TW202300642A - 獲得誘導型多能幹細胞之方法 - Google Patents
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Abstract
本發明提供自造血譜系細胞獲得誘導型多能幹細胞之方法。
Description
細胞療法提供治療各種疾病及病狀之極大前景。於細胞療法中,將自體或同種異體細胞移植至患者中以替換或修復可自大批醫學病狀(包括遺傳病症、癌症、神經學病症、心臟病或眼睛相關組織)產生之缺陷或受損組織或細胞。多能幹細胞尤其可用於細胞療法中,包含從體細胞產生之多能幹細胞。K. Takahashi及S. Yamanaka之有創造力的工作證實,來自經表現四種再程式化轉錄因子Oct3/4、Klf4、Sox2及c-Myc之逆轉錄病毒載體轉導之小鼠纖維母細胞誘導多能幹細胞(
Cell(2006) 126:663-76)。然而,逆轉錄病毒載體可引起宿主基因組之插入突變及因此非用於臨床情境之理想載體。
因此已嘗試基於RNA之方法用於將再程式化因子引入體細胞中。一種此方法利用基於α病毒之病毒RNA複製子。構成披膜病毒科(
Togaviridae)中之超過30種病毒之屬之α病毒為脂質包膜之正義RNA病毒。新世界α病毒包括東部、西部及委內瑞拉馬腦炎病毒(各自為EEEV、WEEV及VEEV)及見於北美及南美。舊世界α病毒包括基孔肯雅(chikungunya) (CHIK)、辛德畢斯(Sindbis)、羅斯河(Ross River)及歐尼恩(O’nyong-nyong)病毒。
α病毒含有長度約14 kb之正義單股RNA基因組。於進入宿主細胞後,α病毒粒子經歷解組裝及釋放基因組RNA至細胞之細胞質中。病毒基因組之轉譯產生非結構多蛋白P1234,其隨後藉由蛋白酶裂解以產生非結構蛋白(nsP1、nsP2、nsP3及nsP4)。該等非結構蛋白涉及RNA複製及轉錄。亞基因組RNA (26S RNA)亦透過轉錄自病毒基因組產生。26S RNA之轉譯產生結構多蛋白,其經裂解以產生結構蛋白(例如,針對VEEV之C、E3、E2、6K及E1)。該等結構蛋白涉及發芽及病毒衣殼化。參見,例如,Shin等人,
PNAS(2012) 109(41):16534-9;Jose等人,
Future Microbiol. (2009) 4:837-56;Hardy及Strauss,
J Virol.(1989) 63(11):4653-64;Melancon及Garoff,
J Virol.(1987) 61(5):1301-9;及Strauss等人,
Virology(1984) 133(1):92-110);及Glanville等人,
PNAS(1976) 73(9):3059-63)。
α病毒複製子不涉及用於複製之DNA中間體及因此提供對若干其他常用病毒載體(包括逆轉錄病毒載體)之更安全替代(Yoshioka等人,
Cell Stem Cell.(2013) 13(2):246-54;Yoshioka及Dowdy,
PLOS ONE(2017) 12:e0182018)。α病毒及特定言之VEE已經開發作為載體以攜帶編碼再程式化體細胞中之外源轉錄因子至誘導型多能幹細胞(iPSC)之基因。然而,僅於纖維母細胞及血液外生內皮細胞(BOEC)中嘗試此方法。無一種細胞類型係臨床上特別吸引人的。自體纖維母細胞係自患者之皮膚穿刺獲得,其為侵入性且疼痛的。儘管源自外周血,但是BOEC為罕見細胞且需要費力且耗時方法來建立。
再程式化之另一無整合方法利用附加型DNA質體載體。Wen等人使用此方法以將外周血單核細胞再程式化為iPSC(
Stem Cell Rep.(2016) 6:873-84)。但是該方法需要仔細校準透過多重載體引入之各種再程式化因子之水平。
亦已使用仙台(Sendai)病毒載體攜帶編碼再程式化因子之基因。該等仙台載體為負股副黏病毒(
Paramyxoviruses);必須將載體封裝至病毒粒子中。此方法更複雜。其涉及封裝細胞系且可將外來劑引入載體產物。
因此,存在對自外周血球獲得iPSC之有效且安全方法之需求。
本發明提供一種自造血譜系之起始細胞獲得誘導型多能幹細胞(iPSC)群體之方法。該方法包括:將編碼BCL-xL及選自OCT家族蛋白、KLF家族蛋白、MYC家族蛋白、SOX家族蛋白、LIN28蛋白、NANOG蛋白及p53顯性負性蛋白之一或多種附加再程式化因子之α病毒RNA表現構築體引入該等起始細胞,及培養該等起始細胞以允許BCL-xL及該一或多種附加再程式化因子之表現,從而誘導該等起始細胞及其後代再程式化為iPSC。
於一個態樣中,本發明提供自造血譜系之起始細胞獲得之誘導型多能幹細胞(iPSC)群體,該等起始細胞經編碼BCL-xL及選自OcT家族蛋白、KLF家族蛋白、MyC家族蛋白、SOX家族蛋白、LIN28蛋白、NANOG蛋白及p53顯性負性蛋白之一或多種附加再程式化因子之α病毒RNA表現構築體轉染。
該等起始細胞可為(例如)人類起源之造血幹細胞、紅血球系祖細胞、淋巴系祖細胞、外周血單核細胞、T淋巴細胞、B淋巴細胞、巨噬細胞、單核細胞、嗜中性白血球、嗜酸性白血球或樹突狀細胞。於一些實施例中,該等起始細胞為藉由在存在促紅血球生成素(EPO)、幹細胞因子(SCF)及IL-3下培養外周血單核細胞(PBMC)視情況持續5至10天或6至7天獲得之紅血球系祖細胞。於另外實施例中,該等PBMC係在存在0.5至5 IU/ml EPO、50至200 ng/mL SCF及1至10 ng/mL IL-3下培養。
於一些實施例中,該RNA表現構築體透過電穿孔引入起始細胞中。於另外實施例中,該等起始細胞係在電穿孔之前利用B18R蛋白培養。
於另一態樣中,本發明提供編碼BCL-xL及選自OCT家族蛋白、KLF家族蛋白、MYC家族蛋白、SOX家族蛋白、LIN28蛋白、NANOG蛋白及p53顯性負性蛋白之一或多種附加再程式化因子之α病毒RNA表現構築體。
亦提供包含本文中之α病毒RNA表現構築體之編碼序列之DNA載體,及包含本文中之α病毒RNA表現構築體或DNA載體之宿主細胞(例如,人類細胞)。
於一些實施例中,該α病毒RNA表現構築體係自我複製及包含足以致使構築體自我複製之一或多種非結構蛋白之基因(例如,nsP1、nsP2、nsP3及nsP4基因)。於另外實施例中,該α病毒RNA表現構築體為委內瑞拉馬腦炎病毒(VEEV) RNA表現構築體及包含VEEV nsP1、nsP2、nsP3及nsP4基因。於某些實施例中,該VEEV RNA表現構築體含有來自野生型VEEV基因組之該(等)對應區域之一或多個(例如,兩個或更多個、三個或更多個、四個或更多個、五個或更多個、或六個或更多個)突變。
於一些實施例中,該OCT家族蛋白為OCT4 (例如,人類OCT4);該KLF家族蛋白為KLF4 (例如,人類KLF4);該SOX家族蛋白為SOX2 (例如,人類SOX2);該LIN28蛋白為LIN28B (例如,人類LIN28B);及/或該MYC家族蛋白為c-MYC (例如,人類c-MYC)。
於特定實施例中,該BCL-xL蛋白包含SEQ ID NO:1或與其至少95%相同之胺基酸序列;該OCT4蛋白包含SEQ ID NO:3或與其至少95%相同之胺基酸序列;該KLF4蛋白包含SEQ ID NO:5或與其至少95%相同之胺基酸序列;該SOX2蛋白包含SEQ ID NO:7或與其至少95%相同之胺基酸序列;及/或該c-MYC蛋白包含SEQ ID NO:9或與其至少95%相同之胺基酸序列。
於一些實施例中,BCL-xL及該一或多種附加再程式化因子之該等編碼序列被2A肽之編碼序列或內部核糖體進入位點(IRES)分隔開。於一些實施例中,BCL-xL及該一或多種附加再程式化因子之該等編碼序列係在共同(common)啟動子(例如,26S啟動子)之轉錄控制下。
於一些實施例中,本文中之α病毒RNA表現構築體指導OCT家族蛋白、SOX家族蛋白、BCL-xL及MYC家族蛋白,及視情況KLF家族蛋白之表現。
於另一態樣中,本發明提供一種於活體外獲得分化細胞之方法,其包括在存在分化促進劑下培養本文中獲得之iPSC。亦提供藉由自iPSC分化獲得之分化細胞。於一些實施例中,本文中獲得之分化細胞為視情況選自T細胞、表現嵌合抗原受體(CAR)之T細胞、抑制性T細胞、骨髓細胞、樹突狀細胞及免疫抑制性巨噬細胞之人類免疫細胞;視情況選自多巴胺能神經元、小膠質細胞、少突膠質細胞、星形膠質細胞、皮質神經元、脊髓或動眼神經元、腸神經元、基板衍生細胞、施旺(Schwann)細胞及三叉神經或感覺神經元之人類神經系統之細胞;視情況選自心肌細胞、內皮細胞及結節細胞之人類心血管系統之細胞;視情況選自肝細胞、膽管細胞及胰β細胞之人類代謝系統之細胞,或視情況選自視網膜色素上皮細胞、感光錐形細胞、感光桿狀細胞、雙極細胞、神經節細胞之人類眼部系統之細胞、免疫細胞、神經細胞、心血管細胞或代謝系統之細胞。於特定實施例中,該分化細胞為外胚層譜系(例如,神經元)。於其他特定實施例中,該分化細胞為中胚層譜系(例如,心肌細胞)。
本發明亦提供醫藥組合物,其包含本文中獲得之分化細胞及醫藥上可接受之載劑。本發明亦提供治療有需要患者之方法,其包括向該患者投與該醫藥組合物;分化細胞用於製造用於治療有需要患者之藥劑之用途;及分化細胞或醫藥組合物用於治療有需要患者。
本發明之其他特徵、目標及優點於跟隨實施方式中顯然。然而,應瞭解,雖然指示本發明之實施例及態樣,但是實施方式僅經由說明提供,非限制性。本發明之範圍內之各種變化及修改將自實施方式對熟習此項技術者變得顯然。
序列表
本申請案含有以ASCII格式電子提交之序列表及其全文係以引用的方式併入本文中。在2022年3月23日創建之該ASCII副本被命名為025450_TW014_SL.txt及大小為62,837字節。
本發明描述將血液衍生細胞(例如,紅血球系祖細胞)再程式化至誘導型多能幹細胞(iPSC)之改良之方法。此等方法涉及使用編碼再程式化因子BCL-xL及一或多種(例如,一種、兩種、三種、四種、五種、六種、七種或所有八種)附加再程式化因子(例如,OCT家族成員、KLF家族成員、SOX家族成員、MYC蛋白、NANOG蛋白、GLIS家族成員、LIN28蛋白及p53顯性負性)之α病毒(例如,VEEV) RNA表現載體(即,或表現構築體)。可將α病毒RNA表現構築體透過本文中所述之改良之方法引入血球中。經轉染之細胞於小於3週內分化成可收穫iPSC。
外周血為用於將體細胞再程式化至iPSC之易得細胞資源。因此,本發明方法極大地提高產生iPSC之效率。由於使用不融合至宿主細胞之基於RNA之表現載體,藉由本發明方法獲得之iPSC具有較藉由先前方法使用逆轉錄病毒載體獲得之彼等更安全臨床特性。
I. 表現再程式化因子之 α 病毒 RNA 構築體
本發明之α病毒RNA表現構築體為自我複製RNA複製子。自我複製RNA複製子或構築體係指表現非結構蛋白基因使得其可指導其於宿主細胞中之自我複製之RNA分子。其可包含5’及3’ α病毒複製識別序列,對RNA複製及轉錄必需之α病毒非結構蛋白之編碼序列(例如,VEE nsP1、nsP2、nsP3及nsP4)及聚腺苷酸化信號序列。其可另外含有指導異源RNA序列(諸如編碼再程式化因子者)之表現之一或多個元件(例如,IRES序列、核心或微型啟動子及類似者)。
於一些實施例中,α病毒RNA構築體為包含以下之VEEV RNA複製子:(i)對複製必要之VEEV非結構蛋白之基因,(ii) 5’及3’病毒複製識別序列,(iii)表現盒,諸如用於表現所關注之再程式化因子之多順反子性表現盒;及(iv)聚腺苷酸化尾。亦參見Yoshioka 2013及2017,見上;及WO 2013/177133,及美國專利10,793,833、10,370,646及9,862,930。複製子可缺少VEEV結構蛋白基因。自我複製VEE RNA構築體可在有限數目之細胞分裂期間在經轉染細胞內部複製。RNA構築體藉由降解損失之時間可進一步藉由自培養基收回之B18R調節。
示例性VEEV RNA構築體表現BCL-xL及其他再程式化因子。再程式化因子為當於體細胞中過度表現時,誘導細胞自分化狀態過渡至多能狀態之蛋白質。本文中所用之再程式化因子可為人類蛋白或保留所需生物效應之其修飾形式。
A. BCL-xL
人類BCL-xL藉由
BCL2L1基因編碼。示例性人類BCL-xL胺基酸序列可見於UniProt寄存編號Q07817且具有下列胺基酸序列:
此蛋白質之功能類似物,即,具有相同或實質上相同生物功能(例如,保留蛋白質之轉錄因子功能之70%或更多,80%或更多,90%或更多,95%或更多,或98%或更多)之分子作為BCL-xL蛋白包含於本發明。例如,該功能類似物可為以上蛋白質之同功異型物或變異體,例如,含有以上蛋白質之一部分具有或不具有另外胺基酸殘基及/或含有相對於以上蛋白質之突變。於一些實施例中,該功能類似物具有與SEQ ID NO:1至少90%、95%、98%或99%之序列同一性。兩個胺基酸序列(或兩個核酸序列)之同一性%可藉由(例如) BLAST®使用默認參數(在美國醫學國家中心之國家圖書館(U.S. National Library of Medicine’s National Center)生物技術資訊網站可得)獲得。於一些實施例中,出於比較目的比對之參考序列之長度為參考序列之至少30% (例如,至少40%、50%、60%、70%、80%或90%)。
示例性VEEV構築體可包含Oct家族蛋白(例如,OCT1、OCT2、OCT4、OCT6、OCT7、OCT8、OCT9及OCT11)之編碼序列。參見,例如,美國專利8,278,104及WO 2013/177133。人類OCT4係藉由
POU5F1基因編碼。示例性人類OCT4胺基酸序列可見於UniProt寄存編號Q01860且具有下列胺基酸序列:
此蛋白質之功能類似物,即,具有相同或實質上相同生物功能(例如,保留蛋白質之轉錄因子功能之70%或更多,80%或更多,90%或更多,95%或更多,或98%或更多)之分子作為OCT4蛋白包含於本發明。例如,該功能類似物可為以上蛋白質之同功異型物或變異體,例如,含有以上蛋白質之一部分具有或不具有另外胺基酸殘基及/或含有相對於以上蛋白質之突變。於一些實施例中,該功能類似物具有與SEQ ID NO:3至少90%、95%、98%或99%之序列同一性。
示例性VEEV構築體可包含KLF家族蛋白(例如,KLF1、KLF2、KLF3、KLF4、KLF5、KLF6、KLF7、KLF8、KLF9、KLF10、KLF11、KLF12、KLF13、KLF14、KLF15、KLF16及KLF17)之編碼序列。參見,例如,美國專利8,278,104及WO 2013/177133。人類KLF4係藉由
KLF4基因編碼。示例性人類KLF4胺基酸序列可見於UniProt寄存編號O43474且具有下列胺基酸序列:
此蛋白質之功能類似物,即,具有相同或實質上相同生物功能(例如,保留蛋白質之轉錄因子功能之70%或更多,80%或更多,90%或更多,95%或更多,或98%或更多)之分子作為KLF4蛋白包含於本發明。例如,該功能類似物可為以上蛋白質之同功異型物或變異體,例如,含有以上蛋白質之一部分具有或不具有另外胺基酸殘基及/或含有相對於以上蛋白質之突變。於一些實施例中,該功能類似物具有與SEQ ID NO:5至少90%、95%、98%或99%之序列同一性。於一些實施例中,ESSRB可代替KLF蛋白使用。於一些實施例中,該KLF4蛋白為SEQ ID NO:5之同功異型物且包含以下所示之SEQ ID NO:6之胺基酸殘基2-471。
示例性VEEV構築體可包含SOX家族蛋白(例如,SOX1、SOX2、SOX3、SOX4、SOX5、SOX6、SOX7、SOX8、SOX9、SXO10、SOX11、SOX12、SOX13、SOX14、SOX15、SOX17、SOX18、SOX21及SOX30)之編碼序列。參見,例如,美國專利8,278,104及WO 2013/177133。人類SOX2係藉由
SOX2基因編碼。示例性人類SOX2胺基酸序列可見於UniProt寄存編號P48431且具有下列胺基酸序列:
此蛋白質之功能類似物,即,具有相同或實質上相同生物功能(例如,保留蛋白質之轉錄因子功能之70%或更多,80%或更多,90%或更多,95%或更多,或98%或更多)之分子作為SOX2蛋白包含於本發明。例如,該功能類似物可為以上蛋白質之同功異型物或變異體,例如,含有以上蛋白質之一部分具有或不具有另外胺基酸殘基及/或含有相對於以上蛋白質之突變。於一些實施例中,該功能類似物具有與SEQ ID NO:7至少90%、95%、98%或99%之序列同一性。
示例性VEEV構築體可包含MYC家族蛋白(例如,c-MYC、n-MYC及l-MYC)之編碼序列。參見,例如,美國專利8,278,104。人類c-MYC係藉由
MYC基因編碼。示例性人類c-MYC胺基酸序列可見於UniProt寄存編號P01106且具有下列胺基酸序列:
此蛋白質之功能類似物,即,具有相同或實質上相同生物功能(例如,保留蛋白質之轉錄因子功能之70%或更多,80%或更多,90%或更多,95%或更多,或98%或更多)之分子作為c-MYC蛋白包含於本發明。例如,該功能類似物可為以上蛋白質之同功異型物或變異體,例如,含有以上蛋白質之一部分具有或不具有另外胺基酸殘基及/或含有相對於以上蛋白質之突變。於一些實施例中,該功能類似物具有與SEQ ID NO:9至少90%、95%、98%或99%之序列同一性。於一些實施例中,具有降低之轉形活性之MYC變異體可代替c-MYC使用。參見,例如,美國專利9,005,967。
示例性VEEV構築體可包含GLIS家族蛋白(例如,GLIS1、GLIS2及GLIS3)之編碼序列。參見,例如,美國專利8,951,801。人類GLIS1係藉由
GLIS1基因編碼。示例性人類GLIS1胺基酸序列可見於UniProt寄存編號Q8NBF1且具有下列胺基酸序列:
此蛋白質之功能類似物,即,具有相同或實質上相同生物功能(例如,保留蛋白質之轉錄因子功能之70%或更多,80%或更多,90%或更多,95%或更多,或98%或更多)之分子作為GLIS1蛋白包含於本發明。例如,該功能類似物可為以上蛋白質之同功異型物或變異體,例如,含有以上蛋白質之一部分具有或不具有另外胺基酸殘基及/或含有相對於以上蛋白質之突變。於一些實施例中,該功能類似物具有與SEQ ID NO:11至少90%、95%、98%或99%之序列同一性。
G. NANOG
本發明VEEV構築體可包含NANOG之編碼序列。參見,例如,美國專利9,506,039。人類NANOG係藉由
NANOG基因編碼。示例性人類NANOG胺基酸序列可見於UniProt寄存編號Q9H9S0且具有下列胺基酸序列:
此序列之功能類似物,即,具有以上蛋白質之相同或實質上相同生物功能(例如,保留蛋白質之轉錄因子功能之70%或更多,80%或更多,90%或更多,95%或更多,或98%或更多)之分子作為NANOG蛋白包含於本發明。例如,該功能類似物可為以上蛋白質之同功異型物或變異體,例如,含有以上蛋白質之一部分具有或不具有另外胺基酸殘基及/或含有相對於以上蛋白質之突變。於一些實施例中,該功能類似物具有與SEQ ID NO:12至少90%、95%、98%或99%之序列同一性。
H. LIN28 蛋白
本發明VEEV構築體可包含LIN28蛋白(例如,LIN28A或LIN28B)之編碼序列。參見,例如,美國專利9,506,039。人類LIN28B係藉由
LIN28B基因編碼。示例性人類LIN28B胺基酸序列可見於UniProt寄存編號A0A1B0GVD3且具有下列胺基酸序列:
此序列之功能類似物,即,具有以上蛋白質之相同或實質上相同生物功能(例如,保留蛋白質之轉錄因子功能之70%或更多,80%或更多,90%或更多,95%或更多,或98%或更多)之分子作為LIN28B蛋白包含於本發明。例如,該功能類似物可為以上蛋白質之同功異型物或變異體,例如,含有以上蛋白質之一部分具有或不具有另外胺基酸殘基及/或含有相對於以上蛋白質之突變。於一些實施例中,該功能類似物具有與SEQ ID NO:13至少90%、95%、98%或99%之序列同一性。
本文中所述之再程式化因子之示例性功能類似物述於(例如) Yang等人,
Asian J Andrology(2015) 17:394-402中,其揭示內容之全文係以引用的方式併入本文中。
I. RNA 表現構築體
於一些實施例中,可將再程式化因子之編碼序列併入一或多個表現盒中,該等表現盒各具有其自身啟動子(例如,26S啟動子)及其他轉錄調節元件。
