利用双向回交导入系定位水稻苗期耐亚铁毒和锌毒的QTL

引用本文

张建, AijazAhmedSOOMRO, 柴路, 崔彦茹, 王小倩, 郑天清, 徐建龙, 黎志康. 利用双向回交导入系定位水稻苗期耐亚铁毒和锌毒的QTL. 作物学报, 2013, 39(10): 1754-1765
[ZHANG Jian, Aijaz Ahmed SOOMRO, CHAI Lu, CUI Yan-Ru, WANG Xiao-Qian, ZHENG Tian-Qing, XU Jian-Long, LI Zhi-Kang. Mapping of QTL for Iron and Zinc Toxicity Tolerance at Seedling Stage Using a Set of Reciprocal Introgression Lines of Rice. Acta Agronomica Sinica, 2013, 39(10): 1754-1765] 复制到剪切板

Permissions

This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

利用双向回交导入系定位水稻苗期耐亚铁毒和锌毒的QTL

张建, AijazAhmedSOOMRO, 柴路, 崔彦茹, 王小倩, 郑天清^*, 徐建龙^*, 黎志康

中国农业科学院作物科学研究所 / 农作物基因资源与基因改良国家重大科学工程, 北京 100081

^*通讯作者(Corresponding author): 徐建龙, E-mail:xujianlong@caas.cn

收稿日期:2013-06-04

基金:本研究由国家高技术研究发展计划(863计划)项目(2012AA101101)和引进国际先进农业科学技术计划(948计划)项目(2011-G2B)资助。

摘要

铁和锌是水稻生长必需的微量元素,但在低洼或酸性土壤中, 过量的二价铁和锌对水稻生长具毒害作用,最终造成生物量和产量下降。为探讨水稻苗期耐亚铁毒、锌毒的遗传机制, 利用优质粳稻品种Lemont和高产籼稻品种特青为亲本构建的高代双向回交导入系和308个在染色体上均匀分布的SNP标记剖析耐亚铁毒、锌毒相关的QTL。从双向导入系共检测到42个影响耐亚铁毒、锌毒相关性状如苗高、苗干重、根干重以及胁迫与对照相对值的QTL, 多数位点增强亚铁毒、锌毒抗性的有利等位基因来自Lemont。其中同时在2个背景下表达的QTL有4个, 占定位QTL总数的9.52%, 说明大多数QTL的表达具有明显的遗传背景效应。同一遗传背景下同时影响耐亚铁毒和锌毒的QTL有9个, 其中QSdw5在2个背景中均被检测到, 其效应大小和方向一致, 说明水稻苗期耐亚铁毒、锌毒之间存在遗传重叠位点。因此, 通过分子标记辅助选择从Lemont中导入或聚合有利的遗传重叠区域, 可以提高特青对亚铁毒、锌毒的抗性水平。

关键词: 耐亚铁毒害; 耐锌毒害; QTL; SNP; 水稻

Mapping of QTL for Iron and Zinc Toxicity Tolerance at Seedling Stage Using a Set of Reciprocal Introgression Lines of Rice

ZHANG Jian, Aijaz Ahmed SOOMRO, CHAI Lu, CUI Yan-Ru, WANG Xiao-Qian, ZHENG Tian-Qing, XU Jian-Long^*, LI Zhi-Kang

Institute of Crop Sciences / National Key Facility for Crop Gene Resources & Genetic Improvement, Chinese Academy of Agricultural Sciences, Beijing 100081, China

Abstract

Ferrum and zinc are essential microelements for rice. However, rice growing in lowland or acid soil is easily subjected to ferrous iron and zinc toxicities which harm plant growth and finally result in deduction of biomass and yield in production. The aim of this study was to identify quantitative trait locus (QTL) underlying ferrous iron toxicity tolerance (FTT) and zinc toxicity tolerance (ZTT) of rice using a reciprocal advanced backcross introgression lines, which was derived from the cross between Lemont (japonica) and Teqing (indica), and 308 evenly distributed single nucleotide polymorphism (SNP) markers developed from the two parents. A total of 42 putative QTLs affecting shoot height (SH), shoot dry weight (SDW), and root dry weight (RDW) under control and stress conditions and for the relative value of the stress to the control were detected, the alleles at most loci improving the tolerance of ferrous and zinc toxicities were from Lemont. Among them, only four QTLs (9.5%) were detected under the two backgrounds, indicating the expression of most QTLs is specific to genetic background. Nine QTLs were detected from the same genetic background affecting the tolerance of both ferrous and zinc toxicities, in whichQSdw5was expressed under the two backgrounds with the same direction and similar quantity of gene additive effect, suggesting that there is a genetic overlap between FTT and ZTT at seedling stage in rice. It is likely, therefore, to improve Teqing’s FTT and ZTT by introgressing and pyramiding Lemont favorable alleles at the overlapping QTLs via marker-assisted selection.

Keyword: Ferrous iron toxicity tolerance; Zinc toxicity tolerance; Quantitative trait locus (QTL); Single nucleotide polymorphism (SNP); Rice

Show Figures

引言

铁和锌是水稻生长必需的微量元素, 但土壤中铁锌的过量积累会对水稻生长带来毒害。其中, 亚铁毒害是低洼地水稻栽培中面临的主要生理病害, 尤其是在热带和亚热带的酸性土壤中^[1]。在厌氧环境中, 铁主要以二价形式存在, 极易被水稻吸收并过量积累而造成亚铁毒害^[2]。而在二价铁污染的土壤中, 水稻也容易发生锌离子毒害^[3]。水稻遭受铁、锌毒害后均会抑制生长, 进而造成生物量和产量下降^[4,5]。

目前, 对水稻苗期耐亚铁毒害的遗传研究较多, 通过考察在Fe²⁺胁迫下的耐亚铁毒害的相关性状, 如叶斑指数(leaf bronzing index, LBI)、苗高、根长、地上部苗干重、根干重、分蘖数以及地上部相对苗干重等, 定位到一些耐亚铁毒害的QTL^{[6,7,8,9,10,11]}。对水稻在耐锌毒害生理方面的研究报道较多^[12,13], 而对其遗传方面的研究较少, Dong^[14]以苗期受害等级为耐锌毒害的指标共检测到3个主效QTL。植物中的锌铁控制运转相关蛋白(zinc-regulated transporters, iron- regulated transporter like protein, ZIP)参与细胞内锌和铁的跨膜运输, 调节植物细胞内锌、铁平衡与分配^[15]。在高浓度Fe²⁺和Zn²⁺胁迫条件下, 水稻中会生成活性氧, 进而破坏细胞膜、蛋白和核酸, 存在的自由基又会氧化叶绿素, 使光合作用降低, 最终导致光合产物下降^[16,17]。目前虽然定位到一些与耐亚铁毒、锌毒相关的QTL^{[6,7,8,9,10,11,14]}, 但由于采用不同的遗传群体和分子标记, 而且评价指标各异, 因此定位到的结果很难相互比较, 也未见有研究水稻苗期耐亚铁毒和锌毒遗传关系的报道。

本试验采用Lemont/特青构建的高代双向回交导入系分别进行水稻苗期耐亚铁毒、锌毒的QTL定位, 试图分析水稻苗期耐亚铁毒和锌毒的遗传机制, 为分子标记辅助改良水稻耐亚铁毒和锌毒害水平提供参考。

1 材料与方法

1.1 试验材料

以美国南部的优质粳稻品种Lemont和我国高产的籼稻品种特青配置杂种F₁, F₁植株分别与双亲回交, 分别形成Lemont和特青背景的双向回交BC₁F₁群体, 从双向回交后代随机选单株分别与各自的轮回亲本进行2~4次不等的连续回交并经自交多代稳定, 最后分别获得183个Lemont背景导入系(LT-IL), 包括30份BC₂F₅、122份BC₃F₄和31份BC₄F₃, 204个特青背景导入系(TQ-IL), 包括117份BC₂F₅、62份BC₃F₄和25份BC₄F₃。

1.2 苗期耐铁、锌毒害的鉴定

试验于2012年6月7日至9月4日在中国农业科学院作物科学研究所温室进行, 通过湿帘和排气扇调节温室内的温、湿度, 在整个生长和胁迫期间的昼夜平均温度分别为30℃和25℃左右, 湿度约为60%。从每株系挑选饱满的种子约50粒浸种、催芽, 选取露白整齐均一的种子播于底部带有尼龙网的泡沫板(10行×13列)里。用pH为5.0的清水培养7 d, 然后换用Yoshida^[18]营养液培养至二叶一心期分别开始进行亚铁毒、锌毒胁迫处理。亚铁毒处理采用FeSO₄·7H₂O, 将营养液Fe²⁺浓度提高至280 mg L^-1(对照为2.0 mg L^-1), 锌毒处理采用ZnSO₄·7H₂O, 将营养液Zn²⁺浓度提高至180 mg L^-1(对照为0.01 mg L^-1), 每5 d更换一次营养液, 每2 d调节pH至5.0。处理和对照均设2次重复, 完全随机排列。处理20 d后, 测量处理和对照的苗高, 地上部苗干重和根干重。

1.3 SNP基因型分析

在北京大学生命科学院完成SNP位点的挑选、芯片制定和SNP基因型分型。先对特青和Lemont进行全基因组重测序, 分别得到6 644 677 680和 5 236 481 956 bp的测序数据量。以日本晴的MSU6.1版的基因组序列作为参考基因组(参考基因组为 372 317 567 bp), 将测序数据与参考基因组比对, 获得基因组序列一致的碱基数, 特青为6 154 118 505 bp, Lemont为4 969 908 313 bp, 特青和Lemont的基因组覆盖率分别为92.39%和93.56%。在一致序列的基础上, 将检测到的基因型与参考序列之间存在多态性的位点过滤, 得到一个初步的SNP数据集。根据reads覆盖度的不同, 在特青和Lemont之间得到了223 001个SNP (reads≥3), 206 433个SNP (reads≥4), 186 504个SNP (reads≥5)。在SNP数据集的基础上, 挑选384个在基因组上均匀分布(平均每5 cM选1个)的SNP, 经美国Illumina公司打分, 从SNP数据集中选分值高的部分SNP进行实验验证, 发现当reads覆盖度≥9的时候, SNP实验验证的成功率高达99%。随后将SNP数据集根据基因组上的分布等分成384份, 并根据Illumina分值和SNP的reads覆盖度从每一份中挑选1个SNP位点。

以筛选出来在双亲间有多态的384个SNP标记送往美国Illumina公司制作芯片, 采用SNP芯片分别鉴定双向导入系群体基因型, 由于存在基因型数据的缺失, 最终选取2个背景下308个相同的SNP位点, 用于构建12条染色体的SNP遗传连锁图。

1.4 数据分析及QTL定位

利用SAS PROC CORR^[19]分析各性状间的相关关系, t测验检验双亲各性状间平均数的差异显著性。以对照与胁迫条件下相关表型数据和相对值[(处理值-对照值)/对照值×100%]为输入数据, 结合308个SNP标记基因型数据, 采用IciMapping 3.2软件^[20]检测影响2种环境下各性状及胁迫与对照相对值的QTL, 选用LOD≥2.5作为阈值来判断QTL的存在与否。同时采用SAS PROC GLM检测各性状的QTL ( P<0.001), 与ICIM定位的结果比较验证。对于QTL初定位, 凡落在5000 kb (相当于20 cM)范围内且基因效应方向相同的归并为同一个QTL。当一个QTL与2个或多个标记连锁时, 以 F值最高的标记作为与QTL连锁标记列出^[21]。为避免使用一个固定的QTL阈值而导致第II类错误的发生, 对所有在一个环境下(对照或胁迫条件)检测到的QTL, 重新检测其在另一个环境下在概率0.05水平的显著性, 只要达到0.05水平显著, 也一并列出该QTL的统计参数^[22]。

