CN110896642A - 基于III型聚酮合酶的新型丙二酰-CoA生物传感器及其用途 - Google Patents
基于III型聚酮合酶的新型丙二酰-CoA生物传感器及其用途 Download PDFInfo
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Abstract
本发明涉及用于丙二酰‑CoA检测的重组微生物，在其中III型聚酮合酶编码基因插入基因组中，或在其中引入了含有所述基因的重组载体；使用所述重组微生物筛选丙二酰‑CoA产生诱导物质的方法；筛选参与增加丙二酰‑CoA产生的基因的方法；以及一种如下方法，其包括在微生物中敲低通过所述方法筛选的基因从而增加所述微生物中丙二酰‑CoA的产生，以及使用丙二酰‑CoA作为前体在所述微生物中产生有用的物质。根据本发明的生物传感器的使用提供了单步信号生成、在各种微生物中的利用、在自荧光微生物中的利用、简单的构建方法和简单的筛选方法。另外，当本发明与高通量筛选相结合时，其优点在于可以非常容易且快速地(约3天)筛选具有增加的丙二酰‑CoA产生能力的菌株，并且所述菌株可以直接应用于有用化合物的基于丙二酰‑CoA的产生。
Description
技术领域
本发明涉及基于III型聚酮合酶的新型丙二酰-CoA生物传感器及其用途，并且更具体地涉及用于丙二酰-CoA检测的重组微生物(其中引入了含有III型聚酮合酶编码基因的重组载体)；使用重组微生物筛选丙二酰-CoA产生诱导物质的方法；筛选参与增加丙二酰-CoA产生的基因的方法；以及一种如下方法，其包括在微生物中敲低通过所述方法筛选的基因从而增加微生物中丙二酰-CoA的产生，以及使用丙二酰-CoA作为前体在微生物中产生有用的物质。
背景技术
由于全球环境问题，有限资源的耗竭以及对环境友好型能源的需求增加，基于可再生生物的细胞工厂的建设越来越受到关注。这些细胞工厂可以通过针对产生所需的代谢物优化的细胞内代谢网络产生所需的代谢物(生物能、环境友好的化学物质、新药物物质等)，并且这一技术过程需要各种分子生物技术。
在可通过微生物细胞工厂产生的各种物质中，丙二酰-CoA尤为重要，因为它是产生非常有用的物质如聚酮、苯丙素类和生物燃料的关键前体。然而，丙二酰-CoA在细胞中引起各种必需的代谢反应(如脂肪酸生物合成)，因此可用于代谢工程化的丙二酰-CoA的量非常小。此外，需要高性能仪器如LC-MS/MS来测量丙二酰-CoA的细胞内浓度，并且需要一种花费很长时间并要求非常小心的采样方法来测量丙二酰-CoA，所述丙二酰-CoA在细胞中快速产生后消失并且对周围环境条件敏感。因此，已经从先前研究开发了基于转录因子的丙二酰-CoA生物传感器，以更快速且更容易地测量丙二酰-CoA的细胞内浓度(Xu P,Li L,ZhangF,Stephanopoulos G,Koffas M(2014),Proc Natl Acad Sci U S A111:11299-11304；LiS,Si T,Wang M,Zhao H(2015),ACS Synth Biol 4:1308-1315)。然而，如上所述基于转录因子的基于荧光蛋白的传感器的缺点在于它们必须经历各种信号传导步骤并且不能用于自荧光微生物如假单胞菌属(Pseudomonas)。
因此，本发明人努力解决上述问题，因此开发出一种基于III型聚酮合酶的新型丙二酰-CoA生物传感器，其利用III型聚酮合酶(包括RppA)通过一步反应将丙二酰-CoA转化为有色物质的特性；并且发现生物传感器的使用可以容易地检测丙二酰-CoA，使得可以筛选产生丙二酰-CoA的菌株、丙二酰-CoA产生诱导物质和参与增加丙二酰-CoA产生的基因，以及使得可以容易地构建一种使用丙二酰-CoA作为底物或前体来产生各种有用物质的方法，从而完成本发明。
发明内容
技术问题
本发明的目的是提供用于丙二酰-CoA检测的新型生物传感器和使用所述生物传感器的各种方法，以及通过使用所述方法增加所述微生物中丙二酰-CoA产生以及使用丙二酰-CoA作为底物或前体在所述微生物中产生有用物质的方法。
技术方案
为了达到上述目的，本发明提供了一种用于丙二酰-CoA检测的重组微生物，在其中III型聚酮合酶编码基因插入所述微生物的基因组中或在其中引入了含有III型聚酮合酶编码基因的重组载体。
本发明还提供了筛选丙二酰-CoA产生诱导物质的方法，其包括以下步骤：
(a)培养上述重组微生物；
(b)向所述重组微生物中添加候选物质；
(c)将添加所述候选物质后的所述重组微生物的培养上清液的颜色与没有添加所述候选物质的所述重组微生物的培养上清液的颜色进行比较；并且
(d)当添加所述候选物质后所述重组微生物的培养上清液显示出比没有添加所述候选物质的所述重组微生物的培养上清液更深的红色时，选择所述候选物质作为所述丙二酰-CoA产生诱导物质。
本发明还提供了筛选参与增加的丙二酰-CoA产生的基因的方法，所述方法包括以下步骤：
(a)将改变重组微生物中的基因表达的基因调节文库引入上述重组微生物中，从而构建所述重组微生物中的基因表达发生改变的重组微生物文库；
(b)培养所构建的重组微生物文库和未引入基因调节文库的重组微生物，从而测量所述培养上清液的吸光度，并比较所培养上清液的颜色；并且
(c)当所述重组微生物文库的培养上清液显示出比未引入所述基因调节文库的重组微生物的培养上清液更深的红色时，选择引入所述重组微生物文库中的基因。
本发明还提供了产生具有增加的丙二酰-CoA产生能力的重组微生物的方法，所述方法包括调节通过上述方法筛选的基因在固有地具有丙二酰-CoA产生能力或外部引入丙二酰-CoA产生能力的微生物中的表达。
本发明还提供了具有增加的丙二酰-CoA产生能力的重组微生物，其中选自以下基因的一种或多种基因：
fabF(3-氧代酰基-[酰基-载体-蛋白质]合酶II)；
yfcY(β-酮酰-CoA硫解酶)；
xapR(xapAB的转录激活因子)；
cytR(deo操纵子、udp、cdd、tsx、nupC和nupG的转录阻遏物)；
fabH(3-氧代酰基-[酰基-载体-蛋白质]合酶III)；
mqo(苹果酸脱氢酶)；
yfiD(丙酮酸甲酸裂解酶亚基)；
fmt(10-甲酰基四氢叶酸：L-甲硫氨酰基-tRNA(fMet)N-甲酰基转移酶)；
pyrF(乳清酸核苷-5'-磷酸脱羧酶)；
araA(L-阿拉伯糖异构酶)；
fadR(fad调节子的负调节因子和fabA的正调节因子)；
pabA(氨基脱氧分支酸合酶，亚基II)；
purB(腺苷酸琥珀酸裂解酶)；和
hycI(参与加工HycE的C末端的蛋白酶)在具有丙二酰-CoA产生能力的微生物中的表达与在野生型微生物中的表达相比降低。
本发明还提供了使用丙二酰-CoA作为底物或中间体产生有用物质的方法，所述方法包括以下步骤：
(a)构建重组微生物，其中另外引入参与所述有用物质产生的基因或增加所述基因的表达，或者另外使参与所述有用物质产生的基因缺失或减少所述基因的表达；
(b)培养所构建的微生物；并且
(c)从所培养的微生物回收所述有用物质。
本发明还提供了用于产生6-甲基水杨酸的重组微生物，其中在上述重组微生物中另外引入了编码6-甲基水杨酸合酶(6MSAS)和4'-磷酸泛酰巯基乙胺基转移酶(Sfp)的基因，或者增加所述基因的表达。
本发明还提供了产生6-甲基水杨酸的方法，所述方法包括以下步骤：
(a)培养上述重组微生物；并且
(b)从所培养的微生物回收所述6-甲基水杨酸。
本发明还提供了用于产生芦荟酮(aloesone)的重组微生物，其中在上述重组微生物中另外引入了编码芦荟酮合酶的基因或增加了所述基因的表达。
本发明还提供了产生芦荟酮的方法，所述方法包括以下步骤：
(a)培养上述重组微生物；并且
(b)从所培养的微生物回收所述芦荟酮。
本发明还提供了用于产生白藜芦醇的重组微生物，其中在上述重组微生物中另外引入了编码酪氨酸氨裂解酶(TAL)、4-香豆酸:CoA连接酶(4CL)和芪合酶(STS)的基因或者增加所述基因的表达。
本发明还提供了产生白藜芦醇的方法，所述方法包括以下步骤：
(a)培养上述重组微生物；并且
(b)从所培养的微生物回收所述白藜芦醇。
本发明还提供了用于产生柚皮素的重组微生物，其中在上述重组微生物中另外引入了编码酪氨酸氨裂解酶(TAL)、4-香豆酸:CoA连接酶(4CL)、查耳酮合酶(CHS)和查耳酮异构酶(CHI)的基因或者增加所述基因的表达。
本发明还提供了产生柚皮素的方法，所述方法包括以下步骤：
(a)培养上述重组微生物；并且
(b)从所培养的微生物回收所述柚皮素。
附图说明
图1示出了基于III型聚酮合酶(RppA)的丙二酰-CoA生物传感器的操作机制。(A)RppA可以将五分子的丙二酰-CoA转化为一分子的红色淡黄霉素。此时，RppA负责将丙二酰-CoA转化为1,3,6,8-四羟基萘(THN)，然后自发转化为淡黄霉素。(B)当通过RppA生物传感器测得红色变深时，丙二酰-CoA的细胞内水平更高。
图2示出了淡黄霉素生物合成。(A)来自五种不同菌株的通过rppA得到的淡黄霉素产量。(B)通过LC-MS和MS/MS分析确认的淡黄霉素产量。(C)淡黄霉素产量的优化。通过优化5'UTR增加淡黄霉素的产量。(D)检验RppA生物传感器的适用性。通过使用浅蓝菌素的间接方法测量由丙二酰-CoA的细胞内水平的变化引起的信号的变化。RppA+表示表达RppA的菌株，而RppA-表示不表达RppA的菌株。
图3示出了RppA生物传感器的适用性和可扩展性。(A)随着添加的浅蓝菌素的量增加，淡黄霉素产量增加，并且(B)这即使用肉眼也可以确认。(C)来自所有16种大肠杆菌(E.coli)菌株的淡黄霉素产量是可能的。(D)在表达RppA的菌株(RppA+)和不表达RppA的对照菌株(RppA-)的情况下，来自对照的340nm处的噪声(由箭头指示)最低并且来自RppA+菌株的信号最强，因此340nm处的吸光度用作RppA生物传感器的信号。
图4示出了作为丙二酰-CoA生物传感器的三种III型聚酮合酶(AaOKS、AaPKS4和AaPKS5)的表征。(A)通过LC-MS(负扫描模式)生成的表达AaOKS、AaPKS4和AaPKS5的大肠杆菌BL21(DE3)菌株的培养物上清液的提取离子色谱图。由色谱图预测的产物如下：5,7-二羟基-2-甲基色酮(m/z191)；芦荟酮(m/z 231)；SEK4和SEK4b(m/z 317)。(B)示出了当向培养基中添加不同浓度的浅蓝菌素时表达AaOKS、AaPKS4和AaPKS5的大肠杆菌菌株的培养上清液的颜色，并且(C)示出了不同波长处的培养上清液的光密度(吸光度)。CT，对照菌株(BL21(DE3)pET-30a(+))。此时，与对照相比，丙二酰-CoA生物传感器菌株在300nm处的信号最强，因此在AaOKS、AaPKS4和AaPKS5的情况下，使用300nm处的吸光度作为信号。数据点表示三个生物学重复的平均值。
图5指示三种III型聚酮合酶(AaOKS、AaPKS4和AaPKS5)可以用作丙二酰-CoA生物传感器。(A)AaOKS、(B)AaPKS4和(C)AaPKS5指示由丙二酰-CoA水平的增加(通过添加浅蓝菌素控制)引起的归一化信号增加。每个信号指示每个包含或不包含PKS的情况。
图6示出了在恶臭假单胞菌(Pseudomonas putida)、谷氨酸棒杆菌(Corynebacterium glutamicum)和混浊红球菌(Rhodococcus opacus)中测试RppA生物传感器的结果。(A)示出了当将指示的质粒转化至恶臭假单胞菌菌株中时淡黄霉素的产量。(B)示出了恶臭假单胞菌pBBR1-rppA传感器菌株的浅蓝菌素浓度依赖性信号强度，并且(C)示出了此时相对的淡黄霉素产量。(D)示出了当将指示的质粒转化至谷氨酸棒杆菌菌株中时淡黄霉素的产量。(E)示出了谷氨酸棒杆菌pCES-His-rppA传感器菌株的浅蓝菌素浓度依赖性信号强度，并且(F)示出了此时相对的淡黄霉素产量。(G)示出了当将指示的质粒转化至混浊红球菌菌株中时培养基的颜色。
图7示出了通过使用RppA生物传感器的高通量筛选来筛选菌株的过程，所述菌株显示增加的丙二酰-CoA产量。
图8示出了当引入大肠杆菌基因组规模的合成对照sRNA文库并且使用RppA生物传感器进行高通量筛选时筛选示出丙二酰-CoA水平增加的菌株的初始结果。
图9示出了当26个最初筛选的参与增加的丙二酰基-CoA产生的合成对照sRNA转化至大肠杆菌传感器菌株中时生成的信号。此时，选择与对照相比显示信号增加70％或更多的14种合成对照sRNA作为最终靶标。
图10示出了为鉴定增加的丙二酰-CoA产量而进行的FVSEOF模拟。(A)示出了与丙二酰-CoA生物合成相关的生物合成途径。粗体字母表示反应，而普通字母表示代谢物。灰色箭头表示代谢通量，而红色箭头表示被鉴定为过表达靶标的代谢通量。(B)示出了当过表达由FVSEOF鉴定的基因靶标时出现的生物传感器信号。(C)示出了由FVSEOF鉴定的基因靶标的列表。
图11示出了6-甲基水杨酸的生物合成途径。此处，红色X表示敲除基因，而蓝色X表示敲低基因。Bla，β-内酰胺酶基因；kanR，卡那霉素抗性基因；p15A，复制起点；ColE1，复制起点；Ptac，tac启动子；PBAD，阿拉伯糖-诱导型启动子；PR，PR启动子；rrnB，rrnBT1T2终止子；T1/TE，终止子。Gly，甘油；Gly-3P，3-磷酸甘油；DHA，二羟丙酮；DHAP，磷酸二羟丙酮；G3P，3-磷酸甘油醛；1,3BPG，1,3-二磷酸甘油酸；3PG，3-磷酸甘油酸；2PG，2-磷酸甘油酸；PEP，磷酸烯醇式丙酮酸；PYR，丙酮酸；OAA，草酰乙酸；E4P，D-赤藓糖4-磷酸；DAHP，3-脱氧-D-阿拉伯庚酮糖酸7-磷酸；CHOR，分支酸；AcCoA，乙酰-CoA；MalCoA，丙二酰-CoA；6MSAS，6MSA合酶；KS，酮基合酶；AT，酰基转移酶；DH，脱水酶；KR，酮还原酶；ACP，酰基载体蛋白。
图12示出了用于产生6-甲基水杨酸的不同基因表达盒。(A)示出了质粒pTac-Pg6MSAS-sfp，并且(B)示出了用质粒pTac-Pg6MSAS在大肠杆菌BAP1菌株的基因组上表达的sfp。(C)示出了为了分析6-甲基水杨酸合酶(Pg6MSAS)和4'-磷酸泛酰巯基乙胺基转移酶(Sfp)的表达而进行的SDS-PAGE的结果。Pg6MSAS(188kDa)，Sfp(26.1kDa)。
图13示出了来自经修饰的大肠杆菌菌株的6-甲基水杨酸的产量。(A)示出了在基于BL21的菌株和基于BAP1的菌株中来自不同浓度的葡萄糖/甘油的6-甲基水杨酸的产量。(B)示出了由大肠杆菌产生的6-甲基水杨酸的LC-MS分析的结果，并且(C)示出了可商购的6-甲基水杨酸化合物的LC-MS分析的结果。(D)示出了在16种大肠杆菌菌株中试管规模的6-甲基水杨酸产量。蓝色部分指示产量为1mg/L或更高。(E)示出了在大肠杆菌BL21(DE3)pTac-Pg6MSAS-sfp pWAS-抗-pabA(pabA敲低sRNA质粒)菌株中6-甲基水杨酸产量的时间历程。(F)示出了在BAP1 pTac-Pg6MSAS pWAS-抗-pabA菌株中6-甲基水杨酸产量的时间历程。(G)示出了相同菌株的补料分批发酵。红色箭头表示IPTG诱导的开始。在(E)、(F)和(G)中，蓝色的线和点表示细胞生长(OD600)，而红色的线和点表示6-甲基水杨酸浓度。
图14示出了当将实施例2.1中选择的14种合成对照sRNA引入6种筛选的大肠杆菌菌株中并对产生的菌株进行试管规模的培养时的6-甲基水杨酸产量。
图15的左图示出了当左图底部的基因在大肠杆菌BAP1 pTac-Pg6MSAS pWAS-抗-pabA(图12中示出最高的6-甲基水杨酸生产力的菌株)中过表达时烧瓶培养物中出现的6-甲基水杨酸的浓度。右图示出了大肠杆菌BAP1pTac-Pg6MSAS pWAS-抗-pabA pBBR1-accBCD1(示出最高生产力的菌株)的补料分批发酵的结果。
图16示出了芦荟酮生物合成途径。此处，红色X表示敲除基因，而蓝色X表示敲低基因。Glc，葡萄糖；G6P，葡萄糖6-磷酸；F6P，果糖6-磷酸；F1,6BP，果糖1,6-二磷酸；DHAP，磷酸二羟丙酮；G3P，3-磷酸甘油醛；1,3BPG，1,3-二磷酸甘油酸；3PG，3-磷酸甘油酸；2PG，2-磷酸甘油酸；PEP，磷酸烯醇式丙酮酸；PYR，丙酮酸；OAA，草酰乙酸；E4P，D-赤藓糖4-磷酸；DAHP，3-脱氧-D-阿拉伯庚酮糖酸7-磷酸；CHOR，分支酸；AcCoA，乙酰-CoA；MalCoA，丙二酰-CoA；ALS，芦荟酮合酶。bla，β-内酰胺酶基因；spcR，大观霉素-抗性基因；CDF，复制起点；ColE1，复制起点；PT7，T7启动子；PBAD，阿拉伯糖-诱导型启动子；PR，PR启动子；rrnB，rrnBT1T2终止子；T1/TE，终止子。
图17示出了经修饰的大肠杆菌菌株中的芦荟酮产量。(A)示出了为了分析芦荟酮合酶(RpALS，43kDa)和芦荟酮合酶(AaPKS3，44kDa)的表达而进行的SDS-PAGE的结果。(B)示出了当使用葡萄糖/甘油作为碳源表达RpALS/AaPKS3时芦荟酮的产量。(C)示出了在大肠杆菌中产生的芦荟酮的LC-MS和MS/MS谱。(D)示出了BL21(DE3)pCDF-RpALS菌株中芦荟酮产量的时间历程，并且(E)示出了BL21(DE3)pCDF-RpALS pWAS-抗-pabA菌株中芦荟酮产量的时间历程。在(D)和(E)中，蓝色的线和点描绘细胞生长(OD600)，而红色的线和点描绘芦荟酮浓度。(F)示出了通过将实施例2.1中选择的14种合成对照sRNA引入两种类型的大肠杆菌菌株中而产生的菌株的试管规模培养的结果。
图18示出了当图底部的基因在BL21(DE3)pCDF-RpALS pWAS-抗-pabA(图17(E)中示出最高的芦荟酮生产力的菌株)中过表达时烧瓶培养物中出现的芦荟酮的浓度。
图19示出了白藜芦醇生物合成途径。此处，红色X表示敲除基因，而蓝色X表示敲低基因。黑色箭头以及用粗体字母书写的基因表示过表达的代谢通量。红色虚线(以及圆圈中的符号-)指示转录抑制。黑色虚线(以及圆圈中的符号+)指示转录激活。Gly，甘油；Gly-3P，3-磷酸甘油；DHA，二羟丙酮；DHAP，磷酸二羟丙酮；G3P，3-磷酸甘油醛；1,3BPG，1,3-二磷酸甘油酸；3PG，3-磷酸甘油酸；2PG，2-磷酸甘油酸；PEP，磷酸烯醇式丙酮酸；PYR，丙酮酸；OAA，草酰乙酸；E4P，D-赤藓糖4-磷酸；DAHP，3-脱氧-D-阿拉伯庚酮糖酸7-磷酸；SHIK，莽草酸；CHOR，分支酸；PPHN，预苯酸；HPP，4-羟基苯丙酮酸；FOR，甲酸；AcCoA，乙酰-CoA；MalCoA，丙二酰-CoA。bla，β-内酰胺酶基因；kanR，卡那霉素-抗性基因；spcR，大观霉素-抗性基因；p15A，复制起点；ColE1，复制起点；CDF，复制起点；Ptac，tac启动子；PBAD，阿拉伯糖-诱导型启动子；PR，PR启动子；Ptrc，trc启动子；rrnB，rrnBT1T2终止子；T1/TE，终止子。
图20示出了通过经修饰的大肠杆菌菌株得到的对香豆酸和白藜芦醇的产量。(A)示出了通过各种经构建用于检验N末端蛋白质标签的作用的质粒得到的对香豆酸的产量。将除了pTY13-HisTAL以外的每一种载体转化至BTY5.13菌株中，并将pTY13-HisTAL转化至BTY5菌株中。(B)示出了为了分析每种蛋白质表达而进行的SDS-PAGE的结果。At4CL1m(61.1kDa)，At4CL3(61.3kDa)，At4CL4(62.6kDa)，Sc4CLm(55.3kDa)，STS(42.8kDa)。CT描绘了对照。M描绘了蛋白质大小标记物。(C)示出了通过STS和不同4CL的组合得到的白藜芦醇产量的结果。(D)示出了通过不同质粒构建策略得到的白藜芦醇产量。此处，1至5描述了质粒，并且每种质粒如下。1，pTac-VvSTS-At4CL1m；2，pTac-At4CL1m-opr-VvSTS(表达操纵子中的两个基因)；3，pTac-At4CL1m-fus-VvSTS(表达At4CL1m和STS的融合蛋白)；4和5，pTacCDF-VvSTS-At4CL1m。此处，将对应于1至4的质粒引入大肠杆菌BL21(DE3)中并将其在2mM对香豆酸和20g/L甘油的存在下培养。将对应于5的质粒引入BTY5 pTY13-HisTAL中，并将其在20g/L甘油的存在下培养。对应于5的菌株用作产生白藜芦醇的碱性菌株，并通过较深的蓝色条指示。(E)示出了通过将实施例2.1中选择的14种合成对照sRNA转化至产生白藜芦醇的碱性菌株(BTY5 pTY13-HisTAL pTacCDF-VvSTS-At4CL1m)中，接着进行烧瓶培养获得的白藜芦醇的产量。与对照相比，展示为较深的红色图的对应于六种菌株的sRNA示出白藜芦醇产量的2.5倍增加，因此将这六种sRNA用于组合双敲低测试中。(F)示出了使用先前选择的六种sRNA的组合的组合双敲低测试的结果。*P<0.05，**P<0.01，***P<0.001(双尾学生t检验)。
图21示出了为了确定由经修饰的大肠杆菌产生的白藜芦醇的真实性而进行的LC-MS的结果。(A)示出了由经修饰的大肠杆菌产生的白藜芦醇的LC-MS谱，并且(B)示出了可商购的白藜芦醇化合物的LC-MS谱。
图22示出了柚皮素生物合成途径。此处，红色X表示敲除基因，而蓝色X表示敲低基因。黑色箭头以及用粗体字母书写的基因表示过表达的代谢通量。红色虚线(以及圆圈中的符号-)指示转录抑制。黑色虚线(以及圆圈中的符号+)指示转录激活。bla，β-内酰胺酶基因；kanR，卡那霉素-抗性基因；spcR，大观霉素-抗性基因；p15A，复制起点；ColE1，复制起点；CDF，复制起点；Ptac，tac启动子；PBAD，阿拉伯糖-诱导型启动子；PR，PR启动子；Ptrc，trc启动子；rrnB，rrnBT1T2终止子；T1/TE，终止子。Gly，甘油；Gly-3P，3-磷酸甘油；DHA，二羟丙酮；DHAP，磷酸二羟丙酮；G3P，3-磷酸甘油醛；1,3BPG，1,3-二磷酸甘油酸；3PG，3-磷酸甘油酸；2PG，2-磷酸甘油酸；PEP，磷酸烯醇式丙酮酸；PYR，丙酮酸；OAA，草酰乙酸；E4P，D-赤藓糖4-磷酸；DAHP，3-脱氧-D-阿拉伯庚酮糖酸7-磷酸；SHIK，莽草酸；CHOR，分支酸；PPHN，预苯酸；HPP，4-羟基苯丙酮酸；FOR，甲酸；AcCoA，乙酰-CoA；MalCoA，丙二酰-CoA；TYR，L-酪氨酸；COU，对香豆酸；CouCoA，对香豆酰-CoA。
图23示出了经修饰的大肠杆菌中的柚皮素产量。(A)示出了含有pTrcCDF-At4CL1m-AtCHI-PhCHS质粒的菌株中的柚皮素产量。此处，在具有对香豆酸途径的菌株的情况下，将载体转化至BTY5 pTY13-HisTAL菌株中，而在其他情况下，将载体转化至BL21(DE3)菌株中，然后将2mM对香豆酸添加至培养基中。添加20g/L的葡萄糖或甘油。(B)示出了异源酶的SDS-PAGE分析。At4CL1m，PhCHS(42.6kDa)，AtCHI(26.6kDa)。
图24示出了为了确定由经修饰的大肠杆菌产生的柚皮素的真实性而进行的LC-MS的结果。(A)示出了由经修饰的大肠杆菌产生的柚皮素的LC-MS谱，并且(B)示出了可商购的柚皮素化合物的LC-MS谱。
图25示出了经修饰的大肠杆菌中的柚皮素产量。(A)示出了将实施例2.1中选择的14种合成对照sRNA转化至碱性柚皮素菌株(BTY5 pTY13-HisTAL pTrcCDF-At4CL1m-AtCHI-PhCHS)中，然后进行烧瓶培养而产生的柚皮素的量。与对照相比，展示为较深的红色条的对应于三种菌株的sRNA示出柚皮素产量增加15％或更多，因此将这三种sRNA用于组合双敲低测试。(B)示出了使用先前选择的三种sRNA的组合的组合双敲低测试的结果。*P<0.05，**P<0.01(双尾学生t检验)。
具体实施方式
除非另外定义，否则本文中使用的所有技术术语和科学术语具有与本发明所属领域的普通技术人员通常所理解的那些相同的含义。通常，本文使用的命名法和下文将描述的实验方法是本领域公知且常用的方法。
在本发明中，已发现与常规的测量丙二酰-CoA浓度的方法相比，使用引入III型聚酮合酶的重组微生物使得能够以在时间、成本、便利性等方面显著改善的方式来测量丙二酰-CoA浓度。
因此，在一方面，本发明涉及用于丙二酰-CoA检测的重组微生物，在其中III型聚酮合酶编码基因插入所述微生物的基因组中或在其中引入了含有III型聚酮合酶编码基因的重组载体。
在本发明中，III型聚酮合酶可以是：
源自选自灰色链霉菌(Streptomyces griseus)、天蓝色链霉菌(Streptomycescoelicolor)、阿维链霉菌(Streptomyces avermitilis)、红色糖多孢菌(Saccharopolyspora erythraea)、波赛链霉菌(Streptomyces peucetius)和针孢链霉菌(Streptomyces aculeolatus)的微生物的RppA；
源自荧光假单胞菌(Pseudomonas fluorescens)的PhlD(聚酮合酶)；
源自地中海拟无枝菌酸菌的DpgA(聚酮合酶)；
源自掌叶大黄的ALS(芦荟酮合酶)；或
源自木立芦荟(Aloe arborescens)的PCS(5,7-二羟基-2-甲基色酮合酶)、OKS(八酮合酶)、PKS3(芦荟酮合酶)、PKS4(八酮合酶2)或PKS5(八酮合酶3)，但不限于这些。
源自上述六种放线菌(Actinomycetes)菌株的III型聚酮合酶RppA使用丙二酰-CoA作为底物产生淡黄霉素作为产物。此外，其他III型聚酮合酶也使用丙二酰-CoA作为底物，但PhlD产生间苯三酚作为产物，DpgA产生二羟基苯乙酸酯作为产物，PCS产生5,7-二羟基-2-甲基色酮作为产物，并且OKS、PKS4或PKS5产生SEK4和SEK4b作为产物。ALS和PKS3产生芦荟酮作为产物。
用于本发明的每种RppA酶的氨基酸序列如下：
灰色链霉菌RppA[SEQ ID NO:93]：
天蓝色链霉菌RppA[SEQ ID NO:94]：
阿维链霉菌RppA[SEQ ID NO:95]：
红色糖多孢菌RppA[SEQ ID NO:96]：
波赛链霉菌RppA[SEQ ID NO:97]：
针孢链霉菌RppA[SEQ ID NO:98]：
用于本发明的其他III型聚酮合酶的氨基酸序列如下：
荧光假单胞菌III型聚酮合酶(PhlD)[SEQ ID NO:99]：
东方拟无枝菌酸菌(Amycolatopsis orientalis)III型聚酮合酶(DpgA)[SEQ IDNO:100]：
掌叶大黄(Rheum palmatum)III型聚酮合酶(芦荟酮合酶，ALS)[SEQ ID NO:101]：
木立芦荟(Aloe arborescens)III型聚酮合酶(5,7-二羟基-2-甲基色酮合酶，PCS)[SEQ ID NO:102]：
木立芦荟III型聚酮合酶(八酮合酶，OKS)[SEQ ID NO:103]：
木立芦荟III型聚酮合酶(芦荟酮合酶，PKS3)[SEQ ID NO:104]：
木立芦荟III型聚酮合酶(八酮合酶2，PKS4)[SEQ ID NO:105]：
木立芦荟III型聚酮合酶(八酮合酶3，PKS5)[SEQ ID NO:106]：
上述III型聚酮合酶的氨基酸序列示出了一些可用于构建根据本发明的用于丙二酰-CoA检测的重组微生物的酶。本发明的技术特征在于可通过引入III型聚酮合酶进行丙二酰-CoA检测。因此，对于本领域技术人员明显的是，本发明使得可以构建引入除上述III型聚酮合酶之外的III型聚酮合酶的重组微生物。
在本发明中，重组微生物的特征可以在于编码III型聚酮合酶的基因与选自tac、trc、T7、BAD、λPR和Anderson合成启动子(但不限于这些)的启动子可操作地连接。
在本发明中，重组微生物可以选自由以下项组成的菌株组：大肠杆菌、根瘤菌属(Rhizobium)、双歧杆菌属(Bifidobacterium)、红球菌属(Rhodococcus)、假丝酵母属(Candida)、欧文氏菌属(Erwinia)、肠杆菌属(Enterobacter)、巴氏杆菌属(Pasteurella)、曼氏杆菌属(Mannheimia)、放线杆菌属(Actinobacillus)、凝聚杆菌属(Aggregatibacter)、黄单胞菌属(Xanthomonas)、弧菌属(Vibrio)、假单胞菌属(Pseudomonas)、固氮菌属(Azotobacter)、不动杆菌属(Acinetobacter)、劳尔氏菌属(Ralstonia)、土壤杆菌属(Agrobacterium)、红杆菌属(Rhodobacter)、发酵单胞菌属(Zymomonas)、芽孢杆菌属(Bacillus)、葡萄球菌属(Staphylococcus)、乳球菌属(Lactococcus)、链球菌属(Streptococcus)、乳杆菌属(Lactobacillus)、梭菌属(Clostridium)、棒杆菌属(Corynebacterium)、链霉菌属(Streptomyces)、双歧杆菌属(Bifidobacterium)、蓝细菌(Cyanobacterium)和圆杆菌属(Cyclobacterium)，但应用本发明的微生物不限于这些。优选地，本发明的重组微生物可以通过将重组载体引入选自大肠杆菌、假单胞菌属物种、棒杆菌属物种和红球菌属物种(但不限于这些)的微生物中来构建。作为大肠杆菌，优选使用选自下表2中所示的大肠杆菌菌株中的一种，但是本领域技术人员将理解，也可以使用预期表现出类似效果的其他大肠杆菌菌株。本发明的重组微生物可以通过使用作为假单胞菌属物种的恶臭假单胞菌、作为棒杆菌属物种的谷氨酸棒杆菌和作为红球菌属物种的混浊红球菌来构建，但这是代表性微生物的例子，并且应用本发明的微生物不限于这些。
同时，在本发明中，当使用用于丙二酰-CoA检测的重组微生物(在其中III型聚酮合酶编码基因插入微生物的基因组中或在其中引入了含有III型聚酮合酶编码基因的重组载体)时，可以通过来自丙二酰-CoA的浓度依赖性显色容易地确认丙二酰-CoA的产量，并且可以容易地筛选丙二酰-CoA产生诱导物质。
因此，在另一方面，本发明涉及筛选丙二酰-CoA产生诱导物质的方法，其包括以下步骤：
(a)培养所述重组微生物；
(b)向所述重组微生物中添加候选物质；
(c)将添加所述候选物质后的所述重组微生物的培养上清液的颜色与没有添加所述候选物质的所述重组微生物的培养上清液的颜色进行比较；并且
(d)当添加所述候选物质后所述重组微生物的培养上清液显示出比没有添加所述候选物质的所述重组微生物的培养上清液更深的红色时，选择所述候选物质作为所述丙二酰-CoA产生诱导物质。
在本发明中，步骤(c)可以包括通过肉眼清楚地观察颜色的变化，或者测量吸光度并定量地表示吸光度的变化。
因此，步骤(c)中颜色之间的比较可以通过肉眼进行。
另外，步骤(c)中颜色之间的比较可以通过测量吸光度来进行，并且步骤(d)可以包括当添加所述候选物质后测量的吸光度高于未添加所述候选物质的情况下测量的吸光度时，选择所述候选物质作为所述丙二酰-CoA产生诱导物质。
在本发明中，可以测量280至450nm、优选300至340nm的吸光度(OD值)，并且OD值随着丙二酰-CoA浓度的增加而增加。
丙二酰-CoA产生诱导物质可以通过如下机制诱导丙二酰-CoA的产生，所述机制调节参与增加丙二酰-CoA产生的基因的表达。
因此，在再另一方面，本发明涉及筛选参与增加丙二酰基-CoA产生的基因的方法，所述方法包括以下步骤：
(a)将改变重组微生物中的基因表达的基因调节文库引入上述重组微生物中，从而构建所述重组微生物中的基因表达发生改变的重组微生物文库；
(b)培养所述构建的重组微生物文库，测量所述重组微生物文库的培养上清液的吸光度，培养所述重组微生物然后将所述基因调节文库引入，并比较所述重组微生物文库的培养上清液的颜色与在将所述基因调节文库引入之前所述重组微生物的培养上清液的颜色；并且
(c)当所述重组微生物文库的培养上清液显示出比在将所述基因调节文库引入之前重组微生物的培养上清液更深的红色时，选择引入所述重组微生物文库中的基因。
在本发明中，步骤(b)可以包括通过肉眼清楚地观察颜色的变化，或者测量吸光度并定量地表示吸光度的变化。
因此，步骤(c)中颜色的比较可以通过肉眼进行。
另外，步骤(b)中的颜色之间的比较是通过测量吸光度来进行，并且步骤(c)可以包括当所述重组微生物文库的培养上清液的吸光度高于在将所述基因调节文库引入之前所述重组微生物的培养上清液的吸光度时，选择在所述重组微生物文库中引入的基因作为参与增加丙二酰-CoA产生的基因。
在本发明中，可以测量280至450nm、优选300至340nm的吸光度(OD值)，并且OD值随着丙二酰-CoA浓度的增加而增加。
在本发明中，基因调节文库可以是选自以下的文库：sRNA文库、基因组文库、cDNA文库、gRNA文库和用于构建敲除或突变菌株的寡核苷酸文库，但不限于这些。
具体地，为了达到本发明的目的，基因调节文库可以适当地选自以下：
sRNA文库或用于构建敲除或突变菌株以抑制重组微生物中的内源基因表达的寡核苷酸文库；
作为针对内源或外源基因过表达的文库的基因组文库或cDNA文库；和
gRNA(指导RNA，其用于CRISPR(规律间隔成簇短回文重复序列)-Cas9技术)文库，它是能够实现基因表达抑制和基因过表达二者的文库。
也就是说，以下实施例2中描述的合成对照sRNA文库的添加是用于描述本发明的例子，并且呈现了使用所述合成对照sRNA文库的方法，并且可以引入表现出类似功能的任何其他文库。
同时，当微生物中通过所述方法筛选的基因被敲低时，可以增加微生物中丙二酰-CoA产量。因此，在又另一个方面，本发明涉及产生具有增加的丙二酰-CoA产生能力的重组微生物的方法，所述方法包括调节通过所述方法筛选的基因在具有丙二酰-CoA产生能力的微生物中的表达。
具有丙二酰-CoA产生能力的微生物意指固有地具有丙二酰-CoA产生能力或外部引入丙二酰-CoA产生能力的微生物。
在本发明中，筛选的基因可以是选自以下项的一种或多种基因：
fabF(3-氧代酰基-[酰基-载体-蛋白质]合酶II)；
yfcY(β-酮酰-CoA硫解酶)；
xapR(xapAB的转录激活因子)；
cytR(deo操纵子、udp、cdd、tsx、nupC和nupG的转录阻遏物)；
fabH(3-氧代酰基-[酰基-载体-蛋白质]合酶III)；
mqo(苹果酸脱氢酶)；
yfiD(丙酮酸甲酸裂解酶亚基)；
fmt(10-甲酰基四氢叶酸：L-甲硫氨酰基-tRNA(fMet)N-甲酰基转移酶)；
pyrF(乳清酸核苷-5'-磷酸脱羧酶)；
araA(L-阿拉伯糖异构酶)；
fadR(fad调节子的负调节因子和fabA的正调节因子)；
pabA(氨基脱氧分支酸合酶，亚基II)；
purB(腺苷酸琥珀酸裂解酶)；和
hycI(参与加工HycE的C末端的蛋白酶)，
并且可以减少所述基因的表达，但所述筛选的基因不限于这些。
在本发明中，对应地，fabF编码3-氧代酰基-[酰基-载体-蛋白质]合酶II，yfcY编码β-酮酰-CoA硫解酶，xapR编码xapAB的转录激活因子，cytR编码deo操纵子、udp、cdd、tsx、nupC和nupG的转录阻遏物，fabH编码3-氧代酰基-[酰基-载体-蛋白质]合酶III，mqo编码苹果酸脱氢酶，yfiD编码丙酮酸甲酸裂解酶亚基，fmt编码10-甲酰基四氢叶酸:L-甲硫氨酰基-tRNA(fMet)N-甲酰基转移酶，pyrF编码乳清酸核苷-5'-磷酸脱羧酶，araA编码L-阿拉伯糖异构酶，fadR编码fad调节子的负调节因子和fabA的正调节因子，pabA编码氨基脱氧分支酸合酶亚基II，purB编码腺苷酸琥珀酸裂解酶，并且hycI编码参与加工HycE的C末端的蛋白酶。
被筛选并用于本发明的每种酶的氨基酸序列如下，但可以在氨基酸序列中具有一个或多个氨基酸的取代、缺失或添加，并且在这种情况下，得到的突变氨基酸序列可以表现出与本发明相同或更好的功能。例如，明显的是，如果突变氨基酸序列的结构构型与野生型氨基酸序列相同或类似，并因此对底物表现出相同或更高的作用，则突变氨基酸序列也落入本发明的范围内。在相同的上下文中，明显的是，除了本发明中使用的酶之外，具有相同或类似功能的其他微生物来源的酶可以适当地应用于本发明，并且相应酶的在可由本领域普通技术人员容易地应用的范围内的应用落入本发明的范围内。
FabF[SEQ ID NO:107]：
MSKRRVVVTGLGMLSPVGNTVESTWKALLAGQSGISLIDHFDTSAYATKFAGLVKDFNCEDIISRKEQRKMDAFIQYGIVAGVQAMQDSGLEITEENATRIGAAIGSGIGGLGLIEENHTSLMNGGPRKISPFFVPSTIVNMVAGHLTIMYGLRGPSISIATACTSGVHNIGHAARIIAYGDADVMVAGGAEKASTPLGVGGFGAARALSTRNDNPQAASRPWDKERDGFVLGDGAGMLVLEEYEHAKKRGAKIYAELVGFGMSSDAYHMTSPPENGAGAALAMANALRDAGIEASQIGYVNAHGTSTPAGDKAEAQAVKTIFGEAASRVLVSSTKSMTGHLLGAAGAVESIYSILALRDQAVPPTINLDNPDEGCDLDFVPHEARQVSGMEYTLCNSFGFGGTNGSLIFKKI
YfcY[SEQ ID NO:108]：
MGQVLPLVTRQGDRIAIVSGLRTPFARQATAFHGIPAVDLGKMVVGELLARSEIPAEVIEQLVFGQVVQMPEAPNIAREIVLGTGMNVHTDAYSVSRACATSFQAVANVAESLMAGTIRAGIAGGADSSSVLPIGVSKKLARVLVDVNKARTMSQRLKLFSRLRLRDLMPVPPAVAEYSTGLRMGDTAEQMAKTYGITREQQDALAHRSHQRAAQAWSDGKLKEEVMTAFIPPYKQPLVEDNNIRGNSSLADYAKLRPAFDRKHGTVTAANSTPLTDGAAAVILMTESRAKELGLVPLGYLRSYAFTAIDVWQDMLLGPAWSTPLALERAGLTMSDLTLIDMHEAFAAQTLANIQLLGSERFAREALGRAHATGEVDDSKFNVLGGSIAYGHPFAATGARMITQTLHELRRRGGGFGLVTACAAGGLGAAMVLEAE
XapR[SEQ ID NO:109]：
MERVYRTDLKLLRYFLAVAEELHFGRAAARLNMSQPPLSIHIKELENQLGTQLFIRHSRSVVLTHAGKILMEESRRLLVNANNVLARIEQIGRGEAGRIELGVVGTAMWGRMRPVMRRFLRENPNVDVLFREKMPAMQMALLERRELDAGIWRMATEPPTGFTSLRLHESAFLVAMPEEHHLSSFSTVPLEALRDEYFVTMPPVYTDWDFLQRVCQQVGFSPVVIREVNEPQTVLAMVSMGIGITLIADSYAQMNWPGVIFRPLKQRIPADLYIVYETQQVTPAMVKLLAALTQ
CytR[SEQ ID NO:110]：
MKAKKQETAATMKDVALKAKVSTATVSRALMNPDKVSQATRNRVEKAAREVGYLPQPMGRNVKRNESRTILVIVPDICDPFFSEIIRGIEVTAANHGYLVLIGDCAHQNQQEKTFIDLIITKQIDGMLLLGSRLPFDASIEEQRNLPPMVMANEFAPELELPTVHIDNLTAAFDAVNYLYEQGHKRIGCIAGPEEMPLCHYRLQGYVQALRRCGIMVDPQYIARGDFTFEAGSKAMQQLLDLPQPPTAVFCHSDVMALGALSQAKRQGLKVPEDLSIIGFDNIDLTQFCDPPLTTIAQPRYEIGREAMLLLLDQMQGQHVGSGSRLMDCELIIRGSTRALP
FabH[SEQ ID NO:111]：
MYTKIIGTGSYLPEQVRTNADLEKMVDTSDEWIVTRTGIRERHIAAPNETVSTMGFEAATRAIEMAGIEKDQIGLIVVATTSATHAFPSAACQIQSMLGIKGCPAFDVAAACAGFTYALSVADQYVKSGAVKYALVVGSDVLARTCDPTDRGTIIIFGDGAGAAVLAASEEPGIISTHLHADGSYGELLTLPNADRVNPENSIHLTMAGNEVFKVAVTELAHIVDETLAANNLDRSQLDWLVPHQANLRIISATAKKLGMSMDNVVVTLDRHGNTSAASVPCALDEAVRDGRIKPGQLVLLEAFGGGFTWGSALVRF
Mqo[SEQ ID NO:112]：
MKKVTAMLFSMAVGLNAVSMAAKAKASEEQETDVLLIGGGIMSATLGTYLRELEPEWSMTMVERLEGVAQESSNGWNNAGTGHSALMELNYTPQNADGSISIEKAVAINEAFQISRQFWAHQVERGVLRTPRSFINTVPHMSFVWGEDNVNFLRARYAALQQSSLFRGMRYSEDHAQIKEWAPLVMEGRDPQQKVAATRTEIGTDVNYGEITRQLIASLQKKSNFSLQLSSEVRALKRNDDNTWTVTVADLKNGTAQNIRAKFVFIGAGGAALKLLQESGIPEAKDYAGFPVGGQFLVSENPDVVNHHLAKVYGKASVGAPPMSVPHIDTRVLDGKRVVLFGPFATFSTKFLKNGSLWDLMSSTTTSNVMPMMHVGLDNFDLVKYLVSQVMLSEEDRFEALKEYYPQAKKEDWRLWQAGQRVQIIKRDAEKGGVLRLGTEVVSDQQGTIAALLGASPGASTAAPIMLNLLEKVFGDRVSSPQWQATLKAIVPSYGRKLNGDVAATERELQYTSEVLGLNYDKPQAADSTPKPQLKPQPVQKEVADIAL
YfiD[SEQ ID NO:113]：
MITGIQITKAANDDLLNSFWLLDSEKGEARCIVAKAGYAEDEVVAVSKLGDIEYREVPVEVKPEVRVEGGQHLNVNVLRRETLEDAVKHPEKYPQLTIRVSGYAVRFNSLTPEQQRDVIARTFTESL
Fmt[SEQ ID NO:114]：
MSESLRIIFAGTPDFAARHLDALLSSGHNVVGVFTQPDRPAGRGKKLMPSPVKVLAEEKGLPVFQPVSLRPQENQQLVAELQADVMVVVAYGLILPKAVLEMPRLGCINVHGSLLPRWRGAAPIQRSLWAGDAETGVTIMQMDVGLDTGDMLYKLSCPITAEDTSGTLYDKLAELGPQGLITTLKQLADGTAKPEVQDETLVTYAEKLSKEEARIDWSLSAAQLERCIRAFNPWPMSWLEIEGQPVKVWKASVIDTATNAAPGTILEANKQGIQVATGDGILNLLSLQPAGKKAMSAQDLLNSRREWFVPGNRLV
PyrF[SEQ ID NO:115]：
MTLTASSSSRAVTNSPVVVALDYHNRDDALAFVDKIDPRDCRLKVGKEMFTLFGPQFVRELQQRGFDIFLDLKFHDIPNTAAHAVAAAADLGVWMVNVHASGGARMMTAAREALVPFGKDAPLLIAVTVLTSMEASDLVDLGMTLSPADYAERLAALTQKCGLDGVVCSAQEAVRFKQVFGQEFKLVTPGIRPQGSEAGDQRRIMTPEQALSAGVDYMVIGRPVTQSVDPAQTLKAINASLQRSA
AraA[SEQ ID NO:116]：
MTIFDNYEVWFVIGSQHLYGPETLRQVTQHAEHVVNALNTEAKLPCKLVLKPLGTTPDEITAICRDANYDDRCAGLVVWLHTFSPAKMWINGLTMLNKPLLQFHTQFNAALPWDSIDMDFMNLNQTAHGGREFGFIGARMRQQHAVVTGHWQDKQAHERIGSWMRQAVSKQDTRHLKVCRFGDNMREVAVTDGDKVAAQIKFGFSVNTWAVGDLVQVVNSISDGDVNALVDEYESCYTMTPATQIHGKKRQNVLEAARIELGMKRFLEQGGFHAFTTTFEDLHGLKQLPGLAVQRLMQQGYGFAGEGDWKTAALLRIMKVMSTGLQGGTSFMEDYTYHFEKGNDLVLGSHMLEVCPSIAAEEKPILDVQHLGIGGKDDPARLIFNTQTGPAIVASLIDLGDRYRLLVNCIDTVKTPHSLPKLPVANALWKAQPDLPTASEAWILAGGAHHTVFSHALNLNDMRQFAEMHDIEITVIDNDTRLPAFKDALRWNEVYYGFRR
ArgC[SEQ ID NO:117]：
MLNTLIVGASGYAGAELVTYVNRHPHMNITALTVSAQSNDAGKLISDLHPQLKGIVDLPLQPMSDISEFSPGVDVVFLATAHEVSHDLAPQFLEAGCVVFDLSGAFRVNDATFYEKYYGFTHQYPELLEQAAYGLAEWCGNKLKEANLIAVPGCYPTAAQLALKPLIDADLLDLNQWPVINATSGVSGAGRKAAISNSFCEVSLQPYGVFTHRHQPEIATHLGADVIFTPHLGNFPRGILETITCRLKSGVTQAQVAQVLQQAYAHKPLVRLYDKGVPALKNVVGLPFCDIGFAVQGEHLIIVATEDNLLKGAAAQAVQCANIRFGYAETQSLI
FadR[SEQ ID NO:118]：
MVIKAQSPAGFAEEYIIESIWNNRFPPGTILPAERELSELIGVTRTTLREVLQRLARDGWLTIQHGKPTKVNNFWETSGLNILETLARLDHESVPQLIDNLLSVRTNISTIFIRTAFRQHPDKAQEVLATANEVADHADAFAELDYNIFRGLAFASGNPIYGLILNGMKGLYTRIGRHYFANPEARSLALGFYHKLSALCSEGAHDQVYETVRRYGHESGEIWHRMQKNLPGDLAIQGR
NudD[SEQ ID NO:119]：
MFLRQEDFATVVRSTPLVSLDFIVENSRGEFLLGKRTNRPAQGYWFVPGGRVQKDETLEAAFERLTMAELGLRLPITAGQFYGVWQHFYDDNFSGTDFTTHYVVLGFRFRVSEEELLLPDEQHDDYRWLTSDALLASDNVHANSRAYFLAEKRTGVPGL
PabA[SEQ ID NO:120]：
MILLIDNYDSFTWNLYQYFCELGADVLVKRNDALTLADIDALKPQKIVISPGPCTPDEAGISLDVIRHYAGRLPILGVCLGHQAMAQAFGGKVVRAAKVMHGKTSPITHNGEGVFRGLANPLTVTRYHSLVVEPDSLPACFDVTAWSETREIMGIRHRQWDLEGVQFHPESILSEQGHQLLANFLHR
PurB[SEQ ID NO:121]：
MELSSLTAVSPVDGRYGDKVSALRGIFSEYGLLKFRVQVEVRWLQKLAAHAAIKEVPAFAADAIGYLDAIVASFSEEDAARIKTIERTTNHDVKAVEYFLKEKVAEIPELHAVSEFIHFACTSEDINNLSHALMLKTARDEVILPYWRQLIDGIKDLAVQYRDIPLLSRTHGQPATPSTIGKEMANVAYRMERQYRQLNQVEILGKINGAVGNYNAHIAAYPEVDWHQFSEEFVTSLGIQWNPYTTQIEPHDYIAELFDCVARFNTILIDFDRDVWGYIALNHFKQKTIAGEIGSSTMPHKVNPIDFENSEGNLGLSNAVLQHLASKLPVSRWQRDLTDSTVLRNLGVGIGYALIAYQSTLKGVSKLEVNRDHLLDELDHNWEVLAEPIQTVMRRYGIEKPYEKLKELTRGKRVDAEGMKQFIDGLALPEEEKARLKAMTPANYIGRAITMVDELK
HycI[SEQ ID NO:122]：
MTDVLLCVGNSMMGDDGAGPLLAEKCAAAPKGNWVVIDGGSAPENDIVAIRELRPTRLLIVDATDMGLNPGEIRIIDPDDIAEMFMMTTHNMPLNYLIDQLKEDIGEVIFLGIQPDIVGFYYPMTQPIKDAVETVYQRLEGWEGNGGFAQLAVEEE
在本发明中，重组微生物可以选自大肠杆菌、根瘤菌属(Rhizobium)、双歧杆菌属(Bifidobacterium)、红球菌属(Rhodococcus)、假丝酵母属(Candida)、欧文氏菌属(Erwinia)、肠杆菌属(Enterobacter)、巴氏杆菌属(Pasteurella)、曼氏杆菌属(Mannheimia)、放线杆菌属(Actinobacillus)、凝聚杆菌属(Aggregatibacter)、黄单胞菌属(Xanthomonas)、弧菌属(Vibrio)、假单胞菌属(Pseudomonas)、固氮菌属(Azotobacter)、不动杆菌属(Acinetobacter)、劳尔氏菌属(Ralstonia)、土壤杆菌属(Agrobacterium)、红杆菌属(Rhodobacter)、发酵单胞菌属(Zymomonas)、芽孢杆菌属(Bacillus)、葡萄球菌属(Staphylococcus)、乳球菌属(Lactococcus)、链球菌属(Streptococcus)、乳杆菌属(Lactobacillus)、梭菌属(Clostridium)、棒杆菌属(Corynebacterium)、链霉菌属(Streptomyces)、双歧杆菌属(Bifidobacterium)、蓝细菌(Cyanobacterium)和圆杆菌属(Cyclobacterium)，但可以应用本发明的微生物不限于这些。优选地，本发明的微生物可以通过将重组载体引入选自大肠杆菌、假单胞菌属物种、棒杆菌属物种和红球菌属物种(但不限于这些)的微生物中来构建。作为大肠杆菌，优选使用选自下表2中所示的大肠杆菌菌株中的一种，但是本领域技术人员将理解，也可以使用预期表现出类似效果的其他大肠杆菌菌株。本发明的重组微生物可以通过使用作为假单胞菌属物种的恶臭假单胞菌、作为棒杆菌属物种的谷氨酸棒杆菌和作为红球菌属物种的混浊红球菌来构建，但这是代表性微生物的例子，并且应用本发明的微生物不限于这些。
此外，利用丙二酰-CoA作为底物或中间体，使用筛选的基因仅通过非常简单的遗传操作能够以高产率产生有用的物质。
因此，在第一个另外的方面，本发明涉及具有增加的丙二酰-CoA产生能力的重组微生物，其中选自以下基因的一种或多种基因：
fabF(3-氧代酰基-[酰基-载体-蛋白质]合酶II)；
yfcY(β-酮酰-CoA硫解酶)；
xapR(xapAB的转录激活因子)；
cytR(deo操纵子、udp、cdd、tsx、nupC和nupG的转录阻遏物)；
fabH(3-氧代酰基-[酰基-载体-蛋白质]合酶III)；
mqo(苹果酸脱氢酶)；
yfiD(丙酮酸甲酸裂解酶亚基)；
fmt(10-甲酰基四氢叶酸：L-甲硫氨酰基-tRNA(fMet)N-甲酰基转移酶)；
pyrF(乳清酸核苷-5'-磷酸脱羧酶)；
araA(L-阿拉伯糖异构酶)；
fadR(fad调节子的负调节因子和fabA的正调节因子)；
pabA(氨基脱氧分支酸合酶，亚基II)；
purB(腺苷酸琥珀酸裂解酶)；和
hycI(参与加工HycE的C末端的蛋白酶)在具有丙二酰-CoA产生能力的微生物中的表达与在野生型微生物中的表达相比降低。
具有丙二酰-CoA产生能力的微生物意指固有地具有丙二酰-CoA产生能力或外部引入丙二酰-CoA产生能力的微生物。
本发明还涉及使用丙二酰-CoA作为底物或中间体产生有用物质的方法，所述方法包括以下步骤：
(a)构建重组微生物，其中另外引入参与所述有用物质产生的基因或增加所述基因的表达，或者另外使参与所述有用物质产生的基因缺失或减少所述基因的表达；
(b)培养所构建的微生物；并且
(c)从所培养的微生物回收所述有用物质。
在本发明中，有用物质可以是选自以下项的一种或多种有用物质：
聚酮化合物，其由放线菌紫素、多柔比星、道诺霉素、土霉素、雷帕霉素、洛伐他汀、SEK4、SEK4b、SEK34、SEK15、SEK26、FK506、DMAC、阿克拉菌酮、阿克拉酸、ε-紫红霉酮、芦荟苦素、芦荟宁、芦荟苷、5,7-二羟基-2-甲基色酮、红霉素、利福霉素、阿维菌素、格尔德霉素、伊维菌素、多西环素、安曲霉素、青霉酸、卡奇霉素、埃博霉素、特曲霉素、富伦菌素、三乙酸内酯、6-甲基水杨酸和芦荟酮构成；
苯丙素类化合物，其由松属素、二氢山柰酚、圣草酚、二氢槲皮素、黄豆苷元、染料木素、芹菜素、木犀草素、山柰酚、槲皮素、儿茶素、天竺葵色素、矢车菊素、阿福豆素、杨梅素、非瑟酮、高良姜素、橙皮素、柑桔黄酮、花翠素、表儿茶素、白杨素、白藜芦醇和柚皮素构成；
生物燃料组，其由戊烷、己烷、庚烷、辛烷、壬烷、癸烷、十一烷、十二烷、十三烷、十四烷、十五烷、十六烷、十七烷、十八烷、十九烷、二十烷、戊醇、己醇、庚醇、辛醇、壬醇、癸醇、十一烷醇、十二烷醇、十三烷醇、十四烷醇、十五烷醇、十六烷醇、十七烷醇、十八烷醇、十九烷醇、二十烷醇、癸酸甲酯、月桂酸甲酯、肉豆蔻酸甲酯、棕榈酸甲酯、棕榈油酸甲酯、硬脂酸甲酯、油酸甲酯、亚油酸甲酯、亚麻酸甲酯、花生酸甲酯、二十烯酸甲酯、芥酸甲酯、癸酸乙酯、月桂酸乙酯、肉豆蔻酸乙酯、棕榈酸乙酯、棕榈油酸乙酯、硬脂酸乙酯、油酸乙酯、亚油酸乙酯、亚麻酸乙酯、花生酸乙酯、二十烯酸乙酯和芥酸甲酯构成；
由神经酰胺、棕榈酸酯和鞘氨醇构成的脂质化合物；和
3-羟基丙酸，但不限于这些。换言之，本发明的生物传感器(即重组微生物)可以应用于高价值产物，如除了上述有用物质之外的其他任何丙二酰-CoA来源的天然物质、化合物、生物燃料等。
本发明还涉及用于产生6-甲基水杨酸的重组微生物，其中在重组微生物中另外引入了编码6-甲基水杨酸合酶(6MSAS)和4'-磷酸泛酰巯基乙胺基转移酶(Sfp)的基因，或者增加所述基因的表达。
6MSAS可以源自展青霉(Penicillium patulum)(或灰黄青霉(Penicilliumgriseofulvum))、土曲霉(Aspergillus terreus)、棘孢曲霉(Aspergillus aculeatus)、黑曲霉(Aspergillus niger)、韦斯特迪克氏曲霉(Aspergillus westerdijkiae)、雪白丝衣霉(Byssochlamys nivea)、Glarea lozoyensis、扩展青霉(Penicillium expansum)或抗生链霉菌(Streptomyces antibioticus)，但不限于这些。
Sfp可以源自枯草芽孢杆菌(Bacillus subtilis)、产氨棒杆菌(Corynebacteriumammoniagenes)、大肠杆菌(Escherichia coli)、智人(Homo sapiens)、结核分枝杆菌(Mycobacterium tuberculosis)、铜绿假单胞菌(Pseudomonas aeruginosa)、蓖麻(Ricinus communis)、酿酒酵母(Saccharomyces cerevisiae)、菠菜(Spinaciaoleracea)、橙黄标桩菌(Stigmatella aurantiaca)、天蓝色链霉菌、肺炎链霉菌(Streptomyces pneumonia)、轮丝链霉菌(Streptomyces verticillus)或哈维氏弧菌(Vibrio harveyi)，但不限于这些。
与野生型微生物中基因的表达相比表达降低的基因可以是选自pabA、fabF、xapR和ytcY的一种或多种，但不限于此。
用于产生6-甲基水杨酸的重组微生物的特征可以在于：在重组微生物中另外引入了编码选自葡萄糖-6-磷酸脱氢酶(Zwf)、苹果酸脱氢酶(Mdh)、磷酸甘油酸脱氢酶(SerA)、乙酰-CoA羧化酶(AccBC和AccD1)、甘油醛3-磷酸脱氢酶(GapA)、磷酸甘油酸激酶(Pgk)、乙酰-CoA合成酶(Acs)和丙酮酸脱氢酶(PDH：AceEF和Lpd)的一种或多种酶的基因或增加所述基因的表达。
Zwf、Mdh、SerA、GapA、Pgk、Acs、AceE、AceF和Lpd可以源自大肠杆菌(E.coli)，但不限于此。AccBC和AccD1可以源自谷氨酸棒杆菌，但不限于此，并且可以扩展至对应于其他微生物来源的乙酰-CoA羧化酶的酶。
被筛选并用于本发明的每种酶的氨基酸序列如下，但可以在氨基酸序列中具有一个或多个氨基酸的取代、缺失或添加，并且在这种情况下，得到的突变氨基酸序列可以表现出与本发明相同或更好的功能。例如，明显的是，如果突变氨基酸序列的结构构型与野生型氨基酸序列相同或类似，并因此对底物表现出相同或更高的作用，则突变氨基酸序列也落入本发明的范围内。在相同的上下文中，明显的是，除了本发明中使用的酶之外，具有相同或类似功能的其他微生物来源的酶可以适当地应用于本发明，并且相应酶的在可由本领域普通技术人员容易地应用的范围内的应用落入本发明的范围内。
Zwf[SEQ ID NO:171]：
MAVTQTAQACDLVIFGAKGDLARRKLLPSLYQLEKAGQLNPDTRIIGVGRADWDKAAYTKVVREALETFMKETIDEGLWDTLSARLDFCNLDVNDTAAFSRLGAMLDQKNRITINYFAMP
PSTFGAICKGLGEAKLNAKPARVVMEKPLGTSLATSQEINDQVGEYFEECQVYRIDHYLGKETVLNLLALRFANSLFVNNWDNRTIDHVEITVAEEVGIEGRWGYFDKAGQMRDMIQNHLLQILCMIAMSPPSDLSADSIRDEKVKVLKSLRRIDRSNVREKTVRGQYTAGFAQGKKVPGYLEEEGANKSSNTETFVAIRVDIDNWRWAGVPFYLRTGKRLPTKCSEVVVYFKTPELNLFKESWQDLPQNKLTIRLQPDEGVDIQVLNKVPGLDHKHNLQITKLDLSYSETFNQTHLADAYERLLLETMRGIQALFVRRDEVEEAWKWVDSITEAWAMDNDAPKPYQAGTWGPVASVAMITRDGRSWNEFE
