苜蓿夜蛾和大豆食心虫气味感受相关蛋白基因的克隆分析及表达
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摘要
苜蓿夜蛾(鳞翅目：夜蛾科)和大豆食心虫(鳞翅目：小卷蛾科)是大豆生长期和结荚期的两种主要害虫,随着近年来免耕和重迎茬面积增加,两种害虫的发生明显加重,成为造成大豆品质和产量降低的主要因素之一。昆虫触角是嗅觉感器分布集中的部位,触角上的各种感器调节着昆虫行为与化学、物理等各种环境刺激因子的关系,在昆虫的寄主定位、识别、取食、觅偶、交配、繁殖、栖息、防御与迁移等过程中起着极为重要的作用。因此,研究其嗅觉感器的形态与结构是探索昆虫嗅觉行为和识别机制的必要前提。昆虫的嗅觉识别过程是非常复杂的,多种蛋白参与了这一过程,这些蛋白主要包括气味结合蛋白、气味受体以及气味降解酶。深入研究嗅觉识别过程中的几个主要蛋白的结构与功能对于阐明嗅觉识别的分子机理非常重要。因此,本试验以苜蓿夜蛾和大豆食心虫为研究对象,利用扫描电镜对触角的超微结构进行了观察,利用RT-PCR和RACE的方法克隆得到了嗅觉识别的相关基因,并对相关基因的功能进行了分析,以期为阐明这两种害虫嗅觉识别的分子机理提供依据。
利用扫描电镜对苜蓿夜蛾触角上的感受器类型进行观察,结果表明,雌雄苜蓿夜蛾的触角类型都是线状触角,由柄节、梗节和鞭节组成。触角上有8种感器分布,分别为毛形感器、刺形感器、锥形感器、腔锥形感器、耳形感器、鳞形感器、栓锥形感器和Bohm氏鬃毛,触角感器主要分布于触角的迎风面和侧面,柄节周围被鳞片包被,鞭节背风面有半包围状整齐排列的鳞片,鳞片结合处有零星分布的磷形感器、刺形感器；在整个触角上,感器主要集中在鞭节,柄节和梗节较少。不同类型感器相比而言,以毛形感器数量最多,刺形感器排列较为整齐和规律。每一鞭小节背风面都覆盖有鳞片,后排鳞片的前缘盖着前排鳞片的基部,层层相叠,绝大多数感器着生在触角的迎风面和侧面。
利用RT-PCR和RACE的方法得到了苜蓿夜蛾和大豆食心虫的五个气味结合蛋白(OBP)和一个气味受体蛋白(Or83b)基因的全长,其中包括两种昆虫的信息素结合蛋白(PBP)和普通气味结合蛋白1(GOBP1),苜蓿夜蛾的普通气味结合蛋白2(GOBP2)和受体蛋白Or83b蛋白。用苜蓿夜蛾的信息素结合蛋白、普通气味结合蛋自1和2三条序列构建了三个基因的原核表达载体,并且在大肠杆菌中成功进行了表达。
苜蓿夜蛾PBP基因cDNA序列的全长为625个碱基,包括一个510个碱基的开放读码框,编码170个氨基酸组成的多肽,推导出的分子量为19.0kD,预测等电点pI为5.65。在27-28位为信号肽切割位点,并存在1个N位糖基化位点。大豆食心虫PBP基因cDNA序列的全长为610个碱基,包括一个504个碱基的开放读码框,编码168个氨基酸组成的多肽,推导出的分子量为19.3kD,预测等电点pI为5.55。在24-25位为信号肽切割位点,并在52的位置上有一个N位糖基化位点。通过和已知昆虫PBP基因cDNA序列的对比,发现两种昆虫的PBP基因氨基酸序列有典型OBP家族特点的六个保守的半胱氨酸,苜蓿夜蛾和大豆食心虫PBP与其他已知鳞翅目昆虫PBP的同源性分别在44%-96%和45%-70%之间,对比过程中发现在已研究昆虫PBPs的研究发现多种昆虫中存在三种PBPs,也就是PBP存在不同的亚型,多种PBP存在的特征可能与昆虫间信息素成分的多样性有关。
苜蓿夜蛾GOBP1基因cDNA序列的全长为973个碱基,包括一个495个碱基的开放读码框,编码164个氨基酸组成的多肽,推导出的分子量为19.0kD,预测等电点pI为5.32。在19-20位为信号肽切割位点。大豆食心虫GOBP1基因cDNA序列的全长为920个碱基,包括一个492个碱基的开放读码框编码163个氨基酸组成的多肽,推导出的分子量为18.8kD,预测等电点pI为5.29。在19-20位为信号肽切割位点。通过和已知昆虫GOBP1基因的cDNA序列对比,发现两种昆虫的GOBP1基因氨基酸序列有典型OBP家族特点的六个保守的半胱氨酸,苜蓿夜蛾和大豆食心虫GOBP1与其他已知鳞翅目昆虫GOBP1的同源性分别在63%-95%和65%-82%之间。
苜蓿夜蛾GOBP2基因cDNA序列全长为624个碱基,包括一个489个碱基的开放读码框,编码162个氨基酸组成的多肽,推导出的分子量为18.2kD,预测等电点pI为5.30。在21-22位为信号肽切割位点。通过和已知昆虫GOBP2基因的cDNA序列对比,发现有典型OBP家族特点的六个保守的半胱氨酸,与已知鳞翅目昆虫GOBP2的同源性在75%-99%,说明GOBP2基因在鳞翅目昆虫中比较保守。
苜蓿夜蛾Or83b基因cDNA序列全长1715个碱基,包括一个1422个碱基的开放读码框,编码473个氨基酸,分子量约为53.5kD,多肽的预测等电点pI为8.37。对苜蓿夜蛾Or83b氨基酸序列进行分析预测,发现苜蓿夜蛾Or83b氨基酸序列中存在2个N位糖基化位点,N-端无信号肽,含有7个跨膜区域,N末端在细胞膜内,而C末端在细胞膜外,符合昆虫气味受体的结构特征。通过和已知昆虫的Or83b基因的cDNA序列对比,发现苜蓿夜蛾的Or83b和其他鳞翅目昆虫Or83b同源性达到96%以上。
对五条气味结合蛋白的生物信息学分析,发现这五条基因编码的蛋白具有昆虫气味结合蛋白的典型特征：小分子量、水溶性酸性蛋白；序列中有6个保守的半胱氨酸,N-末端有信号肽,具有分泌蛋白的特征。苜蓿夜蛾气味受体蛋白Or83b具有昆虫气味受体的典型特征：具有七个跨膜区,N末端在细胞膜内,C末端在细胞膜外,N-无信号肽。
