CN114401730A - 免疫调节组合物和方法 - Google Patents
免疫调节组合物和方法 Download PDFInfo
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- CN114401730A CN114401730A CN202080047260.8A CN202080047260A CN114401730A CN 114401730 A CN114401730 A CN 114401730A CN 202080047260 A CN202080047260 A CN 202080047260A CN 114401730 A CN114401730 A CN 114401730A
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Abstract
本文提供了包含构建体的方法和组合物,所述构建体包括两个或更多个截短的T3SS细菌效应多肽。还提供了包含构建体的药物组合物和基于施用此类构建体治疗炎症性疾病的方法。
Description
相关申请的交叉引用
本申请根据35U.S.C.§119(e)(1)要求2019年5月1日提交的美国临时申请序列号62/841,312的优先权,其内容通过引用并入本文。
序列表
本申请包含序列表,该序列表已以ASCII格式以电子方式提交,并在此以其全文形式通过引用并入本文。所述ASCII副本创建于2020年5月1日,名为G6113-00029_SL.txt,大小为60,849字节。
技术领域
本发明涉及用于治疗炎症性病症的组合物和方法。
背景技术
炎症是生理防御机制,用于识别和去除潜在的有害刺激,例如病原体、刺激物或受损细胞。炎症分为急性或慢性。急性炎症是指身体的即时免疫反应,以帮助防止进一步受伤并促进愈合。急性炎症通常是自限性的。在一些情况下,炎症过程变得持续,导致慢性炎症的发展。慢性炎症会导致慢性疼痛、发红、肿胀、僵硬和对正常组织的损害。慢性炎症与大范围的疾病相关,这些疾病在全球范围内具有显著的发病率和死亡率,例如,关节炎和关节疾病、心血管疾病、过敏症、慢性阻塞性肺病、糖尿病、炎症性肠病和癌症。持续需要治疗炎症性病症的新的和有效的方法。
发明内容
本文公开了包含组合物的构建体,所述组合物包括两种或更多种截短的T3SS细菌效应多肽。本文提供的构建体可以包括两个或更多个截短的T3SS细菌效应多肽,所述截短的T3SS细菌效应多肽包括全长细菌效应多肽YopE、YopJ、YopM、NleE、NleC、NleB、OspZ、IpaH4.5、IpaH7.8和IpaH9.8的一部分。构建体可以进一步包括蛋白质转导结构域。构建体可以配制成用于治疗炎症性病症的药物。
附图说明
本发明的这些和其他特征和优点将在本发明的优选实施方案的以下详细描述中更充分地公开或变得显而易见,其将与附图一起考虑,其中相似的数字表示相似的元件,并且进一步其中:
图1是细菌效应多肽氨基酸和核苷酸序列的列表。
图2是显示包含YopE多肽和OspZ多肽的构建体的图。
图3是显示包含截短的YopM多肽和截短的OspZ多肽的构建体的图。
图4是显示包含截短的YopM多肽和截短的NleC多肽的构建体的图。
图5是显示全长IpaH9.8多肽和全长IpaH4.5多肽的结构域的图。
具体实施方式
优选实施方案的描述旨在结合附图进行阅读,这些附图被认为是本发明的整个书面描述的一部分。附图不一定按比例绘制,并且为了清楚和简洁起见,本发明的某些特征可以按比例放大或以某种示意性的形式显示。在描述中,诸如“水平”、“垂直”、“向上”、“向下”、“顶部”和“底部”等相关术语及其派生词(例如,“水平地”、“向下地”、“向上地”等)应解释为是指当时描述的或正在讨论的附图中所示的方向。这些相对术语是为了便于描述,通常并不旨在需要特定的方向。包括“向内”对“向外”、“纵向”对“横向”等的术语将视情况相对于彼此或相对于伸长轴、或旋转轴或旋转中心来解释。关于附接、耦合等术语,例如“连接的”和“互连的”,是指一种关系,其中结构通过中间结构直接或间接地相互固定或附接,以及可移动或刚性的附接或关系,除非另有明确描述。术语“可操作地连接”是这样的附接、耦合或连接,其允许相关结构凭借该关系如预期那样操作。当仅示出单个机器时,术语“机器”也应被认为包括单独或联合执行一组(或多组)指令以实施本文讨论的任何一种或多种方法的机器的任何集合。在权利要求中,方法加功能条款(如果使用)旨在涵盖由书面描述或附图所描述、建议或显而易见的结构,用于实施所述功能,不仅包括结构等效物,还包括等效结构。
本文公开了用于治疗炎症性病症的组合物和方法。组合物可以包括包含两种或更多种截短的T3SS细菌效应多肽的构建体。在感染过程中,通常通过III型分泌系统(T3SS)将细菌效应多肽注射到宿主细胞中。这些多肽通过靶向宿主炎症信号通路来抑制或禁用宿主免疫反应,允许病原体破坏宿主防御以确保细菌存活。本文公开的组合物和方法具有免疫调节活性,因此可用于治疗炎症。截短的T3SS细菌效应多肽可以提供增强的药代动力学特性和生物利用度,以提高治疗效果。
本文提供的构建体可以包括两个或更多个截短的T3SS细菌效应多肽,所述截短的T3SS细菌效应多肽包括T3SS全长细菌效应多肽的一部分。T3SS全长细菌效应多肽可以具有生物活性,例如E3泛素连接酶活性、RhoGTP酶调节活性、半胱氨酸甲基化酶活性、锌金属蛋白酶活性、乙酰转移酶活性或O-GlcNac转移酶活性。本文提供的构建体可以包括两个或更多个截短的T3SS细菌效应多肽,所述截短的T3SS细菌效应多肽包括全长细菌效应多肽YopE、YopJ、YopM、NleC、NleB、OspZ、IpaH4.5、IpaH7.8和IpaH9.8的一部分。两个或更多个截短的T3SS细菌效应多肽可以相同或不同。示例性构建体可以包括截短的YopE多肽和截短的OspZ多肽;截短的YopM多肽和截短的OspZ多肽;截短的YopM多肽和截短的NleC多肽;截短的YopM多肽和截短的NleB多肽;和截短的IpaH9.8多肽和截短的IpaH4.5多肽。在一些实施方案中,本文提供的构建体可以排除包含全长细菌效应多肽YopE、YopJ、YopM、NleC、NleB、OspZ、IpaH4.5、IpaH7.8和IpaH9.8的一部分的任何一种截短的T3SS细菌效应多肽。
有用的细菌效应多肽可以具有如表1所示的生物化学活性、靶标特异性和细胞效应。
表1:T3SS效应物活性
组合物
本文公开的构建体包括两个或更多个截短的T3SS细菌效应多肽。截短的细菌效应多肽可以是参考的全长多肽的连续或相邻部分(例如,10个氨基酸长度的多肽片段可以是该多肽内的任何10个相邻残基)。因此,截短的T3SS细菌效应多肽可以位于参考的全长多肽内。截短的T3SS细菌效应多肽可以是长度比参考的全长多肽短的至少1、2、3、4、5、6、7、10、15、20、25、30、40、50、60、70、80、90、100或更多个氨基酸残基。在一些实施方案中,相对于参考的全长多肽,截短的T3SS细菌效应多肽的氨基酸序列在C端缺少1、2、3、4、5、6、7、10、15、20、25、30、40、50、60、70、80、90、100或更多个氨基酸残基。在一些实施方案中,相对于参考的全长多肽,截短的T3SS细菌效应多肽的氨基酸序列在N端缺少1、2、3、4、5、6、7、10、15、20、25、30、40、50、60、70、80、90、100或更多个氨基酸残基。
在一些实施方案中,截短的细菌效应多肽保留了参考的全长细菌效应多肽的一种或多种活性。例如,截短的E3泛素连接酶可以保留参考的全长E3泛素连接酶的所有或基本上所有的E3泛素连接酶活性。在一些实施方案中,截短的细菌效应多肽缺乏或基本上缺乏参考的全长细菌效应多肽的一种或多种活性。
参考的全长T3SS细菌效应多肽可以具有如SEQ ID NO.:1;SEQ ID NO.:3;SEQ IDNO.:5;SEQ ID NO.:7;SEQ ID NO.:9;SEQ ID NO.:11;SEQ ID NO.:15;或SEQ ID NO.:21中所示的氨基酸序列。在一些实施方案中,参考的全长T3SS细菌效应多肽可以具有与SEQ IDNO.:1;SEQ ID NO.:3;SEQ ID NO.:5;SEQ ID NO.:7;SEQ ID NO.:9;SEQ ID NO.:11;SEQID NO.:15;或SEQ ID NO.:21中所示的氨基酸序列至少90%相同的氨基酸序列。
构建体可以是包含第一截短的T3SS细菌效应多肽的氨基酸序列和第二截短的细菌效应多肽的氨基酸序列的融合蛋白。在一些实施方案中,第一截短的细菌效应多肽的氨基酸序列和第二截短的细菌效应多肽的氨基酸序列可以是连续的,第一截短的细菌效应多肽的氨基酸序列和第二截短的细菌效应多肽的氨基酸序列通过肽键接合。
在一些实施方案中,第一截短的细菌效应多肽的氨基酸序列和第二截短的细菌效应多肽的氨基酸序列通过接头连接。接头可以是可切割的接头。可切割的接头可以包括pH敏感性接头,例如腙;基于氨基磷酸酯的接头,硫代马来酸;蛋白酶体特异性接头,例如Phe-Lys二肽接头、Val-Cit-PABC接头;酶特异性接头,例如葡糖苷酸-MABC接头或β-葡糖苷酸接头;二硫化物接头,例如:二硫环肽接头、s Nfo-SPDB接头或SPDB接头;金属辅助接头,例如钯接头或铁接头;光可裂解接头,例如硝基苄基接头,或二6-(3-琥珀酰亚胺基羰氧基甲基-4-硝基-苯氧基)-己酸二硫化物二乙醇酯(SCNE)接头。
在一些实施方案中,接头可以包括至少一个氨基酸残基并且可以是至少或大约2、3、4、5、6、7、10、15、20、25、30、40、或50个氨基酸残基的肽。当接头是单个氨基酸残基时,它可以是任何天然或非天然存在的氨基酸(例如,Gly、Cys、Lys、Glu或Asp)或包含两个此类残基的二肽(例如,Gly-Lys)。当接头是短肽时,它可以是富含甘氨酸的肽(其往往是柔性的),例如具有序列[Gly-Gly-Gly-Gly-Ser]n的肽,其中n是1到6的整数(SEQ ID NO.:25)(包括在内),或富含丝氨酸的肽接头。富含丝氨酸的肽接头包括式[X-X-X-X-Gly]y的接头,其中多达两个X是Thr,其余的X是Ser,并且y是1到5的整数(SEQ ID NO.:26)(包括在内)(例如,Ser-Ser-Ser-Ser-Gly(SEQ ID NO.:27),其中y大于1)。其他接头包括刚性接头(例如,PAPAP(SEQ ID NO.:28)和(PT)nP,其中n是2、3、4、5、6或7(SEQ ID NO.:29))和α-螺旋接头(例如,A(EAAAK)nA,其中n是1、2、3、4或5(SEQ ID NO.:30))。当接头是琥珀酸时,其一个羧基可以与氨基酸残基的氨基形成酰胺键,而其另一个羧基可以例如与肽或取代基的氨基形成酰胺键。当接头是Lys、Glu或Asp时,其羧基可以与氨基酸残基的氨基形成酰胺键,而其氨基可以例如与取代基的羧基形成酰胺键。当Lys用作接头时,可以在Lys的ε-氨基和取代基之间插入另外的接头。另外的接头可以是琥珀酸,其可以与Lys的ε-氨基和取代基中存在的氨基形成酰胺键。在一个实施方案中,另外的接头是Glu或Asp(例如,其与Lys的ε-氨基形成酰胺键并且与取代基中存在的羧基形成另一个酰胺键),即,取代基是Nε-酰化的赖氨酸残基。
本文公开的构建体还可包含蛋白质转导结构域(PTD),即介导跨细胞膜易位的氨基酸序列。有用的蛋白转导结构域包括YopM蛋白质转导结构域和IpaH蛋白质转导结构域。示例性的YopM蛋白质转导结构域可以具有如SEQ ID NO.:17中所示的氨基酸序列。示例性的IpaH9.8转导结构域可以具有如SEQ ID NO.11的氨基酸1-57中所示的氨基酸序列。蛋白质转导结构域和包含第一截短的T3SS细菌效应多肽序列和第二截短的T3SS细菌效应多肽序列的构建体可以是融合蛋白。在一些实施方案中,蛋白质转导结构域的氨基酸序列和第一截短的细菌效应多肽和第二截短的细菌效应多肽的氨基酸序列可以是相邻的,蛋白质转导结构域的氨基酸序列和第一截短的细菌效应多肽和第二截短的细菌效应多肽的氨基酸序列通过肽键接合。在一些实施方案中,蛋白质转导结构域和第一截短的细菌效应多肽的氨基酸序列和第二截短的细菌效应多肽的氨基酸序列通过接头连接,即上述任何接头。
示例性的构建体和此类构建体的氨基酸序列在图3和4中示出。图3描绘了包含YopM蛋白质转导结构域、截短的YopM多肽和截短的OspZ多肽的融合蛋白。图3中所示的融合蛋白的氨基酸序列为SEQ ID No.:23。图4描绘了包含YopM蛋白质转导结构域、截短的YopM多肽和截短的NleC多肽的融合蛋白。图4中所示的融合蛋白的氨基酸序列为SEQ ID No.:24。
相对于天然多肽氨基酸序列(本文也称为“变体”T3SS多肽),本文提供的多肽可以具有一个或多个氨基酸添加、减少或取代,可以如本文所述制备和修饰。在一些情况下,可以通过选择在不显著改变其对维持以下方面的作用的取代来进行氨基酸取代:(a)取代区域内肽骨架的结构,(b)靶位点处的分子的电荷或疏水性,或(c)大部分侧链。例如,可以基于侧链特性将天然存在的残基分成几组:(1)疏水性氨基酸(正亮氨酸、甲硫氨酸、丙氨酸、缬氨酸、亮氨酸和异亮氨酸);(2)中性亲水氨基酸(半胱氨酸、丝氨酸和苏氨酸);(3)酸性氨基酸(天冬氨酸和谷氨酸);(4)碱性氨基酸(天冬酰胺、谷氨酰胺、组氨酸、赖氨酸和精氨酸);(5)影响链定向的氨基酸(甘氨酸和脯氨酸);(6)芳香族氨基酸(色氨酸、酪氨酸和苯丙氨酸)。在这些组内进行的取代可以被认为是保守取代。有用的保守取代的非限制性实例可以包括但不限于用缬氨酸取代丙氨酸、赖氨酸取代精氨酸、谷氨酰胺取代天冬酰胺、谷氨酸取代天冬氨酸、丝氨酸取代半胱氨酸、天冬酰胺取代谷氨酰胺、天冬氨酸取代谷氨酸、脯氨酸取代甘氨酸、精氨酸取代组氨酸、亮氨酸取代异亮氨酸、异亮氨酸取代亮氨酸、精氨酸取代赖氨酸、亮氨酸取代甲硫氨酸、亮氨酸取代苯丙氨酸、甘氨酸取代脯氨酸、苏氨酸取代丝氨酸、丝氨酸取代苏氨酸、酪氨酸取代色氨酸、苯丙氨酸取代酪氨酸和/或亮氨酸取代缬氨酸。
