富士苹果短枝型性状相关基因的分离及表达
摘要
芽变选种是优中选优，是果树品种改良的重要途径之一。通过芽变选种，使果树株型、着色等农艺和品质性状得到了有效改良，推动了果树品种结构的优化调整与升级换代。短枝型芽变品种因其优异的株型特征，使矮化密植集约化栽培方式得到有力的推广，推动了世界苹果栽培制度的变革，促进了苹果产业的高效发展。所以，进一步探讨短枝型芽变的形成机理对苹果品种改良及苹果产业发展具有重要意义。
本文以富士苹果短枝型芽变‘龙富短枝’和非短枝型对照品种‘长富二号’为试材，深入研究短枝型芽变形成的调控机理。文中分别测量了两试材枝条的节间长度和果型指数等指标；测定了两试材的内源激素赤霉素含量；克隆了赤霉素合成途径中关键基因，并对其表达模式进行了分析；以‘龙富短枝’为研究材料，‘长富二号’作为参照，采用抑制性差减杂交技术分析了两者基因组表达的差异；克隆了赤霉素受体MdGID1基因和DELLA蛋白，并分析其表达模式。主要研究成果如下：
1.测量了‘龙富短枝’和‘长富二号’枝条的节间长度及果型指数等指标，结果发现，在花后的4个时期，两试材枝条的节间长度均存在明显差异，‘龙富短枝’枝条节间的平均长度为2.0cm，‘长富二号’指条节间的平均长度为2.6cm。二者的果型指数、单果重及果实糖酸比差异均不显著。
2.利用HPLC测定了两试材内源激素赤霉素的含量，结果发现，在花后四个时期，赤霉素含量在两试材中均呈先升高后下降的趋势。在花后80d，两试材的赤霉素含量存在明显差异，‘龙富短枝’中赤霉素含量为110ng·g-1FW，‘长富二号’为249ng·g-1FW。初步结果认为，由于‘龙富短枝’中赤霉素含量的降低，使其发生了短枝型突变现象
3.利用同源克隆技术分离得到赤霉素合成途径中关键基因GA20-氧化酶和贝壳杉烯氧化酶cDNA序列，并分析了其在两试材中的表达模式。结果显示，GA20-氧化酶和贝壳杉烯氧化酶cDNA序列在两试材中相同，无碱基突变、插入或缺失现象；其相对表达量的变化趋势与赤霉素含量的变化趋势相似。初步结果表明，GA20-氧化酶和贝壳杉烯氧化酶基因在两试材中的表达量差异，是造成其赤霉素含量变化的原因。
4.利用抑制性差减杂交结合Real-time PCR技术，分析了在两材料中差异表达的基因，并根据其功能对差异基因进行分类。结果发现，两试材中有大量差异表达的基因，主要参与代谢过程、转录调控、转运、光形态建成和响应胁迫。
5.利用RT-PCR和RACE技术分离得到短枝型芽变‘龙富短枝’中赤霉素受体MdGID1基因cDNA序列。结果显示，MdGID1全长1041bp，编码346个氨基酸；序列比对及进化树分析结果显示MdGID1与其它植物中赤霉素受体GID1同源性很高。初步证明MdGID1是苹果中的赤霉素受体。
6.利用High-Tail技术分离得到MdGID1的基因组及其上游调控序列，并对其进行分析。结果表明，MdGID1含有两个外显子和一个内含子；上游调控序列具有植物激素、光和温度等顺式作用元件。Real-time PCR分析发现，MdGID1在苹果枝条、叶片、果实、花和芽中均有表达，但在枝条和花中表达量较高，在短枝型芽变‘龙富短枝’和对照‘长富二号’中的表达量存在明显差异,在花后80d，在‘长富二号’相对表达量是‘龙富短枝’的2～10倍。
7.构建了原核表达载体pJET-MdGID1和pGEX-4T-MdRGL，并成功表达了目的蛋白，为深入研究其蛋白结构和功能奠定基础。
As one of fruit tree breeding methods, bud sport selection is an important way toimprove the plant-type, coloring and other characterizations, which is good for themodification of varieties composition and upgradation of fruit tree. Because of its excellentplant-type, spur type varieties promotes the dwarf and close planting method, thetransformation of the world apple cultivation system and the efficient development of appleindustry. Therefore, it is necessary to further study the mechanism of spur-type mutantformation, then improve apple varieties and apple industry development.
