CN111164226A - 评估g蛋白活化的系统和方法 - Google Patents
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Abstract
描述了用于在各种细胞区室处，诸如质膜和胞内体处，并以Gα蛋白亚基家族选择性的方式监测G蛋白活化的系统和方法。这些系统和方法还允许监测G蛋白偶联受体(GPCR)介导的以及非受体鸟嘌呤核苷酸交换因子(GEF)介导的G蛋白活化，并且这些系统和方法是基于标记有合适的能量(BRET)供体和接受体的G蛋白的特异性效应子的G蛋白结合结构域和细胞区室标志物的使用。
Description
相关申请的交叉引用
本申请要求于2017年10月18日提交的美国临时申请No.62/573,853的优先权，其通过引用整体并入本文。
技术领域
本发明总体上涉及细胞中G蛋白活化的监测。
背景技术
异三聚体G蛋白是G蛋白偶联受体(GPCR)的经典信号转导伴侣，并且还已知通过Gα相互作用的囊泡相关蛋白(GIV；也称为Girdin)参与其他类型受体的信号转导，特别是受体酪氨酸激酶(RTK)(Ghosh P,2016,Pharmacol Res.105:99-107)。受体活化触发了Gα结合的GDP交换成GTP，导致异三聚体G蛋白的构象重排，从而促进Gα和Gβ/γ亚基的解离。然后GTP结合的Gα和游离Gβ/γ亚基可用于接合特异性效应子。
根据Gα亚基活化后的信号转导结果，G蛋白可分为多个家族。Gs家族(Gαs，Gαolf)通过直接活化腺苷酸环化酶(AC)来促进cAMP的产生。相反，Gi家族(Gαi1、Gαi2、Gαi3、GαoA、GαoB和Gαz)通过抑制特定AC来降低cAMP水平。Gq家族(Gαq、Gα11、Gα14和Gα15)活化磷脂酶Cβ(PLCβ)产生第二种信使二酰基甘油(DAG)和肌醇三磷酸酯(IP3)，随后分别促进了PKC的活化和Ca2+从内质网释放。最后，已知G12/13家族(Gα12和Gα13)控制Rho-GEF，诸如LARG、p115和TRIO，并因此影响与细胞骨架重塑相关的过程(例如，趋化性)。鉴于活化属于不同家族的G蛋白后信号转导的显著结果，需要允许以G蛋白家族选择性方式监测G蛋白活化的系统和测定法，例如，以更好地表征各种配体引起的细胞表面受体活化(例如，由给定配体活化的信号转导通路)以及识别G蛋白的更特异性的调节物。
尽管细胞表面受体活化主要发生在对配体结合应答的质膜(PM)处，但已显示细胞表面受体(例如，GPCR、RTK)的信号转导发生在其他细胞区室中，包括胞内体和高尔基体。活化后，许多细胞表面受体进入胞内体，胞内体中配体-受体复合物的运输提供了一种机制，该机制或通过降解受体来终止信号转导(在溶酶体和蛋白酶体中)，或通过将受体循环回细胞表面来维持信号转导。但是，已证明受体信号转导也可以在胞内体中启动、持续和终止(Murphy等人,2009,PNAS,vol.106no.42,17615-17622；Tsvetanova等人,2015,J.Biol.Chem.290(11),pp.6689-6696；Vilardaga等人,2014,Nature Chemical Biology10,700-706)。类似地，已发现一些G蛋白与以下相关并在该处被活化：高尔基体(Lo等人,2015,Dev.Cell.33:189-203)以及内质网(ER)-高尔基体界面(Bastin等人,Front BioengBiotechnol.2015Sep 1；3:128)。PM和其他区室(诸如胞内体和高尔基体)的受体会产生不同的信号，从而导致截然不同的生理应答，并且不同的机制调节这些区室的受体信号转导。这些不同的信号转导和调节机制提高了基于靶向其他细胞区室(而不是PM)的信号转导的新疗法的可能性，因此需要用于监测不同细胞区室处的G蛋白活化的系统和测定法。
美国专利号9,029,097和WO/2016/058094公开了用于监测G蛋白活化的基于BRET的生物传感器。然而，这些生物传感器需要对一种或多种G蛋白亚基(Gα、Gβ和/或Gγ)和/或GPCR进行标记，这可能会影响这些蛋白质的活性。而且，这些生物传感器检测细胞中的整体G蛋白活化，但不允许监测在不同的细胞区室的G蛋白活化以及以G蛋白家族选择性方式监测G蛋白活化。
本说明书涉及多个文献，其内容通过引用整体并入本文。
发明内容
本公开提供以下1至67项目：
1.一种以Gα蛋白亚基家族选择性方式测量G蛋白活化调节的系统，所述系统包括表达以下的细胞：
(i)第一组分，包括用生物发光供体分子或荧光接受体分子标记的Gα亚基相互作用多肽(GASIP)；
其中：如果所述Gα蛋白亚基家族是Gi，则所述GASIP包含与Gi特异性结合的蛋白结构域；如果所述Gα蛋白亚基家族是Gq，则所述GASIP包含与Gq特异性结合的蛋白结构域；以及如果所述Gα蛋白亚基家族是G12/13，则所述GASIP包含与G12/13特异性结合的蛋白结构域；以及
(ii)第二组分，包括生物发光供体分子或荧光接受体分子标记的质膜(PM)靶向部分、胞内体靶向部分或高尔基体靶向部分；
其中，如果所述GASIP用所述荧光接受体分子标记，则所述PM靶向部分、胞内体靶向部分或高尔基体靶向部分用所述生物发光供体分子标记，以及如果所述GASIP用所述生物发光供体分子标记，则所述PM靶向部分、胞内体靶向部分或高尔基体靶向部分用所述荧光接受体分子标记。
2.项目1的系统，其中所述与Gi特异性结合的蛋白结构域是Rap1GAP的G蛋白结合结构域或G蛋白信号转导调节物(RGS)蛋白的G蛋白结合结构域。
3.项目2的系统，其中所述GASIP包含Rap1GAP的G蛋白结合结构域。
4.项目3的系统，其中所述Rap1GAP的G蛋白结合结构域包含Rap1GAP(SEQ ID NO：8)的1至442位的残基或其变体，其中在位置437、439和441的一个或多个丝氨酸残基突变或缺失。
5.项目4的系统，其中所述Rap1GAP的G蛋白结合结构域包含Rap1GAP的1至420位或1至436位的残基。
6.项目4的系统，其中在位置437、439和441的所有三个丝氨酸残基都突变。
7.项目4或6的系统，其中所述丝氨酸残基被丙氨酸替换。
8.项目2的系统，其中所述GASIP包含RGS蛋白的G蛋白结合结构域。
9.项目8的系统，其中所述RGS蛋白是RGS17、RGS19或RGS20。
10.项目9的系统，其中所述G蛋白结合结构域包含RGS17(SEQ ID NO：17)的64至210位的残基、RGS19(SEQ ID NO：18)的70-217位的残基或RGS20(SEQ ID NO：19)的242-388位的残基。
11.项目1的系统，其中所述与Gq特异性结合的蛋白结构域是P63RhoGEF的G蛋白结合结构域或GRK2的G蛋白结合结构域。
12.项目11的系统，其中所述GASIP包含P63RhoGEF的G蛋白结合结构域。
13.项目12的系统，其中所述P63RhoGEF的G蛋白结合结构域包含P63RhoGEF(SEQID NO：25)的295至502位的残基。
14.项目11的系统，其中所述GASIP包含GRK2的G蛋白结合结构域。
15.项目14的系统，其中所述GRK2的G蛋白结合结构域包含GRK2(SEQ ID NO：27)的30至203位的残基。
16.项目1的系统，其中所述与G12/13特异性结合的蛋白结构域是PDZRhoGEF的G蛋白结合结构域或P115RhoGEF的G蛋白结合结构域。
17.项目16的系统，其中所述GASIP包含PDZRhoGEF的G蛋白结合结构域。
18.项目17的系统，其中所述PDZRhoGEF的G蛋白结合结构域包含PDZRhoGEF(SEQID NO：21)的281至483位的残基。
19.项目16的系统，其中所述GASIP包含P115RhoGEF的G蛋白结合结构域。
20.项目19的系统，其中所述P115RhoGEF的G蛋白结合结构域包含P115RhoGEF(SEQID NO：23)的1至244位的残基。
21.项目1至20中任一项的系统，其中，所述GASIP用所述生物发光供体分子标记，并且所述PM靶向部分、胞内体靶向部分或高尔基体靶向部分用所述荧光接受体分子标记。
22.项目1至21中任一项的系统，其中所述PM靶向部分是定位于PM的PM蛋白或其片段。
23.项目22的系统，其中所述PM蛋白或其片段包含(a)棕榈酰化、肉豆蔻酰化和/或异戊烯化信号序列和/或(b)多碱性序列。
24.项目22或23的系统，其中所述PM靶向部分包含氨基酸序列GCMSCKCVLS(SEQ IDNO：62)、GCMGLPCVVM(SEQ ID NO：63)、CVKIKKCIIM(SEQ ID NO：64)、KKKKKKSKTKCVIM(SEQID NO：65)、KNGKKKRKSLAKRIRERCCIL(SEQ ID NO：45)、CMSCKCCIL(SEQ ID NO：46)或SPKKGLLQRLFKRQHQNNSKS(SEQ ID NO：47)。
25.项目24的系统，其中所述PM靶向部分包含氨基酸序列GKKKKKKSKTKCVIM(SEQID NO：1)。
26.项目1至21中任一项的系统，其中所述胞内体靶向部分是定位于胞内体的胞内体蛋白或其片段。
27.项目26的系统，其中所述胞内体蛋白或其片段包含FYVE结构域。
28.项目27的系统，其中所述胞内体靶向部分包含人endofin的FYVE结构域。
29.项目28的系统，其中所述胞内体靶向部分包含人endofin(SEQ ID NO：39)的739至806位的残基。
30.项目1至21中任一项的系统，其中所述高尔基体靶向部分是定位于高尔基体的高尔基体蛋白或其片段。
31.项目30的系统，其中所述高尔基体靶向部分是定位于高尔基体的eNOS1或其片段。
32.项目31的系统，其中所述高尔基体靶向部分包含人eNOS1(SEQ ID NO：48)的1至73位的残基。
33.项目1至32中任一项的系统，其中所述第一组分还包括在(i)所述GASIP和(ii)所述生物发光供体分子或荧光接受体分子之间的接头。
34.项目1至33中任一项的系统，其中所述第二组分还包括在(i)所述PM靶向部分、胞内体靶向部分或高尔基体靶向部分与(ii)所述生物发光供体分子或荧光接受体分子之间的接头。
35.项目33或34的系统，其中所述接头是5至25个氨基酸连接的肽。
36.项目1至35中任一项的系统，还包括第三组分，第三组分是通过所述G蛋白信号转导的细胞表面受体。
37.项目36的系统，其中所述细胞表面受体是GPCR、RTK或整合素受体。
38.项目1至37中任一项的系统，其中所述G蛋白活化是G蛋白偶联受体(GPCR)介导的G蛋白活化。
39.项目1至38中任一项的系统，还包括第四组分，第四组分为重组Gα亚基多肽。
40.项目39的系统，其中所述G蛋白活化是非受体鸟嘌呤核苷酸交换因子(GEF)介导的G蛋白活化，其中所述重组Gα亚基多肽在所述Gα亚基多肽的羧基(C)末端结构域中包含至少一个突变，并且其中所述C末端结构域对应于所述Gα亚基多肽的最后七个残基。
41.项目40的系统，其中所述突变是至少最后两个C末端残基的缺失或替换。
42.项目41的系统，其中所述突变是至少最后五个C末端残基的缺失或替换。
43.项目40的系统，其中所述突变是在所述C末端结构域中至少一个保守亮氨酸残基的缺失或替换。
44.项目43的系统，其中所述突变是在所述C末端结构域中最后一个保守亮氨酸残基的替换。
45.项目44的系统，其中所述替换是亮氨酸至天冬氨酸、亮氨酸至脯氨酸或亮氨酸至精氨酸的替换。
46.项目40至45中任一项的系统，其中所述GEF为GIV/Girdin，并且其中所述GASIP包含如项目1至7中任一项所限定的Rap1GAP的G蛋白结合结构域。
47.项目46的系统，其中所述GEF被受体酪氨酸激酶(RTK)活化。
48.项目1至47中任一项的系统，其中所述生物发光供体分子是荧光素酶，优选地是海肾(Renilla)荧光素酶蛋白(rLuc)。
49.项目1至48中任一项的系统，其中所述荧光接受体分子是绿色荧光蛋白(GFP)，优选地是海肾GFP(rGFP)。
50.一种或多种核酸，编码项目1至49中任一项的系统的第一组分和/或第二组分。
51.项目50的一种或多种核酸，包括编码项目1至49中任一项的系统的第一组分的第一核酸和编码项目1至49中任一项的系统的第二组分的第二核酸。
52.一种或多种载体，包括项目50或51的一种或多种核酸。
53.宿主细胞，表达项目1至49中任一项所限定的系统的组分。
54.一种确定剂是否调节感兴趣的G蛋白活化的方法，所述方法包括：(a)使项目1至49中任一项的系统与所述生物发光供体分子的底物接触；以及(b)在所述剂存在和不存在下测量系统中的BRET信号；其中在所述剂存在下的所述BRET信号相对于其不存在下的差异表明所述剂调节所述感兴趣的G蛋白活化。
55.项目54的方法，其中所述感兴趣的G蛋白是Gi蛋白亚基家族，并且其中所述GASIP包含项目1至7中任一项所限定的Rap1GAP的G蛋白结合结构域。
56.项目49的方法，其中所述系统包括Gi蛋白亚基家族的重组Gα亚基多肽。
57.项目56的方法，其中所述方法使用多个系统执行，并且其中每个所述系统包括Gi蛋白亚基家族的不同重组Gα亚基多肽。
58.项目54的方法，其中所述感兴趣的G蛋白是Gq蛋白亚基家族，其中所述GASIP包含项目1和11至15中任一项所限定的P63RhoGEF的G蛋白结合结构域或GRK2的G蛋白结合结构域。
59.项目58的方法，其中所述系统包括Gq蛋白亚基家族的重组Gα亚基多肽。
60.项目59的方法，其中所述方法使用多个系统执行，并且其中每个所述系统包括Gq蛋白亚基家族的不同重组Gα亚基多肽。
61.项目54的方法，其中所述感兴趣的G蛋白是G12/13蛋白亚基家族，其中所述GASIP包含项目1和16至20中任一项所限定的PDZRhoGEF的G蛋白结合结构域或P115RhoGEF的G蛋白结合结构域。
62.项目61的方法，其中所述系统包括G12/13蛋白亚基家族的重组Gα亚基多肽。
63.项目62的方法，其中所述方法使用多个系统执行，并且其中每个所述系统包括G12/13蛋白亚基家族的不同重组Gα亚基多肽。
64.一种确定剂是否调节非受体鸟嘌呤核苷酸交换因子(GEF)介导的G蛋白活化的方法，所述方法包括(a)使项目40至47中任一项的系统与所述生物发光供体分子的底物接触；以及(b)在所述剂存在和不存在下测量系统中的BRET信号；其中在所述剂存在下的所述BRET信号相对于其不存在下的差异表明所述剂调节非受体GEF介导的G蛋白活化。
65.项目54至64中的任一项的方法，其中使用读板仪或通过显微术来测量BRET信号。
66.项目54至65中任一项的方法，其中底物是腔肠素底物。
67.项目66的方法，其中所述腔肠素底物是甲氧基e腔肠素。
通过参考附图阅读以下仅以举例方式给出的本发明的特定实施方式的非限制性描述，本发明的其他目的、优点和特征将变得更加明显。
附图说明：
在附图中：
图1A描绘了基于效应子的传感器的原理，其监测GPCR介导的直接G蛋白活化(上小图i)或鸟嘌呤-核苷酸交换因子(GEF)介导的G蛋白活化(下小图ii)。使表达受体，用合适的生物发光共振能量转移(BRET)供体或接受体(例如，rGFP)标记的细胞区室(或亚细胞定位)标志物(诸如质膜(PM)或早期胞内体(EE)标志物)，用合适的BRET供体或接受体(例如，RlucII)标记的特定G蛋白效应子的Gα相互作用结构域的细胞暴露于激动剂以活化共表达的G蛋白。在小图i)中，激动剂诱导的GPCR刺激直接活化了G蛋白，其将标记的效应子从细胞质募集到标记的膜。在小图ii)中，在RTK(例如，EGFR)或整合素α-β复合物活化后，GEF诸如GIV/Girdin的募集介导了G蛋白活化。
图1B-1E示出了用三种不同的G蛋白效应子获得的特异性的实例。与Gq、Gi和G12/13家族成员偶联的受体与这些G蛋白共表达，其中rGFP与PM标志物融合(rGFP-CAAX，GKKKKKKSKTKCVIM，SEQ ID NO：1)，以及G蛋白效应子用RlucII标记。图1B示出使用Rap1GAP(SSS-AAA)-RlucII和rGFP-CAAX(PM标志物)，用血小板活化因子(PAF)/PAFR介导的G蛋白(Gq、G11、G14、G12、GoA、GoB、Gz、Gi1、Gi2、Gi3)活化获得的剂量应答曲线(DRC)。模拟应答代表内源表达的Gi1、Gi2和Gi3蛋白的活化。图1C示出了使用PDZRG-RlucII和rGFP-CAAX，用U46619/TPαR介导的G蛋白(Gq、G11、G14、G15、G12、G13、Gz)活化获得的DRC。模拟应答代表内源表达的G12和G13蛋白的活化。图1D和1E示出了使用P63RG-RlucII和rGFP-CAAX，用PAF/PAFR介导的G蛋白(Gq、G11、G14、G12、GoA、GoB、Gz、Gi1、Gi2、Gi3)活化(图1D)以及U46619/TPαR介导的G蛋白(Gq、G11、G14、G15、G12、G13、Gz)活化(图1E)获得的DRC。模拟应答代表内源表达的Gq和G11蛋白的活化。
图2A描绘了基于Rap1GAP的系统的测试的各种构建体，用于监测Gi家族G蛋白的活化。第一个构建体由在C末端标记有BRET供体Rluc8的Rap1GAP的部分(1-442位残基)组成。使用不同的接头并使用RlucII而不是Rluc8，从该构建体衍生出第二构建体。实验的结果在图2B-2Z示出。接头A＝GSGGGSGGGA(SEQ ID NO：6)，而接头B＝GSAGTGGRAIDIKLPAT(SEQ IDNO：7)。Rap1GAP 1-442＝SEQ ID NO：8；Rap1GAPΔcterm(1-420)＝SEQ ID NO：10；Rap1GAPΔSSS＝SEQ ID NO：9；Rap1GAP SS-AA＝SEQ ID NO：13；Rap1GAP SS-DA＝SEQ ID NO：14；Rap1GAP SS-AD＝SEQ ID NO：15；Rap1GAP SS-DD＝SEQ ID NO：16；Rap1GAP SSS-AAA＝SEQID NO：11；Rap1GAP SSS-TTT＝SEQ ID NO：12。
图2B-2E示出了使用三种不同的Rap1GAP-Rluc构建体：Rap1GAP(1-442)-Rluc8(圆形)、Rap1GAP(1-442)-RlucII(三角形)以及Rap1GAP(ΔCT)-RlucII(菱形)，进行AR-M100390(ARM)刺激后，使用人Mu阿片受体(hMOR1)介导的Gi1活化(图2B)、Gi2活化(图2C)、GoA活化(图2D)以及Gz活化(图2E)的DRC。
图2F-2H示出了使用Rap1GAP(1-442)-Rluc8(图2F)、Rap1GAP(1-442)-RlucII(图2G)或RAP1GAP(ΔCT)-RlucII(图2H)的，在毛喉素(三角形)或媒介物(DMSO/酪氨酸，圆形)存在下使用hMOR1/ARM促进的在PM处的Gz活化的DRC，其促进了cAMP产生的增加以及导致不同蛋白磷酸化的蛋白激酶A的活化。
图2I-2N示出了使用Rap1GAP(1-442)-RlucII(图2I)、截短(1-436位残基)形式的Rap1GAP(ΔSSS)-RlucII(图2J)、Rap1GAP(SS-AD)-RlucII(图2K)、Rap1GAP(SS-DA)-RlucII(图2L)、Rap1GAP(SS-AA)-RlucII(图2M)或Rap1GAP(SS-DD)-RlucII(图2N)，在毛喉素(方形)或媒介物(圆形)存在的情况下使用hMOR1/ARM促进的在PM处的Gi2活化的DRC。
图2O-2Q示出了使用Rap1GAP(1-442)-RlucII(图2O)、Rap1GAP(SSS-TTT)-RlucII(图2P)和Rap1GAP(SSS-AAA)-RlucII(图2Q)，在毛喉素(三角形)或媒介物(圆形)存在的情况下使用hMOR1/ARM促进的在PM处的GoB活化的DRC。
图2R示出了在共表达Gi1、Gi2、Gi3、GoA或GoB的情况下，使用Rap1GAP(SSS-AAA)-RlucII易位至质膜，多巴胺促进的多巴胺D4受体(D4R)刺激后G蛋白活化的DRC。
图2S-2W示出了用配体A412,997、多巴胺、L741,742和Way-100635刺激多巴胺受体D4R(图2S)、D1R(图2T)、D2R(图2U)、D3R(图2U)或D1R(图2W)之后Gz活化的DRC。
图2X-2Z是描绘测定的Z'因子的图，其指示测定的鲁棒性。用D4R、Rap1GAP(SSS-AAA)-RlucII、rGFP-CAAX与WT Gi2(图2X)、与WT GoA(图2Y)或WT Gz(图2Z)一起共转染HEK293，在96孔板中铺板，在37℃下用100nM多巴胺(三角形)或媒介物(DMSO/Tyrode；方形)刺激6-8min。以BRET2评估Rap1GAP(SSS-AAA)-RlucII至PM的募集。在所示图中，BRET值以每孔表示。
图3A描绘了基于本文所述研究中使用的G蛋白信号转导调节物(RGS)蛋白的成员的RGS结构域的用于监测Gi家族的G蛋白的活化的其他构建体。C末端用BRET供体RlucII标记包括RGS17的RGS(Gi结合)结构域(SEQ ID NO：17的64-210位残基)、RGS19的RGS(Gi结合)结构域(SEQ ID NO：18的70-217位残基)和RGS20的RGS(Gi结合)结构域(SEQ ID NO：19的242-388位残基)的片段。接头B'＝GSAGTGGRAIDIKLASAT(SEQ ID NO：20)。
图3B-3D示出了GoA、GoB、Gz、Gi1、Gi2或Gi3共表达的情况下，使用RGS(RGS17)-RlucII(图3B)、RGS(RGS19)-RlucII(图3C)或RGS(RGS20)-RlucII(图3D)易位到PM，在多巴胺促进的D4)刺激后G蛋白活化的DRC。
图4A描绘了用于监测本文所述研究中使用的G12/13家族的G蛋白活化的PDZRhoGEF(PDZRG)-RlucII构建体。用BRET供体RlucII在C末端标记包括PDZRhoGEF的G12/13结合结构域(SEQ ID NO：21的281-483位残基)的片段。接头D＝GIRLREALKLPAT(SEQ IDNO：22)。
图4B和4C示出在用TPαR激动剂I-BOP(CAS编号：128719-90-4)刺激后，用血栓烷受体(TPαR)、PDZRG-RlucII、rGFP-CAAX以及无Gα(模拟)、5ng的Gα、20ng的Gα或100ng的Gα共转染的HEK293细胞中G12(图4B)和G13(图4C)活化的DRC。
图4D和4E示出了使用PDZRG-RlucII，TPαR配体对在PM处G12(图4D)和G13(图4E)活化的DRC。用已知的完全激动剂(U46619、I-BOP、CTA2)，用部分激动剂(U51605)以及拮抗剂I-SAP和SQ 29,558刺激用血栓烷受体(TPαR)、PDZRG-RlucII、rGFP-CAAX共转染的HEK293细胞。将结果归一化并表示为I-BOP应答(n＝4)的百分比(％)+/-SEM。
图4F和4G是描绘使用PDZRG-RlucII构建体的测定的Z'因子的图。用TPαR、PDZRG-RlucII、rGFP-CAAX和用WT G12(图4F)或WT G13(图4G)共转染HEK293，铺在96孔板中，并在37℃下用100nM的TPαR激动剂U46619(三角形)或媒介物(乙酸甲酯/Tyrode；方形)刺激6-8min。以BRET2评价了PDZRG-RlucII至PM的募集。在所示图中，BRET值以每孔表示。
图5A描述了本文所述研究中使用的用于监测G12/13家族的G蛋白活化的P115RhoGEF(P115RG)-RlucII构建体。用BRET供体RlucII在C末端标记包括P115RhoGEF的G12/13结合结构域(SEQ ID NO：23的1-244位残基)的片段。接头C＝RLKLPAT(SEQ ID NO：24)。
图5B和图5C示出了使用P115RG-RlucII，TPαR配体对在PM处的G12(图5B)和G13(图5C)活化的DRC。用U46619、I-BOP、CTA2、U51605、I-SAP和SQ 29,558刺激用血栓烷受体(TPαR)、P115RG-RlucII、rGFP-CAAX共转染的HEK293细胞。将结果归一化并表示为I-BOP应答(n＝4)的百分比(％)+/-SEM。
图5D和5E是描绘使用PDZRG-RlucII构建体的测定的Z'因子的图。用TPαR、P115RG-RlucII、rGFP-CAAX和WT G12(图5D)或WT G13(图5E)共转染HEK293，铺在96孔板中，并在37℃下用100nM的TPαR激动剂U46619(三角形)或媒介物(乙酸甲酯/Tyrode；方形)刺激6-8min。以BRET2评价了P115RG-RlucII至PM的募集。在所示图中，BRET值以每孔表示。
图6A描述了本文所述研究中使用的用于监测Gq家族(Gq、G11、G14&G15)的G蛋白活化的P63RhoGEF(P63RG)构建体。用BRET供体RlucII在C末端标记包括P63RhoGEF的Gq结合结构域(SEQ ID NO：25的295-502位残基)的片段。接头E＝ASGSAGTGGRAIDIKLPAT(SEQ ID NO：26)。
图6B-6E示出了用TPαR激动剂U46619刺激后，用TPαR、P63RG-RlucII、rGFP-CAAX以及无Gα(模拟，来自内源G蛋白获得的应答)或不同量的Gα亚基共转染的HEK293细胞中在PM处的Gq(图6B)、G11(图6C)、G14(图6D)和G15(图6E)活化的DRC。
图6F-6I示出了用TPαR激动剂U46619刺激后，用TPαR、P63RG-RlucII、rGFP-FYVE以及无Gα(模拟，来自内源G蛋白获得的应答)或不同量的Gα亚基共转染的HEK293细胞中在早期胞内体处的Gq(图6F)、G11(图6G)、G14(图6H)和G15(图6I)活化的DRC。
图6J-6M示出了在图6B-6E确定的优化条件下使用P63RG-RlucII，TPαR配体对在PM处Gq(图6J)、G11(图6K)、G14(图6L)和G15(图6M)活化的DRC。用U46619、I-BOP、CTA2、U51605、I-SAP和SQ 29,558刺激用血栓烷受体(TPαR)、P63RG-RlucII、rGFP-CAAX共转染的HEK293细胞。将结果归一化并表示为I-BOP应答(n＝3和n＝5之间)的百分比(％)+/-SEM。
图6N-6Q示出了在图6F-6I中确定的优化条件下使用P63RG-RlucII，TPαR配体对在EE处Gq(图6N)、G11(图6O)、G14(图6P)和G15(图6Q)活化的DRC。用U46619、I-BOP、CTA2、U51605、I-SAP和SQ 29,558刺激用血栓烷受体(TPαR)、P63RG-RlucII、rGFP-FYVE共转染的HEK293细胞。将结果归一化并表示为I-BOP应答(n＝3和n＝5之间)的百分比(％)+/-SEM。
图6R和6S是描绘使用P63RG-RlucII构建体的测定的Z'因子的图。用TPαR、P63RG-RlucII、rGFP-CAAX和用WT Gq(图6R)或WT G11(图6S)共转染HEK293，铺在96孔板中，并在37℃下用100nM的TPαR激动剂U46619(三角形)或媒介物(乙酸甲酯/Tyrode；方形)刺激6-8min。以BRET2评价了P63RG-RlucII至PM的募集。在所示图中，BRET值以每孔表示。
图7A描述了本文所述研究中使用的用于监测Gq家族(Gq、G11、G14&G15)的G蛋白活化的两个RGS(GRK2)构建体。用BRET供体RlucII在N末端RlucII-RGS(GRK2)或C末端(RGS(GRK2)-RlucII)标记包括GRK2的Gq结合结构域(RGS结构域)(SEQ ID NO：27的30-203位残基)的片段。接头B'＝GSAGTGGRAIDIKLASAT(SEQ ID NO：20)。
图7B和7C示出了使用RlucII-RGS(GRK2)，在两个配体/受体系统，即At1AR/ANGII(图7B)和TPαR/U46619(图7C)中在PM处的Gq、G11、G14和G15活化的DRC。
图7D和7E示出了使用RGS(GRK2)-RlucII，在两个配体/受体系统，即At1AR/ANGII(图7D)和TPαR/U46619(图7E)中在PM处的Gq、G11、G14和G15活化的DRC。
图7F是描绘使用RlucII-RGS(GRK2)构建体的测定的Z'因子的图。用TPαR、RlucII-RGS(GRK2)、rGFP-CAAX和用WT Gq共转染HEK293，铺在96孔板中，并在37℃下用100nM的TPαR激动剂U46619(三角形)或媒介物(乙酸甲酯/Tyrode；方形)刺激6-8min。以BRET2评价了RlucII-RGS(GRK2)至PM的募集。在所示图中，BRET值以每孔表示。
图8A描绘了在本文所述的研究中测试的突变的Gi2和GoB蛋白监测GEF介导的G蛋白活化的能力，即，非GPCR介导的G蛋白活化的能力。Gαi2Δ5＝SEQ ID NO：28；Gαi2Δ2＝SEQ ID NO：29；Gαi2 L-2G＝SEQ ID NO：31；Gαi2 L-2P＝SEQ ID NO：33；Gαi2 L-2R＝SEQID NO：34；Gαi2L-2D＝SEQ ID NO：32；Gαi2 L-7G＝SEQ ID NO：30；GαoBΔ5＝SEQ ID NO：36；GαoB L-2G＝SEQ ID NO：35。
图8B-8G示出了Gi2和GoB的缺失突变体(即Gi2Δ2(图8B和8C)、Gi2Δ5(图8D和8E)和GoBΔ5(图8F和8G))的活化的DRC。用编码EGF受体(EGFR，图8B、8D、8F)或缓激肽受体(BKB2R，图8C、8E、8G)的构建体Rap1GAP(SSS-AAA)-RlucII、rGFP-CAAX和WT或突变的Gi2/GoB亚基共转染HEK293。
图8H-8M示出了Gi2和GoB的Leu至Gly突变体(即Gi2 L-7G(图8H和8I)、Gi2 L-2G(图8J和8K)和GoB L-2G(图8L和8M))的活化的DRC。用编码EGFR(图8H、8J、8L)或BKB2R(图8I、8K、8M)的构建体Rap1GAP(SSS-AAA)-RlucII、rGFP-CAAX和WT或突变的Gi2/GoB亚基共转染HEK293。
图8N-8S示出了Gi2的位置-2突变体(即Gi2 L-2D(图8N和8O)、Gi2 L-2P(图8P和8Q)和Gi2 L-2R(图8R和8S))的活化的DRC。用编码EGFR(图8N、8P、8R)或BKB2R(图8O、8Q、8S)的构建体Rap1GAP(SSS-AAA)-RlucII、rGFP-CAAX和WT或突变的Gi2亚基共转染HEK293。
图8T-8U是描述使用Rap1GAP(SSS-AAA)-RlucII构建体监测GEF介导的G蛋白活化的测定的Z'因子的图。用EGFR、Rap1GAP(SSS-AAA)-RlucII、rGFP-CAAX与WT Gi2(图8T)或突变体Gi2L-2P(图8U)共转染HEK293，铺在96孔板中，在室温下用10ng/ml的EGF(三角形)或媒介物(Tyrode；方形)刺激2min。以BRET2评价Rap1GAP(SSS-AAA)-RlucII至PM的募集。在所示图中，BRET值以每孔表示。
图8V-8X示出了使用Rap1GAP(SSS-AAA)-RlucII构建体(圆形)和RGS(RGS17)-RlucII(三角形)监测的，由两种GPCR(在图8V中的δ-阿片样物质受体(DOR)和在图8W中的D2R)以及RTK(EGFR；图8X)活化Gi2的DRC。用编码受体(DOR、D2R或EGFR)的构建体Rap1GAP(SSS-AAA)-RlucII或RGS(RGS17)-RlucII、rGFP-CAAX和WT Gi2共转染HEK293，铺在96孔板中并在室温下以指定剂量用GPCR激动剂刺激8min(图8V、8W)或用EGF刺激7min(图8X)。以BRET2评价Rap1GAP(SSS-AAA)-RlucII和RGS(RGS17)-RlucII至PM的募集。提出的DRC代表了三个独立的实验。
图9A-9D示出了本文所述研究中使用的多肽的氨基酸序列。
具体实施方式
本文所用的遗传学、分子生物学、生物化学和核酸的术语和符号遵循该领域标准论文和教科书的术语和符号，例如Kornberg和Baker，《DNA复制》，第二版(W大学科学丛书，2005年)；Lehninger，生物化学，第6版(WH Freeman&Co(Sd)，纽约，2012年)；Strachan和Read，《人类分子遗传学》，第二版(Wiley-Liss，纽约，1999年)；Eckstein，编辑，《寡核苷酸和类似物：一种实用方法》(牛津大学出版社，纽约，1991年)；Gait，编辑，《寡核苷酸合成：一种实用方法》(IRL出版社，牛津，1984)年；等等。所有术语应以相关领域中确立的典型含义来理解。
冠词“一”和“一个”在本文中用于指代冠词的语法对象中的一个或多个(即指代至少一个)。举例来说，“一个元件”是指一个元件或多于一个元件。在整个说明书中，除非上下文另外要求，否则词语“包括(comprise)”，“包括(comprises)”和“包括(comprising)”将被理解为暗示包括陈述的步骤或元素或步骤或元素的组，但不排除任何其他步骤或元素或者步骤或元素的组。
本说明书中涉及的与Genbank、RefSeq、UniProt、NCBI和/或Ensembl登录号(或任何其他数据库)相对应的信息，包括核苷酸和氨基酸序列，通过引用并入本文。
除非本文另外指出或与上下文明显矛盾，否则本文描述的所有方法可以以任何合适的顺序执行。
除非另外要求，否则本文提供的任何和所有实例或示例性语言(“例如”，“诸如”)的使用仅旨在更好地说明本发明，并且不对本发明的范围构成限制。
在本文中，术语“约”具有其普通含义。术语“约”用于表示一个值包括用于确定该值的设备或方法的误差的固有变化，或涵盖接近所述值的值，例如在所述值(或值的范围)的10％或5％之内。
本发明涵盖本文公开的实施方式和特征的任何和所有组合和子组合。例如，本文确定的2个、3个、4个、5个或更多个基因的任何组合的表达可用于本文所述的方法。
本公开涉及允许以G蛋白家族选择性的方式并且在不同的细胞区室中监测G蛋白活化的系统和测定法。这些系统和测定法基于标记有合适的能量(即BRET)供体和接受体的G蛋白的特异性效应子和细胞区室标志物的使用。这些系统和测定法有利地不需要任何G蛋白亚基(Gα、Gβ或Gγ)的修饰(例如，用BRET供体或接受体或其他可检测的标记来标记/融合)，因为通过评估特异性G蛋白效应子的易位来以G蛋白家族选择性的方式在特定细胞区室处直接检测G蛋白活化，从而最小化G蛋白的功能/活性被改变的风险。用本文描述的系统获得的动态窗口高于用其他系统评估G蛋白活化获得的动态窗口(例如，美国专利号9,029,097和WO/2016/058094)，这对于高通量筛选(HTS)应用(包括鉴定部分激动剂)可能很重要，并且还允许检测内源性G蛋白介导的信号(在G蛋白家族水平)。通过使用对感兴趣的G蛋白亚基(一种或多种)具有特异性的G蛋白效应子的G蛋白结合结构域，诸如针对Gi家族(Gi1、Gi2、Gi3、GoA、GoB、Gz)的Rap1GAP的G蛋白结合域、针对Gq家族(Gq、G11、G14&G15)的P63RhoGEF(P63RG)的G蛋白结合结构域以及针对G12/13家族的PDZRhoGEF或P115RhoGEF的G蛋白结合结构域。当需要时，可以通过感兴趣的G蛋白亚基(一种或多种)与研究的受体共表达来实现G蛋白亚基水平上的特异性。
因此，一方面，本公开提供了一种以Gα蛋白亚基家族选择性方式测量G蛋白活化调节的系统，所述系统包括：
表达以下的细胞：
(i)第一组分，包括用生物发光供体分子或荧光接受体分子标记的Gα亚基相互作用多肽(GASIP)；
其中：
如果所述Gα蛋白亚基家族是Gi，则所述GASIP包含与Gi结合的蛋白结构域；
如果所述Gα蛋白亚基家族是Gq，则所述GASIP包含与Gq结合的蛋白结构域；
如果所述Gα蛋白亚基家族是G12/13，则所述GASIP包含与G12/13结合的蛋白结构域；和
如果所述Gα蛋白亚基家族是Gs，则所述GASIP包含与Gs结合的蛋白结构域；以及
(ii)第二组分，包括用生物发光供体分子或荧光接受体分子标记的质膜(PM)靶向部分、胞内体靶向部分或高尔基体靶向部分；
其中，如果所述GASIP用所述荧光接受体分子标记，则所述PM靶向部分、胞内体靶向部分或高尔基体靶向部分用所述生物发光供体分子标记，以及如果所述GASIP用所述生物发光供体分子标记，则所述PM靶向部分、胞内体靶向部分或高尔基体靶向部分用所述荧光接受体分子标记。
另一方面，本公开提供了一种以Gα蛋白亚基家族选择性方式测量G蛋白活化调节的系统，所述系统包括：
表达以下的细胞：
(i)第一组分，包括用生物发光供体分子或荧光接受体分子标记的Gα亚基相互作用多肽(GASIP)；
其中：
如果所述Gα蛋白亚基家族是Gi，则所述GASIP包含以下的G蛋白结合结构域：Rap1GAP、G蛋白信号转导调节物(RGS)蛋白或TNFAIP8，优选为Rap1GAP或RGS蛋白的G蛋白结合结构域；
如果所述Gα蛋白亚基家族是Gq，则所述GASIP包含以下的G蛋白结合结构域：RhoGEF蛋白(例如，P63RhoGEF或TRIO)、GRK2、GRK3、TPR1(三十四肽重复结构域1(tetratricopeptide repeat domain 1))或RGS蛋白(例如，RGS2)，优选为P63RhoGEF或GRK2/GRK3的G蛋白结合结构域；
如果所述Gα蛋白亚基家族为G12/13，则所述GASIP包含RhoGEF蛋白(PDZRhoGEF，P115RhoGEF或LARG)、JNK相互作用亮氨酸拉链蛋白(JLP)、钙粘蛋白、Axin1、PP2A、SNAP-α、多囊蛋白(polycistin)1、RGS16、AKAP110或HAX1(HS1相关的蛋白X1)的G蛋白结合结构域，优选为PDZRhoGEF或P115RhoGEF的G蛋白结合结构域；
如果所述Gα蛋白亚基家族是Gs(Gαs、XLGαs、Gαolf)，则所述GASIP包含SNX13(RGS-PX1)、AXIN1或TPR1(三十四肽重复结构域1)的G蛋白结合结构域；
(ii)第二组分，包括用生物发光供体分子或荧光接受体分子标记的质膜(PM)靶向部分、胞内体靶向部分或高尔基体靶向部分；
其中，如果所述GASIP用所述荧光接受体分子标记，则所述PM靶向部分、胞内体靶向部分或高尔基体靶向部分用所述生物发光供体分子标记，以及如果所述GASIP用所述生物发光供体分子标记，则所述细胞M靶向部分、胞内体靶向部分或高尔基体靶向部分用所述荧光接受体分子标记。
选择生物发光供体分子(或BRET供体)和荧光接受体分子(或BRET接受体)，使得生物发光供体分子的发射谱与荧光接受体分子的吸收谱重叠。在这样的条件下，由生物发光供体分子传递的光能处于能够激发荧光接受体分子的波长，即生物发光共振能量转移(BRET)。共振能量转移(缩写为RET)是描述具有重叠发射/吸收谱的两个发色团之间能量转移的机制。当两个发色团(“供体”和“接受体”)彼此之间的距离很短(例如，10-100埃)并且它们的跃迁偶极取向正确时，供体发色团能够将其激发态能量通过非辐射偶极-偶极耦合传递给接受体发色团。生物发光共振能量转移(BRET)是基于供体生物发光分子(生物发光酶，诸如海肾荧光素酶)和接受体荧光分子(例如，海肾GFP)之间能量的非辐射转移。
如本文所用，术语生物发光供体分子是指在对合适的底物起作用或由外部源自身激发后能够产生发光的任何分子。在本公开中可以使用许多不同的生物发光供体分子。发光系统是已知的，并从许多发光的生物体分离，包括细菌、原生动物、腔肠动物、贝类、鱼、多足类、蝇类、真菌、蠕虫、甲壳类和甲虫，特别是光叩甲属(Pyrophorus)的叩头虫和Photinus属、Photuris属和熠萤属(Luciola)的萤火虫。表现出生物发光的其他生物体在PCT公开号WO 00/024878和WO 99/049019中列出。在一种实施方式中，生物发光供体分子是萤光素酶。具有荧光素酶活性的生物发光蛋白的实例公开于美国专利号5,229,285、5,219,737、5,843,746、5,196,524和5,670,356。两种最广泛使用的荧光素酶是：(i)海肾荧光素酶和(ii)萤火虫荧光素酶。
在一种实施方式中，生物发光供体分子是海肾荧光素酶(rLuc)。如本文所用的术语海肾(Renilla)荧光素酶是指用于生物发光以及来源自海肾属的生物体(诸如Renillareniformis或Renilla mulleri)的氧化酶。它包括来自海肾属生物体的天然萤光素酶或其变体，例如Renilla reniformis萤光素酶(Rluc)的天然形式(就氨基酸序列而言)或其变体，诸如RlucII、Rluc3、绿色海肾萤光素酶或Rluc8。术语“RlucII”是指Renillareniformis荧光素酶的突变形式，其相对于天然海肾荧光素酶包含以下氨基酸替换：A55T、C124A和M185V。在一种实施方式中，RlucII包含图9C所示的序列。术语“Rluc8”是指Renillareniformis荧光素酶的突变形式，其相对于天然Renilla reniformis荧光素酶包含以下氨基酸替换：A55T、C124A、S130A、K136R、A143M、M185V、M253L和S287L。天然Renilla mulleri萤光素酶的氨基酸序列以GenBank登录号AAG54094.1公开。
这些酶的天然和合成发光底物在本领域中是众所周知的，并且是可商购的。萤光素酶底物的实例包括萤光素(例如，D萤光素及其盐、Latia萤光素、细菌萤光素、双鞭毛虫萤光素等)、腔肠素、腔肠素h、腔肠素e、腔肠素f、腔肠素fcp、腔肠素cp、腔肠素hcp、腔肠素i、腔肠素ip、腔肠素n、腔肠素400a(DeepBlueCTM)、甲氧基e腔肠素(来自NanoLight
的
Purple I)、甲氧基-腔肠素-甲氧基(来自NanoLight
的
Purple II)、甲氧基-腔肠素F(来自NanoLight
的
Purple III)、甲氧基-腔肠素-碘(来自
的
Purple IV)、甲氧基-v-腔肠素-甲氧基(来自
的
Purple V)、ViviRenTM(来自
)。生物发光供体分子的合适底物可以由技术人员基于所需的生物发光蛋白发射的光的波长和/或强度来选择。在一种实施方式中，荧光素酶底物是腔肠素h、甲氧基e腔肠素或腔肠素400A。
如本文所用，术语荧光接受体分子是指可以接受由于生物发光供体分子的活性而发射的能量并将其重新发射为光能的任何化合物。代表性的荧光接受体蛋白可以包括但不限于，绿色荧光蛋白(GFP)、绿色荧光蛋白的变体(诸如GFP10)、蓝色荧光蛋白(BFP)、青色荧光蛋白(CFP)、黄色荧光蛋白(YFP)、增强的GFP(EGFP)、增强的CFP(ECFP)、增强的YFP(EYFP)、GFPS65T、mAmetrine、LSS-mOrange、LSS-mKate、Emerald、Topaz、GFPuv、不稳定EGFP(dEGFP)、不稳定ECFP(dECFP)、不稳定EYFP(dEYFP)、HcRed、t-HcRed、DsRed、DsRed2、mRFPI、pocilloporin、海肾GFP(rGFP)、Monster GFP、paGFP、Kaede蛋白或藻胆蛋白或其任何一种的生物活性变体或片段。最常用的生物发光或荧光团是来自水母维多利亚多管水母(Aequorea victoria)的GFP和许多由例如诱变和嵌合蛋白技术获得的其他变体(GFP)。GFP根据其发色团的独特成分进行分类，每类具有不同的激发和发射波长：1类，中性酚和阴离子酚盐的野生型混合物：2类，酚盐阴离子：3类，中性酚：4类，带有堆叠s电子体系的酚盐阴离子：5类，吲哚：6类，咪唑：以及7类，苯基。
非蛋白质荧光接受体分子的实例是AlexaTM(分子探针)、荧光染料、Bodipy染料TM(生命技术)、Cy染料TM(生命技术)、荧光素、丹磺酰、伞形酮(7-羟基香豆素)、荧光微球、发光纳米晶体、Marina blueTM(生命技术)、Cascade blueTM(生命技术)、Cascade yellowTM(生命技术)、Pacific BlueTM(生命技术)、Oregon greenTM(生命技术)、四甲基罗丹明、罗丹明、Texas redTM(生命技术)、稀土元素螯合物或其任何组合或衍生物。
其他代表性荧光接受体分子可以包括但不限于sgGFP、sgBFP、BFP蓝移GFP(Y66H)、青色GFP、DsRed、单体RFP、EBFP、ECFP、GFP(S65T)、GFP红移(rsGFP)、非UV激发(wtGFP)、UV激发(wtGFP)、GFPuv、HcRed、rsGFP、蓝宝石GFP、sgBFPTM、sgBFPTM(超辉光BFP)、sgGFPTM、sgGFPTM(超辉光GFP)、Yellow GFP、半导体纳米颗粒(例如，拉曼纳米颗粒)、1,5IAEDANS；1,8-ANS；4甲基伞形酮、5-羧基-2,7-二氯荧光素；5-羧基荧光素(5-FAM)；5-羧基萘荧光素(5-Carboxynapthofluorescein)；5-羧基四甲基罗丹明(5-TAMRA)；5-FAM(5-羧基荧光素)；5-HAT(羟色胺)；5-羟色胺(HAT)；5-ROX(羧基-X-罗丹明)；5-TAMRA(5-羧基四甲基罗丹明)；6-羧基罗丹明6G；6-CR 6G；6-JOE；7-氨基-4-甲基香豆素；7-氨基放线菌素D(7-AAD)；7-羟基-4-甲基香豆素；9-氨基-6-氯-2-甲氧基吖啶；ABQ；酸性品红；ACMA(9-氨基-6-氯-2-甲氧基吖啶)；吖啶橙；吖啶红；吖啶黄；吖啶黄素(Acriflavin)；吖啶黄素Feulgen SITS A；水母素(光蛋白)；AFPs(自发荧光蛋白；量子生物技术)；Alexa Fluor 350TM；Alexa Fluor 430TM；Alexa Fluor 488TM；Alexa Fluor 532TM；Alexa Fluor 546TM；Alexa Fluor 568TM；AlexaFluor 594TM；Alexa Fluor 633TM；Alexa Fluor 647TM；Alexa Fluor 660TM；Alexa Fluor680TM；茜素络合酮(Alizarin Complexon)；茜素红；别藻蓝蛋白(APC)；AMC、AMCA-S；AMCA(氨基甲基香豆素)；AMCA-X；氨基放线菌素D；氨基香豆素；氨基甲基香豆素(AMCA)；苯胺蓝(Anilin Blue)；蒽酰基硬脂酸酯(Anthrocyl stearate)；APC(别藻蓝蛋白)；APC-Cy7；APTRA-BTC；APTS；Astrazon艳红4G；Astrazon橙R；Astrazon红6B；Astrazon黄7GLL；阿的平(Atabrine)；ATTO-TAGTMCBQCA；ATTO-TAGTMFQ；金胺；金膦(Aurophosphine)G；金膦；BAO 9(双氨基苯基噁二唑)；BCECF(高pH)；BCECF(低pH)；硫酸小檗碱；β内酰胺酶；双满(Bimane)；双苯甲酰胺(Bisbenzamide)；双苯甲酰亚胺(Bisbenzimide)(Hoechst)；双BTC；布兰科福尔(Blancophor)FFG；布兰科福尔SV；BOBOTM-l；BOBOTM-3；氟硼荧(Bodipy)492/515；氟硼荧493/503；氟硼荧500/510；氟硼荧505/515；氟硼荧530/550；氟硼荧542/563；氟硼荧558/568；氟硼荧564/570；氟硼荧576/589；氟硼荧581/591；氟硼荧630/650-X；氟硼荧650/665-X；氟硼荧665/676；氟硼荧Fl；氟硼荧FL ATP；氟硼荧FI-神经酰胺；氟硼荧R6G SE；氟硼荧TMR；氟硼荧TMR-X缀合物；氟硼荧TMR-X、SE；氟硼荧TR；氟硼荧TR ATP；氟硼荧TR-X SE；B0-PR0TM-1；BO-PROTM-3；亮磺基黄素FF(Brilliant Sulphoflavin FF)；BTC；BTC-5N；钙黄绿素；钙黄绿素蓝；Calcium CrimsonTM；钙绿；钙绿1Ca2+染料；钙绿2Ca2+；钙绿5N Ca2+；钙绿C18Ca2+；钙橙；Calcofluor White；羧基-X-罗丹明(5-ROX)；Cascade BlueTM；级联黄；儿茶酚胺；CCF2(GeneBlazer)；CFDA；叶绿素；色霉素A；色霉素A；CL-NERF；CMFDA；香豆素鬼笔环肽；C藻蓝蛋白；CPM甲基香豆素；CTC；CTC甲瓒；Cy2TM；Cy3.18；Cy3.5TM；Cy3TM；Cy5.18；Cy5.5TM；Cy5TM；Cy7TM；环AMP荧光传感器(FiCRhR)；Dabcyl；丹磺酰(Dansyl)；丹磺酰胺(DansylAmine)；丹磺酰尸胺(Dansyl Cadaverine)；丹磺酰氯(Dansyl Chloride)；丹磺酰DHPE；丹磺酰氟；DAPI；Dapoxyl；Dapoxyl 2；Dapoxyl 3'DCFDA；DCFH(二乙酸二氯二氢荧光素)；DDAO；DHR(二氢罗丹明123)；Di-4-ANEPPS；Di-8-ANEPPS(非比例)；DiA(4-Di-16-ASP)；二乙酸二氯二氢荧光素(DCFH)；DiD-亲脂示踪剂；DiD(DilC18(5))；DIDS；二氢罗丹明123(DHR)；Dil(DilC18(3))；二硝基酚；DiO(DiOC18(3))；DiR；DiR(DilC18(7))；DM-NERF(高pH)；DNP；多巴胺；DTAF；DY-630-NHS；DY-635-NHS；ELF 97；曙红；赤藓红；赤藓红ITC；溴化乙锭；乙锭均二聚物-1(EthD-1)；Euchrysin；EukoLight；氯化铕(III)；EYFP；快蓝；FDA；富尔根(Feulgen)(副玫瑰红)；FIF(甲醛诱导荧光)；FITC；Flazo Orange；Fluo-3；Fluo-4；荧光素(FITC)；二乙酸荧光素；Fluoro-Emerald；Fluoro-Gold(羟芪巴脒)；Fluor-Ruby；FluorX；FM1-43TM；FM 4-46；Fura RedTM(高pH)；Fura RedTM/Fluo-3；Fura-2；Fura-2/BCECF；Genacryl亮红B；Genacryl亮黄10GF；Genacryl粉3G；Genacryl黄5GF；GeneBlazer(CCF2)；吉草酸(Gloxalic Acid)；粒蓝(Granular blue)；血卟啉；赫斯特(Hoechst)33258；赫斯特33342；赫斯特34580；HPTS；羟基香豆素；羟脒芪(FluoroGold)；羟色胺；Indo-1、高钙；Indo-1、低钙；吲哚二碳花菁(DiD)；吲哚三碳花菁(DiR)；Intrawhite Cf；JC-1；JO-JO-1；JO-PRO-1；LaserPro；Laurodan；LDS 751(DNA)；LDS 751(RNA)；雷可福(Leucophor)PAF；雷可福SF；雷可福WS；丽丝胺(Lissamine)罗丹明；丽丝胺罗丹明B；钙黄绿素/乙啶均二聚物；LOLO-1；LO-PRO-1；Lucifer Yellow；溶酶体示踪蓝(Lyso Tracker Blue)；溶酶体示踪蓝白(LysoTracker Blue-White)；溶酶体示绿(Lyso Tracker Green)；溶酶体示踪红(Lyso TrackerRed)；溶酶体示踪黄(Lyso Tracker Yellow)；LysoSensor蓝；LysoSensor绿；LysoSensor黄/蓝；Mag绿；Magdala红(Phloxin B)；Mag-Fura红；Mag-Fura-2；Mag-Fura-5；Mag-Indo-1；镁绿；镁橙；孔雀石绿；滨海蓝(Marina Blue)；Maxilon亮黄素10GFF；Maxilon亮黄素8GFF；部花青(Merocyanin)；甲氧基香豆素；线粒体示踪绿FM；线粒体示踪橙；线粒体示踪红；光辉霉素；单溴二胺；单溴二胺(mBBr-GSH)；单氯二胺；MPS(甲基绿派若宁二苯乙烯)；NBD；NBD胺；尼罗红；硝基苯并噁二唑；去甲肾上腺素；核固红；核黄；Nylosan Brilliant lavinE8G；Oregon绿；Oregon绿488-X；Oregon绿TM；Oregon绿TM488；Oregon绿TM500；Oregon绿TM514；太平洋蓝；副玫瑰苯胺(Feulgen)；PBFI；PE-Cy5；PE-Cy7；PerCP；PerCP-Cy5.5；PE-TexasRed[Red 613]；竹桃红B(Magdala红)；Phorwite AR；Phorwite BKL；Phorwite Rev；PhorwiteRPA；膦3R；光阻剂；藻红蛋白B[PE]；藻红蛋白R[PE]；PKH26(Sigma)；PKH67；PMIA；Pontochrome蓝黑；POPO-1；POPO-3；PO-PRO-1；PO-PRO-3；樱草黄；普施安黄；碘化丙啶(PI)；PyMPO；芘；派若宁；派若宁B；Pyrozal亮黄素7GF；QSY 7；芥子喹吖因；红613[PE-TexasRed]；试卤灵；RH 414；Rhod-2；罗丹明；罗丹明110；罗丹明123；罗丹明5GLD；罗丹明6G；罗丹明B；罗丹明B 200；罗丹明B extra；罗丹明BB；罗丹明BG；罗丹明绿；罗丹明Phallicidine；罗丹明鬼笔环肽；罗丹明红；罗丹明WT；玫瑰红(Rose Bengal)；R-藻青蛋白；R-藻红蛋白(PE)；S65A；S65C；S65L；S65T；SBFI；血清素；Sevron亮红2B；Sevron亮红4G；Sevron亮红B；Sevron橙；Sevron黄L；SITS；SITS(樱草灵)；SITS(二苯乙烯异硫代磺酸)；SNAFL钙黄绿素；SNAFL-1；SNAFL-2；SNARF钙黄绿素；SNARF1；钠绿；SpectrumAqua；SpectrumGreen；SpectrumOrange；Spectrum红；SPQ(6-甲氧基-N-(3-磺丙基)喹啉鎓)；二苯乙烯；磺酰罗丹明B can C；磺酰罗丹明Extra；SYTO 11；SYTO 12；SYTO 13；SYTO 14；SYTO 15；SYTO 16；SYTO 17；SYTO 18；SYTO 20；SYTO 21；SYTO 22；SYTO 23；SYTO 24；SYTO 25；SYTO 40；SYTO41；SYTO 42；SYTO 43；SYTO 44；SYTO 45；SYTO 59；SYTO 60；SYTO 61；SYTO 62；SYTO 63；SYTO 64；SYTO 80；SYTO 81；SYTO 82；SYTO 83；SYTO 84；SYTO 85；SYTOX蓝；SYTOX绿；SYTOX橙；四环素；四甲基罗丹明(TRITC)；Texas RedTM；Texas Red-XTM缀合物；硫二羰花青素(DiSC3)；噻嗪红R；噻唑橙；硫黄素5；硫黄素S；硫黄素TCN；Thiolyte；Thiozole橙；TinopolCBS(Calcofluor白)；TMR；TO-PRO-1；TO-PRO-3；TO-PRO-5；TOTO-1；TOTO-3；TriColor(PE-Cy5)；TRITC四甲基罗丹明异硫氰酸盐；纯蓝(True Blue)；纯红(TruRed)；Ultralite；荧光素钠B；Uvitex SFC；WW 781；X-罗丹明；XRITC；二甲苯橙；Y66F；Y66H；Y66W；YO-PRO-1；YO-PRO-3；YOYO-1；YOYO-3、SYBR绿、以及噻唑橙(互螯合染料)。