在一些实施方案中,多肽可以包括一个或多个非保守取代。非保守取代通常需要将上述类别之一的成员交换为另一类别的成员。这种产生对于提供这种构建体的大量或替代实施方案可能是合乎需要的。通过测定肽变体的比活性可以容易地确定氨基酸变化是否导致功能性多肽。
本文提供的多肽可以通过表达编码多肽的重组核酸或通过化学合成获得。例如,可以使用使用编码本文提供的多肽的表达载体的重组技术。然后可以使用例如亲和层析技术和HPLC纯化所得的多肽。纯化程度可以通过任何合适的方法测量,包括但不限于:柱色谱、聚丙烯酰胺凝胶电泳或高效液相色谱。本文提供的多肽可以被设计或工程化为包含允许多肽被纯化(例如,捕获到亲和基质上)的标签序列。例如,可以使用诸如c-myc、血凝素、多组氨酸或FlagTM标签(Kodak)之类的标签来帮助多肽纯化。此类标签可以插入多肽内的任何位置,包括羧基或氨基末端。
本文公开的多肽可以从宿主细胞内部或外部分离,或从培养细胞的培养基中分离,并纯化为基本上纯且同质的多肽。基本上纯的多肽可以是例如,从产生它的宿主细胞或培养基中去除的多肽,并且可以是至少60%、至少70%、至少80%或至少90%纯的,不含或基本上不含其他成分,例如不相关的多肽、脂质、核酸或碳水化合物。可以通过适当选择和组合(例如,柱层析、过滤、超滤、盐析、溶剂沉淀、溶剂萃取、蒸馏、免疫沉淀、SDS-聚丙烯酰胺凝胶电泳、等电聚焦、透析和重结晶)来分离和纯化多肽。色谱法包括例如亲和层析、离子交换层析、疏水层析、凝胶过滤法、反相色谱法和吸附色谱法。色谱法可以使用液相色谱法(例如HPLC和FPLC)进行。
本文提供的多肽可以通过与药学上可接受的无毒赋形剂或载体混合而配制成药物组合物。此类组合物可以以有效治疗炎症性病症的量施用于对其有需要的受试者。可以制备用于口服或肠胃外给药的药物组合物,例如鼻内、舌下、口腔、动脉内、关节内、心脏内、皮内、肌肉内、眼内、骨内、腹膜内、鞘内、静脉内、囊内、玻璃体内、皮下、经皮、血管周围、脑内、经粘膜给药。关节内给药可以用于治疗关节的炎症性病症。配制用于肠胃外给药的组合物可以是在生理缓冲水溶液中的液体溶液或悬浮液形式;用于口服给药,特别是以片剂或胶囊的形式;或者用于鼻内给药,特别是以粉末、滴鼻剂或气雾剂的形式。
赋形剂或载体可以根据制剂和给药途径而变化。Remington's PharmaceuticalSciences(E.W.Martin)和USP/NF(美国药典和国家处方集)中描述了药物载体。示例性的赋形剂可以包括糖类,例如乳糖、右旋糖、蔗糖、山梨糖醇、甘露糖醇;淀粉、阿拉伯树胶、磷酸钙、藻酸盐、黄蓍胶、明胶、硅酸钙、微晶纤维素、聚乙烯吡咯烷酮、纤维素、水、糖浆和甲基纤维素。在一些实施方案中,制剂可以包括润滑剂、润湿剂、乳化剂、防腐剂、甜味剂或调味剂。
用于肠胃外给药的制剂可以包含无菌水或盐水、聚亚烷基二醇例如聚乙二醇、植物来源的油、氢化萘等作为常见赋形剂。特别地,生物相容的、可生物降解的丙交酯聚合物、丙交酯/乙交酯共聚物或聚氧乙烯-聚氧丙烯共聚物是用于控制体内多肽释放的赋形剂的实例。其他合适的肠胃外递送系统包括乙烯-乙酸乙烯酯共聚物颗粒、渗透泵、可植入的输注系统和脂质体。如果需要,用于吸入给药的制剂可以包含赋形剂,例如乳糖。吸入制剂可以是包含例如聚氧乙烯-9-月桂基醚、甘胆酸盐和脱氧胆酸盐的水溶液,或者它们可以是用于以滴鼻剂形式给药的油性溶液。如果需要,可以将化合物配制成鼻内施用的凝胶。用于肠胃外给药的制剂还可以包括用于口腔给药的甘胆酸盐。
对于口服给药,片剂或胶囊可以通过常规方法与药学上可接受的赋形剂一起制备,例如粘合剂(例如预胶化玉米淀粉、聚乙烯吡咯烷酮或羟丙基甲基纤维素);填充剂(例如乳糖、微晶纤维素或磷酸氢钙);润滑剂(例如硬脂酸镁、滑石粉或二氧化硅);崩解剂(例如马铃薯淀粉或羟基乙酸淀粉钠);或润湿剂(例如,十二烷基硫酸钠)。片剂可以通过本领域已知的方法进行包衣。还可以配制用于口服给药的制剂以提供化合物的受控释放。
鼻用制剂可以以液体形式或作为干燥产品存在。雾化的水性悬浮液或溶液可以包括载体或赋形剂以调节pH和/或张力。
在一些实施方案中,可以配制药物组合物以调节活性成分的释放。还可以配制药物组合物以便在施用于患者后提供活性成分的快速、持续或延迟释放。本文提供的多肽可以配制成持续释放的剂型。例如,可以将多肽配制成受控释放制剂。在一些实施方案中,包衣、包膜或保护性基质可以配制为包含一种或多种本文提供的多肽。在一些实施方案中,此类包衣、包膜或保护性基质可以用于涂覆留置装置,例如支架、导管和腹膜透析管。在一些情况下,本文提供的多肽可以并入聚合物、脂质体、微乳液、微粒、纳米颗粒或蜡中。
还提供了编码本文公开的任何构建体的核酸。分离的核酸是指不直接与衍生出它的生物体的天然存在的基因组中与其直接相邻的两个序列(一个在5'端和一个在3'端)相邻的核酸。例如,分离的核酸可以是但不限于任何长度的重组DNA分子,条件是通常发现的紧接天然存在的基因组中重组DNA分子侧翼的核酸序列之一被去除或不存在。因此,分离的核酸包括但不限于独立于其他序列作为单独分子存在的重组DNA(例如,通过PCR或限制性内切核酸酶处理产生的cDNA或基因组DNA片段),以及并入载体、自主复制的质粒、病毒(例如逆转录病毒、腺病毒或疱疹病毒)或并入原核生物或真核生物的基因组DNA的重组DNA。此外,分离的核酸可以包括作为杂交或融合核酸序列的一部分的重组DNA分子。
分离的核酸还包括任何非天然存在的核酸,因为非天然存在的核酸序列在自然界中未发现并且在天然存在的基因组中不具有直接相邻的序列。例如,非天然存在的核酸(例如工程化的核酸)被认为是分离的核酸。可以使用分子克隆或化学核酸合成技术制备工程化的核酸(例如,编码包含或由如SEQ ID NO.:23和SEQ ID NO.:24中所示的氨基酸序列组成的多肽的核酸)。分离的非天然存在的核酸可以独立于其他序列,或并入载体、自主复制的质粒、病毒(例如逆转录病毒、腺病毒或疱疹病毒),或原核生物或真核生物的基因组DNA。此外,非天然存在的核酸可以包括作为杂交或融合核酸序列的一部分的核酸分子。存在于例如cDNA文库或基因组文库或含有基因组DNA限制性消化的凝胶切片中的数百至数百万其他核酸中的核酸不被视为分离的核酸。
核酸可以是RNA和DNA,包括mRNA、cDNA、基因组DNA、合成的(例如化学合成的)DNA和核酸类似物。核酸可以是双链或单链的,在单链的情况下,可以是有义链或反义链。此外,核酸可以是环状或线性的。核酸类似物可以在碱基部分、糖部分或磷酸骨架上进行修饰以提高例如核酸的稳定性、杂交或溶解度。碱基部分的修饰包括脱氧胸苷修饰为脱氧尿苷,以及脱氧胞苷修饰为5-甲基-2'-脱氧胞苷和5-溴-2'-脱氧胞苷。糖部分的修饰可以包括修饰核糖的2'羟基以形成2'-0-甲基或2'-0-烯丙基糖。可以修饰磷酸脱氧核糖骨架以产生吗啉代核酸,其中每个碱基部分都连接到六元的吗啉代环或肽核酸,其中脱氧磷酸酯骨架被假肽骨架取代,四个碱基保留。此外,脱氧磷酸酯骨架可以被例如硫代磷酸酯或二硫代磷酸酯骨架、亚磷酰胺或烷基磷酸三酯骨架替代。
本文提供的核酸可以包括或由如SEQ ID NO.:2;SEQ ID NO.:4;SEQ ID NO.:6;SEQ ID NO.:8;SEQ ID NO.:10;SEQ ID NO.:12;SEQ ID NO.:14;SEQ ID NO.:16;SEQ IDNO.:18;SEQ ID NO.:20;或SEQ ID NO.:22中所示的序列中所示的任何核酸序列组成。在一些实施方案中,核酸可以包括SEQ ID NO.:2;SEQ ID NO.:4;SEQ ID NO.:6;SEQ ID NO.:8;SEQ ID NO.:10;SEQ ID NO.:12;SEQ ID NO.:14;SEQ ID NO.:16;SEQ ID NO.:18;SEQ IDNO.:20;或SEQ ID NO.:22中的任何一个的截短的核酸。
编码第一截短的T3SS细菌效应多肽序列和第二截短的T3SS细菌效应多肽序列的核酸包括密码子优化的核酸。为了表达,可以将核酸引入载体(例如,质粒或病毒载体)中,并且本发明包括这样的载体。核酸可以与适用于原核或真核系统的调节区域可操作地连接。在具体实施方案中,调节区域可以是例如启动子或增强子。有用的启动子包括细胞类型特异性启动子、组织特异性启动子、组成型活性启动子和广泛表达启动子。本发明还包括宿主细胞,所述宿主细胞包括表达本发明的多肽的载体,这些细胞可以是原核细胞(例如细菌)或真核细胞(例如哺乳动物)。
通常,本文提供的分离的核酸长度至少为10个核苷酸(例如,长度为10、15、20、25、30、35、40、50、75、100、200、300、350、400或更多个核苷酸)。小于全长的核酸分子可以用作例如引物或探针。可以用分子克隆和化学核酸合成技术产生分离的核酸分子。例如,可以使用聚合酶链反应(PCR)技术。也可以化学合成分离的核酸,作为单个核酸分子(例如,使用亚磷酰胺技术在3'到5'方向使用自动DNA合成)或作为一系列寡核苷酸,然后可以将其连接到载体中。
治疗方法
还提供了通过施用治疗有效量的包含本文公开的任何构建体的药物组合物来治疗患有炎症性病症或处于炎症性病症风险中的受试者的方法。在一些实施方案中,需要治疗的受试者(例如,人类患者)被诊断患有、怀疑患有炎症性病症或处于炎症性病症的风险中。示例性的炎症性病症包括但不限于关节炎和关节疾病中发现的炎症性病症,例如类风湿性关节炎和骨关节炎;心血管疾病;过敏;哮喘;慢性阻塞性肺病;糖尿病;胃肠道疾病,例如炎症性肠病、克罗恩病和回肠结肠炎(ileocolitis);癌症,例如肾癌、前列腺癌、卵巢癌、肝细胞癌、胰腺癌、结直肠癌、肺癌和间皮瘤;慢性肾病;和阿尔茨海默病。
一般而言,治疗可以包括抑制正在经历或表现出疾病、病症或疾病的病理学或症候学的个体的炎症性病症的一种或多种(即阻止病理学和/或症候学的进一步发展)。治疗还可以包括改善正在经历或表现出疾病、病症或疾病的病理学或症候学的个体的炎症性病症(即,逆转病理学和/或症候学),例如降低疾病的严重程度或减轻或缓和疾病的一种或多种症状。
受试者可以是人类或非人类动物。示例性的非人类物种包括但不限于非人类灵长类动物;家畜,例如马、猪、牛、羊;猫、狗、小鼠或大鼠。可以通过检测通常与炎症性病症相关的症状来鉴定适合治疗的受试者,例如疼痛、疲劳、胃肠道症状例如便秘、腹泻和胃酸倒流、体重增加和频繁感染。还可以通过实验室测试来鉴定适合治疗的受试者,包括例如血清蛋白电泳(SPE)、高敏C反应蛋白、纤维蛋白原和促炎细胞因子的检测。
治疗有效量可以是引发生物或医学反应的活性化合物或药剂的量,研究人员、兽医、医生或其他临床医生正在组织、系统、动物、个体或人中寻找这样的反应。
本文提供的组合物可以与一种或多种常规治疗剂联合施用,包括治疗关节炎和关节疾病,例如类风湿性关节炎和骨关节炎;心血管疾病;过敏;哮喘;慢性阻塞性肺病;糖尿病;胃肠道疾病,例如炎症性肠病、克罗恩病和回肠结肠炎;癌症,例如肾癌、前列腺癌、卵巢癌、肝细胞癌、胰腺癌、结直肠癌、肺癌和间皮瘤;慢性肾病;和阿尔茨海默病。
本发明的特点
一般而言,本发明的特征在于可以包括两个或更多个截短的T3SS细菌效应多肽的构建体,所述截短的T3SS细菌效应多肽包括全长细菌效应多肽的一部分。在一方面,构建体可以包括与截短的T3SS半胱氨酸甲基转移酶多肽连接的截短的YopM多肽。截短的YopM多肽可以具有与SEQ ID NO.:19中所示的氨基酸序列至少90%相同的氨基酸序列。截短的YopM多肽可以具有如SEQ ID NO.:19中所示的氨基酸序列。截短的T3SS半胱氨酸甲基转移酶多肽可以包括OspZ多肽的一部分,所述OspZ多肽具有如SEQ ID NO.:3中所示的氨基酸序列。截短的OspZ多肽可以具有与SEQ ID NO.:3的氨基酸226-446至少90%相同的氨基酸序列。截短的OspZ多肽可以具有如SEQ ID NO.:3的氨基酸226-446中所示的氨基酸序列。构建体可以进一步包括蛋白质转导结构域,例如,如SEQ ID NO.:17中所示的YopM蛋白质转导结构域。在一些实施方案中,构建体包括如SEQ ID NO.:23中所示的氨基酸序列。
在另一方面,构建体可以包括与截短的T3SS锌金属蛋白酶多肽连接的截短的YopM多肽。截短的YopM多肽可以具有与SEQ ID NO.:19中所示的氨基酸序列至少90%相同的氨基酸序列。截短的YopM多肽可以具有如SEQ ID NO.:19中所示的氨基酸序列。截短的T3SS锌金属蛋白酶多肽可以包括具有如SEQ ID NO.:5中所示的氨基酸序列的NleC多肽的一部分。截短的NleC多肽可以具有与SEQ ID NO.:5的氨基酸2-187至少90%相同的氨基酸序列。在一些实施方案中,截短的NleC多肽可以具有如SEQ ID NO.:5的氨基酸2-187中所示的氨基酸序列。在一些实施方案中,截短的NleC多肽具有如SEQ ID NO.:5的氨基酸2-187中所示的氨基酸序列。构建体可以进一步包括蛋白质转导结构域,例如,如SEQ ID NO.:17中所示的YopM蛋白质转导结构域。在一些实施方案中,构建体具有如SEQ ID NO.:24中所示的氨基酸序列。
