CN113528366A - 一种产beta-丙氨酸酵母菌及其构建方法 - Google Patents
一种产beta-丙氨酸酵母菌及其构建方法 Download PDFInfo
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Abstract
本发明公开了一种产beta‑丙氨酸酵母菌,其为酿酒酵母BY4742衍生菌,在基因组上整合了外源的甲基丙二酰‑CoA变构酶基因、甲基丙二酰‑CoA脱羧酶基因、丙酸CoA‑转移酶基因、乳酰基‑CoA脱水酶alfa亚基基因、乳酰基‑CoA脱水酶beta亚基基因、硫酯裂解酶基因和氨基裂合酶基因,可通过一步发酵法实现β‑丙氨酸的从头合成。
Description
技术领域
本发明属于代谢工程和基因工程领域,具体地说,涉及一种产β-丙氨酸的酿酒酵母工程菌及其构建方法。
背景技术
β-丙氨酸(beta-丙氨酸)是自然界中存在的一种天然β型氨基酸。虽然不是蛋白质氨基酸,但其参与多种功能性物质如肌肽、维生素B5的合成,被广泛应用于医药、食品、化工等行业。例如可用于合成泛酸、泛酸钙、肌肽、帕米磷酸钠、八柳氮等,也可作为一种膳食补充剂为肌肉提供能量。由于β-丙氨酸及其衍生物在医药、美容、食品、饲料及化工等领域有广泛应用,因而市场需求量日渐上升。
β-丙氨酸制备方法有化学法、酶法及发酵法三种方法。其中,化学法和酶法是现在主要的生产方法。化学法包括丙烯腈法、丙烯酸法、β-氨基丙腈法、琥珀酰亚胺降解法等,但这些方法为高温、高压反应,工艺条件苛刻,且反应过程副产物较多,提取过程复杂,成本高,环境不友好,这些不可避免的缺点导致化学法合成β-丙氨酸在市场上的竞争力越来越弱。
酶法催化合成β-丙氨酸主要是L-天冬氨酸在L-天冬氨酸α脱羧酶(ADC)催化下脱羧生成β-丙氨酸,该方法催化效率高、条件温和、提取工艺简单环保,一直是研究的热点,但该方法的缺陷是酶催化底物天冬氨酸比较昂贵,导致生产成本较高,工业化受到很大限制。专利文献CN201710659654.9公开了生物酶法催化丙烯酸加氨合成β-丙氨酸的方法,但因为后处理步骤繁琐、生产成本较高因而无法工业化应用。发明人在前的研究CN201911182183.2中报道了以丙烯酸或丙烯腈为底物、用天冬氨酸氨裂合酶突变体AspB-11-D4催化生产β-丙氨酸,但在工业化生产方面尚需进一步完善。
发酵法采用廉洁易得的葡萄糖作为起始原料,近些年来逐渐成为新研究方向,专利文献CN201910474753.9公开了在大肠杆菌工程菌中实现了以葡萄糖为原料发酵产β-丙氨酸,产量高达50g/L,糖酸转化率达40%,但该方法还仅处于实验室研究阶段。此外,专利文献CN112662609A、CN112625985A、CN103898033A也公开了利用大肠杆菌工程菌发酵来生产β-丙氨酸,但也都仅限于实验室研究阶段,尚无法实现工业化应用。
现有技术的发酵法基本上都是采用大肠杆菌工程菌进行发酵,这存在一个致命缺陷。因为在公众看来,尤其是β-丙氨酸产品的普通消费者看来,大肠杆菌是一种致病菌,发酵产品会存在内毒素,这导致β-丙氨酸产品的用户难以接受,严重影响发酵产品的市场销售。
发明内容
为了探索采用公认的生物安全模式微生物比如谷氨酸棒杆菌、乳酸菌、枯草芽孢杆菌、面包酵母、酿酒酵母等作为菌种通过发酵来生产β-丙氨酸的可行性,我们对于β-丙氨酸在不同种类微生物中的可能代谢途径进行了推演和实验,并在某些微生物中取得了成功。例如,开发出了酿酒酵母BY4742(MATα his3Δ1 leu2Δ0 lys2Δ0 ura3Δ0)生物合成β-丙氨酸的代谢路线,从而使本发明构建的酿酒酵母工程菌能够实现β-丙氨酸的一步发酵生产。
因此,本发明的第一个目的在于提供一种产β-丙氨酸的酿酒酵母,其为酿酒酵母BY4742衍生菌,在基因组上整合了外源的编码甲基丙二酰-CoA变构酶的MMUT基因、编码甲基丙二酰-CoA脱羧酶的YgfG基因、编码丙酸CoA-转移酶(propionate CoA-transferase)的Pct基因、编码乳酰基-CoA脱水酶alfa亚基的lcdA基因、编码乳酰基-CoA脱水酶beta亚基的lcdB基因、编码硫酯裂解酶(tesB)的TesB基因和编码氨基裂合酶例如天冬氨酸氨裂合酶的AspB基因。
上述酿酒酵母BY4742的基因型可以为MATα his3Δ1 leu2Δ0 lys2Δ0 ura3Δ0即BY4742(MATα his3Δ1 leu2Δ0 lys2Δ0 ura3Δ0)。
优选地,上述甲基丙二酰-CoA变构酶来源于人Homo sapiens(Human),其氨基酸序列为SEQ ID NO:1;
上述甲基丙二酰-CoA脱羧酶来源于大肠杆菌Escherichia coli(strain K12),其氨基酸序列为SEQ ID NO:3;
上述丙酸CoA-转移酶来源于大肠杆菌Escherichia coli(strain K12),其氨基酸序列为SEQ ID NO:5;
上述乳酰基-CoA脱水酶alfa亚基来源于丙酸厌氧菌Anaerotignum propionicumDSM 1682,其氨基酸序列为SEQ ID NO:7;
上述乳酰基-CoA脱水酶beta亚基来源于丙酸厌氧菌Anaerotignum propionicumDSM 1682,其氨基酸序列为SEQ ID NO:9;
上述硫酯裂解酶来源于大肠杆菌Escherichia coli(strain K12),其氨基酸序列为SEQ ID NO:11;并且/或者
上述氨基裂合酶可以是来源于芽孢杆菌Bacillus sp.YM55-1的野生型天冬氨酸氨裂合酶(AspB,文献Ruifeng Li.et al,Computational redesign of enzymes forregio-and enantioselective hydroamination.Nat.Chem.Biol.,2018.中报道的序列)或者其(N142V,H188A)突变体,更优选所述(N142V,H188A)突变体,其氨基酸序列为SEQ IDNO:13,已在发明人在前的专利CN201911182183.2中公开,在该专利文献中是氨基酸序列为SEQ ID NO:3,编号是AspB-11-D4(相对应地,野生型酶编号为AspB1),在本文中继续沿用该编号。
在一种实施方式中,上述MMUT基因的核苷酸序列可以为SEQ ID NO:2;
上述YgfG基因的核苷酸序列可以为SEQ ID NO:4;
上述Pct基因的核苷酸序列为SEQ ID NO:6;
上述lcdA基因的核苷酸序列可以为SEQ ID NO:8;
上述lcdB基因的核苷酸序列可以为SEQ ID NO:10;
上述TesB基因的核苷酸序列为SEQ ID NO:12;并且/或者
上述AspB基因的核苷酸序列为SEQ ID NO:14。
本发明的第二个目的在于提供一种构建上述酿酒酵母的方法,其包括以下步骤:
A.构建MMUT基因表达盒,包括启动子pTEF1、MMUT基因、终止子ADH1t,形成pTEF1-MMUT-ADH1t表达盒;
构建YgfG基因表达盒,包括启动子pTPI1、ygfG基因、终止子PRM5t,形成pTPI1-ygfG-PRM5t表达盒;
构建Pct基因表达盒,包括启动子pTEF2、pct基因、终止子IDP1t,形成pTEF2-pct-IDP1t表达盒;
构建lcdA基因表达盒,包括启动子pENO2、lcdA基因、终止子CPS1t,形成pENO2-lcdA-CPS1t表达盒;
构建lcdB基因表达盒,包括启动子pGPM1、lcdB基因、终止子IDP1t,形成pGPM1-lcdB-IDP1t表达盒;
构建TesB基因表达盒,包括启动子pPGK3、tesB基因、终止子PRM9t,形成pPGK3-tesB-PRM9t表达盒;
构建AspB基因表达盒,包括启动子pTDH3、AspB-11-D4基因、终止子SPG5t,形成pTDH3-AspB-11-D4-SPG5t表达盒;
B.将上述步骤A构建的7种基因表达盒装载到适合于在酿酒酵母中表达的质粒载体上,形成基因簇表达质粒;
C.将步骤B得到的基因簇表达质粒转化入酿酒酵母BY4742,筛选基因组中整合了上述7种基因的阳性克隆;
D.筛选得到生产β-丙氨酸的酿酒酵母。
优选地,上述适合于在酿酒酵母中表达的质粒载体可以是pUC57等,但不限于此,所形成的基因簇表达质粒命名为pUC57-pathway质粒。该pUC57-pathway质粒全长序列为16192bp。
上述步骤C中的质粒转化方法选自化学转化(比如氯化钙转化)和电转化,优选电转化。
本发明的第三个目的在于提供上述的酿酒酵母在生产β-丙氨酸中的应用。
具体而言,通过上述酿酒酵母的发酵来生产β-丙氨酸,即通过“一步发酵法”实现β-丙氨酸的从头合成。
在发酵时,发酵培养基中可以添加有氨水或铵盐,所述铵盐选自硫酸铵、氯化铵、磷酸铵、碳酸铵、硝酸铵等。
在一种实施方式中,发酵用的种子培养基可以是YPD培养基,菌体发酵培养基是YPD-NH4培养基,组成为:10g/L酵母提取物,20g/L胰蛋白胨,20g/L葡萄糖,根据需要在发酵期间,例如发酵培养至45-48h,添加15%左右的氨水。
按照酿酒酵母的生长习性,发酵温度是30℃左右。
本发明构建的酿酒酵母工程菌是一种公认的安全模式微生物,能够通过发酵直接产生β-丙氨酸,所得到的发酵产物易于被广大用户接受,具有工业化开发利用价值。
附图说明
图1为本发明构建的酿酒酵母BY4742(MATα his3Δ1 leu2Δ0 lys2Δ0 ura3Δ0)菌株内生物合成β-丙氨酸的代谢路线图。
图2是本发明构建的基因簇表达质粒pUC57-pathway的结构示意图。
具体实施方式
本发明对酿酒酵母BY4742(MATα his3Δ1 leu2Δ0 lys2Δ0 ura3Δ0)菌株内在的代谢途径进行了改变,如图1所示,可以从葡萄糖出发,通过酶系催化实现β-丙氨酸的生物合成。
在该代谢途径中,酿酒酵母菌可以摄入葡萄糖作为碳源,代谢生成丙酮酸和琥珀酰-CoA,其中丙酮酸可在乳酸脱氢酶的催化下生成乳酸;琥珀酰-CoA先后经甲基丙二酰-CoA变构酶和甲基丙二酰-CoA脱羧酶催化生成丙酰-CoA;丙酰-CoA和乳酸经丙酸CoA转移酶催化发生转酰基反应,生成乳酰基-CoA;进一步经乳酰基-CoA脱水酶、硫酯裂解酶催化生成丙烯酸;丙烯酸可在无机铵存在的条件下,经氨基裂合酶催化生成β-丙氨酸。
