CN111073898A - 小麦条锈病抗性基因YrU1及其克隆与应用 - Google Patents
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Abstract
本发明提供了小麦条锈病抗性基因YrU1及其克隆与应用,属于植物基因工程技术领域。所述小麦条锈病抗性基因包括自身启动子和5′‑UTR、3′‑UTR在内的基因全长的核苷酸序列如SEQ ID NO.1所示,其编码区的核苷酸序列如SEQ ID NO.2所示,编码的蛋白其氨基酸序列如SEQ ID NO.3所示。小麦条锈病抗性基因YrU1在小麦抗性育种中具有广阔的应用价值。
Description
技术领域
本发明属于植物基因工程技术领域。具体涉及从乌拉尔图小麦材料中,定位、克隆、验证小麦条锈病抗性基因YrU1以及该基因在条锈病抗性育种中的应用。
背景技术
小麦是重要的粮食作物,为世界上大约35%的人提供主食(Huang and Röder,2004; Zhou et al., 2014)。小麦条锈病是由条形柄锈菌(Puccinia striiformis.)引起的危害中国以及世界其他小麦主产国小麦生长的最严重病害之一,每年都会导致小麦大量减产,在大部分小麦生产地条锈病的流行能导致10%~70%的减产,如果发病较早且持续感染时间长,甚至能导致100%的减产(Chen, 2010; Fu et al., 2009)。由于条锈病的发病区域的扩大和产量损失的增加,小麦条锈病已成为所有作物锈病中最严重的病害(Beddow etal., 2015)。种植具有条锈病抗性的小麦品种是防治条锈病非常有效的、经济节约并且环境友好的措施(Chen, 2005; Line et al., 1995; Roebbelen and Sharp, 1978;Schwessinger, 2016)。
迄今为止,已经有超过80个小麦条锈病抗性位点被鉴定(Pakeerathan et al.,2019),仅有少数基因被克隆,其中Yr5, Yr7和YrSP是一个基因簇,编码BED-NLR蛋白,Yr15具有两个类激酶结构域,Yr18/Lr34编码一个ABC转座蛋白,Yr36具有一个激酶结构域和一个脂类结合结构域,Yr46编码一个己糖转运蛋白,YrAS2388编码一个NLR蛋白(Fu et al.,2009; Klymiuk et al., 2018; Krattinger et al., 2009; Marchal et al., 2018;Moore et al., 2015; Zhang et al., 2019)。Yr5, Yr7,YrSP和YrAS2388编码NLR蛋白,是全生育期抗性基因,Yr15,Yr18/Lr34,Yr36和Yr46编码不同的蛋白家族且具有广泛的条锈病抗性。由于条锈菌的进化,小麦抗病基因的持久性不断受到挑战。
植物通过细胞内NLR受体监控各种病原菌的侵染(Jones et al., 2016)。大部分NLR受体包含三个结构域:C端可变结构域,NB-ARC结构域和LRR结构域,但是少数NLR受体除了这三个结构域外,还具有额外的结构域作为整合的诱饵结构域(Li et al., 2015)。已经发现的NLR受体中,拟南芥的RRS1蛋白包含一个C端的WRKY结构域,水稻RGA5蛋白包含一个C端的RATX1结构域,小麦基因簇Yr5, Yr7和YrSP包含一个N端BED结构域(Cesari et al.,2014; Marchal et al., 2018; Sarris et al., 2015)。
由于小麦的育种方式,现在种植的小麦品种90%以上都对条锈病感病。乌拉尔图小麦是六倍体小麦A基因组的祖先种,可以为小麦育种提供更多的抗病基因资源。乌拉尔图小麦G1812的精细基因组序列已经发表,为克隆基因提供了便利(Ling et al., 2018; Linget al., 2013)。我们以乌拉尔图小麦为材料,定位和克隆新的条锈病抗性基因,并对发现的条锈病抗性基因的抗病机理进行初步探索,为小麦抗病育种提供新的基因资源和理论基础。
发明内容
本发明提供了小麦条锈病抗性基因YrU1的克隆与应用,为小麦条锈病抗性育种提供新颖、高效的抗性基因,培育更加持久高效的条锈病抗性小麦品种,拓展对植物应对病原菌的抗病机制的研究。
为了解决上述问题,本发明克隆了一个来源于乌拉尔图小麦PI428309的条锈病抗性基因YrU1,其编码区核苷酸序列如SEQ ID NO.2中的第1-4731位核苷酸所示。
条锈病抗性基因YrU1的启动子区核苷酸序列如SEQ ID NO.1 中的第1-2104位核苷酸所示。
条锈病抗性基因YrU1的5′-UTR和3′-UTR的核苷酸序列如SEQ ID NO.1 中的第2105-2714位和第9773-10537位核苷酸所示。
乌拉尔图小麦的条锈病抗性基因YrU1的核苷酸序列如SEQ ID NO.1所示,通过对该序列进行一个或者多个核苷酸的替换、缺失或***后得到的仍能编码相同或是相似功能蛋白的核苷酸序列也属于本发明内容。
本发明还提供了条锈病抗性基因YrU1编码的蛋白,其氨基酸序列SEQ ID NO.3所示。
本发明还提供了含有基因YrU1的载体。
本发明还提供了所述小麦条锈病抗性基因YrU1在制备条锈病抗性转基因小麦中的运用。
文中所述抗病或是感病如无特殊说明即指条锈病。所述条锈病是指小麦条锈病,由小麦条锈菌即条形柄锈菌小麦专化型所引起,下文中如无特殊标识小麦条锈菌则具体指广泛流行于中国的小麦条锈菌菌株CYR33, CYR17,CYR31,CYR32和V26(Pst CYR33,PstCYR17,Pst CYR31,Pst CYR32和Pst V26)。
本发明的有益效果在于:
(1)本发明通过图位克隆的方法在乌拉尔图小麦中克隆了一个条锈病抗性基因YrU1,该抗性基因不仅在乌拉尔图小麦中具有条锈病抗性,而且通过转基因方式将自身启动子序列启动的YrU1基因转入普通小麦中同样具有条锈病抗性,说明在不同的麦属植物中具有类似的条锈病抗性应答网络。
(2)源于乌拉尔图小麦的条锈病抗性基因YrU1对我国流行的条锈菌生理小种CYR33,CYR32,CYR17和V26都具有抗性,说明YrU1基因在小麦抗性育种中具有重要的应用价值。
附图说明
图 2条锈病抗性基因YrU1编码一个新的CC-NNS-LRR蛋白,在其氨基端连接一个额外的ANK结构域,羧基端连接一个额外的WRKY结构域。
图3 乌拉尔图小麦PI428309和G1812中YrU1和CG2基因在接种条锈菌CYR33前后的表达量变化图。
图4在乌拉尔图小麦材料PI428309中以大麦花叶条纹病毒诱导沉默YrU1,可导致该材料对PstCYR33感病。
图5以YrU1基因稳定转化普通小麦感病品种Bobwhite,转基因植株获得了对PstCYR33的抗性。
具体实施方式
下述实施例用于阐述、解释本发明,而并非用来局限本发明的范围。
本发明申请人在对一百多个乌拉尔图小麦材料进行条锈病抗性鉴定中,发现了PI427328,PI428294,PI428209,PI428214和PI428309等5个材料对条锈菌生理小种CYR33具有抗性,其中PI428309对条锈菌生理小种CYR17,CYR31,CYR32和V26也具有抗性,另外还发现已经测序的乌拉尔图小麦材料G1812对5个条锈菌生理小种呈现高感的表型。我们构建了PI428309和G1812的杂交群体,通过分析杂交F2群体接种PstCYR33后的抗感分离比,发现在PI428309材料中存在一个显性的条锈病抗性位点(暂命名为Yr)。利用普通小麦A基因组已公布的分子标记和依据G1812的序列设计的分子标记,将此抗性位点初步定位在5AL上0.2cM范围内,并发现一个共分离标记SCF20 (图1a),遗传图谱上的标记引物如表1所示。随后我们进行了更精细的定位、候选基因的克隆和验证,并分析了抗性基因YrU1所编码的蛋白结构。为了便于读者深入理解本发明的内容和目的,将通过以下实施例来详细介绍本发明的具体技术实施步骤。实施例中所述技术手段为本领域内人员所熟悉的常规手段。
表 1引物序列表
实施例1条锈病抗性基因的精细定位
为了更精细的定位条锈病抗性基因,我们通过二穗短柄草基因组序列与共分离分子标记SCF20的共线性序列BLSATN乌拉尔图小麦材料G1812的序列,设计SSR标记和CAPs标记。最终通过3,034个PI428309×G1812 F2纯合群体,将条锈病抗性基因位点Yr定位在G1812基因组的65 Kb范围内(图1)。G1812的基因组序列在该区域内有三个候选基因,TuG1812G0500003718, TuG1812G0500003719和TuG1812G0500003720,且都编码NLR蛋白,这三个基因序列可在上获取。
条锈病抗性位点定位实验中所涉及的SSR分子标记是通过SSR Locator软件设计的,CAPs标记是通过在线软件http://helix.wustl.edu/dcaps/dcaps.html dCAPS Finder2.0设计产生的。分子标记的PCR扩增反应体系为10 μL PCR体系(康为世纪2×Taq MasterMix 5 μL,10μM 的正反向引物各0.5 μL,灭菌去离子水2 μL,DNA 模版2 μL),扩增反应在Eppendorf PCR仪上进行。PCR程序为:95℃预变性5 min,95℃变性30 s,55℃退火30 s,72℃延伸1 、30 s,变性、退火、延伸三个步骤35个循环,最后72℃充分延伸10 min,结束程序,转入4℃保存阶段。SSR分子标记扩增产物以8.0% 非变性聚丙烯酰胺凝胶电泳,CAPs标记扩增片段的酶切产物以4%的琼脂糖凝胶电泳,检测标记在F2群体单株间的多态性。
实施例2 条锈病候选抗性基因的克隆
G1812中的TuG1812G0500003718, TuG1812G0500003719和TuG1812G0500003720均编码R蛋白,为了克隆抗条锈病基因,我们根据这三个基因的序列设计扩增引物,通过根据TuG1812G0500003718和TuG1812G0500003719设计的引物YrU1-C和CG2-C得到了PI428309材料中的两个等位基因,分别命名为YrU1和CG2。我们也通过引物YrU1-C在PI428309的cDNA中扩增得到了YrU1的编码序列(SEQ ID NO.2),其编码的蛋白,氨基酸序列如SEQ ID NO.3所示。CG2在PI428309中不能检测到表达,从而未得到CG2的编码序列。由于G1812和PI428309两个材料之间的序列多态性,在PI428309中不存在TuG1812G0500003720的等位基因。我们比较分析了YrU1和CG2基因在PI428309和G1812之间的序列差异,结果表明YrU1在PI428309中存在额外的~1 Kb的序列,而CG2在PI428309和G1812之间只存在SNP的差异。PI428309中YrU1的~1 Kb额外序列编码一个WRKY结构域,而该结构域在G1812中不存在(图2)。我们检测了PI428309和G1812中YrU1和CG2基因在接种条锈菌CYR33前后的表达量变化,结果表明YrU1基因轻微的受到条锈菌侵染的影响,而CG2基因在接菌前后均未检测到表达(图3)。基于以上结果,我们选定YrU1基因作为候选条锈病抗性基因进行后续功能验证。
实施例3 条锈病抗性基因YrU1的基因功能验证
为了确定乌拉尔图小麦PI428309中是否由YrU1来响应对条锈菌CYR33的抗性,通过大麦花叶条纹病毒介导的基因沉默试验(VIGS)和转基因互补试验进行验证。