Mdh[SEQ ID NO:172]：
MKVAVLGAAGGIGQALALLLKTQLPSGSELSLYDIAPVTPGVAVDLSHIPTAVKIKGFSGEDATPALEGADVVLISAGVARKPGMDRSDLFNVNAGIVKNLVQQVAKTCPKACIGIITNPVNTTVAIAAEVLKKAGVYDKNKLFGVTTLDIIRSNTFVAELKGKQPGEVEVPVIGGHSGVTILPLLSQVPGVSFTEQEVADLTKRIQNAGTEVVEAKAGGGSATLSMGQAAARFGLSLVRALQGEQGVVECAYVEGDGQYARFFSQPLLLGKNGVEERKSIGTLSAFEQNALEGMLDTLKKDIALGEEFVNK
SerA[SEQ ID NO:173]：
MAKVSLEKDKIKFLLVEGVHQKALESLRAAGYTNIEFHKGALDDEQLKESIRDAHFIGLRSRTHLTEDVINAAEKLVAIGCFCIGTNQVDLDAAAKRGIPVFNAPFSNTRSVAELVIGELLLLLRGVPEANAKAHRGVWNKLAAGSFEARGKKLGIIGYGHIGTQLGILAESLGMYVYFYDIENKLPLGNATQVQHLSDLLNMSDVVSLHVPENPSTKNMMGAKEISLMKPGSLLINASRGTVVDIPALCDALASKHLAGAAIDVFPTEPATNSDPFTSPLCEFDNVLLTPHIGGSTQEAQENIGLEVAGKLIKYSDNGSTLSAVNFPEVSLPLHGGRRLMHIHENRPGVLTALNKIFAEQGVNIAAQYLQTSAQMGYVVIDIEADEDVAEKALQAMKAIPGTIRARLLY
GapA[SEQ ID NO:174]：
MTIKVGINGFGRIGRIVFRAAQKRSDIEIVAINDLLDADYMAYMLKYDSTHGRFDGTVEVKDGHLIVNGKKIRVTAERDPANLKWDEVGVDVVAEATGLFLTDETARKHITAGAKKVVMTGPSKDNTPMFVKGANFDKYAGQDIVSNASCTTNCLAPLAKVINDNFGIIEGLMTTVHATTATQKTVDGPSHKDWRGGRGASQNIIPSSTGAAKAVGKVLPELNGKLTGMAFRVPTPNVSVVDLTVRLEKAATYEQIKAAVKAAAEGEMKGVLGYTEDDVVSTDFNGEVCTSVFDAKAGIALNDNFVKLVSWYDNETGYSNKVLDLIAHISK
Pgk[SEQ ID NO:175]：
MSVIKMTDLDLAGKRVFIRADLNVPVKDGKVTSDARIRASLPTIELALKQGAKVMVTSHLGRPTEGEYNEEFSLLPVVNYLKDKLSNPVRLVKDYLDGVDVAEGELVVLENVRFNKGEKKDDETLSKKYAALCDVFVMDAFGTAHRAQASTHGIGKFADVACAGPLLAAELDALGKALKEPARPMVAIVGGSKVSTKLTVLDSLSKIADQLIVGGGIANTFIAAQGHDVGKSLYEADLVDEAKRLLTTCNIPVPSDVRVATEFSETAPATLKSVNDVKADEQILDIGDASAQELAEILKNAKTILWNGPVGVFEFPNFRKGTEIVANAIADSEAFSIAGGGDTLAAIDLFGIADKISYISTGGGAFLEFVEGKVLPAVAMLEERAKK
Acs[SEQ ID NO:176]：
MSQIHKHTIPANIADRCLINPQQYEAMYQQSINVPDTFWGEQGKILDWIKPYQKVKNTSFAPGNVSIKWYEDGTLNLAANCLDRHLQENGDRTAIIWEGDDASQSKHISYKELHRDVCRFANTLLELGIKKGDVVAIYMPMVPEAAVAMLACARIGAVHSVIFGGFSPEAVAGRIIDSNSRLVITSDEGVRAGRSIPLKKNVDDALKNPNVTSVEHVVVLKRTGGKIDWQEGRDLWWHDLVEQASDQHQAEEMNAEDPLFILYTSGSTGKPKGVLHTTGGYLVYAALTFKYVFDYHPGDIYWCTADVGWVTGHSYLLYGPLACGATTLMFEGVPNWPTPARMAQVVDKHQVNILYTAPTAIRALMAEGDKAIEGTDRSSLRILGSVGEPINPEAWEWYWKKIGNEKCPVVDTWWQTETGGFMITPLPGATELKAGSATRPFFGVQPALVDNEGNPLEGATEGSLVITDSWPGQARTLFGDHERFEQTYFSTFKNMYFSGDGARRDEDGYYWITGRVDDVLNVSGHRLGTAEIESALVAHPKIAEAAVVGIPHNIKGQAIYAYVTLNHGEEPSPELYAEVRNWVRKEIGPLATPDVLHWTDSLPKTRSGKIMRRILRKIAAGDTSNLGDTSTLADPGVVEKLLEEKQAIAMPS
AceE[SEQ ID NO:177]：
MSERFPNDVDPIETRDWLQAIESVIREEGVERAQYLIDQLLAEARKGGVNVAAGTGISNYINTIPVEEQPEYPGNLELERRIRSAIRWNAIMTVLRASKKDLELGGHMASFQSSATIYDVCFNHFFRARNEQDGGDLVYFQGHISPGVYARAFLEGRLTQEQLDNFRQEVHGNGLSSYPHPKLMPEFWQFPTVSMGLGPIGAIYQAKFLKYLEHRGLKDTSKQTVYAFLGDGEMDEPESKGAITIATREKLDNLVFVINCNLQRLDGPVTGNGKIINELEGIFEGAGWNVIKVMWGSRWDELLRKDTSGKLIQLMNETVDGDYQTFKSKDGAYVREHFFGKYPETAALVADWTDEQIWALNRGGHDPKKIYAAFKKAQETKGKATVILAHTIKGYGMGDAAEGKNIAHQVKKMNMDGVRHIRDRFNVPVSDADIEKLPYITFPEGSEEHTYLHAQRQKLHGYLPSRQPNFTEKLELPSLQDFGALLEEQSKEISTTIAFVRALNVMLKNKSIKDRLVPIIADEARTFGMEGLFRQIGIYSPNGQQYTPQDREQVAYYKEDEKGQILQEGINELGAGCSWLAAATSYSTNNLPMIPFYIYYSMFGFQRIGDLCWAAGDQQARGFLIGGTSGRTTLNGEGLQHEDGHSHIQSLTIPNCISYDPAYAYEVAVIMHDGLERMYGEKQENVYYYITTLNENYHMPAMPEGAEEGIRKGIYKLETIEGSKGKVQLLGSGSILRHVREAAEILAKDYGVGSDVYSVTSFTELARDGQDCERWNMLHPLETPRVPYIAQVMNDAPAVASTDYMKLFAEQVRTYVPADDYRVLGTDGFGRSDSRENLRHHFEVDASYVVVAALGELAKRGEIDKKVVADAIAKFNIDADKVNPRLA
AceF[SEQ ID NO:178]：
MAIEIKVPDIGADEVEITEILVKVGDKVEAEQSLITVEGDKASMEVPSPQAGIVKEIKVSVGDKTQTGALIMIFDSADGAADAAPAQAEEKKEAAPAAAPAAAAAKDVNVPDIGSDEVEVTEILVKVGDKVEAEQSLITVEGDKASMEVPAPFAGTVKEIKVNVGDKVSTGSLIMVFEVAGEAGAAAPAAKQEAAPAAAPAPAAGVKEVNVPDIGGDEVEVTEVMVKVGDKVAAEQSLITVEGDKASMEVPAPFAGVVKELKVNVGDKVKTGSLIMIFEVEGAAPAAAPAKQEAAAPAPAAKAEAPAAAPAAKAEGKSEFAENDAYVHATPLIRRLAREFGVNLAKVKGTGRKGRILREDVQAYVKEAIKRAEAAPAATGGGIPGMLPWPKVDFSKFGEIEEVELGRIQKISGANLSRNWVMIPHVTHFDKTDITELEAFRKQQNEEAAKRKLDVKITPVVFIMKAVAAALEQMPRFNSSLSEDGQRLTLKKYINIGVAVDTPNGLVVPVFKDVNKKGIIELSRELMTISKKARDGKLTAGEMQGGCFTISSIGGLGTTHFAPIVNAPEVAILGVSKSAMEPVWNGKEFVPRLMLPISLSFDHRVIDGADGARFITIINNTLSDIRRLVM
Lpd[SEQ ID NO:179]：
MSTEIKTQVVVLGAGPAGYSAAFRCADLGLETVIVERYNTLGGVCLNVGCIPSKALLHVAKVIEEAKALAEHGIVFGEPKTDIDKIRTWKEKVINQLTGGLAGMAKGRKVKVVNGLGKFTGANTLEVEGENGKTVINFDNAIIAAGSRPIQLPFIPHEDPRIWDSTDALELKEVPERLLVMGGGIIGLEMGTVYHALGSQIDVVEMFDQVIPAADKDIVKVFTKRISKKFNLMLETKVTAVEAKEDGIYVTMEGKKAPAEPQRYDAVLVAIGRVPNGKNLDAGKAGVEVDDRGFIRVDKQLRTNVPHIFAIGDIVGQPMLAHKGVHEGHVAAEVIAGKKHYFDPKVIPSIAYTEPEVAWVGLTEKEAKEKGISYETATFPWAASGRAIASDCADGMTKLIFDKESHRVIGGAIVGTNGGELLGEIGLAIEMGCDAEDIALTIHAHPTLHESVGLAAEVFEGSITDLPNPKAKKK
AccBC[SEQ ID NO:180]：
VSVETRKITKVLVANRGEIAIRVFRAARDEGIGSVAVYAEPDADAPFVSYADEAFALGGQTSAESYLVIDKIIDAARKSGADAIHPGYGFLAENADFAEAVINEGLIWIGPSPESIRSLGDKVTARHIADTAKAPMAPGTKEPVKDAAEVVAFAEEFGLPIAIKAAFGGGGRGMKVAYKMEEVADLFESATREATAAFGRGECFVERYLDKARHVEAQVIADKHGNVVVAGTRDCSLQRRFQKLVEEAPAPFLTDDQRERLHSSAKAICKEAGYYGAGTVEYLVGSDGLISFLEVNTRLQVEHPVTEETTGIDLVREMFRIAEGHELSIKEDPAPRGHAFEFRINGEDAGSNFMPAPGKITSYREPQGPGVRMDSGVVEGSEISGQFDSMLAKLIVWGDTREQALQRSRRALAEYVVEGMPTVIPFHQHIVENPAFVGNDEGFEIYTKWIEEVWDNPIAPYVDASELDEDEDKTPAQKVVVEINGRRVEVALPGDLALGGTAGPKKKAKKRRAGGAKAGVSGDAVAAPMQGTVIKVNVEEGAEVNEGDTVVVLEAMKMENPVKAHKSGTVTGLTVAAGEGVNKGVVLLEIK
AccD1[SEQ ID NO:181]：
MTISSPLIDVANLPDINTTAGKIADLKARRAEAHFPMGEKAVEKVHAAGRLTARERLDYLLDEGSFIETDQLARHRTTAFGLGAKRPATDGIVTGWGTIDGREVCIFSQDGTVFGGALGEVYGEKMIKIMELAIDTGRPLIGLYEGAGARIQDGAVSLDFISQTFYQNIQASGVIPQISVIMGACAGGNAYGPALTDFVVMVDKTSKMFVTGPDVIKTVTGEEITQEELGGATTHMVTAGNSHYTAATDEEALDWVQDLVSFLPSNNRSYTPLEDFDEEEGGVEENITADDLKLDEIIPDSATVPYDVRDVIECLTDDGEYLEIQADRAENVVIAFGRIEGQSVGFVANQPTQFAGCLDIDSSEKAARFVRTCDAFNIPIVMLVDVPGFLPGAGQEYGGILRRGAKLLYAYGEATVPKITVTMRKAYGGAYCVMGSKGLGSDINLAWPTAQIAVMGAAGAVGFIYRKELMAADAKGLDTVALAKSFEREYEDHMLNPYHAAERGLIDAVILPSETRGQISRNLRLLKHKNVTRPARKHGNMPL
本发明还涉及产生6-甲基水杨酸的方法，所述方法包括以下步骤：
(a)培养上述重组微生物；并且
(b)从所培养的微生物回收所述6-甲基水杨酸。
本发明的产生6-甲基水杨酸的方法可以包括通过添加1-50g/L的葡萄糖或1-100g/L的甘油作为碳源来培养重组微生物。
本发明还涉及用于产生芦荟酮的重组微生物，其中在重组微生物中另外引入了编码芦荟酮合酶的基因或增加所述基因的表达。在本发明中，芦荟酮合酶可以是ALS或PKS3，ALS可以源自掌叶大黄，并且PKS3可以源自木立芦荟，但除了所述芦荟酮合酶以外的芦荟酮合酶可以应用于本发明。
与野生型微生物中基因的表达相比表达降低的基因可以是pabA，但不限于此。
用于产生芦荟酮的重组微生物的特征可以在于：在重组微生物中另外引入了编码选自葡萄糖-6-磷酸脱氢酶(Zwf)、苹果酸脱氢酶(Mdh)、磷酸甘油酸脱氢酶(SerA)、乙酰-CoA羧化酶(AccBC和AccD1)、甘油醛3-磷酸脱氢酶(GapA)、磷酸甘油酸激酶(Pgk)、乙酰-CoA合成酶(Acs)和丙酮酸脱氢酶(PDH：AceEF和Lpd)的一种或多种酶的基因或增加所述基因的表达。
Zwf、Mdh、SerA、GapA、Pgk、Acs、AceE、AceF和Lpd可以源自大肠杆菌(E.coli)，但不限于此。AccBC和AccD1可以源自谷氨酸棒杆菌，但不限于此，并且可以扩展至对应于其他微生物来源的乙酰-CoA羧化酶的酶。
本发明还涉及产生芦荟酮的方法，所述方法包括以下步骤：
(a)培养上述重组微生物；并且
(b)从所培养的微生物回收所述芦荟酮。
本发明的产生芦荟酮的方法可以包括通过添加1-50g/L的葡萄糖或1-100g/L的甘油作为碳源来培养重组微生物。
本发明还涉及用于产生白藜芦醇的重组微生物，其中在重组微生物中另外引入了编码酪氨酸氨裂解酶(TAL)、4-香豆酸:CoA连接酶(4CL)和芪合酶(STS)的基因或者增加所述基因的表达。
TAL可以源自荚膜红细菌(Rhodobacter capsulatus)、蝶豆(Clitoriaternatea)、草莓(Fragaria x ananassa)、类球红细菌(Rhodobacter sphaeroides)、玉米(Zea mays)或西班牙糖丝菌(Saccharothrix espanaensis)，
4CL可以源自拟南芥(Arabidopsis thaliana)、天蓝色链霉菌、伍氏醋酸杆菌(Acetobacterium woodii)、藿香(Agastache rugose)、燕麦(Avena sativa)、野茶树(Camellia sinensis)、百金花(Centaurium erythraea)、翁柱(Cephalocereus senilis)、椰子(Cocos nucifera)、枇杷(Eriobotrya japonica)、鸡冠刺桐(Erythrinacristagalli)、连翘属物种(Forsythia sp.)、草莓(Fragaria x ananassa)、大豆(Glycinemax)、陆地棉(Gossypium hirsutum)、大麻槿(Hibiscus cannabinus)、卡氏落叶松(Larixcajanderi)、兴安落叶松(Larix gmelinii)、日本落叶松(Larix kaempferi)、Larixkamtschatica、西伯利亚落叶松(Larix sibirica)、Larix sukaczewii、紫草(Lithospermum erythrorhizon)、多年生黑麦草(Lolium perenne)、金银花(Lonicerajaponica)、水杉(Metasequoia glyptostroboides)、烟草(Nicotiana tabacum)、罗勒(Ocimum basilicum)、小冠熏(Ocimum tenuiflorum)、稻(Oryza sativa)、毛泡桐(Paulownia tomentosa)、香芹(Petroselinum crispum)、桂竹(Phyllostachysbambusoides)、小立碗藓(Physcomitrella patens)、欧洲云杉(Picea abies)、辐射松(Pinus radiate)、火炬松(Pinus taeda)、豌豆(Pisum sativum)、侧柏(Platycladusorientalis)、Polyporus hispidus、毛白杨(Populus tomentosa)、颤杨(Populustremuloides)、加杨(Populus x canadensis)、欧洲甜樱桃(Prunus avium)、野葛(Pueraria montana)、刺槐(Robinia pseudoacacia)、芸香(Ruta graveolens)、酿酒酵母、垂柳(Salix babylonica)、番茄(Solanum lycopersicum)、马铃薯(Solanum tuberosum)、欧洲花楸(Sorbus aucuparia)、小麦(Triticum aestivum)或葡萄(Vitis vinifera)，并且
STS可以源自落花生(Arachis hypogaea)、赤松(Pinus densiflora)、马尾松(Pinus massoniana)、北美乔松(Pinus strobus)、虎杖(Polygonumcuspidatum)、松叶蕨(Psilotum nudum)或葡萄，但不限于这些。
4-香豆酸:CoA连接酶可以是其中SEQ ID NO:128所示的氨基酸序列中的氨基酸突变为I250L/N404K/I461V的突变酶，或者是其中SEQ ID NO:131所示的氨基酸序列中的氨基酸突变为A294G/A318G的突变酶，但不限于这些。
与野生型微生物中基因的表达相比表达降低的基因可以是选自pabA、yfiD、mqo、xapR、purB、fabH、fabF、ytcY、argC、nudD、araA、fadR、cytR、fmt和pyrF的一种或多种，但不限于这些。
本发明还涉及产生白藜芦醇的方法，所述方法包括以下步骤：
(a)培养上述重组微生物；并且
(b)从所培养的微生物回收所述白藜芦醇。
本发明的产生白藜芦醇的方法可以包括通过添加1-50g/L的葡萄糖或1-100g/L的甘油作为碳源来培养重组微生物。
本发明还涉及用于产生柚皮素的重组微生物，其中在上述重组微生物中另外引入了编码酪氨酸氨裂解酶(TAL)、4-香豆酸:CoA连接酶(4CL)、查耳酮合酶(CHS)和查耳酮异构酶(CHI)的基因或者增加所述基因的表达。
TAL可以源自荚膜红细菌(Rhodobacter capsulatus)、蝶豆(Clitoriaternatea)、草莓(Fragaria x ananassa)、类球红细菌(Rhodobacter sphaeroides)、玉米(Zea mays)或西班牙糖丝菌(Saccharothrix espanaensis)，
4CL可以源自拟南芥(Arabidopsis thaliana)、天蓝色链霉菌、伍氏醋酸杆菌(Acetobacterium woodii)、藿香(Agastache rugose)、燕麦(Avena sativa)、野茶树(Camellia sinensis)、百金花(Centaurium erythraea)、翁柱(Cephalocereus senilis)、椰子(Cocos nucifera)、枇杷(Eriobotrya japonica)、鸡冠刺桐(Erythrinacristagalli)、连翘属物种(Forsythia sp.)、草莓(Fragaria x ananassa)、大豆(Glycinemax)、陆地棉(Gossypium hirsutum)、大麻槿(Hibiscus cannabinus)、卡氏落叶松(Larixcajanderi)、兴安落叶松(Larix gmelinii)、日本落叶松(Larix kaempferi)、Larixkamtschatica、西伯利亚落叶松(Larix sibirica)、Larix sukaczewii、紫草(Lithospermum erythrorhizon)、多年生黑麦草(Lolium perenne)、金银花(Lonicerajaponica)、水杉(Metasequoia glyptostroboides)、烟草(Nicotiana tabacum)、罗勒(Ocimum basilicum)、小冠熏(Ocimum tenuiflorum)、稻(Oryza sativa)、毛泡桐(Paulownia tomentosa)、香芹(Petroselinum crispum)、桂竹(Phyllostachysbambusoides)、小立碗藓(Physcomitrella patens)、欧洲云杉(Picea abies)、辐射松(Pinus radiate)、火炬松(Pinus taeda)、豌豆(Pisum sativum)、侧柏(Platycladusorientalis)、Polyporus hispidus、毛白杨(Populus tomentosa)、颤杨(Populustremuloides)、加杨(Populus x canadensis)、欧洲甜樱桃(Prunus avium)、野葛(Pueraria montana)、刺槐(Robinia pseudoacacia)、芸香(Ruta graveolens)、酿酒酵母、垂柳(Salix babylonica)、番茄(Solanum lycopersicum)、马铃薯(Solanum tuberosum)、欧洲花楸(Sorbus aucuparia)、小麦(Triticum aestivum)或葡萄(Vitis vinifera)，
CHS可以源自小苍兰属杂交品种(Freesia hybrid cultivar)、紫花苜蓿(Medicago sativa)、小立碗藓、钝鳞紫背苔(Plagiochasma appendiculatum)、小麦、葡萄、甜橙(Citrus sinensis)、拟南芥、燕麦、胡萝卜(Daucus carota)、荞麦(Fagopyrumesculentum)、大豆(Glycine max)、棘甘草(Glycyrrhiza echinata)、啤酒花(Humuluslupulus)、金丝桃(Hypericum androsaemum)、香芹、小立碗藓、覆盆子(Rubus idaeus)、黄岑(Scutellaria baicalensis)、Xanthisma gracile、硫华菊(Cosmos sulphureus)、非洲菊杂交品种(Gerbera hybrid cultivar)、大麦(Hordeum vulgare)、胡桃属物种(Juglanssp.)、菜豆(Phaseolus vulgaris)、野葛(Pueraria montana)、黑麦(Secale cereal)、蝇子草属物种(Silene sp.)、白芥子(Sinapis alba)、菠菜、长须银柴胡(Stellarialongipes)、郁金香属杂交品种(Tulipa hybrid cultivar)、马鞭草属物种(Verbena sp.)或矮牵牛(Petunia x hybrida)，并且
CHI可以源自紫苏(Perilla frutescens)、银杏(Ginkgo biloba)、胡芦巴(Trigonella foenumgraecum)、紫花苜蓿、黄岑、大豆、翁柱、甜橙、棘甘草、乌拉尔甘草(Glycyrrhiza uralensis)、白花百合(Lilium candidum)、杨梅(Morella rubra)、矮牵牛、菜豆、Soja hispida、郁金香属杂交品种、拟南芥或烟草，但不限于这些。
4-香豆酸:CoA连接酶可以是其中SEQ ID NO:128所示的氨基酸序列中的氨基酸突变为I250L/N404K/I461V的突变酶，但不限于此。
与野生型微生物中基因的表达相比表达降低的基因可以是选自fadR、hycI和xapR的一种或多种。
本发明还涉及产生柚皮素的方法，所述方法包括以下步骤：
(a)培养上述重组微生物；并且
(b)从所培养的微生物回收所述柚皮素。
本发明的产生柚皮素的方法可以包括通过添加1-50g/L的葡萄糖或1-100g/L的甘油作为碳源来培养重组微生物。
在本发明中，基因的“调节”包括所有基因缺失、表达抑制、表达促进、敲低、启动子替换和通过技术(如调节机制)诱导基因的表达，并且意在包括使生物合成途径中呈现的一种或多种酶进化或突变。
在本发明中，“敲低”意指显著降低基因的表达水平以降低基因的功能，这与完全阻断基因表达的“敲除”不同。它可以在基因转录物水平或蛋白质水平上进行调节。然而，由于本发明在抑制或降低编码参与相应途径的酶的基因的表达方面具有重要意义，因此使用“敲低”和“敲除”中的任何一种都可以实现所需的目的。
如本文所使用的，术语“载体”是指包含与合适的控制序列可操作连接的DNA序列的DNA构建体，其能够在合适的宿主中实现DNA的表达。载体可以是质粒、噬菌体颗粒、或简单地潜在的基因组插入物。一旦掺入合适的宿主中，载体可以独立于宿主基因组复制和起作用，或者在一些情况下可以整合到基因组本身中。在本说明书中，“质粒”和“载体”有时可互换使用，因为质粒是最常用的载体形式。出于本发明的目的，优选使用质粒载体。可用于此目的的典型质粒载体包含：(a)复制起点，通过所述复制起点高效地进行复制，使得每个宿主细胞产生数百个质粒载体；(b)抗生素抗性基因，可以通过所述抗生素抗性基因选择用质粒载体转化的宿主细胞；和(c)可以插入外源DNA片段的限制酶切割位点。即使载体中不存在合适的限制酶切割位点，使用常规合成寡核苷酸衔接子或接头也能够将载体与外源DNA片段之间容易连接。连接后，载体应转化至合适的宿主细胞中。通过氯化钙法或电穿孔(Neumann等人,EMBO J.,1:841,1982)可以容易地实现转化。
载体的启动子可以是组成型或诱导型的，并且可以进一步修饰以实现本发明的效果。此外，表达载体包含用于选择含有所述载体的宿主细胞的选择标记物，并且可复制表达载体包含复制起点。载体可以是自我复制的，或者可以整合至宿主基因组DNA中。优选地，插入载体中的基因不可逆地融合至宿主细胞的基因组中，使得所述基因在细胞中的表达长时间稳定地持续。
当核苷酸序列被放置成与另一核苷酸序列有功能关系时，所述核苷酸序列为“可操作地连接”。核苷酸序列可以是基因和连接的一个或多个控制序列，当所述基因与一个或多个控制序列(例如转录激活蛋白)结合时，所述一个或多个控制序列能够使得基因表达。例如，当表达为参与多肽分泌的前蛋白时，用于前序列或分泌前导序列的DNA与编码多肽的DNA可操作地连接；当影响序列的转录时，启动子或增强子与编码序列可操作地连接；以及当影响序列的转录时，核糖体结合位点(RBS)与编码序列可操作地连接，或当安排为促进翻译时，RBS与编码序列可操作地连接。通常，术语“可操作地连接”是指DNA连接的序列是连续的，并且在分泌前导序列的情况下是连续的并且存在于阅读框中。但是，增强子不一定是连续的。这些序列之间通过在便利的限制性酶切位点处的连接进行连接。然而，当位点不存在时，根据常规方法使用合成的寡核苷酸衔接子或接头。
如本领域熟知的，为了增加转染基因在宿主细胞中的表达水平，相应的基因应该与在选择的表达宿主中操作的转录和翻译表达控制序列可操作地连接。优选地，表达控制序列和相应的基因与细菌选择标记物和复制起点一起被包括在一个重组载体中。当宿主细胞是真核细胞时，重组载体应还包含可用于真核表达宿主的表达标记物。
由上述重组载体转化的宿主细胞构成了本发明的另一个方面。如本文所使用的，术语“转化”是指通过将DNA导入宿主中，DNA可以作为染色体外的因子或通过完整的染色体的完成来复制。
当然，应该理解，根据本发明所有载体和表达控制序列不能等同地发挥作用以表达DNA序列。同样，对于相同的表达系统，所有宿主并非等同地发挥作用。然而，本领域技术人员可以在不脱离本发明的范围或承担过多的实验负担的情况下，从各种载体、表达控制序列和宿主中作出适当的选择。例如，必须在考虑宿主的情况下选择载体，因为必须在宿主中复制载体。具体地，还应考虑载体的拷贝数、调节拷贝数的能力和由相应载体编码的其他蛋白的表达(例如抗生素标记物的表达)。
另外，可以将本发明中引入的基因引入宿主细胞的基因组中并以染色体因子的形式呈现。对于本发明所属领域的技术人员明显的是，即使当将基因插入宿主细胞的基因组染色体中时，所述基因的作用与当如上所述将重组载体引入宿主细胞中时的作用也是相同的。
实施例
在下文将参照实施例进一步详细描述本发明。对于本领域普通技术人员将明显的是，这些实施例仅用于说明目的，而不应被解释为限制本发明的范围。
实施例1.基于III型聚酮合酶的丙二酰-CoA生物传感器的构建
1.1.多种III型聚酮合酶的性能测试
本实施例和以下实施例中使用的限制酶购自New England Labs(美国)或Enzynomics(韩国)，并且PCR聚合酶购自Biofact(韩国)。其他的单独标记。另外，本实施例和以下实施例中引入的外源基因与在其上含有信息的数据库中的登录号及其序列一起总结在下表1中。另外，除非另有说明，否则用于克隆的大肠杆菌是DH5α菌株，将其在LB培养基(每升：10g胰蛋白胨，5g酵母提取物和10g NaCl)中或在LB琼脂板(1/5％，v/v)上培养。此外，如果需要的话，添加适当浓度的抗生素(50μg/mL卡那霉素，100μg/mL氨苄青霉素和/或100μg/mL大观霉素)。
在本发明中，使用III型聚酮合酶RppA由五分子丙二酰-CoA产生红色淡黄霉素，并将其用作细胞内丙二酰-CoA水平的指示剂(图1)。为此，比较和测试了多种III聚酮合酶的性能。克隆了来自总共五种放线菌菌株(灰色链霉菌、天蓝色链霉菌、阿维链霉菌、红色糖多孢菌、针孢链霉菌(Streptomyces aculeolatus))的III型聚酮合酶编码rppA基因，并将每一种基因转化至大肠杆菌BL21(DE3)菌株中。然后，将每种构建的菌株接种至含有补充有10g/L葡萄糖的M9基本培养基的烧瓶中，随后在初始指数期用0.5mM异丙基-β-D-1-硫代半乳糖苷(IPTG)处理。接着，将每种菌株培养48小时，并观察到淡黄霉素的产生。此时，观察到在表达基于灰色链霉菌的酶Sgr_RppA的菌株中淡黄霉素产量最高(25.7mg/L；图2A)。构建的质粒是基于pET-30a(+)(Novagen)，并且分别命名为pET-Sgr_rppA、pET-Sco_rppA、pET-Sma_rppA、pET-Sen_rppA和pET-Sac_rppA。为了构建pET-Sgr_rppA，使用灰色链霉菌的基因组DNA作为模板以及引物对SEQ ID NO:1和SEQ ID NO:2扩增rppA基因。使用Gibson组件将基因在EcoRI和BamHI位点处克隆到pET-30a(+)质粒中(Gibson DG等人(2009),NatureMethods 6:343-345)。其他质粒也通过与上述相同的方法构建。M9基本培养基每升含有以下组分：12.8g Na2HPO4·7H2O、3g KH2PO4、0.5g NaCl、1g NH4Cl、2mM MgSO4、0.1mM CaCl2。
[SEQ ID NO: 1] 5'-CTTTAAGAAGGAGATATACATATGGCGACCCTGTGCCGACC-3'
[SEQ ID NO: 2] 5'-CTTGTCGACGGAGCTCGAATTCATTAGCCGGACAGCGCAACGC-3'
[表1]
关于本发明中提出的外源基因的信息
*描绘了酶的NCBI编号
在图1A中，通过RppA酶将丙二酰-CoA转化为THN。然后，通过自发氧化反应将THN转化为淡黄霉素。据报道，WhiE_ORFIII参与这一最终步骤(Austin MB等人(2004),J BiolChem 279:45162-45174)，并且因此在本发明中，将相应基因另外克隆至pET-30a(+)中，但这导致降低的淡黄霉素产量。这表明rppA单个基因足以产生淡黄霉素。通过LC-MS和MS/MS证实了产生的淡黄霉素(图2B)。此后，除了常规的基于T7启动子的系统之外，在基于tac启动子的质粒中针对其在所有大肠杆菌菌株中的用途建立rppA表达平台。为此，首先构建了基于pCDFDuet-1(Novagen)质粒的pTacCDFS质粒，其含有CDF复制起点并具有大观霉素抗生素抗性和基于tac启动子的基因表达盒。为此，使用pTac15K质粒(Lee SY等人(2008)，美国专利20110269183)作为模板以及引物SEQ ID NO:3和SEQ ID NO:4通过PCR扩增含有tac启动子的DNA片段，并且使用pCDFDuet-1质粒作为模板以及引物SEQ ID NO:5和SEQ ID NO:6将相应质粒线性化。使用Gibson组件(Gibson assembly)将产生的两个DNA片段进行组装，从而构建pTacCDFS质粒。还构建了pTrcCDFS质粒用于进一步使用，并且此质粒与pTacCDFS质粒相同，不同之处在于前者含有trc启动子。另外，以与pTacCDFS质粒相同的方式构建pTrcCDFS质粒，不同之处在于使用pTrc99A质粒(Lee SY等人(2008)，美国专利20110269183)作为模板通过PCR扩增含有trc启动子的DNA片段。此后，通过以下方法构建基于pTacCDFS质粒的rppA表达质粒pTac-Sgr_rppA。首先，使用引物SEQ ID NO:7和SEQ IDNO:8从灰色链霉菌的基因组DNA扩增rppA基因，并且使用引物SEQ ID NO:9和SEQ ID NO:10将pTacCDFS质粒线性化。使用Gibson组件将两个产生的DNA片段进行组装，从而构建pTac-Sgr_rppA质粒。此外，为了优化rppA的表达，进行5'非翻译区(5'UTR)的优化。使用先前报道的UTR设计器(url:https://sbi.postech.ac.kr/utr_designer/)设计5’UTR DNA序列。为了构建相应引物pTac-5'UTR-Sgr_rppA，使用pTac-Sgr_rppA作为模板以及引物SEQ ID NO:9和SEQ ID NO:11进行反向PCR。用DpnI处理产生的线性化质粒以去除模板，并通过T4多核苷酸激酶(PNK)(Enzynomics，韩国)和T4连接酶(Elpis Biotech，韩国)处理进行连接。将得到的质粒引入大肠杆菌BL21(DE3)菌株中，并在M9基本培养基中培养，从而产生17.8mg/L的淡黄霉素(图2C)。
[SEQ ID NO:3] 5'-CGACTCCTGCATTAGGAAATGACTGCACGGTGCACCAATG-3'
[SEQ ID NO:4] 5'-GCGTTTCACTTCTGAGTTCG-3'
[SEQ ID NO:5] 5'-CGAACTCAGAAGTGAAACGCCTGAAACCTCAGGCATTTGAG-3'
[SEQ ID NO:6] 5'-ATTTCCTAATGCAGGAGTCG-3'
[SEQ ID NO:7] 5'-CAATTTCACACAGGAAACAGAATGGCGACCCTGTGCCGACC-3'
[SEQ ID NO:8] 5'-CTCTAGAGGATCCCCGGGTACCATTAGCCGGACAGCGCAACGC-3'
[SEQ ID NO:9] 5'-GGTACCCGGGGATCCTCTAGAG-3'
[SEQ ID NO:10] 5'-TCTGTTTCCTGTGTGAAATTG-3'
[SEQ ID NO:11] 5'-GATGCTCCTTTCTTGTTATTGAAATTGTTATCCGCTCACAATTC-3'
1.2.基于III型聚酮合酶的丙二酰-CoA生物传感器的适用性的检验
在构建基于灰色链霉菌RppA的丙二酰-CoA生物传感器后，进行了以下实验以检验生物传感器的适用性。首先，将RppA丙二酰-CoA生物传感器的信号定义为340nm处培养上清液的吸光度(图3D)。通常，为了表征生物传感器，必须添加不同浓度的目标化合物以检查信号是否增加。然而，因为丙二酰-CoA是不能穿过细胞膜的物质，所以将丙二酰-CoA添加至培养基中是没有意义的。因此，使用已知阻断脂肪酸生物合成途径的浅蓝菌素(Sigma-aldrich，美国)来间接表征生物传感器(Omura S(1976),Bacteriol Rev40:681-697)。浅蓝菌素阻断脂肪酸生物合成途径，从而导致丙二酰-CoA(一种脂肪酸前体)的积累。因此，测量了根据先前研究中所述添加不同浓度的浅蓝菌素时在大肠杆菌中丙二酰-CoA的浓度(XuP,Li L,Zhang F,Stephanopoulos G,Koffas M(2014),Proc Natl Acad Sci U S A111:11299-11304)。将含有质粒pTac-5'UTR-Sgr_rppA的大肠杆菌BL21(DE3)菌株接种至14-mL一次性Falcon圆底试管中的3mL LB培养基中。接着，将菌株在培养箱中于30℃和220rpm下培养16小时，随后接种至另一个14-mL一次性Falcon圆底试管中的3mL改良R/2培养基(pH6.8)中。将改良R/2培养基制备为每升含有以下组分：3g酵母提取物、6.75g KH2PO4、2g(NH4)2HPO4、0.8g MgSO4·7H2O、3g(NH4)2SO4、0.85g柠檬酸、5mL痕量金属溶液[每1L的0.1MHCl：10g FeSO4·7H2O、2.25g ZnSO4·7H2O、0.58g MnSO4·5H2O、1g CuSO4·5H2O、0.1g(NH4)6Mo7O24、4H2O、0.02g Na2B4O7·10H2O、2g CaCl2·2H2O]。向其中添加20g/L葡萄糖、100μg/mL大观霉素、0.2mM IPTG和适当浓度的浅蓝菌素。添加的浅蓝菌素浓度如下：0、5、10、15、20、25、50和100μM。将接种的细胞在培养箱中于30℃和220rpm下培养16小时，随后测量细胞的生长(OD600)和培养上清液在340nm处的吸光度。测量结果显示在图2D中。此时，可以看出，与不表达RppA的菌株相比，表达RppA的菌株生成了随着浅蓝菌素浓度的增加而明显增加的归一化信号。如图3A所示，可以证实，如通过肉眼观察到的那样，淡黄霉素的归一化产量也随着浅蓝菌素浓度的增加而增加(图3B)。此外，证实了当将pTac-5'UTR-Sgr_rppA质粒引入16种不同的大肠杆菌菌株(表2)中时，所有菌株均产生淡黄霉素。这表明本发明中提出的丙二酰-CoA生物传感器适用于所有大肠杆菌菌株(图3C)。
[表2]
此外，为了证明除了基于灰色链霉菌RppA的丙二酰-CoA生物传感器以外的其他III型聚酮合酶也可用作丙二酰-CoA生物传感器，测试了三种III型聚酮合酶。为此，测试了木立芦荟III型聚酮合酶(八酮合酶，OKS)[SEQ ID NO:103]、木立芦荟III型聚酮合酶(八酮合酶2，PKS4)[SEQ ID NO:105]和木立芦荟III型聚酮合酶(八酮合酶3，PKS5)[SEQ ID NO:106]，如下构建质粒pET-AaOKS、pET-AaPKS4和pET-AaPKS5，每一种质粒含有编码每一种酶的基因。首先，使用合成的AaOKS基因(Integrated DNA Technologies,Inc.，美国)作为模板以及引物SEQ ID NO:137和SEQ ID NO:138通过PCR扩增AaOKS，随后将所述AaOKS在NdeI和EcoRI位点处插入pET-30a(+)质粒中。其他质粒也以如上所述的相同方法来构建，不同之处在于AaPKS4是使用引物SEQ ID NO:141和SEQ ID NO:142来扩增，而AaPKS5是使用引物SEQ ID NO:143和SEQ ID NO:144来扩增。
[SEQ ID NO: 135] 5'-CTTTAAGAAGGAGATATACATATGAGTTCACTCTCCAACTCTC-3'
[SEQ ID NO: 136] 5'-CTTGTCGACGGAGCTCGAATTCATTACATGAGAGGCAGGCTGTG-3'
[SEQ ID NO: 137] 5'-CTTTAAGAAGGAGATATACATATGAGTTCACTCTCCAACGCTTC-3'
[SEQ ID NO: 138] 5'-CTTGTCGACGGAGCTCGAATTCATTACATGAGAGGCAGGCTGTG-3'
[SEQ ID NO: 139] 5'-CTTTAAGAAGGAGATATACATATGGGTTCACTCTCCGACTCTAC-3'
[SEQ ID NO: 140] 5'-CTTGTCGACGGAGCTCGAATTCATTACACAGGAGGCAGGCTATG-3'
[SEQ ID NO: 141] 5'-CTTTAAGAAGGAGATATACATATGGGTTCACTCTCCAACTAC-3'
[SEQ ID NO: 142] 5'-CTTGTCGACGGAGCTCGAATTCATTACACAGGGGGCAGGCTG-3'
[SEQ ID NO: 143] 5'-CTTTAAGAAGGAGATATACATATGGGTTCGATCGCCGAG-3'
[SEQ ID NO: 144] 5'-CTTGTCGACGGAGCTCGAATTCATTAGACAGGAGGCAGGCTG-3'
[SEQ ID NO: 145] 5'-GCTCACCATCATTGGGCTCCGTGGCCCCAATG-3'
[SEQ ID NO: 146] 5'-CATTGGGGCCACGGAGCCCAATGATGGTGAGC-3'
当培养含有每一种构建的质粒的大肠杆菌BL21(DE3)菌株时，可以观察到表达AaOKS、AaPKS4和AaPKS5的菌株的颜色变化。因此，通过LC-MS分析对促成颜色变化的聚酮合酶进行了分析(图4A)。如上所述，进行了使用浅蓝菌素的丙二酰-CoA实验，并且结果表明，随着细胞内丙二酰-CoA浓度的增加，由每种菌株生成的信号增加(图4和图5)。此处，信号指示培养上清液在300nm处的吸光度(图4C)。因此，在本发明中，证明了不仅RppA酶，能够由丙二酰-CoA产生基于有色聚酮的化合物的所有其他III型聚酮合酶都可以用作丙二酰-CoA生物传感器。
在以下实施例中，使用上述III型聚酮合酶中的RppA进行另外的实验。
1.3.其他工业上有用的菌株的可扩展性的检验
在上述实施例中证明了在大肠杆菌中的适用性后，检验这种RppA丙二酰-CoA生物传感器对于其他工业上有用的菌株的可扩展性。除了大肠杆菌之外，此检验中使用的菌株的例子包括恶臭假单胞菌(它是代表性的革兰氏阴性菌)以及谷氨酸棒杆菌和混浊红球菌(它们是代表性革兰氏阳性菌)。首先，使用pBBR1MCS2作为基础质粒构建了用于在恶臭假单胞菌菌株中产生淡黄霉素的质粒(Kovach ME等人(1995),Gene 166:175-176)。首先，使用引物SEQ ID NO:12和SEQ ID NO:13将pBBR1MCS2质粒线性化。然后，使用pTac-Sgr_rppA质粒作为模板以及引物SEQ ID NO:14和SEQ ID NO:15通过PCR扩增含有tac启动子的rppA表达盒。使用Gibson组件将获得的两个DNA片段进行组装，从而构建pBBR1-rppA质粒。此外，为了检验N末端聚His标签对酶表达和淡黄霉素产量的影响，首先构建了pTac-His-Sgr_rppA。此时，使用Gibson组件将使用引物SEQ ID NO:9和SEQ ID NO:10通过PCR线性化的pTacCDFS质粒和使用引物SEQ ID NO:16和SEQ ID NO:8从灰色链霉菌的基因组DNA通过PCR扩增的经His标记的rppA基因组装至单个质粒(pTac-His-Sgr_rppA)中。将构建的质粒使用引物SEQID NO:14和SEQ ID NO:15通过PCR再次扩增，并将其使用Gibson组件与线性化pBBR1MCS2质粒进行组装，从而构建pBBR1-His-rppA质粒。此时，将pBBR1-rppA和pBBR1-His-rppA质粒中的每一种转化至恶臭假单胞菌菌株中，随后将每一种得到的菌株接种至含有5mL LB培养基的试管中，然后在30℃和220rpm下培养18小时。接着，将0.8g/L MgSO4·7H2O、10g/L葡萄糖和抗生素添加至5mL LB中，然后进行传代培养。在相同条件下培养25小时后，进行采样。作为结果，在含有pBBR1-rppA质粒的菌株中产生了较大量的淡黄霉素(44.7mg/L；图6A)。因此，使用筛选的恶臭假单胞菌pBBR1-rppA菌株，以与上文实施例1.2中所述相同的方式使用浅蓝菌素进行丙二酰-CoA生物传感器的表征。作为结果，可以观察到淡黄霉素的归一化信号和归一化产量也随着相应菌株中浅蓝菌素浓度的增加而增加(图6B和图6C)。此处，在补充有20g/L葡萄糖、25μg/mL卡那霉素、0.5mM IPTG和适当浓度的浅蓝菌素的2mL MR-MOPS培养基中进行传代培养，并且MR-MOPS培养基的组合物(pH 7.0)每升含有以下组分：6.67gKH2PO4、4g(NH4)2HPO4、0.8g柠檬酸、5mL痕量金属溶液、20.09g MOPS(3-(N-吗啉代)丙磺酸)、0.8g/L MgSO4·7H2O。结果证明RppA丙二酰-CoA生物传感器成功地在恶臭假单胞菌中起作用。
[SEQ ID NO:12] 5'-CGAACTCAGAAGTGAAACGCTGCCTAATGAGTGAGCTAAC-3'
[SEQ ID NO:13] 5'-CATTGGTGCACCGTGCAGTCAAAATTCGCGTTAAATTTTTG-3'
[SEQ ID NO:14] 5'-GACTGCACGGTGCACCAATG-3'
[SEQ ID NO:15] 5'-GCGTTTCACTTCTGAGTTCG-3'
[SEQ ID NO:16] 5'-CAATTTCACACAGGAAACAGAATGCACCATCACCATCACCATGCGACCCTGTGC CGACC-3'
此外，为了将此生物传感器应用于谷氨酸棒杆菌菌株，使用pCES-H36作为基础质粒构建rppA表达质粒(Yim SS,An SJ,Kang M,Lee J,Jeong KJ(2013),Biotechnol Bioeng110:2959-2969)。为此，使用pCES-H36-GFP质粒作为模板以及引物SEQ ID NO:17和SEQ IDNO:18将相应质粒线性化。使用pTac-Sgr_rppA作为模板以及引物SEQ ID NO:19和SEQ IDNO:20通过PCR扩增基于tac启动子的rppA表达盒，并使用pTac-His-Sgr_rppA作为模板以及引物SEQ ID NO:21和SEQ ID NO:20扩增基于tac启动子的N末端His标记的rppA表达盒。使用Gibson组件将两种不同的rppA表达盒与上述线性化pCES-H36质粒进行组装，从而构建pCES-rppA和pCES-His-rppA质粒。将构建的质粒转化至谷氨酸棒杆菌菌株中，然后将其接种至5mL LB培养基中。将每一种所得菌株在培养箱中于30℃和220rpm下培养18小时，然后将其在5mL LB培养基中传代培养48小时，之后进行采样。此时，仅在含有N末端His标记的rppA表达盒(pCES-His-rppA)的菌株中产生了3.9mg/L淡黄霉素(图6D)。因此，使用选择的谷氨酸棒杆菌pCES-His-rppA菌株，以与上文实施例1.2中所述相同的方式使用浅蓝菌素进行丙二酰-CoA生物传感器的表征。作为结果，可以观察到淡黄霉素的归一化信号和归一化产量也随着相应菌株中浅蓝菌素浓度的增加而增加(图6E和图6F)。
[SEQ ID NO:17]5'-CCATTATAATTAGGCCTCGG-3'
[SEQ ID NO:18] 5'-CCATGCTACTCCTACCAACC-3'
[SEQ ID NO:19] 5'-GGTTGGTAGGAGTAGCATGGGATCCATGGCGACCCTGTGCCGACC-3'
[SEQ ID NO:20] 5'-CCGAGGCCTAATTATAATGGATTAGCCGGACAGCGCAACGC-3'
[SEQ ID NO:21] 5'-GGTTGGTAGGAGTAGCATGGGATCCATGCACCATCACCATCACCATGC-3'
在另一个例子中，为了将此生物传感器应用于混浊红球菌菌株，使用pCH作为基础质粒构建rppA表达质粒。使用乙酰胺诱导型启动子(G.Roberts等人,FEMSMicrobiol.Lett.222:131-136,2003)，使用引物SEQ ID NO:22和SEQ ID NO:23通过PCR将其扩增。另外，使用引物SEQ ID NO:24和SEQ ID NO:25通过PCR从灰色链霉菌的基因组DNA扩增rppA表达盒，并且使用引物SEQ ID NO:26和SEQ ID NO:25通过PCR扩增rppA表达盒用于另外表达N末端His标签。使用Gibson组件将每一种rppA表达盒、乙酰胺诱导型启动子DNA片段和通过PstI线性化的pCH质粒组装在一起，从而构建两种载体(pCH-rppA和pCH-His-rppA)。将每一种构建的质粒转化至混浊红球菌菌株中，然后将其接种至5mL LB培养基中。将每一种所得菌株在培养箱中于30℃和220rpm下培养18小时，然后将其在5mL LB培养基中传代培养48小时，之后进行采样。可以看出，红色淡黄霉素仅在含有pCH-rppA的菌株中产生(图6G)。
[SEQ ID NO:22] 5'-CTTGATCAGCTTGCATGCCTGCAGAAGCTTTCTAGCAGAAATAATTC-3'
[SEQ ID NO:23] 5'-GTGCATGTGGACTCCCTTTCTCTTATC-3'
[SEQ ID NO:24] 5'-GATAAGAGAAAGGGAGTCCACATGGCGACCCTGTGCCGACC-3'
[SEQ ID NO:25] 5'-GGATCCTCTAGAGTCGACCTGCAGATTAGCCGGACAGCGCAACGC-3'
[SEQ ID NO:26] 5'-GATAAGAGAAAGGGAGTCCACATGCACCATCACCATCACCATGC-3'
实施例2.通过使用RppA生物传感器对具有增加的丙二酰-CoA产生能力的菌株进行高通量筛选
2.1.通过引入大肠杆菌基因组规模的合成对照sRNA文库对具有增加的丙二酰-
CoA产生能力的菌株进行高通量筛选
在构建RppA丙二酰-CoA生物传感器、确认其成功操作以及证明生物传感器的适用性和可扩展性后，将通过本发明人开发的合成对照sRNA技术引入，以通过使用生物传感器来筛选具有增加的丙二酰-CoA产生能力的菌株(KR 10-1575587、US 9388417、EP13735942.8、CN 201380012767.X、KR10-1690780、KR 10-1750855、US 15317939、CN201480081132.X；Na D等人(2013),Nat Biotechnol 31:170-174；Yoo SM,Na D,Lee SY(2013),Nat Protoc 8:1694-1707)。另外，为了包含大肠杆菌中的所有主要基因，将先前构建的大肠杆菌基因组规模的合成对照sRNA文库(包括大肠杆菌K-12W3110菌株中的1,858个基因)引入，并在基因水平上敲低，并且筛选对于增加丙二酰-CoA产生有效的敲低基因靶标。因此，将大肠杆菌基因组规模的合成对照sRNA文库引入含有pTac-5'UTR_Sgr_rppA质粒的大肠杆菌BL21(DE3)菌株中，然后进行高通量筛选(图7)。为了覆盖所有1,858种sRNA，从获得的克隆选择大小等于大于6倍文库大小的11,488个克隆，然后使用机器人自动高通量筛选系统。为此，使用K3菌落挑选器(KBiosystems，巴斯尔登，英国)，并通过机器人(韩国生物科学与生物技术研究所(Korea Research Institute of Bioscience andBiotechnology，KRIBB)；井邑市，韩国)将每一种菌落接种至96孔微型板中的LB培养基(补充有抗生素和0.2mM IPTG)中。K3菌落挑选器照亮琼脂板并将琼脂板成像。然后，内在程序识别并定位菌落，并且指导尖端带有针的机器人手臂挑选所选择的菌落。将接种的细胞在HT-MegaGrow培养箱(Bioneer，韩国)中于30℃和500rpm下培养24小时。培养后，选择具有相对较强信号的231种菌株。将所选择的菌株在含有3mL LB培养基(补充有抗生素和0.2mMIPTG)的14mL一次性Falcon圆底试管中于30℃和250rpm下培养24小时。其中，70种菌株显示出高于对照菌株(不含sRNA的生物传感器菌株)的信号，并对这些菌株中含有的sRNA进行测序(图8)。此时，两次观察到三种sRNA(argB、fabF、nudD)。另外，其中26种菌株与对照菌株相比，信号增加45％或更多。因此，将对应于26种菌株的sRNA载体转化回原始传感器菌株(图9)，并如上所述再次在3mL LB中培养。作为结果，其中最终选择了显示与对照菌株相比信号增加超过70％的14种sRNA。用于增加丙二酰-CoA产生的14种选择的敲低基因靶标列于下表3中。
[表3]
14种最终选择的敲低靶基因的列表
2.2.通过RppA生物传感器确认用于使用FVSEOF算法鉴定的增加丙二酰-CoA产生
的过表达基因靶标
通过RppA生物传感器不仅证明了敲低基因靶标，还证明了过表达基因靶标。此时，使用了利用FVSEOF算法鉴定的基因靶标(Park JM等人(2012),BMC Syst Biol 6:106)。使用大肠杆菌基因组规模的代谢模型iJO1366进行计算机模拟分析(Orth JD等人(2011),MolSyst Biol 7:535)。因此鉴定的9种基因靶标如下：zwf、mdh、fumA、fumB、fumC、serA、serB、serC和tpiA(图10A和图10C)。将每一种基因靶标在大肠杆菌BL21(DE3)中扩增，在trc启动子下插入pTrc99A质粒中，并转化至BL21(DE3)pTac-5'UTR-Sgr_rppA菌株中。以与上文实施例2.1中所述相同的方式将构建的传感器菌株在试管中培养，并在此时测量信号。作为结果，在9种基因靶标中的8种基因靶标(serA、mdh、zwf、tpiA、serB、fumB、serC和fumC)中观察到与对照相比增加的信号(图10B)。
实施例3.使用通过RppA生物传感器筛选的敲低基因靶标增加有用产物的产量
除非另有说明，否则本实施例和以下实施例中使用的烧瓶培养条件如下。将菌落接种至含有10mL LB培养基的试管中，并在培养箱中于37℃和200rpm下培养。将1mL细胞培养物接种至含有50mL改良R/2培养基的带挡板烧瓶中。当已经在37℃和200rpm下生长的菌株达到0.8的OD600值时，用0.5mM IPTG处理菌株以诱导产生，并将其在30℃和200rpm下培养48小时。添加甘油或葡萄糖作为碳源。
3.1.使用选择的敲低基因靶标增加6-甲基水杨酸的产生
在下一步骤中，将上述实施例中选择的且能够增加丙二酰-CoA产生的敲低基因靶标用于产生有用的产物，以证明本发明的生物传感器可以成功地筛选有效的敲低基因靶标。因此，作为第一种产物，试图由基因工程化大肠杆菌产生据报道由灰黄青霉(或展青霉)菌株产生的6-甲基水杨酸(6MSA)。6-甲基水杨酸具有抗菌和抗真菌活性(Dimroth P,Ringelmann E,Lynen F(1976),Eur J Biochem 68:591-596)，并且由一分子乙酰-CoA和三分子丙二酰-CoA通过真菌中发现的6-甲基水杨酸合酶(6MSAS)(I型迭代聚酮合酶)产生(图11)。
用于本发明的6-甲基水杨酸合酶(6MSAS)的序列如下：
6-甲基水杨酸合酶(6MSAS)[SEQ ID NO:123]：
为了构建能够产生6-甲基水杨酸的大肠杆菌菌株，首先使用引物SEQ ID NO:27/SEQ ID NO:28通过反向PCR将pTac15K质粒线性化，然后使用灰黄青霉的基因组DNA作为模板并顺序地使用引物SEQ ID NO:29和SEQ ID NO:30以及SEQ ID NO:31和SEQ ID NO:30扩增编码6-甲基水杨酸合酶的Pg6MSAS基因。使用Gibson组件将两个DNA片段进行组装，从而构建pTac-Pg6MSAS质粒(图12B)。6MSAS酶在其末端含有酰基-载体蛋白结构域，并且需要4'-磷酸泛酰巯基乙胺基转移酶(Sfp)来激活此结构域。
用于本发明的4'-磷酸泛酰巯基乙胺基转移酶(Sfp)的序列如下：
4'-磷酸泛酰巯基乙胺基转移酶(Sfp)[SEQ ID NO:124]：
使用引物SEQ ID NO:32和SEQ ID NO:33通过PCR从枯草芽孢杆菌的基因组DNA扩增编码4’-磷酸泛酰巯基乙胺基转移酶的sfp基因片段，并将所述sfp基因片段在EcoRI位点处插入pTac15K质粒中。将得到的质粒使用引物SEQ ID NO:34和SEQ ID NO:35通过PCR再次扩增，然后将其在SphI位点处插入pTac-Pg6MSAS质粒中，从而构建pTac-Pg6MSAS-sfp质粒(图12A)。将构建的质粒转化至大肠杆菌BL21(DE3)菌株中，并通过SDS-PAGE检查酶表达(图12C)。然后，菌株的烧瓶培养在不同浓度的葡萄糖或甘油的存在下在改良R/2培养基中进行，并且作为结果，可以看出当添加100g/L的甘油时，产生了4.7mg/L的6-甲基水杨酸(图13A)。通过LC-MS确认所产生的6-甲基水杨酸的真实性(图13B和图13C)。在引入上文实施例中选择的sRNA之前，为了检验不同大肠杆菌菌株中6-甲基水杨酸的产量，将pTac-Pg6MSAS-sfp质粒引入上表1中所示的16种大肠杆菌菌株中，然后在3mL改良R/2培养基中进行每一种菌株的试管规模的培养。在这些菌株中，选择产生超过1mg/L 6MSA的6种菌株(NM522、BL21、S17-1、JM110、HB101和XL1-蓝)(图13D)。然后，将14种选择的sRNA引入每种选择的菌株中，从而构建总共84种菌株，并对每种构建的菌株进行试管规模的培养。作为结果，pabA-敲低BL21(DE3)菌株产生最大量的6-甲基水杨酸(6.1mg/L；图14)。在相同的培养基条件下对此菌株进一步进行烧瓶培养(50mL)，结果产生了8.0mg/L的6-甲基水杨酸(图13E)。
