水稻株型相关性状的QTL分析

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张玲, 李晓楠, 王伟, 杨生龙, 李清, 王嘉宇. 水稻株型相关性状的QTL分析. 作物学报, 40(12): 2128-2135
ZHANG Ling, LI Xiao-Nan, WANG Wei, YANG Sheng-Long, LI Qing, WANG Jia-Yu. Analysis of QTLs for Plant Type Traits in Rice (Oryza sativa) . Acta Agronomica Sinica, 40(12): 2128-2135 复制到剪切板

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作物学报编辑部

水稻株型相关性状的QTL分析

张玲^*, 李晓楠^*, 王伟, 杨生龙, 李清, 王嘉宇^*

沈阳农业大学水稻研究所 / 农业部东北水稻生物学与遗传育种重点实验室, 辽宁沈阳 110866

^* 通讯作者(Corresponding author): 王嘉宇, E-mail:ricewjy@126.com; Tel: 024-88487184

收稿日期:2014-09-16

基金:本研究由辽宁省高等学校优秀人才支持计划项目(LJQ2013075)资助;

摘要

以南方籼型杂交稻恢复系泸恢99和北方粳型超级稻沈农265杂交衍生的重组自交系群体(recombinant inbred lines, RILs)为试验材料, 对株型性状(株高、穗长、分蘖和叶片性状)进行不同环境下的数量性状基因位点(quantitative trait locus, QTL)分析。共检测到39个相关QTL, 分布在水稻第1、第2、第3、第6、第7、第8和第9染色体上, LOD值介于2.50~16.90之间, 有11个QTL能在两年中被检测到。株型相关的QTL在染色体上成簇分布, 主要分布于第1、第6和第9染色体上, 这可能与株型性状间显著或极显著相关有关。其中, 在第9染色体上RM3700B-RM7424区间存在1个QTL簇, 含4个QTL, 即qPH9、qPL9、qFLL9和qSLL9, 这4个QTL在2年中均被检测到。此外, 进一步鉴定出5个能稳定表达的QTL, 其中,qPH8、qFLW6和qSLW6效应较大。这些信息综合反映了株型相关性状遗传的复杂性, 有助于更全面地了解和掌握株型性状的遗传基础。

关键词: 水稻:; 株型性状:; 不同环境:; QTL分析

Analysis of QTLs for Plant Type Traits in Rice (Oryza sativa)

ZHANG Ling^**, LI Xiao-Nan^**, WANG Wei, YANG Sheng-Long, LI Qing, WANG Jia-Yu^*

Rice Research Institute, Shenyang Agricultural University / Key Laboratory of Northeast Rice Biology and Breeding, Ministry of Agriculture, Shenyang 110866, China

Abstract

A recombinant inbred lines (RILs) derived from the across betweenindica restorer line Luhui 99 and superjaponica cultivar Shennong 265, was used to analyze the quantitative trait loci (QTLs) for plant height, panicle length, tillers, and leaf traits in 2012 and 2013. A total of 39 QTLs were detected on chromosomes 1, 2, 3, 6, 7, 8, and 9 with LOD score ranging from 2.50 to 16.90. Eleven of them were detected in both 2012 and 2013. Moreover, QTL clusters were detected on chromosomes 1, 6, and 9, which may be related to significant or highly significant correlations between plant type traits. Among them, the QTL cluster on chromosome 9 contained four QTLs,qPH9,qPL9,qFLL9, andqSLL9in the interval between RM3700 and RM7424, and the four QTLs were detected in both years. In addition, five major QTLs were first reported, among which three QTLs (qPH8,qFLW6, andqSLW6) had the larger effect. The results facilitate further understanding of the genetic basis for plant height, panicle length, tillers and leaf traits.

Keyword: Oryza sativa L.; Plant type traits; Different environments; QTL analysis

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水稻是重要的粮食作物之一, 提高产量是水稻遗传育种学家追求的永恒目标。对水稻高产育种而言, 其基本原理有两条, 即形态改良和机能改进^{[ 1]}。前者主要通过塑造适应本地区的理想株型, 进而构建高光效群体和提高群体物质生产能力, 后者则是通过利用杂种优势改善个体的生理机能, 提高个体生产能力。水稻育种历史也表明, 株型改良对水稻产量潜力的提高发挥了重要作用。无论20世纪60年代的矮化育种, 还是在此基础上兴起的理想株型育种, 核心问题都是株型^{[ 2]}。可见, 良好的株型是水稻新品种选育的重要目标之一, 株型性状的遗传研究对水稻育种的理论、方法和策略都至关重要。

迄今为止, 关于株型的遗传规律研究较多, 一般认为株型性状为多基因控制的数量性状, 易受环境的影响。近年来, 人们加强了对水稻株型的遗传基础研究, 尤其是水稻株型的分子调控机制研究取得了较大进展, 一些株型性状相关基因相继被鉴定和功能注释。水稻半矮秆材料的发现和利用推动了水稻绿色革命, 后来的研究发现, 半矮秆基因 sd1是由3个外显子和2个内含子组成的开放阅读框, 编码由389个氨基酸组成的GA20氧化酶(GA20ox), 参与赤霉素的生物合成^{[ 3]}。Jiao等^{[ 4]}和Miura等^{[ 5]}克隆了理想株型基因 IPA1, 该基因编码含有SBP结构域的转录因子OsSPL14, 在营养生长时期的茎尖和生殖生长时期的枝梗原基中表达量最高, IPA1可以使水稻分蘖数减少, 穗粒数和千粒重增加, 茎秆变得粗壮, 从而有助于水稻产量的提高。 MOC1基因是控制水稻分蘖数量的相关基因, 编码GRAS家族蛋白, 通过决定腋生分生组织起始和生长而控制分蘖的数量^{[ 6]}。Qi等^{[ 7]}深入研究了矮秆窄叶突变体 nal1, NAL1基因编码一个植物特有的未知功能蛋白, 该基因通过改变生长素极性运输和分布来调控水稻的维管排列, 进而影响叶片的生长发育。多蘖矮秆基因 HTD2/ D88/ D14编码一个酯酶, 能抑制水稻分枝发生, 负调节水稻分蘖数^{[ 8, 9, 10]}。矮化多分蘖突变体 d53是一个独脚金内酯不敏感突变体, 对该基因的功能分析表明, D53蛋白作为独脚金内酯信号途径的抑制子参与调控水稻分蘖的生长发育^{[ 11, 12]}。这些基因的鉴定不仅为水稻株型改良提供了重要的理论基础, 而且为水稻籼粳交杂种优势利用提供了有用的基因和材料。尽管人们对水稻的株型性状进行了很多分析, 定位到一些控制株型的QTL/基因, 但多数分析涉及的性状较少, 分析的不够透彻。因此, 有必要对株型相关性状进行更为深入细致的分析。本研究以重组自交系群体为材料, 对9个株型性状进行QTL分析, 同时与已报道的研究结果进行比较, 进一步加深对水稻株型性状遗传多样性的理解, 也为利用分子标记辅助聚合水稻株型性状基因提供理论依据。

1 材料与方法

1.1 试验材料

沈农265是北方粳型超高产品种, 其株型紧凑, 穗短而直立, 泸恢99是籼型杂交稻重要恢复系, 其剑叶和穗长较大, 株型为叶下禾。2007年夏季在沈阳配制杂交组合“泸恢99/沈农265”, 采用单粒传法(single seed descent, SSD)获得RIL群体, 利用RIL群体中151个稳定株系作为试验材料。

1.2 田间试验

试验于2012年和2013年在沈阳农业大学水稻研究所试验田进行, 4月18日至20日育苗, 5月18日至20日移栽, 151个RIL株系, 每小区3行, 每行10株, 栽植的行株距为30.0 cm×13.3 cm, 单苗插植, 其他管理与一般大田相同。

参照《水稻种质资源描述规范和数据标准》^{[ 13]}去除边株选取5穴, 测定株高、分蘖、穗长、剑叶长、剑叶宽、倒二叶长和倒二叶宽, 计算剑叶和倒二叶的叶长宽比, 2次重复取均值。

1.3 统计分析

选择均匀分布于水稻12条染色体上的456对分子标记对泸恢99和沈农265进行差异引物筛选, 其中包括366对SSR (简单重复序列, Simple Sequence Repeat)标记、34对InDel (插入/缺失, Insertion/ Deletion)标记^{[ 14]}和56对SSILP (亚种间特异性内含子长度多态性, Subspecies-Specific Intron Length Poly--morphism)标记^{[ 15]}, 共筛选出179对具有明显差异的多态性标记构建分子标记连锁图谱, 多态性频率为39.25%。图谱覆盖了水稻基因组约1571.3 cM, 标记间的平均距离为8.77 cM, 平均每对染色体上的标记数为14.92个。利用Mapmaker/EXP3.0分析软件构建连锁图谱, 利用Kosambi函数将重组率转化为遗传图距(cM), 利用Mapchart2.2绘制连锁图^{[ 16, 17]}。采用QTL ICI Mapping v3.0的完备区间作图(ICIM)方法进行QTL分析^{[ 18]}, LOD的阈值设为2.5, 当实际求得的LOD值大于LOD阈值时, 就判定该区段存在1个QTL, 同时估算每个QTL的加性效应值和贡献率。QTL的命名遵循McCouch等^{[ 19]}的原则。

