Chinese Journal OF Rice Science ›› 2023, Vol. 37 ›› Issue (2): 113-124.DOI: 10.16819/j.1001-7216.2023.220314
• Research Papers • Next Articles
LIAN Yuanxun1,2,#, WEI Ziyun1,2,#, ZHANG Qiang2, LI Qing2, REN Deyong2, HU Jiang2, ZHU Li2, GAO Zhenyu2, ZHANG Guangheng2, GUO Longbiao2, ZENG Dali2, QIAN Qian2,*(), SHEN Lan2,*()
Received:
2022-03-28
Revised:
2022-05-09
Online:
2023-03-10
Published:
2023-03-10
Contact:
QIAN Qian, SHEN Lan
About author:
First author contact:#These authors contributed equally to this work
廉院训1,2,#, 韦子芸1,2,#, 张强2, 李清2, 任德勇2, 胡江2, 朱丽2, 高振宇2, 张光恒2, 郭龙彪2, 曾大力2, 钱前2,*(), 沈兰2,*()
通讯作者:
钱前,沈兰
作者简介:
第一联系人:#共同第一作者
基金资助:
LIAN Yuanxun, WEI Ziyun, ZHANG Qiang, LI Qing, REN Deyong, HU Jiang, ZHU Li, GAO Zhenyu, ZHANG Guangheng, GUO Longbiao, ZENG Dali, QIAN Qian, SHEN Lan. Identification and Gene Mapping of a Zebra Leaf Mutant zl7 in Rice[J]. Chinese Journal OF Rice Science, 2023, 37(2): 113-124.
廉院训, 韦子芸, 张强, 李清, 任德勇, 胡江, 朱丽, 高振宇, 张光恒, 郭龙彪, 曾大力, 钱前, 沈兰. 水稻斑马叶突变体zl7的鉴定与基因的精细定位[J]. 中国水稻科学, 2023, 37(2): 113-124.
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URL: http://www.ricesci.cn/EN/10.16819/j.1001-7216.2023.220314
标记 Marker | 正向引物序列 Forward primer sequence | 反向引物序列 Reverse primer sequence |
---|---|---|
M1 | GTCCATGCATCCATCTCTAG | ACGGAAGGAATACGTCTGTA |
M2 | TGTGGACAACCTCAACTGAAAGC | CATAATCACCAACATCGGAGAAGC |
M3 | TGTTGAGCTAGAAGAGAGGGG | TGAACACAAAAGGATGCGCT |
M4 | CCAAGTCTTAAGCTACCCCT | CGCAGGGCTTAATAGAATAC |
M5 | GATAGAGCGAGTGAGCAAAC | CCTACCAATTCAACTCCAAC |
M6 | TGGATTGAGGATCAGGATAG | TCTGGAATTTTCCCTAATGA |
M7 | GAAATCAGTCAGAAAGACCG | CCATCTTCTCACTGTGGAGT |
M8 | GCCCTAATTGCTCCAGGTCT | AATTCTAGCAGTGTTCCATTGTG |
M9 | TTCTTCCATGTAGCAAGCATT | CCTACTGCCTGCCAAATCTAT |
Table 1. Primers and sequences used in gene mapping.
