苹果山梨醇代谢相关基因的分子特性研究
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摘要
山梨醇是许多蔷薇科植物中主要的光合产物,也是碳水化合物的主要运输形式和贮藏物质,与植物的生长、果实品质和产量等密切相关。研究表明,山梨醇还可以作为小分子渗透物质当植物遭遇逆境时在体内积累。山梨醇的积累对提高植物抗旱、抗盐碱、抵御低温、抗火疫病、清除活性氧等能力具有重要作用。山梨醇还能与硼形成复合物而提高植物体内硼的运输能力。参与山梨醇代谢的关键酶(蛋白)包括6-磷酸山梨醇脱氢酶(S6PDH)、山梨醇运输蛋白(SOT)和山梨醇脱氢酶(SDH)。本研究以‘嘎拉’苹果(Malus domestica cv. Gala)为材料,克隆了与山梨醇代谢相关的基因S6PDH、SOT和SDH,并利用实时荧光定量PCR分别分析了这些基因在苹果不同时空的表达模式;获得了S6PDH基因启动子序列并进行了活性分析;利用GFP和胶体金免疫电镜技术研究了S6PDH在细胞中的定位。获得的主要结果如下:
1.克隆了S6PDH基因cDNA全长序列,该序列长1080bp,编码310个氨基酸。获得了两个大小分别为3595bp和3402bp的S6PDH基因gDNA序列。结构分析表明,两个gDNA序列都是由6个外显子和5个内含子组成。两个gDNA序列大小不同是由于后三个内含子长度不同造成的。
实时定量RT-PCR分析发现,S6PDH基因在幼叶中相对表达水平较低,随叶龄的增加基因表达水平明显提高,在成熟叶片中达到最大量。随着叶片的衰老,S6PDH基因的表达水平呈现显著的降低趋势。此过程中S6PDH酶活性与基因表达呈现出基本一致的变化趋势。在干旱胁迫和盐胁迫条件下,S6PDH基因表达量随胁迫的加重而上升,当胁迫达一定程度时S6PDH基因表达急剧下降。结果表明S6PDH的表达与叶片从库到源的功能转化功能密切相关,转录水平是S6PDH表达的关键调控步骤。
将苹果S6PDH基因通过农杆菌介导法转入猕猴桃中,对获得的抗性植株进行了PCR检测和Southern杂交检测,共获得了5个转基因株系。证明已将苹果S6PDH基因转入猕猴桃中。
2.洋葱表皮亚细胞定位结果表明,GFP-S6PDH融合蛋白主要定位于细胞质中。免疫细胞化学标记结果表明,经苹果S6PDH抗体孵育和山羊抗兔免疫球蛋白-胶体金标记后,胶体金颗粒主要沉积于细胞的胞质和叶绿体中,表明苹果叶片中S6PDH主要定位于胞质和叶绿体中,在细胞其他部位没有分布。
3.克隆到了S6PDH基因启动子序列。该序列长2396bp,序列分析表明该启动子区包括多种顺式作用元件。将该序列插入表达载体pC0390-GUS中并转化烟草,经GUS组织化学染色,证明本研究获得的启动子具有转录活性。构建了一系列启动子5`端缺失体并转化烟草,经GUS组织化学染色和活性测定,发现启动子序列中-2122~-1719bp区域和-1066~0bp区域在调节基因转录方面起重要作用。
4.克隆到了3个SDH基因家族成员MdSDH7、MdSDH8和MdSDH9,其全长cDNA序列长度分别为1418bp、1446bp和1388bp,编码371、368和368个氨基酸。它们的gDNA序列长度分别为2176bp、1239bp和1246bp。结构分析表明,MdSDH7的gDNA序列由6个外显子和5个内含子组成,而MdSDH8和MdSDH9的gDNA序列都是由2个外显子和1个内含子组成。
实时定量RT-PCR的分析结果发现,在整个果实生长过程中,MdSDH7基因在果实生长后期表达较高,MdSDH8基因在果实生长中、后期表达较高,MdSDH9基因在果实生长中期表达较高。SDH活性在果实生长初期(花后0d和15d)最高,随着果实的成熟活性开始急剧下降,在花后92d活性有所上升,但增加不大,随后又开始下降。果实相对生长速率与SDH活性变化呈现出相同的模式。
MdSDH8和MdSDH9在幼叶中表达量很高,随叶片的成熟表达量显著下降,而MdSDH7则与它们表现出相反的表达变化趋势。幼叶中SDH活性最高,随叶片成熟活性显著下降,但当叶片进入衰老期后活性又再次升高。在幼果中,MdSDH7、MdSDH8和MdSDH9三个基因都在种子中表达量最高,在果肉和果皮中的表达量差异不大。在成熟果中,MdSDH7基因在种子和果皮中的表达量较高,MdSDH8基因在种子中的表达量较高,MdSDH9基因在果肉中的表达量较高。幼果中SDH的活性在种子最高,果肉和果皮中的活性基本一致。相反的是,成熟果果皮和果肉中SDH活性较高,而种子中SDH活性较低。以上结果表明,果实生长过程中SDH活性与RGR变化呈现一致的趋势,表明SDH活性与果实“库”强度密切相关。SDH活性变化与基因表达模式不一致,这可能是由于基因家族成员共同作用的结果,也可能是SDH基因表达存在转录后调控机制。
5.获得了4个SOT基因家族成员MdSOT7、MdSOT8、MdSOT9和MdSOT10,其全长cDNA序列长度分别为1681bp、1755bp、1797bp和1849bp,编码538、526、491和535个氨基酸。用扩增基因全长的引物对苹果基因组DNA进行扩增,发现MdSOT7、MdSOT8、MdSOT9和MdSOT10的gDNA序列长度分别为2616bp、2230bp、2397bp和3013bp。结构分析表明,4个SOT基因gDNA序列都是由3个外显子和2个内含子组成,但是外显子和内含子的长度和位置差异很大。
实时定量RT-PCR分析表明,MdSOT7基因在各时期叶片中表达水平较低且表达量基本不变,MdSOT8、MdSOT9和MdSOT10三个基因在成熟叶中表达较高。MdSOT7在果实发育初期和后期表达较高,MdSOT8在果实发育中期表达较高,MdSOT9在整个果实发育阶段都有表达且变化不大,MdSOT10在果实发育初期表达量较高。
