CN115916961A - T细胞 - Google Patents
T细胞 Download PDFInfo
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- CN115916961A CN115916961A CN202180036346.5A CN202180036346A CN115916961A CN 115916961 A CN115916961 A CN 115916961A CN 202180036346 A CN202180036346 A CN 202180036346A CN 115916961 A CN115916961 A CN 115916961A
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Abstract
本发明涉及T细胞以及用于产生组织驻留记忆T细胞(TRM)的方法。本发明涉及通过本发明的方法获取的组织驻留记忆T细胞(TRM)本身、包含这些TRM细胞的组合物以及这些TRM细胞和组合物在治疗中的应用,例如应用于治疗癌症的免疫疗法。
Description
技术领域
本发明涉及T细胞,特别涉及但不限于用于产生组织驻留记忆T细胞(TRM)的方法、通过本发明的这些方法获取的组织驻留记忆T细胞(TRM)本身、包含这些TRM细胞的组合物、以及这些TRM细胞及组合物在治疗中的应用,例如应用于治疗癌症的免疫疗法。
背景技术
使用免疫检查点抑制剂的免疫疗法,例如阻断针对PD-1和CTLA-4的抗体,显著地提升了无癌生存期。重要的是,过继转移,例如嵌合抗原受体(CAR)T细胞、一般肿瘤检测delta-oneγδT细胞(DOT)[2,3]或是MR-1限制性T细胞[4],已经取得了很有希望的成果。然而,这些方法对于实体肿瘤是否足够有效仍是一个问题,例如对于乳腺癌,只有一小部分患者获得持久的治疗反应。T细胞免疫疗法的成功,特别是在实体肿瘤中,取决于在肿瘤组织内传递和激活具有细胞毒活性的肿瘤特异性淋巴细胞,例如CD8+T细胞。
组织免疫的突破揭示了组织驻留记忆CD8+T(TRM)细胞的存在[5,6],它具有深入组织的能力。近年来,揭示了肿瘤中TRM细胞的存在与患者的阳性预后之间存在强相关性,其中,肿瘤中TRM细胞通过其标记物CD103来识别。TRM细胞与乳腺癌、肺癌、卵巢癌和宫颈癌等实体瘤的总生存期和无病状态延长密切相关,相关性比CD8+T细胞的总数更高[7-15]。事实上,在实体肿瘤中,TRM细胞与增强细胞毒性T细胞的应答直接相关[16,17]。
因此,在探索改进的免疫治疗机制时,关注CD8+T细胞的产生以及将其输送至组织深处是非常重要的,并且是对于用于治疗实体肿瘤的成功的基于细胞的癌症免疫疗法的重要步骤改变。
发明内容
本发明人假设调节性T细胞(TREG)在一种T细胞的产生中是重要的,这种T细胞能够深入组织并对实体肿瘤非常有效。本发明人已经能够描述和识别产生TRM细胞所需的因子,从而能够在体外产生TRM细胞,目的是产生具有组织穿透性的抗肿瘤T细胞。
因此,在本发明的第一方面,提供了一种用于产生组织驻留记忆T细胞(TRM)的方法,该方法包括转化生长因子β(TGFβ)的存在下培养淋巴细胞,以及/或者将该淋巴细胞与调节性T细胞共同培养。
有利的是,如实施例中所述,本发明人开发了一种方案,通过建立组织穿透性T细胞(即组织驻留记忆T细胞)的开发所需的体外要求来产生用于细胞疗法的T细胞。这种细胞的产生将导致TRM细胞的产生,TRM细胞能够在疾病组织(如肿瘤组织及转移性肿瘤)中输送和激活具有细胞毒性活性的疾病特异性淋巴细胞,从而显著拓宽了基于T细胞疗法的治疗工具包。此外,如实施例中所述,本发明人开发了一种新的方案来产生用于细胞疗法的TRM细胞,在其培养物中不包括TREG细胞。
优选地,该方法在体外进行。
优选地,该方法包括在TGFβ的存在下培养淋巴细胞。优选地,在一些实施例中,该方法不包括在调节性T细胞的存在下培养淋巴细胞。
优选地,TGFβ是具有生物活性的(即活化的)。优选地,TGFβ是哺乳动物的。TGFβ可以是啮齿动物、狗、马或猪的TGFβ。啮齿动物可以是大鼠或小鼠。最优选地,TGFβ是人类的TGFβ。
在一个实施例中,TGFβ是如美国国家生物技术信息中心(Genebank)ID No:7040所表示的TGFβ1,在本文中作为SEQ ID No:1提供,如下所示:
MPPSGLRLLPLLLPLLWLLVLTPGRPAAGLSTCKTIDMELVKRKRIEAIRGQILSKLRLASPPSQGEVPPGPLPEAVLALYNSTRDRVAGESAEPEPEPEADYYAKEVTRVLMVETHNEIYDKFKQSTHSIYMFFNTSELREAVPEPVLLSRAELRLLRLKLKVEQHVELYQKYSNNSWRYLSNRLLAPSDSPEWLSFDVTGVVRQWLSRGGEIEGFRLSAHCSCDSRDNTLQVDINGFTTGRRGDLATIHGMNRPFLLLMATPLERAQHLQSSRHRRALDTNYCFSSTEKNCCVRQLYIDFRKDLGWKWIHEPKGYHANFCLGPCPYIWSLDTQYSKVLALYNQHNPGASAAPCCVPQALEPLPIVYYVGRKPKVEQLSNMIVRSCKCS
[SEQ ID No:1]
因此,优选地,TGFβ包含SEQ ID No:1中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,TGFβ是如Genebank ID No:7042所表示的TGFβ2,在本文中作为SEQ ID No:14提供,如下所示:
MHYCVLSAFLILHLVTVALSLSTCSTLDMDQFMRKRIEAIRGQILSKLKLTSPPEDYPEPEEVPPEVISIYNSTRDLLQEKASRRAAACERERSDEEYYAKEVYKIDMPPFFPSENAIPPTFYRPYFRIVRFDVSAMEKNASNLVKAEFRVFRLQNPKARVPEQRIELYQILKSKDLTSPTQRYIDSKVVKTRAEGEWLSFDVTDAVHEWLHHKDRNLGFKISLHCPCCTFVPSNNYIIPNKSEELEARFAGIDGTSTYTSGDQKTIKSTRKKNSGKTPHLLLMLLPSYRLESQQTNRRKKRALDAAYCFRNVQDNCCLRPLYIDFKRDLGWKWIHEPKGYNANFCAGACPYLWSSDTQHSRVLSLYNTINPEASASPCCVSQDLEPLTILYYIGKTPKIEQLSNMIVKSCKCS
[SEQ ID No:14]
因此,优选地,TGFβ包含SEQ ID No:14中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,TGFβ是如Genebank ID No:7043所表示的TGFβ3,在本文中作为SEQ ID No:16提供,如下所示:
MKMHLQRALVVLALLNFATVSLSLSTCTTLDFGHIKKKRVEAIRGQILSKLRLTSPPEPTVMTHVPYQVLALYNSTRELLEEMHGEREEGCTQENTESEYYAKEIHKFDMIQGLAEHNELAVCPKGITSKVFRFNVSSVEKNRTNLFRAEFRVLRVPNPSSKRNEQRIELFQILRPDEHIAKQRYIGGKNLPTRGTAEWLSFDVTDTVREWLLRRESNLGLEISIHCPCHTFQPNGDILENIHEVMEIKFKGVDNEDDHGRGDLGRLKKQKDHHNPHLILMMIPPHRLDNPGQGGQRKKRALDTNYCFRNLEENCCVRPLYIDFRQDLGWKWVHEPKGYYANFCSGPCPYLRSADTTHSTVLGLYNTLNPEASASPCCVPQDLEPLTILYYVGRTPKVEQLSNMVVKSCKCS
[SEQ ID No:16]
因此,优选地,TGFβ包含SEQ ID No:16中基本列出的序列、其片段或变体,或由其组成。
优选地,TGFβ以0.01ng/ml和50ng/ml之间的浓度存在。更优选地,TGFβ以0.1ng/ml和20ng/ml之间的浓度存在,或是以0.1ng/ml和10ng/ml之间的浓度存在。最优选地,TGFβ以0.25ng/ml和5ng/ml之间的浓度存在,并更优选地以0.5ng/ml和5ng/ml之间的浓度存在。
优选地,上述淋巴细胞是幼稚CD8+T淋巴细胞、效应CD8+T淋巴细胞或记忆CD8+T淋巴细胞。
优选地,上述淋巴细胞是幼稚CD8+T淋巴细胞或效应CD8+T淋巴细胞。
优选地,上述淋巴细胞是幼稚CD8+T淋巴细胞。当该淋巴细胞是人类的细胞时,该幼稚CD8+T淋巴细胞可通过分化簇45亚型RA(CD45RA+)、C-C趋化因子受体7型(CCR7+)和/或分化簇27(CD27+)的表达来定义。进一步,该幼稚CD8+T淋巴细胞的特征可表示为缺失分化簇45亚型RO(CD45RO-)的表达。
当淋巴细胞是鼠科的、优选为小鼠的淋巴细胞时,幼稚CD8+T淋巴细胞可通过分化簇67亚型L(CD67L+)、C-C趋化因子受体7型(CCR7+)、分化簇127(CD127+)和/或分化簇27(CD27+)的表达来定义。幼稚的CD8+T淋巴细胞可通过分化簇44(CD44+)的低表达水平进一步定义。
优选地,淋巴细胞是效应CD8+T淋巴细胞。当该淋巴细胞是人类的淋巴细胞时,效应CD8+T淋巴细胞的特征为具有分化簇45同种型RA(CD45RA+)和/或分化簇45异型RO(CD45RO+)的表达。效应CD8+T淋巴细胞的进一步特征为缺失C-C趋化因子受体7型(CCR7-)的表达。
当淋巴细胞是鼠科的、优选为小鼠的淋巴细胞时,效应CD8+T淋巴细胞的特征为分化簇44(CD44+)的高水平表达和/或缺失分化簇62配体(CD62L)的表达。
优选地,淋巴细胞是记忆CD8+T淋巴细胞。记忆CD8+T淋巴细胞可以是中央记忆CD8+T淋巴细胞或效应记忆CD8+T淋巴细胞。
当淋巴细胞是人类的淋巴细胞时,中央记忆CD8+T淋巴细胞的特征为具有分化簇45亚型RO(CD45RO+)的表达。中央记忆CD8+T淋巴细胞的进一步特征为缺失分化簇45亚型RA(CD45RA-)、C-C趋化因子受体7型(CCR7-)分化簇27(CD27-)和/或分化簇62L(CD62L-)的表达。
当淋巴细胞是鼠科的、优选为小鼠的淋巴细胞时,中央记忆CD8+T淋巴细胞的特征为分化簇44(CD44+)和/或分化簇62配体(CD62L)的高水平表达。
当淋巴细胞是人类的淋巴细胞时,效应记忆CD8+T淋巴细胞的特征为具有分化簇45亚型RO(CD45RO+)的表达。效应记忆CD8+T淋巴细胞的进一步特征为缺失分化簇45亚型RA(CD45RA-)、C-C趋化因子受体7型(CCR7-)分化簇27(CD27-)和/或分化簇62L(CD62L-)的表达。
当淋巴细胞是鼠科的、优选为小鼠的淋巴细胞时,效应记忆CD8+T淋巴细胞的特征为具有分化簇44(CD44+)的高水平表达和/或缺失分化簇62配体(CD62L)的表达。
在一个实施例中,CD45RA可以由Genebank ID No:5788表示,其在本文中作为SEQID No:18提供,如下所示:
MTMYLWLKLLAFGFAFLDTEVFVTGQSPTPSPTGLTTAKMPSVPLSSDPLPTHTTAFSPASTFERENDFSETTTSLSPDNTSTQVSPDSLDNASAFNTTGVSSVQTPHLPTHADSQTPSAGTDTQTFSGSAANAKLNPTPGSNAISDVPGERSTASTFPTDPVSPLTTTLSLAHHSSAALPARTSNTTITANTSDAYLNASETTTLSPSGSAVISTTTIATTPSKPTCDEKYANITVDYLYNKETKLFTAKLNVNENVECGNNTCTNNEVHNLTECKNASVSISHNSCTAPDKTLILDVPPGVEKFQLHDCTQVEKADTTICLKWKNIETFTCDTQNITYRFQCGNMIFDNKEIKLENLEPEHEYKCDSEILYNNHKFTNASKIIKTDFGSPGEPQIIFCRSEAAHQGVITWNPPQRSFHNFTLCYIKETEKDCLNLDKNLIKYDLQNLKPYTKYVLSLHAYIIAKVQRNGSAAMCHFTTKSAPPSQVWNMTVSMTSDNSMHVKCRPPRDRNGPHERYHLEVEAGNTLVRNESHKNCDFRVKDLQYSTDYTFKAYFHNGDYPGEPFILHHSTSYNSKALIAFLAFLIIVTSIALLVVLYKIYDLHKKRSCNLDEQQELVERDDEKQLMNVEPIHADILLETYKRKIADEGRLFLAEFQSIPRVFSKFPIKEARKPFNQNKNRYVDILPYDYNRVELSEINGDAGSNYINASYIDGFKEPRKYIAAQGPRDETVDDFWRMIWEQKATVIVMVTRCEEGNRNKCAEYWPSMEEGTRAFGDVVVKINQHKRCPDYIIQKLNIVNKKEKATGREVTHIQFTSWPDHGVPEDPHLLLKLRRRVNAFSNFFSGPIVVHCSAGVGRTGTYIGIDAMLEGLEAENKVDVYGYVVKLRRQRCLMVQVEAQYILIHQALVEYNQFGETEVNLSELHPYLHNMKKRDPPSEPSPLEAEFQRLPSYRSWRTQHIGNQEENKSKNRNSNVIPYDYNRVPLKHELEMSKESEHDSDESSDDDSDSEEPSKYINASFIMSYWKPEVMIAAQGPLKETIGDFWQMIFQRKVKVIVMLTELKHGDQEICAQYWGEGKQTYGDIEVDLKDTDKSSTYTLRVFELRHSKRKDSRTVYQYQYTNWSVEQLPAEPKELISMIQVVKQKLPQKNSSEGNKHHKSTPLLIHCRDGSQQTGIFCALLNLLESAETEEVVDIFQVVKALRKARPGMVSTFEQYQFLYDVIASTYPAQNGQVKKNNHQEDKIEFDNEVDKVKQDANCVNPLGAPEKLPEAKEQAEGSEPTSGTEGPEHSVNGPASPALNQGS
[SEQ ID No:18]
因此,优选地,CD45RA包含SEQ ID No:18中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,CD45RO可以由Genebank ID No:5788表示,其在本文中作为SEQID No:19提供,如下所示:
MTMYLWLKLLAFGFAFLDTEVFVTGQSPTPSPTDAYLNASETTTLSPSGSAVISTTTIATTPSKPTCDEKYANITVDYLYNKETKLFTAKLNVNENVECGNNTCTNNEVHNLTECKNASVSISHNSCTAPDKTLILDVPPGVEKFQLHDCTQVEKADTTICLKWKNIETFTCDTQNITYRFQCGNMIFDNKEIKLENLEPEHEYKCDSEILYNNHKFTNASKIIKTDFGSPGEPQIIFCRSEAAHQGVITWNPPQRSFHNFTLCYIKETEKDCLNLDKNLIKYDLQNLKPYTKYVLSLHAYIIAKVQRNGSAAMCHFTTKSAPPSQVWNMTVSMTSDNSMHVKCRPPRDRNGPHERYHLEVEAGNTLVRNESHKNCDFRVKDLQYSTDYTFKAYFHNGDYPGEPFILHHSTSYNSKALIAFLAFLIIVTSIALLVVLYKIYDLHKKRSCNLDEQQELVERDDEKQLMNVEPIHADILLETYKRKIADEGRLFLAEFQSIPRVFSKFPIKEARKPFNQNKNRYVDILPYDYNRVELSEINGDAGSNYINASYIDGFKEPRKYIAAQGPRDETVDDFWRMIWEQKATVIVMVTRCEEGNRNKCAEYWPSMEEGTRAFGDVVVKINQHKRCPDYIIQKLNIVNKKEKATGREVTHIQFTSWPDHGVPEDPHLLLKLRRRVNAFSNFFSGPIVVHCSAGVGRTGTYIGIDAMLEGLEAENKVDVYGYVVKLRRQRCLMVQVEAQYILIHQALVEYNQFGETEVNLSELHPYLHNMKKRDPPSEPSPLEAEFQRLPSYRSWRTQHIGNQEENKSKNRNSNVIPYDYNRVPLKHELEMSKESEHDSDESSDDDSDSEEPSKYINASFIMSYWKPEVMIAAQGPLKETIGDFWQMIFQRKVKVIVMLTELKHGDQEICAQYWGEGKQTYGDIEVDLKDTDKSSTYTLRVFELRHSKRKDSRTVYQYQYTNWSVEQLPAEPKELISMIQVVKQKLPQKNSSEGNKHHKSTPLLIHCRDGSQQTGIFCALLNLLESAETEEVVDIFQVVKALRKARPGMVSTFEQYQFLYDVIASTYPAQNGQVKKNNHQEDKIEFDNEVDKVKQDANCVNPLGAPEKLPEAKEQAEGSEPTSGTEGPEHSVNGPASPALNQGS
[SEQ ID No:19]
因此,优选地,CD45RO包含SEQ ID No:19中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,CCR7可由Genebank ID No:1236表示,其在本文中作为SEQ IDNo:20提供,如下所示:
MDLGKPMKSVLVVALLVIFQVCLCQDEVTDDYIGDNTTVDYTLFESLCSKKDVRNFKAWFLPIMYSIICFVGLLGNGLVVLTYIYFKRLKTMTDTYLLNLAVADILFLLTLPFWAYSAAKSWVFGVHFCKLIFAIYKMSFFSGMLLLLCISIDRYVAIVQAVSAHRHRARVLLISKLSCVGIWILATVLSIPELLYSDLQRSSSEQAMRCSLITEHVEAFITIQVAQMVIGFLVPLLAMSFCYLVIIRTLLQARNFERNKAIKVIIAVVVVFIVFQLPYNGVVLAQTVANFNITSSTCELSKQLNIAYDVTYSLACVRCCVNPFLYAFIGVKFRNDLFKLFKDLGCLSQEQLRQWSSCRHIRRSSMSVEAETTTTFSP
[SEQ ID No:20]
因此,优选地,CCR7包含SEQ ID No:20中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,CD27可以由Genebank ID No:939表示,其在本文中作为SEQ IDNo:21提供,如下所示:
MARPHPWWLCVLGTLVGLSATPAPKSCPERHYWAQGKLCCQMCEPGTFLVKDCDQHRKAAQCDPCIPGVSFSPDHHTRPHCESCRHCNSGLLVRNCTITANAECACRNGWQCRDKECTECDPLPNPSLTARSSQALSPHPQPTHLPYVSEMLEARTAGHMQTLADFRQLPARTLSTHWPPQRSLCSSDFIRILVIFSGMFLVFTLAGALFLHQRRKYRSNKGESPVEPAEPCHYSCPREEEGSTIPIQEDYRKPEPACSP
[SEQ ID No:21]
因此,优选地,CD27包含SEQ ID No:21中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,CD62L可由Genebank ID No:6402表示,其在本文中作为SEQ IDNo:22提供,如下所示:
MGCRRTREGPSKAMIFPWKCQSTQRDLWNIFKLWGWTMLCCDFLAHHGTDCWTYHYSEKPMNWQRARRFCRDNYTDLVAIQNKAEIEYLEKTLPFSRSYYWIGIRKIGGIWTWVGTNKSLTEEAENWGDGEPNNKKNKEDCVEIYIKRNKDAGKWNDDACHKLKAALCYTASCQPWSCSGHGECVEIINNYTCNCDVGYYGPQCQFVIQCEPLEAPELGTMDCTHPLGNFSFSSQCAFSCSEGTNLTGIEETTCGPFGNWSSPEPTCQVIQCEPLSAPDLGIMNCSHPLASFSFTSACTFICSEGTELIGKKKTICESSGIWSNPSPICQKLDKSFSMIKEGDYNPLFIPVAVMVTAFSGLAFIIWLARRLKKGKKSKRSMNDPY
[SEQ ID No:22]
因此,优选地,CD62L包含SEQ ID No:22中基本列出的序列、其片段或变体,或由其组成。
优选地,淋巴细胞从人类或非人动物的组织中获得。优选地,非人动物是哺乳动物。非人动物可以是啮齿动物、狗、马或猪。啮齿动物可以是大鼠或小鼠。优选地,淋巴细胞是从人类的组织中获得的。组织可选自:血液、脾脏、淋巴结、肺、胃肠道、皮肤、前列腺、乳腺组织、肝脏、骨髓和胰腺。优选地,组织是血液或骨髓。
该方法可包括从人类或非人动物获得的组织获得淋巴细胞。淋巴细胞可以通过本领域已知的任何合适的方法获得。获取方法包括血沉棕黄涂层或密度梯度、荧光活化细胞分选和/或磁活化细胞分选。这些方法为本领域技术人员所知。
优选地,通过本发明的方法产生的组织驻留记忆T细胞(TRM)是组织驻留记忆CD8+T细胞。优选地,使用该方法产生多个组织驻留记忆T细胞(TRM)。
组织驻留记忆CD8+T细胞的特征为具有分化簇8(CD8)、分化簇69(CD69)、锌指蛋白683(ZNF683/HOBIT)、芳基烃受体(AhR)和/或分化簇103(CD103)的表达。组织驻留记忆CD8+(细胞毒性)T细胞的进一步特征为缺失杀伤细胞凝集素样受体亚家族G成员(KLRG1)和/或脱中胚蛋白(Eomesodermin,Eomes)。
优选地,组织驻留记忆CD8+(细胞毒性)T细胞的特征为具有CD8、CD69、Hobit、AhR和CD103的表达。优选地,该组织驻留记忆CD8+的(细胞毒性的)T细胞的特征为具有CD8,CD69,Hobit,AhR,CD103的表达,并且缺失KLRG1和Eomes的表达。
在一个实施例中,CD8可以由Genebank ID No:925表示,其在本文中作为SEQ IDNo:2提供,如下所示:
MALPVTALLLPLALLLHAARPSQFRVSPLDRTWNLGETVELKCQVLLSNPTSGCSWLFQPRGAAASPTFLLYLSQNKPKAAEGLDTQRFSGKRLGDTFVLTLSDFRRENEGYYFCSALSNSIMYFSHFVPVFLPAKPTTTPAPRPPTPAPTIASQPLSLRPEACRPAAGGAVHTRGLDFACDIYIWAPLAGTCGVLLLSLVITLYCNHRNRRRVCKCPRPVVKSGDKPSLSARYV
[SEQ ID No:2]
因此,优选地,CD8包含SEQ ID No:2中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,CD69可由Genebank ID No:969表示,其在本文中作为SEQ IDNo:3提供,如下所示:
MSSENCFVAENSSLHPESGQENDATSPHFSTRHEGSFQVPVLCAVMNVVFITILIIALIALSVGQYNCPGQYTFSMPSDSHVSSCSEDWVGYQRKCYFISTVKRSWTSAQNACSEHGATLAVIDSEKDMNFLKRYAGREEHWVGLKKEPGHPWKWSNGKEFNNWFNVTGSDKCVFLKNTEVSSMECEKNLYWICNKPYK
[SEQ ID No:3]
因此,优选地,CD69包含SEQ ID No:3中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,CD103可由Genebank ID No:3682表示,其在本文中作为SEQ IDNo:4提供,如下所示:
MWLFHTLLCIASLALLAAFNVDVARPWLTPKGGAPFVLSSLLHQDPSTNQTWLLVTSPRTKRTPGPLHRCSLVQDEILCHPVEHVPIPKGRHRGVTVVRSHHGVLICIQVLVRRPHSLSSELTGTCSLLGPDLRPQAQANFFDLENLLDPDARVDTGDCYSNKEGGGEDDVNTARQRRALEKEEEEDKEEEEDEEEEEAGTEIAIILDGSGSIDPPDFQRAKDFISNMMRNFYEKCFECNFALVQYGGVIQTEFDLRDSQDVMASLARVQNITQVGSVTKTASAMQHVLDSIFTSSHGSRRKASKVMVVLTDGGIFEDPLNLTTVINSPKMQGVERFAIGVGEEFKSARTARELNLIASDPDETHAFKVTNYMALDGLLSKLRYNIISMEGTVGDALHYQLAQIGFSAQILDERQVLLGAVGAFDWSGGALLYDTRSRRGRFLNQTAAAAADAEAAQYSYLGYAVAVLHKTCSLSYIAGAPRYKHHGAVFELQKEGREASFLPVLEGEQMGSYFGSELCPVDIDMDGSTDFLLVAAPFYHVHGEEGRVYVYRLSEQDGSFSLARILSGHPGFTNARFGFAMAAMGDLSQDKLTDVAIGAPLEGFGADDGASFGSVYIYNGHWDGLSASPSQRIRASTVAPGLQYFGMSMAGGFDISGDGLADITVGTLGQAVVFRSRPVVRLKVSMAFTPSALPIGFNGVVNVRLCFEISSVTTASESGLREALLNFTLDVDVGKQRRRLQCSDVRSCLGCLREWSSGSQLCEDLLLMPTEGELCEEDCFSNASVKVSYQLQTPEGQTDHPQPILDRYTEPFAIFQLPYEKACKNKLFCVAELQLATTVSQQELVVGLTKELTLNINLTNSGEDSYMTSMALNYPRNLQLKRMQKPPSPNIQCDDPQPVASVLIMNCRIGHPVLKRSSAHVSVVWQLEENAFPNRTADITVTVTNSNERRSLANETHTLQFRHGFVAVLSKPSIMYVNTGQGLSHHKEFLFHVHGENLFGAEYQLQICVPTKLRGLQVVAVKKLTRTQASTVCTWSQERACAYSSVQHVEEWHSVSCVIASDKENVTVAAEISWDHSEELLKDVTELQILGEISFNKSLYEGLNAENHRTKITVVFLKDEKYHSLPIIIKGSVGGLLVLIVILVILFKCGFFKRKYQQLNLESIRKAQLKSENLLEEEN
[SEQ ID No:4]
因此,优选地,CD103包含SEQ ID No:4中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,KLRG1可以由Genebank ID No:10219表示,其在本文中作为SEQID No:5提供,如下所示:
MTDSVIYSMLELPTATQAQNDYGPQQKSSSSRPSCSCLVAIALGLLTAVLLSVLLYQWILCQGSNYSTCASCPSCPDRWMKYGNHCYYFSVEEKDWNSSLEFCLARDSHLLVITDNQEMSLLQVFLSEAFCWIGLRNNSGWRWEDGSPLNFSRISSNSFVQTCGAINKNGLQASSCEVPLHWVCKKCPFADQALF
[SEQ ID No:5]
因此,优选地,KLRG1包含SEQ ID No:5中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,Eomes可由Genebank ID No:8320表示,其在本文中作为SEQ IDNo:6提供,如下所示:
MQLGEQLLVSSVNLPGAHFYPLESARGGSGGSAGHLPSAAPSPQKLDLDKASKKFSGSLSCEAVSGEPAAASAGAPAAMLSDTDAGDAFASAAAVAKPGPPDGRKGSPCGEEELPSAAAAAAAAAAAAAATARYSMDSLSSERYYLQSPGPQGSELAAPCSLFPYQAAAGAPHGPVYPAPNGARYPYGSMLPPGGFPAAVCPPGRAQFGPGAGAGSGAGGSSGGGGGPGTYQYSQGAPLYGPYPGAAAAGSCGGLGGLGVPGSGFRAHVYLCNRPLWLKFHRHQTEMIITKQGRRMFPFLSFNINGLNPTAHYNVFVEVVLADPNHWRFQGGKWVTCGKADNNMQGNKMYVHPESPNTGSHWMRQEISFGKLKLTNNKGANNNNTQMIVLQSLHKYQPRLHIVEVTEDGVEDLNEPSKTQTFTFSETQFIAVTAYQNTDITQLKIDHNPFAKGFRDNYDSSHQIVPGGRYGVQSFFPEPFVNTLPQARYYNGERTVPQTNGLLSPQQSEEVANPPQRWLVTPVQQPGTNKLDISSYESEYTSSTLLPYGIKSLPLQTSHALGYYPDPTFPAMAGWGGRGSYQRKMAAGLPWTSRTSPTVFSEDQLSKEKVKEEIGSSWIETPPSIKSLDSNDSGVYTSACKRRRLSPSNSSNENSPSIKCEDINAEEYSKDTSKGMGGYYAFYTTP
[SEQ ID No:6]
因此,优选地,Eomes包含SEQ ID No:6中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,Hobit可由Genebank ID No:257101表示,其在本文中作为SEQID No:9提供,如下所示:
MKEESAAQLGCCHRPMALGGTGGSLSPSLDFQLFRGDQVFSACRPLPDMVDAHGPSCASWLCPLPLAPGRSALLACLQDLDLNLCTPQPAPLGTDLQGLQEDALSMKHEPPGLQASSTDDKKFTVKYPQNKDKLGKQPERAGEGAPCPAFSSHNSSSPPPLQNRKSPSPLAFCPCPPVNSISKELPFLLHAFYPGYPLLLPPPHLFTYGALPSDQCPHLLMLPQDPSYPTMAMPSLLMMVNELGHPSARWETLLPYPGAFQASGQALPSQARNPGAGAAPTDSPGLERGGMASPAKRVPLSSQTGTAALPYPLKKKNGKILYECNICGKSFGQLSNLKVHLRVHSGERPFQCALCQKSFTQLAHLQKHHLVHTGERPHKCSIPWVPGRNHWKSFQAWREREVCHKRFSSSSNLKTHLRLHSGARPFQCSVCRSRFTQHIHLKLHHRLHAPQPCGLVHTQLPLASLACLAQWHQGALDLMAVASEKHMGYDIDEVKVSSTSQGKARAVSLSSAGTPLVMGQDQNN
[SEQ ID No:9]
因此,优选地,Hobit包含SEQ ID No:9中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,Ahr可由Genebank ID No:196表示,其在本文中作为SEQ ID No:10提供,如下所示:
MNSSSANITYASRKRRKPVQKTVKPIPAEGIKSNPSKRHRDRLNTELDRLASLLPFPQDVINKLDKLSVLRLSVSYLRAKSFFDVALKSSPTERNGGQDNCRAANFREGLNLQEGEFLLQALNGFVLVVTTDALVFYASSTIQDYLGFQQSDVIHQSVYELIHTEDRAEFQRQLHWALNPSQCTESGQGIEEATGLPQTVVCYNPDQIPPENSPLMERCFICRLRCLLDNSSGFLAMNFQGKLKYLHGQKKKGKDGSILPPQLALFAIATPLQPPSILEIRTKNFIFRTKHKLDFTPIGCDAKGRIVLGYTEAELCTRGSGYQFIHAADMLYCAESHIRMIKTGESGMIVFRLLTKNNRWTWVQSNARLLYKNGRPDYIIVTQRPLTDEEGTEHLRKRNTKLPFMFTTGEAVLYEATNPFPAIMDPLPLRTKNGTSGKDSATTSTLSKDSLNPSSLLAAMMQQDESIYLYPASSTSSTAPFENNFFNESMNECRNWQDNTAPMGNDTILKHEQIDQPQDVNSFAGGHPGLFQDSKNSDLYSIMKNLGIDFEDIRHMQNEKFFRNDFSGEVDFRDIDLTDEILTYVQDSLSKSPFIPSDYQQQQSLALNSSCMVQEHLHLEQQQQHHQKQVVVEPQQQLCQKMKHMQVNGMFENWNSNQFVPFNCPQQDPQQYNVFTDLHGISQEFPYKSEMDSMPYTQNFISCNQPVLPQHSKCTELDYPMGSFEPSPYPTTSSLEDFVTCLQLPENQKHGLNPQSAIITPQTCYAGAVSMYQCQPEPQHTHVGQMQYNPVLPGQQAFLNKFQNGVLNETYPAELNNINNTQTTTHLQPLHHPSEARPFPDLTSSGFL
[SEQ ID No:10]
因此,优选地,Ahr包含SEQ ID No:10中基本列出的序列、其片段或变体,或由其组成。
优选地,该方法包括在白细胞介素2、4、7、12、15和/或21(IL-2、IL-4、IL-7、IL-12、IL-15和/或IL-21)的存在下培养上述淋巴细胞。
白细胞介素优选是哺乳动物的,最优选是人类的白细胞介素。
优选地,该方法包括在白细胞介素7(IL-7)存在下培养淋巴细胞。
优选地,IL-7是哺乳动物的。最优选地,IL-7是人类的IL-7。在一个实施例中,IL-7可由Genebank ID No:3574表示,其在本文中作为SEQ ID No:28提供,如下所示:
