CN110592153A - 生产o-乙酰基高丝氨酸的微生物和用其生产o-乙酰基高丝氨酸的方法 - Google Patents
生产o-乙酰基高丝氨酸的微生物和用其生产o-乙酰基高丝氨酸的方法 Download PDFInfo
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Abstract
公开了生产O‑乙酰基高丝氨酸的埃希氏菌属微生物,和利用该微生物高产率生产O‑乙酰基高丝氨酸的方法。
Description
本申请是分案申请,原申请的申请日为2015年6月22日,申请号为201580001286.8,发明名称为“生产O-乙酰基高丝氨酸的微生物和用其生产O-乙酰基高丝氨酸的方法”。
技术领域
本发明涉及生产O-乙酰基高丝氨酸的埃希氏菌属(Escherichia sp.)微生物,和用该微生物高产率生产O-乙酰基高丝氨酸的方法。
背景技术
O-乙酰基高丝氨酸是甲硫氨酸的前体,而甲硫氨酸是身体必需氨基酸之一。甲硫氨酸作为医用输注液的组分和医疗产品原料以及动物饲料和食物添加剂已被广泛使用。
甲硫氨酸可生物学或化学合成。近来,公开了两步法,其中将通过发酵生产的L-甲硫氨酸前体通过酶反应转化成L-甲硫氨酸(国际公开号WO 2008/013432)。在上述两步法中,O-琥珀酰基高丝氨酸和O-乙酰基高丝氨酸可用作甲硫氨酸前体,并且高产率生产O-乙酰基高丝氨酸以大规模成本有效地生产甲硫氨酸非常重要。
发明内容
技术问题
发明人在试图提高O-乙酰基高丝氨酸生产时发现柠檬酸合酶蛋白的表达或活性减少可显著增加O-乙酰基高丝氨酸的生产能力,从而完成本发明。
技术方案
本发明的一个目的是提供O-乙酰基高丝氨酸生产能力提高的O-乙酰基高丝氨酸生产微生物。
本发明的另一目的是提供利用该微生物生产O-乙酰基高丝氨酸的方法。
有利效果
使用根据本发明所述的具有O-乙酰基高丝氨酸生产能力的微生物可比化学合成以更高产率和更加环境友好的方式生产O-乙酰基高丝氨酸。另外,由此生产的O-乙酰基高丝氨酸可通过O-乙酰基高丝氨酸硫化氢解酶,用作甲硫氨酸和乙酸合成的前体,从而实现L-甲硫氨酸的生物转化,并且由此转化的L-甲硫氨酸可广泛用于生产人类食物或食物添加剂以及动物饲料或动物饲料添加剂。
附图说明
图1是用于构建柠檬酸合酶活性减弱的微生物的表达盒设计。
图2是pBAD24-柠檬酸合酶反义RNA(asRNA)载体的限制图谱。
具体实施方式
一方面,本发明提供生产O-乙酰基高丝氨酸的埃希氏菌属微生物,其中内源柠檬酸合酶蛋白的活性被减弱或灭活。
如本文所用,术语“O-乙酰基高丝氨酸”——其是微生物的甲硫氨酸生物合成途径中的具体中间材料——意指L-高丝氨酸的乙酰基衍生物。O-乙酰基高丝氨酸可利用高丝氨酸和乙酰辅酶A作为底物、通过使乙酰基从乙酰辅酶A转移至高丝氨酸的酶活性来生产。
如本文所用,术语“生产O-乙酰基高丝氨酸的微生物”包括这样的微生物:在活体生物内生产O-乙酰基高丝氨酸的真核或原核微生物,具有O-乙酰基高丝氨酸生产能力,而其亲本微生物不具有O-乙酰基高丝氨酸生产能力;或这样的微生物:内源地具有O-乙酰基高丝氨酸生产能力。
可通过物种的改良来提供或促进O-乙酰基高丝氨酸生产能力。具有O-乙酰基高丝氨酸生产能力的微生物可包括属于埃希氏菌属、欧文氏菌属(Erwinia sp.)、沙雷氏菌属(Serratia sp.)、普罗威登斯菌属(Providencia sp.)、棒状杆菌属(Corynebacteriasp.)、假单胞菌属(Pseudomonas sp.)、钩端螺旋体属(Leptospira sp.)、沙门氏菌属(Salmonella sp.)、短杆菌属(Brevibacteria sp.)、Hypomononas sp.、色杆菌属(Chromobacterium sp.)、和诺卡氏菌属(Norcardia sp.)、或真菌、或酵母的微生物;具体地,属于埃希氏菌属、棒状杆菌属、钩端螺旋体属、和酵母的微生物;和更具体地,属于埃希氏菌属的微生物,具体例如,大肠杆菌(Escherichia Coli)。具有O-乙酰基高丝氨酸生产能力的微生物可以是生产下列的微生物:L-赖氨酸、L-苏氨酸、L-异亮氨酸、或L-甲硫氨酸或其衍生物,但不限于此。
如本文所用,术语“柠檬酸合酶(E.C.2.3.3.1)”意指介导草酰乙酸盐(oxaloacetate)和乙酰辅酶A之间的反应的TCA循环第一步中的酶。具体地,柠檬酸合酶介导乙酰辅酶A中具有两个碳原子的乙酸残基和具有四个碳原子的草酰乙酸盐之间的缩合反应,从而生成具有六个碳原子的柠檬酸盐(citrate)。在大肠杆菌中,柠檬酸合酶被命名为GltA,柠檬酸合酶和GltA在本发明中可互换使用。
乙酰辅酶A+草酰乙酸盐+H2O→柠檬酸盐+辅酶A-SH
具体地,柠檬酸合酶可以是源自埃希氏菌属的柠檬酸合酶,更具体地源自大肠杆菌的GltA。柠檬酸合酶可以是这样的蛋白质:包括SEQ ID NO:4表示的氨基酸序列;或与SEQID NO:4的氨基酸序列具有70%或更高,具体地80%或更高、或更具体地90%或更高的同源性的氨基酸序列。另外,作为具有同源性的序列,如果该氨基酸序列是具有与SEQ ID NO:4相同或相应柠檬酸合酶活性的氨基酸序列,则在部分序列中具有敲除、修饰、替换、或添加的氨基酸序列显然也应被包括在本发明的范围内。另外,基于遗传密码简并,编码相同氨基酸序列和其变体的多核苷酸序列也应被包括本发明的范围内。
如本文所用,术语“内源”活性意指蛋白质在微生物中的天然状态或微生物中提供的相应蛋白质在修饰前的活性状态。
“蛋白质活性与其内源活性相比减弱或灭活(失活,inactivation)”意指蛋白质活性在与其天然状态具有的活性相比时降低或消除。减弱是表示如下情况的概念:由于蛋白质编码基因的修饰,蛋白质活性与微生物原本具有的蛋白质活性相比降低;整体蛋白质表达水平低于微生物自然型菌株的整体蛋白质表达水平;或其组合,但不限于此。灭活包括如下情况:编码蛋白质的基因与自然型菌株相比完全不被表达;和基因被表达,但不呈现活性。
蛋白质活性的减弱或灭活可通过本领域公知的各种方法实现。方法实例可包括用修饰基因替换染色体上编码蛋白质的基因从而可降低酶活性(其包括去除蛋白质活性的情况)的方法;在染色体上编码蛋白质的基因的表达调控序列上引入修饰的方法;用具有弱活性或不具有活性的序列替换编码蛋白质的基因的表达调控序列的方法;敲除染色体上编码蛋白质的基因的部分或整体的方法;引入反义寡核苷酸(例如,反义RNA)的方法,该反义寡核苷酸通过互补结合至染色体上的基因的转录物而抑制从mRNA至蛋白质的翻译;致使核糖体附着不能进行的方法——通过在编码蛋白质的基因的SD序列前端人为添加Shine-Dalgarno(SD)序列和其互补序列而形成二级结构;逆转录工程(RTE)方法——添加启动子以在相应序列的开放阅读框(ORF)的3’端逆转录;等,并且也包括其组合,但不限于此。
具体地,敲除编码蛋白质的基因的部分或整体的方法可通过如下进行:经由将染色体插入微生物的载体,用多核苷酸或(在多核苷酸序列部分被敲除时)标记物替换染色体中编码内源目标蛋白质的多核苷酸。例如,可采用通过同源重组进行基因敲除的方法,但不限于此。另外,如本文所用,术语“部分”,虽然可依据多核苷酸种类而变,但可具体意指1个核苷酸至300个核苷酸,更具体地1个核苷酸至100个核苷酸,还更具体地1个核苷酸至50个核苷酸,但不限于此。
另外,修饰表达调控序列的方法可通过如下进行:诱导多核苷酸序列的表达调控序列改变——通过敲除、插入、保守型替换、非保守型替换或其组合,以进一步减弱表达调控序列的活性;或用活性较弱的多核苷酸序列替换多核苷酸序列。多核苷酸序列可包括启动子、操纵子序列、编码核糖体结合域的序列、和调控转录和翻译终止的序列,但不限于此。
另外,修饰染色体上的基因序列的方法可通过如下进行:诱导该序列改变——通过敲除、插入、保守型替换、非保守型替换、或其组合,以进一步减弱表达调控序列的活性;或用具有较弱活性的改良基因序列或不具有活性的改良基因序列替换该序列,但不限于此。
具体地,关于柠檬酸合酶蛋白活性减弱,可用其他氨基酸(一个或多个)取代柠檬酸合酶蛋白的氨基酸序列中部分氨基酸(一个或多个)。更具体地,可包括具有如下氨基酸序列的柠檬酸合酶:柠檬酸合酶蛋白的氨基酸序列中的第145位氨基酸或第167位氨基酸由酪氨酸(Y)或赖氨酸(K)被取代成其他氨基酸(一个或多个)。还更具体地,柠檬酸合酶可以是具有编码如下修饰多肽的基因序列的柠檬酸合酶:其中柠檬酸合酶蛋白的氨基酸序列中第145位氨基酸由酪氨酸(Y)被取代成丙氨酸(A),并且第167位氨基酸由赖氨酸(K)被取代成丙氨酸(A)取代。具体地,在将起始密码子编码的甲硫氨酸的下一位氨基酸设置为第1位氨基酸后,按序列顺序确定氨基酸残基数量。多肽可分别具有SEQ ID NO:1或2表示的氨基酸序列。另外,如果柠檬酸合酶的活性弱于野生型的活性,则柠檬酸合酶可包括与SEQ IDNO:1或2的氨基酸序列具有80%或更高,具体地90%或更高,更具体地95%或更高,还更具体地97%或更高同源性的氨基酸序列。作为具有同源性的序列,如果该氨基酸序列是与SEQID NO:1或2的蛋白质具有基本上相同或相应的生物学活性的氨基酸序列,则在部分序列具有敲除、修饰、替换、或添加的氨基酸序列显然也应被包括在本发明的范围内。
如本文所用,术语“同源性”意指两个多核苷酸或多肽部分之间的同一性百分比。一部分与另一部分的序列之间的同源性可通过本领域已知的技术确定。例如,同源性可通过如下确定:利用计算机程序排列,在两个不同多核苷酸分子或两个不同多肽之间直接排列序列信息和容易地获得序列信息。计算机程序可包括BLAST(NCBI)、CLC Main Workbench(CLC bio)、MegAlignTM(DNASTAR Inc)等。另外,多核苷酸之间的同源性可通过如下确定:在同源区之间形成稳定双链的条件下杂交多核苷酸,利用单链特异性核酸酶进行分解,和确定分解片段。
如本文所用,术语“同源性”意指具有“共同进化起源”的蛋白质之间的关系,包括源自超家族蛋白质的所有语法形式或拼写变型的同源蛋白质,和源自不同物种的同源蛋白质。这些蛋白质(及其编码基因)具有通过高水平序列相似性反映的序列同源性。然而,普遍使用和本发明使用的术语“同源性”意指被诸如“非常高”的形容词修饰的序列相似性,而非意指共同的进化起源。
在本发明的示例性实施方式中,微生物可以是胱硫醚γ合酶(EC 2.5.1.48)、高丝氨酸激酶(EC 2.7.1.39)、或二者的活性弱于其内源活性或灭活的微生物。
如本文所用,术语“胱硫醚γ合酶”意指可利用O-琥珀酰基高丝氨酸和L-半胱氨酸作为底物通过下述化学反应合成胱硫醚的酶。在本发明中,来自大肠杆菌(E.coli)的胱硫醚γ合酶被命名为“MetB”。
O-琥珀酰基-L-高丝氨酸+L-半胱氨酸→L-胱硫醚+琥珀酸盐(succinate)
具体地,来自大肠杆菌的胱硫醚γ合酶——虽然并非具体限于此——可以是包括SEQ ID NO:9表示的氨基酸序列或与SEQ ID NO:9的氨基酸序列具有70%或更高,具体地80%或更高,更具体地90%或更高同源性的氨基酸序列的蛋白质。另外,作为具有同源性的序列,如果该氨基酸序列是与SEQ ID NO:9的氨基酸序列具有相同或相应高丝氨酸激酶活性的氨基酸序列,则在部分序列具有敲除、修饰、替换、或添加的氨基酸序列显然也应被包括在本发明的范围内。另外,基于遗传密码简并,编码相同氨基酸序列和其变体的多核苷酸序列也应被包括在本发明的范围内。
减弱和灭活胱硫醚γ合酶活性的方法可按照上述方法进行。
如本文所用,术语“高丝氨酸激酶”意指致使高丝氨酸磷酸化的酶,其进行下述化学反应。在本发明中,来自大肠杆菌的高丝氨酸激酶被命名为“ThrB”。
ATP+L-高丝氨酸→ADP+O-磷酸-L-高丝氨酸
具体地,来自埃希氏菌属的高丝氨酸激酶——虽然并非具体限于此——可以是包括SEQ ID NO:11表示的氨基酸序列或与SEQ ID NO:11的氨基酸序列具有70%或更高、具体地80%或更高、或更具体地90%或更高同源性的氨基酸序列的蛋白质。另外,作为具有同源性的序列,如果该氨基酸序列是与SEQ ID NO:11的氨基酸序列具有相同或相应高丝氨酸激酶活性的氨基酸序列,则在部分序列具有敲除、修饰、替换、或添加的氨基酸序列显然也应被包括在本发明的范围内。另外,基于遗传密码简并,编码相同氨基酸序列和其变体的多核苷酸序列也应被包括在本发明的范围内。
减弱和灭活高丝氨酸激酶活性的方法可按照上述方法进行。
在本发明的具体方面,微生物可以是这样的微生物:其中高丝氨酸O-乙酰基转移酶的活性被进一步引入或增强,或内源高丝氨酸O-琥珀酰基转移酶被进一步修饰以具有高丝氨酸O-乙酰基转移酶的活性。
如本文所用,术语“高丝氨酸O-乙酰基转移酶(EC 2.3.1.31)”意指具有将乙酰基从乙酰辅酶A转移至高丝氨酸的活性的酶。
具体地,可将高丝氨酸O-乙酰基转移酶活性引入根据本发明的微生物。高丝氨酸O-乙酰基转移酶可源自各种微生物物种,例如,选自下列的微生物:棒状杆菌属、钩端螺旋体属、奇异球菌属(Deinococcus sp.)、奇异球菌属、假单胞菌属、和分支杆菌属(Mycobacterium sp.)。具体地,高丝氨酸O-乙酰基转移酶可以是包括下列表示的氨基酸序列的高丝氨酸O-乙酰基转移酶:SEQ ID NO:13(迈氏钩端螺旋体(Leptospira meyeri))、SEQ ID NO:14(谷氨酸棒状杆菌(Corynebacterium glutamicum))、或SEQ ID NO:15(抗辐射奇异球菌(Deinococcus radiodurans)),但不限于此。另外,高丝氨酸O-乙酰基转移酶可以是包括上述氨基酸序列或与上述氨基酸序列具有70%或更高、具体地80%或更高、或更具体地90%或更高同源性的氨基酸序列的蛋白质。另外,基于遗传密码简并,编码相同氨基酸序列和其变体的多核苷酸序列也应被包括在本发明的范围内。
本发明所用的高丝氨酸O-乙酰基转移酶的实例被公开于韩国专利申请公开号10-2011-0023703和欧洲专利申请公开号EP 2290051,这些专利文件的完整说明书可被本发明包括作为参考。
另外,其中内源高丝氨酸O-琥珀酰基转移酶(EC 2.3.1.46)经修饰具有高丝氨酸O-乙酰基转移酶活性的蛋白质意指这样的多肽:其中具有高丝氨酸O-琥珀酰基转移酶活性的多肽的底物特异性从琥珀酰辅酶A变成乙酰辅酶A。另外,修饰蛋白质——虽然并非具体限于此——可以是通过替换具有高丝氨酸O-琥珀酰基转移酶活性的多肽的部分氨基酸序列而不同于其野生型的具有高丝氨酸O-乙酰基转移酶活性的肽。
高丝氨酸O-琥珀酰基转移酶的实例可以是来自肠细菌属(Enterobacteria sp.)、沙门氏菌属、假单胞菌属、杆菌属(Bacillus sp.)、或埃希氏菌属的多肽,具体地、来自埃希氏菌属的具有高丝氨酸O-琥珀酰基转移酶活性的多肽,例如,来自大肠杆菌的具有高丝氨酸O-琥珀酰基转移酶活性的多肽。更具体地,来自大肠杆菌的高丝氨酸O-琥珀酰基转移酶可具有SEQ ID NO:16表示的氨基酸序列,但不限于此。来自大肠杆菌的高丝氨酸O-琥珀酰基转移酶被命名为“MetA”。
修饰型高丝氨酸O-琥珀酰基转移酶可以是变体多肽,其中SEQ ID NO:16表示的多肽或与SEQ ID NO:16的多核苷酸序列具有95%或以上同源性的多肽的第111位氨基酸被谷氨酸取代,另外,第112位氨基酸被苏氨酸或组氨酸取代。具体地,变体多肽可以是具有SEQID NOS:17至19中任一个的氨基酸序列的多肽。另外,变体多肽可以是包括与上述氨基酸序列具有70%或更高、具体地80%或更高、或更具体地90%或更高同源性的氨基酸序列的蛋白质。另外,基于遗传密码简并,编码相同氨基酸序列和其变体的多核苷酸序列也应被包括在本发明的范围内。关于修饰型高丝氨酸O-琥珀酰基转移酶的信息可从韩国专利申请公开号10-2012-0070531或国际公开号WO2012/087039获得,这些专利文件的完整说明书被本发明包括作为参考。
如本文所用,术语“引入或增强活性”意指提供具体蛋白质的活性给不具有该蛋白质的微生物;或在具有该蛋白质的微生物内增强该蛋白质的胞内活性,和类似情况,并且意指与该蛋白质的内源活性相比增加该蛋白质的胞内活性。
如本文所用,术语“引入或增强蛋白质活性”不仅意指由于增加蛋白质自身活性而导致实现效果高于原始功能,而且意指由于内源基因活性增加、内部或外部因素造成的内源基因扩增、拷贝数增加、从外界引入基因、因替换、修饰或突变造成的酶活性增加等导致的蛋白质活性增加,但不限于此。
在上文中,基因拷贝数增加——虽然并非具体限于此——可在可操作地连接至载体的状态下进行、或通过插入宿主细胞中的染色体进行。具体地,该方法可通过如下进行:将可操作地连接编码本发明蛋白质的多核苷酸并且可独立于宿主进行复制和发挥功能的载体引入宿主细胞;或将可操作地连接多核苷酸、使该多核苷酸插入宿主细胞染色体的载体引入宿主细胞,从而增加在宿主细胞染色体内的基因拷贝数。
载体是这样的DNA构建物:包括编码目标蛋白质的多核苷酸的多核苷酸序列,其可操作地连接至适当的调控序列,以使目标蛋白质可在合适的宿主中表达,其中调控序列包括启动转录的启动子、调控转录的无规操纵子序列、编码适当的mRNA核糖体结合域的序列、和调控转录和翻译的序列。载体在转化到适当的宿主细胞后可独立于宿主基因组进行复制或发挥功能,或可被整合到宿主基因组自身中。
本发明使用的载体可不被具体限制,只要该载体在宿主细胞中可复制,并且可使用本领域已知的任何载体。载体的实例可包括天然或重组质粒、粘粒、病毒、和噬菌体。例如,作为噬菌体载体或粘粒载体,可使用pWE15、M13、λMBL3、λMBL4、λIXII、λASHII、λAPII、λt10、λt11、Charon4A、Charon21A、等;和作为质粒载体,可使用pBR系、pUC系、pBluescriptII系、pGEM系、pTZ系、pCL系、pET系等。具体地,可使用pDZ、pACYC177、pACYC184、pCL、pECCG117、pUC19、pBR322、pMW118、pCC1BAC载体等。
另外,可利用插入微生物染色体的载体用修饰多核苷酸取代编码内源目标蛋白质的多核苷酸。多核苷酸插入染色体可利用本领域已知的方法进行,例如,通过同源重组。由于本发明的载体可通过同源重组插入染色体,可另外包括用于证实插入染色体的选择标记物。利用选择标记物来选择被转化的细胞,即,证实目标多核苷酸是否已被插入,并且可使用提供可选表型如抗药性、营养需求、细胞毒剂抗性、和表面蛋白质表达的标记物,但不限于此。在选择剂处理的情况下,只有表达选择标记物的细胞可存活或表达其他表型特征,因此可容易选择被转化的细胞。
如本文所用,术语“转化”意指将包括编码目标蛋白质的多核苷酸的载体引入宿主细胞,从而实现编码蛋白质的多核苷酸在宿主细胞中表达的过程。关于转化的多核苷酸,其是被插入宿主细胞的染色体并位于其中还是位于染色体外无关紧要,只要其可在宿主细胞中表达即可。另外,多核苷酸包括编码目标蛋白质的DNA和RNA。多核苷酸可以任何方式插入,只要其可被引入宿主细胞并且在其中表达。例如,多核苷酸可以表达盒形式被引入宿主细胞,表达盒是包括自我表达所需的所有必需要素的基因构建物。表达盒可常规地包括可操作地连接至多核苷酸的启动子、转录终止信号、核糖体结合域、和翻译终止信号,并且可以是能够自我复制的表达载体形式。另外,多核苷酸可被原样引入宿主细胞,并且可操作地连接至其在宿主细胞中表达所必需的序列。另外,如本文所用,术语“可操作地连接”意指启动子序列(启动和介导编码目标蛋白质的多核苷酸的转录)和基因序列之间的功能性连接。
然后,可通过如下进行表达调控序列的修饰——虽然并非具体限于此——以增加多核苷酸表达:经由敲除、插入、保守型替换、非保守型替换、或其组合诱导多核苷酸序列改变,以进一步增强表达调控序列的活性;或用活性较强的多核苷酸序列替换多核苷酸序列。表达调控序列——虽然并非具体限于此——可包括启动子、操纵子序列、编码核糖体结合域的序列、和调控转录和翻译终止的序列等。另外,强外源启动子,代替原启动子,可被连接至多核苷酸表达单元的上端。
此外,可通过如下进行染色体上多核苷酸序列的修饰——虽然并非具体限于此:经由敲除、插入、保守型替换、非保守型替换、或其组合诱导多核苷酸序列的表达调控序列改变,以进一步增强多核苷酸序列的活性;或用活性较强的增强型多核苷酸序列替换多核苷酸序列。
总体上,蛋白质活性的引入和增强可使相应蛋白质的活性或强度相对于野生型蛋白质或微生物中的蛋白质的活性或强度增加至少1%、10%、25%、50%、75%、100%、150%、200%、300%、400%、或500%,至多1000%或2000%。
另外,微生物可以是这样的微生物:其中内源高丝氨酸O-琥珀酰基转移酶的活性与内源活性相比减弱或灭活,从而通过阻断由高丝氨酸生物合成O-琥珀酰基高丝氨酸的途径来增强O-乙酰基高丝氨酸的生物合成途径。
高丝氨酸O-琥珀酰基转移酶活性的减弱和灭活可按照上述方法进行。
在本发明的示例性实施方式中,O-乙酰基高丝氨酸生产微生物可这样的微生物:其中从磷酸烯醇丙酮酸盐(phosphoenolpyruvate)至高丝氨酸的生物合成途径所涉及的酶的活性被另外引入或增强,以进一步增加高丝氨酸(O-乙酰基高丝氨酸生物合成的底物)的量。
具体地,上述微生物可以是这样的微生物:其中选自磷酸烯醇丙酮酸羧化酶(ppc,EC 4.1.1.31)、天冬氨酸氨基转移酶(aspC,EC 2.6.1.1)、和天冬氨酸半醛脱氢酶(asd,EC1.2.1.11)的至少一种蛋白质的活性被进一步引入或增强。
例如,编码包括SEQ ID NO:20表示的氨基酸序列的羧酸磷酸烯醇丙酮酸的ppc基因、编码包括SEQ ID NO:21表示的氨基酸序列的天冬氨酸氨基转移酶的aspC基因、和编码包括SEQ ID NO:22表示的氨基酸的天冬氨酸半醛脱氢酶的asd基因可被引入微生物。例如,这三种不同酶的活性可通过如下被引入和增强:使存在于宿主细胞染色体中的编码这三种不同酶的基因的拷贝数全部至少为2,但不限于此。活性的引入和增强可按照上述方法进行。
在本发明的示例性实施方式中,通过各种方法减弱或灭活柠檬酸合酶蛋白的活性,包括敲除生产O-乙酰基高丝氨酸的大肠杆菌微生物中的柠檬酸合酶基因;将编码与野生型相比活性减弱的修饰柠檬酸合酶蛋白的基因引入柠檬酸合酶基因的位置;和引入柠檬酸合酶基因反义RNA的表达载体。结果是,由此构建的柠檬酸合酶蛋白活性减弱或灭活的O-乙酰基高丝氨酸生产微生物,与亲本微生物相比,显示出提高的O-乙酰基高丝氨酸生产能力(实施例1至4)。
另一方面,本发明提供利用O-乙酰基高丝氨酸生产能力提高的O-乙酰基高丝氨酸生产微生物生产O-乙酰基高丝氨酸的方法。具体地,本发明提供生产O-乙酰基高丝氨酸的方法,包括(a)培养微生物;和(b)回收在微生物培养期间生产的O-乙酰基高丝氨酸。
根据本发明所述的具有O-乙酰基高丝氨酸生产能力的大肠杆菌的培养方法可按照本领域已知的适当培养基和培养条件进行。本领域技术人员可容易根据所选微生物调整培养过程。具体地,由于本发明的微生物是柠檬酸合酶(介导TCA循环第一步的酶)的活性减弱或灭活的微生物,培养基可包括谷氨酸盐,但不具体限于此。
培养方法的实例可包括分批培养、连续培养、和补料分批培养,但不限于此。这些不同的方法例如被公开于"Biochemical Engineering",James M.Lee,Prentice-HallInternational Editions,第138-176页中。
培养中使用的培养基可适当地满足具体微生物的要求。具体地,微生物培养基的实例被公开于"Manual of Methods for General Bacteriology",American Society forBacteriology,Washington,DC,1981。培养基可以是包括合适的碳源、磷源、无机化合物、氨基酸、和/或维生素等的培养基,并且培养可在有氧条件下同时调节温度、pH等进行。
碳源的实例可包括碳水化合物,如葡萄糖、乳糖、蔗糖、乳酸、果糖、麦芽糖、淀粉、和纤维素;脂肪,如大豆油、葵花籽油、蓖麻油、berber oil、和椰子油;脂肪酸,如棕榈酸、硬脂酸、和亚油酸;醇,如丙三醇和乙醇;和有机酸,如乙酸。这些碳源可单独或组合使用,但不限于此。
