CN110317768A - 一种消减4-甲基苯酚的乳酸菌及其在酿造体系中的应用 - Google Patents
一种消减4-甲基苯酚的乳酸菌及其在酿造体系中的应用 Download PDFInfo
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- CN110317768A CN110317768A CN201910618450.XA CN201910618450A CN110317768A CN 110317768 A CN110317768 A CN 110317768A CN 201910618450 A CN201910618450 A CN 201910618450A CN 110317768 A CN110317768 A CN 110317768A
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- methylphenol
- lactobacillus brevis
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Abstract
本发明公开了一种消减4‑甲基苯酚的乳酸菌及其在酿造体系中的应用,属于生物技术领域。本发明的消减4‑甲基苯酚的乳酸菌,表达了来源于谷氨酸棒状杆菌的4‑甲基苯基磷酸酯合成酶的编码基因creI与creH,该菌株能够有效的转化4‑甲基苯酚为4‑甲基苯基磷酸酯,增加产物的沸点,降低白酒中的4‑甲基苯酚含量,达到在酿造体系中消减4‑甲基苯酚的能力,对今后完全降解酿造体系中4‑甲基苯酚提供新的策略。
Description
技术领域
本发明涉及一种消减4-甲基苯酚的乳酸菌及其在酿造体系中的应用,属于生物技术领域。
背景技术
4-甲基苯酚(p-cresol)是一种可随水蒸气挥发,呈现窖泥臭、皮革臭、焦皮臭、动物臭的芳香族化合物。4-甲基苯酚是酪氨酸的降解产物,现已发现奶酪、葡萄酒、威士忌、白酒等发酵食品中均含有微量的4-甲基苯酚。4-甲基苯酚阈值很低(46%酒精中的阈值为116.97μg/L),ppm级别的4-甲基苯酚即可对食品的风味造成不良影响。
根据生产工艺不同,白酒可以细分为十二大香型,其中药香型、浓香型、酱香型、兼香型、等七种基于泥窖或半泥窖发酵的白酒中均检测到了4-甲基苯酚,该物质也是产生窖泥臭的重要化合物。目前,甲基苯酚的去除手段包括物理化学法和生物法。物理化学法如催化氧化、吸附剂吸附等,处理成本高,易产生毒性副产物,限制了应用范围。通过利用活性炭吸附虽然能较好消除白酒中的4-甲基苯酚,但同时也会非特异性吸附其他风味物质,对白酒的品质会有较大影响。4-甲基苯酚微生物降解法在环境修复领域得到了广泛的研究,4-甲基苯酚降解菌可从高4-甲基苯酚环境中进行筛选,目前发现的4-甲基苯酚降解菌包括谷氨酸棒杆菌(Corynebacterium glutamicum),黏质沙雷氏菌(Serratia marcescens),产碱杆菌(Advenella sp.),芽孢杆菌(Bacillus sp.),假单胞菌(Pseudomonas sp.),嗜麦芽窄食单胞菌(Stenotrophomonas maltophilia),黏鞭霉菌(Gliomastix indicus)。这些微生物多生长于中性和弱碱性环境中,且当培养基中超过一定浓度的4-甲基苯酚会对微生物生长产生抑制作用。因白酒酿造体系(酒醅)为酸性环境,发酵过程中产生的乳酸、乙酸、丁酸和己酸等使酒醅的pH值低至3.5左右,现发现的4-甲基苯酚降解菌难以在白酒酿造体系中生长。