2 结果与分析

2.1 RIL群体及其亲本的株型性状表现

亲本间9个性状都存在显著差异, 与泸恢99相比沈农265具株型紧凑的直立穗, 其剑叶和倒二叶均为“短-窄”型, 株高稍矮、分蘖稍多, 在重组自交系群体中株型性状均表现出较大的超亲分布, 变异幅度较大, 表现数量性状的遗传特性( 表1), 符合QTL作图要求。

表1 株型性状在亲本和RILs群体中的分布情况 Table 1 Phenotypic performance of plant type traits between parents and the RILs population in 2012 and 2013

性状 Trait	亲本 Parent		重组自交系群体 RIL population
性状 Trait	泸恢99 Luhui 99	沈农265 Shennong 265	均值±标准差 Mean ± SD	变幅 Range	峰度 Skewness	偏度 Kurtosis
2012
株高PH (cm)	112.06±2.08	108.54±2.63^*	112.63±124.46	58.50-158.25	-0.27	-0.69
穗长PL (cm)	33.15±2.18	19.38±2.74^**	23.36±3.86	14.51-33.80	0.19	-0.13
每穴分蘖数TN	9.65±0.58	11.45±0.85^*	12.26±2.90	6.63-22.00	0.74	0.56
剑叶长FLL (cm)	32.63±2.10	24.36±3.35^**	28.11±6.25	17.03-46.11	0.46	-0.36
剑叶宽FLW (cm)	2.28±0.21	1.79±0.15^**	1.99±0.28	1.40-2.95	0.13	-0.18
剑叶长宽比FLWR	14.31±2.32	13.60±1.29^**	14.41±3.72	7.24-24.27	0.51	-0.36
倒二叶长SLL (cm)	53.52±2.02	37.93±2.95^**	42.29±6.64	27.38-60.28	0.20	-0.08
倒二叶宽SLW (cm)	1.96±0.05	1.82±0.06^**	1.73±0.34	0.95-4.09	2.03	1.70
倒二叶长宽比SLWR	27.31±1.45	20.84±1.18^**	24.39±5.77	12.76-46.45	0.64	0.02
2013
株高PH (cm)	109.00±2.73	105.25±2.28^*	119.97±25.97	63.90-175.30	-0.10	-0.90
穗长PL (cm)	34.22±3.03	17.78±2.38^**	23.62±3.60	11.26-37.29	0.17	0.96
每穴分蘖数TN	8.75±1.18	10.25±0.74^*	11.17±2.27	7.10-19.40	1.00	1.45
剑叶长FLL (cm)	34.63±2.40	21.38±2.90^**	30.52±6.91	16.44-51.78	0.48	-0.15
剑叶宽FLW (cm)	2.30±0.22	1.88±0.13^**	1.96±0.31	0.95-2.67	-0.24	-0.01
剑叶长宽比FLWR	15.06±1.67	11.37±1.12^**	15.98±4.49	8.28-30.28	0.90	1.40
倒二叶长SLL (cm)	54.50±3.48	38.13±2.56^**	44.59±7.13	29.90-63.94	0.21	-0.34
倒二叶宽SLW (cm)	1.98±0.08	1.75±0.05^**	1.78±0.26	1.10-2.54	0.05	-0.40
倒二叶长宽比SLWR	27.53±2.45	21.79±1.03^**	26.79±7.23	7.36-46.28	0.59	0.20

^*,^** denote significance of variance at 0.05 and 0.01 probability levels, respectively. PH: plant height; PL: panicle length; TN: tiller number per hill; FLL: flag leaf length; FLW: flag leaf width; FLWR: flag leaf length width rate; SLL: the second leaf length; SLW: the second leaf width; SLWR: the second leaf length to width ratio.

^*,^**分别表示方差在0.05和0.01水平上显著。

表1 株型性状在亲本和RILs群体中的分布情况 Table 1 Phenotypic performance of plant type traits between parents and the RILs population in 2012 and 2013

2.2 株型性状间的相关性

相关分析表明( 表2), 株高与穗长、剑叶长、倒二叶长均呈极显著正相关, 与每穴分蘖数呈极显著负相关, 与剑叶宽的相关性不显著: 穗长与每穴分蘖数呈极显著负相关, 与剑叶长、倒二叶长均呈极显著正相关, 与剑叶宽的相关性不显著: 每穴分蘖数与剑叶长、剑叶宽、倒二叶长均存在极显著的负相关, 但相关系数均相对较小, 与倒二叶宽的相关性不显著: 剑叶长和倒二叶长存在极显著的正相关, 与剑叶宽相关性不显著: 剑叶宽与倒二叶宽呈极显著正相关, 与倒二叶长相关性不显著。此外, 倒二叶宽年际间各相关系数变化较大, 说明其易受环境条件的影响。

两年间各指标呈现极显著相关, 但每穴分蘖数的相关系数相对较小, 表明其可能易受环境影响。株高和剑叶长的相关系数相对较大, 其受环境因素影响很小。

表2 株型性状的相关系数 Table 2 Correlation coefficient between observed traits in the RIL population

2.3 株型性状QTL定位

2.3.1 2012年株型性状QTL分析 2012年检测到19个QTL, 分布在第1、第2、第3、第6、第8和第9染色体上( 表3和图1)。检测到控制株高的QTL 1个, 位于第1染色体上, 增效等位基因来自泸恢99, 贡献率为44.25%。检测到控制穗长的QTL 2个, 分别位于第8和第9染色体, 其中 qPL8增效等位基因来自沈农265, qPL9增效等位基因来自泸恢99, 贡献率分别为6.58%和48.07%。检测到1个与每穴分蘖数相关的QTL qTN2, 贡献率为30.38%, 其增效等位基因来自泸恢99。控制剑叶长度的位点有2个, 位于第8和第9染色体, qFLL9增效等位基因来自泸恢99, 贡献率为44.50%, qFLL8的增效等位基因来自沈农265, 贡献率为12.53%。控制剑叶宽度的位点有2个, 分别位于第1和第6染色体, 增效等位基因都来自于泸恢99, 贡献率分别为11.73%和9.75%: 共检测到2个控制剑叶长宽比的QTL qFLWR1和 qFLWR9, 其中 qFLWR9贡献率最大, 表型贡献率达47.48%, 其增效基因来自泸恢99, qFLWR6增效基因来自沈农265, 贡献率均相对较小。

控制倒二叶长度的位点有4个, 位于第1、第3和第9染色体, 增效等位基因来自沈农265的有 qSFL1和 qSFL3c, 贡献率分别为5.36%和7.86%, qSFL3a和 qSFL9的增效等位基因来自泸恢99, 贡献率为7.53%和36.09%。控制倒二叶宽度的位点有3个, 位于第1、第6和第9染色体, 增效等位基因来自泸恢99的有 qSFW1和 qSFW6, 贡献率分别为7.99%和14.66%, 增效等位基因来自沈农265只有 qSFW9, 贡献率为9.44%。检测到2个控制倒二叶长宽比的QTL qSLWR1和 qSLWR9, 其中 qSLWR9贡献率较大, 其增效基因均来自泸恢99, qSLWR1贡献率均相对较小, 其表型贡献率仅4.12%, 增效基因来自沈农265。

2.3.2 2013年株型性状QTL分析 2013年检测到20个QTL ( 表3和图1), 分布在第1、第2、第6、第7、第8和第9染色体上。检测到控制株高的QTL 4个, 分布在第1、第8和第9染色体上, 其中 qPH1a的效应值最大, qPH1b的效应值最小, 除 qPH9外增效等位基因均来自泸恢99。检测到1个控制穗长的QTL qPL9, 其LOD值、表型贡献率、加性效应值分别为9.19、30.99%和2.03, 其增效等位基因来自泸恢99。控制剑叶长度的位点有2个, 位于第8和第9染色体, LOD值分别为3.91和10.71, 贡献率分别为9.92%和32.03%, 加性效应值分别为 -2.17和3.96, 其中 qFLL9增效等位基因来自泸恢99, qFLL8的增效等位基因来自沈农265。控制剑叶宽度的位点有4个, 位于第2、第6、第7和第9染色体, 增效等位基因来自泸恢99的有 qFLW2和 qFLW6, 贡献率分别为10.69%和12.50%, 增效等位基因来自沈农265的有 qFLW7和 qFLW9, 贡献率分别为9.91%和7.49%。共检测到2个控制剑叶长宽比的QTL, 其中 qFLWR9贡献率较大, 表型贡献率达33.87%, 其增效基因来自泸恢99, qFLWR6贡献率为10.58%, 增效基因来自沈农265。

