植物学报 ›› 2022, Vol. 57 ›› Issue (2): 157-170.DOI: 10.11983/CBB21216
叶涵斐1, 殷文晶1, 管易安1, 杨凯如1, 陈芊羽1, 俞淑颖1, 朱旭东2, 辛德东1, 章薇1,*(), 王跃星2,*(
), 饶玉春1(
)
收稿日期:
2021-12-12
接受日期:
2022-01-20
出版日期:
2022-03-01
发布日期:
2022-03-24
通讯作者:
章薇,王跃星,饶玉春
作者简介:
ryc@zjnu.cn基金资助:
Hanfei Ye1, Wenjing Yin1, Yian Guan1, Kairu Yang1, Qianyu Chen1, Shuying Yu1, Xudong Zhu2, Dedong Xin1, Wei Zhang1,*(), Yuexing Wang2,*(
), Yuchun Rao1(
)
Received:
2021-12-12
Accepted:
2022-01-20
Online:
2022-03-01
Published:
2022-03-24
Contact:
Wei Zhang,Yuexing Wang,Yuchun Rao
摘要:
维生素E (VE)是稻米营养品质的重要指标。水稻(Oryza sativa)是我国种植最广泛的粮食作物, 增加其籽粒的VE含量是实现国民营养强化的一条便捷有效的途径。该研究以籼稻华占(HZ)为父本, 粳稻热研2号(Nekken2)为母本, 构建120个重组自交系(RILs)群体。采用高效液相色谱法(HPLC)对RILs群体的VE各组分含量进行测定, 并基于构建的高密度分子遗传图谱进行QTL定位, 谱系分析后挖掘到122个VE总量和分量相关QTLs, 分布在12条染色体上。其中qT3α/to2-1的LOD值高达10.32, qT3α2-1的LOD值高达9.91, 另有多个控制各异构体含量的主效QTLs, 且区间内包含OsGGR1、OsGGR2、OsTC及OsγTMT等VE生物合成基因。通过qRT-PCR检测亲本中VE合成基因的表达量, 发现在华占中候选基因的表达量均极显著高于热研2号, 推测这些基因的高表达是华占生育酚及生育三烯酚含量高于热研2号的原因。研究挖掘到的QTL数目较多, LOD值也较大, 为进一步筛选和培育高VE含量的水稻新品种奠定了分子基础, 同时为揭示水稻VE生物合成的分子调控机制提供了重要基因资源。
叶涵斐, 殷文晶, 管易安, 杨凯如, 陈芊羽, 俞淑颖, 朱旭东, 辛德东, 章薇, 王跃星, 饶玉春. 水稻籽粒维生素E QTL挖掘及候选基因分析. 植物学报, 2022, 57(2): 157-170.
Hanfei Ye, Wenjing Yin, Yian Guan, Kairu Yang, Qianyu Chen, Shuying Yu, Xudong Zhu, Dedong Xin, Wei Zhang, Yuexing Wang, Yuchun Rao. QTL Mapping and Candidate Gene Analysis of Vitamin E in Rice Grain. Chinese Bulletin of Botany, 2022, 57(2): 157-170.
Primer name | Sequence (5′-3′) | Tm (°C) | Length (bp) |
---|---|---|---|
LOC_Os01g16020-F-qrt | CCATGGCCTACTACGACGAC | 60.05 | 150 |
LOC_Os01g16020-R-qrt | CCGACTGGAGCCTCTTGATC | 59.95 | |
LOC_Os02g07160-F-qrt | AGGAGCAGATCAACGAGTGC | 60.11 | 172 |
LOC_Os02g07160-R-qrt | TTCTGGTACTCCTGCCCACT | 60.28 | |
LOC_Os02g17650-F-qrt | TCGACGCAACAAGCAACATG | 59.95 | 161 |
LOC_Os02g17650-R-qrt | ACCAGGGGGCTTGAGAAATG | 59.75 | |
LOC_Os02g47310-F-qrt | CGGCAGAGCAAGGGTTATCA | 59.96 | 223 |
LOC_Os02g47310-R-qrt | GACTCTTCGGATGGCTCGAG | 60.39 | |
LOC_Os02g51080-F-qrt | GATGATTTCGCCGTCCAACG | 60.05 | 200 |
LOC_Os02g51080-R-qrt | TGTAGTGCACCACGTACGAC | 60.03 | |
LOC_Os06g43880-F-qrt | CCGTGCAAGTCCAGGAAGAT | 59.96 | 196 |
LOC_Os06g43880-R-qrt | TCACCAGCAATGGCATAGCA | 60.00 | |
LOC_Os06g44840-F-qrt | CCTTGGATGGGCTGTTGGAT | 59.99 | 223 |
LOC_Os06g44840-R-qrt | TGGCCTGGTAAAGACTGCTG | 60.08 | |
LOC_Os07g08200-F-qrt | ACCGAGGACATGAGGGATGA | 59.98 | 179 |
LOC_Os07g08200-R-qrt | TTCTGCCTGGCCTTCTCAAG | 60.07 | |
LOC_Os12g42090-F-qrt | CTCAAGGGCGTCACCATCAT | 60.12 | 224 |
LOC_Os12g42090-R-qrt | AGCATCCACATGTCAGCGAA | 59.98 |
表1 qRT-PCR所用引物序列
Table 1 Primer sequences for qRT-PCR
Primer name | Sequence (5′-3′) | Tm (°C) | Length (bp) |
---|---|---|---|
LOC_Os01g16020-F-qrt | CCATGGCCTACTACGACGAC | 60.05 | 150 |
LOC_Os01g16020-R-qrt | CCGACTGGAGCCTCTTGATC | 59.95 | |
LOC_Os02g07160-F-qrt | AGGAGCAGATCAACGAGTGC | 60.11 | 172 |
LOC_Os02g07160-R-qrt | TTCTGGTACTCCTGCCCACT | 60.28 | |
LOC_Os02g17650-F-qrt | TCGACGCAACAAGCAACATG | 59.95 | 161 |
LOC_Os02g17650-R-qrt | ACCAGGGGGCTTGAGAAATG | 59.75 | |
LOC_Os02g47310-F-qrt | CGGCAGAGCAAGGGTTATCA | 59.96 | 223 |
