亲本籼粳成分与两系杂交粳稻杂种优势的关系及遗传基础

2026-01-13 12:56:22 世界杯最佳进球

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于亚辉, 刘郁, 李振宇, 陈广红, 徐正进, 唐亮, 毛艇, 徐海. . 亲本籼粳成分与两系杂交粳稻杂种优势的关系及遗传基础. 作物学报, 2016, 42(5): 648-657

YU Ya-Hui, LIU Yu, LI Zhen-Yu, CHEN Guang-Hong, XU Zheng-Jin, TANG Liang, MAO Ting, XU Hai. . Relationship between Indica-Japonica Index of Parents and Heterosis of Hybrid and Its Genetic Basis in Japonica Two Line Hybrid Rice . ACTA AGRONOMICA SINICA, 2016, 42(5): 648-657

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作物学报编辑部

亲本籼粳成分与两系杂交粳稻杂种优势的关系及遗传基础

于亚辉1,2, 刘郁1, 李振宇1, 陈广红1, 徐正进2, 唐亮2,*, 毛艇1, 徐海2

1辽宁省盐碱地利用研究所, 辽宁盘锦 124010

2沈阳农业大学水稻研究所, 辽宁沈阳 110866

* 通讯作者(Corresponding author): 唐亮, E-mail: tl_rice@126.com

第一作者联系方式: E-mail: yyh666@sina.com

收稿日期:2015-09-11

接受日期:2016-01-11

网络出版日期:2016-02-18

基金:本研究由辽宁省农业领域青年科技创新人才培养计划(2015034, 2014046)和辽宁省科学技术计划项目(2014020004-101)资助

摘要

以籼粳重组自交系(秋光×七山占, RIL)和粳型光温敏核不育系(GB028S)及其杂交F1为材料, 利用程氏指数法和分子标记法分析亲本的籼粳成分与杂种优势的关系及遗传基础。结果表明, 采用程氏指数法和分子标记法在群体籼粳分类的结果上比较一致; RIL偏粳系数与F1产量及其杂种优势均呈显著或极显著的二次曲线关系; F1产量在偏粳系数0.55~0.70区间内出现高峰值, 杂种优势在偏粳系数0.50~0.65区间内出现高峰值, 即RIL偏粳系数为0.55~0.65时F1有形成较高产量及杂种优势的潜力。Chr.8、Chr.11和Chr.12的籼粳成分与F1产量及杂种优势关系密切, 双亲的遗传距离与F1产量和相关性状及杂种优势没有明显的关系。

关键词:

重组自交系; 两系杂交粳稻; 杂种优势; 籼粳成分

Relationship between Indica-Japonica Index of Parents and Heterosis of Hybrid and Its Genetic Basis in Japonica Two Line Hybrid Rice

YU Ya-Hui1,2, LIU Yu1, LI Zhen-Yu1, CHEN Guang-Hong1, XU Zheng-Jin2, TANG Liang2,*, MAO Ting1, XU Hai2

1 Liaoning Institute of Saline-Alkali Land Utilization, Panjin 124010, China

2 Rice Research Institute, Shenyang Agricultural University, Shenyang 110866, China

Fund:This study was supported by the Cultivation Plan for Youth Agricultural Science and Technology Innovative Talents of Liaoning Province (2015034, 2014046) and the Science and Technology Items of Liaoning Province (2014020004-101)

Abstract

In order to further reveal the relationship between composition of subspecies lineage and heterosis in Oryza sativa L., we built a bridge for morphological index, Cheng’s index (Chi) and molecular index, japonica discrimination values ( Dj) based on two populations, a recombinant inbred lines (RILs), derived from the cross between indica cultivar Qishanzhan and japonica cultivar Akihikari, and japonica photo-thermo sensitive genic male sterile (PTGMS) line GB028S and their F1 hybrids. Two indexs achieved relatively consistent results in two populations independently. Significant curvilinear relationship appeared among Dj values, F1 yield and yield heterosis. The peaks of F1 yield and yield heterosis were detected in Dj interval from 0.55 to 0.70 and from 0.50 to 0.65 respectively, indicating that 0.55-0.65 is the preferential Dj interval for producing high yield and heterosis of F1 hybrids. Compositions of subspecies lineage on Chr.8, Chr.11, and Chr.12 were closely related with yield and heterosis of F1,however, the relationship among parent’s genetic distance, F1 yield and heterosis was not significant.

Keyword:

RIL; Two-line hybrid rice; Heterosis; Indica-japonica composition

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自1908年Shull提出“ 杂种优势” 以来, 杂种优势在提高作物产量上被广泛利用。水稻特别是亚洲栽培稻主要分为籼稻和粳稻两个亚种, 其遗传分化程度较大, 在育种研究中广泛认为籼粳交具有较强杂种优势。自1951年以来, 沈阳农业大学杨守仁教授团队开始从事籼粳稻杂交育种的基础研究, 认为籼粳交是产生杂种优势的有效途径[1]。经过育种实践, 育成了应用价值很高的沈农98366、沈农127和沈农1033等。在相同理论的指导下, 我国育种家利用籼粳杂交陆续育成多个新品种, 如浙江的矮粳23、T209、城特232等, 辽宁的辽粳5号和辽粳326, 江苏的紫金粳和南粳35, 北京的中作系列, 湖北的鄂育晚5号等[2]。同时, 国外学者对籼粳杂交育种技术亦高度重视, 20世纪60年代, 韩国育种家利用籼粳杂交育成了统一、水原、密阳等高产矮秆品种。日本于1981开始籼粳交育种, 先后育成了中国91、北陆125、关东146等新品种[2]。籼粳杂交理论不仅在常规育种上得到广泛应用, 更是杂交稻育种的重要手段。20世纪90年代, 我国育种家直接利用籼粳杂交强杂种优势成功培育出两优培九、Y两优2号、Y两优900等一系列两系法杂交水稻组合, 其中Y两优900最高单产达14.82 t hm-2。截至2012年, 两系杂交水稻累计推广3.33× 107 hm2, 增产稻谷1.11× 1010 kg[3, 4, 5]。而在我国北方, 两系杂交水稻受气候环境条件的限制发展较慢。籼粳杂交F1具有大穗、大粒、生育期偏长和生长量大等特点, 较适宜在温光条件较好的南方稻区种植; 但在北方稻区生育期偏长和抗寒性差却是籼粳杂交F1最大的种植障碍, 易引起结实率低、早衰和倒伏等弊病, 使高产稳产受到极大的影响。所以, 在提高籼粳交强优势利用的同时, 避免籼稻血缘引入造成的适应性下降已成为困扰北方两系杂交水稻育种的重要问题。为此, 探讨籼粳交的遗传基础, 研究调节搭配亲本组合及其籼粳成分, 使北方两系杂交水稻能获得高产稳产成为了现阶段研究的重点。本文通过籼粳重组自交系(RIL)与粳型光温敏核不育系杂交获得杂种F1, 利用程氏指数法和分子标记法分析RIL和不育系的籼粳成分及遗传距离, 研究其与F1杂种优势的关系, 旨在为北方两系杂交水稻的发展打下理论基础。

1 材料与方法1.1 试验材料及设计2012年以粳型光温敏核不育系GB028S和高代籼粳交重组自交系(秋光× 七山占) RIL F10为亲本, 杂交构建F1群体, 该杂种群体F1共98个。2013年将杂种F1群体与RIL种植于辽宁省盐碱地利用研究所试验地, 每株系2行, 每行2 m, 行距为30.0 cm, 株距为13.3 cm, 每穴单株, 2次重复, 栽培管理同当地生产田。

杂种F1群体与RIL齐穗后测量株高, 成熟后收获每株系5株, 网室风干后考种, 调查有效穗数、穗长、穗粒数、千粒重、结实率及单株产量等性状。利用中亲优势代表杂种优势, HMP = (F1-MP)/MP× 100, 式中HMP为中亲优势, MP为中亲值。

1.2 以程氏指数法分析亲本籼粳成分调查分析不育系GB028S和籼粳交重组自交系的抽穗时壳色、叶毛性状、1~2穗节长、籽粒长宽比、稃毛、酚反应, 计算程氏指数(Cheng’ s index, Chi)。程氏指数在划分籼粳成分上分4级, 0~8为籼、9~13为偏籼、14~17为偏粳、18~24为粳。

1.3 以SSR分子标记分析亲本籼粳成分及遗传距离均匀选取142个SSR引物用于标记亲本的基因型, 参照毛艇等[6]的方法鉴定籼粳成分, 将与秋光带型相同的带视为一个粳性位点, 以B来表示, 与七山占带型相同的带视为一个籼性位点, 以A来表示。基因型差异程度用粳性位点所占比例Dj值表示, 即每一株系的Dj值(%) = B位点数/(A位点数+B位点数)× 100。按Dj值的含义, 0~25%为籼型、25%~50%为偏籼型、50%~75%为偏粳型、75%~100%为粳型。按照Nei计算标准遗传距离[7], 亲本间的遗传距离GD (genetic distance) = -ln [2Mxy/(Mx+My)], 公式中Mx和My分别为X和Y两材料的总片段数, Mxy为两材料的共片段数。采用SPSS 19.0和Microsoft Excel 2013软件统计分析试验所得数据, 并绘制图表。

2 结果与分析2.1 RIL和GB028S的籼粳表现图1的RIL和GB028S程氏指数整体偏粳分布, 程氏指数主要分布在14~18之间, 其中GB028S的程氏指数为20.5。偏粳系数主要集中在0.60~0.90之间, 整体偏粳, GB028S的偏粳系数为0.875。程氏指数Chi和分子标记的偏粳系数Dj值呈极显著正相关(相关系数r = 0.450), 由此可见利用程氏指数法及分子标记法的偏粳系数在衡量群体籼粳成分上具有较高的相似性, 可以利用它们协同划分目标群体的籼粳成分。

图1Fig. 1Figure OptionViewDownloadNew Window 图1 RIL和GB028S的偏粳系数及程氏指数比较Fig. 1 Comparison of parental Chi and Dj

2.2 RIL的籼粳成分及亲本间遗传距离与F1性状及杂种优势的相关分析由图2可以看出单株产量与偏粳系数存在二次曲线关系, 父本RIL的偏粳系数Dj值在0.55~0.70区间时, F1的单株产量较高, Dj值为0.67时产量达峰值。单株产量杂种优势与偏粳系数也存在二次曲线关系(图2), F1的产量杂种优势的高峰区出现在父本RIL Dj值0.50~0.65区间内。两个高峰区在偏粳系数0.55~0.65重合, 说明就母本GB028S, 当父本偏粳系数为0.55~0.65时有形成较高产量及杂种优势的潜力。父本RIL的程氏指数和Dj值都与杂种F1的每穗粒数呈极显著负相关(表1), 父本的程氏指数与杂种F1的千粒重呈极显著正相关, 与结实率呈显著正相关, 与着粒密度呈极显著负相关。父本RIL的Dj值与杂种F1的单株产量呈极显著负相关。

图2Fig. 2Figure OptionViewDownloadNew Window 图2 RIL的Dj与F1单株产量和杂种优势的相关分析Dj: 偏粳系数; HMP: 中亲优势。Fig. 2 Relationship of RILs’ Dj with yield and HMP in F1Dj: Deviation japonica coefficient; HMP: Mid-parent heterosis.

表1Table 1表1(Table 1) 表1 RIL籼粳成分及亲本间遗传距离与F1产量性状和杂种优势的相关分析 Table 1 Relationship of RILs’ indica-japonica index and parental genetic distance with traits and heterosis in F1性状Trait程氏指数 Cheng’ s index偏粳系数 Dj遗传距离 GDX1X2X1X2X1X2单株产量 Grain weight per plant-0.0510.082-0.287* * -0.211* -0.0490.038穗数 Panicles per plant0.013-0.309* * -0.074-0.164-0.005-0.071穗粒数 Grains per panicle-0.340* * 0.077-0.261* * -0.072-0.130-0.015千粒重 1000-grain weight0.302* * 0.0140.165-0.015-0.051-0.154结实率 Seed setting rate0.255* -0.084-0.003-0.240* 0.084-0.056株高 Plant height-0.1520.041-0.236* -0.1520.1460.103穗长 Panicle length-0.0860.009-0.146-0.080-0.056-0.079着粒密度 Grain density-0.276* * -0.087-0.194-0.166-0.117-0.058* , * * mean significant correlation at the 0.05 and 0.01, probability levels, respectively; X1: Relationship of parental indica-japonica index and GD with F1yield; X2: Relationship of parental indica-japonica index and GD with F1heterosis.