Sorbitol is the primary photosynthetic product, and major translocated and storaged form of photosynthate in many species of Rosaceae. Those plant development and fruit quality depend on sorbitol. In addition, sorbitol is an efficient osmolyte. Its accumulation can improve plant tolerance of salinity, drought, low temperature, fireblight disease and oxidative stress. Sorbitol can also form complex with boron to facilitate phloem boron transport. In metabolism of sorbitol, NADP-dependent sorbitol-6-phosphate dehydrogenase (S6PDH), sorbitol transporter (SOT), NAD-sorbitol dehydrogenase (SDH) play key roles. Up to now, though sorbitol and its metabolic enzymes have been investigated, information about molecular characteristics of those enzymes is still lacking. In present study, cloning, real-time PCR, sub-cellular localization and transformation were used to illustrate the molecular characteristics of enzymes involved in sorbitol metabolism. The main results were as follows:
1. The full-length cDNA sequence of S6PDH gene was cloned from apple. It was 1080 bp long, encoding 314 amino acid residues. Based on the ORF, two genomic DNA sequences of S6PDH gene were also isolated, one is 3595bp, and the other is 3402bp. These two gDNA sequences had six extrons and five introns. The difference between them was the length of last three introns.
Expression pattern of S6PDH during leaf development were detected by real-time RT-PCR. S6PDH mRNA level was low in young leaves, increased markedly with leaf growth, reached maximum in mature leaves, and then reduced in old leaves, paralleling with the enzyme activity during leaf development. Under salt or drought stress conditions, S6PDH mRNA level was increased prominently with the prolongation of stress treatment.Therefore, the expression of S6PDH seemed closely related with the transition from sink to source function of the leaves, was also suggested to be regulated at the transcriptional level.
The binary plasmid vector pBI-S6PDH including the S6PDH coding region was introduced into kiwifruit by agrobacterium-mediated method. The kanamycin resistant plantlets were identified by PCR and southern blot. Five plantlets were obtained for future research.
2. Subcellular localization analysis with GFP using gun-mediated bombardment method revealed that S6PDH protein was located in cytoplasm of onion epidermal cell. Immunocytochemical study showed that after incubated by antibody of S6PDH and labeled by goat anti-rabbit immunoglobulin, colloidal gold mainly sedimentated in cytoplast and chloroplast, which indicated that S6PDH mainly located in cytoplast and chloroplast in apple leaves.