LOC_Os02g47310-R-qrt | GACTCTTCGGATGGCTCGAG | 60.39 | |
LOC_Os02g51080-F-qrt | GATGATTTCGCCGTCCAACG | 60.05 | 200 |
LOC_Os02g51080-R-qrt | TGTAGTGCACCACGTACGAC | 60.03 | |
LOC_Os06g43880-F-qrt | CCGTGCAAGTCCAGGAAGAT | 59.96 | 196 |
LOC_Os06g43880-R-qrt | TCACCAGCAATGGCATAGCA | 60.00 | |
LOC_Os06g44840-F-qrt | CCTTGGATGGGCTGTTGGAT | 59.99 | 223 |
LOC_Os06g44840-R-qrt | TGGCCTGGTAAAGACTGCTG | 60.08 | |
LOC_Os07g08200-F-qrt | ACCGAGGACATGAGGGATGA | 59.98 | 179 |
LOC_Os07g08200-R-qrt | TTCTGCCTGGCCTTCTCAAG | 60.07 | |
LOC_Os12g42090-F-qrt | CTCAAGGGCGTCACCATCAT | 60.12 | 224 |
LOC_Os12g42090-R-qrt | AGCATCCACATGTCAGCGAA | 59.98 |
图1 维生素E各异构体的高效液相色谱分析 α-t3: α-生育三烯酚; γ-t3: γ-生育三烯酚; δ-t3: δ生育三烯酚; α-toco: α-生育酚; γ-toco: γ-生育酚; δ-toco: δ-生育酚; IS: 内标
Figure 1 High performance liquid chromatography analysis of vitamin E isomers α-t3: α-tocotrienol; γ-t3: γ-tocotrienol; δ-t3: δ-tocotrienol; α-toco: α-tocophorel; γ-toco: γ-tocophorel; δ-toco: δ-tocophorel; IS: Internal standard
Trait | RILs | Parents | |||
---|---|---|---|---|---|
Means±SD | Range | Skewness | HZ | Nekken2 | |
Total-VE (μg∙g-1) | 33.9489±8.24636 | 11.64-62.93 | 0.7220 | 39.0169 | 25.1275 |
Total-toco (μg∙g-1) | 13.9571±6.54920 | 4.19-43.16 | 1.7550 | 15.6586 | 7.3524 |
Total-t3 (μg∙g-1) | 19.9918±4.44004 | 7.45-30.41 | 0.0940 | 23.3583 | 17.7751 |
α-toco (μg∙g-1) | 11.0692±6.46111 | 2.34-41.33 | 1.9120 | 11.8855 | 4.7839 |
γ-toco (μg∙g-1) | 2.7072±1.11795 | 0.74-6.73 | 0.3200 | 3.6121 | 2.3390 |
δ-toco (μg∙g-1) | 0.1807±0.12540 | 0.03-0.92 | 2.1220 | 0.1610 | 0.2295 |
α-t3 (μg∙g-1) | 2.3057±1.33266 | 0.60-6.10 | 1.1710 | 1.4780 | 5.1269 |
γ-t3 (μg∙g-1) | 16.0730±4.3067 | 5.76-26.14 | 0.1970 | 20.4400 | 11.2289 |
δ-t3 (μg∙g-1) | 1.6131±0.63116 | 0.56-4.46 | 1.4040 | 1.4404 | 1.4193 |
α-toco/total-toco | 0.7618±0.12004 | 0.36-0.96 | 0.7610 | 0.7596 | 0.6344 |
α-t3/total-t3 | 0.1203±0.07395 | 0.04-0.34 | 1.0980 | 0.0634 | 0.2881 |
表2 水稻亲本及重组自交系群体(RILs)维生素E相关性状表型变异
Table 2 Phenotypic variation of vitamin E related traits in parents and recombinant inbred lines (RILs) of rice
Trait | RILs | Parents | |||
---|---|---|---|---|---|
Means±SD | Range | Skewness | HZ | Nekken2 | |
Total-VE (μg∙g-1) | 33.9489±8.24636 | 11.64-62.93 | 0.7220 | 39.0169 | 25.1275 |
Total-toco (μg∙g-1) | 13.9571±6.54920 | 4.19-43.16 | 1.7550 | 15.6586 | 7.3524 |
Total-t3 (μg∙g-1) | 19.9918±4.44004 | 7.45-30.41 | 0.0940 | 23.3583 | 17.7751 |
α-toco (μg∙g-1) | 11.0692±6.46111 | 2.34-41.33 | 1.9120 | 11.8855 | 4.7839 |
γ-toco (μg∙g-1) | 2.7072±1.11795 | 0.74-6.73 | 0.3200 | 3.6121 | 2.3390 |
δ-toco (μg∙g-1) | 0.1807±0.12540 | 0.03-0.92 | 2.1220 | 0.1610 | 0.2295 |
α-t3 (μg∙g-1) | 2.3057±1.33266 | 0.60-6.10 | 1.1710 | 1.4780 | 5.1269 |
γ-t3 (μg∙g-1) | 16.0730±4.3067 | 5.76-26.14 | 0.1970 | 20.4400 | 11.2289 |
δ-t3 (μg∙g-1) | 1.6131±0.63116 | 0.56-4.46 | 1.4040 | 1.4404 | 1.4193 |
α-toco/total-toco | 0.7618±0.12004 | 0.36-0.96 | 0.7610 | 0.7596 | 0.6344 |
α-t3/total-t3 | 0.1203±0.07395 | 0.04-0.34 | 1.0980 | 0.0634 | 0.2881 |