在一方面,构建体可以包括与截短的T3SS O-GlcNac转移酶连接的截短的YopM多肽。截短的YopM多肽可以具有与SEQ ID NO.:19中所示的氨基酸序列至少90%相同的氨基酸序列。截短的YopM多肽可以具有如SEQ ID NO.:19中所示的氨基酸序列。截短的T3SS O-GlcNac转移酶可以包括具有如SEQ ID NO.:9中所示的氨基酸序列的NleB多肽的一部分。截短的NleB多肽可以具有与SEQ ID NO.:9的氨基酸2-226至少90%的相同的氨基酸序列。截短的NleB多肽可以具有如SEQ ID NO.:9的氨基酸2-226中所示的氨基酸序列。构建体可以进一步包括蛋白质转导结构域,例如,如SEQ ID NO.:17中所示的YopM蛋白质转导结构域。
在一方面,构建体可以包括与截短的第二T3SS E3泛素连接酶连接的截短的第一T3SS E3泛素连接酶多肽。第一和第二截短的T3SS E3泛素连接酶多肽可以不同。截短的第一E3泛素连接酶可以包括具有如SEQ ID NO.:11中所示的氨基酸序列的IpaH9.8多肽的一部分。截短的第一IpaH9.8多肽包括与SEQ ID NO.:11的氨基酸56-228至少90%相同的氨基酸序列。截短的第二E3泛素连接酶包括具有如SEQ ID NO.:13中所示的氨基酸序列的IpaH4.5多肽的一部分。截短的第二IpaH4.5多肽包括与SEQ ID NO.:13的氨基酸62-270至少90%相同的氨基酸序列。构建体可以进一步包括蛋白质转导结构域,例如,如SEQ IDNO.:11的氨基酸1-57中所示的IpaH9.8蛋白质转导结构域。
在一方面,构建体可以包括与半胱氨酸甲基转移酶连接的RhoGTP酶调节剂,其中RhoGTP酶调节剂通过pH敏感性接头与半胱氨酸甲基转移酶连接。RhoGTP酶调节剂可以是具有如SEQ ID NO.:1中所示的氨基酸序列的YopE多肽。半胱氨酸甲基转移酶可以是具有如SEQ ID NO.:5中所示的氨基酸序列的OspZ多肽。pH敏感性接头包括肼、基于氨基磷酸酯的接头或硫代马来酸。
在一方面,构建体可以包括与乙酰转移酶连接的截短的YopM多肽。截短的YopM多肽可以具有与SEQ ID NO.:19中所示的氨基酸序列至少90%相同的氨基酸序列。截短的YopM多肽可以具有如SEQ ID NO.:19中所示的氨基酸序列。乙酰转移酶可以是YopJ多肽,其具有如SEQ ID NO.:9中所示的氨基酸序列,包括在半胱氨酸172处的突变。
在一方面,构建体可以包括与乙酰转移酶连接的截短的YopE多肽。截短的YopE多肽可以包括具有如SEQ ID NO.:1中所示的氨基酸序列的YopE多肽的一部分。构建体可以进一步包括蛋白质转导结构域,例如,如SEQ ID NO.:11的氨基酸1-57中所示的IpaH9.8蛋白质转导结构域。
在一方面,构建体可以进一步包括蛋白质转导结构域。蛋白质转导结构域可以是YopM蛋白质转导结构域。YopM蛋白质转导结构域可以具有SEQ ID NO.:19中所示的氨基酸序列。蛋白转导结构域可以是IpaH9.8蛋白质转导结构域。IpaH9.8蛋白质转导结构域可以具有如SEQ ID NO.:23的氨基酸1-56中所示的氨基酸序列。
在一方面,任何构建体可以包括融合蛋白。第一截短的T3SS细菌效应多肽和第二截短的T3SS细菌效应多肽可以通过接头接合。接头可以是可切割的接头。可切割的接头可以是pH敏感性接头。pH敏感性接头可以选自由肼、基于氨基磷酸酯的接头和硫代马来酸组成的组。
在一方面,还提供了编码本文公开的任何构建体的核酸。
在一方面,构建体可以配制成包含构建体和药学上可接受的载体的药物组合物。
在一方面,本申请的特征在于治疗患有炎症性病症或处于炎症性病症风险中的受试者的方法,所述方法包括向受试者施用治疗有效量的药物组合物,所述药物组合物包括构建体,所述构建体可以包括第一截短的T3SS细菌效应多肽和第二截短的T3SS细菌效应多肽和药学上可接受的载体。方法可以包括鉴定受试者的步骤。炎症性病症可以是胃肠道疾病、肌肉骨骼疾病、自身免疫疾病或皮肤疾病。在一方面,炎症性病症可以包括炎症性肠病、克罗恩病、类风湿性关节炎、骨关节炎、癌症、过敏、心血管疾病、慢性阻塞性肺病和糖尿病。
实施例
实施例1
我们分析了E3泛素连接酶IpaH7.8和IpaH9.8对来自THP-1细胞的细胞因子释放的影响。在存在和不存在递增量(0.25μg、0.5μg和1.0μg)重组IpaH7.8或IpaH9.8的情况下培养THP-1细胞。对于IpaH7.8,0.25μg蛋白质约为3.87nM。对于IpaH9.8,0.25μg蛋白质约为4.0nM。然后用脂多糖(LPS)处理培养物以诱导细胞因子释放。
如表2所示,重组IpaH7.8抑制IL-1β、TNF-α、MCP-1、IL-6、IL-8、IL-23的释放。如表3所示,重组IpaH9.8对IL-1β、TNF-α和MCP-1、IL-6、IL-8、IL-23的释放产生剂量依赖性抑制。这些数据表明,低纳摩尔浓度的E3泛素连接酶IpaH7.8和IpaH9.8可以有效地下调THP-1细胞中的细胞因子水平。
表2:IpaH7.8对细胞因子释放的影响
表3:IpaH9.8对细胞因子释放的影响
序列表
<110> 因内特生物制品有限责任公司
<120> 免疫调节组合物和方法
<130> G6113-00029
<140>
<141>
<150> 62/841,312
<151> 2019-05-01
<160> 30
<170> PatentIn version 3.5
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<210> 7
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Tyr Ser Arg Met Asp Val Glu Val Met Pro Ala Leu Val Ile Gln Ala
50 55 60
Asn Asn Lys Tyr Pro Glu Met Asn Leu Asn Leu Val Thr Ser Pro Leu
65 70 75 80
Asp Leu Ser Ile Glu Ile Lys Asn Val Ile Glu Asn Gly Val Arg Ser
85 90 95
Ser Arg Phe Ile Ile Asn Met Gly Glu Gly Gly Ile His Phe Ser Val
100 105 110
Ile Asp Tyr Lys His Ile Asn Gly Lys Thr Ser Leu Ile Leu Phe Glu
115 120 125
Pro Ala Asn Phe Asn Ser Met Gly Pro Ala Met Leu Ala Ile Arg Thr
130 135 140
Lys Thr Ala Ile Glu Arg Tyr Gln Leu Pro Asp Cys His Phe Ser Met
145 150 155 160
Val Glu Met Asp Ile Gln Arg Ser Ser Ser Glu Cys Gly Ile Phe Ser
165 170 175
Phe Ala Leu Ala Lys Lys Leu Tyr Ile Glu Arg Asp Ser Leu Leu Lys
180 185 190
Ile His Glu Asp Asn Ile Lys Gly Ile Leu Ser Asp Gly Glu Asn Pro
195 200 205
Leu Pro His Asp Lys Leu Asp Pro Tyr Leu Pro Val Thr Phe Tyr Lys
210 215 220
His Thr Gln Gly Lys Lys Arg Leu Asn Glu Tyr Leu Asn Thr Asn Pro
225 230 235 240
Gln Gly Val Gly Thr Val Val Asn Lys Lys Asn Glu Thr Ile Val Asn
245 250 255
Arg Phe Asp Asn Asn Lys Ser Ile Val Asp Gly Lys Glu Leu Ser Val
260 265 270
Ser Val His Lys Lys Arg Ile Ala Glu Tyr Lys Thr Leu Leu Lys Val
275 280 285
<210> 8
<211> 867
<212> DNA
<213> 假结核耶尔森氏菌
<400> 8
atgatcggac caatatcaca aataaatatc tccggtggct tatcagaaaa agagaccagt 60
tctttaatca gtaatgaaga gcttaaaaat atcataacac agttggaaac tgatatatcg 120
gatggatcct ggttccataa aaattattca cgtatggatg tagaagtcat gcccgcattg 180
gtaatccagg cgaacaataa atatccggaa atgaatctta atcttgttac atctccattg 240
gacctttcaa tagaaataaa aaacgtcata gaaaatggag ttagatcttc ccgcttcata 300
attaacatgg gggaaggtgg aatacatttc agtgtaattg attacaaaca tataaatggg 360
aaaacatctc tgatattgtt tgaaccagca aactttaaca gtatggggcc agcgatgctg 420
gcaataagga caaaaacggc tattgaacgt tatcaattac ctgattgcca tttctccatg 480
gtggaaatgg atattcagcg aagctcatct gaatgtggta tttttagttt tgcactggca 540
aaaaaacttt acatcgagag agatagcctg ttgaaaatac atgaagataa tataaaaggt 600
atattaagtg atggtgaaaa tcctttaccc cacgataagt tggacccgta tctcccggta 660
actttttaca aacatactca aggtaaaaaa cgtcttaatg aatatttaaa tactaacccg 720
cagggagttg gtactgttgt taacaaaaaa aatgaaacca tcgttaatag atttgataac 780
aataaatcca ttgtagatgg aaaggaatta tcagtttcgg tacataaaaa gagaatagct 840
gaatataaaa cacttctcaa agtataa 867
<210> 9
<211> 336
<212> PRT
<213> 大肠杆菌
<400> 9
Met Ile Pro Pro Leu Asn Arg Tyr Val Pro Ala Leu Ser Lys Asn Glu
1 5 10 15
Leu Val Lys Thr Val Thr Asn Arg Asp Ile Gln Phe Thr Ser Phe Asn
20 25 30
Gly Lys Asp Tyr Pro Leu Cys Phe Leu Asp Glu Lys Thr Pro Leu Leu
35 40 45
Phe Gln Trp Phe Glu Arg Asn Pro Ala Arg Phe Gly Lys Asn Asp Ile
50 55 60
Pro Ile Ile Asn Thr Glu Lys Asn Pro Tyr Leu Asn Asn Ile Ile Lys
65 70 75 80
Ala Ala Thr Ile Glu Lys Glu Arg Leu Ile Gly Ile Phe Val Asp Gly
85 90 95
Asp Phe Phe Pro Gly Gln Lys Asp Ala Phe Ser Lys Leu Glu Tyr Asp
100 105 110
Tyr Glu Asn Ile Lys Val Ile Tyr Arg Asn Asp Ile Asp Phe Ser Met
115 120 125
Tyr Asp Lys Lys Leu Ser Glu Ile Tyr Met Glu Asn Ile Ser Lys Gln
130 135 140
Glu Ser Met Pro Glu Glu Lys Arg Asp Cys His Leu Leu Gln Leu Leu
145 150 155 160
Lys Lys Glu Leu Ser Asp Ile Gln Glu Gly Asn Asp Ser Leu Ile Lys
165 170 175
Ser Tyr Leu Leu Asp Lys Gly His Gly Trp Phe Asp Phe Tyr Arg Asn
180 185 190
Met Ala Met Leu Lys Ala Gly Gln Leu Phe Leu Glu Ala Asp Lys Val
195 200 205
Gly Cys Tyr Asp Leu Ser Thr Asn Ser Gly Cys Ile Tyr Leu Asp Ala
210 215 220
