CN102816221B - 鸡柔嫩艾美耳球虫ma1基因、载体、重组菌株和蛋白及其应用 - Google Patents
鸡柔嫩艾美耳球虫ma1基因、载体、重组菌株和蛋白及其应用 Download PDFInfo
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- CN102816221B CN102816221B CN201210279040.5A CN201210279040A CN102816221B CN 102816221 B CN102816221 B CN 102816221B CN 201210279040 A CN201210279040 A CN 201210279040A CN 102816221 B CN102816221 B CN 102816221B
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Abstract
本发明公开了鸡柔嫩艾美耳球虫MA1基因、载体、重组菌株和蛋白及其应用。EtMA1蛋白的氨基酸序列如SEQIDNO:2所示，或者是SEQIDNO:2所示的氨基酸序列经取代、缺失和/或增加一个或多个氨基酸和/或末端修饰且具有同等活性的蛋白。本发明还公开了EtMA1膜外功能域蛋白（EtMA1-Outsidedomain），其氨基酸序列如SEQIDNO:4所示，或者是SEQIDNO:4所示的氨基酸序列经取代、缺失和/或增加一个或多个氨基酸和/或末端修饰且具有同等活性的蛋白。EtMA1蛋白和EtMA1膜外功能域蛋白可应用于制备抗艾美耳球虫药物，通过免疫EtMA1膜外功能域蛋白，能有效控制鸡体感染柔嫩艾美耳球虫病，大大降低养鸡场抗球虫药的使用量，有效控制鸡球虫病。
Description
技术领域
本发明属于基因工程技术领域，具体涉及鸡柔嫩艾美耳球虫MA1蛋白（EtMA1蛋白）的膜外功能域蛋白（EtMA1-Outside domain）及其编码基因、含有该基因的载体、重组菌株及其应用；还涉及鸡柔嫩艾美耳球虫MA1蛋白及其编码基因、含有该基因的载体、重组菌株及其应用。
背景技术
鸡球虫病是一种严重危害集约化养鸡业、世界性流行的寄生虫病，在无预防措施或预防失败（如因抗药性问题致药物无效）的情况下，鸡发病率可达30%-100%，致死率可高达80%。全球每年因球虫病引起的经济损失超过35亿美元以上，我国因球虫病所致养鸡业的经济损失估计可高达35亿元人民币以上。
目前，世界公认的鸡球虫有7种：柔嫩艾美耳球虫（E.tenella），毒害艾美耳球虫（E.necatrix），巨型艾美耳球虫（E.maxima），堆型艾美耳球虫（E.acervulina），布氏艾美耳球虫（E.brunette），早熟艾美耳球虫（E.praecox），和缓艾美耳球虫（E.mitis），其中柔嫩艾美耳球虫主要侵害盲肠，是毒力最强、危害最大的鸡球虫，也是研究最多的虫种。
长久以来，国内外鸡球虫病的预防一直采用 “在饲料中添加抗球虫药、实行以饲料厂为中心控制环节”的技术方法，但该方法已遭遇到球虫抗药性的严峻挑战，在某些地区甚至已到无药可用的困难境地。免疫控制技术作为一种“新型的 (novel)可持续的(sustainable)
”的球虫病控制方法已得到越来越多的应用。尽管以鸡球虫病活卵囊疫苗为主要技术载体的免疫控制技术正在逐步推广应用，但这类疫苗完全以鸡体作为唯一制苗载体，普遍存在生产成本高、保存运输困难、接种使用后对鸡体生长有明显抑制、散毒危险和难以质量控制等几乎不可逾越的障碍。
因此，开发新药或新型分子疫苗是球虫病研究的主要方向，然而迄今为止，人们对艾美耳球虫的药物靶标分子或候选疫苗分子都知之甚少。
顶膜抗原（Apical
Membrane Antigen，AMA）是一种在顶复门原虫中高度保守的微线分泌蛋白，可以和棒状体分泌的颈部蛋白共同形成“运动结合体（Moving Junction）”，共同完成虫体与宿主细胞的粘附作用, 是协助虫体进入宿主细胞的关键物质。
近几年，通过对鸡球虫基因文库和蛋白质组的研究，开始对鸡球虫 AMA1 有所了解，但目前为止未有系统性研究的报道。Ng 等在对柔嫩艾美耳球虫的子孢子和裂殖子的 cDNA 文库比较研究时发现，子孢子阶段有一条 EST 序列（基因登陆号：BG589）与 AMA1 具有相似性，而在裂殖子文库中未找到相同的 EST 序列。在Bromley等的蛋白组学研究中发现，柔嫩艾美耳球虫不只一种顶膜抗原，并且出现在不同的发育阶段，所以分析可能这些蛋白分别在不同的入侵阶段发挥着类似的作用。Blake 等利用基因图谱法对球虫中免疫力最强的巨型艾美耳球虫保护性抗原进行筛选，将两株具有不同生物学特性的巨型艾美耳球虫在鸡体内进行杂交后的基因组，与柔嫩艾美耳球虫的基因组进行比对，结果发现与 AMA1 具有同源性的抗原可以优先刺激机体产生免疫力，可以优先作为疫苗和药物开发的研究靶标。本课题组在对广东株柔嫩艾美耳球虫裂殖子cDNA的克隆时，获得了多个类似顶膜抗原的基因，其中鸡柔嫩艾美耳球虫MA1蛋白（EtMA1 ）的氨基酸序列与EtAMA1具有高度保守的序列，推测为EtAMA1的家族蛋白。在对其功能进行研究时，发现免疫EtMA1蛋白的膜外功能结构域可以有效预防鸡体感染鸡球虫病。而迄今为止，尚无任何其他关于该蛋白对艾美耳球虫免疫保护性的研究。
发明内容
本发明的一个目的在于提供一种鸡柔嫩艾美耳球虫MA1蛋白（EtMA1蛋白）。
本发明的另一个目的在于提供编码上述EtMA1蛋白的基因。
本发明的目的在于提供一种EtMA1膜外功能域蛋白（EtMA1-Outside domain）。
本发明的另一个目的在于提供编码上述EtMA1膜外功能域蛋白的基因。
本发明的另一个目的在于提供含有EtMA1蛋白基因或EtMA1膜外功能域蛋白基因的克隆载体。
本发明的另一个目的在于提供一种EtMA1膜外功能域蛋白的制备方法。
本发明的另一个目的在于提供EtMA1蛋白和/或EtMA1膜外功能域蛋白在制备抗艾美耳球虫药物中的应用。
本发明所采用的技术方案是：
