CN112359130B - 一种检测小麦秆锈病抗性基因SrTm5的诊断性分子标记及其育种应用 - Google Patents
一种检测小麦秆锈病抗性基因SrTm5的诊断性分子标记及其育种应用 Download PDFInfo
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Abstract
本发明属于植物生物技术和小麦分子育种领域，具体涉及一种检测小麦秆锈病抗性基因SrTm5的诊断性分子标记及其育种应用。本发明提供的分子标记TM5‑TF2R2‑673bp结合PCR的方法，可准确检测材料中是否存在抗秆锈病基因SrTm5，操作简单，成本低廉，可用于育种的早代选择，促进了对SrTm5基因的育种应用，在育种、科研、生产实践中具有广泛的应用前景和重大的经济价值。
Description
技术领域
本发明属于植物生物技术和小麦分子育种领域，具体涉及一种检测抗秆锈病基因SrTm5诊断性分子标记的开发及其育种应用。
背景技术
秆锈病是危害小麦产量最严重的真菌病害之一，该病主要发生在叶片、茎秆和穗部，发病时严重影响小麦产量。目前控制小麦秆锈病最为经济有效的方法是利用抗秆锈病基因来培育抗病小麦品种。自20世纪70年代，由于抗秆锈病基因在育种中的广泛使用以及寄主小檗属植物的大规模去除，该病害得到了有效控制。
但是，1998年在非洲的乌干达发现了一个致病性极强的秆锈病新小种TTKSK，俗称为Ug99，它使得育种中广泛使用的Sr31和Sr38等抗秆锈病基因都丧失了抗性。目前，Ug99已经进化出10多个变种，并从非洲传到了亚洲，已经扩散至十几个国家。目前，全球超过80％的小麦品种(系)对Ug99没有抗性，因而Ug99的出现使得世界小麦生产又重新处于秆锈病的威胁之下。2006年，我国有118份小麦品种送至肯尼亚进行Ug99抗病性鉴定，结果显示高感品种占98.3％，所以一旦Ug99传入我国，会对我国小麦生产造成严重影响。
目前只有少数几个小麦抗秆锈病基因被成功克隆，且皆为国外研究单位完成。因此，克隆Ug99抗性基因并转育到我国小麦主栽品种，显得尤为重要。抗秆锈病基因SrTm5来源于一粒小麦(T.monococcum)材料PI 306540，对秆锈菌小种TTKSK(Ug99)、TTKST、PKSSC和MCCFC等表现出良好的抗性。目前，在育种中只能使用SrTm5的连锁标记进行分子标记辅助选择，缺少基因内部功能分子标记，这样难以准确、高效地检测小麦抗秆锈病基因SrTm5，极大限制了该基因在小麦育种中的利用，所以亟需对抗秆锈病基因SrTm5进行克隆，并且根据其序列设计基因内高效的诊断性分子标记。
发明内容
本发明提供了一种小麦抗秆锈病基因SrTm5诊断性分子标记的开发与应用，用于准确鉴定小麦材料是否含有抗秆锈病基因SrTm5，可用于育种的早代选择，解决了小麦育种中难以准确检测和有效利用抗秆锈病基因SrTm5的问题，具有广泛的应用前景和重大的经济价值。
本发明提供了一种用于检测小麦抗秆锈病基因SrTm5的诊断性分子标记TM5-TF2R2-673bp，其为显性标记，上下游引物序列分别为：
5’-GCACTGAGACTCCTCGGTGATGT-3’(SEQ ID NO：5)
5’-CACTCATATTACCCCCTTCCTTACC-3’(SEQ ID NO：6)
本发明提供了一种小麦抗秆锈病基因SrTm5的鉴定方法，包括以下操作步骤：
a)提取待测样本DNA；
b)用检测引物对(SEQ ID NO：5和6)进行PCR扩增；
c)对PCR产物进行电泳分析。
若PCR产物可检测到一条673bp的特异条带，则表明待测小麦样本含有抗秆锈病基因SrTm5，样本具有抗秆锈病性状；若无PCR产物，则判断待测小麦材料不含有抗秆锈病基因SrTm5。
优选地，在实施例中，上述鉴定方法，步骤a的DNA提取方式可以是CTAB法。
优选地，在实施例中，上述鉴定方法，步骤b的PCR条件可以是94℃3分钟；94℃30秒、60℃30秒、72℃30秒，共35个循环；72℃10分钟。
优选地，在实施例中，上述鉴定方法，步骤b的PCR反应体系包含100ng DNA模板，12.5μL2×Es Taq MasterMix(康为世纪公司提供，商业化产品，内含Es Taq DNApolymerase，MgCl2和dNTP)，5μM引物。
本发明还提供了一种抗秆锈病基因SrTm5检测的PCR试剂盒，其特征在于所述试剂盒包含一对检测引物，其引物的序列如SEQ ID NO：5和6所示，用于检测小麦材料中是否含有SrTm5基因，以判断小麦材料是否为抗秆锈病材料。
本发明还提供了一种试剂盒的使用方法，所述试剂盒的使用方法包括如下步骤：
(a)提取待测样本DNA；
(b)用检测引物对(SEQ ID NO：5和6)进行PCR扩增；
(c)对PCR产物进行电泳分析。
本发明还提供了一种抗秆锈病小麦材料的分子标记辅助选育方法：其特征在于所述使用方法包括如下步骤：
(a)提取待测样本DNA；
(b)用检测引物对进行PCR扩增；和
(c)对PCR产物进行电泳分析。
若PCR扩增产物可检测出一条673bp的特异条带，则表明待测小麦样本含有抗秆锈病基因SrTm5，样本具有抗秆锈病性状；若无PCR扩增产物，则判断待测小麦材料不含有抗秆锈病基因SrTm5。
本发明相对于现有技术，具有如下的优点及效果：
(1)本发明利用图位克隆方法克隆了抗秆锈病基因SrTm5，并在此基础上开发了基因SrTm5的诊断性分子标记TM5-TF2R2-673bp，该标记是位于基因内部的特异性分子标记。
(2)简便、准确、高效。TM5-TF2R2-673bp是显性分子标记，能够通过PCR方法准确的鉴定小麦材料中是否存在SrTm5基因，从而加快对抗秆锈病基因SrTm5的育种利用。该鉴定操作方法简单，耗费较低，且产物特异性好。
附图说明
图1为不含有基因SrTm5的感病家系和含有基因SrTm5的抗病家系对秆锈菌小种PKSSC和TTKSK的表型反应。
图2为小麦抗秆锈病基因SrTm5的精细定位和克隆。
图3为抗秆锈病基因SrTm5的结构示意图，其中在第二个内含子中有一个15Kb的反转座子插入。
图4为诊断性分子标记TM5-TF2R2-673bp在PI 306540和PI 272557杂交产生的F4分离群体中扩增的PCR电泳胶图。其中泳道1为抗病亲本PI 306540，泳道2为感病亲本PI272557，泳道3-22依次为F4分离群体的部分单株。
图5为诊断性分子标记TM5-TF2R2-673bp在部分二倍体、四倍体和六倍体小麦品种中扩增的PCR电泳胶图。
具体实施方式
下述实施例用于进一步描述、解释本发明，以便于更好地理解本发明，但并不限定本发明。下述实施例中的实验方法，如无特殊说明，均为常规方法，所用到的试剂等如无特殊说明，均可通过商业途径获得。
下面实施例中用到的小麦叶片基因组DNA的提取方法--CTAB法步骤如下：
(1)将适量叶片装入2mL圆底离心管中，加入钢珠，置于液氮中速冻，使用研磨仪将叶片磨为粉末；
(2)加入500μL CTAB提取液，65℃水浴30分钟，期间颠倒混匀两次；
(3)取出离心管冷却至室温，加入500μL的氯仿:异戊醇(24:1)混合液，充分摇匀，静置15分钟，于12000rpm离心10分钟；
(4)吸取上清500μL，加入预冷的异丙醇500μL，摇匀后，4℃静置5min，12000rpm离心10分钟；
(5)弃上清，加入600μL70％乙醇洗涤，然后12000rpm离心5min；
(6)弃上清，风干，加入200μL ddH2O溶解DNA。
实施例1：苗期秆锈病抗性鉴定
在人工气候室种植待鉴定的一粒小麦抗病亲本PI 306540和感病亲本PI 272557，以及二者杂交产生的F4分离群体(F4群体来源于杂合的F3单株。由于PI 306540含有多个抗秆锈病基因，所以需要选择只含有基因SrTm5的杂合F3单株来构建F4分离群体)。
人工气候室条件设置：光周期16小时，白天22℃，夜晚20℃，湿度80-90％。当幼苗长至两叶一心期，采用人工扫抹法将新鲜秆锈菌孢子接种到上述幼苗上。接菌后黑暗、保湿处理24小时。接菌12天后对材料进行秆锈病抗性鉴定，鉴定方法按照0-4级分级标准进行(注：0级为免疫；0；为近免疫；1级为高抗；2级为中抗；3级为中感；4级为高感)。
如图1所示，接菌12天后，不含有基因SrTm5和含有基因SrTm5的家系对秆锈菌小种PKSSC和TTKSK表现出的不同的发病表型。
结果表明：含有基因SrTm5的家系表现为近免疫或高抗(IT＝0；或1)，不含有基因SrTm5的家系表现为中感或高感(IT＝3或4)。表型数据用于后续分析分子标记与基因SrTm5的连锁关系，以及基因的图位克隆。
实施例2：秆锈病抗性基因SrTm5的精细定位、图位克隆
为了精细定位抗秆锈病基因SrTm5，申请人利用中国春参考基因组(https://wheat-urgi.versailles.inra.fr/Seq-Repository/BLAST)，在染色体7AL上开发CAPs或InDel标记。通过利用包含1132个单株的PI 306540X PI 272557的F4分离群体，将抗秆锈病基因SrTm5定位在CAPs标记Pku7A4995F2R2和标记Pku7A5001F6R6之间0.13cM区域内(图2)。该区域在中国春参考基因组中物理区间为289Kb，只含有5个候选基因：TraesCS7A02G499600，TraesCS7A02G499700，TraesCS7A02G499800，TraesCS7A02G499900和TraesCS7A02G500000。基因注释表明，在这些基因中只有基因TraesCS7A02G499600编码NLR(核苷酸结合结构域及富含亮氨酸重复序列，Nucleotide-binding,leucine-rich repeat)蛋白，其它基因都与抗病无关，推测该基因是SrTm5的唯一候选基因。
为了得到TraesCS7A02G499600在抗病亲本PI 306540中的直系同源基因，申请人构建了PI 306540的BAC文库，利用标记筛选，得到阳性克隆BAC-Tm8。对阳性克隆进行WIDESEQ测序，对序列进行组装拼接，得到基因全长核苷酸序列(如SEQ ID NO：1所示)，编码区核苷酸序列(如SEQ ID NO：2所示)，以及其编码蛋白质(如SEQ ID NO：3所示)。
SrTm5基因全长为19715bp，由4个外显子组成，编码蛋白包含938个氨基酸，基因结构如图3所示(方框代表外显子，实线代表内含子，三角形代表反转座子的插入位点)。分析了抗病亲本PI 306540中SrTm5的基因序列与感病亲本PI 272557之间的多态性，结果表明二者序列的相似性为92％。在抗病亲本PI 306540中，基因SrTm5的第二个内含子存在一个15Kb的反转座子插入，而该插入在PI 272557和中国春中都不存在。
实施例3：秆锈病抗性基因SrTm5的诊断性分子标记的开发
利用primer5软件，申请人设计了秆锈病抗性基因SrTm5的诊断性分子标记。具体地，利用第二个内含子序列设计了正向引物(如SEQ ID NO：5所示)，利用插入的反转座子序列设计了反向引物(如SEQ ID NO：6所示)，并将这一诊断性分子标记命名为TM5-TF2R2-673bp，扩增产物序列如SEQ ID NO：4所示。
实施例4：SrTm5诊断性分子标记的应用
1)利用该诊断性分子标记TM5-TF2R2-673bp，对抗病亲本PI 306540、感病亲本PI272557以及它们杂交产生的F4分离群体的192个单株进行基因型和表型鉴定。
具体地，根据设计的分子标记TM5-TF2R2-673bp，对上述小麦样本用CTAB法提取基因组DNA进行PCR扩增，具体反应体系如下：
DNA模板(50ng/μL)2.0μL
2×Es TaqMasterMix(Dye)12.5μL(康为世纪公司)
10μM引物0.5μL
加ddH2O补齐至25μL
PCR扩增反应程序：94℃预变性3分钟，94℃变性30秒，60℃退火30秒，72℃延伸30秒，35个循环，72℃延伸10分钟。
