芝麻香型白酒高温大曲细菌群落多样性研究
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摘要
芝麻香型白酒作为我国传统白酒的创新酒种,以独特的浓、酱、清融合的生产工艺特点和优雅的芝麻香味而闻名。高温制曲为芝麻香型白酒酿造提供丰富的微生物种类、生物酶、各类香味物质及其前驱物质,其质量好坏直接关系到芝麻香型白酒的出酒率和酒质。由于受到传统发酵特点及其研究方法的限制,使得对高温制曲过程中微生物的群落结构研究和认识还很有限。本文以山东扳倒井芝麻香型白酒高温大曲为研究对象,首次将现代分子生物学和分子生态学技术应用于高温大曲细菌群落结构研究中,为探索芝麻香型白酒高温大曲功能性微生物的作用机制提供科学依据。
本文利用非培养PCR-变性梯度凝胶电泳技术(PCR-DGGE)和16Sr DNAV5-V8区克隆文库构建技术,结合可培养细菌的分离筛选和多相鉴定技术,动态分析高温大曲曲房环境、母曲、大曲原料、大曲发酵前期、中期、中后期、后期和出房时的优势细菌群落结构,并对其中高温功能性细菌进行分离筛选和鉴定。研究结果发现高温大曲中细菌种类丰富,主要来源于曲房环境、母曲和大曲原料,随着发酵温度的升高形成以高温细菌为主的群落结构。曲房环境中第一优势细菌为芽孢杆菌属(Bacillus sp.),丰度为21.67%,在高温大曲发酵前期、中期和后期也被检测到,且其丰度分别为13.7%、12.32%和23.15%。高温大曲原料中的第一优势细菌为泛菌属(Pantoea sp),丰度为73.68%,仅在大曲发酵前期少量被检测出,第二优势细菌为魏斯氏菌属(Weissella sp.),丰度为16.99%,第三优势细菌乳杆菌属(Lactobacillus sp.),丰度为4.57%,在大曲发酵的前期、中期和后期被分别被检测到,其丰度分别为前期3.22%和11.29%、中期6.89%和25.11%、后期3.74%和7.47%。大曲发酵前期的第一优势细菌是高温放线菌属(Thermoactinomyces sp.),丰度为47.57%,发酵中期、后期和出房时也检测到该菌属,且丰度分别为4.93%,15.68%和4.35%。大曲发酵结束时的优势细菌是Rubellimicrobium sp.,丰度是31.52%。
研究还从高温大曲中分离、筛选出85株高温细菌,并获得1株产中性蛋白酶菌株M5,中性蛋白酶活力96.06U/g,显著高于对照菌株,经16S rDNA序列分析和gryA基因序列分析鉴定为枯草芽孢杆菌枯草亚种(Bacillus subtilis subsp. Subtilis.);分离筛选出的优势细菌ZM60经多相鉴定为普通高温放线菌(Thermoactinomycesvulgaris),该菌株55℃生长良好,37℃不生长,具白色气生菌丝,产孢子,具有碱性磷酸盐酶、酯酶、类脂酯酶及萘酚-AS-BI-磷酸水解酶的活力。作为高温大曲的优势细菌,该菌株在芝麻香型白酒特殊性风味的形成过程中的作用尚未报道过。
本研究除发现了过去曾研究报道过的高温大曲优势细菌芽孢杆菌属(Bacillussp.),魏斯氏菌属(Weissella sp.)和乳杆菌属(Lactobacillus sp.)外,还发现了尚未在高温大曲研究中报道过的细菌菌种,如泛菌属(Pantoea sp.),(?)嗜盐单胞菌属(Halomonas sp.),高温放线菌属(Thermoactinomyces sp.), Kroppenstedtia sp.,莱西氏菌属(Laceyella sp.),片球菌属(Pediococcus sp.)和志贺氏菌属(Shigellasp.)等,其中泛球菌属(Pantoea sp.)菌种来源于植物性原料中,高温放线菌属(Thermoactinomyces sp.)和莱西氏菌属(Laceyella sp.)为属于芽孢菌科的高温放线菌, Kroppenstedtia sp为近年来新发表菌属,志贺氏菌属(Shigella sp.)为常见致病菌,这些细菌在高温大曲制曲过程中发挥的功能性作用和带来的影响还有待下进一步深入研究。
The Sesame-flavor Liquor is one of innovative liquor types in China, which has been universally popular for its technology characteristics combined with multifold-flavor liquors and elegant sesame flavor. High temperature Daqu is a sacchariferous starter and it can provide large amounts of microorganisms, enzymes, flavor components and its precursors for brewing sesame-flavor liquor. So its quality is directly relevant to the output rate and quality of liquor. Due to the unique features of traditional fermentation technology and the limitation of research methods in this field, the knowledge of microbes community structure occurred in high temperature Daqu is still imperfect. Molecular biological and molecular ecological technologies were firstly used to reveal the bacterial community diversity from high temperature Daqu of Sesame-flavor liquor manufactured by Bandaojing Co. Ltd. Further, the results can also supply scientific data for study of the functional microbes'mechanism.