在一种实施方式中，发光接受体分子是海肾GFP(rGFP)。如本文所用的术语“海肾GFP”(Renilla GFP)是指源自海肾属的生物体，诸如Renilla reniformis或Renillamulleri的绿色荧光蛋白。它包括来自海肾生物体的天然GFP或其变体。在一种实施方式中，海肾GFP是Renilla reniformis GFP，在另外实施方式中，是Renilla reniformis GFP的天然形式(在氨基酸序列方面)。在一种实施方式中，rGFP包含图9D所示的序列。天然Renillamulleri GFP的氨基酸序列以GenBank登录号AAG54098.1公开。可以对海肾萤光素酶和/或海肾GFP的核酸序列进行密码子优化以在人细胞中表达(即“人源化”，参见例如，WO 2002/057451，用于人源化形式的Renilla mulleri GFP)。
适用于BRET的生物发光供体和荧光接受体分子的代表性组合(称为BRET对)包括萤光素酶(Luc)/GFP、Luc/Venus、Luc/Topaz、Luc/GFP-10、Luc/GFP-2、Luc/YFP、Luc/rGFP等。在另一实施方式中，在本文描述的生物传感器和方法中使用以下BRET配置之一：RlucII/coel-400a/增强蓝(EB)FP2、RlucII/coel-400a/超青色荧光蛋白(SCFP3A)、RlucII/coel-400a/mAmetrine、RlucII/coel-400a/rGFP，RlucII/coel-400a/mAmetrine。
在一种实施方式中，生物发光供体分子是海肾荧光素酶，而荧光接受体分子是海肾GFP。
在一种实施方式中，第二组分包括PM靶向部分。如本文所用，术语“质膜(PM)靶向部分”是指能够将生物发光供体或荧光接受体分子(例如，海肾GFP或海肾Luc)募集或螯合到PM的任何部分。因此，生物发光供体或荧光接受体分子可以与质膜上存在的任何蛋白质(例如，在PM上存在的受体或任何其他蛋白质)或其片段融合。这种蛋白的实例是小窝蛋白1(Caveolin-1)，它是在许多细胞类型中存在的小窝(一类对应于质膜的小(50-100nm)内陷的脂筏)的主要成分。已经确定通过CAV1基因的可变剪接产生的小窝蛋白1的两种同工型：小窝蛋白1α(包含残基2-178)和小窝蛋白1β(对应于32-178序列)。此类部分的其他实例包括肽/多肽，其包含用于蛋白质脂质化/脂肪酸酰化(诸如肉豆蔻酰化，棕榈酰化和异戊烯化)的信号序列，以及多碱性结构域。已知有几种蛋白质被肉豆蔻酰化、棕榈酰化和/或异戊烯化(例如，蛋白激酶和磷酸酶，诸如Yes、Fyn、Lyn、Lck、Hck、Fgr、Gα蛋白、一氧化氮合酶、ADP-核糖基化因子(ARF)、钙结合蛋白和膜或与细胞骨架相关的结构蛋白诸如MARCKS(参见例如，Wright等人,J Chem Biol.Mar 2010；3(1):19-35；Alcart-Ramos等人,Biochimicaet Biophysica Acta(BBA)–Biomembranes,Volume 1808,Issue 12,December 2011,Pages2981-2994)，以及因此任何这些蛋白的信号序列的肉豆蔻酰化、棕榈酰化和异戊烯化(例如，香叶基香叶基化)可用于生物传感器。在一种实施方式中，肉豆蔻酰化和/或棕榈酰化序列来自Lyn激酶。
在一种实施方式中，PM膜靶向部分包含CAAX基序(C是半胱氨酸残基，AA是两个脂族残基，并且X代表任何氨基酸。在“CAAX蛋白”中存在CAAX基序，“CAAX蛋白”定义为在C末端具有指导其翻译后修饰的特定氨基酸序列的一组蛋白质。CAAX蛋白涵盖多种分子，其包括核纤层蛋白(中间丝)，诸如前核纤层蛋白A、核纤层蛋白B1和核纤层蛋白B2，Ras和多种GTP结合蛋白(G蛋白)，诸如Ras、Rho、Rac和Cdc42，几种蛋白激酶和磷酸酶等(参见例如，Gao等人,Am J Transl Res.2009；1(3):312-325)。C末端的端部带有CAAX基序或框的蛋白质在该蛋白质迁移到质膜或核膜并发挥不同的功能之前，该蛋白质通常需要异戊二烯化过程。在一种实施方式中，CAAX框源自人RAS家族蛋白，例如HRAS、NRAS、Ral-A、KRAS4A或KRAS4b。RAS、NRAS、KRAS4A或KRAS4b的最后几个C末端残基(称为高变区或HVR)描述如下，假定的最小质膜靶向区域用斜体标出，并在CAAX框上加下划线(参见例如，Ahearn等人,NatureReviews Molecular Cell Biology 13:39-51,January 2012)：HRAS:KLNPPDESGPGCMSCKCVLS(SEQ ID NO：41)；NRAS：KLNSSDDGTQGCMGLPCVVM(SEQ ID NO：42)；KRAS4A：KISKEEKTPGCVKIKKCIIM(SEQ ID NO：43)；KRAS4B：KMSKDGKKKKKKSKTKCVIM(SEQ IDNO：44)；Ral-A/Ral1：KNGKKKRKSLAKRIRERCCIL(SEQ ID NO:45)。在一种实施方式中，PM靶向部分包含氨基酸序列GCMSCKCVLS(SEQ ID NO：60)，GCMGLPCVVM(SEQ ID NO：61)，CVKIKKCIIM(SEQ ID NO：62)，KKKKKKSKTKTKCVIM(SEQ ID NO：63)或KNGKKKRKSLAKRIRERCCIL(SEQ ID NO：45)，优选地，PM靶向部分包含来自KRAS4B的序列GKKKKKKSKTKCVIM(SEQ ID NO：1)。在另一实施方式中，PM靶向部分包含来自HRAS的质膜靶向棕榈酰化序列和来自Ral-A/Ral1的异戊烯化信号序列(序列：CMSCKCCIL，SEQ ID NO：4)。
几种蛋白质还包含靶向PM的非脂质多碱性结构域，诸如Ras小GTP酶、磷酸酶PTEN、非受体酪氨酸激酶Src、肌动蛋白调节剂WASP和MARCKS，以及G蛋白偶联受体激酶(GRK)，诸如GRK5。在一种实施方式中，多碱性结构域来自GRK5，并且包含序列SPKKGLLQRLFKRQHQNNSKS(SEQ ID NO：5)。
在一种实施方式中，第二组分包括胞内体靶向部分。如本文所用，术语“胞内体靶向部分”是指能够将生物发光供体或荧光接受体分子募集或螯合到胞内体例如早期胞内体的任何部分。几种胞内体靶向部分/标志物是本领域已知的，并包括Rab蛋白家族(RAB4、RAB5、RAB7、RAB9和RAB11)，甘露糖6-磷酸受体(M6PR)，小窝蛋白1和小窝蛋白2，转铁蛋白及其受体，网格蛋白以及包含FYVE结构域的蛋白质，诸如早期胞内体自身抗原1(EEA1)，Rabenosyn-5，用于受体活化的Smad锚(SARA)，Vps27p和Endofin。一些标志物对早期胞内体(例如，RAB4、转铁蛋白及其受体，以及包含FYVE结构域的蛋白)更具特异性，其他标志物对晚期胞内体(例如，RAB7，RAB9和M6PR)更具特异性，而另外一些标志物对再循环胞内体(例如，RAB11、RAB4)更具特异性。因此，这些蛋白或其合适的片段可以与生物发光供体分子(例如，海肾Luc)或荧光接受体分子(例如，海肾GFP)融合，以将它们连接/靶向至胞内体定位。
在一种实施方式中，胞内体靶向部分包含FYVE结构域。FYVE结构域由三个保守元素限定：N末端WxxD、中央RR/KHHCR和C末端RVC基序。含有FYVE结构域的人蛋白质的实例包括ANKFY1、EEA1 FGD1、FGD2、FGD3、FGD4、FGD5、FGD6、FYCO1、HGS MTMR3、MTMR4、PIKFYVE、PLEKHF1、PLEKHF2、RUFY1、RUFY2、WDF3、WDFY1、WDFY2、WDFY3、ZFYVE1、ZFYVE16、ZFYVE19、ZFYVE20、ZFYVE21、ZFYVE26、ZFYVE27、ZFYVE28以及ZFYVE9(EMBL-EBI、家族FYVE(PF01363))。在一种实施方式中，胞内体靶向部分包含人ZFYVE16/Endofin(UniProtKB-Q7Z3T8，SEQ ID NO：39)的FYVE结构域，例如人Endofin的大约747至805位残基或大约739至806位残基。
在一种实施方式中，第二组分包括高尔基体靶向部分。如本文所用，术语“高尔基体靶向部分”是指能够将生物发光供体或荧光接受体分子募集或螯合到高尔基体上的任何部分。几种高尔基体靶向部分/标志物是本领域已知的，并且包括eNOS(例如，其N末端部分，J.Liu等人,Biochemistry,35(1996),pp.13277-13281)、GM130、高尔基体蛋白97、58K蛋白、反式高尔基体网络膜蛋白2(TGOLN2)、TGN46、TGN38、甘露糖苷酶2、突触融合蛋白6、GM130(GOLGA2)、高尔基体蛋白160、Membrin(GS27)、GS28、Coatomer蛋白、Rbet1和RCAS1。因此，可以将这些蛋白质或其合适的片段融合到海肾Luc或海肾GFP上，以将它们连接/靶向至高尔基体定位。在一种实施方式中，高尔基体靶向部分是人eNOS蛋白(SEQ ID NO：46)的N末端部分，例如人eNOS1的1至73位残基。
在一种实施方式中，(i)PM靶向部分、胞内体靶向部分或高尔基体靶向部分与(ii)GASIP之间没有直接的蛋白-蛋白相互作用。
生物发光供体或荧光接受体分子可以在相对于靶向部分的N末端，内部或C末端融合。在一种实施方式中，PM靶向部分与所述生物发光供体或荧光接受体分子的C末端融合。在一种实施方式中，PM靶向部分与荧光接受体分子融合，优选与荧光接受体分子的C末端融合。在一种实施方式中，胞内体靶向部分与所述生物发光供体或荧光接受体分子的C末端融合，在另一种实施方式中，与所述荧光接受体分子的C末端融合。
生物发光供体或荧光接受体分子可以在相对于GASIP的N末端、内部或在C末端融合。在一种实施方式中，将生物发光供体或荧光接受体分子融合至GASIP的N末端。在另一实施方式中，将生物发光供体或荧光接受体分子融合至GASIP的C末端。在另一实施方式中，生物发光供体分子与GASIP融合，优选与GASIP的C末端融合。
术语“Rap1GAP的G蛋白结合结构域”是指包含Rap1 GTP酶活化蛋白1(Rap1GAP)蛋白(UniProt登录号P47736，SEQ ID NO：47)的结构域或其变体的多肽，其具有与Gi家族的Gα亚基蛋白(一种或多种)结合的能力。Rap1GAP的G蛋白结合结构域位于Rap1GAP的N末端部分，并且例如包含来自天然Rap1GAP的至少50、100、150、200、250、300、350或400个残基。Rap1GAP的G蛋白结合结构域可以包含不消除与Gα亚基蛋白结合的一个或多个突变。在一种实施方式中，Rap1GAP的G蛋白结合结构域包含天然Rap1GAP的1-420位或1-436位残基或其变体，其保留与Gi家族的Gα亚基蛋白(一种或多种)结合的能力。在另一实施方式中，Rap1GAP的G蛋白结合结构域包含1至442位残基或其变体，其保留与Gi家族的Gα亚基蛋白(一种或多种)结合的能力。在一种实施方式中，Rap1GAP的G蛋白结合结构域融合至生物发光供体分子的N末端。
在另一实施方式中，Rap1GAP的G蛋白结合结构域包含降低其对下游信号转导事件例如激酶(例如，蛋白激酶A)活化的敏感性的突变。可以例如通过引入一个或多个推定的磷酸化位点(例如，位于天然Rap1GAP的431、437、439和/或441位的丝氨酸残基)的突变(一种或多种)(例如缺失、替换)来实现对下游信号转导事件诸如激酶活化的敏感性的这种降低。在另一实施方式中，在Rap1GAP的G蛋白结合结构域中，在位置437、439和/或441上的一个或多个丝氨酸残基被突变，优选在Rap1GAP的G蛋白结合结构域中在位置437、439和/或441上的至少两个丝氨酸残基被突变，以及更优选，在位置437、439和441处的所有三个丝氨酸残基都被突变。在另一实施方式中，突变是替换，例如被不能磷酸化的氨基酸(例如丙氨酸残基)替换。在一种实施方式中，在Rap1GAP的G蛋白结合结构域中，位置437、439和441上的所有三个丝氨酸残基都被丙氨酸替换。在一种实施方式中，Rap1GAP的G蛋白结合结构域包含图9A和图9B所示的序列之一。
如本文所用的术语“G蛋白信号转导调节物(RGS)蛋白的G蛋白结合结构域”是指包含RGS蛋白的结构域或其变体的多肽，其具有与Gi家族的Gα亚基蛋白(一种或多种)结合的能力。RGS蛋白是22个蛋白的家族(RGS1-RGS22)，包含RGS框或RGS结构域(PROSITE条目PS50132)。RGS蛋白的G蛋白结合结构域可包含来自RGS蛋白的至少50、100、150、200、250、300、350或400个残基，或其变体，其保留结合Gi家族的Gα亚基蛋白(一种或多种)的能力。在一种实施方式中，GASIP包含RZ/A亚家族的RGS蛋白的G蛋白结合结构域，优选RGS17(RGSZ2，UniProt登录号Q9UGC6，SEQ ID NO：17)、RGS19(GAIP，UniProt登录号P49795，SEQ ID NO：18)或RGS20(RGSZ1，UniProt登录号O76081，SEQ ID NO：19)或其具有与Gi家族的Gα亚基蛋白(一种或多种)结合的能力的变体。在一种实施方式中，RGS17的G蛋白结合结构域包含天然RGS17的84-200位残基或其变体，其具有与Gi家族的Gα亚基蛋白(一种或多种)结合的能力。在一种实施方式中，RGS17的G蛋白结合结构域包含天然RGS17的64-210位残基或其变体，其具有与Gi家族的Gα亚基蛋白(一种或多种)结合的能力。在一种实施方式中，RGS19的G蛋白结合结构域包含天然RGS19的90-206位残基或其具有与Gi家族的Gα亚基蛋白(一种或多种)结合的能力的变体。在一种实施方式中，RGS19的G蛋白结合结构域包含天然RGS19的70-217位残基或其具有与Gi家族的Gα亚基蛋白(一种或多种)结合的能力的变体。在一种实施方式中，RGS20的G蛋白结合结构域包含天然RGS20的262-378位残基或其变体，其具有与Gi家族的Gα亚基蛋白(一种或多种)结合的能力。在一种实施方式中，RGS20的G蛋白结合结构域包含天然RGS20的242-388位残基或其变体，其具有与Gi家族的Gα亚基蛋白(一种或多种)结合的能力。在一种实施方式中，RGS蛋白的G蛋白结合结构域融合到生物发光供体分子的N末端。
包含可以在本文所述的系统/方法中使用的与Gi家族的Gα亚基蛋白(一种或多种)结合的结构域的另一种蛋白是肿瘤坏死因子α(TNFα)诱导蛋白8(TNFAIP8，UniProtKB登录号O95379，SEQ ID NO：48)。
术语“P63RhoGEF的G蛋白结合结构域”是指包含P63RhoGEF(Rho鸟嘌呤核苷酸交换因子25)蛋白(UniProt登录号Q86VW2，SEQ ID NO：25)的结构域或其变体的多肽，其具有与Gq家族的Gα亚基蛋白(一种或多种)结合的能力。P63RhoGEF的G蛋白结合结构域位于P63RhoGEF的C末端部分，并且例如包含天然P63RhoGEF的至少50、100、150、200、250、300、350或400个残基或其变体，其保留与Gq家族的Gα亚基蛋白(一种或多种)结合的能力。P63RhoGEF的G蛋白结合结构域可以包含不消除与Gα亚基蛋白结合的一个或多个突变。在一种实施方式中，P63RhoGEF的G蛋白结合结构域包含天然P63RhoGEF的348至466位残基，或其变体，其保留结合Gq家族的Gα亚基蛋白(一种或多种)的能力。在另一实施方式中，P63RhoGEF的G蛋白结合结构域包含天然P63RhoGEF的295至502位残基或其变体，其保留与Gq家族的Gα亚基蛋白(一种或多种)结合的能力。在一种实施方式中，P63RhoGEF的G蛋白结合结构域融合至生物发光供体分子的N末端。
术语“GRK2的G蛋白结合结构域”是指包含GRK2(β肾上腺素能受体激酶1)蛋白(UniProt登录号P25098，SEQ ID NO：27)的结构域或其变体的多肽，其具有与Gq家族的Gα亚基蛋白(一种或多种)结合的能力。GRK2的G蛋白结合结构域位于GRK2的N末端部分，并且例如包含天然GRK2的至少50、100、150、200、250、300、350或400个残基或其变体，其保留与Gq家族的Gα亚基蛋白(一种或多种)结合的能力。GRK2的G蛋白结合结构域可以包含不消除与Gα亚基蛋白结合的一个或多个突变。在一种实施方式中，GRK2的G蛋白结合结构域包含天然GRK2的54至175位残基，或其变体，其保留结合Gq家族的Gα亚基蛋白(一种或多种)的能力。在另一实施方式中，GRK2的G蛋白结合结构域包含天然GRK2的30至203位残基或其变体，其保留与Gq家族的Gα亚基蛋白(一种或多种)结合的能力。在一种实施方式中，GRK2的G蛋白结合结构域融合至生物发光供体分子的C末端。
包含可在本文所述的系统/方法中使用的结合Gq家族的Gα亚基蛋白(一种或多种)的结构域的其他蛋白包括GRK3(例如，
位残基)、PLCβ蛋白、RGS蛋白(例如，RGS2，
位残基)、TRP1和其他RhoGEF蛋白(例如，三功能结构域蛋白，TRIO)。包含如上文所定义的这些结构域或其保留结合Gq家族的Gα亚基蛋白(一种或多种)的能力的变体的多肽可以用于本文所述的系统/方法中。
术语“PDZRhoGEF的G蛋白结合结构域”是指包含PDZRhoGEF(Rho鸟嘌呤核苷酸交换因子11)蛋白(UniProt登录号O15085，SEQ ID NO：21)的结构域或其变体的多肽，其具有与G12/13家族的Gα亚基蛋白(一种或多种)结合的能力。PDZRhoGEF的G蛋白结合结构域位于PDZRhoGEF的中央部分，并且例如包含天然PDZRhoGEF的至少50、100、150、200、250、300、350或400个残基或其变体，其保留与G12/13家族的Gα亚基蛋白(一种或多种)结合的能力。PDZRhoGEF的G蛋白结合结构域可以包含不消除与Gα亚基蛋白结合的一个或多个突变。在一种实施方式中，PDZRhoGEF的G蛋白结合结构域包含天然PDZRhoGEF的306至486位残基或其变体，其保留与G12/13家族的Gα亚基蛋白(一种或多种)结合的能力。在另一实施方式中，PDZRhoGEF的G蛋白结合结构域包含天然PDZRhoGEF的281至483位残基或其变体，其保留与G12/13家族的Gα亚基蛋白(一种或多种)结合的能力。在一种实施方式中，PDZRhoGEF的G蛋白结合结构域融合至生物发光供体分子的N末端。
术语“P115RhoGEF的G蛋白结合结构域”是指包含P115RhoGEF(Rho鸟嘌呤核苷酸交换因子1)蛋白(UniProt登录号Q92888，SEQ ID NO：23)的结构域或其变体的多肽，其具有与G12/13家族的Gα亚基蛋白(一种或多种)结合的能力。P115RhoGEF的G蛋白结合结构域位于P115RhoGEF的N末端部分，并且例如包含天然P115RhoGEF的至少50、100、150、200、250、300、350或400个残基或其变体，其保留与G12/13家族的Gα亚基蛋白(一种或多种)结合的能力。P115RhoGEF的G蛋白结合结构域可以包含不消除与Gα亚基蛋白结合的一个或多个突变。在一种实施方式中，P115RhoGEF的G蛋白结合结构域包含天然P115RhoGEF的41至232位残基或其变体，其保留与G12/13家族的Gα亚基蛋白(一种或多种)结合的能力。在另一实施方式中，P115RhoGEF的G蛋白结合结构域包含天然P115RhoGEF的1至244位残基或其变体，其保留与G12/13家族的Gα亚基蛋白(一种或多种)结合的能力。在一种实施方式中，P115RhoGEF的G蛋白结合结构域融合至生物发光供体分子的N末端。
包含可以在本文所述的系统/方法中使用的与G12/13家族的Gα亚基蛋白(一种或多种)结合的结构域的其他蛋白包括RhoGEF，诸如Rho鸟嘌呤核苷酸交换因子12(LARG，UniProtKB登录号Q9NZN5，SEQ ID NO：49)(例如，残基
)，JNK相互作用亮氨酸拉链蛋白(JLP，C末端部分)钙粘蛋白，AXIN1(UniProtKB登录号O15169，SEQ ID NO：50，残基
对G12更具选择性)，PP2A(对G12更具选择性)，α可溶性NSF附着蛋白(αSNAP，UniProtKB登录号P54920，SEQ ID NO：51，N末端部分，对G12更具选择性)，多囊蛋白-1(UniProtKB登录号P98161，SEQ ID NO：52，对G12更具选择性)，RGS16(UniProtKB登录号O15492，SEQ ID NO：53，N末端部分，对G13更具选择性)，AKAP110(UniProtKB登录号O75969，SEQ ID NO：54，C末端部分，对G13更具选择性)，HS1相关蛋白X1(HAX1，UniProtKB登录号O00165，SEQ ID NO：55，残基
对G13更具选择性)。包含如上文所定义的这些结构域或其保留结合G12/13家族的Gα亚基蛋白(一种或多种)的能力的变体的多肽可以用于本文所述的系统/方法中。使用对亚基(G12或G13)之一更具选择性的结构域可用于评估内源性G12或G13蛋白的选择性活化。
结合Gs家族Gα亚基蛋白(一种或多种)的结构域包括SNX13(RGS-PX1，UniProtKB登录号Q9Y5W8，SEQ ID NO：56，残基
)，AXIN1(UniProtKB登录号O15169，SEQ IDNO：50，残基
)或TPR1(三十四肽重复域1，UniProtKB登录号Q99614，SEQ ID NO：57，C末端部分)。在一种实施方式中，Gs家族的Gα亚基蛋白是膜锚定的XL(α)s亚基。
在一种实施方式中，GASIP的长度为约50、75或100个氨基酸至约500、600、700或800个氨基酸，例如约100或150个氨基酸至约200、250、300、350、400或500个氨基酸。
在一种实施方式中，GASIP不包含Rap1GAP、RGS蛋白(例如，RGS17、19或20)、P63RhoGEF、GRK2、PDZRhoGEF或P115RhoGEF的全长序列，即它相对于全长蛋白质包含一个或多个突变(替换、缺失等)。在一种实施方式中，GASIP缺乏参与募集或锚定至PM的残基或结构域，例如，肉豆蔻酰化、棕榈酰化和/或异戊烯化信号序列。使用这些蛋白的截短和/或突变形式有利地允许最小化下游信号转导事件的效应(例如，激酶活化)和/或确保在G蛋白活化时GASIP正确定位(胞浆)以及易位至不同区室，从而提高了测定的可靠性和/或敏感性。
在一种实施方式中，本文所述的系统还包含重组Gα蛋白亚基。本文限定的Gα蛋白亚基包括但不限于17种不同的已知同工型，它们的剪接变体和任何突变的Gα蛋白，例如导致非选择性/混杂Gα的那些。在一种非限制性实施方式中，本文所述的Gα蛋白选自任何天然哺乳动物Gα蛋白中，其包括Gq、Gs、Gi1、Gi2、Gi3、Gt-cone、Gt-rod、Gt-gust、Gz、GoA、GoB、Golf、G11、G12、G13、G14和G15/G16(也称为GNA15)，这些同工型的剪接变体及其功能变体。在一种实施方式中，重组Gα蛋白亚单位是Gi家族的，例如，Gi1、Gi2、Gi3、GoA、GoB、Gt-cone、Gt-rod、Ggus和/或Gz。在一种实施方式中，Gα蛋白亚基是Gq家族的，例如Gq、G11、G14和/或G15/16)。在一种实施方式中，Gα蛋白亚基是G12/13家族的。
在一种实施方式中，重组Gα亚基多肽包含降低或消除通过GPCR的活化的至少一个突变。此类突变的Gα亚基多肽可用于评估非受体鸟嘌呤核苷酸交换因子(GEF)介导的G蛋白活化，即未通过GPCR参与诱导的G蛋白活化。在一种实施方式中，所述突变在羧基(C)末端结构域中，并且是优选在所述Gα亚基多肽的C末端的最后七个残基中的一个或多个中的突变。在一种实施方式中，突变是在C末端的最后的一个、2、3、4、5、6或7个残基的截短。在另一实施方式中，突变是在C末端上的至少的一个、2、3、4、5、6或所有残基的替换。在一种实施方式中，突变是在所述C末端结构域中至少一个保守的亮氨酸残基的缺失或替换，优选在所述C末端结构域中最后的保守亮氨酸残基(天然蛋白质的倒数第二个残基)的缺失或替换。在另一种实施方式中，突变是最后的保守亮氨酸残基的替换，优选替换成天冬氨酸(D)、脯氨酸(P)或精氨酸(R)残基。在一种实施方式中，突变的重组Gα亚基多肽是Gi家族的，例如Gi2或GoB。在另一实施方式中，突变的重组Gα亚基多肽包含图9B和9C中描绘的序列之一，例如Gαi2Δ5(SEQ ID NO：28)；Gαi2Δ2(SEQ ID NO：29)；Gαi2L-2G(SEQ ID NO：31)；Gαi2L-2P(SEQID NO：33)；Gαi2L-2R(SEQ ID NO：34)；Gαi2L-2D(SEQ ID NO：32)；Gαi2L-7G(SEQ ID NO：30)；GαoBΔ5(SEQ ID NO：36)；或GαoBL-2G(SEQ ID NO：35)。
如本文所用，术语“重组”是指由重组核酸分子(即通过分子生物学/基因工程技术制备的核酸)表达的蛋白质分子，例如在用编码该蛋白质的核酸(例如，存在于载体中)转染/转导细胞(或其后代)后表达的蛋白质(与细胞天然表达的蛋白质形成对比)。
在一种实施方式中，本文所述的系统还包括细胞表面受体。生物传感器的细胞可以天然地表达细胞表面受体，或者细胞表面受体可以是重组细胞表面受体(例如，细胞已经被编码细胞表面受体的核酸转染或转化)。如本文所用，术语“细胞表面受体”是指附着或嵌入质膜的蛋白，并且其在配体结合后诱导G蛋白活化。诱导G蛋白活化的细胞表面受体的实例包括G蛋白偶联受体(GPCR)、受体酪氨酸激酶类(RTK)、整合素类。G蛋白活化通常通过配体的GPCR参与进行，而G蛋白活化也可以通过非受体鸟嘌呤核苷酸交换因子(GEF)诸如GIV(与Gα相互作用囊泡相关蛋白，也称为Girdin)、NUCB1(核连蛋白1，也称为calnuc)、NUCB2和DAPLE(散乱相关蛋白)实现。例如，GIV活性与Gi活性的RTK(例如，EGFR)和整合素α-β复合物调节有关。在一种实施方式中，GASIP包含Rap1GAP的G蛋白结合结构域，并且细胞表面受体是GPCR或RTK。在另一种实施方式中，GASIP包含RGS蛋白，优选RGS17的G蛋白结合结构域，并且细胞表面受体是GPCR。
“GPCR”是指全长天然GPCR分子以及突变体GPCR分子。在Foord等人的(2005)Pharmacol Rev.57,279-288给出了GPCR的列表，其以引用方式并入本文，并且GPCR的更新列表可在IUPHAR-DB数据库中获得(Harmar AJ,等人(2009)IUPHAR-DB:the IUPHARdatabase of G protein-coupled receptors and ion channels.Nucl.Acids Res.37(Database issue):D680-D685；Sharman JL,等人,(2013)IUPHAR-DB:updated databasecontent and new features.Nucl.Acids Res.41(Database Issue):D1083-8)。
“RTK”是指全长天然RTK蛋白以及突变体RTK蛋白。RTK(EC2.7.10.1，根据IUBMB酶命名法)是针对许多多肽生长因子、细胞因子和激素的细胞表面受体，其特点为胞内区域包含负责这些受体的激酶活性的催化结构域，其催化受体自磷酸化和RTK底物的酪氨酸磷酸化。人类中已知58种RTK，分布在20个亚家族中((Robinson等人,Oncogene 2000,19(49):5548-57)。
“整合素”是指全长天然整合素蛋白以及突变体整合素蛋白。整合素由称为α和β的两个非共价相关的跨膜糖蛋白亚基组成。由9种类型的β亚基和24种类型的α亚基形成多种人整合素异二聚体。存在于脊椎动物的代表性整合素包括α1β1、α2β1、α3β1、α4β1、α5β1、α6β1、α7β1、αLβ2、αMβ2、αIIbβ3、αVβ1、αVβ3、αVβ5、αVβ6、αVβ8和α6β4。
如本文所用，术语“变体”(或“突变体”)是指具有与参考(例如，天然)序列至少60％的序列同一性并且保留其所需活性(例如结合于Gα亚基或充当BRET供体或接受体的能力)的蛋白/多肽。在其他实施方式中，该变体具有与参考(例如，天然)序列至少65、70、75、80、85、90、91、92、93、94、95、96、97、98或99％的相似性或同一性，并保留其所需活性。“相似性”和“同一性”是指两个多肽分子之间的序列相似性/同一性。可以通过比较比对序列中的每个位置来确定相似性或同一性。氨基酸序列之间的相似性或同一性的程度是该序列共有的位置上匹配或相同氨基酸数的函数。用于比较相似性或同一性的序列最优比对可以使用多种算法进行，诸如局部同源算法(Smith and Waterman,1981,Adv.Appl.Math 2:482)，同源比对算法(Needleman and Wunsch,1970,J.Mol.Biol.48:443)，相似性方法的搜索(Pearson and Lipman,1988,Proc.Natl.Acad.Sci.USA 85:2444)，以及这些算法的计算机化实现(诸如，Wisconsin Genetics Software Package,Genetics Computer Group,Madison,Wis.,U.S.A.中的GAP、BESTFIT、FASTA以及TFASTA)。序列相似性或同一性也可以使用BLAST算法来确定，其在Altschul等人,1990,J.Mol.Biol.215:403-10中描述(使用发布的默认设置)。可以通过美国国家生物技术信息中心网站获得进行BLAST分析的软件。
在实施方式中，本文描述的组分(融合分子)的结构域可以直接(例如，通过肽键)共价连接，或通过合适的接头部分(例如，一个或多个氨基酸的接头或其他类型的化学接头(例如，碳水化合物接头、脂接头、脂肪酸接头、聚醚接头、PEG等)“间接”共价连接。在一种实施方式中，可以在上述结构域之前(N末端)、之中或之后(C末端)插入一个或多个另外的结构域。在一种实施方式中，融合分子的结构域通过肽键共价连接。在另一种实施方式中，融合分子的一个或多个组分通过肽接头连接。在一种实施方式中，接头是肽接头，其长度通常为2至30个氨基酸的范围，例如约5至约20-25个氨基酸或约10至约15-20个氨基酸。每个接头的组成和长度可以根据所需的各种性质例如灵活性和水溶性来选择。例如，肽接头可包含相对小的氨基酸残基，包括但不限于甘氨酸；小氨基酸残基可减少肽接头的空间体积并增加肽接头的灵活性。肽接头也可包含极性氨基酸，包括但不限于丝氨酸。极性氨基酸残基可增加肽接头的水溶性。此外，可以使用程序诸如Globplot 2.3(Linding等人,GlobPlot:exploring protein sequences for globularity and disorder,Nucleic Acid Res2003-Vol.31,No.13,3701-8)来帮助确定无序性和球状性的程度，进而还确定其灵活性程度。在一种实施方式中，肽接头包含以下实施例和/或附图中公开的一个或多个氨基酸序列(SEQ ID NO：6、7、20、22、24和26)。
在一种实施方式中，该系统还包括表达本文限定的各种组分即第一组分和第二组分以及任选地重组Gα蛋白亚基和/或细胞表面受体的细胞。
在另一方面，本公开提供了编码本文限定的上述第一组分和/或第二组分的核酸或多个核酸。在一种实施方式中，核酸(一种或多种)存在于载体/质粒(或多个载体/质粒)中，在另一种实施方式中，存在于表达载体(一种或多种)/质粒(一种或多种)。此类载体包含编码可操作地连接至一个或多个转录调控序列(一种或多种)，例如启动子、增强子和/或其他调控序列的上述第一组分和/或第二组分的核酸(一种或多种)。在一种实施方式中，核酸编码第一组分和第二组分(多顺反子构建体)。
术语“载体”是指其能够转运已经与其连接的其他核酸的核酸分子。一类优选的载体是附加体，即能够进行染色体外复制的核酸。优选的载体是能够使与其连接的核酸自主复制和/或表达的载体。能够指导与其可操作地连接的基因表达的载体在本文中称为“表达载体”。可以通过本领域普通技术人员已知的标准技术来构建本发明的重组表达载体，例如前述Sambrook等人。有多种策略可用于连接DNA片段，策略的选择取决于DNA片段末端的性质，并且本领域技术人员可以容易地确定。本发明的载体还可包含其他序列元件，以促进载体在细菌和宿主细胞中的繁殖和选择。另外，本发明的载体可以包含一个或多个限制性核酸内切酶位点的核苷酸序列。编码序列(诸如选择标志物和报道基因)是本领域技术人员众所周知的。
可以将包含本文限定的一种或多种核酸的重组表达载体引入细胞(宿主细胞)，该细胞可以包括能够从所限定的重组表达载体表达蛋白编码区的活细胞。活细胞可以包括培养的细胞和活生物体内的细胞。因此，本发明还提供了含有本发明的重组表达载体的宿主细胞。术语“细胞”、“宿主细胞”和“重组宿主细胞”在本文可互换使用。这样的术语不仅指特定的宿主细胞，而且还指这种细胞的后代或潜在后代。由于突变或环境影响，某些修饰可能会在后代中发生，因此此类后代实际上可能与亲本细胞不同，但仍包括在本文所用术语的范围内。
可以通过常规转化或转染技术将载体DNA引入细胞。术语“转化”和“转染”是指用于将外来核酸引入宿主细胞的技术，包括磷酸钙或氯化钙共沉淀、DEAE-葡聚糖介导的转染、脂质转染、电穿孔、显微注射和病毒介导的转染。转化或转染宿主细胞的合适方法可以例如在Sambrook等人(Molecular Cloning:A Laboratory Manual,2nd Edition,ColdSpring Harbor Laboratory press(1989))以及其他实验手册中找到。“转录调节序列/元件”是通用术语，是指诱导或控制与其可操作地连接的蛋白质编码序列的转录的DNA序列，诸如起始和终止信号、增强子和启动子，剪接信号，多腺苷酸化信号。当第一核酸序列与第二核酸序列处于功能关系时，第一核酸序列与第二核酸序列“可操作地连接”。例如，如果启动子影响编码序列的转录或表达，则该启动子可操作地连接至编码序列。通常，可操作地连接的DNA序列是邻近的，并且在需要连接两个蛋白质编码区的情况下处于阅读框中。但是，由于例如增强子通常在与启动子相距几千碱基时起作用，并且内含子序列的长度可能是可变的，因此一些多核苷酸元件可以可操作地连接但不邻近。
在另一方面，本公开提供了试剂盒，其包含编码第一组分的第一核酸和编码第二组分的第二核酸，或编码第一组分和第二组分的核酸。在一种实施方式中，试剂盒还包含编码Gα蛋白亚基的核酸。在另一实施方式中，试剂盒还包含编码细胞表面受体的核酸。
在另一方面，本公开提供了一种细胞，其包含或表达以上限定的第一组分和/或第二组分。在一种实施方式中，已经用编码上述第一组分和/或第二组分的核酸转染或转化了细胞。在一种实施方式中，细胞还包含或表达重组Gα蛋白亚基。在一种实施方式中，细胞还包含或表达重组细胞表面受体，例如，GPCR、RTK或整合素。
本公开还提供了重组表达系统、载体和细胞，诸如上述的那些，用于使用例如本领域众所周知的培养基和试剂来表达本发明的第一组分和/或第二组分。所述细胞可以是能够表达以上限定的第一组分和第二组分(一种或多种)的任何细胞。合适的宿主细胞和表达蛋白的方法是本领域众所周知的。可以使用能够表达以上限定的组分(一种或多种)的任何细胞。例如，可以使用真核宿主细胞，诸如哺乳动物细胞(例如，啮齿动物细胞，诸如小鼠、大鼠和仓鼠细胞系、人细胞/细胞系)。在另一种实施方式中，上述细胞是人细胞系，例如胚胎肾细胞系(例如，HEK293或HEK293T细胞)。
在另一方面，本公开提供了一种用于确定剂是否调节感兴趣的G蛋白活化的方法，所述方法包括：
使本文限定的系统与生物发光供体分子的底物接触；
在所述剂存在和不存在下测量系统中的BRET信号；
其中在所述剂存在下相对于其不存在下所述BRET信号的差异表明所述剂调节所述感兴趣的G蛋白活化。
在一种实施方式中，该剂是激动剂，并且所述BRET信号的差异是增加。
在一种实施方式中，感兴趣的G蛋白是Gi蛋白亚基家族，并且其中GASIP包含本文限定的Rap1GAP或RGS蛋白的G蛋白结合结构域。在一种实施方式中，在上述方法中使用的系统包含Gi蛋白亚基家族的重组Gα亚基多肽，即重组Gi1、Gi2、Gi3、GoA、GoB或Gz亚基多肽。为了确定给定的测试剂的特性和特征(即为了确定测试剂活化的特定Gi蛋白亚基(一种或多种))，可以使用多个(即两个或更多个)系统执行上述方法，每个系统都包含不同的Gi蛋白亚基，例如包含重组Gi1的第一系统、包含重组Gi2的第二个系统、包含重组Gi3的第三系统等。
在另一实施方式中，感兴趣的G蛋白是Gq蛋白亚基家族，并且其中GASIP包含本文限定的P63RhoGEF或GRK2的G蛋白结合结构域。在一种实施方式中，在上述方法中使用的系统包含Gq蛋白亚基家族的重组Gα亚基多肽，即重组Gq、G11、G14或G15亚基多肽。为了确定给定的测试剂的特性和特征(即为了确定测试剂活化的特定Gq蛋白亚基(一种或多种))，可以使用多个(即两个或更多个)系统执行上述方法，每个系统都包含不同的Gq蛋白亚基，例如包含重组Gq的第一系统、包含重组G11的第二个系统、包含重组G14的第三系统等。
在另一实施方式中，感兴趣的G蛋白是G12/13蛋白亚基家族，并且其中GASIP包含本文限定的PDZRhoGEF或P115RhoGEF的G蛋白结合结构域。在一种实施方式中，在上述方法中使用的系统包含G12/13蛋白亚基家族的重组Gα亚基多肽，即重组Gq、G12或G13亚基多肽。为了确定给定的测试剂的特性和特征(即为了确定测试剂活化的特定G12/13蛋白亚基(一种或多种))，可以使用多个(即两个或更多个)系统执行上述方法，每个系统都包含不同的G12/13蛋白亚基，例如包含重组G12的第一系统和包含重组G13的第二系统。
在另一方面，本公开提供了一种用于确定剂是否调节非受体鸟嘌呤核苷酸交换因子(GEF)介导的G蛋白活化的方法，所述方法包括：
使本文限定的系统与生物发光供体分子的底物接触；
在所述剂存在和不存在下，在系统中测量BRET信号，其中所述系统包含含有减少或消除通过GPCR的活化的至少一个突变的重组Gα亚基多肽。
其中在所述剂存在下相对于其不存在下所述BRET信号的差异表明所述剂调节非受体GEF介导的G蛋白活化。
在一种实施方式中，该剂是激动剂，并且所述BRET信号的差异是增加(或更高的BRET信号)。
在另一实施方式中，所述剂是拮抗剂，并且所述BRET信号的差异是减少(或更低的BRET信号)。在一种实施方式中，为了确定剂是否为拮抗剂，将系统与已知的激动剂接触，并且在剂的存在下由已知的激动剂诱导的BRET信号的降低表明该剂为拮抗剂。
在另一方面，本公开提供了一种用于确定剂是否诱导GPCR介导的G蛋白活化或RTK/GEF介导的G蛋白活化的方法，该方法包括：
使用第一生物传感器在剂存在下测量BRET信号，其中GASIP包含Rap1GAP的G蛋白结合结构域，优选Rap1GAP的G蛋白结合结构域，该G蛋白结合结构域包括降低其对下游信号转导事件敏感性的突变(例如，激酶活化)，更优选包括在位置437、439和/或441处一个或多个丝氨酸残基的突变(例如，Rap1GAP(SSS-AAA))，如上文所限定；
使用第二生物传感器在所述剂存在下测量BRET信号，其中所述GASIP包含RGS蛋白的G蛋白结合结构域，优选RGS17，如上文限定；
其中在第一生物传感器情况下存在剂时，BRET信号增加(例如，剂量依赖性增加)，而在第二生物传感器情况下存在剂时，BRET信号没有增加，这表明该剂诱导RTK/GEF介导的G蛋白活化；并且其中在第一生物传感器和第二生物传感器情况下在剂的存在下BRET信号的增加(例如，剂量依赖性增加)指示该剂诱导GPCR介导的G蛋白活化。
术语“化合物”、“剂”、“测试化合物”或“测试剂”是指可以通过本文所述的系统/测定法筛选的任何分子(例如，候选药物)，可以从许多来源获得，包括合成的或天然的化合物的库。例如，许多手段可用于多种有机化合物和生物分子的随机和定向合成，包括表达随机寡核苷酸。替代地，可获得或容易产生细菌、真菌、植物和动物提取物形式的天然化合物的库。另外，可以通过常规化学、物理和生化手段容易地修饰天然或合成产生的库和化合物。
在一种实施方式中，本文所用的“增加的信号”或“更高的信号”是指相对于在不存在测试剂的情况下测得的参考信号高至少10、20、30、40、45或50％的信号。在另一种实施方式中，“更高的信号”是通过显示在测试剂存在下测得的信号相对于测试剂不存在的统计学显著差异(使用适当的统计分析确定)来确定的，例如通过组合多个样品获得的结果。用于确定不同数据集之间的显著差异的统计分析(ANOVA、学生t检验、卡方等)在本领域中是已知的，并且可以使用合适的计算机程序来执行这种分析。
阳性对照和阴性对照可用于本文所述的方法/测定法。可以多次进行对照样品和测试样品以获得统计上显著的结果。
在一种实施方式中，上述方法是高通量方法(高通量筛选，HTS)。如本文所用，术语“高通量筛选”(HTS)是指一种允许快速且并行地筛选大量化合物(数百、数千)针对靶分子的结合活性或生物活性的方法。这种HTS方法通常在具有数个孔例如384、1536或3456个孔的微量滴定板中进行。对于HTS，以高灵敏度、准确性和可再现性检测读出信号非常重要。确定方法是否适合HTS或与HTS兼容的一种方式是测量Z因子，如以下实例中所述。理想地，Z因子应为至少0.5(即在0.5和1之间)，并且优选地为至少0.55、0.6、0.65、0.7、0.75或0.8。
测量BRET信号的方法和设备是本领域众所周知的。例如，可以通过测定BRET接受体信号的强度(光强度)和/或通过计算BRET接受体发射的信号或光强度与BRET供体发射的信号或光强度之比(BRET比率)来测量BRET信号。BRET信号可以使用以下测量：酶标仪或显微镜，带有适当滤光片组，以检测BRET供体和/或BRET接受体的光发射。
本文所述的系统和方法可用于HTS中，以确定例如具有调节G蛋白活化(通过GPCR或其他通过G蛋白信号转导的受体)的活性的，并因此是潜在的候选药物或用于治疗涉及G蛋白信号转导的疾患(例如，抑制或刺激G蛋白将是有益)的化合物。本文所述的系统和方法也可以用于进行配体特性分析，例如，用于确定配体调节通过G蛋白的GPCR或其他受体信号转导的途径。
本文所述的系统和方法可用于筛选化合物库以确定用于药物开发的潜在候选物。当本文所述的系统与化合物接触时，BRET信号比率的变化可以确定药物或药物活性化合物，并且可以确定其对细胞途径的作用。而且，本文描述的系统和方法也可以用于确定候选药物的剂量依赖性。
机电读板仪可用于检测信号比率变化。使用本发明的系统，此类读板仪可用于HTS、候选药物筛选和药物剂量依赖性研究。可用于实施本发明的读板仪的实例包括由PerkinElmer(Boston,MA)提供的FusionTM系列读板仪，包括PerkinElmer FusionTM通用微板分析仪装置。PerkinElmer EnVisionTMHTS模型也可以用于实施本文所述的方法。
实施例
通过以下非限制性实施例进一步详细说明本发明。
实施例1：材料和方法
材料.血管紧张素II(AngII；[Asp-Arg-Val-Tyr-Ile-His-Pro-Phe]，SEQ ID NO：58)、Forkolin、多巴胺和聚鸟氨酸来自Sigma-Aldrich。u46619、I-BOP、CTA2、U51605、I-SAP、SQ 29558来自
(Ann Arbor,MI)。人EGF来自Cedarlane实验室。胰激肽([Lys0]-缓激肽)来自Eurogentec。WAY-100635、A 412997、L 741742、SNC-80和AR-M100390来自Tocris Bioscience(Bio-Techne)。血小板活化因子(PAF-C16)来自Enzo LifeSciences，Inc.。鲑鱼精子DNA、杜尔贝科改良伊格尔培养基(DMEM)、胎牛血清、小牛血清、
和其他细胞培养试剂购自Life Technologies(Thermo FisherScientific)以及购自Wisent Inc。25kDa线性聚乙烯亚胺(PEI)来自Polysciences Inc.。Prolume Purple I购自
Technology。Phusion DNA聚合酶来自Thermo FisherScientific。限制性酶、T4 DNA连接酶和吉布森组装混合物获自New England BiolabsLtd.。白色96孔聚苯乙烯细胞培养微孔板，固体底部(CellStar 655 083)来自Greiner。
质粒和构建.质膜标志物rGPF-CAAX和早期胞内体标志物rGFP-FYVE构建体已被描述(Namkung Y.等人2016；Nat Commun.7:12178)。编码TPα受体构建体、所有Gα亚基、Gβ1亚基和Gγ1亚基的质粒来自cdna.org。对带有信号肽：MDSKGSSQKGSRLLLLLVVSNLLLCQGVVS(SEQ ID NO：59)的人受体D4R、At1AR和Mu阿片样受体(hMOR1)的编码序列(cds)进行了序列优化，在GeneART(Thermo Fisher Scientific)合成并通过来自Invitrogen(ThermoFisher Scientific)的pCDNA3.1(+)中的吉布森组装亚克隆；hMOR1被亚克隆到pLVX IRESpuro(Clontech，CA)中。Dr Yosef Yarden提供了编码人EGFR的构建体(Tzahar E.等人1996；Molecular and Cellular Biology 16(10):5276–5287)。将缓激肽B2受体(BKB2R)cds和RAPGAP(1-442)-Rluc8构建体(参见图9A和9B)进行序列优化，在Topgenetech合成并亚克隆到pCDNA3.1(+)中；在RapGAP(1-442)和Rluc8之间存在肽接头GSGGGSGGGA(SEQ IDNO：6)(参见图2A)。通过用pCDNA3.1(+)RapGAP(1-442)-Rluc8进行PCR扩增，并通过吉布森组装亚克隆至pCDNA3.1 Hygro(+)GFP10-RlucII中，得到编码RlucII标记形式的以下构建体：RapGAP(1-442)、Rap1GAPΔCT(1-420)、Rap1GAPΔSSS(1-436)、Rap1GAP SSS-AAA(1-442；S437A/S439A/S441A)、Rap1GAP SSS-TTT(1-442；S437T/S439T/S441T)、Rap1GAP SS-AA(1-442；S437A/S441A)、Rap1GAP SS-DA(1-442；S437A/S441A)、Rap1GAP SS-AD(1-442；S437A/S441D)、Rap1GAP SS-DD(1-442；S437A/S441A)，其中用Rap1GAP的不同变体的编码序列代替GFP10(参见图2A、9A和9B)。肽接头：GSAGTGGRAIDIKLPAT(SEQ ID NO：7)存在于RAPGAP和RlucII之间(参见图2A)。通过从pCDNA 3.1HA-RGS17(来自cDNA.org)进行PCR扩增并通过吉布森组装亚克隆到pCDNA3.1 Hygro(+)GFP10-RlucII(其中用RGS(RGS17 64-210)的cds替换GFP10)得到编码用RlucII标记的人RGS17(残基：64-210)的RGS结合结构域的构建体(见图3A和9D)。通过从pCDNA 3.1HA-RGS19(来自cDNA.org)进行PCR扩增并通过吉布森组装将其亚克隆到pCDNA3.1Hygro(+)GFP10-RlucII(其中用RGS19(70-217)的编码序列替换GFP10)得到编码用RlucII标记的人RGS19(残基：70-217)的RGS结合结构域的构建体(见图3A和9D)。通过从pCDNA 3.1HA-RGS20 var1(来自cDNA.org)进行PCR扩增并通过吉布森组装将其亚克隆到pCDNA3.1 Hygro(+)GFP10-RlucII(其中用RGS(RGS20 242-388)的编码序列替换GFP10)得到编码用RlucII标记的人RGS20 var1(残基：242-388)的RGS结合结构域的构建体(见图3A和9D)。肽接头：GSAGTGGRAIDIKLASAT(SEQ ID NO：20)存在于Rap1GAP和RlucII之间(参见图3A)。通过从IMAGE克隆(OpenBiosystems)进行PCR扩增并通过吉布森组装亚克隆至pCDNA3.1 Hygro(+)GFP10-RlucII(其中替换GFP10)得到编码用RlucII标记的人PDZRhoGEF(残基：281-483)和P115RhoGEF(残基：1-244)的G12/13结合结构域的构建体(见图4A，5A和9D)。肽接头：GILREALKLPAT(SEQ ID NO：22)和RLKLPAT(SEQ ID NO：24)分别存在于RlucII与PDZRhoGEF和P115RhoGEF的G12/13结合结构域之间(参见图4A和5A)。通过从IMAGE克隆(OpenBiosystems)并通过吉布森组装亚克隆至pCDNA3.1 Hygro(+)GFP10-RlucII(其中替换GFP10)得到编码用RlucII标记的人P63RhoGEF(残基：295-502)的Gq结合结构域的构建体(见图6A和9B)。肽接头：ASGSAGTGGRAIDIKLPAT(SEQ ID NO：26)存在于Gq结合结构域和RlucII之间(参见图6A)。通过从pcDNA3.1Z-GRK2-GFP10进行PCR扩增并通过吉布森组装亚克隆到pCDNA3.1 Hygro(+)GFP10-RlucII和pCDNA3.1(+)RlucII-GFP10st2(其中替换GFP10并分别创建RGS(GRK2)-RlucII和RlucII-RGS(GRK2))得到编码用RlucII标记的人GRK2(残基：30-203)的RGS结构域的构建体(见图7A和9D)。在两种构建体中，肽接头：GSAGTGGRAIDIKLASAT(SEQ ID NO：20)存在于RGS(GRK2)结构域和RlucII之间(参见图7A)。通过从pCDNA3.1(+)GNAi2和pCDNA3.1(+)GNAoB(来自cdna.org)进行PCR扩增并通过吉布森组装亚克隆至pCDNA3.1 Zeo(+)中得到编码Gαi2和GαoB突变体的构建体：Gαi2Δ5(＝1-350)、Gαi2Δ2(＝1-353)、Gαi2L-7G(＝L354G)、Gαi2 L-2G(＝L354G)、Gαi2 L-2D(＝L354D)、Gαi2L-2P(＝L354P)、Gαi2 L-2R(＝L354R)、GαoB L-2G(＝L353G)、GαoBΔ5(＝1-349)。
细胞培养和瞬时转染.将人胚胎肾293(HEK293)细胞在37℃下在5％CO2湿润气氛中保持在补充有10％胎牛血清、100单位/ml青霉素/链霉素的杜尔贝科改良伊格尔培养基(DMEM)中。在补充有5％胎牛血清和20μg/ml庆大霉素的DMEM中培养HEK293SL细胞。使细胞在37℃，5％CO2和90％湿度下生长。
