CN113354745B - 一种组合物及规模化生产成纤维细胞生长因子的方法 - Google Patents
一种组合物及规模化生产成纤维细胞生长因子的方法 Download PDFInfo
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Abstract
本发明涉及一种组合物及规模化生产成纤维细胞生长因子的方法。通过将成纤维细胞生长因子与绿色荧光蛋白和小泛素样修饰因子融合实现成纤维细胞生长因子的可溶性大量表达，然后利用三(2‑羰基乙基)磷盐酸盐和硫酸锌的协同作用在促进在冷激启动子的作用下表达的小泛素样修饰因子蛋白酶对融合蛋白进行胞内快速的水解，简化了生产步骤，从而实现了从毫克级到公斤级规模生产具有生物学活性的成纤维细胞生长因子。
Description
技术领域
本发明涉及一种用于发酵培养及纯化成纤维细胞生长因子的组合物，以及含有成纤维细胞生长因子的融合蛋白，特别是规模化培养并获得成纤维细胞生长因子的方法。
背景技术
成纤维细胞生长因子(Fibroblast growth factors,FGFs)超家族包括22个成员，主要分为分泌型和胞内型。分泌型成纤维细胞生长因子(eFGF)包括成纤维细胞生长因子1-10及成纤维细胞生长因子16-23，胞内型成纤维细胞生长因子(iFGF)包括成纤维细胞生长因子11-14。eFGF必需通过结合及活化细胞表面酪氨酸激酶(receptor tyrosine kinases，RTKs)家族成员的成纤维细胞生长因子受体(FGFRs)进而激活下游信号通路，而iFGF则主要与其它蛋白相互作用进而调控细胞钠通道开关。在人体中，成纤维细胞生长因子超家族几乎在所有组织中都有表达，其对胚胎发育、器官形成和再生、组织修复以及机体的新陈代谢都发挥着极为重要的生物学功能。
人和小鼠的成纤维细胞生长因子超家族都包括22个成员，而成纤维细胞生长因子-15与成纤维细胞生长因子-19在脊椎动物中是同源基因，成纤维细胞生长因子-15尚未在人类基因组中发现，而成纤维细胞生长因子-19则相反，在大小鼠体内没有被发现。
利用基因工程技术获得大量有活性的成纤维细胞生长因子是生产成纤维细胞生长因子相关药物的必要条件。相对于酵母、昆虫细胞以及哺乳动物细胞等真核表达***而言，利用原核***(尤其是大肠杆菌)表达重组蛋白，其所需时间、生产成本及场地要求等都是最简便和经济的。然而由于成纤维细胞因子在原核***中表达量低、形成包涵体或对宿主有毒性等问题导致难以规模化生产。此外，尽管可以通过分子伴侣融合的方式增加表达量，但会出现冗余的氨基酸，导致分离纯化工艺难度加大以及生产成本急剧提升。以上这些都对成纤维细胞生长因子成药性造成严重的影响。
发明内容
本发明之一提供了一种组合物，其包括三(2-羰基乙基)磷盐酸盐和硫酸锌。
在一个具体实施方式中，所述三(2-羰基乙基)磷盐酸盐和所述硫酸锌的质量比是1:(1至50)。
本发明之二提供了如本发明之一所述的组合物在生产成纤维细胞生长因子中应用，特别是规模化生产成纤维细胞生长因子中的应用。
本发明之三提供了一种含有成纤维细胞生长因子的融合蛋白，其包括从N端到C端的可视化报告蛋白、小泛素样修饰因子(Small Ubiquitin-like Modifier，SUMO)和成纤维细胞生长因子。
在一个具体实施方式中，所述成纤维细胞生长因子为人源成纤维细胞生长因子-1、人源成纤维细胞生长因子-2、人源成纤维细胞生长因子-3、人源成纤维细胞生长因子-4、人源成纤维细胞生长因子-5、人源成纤维细胞生长因子-6、人源成纤维细胞生长因子-7、人源成纤维细胞生长因子-8、人源成纤维细胞生长因子-9、人源成纤维细胞生长因子-10、人源成纤维细胞生长因子-11、人源成纤维细胞生长因子-12、人源成纤维细胞生长因子-13、人源成纤维细胞生长因子-14、人源成纤维细胞生长因子-16、人源成纤维细胞生长因子-17、人源成纤维细胞生长因子-18、人源成纤维细胞生长因子-19、人源成纤维细胞生长因子-20、人源成纤维细胞生长因子-21、人源成纤维细胞生长因子-22、人源成纤维细胞生长因子-23中的一种。
在一个具体实施方式中，所述人源成纤维细胞生长因子-1的氨基酸序列如SEQ IDNo.1所示，所述人源成纤维细胞生长因子-2的氨基酸序列如SEQ ID No.2所示，所述人源成纤维细胞生长因子-3的氨基酸序列如SEQ ID No.3所示，所述人源成纤维细胞生长因子-4的氨基酸序列如SEQ ID No.4所示，所述人源成纤维细胞生长因子-5的氨基酸序列如SEQID No.5所示，所述人源成纤维细胞生长因子-6的氨基酸序列如SEQ ID No.6所示，所述人源成纤维细胞生长因子-7的氨基酸序列如SEQ ID No.7所示，所述人源成纤维细胞生长因子-8的氨基酸序列如SEQ ID No.8所示，所述人源成纤维细胞生长因子-9的氨基酸序列如SEQ ID No.9所示，所述人源成纤维细胞生长因子-10的氨基酸序列如SEQ ID No.10所示，所述人源成纤维细胞生长因子-11的氨基酸序列如SEQ ID No.11所示，所述人源成纤维细胞生长因子-12的氨基酸序列如SEQ ID No.12所示，所述人源成纤维细胞生长因子-13的氨基酸序列如SEQ ID No.13所示，所述人源成纤维细胞生长因子-14的氨基酸序列如SEQ IDNo.14所示，所述人源成纤维细胞生长因子-16的氨基酸序列如SEQ ID No.15所示，所述人源成纤维细胞生长因子-17的氨基酸序列如SEQ ID No.16所示，所述人源成纤维细胞生长因子-18的氨基酸序列如SEQ ID No.17所示，所述人源成纤维细胞生长因子-19的氨基酸序列如SEQ ID No.18所示，所述人源成纤维细胞生长因子-20的氨基酸序列如SEQ ID No.19所示，所述人源成纤维细胞生长因子-21的氨基酸序列如SEQ ID No.20所示，所述人源成纤维细胞生长因子-22的氨基酸序列如SEQ ID No.21所示，所述人源成纤维细胞生长因子-23的氨基酸序列如SEQ ID No.22所示。
在一个具体实施方式中，所述可视化报告蛋白为荧光蛋白。
在一个具体实施方式中，所述可视化报告蛋白为绿色荧光蛋白、红色荧光蛋白、蓝色荧光蛋白和青色荧光蛋白中的一种；优选地，所述绿色荧光蛋白的氨基酸序列如SEQ IDNo.23所示。
在一个具体实施方式中，所述小泛素样修饰因子的氨基酸序列如SEQ ID No.24所示。
在一个具体实施方式中，所述可视化报告蛋白和所述小泛素样修饰因子之间由多肽接头连接。
在一个具体实施方式中，所述多肽接头的氨基酸序列如SEQ ID No.25所示。
本发明之四提供了一种用于表达如本发明之三中任意一项所述的融合蛋白的表达盒，其包括从5’端到3’端的第一启动子、编码所述融合蛋白的核酸和第一终止子。其中，在所述第一启动子的下游和所述编码所述融合蛋白的核酸的上游还包括RBS，所述RBS的核苷酸序列如SEQ ID No.53所示。在所述第一启动子的下游和所述编码所述融合蛋白的核酸的上游还可以包括酶切位点。其中，酶切位点位于所述RBS的下游。
在一个具体实施方式中，所述第一启动子为T7启动子，所述第一终止子为T7终止子。
在一个具体实施方式中，编码所述人源成纤维细胞生长因子-1的核苷酸如SEQ IDNo.26所示，编码所述人源成纤维细胞生长因子-2的核苷酸序列如SEQ ID No.27所示，编码所述人源成纤维细胞生长因子-3的核苷酸序列如SEQ ID No.28所示，编码所述人源成纤维细胞生长因子-4的核苷酸序列如SEQ ID No.29所示，编码所述人源成纤维细胞生长因子-5的核苷酸序列如SEQ ID No.30所示，编码所述人源成纤维细胞生长因子-6的核苷酸序列如SEQ ID No.31所示，编码所述人源成纤维细胞生长因子-7的核苷酸序列如SEQ ID No.32所示，编码所述人源成纤维细胞生长因子-8的核苷酸序列如SEQ ID No.33所示，编码所述人源成纤维细胞生长因子-9的核苷酸序列如SEQ ID No.34所示，编码所述人源成纤维细胞生长因子-10的核苷酸序列如SEQ ID No.35所示，编码所述人源成纤维细胞生长因子-11的核苷酸序列如SEQ ID No.36所示，编码所述人源成纤维细胞生长因子-12的核苷酸序列如SEQID No.37所示，编码所述人源成纤维细胞生长因子-13的核苷酸序列如SEQ ID No.38所示，编码所述人源成纤维细胞生长因子-14的核苷酸序列如SEQ ID No.39所示，编码所述人源成纤维细胞生长因子-16的核苷酸序列如SEQ ID No.40所示，编码所述人源成纤维细胞生长因子-17的核苷酸序列如SEQ ID No.41所示，编码所述人源成纤维细胞生长因子-18的核苷酸序列如SEQ ID No.42所示，编码所述人源成纤维细胞生长因子-19的核苷酸序列如SEQID No.43所示，编码所述人源成纤维细胞生长因子-20的核苷酸序列如SEQ ID No.44所示，编码所述人源成纤维细胞生长因子-21的核苷酸序列如SEQ ID No.45所示，编码所述人源成纤维细胞生长因子-22的核苷酸序列如SEQ ID No.46所示，编码所述人源成纤维细胞生长因子-23的核苷酸序列如SEQ ID No.47所示。
在一个具体实施方式中，编码所述可视化报告蛋白的核苷酸序列如SEQ ID No.48所示。
在一个具体实施方式中，编码所述小泛素样修饰因子的核苷酸序列如SEQ IDNo.49所示。
在一个具体实施方式中，编码所述多肽接头的核苷酸序列如SEQ ID No.50所示。
本发明之五提供了一套用于表达如本发明之三中任意一项所述的融合蛋白或用于表达如本发明之三中任意一项所述的融合蛋白并纯化所述成纤维细胞生长因子的表达盒，其包括如本发明之四所述的表达盒和用于表达小泛素样修饰因子蛋白酶(SUMO蛋白酶，例如Ulp1)的表达盒。
在一个具体实施方式中，所述用于表达小泛素样修饰因子蛋白酶的表达盒包括从5’端到3’端的第二启动子、3’-UTR、编码小泛素样修饰因子蛋白酶的核酸和5’-UTR。
在一个具体实施方式中，所述第二启动子为CspA启动子，所述3’-UTR为CspA 3’-UTR，所述5’-UTR为CspA5’-UTR。
在一个具体实施方式中，所述小泛素样修饰因子蛋白酶的氨基酸序列如SEQ IDNo.51所示。
在一个具体实施方式中，编码小泛素样修饰因子蛋白酶的核苷酸序列如SEQ IDNo.52所示。
本发明之六提供了一种获得如本发明之三中任意一项所述的融合蛋白或其中的所述成纤维细胞生长因子的方法，其包括如下步骤：
1-1)将如本发明之四所述的表达盒或编码所述融合蛋白的核酸连接到第一表达载体上，得到第一重组表达载体；其中，所述第一表达载体为异丙基-β-D-硫代半乳糖苷(IPTG)诱导型的表达载体；
2-1)将如本发明之五所述的表达盒中的所述用于表达小泛素样修饰因子蛋白酶的表达盒或编码小泛素样修饰因子蛋白酶的核酸连接到所述第一重组表达载体上，得到第二重组表达载体；
3-1)将所述第二重组表达载体转化至第一宿主中，得到第一重组生物体；
4-1)在第一培养液中培养所述第一重组生物体至表达有所述融合蛋白和所述小泛素样修饰因子蛋白酶，得到第一培养物；
5-1)纯化所述第一培养物从而得到所述成纤维细胞生长因子；
或者
1-2)同1-1)；
2-2)将如本发明之五所述的表达盒中的所述用于表达小泛素样修饰因子蛋白酶的表达盒或编码小泛素样修饰因子蛋白酶的核酸连接到所述第二表达载体上，得到第三重组表达载体；
3-2)将所述第一重组表达载体转化至第二宿主中，得到第二重组生物体；将所述第三重组表达载体转化至第三宿主中，得到第三重组生物体；
4-2)在第二培养液培养所述第二重组生物体至表达有所述融合蛋白，得到第二培养物；在第三培养液培养所述第三重组生物体至表达有所述小泛素样修饰因子蛋白酶，得到第三培养物；将所述第二培养物与所述第三培养物混合，得到第四培养物；
5-2)纯化所述第四培养物，从而得到所述成纤维细胞生长因子；
或者
1-3)同1-1)；
2-3)同3-2)中的将所述第一重组表达载体转化至第二宿主中，得到第二重组生物体；
3-3)在第二培养液培养所述第二重组生物体至表达有所述融合蛋白，得到第二培养物；
4-3)纯化所述第二培养物，从而得到所述融合蛋白。
在一个具体实施方式中，所述第一培养液、所述第二培养液和所述第三培养液独立地为LB培养液或含葡萄糖的LB培养液。
在一个具体实施方式中，所述第一表达载体可以为pET系列的表达载体中的一种。
在一个具体实施方式中，所述第一表达载体为pET21a、pET20b和pET28a中的一种；
所述第二表达载体为pCold，其可购自大连宝生物工程有限公司(货号：3360)；
所述第一宿主、所述第二宿主和所述第三宿主独立地为大肠杆菌(Escherichiacoli)；优选地，所述大肠杆菌的菌株为BL21(DE3)。
在一个具体实施方式中，在步骤4-2)或步骤3-3)中，培养所述第二重组生物体的温度为28至37℃，时间为4至8小时；
在步骤4-1)中，在28至37℃的温度下，在异丙基-β-D-硫代半乳糖苷(IPTG)的诱导下培养所述第一重组生物体4至8小时，然后在10至15℃的温度下继续培养所述第一重组生物体15至60分钟；
在步骤4-2)中，培养所述第三重组生物体的温度为10至15℃，时间为15至60分钟。
在一个具体实施方式中，在步骤4-2)或步骤3-3)中，培养所述第二重组生物体的温度为32至37℃，时间为4至6小时；
在步骤4-1)中，在32至37℃的温度下，在IPTG的诱导下培养所述第一重组生物体4至6小时，然后在10至15℃的温度下继续培养所述第一重组生物体15至30分钟；
在步骤4-2)中，培养所述第三重组生物体的温度为10至15℃，时间为15至30分钟。
在一个具体实施方式中，在步骤4-1)中，在培养温度降至所述10至15℃时，在所述第一培养液中加入如本发明之一所述的组合物。
在一个具体实施方式中，所述三(2-羰基乙基)磷盐酸盐在所述第一培养液中的终浓度为0.1至1毫摩尔/升；所述硫酸锌所述第一培养液中的终浓度为1至5毫摩尔/升。
在一个具体实施方式中，在步骤4-2)中，在所述第二培养液中加入如本发明之一所述的组合物。
在一个具体实施方式中，所述三(2-羰基乙基)磷盐酸盐在所述第二培养液中的终浓度为0.1至1毫摩尔/升；所述硫酸锌所述第二培养液中的终浓度为1至5毫摩尔/升。
此外，在一个具体实施方式中，对纯化的成纤维细胞生长因子-1、成纤维细胞生长因子-2、成纤维细胞生长因子-3、成纤维细胞生长因子-4、成纤维细胞生长因子-5、成纤维细胞生长因子-6、成纤维细胞生长因子-7、成纤维细胞生长因子-10、成纤维细胞生长因子-11、成纤维细胞生长因子-12、成纤维细胞生长因子-13、成纤维细胞生长因子-14、成纤维细胞生长因子-16、成纤维细胞生长因子-18、成纤维细胞生长因子-19、成纤维细胞生长因子-20、成纤维细胞生长因子-21、成纤维细胞生长因子-22和成纤维细胞生长因子-23的活性进行了相应的分析，结果显示它们均表现出了各自应有的活性。
本发明的有益效果：
针对现有技术中原核表达所遇到的问题并为了进一步提高成纤维细胞生长因子表达过程中质量控制精度，本发明提供了一种组合物，并设计了与所述组合物配合的含有两种不同诱导表达方式基因表达盒的重组大肠杆菌表达质粒，其中一个表达盒用于表达含有成纤维细胞生长因子的融合蛋白，将成纤维细胞生长因子与荧光蛋白和SUMO融合有利于可溶性融合蛋白的大量表达，同时利用例如绿色荧光蛋白产生的绿色荧光可以对蛋白的表达进行可视化监测；另一个则用于表达小泛素样修饰因子蛋白酶。在融合蛋白诱导结束后，进入低温培养过程时添加本发明的化合物组合，可实现小泛素样修饰因子蛋白酶在被诱导表达的同时对融合蛋白进行水解效应，从而达到可溶性的、与天然成纤维细胞生长因子氨基酸序列完全一致且具有相应生物学功能的成纤维细胞生长因子规模化生产的目的，采用本发明的方法，不但可以获得具生物学功能的成纤维细胞生长因子，且大大简化纯化步骤，显著降低生产成本，因而能满足药品、日用品和科研试剂生产的需要。
本发明的优点在于：1)通过目视绿色荧光，可以实时评估或监测重组融合蛋白的可溶性表达及表达量，而为了达到上述目的，采用的常规方法至少需要6个小时以上；2)与其它蛋白酶识别位点不同，SUMO蛋白酶识别的是SUMO的三维结构，因此，不存在乱切割(水解)的情况；此外，利用SUMO蛋白酶水解时也不会使目的蛋白序列内带入任何冗余的氨基酸，因而可以获得与天然蛋白序列一致的成纤维细胞生长因子；3)利用本发明特有的融合蛋白的方式使得在大肠杆菌宿主中难表达(difficult-to-express)的成纤维细胞生长因子实现可溶性表达，而且还首次通过本发明的组合物间的协同增效作用实现了胞内促SUMO蛋白酶表达的同时对融合蛋白进行快速水解，因而大大简化成纤维细胞生长因子的生产和纯化过程。相对于常规方法所需要的时间，本发明可将纯化时间缩短至少8至48小时；4)采用本发明的工艺，无需复杂的操作工艺和特殊的器械，只需扩大生产规模，例如发酵规模、菌体破碎及离心器械的规模、凝胶层析规模等，即可实现从毫克级到公斤级成纤维细胞生长因子的生产。
附图说明
图1 pET-GSFGF9-CspA-Ulp1重组质粒图谱示意图
图2 pET-GSFGF17-CspA-Ulp1/BL21诱导表达后的荧光观察
A.发酵液在自然光下的观察结果，其中左侧试管(pET-GSFGF17-CspA-Ulp1/BL21IPTG诱导表达后)中的发酵液显示出绿色荧光，右侧试管(pET-GSFGF17-CspA-Ulp1/BL21IPTG诱导表达前)的发酵液不显示绿色荧光；B.pET-GSFGF17-CspA-Ulp1/BL21 IPTG诱导表达后的发酵液在紫外灯光下的观察结果。
图3.融合蛋白GFP-SUMO-FGFn表达分析
A.泳道1：IPTG诱导后的pET-GSFGF8-CspA-Ulp1/BL21总蛋白；泳道2：IPTG诱导后的pET-GSFGF9-CspA-Ulp1/BL21总蛋白；泳道3：IPTG诱导后的pET-GFP-SUMO/BL21总蛋白；泳道M：蛋白质分子量标记；泳道4：IPTG诱导前的pET-GSFGF8-CspA-Ulp1/BL21总蛋白；泳道5：IPTG诱导后的pET-GSFGF3-CspA-Ulp1/BL21总蛋白；泳道6：IPTG诱导后的pET-GSFGF16-CspA-Ulp1/BL21总蛋白；泳道7：IPTG诱导后的pET-GSFGF17-CspA-Ulp1/BL21总蛋白；泳道8：IPTG诱导后的pET-GSFGF19-CspA-Ulp1/BL21总蛋白；泳道9：IPTG诱导后的pET-GSFGF20-CspA-Ulp1/BL21总蛋白；
B.泳道1：IPTG诱导后的pET-GSFGF4-CspA-Ulp1/BL21总蛋白；泳道2：IPTG诱导后的pET-GSFGF6-CspA-Ulp1/BL21总蛋白；泳道3：IPTG诱导后的pET-GSFGF7-CspA-Ulp1/BL21总蛋白；泳道4：IPTG诱导后的pET-GFP-SUMO/BL21总蛋白；泳道M：蛋白质分子量标记；泳道5：IPTG诱导前的pET-GSFGF4-CspA-Ulp1/BL21总蛋白；泳道6：IPTG诱导后的pET-GSFGF10-CspA-Ulp1/BL21总蛋白；泳道7：IPTG诱导后的pET-GSFGF18-CspA-Ulp1/BL21总蛋白；泳道8：IPTG诱导后的pET-GSFGF21-CspA-Ulp1/BL21总蛋白；泳道9：IPTG诱导后的pET-GSFGF22-CspA-Ulp1/BL21总蛋白；
C.泳道1：IPTG诱导后的pET-GSFGF11-CspA-Ulp1/BL21总蛋白；泳道2：IPTG诱导后的pET-GSFGF23-CspA-Ulp1/BL21总蛋白；泳道3：IPTG诱导后的pET-GSFGF5-CspA-Ulp1/BL21总蛋白；泳道4：IPTG诱导后的pET-GFP-SUMO/BL21总蛋白；泳道M：蛋白质分子量标记；泳道5：IPTG诱导前的pET-GSFGF11-CspA-Ulp1/BL21总蛋白；泳道6：IPTG诱导后的pET-GSFGF12-CspA-Ulp1/BL21总蛋白；泳道7：IPTG诱导后的pET-GSFGF13-CspA-Ulp1/BL21总蛋白；泳道8：IPTG诱导后的pET-GSFGF14-CspA-Ulp1/BL21总蛋白。
图4 IPTG诱导表达后的融合蛋白GFP-SUMO-FGFn的可溶性分析
A：泳道1：pET-GSFGF8-CspA-Ulp1/BL21总蛋白；泳道2：pET-GSFGF8-CspA-Ulp1/BL21上清液中的蛋白；泳道3：pET-GSFGF8-CspA-Ulp1/BL21沉淀中的蛋白；泳道M：蛋白质分子量标记；泳道4：pET-GSFGF5-CspA-Ulp1/BL21总蛋白；泳道5：pET-GSFGF5-CspA-Ulp1/BL21上清液中的蛋白；泳道6：pET-GSFGF5-CspA-Ulp1/BL21沉淀中的蛋白；泳道7：pET-GSFGF23-CspA-Ulp1/BL21总蛋白；泳道8：pET-GSFGF23-CspA-Ulp1/BL21上清液中的蛋白；泳道9：pET-GSFGF23-CspA-Ulp1/BL21沉淀中的蛋白；
