CN100439492C - 宿主微生物 - Google Patents
宿主微生物 Download PDFInfo
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- CN100439492C CN100439492C CNB028111184A CN02811118A CN100439492C CN 100439492 C CN100439492 C CN 100439492C CN B028111184 A CNB028111184 A CN B028111184A CN 02811118 A CN02811118 A CN 02811118A CN 100439492 C CN100439492 C CN 100439492C
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- Prior art keywords
- gene
- glu
- ala
- asn
- val
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Abstract
一种已缺失或失活一个或几个参与芽孢形成中期到晚期的基因的微生物;一种利用微生物制得目标产物(蛋白质)的方法。当应用该微生物时,不形成芽孢,由此能够生产目标产物(蛋白质),同时降低能量消耗、副产物的生产和特定生产速度,以减少培养基的不必要的消耗。此外,生产期可以被延长,从而可以有效地制得目标产物(蛋白质)。
Description
技术领域
本发明涉及用于生产有用的蛋白质或多肽的宿主微生物,及重组微生物。
背景技术
微生物已被应用在广泛的有用物质的工业生产中。例如,利用微生物不仅可生产酒精饮料和食品,如味噌(发酵的大豆糊)和酱油(大豆调味汁),还可生产氨基酸、有机酸、与核酸相关物质、抗生素、糖类、脂类、蛋白质,及很多其它类型物质。在广泛领域中都可发现这些物质的用途,包括食品、药物、日常必需品,如清洁剂和化妆品,以及多种可通过化学方法获得产品的原料。
在通过利用微生物进行有用物质的工业生产中,一个重要问题是生产力的提高。因此,作为一种衡量方法,通过传统遗传技术,如突变技术已经进行了可生产物质的微生物的培养。特别地,归功于微生物遗传学和生物工艺学的进展,现在通过利用遗传技术,如遗传工程技术可以比以往更为有效地进行对这些可生产物质的微生物的培养,也发展了用于遗传重组的宿主微生物。例如,发展了由公认安全并优秀的微生物菌株枯草杆菌(Bacillus subtilis)Marburg No.168改良而获得的微生物菌株。
微生物含有各种不同的基因,所以它们可以使自己适应自然界里的环境改变。因此,在应用几类有限的生产培养基进行的蛋白质或类似物质的工业生产方面,微生物的物质生产力并不一定是高效的。
同样,关于特定类型微生物,已经建立了其参与早期芽孢形成的基因单一缺失或失活的菌株。不过,这些菌株在生产力上并没有足够的改进。
因此,本发明的一个目标是提供一种宿主微生物,该微生物是通过从基因组中清除对蛋白质或多肽的生产无用或有害的基因,或使这些基因失活,从而增加蛋白质或多肽产量。本发明的另一个目标是提供一种重组微生物,该重组微生物是通过把编码了蛋白质或多肽,并被连接到转录起始调节区域、翻译起始调节区域、或分泌信号区域的区域下游末端的基因整合到上述宿主微生物而获得的。本发明还有一个目标是提供一种利用重组微生物生产蛋白质或多肽的方法。
发明内容
发明人在多种编码于微生物基因组的基因中,广泛寻找对有用蛋白质或多肽的生产无用或有害的基因,并且发现与未经缺失或失活作用的微生物所获得生产力相比较,可通过如下方法提高目标蛋白质或多肽的生产力,即将参与芽孢形成的特定基因从基因组中缺失或使其失活,接着再把编码了目标蛋白质或多肽,并连接到合适的转录起始调节区域、翻译起始调节区域、或分泌信号区域的基因整合进微生物中。
相应地,本发明提供了一种已缺失或失活了参与芽孢形成中期到晚期的一个或几个基因的微生物;也提供了一种通过将编码了蛋白质或多肽,并连接到转录起始调节区域、翻译起始调节区域、或分泌信号区域的区域下游末端的基因整合进基因缺失或基因失活的微生物中,而获得的重组微生物;以及一种利用重组微生物生产蛋白质或多肽的方法。
实施发明的最佳方式
用于构建本发明微生物的母体微生物只要含有参与芽孢形成的基因就无需强加限制。优选的母体微生物是形成芽孢的微生物。母体微生物可以是野生型或突变型。特定实例包括属于杆菌属的细菌,如枯草杆菌,属于梭状芽孢杆菌属的细菌,和酵母,属于杆菌属的细菌是优选的。在这些微生物中,枯草杆菌是尤其优选的,因为其完整的基因组信息已被阐明,也建立了遗传工程和基因组工程技术,并且属于枯草杆菌的细菌具有胞外分泌蛋白质的能力。
通过利用本发明微生物产生的目标蛋白质或多肽的实例包括:在食品、药物、化妆品、清洁剂、纤维处理、医疗检测用药物等方面有用的酶;及蛋白质和多肽,如生理学活性因子。
分散存在于基因组上的几百个或更多的基因被确定参与了芽孢形成。在这些基因当中,本发明中待缺失或失活的目标基因优选的是促进芽孢形成的基因,这种基因的实例包括:编码了芽孢形成期特异性σ因子的基因,参与任何σ因子基因表达的基因,以及参与激活任何σ因子的基因。另外,通过任何σ因子转录,从而参与促进芽孢形成的基因也被包括在本发明的范畴内。在本专利说明书中,芽孢形成的早期、中期、晚期被分别限定为O-I期、II-III期、IV-V期。已知芽孢形成早期(O-I期)的胞外酶,如蛋白酶和淀粉酶的产量与对数生长阶段产生的量相比是增加的。因此,待缺失或失活的目标基因优选的是特定于芽孢形成中期到晚期表达,从而参与芽孢形成的一个或几个基因。特定地,目标基因优选的是涉及芽孢形成II、III、IV或V期,更为优选的是II或III期,特别优选的是II期的一个或几个基因。本发明发现这些基因不直接涉及目标蛋白质的生产,并且对工业生产应用的普通培养基中的微生物的生长是非必须的。
表1和表2列举了枯草杆菌的这些基因。
本说明书基于Nature 390,249-256(1997)中报导的,及JAFAN:Japan Functional Analysis Network for Bacillus subtilis(BSORF DB)(http://bacillus.genome.ad.jp/)出版的枯草杆菌基因组资料库,描述了基因的名称、位点、碱基数目及功能。
表1
基因 | 位点(kb) | 功能 |
sigE | 1,604 | II期,母细胞特异性σE因子 |
sigF | 2,443 | II期,前芽孢特异性σF因子 |
spoIISB | 1,328 | II期和其后的时期,参与芽孢形成 |
spoIIE | 71 | II期,激活前芽孢特异性σF因子 |
sigG | 1,605 | III-V期,前芽孢特异性σG因子 |
spoIVCB-spoIIIC | 2,652-2,701 | IV-V期,母细胞特异性σK因子 |
表2
基因 | 位点(kb) | 功能 |
spoIIGA | 1,604 | II期,激活母细胞特异性σE因子 |
spoIIAA | 2,444 | II期,参与激活前芽孢特异性σF因子 |
spoIVFB | 2,855 | IV-V期,激活母细胞特异性σK因子 |
spoIIR | 3,794 | II期,参与激活母细胞特异性σE因子 |
spoIIIJ | 4,213 | III-V期,参与激活前芽孢特异性σG因子 |
下列基因被认为与表1所列举基因相当:与表1或表2所列枯草杆菌基因功能相同的基因;来源自其它微生物的基因,优选的是来自杆菌属细菌,并且表现出与表1所列基因中的一个基因有70%或更多同源,优选的是80%或更多,更为优选的是90%或更多同源性。这些基因被包括在本发明的待缺失或失活的基因当中。氨基酸序列之间的同源性是通过Lipman-Pearson方法计算的(Science,227,1435(1985))。