鸡柔嫩艾美耳球虫MA1蛋白（EtMA1蛋白），其氨基酸序列如SEQ ID NO:2所示，或者是SEQ ID NO:2所示的氨基酸序列经取代、缺失和/或增加一个或多个氨基酸和/或末端修饰且具有同等活性的蛋白。
编码上述EtMA1蛋白的基因，其核苷酸序列如SEQ ID NO：1所示。
EtMA1膜外功能域蛋白（EtMA1-Outside domain），其氨基酸序列如SEQ ID
NO:4所示，或者是SEQ ID NO:4所示的氨基酸序列经取代、缺失和/或增加一个或多个氨基酸和/或末端修饰且具有同等活性的蛋白。
编码上述EtMA1膜外功能域蛋白的基因，其核苷酸序列如SEQ ID NO:3所示。
一种克隆载体，含有EtMA1蛋白的编码基因或EtMA1膜外功能域蛋白的编码基因。
一种表达载体，含有EtMA1蛋白的编码基因或EtMA1膜外功能域蛋白的编码基因。
生产EtMA1膜外功能域蛋白的方法，包括将上述的表达载体导入宿主细胞中，表达得到EtMA1膜外功能域蛋白。
EtMA1蛋白或EtMA1膜外功能域蛋白在制备抗艾美耳球虫药物中的应用。
本发明的有益效果在于：
通过免疫EtMA1膜外功能域蛋白，能有效控制鸡体感染柔嫩艾美耳球虫病，大大降低养鸡场抗球虫药的使用量，有效控制鸡球虫病。
附图说明
图1是E.tenella裂殖子总RNA电泳结果；
图2是EtMA1基因全长ORF的PCR扩增鉴定结果（M.DL 2000 Marker；1. EtMA1的PCR产物）；
图3是 EtMA1的菌液PCR鉴定结果（M. DL 2000 Marker；1: EtMA1阳性菌）
图4是 EtMA1信号肽切割位点分析图；
图5是 EtMA1膜外结构域位点分析图；
图6 是EtMA1-Outside domain的PCR产物电泳图（M.DL
2000 Marker；1，2. EtMA1-Outside domain的PCR产物）；
图7是EtMA1-Outside domain的菌液PCR鉴定电泳图（M.DL
2000 Marker；5，6. EtMA1-Outside
domain菌液PCR结果）；
图8是 pColdI–EtMA1-Outside domain的酶切鉴定电泳图；
图9 是pColdI-EtMA1-Outside domain表达产物SDS-PAGE电泳分析图（M.蛋白质分子质量标准；1.未诱导菌液； 2. 诱导后菌液；3-9.纯化的融合蛋白；10.Western blot）。
具体实施方式
下面结合实施例对本发明作进一步的说明，但并不局限于此。
以下实施例中所采用的分子生物学实验技术包括PCR扩增、质粒提取、质粒转化、DNA片段连接、酶切、凝胶电泳等都采用常规的方法，具体可参见《分子克隆实验指南》(第三版) (Sambrook J, Russell
DW，Janssen
K, Argentine J.黄培堂等译，2002，北京：科学出版社)。及《现代分子生物学实验技术》（卢圣栋主编，第2版, 北京:中国协和医科大学出版社, 1999.）
实施例
1
1 EtMA1基因序列的克隆
1.1 PCR引物设计：根据预测的E.tenella棒状体颈部蛋白Et MA1基因的cDNA序列用Premier Primer 5.0软件设计，上海英骏生物工程公司合成：
上游引物MA1F：5’
ATGGAGGCTCTACGGGAAGGCTTC 3’ （SEQ ID NO:5）；
下游引物MA1R：5’
TTAGTAGTAGGCGTCGTGGGCGTTG 3’ （SEQ ID NO:6）。
1. 2 广东株E.tenella裂殖子（由广东省农业科学院兽医研究所寄生生物研究室保存）的分离和纯化：方法参照《鸡球虫病学》（索勋，李国清主编，北京:中国农业大学出版社，1997.）。
1.3 E.tenella裂殖子总RNA的提取：方法参照Invitrogen公司的TRIZOL LS® Reagent试剂盒说明书。分光光度计测定RNA浓度为1µg/ml；电泳结果显示分共为28S，18S和5S等3个大小不同的片段，条带清晰（如图1）。
1.4 RT-PCR扩增EtMA1全长ORF序列
以上述步骤抽提的E.tenella裂殖子的总RNA作为模板，用Takara公司的PrimeScriptTM1st
Strand cDNA Synthesis Kit试剂盒合成cDNA；以cDNA为模板，MA1F，MA1R为引物，TaKaRa LA TaqTM酶进行PCR扩增，反应条件：94℃预变性5min，94℃变性30s，55℃退火30s，72℃延伸2min，72℃延伸10min，35个循环。产物经1%琼脂糖凝胶电泳分析，结果扩增获得一条约1600bp的片段（见图2）。将PCR产物纯化回收后按试剂盒说明书连接至pMD18-T载体（Takara公司），再将连接产物转化入E.coli DH5α感受态细胞，得到E.coli DH5α（pMD18-T- EtMA1）。在Amp抗性平板上生长的白色菌落为可疑重组子，挑单菌落培养，以菌液为模板，以引物EtMA2F，EtMA2R进行菌液PCR扩增，获得大小约为1600bp的条带（见图3）。阳性菌送上海生工生物工程公司测序，EtMA1序列全长为1617bp，如SEQ ID
NO:1所示。
1.5 序列的分析
利用在线Translate tool（http://www.expasy.ch/tools/dna.html）翻译EtMA1序列，推测出其氨基酸序列（SEQ ID NO.2）。 利用信号肽在线分析软件SignalP 4.0
Server分析EtMA1信号肽，利用跨膜结构域在线分析软件TMHMM Server
v. 2.0分析EtMA1氨基酸序列，确定34aa～35aa为信号肽切割位点（图4），1aa-443aa为膜外结构域（图5），根据分析结果，本实验选取35aa~443aa 蛋白（SEQ ID NO:4所示，其编码基因为SEQ ID NO:3）作为EtMA1膜外功能域蛋白（EtMA1-Outside domain），进行表达和动物保护性实验。
2 EtMA1
膜外功能域蛋白
（
EtMA1
-Outside domain
）的制备
2.1 EtMA1-Outside domain的扩增
扩增引物：
EtMA1-Outside domainF：
5’ GGCGGATCCTCTAACGGCTCACAGGTAGCTTCC
3’（SEQ
ID NO:7）；
EtMA1-Outside domain R：
5' GGCAAGCTTTTAAGGGATGCTGCTGCAGTTGCAGT
3'（SEQ
ID NO:8）。
在引物的上游序列中分别引入BamH Ⅰ酶切位点，在下游引物引入HindIII酶切位点；并分别加“GGC”3个碱基以保证内切酶发挥作用。以上引物均由广东英俊生物技术公司合成。
以E.coli DH5α（pMD18-T-EtMA1）为模板，建立如下PCR反应扩增EtMA1-Outside domain的序列：
PCR反应程序：
94℃预变性5min→94℃变性30sec→55℃退火30sec→72℃延伸2min→72℃延伸10min→4℃；共35个循环。
将PCR产物以1%琼脂糖凝胶于100V电泳1h后，紫外检测仪下观察结果（图6）。产物大小约为1200bp，与预期的产物大小一致。
用DNA胶回收试剂盒回收目的片段，并进行酶切反应（表2）：
37℃孵育3h后，以1%琼脂糖凝胶于100V电泳1h后，用DNA胶回收试剂盒回收酶切后的目的片段。
2.2表达载体的处理
取一E.coli表达载体新鲜菌落分别接种于10ml LB液体培养基（含100μg/ml氨苄青霉素），37℃振荡培养12h后，用小量质粒制备试剂盒抽提质粒；建立如下酶切反应，酶切质粒：
37℃孵育3h后，将样品以1%琼脂糖凝胶100V电泳1h。用DNA胶回收试剂盒回收酶切后的质粒；
2.3酶切载体与酶切片段的连接：
酶切后的EtMA1-Outside domain与pCold
I按表4组成建立连接反应，构建pCold I-EtMA1-Outside
domain重组表达载体。
2.4 连接产物转化E.coli DH5α (CaCl2法)
（1）取一新鲜培养的E.coli DH 5α单菌落，接种于一个含有10 mL LB的液体培养基，37℃振荡培养过夜。
（2）取过夜培养的菌液1 mL接种于100 mL的LB液体培养基中，剧烈振摇2 h（300r/min）至OD600=0.4。
（3）将细菌转移到一个无菌的预冷的50 mL离心管中，在冰上放置10 min，使培养物冷却至0 ℃。
（4）于4 ℃以4 100 r/min离心10 min回收细胞。
（5）弃掉培养液，将管倒置1 min。
（6）每50 mL初始培养液用30 mL用冰预冷的0.1 mol/L CaCl2-MgCl2溶液重悬每份细胞沉淀。
（7）于4℃以4 100 r/min离心10 min回收细胞。
（8）弃掉培养液，将管倒置1 min。
（9）每50 mL初始培养物用2 mL预冷的0.1 mol/L CaCl2重悬每份细胞沉淀。
（10）从用CaCl2制备的感受态细胞悬液中吸取200 μL转移到1 mL的离心管中，每管加入连接产物5 μL，冰浴30 min。
（11）将离心管放入预加温至42 ℃水浴中，静置90 s。
（12）快速将管转移到冰浴中，使细胞冷却1～2 min。
（13）每管加800 μL SOC培养基，于37℃缓慢振摇培养温育45 min。
（14）将培养物涂布于LB琼脂平板（含100 μg/mL Amp），将平板置于室温直至液体被吸收，倒置平皿，37 ℃过夜培养（约12～16 h）。
2.5
PCR法鉴定转化克隆
挑取一些菌落分别接种于10ml
LB培养基中（含100μg/ml氨苄青霉素），37℃振荡过夜培养后，取培养物建立如下PCR反应筛选阳性克隆：
PCR反应程序：同上。
取PCR反应产物10μl，以1%琼脂糖凝胶于100V电泳1h后，紫外检测仪下观察结果（图7）。
2.6
PCR阳性克隆的酶切鉴定
取PCR反应阳性的细菌培养物用小量质粒制备试剂盒按操作说明抽提质粒，并进行如下酶切反应：
37℃孵育3h后，以1%琼脂糖凝胶100V电泳1h，用紫外检测仪下检测结果（图8）。
取PCR和限制性酶切鉴定均为阳性的克隆送上海英骏生物技术公司进行重组质粒的测序，分析重组质粒中外源基因的阅读框架是否改变，结果显示扩增所获得序列与预获得序列完全匹配，且正确插入pColdI载体。
2.7 pColdI-EtMA1-Outside domain在表达菌中的诱导表达
（1）将鉴定为阳性的新鲜单个重组菌落接种10ml LB培养基（含100μg/ml氨苄青霉素），37℃振荡培养过夜以获得饱和培养物，抽提重组质粒，转化入表达菌E.coli Rosetta(DE3)。挑取单克隆接种10ml LB培养基（含100μg/ml氨苄青霉素），37℃振荡培养过夜以获得饱和培养物；
（2）取过夜培养物，按1：100接种于500mL含有Amp(100μg/ml)的LB培养液，重组菌的接种液16℃培养；
（3）当培养物OD600值达到0.6左右时，取1ml未诱导的培养物转移至一离心管中，迅速在室温下以10000×g离心1min，弃上清液，细菌沉淀用适量1×SDS凝胶加样缓冲液中，100℃加热5min备用；
（4） 剩余的培养物加入终浓度为0.1mM的IPTG进行诱导表达， 16℃培养组过夜诱导培养。诱导结束后，根据各组OD600的值，分别取与诱导前相同细菌数的菌液，离心后4℃保存。
各重组菌落用IPTG诱导表达后经SDS-PAGE分析，在16℃条件下，使用终浓度为0.1mM的IPTG进行诱导，16h后pColdI-EtMA1-Outside domain重组质粒在E.coli Rosetta(DE3)中有目的蛋白表达（图9）。
2.8表达产物的处理
将诱导结束的培养液分别5500r/min离心15分钟，弃上清夜。菌体用20mL，0.1M Tris-HCL(pH7.4)的缓冲液(含50μL 100mM的PSMF)重悬，-20℃冻存过夜。冻存过夜的裂解液，冰浴解冻后用超声波裂解破碎；4℃ 9000r/min离心20分钟，取上清保存，沉淀用20mL，0.1M Tris-HCL(pH7.4)缓冲液(含50μL 100mM的PSMF)重悬保存。诱导前、诱导后的菌体及裂解后的上清、沉淀经OD600值换算后等量加样进行SDS-PAGE电泳，由BandScan软件分析目的蛋白的表达量，结果显示100mL菌液可以获得约20mg目的蛋白。表达产物经NTA Agarose纯化。
