毛竹细胞骨架微管蛋基因PeTua3和PeTub3研究
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摘要
竹林是我国重要的森林资源，竹材是由各种类型细胞的细胞壁及其包围的空间所组成的复杂生物材料，因其具有强度高、韧性好、硬度大等特点而在建筑、造纸等领域得到广泛的利用。竹子优良的材性特点与其细胞壁的组成及结构密切相关，竹子细胞壁的发育对竹子的生长发育、生理特性、生化代谢以及对外界环境胁迫的有机反应发挥着重要的调节作用，对竹子材性的形成起着决定性作用。细胞骨架指导细胞壁的生物合成，作为细胞骨架的重要组成部分，微管蛋白在细胞壁发育过程中起着重要的调控作用。本研究以我国重要经济竹种--毛竹（Phyllostachys edulis）为材料，分离细胞骨架微管蛋白基因，在对其序列进行充分分析的基础上，通过体外表达和在模式植物中的表达来验证其功能，以其为通过基因工程手段培育竹类植物新品种奠定基础。主要研究结果如下：
1、采用RT-PCR方法从毛竹叶片中克隆了2个微管蛋白基因，分别命名为PeTua3和PeTub3。PeTua3的编码区为1353bp，编码451个氨基酸；PeTub3的编码区1341bp，编码447个氨基酸。
2、Blast分析表明PeTua3和PeTub3与已报道的其它植物的tubulin蛋白具有很高的相似性，分别属于tubulin蛋白的Tua3和Tub3家族，都是疏水性酸性蛋白。通过MOTIFSCAN软件预测蛋白质的功能性位点，结果显示PeTua3和PeTub3序列均包含有N-糖基化位点、磷酸化酪蛋白激酶II位点、N-肉豆蔻酰化位点和磷酸化蛋白激酶位点、GTP酶域和C末端结构域，但是这些位点在蛋白序列中的位置、数量均存在一定的差异，这意味着它们可能存在功能上的不同。应用MEGA4.0软件，构建基于tubulin蛋白的系统进化树，结果表明：PeTua3与水稻的Tua3蛋白在进化关系上最近，Petub3与牛筋草的Tub3蛋白在进化关系上最近，而与大部分双子叶植物Tubulin蛋白的关系较远，这与毛竹属于单子植物禾本科竹亚科的形态学分类相吻合。
3、通过构建PeTua3和PeTub3基因的原核表达载体pET-32b-PeTua3和pET-30a-PeTub3，分别转化大肠杆菌(Escherichia coli)菌株DE3，进行基因的体外表达研究。经过优化诱导表达条件，在37℃，0.4mmol·L~(-1)IPTG诱导2h的条件下获得高丰度的PeTua3可溶性重组蛋白；在28℃，0.4mmol·L~(-1)IPTG诱导4h的条件下获得高丰度的PeTub3可溶性重组蛋白。应用His标签亲和纯化方法，获得PeTua3、PeTub3的重组蛋白，并处理拟南芥种子，幼苗表现为上胚轴明显增粗，侧根增多，这意味着重组蛋白可能具有加快细胞的分裂的生物学活性。
4、构建PeTua3和PeTub3基因的正义和反义植物表达载体，并转化拟南芥，通过抗性筛选获得转基因植株。表型观察发现，转基因拟南芥（正义和反义）和野生拟南芥在发芽和茎的生长上没有因基因的过量表达或抑制而受到影响，但对根系生长影响明显。PeTua3和PeTub3基因的过量表达均促进拟南芥根系生长，侧根增多；石蜡切片显示根系表皮薄壁细胞数量增多，且细胞体积变大，维管柱的体积增大，约是野生拟南芥的2倍，木质部明显加厚。而转反义PeTua3和PeTub3基因的拟南芥根系生长不正常，产生大量须根；石蜡切片显示根系表皮薄壁细胞的数量明显减少，且细胞体积变小，维管柱的体积变小，木质部明显比较薄。
5、在烟草悬浮细胞（BY-2）中过量表达PeTua3和PeTub3，与对照相比转基因BY-2的G2期和S期分裂时间明显缩短，能显著提高烟草细胞的有丝分裂指数，细胞分裂加快。
细胞骨架微管蛋白基因PeTua3和PeTub3的体外表达蛋白能够促进细胞生长，具有生物学活性，这在转PeTua3和PeTub3基因拟南芥中进一步得到证实，对转PeTua3和PeTub3基因烟草悬浮细胞的观察证实，PeTua3和PeTub3基因促进生长是通过缩短细胞分裂周期实现的。由此证明PeTua3和PeTub3基因具有促进细胞发育的功能，这意味着PeTua3和PeTub3基因对于分子育种研究具有重要的潜在应用价值。
Bamboo forest is one of important forest resources. Bamboo timber is a complexbiological material consisted of various types of cell walls forming special spatial structurewhich has high strength, good toughness and hardness, and it is widely used in handcraft,construction, paper making, etc. The excellent material characteristics of bamboo is closelyrelated to the composition and structure of its cell wall. The development of cell wall playsimportant roles in the growth and development, physiological characteristics, biochemicalmetabolism of bamboo and the defence reactions to environmental stress, and it also decidesthe characteristics of bamboo material. The biosynthesis of cell wall is guided by cytoskeletonwhich is composed of microfilament, microtubule and intemediate