CN116457454A - 转化体以及使用其的类胡萝卜素组合物的制造方法 - Google Patents
转化体以及使用其的类胡萝卜素组合物的制造方法 Download PDFInfo
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- C12N5/10—Cells modified by introduction of foreign genetic material
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Abstract
本发明提供一种通过基因重组技术来合成类胡萝卜素组合物的技术。一种转化体,其在宿主细胞中导入有:第一启动子、及与所述第一启动子可操作地连接的包含crtY和crtZ的类胡萝卜素的生物合成基因的上游基因;以及启动子强度比所述第一启动子高的第二启动子、及与所述第二启动子可操作地连接的ZEP基因和CCS基因。
Description
技术领域
本发明涉及一种转化体以及使用其的类胡萝卜素组合物的制造方法。
背景技术
类胡萝卜素是植物、藻类、细菌等生产的天然色素,是在生物中担任多种生理功能的重要的化合物组。迄今为止,许多类胡萝卜素被分离,对于在自然界中仅微量存在的类胡萝卜素,其健康功能也开始受到关注,但由于是微量的,因此商业上的制造成本高而困难。
因此,近年来,用于利用微生物廉价地生产类胡萝卜素的生物技术研究盛行起来。植物中的类胡萝卜素生物合成途径和生物合成基因到2000年为止大致明确。已经进行了大量如下的尝试:从上游鉴定参与类胡萝卜素的生物合成的基因,制作将被鉴定的基因导入微生物的转化体,利用该转化体生产类胡萝卜素。
作为产生类胡萝卜素的宿主,最被研究的微生物是大肠杆菌。作为包含类胡萝卜素的类异戊二烯的基本代谢途径,大肠杆菌通过非甲羟戊酸途径(MEP途径),合成异戊烯基二磷酸(IPP)和作为其异构体的二甲基烯丙基二磷酸(DMAPP)(非专利文献1)。合成的IPP和DMAPP通过经由异戊二烯基转移酶的连续缩合反应,被代谢为香叶基二磷酸(GPP)和法尼基二磷酸(FPP)。
大肠杆菌为类胡萝卜素非生产菌,不具有FPP以后的类胡萝卜素合成所需的酶基因,但通过根据产生目的的类胡萝卜素从上游引入香叶基香叶基焦磷酸(GGPP)合成酶基因(crtE)、八氢番茄红素合成酶基因(crtB)、合成番茄红素的八氢番茄红素不饱和化酶基因(crtI)、合成β胡萝卜素的番茄红素环化酶基因(crtY)、合成β-隐黄质和玉米黄质的β胡萝卜素羟化酶基因(crtZ)这样的类胡萝卜素生物合成基因,能够合成各种类胡萝卜素。
另一方面,即使是判明了能够这样产生的类胡萝卜素,仅导入类胡萝卜素的生物合成基因群时,实际的产生量也大多与实用生产水平相差甚远。这被认为是因为大肠杆菌只能少量产生成为类胡萝卜素合成的前体的FPP。因此,以大量生产类胡萝卜素为目标,研究了以FPP增产为目的的途径工程。
作为途径工程的途径,大致分为改变包含MEP途径的内在代谢途径的方法和导入来自异种生物的甲羟戊酸(MVA)途径基因群的方法。
作为前者的途径的代表例,可举出在高度表达了来自酿酒酵母(S.cerevisiae)或绿藻(雨生红球藻(Haematococcus pluvialis))的1型IPP异构酶基因(idi)的大肠杆菌中,显著增加类胡萝卜素生产量的方法(非专利文献2),例如,报告了导入有idi和来自菠萝泛菌(Pantoea ananatis)的crtE、crtB、crtI、crtY以及crtZ的大肠杆菌能够占优势地产生玉米黄质(非专利文献3)。
作为后者的途径的代表例,发现利用使来自放线菌Streptomyces sp.CL190株的MVA途径基因群(包含2型IPP异构酶基因)进行了功能表达的大肠杆菌,以氘代D-MVA内酯为底物,合成氘代玉米黄质(非专利文献4)。该途径被认为是在用于以大肠杆菌为宿主的类胡萝卜素生产的途径工程研究中,强力且有效的手段(非专利文献5)。
另一方面,判明了玉米黄质环氧化酶(ZEP)基因从玉米黄质合成花药黄质、以及辣椒红色素-辣椒玉红素合成酶(CCS)基因从花药黄质分别合成辣椒红色素(非专利文献6)。然而,虽然这些基因已判明很久,但是没有报告与其他的类胡萝卜素生物合成基因群一起导入宿主,合成辣椒红色素。
现有技术文献
非专利文献
非专利文献1:Rohmer,M.et al.:Biochem.J.,295,517(1993)
非专利文献2:Kajiwara,S.et al.:Biochem.J.,324,421(1997)
非专利文献3:Takemura,M.et al.:Tetrahedron Letters 56,6063(2015).
非专利文献4:Kakinuma,K.et al.:J.Am.Chem.Soc.,123,1238(2001).
非专利文献5:生物工学会志,第93卷,第7号,397(2015).
非专利文献6:Bouvier,F.et al.The Plant Journal,6(1),45(1994)
发明内容
发明所要解决的课题
辣椒红色素与番茄红素、胡萝卜素、隐黄质、玉米黄质不同,是不能通过使用迄今为止已知的重组微生物的方法来合成的。事实上,本发明人即使将CrtE基因、CrtB基因、CrtI基因、CrtY基因、CrtZ基因、ZEP基因、以及CCS基因与IDI基因一起导入到大肠杆菌中,也完全没有生成辣椒红色素。
本发明的目的在于,提供通过基因重组技术来制造包含如辣椒红色素那样迄今为止未能合成的类胡萝卜素的组合物的技术。
用于解决课题的方案
本发明人想到以仅使辣椒红色素的生物合成基因群中的一部分基因的表达量相对变多的方式进行控制的崭新的构想。因此,以相对多地获得辣椒红色素的底物为目的,以使辣椒红色素的生物合成基因群中上游的基因的表达量相对变多的方式构建重组载体,将其进行转化来尝试辣椒红色素的合成。但是,在该方法中,即使引入idi,也完全没有生成辣椒红色素。
因此,尝试转换构想为以使辣椒红色素的生物合成基因群中下游的基因的表达量相对多的方式构建重组载体时,意外地发现了该转化体能够生成辣椒红色素。进而,还没有预期地发现了在将ZEP基因和CCS基因以表达为ZEP和CCS经由特定的亲和蛋白的多聚体间接地结合的方式引入到载体中重组载体中的情况下,还能够生成辣椒玉红素、类胡萝卜黄色素A(Cucurbitaxanthin A)等其他多种类胡萝卜素。本发明是基于该见解并进一步重复进行研究从而完成的发明。
即,本发明提供下述揭示的方式的发明。
项1.一种转化体,其在宿主细胞中导入有:
第一启动子、及与所述第一启动子可操作地连接的类胡萝卜素的生物合成基因的上游基因;以及
启动子强度比所述第一启动子高的第二启动子、及与所述第二启动子可操作地连接的ZEP基因和CCS基因,
上述上游基因包含crtY和crtZ。
项2.根据项1所述的转化体,其中,所述第一启动子与所述第二启动子的启动子强度之差为类胡萝卜素的生物合成基因固有的启动子与PBAD的启动子强度之差以上。
项3.根据项1或2所述的转化体,其中,所述上游基因还包含crtI;crtI和crtB;或crtI、crtB和crtE。
项4.根据项1~3中任一项所述的转化体,其中,所述第一启动子是Plac或类胡萝卜素的生物合成基因固有的启动子。
项5.根据项1~4中任一项所述的转化体,其中,所述第二启动子是PBAD或Ptac。
项6.根据项1~5中任一项所述的转化体,其中,进一步在所述宿主细胞中导入有S蛋白基因、S标签基因(tag gene)和接头基因(linker gene),
所述ZEP基因和所述CCS基因分别与所述S标签基因连接,
所述接头基因连接在所述ZEP基因与所述S标签基因之间和/或所述CCS基因与所述S标签基因之间。
项7.根据项6所述的转化体,其中,所述S标签基因编码的S标签为以下的(1)和(2)中的任一种多肽,
(1)包含编码序列号4所示的氨基酸序列的碱基序列的多肽,
(2)相对于序列号4所示的氨基酸序列的第12位以外的序列一致性为90%以上、且与S蛋白特异性结合的多肽。
项8.根据项1~7中任一项所述的转化体,其中,所述接头基因至少连接在所述ZEP基因与所述S标签基因之间。
项9.根据项1~8中任一项所述的转化体,其中,所述接头基因连接在所述ZEP基因与所述S标签基因之间,而不在所述CCS基因与所述S标签基因之间。
项10.一种类胡萝卜素组合物的制造方法,其包括以下工序:培养项1~9中任一项所述的转化体。
项11.根据项10所述的制造方法,其中,所述类胡萝卜素组合物包含辣椒红色素(capsanthin)。
项12.根据项10或11所述的制造方法,其中,所述类胡萝卜素组合物包含辣椒玉红素(capsorubin)、辣椒红色素3′-乙酸酯、辣椒玉红素3-乙酸酯、类胡萝卜黄色素A(cucurbitaxanthin A)、辣椒玉红素二乙酸酯、辣椒红色素3,6-环氧化物、和/或辣椒红色素3,6-环氧化物3′-乙酸酯。
发明效果
根据本发明,能够通过基因重组技术来制造包含如辣椒红色素那样,迄今为止未能合成的类胡萝卜素的组合物。作为得到这样的效果的机理尚不清楚,但如下进行考察。关于CCS,不仅发现了从花药黄质合成辣椒红色素的活性,还发现了从在类胡萝卜素生物合成途径的上游的番茄红素合成β胡萝卜素的副活性。因此,认为如果使上游的底物增多,则CCS的功能大多被利用于从番茄红素合成β胡萝卜素的反应的催化剂,未发挥显示合成辣椒红色素的活性一方的功能。可以认为,通过相对地降低玉米黄质及其上游的基因的表达量,提高与玉米黄质合成相关的基因的下游的ZEP基因和CCS基因的表达量,能够抑制玉米黄质及其上游的中间体的过度蓄积,使迄今为止无法以显示合成辣椒红色素的活性的方式发挥功能的CCS首次发挥功能。
附图说明
图1表示类胡萝卜素生物合成途径以及介导该途径的酶组。
图2是比较例1中制作的重组载体的示意图。
图3是比较例2(比较例2-1和比较例2-2)中制作的重组载体的示意图。
图4是比较例2(比较例2-1和比较例2-2)中得到的类胡萝卜素组合物的HPLC色谱图。
图5是实施例1中制作的重组载体的示意图。
图6是实施例1中得到的类胡萝卜素组合物的HPLC色谱图。
图7表示实施例1中得到的类胡萝卜素组合物的UPLC色谱图(a)、辣椒红色素的吸收光谱图(b)、以及辣椒红色素的质谱图(c)。
图8是实施例2(实施例2-1和实施例2-2)中制作的重组载体的示意图。
图9表示实施例2(实施例2-1和实施例2-2)中得到的类胡萝卜素组合物的HPLC色谱图。
图10表示实施例2(实施例2-1和实施例2-2)中得到的类胡萝卜素组合物的辣椒红色素的产生量。
图11是实施例3(实施例3-1~实施例3-3)中制作的重组载体的示意图。
图12示意性地表示通过基于实施例3(实施例3-1~实施例3-3)的表达系统而构建的、使用了S蛋白的CCS和ZEP的协同体系。
图13表示实施例3(实施例3-1~实施例3-3)中得到的类胡萝卜素组合物的分析结果。
图14表示实施例3(实施例3-2)中得到的类胡萝卜素组合物的分析结果。
具体实施方式
1.转化体
本发明的转化体的特征在于,在宿主细胞中导入有:第一启动子、及与所述第一启动子可操作地连接的类胡萝卜素的生物合成基因的上游基因;以及启动子强度比所述第一启动子高的第二启动子、及与所述第二启动子可操作地连接的ZEP基因和CCS基因,上述上游基因包含crtY和crtZ。由此,在宿主细胞中表达类胡萝卜素的生物合成酶,能够生成类胡萝卜素组合物。
1-1.类胡萝卜素生物合成酶以及编码其的基因(类胡萝卜素生物合成基因)
图1表示类胡萝卜素生物合成途径以及介导该途径的酶组。应予说明,在图1中,举出了使用重组大肠杆菌生物合成类胡萝卜素的例子。
CrtE是从法尼基二磷酸(FPP)合成香叶基香叶基焦磷酸(GGPP)的GGPP合成酶,以下,将编码CrtE的基因记载为“crtE”。
CrtB是从GGPP合成八氢番茄红素的八氢番茄红素合成酶,以下,将编码CrtB的基因记载为“CrtB”。
CrtI是从八氢番茄红素合成番茄红素的八氢番茄红素不饱和化酶,以下,将编码CrtI的基因记载为“crtI”。
CrtY是从番茄红素合成β胡萝卜素的番茄红素环化酶,以下,将编码CrtY的基因记载为“crtY”。
CrtZ是从β胡萝卜素合成β-隐黄质和玉米黄质的β胡萝卜素羟化酶,以下,将编码CrtZ的基因记载为“crtZ”。
ZEP是从玉米黄质合成花药黄质和紫黄质的玉米黄质环氧化酶。
CCS是已知从花药黄质和紫黄质分别合成辣椒红色素和辣椒玉红素的辣椒红色素/辣椒玉红素合成酶。进而,在由本发明人发现的见解中,CCS也是从花药黄质合成辣椒红色素3'-乙酸酯、类胡萝卜黄色素A,从紫黄质合成辣椒玉红素3-乙酸酯、辣椒玉红素二乙酸酯、辣椒红色素3,6-环氧化物、辣椒红色素3,6-环氧化物3'-乙酸酯的合成酶。
上述的编码类胡萝卜素生物合成酶的基因是公知的基因,其具体的序列可以从官方数据库等中适当取得。
作为成为这些类胡萝卜素生物合成基因的来源的生物,没有特别限定,从产生与这些基因对应的类胡萝卜素的生物中没有特别限制地选择。作为具体的来源生物,不论植物和微生物,具体而言,可举出辣椒(Capsicum)属(例如辣椒(Capsicum annuum))、百合(Lilium)属(例如卷丹百合(Lilium lancifolium))、拟南芥(Arabidopsis)属(例如拟南芥(Arabidopsis thaliana))、玉米黍(Zea)属(例如玉米(Zea Mays))、李(Prunus)属(例如杏(Prunus armeniaca))、蕃薯(Ipomoea)属(例如牵牛(Ipomoea nil))、龙胆(Gentiana)属(例如黄龙胆(Gentiana lutea))、地钱(Marchantia)属(例如地钱(MarchantiaPolymorpha))、泛生菌(Pantoea)属(菠萝泛菌(Pantoea ananas))、农杆菌(Agrobacterium)属(例如Agrobacterium aurantiacum)、红球藻(Heamatococcus)属、单针藻(Monoraphidium)属、法夫酵母(Phaffia)属(Xanthophyllomyces属)、副球菌(Paracoccus)属、短波单胞菌(Brevundimonas)属、戈登氏菌(Gordonia)属(红球菌(Rhodococcus)属)、念珠藻(Nostoc)属、短波单胞菌(Brevundimonas)属、以及粘杆菌(Gloeobacter)属等,优选可举出辣椒(Capsicum)属、百合(Lilium)属、泛生菌(Pantoea)属。另外,不论成为各基因的来源的生物的异同,但优选举出相同的生物。
