CN107109357A - 能生产o‑琥珀酰高丝氨酸或琥珀酸的微生物和用其生产琥珀酸或o‑琥珀酰高丝氨酸方法 - Google Patents
能生产o‑琥珀酰高丝氨酸或琥珀酸的微生物和用其生产琥珀酸或o‑琥珀酰高丝氨酸方法 Download PDFInfo
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Abstract
本发明提供具有生产O‑琥珀酰高丝氨酸或琥珀酸的能力的微生物,以及通过使用所述微生物生产O‑琥珀酰高丝氨酸或琥珀酸的方法。
Description
技术领域
本发明涉及具有生产O-琥珀酰高丝氨酸或琥珀酸的能力的微生物,以及通过使用所述微生物生产O-琥珀酰高丝氨酸或琥珀酸的方法。
背景技术
为了提升在需氧条件下生产O-琥珀酰高丝氨酸或琥珀酸,需要琥珀酰-CoA作为前体。α-酮戊二酸脱氢酶在三羧酸(TCA)循环中起到用于α-酮戊二酸(盐/酯)脱氢成琥珀酸的催化剂的作用。编码这种酶的基因sucAB在染色体上与编码琥珀酸脱氢酶的sdhCDAB和编码琥珀酰-CoA合成酶的sucCD一起成簇表达,并且已知这种表达是由包含在簇中的两个启动子诱导的。簇中的基因表达主要是由sdhC启动子诱导的。此时,以sdhCDAB、sucAB和sucCD的顺序表达。由于TCA循环具有有利于sdhCDAB(编码琥珀酸脱氢酶的基因)的表达的结构,因此琥珀酸脱氢酶的表达较强,所以即使通过sucAB的表达而产生琥珀酰-CoA,琥珀酰-CoA也会易于被快速分解。sucA启动子位于簇的中心,即位于sdhCDAB和sucAB之间。sucA启动子引起sucAB的表达,但在该簇中更重要的启动子是sdhC,其诱导sdhCDAB的表达。sucA启动子比sdhC启动子弱(S J Park,G Chao以及R P Gunsalus,J.Bacteriol.(微生物学杂志)1997;Cunningham,L.&Guest,J.R.(1998)Microbiology(微生物学)144,2113-2123)。因此,有报告显示,通过在需氧条件下在生产琥珀酸的菌株中缺失sdhAB来提高琥珀酸的生产(A.Yu.Skorofkhodova等,Applied Biochemistry and Microbiology(应用生物化学与微生物学),2013年12月,第49卷,第7期,第629-637页)。
在对提升琥珀酸或O-琥珀酰高丝氨酸生产的研究过程中,本发明人开发了能够通过增强α-酮戊二酸脱氢酶的活性来提升O-琥珀酰高丝氨酸或琥珀酸生产的微生物,从而完成本发明。
发明内容
技术问题
本发明的一个目的是提供用于生产O-琥珀酰高丝氨酸或琥珀酸的微生物。
本发明的另一个目的是提供生产O-琥珀酰高丝氨酸或琥珀酸的方法,其包括培养用于生产O-琥珀酰高丝氨酸或琥珀酸的微生物的步骤。
技术方案
根据本发明的一个方面,提供了用于生产O-琥珀酰高丝氨酸或琥珀酸的微生物。
在本发明的一种实施方式中,具有用于生产O-琥珀酰高丝氨酸或琥珀酸的微生物可以是具有如下特征的具有生产O-琥珀酰高丝氨酸或琥珀酸的能力的微生物:α-酮戊二酸脱氢酶的活性与其内源活性水平相比得到增强。
本文中使用时,术语“用于生产O-琥珀酰高丝氨酸或琥珀酸的微生物”表示能够在活体中生产O-琥珀酰高丝氨酸或琥珀酸并且在其中积累或分泌O-琥珀酰高丝氨酸或琥珀酸的原核或真核微生物。例如,用于生产O-琥珀酰高丝氨酸或琥珀酸的微生物可以是属于如下的微生物:埃希氏菌(Escherichia)属、欧文氏菌(Erwinia)属、沙雷氏菌(Serratia)属、普罗威登斯菌(Providencia)属、棒状杆菌(Corynebacteria)属、假单胞菌(Pseudomonas)属、钩端螺旋体(Leptospira)属、沙门氏菌(Salmonella)属、短杆菌(Brevibacterium)属、生丝单胞菌(Hypomononas)属、色杆菌(Chromobacterium)属、诺卡氏菌(Nocardia)属、真菌或酵母。在一种实施方式中,根据本发明的微生物可以是属于埃希氏菌属的微生物。在一种实施方式中,根据本发明的微生物可以是大肠杆菌(E.coli)。
为了提升琥珀酸和O-琥珀酰高丝氨酸的生产,需要琥珀酰-CoA。琥珀酰-CoA是通过在TCA循环中α-酮戊二酸(盐/酯)脱氢而获得的。
α-酮戊二酸脱氢酶是一种催化酶,其催化通过在TCA循环中α-酮戊二酸(盐/酯)脱氢而生产琥珀酸的过程。α-酮戊二酸脱氢酶以包含α-酮戊二酸脱氢酶和二氢硫辛酸-琥珀酰基转移酶的复合体形式存在,并且由sucAB(sucA GenBank Accession No.(基因银行编号)BAA35392.1,sucB GenBank Accession No.BAA35393.1)基因编码。α-酮戊二酸脱氢酶和二氢硫辛酸-琥珀酰基转移酶可以分别具有序列号22和序列号24中分别列出的氨基酸序列,或者具有与上述序列具有至少70%、80%、90%或95%同源性的序列,其由sucA和sucB基因编码的。sucA和sucB可以分别具有序列号23和序列号25中分别列出的序列,或者具有与上述序列具有至少70%、80%、90%或95%同源性的序列。sucAB与sdhCDAB和sucCD一起成簇表达。由于sdhCDAB的强启动子,编码琥珀酸脱氢酶的sdhCDAB高度表达,并从而快速分解琥珀酸和琥珀酰-CoA(Louise Cunningham等,Microbiology(微生物学)(1998),144,第211-2123页)。因此,通过增强酶的活性提高琥珀酸的生产,可以提升琥珀酸和琥珀酰-CoA的生产。
本文中使用时,术语“内源”活性表示在自然条件下或者在突变之前在原始微生物或细胞中的活性。
在本发明的一种实施方式中,上文中的微生物是属于埃希氏菌属的微生物,其中高丝氨酸O-琥珀酰基转移酶的活性进一步与其内源活性水平相比得到增强。
高丝氨酸O-琥珀酰基转移酶是一种起到用于由琥珀酰-CoA和高丝氨酸生产O-琥珀酰高丝氨酸的催化剂作用的酶,其参与甲硫氨酸生物合成途径的第一步。该酶可以具有序列号26中所列的氨基酸序列,或者具有与该序列具有至少70%、80%、90%或95%同源性的氨基酸序列,并且可以由metA基因(GenBank Accession No.(基因银行编号)BAE78015.1)编码。metA基因可以具有序列号27中所列的序列,或具有与该序列具有至少70%、80%、90%或95%同源性的序列。该基因的表达受到甲硫氨酸的反馈控制的抑制。因此,通过消除甲硫氨酸的反馈控制来使metA高度表达的突变体已被用于提升甲硫氨酸生产能力。已解除甲硫氨酸的反馈控制的高丝氨酸O-琥珀酰基转移酶可以具有序列号32所列的氨基酸序列或者具有与上述序列具有至少70%、80%、90%或95%同源性的序列,并且其基因可以由序列号33(metA11:韩国专利公开号2009-0106365)中所列的序列编码或与上述序列具有至少80%、90%或95%同源性的序列编码。
当与本文中所用的序列相关地使用时,术语“同源性”指的是与给定的氨基酸序列或核苷酸序列匹配的程度,并且其可以表示为百分数(%)。在本文中,具有与给定氨基酸序列或核苷酸序列相同或相似的活性的同源性序列用“%同源性”来表示,其可以通过能够使用BLAST 2.0计算机程序的技术人员公知的方法来确定,BLAST 2.0计算机程序计算参数,例如得分、一致性和相似性。
本文中使用时,用于酶活性的术语“增强”指的是酶活性通过编码该酶的基因的过表达而被提高到比内源活性的水平更高的水平,或者通过该基因中的突变而使所编码的酶的活性提高到比内源活性水平更高的水平。例如,酶活性的增强可通过以下方式实现:提高编码该酶的基因的复制次数;或者用较强的启动子替换该基因的内源启动子;或者通过缺失、取代或插入来使染色体上的该基因的部分核苷酸序列或整个核苷酸序列突变,或使整个表达调控序列或一部分表达调控序列突变,或者它们的组合。
本文中使用时,术语“表达调控序列”表示调控基因表达的核苷酸序列,其是能够增加或减少在生物体中特定基因的表达的片段,并且包括启动子、转录因子结合位点等,但不限于此。
在本发明的一种实施方式中,为了提高相应的酶活性,可以用比内源启动子强的启动子替换编码该酶的基因的启动子,或者可以提高基因的复制次数。
在本发明的一种实施方式中,启动子可以是均具有强活性的Ptac、Ptrc、Ppro、PR、PL、Prmf或PcysK,但不限于此。
在本发明的一种实施方式中,微生物可以是属于埃希氏菌属的微生物,其中胱硫醚γ-合酶和高丝氨酸激酶中至少一种酶的活性进一步与相应的内源活性水平相比被减弱或者消除。
胱硫醚γ-合酶具有催化O-琥珀酰高丝氨酸转化成胱硫醚的活性。胱硫醚γ-合酶可以具有序列号28中所列的氨基酸序列或具有与该序列具有至少70%、80%、90%或95%同源性的序列,并且可以由metB基因(GenBank Accession No.BAE77371.1)编码。metB基因可以具有序列号29中所列的序列或具有与该序列具有至少70%、80%、90%或95%同源性的序列。一旦该酶的活性被减弱或消除,则O-琥珀酰高丝氨酸不被转化成胱硫醚,或者其转化率非常低。
高丝氨酸激酶起到用于由高丝氨酸合成O-磷酸高丝氨酸的催化剂的作用,并且可以具有序列号30中所列的氨基酸序列或具有与该序列具有至少70%、80%、90%或95%同源性的序列,并且可以由thrB基因(GenBank Accession No.BAB96580.1)编码。thrB基因可以具有序列号31中所列的序列或具有与该序列至少70%、80%、90%或95%同源性的序列。一旦该酶的活性被减弱或消除,则高丝氨酸不被转化成O-磷酸高丝氨酸,或者其转化率非常低。
本文中使用时,酶活性的“减弱”或“消除”分别表示编码相应酶的基因的表达降低至比内源水平低的水平或者酶活性完全去除。这可以通过如下来实现:通过缺失、替换、插入或者使整个核苷酸序列或一部分核苷酸序列突变、或者使整个或部分表达调控序列突变,或者它们的组合。
在本发明的一种实施方式中,微生物可以是大肠杆菌。该微生物可以是通过如下方式而获得的大肠杆菌:增强α-酮戊二酸脱氢酶的活性至比内源水平高的水平,并且额外地增强高丝氨酸O-琥珀酰基转移酶的活性至比内源水平更高的水平,或者额外地使胱硫醚γ-合酶和高丝氨酸激酶中的至少一种酶的活性减弱到比内源水平更低的水平或者消除。
在本发明的一种实施方式中,在本发明的微生物可以是通过如下方式获得的大肠杆菌:用突变体metA11(序列号33,WO2008-127240 A1)替换大肠杆菌的染色体上的metA基因,其中,metA11是通过消除甲硫氨酸介导的反馈调控而获得的metA基因的变体,并且被含处于比其原始启动子更强的启动子的控制下的sucAB的载体转化,并且使染色体缺失thrB和metB基因缺失。
本发明的一个方面提供生产O-琥珀酰高丝氨酸或琥珀酸的方法,所述方法包括培养埃希氏菌属微生物,其中α-酮戊二酸脱氢酶的活性与其内源活性水平相比得到增强;并且从微生物的培养液回收O-琥珀酰高丝氨酸或琥珀酸。
在本发明的一种实施方式中,在生产O-琥珀酰高丝氨酸或琥珀酸的方法中,可以通过使用本领域技术人员公知的培养条件和合适的培养基来实现培养用于生产O-琥珀酰高丝氨酸或琥珀酸的微生物。该培养过程可以根据本领域技术人员所选择的菌株来容易地调整。培养方法的实例包括间歇式培养、连续式培养和补料分批式培养,但不限于此。包括上述方法的各种培养方法在文献("Biochemical Engineering(生物化学工程)",由JamesM.Lee所著,Prentice-Hall International Editions,第138-176页)中详细说明。
本文中的培养基必须满足培养本文中特定菌株所要求的培养条件。在文献(美国细菌学会(American Society for Bacteriology)的“Manual of Methods for GeneralBacteriology(普通细菌学方法手册)”,Washington D.C.,USA,1981)中描述了各种用于微生物的培养基。这些培养基包含各种碳源、氮源和微量元素。碳源包括碳水化合物,例如葡萄糖、乳糖、蔗糖、果糖、麦芽糖、淀粉和纤维素;脂肪如大豆油、向日葵油、蓖麻油和椰子油;脂肪酸如棕榈酸、硬脂酸和亚油酸;醇如甘油和乙醇;和有机酸如乙酸。碳源可以单独使用或作为至少两种碳源的组合来使用。氮源包括有机氮源,例如蛋白胨、酵母提取物、肉汁、麦芽提取物、玉米浆(CSL)和豆粉;以及无机氮源,例如尿素、硫酸铵、氯化铵、磷酸铵、碳酸铵和硝酸铵。这些氮源可以单独使用或组合使用。此外,在上文的培养基中可以额外包含磷源,并且磷源的实例包括磷酸二氢钾、磷酸氢二钾和其相应的含钠盐。本文中的培养基还可以包含金属如硫酸镁或硫酸铁。此外,也可以添加氨基酸、维生素和适当的前体。
为了在培养液中维持需氧条件,可以将氧气或含氧气体(例如空气)注入培养在中。本文中培养物的温度通常可以为20-45℃,优选地为25-40℃。可以继续培养,直至O-琥珀酰高丝氨酸或琥珀酸的生产达到所期望水平,并且优选的培养时间是10小时-160小时。
有益效果
根据本发明的生产O-琥珀酰高丝氨酸-或琥珀酸的菌株能够有效地生产O-琥珀酰高丝氨酸或琥珀酸,并且由于酶促或化学促转化的可能性,其可以被应用于各种领域中。
附图说明
图1是重组载体pCL_PsucA-sucAB的示意图。
图2是重组载体pCL_Prmf-sucAB的示意图。
图3是重组载体pCL_Ptrc-sucAB的示意图。
图4是重组载体pCL_Pcysk-metA11_PsucA-sucAB的示意图。
具体实施方式
下文中,将参考一些实例更详细地描述本发明。然而,这些实例仅仅用于说明性目的,并且本申请的范围不意图受这些实例限制。
参考实例:生产O-琥珀酰高丝氨酸-或琥珀酸的菌株的构建
1)metB基因缺失
为了提高O-琥珀酰高丝氨酸或琥珀酸的积累,通过使编码L-甲硫氨酸前体降解中所涉及的胱硫醚γ-合酶的metB基因缺失来构建菌株。
使野生型大肠杆菌(K12)W3110菌株中编码胱硫醚γ-合酶的metB基因缺失。已知,胱硫醚γ-合酶与各种甲硫氨酸前体结合,并因此能够产生各种副产物。因此,胱硫醚合酶的过表达可以提高副反应,以降低细胞内反应的效率。为了缺失metB基因,进行FRT-一步-PCR(FRT-one-step-PCR)缺失方法(PNAS(2000),Vol.197,p6640-6645)。为了缺失metB基因,使用pKD3载体(PNAS(2000)Vol.97,p6640-6645)作为模板用由序列号12和序列号13表示的引物进行PCR,得到缺失盒(cassette)的构建。
序列号12:
5'-TTACTCTGGTGCCTGACATTTCACCGACAAAGCCCAGGGAACTTCATCACGTGTAGGCTGGAGCTGCTTC-3'
序列号13:
5'-CGCTGCGCCAGCTCCATACGCGGCACCAGCGTTCGCAACCCACGTAGCAGCATATGAATATCCTCCTTAG-3'
按如下进行PCR:在95℃下变性30秒,在55℃下退火30秒,并在72℃下延伸1分钟,进行30个从变性到延伸的循环。将PCR产物在1.0%琼脂糖凝胶上进行电泳。洗脱并纯化1.1kb带。将回收的DNA片段电穿孔到之前用pKD46载体(PNAS(2000)Vol.97,p6640-6645)转化过的大肠杆菌(K12)W3110菌株中。为了电穿孔,在30℃下在补充有200μg/L氨苄西林和5mM L-阿拉伯糖的LB培养基中培养用pKD46转化过的W3110菌株,直到OD600达到0.5。将菌株用10%甘油洗涤三次。在2500V下进行电穿孔。将回收的菌株涂在含30μg/L氯霉素的LB平板培养基上,然后在37℃下培养1至2天。然后选择抗性菌株。
然后,根据上文所提到的条件,使用由序列号14和序列号15表示的引物,将所选择的菌株作为模板进行PCR。在1.0%琼脂糖凝胶上观察到1.5kb大小的带,表明metB基因缺失。
序列号14:5'-TATTCGCCGCTCCATTCAGC-3'
序列号15:5'-TACCCCTTGTTTGCAGCCCG-3'
使用pCP20载体(PNAS(2000)vol.97,p6640-6645)转化metB基因缺失的菌株,然后将其在补充有100μg/L氨苄西林的LB培养基中培养。通过上文所描述的方式相同的方式进行PCR,并且选择最终的metB基因缺失的菌株,其在1.0%琼脂糖凝胶上表现了收缩的带。证实氯霉素标记的缺失。获得的甲硫氨酸营养缺陷型菌株被命名为CC03-0132。
2)thrB基因缺失
为了提高由高丝氨酸生产的O-琥珀酰高丝氨酸,使编码高丝氨酸激酶的基因thrB缺失。具体地,当使用生产苏氨酸的菌株时,该基因的缺失是必要的,因为高丝氨酸利用活性非常强。为了从上文1)中所制备的CC03-0132菌株缺失thrB基因,进行FRT-一步-PCR缺失方法(PNAS(2000),Vol.197,p6640-6645)。为了缺失thrB基因,使用pKD3载体(PNAS(2000)Vol.97,p6640-6645)作为模板用由序列号16和序列号17表示的引物进行PCR,得到缺失盒(cassette)的构建。
序列号16:
5'-CATGGTTAAAGTTTATGCCCCGGCTTCCAGTGCCAATATGAGCGTCGGGTGTGTAGGCTGGAGCTGCTTC-3'
序列号17:
5'-GGAGATACCGCTCGCTACCGCGCCGATTTCCGCGACCGCCTGCCGCGCCTCATATGAATATCCTCCTTAG-3'
按如下进行PCR:在95℃下变性30秒,在55℃下退火30秒,并在72℃下延伸1分钟,进行30个从变性到延伸的循环。将PCR产物在1.0%琼脂糖凝胶上进行电泳。洗脱并纯化1.1kb带。将回收的DNA片段电穿孔到之前用pKD46载体(PNAS(2000)Vol.97,p6640-6645)转化过的CC03-0132菌株中。为了电穿孔,在30℃下在补充有200μg/L氨苄西林和5mM L-阿拉伯糖的LB培养基中培养用pKD46转化过的CC03-0132菌株,直到OD600达到0.5。将菌株用10%甘油洗涤三次。在2500V下进行电穿孔。将回收的菌株涂在含30μg/L氯霉素的LB平板培养基上,然后在37℃下培养1至2天。然后选择抗性菌株。
然后,根据上文所提到的条件,使用由序列号18和序列号19表示的引物,将所选择的菌株作为模板进行PCR。在1.0%琼脂糖凝胶上观察到1.5kb大小的带,表明thrB基因缺失。
序列号18:5'-ACTCGACGATCTCTTTGCC-3'
序列号19:5'-ACGCCGAGAGGATCTTCGCAG-3'
使用pCP20载体(PNAS(2000)vol.97,p6640-6645)转化所证实的的菌株,然后将其在补充有100μg/L氨苄西林的LB培养基中培养。通过上文所描述的方式相同的方式进行PCR,并且选择最终的thrB基因缺失的菌株,其在1.0%琼脂糖凝胶电泳上表现了较小的带。证实氯霉素标记的缺失。获得的菌株被命名为CC03-0133。
3)用于插入metA11的pSG载体的构建
高丝氨酸琥珀酰转移酶在添加到培养基的痕量甲硫氨酸的反馈控制下,其大部分活性被抑制。为了提高作为L-甲硫氨酸前体的O-琥珀酰高丝氨酸的生产,采用不受甲硫氨酸的反馈控制的突变体进行替换。为了用编码不受甲硫氨酸的反馈控制的突变体的metA11(序列号33)替换大肠杆菌中编码高丝氨酸琥珀酰转移酶的染色体上的野生型metA基因(序列号27),构建插入载体pSG-metA11。根据WO2008/127240 A1中给出的说明,获得metA11基因的核苷酸序列信息,并且基于该信息,合成引物(序列号20和序列号21),其含有从metA11基因的起始密码子ATG开始的开放阅读框(ORF)和限制酶(EcoRI和Sacl)的识别位点。使用WO2008/127240 A1中所示的TF4076BJF metA#11菌株作为模板,用由如下的序列表示的引物进行PCR。
序列号20:5'-ggccgaattcatgccgattcgtgtgccgga-3'
序列号21:5'-ggccgagttcttaatccagcgttggattca-3'