於一些實施例中,可將再程式化因子之編碼序列放入多順反子性表現盒之框中使得其自共同啟動子(例如,26S或SP6啟動子)轉錄。此等編碼序列可被轉譯跳躍序列(即,自裂解肽之框內編碼序列)分隔開,使得自多順反子性盒之mRNA轉錄本之轉譯將導致分開蛋白質。自裂解肽引起在轉譯期間之核糖體跳躍。自裂解肽之實例為2A肽,其為具有18至22個胺基酸之典型長度之病毒衍生肽。2A肽包含T2A、P2A、E2A、F2A及PQR (Lo等人,
Cell Reports(2015) 13:2634-2644)。舉例而言,P2A為19個胺基酸之肽;於裂解後,來自P2A之少數胺基酸殘基留在上游基因上及脯胺酸留在第二基因之開始處。再程式化因子之編碼序列反而亦可被mRNA中之內部核糖體進入位點(IRES)分隔開。IRES亦允許分開多肽自共同RNA轉錄本之轉譯。留在經處理之多肽上之2A殘基不影響該等多肽之功能性。
舉例而言,α病毒RNA構築體可自5’至3’包含:[α病毒5’ UTR] — [α病毒RNA複製酶之基因] — [啟動子] — [再程式化因子1編碼序列] — [2A肽編碼序列] — [再程式化因子2編碼序列] — [2A肽編碼序列] — [再程式化因子3編碼序列] — [IRES或核心啟動子] — [再程式化因子4編碼序列] — [2A肽編碼序列] — [再程式化因子5] — [視情況可選的標誌物] — [α病毒3’ UTR及聚腺苷酸尾]。該聚腺苷酸尾長度可變化(例如,自10至超過200個腺苷酸),及再程式化因子之順序可改變而不影響RNA構築體之再程式化功能。多順反子性再程式化因子表現盒之啟動子可為(例如) 26S內部啟動子。於一些實施例中,該α病毒RNA構築體為VEEV RNA構築體及其複製酶之基因為VEEV RNA複製酶1、2、3及4。
於一些實施例中,該α病毒(例如,VEEV) RNA構築體可具有如
圖 1中所示之結構,自5’至3’視情況包含nsP1、nsP2、nsP3及nsP4之編碼序列,及表現再程式化因子,諸如(i) OCT4、KLF4、SOX2、BCL-xL及c-Myc,(ii) OCT4、SOX2、BCL-xL及c-MYC,或(ii) OCT4、SOX2及BCL-xL之組合之多順反子性表現盒。於多順反子性表現盒中,該表現盒可在26S啟動子之轉錄控制下,及/或再程式化因子之編碼序列可被IRES序列或自裂解2A肽之編碼序列分隔開(IRES位置之非限制性實例示於
圖 1中)。
該α病毒(例如,VEEV) RNA構築體可自DNA範本(例如,DNA質體構築體)產生。舉例而言,RNA構築體可藉由使用SP6 (或T7)活體外轉錄套組自DNA範本轉錄。
VEEV之任何株系可用於提供本發明RNA構築體之主鏈。例如,可使用VEEV之TC-83株系。此株系含有nsP2之P773S突變及因此具有對經轉導細胞之降低之細胞病變效應。可引入ns蛋白中之一或多者之其他或另外突變以改善RNA複製及表現,及/或減弱對RNA基因組之免疫反應。例如,該VEEV RNA表現構築體可包含圖6B中所示之彼等突變中之一或多者(例如,兩者、三者、四者、五者或所有六者)。另外,代替VEEV,亦可使用其他α病毒以提供再程式化自我複製RNA構築體之主鏈。可基於RNA構築體之其他α病毒之非限制性實例為東部馬腦炎病毒(EEEV)、沼澤地(Everglades)病毒、穆坎博(Mucambo)病毒、皮克斯(Pixuna)病毒及西部馬腦炎病毒(WEEV)、辛德畢斯病毒、西門利啟森林(Semliki Forest)病毒、米德爾堡(Middelburg)病毒、基孔肯雅病毒、歐尼恩病毒、羅斯河病毒、巴馬森林(Barmah Forest)病毒、蓋塔(Getah)病毒、鷺山(Sagiyama)病毒、比巴魯(Bebaru)病毒、馬雅羅(Mayaro)病毒、烏納(Una)病毒、奧拉(Aura)病毒、瓦塔羅阿(Whataroa)病毒、巴班肯(Babanki)病毒、孜拉加奇(Kyzylagach)病毒、高地J病毒、摩根堡(Fort Morgan)病毒、恩杜穆(Ndumu)病毒及博吉河(Buggy Creek)病毒。該等RNA構築體可含有超過一種α病毒之序列。
II. 使用 RNA 構築體自造血譜系之細胞產生 iPSC
本發明方法將血球有效再程式化(或稱作「去分化」)以變成誘導型多能幹細胞。
如本文中所用,術語「多能」或「多能性」係指細胞自我更新及分化成三個胚層:內胚層、中胚層或外胚層中之任一者之細胞的能力。「多能幹細胞」或「PSC」包括(例如)源自囊胚之內細胞團或藉由體細胞核移植衍生之胚胎幹細胞及源自非多能細胞之iPSC。
術語「誘導型多能幹細胞」或「iPSC」係指自非多能細胞(諸如成人體細胞)、部分分化細胞或末期分化細胞(諸如纖維母細胞、造血譜系細胞、肌細胞、神經元、上皮細胞或類似者)人工製備之一種類型之多能幹細胞,藉由引入該細胞或使該細胞與一或多種再程式化因子接觸。
PSC誘導之起始細胞群體可自需要細胞療法之患者或健康供體之血液(例如,外周血)獲得。外周血單核細胞(PMBC)可藉由習知方法分離及然後進一步分段及/或濃化以獲得細胞,例如,T淋巴細胞、B淋巴細胞、單核細胞、自然殺手細胞、嗜中性白血球、嗜酸性白血球、樹突狀細胞及各種造血祖細胞(諸如紅血球系祖細胞、淋巴祖細胞及骨髓祖細胞)之子集。
於一些實施例中,將PMBC於補充有促紅血球生成素(EPO)、幹細胞因子(SCF)及IL-3之基礎培養基(例如,StemSpan™ SFEM II培養基;StemCell Technologies)中培養一段時間(例如,3至10天,諸如6、7或8天),以獲得針對紅血球系祖細胞濃化之細胞群體。可將培養基補充(例如) 0.5至5 (例如,1、2、3或4) IU/mL EPO、50至200 (例如,75、100、125、150或175) ng/mL SCF及1至10 (例如,2、3、4、5、6、7、8或9) ng/mL IL-3。亦可使用促進紅血球系祖細胞增殖之其他因子,例如,重組人類胰島素、鐵飽和之人類轉鐵蛋白、硝酸鐵、氫化可的鬆。參見,例如,Neildez-Nguyen等人,
Nat Biotechnol.(2002) 20:467-72;及Filippone等人,
PLoS One(2010) 5(3):e9496。紅血球系祖細胞可藉由(例如)螢光或磁性活化之細胞分選使用結合至紅血球系祖細胞標誌物(諸如CD71及CD36)之試劑(例如,抗體)自細胞培養物進一步分離。
於一些實施例中,該等PBMC係在存在1 IU/ml EPO、約100 ng/mL SCF及約5 ng/mL IL-3下培養。於一些實施例中,該等PBMC係在存在1 IU/ml EPO、約100 ng/mL SCF及約10 ng/mL IL-3下培養。於一些實施例中,該等PBMC係在存在1 IU/ml EPO、約150 ng/mL SCF及約5 ng/mL IL-3下培養。於一些實施例中,該等PBMC係在存在1 IU/ml EPO、約150 ng/mL SCF及約10 ng/mL IL-3下培養。於一些實施例中,該等PBMC係在存在1 IU/ml EPO、約200 ng/mL SCF及約5 ng/mL IL-3下培養。於一些實施例中,該等PBMC係在存在1 IU/ml EPO、約200 ng/mL SCF及約10 ng/mL IL-3下培養。於一些實施例中,該等PBMC係在存在3 IU/ml EPO、約100 ng/mL SCF及約5 ng/mL IL-3下培養。於一些實施例中,該等PBMC係在存在3 IU/ml EPO、約100 ng/mL SCF及約10 ng/mL IL-3下培養。於一些實施例中,該等PBMC係在存在3 IU/ml EPO、約150 ng/mL SCF及約5 ng/mL IL-3下培養。於一些實施例中,該等PBMC係在存在3 IU/ml EPO、約150 ng/mL SCF及約10 ng/mL IL-3下培養。於一些實施例中,該等PBMC係在存在3 IU/ml EPO、約200 ng/mL SCF及約5 ng/mL IL-3下培養。於一些實施例中,該等PBMC係在存在3 IU/ml EPO、約200 ng/mL SCF及約10 ng/mL IL-3下培養。於此等實施例中,可進行該培養6、7或8天。
血球之其他子集亦可藉由透過細胞培養物分段及/或濃化獲得。血球之特定子集之標誌物係熟知,諸如針對T淋巴細胞之CD3及針對B細胞之CD19及CD20。
可藉由許多技術,包括微注射、電穿孔、基因槍粒子遞送、脂質轉染、陽離子聚合物及磷酸鈣沉澱將本發明RNA構築體引入體細胞群體中。於一些實施例中,透過電穿孔將本發明構築體RNA構築體引入體細胞(例如,造血祖細胞及淋巴細胞)中。雖然已知使用α病毒作為載體可藉由干擾素(IFN)藉由先天免疫反應抑制,但是I型IFN抑制劑(諸如B18R或B19R)可用於抑制細胞抗病毒反應,從而啟動細胞中之所需複製子活性。於一些實施例中,可在電穿孔之前將該等細胞用B18R蛋白處理以促進α病毒(例如,VEEV)遞送及隨後複製及/或抑制經轉染細胞中之細胞干擾素反應。可將經電穿孔細胞在存在B18R下培養2至3週,在此期間iPSC出現及可收穫。可藉由標誌物(諸如TRA-1-60、NANOG、SSEA3及SSEA4)檢測iPSC。於一些實施例中,諸如Opti-MEM® (Thermo Fisher)之培養基可用作電穿孔細胞懸浮緩衝液以促進細胞於電穿孔後之生存。於一些實施例中,可將RNA構築體包裝至α病毒病毒粒子及該病毒粒子係用於轉導待再程式化之細胞。
維持iPSC之方法係此項技術中熟知,及此等方法中之許多與維持胚胎幹細胞之方法相似。參見,例如,Thomson等人,
Science(1998) 282(5391):1145-7;Hovatta等人,
Human Reprod.(2003) 18(7):1404-09;Ludwig等人,
Nature Methods(2006) 3:637-46;Kennedy等人,
Blood(2007) 109:2679-87;Chen等人,
Nature Methods(2011) 8:424-9;及Wang等人,
Stem Cell Res. (2013) 11(3):1103-16。亦可將iPSC在使用之前冷凍保存。
III. iPSC 分化為靶細胞類型
iPSC為可於患有許多不同疾病之患者中遞送再生藥物之大量特定細胞類型之潛在產生的起始點。於iPSC之背景下,分化為譜系規格使用細胞特異性協定,以iPSC開始之過程。藉由本發明方法獲得之iPSC可分化成細胞療法所關注之細胞類型,包括內胚層、外胚層及中胚層譜系之細胞。於一些實施例中,可首先將iPSC遺傳工程改造(例如,以產生於患者中有缺陷之功能蛋白、以產生治療性蛋白、以包含自殺開關、或以逃避免疫檢測,從而支持異基因應用),之後分化成所關注之細胞類型。誘導iPSC至各種譜系之細胞之分化及其擴增之方法係此項技術中熟知。以下描述分化之細胞類型之非限制性實例。
A. 免疫細胞
視情況經遺傳修飾之iPSC可分化成免疫細胞,諸如淋巴系細胞(例如,T細胞、B細胞及NK細胞)、骨髓細胞(例如,粒細胞、單核細胞/巨噬細胞及組織駐留巨噬細胞,諸如小神經膠質)及樹突狀細胞(例如,骨髓樣樹突狀細胞及漿細胞樣樹突狀細胞)。於一些實施例中,該等經遺傳修飾之細胞為表現嵌合抗原受體(CAR)之T細胞或CAR T細胞。該等經遺傳修飾之免疫細胞亦可表現免疫調節轉殖基因(諸如HLA-G或HLA-E)。
例如,誘導PSC分化成樹突狀細胞之方法述於Slukvin等人,
J Imm.(2006) 176:2924-32;Su等人,
Clin Cancer Res.(2008) 14(19):6207-17;及Tseng等人,
Regen Med.(2009) 4(4):513-26中。誘導PSC至造血祖細胞、骨髓譜系細胞及T淋巴細胞之方法述於(例如) Kennedy等人,
Cell Rep.(2012) 2:1722-35中。誘導PSC至巨噬細胞之方法述於van Wilgenburg等人,
PLoS One(2013) 8(8):e71098中。
可將免疫細胞,諸如免疫抑制性免疫細胞(例如,調節性T及免疫抑制性巨噬細胞)移植至患有自體免疫性疾病之患者中,該自體免疫性疾病包括(不限於)類風濕性關節炎、多發性硬化、慢性淋巴細胞性甲狀腺炎、胰島素依賴性糖尿病、重症肌無力、慢性潰瘍性結腸炎、潰瘍性結腸炎、克羅恩氏病(Crohn’s disease)、發炎性腸病、古德帕斯氏(Goodpasture’s)症候群、全身性紅斑狼瘡、全身性血管炎、硬皮病、自體免疫性溶血性貧血及自體免疫性甲狀腺病。基於免疫細胞之療法亦可用於治療移植中之移植物排斥,包括治療與移植相關之症狀,諸如纖維化。
B. 神經細胞
視情況經遺傳修飾之iPSC可分化成神經細胞,包括(不限於)不考慮任何特定神經元亞型之神經元及神經元前體細胞(例如,多巴胺能神經元、腸神經元、中間神經元及皮層神經元);不考慮任何特定膠質亞型之膠質細胞及膠質前體細胞(例如,少突膠質細胞、星形膠質細胞、專用少突膠質細胞前體細胞及可產生星形膠質細胞及少突膠質細胞之雙能膠質前體);及小神經膠質及小神經膠質前體細胞。亦涵蓋脊髓或動眼神經元、腸神經元、基板衍生之細胞、施旺細胞及三叉神經或感覺神經元。
可將神經細胞移植至包括(不限於)患有神經退化性疾病之患者。神經退化性疾病之實例尤其為帕金森氏病(Parkinson’s disease)、阿茲海默氏病(Alzheimer’s disease)、癡呆、癲癇、路易體(Lewy body)症候群、亨廷頓氏病(Huntington’s disease)、脊髓肌肉萎縮、弗裡德賴希氏(Friedreich’s)共濟失調、肌萎縮側索硬化、巴登氏病(Batten disease)及多系統萎縮、腦白質營養不良、橫貫性脊髓炎、視神經脊髓炎、溶酶體貯積病(例如,賀勒氏(Hurler)症候群、法布裡病氏(Fabry disease)、戈謝氏病(Gaucher disease)、Sly症候群、GM1及GM2神經節苷脂貯積病、亨特氏(Hunter)症候群、尼曼-匹克氏病(Niemann-Pick disease)、沙費利波(Sanfilippo)症候群)、tau蛋白病(tauopathies)。
針對許多此等疾病,可首先透過雙重SMAD抑制指導iPSC接受原始神經細胞命運(Chambers等人,
Nat Biotechnol. (2009) 27(3):275-80)。原始神經細胞採用前面特徵,因此另外信號之不存在將提供前面/前腦皮層細胞。可阻斷尾化信號以防止可以其他方式產生具有更後部特徵之培養物之副分泌信號(例如,XAV939可阻斷WNT及SU5402可阻斷FGF信號)。背部皮層神經元可藉由阻斷SHH活化來製備,而腹部皮層神經元可透過SHH活化製備。更多尾部細胞類型,諸如血清素能神經元或脊髓運動神經元可藉由透過添加FGF及/或WNT信號尾化培養物來製備。針對一些細胞類型,可添加視黃酸(另一種尾化劑)以後驗培養物。膠質細胞類型之產生一般可跟隨原始神經細胞在含FGF2及/或EGF之培養基中之培養物擴增之前之相同模式。PNS細胞類型可跟隨相同一般原則,但是早在分化過程中具有及時WNT信號。
可透過放入所討論之受損組織中之套管將神經細胞引入患者中。可將細胞製劑放入支持介質中及負載至注射器或可精確遞送製劑之類吸量管裝置中。然後可將套管放入患者之神經系統,通常使用立體定向方法以精確靶向遞送。然後可將細胞以相容之速率驅逐至組織中。
C. 心血管細胞
視情況經遺傳修飾之iPSC可分化成心血管系統之細胞,諸如包含特定心肌細胞亞型(例如,心室或心房)之心肌細胞、心臟纖維母細胞、心臟平滑肌細胞、心臟心外膜細胞、心臟心內膜細胞、心臟內皮細胞、浦肯野(Purkinje)纖維及結節及起搏細胞。將iPSC分化成心肌細胞之許多方法存在,例如,如Kattman等人,
Cell Stem Cell(2011) 8(2):228-40;Lian等人,
PNAS(2012) 109:e1848-57;Lee等人,
Cell Stem Cell(2017) 21:179-94中所示,及如WO 2016/131137、WO 2018/098597及美國專利9,453,201中所示。此項技術中之任何適宜方法可與本文中之方法使用,以獲得PSC衍生之心肌細胞。
於一些實施例中,該等iPSC係於一或多個心臟分化培養基中培育。例如,該培養基可含有變化濃度之骨形態發生蛋白(BMP,諸如BMP4)及活化素(諸如活化素A)。可進行分化因子濃度之滴定以測定達成所需心肌細胞分化必需之最佳濃度。
於一些實施例中,該等分化之心肌細胞表現心臟肌鈣蛋白T (cTnT)及/或肌球蛋白輕鏈2v (MLC2v)中之一或多者。於一些實施例中,不成熟心肌細胞表現肌鈣蛋白T、心臟肌鈣蛋白I、α輔肌動蛋白及/或β-肌球蛋白重鏈中之一或多者。
D. 代謝系統之細胞
視情況經遺傳修飾之iPSC可分化成涉及人類代謝系統之細胞。例如,該等細胞可為胃腸系統之細胞(例如,肝細胞、膽管細胞及胰β細胞)、造血系統之細胞及中樞神經系統之細胞(例如,垂體激素釋放細胞)。舉例而言,為產生垂體激素釋放細胞,將iPSC與BMP4及SB431542 (其阻斷活化素信號傳導)培養,之後添加SHH/FGF8及FGF10;然後將細胞在FGF8或BMP (或二者)之前僅經受SHH/FGF8及FGF10持續延長之時間段,以誘導細胞變成特異性激素釋放細胞。參見,例如,Zimmer等人,
Stem Cell Reports(2016) 6:858-72。
E. 眼部系統之細胞
視情況經遺傳修飾之iPSC可分化成眼部系統之細胞。例如,該等細胞可為視網膜祖細胞、視網膜色素上皮(RPE)祖細胞、RPE細胞、神經視網膜祖細胞、感光祖細胞、感光細胞、雙極細胞、水平細胞、神經節細胞、無長突細胞、穆勒(Mueller)膠質細胞、錐形細胞或桿狀細胞。將iPSC分化成RPE細胞之方法述於(例如) WO 2017/044483中。分離RPE細胞之方法述於(例如) WO 2017/044488中。將iPSC分化成神經視網膜祖細胞之方法述於WO 2019/204817中。識別及分離視網膜祖細胞及RPE細胞之方法述於(例如) WO 2011/028524中。
IV. 醫藥組合物及用途
本文中所述之iPSC衍生細胞可以含有細胞及醫藥上可接受之載劑之醫藥組合物提供。該醫藥上可接受之載劑可為視情況不含有任何動物源組分之細胞培養基。針對儲存及運輸,可將細胞在< -70℃下(例如,在乾冰上或於液氮中)冷凍保存。在使用之前,可將細胞解凍,及於支持所關注之細胞類型之無菌細胞培養基中稀釋。
該等細胞可經全身(例如,透過靜脈內注射或輸注)或經局部(例如,透過直接注射至局部組織,例如,心臟、腦及受損組織之位點)投與至患者。投與細胞至患者之組織或器官之各種方法係此項技術中已知,該等方法包括(不限於)冠狀動脈內投與、心肌內投與、經心內膜投與或顱內投與。
向患者投與治療上有效數目之iPSC衍生細胞。如本文中所用,術語「治療上有效」係指當向患有或易感疾病、病症及/或病狀之人類個體投與時,足以治療、預防及/或延遲疾病、病症及/或病狀之該(等)症狀之發作或進展之細胞數目或醫藥組合物的量。一般技術者應瞭解,治療上有效量通常經由包含至少一個單位劑量之給藥方案投與。
除非本文中另有指定,否則結合本發明所用之科學及技術術語應具有由一般技術者通常所理解之含義。雖然與本文中所述彼等相似或等效之方法及材料亦可用於實踐或測試本發明,但是以下描述示例性方法及材料。於衝突之情況下,以本說明書(包含定義)為準。一般而言,結合本文中所述之細胞及組織培養、分子生物學、免疫學、微生物學、遺傳學、分析化學、合成有機化學、醫藥化學及蛋白質及核酸化學及雜交所用之命名法及其技術為此項技術中熟知且常用之彼等。酶促反應及純化技術係根據製造商之說明進行,如此項技術中通常所實現或如本文中所述。另外,除非上下文另有要求,否則單數術語應包含複數及複數術語應包含單數。整篇本說明書及實施例,單詞「具有(have)」及「包含(comprise)」或變型,諸如「具有(has/having)」、「包含(comprises/comprising)」應理解為暗示指定整數或整數組之納入,但是不排除任何其他整數或整數組。本文中提及之所有公開案及其他參考文獻之全文係以引用的方式併入。雖然本文中引用許多文檔,但是此引用不構成承認此等文檔中之任一者形成此項技術中之常識之部分。
為了可更佳理解本發明,闡述下列實例。此等實例係僅出於說明之目的且不應解釋為以任何方式限制本發明之範圍。
實例
實例 1 : VEEV 設計及 RNA 合成
此實例描述多順反子性VEEV RNA構築體之設計及其合成。用於下列研究中之構築體係基於Yoshioka, 2013,見上中之所述之VEEV RNA主鏈序列。該主鏈序列編碼VEEV之四種非結構性蛋白。修改亞基因組序列以表現再程式化核因子之各種組合(
