重叠, 表明Sat_402~Satt277标记区间对大豆单粒重和α -生育酚合成具有重要作用。另外, 与δ -生育酚含量相关位于第19染色体上的标记Sat_320和Satt313, 前人研究证明Satt313标记与种子含油量和耐铝胁迫相关^{[27, 28]}, 而这个标记未见报道与维生素E组分相关, 这可能是由于遗传背景不同所致; Sat_320标记未见报道与δ -生育酚含量相关, 它可能是控制δ -生育酚含量的新QTL, 其作用有待进一步研究证实。标记Sat_320 (GmComposite2003_M, 32.36 cM)Satt313 (GmComposite2003_M, 34.54 cM)相距2.18 cM。经Soybase网站检索Sat_320~Satt313区间内所公布的基因, 包含锌指蛋白 ZIP 转录因子(Glyma19.G099000), 可能与维生素E合成和调控有关; 该区间同时还包含有K⁺、Cl^-、Ca²⁺调控基因(Glyma19.G098800、Glyma19.G098100和Glyma19.G 098900)以及抗逆境基因(Glyma19.G098200), 这些离子对干旱、盐渍化及其渗透胁迫的调节具有重要的作用, 说明该标记区间与维生素E组分和大豆逆境密切相关。所以, 本文发掘到的Sat_402~Satt277与Sat_320~Satt313标记区间及其标记在大豆品质育种和抗逆育种中具有重要作用。

第9和第1染色体上的标记区间Satt617~ Satt001和Sat_346~Satt436都与γ -生育酚含量和维生素E总含量相关, 均可以被WinQTLCart 2.5检测到, 其加性效应为负值, 说明这4个QTL对γ -生育酚和维生素E总含量起负向作用, 等位基因来源于低含量亲本ZDD2315。维生素E生物合成途径中5个关键酶大致可以分为两类, 即以提高维生素E含量为主要作用的酶, 包括HPPD和HPT; 以改变维生素E组成为主要作用的酶, 包括MPBQMT、TC和γ -TMT。催化γ -异构体转化成α -异构体的酶是合成途径的最后一个关键酶γ -TMT^[29]。本研究中定位到的控制γ -生育酚含量的QTL与大豆γ -TMT基因Glyma12g01690^[30]不在同一个染色体, 而与尿黑酸叶绿基转移酶HPT (GenBank登录号为DQ231061)基因^[31]位于相同染色体(Gm09: 18 579 763~18 585 148), 但与Satt617标记(Gm09: 31 816 370~31 816 414)相距较远。这2个标记区间及标记尚未见报道与维生素E含量相关, 说明本文定位到的这4个QTL (qγ 20-1、qγ 24-1、qTVE20-1和qTVE24-1)可能是控制维生素E含量的新QTL。本研究还定位到qα 3-1和qTVE10-1与前人^{[10, 11]}研究不同, 它们也可能是维生素E及组分含量的新位点, 其作用有待进一步研究证实。

3.2 大豆维生素E及组分间关系的QTL基础

本研究维生素E及组分在内的4种性状一共检测到8个加性QTL和12对上位性QTL。由相关分析可知, α -生育酚和γ -生育酚含量、维生素E总含量存在着极显著正相关(P< 0.001), γ -生育酚与VE总含量存在着极显著正相关(P< 0.001), δ -生育酚和γ -生育酚含量存在着极显著正相关(P< 0.001)。从QTL间的连锁关系可以看出这种相关的遗传基础。(1) α -生育酚和γ -生育酚含量: O-1连锁群上检测到控制α -生育酚含量的加性QTL qα 3-1与控制γ -生育酚含量的上位性QTL γ -Toc-e1, 1位于同一连锁群的不同区间, 加性效应一正一负, 这表明在O-1连锁群这2个区段可能存在同时降低α -生育酚含量和提高γ -生育酚含量的主效QTL和上位性QTL。(2) α -生育酚和维生素E总含量: 控制α -生育酚含量的加性QTL qα 17-1与控制维生素E总含量的上位性QTL TVE-e3, 1位于C2连锁群的同一区间, 加性效应相反。而位于O-1连锁群上控制α -生育酚含量的加性QTLqα 3-1与控制维生素E总含量的上位性QTL (TVE-e1, 1和TVE-e2, 1)定位区间不同, 加性效应有正有负。这表明在C2和O-1连锁群可能存在同时调控α -生育酚和维生素E总含量的主效QTL和互作QTL。(3) γ -生育酚和维生素E总含量: 控制γ -生育酚含量的加性QTL qγ 20-1与控制维生素E总含量的上位性QTL TVE-e3, 2位于K连锁群的不同区间, 均表现负向效应, 这表明在K连锁群这2个区段可能存在同时降低γ -生育酚和维生素E总含量的主效QTL和上位性QTL。(4) δ -生育酚和γ -生育酚含量: 控制δ -生育酚含量的上位性QTL (δ -Toc-e3, 2)和控制γ -生育酚含量的上位性QTL (γ -Toc-e2, 1)位于C2连锁群的不同区间, 均表现负向效应。这表明这2个区段可能存在同时降低δ -生育酚和γ -生育酚含量的互作QTL。QTL的定位结果与性状的表型相关一致性较好, 说明了性状相关的遗传基础可能来源于控制数量性状位点的QTL间的连锁。此外, δ -生育酚和维生素E总含量存在极显著的正相关(P< 0.001), 但是在加性QTL和互作QTL中没有检测到相关QTL间的连锁, 这说明可能还有许多没有检测到的微效QTL在起作用。

所检测到的8个加性QTL中, 加性QTL qα 17-1和qδ 14-1与互作QTLδ -Toc-e3, 2、TVE-e3, 1和δ -Toc-e1, 2在同一区间。此外, 其他加性QTL均未参与互作, 互作大多是非加性QTL之间的互作。这说明控制维生素E及组分含量的一些QTL是独立作用的, 不受其他QTL的影响, 而另一些QTL是和其他QTL一起存在时才有效应。