使用pfu-X DNA聚合酶(Solgent,SPX16-R250)按如下进行PCR:在95℃下变性30秒,在55℃下退火30秒,并在72℃下延伸1分钟,进行30个从变性到延伸的循环。结果,获得PCR产物,在该PCR产物中在两端均包含EcoRI和Sacl的识别位点的metA11 ORF被扩增。用限制酶EcoRI和Sacl处理通过上上述PCR获得的metA11基因,然后连接到已经用限制酶EcoRI和Sacl处理过的pSG76-C载体(J Bacteriol.1997Jul;179(13):4426-8)。最终,构建含克隆的metA11基因的pSG-metA11重组载体。
4)metA11插入菌株的构建
将在参考实例3)中所构建的metA11插入载体pSG-metA11引入到参考实例2)中所获得的菌株中,然后在LB-Cm培养基(酵母提取物10g/L,NaCl 5g/L,胰蛋白胨10g/L,氯霉素30μg/L)中培养。然后,选择氯霉素抗性的菌落。所选择的转化体是在染色体的metA位置处用pSG-metA11载体插入的一次(primary)菌株。使用表达限制酶I-SceI的pASceP载体(JOURNAL OF BACTERIOLOGY,July 1997,4426-4428)转化插入metA11基因的菌株,其中,限制酶I-SceI切割pSG载体中的I-SceI位点。在LB-Amp培养基(酵母提取物10g/L,NaCl 5g/L,胰蛋白胨10g/L,氯霉素100μg/L)中生长后,选择菌株。在所选择的菌株中,用metA11替换野生型metA,并且从中消除所插入的pSG76-C载体。该菌株被命名为大肠杆菌E.coli CC03-0038。
5)基于生产苏氨酸的菌株构建生产O-琥珀酰高丝氨酸的菌株
通过使用国际专利申请WO 2005/075625中所描述的生产苏氨酸的菌株,即大肠杆菌KCCM 10541P,而不使用野生型W3110,通过在上文1)至4)中所描述的方法相同的方法构建具有生产琥珀酸和O-琥珀酰高丝氨酸能力的菌株,并且所构建的菌株被命名为CJM2-A11。
实施例1:构建用于sucAB表达增强的质粒
1-1.构建具有sucA启动子的用于sucAB表达增强的质粒
从美国国家生物技术信息中心(the National Center for BiotechnologyInformation,USA)的数据库获得由sucAB(sucA GenBank Accession No.BAA35392.1:序列号23,sucB GenBank Accession No.BAA35393.1:序列号25)编码的α-酮戊二酸脱氢酶的核苷酸序列信息,基于此,合成由序列号1和序列号2表示的引物,所述引物可分别由限制酶Hindlll和Xbal识别并包含从ATG(sucAB的ORF起始密码子)开始-188范围的序列,从而获得在sucA启动子控制下的sucAB基因。
序列号1:5'-GGCCAAGCTTGCATCAGGCGTAACAAAGAA-3'
序列号2:5'-GGCCTCTAGATGTCCATCCTTCAGTAATCG-3'
使用野生型大肠杆菌W3110的染色体DNA作为模板,用由序列号1和序列号2表示的引物,通过PCR(聚合酶连锁反应)进行sucAB(编码α-酮戊二酸脱氢酶的基因)的克隆。使用pfu-X DNA聚合酶(Solgent,SPX16-R250)按如下进行PCR[Sambrook等,MolecularCloning,a Laboratory Manual(1989),Cold Spring Harbor Laboratories]:在95℃下变性30秒,在55℃下退火30秒,并在72℃下延伸5分钟,进行30个从变性到延伸的循环。结果,获得包含sucA启动子、sucAB基因以及Hindlll和Xbal的识别位点的大约4.5kb PCR产物。用限制酶Hindlll和Xbal处理所获得的PCR产物。通过使用T4DNA连接酶,将PCR产物连接到已经使用限制酶Hindlll和Xbal预处理的pCL1920载体(Lerner,C.G.and Inouye,M.,Nucl.Acids Res.(1990)18:4631)中,从而构建重组载体pCL PsucA-sucAB。图1说明重组载体pCL PsucA-sucAB的示意图。
1-2.构建具有启动子rmf或trc的用于sucAB表达增强的质粒
从美国国家生物技术信息中心的数据库获得sucAB基因(编码α-酮戊二酸脱氢酶的基因)的核苷酸序列信息(sucA GenBank Accession No.BAA35392.1:序列号23,sucBGenBank Accession No.BAA35393.1:序列号25),基于此,合成具有EcoRV和Hindlll识别位点的、由序列号3和序列号4表示的引物,从而获得sucAB基因。
序列号3:5'-ATCATGCAGAACAGCGCTTTGAA-3'
序列号4:5'-GGCCAAGCTTTGTCCATCCTTCAGTAATCG-3'
使用野生型大肠杆菌W3110的染色体DNA作为模板,用由序列号3和序列号4表示的引物,通过PCR进行sucAB的克隆。使用pfu-X DNA聚合酶(Solgent,SPX16-R250)按如下进行PCR[Sambrook等,Molecular Cloning,a Laboratory Manual(1989),Cold Spring HarborLaboratories]:在95℃下变性30秒,在55℃下退火30秒,并在72℃下延伸5分钟,进行30个从变性到延伸的循环。结果,获得包含sucAB基因和限制酶Hindlll的识别位点的大约4.3kb的PCR产物。用限制酶Hindlll处理所获得的PCR产物。为了替换sucA启动子(即sucAB基因的天然启动子),用限制酶EcoRV和Hindlll处理分别包含启动子rmf和trc的载体pCL_Prmf-gfp(序列号5)和pCL_Ptrc-gfp(序列号6),然后使用T4DNA连接酶(Roche:10481220001)连接PCR产物。结果,构建重组载体pCL_Prmf-sucAB和pCL_Ptrc-sucAB。用绿色荧光蛋白基因gfp引入载体pCL_Prmf-gfp和pCL_Ptrc-gfp,从而测量启动子rmf和trc的强度。同时,将sucAB与载体的启动子区域连接,从而构建含sucAB的载体,其会在启动子rmf和trc的控制下表达。图2和图3是分别说明重组载体pCL_Prmf-sucAB和pCL_Ptrc-sucAB的示意图。
实施例2:构建用于sucAB和metA11表达同时增强的质粒
为了合成O-琥珀酰高丝氨酸,构建从同时表达sucAB和metA11的表达载体。基于编码国际专利公开号WO2008/127240 A1中所描述的TF4076BJF metA#11菌株的O-琥珀酰转移酶突变体的氨基酸序列,获得metA11基因的核苷酸序列信息,并且基于核苷酸序列信息,合成在两端具有限制酶EcoRV和Hindlll的识别位点的、由序列号7和序列号8表示的引物,从而使metA11基因的从ATG至TAA的ORF扩增。
序列号7:5'-GAGTGCGATATC atgccgattcgtgtgccggac-3'
序列号8:5'-GCACTCAAGCTT ttaatccagcgttggatacatg-3'
使用TF4076BJF metA#11作为模板,用由序列号7和序列号8表示的引物进行PCR。使用pfu-X DNA聚合酶(Solgent,SPX16-R250)按如下进行PCR:在95℃下变性30秒,在55℃下退火30秒,并在72℃下延伸1分钟,进行30个从变性到延伸的循环。用限制酶EcoRV和Hindlll处理所获得的PCR产物。用限制酶EcoRV和Hindlll处理载体pCL_Pcysk-gfp(序列号9),然后使用T4DNA连接酶(Roche:10481220001)连接。结果,构建重组载体pCL_Pcysk-metA11。为了向上文所示的构建的载体中插入sucAB,合成具有Hindlll的识别位点的、由序列号10和序列号11表示的引物。
序列号10:5'-GGCCAAGCTTGCATCAGGCGTAACAAAGAA-3'
序列号11:5'-GGCCAAGCTTTGTCCATCCTTCAGTAATCG-3'
为了同时表达metA11和sucAB,构建载体pCL_Pcysk-metA11_PsucA-sucAB。首先,使用野生型大肠杆菌W3110的染色体DNA作为模板,用由序列号10和序列号11表示的引物进行PCR。使用pfu-X DNA聚合酶(Solgent,SPX16-R250)按如下进行PCR[Sambrook等,Molecular Cloning,a Laboratory Manual(1989),Cold Spring Harbor Laboratories]:在95℃下变性30秒,在55℃下退火30秒,并在72℃下延伸1分钟,进行30个从变性到延伸的循环。结果,获得具有在两边包含Hindlll的识别位点的PsucA-sucAB的大约4.5kb的PCR产物。用限制酶Hindlll处理所获得的PCR产物,然后使用T4DNA连接酶(Roche:10481220001)连接到已经用限制酶Hindlll预处理过的pCL_Pcysk-metA11载体。结果,构建重组载体pCL_Pcysk-metA11_PsucA-sucAB。图4是说明重组载体pCL_Pcysk-metA11_PsucA-sucAB的示意图。
实施例3:用于生产琥珀酸的发酵
进行玻瓶培养以研究当在参考实例中所构建的生产O-琥珀酰高丝氨酸的菌株中仅增强sucAB的活性时琥珀酸的生产。用实施例1中所构建的质粒pCL_PsucA-sucAB、pCL_Prmf-sucAB和pCL_Ptrc-sucAB转化参考实例4)的菌株CC03-0038和参考实例5)的菌株CJM2-A11。将菌株涂在含壮观霉素(抗生素)的LB平板培养基上,制备插入了sucAB的菌株。作为对照组,制备用pCL1920载体引入的CC03-0038和CJM2-A11菌株。将具有从国际专利公开号WO10/101360中所描述的沙门氏菌菌株获得的pUR 400质粒的scrO序列的pCC1BAC-scrO载体(序列号34)引入到这些菌株中,以使它们使用原糖作为碳源。将所得到的菌株在具有下表2中所示的组成的培养基中培养,以评价琥珀酸的生产。
将每种菌株接种到培养基中并在33℃下培养过夜。将单菌落接种到2ml的含壮观霉素的LB培养基中,然后在33℃下培养2小时。以OD600=0.5的密度将菌株再次接种到含25mL玻瓶培养基的250ml锥形玻瓶中,然后在33℃下在200rpm下培养33小时。进行高效液相色谱(HPLC),以研究琥珀酸的生产。结果示于表3中。
结果,随着sucAB的表达得到增强,琥珀酸的生产得到提高。该结果表明,sucAB的表达增强具有下调谷氨酸的作用,而且提高了琥珀酰-CoA生产的能量,以提高琥珀酸的生产。当使用葡萄糖作为碳源时,琥珀酸的生产最大提高了30%。随着在需氧条件下菌株中琥珀酸合成得到增强,预期琥珀酸的生产会进一步提高。当使用原糖作为碳源时,琥珀酸的生产提高至当使用葡萄糖作为碳源时的生产。结果示于表4中。
表1:构建葡萄糖玻瓶培养基
【表1】
表2:构建原糖玻瓶培养基
【表2】
表3:使用葡萄糖经由玻瓶培养生产琥珀酸
【表3】
表4:使用原糖经由玻瓶培养生产琥珀酸
【表4】
实施例4:用于生产O-琥珀酰高丝氨酸的发酵
为了在CC03-0038和CJM2-A11菌株中同时增强sucAB和metA基因的表达,将这些菌株用实施例2中所构建的pCL_Pcysk-metA11_PsucA-sucAB质粒转化,然后在锥形瓶中培养以研究O-琥珀酰高丝氨酸的生产。玻瓶培养基的组成如表1中所示。参考实施例4)中metA11基因的增强表达是在原始启动子的控制下进行的。因此,为了增强其表达,将metA11克隆到可以由cysK启动子表达的载体中,其中同时诱导sucAB的表达。
用实施例2中所构建的pCL_Pcysk-metA11_PsucA-sucAB质粒转化CC03-0038。将菌株涂在含壮观霉素(抗生素)的LB平板培养基上,以制备经转化的菌株。作为对照组,用pCL1920载体转化CC03-0038和CJM2-A11。通过引入具有从韩国专利申请号2009-0018128中所描述的沙门氏菌菌株获得的pUR 400质粒的scrO序列的pCC1BAC-scrO载体引入到菌株中,构建能够使用原糖作为碳源的菌株。将菌株在具有表2中所示的组成的原糖玻瓶培养基中培养,以评价O-琥珀酰高丝氨酸的生产。为了评价的目的,将上文中所描述的一些菌株接种到培养基中,并在33℃下培养过夜。将单菌落接种到2ml的含壮观霉素的LB培养基中,然后在33℃下培养2小时。以OD600=0.5的密度将菌株再次接种到含25mL玻瓶培养基的250ml锥形玻瓶中,然后在33℃下在200rpm下培养33小时。进行HPLC,以研究O-琥珀酰高丝氨酸的生产。结果示于表5中。
结果表明,在引入pCL_Pcysk-metA11_PsucA-sucAB的菌株中,O-琥珀酰高丝氨酸的浓度提高。与对照组相比,产量(production yield)提高了约40%。随着O-琥珀酰高丝氨酸提高,谷氨酸盐/酯和高丝氨酸的量减少。结果表明,sucAB的表达增强在供应琥珀酰-CoA中具有重要作用。使用琥珀酰-CoA作为模板,当metA表达同时增强时,O-琥珀酰高丝氨酸的浓度迅速提高。
表5:使用葡萄糖经由玻瓶培养生产O-琥珀酰高丝氨酸
【表5】
表6:使用原糖经由玻瓶培养生产O-琥珀酰高丝氨酸
【表6】
当用根据本发明的实施方式用含有sucAB的载体转化属于埃希氏菌属的微生物以增强sucAB的表达时,确认琥珀酰-CoA和琥珀酸的生产能力提高。因此,sucAB的表达增强可以被应用于具有与上文所述的中心碳代谢方面相同的TCA循环的那些微生物,例如诸如酵母和放线菌等的那些微生物。
经改造的CC03-0038/pCL_Pcysk-metA11_PsucA-sucAB菌株生产O-琥珀酰高丝氨酸的能力已被确认,其被命名为CC03-0157,并且根据布达佩斯条约于2013年11月22日保藏在韩国微生物保藏中心(KCCM)(保藏号:KCCM11488P)。
序列表自由文本
本文中使用的由序列号1至序列号34表示的序列示于所附的序列表中。
序列表
<110> CJ第一制糖株式会社
<120> 能生产O-琥珀酰高丝氨酸或琥珀酸的微生物和用其生产琥珀酸或O-琥珀酰高丝氨酸方法
<130> PX049610
<150> KR10-2014-0067787
<151> 2014-06-03
<160> 34
<170> PatentIn version 3.5
<210> 1
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 用于Psuc_sucAB的正向引物
<400> 1
ggccaagctt gcatcaggcg taacaaagaa 30
<210> 2
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 用于Psuc_sucAB的反向引物
<400> 2
ggcctctaga tgtccatcct tcagtaatcg 30
<210> 3
<211> 23
<212> DNA
<213> 人工序列
<220>
<223> 用于sucAB的正向引物
<400> 3
atcatgcaga acagcgcttt gaa 23
<210> 4
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 用于sucAB的反向引物
<400> 4
ggccaagctt tgtccatcct tcagtaatcg 30
<210> 5
<211> 5545
<212> DNA
<213> 人工序列
<220>
<223> pCL_Prmf-gfp
<400> 5
atcgaatcaa agctgccgac aacacgggag ccagtgacgc ctcccgtggg gaaaaaatca 60
tggcaattct ggaagaaata gcgctttcag ccggcaaacc ggctgaagcc ggatctgcga 120
ttctgataac aaactagcaa caccagaaca gcccgtttgc gggcagcaaa acccgtggga 180
attaattccc ctgctcgcgc aggctgggtg ccaagctctc gggtaacatc aaggcccgat 240
ccttggagcc cttgccctcc cgcacgatga tcgtgccgtg atcgaaatcc agatccttga 300
cccgcagttg caaaccctca ctgatccgca tgcccgttcc atacagaagc tgggcgaaca 360
aacgatgctc gccttccaga aaaccgagga tgcgaaccac ttcatccggg gtcagcacca 420
ccggcaagcg ccgcgacggc cgaggtcttc cgatctcctg aagccagggc agatccgtgc 480
acagcacctt gccgtagaag aacagcaagg ccgccaatgc ctgacgatgc gtggagaccg 540
aaaccttgcg ctcgttcgcc agccaggaca gaaatgcctc gacttcgctg ctgcccaagg 600
ttgccgggtg acgcacaccg tggaaacgga tgaaggcacg aacccagtgg acataagcct 660
gttcggttcg taagctgtaa tgcaagtagc gtatgcgctc acgcaactgg tccagaacct 720
tgaccgaacg cagcggtggt aacggcgcag tggcggtttt catggcttgt tatgactgtt 780
tttttggggt acagtctatg cctcgggcat ccaagcagca agcgcgttac gccgtgggtc 840
gatgtttgat gttatggagc agcaacgatg ttacgcagca gggcagtcgc cctaaaacaa 900
agttaaacat catgagggaa gcggtgatcg ccgaagtatc gactcaacta tcagaggtag 960
ttggcgtcat cgagcgccat ctcgaaccga cgttgctggc cgtacatttg tacggctccg 1020
cagtggatgg cggcctgaag ccacacagtg atattgattt gctggttacg gtgaccgtaa 1080
ggcttgatga aacaacgcgg cgagctttga tcaacgacct tttggaaact tcggcttccc 1140
ctggagagag cgagattctc cgcgctgtag aagtcaccat tgttgtgcac gacgacatca 1200
ttccgtggcg ttatccagct aagcgcgaac tgcaatttgg agaatggcag cgcaatgaca 1260
ttcttgcagg tatcttcgag ccagccacga tcgacattga tctggctatc ttgctgacaa 1320
aagcaagaga acatagcgtt gccttggtag gtccagcggc ggaggaactc tttgatccgg 1380
ttcctgaaca ggatctattt gaggcgctaa atgaaacctt aacgctatgg aactcgccgc 1440
ccgactgggc tggcgatgag cgaaatgtag tgcttacgtt gtcccgcatt tggtacagcg 1500
cagtaaccgg caaaatcgcg ccgaaggatg tcgctgccga ctgggcaatg gagcgcctgc 1560
cggcccagta tcagcccgtc atacttgaag ctagacaggc ttatcttgga caagaagaag 1620
atcgcttggc ctcgcgcgca gatcagttgg aagaatttgt ccactacgtg aaaggcgaga 1680
tcaccaaggt agtcggcaaa taatgtctaa caattcgttc aagccgacgc cgcttcgcgg 1740
