利用单片段代换系研究水稻产量相关性状QTL加性及上位性效应

作物学报ACTA AGRONOMICA SINICA 2012, 38(11): 2007?2014 https://www.360docs.net/doc/894202882.html,/zwxb/ ISSN 0496-3490; CODEN TSHPA9E-mail: xbzw@https://www.360docs.net/doc/894202882.html,

DOI: 10.3724/SP.J.1006.2012.02007

利用单片段代换系研究水稻产量相关性状QTL加性及上位性效应

赵芳明1张桂权2曾瑞珍2杨正林1凌英华1桑贤春1何光华1,*

1西南大学水稻研究所 / 转基因植物与安全控制重庆市重点实验室 / 南方山地农业教育部工程研究中心, 重庆 400716; 2华南农业大学广东省植物分子育种重点实验室, 广东广州 510642

摘要: 产量及其相关性状如单株有效穗数、千粒重、穗实粒数、穗总粒数和结实率等是水稻重要的农艺性状, 了

解产量及其相关性状QTL的加性及上位性效应对以分子标记聚合育种改良水稻产量具有重要意义。本文以16个单

片段代换系及15个双片段代换系分析了水稻产量相关性状QTL的加性及上位性效应。共检出影响产量及其相关性

状的13个QTL, 包括产量性状1个、单株有效穗数1个、千粒重4个、穗实粒数4个、穗总粒数2个和结实率1个,

分布于第2、第3、第4、第7和第10染色体上。此外, 检出12对双基因互作。结果显示, 2个正向(或负向)产量性

状QTL聚合, 往往会产生负向(或正向)的上位性效应, 能否产生更大(或更小)的目标性状, 取决于双片段遗传效应(加性效应与上位效应代数和)绝对值与单片段最大加性效应绝对值的差。本研究结果对实施高产分子标记聚合育种

方法有重要参考价值。

关键词: 水稻; 单片段代换系; 产量相关性状QTL; 加性效应; 上位性效应

Epistatic and Additive Effects of QTLs for Yield-Related Traits Using Single Segment Substitution Lines of Rice (Oryza sativa L.)

ZHAO Fang-Ming1, ZHANG Gui-Quan2, ZENG Rui-Zhen2, YANG Zheng-Lin1, LING Ying-Hua1, SANG Xian-Chun1, and HE Guang-Hua1,*

1 Rice Research Institute, Southwest University / Chongqing Key Laboratory of Application and Safety Control of Genetically Modified Crops / En-gineering Research Center of South Upland Agriculture, Ministry of Education, Chongqing 400716, China;

2 Guangdong Key Laboratory of Plant Molecular Breeding, South China Agricultural University, Guangzhou 510642, China

Abstract: Yield-related traits such as panicle number per plant, thousand grain weight, number of grains per panicle, number of spikelets per panicle and seed setting rate are important agronomic traits in rice. Understanding additive and epistatic effects of QTL for yield-related traits are important to increase rice yield using method of pyramiding breeding with molecular marker as-sisted selection. In this paper, additive and epistatic effects of QTLs for rice yield-related traits were analyzed using 16 single segment substitution lines (SSSL) and 15 double segment substitution lines (DSSL). A total of thirteen QTLs for yield-related traits were identified on the chromosomes 2, 3, 4, 7, and 10 respectively, containing one for grain yield per plant (GY), one for panicle number (PN), four for thousand-grain weight (TGW), four for number of grains per panicle (NGP), two for number of spikelets per panicle (NSP) and one for seed-setting rate (SSR). Furthermore, twelve pairs of digenic interactions were detected

for yield-related traits. The results showed that pyramiding two QTLs with positive effects (or two QTLs with negative effects) often results in negative epistatic effects (or positive epistatic effects) in DSSL. Whether larger or smaller value of yield-related traits is produced lies on the difference between the absolute value of genetic effect (algebraic sum of additive and epistatic effects)

in the DSSL and the largest value of additive effect in the SSSL. These results are important to improve yield by pyramiding fa-vorable QTLs for yield-related traits.

Keywords: Rice; Single segment substitution lines; QTLs for yield-related trait; Additive effects; Epistatic effect 单产的提高始终是水稻育种的追求目标, 而产量性状是数量性状, 遗传基础复杂, 易受环境和遗

本研究由西南大学基本科研业务费专项资金(XDJK2010B011)和重庆市自然科学基金项目(CSTC, 2010BB1131)资助。

*通讯作者(Corresponding author): 何光华, E-mail: hegh@https://www.360docs.net/doc/894202882.html,

第一作者联系方式: E-mail: zhaofangming2004@https://www.360docs.net/doc/894202882.html,

Received(收稿日期): 2012-03-18; Accepted(接受日期): 2012-07-05; Published online(网络出版日期): 2012-09-10.