圖 1)。RNA構築體OKS-iBM編碼OCT4、KLF4、SOX2、BCL-xL及c-MYC。RNA構築體OKS-iGM編碼OCT4、KLF4、SOX2、GLIS1及c-MYC。RNA構築體OKS-iG編碼OCT4、KLF4、SOX2及GLIS1。RNA構築體OSB編碼OCT4、SOX2及BCL-xL。RNA構築體OS-iB編碼OCT4、SOX2及BCL-xL。RNA構築體OS-iM編碼OCT4、SOX2及c-MYC。RNA構築體OS-iBM編碼OCT4、SOX2、BCL-xL及c-MYC。名字包含「i」之構築體含有緊接SOX2編碼序列下游之IRES序列。各種再程式化因子之編碼序列被自裂解2A肽之編碼序列分隔開或被IRES分隔開,使得所有再程式因子自相同啟動子表現(Wen等人,
Stem Cell Rep.(2016) 6:873-84;Su等人,
PLoS ONE(2013) 8:e64496)。
該等VEEV RNA構築體自其各自DNA質體範本酶促合成。為最佳化RNA合成,使用AG封蓋類似技術(CleanCap, Trilink)產生5’封端之RNA。
重組VEEV構築體之主鏈序列示於圖6B中(SEQ ID NO:15),其中表現盒之插入位點藉由序列中之星號指示。
實例 2 :紅血球系祖細胞至 iPSC 之再程式化
此實例描述紅血球系祖細胞至iPSC之再程式化之示例性協定。為獲得針對紅血球系祖細胞(EP)濃化之細胞群體,將來自人類供體之PBMC解凍及然後於補充有約3 IU/mL EPO、約100 ng/mL SCF及約5 ng/mL IL-3之培養基(「EP培養基」)中培養5至10天(例如,6天)。此等生長因子支持EP增殖。參見,例如,Wang等人,
Clin Hemorheol Microcirc. (2007) 37(4):291-9。或者,可將PBMC如Wen等人(
Stem Cell Reports(2016) 6:873-84)中所述,例如,於補充有100 ng/ml幹細胞因子(Peprotech;300-07)、10 ng/ml介白素-3 (Peprotech;AF-200-03)、2 U/ml促紅血球生成素(Peprotech;100-64)、20 ng/ml胰島素生長因子-1 (Peprotech;100-11)、1 mM地塞米松(dexamethasone) (Sigma;D4902)及0.2 mM 1-硫代甘油(Sigma;M6145)之包含Stemline® II 造血幹細胞擴增培養基(Sigma;S0192)之培養基中培養。
更具體而言,將經解凍之PBMC接種於EP培養基中以於組織培養物處理板中達成2至3 x 10
6個細胞/mL之細胞密度。於接種後,更換約1/4至3/4之培養基過夜(例如,16至24小時)。在第2天,將細胞轉移至新容器中(超低黏附,經非組織培養物處理)及每日進行25至75%培養基更換。在第5天,藉由添加另外EP培養基將細胞稀釋2倍。在第6天(或第7天),更換一半之培養基,及藉由流動式細胞測量術評價EP細胞之樣品針對CD71及CD36之雙陽性。
然後將CD71
+CD36
+EP細胞用干擾素抑制劑(例如,重組B18R蛋白)培育20分鐘。將細胞離心,用DPBS洗滌,及然後以約2 x 10
7個細胞/mL再懸浮於Opti-MEM™ (Thermo Fisher Scientific)中。針對每個120 μL電穿孔反應,將4 μg VEEV再程式化RNA轉移至冷凍過之1.5 mL微管中。然後將細胞電穿孔及平板接種於補充B18R之EP培養基中及分批進料2天。將板用諸如玻連蛋白(vitronectin)或層連蛋白(laminin)之受質塗覆。在轉染後第3至6天,將細胞用補充B18R之Essential 7或ReproTeSR™培養基分批進料。在轉染後第7天開始,培養物經歷每日完全培養基更換。iPSC之群落在轉染後約第10天出現,及準備在第15天與第20天之間挑選。將經挑選之群落在不存在B18R下擴增及然後冷凍保存(本文中稱作VEE-EP-iPSC)。
如
圖 2A 至 C中所示,利用三種VEEV RNA構築體產生之VEE-EP-iPSC表現與未分化之多能細胞相關聯之核(NANOG)及表面標誌物(TRA-1-60、SSEA-3及SSEA-4)。此等細胞亦具有正常核型。
該等VEE-EP-iPSC藉由下一代定序描繪以評估超過500個癌症相關基因之遺傳變異體之獲取。當將VEE-EP-iPSC系中之超過500個基因之遺傳序列與供體PBMC之起始群體相比時,未觀察到序列差異。此等資料證實在再程式化期間未獲得遺傳變異體。
所有VEE-EP-iPSC細胞系證實分化成代表外胚層之TH
+多巴胺能神經元之能力(
圖 3)。為使VEE-EP-iPSC朝向多巴胺能神經元直接分化,吾人首先藉由自第0天至第7天阻斷TGF-β及BMP信號傳導來誘導iPSC朝向神經外皮層譜系。吾人同時自第0天至第7天經由重組C25II SHH蛋白將細胞模式化至底板命運,及自第0天至第12天使用WNT促效劑小分子CHIR-99021微調細胞之中腦同一性。於此等步驟後,吾人自第10天至第16天使用神經營養因子及自第12天至第16天使用小分子γ-分泌酶抑制劑DAPT使祖細胞成熟為神經元命運。將第16天分化細胞於活體外成熟5天及藉由流動式細胞測量術定量TH
+FOXA2
+多巴胺神經元。
所有VEE-EP-iPSC細胞系亦能分化成代表中胚層分化之心肌肌鈣蛋白(cTNT)陽性心肌細胞(
圖 4)。利用WNT信號傳導之階段特異性調節透過使用WNT促效劑CHIR-99021及WNT拮抗劑內-IWR1使VEE-EP-iPSC系朝向心臟譜系分化。藉由流動式細胞測量術定量心肌細胞之心肌肌鈣蛋白(cTNT)染色。
為測定含有不同轉錄因子組合之VEEV RNA構築體之再程式化效率,將紅血球系祖細胞自PBMC擴增及用再程式化RNA構築體電穿孔。在電穿孔後17天(針對OKS-iBM及附加型構築體)或25天(針對OKS-iGM及OKS-iG構築體)定量TRA-1-60陽性群落。以表現PSC標誌物TRA-1-60之細胞基群落之數目測定再程式化效率(
圖 5A)。當BCL-xL編碼序列包含於VEEV構築體中時,VEEV RNA介導之EP再程式化之效率顯著增加(
圖 5B)。含有BCL-xL編碼序列之VEEV OKS-iBM構築體證實較含有傳統再程式化因子OCT4、SOX2、KLF4、L-MYC、LIN28及p53顯性負性之附加體基對照高四倍的EP再程式化效率(
圖 5B – iBM/Epi5)。OS-iBM構築體亦係活性及形成iPSC群落(數據未顯示)。
當Su等人(2013,見上)包含BCL-xL作為第五再程式化因子(附加型OS+MK+B組合)時,其觀察到與OS+MK (OCT4、SOX2、c-MYC及KLF4)組合相比再程式效率之約8倍增加。然而,當Yoshioka及Dowdy (2017,見上)評價再程式化因子組合時,GLIS1 (VEE-OKS-iGM)之納入與四因子組合(VEE-OKS-iM)相比提高再程式化效率約20倍。因此,意外地將GLIS1用BCL-xL (VEE-OKS-iBM)替換再增加紅血球系祖細胞之再程式化效率8倍(
圖 5B – iBM/iGM)。再程式化效率之此出人意料且急劇增加將低效率紅血球系再程式化轉變成穩健方法。更穩健效率允許跨不同血液樣品之一致再程式化及對EP再程式化用於臨床用途之可行性係關鍵的。
實例 3 : T 細胞至 iPSC 之再程式化
此實例描述將T淋巴細胞再程式化至iPSC之協定。藉由來自兩個獨立供體(AllCells)之外周血之陰性免疫選擇及免疫表現型獲得經純化之CD3
+T細胞(泛T細胞)。將來自兩個供體之泛T細胞解凍及維持於補充有CTS™ GlutaMAX™及100 IU/mL之T細胞完全培養基中。在電穿孔之前,將泛T細胞用0.2 μg/mL重組B18R蛋白處理30分鐘,用冰冷磷酸鹽緩衝鹽水洗滌及再懸浮。
將細胞電穿孔,平板接種及在超低黏附板(Corning)上於以上補充有0.2 μg/mL B18R、CTS™ GlutaMAX™及100 IU/mL IL-2之T細胞培養基中培育24小時。接下來,將細胞再平板接種至塗覆有LN521之新鮮補充0.2 μg/mL B18R之組織培養板上。在電穿孔後第3天與第6天之間,將再程式化培養物用每日補充0.2 μg/mL B18R之StemFit Basic03培養基(Ajinomoto)分批進料。在轉染後第7天開始,培養物經歷利用補充B18R之StemFit Basic03培養基之每日完全培養基更換。iPSC之群落在轉染後約第10天出現,及準備在第15天與第20天之間挑選。將經挑選之群落在不存在B18R下用StemFit Basic03培養基擴增及然後冷凍保存(本文中稱作VEE-T-iPSC)。
總之,吾人已證實除了紅血球系祖細胞外,CD3
+T-細胞可藉由用VEE-OKS-iBM RNA電穿孔再程式化。將兩個不同供體T-細胞批次再程式化以建立iPSC系,其中兩個供體之間之再程式化效率平均值為0.005%。此為足夠細胞系衍生率,因為其將導致50個群落/百萬經轉染細胞。
序列表
下表中提供本發明之示例性序列(SEQ: SEQ ID NO)。
表1
SEQ | 描述 |
1 | 人類BCL-xL胺基酸序列 |
2 | 本文中表現之示例性人類BCL-xL胺基酸序列 |
3 | 人類OCT4胺基酸序列 |
4 | 本文中表現之示例性人類OCT4胺基酸序列 |
5 | 人類KLF4胺基酸序列 |
6 | 本文中表現之示例性人類 KLF4胺基酸序列 |
7 | 人類SOX2胺基酸序列 |
8 | 本文中表現之示例性人類SOX2胺基酸序列 |
9 | 人類c-MYC胺基酸序列 |
10 | 本文中表現之示例性人類c-MYC胺基酸序列 |
11 | 人類GLIS1胺基酸序列 |
12 | 人類NANOG胺基酸序列 |
13 | 人類LIN28B胺基酸序列 |
14 | 野生型VEEV基因組之核苷酸序列 |
15 | 重組VEEV RNA表現載體之核苷酸序列 |
圖 1為說明7種示例性VEEV RNA再程式化構築體(OKS-iBM、OKS-iGM、OKS-iG、OSB、OS-iB、OS-iM及OS-iBM)之示意圖。nsP1-4:非結構蛋白之編碼序列。OCT4:八聚體結合轉錄因子4之編碼序列。KLF4:類Krüppel因子4之編碼序列。SOX2:SRY-盒轉錄因子2之編碼序列。BCL-xL:特大B-細胞淋巴瘤。GLIS1:Glis家族鋅指1之編碼序列。IRES:內部核糖體進入位點。具有星號之灰色條:2A肽之編碼序列。AAA:聚腺苷酸序列。
圖 2A 至 D為顯示多能性相關標誌物於四種VEE-EP-iPSC系(各自為VEE-EP-iPSC-1、-2、-3及-4)中之表現之流動式細胞測量術圖。VEE-EP-iPSC-1、-2、-3及-4藉由紅血球系組(EP)細胞各自利用OKS-iBM、OKS-iGM、OKS-iG及附加體控制之電穿孔產生。EBNA OriP附加體控制含有再程式化因子OCT4、SOX2、KLF4、L-MYC、LIN28及p53顯性負性(Epi5™附加型iPSC再程式化套組,Thermo Fisher;Okita等人,
Nat Meth.(2011) 8:409-12)。流動式細胞測量術係在經培養之細胞上在通道8上進行。
圖 3為顯示首先藉由誘導神經外皮層譜系及然後使祖細胞成熟為神經元命運將四種VEE-EP-iPSC系分化16天之條形圖。TH
+FOXA2
+多巴胺神經元係藉由流動式細胞測量術定量。TH:酪胺酸羥化酶。FOXA2:叉頭盒蛋白A2。
圖 4為顯示四種VEE-EP-iPSC系朝向具有WNT信號之階段特異性調節之心臟譜系分化7天之條形圖。心肌細胞之心肌肌鈣蛋白(cTNT)係藉由流動式細胞測量術定量。
圖 5A及
5B顯示EP之VEE RNA再程式化之效率。將EP用
圖 1中所說明之再程式化構築體電穿孔。使用如上針對
圖 2所述之附加體控制。
圖 5A為顯示TRA-1-60染色之全孔成像之照片組。
圖 5B為定量TRA-1-60
+群落之圖。
圖 6A顯示野生型VEEV RNA基因組序列之核苷酸序列(SEQ ID NO:14) (不同之處在於序列中之T針對RNA為U)。
圖 6B顯示重組VEEV RNA表現載體之核苷酸序列(SEQ ID NO:15) (不同之處在於序列中之T針對RNA為U)。此序列相對於野生型序列含有六個突變(C352G、A1564G、C1567A、T1570C、C1647A及C3917T)。選殖位點由星號指示。
<![CDATA[<110> 美商藍岩醫療公司(BLUEROCK THERAPEUTICS LP)]]> <![CDATA[<120> 獲得誘導型多能幹細胞之方法]]> <![CDATA[<130> 025450.TW014]]> <![CDATA[<140> TW 111111472]]> <![CDATA[<141> 2022-03-25]]> <![CDATA[<150> 63/166,071]]> <![CDATA[<151> 2021-03-25]]> <![CDATA[<160> 15 ]]> <![CDATA[<170> PatentIn version 3.5]]> <![CDATA[<210> 1]]> <![CDATA[<211> 233]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 智人]]> <![CDATA[<400> 1]]> Met Ser Gln Ser Asn Arg Glu Leu Val Val Asp Phe Leu Ser Tyr Lys 1 5 10 15 Leu Ser Gln Lys Gly Tyr Ser Trp Ser Gln Phe Ser Asp Val Glu Glu 20 25 30 Asn Arg Thr Glu Ala Pro Glu Gly Thr Glu Ser Glu Met Glu Thr Pro 35 40 45 Ser Ala Ile Asn Gly Asn Pro Ser Trp His Leu Ala Asp Ser Pro Ala 50 55 60 Val Asn Gly Ala Thr Gly His Ser Ser Ser Leu Asp Ala Arg Glu Val 65 70 75 80 Ile Pro Met Ala Ala Val Lys Gln Ala Leu Arg Glu Ala Gly Asp Glu 85 90 95 Phe Glu Leu Arg Tyr Arg Arg Ala Phe Ser Asp Leu Thr Ser Gln Leu 100 105 110 His Ile Thr Pro Gly Thr Ala Tyr Gln Ser Phe Glu Gln Val Val Asn 115 120 125 Glu Leu Phe Arg Asp Gly Val Asn Trp Gly Arg Ile Val Ala Phe Phe 130 135 140 Ser Phe Gly Gly Ala Leu Cys Val Glu Ser Val Asp Lys Glu Met Gln 145 150 155 160 Val Leu Val Ser Arg Ile Ala Ala Trp Met Ala Thr Tyr Leu Asn Asp 165 170 175 His Leu Glu Pro Trp Ile Gln Glu Asn Gly Gly Trp Asp Thr Phe Val 180 185 190 Glu Leu Tyr Gly Asn Asn Ala Ala Ala Glu Ser Arg Lys Gly Gln Glu 195 200 205 Arg Phe Asn Arg Trp Phe Leu Thr Gly Met Thr Val Ala Gly Val Val 210 215 220 Leu Leu Gly Ser Leu Phe Ser Arg Lys 225 230 <![CDATA[<210> 2]]> <![CDATA[<211> 254]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 人造序列]]> <![CDATA[<220>]]> <![CDATA[<223> 人造序列之描述:合成多肽]]> <![CDATA[<400> 2]]> Met Ser Gln Ser Asn Arg Glu Leu Val Val Asp Phe Leu Ser Tyr Lys 1 5 10 15 Leu Ser Gln Lys Gly Tyr Ser Trp Ser Gln Phe Ser Asp Val Glu Glu 20 25 30 Asn Arg Thr Glu Ala Pro Glu Gly Thr Glu Ser Glu Met Glu Thr Pro 35 40 45 Ser Ala Ile Asn Gly Asn Pro Ser Trp His Leu Ala Asp Ser Pro Ala 50 55 60 Val Asn Gly Ala Thr Gly His Ser Ser Ser Leu Asp Ala Arg Glu Val 65 70 75 80 Ile Pro Met Ala Ala Val Lys Gln Ala Leu Arg Glu Ala Gly Asp Glu 85 90 95 Phe Glu Leu Arg Tyr Arg Arg Ala Phe Ser Asp Leu Thr Ser Gln Leu 100 105 110 His Ile Thr Pro Gly Thr Ala Tyr Gln Ser Phe Glu Gln Val Val Asn 115 120 125 Glu Leu Phe Arg Asp Gly Val Asn Trp Gly Arg Ile Val Ala Phe Phe 130 135 140 Ser Phe Gly Gly Ala Leu Cys Val Glu Ser Val Asp Lys Glu Met Gln 145 150 155 160 Val Leu Val Ser Arg Ile Ala Ala Trp Met Ala Thr Tyr Leu Asn Asp 165 170 175 His Leu Glu Pro Trp Ile Gln Glu Asn Gly Gly Trp Asp Thr Phe Val 180 185 190 Glu Leu Tyr Gly Asn Asn Ala Ala Ala Glu Ser Arg Lys Gly Gln Glu 195 200 205 Arg Phe Asn Arg Trp Phe Leu Thr Gly Met Thr Val Ala Gly Val Val 210 215 220 Leu Leu Gly Ser Leu Phe Ser Arg Lys Gly Ser Gly Ala Thr Asn Phe 225 230 235 240 Ser Leu Leu Lys Gln Ala Gly Asp Val Glu Glu Asn Pro Gly 245 250 <![CDATA[<210> 3]]> <![CDATA[<211> 360]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 智人]]> <![CDATA[<400> 3]]> Met Ala Gly His Leu Ala Ser Asp Phe Ala Phe Ser Pro Pro Pro Gly 1 5 10 15 Gly Gly Gly Asp Gly Pro Gly Gly Pro Glu Pro Gly Trp Val Asp Pro 20 25 30 Arg Thr Trp Leu Ser Phe Gln Gly Pro Pro Gly Gly Pro Gly Ile Gly 35 40 45 Pro Gly Val Gly Pro Gly Ser Glu Val Trp Gly Ile Pro Pro Cys Pro 50 55 60 Pro Pro Tyr Glu Phe Cys Gly Gly Met Ala Tyr Cys Gly Pro Gln Val 65 70 75 80 Gly Val Gly Leu Val Pro Gln Gly Gly Leu Glu Thr Ser Gln Pro Glu 85 90 95 Gly Glu Ala Gly Val Gly Val Glu Ser Asn Ser Asp Gly Ala Ser Pro 100 105 110 Glu Pro Cys Thr Val Thr Pro Gly Ala