盾壳霉产孢相关基因的克隆及腺苷脱氨酶(CMADA)基因的初步研究
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摘要
为了发挥盾壳霉的生防作用,有必要深入研究盾壳霉的分子生长发育、生理代谢途径及产孢调控。本研究以盾壳霉ZS-1菌株为出发菌株,在前期基础上进一步优化农杆菌介导的转化条件,将转化效率提高至50转化子/培养皿,既保证了转化效率,又可避免两个单孢萌发的菌丝长到一起,保证了转化子的纯度。通过这种方法构建了含5000个转化子的盾壳霉T-DNA标记插入突变体库。以期获得大量突变体克隆相关基因,研究基因功能。
随后我们自盾壳霉的插入突变体库中共筛选到了23株表型异常突变体,其中8株突变体在PDA平皿上完全不能产生分生孢子,10株产孢明显减少,6株产生色素异常,3株生长迅速,3株生长缓慢,2株菌丝塌陷。其中产孢缺陷型突变体在菌丝生长速度、产抗生物质、寄生菌核能力和菌落形态等方面存在差异显著性。利用反向PCR、cDNA文库等克隆基因的技术平台,克隆了5株突变体的相关基因,包括ZS-1TN29、ZS-1TN289、ZS-1TN5012、ZS-1TN5498和ZS-1TN6222,利用NCBI保守结构域数据库比较分析,编码蛋白分别推定为CMARGJ、CMGAL、CMHMGR、CMPurD、CMMAPKKK。
对突变体ZS-1TN250进行了详细研究,发现该突变体生长速度迅速,但在PDB中的生物产量比野生菌株低；产孢明显减少,后期有少量孢子产生,显微观察发现大部分孢子没有黑色素化；该突变体能寄生核盘菌,但在菌核上不产生分生孢子,致腐菌核能力显著下降；可以正常产生抗生物质。采用反向PCR技术对ZS-1TN250中T-DNA标记插入位点的侧翼基因组DNA进行克隆,获得了大小为902bp的DNA片段,结合盾壳霉基因组序列分析,利用DNA MANagement软件进行全长序列拼接,得到该基因全长1522bp,包含4个外显子和3个内含子,cDNA全长为996 bp,推定编码含332个氨基酸的蛋白,T-DNA插入到第二个外显子上191nt的位置。通过NCBI BLASTP数据库,对推定编码的氨基酸序列进行分析,与adenosine/AMP deaminase family protein (Pyrenophora tritici-repentis)具有最高相似度,在蛋白水平上相似性达61%,故命名该基因为CMADA。进一步设计引物,自盾壳霉cDNA文库扩增得到3'末端。Southern杂交证实T-DNA标记在ZS-1TN250中为单拷贝。RT-PCR检测到CMADA基因在野生菌株中的积累,但在突变体ZS-1TN250中没有检测信号。用腺苷脱氨酶试剂盒测定ADA的活性,在野生菌株中可检测到该酶的活性,而突变体ZS-1TN250中未能检测到该酶的活性。因此,推定突变体表型的变化是由于T-DNA插入造成的,CMADA基因与盾壳霉的菌丝生长和孢子形成相关。
In this study, the efficient Agrobacterium tumefacies mediated transformation system was established to transform Coniothyrium minitans. Conidia from 14-18 dpi colony developed on PDA slant of ZS-1 were collected after centrifuge, and then the conidial concentration was adjusted and resuspended to bacterial liquid. The results showed that fifty transformants per plate were obtained. A T-DNA insertional library with 5000 transformants of ZS-1 was constructed using the method described above.
From this library,23 mutants were screened out, and among them 8 mutants could not produce any conidia on PDA plate,10 mutants could produce a few conidia,6 mutants produced abnormal pigment as compared with ZS-1,3 mutants grew faster on PDA than ZS-1,3 mutants grew slower than ZS-1, and hyphae of two mutant collapsed and then degraded. These mutants were significantly different in hyphal growth rate, antibiotics production, parasite of sclerotia, colony morphology and so on. Genomic DNA fragment flanking the insertional T-DNA were cloned from six mutants using Inverse-PCR technique, including ZS-1 TN29、ZS-1TN28、ZS-1TN5012、ZS-1TN5498 and ZS-1TN6222. Putative protein were CMARGJ, CMGAL、CMHMGR、CMPurD、CMMAPKKK respectively by conserved domain analysising.
Mutant ZS-1TN250 was studied in detail. The growth rate of mutant ZS-1TN250 was significantly faster than ZS-1, but the biomass in PDB was less. Sporulation was declined significantly and could produce a few conidia on PDA plate. The pycnidia could not break to release conidia, and most of the spores could not mature. This mutant could parasitize sclerotia of Sclerotinia sclerotiorum, but could not produce conidia on sclertia and rotting capacity was decreased significantly. Just like ZS-1, ZS-1TN250 could secrete antibiotic substances. Genomic DNA fragment flanking the insertional T-DNA of ZS-1TN250 was obtained by Inverse-PCR. Based on the flanking DNA, the sequence of a gene was abtained. The full length gene included 1522bp with 4 exons and 3 introns, and cDNA length was 996 bp, encoding a protein with 332 aa. The T-DNA inserted into the second exon, insertion point was at the 191nt position. The deduced amino acid sequence shared 61% identity with adenosine/AMP deaminase family protein of Pyrenophora tritici-repentis. Therefore, we named the gene as CMADA. Further primers were designed, and 3'end was amplified from the cDNA library of C. minitans. Southern blot analysis confirmed that single T-DNA was inserted in the ZS-1TN250. RT-PCR showed that CMADA gene was expressed in the wild strain ZS-lbut not in ZS-1TN250. This indicates that T-DNA insertion induced the mutant phenotype. The protein activity was detected with ADA activity determination kit. The enzyme activity could be detected in the wild strain, whereas could not in mutant ZS-1TN250. The results suggested that CMADA gene is related to mycelial growth and conodia formation of Coniothyrium minitans.