控制倒二叶长度的位点有3个, 位于第2、第8和第9染色体, 增效等位基因来自沈农265的有 qSFL8, 贡献率分别为9.98%, 来自泸恢99的有 qSFL2和 qSFL9, 贡献率为8.60%和26.83%。控制倒二叶宽度的位点有2个, 位于第6和第9染色体, qSFW6增效等位基因来自泸恢, 贡献率仅为6.01%, qSFW9增效等位基因来自沈农265, 贡献率为11.09%。检测到2个控制倒二叶长宽比的QTL qSLWR1和 qSLWR9, 其中 qSLWR9贡献率较大, 能解释表型变异的32.96%, 增效基因来自泸恢99。未检测到与每穴分蘖数相关的QTL, 可能是每穴分蘖数受环境因素影响较大所致。

2.3.3 两年株型性状QTL比较分析两年共检测到39个株型性状相关的QTL, 2012年和2013年分别为19个和20个 QTL, 分布于第1、第2、第3、第6、第7、第8和第9染色体, LOD值介于2.50~16.90之间, 单个QTL贡献率在5.36%~46.16%范围内。其中, 有11个QTL在2年间均能被检测到, 包括1个株高的QTL ( qPH1a), 1个穗长QTL ( qPL9)、2个剑叶长的QTL ( qFLL8和 qFLL9), 1个剑叶宽的QTL ( qFLW6),1个剑叶长宽比QTL( qFLWR9), 1个倒二叶长的QTL ( qSFL9b), 2个倒二叶宽的QTL ( qSFW6和 qSFW9)以及2个倒二叶长宽比QTL ( qSLWR1和 qSLWR9)。就这些QTL分布的染色体区间而言, 控制株型性状的QTL分别在第1、第6和第9染色体上形成QTL簇。其中, 在第1染色体RM1361-RM6321和RM580-RM259区域及附近检测到控制多个株型性状的QTL, 在第6染色体PSM389-RI04969区域检测到能调控剑叶宽、倒二叶宽和剑叶长宽比的QTL, 在第9染色体RM3700B-RM7424A区域检测到同时调控株高、穗长、剑叶长、剑叶宽、倒二叶长和倒二叶宽的QTL, 而其余位点多效性不明显。

表3 2年间9个株型性状的QTL定位结果 Table 3 Identification of QTLs for nine plant type traits in RILs population and their genetic parameters estimated in 2012 and 2013

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	图1 株型性状QTL在染色体上的分布Fig. 1 Location of the QTLs for plant type traits detected in 2012 and 2013 in the genetic map

3 讨论

株型改良是水稻增产的重要途径之一, 通过合理组配调控株高、分蘖和剑叶等株型性状可以提升冠层的光合利用率进而增加单产。株型性状主要包括株高、穗型、分蘖、叶片和根系等性状, 与光合作用和水肥吸收的主要部位相关联, 显著影响产量和品质。株高是与产量相关的重要农艺性状之一, 研究表明株高, 叶长与穗重、千粒重、一次、二次枝梗数等相关性状显著正相关, 但并不是越高(长)越好, 株高过高, 重心也会升高, 易倒伏, 导致产量和品质降低。因此, 适当增加株高和培育壮秆抗倒伏的新品种是超高产育种的主要目标。李仕贵等^{[ 20]}以籼粳交DH群体对剑叶性状进行定位分析, 共检测出13个QTL, 包括控制剑叶长的QTL 4个, 叶宽的QTL 5个, 影响剑叶长宽比的QTL 2个; 岳兵等^{[ 21]}以重组自交系为材料, 分别检测到控制剑叶长、宽和叶面积的QTL 5、3和5个。徐建军等^{[ 22]}利用染色体片段代换系群体定位剑叶形态QTL, 共检测到14个相关QTL, 贡献率介于4.06%~60.40%; 王一平等^{[ 23]}以汕优63重组自交系群体为材料, 对顶部3叶的叶片性状进行QTL分析, 共检测到28个QTL, 同时发现穗重与9个叶片性状存在极显著的正相关。彭茂民等^{[ 24]}认为控制水稻剑叶形态性状的QTL在不同遗传背景下的数量、分布、QTL位置、贡献率和加性效应值都有比较明显的差异。本研究2年共检测到39个QTL, 其中17个在单一年份中被检测到, 11个在2年均能被检测到, 分别占当年定位QTL总数的57.9%和55.0%, 重复性相对较好。根据国家水稻数据中心的基因数据库(http://www. ricedata.cn/gene/)和水稻QTL数据库(http://qtaro. abr.affrc.go.jp/qtab/table)公布的信息, 结合本研究中检测到的12个主效QTL, 对株型相关基因/QTL的图谱整合( 图1)发现, 在第1染色体长臂和第9染色体相应的区域均有相关QTL被报道, 这些基因/QTL可能相同, 也可能是新的基因或等位基因。特别是位于第6染色体控制叶宽的 qFLW6和 qSLW6与已定位的 qPL6和 qPH6位于相近区域, 但还未见克隆的报道。本研究中定位到的 qSLWR1和 qFLL8未见有文献报道, 是2个新的QTL位点。

QTL的多效性指的是同一区间的QTL同时对多个性状产生作用, 这种情况在作物中普遍存在。就泸恢99/沈农265的RILs群体而言, 控制株型性状的QTL主要分布在第1、第6和第9染色体上, 且成簇分布。本研究中效应值较大的QTL在不同的环境中表达比较稳定, 其中第6染色体的 qFLW6和 qSLW6, 第6染色体长臂末端的 qFLW6和 qSLW6与姜树坤等^{[ 25]}检测到的 qFLW6、 qFLL6、 qSLW6a和 qSLW6b, 刘进等^{[ 26]}在第6染色体检测到的 qFLW6和 qSLW6在相近的位置; 第9染色体检测到的 qPH9、 qPL9、 qFLL9和 qSFL9b与姜树坤等^{[ 25]}、Hittalmani等^{[ 27]}、方萍等^{[ 28]}和杜景红等^{[ 29]}在第9染色体上定位到株高、穗长和剑叶长QTL位点相近, 其贡献率和加性效应均相对较大, 能在不同环境中被检测到。此外, 在该区间内存在已被克隆的穗型基因 qPE9-1/ EP/ DEP1^{[ 30, 31, 32]}, 该基因调控株高、穗长和着粒密度等多个性状。当然, 本研究所检测到的QTL是否为 DEP1基因的多效性造成, 还需要进一步精细定位分析来确定。

4 结论

利用泸恢99/沈农265的RIL群体构建了覆盖水稻基因组1571.3 cM的遗传图谱, 标记间平均距离为8.77 cM。对株高、穗长、分蘖、剑叶长、剑叶宽、剑叶长宽比、倒二叶长、倒二叶宽和倒二叶长宽比等9个株型性状进行QTL定位分析, 2年共检测到39个QTL, 包括2012年的19个, 2013年的20个。2年重复检测到的QTL有11个, 占总数的28.21%。部分株型性状的QTL在染色体上成簇分布, 主要分布在第1、第6和第9染色体上, 这可能与株型性状间存在显著或极显著相关有关。特别是位于第9染色体RM3700B-RM7424区间的1个QTL簇包括4个QTL( qPH9、 qPL9、 qFLL9和 qSLL9), 这4个QTL在两年中均被检测到, 且效应较大, 对这些QTL可作进一步深入研究。

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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2007

0.0

... 对水稻高产育种而言, 其基本原理有两条, 即形态改良和机能改进^[1] ...

2012

0.0

0.55

... 无论20世纪60年代的矮化育种, 还是在此基础上兴起的理想株型育种, 核心问题都是株型^[2] ...

2002

38.597

0.0

Sasaki

, Ashikari

, Ueguchi-Tanaka

, Itoh

, Nishimura

, Swapan

, Ishiyama

, Saito

, Kobayashi

, Khush

G S

, Kitano

, Matsuoka

Green revolution: a mutant gibberellin- synthesis gene in rice. Nature, 2002, 416: 701-702

Nature is the international weekly journal of science: a magazine style journal that publishes full-length research papers in all disciplines of science, as well as News and Views, reviews, news, features, commentaries, web focuses and more, covering all branches of science and how science impacts upon all aspects of society and life.

