CN112195129A - 紫色杆菌素生物合成基因簇及其应用 - Google Patents
紫色杆菌素生物合成基因簇及其应用 Download PDFInfo
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- CN112195129A CN112195129A CN202011139097.6A CN202011139097A CN112195129A CN 112195129 A CN112195129 A CN 112195129A CN 202011139097 A CN202011139097 A CN 202011139097A CN 112195129 A CN112195129 A CN 112195129A
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Abstract
本发明公开了一种新的菌株Janthinbacterium sp.B9‑8,并公开了来源于该菌株的紫色杆菌素生物合成相关的基因簇,所述基因序列包括五个紫色杆菌素生物合成途径基因vioA、vioB、vioC、vioD和vioE和一个SD序列,其中vioE在5’端连接一个SD序列。本发明还公开了一种工程菌,其通过将Janthinbacterium sp.B9‑8中紫色杆菌素合成途径中的基因簇构建质粒导入宿主大肠杆菌中得到的。本发明构建的工程菌可用来生产紫色杆菌素,制备紫色杆菌素。本发明发酵使用培养基成分简单,成本低廉,操作简单,产率高,最终紫色杆菌素产量达到0.13g/L,可用于工业化生产。
Description
技术领域
本发明属于基因工程技术,具体涉及一种生产紫色杆菌素的工程菌构建及其应用方法。
背景技术
紫色杆菌素(violacein)是由细菌合成的一种非水溶性次级代谢产物,属于吲哚衍生物,是以色氨酸分子为前体物质氧化缩合而成。近年研究发现,紫色杆菌素不仅可用做染色剂,其还具有抗肿瘤、抗菌、抗病毒、抗氧化等多种重要生物活性,在食品、医疗和印染工业等方面有着广阔的应用前景。紫色杆菌素生物合成途径首先由Pemberton等人研究,并由Balibar和桑切斯等人完全阐明。紫色杆菌素是由两个色氨酸分子通过由一个基因簇编码的五种蛋白质氧化缩合而成,这五个蛋白质的编码基因(vioA,vioB,vioC,vioD,vioE)是在同一方向上进行转录,生物合成总共涉及六个步骤:五个酶促反应和一个非酶促氧化脱羧步骤。
目前,已有多种微生物被证实具有紫色杆菌素合成能力,包括Chromobacteriumviolaceum、Janthinobacterium lividum、Dunganella sp.、Massilia sp.和Pseudoalteromonas luteoviolacea等,它们在系统发育上是多种多样的,从海洋,冰川,河流和土壤等不同环境中筛选获得,具有独特的生存能力。已有多种紫色杆菌素生产菌株如C.violaceum、J.lividum、Pseudoalteromonas tunicata、P.luteoviolacea、Massiliasp.WG5等菌株的全基因组被测序并已在NCBI数据库中收录,不同来源的原始菌株的紫色杆菌素基因簇序列从基因组成、大小上分别存在一定程度的差异。
发明内容
发明目的:根据上述现有技术,本发明提供了一种紫色杆菌素生物合成基因簇及其应用。
技术方案:本发明提供了一种新的菌株Janthinbacterium sp.B9-8,保藏编号为CCTCC NO:M2020183。
本发明所述的紫色杆菌素生物合成基因簇,选自如下1)或2):
1)包括五个紫色杆菌素生物合成途径基因vioA、vioB、vioC、vioD和vioE和一个核糖体结合位点SD序列,其核苷酸序列如SEQ ID NO:6所示,SD序列为5’-taatataagaaggagatata-3’;
2)与1)的基因簇具有80%以上同源性,且编码紫色杆菌素合成途径中的VioA、VioB、VioC、VioD和VioE五种酶的基因簇。
所述紫色杆菌素生物合成基因簇来源于Janthinbacterium sp.B9-8,保藏编号为CCTCC NO:M2020183。
该菌株Janthinbacterium sp.B9-8从新疆罗布泊地区筛选获得,该菌株生长较快,在25℃下经24h能达到一个较高的生物量和紫色杆菌素产量。该菌株生产的紫色杆菌素具有广谱抗菌活性、抗肿瘤活性、抗氧化、抗病毒等强生物活性,具有较好的水溶性和有机溶性。对菌株基因组进行了全测序,并鉴定获得了紫色杆菌素的合成基因簇vioABCDE,其16S rDNA序列与标准模式菌Janthinobacterium lividum DSM 1522T(Y08846)和Janthinbacterium agaricidamnosum W1r3T(Y08845)同源性最高,相似性分别为98.6%和98.1%。Janthinobacterium sp.B9-8中紫色杆菌素合成的基因簇与其他不同来源菌株的紫色杆菌素基因簇相似性不高(64-78%),尤其是与已知产紫色杆菌素的常用菌株C.violaceum和J.lividum相比,基因比对的覆盖率仅在14-17%。
表1:产紫色杆菌素菌株与Janthinobacterium sp.B9-8中的vio基因簇DNA序列对比
进一步的,所述基因vioA、vioB、vioC、vioD和vioE编码紫色杆菌素合成途径中的VioA、VioB、VioC、VioD和VioE五种蛋白质,所述蛋白的氨基酸序列依次如SEQ ID NO:1-5所示。
具体的,所述VioA蛋白具体可如附件的序列1所示。所述VioB蛋白具体可如附件的序列2所示。所述VioC蛋白具体可如附件的序列3所示。所述VioD蛋白具体可如附件的序列4所示。所述VioE蛋白具体可如附件的序列5所示。
所述参与紫色杆菌素途径的各个蛋白的编码基因具体可如附件的序列6所示。其中,所述VioA蛋白的编码基因具体可如序列表的序列6自5’末端第1-1287位核苷酸所示。所述VioB蛋白的编码基因具体可如序列表的序列6自5’末端第1345-4368位核苷酸所示。所述VioC蛋白的编码基因具体可如序列表的序列6自5’末端第4372-5661位核苷酸所示。所述VioD蛋白的编码基因具体可如序列表的序列6自5’末端第5661-6791位核苷酸所示。所述VioE蛋白的编码基因具体可如序列表的序列6自5’末端第6793-7374位核苷酸所示。
本发明还公开了一种含有上述基因簇的重组表达载体。所述重组表达载体为在表达载体pET Duet-1和pRSF Duet-1的多克隆位点插入所述基因簇得到的重组表达载体。
本发明还公开了一种重组菌,将所述基因簇或所述重组表达载体导入宿主细胞获得。所述宿主细胞为大肠杆菌,具体为大肠杆菌BL21(DE3)。
含有上述基因簇的表达盒或转基因细胞系也在本发明的保护范围内。
本发明还公开了所述基因簇、所述重组表达载体或重组菌在生产紫色杆菌素中的应用。
本发明还公开了一种生产紫色杆菌素的方法,包括以下步骤:
(1)将所述基因簇或所述重组表达载体导入宿主细胞大肠杆菌获得重组菌;
(2)步骤(1)所得重组菌发酵培养得到紫色杆菌素。
其中,步骤(2)中,所述发酵培养培养基包括:蛋白胨3-10g/L、酵母粉1-10g/L、葡萄糖0.5-10g/L、硫酸铵0-5g/L,氨苄青霉素0-100mg/mL和卡那霉素0-100mg/mL。其中,所述50mL液体发酵培养基添加50μL卡那霉素和50μL氨苄青霉素溶液。
优选的,所述培养基包括蛋白胨5g/L、酵母粉3g/L、葡萄糖1g/L、硫酸铵1g/L,氨苄青霉素100mg/mL和卡那霉素50mg/mL,pH为4。在37℃培养细胞浓度为OD600至0.8-1.0时,转为15℃发酵培养0-120h。
进一步优选的,步骤(2)中,发酵体系培养温度为15-25℃,先加入诱导剂乳糖,培养10h后加入L-色氨酸,再培养30-98h。
进一步的,所述L-色氨酸浓度为1-2mg/mL,所述乳糖浓度为40-50mg/mL,优选40mg/mL。
在所述发酵培养后,包括如下提取的步骤:将所述发酵体系离心,收集菌体;将所述菌体与无水乙醇混合,破碎菌体,离心,收集上清液,即为紫色杆菌素的提取液。上述过程中,在所述提取后,包括将所述紫色杆菌素的提取液进行高效液相色谱分离的步骤;所述高效液相色谱分离中,所用的色谱柱为Venusil XBP C18液相色谱柱,流动相为乙腈和水的混合溶液,乙腈和水的体积比为4∶6;流速为1.0mL/min;柱温为40℃,根据检测紫色杆菌素标准品,紫色杆菌素的保留时间为4.498min。
有益效果:本发明将合成紫色杆菌素能力较强的Janthinbacterium sp.B9-8中的紫色杆菌素合成基因簇导入大肠杆菌菌株中,使紫色杆菌素合成相关基因簇实现异源表达,构建了能高效合成紫色杆菌素的重组菌。本发明的制备紫色杆菌素的方法简单,发酵生产所使用的培养基成分简单、价格低廉。本发明方法操作简单,产率高,其产量可达到0.13g/L。因此,本发明所构建的重组菌及紫色杆菌素的生产方法具有广阔的应用前景。