在本发明中能够使用的类胡萝卜素的生物合成基因中,不仅包含由能够从官方数据库取得的碱基序列及其密码子优化序列构成的基因,只要是编码具有类胡萝卜素的生物合成酶活性的酶的基因,则也包含由与该序列类似(具体而言,与相对于下述(I-1)、(II-1)、(III-1)的碱基序列的(I-2)、(II-2)、(III-2)和(I-3)、(II-3)、(III-3)的碱基序列的类似性相当的程度地类似)的碱基序列构成的基因。
作为上述类胡萝卜素的生物合成基因的碱基序列的优选例,可举出以下。
(I-1)DDBJ登录号D90087所示的crtE、crtY、crtI、crtB和crtZ基因的碱基序列,
(II-1)DDBJ登录号XM_016705616所示的ZEP基因的碱基序列,
(III-1)DDBJ登录号X76165所示的CCS基因的碱基序列,
(I-2)一种核酸的碱基序列,其为分别编码具有从法尼基二磷酸(FPP)合成香叶基香叶基焦磷酸(GGPP)的GGPP合成酶活性的酶、具有从GGPP合成八氢番茄红素的八氢番茄红素合成酶活性的酶、具有从八氢番茄红素合成番茄红素的八氢番茄红素不饱和化酶活性的酶、具有从番茄红素合成β胡萝卜素的番茄红素环化酶活性的酶、以及具有从β胡萝卜素合成β-隐黄质和玉米黄质的β胡萝卜素羟化酶活性的酶的基因,所述碱基序列与分别由与上述(I-1)所示的碱基序列互补的碱基序列构成的核酸在严格条件下进行杂交,
(II-2)一种核酸的碱基序列,其为编码具有从玉米黄质合成花药黄质和紫黄质的玉米黄质环氧化酶活性的酶的基因,所述碱基序列与由与上述(II-1)所示的碱基序列互补的碱基序列构成的核酸在严格条件下进行杂交,
(III-2)一种核酸的碱基序列,其为编码从花药黄质和紫黄质分别合成辣椒红色素和辣椒玉红素,此外,合成辣椒玉红素、辣椒红色素3′-乙酸酯、辣椒玉红素3-乙酸酯、类胡萝卜黄色素A、辣椒玉红素二乙酸酯、辣椒红色素3,6-环氧化物、辣椒红色素3,6-环氧化物3′-乙酸酯的酶活性的基因,所述碱基序列与由与上述(III-1)所示的碱基序列互补的碱基序列构成的核酸在严格条件下进行杂交,
(I-3)一种碱基序列,其为分别编码具有从法尼基二磷酸(FPP)合成香叶基香叶基焦磷酸(GGPP)的GGPP合成酶活性的酶、具有从GGPP合成八氢番茄红素的八氢番茄红素合成酶活性的酶、具有从八氢番茄红素合成番茄红素的八氢番茄红素不饱和化酶活性的酶、具有从番茄红素合成β胡萝卜素的番茄红素环化酶活性的酶、以及具有从β胡萝卜素合成β-隐黄质和玉米黄质的β胡萝卜素羟化酶活性的酶的基因,所述碱基序列和与上述(I-1)所示的碱基序列互补的碱基序列分别具有90%以上的同源性,
(II-3)一种碱基序列,其为编码具有从玉米黄质合成花药黄质和紫黄质的玉米黄质环氧化酶活性的酶的基因,所述碱基序列与上述(II-1)所示的碱基序列具有90%以上的同源性,
(III-3)一种碱基序列,其为编码从花药黄质和紫黄质分别合成辣椒红色素和辣椒玉红素,此外,合成辣椒玉红素、辣椒红色素3′-乙酸酯、辣椒玉红素3-乙酸酯、类胡萝卜黄色素A、辣椒玉红素二乙酸酯、辣椒红色素3,6-环氧化物、辣椒红色素3,6-环氧化物3′-乙酸酯的酶活性的基因,所述碱基序列与上述(III-1)所示的碱基序列具有90%以上的同源性。
在上述(I-2)、(II-2)、(III-2)中,“严格条件”是指在包含0.5%SDS、5×Denhartz’s〔Denhartz’s,0.1%牛血清白蛋白(BSA)、0.1%聚乙烯吡咯烷酮、0.1%聚蔗糖(Ficoll)400〕和100μg/ml鲑鱼精子DNA的6×SSC(1×SSC为0.15M NaCl、0.015M柠檬酸钠、pH7.0)中在50℃~65℃下保温4小时~一晚的条件。在严格条件下的杂交,具体而言,通过以下方法进行。即,制作使DNA文库或cDNA文库固定化的尼龙膜,在包含6×SSC、0.5%SDS、5×Denhartz’s、100μg/ml鲑鱼精子DNA的预杂交溶液中,在65℃下封闭尼龙膜。然后,加入用32P标记的各探针,在65℃下保温一夜。将该尼龙膜在6×SSC中、室温下清洗10分钟,在包含0.1%SDS的2×SSC中、室温下清洗10分钟,在包含0.1%SDS的0.2×SSC中、在45℃下清洗30分钟后,采用放射自显影术,能够检测出与探针特异性杂交的核酸。
在上述(I-3)、(II-3)、(III-3)中,上述同源性只要为90%以上即可,但可举出优选为95%以上,更优选为98%以上,进一步优选为98.5%以上,进一步优选为99%以上,更进一步优选为99.5%以上,特别优选为99.9%以上。“同源性”是使用具有将基准序列作为参考序列进行比较的算法的公开或市售的软件来计算的。具体而言,可以使用BLAST、FASTA、或GENETYX(GENETYX CORPORATION制造)等,将它们设定为默认参数使用即可。
进而,上述的类胡萝卜素的生物合成基因可以将全长序列引入重组载体中,也可以在除去了宿主中的目标酶的表达中不需要的序列的状态下引入到重组载体中。例如,在成为类胡萝卜素的生物合成基因的来源的生物为植物、宿主为非植物细胞的情况下,从更进一步提高辣椒红色素的产生量的观点出发,优选在除去信号序列的状态下引入到重组载体中。来自各植物的基因的信号序列也可以通过使用官方数据库或各种基因分析工具而容易地取得。
以下,在类胡萝卜素的生物合成基因群中,将crtE、crtB、crtI、crtY以及crtZ分类为“上游基因”,将ZEP基因和CCS基因分类为“下游基因”。
类胡萝卜素的生物合成基因的上游基因与第一启动子可操作地连接,类胡萝卜素的生物合成基因的下游基因与第二启动子可操作地连接。即,第一启动子是进行上游基因的转录控制的区域,第二启动子是进行下游基因的转录控制的区域。另外,可操作地连接是指,控制因子和上述基因以能够在宿主细胞中操作的状态连接。
1-2.启动子
关于第一启动子和第二启动子的关系,第二启动子被设计为启动子强度高于第一启动子。例如,第二启动子的启动子强度优选为第一启动子的启动子强度的3倍以上。启动子强度是指转录开始的频率,例如通过Pbla单位(Pbla unit;Deuschle et al.,EMBO J.,5:2987-2994(1986))、Miller单位(Miller units;也称为β半乳糖苷酶单位;Pbla单位的5000倍(Lanzer et al.,PNAS USA,85,8973-8977(1988))、RPU(Relative promoterunits;Meyer et al.,Nature Chemical Biology 15,196-204(2019))等测定。
在本发明中,利用第一启动子和第二启动子,以类胡萝卜素的生物合成基因的上游基因的表达量相对降低、下游基因的表达量相对变高的方式控制,由此能够合成类胡萝卜素。
从进一步提高类胡萝卜素的合成量的观点出发,第一启动子和第二启动子的启动子强度之差优选为类胡萝卜素的生物合成基因固有的启动子(优选为在菠萝泛菌(Pantoeaananatis)中生来具备的启动子,更优选为包含在DDBJ登录号D90087中的启动子)与PBAD启动子的启动子强度之差以上,更优选为类胡萝卜素的生物合成基因固有(优选为在菠萝泛菌中生来具备的启动子,更优选为包含在DDBJ登录号D90087中的启动子)的启动子与tac启动子的启动子强度之差以上。
作为第一启动子和第二启动子,只要将设计为启动子强度在第二启动子一方相对高作为限度,就没有特别限定,可以根据导入有引入了这些启动子的重组载体的宿主的种类适当选择。第一启动子和第二启动子可以是天然启动子,也可以是人工启动子。
作为用作第一启动子和第二启动子的启动子的具体例,在大肠杆菌等原核生物中,可举出lac启动子(Plac)、类胡萝卜素的生物合成基因固有的启动子、BAD启动子(PBAD)、tac启动子(Ptac)、trc启动子、T7启动子、T7lac启动子、T5启动子、UV5lac启动子、L8-UV5lac启动子、cspA启动子、CAG启动子、CMV启动子、RSV启动子等,在酵母中,可举出GAL1启动子、NMT1启动子、TEF1第一启动子、ADH1启动子、TPI1启动子、HXT7启动子、TDH3启动子、PGK1启动子、PYK1启动子、CAG启动子、CMV启动子、RSV启动子等。
应予说明,类胡萝卜素的生物合成基因固有的启动子是在类胡萝卜素的生物合成中生来具备的启动子。作为类胡萝卜素的生物合成基因固有的启动子,可以没有特别限制地使用,在成为上述类胡萝卜素生物合成基因的来源的生物中,可以使用作为该类胡萝卜素生物合成基因的启动子而生来具备的启动子。作为一例,在类胡萝卜素的生物合成基因固有的启动子的控制下配置的基因来自菠萝泛菌的情况下,作为该类胡萝卜素的生物合成基因固有的启动子,也可以优选使用在菠萝泛菌中生来具备的启动子,更优选使用包含在DDBJ登录号D90087中的启动子。
可以从包含这些例示的各种启动子中选择启动子强度不同的2种,将启动子强度低的一方作为第一启动子,将启动子强度低的一方作为第二启动子而选择。进而,为了进一步提高类胡萝卜素的合成量,可以从包含上述例示的各种启动子中选择启动子强度成为上述的启动子强度之差的2种启动子的组合,将启动子强度低的一方作为第一启动子,启动子强度低的一方作为第二启动子而选择。
进一步优选的是,作为第一启动子,可举出Plac和类胡萝卜素的生物合成基因固有的启动子,作为第二启动子,可举出PBAD和Ptac,进一步优选为Ptac。
另外,通过使用上述那样的启动子强度不同的第一启动子和第二启动子进行的表达量的控制,以抑制基于CCS的从番茄红素合成β胡萝卜素的反应的方式进行。因此,通过第一启动子表达的上游基因中只要至少包含crtY和crtZ即可。
关于其他上游基因,可以与crtY和crtZ相同地配置在第一启动子的控制下,也可以配置在其他启动子的控制下。优选的是,其他上游基因优选配置在第一启动子的控制下。作为其他上游基因配置在第一启动子的控制下的情况的具体例,作为该上游基因,可举出进一步(i)包含crtI的情况;(ii)包含crtI和crtB的情况;以及(iii)包含crtI、crtB和crtE的情况。另外,也可以与其他上游基因一起包含IPP异构酶基因(idi),即,也可以在第一启动子的控制下,进一步配置(iv)crtI、crtB、crtE以及idi。
第一启动子及配置于其控制下的基因、以及第二启动子及配置于其控制下的基因可通过均引入到1个表达载体而构成1个重组载体,也可通过引入到分别的表达载体而分别构成第一重组载体和第二重组载体。
1-3.S用于经由S蛋白的CCS和ZEP的协同体系表达的基因
CCS除了具有辣椒红色素/辣椒玉红素合成活性以外,还具有CrtY所具有的番茄红素β-环化酶活性。通过这样的CCS的双重活性,即使在大肠杆菌体内CCS基因高效率地表达,也存在如下情况:不是与ZEP共同作用,而是与八氢番茄红素脱氢酶CrtI共同作用,由此从CrtI接收番茄红素作为底物而合成β-胡萝卜素的反应优先。因此,为了通过使CCS选择性地与ZEP进行协同反应来进一步提高基于CCS的反应产物的生成量,CCS基因和ZEP基因优选如下述那样以使经由S蛋白的CCS和ZEP的协同体系表达的方式导入宿主细胞。将示意性地说明该协同体系的图示于图12。
在本发明的使经由S蛋白的CCS和ZEP的协同体系表达的方式中,进一步将S蛋白基因、S标签基因以及接头基因导入宿主细胞,分别在该ZEP基因和该CCS基因中连接该S标签基因,该接头基因连接在该ZEP基因与该S标签基因之间和/或该CCS基因与该S标签基因之间。
S标签和S蛋白是彼此特异性的亲和标签与亲和蛋白的关系,是构成一个核糖核酸酶A(RNase A)分子的多肽。用枯草杆菌蛋白酶(Subtilisin)切断核糖核酸酶A而得到的N末端20个氨基酸片段为S标签,由剩余的约100个氨基酸构成的片段为S蛋白。S标签和S蛋白可以非共价地结合而再现原来的高级结构,其结果呈现核糖核酸酶(RNase)活性(Lopez-Alonso,J.P.,Bruix,M.,Font,J.,Ribo,M.,Vilanova,M.,Rico,M.,Gotte,G.,Libonati,M.,Gonzalez,C.and Laurents,D.V.(2006)J.Biol.Chem.281,9400-9406)。
S蛋白和核糖核酸酶A具有如下性质:这些分子的一部分在分子间交换C末端β-链(β-strand),形成如图12所示的均质“二聚体”、均质“三聚体”、三聚体以上的均质“多聚体”那样的多聚体。
另一方面,连接有S标签基因的ZEP基因和CCS基因分别在宿主细胞内作为S标签融合ZEP和S标签融合CCS表达。进而,接头基因连接在ZEP基因与S标签基因之间和/或CCS基因与S标签基因之间,因此在宿主细胞内表达的S标签融合ZEP和S标签融合CCS的任一方或双方采取在S标签与酶之间经由接头的融合蛋白的形态。在图1的示意图中,举出并示出S标签融合ZEP和S标签融合CCS这两者以在S标签与酶之间经由接头的融合蛋白的形态表达(图中“S标签+接头+CCS”以及“S标签+接头+ZEP”)的例子。
这样的S标签融合ZEP和S标签融合CCS与S蛋白共同表达,因此如图12所示,形成融合ZEP和融合CCS经由S标签而间接地结合于S蛋白多聚体的复合体。通过形成这样的复合体,在宿主细胞内ZEP与CCS物理性地接近,因此CCS能够选择性地与ZEP进行协同反应。即,CCS能够顺利地接收基于ZEP的代谢物。由此,能够得到更多的通过CCS代谢生成的类胡萝卜素。更具体而言,能够提高辣椒红色素的生成量、和/或提高辣椒玉红素、辣椒红色素3’-乙酸酯、辣椒玉红素3-乙酸酯、类胡萝卜黄色素A、辣椒玉红素二乙酸酯、辣椒红色素3,6-环氧化物和/或辣椒红色素3,6-环氧化物3’-乙酸酯的生成量。
如上所述,S蛋白已知为核糖核酸酶A的基于枯草杆菌蛋白酶的由约100个氨基酸构成的切断产物,因此其氨基酸序列是公知的(具体而言,可举出序列号2),S蛋白基因的碱基序列也是公知的(具体而言,可举出序列号1)。作为能够在本发明中使用的S蛋白基因的碱基序列,可举出以下的碱基序列。
(IV-1)序列号1所示的碱基序列,
(IV-2)一种核酸的碱基序列,其为编码形成多聚体且与S标签特异性结合的蛋白质的基因,所述核酸与由与上述(IV-1)所示的碱基序列互补的碱基序列构成的核酸在严格条件下杂交,以及
(IV-3)一种碱基序列,其为编码形成多聚体且与S标签特异性结合的蛋白质的基因,所述碱基序列与上述(IV-1)所示的碱基序列具有90%以上的同源性。