氮源的实例可包括有机氮源,如蛋白胨、酵母提取物、肉汤、麦芽提取物、玉米浆(CSL)、和豆粉;和无机氮源,如尿素、硫酸铵、氯化铵、磷酸铵、碳酸铵、和硝酸铵。这些氮源可单独或组合使用、但不限于此。培养基可进一步包括磷酸二氢钾、磷酸氢二钾、和相应的含钠盐作为磷源。培养基可包括金属,如硫酸镁和硫酸铁。另外,可包括氨基酸、维生素和合适的前体。这些培养基或前体可以分批培养或连续培养的形式加入培养物中,但不限于此。
另外,可在培养期间以合适的方式通过添加化合物如氢氧化铵、氢氧化钾、氨、磷酸、和硫酸调节培养物的pH。另外,可利用消泡剂如脂肪酸聚二醇酯在培养期间防止泡沫形成。另外,为维持培养液的有氧条件,可添加氧气或包含氧气的气体(例如,空气)至培养物。培养温度可以为20℃至45℃,具体地25℃至40℃,但不限于此。培养可持续,直到O-乙酰基高丝氨酸的生产达到目标水平,具体地持续10小时至160小时,但不限于此。
本发明的生产O-乙酰基高丝氨酸的方法可进一步包括从培养的微生物或其培养产物回收O-乙酰基高丝氨酸。目标O-乙酰基高丝氨酸的回收可通过根据本发明所述的微生物培养方法进行,例如,本领域已知的适当方法,如分批培养、连续培养、和补料分批培养。
回收可包括纯化步骤。
由此回收的O-乙酰基高丝氨酸可通过两步法(韩国专利号10-0905381)生产甲硫氨酸。
两步法包括通过利用具有O-乙酰基高丝氨酸硫化氢解酶活性的酶或具有这种酶的微生物的酶反应,同时利用L-甲硫氨酸前体生产微生物生产的O-乙酰基高丝氨酸和甲硫醇作为底物生产L-甲硫氨酸和有机酸的过程。
更具体地,本发明提供通过O-乙酰基高丝氨酸硫化氢解酶等的酶反应,利用通过上述方法积累的O-乙酰基高丝氨酸作为底物生产L-甲硫氨酸的方法。
在第二步中,当O-乙酰基高丝氨酸用作L-甲硫氨酸前体时,可使用源自微生物的O-乙酰基高丝氨酸硫化氢解酶,该微生物具体地属于钩端螺旋体属、色杆菌属、和生丝单胞菌属(Hyphomonas sp.),更具体地属于迈氏钩端螺旋体(Leptospira meyeri)、铜绿色假单胞菌(Pseudomonas aurogenosa)、海王生丝单胞菌(Hyphomonas Neptunium)、和紫色色杆菌(Chromobacterium Violaceum)。
反应同下文显示。
生产甲硫氨酸的另外过程被公开于韩国专利号10-0905381中,该专利的整个说明书可作为引用被本发明包括。
用于发明的方式
下文参考下列实施例对本发明进行更详细的描述。然而,这些实施例仅以示例为目的,并非意图本发明受限于这些实施例。
参考实施例:构建O-乙酰基高丝氨酸生产微生物
<1-1>敲除源自野生型大肠杆菌的metB基因(国际公开号WO 2008/013432)利用大肠杆菌(埃希氏菌属的代表性微生物)构建O-乙酰基高丝氨酸生产微生物。为此,使用从美国菌种保藏中心(American Type Culture Collection,ATCC)获得的野生型大肠杆菌K12W3110(ATCC27325)。首先,为阻断O-琥珀酰基-L-高丝氨酸至胱硫醚的合成途径,敲除编码胱硫醚合酶的metB基因(SEQ ID NO:10)。具体地,通过FRT-一步PCR敲除法(PNAS(2000)vol97:P6640-6645)敲除编码胱硫醚合酶的metB基因。
具体地,利用SEQ ID NOS:30和31的引物,基于pKD3载体(PNAS(2000)vol 97:P6640-6645)作为模板,通过PCR反应,如下构建metB缺失盒:94℃变性30秒,55℃退火30秒,和72℃延伸1分钟,30个循环。将所得PCR产物在1.0%琼脂糖凝胶上进行电泳,并纯化由此获得的1.2kb DNA带。将回收的DNA片段电穿孔到已被pKD46载体转化的大肠杆菌(K12)W3110(PNAS(2000)vol 97:P6640-6645)中。关于电穿孔,将pKD46转化的W3110菌株在30℃下在包含100μg/L氨苄西林和5mM阿拉伯糖(L-阿拉伯糖)的LB培养基中培养,直到OD600=0.6,并在用无菌蒸馏水洗涤两次和用10%丙三醇洗涤一次后使用。电穿孔在2500V下进行。将回收的菌株在包含25μg/L氯霉素的LB平板上划线(streak),在37℃下培养过夜,并选择显示抗性的菌株。使选择的菌株在相同条件下,利用相同引物,基于该菌株作为模板,进行PCR反应,并通过观察1.0%琼脂糖凝胶上1.2kb的基因尺寸来证实metB基因的敲除。将由此证实的菌株在经pCP20载体(PNAS(2000)vol 97:P6640-6645)再次转化后在LB培养基中培养,并构建最终的metB基因敲除菌株(其中通过相同条件下进行的PCR,在1.0%琼脂糖凝胶上,基因尺寸减少至150bp),并证实氯霉素标记物的去除。由此构建的菌株被命名为“W3-B”。
<1-2>敲除thrB基因(国际公开号WO 2008/013432)
在增加来源于高丝氨酸的O-琥珀酰基高丝氨酸合成量的工作中,敲除thrB基因——高丝氨酸激酶编码基因。关于从实施例1构建的W3-B菌株敲除thrB基因,采用敲除metB基因时使用的FRT一步PCR敲除法。
利用SEQ ID NOS:32和33的引物,基于pKD4载体(PNAS(2000)vol 97:P6640-6645)作为模板,通过PCR,如下构建thrB缺失盒:94℃变性30秒,55℃退火30秒,和72℃延伸1分钟,30个循环。
将所得PCR产物在1.0%琼脂糖凝胶上进行电泳,并纯化由此获得的1.6kb DNA带。将回收的DNA片段电穿孔到已被pKD46载体转化的W3-B菌株中。将回收的菌株在包含50μg/L卡那霉素的LB平板上划线,在37℃下培养过夜,并选择显示抗性的菌株。
将选择的菌株利用相同的SEQ ID NOS:32和33的引物,直接基于该菌株作为模板,在相同条件下进行PCR反应,并通过选择在1.0%琼脂糖凝胶上具有1.6kb基因尺寸的菌株来证实thrB基因的敲除。将由此证实的菌株经pCP20载体再次转化后在LB培养基中培养,并构建最终的thrB基因敲除菌株(其中通过相同条件下进行的PCR,在1.0%琼脂糖凝胶上,基因尺寸减少至150bp),并证实卡那霉素标记物的去除。由此构建的菌株被命名为“W3-BT”。
<1-3>具有高丝氨酸乙酰基转移酶活性的变体metA(国际公开号WO 2012/087039)
为强化参考实施例<1-2>获得的菌株的高丝氨酸乙酰基转移酶活性,意图引入编码高丝氨酸乙酰基转移酶的突变型metA基因(SEQ ID NOS:24和26)。
首先,为构建活性强化的metA基因变体,利用SEQ ID NOS:34和35的引物,基于野生型菌株W3110的染色体作为模板,进行PCR反应,扩增编码高丝氨酸O-琥珀酰基转移酶的metA基因。
基于在NIH基因库登记的大肠杆菌染色体NC_000913的多核苷酸序列来制备PCR反应中使用的引物,并且SEQ ID NOS:34和35的引物分别具有EcoRV和HindIII限制位点。将由此获得的PCR产物和包括Pcj1的pCL1920质粒分别用EcoRV和HindIII处理,并将PCR产物克隆到pCL1920质粒中。用克隆质粒转化大肠杆菌DH5α,并在包含50μg/mL大观霉素的LB平板上选择转化的大肠杆菌DH5α,从中获得质粒。由此获得的质粒被命名为“pCL_Pcj1_metA”。
然后,利用定点诱变试剂盒(Strata基因,USA),基于以上构建的pCL_Pcj1_metA质粒作为模板,用谷氨酸(Glu)(G111E)取代O-琥珀酰基转移酶的第111位氨基酸,甘氨酸(Gly)。由此构建的包括G111E metA基因变体的质粒被命名为“pCL_Pcj1_metA(EL)”。
另外,为进行O-琥珀酰基转移酶的第111位氨基酸从甘氨酸至谷氨酸取代,和第112位氨基酸从亮氨酸至组氨酸取代,使用SEQ ID NOS:38和39的引物。包括其中第111位氨基酸从甘氨酸至谷氨酸取代并且第112位氨基酸从亮氨酸至组氨酸取代的metA基因的质粒被命名为“pCL_Pcj1_metA(EH)”。
然后,利用pKD3载体作为模板以及SEQ ID NOS:40和41的引物,通过PCR,如下构建用于将metA(EH)替换到菌株中的替换盒:94℃变性30秒,55℃退火30秒,和72℃延伸2分钟,30个循环。利用pCL-Pcj1-metA(EH)作为替换盒metA(EH)部分的模板以及SEQ ID NOS:42和43的引物、和野生型metA部分的SEQ ID NOS:42和45的引物,获得各个PCR产物。利用三种不同PCR产物以及SEQ ID NOS:42和45的引物,构建包括氯霉素标记物部分的metA(EH)替换盒,并将其电穿孔到参考实施例<1-2>构建的已被pKD46载体转化的W3-BT菌株中。将由此证实的菌株经pCP20载体再次转化后在LB培养基中培养,其中氯霉素标记物被去除并且metA基因被metA(EH)取代的菌株被命名为“W3-BTA”。
<1-4>构建具有2个拷贝的ppc、aspC和asd基因的菌株(欧洲专利申请公开号EP2290051)
为增加参考实施例<1-3>构建的W3-BTA菌株的O-乙酰基高丝氨酸生产能力,通过引用此前提交的专利EP 2290051来增强生物合成途径。以与上述EP专利相同的方式,构建如下每种基因具有2个扩增拷贝的菌株:即,编码磷酸烯醇丙酮酸羧化酶的ppc基因,利用SEQ ID NOS:46、47、48和49的引物;编码天冬氨酸氨基转移酶的aspC基因,利用SEQ IDNOS:50和51的引物;和编码天冬氨酸半醛脱氢酶的asd基因,利用SEQ ID NOS:52、53、54和55的引物。具体地,在生产O-乙酰基高丝氨酸时生物合成途径增强的上述菌株被命名为“W3-BTA2PCD”(也被称为“WCJM”)。
<1-5>烧瓶培养实验
通过爱伦美氏烧瓶培养,测试参考实施例<1-3>和<1-4>构建的菌株的O-乙酰基高丝氨酸产量。
具体地,将W3110、W3-BTA、和WCJM菌株接种于LB培养基,并在33℃下培养过夜。然后,将其单个集落接种于3mL LB培养基,在33℃下培养5小时,在包含25mL O-乙酰基高丝氨酸生产培养基的250mL爱伦美氏烧瓶中稀释200倍,在33℃下在200rpm转速下培养30小时,并通过HPLC分析测定O-乙酰基高丝氨酸产量。所用培养基组成显示在下表1中,并且测定的O-乙酰基高丝氨酸产量显示在下表2中。
[表1]O-乙酰基高丝氨酸生产烧瓶培养基的组成
[表2]
结果表明,野生型W3110完全不生产O-乙酰基高丝氨酸,而W3-BTA菌株生产0.9g/LO-乙酰基高丝氨酸,并且生物合成途径强化的WCJM菌株生产1.2g/L O-乙酰基高丝氨酸。
实施例1:去除柠檬酸合酶活性
<1-1>在O-乙酰基高丝氨酸生产微生物中构建柠檬酸合酶基因敲除微生物
柠檬酸合酶(GltA)是TCA循环的第一步中的酶,并且始于草酰乙酸盐和乙酰辅酶A之间的反应。众所周知TCA循环减少导致生长抑制(Meded Rijksuniv Gent FakLandbouwkd Toegep Biol Wet.2001;66(3a):333-6)。然而,为增加乙酰辅酶A(用作O-乙酰基高丝氨酸的底物)的量,将要生产去除了柠檬酸合酶活性的W3-BTA和WCJM菌株。
具体地,利用SEQ ID NOS:56和57的引物,基于pKD4载体作为模板,通过PCR,如下敲除W3-BTA和WCJM菌株中的柠檬酸合酶基因:94℃变性30秒,55℃退火30秒,和72℃延伸2分钟,30个循环。将所得PCR产物在1.0%琼脂糖凝胶上进行电泳,证实基因尺寸为1.6kb,并纯化其DNA。将回收的DNA片段电穿孔到已被pKD46载体转化的W3-BTA和WCJM菌株中。关于电穿孔,将pKD46载体转化的W3-BTA和WCJM菌株在30℃下在包含100μg/L氨苄西林和5mM阿拉伯糖的LB培养基中培养,直到OD600=0.6,并用蒸馏水洗涤两次和10%丙三醇洗涤一次,待用。电穿孔在2500V下进行。将由此回收的菌株在包含50μg/L卡那霉素的LB平板上划线,在37℃下培养,并选择显示抗性的菌株。
将选择的菌株在相同条件下利用SEQ ID NOS:58和59的引物进行PCR,在1.0%琼脂糖凝胶上进行电泳,并观察到基因尺寸为2.5kb,从而证实缺失盒被插入染色体的柠檬酸合酶基因部分。将由此证实的菌株用pCP20载体再次转化,在LB培养基中培养,并通过PCR构建敲除了柠檬酸合酶基因,在1.0%琼脂糖凝胶上尺寸减少至1.1kb的菌株,并且证实卡那霉素标记物被去除。由此构建的菌株分别被命名为“W3-BTA-AD”和“WCJM-AD”。
<1-2>评价O-乙酰基高丝氨酸生产微生物中的柠檬酸合酶基因敲除微生物
W3-BTA-AD和WCJM-AD菌株可在LB培养基中生长,但由于敲除了柠檬酸合酶基因,其不能在包含O-乙酰基高丝氨酸的培养基中生长。为测试O-乙酰基高丝氨酸产量,在培养基现有组成中添加3g/L谷氨酸盐的条件下(表3——培养基中添加谷氨酸盐的组成)进行爱伦美氏烧瓶培养。
具体地,将W3-BTA-AD和WCJM-AD菌株接种于LB培养基,并在33℃下培养过夜。然后,将其单个集落接种于3mL LB培养基,在33℃下培养5小时,在包含25mL O-乙酰基高丝氨酸生产培养基(添加了谷氨酸盐)的250mL爱伦美氏烧瓶中稀释200倍,在33℃下在200rpm转速下培养30小时,并通过HPLC分析测定O-乙酰基高丝氨酸产量。测定的O-乙酰基高丝氨酸产量显示在下表4中。
[表3]在基础培养基中添加谷氨酸盐的培养基的组成
[表4]通过烧瓶培养生产O-乙酰基高丝氨酸
通过烧瓶培养进行O-乙酰基高丝氨酸生产的结果表明,W3-BTA菌株生产0.9g/LO-乙酰基高丝氨酸,而W3-BTA-AD生产1.4g/L O-乙酰基高丝氨酸,增加55.6%,尽管显示其葡萄糖消耗减少。WCJM菌株生产1.3g/L O-乙酰基高丝氨酸,而WCJM-AD菌株生产2.1g/L O-乙酰基高丝氨酸,因此证实由于敲除了柠檬酸合酶基因,O-乙酰基高丝氨酸生产能力提高61.5%。
实施例2:柠檬酸合酶蛋白活性减弱
<2-1>柠檬酸合酶基因修饰种类
实施例<1-1>构建的WCJM-AD菌株显示低培养速率,并且根据在大量参考文献(TheJournal of Biological Chemistry,2003,278,35435-35443)中已知的各种柠檬酸合酶修饰,选择显示活性减弱和乙酰辅酶A结合能力降低的三种不同类型的变体。关于这三种不同变体类型的信息显示在显示修饰基因的表5中,其中第145位氨基酸,酪氨酸(Y),被丙氨酸(A)取代,第167位氨基酸,赖氨酸(K),被丙氨酸(A)取代,和第204位氨基酸,苏氨酸(T),被丙氨酸(A)取代。
[表5]
柠檬酸合酶(gltA)变体评价
<2-2>在O-乙酰基高丝氨酸生产微生物中构建柠檬酸合酶蛋白活性减弱的微生物
发明人意图通过将如实施例<2-1>所示柠檬酸合酶蛋白活性减弱的变体引入O-乙酰基高丝氨酸生产微生物来增加生产能力。
为将这三种不同类型的柠檬酸合酶基因变体引入WCJM-AD菌株,如图1所示设计修饰替换盒。通过用核苷酸取代引物来合成各变体,并通过3种PCR产物构建各盒。关于柠檬酸合酶基因部分,利用W3110菌株作为模板,并且第145位氨基酸修饰,分别利用SEQ ID NOS:60和63以及SEQ ID NOS:62和61的引物,通过进行PCR反应,产生并获得尺寸为514bp和1,112bp的PCR产物。
同样,利用SEQ ID NOS:60和65的引物和SEQ ID NOS:64和61的引物,第167位氨基酸修饰产生尺寸为580bp和1,046bp的PCR产物,而利用SEQ ID NOS:60和67以及SEQ IDNOS:66和61的引物,第204位氨基酸修饰产生尺寸为688bp和936bp的PCR产物。关于共同的卡那霉素部分,利用SEQ ID NOS:68和69的引物,基于pKD4载体作为模板,进行PCR反应。具体地,为插入柠檬酸合酶基因位置,构建包括SEQ ID NO:69中柠檬酸合酶基因的下游多核苷酸序列的盒,并通过电泳获得1,571bp尺寸的PCR产物。分别利用SEQ ID NOS:60和69的引物,基于卡那霉素DNA片段(根据修饰收集的两个DNA片段中每一个的共同部分),如下进行缝合PCR反应(sewing PCR reaction):94℃变性30秒,55℃退火30秒,和72℃延伸4分钟,30个循环。在1.0%琼脂糖凝胶上验证各最终PCR产物,并且三个不同种类的柠檬酸合酶基因修饰盒得到3,115bp尺寸的DNA片段。将收集的DNA片段电穿孔到已被pKD46载体转化的WCJM-AD菌株中。关于电穿孔,将pKD46转化的W3110菌株在30℃下在包含100μg/L氨苄西林和5mM阿拉伯糖的LB培养基中培养,直到OD600=0.6,并且在用无菌蒸馏水洗涤两次和10%丙三醇洗涤一次后使用。电穿孔在2500V下进行。将回收的菌株在包含25μg/L卡那霉素的LB平板上划线,在37℃下培养过夜,并选择显示抗性的菌株。利用相同的SEQ ID NOS:58和59的引物,基于该菌株作为模板,在相同条件下,使选择的菌株进行PCR反应,并通过观察到1.0%琼脂糖凝胶上3.7kb的基因尺寸证实metB基因的敲除,从而证实修饰盒(其中柠檬酸合酶基因的氨基酸被取代)被插入。将由此证实的菌株经pCP20载体再次转化后在LB培养基中培养,并且构建关于柠檬酸合酶活性的三种变体菌株(通过相同条件下进行的PCR,在1.0%琼脂糖凝胶上,基因尺寸减少至2.5kb),并且证实卡那霉素标记物的去除。由此构建的菌株被命名为“WCJM-A145”、“WCJM-A167”、和“WCJM-A204”,并且引入了修饰的柠檬酸合酶基因的序列信息分别显示在SEQ ID NOS:1至3(氨基酸序列)和SEQ ID NOS:5至7(核苷酸序列)中。
<2-3>在O-乙酰基高丝氨酸生产微生物中评价柠檬酸合酶活性减弱的微生物
进行爱伦美氏烧瓶培养以测定柠檬酸合酶基因活性减弱的三种不同菌株WCJM-A145、WCJM-A167和WCJM-A204的O-乙酰基高丝氨酸产量。将四种菌株——即,WCJM-A145、WCJM-A167、和WCJM-A204菌株,包括WCJM菌株——接种于LB培养基,并在33℃下培养过夜。然后,将其单个集落接种于3mL LB培养基,在33℃下培养5小时,在包含25mL O-乙酰基高丝氨酸生产培养基的250mL爱伦美氏烧瓶中稀释200倍,在33℃下在200rpm转速下培养30小时,并通过HPLC分析测定O-乙酰基高丝氨酸产量。结果显示在下表6中。
[表6]通过烧瓶培养生产O-乙酰基高丝氨酸
通过烧瓶培养进行O-乙酰基高丝氨酸生产的结果表明,WCJM菌株生产1.3g/L O-乙酰基高丝氨酸,并且两菌株WCJM-A145和WCJM-A167分别生产2.0g/L和1.9g/L O-乙酰基高丝氨酸,同时随着其吸光度(OD)减少,其葡萄糖消耗量减少。考虑到谷氨酸盐从1.3g/L具体减少至0g/L,证实该结果是由柠檬酸合酶活性减弱导致的TCA循环流减少造成的。然而,WCJM-A204菌株显示谷氨酸盐增加,同时显示O-乙酰基高丝氨酸产量减少至0.2g/L,因此证实该修饰是活性强化修饰。
实施例3:柠檬酸合酶蛋白表达减弱
<3-1>构建柠檬酸合酶基因反义RNA(asRNA)的表达载体
发明人试图利用反义RNA(asRNA)技术来减弱柠檬酸合酶蛋白的表达。反义RNA技术是通过过表达目标基因的柠檬酸合酶mRNA的互补结合部分以阻止柠檬酸合酶mRNA和核糖体之间的结合从而减少蛋白质表达的方法。这种方法的优势在于:其可通过控制与柠檬酸合酶基因的mRNA的结合力调控抑制水平,并且这种方法还可用于构建重组微生物,因为这种方法可通过反义RNA控制基因的表达来有效构建和减少基因表达,而无需常规基因敲除方法。
载体构建参考参考文献(Methods Mol Biol.2012;815:307-19.doi:10.1007/978-1-61779-424-7_23)进行,并且关于过表达,将合酶基因的反义RNA区引入能够进行诱导的pBAD24质粒。pBAD24-柠檬酸合酶asRNA载体图谱显示在图2中。表达柠檬酸合酶基因的反义RNA的区域的尺寸为100bp,包括启动子区域的52bp区域和柠檬酸合酶起始密码子的48bp区域,并且降低反义RNA(asRNA)不稳定性的38bp配对末端(PT)结构连接至两个侧翼区域。利用SEQ ID NOS:70和71的引物获得柠檬酸合酶基因的反义RNA区,并且包括NcoI和HindIII限制位点以克隆到载体中。
由此获得的PCR产物的尺寸为194bp,并且将PCR产物分别经EcoRV和HindIII处理后克隆到pBAD24质粒中。用由此克隆的质粒转化大肠杆菌DH5α,从包含100μg/mL氨苄西林的LB平板选择转化的大肠杆菌DH5α,并从中获得质粒。由此获得的质粒被命名为“pBAD24-gltA asRNA”。
<3-2>柠檬酸合酶基因的反义RNA的表达载体在O-乙酰基高丝氨酸生产微生物中的引入和其评价
将pBAD24-gltA-asRNA,柠檬酸合酶基因的反义RNA的表达载体,转化到WCJM菌株(O-乙酰基高丝氨酸生产微生物)中。在此,转化的菌株被命名为“WCJM/A-asRNA”。具体地,尝试通过控制柠檬酸合酶的反义RNA的表达量来控制柠檬酸合酶蛋白的表达量,并且在此,可根据阿拉伯糖的浓度控制反义RNA的表达量。
结果是,证实在如实施例2减弱柠檬酸合酶活性时,O-乙酰基高丝氨酸产量增加。
另外,进行爱伦美氏烧瓶培养以考察O-乙酰基高丝氨酸产量是否随柠檬酸合酶表达量减少而增加。
具体地,将WCJM和WCJM/A-asRNA菌株接种于LB培养基,并在33℃下培养过夜。然后,将其单个集落接种于3mL LB培养基,在33℃下培养5小时,在包含25mL O-乙酰基高丝氨酸生产培养基的250mL爱伦美氏烧瓶中稀释200倍。具体地,为控制柠檬酸合酶的反义RNA的表达量,添加0mM、2mM、和5mM浓度的阿拉伯糖,并在33℃下在200rpm转速下培养15小时和30小时。通过HPLC分析测定O-乙酰基高丝氨酸产量,结果显示在下表7和8中。
[表7]
[表8]
结果是,证实在培养15小时时,WCJM/A-asRNA菌株显示,随着阿拉伯糖浓度,OD减少约1,而O-乙酰基高丝氨酸的浓度相似。然而,在培养30小时时,WCJM菌株(对照菌株)即使在阿拉伯糖浓度增加时也显示相同的OD和O-乙酰基高丝氨酸浓度,而WCJM/A-asRNA菌株(引入了柠檬酸合酶反义RNA的表达载体的菌株)显示出随着阿拉伯糖浓度增加的显著差异。在阿拉伯糖浓度为0mM时,OD为9.2,而在5mM阿拉伯糖浓度时,OD为7.1,减少5.1,并且O-乙酰基高丝氨酸量增加30.8%,尽管葡萄糖消耗量少。根据这些结果,证实不仅柠檬酸合酶活性减弱,蛋白质表达减弱也呈现相同的结果。
实施例4:O-乙酰基高丝氨酸高产率微生物的柠檬酸合酶活性的减弱和灭活
<4-1>柠檬酸合酶活性灭活的O-乙酰基高丝氨酸高产率微生物的构建和其评价
国际公开号WO 2012/087039详细公开了基于NTG突变由源自野生型W3110菌株的苏氨酸生产微生物构建O-乙酰基高丝氨酸生产微生物的方法。具体地,构建的高产率生产O-乙酰基高丝氨酸的菌株以登录号KCCM 11146P保藏于韩国微生物保藏中心(KoreanCulture Center of Microorganisms,KCCM)。
KCCM11146P菌株可在烧瓶培养期间消耗40g/L葡萄糖,并生产约15g/L至16g/L O-乙酰基高丝氨酸,因此被认为具有高O-乙酰基高丝氨酸生产能力。因此,为考察该菌株在去除柠檬酸合酶活性时是否生产较高产率的O-乙酰基高丝氨酸,对KCCM11146P菌株应用相同的构建方法。构建方法同实施例<1-1>,并且通过此方法,构建去除了柠檬酸合酶活性的KCCM11146P菌株,并将其被命名为“KCCM11146P-AD”。
通过爱伦美氏烧瓶培养测试去除了柠檬酸合酶活性的KCCM11146P菌株的O-乙酰基高丝氨酸产量。将KCCM11146P或KCCM11146P-AD菌株接种于LB培养基并在33℃下培养过夜。然后,将其单个集落接种于3mL LB培养基,在33℃下培养5小时,在包含25mL O-乙酰基高丝氨酸生产培养基(添加了谷氨酸盐)的250mL爱伦美氏烧瓶中稀释200倍,并在33℃下在200rpm转速下培养30小时。通过HPLC分析测定O-乙酰基高丝氨酸产量,结果显示在下表9中。
