鸭肠炎病毒基因组dna的提取及其序列的制作方法

专利名称：：鸭肠炎病毒基因组dna的提取及其序列的制作方法
技术领域：
：本发明涉及鸭肠炎病毒基因组DNA的提取，本发明还涉及鸭肠炎病毒基因组序列，属于生物
技术领域：
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背景技术：
：鸭病毒性肠炎(duckvirusenteritis,DVE)又名鸭瘟(duckplague,DP),是危害鸭、鹅及其他雁形目禽类的一种急性、接触性、败血性传染病，以血管损伤、消化道出血坏死、淋巴器官受损和实质ti器官退形性变化为主要特征。该病在我国流行已近40年，但这期间的流行特点发生了一定变化。根据1963年以前的报道，最易感的鸭群发病率和死亡率均在90%以上，但在随后的20多年中则有明显的下降。自2000年以来，DVE的流行特点又有了新的变化，表现在病毒主要侵害l月龄以下的雏鸭，成年鸭很少发病，常规弱毒疫苗免疫效果不佳，鹅发病的比例明显上升，国内一些学者称之为"新型鸭瘟"。目前人们已经知道，该病的病原为鸭肠炎病毒(duckenteritisvirus,DEV)，与常见的鸡喉气管炎病毒、马立克氏病毒、火鸡疱疹病毒、猪伪狂犬病毒等同属a-疱疹病毒亚科。该亚科目前共分4个属，分别是单纯疱疹病毒属(57历Ae;^&M)，以人疱疹病毒l型(HSV-1)为代表，还包括人疱疹病毒2型、牛疱疹病毒2型和猕猴疱疹病毒1型等；水痘病毒属(Za/7'ce/7ow'nAS),以人疱疹病毒3型为代表，还包括牛疱疹病毒1型、马疱疹病毒1型、马疱疹病毒4型和猪疱疹病毒1型等；类鸡马立克病病毒属(胞rWsA'seae-乃'Aew'/Y^es)，主要包括鸡马立克氏病病毒(MDV)1型、2型和火鸡疱疹病毒(HVT);类鸡传染性喉气管炎病毒属(/"ow'r^)，目前仅有鸡传染性喉气管炎病毒(ILTV)一个成员。疱疹病毒的分类除了根据其病原特征和致病特性外，基因组结构也是主要依据之一，例如，MDV曾因能够引起淋巴细胞肿瘤而被分到Y-疱疹病毒亚科，后来根据其基因组结构和序列才将它归于a-疱疹病毒亚科。DEV由于复制周期短，能在三叉神经结等处形成潜伏感染等特征而被分到a-疱疹病毒亚科，但目前人们还不知道它的具体分类地位，即该将它归于哪个属，或者为它新建一个属，其中的主要原因就是DEV的全基因组序列尚不为人们所知。己知DEV为双股双链DNA病毒，无血凝特性和血细胞吸附作用，除有芯髓、衣壳、囊膜等常见结构外，在衣壳和囊膜间还有疱疹病毒所特有的皮层成分。DEV呈球形，芯髓直径约60nm,衣壳直径约90nm，成熟病毒直径150—380nm,在CsCI中浮密度为1.272g/ml，由限制性内切酶图谱测出的核酸平均分子质量为93.93X106D，但是目前没有DEV的全基因组序列的相关报道。
发明内容本发明目的是通过提取DEV基因组DNA，构建基因组文库，通过大规模测序，获得DEV全基因组核苷酸序列。从而为DEV的分类提供依据，为其生物学特性的深入研究奠定基础，也为DVE的临床防治开辟新的途径。本发明的目的通过以下技术方案来实现-鸭肠炎病毒的增殖将鸭肠炎病毒商品疫苗株(购自中牧实业股份有限公司)接种鸡胚成纤维细胞(CEF),待出现完全病变以后，收集病毒液样品，一8(TC保存备用。鸭肠炎病毒基因组DNA的提取将收集的病毒液样品高速离心，使细胞和病毒粒子同时沉淀下来，然后用去离子水悬浮，冻融3次以后高速离心，沉淀悬浮于预冷的高渗缓冲液以裂解细胞，使细胞DNA充分释放；再次高速离心后，沉淀悬浮于含微球菌核酸酶的缓冲液，37"作用30min,以消化分解细胞DNA;加入EDTA抑制核酸酶活性，加入消化缓冲液，5(TC作用过夜；使用常规酚/氯仿抽提病毒基因组DNA，取水相，加入醋酸铵和无水乙醇沉淀病毒DNA，洗涤晾干后溶解于适量TE缓冲液，测定DNA浓度后，一2(TC保存备用。鸭肠炎病毒基因组DNA样品的PCR鉴定将上述步骤所得病毒基因组DNA样品经PCR方法进行鉴定，同时设立阴性、阳性对照，PCR产物经in/。琼脂糖凝胶电泳，拍照。鸭肠炎病毒基因组文库的构建和序列测定将病毒基因组样品用DNA片段化仪随机剪切，使产生的主要片段在2.02.5kb之间，回收后与pUC19载体连接，转化感受态大肠杆菌，涂布选择性培养基，挑取阳性克隆进行序列测定。使用通用引物M13与MegaBaselOOO测序仪上迸行序列测定，用Phred软件从测序仪生成的色谱图中获取碱基序列，用Phrap软件和Consed软件将各测序片段进行拼接和编辑，最后剩余的Gap则用PCR方法补齐。获得的基因组序列用Lasergene和Blast等生物信息学软件进行0RF确定和同源性分析。测序获得DEV商品疫苗株基因组全长为156512bp，G+C含量为44.85%。呈现典型的疱疹病毒基因组结构基因组由长独特区(UniqueLong,UL)和短独特区(UniqueShort,US)组成。US区两端为一对反向重复序列，分别称为US内部重复序列(internalUSr印eats，IRS)和US末端重复序列(terminalUSr印eats，TRS)，与水痘病毒属相一致。基因组至少编码78个开放阅读框(ORF)，基因编码的氨基酸序列与a-亚科疱疹病毒成员有同源性，其中5个基因仅与目前所知的禽疱疹病毒有同源性，该序列为SEQN0.1,基因组结构示意图在附图l中表示出。本发明与现有技术成果相比具有以下创新点-1.建立了一种用于鸭肠炎病毒基因组DNA的提取方法。本方法不需要纯化病毒，操作步骤相对简单，最大限度地消除了细胞DNA的污染，获得了非常纯净的病毒DNA，保证了下游工作的顺利进行。实验证明该方法对于获得纯净的病毒基因组DNA非常有效，可借鉴到其他有囊膜DNA病毒的基因组提取研究中。2.国内外首次完成了鸭肠炎病毒基因组序列的测定，推测基因组至少编码78个0RFs，绝大多数基因编码的氨基酸序列与其他a-亚科疱疹病毒具有较高同源性，基因组结构与水痘病毒属成员一致，结合病毒复制周期短，能够在三叉神经结形成潜伏感染的生物学特性，支持了将DEV归于水痘病毒属的观点。图l是鸭肠炎病毒基因组结构示意图。图中箭头代表编码基因的开发阅读框(ORF)，目代表长独特区(UL)，,代表短独特区(US),■和,代表短重复区(RS);每行最上端数字表示序列中核苷酸的长度，靠近箭头的数字表示编码基因的序号。图2是鸭肠炎病毒基因组DNA的PCR鉴定图片。其中M是标准分子量DL2000，1、2为基因组样品，3为阳性对照样品，4为阴性对照样品。具体实施例方式(—)鸭肠炎病毒的增殖和基因组DNA的提取1.鸭肠炎病毒的增殖取孵化至9日龄SPF鸡胚，用碘酒消毒蛋壳后，无菌取出胚体，放于培养皿中，除去头、四肢和内脏。胚体用Hanks液洗涤3次，转移至容积适中的小烧杯中，剪成12rnrn小块，洗涤3次后，加入适量胰蛋白酶，室温消化2030min。加入适量细胞生长液，用大口吸管反复吹打，使细胞分散，形成悬液，转入离心管，以lOOOr/min离心10min,弃上清，将细胞沉淀用Hanks液洗涤2次。加入细胞生长液，用吸管吹打制成细胞悬液，计数，稀释成10"个/ml，分装细胞培养瓶或细胞培养板，于37°C、5%0)2培养箱中培养。待细胞长成单层后，即可用于接种病毒或继代培养。将DEV疫苗株(购自中牧实业股份有限公司)接种CEF细胞，同时设立不接毒正常细胞对照，每日观察细胞病变，待出现完全病变后，收获病毒液，一80'C保存备用。2.鸭肠炎病毒基因组DNA的提取(1)将收集的病毒样品30,000Xg离心30min，使细胞和游离病毒粒子沉淀。(2)将步骤(1)所得沉淀用灭菌蒸馏水悬浮，—85。C23。C冻融3次，30，000Xg离心30min，所得沉淀重悬于预冷的Cellpermeabilizationbuffer,置冰浴10min,30，000Xg离心30min。弃上清，取沉淀，重复本歩骤一次。(3)重复步骤(2)离心后，弃上清，将沉淀悬浮于500"1Cellnucleibuffer,再加入500p1含600U微球菌核酸醜的2xNucleasebuffer,混合均匀，37。C作用30min。(4)加入60nl0.2MEDTA终止反应，加入4mlDigestionbuffer,用剪去末端的Tip头轻柔混匀，避免对病毒DNA的剪切作用，50'C过夜。(5)加入等体积酚/氯仿，轻轻混匀，8,000Xg离心5min，水相再用氯仿抽提1次，8,000Xg离心5min。(6)室温条件下，取上层水相，加入0.5倍体积的7.5M醋酸铵和2倍体积无水乙醇，轻轻混匀，16，000Xg离心5min。(7)步骤(6)所得DNA沉淀用70X乙醇洗涤2次，晾干，溶解于100ulTE，测定和计算DNA浓度，标记作样品2(Sa卿le2)，同时用等量未接种病毒的正常细胞以相同程序提取DNA作为对照，标记为对照(Contro12)。一20'C保存备用。(二)鸭肠炎病毒基因组DNA的PCR鉴定用PCR方法对提取的DEV基因组DNA进行鉴定，引物序列为上游引物5'-TCCTGGAACAATCACAAC-3，下游引物5，-TCGCCTGCCAACTTAT-3'组分如下上游引物下游引物dNTP(2.5mmo1/1)MgCU25腦1/1)TaqDNA聚合酶IOX缓冲液模板DNAddH20l.Ou1l.Ou14.0u13.On10.5u15.On110.Ongupto50y1混合物用30.Oil1石蜡油覆盖，置于PCR仪内进行扩增反应，程序为95°C5min、94°Clmin、50°C30s、72°Clmin的反应循环，共30个循环，最后72。C延伸10min，4'C结束反应。同时以正常细胞提取的DNA作为对照。PCR产物经1X琼脂糖凝胶电泳，拍照，照片见附图2。(三)鸭肠炎病毒基因组文库的构建和序列测定1.基因组DNA片段化将基因组DNA样品用TE缓冲液(pH8.0)稀释成40ng/ul，以100yl体积上样，调整片段化仪Hydroshear的速度代码和循环次数两个参数对样品进行随机剪切，使片段化后产生的DNA主要片段在2.02.5kb之间。片段化结束后，将样品于1%琼脂糖电泳，用试剂盒回收2.02.5kb之间的片段，溶于适量灭菌去离子水中。