Val Lys Leu Glu Lys Glu Lys 115 120 125 Leu Glu Gln Asn Pro Glu Glu Ser Gln Asp Ile Lys Ala Leu Gln Lys 130 135 140 Glu Leu Glu Gln Phe Ala Lys Leu Leu Lys Gln Lys Arg Ile Thr Leu 145 150 155 160 Gly Tyr Thr Gln Ala Asp Val Gly Leu Thr Leu Gly Val Leu Phe Gly 165 170 175 Lys Val Phe Ser Gln Thr Thr Ile Cys Arg Phe Glu Ala Leu Gln Leu 180 185 190 Ser Phe Lys Asn Met Cys Lys Leu Arg Pro Leu Leu Gln Lys Trp Val 195 200 205 Glu Glu Ala Asp Asn Asn Glu Asn Leu Gln Glu Ile Cys Lys Ala Glu 210 215 220 Thr Leu Val Gln Ala Arg Lys Arg Lys Arg Thr Ser Ile Glu Asn Arg 225 230 235 240 Val Arg Gly Asn Leu Glu Asn Leu Phe Leu Gln Cys Pro Lys Pro Thr 245 250 255 Leu Gln Gln Ile Ser His Ile Ala Gln Gln Leu Gly Leu Glu Lys Asp 260 265 270 Val Val Arg Val Trp Phe Cys Asn Arg Arg Gln Lys Gly Lys Arg Ser 275 280 285 Ser Ser Asp Tyr Ala Gln Arg Glu Asp Phe Glu Ala Ala Gly Ser Pro 290 295 300 Phe Ser Gly Gly Pro Val Ser Phe Pro Leu Ala Pro Gly Pro His Phe 305 310 315 320 Gly Thr Pro Gly Tyr Gly Ser Pro His Phe Thr Ala Leu Tyr Ser Ser 325 330 335 Val Pro Phe Pro Glu Gly Glu Ala Phe Pro Pro Val Ser Val Thr Thr 340 345 350 Leu Gly Ser Pro Met His Ser Asn 355 360 <![CDATA[<210> 4]]> <![CDATA[<211> 380]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 人造序列]]> <![CDATA[<220>]]> <![CDATA[<223> 人造序列之描述:合成多肽]]> <![CDATA[<400> 4]]> Met Ala Gly His Leu Ala Ser Asp Phe Ala Phe Ser Pro Pro Pro Gly 1 5 10 15 Gly Gly Gly Asp Gly Pro Gly Gly Pro Glu Pro Gly Trp Val Asp Pro 20 25 30 Arg Thr Trp Leu Ser Phe Gln Gly Pro Pro Gly Gly Pro Gly Ile Gly 35 40 45 Pro Gly Val Gly Pro Gly Ser Glu Val Trp Gly Ile Pro Pro Cys Pro 50 55 60 Pro Pro Tyr Glu Phe Cys Gly Gly Met Ala Tyr Cys Gly Pro Gln Val 65 70 75 80 Gly Val Gly Leu Val Pro Gln Gly Gly Leu Glu Thr Ser Gln Pro Glu 85 90 95 Gly Glu Ala Gly Val Gly Val Glu Ser Asn Ser Asp Gly Ala Ser Pro 100 105 110 Glu Pro Cys Thr Val Thr Pro Gly Ala Val Lys Leu Glu Lys Glu Lys 115 120 125 Leu Glu Gln Asn Pro Glu Glu Ser Gln Asp Ile Lys Ala Leu Gln Lys 130 135 140 Glu Leu Glu Gln Phe Ala Lys Leu Leu Lys Gln Lys Arg Ile Thr Leu 145 150 155 160 Gly Tyr Thr Gln Ala Asp Val Gly Leu Thr Leu Gly Val Leu Phe Gly 165 170 175 Lys Val Phe Ser Gln Thr Thr Ile Cys Arg Phe Glu Ala Leu Gln Leu 180 185 190 Ser Phe Lys Asn Met Cys Lys Leu Arg Pro Leu Leu Gln Lys Trp Val 195 200 205 Glu Glu Ala Asp Asn Asn Glu Asn Leu Gln Glu Ile Cys Lys Ala Glu 210 215 220 Thr Leu Val Gln Ala Arg Lys Arg Lys Arg Thr Ser Ile Glu Asn Arg 225 230 235 240 Val Arg Gly Asn Leu Glu Asn Leu Phe Leu Gln Cys Pro Lys Pro Thr 245 250 255 Leu Gln Gln Ile Ser His Ile Ala Gln Gln Leu Gly Leu Glu Lys Asp 260 265 270 Val Val Arg Val Trp Phe Cys Asn Arg Arg Gln Lys Gly Lys Arg Ser 275 280 285 Ser Ser Asp Tyr Ala Gln Arg Glu Asp Phe Glu Ala Ala Gly Ser Pro 290 295 300 Phe Ser Gly Gly Pro Val Ser Phe Pro Leu Ala Pro Gly Pro His Phe 305 310 315 320 Gly Thr Pro Gly Tyr Gly Ser Pro His Phe Thr Ala Leu Tyr Ser Ser 325 330 335 Val Pro Phe Pro Glu Gly Glu Ala Phe Pro Pro Val Ser Val Thr Thr 340 345 350 Leu Gly Ser Pro Met His Ser Asn Gly Ser Gly Glu Gly Arg Gly Ser 355 360 365 Leu Leu Thr Cys Gly Asp Val Glu Glu Asn Pro Gly 370 375 380 <![CDATA[<210> 5]]> <![CDATA[<211> 413]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 智人]]> <![CDATA[<400> 5]]> Met Arg Gln Pro Pro Gly Glu Ser Asp Met Ala Val Ser Asp Ala Leu 1 5 10 15 Leu Pro Ser Phe Ser Thr Phe Ala Ser Gly Pro Ala Gly Arg Glu Lys 20 25 30 Thr Leu Arg Gln Ala Gly Ala Pro Asn Asn Arg Trp Arg Glu Glu Leu 35 40 45 Ser His Met Lys Arg Leu Pro Pro Val Leu Pro Gly Arg Pro Tyr Asp 50 55 60 Leu Ala Ala Ala Thr Val Ala Thr Asp Leu Glu Ser Gly Gly Ala Gly 65 70 75 80 Ala Ala Cys Gly Gly Ser Asn Leu Ala Pro Leu Pro Arg Arg Glu Thr 85 90 95 Glu Glu Phe Asn Asp Leu Leu Asp Leu Asp Phe Ile Leu Ser Asn Ser 100 105 110 Leu Thr His Pro Pro Glu Ser Val Ala Ala Thr Val Ser Ser Ser Ala 115 120 125 Ser Ala Ser Ser Ser Ser Ser Pro Ser Ser Ser Gly Pro Ala Ser Ala 130 135 140 Pro Ser Thr Cys Ser Phe Thr Tyr Pro Ile Arg Ala Gly Asn Asp Pro 145 150 155 160 Gly Val Ala Pro Gly Gly Thr Gly Gly Gly Leu Leu Tyr Gly Arg Glu 165 170 175 Ser Ala Pro Pro Pro Thr Ala Pro Phe Asn Leu Ala Asp Ile Asn Asp 180 185 190 Val Ser Pro Ser Gly Gly Phe Val Ala Glu Leu Leu Arg Pro Glu Leu 195 200 205 Asp Pro Val Tyr Ile Pro Pro Gln Gln Pro Gln Pro Pro Gly Gly Gly 210 215 220 Leu Met Gly Lys Phe Val Leu Lys Ala Ser Leu Ser Ala Pro Gly Ser 225 230 235 240 Glu Tyr Gly Ser Pro Ser Val Ile Ser Val Ser Lys Gly Ser Pro Asp 245 250 255 Gly Ser His Pro Val Val Val Ala Pro Tyr Asn Gly Gly Pro Pro Arg 260 265 270 Thr Cys Pro Lys Ile Lys Gln Glu Ala Val Ser Ser Cys Thr His Leu 275 280 285 Gly Ala Gly Pro Pro Leu Ser Asn Gly His Arg Pro Ala Ala His Asp 290 295 300 Phe Pro Leu Gly Arg Gln Leu Pro Ser Arg Thr Thr Pro Thr Leu Gly 305 310 315 320 Leu Glu Glu Val Leu Ser Ser Arg Asp Cys His Pro Ala Leu Pro Leu 325 330 335 Pro Pro Gly Phe His Pro His Pro Gly Pro Asn Tyr Pro Ser Glu Leu 340 345 350 Met Pro Pro Gly Ser Cys Met Pro Glu Glu Pro Lys Pro Lys Arg Gly 355 360 365 Arg Arg Ser Trp Pro Arg Lys Arg Thr Ala Thr His Thr Cys Asp Tyr 370 375 380 Ala Gly Cys Gly Lys Thr Tyr Thr Lys Ser Ser His Leu Lys Ala His 385 390 395 400 Arg Ser Asp His Leu Ala Leu His Met Lys Arg His Phe 405 410 <![CDATA[<210> 6]]> <![CDATA[<211> 493]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 人造序列]]> <![CDATA[<220>]]> <![CDATA[<223> 人造序列之描述:合成多肽]]> <![CDATA[<400> 6]]> Pro Met Ala Val Ser Asp Ala Leu Leu Pro Ser Phe Ser Thr Phe Ala 1 5 10 15 Ser Gly Pro Ala Gly Arg Glu Lys Thr Leu Arg Gln Ala Gly Ala Pro 20 25 30 Asn Asn Arg Trp Arg Glu Glu Leu Ser His Met Lys Arg Leu Pro Pro 35 40 45 Val Leu Pro Gly Arg Pro Tyr Asp Leu Ala Ala Ala Thr Val Ala Thr 50 55 60 Asp Leu Glu Ser Gly Gly Ala Gly Ala Ala Cys Gly Gly Ser Asn Leu 65 70 75 80 Ala Pro Leu Pro Arg Arg Glu Thr Glu Glu Phe Asn Asp Leu Leu Asp 85 90 95 Leu Asp Phe Ile Leu Ser Asn Ser Leu Thr His Pro Pro Glu Ser Val 100 105 110 Ala Ala Thr Val Ser Ser Ser Ala Ser Ala Ser Ser Ser Ser Ser Pro 115 120 125 Ser Ser Ser Gly Pro Ala Ser Ala Pro Ser Thr Cys Ser Phe Thr Tyr 130 135 140 Pro Ile Arg Ala Gly Asn Asp Pro Gly Val Ala Pro Gly Gly Thr Gly 145 150 155 160 Gly Gly Leu Leu Tyr Gly Arg Glu Ser Ala Pro Pro Pro Thr Ala Pro 165 170 175 Phe Asn Leu Ala Asp Ile Asn Asp Val Ser Pro Ser Gly Gly Phe Val 180 185 190 Ala Glu Leu Leu Arg Pro Glu Leu Asp Pro Val Tyr Ile Pro Pro Gln 195 200 205 Gln Pro Gln Pro Pro Gly Gly Gly Leu Met Gly Lys Phe Val Leu Lys 210 215 220 Ala Ser Leu Ser Ala Pro Gly Ser Glu Tyr Gly Ser Pro Ser Val Ile 225 230 235 240 Ser Val Ser Lys Gly Ser Pro Asp Gly Ser His Pro Val Val Val Ala 245 250 255 Pro Tyr Asn Gly Gly Pro Pro Arg Thr Cys Pro Lys Ile Lys Gln Glu 260 265 270 Ala Val Ser Ser Cys Thr His Leu Gly Ala Gly Pro Pro Leu Ser Asn 275 280 285 Gly His Arg Pro Ala Ala His Asp Phe Pro Leu Gly Arg Gln Leu Pro 290 295 300 Ser Arg Thr Thr Pro Thr Leu Gly Leu Glu Glu Val Leu Ser Ser Arg 305 310 315 320 Asp Cys His Pro Ala Leu Pro Leu Pro Pro Gly Phe His Pro His Pro 325 330 335 Gly Pro Asn Tyr Pro Ser Phe Leu Pro Asp Gln Met Gln Pro Gln Val 340 345 350 Pro Pro Leu His Tyr Gln Glu Leu Met Pro Pro Gly Ser Cys Met Pro 355 360 365 Glu Glu Pro Lys Pro Lys Arg Gly Arg Arg Ser Trp Pro Arg Lys Arg 370 375 380 Thr Ala Thr His Thr Cys Asp Tyr Ala Gly Cys Gly Lys Thr Tyr Thr 385 390 395 400 Lys Ser Ser His Leu Lys Ala His Leu Arg Thr His Thr Gly Glu Lys 405 410 415 Pro Tyr His Cys Asp Trp Asp Gly Cys Gly Trp Lys Phe Ala Arg Ser 420 425 430 Asp Glu Leu Thr Arg His Tyr Arg Lys His Thr Gly His Arg Pro Phe 435 440 445 Gln Cys Gln Lys Cys Asp Arg Ala Phe Ser Arg Ser Asp His Leu Ala 450 455 460 Leu His Met Lys Arg His Phe Gly Ser Gly Gln Cys Thr Asn Tyr Ala 465 470 475 480 Leu Leu Lys Leu Ala Gly Asp Val Glu Ser Asn Pro Gly 485 490 <![CDATA[<210> 7]]> <![CDATA[<211> 317]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 智人]]> <![CDATA[<400> 7]]> Met Tyr Asn Met Met Glu Thr Glu Leu Lys Pro Pro Gly Pro Gln Gln 1 5 10 15 Thr Ser Gly Gly Gly Gly Gly Asn Ser Thr Ala Ala Ala Ala Gly Gly 20 25 30 Asn Gln Lys Asn Ser Pro Asp Arg Val Lys Arg Pro Met Asn Ala Phe 35 40 45 Met Val Trp Ser Arg Gly Gln Arg Arg Lys Met Ala Gln Glu Asn Pro 50 55 60 Lys Met His Asn Ser Glu Ile Ser Lys Arg Leu Gly Ala Glu Trp Lys 65 70 75 80 Leu Leu Ser Glu Thr Glu Lys Arg Pro Phe Ile Asp Glu Ala Lys Arg 85 90 95 Leu Arg Ala Leu His Met Lys Glu His Pro Asp Tyr Lys Tyr Arg Pro 100 105 110 Arg Arg Lys Thr Lys Thr Leu Met Lys Lys Asp Lys Tyr Thr Leu Pro 115 120 125 Gly Gly Leu Leu Ala Pro Gly Gly Asn Ser Met Ala Ser Gly Val Gly 130 135 140 Val Gly Ala Gly Leu Gly Ala Gly Val Asn Gln Arg Met Asp Ser Tyr 145 150 155 160 Ala His Met Asn Gly Trp Ser Asn Gly Ser Tyr Ser Met Met Gln Asp 165 170 175 Gln Leu Gly Tyr Pro Gln His Pro Gly Leu Asn Ala His Gly Ala Ala 180 185 190 Gln Met Gln Pro Met His Arg Tyr Asp Val Ser Ala Leu Gln Tyr Asn 195 200 205 Ser Met