B：泳道1：pET-GSFGF4-CspA-Ulp1/BL21总蛋白；泳道2：pET-GSFGF4-CspA-Ulp1/BL21上清液中的蛋白；泳道3：pET-GSFGF4-CspA-Ulp1/BL21沉淀中的蛋白；泳道M：蛋白质分子量标记；泳道4：pET-GSFGF16-CspA-Ulp1/BL21总蛋白；泳道5：pET-GSFGF16-CspA-Ulp1/BL21上清液中的蛋白；泳道6：pET-GSFGF16-CspA-Ulp1/BL21沉淀中的蛋白；泳道7：pET-GSFGF14-CspA-Ulp1/BL21总蛋白样品；泳道8：pET-GSFGF14-CspA-Ulp1/BL21上清液中的蛋白；泳道9：pET-GSFGF14-CspA-Ulp1/BL21沉淀中的蛋白；
C：泳道1：pET-GSFGF11-CspA-Ulp1/BL21总蛋白；泳道2：pET-GSFGF11-CspA-Ulp1/BL21上清液中的蛋白；泳道3：pET-GSFGF11-CspA-Ulp1/BL21沉淀中的蛋白；泳道4：pET-GSFGF12-CspA-Ulp1/BL21总蛋白；泳道5：pET-GSFGF12-CspA-Ulp1/BL21上清液中的蛋白；泳道6：pET-GSFGF12-CspA-Ulp1/BL21沉淀中的蛋白；泳道M：蛋白质分子量标记；
D：泳道1：pET-GSFGF19-CspA-Ulp1/BL21总蛋白；泳道2：pET-GSFGF19-CspA-Ulp1/BL21上清液中的蛋白；泳道3：pET-GSFGF19-CspA-Ulp1/BL21沉淀中的蛋白；泳道M：蛋白质分子量标记；泳道4：pET-GSFGF17-CspA-Ulp1/BL21总蛋白；泳道5：pET-GSFGF17-CspA-Ulp1/BL21上清液中的蛋白；泳道6：pET-GSFGF17-CspA-Ulp1/BL21沉淀中的蛋白。
图5.三(2-羰基乙基)磷盐酸盐、硫酸锌及组合对小泛素样修饰因子蛋白酶水解融合蛋白的功能分析
黑色箭头指示为融合蛋白GFP-SUMO-FGF12；灰色箭头指示为融合蛋白GFP-SUMO-FGF12水解后的GFP-SUMO；中空箭头指示为融合蛋白GFP-SUMO-FGF12水解后的成纤维细胞生长因子-12。
A：摇瓶1和摇瓶I，15℃低温诱导起始时添加终浓度为5毫摩尔/升硫酸锌对小泛素样修饰因子蛋白酶的水解融合蛋白的影响
泳道1：pET-GSFGF12-CspA-Ulp1/BL21低温诱导30分钟；泳道2：pET-GSFGF12-CspA-Ulp1/BL21低温诱导60分钟；泳道3：pET-GSFGF12-CspA-Ulp1/BL21低温诱导120分钟；泳道4：pET-GFP-SUMO/BL21低温诱导120分钟；
B：摇瓶2，15℃低温诱导起始时添加终浓度为1毫摩尔/升三(2-羰基乙基)磷盐酸盐对小泛素样修饰因子蛋白酶的水解融合蛋白的影响
泳道1：低温诱导0分钟；泳道2：低温诱导15分钟；泳道3：低温诱导30分钟；泳道4：低温诱导60分钟；泳道5：低温诱导90分钟；泳道6：低温诱导120分钟；
C：摇瓶3，15℃低温诱导起始时添加终浓度为1毫摩尔/升三(2-羰基乙基)磷盐酸盐和1毫摩尔/升硫酸锌组合对小泛素样修饰因子蛋白酶的水解融合蛋白的影响
泳道1：低温诱导0分钟；泳道2：低温诱导15分钟；泳道3：低温诱导30分钟；泳道4：低温诱导60分钟；泳道5：低温诱导90分钟；泳道6：低温诱导120分钟；
D：摇瓶4，15℃低温诱导起始时添加终浓度为1毫摩尔/升三(2-羰基乙基)磷盐酸盐和3毫摩尔/升硫酸锌组合对小泛素样修饰因子蛋白酶的水解融合蛋白的影响
泳道1：低温诱导0分钟；泳道2：低温诱导15分钟；泳道3：低温诱导30分钟；泳道4：低温诱导60分钟；泳道5：低温诱导90分钟；泳道6：低温诱导120分钟；
E：摇瓶5，15℃低温诱导起始时添加终浓度为0.1毫摩尔/升三(2-羰基乙基)磷盐酸盐和5毫摩尔/升硫酸锌组合对小泛素样修饰因子蛋白酶水解融合蛋白的影响
泳道1：低温诱导0分钟；泳道2：低温诱导15分钟；泳道3：低温诱导30分钟；泳道4：低温诱导60分钟；泳道5：低温诱导90分钟；泳道6：低温诱导120分钟。
图6 三(2-羰基乙基)磷盐酸盐和硫酸锌组合促进小泛素样修饰因子蛋白酶水解其它融合蛋白GFP-SUMO-FGFn
15℃低温诱导过程中添加终浓度为0.1毫摩尔/升三(2-羰基乙基)磷盐酸盐和5毫摩尔/升硫酸锌，观察小泛素样修饰因子蛋白酶的对GFP-SUMO-FGFn融合蛋白水解能力。
泳道M：蛋白质分子量标记；泳道1：pET-GSFGF9-CspA-Ulp1/BL21低温诱导0分钟；泳道2：pET-GSFGF9-CspA-Ulp1/BL21低温诱导30分钟；泳道3：pET-GSFGF8-CspA-Ulp1/BL21低温诱导30分钟；泳道4：pET-GSFGF16-CspA-Ulp1/BL21低温诱导30分钟；泳道5：pET-GSFGF17-CspA-Ulp1/BL21低温诱导30分钟；泳道6：pET-GSFGF19-CspA-Ulp1/BL21低温诱导30分钟。
图7 水解前后的融合蛋白GFP-SUMO-FGFn的杂交分析
实心箭头表示融合蛋白GFP-SUMO-FGFn；空心箭头表示成纤维细胞生长因子-n。
A：泳道M：蛋白质分子量标记；泳道1：pET-GSFGF9-CspA-Ulp1/BL21 IPTG诱导前；泳道2：pET-GSFGF9-CspA-Ulp1/BL21低温诱导30分钟；泳道3：pET-GSFGF9-CspA-Ulp1/BL21低温诱导起始即0分钟；
B：泳道M：蛋白质分子量标记；泳道1：pET-GSFGF18-CspA-Ulp1/BL21低温诱导30分钟；泳道2：pET-GSFGF18-CspA-Ulp1/BL21低温诱导5分钟；泳道3：pET-GSFGF18-CspA-Ulp1/BL21低温诱导起始即0分钟。
图8 成纤维细胞生长因子-17(FGF-17)的纯化过程分析
A：CM离子层析凝胶分离FGF-17，其中，泳道1：含0.1M氯化钠的缓冲液1；泳道2：含0.3M氯化钠的缓冲液2：泳道3和4：含0.6M氯化钠的缓冲液3；泳道4：含1.0摩尔/升氯化钠的缓冲液6；泳道M：蛋白质分子量标记；
B：肝素亲和层析凝胶纯化FGF-17，其中，泳道1和2：含1.2摩尔/升氯化钠的缓冲液4。
图9 成纤维细胞生长因子促***细胞增殖的活性分析
具体实施方式
以下通过优选的实施案例的形式对本发明的上述内容作进一步的详细说明，但其不构成对本发明的限制。
如无特别说明，本发明的实施例中的试剂、酶类等均可通过商业途径购买。
大肠杆菌pET-21a表达载体(货号：69740)以及大肠杆菌BL21(DE3)(货号：69450)购自Sigma-Aldrich公司。
pCold(货号3360)购自宝生物工程(大连)有限公司。
纯化融合蛋白所用到的肝素亲和层析凝胶(货号：17-0998-03)和离子层析凝胶(CM Sepharose Fast Flow，货号：17-0719-01；Q Sepharose Fast Flow，货号：17-0510-01)购自美国GE公司。
人源成纤维细胞生长因子-1的氨基酸序列如SEQ ID No.1所示，人源成纤维细胞生长因子-2的氨基酸序列如SEQ ID No.2所示，人源成纤维细胞生长因子-3的氨基酸序列如SEQ ID No.3所示，人源成纤维细胞生长因子-4的氨基酸序列如SEQ ID No.4所示，人源成纤维细胞生长因子-5的氨基酸序列如SEQ ID No.5所示，人源成纤维细胞生长因子-6的氨基酸序列如SEQ ID No.6所示，人源成纤维细胞生长因子-7的氨基酸序列如SEQ ID No.7所示，人源成纤维细胞生长因子-8的氨基酸序列如SEQ ID No.8所示，人源成纤维细胞生长因子-9的氨基酸序列如SEQ ID No.9所示，人源成纤维细胞生长因子-10的氨基酸序列如SEQ ID No.10所示，人源成纤维细胞生长因子-11的氨基酸序列如SEQ ID No.11所示，人源成纤维细胞生长因子-12的氨基酸序列如SEQ ID No.12所示，人源成纤维细胞生长因子-13的氨基酸序列如SEQ ID No.13所示，人源成纤维细胞生长因子-14的氨基酸序列如SEQ IDNo.14所示，人源成纤维细胞生长因子-16的氨基酸序列如SEQ ID No.15所示，人源成纤维细胞生长因子-17的氨基酸序列如SEQ ID No.16所示，人源成纤维细胞生长因子-18的氨基酸序列如SEQ ID No.17所示，人源成纤维细胞生长因子-19的氨基酸序列如SEQ ID No.18所示，人源成纤维细胞生长因子-20的氨基酸序列如SEQ ID No.19所示，人源成纤维细胞生长因子-21的氨基酸序列如SEQ ID No.20所示，人源成纤维细胞生长因子-22的氨基酸序列如SEQ ID No.21所示，人源成纤维细胞生长因子-23的氨基酸序列如SEQ ID No.22所示。
绿色荧光蛋白的氨基酸序列如SEQ ID No.23所示；小泛素样修饰因子的氨基酸序列如SEQ ID No.24所示；绿色荧光蛋白和小泛素样修饰因子之间的多肽接头的氨基酸序列如SEQ ID No.25所示。
编码人源成纤维细胞生长因子-1的核苷酸如SEQ ID No.26所示，编码人源成纤维细胞生长因子-2的核苷酸序列如SEQ ID No.27所示，编码人源成纤维细胞生长因子-3的核苷酸序列如SEQ ID No.28所示，编码人源成纤维细胞生长因子-4的核苷酸序列如SEQ IDNo.29所示，编码人源成纤维细胞生长因子-5的核苷酸序列如SEQ ID No.30所示，编码人源成纤维细胞生长因子-6的核苷酸序列如SEQ ID No.31所示，编码人源成纤维细胞生长因子-7的核苷酸序列如SEQ ID No.32所示，编码人源成纤维细胞生长因子-8的核苷酸序列如SEQ ID No.33所示，编码人源成纤维细胞生长因子-9的核苷酸序列如SEQ ID No.34所示，编码人源成纤维细胞生长因子-10的核苷酸序列如SEQ ID No.35所示，编码人源成纤维细胞生长因子-11的核苷酸序列如SEQ ID No.36所示，编码人源成纤维细胞生长因子-12的核苷酸序列如SEQ ID No.37所示，编码人源成纤维细胞生长因子-13的核苷酸序列如SEQ IDNo.38所示，编码人源成纤维细胞生长因子-14的核苷酸序列如SEQ ID No.39所示，编码人源成纤维细胞生长因子-16的核苷酸序列如SEQ ID No.40所示，编码人源成纤维细胞生长因子-17的核苷酸序列如SEQ ID No.41所示，编码人源成纤维细胞生长因子-18的核苷酸序列如SEQ ID No.42所示，编码人源成纤维细胞生长因子-19的核苷酸序列如SEQ ID No.43所示，编码人源成纤维细胞生长因子-20的核苷酸序列如SEQ ID No.44所示，编码人源成纤维细胞生长因子-21的核苷酸序列如SEQ ID No.45所示，编码人源成纤维细胞生长因子-22的核苷酸序列如SEQ ID No.46所示，编码人源成纤维细胞生长因子-23的核苷酸序列如SEQID No.47所示。
编码绿色荧光蛋白的核苷酸序列如SEQ ID No.48所示；编码小泛素样修饰因子的核苷酸序列如SEQ ID No.49所示；编码绿色荧光蛋白和小泛素样修饰因子之间的多肽接头的核苷酸序列如SEQ ID No.50所示。
小泛素样修饰因子蛋白酶的氨基酸序列如SEQ ID No.51所示；编码小泛素样修饰因子蛋白酶的基因的核苷酸序列如SEQ ID No.52所示。CspA启动子、CspA 3’-UTR和CspA5’-UTR位于pCold载体上。
实施例1可视化的成纤维细胞生长因子融合蛋白的原核表达载体的构建和重组菌株的获得
设计融合蛋白GFP-SUMO-FGFn(其中n为1至23，且n不为15，与人源成纤维细胞生长因子-n中的n一致)从N端到C端依次为绿色荧光蛋白、多肽接头、小泛素样修饰因子和人源成纤维细胞生长因子-n(其中n为1至23，且n不为15)，各部分的氨基酸序列如上所述；编码融合蛋白的核酸从5’端到3’端依次为绿色荧光蛋白、多肽接头、小泛素样修饰因子和人源成纤维细胞生长因子-n(其中n为1至23，且n不为15)，各部分的核苷酸序列如上所述。
编码融合蛋白GFP-SUMO-FGFn的核酸由南京金斯瑞生物科技有限公司合成，然后利用基因工程方法将所合成的核酸片段分别克隆到大肠杆菌表达质粒载体pET-21a中，得到重组质粒，转化大肠杆菌。经核酸测序检测核酸序列的正确性。将含有编码融合蛋白GFP-SUMO-FGFn核酸的重组质粒命名为pET-GFP-SUMO-hFGFn(n相应地为1至23，且n不为15)。
将重组质粒pET-GFP-SUMO-hFGFn转化到大肠杆菌宿主BL21(DE3)中，用氨苄青霉素抗性筛选出阳性重组菌，并分别命名为pET-GFP-SUMO-hFGFn/BL21。
实施例2小泛素样修饰因子蛋白酶Ulp1的原核表达载体的构建和重组菌株的获得
小泛素样修饰因子蛋白酶Ulp1的氨基酸序列和编码其的核酸的核苷酸序列上所述。编码小泛素样修饰因子蛋白酶Ulp1的核酸由南京金斯瑞生物科技有限公司合成，然后利用基因工程方法将所合成的核酸片段克隆到大肠杆菌表达质粒载体pCold中，得到重组质粒，转化大肠杆菌。经核酸测序检测核酸序列的正确性，将正确的重组质粒命名为pCold-Ulp1。
利用Pfo I单酶切pCold-Ulp1质粒，获得含有CspA启动子、3’-UTR、编码小泛素样修饰因子蛋白酶的核酸和5’-UTR的小泛素样修饰因子蛋白酶表达盒的DNA片段，然后将该表达盒的DNA片段利用T4聚合酶将酶切位点末端补平，形成平末端。利用平末端限制性内切酶EcoICRI单酶切pET-GFP-SUMO-hFGFn，获得载体骨架，将该载体骨架与小泛素样修饰因子蛋白酶表达盒连接，转化到大肠杆菌。经核酸测序检测核酸序列的正确性，将正确的重组质粒命名为pET-GSFGFn-CspA-Ulp1(n＝1,2,3,4,5,6,7,8,9,10,11,12,13,14,16,17,18,19,20,21,22,23)。以pET-GSFGF9-CspA-Ulp1为例展示质粒图谱如图1所示。
将重组质pET-GSFGFn-CspA-Ulp1转化到大肠杆菌宿主BL21(DE3)中，用氨苄青霉素抗性筛选出阳性重组菌，并命名为pET-GSFGFn-CspA-Ulp1/BL21。
实施例3融合蛋白GFP-SUMO的原核表达载体的构建和重组菌株的获得
设计融合蛋白GFP-SUMO从N端到C端依次为绿色荧光蛋白、多肽接头和小泛素样修饰因子，各部分的氨基酸序列如上所述；编码融合蛋白的核酸从5’端到3’端依次为绿色荧光蛋白、多肽接头和小泛素样修饰因子，各部分的核苷酸序列如上所述。
编码融合蛋白GFP-SUMO的核酸由南京金斯瑞生物科技有限公司合成，然后利用基因工程方法将所合成的核酸片段克隆到大肠杆菌表达质粒载体pET-21a中，得到重组质粒，转化大肠杆菌。经核酸测序检测核酸序列的正确性。将含有编码GFP-SUMO核酸的重组质粒命名为pET-GFP-SUMO。
将重组质粒pET-GFP-SUMO转化到大肠杆菌宿主BL21(DE3)中，用氨苄青霉素抗性筛选出阳性重组菌，并分别命名为pET-GFP-SUMO/BL21，该重组菌株作为pET-GSFGFn-CspA-Ulp1/BL21表达蛋白的对照菌株。
实施例4重组菌株的诱导表达
按0.1％(体积/体积)的接种量将pET-GSFGFn-CspA-Ulp1/BL21接种到50毫升新鲜的液体LB培养基中(蛋白胨10克/升，酵母提取物5克/升，氯化钠10克/升，氨苄青霉素100毫克/升，pH＝7.0)，220转/分钟，37℃振荡培养14小时。然后，按5％(体积/体积)的接种量接种到1000毫升新鲜的液体LB培养基中，220转/分钟，37℃振荡培养至OD600＝1.0时(约2-3小时)，加入终浓度为1毫摩尔/升的异丙基-β-D-硫代半乳糖苷(Isopropyl-beta-D-thiogalactopyranoside，IPTG)，37℃，160转/分钟振荡培养，1小时后，发酵液逐步呈现绿色荧光，如图2所示为pET-GSFGF17-CspA-Ulp1/BL21经IPTG诱导表达后的荧光观察，其中的图A为发酵液在自然光下的观察结果，其中左侧试管中的发酵液显示出明显绿色荧光(pET-GSFGF17-CspA-Ulp1/BL21经IPTG诱导表达1小时后)，右侧试管的发酵液不显示绿色荧光(pET-GSFGF17-CspA-Ulp1/BL21 IPTG诱导表达前)；图B为pET-GSFGF17-CspA-Ulp1/BL21发酵液在紫外灯光下的观察结果，显示出的绿色荧光更为明显。其他成纤维细胞生长因子重组菌株亦有相同的结果。经IPTG诱导4小时后，12000转/分钟离心，收集显示绿色荧光的沉淀物，即诱导表达后的重组菌体，用磷酸缓冲液(137毫摩尔/升的氯化钠，2.7毫摩尔/升的氯化钾，10毫摩尔/升的磷酸二氢钠，2毫摩尔/升的磷酸二氢钾，0.2％NP-40，pH＝7.0)重悬重组菌体，得到菌悬液。将菌悬液制样进行十二烷基硫酸钠-聚丙烯酰胺凝胶电泳(sodium-dodecylsulfatepolyacrylamide gel electrophoresis，SDS-PAGE)检测。SDS-PAGE的具体操作方法参考Sambrook et al.，Molecular Cloning：A Laboratory Manual，Cold SpringHarbor Laboratory Press，Cold Spring Harbor，NY，1989。
以进行同样诱导操作的pET-GFP-SUMO/BL21作为对照；同时以IPTG诱导前的pET-GSFGF8-CspA-Ulp1/BL21(图3A)、IPTG诱导前的pET-GSFGF4-CspA-Ulp1/BL21(图3B)和IPTG诱导前的pET-GSFGF11-CspA-Ulp1/BL21(图3C)作为另外的对照，即在加入IPTG之前，从发酵液中取出1毫升菌液以用于后续总蛋白的SDS-PAGE。
SDS-PAGE的结果见图3。从图3可知，融合蛋白GFP-SUMO-FGFn可大量表达。采用Image J软件对SDS-PAGE结果进行密度灰度扫描显示，融合蛋白GFP-SUMO-FGFn表达量占总蛋白的40％以上。
实施例5重组菌株的诱导表达
IPTG诱导的温度为28℃，诱导的时间为8小时，其他同实施例4。结果与实施例4相当。
实施例6重组菌株的诱导的可溶性分析
以实施例4相同的操作进行融合蛋白GFP-SUMO-FGFn的IPTG诱导表达。诱导表达后，12000转/分钟离心，收集显示绿色荧光的沉淀物，即诱导表达后的重组菌体，并称重。
将上述收集得到的重组菌体，按重组菌体：磷酸缓冲液为1：10的比例(质量/体积)加入磷酸缓冲液(137毫摩尔/升的氯化钠，2.7毫摩尔/升的氯化钾，10毫摩尔/升的磷酸二氢钠，2毫摩尔/升的磷酸二氢钾，0.2％NP-40，pH＝7.0)重悬重组菌体，得到菌悬液。将菌悬液置于高压均质机中，在4℃，先后在800和1200bar(巴)的压强条件下连续均质处理2次，得到裂解液。将裂解液置于无菌离心管中，12000rpm(转/分钟)离心1分钟，取裂解液、上清液和沉淀分别进行SDS-PAGE。
部分pET-GSFGFn-CspA-Ulp1/BL21总蛋白、上清液中的蛋白以及沉淀中的蛋白SDS-PAGE的结果展示于图4。从图4可知，融合蛋白GFP-SUMO-FGF4、GFP-SUMO-FGF5、GFP-SUMO-FGF8、GFP-SUMO-FGF11、GFP-SUMO-FGF12、GFP-SUMO-FGF14、GFP-SUMO-FGF16、GFP-SUMO-FGF17、GFP-SUMO-FGF19和GFP-SUMO-FGF23的可溶性表达量占融合蛋白总量的80％以上；而且这些融合蛋白没有出现被水解的现象，这说明在37℃ IPTG诱导的过程中不会导致小泛素样修饰因子蛋白酶Ulp1的正常表达，因而没有出现融合蛋白GFP-SUMO-FGFn被水解的现象。
未示出的pET-GSFGFn-CspA-Ulp1/BL21以同样地诱导表达操作后结果相当，即可溶性融合蛋白GFP-SUMO-FGFn的表达量占融合蛋白GFP-SUMO-FGFn总量的80％以上；而且这些融合蛋白同样没有出现被水解的现象。
综合上述这些结果表明，利用融合表达的方式可以高效表达可溶性含成纤维细胞生长因子-n的融合蛋白GFP-SUMO-FGFn，且其产量能完全满足后期的成纤维细胞生长因子规模化生产的需要。