在本文中,为了描述简便,有时会将某种酶比如硫酯裂解酶tesB与其编码基因(DNA)名称混用,本领域技术人员应能理解它们在不同描述场合表示不同的物质。本领域技术人员根据语境和上下文容易理解它们的含义。例如,对于TesB,用于描述硫酯裂解酶的功能或类别时,指的是蛋白质;在作为一种基因描述时,指的是编码该酶的基因。
应理解,本发明中整合入酿酒酵母基因组的MMUT基因、YgfG基因、Pct基因、lcdA基因、lcdB基因、TesB基因和AspB基因还可以分别单独地克隆在一个质粒上,然后分别地、或者同时地转入同一个酿酒酵母菌感受态细胞中;也可以将两个以上基因相组合地克隆在一个质粒上;也可以像图2所示的那样,将这7种基因形成基因簇克隆在一个质粒上,然后分别地、或者同时地转入同一个酿酒酵母菌感受态细胞中,经筛选阳性克隆,得到β-丙氨酸基因工程菌。所述质粒载体可以是任何适合于在酿酒酵母BY4742中表达的质粒载体例如pUC57等。
在本文中,对于本发明,术语“β-丙氨酸基因工程生产菌”、“β-丙氨酸基因工程菌”、“丙氨酸生产菌”都表示构建的酿酒酵母BY4742工程菌,尤其是酿酒酵母BY4742(MATαhis3Δ1 leu2Δ0 lys2Δ0 ura3Δ0)工程菌。
为了在酿酒酵母BY4742中表达中最佳地表达甲基丙二酰-CoA变构酶SEQ ID NO:1、甲基丙二酰-CoA脱羧酶SEQ ID NO:3、丙酸CoA-转移酶SEQ ID NO:5、乳酰基-CoA脱水酶alfa亚基SEQ ID NO:7、乳酰基-CoA脱水酶beta亚基SEQ ID NO:9、硫酯裂解酶SEQ ID NO:11和氨基裂合酶SEQ ID NO:13,本发明对它们的表达基因进行了密码子优化。
密码子优化是可用于通过增加感兴趣基因的翻译效率使生物体中蛋白质表达最大化的一种技术。不同的生物体由于突变倾向和天然选择而通常示出对于编码相同氨基酸的一些密码子之一的特殊偏好性。例如,在生长快速的微生物如大肠杆菌中,优化密码子反映出其各自的基因组tRNA库的组成。因此,在生长快速的微生物中,氨基酸的低频率密码子可以用用于相同氨基酸的但高频率的密码子置换。因此,优化的DNA序列的表达在快速生长的微生物中得以改良。
经过针对酿酒酵母进行密码子优化,上述7种酶SEQ ID NO:1、3、5、7、9、11、13的编码基因分别可以是SEQ ID NO:2、4、6、8、10、12、14。
以下结合具体实施例对本发明做进一步详细说明。应理解,以下实施例仅用于说明本发明而非用于限定本发明的范围。
本文中涉及到多种物质的添加量、含量及浓度,其中所述的百分含量,除特别说明外,皆指质量百分含量。
实施例
材料和方法
实施例中的全基因合成、引物合成及测序皆由苏州金唯智生物科技有限公司完成。
本文中的分子生物学实验包括质粒构建、酶切、感受态细胞制备、转化等主要参照《分子克隆实验指南》(第三版),J.萨姆布鲁克,D.W.拉塞尔(美)编著,黄培堂等译,科学出版社,北京,2002)进行。比如感受态细胞转化方法及感受态制备方法均参照第1章96页进行。必要时可以通过简单试验确定具体实验条件。
PCR扩增实验根据质粒或DNA模板供应商提供的反应条件或试剂盒说明书进行。必要时可以通过简单试验予以调整。
培养基:
LB培养基:5g/L酵母提取物,10g/L胰蛋白胨,10g/L氯化钠。(LB固体培养基另加20g/L琼脂粉。)
YPD培养基:10g/L酵母提取物,20g/L胰蛋白胨,20g/L葡萄糖。(固体培养基另加20g/L琼脂粉。)
YPD-NH4培养基:10g/L酵母提取物,20g/L胰蛋白胨,20g/L葡萄糖,根据需要在发酵培养至48h,添加15%的氨水。
以下实施例中,当使用含卡那霉素的培养基时,所述卡那霉素在培养基中的终浓度为50μg/ml。
20X电转母液:80g/L甘氨酸,2%吐温80。
丙烯酸HPLC检测方法:
检测仪器:安捷伦1260高效液相色谱仪,色谱柱:上海月旭OAA色谱柱,流动相A(100%):10mM磷酸二氢钾(0.68g到500ml水)磷酸调至PH2.5,流速1mL/min,柱温箱20℃,检测时间20min,采集波长210nm,进样量5ul。
β-丙氨酸HPLC检测方法:
检测仪器:安捷伦1260高效液相色谱仪,色谱柱:安捷伦SB-C18色谱柱或依利特BDS-C18柱,流动相A:2.871g无水乙酸钠+700ml水,流动相B:甲醇,梯度程序见下表,流速1.0mL/min,柱温箱40℃,检测波长334nm,进样量5ul,检测时间20min。
时间min | A% | B% |
0 | 70 | 30 |
6 | 70 | 30 |
7 | 55 | 45 |
15 | 55 | 45 |
15.5 | 70 | 30 |
20 | 70 | 30 |
衍生剂的配制:称取0.1372g邻苯二甲醛、0.0589g N-乙酰-L-半胱氨酸,加2ml无水乙醇,超声波震荡1min至溶解,加硼酸缓冲液至10ml(容量瓶定容)(P1F3位置)。
硼酸缓冲液配制:称取6.183g硼酸粉末,加250ml蒸馏水超声波震荡至溶解,然后用6mol NaOH溶液调pH至9.5,过滤,备用(P1F2位置)。P1F1位置是水针。衍生方法:1、使用默认补偿值以最大速度从位置P1F2上抽取5.00μl,在冲洗口清洗针3s。2、使用默认补偿值以最大速度从样品中抽取2.00μl。3、混合:将来自空气中的默认体积,以最大速度混合2次。4、等待0.1min,在冲洗口清洗针3s。5、使用默认补偿值以最大速度从位置P1F3上抽取1.00μl。6、混合:将来自空气中的默认体积,以最大速度混合6次。7、等待0.3min。8、在冲洗口清洗针3s。9、以最大速度,从针座上抽取32.00μl。10、将来自针座的默认体积,以最大速度混合2次。11、等待0.5min。12、进样。
实施例中所使用的出发菌株为酿酒酵母BY4742(MATα his3Δ1 leu2Δ0 lys2Δ0ura3Δ0),由上海工业生物技术研发中心惠赠。
实施例中所使用的质粒pUC57-pathway委托苏州金唯智生物科技有限公司合成,任何单位和个人都可以获得这些质粒用于验证本发明,但未经允许不得用作其他用途,包括开发利用、科学研究和教学。
实施例1:β-丙氨酸代谢工程菌的构建
1.1根据图2的设计,将7种基因搭配与酿酒酵母BY4742(MATαhis3Δ1leu2Δ0lys2Δ0ura3Δ0)适配的功能启动子和终止子,形成7个基因表达盒,其中:lcdA基因,使用启动子pENO2,终止子CPS1t,形成pENO2-lcdA-CPS1t;pct基因,使用启动子pTEF2,终止子IDP1t,形成pTEF2-pct-IDP1t;lcdB基因,使用启动子pGPM1,终止子IDP1t,形成pGPM1-lcdB-IDP1t;ygfG基因,使用启动子pTPI1,终止子PRM5t,形成pTPI1-ygfG-PRM5t;tesB基因,使用启动子pPGK3,终止子PRM9t,形成pPGK3-tesB-PRM9t;MMUT基因,使用启动子pTEF1,终止子ADH1t,形成pTEF1-MMUT-ADH1t;AspB-11-D4基因,使用启动子pTDH3,终止子SPG5t,形成pTDH3-AspB-11-D4-SPG5t。
将上述基因簇设计后,送苏州金唯智公司合成,装载到质粒载体pUC57上,形成pUC57-pathway质粒,全长序列为16192bp,如图2所示。
1.2将酿酒酵母BY4742(MATα his3Δ1 leu2Δ0 lys2Δ0 ura3Δ0)菌种在YPD固体培养基上划线,30℃培养2天,挑单菌落转接装有4ml YPD液体培养基的试管。30℃,220rpm过夜培养,转接装有25ml YPD液体培养基的250ml摇瓶,30℃,220rpm培养4~6h,培养至OD600为0.8~1.0,菌液用于制备酿酒酵母转化感受态。感受态的制作和转化,使用Frozen-EZ Yeast Transformation II Kit进行,严格参考其使用说明书。
利用电转化方法将质粒pUC57-pathway转入感受态细胞中。转化产物涂布SC-his平板(葡萄糖20g/l,YNB基本氮源1.7g/l,赖氨酸、亮氨酸、尿嘧啶各50mg/l),30℃培养4天。使用HIS3作为后续转化酿酒酵母宿主的筛选标记,阳性转化子可在缺少组氨酸的筛选平板上生长。
1.3筛选平板上,挑取阳性转化子,进行基因组抽提,以基因组为模板,分别使用下述两对验证引物Yz-1-F/Yz-1-R和Yz-2-F/Yz-2-R对转化子进行PCR验证,PCR反应体系配置严格参考High efficient&High fidelity PCR enzyme KOD FX(购自东洋纺公司)。PCR产物进行DNA凝胶电泳检测,两个PCR产物分别对应7935bp和6928bp大小。
验证引物:
Yz-1-F:ATGAATACTGATGTTAGAAT,
Yz-1-R:ATGTCTTATCAATATGTCAA。
Yz-2-F:TTAATGACCAACAAAATTTG,
Yz-2-R:TTATTCAGCTGCAGCCATAT。
实施例2:β-丙氨酸工程菌发酵验证
对实施例1中鉴定获得的基因型阳性转化子进行摇瓶发酵验证。将转化子分别接种YPD-NH4培养基进行发酵,30℃、220rpm培养3天。阳性转化子的β-丙氨酸最高产量为3g/L。相反,未进行β-丙氨酸代谢途径整合的宿主菌不产生丙烯酸和β-丙氨酸。
上述实施例表明,发明人针对酿酒酵母BY4742(MATα his3Δ1 leu2Δ0 lys2Δ0ura3Δ0)设计的β-丙氨酸代谢途径是成功的。整合了该代谢途径的菌株能够实现发酵液中β-丙氨酸的积累,具有工业化开发潜力。
应理解,上述实施例仅用于举例说明目的,而不是对本发明的限制。本领域技术人员在阅读了本发明的构思之后,对其作出的各种改变或调整,均应落入本发明的保护范围内,这些等价形式同样属于本文所附权利要求书限定的范围。
序列表
<110> 洛阳华荣生物技术有限公司
<120> 一种产beta-丙氨酸酵母菌及其构建方法