随后我们自盾壳霉的插入突变体库中共筛选到了23株表型异常突变体,其中8株突变体在PDA平皿上完全不能产生分生孢子,10株产孢明显减少,6株产生色素异常,3株生长迅速,3株生长缓慢,2株菌丝塌陷。其中产孢缺陷型突变体在菌丝生长速度、产抗生物质、寄生菌核能力和菌落形态等方面存在差异显著性。利用反向PCR、cDNA文库等克隆基因的技术平台,克隆了5株突变体的相关基因,包括ZS-1TN29、ZS-1TN289、ZS-1TN5012、ZS-1TN5498和ZS-1TN6222,利用NCBI保守结构域数据库比较分析,编码蛋白分别推定为CMARGJ、CMGAL、CMHMGR、CMPurD、CMMAPKKK。
对突变体ZS-1TN250进行了详细研究,发现该突变体生长速度迅速,但在PDB中的生物产量比野生菌株低；产孢明显减少,后期有少量孢子产生,显微观察发现大部分孢子没有黑色素化；该突变体能寄生核盘菌,但在菌核上不产生分生孢子,致腐菌核能力显著下降；可以正常产生抗生物质。采用反向PCR技术对ZS-1TN250中T-DNA标记插入位点的侧翼基因组DNA进行克隆,获得了大小为902bp的DNA片段,结合盾壳霉基因组序列分析,利用DNA MANagement软件进行全长序列拼接,得到该基因全长1522bp,包含4个外显子和3个内含子,cDNA全长为996 bp,推定编码含332个氨基酸的蛋白,T-DNA插入到第二个外显子上191nt的位置。通过NCBI BLASTP数据库,对推定编码的氨基酸序列进行分析,与adenosine/AMP deaminase family protein (Pyrenophora tritici-repentis)具有最高相似度,在蛋白水平上相似性达61%,故命名该基因为CMADA。进一步设计引物,自盾壳霉cDNA文库扩增得到3'末端。Southern杂交证实T-DNA标记在ZS-1TN250中为单拷贝。RT-PCR检测到CMADA基因在野生菌株中的积累,但在突变体ZS-1TN250中没有检测信号。用腺苷脱氨酶试剂盒测定ADA的活性,在野生菌株中可检测到该酶的活性,而突变体ZS-1TN250中未能检测到该酶的活性。因此,推定突变体表型的变化是由于T-DNA插入造成的,CMADA基因与盾壳霉的菌丝生长和孢子形成相关。
In this study, the efficient Agrobacterium tumefacies mediated transformation system was established to transform Coniothyrium minitans. Conidia from 14-18 dpi colony developed on PDA slant of ZS-1 were collected after centrifuge, and then the conidial concentration was adjusted and resuspended to bacterial liquid. The results showed that fifty transformants per plate were obtained. A T-DNA insertional library with 5000 transformants of ZS-1 was constructed using the method described above.
From this library,23 mutants were screened out, and among them 8 mutants could not produce any conidia on PDA plate,10 mutants could produce a few conidia,6 mutants produced abnormal pigment as compared with ZS-1,3 mutants grew faster on PDA than ZS-1,3 mutants grew slower than ZS-1, and hyphae of two mutant collapsed and then degraded. These mutants were significantly different in hyphal growth rate, antibiotics production, parasite of sclerotia, colony morphology and so on. Genomic DNA fragment flanking the insertional T-DNA were cloned from six mutants using Inverse-PCR technique, including ZS-1 TN29、ZS-1TN28、ZS-1TN5012、ZS-1TN5498 and ZS-1TN6222. Putative protein were CMARGJ, CMGAL、CMHMGR、CMPurD、CMMAPKKK respectively by conserved domain analysising.
Mutant ZS-1TN250 was studied in detail. The growth rate of mutant ZS-1TN250 was significantly faster than ZS-1, but the biomass in PDB was less. Sporulation was declined significantly and could produce a few conidia on PDA plate. The pycnidia could not break to release conidia, and most of the spores could not mature. This mutant could parasitize sclerotia of Sclerotinia sclerotiorum, but could not produce conidia on sclertia and rotting capacity was decreased significantly. Just like ZS-1, ZS-1TN250 could secrete antibiotic substances. Genomic DNA fragment flanking the insertional T-DNA of ZS-1TN250 was obtained by Inverse-PCR. Based on the flanking DNA, the sequence of a gene was abtained. The full length gene included 1522bp with 4 exons and 3 introns, and cDNA length was 996 bp, encoding a protein with 332 aa. The T-DNA inserted into the second exon, insertion point was at the 191nt position. The deduced amino acid sequence shared 61% identity with adenosine/AMP deaminase family protein of Pyrenophora tritici-repentis. Therefore, we named the gene as CMADA. Further primers were designed, and 3'end was amplified from the cDNA library of C. minitans. Southern blot analysis confirmed that single T-DNA was inserted in the ZS-1TN250. RT-PCR showed that CMADA gene was expressed in the wild strain ZS-lbut not in ZS-1TN250. This indicates that T-DNA insertion induced the mutant phenotype. The protein activity was detected with ADA activity determination kit. The enzyme activity could be detected in the wild strain, whereas could not in mutant ZS-1TN250. The results suggested that CMADA gene is related to mycelial growth and conodia formation of Coniothyrium minitans.
引文
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