关于上述(IV-2)中的“严格条件”,与关于上述(I-2)、(II-2)、(III-2)所述的条件相同。另外,关于上述(IV-3)中的“同源性”及其优选例,与关于上述(I-3)、(II-3)、(III-3)所述的相同。
如上所述,S标签已知为核糖核酸酶A的基于枯草杆菌蛋白酶的N末端的20个氨基酸切断产物,因此其氨基酸序列是公知的(具体而言,可举出序列号4的第12位是代替苯丙氨酸而为组氨酸的氨基酸序列),S蛋白基因的碱基序列也是公知的(具体而言,可举出序列号3的第34~36位是代替编码苯丙氨酸的序列而为编码组氨酸的序列的碱基序列)。
作为能够在本发明中使用的S标签,可举出以下的任一种。
(1)包含编码序列号4所示的氨基酸序列的碱基序列的多肽,
(2)一种多肽,其相对于序列号4所示的氨基酸序列的第12位以外的序列一致性为90%以上,优选为95%以上,且与S蛋白特异性结合。
应予说明,序列号4所示的氨基酸序列以抑制由S标签在宿主细胞内与S蛋白结合引起的对核糖核酸酶活性的影响为目的,第12位由野生型S标签中的组氨酸替换为苯丙氨酸。
在上述(2)的多肽中,相对于序列号4所示的氨基酸序列的序列一致性是与序列号4所示的氨基酸序列进行比较而算出的序列一致性。另外,“序列一致性”表示通过BLASTPACKAGE[sgi32 bit edition,Version2.0.12;available from National Centerfor Biotechnology Information(NCBI)]的bl2seq program(Tatiana A.Tatsusova,Thomas L.Madden,FEMS Microbiol.Lett.,Vol.174,p247-250,1999)得到的氨基酸序列的一致性的值。参数设定为Gap insertion Cost value:11、Gap extension Cost value:1即可。
作为能够在本发明中使用的S标签基因的碱基序列,只要是编码上述S标签的碱基序列,就没有特别限定,但优选举出以下。
(V-1)序列号3所示的碱基序列(即,编码序列号4所示的氨基酸序列的碱基序列),
(V-2)一种碱基序列,其为编码与S蛋白特异性结合的肽的基因,所述碱基序列与上述(V-1)所示的碱基序列(其中,序列号3的第34~36位以外的碱基序列)具有90%以上、优选95%以上的同源性。
关于上述(V-2)中的“同源性”,与关于上述(I-3)、(II-3)、(III-3)所述的相同。
应予说明,S标签基因的上述(V-1)~(V-2)中的序列号3的第34位~第36位以抑制由表达的S标签在宿主细胞内与S蛋白结合引起的对核糖核酸酶活性的影响为目的,由编码野生型S标签中存在的组氨酸的序列替换为编码苯丙氨酸的序列。
另外,作为S标签基因的结合位置,没有特别限定,但从减少对与S标签融合的酶的活性的影响的观点出发,可以设计为使S标签与酶(ZEP和/或CCS)的N末端结合。
作为接头基因,只要是通过在融合蛋白中使接头(肽)在S标签与酶之间保持适度的距离,从而在复合体中能够确保在CCS与ZEP之间的自由度的基因即可,没有特别限定。作为融合蛋白中的接头(肽)的优选的序列,从排除接头(肽)自身的极性的偏差而抑制不期望的吸附等的观点出发,优选为非极性氨基酸与极性氨基酸交替连接的序列,作为该非极性氨基酸和极性氨基酸的组合,优选举出甘氨酸和丝氨酸的组合。作为接头(肽)的长度,例如可举出15~25个氨基酸、优选为18~22个氨基酸。
如上所述,接头基因的连接位置只要是ZEP基因与S标签基因之间以及CCS基因与S标签基因之间的至少任一者即可,但优选为至少连接在ZEP基因与S标签基因之间,特别优选为仅在ZEP基因与S标签基因之间(不在CCS基因与S标签基因之间)。
1-4.其他基因
在重组载体中,除了上述的序列以外,还可以包含第一启动子和第二启动子以外的调节序列。这样的调节序列位于类胡萝卜素的生物合成基因的上游基因的序列和/或类胡萝卜素的生物合成基因的下游基因的序列的上游(5'侧)、该序列内、或下游(3'侧),对转录、RNA加工或稳定性等造成影响。作为调节序列,可以举出翻译前导序列、内含子、聚腺苷酸化识别序列、RNA加工位点、效应结合位点、茎-环结构等。
1-5.表达载体
作为表达载体,可以使用由能够在宿主内自主增殖的质粒、病毒、噬菌体、转座子、IS元件、噬菌粒、粘粒等作为基因重组用而构建的表达载体。作为这样的表达载体,例如可举出pET、pLG、pACYC、pBR、pUC、pKC、pRep、pHS、pKK、pDHE、pPLc、pMBL、pUR、pIN、λgt、pBdCI、pUB、pC、pBD;pSA、pAJ、YEp、YRp、YIp、pYAC、pCDM、pMT2PC、腺病毒载体、腺相关病毒载体、逆转录病毒载体、疱疹病毒载体等。另外,在第一重组载体和第二重组载体中,成为基础的表达载体可以相同,也可以不同。另外,表达载体只要选择与宿主细胞的适当组合来使用即可,例如作为将大肠杆菌作为宿主细胞时的优选例,可举出pACYC载体和pUC载体与JM101(DE3)大肠菌株的组合等。
1-6.宿主细胞
作为宿主,只要是重组载体稳定且能够自主增殖、能够表达外源基因的性状,就没有特别限制,另外,不论类胡萝卜素产生细胞和类胡萝卜素非产生细胞。作为宿主的优选例,例如可举出属于大肠杆菌(Escherichia coli)等的埃希菌(Escherichia)属、枯草芽孢杆菌(Bacillus subtilis)等的芽孢杆菌属、恶臭假单胞菌(Pseudomonas putida)等的假单胞菌属等的细菌;酵母(例如酿酒酵母(Saccharomyces cerevisiae)、裂殖酵母(S.pombe)、产朊假丝酵母(Candida utilis)、巴斯德毕赤酵母(Pichia pastoris)、红法夫酵母(Xanthophyllomyces dendrorhous)等)等,除此以外,也可以是动物细胞、昆虫细胞、植物等。其中,作为这些中特别优选的宿主,可举出大肠杆菌。
1-7.转化体的制作
本发明的转化体可通过向宿主导入重组载体而得到,向宿主导入重组载体的条件根据宿主的种类等适宜设定即可。如果是宿主为细菌的情况下,则例如可举出热休克法、使用钙离子处理得到的感受态细胞的方法和电穿孔法等。如果是宿主为酵母的情况下,则例如可举出电穿孔法(electroporation method)、原生质球法和醋酸锂法等。如果是宿主为动物细胞的情况下,则例如可举出电穿孔法、磷酸钙法和脂质转染法等。如果是宿主为昆虫细胞的情况下,则例如可举出磷酸钙法、脂质转染法和电穿孔法等。如果是宿主为植物细胞的情况下,则可举出例如电穿孔法、农杆菌法、基因枪法、及PEG法等。
重组载体是否被引入到宿主的确认可以通过PCR法、Southern杂交法、以及Northern杂交法等进行。
在通过PCR法确认重组载体是否被引入到宿主的情况下,例如,从转化体中分离、纯化重组载体即可。
重组载体的分离、纯化例如在宿主为细菌的情况下,可以基于将细菌裂解而得到的溶菌物来进行。作为裂解方法,例如可利用溶菌酶(lysozyme)等溶菌酶来实施处理,可根据需要合并使用蛋白酶及其它酶、以及十二烷基硫酸钠(SDS)等表面活性剂。
进而,也可以组合冷冻融解、超声波照射以及弗氏压碎(French press)处理这样的物理破碎方法。从溶菌物中分离、纯化DNA例如可以通过将利用苯酚处理及蛋白酶处理的除蛋白处理、核糖核酸酶处理、醇沉淀处理以及市售的试剂盒适宜组合来进行。
DNA的切断可以按照常规方法、例如使用限制酶处理来进行。作为限制酶,例如可以使用作用于特定的核苷酸序列的II型限制酶。DNA与表达载体的结合例如可以使用DNA连接酶进行。
然后,将分离、纯化的DNA用特定的限制酶切断,进行琼脂糖凝胶电泳、聚丙烯酰胺凝胶电泳、毛细管电泳等,利用溴化乙锭和SYBR Green液等进行染色,根据检测出的带长图案确认转化。
另外,将分离、纯化的DNA作为模板,设计引物进行PCR,由此也能够确认转化。对通过PCR得到的扩增产物进行琼脂糖凝胶电泳、聚丙烯酰胺凝胶电泳、毛细管电泳等,利用溴化乙锭和SYBR Green液等进行染色,然后将扩增产物作为带进行检测,由此能够确认转化。
另外,也可以使用预先用荧光色素等标签的引物进行PCR来检测扩增产物。进而,也可以采用使扩增产物结合于微孔板等固相并通过荧光及酶反应等确认扩增产物的方法。
2.类胡萝卜素组合物的制造方法
本发明的类胡萝卜素组合物的制造方法可以通过包括培养本发明的转化体的工序的制造方法得到。
转化体的培养条件只要考虑宿主的营养生理性质适宜设定即可,但优选举出液体培养。另外,如果是进行工业制造的情况下,则优选通气搅拌培养。
作为培养基的营养源,可使用转化体的生长所必需的营养源。作为碳源,只要是能够资源化的碳化合物即可,例如可举出糖蜜、葡萄糖、果糖、麦芽糖、蔗糖、乳糖、蔗糖、淀粉、乳糖、甘油、丙酮酸等。
作为氮源,只要是能够资源化的氮化合物即可,例如可举出玉米浆、蛋白胨(兽肉蛋白胨、酪蛋白蛋白胨、大豆蛋白胨等)、提取物(肉提取物、酵母提取物)等天然成分;乙酸铵、氯化铵、硫酸铵等铵盐;谷氨酸、天门冬氨酸、甘氨酸等氨基酸等。
除了碳源和氮源之外,例如,也可以根据需要使用磷酸盐、碳酸盐、硫酸盐、镁、钙、钾、铁、锰和锌等盐类、以及特定的维生素等。
培养温度可以在本发明的转化体能够生长且本发明的转化体产生本发明的多肽的范围内适当设定,优选可举出20~40℃左右、优选为30~37℃左右。培养只要估计本发明的多肽达到最高产量的时期而在适当时期完成即可,作为培养时间的例子,可举出24~200小时、优选为60~90小时左右。
作为培养结束后的菌体的回收方法,可举出培养液的过滤、倾析、离心分离等。回收的菌体根据需要可以用水、氯化钠溶液、二甲基甲酰胺等进行处理。
在本发明的制造方法中得到的类胡萝卜素组合物中,典型地包含辣椒红色素,优选进一步含有辣椒玉红素、辣椒红色素3'-乙酸酯、辣椒玉红素3-乙酸酯、类胡萝卜黄色素A、辣椒玉红素二乙酸酯、辣椒红色素3,6-环氧化物和/或辣椒红色素3,6-环氧化物3′-乙酸酯。
从菌体中提取类胡萝卜素组合物可以使用适当的有机溶剂。作为有机溶剂的例子,可举出甲醇、乙醇、异丙醇、丙酮、己烷、二乙基醚、四氢呋喃、甲基乙基酮、甲基异丁基酮、二氯甲烷、氯仿、二甲基甲酰胺、二甲基亚砜、乙酸甲酯、乙酸乙酯等,可以从这些溶剂中选择1种或多种。这些有机溶剂中,优选可举出丙酮、甲醇和二乙基醚。
如上所述那样得到的类胡萝卜素组合物组分可以作为组合物直接进行浓缩和干燥,也可以将各类胡萝卜素进行纯化并浓缩和干燥。
作为纯化方法,例如可以通过适当组合凝胶过滤、吸附色谱法、离子交换色谱法、亲和色谱法等方法来进行。作为干燥方法,可举出冷冻干燥、真空干燥、喷雾干燥等。
实施例
以下,通过实施例对本发明进行更具体的说明,但本发明不限定于这些实施例。应予说明,在以下实施例和比较例中使用的启动子是Plac、类胡萝卜素的生物合成基因固有的启动子(是在菠萝泛菌中生来具备的启动子,具体而言是包含在DDBJ登录号D90087中的启动子)、PBAD或Ptac,启动子强度的关系为Plac<该类胡萝卜素的生物合成基因固有的启动子<PBAD<Ptac,PBAD的启动子强度为该类胡萝卜素的生物合成基因固有的启动子的启动子强度的3倍以上。
[比较例1]
在本比较例中,构建图2所示的pACHP-Zea作为引入的第一重组载体,构建图2所示的pUC-Plac-CaCCS-CaZEP作为第二重组载体。这些重组载体设计为上游基因的表达量利用类胡萝卜素的生物合成基因固有的启动子(来自菠萝泛菌)而相对高,下游基因的表达量利用Plac而相对低。将本比较例的第一重组载体和第二重组载体的概要示于表1。
[表1]
作为第一重组载体的pACHP-Zea是非专利文献3的报告中使用的玉米黄质合成用的载体。pACHP-Zea是在pACYC载体中插入来自雨生红球藻(Haematococcus pluvalis)的idi基因(DDBJ登录号AB019034),并且插入玉米黄质生物合成所需的基因群(来自菠萝泛菌的crtE、crtY、crtI、crtB以及crtZ各基因(DDBJ登录号D90087))而构成。
作为第二重组载体的pUC-Plac-CaCCS-CaZEP通过将来自辣椒(Capsicum annuum)的ZEP基因(以下记载为“CaZEP”;DDBJ登录号XM_016705616)和来自辣椒(Capsicumannuum)的CCS基因(以下记载为“CaCCS”;DDBJ登录号X76165)配置在pUC载体的Plac启动子下游而制作。
将这些第一重组载体和第二重组载体一起通过热休克法导入大肠杆菌JM109株中。
导入重组载体后,将LB培养基上的单个菌落移植到5mL的LB液体培养基中,在200rpm、37℃下振荡培养16小时。将100μL的培养液移植到10mL的TB液体培养基中,在200rpm、30℃下振荡培养24小时后加入10μL的异丙基-β-D-吡喃半乳糖苷(IPTG)诱导基因表达,进而继续培养48小时。另外,作为抗生素,在培养基中加入氯霉素(30mg/L)和卡那霉素(40mg/L)。
将所得到的培养液的1mL以8000g离心分离1分钟,除去上清液,回收细胞组分。将细胞组分悬浮于1%氯化钠溶液中,以8000g离心分离1分钟,除去上清液,回收细胞。在回收的细胞中添加0.5mL的丙酮、1mL的二乙基醚/己烷(1:1(体积基准)),用涡流强烈搅拌。添加1mL水并搅拌后静置5分钟,将上层回收至1.5mL离心管中,利用真空离心浓缩器使其干燥固化。
将得到的干燥固化物供于使用了Aquity H-class UPLC(Waters)的类胡萝卜素的分离。作为柱,使用Aquity UPLC BEH C18柱(2.1x100mm,1.7μm,Waters),作为流动相A,使用乙腈:超纯水(85:15(体积基准)),作为流动相B,使用乙腈:甲醇(65:35(体积基准)),以流动相A:流动相B成为100:0至0:100的方式用8分钟的梯度进行送液。然后,通过将0:100保持5分钟,分离类胡萝卜素。
其结果,无法确认溶出时间与辣椒红色素标准品相同的峰。
[比较例2(比较例2-1和比较例2-2)]
在本比较例中,构建图3所示的pAC-HiEBIY-Z-CaZEP作为引入的第一重组载体,构建图3所示的pUC-CaCCS(将使用该第二重组载体的情况作为比较例2-1)或pUC-LlCCS(将使用该第二重组载体的情况作为比较例2-2)。这些重组载体设计成上游基因和下游基因中的ZEP基因的表达量利用Ptac而相对高,下游基因中的CCS基因的表达量利用PBAD而相对低。将本比较例的第一重组载体和第二重组载体的概要示于表2。
