除草微生物Ophiobolin A毒素合成相关基因研究
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摘要
蛇孢菌素化合物具有除草、抑菌、杀线虫、抗肿瘤细胞等活性，在农业和医药领域都有很好的应用前景。但是由于蛇孢菌素化合物是真菌体内产生的一类次生代谢产物，因此产量极低，限制其商业化生产。因此采用各种手段提高蛇孢菌素的产量，使其达到工业化生产的水平是促进其广泛应用的重要研究方向。
为解决蛇孢菌素产量低的问题，从两个方面开展研究：一是对产蛇孢菌素真菌蟋蟀草平脐蠕孢菌（Biopolaris eleusines）转录组进行测序，获得高质量的EST序列，以此为线索，采用RACE技术克隆蟋蟀草平脐蠕孢菌中蛇孢菌素合成途径关键酶基因，为今后利用基因工程手段提高真菌中蛇孢菌素产量奠定基础；二是初步研究了单一化合物茉莉酸甲酯对蟋蟀草平脐蠕孢菌产蛇孢菌素合成相关基因表达的影响。本研究取得如下结果：
首次对蟋蟀草平脐蠕孢菌转录组进行测序，研究其基因表达谱，挖掘其功能基因。共获得26,555,560条高质量EST，序列平均长度为559bp。所得序列经聚类合并拼接后得到32,100条高质量的单基因簇（Unigene），数据库中的序列同源性比较表明，其中97.9%（31432条）序列与其它生物的已知基因具有不同程度的同源性。通过BLAST与KEGG Pathway分析获得了可能与萜类化合物合成相关的60条Unigene，分别编码合成途径中17个酶。分析还发现转录因子序列76条。这些基因的发现为二倍半萜类化合物蛇孢菌素生物合成研究奠定基础，同时也为蟋蟀草平脐蠕孢菌转录组研究提供了基础数据。
克隆了蟋蟀草平脐蠕孢菌中蛇孢菌素合成途径上第一个限速酶3-羟基-3-甲基戊二酰辅酶A还原酶基因（BeHMGR, GenBank accession No. JQ780844）的全长cDNA序列并对其进行生物信息学分析。该基因全长3906bp，含有一个3474bp的开放阅读框，编码一个具有1157个氨基酸残基的蛋白，BLASTP分析显示，该基因编码的氨基酸与其它真菌的HMGR有较高相似性。其中与小麦黄斑叶枯病原真菌(Pyrenophora tritici-repentis)有93%相似性和86％一致性。荧光定量PCR分析发现，BeHMGR基因的表达受茉莉酸甲酯诱导。
克隆了蟋蟀草平脐蠕孢菌中蛇孢菌素合成途径上重要前体物异戊烯基焦磷酸异构酶基因（BeIPPI, GenBank accession No. JQ780840）的全长cDNA序列，该基因全长1059bp，含有一个672bp的开放阅读框，编码一个具有223个氨基酸残基的蛋白，BLASTP分析显示，该基因编码的氨基酸与其它真菌的IPPI保守结构域以及催化活性位点的氨基酸残基都完全一致，但是在C端比其它蛋白缺失40个左右的氨基酸残基，该基因是否是一个有功能的IPPI基因还有待于下一步基因功能验证。
克隆了蟋蟀草平脐蠕孢菌中蛇孢菌素合成途径上关键酶法尼基焦磷酸合成酶基因（BeFPPS, GenBank accession No. JQ780841）的全长cDNA序列，该基因全长1186bp，含有一个1044bp的开放阅读框，编码一个具有347个氨基酸残基的蛋白。BLASTP分析显示，BeFPPS基因编码的氨基酸与其它真菌的FPPS有较高相似性，其中亲缘关系最近的是大麦网斑病菌（Pyrenophora teres f. teres），氨基酸序列一致性达89%。
克隆了蟋蟀草平脐蠕孢菌中蛇孢菌素合成途径上关键酶香叶基香叶基焦磷酸合成酶基因（BeGGPPS, GenBank accession No. JQ780842）的全长cDNA序列，该基因全长1666bp，含有一个1305bp的开放阅读框，编码一个具有434个氨基酸残基的蛋白。蛋白质序列多重比对结果表明，BeGGPPS与其它真菌来源的GGPPS具有较高同源性，BeGGPPS含有典型的异戊烯基转移酶5个氨基酸保守区（I－V），BeGGPPS系统发育树分析表明，BeGGPPS与小麦黄斑叶枯病菌（P. tritici-repentis）中GGPPS蛋白有更近的亲缘关系。
初步研究了茉莉酸甲酯对蟋蟀草平脐蠕孢菌蛇孢菌素A相关基因表达的影响。研究发现茉莉酸甲酯处理后能诱导蛇孢菌素A合成途径上BeHMGR, BeIPPI, BeFPPS和BeGGPPS基因的高表达。
本研究测定了蟋蟀草平脐蠕孢菌转录组序列，首次从该菌中分离克隆了4个蛇孢菌素生物合成关键酶基因，初步研究了茉莉酸甲酯对蛇孢菌素合成基因表达水平的影响。这些研究结果为深入阐明蛇孢菌素生物合成的分子机制奠定了重要基础，也为蛇孢菌素的代谢工程提供了更多可能的调控靶点，具有重要的理论意义和应用价值。
Ophiobolins are sesterterpenoid-type phytotoxins produced by fungi. They are active on a broadspectrum of organisms including plants, fungi, bacteria, nematodes and tumor cells, so they maybe important candidate for development of new crop-protection and pharmaceutical product.However, due to the low content of bioactive components in ophiobolin-producing fungi, thecommercial applications of ophiobolins are restricted. Hence, a number of studies have beenconducted in attempts to improve the yield of ophiobolins.