MFHVSFRYIFGLPPLILVLLPVASSDCDIEGKDGKQYESVLMVSIDQLLDSMKEIGSNCLNNEFNFFKRHICDANKEGMFLFRAARKLRQFLKMNSTGDFDLHLLKVSEGTTILLNCTGQVKGRKPAALGEAQPTKSLEENKSLKEQKKLNDLCFLKRLLQEIKTCWNKILMGTKEH
[SEQ ID No:28]
因此,优选地,IL-7包含SEQ ID No:28中基本列出的序列、其片段或变体,或由其组成。
优选地,IL-7可以0.1ng/ml和200ng/ml之间的浓度存在。更优选地,IL-7可以2ng/ml和100ng/ml之间的浓度存在。最优选地,IL-7可以10ng/ml和50ng/ml之间的浓度存在。
优选地,该方法还包括在白细胞介素15(IL-15)存在下培养淋巴细胞。
优选地,IL-15是哺乳动物的。最优选地,IL-15是人类的IL-15。在一个实施例中,IL-15可由Genebank ID No:3600表示,其在本文中作为SEQ ID No:7提供,如下所示:
MRISKPHLRSISIQCYLCLLLNSHFLTEAGIHVFILGCFSAGLPKTEANWVNVISDLKKIEDLIQSMHIDATLYTESDVHPSCKVTAMKCFLLELQVISLESGDASIHDTVENLIILANNSLSSNGNVTESGCKECEELEEKNIKEFLQSFVHIVQMFINTS
[SEQ ID No:7]
因此,优选地,IL-15包含SEQ ID No:7中基本列出的序列、其片段或变体,或由其组成。
优选地,IL-15可以1ng/ml和100ng/ml之间的浓度存在。更优选地,IL-15可以5ng/ml和50ng/ml之间的浓度存在。最优选地,IL-15可以10ng/ml和25ng/ml之间的浓度存在。
优选地,该方法还包括在白细胞介素33(IL-33)存在下进行培养。优选地,IL-33是哺乳动物的。最优选地,IL-33是人类的IL-33。在一个实施例中,IL-33可以由Genebank IDNo:90865表示,其在本文中作为SEQ ID No:8提供,如下所示:
MKPKMKYSTNKISTAKWKNTASKALCFKLGKSQQKAKEVCPMYFMKLRSGLMIKKEACYFRRETTKRPSLKTGRKHKRHLVLAACQQQSTVECFAFGISGVQKYTRALHDSSITGISPITEYLASLSTYNDQSITFALEDESYEIYVEDLKKDEKKDKVLLSYYESQHPSNESGDGVDGKMLMVTLSPTKDFWLHANNKEHSVELHKCEKPLPDQAFFVLHNMHSNCVSFECKTDPGVFIGVKDNHLALIKVDSSENLCTENILFKLSET
[SEQ ID No:8]
因此,优选地,IL-33包含SEQ ID No:8中基本列出的序列、其片段或变体,或由其组成。
优选地,IL-33可以0.5ng/ml和100ng/ml之间的浓度存在。更优选地,IL-33可以2ng/ml和50ng/ml之间的浓度存在。最优选地,IL-33可以10ng/ml和25ng/ml之间的浓度存在。
优选地,该方法包括在白细胞介素2(IL-2)存在下培养淋巴细胞。
因此,在一个实施例中,提供了一种用于产生组织驻留记忆T细胞(TRM)的方法,该方法包括在TGFβ存在下培养幼稚CD8+T淋巴细胞。
在另一个实施例中,提供了一种用于产生组织驻留记忆T细胞的方法,该方法包括在TGFβ、IL-15和IL-33的存在下培养幼稚CD8+T淋巴细胞。
因此,在一个实施例中,提供了一种用于产生组织驻留记忆CD8+(细胞毒性)T细胞的方法,该方法包括在TGFβ、IL-15和IL-33的存在下培养幼稚CD8+T淋巴细胞。
优选地,该方法还包括在至少一种白细胞介素1家族成员的存在下进行培养,例如在IL-1α、IL-1β和/或IL-18。
优选地,白细胞介素1家族成员是哺乳动物的。最优选地,白细胞介素1家族成员是人类的。
在一个实施例中,IL-1α可由Genebank ID No:3552表示,其在本文中作为SEQ IDNo:11提供,如下所示:
MAKVPDMFEDLKNCYSENEEDSSSIDHLSLNQKSFYHVSYGPLHEGCMDQSVSLSISETSKTSKLTFKESMVVVATNGKVLKKRRLSLSQSITDDDLEAIANDSEEEIIKPRSAPFSFLSNVKYNFMRIIKYEFILNDALNQSIIRANDQYLTAAALHNLDEAVKFDMGAYKSSKDDAKITVILRISKTQLYVTAQDEDQPVLLKEMPEIPKTITGSETNLLFFWETHGTKNYFTSVAHPNLFIATKQDYWVCLAGGPPSITDFQILENQA
[SEQ ID No:11]
因此,优选地,IL-1α包含SEQ ID No:11中基本列出的序列、其片段或变体,或由其组成。
优选地,IL-1α可以0.1ng/ml和100ng/ml之间的浓度存在。更优选地,IL-1α可以1ng/ml和50ng/ml之间的浓度存在。最优选地IL-1α可以5ng/ml和20ng/ml之间的浓度存在。
在一个实施例中,IL-1β可由Genebank ID No:3553表示,其在本文中作为SEQ IDNo:12提供,如下所示:
MAEVPELASEMMAYYSGNEDDLFFEADGPKQMKCSFQDLDLCPLDGGIQLRISDHHYSKGFRQAASVVVAMDKLRKMLVPCPQTFQENDLSTFFPFIFEEEPIFFDTWDNEAYVHDAPVRSLNCTLRDSQQKSLVMSGPYELKALHLQGQDMEQQVVFSMSFVQGEESNDKIPVALGLKEKNLYLSCVLKDDKPTLQLESVDPKNYPKKKMEKRFVFNKIEINNKLEFESAQFPNWYISTSQAENMPVFLGGTKGGQDITDFTMQFVSS
[SEQ ID No:12]
因此,优选地,IL-1β包含SEQ ID No:12中基本列出的序列、其片段或变体,或由其组成。
优选地,IL-1β可以0.1ng/ml和100ng/ml之间的浓度存在。更优选地,IL-1β可以1ng/ml和50ng/ml之间的浓度存在。最优选地,IL-1β可以5ng/ml和20ng/ml之间的浓度存在。
在一个实施例中,IL-18可由Genebank ID No:3606表示,其在本文中作为SEQ IDNo:13提供,如下所示:
MAAEPVEDNCINFVAMKFIDNTLYFIAEDDENLESDYFGKLESKLSVIRNLNDQVLFIDQGNRPLFEDMTDSDCRDNAPRTIFIISMYKDSQPRGMAVTISVKCEKISTLSCENKIISFKEMNPPDNIKDTKSDIIFFQRSVPGHDNKMQFESSSYEGYFLACEKERDLFKLILKKEDELGDRSIMFTVQNED
[SEQ ID No:13]
因此,优选地,IL-18包含SEQ ID No:13中基本列出的序列、其片段或变体,或由其组成。
优选地,IL-18可以0.1ng/ml和100ng/ml之间的浓度存在。更优选地,IL-18可以1ng/ml和50ng/ml之间的浓度存在。最优选地,IL-18可以5ng/ml和20ng/ml之间的浓度存在。
淋巴细胞可在包含至少一种芳烃受体(AhR)配体的培养基中培养。AhR配体可以是激动剂或拮抗剂。优选地,AhR配体是激动剂。优选地,AhR配体是拮抗剂。
AhR配体可选自卤代芳烃、多环芳烃、膳食衍生芳烃、血红素代谢物、靛蓝、芳烃受体拮抗剂StemRegenin 1和色氨酸代谢物。
卤代芳烃可以是四氯二苯并对二恶英(TCDD)。多环芳烃可以是3-甲基胆蒽。色氨酸代谢物可以是6-甲酰吲哚并[3,2-b]咔唑(FICZ)。膳食衍生芳烃可以是黄酮和/或吲哚衍生物。吲哚衍生物可以是吲哚-3-甲醇(I3C)和/或其产物二吲哚基甲烷(DIM)。
淋巴细胞可在包含至少一种脂质的培养基中培养。优选地,脂质是胆固醇和/或中链脂肪酸(MCFAs)。MCFA可以是油酸。
淋巴细胞可进一步与抗原一起培养。抗原的类型将取决于TRM细胞用于的治疗应用。例如,淋巴细胞可以与肿瘤抗原一起培养。
优选地,在一些实施例中,该方法包括在调节性T细胞或1型调节性T细胞的存在下培养淋巴细胞。
因此,在本发明的另一方面,提供了一种用于产生组织驻留记忆T细胞(TRM)的方法,该方法包括在转化生长因子β(TGFβ)的存在下培养淋巴细胞,以及/或者将淋巴细胞与1型调节性T细胞共同培养。
本领域技术人员能够理解,“调节性T细胞”是作为CD4+T细胞亚群参与外周免疫的一种T细胞。优选地,调节性T细胞的特征在于具有叉头盒P3转录因子(Foxp3)的表达。在另一些实施例中,该方法不包括在调节性T细胞存在下培养淋巴细胞。
本领域技术人员能够理解,“1型调节性T细胞”是作为CD4+T细胞亚群参与外周免疫的一种调节性T淋巴细胞。优选地,1型调节性T细胞的特征在于具有叉头盒P3转录因子(Foxp3)、T盒转录因子21(Tbet)和/或表面分子C-X-C基序趋化因子受体3(CXCR3)的表达。在另一些实施例中,该方法不包括在1型调节性T细胞存在下培养淋巴细胞。
在一个实施例中,Foxp3可由Genebank ID No:50943表示,其在本文中作为SEQ IDNo:23提供,如下所示:
MPNPRPGKPSAPSLALGPSPGASPSWRAAPKASDLLGARGPGGTFQGRDLRGGAHASSSSLNPMPPSQLQLPTLPLVMVAPSGARLGPLPHLQALLQDRPHFMHQLSTVDAHARTPVLQVHPLESPAMISLTPPTTATGVFSLKARPGLPPGINVASLEWVSREPALLCTFPNPSAPRKDSTLSAVPQSSYPLLANGVCKWPGCEKVFEEPEDFLKHCQADHLLDEKGRAQCLLQREMVQSLEQQLVLEKEKLSAMQAHLAGKMALTKASSVASSDKGSCCIVAAGSQGPVVPAWSGPREAPDSLFAVRRHLWGSHGNSTFPEFLHNMDYFKFHNMRPPFTYATLIRWAILEAPEKQRTLNEIYHWFTRMFAFFRNHPATWKNAIRHNLSLHKCFVRVESEKGAVWTVDELEFRKKRSQRPSRCSNPTPGP
[SEQ ID No:23]
因此,优选地,Foxp3包含SEQ ID No:23中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,Tbet可由Genebank ID No:30009表示,其在本文中作为SEQ IDNo:24提供,如下所示:
MGIVEPGCGDMLTGTEPMPGSDEGRAPGADPQHRYFYPEPGAQDADERRGGGSLGSPYPGGALVPAPPSRFLGAYAYPPRPQAAGFPGAGESFPPPADAEGYQPGEGYAAPDPRAGLYPGPREDYALPAGLEVSGKLRVALNNHLLWSKFNQHQTEMIITKQGRRMFPFLSFTVAGLEPTSHYRMFVDVVLVDQHHWRYQSGKWVQCGKAEGSMPGNRLYVHPDSPNTGAHWMRQEVSFGKLKLTNNKGASNNVTQMIVLQSLHKYQPRLHIVEVNDGEPEAACNASNTHIFTFQETQFIAVTAYQNAEITQLKIDNNPFAKGFRENFESMYTSVDTSIPSPPGPNCQFLGGDHYSPLLPNQYPVPSRFYPDLPGQAKDVVPQAYWLGAPRDHSYEAEFRAVSMKPAFLPSAPGPTMSYYRGQEVLAPGAGWPVAPQYPPKMGPASWFRPMRTLPMEPGPGGSEGRGPEDQGPPLVWTEIAPIRPESSDSGLGEGDSKRRRVSPYPSSGDSSSPAGAPSPFDKEAEGQFYNYFPN
[SEQ ID No:24]
因此,优选地,Tbet包含SEQ ID No:24中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,CXCR3可由Genebank ID No:2833表示,其在本文中作为SEQ IDNo:25提供,如下所示:
MELRKYGPGRLAGTVIGGAAQSKSQTKSDSITKEFLPGLYTAPSSPFPPSQVSDHQVLNDAEVAALLENFSSSYDYGENESDSCCTSPPCPQDFSLNFDRAFLPALYSLLFLLGLLGNGAVAAVLLSRRTALSSTDTFLLHLAVADTLLVLTLPLWAVDAAVQWVFGSGLCKVAGALFNINFYAGALLLACISFDRYLNIVHATQLYRRGPPARVTLTCLAVWGLCLLFALPDFIFLSAHHDERLNATHCQYNFPQVGRTALRVLQLVAGFLLPLLVMAYCYAHILAVLLVSRGQRRLRAMRLVVVVVVAFALCWTPYHLVVLVDILMDLGALARNCGRESRVDVAKSVTSGLGYMHCCLNPLLYAFVGVKFRERMWMLLLRLGCPNQRGLQRQPSSSRRDSSWSETSEASYSGL
[SEQ ID No:25]
因此,优选地,CXCR3包含SEQ ID No:25中基本列出的序列、其片段或变体,或由其组成。
优选地,调节性T细胞(优选为1型调节性T细胞)表达整合素alpha V beta 8(αvβ8)。本领域技术人员能够理解,αvβ8是整合素亚基(Itgβ8)和整合素亚基alpha V(Itgav)的二聚体。因此,优选地,调节性T细胞(优选为1型调节性T细胞)表达Itgβ8和Itgav。
在一个实施例中,Itgβ8可由Genebank ID No:3696表示,其在本文中作为SEQ IDNo:26提供,如下所示:
MCGSALAFFTAAFVCLQNDRRGPASFLWAAWVFSLVLGLGQGEDNRCASSNAASCARCLALGPECGWCVQEDFISGGSRSERCDIVSNLISKGCSVDSIEYPSVHVIIPTENEINTQVTPGEVSIQLRPGAEANFMLKVHPLKKYPVDLYYLVDVSASMHNNIEKLNSVGNDLSRKMAFFSRDFRLGFGSYVDKTVSPYISIHPERIHNQCSDYNLDCMPPHGYIHVLSLTENITEFEKAVHRQKISGNIDTPEGGFDAMLQAAVCESHIGWRKEAKRLLLVMTDQTSHLALDSKLAGIVVPNDGNCHLKNNVYVKSTTMEHPSLGQLSEKLIDNNINVIFAVQGKQFHWYKDLLPLLPGTIAGEIESKAANLNNLVVEAYQKLISEVKVQVENQVQGIYFNITAICPDGSRKPGMEGCRNVTSNDEVLFNVTVTMKKCDVTGGKNYAIIKPIGFNETAKIHIHRNCSCQCEDNRGPKGKCVDETFLDSKCFQCDENKCHFDEDQFSSESCKSHKDQPVCSGRGVCVCGKCSCHKIKLGKVYGKYCEKDDFSCPYHHGNLCAGHGECEAGRCQCFSGWEGDRCQCPSAAAQHCVNSKGQVCSGRGTCVCGRCECTDPRSIGRFCEHCPTCYTACKENWNCMQCLHPHNLSQAILDQCKTSCALMEQQHYVDQTSECFSSPSYLRIFFIIFIVTFLIGLLKVLIIRQVILQWNSNKIKSSSDYRVSASKKDKLILQSVCTRAVTYRREKPEEIKMDISKLNAHETFRCNF
[SEQ ID No:26]
因此,优选地,Itgβ8包含SEQ ID No:26中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,Itgav可由Genebank ID No:3685表示,其在本文中作为SEQ IDNo:27提供,如下所示:
MAFPPRRRLRLGPRGLPLLLSGLLLPLCRAFNLDVDSPAEYSGPEGSYFGFAVDFFVPSASSRMFLLVGAPKANTTQPGIVEGGQVLKCDWSSTRRCQPIEFDATGNRDYAKDDPLEFKSHQWFGASVRSKQDKILACAPLYHWRTEMKQEREPVGTCFLQDGTKTVEYAPCRSQDIDADGQGFCQGGFSIDFTKADRVLLGGPGSFYWQGQLISDQVAEIVSKYDPNVYSIKYNNQLATRTAQAIFDDSYLGYSVAVGDFNGDGIDDFVSGVPRAARTLGMVYIYDGKNMSSLYNFTGEQMAAYFGFSVAATDINGDDYADVFIGAPLFMDRGSDGKLQEVGQVSVSLQRASGDFQTTKLNGFEVFARFGSAIAPLGDLDQDGFNDIAIAAPYGGEDKKGIVYIFNGRSTGLNAVPSQILEGQWAARSMPPSFGYSMKGATDIDKNGYPDLIVGAFGVDRAILYRARPVITVNAGLEVYPSILNQDNKTCSLPGTALKVSCFNVRFCLKADGKGVLPRKLNFQVELLLDKLKQKGAIRRALFLYSRSPSHSKNMTISRGGLMQCEELIAYLRDESEFRDKLTPITIFMEYRLDYRTAADTTGLQPILNQFTPANISRQAHILLDCGEDNVCKPKLEVSVDSDQKKIYIGDDNPLTLIVKAQNQGEGAYEAELIVSIPLQADFIGVVRNNEALARLSCAFKTENQTRQVVCDLGNPMKAGTQLLAGLRFSVHQQSEMDTSVKFDLQIQSSNLFDKVSPVVSHKVDLAVLAAVEIRGVSSPDHVFLPIPNWEHKENPETEEDVGPVVQHIYELRNNGPSSFSKAMLHLQWPYKYNNNTLLYILHYDIDGPMNCTSDMEINPLRIKISSLQTTEKNDTVAGQGERDHLITKRDLALSEGDIHTLGCGVAQCLKIVCQVGRLDRGKSAILYVKSLLWTETFMNKENQNHSYSLKSSASFNVIEFPYKNLPIEDITNSTLVTTNVTWGIQPAPMPVPVWVIILAVLAGLLLLAVLVFVMYRMGFFKRVRPPQEEQEREQLQPHENGEGNSET
[SEQ ID No:27]
因此,优选地,Itgav包含SEQ ID No:27中基本列出的序列、其片段或变体,或由其组成。
在一个实施例中,1型调节性T细胞(优选为1型调节性T细胞)可被激活,优选为被抗CD3分子(例如抗CD3抗体)和/或IL-2激活。
在调节性T细胞(优选为1型调节性T细胞)中,αvβ8的表达可以用双歧杆菌素增强。因此,该方法可进一步包括使调节性T细胞(优选为1型调节性T细胞)与双歧杆菌素接触。
该方法可进一步包括用树突细胞培养淋巴细胞。
优选地,淋巴细胞在100E+03细胞/cm2和2000E+03细胞/cm2之间培养。更优选地,该淋巴细胞在250E+03细胞/cm2和1000E+03细胞/cm2之间培养。最优选地,淋巴细胞在250E+03细胞/cm2和1000E+03细胞/cm2之间培养。
在一个实施例中,该方法还包括从培养物中纯化TRM细胞。
本领域技术人员能够理解,本文所述的任何因子,例如细胞因子,可以是哺乳动物的。哺乳动物可以是啮齿动物、狗、马或猪。啮齿动物可以是大鼠或小鼠。然而,优选地,本文描述的因子是人类的因子。
在第二方面,提供了一种扩展组织驻留记忆T细胞(TRM)群体的方法,该方法包括培养如第一方面所定义的组织驻留记忆T细胞群体。
组织驻留记忆T细胞可以如第一方面所定义。
第一方面或第二方面的方法可以进一步包括在IL-2、IL-4、IL-7、IL-12、IL-15和/或IL-21的存在下培养组织驻留记忆T细胞。
优选地,IL-2是哺乳动物的。最优选地,IL-2是人类的IL-2。在一个实施例中,IL-2可由Genebank ID No:3558表示,其在本文中作为SEQ ID No:17提供,如下所示:
MYRMQLLSCIALSLALVTNSAPTSSSTKKTQLQLEHLLLDLQMILNGINNYKNPKLTRMLTFKFYMPKKATELKHLQCLEEELKPLEEVLNLAQSKNFHLRPRDLISNINVIVLELKGSETTFMCEYADETATIVEFLNRWITFCQSIISTLT
[SEQ ID No:17]
因此,优选地,IL-2包含SEQ ID No:17中基本列出的序列、其片段或变体,或由其组成。
优选地,IL-2可以0.1ng/ml和200ng/ml之间的浓度存在。更优选地,IL-2可以2ng/ml和100ng/ml之间的浓度存在。最优选地,IL-2可以10ng/ml和50ng/ml之间的浓度存在。
优选地,IL-21是哺乳动物的。最优选地,IL-21是人类的IL-21。在一个实施例中,IL-21可以由Genebank ID No:59067表示,其在本文中作为SEQ ID No:15提供,如下所示:
MRSSPGNMERIVICLMVIFLGTLVHKSSSQGQDRHMIRMRQLIDIVDQLKNYVNDLVPEFLPAPEDVETNCEWSAFSCFQKAQLKSANTGNNERIINVSIKKLKRKPPSTNAGRRQKHRLTCPSCDSYEKKPPKEFLERFKSLLQKMIHQHLSSRTHGSEDS
[SEQ ID No:15]
因此,优选地,IL-21包含SEQ ID No:15中基本列出的序列、其片段或变体,或由其组成。
优选地,IL-21可以0.1ng/ml和200ng/ml之间的浓度存在。更优选地,IL-21可以5ng/ml和100ng/ml之间的浓度存在。最优选地,IL-21可以10ng/ml和50ng/ml之间的浓度存在。
在第三方面,提供了一种通过第一方面的方法获得的或可获得的组织驻留记忆T细胞。
由本发明方法产生的组织驻留记忆T细胞在治疗应用中特别有用。
T细胞已被证明是根除肿瘤的有力工具。已知肿瘤组织中T细胞的存在,特别是组织驻留记忆T细胞的存在与阳性癌症预后94相关。一些研究特别将实体肿瘤中TRM细胞(通过其CD103的表达确定)的存在与高存活率和总体阳性预后联系起来,即使是在癌症晚期95 , 96。而且,TRM细胞在与其他已知免疫疗法97的联合治疗中已显示出能够显著提高癌症患者的存活率。然而,与其他类型的T细胞相比,天然存在的TRM细胞的数量通常较低,难以观察到显著的治疗效果。因此,在体外产生具有TRM细胞属性及其迁移和组织归巢属性的T细胞,例如具有CD103、CD69和CTLA-4的表达的T细胞,这将使得该T细胞能够穿透肿瘤,将之用于单一或联合治疗将是非常有价值的。
因此,优选地,本发明的TRM细胞具有CD103的表达。优选地,本发明的TRM细胞具有CD69的表达。优选地,本发明的TRM细胞具有CTLA-4的表达。
相应地,在本发明的第四方面,提供了根据第三方面的组织驻留记忆T细胞,可选地其扩展的群体,在治疗方面的应用。
在本发明的第五方面,提供了一种根据第三方面的组织驻留记忆T细胞,可选地其扩展的群体,在T细胞疗法的应用。
应当理解,组织驻留记忆T细胞(TRM)的扩展群体在治疗中特别有用,尤其是在T细胞疗法中。
T细胞疗法可以是CAR-T细胞疗法。T细胞疗法可以是先天性T细胞疗法,例如基于γδT细胞、黏膜相关的不变T细胞或是杀伤T细胞的疗法。
在本发明的第六方面,提供了根据第三方面所述的组织驻留记忆T细胞、可选地其扩展的群体,在预防、治疗或改善癌症或感染的应用。
在本发明的第七方面,提供了一种治疗受试者的癌症或感染的方法,该方法包括向需要这样治疗的受试者施用、或已经施用治疗有效量的根据第三方面所述的组织驻留记忆T细胞,可选地其扩展的群体。
TRM细胞可通过在抗原呈递细胞、白细胞介素(IL)-15和TGFβ存在下体外培养先前激活的T细胞而产生。优选地,添加IL-2或特别是IL-7来增强TRM细胞的迁移特性,例如分化簇(CD)69、CD103和细胞毒性T淋巴细胞相关蛋白4(CTLA-4)的表达,在小鼠模型中过继移植后,还增强了外周器官的细胞恢复。
应当理解,根据本发明产生的组织驻留记忆T细胞可以用于单一疗法(即,仅使用(i)组织驻留记忆T细胞或(ii)包含组织驻留记忆T细胞的治疗组合物)。或者,根据本发明的组织驻留记忆T细胞可以用作治疗、改善或预防疾病(例如癌症)的已知疗法的辅助剂或与之结合。
根据本发明的组织驻留记忆T细胞可以组合成具有多种不同形式的组合物,具体取决于组合物的使用方式。因此,例如,组合物可以是粉末、片剂、胶囊、液体、软膏、乳膏、凝胶、水凝胶、气雾剂、喷雾、胶束溶液、透皮贴剂、脂质体悬浮液或任何其他合适的形式,其可以施用给需要治疗的人或动物。应当理解,根据本发明的药物载体应当是被给予药物的受试者耐受良好的载体。
本发明的组织驻留记忆T细胞可以多种方式使用。例如,可能需要口服给药,在这种情况下,药物可以包含在组合物中,例如可以片剂、胶囊或液体的形式口服摄入。本发明的抗生素组合物和制剂可以通过吸入(例如鼻内)给药。组合物也可配制用于局部使用。例如,配制为面霜或软膏,可以涂抹在皮肤上。
根据本发明的组织驻留记忆T细胞组合物和制剂也可并入缓释或延迟释放装置中。例如,这种装置可以在特定组织位置插入皮肤上或皮肤下,药物可以在数小时、数天、数周甚至数月内释放。该装置可以至少位于治疗部位附近。当需要用根据本发明使用的药剂进行长期治疗并且通常需要频繁给药(例如至少每天给药)时,这种装置可能是特别有利的。
在一个优选的实施例中,根据本发明的组织驻留记忆T细胞可以通过注射到血流中或直接注射到需要治疗的部位来给药给受试者。注射可以是静脉注射(推注或输液)或皮下注射(推或输液),或皮内注射(推剂或输液)或者肌肉注射。优选地,本发明的组织驻留记忆T细胞通过外周血给药。优选地,静脉注射本发明的组织驻留记忆T细胞。
应当理解,所需的组织驻留记忆T细胞的量由其生物活性和生物利用度决定,而生物利用度又取决于给药模式、组织驻留记忆T细胞的物理化学性质,以及它们是用作单一疗法还是用于联合疗法。给药频率还将受到受试者体内组织驻留记忆T细胞半衰期的影响。最佳给药剂量可由本领域技术人员确定,并且将随所使用的特定组合物和制剂、药物组合物的强度、给药方式和待治疗的特定疾病的进展而变化。取决于所治疗的特定受试者的其他因素将导致需要调整剂量,包括受试者年龄、体重、性别、饮食和给药时间。
通常,根据使用的组合物或制剂,可以使用0.001μg/kg体重和10mg/kg体重之间的本发明TRM细胞或制剂的日剂量。更优选地,日剂量在0.01g/kg体重和1mg/kg体重之间,更优选地,在0.1g/kg和100g/kg体重之间,最优选地,在约0.1g/kg和10g/kg体重之间。通常,可以使用本发明的10^5TRM细胞和10^7TRM细胞之间的日剂量。优选地,可以使用本发明的10^5TRM细胞和10^6TRM细胞之间的日剂量。
组合物或制剂可在待治疗疾病发作之前、期间或之后施用。每日剂量可以单次给药(例如,每日单次注射)。或者,组织驻留记忆T细胞可能需要在一天内给药两次或更多次。例如,组织驻留记忆T细胞可以每天两次(或更多次,取决于正在治疗的疾病的严重程度)给药,剂量介于10^5TRM细胞和10^7TRM细胞之间(例如,假设体重为70kg)。接受治疗的患者可以在醒来时服用第一剂,然后在晚上服用第二剂(如果是两剂方案),或者此后每隔3或4小时服用一次。或者,可以使用缓释装置来向患者提供根据本发明的最佳浓度剂量,而无需重复给药。
已知的程序,例如制药工业常规使用的那些程序(例如,体内实验、临床试验等),可用于形成根据本发明的特定制剂和精确的治疗方案(例如,组织驻留记忆T细胞的每日剂量和给药频率)。
在第八方面,提供了一种药物组合物,其包含根据第三方面所述的组织驻留记忆T细胞、可选地其扩展的群体以及药学上可接受的赋形剂。
在本发明的第九方面,还提供了一种用于制备第八方面所述的药物组合物的方法,该方法包括将治疗有效量的第三方面所述的组织驻留记忆T细胞、可选地其扩展的群体与药学上可接受的赋形剂进行组合。
“受试者”可以是脊椎动物、哺乳动物或家畜。因此,根据本发明的药物可用于治疗任何哺乳动物,例如牲畜(例如马)、宠物,或可用于其他兽医应用。最优选的是,受试者是人类。
组织驻留记忆T细胞的“治疗有效量”是指当给药给受试者时,产生所需效果所需的量。
药剂的量可以是约10^5TRM细胞至约10^7TRM细胞,最优选地,为约10^5TRM细胞到约10^6TRM细胞。
本文所指的“药学上可接受的载体”是本领域技术人员已知的任何已知化合物或已知化合物的组合,其可用于配制药物组合物,特别是用于基于T细胞的治疗的制剂。
在一个实施例中,药学上可接受的载体可以是固体,组合物可以是粉末或片剂的形式。固体的、药学上可接受的载体可以包括一种或多种物质,这些物质也可以用作调味剂、润滑剂、增溶剂、悬浮剂、染料、填料、胶质剂、压缩助剂、惰性粘合剂、甜味剂、防腐剂、染料或片剂崩解剂。载体也可以是封装材料。在粉末中,载体是与本发明的细碎活性剂混合的细碎固体。在片剂中,活性剂可与具有必要压缩特性的载体以适当比例混合,并压实成所需的形状和尺寸。粉末和片剂优选地含有高达99%的活性剂。合适的固体载体包括例如磷酸钙、硬脂酸镁、滑石、糖、乳糖、糊精、淀粉、明胶、纤维素、聚乙烯吡咯烷、低熔点蜡和离子交换树脂。在另一个实施例中,药物载体可以是凝胶,组合物可以是乳膏或类似物的形式。
然而,药物载体可以是液体,并且药物组合物是溶液形式。液体载体用于制备溶液、悬浮液、乳液、糖浆、酏剂和加压组合物。本发明的活性剂可以溶解或悬浮在药学上可接受的液体载体中,例如水、有机溶剂、两者的混合物或药学上可以接受的油或脂肪。液体载体可以含有其他合适的药物添加剂,例如增溶剂、乳化剂、缓冲剂、防腐剂、甜味剂、调味剂、悬浮剂、增稠剂、着色剂、粘度调节剂、稳定剂或渗透调节剂。用于口服和肠外给药的液体载体的合适实例包括水(部分含有如上所述的添加剂,例如纤维素衍生物,优选羧甲基纤维素钠溶液)、醇(包括一元醇和多元醇,例如二醇)及其衍生物,以及油(例如分级椰子油和花生油)。对于肠胃外给药,载体也可以是油酸乙酯和肉豆蔻酸异丙酯等油性酯。无菌液体载体可用于肠胃外给药的无菌液体形式组合物。加压组合物的液体载体可以是卤代烃或其他药学上可接受的推进剂。
液体药物组合物是无菌溶液或悬浮液,可通过例如肌肉内、鞘内、硬膜外、腹腔内、静脉内和特别是皮下注射来使用。药剂可制备为无菌固体组合物,其可在施用时使用无菌水、盐水或其他合适的无菌注射介质溶解或悬浮。
本发明的药剂和组合物可以以含有其他溶质或悬浮剂(例如,足够的盐或葡萄糖使溶液等渗)、胆汁盐、阿拉伯胶、明胶、山梨醇单酯、聚山梨醇酯80(山梨醇及其酸酐与环氧乙烷共聚的油酸酯)等的无菌溶液或悬浮液的形式口服给药。根据本发明使用的药剂也可以液体或固体组合物形式口服给药。适用于口服给药的组合物包括固体形式,如药丸、胶囊、颗粒、片剂和粉末,以及液体形式,如溶液、糖浆、酏剂和悬浮液。用于肠胃外给药的形式包括无菌溶液、乳剂和悬浮液。
应当理解,本发明延伸至任何核酸或肽或其变体、衍生物或类似物,其基本上包含本文所述的任何序列的氨基酸或核酸序列,包括其变体或片段。术语“基本上是氨基酸/核苷酸/肽序列”、“变体”和“片段”可以是与本文所述序列中任一序列的氨基酸/核苷酸或肽序列具有至少40%序列同一性的序列,例如与识别为SEQ ID Nos:1-28的序列具有40%同一性,以此类推。
本发明还设想了序列同一性大于65%、更优选地大于70%、进一步更优选地大于75%、进一步更优选地大于80%的氨基酸/多核苷酸/多肽序列。优选地,该氨基酸/多核苷酸/多肽序列与所述序列中的任一序列具有至少85%的序列同一性,更优选地具有至少90%的序列同一性,进一步更优选地具有至少92%的序列同一性,进一步更优选地具有至少95%的序列同一性,进一步更优选地具有至少97%的序列同一性,最优选地与本文所述的任何序列具有至少99%的序列同一性。
本领域技术人员能够理解如何计算两个氨基酸/多核苷酸/多肽序列之间的百分比同一性。为了计算两个氨基酸/多核苷酸/多肽序列之间的百分比同一性,必须首先准备两个序列的比对,然后计算序列同一性值。两个序列的百分比同一性可以取不同的值,具体取决于:-(i)用于比对序列的方法,例如ClustalW、BLAST、FASTA、Smith-Waterman(在不同的程序中实现),或由3D比较来进行结构对齐;以及(ii)比对方法所使用的参数,例如,局部与全局比对,所使用的对得分矩阵(例如BLOSUM62、PAM250、Gonnet等),以及间隙惩罚,例如函数形式和常数。
进行比对后,有许多不同的方法计算两个序列之间的百分比同一性。例如,可以将同一性值除以:(i)最短序列的长度;(ii)比对的长度;(iii)序列的平均长度;(iv)非间隙位置的数量;或(v)不包括悬突的等效位置的数量。此外,可以理解,百分比同一性也强烈依赖于序列长度。因此,一对序列越短,就有越高概率碰巧其具有序列同一性。
因此,应当理解,蛋白质或DNA序列的精确比对是一个复杂的过程。流行的多重比对程序ClustalW(Thompson等人,1994,Nucleic Acids Research,224673-4680;
Thompsson等人,1997,Nucleical Acids Research,244876-4882)是根据本发明产生蛋白质或DNA多重比对的优选方式。ClustalW的合适参数如下:对于DNA比对:Gap OpenPenalty=15.0,Gap Extension Penalty=6.66,and Matrix=Identity。对于蛋白质比对:Gap Open Penalty=10.0,Gap Extension Penalty=0.2,and Matrix=Gonnet。对于DNA和蛋白质比对:ENDGAP=-1,GAPDIST=4。本领域技术人员能够意识到,可能需要改变这些参数和其他参数以实现最佳序列比对。