当一个或几个选自上述基因的基因被缺失或失活时,微生物的芽孢形成所需的化学能消耗便降低了,蛋白质或多肽的生成期被延长,或获得其它益处,使得提高了蛋白质或多肽的生产力。
对被缺失或失活的基因数目无需强加限制,只要有至少一个基因被缺失或失活即可。数目可以是3个或更多,或5个或更多。优选的是2个或3个,尤其优选的是2个。
为构建本发明的微生物,还可缺失或失活除上述基因外的一个或几个基因。通过这样组合,便可预期在改进生产力方面获得更大的效果。
基因的缺失或失活可通过已知方法进行。实例包括顺序地缺失或失活目标基因的方法,以及通过缺失或突变使一个或几个任意DNA片段失活,并通过适当技术根据蛋白质生产力分析和评估相应基因的方法。
通过例如同源重组方法缺失或失活目标基因。特定的是通过利用合适的质粒载体进行克隆而获得含目标基因的DNA片段。其它方法中,是采用如下方式使所获DNA片段突变的,即通过常规基因工程技术使基因的全部区域或目标基因的部分区域缺失,而保留与目标基因的相应末端相连的DNA片段;通过碱基替代、移码突变等方法引起结构基因中的无意义突变;或者通过克隆化或PCR分离目标基因片段,再将DNA片段插入分离的目标基因片段中的方法。随后将突变DNA片段导入母体微生物,从而在与目标基因相应末端邻近的两个区域,与母体微生物的基因组同源重组。这样,基因组上的目标基因便可被已缺失或失活目标基因的DNA片段所替代。
当以枯草杆菌细菌为母体微生物以获得本发明微生物时,通过同源重组缺失或失活目标基因的一些方法已有报导(如Mol.Gen.Genet.,223,268(1990))。本发明的宿主微生物可通过重复这种方法获得。
也可通过与上述类似的方式,利用片段进行克隆和同源重组,从母体微生物获得一个或几个任意DNA片段;或可选地通过γ射线照射母体微生物,从而使一个或几个任意DNA片段缺失或失活。
本发明的重组微生物可通过如下方法获得,即将编码目标蛋白质或多肽的基因(下文见“目标蛋白质或多肽基因”)整合到已缺失或失活一个或几个参与芽孢形成中期到晚期的基因的微生物中。
对目标蛋白质或多肽基因无需强加限制。这种基因的实例包括:工业用酶,如用于生产清洁剂、食品、纤维、饲料、化学制品的酶,医用酶,诊断用酶,以及生理学活性肽。工业用酶基于其功能可分类为:氧化还原酶、转移酶、水解酶、裂解酶、异构酶、连接酶/合成酶及其它。目标蛋白质或多肽基因优选的实例包括:编码水解酶,诸如纤维素酶、α-淀粉酶或蛋白酶的基因。特定实例包括属于多糖水解酶分类中的家族5的纤维素酶(Biochem.J.,280,309(1991))。其中,例证了来源自微生物的纤维素酶,特别是来源自杆菌属细菌的纤维素酶。更多特定实例包括:来源自杆菌属细菌,并具有SEQ ID NO:1或2序列的碱性纤维素酶;及具有特定序列的纤维素酶,该序列具有与SEQ ID NO:1或2序列70%或更多,优选的80%或更多,更为优选的90%或更多的同源性。氨基酸序列之间的同源性是通过Lipman-Pearson方法测定的(Science,227,1435(1985))。α-淀粉酶的实例包括来源自微生物的α-淀粉酶,特别优选的是来源自杆菌属细菌的液化型α-淀粉酶。蛋白酶的实例包括来源自微生物的丝氨酸蛋白酶和金属蛋白酶,特别是来源自杆菌属细菌的蛋白酶。
目标蛋白质或多肽基因在其上游末端被理想地连接到参与基因转录或翻译、或基因产物分泌的调节区域,即含有启动子和转录起始点的转录起始调节区域、含有核糖体结合位点和起始密码子的翻译起始区域、或合适形式的分泌信号肽区域。例如,目标蛋白质或多肽基因被理想地连接到纤维素酶基因包含的上述调节区域中,该纤维素酶基因来源自Japanese Patent Application Laid-Open(kokai)No.2000-210081或Hei 4-190793所描述的杆菌属细菌,以及被连接到纤维素基因上游末端邻近区域包含的上述调节区域中并具有1kb或小于1kb,优选的0.6kb或小于0.6kb的长度。特定的在其它情况当中,目标蛋白质或多肽基因被理想地连接到SEQ ID NO:1或2序列,或与SEQ ID NO:1或2序列一定程度上同源、具有与上述序列相似调节功能的碱基序列中。
本发明的重组微生物可通过如下方法获得,使含有目标蛋白质或多肽基因的DNA片段与合适的质粒载体组合,并通过常规转化方法将重组质粒整合进宿主微生物。可选地,本发明的重组微生物可通过另一方法获得,即利用被连接到宿主微生物基因的合适同源区域中的DNA片段,将其直接整合进宿主微生物基因。
利用本发明的重组微生物进行目标蛋白质或多肽的生产可通过如下方法完成:将重组微生物接种至含有可同化碳源、氮源和其它基本成分的培养基中,通过常规方法培养重组微生物,并于培养完成后,收集并纯化目标蛋白质或多肽。
如上所述,可生成已缺失或失活芽孢形成相关基因的目标宿主微生物,并且通过利用宿主微生物可生产目标重组微生物。另外,利用重组微生物,可有效生产有用的蛋白质或多肽。通过利用枯草杆菌制得α-淀粉酶或纤维素酶的实例情况将于下文特别描述。
例如,当枯草杆菌属细菌编码RNA聚合酶亚基σF因子的sigF基因(768bp)被缺失时,可应用下列操作步骤,该RNA聚合酶亚基σF因子在芽孢形成II期或随后时期出现的前芽孢中表达。
第一步,从枯草杆菌菌株的宿主微生物中提取基因组基因。以基因组基因为模板,通过SOE(重叠延伸拼接法)-PCR(Gene,77,61(1989))或其它方法将标记基因,如抗氯霉素基因插入待连接片段,即位于sigF基因起始密码子上游的DNA片段和位于sigF基因终止密码子下游的DNA片段之间,使二者连接起来。
下一步,通过适当方法利用上一步所获DNA片段转化宿主枯草杆菌属细菌,并基于氯霉素抗性或其它特性,分离转化株,促成sigF基因的上游和下游区域的同源重组,从而获得相应的转化株,该转化株中,位于基因组上的sigF基因被诸如抗氯霉素基因等标记基因所替代。
其后,将含有编码α-淀粉酶或纤维素酶的基因的质粒导入上一步获得的转化株中,并以枯草杆菌的原始细胞系为对照。在合适的条件下,例如在营养培养基中进行震荡,培养已获得的重组体。检测培养溶液上清的α-淀粉酶活性或纤维素酶活性,并将其生产力与枯草杆菌原始细胞系的生产力相比较,从而确认通过缺失sigF基因可获得增加量的目标产物。当分离和纯化培养液时,可获得α-淀粉酶和纤维素酶。
实施例
实施例1
从枯草杆菌168中提取基因组DNA。以提取的基因为模板,使基因组上在sigF基因(碱基号2442658←2443425)上游末端与其邻近的一个长度为1.5-kb的DNA片段(A),和在sigF基因下游末端与其邻近的一个长度为1.5-kb的DNA片段(B)倍增。以质粒pC194为模板,单独倍增含有抗氯霉素基因的0.9-kb DNA片段(C)。通过SOE-PCR按顺序连续连接片段(A)、(C)、(B),从而制得长度为3.9-kb的DNA片段。通过适当方法利用获得的DNA片段转化枯草杆菌168。在含有氯霉素的LB琼脂培养基上培养枯草杆菌168,分离菌落作为转化株。通过PCR和测序证实:该转化株具有的基因组中,含sigF基因(2442632-2443318)的区域缺失,并被抗氯霉素基因替代。基因组的下列区域中的每一个都被分别缺失,并以与上述相似的方式被抗氯霉素基因替代:含sigE基因的区域(1604136-1604976);含spoIISB基因的替代部分(1347913←1348083)的区域(1347781-1348081);含spoIIE基因的替代部分(70536→73019)的区域(70537-73018);含sigG基因的替代部分(1605025→1605807)的区域(1605083-1605877);含spoIVCB基因(2652262→2652732)的区域(2652156-2652723);或含从spoIVCB基因到spoIIIC基因的区域(2652262→2701023)的区域(2652156-2701031),从而制得参与芽孢形成的基因缺失的微生物。