标记 Marker | 正向引物序列 Forward primer sequence | 反向引物序列 Reverse primer sequence |
---|---|---|
M1 | GTCCATGCATCCATCTCTAG | ACGGAAGGAATACGTCTGTA |
M2 | TGTGGACAACCTCAACTGAAAGC | CATAATCACCAACATCGGAGAAGC |
M3 | TGTTGAGCTAGAAGAGAGGGG | TGAACACAAAAGGATGCGCT |
M4 | CCAAGTCTTAAGCTACCCCT | CGCAGGGCTTAATAGAATAC |
M5 | GATAGAGCGAGTGAGCAAAC | CCTACCAATTCAACTCCAAC |
M6 | TGGATTGAGGATCAGGATAG | TCTGGAATTTTCCCTAATGA |
M7 | GAAATCAGTCAGAAAGACCG | CCATCTTCTCACTGTGGAGT |
M8 | GCCCTAATTGCTCCAGGTCT | AATTCTAGCAGTGTTCCATTGTG |
M9 | TTCTTCCATGTAGCAAGCATT | CCTACTGCCTGCCAAATCTAT |
基因 Gene | 基因登录号 Gene ID | 定量正向引物序列 Forward primer sequence | 定量反向引物序列 Reverse primer sequence | 功能 Function |
---|---|---|---|---|
NYC1 | LOC_Os01g12710 | CATGCAACACCAACAAAAGG | GACCATTCCAGGAGAAGCAG | 叶绿素b还原酶基因 |
NOL | LOC_Os03g45194 | CCACGAAAGGTATAGGATATG | TCAAGTCAGTCACCGCAGAT | 叶绿素b还原酶基因 |
NYC3 | LOC_Os06g24730 | TCTATCTAGGTGCCAAAGGC | ATTCTGGCACCTGCTGTTTC | α/β折叠水解酶家族蛋白 |
PAO | LOC_Os03g05310 | AAGCCTCCGATGTTACCGAA | CGAGGGTTTCCAGAATTTGA | 脱镁叶绿酸a加氧酶,叶绿体前体 |
SGR | LOC_Os09g36200 | GCAATGTCGCCAAATGACG | GCTCACCACACTCATTCCCTAAAG | 镁离子去螯合酶 |
RCCR1 | LOC_Os10g25020 | GGATCGACGATTGATTTCATG | GTCGAGGCGTTCAGAAAGAT | 红色叶绿素分解代谢还原酶 |
RCCR2 | LOC_Os10g25040 | TGGCGAGGGACAGGAAGGT | GGATGTGGTGGCGAGAGAAAC | 红色叶绿素分解代谢还原酶 |
LchP2 | LOC_Os09g17740 | GAAGAAGATCAAGAACGGCC | TTGCCGGGGACGAAGTTGGT | 捕光叶绿素a/b结合蛋白基因 |
PsbA | LOC_Os12g19580 | AGAGACGCGAAAGTACAAGC | AAGTTGCGGTCAATAAGGTA | 光合反应中心蛋白 |
RpoC2 | LOC_Os04g16830 | ATGCATCGCAGGTACACCAA | CCCTCGCGTAAATTGCTTTG | DNA定向RNA聚合酶亚单位β |
Rps15 | LOC_Os12g10580 | AGATACGGAGACTTGCTTCA | GCTCCCTAATATCCAACTGACT | 核酮糖二磷酸羧化酶大链前体 |
V1 | LOC_Os03g45400 | AGAATCAGCGCGAGAAGAGAACCT | TACACCAGCTTTGGAGGAGCTGAA | 质体蛋白 |
V2 | LOC_Os03g20460 | AGCAGATCCGTGATTACATGGCGA | TGCCTCTTCACTCTCTGCAACCAA | 鸟苷酸激酶 |
YGL8 | LOC_Os01g17170 | TGGATCTAACATGACACGCACCCA | ACTGTAACGGCATTCTTCTCCGGT | 镁原卟啉IX单酯环化酶的催化亚基 |
CAO1 | LOC_Os10g41780 | TTGGCTCAGTTAATGAGGGCAGAATCC | GGATGCGCACGTTGAGCATCTTTGTGG | 叶绿素a加氧酶 |
PORA | LOC_Os04g58200 | ATGGCTCTCCAAGTTCAG | TGGCTCACGCTAAGGAAC | NADPH:原叶绿素酸酯氧化还原酶A |