对苜蓿夜蛾PBP、GOBP1、GOBP2和Or83b四条基因在雌雄成虫的触角、腹部和足三个器官的表达情况进行了研究,结果表明四条基因在雌雄触角、腹部和足中均有表达,同时通过对苜蓿夜蛾PBP各器官荧光定量分析,发现苜蓿夜蛾PBP主要存在于雄虫触角中,在雄虫触角中表达量是雌虫的30倍以上,而雌虫触角中的表达量也是其他器官的3倍以上。
Heliothis viriplaca (Lepidoptera:Noctuoidea) and Leguminivora glycinivorella (Lepidoptera: Olethreutidae) are important Lepidopteran pests for soybean in grow and pod period. Recently, the two pests more and more serious because of the increase area field of the free cultivate and plant soybean for several years. They are one of the main reasons of the quality and output decline. Sensillae is focus on antennae, and measure relation between the insect behavior and the environment stimulate factor of chemic and physical. It is important to insect to find host plant, food, mate and pertect prey and migrate. Therefore, the morphology and configuration is precondition of explore insect olfactory behavior and identify system. Insect olfactory identify process is very complex and many protein participate in. They are odorant binding protein, odorant receptor and odorant degrading enzymes. The proteins are important to elucidate the olfactory identify molecular mechanism. In this paper, the antenna of H viriplaca was observed by scanning electron microscopy, and use RT-PCR and RACE method to clone and analysis the olfactory related genes form antenna of H viriplaca and L glycinivorella. The Primary results are as follows:
The antenna of H viriplaca was observed by scanning electron microscopy. Female and male moths are filiform and consist of the scape, pedicel and flagellum, and eight types of sensillawere found. These were sensilla trichodea, sensilla chaetica, sensilla basiconca, sensilla coelocomica, Sensilla auricillica, sensillum squamiformia, sensilla styloconica and Bohm's bristles. The Sensillae distributed the rear and side of antenna, the scape was wrapped by squama, half of the flagellum was wrapped in order by squama, and a few of sensillum squamiformia and sensilla chaetica distributed in the squamas. Sensillae focus on flagellum, only a few Sensillae in scape and pedicel. Sensilla trichodea was the most sensillae, and sensilla chaetica was orderly on the antenna. The every little flagellum rear coverd by squamas, so most of the sensillae was in the rear and side of antenna.