将来源自杆菌sp.KSM-S237(Japanese Patent Application Laid-Open(kokai)No.2000-210081)的碱性纤维素酶基因的一个DNA片段(3.1kb)导入穿梭载体pHY300PLK的限制酶BamHI切割点,从而制得重组体质粒pHY-S237。通过原生质体方法将质粒整合进实施例1制得的每个基因缺失微生物内,并以枯草杆菌168作为对照。在LB培养基(10ml)中,37C,震荡条件下培养获得的微生物过夜。将相应的培养液(0.05ml)接种至2×L-麦芽糖培养基中(50ml;2%蛋白胨,1%酵母提取物,1%NaCl,7.5%麦芽糖,7.5-ppm 4-5水合硫酸锰,15-ppm四环素),然后在30℃震荡条件下培养3天。培养完成后,通过离心从培养液中除去细胞,并检测上清液中的碱性纤维素酶活性,以测定培养期间细胞外分泌的碱性纤维素酶的数量。结果如表3所示,发现与对照微生物枯草杆菌168(野生型)相比,所有缺失了参与芽孢形成的基因的微生物都分泌了增加量的碱性纤维素酶。
表3
缺失基因 | 基因位点(kb) | 分泌的碱性纤维素酶数量(相对值) |
sigE | 1,604 | 217 |
sigF | 2,443 | 212 |
spoIISB | 1,328 | 140 |
spoIIE | 71 | 216 |
sigG | 1,605 | 163 |
spoIVCB-spoIIIC | 2,652-2,701 | 141 |
spoIVCB | 2,652 | 141 |
无(野生型) | - | 100 |
工业适用性
当应用本发明微生物时不形成芽孢。因此,本发明能够生产目标蛋白质或目标多肽,同时降低能量损失、副产物的生产和特定生产速度,以大量减少培养基不必要的消耗。此外,蛋白质或多肽的生产期可以被延长,由此可以有效地制得目标产物。
序列表
<110)>花王株式会社
<120>宿主微生物
<130>KS-660
<140>
<141>
<150>JP P2001-160520
<151>2001-05-29
<160>2
<170>PatentIn Ver.2.1
<210>1
<211>3150
<212>DNA
<213>杆菌sp.KSM-S237
<220>
<221>CDS
<222>(573)..(3044)
<220>
<221>sig_肽
<222>(573)..(659)
<220>
<221>mat_肽
<222>(660)..(3044)
<400>1
gatttgccga tgcaacaggc ttatatttag aggaaattto tttttaaatt gaatacggaa 60
taaaatcagg taaacaggtc ctgattttat ttttttgagt tttttagaga actgaagatt 120
gaaataaaag tagaagacaa aggacataag aaaattgcat tagttttaat tatagaaaac 180
gcctttttat aattatttat acctagaacg aaaatactgt ttcgaaagcg gtttactata 240
aaaccttata ttccggctct tttttaaaac agggggtaaa aattcactct agtattctaa 300
tttcaacatg ctataataaa tttgtaagac gcaatatgca tctctttttt tacgatatat 360
gtaagcggtt aaccttgtgc tatatgccga tttaggaagg ggggtagatt gagtcaagta 420
gtaataatat agataactta taagttgttg agaagcagga gagcatctgg gttactcaca 480
agttttttta aaactttaac gaaagcactt tcggtaatgc ttatgaattt agctatttga 540
ttcaattact ttaaaaatat ttaggaggta at atg atg tta aga aag aaa aca 593
Met Met Leu Arg Lys Lys Thr
-25
aag cag ttg att tct tcc att ctt att tta gtt tta ctt cta tct tta 641
Lys Gln Leu Ile Ser Ser Ile Leu Ile Leu Val Leu Leu Leu Ser Leu
-20 -15 -10
ttt ccg gca gct ctt gca gca gaa gga aac act cgt gaa gac aat ttt 689
Phe Pro Ala Ala Leu Ala Ala Glu Gly Asn Thr Arg Glu Asp Asn Phe
-5 -1 1 5 10
aaa cat tta tta ggt aat gac aat gtt aaa cgc cct tct gag gct ggc 737
Lys His Leu Leu Gly Asn Asp Asn Val Lys Arg Pro Ser Glu Ala Gly
15 20 25
gca tta caa tta caa gaa gtc gat gga caa atg aca tta gta gat caa 785
Ala Leu Gln Leu Gln Glu Val Asp Gly Gln Met Thr Leu Val Asp Gln
30 35 40
cat gga gaa aaa att caa tta cgt gga atg agt aca cac gga tta cag 833
His Gly Glu Lys Ile Gln Leu Arg Gly Met Ser Thr His Gly Leu Gln
45 50 55
tgg ttt cct gag atc ttg aat gat aac gca tac aaa gct ctt tct aac 881
Trp Phe Pro Glu Ile Leu Asn Asp Asn Ala Tyr Lys Ala Leu Ser Asn
60 65 70
gat tgg gat tcc aat atg att cgt ctt gct atg tat gta ggt gaa aat 929
Asp Trp Asp Ser Asn Met Ile Arg Leu Ala Met Tyr Val Gly Glu Asn
75 80 85 90
ggg tac gct aca aac cct gag tta atc aaa caa aga gtg att gat gga 977
Gly Tyr Ala Thr Asn Pro Glu Leu Ile Lys Gln Arg Val Ile Asp Gly
95 100 105
att gag tta gcg att gaa aat gac atg tat gtt att gtt gac tgg cat 1025
Ile Glu Leu Ala Ile Glu Asn Asp Met Tyr Val Ile Val Asp Trp His
110 115 120
gtt cat gcg cca ggt gat cct aga gat cct gtt tat gca ggt gct aaa 1073
Val His Ala Pro Gly Asp Pro Arg Asp Pro Val Tyr Ala Gly Ala Lys
125 130 135
gat ttc ttt aga gaa att gca gct tta tac cct aat aat cca cac att 1121