Culture-independent analysis of PCR-denaturing gradient gel electrophoresis (PCR-DGGE) and16S rDNA V5-V8region clone library construction combined with culture-dependent analysis of screening and identification technologies were applied to investigate dynamic bacterial community diversity of high temperature making processes, including environment of Daqu room, Muqu, raw material, prophase fermentation, middle fermentation, late fermentation and new Daqu. And the high temperature functional bacterial also were isolated and identified in this study. The results indicated that abundant bacterial community mainly originated from Daqu room envrionment, Muqu and raw material. The bacterial community structure based high temperature bacteria were formed with the increase of fermentation temperature. The first dominant bacteria (abundant%) Bacillus sp.(21.67%) was detected at the stage of prophase, middle and late fermentation, and the abundant were13.7%,12.32%and23.15%separately. The first dominant bacteria (abundant%) Pantoea sp.(73.68%) in raw material was only found a few amount at prophase fermentation, the second dominant bacteria (abundant%) Weissella sp.(16.99%) and the third dominant bacteria (abundant%) Lactobacillus sp.(4.57%) were detected separately3.22%and11.29%at the prophase fermentation,6.89%and25.11%at middle fermentation,3.74%and7.47%at late fermentation. The first dominant bacteria (abundant%) Thermoactinomyces sp.(47.57%) at prophase fermentation was also detected at the middle fermentation, late fermentation and new Daqu as abundant of4.93%,15.68%and4.35%. The predominant bacteria (abundant%) in new Daqu was Rubellimicrobium sp.(31.52%).
85thennophilic bacteria anti-high-temperature strains were selected according to plate screening and conical flask screening, we obtained a strain of higher neutral protease producing capability, whose enzyme activity reached to96.06U/g. This strain was identified to be Bacillus subtilis subsp. subtilis by16S rDNA and gryA sequencing. ZM60, the dominant thennophilic bacteria, was identified as Thennoactinomyces vulgaris by polyphasic identification. It growth at55℃and not growth at37℃, with white aerial mycelium and spore-forming. Zm60had enzyme activity of Alkaline phosphatase, Esterase, Lipid esterase and Naphthol-AS-BI-phosphate hydrolase. As the dominant bacteria in process of high temperature Daqu, the role of Thermoactinomyces vulgaris during the formation of particularity flavor of sesame-flavor liquor has not been reported before.
