CN112321694B - 小麦抗叶锈病蛋白及其编码基因和应用 - Google Patents
小麦抗叶锈病蛋白及其编码基因和应用 Download PDFInfo
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Abstract
本发明涉及生物技术领域,具体公开一种小麦抗叶锈病蛋白及其编码基因和应用。所述小麦抗叶锈病蛋白其具有如SEQ ID NO:1所示的氨基酸序列,或者具有如SEQ ID NO:1所述的氨基酸序列经缺失、替换或添加一个或多个氨基酸得到的具有提高小麦抗叶锈病功能的氨基酸序列,所述小麦抗叶锈病蛋白或编码其的基因可应用于调控或提高小麦抗叶锈病性能。本发明提供了小麦抗叶锈病蛋白,并具体公开了所述小麦抗叶锈病蛋白的氨基酸序列,在小麦中提高该小麦抗叶锈病蛋白的表达量能够显著提高小麦对叶锈病的抵抗能力,其对于小麦抗叶锈病育种具有重大的应用推广价值。
Description
技术领域
本发明涉及生物技术领域,尤其涉及一种小麦抗叶锈病蛋白及其编码基因和应用。
背景技术
小麦是世界范围内最重要的粮食作物之一,也是主要的食物来源。然而,几千年来小麦锈菌一直威胁着小麦生产,成为小麦生产的主要病害。小麦锈菌分为小麦叶锈菌(Puccinia triticinia)、小麦秆锈菌(P.graminis f.sp.tritici)和小麦条锈菌(P.striiformis f.sp.tritici)。其中由小麦叶锈菌引起的小麦叶锈病较为常见,在世界范围内发生最为广泛,是危害小麦产量的重要因素之一,病害严重流行年份可使小麦减产40%以上,甚至绝收。小麦叶锈病主要发生在长江流域、黄淮流域、华北麦区和东北麦区等,1969年、1973年、1975年和1979年该病害的大面积流行给我国小麦产量造成了严重损失。近几年,华北及黄淮麦区频频发生,尤其是2012年和2015年,小麦叶锈病在我国大部分小麦产区传播,造成了小麦生产的巨大损失。随着全球气候变暖,未来可能会加剧小麦叶锈病的发生和流行,选育和利用抗病品种是控制小麦叶锈病最经济、有效且对环境安全的防控措施。
目前,由于长期大面积种植具有单一或少数基因的抗病品种,对病原菌产生了选择压力。病原菌为了适应这种压力,通过无毒基因或其调控基因的变异,进化出了对这些品种致病的菌株,导致抗病品种抗性被克服。目前国内外已正式命名了79个抗叶锈病基因,这些基因多数为小种专化抗病基因,其随小种的毒性变异容易丧失抗性。在我国,目前只有少数抗病基因属于有效的抗锈病基因。为了控制小麦锈病,一方面需要加快抗病基因的挖掘,培育具有持久抗性品种的培育,另一方面需要利用已有的抗性资源,对含有不同抗病基因的品种进行轮换和布局,尽可能使病原菌群体毒性变异趋于稳定,以延长品种使用年限。因此定位和克隆小麦抗锈病基因并开发其功能标记,可以加速抗病品种的培育,同时为研究小麦抗病基因和锈菌无毒基因互作的分子机理奠定基础。
发明内容
针对现有小麦抗小麦叶锈菌的品种单一,在病菌长期变异过程中,其抗性容易被克服,且目前的抗叶锈病基因多数为小种专化抗病基因,其随小种的毒性变异容易丧失抗性的问题,本发明提供一种小麦抗叶锈病蛋白及其编码基因和应用。
为达到上述发明目的,本发明实施例采用了如下的技术方案:
小麦抗叶锈病蛋白,其具有如SEQ ID NO:1所示的氨基酸序列,或者具有如SEQ IDNO:1所述的氨基酸序列经缺失、替换或添加一个或多个氨基酸得到的具有提高小麦抗叶锈病功能的氨基酸序列。
相对于现有技术,本发明提供了一种小麦抗叶锈病蛋白,并具体公开了所述小麦抗叶锈病蛋白的氨基酸序列,在小麦中提高该小麦抗叶锈病蛋白的表达量能够显著提高小麦对叶锈病的抵抗能力,其对于小麦抗叶锈病育种具有重大的应用推广价值。
优选的,所述小麦抗叶锈病蛋白为在SEQ ID NO:1序列的N端或/和C端连接蛋白标签。
优选的,所述蛋白标签为序列为SEQ ID NO:2的Poly-Arg、序列为SEQ ID NO:3的Poly-His、序列为SEQ ID NO:4的FLAG、序列为SEQ ID NO:5的Strep-tag II、序列为SEQ IDNO:6的c-myc和序列为SEQ ID NO:7的HA中的任意一种。
本发明还提供编码所述小麦抗叶锈病蛋白的核酸分子。
本发明还提供包含所述核酸分子及其表达调控原件的重组表达载体,所述重组表达载体具体可为在现有植物表达载体中插入所述核酸分子得到的重组质粒。其中现有植物表达载体具体可为pTCK303载体。
本发明还提供包含所述核酸分子并进行表达的宿主细胞。
本发明还提供所述小麦抗叶锈病蛋白在调控或提高小麦抗叶锈病性能中的应用。
本发明还提供编码所述小麦抗叶锈病蛋白的核酸分子在抗叶锈病小麦品种遗传育种中的应用。
本发明还提供一种用于扩增编码得到所述小麦抗叶锈病蛋白的基因的引物,其上游引物为HBAU194F:5′-TCTAGAGGATCCCCGGGTACCATGGCCATGATCTTACCAGCC-3′;
下游引物为HBAU194R:5′-ATTCGAGCTCTCTAGAACTAGTTTAATCAGCCCGTGGCCAC-3′。
本发明提供的扩增编码所述小麦抗叶锈病蛋白的基因的引物可以特异性的扩增并准确得到编码小麦抗叶锈病蛋白的基因序列。
本发明还提供一种用于鉴定编码所述小麦抗叶锈病蛋白的基因的引物,其上游引物为BD1-L:5′-TCGTACACTGCATCCTTGATCT-3′;下游引物为BD1-R:5′-AGGCTGAAGACTCGTATTTGGA-3′。
本发明提供的用于鉴定编码所述小麦抗叶锈病蛋白的基因的引物可以特异性的结合到编码小麦抗叶锈病蛋白的基因上,准确检测出编码小麦抗叶锈病蛋白的基因,特异性较强;且上述引物的灵敏度高,检测准确率为100%。
附图说明
图1是本发明实施例中编码小麦抗叶锈病蛋白的基因在小麦品种周麦22(ZM)上接种叶锈菌后不同时间点的表达量柱形图;
图2是本发明实施例中转基因植株在T0代的抗叶锈病表型图,其中,“+”表示阳性,即含有Lr13基因,“-”表示阴性,即不含Lr13基因;
图3是本发明实施例中T1代转基因植株的抗叶锈病表型图,其中,“+”表示阳性,即含有Lr13基因,“-”表示阴性,即不含Lr13基因。
具体实施方式
为了使本发明的目的、技术方案及优点更加清楚明白,以下结合实施例,对本发明进行进一步详细说明。应当理解,此处所描述的具体实施例仅用以解释本发明,并不用于限定本发明。
实施例1
1、材料
普通小麦品种周麦22(ZM):高抗叶锈病的中国小麦地方品种;
小麦品种郑州5389(ZZ):为高感叶锈病品种;
小麦品种Fielder:为叶锈病感病材料品种;
叶锈病致病真菌:叶锈病菌(Puccinia triticinia),FHDS生理小种。
2、小麦抗叶锈病的分级标准
根据表型,植株的叶锈病抗病性侵染型采用0-4级的分级标准。“0”为免疫,“;”为近免疫,“1”为高度抗病,“2”为中度抗病,“3”为中度感病,“4”为高度感病。按大分类来说,0-2为抗病,3-4为感病。各个分级标准相应的表型见表1。
表1
3、蛋白及其编码基因序列的获得
课题组先期在周麦22中定位了一个苗期温敏抗叶锈病新基因LrZH22,距离两侧最近的SSR标记为Xbarc55 and Xgwm374,距离LrZH22分别为2.4cM和4.8cM。进一步利用标记检测7213个周麦22/郑州5389的F2:3家系,获得332个重组家系。结合重组体的表型数据并利用中国春参考序列及混池转录组测序(BSR-Seq)方法设计特异标记引物对LrZH22进行精细定位,构建了LrZH22所在区域的精细遗传连锁图谱,将LrZH22定位到约70kb的物理区段。该区间有2个注释基因,其中1个是典型的与抗病相关的NBS-LRR蛋白。同时选取抗病亲本周麦22和8个纯感EMS突变家系进行MutRenSeq,结果将抗病基因LrZH22成功定位在2BS的特定物理区间,该结果与精细定位的NBS-LRR类抗病基因结果一致。利用诺唯赞高保真DNA聚合酶,通过PCR扩增普通小麦品种周麦22及小麦感病品种郑州5389的候选蛋白基因序列,其表达的蛋白序列如SEQ ID NO:1所示,将该蛋白命名为Lr13蛋白。周麦22的cDNA中编码Lr13蛋白的开放阅读框如SEQ ID NO:8所示。周麦22的基因组DNA中,编码Lr13蛋白的基因序列如SEQID NO:9所示(具有4个外显子)。
4、Lr13基因在不同接种时间的表达分析
供试植株为普通小麦周麦22(ZM)植株。将供试植株培养至一叶一心期,然后接种叶锈菌生理小种FHDS。在恒定温度22度条件下,分别于接种0h、6h、12h、24h、36h、48h和72h后,取植株的第一片叶片,检测Lr13基因的相对表达水平。
接种方式:将感染叶锈病致病菌并充分发病的感病小麦材料郑州5389(ZZ)植株在供试植株上方采用人工扫抹法进行接菌。
检测Lr13基因的相对表达水平的方法:提取供试植株叶片的总RNA,以Actin基因为内参基因,通过qRT-PCR检测Lr13基因的相对表达水平。
用于鉴定Lr13基因表达量的引物如下:
BD1-L(SEQ ID NO:10):5′-TCGTACACTGCATCCTTGATCT-3′;
BD1-R(SEQ ID NO:11):5′-AGGCTGAAGACTCGTATTTGGA-3′。
检测结果如图1所示,Lr13基因的表达受到叶锈致病菌的诱导,接种叶锈菌6h和36h表达量达到最高。
4、转基因验证Lr13蛋白功能
4.1、构建重组质粒
(1)取小麦周麦22的叶片,提取总RNA。
(2)将步骤(1)得到的RNA进行反转录,获得cDNA,以该cDNA作为模板,通过设计特异性引物HBAU194F(SEQ ID NO:12)和HBAU194R(SEQ ID NO:13),用高保真DNA聚合酶对Lr13的全长基因编码区进行扩增,回收PCR扩增产物。
HBAU194F:5′-TCTAGAGGATCCCCGGGTACCATGGCCATGATCTTACCAGCC-3′;
HBAU194R:5′-ATTCGAGCTCTCTAGAACTAGTTTAATCAGCCCGTGGCCAC-3′。
(3)用限制性内切酶KpnI和SpeI分别对步骤(2)回收的PCR扩增产物和pTCK303载体进行双酶切,回收酶切后的PCR扩增产物以及载体骨架。
(4)将酶切后的PCR扩增产物以及载体骨架用pEASY-Uni Seamless Cloning andAssembly Kit(CU101-01)同源重组试剂盒进行重组,得到重组质粒。该重组质粒转化大肠杆菌后,挑选阳性克隆进行测序,结果表明,Lr13基因的CDS片段正确连接到pTPCK303-Hyg载体上,说明成功构建了Lr13的过表达载体pTPCK303-Hyg-Lr13。
4.2、构建转基因植株
(1)将4.1中得到的重组质粒pTPCK303-Hyg-Lr13导入农杆菌EHA105中,得到重组农杆菌。
(2)取步骤(1)得到的重组农杆菌,对小麦Fielder的胚性愈伤组织进行遗传转化,培养获得植株,即为T0代植株。
(3)T0代植株自交获得籽粒,将籽粒培育为植株,即为T1代植株。
(4)从T1代植株中筛选转基因植株。
筛选转基因植株的方法:提取植株叶片的基因组DNA,以基因组DNA为模板,采用22-5OE-1L/R引物对进行PCR扩增来验证转基因植株。
22-5OE-1L(SEQ ID NO:16):5′-CTTCTGCAGGTCGACTCTAG-3′;
22-5OE-1R(SEQ ID NO:17):5′-TAATTATTGGGCTCATCACAGTC-3′。
具体验证方法为:
(1)取转基因T0和T1代小麦材料的叶片,提取DNA。
(2)分别以其DNA作为模板,通过设计的特异性引物22-5OE-1L/R,用高保真DNA聚合酶对转基因后代植株进行扩增,PCR产物经1%琼脂糖凝胶电泳检测。如果得到的PCR扩增产物中含有约518bp的DNA片段,则该小麦含有Lr13基因;否则该小麦不含有Lr13基因。4.3、制备转空载体植株
用pTCK303空载体代替重组质粒pTPCK303-Hyg-Lr13,参照4.2中的步骤(1)-(2)进行操作,得到转空载体植株。
4.4、Lr13蛋白的叶锈病抗病性鉴定
在4.2中的第一批转基因植株中,获得7棵T0代转基因植株,在其长至一叶一心时对其接菌叶锈生理小种FHDS,当充分发病后(10天后),进行表型鉴定和PCR表达检测。发现3株阳性转基因苗与抗病材料周麦22表现抗性,参见图2,其中3株阳性转基因苗分别为T0-1、T0-2和T0-3,并分别获得相应T1代种子,同样对其进行接菌表型鉴定。
T1代转基因植株与周麦22植株表型一致,表现出明显的叶锈病抗病性,如图3所示。转空载体的植株和小麦Fielder植株表型一致,表型出明显的叶锈病感病性。T0-1、T0-2和T0-3的抗病等级均为2级。周麦22的抗病等级为1或2级。转空载体植株和小麦Fielder植株的抗病等级均为4级。