图2 水稻重组自交系中VE异构体含量的分布 α-t3、δ-t3、α-toco和δ-toco同图1。β-t3: β-生育三烯酚; β-toco: β-生育酚
Figure 2 Distribution of vitamin E isomers in recombinant inbred lines of rice α-t3, δ-t3, α-toco, and δ-toco are the same as Figure 1. β-t3: β-tocotrienol; β-toco: β-tocophorel
图3 水稻维生素E各异构体的相关系数 α-t3、γ-t3、δ-t3、α-toco、γ-toco和δ-toco同图1。
Figure 3 Correlation coefficient of vitamin E isomers in rice α-t3, γ-t3, δ-t3, α-toco, γ-toco, and δ-toco are the same as Figure 1.
Trait | QTL | Chromosome | Physical distance (bp) | Position of support (cM) | LOD | |||
---|---|---|---|---|---|---|---|---|
Total | qVE2-1 | 02 | 3467582-15790369 | 14.86-67.69 | 2.82 | |||
qVE2-2 | 02 | 30526267-30670565 | 130.86-131.48 | 3.11 | ||||
qVE5-1 | 05 | 27292002-27936693 | 116.99-119.76 | 3.05 | ||||
qVE7-1 | 07 | 25017683-25489862 | 107.24-109.27 | 3.50 | ||||
qVE8-1 | 08 | 10042712-11311929 | 43.05-48.49 | 3.69 | ||||
qVE8-2 | 08 | 14098425-15488141 | 60.44-66.39 | 4.00 | ||||
qVE8-3 | 08 | 19014129-21020793 | 81.51-90.11 | 4.85 | ||||
qVE10-1 | 10 | 10714090-13003806 | 45.93-55.74 | 3.70 | ||||
Total-t3 | qT3to1-1 | 01 | 7035569-8460131 | 30.16-36.27 | 3.51 | |||
qT3to3-1 | 03 | 11975352-12205246 | 51.34-52.32 | 2.93 | ||||
qT3to5-1 | 05 | 27199002-27292002 | 116.59-116.99 | 2.88 | ||||
qT3to10-1 | 10 | 13003806-15914330 | 55.74-68.22 | 2.96 | ||||
Total-toco | qTto2-1 | 02 | 7154574-16772608 | 30.67-71.90 | 3.63 | |||
qTto8-1 | 08 | 10042712-15565426 | 43.05-66.72 | 2.68 | ||||
qTto8-2 | 08 | 19301150-20349908 | 82.74-87.23 | 3.65 | ||||
qTto10-1 | 10 | 57733-5856857 | 0.25-25.11 | 3.46 | ||||
qTto10-2 | 10 | 10573976-13003806 | 45.33-55.74 | 7.39 | ||||
qTto11-1 | 11 | 9677541-9808325 | 41.48-42.05 | 2.65 | ||||
α-t3 | qT3α1-1 | 01 | 8235815-8460131 | 35.30-36.27 | 3.57 | |||
qT3α2-1 | 02 | 20415024-32383695 | 87.51-138.82 | 9.91 | ||||
qT3α2-2 | 02 | 32754542-35824426 | 140.41-153.57 | 3.64 | ||||
qT3α5 | 05 | 4080644-6619616 | 17.49-28.38 | 3.58 | ||||
qT3α7-1 | 07 | 42757-572873 | 0.18-2.46 | 3.27 | ||||
qT3α8-1 | 08 | 4857563-6004458 | 20.82-25.74 | 3.43 | ||||
qT3α8-2 | 08 | 8917564-15791348 | 38.23-67.69 | 4.66 | ||||
qT3α8-3 | 08 | 24746469-25177340 | 106.08-107.93 | 2.50 | ||||
qT3α8-4 | 08 | 25195867-25340928 | 108.01-108.63 | 2.55 | ||||
qT3α9-1 | 09 | 13620974-14832840 | 58.39-63.58 | 3.10 | ||||
qT3α9-2 | 09 | 16630835-22938953 | 71.29-98.33 | 5.28 | ||||
α-t3/total-t3 | qT3α/to1-1 | 01 | 8235815-9438021 | 35.30-40.46 | 3.66 | |||
qT3α/to2-1 | 02 | 20415024-35824426 | 87.51-153.57 | 10.32 | ||||
qT3α/to3-1 | 03 | 5183108-6234241 | 22.22-26.72 | 3.25 | ||||
qT3α/to3-2 | 03 | 6743718-6982005 | 28.91-29.93 | 3.31 | ||||
qT3α/to5-1 | 05 | 4080644-6619616 | 17.49-28.38 | 3.31 | ||||
qT3α/to6-1 | 06 | 20854685-21217545 | 89.40-90.95 | 2.81 | ||||
qT3α/to6-2 | 06 | 21507916-21913927 | 92.20-93.94 | 2.66 | ||||
qT3α/to6-3 | 06 | 22256655-24198880 | 95.41-103.73 | 3.57 | ||||
qT3α/to7-1 | 07 | 42757-875466 | 0.18-3.75 | 2.88 | ||||
qT3α/to8-1 | 08 | 5469252-5970122 | 23.45-25.59 | 3.53 | ||||
qT3α/to8-2 | 08 | 13109769-13402076 | 56.20-57.45 | 3.79 | ||||
