结球甘蓝BoTCTP基因表达载体的构建及转化芥蓝的研究
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摘要
芥蓝(Brassica alboglabra)是十字花科重要蔬菜之一,在栽培过程中常常会遭受各种逆境的严重影响,给芥蓝的生产造成很大的损失。为提高芥蓝对逆境胁迫的抵抗能力,本研究根据结球甘蓝翻译控制肿瘤蛋白基因(translationally controlled tumor protein cloned from Brassica oleracea, BoTCTP)序列设计特异引物,经PCR扩增得到目的片段,然后构建过表达载体,利用农杆菌(Agrobacterium tumefaciens)介导法导入芥蓝中,并对其转基因植株进行了分子生物学鉴定。研究结果显示,4株外源目的基因已经整合到受体基因组中,1株以双拷贝形式存在,另外3株以单拷贝形式存在。转基因植株的长势旺盛,生长速率加快,叶面积较大,在逆境胁迫(高低温,盐碱,干旱,重金属等)下转基因植株对胁迫的耐受性明显优于对照植株,从而表明BoTCTP基因与芥蓝的生长密切相关,并对芥蓝的抗逆性起调控作用。本研究结果为芥蓝抗逆性的进一步研究提供了科学依据。
Chinese kale(Brassica alboglabra) is one of the important vegetables in the Cruciferous family. It is often subjected to various adverse effects during the cultivation process, causing great losses to the production. In order to improve the resistance of Chinese kale to stress, the specific primers were designed according to BoTCTP(translationally controlled tumor protein cloned from Brassica oleracea) in this research.The interest fragment was amplified by PCR, and then the overexpression vector was constructed and transferred into Chinese kale by Agrobacterium tumefaciens mediated. The transformants were confirmed by molecular methods and biological assay. Four transgenic plants were obtained, and then taken to design the experiments associated with growth and different stress, such as high temperature, low temperature, salt,heavy metal and so on. The results showed that the transgenic plants grew faster, stronger and had biggerleaves than non-transgenic plants. In the different stress condition, the transgenic plants had obviously stronger resistance than the control, which showed that the BoTCTP was closely related to the growth and had a regulatory effect on the stress resistance of Chinese kale. These results provide some scientific basis for the future research on the stress resistance of Chinese kale.翻译控制肿瘤蛋白(translationally controlled tu-桑树(Morus atropurpurea)(雷朋等, 2015)、水稻(Orymor protein, TCTP)广泛存在于动物、植物及酵母中za sativa)(王步勇等, 2016)、刺槐(Robinia pseudoaca-(Yenofsky et al., 1983, Sturzenbaum et al., 1998),是cia)(Chou et al., 2016)、黄 瓜(Cucumis sativus)一类在序列上高度保守且具有很高同源性的蛋白(Meng et al.,2017)、番 茄(Solanum lycopersicum)家族.TCTP 具有很多重要的生物学功能,除了调(Márcio et al., 2017;Fernanda et al., 2017)、小 麦节细胞周期、钙离子结合、微管结合、组胺释放、炎(Triticum aestivum)(麻楠等, 2018)等植物也先后开症反应 、抗凋亡 、抗疟疾等作用(Bommer et al.,展了 TCTP 基因的克隆与表达分析研究.此外,2004; 吕素芳等, 2006; Lucibello et al., 2011),还具AtTCTP的过表达不仅增强了拟南芥对干旱的忍受有参与细胞增殖与分化、细胞骨架重排(Bazile et力,而且在脱落酸(abscisic acid, ABA)介导的气孔运al.,2009; Tsarova et al., 2010; 陈科等, 2012)以及促动中发挥作用(Kim et al., 2012),刺槐 TCTP(简称进受精能力的提高(Jeon et al., 2017)等功能.