扩增产物通过1.5％的琼脂糖凝胶电泳分析，电泳缓冲液为1×TAE，电泳仪的恒定电压为100V。凝胶采用EB显色，使用BIO-RAD成像仪进行拍照。部分结果如图4所示，单株3、4、5、8、9、11、12、13、14、15、18、20、22扩增出与PI 306540(泳道1)一样的条带，说明含有基因SrTm5；而单株6、7、10、16、17、19、21与PI 272557(泳道2)一样，无PCR扩增产物，说明不含有基因SrTm5。
利用秆锈菌小种PKSSC对抗、感亲本和上述192个F4单株进行抗病性鉴定，结果显示，得到的表型与基因型完全一致。
2)利用分子标记TM5-TF2R2-673bp对93份(包含抗病亲本PI 306540和感病亲本PI272557)一粒小麦T.monococcum材料进行基因型和表型鉴定。
具体地，根据设计的分子标记TM5-TF2R2-673bp，对上述小麦材料采用CTAB法提取基因组DNA进行PCR扩增，具体反应体系如下：
DNA模板(50ng/μL)2.0μL
2×Es TaqMasterMix(Dye)12.5μL(康为世纪公司)
10μM引物0.5μL
加ddH2O补齐至25μL
PCR扩增反应程序：94℃预变性3分钟，94℃变性30秒，60℃退火30秒，72℃延伸30秒，35个循环，72℃延伸10分钟。
扩增产物通过1.5％的琼脂糖凝胶电泳分析，电泳缓冲液为1×TAE，电泳仪的恒定电压为100V。凝胶采用EB显色，使用BIO-RAD成像仪进行拍照。
结果显示其中有15份材料PI 277130、PI 306545、PI 355538、CItr 17657、PI277131-2、PI 306544、PI 306547、PI 352480、PI 355536、PI 355541、PI 377668、PI435000、PI 362610、PI 435001和PI 221414能扩增出与抗病亲本PI 306540一样的目标条带，说明这15份T.monococcum材料都含有基因SrTm5。我们用秆锈菌小种TTKSK进行抗病性鉴定，结果显示这15份材料表型与基因型鉴定结果完全一致，这说明本发明所提供的分子标记TM5-TF2R2-673bp可以高效、精确的用于鉴定小麦材料中是否含有抗秆锈病基因SrTm5，统计结果如表1所示。
3)利用分子标记TM5-TF2R2-673bp对来自中国、美国等国家的58份六倍体小麦(T.aestivum)和22份四倍体小麦(T.turgidum)进行基因型鉴定。
具体地，根据设计的分子标记TM5-TF2R2-673bp，对上述小麦材料用CTAB法提取基因组DNA进行PCR扩增，具体反应体系如下：
DNA模板(50ng/μL)2.0μL
2×Es TaqMasterMix(Dye)12.5μL(康为世纪公司提供)
10μM引物0.5μL
加ddH2O补齐至25μL
PCR扩增反应程序：94℃预变性3分钟，94℃变性30秒，60℃退火30秒，72℃延伸30秒，35个循环，72℃延伸10分钟。
扩增产物通过1.5％的琼脂糖凝胶电泳分析，电泳缓冲液为1×TAE，电泳仪的恒定电压为100V。凝胶采用EB显色，使用BIO-RAD成像仪进行拍照。
结果表明，这些材料均未扩增出673bp的目标条带，表明这些材料都不含有基因SrTm5(如表1所示)。图5为分子标记TM5-TF2R2-673bp在部分二倍体、四倍体和六倍体小麦品种中扩增的电泳胶图，其中泳道编号1-15依次为小偃22、中国春、川麦42、Fielder、8155B、Rusty、D447、Kronos、PI 272557、PI 418580、PI 487249、PI 427507、PI 306540、PI377668、PI 362610。其中含有基因SrTm5的抗病材料PI 306540(泳道13)、PI 377668(泳道14)和PI 362610(泳道15)可以扩增出一条673bp的条带，而在其它未携带SrTm5基因的小麦材料中均未扩增出条带。这表明该标记可在不同小麦材料样品中进行PCR扩增，从而可以精确、快速地区分小麦材料中是否携带抗秆锈病基因SrTm5。
表1：诊断性分子标记TM5-TF2R2-673bp鉴定不同的二倍体、四倍体和六倍体小麦材料
以上内容通过一般性的说明和具体实施例对本发明进行了详细的描述，但本发明并不限制于以上描述的具体实施例。对于本领域内的技术人员而言，对本发明进行某些修改和替代是轻而易举的，因此在不脱离本发明的基本内容而做的任何变换和修改都涵盖在本发明要求保护的范围之内。
序列表
<110> 北京大学现代农业研究院
<120> 一种检测小麦秆锈病抗性基因 SrTm5 的诊断性分子标记及其育种应用
<160> 6
<170> SIPOSequenceListing 1.0
<210> 1
<211> 19715
<212> DNA
<213> 一粒小麦(Triticum monococcum)
<400> 1
atggcggaag ttctggtgag cgcatccaca ggggcaatgg gttccctcct gcggaagttg 60
ggagctatgt tgacggatga atacaagctg ctcaagaacg tccgtgggga catcaagttc 120
ctcaaggacg agctggaggt catgtgcgcg ttcctcctca agatgtcaga tgtggaggag 180
cctgacgaac caacaaagct gcgtgtgacg gcggtgcggg agatgtccta taagatagag 240
gacaacatcg acaagttcat ggtcctcgtg gaacacgagt cctcctgctc cgaggcggct 300
catggcgtcg ccaagctcat ggacaagtgc aagaacttgc tgccggacat caagacccgc 360
cgcaggatcg ccaaggaggt caaagacatc aagaaagaaa ttaaggatgt cagcgacaga 420
ttttcaaggt accagatctt gttctcttta cagtactatt agtacctacc atcagatttg 480
tgaccgattt gttgccactc aaggtacaag attgacgagt cctcctcctc tatgccggca 540
aaaggcaagg tcgaccctcg acttcgtgca gtctacaaag acgccgcaga gctcgtcgga 600
attgatggac caaaggatga acttgtcaag tggctgaatg agaaagaggg ccaatcactg 660
aaatctgtct ctattgttgg atatggaggg ttgggcaaga ccacgctcgc caaccagatc 720
cgggtcaacc ttggagcgac cttcgactgc ggggcttttg tctcgatttc acgcaagcct 780
gacatgaagg cgatattaag atctatatta tcgcaaatca ccaagaaaga cgaggcttgc 840
tctagattag atgatataca actcatcatc gacaagattc gagaattcct ccaagacaca 900
aggtatgtat gtgagatcta ctacgaagta acattgctta attagctcat gcataatccg 960
ttgcctagaa ttttgctaat gaagattgct ttgtgtgatt tactttcgtc ctcaacttgt 1020
atgactgcgc ttaaactttc aagtcattgt ttcttgcaac ttatgtcgat ctgttcacta 1080
attaagatat agctaaaatg tgtttgccaa tttctgtcgc cacacatttt ctaatttgat 1140
tgccacaggt ttgttaaact tgcggctatg ttctgtttat taaataattg tgacatggcg 1200
tcagtcaaaa caataaagta catgcaattc cctctctttt gtacaagata aatatccgca 1260
cgcactacat gactgcaaat atatctatct tctaaggacc caaagcactt gcatgtgcag 1320
accattgtta ccggatatgc agtacattag gtttgtgatt agccacttta agaacatagg 1380
gtttctagag gctataaacc tacggttatc agtattataa tcttgaatgg atatcaaaca 1440
tccgttgaag aatctatgcc cataacacaa aaagagaaaa gggaagaatt aatcatagtc 1500
agatcggacc agcctgacta gcgtaacata atgacaacag taggttgttg cgtctttcct 1560
ggtttcgtag caatttgcat gacgaattga ggcgcagagc caccactcaa tatgtggtcc 1620
taccaatcat gaggaaataa cttgttacca aaatgacaaa agaaaatagg tagtccccat 1680
cattgtttca atatctatcc tcttgcgaac atgctattgg cagtggcaaa acattgtacc 1740
ttgccactgg caacagctgc ctttactcgg cctgagcatt catatttttt tcctcttggt 1800
tgttaaatct tgctattttt cgcttgcctt ctatggcaat cttttttttt cttatattgc 1860
caccaaattg tatctaagag caactctaac acgccgacgc aatttttgtc catttagatc 1920
ggccaagcgg acggggtgtc cggttttgcc tttgggtggt gcgcctacgc attagtccgc 1980
gtgggcattt aaaaaactta aaaagcttaa aataattaaa tataaataaa aatgcatatt 2040
aaagcccgtc aaagtccaca taattaaaca taatacatta atatattaat aaaaaacagc 2100
cgccccagcc cgcagcccta ggcgtcttcg tcggaggagt gggtgaggtc gaagaatgcc 2160
ggcagctcag cccacgggaa ggcctgctgg tgggcagcca gcaccgcctg ctccgctgtc 2220
tgcggcggcg gcggccctac ctggcaccgc gcacgcctcc tcctcgcgct cctgccgctc 2280
ctcctcatcg cacgcccaaa gaagttttgg cgctcctccc cctcgggttc ctgctgcatg 2340
cggacctagc cgtcgtaccg ctcctcctcc agcgtgcggt ccttccacgc cacgaggtag 2400
gccgcctcct cctccatcac gcacagccaa atgggcggct tgctcatccg cttgcagacg 2460
atgccctgcc attggtaaac gccggtgggc gcgggtggtg ctagcgctgg tgcaggcggt 2520