This study not only found the dominant bacteria as Bacillus sp., Weissella sp. and Lactobacillus sp. reported before, but also detected predominant bacteria as Pantoea sp., Halomonas sp., Thermoactinomyces sp., Kroppenstedtia sp., Laceyella sp., Pediococcus sp. and Shigella sp. havn't been reported before. Among these genera, Pantoea sp. was common endophytic bacteria, Thennoactinomyces sp. and Laceyella sp. were thennoactinomyces belong to family Bacillus, Kroppenstedtia sp. was new genus published in recent years and Shigella sp. was common pathogenic bacteria. The more research need to be applied in order to reveal the functional role of these dominant bacteria in the process of high temperature Daqu making.
本文利用非培养PCR-变性梯度凝胶电泳技术(PCR-DGGE)和16Sr DNAV5-V8区克隆文库构建技术,结合可培养细菌的分离筛选和多相鉴定技术,动态分析高温大曲曲房环境、母曲、大曲原料、大曲发酵前期、中期、中后期、后期和出房时的优势细菌群落结构,并对其中高温功能性细菌进行分离筛选和鉴定。研究结果发现高温大曲中细菌种类丰富,主要来源于曲房环境、母曲和大曲原料,随着发酵温度的升高形成以高温细菌为主的群落结构。曲房环境中第一优势细菌为芽孢杆菌属(Bacillus sp.),丰度为21.67%,在高温大曲发酵前期、中期和后期也被检测到,且其丰度分别为13.7%、12.32%和23.15%。高温大曲原料中的第一优势细菌为泛菌属(Pantoea sp),丰度为73.68%,仅在大曲发酵前期少量被检测出,第二优势细菌为魏斯氏菌属(Weissella sp.),丰度为16.99%,第三优势细菌乳杆菌属(Lactobacillus sp.),丰度为4.57%,在大曲发酵的前期、中期和后期被分别被检测到,其丰度分别为前期3.22%和11.29%、中期6.89%和25.11%、后期3.74%和7.47%。大曲发酵前期的第一优势细菌是高温放线菌属(Thermoactinomyces sp.),丰度为47.57%,发酵中期、后期和出房时也检测到该菌属,且丰度分别为4.93%,15.68%和4.35%。大曲发酵结束时的优势细菌是Rubellimicrobium sp.,丰度是31.52%。
研究还从高温大曲中分离、筛选出85株高温细菌,并获得1株产中性蛋白酶菌株M5,中性蛋白酶活力96.06U/g,显著高于对照菌株,经16S rDNA序列分析和gryA基因序列分析鉴定为枯草芽孢杆菌枯草亚种(Bacillus subtilis subsp. Subtilis.);分离筛选出的优势细菌ZM60经多相鉴定为普通高温放线菌(Thermoactinomycesvulgaris),该菌株55℃生长良好,37℃不生长,具白色气生菌丝,产孢子,具有碱性磷酸盐酶、酯酶、类脂酯酶及萘酚-AS-BI-磷酸水解酶的活力。作为高温大曲的优势细菌,该菌株在芝麻香型白酒特殊性风味的形成过程中的作用尚未报道过。
本研究除发现了过去曾研究报道过的高温大曲优势细菌芽孢杆菌属(Bacillussp.),魏斯氏菌属(Weissella sp.)和乳杆菌属(Lactobacillus sp.)外,还发现了尚未在高温大曲研究中报道过的细菌菌种,如泛菌属(Pantoea sp.),(?)嗜盐单胞菌属(Halomonas sp.),高温放线菌属(Thermoactinomyces sp.), Kroppenstedtia sp.,莱西氏菌属(Laceyella sp.),片球菌属(Pediococcus sp.)和志贺氏菌属(Shigellasp.)等,其中泛球菌属(Pantoea sp.)菌种来源于植物性原料中,高温放线菌属(Thermoactinomyces sp.)和莱西氏菌属(Laceyella sp.)为属于芽孢菌科的高温放线菌, Kroppenstedtia sp为近年来新发表菌属,志贺氏菌属(Shigella sp.)为常见致病菌,这些细菌在高温大曲制曲过程中发挥的功能性作用和带来的影响还有待下进一步深入研究。
The Sesame-flavor Liquor is one of innovative liquor types in China, which has been universally popular for its technology characteristics combined with multifold-flavor liquors and elegant sesame flavor. High temperature Daqu is a sacchariferous starter and it can provide large amounts of microorganisms, enzymes, flavor components and its precursors for brewing sesame-flavor liquor. So its quality is directly relevant to the output rate and quality of liquor. Due to the unique features of traditional fermentation technology and the limitation of research methods in this field, the knowledge of microbes community structure occurred in high temperature Daqu is still imperfect. Molecular biological and molecular ecological technologies were firstly used to reveal the bacterial community diversity from high temperature Daqu of Sesame-flavor liquor manufactured by Bandaojing Co. Ltd. Further, the results can also supply scientific data for study of the functional microbes'mechanism.