Thr Ser Ser Gln Thr Tyr Met Asn Gly Ser Pro Thr Tyr Ser 210 215 220 Met Ser Tyr Ser Gln Gln Gly Thr Pro Gly Met Ala Leu Gly Ser Met 225 230 235 240 Gly Ser Val Val Lys Ser Glu Ala Ser Ser Ser Pro Pro Val Val Thr 245 250 255 Ser Ser Ser His Ser Arg Ala Pro Cys Gln Ala Gly Asp Leu Arg Asp 260 265 270 Met Ile Ser Met Tyr Leu Pro Gly Ala Glu Val Pro Glu Pro Ala Ala 275 280 285 Pro Ser Arg Leu His Met Ser Gln His Tyr Gln Ser Gly Pro Val Pro 290 295 300 Gly Thr Ala Ile Asn Gly Thr Leu Pro Leu Ser His Met 305 310 315 <![CDATA[<210> 8]]> <![CDATA[<211> 318]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 人造序列]]> <![CDATA[<220>]]> <![CDATA[<223> 人造序列之描述:合成多肽]]> <![CDATA[<400> 8]]> Pro Met Tyr Asn Met Met Glu Thr Glu Leu Lys Pro Pro Gly Pro Gln 1 5 10 15 Gln Thr Ser Gly Gly Gly Gly Gly Asn Ser Thr Ala Ala Ala Ala Gly 20 25 30 Gly Asn Gln Lys Asn Ser Pro Asp Arg Val Lys Arg Pro Met Asn Ala 35 40 45 Phe Met Val Trp Ser Arg Gly Gln Arg Arg Lys Met Ala Gln Glu Asn 50 55 60 Pro Lys Met His Asn Ser Glu Ile Ser Lys Arg Leu Gly Ala Glu Trp 65 70 75 80 Lys Leu Leu Ser Glu Thr Glu Lys Arg Pro Phe Ile Asp Glu Ala Lys 85 90 95 Arg Leu Arg Ala Leu His Met Lys Glu His Pro Asp Tyr Lys Tyr Arg 100 105 110 Pro Arg Arg Lys Thr Lys Thr Leu Met Lys Lys Asp Lys Tyr Thr Leu 115 120 125 Pro Gly Gly Leu Leu Ala Pro Gly Gly Asn Ser Met Ala Ser Gly Val 130 135 140 Gly Val Gly Ala Gly Leu Gly Ala Gly Val Asn Gln Arg Met Asp Ser 145 150 155 160 Tyr Ala His Met Asn Gly Trp Ser Asn Gly Ser Tyr Ser Met Met Gln 165 170 175 Asp Gln Leu Gly Tyr Pro Gln His Pro Gly Leu Asn Ala His Gly Ala 180 185 190 Ala Gln Met Gln Pro Met His Arg Tyr Asp Val Ser Ala Leu Gln Tyr 195 200 205 Asn Ser Met Thr Ser Ser Gln Thr Tyr Met Asn Gly Ser Pro Thr Tyr 210 215 220 Ser Met Ser Tyr Ser Gln Gln Gly Thr Pro Gly Met Ala Leu Gly Ser 225 230 235 240 Met Gly Ser Val Val Lys Ser Glu Ala Ser Ser Ser Pro Pro Val Val 245 250 255 Thr Ser Ser Ser His Ser Arg Ala Pro Cys Gln Ala Gly Asp Leu Arg 260 265 270 Asp Met Ile Ser Met Tyr Leu Pro Gly Ala Glu Val Pro Glu Pro Ala 275 280 285 Ala Pro Ser Arg Leu His Met Ser Gln His Tyr Gln Ser Gly Pro Val 290 295 300 Pro Gly Thr Ala Ile Asn Gly Thr Leu Pro Leu Ser His Met 305 310 315 <![CDATA[<210> 9]]> <![CDATA[<211> 439]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 智人]]> <![CDATA[<400> 9]]> Met Pro Leu Asn Val Ser Phe Thr Asn Arg Asn Tyr Asp Leu Asp Tyr 1 5 10 15 Asp Ser Val Gln Pro Tyr Phe Tyr Cys Asp Glu Glu Glu Asn Phe Tyr 20 25 30 Gln Gln Gln Gln Gln Ser Glu Leu Gln Pro Pro Ala Pro Ser Glu Asp 35 40 45 Ile Trp Lys Lys Phe Glu Leu Leu Pro Thr Pro Pro Leu Ser Pro Ser 50 55 60 Arg Arg Ser Gly Leu Cys Ser Pro Ser Tyr Val Ala Val Thr Pro Phe 65 70 75 80 Ser Leu Arg Gly Asp Asn Asp Gly Gly Gly Gly Ser Phe Ser Thr Ala 85 90 95 Asp Gln Leu Glu Met Val Thr Glu Leu Leu Gly Gly Asp Met Val Asn 100 105 110 Gln Ser Phe Ile Cys Asp Pro Asp Asp Glu Thr Phe Ile Lys Asn Ile 115 120 125 Ile Ile Gln Asp Cys Met Trp Ser Gly Phe Ser Ala Ala Ala Lys Leu 130 135 140 Val Ser Glu Lys Leu Ala Ser Tyr Gln Ala Ala Arg Lys Asp Ser Gly 145 150 155 160 Ser Pro Asn Pro Ala Arg Gly His Ser Val Cys Ser Thr Ser Ser Leu 165 170 175 Tyr Leu Gln Asp Leu Ser Ala Ala Ala Ser Glu Cys Ile Asp Pro Ser 180 185 190 Val Val Phe Pro Tyr Pro Leu Asn Asp Ser Ser Ser Pro Lys Ser Cys 195 200 205 Ala Ser Gln Asp Ser Ser Ala Phe Ser Pro Ser Ser Asp Ser Leu Leu 210 215 220 Ser Ser Thr Glu Ser Ser Pro Gln Gly Ser Pro Glu Pro Leu Val Leu 225 230 235 240 His Glu Glu Thr Pro Pro Thr Thr Ser Ser Asp Ser Glu Glu Glu Gln 245 250 255 Glu Asp Glu Glu Glu Ile Asp Val Val Ser Val Glu Lys Arg Gln Ala 260 265 270 Pro Gly Lys Arg Ser Glu Ser Gly Ser Pro Ser Ala Gly Gly His Ser 275 280 285 Lys Pro Pro His Ser Pro Leu Val Leu Lys Arg Cys His Val Ser Thr 290 295 300 His Gln His Asn Tyr Ala Ala Pro Pro Ser Thr Arg Lys Asp Tyr Pro 305 310 315 320 Ala Ala Lys Arg Val Lys Leu Asp Ser Val Arg Val Leu Arg Gln Ile 325 330 335 Ser Asn Asn Arg Lys Cys Thr Ser Pro Arg Ser Ser Asp Thr Glu Glu 340 345 350 Asn Val Lys Arg Arg Thr His Asn Val Leu Glu Arg Gln Arg Arg Asn 355 360 365 Glu Leu Lys Arg Ser Phe Phe Ala Leu Arg Asp Gln Ile Pro Glu Leu 370 375 380 Glu Asn Asn Glu Lys Ala Pro Lys Val Val Ile Leu Lys Lys Ala Thr 385 390 395 400 Ala Tyr Ile Leu Ser Val Gln Ala Glu Glu Gln Lys Leu Ile Ser Glu 405 410 415 Glu Asp Leu Leu Arg Lys Arg Arg Glu Gln Leu Lys His Lys Leu Glu 420 425 430 Gln Leu Arg Asn Ser Cys Ala 435 <![CDATA[<210> 10]]> <![CDATA[<211> 440]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 人造序列]]> <![CDATA[<220>]]> <![CDATA[<223> 人造序列之描述:合成多肽]]> <![CDATA[<400> 10]]> Pro Met Pro Leu Asn Val Ser Phe Thr Asn Arg Asn Tyr Asp Leu Asp 1 5 10 15 Tyr Asp Ser Val Gln Pro Tyr Phe Tyr Cys Asp Glu Glu Glu Asn Phe 20 25 30 Tyr Gln Gln Gln Gln Gln Ser Glu Leu Gln Pro Pro Ala Pro Ser Glu 35 40 45 Asp Ile Trp Lys Lys Phe Glu Leu Leu Pro Thr Pro Pro Leu Ser Pro 50 55 60 Ser Arg Arg Ser Gly Leu Cys Ser Pro Ser Tyr Val Ala Val Thr Pro 65 70 75 80 Phe Ser Leu Arg Gly Asp Asn Asp Gly Gly Gly Gly Ser Phe Ser Thr 85 90 95 Ala Asp Gln Leu Glu Met Val Thr Glu Leu Leu Gly Gly Asp Met Val 100 105 110 Asn Gln Ser Phe Ile Cys Asp Pro Asp Asp Glu Thr Phe Ile Lys Asn 115 120 125 Ile Ile Ile Gln Asp Cys Met Trp Ser Gly Phe Ser Ala Ala Ala Lys 130 135 140 Leu Val Ser Glu Lys Leu Ala Ser Tyr Gln Ala Ala Arg Lys Asp Ser 145 150 155 160 Gly Ser Pro Asn Pro Ala Arg Gly His Ser Val Cys Ser Thr Ser Ser 165 170 175 Leu Tyr Leu Gln Asp Leu Ser Ala Ala Ala Ser Glu Cys Ile Asp Pro 180 185 190 Ser Val Val Phe Pro Tyr Pro Leu Asn Asp Ser Ser Ser Pro Lys Ser 195 200 205 Cys Ala Ser Gln Asp Ser Ser Ala Phe Ser Pro Ser Ser Asp Ser Leu 210 215 220 Leu Ser Ser Thr Glu Ser Ser Pro Gln Gly Ser Pro Glu Pro Leu Val 225 230 235 240 Leu His Glu Glu Thr Pro Pro Thr Thr Ser Ser Asp Ser Glu Glu Glu 245 250 255 Gln Glu Asp Glu Glu Glu Ile Asp Val Val Ser Val Glu Lys Arg Gln 260 265 270 Ala Pro Gly Lys Arg Ser Glu Ser Gly Ser Pro Ser Ala Gly Gly His 275 280 285 Ser Lys Pro Pro His Ser Pro Leu Val Leu Lys Arg Cys His Val Ser 290 295 300 Thr His Gln His Asn Tyr Ala Ala Pro Pro Ser Thr Arg Lys Asp Tyr 305 310 315 320 Pro Ala Ala Lys Arg Val Lys Leu Asp Ser Val Arg Val Leu Arg Gln 325 330 335 Ile Ser Asn Asn Arg Lys Cys Thr Ser Pro Arg Ser Ser Asp Thr Glu 340 345 350 Glu Asn Val Lys Arg Arg Thr His Asn Val Leu Glu Arg Gln Arg Arg 355 360 365 Asn Glu Leu Lys Arg Ser Phe Phe Ala Leu Arg Asp Gln Ile Pro Glu 370 375 380 Leu Glu Asn Asn Glu Lys Ala Pro Lys Val Val Ile Leu Lys Lys Ala 385 390 395 400 Thr Ala Tyr Ile Leu Ser Val Gln Ala Glu Glu Gln Lys Leu Ile Ser 405 410 415 Glu Glu Asp Leu Leu Arg Lys Arg Arg Glu Gln Leu Lys His Lys Leu 420 425 430 Glu Gln Leu Arg Asn Ser Cys Ala 435 440 <![CDATA[<210> 11]]> <![CDATA[<211> 620]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 智人]]> <![CDATA[<400> 11]]> Met Ala Glu Ala Arg Thr Ser Leu Ser Ala His Cys Arg Gly Pro Leu 1 5 10 15 Ala Thr Gly Leu His Pro Asp Leu Asp Leu Pro Gly Arg Ser Leu Ala 20 25 30 Thr Pro Ala Pro Ser Cys Tyr Leu Leu Gly Ser Glu Pro Ser Ser Gly 35 40 45 Leu Gly Leu Gln Pro Glu Thr His Leu Pro Glu Gly Ser Leu Lys Arg 50 55 60 Cys Cys Val Leu Gly Leu Pro Pro Thr Ser Pro Ala Ser Ser Ser Pro 65 70 75 80 Cys Ala Ser Ser Asp Val Thr Ser Ile Ile Arg Ser Ser Gln Thr Ser 85 90 95 Leu Val Thr Cys Val Asn Gly Leu Arg Ser Pro Pro Leu Thr Gly Asp 100 105 110 Leu Gly Gly Pro Ser Lys Arg Ala Arg Pro Gly Pro Ala Ser Thr Asp 115 120 125 Ser His Glu Gly Ser Leu Gln Leu Glu Ala Cys Arg Lys Ala Ser Phe 130 135 140 Leu Lys Gln Glu Pro Ala Asp Glu Phe Ser Glu Leu Phe Gly Pro His 145 150 155 160 Gln Gln Gly Leu Pro Pro Pro Tyr Pro Leu Ser Gln Leu Pro Pro Gly 165 170 175 Pro Ser Leu Gly Gly Leu Gly Leu Gly Leu Ala Gly Arg Val Val Ala 180 185 190 Gly Arg Gln Ala Cys Arg Trp Val Asp Cys Cys Ala Ala Tyr Glu Gln 195 200 205 Gln Glu Glu Leu Val Arg His Ile Glu Lys Ser His Ile Asp Gln Arg 210 215 220 Lys Gly Glu Asp Phe Thr Cys Phe Trp Ala Gly Cys Val Arg Arg Tyr 225 230 235 240 Lys Pro Phe Asn Ala Arg Tyr Lys Leu Leu Ile His Met Arg Val His 245 250 255 Ser Gly Glu Lys Pro Asn Lys Cys Met Phe Glu Gly Cys Ser Lys Ala 260 265 270 Phe Ser Arg Leu Glu Asn Leu Lys Ile His Leu Arg Ser His Thr Gly 275 280 285 Glu Lys Pro Tyr Leu Cys Gln His Pro Gly Cys Gln Lys Ala Phe Ser 290 295 300 Asn Ser Ser Asp Arg Ala Lys His Gln Arg Thr His Leu Asp Thr Lys 305 310 315 320 Pro Tyr Ala Cys Gln Ile Pro Gly Cys Ser Lys Arg Tyr Thr Asp Pro 325 330 335 Ser Ser Leu Arg Lys His Val Lys Ala His Ser Ala Lys Glu Gln Gln 340 345 350 Val Arg Lys Lys Leu His Ala Gly Pro Asp Thr Glu Ala Asp Val Leu 355 360 365 Thr Glu Cys Leu Val Leu Gln Gln Leu His Thr Ser Thr Gln Leu Ala 370 375 380 Ala Ser Asp Gly Lys Gly Gly Cys Gly Leu Gly Gln Glu Leu Leu Pro 385 390 395 400 Gly Val Tyr Pro Gly Ser Ile Thr Pro His Asn Gly Leu Ala Ser Gly 405 410 415 Leu Leu Pro Pro Ala His Asp Val Pro Ser Arg His His Pro Leu Asp 420 425 430 