* 和* * 表示在0.05、0.01水平上显著相关; X1: RIL籼粳成分及亲本间遗传距离与F1产量性状的相关系数; X2: RIL籼粳成分及亲本间遗传距离与F1产量性状杂种优势的相关系数。

表1 RIL籼粳成分及亲本间遗传距离与F1产量性状和杂种优势的相关分析 Table 1 Relationship of RILs’ indica-japonica index and parental genetic distance with traits and heterosis in F1由表1可以看出, F1产量的杂种优势与父本RIL的偏粳系数Dj值的负相关达到显著水平。F1穗数的杂种优势与父本RIL的程氏指数呈极显著负相关。F1结实率的杂种优势与父本RIL的偏粳系数Dj值呈显著负相关。F1其他农艺性状的杂种优势与父本RIL的程氏指数和Dj偏粳系数的相关均未达到显著水平。亲本间的遗传距离与F1单株产量和部分性状及其杂种优势的相关都未达到显著水平。

2.3 RIL染色体偏粳分布与F1产量和部分性状及杂种优势相关分析根据分子标记所在染色体不同, 分别计算父本RIL各家系每条染色体偏粳系数, 分析父本RIL单一染色体籼粳片段分布与杂种F1产量等性状及杂种优势的关系(表2)。杂种F1的产量与父本RIL的Chr.8和Chr.11的偏粳系数Dj值呈显著负相关, 在产量构成因素中, F1的穗数与父本RIL的Chr.8和Chr.11的Dj值呈显著负相关; F1的千粒重与父本RIL的Chr.1、Chr.11和Chr.12的Dj值的负相关, 及与Chr.9的Dj值的正相关均达到显著或极显著水平, 可见控制千粒重表达的关键籼粳成分基因多分布于Chr.1、Chr.9、Chr.11和Chr.12, 但作用方向各有不同; F1的结实率与父本Chr.3的Dj值呈极显著正相关, 与Chr.4和Chr.12的Dj值呈显著正相关; F1的株高与父本Chr.3和Chr.5的Dj值呈显著负相关, 与Chr.12的Dj值呈极显著负相关, 杂种F1的着粒密度与父本RIL的Chr.1和Chr.11的Dj值呈极显著负相关。

表2Table 2表2(Table 2) 表2 RIL染色体偏粳系数与F1单株产量及部分性状的相关系数 Table 2 Correlation coefficients between chromosomes’ Dj and F1 traits分量来源Component source单株产量Grain weight per plant穗数Panicles per plant穗粒数Grains per panicle千粒重1000-grain weight结实率Seed setting rate株高Plant height穗长Panicle length着粒密度GraindensityChr.1-0.1560.0780.066-0.349* * -0.002-0.126-0.150-0.274* * Chr.2-0.180-0.131-0.129-0.1100.052-0.150-0.215-0.020Chr.3-0.050-0.150-0.1200.0170.280* * -0.232* -0.1250.069Chr.4-0.027-0.0270.012-0.0780.206* 0.0330.112-0.101Chr.5-0.0920.0320.072-0.1300.141-0.234* -0.179-0.063Chr.6-0.186-0.105-0.097-0.143-0.019-0.0040.118-0.173Chr.7-0.0070.0170.133-0.130-0.078-0.1280.041-0.154Chr.8-0.238* -0.217* -0.071-0.101-0.012-0.067-0.056-0.075Chr.90.0020.012-0.0920.198* -0.0420.0020.0050.190Chr.100.0140.0570.0480.0550.0060.047-0.0360.064Chr.11-0.224* -0.237* 0.098-0.285* * 0.0640.037-0.012-0.275* * Chr.12-0.148-0.119-0.129-0.205* 0.212* -0.308* * -0.228-0.119* , * * mean significant correlation at the 0.05 and 0.01 probability levels, respectively.

* 和* * 表示在0.05、0.01水平上显著相关。

表2 RIL染色体偏粳系数与F1单株产量及部分性状的相关系数 Table 2 Correlation coefficients between chromosomes’ Dj and F1 traits表3表明, F1产量的杂种优势与父本Chr.12的偏粳系数Dj值呈极显著负相关。F1穗数的杂种优势与父本Chr.2的Dj值呈显著负相关, F1穗粒数的杂种优势则与父本Chr.4的Dj值呈显著负相关, F1千粒重的杂种优势与父本Chr.2的Dj值的负相关, 及与Chr.4的Dj值的负相关都达到显著水平。F1着粒密度的杂种优势与父本Chr.1和Chr.6的Dj值呈极显著负相关。

表3Table 3表3(Table 3) 表3 RIL染色体成分与F1杂种优势的相关系数 Table 3 Correlation coefficients between chromosomes’ Dj and HMP分量来源Component source单株产量Grain weight per plant穗数Panicles per plant穗粒数Grains per panicle千粒重1000-grain weight结实率Seed setting rate株高Plant height穗长Panicle length着粒密度Grain densityChr.10.018-0.0290.011-0.089-0.040-0.0570.038-0.380* * Chr.2-0.155-0.215* 0.012-0.241* 0.010-0.061-0.040-0.142Chr.3-0.106-0.060-0.0790.124-0.069-0.0900.0650.163Chr.4-0.0620.022-0.216* 0.202* 0.020-0.006-0.0390.073Chr.5-0.159-0.058-0.1860.1050.027-0.1490.0400.046Chr.6-0.035-0.1550.076-0.097-0.018-0.046-0.099-0.270* * Chr.70.0080.024-0.074-0.0760.177-0.103-0.177-0.121Chr.8-0.064-0.114-0.091-0.0800.128-0.068-0.077-0.056Chr.9-0.065-0.0960.095-0.032-0.1100.0430.0520.115Chr.10-0.023-0.0460.0300.037-0.0540.011-0.0060.023Chr.11-0.118-0.006-0.161-0.010-0.1020.0400.050-0.192Chr.12-0.299* * -0.187-0.181-0.055-0.087-0.167-0.040-0.050* , * * mean significant correlation at the 0.05 and 0.01 probability levels, respectively.

* 和* * 表示在0.05、0.01水平上显著相关。

表3 RIL染色体成分与F1杂种优势的相关系数 Table 3 Correlation coefficients between chromosomes’ Dj and HMP以12条染色体的籼粳成分为变量, 对98份F1单株产量进行主成分分析, 经分析提取5个主成分, 分别用X1、X2、X3、X4、X5表示(表4), 第1到第5主成分贡献率分别为17.95%、12.00%、11.11%、9.61%和8.51%, 共累积贡献率为59.20%。第1主成分X1中Chr.5和Chr.8载荷最大, 第2主成分X1中Chr.4载荷最大, 第3主成分X1中Chr.11载荷最大, 第4主成分X1中Chr.10载荷最大, 第5主成分X1中Chr.12载荷最大。从表4可以看出贡献率最高的第1主成分内染色体的载荷值比较接近, 贡献差异较小。

表4Table 4表4(Table 4) 表4 RIL染色体籼粳成分主成分载荷矩阵 Table 4 Load matrix of principal component analysis on chromosome elements of RILs分量来源Component source主成份 Main componentX1X2X3X4X5Chr.10.311-0.576-0.3120.117-0.367Chr.20.4700.149-0.5600.119-0.259Chr.30.4890.2780.249-0.574-0.113Chr.40.4910.5040.248-0.170Chr.50.5470.1270.3200.470Chr.60.427-0.278-0.370-0.152Chr.70.4220.228-0.6470.190Chr.80.5460.272-0.3940.213Chr.90.3320.3820.369-0.133Chr.100.4500.4620.420-0.147Chr.110.265-0.3660.4910.386-0.302Chr.120.377-0.2940.1780.600 表4 RIL染色体籼粳成分主成分载荷矩阵 Table 4 Load matrix of principal component analysis on chromosome elements of RILs综合表2、表3和主成分分析可知, F1的产量及其构成要素和杂种优势与父本RIL Chr.8、Chr.11、Chr.12的偏粳系数Dj值关系密切。根据杂种F1的产量高低和产量杂种优势强弱, 挑选出极端高和极端低两组, 对其父本RIL的Chr.8、Chr.11和Chr.12染色体上的籼粳片段分布情况进行分析(图3)。如图所示, 当父本RIL染色体Chr.8上的分子标记RM408-RM25区间上存在籼性片段时, F1具有较高产量和穗数。当父本RIL染色体Chr.11上的分子标记RM286-RM4和PSM366-RM224之间存在籼性片段时, 杂种F1具有较高的产量、穗数和千粒重。当父本RIL染色体Chr.12上的RM519-RM270区间存在籼性片段时, 杂种F1具有较高的产量杂种优势。

图3Fig. 3Figure OptionViewDownloadNew Window 图3 RIL染色体籼粳成分与杂种F1产量及杂种优势相关的标记位点分布Fig. 3 Markers position for correlation of chromosomes’ Dj with yield and HMP in F1

3 讨论3.1 亲本的籼粳成分与杂种优势的关系程侃生的形态指数法(程氏指数法)在籼粳分类上得到众多学者广泛的认可, 能较圆满地区分籼粳品种, 但无法得知籼粳交育成品种的籼粳成分比例和构成。随分子生物学的飞速发展, 利用分子标记来研究亲本血缘关系及判定籼粳分类成为可能。本研究表明, 利用以形态指标的程氏指数及以分子标记为主的偏粳系数(Dj值)在分类群体籼粳成分上具有一定的相似性, 可作为划分目标材料籼粳成分的依据, 这与陈跃进等[8]的研究结果较一致。卢宝荣等[9]研究认为基于高通量InDel分子标记鉴定籼粳稻与基于程氏指数法鉴定结果在很大程度上表现一致。刘丹等[10]、孙健等[11]研究认为利用InDel、SSILP标记鉴定北方粳稻籼粳成分高效准确, 与程氏指数法鉴定结果较一致。而张培江等[12]、王彦荣等[13]研究认为程氏指数法与SSR分子标记对籼粳分类结果不完全一致。分析认为程氏指数法主要利用形态及部分生理特性, 受环境及人为操作影响较大, 有一定的误差。分子标记能直接反应DNA的差异, 相对准确率较高, SSR、ILP、InDel和SNPs[14, 15, 16, 17]等分子标记被认为是鉴定籼粳成分快速、高效的方法。

在父本籼粳成分构成与F1产量关系的分析中, 杂种F1单株产量及杂种优势与父本的程氏指数的相关分析不显著, 却与父本的偏粳系数呈极显著负相关, 进一步分析表明二者存在二次曲线关系, 说明基于粳型光温敏核不育系GB028S (Chi: 20.5, Dj: 0.875)组配的杂交组合, 当父本偏粳系数在0.55~ 0.65范围时具有形成杂种优势实现高产的潜力。在染色体的籼粳成分与单株产量等性状及相应杂种优势的相关分析中, F1单株产量与父本Chr.8和Chr.11的偏粳系数呈显著负相关, 当父本Chr.8上的RM408-RM337、RM337-PSM150、PSM150-RM38、RM38-RM25和Chr.11上的RM286-RM4、PSM366- PSM418、PSM418-RM224标记之间存在不同程度籼性片段时, 组配后的杂种F1具有较高的产量。很多研究也在上述区间附近发现很多调控产量及其相关的基因或QTL, Luo等[18]在研究杂交籼稻杂种优势上位性遗传基础时将穗数qPP8定位在RM38附近, 陈深广等[19]将控制结实率杂种优势的qHSF-8定位在RM337附近, 赵秀琴等[20]将穗粒数qGN8定位在RM38。陈冰嬬等[21]研究籼稻恢复系蜀恢527籽粒性状QTL时将千粒重qTGW11定位在RM286附近, Li等[22]和Chai等[23]将单株产量qGY11-2和qGYP11.7、赵芳明等[24]和王智权等[25]将千粒重qTGW11、徐建龙等[26]将单株有效穗数qPN11、赵秀琴等[20]将穗粒数qGN11b均定位在RM224附近, 可见标记RM224附近可能存在与产量相关的重要基因, 这需要在以后的研究中进一步筛选挖掘。单株产量的杂种优势与Chr.12的偏粳系数呈极显著负相关, 通过分析筛选出Chr.12上的RM519-PSM189、PSM189-PSM190和PSM190-RM270标记之间存在籼性片段时, F1具有较高的产量杂种优势。辛业云等[27]在研究两系杂交籼稻两优培九杂种优势中将产量qHY12定位在RM519附近, 王智权等[25]将每穗粒数qHSPP12和结实率qHSSR12及穆平等[28]将有效穗数qPN12定位在RM270附近。总之, 北方两系杂交粳稻利用籼粳交提高产量杂种优势, 其亲本选育上应侧重挖掘研究Chr.8、Chr.11和Chr.12在上述标记附近的籼性基因, 有效导入目标基因, 提高分子标记在育种实践上的应用价值。