使用聚乙烯亚胺(PEI)进行转染：实验前两天，将HEK293细胞用不含钙或镁的PBS洗涤，脱附，并使用PEI作为转染剂(比例为3：1，PEI/DNA)用指定的质粒转染。然后将细胞以每孔35000个细胞的密度直接接种在聚L-鸟氨酸氢溴酸盐预处理或未预处理的96孔板上。使用未处理的板在Spark 10M读取仪中得到所有实验读数。
BRET测量.转染后48小时，将细胞用预热的Tyrode缓冲液(140mM NaCl、2.7mMKCl、1mM CaCl2、12mM NaHCO3、5.6mM D-葡萄糖、0.5mM MgCl2，0.37mM NaH2PO4、25mM HEPES，pH 7.4)洗涤两次，之后用各种浓度的配体在室温(RT)或37℃下刺激，如所指出的。在BRET测量之前至少6min(37℃)至10min(在RT)，将细胞可渗透的底物腔肠素紫I(coelenterazine Purple I)加入Tyrode缓冲液至终浓度为1μM。测量是在Spark 10M读取仪(Tecan Life Sciences；供体：360nm/380nm&接受体：505nm/570nm)，LB 941多模式读板仪(Berthold Technologies；供体：400/470-nm和接受体：515/520-nm滤光片)或SynergyNeo(Biotek：(供体：400/480nm，接受体：515/530nm)上进行。通过计算rGFP(接受体)发出的光强度与RlucII(供体)发出的光强度之比来确定BRET信号。所有的BRET测量都是在Tristar读取仪(图1-2M&图2R-7F)中在37℃下，在Spark 10M读取仪(图2N-2Q&图8B-8S)中在RT下中或在Synergy Neo(图8T-8X)中在RT下进行。
Z'因子确定.如Zhang等人所述计算Z'因子值(J Biomol Screen.1999；4(2):67-73)。超过0.4的Z'因子被认为是鲁棒测定法。
数据分析.EC50值的估计是使用
Prism曲线拟合程序计算的。整个研究过程中呈现的曲线代表了最佳拟合，也是使用此
Prism程序生成的。
实施例2：以G蛋白家族选择性方式监测G蛋白活化的系统和测定法的产生和特异性
图1A至1E示出了基于来自G蛋白效应子的Gα亚基相互作用多肽(GASIP)的易位，以G蛋白家族选择性方式并在不同的细胞区室中监测G蛋白活化的系统和测定法的产生和特异性。图1A描绘了基于效应子的传感器监测以下的原理，i)GPCR介导的直接G蛋白活化；以及ii)鸟嘌呤核苷酸交换因子(GEF)介导的G蛋白活化。将表达受体，用rGFP标记的亚细胞定位结构域(例如针对质膜(PM)或针对早期胞内体(EE))，用BRET供体(例如，RlucII)标记的特定效应子的Gα相互作用结构域的细胞暴露于激动剂以活化共表达的G蛋白。在i)中，激动剂诱导的GPCR刺激直接活化了G蛋白，其将标记的效应子从细胞质募集到rGFP标记的膜。在ii)中，在RTK(例如，EGFR)或整合素α-β复合物活化后，由GEF诸如GIV/Girdin的募集介导了G蛋白活化。不需要修饰G蛋白亚基来监测其活化，通过共表达Gα蛋白亚基与研究的受体并通过使用对G蛋白每个家族具有特异性的GASIP(诸如针对Gi家族(Gi1、Gi2、Gi3、GoA、GoB、Gz)的Rap1GAP的G蛋白结合结构域(图1B)，针对Gq家族(Gq、G11、G14&G15)的P63RhoGEF(P63RG)的G蛋白结合结构域(图1C)以及针对G12/13家族的PDZRhoGEF(PDZRG)的G蛋白结合结构域(图1D和1E))可以实现特异性。当使用Rap1GAP的G蛋白结合结构域时，只有Gi家族的成员显示出比模拟条件更大的应答(内源表达的Gi1、i2&i3蛋白的活化)(图1B)，当使用PDZRG时，只有G12/13家族的成员显示出比模拟条件更大的应答(内源表达的G12和G13蛋白的活化)(图1C)，以及当使用P63RG时，仅Gq家族的成员显示出比模拟条件更大的应答(内源表达的Gq和G11蛋白的活化)(图1D和1E)，因此证实了测定法的特异性。
实施例3：用于监测Gi家族的G蛋白的活化的系统和测定法
图2A至2Z示出了用于监测Gi家族(Gi1、i2、i3、oA、oB、z)的G蛋白活化的基于Rap1GAP的BRET传感器的优化和使用。测试的各种构建体示于图2A。示于图2B至2E中的结果显示了使用测试的两种BRET供体(Rluc8和RlucII)获得了可检测的BRET信号，其中相对于Rluc8，RlucII通常给出更好的动态窗口。示于图2F和2G的结果证明Rap1GAP(1-442)构建体对磷酸化敏感，如用毛喉素(其可促进cAMP产生增加和蛋白激酶A的活化，从而导致不同蛋白的磷酸化)预处理细胞时测得的较低应答所证明的那样，这可能影响测定法。产生并测试C末端截短的变体(1-420和1-436)以及包含在推定的磷酸化位点(残基Ser 437、439和441)的Ser至Ala和Ser至Asp替换的组合的不同突变体。RAP1GAP的C末端截短变体(ΔCT)对毛喉素的作用不敏感，但该较短片段(Rap1GAP 1-420)的窗口显著低于Rap1GAP(1-442)的窗口(图2H)。在测试的其他突变体中，仍受毛喉素影响的突变体仅为Rap1GAP(SS-AD)、Rap1GAP(SSS-TTT)和Rap1GAP(SS-AA)，后者显示的动态窗口相当于Rap1GAP(1-442)-RlucII(图2J-2Q)。Rap1GAP(SSS-AAA)用于以下描述的其他实验。
获得多巴胺促进的刺激后多巴胺D4受体(D4R)的G蛋白分析(profiling，谱)，Rap1GAP(SSS-AAA)-RlucII易位至质膜，每种G蛋白(Gi1、Gi2、Gi3、GoA和GoB)显示出具有不同药理学特点的剂量-应答曲线(图2R)。Gz活化被用于分析几种多巴胺受体配体(A412997、多巴胺、L741 742和Way-100635)对5种多巴胺受体的特性，结果在图2S至2W中示出。显示多巴胺可活化所有受体(EC50：D1R＝380nM，D2R＝2.6nM，D3R＝0.50nM，D4R＝0.11nM，D5R＝51nM)，已知的D4R拮抗剂L741,742没有显示对任何多巴胺受体的激动剂或逆性质。WAY-100635仅对D4R(EC50＝0.83)显示出激动剂特性，而A412,997活化了D3R(EC50＝281nM)和D4R(EC50＝0.04nM)二者，但没有活化D1R、D2R或D5R。在图2X-2Z中描绘的结果表明该测定法是鲁棒的，并且与高通量筛选(HTS)兼容，对于Gi2、GoA和Gz活化的Z'因子分别评估为0.812、0.703和0.607。
图3A-3D表明G蛋白信号转导调节物(RGS)蛋白的成员(诸如RGS17、19和20)的RGS结构域可以用作Rap1GAP的替代物以监测G蛋白活化。基于RGS17(残基64-210)、RGS19(残基70-217)和RGS20(残基242-388)蛋白的RGS结构域的RlucII标记的构建体在图3A中给出。仅使用这些蛋白的Gα结合RGS结构域有利地允许在G蛋白活化时进行胞浆定位和易位至不同的区室，而没有影响全长蛋白中存在的其他结构域或这些蛋白质的N末端部分存在的棕榈酰化位点。RGS(RGS17)-RlucII(图3B)、RGS(RGS19)-RlucII(图3C)和RGS(RGS20)-RlucII(图3D)获得的剂量-应答曲线是针对D4R/多巴胺介导的Gi1、Gi2、Gi3、GoA、GoB和Gz在PM处的活化而给出的。使用3种RGS构建体获得了相似的结果(偶联和EC50)，证实了这些基于RGS的构建体可用于监测Gi和Go活化。结果与基于Rap1GAP的构建体所获得的结果相似，不同之处在于Gz的共表达不会导致可与模拟条件区分开的信号，并且基于RGS的构建体相对于基于Rap1GAP-的构建体，其Gi3的动态窗口更小。
实施例4：用于监测G12/13家族的G蛋白的活化的系统和测定法
在图4A-G中描述了用于监测G12/13家族的G蛋白的活化的基于PDZRhoGEF的BRET系统的优化和使用。包含G12/13结合结构域的PDZRhoGEF的片段(残基281-483，PDZRG)在C末端用BRET供体RlucII标记(图4A)。已显示用于测量G12和13活化的动态窗口直接取决于Gα亚基的表达水平，但表达水平不影响I-BOP/TPαR介导的G12和13活化的效力，如用不同量的G12和13获得了相当的(可比较的)LogEC50值证明了这一点(图4B和4C)。PDZRG-RlucII用于分析TPαR配体对PM处G12和G13活化的特性。用已知的完全激动剂(U46619、I-BOP、CTA2)，用一种部分激动剂(U51605)以及拮抗剂I-SAP和SQ 29,558刺激细胞。在图4D和4E中描绘的结果表明SQ 29,558没有作为TPαR激动剂起作用。与它们的已知活性/性质一致，U46619、I-BOP和CTA2是G12和13活化的完全激动剂，证实了监测G12/13活化的测定法的有效性。有趣的是，U51605和I-SAP被显示作为有偏好的配体起作用。已证明U51605是G12活化的部分激动剂(最大的40％)，而几乎是G13的完全激动剂(最大的93％)。相反，I-SAP刺激不能在测定中诱导显著的G12活化(图4D)，但是检测到其对于G13的部分激动剂活性(最大的53％)(图4E)。最后，获得对于G12活化评价为0.645的Z'因子并且对于G13活化评估为0.812的Z'因子(图4F和4G)，再次表明该测定法是鲁棒的并且与基于G12/G13调节的高通量筛选应用兼容。
在图5A-E中描述了用于监测G12/13家族的G蛋白的活化的基于P115RhoGEF的BRET系统的优化和使用。包含G12/13结合结构域的P115RhoGEF的片段(残基1-244，P115RG)在C末端用BRET供体RlucII标记(图5A)。P115RG-RlucII用于分析TPαR配体对PM处G12和G13活化的特性。用U46619、I-BOP、CTA2、U51605、I-SAP和SQ 29,558刺激细胞。在图5B和5C中描绘的结果与用PDZRG-RlucII获得的结果一致，表明SQ 29,558不作为TPαR激动剂起作用，U46619、I-BOP和CTA2是对G12和13活化的完全激动剂，显示U51605和I-SAP作为有偏好的配体起作用。U51605被证明是对G12(最大的14％)和G13(最大的60％)活化的部分激动剂，而I-SAP刺激是对G13(最大的19％)的部分激动剂，对G12活化是中性的。最后，获得对于G12活化评价为0.703的Z'因子并且对于G13活化评估为0.743的Z'因子(图5D和5E)，再次表明该测定法是鲁棒的并且与基于G12/G13调节的高通量筛选应用兼容。
实施例5：用于监测Gq家族的G蛋白的活化的系统和测定法
在图6A-S中描述了用于监测Gq家族(Gq、G11、G14和G15)的G蛋白活化的基于P63RhoGEF的BRET系统的优化和使用。包含Gq结合结构域的P63RhoGEF的片段(残基295-502，P63RG)在C末端用BRET供体RlucII标记(图6A)。全长P63RhoGEF蛋白通过其N末端与PM相关联。与PDZRG-RlucII和P115RG-RlucII一样，使用仅包含P63RhoGEF的G蛋白结合结构域的片段有利地允许在G蛋白活化时胞浆定位并易位至不同的区室。使用不同的膜标志物(针对PM的rGFP-CAAX和针对EE的rGFP-FYVE)来监测PM处和早期胞内体(EE)处的G蛋白活性的优化在图6B-6E(对于PM)和6F-I(对于EE)给出。模拟代表在未用重组Gα转染的细胞中从内源性G蛋白获得的应答。显示用于测量Gq家族成员的活化的动态窗口取决于Gα亚基的表达水平。如关于Gq的图6B和关于G14的图6D所示，随着更多的Gα被共转染，动态窗口会提高。对于G15(图6E)，该作用在5ng的编码Gα15的共转染的构建体的情况下已经达到最大，并且更多转染没有明显的作用。对于G11(在图6C中)，表明转染的DNA水平增加超过5ng会导致动态窗的降低。对于监测EE处的G蛋白活性获得了相似的结果(图6F-I)。
使用优化的条件对在PM(图6J-6M)和EE(图6N-6Q)处的TPαR配体U46619、I-BOP、CTA2、U51605、I-SAP和SQ 29,558的特性进行分析。如图6J-6M所示，U46619、I-BOP和CTA2是对Gq(图6J)、G14(图6K)、G11(图6L)和G15(图6M)的完全激动剂；U51605是部分激动剂(对于Gq的I-BOP最大应答的32％，对于G14为25％，对于G11为24％，以及对于G15为10％)；I-SAP(对于Gq仅为7％)并且SQ 29,558无法在PM处诱导所有四种G蛋白的显著活化。当监测EE处的G蛋白活性时，获得相似的结果(效力和功效)(图6N-6Q)。关于U51605，在EE处观察到对Gq(I-BOP最大应答的71％)、G11(61％)和G14(75％)更高的功效，但对G15没有应答。最后，获得在PM处对于Gq和G11活化分别评价为0.840和0.879的Z'因子(图6R和6S)，再次表明该测定法是鲁棒的并且与基于Gq调节的高通量筛选应用兼容。
在图7A-7F中描述了用于监测Gq家族(Gq、G11、G14和G15)的G蛋白活化的基于GRK2的BRET系统的优化和使用。提出了两种RGS(GRK2)构建体。用BRET供体RlucII在N末端(RlucII-RGS(GRK2))或C末端(RGS(GRK2)-RlucII)标记包括Gq结合结构域(RGS结构域)(残基30-203,RGS(GRK2))的片段。全长GRK2蛋白具有调节其向PM的募集的普列克底物蛋白(PH)结构域和Gβγ相互作用结构域。仅使用GRK2的与Gq结合RGS结构域有利地允许在Gq蛋白质活化时胞浆定位和易位到不同的区室，而不受PIP2或游离Gβγ亚基水平的影响，其可以例如通过其他非Gq G蛋白质活化来调节。在图7B和7C中，呈现了RlucII-RGS(GRK2)获得的通过两个Gq偶联受体(AT1AR，被血管紧张素II刺激(图7B)以及TPαR，被U46619刺激(图7C))在PM处Gq、G11、G14和G15活化的剂量-应答曲线。如图7D和7E所示，尽管动态窗口总体上相对于RlucII-RGS(GRK2)更低，但RGS(GRK2)-RlucII获得了类似的结果(偶联和EC50)。在100nM U46619情况下，TPαR活化后评估RlucII-RGS(GRK2)到PM的募集时，获得的Z'因子评价为0.785，这证实了该测定法是鲁棒的，并且与基于Gq调节的高通量筛选应用兼容。
实施例6：用于监测通过非受体鸟嘌呤核苷酸交换因子(GEF)的G蛋白活化的系统和测定法。
G蛋白活化可以通过非受体鸟嘌呤核苷酸交换因子(GEF)诸如GIV(与Gα相互作用囊泡相关蛋白，也称为Girdin)、NUCB1(核连蛋白1，也称为calnuc)、NUCB2和DAPLE(散乱相关蛋白)实现。GIV/Girdin活性与Gi活性的RTK(例如EGFR)和整合素调节有关。DAPLE通过wnt/Frizzle受体(GPCR)与rac和Gi活化相关。可以使用用于监测本文所述的Gi家族的G蛋白活化的系统/测定法(诸如使用Rap1Gap(SSS-AAA)-RlucII的系统/测定法)来监测GEF介导的WT Gi蛋白的活化。但是，由于已显示GPCR可反式活化RTK(例如EGFR和IGFR)，因此对于HTS应用和配体特性分析研究，能够区分GPCR和GEF介导的G蛋白活化可能是有用的。为了实现该目的，设计和测试了一组具有C末端突变的突变体Gαi2蛋白(图8A)。在剂量应答曲线中，将EGFR介导的WT和突变体Gi2的活化与GPCR(BKB2R)应答进行了比较。缺失Gαi2的最后2个残基(图8B、8C)或Gαi2(图8D、8E)和GαoB(图8F、8G)的最后5个残基足以在维持EGFR介导的Gi2和GoB活化的同时防止GPCR介导的G蛋白活化。对于Gi2，与WT G蛋白相比，对于共表达EGFR的细胞，两种缺失均改变了基础活性。在2个保守的亮氨酸残基(位置-7和位置-2)进行替换，以确定导致与EGFR类似的基础活性，而突变体对GPCR活化仍然不活跃的残基。L-7G突变体示出与具有EGFR的WT不同的基础活性(图8H、8I)。Gαi2的L-2G突变体的结果显示与具有EGFR的WT相似的基础活性，但与BKB2R具有不同的基础活性(图87J、8K)。对于GαoB L-2G突变体，对两种受体的基础活性类似于WT蛋白(图8L、8M)。测试了其他L-2突变体Gαi2蛋白(L-2D、L-2P和L-2R)。尽管如激动剂诱导的Rap1GAP(SSS-AAA)-RlucII易位到PM来测量的它们的动态窗口比WT蛋白的动态窗口小，但相对于Gαi2蛋白的缺失或L-2G突变体而言，它们的基础活性差异非常接近(图8N-8S)。获得了在WT Gαi2情况下评价的Z'因子为0.70(图8T)以及在L-2P Gαi2突变体的情况下为0.73(图8U)，这表明使用突变的和WT G蛋白二者监测非GPCR介导的G蛋白活化的测定法都是鲁棒的。图8V-8X示出Rap1GAP(SSS-AAA)-RlucII可用于监测GPCR(图8V SNC80介导的DOR活化，以及图8W喹吡罗介导的D2R活化)以及GEF介导的G蛋白活化(通过EGFR刺激；图8X)，而基于RGS(RGS17)-RlucII的传感器仅监测GPCR介导的活化，因为在浓度逐渐升高的激动剂EGF存在下，BRET信号没有显著增加。由于GPCR活化可导致RTK反式活化，因此这些结果提供了证据，证明WT Gα/RGS(RGS17)-RlucII和突变体Gα/Rap1GAP(SSS-AAA)-RlucII可用于区分GPCR介导的G蛋白活化与RTG/GEF介导的G蛋白活化。
尽管以上已经通过本发明的特定实施方式描述了本发明，但是在不脱离如所附权利要求所限定的本发明的精神和实质的情况下，可以对其进行修改。在权利要求中，词语“包括”用作开放式术语，基本上等同于短语“包括但不限于”。除非上下文另外明确指出，否则单数形式“一个”、“一种”和“该”包括相应的复数形式。
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<110> 蒙特利尔大学（UNIVERSITÉ DE MONTRÉAL）
多曼治疗学公司（DOMAIN THERAPEUTICS）
<120> 评估G蛋白活化的系统和方法
<130> PN20001989P
<150> US 62/573,853
<151> 2017-10-18
<160> 63
<170> PatentIn version 3.5
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225 230 235 240
Gly Gly Leu Asp Val Thr His Gly Gln Thr Gly Thr Glu Ser Val Tyr
245 250 255
Cys Asn Phe Arg Asn Lys Glu Ile Met Phe His Val Ser Thr Lys Leu
260 265 270
Pro Tyr Thr Glu Gly Asp Ala Gln Gln Leu Gln Arg Lys Arg His Ile
275 280 285
Gly Ala Ala Ile Val Ala Val Val Phe Gln Asp Glu Asn Thr Pro Phe
290 295 300
Val Pro Asp Met Ile Ala Ser Asn Phe Leu His Ala Tyr Val Val Val
305 310 315 320
Gln Ala Glu Gly Gly Gly Pro Asp Gly Pro Leu Tyr Lys Val Ser Val
325 330 335
Thr Ala Arg Asp Asp Val Pro Phe Phe Gly Pro Pro Leu Pro Asp Pro
340 345 350
Ala Val Phe Arg Lys Gly Pro Glu Phe Gln Glu Phe Leu Leu Thr Lys
355 360 365
Leu Ile Asn Ala Glu Tyr Ala Cys Tyr Lys Ala Glu Lys Phe Ala Lys
370 375 380
Leu Glu Glu Arg Thr Arg Ala Ala Leu Leu Glu Thr Leu Tyr Glu Glu
385 390 395 400
Leu His Ile His Ser Gln Ser Met Met Gly Leu Gly Gly Asp Glu Asp
405 410 415
Lys Met Glu Asn Gly Ser Gly Gly Gly Gly Phe Phe Glu Ser Phe Lys
420 425 430
Arg Val Ile Arg
435
<210> 10
<211> 420
<212> PRT
<213> 智人
<400> 10
Met Ile Glu Lys Met Gln Gly Ser Arg Met Asp Glu Gln Arg Cys Ser
1 5 10 15
Phe Pro Pro Pro Leu Lys Thr Glu Glu Asp Tyr Ile Pro Tyr Pro Ser
20 25 30
Val His Glu Val Leu Gly Arg Glu Gly Pro Phe Pro Leu Ile Leu Leu
35 40 45
Pro Gln Phe Gly Gly Tyr Trp Ile Glu Gly Thr Asn His Glu Ile Thr
50 55 60
Ser Ile Pro Glu Thr Glu Pro Leu Gln Ser Pro Thr Thr Lys Val Lys
65 70 75 80
Leu Glu Cys Asn Pro Thr Ala Arg Ile Tyr Arg Lys His Phe Leu Gly
85 90 95
Lys Glu His Phe Asn Tyr Tyr Ser Leu Asp Ala Ala Leu Gly His Leu
100 105 110
Val Phe Ser Leu Lys Tyr Asp Val Ile Gly Asp Gln Glu His Leu Arg
115 120 125
Leu Leu Leu Arg Thr Lys Cys Arg Thr Tyr His Asp Val Ile Pro Ile
130 135 140
Ser Cys Leu Thr Glu Phe Pro Asn Val Val Gln Met Ala Lys Leu Val
145 150 155 160
Cys Glu Asp Val Asn Val Asp Arg Phe Tyr Pro Val Leu Tyr Pro Lys
165 170 175
Ala Ser Arg Leu Ile Val Thr Phe Asp Glu His Val Ile Ser Asn Asn
180 185 190
Phe Lys Phe Gly Val Ile Tyr Gln Lys Leu Gly Gln Thr Ser Glu Glu
195 200 205
Glu Leu Phe Ser Thr Asn Glu Glu Ser Pro Ala Phe Val Glu Phe Leu
210 215 220
Glu Phe Leu Gly Gln Lys Val Lys Leu Gln Asp Phe Lys Gly Phe Arg
225 230 235 240
Gly Gly Leu Asp Val Thr His Gly Gln Thr Gly Thr Glu Ser Val Tyr
245 250 255
Cys Asn Phe Arg Asn Lys Glu Ile Met Phe His Val Ser Thr Lys Leu
260 265 270
Pro Tyr Thr Glu Gly Asp Ala Gln Gln Leu Gln Arg Lys Arg His Ile
275 280 285
Gly Ala Ala Ile Val Ala Val Val Phe Gln Asp Glu Asn Thr Pro Phe
290 295 300
Val Pro Asp Met Ile Ala Ser Asn Phe Leu His Ala Tyr Val Val Val
305 310 315 320
Gln Ala Glu Gly Gly Gly Pro Asp Gly Pro Leu Tyr Lys Val Ser Val
325 330 335
Thr Ala Arg Asp Asp Val Pro Phe Phe Gly Pro Pro Leu Pro Asp Pro
340 345 350
Ala Val Phe Arg Lys Gly Pro Glu Phe Gln Glu Phe Leu Leu Thr Lys
355 360 365
Leu Ile Asn Ala Glu Tyr Ala Cys Tyr Lys Ala Glu Lys Phe Ala Lys
370 375 380
Leu Glu Glu Arg Thr Arg Ala Ala Leu Leu Glu Thr Leu Tyr Glu Glu
385 390 395 400
Leu His Ile His Ser Gln Ser Met Met Gly Leu Gly Gly Asp Glu Asp
405 410 415
Lys Met Glu Asn
420
<210> 11
<211> 442
<212> PRT
<213> 智人
<400> 11
Met Ile Glu Lys Met Gln Gly Ser Arg Met Asp Glu Gln Arg Cys Ser
1 5 10 15
Phe Pro Pro Pro Leu Lys Thr Glu Glu Asp Tyr Ile Pro Tyr Pro Ser
20 25 30
Val His Glu Val Leu Gly Arg Glu Gly Pro Phe Pro Leu Ile Leu Leu
35 40 45
Pro Gln Phe Gly Gly Tyr Trp Ile Glu Gly Thr Asn His Glu Ile Thr
50 55 60
Ser Ile Pro Glu Thr Glu Pro Leu Gln Ser Pro Thr Thr Lys Val Lys
65 70 75 80
Leu Glu Cys Asn Pro Thr Ala Arg Ile Tyr Arg Lys His Phe Leu Gly
85 90 95
Lys Glu His Phe Asn Tyr Tyr Ser Leu Asp Ala Ala Leu Gly His Leu
100 105 110
Val Phe Ser Leu Lys Tyr Asp Val Ile Gly Asp Gln Glu His Leu Arg
115 120 125
Leu Leu Leu Arg Thr Lys Cys Arg Thr Tyr His Asp Val Ile Pro Ile
130 135 140
Ser Cys Leu Thr Glu Phe Pro Asn Val Val Gln Met Ala Lys Leu Val
145 150 155 160
Cys Glu Asp Val Asn Val Asp Arg Phe Tyr Pro Val Leu Tyr Pro Lys
165 170 175
Ala Ser Arg Leu Ile Val Thr Phe Asp Glu His Val Ile Ser Asn Asn
180 185 190
Phe Lys Phe Gly Val Ile Tyr Gln Lys Leu Gly Gln Thr Ser Glu Glu
195 200 205
Glu Leu Phe Ser Thr Asn Glu Glu Ser Pro Ala Phe Val Glu Phe Leu
210 215 220
Glu Phe Leu Gly Gln Lys Val Lys Leu Gln Asp Phe Lys Gly Phe Arg
225 230 235 240
Gly Gly Leu Asp Val Thr His Gly Gln Thr Gly Thr Glu Ser Val Tyr
245 250 255
Cys Asn Phe Arg Asn Lys Glu Ile Met Phe His Val Ser Thr Lys Leu
260 265 270
Pro Tyr Thr Glu Gly Asp Ala Gln Gln Leu Gln Arg Lys Arg His Ile
275 280 285
Gly Ala Ala Ile Val Ala Val Val Phe Gln Asp Glu Asn Thr Pro Phe
290 295 300
Val Pro Asp Met Ile Ala Ser Asn Phe Leu His Ala Tyr Val Val Val
305 310 315 320
Gln Ala Glu Gly Gly Gly Pro Asp Gly Pro Leu Tyr Lys Val Ser Val
325 330 335
Thr Ala Arg Asp Asp Val Pro Phe Phe Gly Pro Pro Leu Pro Asp Pro
340 345 350
Ala Val Phe Arg Lys Gly Pro Glu Phe Gln Glu Phe Leu Leu Thr Lys
355 360 365
Leu Ile Asn Ala Glu Tyr Ala Cys Tyr Lys Ala Glu Lys Phe Ala Lys
370 375 380
Leu Glu Glu Arg Thr Arg Ala Ala Leu Leu Glu Thr Leu Tyr Glu Glu
385 390 395 400
Leu His Ile His Ser Gln Ser Met Met Gly Leu Gly Gly Asp Glu Asp
405 410 415
Lys Met Glu Asn Gly Ser Gly Gly Gly Gly Phe Phe Glu Ser Phe Lys
420 425 430
Arg Val Ile Arg Ala Arg Ala Gln Ala Met
435 440
<210> 12
<211> 442
<212> PRT
<213> 智人
<400> 12
Met Ile Glu Lys Met Gln Gly Ser Arg Met Asp Glu Gln Arg Cys Ser
1 5 10 15
Phe Pro Pro Pro Leu Lys Thr Glu Glu Asp Tyr Ile Pro Tyr Pro Ser
20 25 30
Val His Glu Val Leu Gly Arg Glu Gly Pro Phe Pro Leu Ile Leu Leu
35 40 45
Pro Gln Phe Gly Gly Tyr Trp Ile Glu Gly Thr Asn His Glu Ile Thr
50 55 60
Ser Ile Pro Glu Thr Glu Pro Leu Gln Ser Pro Thr Thr Lys Val Lys
65 70 75 80
Leu Glu Cys Asn Pro Thr Ala Arg Ile Tyr Arg Lys His Phe Leu Gly
85 90 95
Lys Glu His Phe Asn Tyr Tyr Ser Leu Asp Ala Ala Leu Gly His Leu
100 105 110
Val Phe Ser Leu Lys Tyr Asp Val Ile Gly Asp Gln Glu His Leu Arg
115 120 125
Leu Leu Leu Arg Thr Lys Cys Arg Thr Tyr His Asp Val Ile Pro Ile
130 135 140
Ser Cys Leu Thr Glu Phe Pro Asn Val Val Gln Met Ala Lys Leu Val
145 150 155 160
Cys Glu Asp Val Asn Val Asp Arg Phe Tyr Pro Val Leu Tyr Pro Lys
165 170 175
Ala Ser Arg Leu Ile Val Thr Phe Asp Glu His Val Ile Ser Asn Asn
180 185 190
Phe Lys Phe Gly Val Ile Tyr Gln Lys Leu Gly Gln Thr Ser Glu Glu
195 200 205
Glu Leu Phe Ser Thr Asn Glu Glu Ser Pro Ala Phe Val Glu Phe Leu
210 215 220
Glu Phe Leu Gly Gln Lys Val Lys Leu Gln Asp Phe Lys Gly Phe Arg
225 230 235 240
Gly Gly Leu Asp Val Thr His Gly Gln Thr Gly Thr Glu Ser Val Tyr
245 250 255
Cys Asn Phe Arg Asn Lys Glu Ile Met Phe His Val Ser Thr Lys Leu
260 265 270
Pro Tyr Thr Glu Gly Asp Ala Gln Gln Leu Gln Arg Lys Arg His Ile
275 280 285
Gly Ala Ala Ile Val Ala Val Val Phe Gln Asp Glu Asn Thr Pro Phe
290 295 300
Val Pro Asp Met Ile Ala Ser Asn Phe Leu His Ala Tyr Val Val Val
305 310 315 320
Gln Ala Glu Gly Gly Gly Pro Asp Gly Pro Leu Tyr Lys Val Ser Val
325 330 335
Thr Ala Arg Asp Asp Val Pro Phe Phe Gly Pro Pro Leu Pro Asp Pro
340 345 350
Ala Val Phe Arg Lys Gly Pro Glu Phe Gln Glu Phe Leu Leu Thr Lys
355 360 365
Leu Ile Asn Ala Glu Tyr Ala Cys Tyr Lys Ala Glu Lys Phe Ala Lys
370 375 380
Leu Glu Glu Arg Thr Arg Ala Ala Leu Leu Glu Thr Leu Tyr Glu Glu
385 390 395 400
Leu His Ile His Ser Gln Ser Met Met Gly Leu Gly Gly Asp Glu Asp
405 410 415
Lys Met Glu Asn Gly Ser Gly Gly Gly Gly Phe Phe Glu Ser Phe Lys
420 425 430
Arg Val Ile Arg Thr Arg Thr Gln Thr Met
435 440
<210> 13
<211> 442
<212> PRT
<213> 智人
<400> 13
Met Ile Glu Lys Met Gln Gly Ser Arg Met Asp Glu Gln Arg Cys Ser
1 5 10 15
Phe Pro Pro Pro Leu Lys Thr Glu Glu Asp Tyr Ile Pro Tyr Pro Ser
20 25 30
Val His Glu Val Leu Gly Arg Glu Gly Pro Phe Pro Leu Ile Leu Leu
35 40 45
Pro Gln Phe Gly Gly Tyr Trp Ile Glu Gly Thr Asn His Glu Ile Thr
50 55 60
Ser Ile Pro Glu Thr Glu Pro Leu Gln Ser Pro Thr Thr Lys Val Lys
65 70 75 80
Leu Glu Cys Asn Pro Thr Ala Arg Ile Tyr Arg Lys His Phe Leu Gly
85 90 95
Lys Glu His Phe Asn Tyr Tyr Ser Leu Asp Ala Ala Leu Gly His Leu
100 105 110
Val Phe Ser Leu Lys Tyr Asp Val Ile Gly Asp Gln Glu His Leu Arg
115 120 125
Leu Leu Leu Arg Thr Lys Cys Arg Thr Tyr His Asp Val Ile Pro Ile
130 135 140
Ser Cys Leu Thr Glu Phe Pro Asn Val Val Gln Met Ala Lys Leu Val
145 150 155 160
Cys Glu Asp Val Asn Val Asp Arg Phe Tyr Pro Val Leu Tyr Pro Lys
165 170 175
Ala Ser Arg Leu Ile Val Thr Phe Asp Glu His Val Ile Ser Asn Asn
180 185 190
Phe Lys Phe Gly Val Ile Tyr Gln Lys Leu Gly Gln Thr Ser Glu Glu
195 200 205
Glu Leu Phe Ser Thr Asn Glu Glu Ser Pro Ala Phe Val Glu Phe Leu
210 215 220
Glu Phe Leu Gly Gln Lys Val Lys Leu Gln Asp Phe Lys Gly Phe Arg
225 230 235 240
Gly Gly Leu Asp Val Thr His Gly Gln Thr Gly Thr Glu Ser Val Tyr
245 250 255
Cys Asn Phe Arg Asn Lys Glu Ile Met Phe His Val Ser Thr Lys Leu
260 265 270
Pro Tyr Thr Glu Gly Asp Ala Gln Gln Leu Gln Arg Lys Arg His Ile
275 280 285
Gly Ala Ala Ile Val Ala Val Val Phe Gln Asp Glu Asn Thr Pro Phe
290 295 300
Val Pro Asp Met Ile Ala Ser Asn Phe Leu His Ala Tyr Val Val Val
305 310 315 320
Gln Ala Glu Gly Gly Gly Pro Asp Gly Pro Leu Tyr Lys Val Ser Val
325 330 335
Thr Ala Arg Asp Asp Val Pro Phe Phe Gly Pro Pro Leu Pro Asp Pro
340 345 350
Ala Val Phe Arg Lys Gly Pro Glu Phe Gln Glu Phe Leu Leu Thr Lys
355 360 365
Leu Ile Asn Ala Glu Tyr Ala Cys Tyr Lys Ala Glu Lys Phe Ala Lys
370 375 380
Leu Glu Glu Arg Thr Arg Ala Ala Leu Leu Glu Thr Leu Tyr Glu Glu
385 390 395 400
Leu His Ile His Ser Gln Ser Met Met Gly Leu Gly Gly Asp Glu Asp
405 410 415
Lys Met Glu Asn Gly Ser Gly Gly Gly Gly Phe Phe Glu Ser Phe Lys
420 425 430
Arg Val Ile Arg Ala Arg Ser Gln Ala Met
435 440
<210> 14
<211> 442
<212> PRT
<213> 智人
<400> 14
Met Ile Glu Lys Met Gln Gly Ser Arg Met Asp Glu Gln Arg Cys Ser
1 5 10 15
Phe Pro Pro Pro Leu Lys Thr Glu Glu Asp Tyr Ile Pro Tyr Pro Ser
20 25 30
Val His Glu Val Leu Gly Arg Glu Gly Pro Phe Pro Leu Ile Leu Leu
35 40 45
Pro Gln Phe Gly Gly Tyr Trp Ile Glu Gly Thr Asn His Glu Ile Thr
50 55 60
Ser Ile Pro Glu Thr Glu Pro Leu Gln Ser Pro Thr Thr Lys Val Lys
65 70 75 80
Leu Glu Cys Asn Pro Thr Ala Arg Ile Tyr Arg Lys His Phe Leu Gly
85 90 95
Lys Glu His Phe Asn Tyr Tyr Ser Leu Asp Ala Ala Leu Gly His Leu
100 105 110
Val Phe Ser Leu Lys Tyr Asp Val Ile Gly Asp Gln Glu His Leu Arg
115 120 125
Leu Leu Leu Arg Thr Lys Cys Arg Thr Tyr His Asp Val Ile Pro Ile
130 135 140
Ser Cys Leu Thr Glu Phe Pro Asn Val Val Gln Met Ala Lys Leu Val
145 150 155 160
Cys Glu Asp Val Asn Val Asp Arg Phe Tyr Pro Val Leu Tyr Pro Lys
165 170 175
Ala Ser Arg Leu Ile Val Thr Phe Asp Glu His Val Ile Ser Asn Asn
180 185 190
Phe Lys Phe Gly Val Ile Tyr Gln Lys Leu Gly Gln Thr Ser Glu Glu
195 200 205
Glu Leu Phe Ser Thr Asn Glu Glu Ser Pro Ala Phe Val Glu Phe Leu
210 215 220
Glu Phe Leu Gly Gln Lys Val Lys Leu Gln Asp Phe Lys Gly Phe Arg
225 230 235 240
Gly Gly Leu Asp Val Thr His Gly Gln Thr Gly Thr Glu Ser Val Tyr
245 250 255
Cys Asn Phe Arg Asn Lys Glu Ile Met Phe His Val Ser Thr Lys Leu
260 265 270
Pro Tyr Thr Glu Gly Asp Ala Gln Gln Leu Gln Arg Lys Arg His Ile
275 280 285
Gly Ala Ala Ile Val Ala Val Val Phe Gln Asp Glu Asn Thr Pro Phe
290 295 300
Val Pro Asp Met Ile Ala Ser Asn Phe Leu His Ala Tyr Val Val Val
305 310 315 320
Gln Ala Glu Gly Gly Gly Pro Asp Gly Pro Leu Tyr Lys Val Ser Val
325 330 335
Thr Ala Arg Asp Asp Val Pro Phe Phe Gly Pro Pro Leu Pro Asp Pro
340 345 350
Ala Val Phe Arg Lys Gly Pro Glu Phe Gln Glu Phe Leu Leu Thr Lys
355 360 365
Leu Ile Asn Ala Glu Tyr Ala Cys Tyr Lys Ala Glu Lys Phe Ala Lys
370 375 380
Leu Glu Glu Arg Thr Arg Ala Ala Leu Leu Glu Thr Leu Tyr Glu Glu
385 390 395 400
Leu His Ile His Ser Gln Ser Met Met Gly Leu Gly Gly Asp Glu Asp
405 410 415
Lys Met Glu Asn Gly Ser Gly Gly Gly Gly Phe Phe Glu Ser Phe Lys
420 425 430
Arg Val Ile Arg Asp Arg Ser Gln Ala Met
435 440
<210> 15
<211> 442
<212> PRT
<213> 智人
<400> 15
Met Ile Glu Lys Met Gln Gly Ser Arg Met Asp Glu Gln Arg Cys Ser
1 5 10 15
Phe Pro Pro Pro Leu Lys Thr Glu Glu Asp Tyr Ile Pro Tyr Pro Ser
20 25 30
Val His Glu Val Leu Gly Arg Glu Gly Pro Phe Pro Leu Ile Leu Leu
35 40 45
Pro Gln Phe Gly Gly Tyr Trp Ile Glu Gly Thr Asn His Glu Ile Thr
50 55 60
Ser Ile Pro Glu Thr Glu Pro Leu Gln Ser Pro Thr Thr Lys Val Lys
65 70 75 80
Leu Glu Cys Asn Pro Thr Ala Arg Ile Tyr Arg Lys His Phe Leu Gly
85 90 95
Lys Glu His Phe Asn Tyr Tyr Ser Leu Asp Ala Ala Leu Gly His Leu
100 105 110
Val Phe Ser Leu Lys Tyr Asp Val Ile Gly Asp Gln Glu His Leu Arg
115 120 125
Leu Leu Leu Arg Thr Lys Cys Arg Thr Tyr His Asp Val Ile Pro Ile
130 135 140
Ser Cys Leu Thr Glu Phe Pro Asn Val Val Gln Met Ala Lys Leu Val
145 150 155 160
Cys Glu Asp Val Asn Val Asp Arg Phe Tyr Pro Val Leu Tyr Pro Lys
165 170 175
Ala Ser Arg Leu Ile Val Thr Phe Asp Glu His Val Ile Ser Asn Asn
180 185 190
Phe Lys Phe Gly Val Ile Tyr Gln Lys Leu Gly Gln Thr Ser Glu Glu
195 200 205
Glu Leu Phe Ser Thr Asn Glu Glu Ser Pro Ala Phe Val Glu Phe Leu
210 215 220
Glu Phe Leu Gly Gln Lys Val Lys Leu Gln Asp Phe Lys Gly Phe Arg
225 230 235 240
Gly Gly Leu Asp Val Thr His Gly Gln Thr Gly Thr Glu Ser Val Tyr
245 250 255
Cys Asn Phe Arg Asn Lys Glu Ile Met Phe His Val Ser Thr Lys Leu
260 265 270
Pro Tyr Thr Glu Gly Asp Ala Gln Gln Leu Gln Arg Lys Arg His Ile
275 280 285
Gly Ala Ala Ile Val Ala Val Val Phe Gln Asp Glu Asn Thr Pro Phe
290 295 300
Val Pro Asp Met Ile Ala Ser Asn Phe Leu His Ala Tyr Val Val Val
305 310 315 320
Gln Ala Glu Gly Gly Gly Pro Asp Gly Pro Leu Tyr Lys Val Ser Val
325 330 335
Thr Ala Arg Asp Asp Val Pro Phe Phe Gly Pro Pro Leu Pro Asp Pro
340 345 350
Ala Val Phe Arg Lys Gly Pro Glu Phe Gln Glu Phe Leu Leu Thr Lys
355 360 365
Leu Ile Asn Ala Glu Tyr Ala Cys Tyr Lys Ala Glu Lys Phe Ala Lys
370 375 380
Leu Glu Glu Arg Thr Arg Ala Ala Leu Leu Glu Thr Leu Tyr Glu Glu
385 390 395 400
Leu His Ile His Ser Gln Ser Met Met Gly Leu Gly Gly Asp Glu Asp
405 410 415
Lys Met Glu Asn Gly Ser Gly Gly Gly Gly Phe Phe Glu Ser Phe Lys
420 425 430
Arg Val Ile Arg Ala Arg Ser Gln Asp Met
435 440
<210> 16
<211> 442
<212> PRT
<213> 智人
<400> 16
Met Ile Glu Lys Met Gln Gly Ser Arg Met Asp Glu Gln Arg Cys Ser
1 5 10 15
Phe Pro Pro Pro Leu Lys Thr Glu Glu Asp Tyr Ile Pro Tyr Pro Ser
20 25 30
Val His Glu Val Leu Gly Arg Glu Gly Pro Phe Pro Leu Ile Leu Leu
35 40 45
Pro Gln Phe Gly Gly Tyr Trp Ile Glu Gly Thr Asn His Glu Ile Thr
50 55 60
Ser Ile Pro Glu Thr Glu Pro Leu Gln Ser Pro Thr Thr Lys Val Lys
65 70 75 80
Leu Glu Cys Asn Pro Thr Ala Arg Ile Tyr Arg Lys His Phe Leu Gly
85 90 95
Lys Glu His Phe Asn Tyr Tyr Ser Leu Asp Ala Ala Leu Gly His Leu
100 105 110
Val Phe Ser Leu Lys Tyr Asp Val Ile Gly Asp Gln Glu His Leu Arg
115 120 125
Leu Leu Leu Arg Thr Lys Cys Arg Thr Tyr His Asp Val Ile Pro Ile
130 135 140
Ser Cys Leu Thr Glu Phe Pro Asn Val Val Gln Met Ala Lys Leu Val
145 150 155 160
Cys Glu Asp Val Asn Val Asp Arg Phe Tyr Pro Val Leu Tyr Pro Lys
165 170 175
Ala Ser Arg Leu Ile Val Thr Phe Asp Glu His Val Ile Ser Asn Asn
180 185 190
Phe Lys Phe Gly Val Ile Tyr Gln Lys Leu Gly Gln Thr Ser Glu Glu
195 200 205
Glu Leu Phe Ser Thr Asn Glu Glu Ser Pro Ala Phe Val Glu Phe Leu
210 215 220
Glu Phe Leu Gly Gln Lys Val Lys Leu Gln Asp Phe Lys Gly Phe Arg
225 230 235 240
Gly Gly Leu Asp Val Thr His Gly Gln Thr Gly Thr Glu Ser Val Tyr
245 250 255