To significantly increase yields of ophiobolin A from an ophiobolin-producing fungus,Biopolaris eleusines (Be), this study focused on two aspects. The first was that transcriptomesequencing was used to produce a substantial expressed sequence tags (EST) dataset from Be.Based on the analysis of EST, molecular cloning by RACE technology and analysis of related keygenes involved in ophiobolin A biosynthesis pathway were conducted. This would provide someimportant candidate genes for Be genetic engineering to improve the yield of ophiobolin A in thefuture. The second was the stimulatory effect of methyl jasmonate (MeJA) on the expression levelof BeHMGR, BeIPPI, BeFPPS, BeGGPPS involved in biosynthesis of ophiobolin A in Be. Themain results from the study were as follows:
To investigate the profile of gene expression in Be and elucidate its functional gene,high-throughput transcriptome sequencing was firstly used to produce a substantial EST from Be.A total of26,555,560high quality EST with an average read length of559bp were generated.These EST were assembled into32,100unigenes.97.9%of these unigenes (31432) wereannotated and showed different levels of homology with other organisms using BLAST searches(E-value≤1e-5) against the SwissProt, KEGG, Nr, COG and GO databases. Sixty unigenes(encoding17enzymes) were found to be involved in terpenoid biosynthesis. Seventy-six unigeneswere also analyzed be similar to that of transcription factor genes. Data presented in this study notonly provided a number of resources in the profile of gene expression of Be, but also laied thefoundation for cloning and analyzing the encoding gene and regulation gene involved inophiobolins biosynthesis.
A full-length cDNA encoding3-hydroxy-3-methylglutaryl-CoA reductase gene (designated asBeHMGR, GenBank accession No. JQ780844) was cloned from the ophiobolin A-producingfungus Be by rapid amplification of cDNA ends (RACE) technique and was subject to detailedbioinformatic analysis. The full-length cDNA of BeHMGR was3906bp containing a3474bpORF encoding1157amino acids. BLASTP analysis revealed that the deduced BeHMGR hadextensive homology with HMGR of other fungi. Phylogenetic tree analysis indicated thatBeHMGR belongs to the fungi HMGR super-family and had the closest relationship with HMGRfrom Pyrenophora tritici-repentis (86%identity and93%similarity). The expression of BeHMGRwas up-regulated by methyl jasmonate (MeJA), suggesting that it might be elicitor-responsive.
A full-length cDNA encoding isopentenyl diphosphate isomerase gene (designated as BeIPPI, GenBank accession No. JQ780840) was isolated and cloned from the ophiobolin A-producingfungus Be by RACE technique. The bioinformatic analysis found that the full-length cDNA ofBeIPPI was1059bp containing a672bp ORF encoding a polypeptide of223amino acids.Analysis of BeIPPI genomic DNA revealed that it contained two exons and one intron. BLASTPanalysis revealed that the deduced BeIPPI contained all conserved domains and completelyidentical amino acids of catalytic active site compared to other fungi IPPI, but about40aminoacids of BeIPPI was abridged at C-terminus. Functional analysis of BeIPPI will be required toverify that if it is a functional protein.