已有报道发现,在泥窖中强化接种不产或低产4-甲基苯酚的菌株或菌剂,抑制窖泥中的4-甲基苯酚产生菌的生长,控制窖泥中4-甲基苯酚的产生,进而控制白酒中的4-甲基苯酚的含量。虽然该方法也可改善白酒中4-甲基苯酚的含量,但是对于强化菌株或菌剂无法抑制的4-甲基苯酚产生菌仍然会在泥窖中存在,这部分菌株产生的4-甲基苯酚仍然无法得到根本上的去除。而且该方法仅对缓解具有泥窖且泥窖中微生物菌群能与强化菌株或菌剂适宜的情况下有作用,对于强化菌株无法定植的泥窖或其他含有4-甲基苯酚的发酵食品是不适用的,应用范围受到限制。
因此,如何保证在正常发酵且风味产生无影响的情况下,有效消除或减少白酒酿造体系中已产生的4-甲基苯酚仍需深入研究。
发明内容
乳杆菌在白酒酿造过程中逐步占据主导地位,在发酵中后期成为酿造体系中的优势微生物。乳杆菌中的大部分种被公认为安全级微生物,在食品、医药、饲料等相关领域被广泛应用。白酒酿造体系中的乳杆菌不具有降解4-甲基苯酚降解途径。
为了解决上述问题,本发明提供了一种消减4-甲基苯酚的短乳杆菌菌株的构建方法及其消减4-甲基苯酚的应用。
4-甲基苯酚的沸点为202℃,远高于水和乙醇的沸点。4-甲基苯酚之所以能进入白酒酒体中,主要是由于固态蒸馏过程中存在较强的夹带作用,可以将高沸点物质随酒精和水带入酒体中。谷氨酸棒状杆菌作为重要的工业生产菌株,具有区别与其他4-甲基苯酚降解菌株的独特降解途径,菌株首先通过4-甲基苯基磷酸酯合成酶将4-甲基苯酚磷酸酯化成4-甲基苯基磷酸酯,不仅减少了4-甲基苯酚,同时产生的4-甲基苯基磷酸酯更易被cre基因簇编码的其他酶识别代谢,促进了4-甲基苯酚的降解。在本发明中,发明人发现,谷氨酸棒状杆菌对低浓度(7mg/L)的4-甲基苯酚有较优的降解效果,几乎可完全降解,降解率达99%以上,发明人猜测对于白酒酿造体系的4-甲基苯酚的降解选择此降解途径进行降解是较适宜的。但谷氨酸棒状杆菌无法耐受酸性和缺氧的白酒酿造环境,限制了其在酿造体系中对4-甲基苯酚进行降解的应用。
谷氨酸棒状杆菌的4-甲基苯酚降解的关键酶包括4-甲基苯基磷酸酯合成酶(CreHI),磷酸基团识别体(CreJEF),磷酸水解酶(CreD),醇脱氢酶(CreC),醛脱氢酶(CreG)。整个编码基因簇cre长达11kb,而短乳杆菌自身基因组较小,仅3M左右,若将整个cre基因簇均在短乳杆菌中表达,是否会对菌株自生长造成不利于影响尚需深入研究。白酒作为蒸馏酒,含有4-甲基苯酚主要是由于酒醅中产生的4-甲基苯酚被蒸馏到原酒中,将4-甲基苯酚转化成高沸点物质即可减少原酒中的4-甲基苯酚含量。因此,本发明仅将编码4-甲基苯基磷酸酯合成酶的编码基因creI与creH转入短乳杆菌中,表达的4-甲基苯基磷酸酯合成酶可转化4-甲基苯酚(沸点202℃)为4-甲基甲苯磷酸酯(沸点大于347℃),减少酒醅中的4-甲基苯酚进入原酒中。该方法操作简单,可有效降低酿造体系的4-甲基苯酚。
本发明的第一个目的是提供一种消减4-甲基苯酚短乳杆菌,所述消减4-甲基苯酚短乳杆菌是在短乳杆菌宿主中过表达creI与creH基因(4-甲基苯基磷酸酯合成酶的编码基因)。
在一种实施方式中,编码所述creI基因的核苷酸序列如SEQ ID NO:1所示、氨基酸序列如SEQ ID NO:8所示。
在一种实施方式中,编码所述creH基因的核苷酸序列如SEQ ID NO:2所示、氨基酸序列如SEQ ID NO:9所示。
在一种实施方式中,所述短乳杆菌菌株为短乳杆菌D17(保藏编号为CGMCCNO.14385)或者模式菌株短乳杆菌ATCC 367、短乳杆菌NCL912、短乳杆菌CGMCC1306、短乳杆菌145等。
在一种实施方式中,所述过表达载体,是任意一种已知的乳酸菌表达载体,比如pMG36e,pVE5523,pNZ8048,pNZ8148等。
在一种实施方式中,creI与creH基因是通过串联进行表达。
在一种实施方式中,串联creI与creH的RBS序列如SEQ ID NO:3所示、氨基酸序列如SEQ ID NO:10所示。