实施例7含成纤维细胞生长因子的融合蛋白的水解
按实施例4的方法对重组菌株pET-GSFGFn-CspA-Ulp1/BL21进行37℃发酵并利用在IPTG诱导4小时后迅速降低到15℃，取出部分发酵液分装到5个1000毫升体积的摇瓶中，摇瓶编号为摇瓶1至摇瓶5，每个摇瓶加入200毫升的发酵液。
以进行同样诱导操作的pET-GFP-SUMO/BL21作为对照，对照摇瓶仅1个，编号为摇瓶I。
在摇瓶1和摇瓶I中，分别加入硫酸锌水溶液，使硫酸锌在发酵液中的终浓度为5毫摩尔/升。
在摇瓶2I中，加入三(2-羰基乙基)磷盐酸盐水溶液，使三(2-羰基乙基)磷盐酸盐在发酵液中的终浓度为1毫摩尔/升。
在摇瓶3中，加入三(2-羰基乙基)磷盐酸盐水溶液，使三(2-羰基乙基)磷盐酸盐在发酵液中的终浓度为1毫摩尔/升；加入硫酸锌水溶液，使硫酸锌在发酵液中的终浓度为1毫摩尔/升。
在摇瓶4中，加入三(2-羰基乙基)磷盐酸盐水溶液，使三(2-羰基乙基)磷盐酸盐在发酵液中的终浓度为1毫摩尔/升；加入硫酸锌水溶液，使硫酸锌在发酵液中的终浓度为3毫摩尔/升。
在摇瓶5中，加入三(2-羰基乙基)磷盐酸盐水溶液，使三(2-羰基乙基)磷盐酸盐在发酵液中的终浓度为0.1毫摩尔/升；加入硫酸锌水溶液，使硫酸锌在发酵液中的终浓度为5毫摩尔/升。
在加入上述组分之后，于15℃中继续培养摇瓶1至摇瓶5和摇瓶I中的发酵液2小时。其中，并在发酵过程中，在0分钟、30分钟、60分钟、90分钟、120分钟分别取样以备用于后续的SDS-PAGE。
将不同摇瓶不同时间点所取样品分别于12000转/分钟高速离心，并分别收集重组菌体，按重组菌体：磷酸缓冲液为1：10的比例(质量/体积)分别加入磷酸缓冲液重悬重组菌体，得到重组菌悬液。将该悬液置于高压均质机中，在4℃，先后在800和1200bar(巴)压强条件下连续均质处理2次，得到裂解液。将裂解液置于无菌离心管中，12000rpm(转/分钟)离心1分钟，收集上清液进行SDS-PAGE检测分析。其中图5显示的是pET-GSFGF12-CspA-Ulp1/BL21的结果。根据图5的电泳结果显示，单独添加硫酸锌水溶液时，即使低温诱导长达2小时，也完全不能促使SUMO蛋白酶对融合蛋白GFP-SUMO-FGF12的水解(图5A)；单独添加终浓度为1毫摩尔/升的三(2-羰基乙基)磷盐酸盐溶液虽能促进SUMO蛋白酶水解融合蛋白GFP-SUMO-FGF12得到成纤维细胞生长因子-12，但水解效率偏低，在处理120分钟后，仍有约30％至40％融合蛋白没有被水解(图5B)；而当同时添加三(2-羰基乙基)磷盐酸盐和硫酸锌时，被SUMO蛋白酶水解的融合蛋白量明显增加，基本上可以在0.5至1小时内将90％以上的融合蛋白GFP-SUMO-FGFn水解，说明该组合具有协同增效作用，能显著促高SUMO蛋白酶对融合蛋白的水解能力(图5C-E)；其中，在低温诱导表达过程中，添加终浓度为0.1毫摩尔/升三(2-羰基乙基)磷盐酸盐和5毫摩尔/升硫酸锌组合的结果显示融合蛋白在15分钟内即被低温诱导表达的SUMO蛋白酶完全水解(图5E)。
此外，对其它融合蛋白GFP-SUMO-FGFn的测试结果显示与GFP-SUMO-FGF12一致。其中图6显示了利用终浓度为0.1毫摩尔/升的三(2-羰基乙基)磷盐酸盐和5毫摩尔/升的硫酸锌组合在15℃培养30分钟内部分pET-GSFGFn-CspA-Ulp1/BL21的上清液SDS-PAGE，图中显示融合蛋白GFP-SUMO-FGFn在15℃低温诱导的30分钟内均能被小泛素样修饰因子蛋白酶完全水解。
为了更清晰地了解融合蛋白GFP-SUMO-FGFn被SUMO蛋白酶逐步水解的进程，利用抗人成纤维细胞生长因子-9(赛默飞世尔科技(中国)有限公司，货号：PA5-111618)和抗人成纤维细胞生长因子-18的抗体(赛默飞世尔科技(中国)有限公司，货号：PA5-78261)分别对pET-GSFGF9-CspA-Ulp1/BL21和pET-GSFGF18-CspA-Ulp1/BL21的低温诱导5分钟和30分钟的上清液进行蛋白免疫印迹分析。结果见图7，蛋白免疫印迹印证了融合蛋白GFP-SUMO-FGFn能够在30分钟之内完全水解。
实施例8含成纤维细胞生长因子的融合蛋白的水解
按实施例4的方法对重组菌株pET-GSFGFn-CspA-Ulp1/BL21进行32℃发酵并利用在IPTG诱导6小时后迅速降低到10℃，加入三(2-羰基乙基)磷盐酸盐水溶液，使三(2-羰基乙基)磷盐酸盐在发酵液中的终浓度为0.1毫摩尔/升，加入硫酸锌水溶液，使硫酸锌在发酵液中的终浓度为5毫摩尔/升，继续培养发酵液2小时。其中，并在发酵过程中，在0分钟、30分钟、60分钟、90分钟、120分钟分别取样以备用于后续的SDS-PAGE。蛋白免疫印迹结果同样表明融合蛋白GFP-SUMO-FGFn能够在30分钟之内被SUMO蛋白酶完全水解。
实施例9成纤维细胞因子-n的大规模发酵及分离纯化
从-80℃冰箱中取出pET-GSFGF17-CspA-Ulp1/BL21冻存菌种，以1：100的比例(体积/体积)接种到3.7升新鲜LB培养基(蛋白胨10克/升，酵母提取物5克/升，氯化钠10克/升，氨苄青霉素100毫克/升，pH＝7.0)中，37℃、220rpm(转/分钟)振荡培养4小时。将菌液以10％接种量(体积/体积)接种到200L(升)含5％(质量/体积)葡萄糖的LB培养基中，37℃培养4-6小时，直至菌体密度达到OD600≈0.6，全部转入800L(升)含5％(质量/体积)葡萄糖的LB培养基中进行培养4小时，再接入到8000L含5％(质量/体积)葡萄糖的LB培养基中进行培养(总体积约为9000L)。待培养基中葡萄糖含量下降到0.1％(质量/体积)时，开始流加补料，补料培养基为含20％(质量/体积)葡萄糖的LB培养基，设定发酵参数：pH＝7.0-7.2；溶氧>30％；搅拌速度250至650rpm(转/分钟)。当湿菌重达到35±1g/L(克/升)时，添加诱导剂在IPTG并使其终浓度达到1毫摩尔/升，诱导4小时。然后，迅速下降发酵液的温度至15℃，与此同时，在发酵液中加入三(2-羰基乙基)磷盐酸盐水溶液，使三(2-羰基乙基)磷盐酸盐在发酵液中的终浓度为0.1毫摩尔/升，同时加入硫酸锌水溶液，使硫酸锌在发酵液中的终浓度为5毫摩尔/升，并设定参数为：pH＝7.0；溶氧>30％；搅拌速度400rpm(转/分钟)，培养1小时，整个发酵和诱导表达过程结束，最终发酵液的总体积约为10000L，离心收集菌体并称重，共得到513千克(KG)湿重的重组菌体，经计算达到51.3g/L湿重的重组菌体。
由于成纤维细胞生长因子都具有肝素结合域，因此，可以利用肝素亲和层析凝胶直接吸附的方法纯化自融合蛋白水解出的成纤维细胞生长因子。
对收集的全部重组菌体采用同实施例6相同的操作进行高压均质机破碎，得到裂解液。将裂解液置于无菌离心管中，12000rpm(转/分钟)离心1分钟，收集上清液，将上清液上样于CM离子交换凝胶层析柱，用含有不同氯化钠浓度的缓冲液对成纤维细胞生长因子-17进行梯度洗脱。所使用的缓冲液依次为缓冲液1(50毫摩尔/升Tris-HCl，0.1摩尔/升氯化钠，pH＝8.0)；缓冲液2(50毫摩尔/升Tris-HCl，0.3摩尔/升氯化钠，pH＝8.0)；缓冲液3(50毫摩尔/升Tris-HCl，0.6摩尔/升氯化钠，pH＝8.0)和缓冲液6(50毫摩尔/升Tris-HCl，1.0摩尔/升氯化钠，pH＝8.0)。SDS-PAGE检测，结果见图8A，分析显示成纤维细胞生长因子-17主要存在于缓冲液3的洗脱液中。将含有成纤维细胞生长因子-17的洗脱液以8-10毫升/分钟的流速上样于肝素亲和层析凝胶柱中，然后，依次使用缓冲液3(50毫摩尔/升Tris-HCl，0.6摩尔/升氯化钠，pH＝8.0)，缓冲液4(50毫摩尔/升Tris-HCl，1.2摩/升尔氯化钠，pH＝8.0)和缓冲液5(50毫摩尔/升Tris-HCl，2.0摩尔/升氯化钠，pH＝8.0)进行洗脱。SDS-PAGE见图8B，检测结果显示，成纤维细胞生长因子-17位于缓冲液4的洗脱液中，电泳条带密度灰度扫描(ImageJ软件)结果表明，其纯度达到95％以上。
利用二喹啉酸(BCA)蛋白定量法对纯化的成纤维细胞生长因子-17定量，并基于pET-GSFGF17-CspA-Ulp1/BL21重组菌体的湿重进行换算，得出成纤维细胞生长因子-17的得率，最终纯化后的成纤维细胞生长因子-17的得率为5.048毫克/克湿重。
利用相同的发酵方法发酵其他pET-GSFGFn-CspA-Ulp1/BL21并纯化成纤维细胞生长因子-n，可以获得类似的发酵结果。纯化这些成纤维细胞生长因子的得率分别为：成纤维细胞生长因子-1 6.066毫克/克湿重，成纤维细胞生长因子-2 8.62毫克/克湿重，成纤维细胞生长因子-3 9.329毫克/克湿重，成纤维细胞生长因子-4 7.14毫克/克湿重，成纤维细胞生长因子-5 5.224毫克/克湿重，成纤维细胞生长因子-6 1.34毫克/克湿重，成纤维细胞生长因子-7 2.019毫克/克湿重，成纤维细胞生长因子-8 2.46毫克/克湿重，成纤维细胞生长因子-9 4.984毫克/克湿重，成纤维细胞生长因子-10 6.673毫克/克湿重，成纤维细胞生长因子-111.243毫克/克湿重，成纤维细胞生长因子-12 1.758毫克/克湿重，成纤维细胞生长因子-13 3.153毫克/克湿重，成纤维细胞生长因子-14 1.585毫克/克湿重，成纤维细胞生长因子-16 1.892毫克/克湿重，成纤维细胞生长因子-18 2.298毫克/克湿重，成纤维细胞生长因子-19 1.957毫克/克湿重，成纤维细胞生长因子-201.466毫克/克湿重，成纤维细胞生长因子-21 3.428毫克/克湿重，成纤维细胞生长因子-22 1.206毫克/克湿重，成纤维细胞生长因子-23 2.694毫克/克湿重。
实施例10成纤维细胞生长因子-8、成纤维细胞生长因子-9和成纤维细胞生长因子-17对***细胞促增殖活性分析
人体内95％的雄激素都是由***细胞合成的，而***细胞数量的减少会导致人体内雄激素的产量显著下降。
将人原代***细胞(货号：4510，ScienCell Research Laboratories,Inc，美国)，接种到改良杜氏伊格尔培养基(Dulbecco's Modified Eagle Medium，DMEM/F-12，购自美国赛默飞公司，货号：11320033)中，37℃培养至对数生长期，用胰蛋白酶溶液消化后，再转接到96孔细胞培养板中(8000个细胞/孔，200微升体积/孔)，然后分别加入含有终浓度为100纳克/毫升纯化的成纤维细胞生长因子-8(FGF-8)、成纤维细胞生长因子-9(FGF-9)和成纤维细胞生长因子-17(FGF-17)，以添加等体积的PBS缓冲液作为阴性对照。培养24小时后，弃去100微升培养基，再加入100微升含有终浓度为10微摩尔/升的5-乙基-2'-脱氧尿苷(EdU，一种示踪化合物，细胞增殖需要脱氧核糖核酸(DNA)发生复制，在此过程中，EdU会***到新形成的核苷酸链中，在波长为450-540纳米的光照射下发绿色荧光)的新鲜改良杜氏伊格尔培养基，继续培育24小时，荧光显微镜观察，结果见图9。图9的结果显示，相对于阴性对照组，FGF-8、FGF-9和FGF-17可以显著促进***细胞脱氧核糖核酸(DNA)的合成。
序列表
<110> 温州医科大学
苏志坚
<120> 一种组合物及规模化生产成纤维细胞生长因子的方法
<130> LHA2160323
<160> 53
<170> SIPOSequenceListing 1.0
<210> 2
<211> 141
<212> PRT
<213> 智人(Homo sapiens)
<400> 2
Met Phe Asn Leu Pro Pro Gly Asn Tyr Lys Lys Pro Lys Leu Leu Tyr
1 5 10 15
Cys Ser Asn Gly Gly His Phe Leu Arg Ile Leu Pro Asp Gly Thr Val
20 25 30
Asp Gly Thr Arg Asp Arg Ser Asp Gln His Ile Gln Leu Gln Leu Ser
35 40 45
Ala Glu Ser Val Gly Glu Val Tyr Ile Lys Ser Thr Glu Thr Gly Gln
50 55 60
Tyr Leu Ala Met Asp Thr Asp Gly Leu Leu Tyr Gly Ser Gln Thr Pro
65 70 75 80
Asn Glu Glu Cys Leu Phe Leu Glu Arg Leu Glu Glu Asn His Tyr Asn
85 90 95
Thr Tyr Ile Ser Lys Lys His Ala Glu Lys Asn Trp Phe Val Gly Leu
100 105 110
Lys Lys Asn Gly Ser Cys Lys Arg Gly Pro Arg Thr His Tyr Gly Gln
115 120 125
Lys Ala Ile Leu Phe Leu Pro Leu Pro Val Ser Ser Asp
130 135 140
<210> 2
<211> 155
<212> PRT
<213> 智人(Homo sapiens)
<400> 2
Met Ala Ala Gly Ser Ile Thr Thr Leu Pro Ala Leu Pro Glu Asp Gly
1 5 10 15
Gly Ser Gly Ala Phe Pro Pro Gly His Phe Lys Asp Pro Lys Arg Leu
20 25 30
Tyr Cys Lys Asn Gly Gly Phe Phe Leu Arg Ile His Pro Asp Gly Arg
35 40 45
Val Asp Gly Val Arg Glu Lys Ser Asp Pro His Ile Lys Leu Gln Leu
50 55 60
Gln Ala Glu Glu Arg Gly Val Val Ser Ile Lys Gly Val Cys Ala Asn
65 70 75 80
Arg Tyr Leu Ala Met Lys Glu Asp Gly Arg Leu Leu Ala Ser Lys Cys
85 90 95
Val Thr Asp Glu Cys Phe Phe Phe Glu Arg Leu Glu Ser Asn Asn Tyr
100 105 110
Asn Thr Tyr Arg Ser Arg Lys Tyr Thr Ser Trp Tyr Val Ala Leu Lys
115 120 125
Arg Thr Gly Gln Tyr Lys Leu Gly Ser Lys Thr Gly Pro Gly Gln Lys
130 135 140
Ala Ile Leu Phe Leu Pro Met Ser Ala Lys Ser
145 150 155
<210> 3
<211> 224
<212> PRT
<213> 智人(Homo sapiens)
<400> 3
Met Pro Ala Ala Gly Pro Gly Ala Arg Leu Arg Arg Asp Ala Gly Gly
1 5 10 15
Arg Gly Gly Val Tyr Glu His Leu Gly Gly Ala Pro Arg Arg Arg Lys
20 25 30
Leu Tyr Cys Ala Thr Lys Tyr His Leu Gln Leu His Pro Ser Gly Arg
35 40 45
Val Asn Gly Ser Leu Glu Asn Ser Ala Tyr Ser Ile Leu Glu Ile Thr
50 55 60
Ala Val Glu Val Gly Ile Val Ala Ile Arg Gly Leu Phe Ser Gly Arg
65 70 75 80
Tyr Leu Ala Met Asn Lys Arg Gly Arg Leu Tyr Ala Ser Glu His Tyr
85 90 95
Ser Ala Glu Cys Glu Phe Val Glu Arg Ile His Glu Leu Gly Tyr Asn
100 105 110
Thr Tyr Ala Ser Arg Leu Tyr Arg Thr Val Ser Ser Thr Pro Gly Ala
115 120 125
Arg Arg Gln Pro Ser Ala Glu Arg Leu Trp Tyr Val Ser Val Asn Gly
130 135 140
Lys Gly Arg Pro Arg Arg Gly Phe Lys Thr Arg Arg Thr Gln Lys Ser
145 150 155 160
Ser Leu Phe Leu Pro Arg Val Leu Asp His Arg Asp His Glu Met Val
165 170 175
Arg Gln Leu Gln Ser Gly Leu Pro Arg Pro Pro Gly Lys Gly Val Gln
180 185 190
Pro Arg Arg Arg Arg Gln Lys Gln Ser Pro Asp Asn Leu Glu Pro Ser
195 200 205
His Val Gln Ala Ser Arg Leu Gly Ser Gln Leu Glu Ala Ser Ala His
210 215 220
<210> 4
<211> 183
<212> PRT
<213> 智人(Homo sapiens)
<400> 4
Met Gly Arg Gly Gly Ala Ala Ala Pro Thr Ala Pro Asn Gly Thr Leu
1 5 10 15
Glu Ala Glu Leu Glu Arg Arg Trp Glu Ser Leu Val Ala Leu Ser Leu
20 25 30
Ala Arg Leu Pro Val Ala Ala Gln Pro Lys Glu Ala Ala Val Gln Ser
35 40 45
Gly Ala Gly Asp Tyr Leu Leu Gly Ile Lys Arg Leu Arg Arg Leu Tyr
50 55 60
Cys Asn Val Gly Ile Gly Phe His Leu Gln Ala Leu Pro Asp Gly Arg
65 70 75 80
Ile Gly Gly Ala His Ala Asp Thr Arg Asp Ser Leu Leu Glu Leu Ser
85 90 95
Pro Val Glu Arg Gly Val Val Ser Ile Phe Gly Val Ala Ser Arg Phe
100 105 110
Phe Val Ala Met Ser Ser Lys Gly Lys Leu Tyr Gly Ser Pro Phe Phe
115 120 125
Thr Asp Glu Cys Thr Phe Lys Glu Ile Leu Leu Pro Asn Asn Tyr Asn
130 135 140
Ala Tyr Glu Ser Tyr Lys Tyr Pro Gly Met Phe Ile Ala Leu Ser Lys
145 150 155 160
Asn Gly Lys Thr Lys Lys Gly Asn Arg Val Ser Pro Thr Met Lys Val
165 170 175
Thr His Phe Leu Pro Arg Leu
180
<210> 5
<211> 252
<212> PRT
<213> 智人(Homo sapiens)
<400> 5
Met Ala Trp Ala His Gly Glu Lys Arg Leu Ala Pro Lys Gly Gln Pro
1 5 10 15
Gly Pro Ala Ala Thr Asp Arg Asn Pro Ile Gly Ser Ser Ser Arg Gln
20 25 30
Ser Ser Ser Ser Ala Met Ser Ser Ser Ser Ala Ser Ser Ser Pro Ala
35 40 45
Ala Ser Leu Gly Ser Gln Gly Ser Gly Leu Glu Gln Ser Ser Phe Gln
50 55 60
Trp Ser Pro Ser Gly Arg Arg Thr Gly Ser Leu Tyr Cys Arg Val Gly
