牡丹花器官发育相关基因的克隆与表达
摘要
牡丹(Paeonia suffruticosa)是中国十大传统名花之一,也是我国的候选国花,具有极高的观赏价值。利用基因工程技术培育花卉新品种,是世界花卉育种的研究热点之一。迄今为止,尽管许多植物的花器官发育相关基因已被分离和克隆,但是关于牡丹的研究鲜有报道。
本研究以牡丹品种‘赵粉’为试材,通过同源克隆的办法,采用RT-PCR和RACE技术,分别从其萼片、花瓣、雄蕊和心皮中成功分离了牡丹PsAP1、PsAP2、PsPI、PsMADS1、PsMADS9、PsAG和PsMADS5这7个花器官形成相关基因的cDNA全长,并通过相对荧光定量PCR的方法,对这些基因在不同器官及花芽分化过程中的时空表达模式进行了分析。同时,分别构建了这7个基因的正义和反义植物表达载体,转化拟南芥和烟草,经过RT-PCR后,得到部分阳性植株,并对其表型初步进行初步观察。主要研究结果如下:
1.根据已知基因的保守序列设计并引物,采用RT-PCR和RACE相结合的方法,首次从牡丹品种‘赵粉’的萼片、花瓣、雄蕊和心皮中成功获得了7个花器官发育相关基因的cDNA全长,分别命名为PsAP1(GenBank登录号:HM143943)、PsAP2(GenBank登录号: HQ222889)、PsPI(GenBank登录号: HQ878444)、PsMADS1(GenBank登录号: HQ222890)、PsMADS9(GenBank登录号: HQ902183)、PsAG(GenBank登录号: HQ222891)和PsMADS5(GenBank登录号: HQ449569)。序列分析表明,除了PsAP2外,其余6个基因均含有MADS结构域和植物所特有的K结构域,属于MADS-box基因家族。
2.采用相对荧光定量的方法对7个基因在牡丹不同器官中的表达差异进行分析发现,不同的基因在不同的器官中有不同的表达模式,具有一定的组织特异性。其中PsAP1主要在花瓣、萼片和心皮中表达,根和茎中几乎没有表达。PsAP2在各个器官中均有表达,在花瓣中最高。PsPI主要在花瓣和雄蕊中表达,其他器官几乎没有表达。PsMADS1和PsMADS1主要在花瓣、雄蕊和心皮中表达,其他器官几乎没有表达。PsAG主要在雄蕊和心皮中表达,根和萼片中几乎没有表达。PsMADS5在各个器官中均有表达,心皮中表达量最高。
采用相对荧光定量的方法对这7个基因在牡丹花芽分化过程中的表达差异进行分析,结果表明:不同的基因在花芽分化过程中有不同的变化趋势。其中PsAP1在花芽分化前期表达最高,后期表达最低。PsAP2在花芽分化中期表达最高,初期表达最低。PsPI、PsMADS1和PsMADS5在花芽分化过程中的相对表达量均为中期表达最高,初期表达最低。PsMADS9在花芽分化后期表达最高,初期表达最低。PsAG在花芽分化末期表达最高,初期表达最低。
3.成功构建了PsAP1、PsAP2、PsPI、PsMADS1、PsMADS9、PsAG和PsMADS5的正义和反义植物表达载体,采用花序侵染法和叶盘转化法转化拟南芥和烟草,经基因组PCR检测,初步得到部分阳性植株。
4.将PsAP1基因在拟南芥和烟草中超表达,通过对转基因植株表型观察发现,转基因拟南芥和烟草都能够提前开花。初步表明PsAP1基因可能对花期调控具有重要的作用。将PsAG基因在拟南芥中超表达,转基因拟南芥植株与野生型相比,叶片变小,花发育不正常,甚至出现畸形。将PsAG基因在烟草中超表达,转基因烟草,与野生型相比,花型变小,推测PsAG基因可能参与花发育。
Tree peony (Paeonia suffruticosa) is a very popular traditional famous flower and candiate national flower .It has high ornamental value. Molecular breeding has been one of the hotspots in flower type modification. Till now, although many genes and cDNA clones for genes about floral development have been isolated and well characterized, little information, if any, is available concerning molecular aspect of floral organ development in tree peony.
In this study, we firstly isolated the full length cDNA clones encoding PsAP1, PsAP2, PsPI, PsMADS1, PsMADS5, PsAG and PsMADS5 from sepals, petals, stamens and carpels of tree peony cultivar‘Zhao Fen’using RT-PCR and RACE, then, studied their expression patterns in different organs and stages using relative real-time PCR. Meanwhile, in order to validate the function of genes, the sense and antisense plant expression vectors of the seven genes were constructed and transformed into tobacco NC89, respectively. The main results were as follows:
1. Using degenerated primers derived from conserved sequences of previously cloned genes, 7 genes involved in floral organ development were first obtained from sepals, petals, stamens and carpels of tree peony cultivar‘Zhao Fen’by RT-PCR and RACE, and were named PsAP1 (GenBank No.HM14393), PsAP2 (GenBank No. HQ222889), PsPI(GenBank No.HQ878444),PsMADS1(GenBank No.HQ222890),PsMADS9 (GenBank No.HQ902183), PsAG(GenBank No.HQ222891) and PsMADS5(GenBank No.HQ449569), respectively. Sequence analysis showed that these genes all had MADS domains and the plant specific K domains,belonged to MADS-box gene family except PsAP2.
