玉米C_4途径关键酶基因（PPDK、NADP-ME）的克隆及PPDK、PEPC在拟南芥中的表达分析
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摘要
光合作用是植物生物学产量的源泉,通过基因工程手段人为加强“C4特性”在C3遗传背景下的发挥,实现C3植物光合途径的C4化,对于提高作物光能利用率和产量具有重要意义。本试验从酶活和光合均高的玉米自交系中克隆了玉米C4型丙酮酸磷酸二激酶(PPDK)和依赖于NADP的苹果酸酶(NADP-ME)基因,并利用PPDK和本实验室先前克隆的PEPC基因(GeneBank:FJ415327)转化拟南芥,研究了这两个基因的单独及协同作用,通过其生理生化方面的变化,为C3作物光合效率的改善提供了候选基因,为C4途径关键酶导入小麦等C3作物提供了依据。
主要结论如下：
1.利用同源克隆技术,从玉米中克隆了PPDK(GenBank:GU363532)和NADP-ME基因的cDNA,并插入双元表达载体pCAMBIA3301.pCAMBIA1391中,构建了p3301-PPDK、 p3301-ME和p1391-ME融合表达载体。序列测定结果表明,PPDK基因长度为3004Bp,包含一个起始密码子和971个氨基酸的开放阅读框,与己登陆的玉米PPDK基因(GeneBank: BT054438)相比,氨基酸序列同源性可达99.4%；NADP-ME基因长度为2127Bp,包含一个起始密码子和636个氨基酸的开放阅读框,与已登陆的玉米NADP-ME基因(GenBank: NM_001111843.1)相比,氨基酸序列同源性可达99.6%。
2.Southern blot表明玉米PPDK、PEPC基因已整合到拟南芥基因组中,拟南芥转化率为0.69%,双基因共转化率为0.07%,66.7%的双基因拟南芥符合Mendel单基因的遗传规律。利用农杆菌介导法将p3301-PPDK和本实验室曾构建的p3301-PEPC高效表达载体导入拟南芥中,筛选到327株抗性转基因拟南芥,其中PEPC株系125株,PPDK株系175株,PPDK、 PEPC共表达株系12株,转p3301株系15株,筛选到PPDK、PEPC、PPDK+PEPC、p330.纯系各24、26、8、3个。
3.RT-PCR、荧光定量PCR、Western blot分析表明外源基因在拟南芥基因组中得到正确表达,且转PPDK、PEPC基因植株的表达量存在较大差异。荧光定量PCR分析现蕾期PPDK、 PEPC基因的相对表达量分别在1.0-4.2倍和1.0-13.3倍；用Quantity One软件对Western blot进行定量分析表明,PPDK、PEPC基因的蛋白相对表达量分别在1.9-2.5倍和1.6-3.7倍。
4.转单、双C4关键酶基因均可以提高拟南芥光合速率,双基因的平均光合速率高于转PEPC基因的平均光合速率,转PEPC基因的平均光合速率高于转PPDK基因的平均光合速率。T3代转基因拟南芥酶活和光合速率测定表明：T3代转基因拟南芥PPDK、PEPC酶活分别比对照增加0.61-3.63倍和0.77-4.81倍,平均增加1.89和1.48倍；转PPDK基因拟南芥光合速率增加3.3-25.3%,平均增加12.2%,转PEPC基因拟南芥光合速率增加6.5-35.1%,平均增加18.9%,双基因(PPDK+PEPC)拟南芥光合速率增加12.6-45.7%,平均增加26.3%。
5.转单个C4关键酶基因加速了C4微循环。T4代转基因拟南芥酶活测定表明：PK17-1-2、 PK26-3-2的PEPC酶活分别比对照增加18.65%和46.37%,与对照差异极显显著；PC65-4-6、PC73-1-3的PPDK酶活分别比对照增加12.96%和26.54%,与对照差异极显著,说明转PPDK基因拟南芥可引起PEPC活性增加,反之亦然；T4代转基因拟南芥光合测定表明：PK17-1-2、 PK26-3-2、PC65-4-6、PC73-1-3的光合速率分别比对照增加了13.14%、23.29%、18.14%、36.44%,达极显著水平。
6.PPDK、PEPC共表达有协同效应。双基因株系PKC6-5-1的光合速率比PK26-3-2和PC73-1-3的高,但是PKC6-5-1的PEPC酶活比PC73-1-3的低,PKC6-5-1的PPDK酶活比PK26-3-2的低；双基因株系PKC17-1-3的光合速率比PK17-1-2和PC65-4-6的高,但PKC6-5-1的PPDK和PEPC酶活偏低。
ABSTRACT:The efficiency of photosynthesis is related with the crop biomass, since the harvest index and leaf area index for many crops, such as wheat and rice, is approaching a ceiling value, it has been a focus that an increase in yield potential should come from improved photosynthesis. According to differences of mechanism in carbon assimilation pathways in photosynthesis, three major photosynthetic types may be distinguished among green plants:C3, C4and Crassulacean acid metabolism plants. C4plants were evolved from C3plants, they adapt to high light, high tempretaure, lower CO2concentration and achieve higher photosynthetic capacity