2.9 蛋白的纯化
Ni-NTA柱纯化表达的可溶性蛋白，纯化在4℃层析柜中进行。
（1）在10mL的可溶性蛋白中加入5mL 50%的Ni-NTA琼脂后置于4℃ 轻柔混匀1h；
（2）把可溶性蛋白-Ni-NTA琼脂混合物加入到底部封住的纯化柱中；
（3）移去底部的盖子，收集液体；
（4）用含10mM imidazole的Wash
Buffer洗涤纯化柱四次，每次10mL，同时分别收集每次的洗涤液；
（5）用含250mM imidazole的Elution
Buffer洗涤纯化柱四次，每次2mL，同时分别收集每次的洗脱液；
（6）SDS-PAGE GEL电泳检测蛋白纯化效果，结果显示纯化后条带单一，分子量大小与预测大小一致，如图9。
2.10 重组蛋白的免疫印迹分析
参照《分子克隆实验指南》(第三版) (Sambrook J, Russell
DW，Janssen
K, Argentine J.黄培堂等译，2002，北京：科学出版社)。
（1）纯化的融合蛋白进行SDS-PAGE，电泳结束后切下含有纯化蛋白的凝胶条带；
（2）用蒸馏水淋洗石墨板，然后用滤纸吸干电极板上的液滴。同时，戴手套剪切大小与凝胶大小一致的6张Whatman 3MM滤纸和1张硝酸纤维素滤膜，并用铅笔在滤膜的一角作好标记。把硝酸纤维素膜漂浮于一盘去离子水面上，借助毛细作用使之从下往上湿润后，将之浸没于水中，浸泡5min以上以驱除留于滤膜上的气泡。在一浅托盘中加入少量转移缓冲液，把6张3MM滤纸浸泡于其中；
（3）戴手套按说明安装转移装置：平放底部电极（阳极），石墨一边朝上；在其上放置3张用转移缓冲液浸泡过的3MM滤纸，逐张叠放，精确对齐，然后用一试管作滚筒挤出所有气泡。把硝酸纤维素滤膜放在3MM滤纸上，并精确对齐，且在二者之间不留有气泡；
（4）把含有融合蛋白的凝胶转移到一盘去离子水中略为漂洗一下，然后准确平放于硝酸纤维素滤膜上。把凝胶左下角置于硝酸纤维素滤膜的的标记角上，戴手套排出所有气泡；把最后三张滤纸放在凝胶上方，同样确保各层精确对齐并不留气泡；
（5）将靠上方的电极（阴极）放于夹物层上，石墨一边朝下。连接电源，据凝胶面积0.65mA/cm2接通电流，电转移1.5h。断开电源，卸下转移装置。把凝胶转移至盛有考马斯亮兰染液的托盘中，染色检查蛋白质转移是否完全；
（6）把硝酸纤维素滤膜放入可以加热封的塑料袋中，根据滤膜面积以0.1ml/cm2的量加入封闭液，密闭袋口，平放在平缓摇动的摇床平台上于室温温育2h；
（7）弃封闭液，按滤膜面积以0.1ml/cm2的量加入封闭液和适量的第一抗体（小鼠抗His一抗，购自Sigma公司），密封袋口，将滤膜平放在平缓摇动的摇床平台上，于4℃温育2h，然后废弃封闭液和抗体，用250ml PBS漂洗滤膜3次，每次10min；
（8）将硝酸纤维素滤膜转移至200ml 150mM NaCl、50mM Tris·Cl（pH7.5）溶液和托盘中，于室温平缓摇动温育10min；
（9）将滤膜移至一浅托盘中，按滤膜面积以0.1ml/cm2的量加入无磷酸、无叠氮钠的封闭液；
（10）加入终浓度为1μg/ml的辣根过氧化物酶标记的羊抗小鼠IgG抗体（购自Sigma公司），于室温平缓摇动，温育1h；
（11）把滤膜移至一浅托盘上，内装200ml 150mM NaCl、50mM Tris·Cl（pH7.5）溶液。于室温平缓摇动，温育10min，共重复三次；
（12）在9ml 0.01M Tris·Cl（pH7.6）溶液中溶解6mg二氨基联苯胺（OPD），加入1ml 0.03%CoCl2，用Whatman 1号滤纸过滤底物溶液以去除可能形成的沉淀物，加入10μl 30%H2O2，混匀后立即使用；
把经漂洗的硝酸纤维素滤膜移至一浅托盘上，按滤膜面积加入0.1ml/cm2的底物溶液，于室温轻轻摇动，观察显色反应，当蛋白带的颜色深度达到要求（约2-3min）时，立即用蒸馏水略为漂洗，然后把滤膜转移至装有250ml PBS的另一浅托盘中，拍摄滤膜照片，结果显示清晰条带，说明所获得蛋白为EtMA1-Outside domain重组蛋白。见图9。
3
动物保护性实验
3.1 材料：
球虫卵囊：柔嫩艾美耳球虫(Eimeria tenella)广东株孢子化卵囊，由广东省农业科学院兽医研究所保存，使用前在无球虫雏鸡体内复壮。
雏鸡：岭南黄雏鸡，由广东省农业科学院畜牧研究所提供，饲养于已消毒的专用动物房内；鸡笼和所用器皿均严格消毒，自由采食和饮用纯净水；实验前观察雏鸡有无临床症状及连续2d检查粪便有无球虫卵囊，备用。
饲料：由广东省新南都饲料有限公司定制育雏料，不含任何抗球虫药物。
3.2实验方法：
分组：按实验分组及处理将200只岭南黄雏鸡饲养至10日龄，逐只称重，淘汰弱雏和体重过大者，将剩余雏鸡随机分成5组，每组30只，并适当调整使每组雏鸡总体重大致相等。
处理：
EtMA1-Outside domain重组蛋白的乳化： 取2.9纯化后所得的EtMA1 -Outside domain重组蛋白与弗氏佐剂（FCA）按照1：1的比例混匀；用7号针头注射器反复抽吸至滴于水上5min内不扩散为止。
于14、21日龄分别两次经胸部肌肉注射免疫岭南黄鸡慢大鸡。28日龄分别经口感染5×104个新鲜的E.tenella孢子化卵囊。每天观察并记录鸡群的精神状况、采食量、粪便情况等；对死亡雏鸡称重，剖检，若为柔嫩艾美耳球虫感染引起死亡则病变记分为+4分；所有雏鸡于感染后第7天逐只称重，剖检，进行盲肠病变记分；最后计算各组雏鸡的增重和饲料报酬。具体的实验分组如下表：