[SEQ ID NO:27] 5'-GCTGAGAAGCTTGCCAAATAATGGATCCTCTAGAGTCGACCTG-3'
[SEQ ID NO:28] 5'-CTGGGGATGTTTTCCCAGAGGGGTATGTAGAAGTTGCAGCGGAATGCATGAATTCTGTTTCCTGTGTGAAATTG-3'
[SEQ ID NO:29] 5'-ACAGTGAATATGAATTCTCCAACG-3'
[SEQ ID NO:30] 5'-ATTATTTGGCAAGCTTCTCAGC-3'
[SEQ ID NO:31] 5'-CTCTGGGAAAACATCCCCAGCACCAGTCGGAACCCCTGGGACTGAGTACAGTGAATATGAATTCTCCAACG-3'
[SEQ ID NO:32] 5'-TAATAAGAATTCATGAAGATTTACGGAATTTATATG-3'
[SEQ ID NO:33] 5'-TTATTAGAATTCTTATAAAAGCTCTTCGTACGAG-3'
[SEQ ID NO:34] 5'-CTAGAGTCGACCTGCAGGCATGCCACTCCCGTTCTGGATAATG-3'
[SEQ ID NO:35] 5'-CAAAACAGCCAAGCTTGCATGC-3'
为了进一步优化产生6-甲基水杨酸的菌株，用质粒(pTac-Pg6MSAS)仅表达具有长度(5.3kb)的Pg6MSAS基因，并且使用菌株BAP1表达sfp(Pfeifer BA,Admiraal SJ,GramajoH,Cane DE,Khosla C(2001),Science291:1790-1792)，所述菌株BAP1是将sfp插入BL21(DE3)菌株的基因组中的菌株。烧瓶培养的结果显示了增加的6-甲基水杨酸产量为17.6mg/L，并且从SDS-PAGE分析的结果来看，这被认为是由于改善的4'-磷酸泛酰巯基乙胺基转移酶表达和未激活的6-甲基水杨酸合酶的降低的比例(图12C)。因此，当将pabA-敲低sRNA引入大肠杆菌BAP1 pTac-Pg6MSAS菌株中时，可以产生23.9mg/L的量的6-甲基水杨酸，这对应于约35.8％的增加(图13F)。当进行此菌株的补料分批发酵时，在约27小时后可以获得97.8mg/L的增加的6-甲基水杨酸浓度(图13G)。在50g/L甘油作为初始碳源的存在下，在6.6L发酵罐(BioFlo320，Eppendofr，德国)中的1.9L改良R/2培养基中进行补料分批发酵。将菌落接种至含有10mL LB培养基的试管中，然后在37℃和200rpm下培养一天。接着，将细胞接种至两个带挡板的烧瓶中，每个烧瓶含有50mL改良R/2培养基，此时使用50g/L甘油作为碳源。进行烧瓶培养约9小时直至OD600值达到约2，然后将细胞接种至发酵罐中。通过28％(v/v)氨溶液将培养物pH维持在6.8，并且通过以2L/min供应空气和自动控制搅拌速度为1000rpm并且通过增加氧流速，将溶解氧(DO)浓度控制在40％的空气饱和下。使用pH-stat进料策略以提供营养。当pH高于6.86时，自动添加进料溶液。进料溶液每升含有以下组分：800g甘油、6mL痕量金属溶液和12g MgSO4·7H2O。当接种后的OD600值达到2至3时，用0.5mMIPTG诱导异源蛋白表达。
为了进一步增加6-甲基水杨酸的产量，试图增加先前选择的三种FVSEOF基因靶标(P<0.05；zwf、mdh、serA；图10)和以下五种酶的表达：谷氨酸棒杆菌乙酰-CoA羧化酶(AccBC和AccD1)、大肠杆菌甘油醛3-磷酸脱氢酶(GapA)、大肠杆菌磷酸甘油酸激酶(Pgk)、大肠杆菌乙酰-CoA合成酶(Acs)和大肠杆菌丙酮酸脱氢酶(PDH：AceEF和Lpd)。此时，插入相应基因片段并克隆至pBBR1TaC质粒中。pBBR1TaC质粒是由pBBR1MCS构建的基于tac启动子的表达质粒。对于构建pBBR1TaC，首先使用引物SEQ ID NO:147和SEQ ID NO:148通过反向PCR扩增pBBR1MCS，使用pTac15K作为模板以及引物SEQ ID NO:149和SEQ ID NO:150扩增由tac启动子、MCS和rrnBT1T2构成的DNA片段，并使用Gibson组件将所述DNA片段克隆至质粒中。接着，为了将来自上文实施例2.2中构建的pTrc99A-zwf、pTrc99A-mdh和pTrc99A-serA质粒的基因转移至pBBR1TaC，进行以下实验。首先，使用引物SEQ ID NO:151和SEQ ID NO:152扩增zwf、mdh和serA各个基因，使用引物SEQ ID NO:75和SEQ ID NO:76扩增pBBR1TaC质粒，然后进行Gibson组装，从而构建pBBR1-zwf、pBBR1-mdh和pBBR1-serA质粒。谷氨酸棒杆菌乙酰-CoA羧化酶由α亚基AccBC和β亚基AccD1构成，并且将相应基因各自克隆至pBBR1TaC中，从而构建pBBR1-accBC和pBBR1-accD1。此时，使用引物SEQ ID NO:153和SEQ ID NO:154扩增accBC基因，并且使用引物SEQ ID NO:155和SEQ ID NO:156扩增accD1基因。通过Gibson组件将这两个基因片段均插入使用引物SEQ ID NO:9和SEQ ID NO:10扩增的pBBR1TaC质粒中。在这两个构建的质粒中使用pBBR1-accD1作为模板，使用引物SEQ ID NO:157和SEQ IDNO:158扩增含有tac启动子和accD1的DNA片段，然后将所述DNA片段通过Gibson组件插入通过HindIII限制酶消化的pBBR1-accBC质粒中，从而构建pBBR1-accBCD1质粒。大肠杆菌丙酮酸脱氢酶(PDH)由亚基E1(AceE)、E2(AceF)和E3(Lpd)构成。首先，将aceEF基因片段和lpd基因片段克隆至pBBR1TaC质粒中，从而构建pBBR1-aceEF和pBBR1-lpd质粒。使用引物SEQ IDNO:159和SEQ ID NO:160扩增基因片段aceEF，并且使用引物SEQ ID NO:161和SEQ ID NO:162扩增lpd。通过Gibson组件将这两个基因片段各自插入用于构建pBBR1-accBC和pBBR1-accD1质粒的线性化pBBR1TaC质粒中。在这两个构建的质粒中使用pBBR1-lpd作为模板，使用引物SEQ ID NO:163和SEQ ID NO:164扩增包含tac启动子和lpd的基因片段，然后将所述基因片段插入通过SalI限制酶消化的pBBR1-aceEF质粒中，从而构建pBBR1-aceEF-lpd质粒。对于构建分别含有大肠杆菌甘油醛3-磷酸脱氢酶(GapA)、大肠杆菌磷酸甘油酸激酶(Pgk)和大肠杆菌乙酰-CoA合成酶(Acs)的pBBR1-gapA、pBBR1-pgk和pBBR1-acs质粒，使用引物SEQ ID NO:165和SEQ ID NO:166、SEQ ID NO:167和SEQ ID NO:168以及SEQ ID NO:169和SEQ ID NO:170通过PCR扩增gapA、pgk和acs基因片段，然后将其每一种通过Gibson组件插入线性化pBBR1TaC质粒中。
[SEQ ID NO:147] 5'-GACGGATGGCCTTTTACTAGTGCCTGGGGTGCCTAATGAG-3'
[SEQ ID NO:148] 5'-CGATGATTAATTGTCAACTGCTACGCCTGAATAAGTGATAATAAG-3'
[SEQ ID NO:149] 5'-GTTGACAATTAATCATCGGCTC-3'
[SEQ ID NO:150] 5'-ACTAGTAAAAGGCCATCCGTCAGGATG-3'
[SEQ ID NO:151] 5'-GTTGACAATTAATCATCGGC-3'
[SEQ ID NO:152] 5'-CGTTTCACTTCTGAGTTCGG-3'
[SEQ ID NO:153] 5'-CAATTTCACACAGGAAACAGAATTCGTGTCAGTCGAGACTAGGAAG-3'
[SEQ ID NO:154] 5'-CTCTAGAGGATCCCCGGGTACCATTACTTGATCTCGAGGAGAAC-3'
[SEQ ID NO:155] 5'-CAATTTCACACAGGAAACAGAATTCATGACCATTTCCTCACCTTTG-3'
[SEQ ID NO:156] 5'-CTCTAGAGGATCCCCGGGTACCATTACAGTGGCATGTTGCCGTG-3'
[SEQ ID NO:157] 5'-GAGTCGACCTGCAGGCATGCATTGACAATTAATCATCGGCTCG-3'
[SEQ ID NO:158] 5'-CTCATCCGCCAAAACAGCCAAGCTT-3'
[SEQ ID NO:159] 5'-CAATTTCACACAGGAAACAGAATTCATGTCAGAACGTTTCCCAAATG-3'
[SEQ ID NO:160] 5'-CTCTAGAGGATCCCCGGGTACCATTACATCACCAGACGGCGAATG-3'
[SEQ ID NO:161] 5'-CAATTTCACACAGGAAACAGAATTCATGAGTACTGAAATCAAAACTC-3'
[SEQ ID NO:162] 5'-CTCTAGAGGATCCCCGGGTACCATTACTTCTTCTTCGCTTTCGG-3'
[SEQ ID NO:163] 5'-GTACCCGGGGATCCTCTAGAGTTGACAATTAATCATCGGCTCG-3'
[SEQ ID NO:164] 5'-CAAGCTTGCATGCCTGCAGGTCGAC-3'
[SEQ ID NO:165] 5'-CAATTTCACACAGGAAACAGAATTCATGACTATCAAAGTAGGTATCAAC-3'
[SEQ ID NO:166] 5'-CTCTAGAGGATCCCCGGGTACCATTATTTGGAGATGTGAGCGATC-3'
[SEQ ID NO:167] 5'-CAATTTCACACAGGAAACAGAATTCATGTCTGTAATTAAGATGACCGATC-3'
[SEQ ID NO:168] 5'-CTCTAGAGGATCCCCGGGTACCATTACTTCTTAGCGCGCTCTTCG-3'
[SEQ ID NO:169] 5'-CAATTTCACACAGGAAACAGAATTCATGAGCCAAATTCACAAACAC-3'
[SEQ ID NO:170] 5'-CTCTAGAGGATCCCCGGGTACCATTACGATGGCATCGCGATAG-3'
将如上所述构建的8种质粒各自转化至显示最高产量的菌株大肠杆菌BAP1 pTac-Pg6MSAS pWAS-抗-pabA中，然后对菌株进行烧瓶培养。烧瓶培养的结果显示在图15A中。当谷氨酸棒杆菌乙酰-CoA羧化酶过表达时，6-甲基水杨酸的产量增加至63.6mg/L。进行相应菌株的补料分批发酵，结果如图15B所示，来自甘油的6-甲基水杨酸的产量增加至440.3mg/L±30.2mg/L。
3.2.使用选择的敲低基因靶标增加芦荟酮的产生
作为第二种产物，测试了由掌叶大黄(Abe I,Utsumi Y,Oguro S,Noguchi H(2004),FEBS Lett 562:171-176)或木立芦荟(Mizuuchi Y等人(2009),FEBS J 276:2391-2401)III型聚酮合酶产生的芦荟酮。芦荟酮是芦荟苦素的前体，由于其美白效果而广泛用于化妆品领域。然而，尚未报道由芦荟酮产生芦荟苦素的生物合成途径。据报道，芦荟酮由一分子乙酰-CoA和六分子丙二酰-CoA(图16)产生，并且通过掌叶大黄芦荟酮合酶(ALS)或木立芦荟芦荟酮合酶(PKS3)产生。
掌叶大黄芦荟酮合酶(ALS)的序列如下。
掌叶大黄芦荟酮合酶(ALS)[SEQ ID NO:125]：
另外，木立芦荟芦荟酮合酶(PKS3)的序列如下。
木立芦荟芦荟酮合酶PKS3[SEQ ID NO:126]：
对应于这两种酶的基因均通过Integrated DNA Technologies公司(美国)来合成，然后将其通过Gibson组件在NcoI位点处插入pCDFDuet-1(Novagen)质粒中。将这两种构建的质粒pCDF-RpALS和pCDF-AaPKS3各自转化至大肠杆菌BL21(DE3)菌株中，然后通过SDS-PAGE分析异源酶的表达(图17A)。进行菌株的烧瓶培养，结果由20g/L葡萄糖产生20.5mg/L和4.7mg/L芦荟酮(图17B)。因此，将质粒pCDF-RpALS用于以下实验。通过LC-MS和MS/MS分析证实了产生的芦荟酮的真实性(图17C)。接着，如上文实施例3.1中所述，将上述实施例中选择的14种sRNA引入各种大肠杆菌菌株中。为此，将每种sRNA引入BL21(DE3)菌株和W3110(DE3)菌株中，从而构建28种菌株。构建的菌株的试管规模的培养的结果显示在图17F中。此时，pabA敲低BL21(DE3)菌株显示出最高的芦荟酮产量(18.5mg/L)，烧瓶培养中菌株的芦荟酮产量为27.1mg/L，这对应于与对照菌株(不含sRNA)的芦荟酮产量相比32.2％的增加(图17D和图17E)。
为了进一步增加芦荟酮的产生，增加了三种FVSEOF基因靶标(zwf、mdh和serA；图10)和以下用于增加6-甲基水杨酸产生的五种酶：谷氨酸棒杆菌乙酰-CoA羧化酶(AccBC和AccD1)、大肠杆菌甘油醛3-磷酸脱氢酶(GapA)、大肠杆菌磷酸甘油酸激酶(Pgk)、大肠杆菌乙酰-CoA合成酶(Acs)和大肠杆菌丙酮酸脱氢酶(PDH：AceEF和Lpd)。为此，将含有相应基因的八种质粒pBBR1-zwf、pBBR1-mdh、pBBR1-serA、pBBR1-accBCD1、pBBR1-gapA、pBBR1-pgk、pBBR1-acs和pBBR1-aceEF-lpd各自转化至显示最高芦荟酮产量的菌株(具有pCDF-RpALS和pWAS-抗-pabA的大肠杆菌BL21(DE3))中，然后对所得菌株进行烧瓶培养。烧瓶培养的结果显示在图18中。当谷氨酸棒杆菌乙酰-CoA羧化酶过表达时，芦荟酮的产量增加至30.9mg/L。在这种情况下，将20g/L葡萄糖用作碳源。
3.3.使用选择的敲低基因靶标增加白藜芦醇的产生
作为第三种产物，选择了在植物中产生的白藜芦醇。白藜芦醇是属于基于苯丙素类的天然产物中的芪类化合物家族的化合物，它是具有抗氧化、抗老化和抗癌等作用的非常有用的天然产物。白藜芦醇由一分子对香豆酰-CoA和三分子丙二酰-CoA产生(图19)。因此，构建了由简单的碳源产生对香豆酸的菌株。为此，使用先前构建的过度产生酪氨酸的菌株(Kim B,Binkley R,Kim HU,Lee SY(2018),Biotechnol Bioeng)。此菌株(BTY5.13)含有基于pTac15K质粒的质粒(pTY13)，所述质粒在基于大肠杆菌BL21(DE3)的tyrR、tyrP敲低菌株(BTY5)中过表达运动发酵单胞菌(Zymomonas mobilis)tyrC、大肠杆菌aroGfbr和aroL。为了将酪氨酸转化为对香豆酸，应表达酪氨酸氨裂解酶(TAL)。因此，使用引物SEQ ID NO:36和SEQ ID NO:37从西班牙糖丝菌的基因组DNA扩增编码酪氨酸氨裂解酶的基因SeTAL，然后使用引物SEQ ID NO:38和SEQ ID NO:39通过PCR将pTrc99A质粒线性化。接着，使用Gibson组件将这两个DNA片段进行组装，从而构建质粒pTrc-SeTAL。
用于本发明的酪氨酸氨裂解酶的序列如下。
酪氨酸氨裂解酶(TAL)[SEQ ID NO:127]：
为了优化酪氨酸氨裂解酶的表达，将His标签和硫氧还蛋白标签(TrxA)均连接至N末端。对于His标签的附接，使用西班牙糖丝菌的基因组DNA以及引物SEQ ID NO:40和SEQID NO:37通过PCR扩增His-SeTAL，并且使用引物SEQ ID NO:38和SEQ ID NO:41通过反向PCR将pTrc-SeTAL作为模板线性化。使用Gibson组件将这两个DNA片段进行组装，从而构建pTrc-HisTAL。使用大肠杆菌W3110的基因组DNA作为模板以及引物SEQ ID NO:42和SEQ IDNO:43通过PCR扩增trxA基因，并且使用西班牙糖丝菌的基因组DNA作为模板以及引物SEQID NO:44和SEQ ID NO:45通过PCR扩增TrxA-TAL。使用引物SEQ ID NO:46和SEQ ID NO:37，通过PCR对这两个DNA片段进行延伸PCR以获得单个DNA片段。通过Gibson组件将获得的DNA片段与用于构建pTrc-HisTAL的线性化pTrc-SeTAL DNA片段组装，从而构建pTrc-TrxTAL。将三种构建的质粒pTrc-SeTAL、pTrc-HisTAL和pTrc-TrxTAL均转化至BTY5.13菌株中，然后将其接种至含有补充有20g/L葡萄糖的50mL改良MR培养基的烧瓶中，并且将菌株在30℃和200rpm下培养。当细胞达到约0.8的OD600值时，用1mM IPTG处理细胞，然后培养36小时。改良MR培养基每升含有以下组分：6.67g KH2PO4、4g(NH4)2HPO4、0.8g柠檬酸、0.8g MgSO4·7H2O、5mL痕量金属溶液、2g酵母提取物和15g(NH4)2SO4。当表达His-TAL时，可以获得最高水平的对香豆酸产量(0.41g/L)(图20A)。因此，为了通过将pTrc-HisTAL与pTY13质粒组装来构建单个质粒，使用引物SEQ ID NO:47和SEQ ID NO:48扩增HisTAL部分，然后将其通过Gibson组件在NheI位点处插入pTY13质粒中，从而构建pTY13-HisTAL质粒。将构建的质粒转化至BTY5菌株中，然后对菌株进行烧瓶培养。作为结果，由20g/L甘油产生了0.35g/L对香豆酸(图20A)。
[SEQ ID NO:36] 5'-GAATTGTGAGCGGATAACAAAGACCGAGGAAAAGGAGCATCGCAAATGACGCAGGTCGTGGAACGTC-3'
[SEQ ID NO:37] 5'-TAGAGGATCCCCGGGTACTCATCCGAAATCCTTCCCGTC-3'
[SEQ ID NO:38] 5'-GTACCCGGGGATCCTCTAG-3'
[SEQ ID NO:39] 5'-TTGTTATCCGCTCACAATTC-3'
[SEQ ID NO:40] 5'-GAGGAAAAGGAGCATCGCAAATGCACCATCATCATCATCATACGCAGGTCGTGGAACGTC-3'
[SEQ ID NO:41] 5'-TTGCGATGCTCCTTTTCCTC-3'
[SEQ ID NO:42] 5'-ATGAGCGATAAAATTATTCACCTG-3'
[SEQ ID NO:43] 5'-CGCCAGGTTAGCGTCGAGG-3'
[SEQ ID NO:44] 5'-CCTCGACGCTAACCTGGCGACGCAGGTCGTGGAACGTC-3'
[SEQ ID NO:45] 5'-TCATCCGAAATCCTTCCCGTC-3'
[SEQ ID NO:46] 5'-AGACCGAGGAAAAGGAGCATCGCAAATGAGCGATAAAATTATTCACCTG-3'
[SEQ ID NO:47] 5'-GTAAGCCAGTATACACTCCGGACTGCACGGTGCACCAATG-3'
[SEQ ID NO:48] 5'-CTGTTGGGCGCCATCTCCTTGTGTAGAAACGCAAAAAGGCCATC-3'
在成功地如上所述构建产生对香豆酸的菌株后，构建了从对香豆酸开始的下游白藜芦醇生物合成途径，所述生物合成途径由拟南芥突变4-香豆酸:CoA连接酶(4CL)1(At4CL1m)和葡萄芪合酶(STS)构成。首先，使用引物SEQ ID NO:49和SEQ ID NO:50从拟南芥cDNA扩增At4CL1基因，然后将其通过Gibson组件在EcoRI和KpnI位点处插入pTac15K质粒中，从而构建pTac-At4CL1质粒。接着，通过连续三轮定点诱变来构建pTac-At4CL1m(At4CL1m；I250L/N404K/I461V)，以增强底物对于对香豆酰-CoA的特异性。此时，对于连续三轮定点诱变，使用引物对SEQ ID NO:51和SEQ ID NO:52、SEQ ID NO:53和SEQ ID NO:54以及SEQ ID NO:55和SEQ ID NO:56。接着，分别使用引物对SEQ ID NO:57和SEQ ID NO:58以及SEQ ID NO:59和SEQ ID NO:60通过PCR扩增分别编码4-香豆酸:CoA连接酶3和4-香豆酸:CoA连接酶4的来自拟南芥的At4CL3和At4CL4基因，并且使用引物对SEQ ID NO:61和SEQID NO:62通过PCR扩增编码4-香豆酸:CoA连接酶的来自天蓝色链霉菌的Sc4CL基因。以与上述相同的方式克隆扩增的基因，从而构建pTac-At4CL3、pTac-At4CL4和pTac-Sc4CL。对于pTac-Sc4CL，使用引物对SEQ ID NO:63和SEQ ID NO:64进行一轮定点诱变，以增强底物对于对香豆酰-CoA的特异性(Kaneko M,Ohnishi Y,Horinouchi S(2003),J Bacteriol185:20-27)，从而构建pTac-Sc4CLm(Sc4CLm；A294G/A318G)质粒。由Integrated DNATechnologies公司来合成编码葡萄芪合酶的STS基因，并使用引物对SEQ ID NO:65和SEQID NO:66通过PCR扩增所述STS基因，然后使用Gibson组件将所述STS基因在EcoRI和KpnI位点处插入pTac15K质粒中，从而构建pTac-VvSTS质粒。
用于本发明的拟南芥4-香豆酸:CoA连接酶1的序列如下。
拟南芥4-香豆酸:CoA连接酶1[SEQ ID NO:128]：
用于本发明的拟南芥4-香豆酸:CoA连接酶3的序列如下。
拟南芥4-香豆酸:CoA连接酶3[SEQ ID NO:129]：
用于本发明的拟南芥4-香豆酸:CoA连接酶4的序列如下。
拟南芥4-香豆酸:CoA连接酶4[SEQ ID NO:130]：
用于本发明的天蓝色链霉菌4-香豆酸:CoA连接酶的序列如下。
天蓝色链霉菌4-香豆酸:CoA连接酶[SEQ ID NO:131]：
用于本发明的芪合酶的序列如下。
芪合酶[SEQ ID NO:132]：
将每一种上述质粒转化至BL21(DE3)菌株中，然后使用SDS-PAGE分析所得菌株中异源酶的表达。然后，为了将4CL基因与STS基因组装成单个质粒，进行以下操作。首先，使用引物SEQ ID NO:47和SEQ ID NO:48通过PCR从pTac-VvSTS扩增用于芪合酶表达的DNA片段(含有tac启动子)，然后将其通过Gibson组件在NheI位点处插入pTac-At4CL3、pTac-At4CL4l和pTac-Sc4CLm质粒中，从而构建pTac-VvSTS-At4CL3、pTac-VvSTS-At4CL4和pTac-VvSTS-Sc4CLm质粒。对于At4CL1m，使用引物SEQ ID NO:67和SEQ ID NO:68通过PCR从pTac-At4CL1m扩增用于4-香豆酸:CoA连接酶表达的DNA片段，然后将所述DNA片段在PvuII位点处插入pTac-VvSTS质粒中，从而构建pTac-VvSTS-At4CL1m质粒。将构建的质粒pTac-VvSTS-At4CL1m、pTac-VvSTS-At4CL3、pTac-VvSTS-At4CL4和pTac-VvSTS-Sc4CLm转化至BL21(DE3)菌株中，并且在补充有2mM对香豆酸的培养基中对菌株进行烧瓶培养。烧瓶培养的结果显示在图20C中。此时，BL21(DE3)pTac-VvSTS-At4CL1m菌株显示出最高的白藜芦醇产量(18.0mg/L)。如下事实是丙二酰-CoA为瓶颈的间接证据：即使以2mM(328.1mg/L)的浓度添加对香豆酸，也能产生这个量的白藜芦醇。在本发明中，在通过增加丙二酰-CoA浓度根据此假设解决问题之前，通过以各种方式调节用于白藜芦醇产生的外源基因的表达来解决此问题，然后解决丙二酰-CoA。因此，为了在单个操纵子中表达At4CL1m基因和STS基因，使用引物SEQ ID NO:69和SEQ ID NO:70通过PCR扩增STS基因，然后将所述STS基因在SphI位点处插入pTac-At4CL1m质粒中，从而构建pTac-At4CL1m-opr-VvSTS质粒。另外，为了促进通过将这两种酶表达为单个融合酶的底物转化，使用引物对SEQ ID NO:71和SEQ ID NO:72通过PCR扩增At4CL1m基因，然后将所述At4CL1m基因在NdeI位点处插入pTac-VvSTS质粒中，从而构建pTac-At4CL1m-fus-VvSTS质粒。对于融合蛋白的表达，使用甘氨酸-丝氨酸接头(Gly-Gly-Gly-Ser)。然而，这两种策略都没有产生更高水平的白藜芦醇(图20D)。因此，选择pTac-VvSTS-At4CL1m作为最终质粒，此质粒与pTY13-HisTAL不相容(这两个质粒都具有p15A复制起点)，因此将此质粒转移至上述pTacCDFS质粒中。为此，使用引物SEQ ID NO:73和SEQ ID NO:74通过PCR扩增STS表达盒，然后通过Gibson组件将所述表达盒与使用引物SEQ ID NO:75和SEQ ID NO:76线性化的pTacCDFS质粒组装。用PstI限制酶处理构建的质粒，然后通过Gibson组件或T4连接与使用引物SEQ ID NO:75和SEQ ID NO:76扩增的At4CL1m表达盒组装，从而构建单个质粒(pTacCDF-VvSTS-At4CL1m)。将构建的质粒转化至BL21(DE3)菌株中，然后对菌株进行烧瓶培养，从而产生21.2mg/L白藜芦醇(图20D)。然而，这是提供2mM对香豆酸的结果。因此，为了直接由甘油产生白藜芦醇，将pTacCDF-VvSTS-At4CL1m质粒转化至BTY5 pTY13-HisTAL菌株中，并对菌株进行烧瓶培养，作为结果，由20g/L甘油产生12.4mg/L白藜芦醇。通过LC-MS确认产生的白藜芦醇的真实性(图21)。
[SEQ ID NO:49] 5'-CAATTTCACACAGGAAACAGACATATGGCGCCACAAGAACAAGC-3’
[SEQ ID NO:50] 5'-CTCTAGAGGATCCCCGGGTACCATTACAATCCATTTGCTAGTTTTG-3'
[SEQ ID NO:51] 5'-CACAGCGATGACGTCCTACTCTGTGTTTTG-3'
[SEQ ID NO:52] 5'-CAAAACACAGAGTAGGACGTCATCGCTGTG-3'
[SEQ ID NO:53] 5'-CTCTTTCGAGGAAACAACCCGGTGAG-3'
[SEQ ID NO:54] 5'-CTCACCGGGTTGTTTCCTCGAAAGAG-3'
[SEQ ID NO:55] 5'-GATTGAAAGAACTTGTCAAGTATAAAGG-3'
[SEQ ID NO:56] 5'-CCTTTATACTTGACAAGTTCTTTCAATC-3'
[SEQ ID NO:57] 5'-CAATTTCACACAGGAAACAGACATATGATCACTGCAGCTCTACAC-3'
[SEQ ID NO:58] 5'-CTCTAGAGGATCCCCGGGTACCATTAACAAAGCTTAGCTTTGAGG-3'
[SEQ ID NO:59] 5'-CAATTTCACACAGGAAACAGACATATGGTGCTCCAACAACAAACG-3'
[SEQ ID NO:60] 5'-CTCTAGAGGATCCCCGGGTACCATTATTTAGAGCACATGGTTTCC-3'
[SEQ ID NO:61] 5'-CAATTTCACACAGGAAACAGACATATGTTCCGCAGCGAGTACGC-3'
[SEQ ID NO:62] 5'-CTCTAGAGGATCCCCGGGTACCATTATCGCGGCTCCCTGAGCTGTC-3'
[SEQ ID NO:63] 5'-GTACATCGTCAGCGGCGCCGCCCCGCTCGACG-3'
[SEQ ID NO:64] 5'-CGTCGAGCGGGGCGGCGCCGCTGACGATGTAC-3'
[SEQ ID NO:65] 5'-CAATTTCACACAGGAAACAGACATATGGCAAGTGTCGAGGAATTC-3'
[SEQ ID NO:66] 5'-CTCTAGAGGATCCCCGGGTACCATTAATTGGTAACCATCGGAATGG-3'
[SEQ ID NO:67] 5'-GTAAGCCAGTATACACTCCGGACTGCACGGTGCACCAATG-3'
[SEQ ID NO:68] 5'-CTGTTGGGCGCCATCTCCTTGTGTAGAAACGCAAAAAGGCCATC-3'
[SEQ ID NO:69] 5'-GAGTCGACCTGCAGGCATGCAATTTCACACAGGAAACAGA-3'
[SEQ ID NO:70] 5'-CAAAACAGCCAAGCTTGCATG-3'
[SEQ ID NO:71] 5'-TTTCACACAGGAAACAGACATATG-3'
[SEQ ID NO:72] 5'-GAATTCCTCGACACTTGCCATACTACCACCACCCAATCCATTTGCTAGTTTTG-3'
[SEQ ID NO:73] 5'-GTTGACAATTAATCATCGGC-3'
[SEQ ID NO:74] 5'-CGTTTCACTTCTGAGTTCGG-3'
[SEQ ID NO:75] 5'-CCGAACTCAGAAGTGAAACG-3'
[SEQ ID NO:76] 5'-GCCGATGATTAATTGTCAAC-3'
将上文实施例2.1中选择的14种sRNA引入如上所述构建的最终的产白藜芦醇菌株中，并对菌株进行烧瓶培养。烧瓶培养的结果显示在图20E中。作为结果，当敲低pabA时，由甘油产生51.8mg/L白藜芦醇，并且与对照菌株(不具有sRNA的菌株)相比，此产量增加了4.2倍。另外，值得注意的是，pabA是敲低靶标，其显示出如上所述的6-甲基水杨酸和芦荟酮的产量的最大增加。接着，选择能够使白藜芦醇滴度增加超过2.5倍的六种敲低基因靶标，并且构建sRNA质粒，其中每一种质粒都含有基因靶标中的两种基因靶标的组合。将每一种sRNA质粒转化至产生白藜芦醇的菌株中，然后对每一种所得菌株进行烧瓶培养。烧瓶培养的结果显示在图20中。然而，未实现白藜芦醇产量的额外增加，因为当yfiD和purB同时被敲低时最高的白藜芦醇产量为50.0mg/L。
此处，如下构建用于组合双敲低的sRNA质粒。使用引物SEQ ID NO:77和SEQ IDNO:78从亲本质粒扩增有待插入的对合成对照sRNA进行编码的DNA片段，然后通过Gibson组件将所述DNA片段与使用引物SEQ ID NO:79和SEQ ID NO:80通过PCR线性化的含有合成对照sRNA的亲本质粒组装。
[SEQ ID NO:77] 5'-GAATTTTAACAAAATATTAACGAATTCTAACACCGTGCGTG-3'
[SEQ ID NO:78] 5'-GTGCCACCTAAATTGTAAGCGGCGAATTGGGTACCTATAAAC-3'
[SEQ ID NO:79] 5'-GCTTACAATTTAGGTGGCAC-3'
[SEQ ID NO:80] 5'-GTTAATATTTTGTTAAAATTCGCG-3'
3.4.使用选择的敲低基因靶标增加柚皮素的产生
作为第四种产物，选择了在植物中产生的柚皮素。柚皮素是许多药理学上有用的产品的常见前体，其属于基于苯丙素类的天然产物中的类黄酮家族。另外，据报道柚皮素具有抗阿尔茨海默病、抗癌、抗氧化和抗真菌活性。柚皮素由一分子对香豆酰-CoA和三分子丙二酰-CoA(如白藜芦醇)产生(图21)。由于在上文实施例3.3中构建了产生对香豆酸的菌株，因此在本实施例中构建了下游柚皮素生物合成途径。此处，使用拟南芥突变4-香豆酸:CoA连接酶1(At4CL1m)进行从对香豆酸到对香豆酰-CoA的转化，使用矮牵牛查耳酮合酶(CHS)进行从对香豆酰-CoA和丙二酰-CoA到柚皮素查耳酮的转化，并且使用拟南芥查耳酮异构酶(CHI)进行从柚皮素查耳酮到柚皮素的转化。
用于本发明的查耳酮合酶(CHS)的序列如下。
查耳酮合酶(CHS)[SEQ ID NO:133]：
用于本发明的查耳酮异构酶(CHI)的序列如下。
查耳酮异构酶(CHI)[SEQ ID NO:134]：
使用拟南芥cDNA作为模板以及引物SEQ ID NO:81和SEQ ID NO:82通过PCR扩增编码突变4-香豆酸:CoA连接酶1的At4CL1m基因，并且使用拟南芥cDNA作为模板以及引物SEQID NO:83和SEQ ID NO:84扩增编码查耳酮合酶的AtCHI基因。然后，使用引物SEQ ID NO:81和SEQ ID NO:84通过延伸PCR将这两个DNA片段组装成单个DNA片段，随后将所述单个DNA片段在KpnI和BamHI位点处插入pTrc99A质粒中。另外，使用DNA片段(由Integrated DNATechnologies公司合成)作为模板以及引物SEQ ID NO:85和SEQ ID NO:86的引物通过PCR扩增编码查耳酮合酶的PhCHS基因，然后将所述PhCHS基因在BamHI和XbaI位点处插入上述构建的质粒中，从而构建pTrc-At4CL1m-AtCHI-PsCHS质粒。由于与上文实施例3.3中所述相同的原因，用NcoI和PstI限制酶消化质粒pTrc-At4CL1m-AtCHI-PsCHS以使得其与产生对香豆酸的菌株中的质粒相容。通过Gibson组件将产生的DNA片段与使用引物SEQ ID NO:87和SEQ ID NO:88通过反向PCR线性化的pTrcCDFS质粒组装，从而构建pTrcCDF-At4CL1m-AtCHI-PhCHS质粒。
[SEQ ID NO:81] 5'-AGACAGGGTACCATGGCGCCACAAGAACAAG-3'
[SEQ ID NO:82] 5'-ATGTATATCTCCTTCTTAAAGTTAATTACAATCCATTTGCTAGTTTTGCCC-3'
[SEQ ID NO:83] 5'-TTAACTTTAAGAAGGAGATATACATATGTCTTCATCCAACGCCTGC-3'
[SEQ ID NO:84] 5'-AGACAGGGATCCTCAGTTCTCTTTGGCTAGTTTTTCC-3'
[SEQ ID NO:85] 5'-AGAGAGGATCCATAACAATTCCCCATCTTAG-3'
[SEQ ID NO:86] 5'-AGAGATCTAGATTAGGTAGCCACACTATGCAGAACC-3'
[SEQ ID NO:87] 5'-GAAACTGCTTGTTCTTGTGGCGCCATGGTCTGTTTCCTGTGTG-3'
[SEQ ID NO:88] 5'-CTAATCTAGAGTCGACCTGCAGGCATGCAAGCTTG-3'
将如上所述构建的质粒pTrcCDF-At4CL1m-AtCHI-PhCHS引入BTY5pTY13-HisTAL菌株中，然后对菌株进行烧瓶培养。作为结果，分别由葡萄糖和甘油产生37.2mg/L和64.5mg/L柚皮素(图23A)。因此，以下关于柚皮素产生的实验均使用甘油进行。通过SDS-PAGE分析菌株中异源酶的表达(图23B)，并通过LC-MS确认产生的柚皮素的真实性(图24)。将上文实施例2.1中选择的14种sRNA引入构建的产柚皮素菌株中，并对每一种所得菌株进行烧瓶培养。烧瓶培养的结果显示在图25A中。此时，fadR敲低菌株显示出最高的柚皮素产量(92.3mg/L)，与对照菌株(不具有sRNA的产柚皮素菌株)中的柚皮素产量相比，柚皮素产量增加43％。另外，将与对照相比显示出超过15％增加的敲低靶标进行组合并进行组合双敲低实验，并且实验结果显示在图25B中。此时，在fadR和xapR二者均被敲低的菌株中，由甘油产生103.8mg/L柚皮素，与对照中的柚皮素产量相比，柚皮素产量增加61％。
此处，如下构建用于组合双敲低的sRNA质粒。使用引物SEQ ID NO:89和SEQ IDNO:90从亲本质粒扩增有待插入的对合成对照sRNA进行编码的DNA片段，然后通过Gibson组件将所述DNA片段与使用引物SEQ ID NO:91和SEQ ID NO:92通过PCR线性化的含有合成对照sRNA的亲本质粒组装。
[SEQ ID NO:89] 5'-CACTAGATCTCAAATGTGCTGGAATTCTAACACCGTGCGTG-3'
[SEQ ID NO:90] 5'-CCTTATAAATCAAACATGTGCGGCGAATTGGGTACCTATAAAC-3'
[SEQ ID NO:91] 5'-GCACATGTTTGATTTATAAGGG-3'
[SEQ ID NO:92] 5'-CAGCACATTTGAGATCTAGTGG-3'
如上文实施例3中所检验的，通过RppA丙二酰-CoA生物传感器容易且快速地筛选的用于增加丙二酰-CoA产生的敲低靶标都极大地促成有用化合物的基于丙二酰-CoA的产生。在上文实施例中检验的四种产生有用产物的菌株不是通过先前研究中报道的许多代谢工程化策略构建的那些，而是通过构建基本产生途径和简单转化sRNA在短时间内构建的那些。以与如上所述相同的方式构建的菌株显示出卓越产生能力的事实在本领域中是值得注目的。另外，对于本领域技术人员明显的是，RppA生物传感器的实用性和适用性不仅限于上述实施例。
尽管已经参考具体特征详细描述了本发明，但是对于本领域技术人员来说清楚的是，此描述仅用于优选实施方案并且不限制本发明的范围。因此，本发明的实质范围将由所附权利要求及其等效物限定。
实用性
用于测量丙二酰-CoA浓度的常规方法具有各种缺点和局限性，因为它们费时费力，需要熟练的技术，具有低的数据可靠性，难以进行大批量实验，并且需要高性能装置。然而，根据本发明的生物传感器的使用提供了单步信号生成、在各种微生物中使用、在自荧光微生物中使用、简单的构建方法和简单的筛选方法。另外，当本发明与高通量筛选相结合时，其优点在于可以非常容易且快速地(约3天)筛选具有增加的丙二酰-CoA产生能力的菌株，并且所述菌株可以直接应用于有用化合物的基于丙二酰-CoA的产生。
文本文件
参见所附序列表。
序列表
<110> 韩国科学技术院
<120> 基于III型聚酮合酶的新型丙二酰-CoA生物传感器及其用途
<130> PF-B2212
<140> PCT/KR2018/010087
<141> 2018-08-30
<150> KR10-2018-0066323
<151> 2018-06-08
<160> 181
<170> PatentIn版本3.5
<210> 1
<211> 41
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 1
ctttaagaag gagatataca tatggcgacc ctgtgccgac c 41
<210> 2
<211> 43
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 2
cttgtcgacg gagctcgaat tcattagccg gacagcgcaa cgc 43
<210> 3
<211> 40
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 3
cgactcctgc attaggaaat gactgcacgg tgcaccaatg 40
<210> 4
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 4
gcgtttcact tctgagttcg 20
<210> 5
<211> 41
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 5
cgaactcaga agtgaaacgc ctgaaacctc aggcatttga g 41
<210> 6
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 6
atttcctaat gcaggagtcg 20
<210> 7
<211> 41
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 7
caatttcaca caggaaacag aatggcgacc ctgtgccgac c 41
<210> 8
<211> 43
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 8
ctctagagga tccccgggta ccattagccg gacagcgcaa cgc 43
<210> 9
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 9
ggtacccggg gatcctctag ag 22
<210> 10
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 10
tctgtttcct gtgtgaaatt g 21
<210> 11
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 11
gatgctcctt tcttgttatt gaaattgtta tccgctcaca attc 44
<210> 12
<211> 40
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 12
cgaactcaga agtgaaacgc tgcctaatga gtgagctaac 40
<210> 13
<211> 41
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 13
cattggtgca ccgtgcagtc aaaattcgcg ttaaattttt g 41
<210> 14
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 14
gactgcacgg tgcaccaatg 20
<210> 15
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 15
gcgtttcact tctgagttcg 20
<210> 16
<211> 59
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 16
caatttcaca caggaaacag aatgcaccat caccatcacc atgcgaccct gtgccgacc 59
<210> 17
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 17
ccattataat taggcctcgg 20
<210> 18
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 18
ccatgctact cctaccaacc 20
<210> 19
<211> 45
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 19
ggttggtagg agtagcatgg gatccatggc gaccctgtgc cgacc 45
<210> 20
<211> 41
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 20
ccgaggccta attataatgg attagccgga cagcgcaacg c 41
<210> 21
<211> 48
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 21
ggttggtagg agtagcatgg gatccatgca ccatcaccat caccatgc 48
<210> 22
<211> 47
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 22
cttgatcagc ttgcatgcct gcagaagctt tctagcagaa ataattc 47
<210> 23
<211> 27
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 23
gtgcatgtgg actccctttc tcttatc 27
<210> 24
<211> 41
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 24
gataagagaa agggagtcca catggcgacc ctgtgccgac c 41
<210> 25
<211> 45
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 25
ggatcctcta gagtcgacct gcagattagc cggacagcgc aacgc 45
<210> 26
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 26
gataagagaa agggagtcca catgcaccat caccatcacc atgc 44
<210> 27
<211> 43
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 27
gctgagaagc ttgccaaata atggatcctc tagagtcgac ctg 43
<210> 28
<211> 74
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 28
ctggggatgt tttcccagag gggtatgtag aagttgcagc ggaatgcatg aattctgttt 60
cctgtgtgaa attg 74
<210> 29
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 29
acagtgaata tgaattctcc aacg 24
<210> 30
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 30
attatttggc aagcttctca gc 22
<210> 31
<211> 71
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 31
ctctgggaaa acatccccag caccagtcgg aacccctggg actgagtaca gtgaatatga 60
attctccaac g 71
<210> 32
<211> 36
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 32
taataagaat tcatgaagat ttacggaatt tatatg 36
<210> 33
<211> 34
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 33
ttattagaat tcttataaaa gctcttcgta cgag 34
<210> 34
<211> 43
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 34
ctagagtcga cctgcaggca tgccactccc gttctggata atg 43
<210> 35
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 35
caaaacagcc aagcttgcat gc 22
<210> 36
<211> 67
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 36
gaattgtgag cggataacaa agaccgagga aaaggagcat cgcaaatgac gcaggtcgtg 60
gaacgtc 67
<210> 37
<211> 39
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 37
tagaggatcc ccgggtactc atccgaaatc cttcccgtc 39
<210> 38
<211> 19
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 38
gtacccgggg atcctctag 19
<210> 39
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 39
ttgttatccg ctcacaattc 20
<210> 40
<211> 60
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 40
gaggaaaagg agcatcgcaa atgcaccatc atcatcatca tacgcaggtc gtggaacgtc 60
<210> 41
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 41
ttgcgatgct ccttttcctc 20
<210> 42
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 42
atgagcgata aaattattca cctg 24
<210> 43
<211> 19
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 43
cgccaggtta gcgtcgagg 19
<210> 44
<211> 38
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 44
cctcgacgct aacctggcga cgcaggtcgt ggaacgtc 38
<210> 45
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 45
tcatccgaaa tccttcccgt c 21
<210> 46
<211> 49
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 46
agaccgagga aaaggagcat cgcaaatgag cgataaaatt attcacctg 49
<210> 47
<211> 40
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 47
gtaagccagt atacactccg gactgcacgg tgcaccaatg 40
<210> 48
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 48
ctgttgggcg ccatctcctt gtgtagaaac gcaaaaaggc catc 44
<210> 49
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 49
caatttcaca caggaaacag acatatggcg ccacaagaac aagc 44
<210> 50
<211> 46
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 50
ctctagagga tccccgggta ccattacaat ccatttgcta gttttg 46
<210> 51
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 51
cacagcgatg acgtcctact ctgtgttttg 30
<210> 52
<211> 30
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 52
caaaacacag agtaggacgt catcgctgtg 30
<210> 53
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 53
ctctttcgag gaaacaaccc ggtgag 26
<210> 54
<211> 26
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 54
ctcaccgggt tgtttcctcg aaagag 26
<210> 55
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 55
gattgaaaga acttgtcaag tataaagg 28
<210> 56
<211> 28
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 56
cctttatact tgacaagttc tttcaatc 28
<210> 57
<211> 45
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 57
caatttcaca caggaaacag acatatgatc actgcagctc tacac 45
<210> 58
<211> 45
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 58
ctctagagga tccccgggta ccattaacaa agcttagctt tgagg 45
<210> 59
<211> 45
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 59
caatttcaca caggaaacag acatatggtg ctccaacaac aaacg 45
<210> 60
<211> 45
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 60
ctctagagga tccccgggta ccattattta gagcacatgg tttcc 45
<210> 61
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 61
caatttcaca caggaaacag acatatgttc cgcagcgagt acgc 44
<210> 62
<211> 46
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 62
ctctagagga tccccgggta ccattatcgc ggctccctga gctgtc 46
<210> 63
<211> 32
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 63
gtacatcgtc agcggcgccg ccccgctcga cg 32
<210> 64
<211> 32
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 64
cgtcgagcgg ggcggcgccg ctgacgatgt ac 32
<210> 65
<211> 45
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 65
caatttcaca caggaaacag acatatggca agtgtcgagg aattc 45
<210> 66
<211> 46
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 66
ctctagagga tccccgggta ccattaattg gtaaccatcg gaatgg 46
<210> 67
<211> 40
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 67
gtaagccagt atacactccg gactgcacgg tgcaccaatg 40
<210> 68
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 68
ctgttgggcg ccatctcctt gtgtagaaac gcaaaaaggc catc 44
<210> 69
<211> 40
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 69