Culture-independent analysis of PCR-denaturing gradient gel electrophoresis (PCR-DGGE) and16S rDNA V5-V8region clone library construction combined with culture-dependent analysis of screening and identification technologies were applied to investigate dynamic bacterial community diversity of high temperature making processes, including environment of Daqu room, Muqu, raw material, prophase fermentation, middle fermentation, late fermentation and new Daqu. And the high temperature functional bacterial also were isolated and identified in this study. The results indicated that abundant bacterial community mainly originated from Daqu room envrionment, Muqu and raw material. The bacterial community structure based high temperature bacteria were formed with the increase of fermentation temperature. The first dominant bacteria (abundant%) Bacillus sp.(21.67%) was detected at the stage of prophase, middle and late fermentation, and the abundant were13.7%,12.32%and23.15%separately. The first dominant bacteria (abundant%) Pantoea sp.(73.68%) in raw material was only found a few amount at prophase fermentation, the second dominant bacteria (abundant%) Weissella sp.(16.99%) and the third dominant bacteria (abundant%) Lactobacillus sp.(4.57%) were detected separately3.22%and11.29%at the prophase fermentation,6.89%and25.11%at middle fermentation,3.74%and7.47%at late fermentation. The first dominant bacteria (abundant%) Thermoactinomyces sp.(47.57%) at prophase fermentation was also detected at the middle fermentation, late fermentation and new Daqu as abundant of4.93%,15.68%and4.35%. The predominant bacteria (abundant%) in new Daqu was Rubellimicrobium sp.(31.52%).
85thennophilic bacteria anti-high-temperature strains were selected according to plate screening and conical flask screening, we obtained a strain of higher neutral protease producing capability, whose enzyme activity reached to96.06U/g. This strain was identified to be Bacillus subtilis subsp. subtilis by16S rDNA and gryA sequencing. ZM60, the dominant thennophilic bacteria, was identified as Thennoactinomyces vulgaris by polyphasic identification. It growth at55℃and not growth at37℃, with white aerial mycelium and spore-forming. Zm60had enzyme activity of Alkaline phosphatase, Esterase, Lipid esterase and Naphthol-AS-BI-phosphate hydrolase. As the dominant bacteria in process of high temperature Daqu, the role of Thermoactinomyces vulgaris during the formation of particularity flavor of sesame-flavor liquor has not been reported before.
This study not only found the dominant bacteria as Bacillus sp., Weissella sp. and Lactobacillus sp. reported before, but also detected predominant bacteria as Pantoea sp., Halomonas sp., Thermoactinomyces sp., Kroppenstedtia sp., Laceyella sp., Pediococcus sp. and Shigella sp. havn't been reported before. Among these genera, Pantoea sp. was common endophytic bacteria, Thennoactinomyces sp. and Laceyella sp. were thennoactinomyces belong to family Bacillus, Kroppenstedtia sp. was new genus published in recent years and Shigella sp. was common pathogenic bacteria. The more research need to be applied in order to reveal the functional role of these dominant bacteria in the process of high temperature Daqu making.
引文
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