65 70 75 80
Ile Gly Phe His Leu Gln Ile Tyr Pro Asp Gly Lys Val Asn Gly Ser
85 90 95
His Glu Ala Asn Met Leu Ser Val Leu Glu Ile Phe Ala Val Ser Gln
100 105 110
Gly Ile Val Gly Ile Arg Gly Val Phe Ser Asn Lys Phe Leu Ala Met
115 120 125
Ser Lys Lys Gly Lys Leu His Ala Ser Ala Lys Phe Thr Asp Asp Cys
130 135 140
Lys Phe Arg Glu Arg Phe Gln Glu Asn Ser Tyr Asn Thr Tyr Ala Ser
145 150 155 160
Ala Ile His Arg Thr Glu Lys Thr Gly Arg Glu Trp Tyr Val Ala Leu
165 170 175
Asn Lys Arg Gly Lys Ala Lys Arg Gly Cys Ser Pro Arg Val Lys Pro
180 185 190
Gln His Ile Ser Thr His Phe Leu Pro Arg Phe Lys Gln Ser Glu Gln
195 200 205
Pro Glu Leu Ser Phe Thr Val Thr Val Pro Glu Lys Lys Asn Pro Pro
210 215 220
Ser Pro Ile Lys Ser Lys Ile Pro Leu Ser Ala Pro Arg Lys Asn Thr
225 230 235 240
Asn Ser Val Lys Tyr Arg Leu Lys Phe Arg Phe Gly
245 250
<210> 6
<211> 169
<212> PRT
<213> 智人(Homo sapiens)
<400> 6
Met Gly Thr Arg Ala Asn Asn Thr Leu Leu Asp Ser Arg Gly Trp Gly
1 5 10 15
Thr Leu Leu Ser Arg Ser Arg Ala Gly Leu Ala Gly Glu Ile Ala Gly
20 25 30
Val Asn Trp Glu Ser Gly Tyr Leu Val Gly Ile Lys Arg Gln Arg Arg
35 40 45
Leu Tyr Cys Asn Val Gly Ile Gly Phe His Leu Gln Val Leu Pro Asp
50 55 60
Gly Arg Ile Ser Gly Thr His Glu Glu Asn Pro Tyr Ser Leu Leu Glu
65 70 75 80
Ile Ser Thr Val Glu Arg Gly Val Val Ser Leu Phe Gly Val Arg Ser
85 90 95
Ala Leu Phe Val Ala Met Asn Ser Lys Gly Arg Leu Tyr Ala Thr Pro
100 105 110
Ser Phe Gln Glu Glu Cys Lys Phe Arg Glu Thr Leu Leu Pro Asn Asn
115 120 125
Tyr Asn Ala Tyr Glu Ser Asp Leu Tyr Gln Gly Thr Tyr Ile Ala Leu
130 135 140
Ser Lys Tyr Gly Arg Val Lys Arg Gly Ser Lys Val Ser Pro Ile Met
145 150 155 160
Thr Val Thr His Phe Leu Pro Arg Ile
165
<210> 7
<211> 141
<212> PRT
<213> 智人(Homo sapiens)
<400> 7
Met Ser Tyr Asp Tyr Met Glu Gly Gly Asp Ile Arg Val Arg Arg Leu
1 5 10 15
Phe Cys Arg Thr Gln Trp Tyr Leu Arg Ile Asp Lys Arg Gly Lys Val
20 25 30
Lys Gly Thr Gln Glu Met Lys Asn Asn Tyr Asn Ile Met Glu Ile Arg
35 40 45
Thr Val Ala Val Gly Ile Val Ala Ile Lys Gly Val Glu Ser Glu Phe
50 55 60
Tyr Leu Ala Met Asn Lys Glu Gly Lys Leu Tyr Ala Lys Lys Glu Cys
65 70 75 80
Asn Glu Asp Cys Asn Phe Lys Glu Leu Ile Leu Glu Asn His Tyr Asn
85 90 95
Thr Tyr Ala Ser Ala Lys Trp Thr His Asn Gly Gly Glu Met Phe Val
100 105 110
Ala Leu Asn Gln Lys Gly Ile Pro Val Arg Gly Lys Lys Thr Lys Lys
115 120 125
Glu Gln Lys Thr Ala His Phe Leu Pro Met Ala Ile Thr
130 135 140
<210> 8
<211> 194
<212> PRT
<213> 智人(Homo sapiens)
<400> 8
Met Gln Val Thr Val Gln Ser Ser Pro Asn Phe Thr Gln His Val Arg
1 5 10 15
Glu Gln Ser Leu Val Thr Asp Gln Leu Ser Arg Arg Leu Ile Arg Thr
20 25 30
Tyr Gln Leu Tyr Ser Arg Thr Ser Gly Lys His Val Gln Val Leu Ala
35 40 45
Asn Lys Arg Ile Asn Ala Met Ala Glu Asp Gly Asp Pro Phe Ala Lys
50 55 60
Leu Ile Val Glu Thr Asp Thr Phe Gly Ser Arg Val Arg Val Arg Gly
65 70 75 80
Ala Glu Thr Gly Leu Tyr Ile Cys Met Asn Lys Lys Gly Lys Leu Ile
85 90 95
Ala Lys Ser Asn Gly Lys Gly Lys Asp Cys Val Phe Thr Glu Ile Val
100 105 110
Leu Glu Asn Asn Tyr Thr Ala Leu Gln Asn Ala Lys Tyr Glu Gly Trp
115 120 125
Tyr Met Ala Phe Thr Arg Lys Gly Arg Pro Arg Lys Gly Ser Lys Thr
130 135 140
Arg Gln His Gln Arg Glu Val His Phe Met Lys Arg Leu Pro Arg Gly
145 150 155 160
His His Thr Thr Glu Gln Ser Leu Arg Phe Glu Phe Leu Asn Tyr Pro
165 170 175
Pro Phe Thr Arg Ser Leu Arg Gly Ser Gln Arg Thr Trp Ala Pro Glu
180 185 190
Pro Arg
<210> 9
<211> 207
<212> PRT
<213> 智人(Homo sapiens)
<400> 9
Met Pro Leu Gly Glu Val Gly Asn Tyr Phe Gly Val Gln Asp Ala Val
1 5 10 15
Pro Phe Gly Asn Val Pro Val Leu Pro Val Asp Ser Pro Val Leu Leu
20 25 30
Ser Asp His Leu Gly Gln Ser Glu Ala Gly Gly Leu Pro Arg Gly Pro
35 40 45
Ala Val Thr Asp Leu Asp His Leu Lys Gly Ile Leu Arg Arg Arg Gln
50 55 60
Leu Tyr Cys Arg Thr Gly Phe His Leu Glu Ile Phe Pro Asn Gly Thr
65 70 75 80
Ile Gln Gly Thr Arg Lys Asp His Ser Arg Phe Gly Ile Leu Glu Phe
85 90 95
Ile Ser Ile Ala Val Gly Leu Val Ser Ile Arg Gly Val Asp Ser Gly
100 105 110
Leu Tyr Leu Gly Met Asn Glu Lys Gly Glu Leu Tyr Gly Ser Glu Lys
115 120 125
Leu Thr Gln Glu Cys Val Phe Arg Glu Gln Phe Glu Glu Asn Trp Tyr
130 135 140
Asn Thr Tyr Ser Ser Asn Leu Tyr Lys His Val Asp Thr Gly Arg Arg
145 150 155 160
Tyr Tyr Val Ala Leu Asn Lys Asp Gly Thr Pro Arg Glu Gly Thr Arg
165 170 175
Thr Lys Arg His Gln Lys Phe Thr His Phe Leu Pro Arg Pro Val Asp
180 185 190
Pro Asp Lys Val Pro Glu Leu Tyr Lys Asp Ile Leu Ser Gln Ser
195 200 205
<210> 10
<211> 170
<212> PRT
<213> 智人(Homo sapiens)
<400> 10
Met Leu Gly Gln Asp Met Val Ser Pro Glu Ala Thr Asn Ser Ser Ser
1 5 10 15
Ser Ser Phe Ser Ser Pro Ser Ser Ala Gly Arg His Val Arg Ser Tyr
20 25 30
Asn His Leu Gln Gly Asp Val Arg Trp Arg Lys Leu Phe Ser Phe Thr
35 40 45
Lys Tyr Phe Leu Lys Ile Glu Lys Asn Gly Lys Val Ser Gly Thr Lys
50 55 60
Lys Glu Asn Cys Pro Tyr Ser Ile Leu Glu Ile Thr Ser Val Glu Ile
65 70 75 80
Gly Val Val Ala Val Lys Ala Ile Asn Ser Asn Tyr Tyr Leu Ala Met
85 90 95
Asn Lys Lys Gly Lys Leu Tyr Gly Ser Lys Glu Phe Asn Asn Asp Cys
100 105 110
Lys Leu Lys Glu Arg Ile Glu Glu Asn Gly Tyr Asn Thr Tyr Ala Ser
115 120 125
Phe Asn Trp Gln His Asn Gly Arg Gln Met Tyr Val Ala Leu Asn Gly
130 135 140
Lys Gly Ala Pro Arg Arg Gly Gln Lys Thr Arg Arg Lys Asn Thr Ser
145 150 155 160
Ala His Phe Leu Pro Met Val Val His Ser
165 170
<210> 11
<211> 225
<212> PRT
<213> 智人(Homo sapiens)
<400> 11
Met Ala Ala Leu Ala Ser Ser Leu Ile Arg Gln Lys Arg Glu Val Arg
1 5 10 15
Glu Pro Gly Gly Ser Arg Pro Val Ser Ala Gln Arg Arg Val Cys Pro
20 25 30
Arg Gly Thr Lys Ser Leu Cys Gln Lys Gln Leu Leu Ile Leu Leu Ser
35 40 45
Lys Val Arg Leu Cys Gly Gly Arg Pro Ala Arg Pro Asp Arg Gly Pro
50 55 60
Glu Pro Gln Leu Lys Gly Ile Val Thr Lys Leu Phe Cys Arg Gln Gly
65 70 75 80
Phe Tyr Leu Gln Ala Asn Pro Asp Gly Ser Ile Gln Gly Thr Pro Glu
85 90 95
Asp Thr Ser Ser Phe Thr His Phe Asn Leu Ile Pro Val Gly Leu Arg
100 105 110
Val Val Thr Ile Gln Ser Ala Lys Leu Gly His Tyr Met Ala Met Asn
115 120 125
Ala Glu Gly Leu Leu Tyr Ser Ser Pro His Phe Thr Ala Glu Cys Arg
130 135 140
Phe Lys Glu Cys Val Phe Glu Asn Tyr Tyr Val Leu Tyr Ala Ser Ala
145 150 155 160
Leu Tyr Arg Gln Arg Arg Ser Gly Arg Ala Trp Tyr Leu Gly Leu Asp
165 170 175
Lys Glu Gly Gln Val Met Lys Gly Asn Arg Val Lys Lys Thr Lys Ala
180 185 190
Ala Ala His Phe Leu Pro Lys Leu Leu Glu Val Ala Met Tyr Gln Glu
195 200 205
Pro Ser Leu His Ser Val Pro Glu Ala Ser Pro Ser Ser Pro Pro Ala
210 215 220
Pro
225
<210> 12
<211> 243
<212> PRT
<213> 智人(Homo sapiens)
<400> 12
Met Ala Ala Ala Ile Ala Ser Ser Leu Ile Arg Gln Lys Arg Gln Ala
1 5 10 15
Arg Glu Ser Asn Ser Asp Arg Val Ser Ala Ser Lys Arg Arg Ser Ser
20 25 30
Pro Ser Lys Asp Gly Arg Ser Leu Cys Glu Arg His Val Leu Gly Val
35 40 45
Phe Ser Lys Val Arg Phe Cys Ser Gly Arg Lys Arg Pro Val Arg Arg
50 55 60
Arg Pro Glu Pro Gln Leu Lys Gly Ile Val Thr Arg Leu Phe Ser Gln
65 70 75 80
Gln Gly Tyr Phe Leu Gln Met His Pro Asp Gly Thr Ile Asp Gly Thr
85 90 95
Lys Asp Glu Asn Ser Asp Tyr Thr Leu Phe Asn Leu Ile Pro Val Gly
100 105 110
Leu Arg Val Val Ala Ile Gln Gly Val Lys Ala Ser Leu Tyr Val Ala
115 120 125
Met Asn Gly Glu Gly Tyr Leu Tyr Ser Ser Asp Val Phe Thr Pro Glu
130 135 140
Cys Lys Phe Lys Glu Ser Val Phe Glu Asn Tyr Tyr Val Ile Tyr Ser
145 150 155 160
Ser Thr Leu Tyr Arg Gln Gln Glu Ser Gly Arg Ala Trp Phe Leu Gly
165 170 175
Leu Asn Lys Glu Gly Gln Ile Met Lys Gly Asn Arg Val Lys Lys Thr
180 185 190
Lys Pro Ser Ser His Phe Val Pro Lys Pro Ile Glu Val Cys Met Tyr
195 200 205
Arg Glu Pro Ser Leu His Glu Ile Gly Glu Lys Gln Gly Arg Ser Arg
210 215 220
Lys Ser Ser Gly Thr Pro Thr Met Asn Gly Gly Lys Val Val Asn Gln
225 230 235 240
Asp Ser Thr
<210> 13
<211> 245
<212> PRT
<213> 智人(Homo sapiens)
<400> 13
Met Ala Ala Ala Ile Ala Ser Ser Leu Ile Arg Gln Lys Arg Gln Ala
1 5 10 15
Arg Glu Arg Glu Lys Ser Asn Ala Cys Lys Cys Val Ser Ser Pro Ser
20 25 30
Lys Gly Lys Thr Ser Cys Asp Lys Asn Lys Leu Asn Val Phe Ser Arg
35 40 45
Val Lys Leu Phe Gly Ser Lys Lys Arg Arg Arg Arg Arg Pro Glu Pro
50 55 60
Gln Leu Lys Gly Ile Val Thr Lys Leu Tyr Ser Arg Gln Gly Tyr His
65 70 75 80
Leu Gln Leu Gln Ala Asp Gly Thr Ile Asp Gly Thr Lys Asp Glu Asp
85 90 95
Ser Thr Tyr Thr Leu Phe Asn Leu Ile Pro Val Gly Leu Arg Val Val
100 105 110
Ala Ile Gln Gly Val Gln Thr Lys Leu Tyr Leu Ala Met Asn Ser Glu
115 120 125
Gly Tyr Leu Tyr Thr Ser Glu Leu Phe Thr Pro Glu Cys Lys Phe Lys
130 135 140
Glu Ser Val Phe Glu Asn Tyr Tyr Val Thr Tyr Ser Ser Met Ile Tyr
145 150 155 160
Arg Gln Gln Gln Ser Gly Arg Gly Trp Tyr Leu Gly Leu Asn Lys Glu
165 170 175
Gly Glu Ile Met Lys Gly Asn His Val Lys Lys Asn Lys Pro Ala Ala
180 185 190
His Phe Leu Pro Lys Pro Leu Lys Val Ala Met Tyr Lys Glu Pro Ser
195 200 205
Leu His Asp Leu Thr Glu Phe Ser Arg Ser Gly Ser Gly Thr Pro Thr
210 215 220
Lys Ser Arg Ser Val Ser Gly Val Leu Asn Gly Gly Lys Ser Met Ser
225 230 235 240
His Asn Glu Ser Thr
245
<210> 14
<211> 247
<212> PRT
<213> 智人(Homo sapiens)
<400> 14