filament. As a key part ofcytoskeleton, microtubule has an important regulatory role in the development of cell wall. Inthis study, two genes encoding microtubule proteins were isolated from Moso bamboo(Phyllostachys edulis) and their functions were identified through prokaryotic expression invitro and ectopic expression in model plants. This work will lay the basis for molecularengineering to breed new bamboo varieties. The main results are as follows:
1. Two genes encoding microtubule proteins were cloned using RT-PCR from leaves ofMoso bamboo, and named as PeTua3and PeTub3respectively. The open reading frame (ORF)of PeTua3is1353bp encoding451amino acids, while the ORF of PeTub3is1341bpencoding447amino acids.
2. Blast analysis showed that PeTua3and PeTub3had a high similarity with tubilins fromother plants and belonged to α-tubilin and β-tubilin family, respectively. Hydropathic analysisshowed that PeTua3and PeTub3were hydrophobic acidic protein. The protein analysis withMOTIFSCAN showed that Petua3and PeTub3all contained N-glycosylation sites, caseinkinase II phosphorylation sites, N-myristoylation sites, phosphorylation protein kinase sites,GTPase domain and C-terminal domain, however, the positions and quantity of these sites were different in the sequences, which means they might have different functions. Phylogenetic treesbased on tubilins were constructed by using MEGA4.0. The reslut showed that PeTua3andPetub3were closed to α-tubilins and β-tubilins from monocotyledon respectively and wereaway from to those of dicotyledon, the nearest one to PeTua3was that of rice and to PeTub3was that of goose grass, which was consistent with the morphological classification ofBambusoideae in Poaceae of monocotyledon plants.
3. The prokaryotic expression vectors of PeTua3and Petub3were constructed and theplasmids of pET-32b-PeTua3and pET-30a-PeTub3were transformed into Escherichia coli(DE3). The positive clones were selected for further experiments. By optimizing the expressionconditions of induction, the soluble recombinant protein of PeTua3was effectively expressedat37℃induced by0.4mmol·L~(-1)IPTG with2h, while that of PeTub3received higherexpression at28℃induced by0.4mmol·L~(-1)IPTG with4h. The recombinant proteins werepurified by using His·Tag Bind Purification Kit and were used to treat seeds of Arabidopsisthaliana. The result showed that the hypocotyl of the treated seedlings were thicken and thequantity of lateral root increased significantly, which indicated that the recombinant proteinsmight have biological activity for growth stimulation.