对T0-1、T0-2和T0-3和其获得的T1代植株的基因组DNA用22-5OE-1L/R引物进行PCR扩增验证,PCR扩增产物中含有518bp的DNA片段,而空载体植株和小麦Fielder的基因组的PCR扩增产物中未扩增得到518bp的DNA片段。即T0-1、T0-2和T0-3和其获得的T1代小麦中均含有Lr13基因,与该小麦表现出的抗叶锈病表型相一致。
SEQUENCE LISTING
<110> 河北农业大学
<120> 一种小麦抗叶锈病蛋白及其编码基因和应用
<130> 2020
<160> 15
<170> PatentIn version 3.5
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Arg Arg Arg Arg Arg
1 5
<210> 3
<211> 6
<212> PRT
<213> Poly-His
<400> 3
His His His His His His
1 5
<210> 4
<211> 8
<212> PRT
<213> FLAG
<400> 4
Asp Tyr Lys Asp Asp Asp Asp Lys
1 5
<210> 5
<211> 8
<212> PRT
<213> Strep-tag II
<400> 5
Trp Ser His Pro Gln Phe Glu Lys
1 5
<210> 6
<211> 10
<212> PRT
<213> c-myc
<400> 6
Glu Gln Lys Leu Ile Ser Glu Glu Asp Leu
1 5 10
<210> 7
<211> 9
<212> PRT
<213> HA
<400> 7
Tyr Pro Tyr Asp Val Pro Asp Tyr Ala
1 5
<210> 8
<211> 3219
<212> DNA
<213> Lr13编码区序列
<400> 8
atggccatga tcttaccagc ccttcctgta gctgttgagg taatttccag tgtcaatgca 60
ttgaacgaat tcgtcggctg ggcgacacca cacatgaaag cggccatacg ttatgcacgg 120
aacattccac ggtcaccaga tgaagagaac acagctgcac aacgccagat tcaagagaaa 180
attgagcagc taggcgataa gcttgggagg ctgaagactc gtatttggaa gctgaagaca 240
actatgccca agatgcttga cctcattgac agggttgagt ggcaaagcca caaggaacaa 300
gccactgttc tactcccgga gatcaaggat gcagtgtacg atgcagagga tctgcttgat 360
gagtttgatt attatgcgct caagttgaag gttgagtgca acaaaaattt ggggcaagat 420
cacttgcata atgccttcat ggaattctta aacactatag actgtgatga gcccaataat 480
tacaccacga aagtcaatga tattcaagaa agactagatc atctgaatca acagtccaaa 540
gatatgggct tgcacaaagc agagctgagg tttgataaat cagttaggcc agaaacgagt 600
tctttcgtta atgagacaaa aatatttggt cgtaaaaaag aattgaaatc tttgatggag 660
aagctgggag ttaaaattcc agttaaaagt acagggaaga gaaaattggc cgaaggtagt 720
gcaagaatgg aagaattacc tgtgttgcca ataggtaatg caagaatgac agaattgtct 780
gtgttgccaa tagtcggcat gggaggtatg ggaaagacaa caatggctca acaaatctgc 840
agtgattcaa aggtgaagaa gcactttgat ctcataattt ggacatgtgc ttcagatgat 900
tttgacaaaa aaagattaac caaagagatt ctagaacacc tgggacaaga tacatcatcc 960
aataacctga attttcttat gacaaaacta caagattgtg tcaggtccaa aaagttctta 1020
cttgttgttg atgacatttg ggatggtatc ttgaaggatg atggggcggg atggagggag 1080
ttttgtgcac ctctgcaaaa tggttctgaa ggaagtagga ttttggtcac tactagattt 1140
gccgaggttg ctaaagtaat tagccccacg tatcactatg agttaccttg cttaaaaggt 1200
gacgtattct ggaagttctt caaattatgt gcatttggat ccacgagttc tgtcgacaac 1260
cgtgagtcac tagagtgcat tgggaagaat atacttccaa agttgaaggg caacccattg 1320
gctgccaaga ctattggacg tttgttgagg atggacctaa gtacaacaca ttgggaaaat 1380
ataatggaaa gtgaactctg gcaagtagaa caaacggaga atgacatttt gccggccctt 1440
cgactgagct atatgtacct tcctcagaac ctgaaaagat gcttctcaat ctgtgcaatg 1500
tatcccaagg atcacaaatt tgagatgcgt tttttagctg atatttggac tgcacatgga 1560
tatgcggaga atccccaaaa agcatcactc tactttaatg cccttgcaaa tcggtcattc 1620
tttcagaaag catctacata cagttctatg tatgtaattc atgatctgat gcatgatacg 1680
gcaaagcttg tctcaaagga tgagtgcttc atcataaagc atccgcgtga catagataga 1740
gtcccttcaa ctgtccgtca tctgtcaata tttggaaacg gagatattga ttgttctgaa 1800
ctgaagaaaa tatgtaacag caaaaaaata aggaccctgg tatgtgatga taaatttaac 1860
agcggagcag attttgctgg cgtgattgac tcttggttca aggaactgtt gaaaatccgt 1920
gtactgaagt ttagagttgg caaagtaagg aagttacccg agagcatggg caactcaaag 1980
catctgcgtt acttttgttt actggggcct tccaacttcg gcaccctacc ttcaacggtt 2040
tgtcgtctgc ataagctcag aattatagag cgtgccggtg gtgtcattaa aaggtttccg 2100
acaggcttca gtgacctcat cagcttgcag agaattaatt caaaggggtt cacttatgac 2160
agggaccaat ctgacaaact ctgcctagaa tggtcctttg aggttggagc acctagtagt 2220
gagacggtga gaatgatgga aaaccagatg gaagaattac ctcactggaa tatccaacat 2280
ttgcatgtag agaattacga aggtgaatct tgtcctagtt ggctcaagcc caacctcctg 2340
ccagggttac gctcgcttaa atttcgtagg tgtggcaaaa tcaagagcat accatttttt 2400
gtcccgtcag aggagtcacg cgataatgca caaggaccag acaaccttaa tcgtcttgaa 2460
gagttagtcg tggatgcttg ttatgaaatc aattggcaag gctcgatagt cttacctact 2520
tctcttagga agctaacact ttcggcctca ggctgttcca tggataacct tgtgagctgc 2580
ttccgtgacc tcacctccct cactcatatg gaaataaggt tcggcagatc attgacatcc 2640
atccccctcg atgtgtggag tagcaacctt ccatccctgg aggatctatc aattatggat 2700
tgctaccgcc ttgcatcaat aagcgcctct ggagcaagca gcagcagcgg tgttaaagga 2760
ttctcagccc ttgctaagat agaattttac gaatgtggcg cattactgag ccttgatgaa 2820
ttgctgacgc cagactatct acctgccgtg aagacgattt ctgttacaca ctgtgaagcg 2880
ttggcatccg tgtctgtcga caggctccac ggcctggaga acctggaaat tggacgttgc 2940
cacaactgta gaatattgtc agtgccgtcc tctcttaagc agctcgactt gtatagttct 3000
gacggcatag aattcatcga cctcagcaat agccagtctc ggagttcttc agcactgaag 3060
gtacttagca ttacgggctg tccaagcctc aggtccatcg gtggtgaagc agctattgct 3120
gaaataatga atgttggtat aaccgggtgt cccaacctca cggagataca acagccactc 3180
gagcgcggga cggggaggtg gtggccacgg gctgattaa 3219
<210> 9
<211> 27599
<212> DNA
<213> Lr13基因序列
<400> 9
atcattttta tattctttgt tagacggttc atttagttga ctatgttcat tttaactatg 60
cagatgtcat atatcgttca atatgcttat tatgcatttg atgctatatt tttaagataa 120
taaaaacgtt ctttgtaaaa aagaaccgta agcacgaaag aaaagtttag caaaatcttc 180
gtaagggcat ctccaacgcc aatcctcaaa ccgcacacat acgtctgcac tgcattgttt 240
ggaccgtgga agtcatccaa cacggtcctg tatcaatctg tagcatggtc cagacatact 300
ttcttccata aaccgaagac aaacgtgggg gctttgcgag cttcctgaca ccacccacgc 360
ccatttctga ccacactagc ccaccaaaac ccttcctccc tcgactgcgc gcgttcccgc 420
ccggcgccat cccccattca tgccacttta gaacacaccg gtgcacattg aagacggctc 480
aggacagatg cgacctctca cgcctttgcc attgaagcag gactccgccc attgcacgcc 540
cggctgaaca cgcctcctcg ccgcattcaa atgcctccat taaacccgcg tggaaagccg 600
agaaacctac tccggccaca cgtccgttca cgagcgggcc gacattaaac acaacgccag 660
gcaagctctt cccgtccacc cgctatttaa atgaggccag acgccggcca agaatcgcac 720
ctctccccat ctcctccttc taccattttc tccacccttt cgatggcatt cgacgagcaa 780