gagtcgacct gcaggcatgc aatttcacac aggaaacaga 40
<210> 70
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 70
caaaacagcc aagcttgcat g 21
<210> 71
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 71
tttcacacag gaaacagaca tatg 24
<210> 72
<211> 53
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 72
gaattcctcg acacttgcca tactaccacc acccaatcca tttgctagtt ttg 53
<210> 73
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 73
gttgacaatt aatcatcggc 20
<210> 74
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 74
cgtttcactt ctgagttcgg 20
<210> 75
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 75
ccgaactcag aagtgaaacg 20
<210> 76
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 76
gccgatgatt aattgtcaac 20
<210> 77
<211> 41
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 77
gaattttaac aaaatattaa cgaattctaa caccgtgcgt g 41
<210> 78
<211> 42
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 78
gtgccaccta aattgtaagc ggcgaattgg gtacctataa ac 42
<210> 79
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 79
gcttacaatt taggtggcac 20
<210> 80
<211> 24
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 80
gttaatattt tgttaaaatt cgcg 24
<210> 81
<211> 31
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 81
agacagggta ccatggcgcc acaagaacaa g 31
<210> 82
<211> 51
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 82
atgtatatct ccttcttaaa gttaattaca atccatttgc tagttttgcc c 51
<210> 83
<211> 46
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 83
ttaactttaa gaaggagata tacatatgtc ttcatccaac gcctgc 46
<210> 84
<211> 37
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 84
agacagggat cctcagttct ctttggctag tttttcc 37
<210> 85
<211> 31
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 85
agagaggatc cataacaatt ccccatctta g 31
<210> 86
<211> 36
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 86
agagatctag attaggtagc cacactatgc agaacc 36
<210> 87
<211> 43
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 87
gaaactgctt gttcttgtgg cgccatggtc tgtttcctgt gtg 43
<210> 88
<211> 35
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 88
ctaatctaga gtcgacctgc aggcatgcaa gcttg 35
<210> 89
<211> 41
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 89
cactagatct caaatgtgct ggaattctaa caccgtgcgt g 41
<210> 90
<211> 43
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 90
ccttataaat caaacatgtg cggcgaattg ggtacctata aac 43
<210> 91
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 91
gcacatgttt gatttataag gg 22
<210> 92
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 92
cagcacattt gagatctagt gg 22
<210> 93
<211> 372
<212> PRT
<213> 灰色链霉菌(Streptomyces griseus)
<400> 93
Met Ala Thr Leu Cys Arg Pro Ala Ile Ala Val Pro Glu His Val Ile
1 5 10 15
Thr Met Gln Gln Thr Leu Asp Leu Ala Arg Glu Thr His Ala Gly His
20 25 30
Pro Gln Arg Asp Leu Val Leu Arg Leu Ile Gln Asn Thr Gly Val Gln
35 40 45
Thr Arg His Leu Val Gln Pro Ile Glu Lys Thr Leu Ala His Pro Gly
50 55 60
Phe Glu Val Arg Asn Gln Val Tyr Glu Ala Glu Ala Lys Thr Arg Val
65 70 75 80
Pro Glu Val Val Arg Arg Ala Leu Ala Asn Ala Glu Thr Glu Pro Ser
85 90 95
Glu Ile Asp Leu Ile Val Tyr Val Ser Cys Thr Gly Phe Met Met Pro
100 105 110
Ser Leu Thr Ala Trp Ile Ile Asn Ser Met Gly Phe Arg Pro Glu Thr
115 120 125
Arg Gln Leu Pro Ile Ala Gln Leu Gly Cys Ala Ala Gly Gly Ala Ala
130 135 140
Ile Asn Arg Ala His Asp Phe Cys Val Ala Tyr Pro Asp Ser Asn Val
145 150 155 160
Leu Ile Val Ser Cys Glu Phe Cys Ser Leu Cys Tyr Gln Pro Thr Asp
165 170 175
Ile Gly Val Gly Ser Leu Leu Ser Asn Gly Leu Phe Gly Asp Ala Leu
180 185 190
Ser Ala Ala Val Val Arg Gly Gln Gly Gly Thr Gly Met Arg Leu Glu
195 200 205
Arg Asn Gly Ser His Leu Val Pro Asp Thr Glu Asp Trp Ile Ser Tyr
210 215 220
Ala Val Arg Asp Thr Gly Phe His Phe Gln Leu Asp Lys Arg Val Pro
225 230 235 240
Gly Thr Met Glu Met Leu Ala Pro Val Leu Leu Asp Leu Val Asp Leu
245 250 255
His Gly Trp Ser Val Pro Asn Met Asp Phe Phe Ile Val His Ala Gly
260 265 270
Gly Pro Arg Ile Leu Asp Asp Leu Cys His Phe Leu Asp Leu Pro Pro
275 280 285
Glu Met Phe Arg Tyr Ser Arg Ala Thr Leu Thr Glu Arg Gly Asn Ile
290 295 300
Ala Ser Ser Val Val Phe Asp Ala Leu Ala Arg Leu Phe Asp Asp Gly
305 310 315 320
Gly Ala Ala Glu Ser Ala Gln Gly Leu Ile Ala Gly Phe Gly Pro Gly
325 330 335
Ile Thr Ala Glu Val Ala Val Gly Ser Trp Ala Lys Glu Gly Leu Gly
340 345 350
Ala Asp Val Gly Arg Asp Leu Asp Glu Leu Glu Leu Thr Ala Gly Val
355 360 365
Ala Leu Ser Gly
370
<210> 94
<211> 374
<212> PRT
<213> 天蓝色链霉菌(Streptomyces coelicolor)
<400> 94
Met Ala Thr Leu Cys Arg Pro Ser Val Ser Val Pro Glu His Val Ile
1 5 10 15
Thr Met Glu Glu Thr Leu Glu Leu Ala Arg Arg Arg His Thr Asp His
20 25 30
Pro Gln Leu Pro Leu Ala Leu Arg Leu Ile Glu Asn Thr Gly Val Arg
35 40 45
Thr Arg His Ile Val Gln Pro Ile Glu Asp Thr Leu Glu His Pro Gly
50 55 60
Phe Glu Asp Arg Asn Lys Val Tyr Glu Arg Glu Ala Lys Ser Arg Val
65 70 75 80
Pro Ala Val Ile Gln Arg Ala Leu Asp Asp Ala Glu Leu Leu Ala Thr
85 90 95
Asp Ile Asp Val Ile Ile Tyr Val Ser Cys Thr Gly Phe Met Met Pro
100 105 110
Ser Leu Thr Ala Trp Leu Ile Asn Glu Met Gly Phe Asp Ser Thr Thr
115 120 125
Arg Gln Ile Pro Ile Ala Gln Leu Gly Cys Ala Ala Gly Gly Ala Ala
130 135 140
Ile Asn Arg Ala His Asp Phe Cys Thr Ala Tyr Pro Glu Ala Asn Ala
145 150 155 160
Leu Ile Val Ala Cys Glu Phe Cys Ser Leu Cys Tyr Gln Pro Thr Asp
165 170 175
Leu Gly Val Gly Ser Leu Leu Cys Asn Gly Leu Phe Gly Asp Gly Ile
180 185 190
Ala Ala Ala Val Val Arg Gly Arg Gly Gly Thr Gly Val Arg Leu Glu
195 200 205
Arg Asn Gly Ser Tyr Leu Ile Pro Lys Thr Glu Asp Trp Ile Met Tyr
210 215 220
Asp Val Lys Ala Thr Gly Phe His Phe Leu Leu Asp Lys Arg Val Pro
225 230 235 240
Ala Thr Met Glu Pro Leu Ala Pro Ala Leu Lys Glu Leu Ala Gly Glu
245 250 255
His Gly Trp Asp Ala Ser Asp Leu Asp Phe Tyr Ile Val His Ala Gly
260 265 270
Gly Pro Arg Ile Leu Asp Asp Leu Ser Thr Phe Leu Glu Val Asp Pro
275 280 285
His Ala Phe Arg Phe Ser Arg Ala Thr Leu Thr Glu Tyr Gly Asn Ile
290 295 300
Ala Ser Ala Val Val Leu Asp Ala Leu Arg Arg Leu Phe Asp Glu Gly
305 310 315 320
Gly Val Glu Glu Gly Ala Arg Gly Leu Leu Ala Gly Phe Gly Pro Gly
325 330 335
Ile Thr Ala Glu Met Ser Leu Gly Cys Trp Gln Thr Ala Asp Val Arg
340 345 350
Arg Gly Ile Arg Gln Asp Val Thr Arg Thr Ala Ala Arg Gly Val Ser
355 360 365
Arg Arg Val Arg Gln Ala
370
<210> 95
<211> 352
<212> PRT
<213> 阿维链霉菌(Streptomyces avermitilis)
<400> 95
Met Ala Thr Leu Cys Lys Pro Ala Val Ser Val Pro Glu His Val Ile
1 5 10 15
Thr Met Glu Glu Thr Leu Glu Leu Ala Arg Ser Arg His Pro Asp His
20 25 30
Pro Gln Leu Pro Leu Ala Leu Arg Leu Ile Glu Asn Thr Gly Val His
35 40 45
Thr Arg His Ile Val Gln Pro Ile Glu Glu Thr Leu Lys His Pro Gly
50 55 60
Phe Glu Glu Arg Asn His Val Tyr Glu Ala Glu Ala Lys Ala Arg Val
65 70 75 80
Pro Ala Val Val Gln Arg Ala Leu Asp Glu Ala Glu Leu Leu Thr Thr
85 90 95
Asp Ile Asp Val Ile Ile Tyr Val Ser Cys Thr Gly Phe Met Met Pro
100 105 110
Ser Leu Thr Ala Tyr Leu Ile Asn Ser Met Asp Phe Ser Ser Asp Thr
115 120 125
Arg Gln Ile Pro Ile Ala Gln Leu Gly Cys Ala Ala Gly Gly Ser Ala
130 135 140
Ile Asn Arg Ala His Asp Phe Cys Thr Ala Tyr Pro Gln Ala Asn Ala
145 150 155 160
Leu Ile Val Ala Cys Glu Phe Cys Ser Leu Cys Tyr Gln Pro Thr Asp
165 170 175
Leu Gly Val Gly Ser Leu Leu Ser Asn Gly Leu Phe Gly Asp Gly Ile
180 185 190
Ala Ala Ala Ala Val Arg Gly Lys Gly Gly Thr Gly Ile Thr Leu Glu
195 200 205
Arg Asn Ala Ser Tyr Leu Ile Pro Lys Thr Asp Glu Trp Ile Ser Tyr
210 215 220
Asp Val Arg Ala Thr Gly Phe His Phe Leu Leu Asp Lys Arg Val Pro
225 230 235 240
Gly Thr Met Glu Pro Leu Ala Pro Ala Leu Gln Glu Leu Ala Ser Gln
245 250 255
His Gly Trp Asp Ala Ser Asp Leu Asp Phe Tyr Ile Ile His Ala Gly
260 265 270
Gly Pro Arg Ile Leu Asp Asp Leu Ser Lys Phe Leu Arg Val Pro Pro
275 280 285
Glu Ala Phe Arg Phe Ser Arg Ala Thr Leu Thr Glu Tyr Gly Asn Ile
290 295 300
Ala Ser Ala Val Val Leu Asp Ala Leu Arg Arg Leu Phe Asp Glu Gly
305 310 315 320
Gly Ala Glu His Ala Ala Arg Gly Met Leu Ala Gly Phe Gly Pro Gly
325 330 335
Ile Thr Ala Glu Met Ser Leu Gly Arg Trp His Arg Thr Asp Glu Ala
340 345 350
<210> 96
<211> 367
<212> PRT
<213> 红色糖多孢菌(Saccharopolyspora erythraea)
<400> 96
Met Ala Val Leu Cys Thr Pro Ala Val Ala Val Pro Glu His Val Ile
1 5 10 15
Thr Val Glu Glu Thr Leu Asp Leu Ala Arg Arg Val His Ala Asp His
20 25 30
Pro Gln Leu Pro Leu Val Leu Arg Leu Ile Ser Asn Thr Gly Val Arg
35 40 45
Glu Arg His Leu Ile Arg Pro Ile Glu Asp Thr Leu Glu His Pro Gly
50 55 60
Phe Glu Val Arg Asn Arg Ile Tyr Glu Glu Gln Ala Lys Gln Arg Val
65 70 75 80
Pro Ala Val Val Arg Glu Ala Leu Asp Ser Ala Glu Leu Gly Pro Glu
85 90 95
Asp Ile Asp Leu Ile Val Tyr Val Ser Cys Thr Gly Phe Met Met Pro
100 105 110
Ser Leu Thr Ala Trp Leu Ile Asn Ser Met Gly Phe Arg Met Ser Thr
115 120 125
Arg Gln Leu Pro Ile Ala Gln Leu Gly Cys Ala Ala Gly Gly Ala Ala
130 135 140
Ile Asn Arg Ala His Asp Phe Cys Thr Ala Tyr Pro Asp Ala Asn Ala
145 150 155 160
Leu Ile Val Ser Cys Glu Phe Cys Ser Leu Cys Tyr Gln Pro Thr Asp
165 170 175
Asp Asp Ile Gly Ser Leu Leu Ser Asn Gly Leu Phe Gly Asp Ala Val
180 185 190
Gly Ala Ala Val Val Arg Gly His Gly Gly Thr Gly Val Arg Leu Glu
195 200 205
Arg Asn Ala Ser Ser Met Ile Pro Glu Thr Glu Asp Trp Ile Ser Tyr
210 215 220
Ala Val Lys Ala Thr Gly Phe His Phe Gln Leu Asp Lys Arg Val Pro
225 230 235 240
Lys Thr Met Glu Pro Leu Ala Pro Ala Leu Arg Ala Leu Ala Glu Asp
245 250 255
His Arg Trp Asp Val Ala Gly Leu Asp Phe Tyr Val Ile His Ala Gly
260 265 270
Gly Pro Arg Ile Leu Asp Asp Leu Thr Lys Phe Leu Gly Val Pro Ser
275 280 285
Glu Ala Phe Arg His Ser Arg Ala Thr Leu Ala Gln Tyr Gly Asn Ile
290 295 300
Ala Ser Ala Val Val Leu Asp Ala Leu Arg Arg Ile Ile Glu Glu Gly
305 310 315 320
Arg Leu Glu Ser Gly Ala Arg Gly Met Ile Ala Gly Phe Gly Pro Gly
325 330 335
Ile Thr Ala Glu Met Ser Val Gly Thr Trp Val Pro His Asp Val Leu
340 345 350
Leu His Gly Glu His Ser Thr Thr Ser Ala Pro Gly Gly Asn Arg
355 360 365
<210> 97
<211> 377
<212> PRT
<213> 波赛链霉菌(Streptomyces peucetius)
<400> 97
Met Arg Val Pro Val Ala Val Asp Asp Leu Val Ala Pro Ser Thr Met
1 5 10 15
Gly Glu Arg His Thr Val Ile Asp Arg Gly Thr Ser Val Ala Ala Val
20 25 30
His Thr Ala Leu Pro Pro His Arg Tyr Ala Gln Ser Asp Leu Thr Glu
35 40 45
Leu Ile Ala Asp Leu Cys Leu Glu Pro Gly Ala Asp Arg Ala Leu Leu
50 55 60
Arg Arg Leu His Thr Ser Ala Gly Val Arg Thr Arg His Leu Ala Leu
65 70 75 80
Pro Ile Glu Gln Tyr Ala Gly Leu Gly Asp Phe Gly Gln Ala Asn Ala
85 90 95
Ala Trp Leu Thr Val Gly Leu Ala Leu Ala Glu Glu Ala Leu Ser Gly
100 105 110
Ala Leu Asp Ala Ala Gly Leu Thr Ala Ala Asp Ile Asp Leu Leu Val
115 120 125
Cys Thr Ser Ile Thr Gly Val Ala Ala Pro Ser Leu Asp Ala Arg Leu
130 135 140
Ala Val Arg Met Gly Met Arg Ala Asp Val Lys Arg Val Pro Val Phe
145 150 155 160
Gly Leu Gly Cys Val Gly Gly Ala Ala Gly Leu Gly Arg Leu His Asp
165 170 175
Tyr Leu Leu Gly His Pro Asp Asp Thr Ala Val Leu Leu Ser Val Glu
180 185 190
Leu Cys Ser Leu Thr Leu Gln Arg Asp Gly Ser Leu Ala Asn Leu Val
195 200 205
Ala Gly Ala Leu Phe Gly Asp Gly Ala Ala Ala Val Val Ala Arg Gly
210 215 220
Gly Asp Ala Gly Arg Arg Gly Ala Gly Trp Pro Met Val Ala Ala Thr
225 230 235 240
Arg Gly His Leu Tyr Pro Asp Thr Glu His Leu Leu Gly Trp Arg Ile
245 250 255
Gly Ala Ser Gly Phe Arg Val Val Val Asp Ala Gly Ile Pro Asp Val
260 265 270
Val Arg Thr His Leu Gly Gly Asp Leu Arg Asn Phe Leu Ala Thr His
275 280 285
Gly Leu Val Pro Asp Asp Ile Gly Thr Trp Ile Cys His Pro Gly Gly
290 295 300
Pro Lys Val Leu Ala Ala Val Gly Asp Ala Leu Glu Leu Pro Asp Gly
305 310 315 320
Ala Leu Asp Ser Ser Trp Arg Ser Leu Ala Gly Val Gly Asn Leu Ser
325 330 335
Ser Ala Ser Val Leu Arg Val Leu Glu Asp Val Ala Thr Arg Cys Arg
340 345 350
Pro Asp Pro Gly Thr Trp Gly Val Leu Leu Ala Met Gly Pro Gly Phe
355 360 365
Cys Ala Glu Phe Val Leu Leu Arg Trp
370 375
<210> 98
<211> 355
<212> PRT
<213> 针孢链霉菌(Streptomyces aculeolatus)
<400> 98
Met Pro Arg Leu Cys Lys Pro Ala Val Ser Ala Pro Glu Tyr Thr Ile
1 5 10 15
Thr Met Glu Glu Thr Leu Glu Phe Ala Lys Gln Ala His Ala Gly Lys
20 25 30
Pro Gln Leu Pro Leu Ala Leu Arg Leu Ile Arg Asn Thr Gly Val Leu
35 40 45
Lys Arg His Ile Val Gln Pro Ile Glu Lys Thr Leu Gly His Pro Gly
50 55 60
Leu Thr Glu Arg Asn Leu Ile Tyr Glu Ala Glu Ser Lys Lys Met Cys
65 70 75 80
Pro Pro Val Ile Glu Glu Ala Leu Gln Asn Ala Asp Met Thr Ala Arg
85 90 95
Asp Ile Asp Ala Ile Ile Tyr Val Ser Cys Thr Gly Phe Leu Met Pro
100 105 110
Ser Leu Thr Ala Trp Leu Ile Asn Lys Met Gly Phe Arg Ser Asp Thr
115 120 125
Arg Gln Ile Pro Ile Ala Gln Leu Gly Cys Ala Ala Gly Gly Ala Ala
130 135 140
Val Asn Arg Ala His Asp Phe Cys Leu Ala His Pro Gly Ser Asn Val
145 150 155 160
Leu Ile Val Ala Cys Glu Leu Cys Ser Leu Cys Tyr Gln Pro Thr Ala
165 170 175
Asp Asp Ile Gly Ser Leu Leu Ser Asp Gly Leu Phe Gly Asp Ala Val
180 185 190
Ala Ala Ala Val Val Arg Gly Asn Gly Gly Val Gly Ile Glu Val Glu
195 200 205
Arg Asn Ala Ser Tyr Leu Ile Pro Asn Thr Glu Glu Trp Ile Ser Tyr
210 215 220
Ser Val Arg Asp Thr Gly Phe His Phe Gln Leu Asp Arg Arg Val Pro
225 230 235 240
Gly Thr Met Glu Pro Leu Ala Pro Val Leu Arg Glu Phe Ala Lys Asp
245 250 255
His Ser Trp Asp Ala Gly Lys Leu Asp Phe Tyr Ile Val His Ala Gly
260 265 270
Gly Pro Arg Ile Leu Asp Asp Leu Ala Arg Phe Leu Asp Val Asp Arg
275 280 285
Gln Val Phe Arg His Ser Trp Ser Thr Leu Thr Glu Tyr Gly Asn Ile
290 295 300
Ala Ser Ala Val Val Phe Asp Ala Ala Arg Arg Leu Phe Glu Glu Gly
305 310 315 320
Ser Ala Lys Pro Asp Ala Thr Gly Met Ile Ala Gly Phe Gly Pro Gly
325 330 335
Ile Thr Ala Glu Met Ala Leu Gly Thr Trp Gly Thr Asp Gly Thr Gly
340 345 350
Thr Ser Asn
355
<210> 99
<211> 349
<212> PRT
<213> 荧光假单胞菌(Pseudomonas fluorescens)
<400> 99
Met Ser Thr Leu Cys Lys Pro Ser Leu Leu Phe Pro His Tyr Lys Ile
1 5 10 15
Thr Gln Gln Gln Met Ile Asp His Leu Glu Gln Leu His Asp Asp His
20 25 30
Pro Arg Met Ala Leu Ala Lys Arg Met Ile Gln Asn Thr Gln Val Asn
35 40 45
Glu Arg Tyr Leu Val Leu Pro Ile Asp Glu Leu Ala Val His Thr Gly
50 55 60
Phe Thr His Arg Ser Ile Val Tyr Glu Arg Glu Ala Arg Arg Met Ser
65 70 75 80
Ser Ile Ala Ala Arg Gln Ala Ile Glu Asn Ala Gly Leu Thr Thr Asp
85 90 95
Asp Ile Arg Met Val Ala Val Thr Ser Cys Thr Gly Phe Met Met Pro
100 105 110
Ser Leu Thr Ala His Leu Ile Asn Asp Leu Gly Leu Arg Thr Ser Thr
115 120 125
Val Gln Leu Pro Ile Ala Gln Leu Gly Cys Val Ala Gly Ala Ala Ala
130 135 140
Ile Asn Arg Ala Asn Asp Phe Ala Ser Leu Ser Pro Asp Asn His Ala
145 150 155 160
Leu Ile Val Ser Leu Glu Phe Ser Ser Leu Cys Tyr Gln Pro Gln Asp
165 170 175
Thr Lys Leu His Ala Phe Ile Ser Ala Ala Leu Phe Gly Asp Ala Val
180 185 190
Ser Ala Cys Val Met Arg Ala Asp Asp Lys Ala Pro Gly Phe Lys Ile
195 200 205
Ala Lys Thr Gly Ser Tyr Phe Leu Pro Asp Ser Glu His Tyr Ile Lys
210 215 220
Tyr Asp Val Lys Asp Ser Gly Phe His Phe Thr Leu Asp Lys Ala Val
225 230 235 240
Met Asn Ser Ile Lys Asp Val Ala Pro Met Met Glu Glu Leu Asn Phe
245 250 255
Glu Thr Phe Asn Gln His Cys Ala Gln Asn Asp Phe Phe Ile Phe His
260 265 270
Thr Gly Gly Arg Lys Ile Leu Asp Glu Leu Val Leu Gln Leu Asp Leu
275 280 285
Glu Pro Gly Arg Val Ala Gln Ser Arg Asp Ser Leu Ser Glu Ala Gly
290 295 300
Asn Ile Ala Ser Val Val Val Phe Asp Val Leu Lys Arg Gln Phe Asp
305 310 315 320
Ser Gly Pro Ala Asn Gly Ala Thr Gly Met Leu Ala Ala Phe Gly Pro
325 330 335
Gly Phe Thr Ala Glu Met Ala Val Gly Lys Trp Val Ala
340 345
<210> 100
<211> 370
<212> PRT
<213> 东方拟无枝菌酸菌(Amycolatopsis orientalis)
<400> 100
Met Asp Val Ser Met Thr Thr Gly Ile Glu Leu Thr Glu Glu Leu Ser
1 5 10 15
Val Leu Asn Gly Leu Thr Glu Ile Thr Arg Phe Ala Gly Val Gly Thr
20 25 30
Ala Val Ser Glu Thr Ser Tyr Ser Gln Thr Glu Leu Leu Asp Ile Leu
35 40 45
Asp Val Glu Asp Pro Lys Ile Arg Ser Val Phe Leu Asn Ser Ala Ile
50 55 60
Asp Arg Arg Phe Leu Thr Leu Pro Pro Glu Asn Pro Gly Gly Gly Arg
65 70 75 80
Leu Ala Glu Pro Gln Gly Asp Leu Leu Asp Lys His Lys Lys Ile Ala
85 90 95
Val Asp Met Gly Cys Arg Ala Leu Glu Ala Cys Leu Lys Ser Ala Gly
100 105 110
Ala Thr Leu Ser Asp Leu Arg His Leu Cys Cys Val Thr Ser Thr Gly
115 120 125
Phe Leu Thr Pro Gly Leu Ser Ala Leu Ile Ile Arg Glu Met Gly Ile
130 135 140
Asp Pro His Cys Ser Arg Ser Asp Ile Val Gly Met Gly Cys Asn Ala
145 150 155 160
Gly Leu Asn Ala Leu Asn Val Val Ser Gly Trp Ser Ala Ala His Pro
165 170 175
Gly Glu Leu Gly Val Val Leu Cys Ser Glu Ala Cys Ser Ala Ala Tyr
180 185 190
Ala Leu Asp Gly Thr Met Arg Thr Ala Val Val Asn Ser Leu Phe Gly
195 200 205
Asp Gly Ser Ala Ala Leu Ala Val Ile Ser Gly Asp Gly Arg Val Ala
210 215 220
Gly Pro Arg Val Leu Lys Phe Ala Ser Tyr Ile Ile Thr Asp Ala Val
225 230 235 240
Asp Ala Met Arg Tyr Asp Trp Asp Arg Asp Gln Asp Arg Phe Ser Phe
245 250 255
Phe Leu Asp Pro Gln Ile Pro Tyr Val Val Gly Ala His Ala Glu Ile
260 265 270
Val Val Asp Arg Leu Leu Ser Gly Thr Gly Leu Arg Arg Ser Asp Ile
275 280 285
Gly His Trp Leu Val His Ser Gly Gly Lys Lys Val Val Asp Ala Val
290 295 300
Val Val Asn Leu Gly Leu Ser Arg His Asp Val Arg His Thr Thr Gly
305 310 315 320
Val Leu Arg Asp Tyr Gly Asn Leu Ser Ser Gly Ser Phe Leu Phe Ser
325 330 335
Tyr Glu Arg Leu Ser Glu Glu Asp Val Thr Arg Pro Gly Asp Tyr Gly
340 345 350
Val Leu Met Thr Met Gly Pro Gly Ser Thr Ile Glu Met Ala Leu Ile
355 360 365
Gln Trp
370
<210> 101
<211> 391
<212> PRT
<213> 未知
<220>
<223> 掌叶大黄(Rheum palmatum)
<400> 101
Met Ala Asp Val Leu Gln Glu Ile Arg Asn Ser Gln Lys Ala Ser Gly
1 5 10 15
Pro Ala Thr Val Leu Ala Ile Gly Thr Ala His Pro Pro Thr Cys Tyr
20 25 30
Pro Gln Ala Asp Tyr Pro Asp Phe Tyr Phe Arg Val Cys Lys Ser Glu
35 40 45
His Met Thr Lys Leu Lys Lys Lys Met Gln Phe Ile Cys Asp Arg Ser
50 55 60
Gly Ile Arg Gln Arg Phe Met Phe His Thr Glu Glu Asn Leu Gly Lys
65 70 75 80
Asn Pro Gly Met Cys Thr Phe Asp Gly Pro Ser Leu Asn Ala Arg Gln
85 90 95
Asp Met Leu Ile Met Glu Val Pro Lys Leu Gly Ala Glu Ala Ala Glu
100 105 110
Lys Ala Ile Lys Glu Trp Gly Gln Asp Lys Ser Arg Ile Thr His Leu
115 120 125
Ile Phe Cys Thr Thr Thr Ser Asn Asp Met Pro Gly Ala Asp Tyr Gln
130 135 140
Phe Ala Thr Leu Phe Gly Leu Asn Pro Gly Val Ser Arg Thr Met Val
145 150 155 160
Tyr Gln Gln Gly Cys Phe Ala Gly Gly Thr Val Leu Arg Leu Val Lys
165 170 175
Asp Ile Ala Glu Asn Asn Lys Gly Ala Arg Val Leu Val Val Cys Ser
180 185 190
Glu Ile Val Ala Phe Ala Phe Arg Gly Pro His Glu Asp His Ile Asp
195 200 205
Ser Leu Ile Gly Gln Leu Leu Phe Gly Asp Gly Ala Ala Ala Leu Val
210 215 220
Val Gly Thr Asp Ile Asp Glu Ser Val Glu Arg Pro Ile Phe Gln Ile
225 230 235 240
Met Ser Ala Thr Gln Ala Thr Ile Pro Asn Ser Leu His Thr Met Ala
245 250 255
Leu His Leu Thr Glu Ala Gly Leu Thr Phe His Leu Ser Lys Glu Val
260 265 270
Pro Lys Val Val Ser Asp Asn Met Glu Glu Leu Met Leu Glu Ala Phe
275 280 285
Lys Pro Leu Gly Ile Thr Asp Trp Asn Ser Ile Phe Trp Gln Val His
290 295 300
Pro Gly Gly Arg Ala Ile Leu Asp Lys Ile Glu Glu Lys Leu Glu Leu
305 310 315 320
Thr Lys Asp Lys Met Arg Asp Ser Arg Tyr Ile Leu Ser Glu Tyr Gly
325 330 335
Asn Leu Thr Ser Ala Cys Val Leu Phe Val Met Asp Glu Met Arg Lys
340 345 350
Arg Ser Phe Arg Glu Gly Lys Gln Thr Thr Gly Asp Gly Tyr Glu Trp
355 360 365
Gly Val Ala Ile Gly Leu Gly Pro Gly Leu Thr Val Glu Thr Val Val
370 375 380
Leu Arg Ser Val Pro Ile Pro
385 390
<210> 102
<211> 403
<212> PRT
<213> 木立芦荟(Aloe arborescens)
<400> 102
Met Ser Ser Leu Ser Asn Ser Leu Pro Leu Met Glu Asp Val Gln Gly
1 5 10 15
Ile Arg Lys Ala Gln Lys Ala Asp Gly Thr Ala Thr Val Met Ala Ile
20 25 30
Gly Thr Ala His Pro Pro His Ile Phe Pro Gln Asp Thr Tyr Ala Asp
35 40 45
Val Tyr Phe Arg Ala Thr Asn Ser Glu His Lys Val Glu Leu Lys Lys
50 55 60
Lys Phe Asp His Ile Cys Lys Lys Thr Met Ile Gly Lys Arg Tyr Phe
65 70 75 80
Asn Tyr Asp Glu Glu Phe Leu Lys Lys Tyr Pro Asn Ile Thr Ser Tyr
85 90 95
Asp Glu Pro Ser Leu Asn Asp Arg Gln Asp Ile Cys Val Pro Gly Val
100 105 110
Pro Ala Leu Gly Thr Glu Ala Ala Val Lys Ala Ile Glu Glu Trp Gly
115 120 125
Arg Pro Lys Ser Glu Ile Thr His Leu Val Phe Cys Thr Ser Cys Gly
130 135 140
Val Asp Met Pro Ser Ala Asp Phe Gln Cys Ala Lys Leu Leu Gly Leu
145 150 155 160
His Ala Asn Val Asn Lys Tyr Cys Ile Tyr Met Gln Gly Cys Tyr Ala
165 170 175
Gly Gly Thr Val Met Arg Tyr Ala Lys Asp Leu Ala Glu Asn Asn Arg
180 185 190
Gly Ala Arg Val Leu Val Val Cys Ala Glu Leu Thr Ile Met Met Leu
195 200 205
Arg Ala Pro Asn Glu Thr His Leu Asp Asn Ala Ile Gly Ile Ser Leu
210 215 220
Phe Gly Asp Gly Ala Ala Ala Leu Ile Ile Gly Ser Asp Pro Ile Ile
225 230 235 240
Gly Val Glu Lys Pro Met Phe Glu Ile Val Cys Thr Lys Gln Thr Val
245 250 255
Ile Pro Asn Thr Glu Asp Val Ile His Leu His Leu Arg Glu Thr Gly
260 265 270
Met Met Phe Tyr Leu Ser Lys Gly Ser Pro Met Thr Ile Ser Asn Asn
275 280 285
Val Glu Ala Cys Leu Ile Asp Val Phe Lys Ser Val Gly Ile Thr Pro
290 295 300
Pro Glu Asp Trp Asn Ser Leu Phe Trp Ile Pro His Pro Gly Gly Arg
305 310 315 320
Ala Ile Leu Asp Gln Val Glu Ala Lys Leu Lys Leu Arg Pro Glu Lys
325 330 335
Phe Arg Ala Ala Arg Thr Val Leu Trp Asp Tyr Gly Asn Met Val Ser
340 345 350
Ala Ser Val Gly Tyr Ile Leu Asp Glu Met Arg Arg Lys Ser Ala Ala
355 360 365
Lys Gly Leu Glu Thr Tyr Gly Glu Gly Leu Glu Trp Gly Val Leu Leu
370 375 380
Gly Phe Gly Pro Gly Ile Thr Val Glu Thr Ile Leu Leu His Ser Leu
385 390 395 400
Pro Leu Met
<210> 103
<211> 403
<212> PRT
<213> 木立芦荟
<400> 103
Met Ser Ser Leu Ser Asn Ala Ser His Leu Met Glu Asp Val Gln Gly
1 5 10 15
Ile Arg Lys Ala Gln Arg Ala Asp Gly Thr Ala Thr Val Met Ala Ile
20 25 30
Gly Thr Ala His Pro Pro His Ile Phe Pro Gln Asp Thr Tyr Ala Asp
35 40 45
Phe Tyr Phe Arg Ala Thr Asn Ser Glu His Lys Val Glu Leu Lys Lys
50 55 60
Lys Phe Asp Arg Ile Cys Lys Lys Thr Met Ile Gly Lys Arg Tyr Phe
65 70 75 80
Asn Tyr Asp Glu Glu Phe Leu Lys Lys Tyr Pro Asn Ile Thr Ser Phe
85 90 95
Asp Glu Pro Ser Leu Asn Asp Arg Gln Asp Ile Cys Val Pro Gly Val
100 105 110
Pro Ala Leu Gly Ala Glu Ala Ala Val Lys Ala Ile Ala Glu Trp Gly
115 120 125
Arg Pro Lys Ser Glu Ile Thr His Leu Val Phe Cys Thr Ser Cys Gly
130 135 140
Val Asp Met Pro Ser Ala Asp Phe Gln Cys Ala Lys Leu Leu Gly Leu
145 150 155 160
Arg Thr Asn Val Asn Lys Tyr Cys Val Tyr Met Gln Gly Cys Tyr Ala
165 170 175
Gly Gly Thr Val Met Arg Tyr Ala Lys Asp Leu Ala Glu Asn Asn Arg
180 185 190
Gly Ala Arg Val Leu Val Val Cys Ala Glu Leu Thr Ile Ile Gly Leu
195 200 205
Arg Gly Pro Asn Glu Ser His Leu Asp Asn Ala Ile Gly Asn Ser Leu
210 215 220
Phe Gly Asp Gly Ala Ala Ala Leu Ile Val Gly Ser Asp Pro Ile Ile
225 230 235 240
Gly Val Glu Lys Pro Met Phe Glu Ile Val Cys Ala Lys Gln Thr Val
245 250 255
Ile Pro Asn Ser Glu Asp Val Ile His Leu His Met Arg Glu Ala Gly
260 265 270
Leu Met Phe Tyr Met Ser Lys Asp Ser Pro Glu Thr Ile Ser Asn Asn
275 280 285
Val Glu Ala Cys Leu Val Asp Val Phe Lys Ser Val Gly Met Thr Pro
290 295 300
Pro Glu Asp Trp Asn Ser Leu Phe Trp Ile Pro His Pro Gly Gly Arg
305 310 315 320
Ala Ile Leu Asp Gln Val Glu Ala Lys Leu Lys Leu Arg Pro Glu Lys
325 330 335
Phe Arg Ala Thr Arg Thr Val Leu Trp Asp Cys Gly Asn Met Val Ser
340 345 350
Ala Cys Val Leu Tyr Ile Leu Asp Glu Met Arg Arg Lys Ser Ala Asp
355 360 365
Glu Gly Leu Glu Thr Tyr Gly Glu Gly Leu Glu Trp Gly Val Leu Leu
370 375 380
Gly Phe Gly Pro Gly Met Thr Val Glu Thr Ile Leu Leu His Ser Leu
385 390 395 400
Pro Leu Met
<210> 104
<211> 403
<212> PRT
<213> 木立芦荟
<400> 104
Met Gly Ser Leu Ser Asp Ser Thr Pro Leu Met Lys Asp Val Gln Gly
1 5 10 15
Ile Arg Lys Ala Gln Lys Ala Asp Gly Thr Ala Thr Val Met Ala Ile
20 25 30
Gly Thr Ala His Pro Pro His Ile Ile Ser Gln Asp Ser Tyr Ala Asp
35 40 45
Phe Tyr Phe Arg Val Thr Asn Ser Glu His Lys Val Glu Leu Lys Lys
50 55 60
Lys Phe Asp Arg Ile Cys Lys Lys Thr Met Ile Gly Lys Arg Tyr Phe
65 70 75 80
Asn Phe Asp Glu Glu Phe Leu Lys Lys Tyr Pro Asn Ile Thr Ser Phe
85 90 95
Asp Lys Pro Ser Leu Asn Asp Arg His Asp Ile Cys Ile Pro Gly Val
100 105 110
Pro Ala Leu Gly Ala Glu Ala Ala Val Lys Ala Ile Glu Glu Trp Gly
115 120 125
Arg Pro Lys Ser Glu Ile Thr His Leu Val Phe Cys Thr Ser Gly Gly
130 135 140
Val Asp Met Pro Ser Ala Asp Phe Gln Cys Ala Lys Leu Leu Gly Leu
145 150 155 160
Arg Thr Asn Val Asn Lys Tyr Cys Ile Tyr Met Gln Gly Cys Tyr Ala
165 170 175
Gly Gly Thr Val Met Arg Tyr Ala Lys Asp Leu Ala Glu Asn Asn Arg
180 185 190
Gly Ala Arg Val Leu Met Val Cys Ala Glu Leu Thr Ile Ile Ala Leu
195 200 205
Arg Gly Pro Asn Asp Ser His Ile Asp Asn Ala Ile Gly Asn Ser Leu
210 215 220
Phe Gly Asp Gly Ala Ala Ala Leu Ile Val Gly Ser Asp Pro Ile Ile
225 230 235 240
Gly Val Glu Lys Pro Met Phe Glu Ile Val Cys Ala Lys Gln Thr Val
245 250 255
Ile Pro Asn Ser Glu Glu Val Ile His Leu His Leu Arg Glu Ser Gly
260 265 270
Leu Met Phe Tyr Met Thr Lys Asp Ser Ala Ala Thr Ile Ser Asn Asn
275 280 285
Ile Glu Ala Cys Leu Val Asp Val Phe Lys Ser Val Gly Met Thr Pro
290 295 300
Pro Glu Asp Trp Asn Ser Leu Phe Trp Ile Pro His Pro Gly Gly Arg
305 310 315 320
Ala Ile Leu Asp Gln Val Glu Ala Lys Leu Lys Leu Arg Pro Glu Lys
325 330 335
Phe Ser Ala Thr Arg Thr Val Leu Trp Asp Tyr Gly Asn Met Ile Ser
340 345 350
Ala Cys Val Leu Tyr Ile Leu Asp Glu Met Arg Arg Lys Ser Ala Ala
355 360 365
Glu Gly Leu Glu Thr Tyr Gly Glu Gly Leu Glu Trp Gly Val Leu Leu
370 375 380
Gly Phe Gly Pro Gly Met Thr Ile Glu Thr Ile Leu Leu His Ser Leu
385 390 395 400
Pro Pro Val
<210> 105
<211> 403
<212> PRT
<213> 木立芦荟
<400> 105
Met Gly Ser Leu Ser Asn Tyr Ser Pro Val Met Glu Asp Val Gln Ala
1 5 10 15
Ile Arg Lys Ala Gln Lys Ala Asp Gly Thr Ala Thr Val Met Ala Ile
20 25 30
