产酶溶杆菌生防相关基因的克隆、功能分析及工程菌构建
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摘要
产酶溶杆菌(Lysobacter enzymogenes) OH11是本实验室从辣椒根际土壤中分离得到的一种新型生防菌,属溶杆菌属(Lysobacter)、黄单胞科(Xanthomonadaceae)。该菌GC含量较高,有滑行现象,能产生多种胞外酶,包括几丁质酶、蛋白酶、纤维素酶、p-1,3-葡聚糖酶,对许多植物病原真菌、卵菌具有很好的拮抗活性。本研究旨在建立产酶溶杆菌OH11的遗传操作系统,构建转座子随机插入文库,并以此为基础,克隆色素和胞外多糖合成相关基因,并对克隆基因进行功能分析,以明确色素和胞外多糖在该菌定殖、拮抗和生防中的功能。同时,构建具有降解酰基高丝氨酸内酯类信号分子(AHL)能力的产酶溶杆菌工程菌株,用于防治植物细菌病害,拓宽其生防范围。
首次利用mariner转座子对产酶溶杆菌OH11进行了转座诱变,成功构建产酶溶杆菌OH11的转座子随机插入文库。在几丁质选择性培养基上,根据菌落周围是否产生水解圈,成功从OH11突变株文库中筛选出4个几丁质酶缺失突变株。利用建立的任意PCR (abritrary PCR)技术,快速鉴定了mariner转座子在4个几丁质酶缺失突变株中的插入位点。序列分析表明插入位点涉及调控基因clp,几丁质酶编码基因chiA,羧基末端蛋白酶(carboxyl-terminal protease)编码基因cptA和未知蛋白(hyperthetical protein)编码基因。随后,以chiA为报道基因,分析了3种不同类型的自杀质粒(pMW91CM, pJQ200SK, pEX18GM)在OH11中用于完成基因定向敲除的可行性,确定了自杀质粒pEX18GM可以有效地在OH11中完成基因的定向插入失活和缺失敲除,并构建了chiA的缺失突变株。有趣的是,chiA缺失突变株不改变对水稻纹枯病菌(Rhizoctonia solani,)、油菜菌核病菌(Sclerotinia sclerotiorum)、小麦赤霉病菌(Fusarium graminearum)和辣椒疫霉(Phytophthora capsici)的拮抗活性,揭示在防治这四种病原真菌或卵菌时,几丁质酶并不是主要的胞外酶。利用来自于大肠杆菌的组成型强启动子Plpp,实现了chiA基因的互补及外源基因gfp的表达,成功构建了基于质粒的功能基因互补系统和外源基因表达系统。本部分建立的产酶溶杆菌OH11的遗作操作系统,为后续研究OH11的生防分子机理及工程菌的构建奠定了基础。
结合表型筛选和验证,从OH11的突变株文库中筛选到2株色素突变株,分别命名为OH11H和OH11B。在LA平板上,野生型OH11产生黄色素,菌落为黄色。色素突变株OH11H可以产生可溶性的黑色素,但其菌落仍为黄色；OH11B菌落形态为白色,缺失了黄色素的合成。通过任意PCR和亚克隆技术,鉴定了转座子在OH11H和OH11B中的插入位点分别为hmgA基因和acp基因。hmgA基因编码尿黑酸1,2-加氧化酶,参与酪氨酸的代谢。而acp基因编码3-羰基(酰基载体蛋白)合成酶Ⅰ,与脂肪酸生物合成(FAS)相关。在前期以chiA基因作为报告基因筛选自杀载体的基础上,本研究再次以hmgA为报告基因,验证不同类型的自身载体在OH11中完成基因定向敲除的可行性,并再次确认在参试的自杀载体中,只有pEX18GM才能完成靶标基因的定向缺失敲除,与前期筛选结果一致。通过同源重组技术,分别构建了hmgA和acp的定向敲除突变株,并明确hmgA的突变改变了OH11的细胞形态,使细胞从棍棒状变为椭圆状,但不改变菌落形态；提高了OH11生物膜形成能力、在水稻叶片上的定殖能力及对水稻纹枯病的防效,但不改变OH11胞外多糖形成能力、4种胞外酶的产生能力和平板拮抗活性。明确了acp的突变不改变OH11的细胞形状,但使细胞周围“多糖”类物质增加,降低了OH11生物膜形成能力,在水稻叶片上的定殖能力及对水稻纹枯病的防效,同样不改变OH11胞外多糖形成能力、4种胞外酶的产生能力和平板拮抗活性。此外,还初步明确了可溶性黑色素和菌体自身的黄色素可能与细菌抵御紫外光损伤无关。
结合表型筛选和验证,从OH11的突变株文库中筛选到5株胞外多糖(exopolysacchide, EPS)缺失突变株,分别命名为MEPS1-MEPS5。在LAS培养基上,野生型OH11菌落黄色、饱满、光滑,而EPS缺失突变株茵落干燥、皱缩,但菌落仍为黄色。通过任意PCR技术(arbitrary PCR),快速鉴定了转座子在MES1-MES5中的插入位点分别为别是tonB基因、透性酶(permease)编码基因、Ⅳ分泌系统中Pilin蛋白编码基因,以及未知蛋白编码基因,表明众多基因参与了EPS的合成。在任意PCR的基础上,进一步通过亚克隆技术(sub-cloning),克隆了tonB的完整基因。通过同源重组技术,构建了tonB定向缺失突变株,并明确tonB的突变降低了OH11的滑行能力,改变了OH11的菌落形态,但不改变菌体细胞形态；降低了OH11生物膜形成能力、在水稻叶片上的定殖能力及对水稻纹枯病的防效。初步明确了EPS可能与产酶溶杆菌抵御紫外光损伤相关。
利用大肠杆菌中的组成型强启动子Plpp实现了外源基因即AHL类信号分子降解基因aiiA在产酶溶杆菌OH11中的重组表达,构建了抗生素标记的工程菌株OH11A。菌株OH11A能显著降解不同植物病原细菌产生的AHL类信号分子。在离体生防试验中,OH11A能大幅度降低软腐病菌对大白菜的致病性,但不抑制软腐病菌在寄主组织中的生长。在活体生防试验中,OH11A能分别显著降低软腐病菌和西瓜果斑病菌对仙人掌和甜瓜苗的毒性。在后续工作中,我们以lacZ为报告基因,构建了基于广宿主载体pBBR1-CMS5的新型启动子探针pBBRLACZR。利用该探针,从产酶溶杆菌OH11的启动子文库中筛选到一个具有启动子功能的DNA片段,通过启动子亚克隆并结合反复的筛选验证,最终从OH11中克隆了一个组成型强表达的启动子PB500。以前期工作中克隆的hmgA为同源重组靶标基因,以SacB为反向筛选标记,将外源融合基因PB500-aiiA定向整合到hmgA中,构建了无标记、稳定遗传并能显著降解信号分子AHLA的工程菌株OH11PA.工程菌株OH11PA能显著降低软腐病菌在大白菜上的毒性。
Lysobacter enzymogenes strain oH11, isolated from cayenne-root soil, is reported for the first time as a bacterial biological control agent in China. It is characterizaed with a high G+C content and gliding motility, and has been shown to produce extracellular lytic enzymes, including chitinase, protease, cellulase andβ-1,3-glucanas. Furhtermore, it displays in vitro activity against fungal and oomycetous plant pathogens. The objectives of this work are to:(i) establish the genetic manipulation system in strain OH 11 and construct a mutant library by a mariner transposon mutagenesis; (ii) to identify the pigment and exopolysacchide formation related genes, and to investigate the function of these cloned genes in colonization, antimicrobial activity and biocontrol efficiency of strain OH11; (ii) to construct an engineering strain with AHL degradation capacity to biocontrol bacterial plant diseases, broadening the biocontrol spectrum of L. enzymogenes.