gaagagcagt tctccggcat aaaagacggc tgggtggccc gccgagccca ctagatacgg 840
gcgacgcagc tcactgctcc acctccctcg cgtagctcga tggagcaagt tgcccaactc 900
gcgttcccag tcggacacgc gacagcaggg gggggggatc atggcggcat gggtatcgtc 960
gctgtcgtgg atgccggtgt gtagcaccgc gcatcccatg cctacgactg tgccatccgc 1020
cattaatgcg cgtgtagccc cgcgccgccg gcagagaaaa aagccgtctg cgtagacatc 1080
gacgaggtag cgcatcgcgc ccgtgcacgc cgccatcatc gatcagaagt agaaaacggg 1140
aggtcaccgt cgcttgggtc ccatgaaggc caccgctgag gcgacgccgg cgtacatggc 1200
cagtgtcttg ccctctcgcc agccatcctc gtccccttac cctaaaacca actttggtcc 1260
atgcagcgta gggactcttc ccctccggtt gaagcatcag ccggcgggtc cactccgatg 1320
agttgtggga agcgagccgc cctttgtgcc gaagcatgta cctctggggc tcacggctgc 1380
cggacatgga gacgacaatg gagatctgcc gcagccgcag tagatagtcg agggtatcca 1440
tgttgtcgaa atggatagcc gttgcgtcga ctgtagacta gttttatctt agtccggtat 1500
ccttttcagt taaaaatatg gccaaaattt aacgttttca tccggtttat atgaaatctg 1560
tcatatttat ataaaatccg gccgtgttta catgaactcc atccggtttg atgaaaaagt 1620
gtttgcaatg tatgcaagca gtgttggatg atggcttctc gcatcctgtc tgcggactgg 1680
tctctcctgt ccgcagatag atgcgagaga aaatttgtga ttcaccgttg aaaatgccct 1740
aaccaaggtt ggtaggccat agtgaagtga cgcgttgatt gacctgagat gagagacaat 1800
gaagcagaga acgcttcagc ggtgggattt ctcttttttt ttttgagctg caagcggtgg 1860
gaattggttg ggaagaactt cttttttcca cggggaggtt gggaagaaac tattagagca 1920
tctaaataag gtaagagtaa ttagcaattt agagcatgta caattgtgct atcttaaaag 1980
tgtcacgtag aataaataat gaggtggagg agagagaact cataagaaaa ggtttgtctt 2040
ctcttactta agaaaagaca agagatgatc tcttagcaca gtatgtctca tcatgttttt 2100
aagaatatct agttattgaa gataaggcta aaaaatgacc cattatagac aattttattt 2160
gtcatctcta aattacatgc aagacttaag acaagaccgt cttatcaacc attgtacatg 2220
cccttaagcg accaatgagc tgaatctttg tacgaatcaa acgaaggttc ttgacaagtt 2280
cgaaatccct cgaattttgc taagttaaag gagacaataa aaatagaaag ttgtctgaga 2340
aggaaatgag gttgactgac cgaccggagg tgatacaagg aagtgaaatt gggaaatttg 2400
gccgagaacg aactaatcaa cgtgagaatc tagacggtaa caaggtgcaa ctttcgtgtg 2460
atatactccg aaggttcttg gcaagtcttc ggtgaagccc ctccactcct tagtcctcac 2520
accaaaagat aagaagatgt atcagcacct tatgtaacgc cctctggtgc tatccgcggg 2580
agacgcgcga aggcttctcc ttgcgccact caaactgtgc ggtttctctt ctctctgcga 2640
tcaagccatc agcaatttcg tttcggcaac ggtgattgca tcaattggga caggacctcc 2700
ggagtaagcc gggcccttgg caacgaacgg ccggcggccc tcgtgttctt ccaacgcgtg 2760
gggtaaatct gaagaagcag accttttgca gtttatatat ggggtcgaat caatagccca 2820
caccaaggtg catgatccat ctcttctctc tttcgggttt ctgttgatgc tgctagcatg 2880
tagaggtttt gttttccttt cgactaaatt tttacggtaa ttacaactag cgctgggaga 2940
gggctggcaa acggaccctt ttttatggca gttttagttg tgagacgaga tcaaacaatg 3000
acagtttaaa acaaaaaatt cttctagaat agatactttc atgtcgctct gaagaagact 3060
gtcatccctc aaactgaaac tgcccgcaaa atcattcgca aatgccacct cctcctcgcg 3120
aacattcgat agctaagggg gtccattttt ttttaacgaa cgtgcctatg ggaagccaac 3180
cacgtacgta tgcgggtgtt ggaatcggta caacttgtac ccttggcaaa acgtgtcgaa 3240
ctctccacaa gcacaggaac caccaggcag caaagagccc cttctcaggc gacggtaaac 3300
acatggtctc aatcgccaaa gatctttgat ttggtggtgc tcagcccaag atccgtgccg 3360
gagcacgagc atgggcaagt gtgacctctc cgcccacacg aacaaaaact aaaccaaccc 3420
ccgagcagtg aggcaggtcc tagctcgagt actacaccag aacaaggaga aagggggaac 3480
acaagcctcg cccctcgcga aggaaaacta caccaacaat atatactccg caactacatt 3540
gtcagtaagg gcatctccaa cgccgttccg caaagtgcct gcaaccgtcc acaccgcacg 3600
gaccacacgt gttgtgtcat ccaacgcggg ccagtattgg cccgcaagat ggtctagacg 3660
tactttctcc tgcaaaccgg agacaaatat tggggaggca acctctcacg cctccgccat 3720
tgaagcagcg tgccgactga gggtgccgcc cacgacgcgt ctcgggtgtt cacgcctatt 3780
caatactgga gacgtgttat tgtatgggac ggatatctac cacgccattg aatgtctcgc 3840
acgtccgtat ggcaccatta agaaggctcg tcggccggga tacctaccct ggcaccacgt 3900
attgacacac ccggacgcct agtctcaccg aaatccgcta tttaaaggtg gccacacccg 3960
gctaactcca ccccacgaca caagccgctc cacatccttc tcccttgcac tgcatctgtc 4020
atgaatgcag gtggcgaatc tctgtgggag agccttttaa cggatatgaa gcacgaggtg 4080
gccacccttg ccaccgcctg gaagagctgc catgccagga gaatcgaggt cggcctgcca 4140
gccagctctc ccaaggtctt ggacgacgag gacgacctca tgatggagac gagctccaac 4200
gataccgccc cggctctcgc gcatgtgcat gctggcttca ccatggagca tacgcaggct 4260
cactacaatg ccgccatggc ggaggtgcat catgcaccgg cacctttgtt gacgttccag 4320
cagacggagg aggaatagcg gtacaaccaa ttcctcctga aagcagcacc ggctgcagga 4380
gggccagatc tacggcgagt gcgtgagcga gaggtcgtgg cggccatggc cgcagcgaac 4440
cctaacttca tcacggagca gcttgacatc aatggtgtcg tccatgctca agccactggg 4500
cgaacctaca acccactccc tatcctcatc ttgtatggga tgggtagaaa cttctgctaa 4560
gagcatcctg agggcacatc atagagaact gaagggggtg gggggggggt ccacaacccc 4620
cctccccccc cactgtgtct ttcagaccaa agaaaccaca accaaatatt tgcatatata 4680
ctaaaaatag gcctagattg atccatgaaa ttgatcgtaa ggtgcccctc atgtctgtag 4740
attaagcacc gtggatagag tgctagcaac atggcaatgg ttggtatggt ggcaggaaaa 4800
atgggtcaca ccacaaccat cagctccttc tgctacagct ctattacaca acggaggagg 4860
gaggatccac gaaccctgat tcccatccca tagtttgaat tacggacgtt tgaaggatta 4920
ccataagcat tgttgctatc atagttattc tagctgccag atctggagtt ggagtggaag 4980
ctggggttat tgtagttttt gtacccacat gatcttttga agttgtagtt gtagccagat 5040
ccatagttac ttgaggtgag gttgttggag aagctgctag atctagtatt ttgaagttgc 5100
ttgggctaga tccgtggtcg atggctcggg tggatggtgt gggccccatc aggccgcctt 5160
tgggaggaag agcttggagg gtgctgccac cttacgatga tcatattcct tctaccaggg 5220
cgacgatgag ccccacctcc cccgctcagg atccacggga atgtgagcct cctactcgtg 5280
tcgtatcttt tgtcgtagat ccacatcata gtgcaatgat tcatccccaa cattgatctg 5340
cccctcaaag atgcttgccg aagctgatgc agttagtcaa tggcttacaa aacattacaa 5400
ccatagccgt gaaagagcgg gagaggaaag gaggggcagc catctgagag agagagagag 5460
