Bacillus subtilis ZJU15产生的抗菌肽研究
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摘要
从水稻叶片中分出一株具有较好抗菌活性的细菌菌株ZJU15,经常规的生理生化鉴定、显微镜观察和16S rDNA部分序列的检测分析,将该菌株鉴定为枯草芽孢杆菌(Bacillus subtilis)。经作用谱测定,该菌株的培养去菌液能够有效抑制多种食品腐败和致病革兰氏阳性细菌,如单核细胞增生李斯特菌(Listeria monocytogenes)、藤黄微球菌(Micrococcus luteus)、粪肠球菌(Enterococcus feacalis)、金色葡萄球菌(Staphylococcus aureus)、肺炎链球菌(Streptococcus pyogenes)等,但该培养去菌液对革兰氏阴性细菌没有抑制作用。此外,该菌株的培养去菌液对引起柑橘采后腐败的意大利青霉(Penicillium digitatum)、引起多种食品霉变的黄曲霉(Aspergillus flavus)和水稻纹枯病菌(Rhizoctonia solani)等也具有明显的抑制作用,但对酵母样真菌念珠菌(Candida spp.)没有抗菌作用。
通过不同配方培养基的培养比较研究,确定CM培养基最适合ZJU15菌株抗菌物质产生。通过生长曲线和活性曲线测定,发现该菌株在培养30小时左右去菌培养液具有最大的抗菌活性。不同硫酸铵饱和浓度沉淀试验结果显示20%-30%的硫酸铵饱和浓度能使ZJU15菌株产生的拮抗物质得到最大程度的沉淀。在此基础上,将ZJU15菌株接种于液体培养中,35℃下振荡培养30小时,所得发酵液通过离心去菌、硫酸铵沉淀、Superdex 75 prep grade柱层析和RP-HPLC色谱(Hypersil ODS2色谱柱)纯化,获得三个具有抗菌活性的洗脱峰。它们的洗脱时间分别为11.762Min,12.767Min,14.340Min,分别命名为ZJU15A、ZJU15B和ZJU15C。将所得的抗菌肽提交中国人民解放军军事医学科学院生物医学分析中心进行MALDI-TOF-MS质谱扫描分析(扫描范围50-5000Da)、Q-TOF2一级质谱扫描得到LC-ESI-MS图。选择待测离子,然后进行ESI-MS/MS分析。结果显示ZJU15菌株产生的三个抑菌活性成分的分子量分别为1021.70Da,1021.71 Da和1035.69 Da；它们的氨基酸序列分别为LLELLVPLL, LLELLVPLL和LTHMLVPLL.经BLAST序列查询,没有发现SWISS-PROT/TrEMBL数据库中相似抗菌肽序列存在,我们认为它们为新的抗菌肽。
ZJU15菌株的培养去菌液经蛋白酶K处理后完全失去抑菌活性；对蛋白酶E部分敏感,但对胰蛋白酶、胃蚕白酶不敏感。不同pH值处理的结果显示,在pH2-10之间ZJU15菌株的培养去菌液保持抑菌活性不变,在pHl1、pH12处理下抑菌活性显著下降。ZJU15菌株的培养去菌液经37℃、60℃、80℃、100℃、121℃(高压)各处理30min,其抑菌活性保持不变。
将不同浓度的抑菌物质粗提液加入到对数生长中期单增李斯特菌液中,24h后涂板测定茵落总数。结果显示512AU/ml的活性浓度是对对数生长中期单增李斯特菌的最小杀菌浓度,256AU/ml的活性浓度是对对数生长中期单增李斯特菌的最小抑菌浓度。对数生长中期的单增李斯特菌经512AU/ml活性浓度的抑菌物质粗提液处理后,经透射电镜观察发现：单增李斯特菌的菌体边缘模糊、空泡化、胞壁缺陷。此结果表明,ZJU15菌株产生的抑菌物质能够导致单增李斯特菌细胞壁破裂或者形成孔洞,导致内容物泄漏,最终导致细菌细胞的死亡。
棋盘法联合测定抑菌活性研究结果表明：乳酸链球菌素和ZJU15菌株培养去菌液具有协同增效作用。
经ZJU15菌株抑菌物质粗提液对小鼠急性经口灌胃毒性试验,初步表明最高剂量50ml/kg的ZJU15菌株抑菌物质粗提液没有造成雌雄小鼠死亡和异常,即ZJU15菌株抑菌物质粗提液的急性经口LD50>50000mg/kg,ZJU15菌株抑菌物质粗提液对小鼠的急性经口毒性属无毒。
为了评估ZJU15菌株产生的抗菌物质对食品的防腐保鲜效果,我们以腊肉、豆浆和牛奶为材料进行了试验。结果表明经终浓度为512AU/g或1024AU/g的抑菌粗提物处理的未加工腊肉,在37℃保存条件下,7天后出现白色菌斑,而灭菌水对照3天就出现白色菌斑；经终浓度1024AU/ml和512AU/ml的抑菌粗提物处理新鲜豆浆,在37℃的条件下放置16h后没有检测到微生物的存在。商品纯牛奶经终浓度1024AU/ml的抑菌粗提物处理,可有效减少牛奶中的菌落数。以上结果显示出ZJU15菌株产生的抗菌肽在食品加工上有较好的应用前景。
