中国水稻科学 ›› 2021, Vol. 35 ›› Issue (4): 359-372.DOI: 10.16819/j.1001-7216.2021.201205
杜成兴#, 张华丽#, 戴冬青, 吴明月, 梁敏敏, 陈俊宇*(), 马良勇*()
收稿日期:
2020-12-08
修回日期:
2021-01-26
出版日期:
2021-07-10
发布日期:
2021-07-10
通讯作者:
陈俊宇,马良勇
作者简介:
#共同第一作者
基金资助:
Chengxing DU#, Huali ZHANG#, Dongqing DAI, Mingyue WU, Minmin LIANG, Junyu CHEN*(), Liangyong MA*()
Received:
2020-12-08
Revised:
2021-01-26
Online:
2021-07-10
Published:
2021-07-10
Contact:
Junyu CHEN, Liangyong MA
About author:
#These authors contributed equally to this work
摘要:
【目的】粒重粒形对水稻的产量和品质均有重要的影响。本研究通过开展水稻粒重粒形QTL的初步定位,并对新鉴定的第1染色体长臂qTGW1.2/qGL1.2区间进行验证,旨在进一步揭示水稻粒重粒形的遗传调控机制。【方法】以大粒的FM9为父本,小粒的EFT为母本,配组衍生遗传群体,先后获得包含277个株系的F2:3群体和211个株系的重组自交系群体(Recombinant Inbred Lines, RILs),测定千粒重、粒长和粒宽,采用完备区间作图法进行QTL初定位;针对新鉴定的qTGW1.2/qGL1.2区间,筛选2个剩余杂合体单株,自交衍生分离群体,开展QTL效应验证。【结果】初定位分析共检测到35个调控千粒重、粒长和粒宽的QTL,其中,11个能同时在两个群体中被检测到,18个仅在F2:3群体中被检测到,6个仅在RIL群体中被检测到;应用两个剩余杂合体衍生的两套分离群体验证了新鉴定的qTGW1.2/qGL1.2区间对千粒重和粒长的效应,并观察到颖壳细胞长度的显著变化。通过qPCR分析,观察到与细胞周期、生长素代谢和粒形相关基因表达发生了显著变化。【结论】初步定位的35个QTL以及验证的qTGW1.2/qGL1.2有利于进一步揭示水稻粒重粒形的遗传控制基础,也为后续的基因克隆及分子标记辅助选择奠定了基础。
杜成兴, 张华丽, 戴冬青, 吴明月, 梁敏敏, 陈俊宇, 马良勇. 水稻粒重粒形QTL的定位及qTGW1.2/qGL1.2的验证[J]. 中国水稻科学, 2021, 35(4): 359-372.
Chengxing DU, Huali ZHANG, Dongqing DAI, Mingyue WU, Minmin LIANG, Junyu CHEN, Liangyong MA. QTL Analysis for Grain Weight and Shape and Validation of qTGW1.2/qGL1.2 [J]. Chinese Journal OF Rice Science, 2021, 35(4): 359-372.
图2 两个剩余杂合体单株RHL17和RHL124与目标qTGW1.2/qGL1.2区间的基因型组成
Fig. 2. Schematic genotypes of two residual heterozygotes RHL17 and RHL124 and the targeted region of qTGW1.2/qGL1.2.
图3 两个亲本材料EFT和FM9的千粒重、粒长和粒宽的比较**表示在 0.01水平上显著。
Fig. 3. Comparison of 1000-grain weight, grain length and grain width of the parents EFT and FM9.** Significant at 0.01 level.
群体 Population | 性状Trait | F2:3-TGW | F2:3-GL | F2:3-GW | RIL-TGW | RIL-GL |
---|---|---|---|---|---|---|
F2:3 | 粒长GL | 0.679** | ||||
粒宽GW | 0.725** | 0.279** | ||||
RIL | 千粒重TGW | -0.053 | ||||
粒长GL | 0.020 | 0.753** | ||||
粒宽GW | -0.163 | 0.776** | 0.394** |
表1 EFT/FM9 F2:3和RIL群体粒型性状相关性分析
Table 1 Correlationship analysis on grain shape traits in EFT/FM9 F2:3 and RIL population.