agagagagag agagagagag agagagagag agagagagag agagagagag agagagagag 5520
agagagagag agaagcaact aaaatgacgg gcatcagcga cggaagtagg aaaccctaat 5580
gagggattag gagatccaca aactagccat atgtacccct gcggatgggg cgagatgtcc 5640
tggggacaaa aggaacaaat cggtctgaga tagggggatg caccaggtcc acatatatca 5700
gtcaccatgg gaaagtcgag cccactacta gccaaaattg agcataggcc acccatgata 5760
gggccagatc caacaggtct agcaggccca ttgcccatca gggcggcatc tacgagggaa 5820
aatactaggg ttccccttga ttaatgattc accgctgcac caagagaacc aagtgatgat 5880
gccgaggaga aaaactttga cgcacaatca agcatcaaaa actcatcagg agacgacggg 5940
gaagccatac ttggctccag aatgcctgca gtgccctcta gcagcataca aaaaatatcg 6000
cgcctcacta ggatttggcc acaaccaagt cttgggcaag gtcggcctcc tcctgacgcc 6060
agagaggaga taaccatgac cccatgtatg agagcatgac acttacctaa gccaccatct 6120
aggctagtct ccccatgcac cgcacctcta gagccaacat gatgttgaga actgacgacc 6180
tgagcaccta gaagcagctc ccctgagagg agtttcacct cttactagca atacgacacc 6240
tccctcaagt cccagaacaa attcctaggt gcaatagagc ggtatacact aggagagggc 6300
aaagaaggaa gtagcatgcg gtagagggag gggaaagggg gtggcgtgtg agtcacgaga 6360
ggggtcgaca agggagggga gaggcccggg agtatgagag agttattcgc accattgtaa 6420
gcattgatga aaccatggaa tacacaaatg aaaggatgag agaacgcatg cacatgtaca 6480
actactattg attgcttctt catattttca tccaggcaat cttgtctcct tgctcggctg 6540
cgccatacaa gtgggccttt acatgtgtgt gcttatcaat gcatgcccca ttatactatg 6600
tcattgcatc atatagatcc ctcctggggg gcggagagcc ttgatgcaga tatcaacaaa 6660
gttatgtttg acagatttaa cgcatggccc catggtgggt gtcaaatgtt gatgactgaa 6720
tcaaaatatt gtgcatgcgg gtgtatgtat aacttgaata ttgagtcaag caatgcaagg 6780
accccattca agacccttgt tatgatgaag aaaacaataa catggttgtt ttacctaggt 6840
tcagaccctc caatgaggaa aagtcatgcg ccttgctatt gtatattaaa ccgagattct 6900
agtttacaaa ctgtaggttg aatctctccc tcccccaagg gttcttgggg ttttatgtgg 6960
gcactccttg accaacaaac aagtaaatct atctttggat gatcccccat catctactct 7020
tgtaaatgaa aatatctctc ataagaacat aattccaaaa cagcatgcct agattcatct 7080
cctctcagcg tctattttag catgtatgga tgcgaaccat atggtcccta gacacctttc 7140
agtcgactca ctctatccat atccctaggt tatagccctt tgatatttta ttcctaatca 7200
ctgacaatgc catattctat gatattcaca actgctacta ccgtcgtcga aggagaaaga 7260
ttctcttttt acacttccga aagcatcatg aaaacgatct ttgtcattac catgtccata 7320
gtcacaaagc atcgtcaggc gattggtact cataagcact agacactgag ccaaggtgtg 7380
agtcgactta tcatagatat actaatagag cgtcgtggta tagtcagaca acatgcataa 7440
tcagtgtgac atttacaaca ccataatttg gtttgttcct ttagcgaggg cctatccata 7500
tccatcgact ggatatacct ccatatgaat gatggggatg acctacacga tagtcgtaac 7560
tacatgtacc aattgatgaa ccactaggga gcatttaaat agaaaaacac gcaagaccct 7620
tgatgccctt tgtgatccgg tttggatagc agaagattcc cgtacctttt ccccaaggat 7680
tgaccctagg ttaccgaacc cacgagagga tgtgctctaa gataaagagt caaacaagtt 7740
attgttattg aattatattg tagttattta accagacctt taacaatctt tttgggtgta 7800
aacattaata tgaaaatagg catgggtatg aggtcttact ctcaaacctt gtatttgtgc 7860
ttgggatctc atcttgataa tagacatttc tggaagacag ccgttatgat gtgcttgctg 7920
gagtgggaga cgtgtccaat gtccatcttg accgatacgt gacctgcatc aagagtcatt 7980
tgtccaactg atggccctta gcatcgattc acacgaggtt acctcggtta ttaagaataa 8040
acattgagat aaaagcattc ggcgcttaat gactgcacct catttatccc caaggaattg 8100
atttcctcta ccatactaac cccttctttc acttttgttt ggggtagtgt tccaggcatc 8160
ctcactcgat ctatctacgg gatcctgggt acctgcaggg ccttagaata agaagaagac 8220
aagggacacg tccgctgcaa taggacacat acaaataacg taaaatgcgt aaagtcattc 8280
caatgacccc atgcacaaat acatagggtt gcaagacctt tcctcaaccc gttgccatac 8340
cgaagtactc acacatggca aatggatgcc aatatgaact cattgaagat gccataaaga 8400
tgaatgattg attgataata tttcttacaa tagttgttga atgtgataca ccgattacaa 8460
gtatgactaa cttggagaga atggttgtgg tgatgaaggt ggtagcaact atggaggtga 8520
gatgggaaat aggagcagct aaggctggaa gatgggttct cgttctctgt ttcacaaatg 8580
gtgctctccc tattgagaat ttggcatcat cccctttaga gaagttggtt aggtggatgc 8640
cctattgcta atggtgtgta gtacgactgg tggtataaga gggcgcggtt ataccatgca 8700
tcctcttgtg ctctagacta ctccctgtgc ctcctcttga cttgcctctt caaccaaact 8760
tcgtacttcc tcctatgtga tggtattctt ccgtgtttag ggcaatttct tttgaaaaca 8820
cattaaacaa aggatttcgt gatctcttgc attcattggt cattagtggc ggcaatatca 8880
gagtgatttt ctttaaaatt atgggattgt ctagtttaaa caatggtgtc atgagcgcaa 8940
aaatatcacc catcaaccat tcacatggcc ccctactagt cgcttgccat gcttggcagc 9000
ggtaccaaag gactataccc cattgatgga gatggcgtcg gtccagccaa agggtggaag 9060
ctacccgttg gctattacac tagcgcaggc ataggtgtaa tggttagtgg cactaccaaa 9120
actctggagg gtgctctcta tggtggccta gcaatgcatc caaaggcagc aacacaaaac 9180
acactaccaa caacagggga ggccatgctg ccaacaacag gggtttcagg agcaccgacg 9240
cacatggttg aggtttgcaa agcacatgca actaattcat gagccggact taaaccgaca 9300
caagaaacat gagcgggtgt tctttcatct atgttggcag gcaacatgga tcagaggatt 9360
cccgcgaacg ttcaggagca ccatggagga gatgactcta tttggatcta ttgagggttt 9420
ccttgtgacc acatagtcca caagagacgc ttgaaatcca tgtgttaagg tgaatagttt 9480
gaatgaatca tgtcccacat ggcctagcta atgcgtagct ttttccttga ccttgggttt 9540
agggcaatga aagtgtagat gatgtcgatg ccgatgggct agagactttc gcatgcgctt 9600
tcatgcacat ctcctaaaga aactcctcga gcatgggtag tgtgtaatat gggactgagg 9660
aatgggtgcc caacgctcga tgcatgtacc ttggtgcgac ccttttcctc cattggtgac 9720
taagatgagg aggaggaagg catggcattt caaattaaag gcatcactgc actggcctcc 9780
catctctata gagagtgaac aagaaatttc acttttgtgg aaagtgacct ggcacagtca 9840
tggagtgctc ttggtgatgc gatcctggtg ttgggaacca tatatccaac aagaaatcat 9900
gccatgaact tcttcgacat ttcgtcttct ctagtgactc ctagatgcac ctggaatcta 9960
cgataggaaa agaacaatcc aaatgaatgt gatgcagagg aaccacttgg tcagcaacat 10020
gcaccagggg gctagcgttg gcgccacctt ttgcaaatgt cccttctctt ttcaaaaaat 10080
atgtcagcta ctatggcatg taatacatgt taactaccat aatgcatatg ttactccaaa 10140
ggcatctcag gtgaacccca agccactatc gcggccctcg tctcctttct cctctatctc 10200
