VeA调控曲霉发育致病的研究进展
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摘要
全局性调控因子Ve A仅在真菌中存在,具有保守性。Ve A参与细胞的多过程调控,包括发育分化、次生代谢、氧化胁迫应答、侵染宿主致毒致病等。文中总结曲霉Ve A的相关调控功能和作用机理研究,以利于防控真菌传播存活及致毒致病措施的设计,促进抗真菌作物的育种,减少作物被曲霉等真菌及毒素污染;还提出进一步开展ve A基因调控功能研究的方向和作为防控真菌污染靶位点的应用策略。
The global regulator Ve A has only been found in fungi and conserved in numerous fungal species. Ve A is involved in the regulation of diverse cellular processes, including development and differentiation, secondary metabolism, oxidative stress response as well as pathogenesis in numerous fungal species. The review presented here summarized the information on the current understanding of Ve A function and its mechanism of action in Aspergilli, with the aim of facilitating the construction of anti-infection crops and design of effective strategies for controlling fungal dissemination, survival, pathogenesis to decrease the Aspergilli contamination of agricultural commodities. Both the role of ve A gene that needs to be further characterized for future research and the potential target that may be used for controlling mycotoxin contamination were also discussed.
The global regulator Ve A has only been found in fungi and conserved in numerous fungal species. Ve A is involved in the regulation of diverse cellular processes, including development and differentiation, secondary metabolism, oxidative stress response as well as pathogenesis in numerous fungal species. The review presented here summarized the information on the current understanding of Ve A function and its mechanism of action in Aspergilli, with the aim of facilitating the construction of anti-infection crops and design of effective strategies for controlling fungal dissemination, survival, pathogenesis to decrease the Aspergilli contamination of agricultural commodities. Both the role of ve A gene that needs to be further characterized for future research and the potential target that may be used for controlling mycotoxin contamination were also discussed.
引文
[1]Liu ZR,Zhang GY,Yu JH.The function of the Afu4g13170gene in conidiation and gliotoxin production of pathogenic Aspergillus fumigatus[J].Microbiology China,2012,39(1):68-74(in Chinese)刘增然,张光一,Yu JH.致病菌烟曲霉新基因Afu4g13170生孢致毒相关性初步研究[J].微生物学通报,2012,39(1):68-74
[2]Baidya S,Duran R,Lohmar JM,et al.Ve A is associated with the response to oxidative stress in the aflatoxin producer Aspergillus flavus[J].Eukaryotic Cell,2014,13(8):1095-1103
[3]Amaike S,Keller NP.Distinct roles for Ve A and Lae A in development and pathogenesis of Aspergillus flavus[J].Eukaryotic Cell,2009,8(7):1051-1060
[4]Amare MG,Keller NP.Molecular mechanisms of Aspergillus flavus secondary metabolism and development[J].Fungal Genetics and Biology,2014,66:11-18
[5]Palmer JM,Theisen JM,Duran RM,et al.Secondary metabolism and development is mediated by Llm F control of Ve A subcellular localization in Aspergillus nidulans[J].PLo S Genetics,2013,9(1):e1003193
[6]Stinnett SM,Espeso EA,Cobe?o L,et al.Aspergillus nidulans Ve A subcellular localization is dependent on the importinαcarrier and on light[J].Molecular Microbiology,2007,63(1):242-255
[7]Dhingra S,Lind AL,Lin HC,et al.The fumagillin gene cluster,an example of hundreds of genes under Ve A control in Aspergillus fumigatus[J].PLo S One,2013,8(10):e77147
[8]Bayram?,Braus GH.Coordination of secondary metabolism and development in fungi:the velvet family of regulatory proteins[J].FEMS Microbiology Reviews,2012,36(1):1-24
[9]Myung K,Zitomer NC,Duvall M,et al.The conserved global regulator Ve A is necessary for symptom production and mycotoxin synthesis in maize seedlings by Fusarium verticillioides[J].Plant Pathology,2012,61(1):152-160