4. The plant expression vectors of sense/antisense PeTua3/PeTub3were constructed andtransferred into A. thaliana. The sense and antisense transgenic plants were selected withantibiotics and the postive transgenics of T3were used to further studies. The phenotypes ofthe transgenics were observed, the germination was not affected by the overexpression orinhibition of genes. However, the root growth was affected obviously. The root phenotypes ofsense transgenics all grew fast and had more lateral roots, while the antisense transgenics allgrew abnomal and had lots of hair roots. All the roots of sense and antisense transgenics wereembedded in paraffin and sectioned to8~(-1)0m thick for tissue sections making. The roottissue sections of sense transgenics showed that the quantity of epidermal parenchyma cellswas increasing, the volume of the cells was becoming larger, the xylem was significantlythicker and the volume of the vascular cylinder increased about2times than that of wild type. However the quantity of epidermal parenchyma cells in antisense transgenics was significantlyreduced, the volume of the cells became smaller, the xyle was obviously thinner and thevolume of the vascular cylinder became smaller than that of wild type. These results suggestedthat PeTua3and PeTub3could promote cell growth and division to accelerate the growth ofroots.
5. Tobacco (Nicotiana tabacum) BY-2suspension cells were used to study the tubilingenes’ influence on cell division. PeTua3and PeTub3were overexpressed respectively in BY-2suspension cells. The result of microscope observation showed that the mitotic index oftransgenic BY-2cells was increased and the shape of the cells was also changed. Comparedwith BY-2cells, the time of G2and S phase for the transgenic BY-2cells was shortenedsignificantly, which accelerated the cell division.
The recombinant proteins of expression in vitro have biological activity for promoting cellgrowth, which further confirmed in transgenic A. thaliana. PeTua3and PeTub3promotegrowth by reducing the cell division cycle through transgenic BY-2. Thus prove the PeTua3and PeTub3functions to promote cell development has been proved, which means that PeTua3and PeTub3for molecular breeding have important potential applications.
1、采用RT-PCR方法从毛竹叶片中克隆了2个微管蛋白基因，分别命名为PeTua3和PeTub3。PeTua3的编码区为1353bp，编码451个氨基酸；PeTub3的编码区1341bp，编码447个氨基酸。
2、Blast分析表明PeTua3和PeTub3与已报道的其它植物的tubulin蛋白具有很高的相似性，分别属于tubulin蛋白的Tua3和Tub3家族，都是疏水性酸性蛋白。通过MOTIFSCAN软件预测蛋白质的功能性位点，结果显示PeTua3和PeTub3序列均包含有N-糖基化位点、磷酸化酪蛋白激酶II位点、N-肉豆蔻酰化位点和磷酸化蛋白激酶位点、GTP酶域和C末端结构域，但是这些位点在蛋白序列中的位置、数量均存在一定的差异，这意味着它们可能存在功能上的不同。应用MEGA4.0软件，构建基于tubulin蛋白的系统进化树，结果表明：PeTua3与水稻的Tua3蛋白在进化关系上最近，Petub3与牛筋草的Tub3蛋白在进化关系上最近，而与大部分双子叶植物Tubulin蛋白的关系较远，这与毛竹属于单子植物禾本科竹亚科的形态学分类相吻合。
3、通过构建PeTua3和PeTub3基因的原核表达载体pET-32b-PeTua3和pET-30a-PeTub3，分别转化大肠杆菌(Escherichia coli)菌株DE3，进行基因的体外表达研究。经过优化诱导表达条件，在37℃，0.4mmol·L~(-1)IPTG诱导2h的条件下获得高丰度的PeTua3可溶性重组蛋白；在28℃，0.4mmol·L~(-1)IPTG诱导4h的条件下获得高丰度的PeTub3可溶性重组蛋白。应用His标签亲和纯化方法，获得PeTua3、PeTub3的重组蛋白，并处理拟南芥种子，幼苗表现为上胚轴明显增粗，侧根增多，这意味着重组蛋白可能具有加快细胞的分裂的生物学活性。