qT3α/to8-3 | 08 | 24432953-25340928 | 104.74-108.63 | 2.96 | ||||
qT3α/to9-1 | 09 | 13572882-14761639 | 58.18-63.28 | 3.42 | ||||
qT3α/to9-2 | 09 | 16699350-17528726 | 71.59-75.14 | 2.78 | ||||
qT3α/to9-3 | 09 | 17822997-22938953 | 76.40-98.33 | 6.47 | ||||
qT3α/to10-1 | 10 | 12856184-13003806 | 55.11-55.74 | 3.65 | ||||
qT3α/to10-2 | 10 | 19066686-19211967 | 81.73-82.36 | 2.57 | ||||
α-toco | qTα2-1 | 02 | 10650561-10928729 | 45.66-46.85 | 3.41 | |||
qTα2-2 | 02 | 11181757-11319097 | 47.932-48.52 | 2.54 | ||||
qTα8-1 | 08 | 10119315-15791348 | 43.38-67.69 | 2.77 | ||||
qTα8-2 | 08 | 19301150-20108258 | 82.74-86.20 | 3.10 | ||||
qTα10-1 | 10 | 1624313-5776626 | 6.96-24.76 | 3.17 | ||||
qTα10-2 | 10 | 10635436-11008219 | 45.59-47.19 | 3.90 | ||||
qTα11-1 | 11 | 9677541-9808325 | 41.48-42.05 | 2.96 | ||||
qTα11-2 | 11 | 21463452-22167073 | 92.01-95.02 | 3.06 | ||||
α-toco/total-toco | qTα/to1-1 | 01 | 1663060-2070966 | 7.13-8.88 | 2.61 | |||
qTα/to2-1 | 02 | 10650561-10928729 | 45.66-46.85 | 3.83 | ||||
qTα/to2-2 | 02 | 27158525-28132392 | 116.42-120.60 | 2.83 | ||||
qTα/to2-3 | 02 | 28499903-30078981 | 122.17-128.94 | 3.76 | ||||
qTα/to2-4 | 02 | 35381094-35525300 | 151.67-152.29 | 3.24 | ||||
qTα/to3-1 | 03 | 786808-1156051 | 3.37-4.96 | 4.36 | ||||
qTα/to3-2 | 03 | 1531836-1753422 | 6.57-7.52 | 2.81 | ||||
qTα/to4-1 | 04 | 4147493-4739169 | 17.78-20.32 | 4.23 | ||||
qTα/to4-2 | 04 | 6047670-9068118 | 25.92-38.87 | 4.23 | ||||
qTα/to4-3 | 04 | 13341761-14630459 | 57.19-62.72 | 4.18 | ||||
qTα/to5-1 | 05 | 6526040-6673100 | 27.98-28.61 | 2.80 | ||||
qTα/to5-2 | 05 | 7886106-8030564 | 33.81-34.42 | 4.14 | ||||
qTα/to5-3 | 05 | 19534829-19659074 | 83.74-84.27 | 2.67 | ||||
qTα/to6-1 | 06 | 3108218-3378992 | 13.32-14.48 | 3.02 | ||||
qTα/to6-2 | 06 | 16365713-16951387 | 70.16-72.67 | 2.53 | ||||
qTα/to6-3 | 06 | 24956043-25093708 | 106.98-107.57 | 2.79 | ||||
qTα/to6-4 | 06 | 26359832-27156405 | 113.00-116.41 | 3.18 | ||||
qTα/to7-1 | 07 | 13014544-13768860 | 55.79-59.02 | 3.42 | ||||
qTα/to7-2 | 07 | 17441086-17530313 | 74.77-75.15 | 2.70 | ||||
qTα/to8-1 | 08 | 712661-860780 | 3.05-3.69 | 4.14 | ||||
qTα/to8-2 | 08 | 1152527-1753671 | 4.94-7.52 | 4.58 | ||||
qTα/to8-3 | 08 | 5341420-5646427 | 22.90-24.20 | 3.94 | ||||
qTα/to9-1 | 09 | 11654883-11744700 | 49.96-50.35 | 2.52 | ||||
qTα/to11-1 | 11 | 8467466-8766502 | 36.30-37.58 | 3.04 | ||||
qTα/to11-2 | 11 | 21077880-22646003 | 90.36-97.08 | 4.92 | ||||
qTα/to12-1 | 12 | 260226-401477 | 1.12-1.72 | 2.50 | ||||
qTα/to12-2 | 12 | 6824561-7508660 | 29.25-32.19 | 3.34 | ||||
qTα/to12-3 | 12 | 17175063-18091161 | 73.62-77.55 | 3.32 | ||||
γ-t3 | qT3γ1-1 | 01 | 7128319-9521270 | 30.56-40.82 | 3.31 | |||
qT3γ2-1 | 02 | 29846956-31331712 | 127.95-134.31 | 3.26 | ||||
qT3γ2-2 | 02 | 35381094-35675126 | 151.67-152.93 | 3.15 | ||||
qT3γ6-1 | 06 | 27333693-28446541 | 117.17-121.94 | 3.54 | ||||
qT3γ8-1 | 08 | 712661-860780 | 3.05-3.69 | 3.78 | ||||
γ-toco | qTγ1-1 | 01 | 1663060-1775390 | 7.13-7.61 | 2.56 | |||
