石斛属植物亲缘关系研究及ACC氧化酶基因的克隆
摘要
石斛属植物为兰科多年生植物,是兰科最大的属之一,也是重要的园艺植物。石斛属植物除了具有较高的观赏价值外,还有部分品种也是我国传统中药中的重要药材。对石斛属植物的传统分类主要是依靠形态学和地理分布,分类的准确性较低,而且不同学者有不同的分类体系,往往得到的差异较大。近几年,分子标记已经应用在石斛的遗传多样性和亲缘关系分析的研究中。
铁皮石斛又被称为黑节草,是药用石斛中最为珍贵的品种之一,常生长于海拔500-1600M的山间河谷的石头或树上。由于它的生境条件较为独特,自身繁殖能力较低,并且人们过度采挖,现在铁皮石斛的野生资源濒临灭绝。为保护铁皮石斛资源和增加药源,现已有许多学者在铁皮石斛组织培养快繁方面开展研究,但对铁皮石斛人工授粉获得种子后再进行无菌播种的研究和利用分子标记进行亲缘关系的研究相对较少。
ACC氧化酶(1-aminocyclopropane-carboxylic oxidase)是乙烯合成演化中的-个关键酶。乙烯可以诱导如苯丙氨酸解氨酶(PAL)、过氧化物酶、多酚氧化酶和几丁质酶的活性。有报道显示,铁皮石斛的生物碱含量与PAL的活性有着密切的关系。
采用ISSR分子标记技术对收集到的26个石斛属材料进行了亲缘关系的分析,并研究了铁皮石斛人工授粉与无菌萌发的条件。此外,克隆了铁皮石斛的ACC氧化酶基因,并在大肠杆菌中得到了高效表达。主要研究结果如下：
1.采用ISSR标记,对26个材料的亲缘关系进行了分析。从100条ISSR引物中筛选出5条引物对26个石斛材料DNA进行PCR扩增。共扩增出了70条带,其中63条具有多态性,多态性条带百分率为90.00%,平均为每个引物12.6条多态性带。
2利用计算机软件DPS软件统计ISSR数据,得出所研究的26个材料遗传距离介于0.0334-0.7314之间；用UPGMA进行聚类,当遗传距离为0.43时,可以将26份材料聚为6大类群：第一类：尖刀唇石斛、迭鞘石斛、流苏石斛、束花石斛、玫瑰石斛、大苞鞘石斛、重唇石斛、兜唇石斛、金钗石斛、报春石斛、齿瓣石斛、串珠石斛、细茎石斛和铁皮石斛；第二类：美花石斛、长距石斛、短棒石斛、聚石斛、细叶石斛和竹枝石斛；第三类：鼓槌石斛和密花石斛；第四类：黑毛石斛和翅萼石斛；第五类：刀叶石斛；第六类：翅梗石斛。
3.通过采用(1)异株异花授粉；(2)同株不同花序异花；(3)同花序异花授粉；(4)自花授粉；(5)对照(套袋后自然授粉)等方式进行人工授粉,结果表明铁皮石斛花在开放后的第2d至第4d时采用异株异花,且在每天上午10点以前的人工授粉成功率最高。
4.选择通过人工授粉获得的健康种子(8个月种龄)进行无菌播种,结果表明,铁皮石斛种子萌发最佳培养体系：MS+NAA0.5mg/L+土豆汁10%+活性炭0.5%暗处理7天后转光培养。
5.通过PCR的方法,从铁皮石斛的花中克隆了一个同源ACC氧化酶基因。克隆到的cDNA长度为970bp(在NCBI中登录号为JX679494),含有一个编码314个氨基酸的蛋白质。
6.构建了pET-28a-ACO原核表达载体,并转入大肠杆菌(Escherichia coli)中表达,获得1个与预测大小一致的外源蛋白。该研究为生物调控铁皮石斛的生物碱含量提供了理论基础。
Dendrobium species are perennial plants of orchidaceae. Dendrobium is one of the largest genus of the orchid, and also an important horticultural plants. Many species of Dendrobium are precious ornamental plants and tranditional Chinese medicine herbals. The traditional classification of the genus Dendrobium relies mainly on morphology and geographical distribution, but the classification accuracy is relatively low, and different researchers have different classification system, thus there exists a lot of difference.