Ala Thr Thr Ser Ser His His His Leu Ser Pro Leu Pro Met Ala Glu 435 440 445 Ser Thr Arg Asp Gly Leu Gly Pro Gly Leu Leu Ser Pro Ile Val Ser 450 455 460 Pro Leu Lys Gly Leu Gly Pro Pro Pro Leu Pro Pro Ser Ser Gln Ser 465 470 475 480 His Ser Pro Gly Gly Gln Pro Phe Pro Thr Leu Pro Ser Lys Pro Ser 485 490 495 Tyr Pro Pro Phe Gln Ser Pro Pro Pro Pro Pro Leu Pro Ser Pro Gln 500 505 510 Gly Tyr Gln Gly Ser Phe His Ser Ile Gln Ser Cys Phe Pro Tyr Gly 515 520 525 Asp Cys Tyr Arg Met Ala Glu Pro Ala Ala Gly Gly Asp Gly Leu Val 530 535 540 Gly Glu Thr His Gly Phe Asn Pro Leu Arg Pro Asn Gly Tyr His Ser 545 550 555 560 Leu Ser Thr Pro Leu Pro Ala Thr Gly Tyr Glu Ala Leu Ala Glu Ala 565 570 575 Ser Cys Pro Thr Ala Leu Pro Gln Gln Pro Ser Glu Asp Val Val Ser 580 585 590 Ser Gly Pro Glu Asp Cys Gly Phe Phe Pro Asn Gly Ala Phe Asp His 595 600 605 Cys Leu Gly His Ile Pro Ser Ile Tyr Thr Asp Thr 610 615 620 <![CDATA[<210> 12]]> <![CDATA[<211> 305]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 智人]]> <![CDATA[<400> 12]]> Met Ser Val Asp Pro Ala Cys Pro Gln Ser Leu Pro Cys Phe Glu Ala 1 5 10 15 Ser Asp Cys Lys Glu Ser Ser Pro Met Pro Val Ile Cys Gly Pro Glu 20 25 30 Glu Asn Tyr Pro Ser Leu Gln Met Ser Ser Ala Glu Met Pro His Thr 35 40 45 Glu Thr Val Ser Pro Leu Pro Ser Ser Met Asp Leu Leu Ile Gln Asp 50 55 60 Ser Pro Asp Ser Ser Thr Ser Pro Lys Gly Lys Gln Pro Thr Ser Ala 65 70 75 80 Glu Lys Ser Val Ala Lys Lys Glu Asp Lys Val Pro Val Lys Lys Gln 85 90 95 Lys Thr Arg Thr Val Phe Ser Ser Thr Gln Leu Cys Val Leu Asn Asp 100 105 110 Arg Phe Gln Arg Gln Lys Tyr Leu Ser Leu Gln Gln Met Gln Glu Leu 115 120 125 Ser Asn Ile Leu Asn Leu Ser Tyr Lys Gln Val Lys Thr Trp Phe Gln 130 135 140 Asn Gln Arg Met Lys Ser Lys Arg Trp Gln Lys Asn Asn Trp Pro Lys 145 150 155 160 Asn Ser Asn Gly Val Thr Gln Lys Ala Ser Ala Pro Thr Tyr Pro Ser 165 170 175 Leu Tyr Ser Ser Tyr His Gln Gly Cys Leu Val Asn Pro Thr Gly Asn 180 185 190 Leu Pro Met Trp Ser Asn Gln Thr Trp Asn Asn Ser Thr Trp Ser Asn 195 200 205 Gln Thr Gln Asn Ile Gln Ser Trp Ser Asn His Ser Trp Asn Thr Gln 210 215 220 Thr Trp Cys Thr Gln Ser Trp Asn Asn Gln Ala Trp Asn Ser Pro Phe 225 230 235 240 Tyr Asn Cys Gly Glu Glu Ser Leu Gln Ser Cys Met Gln Phe Gln Pro 245 250 255 Asn Ser Pro Ala Ser Asp Leu Glu Ala Ala Leu Glu Ala Ala Gly Glu 260 265 270 Gly Leu Asn Val Ile Gln Gln Thr Thr Arg Tyr Phe Ser Thr Pro Gln 275 280 285 Thr Met Asp Leu Phe Leu Asn Tyr Ser Met Asn Met Gln Pro Glu Asp 290 295 300 Val 305 <![CDATA[<210> 13]]> <![CDATA[<211> 258]]> <![CDATA[<212> PRT]]> <![CDATA[<213> 智人]]> <![CDATA[<400> 13]]> Met Arg Ser Phe Asn Gln Val Ser Ser Ala Pro Gly Gly Ala Ser Lys 1 5 10 15 Gly Gly Gly Glu Glu Pro Gly Lys Leu Pro Glu Pro Ala Glu Glu Glu 20 25 30 Ser Gln Val Leu Arg Gly Thr Gly His Cys Lys Trp Phe Asn Val Arg 35 40 45 Met Gly Phe Gly Phe Ile Ser Met Ile Asn Arg Glu Gly Ser Pro Leu 50 55 60 Asp Ile Pro Val Asp Val Phe Val His Gln Ser Lys Leu Phe Met Glu 65 70 75 80 Gly Phe Arg Ser Leu Lys Glu Gly Glu Pro Val Glu Phe Thr Phe Lys 85 90 95 Lys Ser Ser Lys Gly Leu Glu Ser Ile Arg Val Thr Gly Pro Gly Gly 100 105 110 Ser Pro Cys Leu Gly Ser Glu Arg Arg Pro Lys Gly Lys Thr Leu Gln 115 120 125 Lys Arg Lys Pro Lys Gly Asp Arg Cys Tyr Asn Cys Gly Gly Leu Asp 130 135 140 His His Ala Lys Glu Cys Ser Leu Pro Pro Gln Pro Lys Lys Cys His 145 150 155 160 Tyr Cys Gln Ser Ile Met His Met Val Ala Asn Cys Pro His Lys Asn 165 170 175 Val Ala Gln Pro Pro Ala Ser Ser Gln Gly Arg Gln Glu Ala Glu Ser 180 185 190 Gln Pro Cys Thr Ser Thr Leu Pro Arg Glu Val Gly Gly Gly His Gly 195 200 205 Cys Thr Ser Pro Pro Phe Pro Gln Glu Ala Arg Ala Glu Ile Ser Glu 210 215 220 Arg Ser Gly Arg Ser Pro Gln Glu Ala Ser Ser Thr Lys Ser Ser Ile 225 230 235 240 Ala Pro Glu Glu Gln Ser Lys Lys Gly Pro Ser Val Gln Lys Arg Lys 245 250 255 Lys Thr <![CDATA[<210> 14]]> <![CDATA[<211> 7675]]> <![CDATA[<212> DNA]]> <![CDATA[<213> 委內瑞拉馬腦炎病毒]]> <![CDATA[<400> 14]]> gaaagttcac gttgacatcg aggaagacag cccattcctc agagctttgc agcggagctt 60 cccgcagttt gaggtagaag ccaagcaggt cactgataat gaccatgcta atgccagagc 120 gttttcgcat ctggcttcaa aactgatcga aacggaggtg gacccatccg acacgatcct 180 tgacattgga agtgcgcccg cccgcagaat gtattctaag cacaagtatc attgtatctg 240 tccgatgaga tgtgcggaag atccggacag attgtataag tatgcaacta agctgaagaa 300 aaactgtaag gaaataactg ataaggaatt ggacaagaaa atgaaggagc tcgccgccgt 360 catgagcgac cctgacctgg aaactgagac tatgtgcctc cacgacgacg agtcgtgtcg 420 ctacgaaggg caagtcgctg tttaccagga tgtatacgcg gttgacggac cgacaagtct 480 ctatcaccaa gccaataagg gagttagagt cgcctactgg ataggctttg acaccacccc 540 ttttatgttt aagaacttgg ctggagcata tccatcatac tctaccaact gggccgacga 600 aaccgtgtta acggctcgta acataggcct atgcagctct gacgttatgg agcggtcacg 660 tagagggatg tccattctta gaaagaagta tttgaaacca tccaacaatg ttctattctc 720 tgttggctcg accatctacc acgagaagag ggacttactg aggagctggc acctgccgtc 780 tgtatttcac ttacgtggca agcaaaatta cacatgtcgg tgtgagacta tagttagttg 840 cgacgggtac gtcgttaaaa gaatagctat cagtccaggc ctgtatggga agccttcagg 900 ctatgctgct acgatgcacc gcgagggatt cttgtgctgc aaagtgacag acacattgaa 960 cggggagagg gtctcttttc ccgtgtgcac gtatgtgcca gctacattgt gtgaccaaat 1020 gactggcata ctggcaacag atgtcagtgc ggacgacgcg caaaaactgc tggttgggct 1080 caaccagcgt atagtcgtca acggtcgcac ccagagaaac accaatacca tgaaaaatta 1140 ccttttgccc gtagtggccc aggcatttgc taggtgggca aaggaatata aggaagatca 1200 agaagatgaa aggccactag gactacgaga tagacagtta gtcatggggt gttgttgggc 1260 ttttagaagg cacaagataa catctattta taagcgcccg gatacccaaa ccatcatcaa 1320 agtgaacagc gatttccact cattcgtgct gcccaggata ggcagtaaca cattggagat 1380 cgggctgaga acaagaatca ggaaaatgtt agaggagcac aaggagccgt cacctctcat 1440 taccgccgag gacgtacaag aagctaagtg cgcagccgat gaggctaagg aggtgcgtga 1500 agccgaggag ttgcgcgcag ctctaccacc tttggcagct gatgttgagg agcccactct 1560 ggaagccgat gtcgacttga tgttacaaga ggctggggcc ggctcagtgg agacacctcg 1620 tggcttgata aaggttacca gctacgctgg cgaggacaag atcggctctt acgctgtgct 1680 ttctccgcag gctgtactca agagtgaaaa attatcttgc atccaccctc tcgctgaaca 1740 agtcatagtg ataacacact ctggccgaaa agggcgttat gccgtggaac cataccatgg 1800 taaagtagtg gtgccagagg gacatgcaat acccgtccag gactttcaag ctctgagtga 1860 aagtgccacc attgtgtaca acgaacgtga gttcgtaaac aggtacctgc accatattgc 1920 cacacatgga ggagcgctga acactgatga agaatattac aaaactgtca agcccagcga 1980 gcacgacggc gaatacctgt acgacatcga caggaaacag tgcgtcaaga aagaactagt 2040 cactgggcta gggctcacag gcgagctggt ggatcctccc ttccatgaat tcgcctacga 2100 gagtctgaga acacgaccag ccgctcctta ccaagtacca accatagggg tgtatggcgt 2160 gccaggatca ggcaagtctg gcatcattaa aagcgcagtc accaaaaaag atctagtggt 2220 gagcgccaag aaagaaaact gtgcagaaat tataagggac gtcaagaaaa tgaaagggct 2280 ggacgtcaat gccagaactg tggactcagt gctcttgaat ggatgcaaac accccgtaga 2340 gaccctgtat attgacgaag cttttgcttg tcatgcaggt actctcagag cgctcatagc 2400 cattataaga cctaaaaagg cagtgctctg cggggatccc aaacagtgcg gtttttttaa 2460 catgatgtgc ctgaaagtgc attttaacca cgagatttgc acacaagtct tccacaaaag 2520 catctctcgc cgttgcacta aatctgtgac ttcggtcgtc tcaaccttgt tttacgacaa 2580 aaaaatgaga acgacgaatc cgaaagagac taagattgtg attgacacta ccggcagtac 2640 caaacctaag caggacgatc tcattctcac ttgtttcaga gggtgggtga agcagttgca 2700 aatagattac aaaggcaacg aaataatgac ggcagctgcc tctcaagggc tgacccgtaa 2760 aggtgtgtat gccgttcggt acaaggtgaa tgaaaatcct ctgtacgcac ccacctcaga 2820 acatgtgaac gtcctactga cccgcacgga ggaccgcatc gtgtggaaaa cactagccgg 2880 cgacccatgg ataaaaacac tgactgccaa gtaccctggg aatttcactg ccacgataga 2940 ggagtggcaa gcagagcatg atgccatcat gaggcacatc ttggagagac cggaccctac 3000 cgacgtcttc cagaataagg caaacgtgtg ttgggccaag gctttagtgc cggtgctgaa 3060 gaccgctggc atagacatga ccactgaaca atggaacact gtggattatt ttgaaacgga 3120 caaagctcac tcagcagaga tagtattgaa ccaactatgc gtgaggttct ttggactcga 3180 tctggactcc ggtctatttt ctgcacccac tgttccgtta tccattagga ataatcactg 3240 ggataactcc ccgtcgccta acatgtacgg gctgaataaa gaagtggtcc gtcagctctc 3300 tcgcaggtac ccacaactgc ctcgggcagt tgccactgga agagtctatg acatgaacac 3360 tggtacactg cgcaattatg atccgcgcat aaacctagta cctgtaaaca gaagactgcc 3420 tcatgcttta gtcctccacc ataatgaaca cccacagagt gacttttctt cattcgtcag 3480 caaattgaag ggcagaactg tcctggtggt cggggaaaag ttgtccgtcc caggcaaaat 3540 ggttgactgg ttgtcagacc ggcctgaggc taccttcaga gctcggctgg atttaggcat 3600 cccaggtgat gtgcccaaat atgacataat atttgttaat gtgaggaccc catataaata 3660 ccatcactat cagcagtgtg aagaccatgc cattaagctt agcatgttga ccaagaaagc 3720 ttgtctgcat ctgaatcccg gcggaacctg tgtcagcata ggttatggtt acgctgacag 3780 ggccagcgaa agcatcattg gtgctatagc gcggcagttc aagttttccc gggtatgcaa 3840 accgaaatcc tcacttgaag agacggaagt tctgtttgta ttcattgggt acgatcgcaa 3900 ggcccgtacg cacaatcctt acaagctttc atcaaccttg accaacattt atacaggttc 3960 cagactccac gaagccggat gtgcaccctc atatcatgtg gtgcgagggg atattgccac 4020 ggccaccgaa ggagtgatta taaatgctgc taacagcaaa ggacaacctg gcggaggggt 4080 gtgcggagcg ctgtataaga aattcccgga aagcttcgat ttacagccga tcgaagtagg 4140 aaaagcgcga ctggtcaaag gtgcagctaa acatatcatt