cctgtcattg tgggatgttc ataggtctgc gtatcaagct agccatgaac ggtaactgcg 10260
tacgatgggt ctatgacccc aaggtggcat catgtggcat catcgtttcc aaggttgtca 10320
tggttgcgag ggtagtatgt ggttgtccac ggctatgggt ggcaatgaat tttgacatgc 10380
ttgcctcgat gtggctgatg gtggaaattt ctatgccaag ggtgtcaggc atagttgtag 10440
atggtgatgg tggcagtggg cgccacccta cagctggacc agtccaacaa tgcagtggca 10500
ctcatgctag atccggagct cggggccctg tgttggctat gatacatcgt gtgctcaaca 10560
catgtgttgt ctcatgccaa ttggtttggt gtgcaggcaa gccattggca tgggcaagga 10620
ttattcgggc tttgatggtt gctgccagat ctatggcatg agtatggacg ctcctcgacg 10680
accacatggg tgaccttcaa tgggaggacg tctcacccat ccattagcgc cattacctcc 10740
tctcatcgct ctcatcggct acgggacttt cttccttggc ggggtaggcg gttggttagg 10800
tggcttgcac tactttggcg catggttgca tggcgttcca ggagagtcat ggccctcctt 10860
cgtcggatgc caatgtggtt caacatggat tactggttgg cacaactatc agaataattg 10920
tatgatatgg tgcttagaac tccacttcat gtctaggttg aaaactcttg ttcaggtctt 10980
cgttggttgg tttcagcaac aatgaacttg agtcgctatc ctattgaagg cggcatctcc 11040
gtgccaatgt tctggctttc attgactaaa cttggcgccc aggatgaaaa tctttgtcat 11100
ggtcgtctct ggtaggacat ggtgatgacg catgagtgtg atgtcccttc ttgaaggtga 11160
ttgttgcata agaaattggc ttagtcatag gtgttttttc atatcttgta atattttttc 11220
catagtagtc tacaccttgt actcttcctg ttgatgatta tgcctagttt ctattgtttt 11280
gcaaagtgca catatcgcgc actgagccac caacacacga ttctccttgg atgtgtttgg 11340
ttacctgcat cgttttttgc cactttgcat acttgtccca gttgggccta gctgagcata 11400
tgcaggcaaa aaacctatat ctgcatgttg attggttgcc tgcatcgcat caaggcccag 11460
acgggtgcaa gttgtttggt tgcctgtgtt tgtgctttag ggatgagaag atccaaagca 11520
cagctgtttg gttgtatgca agtatttggt ctgctcaccc ctttcaaatg tggtggactt 11580
atcacccatt tgacagcaca caacacacta tgatcaacaa cagcagaaaa acagcaacaa 11640
caaagcaaaa acaaagcaaa gaacaggcag atcataagct actaggcata cttttaaaac 11700
agcagggcat cccatgcccc tgctggaccc catgttgctg ctccccgggc aaaatatgga 11760
catgttcaca gcaaaacata tgctcaaact taaaagttga tacctaagct atttcagttc 11820
agacttgaca aacgaaccac atggcttgag tgatgttgcc gacgcagctg tgcttcgagc 11880
attggaccct atagatgttc gaggtgttgg catcgaagat ctgcaccttc agccccagcc 11940
cggagagtct gaagaatagg aggtcgttgg ggacgatctt ctgccgatgg aagaagtaca 12000
cccagccgtg cctgagaacc atatgcccct tgaacatctg cacctagacc caaaactcac 12060
atcacttatt ggcggagagg ctgacgacgc gagaagggcc aaggaccaac cattgcttca 12120
tcaccggcgt gaaggcaggc gggatgacga gcatggtgat gtcctcctag tcatggatct 12180
acacgcagaa cctgacaggt tcctgtgcac cctcctcgtg gccggtcctt ggcctcctgc 12240
cagcagtgag cggcatgctc atgtactcaa ccctgtgagg aagcaccacc ctcttcagcc 12300
actggtttgt ggggattacg tcctccatac cctctgtgcc ggcgacgacc tgcgctccgc 12360
caccggctat gtcccacttc ttcttgatta tcttaccagc aaggtggatg acggttagca 12420
acaatgaaag atgatcattg atagtgcaaa tgccaatgat cagtgctacc aatgatcaat 12480
gctaacattt atctctactc ctaatggagt agttggtgaa tagtctccat ggtttatttt 12540
tgctgttttt ttgtctctcc tcccaccacc ctctcccatc ctggtccatt ggtttatttt 12600
tctccccact ctttattaca accaggactt tcctaacgta taacaaatca cggatctaac 12660
ttccttaaat tattcatatc aatcgattcc ttttattggt tcaatttgtc ttacgaaaca 12720
aataacaaaa atttaagaaa caaataatag atttgaatct aactttccta acttatctaa 12780
atcaatcgat ttctctcatt agtgaatttt ttttaggaaa caaataacag acacgtcaca 12840
aactttcctc ccaataaata gtttcttaac atttgcaaac tttcctaatc agacacgtca 12900
caaacgggca ggtactgata tcatgttacc aaacaataaa accccatttg catagaaatc 12960
agttcaaaat cctaactttc ctaaattttt acctttcctt gataacatcc aatatttcct 13020
atgttttcca cctattgaag gaaatcaccc ctcaatcgat attagcagtt ggtgaatagt 13080
ctccacggtt tattttcgct gggttttttc gtctctcctc ccaccacccc ctcccatcct 13140
ggtccatcgg tttatttttc tctccactct ttattacaac caggactttc ctaacgtata 13200
acaaatcacg gatctaactt ccttaaatta ttcatatcaa tcgattcctt ttattggttc 13260
aatttgtctt aggaaacaaa taacaatttt ttaagaaaca aataatagat ttgaatctaa 13320
ctttcctaac ttatctaaat caatcgattt ctcttattag tgaaatttgt tttaggaaac 13380
aaataacaga cacgtcacaa actttcctcc caataaatag tttcttaaca tttgcaaact 13440
ttcctaatca gacacgtcac aaacgggcag gtactgatat aatgttacca aacaataaaa 13500
ccccatttgc ataaaaatca gttcaaaatc ctaactttcc taaattttta gctttccttg 13560
ataacatcca atatttccta tgttttccac ctattgaagg aaatcagcct tccatcaata 13620
ttagcatttt tttggaatga gatctccggg ttatgatttt tttgcgattc tttccaactg 13680
tgacctctcc ttctgctccg gctccttctc gcataatcac ctccaccaca gccagctggc 13740
gccaccctaa acctcttgcc tctccacacc ttctccgcca cctcctctcg tactccattg 13800
acaatccctt cgccacgttt gtccacagcg gtgtcgaggg cagggcagaa cagcatacca 13860
gtggtagagc agcacatagc agcaggaaga agcacgcagc aggcgaggcg agcggccgac 13920
gatgaagggc agagatgcgg ccggcgtggc tgagggggca gccggcaaaa gatggccacg 13980
acaggaggcg gcgcgagaca gggcgcgaca gcggggtgag cgaagggata ggcggcagca 14040
gacggggtga gcgcagccag cgagagcgtg acgcacggcc agggtgtgcg tcacgcgggg 14100
catcgtggtt cggtgcatgt ggcgagcgtg atggcaacgg cgggcacgac ggatgtggtg 14160
tggcacgtgc gtgggcatgg agagccagag agggagtggc cccgcgggcg cggaagaaga 14220
gctgaaggct ctggcatggg cgcacatagg agcgggcatg tgccacgggg tcaatctgag 14280
gggctcaatg gctacccctg caattgccat ggcggacggc ggttctcggc cacggcaaaa 14340
tacttaacga gagcaaacaa gaggggaaac tggggaaagg aaagaggaga tcacggcggt 14400
gtcaatggca ctctaggaga agacagggga gcttggggcg gcgcggatcg aacgggaatg 14460
tcgcggtggc cgaaggttga ggaagacggc aacgacatcg atctgggggt gctggacttg 14520
atctcgttgg cgcagatgaa gtagcggagg cattcggagc tcctcgacaa gctcctagcc 14580
agcagagagg gcagtggccg tgtggacggg gtcggtcatg gtggccgtgg catctgactt 14640
cgtccagatc accgggatcg agcgagtgag gaggagaagt ggatctggga gggggcgtcg 14700
aggaggagtg aggccatggg ggcacgggag gtagggcatc acccttatcc attccccttc 14760
gatgccgacg aggtggtcgg gcgggagccc ggctctgtag cgacgggctc ggggaacagg 14820