2.DNA片段末端的补平与分离在0.2ml灭菌离心管中加入以下组分DNA片段化样品50.0ul10Xbuffer10.0uldNTP(2.5mMeach)10.On10.1%BAS10.0n1ddH20upto92.0n1以上各组分混匀后，先在7(TC保温5min,然后移入37'C恒温槽，加入8ulT4DNA聚合酶，轻柔混匀，继续反应30min。反应结束后，加入等体积酚/氯仿抽提，12000r/min离心20min;上层水相转移到另一灭菌离心管中，加入等体积氯仿混匀，12000r/min离心20min;上层水相转移到另一灭菌离心管中，加入1/10体积3MNaAC和2.5倍体积预冷无水乙醇，12000r/min离心20min;倾弃液体，用70%乙醇洗涤DNA沉淀，晾干，灭菌去离子水溶解，电泳，切胶回收2.0kb2.5kbDNA片段。3.DNA片段与pUC19载体连接常规方法制备pUC19质粒，5^I单酶切后，用小牛肠道碱性磷酸酶去除载体末端5'-磷酸基团，与补平后的基因组DNA样品连接-基因组DNA样品lO.OiUpUC19载体2.OullOXLigasebuffer2.On1T4DANLigase2,0u1ddH20upto20u1以上组分混合均匀后，16'C连接过夜。半透膜脱盐，溶于灭菌去离子水中，使DNA浓度为10ug/ml。4.感受态大肠杆菌制备(1)取活化大肠杆菌DH5a接种LB培养基，于37。C摇振培养至00。达到0.4，迅速将培养物置于冰浴中30min;(2)4'C2500r/min离心15min，弃去培养液，以预冷灭菌去离子水悬浮；(3)4。C2500r/min离心20min,弃上清，用10%甘油重新悬浮;重复本步骤一次;(4)4'C2500r/min离心20min,小心倒掉上清，吸去管壁残留液体，加预冷GYT培养液重新悬浮，测定Ol值，稀释至细菌浓度为3X10"'个/ml。5.重组质粒的电击转化(1)取40ul新鲜制备的感受态DH5a,加入2n1脱盐连接产物，置冰上60s;(2)调节电击仪，使电脉冲为25uF，电压为2.5kV，电阻200Q;(3)上述混合物加至预冷电击杯中，轻弹杯体使混合物位于电击杯底部，将电击杯放入电击仪，启动电脉冲；(4)取出样品，室温下加入lmlSOC培养液，37。C轻柔震荡培养lh;(5)转化产物菌液涂布X-Gal-IPTG-A叩平板，37'C培养过夜，挑取白斑进行培养，提取质粒用于序列测定。6.序列测定和拼接用通用引物M13于megabase1000测序仪上进行序列测定。用Phred软件从测序仪生成的色谱图中获取碱基序列，用Phrap和Consed软件将随机片段进行拼接和编辑，最后剩余的Gap则用PCR方法进行补齐。7.ORF的确定和基因同源性分析全基因组序列用Editseq(Lasergene软件包version6，DNASTAR公司)软件以正反方向6种阅读框方式推导为氨基酸序列，所有编码大于60个氨基酸残基的基因均通过Blast(NationalCenterforBiotechnologyInformation)进行同源性比对，与已知的疱疹病毒有同源性的基因即被认为是DEV的功能基因，并参照HSV-l相应基因名称命名。若是HSV-l没有的基因，则参照同源性最髙的已知基因命名，没有同源性的基因则自行命名。为了验证以上获得的结果，全基因组序列提交在线工具GeneMarks(GeorgiaInstituteofTechnology[http://opal.biology.gatech.edu/GeneMark/genemarks.cgi])进行ORF搜索，并将两种方法获得的结果相互印证。利用blastp、PSI-和PHI-BLAST将推导出的DEV基因氨基酸序列与GenBank中已发表的HSV-1、MDV和PRV的相应基因进行同源性比较。除以上三种病毒以外，通过对比还找出DEV特定基因同源性最高的基因所属的病毒。核苷酸序列表〈110〉山东省农业科学院家禽研究所〈120〉鸭肠炎病毒基因组DNA的提取及其序列<歸1〈210〉1<211>156512〈212〉画<213>鸭肠炎病毒(duckenteritisvirus,DEV)〈220〉<221>gene<222>(l)...(156512)<400>1tccaacgcctctcccggccgcggccgcgcatttgcctgacgtgccttc犯60tgcactgctatggaagggcgttgggggstgggccactcacattc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aagU16320Cgagg33C3t16380ttccggattt16440aacaagccct16500ctttggaagc16560ggaccctata16620函Otcaatgctat16740tatttttcgt16800cggtaacttt16860tggtgctgta16920a肪tcacg幼169803組cggagg17040ggtaagagac17100catggatttg17160gagccgcgta17220tttggcgcgt17280gaatsicatag17340cattctg卿17400aaacgggcgc17460agacaccgcc17520tcatgtaaat17580<formula>formulaseeoriginaldocumentpage17</formula>tcttcattgctgtgttatggtcatcttcaaagtggcat已g20340tgg犯csaatggagtgtc已tctgccatcgttgcagcggttgaacttgttacaatggttca20400ccatcattgtcaateitctggtt犯t3tgstt3tggC鄉3tscatttgttggstt犯ggg20460cggcgcg犯ggatactgttttacaatctgcattgcgttcccaaggacgtttttgccactt20520tattggcagactagcccctacgatgacagtacattcgtggtctg犯atgga犯aggg犯c20580gcg鄉gtggttca犯atggctcacgctcgctc犯ttattactcatggta20640acactatctaaccacagtagggtcagttgatgacccgtttgc卿tacggtggstaactc20700cgacg已cgatctcttaacgatggatatattacacgatatggatcatccat20760cgtacggtcatcgagagcgagtggscctggtgggccttatCgtgg犯t3CtggtC333C￡l208203tcaaaga>cggat3gtg卿tccatatttacatstggattcgcccaaatc20880gcctctgtctatatcttctttt犯已tc犯gggccaggcacaggcgcacgtcttcgtcatc20940atctact犯g3C33CgCgC3atgacccctc3t￡lCtggtttgc鄉cggcg21000cagcgttgcatcgaatggtcatggeicgtat21060attaaacacatc犯cactcgscggaccactttctggtccgggaactagcgctgatgg卿211203tgggCC鄉ggg認gtcgtgtgcttcat21180taccgctacccctactcggaa犯cgct犯ccattgcttcg犯tacgagac21240t^t^Cg3Caccgcaacgggtcctaaatgcgagtcaccagacgatgttt21300agaacgcgcttcatctaactacagttatg3gaatg犯actaagtttgcaaccgttgcttc21360tgatttgggaaggaatcatcgttcaatgtctagaacgagttgcgcttccccaaacgtgtt21420gaattcttaccaccacggatcggcggctcgcctgtacaattcggggacga21480tg咖cggtgcaaitgtc肪gttggag卿cgacggcggcgttcagaccgcgggcatattg21540cctcggtaaacgtgatcgagagcgcg犯ctagctgcagtgcagcgagttatgaaitagact21600gtcatttgggaaat犯ttgatggtttggcgtttggtttatgttgtatcct216603Cttttt8L犯tctggtttatgttgtatccttctaacaaat21720tggtaat犯ttgatttagttaatctgattcstccaaatct21780cgagttttgtagtaattttatta卿tgggtag犯acEicgcatac犯at3acaagtctta21840犯gacgcgccggacatsgcga^C3ttCCELgccacagagccaagtggtct21900tggtttctgacacacagtcgtcgcagtttgtcatctttggcgccagctgc21960atcgattcttgtccgttcaaattcgtccaa3ttgggggg3ggctttctcc22020犯ttgacgtattgtttgcatattcgccgatggctgctagtsatgtttttgcattagcaac22080ggtELELttggttcggcatcaacggccgcgcatagtcctatgacgtcctttgcaatagtatt22140朋gcctgscggC3t3C3CgCatactgtgtctattcctatccsaccgtggtgacaagtgct22200cgtatcccacgctgtagtcggt3tCgC8犯tacgccgacaccaatccgeic22260aa11aatataCCC3Cg3C犯taccgattgtgELttaggcgtgctgttttagC3C3CCC犯C22320tccatggcggttggccgcgstctttcctctagctatttcg22380ccgcgtgttaattggctcgcattccatccggtttgcggtacagcactttc22440gcgcggttcggagactgaaaactcgtctct3g犯tgcacEiacaagttcttcggccgacat22500gggcgccatcattgttagggtgatccgcccgcaatacacggattagccagatggattgat22560tgacgggc犯gtgtccagttatgtatcctaaacataaggttt犯tcgatgagttgtgtga22620tg犯ttagcgccacaacggtcggt犯cgtgtcattctagttatttagtaatattatgatt22680taatgcgtcagattatcttaataatggggttgg犯犯tg3catcactgeitgaatggcctc22740tcagt犯tgttgccccattatgtatatttccaccggacctgggC3gtgtt22800gacatgtc犯agca鄉gsgaacctatcttttacgagagtttaggttatc22860tcgcctcgcttcgatatgccca犯gcgtacagcattcattgttctgcttgcgaagaatgt22920atttcgccgattggctgtgatgcggatttaatttacggacggatgcttagagaatttgta22980aagcgacaatggccggtcaattggttcgggcttcgtgtgtaacacgtaat23040tcgatcggcgtcagcggattggtggcgcatttttcgctag卿gg犯cgctgtacc犯tg23100tttactacatcacgcgttccatggcaactcatcaagaa犯atctatcatcgcgtattcgt23160ttagctgcatcgg犯g犯aB3cgtaBgttacgtactgtccttggggggsc23220aattattcaatgacacccgtaccgcttggcggC3t鄉CgcggcgttggaC3tacttcga23280cgcggtccatcagcgcattttacgataca^gtagattgctgtcctatcagtccgtatatc23340ctacaattggtcgtatttaatggccatatgactgcgctgttgcatgtggatactatgcac23400gtcagcgataa3t3tggc3ttggatggctattgaccg鄉gcggaatcttggcscattat23460tgcgcaaagcgacttggcgtatttgctgatcgtttagcattgctgataggacttgcaatc23520gcgtctagtaaacgtg犯tgtatgcgcacaatcaaagcgacgatgaatactactatggtg23580gtatacccaa3ggttgtctttaggcgttttgcccgttcaataaacgattgg3g犯caaac23640tatgtggctatgattccttttgttcgg3tsctatatg犯gctttggactg23700aacgtgccgaacaatctgtatagacactgccccatcatttttg犯gttgtgctatctata23760aactcB^ttaatttgaatgaCtgt3g犯Ctgtg犯gcgctcgaactacactaccattgtt23820agcgtcccgatgct3tttaaaacgtcgtttatttatc犯aagctttatta23880aacattttata1:tEi犯cc3gatattgtctgctttatctcgcgaataatag239403aaca犯cggccataattaatattatcccgCCggttELEL犯ttccagccgtt犯犯ctgta24000actatgatcatgatcataggcttcccctcgg犯t鄉gggaagggccgca24060tcttcaaatataggcagttcctccggatagccttctacgcgacatgtgt3ctgtgtcacc24120gtgccagtgtattc犯gcgggtaacgag3tct犯tattgacagttccgggatgatcgatg24180cattggcctac犯cttccaccattccgtccttcc已已atgttttcttttagctctcttcca24240accatccatttaattcc犯ccgcagctgtgtcggcgacgcatcttgcattacatactgca24300tggcccgattcgaatgttaagataacgtttggctteig犯ta犯c犯ccgg犯ctgctg犯24360gcaitt3犯tgttcgcgtctccccttcgccattCC￡ltat￡lgctttacatgtgstctctggg24420ggcatag犯ttccgag犯gaagcg犯ccgatgtgtcUct24480tcactttttgcggtag犯taagtccgcactgagtttataggC犯3ttggttg犯tcatcs24540actacccacgtcatacggcctgagccgggggggaa犯犯ttttctaatatacacgttgcc24600cgatatcgttggCCELt犯ELCcatcggatgagcaatgac已g卿tgccggatggtatcttt24660at犯ttttcagtgttattgctccgattgcttctgtccgtctttttgtcttattagtatt已24720ctttcaacaa犯ggcgt卿C犯C3CCC3ttcatac犯tccctcgtctagagcagtcggc24780ttaaggatagtc犯tggaaagct已gagtcttcggtcggttatgtttttca24840tccagtccatttcgcctaggccactatccgcattacttggcatcgcagta24900tagatgattct鄉ttgagcttgcttag犯ccgacgggatcat3ctgcsttccgttccat24960tcgactggaggactgcttgtgttggtttgtaacgaccatt25020gtccaattctt鄉Elttgg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tcggccc犯gcccagaatttcattttctgacatactccgtccttcaccatcagcactgg已犯tcacscgcttcgtctattat犯g3CggCgg犯ttscgttttggtggagacacctttatgacattgaccctctcaatagcgcatactcgttctccaatgtttataccgccacatctcagcataagcgacaaattttctgccgataacg犯tttcg,catgacggagccacag犯ccatatatcag幼ggacasca卿ag犯gacgstgtttatttctggagattttctttcataactgaatcaagttcttcgcgcc￡ttaaaascgsttgtggcagcgcact74340gacgtgttggcatcggttcgtctcttggat74400gcgttctt犯gtgcggctgtcgtcgctgcg74460cgactcttgcgaacgcttaatccggcgcgc74520cgtccgcgtatacctagacgggccgtatgg74580gtcggaggatacccttgcagccactcttta74640tcattttg犯gatgtaataa74700C3t3gC33C3actgacgcaaaagcaattac74760atacagttcctttcatacatatgcatcgac74820attacgctcataattgatcg74880agcagctcgttttgtagtgggggatatgag74940aattcctcgcgaaccccatggtgg犯atat75000tcttgaacgattgagagctcgccaacgtcc75060cgtattacataatatatataagatgtttgt75120aat组gtgggactgtgata75180cgatag犯ag犯aatgttagaagcgcgtga75240gccaagtccatcaacgactctcttgtcact75300atttagsctgcgtaaagtcc已tg犯tgg犯75360tttg犯tgcatattttatggacat犯gtgg75420eicg卿ggc3actaacgcgatggactttac75480gg犯c犯gctgttaaccaatac犯taccga75540ttaaataaagctctgatgggcggttctgeLa75600ccacatttgtEL犯tgC3CgCcttcgctact75660agtatatgttcax;gcgag犯tataatgcgc75720ctgggacctcaccgcggagctagttttctc75780ctaaccggccaactttag犯tgtcgcagct75840tgaggttgta已gtgcggcgtggataggtgt75900tcatcacgttatatcgcc3gatgttcttcg75960tgga>ccaactgggcgtagca76020tccaccgatctgagt犯tct76080cttcgttaccg犯cgaagtt76140鄉gggca^aaattcagatcaattgacta76200ttttgcacgttctctagcatgttctgc犯c76260gaacacgtcgacgatataaa76320tggctc犯atcctcc犯aggacattgctaa76380cg犯gcattttttgatcccaaaacgctcct76440ggatccactggcctcagtttggccagt卿76500aactgatcgcgcggaagctgtggccaatat76560gatggccagttcgcgcccaattg犯attct76620gatatggcctcgtttcgatacagtattcga76680atattggc幼gtttc犯犯a76740卿acttggagccattacagc卿agtgtt76800tctac犯cttc城t犯tgtteitcc犯tgc76860acg犯tcgttacacgtgttgctttgattgc76920c犯gtttacaagtctggaggactacatagc76980aacggacttaccctgatacaaactacacggcgaaccacta3cgcctaga>ccatcgtaatgtgagtccacgactaacatcaattcattagctgtcacagca组tcctgatcataactgcggtcgccagatttccagacatccctttatatagtac朋cttgcgttatacagcgcg犯tttgggtgc郷a3caa3atia11tgttgccgtaaccaatttataacc犯犯ggcattgcctcaacgcgcc肌gacatcgcgtcttgccgtcttctatacgttgtttcaaataaactgtttgacg犯aaagtctattccgagtcagccgcctttgtcccatggacgccaattctcctgagagact鄉tttg犯cttgcaca犯g卿tgtggtgatgtcgctcttattacatcttttttatgcgtcatagaattctgcactagctgcacatggctcggcagctatgtgtactggttactgatcttgcgatacgaggttacgccaacctaaataataaaacact犯tggccttgttcctagacgataacatttaCCgCg3CC3gtccaccggcgatggccgggatcattactatga^gtaitgtcacttgtcttttgaactgttaatgttcctttaatttcggttaagtgtttgcgttcaaatstEitaa犯cacattgtcaaaaca組tgtagcgcattaaatcataattgaatacgtttctgcagcgcgttgatatttgtcgtacgcgcttagaacac犯tatsataaccaacgagctctagcgtacatccccaattaccggcstgaactgac8ggt组gtcggccatgtccatgtttgttutccttgcaacgaactggatttggt卿cgataaatgtactataagctctcg3tgg33Cgtccgtccctaggcggcaatgatcgtcgcggctgcgtgctgatccttcccgattatgttgcttcggtgt犯c犯catgccggcggttttgtataaattcttctatttcctaataccgggcacaataaattattcattg已犯ctgatcccgtataccctaatacccttttgaattt犯tEitatttagtcgtatgactggtgtttcatccatca犯tatagcgatctactagcacttacgcatateictg犯cc3aatagttcccatcctaca犯ccatattcccgtagttttgcCagtC￡LtC￡lttgctttaccagtttcttctccgctgtgtgtaacattatcccatacatcgtgcgccttgggaac3gcca^catggtg鄉ggcctctatatttcagtatg3gg^t3C3ttctaaaatcagagg3cteit3attatttgacgccattgaccatatcgcattgtgttcgtcc卿gtgtctgttctgtatatcactccatttcggaacaatagtatatcgcgta犯ctgstgatgattttattaaacagtattattgccctta^ca^ctatELC3t犯Ctggaccggcgtttatcacatttacgagttcccaaatttctgccctgggccgtccgaatcatccgga卿tgcaatgtata^tcaatgctctatgcgcaaacttcaatgttgcccgggttcgsgatacggtcca8城att犯ggtgcgccgcgttgtgtttaaatactgttccgtactcccct39aaaactgcatgagaaggctaaagagtgatttatggtctggataagtaactc已cacgttgtgctcgUttg犯tgta^ccctggacctatsaactatacggaca犯gatgg犯cgctggstgtgg卿aacccaacggcactaggcgtagtctagttccatactgtgggtatagccgatcttcaatcctggttaacatac由gcggttgtgt3已tCt已3aactcaatgcataattttgccgacccatcaatatatta^gatttt犯gcggttcatca犯taaaacatetagattattgaagtcc鄉卿ttcatctgttCCag￡Ltt￡ltgtctgggcggtgattgggt犯ttgtgcgctaaccaaaccaacEica^gcgggsacgtgaatgtcgttccctccg犯ggagcgcacgtcg犯cgt犯ctggagaccagtatattatggc犯gcgccggcgataatatattccgatttttgeaactacttctgettgc犯g犯attttctccatgtgcttagttcgaggaatacaggaatc鄉aggttaaatctcttttattatatgtt犯caaatattgegtcgtgtaatcattggecatgeacatcatttggttgecttegaggggtatggctt3g犯3tgggt犯tttcattataegtccataaccaatttatctttactgcgatcaattaattgacgagctcagtgcgtagcg卿cgcgteggtagecagcaacgcgcac犯gattcgctctggs犯cgcaatttggatga犯ctg犯ctetattgeatgt8Ct犯t3tagegatcgegEltgt3tttt3Utcg卿8tccagagccgcttgccagtaagetatgegat770407710077160772207728077340774007746077520775807764077700777607782077880779407800078060781207818078240783007836078420784807854078600786607872078780788407890078960790207908079140792007926079320793807944079500795607962079680caaatagtccatcgaccatagcggc犯gtgactgcggcaatgatgtaagacgcgcctgca79740tgeigcgcaaatgcattcctatttggUcgtagaacttgcattagt犯t犯79800ttggtacatcatat犯tttgagttcgaa幼tatatgacccgttgataata79860tcaggcctgacgatatttcgtatactcagcgatacactggtcgcgaactt79920caacctcaagCgCCCCELC3Cagatcacgctcagtacttccatcaccgctatttgtcccgc79980ag卿tcgcgttcgctacttccag犯gtatgcccgagt犯atccaaacta80040cctcaccatccatttcaeLcccccgccgtccaagttcacttcttacat犯t80100ttgacagtaccaaatctccggaggttatccctcgtaatgctataccaaacttggccaatt80160cggcttgtgtgtcactgtcattttccggtgggcgtggtsctgataatatsC3gCC3CC3C802203tgCg犯gt3tgctctcatgccgtcgcgtgtgat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cgcttcgtctgcgttgtctcttagaggcgtc犯t卿gacggga110400tg卿cggcgccggttctgaaccgcgacgtctgcgctcgc犯ctttcgaaccctacccatl薩0gatgctttttCCC犯CC3CCaaatcscctgtccgctgacgtctgctggcg110520gacttgtgtagtttgcgtgcsgcttgaggcatggcagacaca犯acgcctaaagtcatc3110580ctagatgctaatgccgcgcgcccgaacstagttcgcgatcacttcaggactgaggcgtac110640tgacctctattacgttgtcgtacggcagcaaactcgtatal翻0catagactggaactacgggttgggtacggccgtctgcagaacgcgtatagtacttcctcg110760gcttcctagcgtctgtaactgatcgcgtatatacttcgcgtgtactctgc110820aaacatctgcaggcttttattgccgtggccaccctttactgtttgtcttg110880acttactccgcctgttagaggatgacg犯aEtcgacg犯gtcttagtacatgcteittgtaal讓0atttggctaaag犯tgtcgc犯gtccttgcgaatggtatcggtg卿鄉cgtctcccga111000gctcgtc犯tcgacg犯accat犯acat3gtac已tagttttgatcacatgll翻ttgtttcattCgggg3Cgt3ccgcaccgtcctcgttcgtcgtaagtacccgcaggtgccc111120cattctgtatattaacaitcag已tgttagcacttccgcgcctggaccgcggtcgcgttggc111180aatcaaac11acttctactctgttgcccgttcgccgctcccatagaccac3ceictgcata111240atgttcccctttg3tcgatgcgtgtgcaatcax;gttgcgttgtactacattcccaacccc111300犯tcacc犯gtaccgttataatagatggtactggggtgtccatagtcgaggactaagtcc111360aaccagacsggtgagtcgcgtgcccttttt3t犯aggtgtaagttacatccgtaacgtcg111420tttatactgttCC3C犯gga￡tgttgCC￡lgtcaaccggctca6LCgccacgaccttccaggt111480attcattggccttttt犯aa卿tgaatggctttctagacagcggtgatg111540gatggctatatttc犯犯cacaatgcttgcgacaattcggtttgaacgcttcctgagcat111600gcacgccccacagcat犯acactaatccttccttattgtaC3tgt犯3CC8ttgtaatac111660agatttgactatttttgacc犯cc3acgtactacatgcgataccgctggcgcaccacaag111720gcctgcgtcgtttcggcgcctggg已cggagacgggagcctatcgtcggccatggtgtacg111780ggttcacttttgtcatgtcttcggccatatctcgagtctggatcgaatgcataacggatg111840ctggcctgagggtctttg犯gcgatgcatattcgcgtttccgtggc鄉ttctgtcattt111900gggtctccctcgttgacacagccagtacctacaggcgcagtacta犯gctcggcttcccg111960ccccacgctgaattt犯ggctttctg犯犯gccttgacccccacgtcatcacctattgcal匿0acttcagagtccagc3犯aacccagaaatcgcaaggttagggttgaCCCEltaatgcccgc11208051<formula>formulaseeoriginaldocumentpage52</formula>caggc3atttcagcccatctgggtctcttgcatg犯gttttggsgcctcccgcaacgcca114840ttcgtaacaacga^tcgacgagtgtC3g犯Cctggccaactagtctt犯gtttactttaga114900ag犯ccttctgt犯cttscac己cggacccgcctcgagcattatggcagactaaacaagtt114960accc犯tattgt犯C3gg朋gagtagcctc3tttg犯taa115020cEtaccaatgtgccga犯cggcgccattggg115080gccagcggagCCCCCtC3CCg3tCCC3&3g115140gcaccgctctttcatcgttcg3犯gtcattt3tsgtag3t115200cgcggcctatttsgc犯tgctat犯attattattatscttattggttctt115260ttcataaaattgttattcat6LccatattggCtt3犯Cgttttgttaccga115320gcgcccataa犯犯g犯犯tgctcat城ttacgtgtctccgccgacaccgggggcac3t115380catcttggtctctttcacact已gctggactagctccgccgctactgttgg115440ttgtcatttcgtcggccctgttgatc犯tacccgctaattg33C3tt3g3115500CCC3Cga3CCga^aaagcggcgggggtcaaatcttctcatattttgccc115560C3gtt犯C3Ctacacaatatatcgctatag115620tctatgtcacta/ttaaccctccaa/tcattaatalatttgcgacttactttatcgcccgtc115680aaccggtccccggacacccgcgccttaaacaactagaatacaatttgttELctcaccagtg115740C3tgcacatgC3CCCCCCCttttgaacgactcgttccgcg115800ca犯tttaacatcttactttcactttagaa咖ggtcggttcctatgt幼115860c卿ttgtcgctggtgtccattttattc1:cccgcaggcccct犯ac犯"tc115920cccaatataccggagatctatttagccggttcatttcg幼ccgc犯tatcgccggcccgc115980组ELCtgCCCtttgcagctatcccct幼ggt犯gcatccccacccaataaaattattccc116040cttta犯tac城gtcggcaagcctccttaacgccgcggcggtaagaatgac肪ctg3gc116100ggcgcggcgtcgctccccccttttatagtctg鄉ttgcgcaagctcagtgtt333C3gt116160ggcgcccgaceittttgtcatgagcatgcgc6Lg￡L￡ltt￡LC3tagtg鄉gsaElC33gtg3Cg116220tatagaacgaccgccgccceitagttatgactttgcattttgaatctcagcgcagcggcacl翻0tgtsa犯ccgttttta犯cggcatggggccatatagctccattttatattcaacactagc116340CtCCggMtg犯toctgatgC3gtacggtcgtgcgctgcggtgcggtacgtttgtttagt116400tttgtgtatag8CgCt犯C8atggccgcggctgtttggacagtatatccc■60atattgctatgcgtttgccgc犯cttctacgcc鄉gtatgtttaaatggcgcgatcgga116520gggaaggggggggggtt犯tcc鄉cggagaaggacgctatgtttggaaa116580ttaactctttgtttattgct犯tt3gtgttctgttactaccgtataggct画40tgtccagcactgttacatttgcctcggttgttcca犯gtctgcttttagatatatagtgtl謹0aatttccactatttcggaatgC3tCggtgttgttctgtca3a3gCC3t38116760ctatcccgcgatctctggaaatgttgc犯acgttsa肪t3gtatctgtct116820gtcctgtaaccgtgttgaccatctctatttgcat犯cctgtccgtgatgcCC3gg3t3幼116880t3gtaactgagctatttacttgtgttgatactgg鄉tctgactgtcgttaatgtaatgt116940tag3ta犯tcgtcatattgaaggcaaccttgtagcggcggtacaggtatagtttttgtta117000cagtgeigttgagatctgtatagtgttccgtccccgcagattcacatttaa117060a犯tggatccggcstgtttgcca犯ttcatcagtgacagctttctagttg117120tcattggteigcttgtgtata犯tgtsccattttggtattcccgcaggtatccgcgaccgc117180cgg犯卿tgatcctgccgcCgg犯C犯g3ggtmttgg117240accatgcatt犯agcagccgcgac卿幼aagagcctaatgtatatgagt117300ataCELCCCgtagagtacatatcgaaccacttgtagccattactactacaa117360gattggatgtagteiatatctgtactatgcatttcatatatataccattctggtctgtatc117420taatctaaaccagacacataagcattccatacagagtata117480ttgtccctccgg,cctgtcgtccggctgcaaggtcaccagactttgtt117540caacatccaat犯CttCgLtat已ttgagccgggccgac犯attgcg犯cag已gcagagatt117600ctgttggcgggcccagtcgcctaagtatttcttcatgtgttaatgtttcacaaaagcgct117660cattaaaaaacatattctccgattccatttacgtgtagcagcatcacatt117720tacgccctcgtggtccacca3gCtttgCt3ctatatattc3tggCC￡lgtC3gt3gttt3C117780ctggcctaaa犯agtcttggttagacgcgaatattctgcatcctgtttag117840aaattgtattgatgtaactcgcgattgtatctgttgtatagtcagttcag117900tggatggatcaatcgcaactaaccgtttscattttgcccc3gttgUg3gctaaagtcgc117960gacatatcgctactatggccagctttgtct118020gttctacatctagatttagatcaactcgtctatcattgcatccgcaatac118080g3tcgtc3aagctgcttctgtttctactctcggttgtccaaagaccgtaa118140gtgttggtgtcatacgttgatgg鄉cgcaagaatgacatg犯犯C3ttgtctgcg犯tt118200taactccagtccatccatgtgtc犯aagtattatatcccgcccaggtcct118260ctgttataagtgtgttatatatatcgggtccaatagtgtagtttctgtag118320tgatattccagt犯tgtagtgtagcgg犯cctgccgcctccggcatgtattggttctgta118380cttgtgttgagtagcctt卵gcgattccccatcccctaccacacgtggtc118440gttgtttagctgt雄ggtcctggtctgtttccttttactttctttcactttgggatctt118500ctgttttcatattEigccatggatgtgatagttgt组tggttgttgctcactcgaggcgt118560attgtctaatgggggcgacggttgtcggttcttttccctg犯gttggctttgtctttcgt118620cggtagactctggtccatgtcctattatgcctaagt犯at卿atggggg118680ttgtsgtaaatctctcgcgactctsaataatggcaattgttttctgttctt犯3gC3g3tl腿Otacttacc犯g犯g犯tcca犯tattatggcgatgeitaELtcagcactacaatgccgaaga118800ccccagtggaacctcttggcgtgttgctcgcctctttttggcggeLCg犯g118860aacgtgttccggccgcaccstttctttcac犯C3Ct3ttCcttagcgttc118920tacactttctgaggt犯catcaaccatatcaacaccgcgtctttat犯atgtttccctcc118980aaactcctcccatgtgatgttaagtc幼ca犯ggttt犯catgctcattggceigtattta119040tgtgtg城3C3ttgacacagatgcttatgtatttatcagaccatggg犯catttaacac119100cccctgtatgC￡LtCaCggggtgttagatg3accc犯tggtELtgELtgtttg119160caactagcgcgccatatttatagcacc犯tatgc犯ctELgtgcaccatatttatagcacc119220tagactatatctttat犯tatgtcatgtagatggtttagtacgtgcgcgg119280ccgcggccggg卿ggcgttggacccccctttgccatttcccgggccatcl簡Oggcggggcttttttggcggggggccaa已aggggcgtggcttggcaacatgcc犯ctccsc119400ccactgggcggggcttgggctgttgccaacCCCgCCC3C3ata已gcccctcccatgggagl醒Ogggcttggatcattgccaaccccgcccatggg鄉ggcgatcccgcccatggg鄉agtt119520鄉gttggcagggttcc犯aggttttacgg城tagggttagggttggcagggttccaaa119580ggttttacggtattagggttagggttggcagggttccaaaggttttacggtattagggtt119640agggttggcaggttttacggtattagggttagggttggcagggttcc犯a119700ggtttt3CggtattagggttgggttCC犯8ggttttacggta/ttagggtt119760鄉gttggCELgggttccaaaggttttacggtattagggtt郷gttggcagggttcc犯a119820ggttttacggtattagggttagggttggcatt犯gccaagt鄉gttggccgggactgtc119880咖ggg也gggttggccgggctcc犯aggcagaactgtcttagggttagggctggcctt119940atgttattggat郷gttggccctaagccaagtagggttggcaaggttcc120000ctgtcttagggttagggctggccttaagcc33gtaeitgttattggstagg画60gttgggg卿cgggactccg卿gtagttcgctcttaggattggccccatggc犯gcggg120120gcgggtcttcatccgagaactcggc犯鄉attggc犯tscscc犯ataggcggcgactg120180catctgattt鄉gttggcsttcgcccaagtagg^atattgagccgaggcgctaatagg120240gct鄉gttggccatacgccgagt鄉tggtgactgtagccagattctacctggccttta120300tcc犯gtaggtactactgagccgggtagcatagttattaggactgggggcgttgggagga120360gttggagctaaccctaatcc犯ttagatgcctgtaatggcgttatggttggaccattatc120420atctattagggttgtcggttcctttattctttctttcctcccccacgcgg120480actatgcatctgg3ttggccttgttccagaaa^tcgeLctg3gatagtgttt120540gtggttggcatststccaattaactgtagt3gtct3ca^ttgcatcattgacacaggata120600acactgggga犯ttgggcagacgcgttgcc犯caatgctgCggCg3gt3Cgtaatctcca120660ttca/t犯gaatgtgaggtgsttt犯ggtcacatgcacgcccgattgggcgagtcaaatag120720ggcggcccgagtgggcga_￡Ltttgt已cggacaagcgatt犯ctagttcact120780tgt￡LggggC3tacggccatgcatgcttaccatctttttcagagctctgtt120840atcactttattttacctccctctttttgcgtcaattgcagcctatgtgtgttaaggaatt120900aggacctgtggattactatcatg幼tgtcggccagcctctccgcttggta120960ataaagcggtagttgcccgacagccatgatagtctcccatgaacttcggcggUtsgtga121020tctttgtaactaatattcctttacggactt3catagtatttagacttttgacacttattg121080ggatggggtggatggtttggagacagctgcggtggtctatacatctaataaccatccatg121140caatctccaaaattccaccaataactttttggcccggccatattcccgcc121200cgtcactttttac肪gacaggttgcatttatttattctta121260actatcgagctgaat3tgtcg朋gacctctatttcttt犯aaactggataC3tagcgaag121320ctattttctggctgtcta犯犯aatggagaatatataatacggtgcaggt121380BHaaaasstctcttaacaaa犯gtcacgacaagctcccgcgtacggattg121440ccctcatcccccg犯tcatccccatcggactttttatcttatgtggtgct121500ttgggC3ttgg3gtta^ag3tggctgtacggcagtaattgtaacatcatcctcattagat121560tgttcattgggcgggat組agcctgcttcatatctccggcctcgatgctagtagcccta121620gaccccccgtgacatEigttcccaggttattgtaggtggacgttggccgcgctggatagta121680tttgga^ttgaattaccctccatttgcggttcgagtaaagcagctgcacag121740tatgagcggactacttctggeiatgtcgitctggggtctc犯aaggatgcttccggctctct121800gtcaggtaca犯ggcctcagaggtgctcctagaccccatctatttgctgccctgtgatta121860taagcagccccttctaga犯atctggtcctcttatactgagtccatctgcctggaaggcc121920atatccttgcggtcgtattgcgg犯gaaca鄉ccgcgatattcacggggggtcattggc1219803cgcgagtagtgcgaccggcctttgtctcgac犯catstttaacattatcacctctcatc1220403gCtgt犯tgggggttcattBgggtat已gctgagcgaacgcggcttcgacccgcgcaaag122100actccaggccccactatcccacgtgatgtcatcactacccttgacatggcttcacaatct122160ggccatctgtgggccccatgggcctggggagttatagtcgcctttatatatgaatgatag122220ctact犯ttgctggaccgtcggcgggcc犯tccactgtatcaattgtatccaggaa幼tg122280agccgcttgtgggccgagccg3gcctgaggca^gEit^ttc犯ccgcccccgcaaatcct122340aaatctctaacggEitagc已gCagCELCgCCCCtgg6LElttgELgtgcagaaatatctgggacc122