URL: https://www.360docs.net/doc/894202882.html,/kcms/detail/11.1809.S.20120910.1328.007.html

2008作物学报第38卷

传背景的影响。随着分子标记技术的飞速发展, 利用分子标记进行QTL作图, 将复杂的数量性状剖分为若干离散的孟德尔因子所决定的组分, 进而确定其在染色体上的位置及与其他基因的关系, 对于数量性状的逐个改良与聚合有重要意义。近年来, 利用传统作图群体(F2、F3、DH、RIL)大量研究了水稻产量性状QTL, 定位的QTL已涵盖所有12条染色体[1]。但由于QTL对遗传背景和环境影响敏感[2]以及传统作图群体个体间遗传背景差异较大, 使得QTL定位精确度不高, 且较难鉴定出效应较小的QTL[3-4]。为克服以上不足, 构建了一些减少遗传背景干扰的次级作图群体, 如染色体片段代换系(chromosome segment substitution lines, CSSLs) [5-7]和单片段代换系(single segment substitution lines, SSSLs) [8-11], 并已广泛应用于QTL定位[12-15]。单片段代换系只存在供体亲本1个代换片段的差异, 其遗传背景与受体亲本一致[8], 因而利用单片段代换系鉴定QTL, 可提高QTL定位的准确性。

随着QTL定位研究的深入, 近年来对上位性互作的研究也受到极大重视。上位性是指基因位点间的非加性遗传效应, 是偏离单个基因独立效应总和的效应。Eshed和Zamir[16]认为QTL上位性是决定表型值的重要组成。已有大量研究用传统作图群体对产量性状的基因互作进行了上位性分析[17-19], 然而上位性受遗传背景的影响更敏感[2]。利用染色体片段代换系, 既可减少遗传背景的干扰, 又可通过导入不同的遗传组成基因有目的地分析基因间的互作, 因而是研究上位性的有效手段。目前尚没有用近等基因系或单片段代换系对产量相关性状QTL 上位性的研究报道。本研究通过单片段代换系间的杂交而衍生16个次级单片段代换系和15个双片段代换系, 采用随机区组试验设计分析了水稻产量相关性状QTL的加性和上位性效应, 这些结果对于探索通过基因聚合途径提高产量具有重要的参考价值。

1材料与方法

1.1 试验材料

16个单片段代换系(SSSL)、15个双片段代换系(double segment substitution lines, DSSL)和受体亲本华粳籼74均由华南农业大学广东省植物分子育种重点实验室提供, 来源于Zhang等[8]构建的单片段代换系文库。16个SSSL包括7个初级SSSL和9个次级SSSL, 后者是由7个初级SSSL间的杂交分解得到的相互重叠的SSSL; DSSL是由7个初级SSSL间的杂交衍生聚合而成的双片段代换系。材料见表1。

1.2试验方法

采取随机区组试验设计, 每小区种40株, 3次重复。于2008年3月8日种植于西南大学实验地, 4月15日移栽。每小区栽4行, 每行10株, 行、株距分别为26.4 cm和16.5 cm。常规管理。每个小区成熟后分单株收取中间10株, 分别考察单株有效穗数、穗总粒数、穗实粒数、千粒重、结实率和单株产量, 然后以10株的平均数计为该小区的各性状值以备下一步统计分析。

1.3 QTL加性及上位性效应分析

1.3.1 加性效应以SPSS1

2.0统计软件, 对16个SSSL和华粳籼74的3个小区单株产量及其相关性状进行ANOVA分析和Duncan’s多重比较, 如果某一性状与华粳籼74在P<0.01水平上差异显著, 即认为单片段代换系代换片段上存在QTL。同时, 为了提高置信度, 本文还进一步作了单次t检验, 并提供了各QTL的概率值。参照Eshed和Zamir的方法[3]估算各个QTL的加性效应。加性效应值=(单片段代换系的表型值?华粳籼74的表型值)/2。

1.3.2 上位性效应参照Eshed和Zamir的分析方法[16], 其遗传模型为Y =μ + SLH a + SLH b + SLH a×SLH b + ε, 其中, Y表示双片段代换系的某性状遗传值, μ表示受体华粳籼74的某性状平均值, ε表示试验误差, SLH a和SLH b分别表示单片段代换系SSSL(a)和SSSL(b)存在或缺失时的某性状加性遗传效应, SLH a×SLH b表示两代换片段间的某性状上位性互作效应, 以(HJX74+DSSLab)?(SSSLa+ SSSLb)的一半性状表型值进行估计, 其中, 4个参数分别表示受体华粳74、双片段代换系、单片段代换系a和b的性状平均值。上位性效应分析分两步进行, 首先以SPSS1

2.0统计软件选出在某一性状上与华粳籼74差异极显著的DSSL, 然后以t测验检测(HJX74 + DSSLab)与(SSSLa + SSSLb)的显著性, 当二者P<0.01时, 认为两代换片段间存在上位性互作效应。

1.3.3 QTL的命名QTL名称中前面的字母为鉴定性状的英文词首的缩写, 数字为QTL所在染色体的编号, 如果在几个单片段代换系的重叠区段上都检出某一性状的QTL, 参照Paterson等[20]的代换

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2009

表1 所用单片段代换系及双片段代换系 Table 1 SSSL and DSSL used in the experiment

编号 Code 供体来源 Donor

染色体

Chr. 代换片段

Substitution segment 长度 Length (kb)