In recent years, molecular markers methods have been applied in Dendrobium genetic diversity and relationship analysis.
As one kind of Orchidaceae, Dendrobium officinale is also known as Dendrobium candidum, one of the most precious medicinal Dendrobium species. Dendrobium officinale is attached on the wet trunk of trees or in the crack of stones of500-1600meters above sea level. Its habitat conditions is very unique and the ability to reproduce is very weak. In addition,in recent years the exeessive gather and the destruction of natural habitats, Deandidum officinale wildiife resources increasingly decreaseing. To protect the of Dendrobium officinale resources and increase drug source, now there are many scholars do researches on the tissue culture of Dendrobium officinale and the rapid propagation, but there are very few researches on sterile sowing the seeds obtained by artificial pollination and genetic diversity analysis based on molecular markers.
ACC oxidase (1-aminocyclopropane-carboxylic oxidase) is a key enzyme in the biosynthesis of ethylene.Ethylene can induce enzyme involved in metabolite biosynthesis such as phenylalanine ammonia-lyase (PAL), peroxidase, polyphenol oxidase and chitinaseo some researchers reported that PAL is the key factor for the synthesis of Dendrobium alkaloids
ISSR molecular marker technology was used to analysis the genetic relationship between the26materials of the genus Dendrobium, and do some studies on the conditions of Dendrobium officinale artificial pollination and sterile germination. In addition, we cloned and express the ACC oxidase gene of Dendrobium officinale in E. coli. The main findings are as follows:
1.The genetic relationship of the26materials were analyzed by using ISSR Marker.5Primers were chosen from100ISSR-PCR primers for PCR amplification of Dendrobium material DNA,70bands were amplified, and63bands of them were polymorphic(percentage of polymorphic bands was90%) with an average of12.6polymorphic bands per primer.
2.Using DPS software to deal with all date. Cluster analysis indicated the genetic distance of26materials was0.0334-0.7314, and all the materials could be divided into six groups with the genetic distance at0.43. I:Den.heterocarpum Lindl.、Den.denneanum Kerr.、 Den.fimbriatum Hook.、Den.chrysanthum Lindl.、Den.crepidatum Lindl.ex Paxt.、 Den.wardianum Warner.、Den.hercoglossum Rchb.f.、Den. aphyllum(Roxb.)C.E.Fisher.、 Den. nobile Lindl.、Den.primulinum Lindl.、Den.devonianum Paxt.、Den.falconeri Hook.、 Den.moniliforme(L.)Sw.、Den.officinale Kimura et Migo.;Ⅱ:Den.loddigesii Rolfe.、Den. Longicornu Lindl.、Den.Capillipes Rchb.f.、Den.lindleyi Stendel.、Den.hancockii Rolfe. Den.salaccens(Bl.)Lindl.;Ⅲ:Den.chrysotoxum Lindl.、Den.densiflorum Lindl.;Ⅳ: Den.williamsonii Day et Rchb.f.、Den.cariniferum Rchb.f.; V:Den.terminale Par.et Rchb.f.; Ⅵ:Den.trigonopus Rchb.f.
3. Artificial pollination was performed by using:(1) xenogamy;(2) different inflorescence pollination within one plant;(3) different flwers pollination within one inflorescence;(4) the self pollination;(5) the control (natural pollination after bagging), the results showed that the seeding rate reaches the highest under cross pollination before10o 'clock in the morning after blooming2d to4d.
4. Choose healthy seeds obtained by artificial pollination methods (8months old seed) for aseptic seeding, it showed that the optimal medium for seeds germination was MS+NAA0.5mg/L+potato juicel0%+active charcoal0.5%, and cultured for7d in the dark then under light.
5. A homolog of ACC oxidase gene (ACO) was isolated from flowers of Dendrobium officinale kimura et migo by PCR-method. The obtained cDNA of Do ACO is970bp long and contains an open reading frame (ORF) encoding a protein with314amino acid residues (The sequence has been submitted to the GenBank data base, the accession number is JX679494)
6. The constructed prokaryotic expression vector pET-28a-ACO was transformed into E. coli BL21(DE3) and expressed. One foreign protein, which was the same size as expectation, was obtained. This research layed the foundation for studying ethylene biosynthesis and had an implication to improve the alkaloids content in the future.