<130> SHPI2110208
<160> 14
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Phe Ser Thr Val Glu Glu Ser Asn Lys Phe Tyr Lys Asp Asn Ile Lys
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Ala Gly Gln Gln Gly Leu Ser Val Ala Phe Asp Leu Ala Thr His Arg
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Gly Tyr Asp Ser Asp Asn Pro Arg Val Arg Gly Asp Val Gly Met Ala
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Gly Val Ala Ile Asp Thr Val Glu Asp Thr Lys Ile Leu Phe Asp Gly
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Ile Pro Leu Glu Lys Met Ser Val Ser Met Thr Met Asn Gly Ala Val
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Ile Pro Val Leu Ala Asn Phe Ile Val Thr Gly Glu Glu Gln Gly Val
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Asn Ser Ile Ser Ile Ser Gly Tyr His Met Gln Glu Ala Gly Ala Asp
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Phe Gln Pro Lys Asn Ser Lys Ser Leu Leu Leu Arg Ala His Cys Gln
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Gly Ile Pro Lys Val Ala Asp Pro Trp Gly Gly Ser Tyr Met Met Glu
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Ile Glu Glu Met Gly Gly Met Ala Lys Ala Val Ala Glu Gly Ile Pro
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Lys Leu Arg Ile Glu Glu Cys Ala Ala Arg Arg Gln Ala Arg Ile Asp
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atgttgagag ctaaaaatca attgtttttg ttgtctccac attatttgag acaagttaaa 60
gaatcttctg gttctagatt gattcaacaa agattgttgc atcaacagca accattgcat 120
cctgaatggg ctgctttggc taaaaaacaa ttgaaaggta aaaatcctga agatttgatt 180
tggcatactc ctgaaggtat ttctattaaa ccattgtatt ctaaaagaga tactatggat 240
ttgcctgaag aattgcctgg tgttaaacca tttactagag gtccatatcc aactatgtat 300
acttttagac catggactat tagacaatat gctggatttt ctacagtcga ggaatctaac 360
aagttttata aagataatat taaagctggt caacaaggtt tgtctgttgc ttttgatttg 420
gctactcata gaggttatga ttctgataat ccaagagtta gaggtgatgt tggtatggct 480
ggtgttgcta ttgacactgt tgaagatact aaaattttgt ttgatggtat tccattggaa 540
aaaatgtctg tttctatgac tatgaatggt gctgttattc ctgttttggc taattttatt 600
gttactggtg aagaacaagg tgttccaaag gaaaaattga ctggaacaat tcaaaatgat 660
attttgaagg aattcatggt tagaaatact tatatttttc cacctgaacc ttctatgaaa 720
attattgctg atattttcga gtacactgct aaacacatgc caaagtttaa ttctatttct 780
atttctggtt atcatatgca agaagctggt gctgatgcta ttttggaatt ggcttatact 840
ttggctgatg gtttggaata ttctagaact ggtttgcaag ctggtttgac tattgatgaa 900
tttgctccaa gattgtcttt tttttggggt attggtatga atttttatat ggaaattgct 960
aaaatgagag ctggtagaag attgtgggct catttgattg aaaaaatgtt tcaaccaaaa 1020
aattctaaat ctttgttatt gagagctcat tgtcaaactt ctggttggtc tttgactgaa 1080
caagatccat ataataatat tgttagaact gctattgaag ctatggctgc tgtttttggt 1140
ggtactcaat ctttgcatac taattctttt gatgaagctt tgggtttgcc aacagtcaag 1200
tctgctagaa ttgctagaaa tactcaaatt atcattcaag aagaatctgg tattccaaaa 1260
gttgctgatc catggggtgg ttcttatatg atggaatgtt tgactaatga tgtttatgat 1320
gctgctttga aattgattaa tgaaattgaa gaaatgggtg gtatggctaa agctgttgct 1380
gaaggtatcc caaaattgag aattgaagag tgtgctgcta gaaggcaagc tagaattgat 1440
tctggttcag aagttattgt tggtgttaat aaataccaat tggaaaaaga agatgctgtt 1500
gaagttttgg ctatcgataa tacatctgtt agaaataggc aaattgagaa gttgaaaaaa 1560
atcaaatctt ctagagatca agctttggct gaaagatgtt tggctgcttt gactgaatgc 1620
gctgcttctg gtgatggtaa tattttggct ttggctgttg atgcttctag agctagatgt 1680
actgttggtg aaattactga tgctttgaaa aaagtttttg gtgaacataa agctaatgat 1740
agaatggttt ctggtgctta tagacaagaa tttggtgaat ctaaggaaat tacttctgct 1800
attaagagag ttcataaatt tatggaaaga gaaggtagaa gaccaagatt gttggttgct 1860
aaaatgggtc aagatggtca tgatagaggt gctaaagtta ttgctactgg ttttgctgat 1920
ttgggttttg atgttgatat tggtccattg tttcaaactc caagagaagt tgctcaacaa 1980
gctgtcgatg ctgatgttca tgctgttggt atttctactt tggctgctgg tcataaaact 2040
ttggttcctg aattgattaa agaattgaat tctttgggta gacctgatat tttggttatg 2100
tgtggtggtg ttattccacc acaagattat gaatttttgt ttgaagttgg tgtttctaat 2160
gtttttggtc ctggtactag aattccaaaa gctgctgttc aagttttgga tgatattgaa 2220
aaatgtttgg aaaaaaaaca acaatctgtt taa 2253
<210> 3
<211> 261
<212> PRT
<213> Escherichia coli (strain K12)
<400> 3
Met Ser Tyr Gln Tyr Val Asn Val Val Thr Ile Asn Lys Val Ala Val
1 5 10 15
Ile Glu Phe Asn Tyr Gly Arg Lys Leu Asn Ala Leu Ser Lys Val Phe
20 25 30
Ile Asp Asp Leu Met Gln Ala Leu Ser Asp Leu Asn Arg Pro Glu Ile
35 40 45