Cys Asn Phe Arg Asn Lys Glu Ile Met Phe His Val Ser Thr Lys Leu
260 265 270
Pro Tyr Thr Glu Gly Asp Ala Gln Gln Leu Gln Arg Lys Arg His Ile
275 280 285
Gly Ala Ala Ile Val Ala Val Val Phe Gln Asp Glu Asn Thr Pro Phe
290 295 300
Val Pro Asp Met Ile Ala Ser Asn Phe Leu His Ala Tyr Val Val Val
305 310 315 320
Gln Ala Glu Gly Gly Gly Pro Asp Gly Pro Leu Tyr Lys Val Ser Val
325 330 335
Thr Ala Arg Asp Asp Val Pro Phe Phe Gly Pro Pro Leu Pro Asp Pro
340 345 350
Ala Val Phe Arg Lys Gly Pro Glu Phe Gln Glu Phe Leu Leu Thr Lys
355 360 365
Leu Ile Asn Ala Glu Tyr Ala Cys Tyr Lys Ala Glu Lys Phe Ala Lys
370 375 380
Leu Glu Glu Arg Thr Arg Ala Ala Leu Leu Glu Thr Leu Tyr Glu Glu
385 390 395 400
Leu His Ile His Ser Gln Ser Met Met Gly Leu Gly Gly Asp Glu Asp
405 410 415
Lys Met Glu Asn Gly Ser Gly Gly Gly Gly Phe Phe Glu Ser Phe Lys
420 425 430
Arg Val Ile Arg Asp Arg Ser Gln Asp Met
435 440
<210> 17
<211> 210
<212> PRT
<213> 智人
<400> 17
Met Arg Lys Arg Gln Gln Ser Gln Asn Glu Gly Thr Pro Ala Val Ser
1 5 10 15
Gln Ala Pro Gly Asn Gln Arg Pro Asn Asn Thr Cys Cys Phe Cys Trp
20 25 30
Cys Cys Cys Cys Ser Cys Ser Cys Leu Thr Val Arg Asn Glu Glu Arg
35 40 45
Gly Glu Asn Ala Gly Arg Pro Thr His Thr Thr Lys Met Glu Ser Ile
50 55 60
Gln Val Leu Glu Glu Cys Gln Asn Pro Thr Ala Glu Glu Val Leu Ser
65 70 75 80
Trp Ser Gln Asn Phe Asp Lys Met Met Lys Ala Pro Ala Gly Arg Asn
85 90 95
Leu Phe Arg Glu Phe Leu Arg Thr Glu Tyr Ser Glu Glu Asn Leu Leu
100 105 110
Phe Trp Leu Ala Cys Glu Asp Leu Lys Lys Glu Gln Asn Lys Lys Val
115 120 125
Ile Glu Glu Lys Ala Arg Met Ile Tyr Glu Asp Tyr Ile Ser Ile Leu
130 135 140
Ser Pro Lys Glu Val Ser Leu Asp Ser Arg Val Arg Glu Val Ile Asn
145 150 155 160
Arg Asn Leu Leu Asp Pro Asn Pro His Met Tyr Glu Asp Ala Gln Leu
165 170 175
Gln Ile Tyr Thr Leu Met His Arg Asp Ser Phe Pro Arg Phe Leu Asn
180 185 190
Ser Gln Ile Tyr Lys Ser Phe Val Glu Ser Thr Ala Gly Ser Ser Ser
195 200 205
Glu Ser
210
<210> 18
<211> 217
<212> PRT
<213> 智人
<400> 18
Met Pro Thr Pro His Glu Ala Glu Lys Gln Ile Thr Gly Pro Glu Glu
1 5 10 15
Ala Asp Arg Pro Pro Ser Met Ser Ser His Asp Thr Ala Ser Pro Ala
20 25 30
Ala Pro Ser Arg Asn Pro Cys Cys Leu Cys Trp Cys Cys Cys Cys Ser
35 40 45
Cys Ser Trp Asn Gln Glu Arg Arg Arg Ala Trp Gln Ala Ser Arg Glu
50 55 60
Ser Lys Leu Gln Pro Leu Pro Ser Cys Glu Val Cys Ala Thr Pro Ser
65 70 75 80
Pro Glu Glu Val Gln Ser Trp Ala Gln Ser Phe Asp Lys Leu Met His
85 90 95
Ser Pro Ala Gly Arg Ser Val Phe Arg Ala Phe Leu Arg Thr Glu Tyr
100 105 110
Ser Glu Glu Asn Met Leu Phe Trp Leu Ala Cys Glu Glu Leu Lys Ala
115 120 125
Glu Ala Asn Gln His Val Val Asp Glu Lys Ala Arg Leu Ile Tyr Glu
130 135 140
Asp Tyr Val Ser Ile Leu Ser Pro Lys Glu Val Ser Leu Asp Ser Arg
145 150 155 160
Val Arg Glu Gly Ile Asn Lys Lys Met Gln Glu Pro Ser Ala His Thr
165 170 175
Phe Asp Asp Ala Gln Leu Gln Ile Tyr Thr Leu Met His Arg Asp Ser
180 185 190
Tyr Pro Arg Phe Leu Ser Ser Pro Thr Tyr Arg Ala Leu Leu Leu Gln
195 200 205
Gly Pro Ser Gln Ser Ser Ser Glu Ala
210 215
<210> 19
<211> 388
<212> PRT
<213> 智人
<400> 19
Met Pro Gln Leu Ser Gln Asp Asn Gln Glu Cys Leu Gln Lys His Phe
1 5 10 15
Ser Arg Pro Ser Ile Trp Thr Gln Phe Leu Pro Leu Phe Arg Ala Gln
20 25 30
Arg Tyr Asn Thr Asp Ile His Gln Ile Thr Glu Asn Glu Gly Asp Leu
35 40 45
Arg Ala Val Pro Asp Ile Lys Ser Phe Pro Pro Ala Gln Leu Pro Asp
50 55 60
Ser Pro Ala Ala Pro Lys Leu Phe Gly Leu Leu Ser Ser Pro Leu Ser
65 70 75 80
Ser Leu Ala Arg Phe Phe Ser His Leu Leu Arg Arg Pro Pro Pro Glu
85 90 95
Ala Pro Arg Arg Arg Leu Asp Phe Ser Pro Leu Leu Pro Ala Leu Pro
100 105 110
Ala Ala Arg Leu Ser Arg Gly His Glu Glu Leu Pro Gly Arg Leu Ser
115 120 125
Leu Leu Leu Gly Ala Ala Leu Ala Leu Pro Gly Arg Pro Ser Gly Gly
130 135 140
Arg Pro Leu Arg Pro Pro His Pro Val Ala Lys Pro Arg Glu Glu Asp
145 150 155 160
Ala Thr Ala Gly Gln Ser Ser Pro Met Pro Gln Met Gly Ser Glu Arg
165 170 175
Met Glu Met Arg Lys Arg Gln Met Pro Ala Ala Gln Asp Thr Pro Gly
180 185 190
Ala Ala Pro Gly Gln Pro Gly Ala Gly Ser Arg Gly Ser Asn Ala Cys
195 200 205
Cys Phe Cys Trp Cys Cys Cys Cys Ser Cys Ser Cys Leu Thr Val Arg
210 215 220
Asn Gln Glu Asp Gln Arg Pro Thr Ile Ala Ser His Glu Leu Arg Ala
225 230 235 240
Asp Leu Pro Thr Trp Glu Glu Ser Pro Ala Pro Thr Leu Glu Glu Val
245 250 255
Asn Ala Trp Ala Gln Ser Phe Asp Lys Leu Met Val Thr Pro Ala Gly
260 265 270
Arg Asn Ala Phe Arg Glu Phe Leu Arg Thr Glu Phe Ser Glu Glu Asn
275 280 285
Met Leu Phe Trp Met Ala Cys Glu Glu Leu Lys Lys Glu Ala Asn Lys
290 295 300
Asn Ile Ile Glu Glu Lys Ala Arg Ile Ile Tyr Glu Asp Tyr Ile Ser
305 310 315 320
Ile Leu Ser Pro Lys Glu Val Ser Leu Asp Ser Arg Val Arg Glu Val
325 330 335
Ile Asn Arg Asn Met Val Glu Pro Ser Gln His Ile Phe Asp Asp Ala
340 345 350
Gln Leu Gln Ile Tyr Thr Leu Met His Arg Asp Ser Tyr Pro Arg Phe
355 360 365
Met Asn Ser Ala Val Tyr Lys Asp Leu Leu Gln Ser Leu Ser Glu Lys
370 375 380
Ser Ile Glu Ala
385
<210> 20
<211> 18
<212> PRT
<213> 人工序列
<220>
<223> 合成肽接头
<400> 20
Gly Ser Ala Gly Thr Gly Gly Arg Ala Ile Asp Ile Lys Leu Ala Ser
1 5 10 15
Ala Thr
<210> 21
<211> 1522
<212> PRT
<213> 智人
<400> 21
Met Ser Val Arg Leu Pro Gln Ser Ile Asp Arg Leu Ser Ser Leu Ser
1 5 10 15
Ser Leu Gly Asp Ser Ala Pro Glu Arg Lys Ser Pro Ser His His Arg
20 25 30
Gln Pro Ser Asp Ala Ser Glu Thr Thr Gly Leu Val Gln Arg Cys Val
35 40 45
Ile Ile Gln Lys Asp Gln His Gly Phe Gly Phe Thr Val Ser Gly Asp
50 55 60
Arg Ile Val Leu Val Gln Ser Val Arg Pro Gly Gly Ala Ala Met Lys
65 70 75 80
Ala Gly Val Lys Glu Gly Asp Arg Ile Ile Lys Val Asn Gly Thr Met
85 90 95
Val Thr Asn Ser Ser His Leu Glu Val Val Lys Leu Ile Lys Ser Gly
100 105 110
Ala Tyr Val Ala Leu Thr Leu Leu Gly Ser Ser Pro Ser Ser Met Gly
115 120 125
Ile Ser Gly Leu Gln Gln Asp Pro Ser Pro Ala Gly Ala Pro Arg Ile
130 135 140
Thr Ser Val Ile Pro Ser Pro Pro Pro Pro Pro Pro Leu Pro Pro Pro
145 150 155 160
Gln Arg Ile Thr Gly Pro Lys Pro Leu Gln Asp Pro Glu Val Gln Lys
165 170 175
His Ala Thr Gln Ile Leu Arg Asn Met Leu Arg Gln Glu Glu Lys Glu
180 185 190
Leu Gln Asp Ile Leu Pro Leu Tyr Gly Asp Thr Ser Gln Arg Pro Ser
195 200 205
Glu Gly Arg Leu Ser Leu Asp Ser Gln Glu Gly Asp Ser Gly Leu Asp
210 215 220
Ser Gly Thr Glu Arg Phe Pro Ser Leu Ser Glu Ser Leu Met Asn Arg
225 230 235 240
Asn Ser Val Leu Ser Asp Pro Gly Leu Asp Ser Pro Arg Thr Ser Pro
245 250 255
Val Ile Met Ala Arg Val Ala Gln His His Arg Arg Gln Gly Ser Asp
260 265 270
Ala Ala Val Pro Ser Thr Gly Asp Gln Gly Val Asp Gln Ser Pro Lys
275 280 285
Pro Leu Ile Ile Gly Pro Glu Glu Asp Tyr Asp Pro Gly Tyr Phe Asn
290 295 300
Asn Glu Ser Asp Ile Ile Phe Gln Asp Leu Glu Lys Leu Lys Ser Arg
305 310 315 320
Pro Ala His Leu Gly Val Phe Leu Arg Tyr Ile Phe Ser Gln Ala Asp
325 330 335
Pro Ser Pro Leu Leu Phe Tyr Leu Cys Ala Glu Val Tyr Gln Gln Ala
340 345 350
Ser Pro Lys Asp Ser Arg Ser Leu Gly Lys Asp Ile Trp Asn Ile Phe
355 360 365
Leu Glu Lys Asn Ala Pro Leu Arg Val Lys Ile Pro Glu Met Leu Gln
370 375 380
Ala Glu Ile Asp Ser Arg Leu Arg Asn Ser Glu Asp Ala Arg Gly Val
385 390 395 400
Leu Cys Glu Ala Gln Glu Ala Ala Met Pro Glu Ile Gln Glu Gln Ile
405 410 415
His Asp Tyr Arg Thr Lys Arg Thr Leu Gly Leu Gly Ser Leu Tyr Gly
420 425 430
Glu Asn Asp Leu Leu Asp Leu Asp Gly Asp Pro Leu Arg Glu Arg Gln
435 440 445
Val Ala Glu Lys Gln Leu Ala Ala Leu Gly Asp Ile Leu Ser Lys Tyr
450 455 460
Glu Glu Asp Arg Ser Ala Pro Met Asp Phe Ala Leu Asn Thr Tyr Met
465 470 475 480
Ser His Ala Gly Ile Arg Leu Arg Glu Ala Arg Pro Ser Asn Thr Ala
485 490 495
Glu Lys Ala Gln Ser Ala Pro Asp Lys Asp Lys Trp Leu Pro Phe Phe
500 505 510
Pro Lys Thr Lys Lys Ser Ser Asn Ser Lys Lys Glu Lys Asp Ala Leu
515 520 525
Glu Asp Lys Lys Arg Asn Pro Ile Leu Lys Tyr Ile Gly Lys Pro Lys
530 535 540
Ser Ser Ser Gln Ser Thr Phe His Ile Pro Leu Ser Pro Val Glu Val
545 550 555 560
Lys Pro Gly Asn Val Arg Asn Ile Ile Gln His Phe Glu Asn Asn Gln
565 570 575
Gln Tyr Asp Ala Pro Glu Pro Gly Thr Gln Arg Leu Ser Thr Gly Ser
580 585 590
Phe Pro Glu Asp Leu Leu Glu Ser Asp Ser Ser Arg Ser Glu Ile Arg
595 600 605
Leu Gly Arg Ser Glu Ser Leu Lys Gly Arg Glu Glu Met Lys Arg Ser
610 615 620
Arg Lys Ala Glu Asn Val Pro Arg Ser Arg Ser Asp Val Asp Met Asp
625 630 635 640
Ala Ala Ala Glu Ala Thr Arg Leu His Gln Ser Ala Ser Ser Ser Thr
645 650 655
Ser Ser Leu Ser Thr Arg Ser Leu Glu Asn Pro Thr Pro Pro Phe Thr
660 665 670
Pro Lys Met Gly Arg Arg Ser Ile Glu Ser Pro Ser Leu Gly Phe Cys
675 680 685
Thr Asp Thr Leu Leu Pro His Leu Leu Glu Asp Asp Leu Gly Gln Leu
690 695 700
Ser Asp Leu Glu Pro Glu Pro Asp Ala Gln Asn Trp Gln His Thr Val
705 710 715 720
Gly Lys Asp Val Val Ala Gly Leu Thr Gln Arg Glu Ile Asp Arg Gln
725 730 735
Glu Val Ile Asn Glu Leu Phe Val Thr Glu Ala Ser His Leu Arg Thr
740 745 750
Leu Arg Val Leu Asp Leu Ile Phe Tyr Gln Arg Met Lys Lys Glu Asn
755 760 765
Leu Met Pro Arg Glu Glu Leu Ala Arg Leu Phe Pro Asn Leu Pro Glu
770 775 780
Leu Ile Glu Ile His Asn Ser Trp Cys Glu Ala Met Lys Lys Leu Arg
785 790 795 800
Glu Glu Gly Pro Ile Ile Lys Glu Ile Ser Asp Leu Met Leu Ala Arg
805 810 815
Phe Asp Gly Pro Ala Arg Glu Glu Leu Gln Gln Val Ala Ala Gln Phe
820 825 830
Cys Ser Tyr Gln Ser Ile Ala Leu Glu Leu Ile Lys Thr Lys Gln Arg
835 840 845
Lys Glu Ser Arg Phe Gln Leu Phe Met Gln Glu Ala Glu Ser His Pro
850 855 860
Gln Cys Arg Arg Leu Gln Leu Arg Asp Leu Ile Ile Ser Glu Met Gln
865 870 875 880
Arg Leu Thr Lys Tyr Pro Leu Leu Leu Glu Ser Ile Ile Lys His Thr
885 890 895
Glu Gly Gly Thr Ser Glu His Glu Lys Leu Cys Arg Ala Arg Asp Gln
900 905 910
Cys Arg Glu Ile Leu Lys Tyr Val Asn Glu Ala Val Lys Gln Thr Glu
915 920 925
Asn Arg His Arg Leu Glu Gly Tyr Gln Lys Arg Leu Asp Ala Thr Ala
930 935 940
Leu Glu Arg Ala Ser Asn Pro Leu Ala Ala Glu Phe Lys Ser Leu Asp
945 950 955 960
Leu Thr Thr Arg Lys Met Ile His Glu Gly Pro Leu Thr Trp Arg Ile
965 970 975
Ser Lys Asp Lys Thr Leu Asp Leu His Val Leu Leu Leu Glu Asp Leu
980 985 990
Leu Val Leu Leu Gln Lys Gln Asp Glu Lys Leu Leu Leu Lys Cys His
995 1000 1005
Ser Lys Thr Ala Val Gly Ser Ser Asp Ser Lys Gln Thr Phe Ser
1010 1015 1020
Pro Val Leu Lys Leu Asn Ala Val Leu Ile Arg Ser Val Ala Thr
1025 1030 1035
Asp Lys Arg Ala Phe Phe Ile Ile Cys Thr Ser Lys Leu Gly Pro
1040 1045 1050
Pro Gln Ile Tyr Glu Leu Val Ala Leu Thr Ser Ser Asp Lys Asn
1055 1060 1065
Thr Trp Met Glu Leu Leu Glu Glu Ala Val Arg Asn Ala Thr Arg
1070 1075 1080
His Pro Gly Ala Ala Pro Met Pro Val His Pro Pro Pro Pro Gly
1085 1090 1095
Pro Arg Glu Pro Ala Gln Gln Gly Pro Thr Pro Ser Arg Val Glu
1100 1105 1110
Leu Asp Asp Ser Asp Val Phe His Gly Glu Pro Glu Pro Glu Glu
1115 1120 1125
Leu Pro Gly Gly Thr Gly Ser Gln Gln Arg Val Gln Gly Lys His
1130 1135 1140
Gln Val Leu Leu Glu Asp Pro Glu Gln Glu Gly Ser Ala Glu Glu
1145 1150 1155
Glu Glu Leu Gly Val Leu Pro Cys Pro Ser Thr Ser Leu Asp Gly
1160 1165 1170
Glu Asn Arg Gly Ile Arg Thr Arg Asn Pro Ile His Leu Ala Phe
1175 1180 1185
Pro Gly Pro Leu Phe Met Glu Gly Leu Ala Asp Ser Ala Leu Glu
1190 1195 1200
Asp Val Glu Asn Leu Arg His Leu Ile Leu Trp Ser Leu Leu Pro
1205 1210 1215
Gly His Thr Met Glu Thr Gln Ala Ala Gln Glu Pro Glu Asp Asp
1220 1225 1230
Leu Thr Pro Thr Pro Ser Val Ile Ser Val Thr Ser His Pro Trp
1235 1240 1245
Asp Pro Gly Ser Pro Gly Gln Ala Pro Pro Gly Gly Glu Gly Asp
1250 1255 1260
Asn Thr Gln Leu Ala Gly Leu Glu Gly Glu Arg Pro Glu Gln Glu
1265 1270 1275
Asp Met Gly Leu Cys Ser Leu Glu His Leu Pro Pro Arg Thr Arg
1280 1285 1290
Asn Ser Gly Ile Trp Glu Ser Pro Glu Leu Asp Arg Asn Leu Ala
1295 1300 1305
Glu Asp Ala Ser Ser Thr Glu Ala Ala Gly Gly Tyr Lys Val Val
1310 1315 1320
Arg Lys Ala Glu Val Ala Gly Ser Lys Val Val Pro Ala Leu Pro
1325 1330 1335
Glu Ser Gly Gln Ser Glu Pro Gly Pro Pro Glu Val Glu Gly Gly
1340 1345 1350
Thr Lys Ala Thr Gly Asn Cys Phe Tyr Val Ser Met Pro Ser Gly
1355 1360 1365
Pro Pro Asp Ser Ser Thr Asp His Ser Glu Ala Pro Met Ser Pro
1370 1375 1380
Pro Gln Pro Asp Ser Leu Pro Ala Gly Gln Thr Glu Pro Gln Pro
1385 1390 1395
Gln Leu Gln Gly Gly Asn Asp Asp Pro Arg Arg Pro Ser Arg Ser
1400 1405 1410
Pro Pro Ser Leu Ala Leu Arg Asp Val Gly Met Ile Phe His Thr
1415 1420 1425
Ile Glu Gln Leu Thr Leu Lys Leu Asn Arg Leu Lys Asp Met Glu
1430 1435 1440
Leu Ala His Arg Glu Leu Leu Lys Ser Leu Gly Gly Glu Ser Ser
1445 1450 1455
Gly Gly Thr Thr Pro Val Gly Ser Phe His Thr Glu Ala Ala Arg
1460 1465 1470
Trp Thr Asp Gly Ser Leu Ser Pro Pro Ala Lys Glu Pro Leu Ala
1475 1480 1485
Ser Asp Ser Arg Asn Ser His Glu Leu Gly Pro Cys Pro Glu Asp
1490 1495 1500
Gly Ser Asp Ala Pro Leu Glu Asp Ser Thr Ala Asp Ala Ala Ala
1505 1510 1515
Ser Pro Gly Pro
1520
<210> 22
<211> 13
<212> PRT
<213> 人工序列
<220>
<223> 合成肽接头
<400> 22
Gly Ile Arg Leu Arg Glu Ala Leu Lys Leu Pro Ala Thr
1 5 10
<210> 23
<211> 912
<212> PRT
<213> 智人
<400> 23
Met Glu Asp Phe Ala Arg Gly Ala Ala Ser Pro Gly Pro Ser Arg Pro
1 5 10 15
Gly Leu Val Pro Val Ser Ile Ile Gly Ala Glu Asp Glu Asp Phe Glu
20 25 30
Asn Glu Leu Glu Thr Asn Ser Glu Glu Gln Asn Ser Gln Phe Gln Ser
35 40 45
Leu Glu Gln Val Lys Arg Arg Pro Ala His Leu Met Ala Leu Leu Gln
50 55 60
His Val Ala Leu Gln Phe Glu Pro Gly Pro Leu Leu Cys Cys Leu His
65 70 75 80
Ala Asp Met Leu Gly Ser Leu Gly Pro Lys Glu Ala Lys Lys Ala Phe
85 90 95
Leu Asp Phe Tyr His Ser Phe Leu Glu Lys Thr Ala Val Leu Arg Val
100 105 110
Pro Val Pro Pro Asn Val Ala Phe Glu Leu Asp Arg Thr Arg Ala Asp
115 120 125
Leu Ile Ser Glu Asp Val Gln Arg Arg Phe Val Gln Glu Val Val Gln
130 135 140
Ser Gln Gln Val Ala Val Gly Arg Gln Leu Glu Asp Phe Arg Ser Lys
145 150 155 160
Arg Leu Met Gly Met Thr Pro Trp Glu Gln Glu Leu Ala Gln Leu Glu
165 170 175
Ala Trp Val Gly Arg Asp Arg Ala Ser Tyr Glu Ala Arg Glu Arg His
180 185 190
Val Ala Glu Arg Leu Leu Met His Leu Glu Glu Met Gln His Thr Ile
195 200 205
Ser Thr Asp Glu Glu Lys Ser Ala Ala Val Val Asn Ala Ile Gly Leu
210 215 220
Tyr Met Arg His Leu Gly Val Arg Thr Lys Ser Gly Asp Lys Lys Ser
225 230 235 240
Gly Arg Asn Phe Phe Arg Lys Lys Val Met Gly Asn Arg Arg Ser Asp
245 250 255
Glu Pro Ala Lys Thr Lys Lys Gly Leu Ser Ser Ile Leu Asp Ala Ala
260 265 270
Arg Trp Asn Arg Gly Glu Pro Gln Val Pro Asp Phe Arg His Leu Lys
275 280 285
Ala Glu Val Asp Ala Glu Lys Pro Gly Ala Thr Asp Arg Lys Gly Gly
290 295 300
Val Gly Met Pro Ser Arg Asp Arg Asn Ile Gly Ala Pro Gly Gln Asp
305 310 315 320
Thr Pro Gly Val Ser Leu His Pro Leu Ser Leu Asp Ser Pro Asp Arg
325 330 335
Glu Pro Gly Ala Asp Ala Pro Leu Glu Leu Gly Asp Ser Ser Pro Gln
340 345 350
Gly Pro Met Ser Leu Glu Ser Leu Ala Pro Pro Glu Ser Thr Asp Glu
355 360 365
Gly Ala Glu Thr Glu Ser Pro Glu Pro Gly Asp Glu Gly Glu Pro Gly
370 375 380
Arg Ser Gly Leu Glu Leu Glu Pro Glu Glu Pro Pro Gly Trp Arg Glu
385 390 395 400
Leu Val Pro Pro Asp Thr Leu His Ser Leu Pro Lys Ser Gln Val Lys
405 410 415
Arg Gln Glu Val Ile Ser Glu Leu Leu Val Thr Glu Ala Ala His Val
420 425 430
Arg Met Leu Arg Val Leu His Asp Leu Phe Phe Gln Pro Met Ala Glu
435 440 445
Cys Leu Phe Phe Pro Leu Glu Glu Leu Gln Asn Ile Phe Pro Ser Leu
450 455 460
Asp Glu Leu Ile Glu Val His Ser Leu Phe Leu Asp Arg Leu Met Lys
465 470 475 480
Arg Arg Gln Glu Ser Gly Tyr Leu Ile Glu Glu Ile Gly Asp Val Leu
485 490 495
Leu Ala Arg Phe Asp Gly Ala Glu Gly Ser Trp Phe Gln Lys Ile Ser
500 505 510
Ser Arg Phe Cys Ser Arg Gln Ser Phe Ala Leu Glu Gln Leu Lys Ala
515 520 525
Lys Gln Arg Lys Asp Pro Arg Phe Cys Ala Phe Val Gln Glu Ala Glu
530 535 540
Ser Arg Pro Arg Cys Arg Arg Leu Gln Leu Lys Asp Met Ile Pro Thr
545 550 555 560
Glu Met Gln Arg Leu Thr Lys Tyr Pro Leu Leu Leu Gln Ser Ile Gly
565 570 575
Gln Asn Thr Glu Glu Pro Thr Glu Arg Glu Lys Val Glu Leu Ala Ala
580 585 590
Glu Cys Cys Arg Glu Ile Leu His His Val Asn Gln Ala Val Arg Asp
595 600 605
Met Glu Asp Leu Leu Arg Leu Lys Asp Tyr Gln Arg Arg Leu Asp Leu
610 615 620
Ser His Leu Arg Gln Ser Ser Asp Pro Met Leu Ser Glu Phe Lys Asn
625 630 635 640
Leu Asp Ile Thr Lys Lys Lys Leu Val His Glu Gly Pro Leu Thr Trp
645 650 655
Arg Val Thr Lys Asp Lys Ala Val Glu Val His Val Leu Leu Leu Asp
660 665 670
Asp Leu Leu Leu Leu Leu Gln Arg Gln Asp Glu Arg Leu Leu Leu Lys
675 680 685
Ser His Ser Arg Thr Leu Thr Pro Thr Pro Asp Gly Lys Thr Met Leu
690 695 700
Arg Pro Val Leu Arg Leu Thr Ser Ala Met Thr Arg Glu Val Ala Thr
705 710 715 720
Asp His Lys Ala Phe Tyr Val Leu Phe Thr Trp Asp Gln Glu Ala Gln
725 730 735
Ile Tyr Glu Leu Val Ala Gln Thr Val Ser Glu Arg Lys Asn Trp Cys
740 745 750
Ala Leu Ile Thr Glu Thr Ala Gly Ser Leu Lys Val Pro Ala Pro Ala
755 760 765
Ser Arg Pro Lys Pro Arg Pro Ser Pro Ser Ser Thr Arg Glu Pro Leu
770 775 780
Leu Ser Ser Ser Glu Asn Gly Asn Gly Gly Arg Glu Thr Ser Pro Ala
785 790 795 800
Asp Ala Arg Thr Glu Arg Ile Leu Ser Asp Leu Leu Pro Phe Cys Arg
805 810 815
Pro Gly Pro Glu Gly Gln Leu Ala Ala Thr Ala Leu Arg Lys Val Leu
820 825 830
Ser Leu Lys Gln Leu Leu Phe Pro Ala Glu Glu Asp Asn Gly Ala Gly
835 840 845
Pro Pro Arg Asp Gly Asp Gly Val Pro Gly Gly Gly Pro Leu Ser Pro
850 855 860
Ala Arg Thr Gln Glu Ile Gln Glu Asn Leu Leu Ser Leu Glu Glu Thr
865 870 875 880
Met Lys Gln Leu Glu Glu Leu Glu Glu Glu Phe Cys Arg Leu Arg Pro
885 890 895
Leu Leu Ser Gln Leu Gly Gly Asn Ser Val Pro Gln Pro Gly Cys Thr
900 905 910
<210> 24
<211> 7
<212> PRT
<213> 人工序列
<220>
<223> 合成肽接头
<400> 24
Arg Leu Lys Leu Pro Ala Thr
1 5
<210> 25
<211> 580
<212> PRT
<213> 智人
<400> 25
Met Arg Gly Gly His Lys Gly Gly Arg Cys Ala Cys Pro Arg Val Ile
1 5 10 15
Arg Lys Val Leu Ala Lys Cys Gly Cys Cys Phe Ala Arg Gly Gly Arg
20 25 30
Glu Ser Tyr Ser Ile Ala Gly Ser Glu Gly Ser Ile Ser Ala Ser Ala
35 40 45
Ala Ser Gly Leu Ala Ala Pro Ser Gly Pro Ser Ser Gly Leu Ser Ser
50 55 60
Gly Pro Cys Ser Pro Gly Pro Pro Gly Pro Val Ser Gly Leu Arg Arg
65 70 75 80
Trp Leu Asp His Ser Lys His Cys Leu Ser Val Glu Thr Glu Ala Asp
85 90 95
Ser Gly Gln Ala Gly Pro Tyr Glu Asn Trp Met Leu Glu Pro Ala Leu
100 105 110
Ala Thr Gly Glu Glu Leu Pro Glu Leu Thr Leu Leu Thr Thr Leu Leu
115 120 125
Glu Gly Pro Gly Asp Lys Thr Gln Pro Pro Glu Glu Glu Thr Leu Ser
130 135 140
Gln Ala Pro Glu Ser Glu Glu Glu Gln Lys Lys Lys Ala Leu Glu Arg
145 150 155 160
Ser Met Tyr Val Leu Ser Glu Leu Val Glu Thr Glu Lys Met Tyr Val
165 170 175
Asp Asp Leu Gly Gln Ile Val Glu Gly Tyr Met Ala Thr Met Ala Ala
180 185 190
Gln Gly Val Pro Glu Ser Leu Arg Gly Arg Asp Arg Ile Val Phe Gly
195 200 205
Asn Ile Gln Gln Ile Tyr Glu Trp His Arg Asp Tyr Phe Leu Gln Glu
210 215 220
Leu Gln Arg Cys Leu Lys Asp Pro Asp Trp Leu Ala Gln Leu Phe Ile
225 230 235 240
Lys His Glu Arg Arg Leu His Met Tyr Val Val Tyr Cys Gln Asn Lys
245 250 255
Pro Lys Ser Glu His Val Val Ser Glu Phe Gly Asp Ser Tyr Phe Glu
260 265 270
Glu Leu Arg Gln Gln Leu Gly His Arg Leu Gln Leu Asn Asp Leu Leu
275 280 285
Ile Lys Pro Val Gln Arg Ile Met Lys Tyr Gln Leu Leu Leu Lys Asp
290 295 300
Phe Leu Lys Tyr Tyr Asn Arg Ala Gly Met Asp Thr Ala Asp Leu Glu
305 310 315 320
Gln Ala Val Glu Val Met Cys Phe Val Pro Lys Arg Cys Asn Asp Met
325 330 335
Met Thr Leu Gly Arg Leu Arg Gly Phe Glu Gly Lys Leu Thr Ala Gln
340 345 350
Gly Lys Leu Leu Gly Gln Asp Thr Phe Trp Val Thr Glu Pro Glu Ala
355 360 365
Gly Gly Leu Leu Ser Ser Arg Gly Arg Glu Arg Arg Val Phe Leu Phe
370 375 380
Glu Gln Ile Ile Ile Phe Ser Glu Ala Leu Gly Gly Gly Val Arg Gly
385 390 395 400
Gly Thr Gln Pro Gly Tyr Val Tyr Lys Asn Ser Ile Lys Val Ser Cys
405 410 415
Leu Gly Leu Glu Gly Asn Leu Gln Gly Asp Pro Cys Arg Phe Ala Leu
420 425 430
Thr Ser Arg Gly Pro Glu Gly Gly Ile Gln Arg Tyr Val Leu Gln Ala
435 440 445
Ala Asp Pro Ala Ile Ser Gln Ala Trp Ile Lys His Val Ala Gln Ile
450 455 460
Leu Glu Ser Gln Arg Asp Phe Leu Asn Ala Leu Gln Ser Pro Ile Glu
465 470 475 480
Tyr Gln Arg Arg Glu Ser Gln Thr Asn Ser Leu Gly Arg Pro Arg Gly
485 490 495
Pro Gly Val Gly Ser Pro Gly Arg Ile Gln Leu Gly Asp Gln Ala Gln
500 505 510
Gly Ser Thr His Thr Pro Ile Asn Gly Ser Leu Pro Ser Leu Leu Leu
515 520 525
Ser Pro Lys Gly Glu Val Ala Arg Ala Leu Leu Pro Leu Asp Lys Gln
530 535 540
Ala Leu Gly Asp Ile Pro Gln Ala Pro His Asp Ser Pro Pro Val Ser
545 550 555 560
Pro Thr Pro Lys Thr Pro Pro Cys Gln Ala Arg Leu Ala Lys Leu Asp
565 570 575
Glu Asp Glu Leu
580
<210> 26
<211> 19
<212> PRT
<213> 人工序列
<220>
<223> 合成肽接头
<400> 26
Ala Ser Gly Ser Ala Gly Thr Gly Gly Arg Ala Ile Asp Ile Lys Leu
1 5 10 15
Pro Ala Thr
<210> 27
<211> 689
<212> PRT
<213> 智人
<400> 27
Met Ala Asp Leu Glu Ala Val Leu Ala Asp Val Ser Tyr Leu Met Ala
1 5 10 15
Met Glu Lys Ser Lys Ala Thr Pro Ala Ala Arg Ala Ser Lys Lys Ile
20 25 30
Leu Leu Pro Glu Pro Ser Ile Arg Ser Val Met Gln Lys Tyr Leu Glu
35 40 45
Asp Arg Gly Glu Val Thr Phe Glu Lys Ile Phe Ser Gln Lys Leu Gly
50 55 60
Tyr Leu Leu Phe Arg Asp Phe Cys Leu Asn His Leu Glu Glu Ala Arg
65 70 75 80
Pro Leu Val Glu Phe Tyr Glu Glu Ile Lys Lys Tyr Glu Lys Leu Glu
85 90 95
Thr Glu Glu Glu Arg Val Ala Arg Ser Arg Glu Ile Phe Asp Ser Tyr
100 105 110
Ile Met Lys Glu Leu Leu Ala Cys Ser His Pro Phe Ser Lys Ser Ala
115 120 125
Thr Glu His Val Gln Gly His Leu Gly Lys Lys Gln Val Pro Pro Asp
130 135 140
Leu Phe Gln Pro Tyr Ile Glu Glu Ile Cys Gln Asn Leu Arg Gly Asp
145 150 155 160
Val Phe Gln Lys Phe Ile Glu Ser Asp Lys Phe Thr Arg Phe Cys Gln
165 170 175
Trp Lys Asn Val Glu Leu Asn Ile His Leu Thr Met Asn Asp Phe Ser
180 185 190
Val His Arg Ile Ile Gly Arg Gly Gly Phe Gly Glu Val Tyr Gly Cys
195 200 205
Arg Lys Ala Asp Thr Gly Lys Met Tyr Ala Met Lys Cys Leu Asp Lys
210 215 220
Lys Arg Ile Lys Met Lys Gln Gly Glu Thr Leu Ala Leu Asn Glu Arg
225 230 235 240
Ile Met Leu Ser Leu Val Ser Thr Gly Asp Cys Pro Phe Ile Val Cys
245 250 255
Met Ser Tyr Ala Phe His Thr Pro Asp Lys Leu Ser Phe Ile Leu Asp
260 265 270
Leu Met Asn Gly Gly Asp Leu His Tyr His Leu Ser Gln His Gly Val
275 280 285
Phe Ser Glu Ala Asp Met Arg Phe Tyr Ala Ala Glu Ile Ile Leu Gly
290 295 300
Leu Glu His Met His Asn Arg Phe Val Val Tyr Arg Asp Leu Lys Pro
305 310 315 320
Ala Asn Ile Leu Leu Asp Glu His Gly His Val Arg Ile Ser Asp Leu
325 330 335
Gly Leu Ala Cys Asp Phe Ser Lys Lys Lys Pro His Ala Ser Val Gly
340 345 350
Thr His Gly Tyr Met Ala Pro Glu Val Leu Gln Lys Gly Val Ala Tyr
355 360 365
Asp Ser Ser Ala Asp Trp Phe Ser Leu Gly Cys Met Leu Phe Lys Leu
370 375 380
Leu Arg Gly His Ser Pro Phe Arg Gln His Lys Thr Lys Asp Lys His
385 390 395 400
Glu Ile Asp Arg Met Thr Leu Thr Met Ala Val Glu Leu Pro Asp Ser
405 410 415
Phe Ser Pro Glu Leu Arg Ser Leu Leu Glu Gly Leu Leu Gln Arg Asp
420 425 430
Val Asn Arg Arg Leu Gly Cys Leu Gly Arg Gly Ala Gln Glu Val Lys
435 440 445
Glu Ser Pro Phe Phe Arg Ser Leu Asp Trp Gln Met Val Phe Leu Gln
450 455 460
Lys Tyr Pro Pro Pro Leu Ile Pro Pro Arg Gly Glu Val Asn Ala Ala
465 470 475 480
Asp Ala Phe Asp Ile Gly Ser Phe Asp Glu Glu Asp Thr Lys Gly Ile
485 490 495
Lys Leu Leu Asp Ser Asp Gln Glu Leu Tyr Arg Asn Phe Pro Leu Thr
500 505 510
Ile Ser Glu Arg Trp Gln Gln Glu Val Ala Glu Thr Val Phe Asp Thr
515 520 525
Ile Asn Ala Glu Thr Asp Arg Leu Glu Ala Arg Lys Lys Ala Lys Asn
530 535 540
Lys Gln Leu Gly His Glu Glu Asp Tyr Ala Leu Gly Lys Asp Cys Ile
545 550 555 560
Met His Gly Tyr Met Ser Lys Met Gly Asn Pro Phe Leu Thr Gln Trp
565 570 575
Gln Arg Arg Tyr Phe Tyr Leu Phe Pro Asn Arg Leu Glu Trp Arg Gly
580 585 590
Glu Gly Glu Ala Pro Gln Ser Leu Leu Thr Met Glu Glu Ile Gln Ser
