苏云金芽胞杆菌的两类S-层蛋白共存于S-层和伴胞内涵体
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摘要
我们筛选了伴胞晶体组成为S-层蛋白的菌株,从约20株菌株中得到了除CTC菌株之外的4株苏云金芽胞杆菌菌株B22,I13,H67和BMB1152,并且将这类菌株称为PICS菌株(strains whose parasporal inclusions consist of S-layer protein)我们从这4株PICS菌株中克隆到8个典型的S-层蛋白基因,这些菌株的每个基因组都包含两个毗邻的s-层蛋白基因。遗传进化树分析表明这些S-层蛋白在进化树上位于两个大的分支,我们分别称为SLPls和SLP2s。我们对菌株CTC和BMB1152的两类S-层蛋白进行了研究,Western blot以及免疫荧光显微镜实验证明SLP1s和SLP2s都是S-层的组成成分,只是呈现两种表型,即CTC菌株的两个S-层蛋白在对数生长期和稳定期都表达形成一致的S-层；而BMB1152则在对数期主要由SLP1形成S-层,进入稳定期时则刚好相反,主要由SLP2形成S-层。对菌株CTC和BMB1152伴胞内涵体的免疫荧光显微镜检测证明在CTC和BMB1152菌株中,SLP1s和SLP2s都是伴胞内涵体的组成成分,这说明两类S-层蛋白都存在于伴胞内涵体。另外,我们将来自CTC和BMB1152的4个S-层蛋白基因在无晶体突变菌株BMB171进行异源表达,结果表明它们都能形成伴胞内涵体。以上这些结果表明,PICS菌株在苏云金芽胞杆菌具有一定的普遍性,它们的S-层蛋白基因在遗传进化树上分为两类,两类S-层蛋白共存于S-层和伴胞内涵体,两类S-层蛋白基因异源表达亦能形成伴胞内涵体。
苏云金芽胞杆菌幕虫亚种YBT-020被发现含有两种类型的伴胞晶体,一种与芽胞粘连,已知是由位于质粒上的cry26和cry28基因编码；另一种则是游离在芽胞外面,其组成成分尚不清楚。BMB1152是苏云金芽胞杆菌幕虫亚种YBT-020消除质粒的衍生菌株,该菌株不含有Cry蛋白组成的伴胞晶体,已知它的游离晶体成分是S-层蛋白,由此我们推测野生菌株YBT-020的游离晶体很可能是S-层蛋白,则在YBT-020中就会存在两种截然不同的类型的伴胞内涵体,那么他们的共存是否会对S-层蛋白的表达产生影响?为了研究YBT-020中游离晶体的组成以及两种伴胞晶体共存于苏云金芽胞杆菌对于S-层蛋白的表达和调控的影响,我们同时研究和比较了YBT-020和其缺失Cry晶体而仅包含S-层蛋白晶体的衍生菌株BMB1152的S-层蛋白表达情况。Western blot结果表明在YBT-020中存在两个S-层蛋白,但是由S-层蛋白组成的游离伴胞内涵体较其衍生菌株BMB1152不稳定。对三株菌株YBT-020和BMB1152(YBT-020无质粒突变株)以及BMB1151(YBT-020仅缺失cry26所在质粒pBMB26)芽胞期的S-层蛋白进行了SDS-PAGE检测,结果表明在YBT-020中100kDa的S-层蛋白的表达量相当低,而在突变株BMB1151和BMB1152中则大量表达,表明该蛋白的表达与cry26基因所在的质粒(pBMB26)相关,该质粒上可能存在着某些负调控因子影响着着S-层蛋白的表达。综合以上的研究结果表明,在YBT-020中,由cry基因和slp基因编码的两种类型的伴胞晶体同时存在,前者形成粘连型,后者形成游离型。在芽胞期两个s-层蛋白均表达但是都不稳定易降解,此时S-层蛋白的表达受到YBT-020中的质粒pBMB26的负调控。
S-layer proteins (SLPs) are typically thought to be constituents of the bacterial cell surface layer (S-layer). Previously we reported that SLPs can form parasporal inclusions in some Bacillus thuringiensis strains, and these inclusions consist of S-layer protein. But it is still unknown about whether these proteins are encoded by typical S-layer protein genes in phylogenetic level and whether S-layer protein gene directly lead to the formation of parasporal inclusions or S-layer. And it is need to know if S-layer and parasporal inclusions are encoded by the same gene. In this study, we screened four B. thuringiensis strains whose parasporal inclusions consist of S-layer protein (PICS strain) and cloned8slp genes from these strains. It is interesting that each strain contained two adjacent slp genes in each genome. Phylogenetic tree analysis indicated these SLPs could be divided into two groups, designated SLP1s and SLP2s. To confirm which SLPs were present in the S-layer or as a parasporal inclusion, two strains, CTC and BMB1152, were chosen for further study. Western blots using