蝶蛹金小蜂热激蛋白及其体内共生菌Wolbachia分子分型的研究
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摘要
热激蛋白(Heat Shock Proteins，Hsps)又称热休克蛋白或应激蛋白(Heat StressProteins，Hsps)，是普遍存在于原核和真核生物中的一类遗传上高度保守的蛋白。Hsps的表达和调控系统是生物有机体对多种环境胁迫因子产生应激反应达到自我保护的重要物质基础。Wolbachia是广泛分布于节肢动物体内的一类细胞质遗传共生菌，它参与了多种调控其宿主生殖活动的机制，包括诱导细胞质不亲和(CI)、诱导孤雌生殖(PI)、雄性致死和雌性化。本研究克隆了蝶蛹金小蜂体内6个hsps基因，并对其诱导表达模式及功能进行了深入研究。另外，本研究还通过克隆和分析采自中国13个地理蝶蛹金小蜂体内Wolbachia的wsp、ftsZ和16SRNA基因对其感染的Wolbachia进行了分子分型研究。现将主要结果概括如下：
1蝶蛹金小蜂hsps基因的克隆及其序列分析
通过数据库检索、PCR／RT-PCR、RACE和文库筛选等分子手段，共克隆了蝶蛹金小蜂6个热激蛋白成员，涉及了5个热激蛋白家族。
Pphsp90基因ORF长为2148 bp，推测编码716个氨基酸，预测蛋白分子量为82.2 kDa，编码区内不含内含子。Pphsc70-1基因ORF长为1968 bp，推测编码656个氨基酸，预测蛋白分子量为71.3 kDa，该基因含有3个内含子。Pphsc70-2基因ORF长为1923 bp，推测编码641个氨基酸，预测蛋白分子量为70.9 kDa，编码区内含有1个内含子。Pphsp60基因ORF长为1719 bp，推测编码573个氨基酸，预测蛋白分子量为60.6 kDa，编码区内含有3个内含子。Pphsp40基因ORF长为1095 bp，推测编码365个氨基酸，预测蛋白分子量为39.6 kDa。Pphsp20基因ORF长为573 bp，推测编码191个氨基酸，预测蛋白分子量为21.7 kDa。
2不同逆境胁迫对蝶蛹金小蜂体内hsps基因表达的影响
采用TaqMan-MGB探针和特异性引物构建了6个Pphsps基因实时荧光定量RT-PCR检测体系。利用荧光定量RT-PCR方法，检测了蝶蛹金小蜂成虫6个Pphsps基因在高温(30～42℃处理1 h)、低温(3～-15℃处理1 h)、饥饿(0～24 h)及重金属(0.5～5mM Cd~(2+)／Cu~(2+)处理24 h或60 h)等4种逆境胁迫下mRNA的表达情况。
在高温胁迫下，6个Pphsps基因均能被诱导；其中，Pphsp20、Pphsp60、Pphsc70-2和Pphsp90表达量在36℃达到顶峰，而Pphsp40和Pphsc70-1表达量在39℃才达到最高峰。
在低温胁迫下，除Pphsp60基因外，其它5个基因均能被低温胁迫所诱导，并且其表达量均在-3℃时达到最高峰。
在饥饿胁迫下，Pphsp40和Pphsc70-2诱导前后无显著变化；Pphsp20表达量在饥饿6 h时就显著上升，其后随饥饿时间延长，表达量逐渐下降，甚至降至正常水平以下；Pphsp60、Pphsc70-1和Pphsp90表达量均在24 h显著升高。
在重金属Cd~(2+)胁迫下，处理24 h后，Pphsp40表达量随Cd~(2+)浓度的升高而持续升高，而其它5个基因呈先显著上升，其后，随Cd~(2+)浓度的升高，表达量呈下降趋势；处理60 h后，Pphsp40表达量随处理浓度的升高而升高；Pphsp60表达量在Cd~(2+)浓度为0.5 mM时显著上升，而后随Cd~(2+)浓度升高其表达量恢复至对照水平；Pphsc70-1表达量在Cd~(2+)浓度为50 mM时显著上升；其它Pphsps表达量与对照相比无明显差异或显著低于对照水平。
在重金属Cu~(2+)胁迫下，处理24 h后，Pphsp40和Pphsc70-2表达量随处理Cu~(2+)浓度的升高而升高，其它4个基因表达量随处理Cu~(2+)浓度的升高呈先上升后下降趋势；处理60 h后，与对照相比，仅0.5 mM的Cu~(2+)能够诱导Pphsp20和Pphsp90表达量分别上升1.32倍和1.30倍，而大多Pphsps表达量接近于或低于对照水平。
此外，还采用实时定量PCR方法(SYRB Green染料法)检测了蝶蛹金小蜂蛹期Pphsc70-1在40℃热激胁迫0～8 h及恢复1 h或2 h后的表达情况。结果表明，热处理1 h后Pphsc70-1表达量显著上升，然而持续的热激或热激恢复使其诱导的mRNA表达量逐渐降低，甚至显著低于对照水平。