595 600 605
Val Glu Glu Thr Gln Ile Lys Glu Arg Lys Cys Leu Leu Leu Lys Ile
610 615 620
Arg Gly Gly Lys Gln Phe Ile Leu Gln Cys Asp Ser Asp Pro Glu Leu
625 630 635 640
Val Gln Trp Lys Lys Glu Leu Arg Asp Ala Tyr Arg Glu Ala Gln Gln
645 650 655
Leu Val Gln Arg Val Pro Lys Met Lys Asn Lys Pro Arg Ser Pro Val
660 665 670
Val Glu Leu Ser Lys Val Pro Leu Val Gln Arg Gly Ser Ala Asn Gly
675 680 685
Leu
<210> 28
<211> 350
<212> PRT
<213> 智人
<400> 28
Met Gly Cys Thr Val Ser Ala Glu Asp Lys Ala Ala Ala Glu Arg Ser
1 5 10 15
Lys Met Ile Asp Lys Asn Leu Arg Glu Asp Gly Glu Lys Ala Ala Arg
20 25 30
Glu Val Lys Leu Leu Leu Leu Gly Ala Gly Glu Ser Gly Lys Ser Thr
35 40 45
Ile Val Lys Gln Met Lys Ile Ile His Glu Asp Gly Tyr Ser Glu Glu
50 55 60
Glu Cys Arg Gln Tyr Arg Ala Val Val Tyr Ser Asn Thr Ile Gln Ser
65 70 75 80
Ile Met Ala Ile Val Lys Ala Met Gly Asn Leu Gln Ile Asp Phe Ala
85 90 95
Asp Pro Ser Arg Ala Asp Asp Ala Arg Gln Leu Phe Ala Leu Ser Cys
100 105 110
Thr Ala Glu Glu Gln Gly Val Leu Pro Asp Asp Leu Ser Gly Val Ile
115 120 125
Arg Arg Leu Trp Ala Asp His Gly Val Gln Ala Cys Phe Gly Arg Ser
130 135 140
Arg Glu Tyr Gln Leu Asn Asp Ser Ala Ala Tyr Tyr Leu Asn Asp Leu
145 150 155 160
Glu Arg Ile Ala Gln Ser Asp Tyr Ile Pro Thr Gln Gln Asp Val Leu
165 170 175
Arg Thr Arg Val Lys Thr Thr Gly Ile Val Glu Thr His Phe Thr Phe
180 185 190
Lys Asp Leu His Phe Lys Met Phe Asp Val Gly Gly Gln Arg Ser Glu
195 200 205
Arg Lys Lys Trp Ile His Cys Phe Glu Gly Val Thr Ala Ile Ile Phe
210 215 220
Cys Val Ala Leu Ser Ala Tyr Asp Leu Val Leu Ala Glu Asp Glu Glu
225 230 235 240
Met Asn Arg Met His Glu Ser Met Lys Leu Phe Asp Ser Ile Cys Asn
245 250 255
Asn Lys Trp Phe Thr Asp Thr Ser Ile Ile Leu Phe Leu Asn Lys Lys
260 265 270
Asp Leu Phe Glu Glu Lys Ile Thr His Ser Pro Leu Thr Ile Cys Phe
275 280 285
Pro Glu Tyr Thr Gly Ala Asn Lys Tyr Asp Glu Ala Ala Ser Tyr Ile
290 295 300
Gln Ser Lys Phe Glu Asp Leu Asn Lys Arg Lys Asp Thr Lys Glu Ile
305 310 315 320
Tyr Thr His Phe Thr Cys Ala Thr Asp Thr Lys Asn Val Gln Phe Val
325 330 335
Phe Asp Ala Val Thr Asp Val Ile Ile Lys Asn Asn Leu Lys
340 345 350
<210> 29
<211> 353
<212> PRT
<213> 智人
<400> 29
Met Gly Cys Thr Val Ser Ala Glu Asp Lys Ala Ala Ala Glu Arg Ser
1 5 10 15
Lys Met Ile Asp Lys Asn Leu Arg Glu Asp Gly Glu Lys Ala Ala Arg
20 25 30
Glu Val Lys Leu Leu Leu Leu Gly Ala Gly Glu Ser Gly Lys Ser Thr
35 40 45
Ile Val Lys Gln Met Lys Ile Ile His Glu Asp Gly Tyr Ser Glu Glu
50 55 60
Glu Cys Arg Gln Tyr Arg Ala Val Val Tyr Ser Asn Thr Ile Gln Ser
65 70 75 80
Ile Met Ala Ile Val Lys Ala Met Gly Asn Leu Gln Ile Asp Phe Ala
85 90 95
Asp Pro Ser Arg Ala Asp Asp Ala Arg Gln Leu Phe Ala Leu Ser Cys
100 105 110
Thr Ala Glu Glu Gln Gly Val Leu Pro Asp Asp Leu Ser Gly Val Ile
115 120 125
Arg Arg Leu Trp Ala Asp His Gly Val Gln Ala Cys Phe Gly Arg Ser
130 135 140
Arg Glu Tyr Gln Leu Asn Asp Ser Ala Ala Tyr Tyr Leu Asn Asp Leu
145 150 155 160
Glu Arg Ile Ala Gln Ser Asp Tyr Ile Pro Thr Gln Gln Asp Val Leu
165 170 175
Arg Thr Arg Val Lys Thr Thr Gly Ile Val Glu Thr His Phe Thr Phe
180 185 190
Lys Asp Leu His Phe Lys Met Phe Asp Val Gly Gly Gln Arg Ser Glu
195 200 205
Arg Lys Lys Trp Ile His Cys Phe Glu Gly Val Thr Ala Ile Ile Phe
210 215 220
Cys Val Ala Leu Ser Ala Tyr Asp Leu Val Leu Ala Glu Asp Glu Glu
225 230 235 240
Met Asn Arg Met His Glu Ser Met Lys Leu Phe Asp Ser Ile Cys Asn
245 250 255
Asn Lys Trp Phe Thr Asp Thr Ser Ile Ile Leu Phe Leu Asn Lys Lys
260 265 270
Asp Leu Phe Glu Glu Lys Ile Thr His Ser Pro Leu Thr Ile Cys Phe
275 280 285
Pro Glu Tyr Thr Gly Ala Asn Lys Tyr Asp Glu Ala Ala Ser Tyr Ile
290 295 300
Gln Ser Lys Phe Glu Asp Leu Asn Lys Arg Lys Asp Thr Lys Glu Ile
305 310 315 320
Tyr Thr His Phe Thr Cys Ala Thr Asp Thr Lys Asn Val Gln Phe Val
325 330 335
Phe Asp Ala Val Thr Asp Val Ile Ile Lys Asn Asn Leu Lys Asp Cys
340 345 350
Gly
<210> 30
<211> 355
<212> PRT
<213> 智人
<400> 30
Met Gly Cys Thr Val Ser Ala Glu Asp Lys Ala Ala Ala Glu Arg Ser
1 5 10 15
Lys Met Ile Asp Lys Asn Leu Arg Glu Asp Gly Glu Lys Ala Ala Arg
20 25 30
Glu Val Lys Leu Leu Leu Leu Gly Ala Gly Glu Ser Gly Lys Ser Thr
35 40 45
Ile Val Lys Gln Met Lys Ile Ile His Glu Asp Gly Tyr Ser Glu Glu
50 55 60
Glu Cys Arg Gln Tyr Arg Ala Val Val Tyr Ser Asn Thr Ile Gln Ser
65 70 75 80
Ile Met Ala Ile Val Lys Ala Met Gly Asn Leu Gln Ile Asp Phe Ala
85 90 95
Asp Pro Ser Arg Ala Asp Asp Ala Arg Gln Leu Phe Ala Leu Ser Cys
100 105 110
Thr Ala Glu Glu Gln Gly Val Leu Pro Asp Asp Leu Ser Gly Val Ile
115 120 125
Arg Arg Leu Trp Ala Asp His Gly Val Gln Ala Cys Phe Gly Arg Ser
130 135 140
Arg Glu Tyr Gln Leu Asn Asp Ser Ala Ala Tyr Tyr Leu Asn Asp Leu
145 150 155 160
Glu Arg Ile Ala Gln Ser Asp Tyr Ile Pro Thr Gln Gln Asp Val Leu
165 170 175
Arg Thr Arg Val Lys Thr Thr Gly Ile Val Glu Thr His Phe Thr Phe
180 185 190
Lys Asp Leu His Phe Lys Met Phe Asp Val Gly Gly Gln Arg Ser Glu
195 200 205
Arg Lys Lys Trp Ile His Cys Phe Glu Gly Val Thr Ala Ile Ile Phe
210 215 220
Cys Val Ala Leu Ser Ala Tyr Asp Leu Val Leu Ala Glu Asp Glu Glu
225 230 235 240
Met Asn Arg Met His Glu Ser Met Lys Leu Phe Asp Ser Ile Cys Asn
245 250 255
Asn Lys Trp Phe Thr Asp Thr Ser Ile Ile Leu Phe Leu Asn Lys Lys
260 265 270
Asp Leu Phe Glu Glu Lys Ile Thr His Ser Pro Leu Thr Ile Cys Phe
275 280 285
Pro Glu Tyr Thr Gly Ala Asn Lys Tyr Asp Glu Ala Ala Ser Tyr Ile
290 295 300
Gln Ser Lys Phe Glu Asp Leu Asn Lys Arg Lys Asp Thr Lys Glu Ile
305 310 315 320
Tyr Thr His Phe Thr Cys Ala Thr Asp Thr Lys Asn Val Gln Phe Val
325 330 335
Phe Asp Ala Val Thr Asp Val Ile Ile Lys Asn Asn Gly Lys Asp Cys
340 345 350
Gly Leu Phe
355
<210> 31
<211> 355
<212> PRT
<213> 智人
<400> 31
Met Gly Cys Thr Val Ser Ala Glu Asp Lys Ala Ala Ala Glu Arg Ser
1 5 10 15
Lys Met Ile Asp Lys Asn Leu Arg Glu Asp Gly Glu Lys Ala Ala Arg
20 25 30
Glu Val Lys Leu Leu Leu Leu Gly Ala Gly Glu Ser Gly Lys Ser Thr
35 40 45
Ile Val Lys Gln Met Lys Ile Ile His Glu Asp Gly Tyr Ser Glu Glu
50 55 60
Glu Cys Arg Gln Tyr Arg Ala Val Val Tyr Ser Asn Thr Ile Gln Ser
65 70 75 80
Ile Met Ala Ile Val Lys Ala Met Gly Asn Leu Gln Ile Asp Phe Ala
85 90 95
Asp Pro Ser Arg Ala Asp Asp Ala Arg Gln Leu Phe Ala Leu Ser Cys
100 105 110
Thr Ala Glu Glu Gln Gly Val Leu Pro Asp Asp Leu Ser Gly Val Ile
115 120 125
Arg Arg Leu Trp Ala Asp His Gly Val Gln Ala Cys Phe Gly Arg Ser
130 135 140
Arg Glu Tyr Gln Leu Asn Asp Ser Ala Ala Tyr Tyr Leu Asn Asp Leu
145 150 155 160
Glu Arg Ile Ala Gln Ser Asp Tyr Ile Pro Thr Gln Gln Asp Val Leu
165 170 175
Arg Thr Arg Val Lys Thr Thr Gly Ile Val Glu Thr His Phe Thr Phe
180 185 190
Lys Asp Leu His Phe Lys Met Phe Asp Val Gly Gly Gln Arg Ser Glu
195 200 205
Arg Lys Lys Trp Ile His Cys Phe Glu Gly Val Thr Ala Ile Ile Phe
210 215 220
Cys Val Ala Leu Ser Ala Tyr Asp Leu Val Leu Ala Glu Asp Glu Glu
225 230 235 240
Met Asn Arg Met His Glu Ser Met Lys Leu Phe Asp Ser Ile Cys Asn
245 250 255
Asn Lys Trp Phe Thr Asp Thr Ser Ile Ile Leu Phe Leu Asn Lys Lys
260 265 270
Asp Leu Phe Glu Glu Lys Ile Thr His Ser Pro Leu Thr Ile Cys Phe
275 280 285
Pro Glu Tyr Thr Gly Ala Asn Lys Tyr Asp Glu Ala Ala Ser Tyr Ile
290 295 300
Gln Ser Lys Phe Glu Asp Leu Asn Lys Arg Lys Asp Thr Lys Glu Ile
305 310 315 320
Tyr Thr His Phe Thr Cys Ala Thr Asp Thr Lys Asn Val Gln Phe Val
325 330 335
Phe Asp Ala Val Thr Asp Val Ile Ile Lys Asn Asn Leu Lys Asp Cys
340 345 350
Gly Gly Phe
355
<210> 32
<211> 355
<212> PRT
<213> 智人
<400> 32
Met Gly Cys Thr Val Ser Ala Glu Asp Lys Ala Ala Ala Glu Arg Ser
1 5 10 15
Lys Met Ile Asp Lys Asn Leu Arg Glu Asp Gly Glu Lys Ala Ala Arg
20 25 30
Glu Val Lys Leu Leu Leu Leu Gly Ala Gly Glu Ser Gly Lys Ser Thr
35 40 45
Ile Val Lys Gln Met Lys Ile Ile His Glu Asp Gly Tyr Ser Glu Glu
50 55 60
Glu Cys Arg Gln Tyr Arg Ala Val Val Tyr Ser Asn Thr Ile Gln Ser
65 70 75 80
Ile Met Ala Ile Val Lys Ala Met Gly Asn Leu Gln Ile Asp Phe Ala
85 90 95
Asp Pro Ser Arg Ala Asp Asp Ala Arg Gln Leu Phe Ala Leu Ser Cys
100 105 110
Thr Ala Glu Glu Gln Gly Val Leu Pro Asp Asp Leu Ser Gly Val Ile
115 120 125
Arg Arg Leu Trp Ala Asp His Gly Val Gln Ala Cys Phe Gly Arg Ser
130 135 140
Arg Glu Tyr Gln Leu Asn Asp Ser Ala Ala Tyr Tyr Leu Asn Asp Leu
145 150 155 160
Glu Arg Ile Ala Gln Ser Asp Tyr Ile Pro Thr Gln Gln Asp Val Leu
165 170 175
Arg Thr Arg Val Lys Thr Thr Gly Ile Val Glu Thr His Phe Thr Phe
180 185 190
Lys Asp Leu His Phe Lys Met Phe Asp Val Gly Gly Gln Arg Ser Glu
195 200 205
Arg Lys Lys Trp Ile His Cys Phe Glu Gly Val Thr Ala Ile Ile Phe
210 215 220
Cys Val Ala Leu Ser Ala Tyr Asp Leu Val Leu Ala Glu Asp Glu Glu
225 230 235 240
Met Asn Arg Met His Glu Ser Met Lys Leu Phe Asp Ser Ile Cys Asn
245 250 255
Asn Lys Trp Phe Thr Asp Thr Ser Ile Ile Leu Phe Leu Asn Lys Lys
260 265 270
Asp Leu Phe Glu Glu Lys Ile Thr His Ser Pro Leu Thr Ile Cys Phe
275 280 285
Pro Glu Tyr Thr Gly Ala Asn Lys Tyr Asp Glu Ala Ala Ser Tyr Ile
290 295 300
Gln Ser Lys Phe Glu Asp Leu Asn Lys Arg Lys Asp Thr Lys Glu Ile
305 310 315 320
Tyr Thr His Phe Thr Cys Ala Thr Asp Thr Lys Asn Val Gln Phe Val
325 330 335
Phe Asp Ala Val Thr Asp Val Ile Ile Lys Asn Asn Leu Lys Asp Cys
340 345 350
Gly Asp Phe
355
<210> 33
<211> 355
<212> PRT
<213> 智人
<400> 33
Met Gly Cys Thr Val Ser Ala Glu Asp Lys Ala Ala Ala Glu Arg Ser
1 5 10 15
Lys Met Ile Asp Lys Asn Leu Arg Glu Asp Gly Glu Lys Ala Ala Arg
20 25 30
Glu Val Lys Leu Leu Leu Leu Gly Ala Gly Glu Ser Gly Lys Ser Thr
35 40 45
Ile Val Lys Gln Met Lys Ile Ile His Glu Asp Gly Tyr Ser Glu Glu
50 55 60
Glu Cys Arg Gln Tyr Arg Ala Val Val Tyr Ser Asn Thr Ile Gln Ser
65 70 75 80
Ile Met Ala Ile Val Lys Ala Met Gly Asn Leu Gln Ile Asp Phe Ala
85 90 95
Asp Pro Ser Arg Ala Asp Asp Ala Arg Gln Leu Phe Ala Leu Ser Cys
100 105 110
Thr Ala Glu Glu Gln Gly Val Leu Pro Asp Asp Leu Ser Gly Val Ile
115 120 125
Arg Arg Leu Trp Ala Asp His Gly Val Gln Ala Cys Phe Gly Arg Ser
130 135 140
Arg Glu Tyr Gln Leu Asn Asp Ser Ala Ala Tyr Tyr Leu Asn Asp Leu
145 150 155 160
Glu Arg Ile Ala Gln Ser Asp Tyr Ile Pro Thr Gln Gln Asp Val Leu
165 170 175
Arg Thr Arg Val Lys Thr Thr Gly Ile Val Glu Thr His Phe Thr Phe
180 185 190
Lys Asp Leu His Phe Lys Met Phe Asp Val Gly Gly Gln Arg Ser Glu
195 200 205
Arg Lys Lys Trp Ile His Cys Phe Glu Gly Val Thr Ala Ile Ile Phe
210 215 220
Cys Val Ala Leu Ser Ala Tyr Asp Leu Val Leu Ala Glu Asp Glu Glu
225 230 235 240
Met Asn Arg Met His Glu Ser Met Lys Leu Phe Asp Ser Ile Cys Asn
245 250 255
Asn Lys Trp Phe Thr Asp Thr Ser Ile Ile Leu Phe Leu Asn Lys Lys
260 265 270
Asp Leu Phe Glu Glu Lys Ile Thr His Ser Pro Leu Thr Ile Cys Phe
275 280 285
Pro Glu Tyr Thr Gly Ala Asn Lys Tyr Asp Glu Ala Ala Ser Tyr Ile
290 295 300
Gln Ser Lys Phe Glu Asp Leu Asn Lys Arg Lys Asp Thr Lys Glu Ile
305 310 315 320
Tyr Thr His Phe Thr Cys Ala Thr Asp Thr Lys Asn Val Gln Phe Val
325 330 335
Phe Asp Ala Val Thr Asp Val Ile Ile Lys Asn Asn Leu Lys Asp Cys
340 345 350
Gly Pro Phe
355
<210> 34
<211> 355
<212> PRT
<213> 智人
<400> 34
Met Gly Cys Thr Val Ser Ala Glu Asp Lys Ala Ala Ala Glu Arg Ser
1 5 10 15
Lys Met Ile Asp Lys Asn Leu Arg Glu Asp Gly Glu Lys Ala Ala Arg
20 25 30
Glu Val Lys Leu Leu Leu Leu Gly Ala Gly Glu Ser Gly Lys Ser Thr
35 40 45
Ile Val Lys Gln Met Lys Ile Ile His Glu Asp Gly Tyr Ser Glu Glu
50 55 60
Glu Cys Arg Gln Tyr Arg Ala Val Val Tyr Ser Asn Thr Ile Gln Ser
65 70 75 80
Ile Met Ala Ile Val Lys Ala Met Gly Asn Leu Gln Ile Asp Phe Ala
85 90 95
Asp Pro Ser Arg Ala Asp Asp Ala Arg Gln Leu Phe Ala Leu Ser Cys
100 105 110
Thr Ala Glu Glu Gln Gly Val Leu Pro Asp Asp Leu Ser Gly Val Ile
115 120 125
Arg Arg Leu Trp Ala Asp His Gly Val Gln Ala Cys Phe Gly Arg Ser
130 135 140
Arg Glu Tyr Gln Leu Asn Asp Ser Ala Ala Tyr Tyr Leu Asn Asp Leu
145 150 155 160
Glu Arg Ile Ala Gln Ser Asp Tyr Ile Pro Thr Gln Gln Asp Val Leu
165 170 175
Arg Thr Arg Val Lys Thr Thr Gly Ile Val Glu Thr His Phe Thr Phe
180 185 190
Lys Asp Leu His Phe Lys Met Phe Asp Val Gly Gly Gln Arg Ser Glu
195 200 205
Arg Lys Lys Trp Ile His Cys Phe Glu Gly Val Thr Ala Ile Ile Phe
210 215 220
Cys Val Ala Leu Ser Ala Tyr Asp Leu Val Leu Ala Glu Asp Glu Glu
225 230 235 240
Met Asn Arg Met His Glu Ser Met Lys Leu Phe Asp Ser Ile Cys Asn
245 250 255
Asn Lys Trp Phe Thr Asp Thr Ser Ile Ile Leu Phe Leu Asn Lys Lys
260 265 270
Asp Leu Phe Glu Glu Lys Ile Thr His Ser Pro Leu Thr Ile Cys Phe
275 280 285
Pro Glu Tyr Thr Gly Ala Asn Lys Tyr Asp Glu Ala Ala Ser Tyr Ile
290 295 300
Gln Ser Lys Phe Glu Asp Leu Asn Lys Arg Lys Asp Thr Lys Glu Ile
305 310 315 320
Tyr Thr His Phe Thr Cys Ala Thr Asp Thr Lys Asn Val Gln Phe Val
325 330 335
Phe Asp Ala Val Thr Asp Val Ile Ile Lys Asn Asn Leu Lys Asp Cys
340 345 350
Gly Arg Phe
355
<210> 35
<211> 354
<212> PRT
<213> 智人
<400> 35
Met Gly Cys Thr Leu Ser Ala Glu Glu Arg Ala Ala Leu Glu Arg Ser
1 5 10 15
Lys Ala Ile Glu Lys Asn Leu Lys Glu Asp Gly Ile Ser Ala Ala Lys
20 25 30
Asp Val Lys Leu Leu Leu Leu Gly Ala Gly Glu Ser Gly Lys Ser Thr
35 40 45
Ile Val Lys Gln Met Lys Ile Ile His Glu Asp Gly Phe Ser Gly Glu
50 55 60
Asp Val Lys Gln Tyr Lys Pro Val Val Tyr Ser Asn Thr Ile Gln Ser
65 70 75 80
Leu Ala Ala Ile Val Arg Ala Met Asp Thr Leu Gly Ile Glu Tyr Gly
85 90 95
Asp Lys Glu Arg Lys Ala Asp Ala Lys Met Val Cys Asp Val Val Ser
100 105 110
Arg Met Glu Asp Thr Glu Pro Phe Ser Ala Glu Leu Leu Ser Ala Met
115 120 125
Met Arg Leu Trp Gly Asp Ser Gly Ile Gln Glu Cys Phe Asn Arg Ser
130 135 140
Arg Glu Tyr Gln Leu Asn Asp Ser Ala Lys Tyr Tyr Leu Asp Ser Leu
145 150 155 160
Asp Arg Ile Gly Ala Ala Asp Tyr Gln Pro Thr Glu Gln Asp Ile Leu
165 170 175
Arg Thr Arg Val Lys Thr Thr Gly Ile Val Glu Thr His Phe Thr Phe
180 185 190
Lys Asn Leu His Phe Arg Leu Phe Asp Val Gly Gly Gln Arg Ser Glu
195 200 205
Arg Lys Lys Trp Ile His Cys Phe Glu Asp Val Thr Ala Ile Ile Phe
210 215 220
Cys Val Ala Leu Ser Gly Tyr Asp Gln Val Leu His Glu Asp Glu Thr
225 230 235 240
Thr Asn Arg Met His Glu Ser Leu Lys Leu Phe Asp Ser Ile Cys Asn
245 250 255
Asn Lys Trp Phe Thr Asp Thr Ser Ile Ile Leu Phe Leu Asn Lys Lys
260 265 270
Asp Ile Phe Glu Glu Lys Ile Lys Lys Ser Pro Leu Thr Ile Cys Phe
275 280 285
Pro Glu Tyr Thr Gly Pro Ser Ala Phe Thr Glu Ala Val Ala Tyr Ile
290 295 300
Gln Ala Gln Tyr Glu Ser Lys Asn Lys Ser Ala His Lys Glu Ile Tyr
305 310 315 320
Thr His Val Thr Cys Ala Thr Asp Thr Asn Asn Ile Gln Phe Val Phe
325 330 335
Asp Ala Val Thr Asp Val Ile Ile Ala Lys Asn Leu Arg Gly Cys Gly
340 345 350
Gly Tyr
<210> 36
<211> 349
<212> PRT
<213> 智人
<400> 36
Met Gly Cys Thr Leu Ser Ala Glu Glu Arg Ala Ala Leu Glu Arg Ser
1 5 10 15
Lys Ala Ile Glu Lys Asn Leu Lys Glu Asp Gly Ile Ser Ala Ala Lys
20 25 30
Asp Val Lys Leu Leu Leu Leu Gly Ala Gly Glu Ser Gly Lys Ser Thr
35 40 45
Ile Val Lys Gln Met Lys Ile Ile His Glu Asp Gly Phe Ser Gly Glu
50 55 60
Asp Val Lys Gln Tyr Lys Pro Val Val Tyr Ser Asn Thr Ile Gln Ser
65 70 75 80
Leu Ala Ala Ile Val Arg Ala Met Asp Thr Leu Gly Ile Glu Tyr Gly
85 90 95
Asp Lys Glu Arg Lys Ala Asp Ala Lys Met Val Cys Asp Val Val Ser
100 105 110
Arg Met Glu Asp Thr Glu Pro Phe Ser Ala Glu Leu Leu Ser Ala Met
115 120 125
Met Arg Leu Trp Gly Asp Ser Gly Ile Gln Glu Cys Phe Asn Arg Ser
130 135 140
Arg Glu Tyr Gln Leu Asn Asp Ser Ala Lys Tyr Tyr Leu Asp Ser Leu
145 150 155 160
Asp Arg Ile Gly Ala Ala Asp Tyr Gln Pro Thr Glu Gln Asp Ile Leu
165 170 175
Arg Thr Arg Val Lys Thr Thr Gly Ile Val Glu Thr His Phe Thr Phe
180 185 190
Lys Asn Leu His Phe Arg Leu Phe Asp Val Gly Gly Gln Arg Ser Glu
195 200 205
Arg Lys Lys Trp Ile His Cys Phe Glu Asp Val Thr Ala Ile Ile Phe
210 215 220
Cys Val Ala Leu Ser Gly Tyr Asp Gln Val Leu His Glu Asp Glu Thr
225 230 235 240
Thr Asn Arg Met His Glu Ser Leu Lys Leu Phe Asp Ser Ile Cys Asn
245 250 255
Asn Lys Trp Phe Thr Asp Thr Ser Ile Ile Leu Phe Leu Asn Lys Lys
260 265 270
Asp Ile Phe Glu Glu Lys Ile Lys Lys Ser Pro Leu Thr Ile Cys Phe
275 280 285
Pro Glu Tyr Thr Gly Pro Ser Ala Phe Thr Glu Ala Val Ala Tyr Ile
290 295 300
Gln Ala Gln Tyr Glu Ser Lys Asn Lys Ser Ala His Lys Glu Ile Tyr
305 310 315 320
Thr His Val Thr Cys Ala Thr Asp Thr Asn Asn Ile Gln Phe Val Phe
325 330 335
Asp Ala Val Thr Asp Val Ile Ile Ala Lys Asn Leu Arg
340 345
<210> 37
<211> 311
<212> PRT
<213> Renilla reniformis
<400> 37
Met Thr Ser Lys Val Tyr Asp Pro Glu Gln Arg Lys Arg Met Ile Thr
1 5 10 15
Gly Pro Gln Trp Trp Ala Arg Cys Lys Gln Met Asn Val Leu Asp Ser
20 25 30
Phe Ile Asn Tyr Tyr Asp Ser Glu Lys His Ala Glu Asn Ala Val Ile
35 40 45
Phe Leu His Gly Asn Ala Thr Ser Ser Tyr Leu Trp Arg His Val Val
50 55 60
Pro His Ile Glu Pro Val Ala Arg Cys Ile Ile Pro Asp Leu Ile Gly
65 70 75 80
Met Gly Lys Ser Gly Lys Ser Gly Asn Gly Ser Tyr Arg Leu Leu Asp
85 90 95
His Tyr Lys Tyr Leu Thr Ala Trp Phe Glu Leu Leu Asn Leu Pro Lys
100 105 110
Lys Ile Ile Phe Val Gly His Asp Trp Gly Ala Ala Leu Ala Phe His
115 120 125
Tyr Ser Tyr Glu His Gln Asp Lys Ile Lys Ala Ile Val His Ala Glu
130 135 140
Ser Val Val Asp Val Ile Glu Ser Trp Asp Glu Trp Pro Asp Ile Glu
145 150 155 160
Glu Asp Ile Ala Leu Ile Lys Ser Glu Glu Gly Glu Lys Met Val Leu
165 170 175
Glu Asn Asn Phe Phe Val Glu Thr Val Leu Pro Ser Lys Ile Met Arg
180 185 190
Lys Leu Glu Pro Glu Glu Phe Ala Ala Tyr Leu Glu Pro Phe Lys Glu
195 200 205
Lys Gly Glu Val Arg Arg Pro Thr Leu Ser Trp Pro Arg Glu Ile Pro
210 215 220
Leu Val Lys Gly Gly Lys Pro Asp Val Val Gln Ile Val Arg Asn Tyr
225 230 235 240
Asn Ala Tyr Leu Arg Ala Ser Asp Asp Leu Pro Lys Met Phe Ile Glu
245 250 255
Ser Asp Pro Gly Phe Phe Ser Asn Ala Ile Val Glu Gly Ala Lys Lys
260 265 270
Phe Pro Asn Thr Glu Phe Val Lys Val Lys Gly Leu His Phe Ser Gln
275 280 285
Glu Asp Ala Pro Asp Glu Met Gly Lys Tyr Ile Lys Ser Phe Val Glu
290 295 300
Arg Val Leu Lys Asn Glu Gln
305 310
<210> 38
<211> 233
<212> PRT
<213> Renilla reniformis
<400> 38
Met Asp Leu Ala Lys Leu Gly Leu Lys Glu Val Met Pro Thr Lys Ile
1 5 10 15
Asn Leu Glu Gly Leu Val Gly Asp His Ala Phe Ser Met Glu Gly Val
20 25 30
Gly Glu Gly Asn Ile Leu Glu Gly Thr Gln Glu Val Lys Ile Ser Val
35 40 45
Thr Lys Gly Ala Pro Leu Pro Phe Ala Phe Asp Ile Val Ser Val Ala
50 55 60
Phe Ser Tyr Gly Asn Arg Ala Tyr Thr Gly Tyr Pro Glu Glu Ile Ser
65 70 75 80
Asp Tyr Phe Leu Gln Ser Phe Pro Glu Gly Phe Thr Tyr Glu Arg Asn
85 90 95
Ile Arg Tyr Gln Asp Gly Gly Thr Ala Ile Val Lys Ser Asp Ile Ser
100 105 110
Leu Glu Asp Gly Lys Phe Ile Val Asn Val Asp Phe Lys Ala Lys Asp
115 120 125
Leu Arg Arg Met Gly Pro Val Met Gln Gln Asp Ile Val Gly Met Gln
130 135 140
Pro Ser Tyr Glu Ser Met Tyr Thr Asn Val Thr Ser Val Ile Gly Glu
145 150 155 160
Cys Ile Ile Ala Phe Lys Leu Gln Thr Gly Lys His Phe Thr Tyr His
165 170 175
Met Arg Thr Val Tyr Lys Ser Lys Lys Pro Val Glu Thr Met Pro Leu
180 185 190
Tyr His Phe Ile Gln His Arg Leu Val Lys Thr Asn Val Asp Thr Ala
195 200 205
Ser Gly Tyr Val Val Gln His Glu Thr Ala Ile Ala Ala His Ser Thr
210 215 220
Ile Lys Lys Ile Glu Gly Ser Leu Pro
225 230
<210> 39
<211> 1539
<212> PRT
<213> 智人
<400> 39
Met Asp Ser Tyr Phe Lys Ala Ala Val Ser Asp Leu Asp Lys Leu Leu
1 5 10 15
Asp Asp Phe Glu Gln Asn Pro Asp Glu Gln Asp Tyr Leu Gln Asp Val
20 25 30
Gln Asn Ala Tyr Asp Ser Asn His Cys Ser Val Ser Ser Glu Leu Ala
35 40 45
Ser Ser Gln Arg Thr Ser Leu Leu Pro Lys Asp Gln Glu Cys Val Asn
50 55 60
Ser Cys Ala Ser Ser Glu Thr Ser Tyr Gly Thr Asn Glu Ser Ser Leu
65 70 75 80
Asn Glu Lys Thr Leu Lys Gly Leu Thr Ser Ile Gln Asn Glu Lys Asn
85 90 95
Val Thr Gly Leu Asp Leu Leu Ser Ser Val Asp Gly Gly Thr Ser Asp
100 105 110
Glu Ile Gln Pro Leu Tyr Met Gly Arg Cys Ser Lys Pro Ile Cys Asp
115 120 125
Leu Ile Ser Asp Met Gly Asn Leu Val His Ala Thr Asn Ser Glu Glu
130 135 140
Asp Ile Lys Lys Leu Leu Pro Asp Asp Phe Lys Ser Asn Ala Asp Ser
145 150 155 160
Leu Ile Gly Leu Asp Leu Ser Ser Val Ser Asp Thr Pro Cys Val Ser
165 170 175
Ser Thr Asp His Asp Ser Asp Thr Val Arg Glu Gln Gln Asn Asp Ile
180 185 190
Ser Ser Glu Leu Gln Asn Arg Glu Ile Gly Gly Ile Lys Glu Leu Gly
195 200 205
Ile Lys Val Asp Thr Thr Leu Ser Asp Ser Tyr Asn Tyr Ser Gly Thr
210 215 220
Glu Asn Leu Lys Asp Lys Lys Ile Phe Asn Gln Leu Glu Ser Ile Val
225 230 235 240
Asp Phe Asn Met Ser Ser Ala Leu Thr Arg Gln Ser Ser Lys Met Phe
245 250 255
His Ala Lys Asp Lys Leu Gln His Lys Ser Gln Pro Cys Gly Leu Leu
260 265 270
Lys Asp Val Gly Leu Val Lys Glu Glu Val Asp Val Ala Val Ile Thr
275 280 285
Ala Ala Glu Cys Leu Lys Glu Glu Gly Lys Thr Ser Ala Leu Thr Cys
290 295 300
Ser Leu Pro Lys Asn Glu Asp Leu Cys Leu Asn Asp Ser Asn Ser Arg
305 310 315 320
Asp Glu Asn Phe Lys Leu Pro Asp Phe Ser Phe Gln Glu Asp Lys Thr
325 330 335
Val Ile Lys Gln Ser Ala Gln Glu Asp Ser Lys Ser Leu Asp Leu Lys
340 345 350
Asp Asn Asp Val Ile Gln Asp Ser Ser Ser Ala Leu His Val Ser Ser
355 360 365
Lys Asp Val Pro Ser Ser Leu Ser Cys Leu Pro Ala Ser Gly Ser Met
370 375 380
Cys Gly Ser Leu Ile Glu Ser Lys Ala Arg Gly Asp Phe Leu Pro Gln
385 390 395 400
His Glu His Lys Asp Asn Ile Gln Asp Ala Val Thr Ile His Glu Glu
405 410 415
Ile Gln Asn Ser Val Val Leu Gly Gly Glu Pro Phe Lys Glu Asn Asp
420 425 430
Leu Leu Lys Gln Glu Lys Cys Lys Ser Ile Leu Leu Gln Ser Leu Ile
435 440 445
Glu Gly Met Glu Asp Arg Lys Ile Asp Pro Asp Gln Thr Val Ile Arg
450 455 460
Ala Glu Ser Leu Asp Gly Gly Asp Thr Ser Ser Thr Val Val Glu Ser
465 470 475 480
Gln Glu Gly Leu Ser Gly Thr His Val Pro Glu Ser Ser Asp Cys Cys
485 490 495
Glu Gly Phe Ile Asn Thr Phe Ser Ser Asn Asp Met Asp Gly Gln Asp
500 505 510
Leu Asp Tyr Phe Asn Ile Asp Glu Gly Ala Lys Ser Gly Pro Leu Ile
515 520 525
Ser Asp Ala Glu Leu Asp Ala Phe Leu Thr Glu Gln Tyr Leu Gln Thr
530 535 540
Thr Asn Ile Lys Ser Phe Glu Glu Asn Val Asn Asp Ser Lys Ser Gln
545 550 555 560
Met Asn Gln Ile Asp Met Lys Gly Leu Asp Asp Gly Asn Ile Asn Asn
565 570 575
Ile Tyr Phe Asn Ala Glu Ala Gly Ala Ile Gly Glu Ser His Gly Ile
580 585 590
Asn Ile Ile Cys Glu Ile Val Asp Lys Gln Asn Thr Ile Glu Asn Gly
595 600 605
Leu Ser Leu Gly Glu Lys Ser Thr Ile Pro Val Gln Gln Gly Leu Pro
610 615 620
Thr Ser Lys Ser Glu Ile Thr Asn Gln Leu Ser Val Ser Asp Ile Asn
625 630 635 640
Ser Gln Ser Val Gly Gly Ala Arg Pro Lys Gln Leu Phe Ser Leu Pro
645 650 655
Ser Arg Thr Arg Ser Ser Lys Asp Leu Asn Lys Pro Asp Val Pro Asp
660 665 670
Thr Ile Glu Ser Glu Pro Ser Thr Ala Asp Thr Val Val Pro Ile Thr
675 680 685
Cys Ala Ile Asp Ser Thr Ala Asp Pro Gln Val Ser Phe Asn Ser Asn
690 695 700
Tyr Ile Asp Ile Glu Ser Asn Ser Glu Gly Gly Ser Ser Phe Val Thr
705 710 715 720
Ala Asn Glu Asp Ser Val Pro Glu Asn Thr Cys Lys Glu Gly Leu Val
725 730 735
Leu Gly Gln Lys Gln Pro Thr Trp Val Pro Asp Ser Glu Ala Pro Asn
740 745 750
Cys Met Asn Cys Gln Val Lys Phe Thr Phe Thr Lys Arg Arg His His
755 760 765
Cys Arg Ala Cys Gly Lys Val Phe Cys Gly Val Cys Cys Asn Arg Lys
770 775 780
Cys Lys Leu Gln Tyr Leu Glu Lys Glu Ala Arg Val Cys Val Val Cys
785 790 795 800
Tyr Glu Thr Ile Ser Lys Ala Gln Ala Phe Glu Arg Met Met Ser Pro
805 810 815
Thr Gly Ser Asn Leu Lys Ser Asn His Ser Asp Glu Cys Thr Thr Val
820 825 830
Gln Pro Pro Gln Glu Asn Gln Thr Ser Ser Ile Pro Ser Pro Ala Thr
835 840 845
Leu Pro Val Ser Ala Leu Lys Gln Pro Gly Val Glu Gly Leu Cys Ser
850 855 860
Lys Glu Gln Lys Arg Val Trp Phe Ala Asp Gly Ile Leu Pro Asn Gly
865 870 875 880
Glu Val Ala Asp Thr Thr Lys Leu Ser Ser Gly Ser Lys Arg Cys Ser
885 890 895