compared with C3plants(Black,1973). Consequently, the transfer of C4enzyme to C3plants such as rice and wheat is one strategy being adopted for improving their yield. In this experiment, PPDK with an ORF of971amino acids was isolated from maize self lines Z1194and was introduced to Arabidopsis with PEPC, systematical studies on their transcription, translation, eneyzme activities and photosynthetic performences were conducted, which could offer candidate gene and theory for engineering the C4photosynthetic pathway into C3crops such as wheat.
The results were as follows:
1. The full-length cDNA for PPDK and NADP-ME were isolated from Zea mays by homology-based cloning technology. The full-length PPDK and NADP-ME cDNA replaced the (3-glucuronidse(GUS) coding region of the pCAMBIA3301(p3301) or pCAMBIA1391(p1391). The positive clone was named p3301-PPDK, p3301-ME and p1391-ME, respectively. In this study, we have obtained a3004bp cDNA fragment of PPDK from maize Z1194(Zea mays L.), PPDK has a2916bp open reading frame (ORF) and encodes971amino acids with a calculated molecular mass of105kDa polypeptide(GenBank accession number:GU363532). Comparison of GU363532to the C4PPDK gene (GenBank accession number:BT054438) by DNAMAN indicated99.4%similarity between the amino acid sequences. We have also obtained a2127bp cDNA fragment of NADP-ME from maize Zl194, containing one ORF with1911bp which coded a peotide of636amino acids. The identity of the cloned fragment with NADP-ME gene from NM_001111843.1was99.4%by homology alignment with software DNAMAN.
2. The integration of maize PPDK and PEPC gene in transgenic Arabidopsis plants was confirmed by Southern blotting analysis, the transgentic rate and co-transgentic rate was0.69%and9.5%, the ratio of linked-inheritance was66.7%. Arabidopsis plants were transformed by the floral dipping method with tumefaciens GV3101, including p3301-PPDK, p3301-PEPC, p3301-PPDK+p3301-PEPC(PPDK:PEPC was3:2), and p3301. Based on the segregation pattern of the transgenic seedlings by Basta-tolerant and PCR assay. A total of327transgentic plants were obtained, plants containing the PPDK gene, the PEPC gene, the PPDK+PEPC gene, or GUS gene were separated into their respective groups, indicating the following distribution of lines:twenty-four homogenous PPDK lines from175PPDK transgentic lines, twenty-six homogenous PEPC lines from125PPDK transgentic lines, eight homogenous lines linked inheritance with both PPDK and PEPC from12double transgentic lines, and three homogenous p3301lines from15transgentic lines of p3301.
