渐绿木霉基因组中PKS和NRPS基因的多样性分析
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摘要
为了探究渐绿木霉(Trichoderma viridescens)基因组中聚酮合成酶(polyketide synthase,PKS)和非核糖体多肽合成酶(non-ribosomal peptide synthase,NRPS)的多样性,为寻找合成生物活性物质的潜在菌株提供科学依据。本研究通过18S rDNA序列分析,鉴定菌株SC-1为渐绿木霉(T.viridescens);利用兼并性引物对菌株SC-1的PKS和NRPS基因进行PCR扩增及序列测定分析,构建系统发育树,明确菌株SC-1的PKS和NRPS基因序列的系统进化地位。结果表明,菌株SC-1基因组中含有PKS I、NRPS基因片段;经比对,PKS I基因片段与已知Trichoderma atroviride PKS序列相似度为96%,NRPS基因片段与Trichoderma gamsii NRPS序列相似度为97%。本研究结果为渐绿木霉具有产生活性次生代谢产物的潜力,值得进一步开发研究和应用。
To explore the inheritance in Trichoderma viridescens and the diversity of the polyketide synthase(PKS)and non-ribosomal peptide synthase(NRPS) genes and to lay the foundation of looking for the synthesis of the potential strains of bioactive substances.A strain of SC-1 was identified as T.viridescens on the basis of 18S rDNA.PKS and NRPS gene fragments were PCR amplified from genomic DNA with degenerate primers.The biosynthetic potential of strain SC-1 was accessed by bioinformatic and phylogenetic analyses of PKS and NRPS gene sequences.The results showed that the strain of SC-1 contains PKS I and NRPS gene.The PKS I gene sequence had 96% homology compared with Trichoderma atroviride PKS gene.The NRPS gene sequence had 97% homology with Trichoderma gamsii NRPS gene.This study demonstrated that T.viridescens had potential to synthesize bioactive secondary metabolite,therefore,their natural products are worth of further research and development.
To explore the inheritance in Trichoderma viridescens and the diversity of the polyketide synthase(PKS)and non-ribosomal peptide synthase(NRPS) genes and to lay the foundation of looking for the synthesis of the potential strains of bioactive substances.A strain of SC-1 was identified as T.viridescens on the basis of 18S rDNA.PKS and NRPS gene fragments were PCR amplified from genomic DNA with degenerate primers.The biosynthetic potential of strain SC-1 was accessed by bioinformatic and phylogenetic analyses of PKS and NRPS gene sequences.The results showed that the strain of SC-1 contains PKS I and NRPS gene.The PKS I gene sequence had 96% homology compared with Trichoderma atroviride PKS gene.The NRPS gene sequence had 97% homology with Trichoderma gamsii NRPS gene.This study demonstrated that T.viridescens had potential to synthesize bioactive secondary metabolite,therefore,their natural products are worth of further research and development.
引文
Amnuaykanj anasin A.,Punya J.,Paungmoung P.,Rungrod A.,Tachaleat A.,Pongpattanakitshote S.,Cheevadhanarak S.,and Tanticharoen M.,2005,Diversity of type I polyketide synthase genes in the wood-decay fungus Xylaria,sp.BCC1067,FEMS Microbiol.Lett.,251(1):125-136
Bingle L.E.,Simpson T.J.,and Lazarus C.M.,1999,Ketosynthase domain probes identify two subclasses of fungal polyketide synthase genes,Fungal Genet.Biol.,26(3):209-223
Chang Z.,Flatt P.,Gerwick W.H.,Nguyen V.A.,Willis C.L.,and Sherman D.H.,2002,The barbamide biosynthetic gene cluster:a novel marine cyanobacterial system of mixed polyketide synthase(PKS)-non-ribosomal peptide synthetase(NRPS)origin involving an unusual trichloroleucyl starter unit,Gene,296(1-2):235-247