3蝶蛹金小蜂hsps基因在昆虫细胞和原核细胞中表达
采用的Bac-to-Bac杆状病毒表达系统成功地将6个Pphsps基因在昆虫细胞中进行了表达。亚细胞定位分析表明6个基因的表达产物均位于细胞核外的细胞质中。
采用pET28a表达载体将6个Pphsps基因在表达菌株BL21(DE3)中进行原核表达。其中，Pphsp90、Pphsc70-1、Pphsc70-2、Pphsp60和Pphsp20等5个基因在该系统中都得到了不同程度的表达，且它们产生的融合表达蛋白基本上以可溶的形式存在。而Pphsp40基因在该系统中没有表达成功。
4蝶蛹金小蜂Hsps表达产物的纯化及其功能初探
为了研究PpHsps的分子伴侣功能，本实验研究了过表达PpHsps对E．coli细胞在高温下的存活率及E．coli细胞内蛋白热稳定性的影响。结果表明，虽然PpHsp90、PpHsc70-1、PpHsc70-2、PpHsp60和PpHsp20融合蛋白对高温下E．coli细胞内蛋白的稳定性具有一定的保护作用，但仅PpHsp20和PpHsp90对E．coli细胞在高温下的存活有保护作用。因此，我们采用Ni-NTA树脂分别对PpHsp20和PpHsp90融合蛋白进行分离纯化，研究了PpHsp20和PpHsp90对荧光素酶的高温保护能力。结果发现PpHsp20可以在一定程度上抑制高温引起的荧光素酶的聚集现象。
5蝶蛹金小蜂体内Wolbachia wsp、ftsZ和16S RNA基因的克隆及其在不同地理种群中的差异
13地理种群的蝶蛹金小蜂均感染B-Wolbachia。分析从13个地理种群中获得序列，共发现4条wsp基因差异序列(wPup1、wPup2、wPup3和wPup4)、2条ftsZ基因差异序列(fPup1和fPup2)和2条16S rRNA差异序列(16SPup1和16SPup2)；wsp基因在不同Wolbachia株系间存在高度重组。
此外，本研究还发现一个有趣的现象，在蝶蛹金小蜂南方种群中，除上海种群外，仅发现wsp、ftsZ和16S rRNA序列各一条；而在北方种群中，均存在2条或2条以上的wsp、ftsZ和16S rRNA序列。这一结果提示，南方种群(除上海种群)仅感染1个Wolbachia株，而北方种群感染2个Wolbachia株。
Heat shock proteins (Hsps) are a group of conserved proteins in evolution, whichexist in all eukaryotic and prokaryotic organisms. Hsps are very important fororganisms to survive in the stress. Wolbachia are maternally inherited bacteria thatspread widely in arthropods and can induce cytoplasmic incompatibility (CI),parthenogneesis, male killing and feminization. In this study, 6 heat shock genes inthe Pteromalus puparum were cloned and characterized, and their functions were alsostudied. Moreover, an investigation to determine the diversity of Wolbachia in P.puparum from 13 geographically distributed native populations in both southern andnorthern China were undertaken by comparing the sequences of Wolbachia 16S rRNA,ftsZ and wap genes. The results are summarized as follows:
1 Cloning and characterization of the hsps in the P. puparum
Six full-length ORFs coding Hsps involved five Hsp families were cloned by themethods of Blast search, PCR/RT-PCR, RACE and cDNA library screening.