[表2]
在比较例2-1和比较例2-2中,以非专利文献3的报告为基础,如下制作pAC-HiEBIY-Z-CaZEP作为第一重组载体:在pACYC质粒载体中,与来自雨生红球藻(Haematococcus pluvalis)的idi基因一起,将紫黄质生物合成所需的来自菠萝泛菌的crtE、crtB、crtI、crtY、crtZ各基因以及CaZEP的氨基酸残基第64位至第668位(由于来自植物的基因大多在蛋白质的N末端具有在大肠杆菌内不必要的信号序列。简称为CaZEP64-668,在图3中表示为ZEP*)配置于Ptac启动子的下游。
与比较例1相同,将制作得到的第一重组载体pAC-HiEBIY-Z-CaZEP导入大肠杆菌JM101(DE3)并培养,得到上清液。
将得到的上清液供于使用日立制HPLC Chromaster系统的类胡萝卜素的分离。作为柱,使用Inertsil ODS-3(4.6x150mm,5μm,GL Sciences),作为流动相A,使用乙腈,作为流动相B,使用异丙醇,将流动相A:流动相B以100:0送液5分钟后,以流动相A:流动相B成为100:0至50:50(体积基准)的方式用5分钟的梯度进行送液,进一步将50:50保持5分钟,分离类胡萝卜素。
图4中示出得到的HPLC色谱图。如图4所示,确认了在紫黄质和花药黄质以外还生成了玉米黄质和β-胡萝卜素以及其它多种类胡萝卜素中间体。
接着,在比较例2-1和比较例2-2中的比较例2-1中,调查了通过使CaCCS在该类胡萝卜素生产大肠菌株中共表达,能否生产辣椒红色素和辣椒玉红素。预测CaCCS的N末端叶绿体信号序列,设计了切削N末端氨基酸的CaCCS41-498。制作将CaCCS41-498配置于pUC载体的PBAD启动子下游的质粒pUC-CaCCS作为第二重组载体,并导入上述的类胡萝卜素生产大肠菌株。
另外,在比较例2-1和比较例2-2中的比较例2-2中,除了将来自卷丹百合(Liliumlancifolium)的CCS(以下记载为“LlCCS”;DDBJ登录号GU443955)代替CaCCS而导入之外,与比较例2-1相同地制作pUC-LlCCS作为第二重组载体,并导入上述的类胡萝卜素生产大肠菌株。
在比较例2-1和比较例2-2中,导入上述各第二重组载体后,从LB培养基上的单个菌落移植到3mL的LB液体培养基中,在200rpm、37℃下振荡培养16小时,将100μL的培养液移植到10mL的TB液体培养基中,在200rpm、30℃下振荡培养24小时后,加入10μL的异丙基-β-D-吡喃半乳糖苷(IPTG)诱导基因表达,进而继续培养48小时。另外,作为抗生素,在培养基中加入氯霉素(30mg/L)、卡那霉素(40mg/L)。
将所得到的培养液的1mL以8000g离心分离1分钟,除去上清液,回收细胞组分。将细胞组分悬浮于1%氯化钠溶液中,以8000g离心分离1分钟,除去上清液,回收细胞。在回收的细胞中添加丙酮/甲醇(7:3(体积基准)),提取类胡萝卜素,以12000g离心分离2分钟,将上清液用于类胡萝卜素分析。
将得到的上清液供于使用日立制HPLC Chromaster系统的类胡萝卜素的分离。作为柱,使用Inertsil ODS-3(4.6x150mm,5μm,GL Sciences),作为流动相A,使用乙腈,作为流动相B,使用异丙醇,将流动相A:流动相B以100:0送液5分钟后,以流动相A:流动相B成为100:0至50:50(体积基准)的方式用5分钟的梯度进行送液,进一步将50:50保持5分钟。
其结果,如图4的比较例2-1和比较例2-2的结果所示,即使在使CaCCS和LlCCS中的任一者共表达的情况下,所生产的类胡萝卜素也没有变化,无法确认到辣椒红色素的生成。
[实施例1]
在本实施例中,构建图5所示的pACHP-Zea作为引入的第一重组载体,构建图5所示的pUC-CaCCS-CaZEP作为第二重组载体。这些重组载体设计为上游基因的表达量利用类胡萝卜素的生物合成基因固有的启动子(源自菠萝泛菌)而相对低,下游基因的表达量利用PBAD而相对高。将本实施例的第一重组载体和第二重组载体的概要示于表3。
[表3]
作为第一重组载体的pACHP-Zea与比较例1相同地制作。根据非专利专利文献3的报告可知,该pACHP-Zea在基于该转化体的类胡萝卜素组合物中未检测到β-胡萝卜素,玉米黄质占据总类胡萝卜素。
作为第二重组载体的pUC-CaCCS-CaZEP是将CaZEP64-668和CaCCS41-498配置于pUC载体的PBAD启动子下游而制作。
将这些第一重组载体和第二重组载体一起与比较例1相同地导入到大肠杆菌JM101(DE3)。
导入重组载体后,将LB培养基上的单个菌落移植到3mL的LB液体培养基中,在200rpm、37℃下振荡培养16小时。将100μL的培养液移植到10mL的TB液体培养基中,在200rpm、30℃下振荡培养24小时后加入10μL的异丙基-β-D-吡喃半乳糖苷(IPTG)诱导基因表达,进而继续培养48小时。另外,作为抗生素,在培养基中加入氯霉素(30mg/L)和卡那霉素(40mg/L)。
将得到的培养液的1mL以8000g离心分离1分钟,除去上清液,回收细胞组分。将细胞组分悬浮于1%氯化钠溶液中,以8000g离心分离1分钟,除去上清液,回收细胞。添加丙酮/甲醇(7:3(体积基准)),提取类胡萝卜素,以12000g离心分离2分钟,将上清液用于类胡萝卜素分析。
将得到的上清液供于使用日立制HPLC Chromaster系统的类胡萝卜素的分离。作为柱,使用Inertsil ODS-3(4.6x150mm,5μm,GL Sciences),作为流动相A,使用乙腈,作为流动相B,使用异丙醇,将流动相A:流动相B以100:0送液5分钟后,以流动相A:流动相B成为100:0至50:50(体积基准)的方式用5分钟的梯度进行送液,进一步将50:50保持5分钟,分离类胡萝卜素。
另外,还将得到的上清液(类胡萝卜素)供于Aquity H-class UPLC(Waters)。作为柱,使用Aquity UPLC BEH C18柱(2.1x 100mm,1.7μm,Waters),作为流动相A,使用乙腈:超纯水(85:15(体积比)),作为流动相B,使用乙腈:甲醇(65:35(体积比)),以A:B为100:0至0:100用8分钟的梯度进行送液,然后以将0:100保持5分钟的程序分离类胡萝卜素。
将分析结果示于图6。在图6中,示出了单独表达第一表达载体的情况下的HPLC色谱图(上)、和使第一表达载体和第二表达载体共表达的情况下(实施例1)的HPLC色谱图(下)的结果、以及后者(实施例1)中检测出的峰1、2、3、5的吸收光谱。如图6所示,通过第一表达载体和第二表达载体的共表达,能够确认到新的峰5。
将相同的上清液(类胡萝卜素)(样品(sample))用UPLC分析并与标准品(standards)进行比较的结果示于图7a。如图7a所示,通过第一表达载体和第二表达载体的共表达(实施例1),能够确认到新的峰5。另外,将峰5的吸收光谱示于图7b。图7a的利用UPLC色谱图确认到的峰5显示出与辣椒红色素相同的保留时间和吸收光谱。基于摩尔吸光系数由峰强度进行计算的结果,单位培养液的辣椒红色素生产量为292μg/L。
峰5的LC/MS分析使用搭载有Acquity UPLC系统的Waters Xevo G2S Q TOF质谱分析仪(Waters)进行。基于飞行时间型质谱分析装置(ESI-TOF-MS)的分析条件是使用毛细管电压3.2kV、锥孔(cone)电压20eV、120℃、30L/h的氮气,扫描m/z100~1500。MS/MS光谱通过以20V的碰撞能量使用氩的四极TOF MS/MS装置进行测定。UV-VIS吸收光谱使用光电二极管阵列以200nm~600nm进行记录。HPLC分离使用Acquity 1.7μm BEH UPLC C18柱(Waters),使用从乙腈:超纯水(85:15(体积基准))至乙腈:甲醇(65:35(体积基准))的梯度程序(流速0.4mL/分钟、15分钟)。将结果示于图7c。图7c所示的分子离子峰与辣椒红色素的分子量的计算值一致。
另外,峰5的1HNMR(500MHz)光谱将TMS作为内标,使用氘代氯仿使用Varian UNITYINOVA 500分光计(Spectrometer)(Varian Corporation)进行测定。其结果,峰5鉴定为辣椒红色素。
[实施例2(实施例2-1和实施例2-2)]
在本实施例中,构建图8所示的pACHP-Zea作为引入的第一重组载体,构建图8所示的pUC-CaCCS-CaZEP(2种)作为第二重组载体。这些重组载体设计为上游基因的表达量利用类胡萝卜素的生物合成基因固有的启动子(源自菠萝泛菌)而相对低,下游基因的表达量利用Ptac而相对高。将本实施例的第一重组载体和第二重组载体的概要示于表4。
[表4]
作为第一重组载体的pACHP-Zea与比较例1相同地制作。根据非专利文献3的报告可知,该pACHP-Zea在基于该转化体的类胡萝卜素组合物中未检测到β-胡萝卜素,玉米黄质占据总类胡萝卜素。
作为第二重组载体的pUC-CaCCS-CaZEP是将CaZEP(未除去信号序列的序列)和CaCCS(未除去信号序列的序列)或CaCCS41-498配置于pUC载体的Ptac启动子下游而制作。
将这些第一重组载体和第二重组载体一起与比较例1相同地导入到大肠杆菌JM101(DE3)。
导入重组载体后,将LB培养基上的单个菌落移植到0.5mL的LB液体培养基中,在1000rpm、37℃下振荡培养16小时。将20μL培养液移植到2mL的TB液体培养基中,在1000rpm、30℃下振荡培养24小时后加入2μL的1M异丙基-β-D-吡喃半乳糖苷(IPTG)诱导基因表达。进而培养48小时。作为抗生素,在培养基中加入氯霉素(30mg/L)、卡那霉素(40mg/L)。
将得到的培养液的1mL以8000g离心分离1分钟,除去上清液,回收细胞组分。将细胞组分悬浮于1%氯化钠溶液中,以8000g离心分离1分钟,除去上清液,回收细胞。添加丙酮/甲醇(7:3(体积基准)),提取类胡萝卜素,以12000g离心分离2分钟,将上清液用于类胡萝卜素分析。
将得到的上清液供于使用日立制HPLC Chromaster系统的类胡萝卜素的分离。作为柱,使用Inertsil ODS-3(4.6x150mm,5μm,GL Sciences),作为流动相A,使用乙腈,作为流动相B,使用异丙醇,将流动相A:流动相B以100:0送液5分钟后,以流动相A:流动相B成为100:0至50:50(体积基准)的方式用5分钟的梯度进行送液,进一步将50:50保持5分钟,分离类胡萝卜素。
应予说明,为了用于比较,使用除了在上述的第二表达载体中既没有引入CaCCS,也没有引入CaCCS41-498以外相同地制作的比较用的第二表达载体,进行相同的操作。
将分析结果示于图9。在图9中,示出了在使第一表达载体和比较用的第二表达载体共表达的情况下的HPLC色谱图(上)、在使第一表达载体和第二表达载体(引入了全长CaCCS的载体:实施例2-2)共表达的情况下的HPLC色谱图(中)、及在使第一表达载体和第二表达载体(引入了除去了信号序列的CaCCS41-498的载体:实施例2-1)共表达的情况下的HPLC色谱图(下)的结果,以及在使第一表达载体和第二表达载体(引入了除去了信号序列的CaCCS41-498的载体:实施例2-1)共表达的情况下检测出的峰1、2、3、5的吸收光谱。如图9所示,通过第一表达载体和第二表达载体的共表达能够确认到新的峰5、即辣椒红色素。
另外,将图9中的辣椒红色素的峰5的强度进行了对比的图示于图10。如图10所示,与在使用引入了全长CCS的第二表达载体的情况下(实施例2-2)相比,在使用引入了除去了信号序列的CCS41-498的第二表达载体的情况下(实施例2-1),产生了更多的辣椒红色素。
进而,基于摩尔吸光系数由峰强度计算出的峰5的辣椒红色素的产生量的结果,在使用引入了全长CCS的第二表达载体的情况下(实施例2-2)的单位培养液的辣椒红色素生产量为326μg/L,在使用引入了除去了信号序列的CCS41-498的第二表达载体的情况下(实施例2-1)的单位培养液的辣椒红色素生产量为452μg/L。
[实施例3(实施例3-1、实施例3-2、实施例3-3)]
在本实施例中,作为第一重组载体,使用了与比较例1相同的pACHP-Zea。根据非专利文献3的报告可知,该pACHP-Zea在基于该转化体的类胡萝卜素组合物中未检测到β-胡萝卜素,玉米黄质占据总类胡萝卜素。
作为第二重组载体,制作图11所示的pUC-S/slC/sZ、pUC-S/sC/slZ、或pUC-S/slC/slZ(“S”表示S蛋白,“s”表示S-标签(S-Tag),“l”表示接头)。将本实施例的第一重组载体和第二重组载体的概要示于表5。
[表5]
如图12示意性所示,该第二重组载体设计为,CCS和ZEP分别作为与S-标签融合的蛋白质(分别在图12中作为“S标签+接头+CCS”以及“S标签+接头+ZEP”示出)表达后,经由多聚体化的S蛋白(在图12中作为“多聚体”示出)间接地结合而形成复合体(在图12中作为“复合体”示出)。将这样设计的第二重组载体的方式也表现为“引入有S-标签”。
图11所示的pUC-S/slC/sZ、pUC-S/sC/slZ以及pUC-S/slC/slZ这3种第二重组载体设计为具有S-标签与CCS之间、和/或S-标签与ZEP之间有无接头的组合不同的构成。通过在引入有S-标签的第二重组载体内使接头的导入位置这样不同,构建了表达后CCS与ZEP之间的自由度不同的3个条件。
另外,将与引入有S-标签的第二重组载体构建相关的序列信息示于下述表6。
[表6]
碱基序列 | 氨基酸序列 | |
S蛋白 | 序列号1 | 序列号2 |
S-标签 | 序列号3 | 序列号4 |
S-标签/CaCCS | 序列号5 | 序列号6 |
S-标签/接头·/CaCCS | 序列号7 | 序列号8 |
S-标签/CaZEP | 序列号9 | 序列号10 |
S-标签/接头/CaZEP | 序列号11 | 序列号12 |
如序列号8、12所示,作为接头,使用由非极性氨基酸的甘氨酸(G)和极性氨基酸的丝氨酸(S)构成的单元(GS)连续10个单位的由20个氨基酸残基构成的接头({GS}10接头)。另外,S-标签的融合位点为CaCCS以及CaZEP各自的N末端。作为导入S蛋白、S-标签等的构建体,使用pUClac-ptac-CaCCSM40-CaZEP(pMF541)(Furubayashi,M.,Kubo,A.,Takemura,M.,Otani,Y.,Maoka,T.,Terada,Y.,Yaoi,K.,Ohdan,K.,Misawa,N.and Mitani,Y.(2021)J.Agric.Food Chem.,69,5076-5085)。