S1 2 RM3542---RM138-PSM125---Long arm end 39.35 S2 美国茉莉香 American jasmine 3 PSM377---RM6146-RM3646-RM16-PSM127---RM5626 729.14

S3

Basmati 370

4 RM2811---RM401---RM3317

570.52

S4 4 RM5320---RM451---RM303 37.26 S5 4 RM5320---RM451-RM303---RM317 71.01

S6 4 RM5320---RM451-RM303-RM317---RM3217 145.03 S7 4 RM317---RM3217-RM3276---RM5473 137.40 S8

巴西陆稻 IAPAR9

4 RM5320---RM451-RM303-RM317-RM3217-RM3276---RM5473 282.41

S9 6 RM508---RM170-RM190-RM587-RM510---RM225 295.86

S10 6 RM508---RM170---RM190 66.15

S11 6 RM190---RM587-RM510---RM225 109.59 S12 6 RM508---RM170-RM190---RM587 112.73

S13 美国茉莉香 American jasmine 6 RM170---RM190-RM587-RM510---RM225 156.16

S14 Amol 3 7 RM3826---RM336---RM3534 97.44 S15 Lemont 10 RM596---RM258---PSM169 252.83

S16 Lemont

10 RM596---RM258-PSM169---RM147 399.42

D1 3, 6 (Pyramid of S2 and S10) D2 3, 6 (Pyramid of S2 and S12) D3 3, 6 (Pyramid of S2 and S11) D 4 美国茉莉香 American jasmine

3, 6 (Pyramid of S2 and S9) D 5 6, 7 (Pyramid of S10 and S14) D6 Amol 3, 美国茉莉香

Amol 3, American jasmine 6, 7 (Pyramid S9 and S14) D7 6, 2 (Pyramid of S12 and S1) D8 6, 2 (Pyramid of S9 and S1) D9 IRAT261, 美国茉莉香, IRAT261, American jasmine 6, 2 (Pyramid of S11 and S1) D10 3, 4 (Pyramid of S2 and S7) D11 3, 4 (Pyramid of S2 and S8) D12 3, 4 (Pyramid of S2 and S6) D13 6, 4 (Pyramid of S12 and S8) D14 美国茉莉香, 巴西陆稻

American jasmine, IAPAR9 6, 4 (Pyramid of S9 and S8) D15

Lemont, Basmati 370 10, 4

(Pyramid of S16 and S3)

三连线末端为代换片段两侧标记, 三连线区表示重组可能发生的区段, 中间的短连线表示检出的代换片段。

The single hyphen in the middle of markers shows the substitution segment, while the end markers of the triadic hyphens are side markers of substitution segment, in which segments recombination might appear.

作图法命名为相同的名称。 2 结果与分析

2.1 单片段代换系上检出的水稻产量及其相关性状QTL

通过对16个单片段代换系进行产量及其相关性状的QTL 鉴定, 结果在7个单片段代换系上检出影响产量及其相关性状的13个QTL, 包括单株产量的1个, 单株有效穗数1个, 千粒重4个, 穗实粒数

4个, 穗总粒数2个, 结实率1个, 分布于第2、第3、第4、第7和第10染色体上(表2)。

从表2分析可知, 同一产量相关性状QTL 分布于不同的水稻单片段代换系中。如千粒重QTL 能在S2、S3、S7和S16中被检测到, 穗总粒数QTL 能够在S3和S16中被检测到, 穗实粒数QTL 能够在S1、S3、S14和S16中被检测到, 说明水稻产量相关性状QTL 分布于水稻基因组的不同区域。

进一步分析发现, 多数产量相关性状QTL 往

2010

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表2 在单片段代换系上检出的水稻产量相关性状QTL

Table 2 QTL for yield-related traits detected using SSSL of rice

编号

Code 染色体 Chr. 代换区间 Substitution interval

性状 Trait QTL 平均数±标准差 Mean ± SD A P Recipient (Huajingxian 74)

35.92±1.66 S3 4 RM401 单株产量 Grain yield per plant (g)

qGY4 23.01±1.07 –6.46 7.68×10?4

Huajingxian 74

9.20±0.28

S16 10 RM258–PSM169 单株有效穗

Panicle number

qPN10 13.97±0.34 2.39 1.08×10?4

Huajingxian 74

20.20±0.31

S2 3 RM6146–PSM127 qTGW3 23.16±0.34 1.48 8.01×10?4S3 4 RM401

qTGW4.115.53±0.24 –2.34 7.33×10?5S7 4 RM3217–RM3276 qTGW4.2

23.49±0.18 1.65 1.99×10?4S8 4 RM451–RM3276 qTGW4.222.70±0.08 1.25 3.68×10?4S16 10 RM258–PSM169 千粒重 1000-grain weight (g)

qTGW1017.33±0.08 –1.44 2.16×10?4Huajingxian 74

167.07±3.26

S1 2 RM138–PSM125

qNGP2 127.78±4.96 –19.65 7.27×10?4S3 4 RM401 qNGP4 101.57±5.76 –32.75 1.51×10?4S14 7 RM336 qNGP7 125.57±2.04 –20.75 1.08×10?4S16 10 RM258–PSM169 穗实粒数