在一种实施方式中,过表达,具体是:扩增得到creI、creH基因,将creI、creH基因通过RBS序列串联得到creI-RBS-creH片段;然后连接到表达载体pMG36e上获得过表达载体pMG36e-creIH;pMG36e-creIH电转化入短乳杆菌感受态细胞中,筛选得到creI和creH基因表达正确的过表达短乳杆菌菌株。
在一种实施方式中,creI与creH基因在短乳杆菌中的过表达,具体是:以谷氨酸棒状杆菌ATCC 13032的全基因组DNA为模板,分别扩增creI与creH基因。通过重叠PCR方法将creI与creH按基因组的方向creI-creH与设计的RBS序列串联在一起,得到的片段creI-RBS-creH,连接到乳酸菌的表达载体pMG36e上,获得过表达载体pMG36e-creIH。制备短乳杆菌感受态细胞,将过表达载体pMG36e-creIH电转化入短乳杆菌中,通过在红霉素抗性平板上筛选得到含有过表达载体pMG36e-creIH的短乳杆菌菌株,得到creIH过表达短乳杆菌菌株。
本发明的第二个目的是提供一种确定表达载体在本发明的短乳杆菌中的稳定性的方法,所述方法是在无抗性条件下,检测过表达菌株的过表达质粒维持率。
将creIH过表达短乳杆菌株接种于无抗生素的GYP液体培养基中,37℃,静置培养。每24h,48h,72h取样,利用0.9%的NaCl生理盐水梯度稀释后,分别涂布于不含红霉素的GYP平板与含4μg/L红霉素的GYP平板上,37℃静置培养48h后,分别计数。用在有抗平板上的菌落数与在无抗平板上的菌落数的比值作为质粒维持率。通过质粒维持率表征过表达质粒在短乳杆菌中的稳定性。优选的,培养48h,过表达载体的质粒维持率仍为90%以上。
本发明的第三个目的是提供一种消减4-甲基苯酚的方法,所述方法是利用creIH过表达短乳杆菌菌株进行发酵;所述creIH过表达短乳杆菌,过表达了4-甲基苯基磷酸酯合成酶的编码基因。
在一种实施方式中,所述4-甲基苯基磷酸酯合成酶的编码基因为creI与creH基因。可选地,编码所述creI基因的氨基酸序列如SEQ ID NO:8所示。可选地,编码所述creH基因的氨基酸序列如SEQ ID NO:9所示。
在一种实施方式中,所述方法是将creIH过表达短乳杆菌接种于含有4-甲基苯酚的系统中进行培养。
在一种实施方式中,所述含有4-甲基苯酚的系统可以是液态系统,也可以固态系统,比如:酒醅、奶酪、威士忌、白酒等。
在一种实施方式中,将过表达creIH的短乳杆菌接种于的GYP液体发酵培养基中,添加7.3mg/L 4-甲基苯酚,于37℃,静置培养。优选的,培养48h,creIH过表达菌株可消减4-甲基苯酚,消减能力达2130μg/L,消减率达30%。
本发明的第四个目的是提供一种在固态酿造体系消减4-甲基苯酚的方法,所述方法是利用creIH过表达短乳杆菌菌株在固态酿造体系中发酵;所述creIH过表达短乳杆菌,过表达了4-甲基苯基磷酸酯合成酶的编码基因。
在一种实施方式中,所述4-甲基苯基磷酸酯合成酶的编码基因为creI与creH基因。可选地,编码所述creI基因的氨基酸序列如SEQ ID NO:8所示。可选地,编码所述creH基因的氨基酸序列如SEQ ID NO:9所示。
在一种实施方式中,所述发酵是在模拟的固态酿造体系中进行。
在一种实施方式中,所述发酵是在固态酿造的出池酒醅中接种本发明的短乳杆菌菌株,静置培养一段时间。然后将酒醅进行蒸馏。
将creIH过表达短乳杆菌株接种于GYP液体发酵培养基中,37℃培养12h后,离心收集菌体。用10%培养液体积的NaCl生理盐水重悬后,取1mL加入100g的含有4-甲基苯酚的出池酒醅中,搅拌均匀,密封。37℃静置培养。优选的,培养48h,creI-creH过表达菌株可消减酿造体系中的4-甲基苯酚,消减能力达530μg/L,消减率达37.9%。
生物材料:
谷氨酸棒状杆菌ATCC 13032,已在1967年文献公开(Shigeo A.,Ken-Ichiro T,Shukuo Kinoshita.Taxonomical studies on glutamic acid-producing bacteria,1967,13(3):279-301)。
