园艺学报 ›› 2023, Vol. 50 ›› Issue (5): 933-946.doi: 10.16420/j.issn.0513-353x.2022-0159
张琨, 思彬彬, 周军, 任玉锋, 张欣, 徐文娣(), 王佳伟, 乔帅, 王惠冉
收稿日期:
2022-11-04
修回日期:
2023-01-18
出版日期:
2023-05-25
发布日期:
2023-05-31
通讯作者:
*(E-mail:xuwendi2018@nmu.edu.cn)基金资助:
ZHANG Kun, SI Binbin, ZHOU Jun, REN Yufeng, ZHANG Xin, XU Wendi(), WANG Jiawei, QIAO Shuai, WANG Huiran
Received:
2022-11-04
Revised:
2023-01-18
Published:
2023-05-25
Online:
2023-05-31
摘要:
MLO(Mildew Resistance Locus O)家族基因是植物特有的“感病基因”,普遍认为它对白粉病具有负调控作用。目前在单子叶和双子叶植物中发现许多抗白粉病MLO基因,但对于其上游调控因子如何调控其表达而影响白粉病发生的研究却很少。本研究中以易感白粉病的苹果砧木‘青砧1号’叶片为材料,构建其cDNA文库,文库滴度4 × 109 cfu · mL-1,重组率为100%。生物信息分析苹果MdMLO家族基因启动子区共有38种顺式作用元件,包含激素、应激防御和生长发育等响应元件。以TCA和TC-rich repeats为诱饵序列构建诱饵载体,分别命名为Bait-TCA和Bait-TC。利用酵母单杂交的方法筛选其上游调控因子,结果显示Bait-TCA具有自激活现象无法筛选,Bait-TC诱饵载体调取获得30条基因序列,其中6条参与植物抗逆防御反应,推测它们可能通过与TC-rich repeats元件互作来调控MLO的表达,进而调控苹果叶片对于白粉病的响应。
中图分类号:
张琨, 思彬彬, 周军, 任玉锋, 张欣, 徐文娣, 王佳伟, 乔帅, 王惠冉. 苹果砧木‘青砧1号’叶片cDNA文库构建及MdMLO上游调控因子的筛选[J]. 园艺学报, 2023, 50(5): 933-946.
ZHANG Kun, SI Binbin, ZHOU Jun, REN Yufeng, ZHANG Xin, XU Wendi, WANG Jiawei, QIAO Shuai, WANG Huiran. Construction of cDNA Library of Apple Rootstock‘Qingzhen 1’Leaf and Screen of MdMLO Genes’ Upstream Regulator[J]. Acta Horticulturae Sinica, 2023, 50(5): 933-946.
图1 ‘青砧1号’叶片RNA的提取及cDNA的合成 M:Marker;1:总RNA;2:全长cDNA;3:cDNA均一化;4:cDNA短片段去除。
Fig. 1 RNA extraction and cDNA synthesis from‘Qingzhen 1’leaf M:Marker;1:Total RNA;2: Full-length cDNA;3:cDNA homogenization;4:Purifying of the cDNA.
元件名称 Element name | 元件序列 Element sequence | 功能 Function |
---|---|---|
AAAC-motif | CAATCAAAACCT | 光响应元件 Light responsive element |
ACE | GACACGTATG | 参与光反应元件 cis-Acting element involved in light responsiveness |
AE-box | AGAAACAA | 光响应组件的一部分 Part of a module for light response |
AT1-motif | AATTATTTTTTATT | 光响应组件的一部分 Part of a light responsive module |
ATCT-motif | AATCTAATCC | 光响应DNA保守元件的一部分 Part of a conserved DNA module involved in light responsiveness |
chs-Unit 1 m1 | ACCTAACCCGG | 光响应元件的一部分 Part of a light responsive element |
G-Box | CACGTT | 光反应元件 cis-Acting regulatory element involved in light responsiveness |
MRE | AACCTAA | MYB结合位点参与光响应 MYB binding site involved in light responsiveness |
GARE-motif | TCTGTTG | 赤霉素响应元件 Gibberellin-responsive element |
AuxRE | TGTCTCAATAAG | 生长素响应元件的一部分 Part of an auxin-responsive element |
AuxRR-core | GGTCCAT | 生长素响应元件 cis-Acting regulatory element involved in auxin responsiveness |
ABRE | ACGTG | 脱落酸响应元件 cis-Acting element involved in the abscisic acid responsiveness |
P-box | CCTTTTG | 赤霉素响应元件 Gibberellin-responsive element |
TGACG-motif | TGACG | 茉莉酸甲酯响应元件 cis-Acting regulatory element involved in the MeJA-responsiveness |
TGA-element | AACGAC | 生长素响应元件 Auxin-responsive element |
TATC-box | TATCCCA | 赤霉素响应元件 cis-Acting element involved in gibberellin-responsiveness |
circadian | CAAAGATATC | 响应昼夜节律控制元件 cis-Acting regulatory element involved in circadian control |
AACA_motif | TAACAAACTCCA | 胚乳特异性负表达元件 Involved in endosperm-specific negative expression |