3.2 亲本的遗传距离对杂种优势的影响及预测亲本间的遗传差异被普遍认为是子代产生杂种优势的基础, 杂种优势的产生在一定范围内取决于双亲的遗传差异性和性状互补性等因素。而遗传距离作为定量描述遗传差异的方法之一, 被许多学者用于杂种优势的预测。本研究中亲本间的遗传距离与F1单株产量和部分性状及杂种优势的相关分析都未达显著水平, 表明分子标记遗传距离与F1的杂种优势没有明显和必然的联系, 这与李任华等[29]、廖伏明等[30]、孙传清等[31]、朱作峰等[32]和张涛等[33]研究结果比较一致。而蔡建等[34]和赵庆勇等[35]研究认为分子标记遗传距离与杂种优势呈显著正相关。王友林等[36]研究认为Indel标记遗传距离在不同类型品种间具有较好的杂种优势预测能力, 可以用于单穗重的优势预测, 即随着遗传距离的扩大, 杂种优势主要体现在单穗重的增加上。罗小金等[37]认为不同材料、不同遗传距离范围F1的单株产量及杂种优势与遗传距离的相关性存在很大差别, 表现出很复杂的关系, 在特定遗传距离范围内存在相关性。游书梅等[38]研究认为基于单一性状的QTL标记的遗传距离与杂种优势存在正相关。Melchinger等[39]研究认为杂种优势和遗传距离为二次曲线相关, 一定范围内杂种优势因亲本间遗传距离的增加而增强, 但超出这个范围, 杂种优势呈现减弱的现象。也有的研究认为, 同一杂种优势群内的亲本组合中, 亲本遗传距离与杂种一代呈显著相关; 而在不同杂种优势群之间较远的组合中, 相关不显著[40, 41, 42]。Saghai等[43]、Zhao等[44]也认为分子标记遗传距离与杂种优势的关系受材料来源、亲本数目、亲缘关系及所用标记位点的数量等因素的影响, 故而预测性不足。结合上述研究分析主要原因[45, 46, 47]: ①研究的性状及标记的数目较少, 有些性状对杂种优势的表现贡献较低或无贡献。②大多数量性状的表达易受环境影响, 环境(包括同一地点不同年份)的差异导致结果的不同, 使亲本间遗传差异估计的准确性大大降低。③由于上位性的作用, 增加标记的数目可能并不能增加上述相关性, 检测结果只能代表少量基因型, 因此所得遗传差异并不能全部代表亲本间的遗传差异, 而应该寻找与所研究性状有紧密连锁的标记位点。④性状之间相关的复杂性, 也导致遗传距离和杂种优势之间的复杂化。近年来, 洪德林等[48, 49, 50, 51, 52]率先在杂交粳稻上开展亲本产量性状优异配合力的标记基因型筛选工作, 研究得出RM152-165/170和RM2439-150标记是单株日产量和单株有效穗数优异配合力效应最大的标记基因型, RM8254-120/180标记对增加穗部性状的枝梗数贡献较大, 进而为提高粳稻杂种优势预测提供了参考。赵彦宏等[53]研究把杂种优势精确分担到各个QTL位点上, 估算控制性状的QTL 遗传效应进而预测杂种优势。

综上所述, 杂种优势的形成来源于亲本间的遗传差异, 而亲本间的遗传差异可用遗传距离来衡量。虽然基于分子标记的遗传距离用于杂种优势预测在理论和实践上尚无定论, 但普遍认为遗传距离的深化研究是预测杂种优势的有效途径, 只是在度量遗传距离的方法上还有待探讨, 找到能准确有效地度量遗传距离的手段, 将提高遗传距离在杂种优势预测上的应用价值。同时, 挖掘筛选与目标性状杂种优势密切相关的功能基因座位, 利用高通量分子标记精确定位, 分析主效基因及微效基因的作用, 对目标性状开展杂种优势的预测, 提高品种选育的目标性。

4 结论利用程氏指数法及分子标记法的偏粳系数在群体籼粳分类上具有较高的一致性。父本RIL的偏粳系数Dj值与单株产量及其杂种优势存在二次曲线关系。当父本的偏粳系数为0.55~0.65时, 杂种F1单株产量和杂种优势潜力较大。父本Chr.8、Chr.11和Chr.12上部分区段的籼粳成分与杂种F1产量及杂种优势关系密切。双亲的遗传距离与F1产量和相关性状及杂种优势没有明显的关系。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

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2012

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籼粳稻杂交后代易产生超亲变异和优势，蕴藏着巨大的遗传潜力。水稻产量的进一步提升寄希望于籼粳亚种间杂种优势的直接利用，但实现这一目标还面临着许多困难，解决途径有赖于对籼粳分化与杂种优势机理的深入研究。对亚洲栽培稻的起源与分化、籼粳稻分类的方法与评价、籼粳稻杂交育种的历史、存在的主要问题与解决途径、籼粳分化程度与杂种优势的关系、籼粳稻杂交后代中亚种特性与经济性状的关系及生态环境对籼粳交后代的影响等方面的研究进展进行了回顾与展望，为超级稻选育中籼粳亚种间杂种优势的直接利用提供科学依据与参考。

... 自1951年以来, 沈阳农业大学杨守仁教授团队开始从事籼粳稻杂交育种的基础研究, 认为籼粳交是产生杂种优势的有效途径[1] ...

2003

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对水稻籼粳杂交研究的历史进行回顾,总结和分析籼粳稻杂交常规育种、籼粳交杂种优势利用、亚洲栽培稻分类和籼粳杂交遗传规律研究进展,针对籼粳稻杂交育种的现状和发展趋势提出建议.

... 在相同理论的指导下, 我国育种家利用籼粳杂交陆续育成多个新品种, 如浙江的矮粳23、T209、城特232等, 辽宁的辽粳5号和辽粳326, 江苏的紫金粳和南粳35, 北京的中作系列, 湖北的鄂育晚5号等[2] ...

... 日本于1981开始籼粳交育种, 先后育成了中国91、北陆125、关东146等新品种[2] ...

2015

0.0

推广杂交水稻是增加粮食产量的最有效途径;杂交水稻育种是发展杂交水稻的技术核心;杂交水稻育种方法是决定其选育成效的关键.笔者近年在杂交水稻育种方面取得了一定成就,为了总结经验,便于交流,就杂交水稻育种所采用的方法进行回顾.探讨杂交水稻育种方向的确定方法和育种目标确定的原则,认为杂交水稻育种必须符合国家经济建设和社会发展需求,必须符合市场经济规律需求,必须符合企业盈利需求,必须符合未来发展需求;提出杂交水稻育种性状选择要做到“全优、中庸、和谐”,即双亲遗传距离适当,绝大部分性状的选择只选中间类型的,且所选性状要求达到与人和谐、与环境和谐以及性状之间相互和谐;高度重视亲本优良性状选择和杂交种性状全优;突出早发性、理想株形、抗倒性、高结实性、后期强功能性、稻米低垩白粒率等性状选择;必须人为创造或利用自然条件对一些重要性状进行增压选择;少种植育种材料和测交F1,多安排品比组合; 特别重视品比试验的质量和准确性;注重品种审定规则及标准等.

... 1010 kg[3,4,5] ...

2015

0.0

杂交水稻的推广应用为中国粮食安全作出了重大贡献,但近年来随着农业规模化、机械化的迅速发展,杂交水稻生产受到巨大冲击,种植面积减少,且有进一步下降的趋势。分析认为,阻碍中国杂交水稻发展的主要因素有:杂交水稻原有的产量优势、种子价格优势和育秧技术优势正在消失或不复存在,杂交水稻的某些主要性状不适应现代农业生产的要求,种田大户需求变化对杂交水稻提出了更高要求。针对这些问题,提出了相应的解决策略,一是通过培育超高异交结实率的不育系,筛选最佳制种基地和时段进行全程机械化制种,大幅度提高杂交稻制种产量和质量,把杂交稻种子生产成本降低50%;二是政府部门、种子企业和科技人员要各司其职,在推进作物生产方式转型的同时,着力解决杂交水稻生产中存在的问题,确保杂交水稻可持续发展。

... 1010 kg[3,4,5] ...

1980

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... 1010 kg[3,4,5] ...

2009

0.0

A recombinant inbred line(R1L)population of F8 generations derived from the cross between qshanzhan(typical indica) and Akihikari (typical japonica) were used as materials. All of 122 SSR markers containing 94 SSR markers which was distributed on 12 chromosome of rice and 28 indica and japonica specific SSR markers were used to markeranalysis. Based on QTL analysis of Cheng's index,34 markers linked to Cheng's index were found. The 34 markers and 28indica and japonica specific SSR markers were conducted to determine the blood relationship of indica and japonica on thepopulation derived from indica and japonica crossing,and to study the effect of markers types on subspecies classificationof indica and japonica. 3,5,8 SSR markers from per chromo summed to 34,65,90 SSR markers were evenly selectedto determine blood relationship of indica and japonica on the population and to study the effect of markers amount on sub-species classification of indica and japonica. The results showed that the distribution of population derived from indica andjaponica crossing were almost continuous distribution keng (japonica) subspecies and japonica. Indica and hsien (indica)were minority. The consistent degree of classification based on the 28 indica and japonica specific SSR markers and 34markers or all of 122 SSR markers were 96%and 90%by the limit of 50%blood relationship of indica and japonica.The consistent degree of classification based on 34,65,90 markers were 91%,97%and 99%.The consistent degree ofclassification based on 28 indica and japonica specific SSR markers and 65 markers or 122 markers were more than 95%.Using few SSR markers can analyse blood relationship of indica and japopica on the population derived from indica and japomca crossing.

以典型籼稻七山占和典型粳稻秋光构建的F8重组自交系(RIL)为研究对象,选取均匀分布于水稻12条染色体的94对及籼粳特异性28对共122对SSR引物对群体进行标记分析,对与籼粳分类相关的程氏指数各性状进行QTL分析,找出与程氏指数各性状连锁的34对标记,运用这34对标记及28对籼粳特异性标记进行群体籼粳血缘测定,探讨引物类型对分类的影响并按在每条染色体上均匀选取大约3,5,8对引物共计34,65,90对引物进行群体籼粳血缘测定,探讨引物数量对分类的影响。结果表明:几种方法都将群体划分为以偏粳到粳类型居多的连续分布,籼及偏籼呈极少数。按血缘来源为50%的界限进行籼粳划分对比发现28对籼粳特异标记及34对与程氏指数连锁标记与总122对标记的相符度分别为96%和90%,均匀选取34,65,90对引物与总122对标记的相符度分别为91%,97%和99%,28对籼粳特异标记及65对均匀选取标记与122对标记的分类结果相符度均在95%以上,用较少的引物即可以对群体的籼粳血缘进行分析。

... 3 以SSR分子标记分析亲本籼粳成分及遗传距离均匀选取142个SSR引物用于标记亲本的基因型, 参照毛艇等[6]的方法鉴定籼粳成分, 将与秋光带型相同的带视为一个粳性位点, 以B来表示, 与七山占带型相同的带视为一个籼性位点, 以A来表示 ...

1979

0.0

... 按照Nei计算标准遗传距离[7], 亲本间的遗传距离GD (genetic distance) = -ln [2Mxy/(Mx+My)], 公式中Mx和My分别为X和Y两材料的总片段数, Mxy为两材料的共片段数 ...

2007

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为探明水稻品种的亲缘关系,合理利用水稻品种资源,选用55对微卫星标记引物对23个水稻品种的亲缘关系进行了研究,采用UPGMA构建系统树图进行聚类分析和计算遗传距离.结果表明:引物的扩增产物带型清晰,结果稳定;23个品种分为4种类型,其中籼型8个,粳型6个,偏籼型5个,偏粳型4个;籼型品种之间和粳型品种之间的亲缘关系均较近,偏籼型品种之间和偏粳型品种之间的亲缘关系均较远.为了提高籼稻或粳稻品种和杂交种的优势,应注意采用偏籼或偏粳材料作为亲本之一.

... 本研究表明, 利用以形态指标的程氏指数及以分子标记为主的偏粳系数(Dj值)在分类群体籼粳成分上具有一定的相似性, 可作为划分目标材料籼粳成分的依据, 这与陈跃进等[8]的研究结果较一致 ...

2009

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为了建立准确高效的籼稻和粳稻鉴定方法,对基于籼稻(93-11)和粳稻(日本晴)的全基因组DNA序列比对而获得的45个特异插入/缺失(InDcl)位点进行了实验验证.以包括93-11和日本睛在内的44个典型籼稻和典型粳稻品种为实验群体,用45对InDel引物对上述水稻品种的DNA样品进行了PCR扩增和聚丙烯酰胺凝胶电泳,获得了多态的电泳条带.对获得的各InDel位点的基因型数据矩阵进行了中性检测(neutrality test),确定了与栽培稻的籼、粳遗传分化密切相关的34个InDel位点.进一步对来自亚洲11个国家的栽培稻品种和来源不同12个野生稻物种的PCR产物和电泳结果的读取和分析,计算这些栽培稻品种和野生稻DNA样品在这34个InDel位点上的籼型或粳型基因频率,最终确定了不同样品的籼、粳特性.该籼稻和粳稻鉴定方法被称为"InDel分子指数法".与传统基于形态特征鉴定栽培稻籼、粳特性的"程氏指数法"相比,该方法不仅能够准确鉴定籼稻和粳稻,而且还具有更快捷、简便和高效的特点.另外,InDel分子指数法还可以用于野生稻样品的籼、粳特性鉴定,扩大了被检测样品的范围,具有广阔的应用前景.InDel分子指数法的建立为栽培稻育种过程中正确选用籼稻或粳稻种质资源提供了新的技术方法,也为栽培稻的起源、籼-粳遗传分化、以及籼稻和粳稻在驯化过程中如何适应地理环境变化提供了新的研究思路.

... 卢宝荣等[9]研究认为基于高通量InDel分子标记鉴定籼粳稻与基于程氏指数法鉴定结果在很大程度上表现一致 ...

2015

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为了深入剖析水稻粒形性状的遗传调控机理,以典型籼稻品种七山占(Qishanzhan)和典型粳稻品种秋光(Akihikari)为亲本构建的重组自交系群体为材料,于 2011~2013年分别对各株系的粒长、粒宽和粒厚3个粒形性状进行表型测定,并基于完备区间作图法(ICIM)进行粒形性状基因定位研究。试验结果表明:共检测到27个控制粒形性状的QTLs,包括3个粒长QTLs,11个粒宽QTLs和13个粒厚QTLs,它们分布于第1,2,3,4,5,11和 12号染色体上,可分别解释14.45%~38.48%,28.98%~52.36%和38.77%~44.23%的表型变异;进一步分析发现,在第 3,5和12号染色体上检测到的粒形性状QTL位点较多,且呈簇分布;此外,检测到q GL12a,q GL12b,q GW1,q GW5a,q GT11a和q GT12b等6个较新的QTLs位点,其中控制粒宽的q GW5a连续3年表达稳定,是一个重演性极好的QTL位点。以上结果将为水稻粒形性状的QTL克隆和遗传改良奠定基础。

... 刘丹等[10]、孙健等[11]研究认为利用InDel、SSILP标记鉴定北方粳稻籼粳成分高效准确, 与程氏指数法鉴定结果较一致 ...