优选地,两个氨基酸/多核苷酸/多肽序列之间的同一性百分比的计算可以根据(N/T)*100这样的比对来计算,其中N是序列共享相同残基的位置的数目,T是比较的位置的总数,包括间隙和包括或不包括悬突。优选地,悬突包括在计算中。因此,用于计算两个序列之间的百分比同一性的最优选方法包括:(i)使用ClustalW程序,使用合适的参数集,例如,如上所述,准备序列比对;以及(ii)将N和T的值插入以下公式:-序列同一性=(N/T)*100。
本领域技术人员已知鉴定相似序列的替代方法。例如,基本相似的核苷酸序列将由在严格条件下与DNA序列或其互补序列杂交的序列编码。上述严格条件指的是在大约45℃下,核苷酸杂交以在3倍氯化钠/柠檬酸钠(SSC)中过滤结合的DNA或RNA,然后在大约20-65℃的0.2x SSC/0.1%SDS中至少洗涤一次。或者,基本上相似的多肽可以与例如SEQ IDNo:1-28中所示的氨基酸序列中的序列相差至少1个,但少于5、10、20、50或100个氨基酸。
由于遗传密码的简并性,很明显,本文描述的任何核酸序列都可以改变或改变,而基本上不影响由此编码的蛋白质的序列,以提供其功能变体。合适的核苷酸变体是那些序列被编码序列内相同氨基酸的不同密码子取代而改变,从而产生沉默(同义)变化的核苷酸变体。其他合适的变体是那些具有同源核苷酸序列但包含序列的全部或部分的变体,这些变体通过编码氨基酸的不同密码子的替换而改变,所述氨基酸具有与其替代的氨基酸相似的生物物理性质的侧链,以产生保守变化。例如,小的非极性疏水氨基酸包括甘氨酸、丙氨酸、亮氨酸、异亮氨酸、缬氨酸、脯氨酸和蛋氨酸。大型非极性疏水氨基酸包括苯丙氨酸、色氨酸和酪氨酸。极性中性氨基酸包括丝氨酸、苏氨酸、半胱氨酸、天冬酰胺和谷氨酰胺。带正电的(碱性)氨基酸包括赖氨酸、精氨酸和组氨酸。带负电荷的(酸性)氨基酸包括天冬氨酸和谷氨酸。因此,应当理解哪些氨基酸可以被具有类似生物物理性质的氨基酸取代,并且本领域技术人员知道编码这些氨基酸的核苷酸序列。
本文描述的所有特征(包括任何附带的权利要求、摘要和附图)和/或如此公开的任何方法或过程的所有步骤可以以任何组合与上述任何方面组合,但这些特征和/或步骤中的至少一些相互排斥的组合除外。
附图说明
图1显示了Foxp3依赖性的Tbx21切除导致1型Treg细胞数量减少。
(a)通过流式细胞术分析(n=4-7)染色T-bet的肠道TRM或脾脏CD8+T细胞的百分比,对Foxp3WT和Foxp3ΔTbx21小鼠体内的T细胞群进行体外流式细胞术分析。
(b,f)(a)Foxp3WT或(f)Foxp3ΔTbx21小鼠(n=4-12)中表达CXCR3、CCR6或ST2的脾脏、肠系膜淋巴结(mLN)或固有层(LPL)中Treg细胞(CD4+Foxp3+)的比例。
(c,d)CXCR3在CD4+、CD8α+中表达的百分比以及(c)脾脏或(d)胸腺中的Treg细胞(n=5)。
(e)脾脏CD4+T细胞中CXCR3表达的流式细胞术代表图。
(g-j)Foxp3WT(空心圆形)和Foxp3ΔTbx21小鼠(实心三角形)中(g)CD44、(h)Helios、(i)Nrpl1或(j)KLRG1表达的Treg细胞比例(n=4-9)。条形表示平均值,星号条形表示±SEM;为了进行统计分析,使用了Mann-Whitney U检验或多重t检验(g-j);*P<0.05;**P<0.01;***P<0.001;****P<0.0001。
图2显示了Foxp3依赖性Tbx21切除导致CD8+T细胞521群体的改变。
(a,b)脾脏中CD8+T细胞群的流式细胞术分析。
(a)Foxp3WT(空心条)n=8、Foxp3ΔTbx21(实心条)n=4、Foxp2ΔEomes小鼠(灰色条)n=14的脾脏中的幼稚(CD62LhiCD44lo)、中央记忆(CD62LhiCD44hi)和效应记忆(CD62L-CD44hi)CD8+T细胞的百分比;
(b)所示小鼠品系脾脏CD8+T细胞中CD62L和CD44表达的代表性流式细胞术图。
(c-e)肠隔室中T细胞群、CD4+、Foxp3+和CD8α+T细胞的流式细胞术分析。
(c)Foxp3WT(空心符号)或Foxp3ΔTbx21(实心符号)小鼠的上皮内淋巴细胞(IEL);
(d)(c)中描述的相同小鼠的IEL指示亚群数量;
(e)固有层(LPL)(n=8-9)。
(f,g)Foxp3WT和(f)Foxp3ΔTbx21或(g)Foxp3ΔEomes(n=4-12)中指定器官中CD4+Foxp3-和CD8+T细胞的比例。
(h)所示小鼠品系的IEL(上图)或LPL(下图)CD8+T细胞的CD103和CD69表达的代表性点图。
(i)Foxp3WT(空心条),Foxp3ΔTbx21(实心条)和Foxp3ΔEomes(灰色条)小鼠品系(n=5-11)的空肠LPL中总CD8+和CD8+CD103+T细胞的细胞数目。
(j)在所示小鼠品系的LPL中发现的总CD8+T细胞与CD8+CD103+T细胞的比率。
(n=6-13),条形表示平均值,星号条形表示±SEM,对于统计分析,使用多重t检验;*P<0.05;***P<0.01;***P<0.001;****P<0.0001。
图3显示了在缺失1型Treg细胞的情况下TRM细胞的发育减少。
(a-d)从Foxp3WT(空心条),Foxp3ΔTbx21(实心条)和Foxp3ΔEomes(灰色条)小鼠的指定小肠切片分离固有层淋巴细胞,并通过流式细胞术分析。
(a,b)细胞在TCRβ被门控并被检测KLRG1表达(n=4-8)。
(c,d)显示指示的小鼠品系的TCRβ+CD4-CD8α+LPLs中的(c)CD103和KLRG1的表达或(d)Eomes和KLRG1的表达的代表性流式细胞术图。
(e-h)在指定小鼠品系(f)的肝脏和肺中具有KLRG1表达的TCRβ+CD4-CD8α+细胞的比例的代表性图(e)和累积数据(f-h),(g)肝脏(CD69+Eomes-KLRG1-)和(h)肺(CD69+CD103+KLRG1-)(n=7-11)中TRM细胞的比例;条形表示平均值,星号条形表示±SEM,对于统计分析,使用Mann-Whitney U检验或多重t检验(f-h);*P<0.05;***P<0.01;***P<0.001;****P<0.0001。
图4显示了TRM细胞发育减少导致对感染的易感性增加。
(a-b)通过流式细胞术分析来自Foxp3ΔTbx21小鼠的TCRβ+CD8α+固有层淋巴细胞Eomeshi或Eomeslo,对于(a)PD-1和(b)GrzmB表达(n=8-9),在用25μg抗CD3i.p.刺激后48小时进行分析。
(c-h)Foxp3WT(空心符号),(c,e,g)Foxp3ΔTbx21(实心符号)或(d,f,h)Foxp3ΔEomes(灰色符号)小鼠用1000只蠕形艾美球虫的卵囊口服感染。
(c,d)第5-18天从个体小鼠粪便收集的卵囊的累积数量。
(e,f)每只小鼠每天脱落的卵囊数量。
(g,h)蠕形艾美球虫感染过程中的体重变化(每个实验n=3-4,2个生物学重复);条形表示平均值,星号条形表示±SEM。对于统计分析,使用多重t检验或Wilcoxon检验(a,b);*P<0.05;***P<0.01;***P<0.001。
图5显示了1型Treg细胞的募集决定了TRM细胞分化。
(a-b)用来自Ctrl(CD45.1)或Foxp3ΔTbx21(CD45.2)小鼠的骨髓重建Rag2-缺陷型小鼠;评估每个供体对脾脏和TRM(CD8+CD103+KLRG1-)LPL群体中总CD8 T细胞的贡献。(a)代表性点图;(b)评估的个体小鼠概述(n=8)。
(c)收养转让模式的描述;Foxp3WT或Foxp3ΔTbx21小鼠在用蠕形艾美球虫口腔感染前一天静脉内注射C57Bl/6CD45.1+CD8α+T细胞;感染消退后和接种后3-4周,从小肠分离固有层淋巴细胞(LPL)并通过流式细胞术分析。
(d)TRM细胞发育占CD45.1宿主小鼠LPL中总CD8 T细胞的比例,如(c)所述用脾Foxp3ΔTbx21衍生的CD8 T细胞转移并随后进行蠕形艾美球虫攻击(n=6)。
(e)在CD45.1上预先门控的淋巴细胞的代表性点图,以及在TCRβ和CD8α上门控并分析指定小鼠品系中CD103和Eomes的表达。
(f-g)感染后(b)第3周或(c)第9周从小肠获得的总CD8α+LPL和CD103+Eomes-CD8α+LPL的细胞数;如(a)中所述进行门控(n=5-7)。
(h)在将具有或不具有野生型Treg细胞的野生型CD45.1+CD8α+T细胞转移到Foxp3ΔTbx21小鼠后,以及在蠕形艾美球虫攻击后,回收表达CD103的CD8α+CD45.1+T细胞的比例。
(i)Treg细胞亚型的单细胞RNA测序分析,分为1型(85个细胞),2型(35个细胞),3型(28个细胞)或其他未定义的Treg细胞组织;条形表示平均值,星号条形表示±SEM;对于统计分析,使用Mann-Whitney U检验;**P<0.01;****P<0.0001,n.s.=无显著性。
图6显示了1型Treg细胞通过TCRβ的可用性促进TRM细胞发育。
(a,b)Foxp3WT或Foxp3ΔTbx21小鼠在口服感染假结核耶尔森氏菌前一天静脉内接受C57Bl/6CD45.1+CD8α+T细胞;2-3周后,从小肠分离固有层淋巴细胞(LPL)并通过流式细胞术分析。
(a)恢复表达CD103的CD8α+CD45.1+T细胞的比例(n=7-9)。
(b)在CD45.1上预门控的淋巴细胞的代表性点图,以及在TCRβ和CD8α上门控并分析指定小鼠品系中CD103和Eomes的表达。
(c-d)回肠LPL Foxp3WT(空心符号)或Foxp3ΔTbx21(实心符号)小鼠在稳定状态(圆形)或蠕形艾美球虫(Ev)(正方形)感染后10天分析(c)Treg数量、(d)表达CXCR3的Treg数量(n=4-9)。
(e)用蠕形艾美球虫感染或不感染C57BL/6小鼠,并在第10天在回肠中评估超过Hprt的Cxcl10 mRNA水平(2594个生物学重复n=5-8)。
(f-k)Foxp3ΔTbx21小鼠在口腔感染蠕形艾美球虫前一天静脉内接受C57Bl/6CD45.1+CD8α+T细胞;在感染后第3周,从小肠分离LPL并通过流式细胞术分析CD103的表达。除CD8CD45.1细胞外,共转移缺失(f)CXCR3、(h)IL-10、(i)IL-35、(j)整联蛋白8或(k)TGFβ1的野生型Treg细胞或Treg细胞;在面板中累积并使用接受野生型TREG细胞的小鼠(f,h-k)。
(g)所示的小鼠品系在用蠕形艾美球虫口服感染前一天静脉内接受CD45.1+CD8α+T细胞;感染后第10天,对小肠进行CD45.1(绿色),Dapi(蓝色)和Foxp3(红色)染色。从卵囊区域(n=4)显示的代表性免疫组织化学图片(物镜40倍,变焦1.5倍);白色箭头表示CD8 T细胞和TREG紧密接近;条形表示平均值,星号条形表示±SEM;对于统计分析,使用Mann-Whitney U检验;*P<0.05;***P<0.01;***P<0.001;****P<0.0001。
图7显示了Foxp3依赖性Tbx21有条件地缺失导致体外流式细胞分析中1型Treg细胞数量减少。
a)通过C57BL/6脾脏或肠道TRM群中CD8细胞的细胞内染色显示Tbet和Eomes表达的代表性图。
b)针对Foxp3和Tbet染色的CD4 T细胞的代表性图。
c-e)Foxp3WT(空心条)和Foxp3ΔTbx21(实心条)小鼠n=9的胸腺和脾脏中Treg、CD4和CD8 T细胞的数量。
f-h)Foxp3WT(空心条)和Foxp3ΔEomes(灰色条)小鼠中Treg、CD4 T细胞和CD8 T细胞的数量。
i-j)在i)脾或j)胸腺中CD4 CD8α和Treg细胞中CXCR3表达的百分比;通过流式细胞术分析脾脏、肠系膜淋巴结的mLN和固定层中Treg细胞的比例,其具有CXCR3,CCR6或ST2的表达(n≥4,空心符号为Foxp3ΔWT,实心符号为Foxp3ΔTbx21,星号条形代表±SEM,对于统计分析,采用Mann-Whitney U检验或多重t检验,h-m)P<0 05 P<0 01 P<0 001 P<00001。
图8显示了Tregs中的Tbet或Eomes缺陷与小肠中T细胞表型的改变相关。从Foxp3WT、Foxp3ΔTbx21和Foxp3ΔEomes小鼠中分离出固有层淋巴细胞(LPL)并通过流式细胞术进行分析。
a-d)(a)十二指肠(n=8-11)、(b)空肠(n=10-14)、(c)回肠(n=9-13)和(d)结肠(n=4-7)中LPL部分的细胞数,通过对TCRβ+细胞进行门控,然后对CD4+CD8α-YFP-(CD4+)、CD4+CD8α-YFP+(Treg)和CD4-CD8α+T细胞(CD8+)进行门控来确定。
e-f)来自Foxp3WT(空心圆圈)和e)Foxp3ΔTbx21(实心三角形)或(f)Foxp3ΔEomes(实心菱形)(n=8)的个体动物中CD4+Foxp3-T细胞和CD4+YFP+T细胞之间的比率。g)代表性的流式细胞术点图,显示了小鼠品系的CD4+YFP-(上图)或CD8+(下图)LPL细胞的CD103和CD69染色。
h)Foxp3WT(空心条),Foxp3ΔTbx21(实心条)和Foxp3ΔEomes(灰色条)小鼠品系(n=5-11)的固有层中总CD4+T细胞和CD103+CD4+T细胞的数量。
i)在所示小鼠品系(n=6-13)的LPL中发现的Foxp3-CD4+T细胞/CD8+CD103+T细胞的比例;星号条形代表±SEM;对于统计分析,使用了Mann-Whitney U检验;*P<0.05;***P<0.01;***P<0.001;****P<0.0001。
图9显示了缺失1型Tregs导致TRM细胞减少。从Foxp3WT、Foxp3ΔEomes和Foxp3ΔTbx 21小鼠中分离出固有层淋巴细胞,并通过流式细胞术检查。
a)代表性的流式细胞术点图,显示在指定的小鼠品系中CD103和Eomes的对比、KLRG1和CD69的对比。
b)代表性的流式细胞术点图,显示KLRG1和Bcl2蛋白表达的对比。
c)代表性流式细胞术点图,显示CD103和TL四聚体(CD8αα染色)(n=4)的对比。
d)代表性的流式细胞术点图,显示血液和LPL区室中CD8α的体内染色和e概述图(n=5)*P<0.05,**P<0.01,***P<0.001,****P<0.0001。
图10显示了TRM细胞发育减少增加感染易感性。在腹膜内注射CD3ε抗体48小时后,从Foxp3WT和Foxp3ΔTbx 21小鼠中分离出固有层淋巴细胞(LPL)或脾脏CD8 T细胞,并通过流式细胞术检查。
a-c)代表性流式细胞术点图,显示(a)PD1或(b)颗粒酶b与LPLs中Eomes蛋白表达的对比(n=4)。
b)同a),但显示脾脏的累积数据(n=6)。
d-i)口服感染1000蠕形艾美球虫卵囊的(g-i)IL 15R小鼠的C57BL/6(对照)或(d-f)C57BL/6Rag2-/-,(g)对每只小鼠每日粪便中卵囊计数,持续指定天数。
e,h)累积卵囊计数(第6-17天)。
f,i)蠕形艾美球虫感染期间体重的变化(n=5-9,2个生物学重复),星号条形代表±SEM,对于统计分析,采用Mann-Whitney U检验(e,h)、多重t检验(d,f,g,f)或Wilcoxon检验(b);*P<0.05,**P<0.01,***P<0.001,****P<0.0001。
图11显示了免疫激活后Tbet表达和TRM细胞发育。(a-d)在稳定状态或体内抗CD3刺激后24小时后,通过细胞内流式细胞术染色评估指示的T细胞。
a)Foxp3WT(空心符号)和Foxp3ΔTbx21(实心符号)的脾脏的CD4和CD8 T细胞中Tbet表达的概述。
b)通过细胞内染色分析脾脏CD8幼稚及记忆T细胞以及LPL来源的TRM细胞(n=1112)的Tbet表达。来自Foxp3WT和Foxp3ΔEomes小鼠脾脏的CD8 T细胞在活化后24小时(n=35)对(c)Eomes或(d)Tbet进行染色的代表性图。
e-h)Foxp3WT(空心符号)和Foxp3ΔTbx21(实心符号)小鼠过继转移CD8 CD45.1 T细胞,在此基础上一天后用蠕形艾美球虫攻击小鼠。在感染高峰时,即第10天,分离LPL并通过流式细胞术分析CD45.1以得到(e)TRM细胞(CD8+CD103+Eomes-)的比例,以及(f)总CD8+CD45.1+T细胞(n=6 7)的比例。(g)如e)所述,使用0D45.2+Foxp3ΔTbx21小鼠作为宿主,向其转移了另外的CD45.1+Wt Treg细胞,显示LPL中CD4+T细胞、内源性(CD45.2+)和转移的(CD45.1+)CD4和表达CXCR3的Treg细胞的代表性流式细胞术图。
h)同f-g)转移的Treg细胞的CXCR3比例的概述;对于统计分析,使用了Mann-Whitney U检验。
图12显示了选定小鼠品系中的TRM细胞发育。
a,b)在Foxp3WT和Foxp3ΔTbx21小鼠处于稳定状态(圆形)或感染蠕形艾美球虫10天后(方形)(n=5-8),从这些小鼠中分离并计数回肠固有层CD4。
c)指示的小鼠品系在口服Ev感染前一天静脉内接受CD45.1+CD8α+T细胞;在感染后第10天,小肠染色CD45.1(绿色)Dapi(蓝色)和Foxp3(红色);显示代表性的免疫组织化学图片(目标20倍、变焦15倍,显示比例尺50微米,指定区域放大)卵囊是特征性的圆形未染色区域,例如以*标示出的部分,并显示了一个无卵囊区域用于对比(第三面板)。
f)脾脏的CXCR3或CXCR3+Treg细胞的相对表达水平(n=4)。LPL分离自d,e)C57BL6(对照)或IL 10缺陷小鼠、g,j)或骨髓;用C57BL6小鼠或者g,h)IL-35缺陷小鼠、g,i)Foxp3ΔItgβ8、g,j)Foxp3Δtgfβ1小鼠产生嵌合小鼠。显示了代表性的流式细胞术图(f)和表达CD103的细胞的累积比例(h-j)(n=7-10)。条形代表平均值;对于统计分析,使用Mann-Whitney检验***P<0.001,****P<0.0001。
图13显示了流式细胞术分析门控策略。
a)来自C57BL/6Foxp3ΔTbx21小鼠脾脏的代表性流式细胞术图,显示淋巴细胞门控、双重排斥和死细胞排斥,以及基于eYFP和tdRFP检测的CD4选择和Treg选择。
b)来自C57BL/6Foxp3ΔTbx21小鼠的LPL的代表性流式细胞术图,显示淋巴细胞门控、双重排斥和死细胞排斥,以及包含的TCRβ+CD69+和CD8α+;在LPL中,TRM被定义为CD103+。
图14显示了Foxp3ΔTbx21小鼠中的RFP检测。
a-c)来自如图13A所示的C57BL/6Foxp3ΔTbx21小鼠脾脏的代表性流式细胞术图,显示a)在CD4生命门中的eYFP和tdRFP检测,b)在CD4 Foxp3-YFP群体中,以及c)在CD8群体中;显示了2只个体小鼠的两个代表性图。
图15显示了实验布局,详细描述了TRM细胞的生产和测试,从小鼠的初始细胞提取到通过FACS机器分选细胞群。
图16显示了骨髓来源的树突细胞(BMDC)在CD8T细胞中维持CD69表达。当效应CD8T细胞在IL-15和仅TGFβ存在下(圆形)或还在BMDC存在下(方形)与抗CD3一起培养时,在TRM细胞上关键表达的CD69标记在BMDC存在下仍然表达,但在BMDC不存在下丢失。
图17显示了IL-7诱导TRM中的CTLA-4表达。两个独立的重复实验显示当效应CD8 T细胞在不存在IL-7(黑色)和添加IL-7(灰色)的情况下培养时CTLA-4表达的差异。
图18显示了IL-2还诱导TRM中的CTLA-4表达。图中数据显示当效应CD8 T细胞在不存在IL-7和IL-2(黑色)、加入IL-7(浅灰)以及加入IL-2(深灰)的情况下培养时CTLA4的表达。尽管与IL-7、IL-2一起培养的情况下,有相似比例的CD 8T细胞表达CTLA-4,但对于含有IL-7的培养物,CTLA-4表达水平平均来说比含有IL-2的培养物更稳健。
图19显示了详细描述TRM细胞的产生和测试的实验布局,以及攻击后对小鼠器官进行的体内攻击和离体分析。发明人想要确认体外产生的TRM细胞的特征和表型与体内将其植入器官是否相关。
图20显示了用于体内攻击的不同T细胞培养物的标志物CD69、CD103和CTLA-4的表达谱。用于攻击的T细胞在以下条件或组下培养:
组1:CD8+BMDC+aCD3+TGFb+IL-15=TRM;
组2:CD8+BMDC+aCD3+TGFb+IL-15+IL-7=TRM+IL-7;
组3:CD8+BMDC+aCD3+IL-15=效应细胞;
组4:CD8+BMDC+aCD3+TGFb+IL-15+IL-2=TRM+IL-2。
图21显示了在40天的过程中源自被攻击小鼠的脾脏的CD8+T细胞的数量。在图20所示的条件下培养效应CD8 T细胞并转移到完整的C57BL6/J宿主中。在指定的时间,在脾脏中评估转移细胞(CD45.1)的存在。图表显示了两个实验汇集。实验1(黑色符号)不包括组4,并且仅在时间点11-14进行分析。实验2(灰色符号)包括组4,并且在几个时间点进行分析,并且仅显示6只小鼠中的2只,其中细胞在晚期时间点恢复。数据显示与不存在IL-7培养的TRM细胞相比,在IL-7存在下培养的TRM细胞(组2)被大量发现。
图22显示了在40天的过程中来自被攻击小鼠的肠固有层的CD8+T细胞的数量。在图20所示的条件下培养效应CD8 T细胞并转移到完整的C57BL6/J宿主中。在指定的时间,在肠固有层中评估转移细胞(CD45.1)的存在。图表显示了两个实验汇集。实验1(黑色符号)不包括组4,并且仅在时间点11-14进行分析。实验2(灰色符号)包括组4,并且在几个时间点进行分析,并且仅显示6只小鼠中的2只,其中细胞在晚期时间点恢复。数据显示与不存在IL-7培养的TRM细胞相比,在IL-7存在下培养的TRM细胞(组2)被大量发现。
图23显示了在32天的过程中来自被攻击小鼠的肺以及肠的IEL区室的CD8+T细胞的数量。效应CD8 T细胞在图20所示的条件下培养并转移到完整的C57BL6/J宿主中。在指定的时间,在肺以及肠的IEL区室中评估转移细胞(CD45.1)的存在。图表显示了两个器官的实验(实验1–黑色符号;实验2–灰色符号),显示了三组实验,并在多个时间点进行了分析。数据显示与不存在IL-7培养的TRM细胞相比,在IL-7存在下培养的TRM细胞(组2)被大量发现。
图24显示了在40天的过程中来自被攻击小鼠的肠的固有层以及脾的CD8+T细胞的比较数量。在图20所示的条件下培养效应CD8 T细胞并转移到完整的C57BL6/J宿主中。在指定的时间,在脾和肠固有层中评估转移细胞(CD45.1)的存在。数据显示与不存在IL-7培养的TRM细胞相比,在IL-7存在下培养的TRM细胞(组2)被大量发现。
具体实施方式
本发明人假设TREG细胞在T细胞以及TRM细胞的产生中是重要的,该T细胞能够深入组织并且对实体肿瘤非常有效。本发明人旨在确定产生TRM细胞以在体外产生TRM细胞所需的因子,最终目的是适应当前的培养方案以产生抗肿瘤T细胞,使其具有靶向原发性肿瘤的组织穿透性质,并提供针对已迁移至转移的关键器官范围的免疫监视远离原发肿瘤的组织。本发明人还旨在评估如图15中详述的在培养基中不存在TREG细胞时是否可能产生TRM细胞。此外,本发明人评估体外产生的细胞是否保持其治疗性质,特别是它们的如图19所示的在组织内体内迁移和存活的能力。
材料和方法
小鼠:
C57Bl/6J和C57Bl/6J CD45.1小鼠购自法国Charles River公司。Tbx21f/f(Tbx21tm2Srnr)和Eomesfl/fl(Eomestm1Srnr)由Reiner博士14,82提供,Foxp3eYFP-Cre(Foxp3tm4(YFP /icre)Ayr)由Rudensky博士83提供,Rosa26-tdRFP由Fehling博士84提供,Rag2-/-,IL15R-/-(Jackson实验室)。小鼠在葡萄牙里斯本的分子医学研究所饲养。使用8-18周龄的雄性和雌性小鼠,符合研究的年龄和性别。将动物饲养在具有温度控制条件的IVC笼中,处于12小时光照/黑暗循环下,饮用水和食物可自由获取。将所有小鼠保持在无特定病原体的条件下。通过PCR对Foxp3eYFP-Cre Rosa26-tdRFP系中的所有小鼠进行严格的基因分型,并丢弃其中检测到敲除等位基因的那些小鼠(~20%),通过对表达CXCR3的CD4 T细胞进行血型分析,确认存在适当的Tbx21。另外,对小鼠进行了反向筛选,选出与eYFP相关的RFP的不恰当表达(~10%丢弃)(图13-14)。通过对Rag2缺陷小鼠进行亚致死照射(450拉德)产生骨髓嵌合体,随后静脉内注射获得的骨髓细胞。CXCR3-/-(Cxcr3tm1Dgen)85在德国海德堡德国癌症研究中心(DKFZ)繁殖。IL-10-/-86在里斯本Gulbenkian deCiência研究所繁殖,Ebi3-/-87在德国美因茨大学医学中心免疫学研究所培育,与Foxp3yfp-Cre杂交的Itgb8f/f 88和Tgfb1f/f89在法国里昂癌症研究中心免疫病毒学和炎症科培育。所有动物实验均符合葡萄牙食品和兽医总局和当地伦理审查委员会和准则的规定。
细胞分离:如前所述分离肠细胞90。用PBS冲洗肠道以除去内容物并将其纵向打开。切成1cm的片后,将其在含有20mM Hepes,100U/ml青霉素,100μg/ml链霉素,1mM丙酮酸盐,10%FCS,100μg/ml多粘菌素B和10mM EDTA的PBS中在37℃下孵育30分钟,并在此过程中摇动以释放IEL。使用37.5%等渗Percoll进一步纯化IEL单细胞悬液。为了分离LPLs,将肠组织在含有0.5mg/ml胶原酶D(Roche)和0.2mg/ml DNaseI(Roche)的IMDM培养基中在37℃下摇动消化25min。将器官通过70μm过滤器捣碎来分离肝淋巴细胞,然后用37.5%等渗Percoll进行细胞纯化。用剪刀将肺切成小块,并在含有1mg/ml胶原酶D的PBS中在37℃下消化30分钟。含有淋巴细胞的细胞悬浮液通过50μm细胞过滤器后获得。
过继性细胞转移:从脾和淋巴结的单细胞悬液中纯化CD8α+T细胞和/或CD25+细胞(Treg)。简而言之,根据制造商的说明,用抗CD8a-APC或抗CD25-APC抗体标记细胞,并用抗APC MACS玻璃细珠进行选择。计数后,通过流式细胞术测定纯度并调节细胞数。为了确保转移群体中广泛的TCR多样性,至少使用2×106的CD8 T细胞。一些受体小鼠另外接受0.4-1×106的Treg细胞。细胞转移后一天(第0天)进行感染。
感染挑战:如前所述,用蠕形艾美球虫(Ev)感染动物91。简而言之,用去离子水洗涤卵囊3次,漂浮在次氯酸钠中并使用Fuchs-Rosenthal室计数。小鼠通过在100μl水中口服管饲法接受500个卵囊的大肠埃希菌,并在感染清除后(从第3周开始)进行分析。为了确定感染负荷,将动物单独笼养,每天收集粪便直至粪便中不再检测到卵囊。动物感染假结核耶尔森氏菌(Yptb),由T.Bergsbaken博士提供,如前所述58。动物通过口服强饲法在100μl水中感染106Yptb。在第15-19天进行组织分析。
流式细胞术:根据议定的标准92和指定的门控策略(图13),制备来自脾、淋巴结、肠、肺和肝的单细胞悬液并用抗体染色(参见列表)。体内染色通过静脉内注射3μg CD8a-APC抗体进行,5分钟后处死小鼠。TL四聚体由NIH四聚体核心设施提供。样品在FortessaX20细胞计数器(BD Biosciences)上运行并用FlowJo软件(TreeStar)分析。
定量RT-PCR:使用总RNA小提提取试剂盒分离RNA,并使用来自AppliedBiosystems的高容量RNA-to-cDNA试剂盒生成cDNA。使用SYBR Select预混液(AppliedBiosystems)和QuantiTect引物测定Mm_Cxcl10_1_SG,Mm_Tgfb1_1_SG,Mm_Itgb8_1_SG和Mm_Hprt_1_SG(Qiagen)进行扩增。
免疫组织化学和显微镜检查:将肠组织卷成“瑞士卷”,固定在10%福尔马林中,在30%葡萄糖中再水化并在OCT培养基中冷冻。将组织切成10μm的切片,并用4%多聚甲醛处理切片。使用10%BSA进行封闭,并使用以下抗体进行检测:CD45.1(A20,Biolegend)和FOXP3(FJK-16s,eBioscience)。将载玻片安装在Fluoromount(Invitrogen)封片剂中并使用Zeiss LSM 880显微镜成像。使用Fiji软件进行分析。
scRNA-Seq分析:生产并在55中分析原始数据。从最初的数据集中,基于Foxp3选择TREG细胞,不包括Tmems、应激和低质量细胞。为了分析这个子集,我们采用了类似于55的方法,使用R语言的工具包Seurat93。使用“LogNormalize”方法和比例因子105对数据进行归一化;并基于负二项模型和使用UMI对数据进行缩放。TREG的亚型使用以下标准定义:类型1(原始计数分配给基因Tbx21、Stat1和Cxcr3的细胞);类型2(原始计数分配给基因Gata3,Stat6和Il1rl1的细胞);类型3(原始计数分配给基因Rorc、Stat3和Ccr6的细胞);其他(细胞没有原始计数分配给基因Tbx21,Gata3和Rorc)。
体外培养:
从先前腹膜内注射25μg抗CD3ε的C57BL6/J或CD45.1C57BL6/J小鼠获得效应CD8 T细胞。通过AutoMACS珠选择分离细胞,并在IMDM培养基中以每个96孔板平底200000个细胞培养。在指定条件下添加通过标准方案使用GM-CSF培养的100000BMDC。用0.25μg/ml anti-CD3ε、10ng/ml IL-15以及0.5ng/ml TGFβ,并在10-20ng/ml IL-2或IL-7存在下再次激活细胞。细胞生长3天,然后对其进行分析,或将其过继转移到完整的C57BL6/J小鼠中以测试组织归巢。评估细胞的TRM标记CD69、CD103、KLRG-1的缺失以及CTLA-4的表达。
通过静脉内注射将体外分化的细胞转移到小鼠体内。在指定的时间点,处死动物并按照标准方法分离来自脾、肺和小肠的淋巴细胞(IEL和LPL级分)。通过流式细胞术分析细胞群。通过它们的同源标记CD45.1的表达将转移的细胞与内源细胞区分开,并使用流式细胞术计数珠进行细胞计数。
<实施例1>
结果
从FoxP3+细胞中切除Tbx21导致1型TREG细胞减少。
TRM细胞表达T-bet但不表达Eomes(图1a,图7a)37。TREG细胞在不同组织中表达谱系相关的趋化因子受体,具有免疫1、2和3型特征(图1b)。为了测试表达T-bet或Eomes的TREG细胞是否影响TRM细胞,本发明人利用了Foxp3eYFP-Cre Tbx21fl/fl Rosa26tdRFP/tdRFP以及Foxp3eYFP-Cre Eomesfl/fl Rosa26tdRFP/tdRFP小鼠品系(分别参见Foxp3ΔTbx21和Foxp3ΔEomes),以及利用Foxp3eYFP-Cre Rosa26tdRFP/tdRFP小鼠品系作为对照组(Foxp3WT)(方法,图14)。T-bet和Eomes激活在1型免疫应答中重要的基因的转录,例如由T-bet反式激活的趋化因子受体CXCR338。与TREG细胞中缺失T-bet时增强的1型炎症一致35,39(图7b),在Foxp3ΔTbx21小鼠的脾脏中,而不是胸腺,显示了CXCR3+CD4+和CXCR3+CD8+T细胞成比例增加(图1c-d)。胸腺或脾脏中CD4+Foxp3-或TREG(CD4+Foxp3+)细胞的数量相似(图7c-h),但是但脾脏CD8+T细胞在Foxp3ΔTbx21动物中表现出增加的趋势(图7e)29,35,39。在三个月大的小鼠中未观察到自身免疫的迹象。
发明人证实Foxp3特异性靶向,CD4+CXCR3+Foxp3,但非CD4+CXCR3+Foxp3+。T细胞存在于Foxp3ΔTbx21小鼠中(图1c-e),但观察到Foxp3ΔEomes小鼠的外周淋巴器官中CXCR3+TREG细胞的比例和数量增加(图7i-k)。TREG细胞中T-bet的切除导致TREG亚群的分布改进,而非数量改变,并伴随3型TREG细胞的增加(图1f,图7l-n)。与存在于次级淋巴器官(SLO)中的TREG细胞群相比,LPL中的TREG细胞群显示出更活化的表型,表达更高水平的CD44(图1g)。神经粘连因子-1(Nrpl-1)和转录因子Helios主要存在于SLO TREG细胞中,但其在肠中的数量减少。与Foxp3WT对照小鼠相比,TREG细胞在Foxp3ΔTbx21中表现出相似的表型(图1h-i)29。与Foxp3WT对照小鼠相比,在效应T细胞和TEM细胞上表达的共抑制性受体杀伤细胞凝集素样受体G1(KLRG1)40,41在来自Foxp3ΔTbx21的TREG细胞上增加(图1j)。总的来说,这些数据显示,在缺失表达T-bet的TREG细胞的情况下,TREG细胞的数量及其表型保持相似,但是TREG亚群的比例发生了变化。
TREG细胞中Tbx21或Eomes的切除改变了CD8 T细胞的分布。
Foxp3ΔTbx21小鼠的脾脏的CD8+T细胞区室显示效应子(Teff)/TEM T细胞增加(图2a-b)。然而,与Foxp3WT对照相比,Foxp3ΔTbx21小鼠中的肠上皮内部分显示CD4和CD8 T细胞减少(图2c)。在CD8 IEL群体内,在诱导的CD8αβ+内观察到Foxp3ΔTbx21小鼠中显著减少,但在天然CD8αα+IEL群体中未观察到(图2d)。与Foxp3WT对照相比,Foxp3ΔTbx21小鼠的固有层(LP)区室显示CD4+T细胞减少,但CD8+T细胞未减少(图2e)。这种差异在除结肠以外的所有肠切片中都是明显的(图8a-d),其中TREG细胞中的CXCR3和T-bet表达是脱节的42。
尽管LPL区室中CD4+Foxp3-T细胞和TREG细胞的数量发生了变化,并且与Tbx21或Eomes的Foxp3依赖性切除无关,CD4+T细胞和TREG细胞的比例保持稳定(图8e-f)。然而,Foxp3ΔTbx21动物的T细胞群体显示出CD8 T细胞占有明显更大的比例(图2f),而在Foxp3ΔEomes动物中观察到相反的情况,特别是在其近端肠中(图2g)。这些数据表明TREG细胞中T-bet或Eomes的缺失不会改变CD4+Foxp3+和CD4+Foxp3-T细胞之间的比例分布,但对小肠中CD8+T细胞亚群的比例有显著影响。