Five OBPs and one OR genes were cloned from the antenna of H viriplaca and L glycinivorella by RT-PCR and RACE method. They are PBP and GOBP1of H viriplaca and L glycinivorella, GOBP2and Or83b of H viriplaca, and the PBP, GOBP1and GOBP2with vector pet21b were construeted and successed expressed in E. coll induced by IPTG.
The H viriplaca PBP cDNA was625bp in length and contained an open reading frame of510bp. The ORF encoded a polypeptide of170amino acid residues with a predicted molecular weight of19.0kD and a p1of5.65. A signal peptide of27residues was predicted at the N-terminus of the protein, and only one putative N-glycosylation sites was identified at the position34. The L. glycinivorella PBP cDNA was610bp in length and contained an open reading frame of504bp. The ORF encoded a polypeptide of168amino acid residues with a predicted molecular weight of19.3kD and a p1of5.55. A signal peptide of24residues was predicted at the N-terminus of the protein, and only one putative N-glycosylation sites was identified at the position52. Sequence blasted and analysed with other known insects indicated that HvirPBP and LglyPBP were characterized by six conservative Cys, which shared typical feature of OBP family. Identity of HvirPBP and LglyPBP were44%-96%and45%-70%with other known Lepidoptera insects. The kown insects had three subfamilies, and it may be connect with diversity pheromone.
The H viriplaca GOBP1cDNA was973bp in length and contained an open reading frame of495bp. The ORF encoded a polypeptide of164amino acid residues with a predicted molecular weight of19.0kD and a p1of5.32. A signal peptide of19residues was predicted at the N-terminus of the protein. The L. glycinivorella GOBP1cDNA was920bp in length and contained an open reading frame of492bp. The ORF encoded a polypeptide of163amino acid residues with a predicted molecular weight of18.8kD and a p1of5.29. A signal peptide of19residues was predicted at the N-terminus of the protein. Sequence blasted and analysed with other known insects indicated that HvirGOBP1and LglyGOBP1were characterized by six conservative Cys, which shared typical feature of OBP family. Identity of HvirGOBP1and Lgly GOBP1were65%-95%and65%-82%with other known Lepidoptera insects.
The H viriplaca GOBP2cDNA was624bp in length and contained an open reading frame of489bp. The ORF encoded a polypeptide of162amino acid residues with a predicted molecular weight of18.2kD and a pi of5.30. A signal peptide of21residues was predicted at the N-terminus of the protein. Sequence blasted and analysed with other known insects indicated that HvirGOBP2was characterized by six conservative Cys, which shared typical feature of OBP family. Identity of HvirGOBP2was well conserved, and between75%and99%with other known Lepidoptera insects.
The H. viriplaca Or83b cDNA was1715bp in length and contained an open reading frame of1422bp. The ORF encoded a polypeptide of473amino acid residues with a predicted molecular weight of53.5kD and a p1of8.37. Two putative N-glycosylation sites were identified at the position110and170, and no signal peptide at the N-terminus of the protein. Sequence blasted found the sequence of H. viriplaca Or83b was well conserved, and over96%identity with Or83bs of other Lepidoptera moths. Analysed the sequence indicated that HvirOr83b has seven transmembrane domains, which shared typical feature of odorant receptor family.