通过VIGS试验对YrU1的功能进行验证,首先通过引物YrU1-VIGS将一段247 bp的序列(SEQ ID NO.4所示)反向构建到大麦花叶条纹病毒RNAγ链中组成重组质粒BSMV:YrU1,然后通过大麦条纹花叶病毒将YrU1片段导入小麦叶片中(具体的实验方法参照Zhang, Y., Bai, Y., Wu, G., Zou, S., Chen, Y., Gao, C., and Tang, D. (2017).Simultaneous modification of three homoeologs of TaEDR1 by genome editingenhances powdery mildew resistance in wheat. Plant J. 91, 714-724),结果表明在PI428309中瞬时沉默YrU1基因后,降低了PI428309对Pst CYR33的抗性(图4)。
由于乌拉尔图小麦转化技术还不成熟,我们将YrU1基因转入普通感病小麦Bobwhite中。根据TuG1812G0500003718的上下游序列设计引物YrU1-XbaI/PstI,以PI428309基因组DNA为模板,扩增得到YrU1 全长基因组序列(SEQ ID NO.1)(包括自身启动子和5′-UTR、3’-UTR序列),然后通过XbaI和PstI双酶切连接导入到双元表达载体pCAMBIA1300中。包含YrU1基因片段的pCAMBIA1300载体通过基因枪轰击的方法导入到普通感病小麦Bobwhite中,获得了稳定的转基因植株。通过PCR和qRT-PCR验证了转基因植株中YrU1基因的存在。稳定的转基因植株表现出对PstCYR33抗病,证明了候选基因YrU1就是PI428309中的条锈病抗性功能基因(图5)。
以上内容通过一般性的说明和具体的实施方案对本发明进行了详尽的阐述。对于本领域内的工作人员而言,对本发明进行某些更改和衍生是轻而易举的,因此在不偏离本发明基本内容而做任何改动都属于本发明要求保护的范畴。
SEQUENCE LISTING
<110> 福建农林大学
<120> 小麦条锈病抗性基因YrU1及其克隆与应用
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<160> 78
<170> PatentIn version 3.3
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gtgatgtttg atagcacaca aggtattcct ccggtattcg ggagttgcat aatctcatag 60
tcgaaggaat atgtatttga catgaagaaa gcaatagcaa taaactgaac gatcattatg 120
ctaagctaac ggatgggtct tgtccatcac atcattctcc taatgatgtg ctctcgttat 180
caaatgacaa cacatgtcta tggttaggaa accttgacca tatttgatca acgagctagt 240
ctagtagagg cttactaggg acacggtatt tgtctatgta ttcacacatg tatttaagtt 300
tccggtcaat acaattatag catgaataat aaacctttat catggttaag gaaatatagt 360
aataaccatt ttattattgc ctctagggca tatttccatc agaaggaata tgtatttgac 420
atgaagaaag caatagcaat aaactgaacg atcaatatgc taagctaatg aatgggtctt 480
gtccatcaca tcattctcct aatgatgtga tcccgttatt aaatgacaac tcatgtctat 540
ggttaggaaa ccttaaccat ctttgatcaa cgagatagtc tactagaggc tcactaggga 600
cacggtattt gtttatgtat ccacgcatgt atttaagttt ccggtcaaca caattctaac 660
atgaataata aacctttatc atgatttagg aaatataata ataaccattt tattattgcc 720
tctacggcat atttccatca gcgggtcctt gcaatgtccc ttcctcgatg acatgttgga 780
ttttgtccct tggagctatt caaaaaatag aaagtttcct gttcaatcaa cctatgctat 840
ggagtggaaa cacgaacatg ggagaaaact tagaagaatt gataatatgg gacatctaat 900
tattaatcct atttggagtg agatttggaa gttatcttgc ccgttggagg tatttttttt 960
cttcatattt tgcatggaac tctttggcaa atagacacat aaaagtttct cccaattttc 1020
cttcatgttc ctttggatca gaagatacaa aacatatgtt atttcaatgt cacaaagcaa 1080
atgaggtttg gaaagtggcc aggctggatc aagtcattaa aatagcttgc aaagttgatt 1140
atgctggtga ggctgtccta gaatatttac tttcgatacc tgaacatgag ttggctatct 1200
tgggtattca gaatgttcgt gggatgattg caattgctac ttggtatttg tggtgggaaa 1260
ggcgaaaatt ggtccataag gagaaaaccc aagatgctaa ccagatttct atggggattc 1320
acgctattac attaaactat gttataacta tatcacctaa ggctactatg aaaagagggg 1380
gttggacaca acctccgagg gggtttgtaa aacttaatgt ggtatatgct catgtgtcgc 1440
cgagtacaac tgcaacattg ccggatattc aagaaaacag tctaaacgac ccttaaaacc 1500
ttcttatctg ttatcctagg taaatgggac ttctgtaagt cccatactct cattaaccaa 1560
gtgtttgctt cattgttgta ctgtaaatct catactctca ttaaccaagt gtttgcttca 1620
ttgttatact ttaaggttgg ctgcaactgg aacacgtggg tcttcttaat ggtgttttcc 1680
ctaggctttt gatttaagtg tgcatttttg ctgcctctat acatggacct acctctagca 1740
aaaaattctt gacatcgcct tcaagctttt ggttttatta aaatatccat cagtccctac 1800
atgtcatccc ttctgttgag cctcccaaca aaaaaaatga ggctcctaca tgtcatcatg 1860
tcatcggtgc gagccggttg acgtttggga gcaatcagct ggtggatgct aatcatttgc 1920
atctgtattg cggtaccggg ctggatatga ctgagcaggt ggggtgcggc acagggggcc 1980
cggaaggaga aaggagagga gatgaggact gagtggctcc gcatgtagta cagtaatgcg 2040
ttcttccctt cgatttccta cagacctaca tactattcgt agtcgccccc ttttatcgag 2100
ctgtgatgca gatgcttgcg cattggcttt ggcttcagag ggatcgacca agaaaatgaa 2160
gagcacaaga tcgaagactg ccaacgtgcg cgggctcctc tgttcaagca gttatcgtct 2220
tccgctgata agcagcaagc gtggtgactg agacggacgg gaggaagttt ggaggctccc 2280
gacaaggcga caagtgagta cgttcaattt gtttgctgtt tcagtgggat tacccgcgga 2340
actcgttgta atgttaaatt tctgactggt atgcagttcg atgtgtatgt gccgtggggc 2400
agaaggatcc gcctcttggc ctggttgcag cgggagatct ctattgggaa attgagaatt 2460
ttccgtagtg tgtgcggaag tgtcgttctt ctgaatcgag cgaagggagt tataaaggaa 2520
gatcttgccc tattctttga gttgaagctt ggtttcccaa tgttaaccta gatctgcaag 2580
aacacgatct tgcagttttg gacttcgtca caatatatca ccaaaattaa tcaagttctt 2640
gtatttggag atagagaggc acaatccagg ctccgcgctc tagagaaatt gaagaccttc 2700
tggggttctt atccatggag cagcagcaga gcttcgagga gaggacgagc aaagatagcc 2760
ccgagaagat gggtgacagt gcgctccacc ttgcagcaag ggccggaaat gttgcccttg 2820
tgcagaagat ctttgcggac tgcgatccgg agctggtcgc ggaattgact gaccatcaga 2880
accaggacgg tgagacagcg ctgtatgttt ccgcggagat ggggcatgtc gaggttgtgt 2940
gtgaaattct gaaggtttgt gatctgcatt cagcatttct caaggcacac aacagctttg 3000
atgcattcca tattgcagca aagcagggcc atctaggtta gttctttact tctttccttt 3060
catcactaat tcttgtatat tgttgagttt aaccacaatg tgctatgttt gtcaagtcat 3120
tgattggacg attatggttg tgtgaattgg aattcttcag tactccttag ttggattgtt 3180
tatagttgtg attttctttg tcttgtatac agtgacttgt atttcattac ctttgtcccc 3240
aaagcattta ccaggtgtcc gcttttgcat tcaatgttta agaaatttat ctgggtgatt 3300
tggtccatat tccagcctat attgcctggt tttcttggta tggcaggcaa aagtgaccat 3360