附图说明
图1为双酶切重组质粒琼脂糖凝胶电泳结果;
图2为重组质粒pET-AB和重组质粒pRSF-CDE、pRSF-CDE-SD的构建;
图3为紫色杆菌素拟合标准曲线;
图4为重组菌株所产紫色杆菌素提取物;
图5为诱导温度对紫色杆菌素合成的影响结果;
图6为诱导pH对紫色杆菌素合成的影响结果;
图7为不同浓度的乳糖对紫色杆菌素合成的影响结果;
图8为乳糖加入时间对紫色杆菌素合成的影响结果;
图9为L-色氨酸对紫色杆菌素合成的影响结果;
图10为摇瓶发酵试验的结果。
具体实施方式
下面结合具体实施例对本申请做出详细说明。下述实施例中所使用的实验方法如无特殊说明,均为常规方法。下述实施例中所用的材料、试剂等,如无特殊说明,均可从商业途径得到。
实施例1、重组菌的构建
1)制备重组菌
Janthinbacterium sp.B9-8由实验室前期筛选获得,该菌株保藏于中国典型培养物保藏中心(CCTCC),地址:武汉市武昌珞珈山,武汉大学,保藏日期是2020年6月8日,保藏号是CCTCC No:M 2020183。大肠杆菌BL21(DE3)购买自南京诺唯赞生物科技有限公司。pET-Duet和pRSF-Duet质粒购自Novagen公司。
将Janthinbacterium sp.B9-8转接入1/3TGY培养基(0.1%酵母粉、0.167%蛋白胨、0.06%葡萄糖),28℃,200rpm培养36小时,按细菌基因组试剂盒提取Janthinbacteriumsp.B9-8基因组DNA。
根据Janthinbacterium sp.B9-8已测序获得的全基因组测序和注释结果,使用软件Vector NTI 9.0设计引物,扩增引物vioA-F、vioA-R、vioB-F和vioB-R、vioC-F、vioC-R、vioD-F、vioD-R、vioE-F、vioE-R、vioE-SD-F、vioE-SD-R的核苷酸序列见表1,以Janthinbacterium sp.B9-8基因组DNA为模板,扩增vioA和vioB基因构建至pET Duet-1载体上,扩增vioC,vioD,vioE以及vioE-SD基因构建至pRSF-Duet载体上。
利用高保真DNA聚合酶PCR扩增vio片段,PCR产物经PCR纯化试剂盒回收。利用限制性内切酶BglII和KpnI双酶切vioA片段和pET-Duet,片段纯化后进行连接,形成pET Duet-vioA,再用NcoI和HindIII双酶切vio-B片段和pET Duet-vioA,片段纯化后连接形成pETDuet-AB;利用NdeI和KpnI双酶切vio-C片段和pRSF-Duet,片段纯化后进行连接,形成pRSFDuet-vioC,再用NcoI和EcoRI双酶切vio-D片段和pRSF Duet-vioC,片段纯化后连接形成pRSF Duet-CD,之后利用KpnI和XhoI双酶切vio-E/vio-E-SD片段,片段纯化后连接形成pRSF Duet-CDE/pRSF Duet-CDE-SD,双酶切琼脂糖凝胶电泳结果如图1,其中,Lane 1:10000 DNA Marker;Lane 2:pET-AB质粒;Lane 3:BglII、KpnI酶切pET-AB质粒(vioA,1287bp);Lane 4:NcoI、HindIII酶切pET-AB质粒(vioB,3024bp);Lane 5:5000 DNAMarker;Lane 6:pRSF-CDE质粒;Lane 7:NdeI、KpnI酶切pRSF-CDE质粒(vioC,1290bp);Lane8:NcoI、EcoRI酶切pRSF-CDE质粒(vioD,1131bp);Lane 9:KpnI、XhoI酶切pRSF-CDE质粒(vioE,582bp);Lane 10:NdeI、KpnI酶切pRSF-CDE-SD质粒(vioC,1290bp);Lane 11:NcoI、EcoRI酶切pRSF-CDE-SD质粒(vioD,1131bp);Lane 12:KpnI、XhoI酶切pRSF-CDE-SD质粒(vioE-SD,600bp);Lane 13:pRSF-CDE-SD质粒;Lane 14:5000DNA Marker。双酶切后的片段用胶回收试剂盒回收。
表1紫色杆菌素基因扩增引物序列
将连接产物用热激法转化入大肠杆菌BL21(DE3)中,转化产物涂于含氨苄青霉素和卡那霉素的LB平板上,挑取转化子培养后采用质粒提取试剂盒提取质粒,筛选含插入片段的阳性克隆,经测序验证,结果插入的基因序列如序列表1-5中序列,表明插入的基因序列及方向均正确。
将重组表达载体pET Duet-AB、pRSF Duet-CDE、pRSF Duet-CDE-SD(构建过程见图2)通过酶切及测序验证后,将pET Duet-AB转化大肠杆菌BL21(DE3)中,然后将pRSF Duet-CDE、pRSF Duet-CDE-SD分别转化到含pET Duet-AB的重组菌中构建基因工程菌VioABCDE和VioABCDE-SD。转化产物涂布于含氨苄青霉素和卡那霉素的LB平板上,挑取转化子单克隆。接种于含氨苄青霉素(100mg/mL)和卡那霉素(50mg/mL)的LB液体培养基中培养,采用质粒提取试剂盒提取质粒,分别利用表1中引物对质粒进行PCR扩增和酶切的方法进行筛选验证,再进行测序验证。
将含有重组表达载体的大肠杆菌BL21(DE3)菌株,记作VioABCDE和VioABCDE-SD。
含氨苄青霉素(100mg/mL)和卡那霉素(50mg/mL)的LB液体培养基的组成:胰蛋白胨10g/L,酵母粉5g/L,NaCl 10g/L。
含氨苄青霉素(100mg/mL)和卡那霉素(50mg/mL)的LB平板的组成:胰蛋白胨10g/L,酵母粉5g/L,NaCl 10g/L,琼脂20g/L。
实施例2、制备紫色杆菌素
一、发酵菌
种子培养基:为含氨苄青霉素和卡那霉素抗生素的LB液体培养基,组成为胰蛋白胨10g/L,酵母粉5g/L,NaCl 10g/L,氨苄青霉素(100mg/mL)和卡那霉素(50mg/mL)。
发酵培养基:为含氨苄青霉素和卡那霉素抗生素的TGY液体培养基,由胰蛋白胨5g/L,酵母粉3g/L,葡萄糖1g/L、硫酸铵1g/L,氨苄青霉素(100mg/mL)和卡那霉素(50mg/mL)组成。
种子培养:将VioABCDE和VioABCDE-SD单菌落接种至50mL种子培养基,37℃,200rpm振荡培养12h,得到种子培养液。
发酵培养:按4%的接种量,将种子培养液接种至发酵培养基中,37℃培养菌株OD600至0.8-1.0时,转为15℃培养90-108h,收集容器内的所有物质,将容器内的所有物质记作发酵物。
二、提取
取2mL步骤一得到的发酵物,12000rpm离心1min并收集菌体,菌体中加入无水乙醇1mL,用漩涡混合器将其混匀,然后在200W超声波清洗器中振荡2h,随后将发酵物在12000rpm离心1min,收集上清液,即为紫色杆菌素的粗提液。
取上述得到的粗提液,用分光光度计测575nm处吸光值。用紫色杆菌素标准品制作标准曲线如图3,标准曲线方程为Y=0.0292*X+0.0005,R2=0.9996,Y代表紫色杆菌素的浓度(g/L),X代表575nm处吸光值。
重组菌VioABCDE和VioABCDE-SD经发酵提取得到的粗提液通过旋转蒸发至干燥,得到固形物质如图4。
三、高效液相色谱分离纯化
将步骤二中获得的固形物质溶于乙醇中,进行高效液相色谱分析。
所用色谱仪为Agilent 1100型液相色谱仪;色谱柱为Venusil XBP C18液相色谱柱(150mm×4.6mm,5μm),孔径5μm;柱温40℃;进样量20μL;
流动相的组成为:乙腈和水的体积比为4∶6;流速为1.0mL/min;检测波长575nm;
在如上色谱条件下,紫色杆菌素标准品的出峰保留时间为4.498min。
四、紫色杆菌素产量检测
紫色杆菌素含量是通过测定其乙醇溶液在最大吸收波长575nm的吸光度值,根据标准曲线来计算,以无水乙醇作空白对照。
标准曲线:将紫色杆菌素标准品用乙醇配制成不同浓度的溶液,在575nm下检测溶液的吸光度值;以吸光度值对应紫色杆菌素浓度作标准曲线。
色素浓度标准曲线得到相对应的色素浓度值,实验重复3次,实验结果以三次实验的平均值表示。
实施例3、表达紫色杆菌素合成相关基因簇的重组菌高效生产紫色杆菌素
1)诱导温度对紫色杆菌素合成的影响
将工程菌VioABCDE和VioABCDE-SD分别接种至含相应的抗生素的LB液体培养基,37℃,200rpm振荡培养过夜。次日按4%的接种量,接种至新鲜含相应的抗生素的发酵培养基中,37℃继续培养2-3h至OD600为0.8-1.0时,分别在15℃、20℃、25℃、30℃和37℃诱导培养48h后,离心收集菌体,提取紫色杆菌素,定量分析诱导温度对紫色杆菌素合成的影响,如图5所示。
2)诱导pH对紫色杆菌素合成的影响