将上述的第一重组载体和第二重组载体与实施例1相同地导入后,将LB培养基上的单个菌落移植到3mL的2YT液体培养基中,使用Yamato scientific Co.,Ltd.制造的振荡浴(Shaking Bath)BW201以105rpm、28℃振荡培养16小时。将20μL的培养液移植到2.1mL的TB液体培养基中,使用Kuhner公司制造的Lab-Therm LT-X,在180rpm、30℃下振荡培养4小时后,加入0.1mM的异丙基-β-D-吡喃半乳糖苷(IPTG)诱导基因表达,进而继续培养48小时。另外,作为抗生素,在培养基中加入氯霉素(30mg/L)和氨苄青霉素(100mg/L)。
将得到的培养液的1mL以8000g离心分离1分钟,除去上清液,回收细胞组分。将细胞组分悬浮于1%氯化钠溶液中,以8000g离心分离1分钟,除去上清液,回收细胞。添加甲醇和氯仿,提取类胡萝卜素,以12000g离心分离2分钟,将上清液用于类胡萝卜素分析。
将得到的上清液供于使用日立制HPLC Chromaster系统的类胡萝卜素的分离。作为柱,使用Inertsil ODS-3(4.6x150mm,5μm,GL Sciences),作为流动相A,使用乙腈,作为流动相B,使用异丙醇,将流动相A:流动相B以100:0送液5分钟后,以流动相A:流动相B成为100:0至50:50(体积基准)的方式用5分钟的梯度送液,进一步将50:50保持5分钟,分离类胡萝卜素。
将HPLC分析中的对比各类胡萝卜素的峰强度的图表示于图13。在x图13中,一并示出在使用了未引入S-标签的实施例2-1中使用的第二重组载体pUC-CaCCS41-498-CaZEP(图中表示为“C/Z”)的情况下的结果。如图13所示,与未引入S-标签的情况(实施例2-1)相比,在使用了引入有S-标签的第二重组载体的情况下(实施例3-1~实施例3-3)大量蓄积了辣椒红色素。另外,在引入有S-标签的3种第二重组载体中,在使用在S-标签与ZEP之间引入有接头的pUC-S/sC/slZ的情况下(实施例3-2),辣椒红色素蓄积量也显著多。
进而,将基于摩尔吸光系数由峰强度计算出的结果示于表7。与使用未引入S-标签的第二重组载体的情况下(实施例2-1)的单位培养液的辣椒红色素生产量相比,在使用引入有S-标签的第二重组载体的情况下(实施例3-1~3-3)的单位培养液的辣椒红色素生产量提高。特别是,在使用在S-标签与ZEP之间引入有接头的pUC-S/sC/slZ作为第二重组载体的情况下(实施例3-2),不仅单位培养液的辣椒红色素生产量显著提高,而且相对于表7所示的类胡萝卜素的总量,辣椒红色素的比率也显著提高。
[表7]
另外,在将pUC-S/sC/slZ用作第二载体的情况下(实施例3-2)得到的类胡萝卜素通过与实施例1相同的方法进行UPLC分析,将结果示于图14。由图14可知,检测出辣椒红色素3′-乙酸酯、辣椒红色素3,6-环氧化物、辣椒红色素3,6-环氧化物3′-乙酸酯、辣椒玉红素3-乙酸酯、辣椒玉红素二乙酸酯、类胡萝卜黄色素A的各峰。用与实施例1相同的方法将这些类胡萝卜素进行LC/MS分析,结果分子离子峰与各类胡萝卜素的分子量的计算值一致。进而,将由UPLC分析的峰强度算出各类胡萝卜素的比率的结果示于表8。
[表8]
为了NMR测定,类胡萝卜素使用硅胶色谱法,从己烷到醚;到己烷(1:1)、醚:丙酮(1:1)依次提高极性,分取各组分并浓缩后,进一步使用硅胶(silica gel)(Cosmosil5 SL-II)柱10×250mm,以丙酮:己烷(3:7)为流动相,分取以流速2ml/min进行分离得到的各类胡萝卜素峰,将所得样品用与实施例1相同的方法用于1HNMR测定。1HNMR分析的结果,将鉴定的类胡萝卜素(辣椒红色素、辣椒玉红素、类胡萝卜黄色素A及其他衍生物等)的结构式示于下述式。
[化1]
序列表自由文本
序列号3是编码S-标签的碱基序列的第34位~第36位被替换为编码苯丙氨酸的序列而成的。
序列号4是在S-标签的氨基酸序列中第12位被替换为苯丙氨酸而成的。
序列号5和6分别是导入了S-标签的CaCCS的碱基序列和氨基酸序列。
序列号7和8分别是经由接头导入了S-标签的CaCCS的碱基序列和氨基酸序列。
序列号9和10分别是导入了S-标签的CaZEP的碱基序列和氨基酸序列。
序列号11和12分别是经由接头导入了S-标签的CaZEP的碱基序列和氨基酸序列。
序列表
<110> 江崎格力高株式会社
<120> 类胡萝卜素化合物
<130> 21076WO
<150> JP2020-188462
<151> 2020-11-12
<160> 12
<170> PatentIn version 3.5
<210> 1
<211> 321
<212> DNA
<213> 人类
<400> 1
atggattcta gctcaaacta ttgtaatcaa atgatgaaat cccgtaacct gaccaaagac 60
cgctgtaaac ctgtcaatac ctttgtccac gaatctttag ccgatgttca agccgtgtgt 120
tctcaaaaaa atgtagcctg taaaaatggc caaaccaact gttatcaatc atattccacc 180
atgtccatta ccgattgtcg cgaaacagga tcttccaaat atccaaattg tgcatataaa 240
actacccaag cgaacaaaca tattatcgta gcatgtgaag gcaacccgta cgtacctgtc 300
cacttcgacg catcagtgta a 321
<210> 2
<211> 106
<212> PRT
<213> 人类
<400> 2
Met Asp Ser Ser Ser Asn Tyr Cys Asn Gln Met Met Lys Ser Arg Asn
1 5 10 15
Leu Thr Lys Asp Arg Cys Lys Pro Val Asn Thr Phe Val His Glu Ser
20 25 30
Leu Ala Asp Val Gln Ala Val Cys Ser Gln Lys Asn Val Ala Cys Lys
35 40 45
Asn Gly Gln Thr Asn Cys Tyr Gln Ser Tyr Ser Thr Met Ser Ile Thr
50 55 60
Asp Cys Arg Glu Thr Gly Ser Ser Lys Tyr Pro Asn Cys Ala Tyr Lys
65 70 75 80
Thr Thr Gln Ala Asn Lys His Ile Ile Val Ala Cys Glu Gly Asn Pro
85 90 95
Tyr Val Pro Val His Phe Asp Ala Ser Val
100 105
<210> 3
<211> 60
<212> DNA
<213> 人工序列
<220>
<223> 第34位~第36位被替换为编码苯丙氨酸的序列的编码S-标签的碱基序列
<400> 3
aaagaaaccg ctgctgcgaa atttgaacgc cagttcatgg actcgtctac tagcgcagct 60
<210> 4
<211> 20
<212> PRT
<213> 人工序列
<220>
<223> 第12位被替换为苯丙氨酸的S-标签氨基酸序列
<400> 4
Lys Glu Thr Ala Ala Ala Lys Phe Glu Arg Gln Phe Met Asp Ser Ser
1 5 10 15
Thr Ser Ala Ala
20
<210> 5
<211> 1446
<212> DNA
<213> 人工序列
<220>
<223> 导入了S-标签的CaCCS
<400> 5
atggcaaaag aaaccgctgc tgcgaaattt gaacgccagt tcatggactc gtctactagc 60
gcagctatgt ttcattatcg taacaaaagc agcacccatt tttgcagctt tctggatctg 120
gcgccgacca gcaaaccgga aagcctggat gtgaatatca gctgggttga taccgatctg 180
gatggcgccg aatttgatgt gattatcatt ggtaccggtc cggccggcct gcgtctggca 240
gaacaggtta gcaaatatgg tattaaagtg tgctgtgttg atccgagccc gctgagcatg 300
tggccgaaca attatggcgt gtgggttgat gaatttgaaa aactgggtct ggaagattgc 360
ctggatcata aatggccggt gagctgtgtt catatcagcg atcataaaac caaatatctg 420
gatcgcccgt atggtcgcgt gagccgtaaa aaactgaaac tgaaactgct gaacagctgc 480
gtggaaaacc gtgttaaatt ctacaaagcg aaagtgctga aagttaaaca tgaagaattt 540
gaaagcagca ttgtgtgtga tgatggccgc aaaatcagcg gtagcctgat tgttgatgcg 600
agcggctatg ccagcgattt tatcgaatat gataaaccgc gtaatcatgg ctatcaggtg 660
gcgcatggta ttctggccga agttgataat catccgtttg atctggataa aatgatgctg 720
atggattggc gcgatagcca tctgggtaac gaaccgtatc tgcgtgtgaa aaataccaaa 780
gaaccgacct ttctgtatgc gatgccgttt gatcgcaatc tggtgtttct ggaagaaacc 840
agcctggtta gccgtccgat gctgagctat atggaagtga aacgtcgcat ggttgcccgc 900
ctgcgtcatc tgggcattaa agtgcgcagc gttctggaag aagaaaaatg tgtgatcacg 960
atgggcggcc cgctgccgcg tattccgcag aacgttatgg cgatcggcgg taccagcggc 1020
attgtgcatc cgagcagcgg ttatatggtt gcgcgcagca tggcgctggc accggtgctg 1080
gcggaagcca tcgttgaaag cctgggcagc acccgcatga ttcgtggtag ccagctgtat 1140
catcgcgtgt ggaatggtct gtggccgagc gatcgtcgcc gtgttcgtga atgctattgt 1200
tttggcatgg aaaccttatt aaaactggat ctggaaggta cccgccgtct gtttgatgcg 1260
tttttcgatg ttgatccgaa atattggcat ggctttctga gcagccgcct gagcgtgaaa 1320
gaactggccg ttctgagcct gtacctgttt ggtcatgcga gcaacctggc ccgtctggat 1380
atcgtgacca aatgtaccgt gccgctggtt aaactgctgg gcaatctggc gattgaaagc 1440
ctgtaa 1446
<210> 6
<211> 481
<212> PRT
<213> 人工序列
<220>
<223> 导入了S-标签的CaCCS
<400> 6
Met Ala Lys Glu Thr Ala Ala Ala Lys Phe Glu Arg Gln Phe Met Asp
1 5 10 15
Ser Ser Thr Ser Ala Ala Met Phe His Tyr Arg Asn Lys Ser Ser Thr
20 25 30
His Phe Cys Ser Phe Leu Asp Leu Ala Pro Thr Ser Lys Pro Glu Ser
35 40 45
Leu Asp Val Asn Ile Ser Trp Val Asp Thr Asp Leu Asp Gly Ala Glu
50 55 60
Phe Asp Val Ile Ile Ile Gly Thr Gly Pro Ala Gly Leu Arg Leu Ala
65 70 75 80
Glu Gln Val Ser Lys Tyr Gly Ile Lys Val Cys Cys Val Asp Pro Ser
85 90 95
Pro Leu Ser Met Trp Pro Asn Asn Tyr Gly Val Trp Val Asp Glu Phe
100 105 110
Glu Lys Leu Gly Leu Glu Asp Cys Leu Asp His Lys Trp Pro Val Ser
115 120 125
Cys Val His Ile Ser Asp His Lys Thr Lys Tyr Leu Asp Arg Pro Tyr
130 135 140
Gly Arg Val Ser Arg Lys Lys Leu Lys Leu Lys Leu Leu Asn Ser Cys
145 150 155 160
Val Glu Asn Arg Val Lys Phe Tyr Lys Ala Lys Val Leu Lys Val Lys
165 170 175
His Glu Glu Phe Glu Ser Ser Ile Val Cys Asp Asp Gly Arg Lys Ile
180 185 190
Ser Gly Ser Leu Ile Val Asp Ala Ser Gly Tyr Ala Ser Asp Phe Ile