ggattcacag ttttatgtag gatttgttgc tgtcgcgata tgcttataaa atattgatgt 3420
tgatgtatcc aaatcagggt atctgatatg attgattagc gagataatta aatgttgcat 3480
cttcgtgctt tttttttgtg ccagcaattt gtgctagaac aaatcagctc atgcaattta 3540
taatagaagg cagtggagaa gagcaaacca ctgtcctact cctgatccta agttcctaac 3600
tactactccc tcttccataa ttttgaacta aaaccatgac aagaattatg ggacggaggg 3660
agtatgatag ttgtatcttt gtggatgttt ggcttaatgt tattgtcaag ttatttcggc 3720
tcatatacat gtcccatgct ttgtttcgca gttgttttgc aggagctact gaaggctttt 3780
cctgcattag ctatgacaac aaattcagta aatgccacag ctttatacac tgccacactt 3840
cactgccaca ttggtattgt caatcttcta ctggatacag acccaagact tgccaggatt 3900
gcaagaaata atgggaagac agctctgcat atagcagcaa ggctgggcaa tgtggaggtg 3960
gtagtgttgt tgttgaataa agatccggcg actgttttca gaatagacag gaagggacaa 4020
acagcagtgc acatggcttc caaaggccac aatgctgaaa ttctgcttga gctactgaag 4080
cccgatgtct cagtaatcca tttggaagat aacaaggggg acaggccact gcatgttgca 4140
acgcggaagg gaaaaaccat tgtaggttac catatctcta aaatatcttc tgttttcttc 4200
attatttcat tcttcagatg ttaacaattt gttgttgcat attatttctt gttcatccta 4260
tgttcatttg caaagtttgc ctgcttaaac cagcaatatt agattttcta gaacataaag 4320
tatctaaagc aatttacact acattatgta cacaaataat gaaaagactt cctttcatag 4380
acttgagaga actatgaagt tctaagcagt ttcaaccaga atttcagtgg acttatgtat 4440
atgtacttac gattttaaga ttttttctta gtaatttgat cctttttctg atattcgcat 4500
aattccttac agatagttca gacgctaata tccattgaag agattgatat caatgcaatc 4560
aatggagctg gagagaccgc ttttgccatt gcagagaaac tgggtaatga agagcttgta 4620
aacatcctga gggaggctgg tggagtaacc gcagaagagc aagtaaatcc tccgaaatca 4680
atcaagcgtt ttaagcaaac acatgatgtc caatcgcaga tcaagcaaaa gcgtcggaca 4740
aatatgcatt tccacacgat caggaagagt agtcaaaagc tccacactga ggctccagtc 4800
tgcgctttgg cagacgccat gttcagactt cctgcaaagc tcgatgagct actgattagc 4860
cacgtccaca tgcttcctag gggtgcggag gatgagatac ctctcatcaa gcaagatctg 4920
gaagagataa tggccattct gcaggagcac gaccacccag ggagagcgga agaccgtgct 4980
atgacgagca agtgcctgac caaggaggtg cgcgagctgt catacgacat ggaggatagt 5040
gtcgaccagt atgtgcacgc cgtcgacacc aagagaagga ttgttcctcg ccgtaaaaag 5100
tacaagatca cctgtcgtag gggcaagacc actgcgcggc tcccggagaa gcttaagtgg 5160
cgtatatgga tggccaacaa gatcagggag ttcagtgtgc gctcgcaaga ggcgctgcag 5220
cggtacagcc tatttaacca ccctggtgct catgacatca gcacgtctgc tacttctacg 5280
agacatgatg tgtgttttgg ctcttggtat cccacaccgt gtggggagct tgtcggtata 5340
gatggacatt tgaatactct tgaagcgtgg ttgggtaagg atggggagca gcagctcaag 5400
gtggtatctg ttgttggatc tggaggggtt ggtaagacca cactttccaa agagctgtac 5460
cgtagaatca gagggcaatt cgagtgccag gcatttgtga ggacgtcccg gaagcccgac 5520
atcaggaggc ttctcatcag cttgctctca caagtccggc cacaccaaac ccctcacact 5580
tggaaattgc atagtctaat tgccgatatc aggacacatc tccacgataa gaggtataca 5640
tgtctcttga ttcataagaa ttacattttc tggacaggaa tttaatcttg tgagcaggcc 5700
cttagtgtga tttttccatt atacaaacat gttactgcgg catcttagtg caccaaaatg 5760
gtttgcggtt cgatttcttc cttagttcaa cgatatttcg gtggtggtga agatttgaag 5820
tcactgataa tatttttgta tgcagctatg ctgagccact tttttttcct tccagtccag 5880
taagtataga gtggcagatt tccaatgcgt tattaggctg acaactgtgc taaaacattc 5940
ctcaggtact tgatcgtcat tgatgatgta tgggctacac aaacatggga tatcattaat 6000
cgtgctttgc cggctggtaa tctttgcagt agaattctaa taacaacaga agtcgaagat 6060
gtagctctga aatgttgtgg ttatgactct aggcatgttc ttatggtgaa accacttggt 6120
tacgatgatt caagcaaatt atttttcagc acagcttttg gactacaata tgaatgtcct 6180
ccagaactct gtgacgctgc acacaacatt gtgaggaaat gtgctggttc accactagca 6240
atggttactg tggctagtct tttagtaagc cagattggca aaccagagaa atgggattat 6300
gtaaatgaaa tcttaggtca cggtttgagc acatatccta gctcggaagg aatgaaacaa 6360
gtactaaacc ttagttacaa caatcttcct cattatttga aggcatgtgt gatgtatctc 6420
agtatatatg aagaggacta cataattcag aaagatgatt tggtaaagca atggatagct 6480
gaaggtctta tcctagcaac agaagagaaa gacaaagagg aaatatcaag gagatatttt 6540
gatgagctta tcagtagcag aatgatccta cctgtgtata caaatgacaa cgatgatgtt 6600
ttgtcctgca cactgcatca catggtactt gattttatca aacacaagtc cttagaagag 6660
aattttgtca tcccaataga tcatagtcag acaactgcac cactcgcaga caaggttcgt 6720
cgactgtctc tccactttgg taattcagaa gcaacgccac caacaaatat gagactatca 6780
caagttcgga ctctcgcatt tttcggggtc attgagtgtt tgccttccgt tatagagttt 6840
cggcttcttc aagtcctaat cctacatctt tttggcgatg atgaaagtgt cagttttgat 6900
ctcactggaa tatctgagct ttttcggttg agatatttgc atgtcacatg taatgccacc 6960
ttagaagtac cacaaactca gatgcgaggt ttacaatatt tggagacact gaaaatagat 7020
gcaagagtaa gtgcagttcc gtcggacatt gttcatttgc cgagcttgtt gcacctcagt 7080
cttcctgttg ggacaaatct accaaatggt attgaccata tgacatcgct ttgcacactt 7140
gaatattttg atataaatgt taactcaatg gagaatgtgc acagccttgg tgagctgacc 7200
aatcttcagg atcttcggct cacatgttct acagttcctt cttcttactt aaagagtaaa 7260
atcgatagta tgggctctat tcttgcgaac ctcagcaacc tcaggtctgt aactctgaag 7320
tcttcaggta ttctggagag tgaaccttac agcatgatca tttcctgtga tggcttgagc 7380
agcgtttcct ctcctccagc tcttcttcaa agatttgagt ggttgccacg catttgtacc 7440
ttctccagca tccctaagtg gattagccat ctcaacaagc tctgcatttt aaagattggg 7500
cttagggaat tagtgagcaa tgatgtcgct gctctgagag gattgcctgc actaactgtt 7560