药效判定方法及标准抗球虫指数(ACI)按美国默克公司推荐的公式计算：ACI=(相对增重率+存活率)一(卵囊值+病变值)；相对增重率=(实验组增重÷空白对照组增重)×100％；存活率=(各组存活雏鸡数÷各组雏鸡总数)×100％。病变按五级记分：①无卵囊，盲肠正常，为0分；②有卵囊，盲肠黏膜稍增厚，有少量散在出血或少量血样肠内容物，为+1分；③有卵囊，盲肠黏膜增厚，有明显出血或明显血样肠内容物，为+2分；④有卵囊，盲肠黏膜增厚，有大量凝血块或血样肠芯，为+3分；⑤雏鸡因球虫病死亡或有大量卵囊，盲肠外观呈酱油色(或小肠中部有点状坏死灶，黏膜面呈绯红色)，肠管明显肿大，内容物形成明显的血样肠芯，为+4分。组内病变值=每组平均病变记分×20。药效判定标准是ACI<120为无效，120—160为弱效，160—180为中效，180以上为强效。卵囊值由盲肠内容物克卵囊数(OPG)换算而来。
实验结果：
临床症状观察：
接种柔嫩艾美耳球虫孢子化卵囊的红对照组第2天，病鸡出现精神沉郁，呆立，食欲减退；第5天出现明显血便，随后血便逐渐减少，中、低剂量免疫组有部分鸡只拉血便，高剂量免疫组鸡体精神良好，未出现血便。
抗球虫指数：结果见表8。
以上实施例仅为介绍本发明的优选案例，对于本领域技术人员来说，在不背离本发明精神的范围内所进行的任何显而易见的变化和改进，都应被视为本发明的一部分。
<110> 广东省农业科学院兽医研究所
<120> 鸡柔嫩艾美耳球虫MA1基因、载体、重组菌株和蛋白及其应用
<130>
<160> 8
<170> PatentIn version 3.5
<210> 1
<211> 1617
<212> DNA
<213> E.tenella
<400> 1
atggaggctc tacgggaagg cttcggcctg aggcggctgt gctgcatcag
tgccgtggcg 60
gcattttgcc tattcggagc aaagccgagc caagcagcag cttctaacgg
ctcacaggta 120
gcttccaacc catggggaga ttccatgcaa aagttcaaca tttcatacac
tcacggtagc 180
ggggtgtacg tagacctcgg gaacgaaaag acagtgagca acaagaagta
ccgcgagcct 240
gcggggcggt gtcctgtaat gggtaaggag atacggctgc agcagcccac
gacggacagc 300
tctatatggc ctgggaacta cctggagaag gtgcccacaa aggggtcacc
tcgggacacg 360
cggccgctgg ggggtggctt cgccatgtgg gatacgacgc ccgtcaagat
aagccctttg 420
actctgtccg agttggaggc gctggcggag cagcagcggg ccaaaaacga
tcccacctcc 480
cctgcctcgg agaagctggc gaaggtcact gacgggctgg ggctgtgtgc
gtggtgggcg 540
tgggcgacgt acgtgcccaa cgggactacg aacttgaacg acaagtacag
ataccctttt 600
gtctggaacg aagaaaccaa agtgtgcact ttgctaggag tgtccatgca
gctgcttgag 660
ggcgcgggca agtactgctc cgtcggcgac gcctccccag tcctcacttg
gtactgcttc 720
taccccgaaa agaccacccg gccggtttct tacaactctc cttatgtgcg
cgaagaccac 780
gccacggctt gtcccgaaaa agccatttca ggagctcatt tcgggacttg
ggacgggacc 840
acttgccagc gcatgaaggc tgcgaagcaa ataaccgtcc ccaaccccac ggagtgcggc
900
aaagctgtct tcaaagtttc ttcttcagac aaccccacgc agtacacgaa
gccacccacc 960
actgaggcca gcagcagcac cagctccagc aacgcagtgg ccaccatgtg
gcctgtgggc 1020
gccttttcca aggacgaacc ccgcatgcag ggggtgggca ccaactatgc
caactggtac 1080
accaatggta cttgtgagat gtatgacatg gtccccactt gcttcaccct
cgcccctaac 1140
cagttctcgt tcacatccct gggatcggcc gatcccagca ccgccgagct
gcccccctgc 1200
accgaagcca gcgagggctg ggaaatctac ggctactgcg agtgcggaga
cggccactcg 1260
acgccctgga agtgtgagaa cggccagtgg attggcggca gcaacgactg
caactgcagc 1320
agcatccctc ccgtggccct gggcgttagc ttcggcctgc ttgtgccgat
tgccgctctt 1380
atcgcctact tcatctacaa aagaaaaaaa gaaaccagca tcgcaaagaa
ccctaagaag 1440
aaaaagctgc tggacgaaga cgaggagagg gatgaggagt tcctgaaagt
gcaggagaag 1500
aggaagcaca aacaaagtga tttggcccaa gaggccgagc cttcattctg
gggcgaaact 1560
ccccaggacc acacaaacgt tgtggtcgac cacaacgccc acgacgccta
ctactaa 1617
<210> 2
<211> 538
<212> PRT
<213> E.tenella
<400> 2
Met Glu Ala Leu Arg Glu Gly Phe Gly Leu Arg Arg Leu Cys Cys Ile
1
5
10
15