Asp Phe Phe Arg Glu Ile Ala Ala Leu Tyr Pro Asn Asn Pro His Ile
140 145 150
att tat gag tta gcg aat gag ccg agt agt aat aat aat ggt gga gca 1169
Ile Tyr Glu Leu Ala Asn Glu Pro Ser Ser Asn Asn Asn Gly Gly Ala
155 160 165 170
ggg att ccg aat aac gaa gaa ggt tgg aaa gcg gta aaa gaa tat gct 1217
Gly Ile Pro Asn Asn Glu Glu Gly Trp Lys Ala Val Lys Glu Tyr Ala
175 180 185
gat cca att gta gaa atg tta cgt aaa agc ggt aat gca gat gac aac 1265
Asp Pro Ile Val Glu Met Leu Arg Lys Ser Gly Asn Ala Asp Asp Asn
190 195 200
att atc att gtt ggt agt cca aac tgg agt cag cgt ccg gac tta gca 1313
Ile Ile Ile Val Gly Ser Pro Asn Trp Ser Gln Arg Pro Asp Leu Ala
205 210 215
gct gat aat cca att gat gat cac cat aca atg tat act gtt cac ttc 1361
Ala Asp Asn Pro Ile Asp Asp His His Thr Met Tyr Thr Val His Phe
220 225 230
tac act ggt tca cat gct gct tca act gaa agc tat ccg tct gaa act 1409
Tyr Thr Gly Ser His Ala Ala Ser Thr Glu Ser Tyr Pro Ser Glu Thr
235 240 245 250
cct aac tct gaa aga gga aac gta atg agt aac act cgt tat gcg tta 1457
Pro Asn Ser Glu Arg Gly Asn Val Met Ser Asn Thr Arg Tyr Ala Leu
255 260 265
gaa aac gga gta gcg gta ttt gca aca gag tgg gga acg agt caa gct 1505
Gln Asn Gly Val Ala Val Phe Ala Thr Glu Trp Gly Thr Ser Gln Ala
270 275 280
agt gga gac ggt ggt cct tac ttt gat gaa gca gat gta tgg att gaa 1553
Ser Gly Asp Gly Gly Pro Tyr Phe Asp Glu Ala Asp Val Trp Ile Glu
285 290 295
ttt tta aat gaa aac aac att agc tgg gct aac tgg tct tta acg aat 1601
Phe Leu Asn Glu Asn Asn Ile Ser Trp Ala Asn Trp Ser Leu Thr Asn
300 305 310
aaa aat gaa gta tct ggt gca ttt aca cca ttc gag tta ggt aag tct 1649
Lys Asn Glu Val Ser Gly Ala Phe Thr Pro Phe Glu Leu Gly Lys Ser
315 320 325 330
aac gca acc aat ctt gac cca ggt cca gat cat gtg tgg gca cca gaa 1697
Asn Ala Thr Asn Leu Asp Pro Gly Pro Asp His Val Trp Ala Pro Glu
335 340 345
gaa tta agt ctt tct gga gaa tat gta cgt gct cgt att aaa ggt gtg 1745
Glu Leu Ser Leu Ser Gly Glu Tyr Val Arg Ala Arg Ile Lys Gly Val
350 355 360
aac tat gag cca atc gac cgt aca aaa tac acg aaa gta ctt tgg gac 1793
Asn Tyr Glu Pro Ile Aap Arg Thr Lys Tyr Thr Lys Val Leu Trp Asp
365 370 375
ttt aat gat gga acg aag caa gga ttt gga gtg aat tcg gat tct cca 1841
Phe Asn Asp Gly Thr Lys Gln Gly Phe Gly Val Asn Ser Asp Ser Pro
380 385 390
aat aaa gaa ctt att gca gtt gat aat gaa aac aac act ttg aaa gtt 1889
Asn Lys Glu Leu Ile Ala Val Asp Asn Glu Asn Asn Thr Leu Lys Val
395 400 405 410
tcg gga tta gat gta agt aac gat gtt tca gat ggc aac ttc tgg gct 1937
Scr Gly Leu Asp Val Ser Asn Asp Val Ser Asp Gly Asn Phe Trp Ala
415 420 425
aat gct cgt ctt tct gcc aac ggt tgg gga aaa agt gtt gat att tta 1985
Asn Ala Arg Leu Ser Ala Asn Gly Trp Gly Lys Ser Val Asp Ile Leu
430 435 440
aat gct gag aag ctt aca atg gat gtt att gtt gat gaa cca acg acg 2033
Gly Ala Glu Lys Leu Thr Met Asp Val Ile Val Asp Glu Pro Thr Thr
445 450 455
gta gct att gcg gcg att cca caa agt agt aaa agt gga tgg gca aat 2081
Val Ala Ile Ala Ala Ile Pro Gln Ser Ser Lys Ser Gly Trp Ala Asn
460 465 470
cca gag cgt gct gtt cga gtg aac gcg gaa gat ttt gtc cag caa acg 2129
Pro Glu Arg Ala Val Arg Val Asn Ala Glu Asp Phe Val Gln Gln Thr
475 480 485 490
gac ggt aag tat aaa gct gga tta aca att aca gga gaa gat gct cct 2177
Asp Gly Lys Tyr Lys Ala Gly Leu Thr Ile Thr Gly Glu Asp Ala Pro
495 500 505
aac cta aaa aat atc gct ttt cat gaa gaa gat aac aat atg aac aac 2225