gacgatgtgg ctggtggtcc cggcaggtag acactatagc tggcggccga tagctgcagc 2580
gtgtcatcga ggccagacca ctaggagtcc tcgcactgag actcctcggt gatgtgatgg 2640
gccccctcct ccaatgcctt gaggaacgcc gcctcctcct caacctacta tttgaccacc 2700
ggtggcggcg gtggagggca ggcttcgagc tggacggcat cgcggaggcg cacatggtgc 2760
tcgatggtga accagacttc tcagttaggg gagtgtaaca ccctcgatgc gactatatct 2820
cccacgtgtc gaggcacgac ttagaggcat aatcgcattg aaggcataag tcgcaagtga 2880
ggcaatcttc acacaaccca tgtacatata aatataaagg tggaataata ataataattg 2940
gcttacactc gccacgtcac atcaaggtac aaaaatagca ttcatcatac aaacactcat 3000
ggcccgacta cggcgccaaa atagaaaaga ctccccaaca tgcgactaat ccctgaatcg 3060
aaccccgagc aggcaccact actgatcatc aggaaaagaa acatagtatc gctgagagtc 3120
ctcgtcgtac tcccacttca gctcgaacgc gccatccgga ggagtatcat cggtccctgc 3180
atctggtgta atagtaatct gtgagccaca aggactcagc aatctcgcac cctcgcgatc 3240
aagactattt aagcttaata ggtaaggaag ggggtaatat gagtggagct gcagcaagcg 3300
actagcatat atggtggcta acttattcgc aaaagagagc gagaagagaa ggcaaggcac 3360
gagcggaaaa ctaagtggag cagactacgc aaacataact ccaacgccgt gtccacttcc 3420
cggactccgc cgagaagagg ccatcacggc aacacacgca gtagattcat tttaattaat 3480
taagttcaag ttatctacaa ccggacatta acaaatttcc atctgcccat aaccgcgggc 3540
acggctttcg aaagatcaat ccctgcaggg gggtcccaac ttagcccata acaagctctc 3600
acggtcaacg aaggaataga cctccaccca ggacccaccg atcagactcg gtatcccggt 3660
acaacaagat gattcgacag gtcaaaacta gtccagcaac accgcccgaa tgcgccgaca 3720
aatcccgata ggagctgcac atatctcttt ctcagggcac actcagattg tctaaacttc 3780
cggtaggcca gcccagagtt gcccctggtg gccaccggcg gctgacgttc cggaccaaca 3840
ctcagaggag cactggcccg tggggggcta aaataaagat gaccctcggg ctccggaaac 3900
ccaagggaaa gaaaaggtta ggtggcaaat ggtaaaacca atgttgggca ttgctggaaa 3960
agctttaatc aaggcgaact atcaaggggt tcccattata gcccaaccgc gtaagggacg 4020
caacaaaccg ggaacataac accgatatga cggaaactag ggcggcaaga gtggaacaaa 4080
acactaggcg agaggccgag ccttccaccc tttaccaagt atatagataa gtaaagataa 4140
catagcaata taatgatatc ccaacaagta aataaatgat ccaacaagga acggctccaa 4200
tcttcacctg caactagcaa cgctataaga aggggcttga gcaaagcggt aacatagcca 4260
atcaacggtt tgctaggaca aaggtgggtt agaggctcaa tcatggcaat agggaggctg 4320
acaagcaagt ggtaggcatc gtagcattgg caaagcaaaa gagcgagcaa actagcatag 4380
caaagatagt agtgatttca agggtgtgat catcttgcct gcacagttgt cagaattgac 4440
tggatcctca aaagcaaact caacgggctc ctcgttagcg aactcgtctc ccggctctac 4500
ccaacaacac aaacaagcaa caaggataca atcaaccacg tgcaagggca accaataaga 4560
tgcaaagatg atatgctatg cgggatgcga tgcgggatgc aagaatgcaa catgtaacag 4620
gaaaatgcat gaacctggcc tcaacttgga aatccaagtg tgccactgga tagaggagaa 4680
gaaatcccct gaaaacgata taaagatcat ggcaatcgga gctacggtta agaaatggca 4740
agcgttttac taaacgaacc gatctgcgat ttacagcaag taggctccta aatgcaataa 4800
aatgaacatg ccacagccac caaacatgaa aacaaattgc atggcagtga tgtactcatc 4860
aagcttaaca aatcatcatg actgaggcat aggcaaattc atccattaag ggcttcaaac 4920
aagcatggca aaactgaata agcaacacag atttcagact tagcgaaatt aacgctagtc 4980
tgaaatttca gaccacgatg ctctctttgg agcagcaaaa caacatgaag gaaaacctga 5040
acatggcaag taagagcatg gcatggagct actcaacaag cttaacaaaa cacccaaaat 5100
gacctggggc caaaagggtt cacaaaatac tctaccgagc acacgaacat tgatgaaaca 5160
caatcagttt tcagacttag taaaaactgg gacatgctga aataataacc cacggaggca 5220
tgtaaacgag ctcgatgctc tcactacggt gcaagtcatg gcaaggaaag catatatccg 5280
ttaataaggc acaaaatgct agctagacat ggcaagaaca atggcatagc atgcatggat 5340
cactagcaat aactccggca aaatcgcaaa agagttaacg atctgcccaa aacagcaact 5400
atccaaaagt tgagctcgat tgagtcaacc tagggggctc caacataaca agtaaagaca 5460
tggatggatt aagcacaaca agcataacaa atcactctta ctgatcatcc tcaaaagagg 5520
cacggatcac taggaaacaa gccggacata tagcatcgtg aactaaaacg gctcagactt 5580
agtgaaaatc actaagtccc tgaaatctgt aatctcaggt agcccccttc gcaagcttgc 5640
acaagacaac atacacatcc taaaaataca tggaaggcac ctctggaaag aagataacat 5700
gcttaacaat tcatcccaaa ggggcacagg catatcatgc acggattaaa tatggcaaaa 5760
atggcaaaag tgcatgttaa actaacggat ctgcaattta actcaacatg cctccttcca 5820
acagcatttt gggcagcaag atgacctcaa atgcaaatga tgcaatggaa tgaaatgatg 5880
tactcgtcga gacgaagaat ttgacatatc atacgcccaa atcggagcta cggatgcaaa 5940
gttacgggac cccgaacaga ggcatatgga ctgaaaattc ctgggactta gagaaaaatg 6000
caacctccca gatctagggt ttccagattt gcacgcaatc cgaggcggcg gggaggcaga 6060
tcggacacgt ccggcgcgcg gaaagagagg atttagggtt tcggggaagg agatccgaga 6120
tttagaggag ggggtctata tataggcata gagggagcta ggagagtcca aatgaggtgc 6180
ggttttcgcc cacacgatcg tgatcaaacg accgagacga tggagctgag tttggtgggt 6240
tttgggccaa attggagggg tgttgggctg cacacacacg aggccttttc ggtccctcgg 6300
ttaaccgttg gagtatcaaa cgaagtccaa atgacccgaa acttgacagg cggtctaccg 6360
gtagtaaacc aaggccgcat ggcaagtctc ggcccaatcc ggaaatgttt aatccccaca 6420
cacgaaagaa agctagaaat ggccaccgga ggagaacgaa gcgccggaat gcaaaacgga 6480
caacggggaa aatgctcgga tgcatgagat gaacatgtat gcgaatgcaa tgcacgtgat 6540
gacatgatat gagatgcacg agaacaaaaa caacacacga agacaaagac ccgaacccga 6600
gaattaaata taacttaatg ccggaaacgg catgagttgg aatacaattg gggaaggtta 6660
catatggggc gttacaacac tccaccacta cgaaaagatc tcgtcccaag atctaggact 6720
gaaagaactc cgggtactca gaacggaggt gatcctcgcg ttcccaggta gctttacggt 6780
cggaatggtg tgaccacttg actttgagaa acttgattga cttgttgtga gttttgcgtt 6840
