云斑天牛触角cDNA文库的构建及相关嗅觉蛋白的表达及功能分析
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摘要
自然界中,昆虫对化学信号物质的感知主要通过两种化学感受机制：嗅觉和味觉。借助特定的化学感觉机制,昆虫的嗅觉系统能够感知其周围环境中的大量纷繁复杂的化学信息,完成寄主定位、寻找食物配偶、躲避天敌的捕捉、避免有毒物质以及寻找繁殖场所等多种生理行为。阐明昆虫的化学感受机理,可以加深对昆虫与植物、昆虫与昆虫之间相互关系的了解,帮助我们设计新的方法来进行害虫治理。云斑天牛Batocera horsfieldi (Hope)属于鞘翅目天牛科,是一种重要的蛀干害虫,主要以幼虫蛀食树干和成虫补充营方式养危害树木。由于云斑天牛蛀害,导致杨树树势衰弱或全株枯死；同时还造成了杨树材质的损害,造成极大的经济损失。近年来,对云斑天牛的行为学研究比较多,但对云斑天牛嗅觉识别机制的分子机制研究,国内外尚未见报道。为此我们通过分子生物学手段构建cDNA文库并结合批量ESTs测序,对云斑天牛触角功能基因进行了分析。通过对主要的触角气味结合蛋白OBPs和化学感受蛋白CSPs进行基因克隆,分析其在不同组织中的转录表达谱特征,以及对OBP2和CSP2蛋白的表达纯化,初步探讨了其生理功能。主要研究结果如下：
1云斑天牛成虫触角cDNA文库的构建
通过LD-PCR技术合成全长ds-cDNA,电泳后,对0.3-0.6kb ds-cDNA和0.6-1kbds-cDNA分别割胶回收,连接到pDNR-LIB载体上。对扩增文库分别采用PCR扩增和蓝白斑筛选的方法进行鉴定,本研究构建的云斑天牛0.3-0.6kb cDNA文库滴度为6.4×105cfu/mL,库容量为8.9×105克隆,得重组率94.5%;0.6-1kb cDNA文库滴度为4.1×106cfu/mL,库容量为5.8×106克隆,得重组率93.2%,且序列基本上均为全长,这充分说明两个定向片段大小的文库构建得较好。通过测序,成功得到了有效ESTs序列692条,经拼接后分别得到‘'Contigs"和‘'Singlets"(?)序列分别为79个和323个,其中功能基因246个,同源但无注释功能等的基因34个。从拼接后的BLASTx结果看,包括68条云斑天牛气味结合蛋白及化学感受相关的基因,去除重复的序列,得到了4个Odorant Binding proteins (OBPs)机因,3个Chemosensory Proteins (CSPs)基因,2个Pheromone binding proteins (PBP)基因以及9个Minus-OBP基因,为广泛开展云斑天牛分子生物学研究提供了重要的参考和依据。
2云斑天牛气味结合蛋白和化学感受蛋白的克隆及序列分析
利用RT-PCR技术克隆得到云斑天牛3个普通气味结合蛋白(GOBP)基因BhorGOBP1、BhorGOBP2及BhorGOBP3,分别编码142、137和125个氨基酸,预测N-末端分别包含20,19和20个氨基酸组成的信号肽序列。这三个基因编码的氨基酸序列和鞘翅目昆虫GOBP的氨基酸序列比对同源性较高(50%左右),但是BhorGOBP1、BhorGOBP2和BhorOBP3的同源性只有12.68%,说明它们属于不同的GOBP类群；同时克隆得到3个化学感受蛋白(CSP)基因BhorCSP1、BhorCSP2及BhorCSP3的ORF全长,分别编码127、130和125个氨基酸,预测N-末端分别包含20、19和20个氨基酸组成的信号肽序列,所得到的3个CSP基因序列已提交到GenBank,其登录号分别为HQ587041, HQ587040和HQ587042。从序列连配的结果我们发现,在CSPs中存在有3个高度保守的区域结构,分别是位于N-端的Motif A区YTTKYDN[V/I][N/D][L/V] DEIL,中心的Motif B区DGKELKXX [I/L]PDAL,和位于C-端的Motif C区KYDP,进一步分析表明,位于第53、110、123位置的芬芳族残基,以及位于第91/92位的(glutamine/lysine)和129/130位的(lysine/tyrosine)均高度保守。
3云斑天牛气味结合蛋白和化学感受蛋白的时空表达谱研究
依据已克隆得到的云斑天牛Batocera horsfieldi气味结合蛋白OBPs和化学感受蛋白CSPs基因全长序列设计特异引物,利用RT-PCR和Real Time-PCR技术对云斑天牛主要OBPs基因和CSPs基因在云斑天牛成虫的不同组织及羽化后各个发育阶段的转录表达谱进行了研究。基于云斑天牛成虫期RT-PCR的表达谱分析发现,3个OBP基因和3个CSP基因在头部(去除触角)均不表达,而在触角、下颚须、下唇须、翅、足、腹部均有表达。Q-PCR结果显示3个OBP基因在触角中在不同发育阶段的表达量要远高于3个CSP基因；OBPs和CSPs在雄虫不同组织其表达量要高于雌虫(无论交配与否)；在羽化后1-5天以及羽化后30天左右OBPs和CSPs的表达量较高,远高于中间时期的表达量。结果表明BhorCSPs除了嗅觉功能外,可能还在云斑天牛感受外界化合物的过程中发挥着其它重要作用,其在雌雄虫中不同的表达量及不同时期表达量的不同也表明OBPs和CSPs在雄虫寻找配偶及雌虫寻找补充营养寄主及产卵场所的过程中发挥着重要的功能。
4云斑天牛气味结合蛋白OBP2和化学感受蛋白CSP2的原核表达纯化
成功构建了云斑天牛BhorOBP2和BhorCSPSP2(?)勺原核表达质粒,由于目的蛋白含有二硫键,我们最终选择以pET32a原核表达载体来表达目的蛋白,pET32a融合表达系统含有一个硫氧还蛋白标签(Trx),该标签不仅可以增加蛋白可溶性,而且能够催化二硫键的正确形成,因此非常适合含有二硫键的OBPs(3对)和CSPs(2对)蛋白的表达;同时,我们对pET32a进行了人工改造,将pET32a载体自带的Thrombin酶切位点的识别序列突变掉,然后在目的蛋白基因的起始密码子前面加上一个Thrombin识别序列,同时为了将该序列和目的蛋白保持一定的空间距离,在二者之间还添加了一段Linker序列(GGGGSGGG),最终使切除的标签从分子量和pI均和目的蛋白有一定的差别,利于后续的纯化。通过对温度以及IPTG浓度的优化,我们最终确定了pET32a-BhorOBP2/CSP2的最优的可溶性表达条件：16℃,IPTG浓度0.5mmol/L,诱导过夜。经镍离子亲和层析和Resource S/Q(阳/阴)离子交换柱及Superdex75凝胶柱纯化得到纯的蛋白,为制作多克隆抗体,进行嗅觉蛋白在云斑天牛触角嗅觉感器中的免疫定位和荧光竞争结合实验检测其对不同来源种类的气味化学物质的结合能力奠定了物质基础。
