苏云金芽胞杆菌帮助蛋白对杀虫晶体蛋白表达的影响
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摘要
本文主要研究了苏云金芽胞杆菌以色列亚种的20kDa帮助蛋白对库斯塔克亚种的杀虫晶体蛋白基因crylAc的表达及其伴胞晶体形成的影响，针对棉铃虫等重要农业害虫构建并向野生型菌株中转化了含20kDa帮助蛋白基因的位点专一性重组质粒，获得了三株杀虫晶体蛋白高效表达的苏云金芽胞杆菌工程菌。主要结果如下：
在苏云金芽胞杆菌无晶体突变株中，20kDa帮助蛋白对crylAc的表达和伴胞晶体形成有明显的促进作用，毒力也有显著提高。结果表明，在该帮助蛋白的参与下CrylAc的表达量为不含帮助蛋白对照菌的3.5倍，形成的双金字塔形伴胞晶体长达1.85μm、宽0.85μm，体积为对照菌的3倍，同时对初孵棉铃虫幼虫的毒力提高了1.5倍。进一步分析表明，20kDa帮助蛋白是通过防止CrylAc新生肽免受胞内蛋白酶的降解而使得CrylAc表达量提高的。
本试验还利用苏云金芽胞杆菌转座子Tn4430的位点专一性重组系统，构建了含有crylAc10和20kDa蛋白基因的位点专一性重组质粒pBMB1808和仅含有20kDa帮助蛋白基因的位点专一性重组质粒pBMB1820，两质粒均为大肠杆菌／苏云金芽胞杆菌穿梭质粒。导入Bt后，在Tnpl的介导下可发生位点专一性重组、丢掉质粒中的抗生素抗性基因以及其它所有非Bt基因，而获得环境安全的工程菌。
通过电转化将pBMB1808导入到对棉铃虫、小菜蛾等鳞翅目害虫高毒力的三种天然菌株中。所得三种工程菌(BMB21882、BMB15358、BMB83383)的130kDa杀虫晶体蛋白的表达量均为各受体菌的两倍以上，各工程菌的伴胞晶体的平均体积为其受体菌的2-3倍，部分晶体长达4.0μm、宽1.4μm，大小为受体菌晶体的6倍。相反，工程菌芽胞形态由长椭圆形变为小圆球形。与受体菌相比，工程菌的杀虫晶体蛋白的表达曲线也具特点。受体菌晶体蛋白的表达量在芽胞分化开始后短时间内迅速达到最大，但随后缓慢下降，在晶体形成之前有三分之一被降解掉；而工程菌的表达量在整个芽胞形成期内呈持续的积累状态，最后获得超量表达。然而，工程菌的伴胞晶体对棉铃虫初孵幼虫的毒力不但没有上升反而下降。体外活化等试验结果表明，这可能是由伴胞晶体在昆虫体内溶解困难引起的。此外，本试验还发现位点专一性重组质粒在少部分转化子中有被修饰的现象，并导致了位点专一性重组能力的丧失。总之，本实验中将帮助蛋白基因置于位点专一性质粒用于工程菌的构建，获得了杀虫晶体蛋白的超量表达并形成了巨大的伴胞晶体。据我所知，这一结果以及帮助蛋白对CrylAc表达影响的机制皆属首此报导。但是关于如何进一步发挥工程菌巨大伴胞晶体的潜力、以及最终获得无抗性位点专一性重组子还有待于深入研究。
The effects of the 20kDa helper protein (P20) from Bacillus thuringiensis subsp. israelensis were examined on the expression and crystal formation of insecticidal crystal protein CrylAc, and towards the end to construct engineered strains to be both environment-friendly and highly efficient. The major results are described as following:
In the presence of P20, the expression of CrylAclO was 3.5 times of the control without P20, the formed crystals were in an average size of 1.85u.m long and 0.85um wide, shown as a typical bipyramid with 3-time volume of the control crystal. Consequently, the transformant harboring P20 gene and cry 1 Ac 10 exhibited 2.5-time toxicity of the control transformant against neonate larvae of Heliothis unnigera. Further study revealed that serious degradation of CrylAc happened during the whole stage of sporulation, while the 20kDa protein could prevent neonate peptides only instead of the full protoxins from degradation
Two plasmids with the ability of site-specific recombination were obtained, one is pBMB1808 carrying P20 gene and crylAclO between two specific resolution sites (res), the other is pBMB1820 harboring only P20 gene. Also, a Bt replicon was included between these two "res" sites. Other genes such as tnpl. antibiotic resistance genes and an E.coli replicon etc were positioned outside of the two "res" sites. Therefore, a site-specific recombination would happen between two "res" sites in B. thuringiensis, and enable the transformant to get rid of the unwanted DNA fragments. Thus, these two plasmids would be useful in the construction of an environment-friendly engineered strain with high productivity of ICPs.