AT-rich element | ATAGAAATCAA | AT富集DNA结合蛋白位点(ATBP-1) Binding site of AT-rich DNA binding protein(ATBP-1) |
CAT-box | GCCACT | 分生组织表达相关元件 cis-Acting regulatory element related to meristem expression |
GCN4_motif | TGAGTCA | 胚乳表达响应元件 cis-Regulatory element involved in endosperm expression |
HD-Zip 1 | CAAT(A/T)ATTG | 栅栏叶肉细胞分化响应元件 Element involved in differentiation of the palisade mesophyll cells |
HD-Zip 3 | GTAAT(G/C)ATTAC | 蛋白结合位点 Protein binding site |
MSA-like | TCCAACGGT | 细胞周期调节元件cis-Acting element involved in cell cycle regulation |
O2-site | GATGATGTGG | 玉米醇溶蛋白代谢调节元件cis-Acting regulatory element involved in zein metabolism regulation |
RY-element | CATGCATG | 种子特异调控元件 cis-Acting regulatory element involved in seed-specific regulation |
Unnamed_1 | GAATTTAATTAA | 60K蛋白结合位点 60K protein binding site |
DRE | TACCGACAT | 脱水、低温、盐响应元件 cis-Acting element involved in dehydration,low-temp,salt stresses |
CCAAT-box | CAACGG | HvMYB1结合位点 HvMYB1 binding site |
GC-motif | CCCCCG | 增强缺氧特异性诱导元件 Enhancer-like element involved in anoxic specific inducibility |
LTR | CCGAAA | 低温响应元件 cis-Acting element involved in low-temperature responsiveness |
MBS | CAACTG | MYB结合位点参与干旱诱导 MYB binding site involved in drought-inducibility |
TCA-element | CCATCTTTTT | 水杨酸响应元件 cis-Acting element involved in salicylic acid responsiveness |
TC-rich repeats | GTTTTCTTAC | 响应防御和应激反应元件 cis-Acting element involved in defense and stress responsiveness |
ARE | AAACCA | 厌氧诱导关键元件 cis-Acting regulatory element essential for the anaerobic induction |
WUN-motif | AAATTTCCT | 干旱响应元件Wound-responsive element |
CAAT-box | CCAAT | 启动子增强子区域元件Common cis-acting element in promoter and enhancer regions |
TATA-box | TATA | 启动子核心元件Core promoter element around -30 of transcription start |
表1 MLO基因家族顺式作用元件分析
Table 1 cis-Acting element analysis of MLO gene family
元件名称 Element name | 元件序列 Element sequence | 功能 Function |
---|---|---|
AAAC-motif | CAATCAAAACCT | 光响应元件 Light responsive element |
ACE | GACACGTATG | 参与光反应元件 cis-Acting element involved in light responsiveness |
AE-box | AGAAACAA | 光响应组件的一部分 Part of a module for light response |
AT1-motif | AATTATTTTTTATT | 光响应组件的一部分 Part of a light responsive module |
ATCT-motif | AATCTAATCC | 光响应DNA保守元件的一部分 Part of a conserved DNA module involved in light responsiveness |
chs-Unit 1 m1 | ACCTAACCCGG | 光响应元件的一部分 Part of a light responsive element |
G-Box | CACGTT | 光反应元件 cis-Acting regulatory element involved in light responsiveness |
MRE | AACCTAA | MYB结合位点参与光响应 MYB binding site involved in light responsiveness |
GARE-motif | TCTGTTG | 赤霉素响应元件 Gibberellin-responsive element |
AuxRE | TGTCTCAATAAG | 生长素响应元件的一部分 Part of an auxin-responsive element |
AuxRR-core | GGTCCAT | 生长素响应元件 cis-Acting regulatory element involved in auxin responsiveness |
ABRE | ACGTG | 脱落酸响应元件 cis-Acting element involved in the abscisic acid responsiveness |
P-box | CCTTTTG | 赤霉素响应元件 Gibberellin-responsive element |
TGACG-motif | TGACG | 茉莉酸甲酯响应元件 cis-Acting regulatory element involved in the MeJA-responsiveness |
TGA-element | AACGAC | 生长素响应元件 Auxin-responsive element |