Transposon mutagenesis was first performed in Lysobacter enzymogenes strain OH 11 by a mariner transposon and a large-scale mutant library was obtained. Four chitinase-defective mutants, which could not produce hydrolytic zones around colonies in chitine selection medium, were selected from over 2,000 chloramphenicol-resistant (Cm1) mutants. By using arbitrary PCR techonology, the transposon insertion in 4 chitinase-defective mutants were identified as regulatory gene clp, chitinase encoding gene chiA, carboxyl-terminal protease encoding gene cptA and hyprothetical protein encoding gene. Subsequenctly, three different suicide vectors (pMW91CM, pJQ200SK, pEX18GM) were evaluated for the possibility of performing gene mutagenesis in strain OH 11 using the chiA gene (accession number:DQ888611) as a new reporter. Suicide vector pEX18GM was selected, and it was successfully applied for disruption and in-frame deletions in the chiA gene in strain OH11. The chiA-deletion mutant OH11-3 did not have the ability to produce chitinase on chitine selection medium, whereas the corresponding complemented strain OP1 (chiA gene was expressed under an Escherichia coli constitutive promoter Plpp in strain OH11-3) was partially restored in chitinase production in the same medium. Interestingly, the chiA-deletion mutants displayed wild-type antimicrobial activity against Phytophthora capsici, Rhizoctonia solani, Sclerotinia sclerotiorum and Fusarium graminearum. Our data suggest that chitinase might not be a unique lytic enzyme in controlling R. solani, S. sclerotiorum and F. graminearum. Also, the established genetic manipulation system might be explored as a potential tool for functional gene deletion and complementation, and alien gene expression in L. enzymogenes, which will facilitate the molecular study of mechanisms of biological control in L. enzymogenes.
According to the phenotype changes, two pigment changing mutants, named strain OH11H and OH11B were isolated from the mutant library constructed by mariner transposon mutagenesis. In LA agar plates, wild-type OH11 exhibited a yellow colony, while strain OH11H produced dark brown pigment around yellow colonies, and strain OH11B was defective in yellow pigment production, resulting in a white colony. The transposon insertion in strain OH11H and OH11B was identified as hmgA and acp by using arbitrary PCR and sub-cloning technology. Based on the results of suicide-vector selections in a previous work, in this study, the hmgA gene was further used as a new reporter to confirm the results of suicide-vector selections, ultimately confirming that only pEX18GM is an available suicide vector for performing gene deletions in L. enzymogenes OH11. After that, the hmgA and acp disruption mutants were constructed by homologue recombination technology, respectively. Compared to wild-type OH11, the hmgA-disruption mutants changed the cell shape, enhanced the ability of biofilm formation, colonization in rice leaves and bicontrol efficiency against rice sheath blight. However, compared to wild-type OH11, the acp mutants showed a reduction of biofilm formation, colonization in rice leaves and biocontrol efficiency against rice sheat blight. Meanwhile, interestingly, on one hand, the hmgA and acp mutants exhibited wild-type tolerance to ultraviolet light (UV); on the other hand, the hmgA and acp mutation did not affect 4 type lytic enzyme production (chitinase, a-lytic protease, cellulase andβ-1,3-glucanase), and extropolysaccharide formation.
According to the phenotype changes,5 exopolysacchide (EPS)-defective mutants, named MEPS1-MESP5 were isolated from the mutant library constructed by mariner transposon mutagenesis. In LAS agar plates, wild-type OH11 displayed a plump and smooth colony, wherease the EPS-defective mutants exhibited dry, crimple and yellow colonies. By using arbitrary PCR, the transposon insertion in MEPS1-MESP5 was identified as tonB, permease encoding gene, pilin encoding gene and hypothetical protein encoding genes, respectively. After that, the whole tonB gene in strain MESP1 was obtained by using sub-cloning technology. Subsequently, the tonB deletion mutants were constructed by homologue technology. Compared to wild-type OH11, the tonB mutation in strain OH11 decreased gliding motility, tolerance to UV, the ability of biofilm formation, colonization in rice leaves and bicontrol efficiency against rice sheath blight, and changed the bacterial colony. However, the tonB mutants exhibited wild-type cell shape and ability of 4 type lytic enzyme production and antimicrobial activity against R. solani and Phytophthora capsici.