3. The 2396bp promoter of S6PDH gene was cloned. It included many cis-acting elements and showed transcriptional activation identified by transformed tobacco GUS histochemical assay. The result of 5` deletion derivates transformed into tobacco indicates that the regions of -2122~-1719bp and -1066~0bp may play an important role in regulating gene transcription.
4. Three cDNAs encoding different isoforms of NAD-SDH from apple were isolated. The cDNA of MdSDH7, MdSDH8 and MdSDH9 were 1418bp, 1446bp and 1388bp, encoding 371aa, 368aa and 368aa, respectively. gDNA of MdSDH7 was 2176bp including 6 exons and 5 introns, gDNA of MdSDH8 and MdSDH9 were 1239bp and 1246bp respectively and both include 2 exons and 1 intron.
Expression pattern of SDHs was detected by real-time PCR. MdSDH7 expression level was higher at the late developmental stage of apple fruit,MdSDH8 expression level was higher at both middle and late developmental stages,whereas MdSDH7 expression level was higher at the middle developmental stage. The activity of NAD-SDH was high in young fruit (from 0 to 15 DAF). It dropped significantly as fruit developed, but increased slightly in 92DAF, and then dropped again. In general, the activity of NAD-SDH changed in a similar way as that of the relative growth rate of the fruit.
Expression of MdSDH8 and MdSDH9 were high in source leaves whereas that of MdSDH7 was high in sink leaves. The activity of NAD-SDH was high in young leaf, dropped as leaf became mature, and then increased again in senescent leaf.
In young fruit,the expression level of MdSDH7, MdSDH8 and MdSDH9 were all higher in seed than that in flesh or pericarp. In mature fruit,the expression level of MdSDH7 was higher in seed and pericarp, whereas that of MdSDH8 and MdSDH9 was higher in seed and flesh, respectively. The changes in NAD-SDH activity were proportionate to those in RGR, Thus, it is considered that SDH activity was associated with sink strength. The fluctuation pattern of SDH activity did not correspond to that of the transcripts, these results may be due to interaction of SDH isogenes.
5. Four cDNAs encoding different isoforms of SOT from apple were isolated. The cDNA of MdSOT7, MdSOT8, MdSOT9 and MdSOT10 were 1681bp, 1755bp, 1797bp and 1849bp, encoding 538aa, 526aa, 491aa and 535aa, respectively. gDNA of the four genes were 2616bp, 2230bp, 2397bp and 3013bp, respectively, and all of them include 3 exons and 2 introns which differ in length and position. Expression pattern of SOTs were detected by real-time RT-PCR during leaf and fruit development. The expression level of MdSOT7 was consistent with leaf development. The expression level of MdSOT8, MdSOT9 and MdSOT10 was higher in mature leaf. As apple fruit developed, MdSOT7 expression level was higher in the early and late developmental stages,MdSOT8 expression level was higher in the middle stage, MdSOT9 expression level remained unchanged, and MdSOT10 was higher in the early developmental stage.
1.克隆了S6PDH基因cDNA全长序列,该序列长1080bp,编码310个氨基酸。获得了两个大小分别为3595bp和3402bp的S6PDH基因gDNA序列。结构分析表明,两个gDNA序列都是由6个外显子和5个内含子组成。两个gDNA序列大小不同是由于后三个内含子长度不同造成的。