维生素E含有3种主要生育酚, 所以在遗传上维生素E总含量与其组分之间可能会有联系。Li等^[11]利用F_2:7群体合丰25× Bayfield对维生素E及组分含量进行定位, 在3个地区均检测到Dlb连锁群上同时控制维生素E及组分生育酚含量相关的QTL, 2个地区在N、A2和L连锁群上均检测到控制维生素E及组分生育酚含量相关的QTL。本研究利用Essex× ZDD2315 (F_2:7~12)群体对维生素E及组分含量进行定位, α -生育酚和γ -生育酚含量各检测到2个加性QTL, 4对和2对上位性QTL, 其中O-1连锁群上有1个加性QTL和上位性QTL (即前文提到的qα 3-1和γ -Toc-e1, 1)与α -生育酚和γ -生育酚含量相关, 二者效应相反; C2连锁群上控制α -生育酚的加性QTL (qα 17-1)与维生素E总含量的上位性QTL (TVE-e3, 1)均位于同一区间, 但二者效应相反。本研究表明, 在维生素E组分调控体系中控制各组分的基因可能不同。在育种实践中, 需要提高α -生育酚含量或维生素E总含量, 而O-1和C2连锁群上检测到的QTL分别在2个性状中的效应是相反的, 且涉及了上位性, 在实际利用中有一定难度。

3.3 大豆维生素E遗传体系对育种的启示

维生素E及组分生育酚含量表型变异和遗传变异的组成情况见表6和图2。本研究检测到4个性状的加性和上位性所占遗传变异的比例均只有1/4~1/3, 说明维生素E及其生育酚组分之间还存在大量微效位点。另外, 所有加性和上位性均未检测到显著的环境互作, 这可能是因为加性、上位性与环境互作效应较小而不易检出。大豆维生素E与年份方差未达显著水平, 这与李灵娟^[32]研究甘蓝DH群体在多环境下总生育酚的合成量不随生长环境变化而稳定表达相似。

作为一个育种者, 我们希望得到效应大的QTL用于标记辅助选择, 但本文检测到的单个加性QTL和互作QTL的贡献率均小于10%。因此, 如何转移和聚合包括微效基因在内的多基因对育种者来说将是一个挑战。

4 结论

利用QTLNetwork 2.1结合WinQTLCart 2.5和QTL IciMapping 2.0软件共检测到8个和12对控制大豆维生素E及组分生育酚含量的加性和互作QTL。微效基因的方差是遗传方差扣除加性和上位性方差后的剩余部分, 其中α -生育酚含量加性、互作QTL和未定位到QTL的微效基因累计贡献值分别为8.68% (2个)、15.57% (4对)和72.42%, 未检测到的微效基因变异约占遗传变异的3/4。γ -生育酚、δ -生育酚和维生素E总含量的加性QTL累计贡献值分别为8.59% (2)、5.44% (1)和11.39% (3); 互作QTL累计贡献值分别为11.57% (2)、17.61% (3)和9.48% (3); 未定位到的QTL累计贡献值72.43%、66.16%和75.32%, 说明未定位到的微效基因的变异约占2/3以上。各性状的遗传构成中, 未检测出的微效QTL份额最大, 加性QTL和互作QTL贡献相差不大。对这种遗传组成性状的育种要考虑在常规方法聚合微效QTL基础上, 结合标记辅助方法聚合主要QTL。

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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[23]	李广军, 李河南, 程利国, 章元明. 大豆叶绿素含量动态表达的QTL分析. 作物学报, 2010, 36: 242-248 Li G J, Li H N, Cheng L G, Zhang Y M. QTL analysis for dynamic expression of chlorophyll content in soybean. Acta Agron Sin, 2010, 36: 242-248 [本文引用:1] [CJCR: 1.667]
[24]	Rossi M E, Orf J H, Liu L J, Dong Z, Rajcan I. Genetic basis of soybean adaptation to North American vs. Asian mega- environments in two independent populations from Canadian × Chinese crosses. Theor Appl Genet, 2013, 126: 1809-1823 [本文引用:1] [JCR: 3.658]
[25]	Palomeque L, Li J L, Li W, Hedges B, Cober E R, Rajcan I. QTL in mega-environments: II. Agronomic trait QTL co-localized with seed yield QTL detected in a population derived from a cross of high-yielding adapted x high-yielding exotic soybean lines. Theor Appl Genet, 2009, 119: 429-436 [本文引用:1] [JCR: 3.658]
[26]	Han Y, Li D, Zhu D, Li H, Li X, Teng W, Li W. QTL analysis of soybean seed weight across multi-genetic backgrounds and environments. Theor Appl Genet, 2012, 125: 671-683 [本文引用:1] [JCR: 3.658]
[27]	Fasoula, Vasilia A, Harris, Donna K, Boerma H R. Validation and designation of quantitative trait loci for seed protein, seed oil, and seed weight from two soybean populations. Crop Sci, 2004, 44: 1218-1225 [本文引用:1] [JCR: 1.513]
[28]	Korir P, Qi B, Wang Y, Zhao T, Yu D, Chen S, Gai J. A study on relative importance of additive, epistasis and unmapped QTL for aluminum tolerance at seedling stage in soybean. Plant Breed, 2011, 130: 551-562 [本文引用:1] [JCR: 1.175]
[29]	Della P D. Progress in the dissection and manipulation of vitamin E synthesis. Trends Plant Sci, 2005, 10: 574-579 [本文引用:1] [JCR: 11.808]
[30]	Shintani D, Della P D. Elevating the vitamin E content of plants through metabolic engineering. Science, 1998, 282: 2098-2100 [本文引用:1]
[31]	Venkatesh T V, Karunanand aa B, Free D L, Rottnek J M, Baszis S R, Valentin H E. Identification and characterization of an Arabidopsis homogentisate phytyltransferase paralog. Planta, 2006, 223: 1134-1144 [本文引用:1] [JCR: 3.347]
[32]	李灵娟. 甘蓝型油菜基因组中控制种子维生素E含量QTL的检测和分析. 华中农业大学硕士学位论文, 湖北武汉, 2008 Li L J. Identification and Analysis of QTL Controlling Seed Tocopherol Content in the Genome of Brassica napus L. MS Thesis of Huazhong Agricultural University, Hubei, China, 2008 (in Chinese with English abstract) [本文引用:1]

2010

2.428

0.0

... 维生素E参与生物体内多种代谢途径, 不仅参与信号转导、基因表达调控、抗低温胁迫, 而且还具有提高机体免疫力、抗不育、抗癌及预防心血管疾病的作用^[1,2,3,4,5] ...