医学Rpf41)的RNAi干涉载体的导入使其根茎长、鲜重和上,TCTP基因在肿瘤发生与逆转中起关键作用,保结瘤数均减少(Chou et al., 2016),TCTP还参与植物护癌细胞免受外部应激诱导,是癌症治疗的潜在靶的抗虫性(Zhang et al., 2014)和抗病性(Meng et al.,标(Li et al., 2016;Du et al., 2017;Wang et al., 2018).2018)调控.TCTP 的表达水平存在高度差异,这些表达差异主芥蓝(Brassica alboglabra)是十字花科重要蔬菜要取决于细胞或组织的类型、发育阶段、信号反应、之一,风味别致,营养丰富,其中富含的萝卜硫苷具生长信号和细胞因子(Wang et al., 2009).植物有很强的抗癌功效(司雨等, 2009; 孙勃等, 2011;TCTP 可能具备动物 TCTP 相似的功能(曹必好等,Qian et al., 2015).为了提高芥蓝在栽培中对各种2006),不仅能够调控植株的营养生长与生殖生长逆境的抵抗能力,本研究从分子水平研究芥蓝的抗(Hsu et al., 2007; Oliver et al., 2008; Brioudes et al.,逆性.关于芥蓝生理生化指标(温度, 水分, 光照等)2010),同时对逆境胁迫(如温度, 光照, 盐碱和重金的研究报道较多(杨云英, 杨暹, 2002; Issarakraisila属等)的反应也很敏感(林莎等, 2008, Kim et al.,et al., 2007; 杨云英, 杨暹, 2008; Chen et al., 2008;2012; Deng et al., 2016).植物 TCTP 基因的表达同Deng et al., 2017),但其抗逆相关基因的克隆与功能样存在组织特异性,而且在不同部位的表达强度也鉴定研究较少.Zhu等(2016)将拟南芥抗逆基因At-因 植 物 的 不 同 而 可 能 有 所 不 同(Lopez, Franco,EDT1/HDG11导入芥蓝,发现转基因植株的抗旱性2006; 林莎等, 2008; 梁小莲等, 2009).植物 TCTP和渗透胁迫耐受性明显提高.曹必好等(2002)构建在拟南芥(Arabidopsis thaliana)上研究最多,主要包了1个结球甘蓝(Brassica oleracea) cDNA文库,随后括 AtTCTP1 和 AtTCTP2 两条序列,AtTCTP1 在调节在筛选抗病基因时得到1个阳性克隆BoTCTP,该基细胞生长、控制细胞程序化死亡和信号应激反应中因可能与植物抗逆性有关(曹必好等, 2006).干涉起重要作用(Brioudes et al., 2010; Hoepflinger et al.,该基因在芥蓝中的表达,植株生长缓慢,对逆境的2013),AtTCTP2 对提高植株的生存能力和再生能力抵抗力减弱(Cao et al., 2010).目前关于该基因在是必不可少的(Toscano-Morales et al., 2016).近年芥蓝中的过表达反应还未见报道.本研究将来,学者们对山茶花(Camellia japonica)(丁岳炼等,BoTCTP基因克隆到植物表达载体pFGC5941上,利2014)、木薯(Manihot esculent)(Brigida et al., 2014)、用农杆菌(Agrobacterium tumefaciens)介导法将其导
Chinese kale(Brassica alboglabra) is one of the important vegetables in the Cruciferous family. It is often subjected to various adverse effects during the cultivation process, causing great losses to the production. In order to improve the resistance of Chinese kale to stress, the specific primers were designed according to BoTCTP(translationally controlled tumor protein cloned from Brassica oleracea) in this research.The interest fragment was amplified by PCR, and then the overexpression vector was constructed and transferred into Chinese kale by Agrobacterium tumefaciens mediated. The transformants were confirmed by molecular methods and biological assay. Four transgenic plants were obtained, and then taken to design the experiments associated with growth and different stress, such as high temperature, low temperature, salt,heavy metal and so on. The results showed that the transgenic plants grew faster, stronger and had biggerleaves than non-transgenic plants. In the different stress condition, the transgenic plants had obviously stronger resistance than the control, which showed that the BoTCTP was closely related to the growth and had a regulatory effect on the stress resistance of Chinese kale. These results provide some scientific basis for the future research on the stress resistance of Chinese kale.