Analysed the five odorant binding proteins found they had the odorant binding protein typical feature, included small molecular weight, water-solubility acidic protein; six conservative Cys in sequences, A signal peptide was predicted at the N-terminus of the protein which secretory protein character. HvirOr83b has seven transmembrane domains, which shared typical feature of odorant receptor, and no signal peptide at the N-terminus of the protein.
PBP, GOBP1, GOBP2and Or83b four genes expressed in different organs in H. viriplaca, the four genes not only expressed in the antennae of both male and female adults but also expressed in abdomen and legs. Analysed PBP in different organs by RT-PCR method found that PBP gene is main expressed in male antenna, it was30times higher than PBP gene in male antenna, and PBP gene in male antenna was3times higher than PBP other organs.
利用扫描电镜对苜蓿夜蛾触角上的感受器类型进行观察,结果表明,雌雄苜蓿夜蛾的触角类型都是线状触角,由柄节、梗节和鞭节组成。触角上有8种感器分布,分别为毛形感器、刺形感器、锥形感器、腔锥形感器、耳形感器、鳞形感器、栓锥形感器和Bohm氏鬃毛,触角感器主要分布于触角的迎风面和侧面,柄节周围被鳞片包被,鞭节背风面有半包围状整齐排列的鳞片,鳞片结合处有零星分布的磷形感器、刺形感器；在整个触角上,感器主要集中在鞭节,柄节和梗节较少。不同类型感器相比而言,以毛形感器数量最多,刺形感器排列较为整齐和规律。每一鞭小节背风面都覆盖有鳞片,后排鳞片的前缘盖着前排鳞片的基部,层层相叠,绝大多数感器着生在触角的迎风面和侧面。
利用RT-PCR和RACE的方法得到了苜蓿夜蛾和大豆食心虫的五个气味结合蛋白(OBP)和一个气味受体蛋白(Or83b)基因的全长,其中包括两种昆虫的信息素结合蛋白(PBP)和普通气味结合蛋白1(GOBP1),苜蓿夜蛾的普通气味结合蛋白2(GOBP2)和受体蛋白Or83b蛋白。用苜蓿夜蛾的信息素结合蛋白、普通气味结合蛋自1和2三条序列构建了三个基因的原核表达载体,并且在大肠杆菌中成功进行了表达。
苜蓿夜蛾PBP基因cDNA序列的全长为625个碱基,包括一个510个碱基的开放读码框,编码170个氨基酸组成的多肽,推导出的分子量为19.0kD,预测等电点pI为5.65。在27-28位为信号肽切割位点,并存在1个N位糖基化位点。大豆食心虫PBP基因cDNA序列的全长为610个碱基,包括一个504个碱基的开放读码框,编码168个氨基酸组成的多肽,推导出的分子量为19.3kD,预测等电点pI为5.55。在24-25位为信号肽切割位点,并在52的位置上有一个N位糖基化位点。通过和已知昆虫PBP基因cDNA序列的对比,发现两种昆虫的PBP基因氨基酸序列有典型OBP家族特点的六个保守的半胱氨酸,苜蓿夜蛾和大豆食心虫PBP与其他已知鳞翅目昆虫PBP的同源性分别在44%-96%和45%-70%之间,对比过程中发现在已研究昆虫PBPs的研究发现多种昆虫中存在三种PBPs,也就是PBP存在不同的亚型,多种PBP存在的特征可能与昆虫间信息素成分的多样性有关。
苜蓿夜蛾GOBP1基因cDNA序列的全长为973个碱基,包括一个495个碱基的开放读码框,编码164个氨基酸组成的多肽,推导出的分子量为19.0kD,预测等电点pI为5.32。在19-20位为信号肽切割位点。大豆食心虫GOBP1基因cDNA序列的全长为920个碱基,包括一个492个碱基的开放读码框编码163个氨基酸组成的多肽,推导出的分子量为18.8kD,预测等电点pI为5.29。在19-20位为信号肽切割位点。通过和已知昆虫GOBP1基因的cDNA序列对比,发现两种昆虫的GOBP1基因氨基酸序列有典型OBP家族特点的六个保守的半胱氨酸,苜蓿夜蛾和大豆食心虫GOBP1与其他已知鳞翅目昆虫GOBP1的同源性分别在63%-95%和65%-82%之间。
苜蓿夜蛾GOBP2基因cDNA序列全长为624个碱基,包括一个489个碱基的开放读码框,编码162个氨基酸组成的多肽,推导出的分子量为18.2kD,预测等电点pI为5.30。在21-22位为信号肽切割位点。通过和已知昆虫GOBP2基因的cDNA序列对比,发现有典型OBP家族特点的六个保守的半胱氨酸,与已知鳞翅目昆虫GOBP2的同源性在75%-99%,说明GOBP2基因在鳞翅目昆虫中比较保守。
苜蓿夜蛾Or83b基因cDNA序列全长1715个碱基,包括一个1422个碱基的开放读码框,编码473个氨基酸,分子量约为53.5kD,多肽的预测等电点pI为8.37。对苜蓿夜蛾Or83b氨基酸序列进行分析预测,发现苜蓿夜蛾Or83b氨基酸序列中存在2个N位糖基化位点,N-端无信号肽,含有7个跨膜区域,N末端在细胞膜内,而C末端在细胞膜外,符合昆虫气味受体的结构特征。通过和已知昆虫的Or83b基因的cDNA序列对比,发现苜蓿夜蛾的Or83b和其他鳞翅目昆虫Or83b同源性达到96%以上。