2012

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... 刘丹等[10]、孙健等[11]研究认为利用InDel、SSILP标记鉴定北方粳稻籼粳成分高效准确, 与程氏指数法鉴定结果较一致 ...

2000

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用12个水稻亲本按NCⅡ设计配组32F1杂种,以汕优 63为对照,研究播始历期、株高、穗长及产量因素等8个性状的杂种优势.并以12个亲本为DNA样品来源,通过随机引物 PCR扩增基因组DNA的多态性,探索利用RAPD标记水稻亲本遗传距离预测杂种优势的可能性. 由RAPD数据计算的Nei's遗传距离创建聚类树状图.聚类分析结果表明,籼稻和粳稻容易被分开,普通粳稻又容易与光壳稻、爪哇稻分开,但光壳稻和爪哇稻混合聚在一起.F 1 每穗总粒数的优势最强,中亲优势平均为33.46%,竞争优势平均为23.10%.F1播始历期、株高、穗长、有效穗4个性状中亲优势和竞争优势均表现为粳×粳＜粳×偏粳＜粳×籼.每穗总粒数的中亲优势也表现上述趋势,而竞争优势则是粳×粳＜粳×籼＜粳×偏粳.全生育期、株高、穗长的中亲优势和竞争优势与遗传距离之间均达极显著相关.每穗总粒数的中亲优势与遗传距离之间达显著相关,而竞争优势则达极显著相关.根据聚类图发现普通粳稻亚群内杂种优势较弱,亚群间即生态群间的杂种优势较强,群间即籼、粳亚种间杂种优势更强. 利用光壳稻、爪哇稻选育不同生态群方向的恢复系和不育系,配组超强优势的杂交稻组合 .

... 而张培江等[12]、王彦荣等[13]研究认为程氏指数法与SSR分子标记对籼粳分类结果不完全一致 ...

2009

0.0

摘要：以北方杂交粳稻骨干亲本及其组合为材料,采用程氏指数法和微卫星分子标记法,对亲本籼粳成分与杂种产量和稻米品质性状的关系进行了研究。杂交粳稻骨干亲本的程氏指数为13.5～19.3,偏籼系数为0.12～0.38,采用程氏指数和分子标记法对亲本进行籼粳分类的结果并不完全一致。父本程氏指数与产量杂种优势和杂种性状的相关性强于母本,而对于偏籼系数则相反。亲本程氏指数差异与产量杂种优势和杂种性状的相关性强于分子标记差异。亲本籼粳成分及其差异与多数杂种性状和产量杂种优势呈明显的二次曲线关系,存在临界极值;亲本籼粳成分及其差异与杂种产量性状和稻米品质性状的相关性相反。因此,要获得产量杂种优势和杂种表现最佳、高产与优质相结合,父母本籼粳成分应适度搭配。

... 而张培江等[12]、王彦荣等[13]研究认为程氏指数法与SSR分子标记对籼粳分类结果不完全一致 ...

2011

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... 分子标记能直接反应DNA的差异, 相对准确率较高, SSR、ILP、InDel和SNPs[14,15,16,17]等分子标记被认为是鉴定籼粳成分快速、高效的方法 ...

2010

0.0

利用籼粳杂种优势是实现水稻超高产育种目标的重要途径之一。利用根据粳稻日本晴和籼稻93-11基因组序列设计的34对特异性插入/缺失(Insertion /Deletion,InDel)引物,对采自云南省7个地州17个县的181份糯稻品种进行籼粳性分析。通过对这些糯稻品种DNA样品在这34个 InDel位点上的籼型或粳型基因频率,确定各个糯稻品种的籼、粳性。结果表明:141份测试品种被鉴定为典型籼稻,18份为籼稻,8份为典型粳稻,13 份为粳稻,1份为偏粳稻。主成分分析表明,云南糯稻籼粳分化明显。

... 分子标记能直接反应DNA的差异, 相对准确率较高, SSR、ILP、InDel和SNPs[14,15,16,17]等分子标记被认为是鉴定籼粳成分快速、高效的方法 ...

2015

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利用21对特异插入/缺失(Insertion /Deletion,InDel)引物,对自主选育的浙粳4A与22个恢复系及其组合进行水稻籼粳属性分析.根据所配组合的产量性状表现,分析亲本的 InDel遗传距离与杂种优势的相关性,探讨利用InDel标记遗传距离预测杂种优势的可行性.以供试材料在InDel位点上的多态性为原始数据,通过 NTSYS聚类分析进行籼粳属性划分,结果与王明军等的方法所得结果基本一致.结果表明,InDel遗传距离与每穗总粒数、每穗实粒数、单株产量呈极显著正相关,相关系数分别为0.627**,0.602**和0.712**;与单株有效穗数呈显著正相关,相关系数为0.426*;与结实率、千粒重的相关性不显著,相关系数分别为0.217和0.004.表明InDel标记可用于水稻籼粳属性鉴定及杂种优势预测.

... 分子标记能直接反应DNA的差异, 相对准确率较高, SSR、ILP、InDel和SNPs[14,15,16,17]等分子标记被认为是鉴定籼粳成分快速、高效的方法 ...

2012

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Forty SSILP, 30 InDel and 43 SSR markers were used to compare genetic diversity of 42 weedy and cultivated rice entries from Sri Lanka, 9 indica and 7 japonica rice cultivars as well as four representatives of weedy rice from China and other countries(regions). There were 84, 61 and 213 alleles identified by SSILP, InDel and SSR markers, with average polymorphism information content (PIC) values 0.257, 0.255 and 0.543, respectively. The NJ clusters analysis indicated all 42 Sri Lanka weedy and cultivated rice entries belonged to indica type. The genetic distances of SSILP and InDel markers revealed highly correlation (r=0.989). Meanwhile the associations between SSR markers and the other two markers (SSILP and InDel) were lower as 0.812 and 0.808, respectively. With the rising reliability and accuracy of indicajaponica classification resulted from the development of more subspeciesspecific markers, SSILP and InDel markers can be applied extensively for the indicajaponica classification of rice germplasm. And SSR markers can classify better in the subspecies.

采用40对SSILP、30对InDel和43对SSR标记对来自斯里兰卡的28份杂草稻和14栽培稻，来自国内外的9份籼型栽培稻、7份粳型栽培稻和4份代表性杂草稻进行遗传多样性分析。SSILP、InDel和SSR标记分别检测到84、61和213个等位基因，其平均多态性信息含量(PIC)分别为0.257、0.255和0.543。聚类分析显示供试的42份斯里兰卡杂草稻和栽培稻均为籼型。SSILP与InDel标记的相关性很高（r=0.989），而它们与SSR标记相关性较低（0.812和0.808）。结果表明，SSILP和InDel标记可高效鉴别各类稻种资源的籼粳属性，而SSR标记更适用于亚种内的分类。

... 分子标记能直接反应DNA的差异, 相对准确率较高, SSR、ILP、InDel和SNPs[14,15,16,17]等分子标记被认为是鉴定籼粳成分快速、高效的方法 ...

2009

0.0

... 很多研究也在上述区间附近发现很多调控产量及其相关的基因或QTL, Luo等[18]在研究杂交籼稻杂种优势上位性遗传基础时将穗数qPP8定位在RM38附近, 陈深广等[19]将控制结实率杂种优势的qHSF-8定位在RM337附近, 赵秀琴等[20]将穗粒数qGN8定位在RM38 ...

2010

0.0

[目的]利用QTL定位方法检测水稻产量性状杂种优势QTL,并解释杂种优势产生的可能分子机理.[方法]利用重组自交系与亲本协青早B构建BC1杂种群体,通过两地重复试验,以中亲优势考察6个产量性状的杂种优势表型,利用Windows QTL Cartographer 2.5的复合区间作图法检测其QTL.[结果]多数产量性状均表现出较强的杂种优势.在两地试验中,共检测到20个产量性状杂种优势QTL,分布在水稻第 2、3、6、7、8、10等6条染色体上,包括3个控制单株产量杂种优势的QTL、2个控制单株穗数杂种优势的QTL、6个控制每穗总粒数杂种优势的 QTL、4个控制每穗实粒数杂种优势的QTL、4个控制结实率杂种优势的QTL和1个控制千粒重杂种优势的QTL.单个QTL对群体性状表型变异的贡献率为4.90%-12.85%.[结论]检测到控制6个产量性状杂种优势的20个QTL,其中qHNP-3、qHTNSP-7、qHNFGP-7、 qHSF-7、qHTGWT-3 5个QTL在两地试验中稳定表达;检测到的20个杂种优势QTL中,有1 3个与在RIL群体中检测到的QTL重叠,重叠率达65%,因此,认为来自纯系的产量性状加性效应对杂种优势产生具有重要贡献.

2007

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In rice breeding community, drought tolerance (DT) is becoming one of the most important target traits for variety improvement under ever-increasing severe drought situation all over the world. Yield is the most important target focused by breeders, so identification of favorable alleles related to yield and yield components for introgression into rice varieties that suit the specific environments is an efficient practice in plant MAS breeding. In the present study, the QTL conditioning the grain yield per plant (GY) and its component traits including 1000-grain weight (GW) and filled grain number per panicle (GN) in irrigation(control) and rainfed (drought stress) fields were investigated using 254 advanced backcrossing introgression lines (IL) of Lemont with Teqing genetic background and the two parents. Total of 32 main-effect QTLs for GY, GW, and GN were identified in the two conditions which can be grouped into three types based on their behaviors. TypeⅠincluded 10 QTLs which were detected both in the two conditions, type Ⅱ consisted of 14 QTLs which were mapped only in control condition, and type Ⅲ comprised 8 QTLs which were induced by drought and detected only under stress. In addition, nine QTLs affecting trait differences between stress and control were identified ( QGn5 , QGn6 , QGn11a , QGw2 , QGw8 , QGw11a , QGw11b , QGy1 , and QGy11 ). There were three QTLs ( QGn11b , QGn12 , and QGn11b ) which expressed both in the two conditions with same direction and magnitude of gene effect. Therefore the three GN-QTLs and nine QTLs affecting trait differences were considered to directly contribute to drought tolerance (DT). Teqing alleles at the 12 DT-QTLs except QGn5 and QGy1 were associated with DT. As compared with the results from different mapping populations derived from the same parent, Lemont and Teqing, in different environments, seven QTLs including QGn3b , QGw1 , QGw5 , QGy1 , QGy5 , QGy8 , and QGy10 which stably expressed across different genetic backgrounds and environments were identified. The strategy of applying QTL mapping results in rice improvement for DT by marker-assisted selection was discussed.

利用254个Lemont导入到特青背景的高代回交导入系定位了灌溉（对照）与自然降雨（干旱胁迫）环境下影响单株籽粒产量及其穗部相关性状的QTL。在两种环境下共检测到32个影响单株粒籽产量、千粒重和每穗实粒数的主效QTL，根据不同环境下表达的情况将其分成3类，第1类10个QTL，在两种环境下均被检测到；第2类14个QTL，只在对照条件下检测到；第3类8个QTL，受干旱胁迫诱导，只在胁迫条件下被检测到。此外还检测到9个影响胁迫与对照条件下性状差值的QTL。认为在两种条件下均检测到的相对稳定的3个QTL（ QGn11b 、 QGn12 和 QGn11b ）及影响两种条件下性状差值（即性状稳定性）的9个QTL可能对耐旱性有直接贡献。在所有12个耐旱QTL中，除在 QGn5 和 QGy1 的Lemont等位基因减小性状差值（即增强耐旱性）外，其余位点上增强耐旱性的等位基因均来自特青。另外通过与源自相同亲本的不同定位群体在不同环境下定位结果的比较，鉴别出一些受遗传背景和环境影响较小的QTL如 QGn3b 、 QGw1 、 QGw5 、 QGy1 、 QGy5 、 QGy8 和 QGy10 。对应用QTL定位结果进行标记辅助选择培育耐旱品种进行了探讨。

... 7、赵芳明等[24]和王智权等[25]将千粒重qTGW11、徐建龙等[26]将单株有效穗数qPN11、赵秀琴等[20]将穗粒数qGN11b均定位在RM224附近, 可见标记RM224附近可能存在与产量相关的重要基因, 这需要在以后的研究中进一步筛选挖掘 ...