TATC-box | TATCCCA | 赤霉素响应元件 cis-Acting element involved in gibberellin-responsiveness |
circadian | CAAAGATATC | 响应昼夜节律控制元件 cis-Acting regulatory element involved in circadian control |
AACA_motif | TAACAAACTCCA | 胚乳特异性负表达元件 Involved in endosperm-specific negative expression |
AT-rich element | ATAGAAATCAA | AT富集DNA结合蛋白位点(ATBP-1) Binding site of AT-rich DNA binding protein(ATBP-1) |
CAT-box | GCCACT | 分生组织表达相关元件 cis-Acting regulatory element related to meristem expression |
GCN4_motif | TGAGTCA | 胚乳表达响应元件 cis-Regulatory element involved in endosperm expression |
HD-Zip 1 | CAAT(A/T)ATTG | 栅栏叶肉细胞分化响应元件 Element involved in differentiation of the palisade mesophyll cells |
HD-Zip 3 | GTAAT(G/C)ATTAC | 蛋白结合位点 Protein binding site |
MSA-like | TCCAACGGT | 细胞周期调节元件cis-Acting element involved in cell cycle regulation |
O2-site | GATGATGTGG | 玉米醇溶蛋白代谢调节元件cis-Acting regulatory element involved in zein metabolism regulation |
RY-element | CATGCATG | 种子特异调控元件 cis-Acting regulatory element involved in seed-specific regulation |
Unnamed_1 | GAATTTAATTAA | 60K蛋白结合位点 60K protein binding site |
DRE | TACCGACAT | 脱水、低温、盐响应元件 cis-Acting element involved in dehydration,low-temp,salt stresses |
CCAAT-box | CAACGG | HvMYB1结合位点 HvMYB1 binding site |
GC-motif | CCCCCG | 增强缺氧特异性诱导元件 Enhancer-like element involved in anoxic specific inducibility |
LTR | CCGAAA | 低温响应元件 cis-Acting element involved in low-temperature responsiveness |
MBS | CAACTG | MYB结合位点参与干旱诱导 MYB binding site involved in drought-inducibility |
TCA-element | CCATCTTTTT | 水杨酸响应元件 cis-Acting element involved in salicylic acid responsiveness |
TC-rich repeats | GTTTTCTTAC | 响应防御和应激反应元件 cis-Acting element involved in defense and stress responsiveness |
ARE | AAACCA | 厌氧诱导关键元件 cis-Acting regulatory element essential for the anaerobic induction |
WUN-motif | AAATTTCCT | 干旱响应元件Wound-responsive element |
CAAT-box | CCAAT | 启动子增强子区域元件Common cis-acting element in promoter and enhancer regions |
TATA-box | TATA | 启动子核心元件Core promoter element around -30 of transcription start |
图5 TCA- element诱饵质粒载体(1、2)和TC-rich repeats诱饵质粒载体(3、4)
Fig. 5 Construction of TCA-element bait plasmid vector(1,2)and TC-rich repeats bait plasmid vector(3,4)
分类 | 比对结果 | 功能注释 | NCBI登录号 | |
---|---|---|---|---|
Classify | Alignment result | Functional annotation | GenBank ID | |
信号转导 Signal transduc- tion | NADH脱氢酶1 α亚复合体亚基1 NADH dehydrogenase 1 alpha subcomplex subunit 1 | 将电子从NADH转移到线粒体膜呼吸链 Transfer of electrons from NADH to the respiratory chain | XM_029089752.1 | |
空泡分选受体3类似物 Vacuolar-sorting receptor 3-like | 参与网格蛋白包被的囊泡从高尔基体到空泡的分选 Sorting of clathrin-coated vesicles from golgi bodies to vacuoles | XM_008386294.3 | ||
细胞色素b-c1复合体亚基4 Cytochrome b-c1 complex subunit Rieske-4 | 驱动线粒体氧化磷酸化 Drives mitochondrial oxidative phosphorylation | XR_003767947.1 | ||
细胞色素c氧化酶亚基5C-3 Cytochrome c oxidase subunit 5C-3 | 细胞色素c氧化酶的核编码多肽链之一,是线粒体电子传递的末端氧化酶 One of the nucleus-coding polypeptide chains of cytochrome C oxidase,the terminal oxidase of mitochondrial electron transport | XM_008370823.3 | ||
