鸡枞菌与淡色库蚊及家蝇中漆酶编码基因的克隆与表达
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摘要
漆酶(EC 1.10.3.2)是一种含铜的多酚氧化酶,普遍存在于真菌(特别是白腐真菌)、植物、昆虫和细菌中。漆酶的作用底物广泛,有250种左右,主要以酚类和芳香类化合物为主。漆酶在木质素降解、制浆造纸生物漂白、环境保护、食品及饮料加工等领域具有广阔的应用前景。本文以黑翅土白蚁共生鸡枞(实为“土”字旁+“从”)菌、淡色库蚊和家蝇为材料,研究了鸡枞菌中漆酶的性质,产酶诱导和漆酶基因的克隆与异源表达及淡色库蚊和家蝇中漆酶基因的克隆与表达。主要结果如下:
1.分析了黑翅土白蚁工蚁肠道、若蚁、鸡枞菌和菌圃中漆酶的分布,研究了温度、pH值和抑制剂对这些漆酶的影响。实验发现,鸡枞菌具有最高的漆酶活性,达到13.36 U/g;其次为菌圃,为4.24 U/g;在工蚁肠道中,漆酶的活性相对较低,只有1.05 U/g;而若蚁中几乎检测不到漆酶的活性。这些漆酶在酸性条件(pH 3.0-3.5)下表现出最高的活性和较强的热稳定性。一些化合物,如L-半胱氨酸、DTT和氟离子,对漆酶具有强烈的抑制作用。这些结果表明,鸡枞菌可能通过分泌漆酶到菌圃来降解菌圃中的木质素,从而改善菌圃作为工蚁食物的品质。
2.利用不同培养基在不同CO_2浓度环境下对鸡枞菌进行了培养。结果发现,在PDA、CDA、PD和KB(高氮/低氮)培养基中,鸡枞菌都能生长。显微观察发现,这些真菌的形态发生了改变;在PDA和KB(低氮)培养基中培养的鸡枞菌显示出了较低的漆酶活性,两者的活性分别为1.98U/g和0.73U/g。在CO_2浓度为4%、10%和15%的环境中,鸡枞菌均能正常生长,并具有低的漆酶活性,但在CO_2浓度为25%、50%和75%的环境中不能正常生长。这些结果表明,通过人工方式培养可培养出具漆酶活性的共生真菌。
3.通过RACE方法从鸡枞菌中克隆出编码漆酶的基因并在大肠杆菌中对其进行了表达。鸡枞菌中编码漆酶的基因(talcc)全长为2295 bp,编码1551 bp的开放阅读框(ORF)。全长cDNA共由15个外显子组成,预测编码517个氨基酸,推导的蛋白质分子量为55.8KDa,等电点(pI)为5.92。比对发现,鸡枞菌漆酶TaLCC与Cyathus bulleri真菌中漆酶CbLCC的序列最为相似,达到67%;而与来自同属真菌Termitomyces的漆酶LCC1-2序列的相似度只有47%。同时,把含原始信号肽的talcc cDNA克隆到pET-28a表达载体中,然后导入BL21大肠杆菌中,IPTG诱导表达后得到一条60 kDa左右的粗条带,进一步通过western印迹分析鉴定,此条带为重组漆酶蛋白。但经酶活测定发现,纯化的重组蛋白未表现出漆酶活性。
4.通过RACE方法从淡色库蚊和家蝇中克隆漆酶基因。实验发现,淡色库蚊漆酶基因(CpLac2)全长由2289个碱基组成,预测编码762个氨基酸,预测的蛋白含有31个氨基酸的信号肽,说明为分泌型蛋白,其分子量为81.2 kDa,等电点(pI)为6.21。与其他昆虫漆酶比对后发现,CpLac2与昆虫漆酶-2基因存在75%-88%的相似性,属于漆酶-2基因。家蝇漆酶基因片断为1482 bp,此片断也与漆酶-2基因存在极高的相似性,也属于漆酶-2基因。
5.运用RT-PCR和荧光实时PCR分析了CpLac2的表达发育模型及淡色库蚊敏感与抗氰戊菊酯品系的CpLac2表达差异。结果表明,CpLac2基因在淡色库蚊的卵、四龄幼虫和蛹中大量表达,说明CpLac2基因可能参与了淡色库蚊卵鞘、蛹和成虫体表的鞣化过程;CpLac2基因在抗氰戊菊酯淡色库蚊品系中的表达显著高于在敏感品系中的,预示着它可能参与了淡色库蚊对氰戊菊酯抗性的形成,因为CpLac2基因的高水平表达,可能使得淡色库蚊的体表变得更紧密或更坚硬,从而阻止或减缓药剂从体表向体内的渗透。
Laccase(EC 1.10.3.2) is one of copper-contained polyphenol oxidases,and it exits in fungi(especially white-rot fungi),plants,insects,and bacteria.Laccase has broad substrates(about 250 kinds),which mainly are phenolic and aromatic compounds.Laccase has provided great application potential in ligin degradation, biobleaching in pulps,environment protection,processing of foodstuffs and beverages. In this paper,we analyzed the characteristics of laccase from the symbiotic fungi (Termitomyces albuminosus) of Odontotermes formosanus,induced laccase production,cloned a laccase gene from the fungi and expressed it in Escherichia coli. Meanwhile,we also cloned laccase genes from Culex pipiens pallens and Musca domeslica,and analyzed the relation between gene expression of laccase and resistance of fenvalerate in Culex pipiens pallens population.The main results are as follows:
1.Analysis of the distribution of laccase in alimentary tract of workers of O. formosanus,nymph,T.albuminosus,and fungus comb,as well as effect of temperature,pH value and inhibitors to the activity of laccase.The results showed that T.albuminosus exhibited the highest laccase activity,reached 13.36 U/g;followed by the fungus comb,its laccase activity was 4.24 U/g;in alimentary tract of workers,its laccase activity only was 1.05 U/g;in nymph,the laccase activity could not be detected.These laccases from different materials exhibited high activity in acid condition(pH 3.0-3.5) and thermal stability.Also,these laccases could be inhibited strongly by L-cysteine,DTT and fluoride ions.The results suggest that T. albuminosus takes part in lignin degradation of plant litter through secreting laccase into fungus comb in colonies of O.formosanus.
2.Induction of laccase production in environment with different cultivation media and CO_2 concentrations.The results showed that T.albuminosus could be cultured in PDA,CDA,PD,and KB(with LN/HN) media,but the mycelium of these cultured fungi was different from the original fungi.The fungi cultured in PDA and KB(LN) expressed low laccase activity.The T.albuminosus could grow normally in the environments with 4%or 10%and 15%of CO_2 concentration,and expressed low laccase activity,but it could not grow normally in the environments with 25%or 50% and 75%of CO_2 concentration.The results suggest that the laccase activity of T. albuminosus can be induced with cultivation media and CO_2 in laboratory.
3.Cloning of a laccase gene with RACE method from T.albuminosus and expression of this laccase gene in E.coli.The full-length of laccase gene(talcc) from the symbiotic fungi is 2295 bp,contained an open reading frame(ORF) of 1551 bp. This cDNAs was composed of 15 exons,and encoded a putative 517 amino acid protein.The predicted molecular weight(MW) and isoelectric point(pI) for the mature protein of talcc are 55.8 kDa and pH 5.92,respectively.After alignment with other fungus-originated laccases,the TaLCC presented the highest similarity(about 67%) to CbLCC from Cyathus bulleri,but had only 47%similarity to laccase from the same genus fungi.Meanwhile,the cDNAs of talcc with native signal peptide was transformed into the expression vector of pET-28a,and the recombinant vector of pET-28a/talcc was transformed into E.coli of BL21.After induced 1-3 hours with IPTG,the recombinant strain secreted a 60 kDa protein.This protein was the recombinant laccase through analysis of Western blot.Unfortunately,no laccase activity was detected in this purified protein.
4.Cloning of laccase genes from C.pipiens pollens and M.domestica with RACE method.The laccase gene CpLac2 of C.pipiens pollens contained an ORF of 2289 bp that encoded a putative 762 amino acid protein.The predicted MW and pI for the mature protein of CpLac2 were 81.2 kDa and pH 6.21,respectively.The putative protein had a signal peptide of 31 amino acids.After blasted with other insect laccases, CpLac2 had 78-90%identity with the insect laccase 2 family.This means the CpLac2 belongs to the laccase 2 family.A laccase fragment of 1482 bp was cloned from M. domestica.This sequence also presented >90%identity with insect laccase 2 family, and it also means this sequence belongs to the laccase 2 family.