cgcggcttaa ctcaagcgtt agatgcacta agcacataat tgctcacagc caaactatca 1800
ggtcaagtct gcttttatta tttttaagcg tgcataataa gccctacaca aattgggaga 1860
tatatcatga aaggctggct ttttcttgtt atcgcaatag ttggcgaagt aatcgcaaca 1920
tccgcattaa aatctagcga gggctttact aagctcgtca gcgggtgttg gcgggtgtcg 1980
gggctggctt aactatgcgg catcagagca gattgtactg agagtgcacc atatgcggtg 2040
tgaaataccg cacagatgcg taaggagaaa ataccgcatc aggcgccatt cgccattcag 2100
gctgcgcaac tgttgggaag ggcgatcggt gcgggcctct tcgctattac gccagctggc 2160
gaaaggggga tgtgctgcaa ggcgattaag ttgggtaacg ccagggtttt cccagtcacg 2220
acgttgtaaa acgacggcca gtgaattcga gctcggtacc caggaagccg cttctattgc 2280
acaagagata aagcgtctac cttaattata aagatttgta aatataaccg tctccggtat 2340
gttgcctgag gcggtttttt tgtctctaac gtgcggaaaa atttgttcct cttcacattt 2400
tttgtacaac cgacatgccc gtgtagctca caaatatgac agtggcgtga attttgcgca 2460
ttgacggcag ttatgattcg cggtattgct taactgtgat tgcacattta gtaatcactg 2520
ttttcttttc caccagaaac cagtatgagg gaaagatatc atgagtaaag gagaagaact 2580
tttcactgga gttgtcccaa ttcttgttga attagatggt gatgttaatg ggcacaaatt 2640
ttctgtcagt ggagagggtg aaggtgatgc aacatacgga aaacttaccc ttaaatttat 2700
ttgcactact ggaaaactac ctgttccatg gccaacactt gtcactactt tctcttatgg 2760
tgttcaatgc ttttcccgtt atccggatca tatgaaacgg catgactttt tcaagagtgc 2820
catgcccgaa ggttatgtac aggaacgcac tatatctttc aaagatgacg ggaactacaa 2880
gacgcgtgct gaagtcaagt ttgaaggtga tacccttgtt aatcgtatcg agttaaaagg 2940
tattgatttt aaagaagatg gaaacattct cggacacaaa ctcgagtaca actataactc 3000
acacaatgta tacatcacgg cagacaaaca aaagaatgga atcaaagcta acttcaaaat 3060
tcgccacaac attgaagatg gatccgttca actagcagac cattatcaac aaaatactcc 3120
aattggcgat ggccctgtcc ttttaccaga caaccattac ctgtcgacac aatctgccct 3180
ttcgaaagat cccaacgaaa agcgtgacca catggtcctt cttgagtttg taactgctgc 3240
tgggattaca catggcatgg atgagctcta caaataactg caggcatgca agcttggcgt 3300
aatcatggtc atagctgttt cctgtgtgaa attgttatcc gctcacaatt ccacacaaca 3360
tacgagccgg aagcataaag tgtaaagcct ggggtgccta atgagtgagc taactcacat 3420
taattgcgtt gcgctcactg cccgctttcc agtcgggaaa cctgtcgtgc cagctgcatt 3480
aatgaatcgg ccaacgcgaa ttcccgacag taagacgggt aagcctgttg atgataccgc 3540
tgccttactg ggtgcattag ccagtctgaa tgacctgtca cgggataatc cgaagtggtc 3600
agactggaaa atcagagggc aggaactgct gaacagcaaa aagtcagata gcaccacata 3660
gcagacccgc cataaaacgc cctgagaagc ccgtgacggg cttttcttgt attatgggta 3720
gtttccttgc atgaatccat aaaaggcgcc tgtagtgcca tttaccccca ttcactgcca 3780
gagccgtgag cgcagcgaac tgaatgtcac gaaaaagaca gcgactcagg tgcctgatgg 3840
tcggagacaa aaggaatatt cagcgatttg cccgagcttg cgagggtgct acttaagcct 3900
ttagggtttt aaggtctgtt ttgtagagga gcaaacagcg tttgcgacat ccttttgtaa 3960
tactgcggaa ctgactaaag tagtgagtta tacacagggc tgggatctat tctttttatc 4020
tttttttatt ctttctttat tctataaatt ataaccactt gaatataaac aaaaaaaaca 4080
cacaaaggtc tagcggaatt tacagagggt ctagcagaat ttacaagttt tccagcaaag 4140
gtctagcaga atttacagat acccacaact caaaggaaaa ggactagtaa ttatcattga 4200
ctagcccatc tcaattggta tagtgattaa aatcacctag accaattgag atgtatgtct 4260
gaattagttg ttttcaaagc aaatgaacta gcgattagtc gctatgactt aacggagcat 4320
gaaaccaagc taattttatg ctgtgtggca ctactcaacc ccacgattga aaaccctaca 4380
aggaaagaac ggacggtatc gttcacttat aaccaatacg ctcagatgat gaacatcagt 4440
agggaaaatg cttatggtgt attagctaaa gcaaccagag agctgatgac gagaactgtg 4500
gaaatcagga atcctttggt taaaggcttt gagattttcc agtggacaaa ctatgccaag 4560
ttctcaagcg aaaaattaga attagttttt agtgaagaga tattgcctta tcttttccag 4620
ttaaaaaaat tcataaaata taatctggaa catgttaagt cttttgaaaa caaatactct 4680
atgaggattt atgagtggtt attaaaagaa ctaacacaaa agaaaactca caaggcaaat 4740
atagagatta gccttgatga atttaagttc atgttaatgc ttgaaaataa ctaccatgag 4800
tttaaaaggc ttaaccaatg ggttttgaaa ccaataagta aagatttaaa cacttacagc 4860
aatatgaaat tggtggttga taagcgaggc cgcccgactg atacgttgat tttccaagtt 4920
gaactagata gacaaatgga tctcgtaacc gaacttgaga acaaccagat aaaaatgaat 4980
ggtgacaaaa taccaacaac cattacatca gattcctacc tacataacgg actaagaaaa 5040
acactacacg atgctttaac tgcaaaaatt cagctcacca gttttgaggc aaaatttttg 5100
agtgacatgc aaagtaagta tgatctcaat ggttcgttct catggctcac gcaaaaacaa 5160
cgaaccacac tagagaacat actggctaaa tacggaagga tctgaggttc ttatggctct 5220
tgtatctatc agtgaagcat caagactaac aaacaaaagt agaacaactg ttcaccgtta 5280
catatcaaag ggaaaactgt ccatatgcac agatgaaaac ggtgtaaaaa agatagatac 5340
atcagagctt ttacgagttt ttggtgcatt caaagctgtt caccatgaac agatcgacaa 5400
tgtaacagat gaacagcatg taacacctaa tagaacaggt gaaaccagta aaacaaagca 5460
actagaacat gaaattgaac acctgagaca acttgttaca gctcaacagt cacacataga 5520
cagcctgaaa caggcgatgc tgctt 5545
<210> 6
<211> 5727
<212> DNA
<213> 人工序列
<220>
<223> pCL_Ptrc-gfp
<400> 6
tttctttatt ctataaatta taaccacttg aatataaaca aaaaaaacac acaaaggtct 60
agcggaattt acagagggtc tagcagaatt tacaagtttt ccagcaaagg tctagcagaa 120
tttacagata cccacaactc aaaggaaaag gactagtaat tatcattgac tagcccatct 180
caattggtat agtgattaaa atcacctaga ccaattgaga tgtatgtctg aattagttgt 240
tttcaaagca aatgaactag cgattagtcg ctatgactta acggagcatg aaaccaagct 300
aattttatgc tgtgtggcac tactcaaccc cacgattgaa aaccctacaa ggaaagaacg 360
gacggtatcg ttcacttata accaatacgc tcagatgatg aacatcagta gggaaaatgc 420
ttatggtgta ttagctaaag caaccagaga gctgatgacg agaactgtgg aaatcaggaa 480
tcctttggtt aaaggctttg agattttcca gtggacaaac tatgccaagt tctcaagcga 540
aaaattagaa ttagttttta gtgaagagat attgccttat cttttccagt taaaaaaatt 600
cataaaatat aatctggaac atgttaagtc ttttgaaaac aaatactcta tgaggattta 660
tgagtggtta ttaaaagaac taacacaaaa gaaaactcac aaggcaaata tagagattag 720
ccttgatgaa tttaagttca tgttaatgct tgaaaataac taccatgagt ttaaaaggct 780
taaccaatgg gttttgaaac caataagtaa agatttaaac acttacagca atatgaaatt 840
ggtggttgat aagcgaggcc gcccgactga tacgttgatt ttccaagttg aactagatag 900
acaaatggat ctcgtaaccg aacttgagaa caaccagata aaaatgaatg gtgacaaaat 960
accaacaacc attacatcag attcctacct acataacgga ctaagaaaaa cactacacga 1020
tgctttaact gcaaaaattc agctcaccag ttttgaggca aaatttttga gtgacatgca 1080
aagtaagtat gatctcaatg gttcgttctc atggctcacg caaaaacaac gaaccacact 1140
agagaacata ctggctaaat acggaaggat ctgaggttct tatggctctt gtatctatca 1200
gtgaagcatc aagactaaca aacaaaagta gaacaactgt tcaccgttac atatcaaagg 1260
gaaaactgtc catatgcaca gatgaaaacg gtgtaaaaaa gatagataca tcagagcttt 1320
tacgagtttt tggtgcattc aaagctgttc accatgaaca gatcgacaat gtaacagatg 1380
aacagcatgt aacacctaat agaacaggtg aaaccagtaa aacaaagcaa ctagaacatg 1440
aaattgaaca cctgagacaa cttgttacag ctcaacagtc acacatagac agcctgaaac 1500
aggcgatgct gcttatcgaa tcaaagctgc cgacaacacg ggagccagtg acgcctcccg 1560
tggggaaaaa atcatggcaa ttctggaaga aatagcgctt tcagccggca aaccggctga 1620
agccggatct gcgattctga taacaaacta gcaacaccag aacagcccgt ttgcgggcag 1680
caaaacccgt gggaattaat tcccctgctc gcgcaggctg ggtgccaagc tctcgggtaa 1740
catcaaggcc cgatccttgg agcccttgcc ctcccgcacg atgatcgtgc cgtgatcgaa 1800
atccagatcc ttgacccgca gttgcaaacc ctcactgatc cgcatgcccg ttccatacag 1860
aagctgggcg aacaaacgat gctcgccttc cagaaaaccg aggatgcgaa ccacttcatc 1920
cggggtcagc accaccggca agcgccgcga cggccgaggt cttccgatct cctgaagcca 1980
gggcagatcc gtgcacagca ccttgccgta gaagaacagc aaggccgcca atgcctgacg 2040
atgcgtggag accgaaacct tgcgctcgtt cgccagccag gacagaaatg cctcgacttc 2100
gctgctgccc aaggttgccg ggtgacgcac accgtggaaa cggatgaagg cacgaaccca 2160
gtggacataa gcctgttcgg ttcgtaagct gtaatgcaag tagcgtatgc gctcacgcaa 2220
ctggtccaga accttgaccg aacgcagcgg tggtaacggc gcagtggcgg ttttcatggc 2280
ttgttatgac tgtttttttg gggtacagtc tatgcctcgg gcatccaagc agcaagcgcg 2340
ttacgccgtg ggtcgatgtt tgatgttatg gagcagcaac gatgttacgc agcagggcag 2400
tcgccctaaa acaaagttaa acatcatgag ggaagcggtg atcgccgaag tatcgactca 2460
actatcagag gtagttggcg tcatcgagcg ccatctcgaa ccgacgttgc tggccgtaca 2520
tttgtacggc tccgcagtgg atggcggcct gaagccacac agtgatattg atttgctggt 2580
tacggtgacc gtaaggcttg atgaaacaac gcggcgagct ttgatcaacg accttttgga 2640
aacttcggct tcccctggag agagcgagat tctccgcgct gtagaagtca ccattgttgt 2700
gcacgacgac atcattccgt ggcgttatcc agctaagcgc gaactgcaat ttggagaatg 2760
gcagcgcaat gacattcttg caggtatctt cgagccagcc acgatcgaca ttgatctggc 2820
tatcttgctg acaaaagcaa gagaacatag cgttgccttg gtaggtccag cggcggagga 2880
actctttgat ccggttcctg aacaggatct atttgaggcg ctaaatgaaa ccttaacgct 2940
atggaactcg ccgcccgact gggctggcga tgagcgaaat gtagtgctta cgttgtcccg 3000
catttggtac agcgcagtaa ccggcaaaat cgcgccgaag gatgtcgctg ccgactgggc 3060
aatggagcgc ctgccggccc agtatcagcc cgtcatactt gaagctagac aggcttatct 3120
tggacaagaa gaagatcgct tggcctcgcg cgcagatcag ttggaagaat ttgtccacta 3180
cgtgaaaggc gagatcacca aggtagtcgg caaataatgt ctaacaattc gttcaagccg 3240
acgccgcttc gcggcgcggc ttaactcaag cgttagatgc actaagcaca taattgctca 3300
cagccaaact atcaggtcaa gtctgctttt attattttta agcgtgcata ataagcccta 3360
cacaaattgg gagatatatc atgaaaggct ggctttttct tgttatcgca atagttggcg 3420
aagtaatcgc aacatccgca ttaaaatcta gcgagggctt tactaagctc gtcagcgggt 3480
gttggcgggt gtcggggctg gcttaactat gcggcatcag agcagattgt actgagagtg 3540
caccatatgc ggtgtgaaat accgcacaga tgcgtaagga gaaaataccg catcaggcgc 3600