Insects have specialized chemosensory systems detect stimuli from the external environment, including a wide range of chemical compounds. The specialized sensilla in the chemosensory organs (antenna, maxillary palps, labial palps, legs and wings) allows the organism to recognise volatile cues that confer the capacity to detect food sources, predators, mates and oviposition sites. Two major families of soluble proteins, odorant-binding proteins (OBPs) and chemosensory proteins (CSPs), are expressed in chemosensilla of insects and thought to carry lipophilic odorants to the olfactory receptor cells through hydrophilic surroundings and are involved in the initial biochemical step of odour reception. Clarifying the Chemical sense mechanisms of insect not only could be helpful to promote the understanding behavior essence of insects and plants, but also provide theories for controling pest. The long-horned beetle, Batocera horsfieldi (Hope)(Coleoptera:Cerambycidae), is a significant pest of Popolus. The larvae of Batocera horsfieldi cause serious economical loss in result of their eat trunk and covert habitat. In order to further understanding the olfactory mechanism of Batocera horsfieldi and control them by scientific methods, a cDNA library from antenna of B. horsfieldi was constructed and functional from hundreds of expressed sequence tags (ESTs) by patched sequencing were discovered from cDNA library. On the basis of ESTs, OBPs and CSPs of B. horsfieldi were cloned, and the transcriptional profiles in different tissues through different developmental stages were identified, also further recombinant proteins were expressed after construction of expression vectors, meanwhile, the potential function of OBP2was investigated. The main results are as follows:
1The construction of cDNA library with the antenna of B. horsfieldi adults
Through the technique of LD-PCR, ds-cDNA was synthesized, the PCR products were analysed on1%agarose gels, the bands of0.3-0.6kb and0.6-1kb were retrieved respectively and linked into the pDNR-LIB Vector. Through identification of PCR and blue-white selection, the cDNA library was identified high recombinational efficiency, reached above94.5%(hfcbwa) and93.2%(hfcbwb), respectively. And the titer of the two library (hfcbwa and hfcbwb) reached6.4×105pfu/ml and4.1×106pfu/ml.692clones from the cDNA library were selected randomly and sequenced,604high quality sequences more than100bp were produced after removing the vector sequences and contaminated sequences, And the average length of the ESTs (without vector) were386.33bp and468.62bp. Using the Phrap software,79"Contigs" and323"Singlets" were produced from the assembled ESTs, after analyzed by clustering and assemblying, get246functional gene. Compared with NCBI non-redundant protein database using BLASTx,68ESTs of Unigenes matched with olfactory related protein genes, wiped off the repeated ESTs, got4odorant binding proteins,3Chemosensory proteins,2Pheromone binding proteins and9Minus-C odorant binding proteins, which provided foundation for next research.