2008

3.582

0.0

2007

6.555

0.0

1999

1.738

0.0

2000

0.0

Bramley P

, Elmadfa

, Kafatos

, Kelly F

, Manios

, Roxborough H

, Schuch

, Sheehy

P J A

, Wagner

K H

, 2000: Vitamin E. J Sci Food Agric, 2000, 80: 913-938

1 School of Biological Sciences, Royal Holloway, University of London, Egham TW20 0EX, UK 2 Institute of Nutritional Sciences, University of Vienna, A-1090 Vienna, Austria 3 Preventive Medicine and Nutrition Clinic, School of Medicine, University of Crete, PO Box 1393, Iraklion-Crete, Greece 4 Cardiovascular Research, St Thomas' Hospital, London SE1 7EH, UK 5 Astra-Zeneca Wheat Improvement Centre, Norwich Research Park, Norwich NR4 7UA, UK 6 Department of Food Science & Technology, University College, Cork, Ireland * Department of Food Science & Technology, University College, Cork, Ireland

> There is a growing body of evidence to suggest that the dietary intakes of vitamin E are insufficient to protect against the long-term health risks associated with oxidative stress. Traditional plant breeding and food processing technologies have not concerned themselves with maximising the levels of the tocopherols in the diet, and supplementation is necessary both for nutritive reasons and for the protection of fat-rich foods against oxidative rancidity. The paper reviews the potential for improving the tocopherol levels in the diet, particularly α-tocopherol. Genetic technologies have already demonstrated the potential to enhance tocopherol levels by up-regulation of the final steps in the biosynthetic pathway. Other strategies for the enhancement of the vitamin E content of plant foods are considered both from the perspective of improved bioavailability and the levels in processed foods. Finally some priorities for future research in the field are described.

2005

6.555

0.0

... -生育酚大多在种子中发现, 它是大豆油中的一种天然抗氧化剂^[6], 可用来保持油脂的风味、延长油脂的储藏时间以及保证种子长期储藏后的种子活力 ...

2002

0.0

0.3009

米歇尔·琼斯, 田颖. 天然维生素E. 日用化学品科学, 2002, (4): 47-48

Michelle

, Tian

Naturals vitamin E.

Deterg Cosmet, 2002, (4): 47-48 (in Chinese with English abstract)

主要探讨了天然维生素E与人工合成维生素E的区别,及天然维生素E应用在化妆品中的作用和优点.

... -生育酚的大豆品种是满足大豆维生素E市场需求的必要途径^[7,8] ...

2005

1.04

0.0

... -生育酚的大豆品种是满足大豆维生素E市场需求的必要途径^[7,8] ...

2007

0.0

1.667

李卫东, 卢为国, 梁慧珍, 王树峰, 苑保军, 耿臻, 王素阁, 范彦英, 刘亚非, 王令涛. 大豆籽粒维生素E含量与生态因子关系的研究. 作物学报, 2007, 33: 1094-1099

Li W

, Lu W

, Liang H

, Wang S

, Yuan B

, Geng

, Wang S

, Fan Y

, Liu Y

, Wang L

Effects of eco-physiological factors on vitamin E content in soybean seed.

Acta Agron Sin, 2007, 33: 1094-1099 (in Chinese with English abstract)

Vitamin E (VE) is an important nutrient factor for human beings, and soybean seed is one of major available VE recourses. To reveal the correlation between VE content in soybean seed and ecological factors, the present experiment was carried out using soybean cultivar Yudou 25 sown at 5 locations of Henan province on 13 different dates in 2001 and 2002. The data of VE contents for the 109 samples of soybean seed and 33 eco-physiological factors including meteorological factors, soil nutrition and altitudes were obtained and used for statistical analyses. Step-wise regression was used to screen the correlated factors, which showed significant effect on VE content. Results indicated that the VE contents was substantially affected by the environmental conditions and ranged from 500.99 μg g -1 to 2353.68 μg g -1 among all samples. Nine eco-physiological factors were highly correlated with VE content, including 6 meteorological factors, 2 soil nutrition factors and altitude. VE content was negatively correlated with rainfall at seedling (P

1 Institute of Industrial Crops, Henan Academy of Agricultural Sciences, Zhengzhou 450002, Henan; 2 Zhoukou Agricultural Institute, Zhoukou 466001, Henan; 3 Puyang Agricultural Institute, Puyang 457000, Henan; 4 Jiaozuo Agricultural Institute, Jiaozuo 454151, Henan; 5 Seed Administration station of Henan Province, Zhengzhou 450002, Henan, China

维生素E(VE)是人体必须的营养因子。大豆籽粒富含VE，是人类重要的VE来源之一。为了研究生态因子对大豆籽粒中VE含量的影响，2001—2002年在河南省夏大豆主产区的5个试点，以豫豆25为材料分13期播种，将109个大豆样本籽粒的VE含量与气象、土壤养分和海拔等33个生态因子进行了统计分析。诸样本VE总量的变异范围为500.99~2353.68 µg g -1 。逐步回归筛选出9个生态因子与大豆VE含量显著相关。与VE含量呈负相关的因子有幼苗期(P

... 李卫东等^[9]的研究表明, 大豆种子中VE含量受环境影响较大, 受多个基因控制 ...