agaagacgac catgcgggga ctggaccact gggccggttc ggtggcaagg cccgaggggt 14880
agggggaatc ccctttttca tcgtcctttt ggtttttaat gtattctaat ccgtttctcc 14940
ttttaacacc gttttctatt tattcttata caactaaatg aattttgtta attgtgaaac 15000
tatgcacaca attcgagcac aatattttta ggtggcacaa aaagtttgag gccaaaagaa 15060
atacataaaa ttagatttat atttaactgg ttattttaac tactgttgga cctcttatta 15120
atttgacaag atttaatttt tggttcaaat aatgttgggc tcaacatgtc aaagatcagt 15180
ggaatttata gaatatggtg aacattttag ttttactatt tgatgaaata atttatttga 15240
ctttatttta aatttgaatt gggctttgat ttttatcaag tggcagtttg gcttagcaag 15300
cctgaggaca tggcatcatt agggggaggt cactgtagct aatgacttga ggtgttacat 15360
aatcttccca tttctgaaat ttcattgtta aatgattggg gaatatatca cactatccat 15420
tagttcagtc aagctcatca taagtccaca cattattgcc aattgagtaa gtcttattgg 15480
cggggagaat aggctatgtg tgtgtgtggg ttagagagag agagagagag ggagagagag 15540
agtgaggaag agggaggggg agagagtgtg cgtgtgagga tttgatggta aaaaaggccg 15600
ctaatgtcgt aatatcgaac tcttaaagtt aatgtggccc tgttgcaacg cacgggcgtt 15660
cttctagtca agctaacaat gatcactgat aacactgatt ggtgctaaca gtgatcattg 15720
ctagcaatga tcaatgctaa cactgatcaa agctaacact agtcaatgat caatgctaac 15780
actaatcaga gcaagcaatg gtcagtgcta ccatattatt ccaaatgcac atgcattttc 15840
ttactctgat catgtgtctc ctactactat tttgggcatt cgggtatgtc ctactaccat 15900
gcacatgcaa tgtcctatgt tgataatgca ttcacatagc ctaacaacaa caagaacaac 15960
atccaacaca cttgcacatc atacacacta tgttgatcat cctatgaaga acaccactaa 16020
catacacgac aagagcaaca atagaactac acactaagaa caatgctaag cacagcaaga 16080
acaacaacat caacaatgtg gagcaaacaa aacaaacaat gtggcaagaa caacatcaag 16140
atcaatggta agaacagcaa gaacaatgct aacaacatca ctaacaccag caagaacact 16200
gctaagaata gctctaacac tataaagaac gatgctaaga acaacatcaa tagtggcatt 16260
tggagtgaac aaactagtat tccctgtcat ctatctaatc tagcactagc attcatcaca 16320
aacaggaaga acgacatcaa catgataggc atgcacacat attcatccct aaaacggtgg 16380
tttcgtatcg aatccgctcg aatcgaatcg aatcggattg aatccactat gagggtgccg 16440
gtggagctta tgtcggtcgt gagtcacaat ctaatctaac ctaagaaccc taactatgaa 16500
ggaaggggag attagaggag cttacggtgg gggtgccggt ggagcgtatg ccaaccgtgg 16560
agaaaacccc ggcgggaaga ggggtgccgc cgccgaagaa gaagaagaac tggccgtgac 16620
ctcgttgtgc gccggctgga agaggagccc ggcgtcctcg agctcccctc cccttgcacc 16680
ggacgcaggg gagaaaaccc tcgaatggtg taaaaatacc cccataagag gggtcgtcaa 16740
atggcggagt cagcgggtat gaatgcacgc cgttgttgag cccaagaagt ggagctccag 16800
caccggtggt ggaaccggag cgagcggcga cggcgctgca ttggctcgtg gcggcggcat 16860
ggtgcagaac ggagaggaca caaggtttgc caccgcgaga tcccgttcgg aggcggagag 16920
accggcaacc cacctcctgg atgctattgc tgtcggcgat gaccggaatg ggtggggcgc 16980
ggcgagacgg tggctctgcc atcggagaaa ccaaacgggc ggtgaggggt agggagagtg 17040
cgtgcgaact atcgagtgtg gtgacgggaa ggagcaaggc aagttataag atgtttcgtt 17100
gtctcccgtg cttccccacg cgtggctcga gcgaacctgt cttgcaaaaa aggccgatcc 17160
gaccgttctt ccagagccag acccagacct gctttgaggt tcgtgcgggc caagttttag 17220
atgcagctca ggcaaccaaa tgggcccaaa actgctcgcc tgggcctggc tgaacctaat 17280
gcatcccttg tgcttgcggt gtgccttgca tcaaatttta taggatatgg ttgtgcgcgg 17340
ccgaggaaac agatgtcaga aggtttatcc aaagtcatat gctattataa atatatatgc 17400
cacatcagat cactggagac atggagcaaa gaggggaatg aaagagggag tttactgttg 17460
caagaagtgg tgagctagct cgcctagtag caaataaatt tattattaga ggataactta 17520
ctagaaaaat attgaatacc tcagaccata taattagcag ttttatgcaa gatgcgcaca 17580
caaactattg gagggggttg gtactctgta taatatgcac actcattttc atcctaataa 17640
taatggccac agtgaggggg aaaagtgcac ccgagtgcag atcagcaccc aagctccaat 17700
atatgcacct gttatgttat atattatttt tgaaaaatgc atttgtaatt atcaaaaagg 17760
gctgaaatat ataattaaat gtcatttgtt tagacaaaaa aaagatgaag aggaggtcta 17820
aaaaagcaac ttcgaaatcc atatctcgcg catctactag ctggtttaca tgcgttatgt 17880
gtacttatta attcagacat cataaggttg ccctcgatga ccaggtaaaa gctgaaggcg 17940
taagagataa acagggttag tactctgttg ttgcctctgt ttccaccgaa gtgataccaa 18000
taaccaccgc gaacaagaag cattcgtcca cccacaagga aagaaggtaa agcgatatct 18060
tcatgttccg actctgccca ttctaaatgg ttgcctattt tcgataaaat acccttggga 18120
ctaattttgg tttatatgat cacaggagtg agttatttat ggccatgatc ttaccagccc 18180
ttcctgtagc tgttgaggta atttccagtg tcaatgcatt gaacgaattc gtcggctggg 18240
cgacaccaca catgaaagcg gccatacgtt atgcacggaa cattccacgg tcaccagatg 18300
aagagaacac agctgcacaa cgccagattc aagagaaaat tgagcagcta ggcgataagc 18360
ttgggaggct gaagactcgt atttggaagc tgaagacaac tatgcccaag atgcttgacc 18420
tcattgacag ggttgagtgg caaagccaca aggaacaagc cactgttcta ctcccggaga 18480
tcaaggatgc agtgtacgat gcagaggatc tgcttgatga gtttgattat tatgcgctca 18540
agttgaaggt tgagtgcaac aaaaatttgg ggcaagatca cttgcataat gccttcatgg 18600
aattcttaaa cactatagac tgtgatgagc ccaataatta caccacgaaa gtcaatgata 18660
ttcaagaaag actagatcat ctgaatcaac agtccaaaga tatgggcttg cacaaagcag 18720
agctgaggtt tgataaatca gttaggccag aaacgagttc tttcgttaat gagacaaaaa 18780
tatttggtcg taaaaaagaa ttgaaatctt tgatggagaa gctgggagtt aaaattccag 18840
ttaaaagtac agggaagaga aaattggccg aaggtagtgc aagaatggaa gaattacctg 18900
tgttgccaat aggtaatgca agaatgacag aattgtctgt gttgccaata gtcggcatgg 18960
gaggtatggg aaagacaaca atggctcaac aaatctgcag tgattcaaag gtgaagaagc 19020
actttgatct cataatttgg acatgtgctt cagatgattt tgacaaaaaa agattaacca 19080
aagagattct agaacacctg ggacaagata catcatccaa taacctgaat tttcttatga 19140
caaaactaca agattgtgtc aggtccaaaa agttcttact tgttgttgat gacatttggg 19200
atggtatctt gaaggatgat ggggcgggat ggagggagtt ttgtgcacct ctgcaaaatg 19260
gttctgaagg aagtaggatt ttggtcacta ctagatttgc cgaggttgct aaagtaatta 19320
gccccacgta tcactatgag ttaccttgct taaaaggtga cgtattctgg aagttcttca 19380
aattatgtgc atttggatcc acgagttctg tcgacaaccg tgagtcacta gagtgcattg 19440