2 结果与分析

2.1 双向导入系物理图谱及群体结构分析

Lemont和特青之间有多态的308个SNP标记覆盖基因组的大小为360.81 Mb, 平均占国际水稻基因组测序计划^[23]公布的基因组大小(382.15 Mb)的94.40%, 变幅从第9染色体的88.13%到第12染色体的98.78%。根据水稻基因组12条染色体的总遗传距离(1526.8 cM), 308个SNP标记在12条染色体上分布较为均匀, 平均约每5 cM存在1个SNP位点, 2个相邻标记间的最大距离为6.1 Mb。

双向导入系的大部分遗传背景与各自的轮回亲本相同, 其中Lemont背景导入系中轮回亲本Lemont基因组平均占83.86%, 变幅为20.24%~ 100.00%, 特青背景导入系轮回亲本特青基因组平均占91.05%, 变幅为60.95%~100.00% (图1)。

2.2 亲本及导入系耐亚铁毒、锌毒相关性状的表现

由于受环境影响, 双亲在两次试验中表现略有差异, 但变化趋势基本一致(表1)。Lemont和特青在Fe²⁺或Zn²⁺胁迫条件下, 苗高、苗干重、根干重均低于正常条件。在2次试验中, 正常和胁迫条件下特青的苗高、苗干重均显著或极显著高于Lemont; 在Zn²⁺胁迫条件下, 特青仅苗干重极显著高于Lemont。

	Figure Option View Download New Window
	图1 双向导入系群体特青基因组所占比例的频率分布Fig. 1 Frequency distribution of the Teqing genome in the reciprocal introgression line populations

Lemont在Fe²⁺胁迫与正常条件下苗高的相对值显著高于特青, 其余2个性状的相对值差异均未达到显著水平。Lemont在Zn²⁺胁迫与正常条件下苗高、苗干重的相对值显著高于特青, 而根干重的相对值差异未达到显著水平。Lemont在Fe²⁺胁迫和Zn²⁺胁迫的苗干重比正常条件下平均损失分别为9.65%和15.50%, 而特青分别为13.75%和21.10%, 表明Lemont比特青具有较强的耐亚铁毒、锌毒的特性。双向回交导入系群体的耐亚铁毒、锌毒相关性状都表现出超亲分离, 呈连续变异(表1)。

2.3 耐亚铁毒和锌毒相关性状间的相关

正常条件下, 双向回交导入系群体的CSH、CSDW、CRDW彼此呈极显著正相关(表2)。在Fe²⁺或Zn²⁺胁迫条件下, 除Lemont背景导入系FSH与FRDW或ZSH与ZRDW相关不显著外, 其余性状间均显著或极显著正相关。Fe²⁺或Zn²⁺胁迫与对照相对值性状之间, 苗高和苗干重相对值均呈极显著正相关, 而苗高相对值与根干重相对值相关不显著, 表明地上部和地下部对Fe²⁺和Zn²⁺胁迫有不同抗性机制。双向回交导入系群体同一性状在Fe²⁺或Zn²⁺胁迫下与正常条件下呈显著或极显著正相关, 表明SH、SDW与RDW在Fe²⁺或Zn²⁺胁迫与正常条件下的变化趋势一致; 而同一性状Fe²⁺或Zn²⁺胁迫与对照相对值与对照值之间均存在显著或极显著负相关, 表明正常条件下生长量越大则耐Fe²⁺或Zn²⁺能力越差。同一性状在Fe²⁺胁迫与Zn²⁺胁迫之间均呈显著或极显著正相关, 表明双向导入系耐亚铁毒与耐锌毒之间有可能存在一些相同的遗传机制。

表1 双向导入系苗期Fe²⁺和Zn²⁺胁迫下相关性状的表现Table 1 Performance of ferrous and zinc toxicity tolerance related traits in the reciprocal introgression lines

条件 Condition	性状¹⁾ Trait	背景 Background	Lemont (P₁)	特青 (P₂)	P₁-P₂²⁾	平均数±标准差 Mean ± SD	变异系数 CV (%)	变幅 Range
对照 Control	CSH (cm)	TQ	26.5	31.7	-5.2^**	31.2±3.2	10.1	20.8 to 41.5
		LT	24.8	29.2	-4.4^*	24.7±2.0	7.9	18.1 to 31.0
	CSDW (mg)	TQ	48.3	59.4	-11.1^**	54.0±10.1	18.8	28.3 to 84.0
		LT	28.0	35.8	-7.8^**	30.9±6.3	20.4	18.0to 53.2
	CRDW (mg)	TQ	7.5	9.4	-1.9^*	9.4±2.0	21.4	4.3 to 16.3
		LT	6.7	7.8	-1.1	6.8±1.1	16.6	4.3 to 11.3
铁处理 Fe stress	FSH (cm)	TQ	24.5	27.1	-2.6^*	29.8±3.0	10.1	20.7 to 39.1
		LT	22.7	24.8	-2.1^*	22.5±1.7	7.3	16.6 to 26.8
	FSDW (mg)	TQ	43.3	51.5	-8.2^**	48.8±9.7	19.8	22.9 to 78.3
		LT	25.4	30.7	-5.3^**	26.1±5.2	20.1	16.4 to 43.6
	FRDW (mg)	TQ	6.8	8.2	-1.4	8.4±1.8	21.5	3.3 to 14.3
		LT	6.5	7.6	-1.1	5.5±1.1	20.5	2.5 to 10.0
锌处理 Zn stress	ZSH (cm)	TQ	22.2	23.1	-0.9	23.6±2.3	9.8	16.3 to 32.1
		LT	21.3	22.2	-0.9	21.2±1.3	6.4	16.9 to 23.9
	ZSDW (mg)	TQ	41.8	48.4	-6.6^**	41.0±8.4	20.4	18.8 to 63.8
		LT	23.1	27.3	-4.2^**	23.6±4.1	17.5	13.5 to 41.7
	ZRDW (mg)	TQ	6.3	7.1	-0.8	6.1±1.6	25.9	2.9 to 12.0
		LT	6.1	6.9	-0.8	5.3±1.0	18.8	2.5 to 9.3
铁相对值 Fe relative value (%)	FRSH	TQ	-7.5	-14.5	6.96^*	-4.3±2.9	67.4	-27.3 to 0
		LT	-8.5	-15.1	6.60^*	-8.9±5.0	56.8	-27.8 to -0.2
	FRSDW	TQ	-10.4	-13.3	2.95	-9.6±7.4	77.8	-39.2 to 0
		LT	-9.3	-14.2	4.96	-15.1±10.5	69.3	-49.2 to 0
	FRRDW	TQ	-9.3	-12.8	3.43	-11.0±9.2	83.5	-44.4 to 0
		LT	-3.0	-2.6	-0.42	-18.2±12.1	66.6	-50.0 to 0
锌相对值 Zn relative value (%)	ZRSH	TQ	-16.2	-27.1	10.90^*	-24.0±5.1	21.0	-35.1 to -4.2
		LT	-14.1	-24.0	9.86^*	-14.1±5.3	37.5	-30.8 to -0.6
	ZRSDW	TQ	-13.5	-18.5	5.06	-23.5±10.0	43.4	-48.7 to -1.8
		LT	-17.5	-23.7	6.24^*	-22.3±10.8	48.5	-53.0 to -0.3
	ZRRDW	TQ	-16.0	-24.5	8.47	-34.1±15.7	45.9	-49.2 to 0
		LT	-9.0	-11.5	2.58	-20.7±11.6	56.3	-54.3 to 0

¹⁾CSH: 对照条件下的苗高; CSDW: 对照条件下的苗干重; CRDW: 对照条件下的根干重; FSH: 铁处理条件下的苗高; FSDW: 铁处理条件下的苗干重; FRDW: 铁处理条件下的根干重; ZSH: 锌处理条件下的苗高; ZSDW: 锌处理条件下的苗干重; ZRDW: 锌处理条件下的根干重; FRSH: 铁处理与对照的苗高相对值; FRSDW: 铁处理与对照的苗干重相对值; FRRDW: 铁处理与对照的根干重相对值; ZRSH: 锌处理与对照的苗高相对值; ZRSDW: 锌处理与对照的苗干重相对值; ZRRDW: 锌处理与对照的根干重相对值。^{2) *}和^**分别表示差异达0.05和0.01显著水平。

¹⁾CSH: shoot height under control condition; CSDW: shoot dry weight under control condition; CRDW: root dry weight under control condition; FSH: shoot height under ferrous stress condition; FSDW: shoot dry weight under ferrous stress condition; FRDW: root dry weight under ferrous stress condition; ZSH: shoot height under zinc stress condition; ZSDW: shoot dry weight under zinc stress condition; ZRDW: root dry weight under zinc stress condition; FRSH: relative value of shoot height under ferrous condition to control condition; FRSDW: relative value of shoot dry weight under ferrous condition to control condition; FRRDW: relative value of root dry weight under ferrous condition to control condition; ZRSH: relative value of shoot height under zinc condition to control condition; ZRSDW: relative value of shoot dry weight under zinc condition to control condition; ZRRDW: relative value of root dry weight under zinc condition to control condition.^{2) *}and^**indicate the significance at P<0.05 and 0.01 based on t-tests.

表1 双向导入系苗期Fe²⁺和Zn²⁺胁迫下相关性状的表现Table 1 Performance of ferrous and zinc toxicity tolerance related traits in the reciprocal introgression lines

2.4 耐亚铁毒相关性状的QTL定位

从特青背景导入系共检测到对照条件下、Fe²⁺或Zn²⁺胁迫条件下以及处理与对照的相对值影响苗高、苗干重和根干重的23个QTL, 分布在除第4、第10和第12染色体外的9条染色体上。从Lemont背景导入系共检测到上述3种条件下影响苗高、苗干重和根干重的QTL 22个, 分布在除第9和第10染色体外的10条染色体上(图2)。

从特青背景导入系检测到对照和Fe²⁺胁迫下影响SH的QTL 9个, 包括对照条件下的9个和Fe²⁺胁迫下的5个(表3)。除 QSh1、 QSh3a 和 QSh6a外, 其余位点增加苗高的等位基因均来自特青。从Lemont背景导入系检测到对照和Fe²⁺胁迫下影响SH的QTL 6个(表3), 包括对照条件下的5个和Fe²⁺胁迫下的5个。除 QSh5和 QSh11外, 其余位点减小SH的等位基因均来自特青。根据QTL在不同环境下的表达差异, 将这些QTL分成3组。第1组是仅在对照条件下检测到的5个QTL, 包括特青背景下检测到的4个( QSh3a、 QSh6b、 QSh8b和 QSh11)和Lemont背景下检测到的1个( QSh5); 第2组是在对照与Fe²⁺胁迫条件下同时检测到的9个QTL, 包括特青背景下检测到的5个( QSh1、 QSh5、 QSh7、 QSh8a和 QSh9)和Lemont背景下检测到的4个( QSh2a、 QSh3b、 QSh6a和 QSh11); 第3组是仅在Fe²⁺胁迫条件下在Lemont背景下检测到的1个QTL ( QSh2b)。另外, 从双向导入系检测到5个影响Fe²⁺胁迫与对照相对值的QTL, 包括特青背景下检测到的2个( QSh6b和 QSh9)和Lemont背景下检测到的3个( QSh2a、 QSh5和 QSh6a), 除特青背景导入系的 QSh6b外, 其余位点减小SH相对值即降低耐亚铁毒的不利等位基因均来自特青。