An N-acyl homoserine lactonase gene aiiA, transcribed by a strong and constitutive Escherichia coli promoter Plpp (accession no. EU723847), was transformed into strain OH11, creating strain OH11A. The AHL-degradation assay showed that transformant OH11A acquired the ability to degrade AHL molecules produced by Agrobacterium tumefaciens, Pectobacterium carotovorum, Pseudomonas syringae. pv. tomato strain DC3000 and Acidovorax avenae subsp. citrulli. Pathogenicity tests showed that while the parental strain OH11 did not reduce P. carotovorum infection, the transformant OH11A caused a strong reduction of Pectobacterium virulence on Chinese cabbages and cactus, whereas strain OH11A did not seem to interfere with the normal growth of this pathogen in cabbages. In antimicrobial activity assays, strain OH11A and OH11 showed similar antimicrobial activity against Phytophthora capsici and Sclerotinia sclerotiorum. In the following work, a promoter-trapping vector was constructed using lacZ as a reporter, and a strong and constitutive promoter, named PB500, was isolated from the promoter library of strain OH11 by using this promoter-trapping vector. Subsequently, promoter PB500 and aiiA was spliced by overlap PCR amplification, and the spliced gene PB500-aii4 was integrated into hmgA by homologue recombination technology, creating unmarked strain OHl1PA. Similar with strain OH11 A, strain OH11PA exhibited strong degradation ability to AHL signals (AHLA) produced by Agrobacterium tumefaciens, and was able to control soft rot of Chinese cabbages in vitro. This work provided a new strategy for developing genetically engineered multi-functional L. enzymogenes strains that possessed the ability to biologically control fungal pathogens and reduce bacterial pathogenicity.
首次利用mariner转座子对产酶溶杆菌OH11进行了转座诱变,成功构建产酶溶杆菌OH11的转座子随机插入文库。在几丁质选择性培养基上,根据菌落周围是否产生水解圈,成功从OH11突变株文库中筛选出4个几丁质酶缺失突变株。利用建立的任意PCR (abritrary PCR)技术,快速鉴定了mariner转座子在4个几丁质酶缺失突变株中的插入位点。序列分析表明插入位点涉及调控基因clp,几丁质酶编码基因chiA,羧基末端蛋白酶(carboxyl-terminal protease)编码基因cptA和未知蛋白(hyperthetical protein)编码基因。随后,以chiA为报道基因,分析了3种不同类型的自杀质粒(pMW91CM, pJQ200SK, pEX18GM)在OH11中用于完成基因定向敲除的可行性,确定了自杀质粒pEX18GM可以有效地在OH11中完成基因的定向插入失活和缺失敲除,并构建了chiA的缺失突变株。有趣的是,chiA缺失突变株不改变对水稻纹枯病菌(Rhizoctonia solani,)、油菜菌核病菌(Sclerotinia sclerotiorum)、小麦赤霉病菌(Fusarium graminearum)和辣椒疫霉(Phytophthora capsici)的拮抗活性,揭示在防治这四种病原真菌或卵菌时,几丁质酶并不是主要的胞外酶。利用来自于大肠杆菌的组成型强启动子Plpp,实现了chiA基因的互补及外源基因gfp的表达,成功构建了基于质粒的功能基因互补系统和外源基因表达系统。本部分建立的产酶溶杆菌OH11的遗作操作系统,为后续研究OH11的生防分子机理及工程菌的构建奠定了基础。
结合表型筛选和验证,从OH11的突变株文库中筛选到2株色素突变株,分别命名为OH11H和OH11B。在LA平板上,野生型OH11产生黄色素,菌落为黄色。色素突变株OH11H可以产生可溶性的黑色素,但其菌落仍为黄色；OH11B菌落形态为白色,缺失了黄色素的合成。通过任意PCR和亚克隆技术,鉴定了转座子在OH11H和OH11B中的插入位点分别为hmgA基因和acp基因。hmgA基因编码尿黑酸1,2-加氧化酶,参与酪氨酸的代谢。而acp基因编码3-羰基(酰基载体蛋白)合成酶Ⅰ,与脂肪酸生物合成(FAS)相关。在前期以chiA基因作为报告基因筛选自杀载体的基础上,本研究再次以hmgA为报告基因,验证不同类型的自身载体在OH11中完成基因定向敲除的可行性,并再次确认在参试的自杀载体中,只有pEX18GM才能完成靶标基因的定向缺失敲除,与前期筛选结果一致。通过同源重组技术,分别构建了hmgA和acp的定向敲除突变株,并明确hmgA的突变改变了OH11的细胞形态,使细胞从棍棒状变为椭圆状,但不改变菌落形态；提高了OH11生物膜形成能力、在水稻叶片上的定殖能力及对水稻纹枯病的防效,但不改变OH11胞外多糖形成能力、4种胞外酶的产生能力和平板拮抗活性。明确了acp的突变不改变OH11的细胞形状,但使细胞周围“多糖”类物质增加,降低了OH11生物膜形成能力,在水稻叶片上的定殖能力及对水稻纹枯病的防效,同样不改变OH11胞外多糖形成能力、4种胞外酶的产生能力和平板拮抗活性。此外,还初步明确了可溶性黑色素和菌体自身的黄色素可能与细菌抵御紫外光损伤无关。
结合表型筛选和验证,从OH11的突变株文库中筛选到5株胞外多糖(exopolysacchide, EPS)缺失突变株,分别命名为MEPS1-MEPS5。在LAS培养基上,野生型OH11菌落黄色、饱满、光滑,而EPS缺失突变株茵落干燥、皱缩,但菌落仍为黄色。通过任意PCR技术(arbitrary PCR),快速鉴定了转座子在MES1-MES5中的插入位点分别为别是tonB基因、透性酶(permease)编码基因、Ⅳ分泌系统中Pilin蛋白编码基因,以及未知蛋白编码基因,表明众多基因参与了EPS的合成。在任意PCR的基础上,进一步通过亚克隆技术(sub-cloning),克隆了tonB的完整基因。通过同源重组技术,构建了tonB定向缺失突变株,并明确tonB的突变降低了OH11的滑行能力,改变了OH11的菌落形态,但不改变菌体细胞形态；降低了OH11生物膜形成能力、在水稻叶片上的定殖能力及对水稻纹枯病的防效。初步明确了EPS可能与产酶溶杆菌抵御紫外光损伤相关。
利用大肠杆菌中的组成型强启动子Plpp实现了外源基因即AHL类信号分子降解基因aiiA在产酶溶杆菌OH11中的重组表达,构建了抗生素标记的工程菌株OH11A。菌株OH11A能显著降解不同植物病原细菌产生的AHL类信号分子。在离体生防试验中,OH11A能大幅度降低软腐病菌对大白菜的致病性,但不抑制软腐病菌在寄主组织中的生长。在活体生防试验中,OH11A能分别显著降低软腐病菌和西瓜果斑病菌对仙人掌和甜瓜苗的毒性。在后续工作中,我们以lacZ为报告基因,构建了基于广宿主载体pBBR1-CMS5的新型启动子探针pBBRLACZR。利用该探针,从产酶溶杆菌OH11的启动子文库中筛选到一个具有启动子功能的DNA片段,通过启动子亚克隆并结合反复的筛选验证,最终从OH11中克隆了一个组成型强表达的启动子PB500。以前期工作中克隆的hmgA为同源重组靶标基因,以SacB为反向筛选标记,将外源融合基因PB500-aiiA定向整合到hmgA中,构建了无标记、稳定遗传并能显著降解信号分子AHLA的工程菌株OH11PA.工程菌株OH11PA能显著降低软腐病菌在大白菜上的毒性。
