本發(fā)明屬于基因工程
技術(shù)領(lǐng)域：
，具體涉及一種在真核細胞中表達固氮酶基因的方法。
背景技術(shù)：
：氮肥的使用與我國糧食、能源、環(huán)境等可持續(xù)發(fā)展重大問題的矛盾日益尖銳，生物固氮是全球植物所需氮素的主要來源，約占75%，利用生物固氮，可減少氮肥的使用，是解決該矛盾的重要措施之一。然而，迄今為止沒有發(fā)現(xiàn)任何具有固氮功能的真核生物，已知的能夠固氮的生物只限于原核微生物，如細菌和放線菌。生物固氮是指固氮微生物利用其自身的固氮酶進行固氮反應(yīng)的生物過程。固氮酶是將空氣中的氮氣還原的催化反應(yīng)中心，是一種復(fù)雜的多蛋白復(fù)合體，至今僅在原核生物中發(fā)現(xiàn)，沒有在植物中發(fā)現(xiàn)固氮酶基因的報道，也沒有發(fā)現(xiàn)任何具有自主固氮能力的植物。因此開展固氮基因在植物中的表達研究，達到提高生物固氮效率、擴大宿主范圍，發(fā)揮生物固氮在農(nóng)業(yè)生產(chǎn)中的作用，將確保我國的主要農(nóng)作物生產(chǎn)在繼續(xù)提高產(chǎn)量、改良品質(zhì)同時，大幅度地減少化肥、農(nóng)藥的施用量，實現(xiàn)我國農(nóng)業(yè)可持續(xù)發(fā)展。固氮生物的固氮反應(yīng)是在固氮酶的催化作用下實現(xiàn)的。固氮酶非常保守，由nif基因組編碼。多數(shù)固氮生物中的nif基因組包括16-19個基因，總長度為23kb，其中包括8個必需基因：nifhdkenxbq。典型的固氮酶由鐵蛋白和鉬鐵蛋白兩部分組成，通過鐵氧化還原蛋白和黃素氧還蛋白等電子供體還原鐵蛋白，將單電子從鐵蛋白轉(zhuǎn)移到鉬鐵蛋白(要依賴mgatp水解)，最后電子在mofe蛋白內(nèi)部從p簇轉(zhuǎn)移到femo簇底物結(jié)合位點。每一次電子轉(zhuǎn)移都要求鐵蛋白與鉬鐵蛋白組成一個復(fù)合體，轉(zhuǎn)移結(jié)束后兩個組分再分開。目前所有的固氮生物均有一個鉬-鐵固氮酶系統(tǒng)，在鉬缺少的條件下，一些固氮生物會誘導(dǎo)其它輔助因子固氮酶的合成，如釩-鐵或鐵-鐵輔因子固氮酶。固氮酶對氧敏感，有人推測這是由于該二聚物的兩個結(jié)構(gòu)域之間4fe-4s簇的表面暴露所導(dǎo)致的。固氮酶的固氮反應(yīng)具有以下基本特點：1、需要金屬離子和atp；2、對氧敏感，需要在無氧或低氧條件下才能發(fā)揮作用。正是由于固氮酶的以上特點，迄今為止發(fā)現(xiàn)的所有固氮生物只限于原核微生物（細菌和放線菌），這些生物的共同特點是厭氧，除了一個特例——藍細菌。藍細菌是一類放氧性光合生物，在光下，會因光合作用放出的氧而使細胞內(nèi)氧濃度增高，但同時它又有厭氧的固氮系統(tǒng)，它在長期進化過程中形成兩種獨特的抗氧機制。1）分化出特殊的還原性異形胞。某些絲狀藍細菌可在細胞鏈的中間或末端形成異形胞，體積大且有較厚外膜，可防止氧氣進入細胞，此外異形胞只有光合系統(tǒng)ⅰ，因而光合作用不放氧，因此固氮酶在異形胞中可以很好地發(fā)揮固氮作用；2）非異形胞藍細菌固氮酶的保護多種多樣：有的采用將固氮作用與光合作用進行時間上的分隔，黑暗下固氮，光照下進行光合作用，如織線藍菌屬等；有的則形成束狀群體，在其中央處于厭氧環(huán)境下的細胞失去光合系統(tǒng)ⅱ，有利于固氮酶在微氧環(huán)境下進行固氮作用，如束毛藍菌屬；有的則在固氮酶活性高時，細胞內(nèi)用以除去有毒過氧化物的過氧化物酶和超過氧化物酶的活力也均提高，如粘球藍菌屬。lucasj等1985將不產(chǎn)生異型胞的藍藻oscilla-toriasp.，無氮培養(yǎng)基上培養(yǎng)，設(shè)置不同的光暗循環(huán)，結(jié)果在黑暗時間超過8小時條件下，只在黑暗時段固氮；如果黑暗時間少于8小時，固氮酶活性在黑暗開始之前就已經(jīng)有顯示。即使是產(chǎn)生異型胞的藍藻，營養(yǎng)細胞在缺氧的條件下也可以誘導(dǎo)固氮酶，引發(fā)固氮反應(yīng)。這也說明固氮反應(yīng)不僅局限于厭氧細胞中，確切地說這應(yīng)該是一個產(chǎn)氧細胞中的固氮需求與高濃度氧反饋抑制的平衡問題。利用葉綠體具有的與固氮基因相似的原核特性及其它有利因素，程奇提出在葉綠體中表達固氮基因的設(shè)想。程奇等2005年，將klebsiellapneumoniae的固氮酶的鐵蛋白基因nifh轉(zhuǎn)化chlamydomonasreinhardtii葉綠體并替代黑暗條件下在葉綠素合成過程中起作用的dpor蛋白的chll，結(jié)果轉(zhuǎn)化衣藻在黑暗條件下仍然可以合成葉綠素，說明nifh在衣藻葉綠體中可以發(fā)揮功能，這是葉綠體表達固氮酶設(shè)想的成功前期探索。張中林等2001年，成功將固氮酶鐵蛋白nifh基因?qū)氲礁叩饶Ｊ街参餆煵莸娜~綠體中進行表達。但至今為止沒有關(guān)于鉬鐵蛋白成功在植物中表達的相關(guān)報道。目前，也有一些利用體外合成的方法，對固氮酶基因的組分進行研究，l.m.rubio等2008年報道，在鉬，鐵，硫磺和高檸檬酸存在的條件下，僅用nifb,nifen,nifh三個基因即可完成鉬鐵蛋白的體外生物合成。yilinhu等，2009年研究表明，nifen具有乙炔還原活性，卻并不能將n2還原成nh3+，這說明nifen具有與鉬鐵蛋白相似的催化作用，但是它僅僅是鉬鐵蛋白的一個“骨架”，它在結(jié)構(gòu)上更簡單，功能上也更少，因此，它的反應(yīng)底物范圍更窄，底物的還原活性也更低。盡管人們在生物固氮上進行了大量的研究，但目前沒有任何關(guān)于鉬鐵蛋白在植物中成功表達的報道。技術(shù)實現(xiàn)要素：本發(fā)明的目的是為了解決現(xiàn)有技術(shù)的不足，而提供一種在真核細胞中表達固氮酶基因的方法，實現(xiàn)了固氮酶基因在植物中的表達，從而為植物自主固氮奠定基礎(chǔ)。本發(fā)明采用如下技術(shù)方案：一種在真核細胞中表達固氮酶基因的方法，包括以下步驟:步驟一：擴增所述固氮酶基因片段；步驟二，構(gòu)建重組載體：將所述固氮酶基因片段連接至病毒轉(zhuǎn)化載體；步驟三：用步驟二構(gòu)建的重組載體轉(zhuǎn)染所述真核細胞；步驟四：篩選表達所述固氮酶基因的植物細胞。進一步地，步驟一中所述固氮酶基因片段為nifh、nife、nifb和/或nifn。更進一步地，步驟一中擴增所述固氮酶基因片段時的引物如seqidno.1、seqidno.2、seqidno.3、seqidno.4、seqidno.5、seqidno.6、seqidno.7及seqidno.8所示。更進一步地，步驟一中所述擴增時的pcr反應(yīng)體系為：50ul10pmol上游引物，10pmol下游引物，2ul模板dna，4ul2.5mmdntp，5ul10倍extaqtm緩沖液，0.5ultakaraextaqtmpcr反應(yīng)條件：94°c5分鐘94°c30秒，60°c30秒，72°c2分鐘，35個循環(huán)，72°c7分鐘。進一步地，步驟二具體為：將nifh/nife/nifb/nifn基因，分別通過clai/sali，eagi/sali，eagi/sali，clai/sali酶切位點連入載體pvx表達載體pvx-hisg-mf1，得到pvx-hisg-mf1-nifh/nife/nifb/nifn載體。更進一步地，步驟二中所述pvx-hisg-mf1-nifh、pvx-hisg-mf1-nife、pvx-hisg-mf1-nifn、pvx-hisg-mf1-nifb載體的序列如seqidno.13、seqidno.14、seqidno.15、seqidno.16所示，克隆得到的nifh、nife、nifb、nifn基因序列如seqidno.9、seqidno.10、seqidno.11、seqidno.12所示。本發(fā)明與現(xiàn)有技術(shù)相比，其有益效果為：實現(xiàn)了固氮酶結(jié)構(gòu)基因在植物中的表達：本發(fā)明實現(xiàn)了固氮酶結(jié)構(gòu)基因在植物中的表達，從而為植物自主固氮奠定基礎(chǔ)。固氮酶結(jié)構(gòu)基因僅存在于原核生物中，而原核基因在真核生物中的表達并不是特別容易的，目前為止，已報道的在植物中成功表達的固氮酶結(jié)構(gòu)基因僅有nifh。原核生物基因分為編碼區(qū)與非編碼區(qū)。非編碼區(qū)上的基因決定某些性狀是否表達，表達多少次以及何時開始表達。編碼區(qū)能轉(zhuǎn)錄為相應(yīng)的信使rna，進而指導(dǎo)蛋白質(zhì)的合成。非編碼區(qū)則相反，但是非編碼區(qū)對遺傳信息的表達是必不可少的，因為在非編碼區(qū)上有調(diào)控遺傳信息表達的核苷酸序列。非編碼區(qū)位于編碼區(qū)的上游及下游。在調(diào)控遺傳信息表達的核苷酸序列中最重要的是位于編碼區(qū)上游的rna聚合酶結(jié)合位點。rna聚合酶是催化dna轉(zhuǎn)錄為rna，能識別調(diào)控序列中的結(jié)合位點并與其結(jié)合。t7啟動子，是來自于t7噬菌體的能夠?qū)7rna聚合酶有特異性反應(yīng)的強啟動子，是啟動t7噬菌體基因轉(zhuǎn)錄的一段序列。t7rna聚合酶能選擇性的激活t7噬菌體啟動子的轉(zhuǎn)錄，其合成mrna的速度比普通大腸桿菌的rna聚合酶快5倍左右。植物病毒載體（pvx載體）表達外源蛋白表達水平高、表達速度快、宿主范圍廣。本發(fā)明利用t7啟動的pvx載體成功在植物細胞中表達了nifh/nife/nifb/nifn。附圖說明圖1為heliobacteriumchlorum中nifh,nife,nifb,nifn基因的pcr克隆。1.marker1000,2.nifh,3.marker1000,4.nife,5.marker2000,6.nifb,7.marker2000,8.nifn。圖2為pvx/hisg-nifh,pvx/hisg-nife,pvx/hisg-nifb和pvx/hisg-nifn載體的pcr檢測。a1.marker2000,a2.nifh陽性對照,a3-8.nifh1-6單克隆檢測.3號克隆測序結(jié)果與ncbi上公布的nifh序列同源性達100%..a9.陰性對照；b1.marker2000,b2.nife陽性對照,b3-8.nife1-6單克隆檢測.4號克隆測序結(jié)果與ncbi上公布的nife序列同源性達100%.b9.陰性對照；c1.marker2000,c2.nifb陽性對照,c3-8.nifb1-6單克隆檢測.3號克隆測序結(jié)果與ncbi上公布的nifb序列同源性達100%.c9.陰性對照；d1.marker2000,d2.nifn陽性對照,d3-8.nifn1-6單克隆檢測.3號克隆測序結(jié)果與ncbi上公布的nifn序列同源性達100%.d9.陰性對照；圖3nifh/e/n/b蛋白在煙草中表達的western檢測。a1.蛋白marker;a2.轉(zhuǎn)染pvx-hisg-mf1-nifh載體的煙草葉片總蛋白;a3.對照煙草葉片總蛋白;b4.nifh-hiswestern雜交檢測;b5.對照煙草葉片蛋白的western雜交檢測；c6.轉(zhuǎn)染pvx-hisg-mf1-nife載體的煙草葉片總蛋白;c7.對照煙草葉片總蛋白;d8.nife-hiswestern雜交檢測;d9.對照煙草葉片蛋白的western雜交檢測；e1.蛋白marker;e2.對照煙草葉片總蛋白;e3.轉(zhuǎn)染pvx-hisg-mf1-nifb載體的煙草葉片總蛋白;f4.對照煙草葉片蛋白的western雜交檢測;f5.nifb-hiswestern雜交檢測；g6.轉(zhuǎn)染pvx-hisg-mf1-nifn載體的煙草葉片總蛋白;g7.對照煙草葉片總蛋白；h8.nifn-hiswestern雜交檢測;h9.對照煙草葉片蛋白的western雜交檢測。具體實施方式下面結(jié)合實施例對本發(fā)明作進一步的詳細描述。本領(lǐng)域技術(shù)人員將會理解，下列實施例僅用于說明本發(fā)明，而不應(yīng)視為限定本發(fā)明的范圍。heliobacteriumchlorum，厭氧菌，用heliobacteriumchlorummedium370培養(yǎng)基培養(yǎng)。heliobacteriumchlorummedium370培養(yǎng)基：k2hpo41.0g，mgso4x7h2o1.0g，yeastextract10.0g，distilledwater1000.0ml，調(diào)節(jié)ph7.0。充入氮氣，加入sodiumascorbate(0.5g)繼續(xù)充入氮氣。調(diào)節(jié)ph6.8.將45ml培養(yǎng)基倒入50ml容量瓶中，用膠塞旋緊。121°c滅菌15分鐘。低光照條件培養(yǎng)heliobacteriumchlorum。實施例1鉬鐵蛋白結(jié)構(gòu)基因nifh/nife/nifb/nifn的克隆依據(jù)ausubeletal.(1995)的方法提取heliobacteriumchlorum基因組dna。(ausubelfm,brentr,kingstonre,mooredd,seidmanjg,smithja,struhlk(1995)preparationofgenomicdnafrombacteria.in:currentprotocolsinmolecularbiology.wiley,newyork)在ncbi網(wǎng)站nationalcenterforbiotechnologyinformation(http://www.ncbi.nlm.nih.gov/)搜索heliobacteriumchlorum中的nifh/nife/nifb/nifn的同源序列片段，利用primer5軟件設(shè)計引物。引物序列見表一?？寺〉玫搅?57/1388/831/1306bp的nifh/nife/nifb/nifn基因序列，測序結(jié)果顯示與ncbi網(wǎng)上公布的序列同源性100%。pcr反應(yīng)體系：50ul10pmol上游引物，10pmol下游引物，2ul模板dna，4ul2.5mmdntp，5ul10倍extaqtm緩沖液，0.5ultakaraextaqtmpcr反應(yīng)條件：94°c5分鐘94°c30秒，60°c30秒，72°c2分鐘，35個循環(huán)，72°c7分鐘。表1heliobacteriumchlorumnifh/nife/nifn/nifb基因引物序列表引物序列seqidno.nifhncacggaatcgatcgtcagatagccatttacggaaaaggtg1nifhcgtctcagtcgacgattagcccagtttggccatc2nifenacgcgtcggccgtttgtcaatcttaccaaactggacttaaaag3nifecctgtcagtcgaccgttggtgctgccatataggttgattg4nifnncacggaatcgatactggtccggtagctgaacctcgcag5nifncgactcagtcgacttaagtccttatcccgccatggcgtttaag6nifbnacgcgtcggccgggttgcactggttcttgcggaggaaattc7nifbcgtctcagtcgacctatatgatgccgatggcgtcagctc8實施例2構(gòu)建nifh/nife/nifb/nifn病毒轉(zhuǎn)化載體及轉(zhuǎn)染nifh/nife/nifb/nifn基因，分別通過clai/sali，eagi/sali,eagi/sali,clai/sali酶切位點連入載體potatoxpotvirus(pvx)表達載體pvx-hisg-mf1。