A bacterial strain ZJU15, isolated from rice leaf in Hangzhou, China and was identified as Bacillus subtilis ZJU15 according to the conventional physiological biochemistry properties, electron microscope and 16S rDNA sequence analysis. The cell-free supernatant (CFS) of B.subtilis ZJU15 showed antimicrobial activity against several important Gram-positive pathogenic and food-spoilage bacteria, which including Enterococcus feacalis, Listeria monocytogenes, Micrococcus luteus, meticillin-resistant Staphylococcus aureus (MRSA) and S. epidermidis, but it was neither inhibitory to the Gram-Negative bacteria nor the closely related Bacillus bacteria such as B. subtilis, B. pumilus and B. licheniformis. The CFS could also inhibit the growth of some fungal strains such as Aspergillus flavus, Penicillium digitatum and Rhizoctonia solani, but could not inhibit the growth of Candida spp.
The comparison of different culture mediums indicated that CM culture medium was most suitable for the production and extraction of the antibacterial peptides of strain ZJU15. Through the growth curve and the activity detection, it was found that the biggest inhibiting activity reached after the culture of 30 hours. The inhibiting activity of suspension could reach maximum after ammonium sulfate precipitation with 20%-30% saturateion. Based on these results, strain ZJU15 was incubated in CM medium under shake cultivation for 30 hours at 35℃. Three AMPs designated ZJU15A, ZJU15B and ZJU15C were purified from the CFS by sequential ammonium sulphate precipitation with 20% saturateion, gel filtration and reverse phase high-performance liquid chromatography (RP-HPLC).
Matrix-assisted laser desorption/ionization time-of-flight mass spectrometry (MALDI-TOF-MS), Electrospray ionization mass spectrometry (ESI/MS) and Electrospray ionization tandem mass spectrometry (ESI-MS/MS) was performed at the Chinese People's Liberation Army Academy of Military Medical Sciences Biomedicine Analysis Center and the results revealed that the ZJU15A, ZJU15B, and ZJU15C had molecular mass of 1021.70 Da, 1021.71 Da, and 1035.69 Da with the sequences of nine amino acid residues:LLELLVPLL for ZJU15A, LLELLVPLL for ZJU15B and LTHMLVPLL for ZJU15C respectively. There had no significant homology to the known peptides in the database, in our knowledge which indicating that they may be novel antimicrobial peptides.