cagtctcttc aaggattgca actgggtgct cacggtagga gagatcatct tggagctcaa 6900
tgtcctcgaa gttgacggtg cgctcaggcg tcttgaagca cttacggagt tgcgagacgt 6960
ggaacacatc atgaacattg gcaaaggttg caggaagctc gagttgatag gcgaggtcgc 7020
ctctcttgct gacgatcttg aaaggcccga cgtatcttgg ggcaagcttc cctttgatac 7080
cgaagcgacg agtccctttc ataggagaga cgcggaggta gacatgatct ccgatttcga 7140
aagccaaatc acggtgcttg ctgtcgtagt agctcttctg acgagactgg gctgctttga 7200
ggttatcacg aatgagcttg cacatttctt ctgcttctgt gattaggtca ttacccagca 7260
gctgacgttc accggtttca gaccagttga gaggtgtacg gcacttccta ccatacaaaa 7320
tttcaaacgg ggccttgccg gagcttgctt ggaaactgtt gttgtacgag aattcagcat 7380
aaggaaggca atcctcccac ttcatgccga aggagataac acaagccctg agcatatctt 7440
caagaatctg gttgacacgc tcgacttgac cgcttgtttg aggatggaaa gcagtgctga 7500
agcgaatgtt ggtgcccatg gccttctgaa aagaatccca gaacttggag gtaaagatgc 7560
tgccacggtc tgaagatatc acttgaggaa taccgtgcag agagacaatg cgagaggtgt 7620
agagctccgc caattgagct gcagtgattg actctttgat aggcaagaaa tgagccactt 7680
tggtcagctt gtcgatgaca acgaatatag catcattgcc acgcttggac ttagaaaacc 7740
cagtcacgaa gtccatttca atgtggtcaa acttccattc tggaatggcg agaggttgga 7800
ggagacctgc tggcctttgg tgttctgctt tcactcttct gcagacatca cattcattca 7860
cgaattgagc gatctcgcgc ttcattcgag tccaccaata agtttgcttc aggtcttgat 7920
acatcttcgt gctcccaggg tggatggaga ggagagagtt gtgagcctca ttcatgatca 7980
ctttacgaag ttcacctttg ggcacaacaa ttcgatcctc gaagaagaga gtgtccttgt 8040
catcaagtcg gtagcacttg tacttggact gactctttgc aataccaatc ttcacctttt 8100
tcaccatagc atcaaggagc tgggcttggc gaatctggtc ttccaaggta ggagaaactt 8160
gaagattggc aaggaaacct tgaggaacaa cttgcagatt aagtttgcgg aaggcttcac 8220
acagctcggg ttgataaggc ttgagaatca aactgttgca atatgctttt ctgctcaaag 8280
cgtcagcaat cacattagcc ttgcctggag tatactcgat actcggattg tactcttgaa 8340
tcatttcgac ccatcgagtc tgcctgaggt tgaggttagg ctgagtgaag atgtacttga 8400
gactcttgtg atcagtgaag atatccactt tccttcccaa gagaagatgt ctccaagtca 8460
acaatgcgtg cacaactgcc gccaactcga gatcatgagt ggggtagttc ttctcatttg 8520
gcttcaactg gcgagaggta taagcaacaa ctttcttctc ttgcatcagt actgcaccga 8580
gaccttggag agaggcatca caaaagacct cgtacggctt ggtttcatca ggcggagtta 8640
gaactggagc agtgatcaat ttctctttca aagcgttgaa agcaacatca cactctggag 8700
accaaacgta cttgacgtgc ttctgaagaa gatttgagag aggcttcgcg atcttggaga 8760
agttttcgac gaatcttcgg caataacttg caagaccgag gaagctacag agttgcttca 8820
cattctgagg aggttcccaa tttacaattg cagacacctt ctcaggatcc acggaaatgc 8880
ccttggcaga gataatatgc ccaagataaa gaacctcatc gagccaaaac tcacacttag 8940
agaacttggc atagaattga tgttccctta gcttgtcaag aaccaaacgc aagtgtttgg 9000
catgatcttc cttgttcttc gagaaaacca ggatgtcatc gagatagacc aaaacaaagt 9060
cgttggtgta ggggttgaag atgaagttca tcatgcgaga gaacgtcgga ggagcattag 9120
cgaggccaaa agacatcaca gtgtattcat atgacccata gcttgttctg aacgccgtct 9180
tcggaatatc ttgttcacga atgcgaattt gatgataacc catacggaga tcaagcttgg 9240
agaatacttg ggcacccttg agttgttcga atagctcatt aatgttggga agtggatact 9300
tgttcttgat ggtcttcttg ttcaatggac gataatcaac acaaagtcgg tccgttccat 9360
ccttcttctt cacaaaaaga acaccacaac cccacggaga agtactaggc cgaatgagac 9420
ccattctttc ttgcacatcg agctgtttct tcaactcctt caactcttcg gggccgagct 9480
tgtaaggccg tttgcacacc ggttccgtac cgggctcaag ttcaataacg aattcaacgg 9540
gccggtttgg aggcatgcct gggagttctt cagggaagac atcttgatac tcacaaacaa 9600
ccggaatctg cgaaatagca tccaattcac ccttttcatt gagagagaat aaccggatgg 9660
tattatcccg agcagcaaaa acaatgacat cttcaggaga atgagtcagt tgaatctgct 9720
tggctgcaca atcgaggtga gctttgtgtt tagagagcca gtccatccca agaatgagat 9780
tgatatcgga attgccaaga atcatcgggg aagccagaaa cttaaagtca cccagcgaga 9840
tagaaatatc tggaacttca tagttagcaa gcaagcattt gctcggagag acgactgcta 9900
atggctgatg cataacttga gaaaaccacc cgtgcttaga tgcaaaaggc ttcgagatga 9960
aacaatgtga tgcacctgta tcaaaaagta cttttgcagg aacatcgtta acaggaaggt 10020
tacccatgat cacatctgac gaatcctctg cttgagctgc attcaccata ttgaccttgg 10080
cgtgcttggg attgtgcttg accacagctg tacttgccga tctgacagga ggaggaggag 10140
gaagacgcct ctggttgaaa cacttgttgg cgtagtgacc cttctgttgg cacttgttgc 10200
acgtgacctc tgagagcgga cggtgatacg gagcactcgg tcttggagct tgagacgaag 10260
tcttgttctg aaagcctggg ttgggtgggt gggatgaacc actgccacct ttgctcttct 10320
gctgatacgg ctgacggaac ggaggaggag gaagccaata cttctgctgc ttggccactt 10380
gagtagagga cgaagacgaa gtgtccctga ctcgcttcct ggaagcatca cacctgagct 10440
gagcagcctc ttgcttcagt gccatgttgt agaactcatc gtatctcagc ggctcgaaga 10500
ggacaagagc gagctgaatc tcttcacgaa gaccacccct gaactgatag atcatgctct 10560
tctcatcagg aacatcctgc ttggcaaaac gggcgagctt ctggaacaac ttgttgtagt 10620
catagacaga caaagaacct tgcttcagat tgcggaattc ttcacgtttg ctttcagtca 10680
cactttgagg aatgtgatga gcgcggaaat cacgacggaa atcatcccaa gtgatcaaac 10740
gtccacctat ggaatcccgg tactgctgaa accactctgc ggcctgatcc ttgagctgga 10800
aagacgcgaa cttgacgaaa tcttcaggtc tgacgttgct acattcaaaa tgcttgccca 10860
gatcgactaa ccaatcatca gcatcggtag cctcaacgca gttactgaac gtctttggac 10920
catttgcgag gaactgattc agagttgcaa actggttctg gttgctgcct tgattcgctt 10980
gactgctctg attccgctct tggatgattt gcatgatcag ctgagtattt gcattagtag 11040
cagccatcac agcctgccat gcttccggag gaggcggagg tggcggagga tcttgattct 11100
ggttctgact ggtgctctta gcgggagcca tcctgaagag gttgaccacc attagcacac 11160