[10]Bayram?,Krappmann S,Ni M,et al.Vel B/Ve A/Lae A complex coordinates light signal with fungal development and secondary metabolism[J].Science,2008,320(5882):1504-1506
[11]Sarikaya-Bayram?,Bayram?,Valerius O,et al.Lae A control of velvet family regulatory proteins for light-dependent development and fungal cell-type specificity[J].PLo S Genetics,2010,6(12):e1001226
[12]Vienken K,Scherer M,Fischer R.The Zn(II)2Cys6 putative Aspergillus nidulans transcription factor repressor of sexual development inhibits sexual development under low-carbon conditions and in submersed culture[J].Genetics,2005,169(2):619-630
[13]Kong Q,Wang L,Liu ZR,et al.Gβ-like Cpc B plays a crucial role for growth and development of Aspergillus nidulans and Aspergillus fumigatus[J].PLo S One,2013,8(7):e70355
[14]Bayram?,Sarikaya-Bayram?,Ahmed YL,et al.The Aspergillus nidulans MAPK module An Ste11-Ste50-Ste7-Fus3controls development and secondary metabolism[J].PLo S Genetics,2012,8(7):e1002816
[15]Calvo AM,Cary JW.Association of fungal secondary metabolism and sclerotial biology[J].Frontiers in Microbiology,2015,6:62
[16]Park HS,Ni M,Jeong KC,et al.The role,interaction and regulation of the velvet regulator Vel B in Aspergillus nidulans[J].PLo S One,2012,7(9):e45935
[17]Purschwitz J,Müller S,Fischer R.Mapping the interaction sites of Aspergillus nidulans phytochrome Fph A with the global regulator Ve A and the white collar protein Lre B[J].Molecular Genetics and Genomics,2009,281(1):35-42
[18]Sarikaya-Bayram?,Bayram?,Feussner K,et al.Membrane-bound methyltransferase complex Vap A-Vip C-Vap B guides epigenetic control of fungal development[J].Developmental Cell,2014,29(4):406-420
[19]Roze LV,Chanda A,Laivenieks M,et al.Volatile profiling reveals intracellular metabolic changes in Aspergillus parasiticus:ve A regulates branched chain amino acid and ethanol metabolism[J].BMC Biochemistry,2010,11(1):33
[20]Duran RM,Cary JW,Calvo AM.Production of cyclopiazonic acid,aflatrem,and aflatoxin by Aspergillus flavus is regulated by ve A,a gene necessary for sclerotial formation[J].Applied Microbiology and Biotechnology,2007,73(5):1158-1168
[21]Dhingra S,Andes D,Calvo AM.Ve A regulates conidiation,gliotoxin production,and protease activity in the opportunistic human pathogen Aspergillus fumigatus[J].Eukaryotic Cell,2012,11(12):1531-1543
[22]Cary JW,O’Brian GR,Nielsen DM,et al.Elucidation of ve A-dependent genes associated with aflatoxin and sclerotial production in Aspergillus flavus by functional genomics[J].Applied Microbiology and Biotechnology,2007,76(5):1107-1118
[23]Calvo AM.The Ve A regulatory system and its role in morphological and chemical development in fungi[J].Fungal Genetics and Biology,2008,45(7):1053-1061
[24]Park HS,Bayram?,Braus GH,et al.Characterization of the velvet regulators in Aspergillus fumigatus[J].Molecular Microbiology,2012,86(4):937-953
[25]Krappmann S,Bayram?,Braus GH.Deletion and allelic exchange of the Aspergillus fumigatus ve A locus via a novel recyclable marker module[J].Eukaryotic Cell,2005,4(7):1298-1307
[26]Cary JW,Harris-Coward PY,Ehrlich KC,et al.Functional characterization of a Ve A-dependent polyketide synthase gene in Aspergillus flavus necessary for the synthesis of asparasone,a sclerotium-specific pigment[J].Fungal Genetics and Biology,2014,64:25-35
[27]Duran RM,Cary JW,Calvo AM.The role of Ve A in Aspergillus flavus infection of peanut,corn and cotton[J].The Open Mycology Journal,2009,3(1):27-36
[28]Marui J,Ohashi-Kunihiro S,Ando T,et al.Penicillin biosynthesis in Aspergillus oryzae and its overproduction by genetic engineering[J].Journal of Bioscience and Bioengineering,2010,110(1):8-11