Number of grains per

panicle

qNGP10 105.62±4.08 –30.73 7.67×10?5Huajingxian 74

197.19±5.05

S3 4 RM401

qNSP4 120.32±6.15 –38.44 1.66×10?4

S16 10 RM258–PSM169 穗总粒数

Number of spikelets

per panicle

qNSP10 139.80±4.08 –28.70 2.36×10?4Huajingxian 74

84.61±1.38

S14 7 RM336 结实率

Seed-setting rate (%)

qSSR7

73.13±0.68 –5.74 4.58×10?4

A 为加性效应, “正效应”表示增加受体性状值, “负效应”表示减小受体表型值。

A: additive effect; “positive effect” indicates increasing trait values of Huajingxian 74; “negative effect” indicates decreasing trait values of Huajingxian 74.

往成簇分布。如单株产量QTL (qGY4)、千粒重QTL (qTGW4.1)、穗实粒数QTL (qNGP4)和穗总粒数QTL (qNSP4)均在单片段代换系S3上被检测到, 加性效应分别使华粳籼74的相应性状减少12.91 g 、4.67 g 、65.50粒和76.87粒; 穗实粒数QTL (qNGP7)和结实粒QTL (qSSR7)均在单片段代换系S14上被检测到; 单株有效穗数QTL (qPN10)、千粒重QTL (qTGW- 10)、穗实粒数QTL (qNGP10)和穗总粒数QTL (qNSP10)都在单片段代换系S16上被检测到。

2.2 双片段代换系上检出的水稻产量及其相关性状QTL 的上位性效应

由于单片段代换系和双片段代换系是纯合体, 因而理论上单片段代换系的遗传效应只有单片段QTL 的加性效应。双片段代换系的遗传效应理论上是2个代换片段QTL 的加性效应及代换片段QTL 间上位性效应的代数和。本研究利用15个双片段代换系共检出影响产量相关性状的双基因互作12对(表3), 包括单株产量的1对、单株有效穗的1对、千粒重的5对、穗实粒数的1对、穗总粒数的1对

和结实率的3对。从互作方式分析, 4对为2个位点均表现显著加性效应的上位性互作, 3对为仅一个位点表现显著加性效应的上位性互作, 5对为2个位点均没有显著加性效应的上位性互作。从双基因互作效应分析, 在D10中, 第3染色体上加性效应为1.48的粒重QTL (qTGW3)和第4染色体上加性效应为1.65的粒重QTL (qTGW4.2)聚合后, 二者产生的上位性效应为?1.61。这说明双代换片段会使华粳籼74的千粒重在遗传上增加(1.48+1.65?1.61) g, 即 1.52 g, 比单个qTGW4.2可增加千粒重1.65 g 的效应值小, 比单个qTGW3增加千粒重1.48 g 的效应值大, 但更偏向于 1.48 (qTGW3), 没有使籽粒更大, 进一步揭示了聚合2个大粒QTL, 往往产生负向的上位性效应, 双片段代换系的籽粒能否更大, 取决于双片段遗传效应绝对值与单片段最大加性效应绝对值的差, 若为正值, 表明可产生比单片段更大的籽粒; 若为负值, 则表明籽粒不会大于单片段的籽粒。在D15中, 第10染色体上加性效应为?1.44的千粒重QTL (qTGW10)和第4染色体上加性效应为?2.34的千粒

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赵芳明等: 利用单片段代换系研究水稻产量相关性状QTL 加性及上位性效应 2011

重QTL (qTGW4.1)聚合后, 二者产生的上位性效应为1.51, 说明双代换片段会使华粳籼74的千粒重在遗传上减少(?1.44?2.34+1.51) g, 即2.27 g, 由于双代换片段千粒重效应绝对值2.27 g 小于单片段最大加性效应绝对值2.34 g, 这2个小粒QTL 聚合没有产生更小的籽粒。在D15中, 第10染色体上加性效应为?30.73的穗实粒数QTL (qNGP-10)和第4染色

体上加性效应为?32.75的穗实粒数QTL (qNGP-4)聚合后, 二者产生的上位性效应为33.28, 说明双代换片段会使华粳籼74的穗实粒数在遗传上减少(?30.73 ?32.75+33.28)粒, 即30.20粒, 由于双代换片段穗实粒数遗传效应绝对值30.20大于单片段最大加性效应绝对值32.75, 2个少粒QTL 聚合没有产生比单片段更少的籽粒。

表3 双片段代换系上检出的水稻产量相关性状QTL 上位性效应

Table 3 Epistatic effects between QTLs for yield-related traits detected on the double segment substitution lines of rice 编码 Code

染色体 Chr.

代换区间

Substitution interval (i )

染色体 Chr.