钾离子通道AKT2/3 Potassium channel AKT2/3 | 参与韧皮部汁液中输入和卸载钾的作用 Involved in both loading and unloading potassium into/from the phloem sap | XM_029107389.1 | ||
三酰甘油脂肪酶1 Triacylglycerol lipase 1 | 催化三酰基甘油的水解 Catalyzes the hydrolysis of triacylglycerol | XM_008340013.3 | ||
硫氧还原蛋白AAED1类似物 Thioredoxin-like protein AAED1 | 抗氧化活性 Antioxidant activity | XM_008343896.3 | ||
泛酸酯激酶 At2g17340-like | 催化泛酸的磷酸化 Catalyzes the phosphorylation of pantothenate | XM_008384280.3 | ||
非典型N-乙酰转移酶p20类似物 Uncharacterized N-acetyltransferase p20-like | 参与N-乙酰转移酶活性 N-acetyltransferase activity | XM_029096249.1 | ||
转录调控 Transcrip- tional regulation | 核糖核酸内切酶YBEY Endoribonuclease YBEY | 参与晚期70S核糖体质量控制和16S rRNA 3′端成熟 Involved in late-stage 70S ribosome quality control and in maturation of the 3′ terminus of the 16S rRNA | XM_029106325.1 | |
60S核糖体蛋白L10类似物 60S ribosomal protein L10-like | 在核糖体的形成中起积极作用 An active role in the formation of ribosomes | XM_029102547.1 | ||
抗逆防御 Resistance and defense | 脂肪酶PAD4类似物 Lipase-like PAD4 | 促进植物对入侵性生物营养病原体的先天免疫和识别微生物相关分子模式防御机制 Contributing to the plant innate immunity against invasive biotrophic pathogens and to defense mechanisms upon recognition of microbe-associated molecular patterns | XM_029094810.1 | |
应激反应蛋白NST1类似物 Stress response protein NST1-like | 与MSL1一起作为耐盐性的负调节因子 Negative regulator of salt tolerance along with MSL1 | XM_008394867.3 | ||
非特异性脂质转运蛋白3 Non-specific lipid-transfer protein 3 | 可以跨膜转移磷脂和半乳糖脂,参与植株对生物与非生物胁迫的抗性 Phospholipids and galactose lipids can be transferred across the membrane to participate in plant resistance to biological and abiotic stress | XM_008371792.3 | ||
解毒蛋白35类似物 Protein DETOXIFICATION 35-like | 参与类黄酮代谢的多药物和毒素外排转运体 Multidrug and toxin efflux transporters involved in flavonoid metabolism | XM_008381794.3 | ||
单链DNA结合蛋白WHY1 Single stranded DNA binding protein WHY1 | 致病相关基因PR-10a的转录激活因子 Transcriptional activator of the pathogenesis-related gene PR-10a | NM_001294042.1 | ||
脂质转移蛋白VAS类似物 Lipid transfer-like protein VAS | 可以跨膜转移磷脂和半乳糖脂,参与植株对生物与非生物胁迫的抗性 Phospholipids and galactose lipids can be transferred across the membrane to participate in plant resistance to biological and abiotic stress | XM_008342321.3 | ||
生长发育 Grow and develop- ment | MADS-box蛋白 SOC1类似物 MADS-box protein SOC1-like | 调控开花时间的转录因子 Transcription factors that regulate flowering time | XM_008378604.3 | |
质体转录活性蛋白7 Plastid transcriptionally active 7 | 参与质体基因表达,是类囊体形成所必需的 Involved in plastid gene expression,essential for thylakoid formation | XM_008382937.3 | ||
核酸外切酶DPD1 Exonuclease DPD1 | 花粉发育过程中细胞器DNA降解所需的外切酶 Exonuclease enzymes required for degradation of organelle DNA during pollen development | XM_017336537.2 | ||
光合作用NDH亚基的腔定位5 Photosynthetic NDH subunit of lumenal location 5 | 叶绿体NAD(P)H脱氢酶复合物形成和活性所必需 Essential for chloroplast NAD(P)H dehydrogenase complex formation and activity | XM_029091434.1 | ||
锌指蛋白CONSTANS-LIKE13类似物 Zinc finger protein CONSTANS-LIKE 13-like | 促进开花 Accelerate bloom | XM_029100386.1 | ||
二氨基庚二酸表异构酶 Diaminopimelate epimerase | 参与L-赖氨酸生物合成和肽聚糖中五肽的生物合成 Involved in the L-lysine biosynthesis and in the biosynthesis of the pentapeptide incorporated in the peptidoglycan moiety | XM_029101158.1 | ||
内质网素同源物 Endoplasmin homolog | 与芽、根和花分生组织形成及花粉管伸长有关 Related to shoot apical meristem,root apical meristem floral meristem formation and pollen tube elongation | XM_009336652.2 | ||
碳酸酐酶2类似物 Carbonic anhydrase 2-like | 催化二氧化碳的可逆水合作用形成碳酸氢盐 Catalyzes the reversible hydration of carbon dioxide to form bicarbonate | XM_008359520.3 | ||
叶绿体 Chloroplast | 从细胞质向叶绿体输入核编码的光合前蛋白 Nucleo-coded prephotosynthetic proteins are imported from the cytoplasm to chloroplasts | OK545754.1 | ||
LRR受体样丝氨酸/苏氨酸蛋白激酶 LRR receptor-like serine/threonine-protein kinase | 花粉发育的关键调控因子 Key regulator of anther development | XM_029088102.1 | ||
生育酚-O-甲基转移酶 Tocopherol O-methyltransferase | 参与生育酚的合成 Involved in the synthesis of tocopherol | XM_029092547.1 | ||
脂肪酸去饱和酶4 Fatty acid desaturase 4 | 参与生产含有叶绿体特异性的磷脂酰甘油 Involved in the production of chloroplast-specific phosphatidylglycerol | XM_008373042.2 |
表2 阳性克隆测序结果分析
Table 2 Analysis of positive clone sequencing results
分类 | 比对结果 | 功能注释 | NCBI登录号 | |
---|---|---|---|---|
Classify | Alignment result | Functional annotation | GenBank ID | |
信号转导 Signal transduc- tion | NADH脱氢酶1 α亚复合体亚基1 NADH dehydrogenase 1 alpha subcomplex subunit 1 | 将电子从NADH转移到线粒体膜呼吸链 Transfer of electrons from NADH to the respiratory chain | XM_029089752.1 | |
空泡分选受体3类似物 Vacuolar-sorting receptor 3-like | 参与网格蛋白包被的囊泡从高尔基体到空泡的分选 Sorting of clathrin-coated vesicles from golgi bodies to vacuoles | XM_008386294.3 | ||
细胞色素b-c1复合体亚基4 Cytochrome b-c1 complex subunit Rieske-4 | 驱动线粒体氧化磷酸化 Drives mitochondrial oxidative phosphorylation | XR_003767947.1 | ||
细胞色素c氧化酶亚基5C-3 Cytochrome c oxidase subunit 5C-3 | 细胞色素c氧化酶的核编码多肽链之一,是线粒体电子传递的末端氧化酶 One of the nucleus-coding polypeptide chains of cytochrome C oxidase,the terminal oxidase of mitochondrial electron transport | XM_008370823.3 | ||
钾离子通道AKT2/3 Potassium channel AKT2/3 | 参与韧皮部汁液中输入和卸载钾的作用 Involved in both loading and unloading potassium into/from the phloem sap | XM_029107389.1 | ||
三酰甘油脂肪酶1 Triacylglycerol lipase 1 | 催化三酰基甘油的水解 Catalyzes the hydrolysis of triacylglycerol | XM_008340013.3 | ||
硫氧还原蛋白AAED1类似物 Thioredoxin-like protein AAED1 | 抗氧化活性 Antioxidant activity | XM_008343896.3 | ||
泛酸酯激酶 At2g17340-like | 催化泛酸的磷酸化 Catalyzes the phosphorylation of pantothenate | XM_008384280.3 | ||
非典型N-乙酰转移酶p20类似物 Uncharacterized N-acetyltransferase p20-like | 参与N-乙酰转移酶活性 N-acetyltransferase activity | XM_029096249.1 | ||
转录调控 Transcrip- tional regulation | 核糖核酸内切酶YBEY Endoribonuclease YBEY | 参与晚期70S核糖体质量控制和16S rRNA 3′端成熟 Involved in late-stage 70S ribosome quality control and in maturation of the 3′ terminus of the 16S rRNA | XM_029106325.1 | |
60S核糖体蛋白L10类似物 60S ribosomal protein L10-like | 在核糖体的形成中起积极作用 An active role in the formation of ribosomes | XM_029102547.1 | ||