Ser Ala Val Ala Ala Phe Cys Leu Phe Gly Ala Lys Pro Ser Gln Ala
20
25
30
Ala Ala Ser Asn Gly Ser Gln Val Ala Ser Asn Pro Trp Gly Asp Ser
35
40
45
Met Gln Lys Phe Asn Ile Ser Tyr Thr His Gly Ser Gly Val Tyr Val
50
55
60
Asp Leu Gly Asn Glu Lys Thr Val Ser Asn Lys Lys Tyr Arg Glu Pro
65
70
75
80
Ala Gly Arg Cys Pro Val Met Gly Lys Glu Ile Arg Leu Gln Gln Pro
85
90
95
Thr Thr Asp Ser Ser Ile Trp Pro Gly Asn Tyr Leu Glu Lys Val Pro
100
105
110
Thr Lys Gly Ser Pro Arg Asp Thr Arg Pro Leu Gly Gly Gly Phe Ala
115
120
125
Met Trp Asp Thr Thr Pro Val Lys Ile Ser Pro Leu Thr Leu Ser Glu
130
135
140
Leu Glu Ala Leu Ala Glu Gln Gln Arg Ala Lys Asn Asp Pro Thr Ser
145
150
155
160
Pro Ala Ser Glu Lys Leu Ala Lys Val Thr Asp Gly Leu Gly Leu Cys
165
170
175
Ala Trp Trp Ala Trp Ala Thr Tyr Val Pro Asn Gly Thr Thr Asn Leu
180
185
190
Asn Asp Lys Tyr Arg Tyr Pro Phe Val Trp Asn Glu Glu Thr Lys Val
195
200
205
Cys Thr Leu Leu Gly Val Ser Met Gln Leu Leu Glu Gly Ala Gly Lys
210
215
220
Tyr Cys Ser Val Gly Asp Ala Ser Pro Val Leu Thr Trp Tyr Cys Phe
225
230
235 240
Tyr Pro Glu Lys Thr Thr Arg Pro Val Ser Tyr Asn Ser Pro Tyr Val
245
250
255
Arg Glu Asp His Ala Thr Ala Cys Pro Glu Lys Ala Ile Ser Gly Ala
260
265
270
His Phe Gly Thr Trp Asp Gly Thr Thr Cys Gln Arg Met Lys Ala Ala
275
280
285
Lys Gln Ile Thr Val Pro Asn Pro Thr Glu Cys Gly Lys Ala Val Phe
290
295
300
Lys Val Ser Ser Ser Asp Asn Pro Thr Gln Tyr Thr Lys Pro Pro Thr
305
310
315
320
Thr Glu Ala Ser Ser Ser Thr Ser Ser Ser Asn Ala Val Ala Thr Met
325
330
335
Trp Pro Val Gly Ala Phe Ser Lys Asp Glu Pro Arg Met Gln Gly Val
340
345
350
Gly Thr Asn Tyr Ala Asn Trp Tyr Thr Asn Gly Thr Cys Glu Met Tyr
355
360
365
Asp Met Val Pro Thr Cys Phe Thr Leu Ala Pro Asn Gln Phe Ser Phe
370
375
380
Thr Ser Leu Gly Ser Ala Asp Pro Ser Thr Ala Glu Leu Pro Pro Cys
385
390
395
400
Thr Glu Ala Ser Glu Gly Trp Glu Ile Tyr Gly Tyr Cys Glu Cys Gly
405
410
415
Asp Gly His Ser Thr Pro Trp Lys Cys Glu Asn Gly Gln Trp Ile Gly
420
425
430
Gly Ser Asn Asp Cys Asn Cys Ser Ser Ile Pro Pro Val Ala Leu Gly
435
440
445
Val Ser Phe Gly Leu Leu Val Pro Ile Ala Ala Leu Ile Ala Tyr Phe
450
455
460
Ile Tyr Lys Arg Lys Lys Glu Thr Ser Ile Ala Lys Asn Pro Lys Lys
465
470
475
480
Lys Lys Leu Leu Asp Glu Asp Glu Glu Arg Asp Glu Glu Phe Leu Lys
485
490
495
Val Gln Glu Lys Arg Lys His Lys Gln Ser Asp Leu Ala Gln Glu Ala
500
505
510
Glu Pro Ser Phe Trp Gly Glu Thr Pro Gln Asp His Thr Asn Val Val
515
520
525
Val Asp His Asn Ala His Asp Ala Tyr Tyr
530
535
<210> 3
<211> 1227
<212> DNA
<213> E.tenella
<400> 3
tctaacggct cacaggtagc ttccaaccca tggggagatt ccatgcaaaa
gttcaacatt 60