195 200 205
Glu Tyr Asp Lys Pro Arg Asn His Gly Tyr Gln Val Ala His Gly Ile
210 215 220
Leu Ala Glu Val Asp Asn His Pro Phe Asp Leu Asp Lys Met Met Leu
225 230 235 240
Met Asp Trp Arg Asp Ser His Leu Gly Asn Glu Pro Tyr Leu Arg Val
245 250 255
Lys Asn Thr Lys Glu Pro Thr Phe Leu Tyr Ala Met Pro Phe Asp Arg
260 265 270
Asn Leu Val Phe Leu Glu Glu Thr Ser Leu Val Ser Arg Pro Met Leu
275 280 285
Ser Tyr Met Glu Val Lys Arg Arg Met Val Ala Arg Leu Arg His Leu
290 295 300
Gly Ile Lys Val Arg Ser Val Leu Glu Glu Glu Lys Cys Val Ile Thr
305 310 315 320
Met Gly Gly Pro Leu Pro Arg Ile Pro Gln Asn Val Met Ala Ile Gly
325 330 335
Gly Thr Ser Gly Ile Val His Pro Ser Ser Gly Tyr Met Val Ala Arg
340 345 350
Ser Met Ala Leu Ala Pro Val Leu Ala Glu Ala Ile Val Glu Ser Leu
355 360 365
Gly Ser Thr Arg Met Ile Arg Gly Ser Gln Leu Tyr His Arg Val Trp
370 375 380
Asn Gly Leu Trp Pro Ser Asp Arg Arg Arg Val Arg Glu Cys Tyr Cys
385 390 395 400
Phe Gly Met Glu Thr Leu Leu Lys Leu Asp Leu Glu Gly Thr Arg Arg
405 410 415
Leu Phe Asp Ala Phe Phe Asp Val Asp Pro Lys Tyr Trp His Gly Phe
420 425 430
Leu Ser Ser Arg Leu Ser Val Lys Glu Leu Ala Val Leu Ser Leu Tyr
435 440 445
Leu Phe Gly His Ala Ser Asn Leu Ala Arg Leu Asp Ile Val Thr Lys
450 455 460
Cys Thr Val Pro Leu Val Lys Leu Leu Gly Asn Leu Ala Ile Glu Ser
465 470 475 480
Leu
<210> 7
<211> 1506
<212> DNA
<213> 人工序列
<220>
<223> 经由接头导入了S-标签的CaCCS
<400> 7
atgtcgaaag aaaccgctgc tgcgaaattt gaacgccagt tcatggactc gtctactagt 60
gcagctggtt caggtagtgg ttcaggttct ggttcaggtt caggtagtgg ttcaggttct 120
ggttcaatgt ttcattatcg taacaaaagc agcacccatt tttgcagctt tctggatctg 180
gcgccgacca gcaaaccgga aagcctggat gtgaatatca gctgggttga taccgatctg 240
gatggcgccg aatttgatgt gattatcatt ggtaccggtc cggccggcct gcgtctggca 300
gaacaggtta gcaaatatgg tattaaagtg tgctgtgttg atccgagccc gctgagcatg 360
tggccgaaca attatggcgt gtgggttgat gaatttgaaa aactgggtct ggaagattgc 420
ctggatcata aatggccggt gagctgtgtt catatcagcg atcataaaac caaatatctg 480
gatcgcccgt atggtcgcgt gagccgtaaa aaactgaaac tgaaactgct gaacagctgc 540
gtggaaaacc gtgttaaatt ctacaaagcg aaagtgctga aagttaaaca tgaagaattt 600
gaaagcagca ttgtgtgtga tgatggccgc aaaatcagcg gtagcctgat tgttgatgcg 660
agcggctatg ccagcgattt tatcgaatat gataaaccgc gtaatcatgg ctatcaggtg 720
gcgcatggta ttctggccga agttgataat catccgtttg atctggataa aatgatgctg 780
atggattggc gcgatagcca tctgggtaac gaaccgtatc tgcgtgtgaa aaataccaaa 840
gaaccgacct ttctgtatgc gatgccgttt gatcgcaatc tggtgtttct ggaagaaacc 900
agcctggtta gccgtccgat gctgagctat atggaagtga aacgtcgcat ggttgcccgc 960
ctgcgtcatc tgggcattaa agtgcgcagc gttctggaag aagaaaaatg tgtgatcacg 1020
atgggcggcc cgctgccgcg tattccgcag aacgttatgg cgatcggcgg taccagcggc 1080
attgtgcatc cgagcagcgg ttatatggtt gcgcgcagca tggcgctggc accggtgctg 1140
gcggaagcca tcgttgaaag cctgggcagc acccgcatga ttcgtggtag ccagctgtat 1200
catcgcgtgt ggaatggtct gtggccgagc gatcgtcgcc gtgttcgtga atgctattgt 1260
tttggcatgg aaaccttatt aaaactggat ctggaaggta cccgccgtct gtttgatgcg 1320
tttttcgatg ttgatccgaa atattggcat ggctttctga gcagccgcct gagcgtgaaa 1380
gaactggccg ttctgagcct gtacctgttt ggtcatgcga gcaacctggc ccgtctggat 1440
atcgtgacca aatgtaccgt gccgctggtt aaactgctgg gcaatctggc gattgaaagc 1500
ctgtaa 1506
<210> 8
<211> 501
<212> PRT
<213> 人工序列
<220>
<223> 经由接头导入了S-标签的CaCCS
<400> 8
Met Ser Lys Glu Thr Ala Ala Ala Lys Phe Glu Arg Gln Phe Met Asp
1 5 10 15
Ser Ser Thr Ser Ala Ala Gly Ser Gly Ser Gly Ser Gly Ser Gly Ser
20 25 30
Gly Ser Gly Ser Gly Ser Gly Ser Gly Ser Met Phe His Tyr Arg Asn
35 40 45
Lys Ser Ser Thr His Phe Cys Ser Phe Leu Asp Leu Ala Pro Thr Ser
50 55 60
Lys Pro Glu Ser Leu Asp Val Asn Ile Ser Trp Val Asp Thr Asp Leu
65 70 75 80
Asp Gly Ala Glu Phe Asp Val Ile Ile Ile Gly Thr Gly Pro Ala Gly
85 90 95
Leu Arg Leu Ala Glu Gln Val Ser Lys Tyr Gly Ile Lys Val Cys Cys
100 105 110
Val Asp Pro Ser Pro Leu Ser Met Trp Pro Asn Asn Tyr Gly Val Trp
115 120 125
Val Asp Glu Phe Glu Lys Leu Gly Leu Glu Asp Cys Leu Asp His Lys
130 135 140
Trp Pro Val Ser Cys Val His Ile Ser Asp His Lys Thr Lys Tyr Leu
145 150 155 160
Asp Arg Pro Tyr Gly Arg Val Ser Arg Lys Lys Leu Lys Leu Lys Leu
165 170 175
Leu Asn Ser Cys Val Glu Asn Arg Val Lys Phe Tyr Lys Ala Lys Val
180 185 190
Leu Lys Val Lys His Glu Glu Phe Glu Ser Ser Ile Val Cys Asp Asp
195 200 205
Gly Arg Lys Ile Ser Gly Ser Leu Ile Val Asp Ala Ser Gly Tyr Ala
210 215 220
Ser Asp Phe Ile Glu Tyr Asp Lys Pro Arg Asn His Gly Tyr Gln Val
225 230 235 240
Ala His Gly Ile Leu Ala Glu Val Asp Asn His Pro Phe Asp Leu Asp
245 250 255
Lys Met Met Leu Met Asp Trp Arg Asp Ser His Leu Gly Asn Glu Pro
260 265 270
Tyr Leu Arg Val Lys Asn Thr Lys Glu Pro Thr Phe Leu Tyr Ala Met
275 280 285
Pro Phe Asp Arg Asn Leu Val Phe Leu Glu Glu Thr Ser Leu Val Ser
290 295 300
Arg Pro Met Leu Ser Tyr Met Glu Val Lys Arg Arg Met Val Ala Arg
305 310 315 320
Leu Arg His Leu Gly Ile Lys Val Arg Ser Val Leu Glu Glu Glu Lys
325 330 335
Cys Val Ile Thr Met Gly Gly Pro Leu Pro Arg Ile Pro Gln Asn Val
340 345 350
Met Ala Ile Gly Gly Thr Ser Gly Ile Val His Pro Ser Ser Gly Tyr
355 360 365
Met Val Ala Arg Ser Met Ala Leu Ala Pro Val Leu Ala Glu Ala Ile
370 375 380
Val Glu Ser Leu Gly Ser Thr Arg Met Ile Arg Gly Ser Gln Leu Tyr
385 390 395 400
His Arg Val Trp Asn Gly Leu Trp Pro Ser Asp Arg Arg Arg Val Arg
405 410 415
Glu Cys Tyr Cys Phe Gly Met Glu Thr Leu Leu Lys Leu Asp Leu Glu
420 425 430
Gly Thr Arg Arg Leu Phe Asp Ala Phe Phe Asp Val Asp Pro Lys Tyr
435 440 445
Trp His Gly Phe Leu Ser Ser Arg Leu Ser Val Lys Glu Leu Ala Val
450 455 460
Leu Ser Leu Tyr Leu Phe Gly His Ala Ser Asn Leu Ala Arg Leu Asp
465 470 475 480
Ile Val Thr Lys Cys Thr Val Pro Leu Val Lys Leu Leu Gly Asn Leu
485 490 495
Ala Ile Glu Ser Leu
500
<210> 9
<211> 2073
<212> DNA
<213> 人工序列
<220>
<223> 导入了S-标签的CaZEP
<400> 9
atgtcgaaag aaaccgctgc tgcgaaattt gaacgccagt tcatggactc gtctactagt 60
gcagctatga ccatgattac gaattcgatg tatagcaccg tgttttatac cagcgttcat 120
ccgagcacca gcgtgtttag ccgcaaacag ctgccgctgc tgattagcaa agattttccg 180
gccgaactgt atcatagcct gccgtgcaaa agcctggaaa atggtcatat taagaaagtg 240
aaaggcgtta aagccaccct ggccgaagca ccggccaccc cgaccgaaaa atctaacagc 300
gaagtgccgc agaaaaaact gaaagtgctg gttgccggcg gtggcattgg tggcctggtg 360
tttgcgctgg cggcgaaaaa gaaaggcttt gatgtgctgg tttttgaacg tgatctgagc 420