将工程菌VioABCDE和VioABCDE-SD接种至含相应的抗生素的具有不同pH值(pH4、pH5、pH6、pH7和pH8)的发酵培养基中37℃培养2-3h至OD600为0.8-1.0时,在20℃诱导培养48h后,离心收集菌体,提取紫色杆菌素,定量分析诱导pH对紫色杆菌素合成的影响,如图6所示。
3)正交试验确定最佳培养条件
以诱导温度、诱导pH、诱导时间3个因素为试验因素,以紫色杆菌素产量为指标,对VioABCDE和VioABCDE-SD合成紫色杆菌素的能力进行3因子3水平的正交试验,VioABCDE和VioABCDE-SD合成紫色杆菌素能力的结果见表2和表3。
表2菌株VioABCDE关于诱导温度、诱导pH和诱导时间的正交实验直观分析表
注:因素1:温度,1234分别表示15℃、20℃、25℃、30℃
因素2:诱导pH,1234分别表示pH7、pH6、pH5、pH4
因素3:诱导时间,1234分别表示48h、96h、120h、108h
表3 VioABCDE-SD关于诱导温度、诱导pH和诱导时间的正交实验直观分析表
注:因素1:温度,1234分别表示15℃、20℃、25℃、30℃
因素2:诱导pH,1234分别表示pH7、pH6、pH5、pH4
因素3:诱导时间,1234分别表示48h、96h、120h、108h
分析得到最佳实验方案:培养基在pH4条件下,诱导温度为15℃,培养108h后紫色杆菌素得到最大产量。
将工程菌VioABCDE和VioABCDE-SD接种至含相应的抗生素的pH为4的发酵培养基中37℃培养2-3h至OD600为0.8-1.0时,在15℃诱导,离心收集菌体,提取紫色杆菌素,实验结果表明,菌株在pH4的TGY培养基中培养,诱导温度为15℃,诱导时间为108h,VioABCDE和VioABCDE-SD合成紫色杆菌素的含量最高,其中VioABCDE-SD的产量高于VioABCDE。
4)诱导剂乳糖对紫色杆菌素合成的影响
将工程菌VioABCDE和VioABCDE-SD分别接种至含相应的抗生素的LB液体培养基,37℃,200rpm振荡培养过夜。次日按4%的接种量,接种至新鲜含相应的抗生素的发酵培养基中,37℃继续培养2-3h至OD600为0.8-1.0时,加入不同浓度(10-100mg/mL)的诱导剂乳糖,对VioABCDE-SD合成紫色杆菌素的影响如图7所示,表明加入40mg/mL乳糖时VioABCDE-SD产生最大浓度的紫色杆菌素。
将工程菌VioABCDE和VioABCDE-SD分别接种至含相应的抗生素的LB液体培养基,37℃,200rpm振荡培养过夜。次日按4%的接种量,接种至新鲜含相应的抗生素的发酵培养基中,37℃继续培养,在接种后2、4、6、8、10、12、14h分别加入40mg/mL乳糖进行诱导,培养OD600至0.8-1.0时,将温度转为15℃继续培养108h,如图8所示,诱导2h时加入终浓度为40mg/mL的乳糖溶液时紫色杆菌素产量达到最高。
5)L-色氨酸对紫色杆菌素合成的影响
将工程菌VioABCDE和VioABCDE-SD分别接种至含相应的抗生素的LB液体培养基,37℃,200rpm振荡培养过夜。次日按4%的接种量,接种至新鲜含相应的抗生素的发酵培养基中,37℃继续培养,在接种后2h加入40mg/mL乳糖进行诱导,培养OD600至0.8-1.0时,将温度转为15℃继续培养10h时,加入L-色氨酸使其在培养基中达到不同终浓度,继续15℃培养98h。如图9所示,加入L-色氨酸终浓度为2mg/mL时紫色杆菌素产量最高。
实施例4、摇瓶发酵实验
将实施例1制备的工程菌进行摇瓶发酵实验
摇瓶发酵实验以初始基础培养基为对照。
1)初始发酵培养基的溶剂为水,溶质及浓度如下:胰蛋白胨5g/L,酵母粉3g/L,葡萄糖1g/L,硫酸铵1g/L,氨苄青霉素(100mg/mL)和卡那霉素(50mg/mL),pH为7。
①将重组菌单克隆接种至250mL摇瓶(每瓶装有50mL的LB培养基),37℃、200rpm震荡培养12小时。
②取步骤①得到的菌液,按4%的接种量接种至新的250mL摇瓶(每瓶装有50mL的发酵培养基),37℃、200rpm震荡培养2h至OD600为0.8-1.0。
③完成步骤②后,将菌液在20℃、200rpm培养至48h。
2)优化后的发酵培养基的溶剂为水,溶质及浓度如下:蛋白胨5.0g/L,酵母粉3.0g/L,葡萄糖1.0g/L,硫酸铵1g/L,氨苄青霉素(100mg/mL)和卡那霉素(50mg/mL),pH为4。
①将重组菌单克隆接种至250mL摇瓶(每瓶装有50mL的LB培养基),37℃、200rpm震荡培养12小时。
②取步骤①得到的菌液,按4%的接种量接种至新的250mL摇瓶(每瓶装有50mL的TGY培养基),37℃、200rpm震荡培养2h至OD600为0.8-1.0。
③完成步骤②后,加入终浓度为40mg/mL的诱导剂乳糖,15℃、200rpm震荡培养10h后加入终浓度为2mg/mL的L-色氨酸,之后继续培养98h。
④完成步骤③后,将整个发酵体系混匀取样,每个发酵体系取样3mL,分为两份,一份2mL,另一份1mL。
⑤取步骤④得到的2mL样本,12000rpm离心1min并收集沉淀,将沉淀加入1mL无水乙醇,用涡旋振荡器混匀后于超声清洗仪中提取2h,12000rpm离心1min并收集上清液,用分光光度计测575nm处吸光值,根据575nm处吸光值和标准曲线方程,计算步骤③得到的发酵体系中紫色杆菌浓度。
⑥取步骤④得到的1mL样本,12000rpm离心1min并收集沉淀,将沉淀加入1mL生理盐水,震荡混匀,用分光光度计测600nm处吸光值。
摇瓶发酵结果见图10,优化之后的发酵体系,每升发酵液中含有紫色杆菌素0.13g,比优化之前提高了7.5倍。
序列表
<110> 南京师范大学
<120> 紫色杆菌素生物合成基因簇及其应用
<160> 6
<170> SIPOSequenceListing 1.0
<210> 1
<211> 428
<212> PRT
<213> Janthinbacterium sp. B9-8
<400> 1
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Phe Asp His Pro Trp Trp Lys Asp Tyr Glu Leu Glu Asn Lys Val Thr
275 280 285
Ile Val Asp Asn Pro Leu Arg Lys Ile Tyr Phe Lys Ser Glu Lys Tyr
290 295 300
Ile Phe Phe Tyr Thr Asp Ser Ala Tyr Ala Asp Phe Trp Leu Glu Glu
305 310 315 320
Thr Asn Lys Ser Glu Asp Glu Tyr Ile Asn Thr Val Met Gly Leu Ile
325 330 335
Ala Lys Ala Leu Asn Ile Ser Ile Lys Glu Leu Pro Arg Pro Val Ser
340 345 350
Asn Lys Phe Lys Tyr Trp Pro Ser Gly Val Glu Phe Ala Leu Glu Thr
355 360 365
Ser Pro Asp His Pro Pro Val Leu Ser Lys Tyr Asn Gly Lys Val Ile
370 375 380
Ala Thr Ser Asp Ala Tyr Thr Pro His Cys Gly Trp Met Glu Gly Gly
385 390 395 400
Ile Ile Ala Gly Arg Asn Ala Ala Asp His Ile Leu Lys Gln Leu Glu
405 410 415
Lys Ser Asp Val Glu Lys Val Ala Ala Glu Met Asn
420 425
<210> 2
<211> 1007
<212> PRT
<213> Janthinbacterium sp. B9-8
<400> 2
Met Ser Ile Leu Asn Phe Pro Arg Ile His Phe Lys Gly Ala Ala Arg
1 5 10 15
Leu Asn Val Pro Thr Gly Asn Arg Asn Ile Ala Gly Thr Leu Asp Ile
20 25 30
Ala Thr Asn Ser Val Tyr Gln Asp Gly Thr Leu Phe Asp Leu Lys Lys
35 40 45
His Pro Arg Glu Phe His Gln Tyr Leu Arg Gln Leu Pro Pro Arg Phe
50 55 60
Asn Leu Gln Gly Gln Pro Asp Asp Ala Gly Val Phe Asn His Ala Ala