Tbx21+和Eomes+TREG细胞影响CD8 T细胞的记忆区室。
在缺失T-bet足量TREG细胞的情况下,肠中TRM细胞的减少和循环效应物/TEM细胞的比例增加表明这些细胞在TRM细胞的产生或维持中具有潜在作用。尽管IEL数量有所减少(图2c-d),所有CD8+IEL表达TRM细胞标志物CD103和CD69(图2h)。Foxp3ΔTbx21,Foxp3ΔEomes和Foxp3WT小鼠中的LPL区室与CD4+Foxp3-T细胞相似,CD4+Foxp3-T细胞表达具有约一半共表达CD103的高水平的CD69(图8g)。尽管本发明人未观察到CD4 TRM细胞表型的差异,但Foxp3ΔTbx21动物显示CD4+Foxp3-T细胞和CD4+CD103+细胞数量减少的总体趋势(图8h)。
在Foxp3WT和Foxp3ΔEomes动物的LPL区室中,大多数CD8 T细胞表达TRM标记CD69和CD103(图2h-i)。相反,在Foxp3ΔTbx21动物中,超过一半的CD8+T细胞不表达CD103(图2h-i)。因此,尽管CD8+T细胞总数相似,但在缺失表达T-bet的TREG细胞的情况下,这些动物的肠中的CD8+TRM细胞的数量减少,效应物对TRM细胞的比例贡献高(图2i-j),这导致所有三种小鼠品系中CD4 T细胞和CD8 TRM细胞之间的比率恒定(图8i)。这些数据表明肠中的免疫网络被微调,在Foxp3ΔTbx21动物中观察到CD8+T细胞比率增加,这可能是由于不能发育成TRM细胞的CD8+效应T细胞的积累。
Tbx21+TREG细胞影响多种组织中TRM细胞的发育。
对于皮肤感染,已有报道在感染早期、在真皮中发现KLRG1+CD103-CD8+效应T细胞,但在感染晚期和表皮中未发现19。与在稳态条件下在Foxp3ΔTbx21动物的小肠中观察到的CD103-CD8+T细胞群体一致,在小肠的所有切片中观察到显著的KLRG1+CD8+T细胞群,约占总CD8 T细胞群的20%(图3a)。相反地,含有增加的CXCR3+TREG细胞的Foxp3ΔEomes动物(图7h)显示出其近端肠中KLRG1+CD8+T细胞数量减少(图3b)。
与CD103共染色证实KLRG1蛋白与CD103以互斥形式表达19,26,43,并且主要存在于Foxp3ΔTbx21小鼠中(图3c)。存在于Foxp3ΔTbx21动物中的CD103-KLRG1+CD8 T细胞表达高水平的Eomes,而在Foxp3WT中发现很少,在Foxp3ΔEomes动物中则更少(图3d,图9a)。与其效应物状态一致,Foxp3ΔTbx21动物中的KLRG1+Eomes+CD8 T细胞具有降低的促存活蛋白Bcl-2的表达水平,并在TRM细胞成熟期间上调44,45(图9b)。此外,与Foxp3WT对照相比,在Foxp3ΔTbx21动物中,表达CD8αα同二聚体的细胞比例降低,这些细胞与CD8αβ异源二聚体一起表达且具有上皮记忆CD8 T细胞46的特征(图9c)。最后,体内染色证实从LP区室分离的大多数细胞最近没有循环(图9d,e)。总之,在不希望与任何特定理论结合的情况下,这些数据显示,在缺失表达T-bet的TREG细胞的情况下,CD8+效应T细胞积聚在肠屏障处,在该处它们不向TRM细胞发育。
文献中描述了在许多组织中发现了TRM细胞的存在4。与肠的结果一致,与Foxp3ΔEomes和Foxp3WT动物相比,Foxp3ΔTbx21小鼠的肝脏和肺含有比例增加的效应CD8 T细胞,表达高水平的KLRG1和Eomes(图3e-f)。由于CD103表达不被认为是肝脏中TRM细胞的充分标志物,所以本发明人评估了KLRG1和Eomes阴性的CD8+CD69+细胞的比例。在非淋巴组织中的评估中,与Foxp3ΔEomes和Foxp3ΔWT小鼠相比,Foxp3ΔTbx21小鼠含有更少的TRM细胞(图3g-h)。在不希望与任何特定理论结合的情况下,这些数据表明1型TREG细胞对于多种组织中TRM细胞的产生是重要的。
TRM细胞区室受损导致对病原体入侵的保护的降低。
本发明人假设Foxp3ΔTbx21小鼠中TRM细胞数量的减少将导致对新感染的保护降低,因为旁观者介导的TRM细胞活化是限制病原体入侵的重要防御机制47,48,49。我们通过施用抗CD3抗体来测试肠CD8+T细胞的急性应答,施用两天后评估LPL T细胞应答。与具有增加的PD-1表达和减少的颗粒酶B的Eomes-CD8 T细胞相比,Eomes+CD8 T细胞显示出降低的活性谱(图4a,b,图10a-c)50。
接下来,本发明人用细胞内原生寄生虫蠕形艾美球虫(Ev)攻击小鼠,这种寄生虫感染小鼠的小肠上皮细胞。在这种感染模型中,淋巴细胞减少寄生虫负荷(图10d-f),CD8+T细胞和IFNγ在清除中起重要作用10,51,52。尽管缺失1型TREG细胞可能会导致T细胞介导的免疫力增强53,但实际上与Foxp3WT和Foxp3ΔEomes动物相比,Foxp3ΔTbx21小鼠显示对Ev感染的控制受损(图4c-f)。Foxp3ΔTbx21小鼠的表现与没有淋巴细胞的小鼠相反(图10d-e),但与TRM细胞存活所需的IL-15Rα缺陷小鼠26一致,与Foxp3WT,Foxp3ΔEomes和Rag2-/-动物相比,Foxp3ΔTbx21小鼠中存在更多效应细胞,但其具有耗尽表型,Foxp3ΔTbx21小鼠的体重减轻(图4g-h,图10f)。TRM细胞提供的保护,可防止肝脏中的顶复动物亚门寄生虫疟原虫49和皮肤中的病毒26,这些数据与之相符。
1型TREG细胞增强TRM的发育
来自Foxp3ΔTbx21或Foxp3ΔEomes小鼠的T细胞,其在稳定状态下、在激活时或在TRM细胞建立时,与Foxp3WT对照小鼠在Tbet或Eomes的表达上无法区分(图11a-d)。为了评估Foxp3ΔTbx21动物(与Foxp3WT相比)组织中效应T细胞的积累和TRM细胞的减少是否是CD8T细胞固有的,我们用CD45.1对照小鼠和Foxp3ΔTbx21小鼠产生混合的骨髓嵌合体。发现的TRM细胞显示两个供体的贡献相似(图5a-b)。此外,我们将来自Foxp3ΔTbx21小鼠(CD45.2)的CD8 T细胞转移到对照CD45.1动物中。一天后,小鼠接受Ev攻击,两周后清除54。在在寄生虫清除后一周后,Teff细胞减少时,评估CD8CD45.1CD103+TRM细胞的发育(图5c)。在转移的CD45.2+群体内,TRM细胞高效发育(图5d)。总的来说,这些结果表明Foxp3ΔTbx21小鼠中的CD8 T细胞的外在缺陷抑制TRM细胞的产生。为了证实这一点,我们将CD45.1+CD8+T细胞(CD8CD45.1)转移到CD45.2+Foxp3ΔTbx21或Foxp3WT动物中。在感染高峰期(第10天),LPL中存在TRM细胞和效应T细胞(图11e,f)。在Foxp3WT宿主中,大多数转移的CD8CD45.1细胞显示具有低Eomes水平的CD103表达的特征性TRM细胞谱(图5e)。相反地,转移到Foxp3ΔTbx21宿主中的CD8CD45.1 T细胞显示部分TRM细胞形成,这些细胞中的大部分显示效应子表型,具有Eomes表达并缺失CD103表达(图5e)。与Foxp3WT宿主相比,Foxp3ΔTbx21宿主中转移的CD8CD45.1 T细胞的累积数量相似,而Foxp3ΔTbx21宿主中TRM细胞减少(图5f)。对Ev感染后9周的小鼠进行分析,分析结果显示在Foxp3ΔTbx21动物中总体上可检测到的转移的CD8CD45.1 T细胞数量减少(图5g)。这些数据证实在Foxp3ΔTbx21动物中观察到的受损的CD8+TRM细胞分化是CD8+T细胞群体非固有的。本发明人的转移系统能够检验以下假设:TREG细胞通过将CD8CD45.1 T细胞和TREG WT细胞同时转移到Foxp3ΔTbx21动物中促进TRM细胞发育(图11g)。在存在对照TREG细胞的情况下,Foxp3ΔTbx21动物中TRM细胞的产生恢复至在Foxp3WT对照中观察到的水平(图5h),这也支持了上述假设。
为了理解表达T-bet的TREG细胞是否具有可以解释其在TRM发育中的作用的特定功能属性,本发明人利用了最近公开可用的一组单TREG细胞测序数据55。与先前的报道一致,尽管可能存在小的趋势,但发明人并未发现由特征谱系转录因子Tbx21,Gata3或Rorc的存在所定义的TREG细胞亚群中,TREG细胞的效应分子如IL-10,IL-35,TGFβ,CD25,LAG3或CTLA-4的显著差异(图5i)。此外,据报道T-bet缺陷型TREG细胞显示出与对照TREG细胞相似的抑制能力35,37。
TRM的发育依赖于使TGFβ局部可用的TREG的募集。
本发明人的观察依赖于无特定病原体条件下的微生物存在和细胞内小肠寄生虫Ev,这引起了非常局部的反应。本发明人利用我们的CD8CD45.1 T细胞转移系统(图5c),用假结核耶尔森氏菌(Yptb)攻击小鼠,据报道,其可诱导TRM细胞58,59。与使用Ev获得的结果一致,Yptb攻击导致Foxp3WT动物中TRM细胞有效发育,其在Foxp3ΔTbx21动物中显著降低(图6a,b)。
CD8CD45.1 T细胞向Foxp3ΔTbx21小鼠的转移模型使得本发明人能够研究TREG细胞对促进TRM细胞发育的贡献。在缺失T-bet的情况下,TREG细胞表达相似水平的CD103,CCR6和P-选择蛋白35,但不能表达CXCR3,CXCR3对于将T细胞定位于非淋巴组织中的感染区域是重要的7,35,39,60。局部炎症环境控制TRM前体细胞和TREG细胞的募集58,61(图6c)。重要的是,在Ev感染时募集的TREG细胞主要显示表达CXCR3的1型表型(图6d,图11g-h)。考虑到Foxp3ΔTbx21小鼠结肠中的效应细胞和TRM细胞没有改变,其中CXCR3的表达不依赖于T-bet,本发明人假设TREG细胞亚群中的T-bet表达促进这些细胞募集到感染部位并使它们与TRM前体细胞紧密接近。Tbx21的Foxp3依赖性切除导致Ev感染后TREG细胞的募集减少(图6c),主要是由于那些表达CXCR3的细胞(图6d),这与上述假设相符。与募集1型TREG细胞一致,本发明人发现Ev感染后肠组织中CXCR3的配体CXCL10的表达增加(图6e)。在稳态条件下,Foxp3ΔTbx21小鼠的LPL区室中的CD4 T细胞数量减少(图1e),并且可能在TRM细胞产生中起另外的作用。然而,Ev感染导致CD4 T细胞大量募集到由T辅助1表型主导的LPL区室(图12a,b)。利用转移系统(图5a),在与CD8CD45.1细胞伴随转移时,与CXCR3足够的对照细胞相比,CXCR3缺陷型TREG细胞不能支持TRM细胞的有效发育(图6f)。
微生物入侵区域中CD8+和CD4+T细胞与其他免疫细胞的聚集体是常见的58,62,63,64,65,其他免疫细胞例如巨噬细胞和树突细胞,但不包括B细胞。CD4+和CD8+T细胞之间的相互作用虽然不是维持T细胞所必需的,但可能构成可能支持TRM分化的不同微环境。与此一致,本发明人经常在Foxp3WT小鼠中观察到转移的CD8CD45.1 T细胞,该T细胞与表达Foxp3的TREG细胞紧密接近,尽管CD8 T细胞浸润相似,但这种转移的CD8CD45.1 T细胞在Foxp3ΔTbx21动物中不容易观察到(图6g,图12c)。
CXCR3促进TRM发育的要求表明TREG提供短程作用或细胞结合的效应分子。1型细胞因子,例如IL-12,可以维持高水平的T-bet和Eomes,从而阻止TRM细胞的分化和CD103的表达。IL12Rβ2缺陷的CD8+T细胞表达CD103的细胞比例增加,而T细胞簇具有更高的TGFβ转录59。IL-10可以降低IL-12的表达和树突状细胞的成熟66。然而,IL-10缺陷的TREG细胞能够协助TRM细胞的有效发育(图6h),且IL-10缺陷的动物未显示TRM细胞区室减少(图12d,e)。此外,EBI3缺陷的TREG细胞,不能产生IL-35,同样促进TRM细胞的产生(图6i),在EBI3缺陷型骨髓嵌合动物中TRM细胞区室也没有减少(图12g,h)。
许多细胞,特别是粘膜屏障细胞,能够产生TGFβ67。然而,TGFβ作为无活性前体产生,其需要从其潜伏期相关肽切割。TGF-β具有有效的细胞调节活性,作用于多种免疫和非免疫细胞类型,因此其可用性在局部微环境中受到严格调节。最近显示TREG细胞可通过整联蛋白αvβ8激活TGFβ,并且该蛋白在活化/效应TREG细胞中上调,从而减少局部生物活性TGFβ68,并且发明人假设表达CXCR3的TREG细胞的特异性募集,其在稳定状态下不显示对TGFβ1或Itgb8的差异表达(图5f,图12f)42,具有效应CD8 T细胞的位点可以使TGFβ局部释放。与Foxp3ΔTbx21小鼠相似,由Foxp3ΔItgβ8小鼠产生的骨髓嵌合体显示TRM细胞减少和表达KLRG1的效应CD8+T细胞增加(图12g,i)。重要的是,仅在TREG细胞中不存在Itgβ8时,TRM细胞生成的促进效率低下(图6j)。为了理解TREG衍生的TGFβ是否起决定性作用,本发明人使用了Foxp3ΔTgfβ1小鼠。与Foxp3ΔItgβ8小鼠一致,TREG细胞中缺失TGFβ1仅导致效应T细胞增加和TRM细胞比例降低(图12g,j)。在过继转移后,缺乏TGFβ1供应的TREG细胞不能挽救Foxp3ΔTbx21小鼠中TRM细胞的发育(图6k)。总的来说,在不希望与任何特定理论结合的情况下,数据显示TREG细胞通过T-bet诱导的CXCR3表达募集产生TGFβ1,并通过αvβ8整联蛋白的表达使其具有局部生物利用度,以促进发炎组织中的TRM细胞的发育。
讨论
在非淋巴组织中诱导长寿细胞免疫对于防止再感染是重要的,也是疫苗设计的主要目标。本发明人的数据支持一种模型,其中CD8+T细胞作为效应细胞或记忆细胞前体存在于组织中,随后在接收局部线索后分化为TRM细胞69。SLO中的T细胞活化诱导大量组织归巢受体的表达,这些受体将活化的T细胞导向非淋巴组织,确保效应T细胞检查大多数外周组织69,70。TRM细胞的独特特征表明,组织微环境中的因子指导效应细胞分化成TRM细胞。
发明人的数据基于局部感染模型和多克隆TCR库。使用局部感染模型,已经表明最佳的TRM细胞发育,而非维持,需要炎症介导的运输和局部微环境中的同源抗原61,71,72。虽然这与使用系统性病毒感染的观察结果相反,在这些观察中IEL数量保持稳定,但发明人的观察结果与先前报道的小肠感染一致,表明这是局部炎症的一个特征。TRM细胞从募集的效应子或记忆前体T细胞分化可能需要局部提示,例如细胞因子和次级抗原接触。本发明人的数据支持该模型,根据将TREG细胞募集到炎症部位的需要以及其提高生物活性TGFβ水平、从而促进效应物向记忆物发展的能力,还可对模型进行扩展。在总TREG细胞消耗后,病毒感染后中枢神经系统中TRM细胞的数量减少73,表明TREG细胞在TRM细胞发育或维持中的作用。本发明人扩展了这一观察结果,显示1型TREG细胞的局部募集是关键的,其中Itgβ8的表达促进TRM发育,这严重依赖于局部供应的TGFβ及其生物利用度23,26。局部抗原可以将CD8+T细胞保留在与感染细胞形成稳定接触的组织中,随后CD69被重新表达,这一过程需要CXCR3表达74。尽管在非发炎组织中,本发明人报告了CD4和TREG细胞的减少,但是在炎症时募集了CD4 T细胞。尽管TREG细胞的补充足以增强TRM细胞发育,但这并不排除CD4 T细胞的支持作用。本发明人表明,为了将TREG细胞募集到炎症部位,需要CXCR3的表达,也即人体中的1型TREG细胞75,这为上游转录因子T-bet的需求提供了合理性。在缺失CXCR3的情况下,TREG细胞组织募集受到限制,导致1型免疫和免疫病理学增强42,76。虽然TREG细胞的缺失能够增强免疫反应,甚至导致无菌免疫,但是有报道揭示了后续免疫损失53。本发明人在此表明,1型TREG细胞提供TGFβ并使其局部可用于αvβ8整联蛋白的表达,从而促进TRM细胞的发育,支持终身免疫监视并增加针对入侵微生物的组织保护。
本发明人观察到,在缺失表达T-bet的TREG细胞的情况下进行评估,几种组织中TRM分化降低。这突出表明特定的组织微环境在TRM细胞的发育或1型TREG细胞的募集中不起关键作用。然而,组织特异性差异可能会改变TRM发育的幅度或其表型。已知肠中的TRM细胞主要产生IFNγ,而表皮中的TRM细胞已经显示能够在微生物攻击后产生IL-17。再者,额外的组织损伤可以改变表皮TRM细胞功能,促进伤口修复77,78。结肠是明显的例外,在结肠中本发明人在缺失1型TREG细胞的情况下未发现TRM或效应细胞的改变,T细胞亚群中的比例也未改变。尽管TREG细胞需要T-bet表达CXCR3并被募集到炎症部位,但T-bet似乎不能控制结肠中CXCR3的表达42。结肠中T-bet和CXCR3之间的分离表明在含有最大含量微生物的器官中存在替代性免疫调节。
TGFβ是体外和体内CD8+T细胞上CD103表达的有效驱动因子22,并且已显示其能降低KLRG1表达43。此外,慢性感染期间CD103表达减少表明减少炎症对TRM细胞发育的重要性23,79。此外,TGFβRII缺陷的CD8 T细胞不能成为或保持TRM细胞19,58。除CD103+TRM细胞外,还报道了CD103-TRM细胞群22,58,62。该群体的稳定性可能取决于组织类型和抗原持久性。在本发明人模型中,在对照小鼠中,在无特定病原体条件下的稳定状态下,以及Ev攻击后,CD103-T细胞是次要群体。相反地,在Foxp3ΔTbx21小鼠中观察到的CD103-T细胞表达KLRG1和高水平的Eomes,表达CD103并关闭Eomes是T细胞在过渡期的特征80。在不希望与任何特定理论结合的情况下,本发明人的数据确实揭示了1型TREG细胞在TRM细胞发育中的重要作用,但(在缺失TREG细胞情况下)仍然可以产生更小一些的TRM细胞群,这表明其他细胞可能在释放TGFβ中发挥额外作用。已经报道了产生生物活性TGFβ的替代来源,其中包括对维持TRM细胞重要的基质上皮细胞81。
TREG细胞对于抑制过度的免疫反应,从而预防自身免疫和免疫病理学是至关重要的,并且可以降低在血液中测量的感染和疫苗接种后的反应幅度。然而,本发明人的数据突出了TREG细胞在从效应体或记忆前体有效生成组织驻留记忆T细胞方面的重要作用,缺失TREG细胞情况下这些细胞将耗尽。因此,TREG细胞确保关键数量的T细胞可用于组织中的免疫监视,以预防或减少再感染以及减少新感染的病原体负荷。
<实施例2>
结果
本发明人评估了在培养基中缺失TREG细胞时是否可能产生TRM细胞,如图15所示。
参见图16,显示了在具有IL-15、抗原呈递细胞(BMDC)和先前活化的CD8 T细胞的培养物中添加TGFβ足以建立TRM特征,例如在不存在另外的TREG细胞的情况下CD69和CD103的持续表达。
参见图17,显示了IL-15、TGFβ、抗原呈递细胞和先前活化的CD8 T细胞与添加IL-7的组合可以基于其CTLA-4表达增强产生的TRM细胞的迁移能力(前沿免疫学,2018年11月27日;Brunner-Weinzierl和Rudd;Kieke等人,PLOS One,27/5/09)。
参见图18,显示了IL-15,TGFβ,抗原呈递细胞和先前活化的CD8 T细胞与添加IL-2的组合可以基于其CTLA-4表达增强所产生的TRM细胞的迁移能力,但与每个细胞或生物重复中IL-7的添加量没有一致性。
此外,本发明人评估了体外产生的细胞是否保持其治疗性质,特别是它们在组织内体内迁移和存活的能力,相关实验装置在图19中详细示出。
本发明人在由效应CD8+T细胞和骨髓衍生的树突细胞(BMDC)组成的体外装置中重建了体内条件。本发明人表明,添加生物活性TGFβ可以取代TREG细胞在类似TRM细胞的T细胞发育中的作用,具有标志物和组织保留因子CD69和CD103的持续表达(图20)。CD69是通常在T细胞活化后瞬时表达的活化标记。CD69是一种C型凝集素,最有可能参与TRM细胞在非淋巴组织中的滞留,包括在实体肿瘤中98。CD69可以与鞘氨醇-1-磷酸(S1P)1形成复合物,从而阻止其与S1P受体结合,SIP受体会触发T细胞离开组织。此外,数据显示添加IL-7和IL-2诱导强烈的CTLA-4表达(图17和18),其与增强的T细胞迁移相关99 , 100,101。尽管添加IL-2后CTLA-4表达似乎更强,但重复实验显示添加IL-7比添加IL-2具有更好的一致性。而且,添加IL-7导致单个细胞具有更强的CTLA-4表达。
在体内转移体外产生和扩增的T细胞的数据显示,在组织中未发现大量在缺失TGFβ的情况下产生的效应细胞(图21,22,23和24)。另一方面,在所有测试的器官中,例如在肺、肝、固有层以及小肠的上皮内区室中,在转移后40天后都容易发现TRM细胞,特别是在用IL-7刺激时(图21,22,23和24)。
总结
总而言之,本发明人开发了一种高度新颖和创新的方案来产生能够深入穿透肿瘤(特别是实体肿瘤)的T细胞(称为TRM细胞),其可作为针对肿瘤或感染的T细胞疗法的阶段性进展。发明人的方案基于CTLA-4、CD69和CD103的表达产生并扩增具有迁移和组织穿透细胞表型的细胞。在过继转移到完整的小鼠宿主中至少40天后,在各种淋巴和非淋巴组织中容易发现产生的细胞,其符合上述表型且与效应T细胞相反。本发明人的工作将为有效治疗器官感染和患有实体肿瘤的癌症患者做出重要进展,这些病症更难治疗。然而,本发明人相信TRM细胞的组织穿透能力将超出感染或原发性肿瘤的靶向,并且可以提供针对转移的关键器官范围的免疫监视,转移通常迁移到远离原肿瘤的、不易接近的组织。
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Glu Ser Gly Gln Glu Asn Asp Ala Thr Ser Pro His Phe Ser Thr Arg
20 25 30
His Glu Gly Ser Phe Gln Val Pro Val Leu Cys Ala Val Met Asn Val
35 40 45
Val Phe Ile Thr Ile Leu Ile Ile Ala Leu Ile Ala Leu Ser Val Gly
50 55 60
Gln Tyr Asn Cys Pro Gly Gln Tyr Thr Phe Ser Met Pro Ser Asp Ser
65 70 75 80
His Val Ser Ser Cys Ser Glu Asp Trp Val Gly Tyr Gln Arg Lys Cys
85 90 95
Tyr Phe Ile Ser Thr Val Lys Arg Ser Trp Thr Ser Ala Gln Asn Ala
100 105 110
Cys Ser Glu His Gly Ala Thr Leu Ala Val Ile Asp Ser Glu Lys Asp
115 120 125
Met Asn Phe Leu Lys Arg Tyr Ala Gly Arg Glu Glu His Trp Val Gly
130 135 140
Leu Lys Lys Glu Pro Gly His Pro Trp Lys Trp Ser Asn Gly Lys Glu
145 150 155 160
Phe Asn Asn Trp Phe Asn Val Thr Gly Ser Asp Lys Cys Val Phe Leu
165 170 175
Lys Asn Thr Glu Val Ser Ser Met Glu Cys Glu Lys Asn Leu Tyr Trp
180 185 190
Ile Cys Asn Lys Pro Tyr Lys
195
<210> 4
<211> 1179
<212> PRT
<213> 智人(Homo sapiens)
<400> 4
Met Trp Leu Phe His Thr Leu Leu Cys Ile Ala Ser Leu Ala Leu Leu
1 5 10 15
Ala Ala Phe Asn Val Asp Val Ala Arg Pro Trp Leu Thr Pro Lys Gly
20 25 30
Gly Ala Pro Phe Val Leu Ser Ser Leu Leu His Gln Asp Pro Ser Thr
35 40 45
Asn Gln Thr Trp Leu Leu Val Thr Ser Pro Arg Thr Lys Arg Thr Pro
50 55 60
Gly Pro Leu His Arg Cys Ser Leu Val Gln Asp Glu Ile Leu Cys His
65 70 75 80
Pro Val Glu His Val Pro Ile Pro Lys Gly Arg His Arg Gly Val Thr
85 90 95
Val Val Arg Ser His His Gly Val Leu Ile Cys Ile Gln Val Leu Val
100 105 110
Arg Arg Pro His Ser Leu Ser Ser Glu Leu Thr Gly Thr Cys Ser Leu
115 120 125
Leu Gly Pro Asp Leu Arg Pro Gln Ala Gln Ala Asn Phe Phe Asp Leu
130 135 140
Glu Asn Leu Leu Asp Pro Asp Ala Arg Val Asp Thr Gly Asp Cys Tyr
145 150 155 160
Ser Asn Lys Glu Gly Gly Gly Glu Asp Asp Val Asn Thr Ala Arg Gln
165 170 175
Arg Arg Ala Leu Glu Lys Glu Glu Glu Glu Asp Lys Glu Glu Glu Glu
180 185 190
Asp Glu Glu Glu Glu Glu Ala Gly Thr Glu Ile Ala Ile Ile Leu Asp
195 200 205
Gly Ser Gly Ser Ile Asp Pro Pro Asp Phe Gln Arg Ala Lys Asp Phe
210 215 220
Ile Ser Asn Met Met Arg Asn Phe Tyr Glu Lys Cys Phe Glu Cys Asn
225 230 235 240
Phe Ala Leu Val Gln Tyr Gly Gly Val Ile Gln Thr Glu Phe Asp Leu
245 250 255
Arg Asp Ser Gln Asp Val Met Ala Ser Leu Ala Arg Val Gln Asn Ile
260 265 270
Thr Gln Val Gly Ser Val Thr Lys Thr Ala Ser Ala Met Gln His Val
275 280 285
Leu Asp Ser Ile Phe Thr Ser Ser His Gly Ser Arg Arg Lys Ala Ser
290 295 300
Lys Val Met Val Val Leu Thr Asp Gly Gly Ile Phe Glu Asp Pro Leu
305 310 315 320
Asn Leu Thr Thr Val Ile Asn Ser Pro Lys Met Gln Gly Val Glu Arg
325 330 335
Phe Ala Ile Gly Val Gly Glu Glu Phe Lys Ser Ala Arg Thr Ala Arg
340 345 350
Glu Leu Asn Leu Ile Ala Ser Asp Pro Asp Glu Thr His Ala Phe Lys
355 360 365
Val Thr Asn Tyr Met Ala Leu Asp Gly Leu Leu Ser Lys Leu Arg Tyr
370 375 380
Asn Ile Ile Ser Met Glu Gly Thr Val Gly Asp Ala Leu His Tyr Gln
385 390 395 400
Leu Ala Gln Ile Gly Phe Ser Ala Gln Ile Leu Asp Glu Arg Gln Val
405 410 415
Leu Leu Gly Ala Val Gly Ala Phe Asp Trp Ser Gly Gly Ala Leu Leu
420 425 430
Tyr Asp Thr Arg Ser Arg Arg Gly Arg Phe Leu Asn Gln Thr Ala Ala
435 440 445
Ala Ala Ala Asp Ala Glu Ala Ala Gln Tyr Ser Tyr Leu Gly Tyr Ala
450 455 460
Val Ala Val Leu His Lys Thr Cys Ser Leu Ser Tyr Ile Ala Gly Ala
465 470 475 480
Pro Arg Tyr Lys His His Gly Ala Val Phe Glu Leu Gln Lys Glu Gly
485 490 495
Arg Glu Ala Ser Phe Leu Pro Val Leu Glu Gly Glu Gln Met Gly Ser
500 505 510
Tyr Phe Gly Ser Glu Leu Cys Pro Val Asp Ile Asp Met Asp Gly Ser
515 520 525
Thr Asp Phe Leu Leu Val Ala Ala Pro Phe Tyr His Val His Gly Glu
530 535 540
Glu Gly Arg Val Tyr Val Tyr Arg Leu Ser Glu Gln Asp Gly Ser Phe
545 550 555 560
Ser Leu Ala Arg Ile Leu Ser Gly His Pro Gly Phe Thr Asn Ala Arg
565 570 575
Phe Gly Phe Ala Met Ala Ala Met Gly Asp Leu Ser Gln Asp Lys Leu
580 585 590
Thr Asp Val Ala Ile Gly Ala Pro Leu Glu Gly Phe Gly Ala Asp Asp
595 600 605
Gly Ala Ser Phe Gly Ser Val Tyr Ile Tyr Asn Gly His Trp Asp Gly
610 615 620
Leu Ser Ala Ser Pro Ser Gln Arg Ile Arg Ala Ser Thr Val Ala Pro
625 630 635 640
Gly Leu Gln Tyr Phe Gly Met Ser Met Ala Gly Gly Phe Asp Ile Ser
645 650 655
Gly Asp Gly Leu Ala Asp Ile Thr Val Gly Thr Leu Gly Gln Ala Val
660 665 670
Val Phe Arg Ser Arg Pro Val Val Arg Leu Lys Val Ser Met Ala Phe
675 680 685
Thr Pro Ser Ala Leu Pro Ile Gly Phe Asn Gly Val Val Asn Val Arg
690 695 700
Leu Cys Phe Glu Ile Ser Ser Val Thr Thr Ala Ser Glu Ser Gly Leu
705 710 715 720
Arg Glu Ala Leu Leu Asn Phe Thr Leu Asp Val Asp Val Gly Lys Gln
725 730 735
Arg Arg Arg Leu Gln Cys Ser Asp Val Arg Ser Cys Leu Gly Cys Leu
740 745 750
Arg Glu Trp Ser Ser Gly Ser Gln Leu Cys Glu Asp Leu Leu Leu Met
755 760 765
Pro Thr Glu Gly Glu Leu Cys Glu Glu Asp Cys Phe Ser Asn Ala Ser
770 775 780
Val Lys Val Ser Tyr Gln Leu Gln Thr Pro Glu Gly Gln Thr Asp His
785 790 795 800
Pro Gln Pro Ile Leu Asp Arg Tyr Thr Glu Pro Phe Ala Ile Phe Gln
805 810 815
Leu Pro Tyr Glu Lys Ala Cys Lys Asn Lys Leu Phe Cys Val Ala Glu
820 825 830
Leu Gln Leu Ala Thr Thr Val Ser Gln Gln Glu Leu Val Val Gly Leu
835 840 845
Thr Lys Glu Leu Thr Leu Asn Ile Asn Leu Thr Asn Ser Gly Glu Asp
850 855 860
Ser Tyr Met Thr Ser Met Ala Leu Asn Tyr Pro Arg Asn Leu Gln Leu
865 870 875 880
Lys Arg Met Gln Lys Pro Pro Ser Pro Asn Ile Gln Cys Asp Asp Pro
885 890 895
Gln Pro Val Ala Ser Val Leu Ile Met Asn Cys Arg Ile Gly His Pro
900 905 910
Val Leu Lys Arg Ser Ser Ala His Val Ser Val Val Trp Gln Leu Glu
915 920 925
Glu Asn Ala Phe Pro Asn Arg Thr Ala Asp Ile Thr Val Thr Val Thr