[表9]通过烧瓶培养生产O-乙酰基高丝氨酸
通过烧瓶培养进行O-乙酰基高丝氨酸生产的结果表明,KCCM11146P菌株生产14.2g/L O-乙酰基高丝氨酸,而KCCM11146P-AD菌株生产16.7g/L O-乙酰基高丝氨酸,增加17.6%,尽管其显示吸光度(OD)减少。
<4-2>柠檬酸合酶活性减弱的O-乙酰基高丝氨酸高产率微生物的构建和其评价
为测定KCCM11146P菌株——O-乙酰基高丝氨酸高产率菌株——是否甚至在柠檬酸合酶活性减弱时也生产较高产率的O-乙酰基高丝氨酸,将在实施例<2-1>所示的减弱蛋白质活性的三种变体类型中显示最高O-乙酰基高丝氨酸生产能力的第145位氨基酸的修饰(从酪氨酸(Y)至丙氨酸(A))和第167位氨基酸的修饰(从赖氨酸(K)至丙氨酸(A))应用于KCCM11146P菌株。
构建方法同实施例<2-2>,通过该方法,构建了柠檬酸合酶活性减弱的两种KCCM11146P菌株,并将其分别命名为“KCCM11146P-A145”和“KCCM11146P-A167”。
通过爱伦美氏烧瓶培养测试柠檬酸合酶活性减弱的KCCM11146P-A145和KCCM11146P-A167两种菌株的O-乙酰基高丝氨酸产量。将三种菌株(即,KCCM11146P-A145和KCCM11146P-A167菌株和KCCM11146P菌株)接种于LB培养基,并在33℃下培养过夜。然后,将其单个集落接种于3mL LB培养基,在33℃下培养5小时,在包含25mL O-乙酰基高丝氨酸生产培养基的250mL爱伦美氏烧瓶中稀释200倍,并在33℃下在200rpm转速下培养30小时。通过HPLC分析测定O-乙酰基高丝氨酸产量,结果显示在下表10中。
[表10]通过烧瓶培养生产O-乙酰基高丝氨酸
通过烧瓶培养进行O-乙酰基高丝氨酸生产的结果表明,KCCM11146P菌株生产15.0g/L O-乙酰基高丝氨酸,并且KCCM11146P-A145和KCCM11146P-A167两种菌株显示与实施例<2-3>相似的结果。这两种菌株分别生产17.5g/L和17.3g/L O-乙酰基高丝氨酸,增加约16.7%,尽管其都显示吸光度(OD)减少。
根据柠檬酸合酶活性减弱导致的TCA循环流减少,O-乙酰基高丝氨酸高产率菌株显示谷氨酸盐从1.6g/L减少至0g/L。
这些结果证明,柠檬酸合酶活性通过施加减弱的修饰而实现生产O-乙酰基高丝氨酸。另外,其还表明,当基于按照国际公开号WO2008/013432生产的O-乙酰基高丝氨酸作为模板,并且利用另外具有胱硫醚γ合酶、O-琥珀酰基高丝氨酸硫化氢解酶和O-乙酰基高丝氨酸硫化氢解酶活性的转化酶进行转化反应时,可以同时合成L-甲硫氨酸和乙酸盐。
本发明人证实,KCCM11146P菌株,柠檬酸合酶第167位氨基酸变体具有提高的O-乙酰基高丝氨酸产量,将KCCM11146P-A167菌株命名为“CA05-4007”,并且将其在2013年11月22日保藏于布达佩斯条约规定的国际保藏机构,韩国微生物保藏中心(KCCM)(登录号:KCCM11483P)。
根据前文,本发明所属领域技术人员能够理解本发明可以其他具体形式实施,而没有改变本发明的技术思路或重要特征。在这点上,本文公开的示例性实施方式仅以示例为目的,不应被解释为限制本发明的范围。相反,本发明不仅意图包括示例性实施方式,而且包括所附权利要求限定的在本发明精神和范围内可包括的各种可选项、改动、等同形式和其他实施方式。
序列表
<110> CJ第一制糖株式会社
<120> 生产O-乙酰基高丝氨酸的微生物和用其生产O-乙酰基高丝氨酸的方法
<130> OPA15136
<150> KR 10-2014-0076779
<151> 2014-06-23
<160> 71
<170> KopatentIn 2.0
<210> 1
<211> 427
<212> PRT
<213> 人工序列
<220>
<223> GltA A145
<400> 1
Met Ala Asp Thr Lys Ala Lys Leu Thr Leu Asn Gly Asp Thr Ala Val
1 5 10 15
Glu Leu Asp Val Leu Lys Gly Thr Leu Gly Gln Asp Val Ile Asp Ile
20 25 30
Arg Thr Leu Gly Ser Lys Gly Val Phe Thr Phe Asp Pro Gly Phe Thr
35 40 45
Ser Thr Ala Ser Cys Glu Ser Lys Ile Thr Phe Ile Asp Gly Asp Glu
50 55 60
Gly Ile Leu Leu His Arg Gly Phe Pro Ile Asp Gln Leu Ala Thr Asp
65 70 75 80
Ser Asn Tyr Leu Glu Val Cys Tyr Ile Leu Leu Asn Gly Glu Lys Pro
85 90 95
Thr Gln Glu Gln Tyr Asp Glu Phe Lys Thr Thr Val Thr Arg His Thr
100 105 110
Met Ile His Glu Gln Ile Thr Arg Leu Phe His Ala Phe Arg Arg Asp
115 120 125
Ser His Pro Met Ala Val Met Cys Gly Ile Thr Gly Ala Leu Ala Ala
130 135 140
Phe Ala His Asp Ser Leu Asp Val Asn Asn Pro Arg His Arg Glu Ile
145 150 155 160
Ala Ala Phe Arg Leu Leu Ser Lys Met Pro Thr Met Ala Ala Met Cys
165 170 175
Tyr Lys Tyr Ser Ile Gly Gln Pro Phe Val Tyr Pro Arg Asn Asp Leu
180 185 190
Ser Tyr Ala Gly Asn Phe Leu Asn Met Met Phe Ser Thr Pro Cys Glu
195 200 205
Pro Tyr Glu Val Asn Pro Ile Leu Glu Arg Ala Met Asp Arg Ile Leu
210 215 220
Ile Leu His Ala Asp His Glu Gln Asn Ala Ser Thr Ser Thr Val Arg
225 230 235 240
Thr Ala Gly Ser Ser Gly Ala Asn Pro Phe Ala Cys Ile Ala Ala Gly
245 250 255
Ile Ala Ser Leu Trp Gly Pro Ala His Gly Gly Ala Asn Glu Ala Ala
260 265 270
Leu Lys Met Leu Glu Glu Ile Ser Ser Val Lys His Ile Pro Glu Phe
275 280 285
Val Arg Arg Ala Lys Asp Lys Asn Asp Ser Phe Arg Leu Met Gly Phe
290 295 300
Gly His Arg Val Tyr Lys Asn Tyr Asp Pro Arg Ala Thr Val Met Arg
305 310 315 320
Glu Thr Cys His Glu Val Leu Lys Glu Leu Gly Thr Lys Asp Asp Leu
325 330 335
Leu Glu Val Ala Met Glu Leu Glu Asn Ile Ala Leu Asn Asp Pro Tyr
340 345 350
Phe Ile Glu Lys Lys Leu Tyr Pro Asn Val Asp Phe Tyr Ser Gly Ile
355 360 365
Ile Leu Lys Ala Met Gly Ile Pro Ser Ser Met Phe Thr Val Ile Phe
370 375 380
Ala Met Ala Arg Thr Val Gly Trp Ile Ala His Trp Ser Glu Met His
385 390 395 400
Ser Asp Gly Met Lys Ile Ala Arg Pro Arg Gln Leu Tyr Thr Gly Tyr
405 410 415
Glu Lys Arg Asp Phe Lys Ser Asp Ile Lys Arg
420 425
<210> 2
<211> 427
<212> PRT
<213> 人工序列
<220>
<223> GltA A167
<400> 2
Met Ala Asp Thr Lys Ala Lys Leu Thr Leu Asn Gly Asp Thr Ala Val
1 5 10 15
Glu Leu Asp Val Leu Lys Gly Thr Leu Gly Gln Asp Val Ile Asp Ile
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Arg Thr Leu Gly Ser Lys Gly Val Phe Thr Phe Asp Pro Gly Phe Thr
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Ser Thr Ala Ser Cys Glu Ser Lys Ile Thr Phe Ile Asp Gly Asp Glu
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Gly Ile Leu Leu His Arg Gly Phe Pro Ile Asp Gln Leu Ala Thr Asp
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Ser Asn Tyr Leu Glu Val Cys Tyr Ile Leu Leu Asn Gly Glu Lys Pro
85 90 95
Thr Gln Glu Gln Tyr Asp Glu Phe Lys Thr Thr Val Thr Arg His Thr
100 105 110
Met Ile His Glu Gln Ile Thr Arg Leu Phe His Ala Phe Arg Arg Asp
115 120 125
Ser His Pro Met Ala Val Met Cys Gly Ile Thr Gly Ala Leu Ala Ala
130 135 140
Phe Tyr His Asp Ser Leu Asp Val Asn Asn Pro Arg His Arg Glu Ile
145 150 155 160
Ala Ala Phe Arg Leu Leu Ser Ala Met Pro Thr Met Ala Ala Met Cys
165 170 175
Tyr Lys Tyr Ser Ile Gly Gln Pro Phe Val Tyr Pro Arg Asn Asp Leu
180 185 190
Ser Tyr Ala Gly Asn Phe Leu Asn Met Met Phe Ser Thr Pro Cys Glu
195 200 205
Pro Tyr Glu Val Asn Pro Ile Leu Glu Arg Ala Met Asp Arg Ile Leu
210 215 220
Ile Leu His Ala Asp His Glu Gln Asn Ala Ser Thr Ser Thr Val Arg
225 230 235 240
Thr Ala Gly Ser Ser Gly Ala Asn Pro Phe Ala Cys Ile Ala Ala Gly
245 250 255
Ile Ala Ser Leu Trp Gly Pro Ala His Gly Gly Ala Asn Glu Ala Ala
260 265 270
Leu Lys Met Leu Glu Glu Ile Ser Ser Val Lys His Ile Pro Glu Phe
275 280 285
Val Arg Arg Ala Lys Asp Lys Asn Asp Ser Phe Arg Leu Met Gly Phe
290 295 300
Gly His Arg Val Tyr Lys Asn Tyr Asp Pro Arg Ala Thr Val Met Arg
305 310 315 320
Glu Thr Cys His Glu Val Leu Lys Glu Leu Gly Thr Lys Asp Asp Leu
325 330 335
Leu Glu Val Ala Met Glu Leu Glu Asn Ile Ala Leu Asn Asp Pro Tyr
340 345 350
Phe Ile Glu Lys Lys Leu Tyr Pro Asn Val Asp Phe Tyr Ser Gly Ile
355 360 365
Ile Leu Lys Ala Met Gly Ile Pro Ser Ser Met Phe Thr Val Ile Phe
370 375 380
Ala Met Ala Arg Thr Val Gly Trp Ile Ala His Trp Ser Glu Met His
385 390 395 400
Ser Asp Gly Met Lys Ile Ala Arg Pro Arg Gln Leu Tyr Thr Gly Tyr
405 410 415
Glu Lys Arg Asp Phe Lys Ser Asp Ile Lys Arg
420 425
<210> 3
<211> 427
<212> PRT
<213> 人工序列
<220>
<223> GltA A204
<400> 3
Met Ala Asp Thr Lys Ala Lys Leu Thr Leu Asn Gly Asp Thr Ala Val
1 5 10 15
Glu Leu Asp Val Leu Lys Gly Thr Leu Gly Gln Asp Val Ile Asp Ile
20 25 30
Arg Thr Leu Gly Ser Lys Gly Val Phe Thr Phe Asp Pro Gly Phe Thr
35 40 45
Ser Thr Ala Ser Cys Glu Ser Lys Ile Thr Phe Ile Asp Gly Asp Glu
50 55 60
Gly Ile Leu Leu His Arg Gly Phe Pro Ile Asp Gln Leu Ala Thr Asp
65 70 75 80
Ser Asn Tyr Leu Glu Val Cys Tyr Ile Leu Leu Asn Gly Glu Lys Pro
85 90 95
Thr Gln Glu Gln Tyr Asp Glu Phe Lys Thr Thr Val Thr Arg His Thr
100 105 110
Met Ile His Glu Gln Ile Thr Arg Leu Phe His Ala Phe Arg Arg Asp
115 120 125
Ser His Pro Met Ala Val Met Cys Gly Ile Thr Gly Ala Leu Ala Ala
130 135 140
Phe Tyr His Asp Ser Leu Asp Val Asn Asn Pro Arg His Arg Glu Ile
145 150 155 160
Ala Ala Phe Arg Leu Leu Ser Lys Met Pro Thr Met Ala Ala Met Cys
165 170 175
Tyr Lys Tyr Ser Ile Gly Gln Pro Phe Val Tyr Pro Arg Asn Asp Leu
180 185 190
Ser Tyr Ala Gly Asn Phe Leu Asn Met Met Phe Ser Ala Pro Cys Glu
195 200 205
Pro Tyr Glu Val Asn Pro Ile Leu Glu Arg Ala Met Asp Arg Ile Leu
210 215 220
Ile Leu His Ala Asp His Glu Gln Asn Ala Ser Thr Ser Thr Val Arg
225 230 235 240
Thr Ala Gly Ser Ser Gly Ala Asn Pro Phe Ala Cys Ile Ala Ala Gly
245 250 255
Ile Ala Ser Leu Trp Gly Pro Ala His Gly Gly Ala Asn Glu Ala Ala
260 265 270
Leu Lys Met Leu Glu Glu Ile Ser Ser Val Lys His Ile Pro Glu Phe
275 280 285
Val Arg Arg Ala Lys Asp Lys Asn Asp Ser Phe Arg Leu Met Gly Phe
290 295 300
Gly His Arg Val Tyr Lys Asn Tyr Asp Pro Arg Ala Thr Val Met Arg
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Glu Thr Cys His Glu Val Leu Lys Glu Leu Gly Thr Lys Asp Asp Leu
325 330 335
Leu Glu Val Ala Met Glu Leu Glu Asn Ile Ala Leu Asn Asp Pro Tyr
340 345 350
Phe Ile Glu Lys Lys Leu Tyr Pro Asn Val Asp Phe Tyr Ser Gly Ile
355 360 365
Ile Leu Lys Ala Met Gly Ile Pro Ser Ser Met Phe Thr Val Ile Phe
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Ala Met Ala Arg Thr Val Gly Trp Ile Ala His Trp Ser Glu Met His
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Ser Asp Gly Met Lys Ile Ala Arg Pro Arg Gln Leu Tyr Thr Gly Tyr
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Glu Lys Arg Asp Phe Lys Ser Asp Ile Lys Arg
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<210> 4
<211> 427
<212> PRT
<213> 大肠杆菌
<220>
<221> 肽
<222> (1)..(427)
<223> GltA
<400> 4
Met Ala Asp Thr Lys Ala Lys Leu Thr Leu Asn Gly Asp Thr Ala Val
1 5 10 15
Glu Leu Asp Val Leu Lys Gly Thr Leu Gly Gln Asp Val Ile Asp Ile
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Arg Thr Leu Gly Ser Lys Gly Val Phe Thr Phe Asp Pro Gly Phe Thr
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Ser Thr Ala Ser Cys Glu Ser Lys Ile Thr Phe Ile Asp Gly Asp Glu
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Gly Ile Leu Leu His Arg Gly Phe Pro Ile Asp Gln Leu Ala Thr Asp
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Ser Asn Tyr Leu Glu Val Cys Tyr Ile Leu Leu Asn Gly Glu Lys Pro
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Thr Gln Glu Gln Tyr Asp Glu Phe Lys Thr Thr Val Thr Arg His Thr
100 105 110
Met Ile His Glu Gln Ile Thr Arg Leu Phe His Ala Phe Arg Arg Asp
115 120 125
Ser His Pro Met Ala Val Met Cys Gly Ile Thr Gly Ala Leu Ala Ala
130 135 140
Phe Tyr His Asp Ser Leu Asp Val Asn Asn Pro Arg His Arg Glu Ile
145 150 155 160
Ala Ala Phe Arg Leu Leu Ser Lys Met Pro Thr Met Ala Ala Met Cys
165 170 175
Tyr Lys Tyr Ser Ile Gly Gln Pro Phe Val Tyr Pro Arg Asn Asp Leu
180 185 190
Ser Tyr Ala Gly Asn Phe Leu Asn Met Met Phe Ser Thr Pro Cys Glu
195 200 205
Pro Tyr Glu Val Asn Pro Ile Leu Glu Arg Ala Met Asp Arg Ile Leu
210 215 220
Ile Leu His Ala Asp His Glu Gln Asn Ala Ser Thr Ser Thr Val Arg
225 230 235 240
Thr Ala Gly Ser Ser Gly Ala Asn Pro Phe Ala Cys Ile Ala Ala Gly
245 250 255
Ile Ala Ser Leu Trp Gly Pro Ala His Gly Gly Ala Asn Glu Ala Ala
260 265 270
Leu Lys Met Leu Glu Glu Ile Ser Ser Val Lys His Ile Pro Glu Phe
275 280 285
Val Arg Arg Ala Lys Asp Lys Asn Asp Ser Phe Arg Leu Met Gly Phe
290 295 300
Gly His Arg Val Tyr Lys Asn Tyr Asp Pro Arg Ala Thr Val Met Arg
305 310 315 320
Glu Thr Cys His Glu Val Leu Lys Glu Leu Gly Thr Lys Asp Asp Leu
325 330 335
Leu Glu Val Ala Met Glu Leu Glu Asn Ile Ala Leu Asn Asp Pro Tyr
340 345 350
Phe Ile Glu Lys Lys Leu Tyr Pro Asn Val Asp Phe Tyr Ser Gly Ile
355 360 365