2Molecular cloning and characterization of OBPs and CSPs from B. horsfieldi adults
From the B. horsfieldi antennal cDNA library, we obtained three full-length odorant-binding proteins gene by RT-PCR, from the nuclear acid sequences, all of the three BhorOBPs have6conservative cysteines, and a signal peptide which about20amino acids at N-terminus. The multiple sequences alignment of the BhorOBPs with other Coleoptera OBPs showed high sequence identity (50%) with other full-length sequences from GenBank, but the homology of these three BhorOBPs were very low, only12.68%, suggested they belong to different groups, also the results in compliance with the results of phylogenetic tree. Also three full-length cDNAs of CSPs were identified and predicted to encode small proteins (120-130amino acids). The sequences of these genes have been deposited in GenBank under the accession numbers HQ587040, HQ587041and HQ587042. Three key features of the amino acid sequences of the candidate CSP genes were consistent with the CSP family members of other insects:a predicted molecular weight of about13kDa, a hydrophobic N-terminus of about16amino acids encoding a signal peptide and the presence of a highly conserved four-cysteine motif (C-X6-C-X18-C-X2-C). The alignment results also shows the CSPs had three conserved several sequence motifs, including a commonmotif A at their N terminal YTTKYDN[V/I][N/D][L/V]DEIL, central motif B DGKELKXX[L/I]PDAL and C-terminal KYDP, in addition, aromatic residues at position53,110and123that may be functionally important are also highly conserved, along with residues located at91/92(glutamine/lysine) and129/130(lysine/tyrosine).
3Temporal and spatial expression profiles of the BhorOBPs and BhorCSPs genes
Using gene-specific BhorOBPs and BhorCSPs primers and control18s rRNA primers with cDNA prepared from different tissues, we examined the expression patterns at successive developmental stages and in various adult body parts and tissues by RT-PCR and Real time-PCR. Unlike other species, all three B. horsfieldi OBPs and CSPs transcripts were detected in antenna, labial palps, maxillary palps, forelegs, midlegs, mid abdomen and hind abdomen, but no specific products were observed in head (without antenna). On the basis of the normalized relative quantification2-ΔΔCT method for real-time Q-PCR, the transcription profiling of the tissues and developmental phases of the three BhorOBPs and BhorCSPs were obtained. The data presented the transcription levels of the three BhorOBPs were higher than those at the same periods of BhorCSPs in antenna. In general, BhorOBPs and BhorCSPs had the highest expression in antenna, and higher espression in labial palps and maxillary palps, but less in wings, legs and abdomen. The transcript levels of the BhorOBPs and BhorCSPs in all tissues of unmated or mated males were significantly higher than in unmated or mated females in most development stages except in the labial palps and maxillary palps. This is most noticeable for BhorCSPs that were generally expressed highly in the wings and legs. In the different developmental periods, BhorOBPs and BhorCSPs had two highest expression leveral periods, from larva to5day adult and around25/30day adult. These data indicated BhorCSPs maybe had other rols except olfactory sensation, also the difference transcription leveral at different developmental period and in different tissues suggest the BhorOBPs and BhorCSPs have their specifically task in detecting food sources, mates and oviposition sites.