短乳杆菌D17,已经在公布号CN 108034599 A、公开日为2018年05月15日的专利申请2017112754179中公开;其分类学命名为短乳杆菌Lactobacillus brevis,于2017年7月6日保藏于中国微生物菌种保藏管理委员会普通微生物中心,保藏编号为CGMCC NO.14385。
模式菌株短乳杆菌ATCC 367,购自中国微生物保藏中心编号CGMCC1.2028,也可以从美国模式培养物研究所编号ATCC 367购买得到。
本发明涉及的短乳杆菌NCL912、短乳杆菌CGMCC1306及短乳杆菌145已在本专利申请之前的文献中公开(Li,H.X.,Gao,D.D.,et al.A highγ-aminobutyric acid-producing Lactobacillus brevis isolated from Chinese traditional paocai[J].2008,Annals of Microbiology,58(4):649-653.Peng,C.L.,Huang,J.,et al.A Two-stage pH and Temperature Control with Substrate Feeding Strategy forProduction of Gamma-aminobutyric Acid by Lactobacillus brevis CGMCC 1306[J].Chinese Journal of Chemical Engineering,21(10):1190-1194.Wu,Q.,Law,Y.S.,etal.Dairy Streptococcus thermophilus improves cell viability of Lactobacillusbrevis NPS-QW-145and its gamma-aminobutyric acid biosynthesis ability in milk[J].Scientific Report,2015,5(12885):1-12.)。
表达载体pMG36e,如图1所示,已经在1989年的文献中公开(van de Guchte M.,van der Vossen J.M.B.M.,Kok J.,Venema G.Construction of a LactococcalExpression Vector:Expression of Hen Egg White Lysozyme in Lactococcus lactissubsp.lactis.Applied Environmental Microbiology 1989,55(1):224-228.)。
本发明涉及的质粒pVE5523,pNZ8048,pNZ8148已在本专利申请之前的文献中公开(Dieye,Y.,Usai,S.,Clier,F.,Gruss,A.,Piard,J.-C.Design of a protein-targetingsystem for lactic acid bacteria.2001,Journal of Bacteriology,183(14):4157-4166.Ruyter P.G.D.,Kuipers O.P.,Vos W.M.D.Controlled gene expression systemsfor Lactococcus lactis with the food-grade inducer nisin.Applied andEnvironmental Microbiology,1996,62(10):3662-3667.Igor M.,Michiel K..10yearsof the nisin-controlled gene expression system(NICE)in Lactococcuslactis.2005,Applied Microbiology and Biotechnology,68(6):705-717.)。