Asn Leu Lys Asn Ile Ala Phe His Glu Glu Asp Asn Asn Met Asn Asn
510 515 520
atc att ctg ttc gtg gga act gat gca gct gac gtt att tac tta gat 2273
Ile Ile Leu Phe Val Gly Thr Asp Ala Ala Asp Val Ile Tyr Leu Asp
525 530 535
aac att aaa gta att gga aca gaa gtt gaa att cca gtt gtt cat gat 2321
Asn Ile Lys Val Ile Gly Thr Glu Val Glu Ile Pro Val Val His Asp
540 545 550
cca aaa gga gaa gct gtt ctt cct tct gtt ttt gaa gac ggt aca cgt 2369
Pro Lys Gly Glu Ala Val Leu Pro Ser Val Phe Glu Asp Gly Thr Arg
555 560 565 570
caa ggt tgg gac tgg gct gga gag tct ggt gtg aaa aca gct tta aca 2417
Gln Gly Trp Asp Trp Ala Gly Glu Ser Gly Val Lys Thr Ala Leu Thr
575 580 585
att gaa gaa gca aac ggt tct aac gcg tta tca tgg gaa ttt gga tat 2465
Ile Glu Glu Ala Asn Gly Ser Asn Ala Leu Ser Trp Glu Phe Gly Tyr
590 595 600
cca gaa gta aaa cct agt gat aac tgg gca aca gct cca cgt tta gat 2513
Pro Glu Val Lys Pro Ser Asp Asn Trp Ala Thr Ala Pro Arg Leu Asp
605 610 615
ttc tgg aaa tct gac ttg gtt cgc ggt gag aat gat tat gta gct ttt 2561
Phc Trp Lys Ser Asp Leu Val Arg Gly Glu Asn Asp Tyr Val Ala Phe
620 625 630
gat ttc tat cta gat cca gtt cgt gca aca gaa ggc gca atg aat atc 2609
Asp Phe Tyr Leu Asp Pro Val Arg Ala Thr Glu Gly Ala Met Asn Ile
635 640 645 650
aat tta gta ttc cag cca cct act aac ggg tat tgg gta caa gca cca 2657
Asn Leu Val Phe Gln Pro Pro Thr Asn Gly Tyr Trp Val Gln Ala Pro
655 660 665
aaa acg tat acg att aac ttt gat gaa tta gag gaa gcg aat caa gta 2705
Lys Thr Tyr Thr Ile Asn Phe Asp Glu Leu Glu Glu Ala Asn Gln Val
670 675 680
aat ggt tta tat cac tat gaa gtg aaa att aac gta aga gat att aca 2753
Asn Gly Leu Tyr His Tyr Glu Val Lys Ile Asn Val Arg Asp Ile Thr
685 690 695
aac att caa gat gac acg tta cta cgt aac atg atg atc att ttt gca 2801
Asn Ile Gln Asp Asp Thr Leu Leu Arg Asn Met Met Ile Ile Phe Ala
700 705 710
gat gta gaa agt gac ttt gca ggg aga gtc ttt gta gat aat gtt cgt 2849
Asp Val Glu Ser Asp Phe Ala Gly Arg Val Phe Val Asp Asn Val Arg
715 720 725 730
ttt gag ggg gct gct act act gag ccg gtt gaa cca gag cca gtt gat 2897
Phe Glu Gly Ala Ala Thr Thr Glu Pro Val Glu Pro Glu Pro Val Asp
735 740 745
cct ggc gaa gag acg cca cct gtc gat gag aag gaa gcg aaa aaa gaa 2945
Pro Gly Glu Glu Thr Pro Pro Val Asp Glu Lys Glu Ala Lys Lys Glu
750 755 760
caa aaa gaa gca gag aaa gaa gag aaa gaa gca gta aaa gaa gaa aag 2993
Gln Lys Glu Ala Glu Lys Glu Glu Lys Glu Ala Val Lys Glu Glu Lys
765 770 775
aaa gaa gct aaa gaa gaa aag aaa gca gtc aaa aat gag gct aag aaa 3041
Lys Glu Ala Lys Glu Glu Lys Lys Ala Val Lys Asn Glu Ala Lys Lys
780 785 790
aaa taatctatta aactagttat agggttatct aaaggtctga tgtagatctt 3094
Lys
795
ttagataacc tttttcttgc ataactggac acagagttgt tattaaagaa agtaag 3150
<210>2
<211>3332
<212>DNA
<213>杆菌sp.KSM-64
<220>
<221>CDS
<222>(610)..(3075)
<220>
<221>sig_肽
<222>(610)..(696)
<220>
<221>mat_肽
<222>(697)..(3075)
<400>2
agtacttacc attttagagt caaaagatag aagccaagca ggatttgccg atgcaaccgg 60
cttatattta gagggaattt ctttttaaat tgaatacgga ataaaatcag gtaaacaggt 120
cctgatttta tttttttgaa tttttttgag aactaaagat tgaaatagaa gtagaagaca 180
acggacataa gaaaattgta ttagttttaa ttatagaaaa cgcttttcta taattattta 240
tacctagaac gaaaatactg tttcgaaagc ggtttactat aaaaccttat attccggctc 300
tttttttaas cagggggtga aaattcactc tagtattcta atttcaacat gctataataa 360
atttgtaaga cgcaatatac atcttttttt tatgatattt gtaagcggtt aaccttgtgc 420