铁皮石斛又被称为黑节草,是药用石斛中最为珍贵的品种之一,常生长于海拔500-1600M的山间河谷的石头或树上。由于它的生境条件较为独特,自身繁殖能力较低,并且人们过度采挖,现在铁皮石斛的野生资源濒临灭绝。为保护铁皮石斛资源和增加药源,现已有许多学者在铁皮石斛组织培养快繁方面开展研究,但对铁皮石斛人工授粉获得种子后再进行无菌播种的研究和利用分子标记进行亲缘关系的研究相对较少。
ACC氧化酶(1-aminocyclopropane-carboxylic oxidase)是乙烯合成演化中的-个关键酶。乙烯可以诱导如苯丙氨酸解氨酶(PAL)、过氧化物酶、多酚氧化酶和几丁质酶的活性。有报道显示,铁皮石斛的生物碱含量与PAL的活性有着密切的关系。
采用ISSR分子标记技术对收集到的26个石斛属材料进行了亲缘关系的分析,并研究了铁皮石斛人工授粉与无菌萌发的条件。此外,克隆了铁皮石斛的ACC氧化酶基因,并在大肠杆菌中得到了高效表达。主要研究结果如下：
1.采用ISSR标记,对26个材料的亲缘关系进行了分析。从100条ISSR引物中筛选出5条引物对26个石斛材料DNA进行PCR扩增。共扩增出了70条带,其中63条具有多态性,多态性条带百分率为90.00%,平均为每个引物12.6条多态性带。
2利用计算机软件DPS软件统计ISSR数据,得出所研究的26个材料遗传距离介于0.0334-0.7314之间；用UPGMA进行聚类,当遗传距离为0.43时,可以将26份材料聚为6大类群：第一类：尖刀唇石斛、迭鞘石斛、流苏石斛、束花石斛、玫瑰石斛、大苞鞘石斛、重唇石斛、兜唇石斛、金钗石斛、报春石斛、齿瓣石斛、串珠石斛、细茎石斛和铁皮石斛；第二类：美花石斛、长距石斛、短棒石斛、聚石斛、细叶石斛和竹枝石斛；第三类：鼓槌石斛和密花石斛；第四类：黑毛石斛和翅萼石斛；第五类：刀叶石斛；第六类：翅梗石斛。
3.通过采用(1)异株异花授粉；(2)同株不同花序异花；(3)同花序异花授粉；(4)自花授粉；(5)对照(套袋后自然授粉)等方式进行人工授粉,结果表明铁皮石斛花在开放后的第2d至第4d时采用异株异花,且在每天上午10点以前的人工授粉成功率最高。
4.选择通过人工授粉获得的健康种子(8个月种龄)进行无菌播种,结果表明,铁皮石斛种子萌发最佳培养体系：MS+NAA0.5mg/L+土豆汁10%+活性炭0.5%暗处理7天后转光培养。
5.通过PCR的方法,从铁皮石斛的花中克隆了一个同源ACC氧化酶基因。克隆到的cDNA长度为970bp(在NCBI中登录号为JX679494),含有一个编码314个氨基酸的蛋白质。
6.构建了pET-28a-ACO原核表达载体,并转入大肠杆菌(Escherichia coli)中表达,获得1个与预测大小一致的外源蛋白。该研究为生物调控铁皮石斛的生物碱含量提供了理论基础。
Dendrobium species are perennial plants of orchidaceae. Dendrobium is one of the largest genus of the orchid, and also an important horticultural plants. Many species of Dendrobium are precious ornamental plants and tranditional Chinese medicine herbals. The traditional classification of the genus Dendrobium relies mainly on morphology and geographical distribution, but the classification accuracy is relatively low, and different researchers have different classification system, thus there exists a lot of difference.
In recent years, molecular markers methods have been applied in Dendrobium genetic diversity and relationship analysis.