代换区间

Substitution interval (j )

性状 Trait

A i 1)

A j 2)

AA ij 3)

P

D15 10 RM258–PSM169(S16) 4 RM401(S3) 单株产量GY (g) –6.46 2.94 8.81×10?3D15 10 RM258–PSM169(S16) 4 RM401(S3) 有效穗PN

2.39 –2.20 8.80×10?3D3 3 RM6146–PSM127(S2) 6 RM587–RM510(S11)

1.48 –0.90 4.11×10?3D6 6 RM170–RM510(S9) 7 RM336(S14)

1.05 8.92×10?3D9 6 RM587–RM510(S11) 2 RM138–PSM125(S1) 0.97 1.15×10?3D10 3 RM6146–PSM127(S2) 4 RM3217–RM3276(S7) 1.48 1.65 –1.61 3.10×10?4D15 10 RM258–PSM169(S16) 4 RM401(S3) 千粒重TGW (g)–1.44 –

2.34 1.51

3.71×10?4

D15 10 RM258–PSM169(S16) 4 RM401(S3) 实粒数NGP –30.73 –32.75 33.28 6.99×10?4D15 10 RM258–PSM169(S16) 4 RM401(S3) 总粒数NSP

–28.70 –38.44 29.73 2.01×10?3

D1 3 RM6146–PSM127(S2) 6 RM170(S10)

–2.08 3.12×10?3D2 3 RM6146–PSM127(S2) 6 RM170–RM190(S12) –3.18 1.73×10?3D4 3 RM6146–PSM127(S2) 6 RM170–RM510(S13)

结实率SSR (%) –2.95 3.31×10?5

1)

“i ”代换片段加性效应; 2) “j ”代换片段加性效应; 3) 双片段上位性效应。“A i 和A j 中正效应”表示增加华粳籼74性状值; “A i 和A j

中负效应”表示减小华粳籼74性状值。“AA ij 中正效应”表示2个位点间互作基因型与具有正效应的加性基因型方向相同。

1)

additive effect of “i ” substitution segment; 2) additive effect of “j ” substitution segment; 3) the epistatic effect of additive by additive

interaction between substitution segments i and j . “Positive effect in A i and A j ” indicates the increase of trait values in Huajingxian 74; “Posi-tive value in AA ij ” indicates that the two-locus interaction genotypes are syntropic with the additive genotype with positive effect. GY: grain yield per plant; PN: panicle number per plant; TGW: 1000-grain weight; NGP: number of grains per panicle; NSP: number of spikelets per panicle; SSR: seed-setting rate.

3 讨论

利用单片段代换系进行QTL 定位较传统QTL 分析方法具有明显的优势, 其准确度和灵敏度较高, 定位结果实用性强, 可直接用于育种实践[3,8,21]。本研究在7个单片段代换系上检出了13个影响产量及其相关性状的QTL, 平均每个单片段代换系上检出1.85个, 再次表明用单片段代换系鉴定QTL 的高效性。与前人研究[22-44]比较, 本研究结实率QTL (qSSR7)与徐华山等[29]定位的结实率QTL (pss-7)位于相近的染色体区段; 千粒重QTL (qTGW3)与Zhao 等[30]用单片段代换系定位的粒重QTL (Gwt-3-1)、Brondani 等[31]定位的HGW1、

Li 等[32]定位的qGW3-3及已克隆的千粒重QTL (GS3)[33], 是同一个主效QTL, 而且该基因已成功应用于以华粳籼74为背景的分子聚合育种[34]; 千粒重QTL qTGW4.2与Fu

等[35]定位的Kgw4-1、

Li 等[32]定位的qGW4-1在相近的染色体区段。本研究检出的其他10个产量相关性状QTL 尚未被报道, 可能是新的QTL, 这些结果将为利用基因聚合手段改良水稻产量奠定一定基础。

上位性互作是作物产量及其构成的重要遗传组成。利用传统作图群体研究发现, 水稻产量性状的基因位点间普遍存在上位性互作, 有些是有显著加性效应的QTL 间互作, 有些是一个有显著加性效应的QTL 与一个没有显著加性效应的位点间互作, 还有一些是2个都没有显著加性效应的位点间互 作[17-19,25,45]。但这些研究无法有目的地揭示2个QTL 间的上位性互作。通过构建水稻单片段代换系, 并利用含有目标性状QTL 的单片段代换系间杂交构建双片段代换系, 即可实现目标QTL 间的上位性互作研究。本研究利用15对水稻双片段代换系检出12对产量相关性状的QTL 间互作, 其中4对为2个

2012作物学报第38卷

位点均表现显著加性效应的上位性互作, 3对为仅一个位点表现显著加性效应的上位性互作, 5对为2个位点均没有显著加性效应的上位性互作, 互作方式与前人研究相同[17-19,25,45]。但本研究进一步通过分析2个大粒、小粒和少粒QTL的聚合, 发现2个大粒QTL聚合, 往往产生负向的上位性效应, 2个小粒或少粒QTL聚合往往产生正向的上位性效应, 聚合能否产生理想的效果, 取决于双片段加性效应与上位效应代数和的绝对值与单片段最大加性效应绝对值的差。这反映了由于QTL间上位性互作的普遍发生, 另一QTL的导入会导致单个QTL的加性作用相对减弱。Eshed和Zamir [16]的研究也表明, 随着QTL 导入的增多, QTL效应的加性作用明显减弱。尽管存在这样的上位性模式, 但基于相同背景的基因聚合仍是有效的。Gur和Zamir [46]以番茄导入系聚合了3个番茄有利产量性状QTL, 发现聚合系的产量高于对照50%, 证实了单片段代换系是研究上位性和实现基因聚合的理想群体。