2008

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The traditional mapping population with complicated genetic backbround is usually derived from a cross between two parents with a lot of unfavorable agronomic traits. The identified QTLs by traditional mapping can not be applied in genetic improvement of crops due to complicated interactions between QTLs and genetic background. The advanced backcross method for QTL mapping with elite variety as recurrent parent could partially eliminate the background effects and integrate QTL mapping with genetic breeding. In present study, using Shuhui 527, an elite indica restorer in China as recurrent parent and Milagrosa from Philippines as donor, the BC 2 F 2 advanced backcross population with 199 individuals was developed. A total of 85 polymorphic SSR markers evenly distributed on 12 chromosomes were applied to genotype the mapping population. The grain length, grain width, ratio of grain length to width and thousand grain weight of the parents and the BC 2 F 2 population were evaluated after har-vesting. The QTL identification of the above traits was conducted by one-way ANOVA (for single QTL) and two-way ANOVA (for digenic epistatic interaction loci). Among ten QTLs for grain length, grain width, ratio of grain length to width and thousand grain weight detected by one-way ANOVA (P qgl3b on chromosome 3 was a major QTL controlling grain length, ratio of grain length to width and thousand grain weight, which explained 29.37%, 26.15%, and 17.15% of phenotypic variation of the three traits and showed large additive effects (posi-tive allele from Shuhui 527) and negative overdominant effects, qgw8 was also a major QTL controlling grain width with 21.47% contribution to phenotypic variation and a minor QTL controlling thousand grain weight with 5.16% contribution to phenotypic variation. It had large additive effects (positive allele from Shuhui 527) and positive partial dominant effects in both traits. Two-way ANOVA (P<0.005) showed 61 pairs of digenic epistatic interactions involved in 54 QTLs among which 23 loci had pleiotropism affecting two to four traits and eight loci were the same as those detected by one-way ANOVA. It was observed that eight of thirteen pairs of digenic epistatic interactions related to ratio of grain length to width could be found in grain length. The results above will lay a good foundation in fine mapping, cloning and designed molecular breeding of favorable genes related to rice grain size and shape.

1 Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China; 2 International Rice Research Institute, DAPO Box7777, Metro Manila, Philippines; 3 Agricultural College, Anhui Agricultural University, Hefei 230036, Anhui, China

以我国优良籼稻恢复系蜀恢527为轮回亲本, 以来自菲律宾的Milagrosa为供体亲本, 培育了样本容量为199株的BC 2 F 2 高代回交群体。选取85个均匀分布在12条染色体上的多态性SSR标记进行基因型分析, 同时对粒长、粒宽、长宽比和千粒重4种性状进行了表型鉴定。采用性状－标记间的单向和双向方差分析对上述性状进行了QTL定位。单向方差分析(P qgl3b 是一个控制粒长、长宽比和千粒重的主效QTL, 它可以分别解释粒长、长宽比和千粒重表型变异的29.37%、26.15%和17.15%。该QTL对于粒长、长宽比和千粒重均表现较大的加性效应(来自蜀恢527的等位基因为增效)和负向超显性。位于第8染色体的 qgw8 位点是一个控制粒宽的主效QTL, 同时也是控制千粒重的微效QTL, 能解释粒宽表型变异的21.47%和千粒重表型变异的5.16%。该QTL对粒宽和千粒重均具有较大的加性效应(来自蜀恢527的等位基因为增效)和正向部分显性。双向方差分析(P<0.005)共检测到61对显著的上位性互作, 涉及54个QTL, 其中23个是能同时影响2~4个性状的多效位点, 且有8个位点与单向方差分析检测到的相同。控制长宽比的13对上位性互作位点中, 与控制粒长的上位性互作位点完全相同的有8对。以上结果为进一步开展水稻籽粒大小和形状有利基因的精细定位、克隆和分子设计育种奠定了基础。

... 陈冰嬬等[21]研究籼稻恢复系蜀恢527籽粒性状QTL时将千粒重qTGW11定位在RM286附近, Li等[22]和Chai等[23]将单株产量qGY11-2和qGYP11 ...

2002

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... 陈冰嬬等[21]研究籼稻恢复系蜀恢527籽粒性状QTL时将千粒重qTGW11定位在RM286附近, Li等[22]和Chai等[23]将单株产量qGY11-2和qGYP11 ...

2014

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... 陈冰嬬等[21]研究籼稻恢复系蜀恢527籽粒性状QTL时将千粒重qTGW11定位在RM286附近, Li等[22]和Chai等[23]将单株产量qGY11-2和qGYP11 ...

2011

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It is important to accelerate breeding process by pyramiding genes purposefully with molecular marker-assisted selection methods. Additive and epistatic effects of genes are the key to the success in genes pyramiding breeding. In this paper, additive and epistatic effects of QTLs for rice grain shape were analyzed using 16 single segment substitution lines (SSSL) and 15 double segment substitution lines (DSSL). A total of nine QTLs were identified on the Chromosomes 2, 3, 4, and 10 respectively, containing four for grain length, one for grain width and four for ratio of grain length to width. Furthermore, seven pairs of digenic interactions were detected for the grain shape. Among them, three interactions occurred between two loci with obvious effects on the traits, one interaction did not have obvious effects at both two loci, and three interactions occurred between one locus with and another without main effect at the single-locus level. Again, the results also indicated that the epistatic effects were different when the same QTL controlling grain length was pyramided with various SSSLs. Only when the epistatic effects were syntropic with the additive effects of the target genes, the pyramiding effect could be obvious. Moreover, longer or shorter grain was difficult to be obtained by pyramiding two QTLs controlling long grain or two QTLs for short grain. The results are important for improving grain shape by molecular marker assisted selection.

以分子标记辅助选择的手段有目的地进行基因聚合育种，对于加快育种进程具有重要的意义。而基因的加性和上位性效应是决定基因聚合能否成功的关键。本文以16个单片段代换系(SSSL)及15个双片段代换系分析了水稻粒型性状QTL的加性及上位性效应。共检测到9个水稻粒型性状QTL，包括4个粒长QTL、1个粒宽QTL和4个籽粒长宽比QTL，分别位于第2、第3、第4、第7和第10染色体上。此外，还检测出7对双基因互作，其中3对为有显著效应的两座位间互作，1对为两座位均没有显著效应的座位间互作，3对为1个有显著效应的座位与1个没有显著效应的座位间互作。本文结果进一步揭示了同一粒长QTL与不同单片段代换系聚合时会产生不同的互作效应，只有当上位性效应与目标基因的加性效应同向时，才可以达到明显改良粒长的效果。而且，2个长粒或2个短粒QTL聚合很难再产生更长或更短的籽粒。以上结果对于通过分子标记辅助育种手段改良水稻粒型具有重要意义。

2013

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A total of 66 chromosome segment substitution lines, derived from a cross between indica inbred line IR24 (as the recurrent parent) and japonica inbred line Asominori (as the donor parent), were used to investigate the heterotic loci in indica/ japonica intersubspecific rice hybrids. Each line was crossed with the background parent IR24, and the heterosis of F1 hybrids was estimated by comparing the F1 with its two parental lines. Field experiments were carried out across four different environments (2007, 2008 in Nanjing; 2007, 2008 in Nanchang) to evaluate yield and yieldrelated traits in the 66 lines and their 66 corresponding F1 hybrids. Quantitative trait loci (QTL) analyses were conducted using a likelihood ratio test based on the stepwise regression. QTL were detected by statistical software QTL IciMapping using midparental heterosis as basic phenotypic data through a new SSR map. As a result, 53 heterotic loci of yieldcomponent traits were identified with significant effects in different environments. Only one heterotic locus linked closely with marker RM488 on chromosome 1 was detected repeatedly in more than one environment,which might improve the plant height in F1 derivatives. Of all the heterotic loci, 22 (41.51%) showed positive effects, with LOD values ranging from 3.06 to 7.25, explaining 3.74% to 18.5% of the phenotypic variance respetively; 31 loci showed negative effects, with LOD values ranging from 3.07 to 9.70, explaining 0.45% to 30.78% of the phenotypic variance, respectively; Those loci with negative effects mainly affected the traits of grain weight per plant, no. of grains per panicle and seedsetting rate, which were closely related to those genes of hybrid sterility between indica and japonica rice.

1 Research Center of Jiangsu Plant Gene Engineering/State Key Laboratory of Crop Genetics and Germplasm Enhancement, Nanjing Agricultural University, Nanjing 210095, China; 2 Jiangxi Academy of Agricultural Sciences/ National Engineering Laboratory of Rice, Nanchang 330200, China; 3 Crop Science Institute, Chinese Academy of Agricultural Sciences, Beijing 100081, China;

利用以籼稻IR24为受体亲本、粳稻Asominori为供体亲本的66个染色体片段置换系，分别与受体亲本IR24杂交构建1套对应的F1群体，研究杂种优势位点。以中亲优势值作为杂种优势QTL检测的基准表型值，结合新构建的基因型图谱，利用QTL IciMapping软件的逐步回归和极大似然估计结合的方法，检测杂种优势QTL。结果表明，各置换系对应的F1群体，在4个环境（2007年南京、2007年南昌、2008年南京、2008年南昌）下，共检测到53个与产量构成性状相关的杂种优势位点。其中只发现1个杂种优势位点能在多个环境重复检测到，即来自第1染色体与标记RM488紧密连锁的位点。该位点在杂合状态下增加F1的株高。22个位点在杂合状态下具有增效作用，占总位点数的41.51%；LOD值变幅为3.06~7.25，贡献率变幅为3.74%~18.5%。31个位点在杂合状态下具有减效作用，LOD值变幅为3.07~9.70，贡献率变幅为0.45%~3078%，这些具有减效作用的位点主要控制单株产量、每穗实粒数和结实率等性状，与籼粳杂种不育基因密切相关。

... 辛业云等[27]在研究两系杂交籼稻两优培九杂种优势中将产量qHY12定位在RM519附近, 王智权等[25]将每穗粒数qHSPP12和结实率qHSSR12及穆平等[28]将有效穗数qPN12定位在RM270附近 ...

2001

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The genetic mechanism underlying panicle number per plant (PN),spikelet number per panicle (SNP)and its related traits in rice was analysed using 292 F 13 RILs from the cross of Lemont/Teqing and a complete linkage map with 272 molecular markers.The RILs showed tremendous transgressive segregation for all traits studiesd.The weak negative correlation between PN and SNP was observed.Fifty-oneQTLs and 45 epistatic QTL pairs affecting these traits were identified,collectivelyexplaining over 60% of the total variation of individual traits.Almost all SNP-QTLs were attributable to one or more of its contributing components.Branching number traits had greater contributions to SNP than length traits,in which the first had twiceas many QTLs mapped in the same or near regions with SNP as the latter.Only two PN-QTLS were mapped in the near regions with those of related traits of SNP,suggesting a reasonable recombination between PN and SNP would be available by marker-assisted selection (MAS).Some major QTLs including QPn4 for panicle number,QPbn3a,QPbn3b and QPbl4 for panicle branching and length would be of great value in MAS.It was discussed that a new high-yielding panicle type was resulted from reasonably deploying for QTLs of panicle traits by MAS.

应用292个Lemont/特青F13重组自交系(RILS)及其含272标记的遗传连锁图谱剖析单株有效穗数(PN)和每穗粒数(SNP)及其相关性状的遗传。所有性状呈现超亲分离,PN与SNP存在弱的负相关。检测到影响PN和SNP及其相关性状的QTL51个和互作位点45对,它们可以解释60%以上的性状总变异。SNP与其相关性状的QTL定位在一起,比较与SNP-QTL同一或相邻区域的QTL数,穗部枝梗数是长度性状的两倍,故前者对SNP的作用更大。仅有2个PN-QTL与SNP相关性状的QTL相邻,因此通过标记辅助选择有可能实现PN与SNP的有利重组。其中影响PN的QPn4和影响穗枝梗数和长度的QPbn3a,QPbn3和QPbl4等主效QTL,在标记辅助选择中具有重要的应用价值。对通过标记辅助选择合理配置穗部性状QTL产生新的高产穗型进行了讨论。

2014

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【Objective】 The heterosis loci and QTLs of yield and yield components were detected by using a RILsBCF1 population derived from a cross between Pei’ai 64S and 9311. The relationship was explored between the genetic variance of these two parental lines and yield heterosis in the resulted hybrid for predicting hybrid heterosis.【Method】Based on a population of 219 recombinant inbred lines (RILs) of F8 generation produced by single seed descendant method from the Pei’ai 64S×9311 cross, a RILsBCF1 population was generated by backcrossing of RILs to Pei’ai 64S. With a total of 151 polymorphic SSR markers, a linkage map was constructed spanning 1 617.7 cM across the whole genome with an average marker interval of 10.93 cM. The correlation between genetic distances and F1 trait performance of RILsBCF1 and their prediction in yield and yield component traits were conducted respectively by using molecular marker analysis, one-way ANOVA with different freedoms, and composite interval mapping using mix linear model in SAS, together with heterosis prediction and QTL mapping. 【Result】The RILsBCF1 used in the study showed significant diversity with high segregation in multiple traits, and their average performance was significantly higher than that of RILs F8. In this RILsBCF1 population, 74 heterosis positive loci and effect-increasing loci were identified by two-group method and three-group method in yield and yield component traits, respectively. Compared with two-group method, three-group method could get more positive loci or effect-increasing loci to a certain degree and raise efficiency of predicting correlationship between genetic distances of both positive loci and effect-increasing loci and F1 traits’ performances. The result of the effect-increasing loci detecting was the same in both two-group and three-group methods. Six heterosis loci were detected at the same regions for three traits (Sterile lemma per panicle, Grains per panicle and Pencentage seed setting), overlapped with three yield effect-increasing loci clustered on chromosome 7. Based on the relationship between marker-effect values of yield effect-increasing loci using the three-group method and F1 trait performance, four multiple regression prediction models were constructed using a stepwise procedure. A total of 28 markers with heterozygous genotypes were identified to significantly increase the correlation coefficient between the genetic distances and the F1 trait performance from 0.335 to 0.617. Three QTLs for yield heterosis and three QTLs for grain per panicle heterosis were mapped using this RILsBCF1 population. The mapped loci of QTL QGpp7 for grain per panicle heterosis and QHy7 for yield heterosis matched the effect-increasing loci identified by the methods of two-group and three group analyses. 【Conclusion】The approaches of screening more positive loci or effect-increasing loci and specific markers which influent heterosis can increase the correlation coefficient between the genetic distances and the F1 traits performances, and thus can be applied more efficiently in predicting the yield heterosis of rice hybrids with genetic distance of molecular markers. The yield QTL QHy7 located on chromosome 7 with a yield increase contribution of 7.48% can be used for yield heterosis prediction and in hybrid rice breeding. A heading stage QTL located between RM293-RM468 on chromosome 3 with the contribution of 14.9% can be used for rareripe high yield rice breeding.