PORB | LOC_Os10g35370 | CCGCAAGGAGGGAGCGGTG | CCCTCTTGGTGCTAAGGCCG | 原叶绿素酸酯氧化还原酶B |
CHLH | LOC_Os03g20700 | GCACGGGAACTTGGCGTTTCATTA | ACATGTCCTGGAGCTGCTTCTCAT | 镁离子螯合酶H亚基 |
CHLD | LOC_Os03g59640 | TAGCACAGCTGTCAGAGTGGGTTT | TTGCCAGCCACCTCAAGTATCTCA | 镁离子螯合酶D亚基 |
CHLI | LOC_Os03g36540 | AGGGATGCTGAACTCAGGGTGAAA | AAGTAGGACTCACGGAACGCCTTT | 镁离子螯合酶I亚基 |
DVR | LOC_Os03g22780 | AGCCCAGGTTCATCAAGGT | TGATCACCCTCTCGAAGAACT | 联乙烯还原酶基因 |
OsCHLM | LOC_Os06g04150 | GCTTCATCTCCACGCAGTTCTACT | GCAATGACGAATCGAAGACGCACA | 镁原卟啉O-甲基转移酶 |
YGL1 | LOC_Os05g28200 | CCAGCCACTGATGAAAGCAGCAAT | AGAGCGCTAATACACTCGCGAACA | 叶绿素合成酶 |
OsHEMA1 | LOC_Os10g35840 | GATGCAATCACTGCTGGAAAGCGT | CCATCTTGCCAGCACCAATCAACA | 莽草酸/奎宁酸5-脱氢酶 |
OsHEML | LOC_Os08g41990 | AGAACAAAGGGCAGATTGCTGCTG | TGTTTCGTCAAGTCACGGAGAGCA | 转氨酶 |
OsHEMB | LOC_Os06g49110 | TGGCATTGTCAGGGAAGATGGAGT | CCAAAGCAGCACGTATTGCTCCAA | δ-氨基乙酰丙酸脱水酶,叶绿体前体 |
GUN4 | LOC_Os11g16550 | AAGGGAAGGAGAGGCCAAAGTTCA | ACCATGACCAGCATCTCTGCATCA | 镁离子螯合酶H亚基结合蛋白 |
Table 2. Genes and primer sequences used in qRT-PCR.
基因 Gene | 基因登录号 Gene ID | 定量正向引物序列 Forward primer sequence | 定量反向引物序列 Reverse primer sequence | 功能 Function |
---|---|---|---|---|
NYC1 | LOC_Os01g12710 | CATGCAACACCAACAAAAGG | GACCATTCCAGGAGAAGCAG | 叶绿素b还原酶基因 |
NOL | LOC_Os03g45194 | CCACGAAAGGTATAGGATATG | TCAAGTCAGTCACCGCAGAT | 叶绿素b还原酶基因 |
NYC3 | LOC_Os06g24730 | TCTATCTAGGTGCCAAAGGC | ATTCTGGCACCTGCTGTTTC | α/β折叠水解酶家族蛋白 |
PAO | LOC_Os03g05310 | AAGCCTCCGATGTTACCGAA | CGAGGGTTTCCAGAATTTGA | 脱镁叶绿酸a加氧酶,叶绿体前体 |
SGR | LOC_Os09g36200 | GCAATGTCGCCAAATGACG | GCTCACCACACTCATTCCCTAAAG | 镁离子去螯合酶 |
RCCR1 | LOC_Os10g25020 | GGATCGACGATTGATTTCATG | GTCGAGGCGTTCAGAAAGAT | 红色叶绿素分解代谢还原酶 |
RCCR2 | LOC_Os10g25040 | TGGCGAGGGACAGGAAGGT | GGATGTGGTGGCGAGAGAAAC | 红色叶绿素分解代谢还原酶 |
LchP2 | LOC_Os09g17740 | GAAGAAGATCAAGAACGGCC | TTGCCGGGGACGAAGTTGGT | 捕光叶绿素a/b结合蛋白基因 |
PsbA | LOC_Os12g19580 | AGAGACGCGAAAGTACAAGC | AAGTTGCGGTCAATAAGGTA | 光合反应中心蛋白 |
RpoC2 | LOC_Os04g16830 | ATGCATCGCAGGTACACCAA | CCCTCGCGTAAATTGCTTTG | DNA定向RNA聚合酶亚单位β |