对五条气味结合蛋白的生物信息学分析,发现这五条基因编码的蛋白具有昆虫气味结合蛋白的典型特征：小分子量、水溶性酸性蛋白；序列中有6个保守的半胱氨酸,N-末端有信号肽,具有分泌蛋白的特征。苜蓿夜蛾气味受体蛋白Or83b具有昆虫气味受体的典型特征：具有七个跨膜区,N末端在细胞膜内,C末端在细胞膜外,N-无信号肽。
对苜蓿夜蛾PBP、GOBP1、GOBP2和Or83b四条基因在雌雄成虫的触角、腹部和足三个器官的表达情况进行了研究,结果表明四条基因在雌雄触角、腹部和足中均有表达,同时通过对苜蓿夜蛾PBP各器官荧光定量分析,发现苜蓿夜蛾PBP主要存在于雄虫触角中,在雄虫触角中表达量是雌虫的30倍以上,而雌虫触角中的表达量也是其他器官的3倍以上。
Heliothis viriplaca (Lepidoptera:Noctuoidea) and Leguminivora glycinivorella (Lepidoptera: Olethreutidae) are important Lepidopteran pests for soybean in grow and pod period. Recently, the two pests more and more serious because of the increase area field of the free cultivate and plant soybean for several years. They are one of the main reasons of the quality and output decline. Sensillae is focus on antennae, and measure relation between the insect behavior and the environment stimulate factor of chemic and physical. It is important to insect to find host plant, food, mate and pertect prey and migrate. Therefore, the morphology and configuration is precondition of explore insect olfactory behavior and identify system. Insect olfactory identify process is very complex and many protein participate in. They are odorant binding protein, odorant receptor and odorant degrading enzymes. The proteins are important to elucidate the olfactory identify molecular mechanism. In this paper, the antenna of H viriplaca was observed by scanning electron microscopy, and use RT-PCR and RACE method to clone and analysis the olfactory related genes form antenna of H viriplaca and L glycinivorella. The Primary results are as follows:
The antenna of H viriplaca was observed by scanning electron microscopy. Female and male moths are filiform and consist of the scape, pedicel and flagellum, and eight types of sensillawere found. These were sensilla trichodea, sensilla chaetica, sensilla basiconca, sensilla coelocomica, Sensilla auricillica, sensillum squamiformia, sensilla styloconica and Bohm's bristles. The Sensillae distributed the rear and side of antenna, the scape was wrapped by squama, half of the flagellum was wrapped in order by squama, and a few of sensillum squamiformia and sensilla chaetica distributed in the squamas. Sensillae focus on flagellum, only a few Sensillae in scape and pedicel. Sensilla trichodea was the most sensillae, and sensilla chaetica was orderly on the antenna. The every little flagellum rear coverd by squamas, so most of the sensillae was in the rear and side of antenna.
