籽粒苋C_4关键酶基因（NAD-ME、PEPC）的克隆及NAD-ME功能预测
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摘要
与C3植物相比,C4植物在极端条件(强光、高温、干旱等)下具有明显的生长优势、高养分利用率及高生物产量,这是由于其光合速率高、CO2补偿点低,而且几乎没有光呼吸。C4植物这些优点的生物学基础在于其具有C4光合途径。籽粒苋属双子叶C4植物,克隆其C4关键酶基因,以期为C4基因工程提供候选基因。
NAD／NADP-苹果酸酶和磷酸烯醇式丙酮酸羧化酶(PEPC)是C4途径的两个关键酶。NAD／NADP-苹果酸酶催化苹果酸脱羧释放CO2,并产生丙酮酸,丙酮酸再生为PEP,使C4途径得以循环进行。本研究根据已报道的玉米NADP-ME基因序列,设计特异性引物,经RT-PCR扩增获得了籽粒苋NAD-ME基因的cDNA序列。该序列开放可读框长度为1872bp,编码623个氨基酸。通过Blastn和DNAMAN软件比对,发现该基因核苷酸序列与其他植物已报道的NAD／NADP-ME基因有75.1%-80.6%的相似性。另外,对推断氨基酸序列的蛋白保守区、疏水性／亲水性、潜在跨膜片段、信号肽、蛋白固有无序化和蛋白二级结构进行了分析,初步研究了籽粒苋NAD-ME蛋白的理化性质。运用CHIPS、CUSP和CodonW程序分析了籽粒苋NAD-ME基因的密码子的偏好性,并与其他植物的NAD/NADP-ME基因密码子偏好性和大肠杆菌、酵母的基因组的密码子偏好性进行比较。结果表明,所比较的单子叶植物和双子叶植物的NAD/NADP-ME基因与籽粒苋NAD-ME基因都存在类似的密码子使用情况,对同义密码子的使用有明显的偏好性,其密码子多以A或T结尾；籽粒苋NAD-ME基因与大肠杆菌、酵母的基因组密码子偏好性有较大差异。分析结果对选择籽粒苋NAD-ME合适的转基因受体植物以及合适的蛋白质表达系统具有一定的指导意义。
PEPC在Mn2+或Mg2+存在的情况下,催化PEP羧化为OAA。本研究采用分段扩增的方法,首次克隆出了籽粒苋PEPC的基因组DNA全长。经Blast比对分析,结果显示,与本克隆片段相似的序列绝大多数为植物PEPC基因,其中,与F.pringlei的PEPC基因(Z71973.1)相似性高达93%。因此,初步判定实验获得的序列为籽粒苋基因组PEPC基因。
Compared with C3 plant, C4 plant has evident growth advantage, higher rate of nutrition using and higher bio-yield in extreme condition (such as high light intensities, high temperatures and drought) because of efficient photosynthesis and non-photorespiration. Cloning the key enzymes of C4 photosynthetic pathway in Amaranthus hypochondriacus L. as one of C4 dicotyledon provides candidate gene for C4 genetic engineering.
NAD/NADP-malic enzyme and phosphoenolpyruvate carboxylase (PEPC)are two key enzymes of C4 pathway. NAD/NADP-ME catalyzes the oxidative decarboxylation of 1-malate to yield pyruvate, CO2, and NADPH in the presence of a divalent cation. In C4 plants, NAD/NADP-ME release CO2 for fixation by Calvin cycle. In this study, a full-length cDNA encoding Amaranthus hypochondriacus L. NAD-ME was cloned by RT-PCR. The cDNA includes a 1872-bp open reading frame that encodes 623 amino acids. Blastn and DNAMAN analysis reveals that the nucleotide sequence has 75.1%-80.6% identity. Furthermore, the physico-chemical property of NAD-ME of Amaranthus hypochondriacus L.,including the conservative domain, hydrophobicity/hydrophilicity, potential transmembrane segment, signal peptide, intrinsically disordered structure and secondary structure of the hypothetical protein were analyzed. Codon bias of Amaranthus hypochondriacus L. NAD-ME was further analyzed by Codon W, CHIPS and CUSP programs for identifying codon bias. Moreover, the codon bias of Amaranthus hypochondriacus L.NAD-ME were compared with that of NAD/NADP-ME gene of other seven plants and two model organisms, Escherichia coli and yeast. The results showed that codon bias of NAD/NADP-ME of monocotyledons and dicotyledonous plants obviously remarked at some codons which were the A or T in the 3rd position of codons. Codon bias of NAD-ME of Amaranthus hypochondriacus L.was significantly different from those of E.coli and yeast genome.The result has important guidance for choosing appropriate recipient plants and protein exprssion system of NAD-ME gene of Amaranthus hypochondriacus L.
PEPC catalyzes irreversible carboxylation of PEP in the presence of Mn2+ and Mg2+ to yield OAA. In this study, we cloned the genomic gene of PEPC of Amaranthus hypochondriacus L. firstly by PCR. According to Blast analysis, we found most of the cloned DNA sequences similar to that of PEPC gene of other plants. And the identity to the F.pringlei PEPC was 93%.Thus, we determinated the cloned fragment was Amaranthus hypochondriacus L. PEPC preliminarily.