65 70 75 80
Gly His Asn Phe Leu Gly Asn Asn His Phe Ser Trp Glu Asn Thr Gln
85 90 95
Ile Thr Ala Leu Gln Ile Gln Pro Gly Val Leu Leu Asp Asn Asp Pro
100 105 110
Leu Leu Ala Cys Lys Val Glu Leu Trp Gly His Tyr Asn Glu Tyr Leu
115 120 125
Arg Thr Thr Phe Asn Arg Ala Arg Trp Val Asp Ile Asp Pro Ser Arg
130 135 140
Asp Asp Thr Ser Gln Ile Tyr Ala Gly Gln Leu Cys Ile Lys Glu Ala
145 150 155 160
Ala Ala Ala Ala Asn Ala Pro Tyr Leu Phe Ser Ser Asp Ile Asp Cys
165 170 175
Val His Ser Val Arg Trp Tyr Gly Glu Asn His Ile Gln Glu Lys Gln
180 185 190
Pro His Phe Leu Glu Pro Glu Phe Ser Lys Thr Arg Val Phe Gln Phe
195 200 205
Ser Val Ala Lys Asp Ser Glu His Phe Ile Phe Asn Gln Leu Ser Leu
210 215 220
Lys Ser Asp Phe Ile Glu Tyr Leu Gln Gln Glu Leu Lys Arg Glu Asp
225 230 235 240
Val Leu Gly Leu Ser Ile Gln Tyr Ser Val Phe Asn Met Ser Thr Pro
245 250 255
Leu Gln Pro Asp Ile Pro Val Phe Tyr Asp Leu Ser Gly Ser Ile Gly
260 265 270
Leu Trp Leu Lys Gln Asp Met Ala Thr Phe Ala Asn Asp Arg Ile Leu
275 280 285
Tyr Pro Asp Asn Thr Gln Gly Phe Gly Pro Val Ala Ile Lys Ile Gln
290 295 300
Asn Asn Trp Ala Ser Ile Ser Met Pro Cys Ser Ile Pro Phe Thr Arg
305 310 315 320
Arg Asp Ala Val Cys Gly Asn Gln Leu Thr His Pro Leu Gly Pro Lys
325 330 335
His Ser Leu Gly Glu Leu Glu Leu Arg Thr Lys Gly Gly Val Leu Leu
340 345 350
Ala Arg Ile Pro Glu Ser Val Tyr Gly Pro Tyr Trp Gly Ser Ala Gly
355 360 365
Ile Phe Asp Val Pro Leu Leu Asn Asp Ala Ala Glu Asp Ser Leu Ile
370 375 380
Leu Arg Gly Thr Asp Asn Gln Trp Val Glu Ser Asp Trp Leu Ile Gln
385 390 395 400
Ala Glu Gln Asn Val Val Ser Leu Glu Ala Pro Asp Arg Lys Asn Asn
405 410 415
Ala Phe Phe Ser Lys Asp Ile Lys Ile Phe Ser Tyr Phe Arg Gly Glu
420 425 430
Pro Lys Gly Lys Lys Asp Ile Ser Val Phe Ile Glu Asn Lys Ala Val
435 440 445
Val Ser Thr Ser Leu Gln Thr Ile Ser Ser Asn Glu Asn Gly Ile Ala
450 455 460
Phe Leu Ser Ile Val Ser Gln Lys Pro Gly Val Ser Glu Val Phe Val
465 470 475 480
Gly Asp His His Gly Lys Ile Leu Val Arg Val Leu Ser Asp Asp Trp
485 490 495
Ala Leu Leu Glu Val Ala Asp Lys Glu Val Asp Tyr Pro Phe Leu Tyr
500 505 510
Glu Gln Val Met Ser Tyr Tyr Glu Leu Ile Tyr Pro Phe Met Ala Asp
515 520 525
Lys Val Phe Ser Met Ala Asp Gln Cys Lys Cys Glu Thr Tyr Ala Arg
530 535 540
Leu Met Trp Gln Met Cys Asp Pro Leu Asn Arg Asp Lys Ser Tyr Tyr
545 550 555 560
Met Pro Ser Thr Arg Glu Met Ser His Ala Lys Ser Ile Leu Phe Leu
565 570 575
Lys Tyr Leu Ser Asn Val Glu Ala Ser Ala Ile Ala Glu Pro Ala Lys
580 585 590
Ile Ala Leu Ala Thr Glu Val Asn Ser Ser Glu Ile Ser Ser Lys Glu
595 600 605
Glu Leu Val Gln Val Leu Lys Gln Ala Val Asn Leu Glu Leu Ser Ile
610 615 620
Met Leu Gln Tyr Leu Tyr Ala Ala Tyr Ser Leu Pro Thr Phe Ala Ala
625 630 635 640
Gly Glu Gln Leu Leu Ala Ser Asn Lys Trp Thr Ala Glu Gln Leu Ala
645 650 655
Leu Val Cys Gly Gly Ala Asp Arg Arg His Asn Ser Gly Trp Arg Gly
660 665 670
Thr Ile Leu Glu Ile Ala His Glu Glu Met Ile His Tyr Leu Ile Val
675 680 685
Asn Asn Leu Leu Met Ser Leu Gly Glu Pro Phe Phe Pro Gly Thr Pro
690 695 700
Val Leu Gly Leu Lys Ala Ala Glu Ala Phe Gly Leu Asp Thr Asp Phe
705 710 715 720
Ser Phe Glu Pro Phe Ser Glu Asn Val Leu Ala Arg Phe Val Arg Phe
725 730 735
Glu Trp Pro His Tyr Phe Pro Ser Thr Gly Lys Ser Ile Ser Asp Phe
740 745 750
Tyr Ala Glu Ile Arg Lys Ala Phe Val Ser Ile Pro Asp Leu Phe Thr
755 760 765
Thr Pro Leu Ala Lys Thr Ser Gly Glu His His Leu Phe Leu Asn Glu
770 775 780
Leu Thr Asn Arg Lys Phe Pro Ala Tyr Gln Leu Glu Val Asn Asn Arg
785 790 795 800
Glu Ser Ala Leu Phe Ala Ile Asp Phe Val Thr Glu Gln Gly Glu Gly
805 810 815
Ala Ala Val Asp Ser Pro His Tyr Ala Gln Ser His Phe Asn Arg Leu
820 825 830
Arg Glu Ile Ser Arg Thr Leu Leu Ala Ser Glu Thr Pro Phe Glu Pro
835 840 845
Ala Phe Ser Val Leu Lys Asn Pro Val Leu Glu Pro Thr Pro Gly Cys
850 855 860
Asn Thr Val Leu Asp Pro Ala Ala Arg Gln Leu Met Ile Phe Tyr Lys