Met Ala Ala Ala Ile Ala Ser Gly Leu Ile Arg Gln Lys Arg Gln Ala
1 5 10 15
Arg Glu Gln His Trp Asp Arg Pro Ser Ala Ser Arg Arg Arg Ser Ser
20 25 30
Pro Ser Lys Asn Arg Gly Leu Cys Asn Gly Asn Leu Val Asp Ile Phe
35 40 45
Ser Lys Val Arg Ile Phe Gly Leu Lys Lys Arg Arg Leu Arg Arg Gln
50 55 60
Asp Pro Gln Leu Lys Gly Ile Val Thr Arg Leu Tyr Cys Arg Gln Gly
65 70 75 80
Tyr Tyr Leu Gln Met His Pro Asp Gly Ala Leu Asp Gly Thr Lys Asp
85 90 95
Asp Ser Thr Asn Ser Thr Leu Phe Asn Leu Ile Pro Val Gly Leu Arg
100 105 110
Val Val Ala Ile Gln Gly Val Lys Thr Gly Leu Tyr Ile Ala Met Asn
115 120 125
Gly Glu Gly Tyr Leu Tyr Pro Ser Glu Leu Phe Thr Pro Glu Cys Lys
130 135 140
Phe Lys Glu Ser Val Phe Glu Asn Tyr Tyr Val Ile Tyr Ser Ser Met
145 150 155 160
Leu Tyr Arg Gln Gln Glu Ser Gly Arg Ala Trp Phe Leu Gly Leu Asn
165 170 175
Lys Glu Gly Gln Ala Met Lys Gly Asn Arg Val Lys Lys Thr Lys Pro
180 185 190
Ala Ala His Phe Leu Pro Lys Pro Leu Glu Val Ala Met Tyr Arg Glu
195 200 205
Pro Ser Leu His Asp Val Gly Glu Thr Val Pro Lys Pro Gly Val Thr
210 215 220
Pro Ser Lys Ser Thr Ser Ala Ser Ala Ile Met Asn Gly Gly Lys Pro
225 230 235 240
Val Asn Lys Ser Lys Thr Thr
245
<210> 15
<211> 207
<212> PRT
<213> 智人(Homo sapiens)
<400> 15
Met Ala Glu Val Gly Gly Val Phe Ala Ser Leu Asp Trp Asp Leu His
1 5 10 15
Gly Phe Ser Ser Ser Leu Gly Asn Val Pro Leu Ala Asp Ser Pro Gly
20 25 30
Phe Leu Asn Glu Arg Leu Gly Gln Ile Glu Gly Lys Leu Gln Arg Gly
35 40 45
Ser Pro Thr Asp Phe Ala His Leu Lys Gly Ile Leu Arg Arg Arg Gln
50 55 60
Leu Tyr Cys Arg Thr Gly Phe His Leu Glu Ile Phe Pro Asn Gly Thr
65 70 75 80
Val His Gly Thr Arg His Asp His Ser Arg Phe Gly Ile Leu Glu Phe
85 90 95
Ile Ser Leu Ala Val Gly Leu Ile Ser Ile Arg Gly Val Asp Ser Gly
100 105 110
Leu Tyr Leu Gly Met Asn Glu Arg Gly Glu Leu Tyr Gly Ser Lys Lys
115 120 125
Leu Thr Arg Glu Cys Val Phe Arg Glu Gln Phe Glu Glu Asn Trp Tyr
130 135 140
Asn Thr Tyr Ala Ser Thr Leu Tyr Lys His Ser Asp Ser Glu Arg Gln
145 150 155 160
Tyr Tyr Val Ala Leu Asn Lys Asp Gly Ser Pro Arg Glu Gly Tyr Arg
165 170 175
Thr Lys Arg His Gln Lys Phe Thr His Phe Leu Pro Arg Pro Val Asp
180 185 190
Pro Ser Lys Leu Pro Ser Met Ser Arg Asp Leu Phe His Tyr Arg
195 200 205
<210> 16
<211> 195
<212> PRT
<213> 智人(Homo sapiens)
<400> 16
Met Thr Gln Gly Glu Asn His Pro Ser Pro Asn Phe Asn Gln Tyr Val
1 5 10 15
Arg Asp Gln Gly Ala Met Thr Asp Gln Leu Ser Arg Arg Gln Ile Arg
20 25 30
Glu Tyr Gln Leu Tyr Ser Arg Thr Ser Gly Lys His Val Gln Val Thr
35 40 45
Gly Arg Arg Ile Ser Ala Thr Ala Glu Asp Gly Asn Lys Phe Ala Lys
50 55 60
Leu Ile Val Glu Thr Asp Thr Phe Gly Ser Arg Val Arg Ile Lys Gly
65 70 75 80
Ala Glu Ser Glu Lys Tyr Ile Cys Met Asn Lys Arg Gly Lys Leu Ile
85 90 95
Gly Lys Pro Ser Gly Lys Ser Lys Asp Cys Val Phe Thr Glu Ile Val
100 105 110
Leu Glu Asn Asn Tyr Thr Ala Phe Gln Asn Ala Arg His Glu Gly Trp
115 120 125
Phe Met Ala Phe Thr Arg Gln Gly Arg Pro Arg Gln Ala Ser Arg Ser
130 135 140
Arg Gln Asn Gln Arg Glu Ala His Phe Ile Lys Arg Leu Tyr Gln Gly
145 150 155 160
Gln Leu Pro Phe Pro Asn His Ala Glu Lys Gln Lys Gln Phe Glu Phe
165 170 175
Val Gly Ser Ala Pro Thr Arg Arg Thr Lys Arg Thr Arg Arg Pro Gln
180 185 190
Pro Leu Thr
195
<210> 17
<211> 174
<212> PRT
<213> 智人(Homo sapiens)
<400> 17
Met Ala Glu Glu Asn Val Asp Phe Arg Ile His Val Glu Asn Gln Thr
1 5 10 15
Arg Ala Arg Asp Asp Val Ser Arg Lys Gln Leu Arg Leu Tyr Gln Leu
20 25 30
Tyr Ser Arg Thr Ser Gly Lys His Ile Gln Val Leu Gly Arg Arg Ile
35 40 45
Ser Ala Arg Gly Glu Asp Gly Asp Lys Tyr Ala Gln Leu Leu Val Glu
50 55 60
Thr Asp Thr Phe Gly Ser Gln Val Arg Ile Lys Gly Lys Glu Thr Glu
65 70 75 80
Phe Tyr Leu Cys Met Asn Arg Lys Gly Lys Leu Val Gly Lys Pro Asp
85 90 95
Gly Thr Ser Lys Glu Cys Val Phe Ile Glu Lys Val Leu Glu Asn Asn
100 105 110
Tyr Thr Ala Leu Met Ser Ala Lys Tyr Ser Gly Trp Tyr Val Gly Phe
115 120 125
Thr Lys Lys Gly Arg Pro Arg Lys Gly Pro Lys Thr Arg Glu Asn Gln
130 135 140
Gln Asp Val His Phe Met Lys Arg Tyr Pro Lys Gly Gln Pro Glu Leu
145 150 155 160
Gln Lys Pro Phe Lys Tyr Thr Thr Val Thr Lys Arg Ser Arg
165 170
<210> 18
<211> 195
<212> PRT
<213> 智人(Homo sapiens)
<400> 18
Met Arg Pro Leu Ala Phe Ser Asp Ala Gly Pro His Val His Tyr Gly
1 5 10 15
Trp Gly Asp Pro Ile Arg Leu Arg His Leu Tyr Thr Ser Gly Pro His
20 25 30
Gly Leu Ser Ser Cys Phe Leu Arg Ile Arg Ala Asp Gly Val Val Asp
35 40 45
Cys Ala Arg Gly Gln Ser Ala His Ser Leu Leu Glu Ile Lys Ala Val
50 55 60
Ala Leu Arg Thr Val Ala Ile Lys Gly Val His Ser Val Arg Tyr Leu
65 70 75 80
Cys Met Gly Ala Asp Gly Lys Met Gln Gly Leu Leu Gln Tyr Ser Glu
85 90 95
Glu Asp Cys Ala Phe Glu Glu Glu Ile Arg Pro Asp Gly Tyr Asn Val
100 105 110
Tyr Arg Ser Glu Lys His Arg Leu Pro Val Ser Leu Ser Ser Ala Lys
115 120 125
Gln Arg Gln Leu Tyr Lys Asn Arg Gly Phe Leu Pro Leu Ser His Phe
130 135 140
Leu Pro Met Leu Pro Met Val Pro Glu Glu Pro Glu Asp Leu Arg Gly
145 150 155 160
His Leu Glu Ser Asp Met Phe Ser Ser Pro Leu Glu Thr Asp Ser Met
165 170 175
Asp Pro Phe Gly Leu Val Thr Gly Leu Glu Ala Val Arg Ser Pro Ser
180 185 190
Phe Glu Lys
195
<210> 19
<211> 210
<212> PRT
<213> 智人(Homo sapiens)
<400> 19
Met Pro Leu Ala Glu Val Gly Gly Phe Leu Gly Gly Leu Glu Gly Leu
1 5 10 15
Gly Gln Gln Val Gly Ser His Phe Leu Leu Pro Pro Ala Gly Glu Arg
20 25 30
Pro Pro Leu Leu Gly Glu Arg Arg Ser Ala Ala Glu Arg Ser Ala Arg
35 40 45
Gly Gly Pro Gly Ala Ala Gln Leu Ala His Leu His Gly Ile Leu Arg
50 55 60
Arg Arg Gln Leu Tyr Cys Arg Thr Gly Phe His Leu Gln Ile Leu Pro
65 70 75 80
Asp Gly Ser Val Gln Gly Thr Arg Gln Asp His Ser Leu Phe Gly Ile
85 90 95
Leu Glu Phe Ile Ser Val Ala Val Gly Leu Val Ser Ile Arg Gly Val
100 105 110
Asp Ser Gly Leu Tyr Leu Gly Met Asn Asp Lys Gly Glu Leu Tyr Gly
115 120 125
Ser Glu Lys Leu Thr Ser Glu Cys Ile Phe Arg Glu Gln Phe Glu Glu
130 135 140
Asn Trp Tyr Asn Thr Tyr Ser Ser Asn Ile Tyr Lys His Gly Asp Thr
145 150 155 160
Gly Arg Arg Tyr Phe Val Ala Leu Asn Lys Asp Gly Thr Pro Arg Asp
165 170 175
Gly Ala Arg Ser Lys Arg His Gln Lys Phe Thr His Phe Leu Pro Arg
180 185 190
Pro Val Asp Pro Glu Arg Val Pro Glu Leu Tyr Lys Asp Leu Leu Met
195 200 205
Tyr Thr
210
<210> 20
<211> 182
<212> PRT
<213> 智人(Homo sapiens)
<400> 20
Met His Pro Ile Pro Asp Ser Ser Pro Leu Leu Gln Phe Gly Gly Gln
1 5 10 15
Val Arg Gln Arg Tyr Leu Tyr Thr Asp Asp Ala Gln Gln Thr Glu Ala
20 25 30
His Leu Glu Ile Arg Glu Asp Gly Thr Val Gly Gly Ala Ala Asp Gln
35 40 45
Ser Pro Glu Ser Leu Leu Gln Leu Lys Ala Leu Lys Pro Gly Val Ile
50 55 60
Gln Ile Leu Gly Val Lys Thr Ser Arg Phe Leu Cys Gln Arg Pro Asp
65 70 75 80
Gly Ala Leu Tyr Gly Ser Leu His Phe Asp Pro Glu Ala Cys Ser Phe
85 90 95
Arg Glu Leu Leu Leu Glu Asp Gly Tyr Asn Val Tyr Gln Ser Glu Ala
100 105 110
His Gly Leu Pro Leu His Leu Pro Gly Asn Lys Ser Pro His Arg Asp
115 120 125
Pro Ala Pro Arg Gly Pro Ala Arg Phe Leu Pro Leu Pro Gly Leu Pro
130 135 140
Pro Ala Leu Pro Glu Pro Pro Gly Ile Leu Ala Pro Gln Pro Pro Asp
145 150 155 160
Val Gly Ser Ser Asp Pro Leu Ser Met Val Gly Pro Ser Gln Gly Arg
165 170 175
Ser Pro Ser Tyr Ala Ser
180
<210> 21
<211> 150
<212> PRT
<213> 智人(Homo sapiens)
<400> 21
Met Gly Thr Pro Ser Ala Ser Arg Gly Pro Arg Ser Tyr Pro His Leu
1 5 10 15
Glu Gly Asp Val Arg Trp Arg Arg Leu Phe Ser Ser Thr His Phe Phe
20 25 30
Leu Arg Val Asp Pro Gly Gly Arg Val Gln Gly Thr Arg Trp Arg His
35 40 45
Gly Gln Asp Ser Ile Leu Glu Ile Arg Ser Val His Val Gly Val Val
50 55 60
Val Ile Lys Ala Val Ser Ser Gly Phe Tyr Val Ala Met Asn Arg Arg
65 70 75 80
Gly Arg Leu Tyr Gly Ser Arg Leu Tyr Thr Val Asp Cys Arg Phe Arg
85 90 95
Glu Arg Ile Glu Glu Asn Gly His Asn Thr Tyr Ala Ser Gln Arg Trp
100 105 110
Arg Arg Arg Gly Gln Pro Met Phe Leu Ala Leu Asp Arg Arg Gly Gly
115 120 125
Pro Arg Pro Gly Gly Arg Thr Arg Arg Tyr His Leu Ser Ala His Phe
130 135 140
Leu Pro Val Leu Val Ser
145 150
<210> 22
<211> 228
<212> PRT
<213> 智人(Homo sapiens)
<400> 22
Met Tyr Pro Asn Ala Ser Pro Leu Leu Gly Ser Ser Trp Gly Gly Leu
1 5 10 15
Ile His Leu Tyr Thr Ala Thr Ala Arg Asn Ser Tyr His Leu Gln Ile
20 25 30
His Lys Asn Gly His Val Asp Gly Ala Pro His Gln Thr Ile Tyr Ser
35 40 45
Ala Leu Met Ile Arg Ser Glu Asp Ala Gly Phe Val Val Ile Thr Gly
50 55 60
Val Met Ser Arg Arg Tyr Leu Cys Met Asp Phe Arg Gly Asn Ile Phe
65 70 75 80
Gly Ser His Tyr Phe Asp Pro Glu Asn Cys Arg Phe Gln His Gln Thr
85 90 95
Leu Glu Asn Gly Tyr Asp Val Tyr His Ser Pro Gln Tyr His Phe Leu
100 105 110
Val Ser Leu Gly Arg Ala Lys Arg Ala Phe Leu Pro Gly Met Asn Pro
115 120 125
Pro Pro Tyr Ser Gln Phe Leu Ser Arg Arg Asn Glu Ile Pro Leu Ile
130 135 140
His Phe Asn Thr Pro Ile Pro Arg Arg His Thr Arg Ser Ala Glu Asp
145 150 155 160
Asp Ser Glu Arg Asp Pro Leu Asn Val Leu Lys Pro Arg Ala Arg Met
165 170 175
Thr Pro Ala Pro Ala Ser Cys Ser Gln Glu Leu Pro Ser Ala Glu Asp
180 185 190
Asn Ser Pro Met Ala Ser Asp Pro Leu Gly Val Val Arg Gly Gly Arg
195 200 205
Val Asn Thr His Ala Gly Gly Thr Gly Pro Glu Gly Cys Arg Pro Phe
210 215 220
Ala Lys Phe Ile
225
<210> 23
<211> 253