翻译控制肿瘤蛋白(translationally controlled tu-桑树(Morus atropurpurea)(雷朋等, 2015)、水稻(Orymor protein, TCTP)广泛存在于动物、植物及酵母中za sativa)(王步勇等, 2016)、刺槐(Robinia pseudoaca-(Yenofsky et al., 1983, Sturzenbaum et al., 1998),是cia)(Chou et al., 2016)、黄 瓜(Cucumis sativus)一类在序列上高度保守且具有很高同源性的蛋白(Meng et al.,2017)、番 茄(Solanum lycopersicum)家族.TCTP 具有很多重要的生物学功能,除了调(Márcio et al., 2017;Fernanda et al., 2017)、小 麦节细胞周期、钙离子结合、微管结合、组胺释放、炎(Triticum aestivum)(麻楠等, 2018)等植物也先后开症反应 、抗凋亡 、抗疟疾等作用(Bommer et al.,展了 TCTP 基因的克隆与表达分析研究.此外,2004; 吕素芳等, 2006; Lucibello et al., 2011),还具AtTCTP的过表达不仅增强了拟南芥对干旱的忍受有参与细胞增殖与分化、细胞骨架重排(Bazile et力,而且在脱落酸(abscisic acid, ABA)介导的气孔运al.,2009; Tsarova et al., 2010; 陈科等, 2012)以及促动中发挥作用(Kim et al., 2012),刺槐 TCTP(简称进受精能力的提高(Jeon et al., 2017)等功能.医学Rpf41)的RNAi干涉载体的导入使其根茎长、鲜重和上,TCTP基因在肿瘤发生与逆转中起关键作用,保结瘤数均减少(Chou et al., 2016),TCTP还参与植物护癌细胞免受外部应激诱导,是癌症治疗的潜在靶的抗虫性(Zhang et al., 2014)和抗病性(Meng et al.,标(Li et al., 2016;Du et al., 2017;Wang et al., 2018).2018)调控.TCTP 的表达水平存在高度差异,这些表达差异主芥蓝(Brassica alboglabra)是十字花科重要蔬菜要取决于细胞或组织的类型、发育阶段、信号反应、之一,风味别致,营养丰富,其中富含的萝卜硫苷具生长信号和细胞因子(Wang et al., 2009).植物有很强的抗癌功效(司雨等, 2009; 孙勃等, 2011;TCTP 可能具备动物 TCTP 相似的功能(曹必好等,Qian et al., 2015).为了提高芥蓝在栽培中对各种2006),不仅能够调控植株的营养生长与生殖生长逆境的抵抗能力,本研究从分子水平研究芥蓝的抗(Hsu et al., 2007; Oliver et al., 2008; Brioudes et al.,逆性.关于芥蓝生理生化指标(温度, 水分, 光照等)2010),同时对逆境胁迫(如温度, 光照, 盐碱和重金的研究报道较多(杨云英, 杨暹, 2002; Issarakraisila属等)的反应也很敏感(林莎等, 2008, Kim et al.,et al., 2007; 杨云英, 杨暹, 2008; Chen et al., 2008;2012; Deng et al., 2016).植物 TCTP 基因的表达同Deng et al., 2017),但其抗逆相关基因的克隆与功能样存在组织特异性,而且在不同部位的表达强度也鉴定研究较少.Zhu等(2016)将拟南芥抗逆基因At-因 植 物 的 不 同 而 可 能 有 所 不 同(Lopez, Franco,EDT1/HDG11导入芥蓝,发现转基因植株的抗旱性2006; 林莎等, 2008; 梁小莲等, 2009).植物 TCTP和渗透胁迫耐受性明显提高.曹必好等(2002)构建在拟南芥(Arabidopsis thaliana)上研究最多,主要包了1个结球甘蓝(Brassica oleracea) cDNA文库,随后括 AtTCTP1 和 AtTCTP2 两条序列,AtTCTP1 在调节在筛选抗病基因时得到1个阳性克隆BoTCTP,该基细胞生长、控制细胞程序化死亡和信号应激反应中因可能与植物抗逆性有关(曹必好等, 2006).干涉起重要作用(Brioudes et al., 2010; Hoepflinger et al.,该基因在芥蓝中的表达,植株生长缓慢,对逆境的2013),AtTCTP2 对提高植株的生存能力和再生能力抵抗力减弱(Cao et al., 2010).目前关于该基因在是必不可少的(Toscano-Morales et al., 2016).近年芥蓝中的过表达反应还未见报道.本研究将来,学者们对山茶花(Camellia japonica)(丁岳炼等,BoTCTP基因克隆到植物表达载体pFGC5941上,利2014)、木薯(Manihot esculent)(Brigida et al., 2014)、用农杆菌(Agrobacterium tumefaciens)介导法将其导
引文
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Ermolayev V,Weschke W,Manteuffel R.2003.Comparison of Al-induced gene expression in sensitive and tolerant soybean cultivars[J].Journal of Experimental Botany,54(393):2745-2756.