本发明的有益效果:
本发明首次改造酿造体系中的主体微生物-乳酸菌,使其具有4-甲基苯酚的消减能力。该方法简单,便于操作,具有通用性,原则上适合所有的乳杆菌。本发明利用改造的过表达菌株消减酿造体系中的4-甲基苯酚,可有效减少酿造体系中的异味物质,提升白酒品质。该菌株也适用于其他具有4-甲基苯酚异味的发酵食品中。
附图说明
图1.谷氨酸棒状杆菌ATCC 13032降解低浓度的4-甲基苯酚。
图2.用于过表达载体构建的质粒pMG36e。
图3.过表达载体pMG36e-creIH的构建;图3A为creI基因的扩增,图3B为creH基因的扩增,图3C为过表达载体pMG36e-creIH;
附图3A中的标记为:M,2000bp Marker,1,采用引物对F-creI-SacI与R-creI-SmaI扩增短乳杆菌D17基因组,2,采用引物对F-smal-RBS-creH与R-HindIII-creH扩增谷氨酸棒状杆菌基因组;图3B:M,2000bp Marker,1,采用引物对F-smal-RBS-creH与R-HindIII-creH扩增谷氨酸棒状杆菌基因组。
图4.过表达载体pMG36e-creIH转入短乳杆菌的PCR验证图;
附图4中的标记为:M:5000bp Marker;1,采用引物对F-creI-SacI与R-HindIII-creH扩增短乳杆菌D17基因组,2~4采用引物对F-creI-SacI与R-HindIII-creH扩增creI-creH过表达短乳杆菌菌株基因组。
图5.creI-creH短乳杆菌过表达菌株的过表达质粒稳定性。
图6.短乳杆菌D17/pMG36e与短乳杆菌D17/pMG36e-creIH过表达菌株对4-甲基苯酚的消减;
附图6中的标记为:D17/pMG36e为含有空表达载体的短乳杆菌空白对照菌;D17/pMG36e-creIH为creI-creH过表达短乳杆菌菌株。
具体实施方式
MRS培养基:葡萄糖2%,酵母浸粉0.4%,蛋白胨1%,牛肉膏0.5%,吐温80 0.1%,乙酸钠0.5%,磷酸氢二钾0.2%,柠檬酸三铵0.2%,硫酸镁0.02%,硫酸锰0.05%,均为质量-体积分数。调节pH为6.2。固体培养基中添加2%浓度的琼脂。
含红霉素的MRS平板:在MRS培养基基础上加入终浓度为4μg/mL的红霉素;
复苏液:葡萄糖2%,酵母浸粉0.4%,蛋白胨1%,牛肉膏0.5%,吐温80 0.1%,乙酸钠0.5%,磷酸氢二钾0.2%,柠檬酸三铵0.2%,硫酸镁0.02%,硫酸锰0.05%,蔗糖10.26%,均为质量-体积分数。
LB培养基(g/L):液体:酵母浸粉5,蛋白胨10,氯化钠10;固体培养基中添加2%浓度的琼脂。
含红霉素的LB培养基:在LB培养基基础上加入终浓度200μg/mL的红霉素。
GYP培养基:葡萄糖1%,酵母浸粉1%,蛋白胨0.5%,乙酸钠0.2%,硫酸镁0.02%,硫酸锰0.01%,硫酸亚铁0.01%,氯化钠0.01%,均为质量-体积分数。固体培养基中添加2%浓度的琼脂,发酵培养基中添加不同浓度谷氨酸钠。
含红霉素的GYP平板:在GYP培养基基础上加入终浓度为4μg/mL的红霉素;含红霉素的GYP培养基:在GYP培养基基础上加入终浓度1μg/mL的红霉素。
含4-甲基苯酚的样品前处理及SPEM-GC-MS检测4-甲基苯酚含量:
取去除菌体的发酵上清1mL,加入7mL超纯水及3g NaCl,10μL 200μg/L的3,4-二甲基苯酚内标液后置于20mL顶空瓶中,密封。对于酒醅样品的处理,称取5.0g酒醅于50ml离心管中,加入10%乙醇溶液5mL,涡旋混匀并浸泡30min,10000r/min离心5min,取上清液8mL置于20mL顶空瓶中,加入10μL 3,4-二甲基苯酚内标液。采用之前报道的SPEM-GC-MS检测方法分析4-甲基苯酚的含量。SPEM萃取条件:DVB/CAR/PBDS萃取头萃取45min,萃取温度为45℃。GC条件:进样口温度250℃,载气He,流速2mL/min,不分流进样,色谱柱为CP-Wax(60m×0.25m mi.d.×0.25μm,J&W Scientific)。检测时的升温程序为:50℃恒温2min,以6℃/min的速度升温至230℃,保持15min。MS条件:EI电离源,电子能量70eV,离子源温度230℃,扫描范围35.00~350amu。质谱分析用数据库来源于NIST05a.L(Agilent公司)。
实施例1:谷氨酸棒状杆菌降解低含量的4-甲基苯酚
取出-80℃冻存的谷氨酸棒状杆菌ATCC 13032菌株,接种于LB平板上,37℃培养12h。挑取平板上活化后的谷氨酸棒状杆菌单菌落接种至5mL LB培养基中,30℃,静置培养24h,再按10%(V/V)接种至5mL LB培养基中,30℃,静置培养12h,作为种子。种子按10%(V/V)接种至5mL LB培养基中,再加入2μL 4-甲基苯酚(终浓度约为7.3mg/L),37℃培养48h,同时不接菌的作为空白对照。
结果显示,如图1所示,在不添加谷氨酸棒状杆菌的空白对照组中,4-甲基苯酚的含量为初始添加值7.3mg/L,4-甲基苯酚未发生降解。而加入谷氨酸棒状杆菌的实验组中几乎无4-甲基苯酚残留,说明谷氨酸棒状杆菌对于低浓度的4-甲基苯酚有较好的降解作用。
实施例2:creIH基因过表达载体的构建
根据谷氨酸棒状杆菌的模式菌株ATCC13032中creI与creH基因的DNA序列,分别设计引物对F-creI-SacI与R-creI-SmaI,F-smal-RBS-creH与R-HindIII-creH。以谷氨酸棒状杆菌ATCC13032的基因组DNA为模板,分别用引物对F-creI-SacI与R-creI-SmaI和F-smal-RBS-creH与R-HindIII-creH扩增creI基因(序列如SEQ ID NO:1所示)(琼脂糖凝胶电泳结果如图3A所示)与creH基因(序列如SEQ ID NO:2所示)(琼脂糖凝胶电泳结果如图3B所示)。其中引物F-smal-RBS-creH上带有通过RBS Calculator V2.0网站设计的适用于乳杆菌的RBS(核糖体结合位点)(序列如SEQ ID NO:3所示)。PCR条件为98℃预变性30s;98℃变性10s,55℃退火15s,72℃延伸1~2min,30个循环;72℃后延伸10min。采用Omega胶回收试剂盒回收上下游同源臂各1000bp。取等摩尔creI、creH片段加水补足为5μL,再加入5μL的Takara的PrimeSTARMax DNA Polymerase,PCR条件为98℃预变性30s;98℃变性10s,55℃退火2min,72℃延伸2min,15个循环;72℃后延伸10min。以得到的PCR产物为模板,采用引物F-creI-SacI与R-HindIII-creH继续进行PCR。PCR条件为98℃预变性30s;98℃变性10s,55℃退火15s,72℃延伸3min,30个循环;72℃后延伸10min。采用重叠PCR方法得到creI与creH经RBS串联的creI-RBS-creH片段。
利用SacI和HindIII双酶切creI-RBS-creH片段和质粒pMG36e(如图2所示),之后将酶切产物于16℃连接16h后,转化至大肠杆菌Top10中。经菌液PCR及测序鉴定获得阳性菌株,进而得到将creI-creH基因克隆至pMG36e的过表达载体pMG36e-creIH(如图3C所示)。
实施例3:creIH基因过表达短乳杆菌菌株的构建