... 水稻半矮秆材料的发现和利用推动了水稻绿色革命, 后来的研究发现, 半矮秆基因sd1是由3个外显子和2个内含子组成的开放阅读框, 编码由389个氨基酸组成的GA20氧化酶(GA20ox), 参与赤霉素的生物合成^[3] ...

2010

35.209

0.0

JiaoY

, Wang

Y H

, Xue

D W

, Wang

, Yan

M X

, Liu

G F

, Dong

G J

, Zeng

D L

, Lu

Z F

, Zhu

X D

, Qian

, Li

J Y

. Regulation of OsSPL14 by OsmiR156 defines ideal plant architecture in rice. Nat Genet, 2010, 42: 541-544

Increasing crop yield is a major challenge for modern agriculture. The development of new plant types, which is known as ideal plant architecture (IPA), has been proposed as a means to enhance rice yield potential over that of existing high-yield varieties^{1, 2}. Here, we report the cloning and characterization of a semidominant quantitative trait locus, IPA1 (Ideal Plant Architecture 1), which profoundly changes rice plant architecture and substantially enhances rice grain yield. The IPA1 quantitative trait locus encodes OsSPL14 (SOUAMOSA PROMOTER BINDING PROTEIN-LIKE 14) and is regulated by microRNA (miRNA) OsmiR156 in vivo. We demonstrate that a point mutation in OsSPL14 perturbs OsmiR156-directed regulation of OsSPL14, generating an 'ideal' rice plant with a reduced tiller number, increased lodging resistance and enhanced grain yield. Our study suggests that OsSPL14 may help improve rice grain yield by facilitating the breeding of new elite rice varieties.

... Jiao等^[4]和Miura等^[5]克隆了理想株型基因IPA1, 该基因编码含有SBP结构域的转录因子OsSPL14, 在营养生长时期的茎尖和生殖生长时期的枝梗原基中表达量最高, IPA1可以使水稻分蘖数减少, 穗粒数和千粒重增加, 茎秆变得粗壮, 从而有助于水稻产量的提高 ...

2010

35.209

0.0

Kotaro

, Mayuko

, Atsushi

, Song

X J

, Midori

, Kenji

, Makoto

, Hidemi

, Motoyuki

OsSPL14 promotes panicle branching and higher grain productivity in rice. Nat Genet, 2010, 42: 545-549

Identification of alleles that improve crop production and lead to higher-yielding varieties are needed for food security. Here we show that the quantitative trait locus WFP (WEALTHY FARMER'S PANICLE) encodes OsSPL14 (SQUAMOSA PROMOTER BINDING PROTEIN-LIKE 14, also known as IPA1). Higher expression of OsSPL14 in the reproductive stage promotes panicle branching and higher grain yield in rice. OsSPL14 controls shoot branching in the vegetative stage and is affected by microRNA excision. We also demonstrate the feasibility of using the OsSLP14^WFP allele to increase rice crop yield. Introduction of the high-yielding OsSPL14^WFP allele into the standard rice variety Nipponbare resulted in increased rice production.

2003

38.597

0.0

X Y

, Qian

, Fu

Z M

, Wang

Y H

, Xiong

G S

, Zeng

D L

, Wang

X Q

, Liu

X F

, Teng

, Hiroshi

, Yuan

, Luo

, Han

, Li

J Y.

Control of tillering in rice. Nature, 2003, 422: 618-621

... MOC1基因是控制水稻分蘖数量的相关基因, 编码GRAS家族蛋白, 通过决定腋生分生组织起始和生长而控制分蘖的数量^[6] ...

2008

6.555

0.0

... Qi等^[7]深入研究了矮秆窄叶突变体nal1, NAL1基因编码一个植物特有的未知功能蛋白, 该基因通过改变生长素极性运输和分布来调控水稻的维管排列, 进而影响叶片的生长发育 ...

2009

3.518

0.0

Gao

Z Y

, Qian

, Liu

X H

, Yan

M X

, Feng

, Dong

G J

, Liu

, Han

Dwarf 88, a novel putative esterase gene affecting architecture of rice plant. Plant Mol Biol, 2009, 71: 265-276

Rice architecture is an important agronomic trait that affects grain yield. We characterized a tillering dwarf mutant d88 derived from Oryza sativa ssp. japonica cultivar Lansheng treated with EMS. The mutant had excessive shorter tillers and smaller panicles and seeds compared to the wild-type. A reduction in number and size of parenchyma cells around stem marrow cavity as well as a delay in the elongation of parenchyma cells caused slender tillers and dwarfism in the d88 mutant. The D88 gene was isolated via map-based cloning and identified to encode a putative esterase. The gene was expressed in most rice organs, with especially high levels in the vascular tissues. The mutant carried a nucleotide substitution in the first exon of the gene that led to the substitution of arginine for glycine, which presumably disrupted the functionally conserved N-myristoylation domain of the protein. The function of the gene was confirmed by complementation test and antisense analysis. D88, thus, represents a new category of genes that regulates cell growth and organ development and consequently plant architecture. The potential relationship between the tiller formation associated genes and D88 is discussed and future identification of the substrate for D88 may lead to the characterization of new pathways regulating plant development.

1.National Center for Gene Research/Institute of Plant Physiology and Ecology, Shanghai Institutes for Biological Sciences, Chinese Academy of Sciences, 500 Caobao Road, 200233 Shanghai, China 2.State Key Laboratory of Rice Biology, China National Rice Research Institute, Chinese Academy of Agricultural Sciences, 310006 Hangzhou, China

2009

4.134

0.0

2009

3.347

0.0

Liu

W Z

, Wu

, Fu

Y P

, Hu

G C

, Si

H M

, Zhu

, Luan

W J

, He

Z Q

, Sun

Z X.

Identification and characterization of HTD2: a novel gene negatively regulating tiller bud outgrowth in rice. Planta, 2009, 230: 649-658

Tiller number is highly regulated by controlling the formation of tiller bud and its subsequent outgrowth in response to endogenous and environmental signals. Here, we identified a rice mutant htd2 from one of the 15,000 transgenic rice lines, which is characterized by a high tillering and dwarf phenotype. Phenotypic analysis of the mutant showed that the mutation did not affect formation of tiller bud, but promoted the subsequent outgrowth of tiller bud. To isolate the htd2 gene, a map-based cloning strategy was employed and 17 new insertions-deletions (InDels) markers were developed. A high-resolution physical map of the chromosomal region around the htd2 gene was made using the F₂ and F₃ population. Finally, the gene was mapped in 12.8 kb region between marker HT41 and marker HT52 within the BAC clone OSJNBa0009J13. Cloning and sequencing of the target region from the mutant showed that the T-DNA insertion caused a 463 bp deletion between the promoter and first exon of an esterase/lipase/thioesterase family gene in the 12.8 kb region. Furthermore, transgenic rice with reduced expression level of the gene exhibited an enhanced tillering and dwarf phenotype. Accordingly, the esterase/lipase/thioesterase family gene (TIGR locus Os03g10620) was identified as the HTD2 gene. HTD2 transcripts were expressed mainly in leaf. Loss of function of HTD2 resulted in a significantly increased expression of HTD1, D10 and D3, which were involved in the strigolactone biosynthetic pathway. The results suggest that the HTD2 gene could negatively regulate tiller bud outgrowth by the strigolactone pathway.

1.China National Rice Research Institute State Key Laboratory of Rice Biology 359 Tiyuchang Road 310006 Hangzhou Zhejiang China 2.China Three Gorges University Biotechnology Research Center 443002 Yichang Hubei China

2013

38.597

0.0

Zhou

, Lin

Q B

, Zhu

L H

, Ren

Y L

, Zhou

K N

, Nitzan

, Wu

F Q

, Mao

H B

, Dong

, Gan

, Ma

W W

, Gao

, Chen

, Yang

, Wang

, Tan

J J

, Zhang

, Guo

X P

, Wang

J L

, Jiang

, Liu

, Chen

W Q

, Chu

J F

, Yan

C Y

, Kotomi

, Shinsaku

, Tadao

, Cheng

Z J

, Wang

, Lei

C L

, Zhai

H Q

, Wu

C Y

, Wang

H Y

, Zheng

, Wan

J M.