The full-length ORF of the Pphsp90 gene was 2148 bp which was predicted toencode a 716 amino acid peptide with a deduced molecule weight of 82.2 kDa, and nointron was found. The full-length ORF of the Pphsc70-1 gene was 1968 bp which waspredicted to encode a 656 amino acid peptide with a deduced molecule weight of 71.3kDa, and three introns were found. The full-length ORF of the Pphsc70-2 gene was1923 bp which was predicted to encode a 641 amino acid peptide with a deducedmolecule weight of 70.9 kDa, and one intron was found. The full-length ORF of thePphsp60 gene was 1719 bp which was predicted to encode a 573 amino acid peptidewith a deduced molecule weight of 60.6 kDa, and three introns were found. Thefull-length ORF of the Pphsp40 gene was 1095 bp which was predicted to encode a365 amino acid peptide with a deduced molecule weight of 39.6 kDa. The full-lengthORF of the Pphsp20 gene was 573 bp which was predicted to encode a 191 aminoacid peptide with a deduced molecule weight of 21.7 kDa.
2 The Pphsps expression in relation to different stressors
TaqMan-MGB probes and the corresponding primers were designed to constructa system for real-time fluorescence quantitative RT-PCR to detect the changes inexpression levels of the six Pphsps in the adult wasps under the stressors, such as hightemperature (30～42℃for 1h), low temperature (3～15℃for 1h), starvation (0～24h) and heavy metals (0.5～5 mM Cd~(2+)/Cu~(2+) treated 24h or 60h).
Under the stress of high temperature, the expression of six Pphsps could beinduced. The genes, Pphsp20, Pphsp60, Pphsc70-2 and Pphsp90, had highestexpression levels after heat shock at 36℃for 1 h, while the expressional peakscorresponding to the genes, Pphsp40 and Pphsc70-1 were observed at 39℃.
Under the stress of low temperature, with the exclusion of the Pphsp60 gene, theexpression of the other five Pphsps could be induced. The highest expressional levelsof the five genes were observed at -3℃.
Under the stress of starvation, the expressional levels of the Pphsp40 andPphsc70-2 had no significant changes during starvation. The expression levels ofPphsp20 increased significantly at the beginning 6 h of starvation, and then decreasedgradually to the levels that were significantly lower than those of the control. Pphsp60,Pphsc70-1 and Pphsp90 mRNA levels increased significantly after 24 h of starvation.
Under the stress of Cd~(2+), after 24 h treatment, the Pphsp40 mRNA levels weregradually elevated with the increase of the concentrations. However, the other fivePphsps mRNA levels reached the highest peak at the concentration of 0.5 mM, andthen reduced with the increase of the concentrations. After 60 h treatment, thePphsp40 mRNA levels were also gradually elevated with the increase of theconcentrations. Pphsp60 mRNA levels increased significantly at the concentration of0.5 mM. Pphsc70-1 expression levels increased significantly at the concentration of50 mM. The other three genes mRNA levels had no significant difference compare tothe controls or lower than those of controls.
Under the stress of Cu~(2+), after 24 h treatment, the Pphsp40 and Pphsc70-2mRNA levels were gradually elevated with the increase of the concentrations.However, the other four genes mRNA levels reached a maximum at the concentrationof 50 mM. After 60 h treatment, only Pphsp20 and Pphsp90 mRNA levels had1.32-fold and 1.30-fold, respectively, greater than that of the control at theconcentration of 0.5 mM. However, the other treatments of Pphsps wereapproximately identical with or lower than those of the controls.