Gly Thr Ala His Pro Pro His Ile Phe Pro Gln Asp Thr Tyr Ala Asp
35 40 45
Phe Tyr Phe Arg Ala Thr Asn Ser Glu His Lys Val Glu Leu Lys Lys
50 55 60
Lys Phe Asp Arg Ile Cys Lys Lys Thr Met Ile Gly Lys Arg Tyr Phe
65 70 75 80
Asn Tyr Asp Glu Glu Phe Leu Lys Lys Tyr Pro Asn Ile Thr Ser Phe
85 90 95
Asp Glu Pro Ser Leu Asn Asp Arg Gln Asp Ile Cys Val Pro Gly Val
100 105 110
Pro Ala Leu Gly Ala Glu Ala Ala Val Lys Ala Ile Ala Glu Trp Gly
115 120 125
Arg Pro Lys Ser Glu Ile Thr His Leu Val Phe Cys Thr Ser Cys Gly
130 135 140
Val Asp Met Pro Ser Ala Asp Phe Gln Cys Ala Lys Leu Leu Gly Leu
145 150 155 160
Arg Thr Asn Val Asn Lys Tyr Cys Val Tyr Met Gln Gly Cys Tyr Ala
165 170 175
Gly Gly Thr Val Met Arg Tyr Ala Lys Asp Leu Ala Glu Asn Asn Arg
180 185 190
Gly Ala Arg Val Leu Val Val Cys Ala Glu Leu Thr Ile Ile Gly Leu
195 200 205
Arg Gly Pro Asn Glu Ser His Leu Asp Asn Ala Ile Gly Asn Ser Leu
210 215 220
Phe Gly Asp Gly Ala Ala Ala Leu Ile Val Gly Ser Asp Pro Ile Ile
225 230 235 240
Gly Val Glu Arg Pro Met Phe Glu Ile Val Cys Ala Lys Gln Thr Val
245 250 255
Ile Pro Asn Ser Glu Asp Val Ile His Leu His Met Arg Glu Ala Gly
260 265 270
Leu Met Phe Tyr Met Ser Lys Asp Ser Pro Glu Thr Ile Ser Asn Asn
275 280 285
Val Glu Ala Cys Leu Val Asp Val Phe Lys Ser Val Gly Met Thr Pro
290 295 300
Pro Glu Asp Trp Asn Ser Leu Phe Trp Ile Pro His Pro Gly Gly Arg
305 310 315 320
Ala Ile Leu Asp Gln Val Glu Ala Arg Leu Lys Leu Arg Pro Glu Lys
325 330 335
Phe Gly Ala Thr Arg Thr Val Leu Trp Asp Cys Gly Asn Met Val Ser
340 345 350
Ala Cys Val Leu Tyr Ile Leu Asp Glu Met Arg Arg Lys Ser Val Ala
355 360 365
Asp Gly Leu Ala Thr Tyr Gly Glu Gly Leu Glu Trp Gly Val Leu Leu
370 375 380
Gly Phe Gly Pro Gly Met Thr Val Glu Thr Ile Leu Leu His Ser Leu
385 390 395 400
Pro Pro Val
<210> 106
<211> 405
<212> PRT
<213> 木立芦荟
<400> 106
Met Gly Ser Ile Ala Glu Ser Ser Pro Leu Met Ser Arg Glu Asn Val
1 5 10 15
Glu Gly Ile Arg Lys Ala Gln Arg Ala Glu Gly Thr Ala Thr Val Met
20 25 30
Ala Ile Gly Thr Ala His Pro Pro His Ile Phe Pro Gln Asp Thr Tyr
35 40 45
Ala Asp Phe Tyr Phe Arg Ala Thr Asn Ser Glu His Lys Val Glu Leu
50 55 60
Lys Lys Lys Phe Asp Arg Ile Cys Lys Lys Thr Met Ile Gly Lys Arg
65 70 75 80
Tyr Phe Asn Tyr Asp Glu Glu Phe Leu Lys Lys Tyr Pro Asn Ile Thr
85 90 95
Ser Phe Asp Glu Pro Ser Leu Asn Asp Arg Gln Asp Ile Cys Val Pro
100 105 110
Gly Val Pro Ala Leu Gly Lys Glu Ala Ala Leu Lys Ala Ile Glu Glu
115 120 125
Trp Gly Gln Pro Leu Ser Lys Ile Thr His Leu Val Phe Cys Thr Ser
130 135 140
Cys Gly Val Asp Met Pro Ser Ala Asp Phe Gln Leu Ala Lys Leu Leu
145 150 155 160
Gly Leu Asn Thr Asn Val Asn Lys Tyr Cys Val Tyr Met Gln Gly Cys
165 170 175
Tyr Ala Gly Gly Thr Val Leu Arg Tyr Ala Lys Asp Leu Ala Glu Asn
180 185 190
Asn Arg Gly Ser Arg Val Leu Val Val Cys Ala Glu Leu Thr Ile Ile
195 200 205
Gly Leu Arg Gly Pro Asn Glu Ser His Leu Asp Asn Ala Ile Gly Asn
210 215 220
Ser Leu Phe Gly Asp Gly Ala Ala Ala Leu Ile Val Gly Ala Asp Pro
225 230 235 240
Ile Val Gly Ile Glu Lys Pro Ile Phe Glu Ile Val Cys Ala Lys Gln
245 250 255
Thr Val Ile Pro Asp Ser Glu Asp Val Ile His Leu His Leu Arg Glu
260 265 270
Ala Gly Leu Met Phe Tyr Met Ser Lys Asp Ser Pro Glu Thr Ile Ser
275 280 285
Asn Asn Val Glu Gly Cys Leu Val Asp Ile Phe Lys Ser Val Gly Met
290 295 300
Thr Pro Pro Ala Asp Trp Asn Ser Leu Phe Trp Ile Pro His Pro Gly
305 310 315 320
Gly Arg Ala Ile Leu Asp Glu Val Glu Ala Arg Leu Lys Leu Arg Pro
325 330 335
Glu Lys Phe Arg Ala Thr Arg His Val Leu Trp Glu Tyr Gly Asn Met
340 345 350
Val Ser Ala Cys Val Leu Tyr Ile Leu Asp Glu Met Arg Asn Lys Ser
355 360 365
Ala Ala Asp Gly Leu Gly Thr Tyr Gly Glu Gly Leu Glu Trp Gly Val
370 375 380
Leu Leu Gly Phe Gly Pro Gly Met Thr Val Glu Thr Ile Leu Leu His
385 390 395 400
Ser Leu Pro Pro Val
405
<210> 107
<211> 413
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 107
Met Ser Lys Arg Arg Val Val Val Thr Gly Leu Gly Met Leu Ser Pro
1 5 10 15
Val Gly Asn Thr Val Glu Ser Thr Trp Lys Ala Leu Leu Ala Gly Gln
20 25 30
Ser Gly Ile Ser Leu Ile Asp His Phe Asp Thr Ser Ala Tyr Ala Thr
35 40 45
Lys Phe Ala Gly Leu Val Lys Asp Phe Asn Cys Glu Asp Ile Ile Ser
50 55 60
Arg Lys Glu Gln Arg Lys Met Asp Ala Phe Ile Gln Tyr Gly Ile Val
65 70 75 80
Ala Gly Val Gln Ala Met Gln Asp Ser Gly Leu Glu Ile Thr Glu Glu
85 90 95
Asn Ala Thr Arg Ile Gly Ala Ala Ile Gly Ser Gly Ile Gly Gly Leu
100 105 110
Gly Leu Ile Glu Glu Asn His Thr Ser Leu Met Asn Gly Gly Pro Arg
115 120 125
Lys Ile Ser Pro Phe Phe Val Pro Ser Thr Ile Val Asn Met Val Ala
130 135 140
Gly His Leu Thr Ile Met Tyr Gly Leu Arg Gly Pro Ser Ile Ser Ile
145 150 155 160
Ala Thr Ala Cys Thr Ser Gly Val His Asn Ile Gly His Ala Ala Arg
165 170 175
Ile Ile Ala Tyr Gly Asp Ala Asp Val Met Val Ala Gly Gly Ala Glu
180 185 190
Lys Ala Ser Thr Pro Leu Gly Val Gly Gly Phe Gly Ala Ala Arg Ala
195 200 205
Leu Ser Thr Arg Asn Asp Asn Pro Gln Ala Ala Ser Arg Pro Trp Asp
210 215 220
Lys Glu Arg Asp Gly Phe Val Leu Gly Asp Gly Ala Gly Met Leu Val
225 230 235 240
Leu Glu Glu Tyr Glu His Ala Lys Lys Arg Gly Ala Lys Ile Tyr Ala
245 250 255
Glu Leu Val Gly Phe Gly Met Ser Ser Asp Ala Tyr His Met Thr Ser
260 265 270
Pro Pro Glu Asn Gly Ala Gly Ala Ala Leu Ala Met Ala Asn Ala Leu
275 280 285
Arg Asp Ala Gly Ile Glu Ala Ser Gln Ile Gly Tyr Val Asn Ala His
290 295 300
Gly Thr Ser Thr Pro Ala Gly Asp Lys Ala Glu Ala Gln Ala Val Lys
305 310 315 320
Thr Ile Phe Gly Glu Ala Ala Ser Arg Val Leu Val Ser Ser Thr Lys
325 330 335
Ser Met Thr Gly His Leu Leu Gly Ala Ala Gly Ala Val Glu Ser Ile
340 345 350
Tyr Ser Ile Leu Ala Leu Arg Asp Gln Ala Val Pro Pro Thr Ile Asn
355 360 365
Leu Asp Asn Pro Asp Glu Gly Cys Asp Leu Asp Phe Val Pro His Glu
370 375 380
Ala Arg Gln Val Ser Gly Met Glu Tyr Thr Leu Cys Asn Ser Phe Gly
385 390 395 400
Phe Gly Gly Thr Asn Gly Ser Leu Ile Phe Lys Lys Ile
405 410
<210> 108
<211> 436
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 108
Met Gly Gln Val Leu Pro Leu Val Thr Arg Gln Gly Asp Arg Ile Ala
1 5 10 15
Ile Val Ser Gly Leu Arg Thr Pro Phe Ala Arg Gln Ala Thr Ala Phe
20 25 30
His Gly Ile Pro Ala Val Asp Leu Gly Lys Met Val Val Gly Glu Leu
35 40 45
Leu Ala Arg Ser Glu Ile Pro Ala Glu Val Ile Glu Gln Leu Val Phe
50 55 60
Gly Gln Val Val Gln Met Pro Glu Ala Pro Asn Ile Ala Arg Glu Ile
65 70 75 80
Val Leu Gly Thr Gly Met Asn Val His Thr Asp Ala Tyr Ser Val Ser
85 90 95
Arg Ala Cys Ala Thr Ser Phe Gln Ala Val Ala Asn Val Ala Glu Ser
100 105 110
Leu Met Ala Gly Thr Ile Arg Ala Gly Ile Ala Gly Gly Ala Asp Ser
115 120 125
Ser Ser Val Leu Pro Ile Gly Val Ser Lys Lys Leu Ala Arg Val Leu
130 135 140
Val Asp Val Asn Lys Ala Arg Thr Met Ser Gln Arg Leu Lys Leu Phe
145 150 155 160
Ser Arg Leu Arg Leu Arg Asp Leu Met Pro Val Pro Pro Ala Val Ala
165 170 175
Glu Tyr Ser Thr Gly Leu Arg Met Gly Asp Thr Ala Glu Gln Met Ala
180 185 190
Lys Thr Tyr Gly Ile Thr Arg Glu Gln Gln Asp Ala Leu Ala His Arg
195 200 205
Ser His Gln Arg Ala Ala Gln Ala Trp Ser Asp Gly Lys Leu Lys Glu
210 215 220
Glu Val Met Thr Ala Phe Ile Pro Pro Tyr Lys Gln Pro Leu Val Glu
225 230 235 240
Asp Asn Asn Ile Arg Gly Asn Ser Ser Leu Ala Asp Tyr Ala Lys Leu
245 250 255
Arg Pro Ala Phe Asp Arg Lys His Gly Thr Val Thr Ala Ala Asn Ser
260 265 270
Thr Pro Leu Thr Asp Gly Ala Ala Ala Val Ile Leu Met Thr Glu Ser
275 280 285
Arg Ala Lys Glu Leu Gly Leu Val Pro Leu Gly Tyr Leu Arg Ser Tyr
290 295 300
Ala Phe Thr Ala Ile Asp Val Trp Gln Asp Met Leu Leu Gly Pro Ala
305 310 315 320
Trp Ser Thr Pro Leu Ala Leu Glu Arg Ala Gly Leu Thr Met Ser Asp
325 330 335
Leu Thr Leu Ile Asp Met His Glu Ala Phe Ala Ala Gln Thr Leu Ala
340 345 350
Asn Ile Gln Leu Leu Gly Ser Glu Arg Phe Ala Arg Glu Ala Leu Gly
355 360 365
Arg Ala His Ala Thr Gly Glu Val Asp Asp Ser Lys Phe Asn Val Leu
370 375 380
Gly Gly Ser Ile Ala Tyr Gly His Pro Phe Ala Ala Thr Gly Ala Arg
385 390 395 400
Met Ile Thr Gln Thr Leu His Glu Leu Arg Arg Arg Gly Gly Gly Phe
405 410 415
Gly Leu Val Thr Ala Cys Ala Ala Gly Gly Leu Gly Ala Ala Met Val
420 425 430
Leu Glu Ala Glu
435
<210> 109
<211> 294
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 109
Met Glu Arg Val Tyr Arg Thr Asp Leu Lys Leu Leu Arg Tyr Phe Leu
1 5 10 15
Ala Val Ala Glu Glu Leu His Phe Gly Arg Ala Ala Ala Arg Leu Asn
20 25 30
Met Ser Gln Pro Pro Leu Ser Ile His Ile Lys Glu Leu Glu Asn Gln
35 40 45
Leu Gly Thr Gln Leu Phe Ile Arg His Ser Arg Ser Val Val Leu Thr
50 55 60
His Ala Gly Lys Ile Leu Met Glu Glu Ser Arg Arg Leu Leu Val Asn
65 70 75 80
Ala Asn Asn Val Leu Ala Arg Ile Glu Gln Ile Gly Arg Gly Glu Ala
85 90 95
Gly Arg Ile Glu Leu Gly Val Val Gly Thr Ala Met Trp Gly Arg Met
100 105 110
Arg Pro Val Met Arg Arg Phe Leu Arg Glu Asn Pro Asn Val Asp Val
115 120 125
Leu Phe Arg Glu Lys Met Pro Ala Met Gln Met Ala Leu Leu Glu Arg
130 135 140
Arg Glu Leu Asp Ala Gly Ile Trp Arg Met Ala Thr Glu Pro Pro Thr
145 150 155 160
Gly Phe Thr Ser Leu Arg Leu His Glu Ser Ala Phe Leu Val Ala Met
165 170 175
Pro Glu Glu His His Leu Ser Ser Phe Ser Thr Val Pro Leu Glu Ala
180 185 190
Leu Arg Asp Glu Tyr Phe Val Thr Met Pro Pro Val Tyr Thr Asp Trp
195 200 205
Asp Phe Leu Gln Arg Val Cys Gln Gln Val Gly Phe Ser Pro Val Val
210 215 220
Ile Arg Glu Val Asn Glu Pro Gln Thr Val Leu Ala Met Val Ser Met
225 230 235 240
Gly Ile Gly Ile Thr Leu Ile Ala Asp Ser Tyr Ala Gln Met Asn Trp
245 250 255
Pro Gly Val Ile Phe Arg Pro Leu Lys Gln Arg Ile Pro Ala Asp Leu
260 265 270
Tyr Ile Val Tyr Glu Thr Gln Gln Val Thr Pro Ala Met Val Lys Leu
275 280 285
Leu Ala Ala Leu Thr Gln
290
<210> 110
<211> 341
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 110
Met Lys Ala Lys Lys Gln Glu Thr Ala Ala Thr Met Lys Asp Val Ala
1 5 10 15
Leu Lys Ala Lys Val Ser Thr Ala Thr Val Ser Arg Ala Leu Met Asn
20 25 30
Pro Asp Lys Val Ser Gln Ala Thr Arg Asn Arg Val Glu Lys Ala Ala
35 40 45
Arg Glu Val Gly Tyr Leu Pro Gln Pro Met Gly Arg Asn Val Lys Arg
50 55 60
Asn Glu Ser Arg Thr Ile Leu Val Ile Val Pro Asp Ile Cys Asp Pro
65 70 75 80
Phe Phe Ser Glu Ile Ile Arg Gly Ile Glu Val Thr Ala Ala Asn His
85 90 95
Gly Tyr Leu Val Leu Ile Gly Asp Cys Ala His Gln Asn Gln Gln Glu
100 105 110
Lys Thr Phe Ile Asp Leu Ile Ile Thr Lys Gln Ile Asp Gly Met Leu
115 120 125
Leu Leu Gly Ser Arg Leu Pro Phe Asp Ala Ser Ile Glu Glu Gln Arg
130 135 140
Asn Leu Pro Pro Met Val Met Ala Asn Glu Phe Ala Pro Glu Leu Glu
145 150 155 160
Leu Pro Thr Val His Ile Asp Asn Leu Thr Ala Ala Phe Asp Ala Val
165 170 175
Asn Tyr Leu Tyr Glu Gln Gly His Lys Arg Ile Gly Cys Ile Ala Gly
180 185 190
Pro Glu Glu Met Pro Leu Cys His Tyr Arg Leu Gln Gly Tyr Val Gln
195 200 205
Ala Leu Arg Arg Cys Gly Ile Met Val Asp Pro Gln Tyr Ile Ala Arg
210 215 220
Gly Asp Phe Thr Phe Glu Ala Gly Ser Lys Ala Met Gln Gln Leu Leu
225 230 235 240
Asp Leu Pro Gln Pro Pro Thr Ala Val Phe Cys His Ser Asp Val Met
245 250 255
Ala Leu Gly Ala Leu Ser Gln Ala Lys Arg Gln Gly Leu Lys Val Pro
260 265 270
Glu Asp Leu Ser Ile Ile Gly Phe Asp Asn Ile Asp Leu Thr Gln Phe
275 280 285
Cys Asp Pro Pro Leu Thr Thr Ile Ala Gln Pro Arg Tyr Glu Ile Gly
290 295 300
Arg Glu Ala Met Leu Leu Leu Leu Asp Gln Met Gln Gly Gln His Val
305 310 315 320
Gly Ser Gly Ser Arg Leu Met Asp Cys Glu Leu Ile Ile Arg Gly Ser
325 330 335
Thr Arg Ala Leu Pro
340
<210> 111
<211> 317
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 111
Met Tyr Thr Lys Ile Ile Gly Thr Gly Ser Tyr Leu Pro Glu Gln Val
1 5 10 15
Arg Thr Asn Ala Asp Leu Glu Lys Met Val Asp Thr Ser Asp Glu Trp
20 25 30
Ile Val Thr Arg Thr Gly Ile Arg Glu Arg His Ile Ala Ala Pro Asn
35 40 45
Glu Thr Val Ser Thr Met Gly Phe Glu Ala Ala Thr Arg Ala Ile Glu
50 55 60
Met Ala Gly Ile Glu Lys Asp Gln Ile Gly Leu Ile Val Val Ala Thr
65 70 75 80
Thr Ser Ala Thr His Ala Phe Pro Ser Ala Ala Cys Gln Ile Gln Ser
85 90 95
Met Leu Gly Ile Lys Gly Cys Pro Ala Phe Asp Val Ala Ala Ala Cys
100 105 110
Ala Gly Phe Thr Tyr Ala Leu Ser Val Ala Asp Gln Tyr Val Lys Ser
115 120 125
Gly Ala Val Lys Tyr Ala Leu Val Val Gly Ser Asp Val Leu Ala Arg
130 135 140
Thr Cys Asp Pro Thr Asp Arg Gly Thr Ile Ile Ile Phe Gly Asp Gly
145 150 155 160
Ala Gly Ala Ala Val Leu Ala Ala Ser Glu Glu Pro Gly Ile Ile Ser
165 170 175
Thr His Leu His Ala Asp Gly Ser Tyr Gly Glu Leu Leu Thr Leu Pro
180 185 190
Asn Ala Asp Arg Val Asn Pro Glu Asn Ser Ile His Leu Thr Met Ala
195 200 205
Gly Asn Glu Val Phe Lys Val Ala Val Thr Glu Leu Ala His Ile Val
210 215 220
Asp Glu Thr Leu Ala Ala Asn Asn Leu Asp Arg Ser Gln Leu Asp Trp
225 230 235 240
Leu Val Pro His Gln Ala Asn Leu Arg Ile Ile Ser Ala Thr Ala Lys
245 250 255
Lys Leu Gly Met Ser Met Asp Asn Val Val Val Thr Leu Asp Arg His
260 265 270
Gly Asn Thr Ser Ala Ala Ser Val Pro Cys Ala Leu Asp Glu Ala Val
275 280 285
Arg Asp Gly Arg Ile Lys Pro Gly Gln Leu Val Leu Leu Glu Ala Phe
290 295 300
Gly Gly Gly Phe Thr Trp Gly Ser Ala Leu Val Arg Phe
305 310 315
<210> 112
<211> 548
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 112
Met Lys Lys Val Thr Ala Met Leu Phe Ser Met Ala Val Gly Leu Asn
1 5 10 15
Ala Val Ser Met Ala Ala Lys Ala Lys Ala Ser Glu Glu Gln Glu Thr
20 25 30
Asp Val Leu Leu Ile Gly Gly Gly Ile Met Ser Ala Thr Leu Gly Thr
35 40 45
Tyr Leu Arg Glu Leu Glu Pro Glu Trp Ser Met Thr Met Val Glu Arg
50 55 60
Leu Glu Gly Val Ala Gln Glu Ser Ser Asn Gly Trp Asn Asn Ala Gly
65 70 75 80
Thr Gly His Ser Ala Leu Met Glu Leu Asn Tyr Thr Pro Gln Asn Ala
85 90 95
Asp Gly Ser Ile Ser Ile Glu Lys Ala Val Ala Ile Asn Glu Ala Phe
100 105 110
Gln Ile Ser Arg Gln Phe Trp Ala His Gln Val Glu Arg Gly Val Leu
115 120 125
Arg Thr Pro Arg Ser Phe Ile Asn Thr Val Pro His Met Ser Phe Val
130 135 140
Trp Gly Glu Asp Asn Val Asn Phe Leu Arg Ala Arg Tyr Ala Ala Leu
145 150 155 160
Gln Gln Ser Ser Leu Phe Arg Gly Met Arg Tyr Ser Glu Asp His Ala
165 170 175
Gln Ile Lys Glu Trp Ala Pro Leu Val Met Glu Gly Arg Asp Pro Gln
180 185 190
Gln Lys Val Ala Ala Thr Arg Thr Glu Ile Gly Thr Asp Val Asn Tyr
195 200 205
Gly Glu Ile Thr Arg Gln Leu Ile Ala Ser Leu Gln Lys Lys Ser Asn
210 215 220
Phe Ser Leu Gln Leu Ser Ser Glu Val Arg Ala Leu Lys Arg Asn Asp
225 230 235 240
Asp Asn Thr Trp Thr Val Thr Val Ala Asp Leu Lys Asn Gly Thr Ala
245 250 255
Gln Asn Ile Arg Ala Lys Phe Val Phe Ile Gly Ala Gly Gly Ala Ala
260 265 270
Leu Lys Leu Leu Gln Glu Ser Gly Ile Pro Glu Ala Lys Asp Tyr Ala
275 280 285
Gly Phe Pro Val Gly Gly Gln Phe Leu Val Ser Glu Asn Pro Asp Val
290 295 300
Val Asn His His Leu Ala Lys Val Tyr Gly Lys Ala Ser Val Gly Ala
305 310 315 320
Pro Pro Met Ser Val Pro His Ile Asp Thr Arg Val Leu Asp Gly Lys
325 330 335
Arg Val Val Leu Phe Gly Pro Phe Ala Thr Phe Ser Thr Lys Phe Leu
340 345 350
Lys Asn Gly Ser Leu Trp Asp Leu Met Ser Ser Thr Thr Thr Ser Asn
355 360 365
Val Met Pro Met Met His Val Gly Leu Asp Asn Phe Asp Leu Val Lys
370 375 380
Tyr Leu Val Ser Gln Val Met Leu Ser Glu Glu Asp Arg Phe Glu Ala
385 390 395 400
Leu Lys Glu Tyr Tyr Pro Gln Ala Lys Lys Glu Asp Trp Arg Leu Trp
405 410 415
Gln Ala Gly Gln Arg Val Gln Ile Ile Lys Arg Asp Ala Glu Lys Gly
420 425 430
Gly Val Leu Arg Leu Gly Thr Glu Val Val Ser Asp Gln Gln Gly Thr
435 440 445
Ile Ala Ala Leu Leu Gly Ala Ser Pro Gly Ala Ser Thr Ala Ala Pro
450 455 460
Ile Met Leu Asn Leu Leu Glu Lys Val Phe Gly Asp Arg Val Ser Ser
465 470 475 480
Pro Gln Trp Gln Ala Thr Leu Lys Ala Ile Val Pro Ser Tyr Gly Arg
485 490 495
Lys Leu Asn Gly Asp Val Ala Ala Thr Glu Arg Glu Leu Gln Tyr Thr
500 505 510
Ser Glu Val Leu Gly Leu Asn Tyr Asp Lys Pro Gln Ala Ala Asp Ser
515 520 525
Thr Pro Lys Pro Gln Leu Lys Pro Gln Pro Val Gln Lys Glu Val Ala
530 535 540
Asp Ile Ala Leu
545
<210> 113
<211> 127
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 113
Met Ile Thr Gly Ile Gln Ile Thr Lys Ala Ala Asn Asp Asp Leu Leu
1 5 10 15
Asn Ser Phe Trp Leu Leu Asp Ser Glu Lys Gly Glu Ala Arg Cys Ile
20 25 30
Val Ala Lys Ala Gly Tyr Ala Glu Asp Glu Val Val Ala Val Ser Lys
35 40 45
Leu Gly Asp Ile Glu Tyr Arg Glu Val Pro Val Glu Val Lys Pro Glu
50 55 60
Val Arg Val Glu Gly Gly Gln His Leu Asn Val Asn Val Leu Arg Arg
65 70 75 80
Glu Thr Leu Glu Asp Ala Val Lys His Pro Glu Lys Tyr Pro Gln Leu
85 90 95
Thr Ile Arg Val Ser Gly Tyr Ala Val Arg Phe Asn Ser Leu Thr Pro
100 105 110
Glu Gln Gln Arg Asp Val Ile Ala Arg Thr Phe Thr Glu Ser Leu
115 120 125
<210> 114
<211> 315
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 114
Met Ser Glu Ser Leu Arg Ile Ile Phe Ala Gly Thr Pro Asp Phe Ala
1 5 10 15
Ala Arg His Leu Asp Ala Leu Leu Ser Ser Gly His Asn Val Val Gly
20 25 30
Val Phe Thr Gln Pro Asp Arg Pro Ala Gly Arg Gly Lys Lys Leu Met
35 40 45
Pro Ser Pro Val Lys Val Leu Ala Glu Glu Lys Gly Leu Pro Val Phe
50 55 60
Gln Pro Val Ser Leu Arg Pro Gln Glu Asn Gln Gln Leu Val Ala Glu
65 70 75 80
Leu Gln Ala Asp Val Met Val Val Val Ala Tyr Gly Leu Ile Leu Pro
85 90 95
Lys Ala Val Leu Glu Met Pro Arg Leu Gly Cys Ile Asn Val His Gly
100 105 110
Ser Leu Leu Pro Arg Trp Arg Gly Ala Ala Pro Ile Gln Arg Ser Leu
115 120 125
Trp Ala Gly Asp Ala Glu Thr Gly Val Thr Ile Met Gln Met Asp Val
130 135 140
Gly Leu Asp Thr Gly Asp Met Leu Tyr Lys Leu Ser Cys Pro Ile Thr
145 150 155 160
Ala Glu Asp Thr Ser Gly Thr Leu Tyr Asp Lys Leu Ala Glu Leu Gly
165 170 175
Pro Gln Gly Leu Ile Thr Thr Leu Lys Gln Leu Ala Asp Gly Thr Ala
180 185 190
Lys Pro Glu Val Gln Asp Glu Thr Leu Val Thr Tyr Ala Glu Lys Leu
195 200 205
Ser Lys Glu Glu Ala Arg Ile Asp Trp Ser Leu Ser Ala Ala Gln Leu
210 215 220
Glu Arg Cys Ile Arg Ala Phe Asn Pro Trp Pro Met Ser Trp Leu Glu
225 230 235 240
Ile Glu Gly Gln Pro Val Lys Val Trp Lys Ala Ser Val Ile Asp Thr
245 250 255
Ala Thr Asn Ala Ala Pro Gly Thr Ile Leu Glu Ala Asn Lys Gln Gly
260 265 270
Ile Gln Val Ala Thr Gly Asp Gly Ile Leu Asn Leu Leu Ser Leu Gln
275 280 285
Pro Ala Gly Lys Lys Ala Met Ser Ala Gln Asp Leu Leu Asn Ser Arg
290 295 300
Arg Glu Trp Phe Val Pro Gly Asn Arg Leu Val
305 310 315
<210> 115
<211> 245
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 115
Met Thr Leu Thr Ala Ser Ser Ser Ser Arg Ala Val Thr Asn Ser Pro
1 5 10 15
Val Val Val Ala Leu Asp Tyr His Asn Arg Asp Asp Ala Leu Ala Phe
20 25 30
Val Asp Lys Ile Asp Pro Arg Asp Cys Arg Leu Lys Val Gly Lys Glu
35 40 45
Met Phe Thr Leu Phe Gly Pro Gln Phe Val Arg Glu Leu Gln Gln Arg
50 55 60
Gly Phe Asp Ile Phe Leu Asp Leu Lys Phe His Asp Ile Pro Asn Thr
65 70 75 80
Ala Ala His Ala Val Ala Ala Ala Ala Asp Leu Gly Val Trp Met Val
85 90 95
Asn Val His Ala Ser Gly Gly Ala Arg Met Met Thr Ala Ala Arg Glu
100 105 110
Ala Leu Val Pro Phe Gly Lys Asp Ala Pro Leu Leu Ile Ala Val Thr
115 120 125
Val Leu Thr Ser Met Glu Ala Ser Asp Leu Val Asp Leu Gly Met Thr
130 135 140
Leu Ser Pro Ala Asp Tyr Ala Glu Arg Leu Ala Ala Leu Thr Gln Lys
145 150 155 160
Cys Gly Leu Asp Gly Val Val Cys Ser Ala Gln Glu Ala Val Arg Phe
165 170 175
Lys Gln Val Phe Gly Gln Glu Phe Lys Leu Val Thr Pro Gly Ile Arg
180 185 190
Pro Gln Gly Ser Glu Ala Gly Asp Gln Arg Arg Ile Met Thr Pro Glu
195 200 205
Gln Ala Leu Ser Ala Gly Val Asp Tyr Met Val Ile Gly Arg Pro Val
210 215 220
Thr Gln Ser Val Asp Pro Ala Gln Thr Leu Lys Ala Ile Asn Ala Ser
225 230 235 240
Leu Gln Arg Ser Ala
245
<210> 116
<211> 500
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 116
Met Thr Ile Phe Asp Asn Tyr Glu Val Trp Phe Val Ile Gly Ser Gln
1 5 10 15
His Leu Tyr Gly Pro Glu Thr Leu Arg Gln Val Thr Gln His Ala Glu
20 25 30
His Val Val Asn Ala Leu Asn Thr Glu Ala Lys Leu Pro Cys Lys Leu
35 40 45
Val Leu Lys Pro Leu Gly Thr Thr Pro Asp Glu Ile Thr Ala Ile Cys
50 55 60
Arg Asp Ala Asn Tyr Asp Asp Arg Cys Ala Gly Leu Val Val Trp Leu
65 70 75 80
His Thr Phe Ser Pro Ala Lys Met Trp Ile Asn Gly Leu Thr Met Leu
85 90 95
Asn Lys Pro Leu Leu Gln Phe His Thr Gln Phe Asn Ala Ala Leu Pro
100 105 110
Trp Asp Ser Ile Asp Met Asp Phe Met Asn Leu Asn Gln Thr Ala His
115 120 125
Gly Gly Arg Glu Phe Gly Phe Ile Gly Ala Arg Met Arg Gln Gln His
130 135 140
Ala Val Val Thr Gly His Trp Gln Asp Lys Gln Ala His Glu Arg Ile
145 150 155 160
Gly Ser Trp Met Arg Gln Ala Val Ser Lys Gln Asp Thr Arg His Leu
165 170 175
Lys Val Cys Arg Phe Gly Asp Asn Met Arg Glu Val Ala Val Thr Asp
180 185 190
Gly Asp Lys Val Ala Ala Gln Ile Lys Phe Gly Phe Ser Val Asn Thr
195 200 205
Trp Ala Val Gly Asp Leu Val Gln Val Val Asn Ser Ile Ser Asp Gly
210 215 220
Asp Val Asn Ala Leu Val Asp Glu Tyr Glu Ser Cys Tyr Thr Met Thr
225 230 235 240
Pro Ala Thr Gln Ile His Gly Lys Lys Arg Gln Asn Val Leu Glu Ala
245 250 255
Ala Arg Ile Glu Leu Gly Met Lys Arg Phe Leu Glu Gln Gly Gly Phe
260 265 270
His Ala Phe Thr Thr Thr Phe Glu Asp Leu His Gly Leu Lys Gln Leu
275 280 285
Pro Gly Leu Ala Val Gln Arg Leu Met Gln Gln Gly Tyr Gly Phe Ala
290 295 300
Gly Glu Gly Asp Trp Lys Thr Ala Ala Leu Leu Arg Ile Met Lys Val
305 310 315 320
Met Ser Thr Gly Leu Gln Gly Gly Thr Ser Phe Met Glu Asp Tyr Thr
325 330 335
Tyr His Phe Glu Lys Gly Asn Asp Leu Val Leu Gly Ser His Met Leu
340 345 350
Glu Val Cys Pro Ser Ile Ala Ala Glu Glu Lys Pro Ile Leu Asp Val
355 360 365
Gln His Leu Gly Ile Gly Gly Lys Asp Asp Pro Ala Arg Leu Ile Phe
370 375 380
Asn Thr Gln Thr Gly Pro Ala Ile Val Ala Ser Leu Ile Asp Leu Gly
385 390 395 400
Asp Arg Tyr Arg Leu Leu Val Asn Cys Ile Asp Thr Val Lys Thr Pro
405 410 415
His Ser Leu Pro Lys Leu Pro Val Ala Asn Ala Leu Trp Lys Ala Gln
420 425 430
Pro Asp Leu Pro Thr Ala Ser Glu Ala Trp Ile Leu Ala Gly Gly Ala
435 440 445
His His Thr Val Phe Ser His Ala Leu Asn Leu Asn Asp Met Arg Gln
450 455 460
Phe Ala Glu Met His Asp Ile Glu Ile Thr Val Ile Asp Asn Asp Thr
465 470 475 480
Arg Leu Pro Ala Phe Lys Asp Ala Leu Arg Trp Asn Glu Val Tyr Tyr
485 490 495
Gly Phe Arg Arg
500
<210> 117
<211> 334
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 117
Met Leu Asn Thr Leu Ile Val Gly Ala Ser Gly Tyr Ala Gly Ala Glu
1 5 10 15
Leu Val Thr Tyr Val Asn Arg His Pro His Met Asn Ile Thr Ala Leu
20 25 30
Thr Val Ser Ala Gln Ser Asn Asp Ala Gly Lys Leu Ile Ser Asp Leu
35 40 45
His Pro Gln Leu Lys Gly Ile Val Asp Leu Pro Leu Gln Pro Met Ser
50 55 60
Asp Ile Ser Glu Phe Ser Pro Gly Val Asp Val Val Phe Leu Ala Thr
65 70 75 80
Ala His Glu Val Ser His Asp Leu Ala Pro Gln Phe Leu Glu Ala Gly
85 90 95
Cys Val Val Phe Asp Leu Ser Gly Ala Phe Arg Val Asn Asp Ala Thr
100 105 110
Phe Tyr Glu Lys Tyr Tyr Gly Phe Thr His Gln Tyr Pro Glu Leu Leu
115 120 125
Glu Gln Ala Ala Tyr Gly Leu Ala Glu Trp Cys Gly Asn Lys Leu Lys
130 135 140
Glu Ala Asn Leu Ile Ala Val Pro Gly Cys Tyr Pro Thr Ala Ala Gln
145 150 155 160
Leu Ala Leu Lys Pro Leu Ile Asp Ala Asp Leu Leu Asp Leu Asn Gln
165 170 175
Trp Pro Val Ile Asn Ala Thr Ser Gly Val Ser Gly Ala Gly Arg Lys
180 185 190
Ala Ala Ile Ser Asn Ser Phe Cys Glu Val Ser Leu Gln Pro Tyr Gly
195 200 205
Val Phe Thr His Arg His Gln Pro Glu Ile Ala Thr His Leu Gly Ala
210 215 220
Asp Val Ile Phe Thr Pro His Leu Gly Asn Phe Pro Arg Gly Ile Leu
225 230 235 240
Glu Thr Ile Thr Cys Arg Leu Lys Ser Gly Val Thr Gln Ala Gln Val
245 250 255
Ala Gln Val Leu Gln Gln Ala Tyr Ala His Lys Pro Leu Val Arg Leu
260 265 270
Tyr Asp Lys Gly Val Pro Ala Leu Lys Asn Val Val Gly Leu Pro Phe
275 280 285
Cys Asp Ile Gly Phe Ala Val Gln Gly Glu His Leu Ile Ile Val Ala
290 295 300
Thr Glu Asp Asn Leu Leu Lys Gly Ala Ala Ala Gln Ala Val Gln Cys
305 310 315 320
Ala Asn Ile Arg Phe Gly Tyr Ala Glu Thr Gln Ser Leu Ile
325 330
<210> 118
<211> 239
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 118
Met Val Ile Lys Ala Gln Ser Pro Ala Gly Phe Ala Glu Glu Tyr Ile
1 5 10 15
Ile Glu Ser Ile Trp Asn Asn Arg Phe Pro Pro Gly Thr Ile Leu Pro
20 25 30
Ala Glu Arg Glu Leu Ser Glu Leu Ile Gly Val Thr Arg Thr Thr Leu
35 40 45
Arg Glu Val Leu Gln Arg Leu Ala Arg Asp Gly Trp Leu Thr Ile Gln
50 55 60
His Gly Lys Pro Thr Lys Val Asn Asn Phe Trp Glu Thr Ser Gly Leu
65 70 75 80
Asn Ile Leu Glu Thr Leu Ala Arg Leu Asp His Glu Ser Val Pro Gln
85 90 95
Leu Ile Asp Asn Leu Leu Ser Val Arg Thr Asn Ile Ser Thr Ile Phe
100 105 110
Ile Arg Thr Ala Phe Arg Gln His Pro Asp Lys Ala Gln Glu Val Leu
115 120 125
Ala Thr Ala Asn Glu Val Ala Asp His Ala Asp Ala Phe Ala Glu Leu
130 135 140
Asp Tyr Asn Ile Phe Arg Gly Leu Ala Phe Ala Ser Gly Asn Pro Ile
145 150 155 160
Tyr Gly Leu Ile Leu Asn Gly Met Lys Gly Leu Tyr Thr Arg Ile Gly
165 170 175
Arg His Tyr Phe Ala Asn Pro Glu Ala Arg Ser Leu Ala Leu Gly Phe
180 185 190
Tyr His Lys Leu Ser Ala Leu Cys Ser Glu Gly Ala His Asp Gln Val
195 200 205
Tyr Glu Thr Val Arg Arg Tyr Gly His Glu Ser Gly Glu Ile Trp His
210 215 220
Arg Met Gln Lys Asn Leu Pro Gly Asp Leu Ala Ile Gln Gly Arg
225 230 235
<210> 119
<211> 159
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 119
Met Phe Leu Arg Gln Glu Asp Phe Ala Thr Val Val Arg Ser Thr Pro
1 5 10 15
Leu Val Ser Leu Asp Phe Ile Val Glu Asn Ser Arg Gly Glu Phe Leu
20 25 30
Leu Gly Lys Arg Thr Asn Arg Pro Ala Gln Gly Tyr Trp Phe Val Pro
35 40 45
Gly Gly Arg Val Gln Lys Asp Glu Thr Leu Glu Ala Ala Phe Glu Arg
50 55 60
Leu Thr Met Ala Glu Leu Gly Leu Arg Leu Pro Ile Thr Ala Gly Gln
65 70 75 80
Phe Tyr Gly Val Trp Gln His Phe Tyr Asp Asp Asn Phe Ser Gly Thr
85 90 95
Asp Phe Thr Thr His Tyr Val Val Leu Gly Phe Arg Phe Arg Val Ser
100 105 110
Glu Glu Glu Leu Leu Leu Pro Asp Glu Gln His Asp Asp Tyr Arg Trp
115 120 125
Leu Thr Ser Asp Ala Leu Leu Ala Ser Asp Asn Val His Ala Asn Ser
130 135 140
Arg Ala Tyr Phe Leu Ala Glu Lys Arg Thr Gly Val Pro Gly Leu
145 150 155
<210> 120
<211> 187
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 120
