猕猴桃AsA合成相关酶基因及启动子的克隆与功能分析
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摘要
抗坏血酸（AsA）是动植物体内重要的氧化还原剂，参与体内活性氧清除，调控细胞分裂和细胞生长，并作为多个酶的辅因子参与多种生理过程。此外，AsA为人类必需的营养物质，能增强人体相关免疫系统的活性，在由氧胁迫引起的相关疾病的防御方面，发挥着相当重要的作用。AsA还是植物生长调控的必需因子，它能控制植物的开花时间，还能够增强植物体对逆境的抵御能力，比如臭氧毒害、水涝胁迫、高温伤害和病原菌入侵等。
目前，推测AsA合成与积累的关键调控酶：包括GDP-D-甘露糖3’,5’-差向异构酶（GME）、L半乳糖-1-磷酸磷酸酶（GPP）和GDP-L-半乳糖磷酸化酶（GGP）。本研究以山梨猕猴桃（Actinidia rufa）、美味猕猴桃品种“秦美”（A. deliciosa cv.‘qinmei’）和毛花猕猴桃（A. eriantha）为材料，克隆了AsA合成相关酶基因GME、GPP和GGP，并利用实时荧光定量PCR分别分析了这些基因在不同处理条件下的表达模式；克隆了GME、GPP和GGP基因启动子序列并在不同处理下对其进行了活性分析；同时测定了不同处理条件下猕猴桃叶片中AsA含量变化。此外，还研究了猕猴桃果实发育期GGP基因表达水平的变化及其与AsA含量的关系。获得的主要结果如下：
1.从山梨猕猴桃和美味猕猴桃中克隆了GME基因全长，包含1143bp，编码376个氨基酸，把这两个cDNA分别命名为ArGME和AdGME，并登录Genebank，登陆号分别为JN132110和GU339037，二者序列相似性为99.7%。RT-qRCR结果表明：ABA、黑暗和光照处理对ArGME和AdGME基因在叶片中的表达量没有明显影响；GA3处理降低了ArGME和AdGME的表达；SA处理、机械损伤、高温和低温处理，均不同程度的增加了ArGME和AdGME的表达。相同处理对两个猕猴桃种的作用趋势基本一致。
2.利用染色体步移法克隆了山梨猕猴桃和美味猕猴桃GME基因启动子的序列，其长度均为1225-bp，命名为ParGME和PadGME，并登录Genebank，登陆号分别为JQ693500和JQ693499，二者序列相似性为99.16%。瞬时表达实验证明，这两个启动子具有转录活性。光照、机械损伤对GME基因的启动子没有明显的响应，而水杨酸和高温对GME基因启动子的活性却有很强的诱导和激活作用。
3.从美味猕猴桃中克隆了GPP基因并对其表达模式进行了分析，结果发现：持续的黑暗处理降低了GPP基因的表达，紧随其后的光照处理诱导了GPP基因的表达；SA对GPP基因的表达影响甚微；ABA、机械损伤和低氧胁迫能先增加GPP基因的表达而后降低其表达。
4.利用染色体步移法克隆美味猕猴桃GPP基因启动子，长度为1395-bp，瞬时表达实验证明，其具有转录活性。GPP基因启动子5’端缺失体实验表明，光照能够明显诱导GPP基因启动子的活性，重要的光调控元件可能位于-1210到-1047这个区段内。另外，ABA、SA、机械损伤和淹水等处理也能诱导GPP基因启动子的活性。
5.从山梨猕猴桃和毛花猕猴桃中克隆了GGP基因全长cDNA序列，命名为AeGGP(登录号：KC146049)和ArGGP(登录号：KC146048)。猕猴桃果实发育过程中，GGP基因表达与AsA含量变化一致，且毛花猕猴桃最高，山梨猕猴桃最低。GGP基因的表达对光很敏感；MeJA、高温和低氧能诱导GGP基因的表达，ABA降低了其表达。
6.利用染色体步移法克隆了山梨猕猴桃、美味猕猴桃和毛花猕猴桃GGP基因启动子的序列，山梨猕猴桃GGP基因启动子的活性最低，在其序列中发现了一段300bp的插入，启动子缺失体实验研究表明，在山梨猕猴桃GGP基因的启动子中可能存在负调控元件。光、ABA、MeJA、高温和低氧均能诱导GGP基因启动子的活性。根据毛花猕猴桃和美味猕猴桃GGP基因启动子缺失体对不同处理的响应情况，推测G-box motif可能在光暗响应中发挥着极为重要的作用；ABRE在ABA对GGP基因启动子的诱导中发挥了重要作用；CGTCA-motif元件在MeJA对GGP基因启动子的诱导中发挥了重要作用。
Ascorbic acid (AsA) is an important antioxidant in animals and plants, which playsimportant roles in eliminating reactive oxygen species (ROS) while allowing for crucialsignaling to regulate plant growth and defenses and an essential enzyme cofactor in manymetabolism processes. AsA is clearly an essential dietary component for humans, with aprotective role proposed for many disorders through increasing resistance of immune system.Ascorbic acid (AsA) is a vital compound in plants with a range of functions as a growthregulation factor in regulating flowering time, or as a stress response factor, which is involvedin many stress-induced oxidative