[29]Dhingra S,Lind AL,Lin HC,et al.The fumagillin gene cluster,an example of hundreds of genes under Ve A control in Aspergillus fumigatus[J].PLo S One,2013,8(10):e77147
[30]Merhej J,Urban M,Dufresne M,et al.The velvet gene,Fg Ve1,affects fungal development and positively regulates trichothecene biosynthesis and pathogenicity in Fusarium graminearum[J].Molecular Plant Pathology,2012,13(4):363-374
[31]Wiemann P,Brown DW,Kleigrewe K,et al.Ff Vel1 and Ff Lae1,components of a velvet-like complex in Fusarium fujikuroi,affect differentiation,secondary metabolism and virulence[J].Molecular Microbiology,2010,77(4):972-994
[32]Wu DL,Oide S,Zhang N,et al.Ch Lae1 and Ch Vel1 regulate T-toxin production,virulence,oxidative stress response,and development of the maize pathogen Cochliobolus heterostrophus[J].PLo S Pathogens,2012,8(2):e1002542
[33]López-Berges MS,Hera C,Sulyok M,et al.The velvet complex governs mycotoxin production and virulence of Fusarium oxysporum on plant and mammalian hosts[J].Molecular Microbiology,2013,87(1):49-65
[34]Duran RM,Gregersen S,Smith TD,et al.The role of Aspergillus flavus ve A in the production of extracellular proteins during growth on starch substrates[J].Applied Microbiology and Biotechnology,2014,98(11):5081-5094
[35]López-Berges MS,Sch?fer K,Hera C,et al.Combinatorial function of velvet and Are A in transcriptional regulation of nitrate utilization and secondary metabolism[J].Fungal Genetics and Biology,2014,62:78-84
[36]Lan N,Zhang HX,Hu CC,et al.Coordinated and distinct functions of velvet proteins in Fusarium verticillioides[J].Eukaryotic Cell,2014,13(7):909-918
[37]Hong SY,Roze LV,Wee J,et al.Evidence that a transcription factor regulatory network coordinates oxidative stress response and secondary metabolism in aspergilli[J].Microbiology Open,2013,2(1):144-160
[38]Roze LV,Chanda A,Wee J,et al.Stress-related transcription factor Atf B integrates secondary metabolism with oxidative stress response in Aspergilli[J].The Journal of Biological Chemistry,2011,286(40):35137-35148
[39]Duran R,Cary JW,Calvo AM.Role of the osmotic stress regulatory pathway in morphogenesis and secondary metabolism in filamentous fungi[J].Toxins,2010,2(4):367-381
[40]Brown SH,Scott JB,Bhaheetharan J,et al.Oxygenase coordination is required for morphological transition and the host-fungus interaction of Aspergillus flavus[J].Molecular Plant-Microbe Interactions,2009,22(7):882-894
[41]Scarpari M,Punelli M,Scala V,et al.Lipids in Aspergillus flavus-maize interaction[J].Frontiers in Microbiology,2014,5:74
[42]Dyer PS,O’Gorman CM.Sexual development and cryptic sexuality in fungi:insights from Aspergillus species[J].FEMS Microbiology Reviews,2012,36(1):165-192
[43]Tsitsigiannis DI,Kowieski TM,Zarnowski R,et al.Three putative oxylipin biosynthetic genes integrate sexual and asexual development in Aspergillus nidulans[J].Microbiology,2005,151(6):1809-1821
[2]Baidya S,Duran R,Lohmar JM,et al.Ve A is associated with the response to oxidative stress in the aflatoxin producer Aspergillus flavus[J].Eukaryotic Cell,2014,13(8):1095-1103
[3]Amaike S,Keller NP.Distinct roles for Ve A and Lae A in development and pathogenesis of Aspergillus flavus[J].Eukaryotic Cell,2009,8(7):1051-1060
[4]Amare MG,Keller NP.Molecular mechanisms of Aspergillus flavus secondary metabolism and development[J].Fungal Genetics and Biology,2014,66:11-18
[5]Palmer JM,Theisen JM,Duran RM,et al.Secondary metabolism and development is mediated by Llm F control of Ve A subcellular localization in Aspergillus nidulans[J].PLo S Genetics,2013,9(1):e1003193