4、构建PeTua3和PeTub3基因的正义和反义植物表达载体，并转化拟南芥，通过抗性筛选获得转基因植株。表型观察发现，转基因拟南芥（正义和反义）和野生拟南芥在发芽和茎的生长上没有因基因的过量表达或抑制而受到影响，但对根系生长影响明显。PeTua3和PeTub3基因的过量表达均促进拟南芥根系生长，侧根增多；石蜡切片显示根系表皮薄壁细胞数量增多，且细胞体积变大，维管柱的体积增大，约是野生拟南芥的2倍，木质部明显加厚。而转反义PeTua3和PeTub3基因的拟南芥根系生长不正常，产生大量须根；石蜡切片显示根系表皮薄壁细胞的数量明显减少，且细胞体积变小，维管柱的体积变小，木质部明显比较薄。
5、在烟草悬浮细胞（BY-2）中过量表达PeTua3和PeTub3，与对照相比转基因BY-2的G2期和S期分裂时间明显缩短，能显著提高烟草细胞的有丝分裂指数，细胞分裂加快。
细胞骨架微管蛋白基因PeTua3和PeTub3的体外表达蛋白能够促进细胞生长，具有生物学活性，这在转PeTua3和PeTub3基因拟南芥中进一步得到证实，对转PeTua3和PeTub3基因烟草悬浮细胞的观察证实，PeTua3和PeTub3基因促进生长是通过缩短细胞分裂周期实现的。由此证明PeTua3和PeTub3基因具有促进细胞发育的功能，这意味着PeTua3和PeTub3基因对于分子育种研究具有重要的潜在应用价值。
Bamboo forest is one of important forest resources. Bamboo timber is a complexbiological material consisted of various types of cell walls forming special spatial structurewhich has high strength, good toughness and hardness, and it is widely used in handcraft,construction, paper making, etc. The excellent material characteristics of bamboo is closelyrelated to the composition and structure of its cell wall. The development of cell wall playsimportant roles in the growth and development, physiological characteristics, biochemicalmetabolism of bamboo and the defence reactions to environmental stress, and it also decidesthe characteristics of bamboo material. The biosynthesis of cell wall is guided by cytoskeletonwhich is composed of microfilament, microtubule and intemediate filament. As a key part ofcytoskeleton, microtubule has an important regulatory role in the development of cell wall. Inthis study, two genes encoding microtubule proteins were isolated from Moso bamboo(Phyllostachys edulis) and their functions were identified through prokaryotic expression invitro and ectopic expression in model plants. This work will lay the basis for molecularengineering to breed new bamboo varieties. The main results are as follows:
1. Two genes encoding microtubule proteins were cloned using RT-PCR from leaves ofMoso bamboo, and named as PeTua3and PeTub3respectively. The open reading frame (ORF)of PeTua3is1353bp encoding451amino acids, while the ORF of PeTub3is1341bpencoding447amino acids.
2. Blast analysis showed that PeTua3and PeTub3had a high similarity with tubilins fromother plants and belonged to α-tubilin and β-tubilin family, respectively. Hydropathic analysisshowed that PeTua3and PeTub3were hydrophobic acidic protein. The protein analysis withMOTIFSCAN showed that Petua3and PeTub3all contained N-glycosylation sites, caseinkinase II phosphorylation sites, N-myristoylation sites, phosphorylation protein kinase sites,GTPase domain and C-terminal domain, however, the positions and quantity of these sites were different in the sequences, which means they might have different functions. Phylogenetic treesbased on tubilins were constructed by using MEGA4.0. The reslut showed that PeTua3andPetub3were closed to α-tubilins and β-tubilins from monocotyledon respectively and wereaway from to those of dicotyledon, the nearest one to PeTua3was that of rice and to PeTub3was that of goose grass, which was consistent with the morphological classification ofBambusoideae in Poaceae of monocotyledon plants.