Lysobacter enzymogenes strain oH11, isolated from cayenne-root soil, is reported for the first time as a bacterial biological control agent in China. It is characterizaed with a high G+C content and gliding motility, and has been shown to produce extracellular lytic enzymes, including chitinase, protease, cellulase andβ-1,3-glucanas. Furhtermore, it displays in vitro activity against fungal and oomycetous plant pathogens. The objectives of this work are to:(i) establish the genetic manipulation system in strain OH 11 and construct a mutant library by a mariner transposon mutagenesis; (ii) to identify the pigment and exopolysacchide formation related genes, and to investigate the function of these cloned genes in colonization, antimicrobial activity and biocontrol efficiency of strain OH11; (ii) to construct an engineering strain with AHL degradation capacity to biocontrol bacterial plant diseases, broadening the biocontrol spectrum of L. enzymogenes.
Transposon mutagenesis was first performed in Lysobacter enzymogenes strain OH 11 by a mariner transposon and a large-scale mutant library was obtained. Four chitinase-defective mutants, which could not produce hydrolytic zones around colonies in chitine selection medium, were selected from over 2,000 chloramphenicol-resistant (Cm1) mutants. By using arbitrary PCR techonology, the transposon insertion in 4 chitinase-defective mutants were identified as regulatory gene clp, chitinase encoding gene chiA, carboxyl-terminal protease encoding gene cptA and hyprothetical protein encoding gene. Subsequenctly, three different suicide vectors (pMW91CM, pJQ200SK, pEX18GM) were evaluated for the possibility of performing gene mutagenesis in strain OH 11 using the chiA gene (accession number:DQ888611) as a new reporter. Suicide vector pEX18GM was selected, and it was successfully applied for disruption and in-frame deletions in the chiA gene in strain OH11. The chiA-deletion mutant OH11-3 did not have the ability to produce chitinase on chitine selection medium, whereas the corresponding complemented strain OP1 (chiA gene was expressed under an Escherichia coli constitutive promoter Plpp in strain OH11-3) was partially restored in chitinase production in the same medium. Interestingly, the chiA-deletion mutants displayed wild-type antimicrobial activity against Phytophthora capsici, Rhizoctonia solani, Sclerotinia sclerotiorum and Fusarium graminearum. Our data suggest that chitinase might not be a unique lytic enzyme in controlling R. solani, S. sclerotiorum and F. graminearum. Also, the established genetic manipulation system might be explored as a potential tool for functional gene deletion and complementation, and alien gene expression in L. enzymogenes, which will facilitate the molecular study of mechanisms of biological control in L. enzymogenes.
According to the phenotype changes, two pigment changing mutants, named strain OH11H and OH11B were isolated from the mutant library constructed by mariner transposon mutagenesis. In LA agar plates, wild-type OH11 exhibited a yellow colony, while strain OH11H produced dark brown pigment around yellow colonies, and strain OH11B was defective in yellow pigment production, resulting in a white colony. The transposon insertion in strain OH11H and OH11B was identified as hmgA and acp by using arbitrary PCR and sub-cloning technology. Based on the results of suicide-vector selections in a previous work, in this study, the hmgA gene was further used as a new reporter to confirm the results of suicide-vector selections, ultimately confirming that only pEX18GM is an available suicide vector for performing gene deletions in L. enzymogenes OH11. After that, the hmgA and acp disruption mutants were constructed by homologue recombination technology, respectively. Compared to wild-type OH11, the hmgA-disruption mutants changed the cell shape, enhanced the ability of biofilm formation, colonization in rice leaves and bicontrol efficiency against rice sheath blight. However, compared to wild-type OH11, the acp mutants showed a reduction of biofilm formation, colonization in rice leaves and biocontrol efficiency against rice sheat blight. Meanwhile, interestingly, on one hand, the hmgA and acp mutants exhibited wild-type tolerance to ultraviolet light (UV); on the other hand, the hmgA and acp mutation did not affect 4 type lytic enzyme production (chitinase, a-lytic protease, cellulase andβ-1,3-glucanase), and extropolysaccharide formation.