Chen R.Q.,Liao L.,Zhang X.H.,and Chen B.,2014,Cloning and analysis of a halogenase gene of Streptomyces sp.604F from the Arctic Ocean,Weishengwu Xuebao(Acta Microbiologica Sinica),54(6):703-712(陈瑞勤,廖丽,张晓华,陈波,2014,北极海洋链霉菌604F的卤化酶基因克隆及特征,微生物学报,54(6):703-712)
Crawford J.M.,Korman T.P.,Labonte J.W.,Vagstad A.L.,and Hill E.A.,2009,Structural basis for biosynthetic programming of fungal aromatic polyketide cyclization,Nature,461(7267):1139-1143
Demain A.L.,and Sanchez S.,2009,Microbial drug discovery:80 years of progress,J.Antibiot.,62(1):5-16
Feng J.F.,Zhou R.C.,Guo X.T.,and Zhang Y.,2011,Polyketide and its application,Xiandai Nongye Keji(Modern Agricultural Sciences and Technology),(3):24-26(冯健飞,周日成,郭兴庭,张扬,2011,聚酮类化合物及其应用,现代农业科技,(3):24-26)
Harman G.E.,2006,Overview of mechanisms and uses of Trichoderma spp.,Phytopathology,96(2):190-194
Harman G.E.,2011,Trichoderma-not just for biocontrol anymore,Phytoparasitica,39(2):103-108
Harman G.E.,Howell C.R.,Viterbo A.,Chet I.,and Lorito M.,2004,Trichoderma species-opportunistic,avirulent plant symbionts,Nat.Rev.Microbiol.,2(1):43-56
Huang H.,and Hu H.f.,2016,Evolution and diversity of the gene clusters in microbial medicine biosyntheses,Shijie Linchuang Yaowu(World Clinical Drugs),37(2):129-134(黄鹤,胡海峰,2016,微生物药物合成基因簇的进化与结构多样性,世界临床药物,37(2):129-134)
Li J.W.H.,and Vederas J.C.,2009,Drug discovery and natural products:end of an era or an endless frontier,Science,325(5937):161-165
Li Q.,Li X.L.,Huang Y.J.,Chen C.,Huang W.L.,Xiong C.,and Zheng L.Y,2016,Biocontrol effect of Trichoderma virides-%cens strain 18-21 screened from fruiting bodies of Trametes gibbosa,Weishengwuxue Zazhi(Tournal of Microbiology),36(1):17-21(李强,李小林,黄羽佳,陈诚,黄文丽,熊川,郑林用,2016,偏肿栓菌内生木霉菌株18-21抑菌效果研究,微生物学杂志,36(1):17-21)
Lin M.J.,and Ko W.H.,2008,Occurrence of isolates of phytophthora colocasiae in Taiwan with homothallic behavior and its significance,Mycologia,100(5):727-734
Lu Y.,Lu Z.X.,He F.L.,Yuan G.Q.,Chen B.S.,and Liao Y.M.,2013,A research on mating types of Phytophthora colocasiae in south-eastern guangxi province,Jiyinzuxue Yu Yingyong Shengwuxue(Genomics and Applied Biology),32(3):285-290(陆叶,陆志翔,何芳练,袁高庆,陈保善,廖咏梅,2013,桂东南地区芋疫霉的交配型研究,基因组学与应用生物学,32(3):285-290)
Lucas-Elio P.,Hernandez P.,Sanchez-Amat A.,and Solano F.,2005,Purification and partial characterization of marinocine,a new broad-spectrum antibacterial protein produced by marinomonas mediterranea,Biochim.Biophys.Acta.,1721(1-3):193-203
Lv M.,Yang K.,Wang A.,Shi Y.N.,and Xin Z.H.,2016,Isolation and identification based on NRPS functional gene of a fungal epiphytic strain from grapes,Shipin Gongye Keji(Science and Technology of Food Industry),37(18):229-240(吕曼,杨柯,王傲,史雅凝,辛志宏,2016,基于NRPS基因筛选与鉴定葡萄附生真菌,食品工业科技,37(18):229-240)
Maansson M.,Vynne N.G.,Klitgaard A.,Nybo J.L.,Melchiorsen J.,Nguyen D.D.,Sanchez L.M.,Ziemert N.,Dorrestein P.C.,Andersen M.R.,and Gram L.,2016,An integrated metabolomic and genomic mining workflow to uncover the biosynthetic potential of bacteria,American Society for Microbiology,1(3):15-28