Ile Leu Lys Ala Met Gly Ile Pro Ser Ser Met Phe Thr Val Ile Phe
370 375 380
Ala Met Ala Arg Thr Val Gly Trp Ile Ala His Trp Ser Glu Met His
385 390 395 400
Ser Asp Gly Met Lys Ile Ala Arg Pro Arg Gln Leu Tyr Thr Gly Tyr
405 410 415
Glu Lys Arg Asp Phe Lys Ser Asp Ile Lys Arg
420 425
<210> 5
<211> 1284
<212> DNA
<213> 人工序列
<220>
<223> GltA A145
<400> 5
atggctgata caaaagcaaa actcaccctc aacggggata cagctgttga actggatgtg 60
ctgaaaggca cgctgggtca agatgttatt gatatccgta ctctcggttc aaaaggtgtg 120
ttcacctttg acccaggctt cacttcaacc gcatcctgcg aatctaaaat tacttttatt 180
gatggtgatg aaggtatttt gctgcaccgc ggtttcccga tcgatcagct ggcgaccgat 240
tctaactacc tggaagtttg ttacatcctg ctgaatggtg aaaaaccgac tcaggaacag 300
tatgacgaat ttaaaactac ggtgacccgt cataccatga tccacgagca gattacccgt 360
ctgttccatg ctttccgtcg cgactcgcat ccaatggcag tcatgtgtgg tattaccggc 420
gcgctggcgg cgttcgcgca cgactcgctg gatgttaaca atcctcgtca ccgtgaaatt 480
gccgcgttcc gcctgctgtc gaaaatgccg accatggccg cgatgtgtta caagtattcc 540
attggtcagc catttgttta cccgcgcaac gatctctcct acgccggtaa cttcctgaat 600
atgatgttct ccacgccgtg cgaaccgtat gaagttaatc cgattctgga acgtgctatg 660
gaccgtattc tgatcctgca cgctgaccat gaacagaacg cctctacctc caccgtgcgt 720
accgctggct cttcgggtgc gaacccgttt gcctgtatcg cagcaggtat tgcttcactg 780
tggggacctg cgcacggcgg tgctaacgaa gcggcgctga aaatgctgga agaaatcagc 840
tccgttaaac acattccgga atttgttcgt cgtgcgaaag acaaaaatga ttctttccgc 900
ctgatgggct tcggtcaccg cgtgtacaaa aattacgacc cgcgcgccac cgtaatgcgt 960
gaaacctgcc atgaagtgct gaaagagctg ggcacgaagg atgacctgct ggaagtggct 1020
atggagctgg aaaacatcgc gctgaacgac ccgtacttta tcgagaagaa actgtacccg 1080
aacgtcgatt tctactctgg tatcatcctg aaagcgatgg gtattccgtc ttccatgttc 1140
accgtcattt tcgcaatggc acgtaccgtt ggctggatcg cccactggag cgaaatgcac 1200
agtgacggta tgaagattgc ccgtccgcgt cagctgtata caggatatga aaaacgcgac 1260
tttaaaagcg atatcaagcg ttaa 1284
<210> 6
<211> 1284
<212> DNA
<213> 人工序列
<220>
<223> GltA A167
<400> 6
atggctgata caaaagcaaa actcaccctc aacggggata cagctgttga actggatgtg 60
ctgaaaggca cgctgggtca agatgttatt gatatccgta ctctcggttc aaaaggtgtg 120
ttcacctttg acccaggctt cacttcaacc gcatcctgcg aatctaaaat tacttttatt 180
gatggtgatg aaggtatttt gctgcaccgc ggtttcccga tcgatcagct ggcgaccgat 240
tctaactacc tggaagtttg ttacatcctg ctgaatggtg aaaaaccgac tcaggaacag 300
tatgacgaat ttaaaactac ggtgacccgt cataccatga tccacgagca gattacccgt 360
ctgttccatg ctttccgtcg cgactcgcat ccaatggcag tcatgtgtgg tattaccggc 420
gcgctggcgg cgttctatca cgactcgctg gatgttaaca atcctcgtca ccgtgaaatt 480
gccgcgttcc gcctgctgtc ggcgatgccg accatggccg cgatgtgtta caagtattcc 540
attggtcagc catttgttta cccgcgcaac gatctctcct acgccggtaa cttcctgaat 600
atgatgttct ccacgccgtg cgaaccgtat gaagttaatc cgattctgga acgtgctatg 660
gaccgtattc tgatcctgca cgctgaccat gaacagaacg cctctacctc caccgtgcgt 720
accgctggct cttcgggtgc gaacccgttt gcctgtatcg cagcaggtat tgcttcactg 780
tggggacctg cgcacggcgg tgctaacgaa gcggcgctga aaatgctgga agaaatcagc 840
tccgttaaac acattccgga atttgttcgt cgtgcgaaag acaaaaatga ttctttccgc 900
ctgatgggct tcggtcaccg cgtgtacaaa aattacgacc cgcgcgccac cgtaatgcgt 960
gaaacctgcc atgaagtgct gaaagagctg ggcacgaagg atgacctgct ggaagtggct 1020
atggagctgg aaaacatcgc gctgaacgac ccgtacttta tcgagaagaa actgtacccg 1080
aacgtcgatt tctactctgg tatcatcctg aaagcgatgg gtattccgtc ttccatgttc 1140
accgtcattt tcgcaatggc acgtaccgtt ggctggatcg cccactggag cgaaatgcac 1200
agtgacggta tgaagattgc ccgtccgcgt cagctgtata caggatatga aaaacgcgac 1260
tttaaaagcg atatcaagcg ttaa 1284
<210> 7
<211> 1284
<212> DNA
<213> 人工序列
<220>
<223> GltA A204
<400> 7
atggctgata caaaagcaaa actcaccctc aacggggata cagctgttga actggatgtg 60
ctgaaaggca cgctgggtca agatgttatt gatatccgta ctctcggttc aaaaggtgtg 120
ttcacctttg acccaggctt cacttcaacc gcatcctgcg aatctaaaat tacttttatt 180
gatggtgatg aaggtatttt gctgcaccgc ggtttcccga tcgatcagct ggcgaccgat 240
tctaactacc tggaagtttg ttacatcctg ctgaatggtg aaaaaccgac tcaggaacag 300
tatgacgaat ttaaaactac ggtgacccgt cataccatga tccacgagca gattacccgt 360
ctgttccatg ctttccgtcg cgactcgcat ccaatggcag tcatgtgtgg tattaccggc 420
gcgctggcgg cgttctatca cgactcgctg gatgttaaca atcctcgtca ccgtgaaatt 480
gccgcgttcc gcctgctgtc gaaaatgccg accatggccg cgatgtgtta caagtattcc 540
attggtcagc catttgttta cccgcgcaac gatctctcct acgccggtaa cttcctgaat 600
atgatgttct ccgcgccgtg cgaaccgtat gaagttaatc cgattctgga acgtgctatg 660
gaccgtattc tgatcctgca cgctgaccat gaacagaacg cctctacctc caccgtgcgt 720
accgctggct cttcgggtgc gaacccgttt gcctgtatcg cagcaggtat tgcttcactg 780
tggggacctg cgcacggcgg tgctaacgaa gcggcgctga aaatgctgga agaaatcagc 840
tccgttaaac acattccgga atttgttcgt cgtgcgaaag acaaaaatga ttctttccgc 900
ctgatgggct tcggtcaccg cgtgtacaaa aattacgacc cgcgcgccac cgtaatgcgt 960
gaaacctgcc atgaagtgct gaaagagctg ggcacgaagg atgacctgct ggaagtggct 1020
atggagctgg aaaacatcgc gctgaacgac ccgtacttta tcgagaagaa actgtacccg 1080
aacgtcgatt tctactctgg tatcatcctg aaagcgatgg gtattccgtc ttccatgttc 1140
accgtcattt tcgcaatggc acgtaccgtt ggctggatcg cccactggag cgaaatgcac 1200
agtgacggta tgaagattgc ccgtccgcgt cagctgtata caggatatga aaaacgcgac 1260
tttaaaagcg atatcaagcg ttaa 1284
<210> 8
<211> 1284
<212> DNA
<213> 大肠杆菌
<220>
<221> 基因
<222> (1)..(1284)
<223> GltA
<400> 8
atggctgata caaaagcaaa actcaccctc aacggggata cagctgttga actggatgtg 60
ctgaaaggca cgctgggtca agatgttatt gatatccgta ctctcggttc aaaaggtgtg 120
ttcacctttg acccaggctt cacttcaacc gcatcctgcg aatctaaaat tacttttatt 180
gatggtgatg aaggtatttt gctgcaccgc ggtttcccga tcgatcagct ggcgaccgat 240
tctaactacc tggaagtttg ttacatcctg ctgaatggtg aaaaaccgac tcaggaacag 300
tatgacgaat ttaaaactac ggtgacccgt cataccatga tccacgagca gattacccgt 360
ctgttccatg ctttccgtcg cgactcgcat ccaatggcag tcatgtgtgg tattaccggc 420
gcgctggcgg cgttctatca cgactcgctg gatgttaaca atcctcgtca ccgtgaaatt 480
gccgcgttcc gcctgctgtc gaaaatgccg accatggccg cgatgtgtta caagtattcc 540
attggtcagc catttgttta cccgcgcaac gatctctcct acgccggtaa cttcctgaat 600
atgatgttct ccacgccgtg cgaaccgtat gaagttaatc cgattctgga acgtgctatg 660
gaccgtattc tgatcctgca cgctgaccat gaacagaacg cctctacctc caccgtgcgt 720
accgctggct cttcgggtgc gaacccgttt gcctgtatcg cagcaggtat tgcttcactg 780
tggggacctg cgcacggcgg tgctaacgaa gcggcgctga aaatgctgga agaaatcagc 840
tccgttaaac acattccgga atttgttcgt cgtgcgaaag acaaaaatga ttctttccgc 900
ctgatgggct tcggtcaccg cgtgtacaaa aattacgacc cgcgcgccac cgtaatgcgt 960
gaaacctgcc atgaagtgct gaaagagctg ggcacgaagg atgacctgct ggaagtggct 1020
atggagctgg aaaacatcgc gctgaacgac ccgtacttta tcgagaagaa actgtacccg 1080
aacgtcgatt tctactctgg tatcatcctg aaagcgatgg gtattccgtc ttccatgttc 1140
accgtcattt tcgcaatggc acgtaccgtt ggctggatcg cccactggag cgaaatgcac 1200
agtgacggta tgaagattgc ccgtccgcgt cagctgtata caggatatga aaaacgcgac 1260
tttaaaagcg atatcaagcg ttaa 1284
<210> 9
<211> 386
<212> PRT
<213> 大肠杆菌
<220>
<221> 肽
<222> (1)..(386)
<223> metB
<400> 9
Met Thr Arg Lys Gln Ala Thr Ile Ala Val Arg Ser Gly Leu Asn Asp
1 5 10 15
Asp Glu Gln Tyr Gly Cys Val Val Pro Pro Ile His Leu Ser Ser Thr
20 25 30
Tyr Asn Phe Thr Gly Phe Asn Glu Pro Arg Ala His Asp Tyr Ser Arg
35 40 45
Arg Gly Asn Pro Thr Arg Asp Val Val Gln Arg Ala Leu Ala Glu Leu
50 55 60
Glu Gly Gly Ala Gly Ala Val Leu Thr Asn Thr Gly Met Ser Ala Ile
65 70 75 80
His Leu Val Thr Thr Val Phe Leu Lys Pro Gly Asp Leu Leu Val Ala
85 90 95
Pro His Asp Cys Tyr Gly Gly Ser Tyr Arg Leu Phe Asp Ser Leu Ala
100 105 110
Lys Arg Gly Cys Tyr Arg Val Leu Phe Val Asp Gln Gly Asp Glu Gln
115 120 125
Ala Leu Arg Ala Ala Leu Ala Glu Lys Pro Lys Leu Val Leu Val Glu
130 135 140
Ser Pro Ser Asn Pro Leu Leu Arg Val Val Asp Ile Ala Lys Ile Cys
145 150 155 160
His Leu Ala Arg Glu Val Gly Ala Val Ser Val Val Asp Asn Thr Phe
165 170 175
Leu Ser Pro Ala Leu Gln Asn Pro Leu Ala Leu Gly Ala Asp Leu Val
180 185 190
Leu His Ser Cys Thr Lys Tyr Leu Asn Gly His Ser Asp Val Val Ala
195 200 205
Gly Val Val Ile Ala Lys Asp Pro Asp Val Val Thr Glu Leu Ala Trp
210 215 220
Trp Ala Asn Asn Ile Gly Val Thr Gly Gly Ala Phe Asp Ser Tyr Leu
225 230 235 240
Leu Leu Arg Gly Leu Arg Thr Leu Val Pro Arg Met Glu Leu Ala Gln
245 250 255
Arg Asn Ala Gln Ala Ile Val Lys Tyr Leu Gln Thr Gln Pro Leu Val
260 265 270
Lys Lys Leu Tyr His Pro Ser Leu Pro Glu Asn Gln Gly His Glu Ile
275 280 285
Ala Ala Arg Gln Gln Lys Gly Phe Gly Ala Met Leu Ser Phe Glu Leu
290 295 300
Asp Gly Asp Glu Gln Thr Leu Arg Arg Phe Leu Gly Gly Leu Ser Leu
305 310 315 320
Phe Thr Leu Ala Glu Ser Leu Gly Gly Val Glu Ser Leu Ile Ser His
325 330 335
Ala Ala Thr Met Thr His Ala Gly Met Ala Pro Glu Ala Arg Ala Ala
340 345 350
Ala Gly Ile Ser Glu Thr Leu Leu Arg Ile Ser Thr Gly Ile Glu Asp
355 360 365
Gly Glu Asp Leu Ile Ala Asp Leu Glu Asn Gly Phe Arg Ala Ala Asn
370 375 380
Lys Gly
385
<210> 10
<211> 1161
<212> DNA
<213> 大肠杆菌
<220>
<221> 基因
<222> (1)..(1161)
<223> metB
<400> 10
atgacgcgta aacaggccac catcgcagtg cgtagcgggt taaatgacga cgaacagtat 60
ggttgcgttg tcccaccgat ccatctttcc agcacctata actttaccgg atttaatgaa 120
ccgcgcgcgc atgattactc gcgtcgcggc aacccaacgc gcgatgtggt tcagcgtgcg 180
ctggcagaac tggaaggtgg tgctggtgca gtacttacta ataccggcat gtccgcgatt 240
cacctggtaa cgaccgtctt tttgaaacct ggcgatctgc tggttgcgcc gcacgactgc 300
tacggcggta gctatcgcct gttcgacagt ctggcgaaac gcggttgcta tcgcgtgttg 360
tttgttgatc aaggcgatga acaggcatta cgggcagcgc tggcagaaaa acccaaactg 420
gtactggtag aaagcccaag taatccattg ttacgcgtcg tggatattgc gaaaatctgc 480
catctggcaa gggaagtcgg ggcggtgagc gtggtggata acaccttctt aagcccggca 540
ttacaaaatc cgctggcatt aggtgccgat ctggtgttgc attcatgcac gaaatatctg 600
aacggtcact cagacgtagt ggccggcgtg gtgattgcta aagacccgga cgttgtcact 660
gaactggcct ggtgggcaaa caatattggc gtgacgggcg gcgcgtttga cagctatctg 720
ctgctacgtg ggttgcgaac gctggtgccg cgtatggagc tggcgcagcg caacgcgcag 780
gcgattgtga aatacctgca aacccagccg ttggtgaaaa aactgtatca cccgtcgttg 840
ccggaaaatc aggggcatga aattgccgcg cgccagcaaa aaggctttgg cgcaatgttg 900
agttttgaac tggatggcga tgagcagacg ctgcgtcgtt tcctgggcgg gctgtcgttg 960
tttacgctgg cggaatcatt agggggagtg gaaagtttaa tctctcacgc cgcaaccatg 1020
acacatgcag gcatggcacc agaagcgcgt gctgccgccg ggatctccga gacgctgctg 1080
cgtatctcca ccggtattga agatggcgaa gatttaattg ccgacctgga aaatggcttc 1140
cgggctgcaa acaaggggta a 1161
<210> 11
<211> 310
<212> PRT
<213> 大肠杆菌
<220>
<221> 肽
<222> (1)..(310)
<223> thrB
<400> 11
Met Val Lys Val Tyr Ala Pro Ala Ser Ser Ala Asn Met Ser Val Gly
1 5 10 15
Phe Asp Val Leu Gly Ala Ala Val Thr Pro Val Asp Gly Ala Leu Leu
20 25 30
Gly Asp Val Val Thr Val Glu Ala Ala Glu Thr Phe Ser Leu Asn Asn
35 40 45
Leu Gly Arg Phe Ala Asp Lys Leu Pro Ser Glu Pro Arg Glu Asn Ile
50 55 60
