转基因毛白杨生理生化特性及多基因表达载体构建
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摘要
本文以4个转rolB-pttGA20ox双价基因毛白杨株系和1个对照株系PT-16为材料,探讨了rolB-pttGA20ox双价基因对毛白杨生根生理、光合特性、抗逆性以及生长的影响。为进一步提高毛白杨的抗虫性和耐盐碱能力,采用一套基于Cre/loxP重组系统的植物多基因表达载体系统将苏云杆菌毒蛋白基因(BtCrylAc)、甜菜碱醛脱氢酶基因(BADH)、GA20氧化酶基因(pttGA20ox)和rolB基因构建于同一植物表达载体pYL1305上,转化烟草,验证了多基因的表达和功能。主要研究结果如下：
1、转rolB-pttGA20ox双价基因毛白杨试管苗生根过程中内源IAA、GA3、ZR和ABA含量变化与对照差异明显,各项生根指标均高于对照。转基因株系不同程度地增大了对光的生态适应范围,其净光合速率与地径显著相关,与苗高相关不显著。盆栽期,各株系间苗高存在极显著差异,RG-1、RG-2和RG-3显著的高于RG-4和对照PT-16,地径差异不显著；移入大田后,各株系间苗高差异不显著,而地径出现显著差异,RG-4的平均苗高最高,达到了231.25cm,转基因株系中RG-4的平均地径最大,为13.91mm。对各株系的SOD活性和MDA含量测定表明,转基因株系RG-4对逆境的抵抗力最强,但与对照PT-16差异不显著。
rolB-pttGA20ox双价基因的表达可能改变了毛白杨相关的生理代谢途径,同时,转基因株系在光合特性、生长、抗逆性等方面的差异明显,这也为选择优良的转基因株系提供了材料。
2、采用PCR方法,扩增Bt基因,替换植物表达载体pYL1305中的gus基因,得到pYL1305-Bt。扩增CaMV35S-BADH-nos表达盒,并插入到pYL1305-Bt的多克隆位点,得到载体pYL 1305-Bt-BADH。将CaMV35S-GA20ox-nos表达盒、GH3-rolB-nos表达盒分别克隆到供给载体pYL-d1、pYL-d2上,经两轮重组得到多基因单载体pYL1305-Bt-BADH-GA20ox-rolB,并将其转化根癌农杆菌菌株EHA105。
3、采用农杆菌介导的叶盘法将多基因转化烟草,对获得的抗性芽在含25mg／LHyg的培养基中进一步选择后进行分子检测,PCR检测和Southern杂交检测证明多基因已插入到烟草基因组中。进一步的RT-PCR检测证明,外源基因均得到表达,多基因表现为串联在一起插入到同一位点,其中转基因株系T2中各基因的表达量最高。
4、转多基因烟草植株根部、茎部和叶片中IAA的含量均高于对照,分别是是对照的1.90倍、1.92倍和1.17倍；转化植株各部位的GA3含量均高于对照,转化植株中茎部GA3含量最高,叶片最低；转化植株的根、茎部ABA含量略低于对照,叶片中ABA含量高于对照；转化植株各部位的IAA/ABA比值均高于对照,其中根部的IAA/ABA值最高,是对照的1.82倍。
5、转多基因烟草耐盐性试验表明,当NaCl为300mM时,对照生长受到抑制,叶片膨大黄化,不能生根,转基因植株生长正常,叶片保持绿色,并能正常生根。转基因烟草经不同浓度的NaCl长时间胁迫,其SOD活性高于对照,MDA含量变化幅度较小,而对照MDA含量随盐浓度升高而大幅降低。
6、转基因烟草植株的茎、叶中均检测到了Bt毒蛋白,T2株系Bt毒蛋白含量最高,叶片中达到了585.53 ng/g.FW,茎中为573.66 ng/g.FW,而对照中没有检测到Bt毒蛋白。抗虫试验结果表明,饲喂一周后,转基因烟草对2龄舞毒蛾的校正死亡率为78.05%,对3龄斜纹夜蛾的死亡率为30.00%,并明显抑制了存活幼虫的生长发育。
7、移栽后的转基因烟草植株高生长更加明显,节间伸长,表现出了pttGA20ox基因表达的性状特征,转基因植株地径也显著增大、叶片大而深绿、植株健壮,移栽4 W后转化植株平均苗高为19.28cm,地径为5.34mm,分别是对照的1.61倍和1.37倍。
多基因在烟草中均能正常表达,没有出现基因沉默等现象,说明在构建多基因表达载体时,使用一定数目的同源序列的启动子和终止子是可行的,但随着基因的增多应该考虑更换不同启动子或在基因两端添加核基质结合区(MAR)序列以避免转基因沉默。多基因单载体的构建和各基因的正常表达为进一步的毛白杨等林木改良奠定了重要基础。
To reveal the effect of introduced rolB-pttGA20ox double genes on the root physiology, photosynthetic characteristics, stress resistance and growth of transgenic Chinese white poplar (Populus tomentosa Carr.), four Chinese white poplar lines with the indroduction of rolB-pttGA20ox double genes and a control strain PT-16 were invesgitated. To further improve the insect resistance and salt tolerance of Chinese white poplar, Bacillus thuringiensis toxin gene (BtCryIAc), betaine aldehyde dehydrogenase gene (BADH), GA20 oxidase gene (pttGA20ox) and the rolB gene were constructed in one plant expression vector pYL1305 through a set of multi-genes expression vectors based on Cre/loxP recombination system. Furthermore, the multi-gene vector was transformed into tobacco by agrobacterium mediated procedure. The main results of the research are as follows:
1) The difference of endogenous IAA, GA3, ZR and the ABA content changes between transgenic Chinese white poplar with rolB-pttGA20ox and the control was significant during the rooting process, and transgenic Chinese white poplar had better rooting ability as shown in the indexes investigated. In the transgenic lines, the the range of ecological adaptation of light was increased in different degrees. The net photosynthetic rate was significantly correlated with the basal stems, but was of no significant correlation with the height. The height of potted transgenic lines varied greatly and the height of RG-1, RG-2, and RG-3 was significantly higher than that of RG-4 and the control PT-16. Whereas the stem difference was not significant during the potted period. After the lines being moved to farmland, the result was just opposite, with RG-4 being the highest, which reached 231.25cm with the average stem of 13.91mm. The SOD activity and MDA content in different lines indicated that RG-4 has the strongest resistance to adversity, however, the difference was not significant when compared with PT-16.
The introduction of rolB-pttGA20ox double genes may have changed the physiological metabolic pathways of Chinese white poplar. Meanwhile the transgenic lines were significantly different in photosynthetic characteristics, growth, stress resistance, etc., which laid a foundation for the selection of high-quality transgenic plants.
2) pYL1305-Bt was constructed by replacing the gus gene of pYL1305 vector with the Bt gene obtained through PCR. The CaMV35S-BADH-nos expression cassette was amplfied and inserted into pYL 1305-Bt, resulted in the vector pYL 1305-Bt-BADH. The CaMV35S-GA20ox-nos and GH3-rolB-nos expression cassettes were insered into the supplying vector pYL-d1, pYL-d2, respectively; then the multi-gene vector was obtained by two rounds of reconstruction, which was transformed into EHA105.
3) The multi-gene vector was transformed into tobacco through Agrobacterium-mediated leaf disc transformation. The resistant plantlets got were screened by the culture on the medium with 25mg/L of Hyg. PCR analysis and Southern-blot detection to the resistance plantlets selected by antibiotics indicated that the genes had inserted into tobacco genome. Further RT-PCR test showed that all the target genes were transcribed in the transgenic plants, and that the expression of genes in line T2 was the highest. It was also showed that multigene is truly inserted tandemly.