ggaagaatat acttccaaag ttgaagggca acccattggc tgccaagact attggacgtt 19500
tgttgaggat ggacctaagt acaacacatt gggaaaatat aatggaaagt gaactctggc 19560
aagtagaaca aacggagaat gacattttgc cggcccttcg actgagctat atgtaccttc 19620
ctcagaacct gaaaagatgc ttctcaatct gtgcaatgta tcccaaggat cacaaatttg 19680
agatgcgttt tttagctgat atttggactg cacatggata tgcggagaat ccccaaaaag 19740
catcactcta ctttaatgcc cttgcaaatc ggtcattctt tcagaaagca tctacataca 19800
gttctatgta tgtaattcat gatctgatgc atgatacggc aaagcttgtc tcaaaggatg 19860
agtgcttcat cataaagcat ccgcgtgaca tagatagagt cccttcaact gtccgtcatc 19920
tgtcaatatt tggaaacgga gatattgatt gttctgaact gaagaaaata tgtaacagca 19980
aaaaaataag gaccctggta tgtgatgata aatttaacag cggagcagat tttgctggcg 20040
tgattgactc ttggttcaag gaactgttga aaatccgtgt actgaagttt agagttggca 20100
aagtaaggaa gttacccgag agcatgggca actcaaagca tctgcgttac ttttgtttac 20160
tggggccttc caacttcggc accctacctt caacggtttg tcgtctgcat aagctcagaa 20220
ttatagagcg tgccggtggt gtcattaaaa ggtttccgac aggcttcagt gacctcatca 20280
gcttgcagag aattaattca aaggggttca cttatgacag ggaccaatct gacaaactct 20340
gcctagaatg gtcctttgag gttggagcac ctagtagtga gacggtgaga atgatggaaa 20400
accagatgga agaattacct cactggaata tccaacattt gcatgtagag aattacgaag 20460
gtgaatcttg tcctagttgg ctcaagccca acctcctgcc agggttacgc tcgcttaaat 20520
ttcgtaggtg tggcaaaatc aagagcatac cattttttgt cccgtcagag gagtcacgcg 20580
ataatgcaca aggaccagac aaccttaatc gtcttgaaga gttagtcgtg gatgcttgtt 20640
atgaaatcaa ttggcaaggc tcgatagtct tacctacttc tcttaggaag ctaacacttt 20700
cggcctcagg ctgttccatg gataaccttg tgagctgctt ccgtgacctc acctccctca 20760
ctcatatgga aataaggttc ggcagatcat tgacatccat ccccctcgat gtgtggagta 20820
gcaaccttcc atccctggag gatctatcaa ttatggattg ctaccgcctt gcatcaataa 20880
gcgcctctgg agcaagcagc agcagcggtg ttaaaggatt ctcagccctt gctaagatag 20940
aattttacga atgtggcgca ttactgagcc ttgatgaatt gctgacgcca gactatctac 21000
ctgccgtgaa gacgatttct gttacacact gtgaagcgtt ggcatccgtg tctgtcgaca 21060
ggctccacgg cctggagaac ctggaaattg gacgttgcca caactgtaga atattgtcag 21120
tgccgtcctc tcttaagcag ctcgacttgt atagttctga cggcatagaa ttcatcgacc 21180
tcagcaatag ccagtctcgg agttcttcag cactgaaggt acttagcatt acgggctgtc 21240
caagcctcag gtccatcggt ggtgaagcag ctattgctga aataatgaat gttggtataa 21300
ccgggtgtcc caacctcacg gagatacaac agccactcga gcgcgggtac gtatgatgac 21360
agggaattta attgctgtac tttatatgca catcatttgt tctttacaat atatcctgag 21420
atagctgtat ggtaatctcg ttatgtcgct cgttcagctc atcaacattc ttatctggca 21480
accctaatcc gtgaggtgca catgtgctca tatgtctaca ctactttttt ggagcgggtc 21540
tacactacta ctagtgccta tctagcttat tttaattatt acaaataaag aggatgtata 21600
aattaacccc gataggctta aacagtgaat cacaagcgaa catatagatt ttcgtttaga 21660
taatttgcaa gtacatgacg tatttgtata tggattattt accgaaaatt gcctttggag 21720
ctcggcctcc atggaggctt ccaaattcaa atttaaattt catcaaaatt catattgtta 21780
catttcaaaa aaaaatctga aaaaaataca gatatatatg aagtcataac acatatgtgt 21840
gtaaattttc agggcaaact atgttgaaat gaggcctgtg caaaagaaaa tatagggctt 21900
tttaacacat gatactattc atcttctcag gccatgaatt tgactttttt gtacatatag 21960
tatttcaggg tatttcatcc tgaaaattta cacacataca catacacata actacctttt 22020
ttacttacac aaattttgtc agattttttt gaaaccaaaa agtttgaatt tagaattttt 22080
caaaaatttc gagctctagt gagctcggtc accaaacgtc cgttgtcatt atttactatt 22140
tataaattcc acctaggatt ataatcattc catcaatttg aggttgtatt taaatttaag 22200
aaaaaaatca taggaaaaat aactcaaaga ggatgattac atgaagaaaa aaatgtttgt 22260
tctattcaat tatacttgtt tctactttgt gtcgaaataa atgcattttc catttggaaa 22320
acacttgttt gtgcctgact gatctctcga cgtcatcatc caactccagc gatcgacatg 22380
tgttcctgtg tgggtctgca cgtagttgtc gccatctatc cttatctctt cattcgtgag 22440
gcatgtcaac acattctcgt cccccccccc ccccccccca cacacacaca ccccttgatt 22500
tgttcgtttt ctccaccact gctcaccggc actacccaaa ttaccaccga tgttgcacct 22560
gtctcccgcc accgcgttgc gaggatgcta tttctgccac tgcggctgtt gaagagccta 22620
attttcaaat ttggtgggac ggtgcaaaat cgttgttgca tccaccatcg ctgcaagtag 22680
caggatgtac tcaccggcaa cacctgcttc aaagaaacac aggaggaagc cacgtgcttg 22740
cgaagggtgc ggtgatgcaa gggacgaagg tatggcggcg acaagatctc ggcaccaaca 22800
tcactgttgc atcccctcgc cacaacgtcc gacatcgcta cgagcggtgg agcgagctcg 22860
ccgacgacgc atgtcacaat gcaactctgg agggggttgc atgctacaaa gggtgcaaag 22920
gcgctgcttc gtacggtggc ttggtcgccc gtgctgcgat ggagcatcgc tggggcactg 22980
gtgttgcgat gaagcatcat cgggttgtag atgttgggat gtagcaccga cggtgctgtg 23040
atggagcatt atcggggccg ccggtgttgc aacaaagcat catcgagccg tcggtgcttt 23100
catggagcat tttcgggccg ctggtgttct gactgccggg accgcgagga tccagcatca 23160
caggacccga gcgcccgatg agccgaccta tggcgtcgcg gcggacggat ccgtccatcg 23220
gcctgatgct gtcctctcga gagcggcgga gtgcaatgcg ggctgccatt gcatcctcta 23280
acccacacag gatgacaccc cagtcggcgg atccagagtg gtcggagtat gatccactac 23340
ctgccatgcc gaagaaggcc gaaaactgcc gaaggtgagc tcggatgtcg gaggggagtg 23400
gagtgaagtg gctagggttt gctccggtga gcagatgggg atgaatatat gtggggtcgg 23460
atgggccagc atgggccggg tctgacatgg caggcgtgcc cgggcacccc atatccgccc 23520
catatttggg ttagatatga gaggtgccgg tcagcctggg cgtttgaggc ccatttgagg 23580
cgtccgtcta ggccaaaaaa tcgtgaccgg gcgggccgcc tgggcgtttg aggagggttt 23640
aaggcgcccc gctgtagatg ctcacggttc ccatggactg tccgtgcctg ccgctgctta 23700
tgttagaaca tctatagtcg gaccttctat ttcatcctta aacatctgaa acatttcccc 23760
ggtcagttta ataaagaagg tgaccaaaca ctttctcttt aaacatcctg actttttttt 23820
agagaaaagg cgccggccga gcccgagaag agctcaaacc tagaccgtct ttgaattaac 23880
aaagccatca accagccatg attacagggc agcaacatac acaaaggaaa acaacggaaa 23940
gaaggggtac aaggtgctaa gtagaacaac tcaaggcaac ttcatagcgg gcaagaatag 24000
accacgccgt tgccaacaac cgccactata cctacaccag ctaacctact aaggccacgg 24060