Val Tyr Gln Cys Trp Glu Arg Phe Cys Gln Glu Leu Gly Lys Gln Ile
65 70 75 80
Pro Val Ala Met Thr Leu Glu Lys Asn Met Pro Ile Gly Ser Gly Leu
85 90 95
Gly Ser Ser Ala Cys Ser Val Val Ala Ala Leu Met Ala Met Asn Glu
100 105 110
His Cys Gly Lys Pro Leu Asn Asp Thr Arg Leu Leu Ala Leu Met Gly
115 120 125
Glu Leu Glu Gly Arg Ile Ser Gly Ser Ile His Tyr Asp Asn Val Ala
130 135 140
Pro Cys Phe Leu Gly Gly Met Gln Leu Met Ile Glu Glu Asn Asp Ile
145 150 155 160
Ile Ser Gln Gln Val Pro Gly Phe Asp Glu Trp Leu Trp Val Leu Ala
165 170 175
Tyr Pro Gly Ile Lys Val Ser Thr Ala Glu Ala Arg Ala Ile Leu Pro
180 185 190
Ala Gln Tyr Arg Arg Gln Asp Cys Ile Ala His Gly Arg His Leu Ala
195 200 205
Gly Phe Ile His Ala Cys Tyr Ser Arg Gln Pro Glu Leu Ala Ala Lys
210 215 220
Leu Met Lys Asp Val Ile Ala Glu Pro Tyr Arg Glu Arg Leu Leu Pro
225 230 235 240
Gly Phe Arg Gln Ala Arg Gln Ala Val Ala Glu Ile Gly Ala Val Ala
245 250 255
Ser Gly Ile Ser Gly Ser Gly Pro Thr Leu Phe Ala Leu Cys Asp Lys
260 265 270
Pro Glu Thr Ala Gln Arg Val Ala Asp Trp Leu Gly Lys Asn Tyr Leu
275 280 285
Gln Asn Gln Glu Gly Phe Val His Ile Cys Arg Leu Asp Thr Ala Gly
290 295 300
Ala Arg Val Leu Glu Asn
305 310
<210> 12
<211> 933
<212> DNA
<213> 大肠杆菌
<220>
<221> 基因
<222> (1)..(933)
<223> thrB
<400> 12
atggttaaag tttatgcccc ggcttccagt gccaatatga gcgtcgggtt tgatgtgctc 60
ggggcggcgg tgacacctgt tgatggtgca ttgctcggag atgtagtcac ggttgaggcg 120
gcagagacat tcagtctcaa caacctcgga cgctttgccg ataagctgcc gtcagaacca 180
cgggaaaata tcgtttatca gtgctgggag cgtttttgcc aggaactggg taagcaaatt 240
ccagtggcga tgaccctgga aaagaatatg ccgatcggtt cgggcttagg ctccagtgcc 300
tgttcggtgg tcgcggcgct gatggcgatg aatgaacact gcggcaagcc gcttaatgac 360
actcgtttgc tggctttgat gggcgagctg gaaggccgta tctccggcag cattcattac 420
gacaacgtgg caccgtgttt tctcggtggt atgcagttga tgatcgaaga aaacgacatc 480
atcagccagc aagtgccagg gtttgatgag tggctgtggg tgctggcgta tccggggatt 540
aaagtctcga cggcagaagc cagggctatt ttaccggcgc agtatcgccg ccaggattgc 600
attgcgcacg ggcgacatct ggcaggcttc attcacgcct gctattcccg tcagcctgag 660
cttgccgcga agctgatgaa agatgttatc gctgaaccct accgtgaacg gttactgcca 720
ggcttccggc aggcgcggca ggcggtcgcg gaaatcggcg cggtagcgag cggtatctcc 780
ggctccggcc cgaccttgtt cgctctgtgt gacaagccgg aaaccgccca gcgcgttgcc 840
gactggttgg gtaagaacta cctgcaaaat caggaaggtt ttgttcatat ttgccggctg 900
gatacggcgg gcgcacgagt actggaaaac taa 933
<210> 13
<211> 379
<212> PRT
<213> 迈氏钩端螺旋体
<220>
<221> 肽
<222> (1)..(379)
<223> 高丝氨酸O-乙酰基转移酶
<400> 13
Met Pro Thr Ser Glu Gln Asn Glu Phe Ser His Gly Ser Val Gly Val
1 5 10 15
Val Tyr Thr Gln Ser Ile Arg Phe Glu Ser Leu Thr Leu Glu Gly Gly
20 25 30
Glu Thr Ile Thr Pro Leu Glu Ile Ala Tyr Glu Thr Tyr Gly Thr Leu
35 40 45
Asn Glu Lys Lys Asp Asn Ala Ile Leu Val Cys His Ala Leu Ser Gly
50 55 60
Asp Ala His Ala Ala Gly Phe His Glu Gly Asp Lys Arg Pro Gly Trp
65 70 75 80
Trp Asp Tyr Tyr Ile Gly Pro Gly Lys Ser Phe Asp Thr Asn Arg Tyr
85 90 95
Phe Ile Ile Ser Ser Asn Val Ile Gly Gly Cys Lys Gly Ser Ser Gly
100 105 110
Pro Leu Thr Ile Asn Gly Lys Asn Gly Lys Pro Phe Gln Ser Thr Phe
115 120 125
Pro Phe Val Ser Ile Gly Asp Met Val Asn Ala Gln Glu Lys Leu Ile
130 135 140
Ser His Phe Gly Ile His Lys Leu Phe Ala Val Ala Gly Gly Ser Met
145 150 155 160
Gly Gly Met Gln Ala Leu Gln Trp Ser Val Ala Tyr Pro Asp Arg Leu
165 170 175
Lys Asn Cys Ile Val Met Ala Ser Ser Ser Glu His Ser Ala Gln Gln
180 185 190
Ile Ala Phe Asn Glu Val Gly Arg Gln Ala Ile Leu Ser Asp Pro Asn
195 200 205
Trp Asn Gln Gly Leu Tyr Thr Gln Glu Asn Arg Pro Ser Lys Gly Leu
210 215 220
Ala Leu Ala Arg Met Met Gly His Ile Thr Tyr Leu Ser Asp Glu Met
225 230 235 240
Met Arg Glu Lys Phe Gly Arg Lys Pro Pro Lys Gly Asn Ile Gln Ser
245 250 255
Thr Asp Phe Ala Val Gly Ser Tyr Leu Ile Tyr Gln Gly Glu Ser Phe
260 265 270
Val Asp Arg Phe Asp Ala Asn Ser Tyr Ile Tyr Val Thr Lys Ala Leu
275 280 285
Asp His Phe Ser Leu Gly Thr Gly Lys Glu Leu Thr Lys Val Leu Ala
290 295 300
Lys Val Arg Cys Arg Phe Leu Val Val Ala Tyr Thr Ser Asp Trp Leu
305 310 315 320
Tyr Pro Pro Tyr Gln Ser Glu Glu Ile Val Lys Ser Leu Glu Val Asn
325 330 335
Ala Val Pro Val Ser Phe Val Glu Leu Asn Asn Pro Ala Gly Arg His
340 345 350
Asp Ser Phe Leu Leu Pro Ser Glu Gln Gln Asp Ser Ile Leu Arg Asp
355 360 365
Phe Leu Ser Ser Thr Asp Glu Gly Val Phe Leu
370 375
<210> 14
<211> 379
<212> PRT
<213> 谷氨酸棒状杆菌
<220>
<221> 肽
<222> (1)..(379)
<223> 高丝氨酸O-乙酰基转移酶
<400> 14
Met Pro Thr Leu Ala Pro Ser Gly Gln Leu Glu Ile Gln Ala Ile Gly
1 5 10 15
Asp Val Ser Thr Glu Ala Gly Ala Ile Ile Thr Asn Ala Glu Ile Ala
20 25 30
Tyr His Arg Trp Gly Glu Tyr Arg Val Asp Lys Glu Gly Arg Ser Asn
35 40 45
Val Val Leu Ile Glu His Ala Leu Thr Gly Asp Ser Asn Ala Ala Asp
50 55 60
Trp Trp Ala Asp Leu Leu Gly Pro Gly Lys Ala Ile Asn Thr Asp Ile
65 70 75 80
Tyr Cys Val Ile Cys Thr Asn Val Ile Gly Gly Cys Asn Gly Ser Thr
85 90 95
Gly Pro Gly Ser Met His Pro Asp Gly Asn Phe Trp Gly Asn Arg Phe
100 105 110
Pro Ala Thr Ser Ile Arg Asp Gln Val Asn Ala Glu Lys Gln Phe Leu
115 120 125
Asp Ala Leu Gly Ile Thr Thr Val Ala Ala Val Val Leu Leu Gly Gly
130 135 140
Ser Met Gly Gly Ala Arg Thr Leu Glu Trp Ala Ala Met Tyr Pro Glu
145 150 155 160
Thr Val Gly Ala Ala Ala Val Leu Ala Val Ser Ala Arg Ala Ser Ala
165 170 175
Trp Gln Ile Gly Ile Gln Ser Ala Gln Ile Lys Ala Ile Glu Asn Asp
180 185 190
His His Trp His Glu Gly Asn Tyr Tyr Glu Ser Gly Cys Asn Pro Ala
195 200 205
Thr Gly Leu Gly Ala Ala Arg Arg Ile Ala His Leu Thr Tyr Arg Gly
210 215 220
Glu Leu Glu Ile Asp Glu Arg Phe Gly Thr Lys Ala Gln Lys Asn Glu
225 230 235 240
Asn Pro Leu Gly Pro Tyr Arg Lys Pro Asp Gln Arg Phe Ala Val Glu
245 250 255
Ser Tyr Leu Asp Tyr Gln Ala Asp Lys Leu Val Gln Arg Phe Asp Ala
260 265 270
Gly Ser Tyr Val Leu Leu Thr Asp Ala Leu Asn Arg His Asp Ile Gly
275 280 285
Arg Asp Arg Gly Gly Leu Asn Lys Ala Leu Glu Ser Ile Lys Val Pro
290 295 300
Val Leu Val Ala Gly Val Asp Thr Asp Ile Leu Tyr Pro Tyr His Gln
305 310 315 320
Gln Glu His Leu Ser Arg Asn Leu Gly Asn Leu Leu Ala Met Ala Lys
325 330 335
Ile Val Ser Pro Val Gly His Asp Ala Phe Leu Thr Glu Ser Arg Gln
340 345 350
Met Asp Arg Ile Val Arg Asn Phe Phe Ser Leu Ile Ser Pro Asp Glu
355 360 365
Asp Asn Pro Ser Thr Tyr Ile Glu Phe Tyr Ile
370 375
<210> 15
<211> 334
<212> PRT
<213> 抗辐射奇异球菌
<220>
<221> 肽
<222> (1)..(334)
<223> 高丝氨酸O-乙酰基转移酶
<400> 15
Met Thr Ala Val Leu Ala Gly His Ala Ser Ala Leu Leu Leu Thr Glu
1 5 10 15
Glu Pro Asp Cys Ser Gly Pro Gln Thr Val Val Leu Phe Arg Arg Glu
20 25 30
Pro Leu Leu Leu Asp Cys Gly Arg Ala Leu Ser Asp Val Arg Val Ala
35 40 45
Phe His Thr Tyr Gly Thr Pro Arg Ala Asp Ala Thr Leu Val Leu His
50 55 60
Ala Leu Thr Gly Asp Ser Ala Val His Glu Trp Trp Pro Asp Phe Leu
65 70 75 80
Gly Ala Gly Arg Pro Leu Asp Pro Ala Asp Asp Tyr Val Val Cys Ala
85 90 95
Asn Val Leu Gly Gly Cys Ala Gly Thr Thr Ser Ala Ala Glu Leu Ala
100 105 110
Ala Thr Cys Ser Gly Pro Val Pro Leu Ser Leu Arg Asp Met Ala Arg
115 120 125
Val Gly Arg Ala Leu Leu Asp Ser Leu Gly Val Arg Arg Val Arg Val
130 135 140
Ile Gly Ala Ser Met Gly Gly Met Leu Ala Tyr Ala Trp Leu Leu Glu
145 150 155 160
Cys Pro Asp Leu Val Glu Lys Ala Val Ile Ile Gly Ala Pro Ala Arg
165 170 175
His Ser Pro Trp Ala Ile Gly Leu Asn Thr Ala Ala Arg Ser Ala Ile
180 185 190
Ala Leu Ala Pro Gly Gly Glu Gly Leu Lys Val Ala Arg Gln Ile Ala
195 200 205
Met Leu Ser Tyr Arg Ser Pro Glu Ser Leu Ser Arg Thr Gln Ala Gly
210 215 220
Gln Arg Val Pro Gly Val Pro Ala Val Thr Ser Tyr Leu His Tyr Gln
225 230 235 240
Gly Glu Lys Leu Ala Ala Arg Phe Asp Glu Gln Thr Tyr Cys Ala Leu
245 250 255
Thr Trp Ala Met Asp Ala Phe Gln Pro Ser Ser Ala Asp Leu Lys Ala
260 265 270
Val Arg Ala Pro Val Leu Val Val Gly Ile Ser Ser Asp Leu Leu Tyr
275 280 285
Pro Ala Ala Glu Val Arg Ala Cys Ala Ala Glu Leu Pro His Ala Asp
290 295 300
Tyr Trp Glu Leu Gly Ser Ile His Gly His Asp Ala Phe Leu Met Asp
305 310 315 320
Pro Gln Asp Leu Pro Glu Arg Val Gly Ala Phe Leu Arg Ser
325 330
<210> 16
<211> 309
<212> PRT
<213> 大肠杆菌
<220>
<221> 肽
<222> (1)..(309)
<223> 高丝氨酸O-琥珀酰基转移酶
<400> 16
Met Pro Ile Arg Val Pro Asp Glu Leu Pro Ala Val Asn Phe Leu Arg
1 5 10 15
Glu Glu Asn Val Phe Val Met Thr Thr Ser Arg Ala Ser Gly Gln Glu
20 25 30
Ile Arg Pro Leu Lys Val Leu Ile Leu Asn Leu Met Pro Lys Lys Ile
35 40 45
Glu Thr Glu Asn Gln Phe Leu Arg Leu Leu Ser Asn Ser Pro Leu Gln
50 55 60
Val Asp Ile Gln Leu Leu Arg Ile Asp Ser Arg Glu Ser Arg Asn Thr
65 70 75 80
Pro Ala Glu His Leu Asn Asn Phe Tyr Cys Asn Phe Glu Asp Ile Gln
85 90 95
Asp Gln Asn Phe Asp Gly Leu Ile Val Thr Gly Ala Pro Leu Gly Leu
100 105 110
Val Glu Phe Asn Asp Val Ala Tyr Trp Pro Gln Ile Lys Gln Val Leu
115 120 125
Glu Trp Ser Lys Asp His Val Thr Ser Thr Leu Phe Val Cys Trp Ala
130 135 140
Val Gln Ala Ala Leu Asn Ile Leu Tyr Gly Ile Pro Lys Gln Thr Arg
145 150 155 160
Thr Glu Lys Leu Ser Gly Val Tyr Glu His His Ile Leu His Pro His
165 170 175
Ala Leu Leu Thr Arg Gly Phe Asp Asp Ser Phe Leu Ala Pro His Ser
180 185 190
Arg Tyr Ala Asp Phe Pro Ala Ala Leu Ile Arg Asp Tyr Thr Asp Leu
195 200 205
Glu Ile Leu Ala Glu Thr Glu Glu Gly Asp Ala Tyr Leu Phe Ala Ser
210 215 220
Lys Asp Lys Arg Ile Ala Phe Val Thr Gly His Pro Glu Tyr Asp Ala
225 230 235 240
Gln Thr Leu Ala Gln Glu Phe Phe Arg Asp Val Glu Ala Gly Leu Asp
245 250 255
Pro Asp Val Pro Tyr Asn Tyr Phe Pro His Asn Asp Pro Gln Asn Thr
260 265 270
Pro Arg Ala Ser Trp Arg Ser His Gly Asn Leu Leu Phe Thr Asn Trp
275 280 285
Leu Asn Tyr Tyr Val Tyr Gln Ile Thr Pro Tyr Asp Leu Arg His Met
290 295 300
Asn Pro Thr Leu Asp
305
<210> 17
<211> 309
<212> PRT
<213> 人工序列
<220>
<223> 具有高丝氨酸O-乙酰基转移酶活性的变体多肽
metA EL
<400> 17
Met Pro Ile Arg Val Pro Asp Glu Leu Pro Ala Val Asn Phe Leu Arg
1 5 10 15
Glu Glu Asn Val Phe Val Met Thr Thr Ser Arg Ala Ser Gly Gln Glu
20 25 30
Ile Arg Pro Leu Lys Val Leu Ile Leu Asn Leu Met Pro Lys Lys Ile
35 40 45
Glu Thr Glu Asn Gln Phe Leu Arg Leu Leu Ser Asn Ser Pro Leu Gln
50 55 60
Val Asp Ile Gln Leu Leu Arg Ile Asp Ser Arg Glu Ser Arg Asn Thr
65 70 75 80
Pro Ala Glu His Leu Asn Asn Phe Tyr Cys Asn Phe Glu Asp Ile Gln
85 90 95
Asp Gln Asn Phe Asp Gly Leu Ile Val Thr Gly Ala Pro Leu Glu Leu
100 105 110
Val Glu Phe Asn Asp Val Ala Tyr Trp Pro Gln Ile Lys Gln Val Leu
115 120 125
Glu Trp Ser Lys Asp His Val Thr Ser Thr Leu Phe Val Cys Trp Ala
130 135 140
Val Gln Ala Ala Leu Asn Ile Leu Tyr Gly Ile Pro Lys Gln Thr Arg
145 150 155 160
Thr Glu Lys Leu Ser Gly Val Tyr Glu His His Ile Leu His Pro His
165 170 175
Ala Leu Leu Thr Arg Gly Phe Asp Asp Ser Phe Leu Ala Pro His Ser
180 185 190