Hoepflinger M C,Reitsamer J,Geretschlaeger A M,et al.2013.The effect of translationally controlled tumour protein(TCTP)on programmed cell death in plants[J].BMC Plant Biology.13(1):135.
Hsu Y C,Chern J J,Cai Y,et al.2007.Drosophila TCTP is essential for growth and proliferation through regulation of dRheb GTPase[J].Nature,445(7129):785-788.
Issarakraisila M,Ma Q F,Turner D W.2007.Photosynthetic and growth responses of juvenile Chinese kale(Brassica oleracea var.alboglabra)and Caisin(Brassica rapa subsp parachinensis)to water logging and water deficit[J].Scientia Horticulture,111(2):107-113.
Jeon H J,Cui X S,Guo J,et al.2017.TCTP regulates spindle assembly during postovulatory aging and prevents deterioration in mouse oocyte quality[J].Elsevier Journal,1328-1334.
Kim Y M,Han Y J,Hwang O J,et al.2012.Overexpression of Arabidopsis translationally controlled tumor protein gene AtTCTP enhances drought tolerance with rapid ABA-induced stomatal closure[J].Molecules and Cells,33(6):617-626.
Li S,Chen M H,Xiong Q,et al.2016.Characterization of the translationally controlled tumor protein(TCTP)interactome reveals novel binding partners in human cancer cells[J].Journal of proteome research,15(10):3741-3751.
Lopez A P,Franco A R.2006.Cloning and expression of cD-NA encoding translationally controlled tumor protein from strawberry fruits[J].Biologia Plantarum,50(3)447-449.
Lucibello M,Gambacurta A,Zonfrillo M.2011.TCTP is a critical survival factor that protects cancer cells from oxidative stress induced cell-death[J].Experimental Cel Research,317(17):2479-2489.
Márcio de C,Márcio L A,Alessandra V N L,et al.2017.Impacts of the overexpression of a tomato translationally controlled tumor protein(TCTP)in tobacco revealed by phenotypic and transcriptomic analysis[J].Plant Cell Reports,36(6):887-900.
Meng X,Chen Q,Fan H,et al.2017.Molecular characterization,expression analysis and heterologous expression of two translationally controlled tumor protein genes from Cucumis sativus[J].PLoS One,12(9):e0184872
Meng X N,Yu Y,Zhao J Y,et al.2018.The two translationally controlled tumor protein genes,CsTCTP1 and CsT-CTP2,are negative modulators in the Cucumis sativus defense response to sphaerotheca fuliginea[J].In Plan Science,9:544.
Qian H M,Sun B,Miao H Y,et al.2015.Variation of glucosinolates and quinone reductase activity among different varieties of Chinese kale and improvement of glucoraphanin by metabolic engineering[J].Food Chemistry168:321-326.
Santa Brigida A B,Dos Reis S P,Costa C N M,et al.2014Molecular cloning and characterization of a cassava translationally controlled tumor protein gene potentially related to salt stress response[J].Molecular Biology Reports,41(3):1787-1797.
Sturzenbaum S R,Kille P,Morgan A J.1998.Identification of heavy metal induced changes in the expression patterns of the translationally controlled tumour protein(TCTP)in the earthworm Lumbricus rubellus[J].Biochimica e Biophysica Acta,1398(3):294-304.