Met Ile Leu Leu Ile Asp Asn Tyr Asp Ser Phe Thr Trp Asn Leu Tyr
1 5 10 15
Gln Tyr Phe Cys Glu Leu Gly Ala Asp Val Leu Val Lys Arg Asn Asp
20 25 30
Ala Leu Thr Leu Ala Asp Ile Asp Ala Leu Lys Pro Gln Lys Ile Val
35 40 45
Ile Ser Pro Gly Pro Cys Thr Pro Asp Glu Ala Gly Ile Ser Leu Asp
50 55 60
Val Ile Arg His Tyr Ala Gly Arg Leu Pro Ile Leu Gly Val Cys Leu
65 70 75 80
Gly His Gln Ala Met Ala Gln Ala Phe Gly Gly Lys Val Val Arg Ala
85 90 95
Ala Lys Val Met His Gly Lys Thr Ser Pro Ile Thr His Asn Gly Glu
100 105 110
Gly Val Phe Arg Gly Leu Ala Asn Pro Leu Thr Val Thr Arg Tyr His
115 120 125
Ser Leu Val Val Glu Pro Asp Ser Leu Pro Ala Cys Phe Asp Val Thr
130 135 140
Ala Trp Ser Glu Thr Arg Glu Ile Met Gly Ile Arg His Arg Gln Trp
145 150 155 160
Asp Leu Glu Gly Val Gln Phe His Pro Glu Ser Ile Leu Ser Glu Gln
165 170 175
Gly His Gln Leu Leu Ala Asn Phe Leu His Arg
180 185
<210> 121
<211> 456
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 121
Met Glu Leu Ser Ser Leu Thr Ala Val Ser Pro Val Asp Gly Arg Tyr
1 5 10 15
Gly Asp Lys Val Ser Ala Leu Arg Gly Ile Phe Ser Glu Tyr Gly Leu
20 25 30
Leu Lys Phe Arg Val Gln Val Glu Val Arg Trp Leu Gln Lys Leu Ala
35 40 45
Ala His Ala Ala Ile Lys Glu Val Pro Ala Phe Ala Ala Asp Ala Ile
50 55 60
Gly Tyr Leu Asp Ala Ile Val Ala Ser Phe Ser Glu Glu Asp Ala Ala
65 70 75 80
Arg Ile Lys Thr Ile Glu Arg Thr Thr Asn His Asp Val Lys Ala Val
85 90 95
Glu Tyr Phe Leu Lys Glu Lys Val Ala Glu Ile Pro Glu Leu His Ala
100 105 110
Val Ser Glu Phe Ile His Phe Ala Cys Thr Ser Glu Asp Ile Asn Asn
115 120 125
Leu Ser His Ala Leu Met Leu Lys Thr Ala Arg Asp Glu Val Ile Leu
130 135 140
Pro Tyr Trp Arg Gln Leu Ile Asp Gly Ile Lys Asp Leu Ala Val Gln
145 150 155 160
Tyr Arg Asp Ile Pro Leu Leu Ser Arg Thr His Gly Gln Pro Ala Thr
165 170 175
Pro Ser Thr Ile Gly Lys Glu Met Ala Asn Val Ala Tyr Arg Met Glu
180 185 190
Arg Gln Tyr Arg Gln Leu Asn Gln Val Glu Ile Leu Gly Lys Ile Asn
195 200 205
Gly Ala Val Gly Asn Tyr Asn Ala His Ile Ala Ala Tyr Pro Glu Val
210 215 220
Asp Trp His Gln Phe Ser Glu Glu Phe Val Thr Ser Leu Gly Ile Gln
225 230 235 240
Trp Asn Pro Tyr Thr Thr Gln Ile Glu Pro His Asp Tyr Ile Ala Glu
245 250 255
Leu Phe Asp Cys Val Ala Arg Phe Asn Thr Ile Leu Ile Asp Phe Asp
260 265 270
Arg Asp Val Trp Gly Tyr Ile Ala Leu Asn His Phe Lys Gln Lys Thr
275 280 285
Ile Ala Gly Glu Ile Gly Ser Ser Thr Met Pro His Lys Val Asn Pro
290 295 300
Ile Asp Phe Glu Asn Ser Glu Gly Asn Leu Gly Leu Ser Asn Ala Val
305 310 315 320
Leu Gln His Leu Ala Ser Lys Leu Pro Val Ser Arg Trp Gln Arg Asp
325 330 335
Leu Thr Asp Ser Thr Val Leu Arg Asn Leu Gly Val Gly Ile Gly Tyr
340 345 350
Ala Leu Ile Ala Tyr Gln Ser Thr Leu Lys Gly Val Ser Lys Leu Glu
355 360 365
Val Asn Arg Asp His Leu Leu Asp Glu Leu Asp His Asn Trp Glu Val
370 375 380
Leu Ala Glu Pro Ile Gln Thr Val Met Arg Arg Tyr Gly Ile Glu Lys
385 390 395 400
Pro Tyr Glu Lys Leu Lys Glu Leu Thr Arg Gly Lys Arg Val Asp Ala
405 410 415
Glu Gly Met Lys Gln Phe Ile Asp Gly Leu Ala Leu Pro Glu Glu Glu
420 425 430
Lys Ala Arg Leu Lys Ala Met Thr Pro Ala Asn Tyr Ile Gly Arg Ala
435 440 445
Ile Thr Met Val Asp Glu Leu Lys
450 455
<210> 122
<211> 156
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 122
Met Thr Asp Val Leu Leu Cys Val Gly Asn Ser Met Met Gly Asp Asp
1 5 10 15
Gly Ala Gly Pro Leu Leu Ala Glu Lys Cys Ala Ala Ala Pro Lys Gly
20 25 30
Asn Trp Val Val Ile Asp Gly Gly Ser Ala Pro Glu Asn Asp Ile Val
35 40 45
Ala Ile Arg Glu Leu Arg Pro Thr Arg Leu Leu Ile Val Asp Ala Thr
50 55 60
Asp Met Gly Leu Asn Pro Gly Glu Ile Arg Ile Ile Asp Pro Asp Asp
65 70 75 80
Ile Ala Glu Met Phe Met Met Thr Thr His Asn Met Pro Leu Asn Tyr
85 90 95
Leu Ile Asp Gln Leu Lys Glu Asp Ile Gly Glu Val Ile Phe Leu Gly
100 105 110
Ile Gln Pro Asp Ile Val Gly Phe Tyr Tyr Pro Met Thr Gln Pro Ile
115 120 125
Lys Asp Ala Val Glu Thr Val Tyr Gln Arg Leu Glu Gly Trp Glu Gly
130 135 140
Asn Gly Gly Phe Ala Gln Leu Ala Val Glu Glu Glu
145 150 155
<210> 123
<211> 1774
<212> PRT
<213> 荨麻青霉(Penicillium urticae)
<400> 123
Met His Ser Ala Ala Thr Ser Thr Tyr Pro Ser Gly Lys Thr Ser Pro
1 5 10 15
Ala Pro Val Gly Thr Pro Gly Thr Glu Tyr Ser Glu Tyr Glu Phe Ser
20 25 30
Asn Asp Val Ala Val Val Gly Met Ala Cys Arg Val Ala Gly Gly Asn
35 40 45
His Asn Pro Glu Leu Leu Trp Gln Ser Leu Leu Ser Gln Lys Ser Ala
50 55 60
Met Gly Glu Ile Pro Pro Met Arg Trp Glu Pro Tyr Tyr Arg Arg Asp
65 70 75 80
Ala Arg Asn Glu Lys Phe Leu Lys Asn Thr Thr Ser Arg Gly Tyr Phe
85 90 95
Leu Asp Arg Leu Glu Asp Phe Asp Cys Gln Phe Phe Gly Ile Ser Pro
100 105 110
Lys Glu Ala Glu Gln Met Asp Pro Gln Gln Arg Val Ser Leu Glu Val
115 120 125
Ala Ser Glu Ala Leu Glu Asp Ala Gly Ile Pro Ala Lys Ser Leu Ser
130 135 140
Gly Ser Asp Thr Ala Val Phe Trp Gly Val Asn Ser Asp Asp Tyr Ser
145 150 155 160
Lys Leu Val Leu Glu Asp Leu Pro Asn Val Glu Ala Trp Met Gly Ile
165 170 175
Gly Thr Ala Tyr Cys Gly Val Pro Asn Arg Ile Ser Tyr His Leu Asn
180 185 190
Leu Met Gly Pro Ser Thr Ala Val Asp Ala Ala Cys Ala Ser Ser Leu
195 200 205
Val Ala Ile His His Gly Val Gln Ala Ile Arg Leu Gly Glu Ser Lys
210 215 220
Val Ala Ile Val Gly Gly Val Asn Ala Leu Cys Gly Pro Gly Leu Thr
225 230 235 240
Arg Val Leu Asp Lys Ala Gly Ala Ile Ser Ser Asp Gly Ser Cys Lys
245 250 255
Ser Phe Asp Asp Asp Ala His Gly Tyr Ala Arg Gly Glu Gly Ala Gly
260 265 270
Ala Leu Val Leu Lys Ser Leu His Arg Ala Leu Leu Asp His Asp Asn
275 280 285
Val Leu Ala Val Ile Lys Gly Ser Ala Val Cys Gln Asp Gly Lys Thr
290 295 300
Asn Gly Ile Met Ala Pro Asn Ser Val Ala Gln Gln Leu Ala Ala Asn
305 310 315 320
Asn Ala Leu Ser Ala Ala Asn Ile Asp Pro His Thr Val Arg Tyr Val
325 330 335
Glu Ala His Ala Thr Ser Thr Pro Leu Gly Asp Pro Thr Glu Ile Ser
340 345 350
Ala Ile Ala Ser Val Tyr Gly Ala Asp Arg Pro Ala Asp Asp Pro Cys
355 360 365
Tyr Ile Gly Ser Ile Lys Pro Asn Ile Gly His Leu Glu Ala Gly Ala
370 375 380
Gly Val Met Gly Phe Ile Lys Ala Val Leu Ala Ile Gln Lys Gly Val
385 390 395 400
Leu Pro Pro Gln Ala Asn Leu Thr Lys Leu Asn Ser Arg Ile Asp Trp
405 410 415
Lys Thr Ala Gly Val Lys Val Val Gln Glu Ala Thr Pro Trp Pro Glu
420 425 430
Ser Asp Pro Ile Arg Arg Ala Gly Val Cys Ser Tyr Gly Tyr Gly Gly
435 440 445
Thr Val Ser His Ala Val Ile Glu Glu Phe Ser Pro Ile Leu Gln Pro
450 455 460
Asp Pro Leu Gly Asn Gly Ala Val Ser Gly Pro Gly Leu Leu Leu Leu
465 470 475 480
Ser Gly Pro Gln Glu Lys Arg Leu Ala Leu Gln Ala Lys Thr Leu Arg
485 490 495
Asp Trp Met Thr Ala Glu Gly Lys Asp His Asn Leu Ser Asp Ile Leu
500 505 510
Thr Thr Leu Ala Thr Arg Arg Asp His His Asp Tyr Arg Ala Ala Leu
515 520 525
Val Val Asp Asp Tyr Arg Asp Ala Glu Gln Val Leu Gln Ser Leu Ala
530 535 540
Asn Gly Val Asp His Thr Phe Thr Thr Gln Ser Arg Val Leu Gly Ser
545 550 555 560
Asp Ile Ser Lys Asp Val Val Trp Val Phe Ser Gly His Gly Ala Gln
565 570 575
Trp Pro Asp Met Gly Lys Gln Leu Ile His Asn Pro Val Phe Phe Ala
580 585 590
Ala Ile Gln Pro Leu Asp Glu Leu Ile Gln Ala Glu Ile Gly Leu Ser
595 600 605
Pro Ile Glu Leu Leu Arg Thr Gly Asp Phe Glu Ser Ser Asp Arg Val
610 615 620
Gln Ile Leu Thr Tyr Val Met Gln Ile Gly Leu Ser Ala Leu Leu Gln
625 630 635 640
Ser Asn Gly Ile Thr Pro Gln Ala Val Ile Gly His Ser Val Gly Glu
645 650 655
Ile Ala Ala Ser Val Val Ala Gly Ala Leu Ser Pro Ala Glu Gly Ala
660 665 670
Leu Ile Val Thr Arg Arg Ala Leu Leu Tyr Arg Gln Val Met Gly Lys
675 680 685
Gly Gly Met Ile Leu Val Asn Leu Pro Ser Ala Glu Thr Glu Glu Ile
690 695 700
Leu Gly Ser Arg Ser Asp Leu Val Val Ala Ile Asp Ser Ser Pro Ser
705 710 715 720
Ser Cys Val Val Ala Gly Asp Lys Glu Leu Val Ala Glu Thr Ala Glu
725 730 735
Ala Leu Lys Ala Arg Gly Val Lys Thr Phe Thr Val Lys Ser Asp Ile
740 745 750
Ala Phe His Ser Pro Thr Leu Asn Gly Leu Val Asp Pro Leu Arg Asp
755 760 765
Val Leu Ala Glu Thr Leu Ser Pro Val Ser Pro Asn Val Lys Leu Tyr
770 775 780
Ser Thr Ala Leu Ala Asp Pro Arg Gly Gln Asp Leu Arg Asp Val Glu
785 790 795 800
Tyr Trp Ala Gly Asn Met Val Asn Arg Val Arg Leu Thr Ser Ala Val
805 810 815
Lys Ala Ala Val Glu Asp Gly Tyr Arg Leu Phe Leu Glu Val Ser Thr
820 825 830
His Pro Val Val Ser His Ser Ile Asn Glu Thr Leu Met Asp Ala Gly
835 840 845
Met Glu Asp Phe Ala Val Ile Pro Thr Leu Leu Arg Lys Lys Pro Thr
850 855 860
Glu Lys His Ile Leu His Ser Ile Ala Gln Leu His Cys Arg Gly Ala
865 870 875 880
Glu Val Asn Trp Ala Ala Gln Met Pro Gly Arg Trp Ala Thr Gly Val
885 890 895
Pro Thr Thr Thr Trp Met His Lys Pro Ile Trp Arg Lys Ile Glu Thr
900 905 910
Ala Pro Leu His Thr Gly Leu Thr His Asp Val Glu Lys His Thr Leu
915 920 925
Leu Gly Gln Arg Ile Pro Val Pro Gly Thr Asp Thr Tyr Val Tyr Thr
930 935 940
Thr Arg Leu Asp Asn Asp Thr Lys Pro Phe Pro Gly Ser His Pro Leu
945 950 955 960
His Gly Thr Glu Ile Val Pro Ala Ala Gly Leu Ile Asn Thr Phe Leu
965 970 975
Lys Gly Thr Gly Gly Gln Met Leu Gln Asn Val Val Leu Arg Val Pro
980 985 990
Val Ala Ile Asn Ala Pro Arg Ser Val Gln Val Val Val Gln Gln Asp
995 1000 1005
Gln Val Lys Val Val Ser Arg Leu Ile Pro Ser Glu Pro Ser Gln
1010 1015 1020
Leu Asp Asp Asp Ala Ser Trp Val Thr His Thr Thr Ala Tyr Trp
1025 1030 1035
Asp Arg Lys Val Ala Gly Ser Glu Asp Arg Ile Asp Phe Ala Ala
1040 1045 1050
Val Lys Ser Arg Leu Val Thr Lys Leu Ala Asp Asn Phe Ser Ile
1055 1060 1065
Asp Tyr Leu Asp Lys Val Gly Val Ser Ala Met Gly Phe Pro Trp
1070 1075 1080
Ala Val Thr Glu His Tyr Arg Asn Asp Lys Glu Met Leu Ala Arg
1085 1090 1095
Val Asp Val Asn Pro Ala Ile Ser Gly Asp Ala Pro Leu Pro Trp
1100 1105 1110
Asp Ser Ser Ser Trp Ala Pro Val Leu Asp Ala Ala Thr Ser Val
1115 1120 1125
Gly Ser Thr Ile Phe Pro Thr Pro Ala Leu Arg Met Pro Ala Gln
1130 1135 1140
Ile Glu Arg Val Glu Val Phe Thr Ser Gln Asp Pro Pro Lys Ile
1145 1150 1155
Ser Trp Leu Tyr Val Gln Glu Ala Ser Asp Ser Val Pro Thr Ser
1160 1165 1170
His Val Ser Val Val Ser Glu Ala Gly Glu Val Leu Ala Lys Phe
1175 1180 1185
Thr Ala Met Arg Phe Ser Glu Ile Glu Gly Thr Pro Gly Val Ser
1190 1195 1200
Gly Ser Met Glu Ser Leu Val His Gln Ile Ala Trp Pro Pro Ala
1205 1210 1215
Thr Pro Ala Glu Glu Pro Leu Ser Ile Glu Thr Val Ile Leu Val
1220 1225 1230
Ser Pro Asp Ala Thr Thr Arg Ala Leu Tyr Ala Ala Ser Leu Pro
1235 1240 1245
Thr Arg Val Asn Ser Phe Gln Phe Ser Ser Thr Gln Glu Phe Phe
1250 1255 1260
Ser Asn Ala Ser Ser Leu Pro Leu Glu Lys Gly Thr Val Val Thr
1265 1270 1275
Tyr Ile Pro Gly Glu Val Ala Ser Leu Ala Glu Val Pro Ala Ala
1280 1285 1290
Ser Glu Ser Phe Thr Trp Asn Leu Leu Glu Leu Ile Lys Phe Thr
1295 1300 1305
Val Asn Gly Ser Leu Pro Ile Lys Val Phe Thr Leu Thr Ala Asn
1310 1315 1320
Ile Gly Glu Gly Gln Thr Pro Thr Ala Leu Ala Gln Ser Pro Leu
1325 1330 1335
Tyr Gly Leu Ala Arg Val Ile Ala Ser Glu His Pro Asp Leu Gly
1340 1345 1350
Thr Leu Ile Asp Val Glu Glu Pro Val Ile Pro Leu Ser Thr Met
1355 1360 1365
Arg Tyr Ile Gln Gly Ala Asp Ile Ile Arg Ile Asn Asp Gly Ile
1370 1375 1380
Ala Arg Thr Ser Arg Phe Arg Ser Leu Pro Arg Asn Lys Leu Leu
1385 1390 1395
Pro Ala Ser Glu Gly Pro Arg Leu Leu Pro Arg Pro Glu Gly Thr
1400 1405 1410
Tyr Leu Ile Thr Gly Gly Leu Gly Val Leu Gly Leu Glu Val Ala
1415 1420 1425
Asp Phe Leu Val Glu Lys Gly Ala Arg Arg Leu Leu Leu Ile Ser
1430 1435 1440
Arg Arg Ala Leu Pro Pro Arg Arg Thr Trp Asp Gln Val Ser Glu
1445 1450 1455
Asp Leu Gln Pro Thr Ile Ala Lys Ile Arg Leu Leu Glu Ser Arg
1460 1465 1470
Gly Ala Ser Val His Val Leu Pro Leu Asp Ile Thr Lys Pro Asp
1475 1480 1485
Ala Val Glu Gln Leu Thr Thr Ala Leu Asp Arg Leu Ser Leu Pro
1490 1495 1500
Ser Val Gln Gly Val Val His Ala Ala Gly Val Leu Asp Asn Glu
1505 1510 1515
Leu Val Met Gln Thr Thr Arg Asp Ala Phe Asn Arg Val Leu Ala
1520 1525 1530
Pro Lys Ile Ala Gly Ala Leu Ala Leu His Glu Val Phe Pro Pro
1535 1540 1545
Lys Ser Val Asp Phe Phe Val Met Phe Ser Ser Cys Gly Asn Leu
1550 1555 1560
Val Gly Phe Thr Gly Gln Ala Ser Tyr Gly Ser Gly Asn Ala Phe
1565 1570 1575
Leu Asp Thr Leu Ala Thr His Arg Ala Arg Leu Gly Asp Ala Ala
1580 1585 1590
Val Ser Phe Gln Trp Thr Ser Trp Arg Gly Leu Gly Met Gly Ala
1595 1600 1605
Ser Thr Asp Phe Ile Asn Ala Glu Leu Glu Ser Lys Gly Ile Thr
1610 1615 1620
Asp Val Thr Arg Asp Glu Ala Phe Ala Ala Trp Gln His Leu Ala
1625 1630 1635
Lys Tyr Asp Met Asp His Gly Val Val Leu Arg Ser Arg Ala Phe
1640 1645 1650
Glu Asp Gly Glu Pro Ile Pro Val Ser Ile Leu Asn Asp Ile Ala
1655 1660 1665
Val Arg Arg Val Gly Thr Val Ser Asn Thr Ser Pro Ala Ala Ala
1670 1675 1680
Gly Ser Ser Asp Ala Val Pro Thr Ser Gly Pro Glu Leu Lys Ala
1685 1690 1695
Tyr Leu Asp Glu Lys Ile Arg Gly Cys Val Ala Lys Val Leu Gln
1700 1705 1710
Met Thr Ala Glu Asp Val Asp Ser Lys Ala Ala Leu Ala Asp Leu
1715 1720 1725
Gly Val Asp Ser Val Met Thr Val Thr Leu Arg Arg Gln Leu Gln
1730 1735 1740
Leu Thr Leu Lys Ile Ala Val Pro Pro Thr Leu Thr Trp Ser His
1745 1750 1755
Pro Thr Val Ser His Leu Ala Val Trp Phe Ala Glu Lys Leu Ala
1760 1765 1770
Lys
<210> 124
<211> 224
<212> PRT
<213> 枯草芽孢杆菌(Bacillus subtilis)
<400> 124
Met Lys Ile Tyr Gly Ile Tyr Met Asp Arg Pro Leu Ser Gln Glu Glu
1 5 10 15
Asn Glu Arg Phe Met Ser Phe Ile Ser Pro Glu Lys Arg Glu Lys Cys
20 25 30
Arg Arg Phe Tyr His Lys Glu Asp Ala His Arg Thr Leu Leu Gly Asp
35 40 45
Val Leu Val Arg Ser Val Ile Ser Arg Gln Tyr Gln Leu Asp Lys Ser
50 55 60
Asp Ile Arg Phe Ser Thr Gln Glu Tyr Gly Lys Pro Cys Ile Pro Asp
65 70 75 80
Leu Pro Asp Ala His Phe Asn Ile Ser His Ser Gly Arg Trp Val Ile
85 90 95
Cys Ala Phe Asp Ser Gln Pro Ile Gly Ile Asp Ile Glu Lys Thr Lys
100 105 110
Pro Ile Ser Leu Glu Ile Ala Lys Arg Phe Phe Ser Lys Thr Glu Tyr
115 120 125
Ser Asp Leu Leu Ala Lys Asp Lys Asp Glu Gln Thr Asp Tyr Phe Tyr
130 135 140
His Leu Trp Ser Met Lys Glu Ser Phe Ile Lys Gln Glu Gly Lys Gly
145 150 155 160
Leu Ser Leu Pro Leu Asp Ser Phe Ser Val Arg Leu His Gln Asp Gly
165 170 175
Gln Val Ser Ile Glu Leu Pro Asp Ser His Ser Pro Cys Tyr Ile Lys
180 185 190
Thr Tyr Glu Val Asp Pro Gly Tyr Lys Met Ala Val Cys Ala Val His
195 200 205
Pro Asp Phe Pro Glu Asp Ile Thr Met Val Ser Tyr Glu Glu Leu Leu
210 215 220
<210> 125
<211> 391
<212> PRT
<213> 未知
<220>
<223> 掌叶大黄(Rheum palmatum)
<400> 125
Met Ala Asp Val Leu Gln Glu Ile Arg Asn Ser Gln Lys Ala Ser Gly
1 5 10 15
Pro Ala Thr Val Leu Ala Ile Gly Thr Ala His Pro Pro Thr Cys Tyr
20 25 30
Pro Gln Ala Asp Tyr Pro Asp Phe Tyr Phe Arg Val Cys Lys Ser Glu
35 40 45
His Met Thr Lys Leu Lys Lys Lys Met Gln Phe Ile Cys Asp Arg Ser
50 55 60
Gly Ile Arg Gln Arg Phe Met Phe His Thr Glu Glu Asn Leu Gly Lys
65 70 75 80
Asn Pro Gly Met Cys Thr Phe Asp Gly Pro Ser Leu Asn Ala Arg Gln
85 90 95
Asp Met Leu Ile Met Glu Val Pro Lys Leu Gly Ala Glu Ala Ala Glu
100 105 110
Lys Ala Ile Lys Glu Trp Gly Gln Asp Lys Ser Arg Ile Thr His Leu
115 120 125
Ile Phe Cys Thr Thr Thr Ser Asn Asp Met Pro Gly Ala Asp Tyr Gln
130 135 140
Phe Ala Thr Leu Phe Gly Leu Asn Pro Gly Val Ser Arg Thr Met Val
145 150 155 160
Tyr Gln Gln Gly Cys Phe Ala Gly Gly Thr Val Leu Arg Leu Val Lys
165 170 175
Asp Ile Ala Glu Asn Asn Lys Gly Ala Arg Val Leu Val Val Cys Ser
180 185 190
Glu Ile Val Ala Phe Ala Phe Arg Gly Pro His Glu Asp His Ile Asp
195 200 205
Ser Leu Ile Gly Gln Leu Leu Phe Gly Asp Gly Ala Ala Ala Leu Val
210 215 220
Val Gly Thr Asp Ile Asp Glu Ser Val Glu Arg Pro Ile Phe Gln Ile
225 230 235 240
Met Ser Ala Thr Gln Ala Thr Ile Pro Asn Ser Leu His Thr Met Ala
245 250 255
Leu His Leu Thr Glu Ala Gly Leu Thr Phe His Leu Ser Lys Glu Val
260 265 270
Pro Lys Val Val Ser Asp Asn Met Glu Glu Leu Met Leu Glu Ala Phe
275 280 285
Lys Pro Leu Gly Ile Thr Asp Trp Asn Ser Ile Phe Trp Gln Val His
290 295 300
Pro Gly Gly Arg Ala Ile Leu Asp Lys Ile Glu Glu Lys Leu Glu Leu
305 310 315 320
Thr Lys Asp Lys Met Arg Asp Ser Arg Tyr Ile Leu Ser Glu Tyr Gly
325 330 335
Asn Leu Thr Ser Ala Cys Val Leu Phe Val Met Asp Glu Met Arg Lys
340 345 350
Arg Ser Phe Arg Glu Gly Lys Gln Thr Thr Gly Asp Gly Tyr Glu Trp
355 360 365
Gly Val Ala Ile Gly Leu Gly Pro Gly Leu Thr Val Glu Thr Val Val
370 375 380
Leu Arg Ser Val Pro Ile Pro
385 390
<210> 126
<211> 403
<212> PRT
<213> 木立芦荟
<400> 126
Met Gly Ser Leu Ser Asp Ser Thr Pro Leu Met Lys Asp Val Gln Gly
1 5 10 15
Ile Arg Lys Ala Gln Lys Ala Asp Gly Thr Ala Thr Val Met Ala Ile
20 25 30
Gly Thr Ala His Pro Pro His Ile Ile Ser Gln Asp Ser Tyr Ala Asp
35 40 45
Phe Tyr Phe Arg Val Thr Asn Ser Glu His Lys Val Glu Leu Lys Lys
50 55 60
Lys Phe Asp Arg Ile Cys Lys Lys Thr Met Ile Gly Lys Arg Tyr Phe
65 70 75 80
Asn Phe Asp Glu Glu Phe Leu Lys Lys Tyr Pro Asn Ile Thr Ser Phe
85 90 95
Asp Lys Pro Ser Leu Asn Asp Arg His Asp Ile Cys Ile Pro Gly Val
100 105 110
Pro Ala Leu Gly Ala Glu Ala Ala Val Lys Ala Ile Glu Glu Trp Gly
115 120 125
Arg Pro Lys Ser Glu Ile Thr His Leu Val Phe Cys Thr Ser Gly Gly
130 135 140
Val Asp Met Pro Ser Ala Asp Phe Gln Cys Ala Lys Leu Leu Gly Leu
145 150 155 160
Arg Thr Asn Val Asn Lys Tyr Cys Ile Tyr Met Gln Gly Cys Tyr Ala
165 170 175
Gly Gly Thr Val Met Arg Tyr Ala Lys Asp Leu Ala Glu Asn Asn Arg
180 185 190
Gly Ala Arg Val Leu Met Val Cys Ala Glu Leu Thr Ile Ile Ala Leu
195 200 205
Arg Gly Pro Asn Asp Ser His Ile Asp Asn Ala Ile Gly Asn Ser Leu
210 215 220
Phe Gly Asp Gly Ala Ala Ala Leu Ile Val Gly Ser Asp Pro Ile Ile
225 230 235 240
Gly Val Glu Lys Pro Met Phe Glu Ile Val Cys Ala Lys Gln Thr Val
245 250 255
Ile Pro Asn Ser Glu Glu Val Ile His Leu His Leu Arg Glu Ser Gly
260 265 270
Leu Met Phe Tyr Met Thr Lys Asp Ser Ala Ala Thr Ile Ser Asn Asn
275 280 285
Ile Glu Ala Cys Leu Val Asp Val Phe Lys Ser Val Gly Met Thr Pro
290 295 300
Pro Glu Asp Trp Asn Ser Leu Phe Trp Ile Pro His Pro Gly Gly Arg
305 310 315 320
Ala Ile Leu Asp Gln Val Glu Ala Lys Leu Lys Leu Arg Pro Glu Lys
325 330 335
Phe Ser Ala Thr Arg Thr Val Leu Trp Asp Tyr Gly Asn Met Ile Ser
340 345 350
Ala Cys Val Leu Tyr Ile Leu Asp Glu Met Arg Arg Lys Ser Ala Ala
355 360 365
Glu Gly Leu Glu Thr Tyr Gly Glu Gly Leu Glu Trp Gly Val Leu Leu
370 375 380
Gly Phe Gly Pro Gly Met Thr Ile Glu Thr Ile Leu Leu His Ser Leu
385 390 395 400
Pro Pro Val
<210> 127
<211> 510
<212> PRT
<213> 未知
<220>
<223> 西班牙糖丝菌(Saccharothrix espanaensis)
<400> 127
Met Thr Gln Val Val Glu Arg Gln Ala Asp Arg Leu Ser Ser Arg Glu
1 5 10 15
Tyr Leu Ala Arg Val Val Arg Ser Ala Gly Trp Asp Ala Gly Leu Thr
20 25 30
Ser Cys Thr Asp Glu Glu Ile Val Arg Met Gly Ala Ser Ala Arg Thr
35 40 45
Ile Glu Glu Tyr Leu Lys Ser Asp Lys Pro Ile Tyr Gly Leu Thr Gln
50 55 60
Gly Phe Gly Pro Leu Val Leu Phe Asp Ala Asp Ser Glu Leu Glu Gln
65 70 75 80
Gly Gly Ser Leu Ile Ser His Leu Gly Thr Gly Gln Gly Ala Pro Leu
85 90 95
Ala Pro Glu Val Ser Arg Leu Ile Leu Trp Leu Arg Ile Gln Asn Met
100 105 110
Arg Lys Gly Tyr Ser Ala Val Ser Pro Val Phe Trp Gln Lys Leu Ala
115 120 125
Asp Leu Trp Asn Lys Gly Phe Thr Pro Ala Ile Pro Arg His Gly Thr
130 135 140
Val Ser Ala Ser Gly Asp Leu Gln Pro Leu Ala His Ala Ala Leu Ala
145 150 155 160
Phe Thr Gly Val Gly Glu Ala Trp Thr Arg Asp Ala Asp Gly Arg Trp
165 170 175
Ser Thr Val Pro Ala Val Asp Ala Leu Ala Ala Leu Gly Ala Glu Pro
180 185 190
Phe Asp Trp Pro Val Arg Glu Ala Leu Ala Phe Val Asn Gly Thr Gly
195 200 205
Ala Ser Leu Ala Val Ala Val Leu Asn His Arg Ser Ala Leu Arg Leu
210 215 220
Val Arg Ala Cys Ala Val Leu Ser Ala Arg Leu Ala Thr Leu Leu Gly
225 230 235 240
Ala Asn Pro Glu His Tyr Asp Val Gly His Gly Val Ala Arg Gly Gln
245 250 255
Val Gly Gln Leu Thr Ala Ala Glu Trp Ile Arg Gln Gly Leu Pro Arg
260 265 270
Gly Met Val Arg Asp Gly Ser Arg Pro Leu Gln Glu Pro Tyr Ser Leu
275 280 285
Arg Cys Ala Pro Gln Val Leu Gly Ala Val Leu Asp Gln Leu Asp Gly
290 295 300
Ala Gly Asp Val Leu Ala Arg Glu Val Asp Gly Cys Gln Asp Asn Pro
305 310 315 320
Ile Thr Tyr Glu Gly Glu Leu Leu His Gly Gly Asn Phe His Ala Met
325 330 335
Pro Val Gly Phe Ala Ser Asp Gln Ile Gly Leu Ala Met His Met Ala
340 345 350
Ala Tyr Leu Ala Glu Arg Gln Leu Gly Leu Leu Val Ser Pro Val Thr
355 360 365
Asn Gly Asp Leu Pro Pro Met Leu Thr Pro Arg Ala Gly Arg Gly Ala
370 375 380
Gly Leu Ala Gly Val Gln Ile Ser Ala Thr Ser Phe Val Ser Arg Ile
385 390 395 400
Arg Gln Leu Val Phe Pro Ala Ser Leu Thr Thr Leu Pro Thr Asn Gly
405 410 415
Trp Asn Gln Asp His Val Pro Met Ala Leu Asn Gly Ala Asn Ser Val
420 425 430
Phe Glu Ala Leu Glu Leu Gly Trp Leu Thr Val Gly Ser Leu Ala Val
435 440 445
Gly Val Ala Gln Leu Ala Ala Met Thr Gly His Ala Ala Glu Gly Val
450 455 460
Trp Ala Glu Leu Ala Gly Ile Cys Pro Pro Leu Asp Ala Asp Arg Pro
465 470 475 480
Leu Gly Ala Glu Val Arg Ala Ala Arg Asp Leu Leu Ser Ala His Ala
485 490 495
Asp Gln Leu Leu Val Asp Glu Ala Asp Gly Lys Asp Phe Gly
500 505 510
<210> 128
<211> 561
<212> PRT
<213> 拟南芥(Arabidopsis thaliana)
<400> 128
Met Ala Pro Gln Glu Gln Ala Val Ser Gln Val Met Glu Lys Gln Ser
1 5 10 15
Asn Asn Asn Asn Ser Asp Val Ile Phe Arg Ser Lys Leu Pro Asp Ile
20 25 30
Tyr Ile Pro Asn His Leu Ser Leu His Asp Tyr Ile Phe Gln Asn Ile
35 40 45
Ser Glu Phe Ala Thr Lys Pro Cys Leu Ile Asn Gly Pro Thr Gly His
50 55 60
Val Tyr Thr Tyr Ser Asp Val His Val Ile Ser Arg Gln Ile Ala Ala
65 70 75 80
Asn Phe His Lys Leu Gly Val Asn Gln Asn Asp Val Val Met Leu Leu
85 90 95
Leu Pro Asn Cys Pro Glu Phe Val Leu Ser Phe Leu Ala Ala Ser Phe
100 105 110
Arg Gly Ala Thr Ala Thr Ala Ala Asn Pro Phe Phe Thr Pro Ala Glu
115 120 125
Ile Ala Lys Gln Ala Lys Ala Ser Asn Thr Lys Leu Ile Ile Thr Glu
130 135 140
Ala Arg Tyr Val Asp Lys Ile Lys Pro Leu Gln Asn Asp Asp Gly Val
145 150 155 160
Val Ile Val Cys Ile Asp Asp Asn Glu Ser Val Pro Ile Pro Glu Gly
165 170 175
Cys Leu Arg Phe Thr Glu Leu Thr Gln Ser Thr Thr Glu Ala Ser Glu
180 185 190
Val Ile Asp Ser Val Glu Ile Ser Pro Asp Asp Val Val Ala Leu Pro
195 200 205
Tyr Ser Ser Gly Thr Thr Gly Leu Pro Lys Gly Val Met Leu Thr His
210 215 220
Lys Gly Leu Val Thr Ser Val Ala Gln Gln Val Asp Gly Glu Asn Pro
225 230 235 240
Asn Leu Tyr Phe His Ser Asp Asp Val Ile Leu Cys Val Leu Pro Met
245 250 255
Phe His Ile Tyr Ala Leu Asn Ser Ile Met Leu Cys Gly Leu Arg Val
260 265 270
Gly Ala Ala Ile Leu Ile Met Pro Lys Phe Glu Ile Asn Leu Leu Leu
275 280 285
Glu Leu Ile Gln Arg Cys Lys Val Thr Val Ala Pro Met Val Pro Pro
290 295 300
Ile Val Leu Ala Ile Ala Lys Ser Ser Glu Thr Glu Lys Tyr Asp Leu
305 310 315 320
Ser Ser Ile Arg Val Val Lys Ser Gly Ala Ala Pro Leu Gly Lys Glu
325 330 335
Leu Glu Asp Ala Val Asn Ala Lys Phe Pro Asn Ala Lys Leu Gly Gln
340 345 350
Gly Tyr Gly Met Thr Glu Ala Gly Pro Val Leu Ala Met Ser Leu Gly
355 360 365
Phe Ala Lys Glu Pro Phe Pro Val Lys Ser Gly Ala Cys Gly Thr Val
370 375 380
Val Arg Asn Ala Glu Met Lys Ile Val Asp Pro Asp Thr Gly Asp Ser
385 390 395 400
Leu Ser Arg Asn Gln Pro Gly Glu Ile Cys Ile Arg Gly His Gln Ile
405 410 415
Met Lys Gly Tyr Leu Asn Asn Pro Ala Ala Thr Ala Glu Thr Ile Asp
420 425 430
Lys Asp Gly Trp Leu His Thr Gly Asp Ile Gly Leu Ile Asp Asp Asp
435 440 445
Asp Glu Leu Phe Ile Val Asp Arg Leu Lys Glu Leu Ile Lys Tyr Lys
450 455 460
Gly Phe Gln Val Ala Pro Ala Glu Leu Glu Ala Leu Leu Ile Gly His
465 470 475 480
Pro Asp Ile Thr Asp Val Ala Val Val Ala Met Lys Glu Glu Ala Ala
485 490 495
Gly Glu Val Pro Val Ala Phe Val Val Lys Ser Lys Asp Ser Glu Leu
500 505 510
Ser Glu Asp Asp Val Lys Gln Phe Val Ser Lys Gln Val Val Phe Tyr
515 520 525
Lys Arg Ile Asn Lys Val Phe Phe Thr Glu Ser Ile Pro Lys Ala Pro
530 535 540
Ser Gly Lys Ile Leu Arg Lys Asp Leu Arg Ala Lys Leu Ala Asn Gly
545 550 555 560
Leu
<210> 129
<211> 561
<212> PRT
<213> 拟南芥(Arabidopsis thaliana)
<400> 129
Met Ile Thr Ala Ala Leu His Glu Pro Gln Ile His Lys Pro Thr Asp
1 5 10 15
Thr Ser Val Val Ser Asp Asp Val Leu Pro His Ser Pro Pro Thr Pro
20 25 30
Arg Ile Phe Arg Ser Lys Leu Pro Asp Ile Asp Ile Pro Asn His Leu
35 40 45
Pro Leu His Thr Tyr Cys Phe Glu Lys Leu Ser Ser Val Ser Asp Lys
50 55 60
Pro Cys Leu Ile Val Gly Ser Thr Gly Lys Ser Tyr Thr Tyr Gly Glu
65 70 75 80
Thr His Leu Ile Cys Arg Arg Val Ala Ser Gly Leu Tyr Lys Leu Gly
85 90 95
Ile Arg Lys Gly Asp Val Ile Met Ile Leu Leu Gln Asn Ser Ala Glu
100 105 110
Phe Val Phe Ser Phe Met Gly Ala Ser Met Ile Gly Ala Val Ser Thr
115 120 125
Thr Ala Asn Pro Phe Tyr Thr Ser Gln Glu Leu Tyr Lys Gln Leu Lys
130 135 140
Ser Ser Gly Ala Lys Leu Ile Ile Thr His Ser Gln Tyr Val Asp Lys
145 150 155 160
Leu Lys Asn Leu Gly Glu Asn Leu Thr Leu Ile Thr Thr Asp Glu Pro
165 170 175
Thr Pro Glu Asn Cys Leu Pro Phe Ser Thr Leu Ile Thr Asp Asp Glu
180 185 190
Thr Asn Pro Phe Gln Glu Thr Val Asp Ile Gly Gly Asp Asp Ala Ala
195 200 205
Ala Leu Pro Phe Ser Ser Gly Thr Thr Gly Leu Pro Lys Gly Val Val
210 215 220
Leu Thr His Lys Ser Leu Ile Thr Ser Val Ala Gln Gln Val Asp Gly
225 230 235 240
Asp Asn Pro Asn Leu Tyr Leu Lys Ser Asn Asp Val Ile Leu Cys Val
245 250 255
Leu Pro Leu Phe His Ile Tyr Ser Leu Asn Ser Val Leu Leu Asn Ser
260 265 270
Leu Arg Ser Gly Ala Thr Val Leu Leu Met His Lys Phe Glu Ile Gly
275 280 285
Ala Leu Leu Asp Leu Ile Gln Arg His Arg Val Thr Ile Ala Ala Leu
290 295 300
Val Pro Pro Leu Val Ile Ala Leu Ala Lys Asn Pro Thr Val Asn Ser
305 310 315 320
Tyr Asp Leu Ser Ser Val Arg Phe Val Leu Ser Gly Ala Ala Pro Leu
325 330 335
Gly Lys Glu Leu Gln Asp Ser Leu Arg Arg Arg Leu Pro Gln Ala Ile
340 345 350
Leu Gly Gln Gly Tyr Gly Met Thr Glu Ala Gly Pro Val Leu Ser Met
355 360 365
Ser Leu Gly Phe Ala Lys Glu Pro Ile Pro Thr Lys Ser Gly Ser Cys
370 375 380
Gly Thr Val Val Arg Asn Ala Glu Leu Lys Val Val His Leu Glu Thr
385 390 395 400
Arg Leu Ser Leu Gly Tyr Asn Gln Pro Gly Glu Ile Cys Ile Arg Gly
405 410 415
Gln Gln Ile Met Lys Glu Tyr Leu Asn Asp Pro Glu Ala Thr Ser Ala
420 425 430