400ccagtccagttacccgcccatgcgtgactgccgtatg犯caaaggcggtcacacaaggcg122460gctagc鄉a鄉卿ggccacctttttcatCag￡LCC3C3t鄉catgcaagaggccact122520ggggcggcgcttaattcctcccctacgagcgggcagtac3tgeitgaccacctttacatcc122580tcagggtctggaatctggttcatcceiagcacattgtttgcggagcggatcagtgactcgc122640鄉ggcccgtatagtcttccttcgacatggcttccgccacgtcgtgccgccacgcgggcc122700agtgcctccgt犯agcgggtttccagttagg3gg犯3C3tagggtcatct122760犯t犯ttctgcgactatgtcgggctgg認taggtagaccgtgccgactcggttggcagc122820ggagtatgtgttattccggtggggacgcgcctatatcccccgcggggatggggcccgcct122880tgcggcattggaccgagagccttgaccgacggcagtgtccacggtaactcggtttggtta122940tcctttttagaattttgta/tattctgatgttttggcaagccatctggtctatctcggtcc123000tttggggcagtatcatctgtctgggcatatgccccggacccattsctsggcacagaatgg123060cgctttgtttggggcggcccatgaacEitcatcggctccattatctataccgtctcg鄉g123120卿gcttggacgggcat卿tttgcgcttagaagcaaatcUcgtggctgttctcccttc123180ccgttggcccgctcagacccaggtatctgctgacattgcttattgacacgattggcgcca123240tctatcttacgttccctgtcttgcttgtggcagttggtttggcaggccacsctacgggtt123300tggtccttgcgcttagatcccccgtcttgctggatgccgctcccgcttgtcatgtattta123360tcaatggtctttgacgcgattggccagacaagcggcgacgtttgtttgtctttgggggta123420agatgacttaacacccaggatagtttggcgccattgatcttggctgacccatttccgctc123480ggggcag咖agagcgcctctgttsggttccatgcagaatatctacgcctagccctgtcc123540acggaaaagtttacaaagggtctcacagtgctaatcaccgtaccaacagcacgaagcgcc123600attcaatgccctgcgacgatg犯agctcgccccgtagtctcatagcattt1236603t3C3t3t3ggcaacagttta鄉tcctttatccatgatageiagctggacgtgatgccgg123720tggagctgggcagacggaca紐cagcgtcc3gCtCC3gtg3gg33at3CCaggsatgttg123780ccggcaccaccaaagcactcgagcaatscctctattactgcatteicaagctaaaacgcac123840gcatctgccagttcctcggttgagt犯tcagatgcatgttcgggatcacaaaagtcgggg123900tccgccgcgctctgctctgtCgCC3g3tggtccgtggactgcggtgtgttctgcggctct123960atctgcgagtcgccgcgaaccgtcacagtctgcagaccctcggtagcttcgcgcatgctg124020cggcggacagcattcattcggcgataaatggaageiaatgtcatccgttagagatgcggag124080gcstc朋cggcctttgcggggaaaagcatgt犯gcatacgccctccgcagagacatttgc124140agaaattgtttttgcactttatcatagcggcggctcatcgctacggccgcggcaacstgc124200ggtagaccccatagagcatttccagccgccatcgcatctccaatatgtggtagaggtcgc124260gcaacacttccagtaataaatgacccggaaccagtaccacccctcagattgcattgtttt124320tgacaaacttgctccaataacagatcctcaccagttattttgttacattgtatccagccc124380atagcatcgaaactattgctatatagtgcttttacaagagcatcatactgctttgatgaa124440tcccccatctCtCCCElCEl犯wc站ccaccggaccgcggctctcggcataccgctctctg124500gcataccgtatctctggttcatcccacagcccctggcgatagtccccagtaccgccatac124560cgcaccttgcc鄉tggcggtggctcagacCCgggCC3tggatccccatttgtcattagc124620aaaggtccgttcatagtaggcatttctgggacgcctattgacatacaagactcagctccg124680gcgctagatgatgatactcttttcatcgccgatggccgtctccccctacccctacggggg124740cgctggtaaaatgcgtteitttgccgctggtgg鄉ttcgcaggagact幼atctatggtg124800tcatctgatgatgggtgttgatattgagtggactcgcgagggctattcgcgagcatagtt124860ggttgttccatagtctggggctgctgttcggtttcttcttgctgaactgcgcggcgtacg124920ggttgcttgcccttgcgcttgacggtgcccgtggccttgcttgcagcggtgttactggtc124980ggtgggtgtgttaattgctgcgaaaacagttttgataaatC3gtCtttggggaccgctgc125040tccgcattatcctttttacgctgcatggcgggcctacgcttctttggtgcagatggaata125100犯tg3tgg3ggggggcgctttggtgttggagggcggtctggggatggtcc125160gagctcccgtccgaggaggaggaggacgeicgstgtggatg8gCtgg33Ctggacctggac125220cgcccccgatcgacagatctgcgcggcgeicgcggctggcgagcaacatgagactataggt125280agcgcgggcggcgagcggcccctgggggcagaccttggcgatgt卿cggggggccggaa125340cgttccacgacaccagtttgaggcgatcgcgccggcgacctataccaaccggccg鄉ta125400acagaggtcggtgatgacttgggtgtcatggtgcgcccactgtcggcacatctagcaggg125460ggtggcgatgccgatgaagatggtgagcgatggcgcctccgagcggtattcttgggcggt125520gctgctgtcggtgeiccgggc3ggtg幼gCg3tcggtttatcaggagccgatggtcgcccc125580<formula>formulaseeoriginaldocumentpage57</formula>tataggtcgtcgtccgctacgatgcataccgtggttggccgacacctctcatcagatctatggttcatttaacc犯catcatatagtcgc3ta犯aaaaa犯g犯3gc犯8gttgt3tgttgttattcggcaagccagatcggtcgatttgtttagggggcgggggtatcgacccccccggcgtagaggtttgtaccgtcgttaagacaggcctaggatttattttgtaGtcgctttaaatttgagcggataggtcgcctt3g3tagagttattatgttttgcccgggca犯tgtttgtcgggtgtccggcgctgctaatagatcccgtagaacggaaaccctcattaccttctgtgcccctgtatgttgccattcttattaaaaggctgtgtacttaaatacactatgtacacttaagacgacactstccctgtg犯ttga33g3tatgaatcggtcctatagttggttttgcgagacagccatcaactgctttattcagtgctggtgttgttcggccttactttccaggaacgtggcatctgctgtgatgtagcagtttgttcaaact犯ccag3cac朋犯cgtt犯gag犯tstgga犯actataactatgtt犯c犯gttc犯tccagtttaaagcacgatcatcgtt朋cgccaaaacgccagatggca犯atac犯g8tg已gtaactttcattcattattacaaactgtactcgt3C3at3tg犯tg3C3犯agtc3t犯aai3tcggatgtggatcccaatctctcttccccc犯g犯tttttggaggct犯atacgcataaccagccgtatcgt