【目的】对超级杂交稻两优培九影响产量及其构成因素性状的杂种优势位点进行定位，在此基础上探讨亲本培矮64S和9311的遗传差异与水稻产量性状的杂种优势间的关系，以探明水稻产量杂种优势的分子预测途径。【方法】应用经单粒传法获得后续世代的219个培矮64S×9311 F8重组自交系（RILs）株系材料与亲本培矮64S回交，并选用151个分布于水稻基因组12条染色体上的SSR多态性标记，构建回交群体RILs BCF1；构建基因组总长为1 617.7 cM、标记间平均距离10.93 cM和含151个分子标记的遗传图谱；采用分子标记技术和自由度不等的单向分组方差两组法、三组法分析，用SAS软件ANOVA分析、混合线性模型复合区间作图等方法，对回交RILs BCF1群体的产量性状及其构成因素的F1表型值进行相关分析、优势预测与QTL定位。【结果】本回交杂种群体RILs BCF1具备多种基因型，遗传变异丰富，性状平均值均显著高于亲本群体重组自交系RILs F8，共筛选到影响RILs BCF1群体产量及其构成因素性状杂种优势的阳性、增效位点74个；其中，三组法所筛选的阳性、增效位点数高于两组法，用这些阳性、增效位点所预测的遗传距离与产量F1性状值的相关性也显著提高；三组法所筛选产量性状的增效位点与两组法所筛选的增效位点完全一致；连锁紧密的位点有成簇分布的现象，每穗空粒数、每穗实粒数、结实率有6个杂种优势位点相同，并与3个产量杂种优势位点重叠，且均处在第7染色体上；通过逐步回归建立了对4个产量性状进行预测的回归方程模型；筛选到28个杂合型的特异性标记，它们与产量性状的表型值显著相关，使用特异性标记可使遗传距离与产量F1性状值的相关系数由全部标记的0.335提高到0.617；定位到3个与产量杂种优势相关的QTL和3个影响每穗实粒数杂种优势的QTL。其中，在第7染色体上影响每穗实粒数和产量杂种优势的QTL QGpp7和QHy7与影响每穗实粒数和产量杂种优势的增效位点的结果相符。【结论】通过增加筛选产量杂种优势阳性位点或增效位点数量、筛选影响杂种优势特异性分子标记的方法，可显著提高分子标记遗传距离与产量F1性状值的相关性，有效提高用分子标记遗传距离对杂种优势预测效率。定位了3个影响产量杂种优势的QTL及3个影响每穗总粒数杂种优势的QTL，分别在第2、3、7、11和12染色体上，其中，影响产量杂种优势的数量性状位点QHy7，贡献率为7.48%，可用于杂种优势的预测和杂交组合的选配。定位于第3染色体RM293—RM468的表型贡献率为14.9%的抽穗期QTL可用于早熟高产水稻的选育。

2008

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To study the difference of QTLs for rice yield and its component traits under different phosphorus applications, a DH (doubled haploid) population including 116 lines, derived from a cross between phosphorus deficiency tolerant rice IRAT109 and phosphorus deficiency sensitive rice Yuefu, was evaluated for yield per plant (YP) and its components including 1000-grain weight (GW), seed-setting percentage (SP), panicle number per plant (PN), and grain number per panicle (GN) under phosphorus deficiency and normal conditions. No significant differences were found in GW and GN but significant differences were found in SP, PN, and YP for the DH population between the two conditions. This result indicted that SP, PN, and YP were more easily influenced by phosphorus deficiency than GW and GN. A total of seventeen QTLs were detected for plant yield and its components under phosphorus deficiency using the constructed molecular linkage map (including 94 RFLP and 71 SSR markers and covering 1 535 cM). Among them, there were twelve QTLs with general contributions of over 10% to phenotypic variation. Three tightly linked QTL regions on chromosomes 3, 6, and 7 were identified. These QTLs with high general contribution and tightly linked QTL regions might be useful for phosphorus deficiency tolerant and high yielding rice breeding by molecular marker assisted selection.

为研究低磷胁迫对水稻产量及其构成因素QTL表达的影响, 以耐低磷旱稻IRAT109和磷敏感水稻越富杂交的116个株系的DH群体为材料, 在低磷和正常栽培条件下, 调查了千粒重、结实率、有效穗数、穗粒数及单株产量等性状。结果表明, 结实率、有效穗数和单株产量对低磷胁迫的敏感性较大, 而千粒重、穗粒数对低磷胁迫的敏感性较小。利用水稻分子连锁图中94个RFLP标记和71个SSR标记, 依据以上5性状低磷胁迫下与对照的差值进行QTL定位。共检测出17个QTL, 其中12个对表型变异的贡献率大于10%。3、6和7号染色体上3个标记区域存在QTL成簇分布, 这些高贡献率QTL及成簇分布QTL可作为水稻耐低磷产量性状分子育种的重要候选区域。

1998

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摘要：本文利用两套材料，即来自一个籼／粳交（圭６３０／０２４２８）的双单倍体（ＤＨ）系和另一组由多种类型的籼、粳组成的对照组，从分子水平上研究了水稻亲本的遗传分化程度。在此基础上探讨了亲本的遗传分化程度与籼粳杂种优势的关系。两套测交Ｆ１群体的结果一致表明，亲本遗传分化程度对穗数和千粒重的影响相对较小，但是对每穗结实粒数和结实率这两项体现育性的指标影响较大。当亲本遗传分化综合指数ＴＤｊ值为２０￣２５时，育

... 本研究中亲本间的遗传距离与F1单株产量和部分性状及杂种优势的相关分析都未达显著水平, 表明分子标记遗传距离与F1的杂种优势没有明显和必然的联系, 这与李任华等[29]、廖伏明等[30]、孙传清等[31]、朱作峰等[32]和张涛等[33]研究结果比较一致 ...

1998

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Using 7 maintainer lines and 9 restorer lines as the materials, the genetic difference of hybrid rice parents and its relation to F 1 heterosis were studied through principal component analysis and cluster analysis of 11 major agronomic characters. The results showed that among all of the 120 genetic distances of the 16 parents, morethan 80% was below 2; the average genetic distances among the 9 restorers and among the 7 maintainers except Xiangxiang 2B were 0. 4793 and 0. 7723, respectively, which indicates that the genetic difference is comparatively small in the existing hybrid rice parents, especially within restorer lines or within maintainer lines. The analysis of Correlation between the genetic distance of both parents and heterosis of their F 1 hybrids showed that there was not correlation between them among the existing hybrid rice parents. However, it was found that the grain yield of F 1 hybrids was closely related to that of both parents, especially to the mean grain yield of bo th parents, indicating thatparental improvement is of great importance to increase hybrid rice yield. Furthermore, the relation between parental improvement and heterosis, the approaches to the improvement of parents and the principle of choice of pareals for developing hybrids were also discussed.

以目前生产上大面积应用和新育成的16个籼型三系杂交水稻亲本为材料，根据11个主要农艺和经济性状进行主成分分析和聚类分析，以马氏距离为指标，研究了杂交水稻亲本间遗传差异的状况及其与杂种优势的关系。在16个亲本的120个遗传距离中，80%以上小于2；9个恢复系间的平均遗传距离仅为0.4793， 7个保持系间，除湘香2号B外，平均遗传距离为0.7723。说明现有杂交稻亲本间遗传差异较小，尤其是保持系内和恢复系内遗传差异更小。对亲本间遗传距离与杂种优势的相关分析表明，现有籼型三系杂交稻亲本间遗传差异与杂种优势关系不密切，而杂交稻产量的高低与双亲产量尤其是双亲产量平均值的高低密切相关。说明亲本改良在提高杂交稻产量中极为重要。此外，还对亲本改良与杂种优势的关系、杂交稻亲本改良的方法和途径以及杂交稻组合亲本选配原则进行了讨论。

2000

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The four photoperiod-and/or thermo-sensitive genetic male sterile rice (PGMS o r TGMS) lines were test-crossed with 47 improved cultivars used as male parents . Grain yield and yield components of the test-cross F1s exhibited remarked d ifference. It is indicated that N422s/indica from Southern Chin

以培矮64s、 108s、 N422s、 LS2s等4个两系不育系为母本，以韩国籼、中国南方的早中籼(简称中国籼)、东北粳(分东北普通粳和粳杂恢复系)、华北粳、非洲粳、美国粳等6 个生态型的47个育成品种为父本，按照NC-Ⅱ设计，配制188个组合。在所配的组合类型中， N422s/中国籼单株粒重最高，其次是N422s/韩国籼、培矮64

2001

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Genetic divergences of 57 entries from 6 ecotypes of rice were surveyed by PCR analysis with 48 pairs of SSR and by Southern analysis by 50 RFLP probes.It was indicated that SSR markers detected polymorphism effciently in comparison to RFLP markers,considering the allele numbers per locus and average gene diversity.Among all of 1596 pairs,1452 pairs of genetic distances based on SSR data (GD SSR ) were higher than the genetic distance based on RFLP data (GD RFLP )by average of 0.2492.The average GD SSR of entries within indica subspecies was 0.5483,but GD RFLP was only 0.2776.And the GD SSR and GD RFLP of entries with japonica subspecies were 0.577.0.2405 respectively.The average GD SSR was 0.7310.It is suggested that the GD SSR between varieties of introspecies or interspecies are higher than that of GD RFLP .The analysis of correlation between GD SSR and GD RFLP of any entry with other 56 entries indicated that the correlation coefficient for typical Indica or Japonica varieties were higher,but lower for Indica-like or Japonica-like materials.It was also found that the relationships between genetic distance based on these two kinds of markers and theyield of hybrid performance were not significant,but most of the correlation coeffcient between heterosisi and GD RFLP or GD SSR arrived at highly significant level.

(Department of Plant Genetics and Breeding;China Agricultural University,Beijing 100094,China)

用48对SSR引物和50个RFLP探针对来自6个生态型的57份水稻材料之间的遗传差异进行了比较分析,从单个位点的等位基因数和平均基因多样性看,SSR的多态性要高于RFLP。从遗传距离看,在总计1596对材料的遗传距离中,只有144对(占9%)用RFLP标记估计的遗传距离(简称GDRFLP,下同)高于用SSR标记估计的遗传距离(简称GDSSR,下同),平均高出的幅度为0.0371,而有1452对(占91%)材料间的GDssR高于G2RFLP,平均高出幅度为0.2492。无论是籼、粳亚种内还是籼粳亚种间,SSR标记比RFLP估计的遗传距离高。如用SSR标记估计的籼稻亚种内材料之间平均遗传距离为0.5483,而RFLP标记估计的只有0.2776;粳稻亚种内SSR平均遗传距离为0.5777,RFLP的只有0.2405;用SSR和RFLP估计的亚种间的平均遗传距离分别为0.8074和0.731.成对材料的SSR遗传距离与RFLP遗传距离之间的相关分析表明,对于典型的籼粳品种,这两种标记估计的遗传距离相关系数较高,而中间型材料或偏籼偏粳材料的相关程度较低。通过分析4个两系不育系与48个恢复系之间的遗传距离与F1的产量表现及杂种优势之间的相关关系得知,这两种标记估计的遗传距离与杂种表现间的相关均没有达到显著水平,而与杂种优势的相关系数中,除108s与恢复系的GDRFLP和LS2S与恢复系的GDSSR与超亲优势没有达到显著水平外,其他均达到了极显著的水平。

2006

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【Objective】 Correlation between genetic distance and heterosis in an important question in hybrid rice breeding. 【Method】 The genetic distance of the aromatic rice parents and the relativity analysis were studied by SSR in this article. 【Result】 The relativity coefficient was partial to small（-0.257-0.292）between SSR genetic distance and heterosis of four yield traits. The correlation between the genetic distance of the aromatic rice parents selected and the yield heterosis was not significant. 【Conclusion】So, SSR markers selected in this study couldn't predict rice heterosis. The research indicated that different or opposite results would be gained with different materials. Thus, the analysis of genetic distance based on molecular markers predicting the yield heterosis need to be studied further.