In addition, the expression of the Pphsc70-1 gene in the pupae during thermalstress (40℃for 0～8 h) and recovery was also quantified using real-time quantitativePCR (SYBR Green dye). A significantly elevation of Pphsc70-1 expression wasobserved following heat treatment, however, continued exposure to heat shock orrecovery caused the expression of induced mRNA to gradually decline to levels that were significantly lower than those of control pupae.
3 Expression of Pphsps in insect cells and Escherichia coli
The six Pphsps were successfully expressed in the insect cell line Tn-5B1-4 byusing Bac-to-Bac Baculovirus Expression System. By subcellular localizationanalysis, the expressional products of the six Pphsps located in the cytoplasm of theinsect cells.
In addition, the six Pphsps were also expressed in the E. coli strain Bl21 (DE3)for the next functional study on the PpHsps. However, only five genes (except ofPphsp40) were successfully expressed. Further analysis showed that the expressionalproducts of the five Pphsps were dissolved.
4 Purification of PpHsps and their functions under the thermal stress
The effects of overexpressed fusion PpHsps on the survival of E. coli and thestability of E. coli proteins during high-temperature stress were studied. The resultsshowed that the five expressed fusion proteins (PpHsp90, PpHsc70-1, PpHsc70-2,PpHsp60 and PpHsp20) could partly maintain the stability of E. coli proteins up to80℃. However, in functional bioassays (1 h at 50℃) for recombinant PpHsps, only E.coli cells overexpressing PpHsp20 and PpHsp90 proteins had the significantlyincreased survival rate comparing to those lacking PpHsp. Recombinant PpHsp20 andPpHsp90 were purified by Ni-NTA resin. Measuring the light scattering of luciferaserevealed that only PpHsp20 prevented the aggregation of luciferase.
5 Cloning and sequence analysis of wsp, ftsZ and 16S rRNA genes fromWolbachia in the 13 populations of P. puparum in China
The results showed that all the 13 populations were infected with B-Wolbachia.In the 13 populations, we found four distinct sequences for wsp (wPup1, wPup2,wPup3 and wPup4), two distinct sequences for both ftsZ(fPup1 and fPup2) and 16S(16SPup1 and 16SPup2). Moreover, recombination of wsp gene within distinctsequences was found.
In addition, only one sequence was found for wsp, ftsZ and 16S rRNA in the allthe southern populations with the exclusion of Shanghai population. However, two orthree distinct sequences for the three genes were found in the all northern populations.The results indicated that two Wolbachia strains present in the northern populations,whereas only one Wolbachia strain present in the southern populations (exceptShanghai population).
1蝶蛹金小蜂hsps基因的克隆及其序列分析
通过数据库检索、PCR／RT-PCR、RACE和文库筛选等分子手段，共克隆了蝶蛹金小蜂6个热激蛋白成员，涉及了5个热激蛋白家族。
Pphsp90基因ORF长为2148 bp，推测编码716个氨基酸，预测蛋白分子量为82.2 kDa，编码区内不含内含子。Pphsc70-1基因ORF长为1968 bp，推测编码656个氨基酸，预测蛋白分子量为71.3 kDa，该基因含有3个内含子。Pphsc70-2基因ORF长为1923 bp，推测编码641个氨基酸，预测蛋白分子量为70.9 kDa，编码区内含有1个内含子。Pphsp60基因ORF长为1719 bp，推测编码573个氨基酸，预测蛋白分子量为60.6 kDa，编码区内含有3个内含子。Pphsp40基因ORF长为1095 bp，推测编码365个氨基酸，预测蛋白分子量为39.6 kDa。Pphsp20基因ORF长为573 bp，推测编码191个氨基酸，预测蛋白分子量为21.7 kDa。
2不同逆境胁迫对蝶蛹金小蜂体内hsps基因表达的影响
采用TaqMan-MGB探针和特异性引物构建了6个Pphsps基因实时荧光定量RT-PCR检测体系。利用荧光定量RT-PCR方法，检测了蝶蛹金小蜂成虫6个Pphsps基因在高温(30～42℃处理1 h)、低温(3～-15℃处理1 h)、饥饿(0～24 h)及重金属(0.5～5mM Cd~(2+)／Cu~(2+)处理24 h或60 h)等4种逆境胁迫下mRNA的表达情况。
在高温胁迫下，6个Pphsps基因均能被诱导；其中，Pphsp20、Pphsp60、Pphsc70-2和Pphsp90表达量在36℃达到顶峰，而Pphsp40和Pphsc70-1表达量在39℃才达到最高峰。
在低温胁迫下，除Pphsp60基因外，其它5个基因均能被低温胁迫所诱导，并且其表达量均在-3℃时达到最高峰。
在饥饿胁迫下，Pphsp40和Pphsc70-2诱导前后无显著变化；Pphsp20表达量在饥饿6 h时就显著上升，其后随饥饿时间延长，表达量逐渐下降，甚至降至正常水平以下；Pphsp60、Pphsc70-1和Pphsp90表达量均在24 h显著升高。
在重金属Cd~(2+)胁迫下，处理24 h后，Pphsp40表达量随Cd~(2+)浓度的升高而持续升高，而其它5个基因呈先显著上升，其后，随Cd~(2+)浓度的升高，表达量呈下降趋势；处理60 h后，Pphsp40表达量随处理浓度的升高而升高；Pphsp60表达量在Cd~(2+)浓度为0.5 mM时显著上升，而后随Cd~(2+)浓度升高其表达量恢复至对照水平；Pphsc70-1表达量在Cd~(2+)浓度为50 mM时显著上升；其它Pphsps表达量与对照相比无明显差异或显著低于对照水平。
在重金属Cu~(2+)胁迫下，处理24 h后，Pphsp40和Pphsc70-2表达量随处理Cu~(2+)浓度的升高而升高，其它4个基因表达量随处理Cu~(2+)浓度的升高呈先上升后下降趋势；处理60 h后，与对照相比，仅0.5 mM的Cu~(2+)能够诱导Pphsp20和Pphsp90表达量分别上升1.32倍和1.30倍，而大多Pphsps表达量接近于或低于对照水平。
此外，还采用实时定量PCR方法(SYRB Green染料法)检测了蝶蛹金小蜂蛹期Pphsc70-1在40℃热激胁迫0～8 h及恢复1 h或2 h后的表达情况。结果表明，热处理1 h后Pphsc70-1表达量显著上升，然而持续的热激或热激恢复使其诱导的mRNA表达量逐渐降低，甚至显著低于对照水平。
3蝶蛹金小蜂hsps基因在昆虫细胞和原核细胞中表达
采用的Bac-to-Bac杆状病毒表达系统成功地将6个Pphsps基因在昆虫细胞中进行了表达。亚细胞定位分析表明6个基因的表达产物均位于细胞核外的细胞质中。
采用pET28a表达载体将6个Pphsps基因在表达菌株BL21(DE3)中进行原核表达。其中，Pphsp90、Pphsc70-1、Pphsc70-2、Pphsp60和Pphsp20等5个基因在该系统中都得到了不同程度的表达，且它们产生的融合表达蛋白基本上以可溶的形式存在。而Pphsp40基因在该系统中没有表达成功。
4蝶蛹金小蜂Hsps表达产物的纯化及其功能初探
为了研究PpHsps的分子伴侣功能，本实验研究了过表达PpHsps对E．coli细胞在高温下的存活率及E．coli细胞内蛋白热稳定性的影响。结果表明，虽然PpHsp90、PpHsc70-1、PpHsc70-2、PpHsp60和PpHsp20融合蛋白对高温下E．coli细胞内蛋白的稳定性具有一定的保护作用，但仅PpHsp20和PpHsp90对E．coli细胞在高温下的存活有保护作用。因此，我们采用Ni-NTA树脂分别对PpHsp20和PpHsp90融合蛋白进行分离纯化，研究了PpHsp20和PpHsp90对荧光素酶的高温保护能力。结果发现PpHsp20可以在一定程度上抑制高温引起的荧光素酶的聚集现象。
5蝶蛹金小蜂体内Wolbachia wsp、ftsZ和16S RNA基因的克隆及其在不同地理种群中的差异
13地理种群的蝶蛹金小蜂均感染B-Wolbachia。分析从13个地理种群中获得序列，共发现4条wsp基因差异序列(wPup1、wPup2、wPup3和wPup4)、2条ftsZ基因差异序列(fPup1和fPup2)和2条16S rRNA差异序列(16SPup1和16SPup2)；wsp基因在不同Wolbachia株系间存在高度重组。
此外，本研究还发现一个有趣的现象，在蝶蛹金小蜂南方种群中，除上海种群外，仅发现wsp、ftsZ和16S rRNA序列各一条；而在北方种群中，均存在2条或2条以上的wsp、ftsZ和16S rRNA序列。这一结果提示，南方种群(除上海种群)仅感染1个Wolbachia株，而北方种群感染2个Wolbachia株。