4The expression and purification of BhorOBP2and BhorCSP2proteins
The ORF sequence of BhorOBP2and BhorCSP2were subcloned into prokaryotic vector pET32a, transferred into BL21(DE3) and expressed. The recombinant plasmids pET32a have a His-tag and a Trx-tag, the Trx-tag can increase the solubility of the protein, and facilitate forming correct disulfide bond. We deleted the Thrombin sequence of pET32a vector, then added a Thrombin sequence and a linker at the N terminal of BhorOBP2and BhorCSP2. The conditions for soluble expression was optimized by investigating different temperature (16℃), IPTG concentration (0.5mmol/L) and induction time (overnight). Then the buffer condition and chromatography methods were discussed of the proteins. Finally, through nickel affinity column, Resource Q/S ion exchange column and Superdex75gel filtration chromatography, the protein samples with high concentrations and stability were obtained.
1云斑天牛成虫触角cDNA文库的构建
通过LD-PCR技术合成全长ds-cDNA,电泳后,对0.3-0.6kb ds-cDNA和0.6-1kbds-cDNA分别割胶回收,连接到pDNR-LIB载体上。对扩增文库分别采用PCR扩增和蓝白斑筛选的方法进行鉴定,本研究构建的云斑天牛0.3-0.6kb cDNA文库滴度为6.4×105cfu/mL,库容量为8.9×105克隆,得重组率94.5%;0.6-1kb cDNA文库滴度为4.1×106cfu/mL,库容量为5.8×106克隆,得重组率93.2%,且序列基本上均为全长,这充分说明两个定向片段大小的文库构建得较好。通过测序,成功得到了有效ESTs序列692条,经拼接后分别得到‘'Contigs"和‘'Singlets"(?)序列分别为79个和323个,其中功能基因246个,同源但无注释功能等的基因34个。从拼接后的BLASTx结果看,包括68条云斑天牛气味结合蛋白及化学感受相关的基因,去除重复的序列,得到了4个Odorant Binding proteins (OBPs)机因,3个Chemosensory Proteins (CSPs)基因,2个Pheromone binding proteins (PBP)基因以及9个Minus-OBP基因,为广泛开展云斑天牛分子生物学研究提供了重要的参考和依据。
2云斑天牛气味结合蛋白和化学感受蛋白的克隆及序列分析
利用RT-PCR技术克隆得到云斑天牛3个普通气味结合蛋白(GOBP)基因BhorGOBP1、BhorGOBP2及BhorGOBP3,分别编码142、137和125个氨基酸,预测N-末端分别包含20,19和20个氨基酸组成的信号肽序列。这三个基因编码的氨基酸序列和鞘翅目昆虫GOBP的氨基酸序列比对同源性较高(50%左右),但是BhorGOBP1、BhorGOBP2和BhorOBP3的同源性只有12.68%,说明它们属于不同的GOBP类群；同时克隆得到3个化学感受蛋白(CSP)基因BhorCSP1、BhorCSP2及BhorCSP3的ORF全长,分别编码127、130和125个氨基酸,预测N-末端分别包含20、19和20个氨基酸组成的信号肽序列,所得到的3个CSP基因序列已提交到GenBank,其登录号分别为HQ587041, HQ587040和HQ587042。从序列连配的结果我们发现,在CSPs中存在有3个高度保守的区域结构,分别是位于N-端的Motif A区YTTKYDN[V/I][N/D][L/V] DEIL,中心的Motif B区DGKELKXX [I/L]PDAL,和位于C-端的Motif C区KYDP,进一步分析表明,位于第53、110、123位置的芬芳族残基,以及位于第91/92位的(glutamine/lysine)和129/130位的(lysine/tyrosine)均高度保守。
3云斑天牛气味结合蛋白和化学感受蛋白的时空表达谱研究
依据已克隆得到的云斑天牛Batocera horsfieldi气味结合蛋白OBPs和化学感受蛋白CSPs基因全长序列设计特异引物,利用RT-PCR和Real Time-PCR技术对云斑天牛主要OBPs基因和CSPs基因在云斑天牛成虫的不同组织及羽化后各个发育阶段的转录表达谱进行了研究。基于云斑天牛成虫期RT-PCR的表达谱分析发现,3个OBP基因和3个CSP基因在头部(去除触角)均不表达,而在触角、下颚须、下唇须、翅、足、腹部均有表达。Q-PCR结果显示3个OBP基因在触角中在不同发育阶段的表达量要远高于3个CSP基因；OBPs和CSPs在雄虫不同组织其表达量要高于雌虫(无论交配与否)；在羽化后1-5天以及羽化后30天左右OBPs和CSPs的表达量较高,远高于中间时期的表达量。结果表明BhorCSPs除了嗅觉功能外,可能还在云斑天牛感受外界化合物的过程中发挥着其它重要作用,其在雌雄虫中不同的表达量及不同时期表达量的不同也表明OBPs和CSPs在雄虫寻找配偶及雌虫寻找补充营养寄主及产卵场所的过程中发挥着重要的功能。
4云斑天牛气味结合蛋白OBP2和化学感受蛋白CSP2的原核表达纯化
成功构建了云斑天牛BhorOBP2和BhorCSPSP2(?)勺原核表达质粒,由于目的蛋白含有二硫键,我们最终选择以pET32a原核表达载体来表达目的蛋白,pET32a融合表达系统含有一个硫氧还蛋白标签(Trx),该标签不仅可以增加蛋白可溶性,而且能够催化二硫键的正确形成,因此非常适合含有二硫键的OBPs(3对)和CSPs(2对)蛋白的表达;同时,我们对pET32a进行了人工改造,将pET32a载体自带的Thrombin酶切位点的识别序列突变掉,然后在目的蛋白基因的起始密码子前面加上一个Thrombin识别序列,同时为了将该序列和目的蛋白保持一定的空间距离,在二者之间还添加了一段Linker序列(GGGGSGGG),最终使切除的标签从分子量和pI均和目的蛋白有一定的差别,利于后续的纯化。通过对温度以及IPTG浓度的优化,我们最终确定了pET32a-BhorOBP2/CSP2的最优的可溶性表达条件：16℃,IPTG浓度0.5mmol/L,诱导过夜。经镍离子亲和层析和Resource S/Q(阳/阴)离子交换柱及Superdex75凝胶柱纯化得到纯的蛋白,为制作多克隆抗体,进行嗅觉蛋白在云斑天牛触角嗅觉感器中的免疫定位和荧光竞争结合实验检测其对不同来源种类的气味化学物质的结合能力奠定了物质基础。