According to the phenotype changes,5 exopolysacchide (EPS)-defective mutants, named MEPS1-MESP5 were isolated from the mutant library constructed by mariner transposon mutagenesis. In LAS agar plates, wild-type OH11 displayed a plump and smooth colony, wherease the EPS-defective mutants exhibited dry, crimple and yellow colonies. By using arbitrary PCR, the transposon insertion in MEPS1-MESP5 was identified as tonB, permease encoding gene, pilin encoding gene and hypothetical protein encoding genes, respectively. After that, the whole tonB gene in strain MESP1 was obtained by using sub-cloning technology. Subsequently, the tonB deletion mutants were constructed by homologue technology. Compared to wild-type OH11, the tonB mutation in strain OH11 decreased gliding motility, tolerance to UV, the ability of biofilm formation, colonization in rice leaves and bicontrol efficiency against rice sheath blight, and changed the bacterial colony. However, the tonB mutants exhibited wild-type cell shape and ability of 4 type lytic enzyme production and antimicrobial activity against R. solani and Phytophthora capsici.
An N-acyl homoserine lactonase gene aiiA, transcribed by a strong and constitutive Escherichia coli promoter Plpp (accession no. EU723847), was transformed into strain OH11, creating strain OH11A. The AHL-degradation assay showed that transformant OH11A acquired the ability to degrade AHL molecules produced by Agrobacterium tumefaciens, Pectobacterium carotovorum, Pseudomonas syringae. pv. tomato strain DC3000 and Acidovorax avenae subsp. citrulli. Pathogenicity tests showed that while the parental strain OH11 did not reduce P. carotovorum infection, the transformant OH11A caused a strong reduction of Pectobacterium virulence on Chinese cabbages and cactus, whereas strain OH11A did not seem to interfere with the normal growth of this pathogen in cabbages. In antimicrobial activity assays, strain OH11A and OH11 showed similar antimicrobial activity against Phytophthora capsici and Sclerotinia sclerotiorum. In the following work, a promoter-trapping vector was constructed using lacZ as a reporter, and a strong and constitutive promoter, named PB500, was isolated from the promoter library of strain OH11 by using this promoter-trapping vector. Subsequently, promoter PB500 and aiiA was spliced by overlap PCR amplification, and the spliced gene PB500-aii4 was integrated into hmgA by homologue recombination technology, creating unmarked strain OHl1PA. Similar with strain OH11 A, strain OH11PA exhibited strong degradation ability to AHL signals (AHLA) produced by Agrobacterium tumefaciens, and was able to control soft rot of Chinese cabbages in vitro. This work provided a new strategy for developing genetically engineered multi-functional L. enzymogenes strains that possessed the ability to biologically control fungal pathogens and reduce bacterial pathogenicity.
引文
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1. Bourhy, P., Louvel, H., Saint Girons, I., Picardeau, M. (2005) Random Insertional Mutagenesis of Leptospira interrogans, the Agent of Leptospirosis, Using a mariner Transposon. Journal of Bacteriology 189(9):3255-3258.
2. Chen, Q., Wu, H., Fives-Taylorl, P. M. (2002) Construction of a Novel Transposon Mutagenesis System Useful in the Isolation of Streptococcus parasanguis Mutants Defective in Fapl Glycosylation. Infection and Immunity 70(12):6534-6540
3. Chiang, S.L., Mekalanos, J.J. (1999) rfb mutations in Vibrio cholerae do not affect surface production of toxincoregulated pili but still inhibit intestinal colonization. Infect Immun.67: 976-980.
4. Folman, L. B., Postma, J., Van-Veen, J. A. (2003) Characterisation of Lysobacter enzymogenes (Christensen and Cook 1978) strain 3.1T8, a powerful antagonist of fungal diseases of cucumber. Research in Microbiology 158:1-9.
5. Gao, L. Y., Groger, R., Cox, J. S., Beverley, S. M., Lawson, E. H., Brown, E. J. (2003) Transposon Mutagenesis of Mycobacterium marinum Identifies a Locus Linking Pigmentation and Intracellular Survival Locus Linking Pigmentation and Intracellular Survival. Infection and Immunity 71(2): 922-929
6. Gao, Y. J., Zhong, Z. T., Sun, K. J., Wang, H., Zhu, J. (2006) The quorum-sensing system in a plant bacterium Mesorhizobium huakuii affects growth rate and symbiotic nodulation. Plant Soil 286: 53-60
7. Gay, P., Coq, D. L., Steinmetz, M., Berkelman, T., Kado, C. I. (1985) Positive selection procedure for entrapment of insertion sequence elements in gram-negative bacteria. Journal of Bacteriology 164:918-921.
8. Giesler, L. J., Yuen, G. Y. (1998) Evaluation of Stenotrophomonas maltophilia strain C3 for biocontrol of brown patch disease. Crop Protection 17:509-513.
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12. Kobayashi, D. Y., Barrad, N. E. (1996) Selection of bacterial antagonists using enrichment cultures for the control of summer patch disease of Kentucky bluegrass. Current Microbiology 32:106-110.