4结论

利用单片段代换系群体, 共鉴定出13个影响产量及其相关性状QTL和12对产量相关性状双基因互作。2个携带大粒或小粒QTL的水稻单片段代换系聚合没有产生更大或更小粒的籽粒, 而是偏向其中之一。2个含有少粒QTL的单片段代换系聚合也没有产生更少的籽粒。聚合能否最终取得理想的效果, 主要取决于双片段遗传效应(加性效应与上位效应代数和)绝对值与单片段最大加性效应绝对值的差。

References

[1] Xing Y Z, Zhang Q F. Genitic and molecular bases of rice yield.

Annu Rev Plant Biol, 2010, 61: 421–442

[2] Liao C Y, Wu P, Hu B, Yi K K. Effects of genetic background

and environment on QTL and epistasis for rice (Oryza sativa L.) panicle number. Theor Appl Genet, 2001, 103: 104–111

[3] Eshed Y, Zamir D. An introgression line population of Lycoper-

sicon pennellii in the cultivated tomato enables the identification and fine mapping of yield-associated QTL. Genetics, 1995, 141: 1147–1162

[4] Monna L, Lin H X, Kojima S, Sasaki T, Yano M. Genetic dissec-

tion of a genomic region for a quantitative trait locus, Hd3, into two loci, Hd3a and Hd3b, controlling heading date in rice. Theor Appl Genet, 2002, 104: 772–778

[5] Kubo T, Nakamura K, Yoshimura A. Development of a series of

indica chromosome segment substitution lines in japonica back-

ground of rice. Rice Genet Newsl, 1999, 16: 104–106

[6] Ebitani T, Takeuchi Y, Nonoue Y, Yamamoto T, Takeuchi K, Yano

M. Construction and evaluation of chromosome segment substi-

tution lines carrying overlapping chromosome segments of indica

rice cultivar ‘Kasalath’ in a genetic background of japonica elite cultivar ‘Koshihikari’. Breed Sci, 2005, 55: 65–73

[7] Wan J-L(万建林), Zhai H-Q(翟虎渠), Wan J-M(万建民), Yasui

H(安井秀), Yoshimura A(吉村淳). Mapping QTL for traits as-

sociated with resistance to ferrous iron toxicity in rice (Oryza sa-

tiva L.) using japonica chromosome segment substitution lines.

Acta Genet Sin (遗传学报), 2003, 30(10): 893–898 (in Chinese with English abstract)

[8] Zhang G Q, Zeng R Z, Zhang Z M, Ding X H, Li W T, Liu G M,

He F H, Tulukdar A, Huang C F, Xi Z Y, Qin L J, Shi J Q, Zhao

F M, Feng M J, Shan Z L, Chen L, Guo X Q, Zhu H T, Lu Y G.

The construction of a library of single segment substitution lines in rice (Oryza sativa L.). Rice Genet Newsl, 2004, 21: 85–87 [9] Xi Z Y, He F H, Zeng R Z, Zhang Z M, Ding X H, Li W T,

Zhang G Q. Development of a wide population of chromosome single segment substitution lines (SSSL) in the genetic back-

ground of an elite cultivar in rice (Oryza sativa L.). Genome, 2006, 49: 476–484

[10] He F-H(何风华), Xi Z-Y(席章营), Zeng R-Z(曾瑞珍), Zhang

G-Q(张桂权). Developing single segment substitution lines (SSSL) in rice (Oryza sativa L.) using advanced backcrosses and MAS. Acta Genet Sin (遗传学报), 2005, 32(8): 825–831 (in Chinese with English abstract)

[11] Zeng R-Z(曾瑞珍), Tulukdar A, Liu F(刘芳), Zhang G-Q(张桂

权). Mapping of the QTL controlling grain shape in rice using single segment substitution lines. Sci Agric Sin (中国农业科学), 2006, 39(4): 647–654 (in Chinese with English abstract)

[12] Liu G F, Zhu H T, Liu S W, Zeng R Z, Zhang Z M, Li W T, Ding

X H, Zhao F M, Zhang G Q. Unconditional and conditional QTL mapping for the developmental behavior of tiller number in rice (Oryza sativa L.). Genetica, 2010, 138: 885–893

[13] Liu G F, Zeng R Z, Zhu H T, Zhang Z M, Ding X H, Zhao F M,

Li W T, Zhang G Q. Dynamic expression of nine QTLs for tiller number detected with single segment substitution lines in rice.