3. The prokaryotic expression vectors of PeTua3and Petub3were constructed and theplasmids of pET-32b-PeTua3and pET-30a-PeTub3were transformed into Escherichia coli(DE3). The positive clones were selected for further experiments. By optimizing the expressionconditions of induction, the soluble recombinant protein of PeTua3was effectively expressedat37℃induced by0.4mmol·L~(-1)IPTG with2h, while that of PeTub3received higherexpression at28℃induced by0.4mmol·L~(-1)IPTG with4h. The recombinant proteins werepurified by using His·Tag Bind Purification Kit and were used to treat seeds of Arabidopsisthaliana. The result showed that the hypocotyl of the treated seedlings were thicken and thequantity of lateral root increased significantly, which indicated that the recombinant proteinsmight have biological activity for growth stimulation.
4. The plant expression vectors of sense/antisense PeTua3/PeTub3were constructed andtransferred into A. thaliana. The sense and antisense transgenic plants were selected withantibiotics and the postive transgenics of T3were used to further studies. The phenotypes ofthe transgenics were observed, the germination was not affected by the overexpression orinhibition of genes. However, the root growth was affected obviously. The root phenotypes ofsense transgenics all grew fast and had more lateral roots, while the antisense transgenics allgrew abnomal and had lots of hair roots. All the roots of sense and antisense transgenics wereembedded in paraffin and sectioned to8~(-1)0m thick for tissue sections making. The roottissue sections of sense transgenics showed that the quantity of epidermal parenchyma cellswas increasing, the volume of the cells was becoming larger, the xylem was significantlythicker and the volume of the vascular cylinder increased about2times than that of wild type. However the quantity of epidermal parenchyma cells in antisense transgenics was significantlyreduced, the volume of the cells became smaller, the xyle was obviously thinner and thevolume of the vascular cylinder became smaller than that of wild type. These results suggestedthat PeTua3and PeTub3could promote cell growth and division to accelerate the growth ofroots.
5. Tobacco (Nicotiana tabacum) BY-2suspension cells were used to study the tubilingenes’ influence on cell division. PeTua3and PeTub3were overexpressed respectively in BY-2suspension cells. The result of microscope observation showed that the mitotic index oftransgenic BY-2cells was increased and the shape of the cells was also changed. Comparedwith BY-2cells, the time of G2and S phase for the transgenic BY-2cells was shortenedsignificantly, which accelerated the cell division.
The recombinant proteins of expression in vitro have biological activity for promoting cellgrowth, which further confirmed in transgenic A. thaliana. PeTua3and PeTub3promotegrowth by reducing the cell division cycle through transgenic BY-2. Thus prove the PeTua3and PeTub3functions to promote cell development has been proved, which means that PeTua3and PeTub3for molecular breeding have important potential applications.
引文
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Thitamadee S，Tuchihara K．Hashimoto T．Microtubule basis for left-handed helical growth in Arabidopsis.Nature,2002,417:193-196.
Ueda K,Matsuyama T,Hashimoto T.Visualization of microtubules in living cells of transgenic Arabidopsisthaliana. Protoplasma,1999,206:201-206.
Uribe X, Torres MA, Capellades M．Maize α-tubulin genes are expressed according to specific pattern of celldifferentiation．Plant Mol Biol,1998,37:1069-1078.
Vantard M, Blanchon L. Actin polymerization processes in plant cells. Curr Opinin Plant Biol,2002,5:502-506.
Vidali L, Mckenna S T. Actin polymerization is essential for pollen tube growth. Molecular Biology of theCell,2001,12:2534-2545.
Wasteneys G O，Galway M E．Remodeling the cytoskeleton for growth and form: an overview with somenew views. Annual Rev Bio1,2003,54:691-722.
Yamamoto K, Kiss J Z. Disruption of the actin cytoskeleton results in the promotion of gravitropismininflorescence stems and hypocotyls of Arabidopsis. Plant Physiology,2002,128:669-681.
Yoshimura T, Demura T, Igarashi M.,et al.. Differential expression of three genes for different alpha tubulinisotypes during the initial culture of Zinnia mesophyll cells that divide and differentiate into trachearyelements. Plant Cell Physiol,1996,37:1167-1176.
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