Mcdonald J.G.,Smith D.D.,Stiles A.R.,and Russell D.W.,2012,A comprehensive method for extraction and quantitative analysis of sterols and secosteroids from human plasma,J.Lipid Res.,53(7):1399-1409
Misiek M.,and Hoffmeister D.,2007,Fungal genetics,genomics,and secondary metabolites in pharmaceutical sciences,Planta Med.,73(2):103-115
Piel J.,2010,Biosynthesis of polyketides by trans-AT polyketide synthases,Natural Product Reports,27(7):996-1047
Ren D.,Zhang B.,Zhang X.P.,Teng Y.,Li X.L.,and Gu Y.F.,2014,Inheritance in endophytic fungi from Melia toosendan and diversity of genes of PKS and NRPS,Zhongcaoyao(Chinese Traditional and Herbal Drugs),45(10):1461-1467(任丹,张波,张小平,腾芸,李小林,辜运富,2014,川楝内生真菌的遗传及PKS、NRPS基因的多样性,中草药,45(10):1461-1467)
Schümann J.,and Hertweck C.,2006,Advances in cloning,functional analysis and heterologous expression of fungal polyketide synthase genes,J.Biotechnol.,124(4):690-703
Slightom J.L.,Metzger B.P.,Luu H.T.,and Elhammer A.P.,2009,Cloning and molecular characterization of the gene encoding the aureobasidin a biosynthesis complex in aureobasidium pullulans bp-1938,Gene,431(1-2):67-79
Tamura K.,Dudley J.,Nei M.,and Kumar S.,2007,MEGA4:molecular evolutionary genetics analysis(MEGA)software version 4.0,Mol.Biol.Evol.,24(8):1596-1599
Wang S.Y.,2007,Advances in the study of the mechanism and application of nonribosomal peptide synthetases,Weishengwu Xuebao(Acta Microbiologica Sinica),74(4):734-737(王世媛,2007,非核糖体肽合成酶(NRPSs)作用机理与应用的研究进展,微生物学报,74(4):734-737)
Wen Y.P.,2011,Analyses of the antibiotic%Paenimacrolidin and the siderophore paenibactin based on PKS and NRPS genes,Dissertation for Ph.D.,Zheiiang University,Supervisor:Zhao Y.H.,pp.5-7(文艳苹,2011,基于PKS,NRPS基因的抗生素Paenimacrolidin和嗜铁素Paenibactin研究,博士学位论文,浙江大学,导师:赵宇华,pp.5-7)
Xiao J.,Zhang G.T.,Zhu Y.G.,and Zhang C.S.,2012,Progress in biosynthesis of secondary metabolites from marine microorganisms,Zhongguo Kangshengsu Zazhi(Chinese Journal of Antibiotics),37(4):241-253(肖吉,张光涛,朱义广,张长生,2012,海洋微生物次级代谢产物生物合成的研究进展,中国抗生素杂志,37(4):241-253)
Zhang G.Z.,Yand H.T.,Zhang X.J.,Hu J.D.,Guo K.,Huang Y.J.,and Li J.S.,2014,A checklist of known species of trichoderma,Junwu Xuebao(Mycosystema),33(6):1210-1230(张广志,杨合同,张新建,扈进冬,郭凯,黄玉杰,李纪顺,2014,木霉现有种类名录,菌物学报,33(6):1210-1230)
Zhang J.N.,Peng J.,Liu T.X.,and Xin Z.H.,2016,Isolation and identification of endophyte and its secondary metabolites from%Salicornia bigelovii Torr.based on type i polyketide synthase(PKS I)gene,Shipin Kexue(Food Science),37(1):114-119(张俊楠,彭洁,刘天行,辛志宏,2016,基于PKSⅠ基因海芦笋内生真菌及其次级代谢产物筛选鉴定,食品科学,37(1):114-119)
Zhang L.,and Zhang J.Z.,2015,Isolation and purification of active compound from Trichoderma viridescens and its inhibitory activities against phytopathogens,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),48(5):882-888(张量,张敬泽,2015,渐绿木霉抑菌物质的分离纯化及其对植物病原菌的抑制作用,中国农业科学,48(5):882-888)