Tsarova K,Yarmola E G,Bubb M R.2010.Identification of a cofilin-like actin-binding site on translationally controlled tumor protein(TCTP)[J].FEBS Letters,584(23)4756-4760.
Toscano-Morales R,Xoconostle-Cázares B,Martínez-Navarro A C,et al.2014.Long distance movement of an arabidopsis translationally controlled tumor protein(AtTCTP2)mRNA and protein in tobacco[J].Frontiers in Plant Science,(5):705.
Toscano-Morales R,Xoconostle-Cázares B,Cabrera-Ponce JL,et al.2015.AtTCTP2,an Arabidopsis thaliana homolog of translationally controlled tumor protein,enhances in vitro plant regeneration[J].Frontiers in Plant Science,(6):468.
Toscano-Morales R,Xoconostle-Ca′zares B,Mart?′nez-Navarro A C,et al.2016.AtTCTP2 mRNA and protein movement correlates with formation of adventitious roots in tobacco[J].Plant Signaling&Behavior,11(3):e1071003.
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Wang Z Q,Li G Z,Gong Q Q,et al.2015.OsTCTP,encoding a translationally controlled tumor protein,plays an important role in mercury tolerance in rice[J].BMC Plant Biology,15(1):1-15.
Yenofsky R,cerehini S,kraowczynska A,et al.1983.Regulation of mRNA utilization in mouse erythroleukemia cells induced to differentiate by exposure to dimethyl sulfoxide[J].Molecular Cell Bioogy,3(7):1197-1203.
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Deng Z,Chen J,Leclercg J,et al.2016.Expression profiles,characterization and function of HbTCTP in rubber tree(Hevea brasiliensis)[J].Frontiers in plant science,7:789.
Du J H,Yang P,Kong F H,et al.2017.Aberrant expression of translationally controlled tumor protein(TCTP)can lead to radioactive susceptibility and chemosensitivity in lung cancer cells[J].Oncotarget,8(60):101922-101935.
Ermolayev V,Weschke W,Manteuffel R.2003.Comparison of Al-induced gene expression in sensitive and tolerant soybean cultivars[J].Journal of Experimental Botany,54(393):2745-2756.
Hoepflinger M C,Reitsamer J,Geretschlaeger A M,et al.2013.The effect of translationally controlled tumour protein(TCTP)on programmed cell death in plants[J].BMC Plant Biology.13(1):135.
Hsu Y C,Chern J J,Cai Y,et al.2007.Drosophila TCTP is essential for growth and proliferation through regulation of dRheb GTPase[J].Nature,445(7129):785-788.
Issarakraisila M,Ma Q F,Turner D W.2007.Photosynthetic and growth responses of juvenile Chinese kale(Brassica oleracea var.alboglabra)and Caisin(Brassica rapa subsp parachinensis)to water logging and water deficit[J].Scientia Horticulture,111(2):107-113.
Jeon H J,Cui X S,Guo J,et al.2017.TCTP regulates spindle assembly during postovulatory aging and prevents deterioration in mouse oocyte quality[J].Elsevier Journal,1328-1334.
Kim Y M,Han Y J,Hwang O J,et al.2012.Overexpression of Arabidopsis translationally controlled tumor protein gene AtTCTP enhances drought tolerance with rapid ABA-induced stomatal closure[J].Molecules and Cells,33(6):617-626.
Li S,Chen M H,Xiong Q,et al.2016.Characterization of the translationally controlled tumor protein(TCTP)interactome reveals novel binding partners in human cancer cells[J].Journal of proteome research,15(10):3741-3751.
Lopez A P,Franco A R.2006.Cloning and expression of cD-NA encoding translationally controlled tumor protein from strawberry fruits[J].Biologia Plantarum,50(3)447-449.
Lucibello M,Gambacurta A,Zonfrillo M.2011.TCTP is a critical survival factor that protects cancer cells from oxidative stress induced cell-death[J].Experimental Cel Research,317(17):2479-2489.