The cell-free culture of strain ZJU15 lost the inhibiting activity completely after the treatment of proteinase K, and was sensitive to the proteinase E partially, but was stable after the treatment of trypsin, pepsin. The inhibiting activity of the cell-free culture of ZJU15 was invariable at pH2-10, and droped obviously at pHl land pH12. The inhibiting activity of the cell-free culture of strain ZJU15 maintains invariable after treatment at 37℃,60℃,80℃,100℃or 121℃for 30min.
The culture of Listeria monocytogenes at early expression phase was added with different concentration of the Crude Antimicrobial Peptides Supernatant (CAPS) of strain ZJU 15, and the colony-forming units (CFU)/ml was detected after 24h. The results showed that 256AU/ml of the CAPS was the minimum bactericidal concentration (MBC) and 512AU/ml of the CAPS was the minimum inhibitory concentration (MIC). The cells edge blurring, yacuole, wall flaw was observed by transmission electron microscope after the addtion of 512AU/ml of the CAPS to the culture of L. monocytogenes. This result indicated that the CAPS of strain ZJU 15 could cause the cell wall of L. monocytogenes to burst or to form the hole, which resulted in the contents divulging and the cell death finally.
The results indicated that the CFS of strain ZJU 15 and the nisin have the coordination synergized action.
The acute oral toxicity on the mouse showed that the CAPS of strain ZJU15 was not caused death to the tested mouses at the dosage of 50ml/kg.
In order to estimate the preservation efficiency of the antimicrobial peptides produced by ZJU 15 strain on foods, cured meat, fresh soya-bean milk and milk were tested by adding the crude antimicrobial peptides of strain ZJU15. The cured meat, which were treated with 1024 AU/g and 512 AU/g as final concentration of the CAPS or dH2O (as control), were kept at 37℃. After 7d, white microbial spots appeared on the cured meat with the CAPS at 1024 AU/g or 512 AU/g, which was four days later than the time white microbial spots appeared on the cured meat with dH2O. The fresh soya-bean milk, which was treated with 1024 AU/g or 512 AU/g as final concentration of the CAPS or dH2O (as control), was kept at 37℃. After cultivation for16h, no microorganisms was detected in the fresh soya-bean milk with the crude antimicrobial peptides at 1024 AU/g or 512 AU/g. The commercial pure milk, which was treated with 1024 AU/g or 512 AU/g as final concentration of the CAPS or dH2O (as control), was kept at 37℃. After cultivation for 20h, the number of microorganisms was less greatly in the fresh soya-bean milk which incubated with the crude antimicrobial peptides at 1024 AU/g than other treatments.
