苏云金芽胞杆菌S-层蛋白形成内涵体的研究
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摘要
细菌S-层,或叫表面层(Surface layer),简称S-layer,是古菌和真细菌最普通的表层结构,是由一种单分子的蛋白质亚单元组成的晶状二维排列。这种变化多样的最外层结构很好地反映了生物对特殊生态和环境条件的适应性。本文围绕苏云金芽胞杆菌(Bt)的S-层展开研究,主要结果如下:
1、对前人的研究结论进行了补充,证明了Bt.CTC菌株的S-层基因不仅能在细胞表面表达形成典型的S-层结构,而且还可以在细胞内部累积形成内涵体;该内涵体蛋白编码基因与ctc_2高度同源。
2、CTC菌株芽胞期蛋白聚集体(或称“内涵体”)实际上并非传统的伴胞晶体,该蛋白聚集体也并不伴随着芽胞形成而形成,并证明该蛋白聚集体表面披被一层膜状结构。该内涵体在基因组成、表面和内部结构上与传统的伴胞晶体有显著的不同。在Bt.种中,还查明了由S-层蛋白形成内涵体的菌株占总数的22.6%。这些CTC-类似菌株一般不具有生物活性或仅有低生物活性。总之,在本室前人结论的基础上,提出了苏云金芽胞杆菌种中存在一类新的内涵体蛋白。此结论为Bt种内广泛存在的无毒菌株提供了一个新的研究思路。
3、在前述结论上,发现CTC菌株内涵体的稳定表达及其降解与培养条件中的溶氧值有关,即与低氧条件有关。在发酵罐发酵条件下,应使其发酵溶氧浓度控制在2.0 mg/L。此时,经Western blot、RT-PCR和质谱测序表明,由S-层形成的内涵体蛋白为基因ctc2的表达产物。
4、基于苏云金芽胞杆菌野生菌株接收外源质粒比较困难的事实,基本建立了一种基于电穿孔方法的基因转移方法,即在细菌培养过程中,适量加入一定浓度的甘氨酸/溶菌酶,采用SH缓冲液洗涤和悬浮菌体,供体质粒脱盐后,先电穿孔再进行42℃热冲击的方法,转化效率大大提高。
5、通过双交换的方法对Bt.CTC菌株的S-层基因进行了成功缺失,继而对苏云金芽胞杆菌CTC菌株S-层的功能进行了初步探索,发现野生株可以耐受一定的碱性条件,从而证明S-层对于宿主的抗逆性有一定贡献。
This thesis focused on the research of S-layer(surface layer) of Bacillus thuringiensis.S-layer,also called crystalline bacterial cell surface layer,is a layer with two-dimensional lattice structure.It forms the outermost component of many archaeobacteria and eubacteria.Its functions have been proposed to be involved in cell integrity and shape maintenance,molecule or ion traps in cell adhesion,surface recognition,a factor of virulence,or a kind of murein hydrolase and protection from predacious attack by Bdellovibrio.
The results are summarized as following:
1.The S-layer protein resembling as CTC2 of Bt.str.CTC has been proved that it can not only locate on the surface of the cell,but serve as some structural and functional parts inside of the cell during sporulation.
2.Based on the previous research results in our lab,it is first reported that the proteinaceous mass(or termed as "inclusions") of the Bt.str.CTC,accumulated during sporulation,was not a typical crystal protein encoded by cry gene,but a proteinaceous inclusion coded by slp gene.Furthermore,the strains,with the so-called parasporal crystal encoded by S-layer gene during sporulation,are widely distributed and account for 22.6%among the Bt.species.All these strains have provided a new aspect for research of Bt nontoxic strains.All the above results show that a novel group of inclusions encoded by S-layer gene of Bt has been discovered.
3.During the further discovery,the formation of the inclusions coded by S-layer protein gene from Bt str.CTC is related to the dissolved oxygen concentration(DOC). The Bt str.CTC can accumulate the inclusion while growing on the condition of 2.0 mg/L DOC during sporulation.The ctc_2 gene is also testified to code for the inclusions by the method of Western blot,RT-PCR and polypeptide sequencing by mass spectrometry.
4.A new method basing on the electro-poration to transfer the heterogenous gene into the wild type strains among the B.cereus group has been constructed.The method is followed by the following procedures:adding suitable glycine or lysozyme during the cells culture,washing the cells with SH buffer,desalting the plasmid donor, heat-treatment at the temperature of 42℃after the electric pulse.