Insects have specialized chemosensory systems detect stimuli from the external environment, including a wide range of chemical compounds. The specialized sensilla in the chemosensory organs (antenna, maxillary palps, labial palps, legs and wings) allows the organism to recognise volatile cues that confer the capacity to detect food sources, predators, mates and oviposition sites. Two major families of soluble proteins, odorant-binding proteins (OBPs) and chemosensory proteins (CSPs), are expressed in chemosensilla of insects and thought to carry lipophilic odorants to the olfactory receptor cells through hydrophilic surroundings and are involved in the initial biochemical step of odour reception. Clarifying the Chemical sense mechanisms of insect not only could be helpful to promote the understanding behavior essence of insects and plants, but also provide theories for controling pest. The long-horned beetle, Batocera horsfieldi (Hope)(Coleoptera:Cerambycidae), is a significant pest of Popolus. The larvae of Batocera horsfieldi cause serious economical loss in result of their eat trunk and covert habitat. In order to further understanding the olfactory mechanism of Batocera horsfieldi and control them by scientific methods, a cDNA library from antenna of B. horsfieldi was constructed and functional from hundreds of expressed sequence tags (ESTs) by patched sequencing were discovered from cDNA library. On the basis of ESTs, OBPs and CSPs of B. horsfieldi were cloned, and the transcriptional profiles in different tissues through different developmental stages were identified, also further recombinant proteins were expressed after construction of expression vectors, meanwhile, the potential function of OBP2was investigated. The main results are as follows:
1The construction of cDNA library with the antenna of B. horsfieldi adults
Through the technique of LD-PCR, ds-cDNA was synthesized, the PCR products were analysed on1%agarose gels, the bands of0.3-0.6kb and0.6-1kb were retrieved respectively and linked into the pDNR-LIB Vector. Through identification of PCR and blue-white selection, the cDNA library was identified high recombinational efficiency, reached above94.5%(hfcbwa) and93.2%(hfcbwb), respectively. And the titer of the two library (hfcbwa and hfcbwb) reached6.4×105pfu/ml and4.1×106pfu/ml.692clones from the cDNA library were selected randomly and sequenced,604high quality sequences more than100bp were produced after removing the vector sequences and contaminated sequences, And the average length of the ESTs (without vector) were386.33bp and468.62bp. Using the Phrap software,79"Contigs" and323"Singlets" were produced from the assembled ESTs, after analyzed by clustering and assemblying, get246functional gene. Compared with NCBI non-redundant protein database using BLASTx,68ESTs of Unigenes matched with olfactory related protein genes, wiped off the repeated ESTs, got4odorant binding proteins,3Chemosensory proteins,2Pheromone binding proteins and9Minus-C odorant binding proteins, which provided foundation for next research.