930 935 940
Asn Ser Asn Glu Arg Arg Ser Leu Ala Asn Glu Thr His Thr Leu Gln
945 950 955 960
Phe Arg His Gly Phe Val Ala Val Leu Ser Lys Pro Ser Ile Met Tyr
965 970 975
Val Asn Thr Gly Gln Gly Leu Ser His His Lys Glu Phe Leu Phe His
980 985 990
Val His Gly Glu Asn Leu Phe Gly Ala Glu Tyr Gln Leu Gln Ile Cys
995 1000 1005
Val Pro Thr Lys Leu Arg Gly Leu Gln Val Val Ala Val Lys Lys
1010 1015 1020
Leu Thr Arg Thr Gln Ala Ser Thr Val Cys Thr Trp Ser Gln Glu
1025 1030 1035
Arg Ala Cys Ala Tyr Ser Ser Val Gln His Val Glu Glu Trp His
1040 1045 1050
Ser Val Ser Cys Val Ile Ala Ser Asp Lys Glu Asn Val Thr Val
1055 1060 1065
Ala Ala Glu Ile Ser Trp Asp His Ser Glu Glu Leu Leu Lys Asp
1070 1075 1080
Val Thr Glu Leu Gln Ile Leu Gly Glu Ile Ser Phe Asn Lys Ser
1085 1090 1095
Leu Tyr Glu Gly Leu Asn Ala Glu Asn His Arg Thr Lys Ile Thr
1100 1105 1110
Val Val Phe Leu Lys Asp Glu Lys Tyr His Ser Leu Pro Ile Ile
1115 1120 1125
Ile Lys Gly Ser Val Gly Gly Leu Leu Val Leu Ile Val Ile Leu
1130 1135 1140
Val Ile Leu Phe Lys Cys Gly Phe Phe Lys Arg Lys Tyr Gln Gln
1145 1150 1155
Leu Asn Leu Glu Ser Ile Arg Lys Ala Gln Leu Lys Ser Glu Asn
1160 1165 1170
Leu Leu Glu Glu Glu Asn
1175
<210> 5
<211> 195
<212> PRT
<213> 智人(Homo sapiens)
<400> 5
Met Thr Asp Ser Val Ile Tyr Ser Met Leu Glu Leu Pro Thr Ala Thr
1 5 10 15
Gln Ala Gln Asn Asp Tyr Gly Pro Gln Gln Lys Ser Ser Ser Ser Arg
20 25 30
Pro Ser Cys Ser Cys Leu Val Ala Ile Ala Leu Gly Leu Leu Thr Ala
35 40 45
Val Leu Leu Ser Val Leu Leu Tyr Gln Trp Ile Leu Cys Gln Gly Ser
50 55 60
Asn Tyr Ser Thr Cys Ala Ser Cys Pro Ser Cys Pro Asp Arg Trp Met
65 70 75 80
Lys Tyr Gly Asn His Cys Tyr Tyr Phe Ser Val Glu Glu Lys Asp Trp
85 90 95
Asn Ser Ser Leu Glu Phe Cys Leu Ala Arg Asp Ser His Leu Leu Val
100 105 110
Ile Thr Asp Asn Gln Glu Met Ser Leu Leu Gln Val Phe Leu Ser Glu
115 120 125
Ala Phe Cys Trp Ile Gly Leu Arg Asn Asn Ser Gly Trp Arg Trp Glu
130 135 140
Asp Gly Ser Pro Leu Asn Phe Ser Arg Ile Ser Ser Asn Ser Phe Val
145 150 155 160
Gln Thr Cys Gly Ala Ile Asn Lys Asn Gly Leu Gln Ala Ser Ser Cys
165 170 175
Glu Val Pro Leu His Trp Val Cys Lys Lys Cys Pro Phe Ala Asp Gln
180 185 190
Ala Leu Phe
195
<210> 6
<211> 686
<212> PRT
<213> 智人(Homo sapiens)
<400> 6
Met Gln Leu Gly Glu Gln Leu Leu Val Ser Ser Val Asn Leu Pro Gly
1 5 10 15
Ala His Phe Tyr Pro Leu Glu Ser Ala Arg Gly Gly Ser Gly Gly Ser
20 25 30
Ala Gly His Leu Pro Ser Ala Ala Pro Ser Pro Gln Lys Leu Asp Leu
35 40 45
Asp Lys Ala Ser Lys Lys Phe Ser Gly Ser Leu Ser Cys Glu Ala Val
50 55 60
Ser Gly Glu Pro Ala Ala Ala Ser Ala Gly Ala Pro Ala Ala Met Leu
65 70 75 80
Ser Asp Thr Asp Ala Gly Asp Ala Phe Ala Ser Ala Ala Ala Val Ala
85 90 95
Lys Pro Gly Pro Pro Asp Gly Arg Lys Gly Ser Pro Cys Gly Glu Glu
100 105 110
Glu Leu Pro Ser Ala Ala Ala Ala Ala Ala Ala Ala Ala Ala Ala Ala
115 120 125
Ala Ala Thr Ala Arg Tyr Ser Met Asp Ser Leu Ser Ser Glu Arg Tyr
130 135 140
Tyr Leu Gln Ser Pro Gly Pro Gln Gly Ser Glu Leu Ala Ala Pro Cys
145 150 155 160
Ser Leu Phe Pro Tyr Gln Ala Ala Ala Gly Ala Pro His Gly Pro Val
165 170 175
Tyr Pro Ala Pro Asn Gly Ala Arg Tyr Pro Tyr Gly Ser Met Leu Pro
180 185 190
Pro Gly Gly Phe Pro Ala Ala Val Cys Pro Pro Gly Arg Ala Gln Phe
195 200 205
Gly Pro Gly Ala Gly Ala Gly Ser Gly Ala Gly Gly Ser Ser Gly Gly
210 215 220
Gly Gly Gly Pro Gly Thr Tyr Gln Tyr Ser Gln Gly Ala Pro Leu Tyr
225 230 235 240
Gly Pro Tyr Pro Gly Ala Ala Ala Ala Gly Ser Cys Gly Gly Leu Gly
245 250 255
Gly Leu Gly Val Pro Gly Ser Gly Phe Arg Ala His Val Tyr Leu Cys
260 265 270
Asn Arg Pro Leu Trp Leu Lys Phe His Arg His Gln Thr Glu Met Ile
275 280 285
Ile Thr Lys Gln Gly Arg Arg Met Phe Pro Phe Leu Ser Phe Asn Ile
290 295 300
Asn Gly Leu Asn Pro Thr Ala His Tyr Asn Val Phe Val Glu Val Val
305 310 315 320
Leu Ala Asp Pro Asn His Trp Arg Phe Gln Gly Gly Lys Trp Val Thr
325 330 335
Cys Gly Lys Ala Asp Asn Asn Met Gln Gly Asn Lys Met Tyr Val His
340 345 350
Pro Glu Ser Pro Asn Thr Gly Ser His Trp Met Arg Gln Glu Ile Ser
355 360 365
Phe Gly Lys Leu Lys Leu Thr Asn Asn Lys Gly Ala Asn Asn Asn Asn
370 375 380
Thr Gln Met Ile Val Leu Gln Ser Leu His Lys Tyr Gln Pro Arg Leu
385 390 395 400
His Ile Val Glu Val Thr Glu Asp Gly Val Glu Asp Leu Asn Glu Pro
405 410 415
Ser Lys Thr Gln Thr Phe Thr Phe Ser Glu Thr Gln Phe Ile Ala Val
420 425 430
Thr Ala Tyr Gln Asn Thr Asp Ile Thr Gln Leu Lys Ile Asp His Asn
435 440 445
Pro Phe Ala Lys Gly Phe Arg Asp Asn Tyr Asp Ser Ser His Gln Ile
450 455 460
Val Pro Gly Gly Arg Tyr Gly Val Gln Ser Phe Phe Pro Glu Pro Phe
465 470 475 480
Val Asn Thr Leu Pro Gln Ala Arg Tyr Tyr Asn Gly Glu Arg Thr Val
485 490 495
Pro Gln Thr Asn Gly Leu Leu Ser Pro Gln Gln Ser Glu Glu Val Ala
500 505 510
Asn Pro Pro Gln Arg Trp Leu Val Thr Pro Val Gln Gln Pro Gly Thr
515 520 525
Asn Lys Leu Asp Ile Ser Ser Tyr Glu Ser Glu Tyr Thr Ser Ser Thr
530 535 540
Leu Leu Pro Tyr Gly Ile Lys Ser Leu Pro Leu Gln Thr Ser His Ala
545 550 555 560
Leu Gly Tyr Tyr Pro Asp Pro Thr Phe Pro Ala Met Ala Gly Trp Gly
565 570 575
Gly Arg Gly Ser Tyr Gln Arg Lys Met Ala Ala Gly Leu Pro Trp Thr
580 585 590
Ser Arg Thr Ser Pro Thr Val Phe Ser Glu Asp Gln Leu Ser Lys Glu
595 600 605
Lys Val Lys Glu Glu Ile Gly Ser Ser Trp Ile Glu Thr Pro Pro Ser
610 615 620
Ile Lys Ser Leu Asp Ser Asn Asp Ser Gly Val Tyr Thr Ser Ala Cys
625 630 635 640
Lys Arg Arg Arg Leu Ser Pro Ser Asn Ser Ser Asn Glu Asn Ser Pro
645 650 655
Ser Ile Lys Cys Glu Asp Ile Asn Ala Glu Glu Tyr Ser Lys Asp Thr
660 665 670
Ser Lys Gly Met Gly Gly Tyr Tyr Ala Phe Tyr Thr Thr Pro
675 680 685
<210> 7
<211> 162
<212> PRT
<213> 智人(Homo sapiens)
<400> 7
Met Arg Ile Ser Lys Pro His Leu Arg Ser Ile Ser Ile Gln Cys Tyr
1 5 10 15
Leu Cys Leu Leu Leu Asn Ser His Phe Leu Thr Glu Ala Gly Ile His
20 25 30
Val Phe Ile Leu Gly Cys Phe Ser Ala Gly Leu Pro Lys Thr Glu Ala
35 40 45
Asn Trp Val Asn Val Ile Ser Asp Leu Lys Lys Ile Glu Asp Leu Ile
50 55 60
Gln Ser Met His Ile Asp Ala Thr Leu Tyr Thr Glu Ser Asp Val His
65 70 75 80
Pro Ser Cys Lys Val Thr Ala Met Lys Cys Phe Leu Leu Glu Leu Gln
85 90 95
Val Ile Ser Leu Glu Ser Gly Asp Ala Ser Ile His Asp Thr Val Glu
100 105 110
Asn Leu Ile Ile Leu Ala Asn Asn Ser Leu Ser Ser Asn Gly Asn Val
115 120 125
Thr Glu Ser Gly Cys Lys Glu Cys Glu Glu Leu Glu Glu Lys Asn Ile
130 135 140
Lys Glu Phe Leu Gln Ser Phe Val His Ile Val Gln Met Phe Ile Asn
145 150 155 160
Thr Ser
<210> 8
<211> 270
<212> PRT
<213> 智人(Homo sapiens)
<400> 8
Met Lys Pro Lys Met Lys Tyr Ser Thr Asn Lys Ile Ser Thr Ala Lys
1 5 10 15
Trp Lys Asn Thr Ala Ser Lys Ala Leu Cys Phe Lys Leu Gly Lys Ser
20 25 30
Gln Gln Lys Ala Lys Glu Val Cys Pro Met Tyr Phe Met Lys Leu Arg
35 40 45
Ser Gly Leu Met Ile Lys Lys Glu Ala Cys Tyr Phe Arg Arg Glu Thr
50 55 60
Thr Lys Arg Pro Ser Leu Lys Thr Gly Arg Lys His Lys Arg His Leu
65 70 75 80
Val Leu Ala Ala Cys Gln Gln Gln Ser Thr Val Glu Cys Phe Ala Phe
85 90 95
Gly Ile Ser Gly Val Gln Lys Tyr Thr Arg Ala Leu His Asp Ser Ser
100 105 110
Ile Thr Gly Ile Ser Pro Ile Thr Glu Tyr Leu Ala Ser Leu Ser Thr
115 120 125
Tyr Asn Asp Gln Ser Ile Thr Phe Ala Leu Glu Asp Glu Ser Tyr Glu
130 135 140
Ile Tyr Val Glu Asp Leu Lys Lys Asp Glu Lys Lys Asp Lys Val Leu
145 150 155 160
Leu Ser Tyr Tyr Glu Ser Gln His Pro Ser Asn Glu Ser Gly Asp Gly
165 170 175
Val Asp Gly Lys Met Leu Met Val Thr Leu Ser Pro Thr Lys Asp Phe
180 185 190
Trp Leu His Ala Asn Asn Lys Glu His Ser Val Glu Leu His Lys Cys
195 200 205
Glu Lys Pro Leu Pro Asp Gln Ala Phe Phe Val Leu His Asn Met His
210 215 220
Ser Asn Cys Val Ser Phe Glu Cys Lys Thr Asp Pro Gly Val Phe Ile
225 230 235 240
Gly Val Lys Asp Asn His Leu Ala Leu Ile Lys Val Asp Ser Ser Glu
245 250 255
Asn Leu Cys Thr Glu Asn Ile Leu Phe Lys Leu Ser Glu Thr
260 265 270
<210> 9
<211> 524
<212> PRT
<213> 智人(Homo sapiens)
<400> 9
Met Lys Glu Glu Ser Ala Ala Gln Leu Gly Cys Cys His Arg Pro Met
1 5 10 15
Ala Leu Gly Gly Thr Gly Gly Ser Leu Ser Pro Ser Leu Asp Phe Gln
20 25 30
Leu Phe Arg Gly Asp Gln Val Phe Ser Ala Cys Arg Pro Leu Pro Asp
35 40 45
Met Val Asp Ala His Gly Pro Ser Cys Ala Ser Trp Leu Cys Pro Leu
50 55 60
Pro Leu Ala Pro Gly Arg Ser Ala Leu Leu Ala Cys Leu Gln Asp Leu
65 70 75 80
Asp Leu Asn Leu Cys Thr Pro Gln Pro Ala Pro Leu Gly Thr Asp Leu
85 90 95
Gln Gly Leu Gln Glu Asp Ala Leu Ser Met Lys His Glu Pro Pro Gly
100 105 110
Leu Gln Ala Ser Ser Thr Asp Asp Lys Lys Phe Thr Val Lys Tyr Pro
115 120 125
Gln Asn Lys Asp Lys Leu Gly Lys Gln Pro Glu Arg Ala Gly Glu Gly
130 135 140
Ala Pro Cys Pro Ala Phe Ser Ser His Asn Ser Ser Ser Pro Pro Pro
145 150 155 160
Leu Gln Asn Arg Lys Ser Pro Ser Pro Leu Ala Phe Cys Pro Cys Pro
165 170 175
Pro Val Asn Ser Ile Ser Lys Glu Leu Pro Phe Leu Leu His Ala Phe
180 185 190
Tyr Pro Gly Tyr Pro Leu Leu Leu Pro Pro Pro His Leu Phe Thr Tyr
195 200 205
Gly Ala Leu Pro Ser Asp Gln Cys Pro His Leu Leu Met Leu Pro Gln
210 215 220
Asp Pro Ser Tyr Pro Thr Met Ala Met Pro Ser Leu Leu Met Met Val
225 230 235 240
Asn Glu Leu Gly His Pro Ser Ala Arg Trp Glu Thr Leu Leu Pro Tyr
245 250 255
Pro Gly Ala Phe Gln Ala Ser Gly Gln Ala Leu Pro Ser Gln Ala Arg
260 265 270
Asn Pro Gly Ala Gly Ala Ala Pro Thr Asp Ser Pro Gly Leu Glu Arg
275 280 285
Gly Gly Met Ala Ser Pro Ala Lys Arg Val Pro Leu Ser Ser Gln Thr
290 295 300
Gly Thr Ala Ala Leu Pro Tyr Pro Leu Lys Lys Lys Asn Gly Lys Ile
305 310 315 320
Leu Tyr Glu Cys Asn Ile Cys Gly Lys Ser Phe Gly Gln Leu Ser Asn
325 330 335
Leu Lys Val His Leu Arg Val His Ser Gly Glu Arg Pro Phe Gln Cys
340 345 350
Ala Leu Cys Gln Lys Ser Phe Thr Gln Leu Ala His Leu Gln Lys His
355 360 365
His Leu Val His Thr Gly Glu Arg Pro His Lys Cys Ser Ile Pro Trp
370 375 380
Val Pro Gly Arg Asn His Trp Lys Ser Phe Gln Ala Trp Arg Glu Arg
385 390 395 400
Glu Val Cys His Lys Arg Phe Ser Ser Ser Ser Asn Leu Lys Thr His
405 410 415
Leu Arg Leu His Ser Gly Ala Arg Pro Phe Gln Cys Ser Val Cys Arg
420 425 430
Ser Arg Phe Thr Gln His Ile His Leu Lys Leu His His Arg Leu His
435 440 445
Ala Pro Gln Pro Cys Gly Leu Val His Thr Gln Leu Pro Leu Ala Ser
450 455 460
Leu Ala Cys Leu Ala Gln Trp His Gln Gly Ala Leu Asp Leu Met Ala
465 470 475 480
Val Ala Ser Glu Lys His Met Gly Tyr Asp Ile Asp Glu Val Lys Val
485 490 495
Ser Ser Thr Ser Gln Gly Lys Ala Arg Ala Val Ser Leu Ser Ser Ala
500 505 510
Gly Thr Pro Leu Val Met Gly Gln Asp Gln Asn Asn
515 520
<210> 10
<211> 848
<212> PRT
<213> 智人(Homo sapiens)
<400> 10
Met Asn Ser Ser Ser Ala Asn Ile Thr Tyr Ala Ser Arg Lys Arg Arg
1 5 10 15
Lys Pro Val Gln Lys Thr Val Lys Pro Ile Pro Ala Glu Gly Ile Lys
20 25 30
Ser Asn Pro Ser Lys Arg His Arg Asp Arg Leu Asn Thr Glu Leu Asp
35 40 45
Arg Leu Ala Ser Leu Leu Pro Phe Pro Gln Asp Val Ile Asn Lys Leu
50 55 60
Asp Lys Leu Ser Val Leu Arg Leu Ser Val Ser Tyr Leu Arg Ala Lys
65 70 75 80
Ser Phe Phe Asp Val Ala Leu Lys Ser Ser Pro Thr Glu Arg Asn Gly
85 90 95
Gly Gln Asp Asn Cys Arg Ala Ala Asn Phe Arg Glu Gly Leu Asn Leu
100 105 110
Gln Glu Gly Glu Phe Leu Leu Gln Ala Leu Asn Gly Phe Val Leu Val
115 120 125
Val Thr Thr Asp Ala Leu Val Phe Tyr Ala Ser Ser Thr Ile Gln Asp
130 135 140
Tyr Leu Gly Phe Gln Gln Ser Asp Val Ile His Gln Ser Val Tyr Glu
145 150 155 160
Leu Ile His Thr Glu Asp Arg Ala Glu Phe Gln Arg Gln Leu His Trp
165 170 175
Ala Leu Asn Pro Ser Gln Cys Thr Glu Ser Gly Gln Gly Ile Glu Glu
180 185 190
Ala Thr Gly Leu Pro Gln Thr Val Val Cys Tyr Asn Pro Asp Gln Ile
195 200 205
Pro Pro Glu Asn Ser Pro Leu Met Glu Arg Cys Phe Ile Cys Arg Leu
210 215 220
Arg Cys Leu Leu Asp Asn Ser Ser Gly Phe Leu Ala Met Asn Phe Gln
225 230 235 240
Gly Lys Leu Lys Tyr Leu His Gly Gln Lys Lys Lys Gly Lys Asp Gly
245 250 255
Ser Ile Leu Pro Pro Gln Leu Ala Leu Phe Ala Ile Ala Thr Pro Leu
260 265 270
Gln Pro Pro Ser Ile Leu Glu Ile Arg Thr Lys Asn Phe Ile Phe Arg
275 280 285
Thr Lys His Lys Leu Asp Phe Thr Pro Ile Gly Cys Asp Ala Lys Gly
290 295 300
Arg Ile Val Leu Gly Tyr Thr Glu Ala Glu Leu Cys Thr Arg Gly Ser
305 310 315 320
Gly Tyr Gln Phe Ile His Ala Ala Asp Met Leu Tyr Cys Ala Glu Ser
325 330 335
His Ile Arg Met Ile Lys Thr Gly Glu Ser Gly Met Ile Val Phe Arg
340 345 350
Leu Leu Thr Lys Asn Asn Arg Trp Thr Trp Val Gln Ser Asn Ala Arg
355 360 365
Leu Leu Tyr Lys Asn Gly Arg Pro Asp Tyr Ile Ile Val Thr Gln Arg
370 375 380
Pro Leu Thr Asp Glu Glu Gly Thr Glu His Leu Arg Lys Arg Asn Thr
385 390 395 400
Lys Leu Pro Phe Met Phe Thr Thr Gly Glu Ala Val Leu Tyr Glu Ala
405 410 415
Thr Asn Pro Phe Pro Ala Ile Met Asp Pro Leu Pro Leu Arg Thr Lys
420 425 430
Asn Gly Thr Ser Gly Lys Asp Ser Ala Thr Thr Ser Thr Leu Ser Lys
435 440 445
Asp Ser Leu Asn Pro Ser Ser Leu Leu Ala Ala Met Met Gln Gln Asp
450 455 460
Glu Ser Ile Tyr Leu Tyr Pro Ala Ser Ser Thr Ser Ser Thr Ala Pro
465 470 475 480
Phe Glu Asn Asn Phe Phe Asn Glu Ser Met Asn Glu Cys Arg Asn Trp
485 490 495
Gln Asp Asn Thr Ala Pro Met Gly Asn Asp Thr Ile Leu Lys His Glu
500 505 510
Gln Ile Asp Gln Pro Gln Asp Val Asn Ser Phe Ala Gly Gly His Pro
515 520 525
Gly Leu Phe Gln Asp Ser Lys Asn Ser Asp Leu Tyr Ser Ile Met Lys
530 535 540
Asn Leu Gly Ile Asp Phe Glu Asp Ile Arg His Met Gln Asn Glu Lys
545 550 555 560
Phe Phe Arg Asn Asp Phe Ser Gly Glu Val Asp Phe Arg Asp Ile Asp
565 570 575
Leu Thr Asp Glu Ile Leu Thr Tyr Val Gln Asp Ser Leu Ser Lys Ser
580 585 590
Pro Phe Ile Pro Ser Asp Tyr Gln Gln Gln Gln Ser Leu Ala Leu Asn
595 600 605
Ser Ser Cys Met Val Gln Glu His Leu His Leu Glu Gln Gln Gln Gln
610 615 620
His His Gln Lys Gln Val Val Val Glu Pro Gln Gln Gln Leu Cys Gln
625 630 635 640
Lys Met Lys His Met Gln Val Asn Gly Met Phe Glu Asn Trp Asn Ser
645 650 655
Asn Gln Phe Val Pro Phe Asn Cys Pro Gln Gln Asp Pro Gln Gln Tyr
660 665 670
Asn Val Phe Thr Asp Leu His Gly Ile Ser Gln Glu Phe Pro Tyr Lys
675 680 685
Ser Glu Met Asp Ser Met Pro Tyr Thr Gln Asn Phe Ile Ser Cys Asn
690 695 700
Gln Pro Val Leu Pro Gln His Ser Lys Cys Thr Glu Leu Asp Tyr Pro
705 710 715 720
Met Gly Ser Phe Glu Pro Ser Pro Tyr Pro Thr Thr Ser Ser Leu Glu
725 730 735
Asp Phe Val Thr Cys Leu Gln Leu Pro Glu Asn Gln Lys His Gly Leu
740 745 750
Asn Pro Gln Ser Ala Ile Ile Thr Pro Gln Thr Cys Tyr Ala Gly Ala
755 760 765
Val Ser Met Tyr Gln Cys Gln Pro Glu Pro Gln His Thr His Val Gly
770 775 780
Gln Met Gln Tyr Asn Pro Val Leu Pro Gly Gln Gln Ala Phe Leu Asn
785 790 795 800
Lys Phe Gln Asn Gly Val Leu Asn Glu Thr Tyr Pro Ala Glu Leu Asn
805 810 815
Asn Ile Asn Asn Thr Gln Thr Thr Thr His Leu Gln Pro Leu His His
820 825 830
Pro Ser Glu Ala Arg Pro Phe Pro Asp Leu Thr Ser Ser Gly Phe Leu
835 840 845
<210> 11
<211> 271
<212> PRT
<213> 智人(Homo sapiens)
<400> 11
Met Ala Lys Val Pro Asp Met Phe Glu Asp Leu Lys Asn Cys Tyr Ser
1 5 10 15
Glu Asn Glu Glu Asp Ser Ser Ser Ile Asp His Leu Ser Leu Asn Gln
20 25 30
Lys Ser Phe Tyr His Val Ser Tyr Gly Pro Leu His Glu Gly Cys Met
35 40 45
Asp Gln Ser Val Ser Leu Ser Ile Ser Glu Thr Ser Lys Thr Ser Lys
50 55 60
Leu Thr Phe Lys Glu Ser Met Val Val Val Ala Thr Asn Gly Lys Val