从特青背景导入系检测到对照和Fe²⁺胁迫条件下影响SDW的QTL 6个(表3), 包括第2组在对照条件下和Fe²⁺胁迫条件下同时检测到的5个QTL ( QSdw1、 QSdw5、 QSdw7a、 QSdw8a和 QSdw9)和第3组仅在Fe²⁺胁迫条件下检测到的1个QTL ( Qsw8b)。除 QSdw1外, 其余位点增加苗干重的等位基因来自特青。从Lemont背景导入系检测到对照和Fe²⁺胁迫条件下影响SDW的QTL 6个(表3), 包括第1组仅在对照条件下检测到的3个( QSdw4、 QSdw5和 QSdw7a), 第2组在Fe²⁺胁迫和对照条件下同时检测到的1个( QSdw2a)和第3组仅在Fe²⁺胁迫条件下检测到的2个( QSdw6a和 QSdw11)。除 QSdw6a外, 其余位点增加苗干重的等位基因均来自特青。此外, 从双向导入系检测到影响Fe²⁺胁迫与对照相对值的QTL 5个, 包括特青背景下检测到的2个( QSh5和 QSh9) 和Lemont背景下检测到的3个( QSdw2a、 QSdw5和 QSdw6a), 除 QSdw2a外, 其余位点减小SDW相对值即降低耐亚铁毒的不利等位基因均来自特青。

从特青背景导入系检测到对照和Fe²⁺胁迫条件下影响RDW的QTL 4个(表3), 包括第1组的 QRdw2和 QRdw4a, 第2组的 QRdw8和第3组的 QRdw1。除 QRdw1外, 其余位点增加根干重的等位基因均来自特青。从Lemont背景导入系检测到对照和Fe²⁺胁迫条件影响RDW的QTL 7个(表3), 包括第1组的 QRdw1、 QRdw4b、 QRdw5b和 QRdw12, 第2组的 QRdw5a和第3组的 QRdw2和 QRdw6。除 QRdw6外, 其余位点增加干重的等位基因均来自特青。此外, 从双向导入系检测到影响Fe²⁺胁迫与对照相对值的QTL 4个, 包括特青背景下检测到的2个( QRdw1和 QRdw4a)和Lemont背景下检测到的2个( QRdw1和 QRdw6), 除Lemont背景下的 QRdw1外, 其余位点减小RDW相对值即降低耐亚铁毒的不利等位基因均来自特青。

2.5 耐锌毒相关性状的QTL定位

从特青背景导入系检测到对照和Zn²⁺胁迫条件下影响SH的QTL 10个(表3), 包括第1组仅在对照条件下检测到的3个QTL ( QSh5、 QSh8a和 QSh11), 第2组在Zn²⁺胁迫和对照条件下同时检测到的6个QTL ( QSh1、 Qsh3a、 QSh6b、 Qsh7、 QSh8b和 Qsh9)和第3组仅在Zn²⁺胁迫条件下检测到的 QSh2b。除 QSh7、 QSh8b和 QSh9外, 第1组的3个QTL和 Qsh7、 QSh8b和 Qsh9位点增加苗高的等位基因均来自特青, 而其余4个位点增加苗高的等位基因来自Lemont。从Lemont背景导入系检测到对照和Zn²⁺胁迫条件下影响SH的QTL 7个(表3), 包括第1组仅在对照条件下检测到的3个QTL ( QSh3b、 QSh5和 QSh11), 第2组在Zn²⁺胁迫和对照条件下同时检测到的2个QTL ( QSh2a和 QSh6a)和第3组仅在Zn²⁺胁迫条件下检测到的2个QTL ( QSh2b和 QSh7), 除 QSh5和 QSh11外所有QTL增加苗高的等位基因均来自Lemont。另外, 从双向导入系检测到影响Zn²⁺胁迫和对照相对值的5个QTL, 包括特青背景下检测到的3个( QSh2b、 QSh7和 QSh8a)和在Lemont背景下的2个( QSh5和 QSh7), 其减小SH相对值即降低耐锌毒的不利等位基因均来自特青。

从特青背景导入系检测到对照和Zn²⁺胁迫条件下影响SDW的QTL 8个(表3), 包括第1组的3个( QSdw1、 QSdw7a和 QSdw8a), 第2组的2个( QSdw5和 QSdw9)和第3组的3个( QSdw2b、 QSdw7b和 QSdw8b)。除 QSdw1和 QSdw2b外, 其余位点增加苗干重的等位基因均来自特青。从Lemont背景导入系检测到对照和Zn²⁺胁迫条件下影响SDW的QTL 7个(表3), 包括第1组的2个( QSdw2a和 QSdw7a), 第

表2 双向导入系群体苗期Fe²⁺和Zn²⁺胁迫下相关性状的相关系数Table 2 Correlation coefficients between ferrous and zinc toxicity tolerance related traits in the reciprocal introgression lines

	对照苗高 CSH	对照苗干重 CSDW	对照根干重 CRDW	铁处理苗高 FSH	铁处理苗干重 FSDW	铁处理根干重 FRDW	锌处理苗高 ZSH	锌处理苗干重 ZSDW	锌处理根干重 ZRDW	铁苗高相对值 FRSH	铁苗干重相对值 FRSDW	铁根干重相对值 FRRDW	锌苗高相对值 ZRSH	锌苗干重相对值 ZRSDW	锌根干重相对值 ZRRDW
对照苗高 CSH		0.75^***	0.51^***	0.71^***	0.50^***	0.30	0.60^***	0.46^**	0.33	-0.50^***	-0.38	-0.24	-0.63^***	-0.38^*	-0.24
对照苗干重 CSDW	0.85^***		0.77^***	0.51^***	0.72^***	0.44^**	0.34^*	0.65^***	0.51^***	-0.41^**	-0.40^**	-0.35^*	-0.60^***	-0.49^***	-0.34^*
对照根干重 CRDW	0.68^***	0.83^***		0.40^**	0.64^***	0.34^*	0.25	0.64^***	0.58^***	-0.22	-0.21	-0.66	-0.39*	-0.23	-0.58
铁处理苗高 FSH	0.82^***	0.70^***	0.53^***		0.65^***	0.28	0.70^***	0.46^**	0.33	0.25	0.16	-0.15	-0.22	-0.08	-0.10
铁处理苗干重 FSDW	0.76^***	0.80^***	0.65^***	0.84^***		0.45^**	0.32	0.66^***	0.52^***	0.12	0.33	-0.21	-0.30	-0.12	-0.19
铁处理根干重 FRDW	0.58^***	0.62^***	0.53^***	0.58^***	0.80^***		0.20	0.28	0.39^**	-0.08	-0.01	0.45^**	-0.19	-0.23	0.02
锌处理苗高 ZSH	0.76^***	0.59^***	0.45^**	0.67^***	0.65^***	0.50^***		0.47^**	0.29	-0.06	-0.07	-0.08	0.19	0.03	0
锌处理苗干重 ZSDW	0.68^***	0.68^***	0.56^***	0.58^***	0.70^***	0.55^***	0.90^***		0.71^***	-0.07	-0.04	-0.35*	-0.12	0.22	-0.04
锌处理根干重 ZRDW	0.56^***	0.61	0.51^***	0.44^***	0.54^***	0.45^***	0.60^***	0.78		-0.05	-0.04	-0.22	-0.13	0.13	0.30
铁苗高相对值 FRSH	-0.33^*	-0.27	-0.26	0.26	0.11	0.13	-0.15	-0.17	-0.21		0.72^***	0.14	0.59^***	0.41^**	0.19
铁苗干重相对值 FRSDW	-0.29	-0.48^**	-0.39^*	0.05	0.10	0.12	-0.04	-0.12	-0.20	0.59^***		0.20	0.42^**	0.47^**	0.20
铁根干重相对值 FRRDW	-0.14	-0.27	-0.50	0.13	0.11	0.42^**	0.08	-0.04	-0.07	0.47^**	0.60^***		0.23	0.04	0.56^***
锌苗高相对值 ZRSH	-0.40^**	-0.42	-0.37^*	-0.26	-0.22	-0.16	0.28	0.19	0.09	0.30	0.38^*	0.33		0.52^***	0.30
锌苗干重相对值 ZRSDW	-0.26	-0.47^**	-0.37^*	-0.18	-0.19	-0.12	0.27	0.28	0.13	0.15	0.55^***	0.33	0.76^***		0.35^*
锌根干重相对值 ZRRDW	-0.10	-0.20	-0.44^**	-0.06	-0.09	-0.06	0.24	0.22	0.51	0.09	0.19	0.44^***	0.49^**	0.51^***

右上角为Lemont背景导入系性状间的相关，左下角为特青背景导入系性状间的相关。性状缩写见表1。^*,^**和^***分别表示性状间的相关达到0.05、0.01和0.001显著水平。

Numbers below diagonal are for Teqing background-ILs, and those above diagonal for Lemont background-ILs. Trait abbreviations are the same as those given in Table 1.^*,^**, and^*** stand for significant correlation at P≤ 0.05, 0.01, and 0.001, respectively.

表3 双向导入系中检测到影响耐亚铁毒和锌毒相关性状的主效QTLTable 3 Main-effect QTL affecting ferrous and zinc toxicity tolerance related traits detected in the reciprocal introgression lines

	Figure Option View Download New Window
	图2 双向导入系物理图谱及影响水稻苗期耐亚铁毒和锌毒相关性状的主效QTL在染色体上的分布Fig. 2 Distribution of QTL affecting ferrous and zinc toxicity tolerance related traits detected in the two reciprocal introgression lines derived from the cross of Lemont/Teqing

2组的2个( QSdw4和 QSdw5)和第3组的3个( QSdw6b、 QSdw8b和 QSdw11), 所有QTL增加苗干重的等位基因均来自特青。从双向导入系还检测到影响Zn²⁺胁迫SDW和对照相对值的7个QTL, 包括特青背景下检测到的4个( QSdw2b、 QSdw5、 QSdw7b和 QSdw8a)和在Lemont背景下的3个( QSdw2a、 QSdw5和 QSdw6b), 除 QSdw2a和 QSdw6b外, 其余位点减小SDW相对值即降低耐锌毒的不利等位基因均来自特青。

从特青背景导入系在对照和Zn²⁺胁迫条件下检测到影响RDW的QTL 4个, 包括第1组的3个( QRdw2、QRdw4a和 QRdw8)和第3组的1个( QRdw3), 除 QRdw3外, 其增加根干重的等位基因均来自特青。在Lemont背景导入系中, 检测到对照和Zn²⁺胁迫条件下影响RDW的QTL 6个, 包括第1组的3个( QRdw1、QRdw5a和 QRdw5b)、第2组的2个( QRdw4b和 QRdw12)和第3组的1个( QRdw2), 所有位点增加SDW的等位基因均来自特青。另外, 从双向导入系还检测到影响Zn²⁺胁迫RDW和对照相对值的QTL各1个, 即 QRdw8和 QRdw5b, 其减小RDW相对值即降低耐锌毒的不利等位基因均来自特青。

3 讨论

3.1 水稻苗期耐亚铁毒、锌毒QTL及其遗传重叠

利用双向导入系群体检测到影响对照、胁迫及胁迫与对照相对值条件下耐亚铁毒和锌毒相关性状(苗高、地上部苗干重和根干重)的QTL中, 真正对耐亚铁毒和锌毒有贡献的QTL仅有两类, 一是通过胁迫与对照的相对值检测到的QTL (例如 QSdw5), 其解释了由于胁迫所造成表型变化的稳定性; 二是能够在对照和胁迫条件下同时检测到, 且基因效应大小相似、方向一致的稳定表达的QTL (例如 QSh1)。在特青背景下检测到耐亚铁毒的QTL有12个, 耐锌毒的QTL有10个, 其中同时影响耐亚铁毒和锌毒的QTL有5个, 包括 QSh7、 QSh8a、 QSdw5、 QSdw9和 QRdw8, 占QTL总数的22.7%; Lemont背景检测到耐亚铁毒的QTL和耐锌毒的QTL各10个, 其中共同影响耐亚铁毒和锌毒的QTL也有5个, 包括 QSh2a、 QSh5、 QSh6a、 QSdw2a和 QSdw5, 占QTL总数的25%。其中, QSdw5在2个背景下均被检测到, 并且其加性效应的大小和方向一致。这表明水稻苗期耐亚铁毒和锌毒存在一些的遗传重叠位点, 共同影响对亚铁毒和锌毒的抗性。