tcatacactc acggtagcgg ggtgtacgta gacctcggga acgaaaagac
agtgagcaac 120
aagaagtacc gcgagcctgc ggggcggtgt cctgtaatgg gtaaggagat
acggctgcag 180
cagcccacga cggacagctc tatatggcct gggaactacc tggagaaggt
gcccacaaag 240
gggtcacctc gggacacgcg gccgctgggg ggtggcttcg ccatgtggga
tacgacgccc 300
gtcaagataa gccctttgac tctgtccgag ttggaggcgc tggcggagca
gcagcgggcc 360
aaaaacgatc ccacctcccc tgcctcggag aagctggcga aggtcactga
cgggctgggg 420
ctgtgtgcgt ggtgggcgtg ggcgacgtac gtgcccaacg ggactacgaa
cttgaacgac 480
aagtacagat acccttttgt ctggaacgaa gaaaccaaag tgtgcacttt
gctaggagtg 540
tccatgcagc tgcttgaggg cgcgggcaag tactgctccg tcggcgacgc
ctccccagtc 600
ctcacttggt actgcttcta ccccgaaaag accacccggc cggtttctta
caactctcct 660
tatgtgcgcg aagaccacgc cacggcttgt cccgaaaaag ccatttcagg
agctcatttc 720
gggacttggg acgggaccac ttgccagcgc atgaaggctg cgaagcaaat
aaccgtcccc 780
aaccccacgg agtgcggcaa agctgtcttc aaagtttctt cttcagacaa ccccacgcag
840
tacacgaagc cacccaccac tgaggccagc agcagcacca gctccagcaa
cgcagtggcc 900
accatgtggc ctgtgggcgc cttttccaag gacgaacccc gcatgcaggg
ggtgggcacc 960
aactatgcca actggtacac caatggtact tgtgagatgt atgacatggt
ccccacttgc 1020
ttcaccctcg cccctaacca gttctcgttc acatccctgg gatcggccga
tcccagcacc 1080
gccgagctgc ccccctgcac cgaagccagc gagggctggg aaatctacgg
ctactgcgag 1140
tgcggagacg gccactcgac gccctggaag tgtgagaacg gccagtggat
tggcggcagc 1200
aacgactgca actgcagcag
catccct
1227
<210> 4
<211> 409
<212> PRT
<213> E.tenella
<400> 4
Ser Asn Gly Ser Gln Val Ala Ser Asn Pro Trp Gly Asp Ser Met Gln
1
5
10
15
Lys Phe Asn Ile Ser Tyr Thr His Gly Ser Gly Val Tyr Val Asp Leu
20
25
30
Gly Asn Glu Lys Thr Val Ser Asn Lys Lys Tyr Arg Glu Pro Ala Gly
35
40
45
Arg Cys Pro Val Met Gly Lys Glu Ile Arg Leu Gln Gln Pro Thr Thr
50
55
60
Asp Ser Ser Ile Trp Pro Gly Asn Tyr Leu Glu Lys Val Pro Thr Lys
65
70
75 80
Gly Ser Pro Arg Asp Thr Arg Pro Leu Gly Gly Gly Phe Ala Met Trp
85
90
95
Asp Thr Thr Pro Val Lys Ile Ser Pro Leu Thr Leu Ser Glu Leu Glu
100
105
110
Ala Leu Ala Glu Gln Gln Arg Ala Lys Asn Asp Pro Thr Ser Pro Ala
115
120
125
Ser Glu Lys Leu Ala Lys Val Thr Asp Gly Leu Gly Leu Cys Ala Trp
130
135
140
Trp Ala Trp Ala Thr Tyr Val Pro Asn Gly Thr Thr Asn Leu Asn Asp
145
150
155
160
Lys Tyr Arg Tyr Pro Phe Val Trp Asn Glu Glu Thr Lys Val Cys Thr
165
170
175
Leu Leu Gly Val Ser Met Gln Leu Leu Glu Gly Ala Gly Lys Tyr Cys
180
185
190
Ser Val Gly Asp Ala Ser Pro Val Leu Thr Trp Tyr Cys Phe Tyr Pro
195
200
205
Glu Lys Thr Thr Arg Pro Val Ser Tyr Asn Ser Pro Tyr Val Arg Glu
210
215
220
Asp His Ala Thr Ala Cys Pro Glu Lys Ala Ile Ser Gly Ala His Phe
225
230
235
240
Gly Thr Trp Asp Gly Thr Thr Cys Gln Arg Met Lys Ala Ala Lys Gln