D14-SCFD3-dependent degradation of D53 regulates strigolactone signaling. Nature, 2013, 504: 406-410

Strigolactones (SLs), a newly discovered class of carotenoid-derived phytohormones, are essential for developmental processes that shape plant architecture and interactions with parasitic weeds and symbiotic arbuscular mycorrhizal fungi. Despite the rapid progress in elucidating the SL biosynthetic pathway, the perception and signalling mechanisms of SL remain poorly understood. Here we show that DWARF 53 (D53) acts as a repressor of SL signalling and that SLs induce its degradation. We find that the rice (Oryza sativa) d53 mutant, which produces an exaggerated number of tillers compared to wild-type plants, is caused by a gain-of-function mutation and is insensitive to exogenous SL treatment. The D53 gene product shares predicted features with the class I Clp ATPase proteins and can form a complex with the α/β hydrolase protein DWARF 14 (D14) and the F-box protein DWARF 3 (D3), two previously identified signalling components potentially responsible for SL perception. We demonstrate that, in a D14- and D3-dependent manner, SLs induce D53 degradation by the proteasome and abrogate its activity in promoting axillary bud outgrowth. Our combined genetic and biochemical data reveal that D53 acts as a repressor of the SL signalling pathway, whose hormone-induced degradation represents a key molecular link between SL perception and responses.

2013

38.597

0.0

Jiang

, Liu

, Xiong

G S

, Liu

H H

, Chen

F L

, Wang

, Meng

X B

, Liu

G F

, Yu

, Yuan

Y D

, Yi

, Zhao

L H

, Ma

H L

, He

Y Z

, Wu

Z S

, Karsten

, Qian

, Eric

, Wang

Y H

, Li

J Y.

DWARF 53 acts as a repressor of strigolactone signalling in rice. Nature, 2013, 504: 401-405

Strigolactones (SLs) are a group of newly identified plant hormones that control plant shoot branching. SL signalling requires the hormone-dependent interaction of DWARF 14 (D14), a probable candidate SL receptor, with DWARF 3 (D3), an F-box component of the Skp–Cullin–F-box (SCF) E3 ubiquitin ligase complex. Here we report the characterization of a dominant SL-insensitive rice (Oryza sativa) mutant dwarf 53 (d53) and the cloning of D53, which encodes a substrate of the SCF^D3 ubiquitination complex and functions as a repressor of SL signalling. Treatments with GR24, a synthetic SL analogue, cause D53 degradation via the proteasome in a manner that requires D14 and the SCF^D3 ubiquitin ligase, whereas the dominant form of D53 is resistant to SL-mediated degradation. Moreover, D53 can interact with transcriptional co-repressors known as TOPLESS-RELATED PROTEINS. Our results suggest a model of SL signalling that involves SL-dependent degradation of the D53 repressor mediated by the D14–D3 complex.

2006

0.0

... 参照《水稻种质资源描述规范和数据标准》^[13]去除边株选取5穴, 测定株高、分蘖、穗长、剑叶长、剑叶宽、倒二叶长和倒二叶宽, 计算剑叶和倒二叶的叶长宽比, 2次重复取均值 ...

2009

0.0

0.854

B R

, Cai

X X

, Jin

Efficient indica and japonica rice identification based on the InDel molecular method: Its implication in rice breeding and evolutionary research. Prog Nat Sci, 2009, 19: 1241-1252

... 3 统计分析选择均匀分布于水稻12条染色体上的456对分子标记对泸恢99和沈农265进行差异引物筛选, 其中包括366对SSR (简单重复序列, Simple Sequence Repeat)标记、34对InDel (插入/缺失, Insertion/ Deletion)标记^[14]和56对SSILP (亚种间特异性内含子长度多态性, Subspecies-Specific Intron Length Poly--morphism)标记^[15], 共筛选出179对具有明显差异的多态性标记构建分子标记连锁图谱, 多态性频率为39 ...

2008

0.0

Zhao

X Q

, Wu

W R.

Construction of a genetic map based on ILP markers in rice. Hereditas (Beijing), 2008, 30: 225-230

Intron length polymorphism (ILP) is a new type of PCR-based molecular markers with many advantages. We had previously developed 172 ILP markers in rice using the published genome sequence data of indica cultivar 93-11 and ja-ponica cultivar Nipponbare. In order to examine the reliability and the applicability of these ILP markers to genetic map-ping, we constructed a rice genetic map consisting of 172 ILP and 13 SSR markers with a total length of 1 905.7 cM using a BC1F1 (Nipponbare/93-11//Nipponbare) population. Comparison showed that the order of the markers in this map was ex-actly the same as their physical order, verifying the feasibility and efficiency of using ILP markers for genetic mapping. We also investigated the phenomenon of marker segregation distortion and found a region of serious segregation distortion in the short arm of chromosome 6.

1. College of Agriculture and Biotechnology, Zhejiang University, Hangzhou 310029, China; 2. Jiaxing Academy of Agricultural Science, Jiaxing 314016, China

内含子长度多态性(ILP)是一种基于PCR的新型分子标记, 具有许多突出的优点。我们先前利用已公布的籼稻品种93-11和粳稻品种日本晴的基因组序列数据, 已开发了172个水稻ILP标记。为了检验这些ILP标记的可靠性及其在遗传作图中的可用性, 利用一个BC1F1(日本晴/93-11//日本晴)群体, 构建了一张含172个ILP标记座位和13个SSR标记座位的水稻遗传图谱, 总长度为1 905.7 cM。比较显示, 图谱上所有标记的顺序与其物理顺序完全一致, 证明了利用ILP标记进行遗传作图的可行性和有效性。文中还对标记偏分离现象进行了分析, 发现在第6号染色体短臂上存在一个严重偏分离的区域。

1987

3.01

0.0

Land er

E S

, Green

, Abrahamson

, Barlow

, Daly

M J

, Lincoln

S E.

Newburg

MAPMAKER: an interactive computer package for constructing primary genetic linkage maps of experimental and natural populations. Genomics, 1987, 1: 174-181

2011

0.0

2002

1.995

0.0

... 0的完备区间作图(ICIM)方法进行QTL分析^[18], LOD的阈值设为2 ...

1997

0.0

... QTL的命名遵循McCouch等^[19]的原则 ...

2000

0.0

1.667

李仕贵, 何平, 王玉平, 黎汉云, 周开达, 陈英, 朱立煌. 水稻剑叶性状的遗传分析和基因定位. 作物学报, 2000, 26: 261-265

S G

, He

, Wang

Y P

, Li

H Y

, Zhou

K D

, Chen

, Zhu

L H.

Genetic analysis and gene mapping of the leaf traits in rice (Oryza sativa L. ). Acta Agron Sin, 2000, 26: 261-265 (in Chinese with English abstract)

Based on a molecular linkage map of a doubled haploid population derived from the inter-subspecific cross between indica variety (Zhaiyeqing 8) and japonica variety (Jingxi 17). The QTL analysis was conducted for flag leaf traits, including flag leaf length, width, length/width, area and leaf roll.

Rice Research Institute of Sichuan Agricultural University, Wenjiang 611130

本研究以水稻籼粳杂交(窄叶青8号×京系17)F1经花药培养，产生的双单倍体(DH)群体和应用该群体已构建的分子图谱为基础。采用QTL区间作图法对5个剑叶性状进行定位分析，结果表明，在DH群体中，剑叶长、宽、长宽比和叶面积呈连续分布，受微效多基因控制，并且各性状均存在一定数量的超亲遗传类型。 4个性状共检测出13

... 李仕贵等^[20]以籼粳交DH群体对剑叶性状进行定位分析, 共检测出13个QTL, 包括控制剑叶长的QTL 4个, 叶宽的QTL 5个, 影响剑叶长宽比的QTL 2个 ...

2006

2.076

1.323

岳兵, 薛为亚, 罗利军, 邢永忠. 水稻剑叶部分形态生理特性QTL分析以及它们与产量、产量性状的关系. 遗传学报, 2006, 33: 824-832

Yue

, Xue

W Y

, Luo

L J

, Xing

Y Z.

QTL Analysis for flag leaf characteristics and their relationships with yield and yield traits in rice. J Genet Genomics, 2006, 33: 824-832 (in Chinese with English abstract)

Abstract

Photosynthesis of carbohydrate is the primary source of grain yield in rice (Oryza sativa L.). It is important to genetically analyze the morphological and the physiological characteristics of functional leaves, especially flag leaf, in rice improvement. In this study, a recombinant inbred population derived from a cross between an indica (O. sativa L. ssp. indica) cultivar and a japonica (O. sativa L. ssp. japonica) cultivar was employed to map quantitative traits loci (QTLs) for the morphological (i.e., leaf length, width, and area) and physiological (i.e., leaf color rating and stay-green) characteristics of flag leaf and their relationships with yield and yield traits in 2003 and 2004. A total of 17 QTLs for morphological traits (flag leaf length, width, and area), 6 QTLs for degree of greenness and 14 QTLs for stay-green-related traits (retention-degrees of greenness, relative retention of greenness, and retention of the green area) were resolved, and 10 QTLs were commonly detected in both the years. Correlation analysis revealed that flag leaf area increased grain yield by increasing spikelet number per panicle. However, the physiological traits including degree of greenness and stay-green traits were not or negatively correlated to grain yield and yield traits, which may arise from the negative relation between degree of greenness and flag leaf size and the partial sterility occurred in a fraction of the lines in this population. The region RM255-RM349 on chromosome 4 controlled the three leaf morphological traits simultaneously and explained a large part of variation, which was very useful for genetic improvement of grain yield. The region RM422-RM565 on chromosome 3 was associated with the three stay-green traits simultaneously, and the use of this region in genetic improvement of grain yield needs to be assessed by constructing near-isogenic lines.