Five OBPs and one OR genes were cloned from the antenna of H viriplaca and L glycinivorella by RT-PCR and RACE method. They are PBP and GOBP1of H viriplaca and L glycinivorella, GOBP2and Or83b of H viriplaca, and the PBP, GOBP1and GOBP2with vector pet21b were construeted and successed expressed in E. coll induced by IPTG.
The H viriplaca PBP cDNA was625bp in length and contained an open reading frame of510bp. The ORF encoded a polypeptide of170amino acid residues with a predicted molecular weight of19.0kD and a p1of5.65. A signal peptide of27residues was predicted at the N-terminus of the protein, and only one putative N-glycosylation sites was identified at the position34. The L. glycinivorella PBP cDNA was610bp in length and contained an open reading frame of504bp. The ORF encoded a polypeptide of168amino acid residues with a predicted molecular weight of19.3kD and a p1of5.55. A signal peptide of24residues was predicted at the N-terminus of the protein, and only one putative N-glycosylation sites was identified at the position52. Sequence blasted and analysed with other known insects indicated that HvirPBP and LglyPBP were characterized by six conservative Cys, which shared typical feature of OBP family. Identity of HvirPBP and LglyPBP were44%-96%and45%-70%with other known Lepidoptera insects. The kown insects had three subfamilies, and it may be connect with diversity pheromone.
The H viriplaca GOBP1cDNA was973bp in length and contained an open reading frame of495bp. The ORF encoded a polypeptide of164amino acid residues with a predicted molecular weight of19.0kD and a p1of5.32. A signal peptide of19residues was predicted at the N-terminus of the protein. The L. glycinivorella GOBP1cDNA was920bp in length and contained an open reading frame of492bp. The ORF encoded a polypeptide of163amino acid residues with a predicted molecular weight of18.8kD and a p1of5.29. A signal peptide of19residues was predicted at the N-terminus of the protein. Sequence blasted and analysed with other known insects indicated that HvirGOBP1and LglyGOBP1were characterized by six conservative Cys, which shared typical feature of OBP family. Identity of HvirGOBP1and Lgly GOBP1were65%-95%and65%-82%with other known Lepidoptera insects.
The H viriplaca GOBP2cDNA was624bp in length and contained an open reading frame of489bp. The ORF encoded a polypeptide of162amino acid residues with a predicted molecular weight of18.2kD and a pi of5.30. A signal peptide of21residues was predicted at the N-terminus of the protein. Sequence blasted and analysed with other known insects indicated that HvirGOBP2was characterized by six conservative Cys, which shared typical feature of OBP family. Identity of HvirGOBP2was well conserved, and between75%and99%with other known Lepidoptera insects.
The H. viriplaca Or83b cDNA was1715bp in length and contained an open reading frame of1422bp. The ORF encoded a polypeptide of473amino acid residues with a predicted molecular weight of53.5kD and a p1of8.37. Two putative N-glycosylation sites were identified at the position110and170, and no signal peptide at the N-terminus of the protein. Sequence blasted found the sequence of H. viriplaca Or83b was well conserved, and over96%identity with Or83bs of other Lepidoptera moths. Analysed the sequence indicated that HvirOr83b has seven transmembrane domains, which shared typical feature of odorant receptor family.
Analysed the five odorant binding proteins found they had the odorant binding protein typical feature, included small molecular weight, water-solubility acidic protein; six conservative Cys in sequences, A signal peptide was predicted at the N-terminus of the protein which secretory protein character. HvirOr83b has seven transmembrane domains, which shared typical feature of odorant receptor, and no signal peptide at the N-terminus of the protein.
PBP, GOBP1, GOBP2and Or83b four genes expressed in different organs in H. viriplaca, the four genes not only expressed in the antennae of both male and female adults but also expressed in abdomen and legs. Analysed PBP in different organs by RT-PCR method found that PBP gene is main expressed in male antenna, it was30times higher than PBP gene in male antenna, and PBP gene in male antenna was3times higher than PBP other organs.
引文
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