Glu Asp Phe Ser Pro Leu Ser Pro Asp Val Pro Met Thr Val Asn Thr
900 905 910
Val Asp His Ser His Ser Thr Thr Val Glu Lys Pro Asn Asn Glu Thr
915 920 925
Gly Asp Ile Thr Arg Asn Glu Ile Ile Gln Ser Pro Ile Ser Gln Val
930 935 940
Pro Ser Val Glu Lys Leu Ser Met Asn Thr Gly Asn Glu Gly Leu Pro
945 950 955 960
Thr Ser Gly Ser Phe Thr Leu Asp Asp Asp Val Phe Ala Glu Thr Glu
965 970 975
Glu Pro Ser Ser Pro Thr Gly Val Leu Val Asn Ser Asn Leu Pro Ile
980 985 990
Ala Ser Ile Ser Asp Tyr Arg Leu Leu Cys Asp Ile Asn Lys Tyr Val
995 1000 1005
Cys Asn Lys Ile Ser Leu Leu Pro Asn Asp Glu Asp Ser Leu Pro
1010 1015 1020
Pro Leu Leu Val Ala Ser Gly Glu Lys Gly Ser Val Pro Val Val
1025 1030 1035
Glu Glu His Pro Ser His Glu Gln Ile Ile Leu Leu Leu Glu Gly
1040 1045 1050
Glu Ser Phe His Pro Val Thr Phe Val Leu Asn Ala Asn Leu Leu
1055 1060 1065
Val Asn Val Lys Phe Ile Phe Tyr Ser Ser Asp Lys Tyr Trp Tyr
1070 1075 1080
Phe Ser Thr Asn Gly Leu His Gly Leu Gly Gln Ala Glu Ile Ile
1085 1090 1095
Ile Leu Leu Leu Cys Leu Pro Asn Glu Asp Thr Ile Pro Lys Asp
1100 1105 1110
Ile Phe Arg Leu Phe Ile Thr Ile Tyr Lys Asp Ala Leu Lys Gly
1115 1120 1125
Lys Tyr Ile Glu Asn Leu Asp Asn Ile Thr Phe Thr Glu Ser Phe
1130 1135 1140
Leu Ser Ser Lys Asp His Gly Gly Phe Leu Phe Ile Thr Pro Thr
1145 1150 1155
Phe Gln Lys Leu Asp Asp Leu Ser Leu Pro Ser Asn Pro Phe Leu
1160 1165 1170
Cys Gly Ile Leu Ile Gln Lys Leu Glu Ile Pro Trp Ala Lys Val
1175 1180 1185
Phe Pro Met Arg Leu Met Leu Arg Leu Gly Ala Glu Tyr Lys Ala
1190 1195 1200
Tyr Pro Ala Pro Leu Thr Ser Ile Arg Gly Arg Lys Pro Leu Phe
1205 1210 1215
Gly Glu Ile Gly His Thr Ile Met Asn Leu Leu Val Asp Leu Arg
1220 1225 1230
Asn Tyr Gln Tyr Thr Leu His Asn Ile Asp Gln Leu Leu Ile His
1235 1240 1245
Met Glu Met Gly Lys Ser Cys Ile Lys Ile Pro Arg Lys Lys Tyr
1250 1255 1260
Ser Asp Val Met Lys Val Leu Asn Ser Ser Asn Glu His Val Ile
1265 1270 1275
Ser Ile Gly Ala Ser Phe Ser Thr Glu Ala Asp Ser His Leu Val
1280 1285 1290
Cys Ile Gln Asn Asp Gly Ile Tyr Glu Thr Gln Ala Asn Ser Ala
1295 1300 1305
Thr Gly His Pro Arg Lys Val Thr Gly Ala Ser Phe Val Val Phe
1310 1315 1320
Asn Gly Ala Leu Lys Thr Ser Ser Gly Phe Leu Ala Lys Ser Ser
1325 1330 1335
Ile Val Glu Asp Gly Leu Met Val Gln Ile Thr Pro Glu Thr Met
1340 1345 1350
Asn Gly Leu Arg Leu Ala Leu Arg Glu Gln Lys Asp Phe Lys Ile
1355 1360 1365
Thr Cys Gly Lys Val Asp Ala Val Asp Leu Arg Glu Tyr Val Asp
1370 1375 1380
Ile Cys Trp Val Asp Ala Glu Glu Lys Gly Asn Lys Gly Val Ile
1385 1390 1395
Ser Ser Val Asp Gly Ile Ser Leu Gln Gly Phe Pro Ser Glu Lys
1400 1405 1410
Ile Lys Leu Glu Ala Asp Phe Glu Thr Asp Glu Lys Ile Val Lys
1415 1420 1425
Cys Thr Glu Val Phe Tyr Phe Leu Lys Asp Gln Asp Leu Ser Ile
1430 1435 1440
Leu Ser Thr Ser Tyr Gln Phe Ala Lys Glu Ile Ala Met Ala Cys
1445 1450 1455
Ser Ala Ala Leu Cys Pro His Leu Lys Thr Leu Lys Ser Asn Gly
1460 1465 1470
Met Asn Lys Ile Gly Leu Arg Val Ser Ile Asp Thr Asp Met Val
1475 1480 1485
Glu Phe Gln Ala Gly Ser Glu Gly Gln Leu Leu Pro Gln His Tyr
1490 1495 1500
Leu Asn Asp Leu Asp Ser Ala Leu Ile Pro Val Ile His Gly Gly
1505 1510 1515
Thr Ser Asn Ser Ser Leu Pro Leu Glu Ile Glu Leu Val Phe Phe
1520 1525 1530
Ile Ile Glu His Leu Phe
1535
<210> 40
<211> 343
<212> PRT
<213> 智人
<400> 40
Met Trp Pro Asn Gly Ser Ser Leu Gly Pro Cys Phe Arg Pro Thr Asn
1 5 10 15
Ile Thr Leu Glu Glu Arg Arg Leu Ile Ala Ser Pro Trp Phe Ala Ala
20 25 30
Ser Phe Cys Val Val Gly Leu Ala Ser Asn Leu Leu Ala Leu Ser Val
35 40 45
Leu Ala Gly Ala Arg Gln Gly Gly Ser His Thr Arg Ser Ser Phe Leu
50 55 60
Thr Phe Leu Cys Gly Leu Val Leu Thr Asp Phe Leu Gly Leu Leu Val
65 70 75 80
Thr Gly Thr Ile Val Val Ser Gln His Ala Ala Leu Phe Glu Trp His
85 90 95
Ala Val Asp Pro Gly Cys Arg Leu Cys Arg Phe Met Gly Val Val Met
100 105 110
Ile Phe Phe Gly Leu Ser Pro Leu Leu Leu Gly Ala Ala Met Ala Ser
115 120 125
Glu Arg Tyr Leu Gly Ile Thr Arg Pro Phe Ser Arg Pro Ala Val Ala
130 135 140
Ser Gln Arg Arg Ala Trp Ala Thr Val Gly Leu Val Trp Ala Ala Ala
145 150 155 160
Leu Ala Leu Gly Leu Leu Pro Leu Leu Gly Val Gly Arg Tyr Thr Val
165 170 175
Gln Tyr Pro Gly Ser Trp Cys Phe Leu Thr Leu Gly Ala Glu Ser Gly
180 185 190
Asp Val Ala Phe Gly Leu Leu Phe Ser Met Leu Gly Gly Leu Ser Val
195 200 205
Gly Leu Ser Phe Leu Leu Asn Thr Val Ser Val Ala Thr Leu Cys His
210 215 220
Val Tyr His Gly Gln Glu Ala Ala Gln Gln Arg Pro Arg Asp Ser Glu
225 230 235 240
Val Glu Met Met Ala Gln Leu Leu Gly Ile Met Val Val Ala Ser Val
245 250 255
Cys Trp Leu Pro Leu Leu Val Phe Ile Ala Gln Thr Val Leu Arg Asn
260 265 270
Pro Pro Ala Met Ser Pro Ala Gly Gln Leu Ser Arg Thr Thr Glu Lys
275 280 285
Glu Leu Leu Ile Tyr Leu Arg Val Ala Thr Trp Asn Gln Ile Leu Asp
290 295 300
Pro Trp Val Tyr Ile Leu Phe Arg Arg Ala Val Leu Arg Arg Leu Gln
305 310 315 320
Pro Arg Leu Ser Thr Arg Pro Arg Ser Leu Ser Leu Gln Pro Gln Leu
325 330 335
Thr Gln Arg Ser Gly Leu Gln
340
<210> 41
<211> 20
<212> PRT
<213> 智人
<400> 41
Lys Leu Asn Pro Pro Asp Glu Ser Gly Pro Gly Cys Met Ser Cys Lys
1 5 10 15
Cys Val Leu Ser
20
<210> 42
<211> 20
<212> PRT
<213> 智人
<400> 42
Lys Leu Asn Ser Ser Asp Asp Gly Thr Gln Gly Cys Met Gly Leu Pro
1 5 10 15
Cys Val Val Met
20
<210> 43
<211> 20
<212> PRT
<213> 智人
<400> 43
Lys Ile Ser Lys Glu Glu Lys Thr Pro Gly Cys Val Lys Ile Lys Lys
1 5 10 15
Cys Ile Ile Met
20
<210> 44
<211> 20
<212> PRT
<213> 智人
<400> 44
Lys Met Ser Lys Asp Gly Lys Lys Lys Lys Lys Lys Ser Lys Thr Lys
1 5 10 15
Cys Val Ile Met
20
<210> 45
<211> 21
<212> PRT
<213> 智人
<400> 45
Lys Asn Gly Lys Lys Lys Arg Lys Ser Leu Ala Lys Arg Ile Arg Glu
1 5 10 15
Arg Cys Cys Ile Leu
20
<210> 46
<211> 1203
<212> PRT
<213> 智人
<400> 46
Met Gly Asn Leu Lys Ser Val Ala Gln Glu Pro Gly Pro Pro Cys Gly
1 5 10 15
Leu Gly Leu Gly Leu Gly Leu Gly Leu Cys Gly Lys Gln Gly Pro Ala
20 25 30
Thr Pro Ala Pro Glu Pro Ser Arg Ala Pro Ala Ser Leu Leu Pro Pro
35 40 45
Ala Pro Glu His Ser Pro Pro Ser Ser Pro Leu Thr Gln Pro Pro Glu
50 55 60
Gly Pro Lys Phe Pro Arg Val Lys Asn Trp Glu Val Gly Ser Ile Thr
65 70 75 80
Tyr Asp Thr Leu Ser Ala Gln Ala Gln Gln Asp Gly Pro Cys Thr Pro
85 90 95
Arg Arg Cys Leu Gly Ser Leu Val Phe Pro Arg Lys Leu Gln Gly Arg
100 105 110
Pro Ser Pro Gly Pro Pro Ala Pro Glu Gln Leu Leu Ser Gln Ala Arg
115 120 125
Asp Phe Ile Asn Gln Tyr Tyr Ser Ser Ile Lys Arg Ser Gly Ser Gln
130 135 140
Ala His Glu Gln Arg Leu Gln Glu Val Glu Ala Glu Val Ala Ala Thr
145 150 155 160
Gly Thr Tyr Gln Leu Arg Glu Ser Glu Leu Val Phe Gly Ala Lys Gln
165 170 175
Ala Trp Arg Asn Ala Pro Arg Cys Val Gly Arg Ile Gln Trp Gly Lys
180 185 190
Leu Gln Val Phe Asp Ala Arg Asp Cys Arg Ser Ala Gln Glu Met Phe
195 200 205
Thr Tyr Ile Cys Asn His Ile Lys Tyr Ala Thr Asn Arg Gly Asn Leu
210 215 220
Arg Ser Ala Ile Thr Val Phe Pro Gln Arg Cys Pro Gly Arg Gly Asp
225 230 235 240
Phe Arg Ile Trp Asn Ser Gln Leu Val Arg Tyr Ala Gly Tyr Arg Gln
245 250 255
Gln Asp Gly Ser Val Arg Gly Asp Pro Ala Asn Val Glu Ile Thr Glu
260 265 270
Leu Cys Ile Gln His Gly Trp Thr Pro Gly Asn Gly Arg Phe Asp Val
275 280 285
Leu Pro Leu Leu Leu Gln Ala Pro Asp Asp Pro Pro Glu Leu Phe Leu
290 295 300
Leu Pro Pro Glu Leu Val Leu Glu Val Pro Leu Glu His Pro Thr Leu
305 310 315 320
Glu Trp Phe Ala Ala Leu Gly Leu Arg Trp Tyr Ala Leu Pro Ala Val
325 330 335
Ser Asn Met Leu Leu Glu Ile Gly Gly Leu Glu Phe Pro Ala Ala Pro
340 345 350
Phe Ser Gly Trp Tyr Met Ser Thr Glu Ile Gly Thr Arg Asn Leu Cys
355 360 365
Asp Pro His Arg Tyr Asn Ile Leu Glu Asp Val Ala Val Cys Met Asp
370 375 380
Leu Asp Thr Arg Thr Thr Ser Ser Leu Trp Lys Asp Lys Ala Ala Val
385 390 395 400
Glu Ile Asn Val Ala Val Leu His Ser Tyr Gln Leu Ala Lys Val Thr
405 410 415
Ile Val Asp His His Ala Ala Thr Ala Ser Phe Met Lys His Leu Glu
420 425 430
Asn Glu Gln Lys Ala Arg Gly Gly Cys Pro Ala Asp Trp Ala Trp Ile
435 440 445
Val Pro Pro Ile Ser Gly Ser Leu Thr Pro Val Phe His Gln Glu Met
450 455 460
Val Asn Tyr Phe Leu Ser Pro Ala Phe Arg Tyr Gln Pro Asp Pro Trp
465 470 475 480
Lys Gly Ser Ala Ala Lys Gly Thr Gly Ile Thr Arg Lys Lys Thr Phe
485 490 495
Lys Glu Val Ala Asn Ala Val Lys Ile Ser Ala Ser Leu Met Gly Thr
500 505 510
Val Met Ala Lys Arg Val Lys Ala Thr Ile Leu Tyr Gly Ser Glu Thr
515 520 525
Gly Arg Ala Gln Ser Tyr Ala Gln Gln Leu Gly Arg Leu Phe Arg Lys
530 535 540
Ala Phe Asp Pro Arg Val Leu Cys Met Asp Glu Tyr Asp Val Val Ser
545 550 555 560
Leu Glu His Glu Thr Leu Val Leu Val Val Thr Ser Thr Phe Gly Asn
565 570 575
Gly Asp Pro Pro Glu Asn Gly Glu Ser Phe Ala Ala Ala Leu Met Glu
580 585 590
Met Ser Gly Pro Tyr Asn Ser Ser Pro Arg Pro Glu Gln His Lys Ser
595 600 605
Tyr Lys Ile Arg Phe Asn Ser Ile Ser Cys Ser Asp Pro Leu Val Ser
610 615 620
Ser Trp Arg Arg Lys Arg Lys Glu Ser Ser Asn Thr Asp Ser Ala Gly
625 630 635 640
Ala Leu Gly Thr Leu Arg Phe Cys Val Phe Gly Leu Gly Ser Arg Ala
645 650 655
Tyr Pro His Phe Cys Ala Phe Ala Arg Ala Val Asp Thr Arg Leu Glu
660 665 670
Glu Leu Gly Gly Glu Arg Leu Leu Gln Leu Gly Gln Gly Asp Glu Leu
675 680 685
Cys Gly Gln Glu Glu Ala Phe Arg Gly Trp Ala Gln Ala Ala Phe Gln
690 695 700
Ala Ala Cys Glu Thr Phe Cys Val Gly Glu Asp Ala Lys Ala Ala Ala
705 710 715 720
Arg Asp Ile Phe Ser Pro Lys Arg Ser Trp Lys Arg Gln Arg Tyr Arg
725 730 735
Leu Ser Ala Gln Ala Glu Gly Leu Gln Leu Leu Pro Gly Leu Ile His
740 745 750
Val His Arg Arg Lys Met Phe Gln Ala Thr Ile Arg Ser Val Glu Asn
755 760 765
Leu Gln Ser Ser Lys Ser Thr Arg Ala Thr Ile Leu Val Arg Leu Asp
770 775 780
Thr Gly Gly Gln Glu Gly Leu Gln Tyr Gln Pro Gly Asp His Ile Gly
785 790 795 800
Val Cys Pro Pro Asn Arg Pro Gly Leu Val Glu Ala Leu Leu Ser Arg
805 810 815
Val Glu Asp Pro Pro Ala Pro Thr Glu Pro Val Ala Val Glu Gln Leu
820 825 830
Glu Lys Gly Ser Pro Gly Gly Pro Pro Pro Gly Trp Val Arg Asp Pro
835 840 845
Arg Leu Pro Pro Cys Thr Leu Arg Gln Ala Leu Thr Phe Phe Leu Asp
850 855 860
Ile Thr Ser Pro Pro Ser Pro Gln Leu Leu Arg Leu Leu Ser Thr Leu
865 870 875 880
Ala Glu Glu Pro Arg Glu Gln Gln Glu Leu Glu Ala Leu Ser Gln Asp
885 890 895
Pro Arg Arg Tyr Glu Glu Trp Lys Trp Phe Arg Cys Pro Thr Leu Leu
900 905 910
Glu Val Leu Glu Gln Phe Pro Ser Val Ala Leu Pro Ala Pro Leu Leu
915 920 925
Leu Thr Gln Leu Pro Leu Leu Gln Pro Arg Tyr Tyr Ser Val Ser Ser
930 935 940
Ala Pro Ser Thr His Pro Gly Glu Ile His Leu Thr Val Ala Val Leu
945 950 955 960
Ala Tyr Arg Thr Gln Asp Gly Leu Gly Pro Leu His Tyr Gly Val Cys
965 970 975
Ser Thr Trp Leu Ser Gln Leu Lys Pro Gly Asp Pro Val Pro Cys Phe
980 985 990
Ile Arg Gly Ala Pro Ser Phe Arg Leu Pro Pro Asp Pro Ser Leu Pro
995 1000 1005
Cys Ile Leu Val Gly Pro Gly Thr Gly Ile Ala Pro Phe Arg Gly
1010 1015 1020
Phe Trp Gln Glu Arg Leu His Asp Ile Glu Ser Lys Gly Leu Gln
1025 1030 1035
Pro Thr Pro Met Thr Leu Val Phe Gly Cys Arg Cys Ser Gln Leu
1040 1045 1050
Asp His Leu Tyr Arg Asp Glu Val Gln Asn Ala Gln Gln Arg Gly
1055 1060 1065
Val Phe Gly Arg Val Leu Thr Ala Phe Ser Arg Glu Pro Asp Asn
1070 1075 1080
Pro Lys Thr Tyr Val Gln Asp Ile Leu Arg Thr Glu Leu Ala Ala
1085 1090 1095
Glu Val His Arg Val Leu Cys Leu Glu Arg Gly His Met Phe Val
1100 1105 1110
Cys Gly Asp Val Thr Met Ala Thr Asn Val Leu Gln Thr Val Gln
1115 1120 1125
Arg Ile Leu Ala Thr Glu Gly Asp Met Glu Leu Asp Glu Ala Gly
1130 1135 1140
Asp Val Ile Gly Val Leu Arg Asp Gln Gln Arg Tyr His Glu Asp
1145 1150 1155
Ile Phe Gly Leu Thr Leu Arg Thr Gln Glu Val Thr Ser Arg Ile
1160 1165 1170
Arg Thr Gln Ser Phe Ser Leu Gln Glu Arg Gln Leu Arg Gly Ala
1175 1180 1185
Val Pro Trp Ala Phe Asp Pro Pro Gly Ser Asp Thr Asn Ser Pro
1190 1195 1200
<210> 47
<211> 663
<212> PRT
<213> 智人
<400> 47
Met Ile Glu Lys Met Gln Gly Ser Arg Met Asp Glu Gln Arg Cys Ser
1 5 10 15
Phe Pro Pro Pro Leu Lys Thr Glu Glu Asp Tyr Ile Pro Tyr Pro Ser
20 25 30
Val His Glu Val Leu Gly Arg Glu Gly Pro Phe Pro Leu Ile Leu Leu
35 40 45
Pro Gln Phe Gly Gly Tyr Trp Ile Glu Gly Thr Asn His Glu Ile Thr
50 55 60
Ser Ile Pro Glu Thr Glu Pro Leu Gln Ser Pro Thr Thr Lys Val Lys
65 70 75 80
Leu Glu Cys Asn Pro Thr Ala Arg Ile Tyr Arg Lys His Phe Leu Gly
85 90 95
Lys Glu His Phe Asn Tyr Tyr Ser Leu Asp Ala Ala Leu Gly His Leu
100 105 110
Val Phe Ser Leu Lys Tyr Asp Val Ile Gly Asp Gln Glu His Leu Arg
115 120 125
Leu Leu Leu Arg Thr Lys Cys Arg Thr Tyr His Asp Val Ile Pro Ile
130 135 140
Ser Cys Leu Thr Glu Phe Pro Asn Val Val Gln Met Ala Lys Leu Val
145 150 155 160
Cys Glu Asp Val Asn Val Asp Arg Phe Tyr Pro Val Leu Tyr Pro Lys
165 170 175
Ala Ser Arg Leu Ile Val Thr Phe Asp Glu His Val Ile Ser Asn Asn
180 185 190
Phe Lys Phe Gly Val Ile Tyr Gln Lys Leu Gly Gln Thr Ser Glu Glu
195 200 205
Glu Leu Phe Ser Thr Asn Glu Glu Ser Pro Ala Phe Val Glu Phe Leu
210 215 220
Glu Phe Leu Gly Gln Lys Val Lys Leu Gln Asp Phe Lys Gly Phe Arg
225 230 235 240
Gly Gly Leu Asp Val Thr His Gly Gln Thr Gly Thr Glu Ser Val Tyr
245 250 255
Cys Asn Phe Arg Asn Lys Glu Ile Met Phe His Val Ser Thr Lys Leu
260 265 270
Pro Tyr Thr Glu Gly Asp Ala Gln Gln Leu Gln Arg Lys Arg His Ile
275 280 285
Gly Asn Asp Ile Val Ala Val Val Phe Gln Asp Glu Asn Thr Pro Phe
290 295 300
Val Pro Asp Met Ile Ala Ser Asn Phe Leu His Ala Tyr Val Val Val
305 310 315 320
Gln Ala Glu Gly Gly Gly Pro Asp Gly Pro Leu Tyr Lys Val Ser Val
325 330 335
Thr Ala Arg Asp Asp Val Pro Phe Phe Gly Pro Pro Leu Pro Asp Pro
340 345 350
Ala Val Phe Arg Lys Gly Pro Glu Phe Gln Glu Phe Leu Leu Thr Lys
355 360 365
Leu Ile Asn Ala Glu Tyr Ala Cys Tyr Lys Ala Glu Lys Phe Ala Lys
370 375 380
Leu Glu Glu Arg Thr Arg Ala Ala Leu Leu Glu Thr Leu Tyr Glu Glu
385 390 395 400
Leu His Ile His Ser Gln Ser Met Met Gly Leu Gly Gly Asp Glu Asp
405 410 415
Lys Met Glu Asn Gly Ser Gly Gly Gly Gly Phe Phe Glu Ser Phe Lys
420 425 430
Arg Val Ile Arg Ser Arg Ser Gln Ser Met Asp Ala Met Gly Leu Ser
435 440 445
Asn Lys Lys Pro Asn Thr Val Ser Thr Ser His Ser Gly Ser Phe Ala
450 455 460
Pro Asn Asn Pro Asp Leu Ala Lys Ala Ala Gly Ile Ser Leu Ile Val
465 470 475 480
Pro Gly Lys Ser Pro Thr Arg Lys Lys Ser Gly Pro Phe Gly Ser Arg
485 490 495
Arg Ser Ser Ala Ile Gly Ile Glu Asn Ile Gln Glu Val Gln Glu Lys
500 505 510
Arg Glu Ser Pro Pro Ala Gly Gln Lys Thr Pro Asp Ser Gly His Val
515 520 525
Ser Gln Glu Pro Lys Ser Glu Asn Ser Ser Thr Gln Ser Ser Pro Glu
530 535 540
Met Pro Thr Thr Lys Asn Arg Ala Glu Thr Ala Ala Gln Arg Ala Glu
545 550 555 560
Ala Leu Lys Asp Phe Ser Arg Ser Ser Ser Ser Ala Ser Ser Phe Ala
565 570 575
Ser Val Val Glu Glu Thr Glu Gly Val Asp Gly Glu Asp Thr Gly Leu
580 585 590
Glu Ser Val Ser Ser Ser Gly Thr Pro His Lys Arg Asp Ser Phe Ile
595 600 605
Tyr Ser Thr Trp Leu Glu Asp Ser Val Ser Thr Thr Ser Gly Gly Ser
610 615 620
Ser Pro Gly Pro Ser Arg Ser Pro His Pro Asp Ala Gly Lys Leu Gly
625 630 635 640
Asp Pro Ala Cys Pro Glu Ile Lys Ile Gln Leu Glu Ala Ser Glu Gln
645 650 655
His Met Pro Gln Leu Gly Cys
660
<210> 48
<211> 198
<212> PRT
<213> 智人
<400> 48
Met His Ser Glu Ala Glu Glu Ser Lys Glu Val Ala Thr Asp Val Phe
1 5 10 15
Asn Ser Lys Asn Leu Ala Val Gln Ala Gln Lys Lys Ile Leu Gly Lys
20 25 30
Met Val Ser Lys Ser Ile Ala Thr Thr Leu Ile Asp Asp Thr Ser Ser
35 40 45
Glu Val Leu Asp Glu Leu Tyr Arg Val Thr Arg Glu Tyr Thr Gln Asn
50 55 60
Lys Lys Glu Ala Glu Lys Ile Ile Lys Asn Leu Ile Lys Thr Val Ile
65 70 75 80
Lys Leu Ala Ile Leu Tyr Arg Asn Asn Gln Phe Asn Gln Asp Glu Leu
85 90 95
Ala Leu Met Glu Lys Phe Lys Lys Lys Val His Gln Leu Ala Met Thr
100 105 110
Val Val Ser Phe His Gln Val Asp Tyr Thr Phe Asp Arg Asn Val Leu
115 120 125
Ser Arg Leu Leu Asn Glu Cys Arg Glu Met Leu His Gln Ile Ile Gln
130 135 140
Arg His Leu Thr Ala Lys Ser His Gly Arg Val Asn Asn Val Phe Asp
145 150 155 160
His Phe Ser Asp Cys Glu Phe Leu Ala Ala Leu Tyr Asn Pro Phe Gly
165 170 175
Asn Phe Lys Pro His Leu Gln Lys Leu Cys Asp Gly Ile Asn Lys Met
180 185 190
Leu Asp Glu Glu Asn Ile
195
<210> 49
<211> 1544
<212> PRT
<213> 智人
<400> 49
Met Ser Gly Thr Gln Ser Thr Ile Thr Asp Arg Phe Pro Leu Lys Lys
1 5 10 15
Pro Ile Arg His Gly Ser Ile Leu Asn Arg Glu Ser Pro Thr Asp Lys
20 25 30
Lys Gln Lys Val Glu Arg Ile Ala Ser His Asp Phe Asp Pro Thr Asp
35 40 45
Ser Ser Ser Lys Lys Thr Lys Ser Ser Ser Glu Glu Ser Arg Ser Glu
50 55 60
Ile Tyr Gly Leu Val Gln Arg Cys Val Ile Ile Gln Lys Asp Asp Asn
65 70 75 80
Gly Phe Gly Leu Thr Val Ser Gly Asp Asn Pro Val Phe Val Gln Ser
85 90 95
Val Lys Glu Asp Gly Ala Ala Met Arg Ala Gly Val Gln Thr Gly Asp
100 105 110
Arg Ile Ile Lys Val Asn Gly Thr Leu Val Thr His Ser Asn His Leu
115 120 125
Glu Val Val Lys Leu Ile Lys Ser Gly Ser Tyr Val Ala Leu Thr Val
130 135 140
Gln Gly Arg Pro Pro Gly Ser Pro Gln Ile Pro Leu Ala Asp Ser Glu
145 150 155 160
Val Glu Pro Ser Val Ile Gly His Met Ser Pro Ile Met Thr Ser Pro
165 170 175
His Ser Pro Gly Ala Ser Gly Asn Met Glu Arg Ile Thr Ser Pro Val
180 185 190
Leu Met Gly Glu Glu Asn Asn Val Val His Asn Gln Lys Val Glu Ile
195 200 205
Leu Arg Lys Met Leu Gln Lys Glu Gln Glu Arg Leu Gln Leu Leu Gln
210 215 220
Glu Asp Tyr Asn Arg Thr Pro Ala Gln Arg Leu Leu Lys Glu Ile Gln
225 230 235 240
Glu Ala Lys Lys His Ile Pro Gln Leu Gln Glu Gln Leu Ser Lys Ala
245 250 255
Thr Gly Ser Ala Gln Asp Gly Ala Val Val Thr Pro Ser Arg Pro Leu
260 265 270
Gly Asp Thr Leu Thr Val Ser Glu Ala Glu Thr Asp Pro Gly Asp Val
275 280 285
Leu Gly Arg Thr Asp Cys Ser Ser Gly Asp Ala Ser Arg Pro Ser Ser
290 295 300
Asp Asn Ala Asp Ser Pro Lys Ser Gly Pro Lys Glu Arg Ile Tyr Leu
305 310 315 320
Glu Glu Asn Pro Glu Lys Ser Glu Thr Ile Gln Asp Thr Asp Thr Gln
325 330 335
Ser Leu Val Gly Ser Pro Ser Thr Arg Ile Ala Pro His Ile Ile Gly
340 345 350
Ala Glu Asp Asp Asp Phe Gly Thr Glu His Glu Gln Ile Asn Gly Gln
355 360 365
Cys Ser Cys Phe Gln Ser Ile Glu Leu Leu Lys Ser Arg Pro Ala His
370 375 380
Leu Ala Val Phe Leu His His Val Val Ser Gln Phe Asp Pro Ala Thr
385 390 395 400
Leu Leu Cys Tyr Leu Tyr Ser Asp Leu Tyr Lys His Thr Asn Ser Lys
405 410 415
Glu Thr Arg Arg Ile Phe Leu Glu Phe His Gln Phe Phe Leu Asp Arg
420 425 430
Ser Ala His Leu Lys Val Ser Val Pro Asp Glu Met Ser Ala Asp Leu
435 440 445
Glu Lys Arg Arg Pro Glu Leu Ile Pro Glu Asp Leu His Arg His Tyr
450 455 460
Ile Gln Thr Met Gln Glu Arg Val His Pro Glu Val Gln Arg His Leu
465 470 475 480
Glu Asp Phe Arg Gln Lys Arg Ser Met Gly Leu Thr Leu Ala Glu Ser
485 490 495
Glu Leu Thr Lys Leu Asp Ala Glu Arg Asp Lys Asp Arg Leu Thr Leu
500 505 510
Glu Lys Glu Arg Thr Cys Ala Glu Gln Ile Val Ala Lys Ile Glu Glu
515 520 525
Val Leu Met Thr Ala Gln Ala Val Glu Glu Asp Lys Ser Ser Thr Met
530 535 540
Gln Tyr Val Ile Leu Met Tyr Met Lys His Leu Gly Val Lys Val Lys
545 550 555 560
Glu Pro Arg Asn Leu Glu His Lys Arg Gly Arg Ile Gly Phe Leu Pro
565 570 575
Lys Ile Lys Gln Ser Met Lys Lys Asp Lys Glu Gly Glu Glu Lys Gly
580 585 590
Lys Arg Arg Gly Phe Pro Ser Ile Leu Gly Pro Pro Arg Arg Pro Ser
595 600 605
Arg His Asp Asn Ser Ala Ile Gly Arg Ala Met Glu Leu Gln Lys Ala
610 615 620
Arg His Pro Lys His Leu Ser Thr Pro Ser Ser Val Ser Pro Glu Pro
625 630 635 640
Gln Asp Ser Ala Lys Leu Arg Gln Ser Gly Leu Ala Asn Glu Gly Thr
645 650 655
Asp Ala Gly Tyr Leu Pro Ala Asn Ser Met Ser Ser Val Ala Ser Gly
660 665 670
Ala Ser Phe Ser Gln Glu Gly Gly Lys Glu Asn Asp Thr Gly Ser Lys
675 680 685
Gln Val Gly Glu Thr Ser Ala Pro Gly Asp Thr Leu Asp Gly Thr Pro
690 695 700
Arg Thr Leu Asn Thr Val Phe Asp Phe Pro Pro Pro Pro Leu Asp Gln
705 710 715 720
Val Gln Glu Glu Glu Cys Glu Val Glu Arg Val Thr Glu His Gly Thr
725 730 735
Pro Lys Pro Phe Arg Lys Phe Asp Ser Val Ala Phe Gly Glu Ser Gln
740 745 750
Ser Glu Asp Glu Gln Phe Glu Asn Asp Leu Glu Thr Asp Pro Pro Asn
755 760 765
Trp Gln Gln Leu Val Ser Arg Glu Val Leu Leu Gly Leu Lys Pro Cys
770 775 780
Glu Ile Lys Arg Gln Glu Val Ile Asn Glu Leu Phe Tyr Thr Glu Arg
785 790 795 800
Ala His Val Arg Thr Leu Lys Val Leu Asp Gln Val Phe Tyr Gln Arg
805 810 815
Val Ser Arg Glu Gly Ile Leu Ser Pro Ser Glu Leu Arg Lys Ile Phe
820 825 830
Ser Asn Leu Glu Asp Ile Leu Gln Leu His Ile Gly Leu Asn Glu Gln
835 840 845
Met Lys Ala Val Arg Lys Arg Asn Glu Thr Ser Val Ile Asp Gln Ile
850 855 860
Gly Glu Asp Leu Leu Thr Trp Phe Ser Gly Pro Gly Glu Glu Lys Leu
865 870 875 880
Lys His Ala Ala Ala Thr Phe Cys Ser Asn Gln Pro Phe Ala Leu Glu
885 890 895
Met Ile Lys Ser Arg Gln Lys Lys Asp Ser Arg Phe Gln Thr Phe Val
900 905 910
Gln Asp Ala Glu Ser Asn Pro Leu Cys Arg Arg Leu Gln Leu Lys Asp
915 920 925
Ile Ile Pro Thr Gln Met Gln Arg Leu Thr Lys Tyr Pro Leu Leu Leu
930 935 940
Asp Asn Ile Ala Lys Tyr Thr Glu Trp Pro Thr Glu Arg Glu Lys Val
945 950 955 960
Lys Lys Ala Ala Asp His Cys Arg Gln Ile Leu Asn Tyr Val Asn Gln
965 970 975
Ala Val Lys Glu Ala Glu Asn Lys Gln Arg Leu Glu Asp Tyr Gln Arg
980 985 990
Arg Leu Asp Thr Ser Ser Leu Lys Leu Ser Glu Tyr Pro Asn Val Glu
995 1000 1005
Glu Leu Arg Asn Leu Asp Leu Thr Lys Arg Lys Met Ile His Glu
1010 1015 1020
Gly Pro Leu Val Trp Lys Val Asn Arg Asp Lys Thr Ile Asp Leu
1025 1030 1035
Tyr Thr Leu Leu Leu Glu Asp Ile Leu Val Leu Leu Gln Lys Gln
1040 1045 1050
Asp Asp Arg Leu Val Leu Arg Cys His Ser Lys Ile Leu Ala Ser
1055 1060 1065
Thr Ala Asp Ser Lys His Thr Phe Ser Pro Val Ile Lys Leu Ser
1070 1075 1080
Thr Val Leu Val Arg Gln Val Ala Thr Asp Asn Lys Ala Leu Phe
1085 1090 1095
Val Ile Ser Met Ser Asp Asn Gly Ala Gln Ile Tyr Glu Leu Val
1100 1105 1110
Ala Gln Thr Val Ser Glu Lys Thr Val Trp Gln Asp Leu Ile Cys
1115 1120 1125
Arg Met Ala Ala Ser Val Lys Glu Gln Ser Thr Lys Pro Ile Pro
1130 1135 1140
Leu Pro Gln Ser Thr Pro Gly Glu Gly Asp Asn Asp Glu Glu Asp
1145 1150 1155
Pro Ser Lys Leu Lys Glu Glu Gln His Gly Ile Ser Val Thr Gly
1160 1165 1170
Leu Gln Ser Pro Asp Arg Asp Leu Gly Leu Glu Ser Thr Leu Ile
1175 1180 1185
Ser Ser Lys Pro Gln Ser His Ser Leu Ser Thr Ser Gly Lys Ser
1190 1195 1200
Glu Val Arg Asp Leu Phe Val Ala Glu Arg Gln Phe Ala Lys Glu
1205 1210 1215
Gln His Thr Asp Gly Thr Leu Lys Glu Val Gly Glu Asp Tyr Gln
1220 1225 1230
Ile Ala Ile Pro Asp Ser His Leu Pro Val Ser Glu Glu Arg Trp
1235 1240 1245
Ala Leu Asp Ala Leu Arg Asn Leu Gly Leu Leu Lys Gln Leu Leu
1250 1255 1260
Val Gln Gln Leu Gly Leu Thr Glu Lys Ser Val Gln Glu Asp Trp
1265 1270 1275
Gln His Phe Pro Arg Tyr Arg Thr Ala Ser Gln Gly Pro Gln Thr
1280 1285 1290
Asp Ser Val Ile Gln Asn Ser Glu Asn Ile Lys Ala Tyr His Ser
1295 1300 1305
Gly Glu Gly His Met Pro Phe Arg Thr Gly Thr Gly Asp Ile Ala
1310 1315 1320
Thr Cys Tyr Ser Pro Arg Thr Ser Thr Glu Ser Phe Ala Pro Arg
1325 1330 1335
Asp Ser Val Gly Leu Ala Pro Gln Asp Ser Gln Ala Ser Asn Ile
1340 1345 1350
Leu Val Met Asp His Met Ile Met Thr Pro Glu Met Pro Thr Met
1355 1360 1365
Glu Pro Glu Gly Gly Leu Asp Asp Ser Gly Glu His Phe Phe Asp
1370 1375 1380
Ala Arg Glu Ala His Ser Asp Glu Asn Pro Ser Glu Gly Asp Gly
1385 1390 1395
Ala Val Asn Lys Glu Glu Lys Asp Val Asn Leu Arg Ile Ser Gly
1400 1405 1410
Asn Tyr Leu Ile Leu Asp Gly Tyr Asp Pro Val Gln Glu Ser Ser
1415 1420 1425
Thr Asp Glu Glu Val Ala Ser Ser Leu Thr Leu Gln Pro Met Thr
1430 1435 1440
Gly Ile Pro Ala Val Glu Ser Thr His Gln Gln Gln His Ser Pro
1445 1450 1455
Gln Asn Thr His Ser Asp Gly Ala Ile Ser Pro Phe Thr Pro Glu
1460 1465 1470
Phe Leu Val Gln Gln Arg Trp Gly Ala Met Glu Tyr Ser Cys Phe
1475 1480 1485
Glu Ile Gln Ser Pro Ser Ser Cys Ala Asp Ser Gln Ser Gln Ile
1490 1495 1500
Met Glu Tyr Ile His Lys Ile Glu Ala Asp Leu Glu His Leu Lys
1505 1510 1515