... 岳兵等^[21]以重组自交系为材料, 分别检测到控制剑叶长、宽和叶面积的QTL 5、3和5个 ...

2011

0.0

1.494

徐建军, 赵强, 赵元凤, 朱磊, 徐辰武, 顾铭洪, 韩斌, 梁国华. 利用重测序的水稻染色体片段代换系群体定位剑叶形态QTL. 中国水稻科学, 2011, 25: 483-487

J J

, Zhao

Y F

, Zhu

, Xu

C W

, Gu

M H

, Han

, Liang

G H.

Mapping of QTLs for flag leaf shape using whole-genome re-sequenced chromosome segment substitution lines in rice. Chin J Rice Sci, 2011, 25: 483-487 (in Chinese with English abstract)

Flag leaf shape is one of the most important target traits in rice ideal plant type breeding. Identification and localization of new rice flag leaf shape QTLs would be of great significance to facilitate the genetic analysis of complex traits, and provide QTLs/genes to develop new plant type rice varieties via markerassisted selection. One hundred and twenty eight chromosome segment substitution lines with 9311 as recipient and Nipponbare as donor were genotyped by wholegenome resequencing. The pairwise correlation analysis on three flag leaf shape traits was conducted. The correlations between flag leaf length and flag leaf area, and between flag leaf width and flag leaf area were significantly positive. Correlation analysis revealed that the flag leaf length and flag leaf area have a direct positive impact on spikelet number per panicle. By combining the sequencingbased Binmap with multiple linear regression analysis, four, four and six QTLs for flag leaf length, flag leaf width and flag leaf area were mapped, respectively, with contributions from 4.08% to 60.40%. The results provided a foundation for fine mapping and subsequent cloning of the QTLs.

1 Jiangsu Key Laboratory of Crop Genetics and Physiology /Key Laboratory of Plant Function Genomics, Ministry of Education, Yangzhou University, Yangzhou 225009, China; 2 Songjiang Agricultural Technology and Popularization Center of Shanghai, Shanghai 201613, China; 3 Institute of Plant Physiology and Ecology, Chinese Academy of Sciences, Shanghai 200233, China;

剑叶形态性状（剑叶长、剑叶宽和剑叶面积）是水稻理想株型育种的重要目标性状之一。发掘新的控制水稻剑叶形态性状的基因资源，准确鉴定和定位水稻剑叶形态性状QTL，对开展水稻剑叶形态性状分子生物学研究和理想株型分子育种都具有重要意义。以通过高通量测序而准确获知代换片段位置及长度的一套用籼稻品种9311为受体、粳稻品种日本晴为供体构建的，包括128个染色体片段代换系群体为材料，对剑叶形态性状及其与主穗颖花之间的相关性进行分析，结果表明剑叶面积与剑叶长、宽呈极显著正相关，主穗颖花数与剑叶长、剑叶面积呈极显著正相关。利用多元回归分析方法，结合Binmap，共鉴定出与水稻剑叶长、宽和面积相关的QTL分别为4、4和6个，贡献率介于4.08%~60.40%。上述QTL的准确定位，为进一步精细定位及克隆相应QTL以及开展水稻剑叶形态性状分子育种奠定了基础。

... 徐建军等^[22]利用染色体片段代换系群体定位剑叶形态QTL, 共检测到14个相关QTL, 贡献率介于4 ...

2005

0.0

1.494

王一平, 曾建平, 郭龙彪, 邢永忠, 徐才国, 梅捍卫, 应存山, 罗利军. 水稻顶部三叶与穗重的关系及其QTL分析. 中国水稻科学, 2005, 19: 13-20

Wang

Y P

, Zeng

J P

, Guo

L B

, Xing

Y Z

, Xu

C G

, Mei

H W

, Ying

C S

, Luo

L J.

QTL and correlation analysis on characters of top three leaves and panicle weight in rice (Oryza sativa L. ). Chin J Rice Sci, 2005, 19: 13-20 (in Chinese with English abstract)

The correlation and QTL analysis on top three leaves and panicle weight as source-sink traits were conducted on 241 RIL population derived from an elite hybrid rice Shanyou 63. The correlations between grain weight per panicle and all the leaf traits were significantly positive. The biggest correlation coefficient was found between grain weight per panicle and the second leaf weight, followed by the flag leaf weight. The frequency of these traits was approximately normally distributed with tremendous transgressive segregation. A total of 44 main effect QTLs and 43 digenic epistatic loci related to these traits were detected. These main effect QTLs were located on 11 out of 12 chromosomes except chromosome 8, with contributions from 3.19%-26.23%. These digenic epistatic loci were located on all of the 12 chromosomes, with contributions ranging from 2.03%-8.93%. A QTL in marker interval R2510-RM211 on chromosome 2 affected both grain weight per panicle and the second leaf weight, which would be beneficial to super-high-yielding rice improvement.

摘要: 对水稻汕优63重组自交系群体顶部3张叶片的长、宽、重和单穗重等10个性状进行了相关分析和QTL定位。穗重与9个叶片性状存在极显著的正相关，其中与倒2叶重的相关系数最大，剑叶重次之。所有性状在重组自交系群体中均存在双向超亲分离，接近正态分布。共检测到44个主效QTL和43对双位点互作影响上述10个性状。主效QTL分布于水稻的除第8染色体外的其余11条染色体上，贡献率介于3.19%~26.23%；互作分布于水稻的12条染色体上，贡献率变幅为2.03%~8.93%。第2染色体的R2510-RM211标记区间同时检测到控制单穗重和倒2叶重的QTL，该QTL对超级稻株型育种具有应用价值。

... 王一平等^[23]以汕优63重组自交系群体为材料, 对顶部3叶的叶片性状进行QTL分析, 共检测到28个QTL, 同时发现穗重与9个叶片性状存在极显著的正相关 ...

2007

0.0

1.494

彭茂民, 杨国华, 张菁晶, 安保光, 李阳生. 不同遗传背景下水稻剑叶形态性状的QTL分析. 中国水稻科学, 2007, 21: 247-252

Peng

M M

, Yang

G H

, Zhang

J J

, An

B G

, Li

Y S.

QTL Analysis for flag leaf morphological traits in rice (Oryza sativa L. ) under different genetic backgrounds. Chin J Rice Sci, 2007, 21: 247-252 (in Chinese with English abstract)

Zhaiyeqing 8×Jingxi 17 doubled haploid(DH) population and Zhenshan 97×Minghui 63 recombinant inbred line(RIL) population were employed to dissect the genetic basis of the flag leaf morphological traits including length,width,the ratio of length to width,area,and perimeter by using composite interval mapping approaches.The traits were normally distributed with transgressive segregation in both directions,which showed that these traits were controlled by minor polygenes.The significant positive correlation was detected between different traits.In the DH(Zhaiyeqing 8×Jingxi 17) population,eight intervals carrying quantitative trait loci(QTLs) on four chromosomes were identified for the four traits.Two QTLs affected length,perimeter and area of leaf simultaneously.In the RIL(Zhenshan 97×Minghui 63) population,16 intervals carrying QTLs on five chromosomes were identified for the five traits.Among these QTLs,there were one common QTL for length,perimeter and the ratio of length to width,two common QTLs for width and area,one common QTL for length and perimeter,and one common QTL for length,perimeter and area,respectively.These results suggested that there were significant differences in QTLs in different populations.Moreover,there were some common QTLs for the closely relative traits at the same or near chromosome regions.

以典型的籼粳交窄叶青8号/京系17的加倍单倍体（DH）群体和籼籼交珍汕97/明恢63的重组自交系（RIL）群体为材料，并应用这两种群体已构建的分子图谱，对剑叶形态性状中的叶面积、周长、叶长、叶宽、长宽比等进行了QTL分析。在这两个群体中，剑叶的这些形态性状均呈连续性分布，存在一定数量的超亲分离。在DH群体中， 4个剑叶性状共检测到8个QTL，分布在4条染色体上，其中第4、8染色体上各有1个QTL同时影响了叶长、叶周长和面积。在RIL群体中，5个剑叶性状共检测到16个QTL，分布在5条染色体上。其中第1染色体上有1个QTL同时影响叶长、叶周长、长宽比3个性状，第6染色体上有2个影响叶面积的QTL同时也影响叶宽，还有1个QTL同时影响叶长、叶周长2个性状，第11染色体上有1个QTL同时影响叶长、叶周长、叶面积3个性状。研究表明，不同群体的QTL初级定位结果存在较大差异，影响叶片相关性状的一些QTL位于同一染色体的相同或者相邻区域上。

... 彭茂民等^[24]认为控制水稻剑叶形态性状的QTL在不同遗传背景下的数量、分布、QTL位置、贡献率和加性效应值都有比较明显的差异 ...