processes, e.g., responses to water loss, pathogens andoxidizing agents, high temperature, photo-oxidative stress and ozone. Up to now,GDP-mannose-3',5'-epimerase (GME), GDP-L-galactose phosphotylase (GGP), L-galactosephosphatase (GPP) may play key roles in ASA biosynthesis and regulation. In present study,gene cloning, real-time PCR, and promoter characteristics were used to illustrate themolecular characteristics of enzymes involved in ASA biosynthesis under different treatments.To gain a better understanding of the relationship between AsA concentrations andtranscriptional levels of the above genes, and to investigate the regulatory mechanisms fortheir expression in kiwifruit under stress conditions, we performed a systematic investigationof AsA, mRNA expression, and promoter activities in response to various treatments. For this,we evaluated three kiwifruit species (A. eriantha, A. deliciosa cv.‘qinmei’ and A.rufa) thatproduce different amounts of AsA. Furthermore, to evaluate the relationship between levels ofAsA in fruits development and GGP transcripts, we systematically investigated the regulatorymechanisms for GGP expression in the three kiwifruit species. The main results were asfollows:
1. The full-length cDNA sequence of GME gene was cloned from A. deliciosa‘qinmei’and A.rufa, named AdGME (GenBank Accession NO. GU339037) and ArGME (GenBankAccession NO. JN132110). They were both1143-bp long, both encoded a polypeptide of376amino acids. Using the MegAlign program of MEGA, we determined that these amino acidsequences shared99.7%identity. RT-qPCR analyses in two species demonstrated that ABA, dark or light did not seem to alter GME expression, but GA3resulted in a decrease in thetranscript levels of GME. However, application of SA, wounding, cold or heat, GMEtranscript exhibited an increase in both species.
2. The promoters of GME genes were cloned by Genomic DNA-walking method. Theywere both1225-bp long, named ParGME和PadGME. We comparatively analysed thepromoter sequences of ParGME (GenBank Accession NO. JQ693500) and PadGME(GenBank Accession NO. JQ693499). Sequence alignment revealed that they shared99.16%similarity. They showed transcriptional activation identified by transformed tobacco GUShistochemical assay. The promoter activities of GME genes were not significantly inducedwhen the transformed tobacco exposing to light or wounding. While the promoter activities ofGME genes were significantly induced when treated by SA or heat.