在特青背景导入系的3个染色体区段同时定位到耐亚铁毒的2个或3个相关性状的QTL, 包括第1染色体9 765 185~10 822 085区间的 QSh1、 QSdw1和 QRdw1, 第8染色体18 396 373~19 556 500区间的 QSh8a和 QRdw8, 第9染色体5 390 612~7 237 057区间的 QSh9和 QSdw9。从Lemont背景导入系同时定位到耐亚铁毒的2个或3个相关性状的QTL的染色体区段有3个, 包括第2染色体8 454 851~ 9 511 612区间的 QSh2a和 QSdw2a, 第5染色体 17 301 701~19 546 328区间的 QSh5和 QSdw5, 第6染色体8 126 604~9 656 257区间的 QSh6a、 QSdw6a和 QRdw6。从特青背景导入系的3个染色体区段同时定位到耐锌毒2个或3个相关性状的QTL, 包括第2染色体23 557 743~24 626 496区间的 QSh2b和 QSdw2b, 第7染色体27 495 091~29 303 506区间的 QSh7和 QSdw7b, 第8染色体18 396 373~19 556 500区间的 QSh8a、 QSdw8a和 QRdw8。从Lemont背景导入系的2个染色体区段同时定位到耐锌毒的2个以上相关性状的QTL, 包括第2染色体8 454 851~ 9 511 612区间的 QSh2a和 QSdw2b, 第5染色体 17 301 701~19 546 328区间的 QSh5、 QSdw5和 QRdw5b。上述重叠位点中, 除第2染色体区段的 QSh2a和 QSdw2a基因效应方向相反外, 其余位点在不同性状中的基因效应方向一致。

3.2 QTL定位的遗传背景效应

本研究检测到影响苗期耐亚铁毒、锌毒相关性状的42个QTL中, 同时在2个背景下表达的仅有4个(表2), 占9.5%, 说明绝大多数QTL的表达具有明显的遗传背景效应。以往利用Lemont和特青构建的双向导入系群体研究水稻数量性状的试验中也发现相同的现象, 就是虽然双向导入系的亲本相同, 但是在不同回交背景下同时表达的QTL很少^{[24,25,26,27]}。在2个背景同时定位到的QTL中, 虽然遗传背景影响主效QTL的表达, 但并不影响基因作用的方向^[25,27]。在本研究中, 除 QRdw1外, 其余3个位点在2个背景中基因作用方向一致, 即降低耐亚铁毒和锌毒的不利等位基因来自特青, 与上述现象基本一致。

3.3 不同群体定位到耐亚铁毒和锌毒的QTL 比较

根据SNP标记的物理位置, 将本研究定位到的耐亚铁毒和锌毒的QTL与前人的研究结果比较, 发现特青背景导入系影响耐亚铁毒的 QSh1、 QSdw1和 QRdw1与Zn²⁺胁迫相关的 qZNT-1定位在一起^[14]; 影响双向导入系耐亚铁毒的 QSdw5与铁胁迫下影响苗高的 QPh5定位在一起^[9]; 特青背景导入系第9染色体上影响亚铁毒的 QSh9和 QSdw9与影响铁胁迫下相对苗高的 qTFS-9定位在一起^[10]。另外, 耐亚铁毒和锌毒相关的 QSh5、 QSdw5、 QRdw5b与水稻Zn²⁺运输相关的基因 OsZIP2 (23 262 252~23 264 744)^[28]紧邻; 在Lemont背景导入系检测到的与耐锌毒相关的 QRdw4b与Fe²⁺运输相关的基因 OsYSL2 (26 652 325~ 26 655 465)^[29]位于同一区间。在特青背景导入系检测到的影响铁毒的 QSh9和 QSdw9与Fe²⁺运输相关的转录因子 OsWRKY80 (24 734 518~ 24 734 438)^[30]相邻。这些位点的等位性还有待进一步证明。

3.4 不同亚铁毒和锌毒抗性指标性状的评价

水稻苗期耐Fe²⁺和Zn²⁺胁迫的评价体系直接影响到相关抗(耐)基因检出的效果。评价体系包括处理方式、处理时期、处理浓度、处理时间长度和考察性状等几个方面。考察性状有受害等级、苗高、根长、地上部苗干重、根干重、含水量等一般指标^[7,8,9], 也有的考察叶绿素指数、电解质外渗率、MDA含量和离子浓度等生理指标^[1,6,11]。从考察的性状来看, 评价受害等级操作不方便, 而且难于定量比较; 测量相关生理指标又比较繁琐且难于精确测定。通过不同评价体系定位到的QTL均有不同之处^[31], 而且定位结果也难于相互比较, 因此, 建立高效的水稻苗期耐亚铁毒和锌毒胁迫的评价体系显得十分重要。

水稻应对短时间高浓度离子胁迫和长时间低浓度离子胁迫可能存在不同的抗性机制, 借鉴前人的研究方法和预备试验, 选择在二叶一心期胁迫处理20 d, 考察易于测定的苗高、地上部苗干重、根干重等指标, 同时用处理和对照的相对值评价, 这样可以消除幼苗生长本身引起的差异, 其大小可直接反映株系对Fe²⁺和Zn²⁺毒害的抗性。从本研究定位结果来看, 通过苗高和地上部苗干重可以检测到很多共同的位点, 其效应大小和方向也基本一致, 而地上部苗干重较苗高易于考察且试验测量误差较小, 因此我们认为地上部苗干重是较为有效的考察指标。

3.5 利用分子标记辅助选择提高水稻苗期Fe²⁺和Zn²⁺胁迫抗性的启示

从苗高和苗干重来看, 特青显著高于Lemont, 但是在Fe²⁺和Zn²⁺胁迫条件下的干重损失上, 特青低于Lemont, 说明Lemont比特青具有较强的抗性。我们检测到影响Fe²⁺和Zn²⁺胁迫下与对照相对值的QTL, 大部分增加相对值即提高抗性的有利等位基因来自Lemont, 但是在一些位点提高相对值的有利等位基因来自特青, 如 QSh6b、 QSdw2a和 QSdw6b等。因此利用分子标记辅助选择聚合双亲的有利等位基因, 就完全有可能提高品种对亚铁毒和锌毒的抗性水平, 培育出符合育种目标的新品种。

4 结论

利用双向导入系群体分别检测到22个和20个影响水稻苗期耐亚铁毒和锌毒相关性状的QTL, 其中4个在2个背景下共同表达, 9个在同一遗传背景下同时影响耐亚铁毒和锌毒, 即为耐两种金属毒害的遗传重叠位点。利用分子标记辅助选择聚合耐亚铁毒和耐锌毒的遗传重叠位点, 可以提高水稻苗期对亚铁毒和锌毒的抗性水平。

参考文献

View Option

[1]	Zhou J-L((周建林)), Tang J-J((唐建军)). Physiological indices for evaluating iron-toxicity tolerance of rice (Oryza sativa L. ) . Chin J Appl Ecol((应用生态学报)), 2001, 12(1): 159-160 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.7261]
[2]	Gross J, Stein R J, Fett-Neto A G, Fett J P. Iron homeostasis related genes in rice . Genet Mol Biol, 2003, 26: 477-497 [本文引用:1] [JCR: 0.634]
[3]	Song A, Li P, Li Z J, Fan F L, Nikolic M, Liang Y C. The alleviation of zinc toxicity by silicon is related to zinc transport and antioxidative reactions in rice . Plant Soil, 2011, 344: 319-333 [本文引用:1] [JCR: 2.733]
[4]	Fageria N K, Rabelo N A. Tolerance of rice cultivars to iron toxicity . Plant Nutr, 1987, 10: 653-661 [本文引用:1] [JCR: 1.017]
[5]	Ajay Rathore V S. Effect of Zn²⁺ stress in rice on growth and photosynthetic process . Photosynthctica, 1995, 1: 571-584 [本文引用:1]
[6]	Wu P, Hu B, Liao C Y, Zhu J M, Wu Y R, Senadhira D, Paterson A H. Characterization of tissue tolerance to iron by molecular markers in different lines of rice . Plant Soil, 1998, 203: 217-226 [本文引用:3] [JCR: 2.733]
[7]	Wan J L, Zhai H Q, Wan J M, Ikehashi H. Detection and analysis of QTLs for ferrous iron toxicity tolerance in rice, Oryza sativa L . Euphytica, 2003, 131: 201-206 [本文引用:3] [JCR: 1.554]
[8]	Wan J L, Zhai H Q, Wan J M, Yasui H, Yoshimura A. Detection and analysis of QTLs associated with resistance to ferrous iron toxicity in rice (Oryza sativa L. ) using recombinant inbred lines . Acta Agron Sin, 2004, 30: 329-333 [本文引用:3] [CJCR: 1.8267]
[9]	Wan J L, Zhai H Q, Wan J M. Mapping of QTLs for ferrous iron toxicity tolerance in rice (Oryza sativa L. ) . Acta Genet Sin, 2005, 32(11): 1156-1166 [本文引用:4] [JCR: 1.883] [CJCR: 0.6313]
[10]	Ye H-X((叶红霞)), Li M((李梅)), Zhuang J-Y((庄杰云)), Shen S-Q((沈圣泉)). Analysis of gene effects of tolerance to high Fe²⁺ Stress at seedling stage in rice . Mol Plant Breed((分子植物育种)), 2007, 5(1): 105-109 (in Chinese with English abstract) [本文引用:3] [CJCR: 0.9082]
[11]	Dufey I, Hakizimana P, Draye X, Lutts S, Bertin P. QTL mapping for biomass and physiological parameters linked to resistance mechanisms to ferrous iron toxicity in rice . Euphytica, 2009, 167: 143-160 [本文引用:3] [JCR: 1.554]
[12]	Liu M-L((刘美玲)), Liu X-N((刘湘南)), Li T((李婷)), Xiu L-N((修丽娜)). Analysis of hyperspectral singularity of rice under Zn pollution stress . Trans CSAE((农业工程学报)), 2010, 26(3): 191-196 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.7156]
[13]	Xu J-M((徐建明)), Li C-S((李才生)), Mao S-G((毛善国)), Wang X((汪鑫)), Fan C-Y((樊趁英)), Huang P-F((黄鹏飞)), Li Y-W((李耀文)). Effect of Zinc on rice seedlings growth and activity of SOD and POD . J Anhui Agric Sci((安徽农业科学)), 2008, 36(3): 877-878 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.6624]
[14]	Dong Y J, Ogawa T, Lin D Z, Koh H J, Kamiunten H, Matsuo M, Cheng S H. Molecular mapping of quantitative trait loci for zinc toxicity tolerance in rice seedling (Oryza sativa L. ) . Field Crops Res, 2005, 95: 420-425 [本文引用:3] [JCR: 2.474]
[15]	Wang H((汪洪)), Jin J-Y((金继运)). The physiological and molecular mechanisms of Zinc uptake, transport, and hyperaccumulation in plants: a review . Plant Nutr Fert Sci((植物营养与肥料学报)), 2009, 15(1): 225-235 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.4752]
[16]	Doran G, Eberbach P, Helliwell S. The impact of rice plant roots on the reducing conditions in flooded rice soil . Chemosphere, 2006, 63: 1892-1902 [本文引用:1] [JCR: 3.206]
[17]	Chung M J, Walker P A, Brown R W, Hogstrand C. Zincmediated gene expression offers protection against H₂O₂-induced cytotoxicity . Toxicol Appl Pharmacol, 2005, 205: 225-236 [本文引用:1] [JCR: 4.447]
[18]	Yoshida S, Forno D A, Cock J H, Gomez K A. Laboratory Manual for Physiological Studies of Rice, 3rd edn . Manila, The Philippines: IRRI, 1976. pp1-83 [本文引用:1]
[19]	SAS Institute. SAS/STAT User’s Guide . Cary NC, USA: SAS Institute, 1996. pp25-36 [本文引用:1]
[20]	Li H H, Ye G Y, Wang J K. A modified algorithm for the improvement of composite interval mapping . Genetics, 2007, 175: 361-374 [本文引用:1] [JCR: 4.007]
[21]	Xu J L, Lafitte H R, Gao Y M, Fu B Y, Torres R, Li Z K. QTLs for drought avoidance and tolerance identified in a set of rand om introgression lines of rice . Theor Appl Genet, 2005, 111: 1642-1650 [本文引用:1] [JCR: 3.297]
[22]	Li Z K. QTL mapping in rice: a few critical considerations. In: Khush G S, Brar D S, Hardy B, eds. Rice Genetics IV . Science Publishers, Inc. , and International Rice Research Institute, New Delhi, India, and Los Banos, The Philippines, 2001. pp153-172 [本文引用:1]
[23]	International Rice Genome Sequencing Project. The map-based sequence of the rice genome . Nature, 2005, 436: 793-800 [本文引用:1] [JCR: 36.28]
[24]	Mei H W, Xu J L, Li Z K, Luo L J. QTLs influencing panicle size detected in two reciprocal introgressive line (IL) populations in rice (Oryza sativa L. ) . Theor Appl Genet, 2006, 112: 648-656 [本文引用:1] [JCR: 3.297]
[25]	Xie X-W((谢学文)), Xu M-R((许美容)), Zang J-P((藏金萍)), Sun Y((孙勇)), Zhu L-H((朱苓华)), Xu J-L((徐建龙)), Zhou Y-L((周永力)), Li Z-K((黎志康)). Genetic background and environmental effect on expression of QTL for sheath blight resistance in reciprocal intro-gression lines of rice . Acta Agron Sin((作物学报)), 2008, 34(11): 1885-1893 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.8267]
[26]	Yang J((杨静)), Sun Y((孙勇)), Cheng L-R((程立锐)), Zhou Z((周政)), Wang Y((王韵)), Zhu L-H((朱苓华)), Cang J((苍晶)), Xu J-L((徐建龙)), Li Z-K((黎志康)). Genetic background effect on QTL mapping for salt tolerance revealed by a set of reciprocal introgression line populations in rice . Acta Agron Sin((作物学报)), 2009, 35(6): 974-982 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.8267]
[27]	Wang Y((王韵)), Cheng L-R((程立锐)), Sun Y((孙勇)), Zhou Z((周政)), Zhu L-H((朱苓华)), Xu Z-J((徐正进)), Xu J-L((徐建龙)), Li Z-K((黎志康)). Genetic background effect on QTL expression of heading date and plant height and their interaction with environment in reciprocal introgression lines of rice . Acta Agron Sin((作物学报)), 2009, 35(8): 1386-1394 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.8267]
[28]	Ramesh S A, Shin R, Eide D J, Schachtman D P. Differential metal selectivity and gene expression of two zinc transporters from rice . Plant Physiol, 2003, 133: 126-134 [本文引用:1] [JCR: 6.535]
[29]	Koike S, Inoue H, Mizuno D. OsYSL2 is a rice metal-nicotianamine chelates transporter that is regulated by iron and expressed in the phloem . Plant J, 2004, 39: 415-424 [本文引用:1] [JCR: 6.16]
[30]	Ricachenevsky F K, Sperotto R A, Menguer P K, Fett J P. Identification of Fe-excess-induced genes in rice shoots reveals a WRKY transcription factor responsive to Fe, drought and senescence . Mol Biol Rep, 2010, 37: 3735-3745 [本文引用:1] [JCR: 2.929]
[31]	Dufey I, Hiel M P, Hakizimana P, Draye X, Lutts S, Kone B, Drame K N, Konate K A, Sie M, Bertin P. Multienvironment quantitative trait loci mapping and consistency across environments of resistance mechanisms to ferrous iron toxicity in rice . Crop Sci, 2012, 52: 539-550 [本文引用:1] [JCR: 1.641]