Arg Tyr Ala Asp Phe Pro Ala Ala Leu Ile Arg Asp Tyr Thr Asp Leu
195 200 205
Glu Ile Leu Ala Glu Thr Glu Glu Gly Asp Ala Tyr Leu Phe Ala Ser
210 215 220
Lys Asp Lys Arg Ile Ala Phe Val Thr Gly His Pro Glu Tyr Asp Ala
225 230 235 240
Gln Thr Leu Ala Gln Glu Phe Phe Arg Asp Val Glu Ala Gly Leu Asp
245 250 255
Pro Asp Val Pro Tyr Asn Tyr Phe Pro His Asn Asp Pro Gln Asn Thr
260 265 270
Pro Arg Ala Ser Trp Arg Ser His Gly Asn Leu Leu Phe Thr Asn Trp
275 280 285
Leu Asn Tyr Tyr Val Tyr Gln Ile Thr Pro Tyr Asp Leu Arg His Met
290 295 300
Asn Pro Thr Leu Asp
305
<210> 18
<211> 309
<212> PRT
<213> 人工序列
<220>
<223> 具有高丝氨酸O-乙酰基转移酶活性的变体多肽
metA ET
<400> 18
Met Pro Ile Arg Val Pro Asp Glu Leu Pro Ala Val Asn Phe Leu Arg
1 5 10 15
Glu Glu Asn Val Phe Val Met Thr Thr Ser Arg Ala Ser Gly Gln Glu
20 25 30
Ile Arg Pro Leu Lys Val Leu Ile Leu Asn Leu Met Pro Lys Lys Ile
35 40 45
Glu Thr Glu Asn Gln Phe Leu Arg Leu Leu Ser Asn Ser Pro Leu Gln
50 55 60
Val Asp Ile Gln Leu Leu Arg Ile Asp Ser Arg Glu Ser Arg Asn Thr
65 70 75 80
Pro Ala Glu His Leu Asn Asn Phe Tyr Cys Asn Phe Glu Asp Ile Gln
85 90 95
Asp Gln Asn Phe Asp Gly Leu Ile Val Thr Gly Ala Pro Leu Glu Thr
100 105 110
Val Glu Phe Asn Asp Val Ala Tyr Trp Pro Gln Ile Lys Gln Val Leu
115 120 125
Glu Trp Ser Lys Asp His Val Thr Ser Thr Leu Phe Val Cys Trp Ala
130 135 140
Val Gln Ala Ala Leu Asn Ile Leu Tyr Gly Ile Pro Lys Gln Thr Arg
145 150 155 160
Thr Glu Lys Leu Ser Gly Val Tyr Glu His His Ile Leu His Pro His
165 170 175
Ala Leu Leu Thr Arg Gly Phe Asp Asp Ser Phe Leu Ala Pro His Ser
180 185 190
Arg Tyr Ala Asp Phe Pro Ala Ala Leu Ile Arg Asp Tyr Thr Asp Leu
195 200 205
Glu Ile Leu Ala Glu Thr Glu Glu Gly Asp Ala Tyr Leu Phe Ala Ser
210 215 220
Lys Asp Lys Arg Ile Ala Phe Val Thr Gly His Pro Glu Tyr Asp Ala
225 230 235 240
Gln Thr Leu Ala Gln Glu Phe Phe Arg Asp Val Glu Ala Gly Leu Asp
245 250 255
Pro Asp Val Pro Tyr Asn Tyr Phe Pro His Asn Asp Pro Gln Asn Thr
260 265 270
Pro Arg Ala Ser Trp Arg Ser His Gly Asn Leu Leu Phe Thr Asn Trp
275 280 285
Leu Asn Tyr Tyr Val Tyr Gln Ile Thr Pro Tyr Asp Leu Arg His Met
290 295 300
Asn Pro Thr Leu Asp
305
<210> 19
<211> 309
<212> PRT
<213> 人工序列
<220>
<223> 具有高丝氨酸O-乙酰基转移酶活性的变体多肽
metA EH
<400> 19
Met Pro Ile Arg Val Pro Asp Glu Leu Pro Ala Val Asn Phe Leu Arg
1 5 10 15
Glu Glu Asn Val Phe Val Met Thr Thr Ser Arg Ala Ser Gly Gln Glu
20 25 30
Ile Arg Pro Leu Lys Val Leu Ile Leu Asn Leu Met Pro Lys Lys Ile
35 40 45
Glu Thr Glu Asn Gln Phe Leu Arg Leu Leu Ser Asn Ser Pro Leu Gln
50 55 60
Val Asp Ile Gln Leu Leu Arg Ile Asp Ser Arg Glu Ser Arg Asn Thr
65 70 75 80
Pro Ala Glu His Leu Asn Asn Phe Tyr Cys Asn Phe Glu Asp Ile Gln
85 90 95
Asp Gln Asn Phe Asp Gly Leu Ile Val Thr Gly Ala Pro Leu Glu His
100 105 110
Val Glu Phe Asn Asp Val Ala Tyr Trp Pro Gln Ile Lys Gln Val Leu
115 120 125
Glu Trp Ser Lys Asp His Val Thr Ser Thr Leu Phe Val Cys Trp Ala
130 135 140
Val Gln Ala Ala Leu Asn Ile Leu Tyr Gly Ile Pro Lys Gln Thr Arg
145 150 155 160
Thr Glu Lys Leu Ser Gly Val Tyr Glu His His Ile Leu His Pro His
165 170 175
Ala Leu Leu Thr Arg Gly Phe Asp Asp Ser Phe Leu Ala Pro His Ser
180 185 190
Arg Tyr Ala Asp Phe Pro Ala Ala Leu Ile Arg Asp Tyr Thr Asp Leu
195 200 205
Glu Ile Leu Ala Glu Thr Glu Glu Gly Asp Ala Tyr Leu Phe Ala Ser
210 215 220
Lys Asp Lys Arg Ile Ala Phe Val Thr Gly His Pro Glu Tyr Asp Ala
225 230 235 240
Gln Thr Leu Ala Gln Glu Phe Phe Arg Asp Val Glu Ala Gly Leu Asp
245 250 255
Pro Asp Val Pro Tyr Asn Tyr Phe Pro His Asn Asp Pro Gln Asn Thr
260 265 270
Pro Arg Ala Ser Trp Arg Ser His Gly Asn Leu Leu Phe Thr Asn Trp
275 280 285
Leu Asn Tyr Tyr Val Tyr Gln Ile Thr Pro Tyr Asp Leu Arg His Met
290 295 300
Asn Pro Thr Leu Asp
305
<210> 20
<211> 883
<212> PRT
<213> 大肠杆菌
<220>
<221> 肽
<222> (1)..(883)
<223> 磷酸烯醇丙酮酸羧化酶
<400> 20
Met Asn Glu Gln Tyr Ser Ala Leu Arg Ser Asn Val Ser Met Leu Gly
1 5 10 15
Lys Val Leu Gly Glu Thr Ile Lys Asp Ala Leu Gly Glu His Ile Leu
20 25 30
Glu Arg Val Glu Thr Ile Arg Lys Leu Ser Lys Ser Ser Arg Ala Gly
35 40 45
Asn Asp Ala Asn Arg Gln Glu Leu Leu Thr Thr Leu Gln Asn Leu Ser
50 55 60
Asn Asp Glu Leu Leu Pro Val Ala Arg Ala Phe Ser Gln Phe Leu Asn
65 70 75 80
Leu Ala Asn Thr Ala Glu Gln Tyr His Ser Ile Ser Pro Lys Gly Glu
85 90 95
Ala Ala Ser Asn Pro Glu Val Ile Ala Arg Thr Leu Arg Lys Leu Lys
100 105 110
Asn Gln Pro Glu Leu Ser Glu Asp Thr Ile Lys Lys Ala Val Glu Ser
115 120 125
Leu Ser Leu Glu Leu Val Leu Thr Ala His Pro Thr Glu Ile Thr Arg
130 135 140
Arg Thr Leu Ile His Lys Met Val Glu Val Asn Ala Cys Leu Lys Gln
145 150 155 160
Leu Asp Asn Lys Asp Ile Ala Asp Tyr Glu His Asn Gln Leu Met Arg
165 170 175
Arg Leu Arg Gln Leu Ile Ala Gln Ser Trp His Thr Asp Glu Ile Arg
180 185 190
Lys Leu Arg Pro Ser Pro Val Asp Glu Ala Lys Trp Gly Phe Ala Val
195 200 205
Val Glu Asn Ser Leu Trp Gln Gly Val Pro Asn Tyr Leu Arg Glu Leu
210 215 220
Asn Glu Gln Leu Glu Glu Asn Leu Gly Tyr Lys Leu Pro Val Glu Phe
225 230 235 240
Val Pro Val Arg Phe Thr Ser Trp Met Gly Gly Asp Arg Asp Gly Asn
245 250 255
Pro Asn Val Thr Ala Asp Ile Thr Arg His Val Leu Leu Leu Ser Arg
260 265 270
Trp Lys Ala Thr Asp Leu Phe Leu Lys Asp Ile Gln Val Leu Val Ser
275 280 285
Glu Leu Ser Met Val Glu Ala Thr Pro Glu Leu Leu Ala Leu Val Gly
290 295 300
Glu Glu Gly Ala Ala Glu Pro Tyr Arg Tyr Leu Met Lys Asn Leu Arg
305 310 315 320
Ser Arg Leu Met Ala Thr Gln Ala Trp Leu Glu Ala Arg Leu Lys Gly
325 330 335
Glu Glu Leu Pro Lys Pro Glu Gly Leu Leu Thr Gln Asn Glu Glu Leu
340 345 350
Trp Glu Pro Leu Tyr Ala Cys Tyr Gln Ser Leu Gln Ala Cys Gly Met
355 360 365
Gly Ile Ile Ala Asn Gly Asp Leu Leu Asp Thr Leu Arg Arg Val Lys
370 375 380
Cys Phe Gly Val Pro Leu Val Arg Ile Asp Ile Arg Gln Glu Ser Thr
385 390 395 400
Arg His Thr Glu Ala Leu Gly Glu Leu Thr Arg Tyr Leu Gly Ile Gly
405 410 415
Asp Tyr Glu Ser Trp Ser Glu Ala Asp Lys Gln Ala Phe Leu Ile Arg
420 425 430
Glu Leu Asn Ser Lys Arg Pro Leu Leu Pro Arg Asn Trp Gln Pro Ser
435 440 445
Ala Glu Thr Arg Glu Val Leu Asp Thr Cys Gln Val Ile Ala Glu Ala
450 455 460
Pro Gln Gly Ser Ile Ala Ala Tyr Val Ile Ser Met Ala Lys Thr Pro
465 470 475 480
Ser Asp Val Leu Ala Val His Leu Leu Leu Lys Glu Ala Gly Ile Gly
485 490 495
Phe Ala Met Pro Val Ala Pro Leu Phe Glu Thr Leu Asp Asp Leu Asn
500 505 510
Asn Ala Asn Asp Val Met Thr Gln Leu Leu Asn Ile Asp Trp Tyr Arg
515 520 525
Gly Leu Ile Gln Gly Lys Gln Met Val Met Ile Gly Tyr Ser Asp Ser
530 535 540
Ala Lys Asp Ala Gly Val Met Ala Ala Ser Trp Ala Gln Tyr Gln Ala
545 550 555 560
Gln Asp Ala Leu Ile Lys Thr Cys Glu Lys Ala Gly Ile Glu Leu Thr
565 570 575
Leu Phe His Gly Arg Gly Gly Ser Ile Gly Arg Gly Gly Ala Pro Ala
580 585 590
His Ala Ala Leu Leu Ser Gln Pro Pro Gly Ser Leu Lys Gly Gly Leu
595 600 605
Arg Val Thr Glu Gln Gly Glu Met Ile Arg Phe Lys Tyr Gly Leu Pro
610 615 620
Glu Ile Thr Val Ser Ser Leu Ser Leu Tyr Thr Gly Ala Ile Leu Glu
625 630 635 640
Ala Asn Leu Leu Pro Pro Pro Glu Pro Lys Glu Ser Trp Arg Arg Ile
645 650 655
Met Asp Glu Leu Ser Val Ile Ser Cys Asp Val Tyr Arg Gly Tyr Val
660 665 670
Arg Glu Asn Lys Asp Phe Val Pro Tyr Phe Arg Ser Ala Thr Pro Glu
675 680 685
Gln Glu Leu Gly Lys Leu Pro Leu Gly Ser Arg Pro Ala Lys Arg Arg
690 695 700
Pro Thr Gly Gly Val Glu Ser Leu Arg Ala Ile Pro Trp Ile Phe Ala
705 710 715 720
Trp Thr Gln Asn Arg Leu Met Leu Pro Ala Trp Leu Gly Ala Gly Thr
725 730 735
Ala Leu Gln Lys Val Val Glu Asp Gly Lys Gln Ser Glu Leu Glu Ala
740 745 750
Met Cys Arg Asp Trp Pro Phe Phe Ser Thr Arg Leu Gly Met Leu Glu
755 760 765
Met Val Phe Ala Lys Ala Asp Leu Trp Leu Ala Glu Tyr Tyr Asp Gln
770 775 780
Arg Leu Val Asp Lys Ala Leu Trp Pro Leu Gly Lys Glu Leu Arg Asn
785 790 795 800
Leu Gln Glu Glu Asp Ile Lys Val Val Leu Ala Ile Ala Asn Asp Ser
805 810 815
His Leu Met Ala Asp Leu Pro Trp Ile Ala Glu Ser Ile Gln Leu Arg
820 825 830
Asn Ile Tyr Thr Asp Pro Leu Asn Val Leu Gln Ala Glu Leu Leu His
835 840 845
Arg Ser Arg Gln Ala Glu Lys Glu Gly Gln Glu Pro Asp Pro Arg Val
850 855 860
Glu Gln Ala Leu Met Val Thr Ile Ala Gly Ile Ala Ala Gly Met Arg
865 870 875 880
Asn Thr Gly
<210> 21
<211> 396
<212> PRT
<213> 大肠杆菌
<220>
<221> 肽
<222> (1)..(396)
<223> 天冬氨酸氨基转移酶
<400> 21
Met Phe Glu Asn Ile Thr Ala Ala Pro Ala Asp Pro Ile Leu Gly Leu
1 5 10 15
Ala Asp Leu Phe Arg Ala Asp Glu Arg Pro Gly Lys Ile Asn Leu Gly
20 25 30
Ile Gly Val Tyr Lys Asp Glu Thr Gly Lys Thr Pro Val Leu Thr Ser
35 40 45
Val Lys Lys Ala Glu Gln Tyr Leu Leu Glu Asn Glu Thr Thr Lys Asn
50 55 60
Tyr Leu Gly Ile Asp Gly Ile Pro Glu Phe Gly Arg Cys Thr Gln Glu
65 70 75 80
Leu Leu Phe Gly Lys Gly Ser Ala Leu Ile Asn Asp Lys Arg Ala Arg
85 90 95
Thr Ala Gln Thr Pro Gly Gly Thr Gly Ala Leu Arg Val Ala Ala Asp
100 105 110
Phe Leu Ala Lys Asn Thr Ser Val Lys Arg Val Trp Val Ser Asn Pro
115 120 125
Ser Trp Pro Asn His Lys Ser Val Phe Asn Ser Ala Gly Leu Glu Val
130 135 140
Arg Glu Tyr Ala Tyr Tyr Asp Ala Glu Asn His Thr Leu Asp Phe Asp
145 150 155 160
Ala Leu Ile Asn Ser Leu Asn Glu Ala Gln Ala Gly Asp Val Val Leu
165 170 175
Phe His Gly Cys Cys His Asn Pro Thr Gly Ile Asp Pro Thr Leu Glu
180 185 190
Gln Trp Gln Thr Leu Ala Gln Leu Ser Val Glu Lys Gly Trp Leu Pro
195 200 205
Leu Phe Asp Phe Ala Tyr Gln Gly Phe Ala Arg Gly Leu Glu Glu Asp
210 215 220
Ala Glu Gly Leu Arg Ala Phe Ala Ala Met His Lys Glu Leu Ile Val
225 230 235 240
Ala Ser Ser Tyr Ser Lys Asn Phe Gly Leu Tyr Asn Glu Arg Val Gly
245 250 255
Ala Cys Thr Leu Val Ala Ala Asp Ser Glu Thr Val Asp Arg Ala Phe
260 265 270
Ser Gln Met Lys Ala Ala Ile Arg Ala Asn Tyr Ser Asn Pro Pro Ala
275 280 285
His Gly Ala Ser Val Val Ala Thr Ile Leu Ser Asn Asp Ala Leu Arg
290 295 300
Ala Ile Trp Glu Gln Glu Leu Thr Asp Met Arg Gln Arg Ile Gln Arg
305 310 315 320
Met Arg Gln Leu Phe Val Asn Thr Leu Gln Glu Lys Gly Ala Asn Arg
325 330 335
Asp Phe Ser Phe Ile Ile Lys Gln Asn Gly Met Phe Ser Phe Ser Gly
340 345 350
Leu Thr Lys Glu Gln Val Leu Arg Leu Arg Glu Glu Phe Gly Val Tyr
355 360 365
Ala Val Ala Ser Gly Arg Val Asn Val Ala Gly Met Thr Pro Asp Asn
370 375 380
Met Ala Pro Leu Cys Glu Ala Ile Val Ala Val Leu
385 390 395
<210> 22
<211> 367
<212> PRT
<213> 大肠杆菌
<220>
<221> 肽
<222> (1)..(367)
<223> 天冬氨酸半醛脱氢酶
<400> 22
Met Lys Asn Val Gly Phe Ile Gly Trp Arg Gly Met Val Gly Ser Val
1 5 10 15
Leu Met Gln Arg Met Val Glu Glu Arg Asp Phe Asp Ala Ile Arg Pro
20 25 30
Val Phe Phe Ser Thr Ser Gln Leu Gly Gln Ala Ala Pro Ser Phe Gly
35 40 45
Gly Thr Thr Gly Thr Leu Gln Asp Ala Phe Asp Leu Glu Ala Leu Lys
50 55 60