抗逆防御 Resistance and defense | 脂肪酶PAD4类似物 Lipase-like PAD4 | 促进植物对入侵性生物营养病原体的先天免疫和识别微生物相关分子模式防御机制 Contributing to the plant innate immunity against invasive biotrophic pathogens and to defense mechanisms upon recognition of microbe-associated molecular patterns | XM_029094810.1 | |
应激反应蛋白NST1类似物 Stress response protein NST1-like | 与MSL1一起作为耐盐性的负调节因子 Negative regulator of salt tolerance along with MSL1 | XM_008394867.3 | ||
非特异性脂质转运蛋白3 Non-specific lipid-transfer protein 3 | 可以跨膜转移磷脂和半乳糖脂,参与植株对生物与非生物胁迫的抗性 Phospholipids and galactose lipids can be transferred across the membrane to participate in plant resistance to biological and abiotic stress | XM_008371792.3 | ||
解毒蛋白35类似物 Protein DETOXIFICATION 35-like | 参与类黄酮代谢的多药物和毒素外排转运体 Multidrug and toxin efflux transporters involved in flavonoid metabolism | XM_008381794.3 | ||
单链DNA结合蛋白WHY1 Single stranded DNA binding protein WHY1 | 致病相关基因PR-10a的转录激活因子 Transcriptional activator of the pathogenesis-related gene PR-10a | NM_001294042.1 | ||
脂质转移蛋白VAS类似物 Lipid transfer-like protein VAS | 可以跨膜转移磷脂和半乳糖脂,参与植株对生物与非生物胁迫的抗性 Phospholipids and galactose lipids can be transferred across the membrane to participate in plant resistance to biological and abiotic stress | XM_008342321.3 | ||
生长发育 Grow and develop- ment | MADS-box蛋白 SOC1类似物 MADS-box protein SOC1-like | 调控开花时间的转录因子 Transcription factors that regulate flowering time | XM_008378604.3 | |
质体转录活性蛋白7 Plastid transcriptionally active 7 | 参与质体基因表达,是类囊体形成所必需的 Involved in plastid gene expression,essential for thylakoid formation | XM_008382937.3 | ||
核酸外切酶DPD1 Exonuclease DPD1 | 花粉发育过程中细胞器DNA降解所需的外切酶 Exonuclease enzymes required for degradation of organelle DNA during pollen development | XM_017336537.2 | ||
光合作用NDH亚基的腔定位5 Photosynthetic NDH subunit of lumenal location 5 | 叶绿体NAD(P)H脱氢酶复合物形成和活性所必需 Essential for chloroplast NAD(P)H dehydrogenase complex formation and activity | XM_029091434.1 | ||
锌指蛋白CONSTANS-LIKE13类似物 Zinc finger protein CONSTANS-LIKE 13-like | 促进开花 Accelerate bloom | XM_029100386.1 | ||
二氨基庚二酸表异构酶 Diaminopimelate epimerase | 参与L-赖氨酸生物合成和肽聚糖中五肽的生物合成 Involved in the L-lysine biosynthesis and in the biosynthesis of the pentapeptide incorporated in the peptidoglycan moiety | XM_029101158.1 | ||
内质网素同源物 Endoplasmin homolog | 与芽、根和花分生组织形成及花粉管伸长有关 Related to shoot apical meristem,root apical meristem floral meristem formation and pollen tube elongation | XM_009336652.2 | ||
碳酸酐酶2类似物 Carbonic anhydrase 2-like | 催化二氧化碳的可逆水合作用形成碳酸氢盐 Catalyzes the reversible hydration of carbon dioxide to form bicarbonate | XM_008359520.3 | ||
叶绿体 Chloroplast | 从细胞质向叶绿体输入核编码的光合前蛋白 Nucleo-coded prephotosynthetic proteins are imported from the cytoplasm to chloroplasts | OK545754.1 | ||
LRR受体样丝氨酸/苏氨酸蛋白激酶 LRR receptor-like serine/threonine-protein kinase | 花粉发育的关键调控因子 Key regulator of anther development | XM_029088102.1 | ||
生育酚-O-甲基转移酶 Tocopherol O-methyltransferase | 参与生育酚的合成 Involved in the synthesis of tocopherol | XM_029092547.1 | ||
脂肪酸去饱和酶4 Fatty acid desaturase 4 | 参与生产含有叶绿体特异性的磷脂酰甘油 Involved in the production of chloroplast-specific phosphatidylglycerol | XM_008373042.2 |
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