Asp Met Ile Ile Thr Glu Lys Leu Gly Gly Ile Tyr Ile Pro Asp Gly
225 230 235 240
Ile Ala Val His Val Glu Arg Ile Asp Gly Arg Ala Ser Met Glu Asn
245 250 255
Gly Ile Ile Ala Val Asp Arg Asn Asn His Pro Ala Leu Leu Ala Gly
260 265 270
Leu Glu Ile Met His Thr Lys Phe Asp Ala Asp Pro Tyr Ser Asp Gly
275 280 285
Val Cys Asn Gly Ile Arg Lys His Phe Asn Tyr Ser Leu Asn Glu Asp
290 295 300
Tyr Asn Ser Phe Cys Asp Phe Ile Glu Phe Lys His Asp Asn Ile Ile
305 310 315 320
Met Asn Thr Ser Gln Phe Thr Gln Ser Ser Trp Ala Arg His Val Gln
325 330 335
<210> 10
<211> 540
<212> DNA
<213> 大肠杆菌
<400> 10
atgttatctc cattaaatgt tcttcaattt aatttcagag gagagaccgc tttatcagat 60
agtgctcctc tccagactgt ttcctttgct ggaaaagatt attctatgga acccattgat 120
gaaaaaacac ccattctttt tcagtggttt gaagcaaggc cagagcgata cggaaaaggt 180
gaagtaccga tattgaatac caaagagcat ccgtatttga gcaatattat aaatgctgca 240
aaaatagaaa atgagcgcgt aataggagta ctggtagacg gagactttac ttatgagcaa 300
agaaaagaat ttctcagtct tgaagatgaa catcaaaata taaagataat atatcgggaa 360
aatgttgatt tcagtatgta tgataaaaaa ctgtctgata tttatcttga aaatattcat 420
gaacaagaat catatccagc gagtgagaga gataattatc tgttaggctt attaagagaa 480
gagttaaaaa atattccata cggaaaggac tctttgattg aatcatatgc agaaaaaaga 540
<210> 11
<211> 545
<212> PRT
<213> 弗氏志贺氏菌
<400> 11
Met Leu Pro Ile Asn Asn Asn Phe Ser Leu Pro Gln Asn Ser Phe Tyr
1 5 10 15
Asn Thr Ile Ser Gly Thr Tyr Ala Asp Tyr Phe Ser Ala Trp Asp Lys
20 25 30
Trp Glu Lys Gln Ala Leu Pro Gly Glu Glu Arg Asp Glu Ala Val Ser
35 40 45
Arg Leu Lys Glu Cys Leu Ile Asn Asn Ser Asp Glu Leu Arg Leu Asp
50 55 60
Arg Leu Asn Leu Ser Ser Leu Pro Asp Asn Leu Pro Ala Gln Ile Thr
65 70 75 80
Leu Leu Asn Val Ser Tyr Asn Gln Leu Thr Asn Leu Pro Glu Leu Pro
85 90 95
Val Thr Leu Lys Lys Leu Tyr Ser Ala Ser Asn Lys Leu Ser Glu Leu
100 105 110
Pro Val Leu Pro Pro Ala Leu Glu Ser Leu Gln Val Gln His Asn Glu
115 120 125
Leu Glu Asn Leu Pro Ala Leu Pro Asp Ser Leu Leu Thr Met Asn Ile
130 135 140
Ser Tyr Asn Glu Ile Val Ser Leu Pro Ser Leu Pro Gln Ala Leu Lys
145 150 155 160
Asn Leu Arg Ala Thr Arg Asn Phe Leu Thr Glu Leu Pro Ala Phe Ser
165 170 175
Glu Gly Asn Asn Pro Val Val Arg Glu Tyr Phe Phe Asp Arg Asn Gln
180 185 190
Ile Ser His Ile Pro Glu Ser Ile Leu Asn Leu Arg Asn Glu Cys Ser
195 200 205
Ile His Ile Ser Asp Asn Pro Leu Ser Ser His Ala Leu Gln Ala Leu
210 215 220
Gln Arg Leu Thr Ser Ser Pro Asp Tyr His Gly Pro Arg Ile Tyr Phe
225 230 235 240
Ser Met Ser Asp Gly Gln Gln Asn Thr Leu His Arg Pro Leu Ala Asp
245 250 255
Ala Val Thr Ala Trp Phe Pro Glu Asn Lys Gln Ser Asp Val Ser Gln
260 265 270
Ile Trp His Ala Phe Glu His Glu Glu His Ala Asn Thr Phe Ser Ala
275 280 285
Phe Leu Asp Arg Leu Ser Asp Thr Val Ser Ala Arg Asn Thr Ser Gly
290 295 300
Phe Arg Glu Gln Val Ala Ala Trp Leu Glu Lys Leu Ser Ala Ser Ala
305 310 315 320
Glu Leu Arg Gln Gln Ser Phe Ala Val Ala Ala Asp Ala Thr Glu Ser
325 330 335
Cys Glu Asp Arg Val Ala Leu Thr Trp Asn Asn Leu Arg Lys Thr Leu
340 345 350
Leu Val His Gln Ala Ser Glu Gly Leu Phe Asp Asn Asp Thr Gly Ala
355 360 365
Leu Leu Ser Leu Gly Arg Glu Met Phe Arg Leu Glu Ile Leu Glu Asp
370 375 380
Ile Ala Arg Asp Lys Val Arg Thr Leu His Phe Val Asp Glu Ile Glu
385 390 395 400
Val Tyr Leu Ala Phe Gln Thr Met Leu Ala Glu Lys Leu Gln Leu Ser
405 410 415
Thr Ala Val Lys Glu Met Arg Phe Tyr Gly Val Ser Gly Val Thr Ala
420 425 430
Asn Asp Leu Arg Thr Ala Glu Ala Met Val Arg Ser Arg Glu Glu Asn
435 440 445
Glu Phe Thr Asp Trp Phe Ser Leu Trp Gly Pro Trp His Ala Val Leu
450 455 460
Lys Arg Thr Glu Ala Asp Arg Trp Ala Gln Ala Glu Glu Gln Lys Tyr
465 470 475 480
Glu Met Leu Glu Asn Glu Tyr Pro Gln Arg Val Ala Asp Arg Leu Lys
485 490 495
Ala Ser Gly Leu Ser Gly Asp Ala Asp Ala Glu Arg Glu Ala Gly Ala
500 505 510
Gln Val Met Arg Glu Thr Glu Gln Gln Ile Tyr Arg Gln Leu Thr Asp
515 520 525
Glu Val Leu Ala Leu Arg Leu Ser Glu Asn Gly Ser Gln Leu His His
530 535 540
Ser
545
<210> 12
<211> 1822
<212> DNA
<213> 弗氏志贺氏菌
<400> 12
gttaactgaa acagtatcgt ttttttacag ccaattttgt ttatccttat tataataaaa 60
aagtgctgaa gttcatttca tggaatgaac ttcataaaaa ctcctactta tttcttttaa 120
caaagccatt tgtccaccgg ctttaactgg atgcccatca tgttaccgat aaataataac 180
ttttcattgc cccaaaattc tttttataac actatttccg gtacatatgc tgattacttt 240
tcagcatggg ataaatggga aaaacaagcg ctccccggtg aagagcgtga tgaggctgtc 300
tcccgactta aagaatgtct tatcaataat tccgatgaac ttcgactgga ccgtttaaat 360
ctgtcctcgc tacctgacaa cttaccagct cagataacgc tgctcaatgt atcatataat 420
caattaacta acctacctga actgcctgtt acgctaaaaa aattatattc cgccagcaat 480
aaattatcag aattgcccgt gctacctcct gcgctggagt cacttcaggt acaacacaat 540
gagctggaaa acctgccagc tttacccgat tcgttattga ctatgaatat cagctataac 600
gaaatagtct ccttaccatc gctcccacag gctcttaaaa atctcagagc gacccgtaat 660
ttcctcactg agctaccagc attttctgag ggaaataatc ccgttgtcag agagtatttt 720
tttgatagaa atcagataag tcatatcccg gaaagcattc ttaatctgag gaatgaatgt 780
tcaatacata ttagtgataa cccattatca tcccatgctc tgcaagccct gcaaagatta 840
acctcttcgc cggactacca cggcccacgg atttacttct ccatgagtga cggacaacag 900
aatacactcc atcgccccct ggctgatgcc gtgacagcat ggttcccgga aaacaaacaa 960
tctgatgtat cacagatatg gcatgctttt gaacatgaag agcatgccaa caccttttcc 1020
gcgttccttg accgcctttc cgataccgtc tctgcacgca atacctccgg attccgtgaa 1080
caggtcgctg catggctgga aaaactcagt gcctctgcgg agcttcgaca gcagtctttc 1140
gctgttgctg ctgatgccac tgagagctgt gaggaccgtg tcgcgctcac atggaacaat 1200
ctccggaaaa ccctcctggt ccatcaggca tcagaaggcc ttttcgataa tgataccggc 1260
gctctgctct ccctgggcag ggaaatgttc cgcctcgaaa ttctggagga tattgcccgg 1320
gataaagtca gaactctcca ttttgtggat gagatagaag tctacctggc cttccagacc 1380
atgctcgcag agaaacttca gctctccact gccgtgaagg aaatgcgttt ctatggcgtg 1440
tcgggagtga cagcaaatga cctccgcact gccgaagcca tggtcagaag ccgtgaagag 1500
aatgaattta cggactggtt ctccctctgg ggaccatggc atgctgtact gaagcgtacg 1560
gaagctgacc gctgggcgca ggcagaagag cagaaatatg agatgctgga gaatgagtac 1620