245
250
255
Ile Thr Val Pro Asn Pro Thr Glu Cys Gly Lys Ala Val Phe Lys Val
260
265
270
Ser Ser Ser Asp Asn Pro Thr Gln Tyr Thr Lys Pro Pro Thr Thr Glu
275
280
285
Ala Ser Ser Ser Thr Ser Ser Ser Asn Ala Val Ala Thr Met Trp Pro
290
295
300
Val Gly Ala Phe Ser Lys Asp Glu Pro Arg Met Gln Gly Val Gly Thr
305
310
315
320
Asn Tyr Ala Asn Trp Tyr Thr Asn Gly Thr Cys Glu Met Tyr Asp Met
325
330
335
Val Pro Thr Cys Phe Thr Leu Ala Pro Asn Gln Phe Ser Phe Thr Ser
340
345
350
Leu Gly Ser Ala Asp Pro Ser Thr Ala Glu Leu Pro Pro Cys Thr Glu
355
360
365
Ala Ser Glu Gly Trp Glu Ile Tyr Gly Tyr Cys Glu Cys Gly Asp Gly
370
375
380
His Ser Thr Pro Trp Lys Cys Glu Asn Gly Gln Trp Ile Gly Gly Ser
385
390
395
400
Asn Asp Cys Asn Cys Ser Ser Ile Pro
405
<210> 5
<211> 24
<212> DNA
<213> 人工序列
<400> 5
atggaggctc tacgggaagg
cttc
24
<210> 6
<211> 25
<212> DNA
<213> 人工序列
<400> 6
ttagtagtag gcgtcgtggg
cgttg
25
<210> 7
<211> 33
<212> DNA
<213> 人工序列
<400> 7
ggcggatcct ctaacggctc acaggtagct
tcc
33
<210> 8
<211> 35
<212> DNA
<213> 人工序列
<400> 8
ggcaagcttt taagggatgc tgctgcagtt
gcagt
35
Claims (10)
1.鸡柔嫩艾美耳球虫MA1蛋白（EtMA1蛋白），其氨基酸序列如SEQ ID NO:2所示。
2.编码权利要求1所述蛋白的基因。
3.根据权利要求2所述的基因，其特征在于，其核苷酸序列如SEQ ID NO：1所示。
4.EtMA1膜外功能域蛋白（EtMA1-Outside domain），其氨基酸序列如SEQ ID NO:4所示。
5.编码权利要求4所述蛋白的基因。
6.根据权利要求5所述的基因，其核苷酸序列如SEQ ID NO:3所示。
7.一种克隆载体，含有权利要求2、3、5或6所述的基因。
8.一种表达载体，含有权利要求2、3、5或6所述的基因。
9.生产EtMA1膜外功能域蛋白的方法，包括将权利要求8所述的表达载体导入宿主细胞中，表达得到EtMA1膜外功能域蛋白。
10.权利要求1或4所述的蛋白在制备抗艾美耳球虫药物中的应用。
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| CN201210279040.5A CN102816221B (zh) | 2012-08-06 | 2012-08-06 | 鸡柔嫩艾美耳球虫ma1基因、载体、重组菌株和蛋白及其应用 |
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| CN201210279040.5A CN102816221B (zh) | 2012-08-06 | 2012-08-06 | 鸡柔嫩艾美耳球虫ma1基因、载体、重组菌株和蛋白及其应用 |
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| Publication Number | Publication Date |
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| CN102816221B true CN102816221B (zh) | 2014-05-21 |
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Families Citing this family (5)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN106046137B (zh) * | 2016-06-03 | 2019-01-29 | 山东农业大学 | 一种鸡柔嫩艾美尔球虫微线蛋-2突变体EtMIC2-1130 |
| CN106046136B (zh) * | 2016-06-03 | 2019-02-22 | 山东农业大学 | 一种鸡柔嫩艾美尔球虫微线蛋-2突变体EtMIC2-2119 |
| CN110606870B (zh) * | 2019-09-18 | 2021-02-19 | 广东省农业科学院动物卫生研究所 | 用于防治柔嫩艾美耳球虫的多肽及含有其的药物组合物 |
| CN110559431B (zh) * | 2019-10-11 | 2023-02-28 | 南京农业大学 | 一种巨型艾美耳球虫纳米亚单位疫苗及其制备方法和应用 |
| CN113201060B (zh) * | 2021-05-06 | 2022-09-20 | 广东省农业科学院动物卫生研究所 | 用于防治布氏艾美耳球虫的重组多肽及疫苗 |
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