aiiA基因克隆及在尾巨桉中的诱导表达研究
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摘要
桉树原产澳大利亚,是桃金娘科(Myrtaceae)杯果木属(Angophora)、伞房属(Corymbia)和桉属(Eucalyptus)植物的统称。桉树生长快、对环境的适应能力强,是主要的实木与纸浆用材,由于轮伐期短,经济效益高,现已在热带、亚热带地区广泛种植。到2011年,中国桉树种植面积已有368万hm2。尾巨桉是尾叶桉(Eucalyptus urophylla)与巨桉(E. grandis)杂交种,树高50m以上,用途广泛,主要用于造纸与实木用材,是栽培面积最大的桉树人工林无性系。近年来随着桉树人工林的不断扩大,病虫害严重,已成为制约桉树人工林发展的主要因素。传统化学防治方法对环境的副作用大,破坏生态安全。利用包括转基因技术在内的现代生物技术手段进行桉树育种具有高效性和针对性,加快优质高抗新种质的选育。
N-酰基高丝氨酸内酯(N-acyl-homoserine lactones, AHL)是革兰氏阴性病原细菌产生的小分子化合物,也是该类病原菌群体感应系统的关键信号分子。当病原菌产生的AHL达到一定量浓度时,可激活病菌的致病基因表达,使寄主植物感病并表现相应病症。因此,破坏病原菌AHL信号分子可达到降低病原菌致病力的作用。
以尾巨桉优良无性系无菌苗茎段为外植体,通过对噻唑基脲类新型分裂素(N-phenyl-N'-[6-(2-chlorobenzothiazol)-yl] urea, PBU)等多种不同浓度生长调节剂组合的优化,进行胚性愈伤组织诱导及再生研究。结果表明,在添加了2mg-L-1PBU和0.05mg·L-1IAA的改良MS培养基上,茎段外植体5d后愈伤组织诱导率达96%以上。将愈伤组织接种在添加1mg·L-16-BA和0.05mg·L-1NAA的MS培养基上,诱芽率达90.8%。随后在添加了0.8mg·L-1PBU与0.05mg·L-1IAA的1/2MS培养基上诱导芽伸长,用1/2MS培养基附加0.5mg·L-1IBA诱导生根,移栽后得到完整再生植株。通过对尾巨桉胚性与非胚性愈伤组织差异的比较研究,初步揭示了尾巨桉愈伤组织的非胚性化表型的相关基因表达变化以及愈伤组织非胚性化是受环境胁迫影响的本质。
芽孢杆菌中的aiiA基因编码AHL-Lactonase能够水解信号分子AHLs。利用PCR方法从枯草芽孢杆菌中克隆了aiiA基因,序列分析表明该基因由753个碱基组成,编码含有250个氨基酸残基的蛋白质,核苷酸序列与已报道aiiA勺同源性为87%～96%。将该基因置于诱导型启动子PPP3调控下,连接到植物表达载体pCAMB1A1301中,成功构建了aiiA基因的植物诱导表达载体pCAMBIA-PPP3-aii4,为进一步通过转基因技术研究该基因的功能奠定了基础。
本研究选用了pGEX-4T-1表达载体,构建重组原核表达载体pGEX+aiiA。表达条件优化结果表明,在25℃下用0.2mmol·L-1IPTG诱导9h,蛋白的表达量最大,获得AIIA融合蛋白。研究表明AIIA蛋白能够降解细菌的AHL信号分子,猝灭细菌的群体感应,明显减弱病菌的致病力。
根据已报导的PPP3启动子碱基序列设计引物,从烟草基因组中扩增得到启动子PPP3。用PPP3替换pCAMBIA1301中与gus基因相连的35S启动子,构建重组质粒后导入农杆菌GV3101。通过农杆菌介导的瞬时表达技术将受PPP3控制的gus基因导入烟草。分别接种青枯菌和水杨酸,24h后烟草叶片中检测到gus基因转录水平分别提高了27.94和17.69倍。表明PPP3具有诱导表达活性。
经农杆菌介导法将表达载体pCAMBIA+PPP3+aiiA、pCAMBIA+35S+aiiA转化尾巨桉,得到了转基因植株。经PCR和Southern blotting证明aiiA基因成功插入到尾巨桉基因组中。Northern blotting和RT-PCR结果显示,在PPP3的调控下,aiiA受到诱导表达。实时荧光定量PCR结果表明,转基因尾巨桉接种青枯菌、茎腐病和焦枯病后,aiiA基因转录效率分别提高到原来的43.88、30.65和18.95倍。
灌根接种青枯菌和焦枯菌,转aiiA基因尾巨桉抗病性明显的增强,分别提高2.5和2.1个抗性级别,评价为中等抗病水平,表现为发病时间延迟,病情指数降低,抗性相关酶活性显著增强。灌根接种疫霉菌孢子,转基因植株表现出发病延迟、病情进展缓慢等抗病特点。转PPP3+aiiA基因植株抗性提高级别数要高于35S+aiiA基因植株,诱导抗性高于组成型表达。说明通过在植株中诱导表达群体猝灭基因,是提高植物抗病能力的有效策略。
Eucalyptus, belonging to the Myrtaceae family, is the most widely planted hardwood crop in tropical and subtropical regions, because its growth is superior to that of other hardwood crops, it adapts well to the environment, and its wood can be used for multiple purposes. Eucalyptus is the most important forest tree in China, covering more than3million hectares of commercial plantations. Eucalyptus urophylla×E. grandis can reach a height of