2001

0.0

1.7261

... 其中, 亚铁毒害是低洼地水稻栽培中面临的主要生理病害, 尤其是在热带和亚热带的酸性土壤中^[1] ...

... 考察性状有受害等级、苗高、根长、地上部苗干重、根干重、含水量等一般指标^[7,8,9], 也有的考察叶绿素指数、电解质外渗率、MDA含量和离子浓度等生理指标^[1,6,11] ...

2003

0.634

0.0

... 在厌氧环境中, 铁主要以二价形式存在, 极易被水稻吸收并过量积累而造成亚铁毒害^[2] ...

2011

2.733

0.0

Plant Soil. 2011, 344(1-2):319 - 333 DOI:10.1007/s11104-011-0749-3

The alleviation of zinc toxicity by silicon is related to zinc transport and antioxidative reactions in rice

Alin Song(1) Ping Li(1) Zhaojun Li(1) Fenliang Fan(1) Miroslav Nikolic(2) Yongchao Liang(1)

1.Ministry of Agriculture Key Laboratory of Crop Nutrition and Fertilization, Institute of Agricultural Resources and Regional Planning, Chinese Academy of Agricultural Sciences, Beijing, 100081 People’s Republic of China
2.Institute for Multidisciplinary Research, University of Belgrade, Kneza Vi?eslava 1, 11030 Belgrade, Republic of Serbia

The objective of this study is to elucidate the roles of silicon (Si) in enhancing tolerance to excess zinc (Zn) in two contrasting rice (Oryza sativa L.) cultivars: i.e. cv. TY-167 (Zn-resistant) and cv. FYY-326 (Zn-sensitive). Root morphology, antioxidant defense reactions and lipid peroxidation, and histochemical staining were examined in rice plants grown in the nutrient solutions with normal (0.15 μM) and high (2 mM) Zn supply, without or with 1.5 mM Si. Significant inhibitory effects of high Zn treatment on plant growth were observed. Total root length (TRL), total root surface area (TRSA) and total root tip amount (TRTA) of both cultivars were decreased significantly in plants treated with high Zn, whereas these root parameters were significantly increased when Zn-stressed plants were supplied with 1.5 mM Si. Supply of Si also significantly decreased Zn concentration in shoots of both cultivars, indicating lower root-to-shoot translocation of Zn. Moreover, superoxide dismutase (SOD), catalase (CAT), and asorbate peroxidase (APX) activities were increased, whereas malondialdehyde (MDA) and hydrogen peroxide (H₂O₂) concentrations were decreased in Si-supplied plants of both Zn-sensitive and Zn-resistant rice cultivars exposed to Zn stress. These alleviative effects of Si, further confirmed by the histochemical staining methods, were more prominent in the Zn-resistant cultivar than in the Zn-sensitive one. Taken together, all these results suggest that Si-mediated alleviation of Zn toxicity is mainly attributed to Si-mediated antioxidant defense capacity and membrane integrity. The possible role of Si in reduction of root-to-shoot translocation of Zn can also be considered.

... 而在二价铁污染的土壤中, 水稻也容易发生锌离子毒害^[3] ...

1987

1.017

0.0

... 水稻遭受铁、锌毒害后均会抑制生长, 进而造成生物量和产量下降^[4,5] ...

1995

0.0

... 水稻遭受铁、锌毒害后均会抑制生长, 进而造成生物量和产量下降^[4,5] ...

1998

2.733

0.0

... 目前, 对水稻苗期耐亚铁毒害的遗传研究较多, 通过考察在Fe²⁺胁迫下的耐亚铁毒害的相关性状, 如叶斑指数(leaf bronzing index, LBI)、苗高、根长、地上部苗干重、根干重、分蘖数以及地上部相对苗干重等, 定位到一些耐亚铁毒害的QTL^{[6,7,8,9,10,11]} ...

... 目前虽然定位到一些与耐亚铁毒、锌毒相关的QTL^{[6,7,8,9,10,11,14]}, 但由于采用不同的遗传群体和分子标记, 而且评价指标各异, 因此定位到的结果很难相互比较, 也未见有研究水稻苗期耐亚铁毒和锌毒遗传关系的报道 ...

2003

1.554

0.0

Euphytica. 2003, 131(2):201 - 206 DOI:10.1023/A:1023915710103

Detection and analysis of QTLs for ferrous iron toxicity tolerance in rice,Oryza sativa L.

Jian-lin Wan(1) Hu-qu Zhai(2) Jian-min Wan(1) H. Ikehashi(3)

1.Nanjing Agricultural University State Key laboratory of crop genetics and germplasm enhancement Nanjing 210095 People's Republic of China
2.Chinese Academy of Agricultural Sciences Beijing 100081 People's Republic of China
3.Nihon University Department of Breeding 1866 Kameino, Fujisawa-shi Japan

A mapping population of 96 BC₁F₉lines (Backcross Inbred Lines: BILs),derived by a single-seed descent method rom a backcross of Nipponbare (japonica) / Kasalath (indica // Nippon are, was used to detect quantitative trait loci (QTLs) for leaf bronzing index (LBI), stem dry weight (SDW), tiller number (TN) and root dry weight (RDW) under Fe²⁺ stress condition in rice. Two parents and 96 BILs were phenotyped for the traits by growing them in Fe²⁺ toxicity nutrient solution. A total of four QTLs were detected on chromosome 1 and 3, respectively, with LOD of QTLs ranging from 3.17 to 7.03. One QTL controlling LBI, DW, N and RDW was located at the region of C955-C885 on chromosome 1, and their contributions to whole variation were 20.5%, 36.9%, 43.9% and 38.8%,respectively. The QTL located at the region of C955-C885 on chromosome 1 may be important to ferrous iron toxicity tolerance in rice. Another QTL for SDW and RDW was located at the region of C25-C515 on chromosome 3, with respective contributions of 47.9% and 35.0% to whole variation. Further, two QTLs on chromosome 1 were located for RDW at the region of R2329-R210 and for TN at the region of R1928-C178. Comparing with the other mapping results, the QTL located at the region of C955-C885 on chromosome 1 was identical with the results reported previously. There is a linkage between a TL detected under Fe²⁺ stress condition for stem and root dry weight and a QTL detected under phosphorus-deficiency condition for dry weight on chromosome 3.

2004

0.0

1.8267

Acta Agron Sin. 2004, 30(4):329 - 333

Detection and Analysis of QTLs Associated with Resistance to Ferrous Iron Toxicity in Rice ( Oryza sativa L. ),Using Recombinant Inbred Lines

利用重组自交群体检测水稻抗亚铁毒胁迫的QTLs

WAN Jian-Lin;ZHAI Hu-Qu;WAN Jian-Min;Hideshi Yasui;Atsushi Yoshimura

万建林;翟虎渠;万建民;安井秀;吉村醇

State Key laboratory of crop genetics & germplasm enhancement , Jiangsu Plant Gene Engineering Research Center , Nanjing Agricultural University, Nanjing 210095

A mapping population of 81 F11 lines ( recombinant inbred lines: RILs ), derived from a cross between a japonica variety Kinmaze and an indica variety DV85 by the single-seed descent methods, was used to detect quantitative trait loci ( QTLs ) for leaf bronzing index ( LBI ) and plant height ( PH )

潜育性水稻田广泛分布于中国、斯里兰卡、印度、印度尼西亚、塞拉里昂、利比亚、尼日利亚、哥伦比亚和菲律宾等国，其中我国南方稻区就有670万公顷低产潜育性水稻田。该类水稻田还原性强，矿质营养失调，尤以Fe2+ 过量积累，对水稻生长发育产生不良的逆境胁迫作用。培育抗亚铁毒的水稻品种是简便、经济有效地提高稻谷产量的重

2005

1.883

0.6313

J Genet Genomics. 2005, 32(11):1156 - 1166

Mapping of QTLs for ferrous iron toxicity tolerance in rice, Oryza sativa L.