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 23
Met Lys Ser Ser His His His His His His Gly Ser Ser Val Ser Lys
1 5 10 15
Gly Glu Glu Leu Phe Thr Gly Val Val Pro Ile Leu Val Glu Leu Asp
20 25 30
Gly Asp Val Asn Gly His Lys Phe Ser Val Arg Gly Glu Gly Glu Gly
35 40 45
Asp Ala Thr Asn Gly Lys Leu Thr Leu Lys Phe Ile Cys Thr Thr Gly
50 55 60
Lys Leu Pro Val Pro Trp Pro Thr Leu Val Thr Thr Leu Thr Tyr Gly
65 70 75 80
Val Gln Cys Phe Ser Arg Tyr Pro Asp His Met Lys Gln His Asp Phe
85 90 95
Phe Lys Ser Ala Met Pro Glu Gly Tyr Val Gln Glu Arg Thr Ile Ser
100 105 110
Phe Lys Asp Asp Gly Thr Tyr Lys Thr Arg Ala Glu Val Lys Phe Glu
115 120 125
Gly Asp Thr Leu Val Asn Arg Ile Glu Leu Lys Gly Ile Asp Phe Lys
130 135 140
Glu Asp Gly Asn Ile Leu Gly His Lys Leu Glu Tyr Asn Phe Asn Ser
145 150 155 160
His Asn Val Tyr Ile Thr Ala Asp Lys Gln Lys Asn Gly Ile Lys Ala
165 170 175
Asn Phe Lys Ile Arg His Asn Val Glu Asp Gly Ser Val Gln Leu Ala
180 185 190
Asp His Tyr Gln Gln Asn Thr Pro Ile Gly Asp Gly Pro Val Leu Leu
195 200 205
Pro Asp Asn His Tyr Leu Ser Thr Gln Ser Lys Leu Ser Lys Asp Pro
210 215 220
Asn Glu Lys Arg Asp His Met Val Leu Leu Glu Phe Val Thr Ala Ala
225 230 235 240
Gly Ile Thr Leu Gly Met Asp Glu Leu Tyr Lys Gly Ile
245 250
<210> 24
<211> 98
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 24
Met Ser Asp Ser Glu Val Asn Gln Glu Ala Lys Pro Glu Val Lys Pro
1 5 10 15
Glu Val Lys Pro Glu Thr His Ile Asn Leu Lys Val Ser Asp Gly Ser
20 25 30
Ser Glu Ile Phe Phe Lys Ile Lys Lys Thr Thr Pro Leu Arg Arg Leu
35 40 45
Met Glu Ala Phe Ala Lys Arg Gln Gly Lys Glu Met Asp Ser Leu Arg
50 55 60
Phe Leu Tyr Asp Gly Ile Arg Ile Gln Ala Asp Gln Thr Pro Glu Asp
65 70 75 80
Leu Asp Met Glu Asp Asn Asp Ile Ile Glu Ala His Arg Glu Gln Ile
85 90 95
Gly Gly
<210> 25
<211> 8
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 25
Gly Gly Gly Ser Gly Gly Gly Ser
1 5
<210> 26
<211> 426
<212> DNA
<213> 智人(Homo sapiens)
<400> 26
atgttcaacc tgccgccggg caactacaag aaaccgaagc tgctgtattg cagcaacggt 60
ggccactttc tgcgtatcct gccggatggt accgtggatg gtacccgtga ccgtagcgat 120
cagcacatcc agctgcaact gagcgcggag agcgtgggcg aagtttacat taaaagcacc 180
gagaccggcc agtatctggc gatggacacc gatggtctgc tgtacggcag ccaaaccccg 240
aacgaggaat gcctgttcct ggagcgtctg gaggaaaacc actacaacac ctatatcagc 300
aagaaacacg cggaaaagaa ctggtttgtg ggtctgaaga aaaacggcag ctgcaagcgt 360
ggtccgcgta cccactatgg tcaaaaagcg attctgttcc tgccgctgcc ggttagcagc 420
gactaa 426
<210> 27
<211> 468
<212> DNA
<213> 智人(Homo sapiens)
<400> 27
atggcggcgg gtagcatcac caccctgccg gcgctgccgg aagatggtgg cagcggtgcg 60
tttccgccgg gtcacttcaa ggacccgaaa cgtctgtact gcaagaacgg tggcttcttt 120
ctgcgtattc acccggacgg tcgtgtggat ggcgttcgtg aaaagagcga cccgcacatc 180
aaactgcagc tgcaagcgga ggaacgtggt gtggttagca ttaaaggcgt gtgcgcgaac 240
cgttatctgg cgatgaaaga ggacggccgt ctgctggcga gcaaatgcgt taccgatgaa 300
tgcttctttt tcgagcgtct ggaaagcaac aactacaaca cctatcgtag ccgtaagtac 360
accagctggt acgttgcgct gaaacgtacc ggccagtaca agctgggcag caaaaccggt 420
ccgggccaaa aggcgatcct gtttctgccg atgagcgcga aaagctaa 468
<210> 28
<211> 675
<212> DNA
<213> 智人(Homo sapiens)
<400> 28
atgccggcgg cgggtccggg tgcgcgtctg cgtcgtgatg cgggtggccg tggtggcgtg 60
tatgagcacc tgggtggtgc gccgcgtcgt cgtaagctgt actgcgcgac caaatatcac 120
ctgcagctgc acccgagcgg tcgtgtgaac ggcagcctgg aaaacagcgc gtacagcatc 180
ctggagatta ccgcggtgga agttggtatc gttgcgattc gtggtctgtt cagcggccgt 240
tatctggcga tgaacaagcg tggccgtctg tacgcgagcg agcactatag cgcggagtgc 300
gaatttgtgg aacgtatcca cgaactgggt tacaacacct atgcgagccg tctgtaccgt 360
accgttagca gcaccccggg tgcgcgtcgt cagccgagcg cggagcgtct gtggtatgtg 420
agcgttaacg gtaaaggccg tccgcgtcgt ggtttcaaga cccgtcgtac ccagaagagc 480
agcctgtttc tgccgcgtgt gctggaccac cgtgatcacg aaatggttcg tcagctgcaa 540
agcggtctgc cgcgtccgcc gggcaagggc gtgcaaccgc gtcgtcgtcg tcagaaacaa 600
agcccggata acctggaacc gagccatgtt caggcgagcc gtctgggtag ccaactggaa 660
gcgagcgcgc actaa 675
<210> 29
<211> 552
<212> DNA
<213> 智人(Homo sapiens)
<400> 29
atgggtcgtg gtggcgcggc ggcgccgacc gcgccgaacg gtaccctgga ggcggaactg 60
gagcgtcgtt gggaaagcct ggtggcgctg agcctggcgc gtctgccggt tgcggcgcag 120
ccgaaagagg cggcggtgca aagcggtgcg ggtgactacc tgctgggtat caaacgtctg 180
cgtcgtctgt attgcaacgt tggtattggt tttcacctgc aggcgctgcc ggatggtcgt 240
atcggtggcg cgcatgcgga cacccgtgat agcctgctgg aactgagccc ggtggagcgt 300
ggtgtggtta gcatttttgg cgtggcgagc cgtttctttg ttgcgatgag cagcaagggt 360
aaactgtacg gcagcccgtt ctttaccgac gaatgcacct tcaaggagat tctgctgccg 420
aacaactaca acgcgtatga aagctacaaa tatccgggca tgtttatcgc gctgagcaag 480
aacggtaaaa ccaagaaagg caaccgtgtg agcccgacca tgaaggttac ccacttcctg 540
ccgcgtctgt aa 552
<210> 30
<211> 759
<212> DNA
<213> 智人(Homo sapiens)
<400> 30
atggcgtggg cgcatggtga gaagcgtctg gcgccgaagg gtcagccggg tccggcggcg 60
accgaccgta acccgatcgg tagcagcagc cgtcaaagca gcagcagcgc gatgagcagc 120
agcagcgcga gcagcagccc ggcggcgagc ctgggcagcc agggtagcgg tctggaacag 180
agcagctttc agtggagccc gagcggtcgt cgtaccggta gcctgtactg ccgtgttggt 240
atcggctttc acctgcagat ttatccggat ggcaaagtta acggtagcca cgaggcgaac 300
atgctgagcg tgctggaaat tttcgctgtg agccaaggta tcgttggcat tcgtggtgtg 360
ttcagcaaca agtttctggc gatgagcaag aaaggcaagc tgcacgcgag cgcgaaattt 420
accgacgatt gcaagttccg tgagcgtttt caggaaaaca gctacaacac ctatgcgagc 480
gcgatccacc gtaccgagaa aaccggtcgt gaatggtacg tggcgctgaa caagcgtggc 540
aaggcgaaac gtggttgcag cccgcgtgtg aaaccgcaac acattagcac ccacttcctg 600
ccgcgtttta agcagagcga gcaaccggaa ctgagcttca ccgtgaccgt tccggagaag 660
aaaaacccgc cgagcccgat caagagcaaa attccgctga gcgcgccgcg taaaaacacc 720
aacagcgtga agtatcgtct gaaattccgt tttggttaa 759
<210> 31
<211> 510
<212> DNA
<213> 智人(Homo sapiens)
<400> 31
atgggtaccc gtgcgaacaa caccctgctg gatagccgtg gttggggtac cctgctgagc 60
cgtagccgtg cgggtctggc gggtgaaatt gcgggcgtta actgggaaag cggttacctg 120
gtgggcatca agcgtcagcg tcgtctgtat tgcaacgttg gtattggctt ccacctgcaa 180
gtgctgccgg atggtcgtat cagcggcacc cacgaggaaa acccgtacag cctgctggag 240
atcagcaccg ttgaacgtgg tgtggttagc ctgttcggcg tgcgtagcgc gctgtttgtt 300
gcgatgaaca gcaagggtcg tctgtatgcg accccgagct tccaggaaga gtgcaaattt 360
cgtgagaccc tgctgccgaa caactacaac gcgtatgaaa gcgacctgta ccaaggcacc 420
tatattgcgc tgagcaaata cggtcgtgtg aagcgtggca gcaaagttag cccgatcatg 480
accgtgaccc actttctgcc gcgtatttaa 510
<210> 32
<211> 426
<212> DNA
<213> 智人(Homo sapiens)
<400> 32
atgagctacg actatatgga gggtggcgat atccgtgtgc gtcgtctgtt ctgccgtacc 60
cagtggtacc tgcgtattga caagcgtggc aaggttaaag gcacccaaga gatgaagaac 120
aactacaaca tcatggaaat ccgtaccgtg gcggttggta tcgtggcgat taagggcgtt 180
gagagcgaat tctacctggc gatgaacaag gaaggtaaac tgtatgcgaa gaaagagtgc 240
aacgaagatt gcaactttaa ggagctgatc ctggaaaacc actacaacac ctatgcgagc 300
gcgaaatgga cccacaacgg tggcgagatg ttcgtggcgc tgaaccagaa gggtatcccg 360
gttcgtggca agaaaaccaa gaaagaacaa aaaaccgcgc actttctgcc gatggcgatt 420
acctaa 426
<210> 33
<211> 585
<212> DNA
<213> 智人(Homo sapiens)
<400> 33
atgcaggtga ccgttcaaag cagcccgaac ttcacccagc acgtgcgtga gcaaagcctg 60
gttaccgacc agctgagccg tcgtctgatc cgtacctacc aactgtatag ccgtaccagc 120
ggcaagcacg tgcaggttct ggcgaacaaa cgtatcaacg cgatggcgga ggacggtgat 180
ccgttcgcga agctgattgt ggaaaccgac acctttggca gccgtgtgcg tgttcgtggt 240
gcggaaaccg gcctgtacat ctgcatgaac aagaaaggta aactgattgc gaaaagcaac 300
ggcaagggca aagattgcgt gttcaccgag attgttctgg aaaacaacta taccgcgctg 360
caaaacgcga agtacgaggg ctggtatatg gcgttcaccc gtaaaggtcg tccgcgtaag 420
ggcagcaaaa cccgtcagca ccaacgtgaa gttcacttta tgaaacgtct gccgcgtggt 480
caccacacca ccgagcagag cctgcgtttc gaatttctga actacccgcc gtttacccgt 540
agcctgcgtg gcagccaacg tacctgggcg ccggagccgc gttaa 585
<210> 34
<211> 624
<212> DNA
<213> 智人(Homo sapiens)
<400> 34
atgccgctgg gtgaagtggg caactacttc ggcgtgcagg acgcggttcc gtttggtaac 60
gtgccggttc tgccggtgga cagcccggtt ctgctgagcg atcacctggg ccaaagcgag 120
gcgggtggcc tgccgcgtgg tccggcggtt accgacctgg atcacctgaa gggcatcctg 180
cgtcgtcgtc agctgtattg ccgtaccggt ttccacctgg aaatctttcc gaacggtacc 240
attcaaggca cccgtaaaga ccacagccgt ttcggcatcc tggagtttat cagcattgcg 300
gtgggtctgg ttagcattcg tggtgtggat agcggcctgt acctgggtat gaacgagaag 360
ggcgaactgt atggtagcga gaaactgacc caggaatgcg ttttccgtga gcaatttgag 420
gaaaactggt acaacaccta tagcagcaac ctgtacaagc acgtggacac cggtcgtcgt 480
tactatgttg cgctgaacaa agatggcacc ccgcgtgaag gtacccgtac caagcgtcac 540
cagaaattca cccactttct gccgcgtccg gtggacccgg ataaggttcc ggagctgtat 600
aaagatattc tgagccaaag ctaa 624
<210> 35
<211> 513
<212> DNA
<213> 智人(Homo sapiens)
<400> 35
atgctgggtc aggacatggt tagcccggag gcgaccaaca gcagcagcag cagctttagc 60
agcccgagca gcgcgggtcg tcatgtgcgt agctacaacc acctgcaagg cgatgttcgt 120
tggcgtaaac tgttcagctt taccaagtat tttctgaaga tcgagaaaaa cggcaaagtt 180
agcggcacca agaaagaaaa ctgcccgtac agcatcctgg agattaccag cgtggaaatc 240
ggtgtggttg cggttaaagc gattaacagc aactactatc tggcgatgaa caagaaaggt 300
aaactgtatg gcagcaagga gttcaacaac gactgcaagc tgaaagaacg tattgaggaa 360
aacggttaca acacctatgc gagctttaac tggcagcaca acggccgtca aatgtatgtg 420
gcgctgaacg gtaaaggtgc gccgcgtcgt ggtcagaaaa cccgtcgtaa gaacaccagc 480