Arg Cys Ile Ile Leu Arg Ala Pro Ser Gly Ser Lys Val Phe Ser Ala
50 55 60
Gly His Asp Ile His Glu Leu Pro Ser Gly Gly Arg Asp Pro Leu Ser
65 70 75 80
Tyr Asp Asp Pro Leu Arg Gln Ile Thr Arg Met Ile Gln Lys Phe Pro
85 90 95
Lys Pro Ile Ile Ser Met Val Glu Gly Ser Val Trp Gly Gly Ala Phe
100 105 110
Glu Met Ile Met Ser Ser Asp Leu Ile Ile Ala Ala Ser Thr Ser Thr
115 120 125
Phe Ser Met Thr Pro Val Asn Leu Gly Val Pro Tyr Asn Leu Val Gly
130 135 140
Ile His Asn Leu Thr Arg Asp Ala Gly Phe His Ile Val Lys Glu Leu
145 150 155 160
Ile Phe Thr Ala Ser Pro Ile Thr Ala Gln Arg Ala Leu Ala Val Gly
165 170 175
Ile Leu Asn His Val Val Glu Val Glu Glu Leu Glu Asp Phe Thr Leu
180 185 190
Gln Met Ala His His Ile Ser Glu Lys Ala Pro Leu Ala Ile Ala Val
195 200 205
Ile Lys Glu Glu Leu Arg Val Leu Gly Glu Ala His Thr Met Asn Ser
210 215 220
Asp Glu Phe Glu Arg Ile Gln Gly Met Arg Arg Ala Val Tyr Asp Ser
225 230 235 240
Glu Asp Tyr Gln Glu Gly Met Asn Ala Phe Leu Glu Lys Arg Lys Pro
245 250 255
Asn Phe Val Gly His
260
<210> 4
<211> 786
<212> DNA
<213> 人工序列()
<400> 4
atgtcttatc aatatgtcaa tgtcgtcact attaataaag ttgctgttat tgaattcaat 60
tacggaagga aattgaatgc tttgtctaag gttttcattg atgatttgat gcaagctttg 120
tctgatttga atagacctga aattagatgt attattttga gagctccatc tggttctaaa 180
gttttttctg ctggtcatga tattcatgaa ttgccatctg gtggtagaga tccattgtct 240
tatgatgacc cattgagaca aattacaaga atgattcaaa agtttccaaa accaattatt 300
tctatggttg aaggttctgt ttggggtggt gcttttgaaa tgattatgtc ttctgatttg 360
attattgctg cttctacttc tactttttct atgactcctg ttaatttggg tgttccatat 420
aatttggttg gtattcataa tttgactaga gatgctggtt ttcatattgt taaagaattg 480
atttttactg cttctccaat tactgctcaa agagctttgg ctgttggtat tttgaatcat 540
gttgttgaag ttgaagaatt ggaagatttt actttgcaaa tggctcatca tatttctgaa 600
aaagctccat tggctattgc tgttattaaa gaagaattga gagttttggg tgaagctcat 660
actatgaatt ctgatgaatt tgaaagaatt caaggtatga gaagagctgt ttatgattct 720
gaagattatc aagaaggtat gaatgctttt ttggaaaaaa gaaaaccaaa ttttgttggt 780
cattaa 786
<210> 5
<211> 524
<212> PRT
<213> Escherichia coli (strain K12)
<400> 5
Met Arg Lys Val Pro Ile Ile Thr Ala Asp Glu Ala Ala Lys Leu Ile
1 5 10 15
Lys Asp Gly Asp Thr Val Thr Thr Ser Gly Phe Val Gly Asn Ala Ile
20 25 30
Pro Glu Ala Leu Asp Arg Ala Val Glu Lys Arg Phe Leu Glu Thr Gly
35 40 45
Glu Pro Lys Asn Ile Thr Tyr Val Tyr Cys Gly Ser Gln Gly Asn Arg
50 55 60
Asp Gly Arg Gly Ala Glu His Phe Ala His Glu Gly Leu Leu Lys Arg
65 70 75 80
Tyr Ile Ala Gly His Trp Ala Thr Val Pro Ala Leu Gly Lys Met Ala
85 90 95
Met Glu Asn Lys Met Glu Ala Tyr Asn Val Ser Gln Gly Ala Leu Cys
100 105 110
His Leu Phe Arg Asp Ile Ala Ser His Lys Pro Gly Val Phe Thr Lys
115 120 125
Val Gly Ile Gly Thr Phe Ile Asp Pro Arg Asn Gly Gly Gly Lys Val
130 135 140
Asn Asp Ile Thr Lys Glu Asp Ile Val Glu Leu Val Glu Ile Lys Gly
145 150 155 160
Gln Glu Tyr Leu Phe Tyr Pro Ala Phe Pro Ile His Val Ala Leu Ile
165 170 175
Arg Gly Thr Tyr Ala Asp Glu Ser Gly Asn Ile Thr Phe Glu Lys Glu
180 185 190
Val Ala Pro Leu Glu Gly Thr Ser Val Cys Gln Ala Val Lys Asn Ser
195 200 205
Gly Gly Ile Val Val Val Gln Val Glu Arg Val Val Lys Ala Gly Thr
210 215 220
Leu Asp Pro Arg His Val Lys Val Pro Gly Ile Tyr Val Asp Tyr Val
225 230 235 240
Val Val Ala Asp Pro Glu Asp His Gln Gln Ser Leu Asp Cys Glu Tyr
245 250 255
Asp Pro Ala Leu Ser Gly Glu His Arg Arg Pro Glu Val Val Gly Glu
260 265 270
Pro Leu Pro Leu Ser Ala Lys Lys Val Ile Gly Arg Arg Gly Ala Ile
275 280 285
Glu Leu Glu Lys Asp Val Ala Val Asn Leu Gly Val Gly Ala Pro Glu
290 295 300
Tyr Val Ala Ser Val Ala Asp Glu Glu Gly Ile Val Asp Phe Met Thr
305 310 315 320
Leu Thr Ala Glu Ser Gly Ala Ile Gly Gly Val Pro Ala Gly Gly Val
325 330 335
Arg Phe Gly Ala Ser Tyr Asn Ala Asp Ala Leu Ile Asp Gln Gly Tyr
340 345 350
Gln Phe Asp Tyr Tyr Asp Gly Gly Gly Leu Asp Leu Cys Tyr Leu Gly
355 360 365
Leu Ala Glu Cys Asp Glu Lys Gly Asn Ile Asn Val Ser Arg Phe Gly
370 375 380
Pro Arg Ile Ala Gly Cys Gly Gly Phe Ile Asn Ile Thr Gln Asn Thr
385 390 395 400
Pro Lys Val Phe Phe Cys Gly Thr Phe Thr Ala Gly Gly Leu Lys Val
405 410 415
Lys Ile Glu Asp Gly Lys Val Ile Ile Val Gln Glu Gly Lys Gln Lys
420 425 430
Lys Phe Leu Lys Ala Val Glu Gln Ile Thr Phe Asn Gly Asp Val Ala
435 440 445
Leu Ala Asn Lys Gln Gln Val Thr Tyr Ile Thr Glu Arg Cys Val Phe
450 455 460
Leu Leu Lys Glu Asp Gly Leu His Leu Ser Glu Ile Ala Pro Gly Ile
465 470 475 480