4) The IAA levels in the roots, the stems and the leaves of transgenic tobacco plants were 1.90 fold,1.92 fold, and 1.17 fold as high as those of the control, respectively. The GA3 content of transformed plants in all parts was higher than those in the control plants, and that in the stems was the highest while that in the leaves was the lowest. ABA contents in the roots and the stems of transformed plants were slightly lower than those in the control ones, whereas in the leaves, they were higher in the transgenic plants than those in the control plants. IAA/ABA ratios of all parts in transformed plants were higher than those in the control ones, with that in roots being the highest, which was 1.82 times as large as that of the control plants.
5) Under 300mM NaCl.treatment, the gowth of control plants was restrained. The leaves turned yellow and it could not take roots. Whereas the growth of transgenic plants was normal with green leaves and the roots in good conditions. The SOD activity of transgenic plants was higher than that of the control plants when stressed by different concentration of NaCl for long duration, but MDA content changed slightly. While the MDA content of control plants decreased sharply with the increasing concentration of NaCl.
6) The Bt toxic protein was detected in both stems and leaves of the transgenic plants, with that of T2 plants being the highest, of which the Bt toxic protein in the leaves and stems were 585.53 ng/g.FW and 573.66 ng/g.FW respectively. While in the control plants, no Bt toxic protein was detected. A one-week feeding experiments showed that the corrected death rate of 2-year-old tussock moth was 78.05%, and the death rate of 3-year-old tobacco cutworm was 30%, which restrained the growth of larvae obviously.
7) The transgenic tobacco plants grew obviously after being transplanted. The stems were longer, which showed the characteristic of the pttGA20ox genes expression. The basal stem was also bigger, and the leaves were dark green and the plants were strong. After 4 weeks, the average height was 19.28cm, which was 1.61 times as high as that of the control plants; while the basal stem was 5.34mm, which was 1.37 times as big as that of the control plants.
In this study, the multiple genes could be expressed normaly in tobacco without resulting in gene silence, which showed that the construction of vector which has a certain number of homologous promoter and terminator was feasible. But with the increasing of the number of the genes introduced, the replacing of different promoters and the employment of MAR should be considered to avoid gene silence. The successful construction of single multi-gene vector and the normal expression of the genes will lay an important foundation for further improvements of Chinese white poplar and other trees.
1、转rolB-pttGA20ox双价基因毛白杨试管苗生根过程中内源IAA、GA3、ZR和ABA含量变化与对照差异明显,各项生根指标均高于对照。转基因株系不同程度地增大了对光的生态适应范围,其净光合速率与地径显著相关,与苗高相关不显著。盆栽期,各株系间苗高存在极显著差异,RG-1、RG-2和RG-3显著的高于RG-4和对照PT-16,地径差异不显著；移入大田后,各株系间苗高差异不显著,而地径出现显著差异,RG-4的平均苗高最高,达到了231.25cm,转基因株系中RG-4的平均地径最大,为13.91mm。对各株系的SOD活性和MDA含量测定表明,转基因株系RG-4对逆境的抵抗力最强,但与对照PT-16差异不显著。