qTγ2-1 | 02 | 24818487-35305635 | 106.39-151.35 | 6.20 | ||||
qTγ3-1 | 03 | 5859763-6008076 | 25.12-25.75 | 2.50 | ||||
qTγ4-1 | 04 | 5922401-6230638 | 25.39-26.71 | 2.92 | ||||
qTγ4-2 | 04 | 6451666-6599549 | 27.66-28.29 | 2.54 | ||||
qTγ5-1 | 05 | 6983157-10879499 | 29.93-46.64 | 2.91 | ||||
qTγ5-2 | 05 | 27199002-27292002 | 116.59-116.99 | 2.73 | ||||
qTγ6-1 | 06 | 6452921-6720585 | 27.66-28.81 | 3.22 | ||||
qTγ7-1 | 07 | 572873-936652 | 2.46-4.02 | 2.64 | ||||
qTγ8-1 | 08 | 712661-860780 | 3.05-3.69 | 2.72 | ||||
qTγ9-1 | 09 | 21030827-21162733 | 90.15-90.72 | 2.51 | ||||
δ-t3 | qT3δ1-1 | 01 | 19435110-19915117 | 83.31-85.37 | 3.36 | |||
qT3δ1-2 | 01 | 25117984-25257953 | 107.67-108.27 | 5.75 | ||||
qT3δ2-1 | 02 | 7215671-8736288 | 30.93-37.45 | 2.66 | ||||
qT3δ2-2 | 02 | 30228480-32985884 | 129.58-141.40 | 2.66 | ||||
qT3δ3-1 | 03 | 29597075-29806859 | 126.87-127.77 | 2.86 | ||||
qT3δ4-1 | 04 | 17170511-23608260 | 73.61-101.20 | 3.31 | ||||
qT3δ4-2 | 04 | 26410305-31423289 | 113.21-134.70 | 4.71 | ||||
qT3δ8-1 | 08 | 20195421-20349908 | 86.57-87.23 | 2.97 | ||||
qT3δ10-1 | 10 | 57733-13172007 | 0.25-56.46 | 3.88 | ||||
qT3δ10-2 | 10 | 19963553-20188240 | 85.58-86.54 | 4.49 | ||||
qT3δ12-1 | 12 | 4509538-4663215 | 19.33-19.99 | 3.16 | ||||
qT3δ12-2 | 12 | 19235536-26139939 | 82.46-112.05 | 4.90 | ||||
δ-toco | qTδ1-1 | 01 | 38537755-39791276 | 165.20-170.57 | 4.28 | |||
qTδ2-1 | 02 | 7215671-7350401 | 30.93-31.51 | 2.89 | ||||
qTδ2-2 | 02 | 31043939-32985884 | 133.08-141.40 | 2.57 | ||||
qTδ3-1 | 03 | 5785084-6008076 | 24.80-25.75 | 3.49 | ||||
qTδ5-1 | 05 | 853216-1082203 | 3.66-4.64 | 3.23 | ||||
qTδ5-2 | 05 | 6983157-7138974 | 29.93-30.60 | 6.17 | ||||
qTδ5-3 | 05 | 13569180-13776389 | 58.17-59.06 | 2.70 | ||||
qTδ6-1 | 06 | 6452921-6720585 | 27.67-28.81 | 3.80 | ||||
qTδ7-1 | 07 | 7351588-15250540 | 31.51-65.37 | 2.73 | ||||
qTδ8-1 | 08 | 1152527-5163244 | 4.94-22.13 | 3.51 | ||||
qTδ9-1 | 09 | 10102157-12363787 | 43.31-53.00 | 3.73 | ||||
qTδ12-1 | 12 | 9212254-15247966 | 39.49-65.36 | 7.16 |
表3 水稻重组自交系群体籽粒中维生素E异构体的QTL分析
Table 3 QTL analysis of vitamin E isomers in grains of rice recombinant inbred lines
Trait | QTL | Chromosome | Physical distance (bp) | Position of support (cM) | LOD | |||
---|---|---|---|---|---|---|---|---|
Total | qVE2-1 | 02 | 3467582-15790369 | 14.86-67.69 | 2.82 | |||
qVE2-2 | 02 | 30526267-30670565 | 130.86-131.48 | 3.11 | ||||
qVE5-1 | 05 | 27292002-27936693 | 116.99-119.76 | 3.05 | ||||
qVE7-1 | 07 | 25017683-25489862 | 107.24-109.27 | 3.50 | ||||
qVE8-1 | 08 | 10042712-11311929 | 43.05-48.49 | 3.69 | ||||
qVE8-2 | 08 | 14098425-15488141 | 60.44-66.39 | 4.00 | ||||
qVE8-3 | 08 | 19014129-21020793 | 81.51-90.11 | 4.85 | ||||
qVE10-1 | 10 | 10714090-13003806 | 45.93-55.74 | 3.70 | ||||
Total-t3 | qT3to1-1 | 01 | 7035569-8460131 | 30.16-36.27 | 3.51 | |||
qT3to3-1 | 03 | 11975352-12205246 | 51.34-52.32 | 2.93 | ||||
qT3to5-1 | 05 | 27199002-27292002 | 116.59-116.99 | 2.88 | ||||
qT3to10-1 | 10 | 13003806-15914330 | 55.74-68.22 | 2.96 | ||||