2007

1.04

0.0

... Dwiyanti等^[10]用SSR标记分析了F₂和F₃群体中α ...

... 3-1和qTVE10-1与前人^[10,11]研究不同, 它们也可能是维生素E及组分含量的新位点, 其作用有待进一步研究证实 ...

2010

3.658

0.0

Li H

, Liu H

, Han Y

, Wu X

, Teng W

, Liu G

, Li W

Identification of QTL underlying vitamin E contents in soybean seed among multiple environments.

Theor Appl Genet, 2010, 120: 1405-1413

Vitamin E (VE) in soybean seed has value for foods, medicines, cosmetics, and animal husbandry. Selection for higher VE contents in seeds along with agronomic traits was an important goal for many soybean breeders. In order to map the loci controlling the VE content, F 5 -derived F 6 recombinant inbred lines (RILs) were advanced through single-seed-descent (SSD) to generate a population including 144 RILs. The population was derived from a cross between ‘OAC Bayfield’, a soybean cultivar with high VE content, and ‘Hefeng 25’, a soybean cultivar with low VE content. A total of 107 polymorphic simple sequence repeat markers were used to construct a genetic linkage map. Seed VE contents were analyzed by high performance liquid chromatography for multiple years and locations (Harbin in 2007 and 2008, Hulan in 2008 and Suihua in 2008). Four QTL associated with α-Toc (on four linkage groups, LGs), eight QTL associated with γ-Toc (on eight LGs), four QTL associated with δ-Toc (on four LGs) and five QTL associated with total VE (on four LGs) were identified. A major QTL was detected by marker Satt376 on linkage group C2 and associated with α-Toc (0.0012 > P > 0.0001, 5.0% R 2 −1 ), total VE ( P R 2 −1 ). A second QTL detected by marker Satt286 on LG C2 was associated with γ-Toc (0.0003 > P > 0.0001, 6.0% R 2 −1 ) and total VE ( P R 2 −1 ). Another major QTL was detected by marker Satt266 on LG D1b that was associated with α-Toc (0.0002 > P > 0.0001, 4.0% R 2 −1 ) and γ-Toc (0.0032 > P > 0.0001, 3.0% R 2 −1 ). Since beneficial alleles were all from ‘OAC Bayfield’, it was concluded that these three QTL would have great potential value for marker assisted selection for high VE content.

1.Key Laboratory of Forest Tree Genetic Improvement and Biotechnology, Ministry of Education, Northeast Forestry University, Harbin, 150040 China 2.Soybean Research Institute, Key Laboratory of Soybean Biology in Chinese Ministry of Education, Northeast Agricultural University, Harbin, 150030 China

... Li等^[11]利用大豆RIL群体在3个环境发现在Dlb连锁群上同时控制α ...

... 另外, Li等^[11]检测到与α ...

... 3-1和qTVE10-1与前人^[10,11]研究不同, 它们也可能是维生素E及组分含量的新位点, 其作用有待进一步研究证实 ...

... Li等^[11]利用F_2:7群体合丰25× ...

2004

3.658

0.0

Song Q

, Marek L

, Shoemaker R

, Lark K

, Concibido V

, Delannay

, Specht J

, Cregan P

A new integrated genetic linkage map of the soybean.

Theor Appl Genet, 2004, 109: 122-128

A total of 391 simple sequence repeat (SSR) markers designed from genomic DNA libraries, 24 derived from existing GenBank genes or ESTs, and five derived from bacterial artificial chromosome (BAC) end sequences were developed. In contrast to SSRs derived from EST sequences, those derived from genomic libraries were a superior source of polymorphic markers, given that the mean number of tandem repeats in the former was significantly less than that of the latter ( P Minsoy × Noir 1 , Minsoy × Archer , Archer × Noir 1 , Clark × Harosoy , and A81-356022 × PI468916. The JoinMap software package was used to combine the five maps into an integrated genetic map spanning 2,523.6 cM of Kosambi map distance across 20 linkage groups that contained 1,849 markers, including 1,015 SSRs, 709 RFLPs, 73 RAPDs, 24 classical traits, six AFLPs, ten isozymes, and 12 others. The number of new SSR markers added to each linkage group ranged from 12 to 29. In the integrated map, the ratio of SSR marker number to linkage group map distance did not differ among 18 of the 20 linkage groups; however, the SSRs were not uniformly spaced over a linkage group, clusters of SSRs with very limited recombination were frequently present. These clusters of SSRs may be indicative of gene-rich regions of soybean, as has been suggested by a number of recent studies, indicating the significant association of genes and SSRs. Development of SSR markers from map-referenced BAC clones was a very effective means of targeting markers to marker-scarce positions in the genome.

1.USDA-ARS Soybean Genomics and Improvement Laboratory Beltsville MD 20705 USA 2.Iowa State University Department of Agronomy, USDA-ARS-CICG Ames IA 50011 USA 3.Iowa State University Department of Agronomy Ames IA 50011 USA 4.University of Utah Department of Biology Salt Lake City UT 84112 USA 5.Monsanto Company 800 Lindbergh Boulevard St. Louis MO 63167 USA 6.University of Nebraska Department of Agronomy Lincoln NE 68583-0915 USA

... 经与Song等^[12]整合的图谱相比对, 标记排序相对一致 ...

2009

0.0

... 但O、D1b、N和F这4个连锁群由于大的gap分出小连锁群, 标记引物信息、标记试验过程及作图过程见参考文献[13] ...

2007

5.323

0.0

... 1^[14] ...

2001

4.389

0.0

... 采用Piepho和Gauch^[15]提出的标记成对选择方法(marker pair selection)选择复合区间作图中背景标记 ...

1994

0.0

... 5的复合区间作图(CIM)^[17]和QTL IciMapping 2 ...

2007

4.389

0.0

... 0的复合区间作图(ICIM)^[18]定位软件比较QTL定位结果 ...