gcgcacttcc tgccgatggt ggttcacagc taa 513
<210> 36
<211> 678
<212> DNA
<213> 智人(Homo sapiens)
<400> 36
atggcggcgc tggcgagcag cctgatccgt cagaagcgtg aggtgcgcga gccgggtggc 60
agccgtccgg ttagcgcgca acgtcgtgtt tgcccgcgtg gcaccaagag cctgtgccag 120
aaacaactgc tgattctgct gagcaaagtg cgtctgtgcg gtggccgtcc ggcgcgtccg 180
gaccgtggtc cggagccgca gctgaagggt atcgttacca aactgttctg ccgtcagggc 240
ttttacctgc aagcgaaccc ggacggtagc attcaaggca ccccggaaga taccagcagc 300
ttcacccact ttaacctgat cccggttggt ctgcgtgtgg ttaccattca gagcgcgaag 360
ctgggccact acatggcgat gaacgcggag ggtctgctgt atagcagccc gcacttcacc 420
gcggaatgcc gtttcaaaga gtgcgtgttt gaaaactact atgttctgta cgcgagcgcg 480
ctgtatcgtc agcgtcgtag cggtcgtgcg tggtacctgg gtctggataa agagggtcaa 540
gtgatgaaag gtaaccgtgt taagaaaacc aaggcggcgg cgcactttct gccgaaactg 600
ctggaagtgg cgatgtatca agaaccgagc ctgcacagcg ttccggaagc gagcccgagc 660
agcccgccgg cgccgtaa 678
<210> 37
<211> 732
<212> DNA
<213> 智人(Homo sapiens)
<400> 37
atggcggcgg cgatcgcgag cagcctgatt cgtcagaagc gtcaagcgcg tgagagcaac 60
agcgaccgtg tgagcgcgag caagcgtcgt agcagcccga gcaaagatgg ccgtagcctg 120
tgcgaacgtc acgtgctggg tgttttcagc aaagtgcgtt tttgcagcgg ccgtaaacgt 180
ccggttcgtc gtcgtccgga gccgcagctg aaaggtatcg ttacccgtct gttcagccag 240
caaggctact ttctgcaaat gcacccggac ggtaccattg atggcaccaa ggacgaaaac 300
agcgattata ccctgttcaa cctgatcccg gtgggcctgc gtgtggttgc gattcagggt 360
gtgaaagcga gcctgtacgt tgcgatgaac ggcgagggct acctgtatag cagcgacgtt 420
tttaccccgg aatgcaagtt caaagagagc gtgtttgaaa actactatgt tatctacagc 480
agcaccctgt atcgtcagca agagagcggt cgtgcgtggt tcctgggtct gaacaaggaa 540
ggccaaatta tgaaaggtaa ccgtgtgaag aaaaccaagc cgagcagcca ctttgtgccg 600
aaaccgatcg aggtttgcat gtatcgtgaa ccgagcctgc acgagattgg tgaaaagcag 660
ggccgtagcc gtaaaagcag cggcaccccg accatgaacg gtggcaaggt ggttaaccaa 720
gatagcacct aa 732
<210> 38
<211> 738
<212> DNA
<213> 智人(Homo sapiens)
<400> 38
atggcggcgg cgatcgcgag cagcctgatt cgtcaaaagc gtcaggcgcg tgagcgtgaa 60
aagagcaacg cgtgcaaatg cgtgagcagc ccgagcaagg gcaaaaccag ctgcgacaag 120
aacaaactga acgtgttcag ccgtgttaaa ctgtttggta gcaagaagcg tcgtcgtcgt 180
cgtccggagc cgcaactgaa gggcatcgtt accaaactgt acagccgtca gggttatcac 240
ctgcaactgc aggcggacgg taccattgat ggcaccaagg acgaagatag cacctacacc 300
ctgttcaacc tgatcccggt gggcctgcgt gtggttgcga ttcaaggtgt tcagaccaaa 360
ctgtatctgg cgatgaacag cgagggctac ctgtatacca gcgagctgtt taccccggaa 420
tgcaagttca aagagagcgt gtttgaaaac tactatgtta cctacagcag catgatctat 480
cgtcagcaac agagcggtcg tggctggtac ctgggtctga acaaagaggg tgaaattatg 540
aaaggtaacc acgtgaagaa aaacaaaccg gcggcgcact tcctgccgaa gccgctgaaa 600
gttgcgatgt ataaagagcc gagcctgcac gatctgaccg aatttagccg tagcggtagc 660
ggcaccccga ccaagagccg tagcgtgagc ggtgttctga acggtggcaa aagcatgagc 720
cacaacgaaa gcacctaa 738
<210> 39
<211> 744
<212> DNA
<213> 智人(Homo sapiens)
<400> 39
atggcggcgg cgattgcgag cggtctgatt cgtcagaagc gtcaagcgcg tgaacaacac 60
tgggaccgtc cgagcgcgag ccgtcgtcgt agcagcccga gcaagaaccg tggtctgtgc 120
aacggcaacc tggtggacat cttcagcaaa gttcgtattt ttggtctgaa gaaacgtcgt 180
ctgcgtcgtc aggacccgca gctgaaaggc attgtgaccc gtctgtactg ccgtcagggt 240
tactatctgc aaatgcaccc ggacggtgcg ctggatggca ccaaggacga tagcaccaac 300
agcaccctgt tcaacctgat cccggtgggc ctgcgtgtgg ttgcgattca gggtgttaaa 360
accggcctgt atatcgcgat gaacggcgag ggctacctgt atccgagcga gctgtttacc 420
ccggaatgca agttcaaaga gagcgtgttc gaaaactact acgttatcta cagcagcatg 480
ctgtatcgtc agcaagagag cggtcgtgcg tggttcctgg gtctgaacaa ggaaggccaa 540
gcgatgaaag gtaaccgtgt gaagaaaacc aagccggcgg cgcactttct gccgaaaccg 600
ctggaagtgg cgatgtaccg tgaaccgagc ctgcacgatg tgggcgaaac cgttccgaag 660
ccgggtgtga ccccgagcaa aagcaccagc gcgagcgcga tcatgaacgg tggcaagccg 720
gttaacaaga gcaaaaccac ctaa 744
<210> 40
<211> 624
<212> DNA
<213> 智人(Homo sapiens)
<400> 40
atggcggaag tgggtggtgt gttcgcgagc ctggactggg atctgcacgg ctttagcagc 60
agcctgggta acgtgccgct ggcggacagc ccgggcttcc tgaacgagcg tctgggtcag 120
atcgaaggca agctgcaacg tggcagcccg accgattttg cgcacctgaa aggtatcctg 180
cgtcgtcgtc agctgtactg ccgtaccggt ttccacctgg agatttttcc gaacggtacc 240
gtgcatggta cccgtcatga ccacagccgt ttcggcatcc tggaatttat tagcctggcg 300
gtgggtctga tcagcattcg tggtgttgat agcggcctgt acctgggtat gaacgagcgt 360
ggcgaactgt atggtagcaa gaaactgacc cgtgagtgcg ttttccgtga acaatttgag 420
gaaaactggt acaacaccta tgcgagcacc ctgtacaagc acagcgacag cgagcgtcag 480
tactatgtgg cgctgaacaa agatggcagc ccgcgtgaag gttatcgtac caagcgtcac 540
caaaaattca cccactttct gccgcgtccg gttgacccga gcaagctgcc gagcatgagc 600
cgtgacctgt tccactatcg ttaa 624
<210> 41
<211> 588
<212> DNA
<213> 智人(Homo sapiens)
<400> 41
atgacccaag gcgagaacca cccgagcccg aacttcaacc agtacgtgcg tgaccaaggt 60
gcgatgaccg atcagctgag ccgtcgtcaa attcgtgaat accagctgta tagccgtacc 120
agcggcaagc acgtgcaggt taccggtcgt cgtattagcg cgaccgcgga ggacggcaac 180
aagttcgcga aactgatcgt ggaaaccgat acctttggca gccgtgttcg tatcaagggt 240
gcggagagcg aaaaatacat ttgcatgaac aagcgtggta aactgatcgg caaaccgagc 300
ggcaagagca aagactgcgt gttcaccgag attgttctgg aaaacaacta taccgcgttt 360
caaaacgcgc gtcacgaggg ctggttcatg gcgtttaccc gtcagggtcg tccgcgtcag 420
gcgagccgta gccgtcagaa ccaacgtgaa gcgcacttca tcaagcgtct gtatcagggc 480
caactgccgt ttccgaacca cgcggagaag cagaaacaat tcgaatttgt tggtagcgcg 540
ccgacccgtc gtaccaaacg tacccgtcgt ccgcagccgc tgacctaa 588
<210> 42
<211> 525
<212> DNA
<213> 智人(Homo sapiens)
<400> 42
atggcggagg aaaacgtgga cttccgtatc cacgttgaga accagacccg tgcgcgtgac 60
gatgtgagcc gtaagcagct gcgtctgtac caactgtata gccgtaccag cggcaaacac 120
atccaagttc tgggtcgtcg tattagcgcg cgtggtgagg acggcgataa gtacgcgcag 180
ctgctggttg agaccgacac cttcggcagc caagttcgta tcaagggtaa agagaccgaa 240
ttttatctgt gcatgaaccg taagggtaaa ctggtgggca agccggatgg taccagcaaa 300
gaatgcgtgt tcattgagaa ggttctggaa aacaactaca ccgcgctgat gagcgcgaaa 360
tacagcggct ggtacgttgg ttttaccaag aaaggccgtc cgcgtaaggg tccgaaaacc 420
cgtgagaacc agcaagatgt tcacttcatg aagcgttacc cgaaaggcca gccggaactg 480
caaaagccgt ttaaatatac caccgttacc aaacgtagcc gttaa 525
<210> 43
<211> 588
<212> DNA
<213> 智人(Homo sapiens)
<400> 43
atgcgtccgc tggcgtttag cgatgcgggt ccgcatgtgc actatggttg gggcgatccg 60
atccgtctgc gtcacctgta taccagcggt ccgcatggtc tgagcagctg ctttctgcgt 120
attcgtgcgg acggtgtggt tgattgcgcg cgtggtcaga gcgcgcacag cctgctggaa 180
atcaaggcgg tggcgctgcg taccgttgcg attaaaggtg tgcacagcgt tcgttacctg 240
tgcatgggtg cggacggcaa gatgcagggc ctgctgcaat atagcgagga agactgcgcg 300
ttcgaggaag agatccgtcc ggatggctac aacgtgtatc gtagcgagaa acaccgtctg 360
ccggttagcc tgagcagcgc gaagcagcgt caactgtaca aaaaccgtgg tttcctgccg 420
ctgagccact ttctgccgat gctgccgatg gttccggaag agccggaaga cctgcgtggc 480
cacctggaga gcgatatgtt cagcagcccg ctggaaaccg acagcatgga cccgtttggt 540
ctggtgaccg gcctggaagc ggttcgtagc ccgagctttg agaagtaa 588
<210> 44
<211> 633
<212> DNA
<213> 智人(Homo sapiens)
<400> 44
atgccgctgg cggaagtggg tggtttcctg ggtggtctgg aaggtctggg ccagcaagtt 60
ggcagccact ttctgctgcc gccggcgggt gaacgtccgc cgctgctggg cgagcgtcgt 120
agcgcggcgg aacgtagcgc gcgtggtggc ccgggtgcgg cgcagctggc gcacctgcac 180
ggtatcctgc gtcgtcgtca gctgtactgc cgtaccggtt tccacctgca aattctgccg 240
gatggtagcg tgcagggtac ccgtcaagat cacagcctgt tcggcatcct ggaatttatt 300
agcgtggcgg ttggtctggt tagcatccgt ggtgttgaca gcggcctgta cctgggtatg 360
aacgataagg gcgagctgta tggtagcgaa aaactgacca gcgagtgcat cttccgtgaa 420
cagtttgagg aaaactggta caacacctat agcagcaaca tttacaagca cggtgacacc 480
ggccgtcgtt attttgttgc gctgaacaaa gatggtaccc cgcgtgatgg tgcgcgtagc 540
aagcgtcacc aaaaattcac ccactttctg ccgcgtccgg tggacccgga gcgtgttccg 600
gaactgtaca aggatctgct gatgtatacc taa 633
<210> 45
<211> 549
<212> DNA
<213> 智人(Homo sapiens)
<400> 45
atgcatccga ttccggacag cagcccgctg ctgcagtttg gtggccaagt gcgtcaacgt 60
tacctgtata ccgacgatgc gcagcaaacc gaggcgcacc tggagattcg tgaagacggt 120
accgttggtg gcgcggcgga tcagagcccg gaaagcctgc tgcaactgaa ggcgctgaaa 180
ccgggtgtga tccagattct gggcgttaag accagccgtt ttctgtgcca acgtccggat 240
ggtgcgctgt atggcagcct gcacttcgat ccggaggcgt gcagctttcg tgagctgctg 300
ctggaagacg gctacaacgt gtatcaaagc gaagcgcatg gtctgccgct gcacctgccg 360
ggtaacaaga gcccgcaccg tgatccggcg ccgcgtggtc cggcgcgttt cctgccgctg 420
ccgggtctgc cgccggcgct gccggagccg ccgggtatcc tggcgccgca gccgccggac 480
gtgggcagca gcgatccgct gagcatggtt ggtccgagcc aaggccgtag cccgagctat 540
gcgagctaa 549
<210> 46
<211> 453
<212> DNA
<213> 智人(Homo sapiens)
<400> 46
atgggtaccc cgagcgcgag ccgtggtccg cgtagctacc cgcacctgga gggtgacgtt 60
cgttggcgtc gtctgttcag cagcacccac ttctttctgc gtgttgaccc gggtggccgt 120
gtgcagggta cccgttggcg tcacggccaa gatagcatcc tggaaattcg tagcgttcac 180
gtgggtgtgg ttgtgatcaa ggcggttagc agcggcttct atgtggcgat gaaccgtcgt 240
ggtcgtctgt acggcagccg tctgtatacc gtggactgcc gttttcgtga gcgtattgag 300
gaaaacggtc acaacaccta cgcgagccag cgttggcgtc gtcgtggtca accgatgttc 360
ctggcgctgg atcgtcgtgg tggcccgcgt ccgggtggcc gtacccgtcg ttatcacctg 420
agcgcgcact ttctgccggt tctggttagc taa 453
<210> 47
<211> 687
<212> DNA
<213> 智人(Homo sapiens)
<400> 47
atgtatccga acgcgagccc gctgctgggt agcagctggg gtggcctgat ccacctgtac 60