【目的】研究遗传距离与杂种优势的相关性，为杂交水稻育种提供依据。【方法】利用SSR标记检测R527等恢复系与泸香90A等不育系间的遗传距离，并分析遗传距离与产量杂种优势间的相关性。【结果】杂交香稻亲本间遗传距离与产量性状及其各构成因素杂种优势间的相关系数偏小（-0.257～0.292），均未达显著水平。【结论】所选用的杂交水稻亲本间的遗传距离大小并不能反映杂种优势；所选用的SSR标记不能预测水稻产量杂种优势，利用分子标记遗传距离来预测杂交水稻杂种优势的可靠性有待进一步探讨。

2005

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利用AFLP分子标记对46个水稻品种进行遗传多样性分析,继而研究分子标记遗传距离与按照NC设计获得的195个杂交组合的产量及特殊配合力的相关性,探讨预测杂种优势的可能性。结果表明:(1)通过UPGMA聚类分析(图3),可将供试材料分为16个类群,并把来源不明的品种(系)划分到相应类群中,从而对这些材料进行初步鉴定。可见,AFLP分子标记是检测类内品种间遗传差异的有效方法,为水稻品种(系)亲本选配提供理论依据。(2)分子标记遗传距离与杂种产量优势、F1产量、特殊配合力之间都呈显著正相关,相关系数介于0.3235-0.7713之间。但相关程度还不足以预测杂种优势。增效座位和减效座位以及使用与杂种优势有关的QTL连锁标记位点可能提高杂种优势的预测能力,但最终解决,将依赖于杂种优势遗传机理的研究。

... 而蔡建等[34]和赵庆勇等[35]研究认为分子标记遗传距离与杂种优势呈显著正相关 ...

2009

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Genetic diversity among 30 japonica rice was studied using the simple sequence repeat(SSR) markers, and relationship between genetic distance based on SSR markers and yield heterosis was analyzed using the materials consisted of 30 parents and 200 F1 hybrids with a 20×10 diallel crossing design. The results indicated that 64 pairs of SSR primers could produce 185 steady polymorphic bands among 30 parents averaging 2.9 bands for each pair of primers. The polymorphic index content (PIC) values varied from 0.064 to 0.844 with an average of 0.380. Cluster analysis by UPGMA classified 30 parents into seven groups based on genetic similarity, which was coincident with the heterotic groups determined by their pedigree analysis. The correlations between the genetic distance and yield heterosis and hybrid performance were significant except for the total number of spikelets per panicle, and the coefficients of correlations were ranged from -0.361 to 0.359. It is suggested that the molecular markers could be used as a useful tool for assigning heterotic group and measuring genetic diversity, but not for predicting heterosis because the coefficient of relationship between the genetic distance based on SSR markers and yield heterosis was too small.

1 Institute of Food Crops, Jiangsu Academy of Agricultural Sciences/Jiangsu High Quality Rice R & D Center, Nanjing 210014, China; 2 Agricultural College, Nanjing Agricultural University, Nanjing 210095, China；*Corresponding author, E-mail: clwang@jaas.ac.cn

利用SSR分子标记对30个粳稻品种进行遗传多样性分析，继而研究分子标记遗传距离与按照NCⅡ设计获得的200个杂交组合主要产量性状杂种优势的相关性，探讨分子标记遗传距离预测杂种优势的可行性。结果表明，64对SSR引物共检测到185条多态性片段，平均每对引物2.9条，每个SSR位点的多态性信息含量指数（PIC值）变化范围为0.064~0.844，平均为0.380。以SSR标记遗传相似系数为原始数据，按UPGMA聚类方法将30个亲本材料划分为7大类群，分类结果与系谱关系基本相符。分子标记遗传距离与杂种性状平均值的相关除每穗总粒数外均达到显著或极显著水平，与杂种优势的相关均达到极显著水平，相关系数介于-0.361~0.359，说明分子标记可用于水稻杂种优势群的划分和遗传多样性分析，但相关程度还不足以预测产量杂种优势。

... 而蔡建等[34]和赵庆勇等[35]研究认为分子标记遗传距离与杂种优势呈显著正相关 ...

2014

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【Objective】 It is of great significance to identify indica or japonica accurately and predict the heterosis by genetic distance for super high-yielding rice breeding and exploring the indica-japonica inter-subspecies heterosis of rice. 【Method】The indica-japonica attribute of twelve Zheyou hybrid rice lines and their parents were identified by 19 pairs of InDel molecular markers designed on the basis of the comparative genomic DNA sequences between indica variety 9311 and japonica variety Nipponbare. According to the gene frequency of indica/japonica genomics detected by InDel molecular markers, a method was set up modified upon Lu’s report, and all the tested materials were judged for their different attributes, i.e indica, indica-cline, intermediate indica cline, intermediate japonica-cline, japonica-cline and japonica six types. Genetic distances among 13 rice parents (one sterile line and 12 restorer lines) were assessed by InDel molecular markers and calculated by using Nei’s way. The genetic similarity (GS) was reckoned by using UPGMA method to analyze the cluster of genetic similarity. Principal component analyses (PCA) were conducted using SPSS 17.0 statistical analysis software to InDel band assignments. Then, on the basis of the average values of the eigenvectors of the first and second principal components, a scatterplot was created. The control heterosis of six traits, grain yield, effective panicles per hole, spikelet number per panicle, seed setting rate, 1000-grain weight and grain weight per panicle in hybrid F1 were investigated. Furthermore, the correlations between InDel genetic distance and heterosis related to yield traits were analyzed, and possible relation to the performance of Zheyou hybrid rice lines were discussed. 【Result】 The indica-japonica attributes of the tested materials were identified and the results showed that the sterile line Zhe 04A is identified into japonica type. Eight restoring lines were thought of belonging to indica or indica-cline type, and eight combinations derived from them were judged into a range from intermediate indica-cline to japonica-cline type, it was also verified that these eight combinations were typical indica-japonica combinations. Another four restoring lines were divided into the type of japonica-cline, the four combinations from these lines were judged into japonica type. The cluster analyses were indicated that thirty-five materials could be divided into two major groups, indica group, and japonica group, at 0.350 score of GS. At 0.638 score of GS, the japonica group was divided into two sub-groups, intermediate japonica-cline and japonica /japonica-cline group. Zhe 04A and four japonica×japonica hybrids and their restoring lines fall into the latter group. Meanwhile, the indica group had eight indica type restoring lines of indica×japonica hybrids, three indica control and indica hybrid rice Shanyou 63. When all the twelve combinations were analyzed as a whole, the correlations between the InDel genetic distance and economic yield, number of grains per panicle, grain weight per panicle were significantly positively, but the InDel genetic distance was significantly negatively correlated to seed-setting rate. However, when the eight indica-japonica hybrids were tested only, the InDel genetic distance was only significantly positively correlated with grain weight per panicle and 1000-grain weight. 【Conclusion】Eight combinations from twelve tested Zheyou series combinations were verified attributing to the typical hybrids of japonica sterile line×indica restoring line. Genetic distance tested by InDel molecular marker had higher ability to predict heterosis among different breeding types of hybrids rice, especially for the heterosis of grain weight per panicle. With the increase of InDel genetic distance, the heterosis of tested combinations would be mainly expressed in enhancement of grain weight per panicle.

【目的】准确鉴定水稻材料的籼粳属性及利用遗传距离预测杂种优势，为开展水稻超高产育种和籼粳亚种间杂种优势利用提供基础。【方法】利用19对籼粳稻特异插入/缺失（Insertion/Deletion，InDel）引物，对12个浙优系列杂交稻及其双亲的籼粳属性进行了InDel分子标记鉴定，根据被检测水稻样品在多个InDel 位点上的籼型或粳型基因频率，参考卢宝荣的方法，将供试样品分别判定为“籼稻”、“偏籼”、“中间偏籼”、“中间偏粳”、“偏粳”和“粳稻”。采用Nei的方法求算13个亲本间（1个不育系和12个恢复系）的InDel遗传距离，用UPGMA法进行遗传相似性聚类，用SPSS 17.0统计分析软件对InDel条带赋值进行主成分分析, 依据第一和第二主成分的特征向量的平均分量值作平面散点图。统计杂种F1稻谷产量、每穗总粒数、每穴有效穗、结实率、千粒重及单穗重各产量性状的对照优势，以此分析InDel遗传距离与杂种优势的相关性。【结果】（1）参试材料籼粳属性的判别：不育系浙04A被鉴定为“粳稻”，8个恢复系被判定为“籼稻”或“偏籼”，由其所配组的8个杂交组合被判定在“中间偏籼”到“偏粳”之间，证实了这8个组合为典型的籼粳交组合；另外4份恢复系被判定为“偏粳”，所配组的4个杂交组合被判定为“粳稻”。（2）参试材料的聚类分析：当GS 为0.350时，35份试验材料被分为“粳稻”和“籼稻”2个主群，GS为0.638时，粳稻主群又被划分为“粳稻/偏粳”和“中间偏粳”2个亚群。其中，粳稻/偏粳亚群包括不育系浙04A、4个粳粳交恢复系及粳粳交杂种F1、3个粳稻对照种和粳稻秀水09；中间偏粳亚群包括8个籼粳交杂种F1。籼稻主群则包括8个籼粳交组合的恢复系、3个籼稻对照种和籼杂组合汕优63。（3）InDel遗传距离与杂种优势的相关性：在以粳粳交、籼粳交组合整体作为分析对象时，InDel遗传距离与稻谷产量、每穗总粒数、单穗重呈极显著正相关，与结实率呈显著负相关；仅以籼粳交组合为研究对象时，InDel遗传距离仅与单穗重和千粒重呈显著正相关。【结论】供试的12个浙优系列中的8个组合被证实为典型的粳不/籼恢型籼粳交组合，这些组合的双亲间具有较远的遗传距离；InDel遗传距离在不同类型品种间具有较好的杂种优势预测能力，可用于单穗重的优势预测，即随着InDel遗传距离的扩大，杂种优势主要体现在单穗重的增加上。

... 王友林等[36]研究认为Indel标记遗传距离在不同类型品种间具有较好的杂种优势预测能力, 可以用于单穗重的优势预测, 即随着遗传距离的扩大, 杂种优势主要体现在单穗重的增加上 ...

2006

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利用5个光温敏核不育系与40个恢复系（品种）配制了200个组合，应用SSR标记估算了这5个不育系与40个恢复系之间的遗传距离，分析了遗传距离与杂种优势的关系。结果表明：（1）不同材料、不同遗传距离范围之间，遗传距离与单株产量以及有效穗数、穗长、每穗粒敷、着粒密度、结实率、千粒重、单株产量7个性状超亲优势的相关性有很大差别，表现出很复杂的关系。（2）田丰S与父本遗传距离在0．6286～2．5257之间时，F1单株产量及其超亲优势与遗传距离极显著相关；培矮64S与父本遗传距离在0．8247～1．5315之间时，F1单株产量与遗传距离显著相关。（3）所有两系组合亲本间遗传距离在0．5333～1．5之间时，F1单株产量超亲优势与遗传距离显著相关；遗传距离在0．5333～1．0之间时，F1单株产量与遗传距离显著相关，遗传距离分别在1．0～1．5、0．5333～1．5和0．5333～2．5257之间时极显著相关。（4）另外，F1单株产量与遗传距离的相关程度普遍高于其超亲优势与遗传距离的相关程度。

... 罗小金等[37]认为不同材料、不同遗传距离范围F1的单株产量及杂种优势与遗传距离的相关性存在很大差别, 表现出很复杂的关系, 在特定遗传距离范围内存在相关性 ...

2012

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The genetic difference of 10 rice parents (5 CMS lines and 5 The genetic difference of 10 rice parents (5 CMS lines and 5 restorer lines) was analyzed by molecular markers of grain weight loci in rice. The correlation between genetic distance and grain weight and heterosis was studied by reference to the performance of 25 combinations. The results showed that the correlation between genetic distance of grain weight loci and control heterosis wassignificant(r=0.28*) ，correlation between genetic distance of grain weight loci and grain weight was significant(r=0.26*), and correlation between genetic distance of grain weight loci and female parent heterosis wasvery significantly correlated(r=0.41**). The study of molecular markers predict heterosis in hybrid rice.

利用与子粒性状相关的QTL标记分析了10个杂交水稻亲本间（5个不育系和5个恢复系）的遗传差异，结合10个亲本所配25组合的F1表现，研究了基于子粒QTL分子标记遗传距离与粒重及粒重杂种优势的相关性。结果表明分子标记遗传距离与粒重呈显著正相关（r=0.26*），与对照优势呈显著正相关（r=0.28*），与母本优势呈极显著正相关（r=0.41**）。为分子标记预测杂交水稻杂种优势研究奠定了基础。

... 游书梅等[38]研究认为基于单一性状的QTL标记的遗传距离与杂种优势存在正相关 ...

1990

0.0

... Melchinger等[39]研究认为杂种优势和遗传距离为二次曲线相关, 一定范围内杂种优势因亲本间遗传距离的增加而增强, 但超出这个范围, 杂种优势呈现减弱的现象 ...

2012

0.0

... 而在不同杂种优势群之间较远的组合中, 相关不显著[40,41,42] ...

2010

0.0

... 而在不同杂种优势群之间较远的组合中, 相关不显著[40,41,42] ...

2002

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摘要：利用AFLP、RAPD、SSR技术分析了10个恢复系和5个不育系的931个基因座,利用 15个亲本配制了50个杂交组合,在泸州和重庆2个环境下同时种植,考察了产量及其构成因素,从931个基因座中筛选出了与之相关的阳性座位、增效座位、减效座位、非环境型座位,并分析了它们与杂种产量及其构成因素间的关系.结果表明,利用所有座位计算的遗传差异与产量及其构成因素的相关性,绝大多数性状未达显著水平,不能直接用来预测产量及其构成因素.阳性座位在一定程度上可以提高相关系数,因性状不同而存在差异,在多数性状上预测产量及其构成因素还有一定难度;增效座位和减效座位可以大幅度提高相关系数,在不同的环境下也表现一致,可以用来预测产量及其构成因素;非环境型座位计算的相关系数也较高,但低于增效座位和减效座位,说明环境对产量及其构成因素有较大的影响.