3. Analyses by RT-PCR, quantitative real-time PCR and Western blotting all confirmed the effective expression of maize C4-PPDK and C4-PEPC in Arabidopsis. The expression levels of PPDK or PEPC among different transgenic lines varied, which ranged from1-to4.24-fold and1-to13.33-fold by quantitative real-time PCR, respectively. Volume analysis by software Quantity One indicated that the expression levels of PPDK were ranged from1.9-to2.5-fold relevant to the NegCtrl, the expression levels of PEPC were ranged from1.6-to3.7-fold relevant to the NegCtrl, respectively.
4. The photosynthetic performance could be improved by PPDK or PEPC alone, and what's more, Pn (PKC)> Pn (PC)> Pn(PK). In this study, the enzyme activities and the Pn of T3transgentic plants were determined.The enzyme activity of C4-PPDK and C4-PEPC were elevated by0.61-3.63and0.77-4.81times relevant to the controls, the average value was1.89and1.48folds, respectively. The Pn of Ca-PPDK, C4-PEPC and PPDK+PEPC transgenic lines were higher than those of the controls by3.3-25.3%,6.5-35.1%and12.6-45.7%.
5. Tthe introduction of C4-PPDK or C4-PEPC into Arabidopsis could contribute to the strengthen of C4-mini cycle. In this study, the enzyme activities of T4transgentic Arabidopsis were determined. The PEPC enzyme activities of PK17-1-2and PK26-3-2were18.65%and46.37%higher than those of the controls, respectively; the PPDK enzyme activities of PC65-4-6and PC73-1-3were12.96%and26.54%higher than those of the controls, respectively, the PPDK and PEPC enzyme activities could significantly increased by transgentic PPDK and PEPC Arabidopsis. The Pn of PK17-1-2, PK26-3-2, PC65-4-6and PC73-1-3were13.14%,23.29%,18.14%and36.44%higher than those of the controls, respectively.
6. The coexpression of PPDK and PEPC had synergistic effect on photosynthesis rate. To our interest, the Pn of the line PKC6-5-1were the highest among the T4transgentic Arabidopsis, however, its PPDK enzyme activities or its PEPC enzyme activities were not the highest, although the PPDK or PEPC eneyzme activities of the line PKC17-1-3was the lowest among the T4transgentic Arabidopsis, the Pn of the line PKC17-1-3were higher than those of the line PK17-1-2and PC65-4-6.
The results show that both are candidate functional genes in the improvement of Pn of C3Arabidopsis and coexpression of C4-PPDK and C4-PEPC genes had synergistic effects on Pn, which could be of great importance in the agricultural field, especially when considering the limited methods that we currently have in improving crop yields.
主要结论如下：
1.利用同源克隆技术,从玉米中克隆了PPDK(GenBank:GU363532)和NADP-ME基因的cDNA,并插入双元表达载体pCAMBIA3301.pCAMBIA1391中,构建了p3301-PPDK、 p3301-ME和p1391-ME融合表达载体。序列测定结果表明,PPDK基因长度为3004Bp,包含一个起始密码子和971个氨基酸的开放阅读框,与己登陆的玉米PPDK基因(GeneBank: BT054438)相比,氨基酸序列同源性可达99.4%；NADP-ME基因长度为2127Bp,包含一个起始密码子和636个氨基酸的开放阅读框,与已登陆的玉米NADP-ME基因(GenBank: NM_001111843.1)相比,氨基酸序列同源性可达99.6%。
2.Southern blot表明玉米PPDK、PEPC基因已整合到拟南芥基因组中,拟南芥转化率为0.69%,双基因共转化率为0.07%,66.7%的双基因拟南芥符合Mendel单基因的遗传规律。利用农杆菌介导法将p3301-PPDK和本实验室曾构建的p3301-PEPC高效表达载体导入拟南芥中,筛选到327株抗性转基因拟南芥,其中PEPC株系125株,PPDK株系175株,PPDK、 PEPC共表达株系12株,转p3301株系15株,筛选到PPDK、PEPC、PPDK+PEPC、p330.纯系各24、26、8、3个。