Total-toco | qTto2-1 | 02 | 7154574-16772608 | 30.67-71.90 | 3.63 | |||
qTto8-1 | 08 | 10042712-15565426 | 43.05-66.72 | 2.68 | ||||
qTto8-2 | 08 | 19301150-20349908 | 82.74-87.23 | 3.65 | ||||
qTto10-1 | 10 | 57733-5856857 | 0.25-25.11 | 3.46 | ||||
qTto10-2 | 10 | 10573976-13003806 | 45.33-55.74 | 7.39 | ||||
qTto11-1 | 11 | 9677541-9808325 | 41.48-42.05 | 2.65 | ||||
α-t3 | qT3α1-1 | 01 | 8235815-8460131 | 35.30-36.27 | 3.57 | |||
qT3α2-1 | 02 | 20415024-32383695 | 87.51-138.82 | 9.91 | ||||
qT3α2-2 | 02 | 32754542-35824426 | 140.41-153.57 | 3.64 | ||||
qT3α5 | 05 | 4080644-6619616 | 17.49-28.38 | 3.58 | ||||
qT3α7-1 | 07 | 42757-572873 | 0.18-2.46 | 3.27 | ||||
qT3α8-1 | 08 | 4857563-6004458 | 20.82-25.74 | 3.43 | ||||
qT3α8-2 | 08 | 8917564-15791348 | 38.23-67.69 | 4.66 | ||||
qT3α8-3 | 08 | 24746469-25177340 | 106.08-107.93 | 2.50 | ||||
qT3α8-4 | 08 | 25195867-25340928 | 108.01-108.63 | 2.55 | ||||
qT3α9-1 | 09 | 13620974-14832840 | 58.39-63.58 | 3.10 | ||||
qT3α9-2 | 09 | 16630835-22938953 | 71.29-98.33 | 5.28 | ||||
α-t3/total-t3 | qT3α/to1-1 | 01 | 8235815-9438021 | 35.30-40.46 | 3.66 | |||
qT3α/to2-1 | 02 | 20415024-35824426 | 87.51-153.57 | 10.32 | ||||
qT3α/to3-1 | 03 | 5183108-6234241 | 22.22-26.72 | 3.25 | ||||
qT3α/to3-2 | 03 | 6743718-6982005 | 28.91-29.93 | 3.31 | ||||
qT3α/to5-1 | 05 | 4080644-6619616 | 17.49-28.38 | 3.31 | ||||
qT3α/to6-1 | 06 | 20854685-21217545 | 89.40-90.95 | 2.81 | ||||
qT3α/to6-2 | 06 | 21507916-21913927 | 92.20-93.94 | 2.66 | ||||
qT3α/to6-3 | 06 | 22256655-24198880 | 95.41-103.73 | 3.57 | ||||
qT3α/to7-1 | 07 | 42757-875466 | 0.18-3.75 | 2.88 | ||||
qT3α/to8-1 | 08 | 5469252-5970122 | 23.45-25.59 | 3.53 | ||||
qT3α/to8-2 | 08 | 13109769-13402076 | 56.20-57.45 | 3.79 | ||||
qT3α/to8-3 | 08 | 24432953-25340928 | 104.74-108.63 | 2.96 | ||||
qT3α/to9-1 | 09 | 13572882-14761639 | 58.18-63.28 | 3.42 | ||||
qT3α/to9-2 | 09 | 16699350-17528726 | 71.59-75.14 | 2.78 | ||||
qT3α/to9-3 | 09 | 17822997-22938953 | 76.40-98.33 | 6.47 | ||||
qT3α/to10-1 | 10 | 12856184-13003806 | 55.11-55.74 | 3.65 | ||||
qT3α/to10-2 | 10 | 19066686-19211967 | 81.73-82.36 | 2.57 | ||||
α-toco | qTα2-1 | 02 | 10650561-10928729 | 45.66-46.85 | 3.41 | |||
qTα2-2 | 02 | 11181757-11319097 | 47.932-48.52 | 2.54 | ||||
qTα8-1 | 08 | 10119315-15791348 | 43.38-67.69 | 2.77 | ||||
qTα8-2 | 08 | 19301150-20108258 | 82.74-86.20 | 3.10 | ||||
qTα10-1 | 10 | 1624313-5776626 | 6.96-24.76 | 3.17 | ||||
qTα10-2 | 10 | 10635436-11008219 | 45.59-47.19 | 3.90 | ||||
qTα11-1 | 11 | 9677541-9808325 | 41.48-42.05 | 2.96 | ||||
qTα11-2 | 11 | 21463452-22167073 | 92.01-95.02 | 3.06 | ||||
α-toco/total-toco | qTα/to1-1 | 01 | 1663060-2070966 | 7.13-8.88 | 2.61 | |||
qTα/to2-1 | 02 | 10650561-10928729 | 45.66-46.85 | 3.83 | ||||
qTα/to2-2 | 02 | 27158525-28132392 | 116.42-120.60 | 2.83 | ||||
qTα/to2-3 | 02 | 28499903-30078981 | 122.17-128.94 | 3.76 | ||||
qTα/to2-4 | 02 | 35381094-35525300 | 151.67-152.29 | 3.24 | ||||
qTα/to3-1 | 03 | 786808-1156051 | 3.37-4.96 | 4.36 | ||||