rolB-pttGA20ox双价基因的表达可能改变了毛白杨相关的生理代谢途径,同时,转基因株系在光合特性、生长、抗逆性等方面的差异明显,这也为选择优良的转基因株系提供了材料。
2、采用PCR方法,扩增Bt基因,替换植物表达载体pYL1305中的gus基因,得到pYL1305-Bt。扩增CaMV35S-BADH-nos表达盒,并插入到pYL1305-Bt的多克隆位点,得到载体pYL 1305-Bt-BADH。将CaMV35S-GA20ox-nos表达盒、GH3-rolB-nos表达盒分别克隆到供给载体pYL-d1、pYL-d2上,经两轮重组得到多基因单载体pYL1305-Bt-BADH-GA20ox-rolB,并将其转化根癌农杆菌菌株EHA105。
3、采用农杆菌介导的叶盘法将多基因转化烟草,对获得的抗性芽在含25mg／LHyg的培养基中进一步选择后进行分子检测,PCR检测和Southern杂交检测证明多基因已插入到烟草基因组中。进一步的RT-PCR检测证明,外源基因均得到表达,多基因表现为串联在一起插入到同一位点,其中转基因株系T2中各基因的表达量最高。
4、转多基因烟草植株根部、茎部和叶片中IAA的含量均高于对照,分别是是对照的1.90倍、1.92倍和1.17倍；转化植株各部位的GA3含量均高于对照,转化植株中茎部GA3含量最高,叶片最低；转化植株的根、茎部ABA含量略低于对照,叶片中ABA含量高于对照；转化植株各部位的IAA/ABA比值均高于对照,其中根部的IAA/ABA值最高,是对照的1.82倍。
5、转多基因烟草耐盐性试验表明,当NaCl为300mM时,对照生长受到抑制,叶片膨大黄化,不能生根,转基因植株生长正常,叶片保持绿色,并能正常生根。转基因烟草经不同浓度的NaCl长时间胁迫,其SOD活性高于对照,MDA含量变化幅度较小,而对照MDA含量随盐浓度升高而大幅降低。
6、转基因烟草植株的茎、叶中均检测到了Bt毒蛋白,T2株系Bt毒蛋白含量最高,叶片中达到了585.53 ng/g.FW,茎中为573.66 ng/g.FW,而对照中没有检测到Bt毒蛋白。抗虫试验结果表明,饲喂一周后,转基因烟草对2龄舞毒蛾的校正死亡率为78.05%,对3龄斜纹夜蛾的死亡率为30.00%,并明显抑制了存活幼虫的生长发育。
7、移栽后的转基因烟草植株高生长更加明显,节间伸长,表现出了pttGA20ox基因表达的性状特征,转基因植株地径也显著增大、叶片大而深绿、植株健壮,移栽4 W后转化植株平均苗高为19.28cm,地径为5.34mm,分别是对照的1.61倍和1.37倍。
多基因在烟草中均能正常表达,没有出现基因沉默等现象,说明在构建多基因表达载体时,使用一定数目的同源序列的启动子和终止子是可行的,但随着基因的增多应该考虑更换不同启动子或在基因两端添加核基质结合区(MAR)序列以避免转基因沉默。多基因单载体的构建和各基因的正常表达为进一步的毛白杨等林木改良奠定了重要基础。
To reveal the effect of introduced rolB-pttGA20ox double genes on the root physiology, photosynthetic characteristics, stress resistance and growth of transgenic Chinese white poplar (Populus tomentosa Carr.), four Chinese white poplar lines with the indroduction of rolB-pttGA20ox double genes and a control strain PT-16 were invesgitated. To further improve the insect resistance and salt tolerance of Chinese white poplar, Bacillus thuringiensis toxin gene (BtCryIAc), betaine aldehyde dehydrogenase gene (BADH), GA20 oxidase gene (pttGA20ox) and the rolB gene were constructed in one plant expression vector pYL1305 through a set of multi-genes expression vectors based on Cre/loxP recombination system. Furthermore, the multi-gene vector was transformed into tobacco by agrobacterium mediated procedure. The main results of the research are as follows:
1) The difference of endogenous IAA, GA3, ZR and the ABA content changes between transgenic Chinese white poplar with rolB-pttGA20ox and the control was significant during the rooting process, and transgenic Chinese white poplar had better rooting ability as shown in the indexes investigated. In the transgenic lines, the the range of ecological adaptation of light was increased in different degrees. The net photosynthetic rate was significantly correlated with the basal stems, but was of no significant correlation with the height. The height of potted transgenic lines varied greatly and the height of RG-1, RG-2, and RG-3 was significantly higher than that of RG-4 and the control PT-16. Whereas the stem difference was not significant during the potted period. After the lines being moved to farmland, the result was just opposite, with RG-4 being the highest, which reached 231.25cm with the average stem of 13.91mm. The SOD activity and MDA content in different lines indicated that RG-4 has the strongest resistance to adversity, however, the difference was not significant when compared with PT-16.