5.The developmental expression model of CpLac2 in C.pipiens pollens was measured by RT-PCR.The result showed the CpLac2 was abundantly expressed in egg,the 4~(th) instar larva and pupa,which suggested the role of CpLac2 for egg chorion tanning and cuticular sclerotization.Meanwhile,the expression of CpLac2 in fenvalerat-susceptible and -resistant strains of C.pipiens pollens was measured by real-time PCR.The result revealed the CpLac2 was significant higher expressed in resistant strain than in susceptible strain.The overexpression of CpLac2 in resistant strain suggested that resistance could derive from reinforcement of the cuticle,which decreased the penetration of insecticide in cuticle.
1.分析了黑翅土白蚁工蚁肠道、若蚁、鸡枞菌和菌圃中漆酶的分布,研究了温度、pH值和抑制剂对这些漆酶的影响。实验发现,鸡枞菌具有最高的漆酶活性,达到13.36 U/g;其次为菌圃,为4.24 U/g;在工蚁肠道中,漆酶的活性相对较低,只有1.05 U/g;而若蚁中几乎检测不到漆酶的活性。这些漆酶在酸性条件(pH 3.0-3.5)下表现出最高的活性和较强的热稳定性。一些化合物,如L-半胱氨酸、DTT和氟离子,对漆酶具有强烈的抑制作用。这些结果表明,鸡枞菌可能通过分泌漆酶到菌圃来降解菌圃中的木质素,从而改善菌圃作为工蚁食物的品质。
2.利用不同培养基在不同CO_2浓度环境下对鸡枞菌进行了培养。结果发现,在PDA、CDA、PD和KB(高氮/低氮)培养基中,鸡枞菌都能生长。显微观察发现,这些真菌的形态发生了改变;在PDA和KB(低氮)培养基中培养的鸡枞菌显示出了较低的漆酶活性,两者的活性分别为1.98U/g和0.73U/g。在CO_2浓度为4%、10%和15%的环境中,鸡枞菌均能正常生长,并具有低的漆酶活性,但在CO_2浓度为25%、50%和75%的环境中不能正常生长。这些结果表明,通过人工方式培养可培养出具漆酶活性的共生真菌。
3.通过RACE方法从鸡枞菌中克隆出编码漆酶的基因并在大肠杆菌中对其进行了表达。鸡枞菌中编码漆酶的基因(talcc)全长为2295 bp,编码1551 bp的开放阅读框(ORF)。全长cDNA共由15个外显子组成,预测编码517个氨基酸,推导的蛋白质分子量为55.8KDa,等电点(pI)为5.92。比对发现,鸡枞菌漆酶TaLCC与Cyathus bulleri真菌中漆酶CbLCC的序列最为相似,达到67%;而与来自同属真菌Termitomyces的漆酶LCC1-2序列的相似度只有47%。同时,把含原始信号肽的talcc cDNA克隆到pET-28a表达载体中,然后导入BL21大肠杆菌中,IPTG诱导表达后得到一条60 kDa左右的粗条带,进一步通过western印迹分析鉴定,此条带为重组漆酶蛋白。但经酶活测定发现,纯化的重组蛋白未表现出漆酶活性。
4.通过RACE方法从淡色库蚊和家蝇中克隆漆酶基因。实验发现,淡色库蚊漆酶基因(CpLac2)全长由2289个碱基组成,预测编码762个氨基酸,预测的蛋白含有31个氨基酸的信号肽,说明为分泌型蛋白,其分子量为81.2 kDa,等电点(pI)为6.21。与其他昆虫漆酶比对后发现,CpLac2与昆虫漆酶-2基因存在75%-88%的相似性,属于漆酶-2基因。家蝇漆酶基因片断为1482 bp,此片断也与漆酶-2基因存在极高的相似性,也属于漆酶-2基因。
5.运用RT-PCR和荧光实时PCR分析了CpLac2的表达发育模型及淡色库蚊敏感与抗氰戊菊酯品系的CpLac2表达差异。结果表明,CpLac2基因在淡色库蚊的卵、四龄幼虫和蛹中大量表达,说明CpLac2基因可能参与了淡色库蚊卵鞘、蛹和成虫体表的鞣化过程;CpLac2基因在抗氰戊菊酯淡色库蚊品系中的表达显著高于在敏感品系中的,预示着它可能参与了淡色库蚊对氰戊菊酯抗性的形成,因为CpLac2基因的高水平表达,可能使得淡色库蚊的体表变得更紧密或更坚硬,从而阻止或减缓药剂从体表向体内的渗透。
Laccase(EC 1.10.3.2) is one of copper-contained polyphenol oxidases,and it exits in fungi(especially white-rot fungi),plants,insects,and bacteria.Laccase has broad substrates(about 250 kinds),which mainly are phenolic and aromatic compounds.Laccase has provided great application potential in ligin degradation, biobleaching in pulps,environment protection,processing of foodstuffs and beverages. In this paper,we analyzed the characteristics of laccase from the symbiotic fungi (Termitomyces albuminosus) of Odontotermes formosanus,induced laccase production,cloned a laccase gene from the fungi and expressed it in Escherichia coli. Meanwhile,we also cloned laccase genes from Culex pipiens pallens and Musca domeslica,and analyzed the relation between gene expression of laccase and resistance of fenvalerate in Culex pipiens pallens population.The main results are as follows:
1.Analysis of the distribution of laccase in alimentary tract of workers of O. formosanus,nymph,T.albuminosus,and fungus comb,as well as effect of temperature,pH value and inhibitors to the activity of laccase.The results showed that T.albuminosus exhibited the highest laccase activity,reached 13.36 U/g;followed by the fungus comb,its laccase activity was 4.24 U/g;in alimentary tract of workers,its laccase activity only was 1.05 U/g;in nymph,the laccase activity could not be detected.These laccases from different materials exhibited high activity in acid condition(pH 3.0-3.5) and thermal stability.Also,these laccases could be inhibited strongly by L-cysteine,DTT and fluoride ions.The results suggest that T. albuminosus takes part in lignin degradation of plant litter through secreting laccase into fungus comb in colonies of O.formosanus.