cattcgccat tcaggctgcg caactgttgg gaagggcgat cggtgcgggc ctcttcgcta 3660
ttacgccagc tggcgaaagg gggatgtgct gcaaggcgat taagttgggt aacgccaggg 3720
ttttcccagt cacgacgttg taaaacgacg gccagtgaat tcgagctcgg tacccgcttg 3780
ctgcaactct ctcagggcca ggcggtgaag ggcaatcagc tgttgcccgt ctcactggtg 3840
aaaagaaaaa ccaccctggc gcccaatacg caaaccgcct ctccccgcgc gttggccgat 3900
tcattaatgc agctggcacg acaggtttcc cgactggaaa gcgggcagtg agcgcaacgc 3960
aattaatgta agttagcgcg aattgatctg gtttgacagc ttatcatcga ctgcacggtg 4020
caccaatgct tctggcgtca ggcagccatc ggaagctgtg gtatggctgt gcaggtcgta 4080
aatcactgca taattcgtgt cgctcaaggc gcactcccgt tctggataat gttttttgcg 4140
ccgacatcat aacggttctg gcaaatattc tgaaatgagc tgttgacaat taatcatccg 4200
gctcgtataa tgtgtggaat tgtgagcgga taacaatttc acacaggaaa gatatcatga 4260
gtaaaggaga agaacttttc actggagttg tcccaattct tgttgaatta gatggtgatg 4320
ttaatgggca caaattttct gtcagtggag agggtgaagg tgatgcaaca tacggaaaac 4380
ttacccttaa atttatttgc actactggaa aactacctgt tccatggcca acacttgtca 4440
ctactttctc ttatggtgtt caatgctttt cccgttatcc ggatcatatg aaacggcatg 4500
actttttcaa gagtgccatg cccgaaggtt atgtacagga acgcactata tctttcaaag 4560
atgacgggaa ctacaagacg cgtgctgaag tcaagtttga aggtgatacc cttgttaatc 4620
gtatcgagtt aaaaggtatt gattttaaag aagatggaaa cattctcgga cacaaactcg 4680
agtacaacta taactcacac aatgtataca tcacggcaga caaacaaaag aatggaatca 4740
aagctaactt caaaattcgc cacaacattg aagatggatc cgttcaacta gcagaccatt 4800
atcaacaaaa tactccaatt ggcgatggcc ctgtcctttt accagacaac cattacctgt 4860
cgacacaatc tgccctttcg aaagatccca acgaaaagcg tgaccacatg gtccttcttg 4920
agtttgtaac tgctgctggg attacacatg gcatggatga gctctacaaa taactgcagg 4980
catgcaagct tggcgtaatc atggtcatag ctgtttcctg tgtgaaattg ttatccgctc 5040
acaattccac acaacatacg agccggaagc ataaagtgta aagcctgggg tgcctaatga 5100
gtgagctaac tcacattaat tgcgttgcgc tcactgcccg ctttccagtc gggaaacctg 5160
tcgtgccagc tgcattaatg aatcggccaa cgcgaattcc cgacagtaag acgggtaagc 5220
ctgttgatga taccgctgcc ttactgggtg cattagccag tctgaatgac ctgtcacggg 5280
ataatccgaa gtggtcagac tggaaaatca gagggcagga actgctgaac agcaaaaagt 5340
cagatagcac cacatagcag acccgccata aaacgccctg agaagcccgt gacgggcttt 5400
tcttgtatta tgggtagttt ccttgcatga atccataaaa ggcgcctgta gtgccattta 5460
cccccattca ctgccagagc cgtgagcgca gcgaactgaa tgtcacgaaa aagacagcga 5520
ctcaggtgcc tgatggtcgg agacaaaagg aatattcagc gatttgcccg agcttgcgag 5580
ggtgctactt aagcctttag ggttttaagg tctgttttgt agaggagcaa acagcgtttg 5640
cgacatcctt ttgtaatact gcggaactga ctaaagtagt gagttataca cagggctggg 5700
atctattctt tttatctttt tttattc 5727
<210> 7
<211> 33
<212> DNA
<213> 人工序列
<220>
<223> 用于metA11的正向引物
<400> 7
gagtgcgata tcatgccgat tcgtgtgccg gac 33
<210> 8
<211> 34
<212> DNA
<213> 人工序列
<220>
<223> 用于metA11的反向引物
<400> 8
gcactcaagc ttttaatcca gcgttggata catg 34
<210> 9
<211> 5545
<212> DNA
<213> 人工序列
<220>
<223> pCL_Pcysk-gfp
<400> 9
acactacacg atgctttaac tgcaaaaatt cagctcacca gttttgaggc aaaatttttg 60
agtgacatgc aaagtaagta tgatctcaat ggttcgttct catggctcac gcaaaaacaa 120
cgaaccacac tagagaacat actggctaaa tacggaagga tctgaggttc ttatggctct 180
tgtatctatc agtgaagcat caagactaac aaacaaaagt agaacaactg ttcaccgtta 240
catatcaaag ggaaaactgt ccatatgcac agatgaaaac ggtgtaaaaa agatagatac 300
atcagagctt ttacgagttt ttggtgcatt caaagctgtt caccatgaac agatcgacaa 360
tgtaacagat gaacagcatg taacacctaa tagaacaggt gaaaccagta aaacaaagca 420
actagaacat gaaattgaac acctgagaca acttgttaca gctcaacagt cacacataga 480
cagcctgaaa caggcgatgc tgcttatcga atcaaagctg ccgacaacac gggagccagt 540
gacgcctccc gtggggaaaa aatcatggca attctggaag aaatagcgct ttcagccggc 600
aaaccggctg aagccggatc tgcgattctg ataacaaact agcaacacca gaacagcccg 660
tttgcgggca gcaaaacccg tgggaattaa ttcccctgct cgcgcaggct gggtgccaag 720
ctctcgggta acatcaaggc ccgatccttg gagcccttgc cctcccgcac gatgatcgtg 780
ccgtgatcga aatccagatc cttgacccgc agttgcaaac cctcactgat ccgcatgccc 840
gttccataca gaagctgggc gaacaaacga tgctcgcctt ccagaaaacc gaggatgcga 900
accacttcat ccggggtcag caccaccggc aagcgccgcg acggccgagg tcttccgatc 960
tcctgaagcc agggcagatc cgtgcacagc accttgccgt agaagaacag caaggccgcc 1020
aatgcctgac gatgcgtgga gaccgaaacc ttgcgctcgt tcgccagcca ggacagaaat 1080
gcctcgactt cgctgctgcc caaggttgcc gggtgacgca caccgtggaa acggatgaag 1140
gcacgaaccc agtggacata agcctgttcg gttcgtaagc tgtaatgcaa gtagcgtatg 1200
cgctcacgca actggtccag aaccttgacc gaacgcagcg gtggtaacgg cgcagtggcg 1260
gttttcatgg cttgttatga ctgttttttt ggggtacagt ctatgcctcg ggcatccaag 1320
cagcaagcgc gttacgccgt gggtcgatgt ttgatgttat ggagcagcaa cgatgttacg 1380
cagcagggca gtcgccctaa aacaaagtta aacatcatga gggaagcggt gatcgccgaa 1440
gtatcgactc aactatcaga ggtagttggc gtcatcgagc gccatctcga accgacgttg 1500
ctggccgtac atttgtacgg ctccgcagtg gatggcggcc tgaagccaca cagtgatatt 1560
gatttgctgg ttacggtgac cgtaaggctt gatgaaacaa cgcggcgagc tttgatcaac 1620
gaccttttgg aaacttcggc ttcccctgga gagagcgaga ttctccgcgc tgtagaagtc 1680
accattgttg tgcacgacga catcattccg tggcgttatc cagctaagcg cgaactgcaa 1740
tttggagaat ggcagcgcaa tgacattctt gcaggtatct tcgagccagc cacgatcgac 1800
attgatctgg ctatcttgct gacaaaagca agagaacata gcgttgcctt ggtaggtcca 1860
gcggcggagg aactctttga tccggttcct gaacaggatc tatttgaggc gctaaatgaa 1920
accttaacgc tatggaactc gccgcccgac tgggctggcg atgagcgaaa tgtagtgctt 1980
acgttgtccc gcatttggta cagcgcagta accggcaaaa tcgcgccgaa ggatgtcgct 2040
gccgactggg caatggagcg cctgccggcc cagtatcagc ccgtcatact tgaagctaga 2100
caggcttatc ttggacaaga agaagatcgc ttggcctcgc gcgcagatca gttggaagaa 2160
tttgtccact acgtgaaagg cgagatcacc aaggtagtcg gcaaataatg tctaacaatt 2220
cgttcaagcc gacgccgctt cgcggcgcgg cttaactcaa gcgttagatg cactaagcac 2280
ataattgctc acagccaaac tatcaggtca agtctgcttt tattattttt aagcgtgcat 2340
aataagccct acacaaattg ggagatatat catgaaaggc tggctttttc ttgttatcgc 2400
aatagttggc gaagtaatcg caacatccgc attaaaatct agcgagggct ttactaagct 2460
cgtcagcggg tgttggcggg tgtcggggct ggcttaacta tgcggcatca gagcagattg 2520
tactgagagt gcaccatatg cggtgtgaaa taccgcacag atgcgtaagg agaaaatacc 2580
gcatcaggcg ccattcgcca ttcaggctgc gcaactgttg ggaagggcga tcggtgcggg 2640
cctcttcgct attacgccag ctggcgaaag ggggatgtgc tgcaaggcga ttaagttggg 2700
taacgccagg gttttcccag tcacgacgtt gtaaaacgac ggccagtgaa ttcgagctcg 2760
gtaccccagc ctgtttacga tgatcccgct gcttaatctg ttcatcatgc ccgttgccgt 2820
ttgtggcgcg acggcgatgt gggtcgattg ctatcgcgat aaacacgcga tgtggcggta 2880
acaatctacc ggttattttg taaaccgttt gtgtgaaaca ggggtggctt atgccgcccc 2940
ttattccatc ttgcatgtca ttatttccct tctgtatata gatatgctaa atccttactt 3000
ccgcatattc tctgagcggg tatgctacct gttgtatccc aatttcatac agttaaggag 3060
atatcatgag taaaggagaa gaacttttca ctggagttgt cccaattctt gttgaattag 3120
atggtgatgt taatgggcac aaattttctg tcagtggaga gggtgaaggt gatgcaacat 3180
acggaaaact tacccttaaa tttatttgca ctactggaaa actacctgtt ccatggccaa 3240
cacttgtcac tactttctct tatggtgttc aatgcttttc ccgttatccg gatcatatga 3300
aacggcatga ctttttcaag agtgccatgc ccgaaggtta tgtacaggaa cgcactatat 3360
ctttcaaaga tgacgggaac tacaagacgc gtgctgaagt caagtttgaa ggtgataccc 3420
ttgttaatcg tatcgagtta aaaggtattg attttaaaga agatggaaac attctcggac 3480
acaaactcga gtacaactat aactcacaca atgtatacat cacggcagac aaacaaaaga 3540
atggaatcaa agctaacttc aaaattcgcc acaacattga agatggatcc gttcaactag 3600
cagaccatta tcaacaaaat actccaattg gcgatggccc tgtcctttta ccagacaacc 3660
attacctgtc gacacaatct gccctttcga aagatcccaa cgaaaagcgt gaccacatgg 3720
tccttcttga gtttgtaact gctgctggga ttacacatgg catggatgag ctctacaaat 3780
aactgcaggc atgcaagctt ggcgtaatca tggtcatagc tgtttcctgt gtgaaattgt 3840
tatccgctca caattccaca caacatacga gccggaagca taaagtgtaa agcctggggt 3900
gcctaatgag tgagctaact cacattaatt gcgttgcgct cactgcccgc tttccagtcg 3960
ggaaacctgt cgtgccagct gcattaatga atcggccaac gcgaattccc gacagtaaga 4020
cgggtaagcc tgttgatgat accgctgcct tactgggtgc attagccagt ctgaatgacc 4080
tgtcacggga taatccgaag tggtcagact ggaaaatcag agggcaggaa ctgctgaaca 4140
gcaaaaagtc agatagcacc acatagcaga cccgccataa aacgccctga gaagcccgtg 4200
acgggctttt cttgtattat gggtagtttc cttgcatgaa tccataaaag gcgcctgtag 4260
tgccatttac ccccattcac tgccagagcc gtgagcgcag cgaactgaat gtcacgaaaa 4320
agacagcgac tcaggtgcct gatggtcgga gacaaaagga atattcagcg atttgcccga 4380
gcttgcgagg gtgctactta agcctttagg gttttaaggt ctgttttgta gaggagcaaa 4440
cagcgtttgc gacatccttt tgtaatactg cggaactgac taaagtagtg agttatacac 4500
agggctggga tctattcttt ttatcttttt ttattctttc tttattctat aaattataac 4560
cacttgaata taaacaaaaa aaacacacaa aggtctagcg gaatttacag agggtctagc 4620
agaatttaca agttttccag caaaggtcta gcagaattta cagataccca caactcaaag 4680
gaaaaggact agtaattatc attgactagc ccatctcaat tggtatagtg attaaaatca 4740
cctagaccaa ttgagatgta tgtctgaatt agttgttttc aaagcaaatg aactagcgat 4800
tagtcgctat gacttaacgg agcatgaaac caagctaatt ttatgctgtg tggcactact 4860
caaccccacg attgaaaacc ctacaaggaa agaacggacg gtatcgttca cttataacca 4920
atacgctcag atgatgaaca tcagtaggga aaatgcttat ggtgtattag ctaaagcaac 4980
cagagagctg atgacgagaa ctgtggaaat caggaatcct ttggttaaag gctttgagat 5040