实时定量RT-PCR分析发现,S6PDH基因在幼叶中相对表达水平较低,随叶龄的增加基因表达水平明显提高,在成熟叶片中达到最大量。随着叶片的衰老,S6PDH基因的表达水平呈现显著的降低趋势。此过程中S6PDH酶活性与基因表达呈现出基本一致的变化趋势。在干旱胁迫和盐胁迫条件下,S6PDH基因表达量随胁迫的加重而上升,当胁迫达一定程度时S6PDH基因表达急剧下降。结果表明S6PDH的表达与叶片从库到源的功能转化功能密切相关,转录水平是S6PDH表达的关键调控步骤。
将苹果S6PDH基因通过农杆菌介导法转入猕猴桃中,对获得的抗性植株进行了PCR检测和Southern杂交检测,共获得了5个转基因株系。证明已将苹果S6PDH基因转入猕猴桃中。
2.洋葱表皮亚细胞定位结果表明,GFP-S6PDH融合蛋白主要定位于细胞质中。免疫细胞化学标记结果表明,经苹果S6PDH抗体孵育和山羊抗兔免疫球蛋白-胶体金标记后,胶体金颗粒主要沉积于细胞的胞质和叶绿体中,表明苹果叶片中S6PDH主要定位于胞质和叶绿体中,在细胞其他部位没有分布。
3.克隆到了S6PDH基因启动子序列。该序列长2396bp,序列分析表明该启动子区包括多种顺式作用元件。将该序列插入表达载体pC0390-GUS中并转化烟草,经GUS组织化学染色,证明本研究获得的启动子具有转录活性。构建了一系列启动子5`端缺失体并转化烟草,经GUS组织化学染色和活性测定,发现启动子序列中-2122~-1719bp区域和-1066~0bp区域在调节基因转录方面起重要作用。
4.克隆到了3个SDH基因家族成员MdSDH7、MdSDH8和MdSDH9,其全长cDNA序列长度分别为1418bp、1446bp和1388bp,编码371、368和368个氨基酸。它们的gDNA序列长度分别为2176bp、1239bp和1246bp。结构分析表明,MdSDH7的gDNA序列由6个外显子和5个内含子组成,而MdSDH8和MdSDH9的gDNA序列都是由2个外显子和1个内含子组成。
实时定量RT-PCR的分析结果发现,在整个果实生长过程中,MdSDH7基因在果实生长后期表达较高,MdSDH8基因在果实生长中、后期表达较高,MdSDH9基因在果实生长中期表达较高。SDH活性在果实生长初期(花后0d和15d)最高,随着果实的成熟活性开始急剧下降,在花后92d活性有所上升,但增加不大,随后又开始下降。果实相对生长速率与SDH活性变化呈现出相同的模式。
MdSDH8和MdSDH9在幼叶中表达量很高,随叶片的成熟表达量显著下降,而MdSDH7则与它们表现出相反的表达变化趋势。幼叶中SDH活性最高,随叶片成熟活性显著下降,但当叶片进入衰老期后活性又再次升高。在幼果中,MdSDH7、MdSDH8和MdSDH9三个基因都在种子中表达量最高,在果肉和果皮中的表达量差异不大。在成熟果中,MdSDH7基因在种子和果皮中的表达量较高,MdSDH8基因在种子中的表达量较高,MdSDH9基因在果肉中的表达量较高。幼果中SDH的活性在种子最高,果肉和果皮中的活性基本一致。相反的是,成熟果果皮和果肉中SDH活性较高,而种子中SDH活性较低。以上结果表明,果实生长过程中SDH活性与RGR变化呈现一致的趋势,表明SDH活性与果实“库”强度密切相关。SDH活性变化与基因表达模式不一致,这可能是由于基因家族成员共同作用的结果,也可能是SDH基因表达存在转录后调控机制。
5.获得了4个SOT基因家族成员MdSOT7、MdSOT8、MdSOT9和MdSOT10,其全长cDNA序列长度分别为1681bp、1755bp、1797bp和1849bp,编码538、526、491和535个氨基酸。用扩增基因全长的引物对苹果基因组DNA进行扩增,发现MdSOT7、MdSOT8、MdSOT9和MdSOT10的gDNA序列长度分别为2616bp、2230bp、2397bp和3013bp。结构分析表明,4个SOT基因gDNA序列都是由3个外显子和2个内含子组成,但是外显子和内含子的长度和位置差异很大。
实时定量RT-PCR分析表明,MdSOT7基因在各时期叶片中表达水平较低且表达量基本不变,MdSOT8、MdSOT9和MdSOT10三个基因在成熟叶中表达较高。MdSOT7在果实发育初期和后期表达较高,MdSOT8在果实发育中期表达较高,MdSOT9在整个果实发育阶段都有表达且变化不大,MdSOT10在果实发育初期表达量较高。
Sorbitol is the primary photosynthetic product, and major translocated and storaged form of photosynthate in many species of Rosaceae. Those plant development and fruit quality depend on sorbitol. In addition, sorbitol is an efficient osmolyte. Its accumulation can improve plant tolerance of salinity, drought, low temperature, fireblight disease and oxidative stress. Sorbitol can also form complex with boron to facilitate phloem boron transport. In metabolism of sorbitol, NADP-dependent sorbitol-6-phosphate dehydrogenase (S6PDH), sorbitol transporter (SOT), NAD-sorbitol dehydrogenase (SDH) play key roles. Up to now, though sorbitol and its metabolic enzymes have been investigated, information about molecular characteristics of those enzymes is still lacking. In present study, cloning, real-time PCR, sub-cellular localization and transformation were used to illustrate the molecular characteristics of enzymes involved in sorbitol metabolism. The main results were as follows:
1. The full-length cDNA sequence of S6PDH gene was cloned from apple. It was 1080 bp long, encoding 314 amino acid residues. Based on the ORF, two genomic DNA sequences of S6PDH gene were also isolated, one is 3595bp, and the other is 3402bp. These two gDNA sequences had six extrons and five introns. The difference between them was the length of last three introns.