whole-cell associated proteins from strain CTC and BMB1152showed that SLP1s and SLP2s were both constituents of the S-layer. Immunofluorescence reaction with S-layer of strain CTC and BMB1152showed two phynotypes that in strain CTC, SLP1and SLP2expressed in similar grade to form a homologous S-layer in both exponential and stationary phases, but in strain BMB1152, SLP1expression was switched by SLP2gradually so a homologous SLP1-S-layer in exponential phase was replaced by SLP2-S-layer in stationary phase. Immunofluorescence reaction with spore-inclusion mixtures of strain CTC and BMB1152showed SLP1s and SLP2s were also both constituents of parasporal inclusions. When heterogeneously expressed in a crystal negative strain BMB171, four SLPs from strain CTC and BMB1152could also form parasporal inclusions. These mean both groups of SLPs are constituents of the S-layer and parasporal inclusions during different phases (vegetative or sporulation). This temporal and spatial expression is not an occasional phenomenon but ubiquitous in B. thuringiensis strains.
Bacillus thuringiensis subsp finitimus has two types of parasporal inclusions one of which are inside exosporium and coded by cry26and cry28genes that located in plasmids. Another type of inclusions is outside exosporium and constituents are unknown. In order to know the constitute of parasporal inlusions and S-layer of YBT-020, Western blots with S-layer and parasporal inclusions associated proteins were made and results showed that in YBT-020, SLP1expressed early and steady in exponential phase but SLP2expressed considerably late at about10hour so SLP1is the dominant proteins in S-layer formation of YBT-020. In sporulation phase, two S-layer proteins of YBT-020were unsteady and degraded into70kDa proteins different to BMB1152that SLP2protein expressed highly and only SLP1also degraded into70kDa. So in YBT-020free inclusions that coded by S-layer proteins were not steady like in BMB1152and so were not easy to be found like inclusions that codede by cry genes. SDS-PAGE of S-layer proteins of YBT-020and its variant strain BMB1152(crued all plasmids) and BMB1151(crued pBMB26) showed that only in YBT-020,100kDa S-layer protein was expressed lowly and inhibited compared with other two variants, which means that pBMB26may play roles in negatively regulation of S-layer protein expression in YBT-020. Our work showed that in YBT-020, there were two types of parasporal inclusions, one of which were free and consist of S-layer proteins. We also found that the expression of S-layer proteins in YBT-020was negatively regulated by the plasmid pBMB26.