65 70 75 80
Leu Lys Lys Arg Arg Leu Ser Leu Ser Gln Ser Ile Thr Asp Asp Asp
85 90 95
Leu Glu Ala Ile Ala Asn Asp Ser Glu Glu Glu Ile Ile Lys Pro Arg
100 105 110
Ser Ala Pro Phe Ser Phe Leu Ser Asn Val Lys Tyr Asn Phe Met Arg
115 120 125
Ile Ile Lys Tyr Glu Phe Ile Leu Asn Asp Ala Leu Asn Gln Ser Ile
130 135 140
Ile Arg Ala Asn Asp Gln Tyr Leu Thr Ala Ala Ala Leu His Asn Leu
145 150 155 160
Asp Glu Ala Val Lys Phe Asp Met Gly Ala Tyr Lys Ser Ser Lys Asp
165 170 175
Asp Ala Lys Ile Thr Val Ile Leu Arg Ile Ser Lys Thr Gln Leu Tyr
180 185 190
Val Thr Ala Gln Asp Glu Asp Gln Pro Val Leu Leu Lys Glu Met Pro
195 200 205
Glu Ile Pro Lys Thr Ile Thr Gly Ser Glu Thr Asn Leu Leu Phe Phe
210 215 220
Trp Glu Thr His Gly Thr Lys Asn Tyr Phe Thr Ser Val Ala His Pro
225 230 235 240
Asn Leu Phe Ile Ala Thr Lys Gln Asp Tyr Trp Val Cys Leu Ala Gly
245 250 255
Gly Pro Pro Ser Ile Thr Asp Phe Gln Ile Leu Glu Asn Gln Ala
260 265 270
<210> 12
<211> 269
<212> PRT
<213> 智人(Homo sapiens)
<400> 12
Met Ala Glu Val Pro Glu Leu Ala Ser Glu Met Met Ala Tyr Tyr Ser
1 5 10 15
Gly Asn Glu Asp Asp Leu Phe Phe Glu Ala Asp Gly Pro Lys Gln Met
20 25 30
Lys Cys Ser Phe Gln Asp Leu Asp Leu Cys Pro Leu Asp Gly Gly Ile
35 40 45
Gln Leu Arg Ile Ser Asp His His Tyr Ser Lys Gly Phe Arg Gln Ala
50 55 60
Ala Ser Val Val Val Ala Met Asp Lys Leu Arg Lys Met Leu Val Pro
65 70 75 80
Cys Pro Gln Thr Phe Gln Glu Asn Asp Leu Ser Thr Phe Phe Pro Phe
85 90 95
Ile Phe Glu Glu Glu Pro Ile Phe Phe Asp Thr Trp Asp Asn Glu Ala
100 105 110
Tyr Val His Asp Ala Pro Val Arg Ser Leu Asn Cys Thr Leu Arg Asp
115 120 125
Ser Gln Gln Lys Ser Leu Val Met Ser Gly Pro Tyr Glu Leu Lys Ala
130 135 140
Leu His Leu Gln Gly Gln Asp Met Glu Gln Gln Val Val Phe Ser Met
145 150 155 160
Ser Phe Val Gln Gly Glu Glu Ser Asn Asp Lys Ile Pro Val Ala Leu
165 170 175
Gly Leu Lys Glu Lys Asn Leu Tyr Leu Ser Cys Val Leu Lys Asp Asp
180 185 190
Lys Pro Thr Leu Gln Leu Glu Ser Val Asp Pro Lys Asn Tyr Pro Lys
195 200 205
Lys Lys Met Glu Lys Arg Phe Val Phe Asn Lys Ile Glu Ile Asn Asn
210 215 220
Lys Leu Glu Phe Glu Ser Ala Gln Phe Pro Asn Trp Tyr Ile Ser Thr
225 230 235 240
Ser Gln Ala Glu Asn Met Pro Val Phe Leu Gly Gly Thr Lys Gly Gly
245 250 255
Gln Asp Ile Thr Asp Phe Thr Met Gln Phe Val Ser Ser
260 265
<210> 13
<211> 193
<212> PRT
<213> 智人(Homo sapiens)
<400> 13
Met Ala Ala Glu Pro Val Glu Asp Asn Cys Ile Asn Phe Val Ala Met
1 5 10 15
Lys Phe Ile Asp Asn Thr Leu Tyr Phe Ile Ala Glu Asp Asp Glu Asn
20 25 30
Leu Glu Ser Asp Tyr Phe Gly Lys Leu Glu Ser Lys Leu Ser Val Ile
35 40 45
Arg Asn Leu Asn Asp Gln Val Leu Phe Ile Asp Gln Gly Asn Arg Pro
50 55 60
Leu Phe Glu Asp Met Thr Asp Ser Asp Cys Arg Asp Asn Ala Pro Arg
65 70 75 80
Thr Ile Phe Ile Ile Ser Met Tyr Lys Asp Ser Gln Pro Arg Gly Met
85 90 95
Ala Val Thr Ile Ser Val Lys Cys Glu Lys Ile Ser Thr Leu Ser Cys
100 105 110
Glu Asn Lys Ile Ile Ser Phe Lys Glu Met Asn Pro Pro Asp Asn Ile
115 120 125
Lys Asp Thr Lys Ser Asp Ile Ile Phe Phe Gln Arg Ser Val Pro Gly
130 135 140
His Asp Asn Lys Met Gln Phe Glu Ser Ser Ser Tyr Glu Gly Tyr Phe
145 150 155 160
Leu Ala Cys Glu Lys Glu Arg Asp Leu Phe Lys Leu Ile Leu Lys Lys
165 170 175
Glu Asp Glu Leu Gly Asp Arg Ser Ile Met Phe Thr Val Gln Asn Glu
180 185 190
Asp
<210> 14
<211> 414
<212> PRT
<213> 智人(Homo sapiens)
<400> 14
Met His Tyr Cys Val Leu Ser Ala Phe Leu Ile Leu His Leu Val Thr
1 5 10 15
Val Ala Leu Ser Leu Ser Thr Cys Ser Thr Leu Asp Met Asp Gln Phe
20 25 30
Met Arg Lys Arg Ile Glu Ala Ile Arg Gly Gln Ile Leu Ser Lys Leu
35 40 45
Lys Leu Thr Ser Pro Pro Glu Asp Tyr Pro Glu Pro Glu Glu Val Pro
50 55 60
Pro Glu Val Ile Ser Ile Tyr Asn Ser Thr Arg Asp Leu Leu Gln Glu
65 70 75 80
Lys Ala Ser Arg Arg Ala Ala Ala Cys Glu Arg Glu Arg Ser Asp Glu
85 90 95
Glu Tyr Tyr Ala Lys Glu Val Tyr Lys Ile Asp Met Pro Pro Phe Phe
100 105 110
Pro Ser Glu Asn Ala Ile Pro Pro Thr Phe Tyr Arg Pro Tyr Phe Arg
115 120 125
Ile Val Arg Phe Asp Val Ser Ala Met Glu Lys Asn Ala Ser Asn Leu
130 135 140
Val Lys Ala Glu Phe Arg Val Phe Arg Leu Gln Asn Pro Lys Ala Arg
145 150 155 160
Val Pro Glu Gln Arg Ile Glu Leu Tyr Gln Ile Leu Lys Ser Lys Asp
165 170 175
Leu Thr Ser Pro Thr Gln Arg Tyr Ile Asp Ser Lys Val Val Lys Thr
180 185 190
Arg Ala Glu Gly Glu Trp Leu Ser Phe Asp Val Thr Asp Ala Val His
195 200 205
Glu Trp Leu His His Lys Asp Arg Asn Leu Gly Phe Lys Ile Ser Leu
210 215 220
His Cys Pro Cys Cys Thr Phe Val Pro Ser Asn Asn Tyr Ile Ile Pro
225 230 235 240
Asn Lys Ser Glu Glu Leu Glu Ala Arg Phe Ala Gly Ile Asp Gly Thr
245 250 255
Ser Thr Tyr Thr Ser Gly Asp Gln Lys Thr Ile Lys Ser Thr Arg Lys
260 265 270
Lys Asn Ser Gly Lys Thr Pro His Leu Leu Leu Met Leu Leu Pro Ser
275 280 285
Tyr Arg Leu Glu Ser Gln Gln Thr Asn Arg Arg Lys Lys Arg Ala Leu
290 295 300
Asp Ala Ala Tyr Cys Phe Arg Asn Val Gln Asp Asn Cys Cys Leu Arg
305 310 315 320
Pro Leu Tyr Ile Asp Phe Lys Arg Asp Leu Gly Trp Lys Trp Ile His
325 330 335
Glu Pro Lys Gly Tyr Asn Ala Asn Phe Cys Ala Gly Ala Cys Pro Tyr
340 345 350
Leu Trp Ser Ser Asp Thr Gln His Ser Arg Val Leu Ser Leu Tyr Asn
355 360 365
Thr Ile Asn Pro Glu Ala Ser Ala Ser Pro Cys Cys Val Ser Gln Asp
370 375 380
Leu Glu Pro Leu Thr Ile Leu Tyr Tyr Ile Gly Lys Thr Pro Lys Ile
385 390 395 400
Glu Gln Leu Ser Asn Met Ile Val Lys Ser Cys Lys Cys Ser
405 410
<210> 15
<211> 162
<212> PRT
<213> 智人(Homo sapiens)
<400> 15
Met Arg Ser Ser Pro Gly Asn Met Glu Arg Ile Val Ile Cys Leu Met
1 5 10 15
Val Ile Phe Leu Gly Thr Leu Val His Lys Ser Ser Ser Gln Gly Gln
20 25 30
Asp Arg His Met Ile Arg Met Arg Gln Leu Ile Asp Ile Val Asp Gln
35 40 45
Leu Lys Asn Tyr Val Asn Asp Leu Val Pro Glu Phe Leu Pro Ala Pro
50 55 60
Glu Asp Val Glu Thr Asn Cys Glu Trp Ser Ala Phe Ser Cys Phe Gln
65 70 75 80
Lys Ala Gln Leu Lys Ser Ala Asn Thr Gly Asn Asn Glu Arg Ile Ile
85 90 95
Asn Val Ser Ile Lys Lys Leu Lys Arg Lys Pro Pro Ser Thr Asn Ala
100 105 110
Gly Arg Arg Gln Lys His Arg Leu Thr Cys Pro Ser Cys Asp Ser Tyr
115 120 125
Glu Lys Lys Pro Pro Lys Glu Phe Leu Glu Arg Phe Lys Ser Leu Leu
130 135 140
Gln Lys Met Ile His Gln His Leu Ser Ser Arg Thr His Gly Ser Glu
145 150 155 160
Asp Ser
<210> 16
<211> 412
<212> PRT
<213> 智人(Homo sapiens)
<400> 16
Met Lys Met His Leu Gln Arg Ala Leu Val Val Leu Ala Leu Leu Asn
1 5 10 15
Phe Ala Thr Val Ser Leu Ser Leu Ser Thr Cys Thr Thr Leu Asp Phe
20 25 30
Gly His Ile Lys Lys Lys Arg Val Glu Ala Ile Arg Gly Gln Ile Leu
35 40 45
Ser Lys Leu Arg Leu Thr Ser Pro Pro Glu Pro Thr Val Met Thr His
50 55 60
Val Pro Tyr Gln Val Leu Ala Leu Tyr Asn Ser Thr Arg Glu Leu Leu
65 70 75 80
Glu Glu Met His Gly Glu Arg Glu Glu Gly Cys Thr Gln Glu Asn Thr
85 90 95
Glu Ser Glu Tyr Tyr Ala Lys Glu Ile His Lys Phe Asp Met Ile Gln
100 105 110
Gly Leu Ala Glu His Asn Glu Leu Ala Val Cys Pro Lys Gly Ile Thr
115 120 125
Ser Lys Val Phe Arg Phe Asn Val Ser Ser Val Glu Lys Asn Arg Thr
130 135 140
Asn Leu Phe Arg Ala Glu Phe Arg Val Leu Arg Val Pro Asn Pro Ser
145 150 155 160
Ser Lys Arg Asn Glu Gln Arg Ile Glu Leu Phe Gln Ile Leu Arg Pro
165 170 175
Asp Glu His Ile Ala Lys Gln Arg Tyr Ile Gly Gly Lys Asn Leu Pro
180 185 190
Thr Arg Gly Thr Ala Glu Trp Leu Ser Phe Asp Val Thr Asp Thr Val
195 200 205
Arg Glu Trp Leu Leu Arg Arg Glu Ser Asn Leu Gly Leu Glu Ile Ser
210 215 220
Ile His Cys Pro Cys His Thr Phe Gln Pro Asn Gly Asp Ile Leu Glu
225 230 235 240
Asn Ile His Glu Val Met Glu Ile Lys Phe Lys Gly Val Asp Asn Glu
245 250 255
Asp Asp His Gly Arg Gly Asp Leu Gly Arg Leu Lys Lys Gln Lys Asp
260 265 270
His His Asn Pro His Leu Ile Leu Met Met Ile Pro Pro His Arg Leu
275 280 285
Asp Asn Pro Gly Gln Gly Gly Gln Arg Lys Lys Arg Ala Leu Asp Thr
290 295 300
Asn Tyr Cys Phe Arg Asn Leu Glu Glu Asn Cys Cys Val Arg Pro Leu
305 310 315 320
Tyr Ile Asp Phe Arg Gln Asp Leu Gly Trp Lys Trp Val His Glu Pro
325 330 335
Lys Gly Tyr Tyr Ala Asn Phe Cys Ser Gly Pro Cys Pro Tyr Leu Arg
340 345 350
Ser Ala Asp Thr Thr His Ser Thr Val Leu Gly Leu Tyr Asn Thr Leu
355 360 365
Asn Pro Glu Ala Ser Ala Ser Pro Cys Cys Val Pro Gln Asp Leu Glu
370 375 380
Pro Leu Thr Ile Leu Tyr Tyr Val Gly Arg Thr Pro Lys Val Glu Gln
385 390 395 400
Leu Ser Asn Met Val Val Lys Ser Cys Lys Cys Ser
405 410
<210> 17
<211> 153
<212> PRT
<213> 智人(Homo sapiens)
<400> 17
Met Tyr Arg Met Gln Leu Leu Ser Cys Ile Ala Leu Ser Leu Ala Leu
1 5 10 15
Val Thr Asn Ser Ala Pro Thr Ser Ser Ser Thr Lys Lys Thr Gln Leu
20 25 30
Gln Leu Glu His Leu Leu Leu Asp Leu Gln Met Ile Leu Asn Gly Ile
35 40 45
Asn Asn Tyr Lys Asn Pro Lys Leu Thr Arg Met Leu Thr Phe Lys Phe
50 55 60
Tyr Met Pro Lys Lys Ala Thr Glu Leu Lys His Leu Gln Cys Leu Glu
65 70 75 80
Glu Glu Leu Lys Pro Leu Glu Glu Val Leu Asn Leu Ala Gln Ser Lys
85 90 95
Asn Phe His Leu Arg Pro Arg Asp Leu Ile Ser Asn Ile Asn Val Ile
100 105 110
Val Leu Glu Leu Lys Gly Ser Glu Thr Thr Phe Met Cys Glu Tyr Ala
115 120 125
Asp Glu Thr Ala Thr Ile Val Glu Phe Leu Asn Arg Trp Ile Thr Phe
130 135 140
Cys Gln Ser Ile Ile Ser Thr Leu Thr
145 150
<210> 18
<211> 1306
<212> PRT
<213> 智人(Homo sapiens)
<400> 18
Met Thr Met Tyr Leu Trp Leu Lys Leu Leu Ala Phe Gly Phe Ala Phe
1 5 10 15
Leu Asp Thr Glu Val Phe Val Thr Gly Gln Ser Pro Thr Pro Ser Pro
20 25 30
Thr Gly Leu Thr Thr Ala Lys Met Pro Ser Val Pro Leu Ser Ser Asp
35 40 45
Pro Leu Pro Thr His Thr Thr Ala Phe Ser Pro Ala Ser Thr Phe Glu
50 55 60
Arg Glu Asn Asp Phe Ser Glu Thr Thr Thr Ser Leu Ser Pro Asp Asn
65 70 75 80
Thr Ser Thr Gln Val Ser Pro Asp Ser Leu Asp Asn Ala Ser Ala Phe
85 90 95
Asn Thr Thr Gly Val Ser Ser Val Gln Thr Pro His Leu Pro Thr His
100 105 110
Ala Asp Ser Gln Thr Pro Ser Ala Gly Thr Asp Thr Gln Thr Phe Ser
115 120 125
Gly Ser Ala Ala Asn Ala Lys Leu Asn Pro Thr Pro Gly Ser Asn Ala
130 135 140
Ile Ser Asp Val Pro Gly Glu Arg Ser Thr Ala Ser Thr Phe Pro Thr
145 150 155 160
Asp Pro Val Ser Pro Leu Thr Thr Thr Leu Ser Leu Ala His His Ser
165 170 175
Ser Ala Ala Leu Pro Ala Arg Thr Ser Asn Thr Thr Ile Thr Ala Asn
180 185 190
Thr Ser Asp Ala Tyr Leu Asn Ala Ser Glu Thr Thr Thr Leu Ser Pro
195 200 205
Ser Gly Ser Ala Val Ile Ser Thr Thr Thr Ile Ala Thr Thr Pro Ser
210 215 220
Lys Pro Thr Cys Asp Glu Lys Tyr Ala Asn Ile Thr Val Asp Tyr Leu
225 230 235 240
Tyr Asn Lys Glu Thr Lys Leu Phe Thr Ala Lys Leu Asn Val Asn Glu
245 250 255
Asn Val Glu Cys Gly Asn Asn Thr Cys Thr Asn Asn Glu Val His Asn
260 265 270
Leu Thr Glu Cys Lys Asn Ala Ser Val Ser Ile Ser His Asn Ser Cys
275 280 285
Thr Ala Pro Asp Lys Thr Leu Ile Leu Asp Val Pro Pro Gly Val Glu
290 295 300
Lys Phe Gln Leu His Asp Cys Thr Gln Val Glu Lys Ala Asp Thr Thr
305 310 315 320
Ile Cys Leu Lys Trp Lys Asn Ile Glu Thr Phe Thr Cys Asp Thr Gln
325 330 335
Asn Ile Thr Tyr Arg Phe Gln Cys Gly Asn Met Ile Phe Asp Asn Lys
340 345 350
Glu Ile Lys Leu Glu Asn Leu Glu Pro Glu His Glu Tyr Lys Cys Asp
355 360 365
Ser Glu Ile Leu Tyr Asn Asn His Lys Phe Thr Asn Ala Ser Lys Ile
370 375 380
Ile Lys Thr Asp Phe Gly Ser Pro Gly Glu Pro Gln Ile Ile Phe Cys
385 390 395 400
Arg Ser Glu Ala Ala His Gln Gly Val Ile Thr Trp Asn Pro Pro Gln
405 410 415
Arg Ser Phe His Asn Phe Thr Leu Cys Tyr Ile Lys Glu Thr Glu Lys
420 425 430
Asp Cys Leu Asn Leu Asp Lys Asn Leu Ile Lys Tyr Asp Leu Gln Asn
435 440 445
Leu Lys Pro Tyr Thr Lys Tyr Val Leu Ser Leu His Ala Tyr Ile Ile
450 455 460
Ala Lys Val Gln Arg Asn Gly Ser Ala Ala Met Cys His Phe Thr Thr
465 470 475 480
Lys Ser Ala Pro Pro Ser Gln Val Trp Asn Met Thr Val Ser Met Thr
485 490 495
Ser Asp Asn Ser Met His Val Lys Cys Arg Pro Pro Arg Asp Arg Asn
500 505 510
Gly Pro His Glu Arg Tyr His Leu Glu Val Glu Ala Gly Asn Thr Leu
515 520 525
Val Arg Asn Glu Ser His Lys Asn Cys Asp Phe Arg Val Lys Asp Leu
530 535 540
Gln Tyr Ser Thr Asp Tyr Thr Phe Lys Ala Tyr Phe His Asn Gly Asp
545 550 555 560
Tyr Pro Gly Glu Pro Phe Ile Leu His His Ser Thr Ser Tyr Asn Ser
565 570 575
Lys Ala Leu Ile Ala Phe Leu Ala Phe Leu Ile Ile Val Thr Ser Ile
580 585 590
Ala Leu Leu Val Val Leu Tyr Lys Ile Tyr Asp Leu His Lys Lys Arg
595 600 605
Ser Cys Asn Leu Asp Glu Gln Gln Glu Leu Val Glu Arg Asp Asp Glu
610 615 620
Lys Gln Leu Met Asn Val Glu Pro Ile His Ala Asp Ile Leu Leu Glu
625 630 635 640
Thr Tyr Lys Arg Lys Ile Ala Asp Glu Gly Arg Leu Phe Leu Ala Glu
645 650 655
Phe Gln Ser Ile Pro Arg Val Phe Ser Lys Phe Pro Ile Lys Glu Ala
660 665 670
Arg Lys Pro Phe Asn Gln Asn Lys Asn Arg Tyr Val Asp Ile Leu Pro
675 680 685
Tyr Asp Tyr Asn Arg Val Glu Leu Ser Glu Ile Asn Gly Asp Ala Gly
690 695 700
Ser Asn Tyr Ile Asn Ala Ser Tyr Ile Asp Gly Phe Lys Glu Pro Arg
705 710 715 720
Lys Tyr Ile Ala Ala Gln Gly Pro Arg Asp Glu Thr Val Asp Asp Phe
725 730 735
Trp Arg Met Ile Trp Glu Gln Lys Ala Thr Val Ile Val Met Val Thr
740 745 750
Arg Cys Glu Glu Gly Asn Arg Asn Lys Cys Ala Glu Tyr Trp Pro Ser
755 760 765
Met Glu Glu Gly Thr Arg Ala Phe Gly Asp Val Val Val Lys Ile Asn
770 775 780
Gln His Lys Arg Cys Pro Asp Tyr Ile Ile Gln Lys Leu Asn Ile Val
785 790 795 800
Asn Lys Lys Glu Lys Ala Thr Gly Arg Glu Val Thr His Ile Gln Phe
805 810 815
Thr Ser Trp Pro Asp His Gly Val Pro Glu Asp Pro His Leu Leu Leu
820 825 830
Lys Leu Arg Arg Arg Val Asn Ala Phe Ser Asn Phe Phe Ser Gly Pro
835 840 845
Ile Val Val His Cys Ser Ala Gly Val Gly Arg Thr Gly Thr Tyr Ile
850 855 860
Gly Ile Asp Ala Met Leu Glu Gly Leu Glu Ala Glu Asn Lys Val Asp
865 870 875 880
Val Tyr Gly Tyr Val Val Lys Leu Arg Arg Gln Arg Cys Leu Met Val
885 890 895
Gln Val Glu Ala Gln Tyr Ile Leu Ile His Gln Ala Leu Val Glu Tyr
900 905 910
Asn Gln Phe Gly Glu Thr Glu Val Asn Leu Ser Glu Leu His Pro Tyr
915 920 925
Leu His Asn Met Lys Lys Arg Asp Pro Pro Ser Glu Pro Ser Pro Leu
930 935 940
Glu Ala Glu Phe Gln Arg Leu Pro Ser Tyr Arg Ser Trp Arg Thr Gln
945 950 955 960
His Ile Gly Asn Gln Glu Glu Asn Lys Ser Lys Asn Arg Asn Ser Asn
965 970 975
Val Ile Pro Tyr Asp Tyr Asn Arg Val Pro Leu Lys His Glu Leu Glu
980 985 990
Met Ser Lys Glu Ser Glu His Asp Ser Asp Glu Ser Ser Asp Asp Asp
995 1000 1005
Ser Asp Ser Glu Glu Pro Ser Lys Tyr Ile Asn Ala Ser Phe Ile
1010 1015 1020
Met Ser Tyr Trp Lys Pro Glu Val Met Ile Ala Ala Gln Gly Pro
1025 1030 1035
Leu Lys Glu Thr Ile Gly Asp Phe Trp Gln Met Ile Phe Gln Arg
1040 1045 1050
Lys Val Lys Val Ile Val Met Leu Thr Glu Leu Lys His Gly Asp
1055 1060 1065
Gln Glu Ile Cys Ala Gln Tyr Trp Gly Glu Gly Lys Gln Thr Tyr
1070 1075 1080
Gly Asp Ile Glu Val Asp Leu Lys Asp Thr Asp Lys Ser Ser Thr
1085 1090 1095
Tyr Thr Leu Arg Val Phe Glu Leu Arg His Ser Lys Arg Lys Asp
1100 1105 1110
Ser Arg Thr Val Tyr Gln Tyr Gln Tyr Thr Asn Trp Ser Val Glu
1115 1120 1125
Gln Leu Pro Ala Glu Pro Lys Glu Leu Ile Ser Met Ile Gln Val
1130 1135 1140
Val Lys Gln Lys Leu Pro Gln Lys Asn Ser Ser Glu Gly Asn Lys
1145 1150 1155
His His Lys Ser Thr Pro Leu Leu Ile His Cys Arg Asp Gly Ser
1160 1165 1170
Gln Gln Thr Gly Ile Phe Cys Ala Leu Leu Asn Leu Leu Glu Ser
1175 1180 1185
Ala Glu Thr Glu Glu Val Val Asp Ile Phe Gln Val Val Lys Ala