3.RT-PCR、荧光定量PCR、Western blot分析表明外源基因在拟南芥基因组中得到正确表达,且转PPDK、PEPC基因植株的表达量存在较大差异。荧光定量PCR分析现蕾期PPDK、 PEPC基因的相对表达量分别在1.0-4.2倍和1.0-13.3倍；用Quantity One软件对Western blot进行定量分析表明,PPDK、PEPC基因的蛋白相对表达量分别在1.9-2.5倍和1.6-3.7倍。
4.转单、双C4关键酶基因均可以提高拟南芥光合速率,双基因的平均光合速率高于转PEPC基因的平均光合速率,转PEPC基因的平均光合速率高于转PPDK基因的平均光合速率。T3代转基因拟南芥酶活和光合速率测定表明：T3代转基因拟南芥PPDK、PEPC酶活分别比对照增加0.61-3.63倍和0.77-4.81倍,平均增加1.89和1.48倍；转PPDK基因拟南芥光合速率增加3.3-25.3%,平均增加12.2%,转PEPC基因拟南芥光合速率增加6.5-35.1%,平均增加18.9%,双基因(PPDK+PEPC)拟南芥光合速率增加12.6-45.7%,平均增加26.3%。
5.转单个C4关键酶基因加速了C4微循环。T4代转基因拟南芥酶活测定表明：PK17-1-2、 PK26-3-2的PEPC酶活分别比对照增加18.65%和46.37%,与对照差异极显显著；PC65-4-6、PC73-1-3的PPDK酶活分别比对照增加12.96%和26.54%,与对照差异极显著,说明转PPDK基因拟南芥可引起PEPC活性增加,反之亦然；T4代转基因拟南芥光合测定表明：PK17-1-2、 PK26-3-2、PC65-4-6、PC73-1-3的光合速率分别比对照增加了13.14%、23.29%、18.14%、36.44%,达极显著水平。
6.PPDK、PEPC共表达有协同效应。双基因株系PKC6-5-1的光合速率比PK26-3-2和PC73-1-3的高,但是PKC6-5-1的PEPC酶活比PC73-1-3的低,PKC6-5-1的PPDK酶活比PK26-3-2的低；双基因株系PKC17-1-3的光合速率比PK17-1-2和PC65-4-6的高,但PKC6-5-1的PPDK和PEPC酶活偏低。
ABSTRACT:The efficiency of photosynthesis is related with the crop biomass, since the harvest index and leaf area index for many crops, such as wheat and rice, is approaching a ceiling value, it has been a focus that an increase in yield potential should come from improved photosynthesis. According to differences of mechanism in carbon assimilation pathways in photosynthesis, three major photosynthetic types may be distinguished among green plants:C3, C4and Crassulacean acid metabolism plants. C4plants were evolved from C3plants, they adapt to high light, high tempretaure, lower CO2concentration and achieve higher photosynthetic capacity compared with C3plants(Black,1973). Consequently, the transfer of C4enzyme to C3plants such as rice and wheat is one strategy being adopted for improving their yield. In this experiment, PPDK with an ORF of971amino acids was isolated from maize self lines Z1194and was introduced to Arabidopsis with PEPC, systematical studies on their transcription, translation, eneyzme activities and photosynthetic performences were conducted, which could offer candidate gene and theory for engineering the C4photosynthetic pathway into C3crops such as wheat.
The results were as follows:
1. The full-length cDNA for PPDK and NADP-ME were isolated from Zea mays by homology-based cloning technology. The full-length PPDK and NADP-ME cDNA replaced the (3-glucuronidse(GUS) coding region of the pCAMBIA3301(p3301) or pCAMBIA1391(p1391). The positive clone was named p3301-PPDK, p3301-ME and p1391-ME, respectively. In this study, we have obtained a3004bp cDNA fragment of PPDK from maize Z1194(Zea mays L.), PPDK has a2916bp open reading frame (ORF) and encodes971amino acids with a calculated molecular mass of105kDa polypeptide(GenBank accession number:GU363532). Comparison of GU363532to the C4PPDK gene (GenBank accession number:BT054438) by DNAMAN indicated99.4%similarity between the amino acid sequences. We have also obtained a2127bp cDNA fragment of NADP-ME from maize Zl194, containing one ORF with1911bp which coded a peotide of636amino acids. The identity of the cloned fragment with NADP-ME gene from NM_001111843.1was99.4%by homology alignment with software DNAMAN.