水稻耐亚铁毒QTLs的定位

WAN Jian-Lin1, ZHAI Hu-Qu2, WAN Jian-Min1

万建林1;翟虎渠2;万建民1

1.State Key Laboratory of Crop Genetics and Germplasm Enhancement, Nanjing Agricultural University, Nanjing 210095, China; 2.Chinese Academy of Agricultural Sciences, Beijing 100081, China

Ferrous iron toxicity is the main factor limiting the productivity of rice in gleyic paddy soils. In this study, an F2 and an equivalent F3 populations derived from a japonica/indica cross of rice, Longza8503/IR64, were raised under iron-enriched solution cultures , and used to map QTLs controlling ferrous iron toxicity tolerance. A genetic linkage map consisting of 101 SSR markers was constructed to determine the position and nature of quantitative trait loci (QTLs) affecting Fe2+ toxicity tolerance. Three characters, i.e., leaf bronzing index (LBI), plant height (PH) and maximum root length (MRL) were evaluated for the F2 plants and F3 lines and the parents at the seedling stage in nutrient solution. A total of 20 QTLs for LBI, PH and MRL under the Fe2+ stress were detected over 10 of the 12 rice chromosomes, reflecting multigenic control of these traits. QTLs controlling LBI were located at the region of RM315-RM212 on chromosome 1, RM6-RM240 on chromosome 2 and RM252-RM451 on chromosome 4. Comparing with the other mapping results, 1) the QTL for LBI located at the region of RM252-RM451 on chromosome 4 was identical with the QTL for decreased chlorophyll content on a rice function map. Another QTL for LBI located at the region of RM315-RM212 on chromosome 1 was linked with the QTL for chlorophyll content which located at the region of C178-R2635 on a rice function map. 2)The third QTL for LBI located at the region of RM6-RM240 on choromosome 2 was linked with the QTL for potassium uptake located at the region of RZ58-CDO686 under potassium deficiency stress.

亚铁毒是潜育性水稻土中限制水稻产量的主要因子。利用龙杂8503/IR64的F2 和等价的F3群体，在营养液中培养来定位耐亚铁毒的QTLs。通过构建101SSR标记的遗传连锁图谱来确定耐亚铁毒QTLs的位置和特性。借助叶片棕色斑点指数、株高和最大根长三个性状，利用营养液在水稻苗期来评价F2单株、F3群体和亲本龙杂8503、IR64，共检测到叶片棕色斑点指数、株高和最大根长的QTLs20个，分布在水稻的10条染色体上，表明这些性状受多基因控制。控制叶片棕色斑点指数的QTLs分别定位在第1染色体的RM315~RM212、第2染色体的 RM6~RM240 和第4染色体的 RM252~RM451之间。与前人的研究结果比较发现，1）位于第4染色体 RM252~RM451之间的控制叶片棕色斑点指数的QTL与水稻功能图谱上控制叶绿素含量减少的QTL的位置一致。另一个位于第1染色体的RM315-RM212之间的控制叶片棕色斑点指数的QTL与水稻功能图谱上位于C178-R2635之间控制叶绿素含量的QTL连锁。2）位于第2染色体RM6-RM240之间的第3个控制叶片棕色斑点指数的QTL与位于RZ58-CDO686的控制钾吸收的QTL连锁。

... 影响双向导入系耐亚铁毒的QSdw5与铁胁迫下影响苗高的QPh5定位在一起^[9] ...

2007

0.0

0.9082

Mol Plant Breed. 2007, (1):105 - 109

Analysis of Gene Effects of Tolerance to High Fe2+ Stress at Seedling Stage in Rice

水稻幼苗对多浓度Fe2+胁迫的QTL联合检测

YeHongxia^[1];LiMei^[1];ZhuangJieyun^[2];ShenShengquan^[1]

叶红霞^[1];李梅^[1];庄杰云^[2];沈圣泉^[1]

用珍汕97B/密阳46的RIL作图群体,以基本营养液培养为对照(CK),设2种Fe2+(160 mg/L和240 mg/L)作为胁迫处理,以处理20 d的幼苗相对苗高(%)(处理苗高/对照苗高×100%)作为幼苗耐Fe2+胁迫程度的评价指标,进行2个环境的QTL联合检测分析.结果表明,RIL群体株系间对Fe2+胁迫反应差异较大,明显出现超亲分离.共检测到4个耐Fe2+胁迫的主效QTL,即qTFS-1-1、qTFS-3、qTFS-6和qTFS-9,分别位于第1、3、6和9染色体上,可解释14.96%的表型变异,它们与Fe2+胁迫浓度并无显著的G×E互作,表明这4个耐性基因在不同浓度Fe2+胁迫处理下,均可稳定表达.试验还检测到3对耐Fe2+胁迫的加性×加性上位性互作,共可解释7.16%的表型变异,涉及第1、2、7、9、12等5条染色体,该3对上位性互作与Fe2+不同浓度胁迫处理也无显著G×E互作.

... 特青背景导入系第9染色体上影响亚铁毒的QSh9和QSdw9与影响铁胁迫下相对苗高的qTFS-9定位在一起^[10] ...

2009

1.554

0.0

Euphytica. 2009, 167(2):143 - 160 DOI:10.1007/s10681-008-9870-7

QTL mapping for biomass and physiological parameters linked to resistance mechanisms to ferrous iron toxicity in rice

Inès Dufey(1) Patrice Hakizimana(1) Xavier Draye(1) Stanley Lutts(2) Pierre Bertin(1)

1.Université catholique de Louvain Unité d’Ecophysiologie et Amélioration végétale Croix du Sud, 2 bte 11 1348 Louvain-La-Neuve Belgium
2.Université catholique de Louvain Unité de Biologie végétale Croix du Sud, 2 bte 11 1348 Louvain-La-Neuve Belgium

Lowland rice is often subject to iron toxicity which may lead to yield reduction. In order to cope with this nutrient disorder, plants have developed resistance strategies. The aim of this research was to assess morphological and physiological parameters linked to iron toxicity resistance mechanisms and to identify quantitative trait loci (QTLs) involved in their genetic determinism. A segregating population consisting of 164 recombinant inbred lines (RILs) derived from a cross between Azucena and IR64 was tested twice in hydroponics at the vegetative stage at 0 and 250 mg Fe²⁺ l⁻¹. Morphological traits were measured on all 164 RILs. Physiological traits, which were too time-consuming to allow their measurement on all the population, were measured on the two parents and extreme individuals only, selected on the basis of their leaf bronzing index and shoot dry weight. A total of 24 putative QTLs was identified on chromosomes 1, 2, 3, 4, 7 and 11 for leaf bronzing index, shoot water content, shoot and root dry weight, relative variation of shoot and root dry weight, shoot iron concentration, stomatal resistance and chlorophyll content index. Several QTLs were detected in overlapping regions for different parameters. The pertinence of phenotyping extreme RILs only for a QTL analysis is discussed in this study. The QTL analysis allowed to better understand the physiological response of rice in the presence of an excess of ferrous iron, inclusive the relations existing between the stomata closure, the shoot water content reduction and the oxidative stress linked to these growth conditions.

2010

0.0

1.7156

Trans CSAE. 2010, (3):191 - 197

Analysis of hyperspectral singularity of rice under Zn pollution stress

水稻锌污染胁迫的光谱奇异性分析

LiuMeiling;LiuXiangnan;LiTing;XiuLina

刘美玲;刘湘南;李婷;修丽娜

通过研究锌污染胁迫下水稻的光谱奇异性特征来诊断水稻锌污染水平.根据试验区水稻各个生长期的ASD实测高光谱数据和同步获取的作物与农田土壤重金属含量2 a田间试验数据,利用Daubechies小波系中的"Db5"小波函数对锌污染胁迫水稻的350～1 300 nm光谱波段进行5层小波分解,并利用奇异范围、奇异幅度和奇异指数等参数对水稻光谱奇异性进行定量计算与分析.结果表明,"Db5"小波函数分解的第5层小波系数能精准探测水稻光谱的奇异性,其奇异范围集中在480～850 nm波段范围;奇异幅度从出苗期到分蘖期迅速增加,并达到最大值,在随后的分蘖期→拔节期→开花期→成熟期逐渐减少;奇异指数从出苗期→分蘖期→拔节期→开花期→成熟期逐渐增加;水稻叶片锌含量与奇异幅度及奇异指数之间存在较强的相关关系,其决定系数(R~2)为0.8445,均方根误差 (RMSE)为5.60.光谱奇异性能有效地诊断并定量分析水稻锌污染胁迫状况,从而为作物重金属污染胁迫监测提供重要参考.

Abstract：

The aim of this study is to detect the stress levels of rice under Zn pollution by analyzing the characteristics of hyperspectral singularity. Hyperspectral reflectance and heavy metal concentration from different growth stages of rice in two consecutive years were collected in the field experiment. Wavelet transform was performed to hyperspectral reflectance (350-1 300 nm) by using "Daubechies 5" wavelet function, and three categories of singularity parameters as indicators of hyperspectral singularity of rice under Zn stress were explored, including singularity range (SR), singularity amplitude (SA) and singularity exponent (SE). The results showed that the wavelet coefficient of the fifth decomposition level of "Daubechies 5" wavelet function proved successful for accurately identifying the hyperspectral singularity of rice under Zn stress. SR of rice was concentrated on the region around 480-850 nm of spectral signal under Zn stress. Maximum value of SA occurred at the tillering stage, and the SA increased sharply from seeding stage to tillering stage and then decreased from jointing stage to anthesis stage and to mature stage, while the SE increased constantly from seeding stage to mature stage when rice was growing. Zn concentration in rice leaves had better linearity with the SA and SE with correlation coefficient (R~2) of 0.8445 and root mean square error (RMSE) of 5.60. The hyperspectral singularity applying wavelet transform technique has been shown to indentify and quantitatively analyze the stress levels of rice under Zn pollution effectively and provide important reference for detecting other metal-induced stress on crop.

... 对水稻在耐锌毒害生理方面的研究报道较多^[12,13], 而对其遗传方面的研究较少, Dong^[14]以苗期受害等级为耐锌毒害的指标共检测到3个主效QTL ...

2008

0.0

0.6624

J Anhui Agric Sci. 2008, (3):877 - 878

Effect of Zinc on Rice Seedlings Growth and Activity of SOD and POD

锌对水稻幼苗生长及体内SOD、POD活性的影响

徐建明;李才生;毛善国;汪鑫;樊趁英;黄鹏飞;李耀文

研究了水稻幼苗期利用不同浓度锌处理对幼苗生长及体内SOD、POD活性的影响.结果表明:在水稻幼苗2叶1心期,喷施1.0 mg/L的锌溶液能够促进幼苗的生长发育,苗高、根长、苗鲜重、根鲜重及根系活力均显著提高,分别比对照增加23.05%、10.74%、29.77%、23.27%和89.97%;根内的SOD、POD活性比对照增加243.48%和43.60%,叶片中的SOD、POD活性分别比对照增加172.77%和75.35%.根据试验结果建议,在水稻幼苗期喷施锌溶液,浓度以1.0 mg/L为宜.

... 对水稻在耐锌毒害生理方面的研究报道较多^[12,13], 而对其遗传方面的研究较少, Dong^[14]以苗期受害等级为耐锌毒害的指标共检测到3个主效QTL ...

2005

2.474

0.0

... 对水稻在耐锌毒害生理方面的研究报道较多^[12,13], 而对其遗传方面的研究较少, Dong^[14]以苗期受害等级为耐锌毒害的指标共检测到3个主效QTL ...