Heat shock proteins (Hsps) are a group of conserved proteins in evolution, whichexist in all eukaryotic and prokaryotic organisms. Hsps are very important fororganisms to survive in the stress. Wolbachia are maternally inherited bacteria thatspread widely in arthropods and can induce cytoplasmic incompatibility (CI),parthenogneesis, male killing and feminization. In this study, 6 heat shock genes inthe Pteromalus puparum were cloned and characterized, and their functions were alsostudied. Moreover, an investigation to determine the diversity of Wolbachia in P.puparum from 13 geographically distributed native populations in both southern andnorthern China were undertaken by comparing the sequences of Wolbachia 16S rRNA,ftsZ and wap genes. The results are summarized as follows:
1 Cloning and characterization of the hsps in the P. puparum
Six full-length ORFs coding Hsps involved five Hsp families were cloned by themethods of Blast search, PCR/RT-PCR, RACE and cDNA library screening.
The full-length ORF of the Pphsp90 gene was 2148 bp which was predicted toencode a 716 amino acid peptide with a deduced molecule weight of 82.2 kDa, and nointron was found. The full-length ORF of the Pphsc70-1 gene was 1968 bp which waspredicted to encode a 656 amino acid peptide with a deduced molecule weight of 71.3kDa, and three introns were found. The full-length ORF of the Pphsc70-2 gene was1923 bp which was predicted to encode a 641 amino acid peptide with a deducedmolecule weight of 70.9 kDa, and one intron was found. The full-length ORF of thePphsp60 gene was 1719 bp which was predicted to encode a 573 amino acid peptidewith a deduced molecule weight of 60.6 kDa, and three introns were found. Thefull-length ORF of the Pphsp40 gene was 1095 bp which was predicted to encode a365 amino acid peptide with a deduced molecule weight of 39.6 kDa. The full-lengthORF of the Pphsp20 gene was 573 bp which was predicted to encode a 191 aminoacid peptide with a deduced molecule weight of 21.7 kDa.
2 The Pphsps expression in relation to different stressors
TaqMan-MGB probes and the corresponding primers were designed to constructa system for real-time fluorescence quantitative RT-PCR to detect the changes inexpression levels of the six Pphsps in the adult wasps under the stressors, such as hightemperature (30～42℃for 1h), low temperature (3～15℃for 1h), starvation (0～24h) and heavy metals (0.5～5 mM Cd~(2+)/Cu~(2+) treated 24h or 60h).
Under the stress of high temperature, the expression of six Pphsps could beinduced. The genes, Pphsp20, Pphsp60, Pphsc70-2 and Pphsp90, had highestexpression levels after heat shock at 36℃for 1 h, while the expressional peakscorresponding to the genes, Pphsp40 and Pphsc70-1 were observed at 39℃.
Under the stress of low temperature, with the exclusion of the Pphsp60 gene, theexpression of the other five Pphsps could be induced. The highest expressional levelsof the five genes were observed at -3℃.