2. The integration of maize PPDK and PEPC gene in transgenic Arabidopsis plants was confirmed by Southern blotting analysis, the transgentic rate and co-transgentic rate was0.69%and9.5%, the ratio of linked-inheritance was66.7%. Arabidopsis plants were transformed by the floral dipping method with tumefaciens GV3101, including p3301-PPDK, p3301-PEPC, p3301-PPDK+p3301-PEPC(PPDK:PEPC was3:2), and p3301. Based on the segregation pattern of the transgenic seedlings by Basta-tolerant and PCR assay. A total of327transgentic plants were obtained, plants containing the PPDK gene, the PEPC gene, the PPDK+PEPC gene, or GUS gene were separated into their respective groups, indicating the following distribution of lines:twenty-four homogenous PPDK lines from175PPDK transgentic lines, twenty-six homogenous PEPC lines from125PPDK transgentic lines, eight homogenous lines linked inheritance with both PPDK and PEPC from12double transgentic lines, and three homogenous p3301lines from15transgentic lines of p3301.
3. Analyses by RT-PCR, quantitative real-time PCR and Western blotting all confirmed the effective expression of maize C4-PPDK and C4-PEPC in Arabidopsis. The expression levels of PPDK or PEPC among different transgenic lines varied, which ranged from1-to4.24-fold and1-to13.33-fold by quantitative real-time PCR, respectively. Volume analysis by software Quantity One indicated that the expression levels of PPDK were ranged from1.9-to2.5-fold relevant to the NegCtrl, the expression levels of PEPC were ranged from1.6-to3.7-fold relevant to the NegCtrl, respectively.
4. The photosynthetic performance could be improved by PPDK or PEPC alone, and what's more, Pn (PKC)> Pn (PC)> Pn(PK). In this study, the enzyme activities and the Pn of T3transgentic plants were determined.The enzyme activity of C4-PPDK and C4-PEPC were elevated by0.61-3.63and0.77-4.81times relevant to the controls, the average value was1.89and1.48folds, respectively. The Pn of Ca-PPDK, C4-PEPC and PPDK+PEPC transgenic lines were higher than those of the controls by3.3-25.3%,6.5-35.1%and12.6-45.7%.
5. Tthe introduction of C4-PPDK or C4-PEPC into Arabidopsis could contribute to the strengthen of C4-mini cycle. In this study, the enzyme activities of T4transgentic Arabidopsis were determined. The PEPC enzyme activities of PK17-1-2and PK26-3-2were18.65%and46.37%higher than those of the controls, respectively; the PPDK enzyme activities of PC65-4-6and PC73-1-3were12.96%and26.54%higher than those of the controls, respectively, the PPDK and PEPC enzyme activities could significantly increased by transgentic PPDK and PEPC Arabidopsis. The Pn of PK17-1-2, PK26-3-2, PC65-4-6and PC73-1-3were13.14%,23.29%,18.14%and36.44%higher than those of the controls, respectively.
6. The coexpression of PPDK and PEPC had synergistic effect on photosynthesis rate. To our interest, the Pn of the line PKC6-5-1were the highest among the T4transgentic Arabidopsis, however, its PPDK enzyme activities or its PEPC enzyme activities were not the highest, although the PPDK or PEPC eneyzme activities of the line PKC17-1-3was the lowest among the T4transgentic Arabidopsis, the Pn of the line PKC17-1-3were higher than those of the line PK17-1-2and PC65-4-6.
The results show that both are candidate functional genes in the improvement of Pn of C3Arabidopsis and coexpression of C4-PPDK and C4-PEPC genes had synergistic effects on Pn, which could be of great importance in the agricultural field, especially when considering the limited methods that we currently have in improving crop yields.
引文
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