1190 1195 1200
Leu Arg Lys Ala Arg Pro Gly Met Val Ser Thr Phe Glu Gln Tyr
1205 1210 1215
Gln Phe Leu Tyr Asp Val Ile Ala Ser Thr Tyr Pro Ala Gln Asn
1220 1225 1230
Gly Gln Val Lys Lys Asn Asn His Gln Glu Asp Lys Ile Glu Phe
1235 1240 1245
Asp Asn Glu Val Asp Lys Val Lys Gln Asp Ala Asn Cys Val Asn
1250 1255 1260
Pro Leu Gly Ala Pro Glu Lys Leu Pro Glu Ala Lys Glu Gln Ala
1265 1270 1275
Glu Gly Ser Glu Pro Thr Ser Gly Thr Glu Gly Pro Glu His Ser
1280 1285 1290
Val Asn Gly Pro Ala Ser Pro Ala Leu Asn Gln Gly Ser
1295 1300 1305
<210> 19
<211> 1145
<212> PRT
<213> 智人(Homo sapiens)
<400> 19
Met Thr Met Tyr Leu Trp Leu Lys Leu Leu Ala Phe Gly Phe Ala Phe
1 5 10 15
Leu Asp Thr Glu Val Phe Val Thr Gly Gln Ser Pro Thr Pro Ser Pro
20 25 30
Thr Asp Ala Tyr Leu Asn Ala Ser Glu Thr Thr Thr Leu Ser Pro Ser
35 40 45
Gly Ser Ala Val Ile Ser Thr Thr Thr Ile Ala Thr Thr Pro Ser Lys
50 55 60
Pro Thr Cys Asp Glu Lys Tyr Ala Asn Ile Thr Val Asp Tyr Leu Tyr
65 70 75 80
Asn Lys Glu Thr Lys Leu Phe Thr Ala Lys Leu Asn Val Asn Glu Asn
85 90 95
Val Glu Cys Gly Asn Asn Thr Cys Thr Asn Asn Glu Val His Asn Leu
100 105 110
Thr Glu Cys Lys Asn Ala Ser Val Ser Ile Ser His Asn Ser Cys Thr
115 120 125
Ala Pro Asp Lys Thr Leu Ile Leu Asp Val Pro Pro Gly Val Glu Lys
130 135 140
Phe Gln Leu His Asp Cys Thr Gln Val Glu Lys Ala Asp Thr Thr Ile
145 150 155 160
Cys Leu Lys Trp Lys Asn Ile Glu Thr Phe Thr Cys Asp Thr Gln Asn
165 170 175
Ile Thr Tyr Arg Phe Gln Cys Gly Asn Met Ile Phe Asp Asn Lys Glu
180 185 190
Ile Lys Leu Glu Asn Leu Glu Pro Glu His Glu Tyr Lys Cys Asp Ser
195 200 205
Glu Ile Leu Tyr Asn Asn His Lys Phe Thr Asn Ala Ser Lys Ile Ile
210 215 220
Lys Thr Asp Phe Gly Ser Pro Gly Glu Pro Gln Ile Ile Phe Cys Arg
225 230 235 240
Ser Glu Ala Ala His Gln Gly Val Ile Thr Trp Asn Pro Pro Gln Arg
245 250 255
Ser Phe His Asn Phe Thr Leu Cys Tyr Ile Lys Glu Thr Glu Lys Asp
260 265 270
Cys Leu Asn Leu Asp Lys Asn Leu Ile Lys Tyr Asp Leu Gln Asn Leu
275 280 285
Lys Pro Tyr Thr Lys Tyr Val Leu Ser Leu His Ala Tyr Ile Ile Ala
290 295 300
Lys Val Gln Arg Asn Gly Ser Ala Ala Met Cys His Phe Thr Thr Lys
305 310 315 320
Ser Ala Pro Pro Ser Gln Val Trp Asn Met Thr Val Ser Met Thr Ser
325 330 335
Asp Asn Ser Met His Val Lys Cys Arg Pro Pro Arg Asp Arg Asn Gly
340 345 350
Pro His Glu Arg Tyr His Leu Glu Val Glu Ala Gly Asn Thr Leu Val
355 360 365
Arg Asn Glu Ser His Lys Asn Cys Asp Phe Arg Val Lys Asp Leu Gln
370 375 380
Tyr Ser Thr Asp Tyr Thr Phe Lys Ala Tyr Phe His Asn Gly Asp Tyr
385 390 395 400
Pro Gly Glu Pro Phe Ile Leu His His Ser Thr Ser Tyr Asn Ser Lys
405 410 415
Ala Leu Ile Ala Phe Leu Ala Phe Leu Ile Ile Val Thr Ser Ile Ala
420 425 430
Leu Leu Val Val Leu Tyr Lys Ile Tyr Asp Leu His Lys Lys Arg Ser
435 440 445
Cys Asn Leu Asp Glu Gln Gln Glu Leu Val Glu Arg Asp Asp Glu Lys
450 455 460
Gln Leu Met Asn Val Glu Pro Ile His Ala Asp Ile Leu Leu Glu Thr
465 470 475 480
Tyr Lys Arg Lys Ile Ala Asp Glu Gly Arg Leu Phe Leu Ala Glu Phe
485 490 495
Gln Ser Ile Pro Arg Val Phe Ser Lys Phe Pro Ile Lys Glu Ala Arg
500 505 510
Lys Pro Phe Asn Gln Asn Lys Asn Arg Tyr Val Asp Ile Leu Pro Tyr
515 520 525
Asp Tyr Asn Arg Val Glu Leu Ser Glu Ile Asn Gly Asp Ala Gly Ser
530 535 540
Asn Tyr Ile Asn Ala Ser Tyr Ile Asp Gly Phe Lys Glu Pro Arg Lys
545 550 555 560
Tyr Ile Ala Ala Gln Gly Pro Arg Asp Glu Thr Val Asp Asp Phe Trp
565 570 575
Arg Met Ile Trp Glu Gln Lys Ala Thr Val Ile Val Met Val Thr Arg
580 585 590
Cys Glu Glu Gly Asn Arg Asn Lys Cys Ala Glu Tyr Trp Pro Ser Met
595 600 605
Glu Glu Gly Thr Arg Ala Phe Gly Asp Val Val Val Lys Ile Asn Gln
610 615 620
His Lys Arg Cys Pro Asp Tyr Ile Ile Gln Lys Leu Asn Ile Val Asn
625 630 635 640
Lys Lys Glu Lys Ala Thr Gly Arg Glu Val Thr His Ile Gln Phe Thr
645 650 655
Ser Trp Pro Asp His Gly Val Pro Glu Asp Pro His Leu Leu Leu Lys
660 665 670
Leu Arg Arg Arg Val Asn Ala Phe Ser Asn Phe Phe Ser Gly Pro Ile
675 680 685
Val Val His Cys Ser Ala Gly Val Gly Arg Thr Gly Thr Tyr Ile Gly
690 695 700
Ile Asp Ala Met Leu Glu Gly Leu Glu Ala Glu Asn Lys Val Asp Val
705 710 715 720
Tyr Gly Tyr Val Val Lys Leu Arg Arg Gln Arg Cys Leu Met Val Gln
725 730 735
Val Glu Ala Gln Tyr Ile Leu Ile His Gln Ala Leu Val Glu Tyr Asn
740 745 750
Gln Phe Gly Glu Thr Glu Val Asn Leu Ser Glu Leu His Pro Tyr Leu
755 760 765
His Asn Met Lys Lys Arg Asp Pro Pro Ser Glu Pro Ser Pro Leu Glu
770 775 780
Ala Glu Phe Gln Arg Leu Pro Ser Tyr Arg Ser Trp Arg Thr Gln His
785 790 795 800
Ile Gly Asn Gln Glu Glu Asn Lys Ser Lys Asn Arg Asn Ser Asn Val
805 810 815
Ile Pro Tyr Asp Tyr Asn Arg Val Pro Leu Lys His Glu Leu Glu Met
820 825 830
Ser Lys Glu Ser Glu His Asp Ser Asp Glu Ser Ser Asp Asp Asp Ser
835 840 845
Asp Ser Glu Glu Pro Ser Lys Tyr Ile Asn Ala Ser Phe Ile Met Ser
850 855 860
Tyr Trp Lys Pro Glu Val Met Ile Ala Ala Gln Gly Pro Leu Lys Glu
865 870 875 880
Thr Ile Gly Asp Phe Trp Gln Met Ile Phe Gln Arg Lys Val Lys Val
885 890 895
Ile Val Met Leu Thr Glu Leu Lys His Gly Asp Gln Glu Ile Cys Ala
900 905 910
Gln Tyr Trp Gly Glu Gly Lys Gln Thr Tyr Gly Asp Ile Glu Val Asp
915 920 925
Leu Lys Asp Thr Asp Lys Ser Ser Thr Tyr Thr Leu Arg Val Phe Glu
930 935 940
Leu Arg His Ser Lys Arg Lys Asp Ser Arg Thr Val Tyr Gln Tyr Gln
945 950 955 960
Tyr Thr Asn Trp Ser Val Glu Gln Leu Pro Ala Glu Pro Lys Glu Leu
965 970 975
Ile Ser Met Ile Gln Val Val Lys Gln Lys Leu Pro Gln Lys Asn Ser
980 985 990
Ser Glu Gly Asn Lys His His Lys Ser Thr Pro Leu Leu Ile His Cys
995 1000 1005
Arg Asp Gly Ser Gln Gln Thr Gly Ile Phe Cys Ala Leu Leu Asn
1010 1015 1020
Leu Leu Glu Ser Ala Glu Thr Glu Glu Val Val Asp Ile Phe Gln
1025 1030 1035
Val Val Lys Ala Leu Arg Lys Ala Arg Pro Gly Met Val Ser Thr
1040 1045 1050
Phe Glu Gln Tyr Gln Phe Leu Tyr Asp Val Ile Ala Ser Thr Tyr
1055 1060 1065
Pro Ala Gln Asn Gly Gln Val Lys Lys Asn Asn His Gln Glu Asp
1070 1075 1080
Lys Ile Glu Phe Asp Asn Glu Val Asp Lys Val Lys Gln Asp Ala
1085 1090 1095
Asn Cys Val Asn Pro Leu Gly Ala Pro Glu Lys Leu Pro Glu Ala
1100 1105 1110
Lys Glu Gln Ala Glu Gly Ser Glu Pro Thr Ser Gly Thr Glu Gly
1115 1120 1125
Pro Glu His Ser Val Asn Gly Pro Ala Ser Pro Ala Leu Asn Gln
1130 1135 1140
Gly Ser
1145
<210> 20
<211> 378
<212> PRT
<213> 智人(Homo sapiens)
<400> 20
Met Asp Leu Gly Lys Pro Met Lys Ser Val Leu Val Val Ala Leu Leu
1 5 10 15
Val Ile Phe Gln Val Cys Leu Cys Gln Asp Glu Val Thr Asp Asp Tyr
20 25 30
Ile Gly Asp Asn Thr Thr Val Asp Tyr Thr Leu Phe Glu Ser Leu Cys
35 40 45
Ser Lys Lys Asp Val Arg Asn Phe Lys Ala Trp Phe Leu Pro Ile Met
50 55 60
Tyr Ser Ile Ile Cys Phe Val Gly Leu Leu Gly Asn Gly Leu Val Val
65 70 75 80
Leu Thr Tyr Ile Tyr Phe Lys Arg Leu Lys Thr Met Thr Asp Thr Tyr
85 90 95
Leu Leu Asn Leu Ala Val Ala Asp Ile Leu Phe Leu Leu Thr Leu Pro
100 105 110
Phe Trp Ala Tyr Ser Ala Ala Lys Ser Trp Val Phe Gly Val His Phe
115 120 125
Cys Lys Leu Ile Phe Ala Ile Tyr Lys Met Ser Phe Phe Ser Gly Met
130 135 140
Leu Leu Leu Leu Cys Ile Ser Ile Asp Arg Tyr Val Ala Ile Val Gln
145 150 155 160
Ala Val Ser Ala His Arg His Arg Ala Arg Val Leu Leu Ile Ser Lys
165 170 175
Leu Ser Cys Val Gly Ile Trp Ile Leu Ala Thr Val Leu Ser Ile Pro
180 185 190
Glu Leu Leu Tyr Ser Asp Leu Gln Arg Ser Ser Ser Glu Gln Ala Met
195 200 205
Arg Cys Ser Leu Ile Thr Glu His Val Glu Ala Phe Ile Thr Ile Gln
210 215 220
Val Ala Gln Met Val Ile Gly Phe Leu Val Pro Leu Leu Ala Met Ser
225 230 235 240
Phe Cys Tyr Leu Val Ile Ile Arg Thr Leu Leu Gln Ala Arg Asn Phe
245 250 255
Glu Arg Asn Lys Ala Ile Lys Val Ile Ile Ala Val Val Val Val Phe
260 265 270
Ile Val Phe Gln Leu Pro Tyr Asn Gly Val Val Leu Ala Gln Thr Val
275 280 285
Ala Asn Phe Asn Ile Thr Ser Ser Thr Cys Glu Leu Ser Lys Gln Leu
290 295 300
Asn Ile Ala Tyr Asp Val Thr Tyr Ser Leu Ala Cys Val Arg Cys Cys
305 310 315 320
Val Asn Pro Phe Leu Tyr Ala Phe Ile Gly Val Lys Phe Arg Asn Asp
325 330 335
Leu Phe Lys Leu Phe Lys Asp Leu Gly Cys Leu Ser Gln Glu Gln Leu
340 345 350
Arg Gln Trp Ser Ser Cys Arg His Ile Arg Arg Ser Ser Met Ser Val
355 360 365
Glu Ala Glu Thr Thr Thr Thr Phe Ser Pro
370 375
<210> 21
<211> 260
<212> PRT
<213> 智人(Homo sapiens)
<400> 21
Met Ala Arg Pro His Pro Trp Trp Leu Cys Val Leu Gly Thr Leu Val
1 5 10 15
Gly Leu Ser Ala Thr Pro Ala Pro Lys Ser Cys Pro Glu Arg His Tyr
20 25 30
Trp Ala Gln Gly Lys Leu Cys Cys Gln Met Cys Glu Pro Gly Thr Phe
35 40 45
Leu Val Lys Asp Cys Asp Gln His Arg Lys Ala Ala Gln Cys Asp Pro
50 55 60
Cys Ile Pro Gly Val Ser Phe Ser Pro Asp His His Thr Arg Pro His
65 70 75 80
Cys Glu Ser Cys Arg His Cys Asn Ser Gly Leu Leu Val Arg Asn Cys
85 90 95
Thr Ile Thr Ala Asn Ala Glu Cys Ala Cys Arg Asn Gly Trp Gln Cys
100 105 110
Arg Asp Lys Glu Cys Thr Glu Cys Asp Pro Leu Pro Asn Pro Ser Leu
115 120 125
Thr Ala Arg Ser Ser Gln Ala Leu Ser Pro His Pro Gln Pro Thr His
130 135 140
Leu Pro Tyr Val Ser Glu Met Leu Glu Ala Arg Thr Ala Gly His Met
145 150 155 160
Gln Thr Leu Ala Asp Phe Arg Gln Leu Pro Ala Arg Thr Leu Ser Thr
165 170 175
His Trp Pro Pro Gln Arg Ser Leu Cys Ser Ser Asp Phe Ile Arg Ile
180 185 190
Leu Val Ile Phe Ser Gly Met Phe Leu Val Phe Thr Leu Ala Gly Ala
195 200 205
Leu Phe Leu His Gln Arg Arg Lys Tyr Arg Ser Asn Lys Gly Glu Ser
210 215 220
Pro Val Glu Pro Ala Glu Pro Cys His Tyr Ser Cys Pro Arg Glu Glu
225 230 235 240
Glu Gly Ser Thr Ile Pro Ile Gln Glu Asp Tyr Arg Lys Pro Glu Pro
245 250 255
Ala Cys Ser Pro
260
<210> 22
<211> 385
<212> PRT
<213> 智人(Homo sapiens)
<400> 22
Met Gly Cys Arg Arg Thr Arg Glu Gly Pro Ser Lys Ala Met Ile Phe
1 5 10 15
Pro Trp Lys Cys Gln Ser Thr Gln Arg Asp Leu Trp Asn Ile Phe Lys
20 25 30
Leu Trp Gly Trp Thr Met Leu Cys Cys Asp Phe Leu Ala His His Gly
35 40 45
Thr Asp Cys Trp Thr Tyr His Tyr Ser Glu Lys Pro Met Asn Trp Gln
50 55 60
Arg Ala Arg Arg Phe Cys Arg Asp Asn Tyr Thr Asp Leu Val Ala Ile
65 70 75 80
Gln Asn Lys Ala Glu Ile Glu Tyr Leu Glu Lys Thr Leu Pro Phe Ser
85 90 95
Arg Ser Tyr Tyr Trp Ile Gly Ile Arg Lys Ile Gly Gly Ile Trp Thr
100 105 110
Trp Val Gly Thr Asn Lys Ser Leu Thr Glu Glu Ala Glu Asn Trp Gly
115 120 125
Asp Gly Glu Pro Asn Asn Lys Lys Asn Lys Glu Asp Cys Val Glu Ile
130 135 140
Tyr Ile Lys Arg Asn Lys Asp Ala Gly Lys Trp Asn Asp Asp Ala Cys
145 150 155 160
His Lys Leu Lys Ala Ala Leu Cys Tyr Thr Ala Ser Cys Gln Pro Trp
165 170 175
Ser Cys Ser Gly His Gly Glu Cys Val Glu Ile Ile Asn Asn Tyr Thr
180 185 190
Cys Asn Cys Asp Val Gly Tyr Tyr Gly Pro Gln Cys Gln Phe Val Ile
195 200 205
Gln Cys Glu Pro Leu Glu Ala Pro Glu Leu Gly Thr Met Asp Cys Thr
210 215 220
His Pro Leu Gly Asn Phe Ser Phe Ser Ser Gln Cys Ala Phe Ser Cys
225 230 235 240
Ser Glu Gly Thr Asn Leu Thr Gly Ile Glu Glu Thr Thr Cys Gly Pro
245 250 255
Phe Gly Asn Trp Ser Ser Pro Glu Pro Thr Cys Gln Val Ile Gln Cys
260 265 270
Glu Pro Leu Ser Ala Pro Asp Leu Gly Ile Met Asn Cys Ser His Pro
275 280 285
Leu Ala Ser Phe Ser Phe Thr Ser Ala Cys Thr Phe Ile Cys Ser Glu
290 295 300
Gly Thr Glu Leu Ile Gly Lys Lys Lys Thr Ile Cys Glu Ser Ser Gly
305 310 315 320
Ile Trp Ser Asn Pro Ser Pro Ile Cys Gln Lys Leu Asp Lys Ser Phe
325 330 335
Ser Met Ile Lys Glu Gly Asp Tyr Asn Pro Leu Phe Ile Pro Val Ala
340 345 350
Val Met Val Thr Ala Phe Ser Gly Leu Ala Phe Ile Ile Trp Leu Ala
355 360 365
Arg Arg Leu Lys Lys Gly Lys Lys Ser Lys Arg Ser Met Asn Asp Pro
370 375 380
Tyr
385
<210> 23
<211> 431
<212> PRT
<213> 智人(Homo sapiens)
<400> 23
Met Pro Asn Pro Arg Pro Gly Lys Pro Ser Ala Pro Ser Leu Ala Leu
1 5 10 15
Gly Pro Ser Pro Gly Ala Ser Pro Ser Trp Arg Ala Ala Pro Lys Ala
20 25 30
Ser Asp Leu Leu Gly Ala Arg Gly Pro Gly Gly Thr Phe Gln Gly Arg
35 40 45
Asp Leu Arg Gly Gly Ala His Ala Ser Ser Ser Ser Leu Asn Pro Met
50 55 60
Pro Pro Ser Gln Leu Gln Leu Pro Thr Leu Pro Leu Val Met Val Ala
65 70 75 80
Pro Ser Gly Ala Arg Leu Gly Pro Leu Pro His Leu Gln Ala Leu Leu
85 90 95
Gln Asp Arg Pro His Phe Met His Gln Leu Ser Thr Val Asp Ala His
100 105 110
Ala Arg Thr Pro Val Leu Gln Val His Pro Leu Glu Ser Pro Ala Met
115 120 125
Ile Ser Leu Thr Pro Pro Thr Thr Ala Thr Gly Val Phe Ser Leu Lys
130 135 140
Ala Arg Pro Gly Leu Pro Pro Gly Ile Asn Val Ala Ser Leu Glu Trp
145 150 155 160
Val Ser Arg Glu Pro Ala Leu Leu Cys Thr Phe Pro Asn Pro Ser Ala
165 170 175
Pro Arg Lys Asp Ser Thr Leu Ser Ala Val Pro Gln Ser Ser Tyr Pro
180 185 190
Leu Leu Ala Asn Gly Val Cys Lys Trp Pro Gly Cys Glu Lys Val Phe
195 200 205
Glu Glu Pro Glu Asp Phe Leu Lys His Cys Gln Ala Asp His Leu Leu
210 215 220
Asp Glu Lys Gly Arg Ala Gln Cys Leu Leu Gln Arg Glu Met Val Gln
225 230 235 240
Ser Leu Glu Gln Gln Leu Val Leu Glu Lys Glu Lys Leu Ser Ala Met
245 250 255
Gln Ala His Leu Ala Gly Lys Met Ala Leu Thr Lys Ala Ser Ser Val
260 265 270
Ala Ser Ser Asp Lys Gly Ser Cys Cys Ile Val Ala Ala Gly Ser Gln
275 280 285
Gly Pro Val Val Pro Ala Trp Ser Gly Pro Arg Glu Ala Pro Asp Ser
290 295 300
Leu Phe Ala Val Arg Arg His Leu Trp Gly Ser His Gly Asn Ser Thr
305 310 315 320
Phe Pro Glu Phe Leu His Asn Met Asp Tyr Phe Lys Phe His Asn Met
325 330 335
Arg Pro Pro Phe Thr Tyr Ala Thr Leu Ile Arg Trp Ala Ile Leu Glu
340 345 350
Ala Pro Glu Lys Gln Arg Thr Leu Asn Glu Ile Tyr His Trp Phe Thr
355 360 365
Arg Met Phe Ala Phe Phe Arg Asn His Pro Ala Thr Trp Lys Asn Ala
370 375 380
Ile Arg His Asn Leu Ser Leu His Lys Cys Phe Val Arg Val Glu Ser
385 390 395 400
Glu Lys Gly Ala Val Trp Thr Val Asp Glu Leu Glu Phe Arg Lys Lys
405 410 415
Arg Ser Gln Arg Pro Ser Arg Cys Ser Asn Pro Thr Pro Gly Pro
420 425 430
<210> 24
<211> 535
<212> PRT
<213> 智人(Homo sapiens)
<400> 24
Met Gly Ile Val Glu Pro Gly Cys Gly Asp Met Leu Thr Gly Thr Glu
1 5 10 15
Pro Met Pro Gly Ser Asp Glu Gly Arg Ala Pro Gly Ala Asp Pro Gln
20 25 30
His Arg Tyr Phe Tyr Pro Glu Pro Gly Ala Gln Asp Ala Asp Glu Arg
35 40 45
Arg Gly Gly Gly Ser Leu Gly Ser Pro Tyr Pro Gly Gly Ala Leu Val
50 55 60
Pro Ala Pro Pro Ser Arg Phe Leu Gly Ala Tyr Ala Tyr Pro Pro Arg
65 70 75 80
Pro Gln Ala Ala Gly Phe Pro Gly Ala Gly Glu Ser Phe Pro Pro Pro
85 90 95
Ala Asp Ala Glu Gly Tyr Gln Pro Gly Glu Gly Tyr Ala Ala Pro Asp
100 105 110
Pro Arg Ala Gly Leu Tyr Pro Gly Pro Arg Glu Asp Tyr Ala Leu Pro
115 120 125
Ala Gly Leu Glu Val Ser Gly Lys Leu Arg Val Ala Leu Asn Asn His
130 135 140
Leu Leu Trp Ser Lys Phe Asn Gln His Gln Thr Glu Met Ile Ile Thr
145 150 155 160
Lys Gln Gly Arg Arg Met Phe Pro Phe Leu Ser Phe Thr Val Ala Gly
165 170 175
Leu Glu Pro Thr Ser His Tyr Arg Met Phe Val Asp Val Val Leu Val
180 185 190
Asp Gln His His Trp Arg Tyr Gln Ser Gly Lys Trp Val Gln Cys Gly
195 200 205
Lys Ala Glu Gly Ser Met Pro Gly Asn Arg Leu Tyr Val His Pro Asp
210 215 220
Ser Pro Asn Thr Gly Ala His Trp Met Arg Gln Glu Val Ser Phe Gly
225 230 235 240
Lys Leu Lys Leu Thr Asn Asn Lys Gly Ala Ser Asn Asn Val Thr Gln
245 250 255
Met Ile Val Leu Gln Ser Leu His Lys Tyr Gln Pro Arg Leu His Ile
260 265 270
Val Glu Val Asn Asp Gly Glu Pro Glu Ala Ala Cys Asn Ala Ser Asn
275 280 285
Thr His Ile Phe Thr Phe Gln Glu Thr Gln Phe Ile Ala Val Thr Ala