3. The full-length cDNA sequence of AdGPP gene was cloned from A. deliciosa. Theexpression patterns of AdGPP gene were studied when kiwifruit plants were exposing todifferent conditions. RT-qRCR analyses demonstrated: during the first12h of darkness,AdGPP expression gradually but continually declined. After48h of dark treatment, plantswere then exposed to light, and AdGPP expression quickly recovered within1h. Following apeak at Hour2, then AdGPP expression began to decrease to the original level. Treatmentwith SA caused only a slight increase in expression over the entire period. ABA or hypoxicconditions were associated with high AdGPP expression at first, followed by a gradualdecline in transcript levels.
4. The AdGPP promoter was isolated from A. deliciosa genomic DNA by GenomicDNA-walking method. After sequencing, we cloned a1395-bp fragment (GenBank AccessionNo. JX122767). The AdGPP promoter showed transcriptional activation identified bytransformed tobacco GUS histochemical assay. The promoter activities of AdGPP were notsignificantly induced when the transformed tobacco exposing to light or wounding. While thepromoter activities of AdGPP gene were significantly induced when treated by SA or heat.
5. The full-length cDNA sequence of GGP genes were cloned from A.eriantha andA.rufa, named AeGGP (GenBank Accession No. KC146049) and ArGGP (GenBankAccession No. KC146048). Some correlation between relative levels of GGP mRNA andAsA concentrations in three species during fruit development. Transcripts were the mostabundant in A. eriantha, and the lowest in A. rufa. RT-qPCR results with the kiwifruit leavesrevealed that expression of the GGP genes was induced by light, MeJA, heat, hypoxicconditions; and was very sensitive to light conditions.
6. The promoters of GGP genes were cloned from three kiwifruit species by GenomicDNA-walking method. GUS activity of the entire AeGGP promoter was the highest and that of ArGGP promoter was the lowest under normal growing conditions. When compared thethree promoters, we found that ArGGP promoter has a nearly300-bp insertion. The result of5`deletion derivates transformed into tobacco indicates that some negative cis-elements mayexist in the ArGGP promoter. The promoter activities of GGP genes were significantlyinduced when application of ABA or MeJA. Light, heat or hypoxic conditions can also inducethe promoter activities of GGP genes. According to the results of5`deletion derivates ofAeGGP or AdGGP promoter, we deduced that G-box is important for the expression of GGPin kiwifruit under different types of illumination and ABRE or CGTCA-motif may play animportant role in regulating GGP promoter activities after application of ABA or MeJArespectively.
目前，推测AsA合成与积累的关键调控酶：包括GDP-D-甘露糖3’,5’-差向异构酶（GME）、L半乳糖-1-磷酸磷酸酶（GPP）和GDP-L-半乳糖磷酸化酶（GGP）。本研究以山梨猕猴桃（Actinidia rufa）、美味猕猴桃品种“秦美”（A. deliciosa cv.‘qinmei’）和毛花猕猴桃（A. eriantha）为材料，克隆了AsA合成相关酶基因GME、GPP和GGP，并利用实时荧光定量PCR分别分析了这些基因在不同处理条件下的表达模式；克隆了GME、GPP和GGP基因启动子序列并在不同处理下对其进行了活性分析；同时测定了不同处理条件下猕猴桃叶片中AsA含量变化。此外，还研究了猕猴桃果实发育期GGP基因表达水平的变化及其与AsA含量的关系。获得的主要结果如下：
1.从山梨猕猴桃和美味猕猴桃中克隆了GME基因全长，包含1143bp，编码376个氨基酸，把这两个cDNA分别命名为ArGME和AdGME，并登录Genebank，登陆号分别为JN132110和GU339037，二者序列相似性为99.7%。RT-qRCR结果表明：ABA、黑暗和光照处理对ArGME和AdGME基因在叶片中的表达量没有明显影响；GA3处理降低了ArGME和AdGME的表达；SA处理、机械损伤、高温和低温处理，均不同程度的增加了ArGME和AdGME的表达。相同处理对两个猕猴桃种的作用趋势基本一致。