tttccagtgg acaaactatg ccaagttctc aagcgaaaaa ttagaattag tttttagtga 5100
agagatattg ccttatcttt tccagttaaa aaaattcata aaatataatc tggaacatgt 5160
taagtctttt gaaaacaaat actctatgag gatttatgag tggttattaa aagaactaac 5220
acaaaagaaa actcacaagg caaatataga gattagcctt gatgaattta agttcatgtt 5280
aatgcttgaa aataactacc atgagtttaa aaggcttaac caatgggttt tgaaaccaat 5340
aagtaaagat ttaaacactt acagcaatat gaaattggtg gttgataagc gaggccgccc 5400
gactgatacg ttgattttcc aagttgaact agatagacaa atggatctcg taaccgaact 5460
tgagaacaac cagataaaaa tgaatggtga caaaatacca acaaccatta catcagattc 5520
ctacctacat aacggactaa gaaaa 5545
<210> 10
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 用于sucAB的正向引物
<400> 10
ggccaagctt gcatcaggcg taacaaagaa 30
<210> 11
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 用于sucAB的反向引物
<400> 11
ggccaagctt tgtccatcct tcagtaatcg 30
<210> 12
<211> 70
<212> DNA
<213> 人工序列
<220>
<223> metB缺失盒的正向引物
<400> 12
ttactctggt gcctgacatt tcaccgacaa agcccaggga acttcatcac gtgtaggctg 60
gagctgcttc 70
<210> 13
<211> 70
<212> DNA
<213> 人工序列
<220>
<223> metB缺失盒的反向引物
<400> 13
cgctgcgcca gctccatacg cggcaccagc gttcgcaacc cacgtagcag catatgaata 60
tcctccttag 70
<210> 14
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 用于metB缺失的正向引物
<400> 14
tattcgccgc tccattcagc 20
<210> 15
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 用于metB缺失的反向引物
<400> 15
taccccttgt ttgcagcccg 20
<210> 16
<211> 70
<212> DNA
<213> 人工序列
<220>
<223> 用于thrB缺失盒的正向引物
<400> 16
catggttaaa gtttatgccc cggcttccag tgccaatatg agcgtcgggt gtgtaggctg 60
gagctgcttc 70
<210> 17
<211> 70
<212> DNA
<213> 人工序列
<220>
<223> 用于thrB缺失盒的反向引物
<400> 17
ggagataccg ctcgctaccg cgccgatttc cgcgaccgcc tgccgcgcct catatgaata 60
tcctccttag 70
<210> 18
<211> 19
<212> DNA
<213> 人工序列
<220>
<223> 用于thrB缺失的正向引物
<400> 18
actcgacgat ctctttgcc 19
<210> 19
<211> 21
<212> DNA
<213> 用于thrB缺失的反向引物
<400> 19
acgccgagag gatcttcgca g 21
<210> 20
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 用于metA11的正向引物
<400> 20
ggccgaattc atgccgattc gtgtgccgga 30
<210> 21
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 用于metA11的反向引物
<400> 21
ggccgagttc ttaatccagc gttggattca 30
<210> 22
<211> 933
<212> PRT
<213> 大肠杆菌
<400> 22
Met Gln Asn Ser Ala Leu Lys Ala Trp Leu Asp Ser Ser Tyr Leu Ser
1 5 10 15
Gly Ala Asn Gln Ser Trp Ile Glu Gln Leu Tyr Glu Asp Phe Leu Thr
20 25 30
Asp Pro Asp Ser Val Asp Ala Asn Trp Arg Ser Thr Phe Gln Gln Leu
35 40 45
Pro Gly Thr Gly Val Lys Pro Asp Gln Phe His Ser Gln Thr Arg Glu
50 55 60
Tyr Phe Arg Arg Leu Ala Lys Asp Ala Ser Arg Tyr Ser Ser Thr Ile
65 70 75 80
Ser Asp Pro Asp Thr Asn Val Lys Gln Val Lys Val Leu Gln Leu Ile
85 90 95
Asn Ala Tyr Arg Phe Arg Gly His Gln His Ala Asn Leu Asp Pro Leu
100 105 110
Gly Leu Trp Gln Gln Asp Lys Val Ala Asp Leu Asp Pro Ser Phe His
115 120 125
Asp Leu Thr Glu Ala Asp Phe Gln Glu Thr Phe Asn Val Gly Ser Phe
130 135 140
Ala Ser Gly Lys Glu Thr Met Lys Leu Gly Glu Leu Leu Glu Ala Leu
145 150 155 160
Lys Gln Thr Tyr Cys Gly Pro Ile Gly Ala Glu Tyr Met His Ile Thr
165 170 175
Ser Thr Glu Glu Lys Arg Trp Ile Gln Gln Arg Ile Glu Ser Gly Arg
180 185 190
Ala Thr Phe Asn Ser Glu Glu Lys Lys Arg Phe Leu Ser Glu Leu Thr
195 200 205
Ala Ala Glu Gly Leu Glu Arg Tyr Leu Gly Ala Lys Phe Pro Gly Ala
210 215 220
Lys Arg Phe Ser Leu Glu Gly Gly Asp Ala Leu Ile Pro Met Leu Lys
225 230 235 240
Glu Met Ile Arg His Ala Gly Asn Ser Gly Thr Arg Glu Val Val Leu
245 250 255
Gly Met Ala His Arg Gly Arg Leu Asn Val Leu Val Asn Val Leu Gly
260 265 270
Lys Lys Pro Gln Asp Leu Phe Asp Glu Phe Ala Gly Lys His Lys Glu
275 280 285
His Leu Gly Thr Gly Asp Val Lys Tyr His Met Gly Phe Ser Ser Asp
290 295 300
Phe Gln Thr Asp Gly Gly Leu Val His Leu Ala Leu Ala Phe Asn Pro
305 310 315 320
Ser His Leu Glu Ile Val Ser Pro Val Val Ile Gly Ser Val Arg Ala
325 330 335
Arg Leu Asp Arg Leu Asp Glu Pro Ser Ser Asn Lys Val Leu Pro Ile
340 345 350
Thr Ile His Gly Asp Ala Ala Val Thr Gly Gln Gly Val Val Gln Glu
355 360 365
Thr Leu Asn Met Ser Lys Ala Arg Gly Tyr Glu Val Gly Gly Thr Val
370 375 380
Arg Ile Val Ile Asn Asn Gln Val Gly Phe Thr Thr Ser Asn Pro Leu
385 390 395 400
Asp Ala Arg Ser Thr Pro Tyr Cys Thr Asp Ile Gly Lys Met Val Gln
405 410 415
Ala Pro Ile Phe His Val Asn Ala Asp Asp Pro Glu Ala Val Ala Phe
420 425 430
Val Thr Arg Leu Ala Leu Asp Phe Arg Asn Thr Phe Lys Arg Asp Val
435 440 445
Phe Ile Asp Leu Val Cys Tyr Arg Arg His Gly His Asn Glu Ala Asp
450 455 460
Glu Pro Ser Ala Thr Gln Pro Leu Met Tyr Gln Lys Ile Lys Lys His
465 470 475 480
Pro Thr Pro Arg Lys Ile Tyr Ala Asp Lys Leu Glu Gln Glu Lys Val
485 490 495
Ala Thr Leu Glu Asp Ala Thr Glu Met Val Asn Leu Tyr Arg Asp Ala
500 505 510
Leu Asp Ala Gly Asp Cys Val Val Ala Glu Trp Arg Pro Met Asn Met
515 520 525
His Ser Phe Thr Trp Ser Pro Tyr Leu Asn His Glu Trp Asp Glu Glu
530 535 540
Tyr Pro Asn Lys Val Glu Met Lys Arg Leu Gln Glu Leu Ala Lys Arg
545 550 555 560
Ile Ser Thr Val Pro Glu Ala Val Glu Met Gln Ser Arg Val Ala Lys
565 570 575
Ile Tyr Gly Asp Arg Gln Ala Met Ala Ala Gly Glu Lys Leu Phe Asp
580 585 590
Trp Gly Gly Ala Glu Asn Leu Ala Tyr Ala Thr Leu Val Asp Glu Gly
595 600 605
Ile Pro Val Arg Leu Ser Gly Glu Asp Ser Gly Arg Gly Thr Phe Phe
610 615 620
His Arg His Ala Val Ile His Asn Gln Ser Asn Gly Ser Thr Tyr Thr
625 630 635 640
Pro Leu Gln His Ile His Asn Gly Gln Gly Ala Phe Arg Val Trp Asp
645 650 655
Ser Val Leu Ser Glu Glu Ala Val Leu Ala Phe Glu Tyr Gly Tyr Ala
660 665 670
Thr Ala Glu Pro Arg Thr Leu Thr Ile Trp Glu Ala Gln Phe Gly Asp
675 680 685
Phe Ala Asn Gly Ala Gln Val Val Ile Asp Gln Phe Ile Ser Ser Gly
690 695 700
Glu Gln Lys Trp Gly Arg Met Cys Gly Leu Val Met Leu Leu Pro His
705 710 715 720
Gly Tyr Glu Gly Gln Gly Pro Glu His Ser Ser Ala Arg Leu Glu Arg
725 730 735
Tyr Leu Gln Leu Cys Ala Glu Gln Asn Met Gln Val Cys Val Pro Ser
740 745 750
Thr Pro Ala Gln Val Tyr His Met Leu Arg Arg Gln Ala Leu Arg Gly
755 760 765
Met Arg Arg Pro Leu Val Val Met Ser Pro Lys Ser Leu Leu Arg His
770 775 780
Pro Leu Ala Val Ser Ser Leu Glu Glu Leu Ala Asn Gly Thr Phe Leu
785 790 795 800
Pro Ala Ile Gly Glu Ile Asp Glu Leu Asp Pro Lys Gly Val Lys Arg
805 810 815
Val Val Met Cys Ser Gly Lys Val Tyr Tyr Asp Leu Leu Glu Gln Arg
820 825 830
Arg Lys Asn Asn Gln His Asp Val Ala Ile Val Arg Ile Glu Gln Leu
835 840 845
Tyr Pro Phe Pro His Lys Ala Met Gln Glu Val Leu Gln Gln Phe Ala
850 855 860
His Val Lys Asp Phe Val Trp Cys Gln Glu Glu Pro Leu Asn Gln Gly
865 870 875 880
Ala Trp Tyr Cys Ser Gln His His Phe Arg Glu Val Ile Pro Phe Gly
885 890 895
Ala Ser Leu Arg Tyr Ala Gly Arg Pro Ala Ser Ala Ser Pro Ala Val
900 905 910
Gly Tyr Met Ser Val His Gln Lys Gln Gln Gln Asp Leu Val Asn Asp
915 920 925
Ala Leu Asn Val Glu
930
<210> 23
<211> 2802
<212> DNA
<213> 大肠杆菌
<400> 23
atgcagaaca gcgctttgaa agcctggttg gactcttctt acctctctgg cgcaaaccag 60
agctggatag aacagctcta tgaagacttc ttaaccgatc ctgactcggt tgacgctaac 120
tggcgttcga cgttccagca gttacctggt acgggagtca aaccggatca attccactct 180
caaacgcgtg aatatttccg ccgcctggcg aaagacgctt cacgttactc ttcaacgatc 240
tccgaccctg acaccaatgt gaagcaggtt aaagtcctgc agctcattaa cgcataccgc 300
ttccgtggtc accagcatgc gaatctcgat ccgctgggac tgtggcagca agataaagtg 360
gccgatctgg atccgtcttt ccacgatctg accgaagcag acttccagga gaccttcaac 420
gtcggttcat ttgccagcgg caaagaaacc atgaaactcg gcgagctgct ggaagccctc 480
aagcaaacct actgcggccc gattggtgcc gagtatatgc acattaccag caccgaagaa 540
aaacgctgga tccaacagcg tatcgagtct ggtcgcgcga ctttcaatag cgaagagaaa 600
aaacgcttct taagcgaact gaccgccgct gaaggtcttg aacgttacct cggcgcaaaa 660
ttccctggcg caaaacgctt ctcgctggaa ggcggtgacg cgttaatccc gatgcttaaa 720
gagatgatcc gccacgctgg caacagcggc acccgcgaag tggttctcgg gatggcgcac 780
cgtggtcgtc tgaacgtgct ggtgaacgtg ctgggtaaaa aaccgcaaga cttgttcgac 840
gagttcgccg gtaaacataa agaacacctc ggcacgggtg acgtgaaata ccacatgggc 900
ttctcgtctg acttccagac cgatggcggc ctggtgcacc tggcgctggc gtttaacccg 960
tctcaccttg agattgtaag cccggtagtt atcggttctg ttcgtgcccg tctggacaga 1020
cttgatgagc cgagcagcaa caaagtgctg ccaatcacca tccacggtga cgccgcagtg 1080
accgggcagg gcgtggttca ggaaaccctg aacatgtcga aagcgcgtgg ttatgaagtt 1140
ggcggtacgg tacgtatcgt tatcaacaac caggttggtt tcaccacctc taatccgctg 1200
gatgcccgtt ctacgccgta ctgtactgat atcggtaaga tggttcaggc cccgattttc 1260
cacgttaacg cggacgatcc ggaagccgtt gcctttgtga cccgtctggc gctcgatttc 1320
cgtaacacct ttaaacgtga tgtcttcatc gacctggtgt gctaccgccg tcacggccac 1380
aacgaagccg acgagccgag cgcaacccag ccgctgatgt atcagaaaat caaaaaacat 1440
ccgacaccgc gcaaaatcta cgctgacaag ctggagcagg aaaaagtggc gacgctggaa 1500
gatgccaccg agatggttaa cctgtaccgc gatgcgctgg atgctggcga ttgcgtagtg 1560
gcagagtggc gtccgatgaa catgcactct ttcacctggt cgccgtacct caaccacgaa 1620
tgggacgaag agtacccgaa caaagttgag atgaagcgcc tgcaggagct ggcgaaacgc 1680
atcagcacgg tgccggaagc agttgaaatg cagtctcgcg ttgccaagat ttatggcgat 1740