cctcagaggg tggctgaccg gctgaaagca tcaggtctga gcggtgatgc ggatgcggag 1680
agggaagccg gtgcacaggt gatgcgtgag actgaacagc agatttaccg tcagctgact 1740
gacgaggtac tggccctgcg attgtctgaa aacggctcac aactgcacca ttcataatca 1800
catcgcataa aactgtcagc gc 1822
<210> 13
<211> 574
<212> PRT
<213> 弗氏志贺氏菌
<400> 13
Met Lys Pro Ile Asn Asn His Ser Phe Phe Arg Ser Leu Cys Gly Leu
1 5 10 15
Ser Cys Ile Ser Arg Leu Ser Val Glu Glu Gln Cys Thr Arg Asp Tyr
20 25 30
His Arg Ile Trp Asp Asp Trp Ala Arg Glu Gly Thr Thr Thr Glu Asn
35 40 45
Arg Ile Gln Ala Val Arg Leu Leu Lys Ile Cys Leu Asp Thr Arg Glu
50 55 60
Pro Val Leu Asn Leu Ser Leu Leu Lys Leu Arg Ser Leu Pro Pro Leu
65 70 75 80
Pro Leu His Ile Arg Glu Leu Asn Ile Ser Asn Asn Glu Leu Ile Ser
85 90 95
Leu Pro Glu Asn Ser Pro Leu Leu Thr Glu Leu His Val Asn Gly Asn
100 105 110
Asn Leu Asn Ile Leu Pro Thr Leu Pro Ser Gln Leu Ile Lys Leu Asn
115 120 125
Ile Ser Phe Asn Arg Asn Leu Ser Cys Leu Pro Ser Leu Pro Pro Tyr
130 135 140
Leu Gln Ser Leu Ser Ala Arg Phe Asn Ser Leu Glu Thr Leu Pro Glu
145 150 155 160
Leu Pro Ser Thr Leu Thr Ile Leu Arg Ile Glu Gly Asn Arg Leu Thr
165 170 175
Val Leu Pro Glu Leu Pro His Arg Leu Gln Glu Leu Phe Val Ser Gly
180 185 190
Asn Arg Leu Gln Glu Leu Pro Glu Phe Pro Gln Ser Leu Lys Tyr Leu
195 200 205
Lys Val Gly Glu Asn Gln Leu Arg Arg Leu Ser Arg Leu Pro Gln Glu
210 215 220
Leu Leu Ala Leu Asp Val Ser Asn Asn Leu Leu Thr Ser Leu Pro Glu
225 230 235 240
Asn Ile Ile Thr Leu Pro Ile Cys Thr Asn Val Asn Ile Ser Gly Asn
245 250 255
Pro Leu Ser Thr His Val Leu Gln Ser Leu Gln Arg Leu Thr Ser Ser
260 265 270
Pro Asp Tyr His Gly Pro Gln Ile Tyr Phe Ser Met Ser Asp Gly Gln
275 280 285
Gln Asn Thr Leu His Arg Pro Leu Ala Asp Ala Val Thr Ala Trp Phe
290 295 300
Pro Glu Asn Lys Gln Ser Asp Val Ser Gln Ile Trp His Ala Phe Glu
305 310 315 320
His Glu Glu His Ala Asn Thr Phe Ser Ala Phe Leu Asp Arg Leu Ser
325 330 335
Asp Thr Val Ser Ala Arg Asn Thr Ser Gly Phe Arg Glu Gln Val Ala
340 345 350
Ala Trp Leu Glu Lys Leu Ser Ala Ser Ala Glu Leu Arg Gln Gln Ser
355 360 365
Phe Ala Val Ala Ala Asp Ala Thr Glu Ser Cys Glu Asp Arg Val Ala
370 375 380
Leu Thr Trp Asn Asn Leu Arg Lys Thr Leu Leu Val His Gln Ala Ser
385 390 395 400
Glu Gly Leu Phe Asp Asn Asp Thr Gly Ala Leu Leu Ser Leu Gly Arg
405 410 415
Glu Met Phe Arg Leu Glu Ile Leu Glu Asp Ile Ala Arg Asp Lys Val
420 425 430
Arg Thr Leu His Phe Val Asp Glu Ile Glu Val Tyr Leu Ala Phe Gln
435 440 445
Thr Met Leu Ala Glu Lys Leu Gln Leu Ser Thr Ala Val Lys Glu Met
450 455 460
Arg Phe Tyr Gly Val Ser Gly Val Thr Ala Asn Asp Leu Arg Thr Ala
465 470 475 480
Glu Ala Met Val Arg Ser Arg Glu Glu Asn Glu Phe Thr Asp Trp Phe
485 490 495
Ser Leu Trp Gly Pro Trp His Ala Val Leu Lys Arg Thr Glu Ala Asp
500 505 510
Arg Trp Ala Gln Ala Glu Glu Gln Lys Tyr Glu Met Leu Glu Asn Glu
515 520 525
Tyr Ser Gln Arg Val Ala Asp Arg Leu Lys Ala Ser Gly Leu Ser Gly
530 535 540
Asp Ala Asp Ala Glu Arg Glu Ala Gly Ala Gln Val Met Arg Glu Thr
545 550 555 560
Glu Gln Gln Ile Tyr Arg Gln Leu Thr Asp Glu Val Leu Ala
565 570
<210> 14
<211> 1725
<212> DNA
<213> 弗氏志贺氏菌
<400> 14
atgaaaccga tcaacaatca ttcttttttt cgttcccttt gtggcttatc atgtatatct 60
cgtttatcgg tagaagaaca gtgtaccaga gattaccacc gcatctggga tgactgggct 120
agggaaggaa caacaacaga aaatcgcatc caggcggttc gattattgaa aatatgtctg 180
gatacccggg agcctgttct caatttaagc ttactgaaac tacgttcttt accaccactc 240
cctttgcata tacgtgaact taatatttcc aacaatgagt taatctccct acctgaaaat 300
tctccgcttt tgacagaact tcatgtaaat ggtaacaact tgaatatact cccgacactt 360
ccatctcaac tgattaagct taatatttca ttcaatcgaa atttgtcatg tctgccatca 420
ttaccaccat atttacaatc actctcggca cgttttaata gtctggagac gttaccagag 480
cttccatcaa cgctaacaat attacgtatt gaaggtaatc gccttactgt cttgcctgaa 540
ttgcctcata gactacaaga actctttgtt tccggcaaca gactacagga actaccagaa 600
tttcctcaga gcttaaaata tttgaaggta ggtgaaaatc aactacgcag attatccaga 660
ttaccgcaag aactattggc actggatgtt tccaataacc tactaacttc attacccgaa 720
aatataatca cattgcccat ttgtacgaat gttaacattt cagggaatcc attgtcgact 780
cacgttctgc aatccctgca aagattaacc tcttcgccgg actaccacgg cccgcagatt 840
tacttctcca tgagtgacgg acaacagaat acactccatc gccccctggc tgatgccgtg 900
acagcatggt tcccggaaaa caaacaatct gatgtatcac agatatggca tgcttttgaa 960
catgaagagc atgccaacac cttttccgcg ttccttgacc gcctttccga taccgtctct 1020
gcacgcaata cctccggatt ccgtgaacag gtcgctgcat ggctggaaaa actcagtgcc 1080
tctgcggagc ttcgacagca gtctttcgct gttgctgctg atgccactga gagctgtgag 1140
gaccgtgtcg cgctcacatg gaacaatctc cggaaaaccc tcctggtcca tcaggcatca 1200
gaaggccttt tcgataatga taccggcgct ctgctctccc tgggcaggga aatgttccgc 1260
ctcgaaattc tggaggatat tgcccgggat aaagtcagaa ctctccattt tgtggatgag 1320
atagaagtct acctggcctt ccagaccatg ctcgcagaga aacttcagct ctccactgcc 1380
gtgaaggaaa tgcgtttcta tggcgtgtcg ggagtgacag caaatgacct ccgcactgcc 1440
gaagctatgg tcagaagccg tgaagagaat gaatttacgg actggttctc cctctgggga 1500
ccatggcatg ctgtactgaa gcgtacggaa gctgaccgct gggcgcaggc agaagagcag 1560
aagtatgaga tgctggagaa tgagtactct cagagggtgg ctgaccggct gaaagcatca 1620
ggtctgagcg gtgatgcgga tgcggagagg gaagccggtg cacaggtgat gcgtgagact 1680
gaacagcaga tttaccgtca gttgactgac gaggtactgg cctga 1725
<210> 15
<211> 367
<212> PRT
<213> 鼠疫耶尔森氏菌
<400> 15
Met Phe Ile Thr Pro Arg Asn Val Ser Asn Thr Phe Leu Gln Glu Pro
1 5 10 15
Leu Arg His Ser Ser Asp Leu Thr Glu Met Pro Val Glu Ala Glu Asn
20 25 30
Val Lys Ser Lys Thr Glu Tyr Tyr Asn Ala Trp Ala Val Trp Glu Arg
35 40 45
Asn Ala Pro Pro Gly Asn Gly Glu Gln Arg Glu Met Ala Val Ser Arg
50 55 60
Leu Arg Asp Cys Leu Asp Arg Gln Ala His Glu Leu Glu Leu Asn Asn
65 70 75 80
Leu Gly Leu Ser Ser Leu Pro Glu Leu Pro Pro His Leu Glu Ser Leu
85 90 95
Val Ala Ser Cys Asn Ser Leu Thr Glu Leu Pro Glu Leu Pro Gln Ser
100 105 110
Leu Lys Ser Leu Gln Val Asp Asn Asn Asn Leu Lys Ala Leu Ser Asp
115 120 125