2008

0.0

1.667

邢光南, 周斌, 赵团结, 喻德跃, 邢邯, 陈受宜, 盖钧镒. 大豆抗筛豆龟蝽Megacota cribraria (Fabricius)的QTL分析. 作物学报, 2008, 34: 361-368

Xing G

, Zhou

, Zhao T

, Yu D

, Xing

. Chen

S Y

, Gai J

Mapping QTLs of resistance to Megacota cribraria (Fabricius) in soybean.

Acta Agron Sin, 2008, 34: 361-368 (in Chinese with English abstract)

Globular stink bug [Megacota cribraria (Fabricius)] is one of the important pests for soybean [Glycine max (L.) Merr.] in the central and southern China. However, very few reports on soybean resistant to M. cribraria were found in the literature. The objectives of the present study were to find QTLs associated with resistance to M. cribraria in soybean and to analyze the stability of mapped QTLs among years by using two RIL populations, NJRIKY (or KY) and NJRIWT (or WT) derived from the crosses of Kefeng 1 × Nannong 1138-2 and Wan 82-178 × Tongshanbopihuangdoujia, respectively. The 184 lines of KY and 142 lines of WT were tested and the percentage of black mildew on stem and purple spots on leaves was used as indicator to evaluate their resistance to M. cribraria under natural infestation during 2004–2006. There existed significant differences among lines in the two RIL populations during the three years, with heritability of 66.5%–88.8% in KY and 61.3%–84.8% in WT. The composite interval mapping (CIM) of the software Windows QTL Cartographer Version 2.5 was used to map QTLs. The linkage group D1a and C2 in KY and the linkage group H and D1b in WT were found to be related with resistance to L. indicata. The QTL qRMC-d1a-1 on Linkage group D1a was detected consistently associated with reaction to M. cribraria in KY during the three years, which accounted for 7.6%–31.4% of phenotypic variation; the QTL qRMC-c2-1 on linkage group C2 was detected in 2005–2006, which accounted for less phenotypic variation than the former one; the resistance alleles were from Nannong 1138-2. There appeared significant interactions between qRMC-d1a-1 and qRMC-h-1 in KY in 2005 and 2006. The QTL qRMC-h-1 on linkage group H was detected also consistently associated with reaction to M. cribraria in WT during the three years, which accounted for 16.3%–36.2% of phenotypic variation; the QTL qRMC-d1b-2 on linkage group D1b was detected in 2004-2005, which accounted for less phenotypic variation than the former one; the resistant alleles were from Tongshanbopihuangdoujia. Therefore, different QTLs conferred resistance to M. cribraria in KY and WT basically although the two QTLs in the latter population were also identified in one year in the former population. The fact that the QTLs were repeatedly detected in different environments indicated the resistance was controlled stably by the main QTLs. Based on the results, it is inferred that the markers linked to the detected QTLs should be useful for marker-assisted selection for resistance to M. cribraria in soybean.

1 Soybean Research Institute of Nanjing Agricultural University/ National Center for Soybean Improvement/ National Key Laboratory for Crop Genetics and Germplasm Enhancement, Nanjing 210095, Jiangsu, China; 2 Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China

筛豆龟蝽是我国南方大豆的主要害虫之一，本研究旨在定位筛豆龟蝽抗性QTL，分析其稳定性，为大豆抗筛豆龟蝽育种提供参考。以科丰1号×南农1138-2组合衍生的184个重组自交系群体NJRIKY（简称KY）和皖82-178×通山薄皮黄豆甲组合衍生的142个重组自交系群体NJRIWT（简称WT）为材料，2004—2006在田间自然虫源下鉴定了筛豆龟蝽抗性。不同年份内以黑霉程度为指标的方差分析结果表明家系间差异在每年都达极显著水平，遗传变异系数都相当大，遗传率中等偏高。利用Windows QTL Cartographer Version 2.5的复合区间作图法(CIM)，KY群体的抗性QTL主要位于D1a和C2连锁群，WT群体的抗性QTL主要位于H和D1b连锁群。KY群体3年均检测出的qRMC-d1a-1位于D1a连锁群，贡献率为7.6%~31.4%；2005和2006两年均检测出的qRMC-c2-1位于C2连锁群，与环境有互作，效应相对较小；抗性等位基因来自南农1138-2；qRMC-d1a-1和qRMC-h-1在2005年和2006年存在显著的互作。WT群体连锁群H上的qRMC-h-1在3年中都被检测到，贡献率为16.3%~36.2%；D1b连锁群上的qRMC-d1b-2在2004和2005年被检测到，效应相对较小；抗性等位基因来自通山薄皮黄豆甲。虽然WT群体D1b和H连锁群上的这2个QTL在KY群体中也有一年被检测到，但2个群体抗性位点基本上是不同的。QTL在不同环境被重复检出，说明大豆对筛豆龟蝽的抗性由稳定的主效QTL所控制，其2侧邻近标记有希望用于标记辅助选择育种。

... 参考Chantret等的方法^{[19,20,21,22]}判断相同QTL, 并采取更为严格的条件, 即在置信区间相互重叠的基础上, 增加2个QTL位置不超过5 cM, 两侧标记至少有一个相同的条件, 这样的QTL看作同一个QTL ...

2001

3.658

0.0

Chantret

, Mingeot

, Sourdille

, Bernard

, Jacquemin J

, Doussinault

A major QTL for powdery mildew resistance is stable over time and at two development stages in winter wheat.