NAD／NADP-苹果酸酶和磷酸烯醇式丙酮酸羧化酶(PEPC)是C4途径的两个关键酶。NAD／NADP-苹果酸酶催化苹果酸脱羧释放CO2,并产生丙酮酸,丙酮酸再生为PEP,使C4途径得以循环进行。本研究根据已报道的玉米NADP-ME基因序列,设计特异性引物,经RT-PCR扩增获得了籽粒苋NAD-ME基因的cDNA序列。该序列开放可读框长度为1872bp,编码623个氨基酸。通过Blastn和DNAMAN软件比对,发现该基因核苷酸序列与其他植物已报道的NAD／NADP-ME基因有75.1%-80.6%的相似性。另外,对推断氨基酸序列的蛋白保守区、疏水性／亲水性、潜在跨膜片段、信号肽、蛋白固有无序化和蛋白二级结构进行了分析,初步研究了籽粒苋NAD-ME蛋白的理化性质。运用CHIPS、CUSP和CodonW程序分析了籽粒苋NAD-ME基因的密码子的偏好性,并与其他植物的NAD/NADP-ME基因密码子偏好性和大肠杆菌、酵母的基因组的密码子偏好性进行比较。结果表明,所比较的单子叶植物和双子叶植物的NAD/NADP-ME基因与籽粒苋NAD-ME基因都存在类似的密码子使用情况,对同义密码子的使用有明显的偏好性,其密码子多以A或T结尾；籽粒苋NAD-ME基因与大肠杆菌、酵母的基因组密码子偏好性有较大差异。分析结果对选择籽粒苋NAD-ME合适的转基因受体植物以及合适的蛋白质表达系统具有一定的指导意义。
PEPC在Mn2+或Mg2+存在的情况下,催化PEP羧化为OAA。本研究采用分段扩增的方法,首次克隆出了籽粒苋PEPC的基因组DNA全长。经Blast比对分析,结果显示,与本克隆片段相似的序列绝大多数为植物PEPC基因,其中,与F.pringlei的PEPC基因(Z71973.1)相似性高达93%。因此,初步判定实验获得的序列为籽粒苋基因组PEPC基因。
Compared with C3 plant, C4 plant has evident growth advantage, higher rate of nutrition using and higher bio-yield in extreme condition (such as high light intensities, high temperatures and drought) because of efficient photosynthesis and non-photorespiration. Cloning the key enzymes of C4 photosynthetic pathway in Amaranthus hypochondriacus L. as one of C4 dicotyledon provides candidate gene for C4 genetic engineering.
NAD/NADP-malic enzyme and phosphoenolpyruvate carboxylase (PEPC)are two key enzymes of C4 pathway. NAD/NADP-ME catalyzes the oxidative decarboxylation of 1-malate to yield pyruvate, CO2, and NADPH in the presence of a divalent cation. In C4 plants, NAD/NADP-ME release CO2 for fixation by Calvin cycle. In this study, a full-length cDNA encoding Amaranthus hypochondriacus L. NAD-ME was cloned by RT-PCR. The cDNA includes a 1872-bp open reading frame that encodes 623 amino acids. Blastn and DNAMAN analysis reveals that the nucleotide sequence has 75.1%-80.6% identity. Furthermore, the physico-chemical property of NAD-ME of Amaranthus hypochondriacus L.,including the conservative domain, hydrophobicity/hydrophilicity, potential transmembrane segment, signal peptide, intrinsically disordered structure and secondary structure of the hypothetical protein were analyzed. Codon bias of Amaranthus hypochondriacus L. NAD-ME was further analyzed by Codon W, CHIPS and CUSP programs for identifying codon bias. Moreover, the codon bias of Amaranthus hypochondriacus L.NAD-ME were compared with that of NAD/NADP-ME gene of other seven plants and two model organisms, Escherichia coli and yeast. The results showed that codon bias of NAD/NADP-ME of monocotyledons and dicotyledonous plants obviously remarked at some codons which were the A or T in the 3rd position of codons. Codon bias of NAD-ME of Amaranthus hypochondriacus L.was significantly different from those of E.coli and yeast genome.The result has important guidance for choosing appropriate recipient plants and protein exprssion system of NAD-ME gene of Amaranthus hypochondriacus L.
PEPC catalyzes irreversible carboxylation of PEP in the presence of Mn2+ and Mg2+ to yield OAA. In this study, we cloned the genomic gene of PEPC of Amaranthus hypochondriacus L. firstly by PCR. According to Blast analysis, we found most of the cloned DNA sequences similar to that of PEPC gene of other plants. And the identity to the F.pringlei PEPC was 93%.Thus, we determinated the cloned fragment was Amaranthus hypochondriacus L. PEPC preliminarily.
引文
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[3]Kanai R, Edwards G E. The biochemistry of C4 photosynthesis. C4 Plant Biology, 1999,49-87.
[4]Edwrads G E, Andreo C S.NADP-malic enzyme from plants.Phytochemistry,1992, 31,1845-1857.
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[6]Palmer J M. The organization and regulation of electron transport in plant mitochondria. AnnuRev Plant Physiol,1976,27,133-157.
[7]Rustin P, Moreau F, Lance C. Malate oxidation in plant mitochondria via malic enzyme and the cyanide-insensitive electron transport pathway. Plant Physiology, 1980,66,457-462.
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[9]Mariel C, Marcos A. A comprehensive analysis of the NADP malic enzyme gene family of Arabidopsis. Plant Phy,2005,139(1),39-51.
[10]迟伟,阳江华,张青婵.水稻的4个NADP-ME基因具有不同的表达模式.自然科学进展,2004,14(12),1402-1410.
[11]Marion H. O'Leary. Phosphoenolpy ruvatecarboxylase:an enzymologist's view. Annu Rev Plant Physiol,1982,33,297-315.
[12]Raymond C, Jean V, Marion H. PHOSPHOENOLPYRUVATE CARBOXYLASE:A Ubiquitous, Highly Regulated Enzyme in Plants. Annu Rev Plant Physiol Plant Mol Biol,1996,47,273-298.
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