ttgtcgttat atgtccgagc aaagcccgca gaaaaaattg tctttagtag ggcaggattc 7620
ttggttctca agtgcttcaa gttcaggtgc agtgtacctt ggctggaatt tgaggtggat 7680
gcaatgccta atctcttgaa actcaagcta agtttcgatg cccatggagt agatcaacat 7740
ggtactatac ctgtcggcat agtgcactta acaggcctta aggaaatctc tacaaaaatt 7800
tggggtgctg gttacaatga aagaagggct gcaaaatcag cgctgattgc cgctattaag 7860
atgcattcgg gatgtcccac cgccagcata cagtgtttag atgggatgtt cagtggtaag 7920
gatgataata atagcgggat acaagaggaa gaacacatgg ctctgcaaaa gcagcaatac 7980
aatatcatgg aggaagactt gatggaacag cataaagtcc gtcaaaaagg cacgtgggaa 8040
gatgctcaca gaaggtatga tgatgatctc tctcattcat gtgctttgct aagtgagttt 8100
tggtggtggc taaacgcttt gctgccatta ggttttctgc attttcagat tggaaagctg 8160
ttgcggacaa gcaacctaac agtaggtatg tatttgtcag tagtaattaa ttataactga 8220
aataaacata ttttgttctt gctcttgctt cgttcaatat cctaatagat gctaaagcta 8280
ctcagggatg ccacgtctgt tgccttttaa atcctccatg ttttctgatg ggcagagtgc 8340
accatccgtg ccgggttggg aagacgaccg gactgtcaac atcaaggaag tggacagcat 8400
ggctgaagac agagacacag aaatacaaga gcatggataa agactccaat aaacagcatg 8460
gaggtcttcc aaaggaccac atggatgttg cagacaaaca agctgccagt aggtatgtat 8520
ttgtccgttg taactaatta actaacgaat cttctcttct cgctatagct ccgttccata 8580
ccctaatgga tgctaaagct gctcatggat gtcacttgtt ttgcttttta taacttctgt 8640
gtctttggaa cggcagatca cagcacctaa taaccagcat gtagttcttc aaaaggacaa 8700
cgtgcttata ataatccatt agggttcaca aagctatact ctgagaattt atggcatgtt 8760
cctgtttatc tcttagcaac cataaaatag taatagaaga agatttcatg tatttcttta 8820
attactatct atatctatac atgtatatat tgcttttgag agtttatggc atgtttctgt 8880
ttatctcgta gcaacaatcg taggattccc aggcggacca ggaagatgag tagtcggagg 8940
atgcaggtga gggtaggatc cttgcaggac ttgggtttcc tcgacgatgg cttcagctgg 9000
aggaagtacg gccagaagga taccatgggc tccacgtacc caaggtcaga aggaactctc 9060
gcctcaagcg caccaagtat ttctaggccc tgtttgtttg ggcttttgct tttgcttctg 9120
cttttgcagt ttttttcatt ttggccaaaa agccataaaa gctcctaaat agatgctttt 9180
ggagccaaat attgttatat tgattcatca aaatccataa aagctccaaa agcgtctatt 9240
taggagtttt tatgtctttt tggtcaaagc ggaaaagctg caaaagcaga aataaataaa 9300
ggcccaaaca aacagggcct aatcttttcg ctgcacctga cagagactga ctcgaggcac 9360
ttgttgaaat tgaattctca attgcagagc ttatttccgg tgcacgcaca ggcacgttaa 9420
gggctgccta gcgaccaagc aggtgcagcg cacgtctaac gacccgctgc tctttgacgt 9480
ggtgcataac ggggagcaca cgtgcttgga ctctgtcgga tcccctgcga cgtccgggag 9540
ccaggttacc tacgagggga taaatgcaac cgcggcaacg tcttgtggcc aggtcacctt 9600
ggaggaggtg agcagaggta cgcccggagt tggattcatg tccccggcgg tgttcagtag 9660
ccaggttatg tccgaggtgg taagcggaag cgggagtacg gcgggatttt ctggtgatga 9720
ggttggcatg ccagacccgg accgcgatgg ctacgaattt gaccgagttt gaggtcagcg 9780
ctttcttggc ctagtgagat ttgatcatct cctcgcaggg ggcgggcggg gccacgattt 9840
gagcacacgg agggccaagc catgttagta tttatataag gccatgcata tcccatggtg 9900
ccttgttatc ggttctatgg gttgttctgc ttggggcgat ggtgtgtggt tgacccatgg 9960
tgccttgtca gtggggaggg aggaatccgt tgatccaatg tgctcggagg ctaagtgccc 10020
cggcgactgc tcctcttcca ttgctcagtc cccagcacgt ccgatgctta gtccagagga 10080
gccgcttgtt ttcgacgcag ggcgcatgat gggggccttc ggcatgcccg atgttctagt 10140
tgagcctccg gttggggtgt gcggtgttgt tgcgaagggg acacagacta aagcaacgcc 10200
attatctcta gaggattctc tcaccaaggt tacttgtgca ctaccacaac ctctgcttgg 10260
caccccggcg cctttgcacg gggagaagga tgacgagcga caaagtaggc gtctggaaaa 10320
gaagaacaag gtgtgcaaca ttctgacgtc caagtacgca gagcacaggt tgttagagac 10380
ttttggtgag ctgccggagg tcagcgatgc ggagggccct aagcaaaaga tgaaagcata 10440
tctagacatg tataaaaaac cgctctcacc gcaagtgatc aaggggttga ggactttggc 10500
gggcatcaca gaaaagtcaa agttgaacct ttcagcatgt tttccacttg ataatgttcg 10560
cacgtaatgc tttcaggctg agacagaagt gtcatcgata tgtacgattt aatcaccgat 10620
accataatta gagcaactcc aacaattcct tcaaaaaact tcccgatttc gctgaatttt 10680
agaggaagtt tcccgcaaag tactttcgtc gcatcccgta aaacatctgc ccctaaattt 10740
gtttttattt tatttttcat taaacaaaac gaaataatgg cctagggcct gccaacaagt 10800
gacacgcgtg atggccttcg gcggggcgac agtggcaagg tggcgttccg acggcggggc 10860
aatggaaagg cggcgttccg aaggcatggc ggcggcaagg tggtgttctt gcaacgggca 10920
gggcggcggc gatgaagcga tgttccagcg gcgacgacga gcgagtggcg cgtcgagagg 10980
tttcaggtgg ttaccttgcc gccaaaagtt tatgggaaac aaaacttccc ctagacctga 11040
aggaagttgg ggtcaaattt acaagatcct gtaaatttat tgcacgttta cggggttttc 11100
tgaagatgat ttttggccta aattttactt tacggataca tgatctgtta gggttgctct 11160
taggccgggg cataaaagac gagagctcct agttgacgct cgtgcgtcaa ccaggtgtag 11220
atttgctaaa gcctacatga atgggcccac ttcaccaaga acttgttttt taaatatttt 11280
tctctttggt tatttagtca gtttttcagt gaaaaaccag ctattgtttt tcagaagata 11340
gaaaatttta taacctatta aaataggaaa atatattttc aaaaaaatta tattatatgt 11400
tttataaaag ttcaatgtat attaaaaaat tattcaccgt gtattaaaaa tatgttcggt 11460
ttgtataaag aatggtcatc atatgttaaa aaatgttcag tgtatataac aaaaatgttc 11520
atcatatata aaaatatata ttcagtttat gttttgaaga aaatttacct tatgctaaaa 11580
aaagttcagt gtgcttttga aaaatgttca ttgcatgcta aaataggttc agtgtgcatt 11640
tgaaaaagtt caccatatac tgcaaaccta taaaatgcaa aaatataaaa caaaagaaaa 11700