865 870 875 880
Gly Cys His Glu Leu Thr Phe His Met Met Ile Gln His Phe Gly Gln
885 890 895
Lys Pro Leu Gly Ser Leu Arg Arg Ser Arg Leu Met Asn Ala Ala Ile
900 905 910
Asp Ile Met Ala Gly Ile Leu Arg Pro Leu Ser Val Gln Ile Met Ser
915 920 925
Val Pro Ser Gly Val Ser Gly Arg Asn Ala Gly Pro Pro Val Pro Gln
930 935 940
Ala Ile Gln Phe Gln Leu Ile Ala Asp Tyr Gly Ala Gly Cys Glu Val
945 950 955 960
Leu Ala Leu Gln Cys Arg Ala Leu Ala Lys Tyr Ala Lys Glu Met Gln
965 970 975
Leu Ile Lys Pro Ala Met Ala Gln Ile Glu Leu Leu Glu Phe Tyr Asp
980 985 990
Lys Gln Met Met Asp Leu Ala Thr Gly Lys Leu Ser Arg Glu Gly
995 1000 1005
<210> 3
<211> 429
<212> PRT
<213> Janthinbacterium sp. B9-8
<400> 3
Met Lys Lys Ile Ile Ile Val Gly Gly Gly Leu Ala Gly Ser Leu Thr
1 5 10 15
Ala Ile Tyr Leu Ala Lys Arg Gly His Asp Val His Val Phe Glu Lys
20 25 30
Arg Gly Asp Pro Phe Leu Ala Tyr Ser Asp Tyr Ile Asp Gln Val Ser
35 40 45
Ser Arg Ala Ile Gly Val Ser Met Thr Val Arg Gly Ile Gln Ala Val
50 55 60
Leu Asn Ala Gly Ile Pro Lys Glu Glu Leu Asp Leu Cys Gly Ile Pro
65 70 75 80
Ile Ser Gly Met Ser Phe Cys Ile Ala Gly Lys Phe Lys Thr Arg Glu
85 90 95
Leu Ala Pro Val Asp Val Leu Ser Pro Leu Ser Leu Ser Arg Ala Asp
100 105 110
Phe Gln Lys Leu Leu Asn Lys Tyr Ala Glu Ile Asn Asn Val His Tyr
115 120 125
His Tyr Gly Asn Arg Cys Leu Glu Val Asn Leu Asp Asp Lys Ser Ile
130 135 140
Thr Thr Lys Asp Asn Asp Gly Gln Ile Ala Glu His Lys Ala Asp Leu
145 150 155 160
Leu Ile Gly Ala Asp Gly Ala Arg Ser Cys Val Arg Gln Ala Met Gln
165 170 175
Asn Asn Cys Arg Arg Phe Glu Tyr Gln Gln Ser Phe Phe Lys His Gly
180 185 190
Tyr Lys Thr Ile Val Ile Pro Asp Ala Ser Gln Leu Gly Leu Arg Lys
195 200 205
Asp Leu Ile Tyr Phe Phe Gly Met Asp Ser Gly Gly Leu Phe Ala Gly
210 215 220
Arg Ala Ala Thr Ile Pro Asp Gly Ser Ile Ser Phe Ala Val Cys Leu
225 230 235 240
Pro Tyr Gln Gly Glu Ile Ser Leu Gln Ala Lys Asp Ala Gly Thr Met
245 250 255
Gly Arg Phe Phe Asp Arg Tyr Tyr Ser Met Leu Pro Glu Ala Thr Arg
260 265 270
Lys Glu Met Leu Ser Gln Phe Met Glu Lys Pro Ser Asn Asp Leu Ile
275 280 285
Asn Val Arg Ser Ser Val Phe His Tyr Lys Gln His Ala Ile Ile Leu
290 295 300
Gly Asp Ser Ala His Ala Thr Ala Pro Phe Leu Gly Gln Gly Met Asn
305 310 315 320
Met Ala Leu Glu Asp Ala Tyr Ile Leu Asn Leu Leu Phe Glu Lys His
325 330 335
Glu Asp Asn Ile Glu Lys Val Leu Pro Glu Phe Thr Arg Leu Arg Lys
340 345 350
Ala Glu Ala Asp Ala Met Gln Asp Met Ala Ile Asn Asn Tyr Glu Val
355 360 365
Leu Ser Ser Ser Asn Pro Ile Phe Phe Met Arg Ser Lys Tyr Thr Arg
370 375 380
Tyr Met Ser Ser Lys Phe Pro Ala Ser Tyr Pro Pro Asp Met Ala Glu
385 390 395 400
Lys Leu Tyr Phe Thr Ser Met Ala Tyr Ser Glu Leu Arg Asp Ile Gln
405 410 415
Gln Lys Gln Asn Val Trp Tyr Lys Leu Gly Arg Val Asn
420 425
<210> 4
<211> 376
<212> PRT
<213> Janthinbacterium sp. B9-8
<400> 4
Met Lys Ile Leu Val Val Gly Ala Gly Pro Ala Gly Leu Met Phe Ala
1 5 10 15
Ser Gln Leu Lys Lys Leu Lys Gln Asp Trp Asp Ile Ala Ile Val Glu
20 25 30
Lys Asn Thr Val Asp Glu Ile Val Gly Trp Gly Val Val Leu Pro Gly
35 40 45
Lys Ala Pro His His Pro Ala Asn Pro Leu Ser Tyr Leu Pro Asp Tyr
50 55 60
Glu Glu Ile Asp Ala Gln Tyr Ile Asp Glu Phe Cys Leu Val Asn Gln
65 70 75 80
Asp Asp Arg Ala Thr Ala Ser Thr Gly Ile Thr Leu Cys Gly Ala Glu
85 90 95
Arg Lys Ser Leu Val Gly Ala Leu Arg Lys Leu Cys Thr Asn Leu Leu
100 105 110
Ile Pro Ile Thr Tyr Ser Ser Pro Val Phe Asp Glu Asp Gly Leu Asp
115 120 125
Thr Thr Ala Tyr Asp Leu Val Val Ile Ala Asn Gly Ile Asn Asn Ile
130 135 140
Ser Asn Tyr Phe Lys Asp Ala Leu Ala Pro Glu Val Glu Phe Gly Lys
145 150 155 160
Asn Arg Tyr Met Trp Tyr Gly Thr Ser Lys Ile Phe Asp Ala Met Asn
165 170 175
Leu Ile Phe Lys Pro Thr Thr Ser Gly Val Phe Ile Ala His Ala Tyr
180 185 190
Lys Tyr Ser Ser Thr Met Ser Thr Phe Val Val Glu Cys Ser Glu Glu
195 200 205
Thr Tyr His Arg Ser Gly Ile Glu Ser Leu Ser Glu Thr Asp Ala Lys