2010

0.0

1.494

姜树坤, 张喜娟, 黄成, 邢亚南, 郑旭, 徐正进, 陈温福. 基于粳稻F₂和F_{2: 6}群体的连锁图谱及剑叶性状QTL比较分析. 中国水稻科学, 2010, 24: 372-378

Jiang

S K

, Zhang

X J

, Huang

, Xing

Y N

, Zheng

, Xu

Z J

, Chen

W F.

Comparison of genetic linkage map and QTLs controlling flag leaf traits based on F₂ and F_{2: 6} populations derived from japonica rice. Chin J Rice Sci, 2010, 24: 372-378 (in Chinese with English abstract)

A comparative analysis was performed on genetic information and QTLs controlling flag leaf traits including length, width and specific leaf weight of flag leaf between two populations (F2 and F2:6) derived from a cross between two japonica rice cultivars, Shennong 265 and Lijiangxintuanheigu(LTH). On the two genetic linkage maps most markers had the same order along chromosomes, but the genetic distance between two markers was different. Thirty and thirteen markers showed genetic distortion significantly and extremely significantly in F2 population, separately. Nineteen and eleven markers deviated toward Shennong 265 and LTH, separately. Sixtytwo and thirtyeight markers showed genetic distortion significantly and extremely significantly in F2:6 population, separately. Fortythree and nineteen markers deviated toward Shennong 265 and LTH, separately. These distortional markers formed ten segregation distortion regions (SDRs). Six of them were detected in both F2 and F2:6 populations. F2:6 population had more powerful detective ability than F2 population in QTL dectection. Seven QTLs controlling flag leaf traits including two controlling leaf length, four controlling leaf width and one controlling specific weight were detected in F2 population. Whereas seventeen QTLs for these traits (seven for leaf length, five for leaf width and five for specific leaf weight) were detected in F2:6 population. Four QTLs were detected in both populations including qFLL9 controlling flag leaf length on chromosome 9, qFLW4 controlling flag leaf width on chromosome 4, qFLW12.1 controlling flag leaf width on chromosome 12 and qSLW6 controlling specific leaf weight on chromosome 6. Among them, qSLW6 (Additive effect was 1.956 mg/cm2) for specific leaf weight has a high research and application value.

1Key Laboratory of Crop Physiology, Ecology, Genetics and Breeding, Ministry of Agriculture/Key Laboratory of Northern Japonica Rice Breeding of Liaoning Province, Shenyang Agricultural University, Shenyang 110161, China; 2Dandong Academy of Agricultural Sciences, Fengcheng 118109, China; 3Liaoning Fengmin Agricultural HighTech Co. Ltd, Shenyang 110101, China; *Corresponding author, E-mail: xuzhengjin@126.com

以粳稻沈农265和丽江新团黑谷 (LTH) 为亲本，分别构建了F2群体和F2:6群体，分析并比较了两群体的遗传信息和控制剑叶相关性状(剑叶长、宽和比叶重)的数量性状基因座。结果表明：1)多数标记在染色体上的顺序相同，但标记间距不同。F2群体中30个标记显著偏离预期的1∶2∶1孟德尔分离比例，13个标记极显著偏离预期的1∶2∶1孟德尔分离比例，其中19个偏向沈农265，11个偏向LTH。F2:6群体中62个标记显著偏离预期的1∶1孟德尔分离比例，38个标记极显著偏离预期的1∶1孟德尔分离比例，其中43个偏向沈农265，19个偏向LTH。偏分离标记共形成10个偏分离区域，其中有6个区域同时出现在两个群体中。2) F2:6群体检测QTL的能力强于F2群体。F2群体共检测到7个控制剑叶性状的QTL (2个控制剑叶长，4个控制剑叶宽，1个控制比叶重)，而F2:6群体共检测到17个控制剑叶性状的QTL (7个控制剑叶长，5个控制剑叶宽，5个控制比叶重)，其中有4个QTL在两群体中同时检测到，分别是第9染色体上控制剑叶长的qFLL9，第4染色体上控制剑叶宽的qFLW4，第12染色体上控制剑叶宽的qFLW12.1和第6染色体上控制比叶重的qSLW6。其中，控制比叶重的qSLW6 (加性效应值为1.956 mg/cm2)，极富研究与应用价值。

... 本研究中效应值较大的QTL在不同的环境中表达比较稳定, 其中第6染色体的qFLW6和qSLW6, 第6染色体长臂末端的qFLW6和qSLW6与姜树坤等^[25]检测到的qFLW6、qFLL6、qSLW6a和qSLW6b, 刘进等^[26]在第6染色体检测到的qFLW6和qSLW6在相近的位置 ...

... 第9染色体检测到的qPH9、qPL9、qFLL9和qSFL9b与姜树坤等^[25]、Hittalmani等^[27]、方萍等^[28]和杜景红等^[29]在第9染色体上定位到株高、穗长和剑叶长QTL位点相近, 其贡献率和加性效应均相对较大, 能在不同环境中被检测到 ...

2012

0.0

0.951

2002

1.643

0.0

Hittalmani

, Shashidhar

H E

, Bagali

P G

, Huang

, Sidhu

J S

, Singh

V P

, Khush

G S.

Molecular mapping of quantitative trait loci for plant growth, yield and yield related traits across three diverse locations in a doubled haploid rice population. Euphytica, 2002, 125: 207-214

A doubled haploid (DH) population of 125lines derived from IR64 × Azucena, an indica – japonica cross were grown in three different locations in India during the wet season of 1995. The parents of mapping population had diverse phenotypic values for the eleven traits observed. The DH lines exhibited considerable amount of variation for all the traits. Transgressive segregants were observed. Interval analysis with threshold LOD > 3.00 detected a total of thirty four quantitative trait loci (QTL) for eleven traits across three locations. The maximum number of twenty QTL were detected at Punjab location of North India. A total of seven QTL were identified for panicle length followed by six QTL for plant height. Eight QTL were identified on three chromosomes which were common across locations. A maximum of seven QTL were identified for panicle length with the peak LOD score of 6.01 and variance of 26.80%. The major QTL for plant height was located on Chromosome 1 with peak LOD score of 16.06 flanked by RZ730-RZ801 markers. Plant height had the maximum number of common QTL across environment at the same marker interval. One QTL was identified for grain yield per plant and four QTL for thousand grain weight. Clustering of QTL for different traits at the same marker intervals was observed for plant height, panicle exsertion, panicle number, panicle length and biomass production. This suggests that pleiotropism and or tight linkage of different traits could be the plausible reason for the congruence of several QTL. Common QTL identified across locations and environment provide an excellent opportunity for selecting stable chromosomal regions contributing to yield and yield components to develop QTL introgressed lines that can be deployed in rice breeding program.

1.Marker-Assisted Selection Laboratory, Department of Genetics and Plant Breeding, University of Agricultural Sciences, GKVK, Bangalore, 560 065 India 2.Applied Phytologics INC, 4110N Freeway, Sacramento, CA 95834, U.S.A 3.Biotechnology Centre, Punjab Agricultural University, Ludhiana, 141 004 India 4.Indian Agricultural Research Institute, Pusa Road, New Delhi, 110 012 India 5.International Rice Research Institute, P.O. Box 933, 1099 Manila, Philippines

2002

0.0

1.494

方萍, 季天委, 陶勤南, 吴平. 两种供氮水平下水稻穗长QTLs的检测. 中国水稻科学, 2002, 16: 176-178

Fang

, Ji

T W

, Tao

Q N

, Wu

Detecting QTLs for rice panicle length under two nitrogen levels. Chin J Rice Sci, 2002, 16: 176-178 (in Chinese with English abstract)

Quantitative trait loci (QTLs) for rice panicle length under two nitrogen levels were mapped on to the molecular marker linkage map by using a rice doubled haploid population derived from a cross between IR64 and Azucena. Two QTLs for panicle length, one on chromosome 1 and the other on chromosome 4 were detected at high N level, while five QTLs for panicle length were identified at low N level. The interaction between N levels and QTL for rice panicle length was therefore suggested. Based on this interaction and different response to low N stress between two parents, it was supposed that some QTLs only detected at low N level might be related with the ability of rice to tolerate low N stress.