cacaacatct gagaaaagaa aacgtaaaaa acaaaataga agcagctagt taagtgggct 11760
ggcccatctg aaggttcctt cagtgaagtc tcccttatat cacgcactcg ggcgtcacct 11820
aggattgacc ataaaagacc gagtagagag gatggcat 11858
<210> 2
<211> 4731
<212> DNA
<213> 人工序列
<400> 2
atggagcagc agcagagctt cgaggagagg acgagcaaag atagccccga gaagatgggt 60
gacagtgcgc tccaccttgc agcaagggcc ggaaatgttg cccttgtgca gaagatcttt 120
gcggactgcg atccggagct ggtcgcggaa ttgactgacc atcagaacca ggacggtgag 180
acagcgctgt atgtttccgc ggagatgggg catgtcgagg ttgtgtgtga aattctgaag 240
gtttgtgatc tgcattcagc atttctcaag gcacacaaca gctttgatgc attccatatt 300
gcagcaaagc agggccatct agttgttttg caggagctac tgaaggcttt tcctgcatta 360
gctatgacaa caaattcagt aaatgccaca gctttataca ctgccacact tcactgccac 420
attggtattg tcaatcttct actggataca gacccaagac ttgccaggat tgcaagaaat 480
aatgggaaga cagctctgca tatagcagca aggctgggca atgtggaggt ggtagtgttg 540
ttgttgaata aagatccggc gactgttttc agaatagaca ggaagggaca aacagcagtg 600
cacatggctt ccaaaggcca caatgctgaa attctgcttg agctactgaa gcccgatgtc 660
tcagtaatcc atttggaaga taacaagggg gacaggccac tgcatgttgc aacgcggaag 720
ggaaaaacca ttatagttca gacgctaata tccattgaag agattgatat caatgcaatc 780
aatggagctg gagagaccgc ttttgccatt gcagagaaac tgggtaatga agagcttgta 840
aacatcctga gggaggctgg tggagtaacc gcagaagagc aagtaaatcc tccgaaatca 900
atcaagcgtt ttaagcaaac acatgatgtc caatcgcaga tcaagcaaaa gcgtcggaca 960
aatatgcatt tccacacgat caggaagagt agtcaaaagc tccacactga ggctccagtc 1020
tgcgctttgg cagacgccat gttcagactt cctgcaaagc tcgatgagct actgattagc 1080
cacgtccaca tgcttcctag gggtgcggag gatgagatac ctctcatcaa gcaagatctg 1140
gaagagataa tggccattct gcaggagcac gaccacccag ggagagcgga agaccgtgct 1200
atgacgagca agtgcctgac caaggaggtg cgcgagctgt catacgacat ggaggatagt 1260
gtcgaccagt atgtgcacgc cgtcgacacc aagagaagga ttgttcctcg ccgtaaaaag 1320
tacaagatca cctgtcgtag gggcaagacc actgcgcggc tcccggagaa gcttaagtgg 1380
cgtatatgga tggccaacaa gatcagggag ttcagtgtgc gctcgcaaga ggcgctgcag 1440
cggtacagcc tatttaacca ccctggtgct catgacatca gcacgtctgc tacttctacg 1500
agacatgatg tgtgttttgg ctcttggtat cccacaccgt gtggggagct tgtcggtata 1560
gatggacatt tgaatactct tgaagcgtgg ttgggtaagg atggggagca gcagctcaag 1620
gtggtatctg ttgttggatc tggaggggtt ggtaagacca cactttccaa agagctgtac 1680
cgtagaatca gagggcaatt cgagtgccag gcatttgtga ggacgtcccg gaagcccgac 1740
atcaggaggc ttctcatcag cttgctctca caagtccggc cacaccaaac ccctcacact 1800
tggaaattgc atagtctaat tgccgatatc aggacacatc tccacgataa gaggtacttg 1860
atcgtcattg atgatgtatg ggctacacaa acatgggata tcattaatcg tgctttgccg 1920
gctggtaatc tttgcagtag aattctaata acaacagaag tcgaagatgt agctctgaaa 1980
tgttgtggtt atgactctag gcatgttctt atggtgaaac cacttggtta cgatgattca 2040
agcaaattat ttttcagcac agcttttgga ctacaatatg aatgtcctcc agaactctgt 2100
gacgctgcac acaacattgt gaggaaatgt gctggttcac cactagcaat ggttactgtg 2160
gctagtcttt tagtaagcca gattggcaaa ccagagaaat gggattatgt aaatgaaatc 2220
ttaggtcacg gtttgagcac atatcctagc tcggaaggaa tgaaacaagt actaaacctt 2280
agttacaaca atcttcctca ttatttgaag gcatgtgtga tgtatctcag tatatatgaa 2340
gaggactaca taattcagaa agatgatttg gtaaagcaat ggatagctga aggtcttatc 2400
ctagcaacag aagagaaaga caaagaggaa atatcaagga gatattttga tgagcttatc 2460
agtagcagaa tgatcctacc tgtgtataca aatgacaacg atgatgtttt gtcctgcaca 2520
ctgcatcaca tggtacttga ttttatcaaa cacaagtcct tagaagagaa ttttgtcatc 2580
ccaatagatc atagtcagac aactgcacca ctcgcagaca aggttcgtcg actgtctctc 2640
cactttggta attcagaagc aacgccacca acaaatatga gactatcaca agttcggact 2700
ctcgcatttt tcggggtcat tgagtgtttg ccttccgtta tagagtttcg gcttcttcaa 2760
gtcctaatcc tacatctttt tggcgatgat gaaagtgtca gttttgatct cactggaata 2820
tctgagcttt ttcggttgag atatttgcat gtcacatgta atgccacctt agaagtacca 2880
caaactcaga tgcgaggttt acaatatttg gagacactga aaatagatgc aagagtaagt 2940
gcagttccgt cggacattgt tcatttgccg agcttgttgc acctcagtct tcctgttggg 3000
acaaatctac caaatggtat tgaccatatg acatcgcttt gcacacttga atattttgat 3060
ataaatgtta actcaatgga gaatgtgcac agccttggtg agctgaccaa tcttcaggat 3120
cttcggctca catgttctac agttccttct tcttacttaa agagtaaaat cgatagtatg 3180
ggctctattc ttgcgaacct cagcaacctc aggtctgtaa ctctgaagtc ttcaggtatt 3240
ctggagagtg aaccttacag catgatcatt tcctgtgatg gcttgagcag cgtttcctct 3300
cctccagctc ttcttcaaag atttgagtgg ttgccacgca tttgtacctt ctccagcatc 3360
cctaagtgga ttagccatct caacaagctc tgcattttaa agattgggct tagggaatta 3420
gtgagcaatg atgtcgctgc tctgagagga ttgcctgcac taactgtttt gtcgttatat 3480
gtccgagcaa agcccgcaga aaaaattgtc tttagtaggg caggattctt ggttctcaag 3540
tgcttcaagt tcaggtgcag tgtaccttgg ctggaatttg aggtggatgc aatgcctaat 3600
ctcttgaaac tcaagctaag tttcgatgcc catggagtag atcaacatgg tactatacct 3660
gtcggcatag tgcacttaac aggccttaag gaaatctcta caaaaatttg gggtgctggt 3720
tacaatgaaa gaagggctgc aaaatcagcg ctgattgccg ctattaagat gcattcggga 3780
tgtcccaccg ccagcataca gtgtttagat gggatgttca gtggtaagga tgataataat 3840
agcgggatac aagaggaaga acacatggct ctgcaaaagc agcaatacaa tatcatggag 3900
gaagacttga tggaacagca taaagtccgt caaaaaggca cgtgggaaga tgctcacaga 3960
aggttttctg cattttcaga ttggaaagct gttgcggaca agcaacctaa cagtagagtg 4020