[6]Stinnett SM,Espeso EA,Cobe?o L,et al.Aspergillus nidulans Ve A subcellular localization is dependent on the importinαcarrier and on light[J].Molecular Microbiology,2007,63(1):242-255
[7]Dhingra S,Lind AL,Lin HC,et al.The fumagillin gene cluster,an example of hundreds of genes under Ve A control in Aspergillus fumigatus[J].PLo S One,2013,8(10):e77147
[8]Bayram?,Braus GH.Coordination of secondary metabolism and development in fungi:the velvet family of regulatory proteins[J].FEMS Microbiology Reviews,2012,36(1):1-24
[9]Myung K,Zitomer NC,Duvall M,et al.The conserved global regulator Ve A is necessary for symptom production and mycotoxin synthesis in maize seedlings by Fusarium verticillioides[J].Plant Pathology,2012,61(1):152-160
[10]Bayram?,Krappmann S,Ni M,et al.Vel B/Ve A/Lae A complex coordinates light signal with fungal development and secondary metabolism[J].Science,2008,320(5882):1504-1506
[11]Sarikaya-Bayram?,Bayram?,Valerius O,et al.Lae A control of velvet family regulatory proteins for light-dependent development and fungal cell-type specificity[J].PLo S Genetics,2010,6(12):e1001226
[12]Vienken K,Scherer M,Fischer R.The Zn(II)2Cys6 putative Aspergillus nidulans transcription factor repressor of sexual development inhibits sexual development under low-carbon conditions and in submersed culture[J].Genetics,2005,169(2):619-630
[13]Kong Q,Wang L,Liu ZR,et al.Gβ-like Cpc B plays a crucial role for growth and development of Aspergillus nidulans and Aspergillus fumigatus[J].PLo S One,2013,8(7):e70355
[14]Bayram?,Sarikaya-Bayram?,Ahmed YL,et al.The Aspergillus nidulans MAPK module An Ste11-Ste50-Ste7-Fus3controls development and secondary metabolism[J].PLo S Genetics,2012,8(7):e1002816
[15]Calvo AM,Cary JW.Association of fungal secondary metabolism and sclerotial biology[J].Frontiers in Microbiology,2015,6:62
[16]Park HS,Ni M,Jeong KC,et al.The role,interaction and regulation of the velvet regulator Vel B in Aspergillus nidulans[J].PLo S One,2012,7(9):e45935
[17]Purschwitz J,Müller S,Fischer R.Mapping the interaction sites of Aspergillus nidulans phytochrome Fph A with the global regulator Ve A and the white collar protein Lre B[J].Molecular Genetics and Genomics,2009,281(1):35-42
[18]Sarikaya-Bayram?,Bayram?,Feussner K,et al.Membrane-bound methyltransferase complex Vap A-Vip C-Vap B guides epigenetic control of fungal development[J].Developmental Cell,2014,29(4):406-420
[19]Roze LV,Chanda A,Laivenieks M,et al.Volatile profiling reveals intracellular metabolic changes in Aspergillus parasiticus:ve A regulates branched chain amino acid and ethanol metabolism[J].BMC Biochemistry,2010,11(1):33
[20]Duran RM,Cary JW,Calvo AM.Production of cyclopiazonic acid,aflatrem,and aflatoxin by Aspergillus flavus is regulated by ve A,a gene necessary for sclerotial formation[J].Applied Microbiology and Biotechnology,2007,73(5):1158-1168
[21]Dhingra S,Andes D,Calvo AM.Ve A regulates conidiation,gliotoxin production,and protease activity in the opportunistic human pathogen Aspergillus fumigatus[J].Eukaryotic Cell,2012,11(12):1531-1543
[22]Cary JW,O’Brian GR,Nielsen DM,et al.Elucidation of ve A-dependent genes associated with aflatoxin and sclerotial production in Aspergillus flavus by functional genomics[J].Applied Microbiology and Biotechnology,2007,76(5):1107-1118
[23]Calvo AM.The Ve A regulatory system and its role in morphological and chemical development in fungi[J].Fungal Genetics and Biology,2008,45(7):1053-1061
[24]Park HS,Bayram?,Braus GH,et al.Characterization of the velvet regulators in Aspergillus fumigatus[J].Molecular Microbiology,2012,86(4):937-953