pvx-hisg-mf1-nifh/nife/nifb/nifn載體經(jīng)測序驗證，已成功將基因克隆到表達載體。pvx-hisg-mf1-nifh/nife/nifb/nifn經(jīng)體外轉(zhuǎn)錄試劑盒作用后，用于煙草nicotianabenthamiana的轉(zhuǎn)染實驗。轉(zhuǎn)染之后，在20-26度，12小時光照，12小時黑暗條件下培養(yǎng)煙草2-3周，篩選具有病毒侵染斑點癥狀的煙草葉片進行蛋白質(zhì)sds-page及western雜交檢測。實施例3轉(zhuǎn)基因煙草sds-page檢測及westernblot分析轉(zhuǎn)基因及非轉(zhuǎn)基因煙草葉片總蛋白用植物蛋白提取試劑盒進行提取。得到的轉(zhuǎn)基因及非轉(zhuǎn)基因煙草葉片總蛋白進行sds-page電泳檢測。利用anti-hisantibody(invitrogen)抗體及horseradishperoxidase-conjugatedsecondaryantibody(invitrogen)二抗進行western雜交檢測。結(jié)果顯示nifh/nife/nifb/nifn蛋白均在煙草中獲得了成功表達。本發(fā)明中涉及的其他序列如下：nifh序列（seqidno.9）：atgcgtcagatagccatttacggaaaaggtggtatcggtaaatctacgaccacccaaaacaccgtctccgccctcgccgagatgggtaaaaaaatcatgatcgtcggttgtgaccccaaagccgactctactcgtttgattcttcactccaaagcccaagcgacggttatggacctggcccgtgaaaaaggtactgtggaagacctcgaactggaagacgttctcctcaccggtttcgccgacatccgctgtgccgagtccggcggtcccgaacctggtgtaggctgtgccggccgcggtgttatcaccgccatcaacttcctcgaagaaaacggcgcttacacccctgacctggactatgtgttctacgacgttctcggtgacgttgtctgcggcggtttcgccatgcctatccgcgaaaacaaagcccaagaaatctacatcgtgacctccggtgaaatgatggccatgtacgctgccaacaacatcgcccgcggtatccttaaatacgctgctaccggtaaagttcgcctcggcggcctcatctgtaacagccgtaaaaccgacaaggaatacgaactgattgccgagttggccagccgtctgggcacgcaaatgattcacttcctgccccgtgacaaccaagtgcaacgggccgagctgcgccgcatgaccgttatcgaatactctcccgaacatccccaagccgacgaatatcgcgctctggccaagaaaatcgacgagaacaaaatgctcgttgttcccacaccgctttccatggacgaactggaagacctcctcatctcctacggcatcctggaagacgaagaaacggcgatggccaaactgggctaanife序列（seqidno.10）：gtgtttgtcaatcttaccaaactggacttaaaagaaacatgtccccttactgataaaggggcgccgaaactgtgtatgaaagctctgcccggtgaaggtgctgagcgttcctgtgcattcgacggcgcaagagtcgtcctcatgccggtgaccgatgccgcccacgttgtccatggtcccatcgcttgcgcggggaactcctgggacaaccggggcgccaagtcttcgggaccccaactctatcgccgcggattgacgacagacctgatggaaatggatgtcgtctacggtggagagaagaaactcattcaagccatcatcgatgtggcccggaaacaccagcccaaagcggtattcgtctatgccacctgtgtctcggctttaatcggagacgatctggaaacggcctgtaaggaagcgcaaaaagacgtctcagtcccggtcatcccggtaaacgctcctggttttatgggcgataaaaacatcggcaaccgtgtcgccggcgaagtcctcttccgccagatcatcggcacggctgaaccagaaacagtccaccctttcgccatcaacttcatcggtgaatacaacatcgccggcgacttgtgggggatccagccctccttagaggctatcggtattaagctgcaagcgtgcatcagcggcgatgctcgttttgatgacatccgtacggcacaccgagccaaactgaatgtgctggtttgctcgaagagcttgaccaacctggtcaaacagatggaacaaaaatatggaattccatacatcgaaggatccttctatggcatccacgatacaagtgctacactccgagccatcgccaaagctttcggagacgccggcctgatcgaacggacggaagcccatattctccagaaggaagcggaaacgcgagccgtcatcgagccccttaaggcgcaactcagcggcaagagagccgtcctcttcacgggtggcgtgaaaacctggtccatggtcacgaccctgcgggaattagggatcgacgtactggctggtggtacgcagaactccacaccggatgacttcttgcggatgaaagccttgatggacccgacggctcatatcatcgaagataccagtgccgccggatttctaaccctgatcaaagaaaaaagccccgacctcatcgtggccggaggcaaaacgaagtacctggcccataagactcggactccttttctggatatcaaccacggacgcagccttccctatgccggctacgccgggatgatcacctttgccaacgccgttgcccgtactgtattcagccccgtctgggaaaaaatgagggtagcttttccgccggaggtgacggtcaatgagtaacgcaatcaacctatatggcagcaccaacgnifb序列（seqidno.11）：atgggttgcactggttcttgcggaggaaattcaacgaaagacgtatccaagcacccctgtttttcacctcatggccacggaaaatcaggaaggattcatctgcctgtcgctcctggatgtaacatcgcctgtggttattgtgtgcgtaaattcgattgtgccaatgagtcacggcctggcgttaccagccgagtgatcactcccgaacaggccctctggcgcgtagaacaagccttggcctcagacatcggtccttacctgcaagtcatcggcatcgccggccctggagagcctctagctaatccagcgacctataccacattggaactcatccaagagcatcatccccatctcatccgctgcatcagctcgaacgggttgctgttgcctgagcggttgtccgacttgctccgctgcggtgtatcccatgtgacggttaccttaaataccttggatccggctgttggagcccagatctacaggtatgttcgctggcagggccgaaagctgaccggagaaatgggcgcccagatcctcttagaacagcagttgcttggtattgaattggctgcacaagcaggtatgaccgtcaaggtcaacaccgtcgtcatcccaggactgaacgacaaacacctggaagaactggcttgggaagttagagctcgaggagcctctattctcaacctgatgccgttgatcaaccagggaattttcgctgacgtccaaccgccgactccagaggaaatgcacacctaccggcggcaggcagaagcgattttgccgcaactgtctcactgtcgccaatgccgagctgacgccatcggcatcatatagnifn序列（seqidno.12）：atgactggtccggtagctgaacctcgcagctacaccggcaaccctcagaaaaacagccctgccttaggtgccaccatcgcctacatgggcgtcgatggtctcttagcccttttgcacggatcacaaggttgctccacctttatccgtttacagctctcgcgacacttcaaggagcccattcccttaaactcgacagcccttctcgaagatacagccatcttcggaggctgggaacacttgaaaaaaggtatcgccaaagcggctgataaatttcagccacgcatcatcggtgtcatgagttccggcttaacggaaaccttcggcgacgacatggtcagcgccttggccagcttgcgtcaggagcggcccgatttgatcgaaaaaccggtcgtgttggctaaaacccccgactacatcggttccctgcaagaagggtaccagcgcacggttctcgccttgttggatacgcttacggttgacccctctcctagggagtcaaatacggtagccttgttccccggagcccattttactccggccgatgtggaggaactcaaagacacagtttctctgtttgggttgacgcccttagtccttcccgacatctccacatctcttgacggccattttgagctagaaccggcaccggtggttcaagggggcattacgctggaagatatccggcgaattcccgggtgcatcgcctccttctcttttgggcccagcatggccgtcgtcggagaatccttgcaaaagcggcacagtgtaccccatatcgccgtaccttctctgaccggtctcgaagcggtggataccttctaccttaaactttgccaactctcaggccggcccattccggagcggttgctccggcagcggagtcgcctgttagatactttggccgattatcatgaccaactcggccaaaaacgaatcgctatcgccttagaatcagacctgctcatctctttagccaactgcctcgtcgaagtcggagcagtcattcaagtagccctagcggcgacggcactccatgaaaacgtcctccccaacttcaccttctcatctcaatacactgttgaagtgggcgatctggagcacctggaagagcgagcccctggtagtcatctgatcatcgcaaactccaatgctcgtcaagcagcgcttcctctaaaaatccctcatctgcgagccgggctgccggtctttgaccgggccgggatgtcccaaaaagtatggatcggctatcgcggtacacagcagttcttatatgattgcttaaacgccatggcgggataapvx-hisg-mf1-nifh序列（seqidno.13）：gaaaactaaaccatacaccaccaacacaaccaaacccaccacgcccaattgttacacacccgcttgaaaaagaaagtttaacaaatggccaaggtgcgcgaggtttaccaatcttttacagactccaccacaaaaactctcatccaagatgaggcttatagaaacattcgccccatcatggaaaaacacaaactagctaacccttacgctcaaacggttgaagcggctaatgatctagaggggttcggcatagccaccaatccctatagcattgaattgcatacacatgcagccgctaagaccatagagaataaacttctagaggtgcttggttccatcctaccacaagaacctgttacatttatgtttcttaaacccagaaagctaaactacatgagaagaaacccgcggatcaaggacattttccaaaatgttgccattgaaccaagagacgtagccaggtaccccaaggaaacaataattgacaaactcacagagatcacaacggaaacagcatacattagtgacactctgcacttcttggatccgagctacatagtggagacattccaaaactgcccaaaattgcaaacattgtatgcgaccttagttctccccgttgaggcagcctttaaaatggaaagcactcacccgaacatatacagcctcaaatacttcggagatggtttccagtatataccaggcaaccatggtggcggggcataccatcatgaattcgctcatctacaatggctcaaagtgggaaagatcaagtggagggaccccaaggatagctttctcggacatctcaattacacgactgagcaggttgagatgcacacagtgacagtacagttgcaggaatcgttcgcggcaaaccacttgtactgcatcaggagaggagacttgctcacaccggaggtgcgcactttcggccaacctgacaggtacgtgattccaccacagatcttcctcccaaaagttcacaactgcaagaagccgattctcaagaaaactatgatgcagctcttcttgtatgttaggacagtcaaggtcgcaaaaaattgtgacatttttgccaaagtcagacaattaattaaatcatctgacttggacaaatactctgctgtggaactggtttacttagtaagctacatggagttccttgccgatttacaagctaccacctgcttctcagacacactttctggtggcttgctaacaaagacccttgcaccggtgagggcttggatacaagagaaaaagatgcagctgtttggtcttgaggactacgcgaagttagtcaaagcagttgatttccacccggtggatttttctttcaaagtggaaacttgggacttcagattccaccccttgcaagcgtggaaagccttccgaccaagggaagtgtcggatgtagaggaaatggaaagtttgttctcagatggggacctgcttgattgcttcacaagaatgccagcttatgcggtaaacgcagaggaagatttagctgcaatcaggaaaacgcccgagatggatgtcggtcaagaagttaaagagcctgcaggagacagaaatcaatactcaaaccctgcagaaactttcctcaacaagctccacaggaaacacagtagggaggtgaaacaccaggccgcaaagaaagctaaacgcctagctgaaatccaggagtcaatgagagctgaaggtgatgccgaaccaaatgaaataagcgggacgatgggggcaatacccagcaacgccgaacttcctggcacgaatgatgccagacaagaactcacactcccaaccactaaacctgtccctgcaaggtgggaagatgcttcattcacagattctagtgtggaagaggagcaggttaaactccttggaaaagaaaccgttgaaacagcgacgcaacaagtcatcgaaggacttccttggaaacactggattcctcaattaaatgctgttggattcaaggcgctggaaattcagagggataggagtggaacaatgatcatgcccatcacagaaatggtctccgggctggaaaaagaggacttccctgaaggaactccaaaagagttggcacgagaattgttcgctatgaacagaagccctgccaccatccctttggacctgcttagagccagagactacggcagtgatgtaaagaacaagagaattggtgccatcacaaagacacaggcaacgagttggggcgaatacttgacaggaaagatagaaagcttaactgagaggaaagttgcgacttgtgtcattcatggagctggaggttctggaaaaagtcatgccatccagaaggcattgagagaaattggcaagggctcggacatcactgtagtcctgccgaccaatgaactgcggctagattggagtaagaaagtgcctaacactgagccctatatgttcaagacctctgaaaaggcgttaattgggggaacaggcagcatagtcatctttgacgattactcaaaacttcctcccggttacatagaagccttagtctgtttctactctaaaatcaagctaatcattctaacaggagatagcagacaaagcgtctaccatgaaactgctgaggacgcctccatcaggcatttgggaccagcaacagagtacttctcaaaatactgccgatactatctcaatgccacacaccgcaacaagaaagatcttgcgaacatgcttggtgtctacagtgagagaacgggagtcaccgaaatcagcatgagcgccgagttcttagaaggaatcccaactttggtaccctcggatgagaagagaaagctgtacatgggcaccgggaggaatgacacgttcacatacgctggatgccaggggctaactaagccgaaggtacaaatagtgttggaccacaacacccaagtgtgtagcgcgaatgtgatgtacacggcactttctagagccaccgataggattcacttcgtgaacacaagtgcaaattcctctgccttctgggaaaagttggacagcaccccttacctcaagactttcctatcagtggtgagagaacaagcactcagggagtacgagccggcaga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gtgtaggtcgttcgctccaagctgggctgtgtgcacgaaccccccgttcagcccgaccgctgcgccttatccggtaactatcgtcttgagtccaacccggtaagacacgacttatcgccactggcagcagccactggtaacaggattagcagagcgaggtatgtaggcggtgctacagagttcttgaagtggtggcctaactacggctacactagaaggacagtatttggtatctgcgctctgctgaagccagttaccttcggaaaaagagttggtagctcttgatccggcaaacaaaccaccgctggtagcggtggtttttttgtttgcaagcagcagattacgcgcagaaaaaaaggatctcaagaagatcctttgatcttttctacggggtctgacgctcagtggaacgaaaactcacgttaagggattttggtcatgagattatcaaaaaggatcttcacctagatccttttaaattaaaaatgaagttttaaatcaatctaaagtatatatgagtaaacttggtctgacagttaccaatgcttaatcagtgaggcacctatctcagcgatctgtctatttcgttcatccatagttgcctgactccccgtcgtgtagataactacgatacgggagggcttaccatctggccccagtgctgcaatgataccgcgagacccacgctcaccggctccagatttatcagcaataaaccagccagccggaagggccgagcgcagaagtggtcctgcaactttatccgcctccatccagtctattaattgttgccgggaagctagagtaagtagttcgccagttaatagtttgcgcaacgttgttgccattgctacaggcatcgtggtgtcacgctcgtcgtttggtatggcttcattcagctccggttcccaacgatcaaggcgagttacatgatcccccatgttgtgcaaaaaagcggttagctccttcggtcctccgatcgttgtcagaagtaagttggccgcagtgttatcactcatggttatggcagcactgcataattctcttactgtcatgccatccgtaagatgcttttctgtgactggtgagtactcaaccaagtcattctgagaatagtgtatgcggcgaccgagttgctcttgcccggcgtcaatacgggataataccgcgccacatagcagaactttaaaagtgctcatcattggaaaacgttcttcggggcgaaaactctcaaggatcttaccgctgttgagatccagttcgatgtaacccactcgtgcacccaactgatcttcagcatcttttactttcaccagcgtttctgggtgagcaaaaacaggaaggcaaaatgccgcaaaaaagggaataagggcgacacggaaatgttgaatactcatactcttcctttttcaatattattgaagcatttatcagggttattgtctcatgagcggatacatatttgaatgtatttagaaaaataaacaaataggggttccgcgcacatttccccgaaaagtgccacctgacgtctaagaaaccattattatcatgacattaacctataaaaataggcgtatcacgaggccctttcgtctcgcgcgtttcggtgatgacggtgaaaacctctgacacatgcagctcccggagacggtcacagcttgtctgtaagcggatgccgggagcagacaagcccgtcagggcgcgtcagcgggtgttggcgggtgtcggggctggcttaactatgcggcatcagagcagattgtactgagagtgcaccatatgcggtgtgaaataccgcacagatgcgtaaggagaaaataccgcatcaggaaattgtaaacgttaatattttgttaaaattcgcgttaaatttttgttaaatcagctcattttttaaccaataggccgaaatcggcaaaatcccttataaatcaaaagaatagaccgagatagggttgagtgttgttccagtttggaacaagagtccactattaaagaacgtggactccaacgtcaaagggcgaaaaaccgtctatcagggcgatggcccactacgtgaaccatcaccctaatcaagttttttggggtcgaggtgccgtaaagcactaaatcggaaccctaaagggagcccccgatttagagcttgacggggaaagccggcgaacgtggcgagaaaggaagggaagaaagcgaaaggagcgggcgctagggcgctggcaagtgtagcggtcacgctgcgcgtaaccaccacacccgccgcgcttaatgcgccgctacagggcgcgtcgcgccattcgccattcaggctgcgcaactgttgggaagggcgatcggtgcgggcctcttcgctattacgccagctggcgaaagggggatgtgctgcaaggcgattaagttgggtaacgccagggttttcccagtcacgacgttgtaaaacgacggccagtgaattgtaatacgactcactatapvx-hisg-mf1-nife序列（seqidno.14）：gaaaactaaaccatacaccaccaacacaaccaaacccaccacgcccaattgttacacacccgcttgaaaaagaaagtttaacaaatggccaaggtgcgcgaggtttaccaatcttttacagactccaccacaaaaactctcatccaagatgaggcttatagaaacattcgccccatcatggaaaaacacaaactagctaacccttacgctcaaacggttgaagcggctaatgatctagaggggttcggcatagccaccaatccctatagcattgaattgcatacacatgcagccgctaagaccatagagaataaacttctagaggtgcttggttccatcctaccacaagaacctgttacatttatgtttcttaaacccagaaagctaaactacatgagaagaaacccgcggatcaaggacattttccaaaatgttgccattgaaccaagagacgtagccaggtaccccaaggaaacaataattgacaaactcacagagatcacaacggaaacagcatacattagtgacactctgcacttcttggatccgagctacatagtggagacattccaaaactgcccaaaattgcaaacattgtatgcgaccttagttctccccgttgaggcagcctttaaaatggaaagcactcacccgaacatatacagcctcaaatacttcggagatggtttccagtatataccaggcaaccatggtggcggggcataccatcatgaattcgctcatctacaatggctcaaagtgggaaagatcaagtggagggaccccaaggatagctttctcggacatctcaattacacgactgagcaggttgagatgcacacagtgacagtacagttgcaggaatcgttcgcggcaaaccacttgtactgcatcaggagaggagacttgctcacaccggaggtgcgcactttcggccaacctgacaggtacgtgattccaccacagatcttcctcccaaaagttcacaactgcaagaagccgattctcaagaaaactatgatgcagctcttcttgtatgttaggacagtcaaggtcgcaaaaaattgtgacatttttgccaaagtcagacaattaattaaatcatctgacttggacaaatactctgctgtggaactggtttacttagtaagctacatggagttccttgccgatttacaagctaccacctgcttctcagacacactttctggtggcttgctaacaaagacccttgcaccggtgagggcttggatacaagagaaaaagatgcagctgtttggtcttgaggactacgcgaagttagtcaaagcagttgatttccacccggtggatttttctttcaaagtggaaacttgggacttcagattccaccccttgcaagcgtggaaagccttccgaccaagggaagtgtcggatgtagaggaaatggaaagtttgttctcagatggggacctgcttgattgcttcacaagaatgccagcttatgcggtaaacgcagaggaagatttagctgcaatcaggaaaacgcccgagatggatgtcggtcaagaagttaaagagcctgcaggagacagaaatcaatactcaaaccctgcagaaactttcctcaacaagctccacaggaaacacagtagggaggtgaaacaccaggccgcaaagaaagctaaacgcctagctgaaatccaggagtcaatgagagctgaaggtgatgccgaaccaaatgaaataagcgggacgatgggggcaatacccagcaacgccgaacttcctggcacgaatgatgccagacaagaactcacactcccaaccactaaacctgtccctgcaaggtgggaagatgcttcattcacagattctagtgtggaagaggagcaggttaaactccttggaaaagaaaccgttgaaacagcgacgcaacaagtcatcgaaggacttccttggaaacactggattcctcaattaaatgctgttggattcaaggcgctggaaattcagagggataggagtggaacaatgatcatgcccatcacagaaatggtctccgggctggaaaaagaggacttccctgaaggaactccaaaagagttggcacgagaattgttcgctatgaacagaagccctgccaccatccctttggacctgcttagagccagagactacggcagtgatgtaaagaacaagagaattggtgccatcacaaagacacaggcaacgagttggggcgaatacttgacaggaaagatagaaagcttaactgagaggaaagttgcgacttgtgtcattcatggagctggaggttctggaaaaagtcatgccatccagaaggcattgagagaaattggcaagggctcggacatcactgtagtcctgccgaccaatgaactgcggctagattggagtaagaaagtgcctaacactgagccctatatgttcaagacctctgaaaaggcgttaattgggggaacaggcagcatagtcatctttgacgattactc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taccttcggaaaaagagttggtagctcttgatccggcaaacaaaccaccgctggtagcggtggtttttttgtttgcaagcagcagattacgcgcagaaaaaaaggatctcaagaagatcctttgatcttttctacggggtctgacgctcagtggaacgaaaactcacgttaagggattttggtcatgagattatcaaaaaggatcttcacctagatccttttaaattaaaaatgaagttttaaatcaatctaaagtatatatgagtaaacttggtctgacagttaccaatgcttaatcagtgaggcacctatctcagcgatctgtctatttcgttcatccatagttgcctgactccccgtcgtgtagataactacgatacgggagggcttaccatctggccccagtgctgcaatgataccgcgagacccacgctcaccggctccagatttatcagcaataaaccagccagccggaagggccgagcgcagaagtggtcctgcaactttatccgcctccatccagtctattaattgttgccgggaagctagagtaagtagttcgccagttaatagtttgcgcaacgttgttgccattgctacaggcatcgtggtgtcacgctcgtcgtttggtatggcttcattcagctccggttcccaacgatcaaggcgagttacatgatcccccatgttgtgcaaaaaagcggttagctccttcggtcctccgatcgttgtcagaagtaagttggccgcagtgttatcactcatggttatggcagcactgcataattctcttactgtcatgccatccgtaagatgcttttctgtgactggtgagtactcaaccaagtcattctgagaatagtgtatgcggcgaccgagttgctcttgcccggcgtcaatacgggataataccgcgccacatagcagaactttaaaagtgctcatcattggaaaacgttcttcggggcgaaaactctcaaggatcttaccgctgttgagatccagttcgatgtaacccactcgtgcacccaactgatcttcagcatcttttactttcaccagcgtttctgggtgagcaaaaacaggaaggcaaaatgccgcaaaaaagggaataagggcgacacggaaatgttgaatactcatactcttcctttttcaatattattgaagcatttatcagggttattgtctcatgagcggatacatatttgaatgtatttagaaaaataaacaaataggggttccgcgcacatttccccgaaaagtgccacctgacgtctaagaaaccattattatcatgacattaacctataaaaataggcgtatcacgaggccctttcgtctcgcgcgtttcggtgatgacggtgaaaacctctgacacatgcagctcccggagacggtcacagcttgtctgtaagcggatgccgggagcagacaagcccgtcagggcgcgtcagcgggtgttggcgggtgtcggggctggcttaactatgcggcatcagagcagattgtactgagagtgcaccatatgcggtgtgaaataccgcacagatgcgtaaggagaaaataccgcatcaggaaattgtaaacgttaatattttgttaaaattcgcgttaaatttttgttaaatcagctcattttttaaccaataggccgaaatcggcaaaatcccttataaatcaaaagaatagaccgagatagggttgagtgttgttccagtttggaacaagagtccactattaaagaacgtggactccaacgtcaaagggcgaaaaaccgtctatcagggcgatggcccactacgtgaaccatcaccctaatcaagttttttggggtcgaggtgccgtaaagcactaaatcggaaccctaaagggagcccccgatttagagcttgacggggaaagccggcgaacgtggcgagaaaggaagggaagaaagcgaaaggagcgggcgctagggcgctggcaagtgtagcggtcacgctgcgcgtaaccaccacacccgccgcgcttaatgcgccgctacagggcgcgtcgcgccattcgccattcaggctgcgcaactgttgggaagggcgatcggtgcgggcctcttcgctattacgccagctggcgaaagggggatgtgctgcaaggcgattaagttgggtaacgccagggttttcccagtcacgacgttgtaaaacgacggccagtgaattgtaatacgactcactatapvx-hisg-mf1-nifn序列（seqidno.15）：gaaaactaaaccatacaccaccaacacaaccaaacccaccacgcccaattgttacacacccgcttgaaaaagaaagtttaacaaatggccaaggtgcgcgaggtttaccaatcttttacagactccaccacaaaaactctcatccaagatgaggcttatagaaacattcgccccatcatggaaaaacacaaactagctaacccttacgctcaaacggttgaagcggctaatgatctagaggggttcggcatagccaccaatccctatagcattgaattgcatacacatgcagccgctaagaccatagagaataaacttctagaggtgcttggttccatcctaccacaagaacctgttacatttatgtttcttaaacccagaaagctaaactacatgagaagaaacccgcggatcaaggacattttccaaaatgttgccattgaaccaagagacgtagccaggtaccccaaggaaacaataattgacaaactcacagagatcacaacggaaacagcatacattagtgacactctgcacttcttggatccgagctacatagtggagacattccaaaactgcccaaaattgcaaacattgtatgcgaccttagttctccccgttgaggcagcctttaaaatggaaagcactcacccgaacatatacagcctcaaatacttcggagatggtttccagtatataccaggcaaccatggtggcggggcataccatcatgaattcgctcatctacaatggctcaaagtgggaaagatcaagtggagggaccccaaggatagctttctcggacatctcaattacacgactgagcaggttgagatgcacacagtgacagtacagttgcaggaatcgttcgcggcaaaccacttgtactgcatcaggagaggagacttgctcacaccggaggtgcgcactttcggccaacctgacaggtacgtgattccaccacagatcttcctcccaaaagttcacaactgcaagaagccgattctcaagaaaactatgatgcagctcttcttgtatgttaggacagtcaaggtcgcaaaaaattgtgacatttttgccaaagtcagacaattaattaaatcatctgacttggacaaatactctgctgtggaactggtttacttagtaagctacatggagttccttgccgatttacaagctaccacctgcttctcagacacactttctggtggcttgctaacaaagacccttgcaccggtgagggcttggatacaagagaaaaagatgcagctgtttggtcttgaggactacgcgaagttagtcaaagcagttgatttccacccggtggatttttctttcaaagtggaaacttgggacttcagattccaccccttgcaagcgtggaaagccttccgaccaagggaagtgtcggatgtagaggaaatggaaagtttgttctcagatggggacctgcttgattgcttcacaagaatgccagcttatgcggtaaacgcagaggaagatttagctgcaatcaggaaaacgcccgagatggatgtcggtcaagaagttaaagagcctgcaggagacagaaatcaatactcaaaccctgcagaaactttcctcaacaagctccacaggaaacacagtagggaggtgaaacaccaggccgcaaagaaagctaaacgcctagctgaaatccaggagtcaatgagagctgaaggtgatgccgaaccaaatgaaataagcgggacgatgggggcaatacccagcaacgccgaacttcctggcacgaatgatgccagacaagaactcacactcccaaccactaaacctgtccctgcaaggtgggaagatgcttcattcacagattctagtgtggaagaggagcaggttaaactccttggaaaagaaaccgttgaaacagcgacgcaacaagtcatcgaaggacttccttggaaacactggattcctcaattaaatgctgttggattcaaggcgctggaaattcagagggataggagtggaacaatgatcatgcccatcacagaaatggtctccgggctggaaaaagaggacttccctgaaggaactccaaaagagttggcacgagaattgttcgctatgaacagaagccctgccaccatccctttggacctgcttagagccagagactacggcagtgatgtaaagaacaagagaattggtgccatcacaaagacacaggcaacgagttggggcgaatacttgacaggaaagatagaaagcttaactgagaggaaagttgcgacttgtgtcattcatggagctggaggttctggaaaaagtcatgccatccagaaggcattgagagaaattggcaagggctcggacatcactgtagtcctgccgaccaatgaactgcggctagattggagtaagaaagtgcctaacactgagccctatatgttcaagacctctgaaaaggcgttaattgggggaacaggcagcatagtcatctttgacgattactcaaaacttcctcccggttacatagaagccttagtctgtttctactctaaaatcaagctaatcattctaacaggagatagcagacaaagcgtctaccatgaaactgctgaggacgcctccatcaggcatttgggaccagcaacagagtacttctcaaaatactgccgatactatctcaatgccacacaccgcaacaagaaagatcttgcgaacatgcttggtgtctacagtgagagaacgggagtcaccgaaatca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ggcaccaaagcaatcttgtacaactccccaaatctagggtcacgagtgagtctacacaacggaaagaacgcagcatttgctgccgttttgctactgactttgctgatctatggaagtaaatacatatctcaacgcaatcatacttgtgcttgtggtaacaatcatagcagtcattagcacttccttagtgaggactgaaccttgtgtcatcaagattactggggaatcaatcacagtgttggcttgcaaactagatgcagaaaccataagggccattgccgatctcaagccactctccgttgaacggttaagtttccattgatactcgaaagaggtcagcggccgggttgcactggttcttgcggaggaaattcaacgaaagacgtatccaagcacccctgtttttcacctcatggccacggaaaatcaggaaggattcatctgcctgtcgctcctggatgtaacatcgcctgtggttattgtgtgcgtaaattcgattgtgccaatgagtcacggcctggcgttaccagccgagtgatcactcccgaacaggccctctggcgcgtagaacaagccttggcctcagacatcggtccttacctgcaagtcatcggcatcgccggccctggagagcctctagctaatccagcgacctataccacattggaactcatccaagagcatcatccccatctcatccgctgcatcagctcgaacgggttgctgttgcctgagcggttgtccgacttgctccgctgcggtgtatcccatgtgacggttaccttaaataccttggatccggctgttggagcccagatctacaggtatgttcgctggcagggccgaaagctgaccggagaaatgggcgcccagatcctcttagaacagcagttgcttggtattgaattggctgcacaagcaggtatgaccgtcaaggtcaacaccgtcgtcatcccaggactgaacgacaaacacctggaagaactggcttgggaagttagagctcgaggagcctctattctcaacctgatgccgttgatcaaccagggaattttcgctgacgtccaaccgccgactccagaggaaatgcacacctaccggcggcaggcagaagcgattttgccgcaactgtctcactgtcgccaatgccgagctgacgccatcggcatcatataggtcgaccgccgatgaacggttaagtttccattgatactcgaaagatgtcagcaccagctagcacaacacagcccatagggtcaactacctcaactaccacaaaaactgcaggcgcaactcctgccacagcttcaggcctgttcaccatcccggatggggatttctttagtacagcccgtgccatagtagccagcaatgctgtcgcaacaaatgaggacctcagcaagattgaggctatttggaaggacatgaaggtgcccacagacactatggcacaggctgcttgggacttagtcagacactgtgctgatgtaggatcatccgctcaaacagaaatgatagatacaggtccctattccaacggcatcagcagagctagactggcagcagcaattaaagaggtgtgcacacttaggcaattttgcatgaagtatgctccagtggtatggaactggatgttaactaacaacagtccacctgctaactggcaagcacaaggtttcaagcctgagcacaaattcgctgcattcgacttcttcaatggagtcaccaacccagctgccatcatgcccaaagaggggctcatccggccaccgtctgaagctgaaatgaatgctgcccaaactgctgcctttgtgaagattacaaaggccagggcacaatccaacgactttgccagcctagatgcagctgtcactcgaggtcgtatcactggaacaacaaccgctgaggctgttgtcactctaccaccaccataactacgtctacataaccgacgcctaccccagtttcatagtattttctggtttgattgtatgaataatataaataaaaaaaaaaaaaaaaaaaaaaaactagtggtaccgagctcggtaccccatgcaagcttttgttccctttagtgagggttaattccgagcttggcgtaatcatggtcatagctgtttcctgtgtgaaattgttatccgctcacaattccacacaacatacgagccggaagcataaagtgtaaagcctggggtgcctaatgagtgagctaactcacattaattgcgttgcgctcactgcccgctttccagtcgggaaacctgtcgtgccagctgcattaatgaatcggccaacgcgcggggagaggcggtttgcgtattgggcgctcttccgcttcctcgctcactgactcgctgcgctcggtcgttcggctgcggcgagcggtatcagctcactcaaaggcggtaatacggttatccacagaatcaggggataacgcaggaaagaacatgtgagcaaaaggccagcaaaaggccaggaaccgtaaaaaggccgcgttgctggcgtttttccataggctccgcccccctgacgagcatcacaaaaatcgacgctcaagtcagaggtggcgaaacccgacaggactataaagataccaggcgtttccccctggaagctccctcgtgcgctctcctgttccgaccctgccgcttaccggatacctgtccgcctttctcccttcgggaagcgtggcgctttctcatagctcacgctgtaggtatctcagttcggtgtaggtcgttcgctccaagctgggctgtgtgcacgaaccccccgttcagcccgaccgctgcgccttatccggtaactatcgtcttgagtccaacccggtaagacacgacttatcgccactggcagcagccactggtaacaggattagcagagcgaggtatgtaggcggtgctacagagttcttgaagtggtggcctaactacggctacactagaaggacagtatttggtatctgcgctctgctgaagccagttaccttcggaaaaagagttggtagctcttgatccggcaaacaaaccaccgctggtagcggtggtttttttgtttgcaagcagcagattacgcgcagaaaaaaaggatctcaagaagatcctttgatcttttctacggggtctgacgctcagtggaacgaaaactcacgttaagggattttggtcatgagattatcaaaaaggatcttcacctagatccttttaaattaaaaatgaagttttaaatcaatctaaagtatatatgagtaaacttggtctgacagttaccaatgcttaatcagtgaggcacctatctcagcgatctgtctatttcgttcatccatagttgcctgactccccgtcgtgtagataactacgatacgggagggcttaccatctggccccagtgctgcaatgataccgcgagacccacgctcaccggctccagatttatcagcaataaaccagccagccggaagggccgagcgcagaagtggtcctgcaactttatccgcctccatccagtctattaattgttgccgggaagctagagtaagtagttcgccagttaatagtttgcgcaacgttgttgccattgctacaggcatcgtggtgtcacgctcgtcgtttggtatggcttcattcagctccggttcccaacgatcaaggcgagttacatgatcccccatgttgtgcaaaaaagcggttagctccttcggtcctccgatcgttgtcagaagtaagttggccgcagtgttatcactcatggttatggcagcactgcataattctcttactgtcatgccatccgtaagatgcttttctgtgactggtgagtactcaaccaagtcattctgagaatagtgtatgcggcgaccgagttgctcttgcccggcgtcaatacgggataataccgcgccacatagcagaactttaaaagtgctcatcattggaaaacgttcttcggggcgaaaactctcaaggatcttaccgctgttgagatccagttcgatgtaacccactcgtgcacccaactgatcttcagcatcttttactttcaccagcgtttctgggtgagcaaaaacaggaaggcaaaatgccgcaaaaaagggaataagggcgacacggaaatgttgaatactcatactcttcctttttcaatattattgaagcatttatcagggttattgtctcatgagcggatacatatttgaatgtatttagaaaaataaacaaataggggttccgcgcacatttccccgaaaagtgccacctgacgtctaagaaaccattattatcatgacattaacctataaaaataggcgtatcacgaggccctttcgtctcgcgcgtttcggtgatgacggtgaaaacctctgacacatgcagctcccggagacggtcacagcttgtctgtaagcggatgccgggagcagacaagcccgtcagggcgcgtcagcgggtgttggcgggtgtcggggctggcttaactatgcggcatcagagcagattgtactgagagtgcaccatatgcggtgtgaaataccgcacagatgcgtaaggagaaaataccgcatcaggaaattgtaaacgttaatattttgttaaaattcgcgttaaatttttgttaaatcagctcattttttaaccaataggccgaaatcggcaaaatcccttataaatcaaaagaatagaccgagatagggttgagtgttgttccagtttggaacaagagtccactattaaagaacgtggactccaacgtcaaagggcgaaaaaccgtctatcagggcgatggcccactacgtgaaccatcaccctaatcaagttttttggggtcgaggtgccgtaaagcactaaatcggaaccctaaagggagcccccgatttagagcttgacggggaaagccggcgaacgtggcgagaaaggaagggaagaaagcgaaaggagcgggcgctagggcgctggcaagtgtagcggtcacgctgcgcgtaaccaccacacccgccgcgcttaatgcgccgctacagggcgcgtcgcgccattcgccattcaggctgcgcaactgttgggaagggcgatcggtgcgggcctcttcgctattacgccagctggcgaaagggggatgtgctgcaaggcgattaagttgggtaacgccagggttttcccagtcacgacgttgtaaaacgacggccagtgaattgtaatacgactcactata本行業(yè)的技術(shù)人員應(yīng)該了解，本發(fā)明不受上述實施例的限制，上述實施例和說明書中描述的只是說明本發(fā)明的原理，在不脫離本發(fā)明精神和范圍的前提下，本發(fā)明還會有各種變化和改進，這些變化和改進都落入要求保護的本發(fā)明范圍內(nèi)。本發(fā)明要求保護范圍由所附的權(quán)利要求書及其等效物界定。引物序列seqidno.nifhncacggaatcgatcgtcagatagccatttacggaaaaggtg1nifhcgtctcagtcgacgattagcccagtttggccatc2nifenacgcgtcggccgtttgtcaatcttaccaaactggacttaaaag3nifecctgtcagtcgaccgttggtgctgccatataggttgattg4nifnncacggaatcgatactggtccggtagctgaacctcgcag5nifncgactcagtcgacttaagtccttatcccgccatggcgtttaag6nifbnacgcgtcggccgggttgcactggttcttgcggaggaaattc7nifbcgtctcagtcgacctatatgatgccgatggcgtcagctc8nifh序列（seqidno.9）：atgcgtcagatagccatttacggaaaaggtggtatcggtaaatctacgaccacccaaaacaccgtctccgccctcgccgagatgggtaaaaaaatcatgatcgtcggttgtgaccccaaagccgactctactcgtttgattcttcactccaaagcccaagcgacggttatggacctggcccgtgaaaaaggtactgtggaagacctcgaactggaagacgttctcctcaccggtttcgccgacatccgctgtgccgagtccggcggtcccgaacctggtgtaggctgtgccggccgcggtgttatcaccgccatcaacttcctcgaagaaaacggcgcttacacccctgacctggactatgtgttctacgacgttctcggtgacgttgtctgcggcggtttcgccatgcctatccgcgaaaacaaagcccaagaaatctacatcgtgacctccggtgaaatgatggccatgtacgctgccaacaacatcgcccgcggtatccttaaatacgctgctaccggtaaagttcgcctcggcggcctcatctgtaacagccgtaaaaccgacaaggaatacgaactgattgccgagttggccagccgtctgggcacgcaaatgattcacttcctgccccgtgacaaccaagtgcaacgggccgagctgcgccgcatgaccgttatcgaatactctcccgaacatccccaagccgacgaatatcgcgctctggccaagaaaatcgacgagaacaaaatgctcgttgttcccacaccgctttccatggacgaactggaagacctcctcatctcctacggcatcctggaagacgaagaaacggcgatggccaaactgggctaanife序列（seqidno.10）：gtgtttgtcaatcttaccaaactggacttaaaagaaacatgtccccttactgataaaggggcgccgaaactgtgtatgaaagctctgcccggtgaaggtgctgagcgttcctgtgcattcgacggcgcaagagtcgtcctcatgccggtgaccgatgccgcccacgttgtccatggtcccatcgcttgcgcggggaactcctgggacaaccggggcgccaagtcttcgggaccccaactctatcgccgcggattgacgacagacctgatggaaatggatgtcgtctacggtggagagaagaaactcattcaagccatcatcgatgtggcccggaaacaccagcccaaagcggtattcgtctatgccacctgtgtctcggctttaatcggagacgatctggaaacggcctgtaaggaagcgcaaaaagacgtctcagtcccggtcatcccggtaaacgctcctggttttatgggcgataaaaacatcggcaaccgtgtcgccggcgaagtcctcttccgccagatcatcggcacggctgaaccagaaacagtccaccctttcgccatcaacttcatcggtgaatacaacatcgccggcgacttgtgggggatccagccctccttagaggctatcggtattaagctgcaagcgtgcatcagcggcgatgctcgttttgatgacatccgtacggcacaccgagccaaactgaatgtgctggtttgctcgaagagcttgaccaacctggtcaaacagatggaacaaaaatatggaattccatacatcgaaggatccttctatggcatccacgatacaagtgctacactccgagccatcgccaaagctttcggagacgccggcctgatcgaacggacggaagcccatattctccagaaggaagcggaaacgcgagccgtcatcgagccccttaaggcgcaactcagcggcaagagagccgtcctcttcacgggtggcgtgaaaacctggtccatggtcacgaccctgcgggaattagggatcgacgtactggctggtggtacgcagaactccacaccggatgacttcttgcggatgaaagccttgatggacccgacggctcatatcatcgaagataccagtgccgccggatttctaaccctgatcaaagaaaaaagccccgacctcatcgtggccggaggcaaaacgaagtacctggcccataagactcggactccttttctggatatcaaccacggacgcagccttccctatgccggctacgccgggatgatcacctttgccaacgccgttgcccgtactgtattcagccccgtctgggaaaaaatgagggtagcttttccgccggaggtgacggtcaatgagtaacgcaatcaacctatatggcagcaccaacgnifb序列（seqidno.11）：atgggttgcactggttcttgcggaggaaattcaacgaaagacgtatccaagcacccctgtttttcacctcatggccacggaaaatcaggaaggattcatctgcctgtcgctcctggatgtaacatcgcctgtggttattgtgtgcgtaaattcgattgtgccaatgagtcacggcctggcgttaccagccgagtgatcactcccgaacaggccctctggcgcgtagaacaagccttggcctcagacatcggtccttacctgcaagtcatcggcatcgccggccctggagagcctctagctaatccagcgacctataccacattggaactcatccaagagcatcatccccatctcatccgctgcatcagctcgaacgggttgctgttgcctgagcggttgtccgacttgctccgctgcggtgtatcccatgtgacggttaccttaaataccttggatccggctgttggagcccagatctacaggtatgttcgctggcagggccgaaagctgaccggagaaatgggcgcccagatcctcttagaacagcagttgcttggtattgaattggctgcacaagcaggtatgaccgtcaaggtcaacaccgtcgtcatcccaggactgaacgacaaacacctggaagaactggcttgggaagttagagctcgaggagcctctattctcaacctgatgccgttgatcaaccagggaattttcgctgacgtccaaccgccgactccagaggaaatgcacacctaccggcggcaggcagaagcgattttgccgcaactgtctcactgtcgccaatgccgagctgacgccatcggcatcatatagnifn序列（seqidno.12）：atgactggtccggtagctgaacctcgcagctacaccggcaaccctcagaaaaacagccctgccttaggtgccaccatcgcctacatgggcgtcgatggtctcttagcccttttgcacggatcacaaggttgctccacctttatccgtttacagctctcgcgacacttcaaggagcccattcccttaaactcgacagcccttctcgaagatacagccatcttcggaggctgggaacacttgaaaaaaggtatcgccaaagcggctgataaatttcagccacgcatcatcggtgtcatgagttccggcttaacggaaaccttcggcgacgacatggtcagcgccttggccagcttgcgtcaggagcggcccgatttgatcgaaaaaccggtcgtgttggctaaaacccccgactacatcggttccctgcaagaagggtaccagcgcacggttctcgccttgttggatacgcttacggttgacccctctcctagggagtcaaatacggtagccttgttccccggagcccattttactccggccgatgtggaggaactcaaagacacagtttctctgtttgggttgacgcccttagtccttcccgacatctccacatctcttgacggccattttgagctagaaccggcaccggtggttcaagggggcattacgctggaagatatccggcgaattcccgggtgcatcgcctccttctcttttgggcccagcatggccgtcgtcggagaatccttgcaaaagcggcacagtgtaccccatatcgccgtaccttctctgaccggtctcgaagcggtggataccttctaccttaaactttgccaactctcaggccggcccattccggagcggttgctccggcagcggagtcgcctgttagatactttggccgattatcatgaccaactcggccaaaaacgaatcgctatcgccttagaatcagacctgctcatctctttagccaactgcctcgtcgaagtcggagcagtcattcaagtagccctagcggcgacggcactccatgaaaacgtcctccccaacttcaccttctcatctcaatacactgttgaagtgggcgatctggagcacctggaagagcgagcccctggtagtcatctgatcatcgcaaactccaatgctcgtcaagcagcgcttcctctaaaaatccctcatctgcgagccgggctgccggtctttgaccgggccgggatgtcccaaaaagtatggatcggctatcgcggtacacagcagttcttatatgattgcttaaacgccatggcgggataapvx-hisg-mf1-nifh序列（seqidno.13）：gaaaactaaaccatacaccaccaacacaaccaaacccaccacgcccaattgttacacacccgcttgaaaaagaaagtttaacaaatggccaaggtgcgcgaggtttaccaatcttttacagactccaccacaaaaactctcatccaagatgaggcttatagaaacattcgccccatcatggaaaaacacaaactagctaacccttacgctcaaacggttgaagcggctaatgatctagaggggttcggcatagccaccaatccctatagcattgaattgcatacacatgcagccgctaagaccatagagaataaacttctagaggtgcttggttccatcctaccacaagaacctgttacatttatgtttcttaaacccagaaagctaaactacatgagaagaaacccgcggatcaaggacattttccaaaatgttgccattgaaccaagagacgtagccaggtaccccaaggaaacaataattgacaaactcacagagatcacaacggaaacagcatacattagtgacactctgcacttcttggatccgagctacatagtggagacattccaaaactgcccaaaattgcaaacattgtatgcgaccttagttctccccgttgaggcagcctttaaaatggaaagcactcacccgaacatatacagcctcaaatacttcggagatggtttccagtatataccaggcaaccatggtggcggggcataccatcatgaattcgctcatctacaatggctcaaagtgggaaagatcaagtggagggaccccaaggatagctttctcggacatctcaattacacgactgagcaggttgagatgcacacagtgacagtacagttgcaggaatcgttcgcggcaaaccacttgtactgcatcaggagaggagacttgctcacaccggaggtgcgcactttcggccaacctgacaggtacgtgattccaccacagatcttcctcccaaaagttcacaactgcaagaagccgattctcaagaaaactatgatgcagctcttcttgtatgttaggacagtcaaggtcgcaaaaaattgtgacatttttgccaaagtcagacaattaattaaatcatctgacttggacaaatactctgctgtggaactggtttacttagtaagctacatggagttccttgccgatttacaagctaccacctgcttctcagacacactttctggtggcttgctaacaaagacccttgcaccggtgagggcttggatacaagagaaaaagatgcagctgtttggtcttgaggactacgcgaagttagtcaaagcagttgatttccacccggtggatttttctttcaaagtggaaacttgggacttcagattccaccccttgcaagcgtggaaagccttccgaccaagggaagtgtcggatgtagaggaaatggaaagtttgttctcagatggggacctgcttgattgcttcacaagaatgccagcttatgcggtaaacgcagaggaagatttagctgcaatcaggaaaacgcccgagatggatgtcggtcaagaagttaaagagcctgcaggagacagaaatcaatactcaaaccctgcagaaactttcctcaacaagctccacaggaaacacagtagggaggtgaaacaccaggccgcaaagaaagctaaacgcctagctgaaatccaggagtcaatgagagctgaaggtgatgccgaaccaaatgaaataagcgggacgatgggggcaatacccagcaacgccgaacttcctggcacgaatgatgccagacaagaactcacactcccaaccactaaacctgtccctgcaaggtgggaagatgcttcattcacagattctagtgtggaagaggagcaggttaaactccttggaaaagaaaccgttgaaacagcgacgcaacaagtcatcgaaggacttccttggaaacactggattcctcaattaaatgctgttggattcaaggcgctggaaattcagagggataggagtggaacaatgatcatgcccatcacagaaatggtctccgggctggaaaaagaggacttccctgaaggaactccaaaagagttggcacgagaattgttcgctatgaacagaagccctgccaccatccctttggacctgcttagagccagagactacggcagtgatgtaaagaacaagagaattggtgccatcacaaagacacaggcaacgagttggggcgaatacttgacaggaaagatagaaagcttaactgagaggaaagttgcgacttgtgtcattcatggagctggaggttctggaaaaagtcatgccatccagaaggcattgagagaaattggcaagggctcggacatcactgtagtcctgccgaccaatgaactgcggctagattggagtaagaaagtgcctaacactgagccctatatgttcaagacctctgaaaaggcgttaattgggggaacaggcagcatagtcatctttgacgattactcaaaacttcctcccggttacatagaagccttagtctgtttctactctaaaatcaagctaatcattctaacaggagatagcagacaaagcgtctaccatgaaactgctgaggacgcctccatcaggcatttgggaccagcaacagagtacttctcaaaatactgccgatactatctcaatgccacacaccgcaacaagaaagatcttgcgaacatgcttggtgtctacagtgagagaacgggagtcaccgaaatcagcatgagcgccgagttcttagaaggaatcccaactttggtaccctcggatgagaagagaaagctgtacatgggcaccgggaggaatgacacgttcacatacgctggatgccaggggctaactaagccgaaggtacaaatagtgttggaccacaacacccaagtgtgtagcgcgaatgtgatgtacacggcactttctagagccaccgataggattcacttcgtgaacacaagtgcaaattcctctgccttctgggaaaagttggacagcaccccttacctcaagactttcctatcagtggtgagagaacaagcactcagggagtacgagccggcagaggcagagccaattcaagagcctgagccccagacacacatgtgtgtcgagaatgaggagtccgtgctagaagagtacaaagaggaactcttggaaaagtttgacagagagatccactctgaatcccatggtcattcaaactgtgtccaaactgaagacacaaccattcagttgttttcgcatcaacaagcaaaagatgagaccctcctctgggcgactatagatgcgcggctcaagaccagcaatcaagaaacaaacttccgagaattcctgagcaagaaggacattggggacgttctgtttttaaactaccaaaaagctatgggtttacccaaagagcgtattcctttttcccaagaggtctgggaagcttgtgcccacgaagtacaaagcaagtacctcagcaagtcaaagtgcaacttgatcaatgggactgtgagacagagcccagacttcgatgaaaataagattatggtattcctcaagtcgcagtgggtcacaaaggtggaaaaactaggtctacccaagattaagccaggtcaaaccatagcagccttttaccagcagactgtgatgctttttggaactatggctaggtacatgcgatggttcagacaggctttccagccaaaagaagtcttcataaactgtgagaccacgccagatgacatgtctgcatgggccttgaacaactggaatttcagcagacctagcttggctaatgactacacagctttcgaccagtctcaggatggagccatgttgcaatttgaggtgctcaaagccaaacaccactgcataccagaggaaatcattcaggcatacatagatattaagactaatgcacagattttcctaggcacgttatcaattatgcgcctgactggtgaaggtcccacttttgatgcaaacactgagtgcaacatagcttacacccatacaaagtttgacatcccagccggaactgctcaagtttatgcaggagacgactccgcactggactgtgttccagaagtgaagcatagtttccacaggcttgaggacaaattactcctaaagtcaaagcctgtaatcacgcagcaaaagaagggcagttggcctgagttttgtggttggctgatcacaccaaaaggggtgatgaaagacccaattaagctccatgttagcttaaaattggctgaagctaagggtgaactcaagaaatgtcaagattcctatgaaattgatctgagttatgcctatgaccacaaggactctctgcatgacttgttcgatgagaaacagtgtcaggcacacacactcacttgcagaacactaatcaagtcagggagaggcactgtctcactttcccgcctcagaaactttctttaaccgttaagttaccttagagatttgaataagatggatattctcatcagtagtttgaaaagtttaggttattctaggacttccaaatctttagattcaggacctttggtagtacatgcagtagccggagccggtaagtccacagccctaaggaagttgatcctcagacacccaacattcaccgtgcatacactcggtgtccctgacaaggtgagtatcagaactagaggcatacagaagccaggacctattcctgagggcaacttcgcaatcctcgatgagtatactttggacaacaccacaaggaactctaaccaggcactttttgctgacccttatcaggcaccggagtttagcctagagccccacttctacttggaaacatcatttcgagttccgaggaaagtggcagatttgatagctggctgtggcttcgatttcgagaccaactcaccggaagaagggcacttagagatcactggcatattcaaagggcccctactcggaaaggtgatagccattgatgaggagtctgagacaacactgtccaggcatggtgttgagtttgttaagccctgccaagtgacgggacttgagttcaaagtagtcactattgtgtctgccgcaccaatagaggaaattggccagtccacagctttctacaacgctatcaccaggtcaaagggattgacatatgtccgcgcagggccataggctgaccgctccggtcaattctgaaaaagtgtacatagtattaggtctatcatttgctttagtttcaattacctttctgctttctagaaatagcttaccccacgtcggtgacaacattcacagcttgccacacggaggagcttacagagacggcaccaaagcaatcttgtacaactccccaaatctagggtcacgagtgagtctacacaacggaaagaacgcagcatttgctgccgttttgctactgactttgctgatctatggaagtaaatacatatctcaacgcaatcatacttgtgcttgtggtaacaatcatagcagtcattagcacttccttagtgaggactgaaccttgtgtcatcaagattactggggaatcaatcacagtgttggcttgcaaactagatgcagaaaccataagggccattgccgatctcaagccactctccgttgaacggttaagtttccattgatactcgaaagaggtcagcaccagctagcatcgatcgtcagatagccatttacggaaaaggtggtatcggtaaatctacgaccacccaaaacaccgtctccgccctcgccgagatgggtaaaaaaatcatgatcgtcggttgtgaccccaaagccgactctactcgtttgattcttcactccaaagcccaagcgacggttatggacctggcccgtgaaaaaggtactgtggaagacctcgaactggaagacgttctcctcaccggtttcgccgacatccgctgtgccgagtccggcggtcccgaacctggtgtaggctgtgccggccgcggtgttatcaccgccatcaacttcctcgaagaaaacggcgcttacacccctgacctggactatgtgttctacgacgttctcggtgacgttgtctgcggcggtttcgccatgcctatccgcgaaaacaaagcccaagaaatctacatcgtgacctccggtgaaatgatggccatgtacgctgccaacaacatcgcccgcggtatccttaaatacgctgctaccggtaaagttcgcctcggcggcctcatctgtaacagccgtaaaaccgacaaggaatacgaactgattgccgagttggccagccgtctgggcacgcaaatgattcacttcctgccccgtgacaaccaagtgcaacgggccgagctgcgccgcatgaccgttatcgaatactctcccgaacatccccaagccgacgaatatcgcgctctggccaagaaaatcgacgagaacaaaatgctcgttgttcccacaccgctttccatggacgaactggaagacctcctcatctcctacggcatcctggaagacgaagaaacggcgatggccaaactgggctaatcgtcgaccgccgatgaacggttaagtttccattgatactcgaaagatgtcagcaccagctagcacaacacagcccatagggtcaactacctcaactaccacaaaaactgcaggcgcaactcctgccacagcttcaggcctgttcaccatcccggatggggatttctttagtacagcccgtgccatagtagccagcaatgctgtcgcaacaaatgaggacctcagcaagattgaggctatttggaaggacatgaaggtgcccacagacactatggcacaggctgcttgggacttagtcagacactgtgctgatgtaggatcatccgctcaaacagaaatgatagatacaggtccctattccaacggcatcagcagagctagactggcagcagcaattaaagaggtgtgcacacttaggcaattttgcatgaagtatgctccagtggtatggaactggatgttaactaacaacagtccacctgctaactggcaagcacaaggtttcaagcctgagcacaaattcgctgcattcgacttcttcaatggagtcaccaacccagctgccatcatgcccaaagaggggctcatccggccaccgtctgaagctgaaatgaatgctgcccaaactgctgcctttgtgaagattacaaaggccagggcacaatccaacgactttgccagcctagatgcagctgtcactcgaggtcgtatcactggaacaacaaccgctgaggctgttgtcactctaccaccaccataactacgtctacataaccgacgcctaccccagtttcatagtattttctggtttgattgtatgaataatataaataaaaaaaaaaaaaaaaaaaaaaaactagtggtaccgagctcggtaccccatgcaagcttttgttccctttagtgagggttaattccgagcttggcgtaatcatggtcatagctgtttcctgtgtgaaattgttatccgctcacaattccacacaacatacgagccggaagcataaagtgtaaagcctggggtgcctaatgagtgagctaactcacattaattgcgttgcgctcactgcccgctttccagtcgggaaacctgtcgtgccagctgcattaatgaatcggccaacgcgcggggagaggcggtttgcgtattgggcgctcttccgcttcctcgctcactgactcgctgcgctcggtcgttcggctgcggcgagcggtatcagctcactcaaaggcggtaatacggttatccacagaatcaggggataacgcaggaaagaacatgtgagcaaaaggccagcaaaaggccaggaaccgtaaaaaggccgcgttgctggcgtttttccataggctccgcccccctgacgagcatcacaaaaatcgacgctcaagtcagaggtggcgaaacccgacaggactataaagataccaggcgtttccccctggaagctccctcgtgcgctctcctgttccgaccctgccgcttaccggatacctgtccgcctttctcccttcgggaagcgtggcgctttctcatagctcacgctgtaggtatctcagttcggtgtaggtcgttcgctccaagctgggctgtgtgcacgaaccccccgttcagcccgaccgctgcgccttatccggtaactatcgtcttgagtccaacccggtaagacacgacttatcgccactggcagcagccactggtaacaggattagcagagcgaggtatgtaggcggtgctacagagttcttgaagtggtggcctaactacggctacactagaaggacagtatttggtatctgcgctctgctgaagccagttaccttcggaaaaagagttggtagctcttgatccggcaaacaaaccaccgctggtagcggtggtttttttgtttgcaagcagcagattacgcgcagaaaaaaaggatctcaagaagatcctttgatcttttctacggggtctgacgctcagtggaacgaaaactcacgttaagggattttggtcatgagattatcaaaaaggatcttcacctagatccttttaaattaaaaatgaagttttaaatcaatctaaagtatatatgagtaaacttggtctgacagttaccaatgcttaatcagtgaggcacctatctcagcgatctgtctatttcgttcatccatagttgcctgactccccgtcgtgtagataactacgatacgggagggcttaccatctggccccagtgctgcaatgataccgcgagacccacgctcaccggctccagatttatcagcaataaaccagccagccggaagggccgagcgcagaagtggtcctgcaactttatccgcctccatccagtctattaattgttgccgggaagctagagtaagtagttcgccagttaatagtttgcgcaacgttgttgccattgctacaggcatcgtggtgtcacgctcgtcgtttggtatggcttcattcagctccggttcccaacgatcaaggcgagttacatgatcccccatgttgtgcaaaaaagcggttagctccttcggtcctccgatcgttgtcagaagtaagttggccgcagtgttatcactcatggttatggcagcactgcataattctcttactgtcatgccatccgtaagatgcttttctgtgactggtgagtactcaaccaagtcattctgagaatagtgtatgcggcgaccgagttgctcttgcccggcgtcaatacgggataataccgcgccacatagcagaactttaaaagtgctcatcattggaaaacgttcttcggggcgaaaactctcaaggatcttaccgctgttgagatccagttcgatgtaacccactcgtgcacccaactgatcttcagcatcttttactttcaccagcgtttctgggtgagcaaaaacaggaaggcaaaatgccgcaaaaaagggaataagggcgacacggaaatgttgaatactcatactcttcctttttcaatattattgaagcatttatcagggttattgtctcatgagcggatacatatttgaatgtatttagaaaaataaacaaataggggttccgcgcacatttccccgaaaagtgccacctgacgtctaagaaaccattattatcatgacattaacctataaaaataggcgtatcacgaggccctttcgtctcgcgcgtttcggtgatgacggtgaaaacctctgacacatgcagctcccggagacggtcacagcttgtctgtaagcggatgccgggagcagacaagcccgtcagggcgcgtcagcgggtgttggcgggtgtcggggctggcttaactatgcggcatcagagcagattgtactgagagtgcaccatatgcggtgtgaaataccgcacagatgcgtaaggagaaaataccgcatcaggaaattgtaaacgttaatattttgttaaaattcgcgttaaatttttgttaaatcagctcattttttaaccaataggccgaaatcggcaaaatcccttataaatcaaaagaatagaccgagatagggttgagtgttgttccagtttggaacaagagtccactattaaagaacgtggactccaacgtcaaagggcgaaaaaccgtctatcagggcgatggcccactacgtgaaccatcaccctaatcaagttttttggggtcgaggtgccgtaaagcactaaatcggaaccctaaagggagcccccgatttagagcttgacggggaaagccggcgaacgtggcgagaaaggaagggaagaaagcgaaaggagcgggcgctagggcgctggcaagtgtagcggtcacgctgcgcgtaaccaccacacccgccgcgcttaatgcgccgctacagggcgcgtcgcgccattcgccattcaggctgcgcaactgttgggaagggcgatcggtgcgggcctcttcgctattacgccagctggcgaaagggggatgtgctgcaaggcgattaagttgggtaacgccagggttttcccagtcacgacgttgtaaaacgacggccagtgaattgtaatacgactcactatapvx-hisg-mf1-nife序列（seqidno.14）：gaaaactaaaccatacaccaccaacacaaccaaacccaccacgcccaattgttacacacccgcttgaaaaagaaagtttaacaaatggccaaggtgcgcgaggtttaccaatcttttacagactccaccacaaaaactctcatccaagatgaggcttatagaaacattcgccccatcatggaaaaacacaaactagctaacccttacgctcaaacggttgaagcggctaatgatctagaggggttcggcatagccaccaatccctatagcattgaattgcatacacatgcagccgctaagaccatagagaataaacttctagaggtgcttggttccatcctaccacaagaacctgttacatttatgtttcttaaacccagaaagctaaactacatgagaagaaacccgcggatcaaggacattttccaaaatgttgccattgaaccaagagacgtagccaggtaccccaaggaaacaataattgacaaactcacagagatcacaacggaaacagcatacattagtgacactctgcacttcttggatccgagctacatagtggagacattccaaaactgcccaaaattgcaaacattgtatgcgaccttagttctccccgttgaggcagcctttaaaatggaaagcactcacccgaacatatacagcctcaaatacttcggagatggtttccagtatataccaggcaaccatggtggcggggcataccatcatgaattcgctcatctacaatggctcaaagtgggaaagatcaagtggagggaccccaaggatagctttctcggacatctcaattacacgactgagcaggttgagatgcacacagtgacagtacagttgcaggaatcgttcgcggcaaaccacttgtactgcatcaggagaggagacttgctcacaccggaggtgcgcactttcggccaacctgacaggtacgtgattccaccacagatcttcctcccaaaagttcacaactgcaagaagccgattctcaagaaaactatgatgcagctcttcttgtatgttaggacagtcaaggtcgcaaaaaattgtgacatttttgccaaagtcagacaattaattaaatcatctgacttggacaaatactctgctgtggaactggtttacttagtaagctacatggagttccttgccgatttacaagctaccacctgcttctcagacacactttctggtggcttgctaacaaagacccttgcaccggtgagggcttggatacaagagaaaaagatgcagctgtttggtcttgaggactacgcgaagttagtcaaagcagttgatttccacccggtggatttttctttcaaagtggaaacttgggacttcagattccaccccttgcaagcgtggaaagccttccgaccaagggaagtgtcggatgtagaggaaatggaaagtttgttctcagatggggacctgcttgattgcttcacaagaatgccagcttatgcggtaaacgcagaggaagatttagctgcaatcaggaaaacgcccgagatggatgtcggtcaagaagttaaagagcctgcaggagacagaaatcaatactcaaaccctgcagaaactttcctcaacaagctccacaggaaacacagtagggaggtgaaacaccaggccgcaaagaaagctaaacgcctagctgaaatccaggagtcaatgagagctgaaggtgatgccgaaccaaatgaaataagcgggacgatgggggcaatacccagcaacgccgaacttcctggcacgaatgatgccagacaagaactcacactcccaaccactaaacctgtccctgcaaggtgggaagatgcttcattcacagattctagtgtggaagaggagcaggttaaactccttggaaaagaaaccgttgaaacagcgacgcaacaagtcatcgaaggacttccttggaaacactggattcctcaattaaatgctgttggattcaaggcgctggaaattcagagggataggagtggaacaatgatcatgcccatcacagaaatggtctccgggctggaaaaagaggacttccctgaaggaactccaaaagagttggcacgagaattgttcgctatgaacagaagccctgccaccatccctttggacctgcttagagccagagactacggcagtgatgtaaagaacaagagaattggtgccatcacaaagacacaggcaacgagttggggcgaatacttgacaggaaagatagaaagcttaactgagaggaaagttgcgacttgtgtcattcatggagctggaggttctggaaaaagtcatgccatccagaaggcattgagagaaattggcaagggctcggacatcactgtagtcctgccgaccaatgaactgcggctagattggagtaagaaagtgcctaacactgagccctatatgttcaagacctctgaaaaggcgttaattgggggaacaggcagcatagtcatctttgacgattactcaaaacttcctcccggttacatagaagccttagtctgtttctactctaaaatcaagctaatcattctaacaggagatagcagacaaagcgtctaccatgaaactgctgaggacgcctccatcaggcatttgggaccagcaacagagtacttctcaaaatactgccgatactatctcaatgccacacaccgcaacaagaaagatcttgcgaacatgcttggtgtctacagtgagagaacgggagtcaccgaaatcagcatgagcgccgagttcttagaaggaatcccaactttggtaccctcggatgagaagagaaagctgtacatgggcaccgggaggaatgacacgttcacatacgctggatgccaggggctaactaagccgaaggtacaaatagtgttggaccacaacacccaagtgtgtagcgcgaatgtgatgtacacggcactttctagagccaccgataggattcacttcgtgaacacaagtgcaaattcctctgccttctgggaaaagttggacagcaccccttacctcaagactttcctatcagtggtgagagaacaagcactcagggagtacgagccggcagaggcagagccaattcaagagcctgagccccagacacacatgtgtgtcgagaatgaggagtccgtgctagaagagtacaaagaggaactcttggaaaagtttgacagagagatccactctgaatcccatggtcattcaaactgtgtccaaactgaagacacaaccattcagttgttttcgcatcaacaagcaaaagatgagaccctcctctgggcgactatagatgcgcggctcaagaccagcaatcaagaaacaaacttccgagaattcctgagcaagaaggacattggggacgttctgtttttaaactaccaaaaagctatgggtttacccaaagagcgtattcctttttcccaagaggtctgggaagcttgtgcccacgaagtacaaagcaagtacctcagcaagtcaaagtgcaacttgatcaatgggactgtgagacagagcccagacttcgatgaaaataagattatggtattcctcaagtcgcagtgggtcacaaaggtggaaaaactaggtctacccaagattaagccaggtcaaaccatagcagccttttaccagcagactgtgatgctttttggaactatggctaggtacatgcgatggttcagacaggctttccagccaaaagaagtcttcataaactgtgagaccacgccagatgacatgtctgcatgggccttgaacaactggaatttcagcagacctagcttggctaatgactacacagctttcgaccagtctcaggatggagccatgttgcaatttgaggtgctcaaagccaaacaccactgcataccagaggaaatcattcaggcatacatagatattaagactaatgcacagattttcctaggcacgttatcaattatgcgcctgactggtgaaggtcccacttttgatgcaaacactgagtgcaacatagcttacacccatacaaagtttgacatcccagccggaactgctcaagtttatgcaggagacgactccgcactggactgtgttccagaagtgaagcatagtttccacaggcttgaggacaaattactcctaaagtcaaagcctgtaatcacg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aaatcccttataaatcaaaagaatagaccgagatagggttgagtgttgttccagtttggaacaagagtccactattaaagaacgtggactccaacgtcaaagggcgaaaaaccgtctatcagggcgatggcccactacgtgaaccatcaccctaatcaagttttttggggtcgaggtgccgtaaagcactaaatcggaaccctaaagggagcccccgatttagagcttgacggggaaagccggcgaacgtggcgagaaaggaagggaagaaagcgaaaggagcgggcgctagggcgctggcaagtgtagcggtcacgctgcgcgtaaccaccacacccgccgcgcttaatgcgccgctacagggcgcgtcgcgccattcgccattcaggctgcgcaactgttgggaagggcgatcggtgcgggcctcttcgctattacgccagctggcgaaagggggatgtgctgcaaggcgattaagttgggtaacgccagggttttcccagtcacgacgttgtaaaacgacggccagtgaattgtaatacgactcactatapvx-hisg-mf1-nifn序列（seqidno.15）：gaaaactaaaccatacaccaccaacacaaccaaacccaccacgcccaattgttacacacccgcttgaaaaagaaagtttaacaaatggccaaggtgcgcgaggtttaccaatcttttacagactccaccacaaaaactctcatccaagatgaggcttatagaaacattcgccccatcatggaaaaacacaaactagctaacccttacgctcaaacggttgaagcggctaatgatctagaggggttcggcatagccaccaatccctatagcattgaattgcatacacatgcagccgctaagaccatagagaataaacttctagaggtgcttggttccatcctaccacaagaacctgttacatttatgtttcttaaacccagaaagctaaactacatgagaagaaacccgcggatcaaggacattttccaaaatgttgccattgaaccaagagacgtagccaggtaccccaaggaaacaataattgacaaactcacagagatcacaacggaaacagcatacattagtgacactctgcacttcttggatccgagctacatagtggagacattccaaaactgcccaaaattgcaaacattgtatgcgaccttagttctccccgttgaggcagcctttaaaatggaaagcactcacccgaacatatacagcctcaaatacttcggagatggtttccagtatataccaggcaaccatggtggcggggcataccatcatgaattcgctcatctacaatggctcaaagtgggaaagatcaagtggagggaccccaaggatagctttctcggacatctcaattacacgactgagcaggttgagatgcacacagtgacagtacagttgcaggaatcgttcgcggcaaaccacttgtactgcatcaggagaggagacttgctcacaccggaggtgcgcactttcggccaacctgacaggtacgtgattccaccacagatcttcctcccaaaagttcacaactgcaagaagccgattctcaagaaaactatgatgcagctcttcttgtatgttaggacagtcaaggtcgcaaaaaattgtgacatttttgccaaagtcagacaattaattaaatcatctgacttggacaaatactctgctgtggaactggtttacttagtaagctacatggagttccttgccgatttacaagctaccacctgcttctcagacacactttctggtggcttgctaacaaagacccttgcaccggtgagggcttggatacaagagaaaaagatgcagctgtttggtcttgaggactacgcgaagttagtcaaagcagttgatttccacccggtggatttttctttcaaagtggaaacttgggacttcagattccaccccttgcaagcgtggaaagccttccgaccaagggaagtgtcggatgtagaggaaatggaaagtttgttctcagatggggacctgcttgattgcttcacaagaatgccagcttatgcggtaaacgcagaggaagatttagctgcaatcaggaaaacgcccgagatggatgtcggtcaagaagttaaagagcctgcaggagacagaaatcaatactcaaaccctgcagaaactttcctcaacaagctccacaggaaacacagtagggaggtgaaacaccaggccgcaaagaaagctaaacgcctagctgaaatccaggagtcaatgagagctgaaggtgatgccgaaccaaatgaaataagcgggacgatgggggcaatacccagcaacgccgaacttcctggcacgaatgatgccagacaagaactcacactcccaaccactaaacctgtccctgcaaggtgggaagatgcttcattcacagattctagtgtggaagaggagcaggttaaactccttggaaaagaaaccgttgaaacagcgacgcaacaagtcatcgaaggacttccttggaaacactggattcctcaattaaatgctgttggattcaaggcgctggaaattcagagggataggagtggaacaatgatcatgcccatcacagaaatggtctccgggctggaaaaagaggacttccctgaaggaactccaaaagagttggcacgagaattgttcgctatgaacagaagccctgccaccatccctttggacctgcttagagccagagactacggcagtgatgtaaagaacaagagaattggtgccatcacaaagacacaggcaacgagttggggcgaatacttgacaggaaagatagaaagcttaactgagaggaaagttgcgacttgtgtcattcatggagctggaggttctggaaaaagtcatgccatccagaaggcattgagagaaattggcaagggctcggacatcactgtagtcctgccgaccaatgaactgcggctagattggagtaagaaagtgcctaacactgagccctatatgttcaagacctctgaaaaggcgttaattgggggaacaggcagcatagtcatctttgacgattactcaaaacttcctcccggttacatagaagccttagtctgtttctactctaaaatcaagctaatcattctaacaggagatagcagacaaagcgtctaccatgaaactgctgaggacgcctccatcaggcatttgggaccagcaacagagtacttctcaaaatactgccgatactatctcaatgccacacaccgcaacaagaaagatcttgcgaacatgcttggtgtctacagtgagagaacgggagtcaccgaaatcagcatgagcgccgagttcttagaaggaatcccaactttggtaccctcggatgagaagagaaagctgtacatgggcaccgggaggaatgacacgttcacatacgctggatgccaggggctaactaagccgaaggtacaaatagtgttggaccacaacacccaagtgtgtagcgcgaatgtgatgtacacggcactttctagagccaccgataggattcacttcgtgaacacaagtgcaaattcctctgccttctgggaaaagttggacagcaccccttacctcaagactttcctatcagtggtgagagaacaagcactcagggagtacgagccggcagaggcagagccaattcaagagcctgagccccagacacacatgtgtgtcgagaatgaggagtccgtgctagaagagtacaaagaggaactcttggaaaagtttgacagagagatccactctgaatcccatggtcattcaaactgtgtccaaactgaagacacaaccattcagttgttttcgcatcaacaagcaaaagatgagaccctcctctgggcgactatagatgcgcggctcaagaccagcaatcaagaaacaaacttccgagaattcctgagcaagaaggacattggggacgttctgtttttaaactaccaaaaagctatgggtttacccaaagagcgtattcctttttcccaagaggtctgggaagcttgtgcccacgaagtacaaagcaagtacctcagcaagtcaaagtgcaacttgatcaatgggactgtgagacagagcccagacttcgatgaaaataagattatggtattcctcaagtcgcagtgggtcacaaaggtggaaaaactaggtctacccaagattaagccaggtcaaaccatagcagccttttaccagcagactgtgatgctttttggaactatggctaggtacatgcgatggttcagacaggctttccagccaaaagaagtcttcataaactgtgagaccacgccagatgacatgtctgcatgggccttgaacaactggaatttcagcagacctagcttggctaatgactacacagctttcgaccagtctcaggatggagccatgttgcaatttgaggtgctcaaagccaaacaccactgcataccagaggaaatcattcaggcatacatagatattaagactaatgcacagattttcctaggcacgttatcaattatgcgcctgactggtgaaggtcccacttttgatgcaaacactgagtgcaacatagcttacacccatacaaagtttgacatcccagccggaactgctcaagtttatgcaggagacgactccgcactggactgtgttccagaagtgaagcatagtttccacaggcttgaggacaaattactcctaaagtcaaagcctgtaatcacgcagcaaaagaagggcagttggcctgagttttgtggttggctgatcacaccaaaaggggtgatgaaagacccaattaagctccatgttagcttaaaattggctgaagctaagggtgaactcaagaaatgtcaagattcctatgaaattgatctgagttatgcctatgaccacaaggactctctgcatgacttgttcgatgagaaacagtgtcaggcacacacactcacttgcagaacactaatcaagtcagggag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