Theor Appl Genet, 2001, 103: 962-971

Despite the large impact of powdery mildew in wheat cultivated areas, little has been done to study powdery mildew resistance by QTL analysis up to now. The objective of the present paper is to present how the genetic basis of powdery mildew resistance in the resistant wheat line RE714 have been studied by QTL analysis at the adult plant stage over the course of 3 years, and at the vernalized seedling plant stage, and a comparison between the results obtained. Two segregating populations (DH and F 2:3 ) were derived from the cross between the resistant line (RE714), and a susceptible line (Hardi); these were analysed for powdery mildew resistance at the adult plant stage in the field under natural infection conditions in 1996, 1997 and 1998. The DH population was also tested for powdery mildew resistance at the vernalized seedling stage with four different isolates of powdery mildew. At the adult plant stage, a total of three QTLs (on chromosomes 5D, 4A and 6A) and five QTLs (on chromosomes 5D, 6A, 7A and 7B) were found for the DH and F 2:3 populations, respectively. The genetic control of resistance was found to be polygenic but involved a major QTL (on chromosome 5D), which was detected each year and which explained a high proportion of the variability observed (28.1%–37.9%). At the vernalized seedling stage, two QTLs were found (on chromosomes 5D and 7B) and the QTL detected on chromosome 5D was common to the four isolates tested. The comparison between the two development stages showed that the QTL on chromosome 5D was detected in all the different environments tested and again explained a high proportion of the variability. Different molecular interpretations of this QTL have also been discussed.

1.I.N.R.A. Station d’Amélioration des Plantes, Domaine de la Motte, 35653 Le Rheu Cedex, France e-mail: chantret@rennes.inra.fr Fax: (33)-2-23-48-51-20 FR 2.Centre de Recherches agronomiques – CRA, Départmente de Biotechnologie, 234, Chaussée de Charleroi, 5030 Gembloux, Belgium BE 3.I.N.R.A. Station d’Amélioration des Plantes, Domaine de Crouelle, 234 Avenue du Brézet, 63039 Clermont-Ferrand Cedex 2, France FR

2005

4.389

0.0

2007

1.643

0.0

Tucker D

, Griffey C

, Liu

, Brown-Guedira

, Marshall D

, Saghai Maroof

M A

. Confirmation of three quantitative trait loci conferring adult plant resistance to powdery mildew in two winter wheat populations. Euphytica, 2007, 155: 1-13

Hypersensitive, race specific genes primarily have been deployed to control powdery mildew ( Blumeria graminis (DC) EO Speer f. sp. tritici ) in wheat ( Triticum aestivum L.); however, recent efforts have shifted to breeding for more durable resistance. Previously, three quantitative trait loci (QTL) for adult plant resistance (APR) to powdery mildew in the winter wheat cultivar Massey were identified in a Becker/Massey (BM) F 2:3 population. Fourteen new simple sequence repeat (SSR) markers were added to the pre-existing BM F 2:3 linkage maps near the QTL for APR on chromosomes 1BL ( QPm.vt-1BL ), 2AL ( QPm.vt-2AL ), and 2BL ( QPm.vt-2BL ). Genetic linkage maps comprised of 17 previously and newly mapped SSRs from the BM population on chromosomes 1BL, 2AL, and 2BL were constructed in a USG 3209/Jaypee (UJ) F 6:7 recombinant inbred line (RIL) confirmation population, wherein the APR resistance of USG 3209 was derived from Massey. Interval mapping analysis of mildew severity data collected in 2002 ( F 5:6 ) and 2003 ( F 6:7 ) field experiments with marker genotypic data obtained in 2003 ( F 6:7 ) confirmed the presence of the three QTL governing APR to powdery mildew in the UJ RILs. The QTL QPm.vt-1BL , QPm.vt-2AL , and QPm.vt-2BL explained 12–13, 59–69, and 22–48% of the phenotypic variance for powdery mildew severity in the UJ confirmation populations, respectively, in two field experiments. The current study verified that the elite wheat cultivar USG 3209 possesses the same QTL for APR as its parent Massey.

1.Department of Crop and Soil Environmental Science, Virginia Polytechnic Institute and State University, Smyth Hall (0404), Blacksburg, VA 24061, USA 2.Department of Agronomy and Plant Genetics, University of Minnesota, St. Paul, MN 55108, USA 3.USDA-ARS Plant Science Research Unit, Raleigh, NC 27695, USA

2010

0.0

1.667

李广军, 李河南, 程利国, 章元明. 大豆叶绿素含量动态表达的QTL分析. 作物学报, 2010, 36: 242-248

Li G

, Li H

, Cheng L

, Zhang Y

QTL analysis for dynamic expression of chlorophyll content in soybean.

Acta Agron Sin, 2010, 36: 242-248

Abstract The objective of this study was to map quantitative trait loci (QTLs) for chlorophyll content in the F 2:3 and F 2:4 populations derived from soybean ( Glycine max L. Merr.) varieties Lishuizhongzihuangdou and Nannong 493-1. Chlorophyll contents for the function leaf in vivo were measured from seedling to blooming stage in the 244 F 2:3 and F 2:4 families growing in 2 environments. The developmental behaviors of chlorophyll content across environments were analyzed using composite interval mapping in the software of Windows QTL Cartographer v2.5. A total of 20 QTLs were detected at various stages in 2 environments, of which only several QTLs were common across stages, years, or test sites. QTLs distributing in linkage groups N, D1a, F, and K were repeatedly detected for 3 to 4 times. QTL qchl-D1a-1 , which was located on linkage group D1a between markers sat_160 and satt147, was only identified at Jiangpu site in both years. Three common QTLs were mapped on linkage groups K, M, and N at both Jiangpu and Linyi sites. These results provide a theoretical basis for genetic analysis of chlorophyll traits and marker-assisted breeding in soybean. 摘要采用溧水中子黄豆×南农493-1后代衍生的244个F 2 单株及筛选的150个SSR分子标记构建的连锁遗传图谱, 在苗期至开花期测定F 2 衍生F 2:3 和F 2:4 家系生长正常单株的倒3复叶功能叶(非离体)的叶绿素含量13次, 通过Windows QTL Cartographer v2.5软件包的复合区间法, 动态定位了大豆叶绿素含量的QTL。结果表明, 不同时间点共检测到20个QTL, 其中, 不同发育阶段间、年份间和地点间共同的QTL较少, 不同时间点上的QTL差异较大, 重复出现在N、D1a、F和K连锁群的QTL有3∼4次。这些结果为叶绿素含量的遗传剖析和标记辅助育种提供理论依据。

... -生育酚含量相关的位于第6染色体上的标记Sat_402和Satt277, Li等^[23]定位到标记Sat_402与叶绿素的合成有关, Satt277被认为与大豆花发育、种子蛋白质含量、油分、荚宽、荚成熟度、种子重量有关^[24,25], 说明在大豆营养生长和种子发育各个阶段, 可以通过有效措施调控α ...