caccatccgt gccgggttgg gaagacgacc ggactgtcaa catcaaggaa gtggacagca 4080
tggctgaaga cagagacaca gaaatacaag agcatggata aagactccaa taaacagcat 4140
ggaggtcttc caaaggacca catggatgtt gcagacaaac aagctgccag tagcaacaat 4200
cgtaggattc ccaggcggac caggaagatg agtagtcgga ggatgcaggt gagggtagga 4260
tccttgcagg acttgggttt cctcgacgat ggcttcagct ggaggaagta cggccagaag 4320
gataccatgg gctccacgta cccaagagct tatttccggt gcacgcacag gcacgttaag 4380
ggctgcctag cgaccaagca ggtgcagcgc acgtctaacg acccgctgct ctttgacgtg 4440
gtgcataacg gggagcacac gtgcttggac tctgtcggat cccctgcgac gtccgggagc 4500
caggttacct acgaggggat aaatgcaacc gcggcaacgt cttgtggcca ggtcaccttg 4560
gaggaggtga gcagaggtac gcccggagtt ggattcatgt ccccggcggt gttcagtagc 4620
caggttatgt ccgaggtggt aagcggaagc gggagtacgg cgggattttc tggtgatgag 4680
gttggcatgc cagacccgga ccgcgatggc tacgaatttg accgagtttg a 4731
<210> 3
<211> 1576
<212> PRT
<213> 人工序列
<400> 3
Met Glu Gln Gln Gln Ser Phe Glu Glu Arg Thr Ser Lys Asp Ser Pro
1 5 10 15
Glu Lys Met Gly Asp Ser Ala Leu His Leu Ala Ala Arg Ala Gly Asn
20 25 30
Val Ala Leu Val Gln Lys Ile Phe Ala Asp Cys Asp Pro Glu Leu Val
35 40 45
Ala Glu Leu Thr Asp His Gln Asn Gln Asp Gly Glu Thr Ala Leu Tyr
50 55 60
Val Ser Ala Glu Met Gly His Val Glu Val Val Cys Glu Ile Leu Lys
65 70 75 80
Val Cys Asp Leu His Ser Ala Phe Leu Lys Ala His Asn Ser Phe Asp
85 90 95
Ala Phe His Ile Ala Ala Lys Gln Gly His Leu Val Val Leu Gln Glu
100 105 110
Leu Leu Lys Ala Phe Pro Ala Leu Ala Met Thr Thr Asn Ser Val Asn
115 120 125
Ala Thr Ala Leu Tyr Thr Ala Thr Leu His Cys His Ile Gly Ile Val
130 135 140
Asn Leu Leu Leu Asp Thr Asp Pro Arg Leu Ala Arg Ile Ala Arg Asn
145 150 155 160
Asn Gly Lys Thr Ala Leu His Ile Ala Ala Arg Leu Gly Asn Val Glu
165 170 175
Val Val Val Leu Leu Leu Asn Lys Asp Pro Ala Thr Val Phe Arg Ile
180 185 190
Asp Arg Lys Gly Gln Thr Ala Val His Met Ala Ser Lys Gly His Asn
195 200 205
Ala Glu Ile Leu Leu Glu Leu Leu Lys Pro Asp Val Ser Val Ile His
210 215 220
Leu Glu Asp Asn Lys Gly Asp Arg Pro Leu His Val Ala Thr Arg Lys
225 230 235 240
Gly Lys Thr Ile Ile Val Gln Thr Leu Ile Ser Ile Glu Glu Ile Asp
245 250 255
Ile Asn Ala Ile Asn Gly Ala Gly Glu Thr Ala Phe Ala Ile Ala Glu
260 265 270
Lys Leu Gly Asn Glu Glu Leu Val Asn Ile Leu Arg Glu Ala Gly Gly
275 280 285
Val Thr Ala Glu Glu Gln Val Asn Pro Pro Lys Ser Ile Lys Arg Phe
290 295 300
Lys Gln Thr His Asp Val Gln Ser Gln Ile Lys Gln Lys Arg Arg Thr
305 310 315 320
Asn Met His Phe His Thr Ile Arg Lys Ser Ser Gln Lys Leu His Thr
325 330 335
Glu Ala Pro Val Cys Ala Leu Ala Asp Ala Met Phe Arg Leu Pro Ala
340 345 350
Lys Leu Asp Glu Leu Leu Ile Ser His Val His Met Leu Pro Arg Gly
355 360 365
Ala Glu Asp Glu Ile Pro Leu Ile Lys Gln Asp Leu Glu Glu Ile Met
370 375 380
Ala Ile Leu Gln Glu His Asp His Pro Gly Arg Ala Glu Asp Arg Ala
385 390 395 400
Met Thr Ser Lys Cys Leu Thr Lys Glu Val Arg Glu Leu Ser Tyr Asp
405 410 415
Met Glu Asp Ser Val Asp Gln Tyr Val His Ala Val Asp Thr Lys Arg
420 425 430
Arg Ile Val Pro Arg Arg Lys Lys Tyr Lys Ile Thr Cys Arg Arg Gly
435 440 445
Lys Thr Thr Ala Arg Leu Pro Glu Lys Leu Lys Trp Arg Ile Trp Met
450 455 460
Ala Asn Lys Ile Arg Glu Phe Ser Val Arg Ser Gln Glu Ala Leu Gln
465 470 475 480
Arg Tyr Ser Leu Phe Asn His Pro Gly Ala His Asp Ile Ser Thr Ser
485 490 495
Ala Thr Ser Thr Arg His Asp Val Cys Phe Gly Ser Trp Tyr Pro Thr
500 505 510
Pro Cys Gly Glu Leu Val Gly Ile Asp Gly His Leu Asn Thr Leu Glu
515 520 525
Ala Trp Leu Gly Lys Asp Gly Glu Gln Gln Leu Lys Val Val Ser Val
530 535 540
Val Gly Ser Gly Gly Val Gly Lys Thr Thr Leu Ser Lys Glu Leu Tyr
545 550 555 560
Arg Arg Ile Arg Gly Gln Phe Glu Cys Gln Ala Phe Val Arg Thr Ser
565 570 575
Arg Lys Pro Asp Ile Arg Arg Leu Leu Ile Ser Leu Leu Ser Gln Val
580 585 590
Arg Pro His Gln Thr Pro His Thr Trp Lys Leu His Ser Leu Ile Ala
595 600 605
Asp Ile Arg Thr His Leu His Asp Lys Arg Tyr Leu Ile Val Ile Asp
610 615 620
Asp Val Trp Ala Thr Gln Thr Trp Asp Ile Ile Asn Arg Ala Leu Pro
625 630 635 640
Ala Gly Asn Leu Cys Ser Arg Ile Leu Ile Thr Thr Glu Val Glu Asp
645 650 655
Val Ala Leu Lys Cys Cys Gly Tyr Asp Ser Arg His Val Leu Met Val
660 665 670
Lys Pro Leu Gly Tyr Asp Asp Ser Ser Lys Leu Phe Phe Ser Thr Ala
675 680 685
Phe Gly Leu Gln Tyr Glu Cys Pro Pro Glu Leu Cys Asp Ala Ala His
690 695 700
Asn Ile Val Arg Lys Cys Ala Gly Ser Pro Leu Ala Met Val Thr Val
705 710 715 720
Ala Ser Leu Leu Val Ser Gln Ile Gly Lys Pro Glu Lys Trp Asp Tyr
725 730 735
Val Asn Glu Ile Leu Gly His Gly Leu Ser Thr Tyr Pro Ser Ser Glu
740 745 750
Gly Met Lys Gln Val Leu Asn Leu Ser Tyr Asn Asn Leu Pro His Tyr
755 760 765
Leu Lys Ala Cys Val Met Tyr Leu Ser Ile Tyr Glu Glu Asp Tyr Ile
770 775 780
Ile Gln Lys Asp Asp Leu Val Lys Gln Trp Ile Ala Glu Gly Leu Ile
785 790 795 800
Leu Ala Thr Glu Glu Lys Asp Lys Glu Glu Ile Ser Arg Arg Tyr Phe
805 810 815
Asp Glu Leu Ile Ser Ser Arg Met Ile Leu Pro Val Tyr Thr Asn Asp
820 825 830
Asn Asp Asp Val Leu Ser Cys Thr Leu His His Met Val Leu Asp Phe
835 840 845
Ile Lys His Lys Ser Leu Glu Glu Asn Phe Val Ile Pro Ile Asp His