Theor Appl Genet, 2009, 118: 443–453

[14] Zhao F M, Liu G F, Zhu H T, Ding X H, Zeng R Z, Zhang Z M,

Li W T, Zhang G Q. Unconditional and conditional QTL map-

ping for tiller number at various stages by using single segment substitution lines in rice (Oryza sativa L.). Sci Agric Sin, 2008, 7:

第11期赵芳明等: 利用单片段代换系研究水稻产量相关性状QTL加性及上位性效应2013

257–265

[15] Zhao F-M(赵芳明), Zhang G-Q(张桂权), Zeng R-Z(曾瑞珍),

Yang Z-L(杨正林), Zhu H-T(朱海涛), Zhong B-Q(钟秉强), Ling Y-H(凌英华), He G-H(何光华). Additive effects and epis-tasis effects of QTL for plant height and its components using single segment substitution lines (SSSLs) in rice. Acta Agron Sin (作物学报), 2009, 35(1): 48–56 (in Chinese with English ab-stract)

[16] Eshed Y, Zamir D. Less-than-additive epistatic interactions of

quantitative trait loci in tomato. Genetics, 1996, 143: 1807–1817 [17] Zhuang J Y, Fan Y Y, Rao Z M, Wu J L, Xia Y W, Zheng K L.

Analysis on additive effects and analysis on additive effects and additive-by-additive epistatic effects of QTLs for yield traits in a recombinant inbred line population of rice. Theor Appl Genet, 2002, 105: 1137–1145

[18] Yu S B, Li J X, Xu C G, Tan Y F, Gao Y J, Li X H, Zhang Q F,

Saghai Maroof M A. importance of epistasis as the genetic basis of heterosis in an elite rice hybrid. Proc Natl Acad Sci USA, 1997, 94: 9226–9231

[19] Septiningsih E M, Prasetiyono J, Lubis E, Tai T H, Tjubaryat T,

Moeljopawiro S, McCouch S R. Identification of quantitative trait loci for yield and yield components in an advanced back-cross population derived from the Oryza sativa variety IR64 and the wild relative O. rufipogon. Theor Appl Genet, 2003, 107: 1419–1432

[20] Paterson A H, Deverna J W, Lanini B, Tanksley S D. Fine map-

ping of quantitative trait loci using selected overlapping recom-binant chromosomes in an interspecies cross of tomato. Genetics, 1990, 124: 735–742

[21] Zamir D. Improving plant breeding with exotic genetic libraries.

Nat Rev Genet, 2001, 2: 983–989

[22] Hittalmani S, Shashidhar H E, Bagali P G, Huang N, 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

[23] Mebrouk B, Zhu J, Yan J Q, He C X, Wu P. QTL analysis of ge-

netic main effects and genotype (environment interaction effects for yield components in rice (Oryza sativa L.). J Zhejiang Univ (Agric & Life Sci), 2008, 28: 127–135

[24] Moncada P, Martinez C P, Borrero J, Chatel M, Gauch H, Gui-

maraes E P, Tohme J, McCouch S R. Quantitative trait loci for yield and yield components in an Oryza sativa × Oryza rufipogon BC2F2 population evaluated in an upland environment. Theor Appl Genet, 2001, 102: 41–52 [25] Xing Y Z, Tan Y F, Hua J P, Sun C G, Xu C G, Zhang Q. Char-

acterization of the main effects, epistatic effects and their environmental interactions of QTL on the genetic basis of yield

traits in rice. Theor Appl Genet, 2002, 105: 248–257

[26] Tanabe K, Miedal M, Ashikari M, Kitano H, Iwasaki Y. Mapping

of Small and Round Seed 1 gene in rice. Rice Genet Newsl, 2006,

23: 44–47

[27] Jiang G H, Xu C G, Li X H, He Y Q. Characterization of the ge-

netic basis for yield and its component traits of rice revealed by

doubled haploid population. Acta Genet Sin, 2004, 31: 63–72 [28] Cao L-Y(曹立勇), Zhan X-D(占小登), Zhuang J-Y(庄杰云),

Zheng K-L(郑康乐), Cheng S-H(程式华). QTL mapping and

epistasis analysis for yield componentsin a RIL population of rice

(Oryza sativa L. subsp. indica). Sci Agric Sin(中国农业科学),

2003, 36(11): 1241–1247 (in Chinese with English abstract) [29] Xu H-S(徐华山), Sun Y-J(孙永建), Zhou H-J(周红菊), Yu

S-B(余四斌). Development and characterization of contiguous

segment substitution lines with background of an elite restorer

line. Acta Agron Sin (作物学报), 2007, 33(6): 979–986 (in Chi-

nese with English abstract)

[30] Zhao F-M(赵芳明), Zhu H-T(朱海涛), Ding X-H(丁效华), Zeng

R-Z(曾瑞珍), Zhang Z-M(张泽民), Li W-T(李文涛), Zhang G-Q(张桂权). Detection of QTLs for important agronomic traits

and analysis of their stabilities using SSSLs in rice. Sci Agric Sin

(中国农业科学), 2007, 40(3): 447–456 (in Chinese with English

abstract)