Lys Val Glu Glu Ser Tyr Thr Ile Leu Cys Gln Arg Leu Ala Gly
1520 1525 1530
Ser Ala Leu Thr Asp Lys His Ser Asp Lys Ser
1535 1540
<210> 50
<211> 862
<212> PRT
<213> 智人
<400> 50
Met Asn Ile Gln Glu Gln Gly Phe Pro Leu Asp Leu Gly Ala Ser Phe
1 5 10 15
Thr Glu Asp Ala Pro Arg Pro Pro Val Pro Gly Glu Glu Gly Glu Leu
20 25 30
Val Ser Thr Asp Pro Arg Pro Ala Ser Tyr Ser Phe Cys Ser Gly Lys
35 40 45
Gly Val Gly Ile Lys Gly Glu Thr Ser Thr Ala Thr Pro Arg Arg Ser
50 55 60
Asp Leu Asp Leu Gly Tyr Glu Pro Glu Gly Ser Ala Ser Pro Thr Pro
65 70 75 80
Pro Tyr Leu Lys Trp Ala Glu Ser Leu His Ser Leu Leu Asp Asp Gln
85 90 95
Asp Gly Ile Ser Leu Phe Arg Thr Phe Leu Lys Gln Glu Gly Cys Ala
100 105 110
Asp Leu Leu Asp Phe Trp Phe Ala Cys Thr Gly Phe Arg Lys Leu Glu
115 120 125
Pro Cys Asp Ser Asn Glu Glu Lys Arg Leu Lys Leu Ala Arg Ala Ile
130 135 140
Tyr Arg Lys Tyr Ile Leu Asp Asn Asn Gly Ile Val Ser Arg Gln Thr
145 150 155 160
Lys Pro Ala Thr Lys Ser Phe Ile Lys Gly Cys Ile Met Lys Gln Leu
165 170 175
Ile Asp Pro Ala Met Phe Asp Gln Ala Gln Thr Glu Ile Gln Ala Thr
180 185 190
Met Glu Glu Asn Thr Tyr Pro Ser Phe Leu Lys Ser Asp Ile Tyr Leu
195 200 205
Glu Tyr Thr Arg Thr Gly Ser Glu Ser Pro Lys Val Cys Ser Asp Gln
210 215 220
Ser Ser Gly Ser Gly Thr Gly Lys Gly Ile Ser Gly Tyr Leu Pro Thr
225 230 235 240
Leu Asn Glu Asp Glu Glu Trp Lys Cys Asp Gln Asp Met Asp Glu Asp
245 250 255
Asp Gly Arg Asp Ala Ala Pro Pro Gly Arg Leu Pro Gln Lys Leu Leu
260 265 270
Leu Glu Thr Ala Ala Pro Arg Val Ser Ser Ser Arg Arg Tyr Ser Glu
275 280 285
Gly Arg Glu Phe Arg Tyr Gly Ser Trp Arg Glu Pro Val Asn Pro Tyr
290 295 300
Tyr Val Asn Ala Gly Tyr Ala Leu Ala Pro Ala Thr Ser Ala Asn Asp
305 310 315 320
Ser Glu Gln Gln Ser Leu Ser Ser Asp Ala Asp Thr Leu Ser Leu Thr
325 330 335
Asp Ser Ser Val Asp Gly Ile Pro Pro Tyr Arg Ile Arg Lys Gln His
340 345 350
Arg Arg Glu Met Gln Glu Ser Val Gln Val Asn Gly Arg Val Pro Leu
355 360 365
Pro His Ile Pro Arg Thr Tyr Arg Val Pro Lys Glu Val Arg Val Glu
370 375 380
Pro Gln Lys Phe Ala Glu Glu Leu Ile His Arg Leu Glu Ala Val Gln
385 390 395 400
Arg Thr Arg Glu Ala Glu Glu Lys Leu Glu Glu Arg Leu Lys Arg Val
405 410 415
Arg Met Glu Glu Glu Gly Glu Asp Gly Asp Pro Ser Ser Gly Pro Pro
420 425 430
Gly Pro Cys His Lys Leu Pro Pro Ala Pro Ala Trp His His Phe Pro
435 440 445
Pro Arg Cys Val Asp Met Gly Cys Ala Gly Leu Arg Asp Ala His Glu
450 455 460
Glu Asn Pro Glu Ser Ile Leu Asp Glu His Val Gln Arg Val Leu Arg
465 470 475 480
Thr Pro Gly Arg Gln Ser Pro Gly Pro Gly His Arg Ser Pro Asp Ser
485 490 495
Gly His Val Ala Lys Met Pro Val Ala Leu Gly Gly Ala Ala Ser Gly
500 505 510
His Gly Lys His Val Pro Lys Ser Gly Ala Lys Leu Asp Ala Ala Gly
515 520 525
Leu His His His Arg His Val His His His Val His His Ser Thr Ala
530 535 540
Arg Pro Lys Glu Gln Val Glu Ala Glu Ala Thr Arg Arg Ala Gln Ser
545 550 555 560
Ser Phe Ala Trp Gly Leu Glu Pro His Ser His Gly Ala Arg Ser Arg
565 570 575
Gly Tyr Ser Glu Ser Val Gly Ala Ala Pro Asn Ala Ser Asp Gly Leu
580 585 590
Ala His Ser Gly Lys Val Gly Val Ala Cys Lys Arg Asn Ala Lys Lys
595 600 605
Ala Glu Ser Gly Lys Ser Ala Ser Thr Glu Val Pro Gly Ala Ser Glu
610 615 620
Asp Ala Glu Lys Asn Gln Lys Ile Met Gln Trp Ile Ile Glu Gly Glu
625 630 635 640
Lys Glu Ile Ser Arg His Arg Arg Thr Gly His Gly Ser Ser Gly Thr
645 650 655
Arg Lys Pro Gln Pro His Glu Asn Ser Arg Pro Leu Ser Leu Glu His
660 665 670
Pro Trp Ala Gly Pro Gln Leu Arg Thr Ser Val Gln Pro Ser His Leu
675 680 685
Phe Ile Gln Asp Pro Thr Met Pro Pro His Pro Ala Pro Asn Pro Leu
690 695 700
Thr Gln Leu Glu Glu Ala Arg Arg Arg Leu Glu Glu Glu Glu Lys Arg
705 710 715 720
Ala Ser Arg Ala Pro Ser Lys Gln Arg Tyr Val Gln Glu Val Met Arg
725 730 735
Arg Gly Arg Ala Cys Val Arg Pro Ala Cys Ala Pro Val Leu His Val
740 745 750
Val Pro Ala Val Ser Asp Met Glu Leu Ser Glu Thr Glu Thr Arg Ser
755 760 765
Gln Arg Lys Val Gly Gly Gly Ser Ala Gln Pro Cys Asp Ser Ile Val
770 775 780
Val Ala Tyr Tyr Phe Cys Gly Glu Pro Ile Pro Tyr Arg Thr Leu Val
785 790 795 800
Arg Gly Arg Ala Val Thr Leu Gly Gln Phe Lys Glu Leu Leu Thr Lys
805 810 815
Lys Gly Ser Tyr Arg Tyr Tyr Phe Lys Lys Val Ser Asp Glu Phe Asp
820 825 830
Cys Gly Val Val Phe Glu Glu Val Arg Glu Asp Glu Ala Val Leu Pro
835 840 845
Val Phe Glu Glu Lys Ile Ile Gly Lys Val Glu Lys Val Asp
850 855 860
<210> 51
<211> 295
<212> PRT
<213> 智人
<400> 51
Met Asp Asn Ser Gly Lys Glu Ala Glu Ala Met Ala Leu Leu Ala Glu
1 5 10 15
Ala Glu Arg Lys Val Lys Asn Ser Gln Ser Phe Phe Ser Gly Leu Phe
20 25 30
Gly Gly Ser Ser Lys Ile Glu Glu Ala Cys Glu Ile Tyr Ala Arg Ala
35 40 45
Ala Asn Met Phe Lys Met Ala Lys Asn Trp Ser Ala Ala Gly Asn Ala
50 55 60
Phe Cys Gln Ala Ala Gln Leu His Leu Gln Leu Gln Ser Lys His Asp
65 70 75 80
Ala Ala Thr Cys Phe Val Asp Ala Gly Asn Ala Phe Lys Lys Ala Asp
85 90 95
Pro Gln Glu Ala Ile Asn Cys Leu Met Arg Ala Ile Glu Ile Tyr Thr
100 105 110
Asp Met Gly Arg Phe Thr Ile Ala Ala Lys His His Ile Ser Ile Ala
115 120 125
Glu Ile Tyr Glu Thr Glu Leu Val Asp Ile Glu Lys Ala Ile Ala His
130 135 140
Tyr Glu Gln Ser Ala Asp Tyr Tyr Lys Gly Glu Glu Ser Asn Ser Ser
145 150 155 160
Ala Asn Lys Cys Leu Leu Lys Val Ala Gly Tyr Ala Ala Leu Leu Glu
165 170 175
Gln Tyr Gln Lys Ala Ile Asp Ile Tyr Glu Gln Val Gly Thr Asn Ala
180 185 190
Met Asp Ser Pro Leu Leu Lys Tyr Ser Ala Lys Asp Tyr Phe Phe Lys
195 200 205
Ala Ala Leu Cys His Phe Cys Ile Asp Met Leu Asn Ala Lys Leu Ala
210 215 220
Val Gln Lys Tyr Glu Glu Leu Phe Pro Ala Phe Ser Asp Ser Arg Glu
225 230 235 240
Cys Lys Leu Met Lys Lys Leu Leu Glu Ala His Glu Glu Gln Asn Val
245 250 255
Asp Ser Tyr Thr Glu Ser Val Lys Glu Tyr Asp Ser Ile Ser Arg Leu
260 265 270
Asp Gln Trp Leu Thr Thr Met Leu Leu Arg Ile Lys Lys Thr Ile Gln
275 280 285
Gly Asp Glu Glu Asp Leu Arg
290 295
<210> 52
<211> 4303
<212> PRT
<213> 智人
<400> 52
Met Pro Pro Ala Ala Pro Ala Arg Leu Ala Leu Ala Leu Gly Leu Gly
1 5 10 15
Leu Trp Leu Gly Ala Leu Ala Gly Gly Pro Gly Arg Gly Cys Gly Pro
20 25 30
Cys Glu Pro Pro Cys Leu Cys Gly Pro Ala Pro Gly Ala Ala Cys Arg
35 40 45
Val Asn Cys Ser Gly Arg Gly Leu Arg Thr Leu Gly Pro Ala Leu Arg
50 55 60
Ile Pro Ala Asp Ala Thr Ala Leu Asp Val Ser His Asn Leu Leu Arg
65 70 75 80
Ala Leu Asp Val Gly Leu Leu Ala Asn Leu Ser Ala Leu Ala Glu Leu
85 90 95
Asp Ile Ser Asn Asn Lys Ile Ser Thr Leu Glu Glu Gly Ile Phe Ala
100 105 110
Asn Leu Phe Asn Leu Ser Glu Ile Asn Leu Ser Gly Asn Pro Phe Glu
115 120 125
Cys Asp Cys Gly Leu Ala Trp Leu Pro Arg Trp Ala Glu Glu Gln Gln
130 135 140
Val Arg Val Val Gln Pro Glu Ala Ala Thr Cys Ala Gly Pro Gly Ser
145 150 155 160
Leu Ala Gly Gln Pro Leu Leu Gly Ile Pro Leu Leu Asp Ser Gly Cys
165 170 175
Gly Glu Glu Tyr Val Ala Cys Leu Pro Asp Asn Ser Ser Gly Thr Val
180 185 190
Ala Ala Val Ser Phe Ser Ala Ala His Glu Gly Leu Leu Gln Pro Glu
195 200 205
Ala Cys Ser Ala Phe Cys Phe Ser Thr Gly Gln Gly Leu Ala Ala Leu
210 215 220
Ser Glu Gln Gly Trp Cys Leu Cys Gly Ala Ala Gln Pro Ser Ser Ala
225 230 235 240
Ser Phe Ala Cys Leu Ser Leu Cys Ser Gly Pro Pro Pro Pro Pro Ala
245 250 255
Pro Thr Cys Arg Gly Pro Thr Leu Leu Gln His Val Phe Pro Ala Ser
260 265 270
Pro Gly Ala Thr Leu Val Gly Pro His Gly Pro Leu Ala Ser Gly Gln
275 280 285
Leu Ala Ala Phe His Ile Ala Ala Pro Leu Pro Val Thr Ala Thr Arg
290 295 300
Trp Asp Phe Gly Asp Gly Ser Ala Glu Val Asp Ala Ala Gly Pro Ala
305 310 315 320
Ala Ser His Arg Tyr Val Leu Pro Gly Arg Tyr His Val Thr Ala Val
325 330 335
Leu Ala Leu Gly Ala Gly Ser Ala Leu Leu Gly Thr Asp Val Gln Val
340 345 350
Glu Ala Ala Pro Ala Ala Leu Glu Leu Val Cys Pro Ser Ser Val Gln
355 360 365
Ser Asp Glu Ser Leu Asp Leu Ser Ile Gln Asn Arg Gly Gly Ser Gly
370 375 380
Leu Glu Ala Ala Tyr Ser Ile Val Ala Leu Gly Glu Glu Pro Ala Arg
385 390 395 400
Ala Val His Pro Leu Cys Pro Ser Asp Thr Glu Ile Phe Pro Gly Asn
405 410 415
Gly His Cys Tyr Arg Leu Val Val Glu Lys Ala Ala Trp Leu Gln Ala
420 425 430
Gln Glu Gln Cys Gln Ala Trp Ala Gly Ala Ala Leu Ala Met Val Asp
435 440 445
Ser Pro Ala Val Gln Arg Phe Leu Val Ser Arg Val Thr Arg Ser Leu
450 455 460
Asp Val Trp Ile Gly Phe Ser Thr Val Gln Gly Val Glu Val Gly Pro
465 470 475 480
Ala Pro Gln Gly Glu Ala Phe Ser Leu Glu Ser Cys Gln Asn Trp Leu
485 490 495
Pro Gly Glu Pro His Pro Ala Thr Ala Glu His Cys Val Arg Leu Gly
500 505 510
Pro Thr Gly Trp Cys Asn Thr Asp Leu Cys Ser Ala Pro His Ser Tyr
515 520 525
Val Cys Glu Leu Gln Pro Gly Gly Pro Val Gln Asp Ala Glu Asn Leu
530 535 540
Leu Val Gly Ala Pro Ser Gly Asp Leu Gln Gly Pro Leu Thr Pro Leu
545 550 555 560
Ala Gln Gln Asp Gly Leu Ser Ala Pro His Glu Pro Val Glu Val Met
565 570 575
Val Phe Pro Gly Leu Arg Leu Ser Arg Glu Ala Phe Leu Thr Thr Ala
580 585 590
Glu Phe Gly Thr Gln Glu Leu Arg Arg Pro Ala Gln Leu Arg Leu Gln
595 600 605
Val Tyr Arg Leu Leu Ser Thr Ala Gly Thr Pro Glu Asn Gly Ser Glu
610 615 620
Pro Glu Ser Arg Ser Pro Asp Asn Arg Thr Gln Leu Ala Pro Ala Cys
625 630 635 640
Met Pro Gly Gly Arg Trp Cys Pro Gly Ala Asn Ile Cys Leu Pro Leu
645 650 655
Asp Ala Ser Cys His Pro Gln Ala Cys Ala Asn Gly Cys Thr Ser Gly
660 665 670
Pro Gly Leu Pro Gly Ala Pro Tyr Ala Leu Trp Arg Glu Phe Leu Phe
675 680 685
Ser Val Pro Ala Gly Pro Pro Ala Gln Tyr Ser Val Thr Leu His Gly
690 695 700
Gln Asp Val Leu Met Leu Pro Gly Asp Leu Val Gly Leu Gln His Asp
705 710 715 720
Ala Gly Pro Gly Ala Leu Leu His Cys Ser Pro Ala Pro Gly His Pro
725 730 735
Gly Pro Arg Ala Pro Tyr Leu Ser Ala Asn Ala Ser Ser Trp Leu Pro
740 745 750
His Leu Pro Ala Gln Leu Glu Gly Thr Trp Ala Cys Pro Ala Cys Ala
755 760 765
Leu Arg Leu Leu Ala Ala Thr Glu Gln Leu Thr Val Leu Leu Gly Leu
770 775 780
Arg Pro Asn Pro Gly Leu Arg Leu Pro Gly Arg Tyr Glu Val Arg Ala
785 790 795 800
Glu Val Gly Asn Gly Val Ser Arg His Asn Leu Ser Cys Ser Phe Asp
805 810 815
Val Val Ser Pro Val Ala Gly Leu Arg Val Ile Tyr Pro Ala Pro Arg
820 825 830
Asp Gly Arg Leu Tyr Val Pro Thr Asn Gly Ser Ala Leu Val Leu Gln
835 840 845
Val Asp Ser Gly Ala Asn Ala Thr Ala Thr Ala Arg Trp Pro Gly Gly
850 855 860
Ser Val Ser Ala Arg Phe Glu Asn Val Cys Pro Ala Leu Val Ala Thr
865 870 875 880
Phe Val Pro Gly Cys Pro Trp Glu Thr Asn Asp Thr Leu Phe Ser Val
885 890 895
Val Ala Leu Pro Trp Leu Ser Glu Gly Glu His Val Val Asp Val Val
900 905 910
Val Glu Asn Ser Ala Ser Arg Ala Asn Leu Ser Leu Arg Val Thr Ala
915 920 925
Glu Glu Pro Ile Cys Gly Leu Arg Ala Thr Pro Ser Pro Glu Ala Arg
930 935 940
Val Leu Gln Gly Val Leu Val Arg Tyr Ser Pro Val Val Glu Ala Gly
945 950 955 960
Ser Asp Met Val Phe Arg Trp Thr Ile Asn Asp Lys Gln Ser Leu Thr
965 970 975
Phe Gln Asn Val Val Phe Asn Val Ile Tyr Gln Ser Ala Ala Val Phe
980 985 990
Lys Leu Ser Leu Thr Ala Ser Asn His Val Ser Asn Val Thr Val Asn
995 1000 1005
Tyr Asn Val Thr Val Glu Arg Met Asn Arg Met Gln Gly Leu Gln
1010 1015 1020
Val Ser Thr Val Pro Ala Val Leu Ser Pro Asn Ala Thr Leu Ala
1025 1030 1035
Leu Thr Ala Gly Val Leu Val Asp Ser Ala Val Glu Val Ala Phe
1040 1045 1050
Leu Trp Thr Phe Gly Asp Gly Glu Gln Ala Leu His Gln Phe Gln
1055 1060 1065
Pro Pro Tyr Asn Glu Ser Phe Pro Val Pro Asp Pro Ser Val Ala
1070 1075 1080
Gln Val Leu Val Glu His Asn Val Met His Thr Tyr Ala Ala Pro
1085 1090 1095
Gly Glu Tyr Leu Leu Thr Val Leu Ala Ser Asn Ala Phe Glu Asn
1100 1105 1110
Leu Thr Gln Gln Val Pro Val Ser Val Arg Ala Ser Leu Pro Ser
1115 1120 1125
Val Ala Val Gly Val Ser Asp Gly Val Leu Val Ala Gly Arg Pro
1130 1135 1140
Val Thr Phe Tyr Pro His Pro Leu Pro Ser Pro Gly Gly Val Leu
1145 1150 1155
Tyr Thr Trp Asp Phe Gly Asp Gly Ser Pro Val Leu Thr Gln Ser
1160 1165 1170
Gln Pro Ala Ala Asn His Thr Tyr Ala Ser Arg Gly Thr Tyr His
1175 1180 1185
Val Arg Leu Glu Val Asn Asn Thr Val Ser Gly Ala Ala Ala Gln
1190 1195 1200
Ala Asp Val Arg Val Phe Glu Glu Leu Arg Gly Leu Ser Val Asp
1205 1210 1215
Met Ser Leu Ala Val Glu Gln Gly Ala Pro Val Val Val Ser Ala
1220 1225 1230
Ala Val Gln Thr Gly Asp Asn Ile Thr Trp Thr Phe Asp Met Gly
1235 1240 1245
Asp Gly Thr Val Leu Ser Gly Pro Glu Ala Thr Val Glu His Val
1250 1255 1260
Tyr Leu Arg Ala Gln Asn Cys Thr Val Thr Val Gly Ala Ala Ser
1265 1270 1275
Pro Ala Gly His Leu Ala Arg Ser Leu His Val Leu Val Phe Val
1280 1285 1290
Leu Glu Val Leu Arg Val Glu Pro Ala Ala Cys Ile Pro Thr Gln
1295 1300 1305
Pro Asp Ala Arg Leu Thr Ala Tyr Val Thr Gly Asn Pro Ala His
1310 1315 1320
Tyr Leu Phe Asp Trp Thr Phe Gly Asp Gly Ser Ser Asn Thr Thr
1325 1330 1335
Val Arg Gly Cys Pro Thr Val Thr His Asn Phe Thr Arg Ser Gly
1340 1345 1350
Thr Phe Pro Leu Ala Leu Val Leu Ser Ser Arg Val Asn Arg Ala
1355 1360 1365
His Tyr Phe Thr Ser Ile Cys Val Glu Pro Glu Val Gly Asn Val
1370 1375 1380
Thr Leu Gln Pro Glu Arg Gln Phe Val Gln Leu Gly Asp Glu Ala
1385 1390 1395
Trp Leu Val Ala Cys Ala Trp Pro Pro Phe Pro Tyr Arg Tyr Thr
1400 1405 1410
Trp Asp Phe Gly Thr Glu Glu Ala Ala Pro Thr Arg Ala Arg Gly
1415 1420 1425
Pro Glu Val Thr Phe Ile Tyr Arg Asp Pro Gly Ser Tyr Leu Val
1430 1435 1440
Thr Val Thr Ala Ser Asn Asn Ile Ser Ala Ala Asn Asp Ser Ala
1445 1450 1455
Leu Val Glu Val Gln Glu Pro Val Leu Val Thr Ser Ile Lys Val
1460 1465 1470
Asn Gly Ser Leu Gly Leu Glu Leu Gln Gln Pro Tyr Leu Phe Ser
1475 1480 1485
Ala Val Gly Arg Gly Arg Pro Ala Ser Tyr Leu Trp Asp Leu Gly
1490 1495 1500
Asp Gly Gly Trp Leu Glu Gly Pro Glu Val Thr His Ala Tyr Asn
1505 1510 1515
Ser Thr Gly Asp Phe Thr Val Arg Val Ala Gly Trp Asn Glu Val
1520 1525 1530
Ser Arg Ser Glu Ala Trp Leu Asn Val Thr Val Lys Arg Arg Val
1535 1540 1545
Arg Gly Leu Val Val Asn Ala Ser Arg Thr Val Val Pro Leu Asn
1550 1555 1560
Gly Ser Val Ser Phe Ser Thr Ser Leu Glu Ala Gly Ser Asp Val
1565 1570 1575
Arg Tyr Ser Trp Val Leu Cys Asp Arg Cys Thr Pro Ile Pro Gly
1580 1585 1590
Gly Pro Thr Ile Ser Tyr Thr Phe Arg Ser Val Gly Thr Phe Asn
1595 1600 1605
Ile Ile Val Thr Ala Glu Asn Glu Val Gly Ser Ala Gln Asp Ser
1610 1615 1620
Ile Phe Val Tyr Val Leu Gln Leu Ile Glu Gly Leu Gln Val Val
1625 1630 1635
Gly Gly Gly Arg Tyr Phe Pro Thr Asn His Thr Val Gln Leu Gln
1640 1645 1650
Ala Val Val Arg Asp Gly Thr Asn Val Ser Tyr Ser Trp Thr Ala
1655 1660 1665
Trp Arg Asp Arg Gly Pro Ala Leu Ala Gly Ser Gly Lys Gly Phe
1670 1675 1680
Ser Leu Thr Val Leu Glu Ala Gly Thr Tyr His Val Gln Leu Arg
1685 1690 1695
Ala Thr Asn Met Leu Gly Ser Ala Trp Ala Asp Cys Thr Met Asp
1700 1705 1710
Phe Val Glu Pro Val Gly Trp Leu Met Val Ala Ala Ser Pro Asn
1715 1720 1725
Pro Ala Ala Val Asn Thr Ser Val Thr Leu Ser Ala Glu Leu Ala
1730 1735 1740
Gly Gly Ser Gly Val Val Tyr Thr Trp Ser Leu Glu Glu Gly Leu
1745 1750 1755
Ser Trp Glu Thr Ser Glu Pro Phe Thr Thr His Ser Phe Pro Thr
1760 1765 1770
Pro Gly Leu His Leu Val Thr Met Thr Ala Gly Asn Pro Leu Gly
1775 1780 1785
Ser Ala Asn Ala Thr Val Glu Val Asp Val Gln Val Pro Val Ser
1790 1795 1800
Gly Leu Ser Ile Arg Ala Ser Glu Pro Gly Gly Ser Phe Val Ala
1805 1810 1815
Ala Gly Ser Ser Val Pro Phe Trp Gly Gln Leu Ala Thr Gly Thr
1820 1825 1830
Asn Val Ser Trp Cys Trp Ala Val Pro Gly Gly Ser Ser Lys Arg
1835 1840 1845
Gly Pro His Val Thr Met Val Phe Pro Asp Ala Gly Thr Phe Ser
1850 1855 1860
Ile Arg Leu Asn Ala Ser Asn Ala Val Ser Trp Val Ser Ala Thr
1865 1870 1875
Tyr Asn Leu Thr Ala Glu Glu Pro Ile Val Gly Leu Val Leu Trp
1880 1885 1890
Ala Ser Ser Lys Val Val Ala Pro Gly Gln Leu Val His Phe Gln
1895 1900 1905
Ile Leu Leu Ala Ala Gly Ser Ala Val Thr Phe Arg Leu Gln Val
1910 1915 1920
Gly Gly Ala Asn Pro Glu Val Leu Pro Gly Pro Arg Phe Ser His
1925 1930 1935
Ser Phe Pro Arg Val Gly Asp His Val Val Ser Val Arg Gly Lys
1940 1945 1950
Asn His Val Ser Trp Ala Gln Ala Gln Val Arg Ile Val Val Leu
1955 1960 1965
Glu Ala Val Ser Gly Leu Gln Val Pro Asn Cys Cys Glu Pro Gly
1970 1975 1980
Ile Ala Thr Gly Thr Glu Arg Asn Phe Thr Ala Arg Val Gln Arg
1985 1990 1995
Gly Ser Arg Val Ala Tyr Ala Trp Tyr Phe Ser Leu Gln Lys Val
2000 2005 2010
Gln Gly Asp Ser Leu Val Ile Leu Ser Gly Arg Asp Val Thr Tyr
2015 2020 2025
Thr Pro Val Ala Ala Gly Leu Leu Glu Ile Gln Val Arg Ala Phe
2030 2035 2040
Asn Ala Leu Gly Ser Glu Asn Arg Thr Leu Val Leu Glu Val Gln
2045 2050 2055
Asp Ala Val Gln Tyr Val Ala Leu Gln Ser Gly Pro Cys Phe Thr
2060 2065 2070
Asn Arg Ser Ala Gln Phe Glu Ala Ala Thr Ser Pro Ser Pro Arg
2075 2080 2085
Arg Val Ala Tyr His Trp Asp Phe Gly Asp Gly Ser Pro Gly Gln
2090 2095 2100
Asp Thr Asp Glu Pro Arg Ala Glu His Ser Tyr Leu Arg Pro Gly
2105 2110 2115
Asp Tyr Arg Val Gln Val Asn Ala Ser Asn Leu Val Ser Phe Phe
2120 2125 2130
Val Ala Gln Ala Thr Val Thr Val Gln Val Leu Ala Cys Arg Glu
2135 2140 2145
Pro Glu Val Asp Val Val Leu Pro Leu Gln Val Leu Met Arg Arg
2150 2155 2160
Ser Gln Arg Asn Tyr Leu Glu Ala His Val Asp Leu Arg Asp Cys
2165 2170 2175
Val Thr Tyr Gln Thr Glu Tyr Arg Trp Glu Val Tyr Arg Thr Ala
2180 2185 2190
Ser Cys Gln Arg Pro Gly Arg Pro Ala Arg Val Ala Leu Pro Gly
2195 2200 2205
Val Asp Val Ser Arg Pro Arg Leu Val Leu Pro Arg Leu Ala Leu
2210 2215 2220
Pro Val Gly His Tyr Cys Phe Val Phe Val Val Ser Phe Gly Asp
2225 2230 2235
Thr Pro Leu Thr Gln Ser Ile Gln Ala Asn Val Thr Val Ala Pro
2240 2245 2250
Glu Arg Leu Val Pro Ile Ile Glu Gly Gly Ser Tyr Arg Val Trp
2255 2260 2265
Ser Asp Thr Arg Asp Leu Val Leu Asp Gly Ser Glu Ser Tyr Asp
2270 2275 2280
Pro Asn Leu Glu Asp Gly Asp Gln Thr Pro Leu Ser Phe His Trp
2285 2290 2295
Ala Cys Val Ala Ser Thr Gln Arg Glu Ala Gly Gly Cys Ala Leu
2300 2305 2310
Asn Phe Gly Pro Arg Gly Ser Ser Thr Val Thr Ile Pro Arg Glu
2315 2320 2325
Arg Leu Ala Ala Gly Val Glu Tyr Thr Phe Ser Leu Thr Val Trp
2330 2335 2340
Lys Ala Gly Arg Lys Glu Glu Ala Thr Asn Gln Thr Val Leu Ile
2345 2350 2355
Arg Ser Gly Arg Val Pro Ile Val Ser Leu Glu Cys Val Ser Cys
2360 2365 2370
Lys Ala Gln Ala Val Tyr Glu Val Ser Arg Ser Ser Tyr Val Tyr
2375 2380 2385
Leu Glu Gly Arg Cys Leu Asn Cys Ser Ser Gly Ser Lys Arg Gly
2390 2395 2400
Arg Trp Ala Ala Arg Thr Phe Ser Asn Lys Thr Leu Val Leu Asp
2405 2410 2415
Glu Thr Thr Thr Ser Thr Gly Ser Ala Gly Met Arg Leu Val Leu
2420 2425 2430
Arg Arg Gly Val Leu Arg Asp Gly Glu Gly Tyr Thr Phe Thr Leu
2435 2440 2445
Thr Val Leu Gly Arg Ser Gly Glu Glu Glu Gly Cys Ala Ser Ile
2450 2455 2460
Arg Leu Ser Pro Asn Arg Pro Pro Leu Gly Gly Ser Cys Arg Leu
2465 2470 2475
Phe Pro Leu Gly Ala Val His Ala Leu Thr Thr Lys Val His Phe
2480 2485 2490
Glu Cys Thr Gly Trp His Asp Ala Glu Asp Ala Gly Ala Pro Leu
2495 2500 2505
Val Tyr Ala Leu Leu Leu Arg Arg Cys Arg Gln Gly His Cys Glu
2510 2515 2520
Glu Phe Cys Val Tyr Lys Gly Ser Leu Ser Ser Tyr Gly Ala Val
2525 2530 2535
Leu Pro Pro Gly Phe Arg Pro His Phe Glu Val Gly Leu Ala Val
2540 2545 2550
Val Val Gln Asp Gln Leu Gly Ala Ala Val Val Ala Leu Asn Arg
2555 2560 2565
Ser Leu Ala Ile Thr Leu Pro Glu Pro Asn Gly Ser Ala Thr Gly
2570 2575 2580
Leu Thr Val Trp Leu His Gly Leu Thr Ala Ser Val Leu Pro Gly
2585 2590 2595
Leu Leu Arg Gln Ala Asp Pro Gln His Val Ile Glu Tyr Ser Leu
2600 2605 2610
Ala Leu Val Thr Val Leu Asn Glu Tyr Glu Arg Ala Leu Asp Val
2615 2620 2625
Ala Ala Glu Pro Lys His Glu Arg Gln His Arg Ala Gln Ile Arg
2630 2635 2640
Lys Asn Ile Thr Glu Thr Leu Val Ser Leu Arg Val His Thr Val
2645 2650 2655
Asp Asp Ile Gln Gln Ile Ala Ala Ala Leu Ala Gln Cys Met Gly
2660 2665 2670
Pro Ser Arg Glu Leu Val Cys Arg Ser Cys Leu Lys Gln Thr Leu
2675 2680 2685
His Lys Leu Glu Ala Met Met Leu Ile Leu Gln Ala Glu Thr Thr
2690 2695 2700
Ala Gly Thr Val Thr Pro Thr Ala Ile Gly Asp Ser Ile Leu Asn
2705 2710 2715
Ile Thr Gly Asp Leu Ile His Leu Ala Ser Ser Asp Val Arg Ala
2720 2725 2730
Pro Gln Pro Ser Glu Leu Gly Ala Glu Ser Pro Ser Arg Met Val
2735 2740 2745
Ala Ser Gln Ala Tyr Asn Leu Thr Ser Ala Leu Met Arg Ile Leu
2750 2755 2760
Met Arg Ser Arg Val Leu Asn Glu Glu Pro Leu Thr Leu Ala Gly
2765 2770 2775
Glu Glu Ile Val Ala Gln Gly Lys Arg Ser Asp Pro Arg Ser Leu
2780 2785 2790
Leu Cys Tyr Gly Gly Ala Pro Gly Pro Gly Cys His Phe Ser Ile
2795 2800 2805
Pro Glu Ala Phe Ser Gly Ala Leu Ala Asn Leu Ser Asp Val Val
2810 2815 2820
Gln Leu Ile Phe Leu Val Asp Ser Asn Pro Phe Pro Phe Gly Tyr
2825 2830 2835
Ile Ser Asn Tyr Thr Val Ser Thr Lys Val Ala Ser Met Ala Phe
2840 2845 2850
Gln Thr Gln Ala Gly Ala Gln Ile Pro Ile Glu Arg Leu Ala Ser
2855 2860 2865
Glu Arg Ala Ile Thr Val Lys Val Pro Asn Asn Ser Asp Trp Ala
2870 2875 2880
Ala Arg Gly His Arg Ser Ser Ala Asn Ser Ala Asn Ser Val Val
2885 2890 2895
Val Gln Pro Gln Ala Ser Val Gly Ala Val Val Thr Leu Asp Ser
2900 2905 2910
Ser Asn Pro Ala Ala Gly Leu His Leu Gln Leu Asn Tyr Thr Leu
2915 2920 2925
Leu Asp Gly His Tyr Leu Ser Glu Glu Pro Glu Pro Tyr Leu Ala
2930 2935 2940
Val Tyr Leu His Ser Glu Pro Arg Pro Asn Glu His Asn Cys Ser
2945 2950 2955
Ala Ser Arg Arg Ile Arg Pro Glu Ser Leu Gln Gly Ala Asp His
2960 2965 2970
Arg Pro Tyr Thr Phe Phe Ile Ser Pro Gly Ser Arg Asp Pro Ala
2975 2980 2985
Gly Ser Tyr His Leu Asn Leu Ser Ser His Phe Arg Trp Ser Ala
2990 2995 3000
Leu Gln Val Ser Val Gly Leu Tyr Thr Ser Leu Cys Gln Tyr Phe
3005 3010 3015
Ser Glu Glu Asp Met Val Trp Arg Thr Glu Gly Leu Leu Pro Leu
3020 3025 3030
Glu Glu Thr Ser Pro Arg Gln Ala Val Cys Leu Thr Arg His Leu
3035 3040 3045
Thr Ala Phe Gly Ala Ser Leu Phe Val Pro Pro Ser His Val Arg
3050 3055 3060
Phe Val Phe Pro Glu Pro Thr Ala Asp Val Asn Tyr Ile Val Met
3065 3070 3075
Leu Thr Cys Ala Val Cys Leu Val Thr Tyr Met Val Met Ala Ala
3080 3085 3090
Ile Leu His Lys Leu Asp Gln Leu Asp Ala Ser Arg Gly Arg Ala
3095 3100 3105
Ile Pro Phe Cys Gly Gln Arg Gly Arg Phe Lys Tyr Glu Ile Leu
3110 3115 3120
Val Lys Thr Gly Trp Gly Arg Gly Ser Gly Thr Thr Ala His Val
3125 3130 3135
Gly Ile Met Leu Tyr Gly Val Asp Ser Arg Ser Gly His Arg His
3140 3145 3150
Leu Asp Gly Asp Arg Ala Phe His Arg Asn Ser Leu Asp Ile Phe
3155 3160 3165
Arg Ile Ala Thr Pro His Ser Leu Gly Ser Val Trp Lys Ile Arg
3170 3175 3180
Val Trp His Asp Asn Lys Gly Leu Ser Pro Ala Trp Phe Leu Gln
3185 3190 3195
His Val Ile Val Arg Asp Leu Gln Thr Ala Arg Ser Ala Phe Phe
3200 3205 3210
Leu Val Asn Asp Trp Leu Ser Val Glu Thr Glu Ala Asn Gly Gly
3215 3220 3225
Leu Val Glu Lys Glu Val Leu Ala Ala Ser Asp Ala Ala Leu Leu
3230 3235 3240
Arg Phe Arg Arg Leu Leu Val Ala Glu Leu Gln Arg Gly Phe Phe
3245 3250 3255
Asp Lys His Ile Trp Leu Ser Ile Trp Asp Arg Pro Pro Arg Ser
3260 3265 3270
Arg Phe Thr Arg Ile Gln Arg Ala Thr Cys Cys Val Leu Leu Ile
3275 3280 3285
Cys Leu Phe Leu Gly Ala Asn Ala Val Trp Tyr Gly Ala Val Gly
3290 3295 3300
Asp Ser Ala Tyr Ser Thr Gly His Val Ser Arg Leu Ser Pro Leu
3305 3310 3315
Ser Val Asp Thr Val Ala Val Gly Leu Val Ser Ser Val Val Val
3320 3325 3330
Tyr Pro Val Tyr Leu Ala Ile Leu Phe Leu Phe Arg Met Ser Arg
3335 3340 3345
Ser Lys Val Ala Gly Ser Pro Ser Pro Thr Pro Ala Gly Gln Gln
3350 3355 3360
Val Leu Asp Ile Asp Ser Cys Leu Asp Ser Ser Val Leu Asp Ser
3365 3370 3375
Ser Phe Leu Thr Phe Ser Gly Leu His Ala Glu Gln Ala Phe Val
3380 3385 3390
Gly Gln Met Lys Ser Asp Leu Phe Leu Asp Asp Ser Lys Ser Leu
3395 3400 3405
Val Cys Trp Pro Ser Gly Glu Gly Thr Leu Ser Trp Pro Asp Leu
3410 3415 3420
Leu Ser Asp Pro Ser Ile Val Gly Ser Asn Leu Arg Gln Leu Ala
3425 3430 3435
Arg Gly Gln Ala Gly His Gly Leu Gly Pro Glu Glu Asp Gly Phe
3440 3445 3450
Ser Leu Ala Ser Pro Tyr Ser Pro Ala Lys Ser Phe Ser Ala Ser
3455 3460 3465
Asp Glu Asp Leu Ile Gln Gln Val Leu Ala Glu Gly Val Ser Ser
3470 3475 3480