通过不同配方培养基的培养比较研究,确定CM培养基最适合ZJU15菌株抗菌物质产生。通过生长曲线和活性曲线测定,发现该菌株在培养30小时左右去菌培养液具有最大的抗菌活性。不同硫酸铵饱和浓度沉淀试验结果显示20%-30%的硫酸铵饱和浓度能使ZJU15菌株产生的拮抗物质得到最大程度的沉淀。在此基础上,将ZJU15菌株接种于液体培养中,35℃下振荡培养30小时,所得发酵液通过离心去菌、硫酸铵沉淀、Superdex 75 prep grade柱层析和RP-HPLC色谱(Hypersil ODS2色谱柱)纯化,获得三个具有抗菌活性的洗脱峰。它们的洗脱时间分别为11.762Min,12.767Min,14.340Min,分别命名为ZJU15A、ZJU15B和ZJU15C。将所得的抗菌肽提交中国人民解放军军事医学科学院生物医学分析中心进行MALDI-TOF-MS质谱扫描分析(扫描范围50-5000Da)、Q-TOF2一级质谱扫描得到LC-ESI-MS图。选择待测离子,然后进行ESI-MS/MS分析。结果显示ZJU15菌株产生的三个抑菌活性成分的分子量分别为1021.70Da,1021.71 Da和1035.69 Da；它们的氨基酸序列分别为LLELLVPLL, LLELLVPLL和LTHMLVPLL.经BLAST序列查询,没有发现SWISS-PROT/TrEMBL数据库中相似抗菌肽序列存在,我们认为它们为新的抗菌肽。
ZJU15菌株的培养去菌液经蛋白酶K处理后完全失去抑菌活性；对蛋白酶E部分敏感,但对胰蛋白酶、胃蚕白酶不敏感。不同pH值处理的结果显示,在pH2-10之间ZJU15菌株的培养去菌液保持抑菌活性不变,在pHl1、pH12处理下抑菌活性显著下降。ZJU15菌株的培养去菌液经37℃、60℃、80℃、100℃、121℃(高压)各处理30min,其抑菌活性保持不变。
将不同浓度的抑菌物质粗提液加入到对数生长中期单增李斯特菌液中,24h后涂板测定茵落总数。结果显示512AU/ml的活性浓度是对对数生长中期单增李斯特菌的最小杀菌浓度,256AU/ml的活性浓度是对对数生长中期单增李斯特菌的最小抑菌浓度。对数生长中期的单增李斯特菌经512AU/ml活性浓度的抑菌物质粗提液处理后,经透射电镜观察发现：单增李斯特菌的菌体边缘模糊、空泡化、胞壁缺陷。此结果表明,ZJU15菌株产生的抑菌物质能够导致单增李斯特菌细胞壁破裂或者形成孔洞,导致内容物泄漏,最终导致细菌细胞的死亡。
棋盘法联合测定抑菌活性研究结果表明：乳酸链球菌素和ZJU15菌株培养去菌液具有协同增效作用。
经ZJU15菌株抑菌物质粗提液对小鼠急性经口灌胃毒性试验,初步表明最高剂量50ml/kg的ZJU15菌株抑菌物质粗提液没有造成雌雄小鼠死亡和异常,即ZJU15菌株抑菌物质粗提液的急性经口LD50>50000mg/kg,ZJU15菌株抑菌物质粗提液对小鼠的急性经口毒性属无毒。
为了评估ZJU15菌株产生的抗菌物质对食品的防腐保鲜效果,我们以腊肉、豆浆和牛奶为材料进行了试验。结果表明经终浓度为512AU/g或1024AU/g的抑菌粗提物处理的未加工腊肉,在37℃保存条件下,7天后出现白色菌斑,而灭菌水对照3天就出现白色菌斑；经终浓度1024AU/ml和512AU/ml的抑菌粗提物处理新鲜豆浆,在37℃的条件下放置16h后没有检测到微生物的存在。商品纯牛奶经终浓度1024AU/ml的抑菌粗提物处理,可有效减少牛奶中的菌落数。以上结果显示出ZJU15菌株产生的抗菌肽在食品加工上有较好的应用前景。
A bacterial strain ZJU15, isolated from rice leaf in Hangzhou, China and was identified as Bacillus subtilis ZJU15 according to the conventional physiological biochemistry properties, electron microscope and 16S rDNA sequence analysis. The cell-free supernatant (CFS) of B.subtilis ZJU15 showed antimicrobial activity against several important Gram-positive pathogenic and food-spoilage bacteria, which including Enterococcus feacalis, Listeria monocytogenes, Micrococcus luteus, meticillin-resistant Staphylococcus aureus (MRSA) and S. epidermidis, but it was neither inhibitory to the Gram-Negative bacteria nor the closely related Bacillus bacteria such as B. subtilis, B. pumilus and B. licheniformis. The CFS could also inhibit the growth of some fungal strains such as Aspergillus flavus, Penicillium digitatum and Rhizoctonia solani, but could not inhibit the growth of Candida spp.