850 855 860
Ser Gln Thr Thr Ala Pro Leu Ala Asp Lys Val Arg Arg Leu Ser Leu
865 870 875 880
His Phe Gly Asn Ser Glu Ala Thr Pro Pro Thr Asn Met Arg Leu Ser
885 890 895
Gln Val Arg Thr Leu Ala Phe Phe Gly Val Ile Glu Cys Leu Pro Ser
900 905 910
Val Ile Glu Phe Arg Leu Leu Gln Val Leu Ile Leu His Leu Phe Gly
915 920 925
Asp Asp Glu Ser Val Ser Phe Asp Leu Thr Gly Ile Ser Glu Leu Phe
930 935 940
Arg Leu Arg Tyr Leu His Val Thr Cys Asn Ala Thr Leu Glu Val Pro
945 950 955 960
Gln Thr Gln Met Arg Gly Leu Gln Tyr Leu Glu Thr Leu Lys Ile Asp
965 970 975
Ala Arg Val Ser Ala Val Pro Ser Asp Ile Val His Leu Pro Ser Leu
980 985 990
Leu His Leu Ser Leu Pro Val Gly Thr Asn Leu Pro Asn Gly Ile Asp
995 1000 1005
His Met Thr Ser Leu Cys Thr Leu Glu Tyr Phe Asp Ile Asn Val
1010 1015 1020
Asn Ser Met Glu Asn Val His Ser Leu Gly Glu Leu Thr Asn Leu
1025 1030 1035
Gln Asp Leu Arg Leu Thr Cys Ser Thr Val Pro Ser Ser Tyr Leu
1040 1045 1050
Lys Ser Lys Ile Asp Ser Met Gly Ser Ile Leu Ala Asn Leu Ser
1055 1060 1065
Asn Leu Arg Ser Val Thr Leu Lys Ser Ser Gly Ile Leu Glu Ser
1070 1075 1080
Glu Pro Tyr Ser Met Ile Ile Ser Cys Asp Gly Leu Ser Ser Val
1085 1090 1095
Ser Ser Pro Pro Ala Leu Leu Gln Arg Phe Glu Trp Leu Pro Arg
1100 1105 1110
Ile Cys Thr Phe Ser Ser Ile Pro Lys Trp Ile Ser His Leu Asn
1115 1120 1125
Lys Leu Cys Ile Leu Lys Ile Gly Leu Arg Glu Leu Val Ser Asn
1130 1135 1140
Asp Val Ala Ala Leu Arg Gly Leu Pro Ala Leu Thr Val Leu Ser
1145 1150 1155
Leu Tyr Val Arg Ala Lys Pro Ala Glu Lys Ile Val Phe Ser Arg
1160 1165 1170
Ala Gly Phe Leu Val Leu Lys Cys Phe Lys Phe Arg Cys Ser Val
1175 1180 1185
Pro Trp Leu Glu Phe Glu Val Asp Ala Met Pro Asn Leu Leu Lys
1190 1195 1200
Leu Lys Leu Ser Phe Asp Ala His Gly Val Asp Gln His Gly Thr
1205 1210 1215
Ile Pro Val Gly Ile Val His Leu Thr Gly Leu Lys Glu Ile Ser
1220 1225 1230
Thr Lys Ile Trp Gly Ala Gly Tyr Asn Glu Arg Arg Ala Ala Lys
1235 1240 1245
Ser Ala Leu Ile Ala Ala Ile Lys Met His Ser Gly Cys Pro Thr
1250 1255 1260
Ala Ser Ile Gln Cys Leu Asp Gly Met Phe Ser Gly Lys Asp Asp
1265 1270 1275
Asn Asn Ser Gly Ile Gln Glu Glu Glu His Met Ala Leu Gln Lys
1280 1285 1290
Gln Gln Tyr Asn Ile Met Glu Glu Asp Leu Met Glu Gln His Lys
1295 1300 1305
Val Arg Gln Lys Gly Thr Trp Glu Asp Ala His Arg Arg Phe Ser
1310 1315 1320
Ala Phe Ser Asp Trp Lys Ala Val Ala Asp Lys Gln Pro Asn Ser
1325 1330 1335
Arg Val His His Pro Cys Arg Val Gly Lys Thr Thr Gly Leu Ser
1340 1345 1350
Thr Ser Arg Lys Trp Thr Ala Trp Leu Lys Thr Glu Thr Gln Lys
1355 1360 1365
Tyr Lys Ser Met Asp Lys Asp Ser Asn Lys Gln His Gly Gly Leu
1370 1375 1380
Pro Lys Asp His Met Asp Val Ala Asp Lys Gln Ala Ala Ser Ser
1385 1390 1395
Asn Asn Arg Arg Ile Pro Arg Arg Thr Arg Lys Met Ser Ser Arg
1400 1405 1410
Arg Met Gln Val Arg Val Gly Ser Leu Gln Asp Leu Gly Phe Leu
1415 1420 1425
Asp Asp Gly Phe Ser Trp Arg Lys Tyr Gly Gln Lys Asp Thr Met
1430 1435 1440
Gly Ser Thr Tyr Pro Arg Ala Tyr Phe Arg Cys Thr His Arg His
1445 1450 1455
Val Lys Gly Cys Leu Ala Thr Lys Gln Val Gln Arg Thr Ser Asn
1460 1465 1470
Asp Pro Leu Leu Phe Asp Val Val His Asn Gly Glu His Thr Cys
1475 1480 1485
Leu Asp Ser Val Gly Ser Pro Ala Thr Ser Gly Ser Gln Val Thr
1490 1495 1500
Tyr Glu Gly Ile Asn Ala Thr Ala Ala Thr Ser Cys Gly Gln Val
1505 1510 1515
Thr Leu Glu Glu Val Ser Arg Gly Thr Pro Gly Val Gly Phe Met
1520 1525 1530
Ser Pro Ala Val Phe Ser Ser Gln Val Met Ser Glu Val Val Ser
1535 1540 1545
Gly Ser Gly Ser Thr Ala Gly Phe Ser Gly Asp Glu Val Gly Met
1550 1555 1560
Pro Asp Pro Asp Arg Asp Gly Tyr Glu Phe Asp Arg Val
1565 1570 1575
<210> 4
<211> 247
<212> DNA
<213> 人工序列
<400> 4
gcacactgca tcacatggta cttgatttta tcaaacacaa gtccttagaa gagaattttg 60
tcatcccaat agatcatagt cagacaactg caccactcgc agacaaggtt cgtcgactgt 120
ctctccactt tggtaattca gaagcaacgc caccaacaaa tatgagacta tcacaagttc 180
ggactctcgc atttttcggg gtcattgagt gtttgccttc cgttatagag tttcggcttc 240
ttcaagt 247
<210> 5
<211> 41
<212> DNA
<213> 人工序列
<400> 5
ccggggatcc tctagagtga tgtttgatag cacacaaggt a 41
<210> 6
<211> 45
<212> DNA
<213> 人工序列
<400> 6
ccaagcttgc atgcctgcag atgccatcct ctctactcgg tcttt 45
<210> 7
<211> 25
<212> DNA
<213> 人工序列
<400> 7
atggagcagc agcagagctt cgagg 25
<210> 8
<211> 25
<212> DNA
<213> 人工序列
<400> 8
ttacgtgcta acattatcaa gtgga 25
<210> 9
<211> 20
<212> DNA
<213> 人工序列
<400> 9
aaggaggtgc gcgagctgtc 20
<210> 10
<211> 20
<212> DNA
<213> 人工序列
<400> 10
tgacaagtgc tcgatgccaa 20
<210> 11
<211> 20
<212> DNA
<213> 人工序列
<400> 11
gcagagcctg gttcaaaaag 20
<210> 12
<211> 20
<212> DNA
<213> 人工序列
<400> 12
cgcctctagc gagagctatg 20
<210> 13
<211> 20
<212> DNA
<213> 人工序列
<400> 13
gaaaccagca aagaaagaaa 20
<210> 14
<211> 20
<212> DNA
<213> 人工序列
<400> 14
cgtagcagtc atgcaattta 20
<210> 15
<211> 20
<212> DNA
<213> 人工序列
<400> 15
gttgctgcaa acaacaataa 20
<210> 16
<211> 20
<212> DNA
<213> 人工序列
<400> 16
atgtagatcg caacccatag 20
<210> 17
<211> 20
<212> DNA
<213> 人工序列
<400> 17
ctagaacctt ttcagaaccg 20
<210> 18
<211> 21
<212> DNA
<213> 人工序列
<400> 18