In this study, the shoot internode length, fruit shape index, and plant hormonegibberellins acid of common apple ‘Changfu’ and its spur type bud sport ‘Longfu’ have beendetermined; two key genes in gibberellins acid synthesis pathway, GA20ox and KO, wereisolated from ‘Longfu’ and ‘Changfu2’, the expression pattern of GA20ox and KO were alsostudied; The suppression subtractive hybridization method have been used to identify thedifferentially expressed genes in ‘Longfu’ and ‘Changfu2’; a gibberellins acid receptor,MdGID1, regulatly the gibberellins acid response, was isolated from ‘Longfu’, itsexpressionpattern also studied. The main results are as follows:
1. The data showed that there was no significant difference between two species in fruitshape index, fruit weight and ratio of sugar to acid during the four stages after flowering.But the obvious difference was found in the internode length. The average length of‘Longfu’ was2.0cm, which was shorter than that of ‘Changfu2’ by0.6cm.
2. The result of gibberellin (GA) content indicated that in the whole process, the change ofGA content was similar in two materials, namely first increased, then was in a downwardtrend. However, there was a large difference of GA content between two materials at the80-day after flowering. The GA content in the branch of ‘Longfu’ was110ng·g-1FW.that was in ‘Changfu2’249ng·g-1FW, which suggested that the low GA content in‘Longfu’ caused the spur type phenomenon.
3. Based on the homologous clone, the genes of GA20-oxidase and kaurene oxidase wereisolated, which were two key enzymes of GA synthesis pathway. The sequence analysisshowed that there were no mutantion, insertion and deletion in the sequences of two genes between two species. The expression profiles indicated that their expressions wereparalleled with the change of GA content, which caused the GA content difference in twospecies.
4. Using suppression subtractive hybridization (SSH) and real-time quantitive PCRtechnologies, it was found that the expression difference of a large number of genesmainly involved in metabolic processes, transcriptional regulation, transportation,photomorphogenesis and stress response.
5. By RT-PCE and RACE techniques, a GA receptor, MdGID1was isolated. The sequencephylogenetic analysis indicated that the full-length of MdGID1was1041bp, encoding346amino acids, had identity with GID1in other plants, which suggested it was probablythe GA receptor GID1.
6. The upstream sequence of MdGID1was isolated by the High-Tail method. The sequenceanalysis showed there were two exons and one intron in MdGID1. Its upstream sequencecontained several cis-acting elements, including hormones, light and temperature.Real-time quantitive analysis indicated that MdGID1were expressed in all tested tissues,especially high in branch and flower. And its expression was different between twospecies. At the80-day after flowering, its expression was higher in’Changfu2’ than thatin ‘Longfu’ by ten times.
7. For the further research of the structure and function of MdGID1and MdRGL, theprokaryotic expression vector pJET-MdGID1and pGEX-4T-MdRGL were constructedand successfully expressed the target protein.
本文以富士苹果短枝型芽变‘龙富短枝’和非短枝型对照品种‘长富二号’为试材，深入研究短枝型芽变形成的调控机理。文中分别测量了两试材枝条的节间长度和果型指数等指标；测定了两试材的内源激素赤霉素含量；克隆了赤霉素合成途径中关键基因，并对其表达模式进行了分析；以‘龙富短枝’为研究材料，‘长富二号’作为参照，采用抑制性差减杂交技术分析了两者基因组表达的差异；克隆了赤霉素受体MdGID1基因和DELLA蛋白，并分析其表达模式。主要研究成果如下：
1.测量了‘龙富短枝’和‘长富二号’枝条的节间长度及果型指数等指标，结果发现，在花后的4个时期，两试材枝条的节间长度均存在明显差异，‘龙富短枝’枝条节间的平均长度为2.0cm，‘长富二号’指条节间的平均长度为2.6cm。二者的果型指数、单果重及果实糖酸比差异均不显著。
2.利用HPLC测定了两试材内源激素赤霉素的含量，结果发现，在花后四个时期，赤霉素含量在两试材中均呈先升高后下降的趋势。在花后80d，两试材的赤霉素含量存在明显差异，‘龙富短枝’中赤霉素含量为110ng·g-1FW，‘长富二号’为249ng·g-1FW。初步结果认为，由于‘龙富短枝’中赤霉素含量的降低，使其发生了短枝型突变现象
3.利用同源克隆技术分离得到赤霉素合成途径中关键基因GA20-氧化酶和贝壳杉烯氧化酶cDNA序列，并分析了其在两试材中的表达模式。结果显示，GA20-氧化酶和贝壳杉烯氧化酶cDNA序列在两试材中相同，无碱基突变、插入或缺失现象；其相对表达量的变化趋势与赤霉素含量的变化趋势相似。初步结果表明，GA20-氧化酶和贝壳杉烯氧化酶基因在两试材中的表达量差异，是造成其赤霉素含量变化的原因。
4.利用抑制性差减杂交结合Real-time PCR技术，分析了在两材料中差异表达的基因，并根据其功能对差异基因进行分类。结果发现，两试材中有大量差异表达的基因，主要参与代谢过程、转录调控、转运、光形态建成和响应胁迫。
5.利用RT-PCR和RACE技术分离得到短枝型芽变‘龙富短枝’中赤霉素受体MdGID1基因cDNA序列。结果显示，MdGID1全长1041bp，编码346个氨基酸；序列比对及进化树分析结果显示MdGID1与其它植物中赤霉素受体GID1同源性很高。初步证明MdGID1是苹果中的赤霉素受体。