2013

3.658

0.0

Rossi M

, Orf J

, Liu L

, Dong

, Rajcan

Genetic basis of soybean adaptation to North American vs. Asian mega- environments in two independent populations from Canadian × Chinese crosses.

Theor Appl Genet, 2013, 126: 1809-1823

Abstract One of the goals of plant breeding is to increase yield with improved quality characters. Plant introductions (PI) are a rich source of favorable alleles that could improve different characters in modern soybean [ Glycine max (L.) Merril] including yield. The objectives of this study were to identify yield QTL underlying the genetic basis for differential adaptation of soybeans to the Canadian, United States or Chinese mega-environments (ME) and to evaluate the relationship and colocalization between yield and agronomic traits QTL. Two crosses between high-yielding Canadian cultivars and elite Chinese cultivars, OAC Millennium × Heinong 38 and Pioneer 9071 × #8902, were used to develop two recombinant inbred line (RIL) populations. Both populations were evaluated at different locations in Ontario, Canada; Minnesota, United States (US), Heilongjiang and Jilin, China, in 2009 and 2010. Significant variation for yield was observed among the RILs of both populations across the three hypothetical ME. Two yield QTL (linked to the interval Satt364–Satt591 and Satt277) and one yield QTL (linked to marker Sat_341) were identified by single-factor ANOVA and interval mapping across all ME in populations 1 and 2, respectively. The most frequent top ten high-yielding lines across all ME carried most of the high-yielding alleles of the QTL that were identified in two and three ME. Both parents contributed favorable alleles, which suggests that not only the adapted parent but also the PI parents are potential sources of beneficial alleles in reciprocal environments. Other QTL were detected also at two and one ME. Most of the yield QTL were co-localized with a QTL associated with an agronomic trait in one, two, or three ME in just one or in both populations. Results suggested that most of the variation observed in seed yield can be explained by the variation of different agronomic traits such a maturity, lodging and height. Novel alleles coming from PI can favorably contribute, directly or indirectly, to seed yield and the utilization of QTL detected across one, two or three ME would facilitate the new allele introgression into breeding populations in both North America and China.

1. Department of Plant Agriculture, Crop Science Building, University of Guelph, 50 Stone Road East, Guelph, ON, N1G 2W1, Canada 2. Department of Agronomy and Plant Genetics, University of Minnesota, 411 Borlaug Hall, 1991 Buford Circle, St. Paul, MN, 55108, USA 3. Soybean Research Institute of Heilongjiang, Harbin, Heilongjiang, People’s Republic of China 4. Jilin Academy of Agricultural Sciences, Changchun, Jilin, People’s Republic of China 5. Soybean Research Institute of the Northeast Agricultural University, Harbin, Heilongjiang, People’s Republic of China

2009

3.658

0.0

Palomeque

, Li J

, Li

, Hedges

, Cober E

, Rajcan

QTL in mega-environments: II. Agronomic trait QTL co-localized with seed yield QTL detected in a population derived from a cross of high-yielding adapted x high-yielding exotic soybean lines.

Theor Appl Genet, 2009, 119: 429-436

Seed yield mega-environment-universal and specific QTL (QTL U and QTL SP , respectively) linked to Satt100, Satt130, Satt162, Satt194, Satt259 Satt277 and Sat_126, have been identified in a population derived from a cross between a Chinese and a Canadian soybean [ Glycine max (L.) Merrill] elite line. The variation observed in yield could be the consequence of the variation of agronomic traits. Yield-component traits have been reported in the literature, but a better understanding of their impact at the molecular level is still lacking. Therefore, the objectives of this study were to identify traits correlated with yield and to determine if the yield QTL U and QTL SP were co-localized with QTL U and QTL SP associated with an agronomic trait. A recombinant inbred line (RIL) population was developed from a cross between a high-yielding adapted Canadian and a high-yielding exotic Chinese soybean elite line. The RIL were evaluated in multiple environments in China and Canada during the period from 2004 to 2006. Four yield QTL U, tagged by markers Satt100, Satt277, Satt162 and Sat_126, were co-localized with a QTL associated with an agronomic trait, behaving as either QTL U or QTL SP for the agronomic trait. For example, the yield QTL U, tagged by marker Satt100 was associated also with 100 seed weight, pods per plant, pods per node, plant height, R1, R5, R8, oil content and protein content in all Canadian environments, but only with pods per plant, pods per node, plant height, R1, R5, R8 and oil content in two or more Chinese environments. No agronomic traits QTL were co-localized with the yield QTL U tagged by the marker Satt139 or the yield QTL SP tagged by Satt259, suggesting a physiological basis of the yield in these QTL. The results suggest that a successful introgression of crop productivity alleles from plant introductions into an adapted germplasm could be facilitated by the use of both the QTL U and QTL SP because each type of QTL contributed either directly or indirectly through yield-component traits to seed yield of RILs.

1.Department of Plant Agriculture, Crop Science Building, University of Guelph, 50 Stone Road East, Guelph, ON N1G 2W1, Canada 2.Soybean Research Institute of Heilongjiang, Harbin, Heilongjiang People’s Republic of China 3.Soybean Research Institute of the Northeast Agricultural University, Harbin, Heilongiang People’s Republic of China 4.Pioneer Hi-bred Ltd, A DuPont Company, Chatham, ON Canada 5.Agriculture and Agrifood Canada, Eastern Cereal and Oilseed Crop Research Centre, Ottawa, ON Canada

2012

3.658

0.0

Han

, Li

, Zhu

, Li

, Teng

, Li

QTL analysis of soybean seed weight across multi-genetic backgrounds and environments.