将构建好的过表达载体pMG36e-creIH电转化入短乳杆菌D17的感受态细胞,其转化条件为电压2000V~2500V电击5ms,将菌液涂布在含红霉素(终浓度为4μg/μL)的MRS平板上37℃培养,直至长出单克隆。挑选单克隆,用引物对F-creI-SacI与R-HindIII-creH进行PCR验证。PCR条件为98℃预变性30s;98℃变性10s,55℃退火15s,72℃延伸3min,30个循环;72℃后延伸10min。琼脂糖凝胶电泳结果如图4所示。挑选含有过表达载体的短乳杆菌菌株。
如图4A所示,采用引物对F-creI-SacI与R-HindIII-creH,利用野生菌基因组未扩增出任何PCR产物,而利用潜在的过表达菌株的基因组可扩增出约3000bp的PCR产物(为creIH全长),说明在挑选的短乳杆菌过表达菌株中均有过表达质粒,获得creIH过表达短乳杆菌菌株。
表1引物
实施例4:短乳杆菌creIH过表达菌株过表达载体的稳定性测试
在无菌条件下,将-80℃保存的短乳杆菌D17/pMG36e与短乳杆菌D17/pMG36e-creIH取出,分别划线于GYP固体平板上,37℃静置培养。
待单菌落长出后,从活化的GYP固体平板上挑取creIH过表达短乳杆菌菌株D17/pMG36e-creIH,接种于GYP液体培养基中,37℃静置培养24h,制成一级种子培养液。按10%的接种量,将一级种子液接种于新鲜的GYP培养基中,37℃×200rpm培养15h,制成二级种子培养液。将二级种子培养液按10%接种量,接种于GYP的发酵培养基中,37℃静置培养。每24h取样。取的样品用无菌的0.9%NaCl生理盐水按10-1~10-5梯度稀释,选择合适的梯度,取100μL稀释液分别涂于GYP平板与含有红霉素的GYP平板上,37℃静置培养48h,记录平板上的菌落数。质粒维持率为有抗平板上的菌落数与无抗性平板的菌落数的比值。
结果显示:如图5所示,短乳杆菌D17/pMG36e-creIH菌株,培养24h,质粒维持率为94.3%,培养48h,质粒维持率为91.3%,培养72h,质粒维持率为78.1%。培养48h,质粒维持率仍高于90%,说明在不添加红霉素抗性的条件下,发酵48h,creIH过表达短乳杆菌菌株的过表达载体仍保持稳定。
综上说明,不添加红霉素的条件下,质粒可保持稳定48h以上。在发酵食品中,可不添加红霉素,使用creIH过表达短乳杆菌用于4-甲基苯酚的消减。
实施例5:短乳杆菌creIH过表达菌株消减4-甲基苯酚的应用
挑取斜面上活化的空白对照菌短乳杆菌D17/pMG36e与creIH过表达短乳杆菌菌株D17/pMG36e-creIH单菌落,分别接种于含有1μg/L红霉素的GYP种子培养基中,37℃静置培养24h。取种子培养液,按10%的接种量接种于新的含有1μg/L红霉素的GYP种子培养基中,37℃静置培养12h后,培养液作为发酵种子。在250mL三角瓶中装入100mL的添加7.3mg/L 4-甲基苯酚的含有1μg/L红霉素的GYP发酵培养基,取培养12h的发酵种子按10%的接种量接种,于37℃静置培养48h。发酵上清液经SPEM-GC-MS检测4-甲基苯酚含量。
结果显示:如图6所示,空白对照菌短乳杆菌D17/pMG36e不消减4-甲基苯酚,creIH过表达短乳杆菌菌株D17/pMG36e-creIH消减4-甲基苯酚2130μg/L,且消减4-甲基苯酚的比率达30%。综上所述,creIH过表达短乳杆菌菌株可消减液体环境中的4-甲基苯酚。
实施例6:短乳杆菌creIH过表达菌株在酿造体系中消减4-甲基苯酚的应用
挑取斜面上活化的空白对照菌短乳杆菌D17/pMG36e与creIH过表达短乳杆菌菌株D17/pMG36e-creIH单菌落,分别接种于GYP液体培养基中,37℃静置培养24h,制成一级种子培养液。按10%的接种量,将一级种子液接种于新鲜的GYP培养基中,37℃×200rpm培养15h,制成二级种子培养液。将二级种子培养液按10%接种量,接种于GYP的培养基中,37℃×200rpm静置培养12h。9000g,4℃离心5min,获得的菌体用10%体积的生理盐水重悬。