2.利用染色体步移法克隆了山梨猕猴桃和美味猕猴桃GME基因启动子的序列，其长度均为1225-bp，命名为ParGME和PadGME，并登录Genebank，登陆号分别为JQ693500和JQ693499，二者序列相似性为99.16%。瞬时表达实验证明，这两个启动子具有转录活性。光照、机械损伤对GME基因的启动子没有明显的响应，而水杨酸和高温对GME基因启动子的活性却有很强的诱导和激活作用。
3.从美味猕猴桃中克隆了GPP基因并对其表达模式进行了分析，结果发现：持续的黑暗处理降低了GPP基因的表达，紧随其后的光照处理诱导了GPP基因的表达；SA对GPP基因的表达影响甚微；ABA、机械损伤和低氧胁迫能先增加GPP基因的表达而后降低其表达。
4.利用染色体步移法克隆美味猕猴桃GPP基因启动子，长度为1395-bp，瞬时表达实验证明，其具有转录活性。GPP基因启动子5’端缺失体实验表明，光照能够明显诱导GPP基因启动子的活性，重要的光调控元件可能位于-1210到-1047这个区段内。另外，ABA、SA、机械损伤和淹水等处理也能诱导GPP基因启动子的活性。
5.从山梨猕猴桃和毛花猕猴桃中克隆了GGP基因全长cDNA序列，命名为AeGGP(登录号：KC146049)和ArGGP(登录号：KC146048)。猕猴桃果实发育过程中，GGP基因表达与AsA含量变化一致，且毛花猕猴桃最高，山梨猕猴桃最低。GGP基因的表达对光很敏感；MeJA、高温和低氧能诱导GGP基因的表达，ABA降低了其表达。
6.利用染色体步移法克隆了山梨猕猴桃、美味猕猴桃和毛花猕猴桃GGP基因启动子的序列，山梨猕猴桃GGP基因启动子的活性最低，在其序列中发现了一段300bp的插入，启动子缺失体实验研究表明，在山梨猕猴桃GGP基因的启动子中可能存在负调控元件。光、ABA、MeJA、高温和低氧均能诱导GGP基因启动子的活性。根据毛花猕猴桃和美味猕猴桃GGP基因启动子缺失体对不同处理的响应情况，推测G-box motif可能在光暗响应中发挥着极为重要的作用；ABRE在ABA对GGP基因启动子的诱导中发挥了重要作用；CGTCA-motif元件在MeJA对GGP基因启动子的诱导中发挥了重要作用。
Ascorbic acid (AsA) is an important antioxidant in animals and plants, which playsimportant roles in eliminating reactive oxygen species (ROS) while allowing for crucialsignaling to regulate plant growth and defenses and an essential enzyme cofactor in manymetabolism processes. AsA is clearly an essential dietary component for humans, with aprotective role proposed for many disorders through increasing resistance of immune system.Ascorbic acid (AsA) is a vital compound in plants with a range of functions as a growthregulation factor in regulating flowering time, or as a stress response factor, which is involvedin many stress-induced oxidative processes, e.g., responses to water loss, pathogens andoxidizing agents, high temperature, photo-oxidative stress and ozone. Up to now,GDP-mannose-3',5'-epimerase (GME), GDP-L-galactose phosphotylase (GGP), L-galactosephosphatase (GPP) may play key roles in ASA biosynthesis and regulation. In present study,gene cloning, real-time PCR, and promoter characteristics were used to illustrate themolecular characteristics of enzymes involved in ASA biosynthesis under different treatments.To gain a better understanding of the relationship between AsA concentrations andtranscriptional levels of the above genes, and to investigate the regulatory mechanisms fortheir expression in kiwifruit under stress conditions, we performed a systematic investigationof AsA, mRNA expression, and promoter activities in response to various treatments. For this,we evaluated three kiwifruit species (A. eriantha, A. deliciosa cv.‘qinmei’ and A.rufa) thatproduce different amounts of AsA. Furthermore, to evaluate the relationship between levels ofAsA in fruits development and GGP transcripts, we systematically investigated the regulatorymechanisms for GGP expression in the three kiwifruit species. The main results were asfollows:
1. The full-length cDNA sequence of GME gene was cloned from A. deliciosa‘qinmei’and A.rufa, named AdGME (GenBank Accession NO. GU339037) and ArGME (GenBankAccession NO. JN132110). They were both1143-bp long, both encoded a polypeptide of376amino acids. Using the MegAlign program of MEGA, we determined that these amino acidsequences shared99.7%identity. RT-qPCR analyses in two species demonstrated that ABA, dark or light did not seem to alter GME expression, but GA3resulted in a decrease in thetranscript levels of GME. However, application of SA, wounding, cold or heat, GMEtranscript exhibited an increase in both species.