up to50m, and its wood is widely used as timber for heavy structures and bridges. It is highly prized in China for its superior wood properties, rooting ability and disease resistance. Plantation forestry of E. grandis×E. urophylla supplies high-quality raw material for pulp, paper,"wood, and energy while it reduces the pressure on native forests and associated biodiversity. However, diseases such as bacterial wilt, fungal infection, and gray mold disease have seriously endangered the Eucalyptus crop in China, especially that of the hybrid E. grandis×E. urophylla. Traditional methods for the prevention and treatment of these diseases involve the use of chemical pesticides, which often cause environmental pollution and ecological damage. In contrast, the process of producing adevelopment of disease-resistant trees through genetic engineering can potentially be faster, more controllable and predictable and cheaper than traditional breeding.
N-acyl-homoserine lactones(AHL) are metabolites of mostly gram-negative bacteria critical and signaling molecules in bacterial quorum-sensing systems. They can activate the expression of pathogenic genes and induce diseases when AHL reach a threshold concentration. Therefore, reducing the concentration of AHLs is a key point of the diseases' control in plants.
The callus induction and shoot regeneration were studied systematically in elite clone of Eucalyptus urophylla×E. grandis by comparing the variety of combinations of different plant growth regulators, such as N-phenyl-N'-[6-(2-chlorobenzothiazol)-yl] urea (PBU). The results showed that sterile clones of seedling stem segments were used as explants and cultured in a modified MS medium supplemented with the synthetic growth regulator2mg·L-1PBU and0.05mg·L-1IAA. After cultivation for5d,96%of explants formed callus. Callus were transferred to MS medium containing1mg·L-16-BA and0.05mg·L-1NAA to induce bud formation, and90.8%of the callus induced by PBU exhibited adventitious bud formation. Shoot elongation was then stimulated on1/2MS supplemented with0.8mg-L-1PBU and0.05mg·L-1IAA for20d. For rooting, the elongated shoots were cultivated on root induction medium containing0.5mg·L-1IBA. Plantlets were then successfully transplanted to a greenhouse for growing into mature plants. Preliminary reveal callus embryonic phenomenon is affected by environmental stress also reveal the molecular mechanism of non-embryonic phenomenon.