Leu Pro Pro Ser Leu Glu Phe Leu Ala Ala Gly Asn Asn Gln Leu Glu
130 135 140
Glu Leu Pro Glu Leu Gln Asn Ser Ser Phe Leu Lys Ile Ile Asp Val
145 150 155 160
Asp Asn Asn Ser Leu Lys Lys Leu Pro Asp Leu Pro Pro Ser Leu Glu
165 170 175
Phe Leu Ala Ala Gly Asn Asn Gln Leu Glu Glu Leu Ser Glu Leu Gln
180 185 190
Asn Leu Pro Phe Leu Thr Glu Ile His Ala Asp Asn Asn Ser Leu Lys
195 200 205
Thr Leu Pro Asp Leu Pro Pro Ser Leu Lys Thr Leu Asn Val Arg Glu
210 215 220
Asn Tyr Leu Thr Asp Leu Pro Glu Leu Pro Gln Ser Leu Thr Phe Leu
225 230 235 240
Asp Val Ser Asp Asn Ile Phe Ser Gly Leu Ser Glu Leu Pro Pro Asn
245 250 255
Leu Tyr Tyr Leu Asp Ala Ser Ser Asn Gly Ile Arg Ser Leu Cys Asp
260 265 270
Leu Pro Pro Ser Leu Val Glu Leu Asp Val Arg Asp Asn Gln Leu Ile
275 280 285
Glu Leu Pro Ala Leu Pro Pro His Leu Glu Arg Leu Ile Ala Ser Leu
290 295 300
Asn His Leu Ala Glu Val Pro Glu Leu Pro Gln Asn Leu Lys Gln Leu
305 310 315 320
His Val Glu His Asn Ala Leu Arg Glu Phe Pro Asp Ile Pro Glu Ser
325 330 335
Val Glu Asp Leu Arg Met Asp Ser Glu Arg Val Thr Asp Thr Tyr Glu
340 345 350
Phe Ala His Glu Thr Thr Asp Lys Leu Glu Asp Asp Val Phe Glu
355 360 365
<210> 16
<211> 1104
<212> DNA
<213> 鼠疫耶尔森氏菌
<400> 16
atgtttataa ctccaagaaa tgtatctaat acttttttgc aagaaccatt acgtcattct 60
tctgatttaa ctgagatgcc ggttgaagca gaaaatgtta aatctaagac tgaatattat 120
aatgcatggg cggtatggga acgaaatgcc cctccgggga atggtgaaca gagggaaatg 180
gcggtttcaa ggttacgcga ttgcctggac cgacaagccc atgagctaga actaaataat 240
ctggggctga gttctttgcc ggaattacct ccgcatttag agagtttagt ggcgtcatgt 300
aattctctta cagaattacc ggaattgccg cagagcctga aatcacttca agttgataat 360
aacaatctga aggcattatc cgatttacct ccttcactgg aatttcttgc tgctggtaat 420
aatcagctgg aagaattgcc agagttgcaa aactcgtcct tcttgaaaat tattgatgtt 480
gataacaatt cactgaaaaa actacctgat ttacctcctt cactggaatt tcttgctgct 540
ggtaataatc agctggaaga attgtcagag ttacaaaact tgccattctt gactgagatt 600
catgctgata acaattcact gaaaacatta cccgatttac ccccttccct gaaaacactt 660
aatgtcagag aaaattattt aactgatctg ccagaattac cgcagagttt aaccttctta 720
gatgtttctg ataatatttt ttctggatta tcggaattgc caccaaactt gtattatctc 780
gatgcatcca gcaatggaat aagatcctta tgcgatttac ccccttcact ggtagaactt 840
gatgtcagag ataatcagtt gatcgaactg ccagcgttac ctccacactt agaacgttta 900
atcgcttcac ttaatcatct tgctgaagta cctgaattgc cgcaaaacct gaaacagctc 960
cacgtagagc acaacgctct aagagagttt cccgatatac ctgagtcagt ggaagatctt 1020
cggatggact ctgaacgtgt aactgataca tatgaatttg ctcatgagac tacagacaaa 1080
cttgaagatg atgtatttga gtag 1104
<210> 17
<211> 86
<212> PRT
<213> 鼠疫耶尔森氏菌
<400> 17
Met Phe Ile Asn Pro Arg Asn Val Ser Asn Thr Phe Leu Gln Glu Pro
1 5 10 15
Leu Arg His Ser Ser Asn Leu Thr Glu Met Pro Val Glu Ala Glu Asn
20 25 30
Val Lys Ser Lys Thr Glu Tyr Tyr Asn Ala Trp Ser Glu Trp Glu Arg
35 40 45
Asn Ala Pro Pro Gly Asn Gly Glu Gln Arg Glu Met Ala Val Ser Arg
50 55 60
Leu Arg Asp Cys Leu Asp Arg Gln Ala His Glu Leu Glu Leu Asn Asn
65 70 75 80
Leu Gly Leu Ser Ser Leu
85
<210> 18
<211> 258
<212> DNA
<213> 鼠疫耶尔森氏菌
<400> 18
atgttcatca ctccacgcaa tgtatctaac acctttctgc aggaaccgct gcgtcattct 60
agcgacctga ccgaaatgcc agttgaagcg gagaacgtga aatctaagac tgaatactac 120
aacgcgtggg cagtatggga gcgcaatgca ccaccaggta acggtgaaca gcgtgaaatg 180
gcagtaagcc gtctgcgtga ttgcctggat cgccaggctc acgagctgga gctgaacaac 240
ctgggtctgt ctagcctg 258
<210> 19
<211> 131
<212> PRT
<213> 鼠疫耶尔森氏菌
<400> 19
Met Phe Ile Asn Pro Arg Asn Val Ser Asn Thr Phe Leu Gln Glu Pro
1 5 10 15
Leu Arg His Ser Ser Asn Leu Thr Glu Met Pro Val Glu Ala Glu Asn
20 25 30
Val Lys Ser Lys Thr Glu Tyr Tyr Asn Ala Trp Ser Glu Trp Glu Arg
35 40 45
Asn Ala Pro Pro Gly Asn Gly Glu Gln Arg Glu Met Ala Val Ser Arg
50 55 60
Leu Arg Asp Cys Leu Asp Arg Gln Ala His Glu Leu Glu Leu Asn Asn
65 70 75 80
Leu Gly Leu Ser Ser Leu Pro Glu Leu Pro Pro His Leu Glu Ser Leu
85 90 95
Val Ala Ser Cys Asn Ser Leu Thr Glu Leu Pro Glu Leu Pro Gln Ser
100 105 110
Leu Lys Ser Leu Leu Val Asp Asn Asn Asn Leu Lys Ala Leu Ser Asp
115 120 125
Leu Pro Pro
130
<210> 20
<211> 393
<212> DNA
<213> 鼠疫耶尔森氏菌
<400> 20
atgtttatta acccgcgcaa cgtgagcaac acctttctgc aggaaccgct gcgccatagc 60
agcaacctga ccgaaatgcc ggtggaagcg gaaaacgtga aaagcaaaac cgaatattat 120
aacgcgtgga gcgaatggga acgcaacgcg ccgccgggca acggcgaaca gcgcgaaatg 180
gcggtgagcc gcctgcgcga ttgcctggat cgccaggcgc atgaactgga actgaacaac 240
ctgggcctga gcagcctgcc ggaactgccg ccgcatctgg aaagcctggt ggcgagctgc 300
aacagcctga ccgaactgcc ggaactgccg cagagcctga aaagcctgct ggtggataac 360
aacaacctga aagcgctgag cgatctgccg ccg 393
<210> 21
<211> 565
<212> PRT
<213> 弗氏志贺氏菌
<400> 21
Met Phe Ser Val Asn Asn Thr His Ser Ser Val Ser Cys Ser Pro Ser
1 5 10 15
Ile Asn Ser Asn Ser Thr Ser Asn Glu His Tyr Leu Arg Ile Leu Thr
20 25 30
Glu Trp Glu Lys Asn Ser Ser Pro Gly Glu Glu Arg Gly Ile Ala Phe
35 40 45
Asn Arg Leu Ser Gln Cys Phe Gln Asn Gln Glu Ala Val Leu Asn Leu
50 55 60
Ser Asp Leu Asn Leu Thr Ser Leu Pro Glu Leu Pro Lys His Ile Ser
65 70 75 80
Ala Leu Ile Val Glu Asn Asn Lys Leu Thr Ser Leu Pro Lys Leu Pro
85 90 95
Ala Phe Leu Lys Glu Leu Asn Ala Asp Asn Asn Arg Leu Ser Val Ile
100 105 110
Pro Glu Leu Pro Glu Ser Leu Thr Thr Leu Ser Val Arg Ser Asn Gln
115 120 125
Leu Glu Asn Leu Pro Val Leu Pro Asn His Leu Thr Ser Leu Phe Val
130 135 140
Glu Asn Asn Arg Leu Tyr Asn Leu Pro Ala Leu Pro Glu Lys Leu Lys
145 150 155 160
Phe Leu His Val Tyr Tyr Asn Arg Leu Thr Thr Leu Pro Asp Leu Pro
165 170 175
Asp Lys Leu Glu Ile Leu Cys Ala Gln Arg Asn Asn Leu Val Thr Phe
180 185 190
Pro Gln Phe Ser Asp Arg Asn Asn Ile Arg Gln Lys Glu Tyr Tyr Phe
195 200 205
His Phe Asn Gln Ile Thr Thr Leu Pro Glu Ser Phe Ser Gln Leu Asp
210 215 220
Ser Ser Tyr Arg Ile Asn Ile Ser Gly Asn Pro Leu Ser Thr Arg Val
225 230 235 240
Leu Gln Ser Leu Gln Arg Leu Thr Ser Ser Pro Asp Tyr His Gly Pro
245 250 255
Gln Ile Tyr Phe Ser Met Ser Asp Gly Gln Gln Asn Thr Leu His Arg
260 265 270