The introduction of rolB-pttGA20ox double genes may have changed the physiological metabolic pathways of Chinese white poplar. Meanwhile the transgenic lines were significantly different in photosynthetic characteristics, growth, stress resistance, etc., which laid a foundation for the selection of high-quality transgenic plants.
2) pYL1305-Bt was constructed by replacing the gus gene of pYL1305 vector with the Bt gene obtained through PCR. The CaMV35S-BADH-nos expression cassette was amplfied and inserted into pYL 1305-Bt, resulted in the vector pYL 1305-Bt-BADH. The CaMV35S-GA20ox-nos and GH3-rolB-nos expression cassettes were insered into the supplying vector pYL-d1, pYL-d2, respectively; then the multi-gene vector was obtained by two rounds of reconstruction, which was transformed into EHA105.
3) The multi-gene vector was transformed into tobacco through Agrobacterium-mediated leaf disc transformation. The resistant plantlets got were screened by the culture on the medium with 25mg/L of Hyg. PCR analysis and Southern-blot detection to the resistance plantlets selected by antibiotics indicated that the genes had inserted into tobacco genome. Further RT-PCR test showed that all the target genes were transcribed in the transgenic plants, and that the expression of genes in line T2 was the highest. It was also showed that multigene is truly inserted tandemly.
4) The IAA levels in the roots, the stems and the leaves of transgenic tobacco plants were 1.90 fold,1.92 fold, and 1.17 fold as high as those of the control, respectively. The GA3 content of transformed plants in all parts was higher than those in the control plants, and that in the stems was the highest while that in the leaves was the lowest. ABA contents in the roots and the stems of transformed plants were slightly lower than those in the control ones, whereas in the leaves, they were higher in the transgenic plants than those in the control plants. IAA/ABA ratios of all parts in transformed plants were higher than those in the control ones, with that in roots being the highest, which was 1.82 times as large as that of the control plants.
5) Under 300mM NaCl.treatment, the gowth of control plants was restrained. The leaves turned yellow and it could not take roots. Whereas the growth of transgenic plants was normal with green leaves and the roots in good conditions. The SOD activity of transgenic plants was higher than that of the control plants when stressed by different concentration of NaCl for long duration, but MDA content changed slightly. While the MDA content of control plants decreased sharply with the increasing concentration of NaCl.
6) The Bt toxic protein was detected in both stems and leaves of the transgenic plants, with that of T2 plants being the highest, of which the Bt toxic protein in the leaves and stems were 585.53 ng/g.FW and 573.66 ng/g.FW respectively. While in the control plants, no Bt toxic protein was detected. A one-week feeding experiments showed that the corrected death rate of 2-year-old tussock moth was 78.05%, and the death rate of 3-year-old tobacco cutworm was 30%, which restrained the growth of larvae obviously.
7) The transgenic tobacco plants grew obviously after being transplanted. The stems were longer, which showed the characteristic of the pttGA20ox genes expression. The basal stem was also bigger, and the leaves were dark green and the plants were strong. After 4 weeks, the average height was 19.28cm, which was 1.61 times as high as that of the control plants; while the basal stem was 5.34mm, which was 1.37 times as big as that of the control plants.
In this study, the multiple genes could be expressed normaly in tobacco without resulting in gene silence, which showed that the construction of vector which has a certain number of homologous promoter and terminator was feasible. But with the increasing of the number of the genes introduced, the replacing of different promoters and the employment of MAR should be considered to avoid gene silence. The successful construction of single multi-gene vector and the normal expression of the genes will lay an important foundation for further improvements of Chinese white poplar and other trees.
引文
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