Thr Ile Asp Glu Glu Gly Trp Leu His Thr Gly Asp Ile Gly Tyr Val
435 440 445
Asp Glu Asp Asp Glu Ile Phe Ile Val Asp Arg Leu Lys Glu Val Ile
450 455 460
Lys Phe Lys Gly Phe Gln Val Pro Pro Ala Glu Leu Glu Ser Leu Leu
465 470 475 480
Ile Asn His His Ser Ile Ala Asp Ala Ala Val Val Pro Gln Asn Asp
485 490 495
Glu Val Ala Gly Glu Val Pro Val Ala Phe Val Val Arg Ser Asn Gly
500 505 510
Asn Asp Ile Thr Glu Glu Asp Val Lys Glu Tyr Val Ala Lys Gln Val
515 520 525
Val Phe Tyr Lys Arg Leu His Lys Val Phe Phe Val Ala Ser Ile Pro
530 535 540
Lys Ser Pro Ser Gly Lys Ile Leu Arg Lys Asp Leu Lys Ala Lys Leu
545 550 555 560
Cys
<210> 130
<211> 570
<212> PRT
<213> 拟南芥(Arabidopsis thaliana)
<400> 130
Met Val Leu Gln Gln Gln Thr His Phe Leu Thr Lys Lys Ile Asp Gln
1 5 10 15
Glu Asp Glu Glu Glu Glu Pro Ser His Asp Phe Ile Phe Arg Ser Lys
20 25 30
Leu Pro Asp Ile Phe Ile Pro Asn His Leu Pro Leu Thr Asp Tyr Val
35 40 45
Phe Gln Arg Phe Ser Gly Asp Gly Asp Gly Asp Ser Ser Thr Thr Cys
50 55 60
Ile Ile Asp Gly Ala Thr Gly Arg Ile Leu Thr Tyr Ala Asp Val Gln
65 70 75 80
Thr Asn Met Arg Arg Ile Ala Ala Gly Ile His Arg Leu Gly Ile Arg
85 90 95
His Gly Asp Val Val Met Leu Leu Leu Pro Asn Ser Pro Glu Phe Ala
100 105 110
Leu Ser Phe Leu Ala Val Ala Tyr Leu Gly Ala Val Ser Thr Thr Ala
115 120 125
Asn Pro Phe Tyr Thr Gln Pro Glu Ile Ala Lys Gln Ala Lys Ala Ser
130 135 140
Ala Ala Lys Met Ile Ile Thr Lys Lys Cys Leu Val Asp Lys Leu Thr
145 150 155 160
Asn Leu Lys Asn Asp Gly Val Leu Ile Val Cys Leu Asp Asp Asp Gly
165 170 175
Asp Asn Gly Val Val Ser Ser Ser Asp Asp Gly Cys Val Ser Phe Thr
180 185 190
Glu Leu Thr Gln Ala Asp Glu Thr Glu Leu Leu Lys Pro Lys Ile Ser
195 200 205
Pro Glu Asp Thr Val Ala Met Pro Tyr Ser Ser Gly Thr Thr Gly Leu
210 215 220
Pro Lys Gly Val Met Ile Thr His Lys Gly Leu Val Thr Ser Ile Ala
225 230 235 240
Gln Lys Val Asp Gly Glu Asn Pro Asn Leu Asn Phe Thr Ala Asn Asp
245 250 255
Val Ile Leu Cys Phe Leu Pro Met Phe His Ile Tyr Ala Leu Asp Ala
260 265 270
Leu Met Leu Ser Ala Met Arg Thr Gly Ala Ala Leu Leu Ile Val Pro
275 280 285
Arg Phe Glu Leu Asn Leu Val Met Glu Leu Ile Gln Arg Tyr Lys Val
290 295 300
Thr Val Val Pro Val Ala Pro Pro Val Val Leu Ala Phe Ile Lys Ser
305 310 315 320
Pro Glu Thr Glu Arg Tyr Asp Leu Ser Ser Val Arg Ile Met Leu Ser
325 330 335
Gly Ala Ala Thr Leu Lys Lys Glu Leu Glu Asp Ala Val Arg Leu Lys
340 345 350
Phe Pro Asn Ala Ile Phe Gly Gln Gly Tyr Gly Met Thr Glu Ser Gly
355 360 365
Thr Val Ala Lys Ser Leu Ala Phe Ala Lys Asn Pro Phe Lys Thr Lys
370 375 380
Ser Gly Ala Cys Gly Thr Val Ile Arg Asn Ala Glu Met Lys Val Val
385 390 395 400
Asp Thr Glu Thr Gly Ile Ser Leu Pro Arg Asn Lys Ser Gly Glu Ile
405 410 415
Cys Val Arg Gly His Gln Leu Met Lys Gly Tyr Leu Asn Asp Pro Glu
420 425 430
Ala Thr Ala Arg Thr Ile Asp Lys Asp Gly Trp Leu His Thr Gly Asp
435 440 445
Ile Gly Phe Val Asp Asp Asp Asp Glu Ile Phe Ile Val Asp Arg Leu
450 455 460
Lys Glu Leu Ile Lys Phe Lys Gly Tyr Gln Val Ala Pro Ala Glu Leu
465 470 475 480
Glu Ala Leu Leu Ile Ser His Pro Ser Ile Asp Asp Ala Ala Val Val
485 490 495
Ala Met Lys Asp Glu Val Ala Asp Glu Val Pro Val Ala Phe Val Ala
500 505 510
Arg Ser Gln Gly Ser Gln Leu Thr Glu Asp Asp Val Lys Ser Tyr Val
515 520 525
Asn Lys Gln Val Val His Tyr Lys Arg Ile Lys Met Val Phe Phe Ile
530 535 540
Glu Val Ile Pro Lys Ala Val Ser Gly Lys Ile Leu Arg Lys Asp Leu
545 550 555 560
Arg Ala Lys Leu Glu Thr Met Cys Ser Lys
565 570
<210> 131
<211> 522
<212> PRT
<213> 天蓝色链霉菌(Streptomyces coelicolor)
<400> 131
Met Phe Arg Ser Glu Tyr Ala Asp Val Pro Pro Val Asp Leu Pro Ile
1 5 10 15
His Asp Ala Val Leu Gly Gly Ala Ala Ala Phe Gly Ser Thr Pro Ala
20 25 30
Leu Ile Asp Gly Thr Asp Gly Thr Thr Leu Thr Tyr Glu Gln Val Asp
35 40 45
Arg Phe His Arg Arg Val Ala Ala Ala Leu Ala Glu Thr Gly Val Arg
50 55 60
Lys Gly Asp Val Leu Ala Leu His Ser Pro Asn Thr Val Ala Phe Pro
65 70 75 80
Leu Ala Phe Tyr Ala Ala Thr Arg Ala Gly Ala Ser Val Thr Thr Val
85 90 95
His Pro Leu Ala Thr Ala Glu Glu Phe Ala Lys Gln Leu Lys Asp Ser
100 105 110
Ala Ala Arg Trp Ile Val Thr Val Ser Pro Leu Leu Ser Thr Ala Arg
115 120 125
Arg Ala Ala Glu Leu Ala Gly Gly Val Gln Glu Ile Leu Val Cys Asp
130 135 140
Ser Ala Pro Gly His Arg Ser Leu Val Asp Met Leu Ala Ser Thr Ala
145 150 155 160
Pro Glu Pro Ser Val Ala Ile Asp Pro Ala Glu Asp Val Ala Ala Leu
165 170 175
Pro Tyr Ser Ser Gly Thr Thr Gly Thr Pro Lys Gly Val Met Leu Thr
180 185 190
His Arg Gln Ile Ala Thr Asn Leu Ala Gln Leu Glu Pro Ser Met Pro
195 200 205
Ser Ala Pro Gly Asp Arg Val Leu Ala Val Leu Pro Phe Phe His Ile
210 215 220
Tyr Gly Leu Thr Ala Leu Met Asn Ala Pro Leu Arg Leu Gly Ala Thr
225 230 235 240
Val Val Val Leu Pro Arg Phe Asp Leu Glu Gln Phe Leu Ala Ala Ile
245 250 255
Gln Asn His Arg Ile Thr Ser Leu Tyr Val Ala Pro Pro Ile Val Leu
260 265 270
Ala Leu Ala Lys His Pro Leu Val Ala Asp Tyr Asp Leu Ser Ser Leu
275 280 285
Arg Tyr Ile Val Ser Ala Ala Ala Pro Leu Asp Ala Arg Leu Ala Ala
290 295 300
Ala Cys Ser Gln Arg Leu Gly Leu Pro Pro Val Gly Gln Ala Tyr Gly
305 310 315 320
Met Thr Glu Leu Ser Pro Gly Thr His Val Val Pro Leu Asp Ala Met
325 330 335
Ala Asp Ala Pro Pro Gly Thr Val Gly Arg Leu Ile Ala Gly Thr Glu
340 345 350
Met Arg Ile Val Ser Leu Thr Asp Pro Gly Thr Asp Leu Pro Ala Gly
355 360 365
Glu Ser Gly Glu Ile Leu Ile Arg Gly Pro Gln Ile Met Lys Gly Tyr
370 375 380
Leu Gly Arg Pro Asp Ala Thr Ala Ala Met Ile Asp Glu Glu Gly Trp
385 390 395 400
Leu His Thr Gly Asp Val Gly His Val Asp Ala Asp Gly Trp Leu Phe
405 410 415
Val Val Asp Arg Val Lys Glu Leu Ile Lys Tyr Lys Gly Phe Gln Val
420 425 430
Ala Pro Ala Glu Leu Glu Ala His Leu Leu Thr His Pro Gly Val Ala
435 440 445
Asp Ala Ala Val Val Gly Ala Tyr Asp Asp Asp Gly Asn Glu Val Pro
450 455 460
His Ala Phe Val Val Arg Gln Pro Ala Ala Pro Gly Leu Ala Glu Ser
465 470 475 480
Glu Ile Met Met Tyr Val Ala Glu Arg Val Ala Pro Tyr Lys Arg Val
485 490 495
Arg Arg Val Thr Phe Val Asp Ala Val Pro Arg Ala Ala Ser Gly Lys
500 505 510
Ile Leu Arg Arg Gln Leu Arg Glu Pro Arg
515 520
<210> 132
<211> 392
<212> PRT
<213> 葡萄(Vitis vinifera)
<400> 132
Met Ala Ser Val Glu Glu Phe Arg Asn Ala Gln Arg Ala Lys Gly Pro
1 5 10 15
Ala Thr Ile Leu Ala Ile Gly Thr Ala Thr Pro Asp His Cys Val Tyr
20 25 30
Gln Ser Asp Tyr Ala Asp Phe Tyr Phe Arg Val Thr Lys Ser Glu His
35 40 45
Met Thr Ala Leu Lys Lys Lys Phe Asn Arg Ile Cys Asp Lys Ser Met
50 55 60
Ile Lys Lys Arg Tyr Ile His Leu Thr Glu Glu Met Leu Glu Glu His
65 70 75 80
Pro Asn Ile Gly Ala Tyr Met Ala Pro Ser Leu Asn Ile Arg Gln Glu
85 90 95
Ile Ile Thr Ala Glu Val Pro Lys Leu Gly Lys Glu Ala Ala Leu Lys
100 105 110
Ala Leu Lys Glu Trp Gly Gln Pro Lys Ser Lys Ile Thr His Leu Val
115 120 125
Phe Cys Thr Thr Ser Gly Val Glu Met Pro Gly Ala Asp Tyr Lys Leu
130 135 140
Ala Asn Leu Leu Gly Leu Glu Pro Ser Val Arg Arg Val Met Leu Tyr
145 150 155 160
His Gln Gly Cys Tyr Ala Gly Gly Thr Val Leu Arg Thr Ala Lys Asp
165 170 175
Leu Ala Glu Asn Asn Ala Gly Ala Arg Val Leu Val Val Cys Ser Glu
180 185 190
Ile Thr Val Val Thr Phe Arg Gly Pro Ser Glu Asp Ala Leu Asp Ser
195 200 205
Leu Val Gly Gln Ala Leu Phe Gly Asp Gly Ser Ala Ala Val Ile Val
210 215 220
Gly Ser Asp Pro Asp Ile Ser Ile Glu Arg Pro Leu Phe Gln Leu Val
225 230 235 240
Ser Ala Ala Gln Thr Phe Ile Pro Asn Ser Ala Gly Ala Ile Ala Gly
245 250 255
Asn Leu Arg Glu Val Gly Leu Thr Phe His Leu Trp Pro Asn Val Pro
260 265 270
Thr Leu Ile Ser Glu Asn Ile Glu Lys Cys Leu Thr Gln Ala Phe Asp
275 280 285
Pro Leu Gly Ile Ser Asp Trp Asn Ser Leu Phe Trp Ile Ala His Pro
290 295 300
Gly Gly Pro Ala Ile Leu Asp Ala Val Glu Ala Lys Leu Asn Leu Asp
305 310 315 320
Lys Lys Lys Leu Glu Ala Thr Arg His Val Leu Ser Glu Tyr Gly Asn
325 330 335
Met Ser Ser Ala Cys Val Leu Phe Ile Leu Asp Glu Met Arg Lys Lys
340 345 350
Ser Leu Lys Gly Glu Arg Ala Thr Thr Gly Glu Gly Leu Asp Trp Gly
355 360 365
Val Leu Phe Gly Phe Gly Pro Gly Leu Thr Ile Glu Thr Val Val Leu
370 375 380
His Ser Ile Pro Met Val Thr Asn
385 390
<210> 133
<211> 389
<212> PRT
<213> 矮牵牛(Petunia x hybrida)
<400> 133
Met Val Thr Val Glu Glu Tyr Arg Lys Ala Gln Arg Ala Glu Gly Pro
1 5 10 15
Ala Thr Val Met Ala Ile Gly Thr Ala Thr Pro Thr Asn Cys Val Asp
20 25 30
Gln Ser Thr Tyr Pro Asp Tyr Tyr Phe Arg Ile Thr Asn Ser Glu His
35 40 45
Lys Thr Asp Leu Lys Glu Lys Phe Lys Arg Met Cys Glu Lys Ser Met
50 55 60
Ile Lys Lys Arg Tyr Met His Leu Thr Glu Glu Ile Leu Lys Glu Asn
65 70 75 80
Pro Ser Met Cys Glu Tyr Met Ala Pro Ser Leu Asp Ala Arg Gln Asp
85 90 95
Ile Val Val Val Glu Val Pro Lys Leu Gly Lys Glu Ala Ala Gln Lys
100 105 110
Ala Ile Lys Glu Trp Gly Gln Pro Lys Ser Lys Ile Thr His Leu Val
115 120 125
Phe Cys Thr Thr Ser Gly Val Asp Met Pro Gly Cys Asp Tyr Gln Leu
130 135 140
Thr Lys Leu Leu Gly Leu Arg Pro Ser Val Lys Arg Leu Met Met Tyr
145 150 155 160
Gln Gln Gly Cys Phe Ala Gly Gly Thr Val Leu Arg Leu Ala Lys Asp
165 170 175
Leu Ala Glu Asn Asn Lys Gly Ala Arg Val Leu Val Val Cys Ser Glu
180 185 190
Ile Thr Ala Val Thr Phe Arg Gly Pro Asn Asp Thr His Leu Asp Ser
195 200 205
Leu Val Gly Gln Ala Leu Phe Gly Asp Gly Ala Gly Ala Ile Ile Ile
210 215 220
Gly Ser Asp Pro Ile Pro Gly Val Glu Arg Pro Leu Phe Glu Leu Val
225 230 235 240
Ser Ala Ala Gln Thr Leu Leu Pro Asp Ser His Gly Ala Ile Asp Gly
245 250 255
His Leu Arg Glu Val Gly Leu Thr Phe His Leu Leu Lys Asp Val Pro
260 265 270
Gly Leu Ile Ser Lys Asn Ile Glu Lys Ser Leu Glu Glu Ala Phe Arg
275 280 285
Pro Leu Ser Ile Ser Asp Trp Asn Ser Leu Phe Trp Ile Ala His Pro
290 295 300
Gly Gly Pro Ala Ile Leu Asp Gln Val Glu Ile Lys Leu Gly Leu Lys
305 310 315 320
Pro Glu Lys Leu Lys Ala Thr Arg Asn Val Leu Ser Asn Tyr Gly Asn
325 330 335
Met Ser Ser Ala Cys Val Leu Phe Ile Leu Asp Glu Met Arg Lys Ala
340 345 350
Ser Ala Lys Glu Gly Leu Gly Thr Thr Gly Glu Gly Leu Glu Trp Gly
355 360 365
Val Leu Phe Gly Phe Gly Pro Gly Leu Thr Val Glu Thr Val Val Leu
370 375 380
His Ser Val Ala Thr
385
<210> 134
<211> 391
<212> PRT
<213> 拟南芥(Arabidopsis thaliana)
<400> 134
Met Ala Asp Val Leu Gln Glu Ile Arg Asn Ser Gln Lys Ala Ser Gly
1 5 10 15
Pro Ala Thr Val Leu Ala Ile Gly Thr Ala His Pro Pro Thr Cys Tyr
20 25 30
Pro Gln Ala Asp Tyr Pro Asp Phe Tyr Phe Arg Val Cys Lys Ser Glu
35 40 45
His Met Thr Lys Leu Lys Lys Lys Met Gln Phe Ile Cys Asp Arg Ser
50 55 60
Gly Ile Arg Gln Arg Phe Met Phe His Thr Glu Glu Asn Leu Gly Lys
65 70 75 80
Asn Pro Gly Met Cys Thr Phe Asp Gly Pro Ser Leu Asn Ala Arg Gln
85 90 95
Asp Met Leu Ile Met Glu Val Pro Lys Leu Gly Ala Glu Ala Ala Glu
100 105 110
Lys Ala Ile Lys Glu Trp Gly Gln Asp Lys Ser Arg Ile Thr His Leu
115 120 125
Ile Phe Cys Thr Thr Thr Ser Asn Asp Met Pro Gly Ala Asp Tyr Gln
130 135 140
Phe Ala Thr Leu Phe Gly Leu Asn Pro Gly Val Ser Arg Thr Met Val
145 150 155 160
Tyr Gln Gln Gly Cys Phe Ala Gly Gly Thr Val Leu Arg Leu Val Lys
165 170 175
Asp Ile Ala Glu Asn Asn Lys Gly Ala Arg Val Leu Val Val Cys Ser
180 185 190
Glu Ile Val Ala Phe Ala Phe Arg Gly Pro His Glu Asp His Ile Asp
195 200 205
Ser Leu Ile Gly Gln Leu Leu Phe Gly Asp Gly Ala Ala Ala Leu Val
210 215 220
Val Gly Thr Asp Ile Asp Glu Ser Val Glu Arg Pro Ile Phe Gln Ile
225 230 235 240
Met Ser Ala Thr Gln Ala Thr Ile Pro Asn Ser Leu His Thr Met Ala
245 250 255
Leu His Leu Thr Glu Ala Gly Leu Thr Phe His Leu Ser Lys Glu Val
260 265 270
Pro Lys Val Val Ser Asp Asn Met Glu Glu Leu Met Leu Glu Ala Phe
275 280 285
Lys Pro Leu Gly Ile Thr Asp Trp Asn Ser Ile Phe Trp Gln Val His
290 295 300
Pro Gly Gly Arg Ala Ile Leu Asp Lys Ile Glu Glu Lys Leu Glu Leu
305 310 315 320
Thr Lys Asp Lys Met Arg Asp Ser Arg Tyr Ile Leu Ser Glu Tyr Gly
325 330 335
Asn Leu Thr Ser Ala Cys Val Leu Phe Val Met Asp Glu Met Arg Lys
340 345 350
Arg Ser Phe Arg Glu Gly Lys Gln Thr Thr Gly Asp Gly Tyr Glu Trp
355 360 365
Gly Val Ala Ile Gly Leu Gly Pro Gly Leu Thr Val Glu Thr Val Val
370 375 380
Leu Arg Ser Val Pro Ile Pro
385 390
<210> 135
<211> 43
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 135
ctttaagaag gagatataca tatgagttca ctctccaact ctc 43
<210> 136
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 136
cttgtcgacg gagctcgaat tcattacatg agaggcaggc tgtg 44
<210> 137
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 137
ctttaagaag gagatataca tatgagttca ctctccaacg cttc 44
<210> 138
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 138
cttgtcgacg gagctcgaat tcattacatg agaggcaggc tgtg 44
<210> 139
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 139
ctttaagaag gagatataca tatgggttca ctctccgact ctac 44
<210> 140
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 140
cttgtcgacg gagctcgaat tcattacaca ggaggcaggc tatg 44
<210> 141
<211> 42
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 141
ctttaagaag gagatataca tatgggttca ctctccaact ac 42
<210> 142
<211> 42
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 142
cttgtcgacg gagctcgaat tcattacaca gggggcaggc tg 42
<210> 143
<211> 39
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 143
ctttaagaag gagatataca tatgggttcg atcgccgag 39
<210> 144
<211> 42
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 144
cttgtcgacg gagctcgaat tcattagaca ggaggcaggc tg 42
<210> 145
<211> 32
<212> DNA
<213> 人工序列
<220>
<223> 引物.
<400> 145
gctcaccatc attgggctcc gtggccccaa tg 32
<210> 146
<211> 32
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 146
cattggggcc acggagccca atgatggtga gc 32
<210> 147
<211> 40
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 147
gacggatggc cttttactag tgcctggggt gcctaatgag 40
<210> 148
<211> 45
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 148
cgatgattaa ttgtcaactg ctacgcctga ataagtgata ataag 45
<210> 149
<211> 22
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 149
gttgacaatt aatcatcggc tc 22
<210> 150
<211> 27
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 150
actagtaaaa ggccatccgt caggatg 27
<210> 151
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 151
gttgacaatt aatcatcggc 20
<210> 152
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 152
cgtttcactt ctgagttcgg 20
<210> 153
<211> 46
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 153
caatttcaca caggaaacag aattcgtgtc agtcgagact aggaag 46
<210> 154
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 154
ctctagagga tccccgggta ccattacttg atctcgagga gaac 44
<210> 155
<211> 46
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 155
caatttcaca caggaaacag aattcatgac catttcctca cctttg 46
<210> 156
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 156
ctctagagga tccccgggta ccattacagt ggcatgttgc cgtg 44
<210> 157
<211> 43
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 157
gagtcgacct gcaggcatgc attgacaatt aatcatcggc tcg 43
<210> 158
<211> 25
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 158
ctcatccgcc aaaacagcca agctt 25
<210> 159
<211> 47
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 159
caatttcaca caggaaacag aattcatgtc agaacgtttc ccaaatg 47
<210> 160
<211> 45
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 160
ctctagagga tccccgggta ccattacatc accagacggc gaatg 45
<210> 161
<211> 47
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 161
caatttcaca caggaaacag aattcatgag tactgaaatc aaaactc 47
<210> 162
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 162
ctctagagga tccccgggta ccattacttc ttcttcgctt tcgg 44
<210> 163
<211> 43
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 163
gtacccgggg atcctctaga gttgacaatt aatcatcggc tcg 43
<210> 164
<211> 25
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 164
caagcttgca tgcctgcagg tcgac 25
<210> 165
<211> 49
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 165
caatttcaca caggaaacag aattcatgac tatcaaagta ggtatcaac 49
<210> 166
<211> 45
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 166
ctctagagga tccccgggta ccattatttg gagatgtgag cgatc 45
<210> 167
<211> 50
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 167
caatttcaca caggaaacag aattcatgtc tgtaattaag atgaccgatc 50
<210> 168
<211> 45
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 168
ctctagagga tccccgggta ccattacttc ttagcgcgct cttcg 45
<210> 169
<211> 46
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 169
caatttcaca caggaaacag aattcatgag ccaaattcac aaacac 46
<210> 170
<211> 43
<212> DNA
<213> 人工序列
<220>
<223> 引物
<400> 170
ctctagagga tccccgggta ccattacgat ggcatcgcga tag 43
<210> 171
<211> 491
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 171
Met Ala Val Thr Gln Thr Ala Gln Ala Cys Asp Leu Val Ile Phe Gly
1 5 10 15
Ala Lys Gly Asp Leu Ala Arg Arg Lys Leu Leu Pro Ser Leu Tyr Gln
20 25 30
Leu Glu Lys Ala Gly Gln Leu Asn Pro Asp Thr Arg Ile Ile Gly Val
35 40 45
Gly Arg Ala Asp Trp Asp Lys Ala Ala Tyr Thr Lys Val Val Arg Glu
50 55 60
Ala Leu Glu Thr Phe Met Lys Glu Thr Ile Asp Glu Gly Leu Trp Asp
65 70 75 80
Thr Leu Ser Ala Arg Leu Asp Phe Cys Asn Leu Asp Val Asn Asp Thr
85 90 95
Ala Ala Phe Ser Arg Leu Gly Ala Met Leu Asp Gln Lys Asn Arg Ile
100 105 110
Thr Ile Asn Tyr Phe Ala Met Pro Pro Ser Thr Phe Gly Ala Ile Cys
115 120 125
Lys Gly Leu Gly Glu Ala Lys Leu Asn Ala Lys Pro Ala Arg Val Val
130 135 140
Met Glu Lys Pro Leu Gly Thr Ser Leu Ala Thr Ser Gln Glu Ile Asn
145 150 155 160
Asp Gln Val Gly Glu Tyr Phe Glu Glu Cys Gln Val Tyr Arg Ile Asp
165 170 175
His Tyr Leu Gly Lys Glu Thr Val Leu Asn Leu Leu Ala Leu Arg Phe
180 185 190
Ala Asn Ser Leu Phe Val Asn Asn Trp Asp Asn Arg Thr Ile Asp His
195 200 205
Val Glu Ile Thr Val Ala Glu Glu Val Gly Ile Glu Gly Arg Trp Gly
210 215 220
Tyr Phe Asp Lys Ala Gly Gln Met Arg Asp Met Ile Gln Asn His Leu
225 230 235 240
Leu Gln Ile Leu Cys Met Ile Ala Met Ser Pro Pro Ser Asp Leu Ser
245 250 255
Ala Asp Ser Ile Arg Asp Glu Lys Val Lys Val Leu Lys Ser Leu Arg
260 265 270
Arg Ile Asp Arg Ser Asn Val Arg Glu Lys Thr Val Arg Gly Gln Tyr
275 280 285
Thr Ala Gly Phe Ala Gln Gly Lys Lys Val Pro Gly Tyr Leu Glu Glu
290 295 300
Glu Gly Ala Asn Lys Ser Ser Asn Thr Glu Thr Phe Val Ala Ile Arg
305 310 315 320
Val Asp Ile Asp Asn Trp Arg Trp Ala Gly Val Pro Phe Tyr Leu Arg
325 330 335
Thr Gly Lys Arg Leu Pro Thr Lys Cys Ser Glu Val Val Val Tyr Phe
340 345 350
Lys Thr Pro Glu Leu Asn Leu Phe Lys Glu Ser Trp Gln Asp Leu Pro
355 360 365
Gln Asn Lys Leu Thr Ile Arg Leu Gln Pro Asp Glu Gly Val Asp Ile
370 375 380
Gln Val Leu Asn Lys Val Pro Gly Leu Asp His Lys His Asn Leu Gln
385 390 395 400
Ile Thr Lys Leu Asp Leu Ser Tyr Ser Glu Thr Phe Asn Gln Thr His
405 410 415
Leu Ala Asp Ala Tyr Glu Arg Leu Leu Leu Glu Thr Met Arg Gly Ile
420 425 430
Gln Ala Leu Phe Val Arg Arg Asp Glu Val Glu Glu Ala Trp Lys Trp
435 440 445
Val Asp Ser Ile Thr Glu Ala Trp Ala Met Asp Asn Asp Ala Pro Lys
450 455 460
Pro Tyr Gln Ala Gly Thr Trp Gly Pro Val Ala Ser Val Ala Met Ile
465 470 475 480
Thr Arg Asp Gly Arg Ser Trp Asn Glu Phe Glu
485 490
<210> 172
<211> 312
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 172
Met Lys Val Ala Val Leu Gly Ala Ala Gly Gly Ile Gly Gln Ala Leu
1 5 10 15
Ala Leu Leu Leu Lys Thr Gln Leu Pro Ser Gly Ser Glu Leu Ser Leu
20 25 30
Tyr Asp Ile Ala Pro Val Thr Pro Gly Val Ala Val Asp Leu Ser His
35 40 45
Ile Pro Thr Ala Val Lys Ile Lys Gly Phe Ser Gly Glu Asp Ala Thr
50 55 60
Pro Ala Leu Glu Gly Ala Asp Val Val Leu Ile Ser Ala Gly Val Ala
65 70 75 80
Arg Lys Pro Gly Met Asp Arg Ser Asp Leu Phe Asn Val Asn Ala Gly
85 90 95
Ile Val Lys Asn Leu Val Gln Gln Val Ala Lys Thr Cys Pro Lys Ala
100 105 110
Cys Ile Gly Ile Ile Thr Asn Pro Val Asn Thr Thr Val Ala Ile Ala
115 120 125
Ala Glu Val Leu Lys Lys Ala Gly Val Tyr Asp Lys Asn Lys Leu Phe
130 135 140
Gly Val Thr Thr Leu Asp Ile Ile Arg Ser Asn Thr Phe Val Ala Glu
145 150 155 160
Leu Lys Gly Lys Gln Pro Gly Glu Val Glu Val Pro Val Ile Gly Gly
165 170 175
His Ser Gly Val Thr Ile Leu Pro Leu Leu Ser Gln Val Pro Gly Val
180 185 190
Ser Phe Thr Glu Gln Glu Val Ala Asp Leu Thr Lys Arg Ile Gln Asn
195 200 205
Ala Gly Thr Glu Val Val Glu Ala Lys Ala Gly Gly Gly Ser Ala Thr
210 215 220
Leu Ser Met Gly Gln Ala Ala Ala Arg Phe Gly Leu Ser Leu Val Arg
225 230 235 240
Ala Leu Gln Gly Glu Gln Gly Val Val Glu Cys Ala Tyr Val Glu Gly
245 250 255
Asp Gly Gln Tyr Ala Arg Phe Phe Ser Gln Pro Leu Leu Leu Gly Lys
260 265 270
Asn Gly Val Glu Glu Arg Lys Ser Ile Gly Thr Leu Ser Ala Phe Glu
275 280 285
Gln Asn Ala Leu Glu Gly Met Leu Asp Thr Leu Lys Lys Asp Ile Ala
290 295 300
Leu Gly Glu Glu Phe Val Asn Lys
305 310
<210> 173
<211> 410
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 173
Met Ala Lys Val Ser Leu Glu Lys Asp Lys Ile Lys Phe Leu Leu Val
1 5 10 15
Glu Gly Val His Gln Lys Ala Leu Glu Ser Leu Arg Ala Ala Gly Tyr
20 25 30
Thr Asn Ile Glu Phe His Lys Gly Ala Leu Asp Asp Glu Gln Leu Lys
35 40 45
Glu Ser Ile Arg Asp Ala His Phe Ile Gly Leu Arg Ser Arg Thr His
50 55 60
Leu Thr Glu Asp Val Ile Asn Ala Ala Glu Lys Leu Val Ala Ile Gly
65 70 75 80
Cys Phe Cys Ile Gly Thr Asn Gln Val Asp Leu Asp Ala Ala Ala Lys
85 90 95
Arg Gly Ile Pro Val Phe Asn Ala Pro Phe Ser Asn Thr Arg Ser Val
100 105 110
Ala Glu Leu Val Ile Gly Glu Leu Leu Leu Leu Leu Arg Gly Val Pro
115 120 125
Glu Ala Asn Ala Lys Ala His Arg Gly Val Trp Asn Lys Leu Ala Ala
130 135 140
Gly Ser Phe Glu Ala Arg Gly Lys Lys Leu Gly Ile Ile Gly Tyr Gly
145 150 155 160
His Ile Gly Thr Gln Leu Gly Ile Leu Ala Glu Ser Leu Gly Met Tyr
165 170 175
Val Tyr Phe Tyr Asp Ile Glu Asn Lys Leu Pro Leu Gly Asn Ala Thr
180 185 190
Gln Val Gln His Leu Ser Asp Leu Leu Asn Met Ser Asp Val Val Ser
195 200 205
Leu His Val Pro Glu Asn Pro Ser Thr Lys Asn Met Met Gly Ala Lys
210 215 220
Glu Ile Ser Leu Met Lys Pro Gly Ser Leu Leu Ile Asn Ala Ser Arg
225 230 235 240
Gly Thr Val Val Asp Ile Pro Ala Leu Cys Asp Ala Leu Ala Ser Lys
245 250 255
His Leu Ala Gly Ala Ala Ile Asp Val Phe Pro Thr Glu Pro Ala Thr
260 265 270
Asn Ser Asp Pro Phe Thr Ser Pro Leu Cys Glu Phe Asp Asn Val Leu
275 280 285
Leu Thr Pro His Ile Gly Gly Ser Thr Gln Glu Ala Gln Glu Asn Ile
290 295 300
Gly Leu Glu Val Ala Gly Lys Leu Ile Lys Tyr Ser Asp Asn Gly Ser
305 310 315 320
Thr Leu Ser Ala Val Asn Phe Pro Glu Val Ser Leu Pro Leu His Gly
325 330 335
Gly Arg Arg Leu Met His Ile His Glu Asn Arg Pro Gly Val Leu Thr
340 345 350
Ala Leu Asn Lys Ile Phe Ala Glu Gln Gly Val Asn Ile Ala Ala Gln
355 360 365
Tyr Leu Gln Thr Ser Ala Gln Met Gly Tyr Val Val Ile Asp Ile Glu
370 375 380
Ala Asp Glu Asp Val Ala Glu Lys Ala Leu Gln Ala Met Lys Ala Ile
385 390 395 400
Pro Gly Thr Ile Arg Ala Arg Leu Leu Tyr
405 410
<210> 174
<211> 331
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 174
Met Thr Ile Lys Val Gly Ile Asn Gly Phe Gly Arg Ile Gly Arg Ile
1 5 10 15
Val Phe Arg Ala Ala Gln Lys Arg Ser Asp Ile Glu Ile Val Ala Ile
20 25 30
Asn Asp Leu Leu Asp Ala Asp Tyr Met Ala Tyr Met Leu Lys Tyr Asp
35 40 45
Ser Thr His Gly Arg Phe Asp Gly Thr Val Glu Val Lys Asp Gly His
50 55 60
Leu Ile Val Asn Gly Lys Lys Ile Arg Val Thr Ala Glu Arg Asp Pro
65 70 75 80
Ala Asn Leu Lys Trp Asp Glu Val Gly Val Asp Val Val Ala Glu Ala
85 90 95
Thr Gly Leu Phe Leu Thr Asp Glu Thr Ala Arg Lys His Ile Thr Ala
100 105 110
Gly Ala Lys Lys Val Val Met Thr Gly Pro Ser Lys Asp Asn Thr Pro
115 120 125
Met Phe Val Lys Gly Ala Asn Phe Asp Lys Tyr Ala Gly Gln Asp Ile
130 135 140
Val Ser Asn Ala Ser Cys Thr Thr Asn Cys Leu Ala Pro Leu Ala Lys
145 150 155 160
Val Ile Asn Asp Asn Phe Gly Ile Ile Glu Gly Leu Met Thr Thr Val
165 170 175
His Ala Thr Thr Ala Thr Gln Lys Thr Val Asp Gly Pro Ser His Lys
180 185 190
Asp Trp Arg Gly Gly Arg Gly Ala Ser Gln Asn Ile Ile Pro Ser Ser
195 200 205
Thr Gly Ala Ala Lys Ala Val Gly Lys Val Leu Pro Glu Leu Asn Gly
210 215 220
Lys Leu Thr Gly Met Ala Phe Arg Val Pro Thr Pro Asn Val Ser Val
225 230 235 240
Val Asp Leu Thr Val Arg Leu Glu Lys Ala Ala Thr Tyr Glu Gln Ile
245 250 255
Lys Ala Ala Val Lys Ala Ala Ala Glu Gly Glu Met Lys Gly Val Leu
260 265 270
Gly Tyr Thr Glu Asp Asp Val Val Ser Thr Asp Phe Asn Gly Glu Val
275 280 285
Cys Thr Ser Val Phe Asp Ala Lys Ala Gly Ile Ala Leu Asn Asp Asn
290 295 300
Phe Val Lys Leu Val Ser Trp Tyr Asp Asn Glu Thr Gly Tyr Ser Asn
305 310 315 320
Lys Val Leu Asp Leu Ile Ala His Ile Ser Lys
325 330
<210> 175
<211> 387
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 175
Met Ser Val Ile Lys Met Thr Asp Leu Asp Leu Ala Gly Lys Arg Val
1 5 10 15
Phe Ile Arg Ala Asp Leu Asn Val Pro Val Lys Asp Gly Lys Val Thr
20 25 30
Ser Asp Ala Arg Ile Arg Ala Ser Leu Pro Thr Ile Glu Leu Ala Leu
35 40 45
Lys Gln Gly Ala Lys Val Met Val Thr Ser His Leu Gly Arg Pro Thr
50 55 60
Glu Gly Glu Tyr Asn Glu Glu Phe Ser Leu Leu Pro Val Val Asn Tyr
65 70 75 80
Leu Lys Asp Lys Leu Ser Asn Pro Val Arg Leu Val Lys Asp Tyr Leu
85 90 95
Asp Gly Val Asp Val Ala Glu Gly Glu Leu Val Val Leu Glu Asn Val
100 105 110
Arg Phe Asn Lys Gly Glu Lys Lys Asp Asp Glu Thr Leu Ser Lys Lys
115 120 125
Tyr Ala Ala Leu Cys Asp Val Phe Val Met Asp Ala Phe Gly Thr Ala
130 135 140
His Arg Ala Gln Ala Ser Thr His Gly Ile Gly Lys Phe Ala Asp Val
145 150 155 160
Ala Cys Ala Gly Pro Leu Leu Ala Ala Glu Leu Asp Ala Leu Gly Lys
165 170 175
Ala Leu Lys Glu Pro Ala Arg Pro Met Val Ala Ile Val Gly Gly Ser
180 185 190
Lys Val Ser Thr Lys Leu Thr Val Leu Asp Ser Leu Ser Lys Ile Ala
195 200 205
Asp Gln Leu Ile Val Gly Gly Gly Ile Ala Asn Thr Phe Ile Ala Ala
210 215 220
Gln Gly His Asp Val Gly Lys Ser Leu Tyr Glu Ala Asp Leu Val Asp
225 230 235 240
Glu Ala Lys Arg Leu Leu Thr Thr Cys Asn Ile Pro Val Pro Ser Asp
245 250 255