Rps15 | LOC_Os12g10580 | AGATACGGAGACTTGCTTCA | GCTCCCTAATATCCAACTGACT | 核酮糖二磷酸羧化酶大链前体 |
V1 | LOC_Os03g45400 | AGAATCAGCGCGAGAAGAGAACCT | TACACCAGCTTTGGAGGAGCTGAA | 质体蛋白 |
V2 | LOC_Os03g20460 | AGCAGATCCGTGATTACATGGCGA | TGCCTCTTCACTCTCTGCAACCAA | 鸟苷酸激酶 |
YGL8 | LOC_Os01g17170 | TGGATCTAACATGACACGCACCCA | ACTGTAACGGCATTCTTCTCCGGT | 镁原卟啉IX单酯环化酶的催化亚基 |
CAO1 | LOC_Os10g41780 | TTGGCTCAGTTAATGAGGGCAGAATCC | GGATGCGCACGTTGAGCATCTTTGTGG | 叶绿素a加氧酶 |
PORA | LOC_Os04g58200 | ATGGCTCTCCAAGTTCAG | TGGCTCACGCTAAGGAAC | NADPH:原叶绿素酸酯氧化还原酶A |
PORB | LOC_Os10g35370 | CCGCAAGGAGGGAGCGGTG | CCCTCTTGGTGCTAAGGCCG | 原叶绿素酸酯氧化还原酶B |
CHLH | LOC_Os03g20700 | GCACGGGAACTTGGCGTTTCATTA | ACATGTCCTGGAGCTGCTTCTCAT | 镁离子螯合酶H亚基 |
CHLD | LOC_Os03g59640 | TAGCACAGCTGTCAGAGTGGGTTT | TTGCCAGCCACCTCAAGTATCTCA | 镁离子螯合酶D亚基 |
CHLI | LOC_Os03g36540 | AGGGATGCTGAACTCAGGGTGAAA | AAGTAGGACTCACGGAACGCCTTT | 镁离子螯合酶I亚基 |
DVR | LOC_Os03g22780 | AGCCCAGGTTCATCAAGGT | TGATCACCCTCTCGAAGAACT | 联乙烯还原酶基因 |
OsCHLM | LOC_Os06g04150 | GCTTCATCTCCACGCAGTTCTACT | GCAATGACGAATCGAAGACGCACA | 镁原卟啉O-甲基转移酶 |
YGL1 | LOC_Os05g28200 | CCAGCCACTGATGAAAGCAGCAAT | AGAGCGCTAATACACTCGCGAACA | 叶绿素合成酶 |
OsHEMA1 | LOC_Os10g35840 | GATGCAATCACTGCTGGAAAGCGT | CCATCTTGCCAGCACCAATCAACA | 莽草酸/奎宁酸5-脱氢酶 |
OsHEML | LOC_Os08g41990 | AGAACAAAGGGCAGATTGCTGCTG | TGTTTCGTCAAGTCACGGAGAGCA | 转氨酶 |
OsHEMB | LOC_Os06g49110 | TGGCATTGTCAGGGAAGATGGAGT | CCAAAGCAGCACGTATTGCTCCAA | δ-氨基乙酰丙酸脱水酶,叶绿体前体 |
GUN4 | LOC_Os11g16550 | AAGGGAAGGAGAGGCCAAAGTTCA | ACCATGACCAGCATCTCTGCATCA | 镁离子螯合酶H亚基结合蛋白 |
Fig. 1. Phenotypes of wild type CJ06 and zl7 at different growth stages. A, B and C are the phenotypes of wild-type Chunjiang06 and zl7 plants at seedling stage, tillering stage, and heading stage. The white box in figure B and C are enlarged leaves. D, E and F are chlorophyll contents at seedling stage, tillering stage and heading stage, respectively; Chla, Chlorophyll a; Chlb, Chlorophyll b; Car, Carotenoid; Means±SD(n=3); *,** indicates that there is significant difference between the wild type and the mutant at the level of 0.05 and 0.01 (t-test).