A full-length cDNA encoding farnesyl diphosphate synthase gene (designated as BeFPPS,GenBank accession No. JQ780841) was obtained from the ophiobolin A-producing fungus Be byRACE and was subject to detailed bioinformatic analysis. The full-length cDNA of BeFPPS was1186bp containing a1044bp ORF encoding a polypeptide of347amino acids. Analysis ofBeFPPS genomic DNA revealed that it contained two exons and one intron. BLASTP analysisrevealed that the deduced BeFPPS had extensive homology with other fungi FPPS. Phylogenetictree analysis indicated that BeFPPS belongs to the fungi GGPPS super-family and had the closestrelationship with FPPS from P. teres f. teres（89%identity）.
A full-length cDNA encoding geranylgeranlyl diphosphate synthase gene (designated asBeGGPPS, GenBank accession No. JQ780842) was cloned from Be by RACE. The bioinformaticanalysis indicated that the full-length cDNA of BeGGPPS was1666bp containing a1305bp ORFencoding a polypeptide of434amino acids. Analysis of BeGGPPS genomic DNA revealed that itcontained three exons and two introns. BLASTP analysis revealed that the deduced BeGGPPS hadextensive homology with other fungi GGPPS contained all five conserved domains and functionalaspartate-rich motifs of the prenyltransferases. Phylogenetic tree analysis indicated that BeGGPPSbelongs to the fungi GGPPS super-family and had the closest relationship with GGPPS from P.tritici-repentis.
The stimulatory effect of MeJA on the transcriptional level of ophiobolin A gene in Be was alsopreliminarily studied. QRT-PCR analysis indicated that gene expression of BeHMGR, BeIPPI,BeFPPS and BeGGPPS, four key enzymes in ophiobolin A biosynthesis pathway, wasdramatically raised by MeJA treatment.
Transcriptome sequences of Be was firstly determined. BeHMGR, BeIPPI, BeFPPS andBeGGPPS of four key enzymes involved in ophiobolin A biosynthesis pathway were isolated fromophiobolin A-producing fungus stratin Be for the first time. Furthermore, the induction of MeJAon expression level of BeHMGR, BeIPPI, BeFPPS and BeGGPPS was preliminarily studied.These information might be helpful not only for theoretical research of further defining themechanism of ophiobolin A biosynthesis, but also supply more clues of target enzymes tomodulate the production of ophiobolin A for practical application.
为解决蛇孢菌素产量低的问题，从两个方面开展研究：一是对产蛇孢菌素真菌蟋蟀草平脐蠕孢菌（Biopolaris eleusines）转录组进行测序，获得高质量的EST序列，以此为线索，采用RACE技术克隆蟋蟀草平脐蠕孢菌中蛇孢菌素合成途径关键酶基因，为今后利用基因工程手段提高真菌中蛇孢菌素产量奠定基础；二是初步研究了单一化合物茉莉酸甲酯对蟋蟀草平脐蠕孢菌产蛇孢菌素合成相关基因表达的影响。本研究取得如下结果：
首次对蟋蟀草平脐蠕孢菌转录组进行测序，研究其基因表达谱，挖掘其功能基因。共获得26,555,560条高质量EST，序列平均长度为559bp。所得序列经聚类合并拼接后得到32,100条高质量的单基因簇（Unigene），数据库中的序列同源性比较表明，其中97.9%（31432条）序列与其它生物的已知基因具有不同程度的同源性。通过BLAST与KEGG Pathway分析获得了可能与萜类化合物合成相关的60条Unigene，分别编码合成途径中17个酶。分析还发现转录因子序列76条。这些基因的发现为二倍半萜类化合物蛇孢菌素生物合成研究奠定基础，同时也为蟋蟀草平脐蠕孢菌转录组研究提供了基础数据。
克隆了蟋蟀草平脐蠕孢菌中蛇孢菌素合成途径上第一个限速酶3-羟基-3-甲基戊二酰辅酶A还原酶基因（BeHMGR, GenBank accession No. JQ780844）的全长cDNA序列并对其进行生物信息学分析。该基因全长3906bp，含有一个3474bp的开放阅读框，编码一个具有1157个氨基酸残基的蛋白，BLASTP分析显示，该基因编码的氨基酸与其它真菌的HMGR有较高相似性。其中与小麦黄斑叶枯病原真菌(Pyrenophora tritici-repentis)有93%相似性和86％一致性。荧光定量PCR分析发现，BeHMGR基因的表达受茉莉酸甲酯诱导。