Expression pattern of S6PDH during leaf development were detected by real-time RT-PCR. S6PDH mRNA level was low in young leaves, increased markedly with leaf growth, reached maximum in mature leaves, and then reduced in old leaves, paralleling with the enzyme activity during leaf development. Under salt or drought stress conditions, S6PDH mRNA level was increased prominently with the prolongation of stress treatment.Therefore, the expression of S6PDH seemed closely related with the transition from sink to source function of the leaves, was also suggested to be regulated at the transcriptional level.
The binary plasmid vector pBI-S6PDH including the S6PDH coding region was introduced into kiwifruit by agrobacterium-mediated method. The kanamycin resistant plantlets were identified by PCR and southern blot. Five plantlets were obtained for future research.
2. Subcellular localization analysis with GFP using gun-mediated bombardment method revealed that S6PDH protein was located in cytoplasm of onion epidermal cell. Immunocytochemical study showed that after incubated by antibody of S6PDH and labeled by goat anti-rabbit immunoglobulin, colloidal gold mainly sedimentated in cytoplast and chloroplast, which indicated that S6PDH mainly located in cytoplast and chloroplast in apple leaves.
3. The 2396bp promoter of S6PDH gene was cloned. It included many cis-acting elements and showed transcriptional activation identified by transformed tobacco GUS histochemical assay. The result of 5` deletion derivates transformed into tobacco indicates that the regions of -2122~-1719bp and -1066~0bp may play an important role in regulating gene transcription.
4. Three cDNAs encoding different isoforms of NAD-SDH from apple were isolated. The cDNA of MdSDH7, MdSDH8 and MdSDH9 were 1418bp, 1446bp and 1388bp, encoding 371aa, 368aa and 368aa, respectively. gDNA of MdSDH7 was 2176bp including 6 exons and 5 introns, gDNA of MdSDH8 and MdSDH9 were 1239bp and 1246bp respectively and both include 2 exons and 1 intron.
Expression pattern of SDHs was detected by real-time PCR. MdSDH7 expression level was higher at the late developmental stage of apple fruit,MdSDH8 expression level was higher at both middle and late developmental stages,whereas MdSDH7 expression level was higher at the middle developmental stage. The activity of NAD-SDH was high in young fruit (from 0 to 15 DAF). It dropped significantly as fruit developed, but increased slightly in 92DAF, and then dropped again. In general, the activity of NAD-SDH changed in a similar way as that of the relative growth rate of the fruit.
Expression of MdSDH8 and MdSDH9 were high in source leaves whereas that of MdSDH7 was high in sink leaves. The activity of NAD-SDH was high in young leaf, dropped as leaf became mature, and then increased again in senescent leaf.
In young fruit,the expression level of MdSDH7, MdSDH8 and MdSDH9 were all higher in seed than that in flesh or pericarp. In mature fruit,the expression level of MdSDH7 was higher in seed and pericarp, whereas that of MdSDH8 and MdSDH9 was higher in seed and flesh, respectively. The changes in NAD-SDH activity were proportionate to those in RGR, Thus, it is considered that SDH activity was associated with sink strength. The fluctuation pattern of SDH activity did not correspond to that of the transcripts, these results may be due to interaction of SDH isogenes.
5. Four cDNAs encoding different isoforms of SOT from apple were isolated. The cDNA of MdSOT7, MdSOT8, MdSOT9 and MdSOT10 were 1681bp, 1755bp, 1797bp and 1849bp, encoding 538aa, 526aa, 491aa and 535aa, respectively. gDNA of the four genes were 2616bp, 2230bp, 2397bp and 3013bp, respectively, and all of them include 3 exons and 2 introns which differ in length and position. Expression pattern of SOTs were detected by real-time RT-PCR during leaf and fruit development. The expression level of MdSOT7 was consistent with leaf development. The expression level of MdSOT8, MdSOT9 and MdSOT10 was higher in mature leaf. As apple fruit developed, MdSOT7 expression level was higher in the early and late developmental stages,MdSOT8 expression level was higher in the middle stage, MdSOT9 expression level remained unchanged, and MdSOT10 was higher in the early developmental stage.
引文
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