中国水稻科学 ›› 2023, Vol. 37 ›› Issue (2): 113-124.DOI: 10.16819/j.1001-7216.2023.220314
• 研究报告 • 下一篇
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,*()
收稿日期:
2022-03-28
修回日期:
2022-05-09
出版日期:
2023-03-10
发布日期:
2023-03-10
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 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
摘要:
【目的】斑马叶突变体作为水稻叶色突变体的重要种质资源,是研究植物光合作用机制和高光效育种的理想材料,对于解析光合作用机理和提高水稻产量具有重要意义。【方法】用甲基磺酸乙酯(EMS)诱变粳稻品种春江06建立突变体库,从突变体库中筛选到1份苗期为斑马叶的突变体,该突变体被命名为zl7 (zebra leaf 7)。在常规大田种植条件下分别比较突变体与野生型在苗期、抽穗期和成熟期叶色表型和产量性状差异,通过透射电镜实验分析叶片叶绿体发育情况,利用图位克隆方法克隆候选基因,利用荧光定量PCR 方法分析参与叶绿素合成和叶绿体发育相关基因的表达水平。【结果】从苗期开始,突变体zl7表现出典型的斑马叶,叶绿素含量降低,直到抽穗期,斑马叶表型消失,叶片逐渐复绿,叶绿素含量无明显差异。光合速率测定和电镜观察结果显示,突变体zl7的光合速率、气孔导度下降,叶绿体发育异常。与野生型相比,突变体的株高、分蘖、穗长、一次枝梗、二次枝梗和每穗粒数均显著降低,而粒长、粒宽和千粒重均略有增加。荧光定量PCR结果表明突变体中参与叶绿素降解相关基因的表达量显著升高,而参与叶绿素合成和叶绿体发育相关基因的表达量显著降低。遗传分析表明,该突变体受一对隐性核基因调控。通过图位克隆将该基因定位在第7染色体,测序发现突变体的目标基因ZL7编码区发生单碱基替换,导致一个氨基酸由丝氨酸变为天冬酰胺。【结论】ZL7 (Zebra Leaf 7)突变导致叶绿体发育异常,水稻叶片出现斑马叶表型,该基因在叶绿素合成及叶绿体发育中起重要作用。
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.
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.
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 |
表1 基因定位所用引物及序列
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 |
表2 qRT-PCR所用基因及引物序列
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 |
图1 野生型和突变体zl7不同生育期的表型 A,B,C分别为苗期、分蘖期和抽穗期野生型春江06和突变体zl7植株表型,B, C白框中为放大的叶片。D,E,F分别为苗期、分蘖期和抽穗期的叶绿素含量;Chla―叶绿素a,Chlb―叶绿素b,Car―类胡萝卜素;误差线表示3次生物学重复试验的标准差;*,**分别表示野生型与突变体在0.05和0.01水平下差异显著(t检验)。
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 |
表3 野生型春江06与突变体zl7的主要农艺性状比较
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* |
表4 野生型CJ06和突变体zl7分蘖盛期光合参数
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 |
表5 野生型CJ06和突变体zl7抽穗期光合参数
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 |
表6 突变体zl7的遗传分析
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 |
表7 开放阅读框基因注释
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 |
图4 野生型春江06(CJ06)及其突变体zl7叶绿体发育相关基因的表达量分析 误差线表示3次生物学重复试验的标准差。*和**分别表示野生型春江 06(CJ06) 和突变体间差异达 0.05 和 0.01 显著水平(t 检验)。
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)
图5 野生型春江06(CJ06)及其突变体zl7叶绿素合成相关基因的表达量分析
Fig. 5. Expression analysis of chlorophyll synthesis related genes of the wild type Chunjiang 06(CJ06) and its mutant zl7.
图6 野生型春江06(CJ06)及其突变体zl7叶绿素降解相关基因的表达量分析 误差线表示 3 次生物学重复试验的标准差。*和**分别表示野生型春江 06 和突变体间差异达 0.05 和 0.01 显著水平(t 检验)。
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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