As one kind of Orchidaceae, Dendrobium officinale is also known as Dendrobium candidum, one of the most precious medicinal Dendrobium species. Dendrobium officinale is attached on the wet trunk of trees or in the crack of stones of500-1600meters above sea level. Its habitat conditions is very unique and the ability to reproduce is very weak. In addition,in recent years the exeessive gather and the destruction of natural habitats, Deandidum officinale wildiife resources increasingly decreaseing. To protect the of Dendrobium officinale resources and increase drug source, now there are many scholars do researches on the tissue culture of Dendrobium officinale and the rapid propagation, but there are very few researches on sterile sowing the seeds obtained by artificial pollination and genetic diversity analysis based on molecular markers.
ACC oxidase (1-aminocyclopropane-carboxylic oxidase) is a key enzyme in the biosynthesis of ethylene.Ethylene can induce enzyme involved in metabolite biosynthesis such as phenylalanine ammonia-lyase (PAL), peroxidase, polyphenol oxidase and chitinaseo some researchers reported that PAL is the key factor for the synthesis of Dendrobium alkaloids
ISSR molecular marker technology was used to analysis the genetic relationship between the26materials of the genus Dendrobium, and do some studies on the conditions of Dendrobium officinale artificial pollination and sterile germination. In addition, we cloned and express the ACC oxidase gene of Dendrobium officinale in E. coli. The main findings are as follows:
1.The genetic relationship of the26materials were analyzed by using ISSR Marker.5Primers were chosen from100ISSR-PCR primers for PCR amplification of Dendrobium material DNA,70bands were amplified, and63bands of them were polymorphic(percentage of polymorphic bands was90%) with an average of12.6polymorphic bands per primer.
2.Using DPS software to deal with all date. Cluster analysis indicated the genetic distance of26materials was0.0334-0.7314, and all the materials could be divided into six groups with the genetic distance at0.43. I:Den.heterocarpum Lindl.、Den.denneanum Kerr.、 Den.fimbriatum Hook.、Den.chrysanthum Lindl.、Den.crepidatum Lindl.ex Paxt.、 Den.wardianum Warner.、Den.hercoglossum Rchb.f.、Den. aphyllum(Roxb.)C.E.Fisher.、 Den. nobile Lindl.、Den.primulinum Lindl.、Den.devonianum Paxt.、Den.falconeri Hook.、 Den.moniliforme(L.)Sw.、Den.officinale Kimura et Migo.;Ⅱ:Den.loddigesii Rolfe.、Den. Longicornu Lindl.、Den.Capillipes Rchb.f.、Den.lindleyi Stendel.、Den.hancockii Rolfe. Den.salaccens(Bl.)Lindl.;Ⅲ:Den.chrysotoxum Lindl.、Den.densiflorum Lindl.;Ⅳ: Den.williamsonii Day et Rchb.f.、Den.cariniferum Rchb.f.; V:Den.terminale Par.et Rchb.f.; Ⅵ:Den.trigonopus Rchb.f.
3. Artificial pollination was performed by using:(1) xenogamy;(2) different inflorescence pollination within one plant;(3) different flwers pollination within one inflorescence;(4) the self pollination;(5) the control (natural pollination after bagging), the results showed that the seeding rate reaches the highest under cross pollination before10o 'clock in the morning after blooming2d to4d.
4. Choose healthy seeds obtained by artificial pollination methods (8months old seed) for aseptic seeding, it showed that the optimal medium for seeds germination was MS+NAA0.5mg/L+potato juicel0%+active charcoal0.5%, and cultured for7d in the dark then under light.
5. A homolog of ACC oxidase gene (ACO) was isolated from flowers of Dendrobium officinale kimura et migo by PCR-method. The obtained cDNA of Do ACO is970bp long and contains an open reading frame (ORF) encoding a protein with314amino acid residues (The sequence has been submitted to the GenBank data base, the accession number is JX679494)
6. The constructed prokaryotic expression vector pET-28a-ACO was transformed into E. coli BL21(DE3) and expressed. One foreign protein, which was the same size as expectation, was obtained. This research layed the foundation for studying ethylene biosynthesis and had an implication to improve the alkaloids content in the future.
引文
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