gaaaaagaaa aatgttcacg a 21
<210> 19
<211> 20
<212> DNA
<213> 人工序列
<400> 19
aaaacaaata atggtgaccg 20
<210> 20
<211> 20
<212> DNA
<213> 人工序列
<400> 20
agccagaaat atgctgatgt 20
<210> 21
<211> 20
<212> DNA
<213> 人工序列
<400> 21
atgtttgtaa ccgactctgg 20
<210> 22
<211> 20
<212> DNA
<213> 人工序列
<400> 22
ttttcactag caaaagggtc 20
<210> 23
<211> 20
<212> DNA
<213> 人工序列
<400> 23
caaccaccca gtattgttct 20
<210> 24
<211> 20
<212> DNA
<213> 人工序列
<400> 24
agctgcagac ttttaacgac 20
<210> 25
<211> 22
<212> DNA
<213> 人工序列
<400> 25
gtttgtagca gacaacaaca ac 22
<210> 26
<211> 22
<212> DNA
<213> 人工序列
<400> 26
accaaagagt tagctatgga ca 22
<210> 27
<211> 20
<212> DNA
<213> 人工序列
<400> 27
ggatctcaga catgtacgct 20
<210> 28
<211> 20
<212> DNA
<213> 人工序列
<400> 28
cattactagg taccggcttg 20
<210> 29
<211> 20
<212> DNA
<213> 人工序列
<400> 29
aactgctcaa gagctgtagg 20
<210> 30
<211> 20
<212> DNA
<213> 人工序列
<400> 30
aggaacctgc ttacaaatca 20
<210> 31
<211> 20
<212> DNA
<213> 人工序列
<400> 31
tctttgttca caaatcacga 20
<210> 32
<211> 20
<212> DNA
<213> 人工序列
<400> 32
aatatgacca agcctgagtg 20
<210> 33
<211> 20
<212> DNA
<213> 人工序列
<400> 33
gtaggagtac cacgcaacat 20
<210> 34
<211> 20
<212> DNA
<213> 人工序列
<400> 34
gtaatttgtt tatccaccgc 20
<210> 35
<211> 22
<212> DNA
<213> 人工序列
<400> 35
atgtgtatgt gtgaatggtt gt 22
<210> 36
<211> 22
<212> DNA
<213> 人工序列
<400> 36
gttttaggat tttaggggtt tt 22
<210> 37
<211> 20
<212> DNA
<213> 人工序列
<400> 37
tgctgtgttg ctgagataag 20
<210> 38
<211> 20
<212> DNA
<213> 人工序列
<400> 38
tatgttacac ccaccggtat 20
<210> 39
<211> 20
<212> DNA
<213> 人工序列
<400> 39
gctgctcctg ctctagtaaa 20
<210> 40
<211> 20
<212> DNA
<213> 人工序列
<400> 40
aagtaataag cgcaccaaac 20
<210> 41
<211> 20
<212> DNA
<213> 人工序列
<400> 41
aggagagcgg caaggactac 20
<210> 42
<211> 20
<212> DNA
<213> 人工序列
<400> 42
gagaatcaca ggaaggcacg 20
<210> 43
<211> 20
<212> DNA
<213> 人工序列
<400> 43
aatccatcga tagcaagaga 20
<210> 44
<211> 20
<212> DNA
<213> 人工序列
<400> 44
tcattcatcc aaaacatcaa 20
<210> 45
<211> 20
<212> DNA
<213> 人工序列
<400> 45
ggtcaatcaa cgaccagtat 20
<210> 46
<211> 20
<212> DNA
<213> 人工序列
<400> 46
acacaagaag aaaacggaaa 20
<210> 47
<211> 20
<212> DNA
<213> 人工序列
<400> 47
gttctgcatt acctggaaag 20
<210> 48
<211> 20
<212> DNA
<213> 人工序列
<400> 48
ccaattatgg ttggtgttct 20
<210> 49
<211> 20
<212> DNA
<213> 人工序列
<400> 49
tcgatggtca aacctaaaat 20
<210> 50
<211> 20
<212> DNA
<213> 人工序列
<400> 50
acacttttgc aggttgaaat 20
<210> 51
<211> 20
<212> DNA
<213> 人工序列
<400> 51
cacctgttga gaggaagaag 20
<210> 52
<211> 20
<212> DNA
<213> 人工序列
<400> 52
agaatgccaa actgattcat 20
<210> 53
<211> 24
<212> DNA
<213> 人工序列
<400> 53
agaaccatcc gaataggccg agac 24
<210> 54
<211> 23
<212> DNA
<213> 人工序列
<400> 54
agaggtggat tgggcatgga aac 23
<210> 55
<211> 21
<212> DNA
<213> 人工序列
<400> 55
cggtggtggt gaagatttga a 21
<210> 56
<211> 21
<212> DNA
<213> 人工序列
<400> 56
gacgaacctt gtctgcgagt g 21
<210> 57
<211> 24
<212> DNA
<213> 人工序列
<400> 57
ggcaaagcaa tggatagccg aagg 24
<210> 58
<211> 23
<212> DNA
<213> 人工序列
<400> 58
tgcaacaagc tgggcaaatg aac 23
<210> 59
<211> 21
<212> DNA
<213> 人工序列
<400> 59
cgaggacacc aaccagatag c 21
<210> 60
<211> 21
<212> DNA
<213> 人工序列
<400> 60
ccaggcagaa agacaggaaa a 21
<210> 61
<211> 20
<212> DNA
<213> 人工序列
<400> 61
ttggatgggc atttttccgc 20
<210> 62
<211> 21
<212> DNA
<213> 人工序列
<400> 62
tccaagatac aagtgtccgg c 21
<210> 63
<211> 25
<212> DNA
<213> 人工序列
<400> 63
gtcccttgcc aacctgtcag gatag 25
<210> 64
<211> 25
<212> DNA
<213> 人工序列
<400> 64
ctgcatgcca ttcttcagaa gctaa 25
<210> 65
<211> 19
<212> DNA
<213> 人工序列
<400> 65
tttgttacaa cccaggggg 19
<210> 66
<211> 20
<212> DNA
<213> 人工序列
<400> 66
ttgtgtggcg aaagaaacag 20
<210> 67
<211> 20
<212> DNA
<213> 人工序列
<400> 67
gcatagcatc gcatatgcat 20
<210> 68
<211> 20
<212> DNA
<213> 人工序列
<400> 68
gccacgcttg gacaagatat 20
<210> 69
<211> 25
<212> DNA
<213> 人工序列
<400> 69
agagtgcacc atccgtgccg ggttg 25
<210> 70
<211> 25
<212> DNA
<213> 人工序列
<400> 70
actcccgctt ccgcttacca cctcg 25
<210> 71
<211> 23
<212> DNA
<213> 人工序列
<400> 71
tgggaagatg ctcacagaag gta 23
<210> 72
<211> 23
<212> DNA
<213> 人工序列
<400> 72
ctcaaactcg gtcaaattcg tag 23
<210> 73
<211> 41
<212> DNA
<213> 人工序列
<400> 73
tttttttttt ttttagctag cgcacactgc atcacatggt a 41
<210> 74
<211> 41
<212> DNA
<213> 人工序列
<400> 74
gattcttctt ccgttgctag cacttgaaga agccgaaact c 41
<210> 75
<211> 25
<212> DNA
<213> 人工序列
<400> 75
tgctcttgta tttctgtgtc tctgt 25
<210> 76
<211> 25
<212> DNA
<213> 人工序列
<400> 76
cgtcaaaaag gcacgtggga agatg 25
<210> 77
<211> 21
<212> DNA
<213> 人工序列
<400> 77
gctactgaag cccgatgtct c 21
<210> 78
<211> 20
<212> DNA
<213> 人工序列
<400> 78
tctccagctc cattgattgc 20
Claims (5)
1.小麦条锈病抗性基因YrU1,其特征在于:其包括自身启动子和5′-UTR、3′-UTR在内的基因全长的核苷酸序列如SEQ ID NO.1所示。
2.根据权利按要求1所述的小麦条锈病抗性基因YrU1,其特征在于:其编码区的核苷酸序列如SEQ ID NO.2所示。
3.由权利要求2所述的小麦条锈病抗性基因YrU1编码的蛋白,其特征在于:其氨基酸序列如SEQ ID NO.3 所示。
4.含有如权利要求1所述小麦条锈病抗性基因YrU1的载体。
5.如权利要求1所述小麦条锈病抗性基因YrU1在小麦抗性育种中的应用。
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CN111073898A true CN111073898A (zh) | 2020-04-28 |
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