2. The promoters of GME genes were cloned by Genomic DNA-walking method. Theywere both1225-bp long, named ParGME和PadGME. We comparatively analysed thepromoter sequences of ParGME (GenBank Accession NO. JQ693500) and PadGME(GenBank Accession NO. JQ693499). Sequence alignment revealed that they shared99.16%similarity. They showed transcriptional activation identified by transformed tobacco GUShistochemical assay. The promoter activities of GME genes were not significantly inducedwhen the transformed tobacco exposing to light or wounding. While the promoter activities ofGME genes were significantly induced when treated by SA or heat.
3. The full-length cDNA sequence of AdGPP gene was cloned from A. deliciosa. Theexpression patterns of AdGPP gene were studied when kiwifruit plants were exposing todifferent conditions. RT-qRCR analyses demonstrated: during the first12h of darkness,AdGPP expression gradually but continually declined. After48h of dark treatment, plantswere then exposed to light, and AdGPP expression quickly recovered within1h. Following apeak at Hour2, then AdGPP expression began to decrease to the original level. Treatmentwith SA caused only a slight increase in expression over the entire period. ABA or hypoxicconditions were associated with high AdGPP expression at first, followed by a gradualdecline in transcript levels.
4. The AdGPP promoter was isolated from A. deliciosa genomic DNA by GenomicDNA-walking method. After sequencing, we cloned a1395-bp fragment (GenBank AccessionNo. JX122767). The AdGPP promoter showed transcriptional activation identified bytransformed tobacco GUS histochemical assay. The promoter activities of AdGPP were notsignificantly induced when the transformed tobacco exposing to light or wounding. While thepromoter activities of AdGPP gene were significantly induced when treated by SA or heat.
5. The full-length cDNA sequence of GGP genes were cloned from A.eriantha andA.rufa, named AeGGP (GenBank Accession No. KC146049) and ArGGP (GenBankAccession No. KC146048). Some correlation between relative levels of GGP mRNA andAsA concentrations in three species during fruit development. Transcripts were the mostabundant in A. eriantha, and the lowest in A. rufa. RT-qPCR results with the kiwifruit leavesrevealed that expression of the GGP genes was induced by light, MeJA, heat, hypoxicconditions; and was very sensitive to light conditions.
6. The promoters of GGP genes were cloned from three kiwifruit species by GenomicDNA-walking method. GUS activity of the entire AeGGP promoter was the highest and that of ArGGP promoter was the lowest under normal growing conditions. When compared thethree promoters, we found that ArGGP promoter has a nearly300-bp insertion. The result of5`deletion derivates transformed into tobacco indicates that some negative cis-elements mayexist in the ArGGP promoter. The promoter activities of GGP genes were significantlyinduced when application of ABA or MeJA. Light, heat or hypoxic conditions can also inducethe promoter activities of GGP genes. According to the results of5`deletion derivates ofAeGGP or AdGGP promoter, we deduced that G-box is important for the expression of GGPin kiwifruit under different types of illumination and ABRE or CGTCA-motif may play animportant role in regulating GGP promoter activities after application of ABA or MeJArespectively.
引文
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