cgccaggcga tggctgccgg tgagaaactg ttcgactggg gcggtgcgga aaacctcgct 1800
tacgccacgc tggttgatga aggcattccg gttcgcctgt cgggtgaaga ctccggtcgc 1860
ggtaccttct tccaccgcca cgcggtgatc cacaaccagt ctaacggttc cacttacacg 1920
ccgctgcaac atatccataa cgggcagggc gcgttccgtg tctgggactc cgtactgtct 1980
gaagaagcag tgctggcgtt tgaatatggt tatgccaccg cagaaccacg cactctgacc 2040
atctgggaag cgcagttcgg tgacttcgcc aacggtgcgc aggtggttat cgaccagttc 2100
atctcctctg gcgaacagaa atggggccgg atgtgtggtc tggtgatgtt gctgccgcac 2160
ggttacgaag ggcaggggcc ggagcactcc tccgcgcgtc tggaacgtta tctgcaactt 2220
tgtgctgagc aaaacatgca ggtttgcgta ccgtctaccc cggcacaggt ttaccacatg 2280
ctgcgtcgtc aggcgctgcg cgggatgcgt cgtccgctgg tcgtgatgtc gccgaaatcc 2340
ctgctgcgtc atccgctggc ggtttccagc ctcgaagaac tggcgaacgg caccttcctg 2400
ccagccatcg gtgaaatcga cgagcttgat ccgaagggcg tgaagcgcgt agtgatgtgt 2460
tctggtaagg tttattacga cctgctggaa cagcgtcgta agaacaatca acacgatgtc 2520
gccattgtgc gtatcgagca actctacccg ttcccgcata aagcgatgca ggaagtgttg 2580
cagcagtttg ctcacgtcaa ggattttgtc tggtgccagg aagagccgct caaccagggc 2640
gcatggtact gcagccagca tcatttccgt gaagtgattc cgtttggggc ttctctgcgt 2700
tatgcaggcc gcccggcctc cgcctctccg gcggtagggt atatgtccgt tcaccagaaa 2760
cagcaacaag atctggttaa tgacgcgctg aacgtcgaat aa 2802
<210> 24
<211> 405
<212> PRT
<213> 大肠杆菌
<400> 24
Met Ser Ser Val Asp Ile Leu Val Pro Asp Leu Pro Glu Ser Val Ala
1 5 10 15
Asp Ala Thr Val Ala Thr Trp His Lys Lys Pro Gly Asp Ala Val Val
20 25 30
Arg Asp Glu Val Leu Val Glu Ile Glu Thr Asp Lys Val Val Leu Glu
35 40 45
Val Pro Ala Ser Ala Asp Gly Ile Leu Asp Ala Val Leu Glu Asp Glu
50 55 60
Gly Thr Thr Val Thr Ser Arg Gln Ile Leu Gly Arg Leu Arg Glu Gly
65 70 75 80
Asn Ser Ala Gly Lys Glu Thr Ser Ala Lys Ser Glu Glu Lys Ala Ser
85 90 95
Thr Pro Ala Gln Arg Gln Gln Ala Ser Leu Glu Glu Gln Asn Asn Asp
100 105 110
Ala Leu Ser Pro Ala Ile Arg Arg Leu Leu Ala Glu His Asn Leu Asp
115 120 125
Ala Ser Ala Ile Lys Gly Thr Gly Val Gly Gly Arg Leu Thr Arg Glu
130 135 140
Asp Val Glu Lys His Leu Ala Lys Ala Pro Ala Lys Glu Ser Ala Pro
145 150 155 160
Ala Ala Ala Ala Pro Ala Ala Gln Pro Ala Leu Ala Ala Arg Ser Glu
165 170 175
Lys Arg Val Pro Met Thr Arg Leu Arg Lys Arg Val Ala Glu Arg Leu
180 185 190
Leu Glu Ala Lys Asn Ser Thr Ala Met Leu Thr Thr Phe Asn Glu Val
195 200 205
Asn Met Lys Pro Ile Met Asp Leu Arg Lys Gln Tyr Gly Glu Ala Phe
210 215 220
Glu Lys Arg His Gly Ile Arg Leu Gly Phe Met Ser Phe Tyr Val Lys
225 230 235 240
Ala Val Val Glu Ala Leu Lys Arg Tyr Pro Glu Val Asn Ala Ser Ile
245 250 255
Asp Gly Asp Asp Val Val Tyr His Asn Tyr Phe Asp Val Ser Met Ala
260 265 270
Val Ser Thr Pro Arg Gly Leu Val Thr Pro Val Leu Arg Asp Val Asp
275 280 285
Thr Leu Gly Met Ala Asp Ile Glu Lys Lys Ile Lys Glu Leu Ala Val
290 295 300
Lys Gly Arg Asp Gly Lys Leu Thr Val Glu Asp Leu Thr Gly Gly Asn
305 310 315 320
Phe Thr Ile Thr Asn Gly Gly Val Phe Gly Ser Leu Met Ser Thr Pro
325 330 335
Ile Ile Asn Pro Pro Gln Ser Ala Ile Leu Gly Met His Ala Ile Lys
340 345 350
Asp Arg Pro Met Ala Val Asn Gly Gln Val Glu Ile Leu Pro Met Met
355 360 365
Tyr Leu Ala Leu Ser Tyr Asp His Arg Leu Ile Asp Gly Arg Glu Ser
370 375 380
Val Gly Phe Leu Val Thr Ile Lys Glu Leu Leu Glu Asp Pro Thr Arg
385 390 395 400
Leu Leu Leu Asp Val
405
<210> 25
<211> 1218
<212> DNA
<213> 大肠杆菌
<400> 25
atgagtagcg tagatattct ggtccctgac ctgcctgaat ccgtagccga tgccaccgtc 60
gcaacctggc ataaaaaacc cggcgacgca gtcgtacgtg atgaagtgct ggtagaaatc 120
gaaactgaca aagtggtact ggaagtaccg gcatcagcag acggcattct ggatgcggtt 180
ctggaagatg aaggtacaac ggtaacgtct cgtcagatcc ttggtcgcct gcgtgaaggc 240
aacagcgccg gtaaagaaac cagcgccaaa tctgaagaga aagcgtccac tccggcgcaa 300
cgccagcagg cgtctctgga agagcaaaac aacgatgcgt taagcccggc gatccgtcgc 360
ctgctggctg aacacaatct cgacgccagc gccattaaag gcaccggtgt gggtggtcgt 420
ctgactcgtg aagatgtgga aaaacatctg gcgaaagccc cggcgaaaga gtctgctccg 480
gcagcggctg ctccggcggc gcaaccggct ctggctgcac gtagtgaaaa acgtgtcccg 540
atgactcgcc tgcgtaagcg tgtggcagag cgtctgctgg aagcgaaaaa ctccaccgcc 600
atgctgacca cgttcaacga agtcaacatg aagccgatta tggatctgcg taagcagtac 660
ggtgaagcgt ttgaaaaacg ccacggcatc cgtctgggct ttatgtcctt ctacgtgaaa 720
gcggtggttg aagccctgaa acgttacccg gaagtgaacg cttctatcga cggcgatgac 780
gtggtttacc acaactattt cgacgtcagc atggcggttt ctacgccgcg cggcctggtg 840
acgccggttc tgcgtgatgt cgataccctc ggcatggcag acatcgagaa gaaaatcaaa 900
gagctggcag tcaaaggccg tgacggcaag ctgaccgttg aagatctgac cggtggtaac 960
ttcaccatca ccaacggtgg tgtgttcggt tccctgatgt ctacgccgat catcaacccg 1020
ccgcagagcg caattctggg tatgcacgct atcaaagatc gtccgatggc ggtgaatggt 1080
caggttgaga tcctgccgat gatgtacctg gcgctgtcct acgatcaccg tctgatcgat 1140
ggtcgcgaat ccgtgggctt cctggtaacg atcaaagagt tgctggaaga tccgacgcgt 1200
ctgctgctgg acgtgtag 1218
<210> 26
<211> 309
<212> PRT
<213> 大肠杆菌
<400> 26
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> 27
<211> 930
<212> DNA
<213> 大肠杆菌
<400> 27
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 ggcctggtgg 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> 28
<211> 386
<212> PRT
<213> 大肠杆菌
<400> 28
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> 29
<211> 1161
<212> DNA
<213> 大肠杆菌
<400> 29
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> 30
<211> 310
<212> PRT
<213> 大肠杆菌
<400> 30
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> 31
<211> 933
<212> DNA
<213> 大肠杆菌
<400> 31
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> 32
<211> 309
<212> PRT
<213> 人工序列
<220>
<223> MetA11
<400> 32
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 Pro 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 Gly 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 Ser 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> 33
<211> 930
<212> DNA
<213> 人工序列
<220>
<223> metA11
<400> 33
atgccgattc gtgtgccgga cgagctaccc gccgtcaatt tcttgcgtga agaaaacgtc 60
tttgtgatga caacttctcg tgcgcctggt 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 ggcctggtgg ggtttaatga tgtcgcttac 360
tggccgcaga tcaaacaggt gctggagtgg tcgaaagatc acgtcacctc gacgctgtct 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> 34
<211> 15351
<212> DNA
<213> 人工序列
<220>
<223> pCC1BAC_srcO载体
<400> 34
cctggcactg ctacgatctg ataagccagc tgatcgaggg cgatacgccc gaaacgctac 60
aacgctacct gcccgcaacg ctgcagtttc ggcatcagta ataactgagt atttttgaat 120
agcgacgctg gatgccgatt ggtctgaacg gcccggcgat cgccgggcat accattggcg 180
ccggcatcgt gctggtaacg accatattag ggcgtttgag cagttgcctg gcctcacgca 240
gggagactgg attaactgac cgctgacaaa atgcctgaca atgacagtcc gttttgtgcc 300
ggcatgcaag cttgagtatt ctatagtctc acctaaatag cttggcgtaa tcatggtcat 360
agctgtttcc tgtgtgaaat tgttatccgc tcacaattcc acacaacata cgagccggaa 420
gcataaagtg taaagcctgg ggtgcctaat gagtgagcta actcacatta attgcgttgc 480
gctcactgcc cgctttccag tcgggaaacc tgtcgtgcca gctgcattaa tgaatcggcc 540
aacgcgaacc ccttgcggcc gcccgggccg tcgaccaatt ctcatgtttg acagcttatc 600
atcgaatttc tgccattcat ccgcttatta tcacttattc aggcgtagca accaggcgtt 660
taagggcacc aataactgcc ttaaaaaaat tacgccccgc cctgccactc atcgcagtac 720
tgttgtaatt cattaagcat tctgccgaca tggaagccat cacaaacggc atgatgaacc 780
tgaatcgcca gcggcatcag caccttgtcg ccttgcgtat aatatttgcc catggtgaaa 840
acgggggcga agaagttgtc catattggcc acgtttaaat caaaactggt gaaactcacc 900
cagggattgg ctgagacgaa aaacatattc tcaataaacc ctttagggaa ataggccagg 960
ttttcaccgt aacacgccac atcttgcgaa tatatgtgta gaaactgccg gaaatcgtcg 1020
tggtattcac tccagagcga tgaaaacgtt tcagtttgct catggaaaac ggtgtaacaa 1080
gggtgaacac tatcccatat caccagctca ccgtctttca ttgccatacg aaattccgga 1140
tgagcattca tcaggcgggc aagaatgtga ataaaggccg gataaaactt gtgcttattt 1200
ttctttacgg tctttaaaaa ggccgtaata tccagctgaa cggtctggtt ataggtacat 1260
tgagcaactg actgaaatgc ctcaaaatgt tctttacgat gccattggga tatatcaacg 1320
gtggtatatc cagtgatttt tttctccatt ttagcttcct tagctcctga aaatctcgat 1380
aactcaaaaa atacgcccgg tagtgatctt atttcattat ggtgaaagtt ggaacctctt 1440
acgtgccgat caacgtctca ttttcgccaa aagttggccc agggcttccc ggtatcaaca 1500
gggacaccag gatttattta ttctgcgaag tgatcttccg tcacaggtat ttattcgcga 1560
taagctcatg gagcggcgta accgtcgcac aggaaggaca gagaaagcgc ggatctggga 1620
agtgacggac agaacggtca ggacctggat tggggaggcg gttgccgccg ctgctgctga 1680
cggtgtgacg ttctctgttc cggtcacacc acatacgttc cgccattcct atgcgatgca 1740
catgctgtat gccggtatac cgctgaaagt tctgcaaagc ctgatgggac ataagtccat 1800
cagttcaacg gaagtctaca cgaaggtttt tgcgctggat gtggctgccc ggcaccgggt 1860
gcagtttgcg atgccggagt ctgatgcggt tgcgatgctg aaacaattat cctgagaata 1920
aatgccttgg cctttatatg gaaatgtgga actgagtgga tatgctgttt ttgtctgtta 1980
aacagagaag ctggctgtta tccactgaga agcgaacgaa acagtcggga aaatctccca 2040
ttatcgtaga gatccgcatt attaatctca ggagcctgtg tagcgtttat aggaagtagt 2100
gttctgtcat gatgcctgca agcggtaacg aaaacgattt gaatatgcct tcaggaacaa 2160
tagaaatctt cgtgcggtgt tacgttgaag tggagcggat tatgtcagca atggacagaa 2220
caacctaatg aacacagaac catgatgtgg tctgtccttt tacagccagt agtgctcgcc 2280