210 215 220
Ala Phe Ile Ala Asp Val Phe Ser Val Glu Leu Asp Gly Leu Pro Val
225 230 235 240
Glu Val Gln Pro Gly Leu Gln Trp Arg Asn Phe Val Thr Leu Ser His
245 250 255
Thr Lys Ala Tyr Glu Asp Asn Leu Val Leu Leu Gly Asp Ala Leu Gln
260 265 270
Thr Gly His Phe Ser Ile Gly His Gly Thr Thr Met Ala Val Val Ala
275 280 285
Ala Gln Met Leu Val Lys Ala Leu Tyr Glu His Ala Glu Val Asn His
290 295 300
Ala Leu Glu Asp Phe Asn Gln Lys Val Met Pro Leu Met Gln Leu Phe
305 310 315 320
Ser Gly His Ala Gly Ile Ser Arg Thr Trp Phe Glu Thr Ala Asp Glu
325 330 335
Arg Met Asp Leu Thr Ala Ala Glu Leu Ala Lys Ser Phe Ser Glu Arg
340 345 350
Arg Glu Gln Leu Pro Pro Leu Pro Ala Ala Leu Gly Gln Ala Leu Gly
355 360 365
Met Ala Leu Thr Arg Lys Glu Ser
370 375
<210> 5
<211> 193
<212> PRT
<213> Janthinbacterium sp. B9-8
<400> 5
Met Ile Ala Glu Lys Met Leu Pro Pro Arg Leu Pro Glu Gln Trp Ser
1 5 10 15
Ser Ser Tyr Ile Ser Tyr Trp Gln Pro Met Gln Lys Glu Asp Gln Ile
20 25 30
Thr Ser Gly Ile Cys Trp Phe Asp Tyr Glu Gln Asn Arg Cys Arg Ile
35 40 45
Asp Gly Leu Phe Asn Pro Trp Ser Glu Glu Lys Thr Gly His Arg Leu
50 55 60
Trp Met Ser Glu Ile Val Thr Ala Gly Glu Gly Lys Thr Lys Lys Ser
65 70 75 80
Lys Ile Ala Tyr Cys Arg Glu Ser Pro Leu Gly Glu Gly Glu Tyr Glu
85 90 95
Ala Val Val Leu Asp Asp Asp Leu Glu Ser Cys His Glu Val Leu Leu
100 105 110
Thr Gln Asp Val Leu Leu Gln Tyr Gln Ala Ser Tyr Val Gly Ser Gln
115 120 125
His Leu Leu Gly Arg Glu Thr Glu Ala Trp Thr Phe Thr Lys Pro Gly
130 135 140
Lys Gly Pro Ser Thr Tyr Tyr Phe Val Lys Gly Thr Asn Gln Leu Leu
145 150 155 160
Arg Met Val Thr Gly Asp Pro Ala Val His Ala Ser Ile Arg Asp Phe
165 170 175
Pro Asn Phe Thr Thr His Thr Ile Pro Ala Glu Ile Phe Ser Ser Ser
180 185 190
Ile
<210> 6
<211> 7374
<212> DNA
<213> Janthinbacterium sp. B9-8
<400> 6
gtgataaaca acaaagtaat ctctatagta ggtgcaggaa tatcagggct aagctgtgca 60
ttgaaactgg ctggctcaaa attaacaaaa gactacacgc ttcgtgtttt tgaaaacgga 120
gagagaattg gtggccgggc tcattcaatt aaagtggatg attttagtat tgatttagga 180
gcagggcgtt tttctcctgc cctacatcct aatgttgccc agttgcttag tgagctaaaa 240
gaggaaatag aagtatttcc ttttactaaa attgtctgtc ctcacccaca gcatgaaggc 300
ttgaaagaga tactggctca attaaagcta aaaattgaga atagccataa tgaatcattc 360
tttcaatttt tatgccagca tatgggcaat gaaaaatcgc acgcaatcat taatgcttta 420
ggctacgatt cattatatct tccgcaaatc tctcctaaga ttgcatatga tattattgaa 480
aagcatccgg aaatacagtg cttctcagaa aatgaaggct atgagtggtt taatttagtt 540
gacgggtttg ctgctttggc taaaagtcta tatcagcaag ccgctaaatc gggtgtcgag 600
ttttattttg aacaccagtt aattaatttc caaacaaaat cagccaatac tctgcttgag 660
tttgcaggat cagaacagca agaaatttgg cacaatagtg cttataccgt tttagctctt 720
cctccaacag cgatgtcatc gcttaatata gattttccaa atagctggag tgatttttcc 780
tacggttcta tcccattatt taaaggtttt atatttttcg atcatccatg gtggaaagat 840
tatgagcttg aaaataaagt aaccattgtt gataacccat tacgaaaaat atattttaaa 900
agcgaaaaat atatattctt ttacactgat agcgcctatg ctgatttttg gttggaagaa 960
accaataaaa gcgaagatga atatataaat acagtgatgg gtttaattgc taaagcatta 1020
aatatatcta ttaaagaatt accacgccct gtaagcaata agttcaaata ctggcctagt 1080
ggtgtagagt ttgctttaga aacatcacca gatcaccctc ctgttttatc taaatataat 1140
ggcaaagtga ttgctacttc agatgcttat accccgcact gtggctggat ggaaggtggg 1200
attattgcgg gtaggaacgc agcagatcat atcttaaaac aattagagaa atcagatgta 1260
gaaaaagtag cggcagaaat gaattaattt ttgtttgaaa atgtaatggt tttttttcta 1320
attttaaact ggtattgatg cttaatgagt attctgaatt ttccacgtat tcactttaaa 1380
ggtgctgcac ggcttaatgt gccaactggc aatagaaata tcgccggcac tttagacatt 1440
gcaacaaaca gtgtttatca agatggcaca ttatttgatt taaaaaagca ccctagggag 1500
tttcatcagt atctaaggca attgccacca agatttaatc ttcaggggca gccagatgat 1560
gcaggtgttt ttaatcatgc ggcagggcat aactttttag ggaataatca tttttcttgg 1620
gaaaatactc agattaccgc tttgcaaata cagccgggtg ttttgctgga taacgatccg 1680
ctacttgctt gtaaagtgga attatggggg cactataatg agtatttgcg caccacattt 1740