tatatgccga tttaggaagg gggtagattg agtcaagtag tcataattta gataacttat 480
aagttgttga gaagcaggag agaatctggg ttactcacaa gttttttaaa acattatcga 540
aagcactttc ggttatgctt atgaatttag ctatttgatt caattacttt aataatttta 600
ggaggtaat atg atg tta aga aag aaa aca aag cag ttg att tct tcc att 651
Met Met Leu Arg Lys Lys Thr Lys Gln Leu Ile Ser Ser Ile
-25 -20
ctt att tta gtt tta ctt cta tct tta ttt ccg aca gct ctt gca gca 699
Lcu Ile Leu Val Leu Leu Leu Ser Leu Phe Pro Thr Ala Leu Ala Ala
-15 -10 -5 -1 1
gaa gga aac act cgt gaa gac aat ttt aaa cat tta tta ggt aat gac 747
Glu Gly Asn Thr Arg Glu Asp Asn Phe Lys His Leu Leu Gly Asn Asp
5 10 15
aat gt t aaa cgc cct tct gag gct ggc gca tta caa tta caa gaa gtc 795
Asn Val Lys Arg Pro Ser Glu Ala Gly Ala Leu Gln Leu Gln Glu Val
20 25 30
gat gga caa atg aca tta gta gat caa cat gga gaa aaa att caa tta 843
Asp Gly Gln Met Thr Leu Val Asp Gln His Gly Glu Lys Ile Gln Leu
35 40 45
cgt gga a tg agt aca cac gga tta caa tgg ttt cct gag atc ttg aat 891
Arg Gly Met Ser Thr His Gly Leu Gln Trp Phe Pro Glu Ile Leu Asn
50 55 60 65
gat aac gca tac aaa gct ctt gct aac gat tgg gaa tca aat atg att 939
Asp Asn Ala Tyr Lys Ala Leu Ala Asn Asp Trp Glu Ser Asn Met Ile
70 75 80
cgt cta gct atg tat gtc ggt gaa aat ggc tat gct tca aat cca gag 987
Arg Leu Ala Met Tyr Val Gly Glu Asn Gly Tyr Ala Ser Asn Pro Glu
85 90 95
tta att aaa agc aga gtc att aaa gga ata gat ctt gct att gaa aat 1035
Lcu Ile Lys Ser Arg Val Ile Lys Gly Ile Asp Leu Ala Ile Glu Asn
100 105 110
gac atg tat gtc atc gtt gat tgg cat gta cat gca cct ggt gat cct 1083
Asp Mot Tyr Val Ilc Val Asp Trp His Val His Ala Pro Gly Asp Pro
115 120 125
aga gat ccc gtt tac gct gga gca gaa gat ttc ttt aga gat att gca 1131
Arg Asp Pro Val Tyr Ala Gly Ala Glu Asp Phe Phe Arg Asp Ile Ala
130 135 140 145
gca tta tat cct aac aat cca cac att att tat gag tta gcg aat gag 1179
Ala Leu Tyr Pro Asn Asn Pro His Ile Ile Tyr Glu Leu Ala Asn Glu
150 155 160
cca agt agt aac aat aat ggt gga gct ggg att cca aat aat gaa gaa 1227
Pro Ser Ser Asn Asn Asn Gly Gly Ala Gly Ile Pro Asn Asn Glu Glu
165 170 175
ggt tgg aat gcg gta aaa gaa tac gct gat cca att gta gaa atg tta 1275
Gly Trp Asn Ala Val Lys Glu Tyr Ala Asp Pro Ile Val Glu Met Leu
180 185 190
cgt gat agc ggg aac gca gat gac aat att atc att gtg ggt agt cca 1323
Arg Asp Ser Gly Asn Ala Asp Asp Asn Ile Ile Ile Val Gly Ser Pro
195 200 205
aac tgg agt cag cgt cct gac tta gca gct gat aat cca att gat gat 1371
Asn Trp Ser Gln Arg Pro Asp Leu Ala Ala Asp Asn Pro Ile Asp Asp
210 215 220 225
cac cat aca atg tat act gtt cac ttc tac act ggt tca cat gct gct 1419
His His Thr Met Tyr Thr Val His Phe Tyr Thr Gly Ser His Ala Ala
230 235 240
tca act gaa agc tat ccg cct gaa act cct aac tct gaa aga gga aac 1467
Ser Thr Glu Ser Tyr Pro Pro Glu Thr Pro Asn Ser Glu Arg Gly Asn
245 250 255
gta atg agt aac act cgt tat gcg tta gaa aac gga gta gca gta ttt 1515
Val Met Ser Asn Thr Arg Tyr Ala Leu Glu Asn Gly Val Ala Val Phe
260 265 270
gca aca gag tgg gga act agc caa gca aat gga gat ggt ggt cct tac 1563
Ala Thr Glu Trp Gly Thr Ser Gln Ala Asn Gly Asp Gly Gly Pro Tyr
275 280 285
ttt gat gaa gca gat gta tgg att gag ttt tta aat gaa aac aac att 1611
Phe Asp Glu Ala Asp Val Trp Ile Glu Phe Leu Asn Glu Asn Asn Ile
290 295 300 305
agc tgg gct aac tgg tct tta acg aat aaa aat gaa gta tct ggt gca 1659
Ser Trp Ala Asn Trp Ser Leu Thr Asn Lys Asn Glu Val Ser Gly Ala