aagtattttg tttgttgaca gccattgaac gaccaaacat gggcgataat gatttgtctc 24120
tgccgccgca aagggccacc atcctttcac ccccaggctc gatgggtgcc gcaccggcgg 24180
ccggcagtgt ggttccccca gaacccaaat ccaggtgtcc aagctgccca aagagagacg 24240
ccaggagcag acccacacgc aacggaggcc ctaccacccg aacatcatca ccggcagcga 24300
caagagataa cgccggctga gagatgtcga acgaaggcgc taaaccacga agaaggccga 24360
aaatggcagc accatacaat ggtcaatgat aggagtcgcc ggatgaccgg cgccgcaaga 24420
aggacaccga tcccttcaca ccaatggccg ggaccaaccg caccagggac cgggcaagcg 24480
ccggagatgt agctggccac gccgtcatca aaccatcgcc acctccacca ccgcttgagc 24540
cgccctcgaa cgacgccttc aggaaggatc acgccaccac agtgtcgtcg tcgcttggaa 24600
cagggggtct gaggctttca cgtgagctcc tgggagggat gaaaggaagg ggttcctctc 24660
tgaagccagc cagtgtgtgc ttccgatggg tggagccggc cggaaatgtg tgatacagtg 24720
tgttcttgag gtgatgtcgc tgttgtgtgt gctcttcacg tgctccgact gtgtcgcaca 24780
caatctgttc gatgaaatgt tgcatttagg aaaaaaaatg agggttaaaa agtaggggtt 24840
gtggttattc tgcagaagaa aaatgtgaaa tcaagtccgc aaacatttga gaggccaatt 24900
tagccaggtg ccctatagat gctcttatgt gggaattatg ttctctgcca tcagcttgtt 24960
accgctcatc tcgcgctgtg ttcccctctc atggcgctcg gtcctgttgt gccaagccat 25020
ccagcgcgtg gcctccaaag catacaaggg gcgtggccgt gtgggaacag gcgactcgaa 25080
gacgttgtca cagtaactta ttattgtagg aatatttgac ccgggttata atttttgttt 25140
cctcaaatgg ttccagatga tttttacttt atcttttaca ttttgtatac gtatagtcaa 25200
ttgcttttga ggatccaata ttaatgcact catatgtgct tacaggacgg ggaggtggtg 25260
gccacgggct gattaattca agtggacgag aagaaatatg aacagcgtct tagatactca 25320
agaggcgaag agagcaccga taaaggagga caagtataca gcaacggtct agtttcaaca 25380
aaaaagagta cacagcggcg gtgtaggggg gccttctcgt ttgcgatcga atgccggaat 25440
aattgctctc aataataatg gaagttgcat catgctgctt gatgtttgtt acatttactt 25500
tgccctgaaa tttgagttgc attgtatgta ttttgctagt tgtgaagaaa ttctattttc 25560
agttccagag atttcaaata atgtgccact aagaacaccg gatttgcttg tttattgcct 25620
tgtaataata tccattgaag tactgtatat tattaagacc cattcattat tattattatt 25680
attattatta gaactagtca tcgacccatg gactagcaga gttgttacat atacaaaaga 25740
catttaactg ctcataaata ctccttcata tgtactctct tatatgcgac cccattgagt 25800
gtggatcaac gactttttgt gttctgaaca atatcaatag atccatgtga aacgtagttc 25860
ttaaatatgg ttcttgtata taccccagaa caagagtacc caacttgtgt aatgtagtac 25920
cattcagaga acgacctagc gccctccccc tcgcgtcgcc tcccctcggg cggcgccagg 25980
gcccccgaaa ccctagcgcc gccagctccc tcttgccgcc tcctcctgcc gccgctgccg 26040
ccggcgtggg ccgtggtggc agcgggcccg gtgccgaagg tcttcgggcg ggtggatgca 26100
gcggatccca tctgaggcgg cgggggcggc tcctctcgtg agatccaacg gcagcggctg 26160
ggacggcgaa tccgggctgt gcggcggggg ccggagcacc atcgccggcg gagtggcggc 26220
cctgcatgga tggcggcggt ggcagcaccc catctggcgg ggtgggctgt cctggttgtg 26280
atggtgacgg cggtcactgc ggatccaacg ccccgtttgg atccgtcgcg gggaggcctt 26340
ccgccgaccg atggcgcgtg gagggaccgg tgaggagatg ccggatctcc gccggagtac 26400
cgacggcgac atacgtcttc atggcgaggt ttcgctcggt tccagccggc cccgaggtcg 26460
ggaagacgtg gcttacggtg aaaaccgcgc ctgactgcgg tcttgacgga cgatggcggc 26520
gtctcagacg tcgttccctt ttggaggcat cgtcgttgca ggtcacatca acctgatcgg 26580
gaagttccgg gggaaaccct agatctgggt ctaccggatc ggacgatgac ggggccttcg 26640
gtatcgttct ccctcatggg ggcatcgtct tggagcaagt gctggttgga ggggacaaga 26700
ggaggagcgg tgatttatct ggtccattga tgacgacgga cctcggcggc atggcgcagt 26760
ggagacccgg cgcctgatgt gcggagatgg actcgcgcag gagggtgacg ctgcctggcg 26820
tcgtggtggc gtcggcggca gctagaccgg gcaaggttga tgcagcagta cagctctgaa 26880
gatggattgg tggcaggtgg ctgcggcggc ctcatacccg gcaggcgtcc tggttgagaa 26940
gcacgccgga ctaatgggtg cccacaccag gcaggcgtcc tggttgggac ctcaagtctt 27000
agatgttagg tttggctgca aggtctattt ggtattaggc ccagactttc agcgccccta 27060
catcaactgg atagggatag cgatagatgt tgcttagttg gtggctttag gcttattgtt 27120
gtatgactct gtaaggtctt gtgtcaataa ttaataaaat agccgtatgc attgtccaga 27180
tgcagaggcc gggggtcgtc ctccttttct aaaaaaaaaa tcaaagaacg acctatacct 27240
aacaaaacca gatcaggtac caaaagtgtc agtaccatat taatcactgg tgttattaat 27300
tccaaaatct gggcgtgtaa tttcaaacag aacatgatac acactgactg aaagaatcag 27360
atcacttaca tcaaaatcct atctacaggt tgaaagaaca tataagctta gaagatctgc 27420
ttttcaaaag aggaaaaatc agccaacaca tgtcctgcaa gcaacaatgt gcaagtgtat 27480
aacacataca tacatggcaa acttgtaaca atttggggaa tgaagtatca ttttttcacc 27540
ccctatttat ttgtggagca aacatcaact ctcatttcca taagaagaaa gcaacaaat 27599
<210> 10
<211> 22
<212> DNA
<213> BD1-L
<400> 10
tcgtacactg catccttgat ct 22
<210> 11
<211> 22
<212> DNA
<213> BD1-R
<400> 11
aggctgaaga ctcgtatttg ga 22
<210> 12
<211> 42
<212> DNA
<213> HBAU194F
<400> 12
tctagaggat ccccgggtac catggccatg atcttaccag cc 42
<210> 13
<211> 41
<212> DNA
<213> HBAU194R
<400> 13
attcgagctc tctagaacta gtttaatcag cccgtggcca c 41
<210> 14
<211> 20
<212> DNA
<213> 22-5OE-1L
<400> 14
cttctgcagg tcgactctag 20
<210> 15
<211> 23
<212> DNA
<213> 22-5OE-1R
<400> 15
taattattgg gctcatcaca gtc 23
Claims (9)
1.小麦抗叶锈病蛋白,其特征在于:其氨基酸序列如SEQ ID NO:1所示。
2.一种小麦抗叶锈病蛋白,其特征在于:所述小麦抗叶锈病蛋白为在SEQ ID NO:1序列的N端或/和C端连接蛋白标签。
3.如权利要求2所述的小麦抗叶锈病蛋白,其特征在于:所述蛋白标签为序列为SEQ IDNO:2的Poly-Arg、序列为SEQ ID NO:3的Poly-His、序列为SEQ ID NO:4的FLAG、序列为SEQID NO:5的Strep-tag II、序列为SEQ ID NO:6的c-myc和序列为SEQ ID NO:7的HA中的任意一种。
4.编码权利要求1-3任一项所述的小麦抗叶锈病蛋白的核酸分子。
5.包含权利要求4所述的核酸分子及其表达调控元件 的重组表达载体。
6.过表达权利要求1-3任一项所述的小麦抗叶锈病蛋白在提高小麦抗叶锈病性能中的应用。
7.权利要求4所述的编码所述小麦抗叶锈病蛋白的核酸分子在抗叶锈病小麦品种遗传育种中的应用。
8.一种用于扩增得到编码权利要求1-3任一项所述的小麦抗叶锈病蛋白的基因的引物,其特征在于:其上游引物为HBAU194F:5´-TCTAGAGGATCCCCGGGTACCATGGCCATGATCTTACCAGCC-3´;下游引物为HBAU194R:5´-ATTCGAGCTCTCTAGAACTAGTTTAATCAGCCCGTGGCCAC-3´。
9.一种用于鉴定编码权利要求1-3任一项所述的小麦抗叶锈病蛋白的基因的引物,其特征在于:其上游引物为BD1-L:5´-TCGTACACTGCATCCTTGATCT-3´;下游引物为BD1-R:5´-AGGCTGAAGACTCGTATTTGGA-3´。
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