5.The role of S-layer of Bacillus thuringiensis strain CTC has been discussed. The genes,ctc_1 and ctc_2,responsible for the surface layer(S-layer) protein synthesis in Bt str.CTC,were mutated through the method of double crossover and resulted in Bt str.BMB0549.The results show that the S-layer proteins might function as some characteristics adaption to the basic environment.
1、对前人的研究结论进行了补充,证明了Bt.CTC菌株的S-层基因不仅能在细胞表面表达形成典型的S-层结构,而且还可以在细胞内部累积形成内涵体;该内涵体蛋白编码基因与ctc_2高度同源。
2、CTC菌株芽胞期蛋白聚集体(或称“内涵体”)实际上并非传统的伴胞晶体,该蛋白聚集体也并不伴随着芽胞形成而形成,并证明该蛋白聚集体表面披被一层膜状结构。该内涵体在基因组成、表面和内部结构上与传统的伴胞晶体有显著的不同。在Bt.种中,还查明了由S-层蛋白形成内涵体的菌株占总数的22.6%。这些CTC-类似菌株一般不具有生物活性或仅有低生物活性。总之,在本室前人结论的基础上,提出了苏云金芽胞杆菌种中存在一类新的内涵体蛋白。此结论为Bt种内广泛存在的无毒菌株提供了一个新的研究思路。
3、在前述结论上,发现CTC菌株内涵体的稳定表达及其降解与培养条件中的溶氧值有关,即与低氧条件有关。在发酵罐发酵条件下,应使其发酵溶氧浓度控制在2.0 mg/L。此时,经Western blot、RT-PCR和质谱测序表明,由S-层形成的内涵体蛋白为基因ctc2的表达产物。
4、基于苏云金芽胞杆菌野生菌株接收外源质粒比较困难的事实,基本建立了一种基于电穿孔方法的基因转移方法,即在细菌培养过程中,适量加入一定浓度的甘氨酸/溶菌酶,采用SH缓冲液洗涤和悬浮菌体,供体质粒脱盐后,先电穿孔再进行42℃热冲击的方法,转化效率大大提高。
5、通过双交换的方法对Bt.CTC菌株的S-层基因进行了成功缺失,继而对苏云金芽胞杆菌CTC菌株S-层的功能进行了初步探索,发现野生株可以耐受一定的碱性条件,从而证明S-层对于宿主的抗逆性有一定贡献。
This thesis focused on the research of S-layer(surface layer) of Bacillus thuringiensis.S-layer,also called crystalline bacterial cell surface layer,is a layer with two-dimensional lattice structure.It forms the outermost component of many archaeobacteria and eubacteria.Its functions have been proposed to be involved in cell integrity and shape maintenance,molecule or ion traps in cell adhesion,surface recognition,a factor of virulence,or a kind of murein hydrolase and protection from predacious attack by Bdellovibrio.
The results are summarized as following:
1.The S-layer protein resembling as CTC2 of Bt.str.CTC has been proved that it can not only locate on the surface of the cell,but serve as some structural and functional parts inside of the cell during sporulation.
2.Based on the previous research results in our lab,it is first reported that the proteinaceous mass(or termed as "inclusions") of the Bt.str.CTC,accumulated during sporulation,was not a typical crystal protein encoded by cry gene,but a proteinaceous inclusion coded by slp gene.Furthermore,the strains,with the so-called parasporal crystal encoded by S-layer gene during sporulation,are widely distributed and account for 22.6%among the Bt.species.All these strains have provided a new aspect for research of Bt nontoxic strains.All the above results show that a novel group of inclusions encoded by S-layer gene of Bt has been discovered.
3.During the further discovery,the formation of the inclusions coded by S-layer protein gene from Bt str.CTC is related to the dissolved oxygen concentration(DOC). The Bt str.CTC can accumulate the inclusion while growing on the condition of 2.0 mg/L DOC during sporulation.The ctc_2 gene is also testified to code for the inclusions by the method of Western blot,RT-PCR and polypeptide sequencing by mass spectrometry.
4.A new method basing on the electro-poration to transfer the heterogenous gene into the wild type strains among the B.cereus group has been constructed.The method is followed by the following procedures:adding suitable glycine or lysozyme during the cells culture,washing the cells with SH buffer,desalting the plasmid donor, heat-treatment at the temperature of 42℃after the electric pulse.
5.The role of S-layer of Bacillus thuringiensis strain CTC has been discussed. The genes,ctc_1 and ctc_2,responsible for the surface layer(S-layer) protein synthesis in Bt str.CTC,were mutated through the method of double crossover and resulted in Bt str.BMB0549.The results show that the S-layer proteins might function as some characteristics adaption to the basic environment.
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