... 而在不同杂种优势群之间较远的组合中, 相关不显著[40,41,42] ...

1997

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... Saghai等[43]、Zhao等[44]也认为分子标记遗传距离与杂种优势的关系受材料来源、亲本数目、亲缘关系及所用标记位点的数量等因素的影响, 故而预测性不足 ...

1999

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2008

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... 结合上述研究分析主要原因[45,46,47]: ①研究的性状及标记的数目较少, 有些性状对杂种优势的表现贡献较低或无贡献 ...

2014

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摘要：以80份杂交籼稻亲本和部分新育成品系为试验材料，采用24对籼粳特异性分子标记对其进行材料间遗传多态性的划分，计算各材料间遗传距离，并以此为依据，通过UPGMA法进行相关聚类分析。结果表明：按遗传距离值的大小可将试验材料划分为5个大群、12个亚群和28个组群，大面积生产上主要应用的亲本80％以上（保持系、恢复系）均聚集在第1大群内，分类结果和系谱分析基本吻合。结合杂交稻育种实践，群间优势普遍大于群内优势，表明利用籼粳特异分子标记在一定程度上区分杂交亲本优势群可行。

... 结合上述研究分析主要原因[45,46,47]: ①研究的性状及标记的数目较少, 有些性状对杂种优势的表现贡献较低或无贡献 ...

1996

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... 结合上述研究分析主要原因[45,46,47]: ①研究的性状及标记的数目较少, 有些性状对杂种优势的表现贡献较低或无贡献 ...

2010

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我国杂交粳稻经过30多年的研究，已取得一定的成绩，但是相对于杂交籼稻的发展而言，却显严重滞后。杂交籼稻的种植面积已占中国水稻种植总面积的57％，而杂交粳稻仅占3％。杂交粳稻还有很大的发展空间。杂交粳稻发展缓慢的原因是多方面的：一是纯系粳稻品种的不断改良，产量接近、米质优于杂交粳稻组合，让杂交粳稻的推广面临严峻的挑战；二是杂交粳稻育种中缺乏产量和米质相关性状高配合力的恢复系，进而导致杂交粳稻组合的竞争优势不强，这是其根本原因。这与早期杂交粳稻的育种基础有关。以往大量关于恢复系的配合力分析虽有助于鉴别恢复系的配合力高低和提...

... 近年来, 洪德林等[48,49,50,51,52]率先在杂交粳稻上开展亲本产量性状优异配合力的标记基因型筛选工作, 研究得出RM152-165/170和RM2439-150标记是单株日产量和单株有效穗数优异配合力效应最大的标记基因型, RM8254-120/180标记对增加穗部性状的枝梗数贡献较大, 进而为提高粳稻杂种优势预测提供了参考 ...

2011

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提高杂交粳稻竞争优势的关键是选育具有高配合力的大穗型杂交粳稻恢复系.为使配合力改良更有效,选用115对SSR引物扩增6个粳稻BT型不育系和12个恢复系的标记基因型,并按NC Ⅱ(Noah Carolina Ⅱ)遗传交配设计配制72个F1,组合,分析18个亲本的一次枝梗数、二次枝梗数和穗长3个穗部性状的配合力,结合亲本SSR分子标记数据和性状配合力数据筛选3个穗部性状优异配合力的标记基因型.结果发现:23个标记基因型与亲本3个穗部性状配合力显著相关.其中1个标记基因型与亲本3个性状配合力都相关;7个标记基因型与亲本2个性状配合力都相关;15个标记基因型与亲本单个性状配合力相关.标记基因型RM8254-120/180对3个性状的配合力效应都是正的,可使F1的每穗一次枝梗数、二次枝梗数和穗长3个性状值分别增加21.63%、21.12%和16.12%.与亲本2个性状配合力相关的7个标记基因型中有6个标记基因型对亲本的配合力效应是正值,1个标记基因型对亲本的配合力效应是负值.

2012

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To understand the types of gene action controlling yield-related traits and their mid-parental heterosis in japonica rice, we carried out quantitative trait locus (QTL) mapping by using 254 recombinant inbred lines (RILs) derived from a cross of japonica rice varieties Xiushui 79, C Bao and two backcross hybrid (BCF 1 ) populations derived from these RILs. Phenotypic values and mid-parental heterosis values ( H MP ) were investigated for eight traits in the three populations. The eight traits were plant height, growth duration, productive panicle number per plant, panicle length, spikelet number per panicle, percentage of spikelet fertility, primary branch number per panicle, and secondary branch number per panicle. A total of 58 main-effect QTLs (M- QTL ) were detected in the three populations. The percentage of phenotypic variance explained by single M-QTL ranged from 3.3% to 41.9%. Among the 58 M-QTLs detected, 45 M-QTLs (77.6%) showed additive effects, 9 M-QTLs (15.5%) showed partial-to-complete dominant effects, and 4 M-QTLs (6.9%) showed over-dominant effects. Ninety pairs of digenic epistatic QTL (E-QTL) were detected in the three populations. Among them, 44 pairs of E-QTLs were detected in RIL population, and t he percentage of phenotypic variance explained by single pair of QTL ranged from 1.7% to 8.0 % , with an average of 3.7 % . In XSBCF 1 population (Xiushui 79 as recurrent parent), 27 pairs of E-QTL were detected. Sixteen pairs of E-QTL were detected by using BCF 1 phenotypic values, and t he percentage of phenotypic variance explained by single E- QTL ranged from 12.7% to 78.5 % , with an average of 29.2 % . Eleven pairs of E-QTL were detected by using H MP values , and t he percentage of phenotypic variance explained by single E- QTL ranged from 15.0% to 71.8 % , with an average of 40.1 % . In CBBCF 1 population (C Bao as recurrent parent), 19 pairs of E-QTL were detected. Twelve pairs of E-QTL were detected by using BCF 1 phenotypic value, and t he percentage of phenotypic variance explained by single pair of E- QTL ranged from 2.7% to 64.4 % , with an average of 30.1 % . Nine pairs of E-QTL were detected by using H MP value , and t he percentage of phenotypic variance explained by single pair of E- QTL ranged from 21.7% to 64.1 % , with an average of 40.0 % . Two pairs of E-QTL were detected by using both BCF 1 phenotypic value and H MP value in CBBCF 1 population. These results indicated that epistatic effects were the primary genetic basis of heterosis in this cross in japonica rice.

为了解控制粳稻产量相关性状及其中亲优势的基因作用类型 , 利用秀堡 RIL 群体及其 2 个回交 (BCF 1 ) 群体对株高、生育期、单株有效穗数、穗长、每穗颖花数、结实率、一次枝梗数和二次枝梗数 8 个性状及其中亲杂种优势进行 QTL 定位。共检测到 58 个显著的主效 QTL (M-QTL), 单个 M-QTL 的贡献率变幅为 3.3%~41.9% 。 77.6% 的 M-QTL 表现为加性效应 , 15.5% 的 M-QTL 表现为部分或完全显性效应 , 6.9% 的 M-QTL 表现为超显性效应。共检测到 90 对显著的双基因上位性 QTL(E-QTL) 。在 RIL 群体中检测到 44 对 E-QTL, 单对 E-QTL 的贡献率变幅为 1.7%~8.0%, 平均 3.7% 。在 XSBCF 1 群体中检测到 27 对 E-QTL, 其中利用 BCF 1 表型值检测到 16 对 E-QTL, 单对 E-QTL 的贡献率变幅为 12.7%~78.5%, 平均 29.2%; 利用中亲优势值检测到 11 对 E-QTL, 单对 E-QTL 的贡献率变幅为 15.0%~71.8%, 平均 40.1% 。在 CBBCF 1 群体中检测到 19 对 E-QTL, 其中利用 BCF 1 表型值检测到 12 对 E- QTL, 单对 E-QTL 的贡献率变幅为 2.7%~64.4%, 平均 30.1%; 利用中亲优势值检测到 9 对 E-QTL, 单对 E-QTL 的贡献率变幅为 21.7%~64.1%, 平均 40.0% 。在 CBBCF 1 群体中 , 利用 BCF 1 表型值和中亲优势值都检测到的 E-QTL 有 2 对。上述结果表明上位性效应是粳稻秀堡组合杂种优势的主要遗传基础。

2013

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To understand the types of gene action controlling outcrossrelated traits and its midparental heterosis, QTL mapping of eight outcrossingrelated traits including panicle exsertion degree (PED), flag leaf length (FLL), flag leaf angle (FLA), vertical distance between panicle tip and flag leaf tip (DPFT), second leaf length (SLL), second leaf angle (SLA), vertical distance between panicle tip and second leaf tip (DPST) and length of the first internode (LFI) from the top was conducted using 254 Xiushui 79/C Bao recombinant inbred lines (RILs) and two backcross hybrid populations derived from these RILs. 45 maineffect QTLs (MQTL) were identified in the three populations. The percentage of phenotypic variance explained by each QTL ranged from 1.5% to 80.3%. Among them, 73.3% QTL showed an additive effect, 4.5% a partialtocomplete dominant effect, and 22.2% an overdominant effect. Eightytwo pairs of digenic epistatic QTL (EQTL) were detected in the 3 populations,of which 43 pairs were detected in RIL population, and the percentage of phenotypic variance explained by each pair of QTL ranged from 1.0% to 7.0%, with an average 2.7%. Sixteen pairs of EQTL were detected in XSBCF1 population. Among them, 11 pairs were detected by using XSBCF1 phenotypic value, and the percentage of phenotypic variance explained by each EQTL ranged from 11.2% to 36.8%, with an average 21.0%. Six pairs of EQTL were detected by using midparental heterosis value (HMP), and the percentage of phenotypic variance explained by each EQTL ranged from 33.1% to 76.8%, with an average 55.0%. Twentythree pairs of EQTL were detected in CBBCF1 population. Among them, 16 pairs of EQTL were detected by using BCF1 phenotypic value, and the percentage of phenotypic variance explained by each pair of EQTL ranged from 6.2% to 60.0%, with an average 24.0%. Seven pairs of EQTL were detected by using the midparental heterosis value, and the percentage of phenotypic variance explained by each pair of EQTL ranged from 21.3% to 44.4%, with an average 31.0%. These results showed that outcrossingrelated traits themselves were mainly controlled by multiple loci with additive effects, and the primary genetic basis of midparent heterosis for outcrossingrelated traits was overdominant and epistatic effect in japonica rice.

为探明控制粳稻异交相关性状及其中亲优势的基因作用类型，利用秀堡RIL群体及其2个回交(BCF1)群体对穗抽出度、剑叶长、剑叶角度、穗剑高度差、倒2叶长、倒2叶角度、穗与倒2叶（穗二）高度差和倒1节间长8个异交相关性状及其中亲优势进行QTL定位。3个群体中共检测到45个显著的主效QTL（MQTL），单个MQTL的贡献率变幅为1.5%～803%。73.3%的MQTL表现为加性效应，4.5%的MQTL表现为部分或完全显性效应，22.2%的MQTL表现为超显性效应。3个群体中共检测到82对显著的双基因上位性QTL（EQTL）。RIL群体中检测到43对EQTL，单对EQTL的贡献率变幅为1.0%～7.0%，平均2.7%。在以秀水79为父本、与秀堡RIL群体回交的后代（XSBCF1群体）中检测到16对EQTL，其中利用BCF1表型值检测到11对EQTL，单对EQTL的贡献率变幅为11.2%～36.8%，平均21.0%；利用中亲优势值检测到6对EQTL，单对EQTL的贡献率变幅为33.1%～76.8%，平均55.0%。在以C堡为父本、与秀堡RIL群体回交的后代（CBBCF1群体）中检测到23对EQTL，其中利用BCF1表型值检测到16对EQTL，单对EQTL的贡献率变幅为6.2%～60.0%，平均24.0%；利用中亲优势值检测到7对EQTL，单对EQTL的贡献率变幅为21.3%～44.4%，平均31.0%。上述结果表明，粳稻异交相关性状是由多位点控制的，基因对性状本身的作用类型以加性效应为主；粳稻异交相关性状中亲优势主要遗传基础为超显性效应和上位性效应。

2015

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2007

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An immortalized F2 population in rice derived from F1 hybrid between Zhenshan 97B and Minghui 63,was used to QTL mapping on effective panicle number per plant by using QTLNetwork-2.0.A total of eight QTLs were detected on 6 chromosomes.Both general heterosis and environmental interaction heterosis of effective panicle number per plant in two different environments for F1,F2 and F3 generations were predicted based on QTL effects.It was revealed that QTLs with epistatic effects were prominent contributors to the heterosis for effective panicle number per plant,suggesting that the method of heterosis prediction based on QTL mapping was a powerful tool for screening superior hybrids by means of molecular marker assisted selection.

以水稻杂交组合珍汕97B×明恢63所衍生的永久F2（IF2）作图群体为遗传材料，采用QTLNetwork2.0定位软件对有效穗数进行了QTL分析，共检测到8个QTL，分布于6条染色体上。同时根据有效穗数的QTL遗传效应，估算了F1、F2和F3三个杂种世代的有效穗数杂种优势，在预测普通杂种优势的同时，也对环境互作杂种优势进行了预测。上位性QTL对有效穗数杂种优势的形成有重要的贡献。表明基于QTL定位结果的杂种优势预测方法有助于分子标记辅助选育强优势组合。

... 赵彦宏等[53]研究把杂种优势精确分担到各个QTL位点上, 估算控制性状的QTL 遗传效应进而预测杂种优势 ...