Asp Leu Gln Thr Gln Ile Leu Asp Val Met Asp Phe Ala Pro Ile Ile
485 490 495
Asp Arg Asp Ala Asn Gly Gln Ile Lys Leu Met Asp Ala Ala Leu Phe
500 505 510
Ala Glu Gly Leu Met Gly Leu Lys Glu Met Lys Ser
515 520
<210> 6
<211> 1575
<212> DNA
<213> 人工序列()
<400> 6
atgagaaaag ttccaattat tactgctgat gaagctgcta aattgattaa agatggtgat 60
actgttacta cttctggttt tgttggtaat gctattcctg aagctttgga tagagctgtt 120
gaaaaaagat ttttggaaac tggtgaacca aagaatatta catatgttta ttgtggttct 180
caaggtaata gagatggtag aggtgctgaa cattttgctc atgaaggttt gttgaaaaga 240
tatattgctg gtcattgggc tactgttcct gctttgggta aaatggctat ggaaaataaa 300
atggaagctt ataatgtttc tcaaggtgct ttgtgtcatt tgtttagaga tattgcttct 360
cataaacctg gtgtttttac taaagttggt attggtactt ttattgatcc aaggaatggt 420
ggaggtaaag ttaatgatat tactaaggag gatattgttg aattagttga aattaagggt 480
caagaatatt tgttttatcc tgcttttcca attcatgttg ctttgattag aggtacttat 540
gctgatgaat ctggtaatat tacttttgaa aaagaagttg ctccattgga aggtacttct 600
gtttgtcaag ctgttaaaaa ttctggtggt attgttgtcg ttcaagttga aagagttgtt 660
aaagctggta ctttggatcc aagacatgtt aaagttcctg gtatttatgt tgattatgtt 720
gtcgttgctg atcctgaaga tcatcaacaa tctttggatt gtgaatatga tcctgctttg 780
tctggtgaac atagaagacc tgaagttgtt ggtgaaccat tgccattgtc tgctaaaaaa 840
gttattggta gaagaggtgc tattgaattg gaaaaagatg ttgctgttaa tttgggtgtt 900
ggtgctcctg aatatgttgc ttctgttgct gatgaagaag gtattgttga ttttatgact 960
ttgactgctg aatctggtgc tattggtggt gttcctgctg gtggtgttag atttggtgct 1020
tcttataatg ctgatgcttt gattgatcaa ggttatcaat ttgattatta tgatggtgga 1080
ggtttggatt tgtgttattt gggtttggct gaatgtgatg aaaaaggtaa tattaatgtt 1140
tctagatttg gtccaagaat tgctggttgt ggtggtttta ttaatattac tcaaaatact 1200
ccaaaagttt ttttttgtgg tacttttact gctggtggat tgaaggttaa aatcgaagat 1260
ggaaaggtca ttatcgttca agaaggtaag cagaaaaaat tcttgaaagc tgttgaacaa 1320
attactttta atggtgatgt tgctttggct aataaacaac aagtcactta tattactgaa 1380
agatgtgttt ttttgttgaa agaagatggt ttgcatttgt ctgaaattgc tcctggtatt 1440
gatttgcaaa ctcaaatttt ggatgttatg gattttgctc caattattga tagagatgct 1500
aatggtcaaa ttaaattgat ggatgctgct ttgtttgctg aaggtttgat gggtttgaaa 1560
gaaatgaaat cttaa 1575
<210> 7
<211> 422
<212> PRT
<213> Anaerotignum propionicum DSM 1682
<400> 7
Met Ser Leu Thr Gln Gly Met Lys Ala Lys Gln Leu Leu Ala Tyr Phe
1 5 10 15
Gln Gly Lys Ala Asp Gln Asp Ala Arg Glu Ala Lys Ala Arg Gly Glu
20 25 30
Leu Val Cys Trp Ser Ala Ser Val Ala Pro Pro Glu Phe Cys Val Thr
35 40 45
Met Gly Ile Ala Met Ile Tyr Pro Glu Thr His Ala Ala Gly Ile Gly
50 55 60
Ala Arg Lys Gly Ala Met Asp Met Leu Glu Val Ala Asp Arg Lys Gly
65 70 75 80
Tyr Asn Val Asp Cys Cys Ser Tyr Gly Arg Val Asn Met Gly Tyr Met
85 90 95
Glu Cys Leu Lys Glu Ala Ala Ile Thr Gly Val Lys Pro Glu Val Leu
100 105 110
Val Asn Ser Pro Ala Ala Asp Val Pro Leu Pro Asp Leu Val Ile Thr
115 120 125
Cys Asn Asn Ile Cys Asn Thr Leu Leu Lys Trp Tyr Glu Asn Leu Ala
130 135 140
Ala Glu Leu Asp Ile Pro Cys Ile Val Ile Asp Val Pro Phe Asn His
145 150 155 160
Thr Met Pro Ile Pro Glu Tyr Ala Lys Ala Tyr Ile Ala Asp Gln Phe
165 170 175
Arg Asn Ala Ile Ser Gln Leu Glu Val Ile Cys Gly Arg Pro Phe Asp
180 185 190
Trp Lys Lys Phe Lys Glu Val Lys Asp Gln Thr Gln Arg Ser Val Tyr
195 200 205
His Trp Asn Arg Ile Ala Glu Met Ala Lys Tyr Lys Pro Ser Pro Leu
210 215 220
Asn Gly Phe Asp Leu Phe Asn Tyr Met Ala Leu Ile Val Ala Cys Arg
225 230 235 240
Ser Leu Asp Tyr Ala Glu Ile Thr Phe Lys Ala Phe Ala Asp Glu Leu
245 250 255
Glu Glu Asn Leu Lys Ala Gly Ile Tyr Ala Phe Lys Gly Ala Glu Lys
260 265 270
Thr Arg Phe Gln Trp Glu Gly Ile Ala Val Trp Pro His Leu Gly His
275 280 285
Thr Phe Lys Ser Met Lys Asn Leu Asn Ser Ile Met Thr Gly Thr Ala
290 295 300
Tyr Pro Ala Leu Trp Asp Leu His Tyr Asp Ala Asn Asp Glu Ser Met
305 310 315 320
His Ser Met Ala Glu Ala Tyr Thr Arg Ile Tyr Ile Asn Thr Cys Leu
325 330 335
Gln Asn Lys Val Glu Val Leu Leu Gly Ile Met Glu Lys Gly Gln Val
340 345 350
Asp Gly Thr Val Tyr His Leu Asn Arg Ser Cys Lys Leu Met Ser Phe
355 360 365
Leu Asn Val Glu Thr Ala Glu Ile Ile Lys Glu Lys Asn Gly Leu Pro
370 375 380
Tyr Val Ser Ile Asp Gly Asp Gln Thr Asp Pro Arg Val Phe Ser Pro
385 390 395 400
Ala Gln Phe Asp Thr Arg Val Gln Ala Leu Val Glu Met Met Glu Ala
405 410 415
Asn Met Ala Ala Ala Glu
420
<210> 8
<211> 1269
<212> DNA
<213> 人工序列()
<400> 8
atgtctttga ctcaaggtat gaaagctaaa caattgttgg cttattttca aggtaaagct 60
gatcaagatg ctagagaagc taaagctaga ggtgaattgg tttgttggtc tgcttctgtt 120