The AHL-Lactonase, expressed by aiiA gene which widespread in Bacillus sp,can hydrolyze AHLs. The study have cloned the aiiA gene from Bacillus subtilis by PCR, The sequence analysis indicated that the clone was consisted of751nucleotides (nt), coding250amino acids.The nucleotide sequence showed87%-96%identities with those of the aiiA gene that have reported. A plant expression vector of the aiiA gene was constructed and named PCAMBIAb-PPP3-aiiA which the aiiA was controlled by the PPP3promoter. This work laid the foundations for future transgenic research on aiiA gene function.
pGEX-4T-1was constructed and transformed into bacteria strain of E.coli BL21for construction of genetically engineeringed bacterium strain. After9h induction with0.2mmol·L-1IPTG at25℃, the expression of AIIA protein reached to maximal level. AIIA fusion protein was obtained in E.coli system. It was observed that recombinant Escherichia coli producing AIIA proteins had AHL-degrading activity and could attenuate the plant pathogenicity of pathogen.
According to the sequences of pathogen-inducible plant promoters PPP3at GenBank, PPP3promoters were cloned from tobacco genome. It was used to replace cauliflower mosaic virus35S promoter of pCAMBIA1301. The recombinant plasmid was used to transform Agrobacterium tumefaciens GV3101. The inducibility of the PPP3promoters in tobacco leaf was evaluated by Agrobacterium tumefaciens-transient genetic transformation assye. Real-time quantitative PCR was used to screen the PPP3promoters with high inducible expression. The results showed that the gus transcript level under the control of PPP3promoters increased respectively27.94fold and17.69fold after inoculation with Ralstonia solanacearum and SA. The PPP3promoter had the advantages such as low basal activity and high expression activity.
The recombined plasmid was transformed into Eucalyptus urophylla x E.grandis by Agrobacterium-mediated transformation. The results of PCR and Southern blotting showed that the aiiA genes were successfully integrated into the Eucalyptus urophylla×E.grandis genome. The result of reverse transcription polymerase chain reaction (RT-PCR) showed aiiA had partly basal expression. The transcript efficiency of aiiA was calculated by real-time quantitative PCR. The transcript efficiency of aiiA under the control of PPP3promoters increased respectively43.88,30.65and18.95fold after inoculation with Ralstonia solanacearum, E.carotovoraf.sp.zeaeSabet and C.quinqueseptatun.
The result of plant inoculation showed that ransgenic Eucalyptus urophylla×E. grandis expressing AIIA exhibit significantly enhanced resistance to disease comparing to non-transgenic ones. The transgenic plants could delay the wilt symptom development and make disease index reduced. The activity of PPO, PAL, PPO and SOD increased significantly than non-transgenic Eucalyptus urophylla×E.grandis. Inoculated with spores of Phytophthora capsici, transgenic E.urophylla×E.grandis showed markedly enhancement of disease resistance. All of the results indicated that the resistance of plants could be enhanced by introducing aiiA gene into the genomic of plants.
N-酰基高丝氨酸内酯(N-acyl-homoserine lactones, AHL)是革兰氏阴性病原细菌产生的小分子化合物,也是该类病原菌群体感应系统的关键信号分子。当病原菌产生的AHL达到一定量浓度时,可激活病菌的致病基因表达,使寄主植物感病并表现相应病症。因此,破坏病原菌AHL信号分子可达到降低病原菌致病力的作用。