Ala Leu Asp Ile Ile Val Thr Cys Gln Gly Gly Asp Tyr Thr Asn Glu
65 70 75 80
Ile Tyr Pro Lys Leu Arg Glu Ser Gly Trp Gln Gly Tyr Trp Ile Asp
85 90 95
Ala Ala Ser Ser Leu Arg Met Lys Asp Asp Ala Ile Ile Ile Leu Asp
100 105 110
Pro Val Asn Gln Asp Val Ile Thr Asp Gly Leu Asn Asn Gly Ile Arg
115 120 125
Thr Phe Val Gly Gly Asn Cys Thr Val Ser Leu Met Leu Met Ser Leu
130 135 140
Gly Gly Leu Phe Ala Asn Asp Leu Val Asp Trp Val Ser Val Ala Thr
145 150 155 160
Tyr Gln Ala Ala Ser Gly Gly Gly Ala Arg His Met Arg Glu Leu Leu
165 170 175
Thr Gln Met Gly His Leu Tyr Gly His Val Ala Asp Glu Leu Ala Thr
180 185 190
Pro Ser Ser Ala Ile Leu Asp Ile Glu Arg Lys Val Thr Thr Leu Thr
195 200 205
Arg Ser Gly Glu Leu Pro Val Asp Asn Phe Gly Val Pro Leu Ala Gly
210 215 220
Ser Leu Ile Pro Trp Ile Asp Lys Gln Leu Asp Asn Gly Gln Ser Arg
225 230 235 240
Glu Glu Trp Lys Gly Gln Ala Glu Thr Asn Lys Ile Leu Asn Thr Ser
245 250 255
Ser Val Ile Pro Val Asp Gly Leu Cys Val Arg Val Gly Ala Leu Arg
260 265 270
Cys His Ser Gln Ala Phe Thr Ile Lys Leu Lys Lys Asp Val Ser Ile
275 280 285
Pro Thr Val Glu Glu Leu Leu Ala Ala His Asn Pro Trp Ala Lys Val
290 295 300
Val Pro Asn Asp Arg Glu Ile Thr Met Arg Glu Leu Thr Pro Ala Ala
305 310 315 320
Val Thr Gly Thr Leu Thr Thr Pro Val Gly Arg Leu Arg Lys Leu Asn
325 330 335
Met Gly Pro Glu Phe Leu Ser Ala Phe Thr Val Gly Asp Gln Leu Leu
340 345 350
Trp Gly Ala Ala Glu Pro Leu Arg Arg Met Leu Arg Gln Leu Ala
355 360 365
<210> 23
<211> 933
<212> DNA
<213> 大肠杆菌
<220>
<221> 基因
<222> (1)..(933)
<223> 高丝氨酸O-琥珀酰基转移酶
<400> 23
atggttaaag tttatgcccc ggcttccagt gccaatatga gcgtcgggtt tgatgtgctc 60
ggggcggcgg tgacacctgt tgatggtgca ttgctcggag atgtagtcac ggttgaggcg 120
gcagagacat tcagtctcaa caacctcgga cgctttgccg ataagctgcc gtcagaacca 180
cgggaaaata tcgtttatca gtgctgggag cgtttttgcc aggaactggg taagcaaatt 240
ccagtggcga tgaccctgga aaagaatatg ccgatcggtt cgggcttagg ctccagtgcc 300
tgttcggtgg tcgcggcgct gatggcgatg aatgaacact gcggcaagcc gcttaatgac 360
actcgtttgc tggctttgat gggcgagctg gaaggccgta tctccggcag cattcattac 420
gacaacgtgg caccgtgttt tctcggtggt atgcagttga tgatcgaaga aaacgacatc 480
atcagccagc aagtgccagg gtttgatgag tggctgtggg tgctggcgta tccggggatt 540
aaagtctcga cggcagaagc cagggctatt ttaccggcgc agtatcgccg ccaggattgc 600
attgcgcacg ggcgacatct ggcaggcttc attcacgcct gctattcccg tcagcctgag 660
cttgccgcga agctgatgaa agatgttatc gctgaaccct accgtgaacg gttactgcca 720
ggcttccggc aggcgcggca ggcggtcgcg gaaatcggcg cggtagcgag cggtatctcc 780
ggctccggcc cgaccttgtt cgctctgtgt gacaagccgg aaaccgccca gcgcgttgcc 840
gactggttgg gtaagaacta cctgcaaaat caggaaggtt ttgttcatat ttgccggctg 900
gatacggcgg gcgcacgagt actggaaaac taa 933
<210> 24
<211> 930
<212> DNA
<213> 人工序列
<220>
<223> 具有高丝氨酸O-乙酰基转移酶活性的变体多肽
metA EL
<400> 24
atgccgattc gtgtgccgga cgagctaccc gccgtcaatt tcttgcgtga agaaaacgtc 60
tttgtgatga caacttctcg tgcgtctggt caggaaattc gtccacttaa ggttctgatc 120
cttaacctga tgccgaagaa gattgaaact gaaaatcagt ttctgcgcct gctttcaaac 180
tcacctttgc aggtcgatat tcagctgttg cgcatcgatt cccgtgaatc gcgcaacacg 240
cccgcagagc atctgaacaa cttctactgt aactttgaag atattcagga tcagaacttt 300
gacggtttga ttgtaactgg tgcgccgctg gaactggtgg agtttaatga tgtcgcttac 360
tggccgcaga tcaaacaggt gctggagtgg tcgaaagatc acgtcacctc gacgctgttt 420
gtctgctggg cggtacaggc cgcgctcaat atcctctacg gcattcctaa gcaaactcgc 480
accgaaaaac tctctggcgt ttacgagcat catattctcc atcctcatgc gcttctgacg 540
cgtggctttg atgattcatt cctggcaccg cattcgcgct atgctgactt tccggcagcg 600
ttgattcgtg attacaccga tctggaaatt ctggcagaga cggaagaagg ggatgcatat 660
ctgtttgcca gtaaagataa gcgcattgcc tttgtgacgg gccatcccga atatgatgcg 720
caaacgctgg cgcaggaatt tttccgcgat gtggaagccg gactagaccc ggatgtaccg 780
tataactatt tcccgcacaa tgatccgcaa aatacaccgc gagcgagctg gcgtagtcac 840
ggtaatttac tgtttaccaa ctggctcaac tattacgtct accagatcac gccatacgat 900
ctacggcaca tgaatccaac gctggattaa 930
<210> 25
<211> 930
<212> DNA
<213> 人工序列
<220>
<223> 具有高丝氨酸O-乙酰基转移酶活性的变体多肽
metA ET
<400> 25
atgccgattc gtgtgccgga cgagctaccc gccgtcaatt tcttgcgtga agaaaacgtc 60
tttgtgatga caacttctcg tgcgtctggt caggaaattc gtccacttaa ggttctgatc 120
cttaacctga tgccgaagaa gattgaaact gaaaatcagt ttctgcgcct gctttcaaac 180
tcacctttgc aggtcgatat tcagctgttg cgcatcgatt cccgtgaatc gcgcaacacg 240
cccgcagagc atctgaacaa cttctactgt aactttgaag atattcagga tcagaacttt 300
gacggtttga ttgtaactgg tgcgccgctg gaaaccgtgg agtttaatga tgtcgcttac 360
tggccgcaga tcaaacaggt gctggagtgg tcgaaagatc acgtcacctc gacgctgttt 420
gtctgctggg cggtacaggc cgcgctcaat atcctctacg gcattcctaa gcaaactcgc 480
accgaaaaac tctctggcgt ttacgagcat catattctcc atcctcatgc gcttctgacg 540
cgtggctttg atgattcatt cctggcaccg cattcgcgct atgctgactt tccggcagcg 600
ttgattcgtg attacaccga tctggaaatt ctggcagaga cggaagaagg ggatgcatat 660
ctgtttgcca gtaaagataa gcgcattgcc tttgtgacgg gccatcccga atatgatgcg 720
caaacgctgg cgcaggaatt tttccgcgat gtggaagccg gactagaccc ggatgtaccg 780
tataactatt tcccgcacaa tgatccgcaa aatacaccgc gagcgagctg gcgtagtcac 840
ggtaatttac tgtttaccaa ctggctcaac tattacgtct accagatcac gccatacgat 900
ctacggcaca tgaatccaac gctggattaa 930
<210> 26
<211> 930
<212> DNA
<213> 人工序列
<220>
<223> 具有高丝氨酸O-乙酰基转移酶活性的变体多肽
metA EH
<400> 26
atgccgattc gtgtgccgga cgagctaccc gccgtcaatt tcttgcgtga agaaaacgtc 60
tttgtgatga caacttctcg tgcgtctggt caggaaattc gtccacttaa ggttctgatc 120
cttaacctga tgccgaagaa gattgaaact gaaaatcagt ttctgcgcct gctttcaaac 180
tcacctttgc aggtcgatat tcagctgttg cgcatcgatt cccgtgaatc gcgcaacacg 240
cccgcagagc atctgaacaa cttctactgt aactttgaag atattcagga tcagaacttt 300
gacggtttga ttgtaactgg tgcgccgctg gaacatgtgg agtttaatga tgtcgcttac 360
tggccgcaga tcaaacaggt gctggagtgg tcgaaagatc acgtcacctc gacgctgttt 420
gtctgctggg cggtacaggc cgcgctcaat atcctctacg gcattcctaa gcaaactcgc 480
accgaaaaac tctctggcgt ttacgagcat catattctcc atcctcatgc gcttctgacg 540
cgtggctttg atgattcatt cctggcaccg cattcgcgct atgctgactt tccggcagcg 600
ttgattcgtg attacaccga tctggaaatt ctggcagaga cggaagaagg ggatgcatat 660
ctgtttgcca gtaaagataa gcgcattgcc tttgtgacgg gccatcccga atatgatgcg 720
caaacgctgg cgcaggaatt tttccgcgat gtggaagccg gactagaccc ggatgtaccg 780
tataactatt tcccgcacaa tgatccgcaa aatacaccgc gagcgagctg gcgtagtcac 840
ggtaatttac tgtttaccaa ctggctcaac tattacgtct accagatcac gccatacgat 900
ctacggcaca tgaatccaac gctggattaa 930
<210> 27
<211> 2652
<212> DNA
<213> 大肠杆菌
<220>
<221> 基因
<222> (1)..(2652)
<223> 磷酸烯醇丙酮酸羧化酶
<400> 27
atgaacgaac aatattccgc attgcgtagt aatgtcagta tgctcggcaa agtgctggga 60
gaaaccatca aggatgcgtt gggagaacac attcttgaac gcgtagaaac tatccgtaag 120
ttgtcgaaat cttcacgcgc tggcaatgat gctaaccgcc aggagttgct caccacctta 180
caaaatttgt cgaacgacga gctgctgccc gttgcgcgtg cgtttagtca gttcctgaac 240
ctggccaaca ccgccgagca ataccacagc atttcgccga aaggcgaagc tgccagcaac 300
ccggaagtga tcgcccgcac cctgcgtaaa ctgaaaaacc agccggaact gagcgaagac 360
accatcaaaa aagcagtgga atcgctgtcg ctggaactgg tcctcacggc tcacccaacc 420
gaaattaccc gtcgtacact gatccacaaa atggtggaag tgaacgcctg tttaaaacag 480
ctcgataaca aagatatcgc tgactacgaa cacaaccagc tgatgcgtcg cctgcgccag 540
ttgatcgccc agtcatggca taccgatgaa atccgtaagc tgcgtccaag cccggtagat 600
gaagccaaat ggggctttgc cgtagtggaa aacagcctgt ggcaaggcgt accaaattac 660
ctgcgcgaac tgaacgaaca actggaagag aacctcggct acaaactgcc cgtcgaattt 720
gttccggtcc gttttacttc gtggatgggc ggcgaccgcg acggcaaccc gaacgtcact 780
gccgatatca cccgccacgt cctgctactc agccgctgga aagccaccga tttgttcctg 840
aaagatattc aggtgctggt ttctgaactg tcgatggttg aagcgacccc tgaactgctg 900
gcgctggttg gcgaagaagg tgccgcagaa ccgtatcgct atctgatgaa aaacctgcgt 960
tctcgcctga tggcgacaca ggcatggctg gaagcgcgcc tgaaaggcga agaactgcca 1020
aaaccagaag gcctgctgac acaaaacgaa gaactgtggg aaccgctcta cgcttgctac 1080
cagtcacttc aggcgtgtgg catgggtatt atcgccaacg gcgatctgct cgacaccctg 1140
cgccgcgtga aatgtttcgg cgtaccgctg gtccgtattg atatccgtca ggagagcacg 1200
cgtcataccg aagcgctggg cgagctgacc cgctacctcg gtatcggcga ctacgaaagc 1260
tggtcagagg ccgacaaaca ggcgttcctg atccgcgaac tgaactccaa acgtccgctt 1320
ctgccgcgca actggcaacc aagcgccgaa acgcgcgaag tgctcgatac ctgccaggtg 1380
attgccgaag caccgcaagg ctccattgcc gcctacgtga tctcgatggc gaaaacgccg 1440
tccgacgtac tggctgtcca cctgctgctg aaagaagcgg gtatcgggtt tgcgatgccg 1500
gttgctccgc tgtttgaaac cctcgatgat ctgaacaacg ccaacgatgt catgacccag 1560
ctgctcaata ttgactggta tcgtggcctg attcagggca aacagatggt gatgattggc 1620
tattccgact cagcaaaaga tgcgggagtg atggcagctt cctgggcgca atatcaggca 1680
caggatgcat taatcaaaac ctgcgaaaaa gcgggtattg agctgacgtt gttccacggt 1740
cgcggcggtt ccattggtcg cggcggcgca cctgctcatg cggcgctgct gtcacaaccg 1800
ccaggaagcc tgaaaggcgg cctgcgcgta accgaacagg gcgagatgat ccgctttaaa 1860
tatggtctgc cagaaatcac cgtcagcagc ctgtcgcttt ataccggggc gattctggaa 1920
gccaacctgc tgccaccgcc ggagccgaaa gagagctggc gtcgcattat ggatgaactg 1980
tcagtcatct cctgcgatgt ctaccgcggc tacgtacgtg aaaacaaaga ttttgtgcct 2040
tacttccgct ccgctacgcc ggaacaagaa ctgggcaaac tgccgttggg ttcacgtccg 2100
gcgaaacgtc gcccaaccgg cggcgtcgag tcactacgcg ccattccgtg gatcttcgcc 2160
tggacgcaaa accgtctgat gctccccgcc tggctgggtg caggtacggc gctgcaaaaa 2220
gtggtcgaag acggcaaaca gagcgagctg gaggctatgt gccgcgattg gccattcttc 2280
tcgacgcgtc tcggcatgct ggagatggtc ttcgccaaag cagacctgtg gctggcggaa 2340
tactatgacc aacgcctggt agacaaagca ctgtggccgt taggtaaaga gttacgcaac 2400
ctgcaagaag aagacatcaa agtggtgctg gcgattgcca acgattccca tctgatggcc 2460
gatctgccgt ggattgcaga gtctattcag ctacggaata tttacaccga cccgctgaac 2520
gtattgcagg ccgagttgct gcaccgctcc cgccaggcag aaaaagaagg ccaggaaccg 2580
gatcctcgcg tcgaacaagc gttaatggtc actattgccg ggattgcggc aggtatgcgt 2640
aataccggct aa 2652
<210> 28
<211> 1191
<212> DNA
<213> 大肠杆菌
<220>
<221> 基因
<222> (1)..(1191)
<223> 天冬氨酸氨基转移酶
<400> 28
atgtttgaga acattaccgc cgctcctgcc gacccgattc tgggcctggc cgatctgttt 60
cgtgccgatg aacgtcccgg caaaattaac ctcgggattg gtgtctataa agatgagacg 120
ggcaaaaccc cggtactgac cagcgtgaaa aaggctgaac agtatctgct cgaaaatgaa 180
accaccaaaa attacctcgg cattgacggc atccctgaat ttggtcgctg cactcaggaa 240
ctgctgtttg gtaaaggtag cgccctgatc aatgacaaac gtgctcgcac ggcacagact 300
ccggggggca ctggcgcact acgcgtggct gccgatttcc tggcaaaaaa taccagcgtt 360
aagcgtgtgt gggtgagcaa cccaagctgg ccgaaccata agagcgtctt taactctgca 420
ggtctggaag ttcgtgaata cgcttattat gatgcggaaa atcacactct tgacttcgat 480
gcactgatta acagcctgaa tgaagctcag gctggcgacg tagtgctgtt ccatggctgc 540
tgccataacc caaccggtat cgaccctacg ctggaacaat ggcaaacact ggcacaactc 600
tccgttgaga aaggctggtt accgctgttt gacttcgctt accagggttt tgcccgtggt 660
ctggaagaag atgctgaagg actgcgcgct ttcgcggcta tgcataaaga gctgattgtt 720
gccagttcct actctaaaaa ctttggcctg tacaacgagc gtgttggcgc ttgtactctg 780