gctccacctg aattttgtgt tactatgggt attgctatga tttatcctga aactcatgct 180
gctggtattg gtgctagaaa aggtgctatg gatatgttgg aagttgctga tagaaaaggt 240
tataatgttg attgttgttc ttatggtaga gttaatatgg gttatatgga atgtttgaaa 300
gaagctgcta ttactggtgt taaacctgaa gttttggtta attctcctgc tgctgatgtt 360
cctttgcctg atttagtcat tacttgcaat aatatctgta atactttgtt gaaatggtat 420
gaaaatttgg ctgctgaatt ggatattcca tgtattgtta ttgatgttcc attcaatcat 480
actatgccaa tccctgaata tgctaaggct tatattgctg atcagtttag aaatgctatt 540
tctcagttgg aagttatttg tggtagacca ttcgattgga aaaaatttaa agaggtcaaa 600
gaccaaactc aaagatctgt ttatcattgg aatagaattg ctgaaatggc aaaatataaa 660
ccttctccat tgaatggttt tgatttgttt aattacatgg ctttgattgt tgcttgtaga 720
tctttggatt atgctgaaat tacttttaaa gcttttgctg atgaattgga agaaaatttg 780
aaagctggta tttatgcttt taaaggtgct gaaaaaacta gatttcaatg ggaaggtatt 840
gctgtttggc cacatttggg tcatactttt aaatctatga aaaatttgaa ttctattatg 900
actggtactg cttatcctgc tttgtgggat ttgcattatg atgctaatga tgaatcaatg 960
cactctatgg ctgaagctta cactaggatt tatattaata catgtttgca aaataaagtt 1020
gaagttttgt tgggtattat ggaaaaaggt caagttgacg gaactgttta tcacttgaac 1080
agatcttgca aattaatgtc tttcttaaat gttgaaactg cagaaattat taaggagaaa 1140
aatggtttgc catatgtttc tattgatggt gatcaaactg atccaagagt tttttctcct 1200
gctcaatttg atactagagt tcaagctttg gttgaaatga tggaagctaa tatggctgca 1260
gctgaataa 1269
<210> 9
<211> 374
<212> PRT
<213> Anaerotignum propionicum DSM 1682
<400> 9
Met Ser Arg Val Glu Ala Ile Leu Ser Gln Leu Lys Asp Val Ala Ala
1 5 10 15
Asn Pro Lys Lys Ala Met Asp Asp Tyr Lys Ala Glu Thr Gly Lys Gly
20 25 30
Ala Val Gly Ile Met Pro Ile Tyr Ser Pro Glu Glu Met Val His Ala
35 40 45
Ala Gly Tyr Leu Pro Met Gly Ile Trp Gly Ala Gln Gly Lys Thr Ile
50 55 60
Ser Lys Ala Arg Thr Tyr Leu Pro Ala Phe Ala Cys Ser Val Met Gln
65 70 75 80
Gln Val Met Glu Leu Gln Cys Glu Gly Ala Tyr Asp Asp Leu Ser Ala
85 90 95
Val Ile Phe Ser Val Pro Cys Asp Thr Leu Lys Cys Leu Ser Gln Lys
100 105 110
Trp Lys Gly Thr Ser Pro Val Ile Val Phe Thr His Pro Gln Asn Arg
115 120 125
Gly Leu Glu Ala Ala Asn Gln Phe Leu Val Thr Glu Tyr Glu Leu Val
130 135 140
Lys Ala Gln Leu Glu Ser Val Leu Gly Val Lys Ile Ser Asn Ala Ala
145 150 155 160
Leu Glu Asn Ser Ile Ala Ile Tyr Asn Glu Asn Arg Ala Val Met Arg
165 170 175
Glu Phe Val Lys Val Ala Ala Asp Tyr Pro Gln Val Ile Asp Ala Val
180 185 190
Ser Arg His Ala Val Phe Lys Ala Arg Gln Phe Met Leu Lys Glu Lys
195 200 205
His Thr Ala Leu Val Lys Glu Leu Ile Ala Glu Ile Lys Ala Thr Pro
210 215 220
Val Gln Pro Trp Asp Gly Lys Lys Val Val Val Thr Gly Ile Leu Leu
225 230 235 240
Glu Pro Asn Glu Leu Leu Asp Ile Phe Asn Glu Phe Lys Ile Ala Ile
245 250 255
Val Asp Asp Asp Leu Ala Gln Glu Ser Arg Gln Ile Arg Val Asp Val
260 265 270
Leu Asp Gly Glu Gly Gly Pro Leu Tyr Arg Met Ala Lys Ala Trp Gln
275 280 285
Gln Met Tyr Gly Cys Ser Leu Ala Thr Asp Thr Lys Lys Gly Arg Gly
290 295 300
Arg Met Leu Ile Asn Lys Thr Ile Gln Thr Gly Ala Asp Ala Ile Val
305 310 315 320
Val Ala Met Met Lys Phe Cys Asp Pro Glu Glu Trp Asp Tyr Pro Val
325 330 335
Met Tyr Arg Glu Phe Glu Glu Lys Gly Val Lys Ser Leu Met Ile Glu
340 345 350
Val Asp Gln Glu Val Ser Ser Phe Glu Gln Ile Lys Thr Arg Leu Gln
355 360 365
Ser Phe Val Glu Met Leu
370
<210> 10
<211> 1125
<212> DNA
<213> 人工序列()
<400> 10
atgtctagag ttgaagctat tttgtctcaa ttgaaagatg ttgctgctaa tccaaaaaaa 60
gctatggatg attataaagc tgaaactggt aaaggtgctg ttggtattat gccaatttat 120
tctcctgaag aaatggttca tgctgctggt tatttgccaa tgggtatttg gggtgctcaa 180
ggtaaaacta tttctaaagc tagaacttat ttgcctgctt ttgcttgttc tgttatgcaa 240
caagttatgg aattgcaatg tgaaggtgct tatgatgatt tgtctgctgt tattttttct 300
gttccatgtg atactttgaa atgtttgtct caaaaatgga aaggtacttc tcctgttatt 360
gtttttactc atccacaaaa tagaggtttg gaagctgcta atcaattttt ggttactgaa 420
tatgaattgg ttaaagctca attggaatct gtcttgggag ttaaaatctc taatgctgct 480
ttggagaatt caattgctat ctataatgaa aatagagctg ttatgagaga atttgttaaa 540
gttgctgctg attatccaca agttattgat gctgtttcta gacatgctgt ttttaaagct 600
agacaattta tgttgaaaga aaaacatact gctttggtta aagaattgat tgctgaaatt 660
aaagctactc ctgttcaacc atgggatggt aaaaaagttg ttgtcactgg tattttgtta 720
gaaccaaacg agttgttaga tatctttaat gaattcaaaa ttgctattgt tgatgacgat 780
ttggctcaag aatctagaca aattagagtt gatgttttgg atggtgaagg tggtccattg 840
tatagaatgg ctaaagcttg gcaacaaatg tatggttgtt ctttggctac tgatactaaa 900