以尾巨桉优良无性系无菌苗茎段为外植体,通过对噻唑基脲类新型分裂素(N-phenyl-N'-[6-(2-chlorobenzothiazol)-yl] urea, PBU)等多种不同浓度生长调节剂组合的优化,进行胚性愈伤组织诱导及再生研究。结果表明,在添加了2mg-L-1PBU和0.05mg·L-1IAA的改良MS培养基上,茎段外植体5d后愈伤组织诱导率达96%以上。将愈伤组织接种在添加1mg·L-16-BA和0.05mg·L-1NAA的MS培养基上,诱芽率达90.8%。随后在添加了0.8mg·L-1PBU与0.05mg·L-1IAA的1/2MS培养基上诱导芽伸长,用1/2MS培养基附加0.5mg·L-1IBA诱导生根,移栽后得到完整再生植株。通过对尾巨桉胚性与非胚性愈伤组织差异的比较研究,初步揭示了尾巨桉愈伤组织的非胚性化表型的相关基因表达变化以及愈伤组织非胚性化是受环境胁迫影响的本质。
芽孢杆菌中的aiiA基因编码AHL-Lactonase能够水解信号分子AHLs。利用PCR方法从枯草芽孢杆菌中克隆了aiiA基因,序列分析表明该基因由753个碱基组成,编码含有250个氨基酸残基的蛋白质,核苷酸序列与已报道aiiA勺同源性为87%～96%。将该基因置于诱导型启动子PPP3调控下,连接到植物表达载体pCAMB1A1301中,成功构建了aiiA基因的植物诱导表达载体pCAMBIA-PPP3-aii4,为进一步通过转基因技术研究该基因的功能奠定了基础。
本研究选用了pGEX-4T-1表达载体,构建重组原核表达载体pGEX+aiiA。表达条件优化结果表明,在25℃下用0.2mmol·L-1IPTG诱导9h,蛋白的表达量最大,获得AIIA融合蛋白。研究表明AIIA蛋白能够降解细菌的AHL信号分子,猝灭细菌的群体感应,明显减弱病菌的致病力。
根据已报导的PPP3启动子碱基序列设计引物,从烟草基因组中扩增得到启动子PPP3。用PPP3替换pCAMBIA1301中与gus基因相连的35S启动子,构建重组质粒后导入农杆菌GV3101。通过农杆菌介导的瞬时表达技术将受PPP3控制的gus基因导入烟草。分别接种青枯菌和水杨酸,24h后烟草叶片中检测到gus基因转录水平分别提高了27.94和17.69倍。表明PPP3具有诱导表达活性。
经农杆菌介导法将表达载体pCAMBIA+PPP3+aiiA、pCAMBIA+35S+aiiA转化尾巨桉,得到了转基因植株。经PCR和Southern blotting证明aiiA基因成功插入到尾巨桉基因组中。Northern blotting和RT-PCR结果显示,在PPP3的调控下,aiiA受到诱导表达。实时荧光定量PCR结果表明,转基因尾巨桉接种青枯菌、茎腐病和焦枯病后,aiiA基因转录效率分别提高到原来的43.88、30.65和18.95倍。
灌根接种青枯菌和焦枯菌,转aiiA基因尾巨桉抗病性明显的增强,分别提高2.5和2.1个抗性级别,评价为中等抗病水平,表现为发病时间延迟,病情指数降低,抗性相关酶活性显著增强。灌根接种疫霉菌孢子,转基因植株表现出发病延迟、病情进展缓慢等抗病特点。转PPP3+aiiA基因植株抗性提高级别数要高于35S+aiiA基因植株,诱导抗性高于组成型表达。说明通过在植株中诱导表达群体猝灭基因,是提高植物抗病能力的有效策略。
Eucalyptus, belonging to the Myrtaceae family, is the most widely planted hardwood crop in tropical and subtropical regions, because its growth is superior to that of other hardwood crops, it adapts well to the environment, and its wood can be used for multiple purposes. Eucalyptus is the most important forest tree in China, covering more than3million hectares of commercial plantations. Eucalyptus urophylla×E. grandis can reach a height of up to50m, and its wood is widely used as timber for heavy structures and bridges. It is highly prized in China for its superior wood properties, rooting ability and disease resistance. Plantation forestry of E. grandis×E. urophylla supplies high-quality raw material for pulp, paper,"wood, and energy while it reduces the pressure on native forests and associated biodiversity. However, diseases such as bacterial wilt, fungal infection, and gray mold disease have seriously endangered the Eucalyptus crop in China, especially that of the hybrid E. grandis×E. urophylla. Traditional methods for the prevention and treatment of these diseases involve the use of chemical pesticides, which often cause environmental pollution and ecological damage. In contrast, the process of producing adevelopment of disease-resistant trees through genetic engineering can potentially be faster, more controllable and predictable and cheaper than traditional breeding.
N-acyl-homoserine lactones(AHL) are metabolites of mostly gram-negative bacteria critical and signaling molecules in bacterial quorum-sensing systems. They can activate the expression of pathogenic genes and induce diseases when AHL reach a threshold concentration. Therefore, reducing the concentration of AHLs is a key point of the diseases' control in plants.