catgccgtag gaccaaactt 4200 caacaaagtt tcggaggttg aaggtgacaa acagttggca gaggcttatg agtccatcgc 4260 taagattgtc aacgataaca attacaagtc agtagcgatt ccactgttgt ccaccggcat 4320 cttttccggg aacaaagatc gactaaccca atcattgaac catttgctga cagctttaga 4380 caccactgat gcagatgtag ccatatactg cagggacaag aaatgggaaa tgactctcaa 4440 ggaagcagtg gctaggagag aagcagtgga ggagatatgc atatccgacg actcttcagt 4500 gacagaacct gatgcagagc tggtgagggt gcatccgaag agttctttgg ctggaaggaa 4560 gggctacagc acaagcgatg gcaaaacttt ctcatatttg gaagggacca agtttcacca 4620 ggcggccaag gatatagcag aaattaatgc catgtggccc gttgcaacgg aggccaatga 4680 gcaggtatgc atgtatatcc tcggagaaag catgagcagt attaggtcga aatgccccgt 4740 cgaagagtcg gaagcctcca caccacctag cacgctgcct tgcttgtgca tccatgccat 4800 gactccagaa agagtacagc gcctaaaagc ctcacgtcca gaacaaatta ctgtgtgctc 4860 atcctttcca ttgccgaagt atagaatcac tggtgtgcag aagatccaat gctcccagcc 4920 tatattgttc tcaccgaaag tgcctgcgta tattcatcca aggaagtatc tcgtggaaac 4980 accaccggta gacgagactc cggagccatc ggcagagaac caatccacag aggggacacc 5040 tgaacaacca ccacttataa ccgaggatga gaccaggact agaacgcctg agccgatcat 5100 catcgaagag gaagaagagg atagcataag tttgctgtca gatggcccga cccaccaggt 5160 gctgcaagtc gaggcagaca ttcacgggcc gccctctgta tctagctcat cctggtccat 5220 tcctcatgca tccgactttg atgtggacag tttatccata cttgacaccc tggagggagc 5280 tagcgtgacc agcggggcaa cgtcagccga gactaactct tacttcgcaa agagtatgga 5340 gtttctggcg cgaccggtgc ctgcgcctcg aacagtattc aggaaccctc cacatcccgc 5400 tccgcgcaca agaacaccgt cacttgcacc cagcagggcc tgctcgagaa ccagcctagt 5460 ttccaccccg ccaggcgtga atagggtgat cactagagag gagctcgagg cgcttacccc 5520 gtcacgcact cctagcaggt cggtctcgag aaccagcctg gtctccaacc cgccaggcgt 5580 aaatagggtg attacaagag aggagtttga ggcgttcgta gcacaacaac aatgacggtt 5640 tgatgcgggt gcatacatct tttcctccga caccggtcaa gggcatttac aacaaaaatc 5700 agtaaggcaa acggtgctat ccgaagtggt gttggagagg accgaattgg agatttcgta 5760 tgccccgcgc ctcgaccaag aaaaagaaga attactacgc aagaaattac agttaaatcc 5820 cacacctgct aacagaagca gataccagtc caggaaggtg gagaacatga aagccataac 5880 agctagacgt attctgcaag gcctagggca ttatttgaag gcagaaggaa aagtggagtg 5940 ctaccgaacc ctgcatcctg ttcctttgta ttcatctagt gtgaaccgtg ccttttcaag 6000 ccccaaggtc gcagtggaag cctgtaacgc catgttgaaa gagaactttc cgactgtggc 6060 ttcttactgt attattccag agtacgatgc ctatttggac atggttgacg gagcttcatg 6120 ctgcttagac actgccagtt tttgccctgc aaagctgcgc agctttccaa agaaacactc 6180 ctatttggaa cccacaatac gatcggcagt gccttcagcg atccagaaca cgctccagaa 6240 cgtcctggca gctgccacaa aaagaaattg caatgtcacg caaatgagag aattgcccgt 6300 attggattcg gcggccttta atgtggaatg cttcaagaaa tatgcgtgta ataatgaata 6360 ttgggaaacg tttaaagaaa accccatcag gcttactgaa gaaaacgtgg taaattacat 6420 taccaaatta aaaggaccaa aagctgctgc tctttttgcg aagacacata atttgaatat 6480 gttgcaggac ataccaatgg acaggtttgt aatggactta aagagagacg tgaaagtgac 6540 tccaggaaca aaacatactg aagaacggcc caaggtacag gtgatccagg ctgccgatcc 6600 gctagcaaca gcgtatctgt gcggaatcca ccgagagctg gttaggagat taaatgcggt 6660 cctgcttccg aacattcata cactgtttga tatgtcggct gaagactttg acgctattat 6720 agccgagcac ttccagcctg gggattgtgt tctggaaact gacatcgcgt cgtttgataa 6780 aagtgaggac gacgccatgg ctctgaccgc gttaatgatt ctggaagact taggtgtgga 6840 cgcagagctg ttgacgctga ttgaggcggc tttcggcgaa atttcatcaa tacatttgcc 6900 cactaaaact aaatttaaat tcggagccat gatgaaatct ggaatgttcc tcacactgtt 6960 tgtgaacaca gtcattaaca ttgtaatcgc aagcagagtg ttgagagaac ggctaaccgg 7020 atcaccatgt gcagcattca ttggagatga caatatcgtg aaaggagtca aatcggacaa 7080 attaatggca gacaggtgcg ccacctggtt gaatatggaa gtcaagatta tagatgctgt 7140 ggtgggcgag aaagcgcctt atttctgtgg agggtttatt ttgtgtgact ccgtgaccgg 7200 cacagcgtgc cgtgtggcag accccctaaa aaggctgttt aagcttggca aacctctggc 7260 agcagacgat gaacatgatg atgacaggag aagggcattg catgaagagt caacacgctg 7320 gaaccgagtg ggtattcttt cagagctgtg caaggcagta gaatcaaggt atgaaaccgt 7380 aggaacttcc atcatagtta tggccatgac tactctagct agcagtgtta aatcattcag 7440 ctacctgaga ggggccccta taactctcta cggctaacct gaatggacta cgacatagtc 7500 tagtccgcca agtctagcat atgggcgcgt gaattcgccg cgaattggca agctgcttac 7560 atagaactcg cggcgattgg catgccgcct taaaattttt attttatttt tcttttcttt 7620 tccgaatcgg attttgtttt taatatttca aaaaaaaaaa aaaaaaaaaa aaaaa 7675 <![CDATA[<210> 15]]> <![CDATA[<211> 7675]]> <![CDATA[<212> DNA]]> <![CDATA[<213> 人造序列]]> <![CDATA[<220>]]> <![CDATA[<223> 人造序列之描述:合成多核苷酸]]> <![CDATA[<400> 15]]> gaaagttcac gttgacatcg aggaagacag cccattcctc agagctttgc agcggagctt 60 cccgcagttt gaggtagaag ccaagcaggt cactgataat gaccatgcta atgccagagc 120 gttttcgcat ctggcttcaa aactgatcga aacggaggtg gacccatccg acacgatcct 180 tgacattgga agtgcgcccg cccgcagaat gtattctaag cacaagtatc attgtatctg 240 tccgatgaga tgtgcggaag atccggacag attgtataag tatgcaacta agctgaagaa 300 aaactgtaag gaaataactg ataaggaatt ggacaagaaa atgaaggagc tggccgccgt 360 catgagcgac cctgacctgg aaactgagac tatgtgcctc cacgacgacg agtcgtgtcg 420 ctacgaaggg caagtcgctg tttaccagga tgtatacgcg gttgacggac cgacaagtct 480 ctatcaccaa gccaataagg gagttagagt cgcctactgg ataggctttg acaccacccc 540 ttttatgttt aagaacttgg ctggagcata tccatcatac tctaccaact gggccgacga 600 aaccgtgtta acggctcgta acataggcct atgcagctct gacgttatgg agcggtcacg 660 tagagggatg tccattctta gaaagaagta tttgaaacca tccaacaatg ttctattctc 720 tgttggctcg accatctacc acgagaagag ggacttactg aggagctggc acctgccgtc 780 tgtatttcac ttacgtggca agcaaaatta cacatgtcgg tgtgagacta tagttagttg 840 cgacgggtac gtcgttaaaa gaatagctat cagtccaggc ctgtatggga agccttcagg 900 ctatgctgct acgatgcacc gcgagggatt cttgtgctgc aaagtgacag acacattgaa 960 cggggagagg gtctcttttc ccgtgtgcac gtatgtgcca gctacattgt gtgaccaaat 1020 gactggcata ctggcaacag atgtcagtgc ggacgacgcg caaaaactgc tggttgggct 1080 caaccagcgt atagtcgtca acggtcgcac ccagagaaac accaatacca tgaaaaatta 1140 ccttttgccc gtagtggccc aggcatttgc taggtgggca aaggaatata aggaagatca 1200 agaagatgaa aggccactag gactacgaga tagacagtta gtcatggggt gttgttgggc 1260 ttttagaagg cacaagataa catctattta taagcgcccg gatacccaaa ccatcatcaa 1320 agtgaacagc gatttccact cattcgtgct gcccaggata ggcagtaaca cattggagat 1380 cgggctgaga acaagaatca ggaaaatgtt agaggagcac aaggagccgt cacctctcat 1440 taccgccgag gacgtacaag aagctaagtg cgcagccgat gaggctaagg aggtgcgtga 1500 agccgaggag ttgcgcgcag ctctaccacc tttggcagct gatgttgagg agcccactct 1560 ggaggcagac gtcgacttga tgttacaaga ggctggggcc ggctcagtgg agacacctcg 1620 tggcttgata aaggttacca gctacgatgg cgaggacaag atcggctctt acgctgtgct 1680 ttctccgcag gctgtactca agagtgaaaa attatcttgc atccaccctc tcgctgaaca 1740 agtcatagtg ataacacact ctggccgaaa agggcgttat gccgtggaac cataccatgg 1800 taaagtagtg gtgccagagg gacatgcaat acccgtccag gactttcaag ctctgagtga 1860 aagtgccacc attgtgtaca acgaacgtga gttcgtaaac aggtacctgc accatattgc 1920 cacacatgga ggagcgctga acactgatga agaatattac aaaactgtca agcccagcga 1980 gcacgacggc gaatacctgt acgacatcga caggaaacag tgcgtcaaga aagaactagt 2040 cactgggcta gggctcacag gcgagctggt ggatcctccc ttccatgaat tcgcctacga 2100 gagtctgaga acacgaccag ccgctcctta ccaagtacca accatagggg tgtatggcgt 2160 gccaggatca ggcaagtctg gcatcattaa aagcgcagtc accaaaaaag atctagtggt 2220 gagcgccaag aaagaaaact gtgcagaaat tataagggac gtcaagaaaa tgaaagggct 2280 ggacgtcaat gccagaactg tggactcagt gctcttgaat ggatgcaaac accccgtaga 2340 gaccctgtat attgacgaag cttttgcttg tcatgcaggt actctcagag cgctcatagc 2400 cattataaga cctaaaaagg cagtgctctg cggggatccc aaacagtgcg gtttttttaa 2460 catgatgtgc ctgaaagtgc attttaacca cgagatttgc acacaagtct tccacaaaag 2520 catctctcgc cgttgcacta aatctgtgac ttcggtcgtc tcaaccttgt tttacgacaa 2580 aaaaatgaga acgacgaatc cgaaagagac taagattgtg attgacacta ccggcagtac 2640 caaacctaag caggacgatc tcattctcac ttgtttcaga gggtgggtga agcagttgca 2700 aatagattac aaaggcaacg aaataatgac ggcagctgcc tctcaagggc tgacccgtaa 2760 aggtgtgtat gccgttcggt acaaggtgaa tgaaaatcct ctgtacgcac ccacctcaga 2820 acatgtgaac gtcctactga cccgcacgga ggaccgcatc gtgtggaaaa cactagccgg 2880 cgacccatgg ataaaaacac tgactgccaa gtaccctggg aatttcactg ccacgataga 2940 ggagtggcaa gcagagcatg atgccatcat gaggcacatc ttggagagac cggaccctac 3000 cgacgtcttc cagaataagg caaacgtgtg ttgggccaag gctttagtgc cggtgctgaa 3060 gaccgctggc atagacatga ccactgaaca atggaacact gtggattatt ttgaaacgga 3120 caaagctcac tcagcagaga tagtattgaa ccaactatgc gtgaggttct ttggactcga 3180 tctggactcc ggtctatttt ctgcacccac tgttccgtta tccattagga ataatcactg 3240 ggataactcc ccgtcgccta acatgtacgg gctgaataaa gaagtggtcc gtcagctctc 3300 tcgcaggtac ccacaactgc ctcgggcagt tgccactgga agagtctatg acatgaacac 3360 tggtacactg cgcaattatg atccgcgcat aaacctagta cctgtaaaca gaagactgcc 3420 tcatgcttta gtcctccacc ataatgaaca cccacagagt gacttttctt cattcgtcag 3480 caaattgaag ggcagaactg tcctggtggt cggggaaaag ttgtccgtcc caggcaaaat 3540 ggttgactgg ttgtcagacc ggcctgaggc taccttcaga gctcggctgg atttaggcat 3600 cccaggtgat gtgcccaaat atgacataat atttgttaat gtgaggaccc catataaata 3660 ccatcactat cagcagtgtg aagaccatgc cattaagctt agcatgttga ccaagaaagc 3720 ttgtctgcat ctgaatcccg gcggaacctg tgtcagcata ggttatggtt acgctgacag 3780 ggccagcgaa agcatcattg gtgctatagc gcggcagttc aagttttccc gggtatgcaa 3840 