6.利用High-Tail技术分离得到MdGID1的基因组及其上游调控序列，并对其进行分析。结果表明，MdGID1含有两个外显子和一个内含子；上游调控序列具有植物激素、光和温度等顺式作用元件。Real-time PCR分析发现，MdGID1在苹果枝条、叶片、果实、花和芽中均有表达，但在枝条和花中表达量较高，在短枝型芽变‘龙富短枝’和对照‘长富二号’中的表达量存在明显差异,在花后80d，在‘长富二号’相对表达量是‘龙富短枝’的2～10倍。
7.构建了原核表达载体pJET-MdGID1和pGEX-4T-MdRGL，并成功表达了目的蛋白，为深入研究其蛋白结构和功能奠定基础。
As one of fruit tree breeding methods, bud sport selection is an important way toimprove the plant-type, coloring and other characterizations, which is good for themodification of varieties composition and upgradation of fruit tree. Because of its excellentplant-type, spur type varieties promotes the dwarf and close planting method, thetransformation of the world apple cultivation system and the efficient development of appleindustry. Therefore, it is necessary to further study the mechanism of spur-type mutantformation, then improve apple varieties and apple industry development.
In this study, the shoot internode length, fruit shape index, and plant hormonegibberellins acid of common apple ‘Changfu’ and its spur type bud sport ‘Longfu’ have beendetermined; two key genes in gibberellins acid synthesis pathway, GA20ox and KO, wereisolated from ‘Longfu’ and ‘Changfu2’, the expression pattern of GA20ox and KO were alsostudied; The suppression subtractive hybridization method have been used to identify thedifferentially expressed genes in ‘Longfu’ and ‘Changfu2’; a gibberellins acid receptor,MdGID1, regulatly the gibberellins acid response, was isolated from ‘Longfu’, itsexpressionpattern also studied. The main results are as follows:
1. The data showed that there was no significant difference between two species in fruitshape index, fruit weight and ratio of sugar to acid during the four stages after flowering.But the obvious difference was found in the internode length. The average length of‘Longfu’ was2.0cm, which was shorter than that of ‘Changfu2’ by0.6cm.
2. The result of gibberellin (GA) content indicated that in the whole process, the change ofGA content was similar in two materials, namely first increased, then was in a downwardtrend. However, there was a large difference of GA content between two materials at the80-day after flowering. The GA content in the branch of ‘Longfu’ was110ng·g-1FW.that was in ‘Changfu2’249ng·g-1FW, which suggested that the low GA content in‘Longfu’ caused the spur type phenomenon.
3. Based on the homologous clone, the genes of GA20-oxidase and kaurene oxidase wereisolated, which were two key enzymes of GA synthesis pathway. The sequence analysisshowed that there were no mutantion, insertion and deletion in the sequences of two genes between two species. The expression profiles indicated that their expressions wereparalleled with the change of GA content, which caused the GA content difference in twospecies.
4. Using suppression subtractive hybridization (SSH) and real-time quantitive PCRtechnologies, it was found that the expression difference of a large number of genesmainly involved in metabolic processes, transcriptional regulation, transportation,photomorphogenesis and stress response.
5. By RT-PCE and RACE techniques, a GA receptor, MdGID1was isolated. The sequencephylogenetic analysis indicated that the full-length of MdGID1was1041bp, encoding346amino acids, had identity with GID1in other plants, which suggested it was probablythe GA receptor GID1.
6. The upstream sequence of MdGID1was isolated by the High-Tail method. The sequenceanalysis showed there were two exons and one intron in MdGID1. Its upstream sequencecontained several cis-acting elements, including hormones, light and temperature.Real-time quantitive analysis indicated that MdGID1were expressed in all tested tissues,especially high in branch and flower. And its expression was different between twospecies. At the80-day after flowering, its expression was higher in’Changfu2’ than thatin ‘Longfu’ by ten times.
7. For the further research of the structure and function of MdGID1and MdRGL, theprokaryotic expression vector pJET-MdGID1and pGEX-4T-MdRGL were constructedand successfully expressed the target protein.
引文
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