群体 Population | 性状Trait | F2:3-TGW | F2:3-GL | F2:3-GW | RIL-TGW | RIL-GL |
---|---|---|---|---|---|---|
F2:3 | 粒长GL | 0.679** | ||||
粒宽GW | 0.725** | 0.279** | ||||
RIL | 千粒重TGW | -0.053 | ||||
粒长GL | 0.020 | 0.753** | ||||
粒宽GW | -0.163 | 0.776** | 0.394** |
性状 | 数量性状基因座 | 标记区间 | F2:3 | RIL | 已克隆QTLa Cloned QTLa | |||||
---|---|---|---|---|---|---|---|---|---|---|
Trait | QTL | Marker interval | LOD | PVE/% | Add | LOD | PVE/% | Add | ||
千粒重 | qTGW1.1 | RM246-RM1061 | 3.97 | 3.25 | -0.92 | |||||
TGW | qTGW1.2 | RM315-RM12138 | 5.28 | 3.95 | -1.08 | 4.51 | 4.15 | -0.94 | ||
qTGW2 | RM145-RM71 | 59.83 | 38.33 | -3.70 | 31.8 | 34.24 | -3.00 | GW2[ | ||
qTGW3.1 | THI351-THI468 | 4.25 | 4.24 | -1.07 | ||||||
qTGW3.2 | THI1668-THI1701 | 16.05 | 6.74 | -1.49 | 8.68 | 7.60 | -1.41 | GS3[ | ||
qTGW3.3 | RM168-RM3867 | 11.15 | 5.89 | -1.39 | 5.82 | 5.92 | -1.24 | GSA1[ | ||
qTGW5.1 | FIF232-FIF345 | 8.53 | 3.67 | 0.95 | GS5[ | |||||
qTGW5.2 | FIF426-FIF738 | 9.86 | 4.64 | 1.11 | 5.95 | 4.97 | 1.14 | GSE5/GW5/qSW5[ | ||
qTGW6.1 | RM136-RM3 | 4.19 | 2.25 | -0.87 | ||||||
qTGW6.2 | RM30-RM340 | 6.12 | 2.80 | -1.04 | ||||||
qTGW7 | RM420-RM172 | 4.35 | 1.76 | -0.76 | ||||||
qTGW9.1 | RM296-RM7424 | 4.97 | 2.68 | -0.96 | DEP1/DN1/qNGR9[ | |||||
qTGW9.2 | RM7048-RM328 | 3.26 | 3.16 | -0.91 | ||||||
qTGW10.1 | RM228-RM590 | 6.70 | 2.64 | -0.92 | ||||||
qTGW10.2 | RM474-TEN400 | 4.12 | 3.43 | -0.96 | ||||||
qTGW12 | RM1103-RM3226 | 6.26 | 2.62 | -0.93 | ||||||
粒长 | qGL1.1 | RM1061-RM128 | 8.29 | 4.05 | -0.16 | |||||
GL | qGL1.2 | RM315-RM12138 | 7.47 | 3.83 | -0.15 | 9.10 | 8.25 | -0.21 | ||
qGL1.3 | RM12283-RM12333 | 2.99 | 2.42 | -0.12 | ||||||
qGL2 | RM145-RM71 | 12.09 | 6.35 | -0.21 | 12.82 | 18.24 | -0.32 | GW2[ | ||
qGL3.1 | THI156-THI214 | 4.86 | 4.01 | -0.15 | ||||||
qGL3.2 | THI301-THI351 | 5.06 | 2.36 | -0.12 | ||||||
qGL3.3 | THI1668-THI1701 | 45.92 | 32.65 | -0.44 | 24.88 | 25.95 | -0.38 | GS3[ | ||
qGL3.4 | RM168-RM3867 | 7.62 | 5.15 | -0.18 | 2.95 | 2.78 | -0.12 | GSA1[ | ||
qGL5 | RM538-RM480 | 5.14 | 2.45 | 0.13 | ||||||
qGL6 | RM589-RM587 | 4.26 | 2.27 | 0.12 | ||||||
qGL7 | RM351-RM234 | 5.78 | 3.32 | -0.14 | ||||||
qGL9 | RM296-RM7424 | 10.43 | 7.67 | -0.22 | DEP1/DN1/qNGR9[ | |||||
qGL11 | RM1355-RM209 | 7.18 | 3.70 | -0.16 | ||||||
粒宽 | qGW2 | RM145-RM71 | 48.09 | 30.95 | -0.20 | 17.85 | 24.96 | -0.17 | GW2[ | |
GW | qGW3 | RM3867-RM571 | 5.30 | 2.93 | -0.05 | GSA1[ | ||||
qGW5.1 | FIF426-FIF738 | 29.68 | 19.25 | 0.15 | 10.84 | 15.49 | 0.13 | GSE5/GW5/qSW5[ | ||
qGW5.2 | FIF738-FIF801 | 9.30 | 4.42 | 0.07 | ||||||
qGW11 | RM21-RM254 | 4.44 | 3.10 | -0.06 | ||||||
qGW12 | RM1103-RM3226 | 7.88 | 4.01 | -0.07 |
表2 F2:3和RIL群体中检测到的粒重粒形QTL
Table 2 QTLs for grain weight and shape detected in the F2:3 and RIL populations.