Pro Leu Ala Asp Ala Val Thr Ala Trp Phe Pro Glu Asn Lys Gln Ser
275 280 285
Asp Val Ser Gln Ile Trp His Ala Phe Glu His Glu Glu His Ala Asn
290 295 300
Thr Phe Ser Ala Phe Leu Asp Arg Leu Ser Asp Thr Val Ser Ala Arg
305 310 315 320
Asn Thr Ser Gly Phe Arg Glu Gln Val Ala Ala Trp Leu Glu Lys Leu
325 330 335
Ser Ala Ser Ala Glu Leu Arg Gln Gln Ser Phe Ala Val Ala Ala Asp
340 345 350
Ala Thr Glu Ser Cys Glu Asp Arg Val Ala Leu Thr Trp Asn Asn Leu
355 360 365
Arg Lys Thr Leu Leu Val His Gln Ala Ser Glu Gly Leu Phe Asp Asn
370 375 380
Asp Thr Gly Ala Leu Leu Ser Leu Gly Arg Glu Met Phe Arg Leu Glu
385 390 395 400
Ile Leu Glu Asp Ile Ala Arg Asp Lys Val Arg Thr Leu His Phe Val
405 410 415
Asp Glu Ile Glu Val Tyr Leu Ala Phe Gln Thr Met Leu Ala Glu Lys
420 425 430
Leu Gln Leu Ser Thr Ala Val Lys Glu Met Arg Phe Tyr Gly Val Ser
435 440 445
Gly Val Thr Ala Asn Asp Leu Arg Thr Ala Glu Ala Met Val Arg Ser
450 455 460
Arg Glu Glu Asn Glu Phe Thr Asp Trp Phe Ser Leu Trp Gly Pro Trp
465 470 475 480
His Ala Val Leu Lys Arg Thr Glu Ala Asp Arg Trp Ala Gln Ala Glu
485 490 495
Glu Gln Lys Tyr Glu Met Leu Glu Asn Glu Tyr Ser Gln Arg Val Ala
500 505 510
Asp Arg Leu Lys Ala Ser Gly Leu Ser Gly Asp Ala Asp Ala Glu Arg
515 520 525
Glu Ala Gly Ala Gln Val Met Arg Glu Thr Glu Gln Gln Ile Tyr Arg
530 535 540
Gln Leu Thr Asp Glu Val Leu Ala Leu Arg Leu Ser Glu Asn Gly Ser
545 550 555 560
Arg Leu His His Ser
565
<210> 22
<211> 1698
<212> DNA
<213> 弗氏志贺氏菌
<400> 22
atgttctctg taaataatac acactcatca gtttcttgct ccccctctat taactcaaac 60
tcaaccagta atgaacatta tctgagaatc ctgactgaat gggaaaagaa ctcttctccc 120
ggggaagagc gaggcattgc ttttaacaga ctctcccagt gctttcagaa tcaagaagca 180
gtattaaatt tatcagacct aaatttgacg tctcttcccg aattaccaaa gcatatttct 240
gctttgattg tagaaaataa taaattaaca tcattgccaa agctgcctgc atttcttaaa 300
gaacttaatg ctgataataa caggctttct gtgataccag aacttcctga gtcattaaca 360
actttaagtg ttcgttctaa tcaactggaa aaccttcctg ttttgccaaa ccatttaaca 420
tcattatttg ttgaaaataa caggctatat aacttaccgg ctcttcccga aaaattgaaa 480
tttttacatg tttattataa caggctgaca acattacccg acttaccgga taaactggaa 540
attctctgtg ctcagcgcaa taatctggtt acttttcctc aattttctga tagaaacaat 600
atcagacaaa aggaatatta ttttcatttt aatcagataa ccactcttcc ggagagtttt 660
tcacaattag attcaagtta caggattaat atttcaggga atccattgtc gactcgcgtt 720
ctgcaatccc tgcaaagatt aacctcttcg ccggactacc acggcccgca gatttacttc 780
tccatgagtg acggacaaca gaatacactc catcgccccc tggctgatgc cgtgacagca 840
tggttcccgg aaaacaaaca atctgatgta tcacagatat ggcatgcttt tgaacatgaa 900
gagcatgcca acaccttttc cgcgttcctt gaccgccttt ccgataccgt ctctgcacgc 960
aatacctccg gattccgtga acaggtcgct gcatggctgg aaaaactcag tgcctctgcg 1020
gagcttcgac agcagtcttt cgctgttgct gctgatgcca ctgagagctg tgaggaccgt 1080
gtcgcgctca catggaacaa tctccggaaa accctcctgg tccatcaggc atcagaaggc 1140
cttttcgata atgataccgg cgctctgctc tccctgggca gggaaatgtt ccgcctcgaa 1200
attctggagg acattgcccg ggataaagtc agaactctcc attttgtgga tgagatagaa 1260
gtctacctgg ccttccagac catgctcgca gagaaacttc agctctccac tgccgtgaag 1320
gaaatgcgtt tctatggcgt gtcgggagtg acagcaaatg acctccgcac tgccgaagct 1380
atggtcagaa gccgtgaaga gaatgaattt acggactggt tctccctctg gggaccatgg 1440
catgctgtac tgaagcgtac ggaagctgac cgctgggcgc aggcagaaga gcagaagtat 1500
gagatgctgg agaatgagta ctctcagagg gtggctgacc ggctgaaagc atcaggtctg 1560
agcggtgatg cggatgcgga gagggaagcc ggtgcacagg tgatgcgtga gactgaacag 1620
cagatttacc gtcagctgac tgacgaggta ctggccctgc gattgtctga aaacggctca 1680
cgactgcacc attcataa 1698
<210> 23
<211> 379
<212> PRT
<213> 人工序列
<220>
<223> 人工序列的描述:合成的多肽
<400> 23
Met Phe Ile Thr Pro Arg Asn Val Ser Asn Thr Phe Leu Gln Glu Pro
1 5 10 15
Leu Arg His Ser Ser Asp Leu Thr Glu Met Pro Val Glu Ala Glu Asn
20 25 30
Val Lys Ser Lys Thr Glu Tyr Tyr Asn Ala Trp Ala Val Trp Glu Arg
35 40 45
Asn Ala Pro Pro Gly Asn Gly Glu Gln Arg Glu Met Ala Val Ser Arg
50 55 60
Leu Arg Asp Cys Leu Asp Arg Gln Ala His Glu Leu Glu Leu Asn Asn
65 70 75 80
Leu Gly Leu Ser Ser Leu Pro Glu Leu Pro Pro His Leu Glu Ser Leu
85 90 95
Val Ala Ser Cys Asn Ser Leu Thr Glu Leu Pro Glu Leu Pro Gln Ser
100 105 110
Leu Lys Ser Leu Gln Val Asp Asn Asn Asn Leu Lys Ala Leu Ser Asp
115 120 125
Leu Pro Pro Ser Leu Glu Phe Leu Ala Ala Gly Asn Asn Gln Leu Glu
130 135 140
Glu Leu Pro Glu Leu Gln Asn Ser Ser Phe Leu Lys Ile Ile Asp Met
145 150 155 160
Ser Ile Glu Ile Lys Met Ile Ser Pro Ile Lys Asn Ile Lys Asn Val
165 170 175
Phe Pro Ile Asn Thr Ala Asn Thr Glu Tyr Ile Val Arg Asn Ile Tyr
180 185 190
Pro Arg Val Glu His Gly Tyr Phe Asn Glu Ser Pro Asn Ile Tyr Asp
195 200 205
Lys Lys Tyr Ile Ser Gly Ile Thr Arg Ser Met Ala Gln Leu Lys Ile
210 215 220
Glu Glu Phe Ile Asn Glu Lys Ser Arg Arg Leu Asn Tyr Met Lys Thr
225 230 235 240
Met Tyr Ser Pro Cys Pro Glu Asp Phe Gln Pro Ile Ser Arg Asp Glu
245 250 255
Ala Ser Thr Pro Glu Gly Ser Trp Leu Thr Val Ile Ser Gly Lys Arg
260 265 270
Pro Met Gly Gln Phe Ser Val Asp Ser Leu Tyr His Pro Asp Leu His
275 280 285
Ala Leu Cys Glu Leu Pro Glu Ile Ser Cys Lys Ile Phe Pro Lys Glu
290 295 300
Asn Ser Asp Phe Leu Tyr Ile Ile Val Val Phe Arg Asn Asp Ser Pro
305 310 315 320
Gln Gly Glu Leu Arg Ala Asn Arg Phe Ile Glu Leu Tyr Asp Ile Lys
325 330 335
Arg Glu Ile Met Gln Val Leu Arg Asp Glu Ser Pro Glu Leu Lys Ser
340 345 350
Ile Lys Ser Glu Ile Ile Ile Ala Arg Glu Met Gly Glu Leu Phe Ser
355 360 365
Tyr Ala Ser Glu Glu Ile Asp Ser Tyr Ile Lys
370 375
<210> 24
<211> 345
<212> PRT
<213> 人工序列
<220>
<223> 人工序列的描述:合成的多肽
<400> 24
Met Phe Ile Thr Pro Arg Asn Val Ser Asn Thr Phe Leu Gln Glu Pro
1 5 10 15
Leu Arg His Ser Ser Asp Leu Thr Glu Met Pro Val Glu Ala Glu Asn
20 25 30
Val Lys Ser Lys Thr Glu Tyr Tyr Asn Ala Trp Ala Val Trp Glu Arg
35 40 45
Asn Ala Pro Pro Gly Asn Gly Glu Gln Arg Glu Met Ala Val Ser Arg
50 55 60
Leu Arg Asp Cys Leu Asp Arg Gln Ala His Glu Leu Glu Leu Asn Asn
65 70 75 80