The callus induction and shoot regeneration were studied systematically in elite clone of Eucalyptus urophylla×E. grandis by comparing the variety of combinations of different plant growth regulators, such as N-phenyl-N'-[6-(2-chlorobenzothiazol)-yl] urea (PBU). The results showed that sterile clones of seedling stem segments were used as explants and cultured in a modified MS medium supplemented with the synthetic growth regulator2mg·L-1PBU and0.05mg·L-1IAA. After cultivation for5d,96%of explants formed callus. Callus were transferred to MS medium containing1mg·L-16-BA and0.05mg·L-1NAA to induce bud formation, and90.8%of the callus induced by PBU exhibited adventitious bud formation. Shoot elongation was then stimulated on1/2MS supplemented with0.8mg-L-1PBU and0.05mg·L-1IAA for20d. For rooting, the elongated shoots were cultivated on root induction medium containing0.5mg·L-1IBA. Plantlets were then successfully transplanted to a greenhouse for growing into mature plants. Preliminary reveal callus embryonic phenomenon is affected by environmental stress also reveal the molecular mechanism of non-embryonic phenomenon.
The AHL-Lactonase, expressed by aiiA gene which widespread in Bacillus sp,can hydrolyze AHLs. The study have cloned the aiiA gene from Bacillus subtilis by PCR, The sequence analysis indicated that the clone was consisted of751nucleotides (nt), coding250amino acids.The nucleotide sequence showed87%-96%identities with those of the aiiA gene that have reported. A plant expression vector of the aiiA gene was constructed and named PCAMBIAb-PPP3-aiiA which the aiiA was controlled by the PPP3promoter. This work laid the foundations for future transgenic research on aiiA gene function.
pGEX-4T-1was constructed and transformed into bacteria strain of E.coli BL21for construction of genetically engineeringed bacterium strain. After9h induction with0.2mmol·L-1IPTG at25℃, the expression of AIIA protein reached to maximal level. AIIA fusion protein was obtained in E.coli system. It was observed that recombinant Escherichia coli producing AIIA proteins had AHL-degrading activity and could attenuate the plant pathogenicity of pathogen.
According to the sequences of pathogen-inducible plant promoters PPP3at GenBank, PPP3promoters were cloned from tobacco genome. It was used to replace cauliflower mosaic virus35S promoter of pCAMBIA1301. The recombinant plasmid was used to transform Agrobacterium tumefaciens GV3101. The inducibility of the PPP3promoters in tobacco leaf was evaluated by Agrobacterium tumefaciens-transient genetic transformation assye. Real-time quantitative PCR was used to screen the PPP3promoters with high inducible expression. The results showed that the gus transcript level under the control of PPP3promoters increased respectively27.94fold and17.69fold after inoculation with Ralstonia solanacearum and SA. The PPP3promoter had the advantages such as low basal activity and high expression activity.
The recombined plasmid was transformed into Eucalyptus urophylla x E.grandis by Agrobacterium-mediated transformation. The results of PCR and Southern blotting showed that the aiiA genes were successfully integrated into the Eucalyptus urophylla×E.grandis genome. The result of reverse transcription polymerase chain reaction (RT-PCR) showed aiiA had partly basal expression. The transcript efficiency of aiiA was calculated by real-time quantitative PCR. The transcript efficiency of aiiA under the control of PPP3promoters increased respectively43.88,30.65and18.95fold after inoculation with Ralstonia solanacearum, E.carotovoraf.sp.zeaeSabet and C.quinqueseptatun.
The result of plant inoculation showed that ransgenic Eucalyptus urophylla×E. grandis expressing AIIA exhibit significantly enhanced resistance to disease comparing to non-transgenic ones. The transgenic plants could delay the wilt symptom development and make disease index reduced. The activity of PPO, PAL, PPO and SOD increased significantly than non-transgenic Eucalyptus urophylla×E.grandis. Inoculated with spores of Phytophthora capsici, transgenic E.urophylla×E.grandis showed markedly enhancement of disease resistance. All of the results indicated that the resistance of plants could be enhanced by introducing aiiA gene into the genomic of plants.
引文
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