Theor Appl Genet, 2012, 125: 671-683

1. Key Laboratory of Soybean Biology in Chinese Ministry of Education, (Key Laboratory of Biology and Genetics & Breeding for Soybean in Northeast China, Ministry of Agriculture), Northeast Agricultural University, Harbin, 150030, China 2. Soybean Research Institute (Key Laboratory of Soybean Biology in Chinese Ministry of Education), Northeast Agricultural University, Harbin, 150030, China

Seed weight, measured as mass per seed, is an important yield component of soybean and is generally positively correlated with seed yield (Burton et al, Crop Sci 27:1093, 1987 ). In previous reports, quantitative trait loci (QTL) associated with seed weight, were identified in single genetic background. The objective of the present study was to identify QTL and epistatic QTL underlying soybean seed weight in three RIL populations (with one common male parent ‘Hefeng25’) and across three different environments. Overall, 18, 11, and 17 seed weight QTL were identified in HC (‘Hefeng25’ × ‘Conrad’), HM (‘Hefeng25’ × ‘Maple Arrow’), and HB (‘Hefeng25’ × ‘Bayfield’) populations, respectively. The amount of phenotypic variation explained by a single QTL underlying seed weight was usually less than 10 %. The environment and background-independent QTL often had higher additive ( a ) effects. In contrast, the environment or background-dependent QTL were probably due to weak expression of QTL. QTL by environment interaction effects were in the opposite direction of a effects and/or epistasis effects. Four QTL and one QTL could be identified (2.0 < LOD < 9.06) in the HC and HB populations, respectively, across three environments (swHCA2-1, swHCC2-1, swHCD1b-1, swHCA2-2 (linked to Satt233, Satt424, Satt460, Satt428, respectively) and swHBA1-1(Satt449). Seven QTL could be identified in all three RIL populations in at least one location. Two QTL could be identified in the three RIL populations across three environments. These two QTL may have greater potential for use in marker-assisted selection of seed weight in soybean.

... 标记Satt376还被Han等^[26]定位到与大豆单粒重相关, 并且定位区间为Satt376~Satt277, 与本文定位到控制α ...

2004

1.513

0.0

... -生育酚含量相关位于第19染色体上的标记Sat_320和Satt313, 前人研究证明Satt313标记与种子含油量和耐铝胁迫相关^[27,28], 而这个标记未见报道与维生素E组分相关, 这可能是由于遗传背景不同所致 ...

2011

1.175

0.0

Korir

, Qi

, Wang

, Zhao

, Yu

, Chen

, Gai

A study on relative importance of additive, epistasis and unmapped QTL for aluminum tolerance at seedling stage in soybean.

Plant Breed, 2011, 130: 551-562

1 Soybean Research Institute of Nanjing Agricultural University, National Center for Soybean Improvement, National Key Laboratory for Crop Genetics and Germplasm Enhancement, Nanjing, 210095, China 2 Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China 3 Corresponding author, E-mail: sri@njau.edu.cn

> With 3 figures and 6 tables

2005

11.808

0.0

Della P

Progress in the dissection and manipulation of vitamin E synthesis. Trends Plant Sci, 2005, 10: 574-579

... -TMT^[29] ...

1998

0.0

Shintani

, Della P

Elevating the vitamin E content of plants through metabolic engineering.

Science, 1998, 282: 2098-2100

Department of Biochemistry, Mail Stop 200, University of Nevada, Reno, NV 89557, USA.

... -TMT基因Glyma12g01690^[30]不在同一个染色体, 而与尿黑酸叶绿基转移酶HPT (GenBank登录号为DQ231061)基因^[31]位于相同染色体(Gm09: 18 579 763~18 585 148), 但与Satt617标记(Gm09: 31 816 370~31 816 414)相距较远 ...

2006

3.347

0.0

Venkatesh T

, Karunanand aa

, Free D

, Rottnek J

, Baszis S

, Valentin H

Identification and characterization of an Arabidopsis homogentisate phytyltransferase paralog.

Planta, 2006, 223: 1134-1144

Tocochromanols (tocopherols and tocotrienols) are micronutrients with antioxidant properties synthesized by photosynthetic bacteria and plants that play important roles in animal and human nutrition. There is considerable interest in identifying the genes involved in tocochromanol biosynthesis to allow transgenic modification of both tocochromanol levels and tocochromanol composition in agricultural crops. The first committed reaction in tocopherol biosynthesis is the condensation of homogentisic acid (HGA) with phytyldiphosphate or geranylgeranyldiphosphate, catalyzed by the homogentisate phytyltransferase (VTE2) or by the homogentisate geranylgeranyl transferase (HGGT). In this study, we describe the identification of conserved amino acid sequences within VTE2 and HGGT and the application of these conserved sequences for a motif analysis resulting in the discovery of a VTE2 -paralog in the Arabidopsis genome. We designated this new gene VTE2-2 and renamed the old VTE2 to VTE2-1 . Seed-specific expression of VTE2-2 in Arabidopsis resulted in increased seed-tocopherol levels, similar to the transgenic expression of VTE2-1 . Bioinformatics analysis revealed that VTE2-2 is conserved in both monocotyledonous and dicotyledonous plants and is distinct from VTE2-1 and HGGT.

1.Monsanto Company 800 N. Lindbergh Boulevard St. Louis MO 63167 USA 2.Monsanto Company 700 Chesterfield Parkway West Chesterfield MO 63017 USA 3.Monsanto Company Calgene Campus, 1920 Fifth Street Davis CA 95616 USA

2008

0.0

... 大豆维生素E与年份方差未达显著水平, 这与李灵娟^[32]研究甘蓝DH群体在多环境下总生育酚的合成量不随生长环境变化而稳定表达相似 ...