qTα/to3-2 | 03 | 1531836-1753422 | 6.57-7.52 | 2.81 | ||||
qTα/to4-1 | 04 | 4147493-4739169 | 17.78-20.32 | 4.23 | ||||
qTα/to4-2 | 04 | 6047670-9068118 | 25.92-38.87 | 4.23 | ||||
qTα/to4-3 | 04 | 13341761-14630459 | 57.19-62.72 | 4.18 | ||||
qTα/to5-1 | 05 | 6526040-6673100 | 27.98-28.61 | 2.80 | ||||
qTα/to5-2 | 05 | 7886106-8030564 | 33.81-34.42 | 4.14 | ||||
qTα/to5-3 | 05 | 19534829-19659074 | 83.74-84.27 | 2.67 | ||||
qTα/to6-1 | 06 | 3108218-3378992 | 13.32-14.48 | 3.02 | ||||
qTα/to6-2 | 06 | 16365713-16951387 | 70.16-72.67 | 2.53 | ||||
qTα/to6-3 | 06 | 24956043-25093708 | 106.98-107.57 | 2.79 | ||||
qTα/to6-4 | 06 | 26359832-27156405 | 113.00-116.41 | 3.18 | ||||
qTα/to7-1 | 07 | 13014544-13768860 | 55.79-59.02 | 3.42 | ||||
qTα/to7-2 | 07 | 17441086-17530313 | 74.77-75.15 | 2.70 | ||||
qTα/to8-1 | 08 | 712661-860780 | 3.05-3.69 | 4.14 | ||||
qTα/to8-2 | 08 | 1152527-1753671 | 4.94-7.52 | 4.58 | ||||
qTα/to8-3 | 08 | 5341420-5646427 | 22.90-24.20 | 3.94 | ||||
qTα/to9-1 | 09 | 11654883-11744700 | 49.96-50.35 | 2.52 | ||||
qTα/to11-1 | 11 | 8467466-8766502 | 36.30-37.58 | 3.04 | ||||
qTα/to11-2 | 11 | 21077880-22646003 | 90.36-97.08 | 4.92 | ||||
qTα/to12-1 | 12 | 260226-401477 | 1.12-1.72 | 2.50 | ||||
qTα/to12-2 | 12 | 6824561-7508660 | 29.25-32.19 | 3.34 | ||||
qTα/to12-3 | 12 | 17175063-18091161 | 73.62-77.55 | 3.32 | ||||
γ-t3 | qT3γ1-1 | 01 | 7128319-9521270 | 30.56-40.82 | 3.31 | |||
qT3γ2-1 | 02 | 29846956-31331712 | 127.95-134.31 | 3.26 | ||||
qT3γ2-2 | 02 | 35381094-35675126 | 151.67-152.93 | 3.15 | ||||
qT3γ6-1 | 06 | 27333693-28446541 | 117.17-121.94 | 3.54 | ||||
qT3γ8-1 | 08 | 712661-860780 | 3.05-3.69 | 3.78 | ||||
γ-toco | qTγ1-1 | 01 | 1663060-1775390 | 7.13-7.61 | 2.56 | |||
qTγ2-1 | 02 | 24818487-35305635 | 106.39-151.35 | 6.20 | ||||
qTγ3-1 | 03 | 5859763-6008076 | 25.12-25.75 | 2.50 | ||||
qTγ4-1 | 04 | 5922401-6230638 | 25.39-26.71 | 2.92 | ||||
qTγ4-2 | 04 | 6451666-6599549 | 27.66-28.29 | 2.54 | ||||
qTγ5-1 | 05 | 6983157-10879499 | 29.93-46.64 | 2.91 | ||||
qTγ5-2 | 05 | 27199002-27292002 | 116.59-116.99 | 2.73 | ||||
qTγ6-1 | 06 | 6452921-6720585 | 27.66-28.81 | 3.22 | ||||
qTγ7-1 | 07 | 572873-936652 | 2.46-4.02 | 2.64 | ||||
qTγ8-1 | 08 | 712661-860780 | 3.05-3.69 | 2.72 | ||||
qTγ9-1 | 09 | 21030827-21162733 | 90.15-90.72 | 2.51 | ||||
δ-t3 | qT3δ1-1 | 01 | 19435110-19915117 | 83.31-85.37 | 3.36 | |||
qT3δ1-2 | 01 | 25117984-25257953 | 107.67-108.27 | 5.75 | ||||
qT3δ2-1 | 02 | 7215671-8736288 | 30.93-37.45 | 2.66 | ||||
qT3δ2-2 | 02 | 30228480-32985884 | 129.58-141.40 | 2.66 | ||||
qT3δ3-1 | 03 | 29597075-29806859 | 126.87-127.77 | 2.86 | ||||
qT3δ4-1 | 04 | 17170511-23608260 | 73.61-101.20 | 3.31 | ||||
qT3δ4-2 | 04 | 26410305-31423289 | 113.21-134.70 | 4.71 | ||||
qT3δ8-1 | 08 | 20195421-20349908 | 86.57-87.23 | 2.97 | ||||
qT3δ10-1 | 10 | 57733-13172007 | 0.25-56.46 | 3.88 | ||||
qT3δ10-2 | 10 | 19963553-20188240 | 85.58-86.54 | 4.49 | ||||
qT3δ12-1 | 12 | 4509538-4663215 | 19.33-19.99 | 3.16 | ||||
qT3δ12-2 | 12 | 19235536-26139939 | 82.46-112.05 | 4.90 | ||||