13. Kobayashi, D.Y., Reedy, R. M., Palumbo, J. D., Zhou, J. M., Yuen, G. Y.2005. A clp Gene Homologue Belonging to the Crp Gene Family Globally Regulates Lytic Enzyme Production, Antimicrobial Activity, and Biological Control Activity Expressed by Lysobacter enzymogenes Strain C3. Applied and Enviromental Microbilogy 71:261-269
14. Kovach, M. E., Elzer, P. H., Hill, D. S., Robertson, G. T., Farris, M. A., Roop, R. M., Peterson, K. M (1995) Four new derivatives of the broad-host-range cloning vector pBBR1MCS, carrying different antibiotic-resistance cassettes. Gene 166:175-176.
15. Metcalf, W.W., Jiang, W., Daniels, L.L., Kim, S.K., Haldimann, A., Wanner, B.L. (1996) Conditionally replicative and conjugative plasmids carrying lacZ alpha for cloning, mutagenesis, and allele replacement in bacteria. Plasmid 35:1-13.
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17. Palumbo, J. D., Sullivan, R. F., Kobayashi, D. Y. (2003) Molecular Characterization and Expression in Escherichia coli of Three β-1,3-Glucanase Genes from Lysobacter enzymogenes Strain N4-7. Journal of Bacteriology 185:4362-4370
18. Palumbo, J. D., Yuen, G. Y., Jochum, C. C., Tatum, K., Kobayashi, D. Y. (2005) Mutagenesis of (3-1,3-Glucanase Genes in Lysobacter enzymogenes strain C3 Results in Reduced Biological Control Activity Toward Bipolaris Leaf Spot of Tall Fescue and Phythium Damping-off of Sugar Beet. Phytopathology 95(6):701-707.
19. Quandt, J., Hynes, M.F. (1993) Versatile suicide vectors which allow direct selection for gene replacement in gram-negative bacteria. Gene 127:15-21.
20. Reimmann, C., Ginet, N., Michel, L., Keel, C., Michaux, P., Krishnapillai, V., Zala, M., Heurlier, K., Triandafillu, K., Harms,H., Defago,G., Haas. D. (2002) Genetically programmed autoinducer destruction reduces virulence gene expression and swarming motility in Pseudomonas aeruginosa PAO1.Microbiology 148:923-932.
21. Sullivan, R. F., Holtman, M. A., Zylstra, G. J., White, J. F., J. r., Kobayashi, D. Y. (2003) Taxonomic positioning of two biological control agents for plant diseases as Lysobacter enzymogenes based on phylogenetic analysis of 16S rDNA, fatty acid composition and phenotypic characteristics. Journal of Applied Microbiology 94:1079-1086.
22. Yu, F., Zaleta-Rivera, K., Zhu, X., Huffman, J., Millet, J. C., Harris, S. D., Yuen, G., Li, X. C., Du, L. (2007) Structure and Biosynthesis of Heat-Stable Antifungal Factor (HSAF), a Broad-Spectrum Antimycotic with a Novel Mode of Action. Anitimicrobial Agents and Chemotherapy 51(1):64-72
23. Zhang, Z. G., Yuen, G. Y. (2000) The role of chitinase production by Stenotrophomonas maltophilia strain C3 in biological control of Bipolaris sorokiniana. Phytopathology.90 (4):384-389
24. Zhang, Z., Yuen, G. Y., Sarath, G., Penheiter, A. R. (2001) Chitinase from the plant disease biocontrol agent Stenotrophomonas maltophilia C3. Phytopathology 91 (2): 204-211.
25.韩冰(2008)产酶溶杆菌OH11菌株clp基因的克隆及功能分析。硕士学位论文。南京农业大学。
26.姜英华(2006)一种新型生防菌菌株OH11的鉴定及生防效果研究。硕士学位论文。南京农业大学。
27.王云霞,钱国良,胡白石,刘凤权(2008)产酶溶杆菌OH11菌株摇瓶发酵条件研究。中国生物防治24：267-271。
28.朱慧,王云霞,胡白石,刘凤权(2007)产酶溶杆菌OH11菌株α-水解蛋白酶基因的克隆与表达。中国生物防治23：358-362。
29.朱慧,王云霞,胡白石,刘凤权(2008)产酶溶杆菌OH11菌株β-1,3-葡聚糖酶基因的克隆与表达。江苏农业科学24：136-141。
30.朱慧,王云霞,胡白石,刘凤权(2008)产酶溶杆菌OH11菌株几丁质酶基因的克隆与表达。南京农业大学学报31：47-50。
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3. Giesler, L. J., Yuen, G. Y. (1998) Evaluation of Stenotrophomonas maltophilia strain C3 for biocontrol of brown patch disease. Crop Protection 17:509-513.
4. Hoang, T.T., Karkhoff-Schweizer, R.R., Kutchma, A.J., Schweizer, H.P. (1998) A broad-host-range p-FRT recombination system for site-specific excision of chromosomally-located DNA sequences: application for isolation of unmarked Pseudomonas aeruginosa mutants. Gene 212:77-86.
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7. Kobayashi, D. Y., Barrad, N. E. (1996) Selection of bacterial antagonists using enrichment cultures for the control of summer patch disease of Kentucky bluegrass. Current Microbiology 32:106-110.
8. Kobayashi, D.Y., Reedy, R. M., Palumbo, J. D., Zhou, J. M., Yuen, G. Y.2005. A clp gene homologue belonging to the Crp Gene Family globally regulates lytic enzyme production, antimicrobial activity, and biological control activity expressed by Lysobacter enzymogenes Strain C3. Applied and Environmental Microbiology 71:261-269
9. Kovach, M. E., Elzer, P. H., Hill, D. S., Robertson, G. T., Farris, M. A., Roop, R. M., Peterson, K. M (1995) Four new derivatives of the broad-host-range cloning vector pBBR1MCS, carrying different antibiotic-resistance cassettes. Gene 166:175-176.