aaatagacaa atgtagaagg tgaatccaac ggaatgaaaa attgcaacat aagtcttcac 11220
atccgaacaa aatgaacgaa tgcacttctc ttgaagtggt cacatattcc taaattgcga 11280
agccacgaag aatcatgtag aagaggtaca tcaacgaggg atggaatcaa catatgacgt 11340
cggagaagaa cgaataccac aactcgaaat aaaaccaagg gttggcttgc ggaataagcc 11400
ggaacaaaca cataatagaa atttcgtccg aagtgttcgt ggtaggacct acacgggctc 11460
gaccgtacag caccatcatg tacaaggcag tgcacatgac atacgaagcg tccccgagtc 11520
agcatagcca aggactcttt aagacacaac gaggccactg taaaatcgac cgtgaatagg 11580
cgaaccacta gacgtcgaac cccaatttca tatcacacat ctgttggaaa gatatcctaa 11640
ggctacttga attcccacct atgaaatttc cgaaactttc cggttatgta atcaggtgtt 11700
gaggatacag gggaagcata atatctcacc caagctagca aatcctacat ccagctgcat 11760
ccatccttca acacataacc aagaaacctt cggaaaccat ctatctcaac cttcgaaaag 11820
catccgttat acaagttatg gcgatactct cgaactcccg ccccagtact gggtggcgtc 11880
gaggttatct caccaacgaa ctgcataaag gagattttcg atgtcgcgca catatctcaa 11940
gtatgccaga attgcaacga tagaattatg atgacaacac ctcagagctc aactccccgg 12000
gacacttcca ctaaacccct gacaggaggc accaagacag tattctcatc ataagccatc 12060
ggaacaaaac taagatactc gcgtgatcct aaaatttttt ttagtgaaat ttgagaagag 12120
aagagtcaaa actctacgtc aggatgcctt tccagagcga tgaggagact gggaagtaaa 12180
aagaattcct aagctctctg atatataatt cctaaggact caaaacattt ttctagacac 12240
cactcggctg ctaaaacgat caagcaatgg ggctcctaag gtcggggaag gctctgatac 12300
caacttgtaa caccctcgat gcgactatat ctcccacgtg tcgaggcacg acttagaggc 12360
ataatcgcat tgaaggcata agtcgcaagt gaggcaatct tcacacaacc catgtacata 12420
taaatataaa ggtggaataa taataataat tggcttacac tcgccacgtc acatcaaggt 12480
acaaaaatag cattcatcat acaaacactc atggcccgac tacggcgcca aaatagaaaa 12540
gactccccaa catgcgacta atccctgaat cgaaccccga gcaggcacca ctactgatca 12600
tcaggaaaag aaacatagta tcgctgagag tcctcgtcgt actcccactt cagctcgaac 12660
gcgccatccg gaggagtatc attggtccct gcatctggtg taatagtaat ctgtgagcca 12720
caaggactca gcaatctcgc accctcgcga tcaagactat ttaagcttaa taggtaagga 12780
agggggtaat atgagtggag ctgcagcaag cgactagcat atatggtggc taacttattc 12840
gcaaaagaga gcgagaagag aaggcaaggc acgagcggaa aactaagtgg agcagactac 12900
gcaaacataa ctccaacgtc gtgtccactt cccggactcc gtcgagaaga ggccatcacg 12960
gcaacacacg cagtagattc attttaatta attaagttca agttatctac aaccggacat 13020
taacaaattt ccatctgccc ataaccgcgg gcacggcttt cgaaagatca atccctgcag 13080
gggggttccc aacttagccc ataacaagct ctcacggtca acgaaggaat agacctccac 13140
ccaggaccca ccgatcagac tcggtatccc ggtacaacaa gatgattcga caggtcaaaa 13200
ctagtccagc aacaccgccc gaatgcgccg acaaatcccg ataggagctg cacatatctc 13260
tttttcaggg cacactcaga ttgtctaaac ttccgttagg ccagcccaga gttgcccctg 13320
gtggccaccg gcggctgacg ttccggacca acactcagag gagcactggc ccgtgggggg 13380
ctaaaataaa gatgaccctc gggctccgga aacccaaggg aaagaaaagg ttaggtggca 13440
aatggtaaaa ccaatgttgg gcattgctgg aaaagcttta atcaaggcga actatcaatg 13500
ggttcccatt atagcccaac cgcgtaaggg acgcaacaaa ctgggaacat aacaccgata 13560
tgacggaaac tagggcggca agagtggaac aaaacactag gcgagaggcc gagccttcca 13620
ccctttacca agtatataga taagtaaaga taacatagca atataatgat atcccaacaa 13680
gtaaataaat gatccaacaa ggaacggctc caatcttcac ctgcaactag caacgctata 13740
agaaggggct tgagcaaagc ggtaacatag ccaatcaacg gtttgctagg acaaaggtgg 13800
gttagaggct caatcatggc aatagggagg ctgacaagca agtggtaggc atcgtagcat 13860
tggcaaagca aaagagcgag caaactagca tagcaaagat agtagtgatt tcaagggtgt 13920
gatcatcttg cctgcacagt tgtcagaatt gactggatcc tcaaaagcaa actcaacggg 13980
ctcctcgtta gcgaactcgt ctcccggctc tacccaacaa cacaaacaag caacaaggat 14040
acaatcaacc acgtgcaagg gcaaccaata agatgcaaag atgatatgct atgcgggatg 14100
cgatgcggga tgcaagaatg caacatgtaa caggaaaatg catgaacctg gcctcaactt 14160
ggaaatccaa gtgtgccact ggatagagga gaagaaatcc cctgaaaacg atataaagat 14220
catggcaatc ggagctacgg ttaagaaatg gcaagcgttt tactaaacga accgatctgc 14280
gatttacagc aagtaggctg ctaaatgcaa caaaatgaac atgccacagc caccaacatg 14340
aaaacaaatt gcatggcagt gatgtactca tcaagcttaa caaatcatca tgactgaggc 14400
ataggcaaat tcatccatta agggcttcaa acaagcatgg caaaactgaa taagcaacac 14460
agatttcaga cttagcgaaa ttaacgctag tctgaaattt caaaccacga tgctctcttc 14520
ggagcagcaa aacaacatga aggaaaacct gaacatggca agtaagagca tggcatggag 14580
ctactcaaca agcttaacaa aacacccaaa atgacctggg gccaaaaggg ttcacaaaat 14640
actctaccga gcacacgaac attgatgaaa cacaatcagt tttcagactt agtgaaaact 14700
gggacatgct gaaataataa cgcacggagg catgtaaacg agctcgatgc tctcactacg 14760
gtgcaagtta tggcaaggca agcatatatc cattaagaag gcacaaaatg atagctagac 14820
atggcaagaa caatggcata gcatgcatgg atcactagca ataactccgg caaaatcgca 14880
aaagagttaa cgatctgccc aaaacagcaa ctatccaaaa gttgagctcg attgagtcaa 14940
cctagggggc tccaacataa caagtaaaga catggatgga ttaagcacaa caagcataac 15000
aaatcactct tactgatcat cctcaaaaga ggcacggatc actaggaaac aagccggaca 15060
tatagcatcg tgaactaaaa cggctcagac ttagtgaaaa tcactaagtc cctgaaatct 15120
gtaatctcag gtagccccct tcgcaagctt gcacaagaca acatacacat cctaaaaata 15180
catggaaggc acctctggaa agaagataac atgcttaata attcatccca aaggggcata 15240
ggcatatcat gcacggatta aatatggcaa aaatggcaaa agtgcatgtt aaactaacgg 15300
atctgcaatt taactcaaca tgcctccttc caacagcatt ttgggcagca agatgacctc 15360
aaatgcaaat gatgcaatgg aatgaaatga tgtactcgtc gagacgaaga atttgacata 15420