Márcio de C,Márcio L A,Alessandra V N L,et al.2017.Impacts of the overexpression of a tomato translationally controlled tumor protein(TCTP)in tobacco revealed by phenotypic and transcriptomic analysis[J].Plant Cell Reports,36(6):887-900.
Meng X,Chen Q,Fan H,et al.2017.Molecular characterization,expression analysis and heterologous expression of two translationally controlled tumor protein genes from Cucumis sativus[J].PLoS One,12(9):e0184872
Meng X N,Yu Y,Zhao J Y,et al.2018.The two translationally controlled tumor protein genes,CsTCTP1 and CsT-CTP2,are negative modulators in the Cucumis sativus defense response to sphaerotheca fuliginea[J].In Plan Science,9:544.
Qian H M,Sun B,Miao H Y,et al.2015.Variation of glucosinolates and quinone reductase activity among different varieties of Chinese kale and improvement of glucoraphanin by metabolic engineering[J].Food Chemistry168:321-326.
Santa Brigida A B,Dos Reis S P,Costa C N M,et al.2014Molecular cloning and characterization of a cassava translationally controlled tumor protein gene potentially related to salt stress response[J].Molecular Biology Reports,41(3):1787-1797.
Sturzenbaum S R,Kille P,Morgan A J.1998.Identification of heavy metal induced changes in the expression patterns of the translationally controlled tumour protein(TCTP)in the earthworm Lumbricus rubellus[J].Biochimica e Biophysica Acta,1398(3):294-304.
Tsarova K,Yarmola E G,Bubb M R.2010.Identification of a cofilin-like actin-binding site on translationally controlled tumor protein(TCTP)[J].FEBS Letters,584(23)4756-4760.
Toscano-Morales R,Xoconostle-Cázares B,Martínez-Navarro A C,et al.2014.Long distance movement of an arabidopsis translationally controlled tumor protein(AtTCTP2)mRNA and protein in tobacco[J].Frontiers in Plant Science,(5):705.
Toscano-Morales R,Xoconostle-Cázares B,Cabrera-Ponce JL,et al.2015.AtTCTP2,an Arabidopsis thaliana homolog of translationally controlled tumor protein,enhances in vitro plant regeneration[J].Frontiers in Plant Science,(6):468.
Toscano-Morales R,Xoconostle-Ca′zares B,Mart?′nez-Navarro A C,et al.2016.AtTCTP2 mRNA and protein movement correlates with formation of adventitious roots in tobacco[J].Plant Signaling&Behavior,11(3):e1071003.
Tuynder M,Susini L,Prieur S,et al.2002.Biological models and genes of tumor reversion:Cellular reprogramming through tpt1/TCTP and SIAH-1[J].Proceedings of the National Academy of Sciences,99(23):14976-14981.
Wang L L,Tang Y F,Zhao M J,et al..2018.Knockdown of translationally controlled tumor protein inhibits growth,migration and invasion of lung cancer cells[J].Life Sciences,193:292-299.
Wang S,Zhao X F,Wang J X.2009.Molecular cloning and characterization of the translationally controlled tumor protein from Fenneropenaeus Chinensis[J].Molecular Biology Reports,36(7):1683-1693.
Wang Z Q,Li G Z,Gong Q Q,et al.2015.OsTCTP,encoding a translationally controlled tumor protein,plays an important role in mercury tolerance in rice[J].BMC Plant Biology,15(1):1-15.
Yenofsky R,cerehini S,kraowczynska A,et al.1983.Regulation of mRNA utilization in mouse erythroleukemia cells induced to differentiate by exposure to dimethyl sulfoxide[J].Molecular Cell Bioogy,3(7):1197-1203.
Zhang J M,Ma L F,Li W,et al.2014.Cotton TCTP1 gene encoding a translationally controlled tumor protein participates in plant response and tolerance to aphids[J].Plant Cell Tissue and Organ Culture,117(2):145-156
Zhu Z S,Sun B M,Xu X X,et al.2016.Overexpression of AtEDT1/HDG11 in Chinese kale(Brassica oleracea var.alboglabra)enhances drought and osmotic stress tolerance[J].Frontiers in Plant Science,(7):1285