[31] Brondani C, Rangel P H N, Brondani R P V, Ferreira M E. QTL

mapping and introgression of yield-related traits from Oryza

glumaepatula to cultivated rice (Oryza sativa) using microsatellite markers. Theor Appl Genet, 2002, 104: 1192–1203 [32] Li D J, Sun C Q, Fu Y C, Li C, Zhu Z F, Chen L, Cai H W, Wang X

K. Identification and mapping of genes for improving yield from

Chinese common wild rice (O. rufipogon Griff.) using advanced

backcross QTL analysis. Chin Sci Bull, 2002, 47: 1533–1537

[33] Fan C, Xing Y, Mao H, Lu T, Han B, Xu C, Li X, Zhang Q. GS3,

a major QTL for grain length and weight and minor QTL for

grain width and thickness in rice, encodes a putative transmembrane protein. Theor Appl Genet, 2006, 112: 1164–1171

[34] Yang T, Zeng R, Zhu H, Chen L, Zhang Z, Ding X, Li W, Zhang

G. Effect of grain length gene GS3 in pyramiding breeding of rice.

Mol Plant Breed, 2010, 1: 59–66

[35] Fu Q, Zhang P, Tan L, Zhu Z, Ma D, Fu Y, Zhan X, Cai H, Sun C.

Analysis of QTLs for yield-related traits in Yuanjiang common

wild rice (Oryza rufipogon Griff.). J Genet Genomics, 2010, 37:

147–157

2014作物学报第38卷

[36] Yao G-X(姚国新), Li Z-C(李自超), Li J-J(李金杰), Zhang Q(张

强), Hu G-L(胡广隆), Chen C(陈超), Tang B(汤波), Zhang H-L(张洪亮). Mapping QTLs for grain weight and shape using four sister near isogenic lines in rice (Oryza sativa L.). Acta Agron Sin (作物学报), 2010, 36(8): 1310–1317 (in Chinese with English abstract)

[37] Song X J, Huang W, Shi M, Zhu M Z, Lin H X. A QTL for rice

grain width and weight encodes a previously unknown RING- type E3 ubiquitin ligase. Nat Genet, 2007, 39: 623–630

[38] Liu M-W(刘明伟), Liu Y(刘勇), Wang S-Q(王世全), Deng

Q-M(邓其明), Li P(李平). Genetic mapping of the dominant minute grain gene Mi3(t) in rice (Oryza sativa). Chin J Rice Sci (中国水稻科学), 2005, 19(6): 511–515 (in Chinese with English abstract)

[39] Chen B-X(嬬

陈冰), Shi Y-Y(石英尧), Cui J-T(崔金腾), Qian Y-L(钱益亮), Liu H-Y(刘海燕), Zhang L-K(张力科), Wang H(王辉), Gao Y-M(高用明), Zhu L-H(朱苓华), Li Z-K(黎志康).

QTL detection of grain size and shape with BC2F2 advanced backcross population of rice (Oryza sativa L.). Acta Agron Sin (作物学报), 2008, 34(8): 1299–1307 (in Chinese with English abstract)

[40] Cheng G-P(程桂平), Feng J-H(冯九焕), Liang G-H(梁国华),

Liu X-D(刘向东), Li J-Q(李金泉). Identification of QTLs for agronomic traits associated with yield in a BC2F2 population be-tween Oryza sativa and Oryza rufipogon. Chin J Rice Sci (中国

水稻科学), 2006, 20(5): 553–556 (in Chinese with English ab-

stract)

[41] Guo Y-M(郭咏梅), Mu P(穆平), Liu J-F(刘家富), Li Z-C(李自

超), Lu Y-X(卢义宣). Correlation analysis and QTL mapping of grain shape and grain weight in rice under upland and lowland environments. Acta Agron Sin(作物学报), 2007, 33(1): 50–56 (in Chinese with English abstract)

[42] Zhu W-Y(朱文银), Wang C-L(王才林), Yang L-Q(杨连群).

Analysis of substituted segments in five advanced backcross substitution lines in an indica background in rice (Oryza sativa).

Chin Bull Bot(植物学报), 2009, 44(6): 666–672 (in Chinese with English abstract)

[43] Wang B(汪斌), Lan T(兰涛), Wu W-R(吴为人). Identification of

QTL underlying grain traits in rice using SSLP linkage map.

Fujian J Agric Sci(福建农业学报), 2003, 18(1): 11–15 (in Chinese with English abstract)

[44] Yuan A-P(袁爱平), Cao L-Y(曹立勇), Zhuang J-Y(庄杰云), Li

R-Z(李润植), Zheng K-L(郑康乐), Zhu J(朱军), Cheng S-H(程

式华). Analysis of additive and AE interaction effects of QTLs controlling plant height, heading date and panicle number in rice (Oryza sativa L.). Acta Genet Sin(遗传学报), 2003, 30(10): 899–906 (in Chinese with English abstract)

[45] Ma X Q, Tang J H, Teng W T, Yan J B, Meng Y J, Li J S.

Epistatic interaction is an important genetic basis of grain yield and its components in maize. Mol Breed, 2007, 20: 41–51 [46] Gur A, Zamir D. Unused natural variation can lift yield barriers

in plant breeding. PLoS Biol, 2004, 2: 1610–1615