Under the stress of starvation, the expressional levels of the Pphsp40 andPphsc70-2 had no significant changes during starvation. The expression levels ofPphsp20 increased significantly at the beginning 6 h of starvation, and then decreasedgradually to the levels that were significantly lower than those of the control. Pphsp60,Pphsc70-1 and Pphsp90 mRNA levels increased significantly after 24 h of starvation.
Under the stress of Cd~(2+), after 24 h treatment, the Pphsp40 mRNA levels weregradually elevated with the increase of the concentrations. However, the other fivePphsps mRNA levels reached the highest peak at the concentration of 0.5 mM, andthen reduced with the increase of the concentrations. After 60 h treatment, thePphsp40 mRNA levels were also gradually elevated with the increase of theconcentrations. Pphsp60 mRNA levels increased significantly at the concentration of0.5 mM. Pphsc70-1 expression levels increased significantly at the concentration of50 mM. The other three genes mRNA levels had no significant difference compare tothe controls or lower than those of controls.
Under the stress of Cu~(2+), after 24 h treatment, the Pphsp40 and Pphsc70-2mRNA levels were gradually elevated with the increase of the concentrations.However, the other four genes mRNA levels reached a maximum at the concentrationof 50 mM. After 60 h treatment, only Pphsp20 and Pphsp90 mRNA levels had1.32-fold and 1.30-fold, respectively, greater than that of the control at theconcentration of 0.5 mM. However, the other treatments of Pphsps wereapproximately identical with or lower than those of the controls.
In addition, the expression of the Pphsc70-1 gene in the pupae during thermalstress (40℃for 0～8 h) and recovery was also quantified using real-time quantitativePCR (SYBR Green dye). A significantly elevation of Pphsc70-1 expression wasobserved following heat treatment, however, continued exposure to heat shock orrecovery caused the expression of induced mRNA to gradually decline to levels that were significantly lower than those of control pupae.
3 Expression of Pphsps in insect cells and Escherichia coli
The six Pphsps were successfully expressed in the insect cell line Tn-5B1-4 byusing Bac-to-Bac Baculovirus Expression System. By subcellular localizationanalysis, the expressional products of the six Pphsps located in the cytoplasm of theinsect cells.
In addition, the six Pphsps were also expressed in the E. coli strain Bl21 (DE3)for the next functional study on the PpHsps. However, only five genes (except ofPphsp40) were successfully expressed. Further analysis showed that the expressionalproducts of the five Pphsps were dissolved.
4 Purification of PpHsps and their functions under the thermal stress
The effects of overexpressed fusion PpHsps on the survival of E. coli and thestability of E. coli proteins during high-temperature stress were studied. The resultsshowed that the five expressed fusion proteins (PpHsp90, PpHsc70-1, PpHsc70-2,PpHsp60 and PpHsp20) could partly maintain the stability of E. coli proteins up to80℃. However, in functional bioassays (1 h at 50℃) for recombinant PpHsps, only E.coli cells overexpressing PpHsp20 and PpHsp90 proteins had the significantlyincreased survival rate comparing to those lacking PpHsp. Recombinant PpHsp20 andPpHsp90 were purified by Ni-NTA resin. Measuring the light scattering of luciferaserevealed that only PpHsp20 prevented the aggregation of luciferase.
5 Cloning and sequence analysis of wsp, ftsZ and 16S rRNA genes fromWolbachia in the 13 populations of P. puparum in China
The results showed that all the 13 populations were infected with B-Wolbachia.In the 13 populations, we found four distinct sequences for wsp (wPup1, wPup2,wPup3 and wPup4), two distinct sequences for both ftsZ(fPup1 and fPup2) and 16S(16SPup1 and 16SPup2). Moreover, recombination of wsp gene within distinctsequences was found.
In addition, only one sequence was found for wsp, ftsZ and 16S rRNA in the allthe southern populations with the exclusion of Shanghai population. However, two orthree distinct sequences for the three genes were found in the all northern populations.The results indicated that two Wolbachia strains present in the northern populations,whereas only one Wolbachia strain present in the southern populations (exceptShanghai population).
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