Three engineered Bt strains BMB21882, BMB83383 and BMB15358 were obtained by electro-poration of pBMB1808 into three natural isolates with a high productivity of ICPs and high toxicity against H. armigera and Plutella. xylostella. Quantification results revealed that two times of CrylA protoxin produced in the transformants relative to their receipts. And larger bipyramidal crystals were resulted in a size of 2.40 u.m long and 1.05am wide, with 2-3 time volumes of their receipts' crystals. In contrast, the transformants' spores became shorter or spherical instead of long rods. Further study showed that continuous accumulation of CrylA protoxins during the whole sporulation stage was obtained in the engineered strains, while in natural strains a serious hydrolysis of 130kDa protoxin occurred before crystal formation.
Despite of the high expression of ICPs in these transformants, bioassay results failed to show the expected toxicity against their target insects, especially in the case of neonate larvae. However, in vitro activation of crystals in H. armigera midgut juice showed that much more toxic peptides were generated by transformants than the receipts. Further study suggested that the lowered toxicity might be resulted from the poor solubility of crystals in vivo.
Unfortunately, the obtained transformants failed to eliminate the antibiotic resistance gene harbored in pBMB1808. due to both the prompt excision in most transformants before antibiotic selection, and DNA modification on pBMB1808 in the obtained transformants. Therefore, more efforts needed to obtain an ideal engineered strain.
在苏云金芽胞杆菌无晶体突变株中，20kDa帮助蛋白对crylAc的表达和伴胞晶体形成有明显的促进作用，毒力也有显著提高。结果表明，在该帮助蛋白的参与下CrylAc的表达量为不含帮助蛋白对照菌的3.5倍，形成的双金字塔形伴胞晶体长达1.85μm、宽0.85μm，体积为对照菌的3倍，同时对初孵棉铃虫幼虫的毒力提高了1.5倍。进一步分析表明，20kDa帮助蛋白是通过防止CrylAc新生肽免受胞内蛋白酶的降解而使得CrylAc表达量提高的。
本试验还利用苏云金芽胞杆菌转座子Tn4430的位点专一性重组系统，构建了含有crylAc10和20kDa蛋白基因的位点专一性重组质粒pBMB1808和仅含有20kDa帮助蛋白基因的位点专一性重组质粒pBMB1820，两质粒均为大肠杆菌／苏云金芽胞杆菌穿梭质粒。导入Bt后，在Tnpl的介导下可发生位点专一性重组、丢掉质粒中的抗生素抗性基因以及其它所有非Bt基因，而获得环境安全的工程菌。
通过电转化将pBMB1808导入到对棉铃虫、小菜蛾等鳞翅目害虫高毒力的三种天然菌株中。所得三种工程菌(BMB21882、BMB15358、BMB83383)的130kDa杀虫晶体蛋白的表达量均为各受体菌的两倍以上，各工程菌的伴胞晶体的平均体积为其受体菌的2-3倍，部分晶体长达4.0μm、宽1.4μm，大小为受体菌晶体的6倍。相反，工程菌芽胞形态由长椭圆形变为小圆球形。与受体菌相比，工程菌的杀虫晶体蛋白的表达曲线也具特点。受体菌晶体蛋白的表达量在芽胞分化开始后短时间内迅速达到最大，但随后缓慢下降，在晶体形成之前有三分之一被降解掉；而工程菌的表达量在整个芽胞形成期内呈持续的积累状态，最后获得超量表达。然而，工程菌的伴胞晶体对棉铃虫初孵幼虫的毒力不但没有上升反而下降。体外活化等试验结果表明，这可能是由伴胞晶体在昆虫体内溶解困难引起的。此外，本试验还发现位点专一性重组质粒在少部分转化子中有被修饰的现象，并导致了位点专一性重组能力的丧失。总之，本实验中将帮助蛋白基因置于位点专一性质粒用于工程菌的构建，获得了杀虫晶体蛋白的超量表达并形成了巨大的伴胞晶体。据我所知，这一结果以及帮助蛋白对CrylAc表达影响的机制皆属首此报导。但是关于如何进一步发挥工程菌巨大伴胞晶体的潜力、以及最终获得无抗性位点专一性重组子还有待于深入研究。
The effects of the 20kDa helper protein (P20) from Bacillus thuringiensis subsp. israelensis were examined on the expression and crystal formation of insecticidal crystal protein CrylAc, and towards the end to construct engineered strains to be both environment-friendly and highly efficient. The major results are described as following:
In the presence of P20, the expression of CrylAclO was 3.5 times of the control without P20, the formed crystals were in an average size of 1.85u.m long and 0.85um wide, shown as a typical bipyramid with 3-time volume of the control crystal. Consequently, the transformant harboring P20 gene and cry 1 Ac 10 exhibited 2.5-time toxicity of the control transformant against neonate larvae of Heliothis unnigera. Further study revealed that serious degradation of CrylAc happened during the whole stage of sporulation, while the 20kDa protein could prevent neonate peptides only instead of the full protoxins from degradation
Two plasmids with the ability of site-specific recombination were obtained, one is pBMB1808 carrying P20 gene and crylAclO between two specific resolution sites (res), the other is pBMB1820 harboring only P20 gene. Also, a Bt replicon was included between these two "res" sites. Other genes such as tnpl. antibiotic resistance genes and an E.coli replicon etc were positioned outside of the two "res" sites. Therefore, a site-specific recombination would happen between two "res" sites in B. thuringiensis, and enable the transformant to get rid of the unwanted DNA fragments. Thus, these two plasmids would be useful in the construction of an environment-friendly engineered strain with high productivity of ICPs.
Three engineered Bt strains BMB21882, BMB83383 and BMB15358 were obtained by electro-poration of pBMB1808 into three natural isolates with a high productivity of ICPs and high toxicity against H. armigera and Plutella. xylostella. Quantification results revealed that two times of CrylA protoxin produced in the transformants relative to their receipts. And larger bipyramidal crystals were resulted in a size of 2.40 u.m long and 1.05am wide, with 2-3 time volumes of their receipts' crystals. In contrast, the transformants' spores became shorter or spherical instead of long rods. Further study showed that continuous accumulation of CrylA protoxins during the whole sporulation stage was obtained in the engineered strains, while in natural strains a serious hydrolysis of 130kDa protoxin occurred before crystal formation.
Despite of the high expression of ICPs in these transformants, bioassay results failed to show the expected toxicity against their target insects, especially in the case of neonate larvae. However, in vitro activation of crystals in H. armigera midgut juice showed that much more toxic peptides were generated by transformants than the receipts. Further study suggested that the lowered toxicity might be resulted from the poor solubility of crystals in vivo.
Unfortunately, the obtained transformants failed to eliminate the antibiotic resistance gene harbored in pBMB1808. due to both the prompt excision in most transformants before antibiotic selection, and DNA modification on pBMB1808 in the obtained transformants. Therefore, more efforts needed to obtain an ideal engineered strain.
引文
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