310 315 320
ttt aca cca ttc gag tta ggt aag tct aac gca aca agt ctt gac cca 1707
Phe Thr Pro Phe Glu Leu Gly Lys Ser Asn Ala Thr Ser Leu Asp Pro
325 330 335
ggg cca gac caa gta tgg gta cca gaa gag tta agt ctt tct gga gaa 1755
Gly Pro Asp Gln Val Trp Val Pro Glu Glu Leu Ser Leu Ser Gly Glu
340 345 350
tat gta cgt gct cgt att aaa ggt gtg aac tat gag cca atc gac cgt 1803
Tyr Val Arg Ala Arg Ile Lys Gly Val Asn Tyr Glu Pro Ile Asp Arg
355 360 365
aca aaa tac acg aaa gta ctt tgg gac ttt aat gat gga acg aag caa 1851
Thr Lys Tyr Thr Lys Val Leu Trp Asp Phe Asn Asp Gly Thr Lys Gln
370 375 380 385
gga ttt gga gtg aat gga gat tct cca gtt gaa gat gta gtt att gag 1899
Gly Phc Gly Val Asn Gly Asp Ser Pro Val Glu Asp Val Val Ile Glu
390 395 400
aat gaa gcg ggc gct tta aaa ctt tca gga tta gat gca agt aat gat 1947
Asn Glu Ala Gly Ala Leu Lys Leu Ser Gly Leu Asp Ala Ser Asn Asp
405 410 415
gtt tct gaa ggt aat tac tgg gct aat gct cgt ctt tct gcc gac ggt 1995
Val Ser Glu Gly Asn Tyr Trp Ala Asn Ala Arg Leu Ser Ala Asp Gly
420 425 430
tgg gga aaa agt gtt gat att tta ggt gct gaa aaa ctt act atg gat 2043
Trp Gly Lys Ser Val Asp Ile Leu Gly Ala Glu Lys Leu Thr Met Asp
435 440 445
gtg att gtt gat gag ccg acc acg gta tca att gct gca att cca caa 2091
Val Ile Val Asp Glu Pro Thr Thr Val Ser Ile Ala Ala Ile Pro Gln
450 455 460 465
ggg cca tca gcc aat tgg gtt aat cca aat cgt gca att aag gtt gag 2139
Gly Pro Ser Ala Asn Trp Val Asn Pro Asn Arg Ala Ile Lys Val Glu
470 475 480
cca act aat ttc gta ccg tta gga gat aag ttt aaa gcg gaa tta act 2187
Pro Thr Asn Phe Val Pro Leu Gly Asp Lys Phe Lys Ala Glu Leu Thr
485 490 495
ata act tca gct gac tct cca tcg tta gaa gct att gcg atg cat gct 2235
Ile Thr Ser Ala Asp Ser Pro Ser Leu Glu Ala Ile Ala Met His Ala
500 505 510
gaa aat aac aac atc aac aac atc att ctt ttt gta gga act gaa ggt 2283
Glu Asu Asu Asn Ile Asn Asn Ile Ile Leu Phe Val Gly Thr Glu Gly
515 520 525
gct gat gtt atc tat tta gat aac att aaa gta att gga aca gaa gtt 2331
Ala Asp Val Ile Tyr Leu Asp Asn Ile Lys Val Ile Gly Thr Glu Val
530 535 540 545
gaa att cca gtt gtt cat gat cca aaa gga gaa gct gtt ctt cct tct 2379
Glu Ile Pro Val Val His Asp Pro Lys Gly Glu Ala Val Leu Pro Ser
550 555 560
gtt ttt gaa gac ggt aca cgt caa ggt tgg gac tgg gct gga gag tct 2427
Val Phe Glu Asp Gly Thr Arg Gln Gly Trp Asp Trp Ala Gly Glu Ser
565 570 575
ggt gtg aaa aca gct tta aca att gaa gaa gca aac ggt tct aac gcg 2475
Gly Val Lys Thr Ala Leu Thr Ile Glu Glu Ala Asn Gly Ser Asn Ala
580 585 590
tta tca tgg gaa ttt gga tac cca gaa gta aaa cct agt gat aac tgg 2523
Leu Ser Trp Glu Phe Gly Tyr Pro Glu Val Lys Pro Ser Asp Asn Trp
595 600 605
gca aca gct cca cgt tta gat ttc tgg aaa tct gac ttg gtt cgc ggt 2571
Ala Thr Ala Pro Arg Leu Asp Phe Trp Lys Ser Asp Leu Val Arg Gly
610 615 620 625
gaa aat gat tat gta act ttt gat ttc tat cta gat cca gtt cgt gca 2619
Glu Asn Asp Tyr Val Thr Phe Asp Phe Tyr Leu Asp Pro Val Arg Ala
630 635 640
aca gaa ggc gca atg aat atc aat tta gta ttc cag cca cct act aac 2667
Thr Glu Gly Ala Met Asn Ile Asn Leu Val Phe Gln Pro Pro Thr Asn
645 650 655
ggg tat tgg gta caa gca cca aaa acg tat acg att aac ttt gat gaa 2715
Gly Tyr Trp Val Gln Ala Pro Lys Thr Tyr Thr Ile Asn Phe Asp Glu
660 665 670
tra gag gaa gcg aat caa gta aat ggt tta tat cac tat gaa gtg aaa 2763
Leu Glu Glu Ala Asn Gln Val Asn Gly Leu Tyr His Tyr Glu Val Lys