aaccgggcaa gatgggtaga tattgatcca agccgggacg atacttcgca aatctatgcc 1800
gggcagctgt gcataaaaga agcggctgct gcggcaaatg ccccctattt attttcatct 1860
gacattgatt gcgttcactc tgttcgctgg tatggcgaga atcatattca ggaaaagcag 1920
ccgcatttct tggagcctga attttctaag acaagggtgt ttcagttttc tgtggctaag 1980
gattctgagc attttatttt taatcagctt tcgcttaaat cagattttat tgagtattta 2040
caacaagagc taaaaagaga agatgtgctc gggctgagta ttcaatattc tgtatttaat 2100
atgtctactc cgctgcagcc agatatacct gtgttttatg atttaagtgg cagtattggt 2160
ttgtggttaa agcaagatat ggctactttt gccaatgata ggattttata tcctgataat 2220
actcagggct ttggccctgt ggcgatcaag attcaaaata actgggctag tatcagcatg 2280
ccttgcagta ttccttttac tcgccgtgat gctgtttgtg gcaatcaact gactcaccct 2340
cttggcccta agcattcatt aggtgaatta gagctaagga ctaaaggtgg tgttttactg 2400
gcaagaattc ctgaatcggt atatggccca tattggggaa gcgccgggat ttttgatgtg 2460
cccttattaa atgatgccgc tgaagattct ttaatactgc gtggcacaga caatcaatgg 2520
gtagagtcag attggctgat tcaagcagaa cagaatgtgg tttctttgga agcgcctgat 2580
agaaaaaata atgctttttt ctctaaagac atcaaaatat ttagttattt ccgtggcgag 2640
cctaaaggaa aaaaagatat tagtgttttt attgagaata aagcggttgt cagcacttcg 2700
ttgcagacga tttcaagcaa tgaaaacgga attgcatttc tttctatcgt cagtcagaag 2760
cccggcgtta gcgaggtttt tgtaggggac catcatggca agattctggt gcgtgtactt 2820
tctgatgatt gggctttatt agaggtagcg gataaggagg ttgactatcc ttttctgtat 2880
gagcaagtaa tgagttatta cgagctgatt tatccgttta tggcggataa agtgttcagc 2940
atggcagatc agtgcaaatg tgaaacctat gctcgcttga tgtggcaaat gtgtgatcca 3000
ttaaaccgtg ataaaagtta ttacatgcct agtaccagag aaatgtctca tgccaaatca 3060
atattatttc ttaagtattt aagtaatgtt gaagcgtctg ccattgcgga gccagcaaag 3120
attgctttgg ccaccgaggt aaatagtagc gaaatcagca gtaaagaaga attagttcag 3180
gtattaaagc aggcggtgaa tttagagcta tctattatgc tgcaatattt atatgcggct 3240
tattctttgc cgacttttgc ggcaggtgag caattgcttg cgagtaataa atggactgct 3300
gagcaattgg cactggtgtg tggcggggca gatcgtcggc ataattcggg ctggcgtggc 3360
acaattctgg aaatcgccca tgaagaaatg atccactatc taattgtgaa taatttactg 3420
atgtctttag gcgagccatt ttttcctgga actcccgtat tgggtctcaa agccgctgaa 3480
gcctttggcc ttgatacaga tttttctttt gaaccttttt cagaaaatgt gctggctcgc 3540
tttgtcaggt ttgaatggcc acattatttt ccatcaacag gaaaatcaat ttctgatttt 3600
tatgcggaaa ttcgcaaggc atttgtcagt attccggatt tatttaccac cccattagct 3660
aaaacaagcg gggagcatca tttattttta aatgaattaa ccaacagaaa gtttcctgct 3720
taccagttag aagtcaataa tagagaaagt gcgctgtttg ccattgattt tgtaacagaa 3780
caaggcgagg gtgcagcagt agattcacca cattatgctc aaagccattt taatcgatta 3840
agagagatct cccgcacttt actggctagc gagactcctt ttgagcctgc attttctgta 3900
ttaaaaaatc ctgtattaga accgactccc ggctgtaata cggtgcttga tcctgccgcc 3960
agacaattaa tgatttttta taaaggctgc catgagctga cttttcatat gatgattcag 4020
cactttgggc aaaagccttt agggagtttg cgccgttcca gattaatgaa tgcggcgatt 4080
gatattatgg ccggaatatt aaggccttta tcggtgcaaa ttatgtctgt tccttctggt 4140
gtgtcgggcc gtaatgccgg gccgcctgtg ccacaggcga ttcaatttca attgattgcc 4200
gattatggcg ctggctgtga ggtactcgcc ttgcaatgcc gcgcgcttgc aaaatacgca 4260
aaagaaatgc aattaattaa accggctatg gctcaaattg aattacttga attttatgat 4320
aagcagatga tggatcttgc aacaggtaaa ctttcacggg aaggctgaaa aatgaaaaaa 4380
atcattattg ttggcggtgg cttggcaggt agcttaaccg ctatctattt agctaagcgc 4440
ggtcatgatg tgcatgtttt tgaaaaacgg ggcgatccgt ttcttgcata ttcagattat 4500
atcgatcagg ttagctcccg tgcgattggc gtgagtatga ctgtgcgtgg tattcaggcg 4560
gtattaaatg caggcattcc taaggaagag ctcgatttat gcgggattcc tatctccgga 4620
atgtcttttt gtattgcagg taaatttaaa accagagaat tagcaccggt tgatgtgctt 4680
tctccgcttt cattaagcag ggcagatttc caaaagttat taaataaata tgcagaaatt 4740
aataatgttc attaccacta tgggaatcgc tgcttggaag tgaatttaga cgataagagc 4800
attaccacta aagataatga cgggcagatt gccgagcata aagcggattt attgattggt 4860
gcagatggtg ctcgctcttg cgtacgtcag gccatgcaaa ataattgccg ccggtttgaa 4920