Pro Ala Pro Thr Gln Asp Thr His Met Glu Thr Asp Leu Leu Ser
3485 3490 3495
Ser Leu Ser Ser Thr Pro Gly Glu Lys Thr Glu Thr Leu Ala Leu
3500 3505 3510
Gln Arg Leu Gly Glu Leu Gly Pro Pro Ser Pro Gly Leu Asn Trp
3515 3520 3525
Glu Gln Pro Gln Ala Ala Arg Leu Ser Arg Thr Gly Leu Val Glu
3530 3535 3540
Gly Leu Arg Lys Arg Leu Leu Pro Ala Trp Cys Ala Ser Leu Ala
3545 3550 3555
His Gly Leu Ser Leu Leu Leu Val Ala Val Ala Val Ala Val Ser
3560 3565 3570
Gly Trp Val Gly Ala Ser Phe Pro Pro Gly Val Ser Val Ala Trp
3575 3580 3585
Leu Leu Ser Ser Ser Ala Ser Phe Leu Ala Ser Phe Leu Gly Trp
3590 3595 3600
Glu Pro Leu Lys Val Leu Leu Glu Ala Leu Tyr Phe Ser Leu Val
3605 3610 3615
Ala Lys Arg Leu His Pro Asp Glu Asp Asp Thr Leu Val Glu Ser
3620 3625 3630
Pro Ala Val Thr Pro Val Ser Ala Arg Val Pro Arg Val Arg Pro
3635 3640 3645
Pro His Gly Phe Ala Leu Phe Leu Ala Lys Glu Glu Ala Arg Lys
3650 3655 3660
Val Lys Arg Leu His Gly Met Leu Arg Ser Leu Leu Val Tyr Met
3665 3670 3675
Leu Phe Leu Leu Val Thr Leu Leu Ala Ser Tyr Gly Asp Ala Ser
3680 3685 3690
Cys His Gly His Ala Tyr Arg Leu Gln Ser Ala Ile Lys Gln Glu
3695 3700 3705
Leu His Ser Arg Ala Phe Leu Ala Ile Thr Arg Ser Glu Glu Leu
3710 3715 3720
Trp Pro Trp Met Ala His Val Leu Leu Pro Tyr Val His Gly Asn
3725 3730 3735
Gln Ser Ser Pro Glu Leu Gly Pro Pro Arg Leu Arg Gln Val Arg
3740 3745 3750
Leu Gln Glu Ala Leu Tyr Pro Asp Pro Pro Gly Pro Arg Val His
3755 3760 3765
Thr Cys Ser Ala Ala Gly Gly Phe Ser Thr Ser Asp Tyr Asp Val
3770 3775 3780
Gly Trp Glu Ser Pro His Asn Gly Ser Gly Thr Trp Ala Tyr Ser
3785 3790 3795
Ala Pro Asp Leu Leu Gly Ala Trp Ser Trp Gly Ser Cys Ala Val
3800 3805 3810
Tyr Asp Ser Gly Gly Tyr Val Gln Glu Leu Gly Leu Ser Leu Glu
3815 3820 3825
Glu Ser Arg Asp Arg Leu Arg Phe Leu Gln Leu His Asn Trp Leu
3830 3835 3840
Asp Asn Arg Ser Arg Ala Val Phe Leu Glu Leu Thr Arg Tyr Ser
3845 3850 3855
Pro Ala Val Gly Leu His Ala Ala Val Thr Leu Arg Leu Glu Phe
3860 3865 3870
Pro Ala Ala Gly Arg Ala Leu Ala Ala Leu Ser Val Arg Pro Phe
3875 3880 3885
Ala Leu Arg Arg Leu Ser Ala Gly Leu Ser Leu Pro Leu Leu Thr
3890 3895 3900
Ser Val Cys Leu Leu Leu Phe Ala Val His Phe Ala Val Ala Glu
3905 3910 3915
Ala Arg Thr Trp His Arg Glu Gly Arg Trp Arg Val Leu Arg Leu
3920 3925 3930
Gly Ala Trp Ala Arg Trp Leu Leu Val Ala Leu Thr Ala Ala Thr
3935 3940 3945
Ala Leu Val Arg Leu Ala Gln Leu Gly Ala Ala Asp Arg Gln Trp
3950 3955 3960
Thr Arg Phe Val Arg Gly Arg Pro Arg Arg Phe Thr Ser Phe Asp
3965 3970 3975
Gln Val Ala Gln Leu Ser Ser Ala Ala Arg Gly Leu Ala Ala Ser
3980 3985 3990
Leu Leu Phe Leu Leu Leu Val Lys Ala Ala Gln Gln Leu Arg Phe
3995 4000 4005
Val Arg Gln Trp Ser Val Phe Gly Lys Thr Leu Cys Arg Ala Leu
4010 4015 4020
Pro Glu Leu Leu Gly Val Thr Leu Gly Leu Val Val Leu Gly Val
4025 4030 4035
Ala Tyr Ala Gln Leu Ala Ile Leu Leu Val Ser Ser Cys Val Asp
4040 4045 4050
Ser Leu Trp Ser Val Ala Gln Ala Leu Leu Val Leu Cys Pro Gly
4055 4060 4065
Thr Gly Leu Ser Thr Leu Cys Pro Ala Glu Ser Trp His Leu Ser
4070 4075 4080
Pro Leu Leu Cys Val Gly Leu Trp Ala Leu Arg Leu Trp Gly Ala
4085 4090 4095
Leu Arg Leu Gly Ala Val Ile Leu Arg Trp Arg Tyr His Ala Leu
4100 4105 4110
Arg Gly Glu Leu Tyr Arg Pro Ala Trp Glu Pro Gln Asp Tyr Glu
4115 4120 4125
Met Val Glu Leu Phe Leu Arg Arg Leu Arg Leu Trp Met Gly Leu
4130 4135 4140
Ser Lys Val Lys Glu Phe Arg His Lys Val Arg Phe Glu Gly Met
4145 4150 4155
Glu Pro Leu Pro Ser Arg Ser Ser Arg Gly Ser Lys Val Ser Pro
4160 4165 4170
Asp Val Pro Pro Pro Ser Ala Gly Ser Asp Ala Ser His Pro Ser
4175 4180 4185
Thr Ser Ser Ser Gln Leu Asp Gly Leu Ser Val Ser Leu Gly Arg
4190 4195 4200
Leu Gly Thr Arg Cys Glu Pro Glu Pro Ser Arg Leu Gln Ala Val
4205 4210 4215
Phe Glu Ala Leu Leu Thr Gln Phe Asp Arg Leu Asn Gln Ala Thr
4220 4225 4230
Glu Asp Val Tyr Gln Leu Glu Gln Gln Leu His Ser Leu Gln Gly
4235 4240 4245
Arg Arg Ser Ser Arg Ala Pro Ala Gly Ser Ser Arg Gly Pro Ser
4250 4255 4260
Pro Gly Leu Arg Pro Ala Leu Pro Ser Arg Leu Ala Arg Ala Ser
4265 4270 4275
Arg Gly Val Asp Leu Ala Thr Gly Pro Ser Arg Thr Pro Leu Arg
4280 4285 4290
Ala Lys Asn Lys Val His Pro Ser Ser Thr
4295 4300
<210> 53
<211> 202
<212> PRT
<213> 智人
<400> 53
Met Cys Arg Thr Leu Ala Ala Phe Pro Thr Thr Cys Leu Glu Arg Ala
1 5 10 15
Lys Glu Phe Lys Thr Arg Leu Gly Ile Phe Leu His Lys Ser Glu Leu
20 25 30
Gly Cys Asp Thr Gly Ser Thr Gly Lys Phe Glu Trp Gly Ser Lys His
35 40 45
Ser Lys Glu Asn Arg Asn Phe Ser Glu Asp Val Leu Gly Trp Arg Glu
50 55 60
Ser Phe Asp Leu Leu Leu Ser Ser Lys Asn Gly Val Ala Ala Phe His
65 70 75 80
Ala Phe Leu Lys Thr Glu Phe Ser Glu Glu Asn Leu Glu Phe Trp Leu
85 90 95
Ala Cys Glu Glu Phe Lys Lys Ile Arg Ser Ala Thr Lys Leu Ala Ser
100 105 110
Arg Ala His Gln Ile Phe Glu Glu Phe Ile Cys Ser Glu Ala Pro Lys
115 120 125
Glu Val Asn Ile Asp His Glu Thr His Glu Leu Thr Arg Met Asn Leu
130 135 140
Gln Thr Ala Thr Ala Thr Cys Phe Asp Ala Ala Gln Gly Lys Thr Arg
145 150 155 160
Thr Leu Met Glu Lys Asp Ser Tyr Pro Arg Phe Leu Lys Ser Pro Ala
165 170 175
Tyr Arg Asp Leu Ala Ala Gln Ala Ser Ala Ala Ser Ala Thr Leu Ser
180 185 190
Ser Cys Ser Leu Asp Glu Pro Ser His Thr
195 200
<210> 54
<211> 853
<212> PRT
<213> 智人
<400> 54
Met Ser Glu Lys Val Asp Trp Leu Gln Ser Gln Asn Gly Val Cys Lys
1 5 10 15
Val Asp Val Tyr Ser Pro Gly Asp Asn Gln Ala Gln Asp Trp Lys Met
20 25 30
Asp Thr Ser Thr Asp Pro Val Arg Val Leu Ser Trp Leu Arg Arg Asp
35 40 45
Leu Glu Lys Ser Thr Ala Glu Phe Gln Asp Val Arg Phe Lys Pro Gly
50 55 60
Glu Ser Phe Gly Gly Glu Thr Ser Asn Ser Gly Asp Pro His Lys Gly
65 70 75 80
Phe Ser Val Asp Tyr Tyr Asn Thr Thr Thr Lys Gly Thr Pro Glu Arg
85 90 95
Leu His Phe Glu Met Thr His Lys Glu Ile Pro Cys Gln Gly Pro Arg
100 105 110
Ala Gln Leu Gly Asn Gly Ser Ser Val Asp Glu Val Ser Phe Tyr Ala
115 120 125
Asn Arg Leu Thr Asn Leu Val Ile Ala Met Ala Arg Lys Glu Ile Asn
130 135 140
Glu Lys Ile Asp Gly Ser Glu Asn Lys Cys Val Tyr Gln Ser Leu Tyr
145 150 155 160
Met Gly Asn Glu Pro Thr Pro Thr Lys Ser Leu Ser Lys Ile Ala Ser
165 170 175
Glu Leu Val Asn Glu Thr Val Ser Ala Cys Ser Arg Asn Ala Ala Pro
180 185 190
Asp Lys Ala Pro Gly Ser Gly Asp Arg Val Ser Gly Ser Ser Gln Ser
195 200 205
Pro Pro Asn Leu Lys Tyr Lys Ser Thr Leu Lys Ile Lys Glu Ser Thr
210 215 220
Lys Glu Arg Gln Gly Pro Asp Asp Lys Pro Pro Ser Lys Lys Ser Phe
225 230 235 240
Phe Tyr Lys Glu Val Phe Glu Ser Arg Asn Gly Asp Tyr Ala Arg Glu
245 250 255
Gly Gly Arg Phe Phe Pro Arg Glu Arg Lys Arg Phe Arg Gly Gln Glu
260 265 270
Arg Pro Asp Asp Phe Thr Ala Ser Val Ser Glu Gly Ile Met Thr Tyr
275 280 285
Ala Asn Ser Val Val Ser Asp Met Met Val Ser Ile Met Lys Thr Leu
290 295 300
Lys Ile Gln Val Lys Asp Thr Thr Ile Ala Thr Ile Leu Leu Lys Lys
305 310 315 320
Val Leu Leu Lys His Ala Lys Glu Val Val Ser Asp Leu Ile Asp Ser
325 330 335
Phe Leu Arg Asn Leu His Ser Val Thr Gly Thr Leu Met Thr Asp Thr
340 345 350
Gln Phe Val Ser Ala Val Lys Arg Thr Val Phe Ser His Gly Ser Gln
355 360 365
Lys Ala Thr Asp Ile Met Asp Ala Met Leu Arg Lys Leu Tyr Asn Val
370 375 380
Met Phe Ala Lys Lys Val Pro Glu His Val Arg Lys Ala Gln Asp Lys
385 390 395 400
Ala Glu Ser Tyr Ser Leu Ile Ser Met Lys Gly Met Gly Asp Pro Lys
405 410 415
Asn Arg Asn Val Asn Phe Ala Met Lys Ser Glu Thr Lys Leu Arg Glu
420 425 430
Lys Met Tyr Ser Glu Pro Lys Ser Glu Glu Glu Thr Cys Ala Lys Thr
435 440 445
Leu Gly Glu His Ile Ile Lys Glu Gly Leu Thr Leu Trp His Lys Thr
450 455 460
Gln Gln Lys Glu Cys Lys Ser Leu Gly Phe Gln His Ala Ala Phe Glu
465 470 475 480
Ala Pro Asn Thr Gln Arg Lys Pro Ala Ser Asp Ile Ser Phe Glu Tyr
485 490 495
Pro Glu Asp Ile Gly Asn Leu Ser Leu Pro Pro Tyr Pro Pro Glu Lys
500 505 510
Pro Glu Asn Phe Met Tyr Asp Ser Asp Ser Trp Ala Glu Asp Leu Ile
515 520 525
Val Ser Ala Leu Leu Leu Ile Gln Tyr His Leu Ala Gln Gly Gly Arg
530 535 540
Arg Asp Ala Arg Ser Phe Val Glu Ala Ala Gly Thr Thr Asn Phe Pro
545 550 555 560
Ala Asn Glu Pro Pro Val Ala Pro Asp Glu Ser Cys Leu Lys Ser Ala
565 570 575
Pro Ile Val Gly Asp Gln Glu Gln Ala Glu Lys Lys Asp Leu Arg Ser
580 585 590
Val Phe Phe Asn Phe Ile Arg Asn Leu Leu Ser Glu Thr Ile Phe Lys
595 600 605
Arg Asp Gln Ser Pro Glu Pro Lys Val Pro Glu Gln Pro Val Lys Glu
610 615 620
Asp Arg Lys Leu Cys Glu Arg Pro Leu Ala Ser Ser Pro Pro Arg Leu
625 630 635 640
Tyr Glu Asp Asp Glu Thr Pro Gly Ala Leu Ser Gly Leu Thr Lys Met
645 650 655
Ala Val Ser Gln Ile Asp Gly His Met Ser Gly Gln Met Val Glu His
660 665 670
Leu Met Asn Ser Val Met Lys Leu Cys Val Ile Ile Ala Lys Ser Cys
675 680 685
Asp Ala Ser Leu Ala Glu Leu Gly Asp Asp Lys Ser Gly Asp Ala Ser
690 695 700
Arg Leu Thr Ser Ala Phe Pro Asp Ser Leu Tyr Glu Cys Leu Pro Ala
705 710 715 720
Lys Gly Thr Gly Ser Ala Glu Ala Val Leu Gln Asn Ala Tyr Gln Ala
725 730 735
Ile His Asn Glu Met Arg Gly Thr Ser Gly Gln Pro Pro Glu Gly Cys
740 745 750
Ala Ala Pro Thr Val Ile Val Ser Asn His Asn Leu Thr Asp Thr Val
755 760 765
Gln Asn Lys Gln Leu Gln Ala Val Leu Gln Trp Val Ala Ala Ser Glu
770 775 780
Leu Asn Val Pro Ile Leu Tyr Phe Ala Gly Asp Asp Glu Gly Ile Gln
785 790 795 800
Glu Lys Leu Leu Gln Leu Ser Ala Ala Ala Val Asp Lys Gly Cys Ser
805 810 815
Val Gly Glu Val Leu Gln Ser Val Leu Arg Tyr Glu Lys Glu Arg Gln
820 825 830
Leu Asn Glu Ala Val Gly Asn Val Thr Pro Leu Gln Leu Leu Asp Trp
835 840 845
Leu Met Val Asn Leu
850
<210> 55
<211> 279
<212> PRT
<213> 智人
<400> 55
Met Ser Leu Phe Asp Leu Phe Arg Gly Phe Phe Gly Phe Pro Gly Pro
1 5 10 15
Arg Ser His Arg Asp Pro Phe Phe Gly Gly Met Thr Arg Asp Glu Asp
20 25 30
Asp Asp Glu Glu Glu Glu Glu Glu Gly Gly Ser Trp Gly Arg Gly Asn
35 40 45
Pro Arg Phe His Ser Pro Gln His Pro Pro Glu Glu Phe Gly Phe Gly
50 55 60
Phe Ser Phe Ser Pro Gly Gly Gly Ile Arg Phe His Asp Asn Phe Gly
65 70 75 80
Phe Asp Asp Leu Val Arg Asp Phe Asn Ser Ile Phe Ser Asp Met Gly
85 90 95
Ala Trp Thr Leu Pro Ser His Pro Pro Glu Leu Pro Gly Pro Glu Ser
100 105 110
Glu Thr Pro Gly Glu Arg Leu Arg Glu Gly Gln Thr Leu Arg Asp Ser
115 120 125
Met Leu Lys Tyr Pro Asp Ser His Gln Pro Arg Ile Phe Gly Gly Val
130 135 140
Leu Glu Ser Asp Ala Arg Ser Glu Ser Pro Gln Pro Ala Pro Asp Trp
145 150 155 160
Gly Ser Gln Arg Pro Phe His Arg Phe Asp Asp Val Trp Pro Met Asp
165 170 175
Pro His Pro Arg Thr Arg Glu Asp Asn Asp Leu Asp Ser Gln Val Ser
180 185 190
Gln Glu Gly Leu Gly Pro Val Leu Gln Pro Gln Pro Lys Ser Tyr Phe
195 200 205
Lys Ser Ile Ser Val Thr Lys Ile Thr Lys Pro Asp Gly Ile Val Glu
210 215 220
Glu Arg Arg Thr Val Val Asp Ser Glu Gly Arg Thr Glu Thr Thr Val
225 230 235 240
Thr Arg His Glu Ala Asp Ser Ser Pro Arg Gly Asp Pro Glu Ser Pro
245 250 255
Arg Pro Pro Ala Leu Asp Asp Ala Phe Ser Ile Leu Asp Leu Phe Leu
260 265 270
Gly Arg Trp Phe Arg Ser Arg
275
<210> 56
<211> 968
<212> PRT
<213> 智人
<400> 56
Met Leu Thr Glu Ala Ser Leu Ser Ile Trp Gly Trp Gly Ser Leu Gly
1 5 10 15
Ile Val Leu Phe Leu Ile Thr Phe Gly Pro Phe Val Ile Phe Tyr Leu
20 25 30
Thr Phe Tyr Ile Leu Cys Phe Val Gly Gly Gly Leu Val Val Thr Leu
35 40 45
Leu Phe Gly Lys Thr Asn Ser Glu Lys Tyr Leu Glu Gln Cys Glu His
50 55 60
Ser Phe Leu Pro Pro Thr Ser Pro Gly Val Pro Lys Cys Leu Glu Glu
65 70 75 80
Met Lys Arg Glu Ala Arg Thr Ile Lys Ile Asp Arg Arg Leu Thr Gly
85 90 95
Ala Asn Ile Ile Asp Glu Pro Leu Gln Gln Val Ile Gln Phe Ser Leu
100 105 110
Arg Asp Tyr Val Gln Tyr Trp Tyr Tyr Thr Leu Ser Asp Asp Glu Ser
115 120 125
Phe Leu Leu Glu Ile Arg Gln Thr Leu Gln Asn Ala Leu Ile Gln Phe
130 135 140
Ala Thr Arg Ser Lys Glu Ile Asp Trp Gln Pro Tyr Phe Thr Thr Arg
145 150 155 160
Ile Val Asp Asp Phe Gly Thr His Leu Arg Val Phe Arg Lys Ala Gln
165 170 175
Gln Lys Ile Thr Glu Lys Asp Asp Gln Val Lys Gly Thr Ala Glu Asp
180 185 190
Leu Val Asp Thr Phe Phe Glu Val Glu Val Glu Met Glu Lys Glu Val
195 200 205
Cys Arg Asp Leu Val Cys Thr Ser Pro Lys Asp Glu Glu Gly Phe Leu
210 215 220
Arg Asp Leu Cys Glu Val Leu Leu Tyr Leu Leu Leu Pro Pro Gly Asp
225 230 235 240
Phe Gln Asn Lys Ile Met Arg Tyr Phe Val Arg Glu Ile Leu Ala Arg
245 250 255
Gly Ile Leu Leu Pro Leu Ile Asn Gln Leu Ser Asp Pro Asp Tyr Ile
260 265 270
Asn Gln Tyr Val Ile Trp Met Ile Arg Asp Ser Asn Cys Asn Tyr Glu
275 280 285
Ala Phe Met Asn Ile Ile Lys Leu Ser Asp Asn Ile Gly Glu Leu Glu
290 295 300
Ala Val Arg Asp Lys Ala Ala Glu Glu Leu Gln Tyr Leu Arg Ser Leu
305 310 315 320
Asp Thr Ala Gly Asp Asp Ile Asn Thr Ile Lys Asn Gln Ile Asn Ser
325 330 335
Leu Leu Phe Val Lys Lys Val Cys Asp Ser Arg Ile Gln Arg Leu Gln
340 345 350
Ser Gly Lys Glu Ile Asn Thr Val Lys Leu Ala Ala Asn Phe Gly Lys
355 360 365
Leu Cys Thr Val Pro Leu Asp Ser Ile Leu Val Asp Asn Val Ala Leu
370 375 380
Gln Phe Phe Met Asp Tyr Met Gln Gln Thr Gly Gly Gln Ala His Leu
385 390 395 400
Phe Phe Trp Met Thr Val Glu Gly Tyr Arg Val Thr Ala Gln Gln Gln
405 410 415
Leu Glu Val Leu Leu Ser Arg Gln Arg Asp Gly Lys His Gln Thr Asn
420 425 430
Gln Thr Lys Gly Leu Leu Arg Ala Ala Ala Val Gly Ile Tyr Glu Gln
435 440 445
Tyr Leu Ser Glu Lys Ala Ser Pro Arg Val Thr Val Asp Asp Tyr Leu
450 455 460
Val Ala Lys Leu Ala Asp Thr Leu Asn His Glu Asp Pro Thr Pro Glu
465 470 475 480
Ile Phe Asp Asp Ile Gln Arg Lys Val Tyr Glu Leu Met Leu Arg Asp
485 490 495
Glu Arg Phe Tyr Pro Ser Phe Arg Gln Asn Ala Leu Tyr Val Arg Met
500 505 510
Leu Ala Glu Leu Asp Met Leu Lys Asp Pro Ser Phe Arg Gly Ser Asp
515 520 525
Asp Gly Asp Gly Glu Ser Phe Asn Gly Ser Pro Thr Gly Ser Ile Asn
530 535 540
Leu Ser Leu Asp Asp Leu Ser Asn Val Ser Ser Asp Asp Ser Val Gln
545 550 555 560
Leu His Ala Tyr Ile Ser Asp Thr Val Tyr Ala Asp Tyr Asp Pro Tyr
565 570 575
Ala Val Ala Gly Val Cys Asn Asp His Gly Lys Thr Tyr Ala Leu Tyr
580 585 590
Ala Ile Thr Val His Arg Arg Asn Leu Asn Ser Glu Glu Met Trp Lys
595 600 605
Thr Tyr Arg Arg Tyr Ser Asp Phe His Asp Phe His Met Arg Ile Thr
610 615 620
Glu Gln Phe Glu Ser Leu Ser Ser Ile Leu Lys Leu Pro Gly Lys Lys
625 630 635 640
Thr Phe Asn Asn Met Asp Arg Asp Phe Leu Glu Lys Arg Lys Lys Asp
645 650 655
Leu Asn Ala Tyr Leu Gln Leu Leu Leu Ala Pro Glu Met Met Lys Ala
660 665 670
Ser Pro Ala Leu Ala His Tyr Val Tyr Asp Phe Leu Glu Asn Lys Ala
675 680 685
Tyr Ser Lys Gly Lys Gly Asp Phe Ala Arg Lys Met Asp Thr Phe Val
690 695 700
Asn Pro Leu Arg Asn Ser Met Arg Asn Val Ser Asn Ala Val Lys Ser
705 710 715 720
Leu Pro Asp Ser Leu Ala Glu Gly Met Thr Lys Met Ser Asp Asn Met
725 730 735
Gly Lys Met Ser Glu Arg Leu Gly Gln Asp Ile Lys Gln Ser Phe Phe
740 745 750
Lys Val Pro Pro Leu Ile Pro Lys Thr Asp Ser Asp Pro Glu His Arg
755 760 765
Arg Val Ser Ala Gln Leu Asp Asp Asn Val Asp Asp Asn Ile Pro Leu
770 775 780
Arg Val Met Leu Leu Leu Met Asp Glu Val Phe Asp Leu Lys Glu Arg
785 790 795 800
Asn Gln Trp Leu Arg Arg Asn Ile Lys Asn Leu Leu Gln Gln Leu Ile
805 810 815
Arg Ala Thr Tyr Gly Asp Thr Ile Asn Arg Lys Ile Val Asp His Val
820 825 830
Asp Trp Met Thr Ser Pro Glu Gln Val Ala Asp Ser Val Lys Arg Phe
835 840 845
Arg Asp Ala Phe Trp Pro Asn Gly Ile Leu Ala Glu Ala Val Pro Cys
850 855 860
Arg Asp Lys Ser Ile Arg Met Arg Thr Arg Val Ala Gly Lys Thr Lys
865 870 875 880
Leu Leu Ala Ile Met Pro Asp Glu Leu Lys His Ile Ile Gly Ala Glu
885 890 895
Thr Thr Arg Lys Gly Ile Leu Arg Val Phe Glu Met Phe Gln His Asn
900 905 910
Gln Leu Asn Arg Arg Met Val Tyr Val Phe Leu Glu Gly Phe Leu Glu
915 920 925
Thr Leu Phe Pro Gln Tyr Lys Phe Arg Glu Leu Phe Asn Lys Leu His
930 935 940
Ser Arg Ser Lys Gln Met Gln Lys Tyr Lys Gln Lys Leu Gln Thr Thr
945 950 955 960
Gln Ala Pro Ser Leu Gln Lys Arg
965
<210> 57
<211> 292
<212> PRT
<213> 智人
<400> 57
Met Gly Glu Lys Ser Glu Asn Cys Gly Val Pro Glu Asp Leu Leu Asn
1 5 10 15
Gly Leu Lys Val Thr Asp Thr Gln Glu Ala Glu Cys Ala Gly Pro Pro
20 25 30
Val Pro Asp Pro Lys Asn Gln His Ser Gln Ser Lys Leu Leu Arg Asp
35 40 45
Asp Glu Ala His Leu Gln Glu Asp Gln Gly Glu Glu Glu Cys Phe His
50 55 60
Asp Cys Ser Ala Ser Phe Glu Glu Glu Pro Gly Ala Asp Lys Val Glu
65 70 75 80
Asn Lys Ser Asn Glu Asp Val Asn Ser Ser Glu Leu Asp Glu Glu Tyr
85 90 95
Leu Ile Glu Leu Glu Lys Asn Met Ser Asp Glu Glu Lys Gln Lys Arg
100 105 110
Arg Glu Glu Ser Thr Arg Leu Lys Glu Glu Gly Asn Glu Gln Phe Lys
115 120 125
Lys Gly Asp Tyr Ile Glu Ala Glu Ser Ser Tyr Ser Arg Ala Leu Glu
130 135 140
Met Cys Pro Ser Cys Phe Gln Lys Glu Arg Ser Ile Leu Phe Ser Asn
145 150 155 160
Arg Ala Ala Ala Arg Met Lys Gln Asp Lys Lys Glu Met Ala Ile Asn
165 170 175
Asp Cys Ser Lys Ala Ile Gln Leu Asn Pro Ser Tyr Ile Arg Ala Ile
180 185 190
Leu Arg Arg Ala Glu Leu Tyr Glu Lys Thr Asp Lys Leu Asp Glu Ala
195 200 205
Leu Glu Asp Tyr Lys Ser Ile Leu Glu Lys Asp Pro Ser Ile His Gln
210 215 220
Ala Arg Glu Ala Cys Met Arg Leu Pro Lys Gln Ile Glu Glu Arg Asn
225 230 235 240
Glu Arg Leu Lys Glu Glu Met Leu Gly Lys Leu Lys Asp Leu Gly Asn
245 250 255
Leu Val Leu Arg Pro Phe Gly Leu Ser Thr Glu Asn Phe Gln Ile Lys
260 265 270
Gln Asp Ser Ser Thr Gly Ser Tyr Ser Ile Asn Phe Val Gln Asn Pro
275 280 285
Asn Asn Asn Arg
290
<210> 58
<211> 8
<212> PRT
<213> 智人
<400> 58
Asp Arg Val Tyr Ile His Pro Phe
1 5
<210> 59
<211> 30
<212> PRT
<213> 智人
<400> 59
Met Asp Ser Lys Gly Ser Ser Gln Lys Gly Ser Arg Leu Leu Leu Leu
1 5 10 15
Leu Val Val Ser Asn Leu Leu Leu Cys Gln Gly Val Val Ser
20 25 30
<210> 60
<211> 10
<212> PRT
<213> 智人
<400> 60
Gly Cys Met Ser Cys Lys Cys Val Leu Ser
1 5 10
<210> 61
<211> 10
<212> PRT
<213> 智人
<400> 61
Gly Cys Met Gly Leu Pro Cys Val Val Met
1 5 10
<210> 62
<211> 10
<212> PRT
<213> 智人
<400> 62
Cys Val Lys Ile Lys Lys Cys Ile Ile Met
1 5 10
<210> 63
<211> 14
<212> PRT
<213> 智人
<400> 63
Lys Lys Lys Lys Lys Lys Ser Lys Thr Lys Cys Val Ile Met
1 5 10
Claims (67)
1.一种以Gα蛋白亚基家族选择性方式测量G蛋白活化调节的系统，所述系统包括表达以下的细胞：
(i)第一组分，包括用生物发光供体分子或荧光接受体分子标记的Gα亚基相互作用多肽(GASIP)；
其中：
如果所述Gα蛋白亚基家族是Gi，则所述GASIP包含与Gi特异性结合的蛋白结构域；
如果所述Gα蛋白亚基家族是Gq，则所述GASIP包含与Gq特异性结合的蛋白结构域；和
如果所述Gα蛋白亚基家族是G12/13，则所述GASIP包含与G12/13特异性结合的蛋白结构域；以及
(ii)第二组分，包括用生物发光供体分子或荧光接受体分子标记的质膜(PM)靶向部分、胞内体靶向部分或高尔基体靶向部分；
其中，如果所述GASIP用所述荧光接受体分子标记，则所述PM靶向部分、胞内体靶向部分或高尔基体靶向部分用所述生物发光供体分子标记，以及如果所述GASIP用所述生物发光供体分子标记，则所述PM靶向部分、胞内体靶向部分或高尔基体靶向部分用所述荧光接受体分子标记。
2.根据权利要求1所述的系统，其中，所述与Gi特异性结合的蛋白结构域是Rap1GAP的G蛋白结合结构域或G蛋白信号转导调节物(RGS)蛋白的G蛋白结合结构域。
3.根据权利要求2所述的系统，其中，所述GASIP包含所述Rap1GAP的G蛋白结合结构域。
4.根据权利要求3所述的系统，其中，所述Rap1GAP的G蛋白结合结构域包含Rap1GAP(SEQ ID NO：8)的1至442位的残基或其变体，其中在位置437、439和441的一个或多个丝氨酸残基突变或缺失。
5.根据权利要求4所述的系统，其中，所述Rap1GAP的G蛋白结合结构域包含Rap1GAP的1至420位或1至436位的残基。
6.根据权利要求4所述的系统，其中，在位置437、439和441的所有三个丝氨酸残基都突变。
7.根据权利要求4或6所述的系统，其中，所述丝氨酸残基被丙氨酸替换。
8.根据权利要求2所述的系统，其中，所述GASIP包含RGS蛋白的G蛋白结合结构域。
9.根据权利要求8所述的系统，其中，所述RGS蛋白是RGS17、RGS19或RGS20。
10.根据权利要求9所述的系统，其中，所述G蛋白结合结构域包含RGS17(SEQ ID NO：17)的64至210位的残基、RGS19(SEQ ID NO：18)的70-217位的残基或RGS20(SEQ ID NO：19)的242-388位的残基。
11.根据权利要求1所述的系统，其中，所述与Gq特异性结合的蛋白结构域是P63RhoGEF的G蛋白结合结构域或GRK2的G蛋白结合结构域。
12.根据权利要求11所述的系统，其中，所述GASIP包含所述P63RhoGEF的G蛋白结合结构域。
13.根据权利要求12所述的系统，其中，所述P63RhoGEF的G蛋白结合结构域包含P63RhoGEF(SEQ ID NO：25)的295至502位的残基。
14.根据权利要求11所述的系统，其中，所述GASIP包含所述GRK2的G蛋白结合结构域。
15.根据权利要求14所述的系统，其中，所述GRK2的G蛋白结合结构域包含GRK2(SEQ IDNO：27)的30至203位的残基。
16.根据权利要求1所述的系统，其中，所述与G12/13特异性结合的蛋白结构域是PDZRhoGEF的G蛋白结合结构域或P115RhoGEF的G蛋白结合结构域。
17.根据权利要求16所述的系统，其中，所述GASIP包含所述PDZRhoGEF的G蛋白结合结构域。
18.根据权利要求17所述的系统，其中，所述PDZRhoGEF的G蛋白结合结构域包含PDZRhoGEF(SEQ ID NO：21)的281至483位的残基。
19.根据权利要求16所述的系统，其中，所述GASIP包含所述P115RhoGEF的G蛋白结合结构域。
20.根据权利要求19所述的系统，其中，所述P115RhoGEF的G蛋白结合结构域包含P115RhoGEF(SEQ ID NO：23)的1至244位的残基。
21.根据权利要求1至20中任一项所述的系统，其中，所述GASIP用所述生物发光供体分子标记，并且所述PM靶向部分、胞内体靶向部分或高尔基体靶向部分用所述荧光接受体分子标记。
22.根据权利要求1至21中任一项所述的系统，其中，所述PM靶向部分是定位于所述PM的PM蛋白或其片段。
23.根据权利要求22所述的系统，其中，所述PM蛋白或其片段包含(a)棕榈酰化、肉豆蔻酰化和/或异戊烯化信号序列和/或(b)多碱性序列。
24.根据权利要求22或23所述的系统，其中，所述PM靶向部分包含氨基酸序列GCMSCKCVLS(SEQ ID NO：60)、GCMGLPCVVM(SEQ ID NO：61)、CVKIKKCIIM(SEQ ID NO：62)、KKKKKKSKTKCVIM(SEQ ID NO：63)或KNGKKKRKSLAKRIRERCCIL(SEQ ID NO：45)、CMSCKCCIL(SEQ ID NO：4)或SPKKGLLQRLFKRQHQNNSKS(SEQ ID NO：5)。
25.根据权利要求24所述的系统，其中，所述PM靶向部分包含氨基酸序列GKKKKKKSKTKCVIM(SEQ ID NO：1)。
26.根据权利要求1至21中任一项所述的系统，其中，所述胞内体靶向部分是定位于所述胞内体的胞内体蛋白或其片段。
27.根据权利要求26所述的系统，其中，所述胞内体蛋白或其片段包含FYVE结构域。
28.根据权利要求27所述的系统，其中，所述胞内体靶向部分包含人endofin的FYVE结构域。
29.根据权利要求28所述的系统，其中，所述胞内体靶向部分包含人endofin(SEQ IDNO：39)的739至806位的残基。
30.根据权利要求1至21中任一项所述的系统，其中，所述高尔基体靶向部分是定位于所述高尔基体的高尔基体蛋白或其片段。
31.根据权利要求30所述的系统，其中，所述高尔基体靶向部分是定位于所述高尔基体的eNOS1或其片段。
32.根据权利要求31所述的系统，其中，所述高尔基体靶向部分包含人eNOS1(SEQ IDNO：46)的1至73位的残基。
33.根据权利要求1至32中任一项所述的系统，其中，所述第一组分还包括在(i)所述GASIP和(ii)所述生物发光供体分子或荧光接受体分子之间的接头。
34.根据权利要求1至33中任一项所述的系统，其中，所述第二组分还包括在(i)所述PM靶向部分、胞内体靶向部分或高尔基体靶向部分和(ii)所述生物发光供体分子或荧光接受体分子之间的接头。
35.根据权利要求33或34所述的系统，其中，所述接头是5至25个氨基酸连接的肽。
36.根据权利要求1至35中任一项所述的系统，还包括第三组分，所述第三组分是通过所述G蛋白信号转导的细胞表面受体。
37.根据权利要求36所述的系统，其中，所述细胞表面受体是GPCR、RTK或整合素受体。
38.根据权利要求1至37中任一项所述的系统，其中，所述G蛋白活化是G蛋白偶联受体(GPCR)介导的G蛋白活化。
39.根据权利要求1至38中任一项所述的系统，还包括第四组分，所述第四组分为重组Gα亚基多肽。
40.根据权利要求39所述的系统，其中，所述G蛋白活化是非受体鸟嘌呤核苷酸交换因子(GEF)介导的G蛋白活化，其中所述重组Gα亚基多肽在所述Gα亚基多肽的羧基(C)末端结构域中包含至少一个突变，并且其中所述C末端结构域对应于所述Gα亚基多肽的最后七个残基。
41.根据权利要求40所述的系统，其中，所述突变是至少最后两个C末端残基的缺失或替换。
42.根据权利要求41所述的系统，其中，所述突变是至少最后五个C末端残基的缺失或替换。
43.根据权利要求40所述的系统，其中，所述突变是在所述C末端结构域中至少一个保守亮氨酸残基的缺失或替换。
44.根据权利要求43所述的系统，其中，所述突变是在所述C末端结构域中最后一个保守亮氨酸残基的替换。
45.根据权利要求44所述的系统，其中，所述替换是亮氨酸至天冬氨酸、亮氨酸至脯氨酸或亮氨酸至精氨酸的替换。
46.根据权利要求40至45中任一项所述的系统，其中，所述GEF为GIV/Girdin，并且其中所述GASIP包含如权利要求1至7中任一项所限定的Rap1GAP的G蛋白结合结构域。
47.根据权利要求46所述的系统，其中，所述GEF被受体酪氨酸激酶(RTK)活化。
48.根据权利要求1至47中任一项所述的系统，其中，所述生物发光供体分子是荧光素酶，优选地是海肾荧光素酶蛋白(rLuc)。
49.根据权利要求1至48中任一项所述的系统，其中，所述荧光接受体分子是绿色荧光蛋白(GFP)，优选地是海肾GFP(rGFP)。
50.一种或多种核酸，编码权利要求1至49中任一项的系统的所述第一组分和/或第二组分。
51.根据权利要求50所述的一种或多种核酸，包括编码权利要求1至49中任一项的系统的所述第一组分的第一核酸和编码权利要求1至49中任一项的系统的所述第二组分的第二核酸。
52.一种或多种载体，包括权利要求50或51的一种或多种核酸。
53.宿主细胞，表达权利要求1至49中任一项所限定的系统的组分。
54.一种用于确定剂是否调节感兴趣的G蛋白活化的方法，所述方法包括：
(a)使权利要求1至49中任一项的系统与所述生物发光供体分子的底物接触；以及
(b)在所述剂存在和不存在下测量所述系统中的BRET信号；
其中，在所述剂存在下的所述BRET信号相对于其不存在下的差异表明所述剂调节所述感兴趣的G蛋白活化。
55.根据权利要求54所述的方法，其中，所述感兴趣的G蛋白是所述Gi蛋白亚基家族，并且其中所述GASIP包含权利要求1至7中任一项所限定的Rap1GAP的G蛋白结合结构域。
56.根据权利要求49所述的方法，其中，所述系统包括所述Gi蛋白亚基家族的重组Gα亚基多肽。
57.根据权利要求56所述的方法，其中，所述方法使用多个系统执行，并且其中每个所述系统包括所述Gi蛋白亚基家族的不同重组Gα亚基多肽。
58.根据权利要求54所述的方法，其中，所述感兴趣的G蛋白是所述Gq蛋白亚基家族，其中所述GASIP包含权利要求1和11至15中任一项所限定的P63RhoGEF的G蛋白结合结构域或GRK2的G蛋白结合结构域。
59.根据权利要求58所述的方法，其中，所述系统包括所述Gq蛋白亚基家族的重组Gα亚基多肽。
60.根据权利要求59所述的方法，其中，所述方法使用多个系统执行，并且其中每个所述系统包括所述Gq蛋白亚基家族的不同重组Gα亚基多肽。
61.根据权利要求54所述的方法，其中，所述感兴趣的G蛋白是所述G12/13蛋白亚基家族，其中所述GASIP包含权利要求1和16至20中任一项所限定的PDZRhoGEF的G蛋白结合结构域或P115RhoGEF的G蛋白结合结构域。
62.根据权利要求61所述的方法，其中，所述系统包括所述G12/13蛋白亚基家族的重组Gα亚基多肽。
63.根据权利要求62所述的方法，其中，所述方法使用多个系统执行，并且其中每个所述系统包括所述G12/13蛋白亚基家族的不同重组Gα亚基多肽。
64.一种用于确定剂是否调节非受体鸟嘌呤核苷酸交换因子(GEF)介导的G蛋白活化的方法，所述方法包括：
(a)使权利要求40至47中任一项的系统与所述生物发光供体分子的底物接触；以及
(b)在所述剂存在和不存在下测量所述系统中的BRET信号；
其中，在所述剂存在下的所述BRET信号相对于其不存在下的差异表明所述剂调节非受体GEF介导的G蛋白活化。
65.根据权利要求54至64中任一项所述的方法，其中，使用读板仪或通过显微术来测量所述BRET信号。
66.根据权利要求54至65中任一项所述的方法，其中，所述底物是腔肠素底物。
67.根据权利要求66所述的方法，其中，所述腔肠素底物是甲氧基e腔肠素。
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| WO2016058094A1 (en) * | 2014-10-14 | 2016-04-21 | Université de Montréal | BIOSENSOR BASED ON GβΥ-INTERACTING PROTEINS TO MONITOR G-PROTEIN ACTIVATION |
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| AU765703B2 (en) | 1998-03-27 | 2003-09-25 | Bruce J. Bryan | Luciferases, fluorescent proteins, nucleic acids encoding the luciferases and fluorescent proteins and the use thereof in diagnostics, high throughput screening and novelty items |
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| CN117024557B (zh) * | 2023-10-10 | 2024-01-30 | 天津外泌体科技有限公司 | 14-3-3蛋白theta异构体作为细胞外囊泡支架蛋白的应用和细胞外囊泡 |
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