... 3 不同群体定位到耐亚铁毒和锌毒的QTL 比较根据SNP标记的物理位置, 将本研究定位到的耐亚铁毒和锌毒的QTL与前人的研究结果比较, 发现特青背景导入系影响耐亚铁毒的QSh1、QSdw1和QRdw1与Zn²⁺胁迫相关的qZNT-1定位在一起^[14] ...

2009

0.0

1.4752

Plant Nutr Fert Sci. 2009, (1):225 - 235

The physiological and molecular mechanisms of zinc uptake,transport,and hyperaccumulation in plants:A review

植物对锌吸收运输及积累的生理与分子机制

WANGHong;JINJi-yun

汪洪;金继运

锌是植物必需的营养元素,也是重金属污染元素之一.现代分子生物学的发展,极大地推动了植物体内与锌吸收运输有关转运蛋白的研究.目前发现,锌铁控制运转相关蛋白(ZIP)、自然抵抗相关巨噬细胞蛋白NRAMP、重金属ATPase酶、阳离子扩散协助蛋白CDF、Mg2+/H+的反向交换转运蛋白MHX等运输蛋白参与细胞内Zn2+离子的跨膜运输,调节植物细胞内zn2+平衡与分配.利用数量遗传学手段在在水稻上已找到与缺锌植株死亡率和叶片青铜病发牛率有关QTL位点.而在Thlaspi caerulescens和Arabidopsis halleri植物上鉴定出控制锌含量的QTLs,为寻找控制植物高效积累zn的遗传基础规律打下了基础.

... 植物中的锌铁控制运转相关蛋白(zinc-regulated transporters, iron- regulated transporter like protein, ZIP)参与细胞内锌和铁的跨膜运输, 调节植物细胞内锌、铁平衡与分配^[15] ...

2006

3.206

0.0

Chemosphere. 2006, 63(11):1892 - 1902 DOI:10.1016/j.chemosphere.2005.10.027

The impact of rice plant roots on the reducing conditions in flooded rice soils

Abstract

The impact of oxygen diffusion from plant roots on the soil redox in the root zone in flooded rice bays was investigated using two Australian rice growing soils. The rates of production of Fe(II) and Mn(II) in pore water resulting from the reduction of soil minerals was used to gauge the extent of development of anaerobic conditions and the time taken for equilibrium to establish. Soil concentrations of readily reducible Fe were 13–28 times greater than Mn, making Fe a more reliable indicator of redox conditions than Mn. In addition, Mn(II) concentrations reached equilibrium far more rapidly than Fe, which made the identification of any contribution to soil redox by oxygen diffusing from rice plant roots difficult to observe. Dissection of soil cores showed that more than 80% of the rice root mass occurred in the top 4 cm of soil, suggesting that any contribution roots may make to the redox potential of the flooded soils would occur in this region. However, studies conducted indicated that the diffusion of oxygen from the surface floodwater into soil pore water in the 2.5 cm layer of soil was so substantial that it would mask any contribution made by rice plant roots to the overall soil redox in this root zone.

... 在高浓度Fe²⁺和Zn²⁺胁迫条件下, 水稻中会生成活性氧, 进而破坏细胞膜、蛋白和核酸, 存在的自由基又会氧化叶绿素, 使光合作用降低, 最终导致光合产物下降^[16,17] ...

2005

4.447

0.0

1976

0.0

... 0的清水培养7 d, 然后换用Yoshida^[18]营养液培养至二叶一心期分别开始进行亚铁毒、锌毒胁迫处理 ...

1996

0.0

... 4 数据分析及QTL定位利用SAS PROC CORR^[19]分析各性状间的相关关系, t测验检验双亲各性状间平均数的差异显著性 ...

2007

4.007

0.0

... 2软件^[20]检测影响2种环境下各性状及胁迫与对照相对值的QTL, 选用LOD≥2 ...

2005

3.297

0.0

Theor Appl Genet. 2005, 111(8):1642 - 1650 DOI:10.1007/s00122-005-0099-8

QTLs for drought escape and tolerance identified in a set of random introgression lines of rice

J. L. Xu(12) H. R. Lafitte(2) Y. M. Gao(1) B. Y. Fu(1) R. Torres(2) Z. K. Li(12)

1.Chinese Academy of Agricultural Sciences Institute of Crop Sciences/National Key Facility for Crop Gene Resources and Genetic Improvement 12 South Zhong-Guan-Cun Street 100081 Beijing China
2.International Rice Research Institute DAPO Box 7777 Metro Manila Philippines

A large set of 254 introgression lines in an elite indica genetic background were evaluated for grain yield (GY) and related traits under the irrigated (control) and drought (stress) conditions in two consecutive years for genetic dissection of adaptive strategies of rice to water stress. A total of 36 quantitative trait loci (QTLs) affecting heading date (HD), plant height (PH), GY and yield components were identified and most QTLs showed pronounced differential expression either qualitatively or quantitatively in response to drought. These QTLs could be grouped into three major types based on their behaviors under control and stress conditions. Type I included 12 QTLs that expressed under both the stress and non-stress conditions. Type II comprised 17 QTLs that expressed under irrigation but not under stress. Type III included seven QTLs that were apparently induced by stress. The observation that the Lemont (japonica) alleles at all HD QTLs exceptQHd5 resulted in early heading under stress appeared to be responsible for the putative adaptation of Lemont to drought by escaping, whereas the Teqing (indica) alleles at most PH/GY QTLs were consistently associated with increased yield potential and trait stability and thus contributed to DT. Our result that most DT QTLs were non-allelic with QTLs for drought escaping suggests that the two adaptive strategies in the parental lines are under possible negative regulation of two largely non-overlapping genetic systems.

... 同时采用SAS PROC GLM检测各性状的QTL (PF值最高的标记作为与QTL连锁标记列出^[21] ...

2001

0.0

... 05水平显著, 也一并列出该QTL的统计参数^[22] ...

2005

36.28

0.0

... 81 Mb, 平均占国际水稻基因组测序计划^[23]公布的基因组大小(382 ...

2006

3.297

0.0

Theor Appl Genet. 2006, 112(4):648 - 656 DOI:10.1007/s00122-005-0167-0

QTLs influencing panicle size detected in two reciprocal introgressive line (IL) populations in rice (Oryza sativa L.)

H. W. Mei(1) J. L. Xu(2) Z. K. Li(24) X. Q. Yu(1) L. B. Guo(3) Y. P. Wang(3) C. S. Ying(3) L. J. Luo(1)

1.Shanghai Agrobiological Gene Center, 2901 Beidi Road, 201106 Shanghai, China
2.Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, 12 South Zhongguancun St., 100081 Beijing, China
3.China National Rice Research Institute, 359 Tiyuchang Road, 310006 Hangzhou, China
4.Plant Breeding, Genetics and Biotechnology Division, International Rice Research Institute, DAPO Box 7777, Metro Manila, Philippines

Two sets of reciprocal introgression line (IL) populations, i.e., ILs with Lemont as recurrent parent (IL_LT) and ILs with Teqing as recurrent parent (IL_TQ), were developed and evaluated for traits representing panicle size, including primary branch number (PBN), secondary branch number (SBN), and spikelet number per panicle (SNP). Together with the regression to recurrent parent by advanced backcross, transgressive segregations were observed for all traits. Correlation and regression analysis showed that SBN had much higher contribution to SNP than PBN. It was confirmed by the QTL analysis that many common loci were detected between SBN and SNP, in comparison with single common locus between PBN and SNP. One and three main effect QTLs (M-QTLs) were detected for PBN in IL_LT and IL_TQ, respectively. Six M-QTLs per trait per populations were associated with SBN and SNP. Less number and lower contribution of epistasis were detected in IL populations in comparison with mapping result fromF₂ or RI population. There were only four QTLs in fourteen loci (near 30%) commonly detected in both reciprocal IL populations implying the large impact of genetic background on QTLs expression.

... 以往利用Lemont和特青构建的双向导入系群体研究水稻数量性状的试验中也发现相同的现象, 就是虽然双向导入系的亲本相同, 但是在不同回交背景下同时表达的QTL很少^{[24,25,26,27]} ...

2008

0.0

1.8267

Acta Agron Sin. 2008, 34(11):1885 - 1893 DOI:10.3724/SP.J.1006.2008.01885

Genetic Background and Environmental Effects on Expression of QTL for Sheath Blight Resistance in Reciprocal Introgression Lines of Rice

水稻抗纹枯病QTL表达的遗传背景及环境效应

XIE Xue-Wen¹,XU Mei-Rong¹,ZANG Jin-Ping¹,SUN Yong¹,ZHU Ling-Hua¹,XU Jian-Long^1*,ZHOU Yong-Li^1*,LI Zhi-Kang¹²

谢学文¹;许美容¹;藏金萍¹;孙勇¹;朱苓华¹;徐建龙^1,*;周永力^1,*;黎志康^1,2

1 Institute of Crop Sciences, Chinese Academy of Agricultural Sciences / National Key Facility for Crop Gene Resources and Genetic Improvement, Beijing 100081, China; 2 International Rice Research Insititute, DAPO Box 7777, Metro Manila, Philippines

Genetic background and environmental effects of QTL for sheath blight resistance to the isolate RH-9 (Rhizoctonia solani kuhn) were revealed using the reciprocal introgression line populations derived from the cross of Lemont/Teqing. Lemont is highly susceptible while Teqing resistant to RH-9. The relative lesion height (a ratio of lesion height to plant height, RLH) of TQ-ILs was normally distributed whereas that of LT-ILs was apparently inclined to the susceptible parent, Lemont. Total 10 main-effect QTLs and 13 epistatic QTLs affecting sheath blight resistance were mapped under different years and genetic backgrounds. Among them, six main-effect QTLs detected in 2006 were all verified in 2007, suggesting these QTLs had reliable performance across years.QSh4 was the only one QTL expressed under the reciprocal backgrounds and Teqing allele at this locus decreased RLH, suggesting the improvement of resistance level.QSh10a detected in the TQ-ILs and located in the region of RM216–RM311 on the chromosome 10 and QSh10b detected in the LT-ILs in the neighboring region of RM222–RM216 were not the same gene but existed tight linkage as regards to different gene directions in different backgrounds. Most QTLs identified in TQ-ILs were not expressed in LT-ILs, indicating there was evident genetic background effect. By comparative mapping, 8 QTLs detected in this study were located in the same or near regions that associated with sheath blight resistance identified in the previous studies, suggesting these main-effect QTLs could be applied in rice breeding for sheath blight resistance by marker-assisted selection. As indicated in this study, it is an effective way to further improve sheath blight resistance by selecting the QTL stably expressed in different backgrounds or pyramiding different main-effect QTLs.

利用水稻纹枯病菌强致病菌系RH-9人工接种Lemont导入到特青背景的213个近等基因导入系(TQ-ILs)群体和特青导入到Lemont背景的195个近等基因导入系(LT-ILs)群体，定位和分析了水稻抗纹枯病数量性状座位(quantitative trait loci, QTL)及其表达的环境与遗传背景效应。亲本Lemont对RH-9表现为高度感病，特青表现为中等抗病。人工接种后TQ-ILs群体的相对病斑高度(病斑高度与株高比)呈连续正态分布，LT-IL群体则明显偏向感病亲本Lemont。在不同年份和遗传背景下检测到影响纹枯病相对病斑高度的主效QTL 10个和互作QTL 13个，其中2006年在TQ-IL群体定位到的6个主效QTL在2007年均得到验证，表明这些QTL具有较好年度间的重复性。QSh4是唯一在双向导入系背景下表达的QTL，该位点特青等位基因降低相对病斑高度，提高抗性水平。在TQ-ILs群体中定位到位于第10染色体RM216~RM311区间的QSb10a与在LT-IL群体中定位到的位于相邻区间RM222~RM216的QSb10b的基因作用方向不同，推断这两个QTL存在紧密连锁关系。绝大多数在TQ-IL群体中表达的主效及互作QTL在LT-ILs群体中不表达，表明水稻抗纹枯病QTL具有明显的遗传背景效应。通过比较作图，本研究定位到的其中8个QTL在以往不同群体中同样被检测到，这些主效QTL对通过分子标记辅助选择(marker-assisted selection, MAS)培育水稻抗纹枯病育种可能具有应用价值。研究指出，标记辅助选择在不同遗传背景中能稳定表达的QTL或通过聚合不同抗病QTL是进一步提高水稻纹枯病抗性水平的一个有效途径。

... 在2个背景同时定位到的QTL中, 虽然遗传背景影响主效QTL的表达, 但并不影响基因作用的方向^[25,27] ...