675 680 685
att aac gta aga gat att aca aac att caa gat gac acg tta cta cgt 2811
Ile Asn Val Arg Asp Ile Thr Asn Ile Gln Asp Asp Thr Leu Leu Arg
690 695 700 705
aac atg atg atc att ttt gca gat gta gaa agt gac ttt gca ggg aga 2859
Asn Met Met Ile Ile Phe Ala Asp Val Glu Ser Asp Phe Ala Gly Arg
710 715 720
gtc ttt gta gat aat gtt cgt ttt gag ggg gct gct act act gag ccg 2907
Val Phe Val Asp Asn Val Arg Phe Glu Gly Ala Ala Thr Thr Glu Pro
725 730 735
gtt gaa cca gag cca gtt gat cct ggc gaa gag acg ccg cct gtc gat 2955
Val Glu Pro Glu Pro Val Asp Pro Gly Glu Glu Thr Pro Pro Val Asp
740 745 750
gag aag gaa gcg aaa aaa gaa caa aaa gaa gca gag aaa gaa gag aaa 3003
Glu Lys Glu Ala Lys Lys Glu Gln Lys Glu Ala Glu Lys Glu Glu Lys
755 760 765
gaa gca gta aaa gaa gaa aag aaa gaa gct aaa gaa gaa aag aaa gca 3051
Glu Ala Val Lys Glu Glu Lys Lys Glu Ala Lys Glu Glu Lys Lys Ala
770 775 780 785
atc aaa aat gag gct acg aaa aaa taatctaata aactagttat agggttatct 3105
Ile Lys Asn Glu Ala Thr Lys Lys
790
aaaggtctga tgcagatctt ttagataacc tttttttgca taactggaca tagaatggtt 3165
attaaagaaa gcaaggtgtt tatacgatat taaaaaggta gcgattttaa attgaaacct 3225
ttaataatgt cttgtgatag aatgatgaag taatttaaga gggggaaacg aagtgaaaac 3285
ggaaatttct agtagaagaa aaacagacca agaaatactg caagctt 3332
Claims (2)
1.一种通过如下方法获得的重组微生物:将编码目标蛋白质或多肽,并连接到转录起始调节区域、翻译起始调节区域或分泌信号区域的下游末端的基因整合进已缺失或失活了一个或几个参与芽孢形成中期到晚期的基因的微生物中,其中所述的转录起始调节区域、翻译起始调节区域、或分泌信号区域来自SEQ ID NO:1或2的序列,所述的微生物是枯草杆菌,所述的已缺失或失活的基因选自枯草杆菌的sigE基因、sigF基因、spoIIE基因、spoIISB基因、sigG基因、属于从枯草杆菌的spoIVCB到spoIIIC区域中的基因或与这些基因中的任意一个或几个等同的基因。
2.一种通过利用权利要求1所述的重组微生物生产蛋白质或多肽的方法。
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JP2001160520 | 2001-05-29 | ||
JP160520/2001 | 2001-05-29 |
Publications (2)
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CN1513057A CN1513057A (zh) | 2004-07-14 |
CN100439492C true CN100439492C (zh) | 2008-12-03 |
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Application Number | Title | Priority Date | Filing Date |
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Country Status (5)
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US (1) | US7585674B2 (zh) |
EP (1) | EP1391502B1 (zh) |
CN (1) | CN100439492C (zh) |
DK (1) | DK1391502T3 (zh) |
WO (1) | WO2002097064A1 (zh) |
Families Citing this family (30)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
FI112666B (fi) * | 2001-11-06 | 2003-12-31 | Ipsat Therapies Oy | Itiöimätön Bacillus subtilis, sen valmistus ja käyttö |
CN1875095B (zh) | 2003-11-07 | 2011-04-06 | 花王株式会社 | 重组微生物 |
DE102004013988A1 (de) * | 2004-03-19 | 2005-10-13 | Henkel Kgaa | Der Faktor RecA aus Bacillus licheniformis und recA-inaktivierte Sicherheitsstämme für die biotechnologische Produktion |
US7432090B2 (en) * | 2004-07-01 | 2008-10-07 | Rice University | Blocking sporulation by inhibiting SpoIIE |
JP4485341B2 (ja) * | 2004-12-20 | 2010-06-23 | 花王株式会社 | 組換え微生物 |
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- 2002-05-28 DK DK02728174.0T patent/DK1391502T3/da active
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WO2002097064A1 (fr) | 2002-12-05 |
US20040248279A1 (en) | 2004-12-09 |
US7585674B2 (en) | 2009-09-08 |
DK1391502T3 (da) | 2012-05-21 |
EP1391502A1 (en) | 2004-02-25 |
EP1391502B1 (en) | 2012-04-25 |
EP1391502A4 (en) | 2005-01-26 |
CN1513057A (zh) | 2004-07-14 |
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