tcatacgccc aaatcggagc tacggatgca aagttacggg gccccgaaca gaggcatatg 15480
gactgaaaat tcctgggact tagagaaaaa tgcaacctcc cagatctagg gtttccagat 15540
ttgcacgcaa tccgaggcgg cggcgcgagc actgttcatc cgaagctccc cggaatcctc 15600
atcgtcgccg gagtggagag gagaggccgg gccgggcagc ggtggccggg cggcgagcgc 15660
tggcggggaa ggacgccccg gtggcgcgcc ttacggctcc ggccaccgga gccgaaggac 15720
ggggcggcgc ggaacttgag gctccggcca ccggagtcgc gggcggcggc agaggccggt 15780
caaggtccgc ggcggcgccg gcggctcgat ggcggcggga ggaaggagac ggcggcgcgg 15840
agcccaggcg cgggcgcggg gcggcttggg ccgcgggggc ctgccccggg ccgtgcgggc 15900
cggtgcggtg ggcgacggtg gcggcgaacg gcggcgtgac acgtggcggc ggggaggcag 15960
atcggacacg tccggcgccg gcggacgttg tccggcgcgc ggaaagagag gatttagggt 16020
ttcggggaag gagatccgag atttagagga gggggtctat atataggcat agagggagct 16080
aggagagtcc aaatgaggtg cggttttcgc ccacacgatc gtgatcaaac gaccgagacg 16140
atggagctga gtttggtggg ttttgggcca aattggaggg gtgttgggct gcacacacac 16200
gaggcctttt cggtccctcg gttaaccgtt ggagtatcaa acgaagtcca aatgacccga 16260
aacttgacag gcggtctacc ggtagtaaac caaggccgca tggcaagtct cggcccaatc 16320
cggaaatgtt taatccccac acacgaaaga aagctagaaa tggccaccgg aggagaacga 16380
agcgccggaa tgcaaaacgg acaacgggga aaatgctcgg atgcatgaga tgaacatgta 16440
tgcgaatgca atgcacgtga tgacatgata tgagatgcac gaaaacaaaa acaacacacg 16500
gagacaaaga cccgaacccg agaattaaat ataatttaat gccggaaacg gcatgagttg 16560
gagtataatt ggggaaggtt acatatgggg cgttacaggg agtctggcct cgatgggtgg 16620
agtcgcgacg ctgctctggc gtcaggagag cacgccgggc ccaaatctcg ttggcgcgcg 16680
cacgctcgga ctgtggcacc gccggcactg gaatgtggtg ggggtcgagg tgccagccgt 16740
gggggaagtt cacgtccggc cacggcagtg agacgcggta ctcccagtgg tactacgcgt 16800
ggtgcaccgg gatgtactgg tggtcccgtg gcgctgctgc cctcccatcc cgcagcgccg 16860
acaccaatgc gaaccccttc gctgccggcc tcaccggccc ttgcaacgag ccggcctcgt 16920
ggtcgtgctt gccttgccct tacctttgcc aaagaaaccc atgacggcta gggttttctg 16980
tcaccggcga gggagcagat gtggactaga gtgggaggag aagtggatga ggcccgcccc 17040
gccgcaccct tgcaattaaa aagggcgtga tccggacgtt tctagtcact gccatgcggg 17100
cccaaggtag gtgggcggca ttgactacgt ccgtggtggt agttggctgg cagacacgcc 17160
agcccgaggc gcacaccgag cggacgcgca acacgtccgt ctcgtgttcg tgtgaaaaca 17220
aatcaggcac aaatttgggc tggaaatgcg tctacatgga tgccgaacgg acgtgatttg 17280
ggtttgtgtc cgcacgttgg gccggcgttt ttgtgccttt cgaccctaat ggacacacac 17340
ggaccaaatc cgttggcgtg ttagagttgc ttgtacctag tactactatg tgtggagttg 17400
ttatctttga gtttctcccc catctgtagc tgtgtattat tgtgagatgc tctcaggatt 17460
tcacttattt tagataatga caactttctt gatgctagac aattacatca aggtgtatta 17520
tatctaggat gcacacagat gtgagattgt ttcaggattt taggttactt ggtaggtgtt 17580
actataaata tctaatgagc tctaatgtgt atgttcatga aactgtgttg caggtacttt 17640
atcataattg atgatatatg ggagttagga acatgggaaa ctttgaaatg tgcatttgtc 17700
aagaatacat tgggcagcag aataattatc acaacacgta tagttgatgt tgccaaatct 17760
tgttctcctt ctagtgaaga tcttgtctat gagatgaaac cactcagtga ggctgactcg 17820
aagaaattgt ttttcaagag gatatttggc tgtgaagaaa gctgtcctga tagcttgaaa 17880
gaagctgcca atgatatttt gaaaaaatgc cgtggcctac ctctggccat caatgccatt 17940
tctagcgtgt tggtcacaac aagggaaaca aaagaagagt gggatcgggt gcgacattca 18000
atacgctctt caaaggtcaa aagtgatata atagagacca tgaattacat attgtctctt 18060
agttattttg atcttcccca ccacctaaga agctgcttat tgtacctggc tctgtttcct 18120
gaagaccaat tgattggaag gaagcggtta gtacgcagat ggatttctga aggctttatc 18180
catggagaaa gtggacaaga tctcatggaa ttaggagagg aatattttca ccagcttgtg 18240
aacagaagtc taatacagcc cggcaacata gggtatgacg gcaaggcaat gtattgccga 18300
gtccatgaca ccatccttga tttcctaatt gacaaatcct ctgaagagaa catgtgtact 18360
gtcctaaaga aacaatgcaa gcctaatggc atagttcgtc ggctctctct aatggggaat 18420
gaggatgaag aaatcgtgga gcaattggat ctgtcacacg ctcggtcaat cagtgctttt 18480
ggggatatca agctattgcc ttcgcttggg aggtcaaaat gcttgcgcgt gctagacttg 18540
caaggctgcg atcagttgga aaatcatcat atcaaggata ttgaaagact ctatcagcta 18600
agatacttgg atatctcttc tacaggaatc acggagcttc ccaggcaaat tggggagttg 18660
ctgtatctag agacgctagt tgcctatgga ttacgtgagc ttcctgaaag cacaagtcgg 18720
ctacaacgac tggcacgttt gtttgtttac tctggttgta agcttccagg tgggttggga 18780
aacttgataa acttgcaaga gcttgactgc gttgatgcct tgcatttgaa gcatgtggaa 18840
gagctcggca aactaacaaa tctgagaaag ctgagtatta aattggacac tggtgggatt 18900
gaaggcaaca aattagaaga gtccaaggaa aagctggtgt cctctctctg taaactggac 18960
gaatgcggcc tccgttccct gagtattcat tattatctga gagaaaagga tggagaagag 19020
ccgttcctcc ctgctttggg ttgtatccag gaggtttgtg tatatggaca agatatcagc 19080
cggattagca gatggctcgc gtcacttccc aacctacaca ggttgttttt gaaccaaccg 19140
aagatagagc agcaggatat tgagatgatt ggattgatac ccaatctgat cgatttgtca 19200
ctgtctttat acaaaacaga cgacgctggg cggctcatca tcaggcgtga aggatttcaa 19260
cagttgcaga gttttcgggt ttaccatact cgtatgggag ttttaatgtt tgaaccgggg 19320
gctatgccga ggctcaagga gcttaaattc tatgacttca tcggaaagcc caaatctggt 19380
gcggtcgatt tcgactttgg catccaacac ctctccagcc ttgctcgcct cactgtcgac 19440
ttatcctgcg ttggctggac ggcggcggaa gtagaggctg cggaggatgc tttcaagagc 19500
atggccgagg caaaccccaa ccgtccaata ctggagatga caagactttt tccgcaggat 19560
atgttacgag acgagcaaat agatatggca ggctcaggta ctattaacaa gtttgtactt 19620
tttgtcttgg tgagcagtag atcaaaagat gtgcatatgc ttcattcttt ttccggtaat 19680