向含有4-甲基苯酚的待蒸馏的酒醅中加入10%菌悬液(体积/重量),密封。于37℃静置培养48h。称取培养后的5.0g酒醅于50ml离心管中,加入10%乙醇溶液5ml,涡旋混匀并浸泡30min,10000r/min离心5min,取上清液8ml置于20ml顶空瓶中,加入10ul 3,4-二甲基苯酚内标液。采用SPEM-GC-MS检测方法分析酒醅中的4-甲基苯酚的含量。
结果如表2所示,creIH过表达短乳杆菌菌株D17/pMG36e-creIH可消减酿造体系中的4-甲基苯酚,消减能力为530μg/kg,消减率达37.9%。
表2短乳杆菌过表达菌株消减酿造体系中的4-甲基苯酚
SEQUENCE LISTING
<110> 江南大学
<120> 一种消减4-甲基苯酚的乳酸菌及其在酿造体系中的应用
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gcaccttctt ggggcccaat ctttggcaaa atcaaggcaa cagtcactga tattggtggc 1680
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Claims (10)
1.一种消减4-甲基苯酚短乳杆菌,其特征在于,所述消减4-甲基苯酚短乳杆菌是在短乳杆菌宿主中过表达4-甲基苯基磷酸酯合成酶的编码基因。
2.根据权利要求1所述的短乳杆菌,其特征在于,所述4为-甲基苯基磷酸酯合成酶的编码基因creI与creH基因;其中creI的氨基酸序列如SEQ ID NO:8所示,creH的氨基酸序列如SEQ ID NO:9所示。
3.根据权利要求1所述的短乳杆菌,其特征在于,所述宿主为短乳杆菌CGMCCNO.14385、短乳杆菌ATCC 367、短乳杆菌NCL912、短乳杆菌CGMCC1306、短乳杆菌145中的任意一种。
4.根据权利要求1-3任一所述的短乳杆菌,其特征在于,creI与creH基因是通过串联进行表达。
5.一种确定表达载体在权利要求1所述的短乳杆菌中的稳定性的方法,其特征在于,所述方法是在无抗性条件下,检测权利要求1所述的短乳杆菌的过表达质粒维持率。
6.一种消减4-甲基苯酚的方法,其特征在于,所述方法是利用creIH过表达短乳杆菌菌株进行发酵;所述creIH过表达短乳杆菌,过表达了4-甲基苯基磷酸酯合成酶的编码基因。
7.根据权利要求6所述的方法,其特征在于,所述4-甲基苯基磷酸酯合成酶的编码基因为creI与creH基因。可选地,编码所述creI基因的氨基酸序列如SEQ ID NO:8所示。可选地,编码所述creH基因的氨基酸序列如SEQ ID NO:9所示。
8.根据权利要求6-7任一所述的方法,其特征在于,所述方法是将creIH过表达短乳杆菌接种于含有4-甲基苯酚的系统中进行培养。可选地,所述含有4-甲基苯酚的系统可以是液态系统,也可以固态系统,比如:酒醅、奶酪、威士忌、白酒等。
9.一种在固态酿造体系消减4-甲基苯酚的方法,其特征在于,所述方法是利用creIH过表达短乳杆菌菌株在固态酿造体系中发酵;所述creIH过表达短乳杆菌,过表达了4-甲基苯基磷酸酯合成酶的编码基因。
10.根据权利要求9所述的方法,其特征在于,所述发酵是在固态酿造的出池酒醅中接种creIH过表达短乳杆菌菌株,静置培养一段时间。可选地,所述发酵是在模拟的固态酿造体系中进行。
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CN108315288A (zh) * | 2017-11-22 | 2018-07-24 | 华南理工大学 | 一种表达甲酰胺酶与亚磷酸脱氢酶融合蛋白的重组大肠杆菌及其构建方法与应用 |
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