2. The expressions of these seven genes were detected in different organs of tree peony by using relative real-time PCR. The results showed that different genes had different expression patterns and tissue specicity. PsAP1 mainly expressed in petals,sepals and carpels,but hardly expressed in roots and stems. PsAP2 expressed in all the organs,hinghest in petals. PsPI mainly expressed in petals and stamens, hardly expressed in any other organs. PsMADS1 and PsAMDS9 mainly expressed in petals stamens and carprls, hardly expressed in any other organs. PsAG mainly expressed in stamens and carpels, hardly expressed in roots and sepals. PsMADS5 expressed in all the organs, hinghest in petals.
The expressions of these seven genes were detected in different stages of tree peony by using relative real-time PCR. The results indicated that different genes had different expression patterns. PsAP1 were expressioned at the highest level at prophase and the lowest level at anaphase in the process of flower bud differentiation. PsAP2 were expressioned at the highest level at prophase and the lowest level at the initial stage in the process of flower bud differentiation. PsPI PsMADS1 and PsMADS5 were all expressioned at the highest level at metaphase and the lowest level at the initial stage in the process of flower bud differentiation. PsMADS9 were expressioned at the highest level at anaphase and the lowest level at the initial stage in the process of flower bud differentiation. PsAG were expressioned at the highest level at terminal stage and the lowest level at the initial stage in the process of flower bud differentiation.
3 The sense and antisense plant expression vectors of these seven genes(PsAP1,PsAP2, PsPI,PsMADS1,PsMADS9,PsAG,PsMADS5) were constructed and transformed into Arabidopsis thaliana and Nicotiana tabacum via Agrobacterium tumefaciens-mediated flower-dipping and leaf disk transformation method, respectively. Parts of transformed plants were obtained after genomic PCR detection.
4. Over expression of PsAP1 in transgenic Arabidopsis thaliana and Nicotiana tabacum both resulted early flowering. It indicated preliminarily that PsAP1 plays an essential role in the regulation and control of flowering time.
Over expression of PsAG in transgenic Arabidopsis thaliana, the transgenic plants appeared smaller leaves and abnormal flowers compared with wild type Arabidopsis thaliana. Over expression of PsAG in transgenic Nicotiana tabacum, the transgenic plants showed smaller flowers compared with wild type Nicotiana tabacum. It preliminarily indicated that PsAG may be related to flower development.
本研究以牡丹品种‘赵粉’为试材,通过同源克隆的办法,采用RT-PCR和RACE技术,分别从其萼片、花瓣、雄蕊和心皮中成功分离了牡丹PsAP1、PsAP2、PsPI、PsMADS1、PsMADS9、PsAG和PsMADS5这7个花器官形成相关基因的cDNA全长,并通过相对荧光定量PCR的方法,对这些基因在不同器官及花芽分化过程中的时空表达模式进行了分析。同时,分别构建了这7个基因的正义和反义植物表达载体,转化拟南芥和烟草,经过RT-PCR后,得到部分阳性植株,并对其表型初步进行初步观察。主要研究结果如下:
1.根据已知基因的保守序列设计并引物,采用RT-PCR和RACE相结合的方法,首次从牡丹品种‘赵粉’的萼片、花瓣、雄蕊和心皮中成功获得了7个花器官发育相关基因的cDNA全长,分别命名为PsAP1(GenBank登录号:HM143943)、PsAP2(GenBank登录号: HQ222889)、PsPI(GenBank登录号: HQ878444)、PsMADS1(GenBank登录号: HQ222890)、PsMADS9(GenBank登录号: HQ902183)、PsAG(GenBank登录号: HQ222891)和PsMADS5(GenBank登录号: HQ449569)。序列分析表明,除了PsAP2外,其余6个基因均含有MADS结构域和植物所特有的K结构域,属于MADS-box基因家族。
2.采用相对荧光定量的方法对7个基因在牡丹不同器官中的表达差异进行分析发现,不同的基因在不同的器官中有不同的表达模式,具有一定的组织特异性。其中PsAP1主要在花瓣、萼片和心皮中表达,根和茎中几乎没有表达。PsAP2在各个器官中均有表达,在花瓣中最高。PsPI主要在花瓣和雄蕊中表达,其他器官几乎没有表达。PsMADS1和PsMADS1主要在花瓣、雄蕊和心皮中表达,其他器官几乎没有表达。PsAG主要在雄蕊和心皮中表达,根和萼片中几乎没有表达。PsMADS5在各个器官中均有表达,心皮中表达量最高。
采用相对荧光定量的方法对这7个基因在牡丹花芽分化过程中的表达差异进行分析,结果表明:不同的基因在花芽分化过程中有不同的变化趋势。其中PsAP1在花芽分化前期表达最高,后期表达最低。PsAP2在花芽分化中期表达最高,初期表达最低。PsPI、PsMADS1和PsMADS5在花芽分化过程中的相对表达量均为中期表达最高,初期表达最低。PsMADS9在花芽分化后期表达最高,初期表达最低。PsAG在花芽分化末期表达最高,初期表达最低。
3.成功构建了PsAP1、PsAP2、PsPI、PsMADS1、PsMADS9、PsAG和PsMADS5的正义和反义植物表达载体,采用花序侵染法和叶盘转化法转化拟南芥和烟草,经基因组PCR检测,初步得到部分阳性植株。
4.将PsAP1基因在拟南芥和烟草中超表达,通过对转基因植株表型观察发现,转基因拟南芥和烟草都能够提前开花。初步表明PsAP1基因可能对花期调控具有重要的作用。将PsAG基因在拟南芥中超表达,转基因拟南芥植株与野生型相比,叶片变小,花发育不正常,甚至出现畸形。将PsAG基因在烟草中超表达,转基因烟草,与野生型相比,花型变小,推测PsAG基因可能参与花发育。
Tree peony (Paeonia suffruticosa) is a very popular traditional famous flower and candiate national flower .It has high ornamental value. Molecular breeding has been one of the hotspots in flower type modification. Till now, although many genes and cDNA clones for genes about floral development have been isolated and well characterized, little information, if any, is available concerning molecular aspect of floral organ development in tree peony.