Leu Gly Leu Ser Ser Leu Pro Glu Leu Pro Pro His Leu Glu Ser Leu
85 90 95
Val Ala Ser Cys Asn Ser Leu Thr Glu Leu Pro Glu Leu Pro Gln Ser
100 105 110
Leu Lys Ser Leu Gln Val Asp Asn Asn Asn Leu Lys Ala Leu Ser Asp
115 120 125
Leu Pro Pro Ser Leu Glu Phe Leu Ala Ala Gly Asn Asn Gln Leu Glu
130 135 140
Glu Leu Pro Glu Leu Gln Asn Ser Ser Phe Leu Lys Ile Ile Asp Lys
145 150 155 160
Ile Pro Ser Leu Gln Ser Asn Phe Asn Phe Ser Ala Pro Ala Gly Tyr
165 170 175
Ser Ala Pro Ile Ala Pro Asn Arg Ala Glu Asn Ala Tyr Ala Asp Tyr
180 185 190
Val Leu Asp Ile Gly Lys Arg Ile Pro Leu Ser Ala Ala Asp Leu Ser
195 200 205
Asn Val Tyr Glu Ser Val Ile Arg Ala Val His Asp Ser Arg Ser Arg
210 215 220
Leu Ile Asp Gln His Thr Val Asp Met Ile Gly Asn Thr Val Leu Asp
225 230 235 240
Ala Leu Ser Arg Ser Gln Thr Phe Arg Asp Ala Val Ser Tyr Gly Ile
245 250 255
His Asn Glu Lys Val His Ile Gly Cys Ile Lys Tyr Arg Asn Glu Tyr
260 265 270
Glu Leu Asn Glu Glu Ser Ser Val Lys Ile Asp Asp Ile Gln Ser Leu
275 280 285
Thr Cys Asn Glu Leu Tyr Glu Tyr Asp Val Gly Gln Glu Pro Ile Phe
290 295 300
Pro Ile Cys Glu Ala Gly Glu Asn Asp Asn Glu Glu Pro Tyr Val Ser
305 310 315 320
Phe Ser Val Ala Pro Asp Thr Asp Ser Tyr Glu Met Pro Ser Trp Gln
325 330 335
Glu Gly Leu Ile His Glu Ile Ile His
340 345
<210> 25
<211> 30
<212> PRT
<213> 人工序列
<220>
<223> 人工序列的描述:合成的多肽
<220>
<221> MISC_FEATURE
<222> (1)..(30)
<223> 该序列可包括1-6个"Gly Gly Gly Gly Ser"重复单元
<400> 25
Gly Gly Gly Gly Ser Gly Gly Gly Gly Ser Gly Gly Gly Gly Ser Gly
1 5 10 15
Gly Gly Gly Ser Gly Gly Gly Gly Ser Gly Gly Gly Gly Ser
20 25 30
<210> 26
<211> 25
<212> PRT
<213> 人工序列
<220>
<223> 人工序列的描述:合成肽
<220>
<221> MOD_RES
<222> (1)..(4)
<223> Thr或Ser
<220>
<221> MOD_RES
<222> (6)..(9)
<223> Thr或Ser
<220>
<221> MOD_RES
<222> (11)..(14)
<223> Thr或Ser
<220>
<221> MOD_RES
<222> (16)..(19)
<223> Thr或Ser
<220>
<221> MOD_RES
<222> (21)..(24)
<223> Thr或Ser
<220>
<221> MISC_FEATURE
<222> (1)..(25)
<223> 该序列可包括1-5个"Xaa Xaa Xaa Xaa Gly"重复单元,其中Xaa可以是Thr或Ser
<220>
<223> 有关替换和优选实施方案的详细说明,请参见提交的说明书
<400> 26
Xaa Xaa Xaa Xaa Gly Xaa Xaa Xaa Xaa Gly Xaa Xaa Xaa Xaa Gly Xaa
1 5 10 15
Xaa Xaa Xaa Gly Xaa Xaa Xaa Xaa Gly
20 25
<210> 27
<211> 5
<212> PRT
<213> 人工序列
<220>
<223> 人工序列的描述:合成肽
<400> 27
Ser Ser Ser Ser Gly
1 5
<210> 28
<211> 5
<212> PRT
<213> 人工序列
<220>
<223> 人工序列的描述:合成肽
<400> 28
Pro Ala Pro Ala Pro
1 5
<210> 29
<211> 15
<212> PRT
<213> 人工序列
<220>
<223> 人工序列的描述:合成肽
<220>
<221> MISC_FEATURE
<222> (1)..(14)
<223> 该区域可包括2-7个“Pro Thr”重复单元
<400> 29
Pro Thr Pro Thr Pro Thr Pro Thr Pro Thr Pro Thr Pro Thr Pro
1 5 10 15
<210> 30
<211> 27
<212> PRT
<213> 人工序列
<220>
<223> 人工序列的描述:合成肽
<220>
<221> MISC_FEATURE
<222> (2)..(26)
<223> 该区域可包括1-5个"Glu Ala Ala Ala Lys"重复单元
<400> 30
Ala Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys
1 5 10 15
Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys Ala
20 25
Claims (47)
1.一种包含两个或更多个截短的T3SS细菌效应多肽的构建体,其中每个截短的T3SS细菌效应多肽包括相应的全长T3SS细菌效应多肽的一部分。
2.根据权利要求1所述的构建体,其中所述截短的T3SS细菌效应多肽保留所述相应的全长T3SS细菌效应多肽的一种或多种活性。
3.根据权利要求1所述的构建体,其中所述T3SS细菌效应多肽选自由E3泛素连接酶、RhoGTP酶调节剂、半胱氨酸甲基转移酶、锌金属蛋白酶、乙酰转移酶、O-GlcNac转移酶和LRR基序结合和螯合多肽组成的组。
4.根据权利要求3所述的构建体,其中所述E3泛素连接酶是IpaH7.8或IpaH9.8;所述RhoGTP酶调节剂是YopE;所述半胱氨酸甲基转移酶是OspZ或NleE;所述锌金属蛋白酶是NleC;所述乙酰转移酶是YopJ;所述O-GlcNac转移酶是NleB;并且所述LRR基序结合和螯合多肽是YopM。
5.根据权利要求1所述的构建体,其中所述构建体包括与截短的T3SS半胱氨酸甲基转移酶多肽连接的截短的YopM多肽。
6.根据权利要求5所述的构建体,其中所述截短的T3SS半胱氨酸甲基转移酶多肽包括具有如SEQ ID NO.:3中所示的氨基酸序列的OspZ多肽的一部分。
7.根据权利要求6所述的构建体,其中所述截短的OspZ多肽包括与SEQ ID NO.:3的氨基酸226-446至少90%相同的氨基酸序列。
8.根据权利要求7所述的构建体,其中所述截短的OspZ多肽具有如SEQ ID NO.:3的氨基酸226-446中所示的氨基酸序列。
9.根据权利要求5所述的构建体,其中所述构建体包括如SEQ ID NO.:23中所示的氨基酸序列。
10.根据权利要求1所述的构建体,其中所述构建体包括与截短的T3SS锌金属蛋白酶多肽连接的截短的YopM多肽。
11.根据权利要求10所述的构建体,其中所述截短的T3SS锌金属蛋白酶多肽包括具有如SEQ ID NO.:5中所示的氨基酸序列的NleC多肽的一部分。
12.根据权利要求11所述的构建体,其中所述截短的NleC多肽包括与SEQ ID NO.:5的氨基酸2-187至少90%相同的氨基酸序列。
13.根据权利要求12所述的构建体,其中所述截短的NleC多肽具有如SEQ ID NO.:5的氨基酸2-187中所示的氨基酸序列。
14.根据权利要求10所述的构建体,其中所述构建体包括如SEQ ID NO.:24中所示的氨基酸序列。
15.根据权利要求1所述的构建体,其中所述构建体包括与截短的T3SS O-GlcNac转移酶连接的截短的YopM多肽。
16.根据权利要求15所述的构建体,其中所述截短的T3SS O-GlcNac转移酶包括具有如SEQ ID NO.:9中所示的氨基酸序列的NleB多肽的一部分。
17.根据权利要求16所述的构建体,其中所述截短的NleB多肽包括与SEQ ID NO.:9的氨基酸2-226至少90%相同的氨基酸序列。
18.根据权利要求17所述的构建体,其中所述截短的NleB多肽具有如SEQ ID NO.:9的氨基酸2-226中所示的氨基酸序列。
19.根据权利要求1所述的构建体,其中所述构建体包括与截短的第二T3SS E3泛素连接酶连接的截短的第一T3SS E3泛素连接酶多肽。
20.根据权利要求19所述的构建体,其中所述第一和第二截短的T3SS E3泛素连接酶多肽是不同的。
21.根据权利要求19所述的构建体,其中所述截短的第一E3泛素连接酶包括具有如SEQID NO.:11中所示的氨基酸序列的IpaH9.8多肽的一部分。
22.根据权利要求20所述的构建体,其中所述截短的第一IpaH9.8多肽包括与SEQ IDNO.:11的氨基酸56-228至少90%相同的氨基酸序列。
23.根据权利要求22所述的构建体,其中所述截短的第二E3泛素连接酶包括具有如SEQID NO.:13中所示的氨基酸序列的IpaH4.5多肽的一部分。
24.根据权利要求23所述的构建体,其中所述截短的第二IpaH4.5多肽包括与SEQ IDNO.:13的氨基酸62-270至少90%相同的氨基酸序列。
25.根据权利要求1所述的构建体,其中所述构建体包括与半胱氨酸甲基转移酶连接的RhoGTP酶调节剂。
26.根据权利要求25所述的构建体,其中RhoGTP酶调节剂是具有如SEQ ID NO.:1中所示的氨基酸序列的YopE多肽。
27.根据权利要求26所述的构建体,其中所述半胱氨酸甲基转移酶是具有如SEQ IDNO.:5中所示的氨基酸序列的OspZ多肽。
28.根据权利要求1所述的构建体,其中所述构建体包括与截短的乙酰转移酶多肽连接的截短的YopM多肽。
29.根据权利要求28所述的构建体,其中所述截短的乙酰转移酶包括具有如SEQ IDNO.:9中所示的氨基酸序列的YopJ多肽的一部分。
30.根据权利要求28所述的构建体,其中所述截短的YopJ多肽包括在SEQ ID NO.:9的半胱氨酸172处的点突变。
31.根据权利要求1-18中任一项所述的构建体,其中所述截短的YopM多肽具有与如SEQID NO.:19中所示的氨基酸序列至少90%相同的氨基酸序列。
32.根据权利要求31所述的构建体,其中所述截短的YopM多肽具有如SEQ ID NO.:19中所示的氨基酸序列。
33.根据权利要求1-32中任一项所述的构建体,进一步包括蛋白质转导结构域。
34.根据权利要求27所述的构建体,其中所述蛋白质转导结构域是YopM蛋白质转导结构域。
35.根据权利要求34所述的构建体,其中所述YopM蛋白质转导结构域具有如SEQ IDNO.:17中所示的氨基酸序列。
36.根据权利要求27所述的构建体,其中所述蛋白质转导结构域是IpaH9.8蛋白质转导结构域。
37.根据权利要求27所述的构建体,其中所述IpaH9.8蛋白质转导结构域具有如SEQ IDNO.:11的氨基酸2-56中所示的氨基酸序列。
38.根据权利要求1-37中任一项所述的构建体,其中所述构建体包括融合蛋白。
39.根据权利要求1-38中任一项所述的构建体,其中所述两个或更多个截短的T3SS细菌效应多肽通过接头接合。
40.根据权利要求39所述的构建体,其中所述接头是可切割的接头。
41.根据权利要求40所述的构建体,其中所述可切割的接头是pH敏感性接头。
42.根据权利要求41所述的构建体,其中所述pH敏感性接头包括肼、基于氨基磷酸酯的接头或硫代马来酸。
43.根据权利要求39所述的构建体,其中所述接头是共价键。
44.一种核酸,所述核酸编码权利要求1-43中任一项的构建体。
45.一种药物组合物,所述药物组合物包括权利要求1-44中任一项的构建体和药学上可接受的载体。
46.一种治疗患有炎症性疾病的受试者的方法,所述方法包括施用治疗有效量的权利要求45的药物组合物。
47.根据权利要求46所述的方法,其中所述炎症性疾病是皮肤疾病、胃肠疾病或肌肉骨骼疾病。
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