gccatccgcg gcgaaggcca gtatcgtggc ccgattcaga tccagagcaa tgccctggcg 480
gccctggaag cgattgatat ggatgtggcc gaagaaatca tgaacgcggg ttgcattacc 540
ggccagcgta tcaatggtct ggttgatggt attagcggca actggtattg taaatttgat 600
acctttaccc cggccgtgga acgcggcctg ccggtgaccc gtgttattag ccgcatgacc 660
ctgcagcaga tcctggcacg tgccgtgggt gaagatgtta ttatgaacga aagcaatgtg 720
gttaactttg aagatgatgg cgaaaaagtg accgtggttc tggaaaatgg tcagcgcttt 780
accggcgatc tgctggttgg tgccgatggc atccgtagca aagtgcgcac caacctgttt 840
ggtcatagcg aagcgaccta tagcggttat acctgttata ccggcattgc cgattttgtt 900
ccggcggata tcgataccgt gggttatcgt gtttttctgg gccataaaca gtattttgtg 960
agcagcgatg ttggtggcgg taaaatgcag tggtatgcgt ttcataatga accggccggc 1020
ggtgtggatg ccccgaatgg caagaaagaa cgcctgctga aaatttttgg cggttggtgc 1080
gataatgtga tcgatctgct ggttgccacc gatgaagatg cgattctgcg tcgcgatatc 1140
tatgatcgtc cgccgacctt taattggggt cgtggtcgtg tgaccctgct gggtgatagc 1200
gttcatgcca tgcagccgaa cctgggtcag ggcggttgta tggccattga agatagctat 1260
cagctggcgc tggaactgga aaaagcctgg agccgcagcg ccgaaagcgg tagcccgatg 1320
gatgttatca gcagcctgcg tagctatgaa agcgcccgta aactgcgcgt gggtgttatt 1380
catggcctgg cccgtatggc cgcaatcatg gcaagcacct ataaagcgta tctgggcgtg 1440
ggtctgggtc cgctgagctt tctgaccaaa tatcgtattc cgcacccggg tcgtgttggc 1500
ggtcgtgtgt ttgttgatct gggcatgccg ctgatgctga gctgggtgct gggcggtaat 1560
ggtgataaac tggaaggccg tatccagcat tgccgcctga gcgaaaaagc caacgatcag 1620
ctgcgtcgct ggtttgaaga tgatgatgcg ctggaacgcg ccaccgatgc cgaatggctg 1680
ctgctgccgg ccgcaaatgg caatagcgcg ctggaaacca ttgtgctgag ccgtgatgaa 1740
gatgttccgt gcaccattgg tagcgtgagc cataccaaca tcccgggcaa aagcgtggtt 1800
ctgccgctgc cgcaggttag cgaaatgcat gcccagatta gctgtaaaaa caatgcgttt 1860
ttcgtgaccg attttcagag cgaacatggt acctgggtta tcgataatga aggccgtcgc 1920
tatcgcgtga gcccgaactt tccgatgcgt tttcatagca gcgatgtgat tgtttttggt 1980
agcgataaag cggcctttcg cgtgaaaacc atgaaatttc cgagcaaaac cgccgaagcg 2040
aaagaagaac gtaaagtggt tggcaccgcg taa 2073
<210> 10
<211> 690
<212> PRT
<213> 人工序列
<220>
<223> 导入了S-标签的CaZEP
<400> 10
Met Ser Lys Glu Thr Ala Ala Ala Lys Phe Glu Arg Gln Phe Met Asp
1 5 10 15
Ser Ser Thr Ser Ala Ala Met Thr Met Ile Thr Asn Ser Met Tyr Ser
20 25 30
Thr Val Phe Tyr Thr Ser Val His Pro Ser Thr Ser Val Phe Ser Arg
35 40 45
Lys Gln Leu Pro Leu Leu Ile Ser Lys Asp Phe Pro Ala Glu Leu Tyr
50 55 60
His Ser Leu Pro Cys Lys Ser Leu Glu Asn Gly His Ile Lys Lys Val
65 70 75 80
Lys Gly Val Lys Ala Thr Leu Ala Glu Ala Pro Ala Thr Pro Thr Glu
85 90 95
Lys Ser Asn Ser Glu Val Pro Gln Lys Lys Leu Lys Val Leu Val Ala
100 105 110
Gly Gly Gly Ile Gly Gly Leu Val Phe Ala Leu Ala Ala Lys Lys Lys
115 120 125
Gly Phe Asp Val Leu Val Phe Glu Arg Asp Leu Ser Ala Ile Arg Gly
130 135 140
Glu Gly Gln Tyr Arg Gly Pro Ile Gln Ile Gln Ser Asn Ala Leu Ala
145 150 155 160
Ala Leu Glu Ala Ile Asp Met Asp Val Ala Glu Glu Ile Met Asn Ala
165 170 175
Gly Cys Ile Thr Gly Gln Arg Ile Asn Gly Leu Val Asp Gly Ile Ser
180 185 190
Gly Asn Trp Tyr Cys Lys Phe Asp Thr Phe Thr Pro Ala Val Glu Arg
195 200 205
Gly Leu Pro Val Thr Arg Val Ile Ser Arg Met Thr Leu Gln Gln Ile
210 215 220
Leu Ala Arg Ala Val Gly Glu Asp Val Ile Met Asn Glu Ser Asn Val
225 230 235 240
Val Asn Phe Glu Asp Asp Gly Glu Lys Val Thr Val Val Leu Glu Asn
245 250 255
Gly Gln Arg Phe Thr Gly Asp Leu Leu Val Gly Ala Asp Gly Ile Arg
260 265 270
Ser Lys Val Arg Thr Asn Leu Phe Gly His Ser Glu Ala Thr Tyr Ser
275 280 285
Gly Tyr Thr Cys Tyr Thr Gly Ile Ala Asp Phe Val Pro Ala Asp Ile
290 295 300
Asp Thr Val Gly Tyr Arg Val Phe Leu Gly His Lys Gln Tyr Phe Val
305 310 315 320
Ser Ser Asp Val Gly Gly Gly Lys Met Gln Trp Tyr Ala Phe His Asn
325 330 335
Glu Pro Ala Gly Gly Val Asp Ala Pro Asn Gly Lys Lys Glu Arg Leu
340 345 350
Leu Lys Ile Phe Gly Gly Trp Cys Asp Asn Val Ile Asp Leu Leu Val
355 360 365
Ala Thr Asp Glu Asp Ala Ile Leu Arg Arg Asp Ile Tyr Asp Arg Pro
370 375 380
Pro Thr Phe Asn Trp Gly Arg Gly Arg Val Thr Leu Leu Gly Asp Ser
385 390 395 400
Val His Ala Met Gln Pro Asn Leu Gly Gln Gly Gly Cys Met Ala Ile
405 410 415
Glu Asp Ser Tyr Gln Leu Ala Leu Glu Leu Glu Lys Ala Trp Ser Arg
420 425 430
Ser Ala Glu Ser Gly Ser Pro Met Asp Val Ile Ser Ser Leu Arg Ser
435 440 445
Tyr Glu Ser Ala Arg Lys Leu Arg Val Gly Val Ile His Gly Leu Ala
450 455 460
Arg Met Ala Ala Ile Met Ala Ser Thr Tyr Lys Ala Tyr Leu Gly Val
465 470 475 480
Gly Leu Gly Pro Leu Ser Phe Leu Thr Lys Tyr Arg Ile Pro His Pro
485 490 495
Gly Arg Val Gly Gly Arg Val Phe Val Asp Leu Gly Met Pro Leu Met
500 505 510
Leu Ser Trp Val Leu Gly Gly Asn Gly Asp Lys Leu Glu Gly Arg Ile
515 520 525
Gln His Cys Arg Leu Ser Glu Lys Ala Asn Asp Gln Leu Arg Arg Trp
530 535 540
Phe Glu Asp Asp Asp Ala Leu Glu Arg Ala Thr Asp Ala Glu Trp Leu
545 550 555 560
Leu Leu Pro Ala Ala Asn Gly Asn Ser Ala Leu Glu Thr Ile Val Leu
565 570 575
Ser Arg Asp Glu Asp Val Pro Cys Thr Ile Gly Ser Val Ser His Thr
580 585 590
Asn Ile Pro Gly Lys Ser Val Val Leu Pro Leu Pro Gln Val Ser Glu
595 600 605
Met His Ala Gln Ile Ser Cys Lys Asn Asn Ala Phe Phe Val Thr Asp
610 615 620
Phe Gln Ser Glu His Gly Thr Trp Val Ile Asp Asn Glu Gly Arg Arg
625 630 635 640
Tyr Arg Val Ser Pro Asn Phe Pro Met Arg Phe His Ser Ser Asp Val
645 650 655
Ile Val Phe Gly Ser Asp Lys Ala Ala Phe Arg Val Lys Thr Met Lys
660 665 670
Phe Pro Ser Lys Thr Ala Glu Ala Lys Glu Glu Arg Lys Val Val Gly
675 680 685
Thr Ala
690
<210> 11
<211> 2133
<212> DNA
<213> 人工序列
<220>
<223> 经由接头导入了S-标签的CaZEP
<400> 11
atgtcgaaag aaaccgctgc tgcgaaattt gaacgccagt tcatggactc gtctactagt 60
gcagctggtt caggtagtgg ttcaggttct ggttcaggtt caggtagtgg ttcaggttct 120
ggttcaatga ccatgattac gaattcgatg tatagcaccg tgttttatac cagcgttcat 180
ccgagcacca gcgtgtttag ccgcaaacag ctgccgctgc tgattagcaa agattttccg 240
gccgaactgt atcatagcct gccgtgcaaa agcctggaaa atggtcatat taagaaagtg 300
aaaggcgtta aagccaccct ggccgaagca ccggccaccc cgaccgaaaa atctaacagc 360
gaagtgccgc agaaaaaact gaaagtgctg gttgccggcg gtggcattgg tggcctggtg 420
tttgcgctgg cggcgaaaaa gaaaggcttt gatgtgctgg tttttgaacg tgatctgagc 480
gccatccgcg gcgaaggcca gtatcgtggc ccgattcaga tccagagcaa tgccctggcg 540
gccctggaag cgattgatat ggatgtggcc gaagaaatca tgaacgcggg ttgcattacc 600
ggccagcgta tcaatggtct ggttgatggt attagcggca actggtattg taaatttgat 660
acctttaccc cggccgtgga acgcggcctg ccggtgaccc gtgttattag ccgcatgacc 720