Val Arg Val Ala Thr Glu Phe Ser Glu Thr Ala Pro Ala Thr Leu Lys
260 265 270
Ser Val Asn Asp Val Lys Ala Asp Glu Gln Ile Leu Asp Ile Gly Asp
275 280 285
Ala Ser Ala Gln Glu Leu Ala Glu Ile Leu Lys Asn Ala Lys Thr Ile
290 295 300
Leu Trp Asn Gly Pro Val Gly Val Phe Glu Phe Pro Asn Phe Arg Lys
305 310 315 320
Gly Thr Glu Ile Val Ala Asn Ala Ile Ala Asp Ser Glu Ala Phe Ser
325 330 335
Ile Ala Gly Gly Gly Asp Thr Leu Ala Ala Ile Asp Leu Phe Gly Ile
340 345 350
Ala Asp Lys Ile Ser Tyr Ile Ser Thr Gly Gly Gly Ala Phe Leu Glu
355 360 365
Phe Val Glu Gly Lys Val Leu Pro Ala Val Ala Met Leu Glu Glu Arg
370 375 380
Ala Lys Lys
385
<210> 176
<211> 652
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 176
Met Ser Gln Ile His Lys His Thr Ile Pro Ala Asn Ile Ala Asp Arg
1 5 10 15
Cys Leu Ile Asn Pro Gln Gln Tyr Glu Ala Met Tyr Gln Gln Ser Ile
20 25 30
Asn Val Pro Asp Thr Phe Trp Gly Glu Gln Gly Lys Ile Leu Asp Trp
35 40 45
Ile Lys Pro Tyr Gln Lys Val Lys Asn Thr Ser Phe Ala Pro Gly Asn
50 55 60
Val Ser Ile Lys Trp Tyr Glu Asp Gly Thr Leu Asn Leu Ala Ala Asn
65 70 75 80
Cys Leu Asp Arg His Leu Gln Glu Asn Gly Asp Arg Thr Ala Ile Ile
85 90 95
Trp Glu Gly Asp Asp Ala Ser Gln Ser Lys His Ile Ser Tyr Lys Glu
100 105 110
Leu His Arg Asp Val Cys Arg Phe Ala Asn Thr Leu Leu Glu Leu Gly
115 120 125
Ile Lys Lys Gly Asp Val Val Ala Ile Tyr Met Pro Met Val Pro Glu
130 135 140
Ala Ala Val Ala Met Leu Ala Cys Ala Arg Ile Gly Ala Val His Ser
145 150 155 160
Val Ile Phe Gly Gly Phe Ser Pro Glu Ala Val Ala Gly Arg Ile Ile
165 170 175
Asp Ser Asn Ser Arg Leu Val Ile Thr Ser Asp Glu Gly Val Arg Ala
180 185 190
Gly Arg Ser Ile Pro Leu Lys Lys Asn Val Asp Asp Ala Leu Lys Asn
195 200 205
Pro Asn Val Thr Ser Val Glu His Val Val Val Leu Lys Arg Thr Gly
210 215 220
Gly Lys Ile Asp Trp Gln Glu Gly Arg Asp Leu Trp Trp His Asp Leu
225 230 235 240
Val Glu Gln Ala Ser Asp Gln His Gln Ala Glu Glu Met Asn Ala Glu
245 250 255
Asp Pro Leu Phe Ile Leu Tyr Thr Ser Gly Ser Thr Gly Lys Pro Lys
260 265 270
Gly Val Leu His Thr Thr Gly Gly Tyr Leu Val Tyr Ala Ala Leu Thr
275 280 285
Phe Lys Tyr Val Phe Asp Tyr His Pro Gly Asp Ile Tyr Trp Cys Thr
290 295 300
Ala Asp Val Gly Trp Val Thr Gly His Ser Tyr Leu Leu Tyr Gly Pro
305 310 315 320
Leu Ala Cys Gly Ala Thr Thr Leu Met Phe Glu Gly Val Pro Asn Trp
325 330 335
Pro Thr Pro Ala Arg Met Ala Gln Val Val Asp Lys His Gln Val Asn
340 345 350
Ile Leu Tyr Thr Ala Pro Thr Ala Ile Arg Ala Leu Met Ala Glu Gly
355 360 365
Asp Lys Ala Ile Glu Gly Thr Asp Arg Ser Ser Leu Arg Ile Leu Gly
370 375 380
Ser Val Gly Glu Pro Ile Asn Pro Glu Ala Trp Glu Trp Tyr Trp Lys
385 390 395 400
Lys Ile Gly Asn Glu Lys Cys Pro Val Val Asp Thr Trp Trp Gln Thr
405 410 415
Glu Thr Gly Gly Phe Met Ile Thr Pro Leu Pro Gly Ala Thr Glu Leu
420 425 430
Lys Ala Gly Ser Ala Thr Arg Pro Phe Phe Gly Val Gln Pro Ala Leu
435 440 445
Val Asp Asn Glu Gly Asn Pro Leu Glu Gly Ala Thr Glu Gly Ser Leu
450 455 460
Val Ile Thr Asp Ser Trp Pro Gly Gln Ala Arg Thr Leu Phe Gly Asp
465 470 475 480
His Glu Arg Phe Glu Gln Thr Tyr Phe Ser Thr Phe Lys Asn Met Tyr
485 490 495
Phe Ser Gly Asp Gly Ala Arg Arg Asp Glu Asp Gly Tyr Tyr Trp Ile
500 505 510
Thr Gly Arg Val Asp Asp Val Leu Asn Val Ser Gly His Arg Leu Gly
515 520 525
Thr Ala Glu Ile Glu Ser Ala Leu Val Ala His Pro Lys Ile Ala Glu
530 535 540
Ala Ala Val Val Gly Ile Pro His Asn Ile Lys Gly Gln Ala Ile Tyr
545 550 555 560
Ala Tyr Val Thr Leu Asn His Gly Glu Glu Pro Ser Pro Glu Leu Tyr
565 570 575
Ala Glu Val Arg Asn Trp Val Arg Lys Glu Ile Gly Pro Leu Ala Thr
580 585 590
Pro Asp Val Leu His Trp Thr Asp Ser Leu Pro Lys Thr Arg Ser Gly
595 600 605
Lys Ile Met Arg Arg Ile Leu Arg Lys Ile Ala Ala Gly Asp Thr Ser
610 615 620
Asn Leu Gly Asp Thr Ser Thr Leu Ala Asp Pro Gly Val Val Glu Lys
625 630 635 640
Leu Leu Glu Glu Lys Gln Ala Ile Ala Met Pro Ser
645 650
<210> 177
<211> 887
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 177
Met Ser Glu Arg Phe Pro Asn Asp Val Asp Pro Ile Glu Thr Arg Asp
1 5 10 15
Trp Leu Gln Ala Ile Glu Ser Val Ile Arg Glu Glu Gly Val Glu Arg
20 25 30
Ala Gln Tyr Leu Ile Asp Gln Leu Leu Ala Glu Ala Arg Lys Gly Gly
35 40 45
Val Asn Val Ala Ala Gly Thr Gly Ile Ser Asn Tyr Ile Asn Thr Ile
50 55 60
Pro Val Glu Glu Gln Pro Glu Tyr Pro Gly Asn Leu Glu Leu Glu Arg
65 70 75 80
Arg Ile Arg Ser Ala Ile Arg Trp Asn Ala Ile Met Thr Val Leu Arg
85 90 95
Ala Ser Lys Lys Asp Leu Glu Leu Gly Gly His Met Ala Ser Phe Gln
100 105 110
Ser Ser Ala Thr Ile Tyr Asp Val Cys Phe Asn His Phe Phe Arg Ala
115 120 125
Arg Asn Glu Gln Asp Gly Gly Asp Leu Val Tyr Phe Gln Gly His Ile
130 135 140
Ser Pro Gly Val Tyr Ala Arg Ala Phe Leu Glu Gly Arg Leu Thr Gln
145 150 155 160
Glu Gln Leu Asp Asn Phe Arg Gln Glu Val His Gly Asn Gly Leu Ser
165 170 175
Ser Tyr Pro His Pro Lys Leu Met Pro Glu Phe Trp Gln Phe Pro Thr
180 185 190
Val Ser Met Gly Leu Gly Pro Ile Gly Ala Ile Tyr Gln Ala Lys Phe
195 200 205
Leu Lys Tyr Leu Glu His Arg Gly Leu Lys Asp Thr Ser Lys Gln Thr
210 215 220
Val Tyr Ala Phe Leu Gly Asp Gly Glu Met Asp Glu Pro Glu Ser Lys
225 230 235 240
Gly Ala Ile Thr Ile Ala Thr Arg Glu Lys Leu Asp Asn Leu Val Phe
245 250 255
Val Ile Asn Cys Asn Leu Gln Arg Leu Asp Gly Pro Val Thr Gly Asn
260 265 270
Gly Lys Ile Ile Asn Glu Leu Glu Gly Ile Phe Glu Gly Ala Gly Trp
275 280 285
Asn Val Ile Lys Val Met Trp Gly Ser Arg Trp Asp Glu Leu Leu Arg
290 295 300
Lys Asp Thr Ser Gly Lys Leu Ile Gln Leu Met Asn Glu Thr Val Asp
305 310 315 320
Gly Asp Tyr Gln Thr Phe Lys Ser Lys Asp Gly Ala Tyr Val Arg Glu
325 330 335
His Phe Phe Gly Lys Tyr Pro Glu Thr Ala Ala Leu Val Ala Asp Trp
340 345 350
Thr Asp Glu Gln Ile Trp Ala Leu Asn Arg Gly Gly His Asp Pro Lys
355 360 365
Lys Ile Tyr Ala Ala Phe Lys Lys Ala Gln Glu Thr Lys Gly Lys Ala
370 375 380
Thr Val Ile Leu Ala His Thr Ile Lys Gly Tyr Gly Met Gly Asp Ala
385 390 395 400
Ala Glu Gly Lys Asn Ile Ala His Gln Val Lys Lys Met Asn Met Asp
405 410 415
Gly Val Arg His Ile Arg Asp Arg Phe Asn Val Pro Val Ser Asp Ala
420 425 430
Asp Ile Glu Lys Leu Pro Tyr Ile Thr Phe Pro Glu Gly Ser Glu Glu
435 440 445
His Thr Tyr Leu His Ala Gln Arg Gln Lys Leu His Gly Tyr Leu Pro
450 455 460
Ser Arg Gln Pro Asn Phe Thr Glu Lys Leu Glu Leu Pro Ser Leu Gln
465 470 475 480
Asp Phe Gly Ala Leu Leu Glu Glu Gln Ser Lys Glu Ile Ser Thr Thr
485 490 495
Ile Ala Phe Val Arg Ala Leu Asn Val Met Leu Lys Asn Lys Ser Ile
500 505 510
Lys Asp Arg Leu Val Pro Ile Ile Ala Asp Glu Ala Arg Thr Phe Gly
515 520 525
Met Glu Gly Leu Phe Arg Gln Ile Gly Ile Tyr Ser Pro Asn Gly Gln
530 535 540
Gln Tyr Thr Pro Gln Asp Arg Glu Gln Val Ala Tyr Tyr Lys Glu Asp
545 550 555 560
Glu Lys Gly Gln Ile Leu Gln Glu Gly Ile Asn Glu Leu Gly Ala Gly
565 570 575
Cys Ser Trp Leu Ala Ala Ala Thr Ser Tyr Ser Thr Asn Asn Leu Pro
580 585 590
Met Ile Pro Phe Tyr Ile Tyr Tyr Ser Met Phe Gly Phe Gln Arg Ile
595 600 605
Gly Asp Leu Cys Trp Ala Ala Gly Asp Gln Gln Ala Arg Gly Phe Leu
610 615 620
Ile Gly Gly Thr Ser Gly Arg Thr Thr Leu Asn Gly Glu Gly Leu Gln
625 630 635 640
His Glu Asp Gly His Ser His Ile Gln Ser Leu Thr Ile Pro Asn Cys
645 650 655
Ile Ser Tyr Asp Pro Ala Tyr Ala Tyr Glu Val Ala Val Ile Met His
660 665 670
Asp Gly Leu Glu Arg Met Tyr Gly Glu Lys Gln Glu Asn Val Tyr Tyr
675 680 685
Tyr Ile Thr Thr Leu Asn Glu Asn Tyr His Met Pro Ala Met Pro Glu
690 695 700
Gly Ala Glu Glu Gly Ile Arg Lys Gly Ile Tyr Lys Leu Glu Thr Ile
705 710 715 720
Glu Gly Ser Lys Gly Lys Val Gln Leu Leu Gly Ser Gly Ser Ile Leu
725 730 735
Arg His Val Arg Glu Ala Ala Glu Ile Leu Ala Lys Asp Tyr Gly Val
740 745 750
Gly Ser Asp Val Tyr Ser Val Thr Ser Phe Thr Glu Leu Ala Arg Asp
755 760 765
Gly Gln Asp Cys Glu Arg Trp Asn Met Leu His Pro Leu Glu Thr Pro
770 775 780
Arg Val Pro Tyr Ile Ala Gln Val Met Asn Asp Ala Pro Ala Val Ala
785 790 795 800
Ser Thr Asp Tyr Met Lys Leu Phe Ala Glu Gln Val Arg Thr Tyr Val
805 810 815
Pro Ala Asp Asp Tyr Arg Val Leu Gly Thr Asp Gly Phe Gly Arg Ser
820 825 830
Asp Ser Arg Glu Asn Leu Arg His His Phe Glu Val Asp Ala Ser Tyr
835 840 845
Val Val Val Ala Ala Leu Gly Glu Leu Ala Lys Arg Gly Glu Ile Asp
850 855 860
Lys Lys Val Val Ala Asp Ala Ile Ala Lys Phe Asn Ile Asp Ala Asp
865 870 875 880
Lys Val Asn Pro Arg Leu Ala
885
<210> 178
<211> 630
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 178
Met Ala Ile Glu Ile Lys Val Pro Asp Ile Gly Ala Asp Glu Val Glu
1 5 10 15
Ile Thr Glu Ile Leu Val Lys Val Gly Asp Lys Val Glu Ala Glu Gln
20 25 30
Ser Leu Ile Thr Val Glu Gly Asp Lys Ala Ser Met Glu Val Pro Ser
35 40 45
Pro Gln Ala Gly Ile Val Lys Glu Ile Lys Val Ser Val Gly Asp Lys
50 55 60
Thr Gln Thr Gly Ala Leu Ile Met Ile Phe Asp Ser Ala Asp Gly Ala
65 70 75 80
Ala Asp Ala Ala Pro Ala Gln Ala Glu Glu Lys Lys Glu Ala Ala Pro
85 90 95
Ala Ala Ala Pro Ala Ala Ala Ala Ala Lys Asp Val Asn Val Pro Asp
100 105 110
Ile Gly Ser Asp Glu Val Glu Val Thr Glu Ile Leu Val Lys Val Gly
115 120 125
Asp Lys Val Glu Ala Glu Gln Ser Leu Ile Thr Val Glu Gly Asp Lys
130 135 140
Ala Ser Met Glu Val Pro Ala Pro Phe Ala Gly Thr Val Lys Glu Ile
145 150 155 160
Lys Val Asn Val Gly Asp Lys Val Ser Thr Gly Ser Leu Ile Met Val
165 170 175
Phe Glu Val Ala Gly Glu Ala Gly Ala Ala Ala Pro Ala Ala Lys Gln
180 185 190
Glu Ala Ala Pro Ala Ala Ala Pro Ala Pro Ala Ala Gly Val Lys Glu
195 200 205
Val Asn Val Pro Asp Ile Gly Gly Asp Glu Val Glu Val Thr Glu Val
210 215 220
Met Val Lys Val Gly Asp Lys Val Ala Ala Glu Gln Ser Leu Ile Thr
225 230 235 240
Val Glu Gly Asp Lys Ala Ser Met Glu Val Pro Ala Pro Phe Ala Gly
245 250 255
Val Val Lys Glu Leu Lys Val Asn Val Gly Asp Lys Val Lys Thr Gly
260 265 270
Ser Leu Ile Met Ile Phe Glu Val Glu Gly Ala Ala Pro Ala Ala Ala
275 280 285
Pro Ala Lys Gln Glu Ala Ala Ala Pro Ala Pro Ala Ala Lys Ala Glu
290 295 300
Ala Pro Ala Ala Ala Pro Ala Ala Lys Ala Glu Gly Lys Ser Glu Phe
305 310 315 320
Ala Glu Asn Asp Ala Tyr Val His Ala Thr Pro Leu Ile Arg Arg Leu
325 330 335
Ala Arg Glu Phe Gly Val Asn Leu Ala Lys Val Lys Gly Thr Gly Arg
340 345 350
Lys Gly Arg Ile Leu Arg Glu Asp Val Gln Ala Tyr Val Lys Glu Ala
355 360 365
Ile Lys Arg Ala Glu Ala Ala Pro Ala Ala Thr Gly Gly Gly Ile Pro
370 375 380
Gly Met Leu Pro Trp Pro Lys Val Asp Phe Ser Lys Phe Gly Glu Ile
385 390 395 400
Glu Glu Val Glu Leu Gly Arg Ile Gln Lys Ile Ser Gly Ala Asn Leu
405 410 415
Ser Arg Asn Trp Val Met Ile Pro His Val Thr His Phe Asp Lys Thr
420 425 430
Asp Ile Thr Glu Leu Glu Ala Phe Arg Lys Gln Gln Asn Glu Glu Ala
435 440 445
Ala Lys Arg Lys Leu Asp Val Lys Ile Thr Pro Val Val Phe Ile Met
450 455 460
Lys Ala Val Ala Ala Ala Leu Glu Gln Met Pro Arg Phe Asn Ser Ser
465 470 475 480
Leu Ser Glu Asp Gly Gln Arg Leu Thr Leu Lys Lys Tyr Ile Asn Ile
485 490 495
Gly Val Ala Val Asp Thr Pro Asn Gly Leu Val Val Pro Val Phe Lys
500 505 510
Asp Val Asn Lys Lys Gly Ile Ile Glu Leu Ser Arg Glu Leu Met Thr
515 520 525
Ile Ser Lys Lys Ala Arg Asp Gly Lys Leu Thr Ala Gly Glu Met Gln
530 535 540
Gly Gly Cys Phe Thr Ile Ser Ser Ile Gly Gly Leu Gly Thr Thr His
545 550 555 560
Phe Ala Pro Ile Val Asn Ala Pro Glu Val Ala Ile Leu Gly Val Ser
565 570 575
Lys Ser Ala Met Glu Pro Val Trp Asn Gly Lys Glu Phe Val Pro Arg
580 585 590
Leu Met Leu Pro Ile Ser Leu Ser Phe Asp His Arg Val Ile Asp Gly
595 600 605
Ala Asp Gly Ala Arg Phe Ile Thr Ile Ile Asn Asn Thr Leu Ser Asp
610 615 620
Ile Arg Arg Leu Val Met
625 630
<210> 179
<211> 474
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 179
Met Ser Thr Glu Ile Lys Thr Gln Val Val Val Leu Gly Ala Gly Pro
1 5 10 15
Ala Gly Tyr Ser Ala Ala Phe Arg Cys Ala Asp Leu Gly Leu Glu Thr
20 25 30
Val Ile Val Glu Arg Tyr Asn Thr Leu Gly Gly Val Cys Leu Asn Val
35 40 45
Gly Cys Ile Pro Ser Lys Ala Leu Leu His Val Ala Lys Val Ile Glu
50 55 60
Glu Ala Lys Ala Leu Ala Glu His Gly Ile Val Phe Gly Glu Pro Lys
65 70 75 80
Thr Asp Ile Asp Lys Ile Arg Thr Trp Lys Glu Lys Val Ile Asn Gln
85 90 95
Leu Thr Gly Gly Leu Ala Gly Met Ala Lys Gly Arg Lys Val Lys Val
100 105 110
Val Asn Gly Leu Gly Lys Phe Thr Gly Ala Asn Thr Leu Glu Val Glu
115 120 125
Gly Glu Asn Gly Lys Thr Val Ile Asn Phe Asp Asn Ala Ile Ile Ala
130 135 140
Ala Gly Ser Arg Pro Ile Gln Leu Pro Phe Ile Pro His Glu Asp Pro
145 150 155 160
Arg Ile Trp Asp Ser Thr Asp Ala Leu Glu Leu Lys Glu Val Pro Glu
165 170 175
Arg Leu Leu Val Met Gly Gly Gly Ile Ile Gly Leu Glu Met Gly Thr
180 185 190
Val Tyr His Ala Leu Gly Ser Gln Ile Asp Val Val Glu Met Phe Asp
195 200 205
Gln Val Ile Pro Ala Ala Asp Lys Asp Ile Val Lys Val Phe Thr Lys
210 215 220
Arg Ile Ser Lys Lys Phe Asn Leu Met Leu Glu Thr Lys Val Thr Ala
225 230 235 240
Val Glu Ala Lys Glu Asp Gly Ile Tyr Val Thr Met Glu Gly Lys Lys
245 250 255
Ala Pro Ala Glu Pro Gln Arg Tyr Asp Ala Val Leu Val Ala Ile Gly
260 265 270
Arg Val Pro Asn Gly Lys Asn Leu Asp Ala Gly Lys Ala Gly Val Glu
275 280 285
Val Asp Asp Arg Gly Phe Ile Arg Val Asp Lys Gln Leu Arg Thr Asn
290 295 300
Val Pro His Ile Phe Ala Ile Gly Asp Ile Val Gly Gln Pro Met Leu
305 310 315 320
Ala His Lys Gly Val His Glu Gly His Val Ala Ala Glu Val Ile Ala
325 330 335
Gly Lys Lys His Tyr Phe Asp Pro Lys Val Ile Pro Ser Ile Ala Tyr
340 345 350
Thr Glu Pro Glu Val Ala Trp Val Gly Leu Thr Glu Lys Glu Ala Lys
355 360 365
Glu Lys Gly Ile Ser Tyr Glu Thr Ala Thr Phe Pro Trp Ala Ala Ser
370 375 380
Gly Arg Ala Ile Ala Ser Asp Cys Ala Asp Gly Met Thr Lys Leu Ile
385 390 395 400
Phe Asp Lys Glu Ser His Arg Val Ile Gly Gly Ala Ile Val Gly Thr
405 410 415
Asn Gly Gly Glu Leu Leu Gly Glu Ile Gly Leu Ala Ile Glu Met Gly
420 425 430
Cys Asp Ala Glu Asp Ile Ala Leu Thr Ile His Ala His Pro Thr Leu
435 440 445
His Glu Ser Val Gly Leu Ala Ala Glu Val Phe Glu Gly Ser Ile Thr
450 455 460
Asp Leu Pro Asn Pro Lys Ala Lys Lys Lys
465 470
<210> 180
<211> 591
<212> PRT
<213> Corynebacterium glutamicum
<400> 180
Val Ser Val Glu Thr Arg Lys Ile Thr Lys Val Leu Val Ala Asn Arg
1 5 10 15
Gly Glu Ile Ala Ile Arg Val Phe Arg Ala Ala Arg Asp Glu Gly Ile
20 25 30
Gly Ser Val Ala Val Tyr Ala Glu Pro Asp Ala Asp Ala Pro Phe Val
35 40 45
Ser Tyr Ala Asp Glu Ala Phe Ala Leu Gly Gly Gln Thr Ser Ala Glu
50 55 60
Ser Tyr Leu Val Ile Asp Lys Ile Ile Asp Ala Ala Arg Lys Ser Gly
65 70 75 80
Ala Asp Ala Ile His Pro Gly Tyr Gly Phe Leu Ala Glu Asn Ala Asp
85 90 95
Phe Ala Glu Ala Val Ile Asn Glu Gly Leu Ile Trp Ile Gly Pro Ser
100 105 110
Pro Glu Ser Ile Arg Ser Leu Gly Asp Lys Val Thr Ala Arg His Ile
115 120 125
Ala Asp Thr Ala Lys Ala Pro Met Ala Pro Gly Thr Lys Glu Pro Val
130 135 140
Lys Asp Ala Ala Glu Val Val Ala Phe Ala Glu Glu Phe Gly Leu Pro
145 150 155 160
Ile Ala Ile Lys Ala Ala Phe Gly Gly Gly Gly Arg Gly Met Lys Val
165 170 175
Ala Tyr Lys Met Glu Glu Val Ala Asp Leu Phe Glu Ser Ala Thr Arg
180 185 190
Glu Ala Thr Ala Ala Phe Gly Arg Gly Glu Cys Phe Val Glu Arg Tyr
195 200 205
Leu Asp Lys Ala Arg His Val Glu Ala Gln Val Ile Ala Asp Lys His
210 215 220
Gly Asn Val Val Val Ala Gly Thr Arg Asp Cys Ser Leu Gln Arg Arg
225 230 235 240
Phe Gln Lys Leu Val Glu Glu Ala Pro Ala Pro Phe Leu Thr Asp Asp
245 250 255
Gln Arg Glu Arg Leu His Ser Ser Ala Lys Ala Ile Cys Lys Glu Ala
260 265 270
Gly Tyr Tyr Gly Ala Gly Thr Val Glu Tyr Leu Val Gly Ser Asp Gly
275 280 285
Leu Ile Ser Phe Leu Glu Val Asn Thr Arg Leu Gln Val Glu His Pro
290 295 300
Val Thr Glu Glu Thr Thr Gly Ile Asp Leu Val Arg Glu Met Phe Arg
305 310 315 320
Ile Ala Glu Gly His Glu Leu Ser Ile Lys Glu Asp Pro Ala Pro Arg
325 330 335
Gly His Ala Phe Glu Phe Arg Ile Asn Gly Glu Asp Ala Gly Ser Asn
340 345 350
Phe Met Pro Ala Pro Gly Lys Ile Thr Ser Tyr Arg Glu Pro Gln Gly
355 360 365
Pro Gly Val Arg Met Asp Ser Gly Val Val Glu Gly Ser Glu Ile Ser
370 375 380
Gly Gln Phe Asp Ser Met Leu Ala Lys Leu Ile Val Trp Gly Asp Thr
385 390 395 400
Arg Glu Gln Ala Leu Gln Arg Ser Arg Arg Ala Leu Ala Glu Tyr Val
405 410 415
Val Glu Gly Met Pro Thr Val Ile Pro Phe His Gln His Ile Val Glu
420 425 430
Asn Pro Ala Phe Val Gly Asn Asp Glu Gly Phe Glu Ile Tyr Thr Lys
435 440 445
Trp Ile Glu Glu Val Trp Asp Asn Pro Ile Ala Pro Tyr Val Asp Ala
450 455 460
Ser Glu Leu Asp Glu Asp Glu Asp Lys Thr Pro Ala Gln Lys Val Val
465 470 475 480
Val Glu Ile Asn Gly Arg Arg Val Glu Val Ala Leu Pro Gly Asp Leu
485 490 495
Ala Leu Gly Gly Thr Ala Gly Pro Lys Lys Lys Ala Lys Lys Arg Arg
500 505 510
Ala Gly Gly Ala Lys Ala Gly Val Ser Gly Asp Ala Val Ala Ala Pro
515 520 525
Met Gln Gly Thr Val Ile Lys Val Asn Val Glu Glu Gly Ala Glu Val
530 535 540
Asn Glu Gly Asp Thr Val Val Val Leu Glu Ala Met Lys Met Glu Asn
545 550 555 560
Pro Val Lys Ala His Lys Ser Gly Thr Val Thr Gly Leu Thr Val Ala
565 570 575
Ala Gly Glu Gly Val Asn Lys Gly Val Val Leu Leu Glu Ile Lys
580 585 590
<210> 181
<211> 543
<212> PRT
<213> Corynebacterium glutamicum
<400> 181
Met Thr Ile Ser Ser Pro Leu Ile Asp Val Ala Asn Leu Pro Asp Ile
1 5 10 15
Asn Thr Thr Ala Gly Lys Ile Ala Asp Leu Lys Ala Arg Arg Ala Glu
20 25 30
Ala His Phe Pro Met Gly Glu Lys Ala Val Glu Lys Val His Ala Ala
35 40 45
Gly Arg Leu Thr Ala Arg Glu Arg Leu Asp Tyr Leu Leu Asp Glu Gly
50 55 60
Ser Phe Ile Glu Thr Asp Gln Leu Ala Arg His Arg Thr Thr Ala Phe
65 70 75 80
Gly Leu Gly Ala Lys Arg Pro Ala Thr Asp Gly Ile Val Thr Gly Trp
85 90 95
Gly Thr Ile Asp Gly Arg Glu Val Cys Ile Phe Ser Gln Asp Gly Thr
100 105 110
Val Phe Gly Gly Ala Leu Gly Glu Val Tyr Gly Glu Lys Met Ile Lys
115 120 125
Ile Met Glu Leu Ala Ile Asp Thr Gly Arg Pro Leu Ile Gly Leu Tyr
130 135 140
Glu Gly Ala Gly Ala Arg Ile Gln Asp Gly Ala Val Ser Leu Asp Phe
145 150 155 160
Ile Ser Gln Thr Phe Tyr Gln Asn Ile Gln Ala Ser Gly Val Ile Pro
165 170 175
Gln Ile Ser Val Ile Met Gly Ala Cys Ala Gly Gly Asn Ala Tyr Gly
180 185 190
Pro Ala Leu Thr Asp Phe Val Val Met Val Asp Lys Thr Ser Lys Met
195 200 205
Phe Val Thr Gly Pro Asp Val Ile Lys Thr Val Thr Gly Glu Glu Ile
210 215 220
Thr Gln Glu Glu Leu Gly Gly Ala Thr Thr His Met Val Thr Ala Gly
225 230 235 240
Asn Ser His Tyr Thr Ala Ala Thr Asp Glu Glu Ala Leu Asp Trp Val
245 250 255
Gln Asp Leu Val Ser Phe Leu Pro Ser Asn Asn Arg Ser Tyr Thr Pro
260 265 270
Leu Glu Asp Phe Asp Glu Glu Glu Gly Gly Val Glu Glu Asn Ile Thr
275 280 285
Ala Asp Asp Leu Lys Leu Asp Glu Ile Ile Pro Asp Ser Ala Thr Val
290 295 300
Pro Tyr Asp Val Arg Asp Val Ile Glu Cys Leu Thr Asp Asp Gly Glu
305 310 315 320
Tyr Leu Glu Ile Gln Ala Asp Arg Ala Glu Asn Val Val Ile Ala Phe
325 330 335
Gly Arg Ile Glu Gly Gln Ser Val Gly Phe Val Ala Asn Gln Pro Thr
340 345 350
Gln Phe Ala Gly Cys Leu Asp Ile Asp Ser Ser Glu Lys Ala Ala Arg
355 360 365
Phe Val Arg Thr Cys Asp Ala Phe Asn Ile Pro Ile Val Met Leu Val
370 375 380
Asp Val Pro Gly Phe Leu Pro Gly Ala Gly Gln Glu Tyr Gly Gly Ile
385 390 395 400
Leu Arg Arg Gly Ala Lys Leu Leu Tyr Ala Tyr Gly Glu Ala Thr Val
405 410 415
Pro Lys Ile Thr Val Thr Met Arg Lys Ala Tyr Gly Gly Ala Tyr Cys
420 425 430
Val Met Gly Ser Lys Gly Leu Gly Ser Asp Ile Asn Leu Ala Trp Pro
435 440 445
Thr Ala Gln Ile Ala Val Met Gly Ala Ala Gly Ala Val Gly Phe Ile
450 455 460
Tyr Arg Lys Glu Leu Met Ala Ala Asp Ala Lys Gly Leu Asp Thr Val
465 470 475 480
Ala Leu Ala Lys Ser Phe Glu Arg Glu Tyr Glu Asp His Met Leu Asn
485 490 495
Pro Tyr His Ala Ala Glu Arg Gly Leu Ile Asp Ala Val Ile Leu Pro
500 505 510
Ser Glu Thr Arg Gly Gln Ile Ser Arg Asn Leu Arg Leu Leu Lys His
515 520 525
Lys Asn Val Thr Arg Pro Ala Arg Lys His Gly Asn Met Pro Leu
530 535 540
Claims (33)
1.一种用于丙二酰-CoA检测的重组微生物，在其中III型聚酮合酶编码基因插入所述微生物的基因组中，或在其中引入了含有所述III型聚酮合酶编码基因的重组载体。
2.权利要求1的重组微生物，其中所述III型聚酮合酶是：
源自选自灰色链霉菌、天蓝色链霉菌、阿维链霉菌、红色糖多孢菌、波赛链霉菌和针孢链霉菌的微生物的RppA；
源自荧光假单胞菌的PhlD(聚酮合酶)；
源自地中海拟无枝菌酸菌的DpgA(聚酮合酶)；
源自掌叶大黄的ALS(芦荟酮合酶)；或
源自木立芦荟的PCS(5,7-二羟基-2-甲基色酮合酶)、OKS(八酮合酶)、PKS3(芦荟酮合酶)、PKS4(八酮合酶2)或PKS5(八酮合酶3)。
3.权利要求1的重组微生物，其中编码所述III型聚酮合酶的基因与选自tac、trc、T7、BAD、λPR和Anderson合成启动子的启动子可操作地连接。
4.权利要求1的重组微生物，其中所述重组微生物选自大肠杆菌、根瘤菌属、双歧杆菌属、红球菌属、假丝酵母属、欧文氏菌属、肠杆菌属、巴氏杆菌属、曼氏杆菌属、放线杆菌属、凝聚杆菌属、黄单胞菌属、弧菌属、假单胞菌属、固氮菌属、不动杆菌属、劳尔氏菌属、土壤杆菌属、红杆菌属、发酵单胞菌属、芽孢杆菌属、葡萄球菌属、乳球菌属、链球菌属、乳杆菌属、梭菌属、棒杆菌属、链霉菌属、双歧杆菌属、蓝细菌和圆杆菌属。
5.一种筛选丙二酰-CoA产生诱导物质的方法，其包括以下步骤：
(a)培养权利要求1的重组微生物；
(b)向所述重组微生物中添加候选物质；
(c)将添加所述候选物质后的所述重组微生物的培养上清液的颜色与没有添加所述候选物质的所述重组微生物的培养上清液的颜色进行比较；并且
(d)当添加所述候选物质后所述重组微生物的培养上清液显示出比没有添加所述候选物质的所述重组微生物的培养上清液更深的红色时，选择所述候选物质作为所述丙二酰-CoA产生诱导物质。
6.权利要求5的方法，其中步骤(c)中颜色之间的比较是通过肉眼进行。
7.权利要求5的方法，其中步骤(c)中颜色之间的比较是通过测量吸光度来进行，并且步骤(d)包括当添加所述候选物质后测量的吸光度高于未添加所述候选物质的情况下测量的吸光度时，选择所述候选物质作为所述丙二酰-CoA产生诱导物质。
8.一种筛选参与增加丙二酰-CoA产生的基因的方法，所述方法包括以下步骤：
(a)将改变重组微生物中的基因表达的基因调节文库引入权利要求1的重组微生物中，从而构建所述重组微生物中的基因表达发生改变的重组微生物文库；
(b)培养所述构建的重组微生物文库，测量所述重组微生物文库的培养上清液的吸光度，培养所述重组微生物然后将所述基因调节文库引入，并比较所述重组微生物文库的培养上清液的颜色与在将所述基因调节文库引入之前所述重组微生物的培养上清液的颜色；并且
(c)当所述重组微生物文库的培养上清液显示出比在将所述基因调节文库引入之前的重组微生物的培养上清液更深的红色时，选择引入所述重组微生物文库中的基因作为参与增加丙二酰-CoA产生的基因。
9.权利要求8的方法，其中步骤(b)中颜色之间的比较是通过肉眼进行。
10.权利要求8的方法，其中步骤(b)中的颜色之间的比较是通过测量吸光度来进行，并且步骤(c)包括当所述重组微生物文库的培养上清液的吸光度高于在将所述基因调节文库引入之前所述重组微生物的培养上清液的吸光度时，选择在所述重组微生物文库中引入的基因。
11.权利要求8的方法，其中所述基因调节文库是选自以下的文库：sRNA文库、基因组文库、cDNA文库、gRNA文库和用于构建敲除或突变菌株的寡核苷酸文库。
12.一种产生具有增加的丙二酰-CoA产生能力的重组微生物的方法，所述方法包括调节通过权利要求8至11中任一项的方法筛选的基因在具有丙二酰-CoA产生能力的微生物中的表达。
13.权利要求12的方法，其中所述筛选的基因是选自以下项的一种或多种基因：
fabF(3-氧代酰基-[酰基-载体-蛋白质]合酶II)；
yfcY(β-酮酰-CoA硫解酶)；
xapR(xapAB的转录激活因子)；
cytR(deo操纵子、udp、cdd、tsx、nupC和nupG的转录阻遏物)；
fabH(3-氧代酰基-[酰基-载体-蛋白质]合酶III)；
mqo(苹果酸脱氢酶)；
yfiD(丙酮酸甲酸裂解酶亚基)；
fmt(10-甲酰基四氢叶酸：L-甲硫氨酰基-tRNA(fMet)N-甲酰基转移酶)；
pyrF(乳清酸核苷-5'-磷酸脱羧酶)；
araA(L-阿拉伯糖异构酶)；
fadR(fad调节子的负调节因子和fabA的正调节因子)；
pabA(氨基脱氧分支酸合酶，亚基II)；
purB(腺苷酸琥珀酸裂解酶)；和
hycI(参与加工HycE的C末端的蛋白酶)，
并且减少所述基因的表达。
14.权利要求12的方法，其中所述微生物是选自以下项的微生物：大肠杆菌、根瘤菌属、双歧杆菌属、红球菌属、假丝酵母属、欧文氏菌属、肠杆菌属、巴氏杆菌属、曼氏杆菌属、放线杆菌属、凝聚杆菌属、黄单胞菌属、弧菌属、假单胞菌属、固氮菌属、不动杆菌属、劳尔氏菌属、土壤杆菌属、红杆菌属、发酵单胞菌属、芽孢杆菌属、葡萄球菌属、乳球菌属、链球菌属、乳杆菌属、梭菌属、棒杆菌属、链霉菌属、双歧杆菌属、蓝细菌和圆杆菌属。
15.一种具有增加的丙二酰-CoA产生能力的重组微生物，其中选自以下基因的一种或多种基因：
fabF(3-氧代酰基-[酰基-载体-蛋白质]合酶II)；
yfcY(β-酮酰-CoA硫解酶)；
xapR(xapAB的转录激活因子)；
cytR(deo操纵子、udp、cdd、tsx、nupC和nupG的转录阻遏物)；
fabH(3-氧代酰基-[酰基-载体-蛋白质]合酶III)；
mqo(苹果酸脱氢酶)；
yfiD(丙酮酸甲酸裂解酶亚基)；
fmt(10-甲酰基四氢叶酸：L-甲硫氨酰基-tRNA(fMet)N-甲酰基转移酶)；
pyrF(乳清酸核苷-5'-磷酸脱羧酶)；
araA(L-阿拉伯糖异构酶)；
fadR(fad调节子的负调节因子和fabA的正调节因子)；
pabA(氨基脱氧分支酸合酶，亚基II)；
purB(腺苷酸琥珀酸裂解酶)；和
hycI(参与加工HycE的C末端的蛋白酶)在具有丙二酰-CoA产生能力的微生物中的表达与在野生型微生物中的表达相比降低。
16.一种使用丙二酰-CoA作为底物或中间体产生有用物质的方法，所述方法包括以下步骤：
(a)构建重组微生物，其中在权利要求15的重组微生物中另外引入参与所述有用物质产生的基因或增加所述基因的表达，或者另外使参与所述有用物质产生的基因缺失或减少所述基因的表达；
(b)培养所构建的微生物；并且
(c)从所培养的微生物回收所述有用物质。
17.权利要求16的方法，其中所述有用物质是选自以下项的一种或多种：
放线菌紫素、多柔比星、道诺霉素、土霉素、雷帕霉素、洛伐他汀、SEK4、SEK4b、SEK34、SEK15、SEK26、FK506、DMAC、阿克拉菌酮、阿克拉酸、ε-紫红霉酮、芦荟苦素、芦荟宁、芦荟苷、5,7-二羟基-2-甲基色酮、红霉素、利福霉素、阿维菌素、格尔德霉素、伊维菌素、多西环素、安曲霉素、青霉酸、卡奇霉素、埃博霉素、特曲霉素、富伦菌素、三乙酸内酯、6-甲基水杨酸或芦荟酮的聚酮化合物；
松属素、二氢山柰酚、圣草酚、二氢槲皮素、黄豆苷元、染料木素、芹菜素、木犀草素、山柰酚、槲皮素、儿茶素、天竺葵色素、矢车菊素、阿福豆素、杨梅素、非瑟酮、高良姜素、橙皮素、柑桔黄酮、花翠素、表儿茶素、白杨素、白藜芦醇或柚皮素的苯丙素类化合物；
戊烷、己烷、庚烷、辛烷、壬烷、癸烷、十一烷、十二烷、十三烷、十四烷、十五烷、十六烷、十七烷、十八烷、十九烷、二十烷、戊醇、己醇、庚醇、辛醇、壬醇、癸醇、十一烷醇、十二烷醇、十三烷醇、十四烷醇、十五烷醇、十六烷醇、十七烷醇、十八烷醇、十九烷醇、二十烷醇、癸酸甲酯、月桂酸甲酯、肉豆蔻酸甲酯、棕榈酸甲酯、棕榈油酸甲酯、硬脂酸甲酯、油酸甲酯、亚油酸甲酯、亚麻酸甲酯、花生酸甲酯、二十烯酸甲酯、芥酸甲酯、癸酸乙酯、月桂酸乙酯、肉豆蔻酸乙酯、棕榈酸乙酯、棕榈油酸乙酯、硬脂酸乙酯、油酸乙酯、亚油酸乙酯、亚麻酸乙酯、花生酸乙酯、二十烯酸乙酯或芥酸甲酯的生物燃料组；
神经酰胺、棕榈酸酯或鞘氨醇的脂质化合物；和
3-羟基丙酸。
18.一种用于产生6-甲基水杨酸的重组微生物，其中在权利要求15的重组微生物中另外引入了编码6-甲基水杨酸合酶(6MSAS)和4'-磷酸泛酰巯基乙胺基转移酶(Sfp)的基因，或者增加所述基因的表达。
19.权利要求18的重组微生物，其中所敲低的基因是选自pabA、fabF、xapR和ytcY的一种或多种。
20.权利要求18的重组微生物，其中在所述重组微生物中另外引入了编码选自葡萄糖-6-磷酸脱氢酶(Zwf)、苹果酸脱氢酶(Mdh)、磷酸甘油酸脱氢酶(SerA)、乙酰-CoA羧化酶(AccBC和AccD1)、甘油醛3-磷酸脱氢酶(GapA)、磷酸甘油酸激酶(Pgk)、乙酰-CoA合成酶(Acs)和丙酮酸脱氢酶(AceEF和Lpd)的一种或多种酶的基因或增加所述基因的表达。
21.一种产生6-甲基水杨酸的方法，所述方法包括以下步骤：
(a)培养权利要求18的重组微生物；并且
(b)从所培养的微生物回收所述6-甲基水杨酸。
22.一种用于产生芦荟酮的重组微生物，其中在权利要求15的重组微生物中另外引入了编码芦荟酮合酶的基因或增加所述基因的表达。
23.权利要求22的重组微生物，其中所敲低的基因是pabA。
24.权利要求22的重组微生物，其中在所述重组微生物中另外引入了编码选自葡萄糖-6-磷酸脱氢酶(Zwf)、苹果酸脱氢酶(Mdh)、磷酸甘油酸脱氢酶(SerA)、乙酰-CoA羧化酶(AccBC和AccD1)、甘油醛3-磷酸脱氢酶(GapA)、磷酸甘油酸激酶(Pgk)、乙酰-CoA合成酶(Acs)和丙酮酸脱氢酶(AceEF和Lpd)的一种或多种酶的基因或增加所述基因的表达。
25.一种产生芦荟酮的方法，所述方法包括以下步骤：
(a)培养权利要求22的重组微生物；并且
(b)从所培养的微生物回收所述芦荟酮。
26.一种用于产生白藜芦醇的重组微生物，其中在权利要求15的重组微生物中另外引入了编码酪氨酸氨裂解酶(TAL)、4-香豆酸:CoA连接酶(4CL)和芪合酶(STS)的基因或者增加所述基因的表达。
27.权利要求26的重组微生物，其中所述4-香豆酸:CoA连接酶是其中SEQ ID NO:128所示的氨基酸序列中的氨基酸突变为I250L/N404K/I461V的突变酶，或者是其中SEQ ID NO:131所示的氨基酸序列中的氨基酸突变为A294G/A318G的突变酶。
28.权利要求26的重组微生物，其中所敲低的基因是选自pabA、yfiD、mqo、xapR、purB、fabH、fabF、yfcY、fadR、cytR、fmt和pyrF中的一种或多种。
29.一种产生白藜芦醇的方法，所述方法包括以下步骤：
(a)培养权利要求26的重组微生物；并且
(b)从所培养的微生物回收所述白藜芦醇。
30.一种用于产生柚皮素的重组微生物，其中在权利要求15的重组微生物中另外引入了编码酪氨酸氨裂解酶(TAL)、4-香豆酸:CoA连接酶(4CL)、查耳酮合酶(CHS)和查耳酮异构酶(CHI)的基因或者增加所述基因的表达。
31.权利要求30的重组微生物，其中所述4-香豆酸:CoA连接酶是其中SEQ ID NO:128所示的氨基酸序列中的氨基酸突变为I250L/N404K/I461V的突变酶。
32.权利要求30的重组微生物，其中所敲低的基因是选自fadR、hycI和xapR中的一种或多种。
33.一种产生柚皮素的方法，所述方法包括以下步骤：
(a)培养权利要求30的重组微生物；并且
(b)从所培养的微生物回收所述柚皮素。
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| CN110343639A (zh) * | 2019-07-23 | 2019-10-18 | 中国医药集团总公司四川抗菌素工业研究所 | 一株产15(s)-o-乙基雷帕霉素的链霉菌 |
| CN111979209A (zh) * | 2020-08-24 | 2020-11-24 | 中国科学院大学 | 一种聚八酮合酶及其编码基因与应用 |
| CN112592940A (zh) * | 2020-12-16 | 2021-04-02 | 江西邦泰绿色生物合成生态产业园发展有限公司 | 一种n-笏甲氧羰基-葵烷基乙醛的生物酶合成方法 |
| CN113527245A (zh) * | 2021-06-28 | 2021-10-22 | 海南医学院 | 一种芦荟松的合成方法及其用途 |
| CN114574458A (zh) * | 2022-02-28 | 2022-06-03 | 江南大学 | 可以提高柚皮素产量的查尔酮合成酶突变体 |
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| CN113430215B (zh) * | 2021-06-03 | 2023-04-18 | 昆明理工大学 | 乙酰CoA合成酶基因RKACS1及其应用 |
| CN115433685B (zh) * | 2022-06-08 | 2023-03-03 | 安徽大学 | 一种通过改造糖多孢红霉菌sace_5812基因途径提高红霉素产量的方法 |
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| US20210277428A1 (en) | 2021-09-09 |
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| JP2020530757A (ja) | 2020-10-29 |
| EP3608417A4 (en) | 2020-12-09 |
| WO2019235688A1 (ko) | 2019-12-12 |
| KR20190139727A (ko) | 2019-12-18 |
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