克隆了蟋蟀草平脐蠕孢菌中蛇孢菌素合成途径上重要前体物异戊烯基焦磷酸异构酶基因（BeIPPI, GenBank accession No. JQ780840）的全长cDNA序列，该基因全长1059bp，含有一个672bp的开放阅读框，编码一个具有223个氨基酸残基的蛋白，BLASTP分析显示，该基因编码的氨基酸与其它真菌的IPPI保守结构域以及催化活性位点的氨基酸残基都完全一致，但是在C端比其它蛋白缺失40个左右的氨基酸残基，该基因是否是一个有功能的IPPI基因还有待于下一步基因功能验证。
克隆了蟋蟀草平脐蠕孢菌中蛇孢菌素合成途径上关键酶法尼基焦磷酸合成酶基因（BeFPPS, GenBank accession No. JQ780841）的全长cDNA序列，该基因全长1186bp，含有一个1044bp的开放阅读框，编码一个具有347个氨基酸残基的蛋白。BLASTP分析显示，BeFPPS基因编码的氨基酸与其它真菌的FPPS有较高相似性，其中亲缘关系最近的是大麦网斑病菌（Pyrenophora teres f. teres），氨基酸序列一致性达89%。
克隆了蟋蟀草平脐蠕孢菌中蛇孢菌素合成途径上关键酶香叶基香叶基焦磷酸合成酶基因（BeGGPPS, GenBank accession No. JQ780842）的全长cDNA序列，该基因全长1666bp，含有一个1305bp的开放阅读框，编码一个具有434个氨基酸残基的蛋白。蛋白质序列多重比对结果表明，BeGGPPS与其它真菌来源的GGPPS具有较高同源性，BeGGPPS含有典型的异戊烯基转移酶5个氨基酸保守区（I－V），BeGGPPS系统发育树分析表明，BeGGPPS与小麦黄斑叶枯病菌（P. tritici-repentis）中GGPPS蛋白有更近的亲缘关系。
初步研究了茉莉酸甲酯对蟋蟀草平脐蠕孢菌蛇孢菌素A相关基因表达的影响。研究发现茉莉酸甲酯处理后能诱导蛇孢菌素A合成途径上BeHMGR, BeIPPI, BeFPPS和BeGGPPS基因的高表达。
本研究测定了蟋蟀草平脐蠕孢菌转录组序列，首次从该菌中分离克隆了4个蛇孢菌素生物合成关键酶基因，初步研究了茉莉酸甲酯对蛇孢菌素合成基因表达水平的影响。这些研究结果为深入阐明蛇孢菌素生物合成的分子机制奠定了重要基础，也为蛇孢菌素的代谢工程提供了更多可能的调控靶点，具有重要的理论意义和应用价值。
Ophiobolins are sesterterpenoid-type phytotoxins produced by fungi. They are active on a broadspectrum of organisms including plants, fungi, bacteria, nematodes and tumor cells, so they maybe important candidate for development of new crop-protection and pharmaceutical product.However, due to the low content of bioactive components in ophiobolin-producing fungi, thecommercial applications of ophiobolins are restricted. Hence, a number of studies have beenconducted in attempts to improve the yield of ophiobolins.
To significantly increase yields of ophiobolin A from an ophiobolin-producing fungus,Biopolaris eleusines (Be), this study focused on two aspects. The first was that transcriptomesequencing was used to produce a substantial expressed sequence tags (EST) dataset from Be.Based on the analysis of EST, molecular cloning by RACE technology and analysis of related keygenes involved in ophiobolin A biosynthesis pathway were conducted. This would provide someimportant candidate genes for Be genetic engineering to improve the yield of ophiobolin A in thefuture. The second was the stimulatory effect of methyl jasmonate (MeJA) on the expression levelof BeHMGR, BeIPPI, BeFPPS, BeGGPPS involved in biosynthesis of ophiobolin A in Be. Themain results from the study were as follows:
To investigate the profile of gene expression in Be and elucidate its functional gene,high-throughput transcriptome sequencing was firstly used to produce a substantial EST from Be.A total of26,555,560high quality EST with an average read length of559bp were generated.These EST were assembled into32,100unigenes.97.9%of these unigenes (31432) wereannotated and showed different levels of homology with other organisms using BLAST searches(E-value≤1e-5) against the SwissProt, KEGG, Nr, COG and GO databases. Sixty unigenes(encoding17enzymes) were found to be involved in terpenoid biosynthesis. Seventy-six unigeneswere also analyzed be similar to that of transcription factor genes. Data presented in this study notonly provided a number of resources in the profile of gene expression of Be, but also laied thefoundation for cloning and analyzing the encoding gene and regulation gene involved inophiobolins biosynthesis.