性状 | 数量性状基因座 | 标记区间 | F2:3 | RIL | 已克隆QTLa Cloned QTLa | |||||
---|---|---|---|---|---|---|---|---|---|---|
Trait | QTL | Marker interval | LOD | PVE/% | Add | LOD | PVE/% | Add | ||
千粒重 | qTGW1.1 | RM246-RM1061 | 3.97 | 3.25 | -0.92 | |||||
TGW | qTGW1.2 | RM315-RM12138 | 5.28 | 3.95 | -1.08 | 4.51 | 4.15 | -0.94 | ||
qTGW2 | RM145-RM71 | 59.83 | 38.33 | -3.70 | 31.8 | 34.24 | -3.00 | GW2[ | ||
qTGW3.1 | THI351-THI468 | 4.25 | 4.24 | -1.07 | ||||||
qTGW3.2 | THI1668-THI1701 | 16.05 | 6.74 | -1.49 | 8.68 | 7.60 | -1.41 | GS3[ | ||
qTGW3.3 | RM168-RM3867 | 11.15 | 5.89 | -1.39 | 5.82 | 5.92 | -1.24 | GSA1[ | ||
qTGW5.1 | FIF232-FIF345 | 8.53 | 3.67 | 0.95 | GS5[ | |||||
qTGW5.2 | FIF426-FIF738 | 9.86 | 4.64 | 1.11 | 5.95 | 4.97 | 1.14 | GSE5/GW5/qSW5[ | ||
qTGW6.1 | RM136-RM3 | 4.19 | 2.25 | -0.87 | ||||||
qTGW6.2 | RM30-RM340 | 6.12 | 2.80 | -1.04 | ||||||
qTGW7 | RM420-RM172 | 4.35 | 1.76 | -0.76 | ||||||
qTGW9.1 | RM296-RM7424 | 4.97 | 2.68 | -0.96 | DEP1/DN1/qNGR9[ | |||||
qTGW9.2 | RM7048-RM328 | 3.26 | 3.16 | -0.91 | ||||||
qTGW10.1 | RM228-RM590 | 6.70 | 2.64 | -0.92 | ||||||
qTGW10.2 | RM474-TEN400 | 4.12 | 3.43 | -0.96 | ||||||
qTGW12 | RM1103-RM3226 | 6.26 | 2.62 | -0.93 | ||||||
粒长 | qGL1.1 | RM1061-RM128 | 8.29 | 4.05 | -0.16 | |||||
GL | qGL1.2 | RM315-RM12138 | 7.47 | 3.83 | -0.15 | 9.10 | 8.25 | -0.21 | ||
qGL1.3 | RM12283-RM12333 | 2.99 | 2.42 | -0.12 | ||||||
qGL2 | RM145-RM71 | 12.09 | 6.35 | -0.21 | 12.82 | 18.24 | -0.32 | GW2[ | ||
qGL3.1 | THI156-THI214 | 4.86 | 4.01 | -0.15 | ||||||
qGL3.2 | THI301-THI351 | 5.06 | 2.36 | -0.12 | ||||||
qGL3.3 | THI1668-THI1701 | 45.92 | 32.65 | -0.44 | 24.88 | 25.95 | -0.38 | GS3[ | ||