Zhong Z.M.,Lai X.P.,Huang S.,and Zhang G.F.,2018,Optimization of DNA extraction methods and molecular identification of endophytic Xylaria from Dendrobium,Junwu Xuebao(Mycosystema),37(1):73-78(钟志敏,赖小平,黄松张桂芳,2018,石斛内生炭角菌DNA提取方法优化及分子鉴定,菌物学报,37(1):73-78)
Zhou P.J.,Xu Y.,Ma Y.Q.,and Liu H.D.,1994,Amplification of16S r DNAs from halobacteria by means of PCR technique Weishengwu Xuebao(Acta Microbiologica Sinica),34(1):6-8(周培瑾,徐毅,马允卿,刘宏迪,1994,极端嗜盐菌16Sr DNA的PCR扩增,微生物学报,34(1):6-8)
Zhu M.Z.,Cui X.L.,Li M.G.,Li Y.Q.,Zhao J.Y.,Peng Q.,and Wen M.L.,2006,A new approach for microbial drugs discovery:from genes to novel compounds,Tianran Chanwu Yanjiu Yu Fazhan(Natural Product Research and Development)18(5):854-857(朱孟沼,崔晓龙,李铭刚,李一青,赵江源彭谦,文孟良,2006,发现微生物药物的新途径:从基因到新化合物,天然产物研究与开发,18(5):854-857)
Zhu P.,2008,Screening and studying modular structure of PKSand NRPS genes in marine sponge Hymeniacidon perleve associated bacteria,Dissertation for Ph.D.,School of Life Sciences,Zhejiang University,Supervisor:Shao J.Z.,pp.27-29(朱鹏,2008,珍珠膜海绵共附生微生物PKS与NRPS的筛选与模块结构研究,博士学位论文,浙江大学,导师:邵健忠,pp.27-29)
Zhu P.,Zheng L.,Li J.,Shao J.Z.,and Yan X.J.,2007,Screening and characterization of marine bacteria with antibacterial and cytotoxic activities,and existence of PKSⅠand NRPSgenes in bioactive strains,Weishengwu Xuebao(Acta Microbiologica Sinica),47(2):228-234(朱鹏,郑立,林晶,邵健忠,严小军,2007,抗菌和细胞毒活性海洋细菌的筛选及其次生代谢基因证据,微生物学报,47(2):228-234)
Bingle L.E.,Simpson T.J.,and Lazarus C.M.,1999,Ketosynthase domain probes identify two subclasses of fungal polyketide synthase genes,Fungal Genet.Biol.,26(3):209-223
Chang Z.,Flatt P.,Gerwick W.H.,Nguyen V.A.,Willis C.L.,and Sherman D.H.,2002,The barbamide biosynthetic gene cluster:a novel marine cyanobacterial system of mixed polyketide synthase(PKS)-non-ribosomal peptide synthetase(NRPS)origin involving an unusual trichloroleucyl starter unit,Gene,296(1-2):235-247
Chen R.Q.,Liao L.,Zhang X.H.,and Chen B.,2014,Cloning and analysis of a halogenase gene of Streptomyces sp.604F from the Arctic Ocean,Weishengwu Xuebao(Acta Microbiologica Sinica),54(6):703-712(陈瑞勤,廖丽,张晓华,陈波,2014,北极海洋链霉菌604F的卤化酶基因克隆及特征,微生物学报,54(6):703-712)
Crawford J.M.,Korman T.P.,Labonte J.W.,Vagstad A.L.,and Hill E.A.,2009,Structural basis for biosynthetic programming of fungal aromatic polyketide cyclization,Nature,461(7267):1139-1143
Demain A.L.,and Sanchez S.,2009,Microbial drug discovery:80 years of progress,J.Antibiot.,62(1):5-16
Feng J.F.,Zhou R.C.,Guo X.T.,and Zhang Y.,2011,Polyketide and its application,Xiandai Nongye Keji(Modern Agricultural Sciences and Technology),(3):24-26(冯健飞,周日成,郭兴庭,张扬,2011,聚酮类化合物及其应用,现代农业科技,(3):24-26)
Harman G.E.,2006,Overview of mechanisms and uses of Trichoderma spp.,Phytopathology,96(2):190-194
Harman G.E.,2011,Trichoderma-not just for biocontrol anymore,Phytoparasitica,39(2):103-108
Harman G.E.,Howell C.R.,Viterbo A.,Chet I.,and Lorito M.,2004,Trichoderma species-opportunistic,avirulent plant symbionts,Nat.Rev.Microbiol.,2(1):43-56
Huang H.,and Hu H.f.,2016,Evolution and diversity of the gene clusters in microbial medicine biosyntheses,Shijie Linchuang Yaowu(World Clinical Drugs),37(2):129-134(黄鹤,胡海峰,2016,微生物药物合成基因簇的进化与结构多样性,世界临床药物,37(2):129-134)