gcagtcgagc gacagggcga agccctcgag ctggttgccc tcgccgctgg gctggcggcc 2340
gtctatggcc ctgcaaacgc gccagaaacg ccgtcgaagc cgtgtgcgag acaccgcggc 2400
cggccgccgg cgttgtggat acctcgcgga aaacttggcc ctcactgaca gatgaggggc 2460
ggacgttgac acttgagggg ccgactcacc cggcgcggcg ttgacagatg aggggcaggc 2520
tcgatttcgg ccggcgacgt ggagctggcc agcctcgcaa atcggcgaaa acgcctgatt 2580
ttacgcgagt ttcccacaga tgatgtggac aagcctgggg ataagtgccc tgcggtattg 2640
acacttgagg ggcgcgacta ctgacagatg aggggcgcga tccttgacac ttgaggggca 2700
gagtgctgac agatgagggg cgcacctatt gacatttgag gggctgtcca caggcagaaa 2760
atccagcatt tgcaagggtt tccgcccgtt tttcggccac cgctaacctg tcttttaacc 2820
tgcttttaaa ccaatattta taaaccttgt ttttaaccag ggctgcgccc tgtgcgcgtg 2880
accgcgcacg ccgaaggggg gtgccccccc ttctcgaacc ctcccggtcg agtgagcgag 2940
gaagcaccag ggaacagcac ttatatattc tgcttacaca cgatgcctga aaaaacttcc 3000
cttggggtta tccacttatc cacggggata tttttataat tatttttttt atagttttta 3060
gatcttcttt tttagagcgc cttgtaggcc tttatccatg ctggttctag agaaggtgtt 3120
gtgacaaatt gccctttcag tgtgacaaat caccctcaaa tgacagtcct gtctgtgaca 3180
aattgccctt aaccctgtga caaattgccc tcagaagaag ctgttttttc acaaagttat 3240
ccctgcttat tgactctttt ttatttagtg tgacaatcta aaaacttgtc acacttcaca 3300
tggatctgtc atggcggaaa cagcggttat caatcacaag aaacgtaaaa atagcccgcg 3360
aatcgtccag tcaaacgacc tcactgaggc ggcatatagt ctctcccggg atcaaaaacg 3420
tatgctgtat ctgttcgttg accagatcag aaaatctgat ggcaccctac aggaacatga 3480
cggtatctgc gagatccatg ttgctaaata tgctgaaata ttcggattga cctctgcgga 3540
agccagtaag gatatacggc aggcattgaa gagtttcgcg gggaaggaag tggtttttta 3600
tcgccctgaa gaggatgccg gcgatgaaaa aggctatgaa tcttttcctt ggtttatcaa 3660
acgtgcgcac agtccatcca gagggcttta cagtgtacat atcaacccat atctcattcc 3720
cttctttatc gggttacaga accggtttac gcagtttcgg cttagtgaaa caaaagaaat 3780
caccaatccg tatgccatgc gtttatacga atccctgtgt cagtatcgta agccggatgg 3840
ctcaggcatc gtctctctga aaatcgactg gatcatagag cgttaccagc tgcctcaaag 3900
ttaccagcgt atgcctgact tccgccgccg cttcctgcag gtctgtgtta atgagatcaa 3960
cagcagaact ccaatgcgcc tctcatacat tgagaaaaag aaaggccgcc agacgactca 4020
tatcgtattt tccttccgcg atatcacttc catgacgaca ggatagtctg agggttatct 4080
gtcacagatt tgagggtggt tcgtcacatt tgttctgacc tactgagggt aatttgtcac 4140
agttttgctg tttccttcag cctgcatgga ttttctcata ctttttgaac tgtaattttt 4200
aaggaagcca aatttgaggg cagtttgtca cagttgattt ccttctcttt cccttcgtca 4260
tgtgacctga tatcgggggt tagttcgtca tcattgatga gggttgatta tcacagttta 4320
ttactctgaa ttggctatcc gcgtgtgtac ctctacctgg agtttttccc acggtggata 4380
tttcttcttg cgctgagcgt aagagctatc tgacagaaca gttcttcttt gcttcctcgc 4440
cagttcgctc gctatgctcg gttacacggc tgcggcgagc gctagtgata ataagtgact 4500
gaggtatgtg ctcttcttat ctccttttgt agtgttgctc ttattttaaa caactttgcg 4560
gttttttgat gactttgcga ttttgttgtt gctttgcagt aaattgcaag atttaataaa 4620
aaaacgcaaa gcaatgatta aaggatgttc agaatgaaac tcatggaaac acttaaccag 4680
tgcataaacg ctggtcatga aatgacgaag gctatcgcca ttgcacagtt taatgatgac 4740
agcccggaag cgaggaaaat aacccggcgc tggagaatag gtgaagcagc ggatttagtt 4800
ggggtttctt ctcaggctat cagagatgcc gagaaagcag ggcgactacc gcacccggat 4860
atggaaattc gaggacgggt tgagcaacgt gttggttata caattgaaca aattaatcat 4920
atgcgtgatg tgtttggtac gcgattgcga cgtgctgaag acgtatttcc accggtgatc 4980
ggggttgctg cccataaagg tggcgtttac aaaacctcag tttctgttca tcttgctcag 5040
gatctggctc tgaaggggct acgtgttttg ctcgtggaag gtaacgaccc ccagggaaca 5100
gcctcaatgt atcacggatg ggtaccagat cttcatattc atgcagaaga cactctcctg 5160
cctttctatc ttggggaaaa ggacgatgtc acttatgcaa taaagcccac ttgctggccg 5220
gggcttgaca ttattccttc ctgtctggct ctgcaccgta ttgaaactga gttaatgggc 5280
aaatttgatg aaggtaaact gcccaccgat ccacacctga tgctccgact ggccattgaa 5340
actgttgctc atgactatga tgtcatagtt attgacagcg cgcctaacct gggtatcggc 5400
acgattaatg tcgtatgtgc tgctgatgtg ctgattgttc ccacgcctgc tgagttgttt 5460
gactacacct ccgcactgca gtttttcgat atgcttcgtg atctgctcaa gaacgttgat 5520
cttaaagggt tcgagcctga tgtacgtatt ttgcttacca aatacagcaa tagcaatggc 5580
tctcagtccc cgtggatgga ggagcaaatt cgggatgcct ggggaagcat ggttctaaaa 5640
aatgttgtac gtgaaacgga tgaagttggt aaaggtcaga tccggatgag aactgttttt 5700
gaacaggcca ttgatcaacg ctcttcaact ggtgcctgga gaaatgctct ttctatttgg 5760
gaacctgtct gcaatgaaat tttcgatcgt ctgattaaac cacgctggga gattagataa 5820
tgaagcgtgc gcctgttatt ccaaaacata cgctcaatac tcaaccggtt gaagatactt 5880
cgttatcgac accagctgcc ccgatggtgg attcgttaat tgcgcgcgta ggagtaatgg 5940
ctcgcggtaa tgccattact ttgcctgtat gtggtcggga tgtgaagttt actcttgaag 6000
tgctccgggg tgatagtgtt gagaagacct ctcgggtatg gtcaggtaat gaacgtgacc 6060
aggagctgct tactgaggac gcactggatg atctcatccc ttcttttcta ctgactggtc 6120
aacagacacc ggcgttcggt cgaagagtat ctggtgtcat agaaattgcc gatgggagtc 6180
gccgtcgtaa agctgctgca cttaccgaaa gtgattatcg tgttctggtt ggcgagctgg 6240
atgatgagca gatggctgca ttatccagat tgggtaacga ttatcgccca acaagtgctt 6300
atgaacgtgg tcagcgttat gcaagccgat tgcagaatga atttgctgga aatatttctg 6360
cgctggctga tgcggaaaat atttcacgta agattattac ccgctgtatc aacaccgcca 6420
aattgcctaa atcagttgtt gctctttttt ctcaccccgg tgaactatct gcccggtcag 6480
gtgatgcact tcaaaaagcc tttacagata aagaggaatt acttaagcag caggcatcta 6540
accttcatga gcagaaaaaa gctggggtga tatttgaagc tgaagaagtt atcactcttt 6600
taacttctgt gcttaaaacg tcatctgcat caagaactag tttaagctca cgacatcagt 6660
ttgctcctgg agcgacagta ttgtataagg gcgataaaat ggtgcttaac ctggacaggt 6720
ctcgtgttcc aactgagtgt atagagaaaa ttgaggccat tcttaaggaa cttgaaaagc 6780
cagcaccctg atgcgaccac gttttagtct acgtttatct gtctttactt aatgtccttt 6840
gttacaggcc agaaagcata actggcctga atattctctc tgggcccact gttccacttg 6900
tatcgtcggt ctgataatca gactgggacc acggtcccac tcgtatcgtc ggtctgatta 6960
ttagtctggg accacggtcc cactcgtatc gtcggtctga ttattagtct gggaccacgg 7020
tcccactcgt atcgtcggtc tgataatcag actgggacca cggtcccact cgtatcgtcg 7080
gtctgattat tagtctggga ccatggtccc actcgtatcg tcggtctgat tattagtctg 7140
ggaccacggt cccactcgta tcgtcggtct gattattagt ctggaaccac ggtcccactc 7200
gtatcgtcgg tctgattatt agtctgggac cacggtccca ctcgtatcgt cggtctgatt 7260
attagtctgg gaccacgatc ccactcgtgt tgtcggtctg attatcggtc tgggaccacg 7320
gtcccacttg tattgtcgat cagactatca gcgtgagact acgattccat caatgcctgt 7380
caagggcaag tattgacatg tcgtcgtaac ctgtagaacg gagtaacctc ggtgtgcggt 7440
tgtatgcctg ctgtggattg ctgctgtgtc ctgcttatcc acaacatttt gcgcacggtt 7500
atgtggacaa aatacctggt tacccaggcc gtgccggcac gttaaccggg ctgcatccga 7560
tgcaagtgtg tcgctgtcga cgagctcgcg agctcggaca tgaggttgcc ccgtattcag 7620
tgtcgctgat ttgtattgtc tgaagttgtt tttacgttaa gttgatgcag atcaattaat 7680
acgatacctg cgtcataatt gattatttga cgtggtttga tggcctccac gcacgttgtg 7740
atatgtagat gataatcatt atcactttac gggtcctttc cggtgatccg acaggttacg 7800
gggcggcgac ctcgcgggtt ttcgctattt atgaaaattt tccggtttaa ggcgtttccg 7860
ttcttcttcg tcataactta atgtttttat ttaaaatacc ctctgaaaag aaaggaaacg 7920
acaggtgctg aaagcgagct ttttggcctc tgtcgtttcc tttctctgtt tttgtccgtg 7980
gaatgaacaa tggaagtccg agctcatcgc taataacttc gtatagcata cattatacga 8040
agttatattc gatgcggccg caaggggttc gcgtcagcgg gtgttggcgg gtgtcggggc 8100
tggcttaact atgcggcatc agagcagatt gtactgagag tgcaccatat gcggtgtgaa 8160
ataccacaca gatgcgtaag gagaaaatac cgcatcaggc gccattcgcc attcagctgc 8220
gcaactgttg ggaagggcga tcggtgcggg cctcttcgct attacgccag ctggcgaaag 8280
ggggatgtgc tgcaaggcga ttaagttggg taacgccagg gttttcccag tcacgacgtt 8340
gtaaaacgac ggccagtgaa ttgtaatacg actcactata gggcgaattc gagctcggta 8400
cccggggatc ctctagagtc gacctgcagg catgcaagct tattcaggct gcgcaactgt 8460
tgggaagggc gatcggtgcg ggcctcttcg ctattacgcc agctggcgaa agggggatgt 8520
gctgcaaggc gattaagttg ggtaacgcca gggttttccc agtcacgacg ttgtaaaacg 8580
acggccagtg aattcgagct cggtacccgg ggatcctcta gaggaattca tccgccagtt 8640
catccgggaa cggctctggc aggcagtgcc cgcagtcata catgaccgac ctgataacgt 8700
ctggccttct tcgctcagta acgttttgtg tccaggacgt cagtcttcac agctgtcaaa 8760
tcaacgctat ttcataaggc ggctcacgtt tttatgtcat gagccgcgct tttccggctg 8820
gaacgttcca cccactcggt cactgtgatt taatatgcta aaccggttta attaagttta 8880
gctttggagg gttttagcat gaatgcaaaa gtttgggttc tgggcgacgc ggtggtggac 8940
ctgctgccgg agagcgaagg gcgcctgctg cagtgccctg gaggcgcgcc ggctaacgtg 9000
gcggtagggg ttgcccgcct tggcggcaac agcggattta tcggccgcgt cggcggtgac 9060
ccgtttggcc gctacatgcg tcataccctg caacaggagc aggtcgacgt cagccatatg 9120
tatctcgacg atcagcaccg cacgtccact gtggtcgtcg accttgacga ccagggggaa 9180
cgcaccttta cctttatggt acgccccagc gcggacctgt tcctggttga agaagacctg 9240
ccacagtttg ccgccggaca gtggttgcac gtctgctcca tcgcgctcag cgccgagccc 9300
agccgtagca ctaccttcgc ggcgatggag agcatcaggt ctgccggcgg tcgggtcagc 9360
tttgacccta atattcgtcc cgatctctgg caggatcagg ctttgctgct agcctgcctc 9420
gatcgcgctt tgcacatggc caacgcggta aagctatcgg aagaggagct ggtcttcatc 9480
agcagcagta atgatttagc atacggaatc gccagcgtaa cggagcgcta tcagccagaa 9540
ttgctactgg tgacccgggg caaagcgggg gtgcttgccg cgtttcagca gaagtttacc 9600
catttcaacg cccggcctgt ggccagcgtg gacaccaccg gcgcgggaga cgcatttgtc 9660