[25]Krappmann S,Bayram?,Braus GH.Deletion and allelic exchange of the Aspergillus fumigatus ve A locus via a novel recyclable marker module[J].Eukaryotic Cell,2005,4(7):1298-1307
[26]Cary JW,Harris-Coward PY,Ehrlich KC,et al.Functional characterization of a Ve A-dependent polyketide synthase gene in Aspergillus flavus necessary for the synthesis of asparasone,a sclerotium-specific pigment[J].Fungal Genetics and Biology,2014,64:25-35
[27]Duran RM,Cary JW,Calvo AM.The role of Ve A in Aspergillus flavus infection of peanut,corn and cotton[J].The Open Mycology Journal,2009,3(1):27-36
[28]Marui J,Ohashi-Kunihiro S,Ando T,et al.Penicillin biosynthesis in Aspergillus oryzae and its overproduction by genetic engineering[J].Journal of Bioscience and Bioengineering,2010,110(1):8-11
[29]Dhingra S,Lind AL,Lin HC,et al.The fumagillin gene cluster,an example of hundreds of genes under Ve A control in Aspergillus fumigatus[J].PLo S One,2013,8(10):e77147
[30]Merhej J,Urban M,Dufresne M,et al.The velvet gene,Fg Ve1,affects fungal development and positively regulates trichothecene biosynthesis and pathogenicity in Fusarium graminearum[J].Molecular Plant Pathology,2012,13(4):363-374
[31]Wiemann P,Brown DW,Kleigrewe K,et al.Ff Vel1 and Ff Lae1,components of a velvet-like complex in Fusarium fujikuroi,affect differentiation,secondary metabolism and virulence[J].Molecular Microbiology,2010,77(4):972-994
[32]Wu DL,Oide S,Zhang N,et al.Ch Lae1 and Ch Vel1 regulate T-toxin production,virulence,oxidative stress response,and development of the maize pathogen Cochliobolus heterostrophus[J].PLo S Pathogens,2012,8(2):e1002542
[33]López-Berges MS,Hera C,Sulyok M,et al.The velvet complex governs mycotoxin production and virulence of Fusarium oxysporum on plant and mammalian hosts[J].Molecular Microbiology,2013,87(1):49-65
[34]Duran RM,Gregersen S,Smith TD,et al.The role of Aspergillus flavus ve A in the production of extracellular proteins during growth on starch substrates[J].Applied Microbiology and Biotechnology,2014,98(11):5081-5094
[35]López-Berges MS,Sch?fer K,Hera C,et al.Combinatorial function of velvet and Are A in transcriptional regulation of nitrate utilization and secondary metabolism[J].Fungal Genetics and Biology,2014,62:78-84
[36]Lan N,Zhang HX,Hu CC,et al.Coordinated and distinct functions of velvet proteins in Fusarium verticillioides[J].Eukaryotic Cell,2014,13(7):909-918
[37]Hong SY,Roze LV,Wee J,et al.Evidence that a transcription factor regulatory network coordinates oxidative stress response and secondary metabolism in aspergilli[J].Microbiology Open,2013,2(1):144-160
[38]Roze LV,Chanda A,Wee J,et al.Stress-related transcription factor Atf B integrates secondary metabolism with oxidative stress response in Aspergilli[J].The Journal of Biological Chemistry,2011,286(40):35137-35148
[39]Duran R,Cary JW,Calvo AM.Role of the osmotic stress regulatory pathway in morphogenesis and secondary metabolism in filamentous fungi[J].Toxins,2010,2(4):367-381
[40]Brown SH,Scott JB,Bhaheetharan J,et al.Oxygenase coordination is required for morphological transition and the host-fungus interaction of Aspergillus flavus[J].Molecular Plant-Microbe Interactions,2009,22(7):882-894
[41]Scarpari M,Punelli M,Scala V,et al.Lipids in Aspergillus flavus-maize interaction[J].Frontiers in Microbiology,2014,5:74
[42]Dyer PS,O’Gorman CM.Sexual development and cryptic sexuality in fungi:insights from Aspergillus species[J].FEMS Microbiology Reviews,2012,36(1):165-192
[43]Tsitsigiannis DI,Kowieski TM,Zarnowski R,et al.Three putative oxylipin biosynthetic genes integrate sexual and asexual development in Aspergillus nidulans[J].Microbiology,2005,151(6):1809-1821