accgaaatcc tcacttgaag agacggaagt tctgtttgta ttcattgggt acgatcgcaa 3900 ggcccgtacg cacaattctt acaagctttc atcaaccttg accaacattt atacaggttc 3960 cagactccac gaagccggat gtgcaccctc atatcatgtg gtgcgagggg atattgccac 4020 ggccaccgaa ggagtgatta taaatgctgc taacagcaaa ggacaacctg gcggaggggt 4080 gtgcggagcg ctgtataaga aattcccgga aagcttcgat ttacagccga tcgaagtagg 4140 aaaagcgcga ctggtcaaag gtgcagctaa acatatcatt catgccgtag gaccaaactt 4200 caacaaagtt tcggaggttg aaggtgacaa acagttggca gaggcttatg agtccatcgc 4260 taagattgtc aacgataaca attacaagtc agtagcgatt ccactgttgt ccaccggcat 4320 cttttccggg aacaaagatc gactaaccca atcattgaac catttgctga cagctttaga 4380 caccactgat gcagatgtag ccatatactg cagggacaag aaatgggaaa tgactctcaa 4440 ggaagcagtg gctaggagag aagcagtgga ggagatatgc atatccgacg actcttcagt 4500 gacagaacct gatgcagagc tggtgagggt gcatccgaag agttctttgg ctggaaggaa 4560 gggctacagc acaagcgatg gcaaaacttt ctcatatttg gaagggacca agtttcacca 4620 ggcggccaag gatatagcag aaattaatgc catgtggccc gttgcaacgg aggccaatga 4680 gcaggtatgc atgtatatcc tcggagaaag catgagcagt attaggtcga aatgccccgt 4740 cgaagagtcg gaagcctcca caccacctag cacgctgcct tgcttgtgca tccatgccat 4800 gactccagaa agagtacagc gcctaaaagc ctcacgtcca gaacaaatta ctgtgtgctc 4860 atcctttcca ttgccgaagt atagaatcac tggtgtgcag aagatccaat gctcccagcc 4920 tatattgttc tcaccgaaag tgcctgcgta tattcatcca aggaagtatc tcgtggaaac 4980 accaccggta gacgagactc cggagccatc ggcagagaac caatccacag aggggacacc 5040 tgaacaacca ccacttataa ccgaggatga gaccaggact agaacgcctg agccgatcat 5100 catcgaagag gaagaagagg atagcataag tttgctgtca gatggcccga cccaccaggt 5160 gctgcaagtc gaggcagaca ttcacgggcc gccctctgta tctagctcat cctggtccat 5220 tcctcatgca tccgactttg atgtggacag tttatccata cttgacaccc tggagggagc 5280 tagcgtgacc agcggggcaa cgtcagccga gactaactct tacttcgcaa agagtatgga 5340 gtttctggcg cgaccggtgc ctgcgcctcg aacagtattc aggaaccctc cacatcccgc 5400 tccgcgcaca agaacaccgt cacttgcacc cagcagggcc tgctcgagaa ccagcctagt 5460 ttccaccccg ccaggcgtga atagggtgat cactagagag gagctcgagg cgcttacccc 5520 gtcacgcact cctagcaggt cggtctcgag aaccagcctg gtctccaacc cgccaggcgt 5580 aaatagggtg attacaagag aggagtttga ggcgttcgta gcacaacaac aatgacggtt 5640 tgatgcgggt gcatacatct tttcctccga caccggtcaa gggcatttac aacaaaaatc 5700 agtaaggcaa acggtgctat ccgaagtggt gttggagagg accgaattgg agatttcgta 5760 tgccccgcgc ctcgaccaag aaaaagaaga attactacgc aagaaattac agttaaatcc 5820 cacacctgct aacagaagca gataccagtc caggaaggtg gagaacatga aagccataac 5880 agctagacgt attctgcaag gcctagggca ttatttgaag gcagaaggaa aagtggagtg 5940 ctaccgaacc ctgcatcctg ttcctttgta ttcatctagt gtgaaccgtg ccttttcaag 6000 ccccaaggtc gcagtggaag cctgtaacgc catgttgaaa gagaactttc cgactgtggc 6060 ttcttactgt attattccag agtacgatgc ctatttggac atggttgacg gagcttcatg 6120 ctgcttagac actgccagtt tttgccctgc aaagctgcgc agctttccaa agaaacactc 6180 ctatttggaa cccacaatac gatcggcagt gccttcagcg atccagaaca cgctccagaa 6240 cgtcctggca gctgccacaa aaagaaattg caatgtcacg caaatgagag aattgcccgt 6300 attggattcg gcggccttta atgtggaatg cttcaagaaa tatgcgtgta ataatgaata 6360 ttgggaaacg tttaaagaaa accccatcag gcttactgaa gaaaacgtgg taaattacat 6420 taccaaatta aaaggaccaa aagctgctgc tctttttgcg aagacacata atttgaatat 6480 gttgcaggac ataccaatgg acaggtttgt aatggactta aagagagacg tgaaagtgac 6540 tccaggaaca aaacatactg aagaacggcc caaggtacag gtgatccagg ctgccgatcc 6600 gctagcaaca gcgtatctgt gcggaatcca ccgagagctg gttaggagat taaatgcggt 6660 cctgcttccg aacattcata cactgtttga tatgtcggct gaagactttg acgctattat 6720 agccgagcac ttccagcctg gggattgtgt tctggaaact gacatcgcgt cgtttgataa 6780 aagtgaggac gacgccatgg ctctgaccgc gttaatgatt ctggaagact taggtgtgga 6840 cgcagagctg ttgacgctga ttgaggcggc tttcggcgaa atttcatcaa tacatttgcc 6900 cactaaaact aaatttaaat tcggagccat gatgaaatct ggaatgttcc tcacactgtt 6960 tgtgaacaca gtcattaaca ttgtaatcgc aagcagagtg ttgagagaac ggctaaccgg 7020 atcaccatgt gcagcattca ttggagatga caatatcgtg aaaggagtca aatcggacaa 7080 attaatggca gacaggtgcg ccacctggtt gaatatggaa gtcaagatta tagatgctgt 7140 ggtgggcgag aaagcgcctt atttctgtgg agggtttatt ttgtgtgact ccgtgaccgg 7200 cacagcgtgc cgtgtggcag accccctaaa aaggctgttt aagcttggca aacctctggc 7260 agcagacgat gaacatgatg atgacaggag aagggcattg catgaagagt caacacgctg 7320 gaaccgagtg ggtattcttt cagagctgtg caaggcagta gaatcaaggt atgaaaccgt 7380 aggaacttcc atcatagtta tggccatgac tactctagct agcagtgtta aatcattcag 7440 ctacctgaga ggggccccta taactctcta cggctaacct gaatggacta cgacatagtc 7500 tagtccgcca agtctagcat atgggcgcgt gaattcgccg cgaattggca agctgcttac 7560 atagaactcg cggcgattgg catgccgcct taaaattttt attttatttt tcttttcttt 7620 tccgaatcgg attttgtttt taatatttca aaaaaaaaaa aaaaaaaaaa aaaaa 7675
Claims (30)
- 一種自造血譜系之起始細胞獲得誘導型多能幹細胞(iPSC)群體之方法,其包括: 將編碼BCL-xL及選自OCT家族蛋白、KLF家族蛋白、MYC家族蛋白、SOX家族蛋白、LIN28蛋白、NANOG蛋白及p53顯性負性蛋白之一或多種附加再程式化因子之α病毒RNA表現構築體引入該等起始細胞,及 培養該等起始細胞以允許BCL-xL及該一或多種附加再程式化因子之表現,從而誘導該等起始細胞及其後代再程式化為iPSC。
- 如請求項1之方法,其中該等起始細胞為人類起源之造血幹細胞、紅血球系祖細胞、淋巴系祖細胞、外周血單核細胞、T淋巴細胞、B淋巴細胞、巨噬細胞、單核細胞、嗜中性白血球、嗜酸性白血球或樹突狀細胞。
- 如請求項2之方法,其中該等起始細胞為藉由在存在促紅血球生成素、幹細胞因子及IL-3下培養外周血單核細胞(PBMC)視情況持續5至10天或6至7天獲得之紅血球系祖細胞。
- 如請求項3之方法,其中該等PBMC係在存在0.5至5 IU/ml促紅血球生成素、50至200 ng/mL幹細胞因子及1至10 ng/mL IL-3下培養。
- 如請求項1至5中任一項之方法,其中該RNA表現構築體透過電穿孔引入該等起始細胞。
- 如請求項5之方法,其包括在電穿孔之前使該等起始細胞與B18R蛋白接觸。
- 一種自如請求項1至6中任一項之方法獲得之誘導型多能幹細胞(iPSC)之群體。
- 一種自造血譜系之起始細胞獲得之誘導型多能幹細胞(iPSC)的群體,該等起始細胞經編碼BCL-xL及選自OcT家族蛋白、KLF家族蛋白、MyC家族蛋白、SOX家族蛋白、LIN28蛋白、NANOG蛋白及p53顯性負性蛋白之一或多種附加再程式化因子之α病毒RNA表現構築體轉染。
- 如請求項8之iPSC,其中該等起始細胞為人類起源之造血幹細胞、紅血球系祖細胞、淋巴系祖細胞、外周血單核細胞、T淋巴細胞、B淋巴細胞、巨噬細胞、單核細胞、嗜中性白血球、嗜酸性白血球或樹突狀細胞。
- 如請求項9之iPSC,其中該等起始細胞為藉由在存在EPO、SCF及IL-3下培養外周血單核細胞(PBMC)視情況持續5至10天或6至7天獲得之紅血球系祖細胞。
- 一種編碼BCL-xL及選自OCT家族蛋白、KLF家族蛋白、MYC家族蛋白、SOX家族蛋白、LIN28蛋白、NANOG蛋白及p53顯性負性蛋白之一或多種附加再程式化因子之α病毒RNA表現構築體。
- 一種DNA載體,其包含如請求項11之α病毒RNA表現構築體之編碼序列。
- 一種包含如請求項11之α病毒RNA表現構築體或如請求項12之DNA載體之宿主細胞,視情況其中該宿主細胞為人類細胞。
- 如請求項1至6中任一項之方法,如請求項7至10中任一項之iPSC,如請求項11之α病毒RNA表現構築體,如請求項12之DNA載體或如請求項13之宿主細胞,其中該α病毒RNA表現構築體係自我複製型且包含nsP1、nsP2、nsP3及nsP4基因。
- 如上述請求項中任一項之方法、iPSC、α病毒RNA表現構築體、DNA載體或宿主細胞,其中該α病毒RNA表現構築體為委內瑞拉馬腦炎病毒(VEEV) RNA表現構築體且包含VEEV nsP1、nsP2、nsP3及nsP4基因, 視情況其中該VEEV RNA表現構築體含有來自野生型VEEV基因組之該(等)對應區域之一或多個,視情況兩個或更多個、三個或更多個、四個或更多個、五個或更多個、或六個或更多個突變。
- 如上述請求項中任一項之方法、iPSC、α病毒RNA表現構築體、DNA載體或宿主細胞,其中 該OCT家族蛋白為OCT4,視情況人類OCT4, 該KLF家族蛋白為KLF4,視情況人類KLF4, 該SOX家族蛋白為SOX2,視情況人類SOX2, 該LIN28蛋白為LIN28B,視情況人類LIN28B,及/或 該MYC家族蛋白為c-MYC,視情況人類c-MYC。
- 如上述請求項中任一項之方法、iPSC、α病毒RNA表現構築體、DNA載體或宿主細胞,其中該α病毒RNA表現構築體指導以下之表現: 包含SEQ ID NO:1或與其至少95%相同之胺基酸序列之BCL-xL蛋白, 包含SEQ ID NO:3或與其至少95%相同之胺基酸序列之OCT4蛋白, 包含SEQ ID NO:5或與其至少95%相同之胺基酸序列之KLF4蛋白, 包含SEQ ID NO:7或與其至少95%相同之胺基酸序列之SOX2蛋白,及/或 包含SEQ ID NO:9或與其至少95%相同之胺基酸序列之c-MYC蛋白。
- 如上述請求項中任一項之方法、iPSC、α病毒RNA表現構築體、DNA載體或宿主細胞,其中該α病毒RNA表現構築體指導OCT家族蛋白、SOX家族蛋白、BCL-xL及MYC家族蛋白及視情況KLF家族蛋白之表現。
- 如上述請求項中任一項之方法、iPSC、α病毒RNA表現構築體、DNA載體或宿主細胞,其中BCL-xL及該一或多種附加再程式化因子之該等編碼序列被2A肽之編碼序列或內部核糖體進入位點(IRES)分隔開。
- 如上述請求項中任一項之方法、iPSC、α病毒RNA表現構築體、DNA載體或宿主細胞,其中BCL-xL及該一或多種附加再程式化因子之該等編碼序列係在共同(common)啟動子,視情況26S啟動子之轉錄控制下。
- 一種於活體外獲得分化細胞之方法,其包括在存在分化促進劑下培養如請求項7至10及14至20中任一項之iPSC。
- 一種藉由如請求項21之方法獲得之分化細胞。
- 一種自分化如請求項7至10及14至20中任一項之iPSC獲得之分化細胞。
- 如請求項21之方法或如請求項22或23之分化細胞,其中該分化細胞為 人類免疫細胞,視情況選自T細胞、表現嵌合抗原受體(CAR)之T細胞、抑制性T細胞、骨髓細胞、樹突狀細胞及免疫抑制性巨噬細胞; 人類神經系統中之細胞,視情況選自多巴胺能神經元、小膠質細胞、少突膠質細胞、星形膠質細胞、皮質神經元、脊髓或動眼神經元、腸神經元、基板衍生細胞、施旺(Schwann)細胞及三叉神經或感覺神經元; 人類心血管系統中之細胞,視情況選自心肌細胞、內皮細胞及結節細胞; 人類代謝系統中之細胞,視情況選自肝細胞、膽管細胞及胰β細胞,或 人類眼部系統中之細胞,視情況選自視網膜色素上皮細胞、感光錐形細胞、感光桿狀細胞、雙極細胞或神經節細胞。
- 如請求項21至24中任一項之方法或細胞,其中該分化細胞為外胚層譜系,視情況其中該分化細胞為神經元。
- 如請求項21至24中任一項之方法或細胞,其中該分化細胞為中胚層譜系,視情況其中該分化細胞為心肌細胞。
- 一種醫藥組合物,其包含如請求項22至26中任一項之分化細胞及醫藥上可接受之載劑。
- 一種治療有需要患者之方法,其包括向該患者投與如請求項22至26中任一項之分化細胞或如請求項27之醫藥組合物。
- 一種如請求項22至26中任一項之分化細胞之用途,其用於製造用於治療有需要患者之藥劑。
- 如請求項22至26中任一項之分化細胞或如請求項27之醫藥組合物,其用於治療有需要患者。
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JP6166900B2 (ja) | 2009-08-24 | 2017-07-19 | ウイスコンシン アラムナイ リサーチ ファウンデーシヨンWisconsin Alumni Research Foundation | 実質的に純粋なヒト網膜前駆細胞培養物、前脳前駆細胞培養物、網膜色素上皮細胞培養物、及び、それらの製造方法 |
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WO2013056072A1 (en) | 2011-10-13 | 2013-04-18 | Wisconsin Alumni Research Foundation | Generation of cardiomyocytes from human pluripotent stem cells |
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EP3259348A4 (en) | 2015-02-17 | 2018-07-18 | University Health Network | Methods for making and using sinoatrial node-like pacemaker cardiomyocytes and ventricular-like cardiomyocytes |
CA2985714C (en) * | 2015-06-12 | 2024-02-06 | Lonza Walkersville, Inc. | Methods for nuclear reprogramming using synthetic transcription factors |
CN114807035B (zh) | 2015-09-08 | 2024-02-02 | (由卫生与公众服务部部长代表的)美利坚合众国 | 临床级别视网膜色素上皮细胞的可再现的分化方法 |
WO2017044488A1 (en) | 2015-09-08 | 2017-03-16 | Cellular Dynamics International, Inc. | Macs-based purification of stem cell-derived retinal pigment epithelium |
WO2018098597A1 (en) | 2016-12-04 | 2018-06-07 | University Health Network | Generating atrial and ventricular cardiomyocyte lineages from human pluripotent stem cells |
CN112204134A (zh) | 2018-04-20 | 2021-01-08 | 富士胶片细胞动力公司 | 眼细胞的分化方法及其用途 |
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