qGL3.4 | RM168-RM3867 | 7.62 | 5.15 | -0.18 | 2.95 | 2.78 | -0.12 | GSA1[ | ||
qGL5 | RM538-RM480 | 5.14 | 2.45 | 0.13 | ||||||
qGL6 | RM589-RM587 | 4.26 | 2.27 | 0.12 | ||||||
qGL7 | RM351-RM234 | 5.78 | 3.32 | -0.14 | ||||||
qGL9 | RM296-RM7424 | 10.43 | 7.67 | -0.22 | DEP1/DN1/qNGR9[ | |||||
qGL11 | RM1355-RM209 | 7.18 | 3.70 | -0.16 | ||||||
粒宽 | qGW2 | RM145-RM71 | 48.09 | 30.95 | -0.20 | 17.85 | 24.96 | -0.17 | GW2[ | |
GW | qGW3 | RM3867-RM571 | 5.30 | 2.93 | -0.05 | GSA1[ | ||||
qGW5.1 | FIF426-FIF738 | 29.68 | 19.25 | 0.15 | 10.84 | 15.49 | 0.13 | GSE5/GW5/qSW5[ | ||
qGW5.2 | FIF738-FIF801 | 9.30 | 4.42 | 0.07 | ||||||
qGW11 | RM21-RM254 | 4.44 | 3.10 | -0.06 | ||||||
qGW12 | RM1103-RM3226 | 7.88 | 4.01 | -0.07 |
图6 高世代分离群体D1(A-C)和D2群体(D-F)中qTGW1.2/qGL1.2不同基因型间的株型、粒形和粒重的比较**表示在0.01水平上差异显著。
Fig. 6. Comparison of plant architecture, grain shape and grain weight among different genotypes of qTGW1.2/qGL1.2 in the D1 population (A-C) and the D2 population (D-F) of advanced generations. **Significant at 0.01 level.
群体 Population | 分离区别 Segregated region | 性状 Trait | qTGW1.2/qGL1.2EFT | qTGW1.2/qGL1.2FM9 | P值 P value | 加性效应 Additive effect |
---|---|---|---|---|---|---|
E1 | FIR3596-FIR3892 | 千粒重TGW/g | 33.00±1.09 | 34.77±1.03 | 0.0014 | -0.89 |
粒长GL/mm | 9.42±0.17 | 9.85±0.13 | <0.0001 | -0.21 | ||
粒宽GW/mm | 3.09±0.02 | 3.08±0.02 | 0.1703 | |||
E2 | FIR3596-FIR3892 | 千粒重TGW/g | 33.63±0.87 | 35.99±0.88 | <0.0001 | -1.19 |
粒长GL/mm | 9.67±0.17 | 10.11±0.11 | <0.0001 | -0.22 | ||
粒宽GW/mm | 3.10±0.03 | 3.09±0.04 | 0.7305 |
表3 高世代分离群体E1和E2中qTGW1.2/qGL1.2的效应分析
Table 3 Effect of qTGW1.2/qGL1.2 estimated in E1 and E2 populations of advanced generations.