accgcgaccg cgcgtaacag ctatcacctg cagattcaca aaaacggtca tgtggatggt 120
gcgccgcacc aaaccatcta tagcgcgctg atgattcgta gcgaggatgc gggttttgtg 180
gttatcaccg gcgttatgag ccgtcgttac ctgtgcatgg acttccgtgg taacattttt 240
ggcagccact atttcgatcc ggagaactgc cgtttccagc accaaaccct ggaaaacggc 300
tacgacgtgt atcacagccc gcagtaccac tttctggtta gcctgggtcg tgcgaagcgt 360
gcgttcctgc cgggtatgaa cccgccgccg tatagccaat ttctgagccg tcgtaacgag 420
atcccgctga ttcacttcaa caccccgatt ccgcgtcgtc acacccgtag cgcggaggac 480
gatagcgaac gtgatccgct gaacgtgctg aaaccgcgtg cgcgtatgac cccggcgccg 540
gcgagctgca gccaggagct gccgagcgcg gaagacaaca gcccgatggc gagcgatccg 600
ctgggtgtgg ttcgtggtgg ccgtgttaac acccatgcgg gtggcaccgg tccggaaggt 660
tgccgtccgt tcgcgaaatt catctaa 687
<210> 48
<211> 762
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 48
atgaagagca gccatcacca ccaccaccat ggcagcagcg tgagcaaagg cgaggaactg 60
ttcaccggcg tggttccgat cctggtggaa ctggacggcg atgttaacgg tcacaagttt 120
agcgttcgtg gtgagggcga aggtgatgcg accaacggca agctgaccct gaaattcatt 180
tgcaccaccg gtaaactgcc ggtgccgtgg ccgaccctgg ttaccaccct gacctacggt 240
gtgcagtgct ttagccgtta tccggaccac atgaaacaac acgatttctt taagagcgcg 300
atgccggaag gttacgttca ggagcgtacc atcagcttca aagacgatgg cacctataag 360
acccgtgcgg aagtgaaatt tgaaggtgat accctggtta accgtatcga actgaaaggc 420
attgacttca aagaggacgg caacatcctg ggtcacaagc tggaatacaa ctttaacagc 480
cacaacgtgt atattaccgc ggacaagcag aaaaacggta tcaaggcgaa ctttaaaatt 540
cgtcacaacg tggaggacgg cagcgttcaa ctggcggatc actaccagca aaacaccccg 600
attggtgatg gtccggttct gctgccggat aaccactatc tgagcaccca gagcaagctg 660
agcaaagacc cgaacgaaaa gcgtgatcac atggtgctgc tggagttcgt taccgcggcg 720
ggcatcaccc tgggtatgga tgaactgtac aaaggtattg ag 762
<210> 49
<211> 294
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 49
atgagcgaca gcgaggtgaa ccaagaagcg aagccggaag tgaaaccgga agttaagccg 60
gagacccaca tcaacctgaa agttagcgac ggcagcagcg agatcttctt taaaattaag 120
aaaaccaccc cgctgcgtcg tctgatggaa gcgttcgcga aacgtcaggg caaggagatg 180
gacagcctgc gttttctgta tgatggcatc cgtattcagg cggaccaaac cccggaagac 240
ctggatatgg aggacaacga tatcattgaa gcgcaccgtg agcaaattgg tggc 294
<210> 50
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 50
ggcggcggct ccggcggcgg ctcc 24
<210> 51
<211> 232
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 51
Met Lys Ser Ser His His His His His His Gly Ser Ser Leu Val Pro
1 5 10 15
Glu Leu Asn Glu Lys Asp Asp Asp Gln Val Gln Lys Ala Leu Ala Ser
20 25 30
Arg Glu Asn Thr Gln Leu Met Asn Arg Asp Asn Ile Glu Ile Thr Val
35 40 45
Arg Asp Phe Lys Thr Leu Ala Pro Arg Arg Trp Leu Asn Asp Thr Ile
50 55 60
Ile Glu Phe Phe Met Lys Tyr Ile Glu Lys Ser Thr Pro Asn Thr Val
65 70 75 80
Ala Phe Asn Ser Phe Phe Tyr Thr Asn Leu Ser Glu Arg Gly Tyr Gln
85 90 95
Gly Val Arg Arg Trp Met Lys Arg Lys Lys Thr Gln Ile Asp Lys Leu
100 105 110
Asp Lys Ile Phe Thr Pro Ile Asn Leu Asn Gln Ser His Trp Ala Leu
115 120 125
Gly Ile Ile Asp Leu Lys Lys Lys Thr Ile Gly Tyr Val Asp Ser Leu
130 135 140
Ser Asn Gly Pro Asn Ala Met Ser Phe Ala Ile Leu Thr Asp Leu Gln
145 150 155 160
Lys Tyr Val Met Glu Glu Ser Lys His Thr Ile Gly Glu Asp Phe Asp
165 170 175
Leu Ile His Leu Asp Cys Pro Gln Gln Pro Asn Gly Tyr Asp Cys Gly
180 185 190
Ile Tyr Val Cys Met Asn Thr Leu Tyr Gly Ser Ala Asp Ala Pro Leu
195 200 205
Asp Phe Asp Tyr Lys Asp Ala Ile Arg Met Arg Arg Phe Ile Ala His
210 215 220
Leu Ile Leu Thr Asp Ala Leu Lys
225 230
<210> 52
<211> 699
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 52
atgaaaagca gccaccacca ccaccaccac ggtagcagcc tggtgccgga gctgaacgaa 60
aaagacgatg accaggttca aaaggcgctg gcgagccgtg agaacaccca gctgatgaac 120
cgtgataaca tcgaaattac cgtgcgtgac ttcaagaccc tggcgccgcg tcgttggctg 180
aacgatacca tcatcgagtt ctttatgaag tacatcgaaa agagcacccc gaacaccgtg 240
gcgtttaaca gcttctttta caccaacctg agcgagcgtg gttatcaggg cgttcgtcgt 300
tggatgaagc gtaagaaaac ccaaatcgat aaactggaca agatcttcac cccgattaac 360
ctgaaccaga gccactgggc gctgggcatc attgatctga agaaaaagac catcggttac 420
gtggacagcc tgagcaacgg cccgaacgcg atgagcttcg cgattctgac cgatctgcaa 480
aaatatgtta tggaggaaag caagcacacc atcggtgaag attttgacct gattcacctg 540
gattgcccgc agcaaccgaa cggttacgac tgcggcatct atgtttgcat gaacaccctg 600
tatggcagcg cggatgcgcc gctggatttc gactataaag acgcgattcg tatgcgtcgt 660
tttatcgcgc acctgattct gaccgacgcg ctgaagtaa 699
<210> 53
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 53
tttgtttaac tttaagaagg aga 23
Claims (26)
1.一种组合物，其包括三(2-羰基乙基)磷盐酸盐和硫酸锌，所述三(2-羰基乙基)磷盐酸盐和所述硫酸锌的质量比是1:（1至50）。
2.一种获得含有成纤维细胞生长因子的融合蛋白或其中的所述成纤维细胞生长因子的方法，所述融合蛋白包括从N端到C端的可视化报告蛋白、小泛素样修饰因子和成纤维细胞生长因子，所述方法包括如下步骤：
1-1）将所述融合蛋白的表达盒或编码所述融合蛋白的核酸连接到第一表达载体上，得到第一重组表达载体；其中，所述第一表达载体为异丙基-β-D-硫代半乳糖苷诱导型的表达载体；
2-1）将用于表达小泛素样修饰因子蛋白酶的表达盒或编码小泛素样修饰因子蛋白酶的核酸连接到所述第一重组表达载体上，得到第二重组表达载体；
3-1）将所述第二重组表达载体转化至第一宿主中，得到第一重组生物体；
4-1）在第一培养液中培养所述第一重组生物体至表达有所述融合蛋白和所述小泛素样修饰因子蛋白酶，得到第一培养物；
其中，在培养温度降至10至15°C时，在所述第一培养液中加入如权利要求1所述的组合物；
5-1）纯化所述第一培养物从而得到所述成纤维细胞生长因子；
或者
1-2）同1-1）；
2-2）将所述用于表达小泛素样修饰因子蛋白酶的表达盒或编码小泛素样修饰因子蛋白酶的核酸连接到第二表达载体上，得到第三重组表达载体；
3-2）将所述第一重组表达载体转化至第二宿主中，得到第二重组生物体；将所述第三重组表达载体转化至第三宿主中，得到第三重组生物体；
4-2）在第二培养液培养所述第二重组生物体至表达有所述融合蛋白，得到第二培养物；在第三培养液培养所述第三重组生物体至表达有所述小泛素样修饰因子蛋白酶，得到第三培养物；将所述第二培养物与所述第三培养物混合，得到第四培养物；
其中，在所述第二培养液中加入如权利要求1所述的组合物；
5-2）纯化所述第四培养物，从而得到所述成纤维细胞生长因子。
3.根据权利要求2所述的方法，其特征在于，所述成纤维细胞生长因子为人源成纤维细胞生长因子-1、人源成纤维细胞生长因子-2、人源成纤维细胞生长因子-3、人源成纤维细胞生长因子-4、人源成纤维细胞生长因子-5、人源成纤维细胞生长因子-6、人源成纤维细胞生长因子-7、人源成纤维细胞生长因子-8、人源成纤维细胞生长因子-9、人源成纤维细胞生长因子-10、人源成纤维细胞生长因子-11、人源成纤维细胞生长因子-12、人源成纤维细胞生长因子-13、人源成纤维细胞生长因子-14、人源成纤维细胞生长因子-16、人源成纤维细胞生长因子-17、人源成纤维细胞生长因子-18、人源成纤维细胞生长因子-19、人源成纤维细胞生长因子-20、人源成纤维细胞生长因子-21、人源成纤维细胞生长因子-22、人源成纤维细胞生长因子-23中的一种。
4.根据权利要求3所述的方法，其特征在于，所述人源成纤维细胞生长因子-1的氨基酸序列如SEQ ID No. 1所示，所述人源成纤维细胞生长因子-2的氨基酸序列如SEQ ID No. 2所示，所述人源成纤维细胞生长因子-3的氨基酸序列如SEQ ID No. 3所示，所述人源成纤维细胞生长因子-4的氨基酸序列如SEQ ID No. 4所示，所述人源成纤维细胞生长因子-5的氨基酸序列如SEQ ID No. 5 所示，所述人源成纤维细胞生长因子-6的氨基酸序列如SEQID No. 6所示，所述人源成纤维细胞生长因子-7的氨基酸序列如SEQ ID No. 7所示，所述人源成纤维细胞生长因子-8的氨基酸序列如SEQ ID No.8所示，所述人源成纤维细胞生长因子-9的氨基酸序列如SEQ ID No. 9 所示，所述人源成纤维细胞生长因子-10的氨基酸序列如SEQ ID No. 10所示，所述人源成纤维细胞生长因子-11的氨基酸序列如SEQ ID No.11所示，所述人源成纤维细胞生长因子-12的氨基酸序列如SEQ ID No. 12 所示，所述人源成纤维细胞生长因子-13的氨基酸序列如SEQ ID No. 13 所示，所述人源成纤维细胞生长因子-14的氨基酸序列如SEQ ID No. 14所示，所述人源成纤维细胞生长因子-16的氨基酸序列如SEQ ID No. 15 所示，所述人源成纤维细胞生长因子-17的氨基酸序列如SEQ IDNo. 16所示，所述人源成纤维细胞生长因子-18的氨基酸序列如SEQ ID No. 17所示，所述人源成纤维细胞生长因子-19的氨基酸序列如SEQ ID No. 18 所示，所述人源成纤维细胞生长因子-20的氨基酸序列如SEQ ID No. 19所示，所述人源成纤维细胞生长因子-21的氨基酸序列如SEQ ID No. 20所示，所述人源成纤维细胞生长因子-22的氨基酸序列如SEQ IDNo. 21 所示，所述人源成纤维细胞生长因子-23的氨基酸序列如SEQ ID No. 22 所示。
5.根据权利要求4所述的方法，其特征在于，编码所述人源成纤维细胞生长因子-1的核苷酸如SEQ ID No. 26所示，编码所述人源成纤维细胞生长因子-2的核苷酸序列如SEQ IDNo. 27所示，编码所述人源成纤维细胞生长因子-3的核苷酸序列如SEQ ID No. 28所示，编码所述人源成纤维细胞生长因子-4的核苷酸序列如SEQ ID No. 29所示，编码所述人源成纤维细胞生长因子-5的核苷酸序列如SEQ ID No. 30所示，编码所述人源成纤维细胞生长因子-6的核苷酸序列如SEQ ID No. 31所示，编码所述人源成纤维细胞生长因子-7的核苷酸序列如SEQ ID No. 32所示，编码所述人源成纤维细胞生长因子-8的核苷酸序列如SEQID No. 33所示，编码所述人源成纤维细胞生长因子-9的核苷酸序列如SEQ ID No. 34所示，编码所述人源成纤维细胞生长因子-10的核苷酸序列如SEQ ID No. 35所示，编码所述人源成纤维细胞生长因子-11的核苷酸序列如SEQ ID No. 36所示，编码所述人源成纤维细胞生长因子-12的核苷酸序列如SEQ ID No. 37所示，编码所述人源成纤维细胞生长因子-13的核苷酸序列如SEQ ID No. 38所示，编码所述人源成纤维细胞生长因子-14的核苷酸序列如SEQ ID No. 39所示，编码所述人源成纤维细胞生长因子-16的核苷酸序列如SEQ IDNo. 40所示，编码所述人源成纤维细胞生长因子-17的核苷酸序列如SEQ ID No. 41 所示，编码所述人源成纤维细胞生长因子-18的核苷酸序列如SEQ ID No. 42所示，编码所述人源成纤维细胞生长因子-19的核苷酸序列如SEQ ID No. 43所示，编码所述人源成纤维细胞生长因子-20的核苷酸序列如SEQ ID No. 44所示，编码所述人源成纤维细胞生长因子-21的核苷酸序列如SEQ ID No. 45所示，编码所述人源成纤维细胞生长因子-22的核苷酸序列如SEQ ID No. 46所示，编码所述人源成纤维细胞生长因子-23的核苷酸序列如SEQ ID No.47所示。
6.根据权利要求2所述的方法，其特征在于，所述融合蛋白的表达盒包括从5’端到3’端的第一启动子、编码所述融合蛋白的核酸和第一终止子。
7.根据权利要求2所述的方法，其特征在于，所述可视化报告蛋白为荧光蛋白。
8.根据权利要求2所述的方法，其特征在于，所述可视化报告蛋白为绿色荧光蛋白、红色荧光蛋白、蓝色荧光蛋白和青色荧光蛋白中的一种。
9.根据权利要求8所述的方法，其特征在于，所述绿色荧光蛋白的氨基酸序列如SEQ IDNo. 23所示。
10.根据权利要求9所述的方法，其特征在于，编码所述绿色荧光蛋白的核苷酸序列如SEQ ID No. 48所示。
11.根据权利要求2所述的方法，其特征在于，所述小泛素样修饰因子的氨基酸序列如SEQ ID No. 24 所示。
12.根据权利要求11所述的方法，其特征在于，编码所述小泛素样修饰因子的核苷酸序列如SEQ ID No. 49所示。
13.根据权利要求2所述的方法，其特征在于，所述可视化报告蛋白和所述小泛素样修饰因子之间由多肽接头连接。
14.根据权利要求13所述的方法，其特征在于，所述多肽接头的氨基酸序列如SEQ IDNo.25所示。
15.根据权利要求14所述的方法，其特征在于，编码所述多肽接头的核苷酸序列如SEQID No. 50所示。
16.根据权利要求6所述的方法，其特征在于，所述第一启动子为T7启动子，所述第一终止子为T7终止子。
17.根据权利要求2所述的方法，其特征在于，所述用于表达小泛素样修饰因子蛋白酶的表达盒包括从5’端到3’端的第二启动子、3’-UTR、编码小泛素样修饰因子蛋白酶的核酸和5’-UTR。
18.根据权利要求17所述的方法，其特征在于，所述第二启动子为CspA启动子，所述3’-UTR为CspA 3’-UTR，所述5’-UTR为CspA5’-UTR。
19.根据权利要求2或17所述的方法，其特征在于，所述小泛素样修饰因子蛋白酶的氨基酸序列如SEQ ID No. 51所示。
20.根据权利要求19所述的方法，其特征在于，编码小泛素样修饰因子蛋白酶的核苷酸序列如SEQ ID No. 52所示。
21.根据权利要求2所述的方法，其特征在于，所述第一培养液、所述第二培养液和所述第三培养液独立地为LB培养液或含葡萄糖的LB培养液。
22.根据权利要求2所述的方法，其特征在于，所述第一表达载体为pET系列的表达载体中的一种；
所述第二表达载体为pCold；
所述第一宿主、所述第二宿主和所述第三宿主独立地为大肠杆菌（Escherichiacoli）。
23.根据权利要求22所述的方法，其特征在于，所述第一表达载体为pET21a、pET20b和pET28a中的一种。
24.根据权利要求22所述的方法，其特征在于，所述大肠杆菌的菌株为BL21（DE3）。
25.根据权利要求2所述的方法，其特征在于，在步骤4-2）或步骤3-3）中，培养所述第二重组生物体的温度为28至37°C，时间为4至8小时；
在步骤4-1）中，在28至37°C的温度下，在异丙基-β-D-硫代半乳糖苷的诱导下培养所述第一重组生物体4至8小时，然后在10至15°C的温度下继续培养所述第一重组生物体15至60分钟；
在步骤4-2）中，培养所述第三重组生物体的温度为10至15°C，时间为15至60分钟。
26.根据权利要求2所述的方法，其特征在于，在步骤4-1）中，所述三(2-羰基乙基)磷盐酸盐在所述第一培养液中的终浓度为0.1至1 毫摩尔/升；所述硫酸锌所述第一培养液中的终浓度为1至5 毫摩尔/升；
在步骤4-2）中，所述三(2-羰基乙基)磷盐酸盐在所述第二培养液中的终浓度为0.1至1毫摩尔/升；所述硫酸锌所述第二培养液中的终浓度为1至5 毫摩尔/升。
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