The comparison of different culture mediums indicated that CM culture medium was most suitable for the production and extraction of the antibacterial peptides of strain ZJU15. Through the growth curve and the activity detection, it was found that the biggest inhibiting activity reached after the culture of 30 hours. The inhibiting activity of suspension could reach maximum after ammonium sulfate precipitation with 20%-30% saturateion. Based on these results, strain ZJU15 was incubated in CM medium under shake cultivation for 30 hours at 35℃. Three AMPs designated ZJU15A, ZJU15B and ZJU15C were purified from the CFS by sequential ammonium sulphate precipitation with 20% saturateion, gel filtration and reverse phase high-performance liquid chromatography (RP-HPLC).
Matrix-assisted laser desorption/ionization time-of-flight mass spectrometry (MALDI-TOF-MS), Electrospray ionization mass spectrometry (ESI/MS) and Electrospray ionization tandem mass spectrometry (ESI-MS/MS) was performed at the Chinese People's Liberation Army Academy of Military Medical Sciences Biomedicine Analysis Center and the results revealed that the ZJU15A, ZJU15B, and ZJU15C had molecular mass of 1021.70 Da, 1021.71 Da, and 1035.69 Da with the sequences of nine amino acid residues:LLELLVPLL for ZJU15A, LLELLVPLL for ZJU15B and LTHMLVPLL for ZJU15C respectively. There had no significant homology to the known peptides in the database, in our knowledge which indicating that they may be novel antimicrobial peptides.
The cell-free culture of strain ZJU15 lost the inhibiting activity completely after the treatment of proteinase K, and was sensitive to the proteinase E partially, but was stable after the treatment of trypsin, pepsin. The inhibiting activity of the cell-free culture of ZJU15 was invariable at pH2-10, and droped obviously at pHl land pH12. The inhibiting activity of the cell-free culture of strain ZJU15 maintains invariable after treatment at 37℃,60℃,80℃,100℃or 121℃for 30min.
The culture of Listeria monocytogenes at early expression phase was added with different concentration of the Crude Antimicrobial Peptides Supernatant (CAPS) of strain ZJU 15, and the colony-forming units (CFU)/ml was detected after 24h. The results showed that 256AU/ml of the CAPS was the minimum bactericidal concentration (MBC) and 512AU/ml of the CAPS was the minimum inhibitory concentration (MIC). The cells edge blurring, yacuole, wall flaw was observed by transmission electron microscope after the addtion of 512AU/ml of the CAPS to the culture of L. monocytogenes. This result indicated that the CAPS of strain ZJU 15 could cause the cell wall of L. monocytogenes to burst or to form the hole, which resulted in the contents divulging and the cell death finally.
The results indicated that the CFS of strain ZJU 15 and the nisin have the coordination synergized action.
The acute oral toxicity on the mouse showed that the CAPS of strain ZJU15 was not caused death to the tested mouses at the dosage of 50ml/kg.
In order to estimate the preservation efficiency of the antimicrobial peptides produced by ZJU 15 strain on foods, cured meat, fresh soya-bean milk and milk were tested by adding the crude antimicrobial peptides of strain ZJU15. The cured meat, which were treated with 1024 AU/g and 512 AU/g as final concentration of the CAPS or dH2O (as control), were kept at 37℃. After 7d, white microbial spots appeared on the cured meat with the CAPS at 1024 AU/g or 512 AU/g, which was four days later than the time white microbial spots appeared on the cured meat with dH2O. The fresh soya-bean milk, which was treated with 1024 AU/g or 512 AU/g as final concentration of the CAPS or dH2O (as control), was kept at 37℃. After cultivation for16h, no microorganisms was detected in the fresh soya-bean milk with the crude antimicrobial peptides at 1024 AU/g or 512 AU/g. The commercial pure milk, which was treated with 1024 AU/g or 512 AU/g as final concentration of the CAPS or dH2O (as control), was kept at 37℃. After cultivation for 20h, the number of microorganisms was less greatly in the fresh soya-bean milk which incubated with the crude antimicrobial peptides at 1024 AU/g than other treatments.
引文
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