2.Induction of laccase production in environment with different cultivation media and CO_2 concentrations.The results showed that T.albuminosus could be cultured in PDA,CDA,PD,and KB(with LN/HN) media,but the mycelium of these cultured fungi was different from the original fungi.The fungi cultured in PDA and KB(LN) expressed low laccase activity.The T.albuminosus could grow normally in the environments with 4%or 10%and 15%of CO_2 concentration,and expressed low laccase activity,but it could not grow normally in the environments with 25%or 50% and 75%of CO_2 concentration.The results suggest that the laccase activity of T. albuminosus can be induced with cultivation media and CO_2 in laboratory.
3.Cloning of a laccase gene with RACE method from T.albuminosus and expression of this laccase gene in E.coli.The full-length of laccase gene(talcc) from the symbiotic fungi is 2295 bp,contained an open reading frame(ORF) of 1551 bp. This cDNAs was composed of 15 exons,and encoded a putative 517 amino acid protein.The predicted molecular weight(MW) and isoelectric point(pI) for the mature protein of talcc are 55.8 kDa and pH 5.92,respectively.After alignment with other fungus-originated laccases,the TaLCC presented the highest similarity(about 67%) to CbLCC from Cyathus bulleri,but had only 47%similarity to laccase from the same genus fungi.Meanwhile,the cDNAs of talcc with native signal peptide was transformed into the expression vector of pET-28a,and the recombinant vector of pET-28a/talcc was transformed into E.coli of BL21.After induced 1-3 hours with IPTG,the recombinant strain secreted a 60 kDa protein.This protein was the recombinant laccase through analysis of Western blot.Unfortunately,no laccase activity was detected in this purified protein.
4.Cloning of laccase genes from C.pipiens pollens and M.domestica with RACE method.The laccase gene CpLac2 of C.pipiens pollens contained an ORF of 2289 bp that encoded a putative 762 amino acid protein.The predicted MW and pI for the mature protein of CpLac2 were 81.2 kDa and pH 6.21,respectively.The putative protein had a signal peptide of 31 amino acids.After blasted with other insect laccases, CpLac2 had 78-90%identity with the insect laccase 2 family.This means the CpLac2 belongs to the laccase 2 family.A laccase fragment of 1482 bp was cloned from M. domestica.This sequence also presented >90%identity with insect laccase 2 family, and it also means this sequence belongs to the laccase 2 family.
5.The developmental expression model of CpLac2 in C.pipiens pollens was measured by RT-PCR.The result showed the CpLac2 was abundantly expressed in egg,the 4~(th) instar larva and pupa,which suggested the role of CpLac2 for egg chorion tanning and cuticular sclerotization.Meanwhile,the expression of CpLac2 in fenvalerat-susceptible and -resistant strains of C.pipiens pollens was measured by real-time PCR.The result revealed the CpLac2 was significant higher expressed in resistant strain than in susceptible strain.The overexpression of CpLac2 in resistant strain suggested that resistance could derive from reinforcement of the cuticle,which decreased the penetration of insecticide in cuticle.
引文
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