10. Li, S. J., Jochum, C. C., Yu, F. G., Zaleta-Rivera, K., Du, L. C., Harris, S. D., Yuen, G. Y. (2008) An antibiotic complex from Lysobacter enzymogenes strain C3:antimicrobial activity and role in plant disease control. Phytopathology 98(6):695-701
11. Metcalf, W.W., Jiang, W., Daniels, L.L., Kim, S.K., Haldimann, A., Wanner, B.L. (1996) Conditionally replicative and conjugative plasmids carrying lacZ alpha for cloning, mutagenesis, and allele replacement in bacteria. Plasmid 35:1-13.
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16. Palumbo, J. D., Yuen, G. Y., Jochum, C. C, Tatum, K., Kobayashi, D. Y. (2005) Mutagenesis of β-1,3-Glucanase Genes in Lysobacter enzymogenes strain C3 Results in Reduced Biological Control Activity Toward Bipolaris Leaf Spot of Tall Fescue and Phythium Damping-off of Sugar Beet. Phytopathology 95(6):701-707.
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27. Zhang, Z. G., Yuen, G. Y. (2000) The role of chitinase production by Stenotrophomonas maltophilia strain C3 in biological control of Bipolaris sorokiniana. Phytopathology.90 (4):384-389
28. Zhang, Z., Yuen, G. Y, Sarath, G., Penheiter, A. R. (2001) Chitinase from the plant disease biocontrol agent Stenotrophomonas maltophilia C3. Phytopathology 91 (2):204-211.
29.韩冰(2008)产酶溶杆菌OH11菌株clp基因的克隆及功能分析。硕士学位论文。南京农业大学。
30.姜英华(2006)一种新型生防菌菌株OH11的鉴定及生防效果研究。硕士学位论文。南京农业大学。
31.姜英华,胡白石,刘凤权(2005)植物土传病原菌拮抗细菌的筛选与鉴定。中国生物防治21(4)：260-264
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35.朱慧,王云霞,胡白石,刘凤权(2007)产酶溶杆菌OH11菌株α-水解蛋白酶基因的克隆与表达。中国生物防治23：358-362。
36.朱慧,王云霞,胡白石,刘凤权(2008)产酶溶杆菌OH11菌株β-1,3-葡聚糖酶基因的克隆与表达。江苏农业科学24：136-141。
37.朱慧,王云霞,胡白石,刘凤权(2008)产酶溶杆菌OH11菌株几丁质酶基因的克隆与表达。南京农业大学学报31：47-50。
1. Bellemann, P., Geider, K. (1992) Localization of transposon insertions in pathogenicity mutants of Erwinia amylovora and their biochemical characterization. Journal of General Microbiology 138: 931-940.
2. Bourhy, P., Louvel, H., Saint Girons, I., Picardeau, M. (2005) Random Insertional Mutagenesis of Leptospira interrogans, the Agent of Leptospirosis, Using a mariner Transposon. Journal of Bacteriology 189(9):3255-3258.
3. Chen, Q., Wu, H., Fives-Taylorl, P. M. (2002) Construction of a Novel Transposon Mutagenesis System Useful in the Isolation of Streptococcus parasanguis Mutants Defective in Fapl Glycosylation. Infection and Immunity 70(12):6534-6540
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5. Folman, L. B., Postma, J., Van-Veen, J. A. (2003) Characterisation of Lysobacter enzymogenes (Christensen and Cook 1978) strain 3.1T8, a powerful antagonist of fungal diseases of cucumber. Research in Microbiology 158:1-9.
6. Gao, L. Y., Groger, R., Cox, J. S., Beverley, S. M., Lawson, E. H., Brown, E. J. (2003) Transposon Mutagenesis of Mycobacterium marinum Identifies a Locus Linking Pigmentation and Intracellular Survival Locus Linking Pigmentation and Intracellular Survival. Infection and Immunity 71(2): 922-929
7. Gao, Y. J., Zhong, Z. T., Sun, K. J., Wang, H., Zhu, J. (2006) The quorum-sensing system in a plant bacterium Mesorhizobium huakuii affects growth rate and symbiotic nodulation. Plant Soil 286: 53-60
8. Gay, P., Coq, D. L., Steinmetz, M., Berkelman, T., Kado, C. I. (1985) Positive selection procedure for entrapment of insertion sequence elements in gram-negative bacteria. Journal of Bacteriology 164:918-921.
9. Giesler, L. J., Yuen, G. Y. (1998) Evaluation of Stenotrophomonas maltophilia strain C3 for biocontrol of brown patch disease. Crop Protection 17:509-513.
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11. Knobloch, J. K. M., Nedelmann, M., Kiel, K., Bartscht, K., Horstkotte, M. A., Dobinsky, S., Rohde, H., Mack, D. (2003) Establishment of an Arbitrary PCR for Rapid Identification of Tn917 Insertion Sites in Staphylococcus epidermidis:Characterization of Biofilm-Negative and Nonmucoid Mutants. Applied and Enviromental Microbilogy 69(10):5812-5818.
12. Kobayashi, D. Y., Barrad, N. E. (1996) Selection of bacterial antagonists using enrichment cultures for the control of summer patch disease of Kentucky bluegrass. Current Microbiology 32:106-110.
13. Kobayashi, D.Y., Reedy, R. M., Palumbo, J. D., Zhou, J. M., Yuen, G. Y.2005. A clp Gene Homologue Belonging to the Crp Gene Family Globally Regulates Lytic Enzyme Production, Antimicrobial Activity, and Biological Control Activity Expressed by Lysobacter enzymogenes Strain C3.Appl Environ Microbiol71:261-269
14. Kovach, M. E., Elzer, P. H., Hill, D. S., Robertson, G. T., Farris, M. A., Roop, R. M., Peterson, K. M (1995) Four new derivatives of the broad-host-range cloning vector pBBR1MCS, carrying different antibiotic-resistance cassettes. Gene 166:175-176.