tatcagcagt cattttttaa gcatggctat aaaacaattg ttatccctga tgccagccag 4980
cttggcctca gaaaagattt aatttatttc tttggtatgg attctggcgg tttatttgcg 5040
gggcgtgcgg cgactattcc tgatggcagt attagttttg ctgtttgttt gccttatcaa 5100
ggagaaatta gtttacaggc taaagatgca ggtacgatgg ggcgtttttt tgatcgatat 5160
tattcgatgc tgcctgaagc caccagaaaa gaaatgctat cgcaatttat ggagaagcca 5220
agtaatgatc tgattaatgt gaggtcgtcg gtttttcatt ataagcagca cgcaatcatt 5280
ttaggcgatt cagcccatgc caccgctcca tttcttgggc agggaatgaa tatggcattg 5340
gaagacgctt atatccttaa tttattattt gaaaaacatg aagataatat tgaaaaagta 5400
ctgcctgaat ttactcggct gagaaaagcc gaggcagatg cgatgcaaga tatggctatt 5460
aataactatg aggtgctgag cagttcaaat cctattttct ttatgcgttc aaaatatact 5520
cgttatatga gtagtaaatt ccctgccagt tatcctccgg atatggctga aaagctttat 5580
ttcacctcga tggcttatag tgagctaaga gatatccaac agaaacaaaa tgtttggtac 5640
aaactaggga gagtaaacta atgaagatct tagtcgttgg agcagggcca gcgggcttaa 5700
tgtttgccag ccagttaaag aaacttaagc aagattggga tattgcaatt gtagaaaaaa 5760
atacagttga tgaaattgta gggtggggtg ttgtattgcc tggaaaagcg ccccatcatc 5820
cggcaaatcc gcttagctat ttgcctgatt atgaagaaat tgatgcgcaa tatattgatg 5880
aattttgcct ggttaatcaa gatgatcgtg ctacagcaag caccggtatt actttatgcg 5940
gggcagaaag aaaatctttg gtaggcgctt taagaaaact ctgtaccaat ttattgattc 6000
cgattactta ttcttcgcca gtttttgatg aagatggctt ggataccacg gcatatgatc 6060
tggttgtgat tgccaatggg attaataata tctcgaatta ctttaaagat gctttagccc 6120
cagaggtaga gttcggtaaa aatcgctata tgtggtacgg cacatcaaag atttttgatg 6180
caatgaattt gatttttaag ccaaccactt caggtgtatt tattgcccat gcttataaat 6240
actccagcac aatgagcact tttgtagtgg agtgtagcga ggaaacgtat catcgctctg 6300
gtattgagtc tttatcagaa actgatgcaa aagcatttat tgctgatgta tttagtgttg 6360
agctagatgg tctgcctgtt gaggttcagc cgggcttgca atggcgaaat tttgtaacgc 6420
tgagccatac aaaagcttat gaagataatt tggtgctatt gggcgatgcc ttgcaaacag 6480
gccatttttc tatcgggcat ggcacaacaa tggcggtggt tgcggctcag atgttggtaa 6540
aagcattata tgagcatgca gaagtgaacc atgctttaga agattttaat caaaaggtga 6600
tgccgctgat gcaattattt agcgggcatg ccggcattag ccgcacttgg tttgaaacgg 6660
cggatgaaag aatggattta actgccgcag agttagcaaa gagtttttca gagcgccgcg 6720
agcaattgcc tcctttacct gctgctttgg ggcaagcgct gggtatggcg cttactcgta 6780
aggaatctta agatgattgc tgagaaaatg ttaccccctc gcttacctga gcaatggagc 6840
agtagttata tttcttactg gcagcctatg cagaaagaag atcaaattac gtcaggtatt 6900
tgctggtttg attatgaaca aaatcgctgc cgcattgatg gtttgtttaa cccttggtct 6960
gaggaaaaaa caggccatcg cttgtggatg tcagaaattg tgaccgcggg cgaggggaaa 7020
acaaagaaat ctaaaattgc gtattgccgt gaatcccctc ttggcgaagg ggaatatgag 7080
gcggtggtgc ttgatgatga tttggagtct tgccacgaag tgttattaac gcaggacgtt 7140
ttattgcaat atcaggcaag ctatgtggga agtcagcatt tgcttggcag agaaacggaa 7200
gcttggactt ttaccaaacc gggtaagggc ccttcaactt attactttgt aaaagggact 7260
aatcaattgc tgcgcatggt gacgggagac cctgcggtgc atgcttcaat tcgtgatttt 7320
cctaatttca ctactcatac gattccggcc gaaatatttt caagcagtat ttaa 7374
Claims (10)
1.菌株Janthinbacterium sp.B9-8,其特征在于,保藏编号为CCTCC NO:M2020183。
2.紫色杆菌素生物合成基因簇,其特征在于,选自如下1)或2):
1)包括五个紫色杆菌素生物合成途径基因vioA、vioB、vioC、vioD和vioE和一个核糖体结合位点SD序列,其核苷酸序列如SEQ ID NO:6所示,SD序列为5’-taatataagaaggagatata-3’;
2)与1)的基因簇具有80%以上同源性,且编码紫色杆菌素合成途径中的VioA、VioB、VioC、VioD和VioE五种酶的基因簇。
3.根据权利要求2所述的紫色杆菌素生物合成基因簇,其特征在于,所述紫色杆菌素生物合成基因簇来源于菌株Janthinbacterium sp.B9-8,保藏编号为CCTCC NO:M2020183。
4.根据权利要求2或3所述的紫色杆菌素生物合成基因簇,其特征在于,所述紫色杆菌素合成途径中的VioA、VioB、VioC、VioD和VioE五种酶的氨基酸序列依次如SEQ ID NO:1-5所示。
5.含有权利要求2或3所述基因簇的重组表达载体。
6.一种重组菌,将权利要求2或3所述基因簇或权利要求5所述重组表达载体导入宿主细胞获得。
7.含有权利要求2或3所述基因簇的表达盒或转基因细胞系。
8.权利要求2或3所述基因簇、权利要求5所述重组表达载体或权利要求6重组菌在生产紫色杆菌素中的应用。
9.一种生产紫色杆菌素的方法,其特征在于,包括以下步骤:
(1)将权利要求2或3所述基因簇或权利要求5所述重组表达载体导入宿主细胞大肠杆菌获得重组菌;
(2)步骤(1)所得重组菌发酵培养得到紫色杆菌素。
10.根据权利要求9所述的生产紫色杆菌素的方法,其特征在于,步骤(2)中,所述发酵培养培养基包括:蛋白胨3-10g/L、酵母粉1-10g/L、葡萄糖0.5-10g/L、硫酸铵0-5g/L,氨苄青霉素0-100mg/mL和卡那霉素0-100mg/mL;发酵体系培养温度为10-40℃,在加入诱导剂和L-色氨酸后,培养0-120h。
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