2Molecular cloning and characterization of OBPs and CSPs from B. horsfieldi adults
From the B. horsfieldi antennal cDNA library, we obtained three full-length odorant-binding proteins gene by RT-PCR, from the nuclear acid sequences, all of the three BhorOBPs have6conservative cysteines, and a signal peptide which about20amino acids at N-terminus. The multiple sequences alignment of the BhorOBPs with other Coleoptera OBPs showed high sequence identity (50%) with other full-length sequences from GenBank, but the homology of these three BhorOBPs were very low, only12.68%, suggested they belong to different groups, also the results in compliance with the results of phylogenetic tree. Also three full-length cDNAs of CSPs were identified and predicted to encode small proteins (120-130amino acids). The sequences of these genes have been deposited in GenBank under the accession numbers HQ587040, HQ587041and HQ587042. Three key features of the amino acid sequences of the candidate CSP genes were consistent with the CSP family members of other insects:a predicted molecular weight of about13kDa, a hydrophobic N-terminus of about16amino acids encoding a signal peptide and the presence of a highly conserved four-cysteine motif (C-X6-C-X18-C-X2-C). The alignment results also shows the CSPs had three conserved several sequence motifs, including a commonmotif A at their N terminal YTTKYDN[V/I][N/D][L/V]DEIL, central motif B DGKELKXX[L/I]PDAL and C-terminal KYDP, in addition, aromatic residues at position53,110and123that may be functionally important are also highly conserved, along with residues located at91/92(glutamine/lysine) and129/130(lysine/tyrosine).
3Temporal and spatial expression profiles of the BhorOBPs and BhorCSPs genes
Using gene-specific BhorOBPs and BhorCSPs primers and control18s rRNA primers with cDNA prepared from different tissues, we examined the expression patterns at successive developmental stages and in various adult body parts and tissues by RT-PCR and Real time-PCR. Unlike other species, all three B. horsfieldi OBPs and CSPs transcripts were detected in antenna, labial palps, maxillary palps, forelegs, midlegs, mid abdomen and hind abdomen, but no specific products were observed in head (without antenna). On the basis of the normalized relative quantification2-ΔΔCT method for real-time Q-PCR, the transcription profiling of the tissues and developmental phases of the three BhorOBPs and BhorCSPs were obtained. The data presented the transcription levels of the three BhorOBPs were higher than those at the same periods of BhorCSPs in antenna. In general, BhorOBPs and BhorCSPs had the highest expression in antenna, and higher espression in labial palps and maxillary palps, but less in wings, legs and abdomen. The transcript levels of the BhorOBPs and BhorCSPs in all tissues of unmated or mated males were significantly higher than in unmated or mated females in most development stages except in the labial palps and maxillary palps. This is most noticeable for BhorCSPs that were generally expressed highly in the wings and legs. In the different developmental periods, BhorOBPs and BhorCSPs had two highest expression leveral periods, from larva to5day adult and around25/30day adult. These data indicated BhorCSPs maybe had other rols except olfactory sensation, also the difference transcription leveral at different developmental period and in different tissues suggest the BhorOBPs and BhorCSPs have their specifically task in detecting food sources, mates and oviposition sites.
4The expression and purification of BhorOBP2and BhorCSP2proteins
The ORF sequence of BhorOBP2and BhorCSP2were subcloned into prokaryotic vector pET32a, transferred into BL21(DE3) and expressed. The recombinant plasmids pET32a have a His-tag and a Trx-tag, the Trx-tag can increase the solubility of the protein, and facilitate forming correct disulfide bond. We deleted the Thrombin sequence of pET32a vector, then added a Thrombin sequence and a linker at the N terminal of BhorOBP2and BhorCSP2. The conditions for soluble expression was optimized by investigating different temperature (16℃), IPTG concentration (0.5mmol/L) and induction time (overnight). Then the buffer condition and chromatography methods were discussed of the proteins. Finally, through nickel affinity column, Resource Q/S ion exchange column and Superdex75gel filtration chromatography, the protein samples with high concentrations and stability were obtained.
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