(1College of Environmental and Resources Science; Zhejiang University; Hangzhou 310029; China; E-mail: fangping@mail.hz.zj.cn; 2College of Life Science; Hangzhou 310029, China)

在水稻(IR64×Azucena) DH群体中应用分子标记连锁图，对不同供氮水平下的穗长性状进行QTL区间作图分析。在高氮水平下检测到控制穗长的QTLs 2个，分别位于第1和第4染色体上；在低氮水平下检测到5个穗长QTLs，其中3个分别位于第1、4和6染色体上，2个位于第3 染色体的不同部位，说明穗长QTL与供氮水平之间可能存在交互效应。基于这种互作效应及研究中观察到的两供试亲本对低氮胁迫的耐性差异，推测仅在低氮水平下检测到的穗长QTLs可能跟水稻对低氮胁迫耐性有一定的关联。

2008

0.0

1.494

杜景红, 樊叶杨, 王磊, 庄杰云. 应用剩余杂合体衍生的近等基因系分解水稻产量性状QTL. 中国水稻科学, 2008, 22: 1-7

J H

, Fan

Y Y

, Wang

, Zhuang

J Y.

Dissection of QTLs for yield traits by using near isogenic lines derived from residual heterozygous lines in rice. Chin J Rice Sci, 2008, 22: 1-7 (in Chinese with English abstract)

Three residual heterozygous lines(RHLs)carrying heterozygous segments in intervals RM587-RM225,RM204-RM6119 and RM6119-RM402 on the short arm of rice chromosome 6,respectively,were selected from a rice population derived from an RHL for interval RM587-RM402.Ten maternal homozygotes,10 paternal homozygotes and 20 heterozygotes were selected from each of the F2 populations derived from the three RHLs.The three sets of near isogenic lines(NILs)were grown for phenotyping on grain yield per plant,number of panicles per plant,number of filled grains per panicle and 1000-grain weight.With analysis on the phenotype differences among the three genotype groups in each NIL set and that among overlapping chromosome segment substitution lines,three QTLs for filled grain number per panicle and two QTLs for grain yield were resolved.They were located in intervals ranging from 0.66 Mb to 2.49 Mb.The additive effect was higher than the dominance effect at each locus.The allele for increasing the trait value was derived from the paternal parent at qNFGP6-1,and from the maternal parent at the other QTLs.Following the present study,an approach for the construction of new genetic resource to facilitate QTL fine-mapping in rice was proposed.

以杂合区间为RM587-RM402的水稻剩余杂合体（RHL）衍生群体为材料，应用SSR标记检测，筛选到杂合区间分别为RM587-RM225、RM204-RM6119和RM6119-RM402的3个单株，进一步检测其F2群体，分别获得母本纯合型材料10株、父本纯合型材料10株和杂合型材料20株。种植这3套近等基因系材料，考查单株产量及其构成因子每株穗数、每穗实粒数和千粒重。经应用目标区间内等位基因效应分析和交迭重组染色体片段代换系分析，分解出3个控制每穗实粒数的QTL和2个控制单株产量的QTL，这些QTL分别位于物理距离为0.66 ~ 2.49 Mb的区间中，全部表现为加性作用为主，增效等位基因除qNFGP61来自父本密阳46外，其余均来自母本珍汕97B。提出了构建新型遗传材料，提高水稻QTL精细定位效率的策略。

2007

3.658

0.0

Yan

C J

, Zhou

J H

, Yan

, Chen

, Yeboah

, Tang

S Z

, Liang

G H

, Gu

M H.

Identification and characterization of a major QTL responsible for erect panicle trait in japonica rice (Oryza sativa L. ). Theor Appl Genet, 2007, 115: 1093-1100

Panicle erectness (PE) is one of the most important traits for high-yielding japonica cultivars. Although several cultivars with PE trait have been developed and released for commercial production in China, there is little information on the inheritance of PE traits in rice. In the present study, 69 widely cultivated japonica cultivars and a double haploid (DH) population derived from a cross between a PE cultivar (Wuyunjing 8) and a drooping panicle cultivar (Nongken 57) were utilized to elucidate the mechanisms of PE formation and to map PE associated genes. Our data suggested that panicle length (PL) and plant height (PH) significantly affected panicle curvature (PC), with shorter PL and PH resulting in smaller PC and consequently more erect. A putative major gene was identified on chromosome 9 by molecular markers and bulk segregant analysis in DH population. In order to finely map the major gene, all simple sequence repeats (SSR) markers on chromosome 9 as well as 100 newly developed sequence-tagged site (STS) markers were used to construct a linkage group for quantitative trait locus (QTL) mapping. A major QTL, qPE9-1, between STS marker H90 and SSR marker RM5652, was detected, and accounted for 41.72% of PC variation with pleiotropic effect on PH and PL. another QTL, qPE9-2, was also found to be adjacent to qPE9-1. In addition, we found that H90, the nearest marker to qPE9-1, used for genotyping 38 cultivars with extremely erect and drooping panicles, segregated in agreement with PC, suggesting the H90 product was possibly part of the qPE9-1 gene or closely related to it. These data demonstrated that H90 could be used for marker-aided selection for the PE trait in breeding and in the cloning of qPE9-1.

1.Ministry of Education of China The Key Laboratory of Plant Functional Genomics Yangzhou 225009 Jiangsu People’s Republic of China 2.Agricultural College of Yangzhou University Jiangsu Key Laboratory of Crop Genetics and Physiology 12 East Wenhui Road Yangzhou 225009 Jiangsu People’s Republic of China 3.Savanna Agricultural Research Institute (SARI) Tamale Ghana

2009

3.658

0.0

Wang

J Y

, Nakazaki

, Chen

S Q

, Chen

W F

, Saito

, Tsukiyama

, Okumoto

, Xu

Z J

, Tanisaka

Identification and characterization of the erect-pose panicle gene EP conferring high grain yield in rice (Oryza sativa L. ). Theor Appl Genet, 2009, 119: 85-91

The breeding of japonica varieties with erect-pose panicle (EP) has recently progressed in the northern part of China, because these varieties exhibit a far higher grain yield than the varieties with normal-pose panicle (NP). A genetic analysis using the F₂ population from the cross between Liaojing5, the first japonica EP variety in China, and the Japanese japonica NP variety Toyonishiki revealed that EP is governed by a single dominant gene EP. Based on previous studies, map-based cloning of EP locus was conducted using Liaojing5, Toyonishiki, their F₂ population, and a pair of near-isogenic lines for EP locus (ZF14 and WF14) derived from the cross between the two varieties; consequently, the STS marker H90 was found to completely cosegregate with panicle pose. The H90 is located in the coding sequence AK101247 in the database, and the AK101247 of Liaojing5 has a 12 bp sequence in exon 5 replaced with a 637 bp sequence of its wild type allele. It was therefore considered that the AK101247 encodes the protein of the wild type allele at EP locus, and that the sequence substitution in exon 5 of Liaojing5 is crucial for expression of the EP phenotype. The effects of EP gene on agronomic traits were investigated using two pairs of near-isogenic lines (ZF6 vs. WF6 and ZF14 vs. WF14) derived from the cross between the two varieties. Experimental results showed that EP gene markedly enhanced grain yield, chiefly by increasing number of secondary branches and number of grains on the secondary branch. EP gene also produced a remarkable increase in grain density.

1.Shenyang Agricultural University Key Laboratory of Crop Physiology, Ecology, Genetics and Breeding, Ministry of Agriculture/Key Laboratory of Northern Japonica Rice Breeding of Liaoning 110161 Shenyang China 2.Kyoto University Laboratory of Plant Breeding, Graduate School of Agriculture Kyoto 606-8502 Japan

2009

35.209

0.0

Huang

X Z

, Qian

, Liu

Z B

, Sun

H Y

, He

S Y

, Luo

, Xia

G M

, Chu

C C

, Li

J Y

, Fu

X D.

Natural variation at the DEP1 locus enhances grain yield in rice. Nat Genet, 2009, 41: 494-497

Grain yield is controlled by quantitative trait loci (QTLs) derived from natural variations in many crop plants. Here we report the molecular characterization of a major rice grain yield QTL that acts through the determination of panicle architecture. The dominant allele at the DEP1 locus is a gain-of-function mutation causing truncation of a phosphatidylethanolamine-binding protein-like domain protein. The effect of this allele is to enhance meristematic activity, resulting in a reduced length of the inflorescence internode, an increased number of grains per panicle and a consequent increase in grain yield. This allele is common to many Chinese high-yielding rice varieties and likely represents a relatively recent introduction into the cultivated rice gene pool. We also show that a functionally equivalent allele is present in the temperate cereals and seems to have arisen before the divergence of the wheat and barley lineages.