aaaggtagag gtagaatgtt gattaataaa actattcaaa ctggtgctga tgctattgtt 960
gttgctatga tgaaattttg tgatcctgaa gaatgggatt atcctgttat gtatagagaa 1020
ttcgaagaga aaggagttaa atctttaatg attgaagttg accaagaagt ttcatctttt 1080
gagcagatta aaactagatt gcaatctttt gttgaaatgt tgtaa 1125
<210> 11
<211> 286
<212> PRT
<213> Escherichia coli (strain K12)
<400> 11
Met Ser Gln Ala Leu Lys Asn Leu Leu Thr Leu Leu Asn Leu Glu Lys
1 5 10 15
Ile Glu Glu Gly Leu Phe Arg Gly Gln Ser Glu Asp Leu Gly Leu Arg
20 25 30
Gln Val Phe Gly Gly Gln Val Val Gly Gln Ala Leu Tyr Ala Ala Lys
35 40 45
Glu Thr Val Pro Glu Glu Arg Leu Val His Ser Phe His Ser Tyr Phe
50 55 60
Leu Arg Pro Gly Asp Ser Lys Lys Pro Ile Ile Tyr Asp Val Glu Thr
65 70 75 80
Leu Arg Asp Gly Asn Ser Phe Ser Ala Arg Arg Val Ala Ala Ile Gln
85 90 95
Asn Gly Lys Pro Ile Phe Tyr Met Thr Ala Ser Phe Gln Ala Pro Glu
100 105 110
Ala Gly Phe Glu His Gln Lys Thr Met Pro Ser Ala Pro Ala Pro Asp
115 120 125
Gly Leu Pro Ser Glu Thr Gln Ile Ala Gln Ser Leu Ala His Leu Leu
130 135 140
Pro Pro Val Leu Lys Asp Lys Phe Ile Cys Asp Arg Pro Leu Glu Val
145 150 155 160
Arg Pro Val Glu Phe His Asn Pro Leu Lys Gly His Val Ala Glu Pro
165 170 175
His Arg Gln Val Trp Ile Arg Ala Asn Gly Ser Val Pro Asp Asp Leu
180 185 190
Arg Val His Gln Tyr Leu Leu Gly Tyr Ala Ser Asp Leu Asn Phe Leu
195 200 205
Pro Val Ala Leu Gln Pro His Gly Ile Gly Phe Leu Glu Pro Gly Ile
210 215 220
Gln Ile Ala Thr Ile Asp His Ser Met Trp Phe His Arg Pro Phe Asn
225 230 235 240
Leu Asn Glu Trp Leu Leu Tyr Ser Val Glu Ser Thr Ser Ala Ser Ser
245 250 255
Ala Arg Gly Phe Val Arg Gly Glu Phe Tyr Thr Gln Asp Gly Val Leu
260 265 270
Val Ala Ser Thr Val Gln Glu Gly Val Met Arg Asn His Asn
275 280 285
<210> 12
<211> 861
<212> DNA
<213> 人工序列()
<400> 12
atgtctcaag ctttgaaaaa tttgttgact ttgttgaatt tggaaaaaat tgaagaaggt 60
ttgtttagag gtcaatctga agatttgggt ttgagacaag tttttggtgg tcaagttgtt 120
ggtcaagctt tgtatgctgc taaagaaact gttcctgaag aaagattggt tcattctttt 180
cactcttatt ttttgagacc tggtgattct aaaaaaccaa ttatttatga cgttgaaact 240
ttgagagatg gtaattcttt ttctgctaga agagttgctg ctattcaaaa tggtaaacca 300
attttttata tgactgcttc ttttcaagct cctgaagctg gttttgaaca tcaaaaaact 360
atgccatctg ctcctgctcc tgatggtttg ccatctgaaa ctcaaattgc tcaatctttg 420
gctcatttgt tgccacctgt tttgaaagat aaatttattt gtgatagacc attggaagtt 480
agacctgttg aatttcataa tccattgaaa ggtcatgttg ctgaaccaca tagacaagtt 540
tggattagag ctaatggttc tgttcctgat gatttgagag ttcatcaata tttgttgggt 600
tatgcttctg atttgaattt tttgcctgtt gctttgcaac cacatggtat tggttttttg 660
gaacctggta ttcaaattgc tactattgat cattctatgt ggtttcatag accatttaat 720
ttgaatgaat ggttgttgta ttctgttgaa tctacttctg cttcttctgc tagaggtttt 780
gttagaggtg aattttatac tcaagatggt gttttggttg cttctactgt tcaagaaggt 840
gttatgagaa atcataatta a 861
<210> 13
<211> 468
<212> PRT
<213> 人工序列()
<400> 13
Met Asn Thr Asp Val Arg Ile Glu Lys Asp Phe Leu Gly Glu Lys Glu
1 5 10 15
Ile Pro Lys Asp Ala Tyr Tyr Gly Val Gln Thr Ile Arg Ala Thr Glu
20 25 30
Asn Phe Pro Ile Thr Gly Tyr Arg Ile His Pro Glu Leu Ile Lys Ser
35 40 45
Leu Gly Ile Val Lys Lys Ser Ala Ala Leu Ala Asn Met Glu Val Gly
50 55 60
Leu Leu Asp Lys Glu Val Gly Gln Tyr Ile Val Lys Ala Ala Asp Glu
65 70 75 80
Val Ile Glu Gly Lys Trp Asn Asp Gln Phe Ile Val Asp Pro Ile Gln
85 90 95
Gly Gly Ala Gly Thr Ser Ile Asn Met Asn Ala Asn Glu Val Ile Ala
100 105 110
Asn Arg Ala Leu Glu Leu Met Gly Glu Glu Lys Gly Asn Tyr Ser Lys
115 120 125
Ile Ser Pro Asn Ser His Val Asn Met Ser Gln Ser Thr Val Asp Ala
130 135 140
Phe Pro Thr Ala Thr His Ile Ala Val Leu Ser Leu Leu Asn Gln Leu
145 150 155 160
Ile Glu Thr Thr Lys Tyr Met Gln Gln Glu Phe Met Lys Lys Ala Asp
165 170 175
Glu Phe Ala Gly Val Ile Lys Met Gly Arg Cys Ala Leu Gln Asp Ala
180 185 190
Val Pro Ile Leu Leu Gly Gln Glu Phe Glu Ala Tyr Ala Arg Val Ile
195 200 205
Ala Arg Asp Ile Glu Arg Ile Ala Asn Thr Arg Asn Asn Leu Tyr Asp
210 215 220
Ile Asn Met Gly Ala Thr Ala Val Gly Thr Gly Leu Asn Ala Asp Pro
225 230 235 240
Glu Tyr Ile Ser Ile Val Thr Glu His Leu Ala Lys Phe Ser Gly His
245 250 255
Pro Leu Arg Ser Ala Gln His Leu Val Asp Ala Thr Gln Asn Thr Asp
260 265 270
Cys Tyr Thr Glu Val Ser Ser Ala Leu Lys Val Cys Met Ile Asn Met
275 280 285
Ser Lys Ile Ala Asn Asp Leu Arg Leu Met Ala Ser Gly Pro Arg Ala
290 295 300
Gly Leu Ser Glu Ile Val Leu Pro Ala Arg Gln Pro Gly Ser Ser Ile
305 310 315 320