苏云金芽胞杆菌幕虫亚种YBT-020被发现含有两种类型的伴胞晶体,一种与芽胞粘连,已知是由位于质粒上的cry26和cry28基因编码；另一种则是游离在芽胞外面,其组成成分尚不清楚。BMB1152是苏云金芽胞杆菌幕虫亚种YBT-020消除质粒的衍生菌株,该菌株不含有Cry蛋白组成的伴胞晶体,已知它的游离晶体成分是S-层蛋白,由此我们推测野生菌株YBT-020的游离晶体很可能是S-层蛋白,则在YBT-020中就会存在两种截然不同的类型的伴胞内涵体,那么他们的共存是否会对S-层蛋白的表达产生影响?为了研究YBT-020中游离晶体的组成以及两种伴胞晶体共存于苏云金芽胞杆菌对于S-层蛋白的表达和调控的影响,我们同时研究和比较了YBT-020和其缺失Cry晶体而仅包含S-层蛋白晶体的衍生菌株BMB1152的S-层蛋白表达情况。Western blot结果表明在YBT-020中存在两个S-层蛋白,但是由S-层蛋白组成的游离伴胞内涵体较其衍生菌株BMB1152不稳定。对三株菌株YBT-020和BMB1152(YBT-020无质粒突变株)以及BMB1151(YBT-020仅缺失cry26所在质粒pBMB26)芽胞期的S-层蛋白进行了SDS-PAGE检测,结果表明在YBT-020中100kDa的S-层蛋白的表达量相当低,而在突变株BMB1151和BMB1152中则大量表达,表明该蛋白的表达与cry26基因所在的质粒(pBMB26)相关,该质粒上可能存在着某些负调控因子影响着着S-层蛋白的表达。综合以上的研究结果表明,在YBT-020中,由cry基因和slp基因编码的两种类型的伴胞晶体同时存在,前者形成粘连型,后者形成游离型。在芽胞期两个s-层蛋白均表达但是都不稳定易降解,此时S-层蛋白的表达受到YBT-020中的质粒pBMB26的负调控。
S-layer proteins (SLPs) are typically thought to be constituents of the bacterial cell surface layer (S-layer). Previously we reported that SLPs can form parasporal inclusions in some Bacillus thuringiensis strains, and these inclusions consist of S-layer protein. But it is still unknown about whether these proteins are encoded by typical S-layer protein genes in phylogenetic level and whether S-layer protein gene directly lead to the formation of parasporal inclusions or S-layer. And it is need to know if S-layer and parasporal inclusions are encoded by the same gene. In this study, we screened four B. thuringiensis strains whose parasporal inclusions consist of S-layer protein (PICS strain) and cloned8slp genes from these strains. It is interesting that each strain contained two adjacent slp genes in each genome. Phylogenetic tree analysis indicated these SLPs could be divided into two groups, designated SLP1s and SLP2s. To confirm which SLPs were present in the S-layer or as a parasporal inclusion, two strains, CTC and BMB1152, were chosen for further study. Western blots using whole-cell associated proteins from strain CTC and BMB1152showed that SLP1s and SLP2s were both constituents of the S-layer. Immunofluorescence reaction with S-layer of strain CTC and BMB1152showed two phynotypes that in strain CTC, SLP1and SLP2expressed in similar grade to form a homologous S-layer in both exponential and stationary phases, but in strain BMB1152, SLP1expression was switched by SLP2gradually so a homologous SLP1-S-layer in exponential phase was replaced by SLP2-S-layer in stationary phase. Immunofluorescence reaction with spore-inclusion mixtures of strain CTC and BMB1152showed SLP1s and SLP2s were also both constituents of parasporal inclusions. When heterogeneously expressed in a crystal negative strain BMB171, four SLPs from strain CTC and BMB1152could also form parasporal inclusions. These mean both groups of SLPs are constituents of the S-layer and parasporal inclusions during different phases (vegetative or sporulation). This temporal and spatial expression is not an occasional phenomenon but ubiquitous in B. thuringiensis strains.
Bacillus thuringiensis subsp finitimus has two types of parasporal inclusions one of which are inside exosporium and coded by cry26and cry28genes that located in plasmids. Another type of inclusions is outside exosporium and constituents are unknown. In order to know the constitute of parasporal inlusions and S-layer of YBT-020, Western blots with S-layer and parasporal inclusions associated proteins were made and results showed that in YBT-020, SLP1expressed early and steady in exponential phase but SLP2expressed considerably late at about10hour so SLP1is the dominant proteins in S-layer formation of YBT-020. In sporulation phase, two S-layer proteins of YBT-020were unsteady and degraded into70kDa proteins different to BMB1152that SLP2protein expressed highly and only SLP1also degraded into70kDa. So in YBT-020free inclusions that coded by S-layer proteins were not steady like in BMB1152and so were not easy to be found like inclusions that codede by cry genes. SDS-PAGE of S-layer proteins of YBT-020and its variant strain BMB1152(crued all plasmids) and BMB1151(crued pBMB26) showed that only in YBT-020,100kDa S-layer protein was expressed lowly and inhibited compared with other two variants, which means that pBMB26may play roles in negatively regulation of S-layer protein expression in YBT-020. Our work showed that in YBT-020, there were two types of parasporal inclusions, one of which were free and consist of S-layer proteins. We also found that the expression of S-layer proteins in YBT-020was negatively regulated by the plasmid pBMB26.
引文
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