幼犯gtggcggcgggagcgcgccctcgtcgccggaaacggttcgtaggcgattaggttgtatcccattttagtccacactcgcgatct犯aagEigtgg犯犯Eiac犯t犯aaccgagtctcagttcga犯c犯tcgctatcgtttatgagaaagggggccgtgtagtcccgtatcgttacgtcttgaatggtcgctcctgggctcgggtgccgctttttatagtctca￡LCCCgg￡LCCgttCgCELCtCtggtttCC犯tgtgCg犯CgCCagatttgaCC犯tCgCElttcacatattcgcactccga犯tactgtgttggaactct犯tgcatattcataaccccacgeicgtggttattcccagagctgccttca犯ctgcgatcgcgtgtatataattatatcattgccaatatgaccgtaacttgacctttcctcctcaccgcacgctagtgtctatactgttcgacattgatatacggttcgcattcctg犯t犯aggaatcgcgtttaggatggcacc犯act犯tagtatacgagacggttaatttgcggttac128340gt3犯tCttgccgatatcat128400acatgcgatcgtgtgtatcgattaataaaa128460ctctgagtacttactgcgcggcgcctgcga128520ttcatggccgatagtaa犯atatttgccca128580ggcgt卿犯cgggca^犯aaccaatttt128640ttcagccgttcgtgcattacatttatataa128700catactgtgtcatgtttcaaactttccgcc128760tcaatttgattgggtg卿c3tataggcgc128820atc犯ggcggcagtattgacaaagaatcgg128880tacaggcaataaaacacaat128940ccgg犯taggataattagcg1290003gtC3Ct3tgttCtg3gg犯33tatatccg129060C犯3Ctg3gCggtttatg犯aaaccattaa129120atctaggtccaccattcccctgattat3tg129180gcttt幼attgccccc犯aa129240gccgccagccctcaattcccatcccgctgc129300gcctattagtttsaagcagc129360gcagc3ga幼129420attttattatactcgcttata3gcg3gtt3129480g3cac8犯gt3Ctggctaaa1295403cacggtacagttgacgctc129600caac3ccg犯gg3c犯gatcgctatcggca129660ctcgaccctgtg8t幼8犯Cgccattattc129720a犯tcattccttgcagtcaccatcgtaatg129780atcccgatagttatagtcgg129840taaatgcagtgatagtcactgacagcggtg12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t犯aggtggtcatcatgtactgcccgctcgtaggggagga154860attaagcgccgccccagtggcctcttgcatgcct已tgtggtctgatgaaa鄉gtggcct154920ctccttcctgctagccgccttgtgtgaccgcctttgttcatacggcagtcacgcatgggc154980gggt犯ctggactggggtcccagatatttctgcactcaattccaggggcgtgctgctgct1550403tCCgtt3g3g3tttaggatttgcgggggcggttg組atctatgcctcaggctcggctc155100ggcccacaagcggctcattttcctggatac￡t￡Lttg￡lt￡LC3gtggattggcccgccgacgg155160tccagcaattagtagctatcattcat已tstaaaggcgactataactccccaggcccatgg155220ggcccacagatggccagattgtg犯gccatgtca鄉gtagtgatgacatcacgtgggat155280agtggggcctggagtctttgcgcgggtcgaacccccattacagctgaxgagaggtgataatcgcactactcgcgtgccaatgaccccccgcgaccgcaaggatatggccttccaggcagaagaaggggctgcttataatcacagggcagcgcctttgtacctgacagagagccggaagcaagtagtccgctcatactgtgcagctgcttttgtggtcaatccaaatactatccagcgcggatgtcacggggggtctagggctactagcatcccgccc犯tg犯c犯tctaatgaggatgaaactccaatgcccaaagcaccacataagatttcgggggstg鄉gc幼tccgtacgcgggtaagagattttttttacctgcaccgtattaatacacagaaaatsgcttcgctatgtatccattcagctcgatagttaagaataaagtctt城ttaaaagtgacgggcgggaatatggccgttgtttggagattgc已tggatggttattaccatccaccccatcccaatsagtgtc犯aatattagttaca犯gatcacta犯ccgccgacaactaccgctttatatttcgtttgtaccataatccacaggtcctaattccttaacacac已gccgcgttcgctcagctataccct犯tga155340gttgtcgagacaa^ggccgg155400tgaatatcgcggccttgttcttccgcaata155460tggactcagtcagattttct1555203^tagatggggtctaggagcacctctgag155580tccttttgagaccccagatgacattccaga155640actcgaaccgC3幼tgg鄉gt肪tgtstg155700ccaacgtccacctac犯taacctgggaact155760cgaggccggag3t3tg犯gCsggctattat155820tgttacaMtactgccgtacagccatcta/t155880atcttgtccgatgggg3tga155940昭cttgtcgtgactttttctgattttttgt156000tatattctccatttttttagacagcctttt156060agttttt犯atcttcgacat156120gc犯cctgtcttgtagttct156180gggcc犯aaagmttggtgg犯ttggtag156240gatgtatagaccaccgcagctgtctccaaa156300gtctaaatactatgt犯gtccgta犯gg^1563603gttC3tgggagactatcatggctgtcggg156420agcgg卿ggctggccgacattcatgatag156480at15651权利要求1.鸭肠炎病毒基因组DNA序列，其特征在于，全部核苷酸序列长度为156512bp，至少编码78个开发阅读框，基因编码的氨基酸序列与α-亚科疱疹病毒具有同源性，其中5个基因仅与目前所知的禽疱疹病毒有同源性，基因组结构与水痘病毒属相一致，该序列为SEQNO.1。2.—种鸭肠炎病毒基因组DNA的提取方法，其特征在于，具体步骤如下(1)将收集的病毒样品30,000Xg离心30min，使细胞和游离病毒粒子沉淀；(2)将歩骤(1)所得沉淀用灭菌蒸馏水悬浮，一85'C23。C冻融3次，30,000Xg离心30min，所得沉淀重悬于预冷的Cellpermeabilizationbuffer,置冰浴10min，30，000Xg离心30min,弃上清，取沉淀，重复本步骤一次；(3)重复歩骤(2)离心后，弃上清，将沉淀悬浮于500iUCellnuc]eibuffer,再加入500ixl含600U微球菌核酸酶的2xNucleasebuffer,混合均匀，37。C作用30min;(4)加入60ul0.2MEDTA终止反应，加入4mlDigestionbuffer,用剪去末端的Tip头轻柔混匀，避免对病毒DNA的剪切作用，5(TC过夜；(5)加入等体积酚/氯仿，轻轻混匀，8，000Xg离心5min，水相再用氯仿抽提1次，8,OOOXg离心5min;(6)室温条件下，取上层水相，加入0.5倍体积的7.5M醋酸铵和2倍体积无水乙醇，轻轻混匀，16,000Xg离心5min;(7)歩骤(6)所得DNA沉淀用70X乙醇洗涤2次，晾干，即可获得鸭肠炎病毒基因组DNA。全文摘要本发明公开了一种鸭肠炎病毒(DuckEnteritisVirus，DEV)基因组DNA的提取方法和基因组序列。首先利用高渗缓冲液裂解感染DEV的鸡胚成纤维细胞(CEF)，然后加入核酸酶消化细胞DNA，再用酚/氯仿抽提的方法获得病毒基因组DNA。将所得样品随机打断为2.0～2.5kb片段，连接入pUC19载体，转化宿主大肠杆菌构建基因组文库。对文库中的阳性克隆进行随机测序，测序总长度对病毒基因组的覆盖率为6倍。测序结果经拼接和组装，获得了全长为156512bp的DEV全基因组序列，分析发现基因组结构与α-疱疹病毒亚科水痘病毒属成员一致，为DEV的分类提供了理论依据。文档编号C12N15/33GK101182526SQ20071011422公开日2008年5月21日申请日期2007年11月12日优先权日2007年11月12日发明者于可响,史玉颖,静吴,宋敏训,李玉峰,马秀丽,兵黄申请人:山东省农业科学院家禽研究所