δ-toco | qTδ1-1 | 01 | 38537755-39791276 | 165.20-170.57 | 4.28 | |||
qTδ2-1 | 02 | 7215671-7350401 | 30.93-31.51 | 2.89 | ||||
qTδ2-2 | 02 | 31043939-32985884 | 133.08-141.40 | 2.57 | ||||
qTδ3-1 | 03 | 5785084-6008076 | 24.80-25.75 | 3.49 | ||||
qTδ5-1 | 05 | 853216-1082203 | 3.66-4.64 | 3.23 | ||||
qTδ5-2 | 05 | 6983157-7138974 | 29.93-30.60 | 6.17 | ||||
qTδ5-3 | 05 | 13569180-13776389 | 58.17-59.06 | 2.70 | ||||
qTδ6-1 | 06 | 6452921-6720585 | 27.67-28.81 | 3.80 | ||||
qTδ7-1 | 07 | 7351588-15250540 | 31.51-65.37 | 2.73 | ||||
qTδ8-1 | 08 | 1152527-5163244 | 4.94-22.13 | 3.51 | ||||
qTδ9-1 | 09 | 10102157-12363787 | 43.31-53.00 | 3.73 | ||||
qTδ12-1 | 12 | 9212254-15247966 | 39.49-65.36 | 7.16 |
图4 水稻重组自交系维生素E异构体含量的QTL定位 α-t3、γ-t3、δ-t3、α-toco、γ-toco和δ-toco同图1。
Figure 4 QTL mapping of vitamin E isomers content in recombinant inbred lines of rice α-t3, γ-t3, δ-t3, α-toco, γ-toco, and δ-toco are the same as Figure 1.
Chromosome | QTL locus | Gene ID | Gene name | Functional annotation | |
---|---|---|---|---|---|
1 | qT3α/to1, qT3γ1-1 | LOC_Os01g16020 | OsGGR2 | FAD binding domain containing protein, expressed | |
2 | qVE2-1 | LOC_Os02g07160 | OsHPPD | Glyoxalase family protein, putative, expressed | |
2 | qVE2-1, qTto2-1 | LOC_Os02g17650 | OsTC | Tocopherol cyclase, chloroplast precursor, pu- tative, expressed | |
2 | qT3α2-1, qT3α/to2-1, qTγ2-1, qTα/to2-3 | LOC_Os02g47310 | OsγTMT | Cyclopropane-fatty-acyl-phospholipid synthase, putative, expressed | |
2 | qT3α2-1, qT3α/to2-1, qTγ2-1, qTδ2-2, qT3δ2-2, qT3γ2-1 | LOC_Os02g51080 | OsGGR1 | Encodes geranylgeranyl reductase (Li, | |
6 | qTα/to6-4 | LOC_Os06g43880 | OsHGGT | Prenyltransferase, putative, expressed | |
6 | qTα/to6-4 | LOC_Os06g44840 | OsHPT | Encodes homogentisate prenyltransferase (Wang, | |
12 | qT3δ12-2 | LOC_Os12g42090 | OsMPBQ/MT2 | Methyltransferase domain containing protein, putative, expressed |
表4 水稻维生素E积累相关候选基因的功能
Table 4 The function of candidate gene related to accumulation of vitamin E in rice
Chromosome | QTL locus | Gene ID | Gene name | Functional annotation | |
---|---|---|---|---|---|
1 | qT3α/to1, qT3γ1-1 | LOC_Os01g16020 | OsGGR2 | FAD binding domain containing protein, expressed | |
2 | qVE2-1 | LOC_Os02g07160 | OsHPPD | Glyoxalase family protein, putative, expressed | |
2 | qVE2-1, qTto2-1 | LOC_Os02g17650 | OsTC | Tocopherol cyclase, chloroplast precursor, pu- tative, expressed | |
2 | qT3α2-1, qT3α/to2-1, qTγ2-1, qTα/to2-3 | LOC_Os02g47310 | OsγTMT | Cyclopropane-fatty-acyl-phospholipid synthase, putative, expressed | |
2 | qT3α2-1, qT3α/to2-1, qTγ2-1, qTδ2-2, qT3δ2-2, qT3γ2-1 | LOC_Os02g51080 | OsGGR1 | Encodes geranylgeranyl reductase (Li, | |
6 | qTα/to6-4 | LOC_Os06g43880 | OsHGGT | Prenyltransferase, putative, expressed | |
6 | qTα/to6-4 | LOC_Os06g44840 | OsHPT | Encodes homogentisate prenyltransferase (Wang, | |
12 | qT3δ12-2 | LOC_Os12g42090 | OsMPBQ/MT2 | Methyltransferase domain containing protein, putative, expressed |
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