ctatttcagc tacaacgccg gccgtcaagt cctag 19715
<210> 2
<211> 2817
<212> DNA
<213> 一粒小麦(Triticum monococcum)
<400> 2
atggcggaag ttctggtgag cgcatccaca ggggcaatgg gttccctcct gcggaagttg 60
ggagctatgt tgacggatga atacaagctg ctcaagaacg tccgtgggga catcaagttc 120
ctcaaggacg agctggaggt catgtgcgcg ttcctcctca agatgtcaga tgtggaggag 180
cctgacgaac caacaaagct gcgtgtgacg gcggtgcggg agatgtccta taagatagag 240
gacaacatcg acaagttcat ggtcctcgtg gaacacgagt cctcctgctc cgaggcggct 300
catggcgtcg ccaagctcat ggacaagtgc aagaacttgc tgccggacat caagacccgc 360
cgcaggatcg ccaaggaggt caaagacatc aagaaagaaa ttaaggatgt cagcgacaga 420
ttttcaaggt acaagattga cgagtcctcc tcctctatgc cggcaaaagg caaggtcgac 480
cctcgacttc gtgcagtcta caaagacgcc gcagagctcg tcggaattga tggaccaaag 540
gatgaacttg tcaagtggct gaatgagaaa gagggccaat cactgaaatc tgtctctatt 600
gttggatatg gagggttggg caagaccacg ctcgccaacc agatccgggt caaccttgga 660
gcgaccttcg actgcggggc ttttgtctcg atttcacgca agcctgacat gaaggcgata 720
ttaagatcta tattatcgca aatcaccaag aaagacgagg cttgctctag attagatgat 780
atacaactca tcatcgacaa gattcgagaa ttcctccaag acacaaggta ctttatcata 840
attgatgata tatgggagtt aggaacatgg gaaactttga aatgtgcatt tgtcaagaat 900
acattgggca gcagaataat tatcacaaca cgtatagttg atgttgccaa atcttgttct 960
ccttctagtg aagatcttgt ctatgagatg aaaccactca gtgaggctga ctcgaagaaa 1020
ttgtttttca agaggatatt tggctgtgaa gaaagctgtc ctgatagctt gaaagaagct 1080
gccaatgata ttttgaaaaa atgccgtggc ctacctctgg ccatcaatgc catttctagc 1140
gtgttggtca caacaaggga aacaaaagaa gagtgggatc gggtgcgaca ttcaatacgc 1200
tcttcaaagg tcaaaagtga tataatagag accatgaatt acatattgtc tcttagttat 1260
tttgatcttc cccaccacct aagaagctgc ttattgtacc tggctctgtt tcctgaagac 1320
caattgattg gaaggaagcg gttagtacgc agatggattt ctgaaggctt tatccatgga 1380
gaaagtggac aagatctcat ggaattagga gaggaatatt ttcaccagct tgtgaacaga 1440
agtctaatac agcccggcaa catagggtat gacggcaagg caatgtattg ccgagtccat 1500
gacaccatcc ttgatttcct aattgacaaa tcctctgaag agaacatgtg tactgtccta 1560
aagaaacaat gcaagcctaa tggcatagtt cgtcggctct ctctaatggg gaatgaggat 1620
gaagaaatcg tggagcaatt ggatctgtca cacgctcggt caatcagtgc ttttggggat 1680
atcaagctat tgccttcgct tgggaggtca aaatgcttgc gcgtgctaga cttgcaaggc 1740
tgcgatcagt tggaaaatca tcatatcaag gatattgaaa gactctatca gctaagatac 1800
ttggatatct cttctacagg aatcacggag cttcccaggc aaattgggga gttgctgtat 1860
ctagagacgc tagttgccta tggattacgt gagcttcctg aaagcacaag tcggctacaa 1920
cgactggcac gtttgtttgt ttactctggt tgtaagcttc caggtgggtt gggaaacttg 1980
ataaacttgc aagagcttga ctgcgttgat gccttgcatt tgaagcatgt ggaagagctc 2040
ggcaaactaa caaatctgag aaagctgagt attaaattgg acactggtgg gattgaaggc 2100
aacaaattag aagagtccaa ggaaaagctg gtgtcctctc tctgtaaact ggacgaatgc 2160
ggcctccgtt ccctgagtat tcattattat ctgagagaaa aggatggaga agagccgttc 2220
ctccctgctt tgggttgtat ccaggaggtt tgtgtatatg gacaagatat cagccggatt 2280
agcagatggc tcgcgtcact tcccaaccta cacaggttgt ttttgaacca accgaagata 2340
gagcagcagg atattgagat gattggattg atacccaatc tgatcgattt gtcactgtct 2400
ttatacaaaa cagacgacgc tgggcggctc atcatcaggc gtgaaggatt tcaacagttg 2460
cagagttttc gggtttacca tactcgtatg ggagttttaa tgtttgaacc gggggctatg 2520
ccgaggctca aggagcttaa attctatgac ttcatcggaa agcccaaatc tggtgcggtc 2580
gatttcgact ttggcatcca acacctctcc agccttgctc gcctcactgt cgacttatcc 2640
tgcgttggct ggacggcggc ggaagtagag gctgcggagg atgctttcaa gagcatggcc 2700
gaggcaaacc ccaaccgtcc aatactggag atgacaagac tttttccgca ggatatgtta 2760
cgagacgagc aaatagatat ggcaggctca gctacaacgc cggccgtcaa gtcctag 2817
<210> 3
<211> 938
<212> PRT
<213> 一粒小麦(Triticum monococcum)
<400> 3
Met Ala Glu Val Leu Val Ser Ala Ser Thr Gly Ala Met Gly Ser Leu
1 5 10 15
Leu Arg Lys Leu Gly Ala Met Leu Thr Asp Glu Tyr Lys Leu Leu Lys
20 25 30
Asn Val Arg Gly Asp Ile Lys Phe Leu Lys Asp Glu Leu Glu Val Met
35 40 45
Cys Ala Phe Leu Leu Lys Met Ser Asp Val Glu Glu Pro Asp Glu Pro
50 55 60
Thr Lys Leu Arg Val Thr Ala Val Arg Glu Met Ser Tyr Lys Ile Glu
65 70 75 80
Asp Asn Ile Asp Lys Phe Met Val Leu Val Glu His Glu Ser Ser Cys
85 90 95
Ser Glu Ala Ala His Gly Val Ala Lys Leu Met Asp Lys Cys Lys Asn
100 105 110
Leu Leu Pro Asp Ile Lys Thr Arg Arg Arg Ile Ala Lys Glu Val Lys
115 120 125
Asp Ile Lys Lys Glu Ile Lys Asp Val Ser Asp Arg Phe Ser Arg Tyr
130 135 140
Lys Ile Asp Glu Ser Ser Ser Ser Met Pro Ala Lys Gly Lys Val Asp
145 150 155 160
Pro Arg Leu Arg Ala Val Tyr Lys Asp Ala Ala Glu Leu Val Gly Ile
165 170 175
Asp Gly Pro Lys Asp Glu Leu Val Lys Trp Leu Asn Glu Lys Glu Gly
180 185 190
Gln Ser Leu Lys Ser Val Ser Ile Val Gly Tyr Gly Gly Leu Gly Lys
195 200 205
Thr Thr Leu Ala Asn Gln Ile Arg Val Asn Leu Gly Ala Thr Phe Asp
210 215 220
Cys Gly Ala Phe Val Ser Ile Ser Arg Lys Pro Asp Met Lys Ala Ile
225 230 235 240
Leu Arg Ser Ile Leu Ser Gln Ile Thr Lys Lys Asp Glu Ala Cys Ser
245 250 255
Arg Leu Asp Asp Ile Gln Leu Ile Ile Asp Lys Ile Arg Glu Phe Leu
260 265 270
Gln Asp Thr Arg Tyr Phe Ile Ile Ile Asp Asp Ile Trp Glu Leu Gly
275 280 285
Thr Trp Glu Thr Leu Lys Cys Ala Phe Val Lys Asn Thr Leu Gly Ser
290 295 300
Arg Ile Ile Ile Thr Thr Arg Ile Val Asp Val Ala Lys Ser Cys Ser
305 310 315 320
Pro Ser Ser Glu Asp Leu Val Tyr Glu Met Lys Pro Leu Ser Glu Ala
325 330 335