ctgcagcaga tcctggcacg tgccgtgggt gaagatgtta ttatgaacga aagcaatgtg 780
gttaactttg aagatgatgg cgaaaaagtg accgtggttc tggaaaatgg tcagcgcttt 840
accggcgatc tgctggttgg tgccgatggc atccgtagca aagtgcgcac caacctgttt 900
ggtcatagcg aagcgaccta tagcggttat acctgttata ccggcattgc cgattttgtt 960
ccggcggata tcgataccgt gggttatcgt gtttttctgg gccataaaca gtattttgtg 1020
agcagcgatg ttggtggcgg taaaatgcag tggtatgcgt ttcataatga accggccggc 1080
ggtgtggatg ccccgaatgg caagaaagaa cgcctgctga aaatttttgg cggttggtgc 1140
gataatgtga tcgatctgct ggttgccacc gatgaagatg cgattctgcg tcgcgatatc 1200
tatgatcgtc cgccgacctt taattggggt cgtggtcgtg tgaccctgct gggtgatagc 1260
gttcatgcca tgcagccgaa cctgggtcag ggcggttgta tggccattga agatagctat 1320
cagctggcgc tggaactgga aaaagcctgg agccgcagcg ccgaaagcgg tagcccgatg 1380
gatgttatca gcagcctgcg tagctatgaa agcgcccgta aactgcgcgt gggtgttatt 1440
catggcctgg cccgtatggc cgcaatcatg gcaagcacct ataaagcgta tctgggcgtg 1500
ggtctgggtc cgctgagctt tctgaccaaa tatcgtattc cgcacccggg tcgtgttggc 1560
ggtcgtgtgt ttgttgatct gggcatgccg ctgatgctga gctgggtgct gggcggtaat 1620
ggtgataaac tggaaggccg tatccagcat tgccgcctga gcgaaaaagc caacgatcag 1680
ctgcgtcgct ggtttgaaga tgatgatgcg ctggaacgcg ccaccgatgc cgaatggctg 1740
ctgctgccgg ccgcaaatgg caatagcgcg ctggaaacca ttgtgctgag ccgtgatgaa 1800
gatgttccgt gcaccattgg tagcgtgagc cataccaaca tcccgggcaa aagcgtggtt 1860
ctgccgctgc cgcaggttag cgaaatgcat gcccagatta gctgtaaaaa caatgcgttt 1920
ttcgtgaccg attttcagag cgaacatggt acctgggtta tcgataatga aggccgtcgc 1980
tatcgcgtga gcccgaactt tccgatgcgt tttcatagca gcgatgtgat tgtttttggt 2040
agcgataaag cggcctttcg cgtgaaaacc atgaaatttc cgagcaaaac cgccgaagcg 2100
aaagaagaac gtaaagtggt tggcaccgcg taa 2133
<210> 12
<211> 710
<212> PRT
<213> 人工序列
<220>
<223> 经由接头导入了S-标签的CaZEP
<400> 12
Met Ser Lys Glu Thr Ala Ala Ala Lys Phe Glu Arg Gln Phe Met Asp
1 5 10 15
Ser Ser Thr Ser Ala Ala Gly Ser Gly Ser Gly Ser Gly Ser Gly Ser
20 25 30
Gly Ser Gly Ser Gly Ser Gly Ser Gly Ser Met Thr Met Ile Thr Asn
35 40 45
Ser Met Tyr Ser Thr Val Phe Tyr Thr Ser Val His Pro Ser Thr Ser
50 55 60
Val Phe Ser Arg Lys Gln Leu Pro Leu Leu Ile Ser Lys Asp Phe Pro
65 70 75 80
Ala Glu Leu Tyr His Ser Leu Pro Cys Lys Ser Leu Glu Asn Gly His
85 90 95
Ile Lys Lys Val Lys Gly Val Lys Ala Thr Leu Ala Glu Ala Pro Ala
100 105 110
Thr Pro Thr Glu Lys Ser Asn Ser Glu Val Pro Gln Lys Lys Leu Lys
115 120 125
Val Leu Val Ala Gly Gly Gly Ile Gly Gly Leu Val Phe Ala Leu Ala
130 135 140
Ala Lys Lys Lys Gly Phe Asp Val Leu Val Phe Glu Arg Asp Leu Ser
145 150 155 160
Ala Ile Arg Gly Glu Gly Gln Tyr Arg Gly Pro Ile Gln Ile Gln Ser
165 170 175
Asn Ala Leu Ala Ala Leu Glu Ala Ile Asp Met Asp Val Ala Glu Glu
180 185 190
Ile Met Asn Ala Gly Cys Ile Thr Gly Gln Arg Ile Asn Gly Leu Val
195 200 205
Asp Gly Ile Ser Gly Asn Trp Tyr Cys Lys Phe Asp Thr Phe Thr Pro
210 215 220
Ala Val Glu Arg Gly Leu Pro Val Thr Arg Val Ile Ser Arg Met Thr
225 230 235 240
Leu Gln Gln Ile Leu Ala Arg Ala Val Gly Glu Asp Val Ile Met Asn
245 250 255
Glu Ser Asn Val Val Asn Phe Glu Asp Asp Gly Glu Lys Val Thr Val
260 265 270
Val Leu Glu Asn Gly Gln Arg Phe Thr Gly Asp Leu Leu Val Gly Ala
275 280 285
Asp Gly Ile Arg Ser Lys Val Arg Thr Asn Leu Phe Gly His Ser Glu
290 295 300
Ala Thr Tyr Ser Gly Tyr Thr Cys Tyr Thr Gly Ile Ala Asp Phe Val
305 310 315 320
Pro Ala Asp Ile Asp Thr Val Gly Tyr Arg Val Phe Leu Gly His Lys
325 330 335
Gln Tyr Phe Val Ser Ser Asp Val Gly Gly Gly Lys Met Gln Trp Tyr
340 345 350
Ala Phe His Asn Glu Pro Ala Gly Gly Val Asp Ala Pro Asn Gly Lys
355 360 365
Lys Glu Arg Leu Leu Lys Ile Phe Gly Gly Trp Cys Asp Asn Val Ile
370 375 380
Asp Leu Leu Val Ala Thr Asp Glu Asp Ala Ile Leu Arg Arg Asp Ile
385 390 395 400
Tyr Asp Arg Pro Pro Thr Phe Asn Trp Gly Arg Gly Arg Val Thr Leu
405 410 415
Leu Gly Asp Ser Val His Ala Met Gln Pro Asn Leu Gly Gln Gly Gly
420 425 430
Cys Met Ala Ile Glu Asp Ser Tyr Gln Leu Ala Leu Glu Leu Glu Lys
435 440 445
Ala Trp Ser Arg Ser Ala Glu Ser Gly Ser Pro Met Asp Val Ile Ser
450 455 460
Ser Leu Arg Ser Tyr Glu Ser Ala Arg Lys Leu Arg Val Gly Val Ile
465 470 475 480
His Gly Leu Ala Arg Met Ala Ala Ile Met Ala Ser Thr Tyr Lys Ala
485 490 495
Tyr Leu Gly Val Gly Leu Gly Pro Leu Ser Phe Leu Thr Lys Tyr Arg
500 505 510
Ile Pro His Pro Gly Arg Val Gly Gly Arg Val Phe Val Asp Leu Gly
515 520 525
Met Pro Leu Met Leu Ser Trp Val Leu Gly Gly Asn Gly Asp Lys Leu
530 535 540
Glu Gly Arg Ile Gln His Cys Arg Leu Ser Glu Lys Ala Asn Asp Gln
545 550 555 560
Leu Arg Arg Trp Phe Glu Asp Asp Asp Ala Leu Glu Arg Ala Thr Asp
565 570 575
Ala Glu Trp Leu Leu Leu Pro Ala Ala Asn Gly Asn Ser Ala Leu Glu
580 585 590
Thr Ile Val Leu Ser Arg Asp Glu Asp Val Pro Cys Thr Ile Gly Ser
595 600 605
Val Ser His Thr Asn Ile Pro Gly Lys Ser Val Val Leu Pro Leu Pro
610 615 620
Gln Val Ser Glu Met His Ala Gln Ile Ser Cys Lys Asn Asn Ala Phe
625 630 635 640
Phe Val Thr Asp Phe Gln Ser Glu His Gly Thr Trp Val Ile Asp Asn
645 650 655
Glu Gly Arg Arg Tyr Arg Val Ser Pro Asn Phe Pro Met Arg Phe His
660 665 670
Ser Ser Asp Val Ile Val Phe Gly Ser Asp Lys Ala Ala Phe Arg Val
675 680 685
Lys Thr Met Lys Phe Pro Ser Lys Thr Ala Glu Ala Lys Glu Glu Arg
690 695 700
Lys Val Val Gly Thr Ala
705 710
Claims (12)
1.一种转化体,其特征在于,在宿主细胞中导入有:
第一启动子、及与所述第一启动子可操作地连接的类胡萝卜素的生物合成基因的上游基因;以及
启动子强度比所述第一启动子高的第二启动子、及与所述第二启动子可操作地连接的ZEP基因和CCS基因,
所述上游基因包含crtY和crtZ。
2.根据权利要求1所述的转化体,其中,所述第一启动子与所述第二启动子的启动子强度之差为类胡萝卜素的生物合成基因固有的启动子与PBAD的启动子强度之差以上。
3.根据权利要求1或2所述的转化体,其中,所述上游基因还包含crtI;crtI和crtB;或crtI、crtB和crtE。
4.根据权利要求1~3中任一项所述的转化体,其中,所述第一启动子是Plac或类胡萝卜素的生物合成基因固有的启动子。
5.根据权利要求1~4中任一项所述的转化体,其中,所述第二启动子是PBAD或Ptac。
6.根据权利要求1~5中任一项所述的转化体,其中,在所述宿主细胞中进一步导入有S蛋白基因、S标签基因和接头基因,
所述ZEP基因和所述CCS基因分别与所述S标签基因连接,
所述接头基因连接在所述ZEP基因与所述S标签基因之间、和/或、所述CCS基因与所述S标签基因之间。
7.根据权利要求6所述的转化体,其中,所述S标签基因编码的S标签为以下的(1)和(2)中的任一种多肽,
(1)包含编码序列号4所示的氨基酸序列的碱基序列的多肽;
(2)相对于序列号4所示的氨基酸序列的第12位以外的序列一致性为90%以上、且与S蛋白特异性结合的多肽。
8.根据权利要求1~7中任一项所述的转化体,其中,所述接头基因至少连接在所述ZEP基因与所述S标签基因之间。
9.根据权利要求1~8中任一项所述的转化体,其中,所述接头基因连接在所述ZEP基因与所述S标签基因之间,而不在所述CCS基因与所述S标签基因之间。
10.一种类胡萝卜素组合物的制造方法,其特征在于,包括以下工序:培养权利要求1~9中任一项所述的转化体。
11.根据权利要求10所述的制造方法,其中,所述类胡萝卜素组合物包含辣椒红色素。
12.根据权利要求10或11所述的制造方法,其中,所述类胡萝卜素组合物包含辣椒玉红素、辣椒红色素3′-乙酸酯、辣椒玉红素3-乙酸酯、类胡萝卜黄色素A、辣椒玉红素二乙酸酯、辣椒红色素3,6-环氧化物、和/或、辣椒红色素3,6-环氧化物3′-乙酸酯。
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