gttgctgccg acagtgaaac cgttgatcgc gcattcagcc aaatgaaagc ggcgattcgc 840
gctaactact ctaacccacc agcacacggc gcttctgttg ttgccaccat cctgagcaac 900
gatgcgttac gtgcgatttg ggaacaagag ctgactgata tgcgccagcg tattcagcgt 960
atgcgtcagt tgttcgtcaa tacgctgcag gaaaaaggcg caaaccgcga cttcagcttt 1020
atcatcaaac agaacggcat gttctccttc agtggcctga caaaagaaca agtgctgcgt 1080
ctgcgcgaag agtttggcgt atatgcggtt gcttctggtc gcgtaaatgt ggccgggatg 1140
acaccagata acatggctcc gctgtgcgaa gcgattgtgg cagtgctgta a 1191
<210> 29
<211> 1104
<212> DNA
<213> 大肠杆菌
<220>
<221> 基因
<222> (1)..(1104)
<223> 天冬氨酸半醛脱氢酶
<400> 29
atgaaaaatg ttggttttat cggctggcgc ggtatggtcg gctccgttct catgcaacgc 60
atggttgaag agcgcgactt cgacgccatt cgccctgtct tcttttctac ttctcagctt 120
ggccaggctg cgccgtcttt tggcggaacc actggcacac ttcaggatgc ctttgatctg 180
gaggcgctaa aggccctcga tatcattgtg acctgtcagg gcggcgatta taccaacgaa 240
atctatccaa agcttcgtga aagcggatgg caaggttact ggattgacgc agcatcgtct 300
ctgcgcatga aagatgacgc catcatcatt cttgaccccg tcaatcagga cgtcattacc 360
gacggattaa ataatggcat caggactttt gttggcggta actgtaccgt aagcctgatg 420
ttgatgtcgt tgggtggttt attcgccaat gatcttgttg attgggtgtc cgttgcaacc 480
taccaggccg cttccggcgg tggtgcgcga catatgcgtg agttattaac ccagatgggc 540
catctgtatg gccatgtggc agatgaactc gcgaccccgt cctctgctat tctcgatatc 600
gaacgcaaag tcacaacctt aacccgtagc ggtgagctgc cggtggataa ctttggcgtg 660
ccgctggcgg gtagcctgat tccgtggatc gacaaacagc tcgataacgg tcagagccgc 720
gaagagtgga aagggcaggc ggaaaccaac aagatcctca acacatcttc cgtaattccg 780
gtagatggtt tatgtgtgcg tgtcggggca ttgcgctgcc acagccaggc attcactatt 840
aaattgaaaa aagatgtgtc tattccgacc gtggaagaac tgctggctgc gcacaatccg 900
tgggcgaaag tcgttccgaa cgatcgggaa atcactatgc gtgagctaac cccagctgcc 960
gttaccggca cgctgaccac gccggtaggc cgcctgcgta agctgaatat gggaccagag 1020
ttcctgtcag cctttaccgt gggcgaccag ctgctgtggg gggccgcgga gccgctgcgt 1080
cggatgcttc gtcaactggc gtaa 1104
<210> 30
<211> 70
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 30
ttactctggt gcctgacatt tcaccgacaa agcccaggga acttcatcac gtgtaggctg 60
gagctgcttc 70
<210> 31
<211> 70
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 31
ttaccccttg tttgcagccc ggaagccatt ttccaggtcg gcaattaaat catatgaata 60
tcctccttag 70
<210> 32
<211> 70
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 32
aaagaatatg ccgatcggtt cgggcttagg ctccagtgcc tgttcggtgg gtgtaggctg 60
gagctgcttc 70
<210> 33
<211> 70
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 33
agacaaccga catcgctttc aacattggcg accggagccg ggaaggcaaa catatgaata 60
tcctccttag 70
<210> 34
<211> 29
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 34
aattgatatc atgccgattc gtgtgccgg 29
<210> 35
<211> 34
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 35
aattaagctt ttaatccagc gttggattca tgtg 34
<210> 36
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 36
ttgtaactgg tgcgccgctg gaactggtgg ggtttaatga tgtc 44
<210> 37
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 37
gacatcatta aaccccacca gttccagcgg cgcaccagtt acaa 44
<210> 38
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 38
tgtaactggt gcgccgctgg aacatgtggg gtttaatgat gtcg 44
<210> 39
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 39
cgacatcatt aaaccccaca tgttccagcg gcgcaccagt taca 44
<210> 40
<211> 42
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 40
tttccgaaac gtacctcagc aggtgtaggc tggagctgct tc 42
<210> 41
<211> 43
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 41
gaataaaatt tattcacctg ctgcatatga atatcctcct tag 43
<210> 42
<211> 29
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 42
aattgatatc atgccgattc gtgtgccgg 29
<210> 43
<211> 27
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 43
aattaagcct gctgaggtac gtttcgg 27
<210> 44
<211> 23
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 44
cagcaggtga ataaatttta ttc 23
<210> 45
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 45
cgcgaatgga agctgtttcc 20
<210> 46
<211> 31
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 46
gccggaattc tgtcggatgc gatacttgcg c 31
<210> 47
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 47
gaaggagctc agaaaaccct cgcgcaaaag 30
<210> 48
<211> 31
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 48
gccggagctc tgtcggatgc gatacttgcg c 31
<210> 49
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 49
gaagggtacc agaaaaccct cgcgcaaaag 30
<210> 50
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 50
tccgagctca taagcgtagc gcatcaggca 30
<210> 51
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 51
tccgagctcg tccacctatg ttgactacat 30
<210> 52
<211> 27
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 52
ccggaattcc caggagagca ataagca 27
<210> 53
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 53
ctagtctaga tgctctattt aactcccg 28
<210> 54
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 54
ctagtctaga ccaggagagc aataagca 28
<210> 55
<211> 27
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 55
ccggaattct gctctattta actcccg 27
<210> 56
<211> 70
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 56
aggcgctaag gagaccttaa atggctgata caaaagcaaa actcaccctc tgtaggctgg 60
agctgcttcg 70
<210> 57
<211> 70
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 57
gggttaaaat atttacaact tagcaatcaa ccattaacgc ttgatatcgc atgggaatta 60
gccatggtcc 70
<210> 58
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 58
cctttctata actgcgcgtc at 22
<210> 59
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 59
aggggtatag ctacgccaga a 21
<210> 60
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 60
gaaggcaaat ttaagttccg 20
<210> 61
<211> 40
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 61
cgaagcagct ccagcctaca ggtatagata gacgtcattt 40
<210> 62
<211> 42
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 62
cggcgcgctg gcggcgttcg cgcacgactc gctggatgtt aa 42
<210> 63
<211> 42
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 63
ttaacatcca gcgagtcgtg cgcgaacgcc gccagcgcgc cg 42
<210> 64
<211> 42
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 64
cgcgttccgc ctgctgtcgg cgatgccgac catggccgcg at 42
<210> 65
<211> 42
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 65
atcgcggcca tggtcggcat cgccgacagc aggcggaacg cg 42
<210> 66
<211> 40
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 66
tcctgaatat gatgttctcc gcgccgtgcg aaccgtatga 40
<210> 67
<211> 40
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 67
tcatacggtt cgcacggcgc ggagaacatc atattcagga 40
<210> 68
<211> 40
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 68
aaatgacgtc tatctatacc tgtaggctgg agctgcttcg 40
<210> 69
<211> 76
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 69
gcaaaactaa attattggta tcatgaattt gttgtatgat gaataaaata taggggatgg 60
gaattagcca tggtcc 76
<210> 70
<211> 67
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 70
ccatggagga ggaattaacc atgcagtggt ggtggtggtg gtgcgataac agctgtatcc 60
ccgttga 67
<210> 71
<211> 67
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 71
aagcttagga ggaattaacc atgcagtggt ggtggtggtg gtgcgatgca aatttaagtt 60
ccggcag 67
Claims (10)
1.生产L-甲硫氨酸方法,包括:
(a)培养埃希氏菌属(Escherichia sp.)的微生物,生成O-乙酰基高丝氨酸,其中所述微生物的柠檬酸合酶的内源活性减弱或失活;和
(b)使步骤(a)中生成的O-乙酰基高丝氨酸与O-乙酰基高丝氨酸硫化氢解酶或具有O-乙酰基高丝氨酸硫化氢解酶的微生物接触。
2.根据权利要求1所述的方法,其中柠檬酸合酶内源活性减弱的所述微生物具有由SEQID NO:1或SEQ ID NO:2表示的氨基酸序列。
3.根据权利要求1所述的方法,其中(a)的微生物进一步具有与其内源活性相比减弱或失活的胱硫醚γ合酶、高丝氨酸激酶或两者的活性。
4.根据权利要求1所述的方法,其中(a)的微生物进一步具有引入或增强的高丝氨酸O-乙酰基转移酶活性,或具有内源高丝氨酸O-琥珀酰基转移酶,所述内源高丝氨酸O-琥珀酰基转移酶被进一步修饰成与SEQ ID NO:16具有95%或以上同一性的多肽并且进一步具有在G111E位和L112T或L112H位的突变以具有高丝氨酸O-乙酰基转移酶活性。
5.根据权利要求1所述的方法,其中(a)的微生物进一步具有引入或增强的至少一种蛋白质的活性,所述至少一种蛋白质选自磷酸烯醇丙酮酸羧化酶、天冬氨酸氨基转移酶和天冬氨酸半醛脱氢酶。
6.根据权利要求1所述的方法,其中(a)的微生物是大肠杆菌(Escherichia coli)。
7.根据权利要求1所述的方法,其中O-乙酰基高丝氨酸硫化氢解酶衍生自钩端螺旋体属(Leptospira sp.)、色杆菌属(Chromobacterium sp.)、或生丝单胞菌属(Hyphomonassp.)。
8.根据权利要求1所述的方法,进一步包括在步骤(b)中加入甲基硫醇作为底物。
9.根据权利要求1所述的方法,进一步包括回收步骤(a)中生成的O-乙酰基高丝氨酸。
10.根据权利要求1所述的方法,进一步包括回收步骤(b)中生成的L-甲硫氨酸。
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KR1020140076779A KR101641770B1 (ko) | 2014-06-23 | 2014-06-23 | O-아세틸 호모세린을 생산하는 미생물 및 상기 미생물을 이용하여 o-아세틸 호모세린을 생산하는 방법 |
KR10-2014-0076779 | 2014-06-23 | ||
CN201580001286.8A CN105392880B (zh) | 2014-06-23 | 2015-06-22 | 生产o-乙酰基高丝氨酸的微生物和用其生产o-乙酰基高丝氨酸的方法 |
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CN112501144A (zh) * | 2021-02-02 | 2021-03-16 | 中国科学院天津工业生物技术研究所 | 高丝氨酸乙酰转移酶突变体及其在生产o-乙酰高丝氨酸中的应用 |
CN113122486A (zh) * | 2019-12-31 | 2021-07-16 | 江南大学 | 一种丙二酸的全生物合成的方法 |
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WO2019004779A2 (ko) * | 2017-06-30 | 2019-01-03 | 씨제이제일제당 (주) | 신규한 o-숙시닐 호모세린 트랜스퍼라제 변이체 및 이를 이용한 o-숙시닐 호모세린의 제조방법 |
CN110129294B (zh) * | 2018-02-02 | 2021-12-24 | 中国科学院天津工业生物技术研究所 | 高丝氨酸乙酰基转移酶突变体及其宿主细胞和应用 |
KR101915433B1 (ko) | 2018-02-13 | 2018-11-05 | 씨제이제일제당 (주) | 시트레이트 신타아제 (Citrate synthase)의 활성이 약화된 변이형 폴리펩타이드 및 이를 이용한 L-아미노산 생산방법 |
KR102525073B1 (ko) * | 2021-03-10 | 2023-04-24 | 씨제이제일제당 주식회사 | 신규한 시트레이트 신타아제 변이체 및 이를 이용한 l-발린 생산 방법 |
KR102525074B1 (ko) * | 2021-03-10 | 2023-04-24 | 씨제이제일제당 주식회사 | 신규한 시트레이트 신타아제 변이체 및 이를 이용한 o-아세틸-l-호모세린 또는 l-메티오닌 생산 방법 |
KR102525072B1 (ko) * | 2021-03-10 | 2023-04-24 | 씨제이제일제당 주식회사 | 신규한 시트레이트 신타아제 변이체 및 이를 이용한 l-아미노산 생산 방법 |
CN113388564B (zh) * | 2021-06-04 | 2022-03-01 | 浙江工业大学 | 一种o-乙酰-l-高丝氨酸生产菌、构建方法及应用 |
KR20230139597A (ko) * | 2022-03-28 | 2023-10-05 | 씨제이제일제당 (주) | O-아세틸 호모세린 생산 미생물 및 이를 이용한 o-아세틸 호모세린 또는 l-메치오닌 생산 방법 |
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EP2998388A4 (en) | 2017-03-22 |
US10465217B2 (en) | 2019-11-05 |
US20170101655A1 (en) | 2017-04-13 |
EP2998388A1 (en) | 2016-03-23 |
AU2015264911B2 (en) | 2017-11-23 |
AU2015264911A1 (en) | 2016-01-21 |
PL2998388T3 (pl) | 2021-05-17 |
WO2015199396A1 (ko) | 2015-12-30 |
CN105392880A (zh) | 2016-03-09 |
CN105392880B (zh) | 2019-11-08 |
US20200024626A1 (en) | 2020-01-23 |
KR101641770B1 (ko) | 2016-07-22 |
BR112015029716A2 (pt) | 2017-09-26 |
JP2016526927A (ja) | 2016-09-08 |
JP2018046875A (ja) | 2018-03-29 |
JP6607974B2 (ja) | 2019-11-20 |
MY185764A (en) | 2021-06-04 |
EP2998388B1 (en) | 2020-10-07 |
US10815509B2 (en) | 2020-10-27 |
EP3783096A1 (en) | 2021-02-24 |
ES2833432T3 (es) | 2021-06-15 |
BR112015029716B1 (pt) | 2022-07-19 |
RU2614253C1 (ru) | 2017-03-24 |
EP3783096B1 (en) | 2024-03-13 |
KR20160000098A (ko) | 2016-01-04 |
DK2998388T3 (da) | 2020-11-23 |
SG11201509451WA (en) | 2016-01-28 |
BR122020005905B1 (pt) | 2022-07-26 |
JP6336074B2 (ja) | 2018-06-06 |
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