Li J.W.H.,and Vederas J.C.,2009,Drug discovery and natural products:end of an era or an endless frontier,Science,325(5937):161-165
Li Q.,Li X.L.,Huang Y.J.,Chen C.,Huang W.L.,Xiong C.,and Zheng L.Y,2016,Biocontrol effect of Trichoderma virides-%cens strain 18-21 screened from fruiting bodies of Trametes gibbosa,Weishengwuxue Zazhi(Tournal of Microbiology),36(1):17-21(李强,李小林,黄羽佳,陈诚,黄文丽,熊川,郑林用,2016,偏肿栓菌内生木霉菌株18-21抑菌效果研究,微生物学杂志,36(1):17-21)
Lin M.J.,and Ko W.H.,2008,Occurrence of isolates of phytophthora colocasiae in Taiwan with homothallic behavior and its significance,Mycologia,100(5):727-734
Lu Y.,Lu Z.X.,He F.L.,Yuan G.Q.,Chen B.S.,and Liao Y.M.,2013,A research on mating types of Phytophthora colocasiae in south-eastern guangxi province,Jiyinzuxue Yu Yingyong Shengwuxue(Genomics and Applied Biology),32(3):285-290(陆叶,陆志翔,何芳练,袁高庆,陈保善,廖咏梅,2013,桂东南地区芋疫霉的交配型研究,基因组学与应用生物学,32(3):285-290)
Lucas-Elio P.,Hernandez P.,Sanchez-Amat A.,and Solano F.,2005,Purification and partial characterization of marinocine,a new broad-spectrum antibacterial protein produced by marinomonas mediterranea,Biochim.Biophys.Acta.,1721(1-3):193-203
Lv M.,Yang K.,Wang A.,Shi Y.N.,and Xin Z.H.,2016,Isolation and identification based on NRPS functional gene of a fungal epiphytic strain from grapes,Shipin Gongye Keji(Science and Technology of Food Industry),37(18):229-240(吕曼,杨柯,王傲,史雅凝,辛志宏,2016,基于NRPS基因筛选与鉴定葡萄附生真菌,食品工业科技,37(18):229-240)
Maansson M.,Vynne N.G.,Klitgaard A.,Nybo J.L.,Melchiorsen J.,Nguyen D.D.,Sanchez L.M.,Ziemert N.,Dorrestein P.C.,Andersen M.R.,and Gram L.,2016,An integrated metabolomic and genomic mining workflow to uncover the biosynthetic potential of bacteria,American Society for Microbiology,1(3):15-28
Mcdonald J.G.,Smith D.D.,Stiles A.R.,and Russell D.W.,2012,A comprehensive method for extraction and quantitative analysis of sterols and secosteroids from human plasma,J.Lipid Res.,53(7):1399-1409
Misiek M.,and Hoffmeister D.,2007,Fungal genetics,genomics,and secondary metabolites in pharmaceutical sciences,Planta Med.,73(2):103-115
Piel J.,2010,Biosynthesis of polyketides by trans-AT polyketide synthases,Natural Product Reports,27(7):996-1047
Ren D.,Zhang B.,Zhang X.P.,Teng Y.,Li X.L.,and Gu Y.F.,2014,Inheritance in endophytic fungi from Melia toosendan and diversity of genes of PKS and NRPS,Zhongcaoyao(Chinese Traditional and Herbal Drugs),45(10):1461-1467(任丹,张波,张小平,腾芸,李小林,辜运富,2014,川楝内生真菌的遗传及PKS、NRPS基因的多样性,中草药,45(10):1461-1467)
Schümann J.,and Hertweck C.,2006,Advances in cloning,functional analysis and heterologous expression of fungal polyketide synthase genes,J.Biotechnol.,124(4):690-703
Slightom J.L.,Metzger B.P.,Luu H.T.,and Elhammer A.P.,2009,Cloning and molecular characterization of the gene encoding the aureobasidin a biosynthesis complex in aureobasidium pullulans bp-1938,Gene,431(1-2):67-79
Tamura K.,Dudley J.,Nei M.,and Kumar S.,2007,MEGA4:molecular evolutionary genetics analysis(MEGA)software version 4.0,Mol.Biol.Evol.,24(8):1596-1599