290 295 300
Tyr Gln Asn Ala Glu Ile Thr Gln Leu Lys Ile Asp Asn Asn Pro Phe
305 310 315 320
Ala Lys Gly Phe Arg Glu Asn Phe Glu Ser Met Tyr Thr Ser Val Asp
325 330 335
Thr Ser Ile Pro Ser Pro Pro Gly Pro Asn Cys Gln Phe Leu Gly Gly
340 345 350
Asp His Tyr Ser Pro Leu Leu Pro Asn Gln Tyr Pro Val Pro Ser Arg
355 360 365
Phe Tyr Pro Asp Leu Pro Gly Gln Ala Lys Asp Val Val Pro Gln Ala
370 375 380
Tyr Trp Leu Gly Ala Pro Arg Asp His Ser Tyr Glu Ala Glu Phe Arg
385 390 395 400
Ala Val Ser Met Lys Pro Ala Phe Leu Pro Ser Ala Pro Gly Pro Thr
405 410 415
Met Ser Tyr Tyr Arg Gly Gln Glu Val Leu Ala Pro Gly Ala Gly Trp
420 425 430
Pro Val Ala Pro Gln Tyr Pro Pro Lys Met Gly Pro Ala Ser Trp Phe
435 440 445
Arg Pro Met Arg Thr Leu Pro Met Glu Pro Gly Pro Gly Gly Ser Glu
450 455 460
Gly Arg Gly Pro Glu Asp Gln Gly Pro Pro Leu Val Trp Thr Glu Ile
465 470 475 480
Ala Pro Ile Arg Pro Glu Ser Ser Asp Ser Gly Leu Gly Glu Gly Asp
485 490 495
Ser Lys Arg Arg Arg Val Ser Pro Tyr Pro Ser Ser Gly Asp Ser Ser
500 505 510
Ser Pro Ala Gly Ala Pro Ser Pro Phe Asp Lys Glu Ala Glu Gly Gln
515 520 525
Phe Tyr Asn Tyr Phe Pro Asn
530 535
<210> 25
<211> 415
<212> PRT
<213> 智人(Homo sapiens)
<400> 25
Met Glu Leu Arg Lys Tyr Gly Pro Gly Arg Leu Ala Gly Thr Val Ile
1 5 10 15
Gly Gly Ala Ala Gln Ser Lys Ser Gln Thr Lys Ser Asp Ser Ile Thr
20 25 30
Lys Glu Phe Leu Pro Gly Leu Tyr Thr Ala Pro Ser Ser Pro Phe Pro
35 40 45
Pro Ser Gln Val Ser Asp His Gln Val Leu Asn Asp Ala Glu Val Ala
50 55 60
Ala Leu Leu Glu Asn Phe Ser Ser Ser Tyr Asp Tyr Gly Glu Asn Glu
65 70 75 80
Ser Asp Ser Cys Cys Thr Ser Pro Pro Cys Pro Gln Asp Phe Ser Leu
85 90 95
Asn Phe Asp Arg Ala Phe Leu Pro Ala Leu Tyr Ser Leu Leu Phe Leu
100 105 110
Leu Gly Leu Leu Gly Asn Gly Ala Val Ala Ala Val Leu Leu Ser Arg
115 120 125
Arg Thr Ala Leu Ser Ser Thr Asp Thr Phe Leu Leu His Leu Ala Val
130 135 140
Ala Asp Thr Leu Leu Val Leu Thr Leu Pro Leu Trp Ala Val Asp Ala
145 150 155 160
Ala Val Gln Trp Val Phe Gly Ser Gly Leu Cys Lys Val Ala Gly Ala
165 170 175
Leu Phe Asn Ile Asn Phe Tyr Ala Gly Ala Leu Leu Leu Ala Cys Ile
180 185 190
Ser Phe Asp Arg Tyr Leu Asn Ile Val His Ala Thr Gln Leu Tyr Arg
195 200 205
Arg Gly Pro Pro Ala Arg Val Thr Leu Thr Cys Leu Ala Val Trp Gly
210 215 220
Leu Cys Leu Leu Phe Ala Leu Pro Asp Phe Ile Phe Leu Ser Ala His
225 230 235 240
His Asp Glu Arg Leu Asn Ala Thr His Cys Gln Tyr Asn Phe Pro Gln
245 250 255
Val Gly Arg Thr Ala Leu Arg Val Leu Gln Leu Val Ala Gly Phe Leu
260 265 270
Leu Pro Leu Leu Val Met Ala Tyr Cys Tyr Ala His Ile Leu Ala Val
275 280 285
Leu Leu Val Ser Arg Gly Gln Arg Arg Leu Arg Ala Met Arg Leu Val
290 295 300
Val Val Val Val Val Ala Phe Ala Leu Cys Trp Thr Pro Tyr His Leu
305 310 315 320
Val Val Leu Val Asp Ile Leu Met Asp Leu Gly Ala Leu Ala Arg Asn
325 330 335
Cys Gly Arg Glu Ser Arg Val Asp Val Ala Lys Ser Val Thr Ser Gly
340 345 350
Leu Gly Tyr Met His Cys Cys Leu Asn Pro Leu Leu Tyr Ala Phe Val
355 360 365
Gly Val Lys Phe Arg Glu Arg Met Trp Met Leu Leu Leu Arg Leu Gly
370 375 380
Cys Pro Asn Gln Arg Gly Leu Gln Arg Gln Pro Ser Ser Ser Arg Arg
385 390 395 400
Asp Ser Ser Trp Ser Glu Thr Ser Glu Ala Ser Tyr Ser Gly Leu
405 410 415
<210> 26
<211> 769
<212> PRT
<213> 智人(Homo sapiens)
<400> 26
Met Cys Gly Ser Ala Leu Ala Phe Phe Thr Ala Ala Phe Val Cys Leu
1 5 10 15
Gln Asn Asp Arg Arg Gly Pro Ala Ser Phe Leu Trp Ala Ala Trp Val
20 25 30
Phe Ser Leu Val Leu Gly Leu Gly Gln Gly Glu Asp Asn Arg Cys Ala
35 40 45
Ser Ser Asn Ala Ala Ser Cys Ala Arg Cys Leu Ala Leu Gly Pro Glu
50 55 60
Cys Gly Trp Cys Val Gln Glu Asp Phe Ile Ser Gly Gly Ser Arg Ser
65 70 75 80
Glu Arg Cys Asp Ile Val Ser Asn Leu Ile Ser Lys Gly Cys Ser Val
85 90 95
Asp Ser Ile Glu Tyr Pro Ser Val His Val Ile Ile Pro Thr Glu Asn
100 105 110
Glu Ile Asn Thr Gln Val Thr Pro Gly Glu Val Ser Ile Gln Leu Arg
115 120 125
Pro Gly Ala Glu Ala Asn Phe Met Leu Lys Val His Pro Leu Lys Lys
130 135 140
Tyr Pro Val Asp Leu Tyr Tyr Leu Val Asp Val Ser Ala Ser Met His
145 150 155 160
Asn Asn Ile Glu Lys Leu Asn Ser Val Gly Asn Asp Leu Ser Arg Lys
165 170 175
Met Ala Phe Phe Ser Arg Asp Phe Arg Leu Gly Phe Gly Ser Tyr Val
180 185 190
Asp Lys Thr Val Ser Pro Tyr Ile Ser Ile His Pro Glu Arg Ile His
195 200 205
Asn Gln Cys Ser Asp Tyr Asn Leu Asp Cys Met Pro Pro His Gly Tyr
210 215 220
Ile His Val Leu Ser Leu Thr Glu Asn Ile Thr Glu Phe Glu Lys Ala
225 230 235 240
Val His Arg Gln Lys Ile Ser Gly Asn Ile Asp Thr Pro Glu Gly Gly
245 250 255
Phe Asp Ala Met Leu Gln Ala Ala Val Cys Glu Ser His Ile Gly Trp
260 265 270
Arg Lys Glu Ala Lys Arg Leu Leu Leu Val Met Thr Asp Gln Thr Ser
275 280 285
His Leu Ala Leu Asp Ser Lys Leu Ala Gly Ile Val Val Pro Asn Asp
290 295 300
Gly Asn Cys His Leu Lys Asn Asn Val Tyr Val Lys Ser Thr Thr Met
305 310 315 320
Glu His Pro Ser Leu Gly Gln Leu Ser Glu Lys Leu Ile Asp Asn Asn
325 330 335
Ile Asn Val Ile Phe Ala Val Gln Gly Lys Gln Phe His Trp Tyr Lys
340 345 350
Asp Leu Leu Pro Leu Leu Pro Gly Thr Ile Ala Gly Glu Ile Glu Ser
355 360 365
Lys Ala Ala Asn Leu Asn Asn Leu Val Val Glu Ala Tyr Gln Lys Leu
370 375 380
Ile Ser Glu Val Lys Val Gln Val Glu Asn Gln Val Gln Gly Ile Tyr
385 390 395 400
Phe Asn Ile Thr Ala Ile Cys Pro Asp Gly Ser Arg Lys Pro Gly Met
405 410 415
Glu Gly Cys Arg Asn Val Thr Ser Asn Asp Glu Val Leu Phe Asn Val
420 425 430
Thr Val Thr Met Lys Lys Cys Asp Val Thr Gly Gly Lys Asn Tyr Ala
435 440 445
Ile Ile Lys Pro Ile Gly Phe Asn Glu Thr Ala Lys Ile His Ile His
450 455 460
Arg Asn Cys Ser Cys Gln Cys Glu Asp Asn Arg Gly Pro Lys Gly Lys
465 470 475 480
Cys Val Asp Glu Thr Phe Leu Asp Ser Lys Cys Phe Gln Cys Asp Glu
485 490 495
Asn Lys Cys His Phe Asp Glu Asp Gln Phe Ser Ser Glu Ser Cys Lys
500 505 510
Ser His Lys Asp Gln Pro Val Cys Ser Gly Arg Gly Val Cys Val Cys
515 520 525
Gly Lys Cys Ser Cys His Lys Ile Lys Leu Gly Lys Val Tyr Gly Lys
530 535 540
Tyr Cys Glu Lys Asp Asp Phe Ser Cys Pro Tyr His His Gly Asn Leu
545 550 555 560
Cys Ala Gly His Gly Glu Cys Glu Ala Gly Arg Cys Gln Cys Phe Ser
565 570 575
Gly Trp Glu Gly Asp Arg Cys Gln Cys Pro Ser Ala Ala Ala Gln His
580 585 590
Cys Val Asn Ser Lys Gly Gln Val Cys Ser Gly Arg Gly Thr Cys Val
595 600 605
Cys Gly Arg Cys Glu Cys Thr Asp Pro Arg Ser Ile Gly Arg Phe Cys
610 615 620
Glu His Cys Pro Thr Cys Tyr Thr Ala Cys Lys Glu Asn Trp Asn Cys
625 630 635 640
Met Gln Cys Leu His Pro His Asn Leu Ser Gln Ala Ile Leu Asp Gln
645 650 655
Cys Lys Thr Ser Cys Ala Leu Met Glu Gln Gln His Tyr Val Asp Gln
660 665 670
Thr Ser Glu Cys Phe Ser Ser Pro Ser Tyr Leu Arg Ile Phe Phe Ile
675 680 685
Ile Phe Ile Val Thr Phe Leu Ile Gly Leu Leu Lys Val Leu Ile Ile
690 695 700
Arg Gln Val Ile Leu Gln Trp Asn Ser Asn Lys Ile Lys Ser Ser Ser
705 710 715 720
Asp Tyr Arg Val Ser Ala Ser Lys Lys Asp Lys Leu Ile Leu Gln Ser
725 730 735
Val Cys Thr Arg Ala Val Thr Tyr Arg Arg Glu Lys Pro Glu Glu Ile
740 745 750
Lys Met Asp Ile Ser Lys Leu Asn Ala His Glu Thr Phe Arg Cys Asn
755 760 765
Phe
<210> 27
<211> 1048
<212> PRT
<213> 智人(Homo sapiens)
<400> 27
Met Ala Phe Pro Pro Arg Arg Arg Leu Arg Leu Gly Pro Arg Gly Leu
1 5 10 15
Pro Leu Leu Leu Ser Gly Leu Leu Leu Pro Leu Cys Arg Ala Phe Asn
20 25 30
Leu Asp Val Asp Ser Pro Ala Glu Tyr Ser Gly Pro Glu Gly Ser Tyr
35 40 45
Phe Gly Phe Ala Val Asp Phe Phe Val Pro Ser Ala Ser Ser Arg Met
50 55 60
Phe Leu Leu Val Gly Ala Pro Lys Ala Asn Thr Thr Gln Pro Gly Ile
65 70 75 80
Val Glu Gly Gly Gln Val Leu Lys Cys Asp Trp Ser Ser Thr Arg Arg
85 90 95
Cys Gln Pro Ile Glu Phe Asp Ala Thr Gly Asn Arg Asp Tyr Ala Lys
100 105 110
Asp Asp Pro Leu Glu Phe Lys Ser His Gln Trp Phe Gly Ala Ser Val
115 120 125
Arg Ser Lys Gln Asp Lys Ile Leu Ala Cys Ala Pro Leu Tyr His Trp
130 135 140
Arg Thr Glu Met Lys Gln Glu Arg Glu Pro Val Gly Thr Cys Phe Leu
145 150 155 160
Gln Asp Gly Thr Lys Thr Val Glu Tyr Ala Pro Cys Arg Ser Gln Asp
165 170 175
Ile Asp Ala Asp Gly Gln Gly Phe Cys Gln Gly Gly Phe Ser Ile Asp
180 185 190
Phe Thr Lys Ala Asp Arg Val Leu Leu Gly Gly Pro Gly Ser Phe Tyr
195 200 205
Trp Gln Gly Gln Leu Ile Ser Asp Gln Val Ala Glu Ile Val Ser Lys
210 215 220
Tyr Asp Pro Asn Val Tyr Ser Ile Lys Tyr Asn Asn Gln Leu Ala Thr
225 230 235 240
Arg Thr Ala Gln Ala Ile Phe Asp Asp Ser Tyr Leu Gly Tyr Ser Val
245 250 255
Ala Val Gly Asp Phe Asn Gly Asp Gly Ile Asp Asp Phe Val Ser Gly
260 265 270
Val Pro Arg Ala Ala Arg Thr Leu Gly Met Val Tyr Ile Tyr Asp Gly
275 280 285
Lys Asn Met Ser Ser Leu Tyr Asn Phe Thr Gly Glu Gln Met Ala Ala
290 295 300
Tyr Phe Gly Phe Ser Val Ala Ala Thr Asp Ile Asn Gly Asp Asp Tyr
305 310 315 320
Ala Asp Val Phe Ile Gly Ala Pro Leu Phe Met Asp Arg Gly Ser Asp
325 330 335
Gly Lys Leu Gln Glu Val Gly Gln Val Ser Val Ser Leu Gln Arg Ala
340 345 350
Ser Gly Asp Phe Gln Thr Thr Lys Leu Asn Gly Phe Glu Val Phe Ala
355 360 365
Arg Phe Gly Ser Ala Ile Ala Pro Leu Gly Asp Leu Asp Gln Asp Gly
370 375 380
Phe Asn Asp Ile Ala Ile Ala Ala Pro Tyr Gly Gly Glu Asp Lys Lys
385 390 395 400
Gly Ile Val Tyr Ile Phe Asn Gly Arg Ser Thr Gly Leu Asn Ala Val
405 410 415
Pro Ser Gln Ile Leu Glu Gly Gln Trp Ala Ala Arg Ser Met Pro Pro
420 425 430
Ser Phe Gly Tyr Ser Met Lys Gly Ala Thr Asp Ile Asp Lys Asn Gly
435 440 445
Tyr Pro Asp Leu Ile Val Gly Ala Phe Gly Val Asp Arg Ala Ile Leu
450 455 460
Tyr Arg Ala Arg Pro Val Ile Thr Val Asn Ala Gly Leu Glu Val Tyr
465 470 475 480
Pro Ser Ile Leu Asn Gln Asp Asn Lys Thr Cys Ser Leu Pro Gly Thr
485 490 495
Ala Leu Lys Val Ser Cys Phe Asn Val Arg Phe Cys Leu Lys Ala Asp
500 505 510
Gly Lys Gly Val Leu Pro Arg Lys Leu Asn Phe Gln Val Glu Leu Leu
515 520 525
Leu Asp Lys Leu Lys Gln Lys Gly Ala Ile Arg Arg Ala Leu Phe Leu
530 535 540
Tyr Ser Arg Ser Pro Ser His Ser Lys Asn Met Thr Ile Ser Arg Gly
545 550 555 560
Gly Leu Met Gln Cys Glu Glu Leu Ile Ala Tyr Leu Arg Asp Glu Ser
565 570 575
Glu Phe Arg Asp Lys Leu Thr Pro Ile Thr Ile Phe Met Glu Tyr Arg
580 585 590
Leu Asp Tyr Arg Thr Ala Ala Asp Thr Thr Gly Leu Gln Pro Ile Leu
595 600 605
Asn Gln Phe Thr Pro Ala Asn Ile Ser Arg Gln Ala His Ile Leu Leu
610 615 620
Asp Cys Gly Glu Asp Asn Val Cys Lys Pro Lys Leu Glu Val Ser Val
625 630 635 640
Asp Ser Asp Gln Lys Lys Ile Tyr Ile Gly Asp Asp Asn Pro Leu Thr
645 650 655
Leu Ile Val Lys Ala Gln Asn Gln Gly Glu Gly Ala Tyr Glu Ala Glu
660 665 670
Leu Ile Val Ser Ile Pro Leu Gln Ala Asp Phe Ile Gly Val Val Arg
675 680 685
Asn Asn Glu Ala Leu Ala Arg Leu Ser Cys Ala Phe Lys Thr Glu Asn
690 695 700
Gln Thr Arg Gln Val Val Cys Asp Leu Gly Asn Pro Met Lys Ala Gly
705 710 715 720
Thr Gln Leu Leu Ala Gly Leu Arg Phe Ser Val His Gln Gln Ser Glu
725 730 735
Met Asp Thr Ser Val Lys Phe Asp Leu Gln Ile Gln Ser Ser Asn Leu
740 745 750
Phe Asp Lys Val Ser Pro Val Val Ser His Lys Val Asp Leu Ala Val
755 760 765
Leu Ala Ala Val Glu Ile Arg Gly Val Ser Ser Pro Asp His Val Phe
770 775 780
Leu Pro Ile Pro Asn Trp Glu His Lys Glu Asn Pro Glu Thr Glu Glu
785 790 795 800
Asp Val Gly Pro Val Val Gln His Ile Tyr Glu Leu Arg Asn Asn Gly
805 810 815
Pro Ser Ser Phe Ser Lys Ala Met Leu His Leu Gln Trp Pro Tyr Lys
820 825 830
Tyr Asn Asn Asn Thr Leu Leu Tyr Ile Leu His Tyr Asp Ile Asp Gly
835 840 845
Pro Met Asn Cys Thr Ser Asp Met Glu Ile Asn Pro Leu Arg Ile Lys
850 855 860
Ile Ser Ser Leu Gln Thr Thr Glu Lys Asn Asp Thr Val Ala Gly Gln
865 870 875 880
Gly Glu Arg Asp His Leu Ile Thr Lys Arg Asp Leu Ala Leu Ser Glu
885 890 895
Gly Asp Ile His Thr Leu Gly Cys Gly Val Ala Gln Cys Leu Lys Ile
900 905 910
Val Cys Gln Val Gly Arg Leu Asp Arg Gly Lys Ser Ala Ile Leu Tyr
915 920 925
Val Lys Ser Leu Leu Trp Thr Glu Thr Phe Met Asn Lys Glu Asn Gln
930 935 940
Asn His Ser Tyr Ser Leu Lys Ser Ser Ala Ser Phe Asn Val Ile Glu
945 950 955 960
Phe Pro Tyr Lys Asn Leu Pro Ile Glu Asp Ile Thr Asn Ser Thr Leu
965 970 975
Val Thr Thr Asn Val Thr Trp Gly Ile Gln Pro Ala Pro Met Pro Val
980 985 990
Pro Val Trp Val Ile Ile Leu Ala Val Leu Ala Gly Leu Leu Leu Leu
995 1000 1005
Ala Val Leu Val Phe Val Met Tyr Arg Met Gly Phe Phe Lys Arg
1010 1015 1020
Val Arg Pro Pro Gln Glu Glu Gln Glu Arg Glu Gln Leu Gln Pro
1025 1030 1035
His Glu Asn Gly Glu Gly Asn Ser Glu Thr
1040 1045
<210> 28
<211> 177
<212> PRT
<213> 智人(Homo sapiens)
<400> 28
Met Phe His Val Ser Phe Arg Tyr Ile Phe Gly Leu Pro Pro Leu Ile
1 5 10 15
Leu Val Leu Leu Pro Val Ala Ser Ser Asp Cys Asp Ile Glu Gly Lys
20 25 30
Asp Gly Lys Gln Tyr Glu Ser Val Leu Met Val Ser Ile Asp Gln Leu
35 40 45
Leu Asp Ser Met Lys Glu Ile Gly Ser Asn Cys Leu Asn Asn Glu Phe
50 55 60
Asn Phe Phe Lys Arg His Ile Cys Asp Ala Asn Lys Glu Gly Met Phe
65 70 75 80
Leu Phe Arg Ala Ala Arg Lys Leu Arg Gln Phe Leu Lys Met Asn Ser
85 90 95
Thr Gly Asp Phe Asp Leu His Leu Leu Lys Val Ser Glu Gly Thr Thr
100 105 110
Ile Leu Leu Asn Cys Thr Gly Gln Val Lys Gly Arg Lys Pro Ala Ala
115 120 125
Leu Gly Glu Ala Gln Pro Thr Lys Ser Leu Glu Glu Asn Lys Ser Leu
130 135 140
Lys Glu Gln Lys Lys Leu Asn Asp Leu Cys Phe Leu Lys Arg Leu Leu
145 150 155 160
Gln Glu Ile Lys Thr Cys Trp Asn Lys Ile Leu Met Gly Thr Lys Glu
165 170 175
His
Claims (28)
1.一种用于产生组织驻留记忆T细胞(TRM)的方法,该方法包括在转化生长因子β(TGFβ)的存在下培养淋巴细胞,以及/或者将该淋巴细胞与调节性T细胞共同培养。
2.根据权利要求1所述的方法,其中,所述淋巴细胞在TGFβ的存在下培养,优选地,其中,所述淋巴细胞不与调节性T细胞共同培养。
3.根据权利要求1或2所述的方法,其中,所述淋巴细胞是幼稚CD8+T淋巴细胞、效应CD8+T淋巴细胞或记忆CD8+T淋巴细胞。
4.根据前述权利要求中任意一项所述的方法,其中,所述TGFβ以0.01ng/ml和50ng/ml之间的浓度存在。
5.根据前述权利要求中任意一项所述的方法,其中,所述淋巴细胞从人或非人动物的组织获得,可选地,所述组织选自:血液、脾脏、淋巴结、肺、胃肠道、皮肤、前列腺乳腺组织、肝脏、骨髓和胰腺。
6.根据前述权利要求中任意一项所述的方法,其中,所述TRM的特征在于分化簇8(CD8)、分化簇69(CD69)、Hobit、芳烃受体(AhR)和/或分化簇103(CD103)。
7.根据前述权利要求中任意一项所述的方法,其中,所述TRM的特征在于不存在杀伤细胞凝集素样受体亚家族G成员(KLRG1)和/或脱中胚蛋白(Eomes)的表达。
8.根据前述权利要求中任意一项所述的方法,其中,该方法包括在IL-2、IL-4、IL-7、IL-12、IL-15和/或IL-21存在下培养所述淋巴细胞。
9.根据前述权利要求中任意一项所述的方法,进一步包括在白细胞介素15(IL-15)的存在下培养所述淋巴细胞。
10.根据前述权利要求中任意一项所述的方法,进一步包括在白细胞介素33(IL-33)的存在下培养所述淋巴细胞。
11.根据前述权利要求中任意一项所述的方法,其中,该方法包括在白细胞介素7(IL-7)的存在下培养所述淋巴细胞。
12.根据前述权利要求中任意一项所述的方法,其中,该方法包括在白细胞介素2(IL-2)的存在下培养所述淋巴细胞。
13.根据前述权利要求中任意一项所述的方法,进一步包括在至少一种白细胞介素1家族成员的存在下培养所述淋巴细胞。
14.根据权利要求13所述的方法,其中,所述至少一种白细胞介素1家族成员为IL-1a、IL-1b和/或IL-18。
15.根据前述权利要求中任意一项所述的方法,其中,所述淋巴细胞在包含至少一种芳烃受体(AhR)配体的培养基中培养。
16.根据权利要求15所述的方法,其中,所述AhR配体选自卤代芳烃、多环芳烃、膳食衍生芳烃、血红素代谢物、靛蓝、StemRegenin1和色氨酸代谢物。
17.根据前述权利要求中任意一项所述的方法,其中,所述淋巴细胞在包含至少一种脂质的培养基中培养。
18.根据前述权利要求中任意一项所述的方法,其中,所述淋巴细胞在包含抗原的培养基中培养,可选地,其中,所述抗原为肿瘤抗原。
19.根据前述权利要求中任意一项所述的方法,其中,所述调节性T细胞的特征在于Foxp3的表达,或者缺失。
20.根据前述权利要求中任意一项所述的方法,其中,该方法还包括用树突细胞培养所述淋巴细胞。
21.根据前述权利要求中任意一项所述的方法,其中,该方法还包括扩展组织驻留记忆T细胞(TRM)的群体。
22.根据权利要求21所述的方法,其中,该方法进一步包括在IL-2、IL-4、IL-7、IL-12、IL-15和/或IL-21的存在下培养所述TRM细胞。
23.一种组织驻留记忆T细胞(TRM),通过如权利要求1-22中任意一项所述的方法获得,或者是可通过所述方法获得的。
24.根据权利要求23所述的组织驻留记忆T细胞(TRM),可选地,其扩展的群体,用于治疗。
25.根据权利要求23所述的组织驻留记忆T细胞(TRM),可选地,其扩展的群体,用于T细胞疗法。
26.根据权利要求23所述的组织驻留记忆T细胞(TRM),用于预防、治疗或改善癌症或感染。
27.一种药物组合物,包含如权利要求23所述的组织驻留记忆T细胞,可选地,包含其扩展的群体以及药学上可接受的赋形剂。
28.一种制备如权利要求27所述的药物组合物的方法,该方法包括将有效治疗量的如权利要求23所述的组织驻留记忆T细胞、可选地其扩展的群体,与药学上可接受的赋形剂组合。
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