农艺性状 Agronomic trait | 野生型 Wild type | 突变体 Mutant zl7 |
---|---|---|
株高 Plant height/cm | 91.2±4.0 | 79.5±2.7** |
分蘖 Tillering number | 10.0±1.2 | 8.0±0.8* |
穗长 Panicle length/cm | 22.5±1.3 | 20.2±0.8* |
一次枝梗 Primary rachis branches | 17.0±0.5 | 14.0±1.1* |
二次枝梗 Secondary rachis branches | 28.0±1.9 | 23.0±1.6* |
每穗粒数 No. of grains per panicle | 152.0±6.0 | 126.0±4.7** |
结实率Seed setting rate/% | 93.4±0.0 | 93.7±0.0 |
千粒重 1000-grain weight/g | 23.2±0.6 | 24.9±0.4 |
粒长 Grain length/mm | 7.35±0.40 | 7.67±0.63 |
粒宽 Grain width/mm | 3.08±0.23 | 3.17±0.24 |
Table 3. Comparison of main agronomic traits between wild type Chunjiang 06 and mutant zl7.
农艺性状 Agronomic trait | 野生型 Wild type | 突变体 Mutant zl7 |
---|---|---|
株高 Plant height/cm | 91.2±4.0 | 79.5±2.7** |
分蘖 Tillering number | 10.0±1.2 | 8.0±0.8* |
穗长 Panicle length/cm | 22.5±1.3 | 20.2±0.8* |
一次枝梗 Primary rachis branches | 17.0±0.5 | 14.0±1.1* |
二次枝梗 Secondary rachis branches | 28.0±1.9 | 23.0±1.6* |
每穗粒数 No. of grains per panicle | 152.0±6.0 | 126.0±4.7** |
结实率Seed setting rate/% | 93.4±0.0 | 93.7±0.0 |
千粒重 1000-grain weight/g | 23.2±0.6 | 24.9±0.4 |
粒长 Grain length/mm | 7.35±0.40 | 7.67±0.63 |
粒宽 Grain width/mm | 3.08±0.23 | 3.17±0.24 |
材料 Material | 光合作用速率 Pn/ (μmol·m−2 s−1) | 气孔导度 Gs / (mol·m−2 s−1) | 胞间CO2浓度 Ci / (μmol·mol−1) | 蒸腾速率 Tr/ (mol·m−2 s−1) |
---|---|---|---|---|
野生型WT | 26.41±0.34 | 0.06±0.01 | 442.34±24.64 | 2.94±0.08 |
Mutant-g | 21.67±0.51 ** | 0.05±0.01 | 246.63±29.41** | 2.83±0.32 |
Mutant-y | 13.21±0.47 ** | 0.04±0.01 | 133.69±7.49** | 2.34±0.17* |
Table 4. Photosynthetic parameters of the wild type CJ06 and the mutant zl7 at peak tillering stage.
材料 Material | 光合作用速率 Pn/ (μmol·m−2 s−1) | 气孔导度 Gs / (mol·m−2 s−1) | 胞间CO2浓度 Ci / (μmol·mol−1) | 蒸腾速率 Tr/ (mol·m−2 s−1) |
---|---|---|---|---|
野生型WT | 26.41±0.34 | 0.06±0.01 | 442.34±24.64 | 2.94±0.08 |
Mutant-g | 21.67±0.51 ** | 0.05±0.01 | 246.63±29.41** | 2.83±0.32 |
Mutant-y | 13.21±0.47 ** | 0.04±0.01 | 133.69±7.49** | 2.34±0.17* |
材料 Material | 光合作用速率 Pn/ (μmol·m−2·s−1) | 气孔导度 Gs / (mol·m−2·s−1) | 胞间CO2浓度 Ci / (μmol·mol−1) | 蒸腾速率 Tr/ (mol·m−2·s−1) |
---|---|---|---|---|
WT | 16.06±0.56 | 0.15±0.02 | 204.24±17.42 | 3.29±0.30 |
Mutant-g | 18.34±0.64** | 0.28±0.02* | 278.66±9.48** | 4.79±0.26** |
Mutant-y | 9.93±1.41** | 0.13±0.03 | 262.71±15.41** | 2.71±0.58 |
Table 5. Photosynthetic parameters of wild type CJ06 and mutant zl7 at heading stage.