In this study, we firstly isolated the full length cDNA clones encoding PsAP1, PsAP2, PsPI, PsMADS1, PsMADS5, PsAG and PsMADS5 from sepals, petals, stamens and carpels of tree peony cultivar‘Zhao Fen’using RT-PCR and RACE, then, studied their expression patterns in different organs and stages using relative real-time PCR. Meanwhile, in order to validate the function of genes, the sense and antisense plant expression vectors of the seven genes were constructed and transformed into tobacco NC89, respectively. The main results were as follows:
1. Using degenerated primers derived from conserved sequences of previously cloned genes, 7 genes involved in floral organ development were first obtained from sepals, petals, stamens and carpels of tree peony cultivar‘Zhao Fen’by RT-PCR and RACE, and were named PsAP1 (GenBank No.HM14393), PsAP2 (GenBank No. HQ222889), PsPI(GenBank No.HQ878444),PsMADS1(GenBank No.HQ222890),PsMADS9 (GenBank No.HQ902183), PsAG(GenBank No.HQ222891) and PsMADS5(GenBank No.HQ449569), respectively. Sequence analysis showed that these genes all had MADS domains and the plant specific K domains,belonged to MADS-box gene family except PsAP2.
2. The expressions of these seven genes were detected in different organs of tree peony by using relative real-time PCR. The results showed that different genes had different expression patterns and tissue specicity. PsAP1 mainly expressed in petals,sepals and carpels,but hardly expressed in roots and stems. PsAP2 expressed in all the organs,hinghest in petals. PsPI mainly expressed in petals and stamens, hardly expressed in any other organs. PsMADS1 and PsAMDS9 mainly expressed in petals stamens and carprls, hardly expressed in any other organs. PsAG mainly expressed in stamens and carpels, hardly expressed in roots and sepals. PsMADS5 expressed in all the organs, hinghest in petals.
The expressions of these seven genes were detected in different stages of tree peony by using relative real-time PCR. The results indicated that different genes had different expression patterns. PsAP1 were expressioned at the highest level at prophase and the lowest level at anaphase in the process of flower bud differentiation. PsAP2 were expressioned at the highest level at prophase and the lowest level at the initial stage in the process of flower bud differentiation. PsPI PsMADS1 and PsMADS5 were all expressioned at the highest level at metaphase and the lowest level at the initial stage in the process of flower bud differentiation. PsMADS9 were expressioned at the highest level at anaphase and the lowest level at the initial stage in the process of flower bud differentiation. PsAG were expressioned at the highest level at terminal stage and the lowest level at the initial stage in the process of flower bud differentiation.
3 The sense and antisense plant expression vectors of these seven genes(PsAP1,PsAP2, PsPI,PsMADS1,PsMADS9,PsAG,PsMADS5) were constructed and transformed into Arabidopsis thaliana and Nicotiana tabacum via Agrobacterium tumefaciens-mediated flower-dipping and leaf disk transformation method, respectively. Parts of transformed plants were obtained after genomic PCR detection.
4. Over expression of PsAP1 in transgenic Arabidopsis thaliana and Nicotiana tabacum both resulted early flowering. It indicated preliminarily that PsAP1 plays an essential role in the regulation and control of flowering time.
Over expression of PsAG in transgenic Arabidopsis thaliana, the transgenic plants appeared smaller leaves and abnormal flowers compared with wild type Arabidopsis thaliana. Over expression of PsAG in transgenic Nicotiana tabacum, the transgenic plants showed smaller flowers compared with wild type Nicotiana tabacum. It preliminarily indicated that PsAG may be related to flower development.
引文
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