Wang S.Y.,2007,Advances in the study of the mechanism and application of nonribosomal peptide synthetases,Weishengwu Xuebao(Acta Microbiologica Sinica),74(4):734-737(王世媛,2007,非核糖体肽合成酶(NRPSs)作用机理与应用的研究进展,微生物学报,74(4):734-737)
Wen Y.P.,2011,Analyses of the antibiotic%Paenimacrolidin and the siderophore paenibactin based on PKS and NRPS genes,Dissertation for Ph.D.,Zheiiang University,Supervisor:Zhao Y.H.,pp.5-7(文艳苹,2011,基于PKS,NRPS基因的抗生素Paenimacrolidin和嗜铁素Paenibactin研究,博士学位论文,浙江大学,导师:赵宇华,pp.5-7)
Xiao J.,Zhang G.T.,Zhu Y.G.,and Zhang C.S.,2012,Progress in biosynthesis of secondary metabolites from marine microorganisms,Zhongguo Kangshengsu Zazhi(Chinese Journal of Antibiotics),37(4):241-253(肖吉,张光涛,朱义广,张长生,2012,海洋微生物次级代谢产物生物合成的研究进展,中国抗生素杂志,37(4):241-253)
Zhang G.Z.,Yand H.T.,Zhang X.J.,Hu J.D.,Guo K.,Huang Y.J.,and Li J.S.,2014,A checklist of known species of trichoderma,Junwu Xuebao(Mycosystema),33(6):1210-1230(张广志,杨合同,张新建,扈进冬,郭凯,黄玉杰,李纪顺,2014,木霉现有种类名录,菌物学报,33(6):1210-1230)
Zhang J.N.,Peng J.,Liu T.X.,and Xin Z.H.,2016,Isolation and identification of endophyte and its secondary metabolites from%Salicornia bigelovii Torr.based on type i polyketide synthase(PKS I)gene,Shipin Kexue(Food Science),37(1):114-119(张俊楠,彭洁,刘天行,辛志宏,2016,基于PKSⅠ基因海芦笋内生真菌及其次级代谢产物筛选鉴定,食品科学,37(1):114-119)
Zhang L.,and Zhang J.Z.,2015,Isolation and purification of active compound from Trichoderma viridescens and its inhibitory activities against phytopathogens,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),48(5):882-888(张量,张敬泽,2015,渐绿木霉抑菌物质的分离纯化及其对植物病原菌的抑制作用,中国农业科学,48(5):882-888)
Zhong Z.M.,Lai X.P.,Huang S.,and Zhang G.F.,2018,Optimization of DNA extraction methods and molecular identification of endophytic Xylaria from Dendrobium,Junwu Xuebao(Mycosystema),37(1):73-78(钟志敏,赖小平,黄松张桂芳,2018,石斛内生炭角菌DNA提取方法优化及分子鉴定,菌物学报,37(1):73-78)
Zhou P.J.,Xu Y.,Ma Y.Q.,and Liu H.D.,1994,Amplification of16S r DNAs from halobacteria by means of PCR technique Weishengwu Xuebao(Acta Microbiologica Sinica),34(1):6-8(周培瑾,徐毅,马允卿,刘宏迪,1994,极端嗜盐菌16Sr DNA的PCR扩增,微生物学报,34(1):6-8)
Zhu M.Z.,Cui X.L.,Li M.G.,Li Y.Q.,Zhao J.Y.,Peng Q.,and Wen M.L.,2006,A new approach for microbial drugs discovery:from genes to novel compounds,Tianran Chanwu Yanjiu Yu Fazhan(Natural Product Research and Development)18(5):854-857(朱孟沼,崔晓龙,李铭刚,李一青,赵江源彭谦,文孟良,2006,发现微生物药物的新途径:从基因到新化合物,天然产物研究与开发,18(5):854-857)
Zhu P.,2008,Screening and studying modular structure of PKSand NRPS genes in marine sponge Hymeniacidon perleve associated bacteria,Dissertation for Ph.D.,School of Life Sciences,Zhejiang University,Supervisor:Shao J.Z.,pp.27-29(朱鹏,2008,珍珠膜海绵共附生微生物PKS与NRPS的筛选与模块结构研究,博士学位论文,浙江大学,导师:邵健忠,pp.27-29)
Zhu P.,Zheng L.,Li J.,Shao J.Z.,and Yan X.J.,2007,Screening and characterization of marine bacteria with antibacterial and cytotoxic activities,and existence of PKSⅠand NRPSgenes in bioactive strains,Weishengwu Xuebao(Acta Microbiologica Sinica),47(2):228-234(朱鹏,郑立,林晶,邵健忠,严小军,2007,抗菌和细胞毒活性海洋细菌的筛选及其次生代谢基因证据,微生物学报,47(2):228-234)
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