材料 Material | 光合作用速率 Pn/ (μmol·m−2·s−1) | 气孔导度 Gs / (mol·m−2·s−1) | 胞间CO2浓度 Ci / (μmol·mol−1) | 蒸腾速率 Tr/ (mol·m−2·s−1) |
---|---|---|---|---|
WT | 16.06±0.56 | 0.15±0.02 | 204.24±17.42 | 3.29±0.30 |
Mutant-g | 18.34±0.64** | 0.28±0.02* | 278.66±9.48** | 4.79±0.26** |
Mutant-y | 9.93±1.41** | 0.13±0.03 | 262.71±15.41** | 2.71±0.58 |
杂交组合 Hybrid combination | F1 | F2 | χ2 (3:1) | ||
---|---|---|---|---|---|
正常表型 Wild-type phenotype | 突变表型 zl7 phenotype | 正常表型 Wild-type phenotype | 突变表型 zl7 phenotype | ||
zl7/93-11 | 13 | 0 | 2065 | 655 | 1.225 |
93-11/ zl7 | 8 | 0 | 822 | 271 | 0.025 |
Table 6. Genetic analysis of the mutant zl7.
杂交组合 Hybrid combination | F1 | F2 | χ2 (3:1) | ||
---|---|---|---|---|---|
正常表型 Wild-type phenotype | 突变表型 zl7 phenotype | 正常表型 Wild-type phenotype | 突变表型 zl7 phenotype | ||
zl7/93-11 | 13 | 0 | 2065 | 655 | 1.225 |
93-11/ zl7 | 8 | 0 | 822 | 271 | 0.025 |
预测基因 Predictive gene | 基因功能注释 Gene function annotation |
---|---|
ORF1 | 转座子蛋白 |
ORF2 | 表达蛋白 |
ORF3 | ζ-胡萝卜素脱氢酶 |
ORF4 | 表达蛋白 |
ORF5 | 细胞色素b-c1复合亚单位7 |
ORF6 | 表达蛋白 |
ORF7 | 核苷酸结合蛋白 |
ORF8 | 羧基末端肽酶 |
ORF9 | 表达蛋白 |
ORF10 | 肽酶,M24家族蛋白 |
ORF11 | 细胞周期蛋白相关蛋白 |
ORF12 | 表达蛋白 |
Table 7. Gene function annotation of ORFs.
预测基因 Predictive gene | 基因功能注释 Gene function annotation |
---|---|
ORF1 | 转座子蛋白 |
ORF2 | 表达蛋白 |
ORF3 | ζ-胡萝卜素脱氢酶 |
ORF4 | 表达蛋白 |
ORF5 | 细胞色素b-c1复合亚单位7 |
ORF6 | 表达蛋白 |
ORF7 | 核苷酸结合蛋白 |
ORF8 | 羧基末端肽酶 |
ORF9 | 表达蛋白 |
ORF10 | 肽酶,M24家族蛋白 |
ORF11 | 细胞周期蛋白相关蛋白 |
ORF12 | 表达蛋白 |
Fig. 4. Expression analysis of chloroplast development related genes of the wild type Chunjiang 06(CJ06) and its mutant zl7. Mean±SD(n=3). *, ** indicates that there is significant difference between wild type(CJ06) and mutant at the level of 0.05 and 0.01 respectively (t-test)
Fig. 6. Expression analysis of chlorophyll degradation related genes of the wild type Chunjiang 06(CJ06) and its mutant zl7. Mean±SD (n=3). *, ** indicate that there is significant difference between the wild type and the mutant at the level of 0.05 and 0.01 respectively (t-test).
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