Asp Ser Lys Lys Leu Phe Phe Lys Arg Ile Phe Gly Cys Glu Glu Ser
340 345 350
Cys Pro Asp Ser Leu Lys Glu Ala Ala Asn Asp Ile Leu Lys Lys Cys
355 360 365
Arg Gly Leu Pro Leu Ala Ile Asn Ala Ile Ser Ser Val Leu Val Thr
370 375 380
Thr Arg Glu Thr Lys Glu Glu Trp Asp Arg Val Arg His Ser Ile Arg
385 390 395 400
Ser Ser Lys Val Lys Ser Asp Ile Ile Glu Thr Met Asn Tyr Ile Leu
405 410 415
Ser Leu Ser Tyr Phe Asp Leu Pro His His Leu Arg Ser Cys Leu Leu
420 425 430
Tyr Leu Ala Leu Phe Pro Glu Asp Gln Leu Ile Gly Arg Lys Arg Leu
435 440 445
Val Arg Arg Trp Ile Ser Glu Gly Phe Ile His Gly Glu Ser Gly Gln
450 455 460
Asp Leu Met Glu Leu Gly Glu Glu Tyr Phe His Gln Leu Val Asn Arg
465 470 475 480
Ser Leu Ile Gln Pro Gly Asn Ile Gly Tyr Asp Gly Lys Ala Met Tyr
485 490 495
Cys Arg Val His Asp Thr Ile Leu Asp Phe Leu Ile Asp Lys Ser Ser
500 505 510
Glu Glu Asn Met Cys Thr Val Leu Lys Lys Gln Cys Lys Pro Asn Gly
515 520 525
Ile Val Arg Arg Leu Ser Leu Met Gly Asn Glu Asp Glu Glu Ile Val
530 535 540
Glu Gln Leu Asp Leu Ser His Ala Arg Ser Ile Ser Ala Phe Gly Asp
545 550 555 560
Ile Lys Leu Leu Pro Ser Leu Gly Arg Ser Lys Cys Leu Arg Val Leu
565 570 575
Asp Leu Gln Gly Cys Asp Gln Leu Glu Asn His His Ile Lys Asp Ile
580 585 590
Glu Arg Leu Tyr Gln Leu Arg Tyr Leu Asp Ile Ser Ser Thr Gly Ile
595 600 605
Thr Glu Leu Pro Arg Gln Ile Gly Glu Leu Leu Tyr Leu Glu Thr Leu
610 615 620
Val Ala Tyr Gly Leu Arg Glu Leu Pro Glu Ser Thr Ser Arg Leu Gln
625 630 635 640
Arg Leu Ala Arg Leu Phe Val Tyr Ser Gly Cys Lys Leu Pro Gly Gly
645 650 655
Leu Gly Asn Leu Ile Asn Leu Gln Glu Leu Asp Cys Val Asp Ala Leu
660 665 670
His Leu Lys His Val Glu Glu Leu Gly Lys Leu Thr Asn Leu Arg Lys
675 680 685
Leu Ser Ile Lys Leu Asp Thr Gly Gly Ile Glu Gly Asn Lys Leu Glu
690 695 700
Glu Ser Lys Glu Lys Leu Val Ser Ser Leu Cys Lys Leu Asp Glu Cys
705 710 715 720
Gly Leu Arg Ser Leu Ser Ile His Tyr Tyr Leu Arg Glu Lys Asp Gly
725 730 735
Glu Glu Pro Phe Leu Pro Ala Leu Gly Cys Ile Gln Glu Val Cys Val
740 745 750
Tyr Gly Gln Asp Ile Ser Arg Ile Ser Arg Trp Leu Ala Ser Leu Pro
755 760 765
Asn Leu His Arg Leu Phe Leu Asn Gln Pro Lys Ile Glu Gln Gln Asp
770 775 780
Ile Glu Met Ile Gly Leu Ile Pro Asn Leu Ile Asp Leu Ser Leu Ser
785 790 795 800
Leu Tyr Lys Thr Asp Asp Ala Gly Arg Leu Ile Ile Arg Arg Glu Gly
805 810 815
Phe Gln Gln Leu Gln Ser Phe Arg Val Tyr His Thr Arg Met Gly Val
820 825 830
Leu Met Phe Glu Pro Gly Ala Met Pro Arg Leu Lys Glu Leu Lys Phe
835 840 845
Tyr Asp Phe Ile Gly Lys Pro Lys Ser Gly Ala Val Asp Phe Asp Phe
850 855 860
Gly Ile Gln His Leu Ser Ser Leu Ala Arg Leu Thr Val Asp Leu Ser
865 870 875 880
Cys Val Gly Trp Thr Ala Ala Glu Val Glu Ala Ala Glu Asp Ala Phe
885 890 895
Lys Ser Met Ala Glu Ala Asn Pro Asn Arg Pro Ile Leu Glu Met Thr
900 905 910
Arg Leu Phe Pro Gln Asp Met Leu Arg Asp Glu Gln Ile Asp Met Ala
915 920 925
Gly Ser Ala Thr Thr Pro Ala Val Lys Ser
930 935
<210> 4
<211> 673
<212> DNA
<213> 一粒小麦(Triticum monococcum)
<400> 4
gcactgagac tcctcggtga tgtgatgggc cccctcctcc aatgccttga ggaacgccgc 60
ctcctcctca acctactatt tgaccaccgg tggcggcggt ggagggcagg cttcgagctg 120
gacggcatcg cggaggcgca catggtgctc gatggtgaac cagacttctc agttagggga 180
gtgtaacacc ctcgatgcga ctatatctcc cacgtgtcga ggcacgactt agaggcataa 240
tcgcattgaa ggcataagtc gcaagtgagg caatcttcac acaacccatg tacatataaa 300
tataaaggtg gaataataat aataattggc ttacactcgc cacgtcacat caaggtacaa 360
aaatagcatt catcatacaa acactcatgg cccgactacg gcgccaaaat agaaaagact 420
ccccaacatg cgactaatcc ctgaatcgaa ccccgagcag gcaccactac tgatcatcag 480
gaaaagaaac atagtatcgc tgagagtcct cgtcgtactc ccacttcagc tcgaacgcgc 540
catccggagg agtatcatcg gtccctgcat ctggtgtaat agtaatctgt gagccacaag 600
gactcagcaa tctcgcaccc tcgcgatcaa gactatttaa gcttaatagg taaggaaggg 660
ggtaatatga gtg 673
<210> 5
<211> 23
<212> DNA
<213> 人工合成(Artificial Synthesis)
<400> 5
gcactgagac tcctcggtga tgt 23
<210> 6
<211> 25
<212> DNA
<213> 人工合成(Artificial Synthesis)
<400> 6
cactcatatt acccccttcc ttacc 25
Claims (3)
1.一种检测小麦抗秆锈病基因SrTm5的引物对，其特征在于，所述引物对的核苷酸序列如下：
正向引物序列：5’-GCACTGAGACTCCTCGGTGATGT-3’；
反向引物序列：5’-CACTCATATTACCCCCTTCCTTACC-3’。
2.一种小麦抗秆锈病基因SrTm5的鉴定方法，其特征在于，所述方法包括以下步骤：
(a)提取待测样本DNA；
(b)用检测引物对进行PCR扩增；和
(c)对PCR产物进行电泳分析；
其中检测引物对的正向序列是：5’-GCACTGAGACTCCTCGGTGATGT-3’，反向序列是5’-CACTCATATTACCCCCTTCCTTACC-3；
其中若所述PCR产物可检测出一条673bp的特异条带，则表明待测样本含有所述小麦抗秆锈病基因SrTm5；若无PCR扩增产物，则判断所述待测样本不含有所述小麦抗秆锈病基因SrTm5。
3.一种抗秆锈病基因SrTm5的检测试剂盒，其特征在于，所述检测试剂盒中包含一对检测引物，所述检测引物的正向序列是：5’-GCACTGAGACTCCTCGGTGATGT-3’，反向序列是5’-CACTCATATTACCCCCTTCCTTACC-3’。
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| CN111073898A (zh) * | 2020-01-21 | 2020-04-28 | 福建农林大学 | 小麦条锈病抗性基因YrU1及其克隆与应用 |
| CN111719008A (zh) * | 2019-03-19 | 2020-09-29 | 中国科学院遗传与发育生物学研究所 | 一种与小麦白粉病抗病基因Pm5e共分离的SNP及其应用 |
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| CN111719008A (zh) * | 2019-03-19 | 2020-09-29 | 中国科学院遗传与发育生物学研究所 | 一种与小麦白粉病抗病基因Pm5e共分离的SNP及其应用 |
| CN111073898A (zh) * | 2020-01-21 | 2020-04-28 | 福建农林大学 | 小麦条锈病抗性基因YrU1及其克隆与应用 |
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| 小麦远缘杂交现状、抗病基因转移及利用研究进展;刘成等;《中国农业科学》;20200401(第07期);第1287-1308页 * |
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