gccggactgc tcgccagcct tgcggctaac gggatgccaa cggacatgac cgcactggaa 9720
ccgacactca cgcttgcaca gacctgcggc gccctggcca ccacagccaa aggtgcgatg 9780
accgccttgc cttatcagcg cgatctcaac cgtcagtttt aatccttaaa gccgctttgc 9840
gcggctcact ttgttgcatg catcacattt attaaaccgg tttagcatat ttgttttaag 9900
aaaaacaaag gtcgggctta gcatagcgcc taaaccggtt tagcaaaaat tataattttc 9960
catttttact tttgggatgc caacagcatg tacagaaaaa gcacacttgc gatgcttatc 10020
gctttgctaa ccagcgctgc ctcagcccat gcgcaaacgg atataagcac cattgaagcc 10080
cgactcaacg cgctggaaaa acgcctgcag gaggcagaaa acagggcgca aacggcggaa 10140
aaccgcgccg gggcggcgga gaaaaaagtt cagcaactca ccgcgcagca gcaaaaaaac 10200
cagaactcga ctcaggaagt ggctcagcgt accgccagac ttgagaaaaa agccgatgac 10260
aaaagcggat ttgagtttca cggttacgcc cgctccggcg tgataatgaa tgattccggc 10320
gccagcacca aatccggagc ctacataacg ccggcaggtg aaaccggcgg agctatcggc 10380
cgtctgggaa accaggccga tacctatgtt gaaatgaatc ttgaacataa gcagaccctg 10440
gataatgggg ccacgacccg ctttaaggtg atggtcgccg acgggcaaac ctcttataac 10500
gactggactg caagcattag cgatctgaac gttcgtcagg cctttgtcga attgggtaac 10560
ctgccgacgt tcgctgggcc atttaagggc tccaccctgt gggccgggaa acgtttcgac 10620
cgcgacaatt tcgatattca ctggattgac tctgatgtcg tgttcctcgc cggtaccggt 10680
ggtggtatct atgacgtgaa gtggaacgac ggcctgcgga gtaatttctc cctgtacggg 10740
cgtaacttcg gcgacattga tgattccagc aacagcgtgc agaactatat cctcaccatg 10800
aatcacttcg caggtccgct gcagatgatg gtcagcggtc tgcggtctgc gggcgaagga 10860
taacgacgag cgtaaagata gcaacggcaa tctggtaaaa ggcgatgcgg caaacaccgg 10920
cgtgcatgcg ctgctcggcc tgcataacga cagtttctac ggcctgcgcg acggtagcag 10980
taaaaccgct ctgctttatg gtcatggtct gggcgcagag gttaaaggta tcggatctga 11040
tggcgcactt cgtccgggag ccgacacatg gcgcattgcc agttacggca ccacgccgct 11100
cagcgaaaac tggtctgttg ccccggcaat gctggcgcaa cgcagtaaag accgctatgc 11160
cgatggcgac agctatcagt gggcaacatt caacctgcgt ctgattcagg caatcaatca 11220
gaatttcgct ctcgcctacg gaggcagcta ccagtacatg gatcttaaac ccgaaggtta 11280
taacgatcgt caggcggtga acggtagctt ctacaagctc accttcgccc cgacatttaa 11340
ggtcggcagt attggtgatt tcttcagtcg cccggagatt cgtttctata cctcctggat 11400
ggactggagc aaaaaactga ataattacgc cagcgacgac gccctgggca gtgacggttt 11460
taactcgggc ggcgaatggt ctttcggtgt gcagatggaa acctggttct gacggcttac 11520
gcctgatgac aggaatagcc gggggtcaga gcatctttgt caccccggac tcaactaaga 11580
cgcagaaaaa gcgctcccgt gaacgcggga cgacaacata aaaatgttta agccttaaga 11640
gggtactatg gattttgaac agatttcctg ctcgctgctt ccgcttcttg gaggcaaaga 11700
aaatatcgcc agcgccgcgc actgcgccac gcgcctgcgc ctggtgctgg tcgatgattc 11760
gctggccgac cagcaggcca tcggcaaagt tgaaggggtg aagggctgtt ttcgtaatgc 11820
cggacagatg cagattattt tcggcaccgg ggtggtaaat aaggtctacg ctgcctttac 11880
tcaggcggcg ggtattagcg aatccagcaa atcggaagcc gccgacattg cggcaaaaaa 11940
gctcaatccg ttccagcgca tcgcccgcct gctatcaaac atcttcgtgc cgataatccc 12000
tgccatcgtc gcctctggtc tgctgatggg cctgctggga atggtcaaaa catacggctg 12060
ggttgacccg ggcaacgcca tctacatcat gctggatatg tgcagctcgg cggcatttat 12120
cattctgccg attctgattg gctttaccgc cgcccgcgaa ttcggcggta atccttatct 12180
cggcgcgacg cttggcggca ttctgactca tccagcgctg actaacgcct ggggcgtggc 12240
cgcgggtttc cacaccatga actttttcgg cttcgaaatt gccatgatcg gctatcaggg 12300
tacggtgttc ccggtactgc tggcagtatg gtttatgagc atcgttgaga agcagttgcg 12360
tcgcgcaatc cccgatgccc tggatttgat cctgacgccg ttcctgacgg tgattatatc 12420
cggttttatc gccctgttga ttatcggccc ggccggtcgc gcactgggcg acggtatctc 12480
gtttgtcctc agcaccctga ttagccacgc cggctggctc gccgggttac tgtttggcgg 12540
tctctattca gttatcgtca ttaccggtat tcatcacagc ttccatgcgg ttgaagccgg 12600
gttgctgggc aatccctcca tcggcgtcaa cttcctgctg ccgatttggg cgatggccaa 12660
cgtcgctcag ggcggagcct gtctggcggt gtggttcaaa accaaagatg caaaaattaa 12720
agccattact ctgccctcgg cgttttccgc catgctgggc atcaccgagg cggcgatttt 12780
tggtattaac ctgcgctttg tgaagccatt tattgcggcg ctgattggtg gtgcggcggg 12840
cggcgcatgg gtggtatctg tacacgtcta catgaccgcg gtcggcttga cagcgatccc 12900
cggcatggcc atcgtgcagg ccagttcgct gttgaactac attatcggga tggttatcgc 12960
ctttggcgtc gcctttacgg tctccctggt tttgaaatac aaaacggacg ctgaataatg 13020
tctcttccat cacgactgcc tgcgattttg caggccgtaa tgcagggcca gccgcgcgcg 13080
ctggccgata gccactatcc gcgctggcac catgcgccgg tcaccgggct gatgaacgac 13140
cccaacggct ttatcgaatt tgccggacgc tatcatctgt tttatcagtg gaacccgctc 13200
gcctgcgatc atacgtttaa gtgctgggcg cactggagtt ccatcgatct gctgcactgg 13260
cagcatgagc ccattgcgct gatgccggac gaagagtatg accgtaacgg ctgctactcc 13320
ggcagcgcgg tggataacaa cggtacgctt accctgtgct ataccggcaa cgtgaagttt 13380
gccgagggag ggcgaaccgc ctggcaatgc ctggcaacgg aaaacgctga cggcaccttc 13440
cgcaaaatcg gtccggtcct gccgctgccg gagggctaca ccggccacgt gcgcgaccca 13500
aaagtctggc gacacgaaga cctgtggtac atggtgctgg gcgcgcaggg tcggcaaaag 13560
cgcggcaagg tgctgctgtt cagctctgcg gatctccatc agtggacgag tatgggtgaa 13620
atcgccggcc acggcatcaa tggcctcgac gacgtcggct atatgtggga gtgcccggat 13680
ctttttccac tcggcgacca gcatattcta acctgctgtc cgcaggggat tgcccgtgag 13740
gaagagtgct acctgaacac ctacccggca gtatggatgg cgggcgagtt tgattacgct 13800
gctggcgctt tcagacacgg cgaactgcac gaactggacg ccgggtttga gttctacgcc 13860
ccgcaaacca tgcttaccag tgatggccgt cgtctgctgg tcggctggat gggcgtgccg 13920
gagggcgaag agatgcttca gccgaccctg aacaacggct ggatccatca gatgacctgc 13980
ctgcgtgagc tggagtttat caacggtcag ctctatcagc gtccgctacg ggaactgagc 14040
gccctgcgcg gtgaagcgaa cggctggtcg gggaacgccc tgccgctggc gccgatggaa 14100
atcgatttgc aaacccgcgg gggcgatatg ttgagcctcg attttggcgg cgtattaacc 14160
cttgagtgcg atgccagcgg actccgcctg gcccgacgca gtctcgccag tgacgagatg 14220
cattatcgtt actggcgcgg aaacgtccgc tcgctgcgtg ttttcatcga ccagtcgagc 14280
gtggagattt tcataaacgg cggtgaaggg gtgatgagca gccgctactt cccggcctgc 14340
tccggtcagc taacattctc cggcatcacg ccggacgcat tctgctactg gccgctgcga 14400
acttgcatgg tagaataagc gttttgcttc aggctcatgg cgtcgtaatg aaaaccaaac 14460
gcgtaaccat taaagatata gccgaacagg ctggcgtctc caaagcgacc gccagcttgg 14520
tactgaatgg tcgtggcaag gagctgcgcg tggcgcagga aacgcgtgag cgcgtactgt 14580
cgattgcccg taagcatcac tatcagccaa gcattcatgc ccgctcgctg cgcaacaacc 14640
gcagccacac catcgggctg gtggtgccgg agatcaccaa ccacggcttt gcggtctttg 14700
cccatgagct ggagatgctg tgccgcgagg cgggcgtcca gctgttgatc tcttgtactg 14760
atgaaaaccc cggtcaggag agcgtggtgg tcaataatat gattgcccgc caggtcgacg 14820
ggatgatcgt cgcttcctgt atgcacaacg atgccgacta tctcaaattc agccaacagc 14880
tgccagtggt gctgtttgac cggtgcccca atgaaagcgc gctgccgctg gtaatgaccg 14940
attcgattac cccaacggcg gaactgattt cccgcatcgc gcctcagcat agcgatgagt 15000
tctggttttt aggcggtcag gcgcgtctgt cgccctcccg cgatcgtctg accgggttca 15060
cgcagggttt ggctcaggcg ggtattgccc tgcgcccgga atgggtgatc aacggcaatt 15120
accaccccag ctccggctat gagatgtttg ccgcactctg cgcgcgcctt gggcggccgc 15180
ctaaggcgct attcaccgcc gcctgcgggc tgctcgaagg ggttctgcgc tatatgagcc 15240
agcaccattt actcgattcc gatattcatc tgacgagctt tgacgatcac tatctttatg 15300
attcgctgtc gctgcgtatc gacactgtcc agcaggataa tcgccagctg g 15351
Claims (7)
1.一种埃希氏菌属微生物,所述埃希氏菌属微生物具有生产O-琥珀酰高丝氨酸或琥珀酸的能力,其中所述埃希氏菌属微生物的α-酮戊二酸脱氢酶的活性与所述α-酮戊二酸脱氢酶的内源活性水平相比得到增强。
2.根据权利要求1所述的埃希氏菌属微生物,其中所述α-酮戊二酸脱氢酶包含序列号22和序列号24中所列出的氨基酸序列。
3.根据权利要求1所述的埃希氏菌属微生物,其中所述埃希氏菌属微生物的高丝氨酸O-琥珀酰基转移酶的活性进一步与所述高丝氨酸O-琥珀酰基转移酶的内源活性水平相比得到增强。
4.根据权利要求1所述的埃希氏菌属微生物,其中所述埃希氏菌属微生物的胱硫醚γ-合酶和高丝氨酸激酶中至少一种的活性进一步与相应的内源活性水平相比被减弱或者消除。
5.根据权利要求1-4中任意一项所述的埃希氏菌属微生物,其中所述埃希氏菌属微生物是大肠杆菌。
6.一种生产O-琥珀酰高丝氨酸或琥珀酸的方法,其包括:
培养用于生产O-琥珀酰高丝氨酸或琥珀酸的埃希氏菌属微生物,其中所述埃希氏菌属微生物的α-酮戊二酸脱氢酶的活性与所述α-酮戊二酸脱氢酶的内源活性水平相比得到增强;并且
从所述微生物的培养液回收O-琥珀酰高丝氨酸或琥珀酸。
7.根据权利要求6所述的生产O-琥珀酰高丝氨酸或琥珀酸的方法,其中所述埃希氏菌属微生物是大肠杆菌。
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- 2015-06-03 US US15/316,476 patent/US20170247727A1/en not_active Abandoned
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- 2015-06-03 MY MYPI2016002131A patent/MY175821A/en unknown
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- 2015-06-03 JP JP2016571317A patent/JP6506314B2/ja active Active
- 2015-06-03 WO PCT/KR2015/005548 patent/WO2015186958A1/ko active Application Filing
- 2015-06-03 BR BR112016028432A patent/BR112016028432A2/pt not_active Application Discontinuation
- 2015-06-03 CN CN201580029967.5A patent/CN107109357A/zh active Pending
- 2015-06-03 RU RU2016149075A patent/RU2674891C2/ru active
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Publication number | Publication date |
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WO2015186958A1 (ko) | 2015-12-10 |
EP3153575A4 (en) | 2017-04-19 |
KR101547651B1 (ko) | 2015-08-26 |
EP3153575B1 (en) | 2019-04-24 |
RU2016149075A (ru) | 2018-07-09 |
US20170247727A1 (en) | 2017-08-31 |
JP2017516488A (ja) | 2017-06-22 |
MY175821A (en) | 2020-07-10 |
EP3153575A1 (en) | 2017-04-12 |
RU2016149075A3 (zh) | 2018-07-09 |
PL3153575T3 (pl) | 2020-02-28 |
RU2674891C2 (ru) | 2018-12-13 |
BR112016028432A2 (pt) | 2017-10-24 |
ES2734412T3 (es) | 2019-12-05 |
JP6506314B2 (ja) | 2019-04-24 |
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