2009

0.0

1.8267

Acta Agron Sin. 2009, 35(6):974 - 982

Genetic Background Effect on QTL Mapping for Salt Tolerance Revealed by a Set of Reciprocal Introgression Line Populations in Rice

利用双向导入群体检测遗传背景对耐盐QTL定位的影响

YANG Jing¹,SUN Yong²,CHENG Li-Rui²,ZHOU Zheng²,WANG Yun²,ZHU Ling-Hua²,CANG Jing¹,XU Jian-Long^2*,LI Zhi-Kang²³

杨静¹²，孙勇²，程立锐²，周政²，王韵²，朱苓华²，苍晶¹，徐建龙^2*，黎志康²³

1College of Life Science,Northeast Agricultural University,Harbin 150030,China;2 Institute of Crop Sciences/National Key Facility for Crop Gene Resources and Genetic Improvement,Chinese Academy of Agricultural Sciences,Beijing 100081,China;3 International Rice Research Institute,DAPO Box 777,Metro Manila,Philippines

QTLs for salt tolerance (ST) related traits including score of salt toxicity of leaves (SST), survival days of seedlings (SDS), shoot K⁺concentration (SKC) and shoot Na⁺ concentration (SNC) were detected using the reciprocal introgression lines (ILs) derived from a cross between japonica variety “Lemont” andindica variety “Teqing” under salt stress treatment with the concentration of 140 mmol L^-1 NaCL in green house and phytotron. Both genetic backgrounds of the two sets of ILs were similar to their respective parents. On the average, Lemont genome accounted for 83.8% in Lemont background-ILs whereas 88.9% of Teqing genome in Teqing background-ILs. Continuous variation and transgressive segregation for all ST-related traits were observed in the two IL populations with overlaps of frequency distributions for most traits, suggesting the gene effect of reciprocal introgression of the two parents on each ST-related trait was almost equivalent. Eighteen QTLs for all ST-related traits were identified for each of the two IL populations. No any a common QTL for the same trait was detected in the reciprocal IL populations, indicating there was a strong genetic background effect on expression of ST-QTLs, and also suggesting these ST-QTLs had relatively small phenotypic effect. Only one common QTL affecting SKC was identified in Teqing background-ILs under green house and phototron environments, indicating there was a strong interaction of ST-QTL with environment. Although the two parents are moderately susceptible to salt stress, QTL mapping results indicated that some favorable alleles beneficial to improvement of ST-related traits do exist in the parents. Therefore, it is possible to develop a new variety with improved ST by identifying and mining this kind of “hidden” ST-genes existed in the modern varieties through molecular marker technology and further pyramiding these non-allelic alleles via marker-assisted selection.

以优质粳稻品种Lemont与高产籼稻品种特青为亲本培育的高代双向回交导入系为材料，在温室140 mmol L^-1 NaCl胁迫条件下定位影响苗期叶片盐害级别(SST)、幼苗存活天数(SDS)、地上部K⁺浓度(SKC)和地上部Na⁺浓度(SNC)及人工气候室条件下影响地上部K⁺、Na⁺浓度的QTL。双向导入系的大部分遗传背景与各自的受体亲本相同，其中Lemont背景导入系中轮回亲本Lemont的基因组平均占83.8%，特青背景导入系中轮回亲本特青基因组平均占88.9%。各耐盐相关性状在两个背景群体中均出现超亲分离，多数性状的频率分布呈相互重叠状态，表明双亲作为供体相互导入各耐盐性状基因的效应大致相当。两个背景导入系群体中分别检测到影响上述耐盐相关性状的QTL各18个，同一性状在两个背景导入系中未能检测到任何相同表达的QTL，表明耐盐QTL表达具有很强的遗传背景效应，同时也说明这些耐盐QTL的效应可能较小。温室和人工气候室两种环境下仅在特青背景导入系中检测到1个影响SKC的相同QTL，表明耐盐QTL与环境的互作非常明显。虽然双亲均表现中等感盐，但QTL定位结果表明双亲中都存在一些提高耐盐相关性状的有利等位基因。研究认为，利用分子标记技术挖掘“隐蔽”于育成品种中的耐盐基因，进一步利用分子标记辅助选择技术对这些非等位耐盐基因进行聚合，完全有可能提高育成品种的耐盐水平。

2009

0.0

1.8267

Acta Agron Sin. 2009, 35(8):1386 - 1394 DOI:10.3724/SP.J.1006.2009.01386

Genetic Background Effect on QTL Expression of Heading Date and Plant Height and Their Interaction with Environment in Reciprocal Introgression Lines of Rice

利用双向导入系解析水稻抽穗期和株高QTL及其与环境互作表达的遗传背景效应

WANG Yun^1,2, CHENG Li-Rui², SUN Yong², ZHOU Zheng², ZHU Ling-Hua², XU Zheng-Jin¹, XU Jian-Long^2,*,LI Zhi-Kang^2,3

王韵^1,2,程立锐²,孙勇²,周政²,朱苓华²,徐正进¹,徐建龙^2,*,黎志康^2,3

¹ Shenyang Agricultural University / Key Laboratory of Crop Physiology, Ecology, Genetics and Breeding, Ministry of Agriculture, Shenyang 110161, China;
² Institute of Crop Sciences / National Key Facility for Crop Gene Resources and Genetic Improvement, Chinese Academy of Agricultural Sciences, Beijing 100081, China;³International Rice Research Institute, DAPO Box 7777, Metro Manila, Philippines

Expression of quantitative trait is affected by genetic background and environment. Genetic background effect on QTL mapping and QTL by environment interaction for heading date (HD) and plant height (PH) in Beijing and Hainan environments were dissected using a large set of reciprocal introgression lines (ILs) derived from ajaponica variety “Lemont” andindicavariety “Teqing”.The two sets of ILs showed transgressive segregation for the two traits. Total 16 and 17 main-effect QTLs were identified for HD and PH in the two environments, respectively. Among them, only five main-effect QTLs (QHd2,QHd8a,QPh3, QPh5,andQPh12) were detected under the two backgrounds, indicating expression of most main-effect QTLs are specific to genetic background. Three main-effect QTLs (QHd8a, QHd9, andQHd10b) by environment interactions for HD were significantly detected under the two backgrounds, of which that ofQHd8a had earlier heading for 2–3 days in Hainan, but delayed heading for 2–3 days in Beijing. Therefore,QHd8a could be considered as an important main-effect QTL for HD. By comparison with the QTL mapping results previously identified in the seven different mapping populations derived from the same parents in different environments, some stably expressed main-effect QTLs includingQHd3,QHd8a,QPh3,andQPh4 were identified under different backgrounds and environments, suggesting these QTLs could be used in marker-assisted breeding for HD and PH. On the basis of the mapping information, marker-assisted improvement of HD for a rice variety under different environments was deeply discussed.

利用粳稻Lemont和籼稻特青相互导入构建的遗传背景基本一致的双向回交导入系群体，分别在北京和海南环境定位影响抽穗期和株高的主效QTL及其环境互作，分析QTL及其与环境互作表达的遗传背景效应。在北京和海南分别检测到影响抽穗期和株高的主效QTL 16个和17个，其中有5个主效QTL (QHd2、QHd8a、QPh3、QPh5和QPh12)在两种背景下同时被检测到，表明多数主效QTL的表达具有遗传背景特异性。两种背景下检测到影响抽穗期的3个主效QTL (QHd8a、QHd9和QHd10b)存在环境互作，其中QHd8a与海南环境的互作在两种背景下提早抽穗2~3 d，与北京环境的互作则延迟抽穗2~3 d，是影响抽穗期的一个重要主效QTL。通过与以往相同亲本来源的7个不同定位群体在不同环境下定位结果的比较，鉴定出一些在不同遗传背景和环境下稳定表达的主效QTL，如QHd3、QHd8a、QPh3和QPh4，适宜用于水稻抽穗期和株高的分子标记改良。基于QTL定位结果，本文对如何通过分子标记辅助改良品种在不同环境下的抽穗期进行了深入探讨。

... 在2个背景同时定位到的QTL中, 虽然遗传背景影响主效QTL的表达, 但并不影响基因作用的方向^[25,27] ...

2003

6.535

0.0

... 另外, 耐亚铁毒和锌毒相关的QSh5、QSdw5、QRdw5b与水稻Zn²⁺运输相关的基因OsZIP2 (23 262 252~23 264 744)^[28]紧邻 ...

2004

6.16

0.0

... 在Lemont背景导入系检测到的与耐锌毒相关的QRdw4b与Fe²⁺运输相关的基因OsYSL2 (26 652 325~ 26 655 465)^[29]位于同一区间 ...

2010

2.929

0.0

Mol Biol Rep. 2010, 37(8):3735 - 3745 DOI:10.1007/s11033-010-0027-0

Identification of Fe-excess-induced genes in rice shoots reveals a WRKY transcription factor responsive to Fe, drought and senescence

Felipe Klein Ricachenevsky(1) Raul Antonio Sperotto(1) Paloma Koprovski Menguer(2) Janette Palma Fett(12)

1.Centro de Biotecnologia, Universidade Federal do Rio Grande do Sul, P.O. Box 15005, Porto Alegre, RS 91501-970, Brazil
2.Departamento de Botanica, Universidade Federal do Rio Grande do Sul, P.O. Box 15005, Porto Alegre, RS 91501-970, Brazil

Fe participates in several important reactions in plant metabolism. However, Fe homeostasis in plants is not completely understood, and molecular studies on Fe-excess stress are scarce. Rice (Oryza sativa L. ssp.indica) is largely cultivated in submerged conditions, where the extremely reductive environment can lead to severe Fe overload. In this work, we used representational difference analysis (RDA) to isolate sequences up-regulated in rice shoots after exposure to Fe-excess. We isolated 24 sequences which have putative functions in distinct cellular processes, such as transcription regulation (OsWRKY80), stress response (OsGAP1,DEAD-BOX RNA helicase), proteolysis (oryzain-α,rhomboid protein), photosynthesis (chlorophyll a/b binding protein), sugar metabolism (βglucosidase) and electron transport (NADH ubiquinone oxireductase). We show that the putative WRKY transcription factorOsWRKY80 is up-regulated in rice leaves, stems and roots after Fe-excess treatment. This up-regulation is also observed after dark-induced senescence and drought stress, indicating thatOsWRKY80 could be a general stress-responsive gene. To our knowledge, this is the first report of an Fe-excess-induced transcription factor in plants.

... 在特青背景导入系检测到的影响铁毒的QSh9和QSdw9与Fe²⁺运输相关的转录因子OsWRKY80 (24 734 518~ 24 734 438)^[30]相邻 ...

2012

1.641

0.0

... 通过不同评价体系定位到的QTL均有不同之处^[31], 而且定位结果也难于相互比较, 因此, 建立高效的水稻苗期耐亚铁毒和锌毒胁迫的评价体系显得十分重要 ...