群体 Population | 分离区别 Segregated region | 性状 Trait | qTGW1.2/qGL1.2EFT | qTGW1.2/qGL1.2FM9 | P值 P value | 加性效应 Additive effect |
---|---|---|---|---|---|---|
E1 | FIR3596-FIR3892 | 千粒重TGW/g | 33.00±1.09 | 34.77±1.03 | 0.0014 | -0.89 |
粒长GL/mm | 9.42±0.17 | 9.85±0.13 | <0.0001 | -0.21 | ||
粒宽GW/mm | 3.09±0.02 | 3.08±0.02 | 0.1703 | |||
E2 | FIR3596-FIR3892 | 千粒重TGW/g | 33.63±0.87 | 35.99±0.88 | <0.0001 | -1.19 |
粒长GL/mm | 9.67±0.17 | 10.11±0.11 | <0.0001 | -0.22 | ||
粒宽GW/mm | 3.10±0.03 | 3.09±0.04 | 0.7305 |
图7 携带qTGW1.2/qGL1.2的FM9纯合型(qTGW1.2/qGL1.2FM9)和EFT纯合型(qTGW1.2/qGL1.2EFT)材料的颖壳表皮细胞扫描电镜观察和相关基因的表达分析A- qTGW1.2/qGL1.2FM9和qTGW1.2/qGL1.2EFT的颖壳细胞(标尺:200 μm);B-外颖细胞长度和宽度的比较(*, **分别表示在 0.05 和 0.01 水平上显著相关,n=30);C- qTGW1.2/qGL1.2FM9和qTGW1.2/qGL1.2EFT幼穗中细胞周期与粒形相关基因的表达水平比较。CDKA1-A型细胞周期蛋白依赖性激酶1;CDKB2;1-B型周期蛋白依赖性激酶2;CYCA1;1- A型周期蛋白依赖性激酶1-1;CYCA2; 2-A型周期蛋白依赖性激酶2-1;CYCB1;1-B型周期蛋白依赖性激酶1-1;CYCB2; 2-B型周期蛋白依赖性激酶2-2;CYCD4;1-D型周期蛋白依赖性激酶4-1;CYCD4; 2-D型周期蛋白依赖性激酶4-2;KN-突触融合蛋白相关蛋白KNOLLE;CYCT1- T型细胞周期蛋白依赖性激酶1;CYCU4; 3-U型细胞周期蛋白依赖性激酶4-3;CDT2-U型细胞周期蛋白依赖性激酶4-1;MAD2-细胞扩增相关基因MAD2;E2F2- E2F转录因子2;MAPK-丝裂原活化蛋白激酶;MCM2-微型染色体维持蛋白2;MCM3-微型染色体维持蛋白3;MCM4-微型染色体维持蛋白4;GW2-粒宽基因;GS3-粒长基因;GW5 -粒宽基因;OsPIN1a-PIN蛋白1a;OsPIN1- PIN蛋白1;BG1-大粒基因;OsIAA11-Aux/IAA蛋白11;OsARF19-生长素应答因子19;TSG1-色氨酸转氨酶。
Fig. 7. Scanning electron observation and quantitative expression analysis of related genes in qTGW1.2/qGL1.2FM9 and qTGW1.2/qGL1.2EFT. A, Local outer surfaces of qTGW1.2/qGL1.2FM9 and qTGW1.2/qGL1.2EFT spikelet hulls. (Bars = 200 µm); B, Comparison of the cell length and width of outer glume cells (*, **Significant at 0.05 and 0.01 levels, respectively, n=30); C, Two parental homozygous genotypes qTGW1.2/qGL1.2FM9 and qTGW1.2/qGL1.2EFT are involved in the expression levels of cell cycle- and grain shape-related genes in young panicles. CDKA1, Cyclin-dependent kinase A-1; CDKB2;1, B-type cyclin-dependent kinase 2;1; CYCA1;1, A-type cyclin 1;1; CYCA2; 2, A-type cyclin 3;1; CYCB1;1, B-type cyclin 1; CYCB2; 2, B-type cyclin 2;2; CYCD4;1, D-type cyclin 4;1; CYCD4; 2, D-type cyclin 4;2; KN, Syntaxin-related protein KNOLLE; CYCT1, T-type cyclin 1; CYCU4; 3, U-type cyclin 4;3; CDT2, Cadmium tolerant 2; MAD2, Cell expansion-related gene MAD2; E2F2, E2F transcription factor 2; MAPK, Mitogen-activated protein kinase; MCM2, Mini-chromosome maintenance protein 2; MCM3, Mini-chromosome maintenance protein 3; MCM4, Mini-chromosome maintenance protein 4; GW2, Grain weight 2; GS3, Grain size 3; GW5, Grain width 5; OsPIN1a, Pin protein 1a; OsPIN1, Pin-formed 1b; BG1, Big grain 1; OsIAA11, Aux/IAA protein 11; OsARF19, Auxin response factor 19; TSG1, Tillering and small grain 1.
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