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17. Palumbo, J. D., Sullivan, R. F., Kobayashi, D. Y. (2003) Molecular Characterization and Expression in Escherichia coli of Three β-1,3-Glucanase Genes from Lysobacter enzymogenes Strain N4-7. Journal of Bacteriology 185:4362-4370
18. Palumbo, J. D., Yuen, G. Y., Jochum, C. C., Tatum, K., Kobayashi, D. Y. (2005) Mutagenesis of (3-1,3-Glucanase Genes in Lysobacter enzymogenes strain C3 Results in Reduced Biological Control Activity Toward Bipolaris Leaf Spot of Tall Fescue and Phythium Damping-off of Sugar Beet. Phytopathology 95(6):701-707.
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23. Yu, F., Zaleta-Rivera, K., Zhu, X., Huffman, J., Millet, J. C., Harris, S. D., Yuen, G., Li, X. C., Du, L. (2007) Structure and Biosynthesis of Heat-Stable Antifungal Factor (HSAF), a Broad-Spectrum Antimycotic with a Novel Mode of Action. Anitimicrobial Agents and Chemotherapy 51(1):64-72
24. Zhang, Z. G., Yuen, G. Y. (2000) The role of chitinase production by Stenotrophomonas maltophilia strain C3 in biological control of Bipolaris sorokiniana. Phytopathology.90 (4):384-389
25. Zhang, Z., Yuen, G. Y., Sarath, G., Penheiter, A. R. (2001) Chitinase from the plant disease biocontrol agent Stenotrophomonas maltophilia C3. Phytopathology 91 (2):204-211.
26.韩冰(2008)产酶溶杆菌OH11菌株clp基因的克隆及功能分析。硕士学位论文。南京农业大学。
27.姜英华(2006)一种新型生防菌菌株OH11的鉴定及生防效果研究。硕士学位论文。南京农业大学。
28.王云霞,钱国良,胡白石,刘凤权(2008)产酶溶杆菌OH11菌株摇瓶发酵条件研究。中国生物防治24：267-271。
29.朱慧,王云霞,胡白石,刘凤权(2007)产酶溶杆菌OH11菌株α-水解蛋白酶基因的克隆与表达。中国生物防治23：358-362。
30.朱慧,王云霞,胡白石,刘凤权(2008)产酶溶杆菌OH11菌株β-1,3-葡聚糖酶基因的克隆与表达。江苏农业科学24：136-141。
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1. Bourhy, P., Louvel, H., Saint Girons, I., Picardeau, M. (2005) Random Insertional Mutagenesis of Leptospira interrogans, the Agent of Leptospirosis, Using a mariner Transposon. Journal of Bacteriology 189(9):3255-3258.
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18. Palumbo, J. D., Yuen, G. Y., Jochum, C. C., Tatum, K., Kobayashi, D. Y. (2005) Mutagenesis of (3-1,3-Glucanase Genes in Lysobacter enzymogenes strain C3 Results in Reduced Biological Control Activity Toward Bipolaris Leaf Spot of Tall Fescue and Phythium Damping-off of Sugar Beet. Phytopathology 95(6):701-707.
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22. Yu, F., Zaleta-Rivera, K., Zhu, X., Huffman, J., Millet, J. C., Harris, S. D., Yuen, G., Li, X. C., Du, L. (2007) Structure and Biosynthesis of Heat-Stable Antifungal Factor (HSAF), a Broad-Spectrum Antimycotic with a Novel Mode of Action. Anitimicrobial Agents and Chemotherapy 51(1):64-72
23. Zhang, Z. G., Yuen, G. Y. (2000) The role of chitinase production by Stenotrophomonas maltophilia strain C3 in biological control of Bipolaris sorokiniana. Phytopathology.90 (4):384-389
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25.韩冰(2008)产酶溶杆菌OH11菌株clp基因的克隆及功能分析。硕士学位论文。南京农业大学。
26.姜英华(2006)一种新型生防菌菌株OH11的鉴定及生防效果研究。硕士学位论文。南京农业大学。
27.王云霞,钱国良,胡白石,刘凤权(2008)产酶溶杆菌OH11菌株摇瓶发酵条件研究。中国生物防治24：267-271。
28.朱慧,王云霞,胡白石,刘凤权(2007)产酶溶杆菌OH11菌株α-水解蛋白酶基因的克隆与表达。中国生物防治23：358-362。
29.朱慧,王云霞,胡白石,刘凤权(2008)产酶溶杆菌OH11菌株β-1,3-葡聚糖酶基因的克隆与表达。江苏农业科学24：136-141。
30.朱慧,王云霞,胡白石,刘凤权(2008)产酶溶杆菌OH11菌株几丁质酶基因的克隆与表达。南京农业大学学报31：47-50。
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3. Giesler, L. J., Yuen, G. Y. (1998) Evaluation of Stenotrophomonas maltophilia strain C3 for biocontrol of brown patch disease. Crop Protection 17:509-513.
4. Hoang, T.T., Karkhoff-Schweizer, R.R., Kutchma, A.J., Schweizer, H.P. (1998) A broad-host-range p-FRT recombination system for site-specific excision of chromosomally-located DNA sequences: application for isolation of unmarked Pseudomonas aeruginosa mutants. Gene 212:77-86.
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27. Zhang, Z. G., Yuen, G. Y. (2000) The role of chitinase production by Stenotrophomonas maltophilia strain C3 in biological control of Bipolaris sorokiniana. Phytopathology.90 (4):384-389
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3. Chen, Q., Wu, H., Fives-Taylorl, P. M. (2002) Construction of a Novel Transposon Mutagenesis System Useful in the Isolation of Streptococcus parasanguis Mutants Defective in Fapl Glycosylation. Infection and Immunity 70(12):6534-6540
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