Ile Pro Gly Leu Val Ala Pro Val Met Pro Glu Val Met Asn Gln Val
325 330 335
Ala Phe Gln Val Phe Gly Asn Asp Leu Thr Ile Thr Ser Ala Ser Glu
340 345 350
Ala Gly Gln Phe Glu Leu Asn Val Met Glu Pro Val Leu Phe Phe Asn
355 360 365
Leu Ile Gln Ser Ile Ser Ile Met Thr Asn Val Phe Lys Ser Phe Thr
370 375 380
Glu Asn Cys Leu Lys Gly Ile Lys Ala Asn Glu Glu Arg Met Lys Glu
385 390 395 400
Tyr Val Glu Lys Ser Ile Gly Ile Ile Thr Ala Ile Asn Pro His Val
405 410 415
Gly Tyr Glu Thr Ala Ala Lys Leu Ala Arg Glu Ala Tyr Leu Thr Gly
420 425 430
Glu Ser Ile Arg Glu Leu Cys Ile Lys Tyr Gly Val Leu Thr Glu Glu
435 440 445
Gln Leu Asn Glu Ile Leu Asn Pro Tyr Glu Met Thr His Pro Gly Ile
450 455 460
Ala Gly Arg Lys
465
<210> 14
<211> 1407
<212> DNA
<213> 人工序列()
<400> 14
atgaatactg atgttagaat tgaaaaagat tttttgggtg aaaaagaaat tccaaaagat 60
gcttattacg gtgttcaaac tattagagct actgagaatt ttccaattac tggttataga 120
attcatcctg aattgattaa atctttgggt attgttaaaa aatctgctgc tttggctaat 180
atggaagttg gtttgttgga taaagaagtt ggtcaatata ttgttaaagc tgctgatgag 240
gtcattgaag gtaaatggaa tgatcaattt attgttgatc caattcaagg tggtgctggt 300
acttctatta atatgaatgc taatgaagtt attgctaata gagctttgga attgatggga 360
gaagaaaaag gtaactattc taaaatctct ccaaattctc atgttaatat gtctcaatct 420
actgttgatg cttttccaac tgctactcat attgcagttt tatctttgtt aaatcaattg 480
attgaaacaa ctaaatatat gcagcaagaa tttatgaaaa aagctgatga atttgctggt 540
gttattaaaa tgggtagatg tgctttgcaa gatgctgttc caattttgtt gggtcaagaa 600
tttgaagctt atgctagggt tatcgctagg gatattgaaa gaattgcaaa tacaaggaat 660
aatttgtatg atattaatat gggtgctact gctgttggta ctggtttgaa tgctgatcct 720
gaatatattt ctattgttac tgaacatttg gctaaatttt ctggtcatcc attgagatct 780
gctcaacatt tggttgatgc tactcaaaat actgattgtt atactgaggt ctcttctgct 840
ttgaaagttt gtatgattaa catgtctaaa attgctaatg atttgagatt gatggcttct 900
ggtccaagag ctggtttgtc tgaaattgtt ttgcctgcta gacaacctgg ttcttctatt 960
attcctggtt tggttgctcc tgttatgcct gaagttatga atcaagttgc ttttcaagtt 1020
tttggtaatg atttgactat tacttctgct tctgaagctg gtcagttcga attgaacgtc 1080
atggagcctg ttttattttt caatttgatt cagtctatct caattatgac taacgttttc 1140
aagtctttta ctgaaaattg tttgaaggga attaaagcta atgaggaaag aatgaaggaa 1200
tatgtcgaaa agtctattgg tattatcact gctattaatc cacatgttgg ttatgaaact 1260
gctgctaaat tggctagaga agcttatttg actggtgaat ctattagaga attgtgtatt 1320
aagtatggtg tcttgactga agaacaattg aatgaaattt tgaatccata tgaaatgact 1380
catcctggta ttgctggtag aaaataa 1407
Claims (10)
1.一种产β-丙氨酸的酿酒酵母,其为酿酒酵母BY4742衍生菌,在基因组上整合了外源的编码甲基丙二酰-CoA变构酶的MMUT基因、编码甲基丙二酰-CoA脱羧酶的YgfG基因、编码丙酸CoA-转移酶的Pct基因、编码乳酰基-CoA脱水酶alfa亚基的lcdA基因、编码乳酰基-CoA脱水酶beta亚基的lcdB基因、编码硫酯裂解酶的TesB基因和编码氨基裂合酶的AspB基因。
2.如权利要求1所述的酿酒酵母,其特征在于,所述酿酒酵母BY4742的基因型为MATαhis3Δ1 leu2Δ0 lys2Δ0 ura3Δ0即BY4742(MATα his3Δ1 leu2Δ0 lys2Δ0 ura3Δ0)。
3.如权利要求1所述的酿酒酵母,其特征在于,
所述甲基丙二酰-CoA变构酶来源于人Homo sapiens(Human),其氨基酸序列为SEQ IDNO:1;
所述甲基丙二酰-CoA脱羧酶来源于大肠杆菌Escherichia coli(strain K12),其氨基酸序列为SEQ ID NO:3;
所述丙酸CoA-转移酶来源于大肠杆菌Escherichia coli(strain K12),其氨基酸序列为SEQ ID NO:5;
所述乳酰基-CoA脱水酶alfa亚基来源于丙酸厌氧菌Anaerotignum propionicum DSM1682,其氨基酸序列为SEQ ID NO:7;
所述乳酰基-CoA脱水酶beta亚基来源于丙酸厌氧菌Anaerotignum propionicum DSM1682,其氨基酸序列为SEQ ID NO:9;
所述硫酯裂解酶来源于大肠杆菌Escherichia coli(strain K12),其氨基酸序列为SEQ ID NO:11;并且/或者
所述氨基裂合酶氨基酸序列为SEQ ID NO:13。
4.如权利要求3所述的酿酒酵母,其特征在于,
所述MMUT基因的核苷酸序列为SEQ ID NO:2;
所述YgfG基因的核苷酸序列为SEQ ID NO:4;
所述Pct基因的核苷酸序列为SEQ ID NO:6;
所述lcdA基因的核苷酸序列为SEQ ID NO:8;
所述lcdB基因的核苷酸序列为SEQ ID NO:10;
所述TesB基因的核苷酸序列为SEQ ID NO:12;并且/或者
所述AspB基因的核苷酸序列为SEQ ID NO:14。
5.一种构建如权利要求1-4中任一项所述酿酒酵母的方法,其特征在于,包括以下步骤:
A.构建MMUT基因表达盒,包括启动子pTEF1、MMUT基因、终止子ADH1t,形成pTEF1-MMUT-ADH1t表达盒;
构建YgfG基因表达盒,包括启动子pTPI1、ygfG基因、终止子PRM5t,形成pTPI1-ygfG-PRM5t表达盒;
构建Pct基因表达盒,包括启动子pTEF2、pct基因、终止子IDP1t,形成pTEF2-pct-IDP1t表达盒;
构建lcdA基因表达盒,包括启动子pENO2、lcdA基因、终止子CPS1t,形成pENO2-lcdA-CPS1t表达盒;
构建lcdB基因表达盒,包括启动子pGPM1、lcdB基因、终止子IDP1t,形成pGPM1-lcdB-IDP1t表达盒;
构建TesB基因表达盒,包括启动子pPGK3、tesB基因、终止子PRM9t,形成pPGK3-tesB-PRM9t表达盒;
构建AspB基因表达盒,包括启动子pTDH3、AspB-11-D4基因、终止子SPG5t,形成pTDH3-AspB-11-D4-SPG5t表达盒;
B.将上述步骤A构建的7种基因表达盒装载到适合于在酿酒酵母中表达的质粒载体上,形成基因簇表达质粒;
C.将步骤B得到的基因簇表达质粒转化入酿酒酵母BY4742,筛选基因组中整合了上述7种基因的阳性克隆;
D.筛选得到生产β-丙氨酸的酿酒酵母。
6.如权利要求5所述的方法,其特征在于,所述适合于在酿酒酵母中表达的质粒载体是pUC57,所形成的基因簇表达质粒命名为pUC57-pathway质粒。
7.如权利要求1-4中任一项所述的酿酒酵母在生产β-丙氨酸中的应用。
8.如权利要求7所述的应用,其特征在于,通过所述酿酒酵母的发酵来生产β-丙氨酸。
9.如权利要求8所述的应用,其特征在于,发酵培养基中添加有氨水或铵盐。
10.如权利要求9所述的应用,其特征在于,发酵培养基是YPD-NH4培养基,组成为:10g/L酵母提取物,20g/L胰蛋白胨,20g/L葡萄糖,在发酵培养期间,添加15%的氨水。
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