A full-length cDNA encoding3-hydroxy-3-methylglutaryl-CoA reductase gene (designated asBeHMGR, GenBank accession No. JQ780844) was cloned from the ophiobolin A-producingfungus Be by rapid amplification of cDNA ends (RACE) technique and was subject to detailedbioinformatic analysis. The full-length cDNA of BeHMGR was3906bp containing a3474bpORF encoding1157amino acids. BLASTP analysis revealed that the deduced BeHMGR hadextensive homology with HMGR of other fungi. Phylogenetic tree analysis indicated thatBeHMGR belongs to the fungi HMGR super-family and had the closest relationship with HMGRfrom Pyrenophora tritici-repentis (86%identity and93%similarity). The expression of BeHMGRwas up-regulated by methyl jasmonate (MeJA), suggesting that it might be elicitor-responsive.
A full-length cDNA encoding isopentenyl diphosphate isomerase gene (designated as BeIPPI, GenBank accession No. JQ780840) was isolated and cloned from the ophiobolin A-producingfungus Be by RACE technique. The bioinformatic analysis found that the full-length cDNA ofBeIPPI was1059bp containing a672bp ORF encoding a polypeptide of223amino acids.Analysis of BeIPPI genomic DNA revealed that it contained two exons and one intron. BLASTPanalysis revealed that the deduced BeIPPI contained all conserved domains and completelyidentical amino acids of catalytic active site compared to other fungi IPPI, but about40aminoacids of BeIPPI was abridged at C-terminus. Functional analysis of BeIPPI will be required toverify that if it is a functional protein.
A full-length cDNA encoding farnesyl diphosphate synthase gene (designated as BeFPPS,GenBank accession No. JQ780841) was obtained from the ophiobolin A-producing fungus Be byRACE and was subject to detailed bioinformatic analysis. The full-length cDNA of BeFPPS was1186bp containing a1044bp ORF encoding a polypeptide of347amino acids. Analysis ofBeFPPS genomic DNA revealed that it contained two exons and one intron. BLASTP analysisrevealed that the deduced BeFPPS had extensive homology with other fungi FPPS. Phylogenetictree analysis indicated that BeFPPS belongs to the fungi GGPPS super-family and had the closestrelationship with FPPS from P. teres f. teres（89%identity）.
A full-length cDNA encoding geranylgeranlyl diphosphate synthase gene (designated asBeGGPPS, GenBank accession No. JQ780842) was cloned from Be by RACE. The bioinformaticanalysis indicated that the full-length cDNA of BeGGPPS was1666bp containing a1305bp ORFencoding a polypeptide of434amino acids. Analysis of BeGGPPS genomic DNA revealed that itcontained three exons and two introns. BLASTP analysis revealed that the deduced BeGGPPS hadextensive homology with other fungi GGPPS contained all five conserved domains and functionalaspartate-rich motifs of the prenyltransferases. Phylogenetic tree analysis indicated that BeGGPPSbelongs to the fungi GGPPS super-family and had the closest relationship with GGPPS from P.tritici-repentis.
The stimulatory effect of MeJA on the transcriptional level of ophiobolin A gene in Be was alsopreliminarily studied. QRT-PCR analysis indicated that gene expression of BeHMGR, BeIPPI,BeFPPS and BeGGPPS, four key enzymes in ophiobolin A biosynthesis pathway, wasdramatically raised by MeJA treatment.
Transcriptome sequences of Be was firstly determined. BeHMGR, BeIPPI, BeFPPS andBeGGPPS of four key enzymes involved in ophiobolin A biosynthesis pathway were isolated fromophiobolin A-producing fungus stratin Be for the first time. Furthermore, the induction of MeJAon expression level of BeHMGR, BeIPPI, BeFPPS and BeGGPPS was preliminarily studied.These information might be helpful not only for theoretical research of further defining themechanism of ophiobolin A biosynthesis, but also supply more clues of target enzymes tomodulate the production of ophiobolin A for practical application.
引文
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