CN108342401A - 一个kinase基因LeRLK1-V及其表达载体和应用 - Google Patents
一个kinase基因LeRLK1-V及其表达载体和应用 Download PDFInfo
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Abstract
本发明属于基因工程领域，公开了一个kinase基因LeRLK1‑V及其表达载体和应用。LeRLK1‑V的cDNA序列为SEQ ID NO.1及其编码的氨基酸序列为SEQ ID NO.2。该基因来自小麦‑簇毛麦易位系6VS/6AL南农9918的6VS染色体臂上。LeRLK1‑V在抗白粉病小麦品种南农9918中受白粉菌诱导表达增强。通过瞬间表达将该基因转化感病小麦品种扬麦158，结果表明LeRLK1‑V的过量表达可以降低扬麦158的吸器指数。因此，LeRLK1‑V可望用于基因工程育种，将其导入易感白粉病小麦品种中，有望提高小麦的白粉病抗性。
Description
技术领域
本发明属于基因工程领域，公开了一个kinase基因LeRLK1-V及其表达载体和应用。
背景技术
小麦(Triticum aestivum)的整个生育期受到多种病虫的危害，其中由小麦白粉病菌(Blurmeria graminis f.sp.tritici，Bgt)侵染引起的小麦白粉病是小麦最严重的真菌病害之一。由于小麦白粉病菌生理小种多、毒力变异快，一旦产生新的毒性小种或小种种群发生变化，会导致小麦白粉病的大爆发，给农业生产带来灾难性后果。广谱、持久抗病基因的发掘和利用对白粉病防治具有重要意义。
簇毛麦(Haynaldia villosa，2n＝2x＝14，VV)携带的抗白粉病基因Pm21对小麦白粉病具有广谱、高抗特性，该基因定位于6V染色体短臂上。南京农业大学细胞遗传研究所利用四倍体圆锥小麦与二倍体簇毛麦杂交，再与普通小麦回交多代后，选育出了普通小麦-簇毛麦6VS/6AL易位系，将含有Pm21基因的6VS导入普通小麦。普通小麦-簇毛麦6VS/6AL易位系自1994年进入国内育种利用以来，在白粉病常发区进行了多年种植，对白粉病表现出了广谱、高抗特性，以之为亲本已经选育出一批高抗白粉病的小麦新品种，南农9918即选育出的其中一个品种，Pm21在育种中得到广泛应用。在6VS上筛选和鉴定具有白粉病抗性功能的基因，对分析Pm21介导的广谱抗性机理和克隆其候选基因对于利用基因工程手段培育具有白粉病广谱抗性的小麦品种具有重要意义。
发明内容
本发明的目的是针对现有技术的上述缺陷，提供一个kinase基因LeRLK1-V及其表达载体和应用。
本发明的目的可通过如下技术方案实现：
LeRLK1-V基因来自普通小麦(Triticum asetivum L.)南农9918，其核苷酸序列为SEQ IDNO.1。
该LeRLK1-V基因的蛋白质为LeRLK1-V，其氨基酸序列为SEQ ID NO.2。
含有所述的LeRLK1-V的重组表达载体pBI220:LeRLK1-V。
所述的LeRLK1-V基因的重组表达载体优选以pBI220为出发载体，将LeRLK1-V基因***pBI220的BamHI和StuI酶切位点间所得。
所述的LeRLK1-V基因在构建抗白粉病小麦品种中的应用。
所述的LeRLK1-V基因的表达载体在构建抗白粉病小麦品种中的应用。
有益效果：
本发明从小麦-簇毛麦易位系6VS/6AL南农9918的6VS染色体臂上克隆获得小麦中克隆得到了一个kinase基因LeRLK1-V及其所编码的蛋白质LeRLK1-V，将其***表达载体pBI220，得到的该基因的过量表达载体导入感病小麦品种中，可以提高感白粉病小麦品种对白粉病的抗性。LeRLK1-V的超量表达载体用于基因工程育种，将其导入易感白粉病小麦品种中，能够获得具备白粉病抗性的小麦种质。
附图说明
图1 LeRLK1-V在抗白粉病南农9918中受白粉菌诱导表达
图2 pBI220:LeRLK1-V超量表达载体图谱
图3与白粉菌互作的表达GUS基因的表皮细胞
A图:白粉菌侵入GUS表达的表皮细胞后未形成吸器；B图:白粉菌侵入GUS表达的表皮细胞后形成吸器；其中，co:分生孢子；pp:侵入钉；ha:吸器；hy:菌丝
图4利用瞬间表达研究LeRLK1-V基因对吸器形成的影响和抗白粉病效应
图5 LeRLK1-V转化扬麦158的T0代阳性植株的PCR分子鉴定
泳道1为DNA ladder，泳道2为水对照，泳道3为未转化扬麦158对照，泳道4为包含目的基因的载体对照，泳道5-10为阳性转化植株。
图6 LeRLK1-V转化扬麦158的T0代阳性植株的白粉病抗性鉴定
上排是重复1的鉴定，下排是重复2的鉴定
具体实施方式
实施例1小麦-簇毛麦易位系6VS/6AL南农9918中一个具有lectin和LRR结构域的kinase基因LeRLK1-V的克隆
小麦-簇毛麦易位系6VS/6AL南农9918是南京农业大学细胞遗传所育成的含有广谱抗白粉病基因Pm21的小麦品种(公知公用，陈佩度，张守忠，王秀娥，王苏玲，周波，冯祎高，刘大钧.抗白粉病高产小麦新品种南农9918.南京农业大学学报,2002,25(4):1438-1444)。前期已经在南农9918的6VS上开发了多个6VS的转化序列，进一步利用小片段***易位系NAU418将Pm21限定在6EST258和CINAU15之间，并且在Pm21所在染色体区间还有6EST243和6EST238两个标记(公知公用，Radiation-induced translocations withreduced Haynaldia villosa chromatin at the Pm21locus for powdery mildewresistance in wheat.Molecular breeding,2013,31:477–484)。
将6EST258、CINAU15、6EST243、6EST238序列与水稻(http://rice.plantbiology.msu.edu)、大麦(http://webblast.ipk-gatersleben.de/barley/viroblast.php)、短柄草(http://plants.ensembl.org/Brachypodium_distachyon/Info/Index)的序列数据库进行Blastn比对分析，分别提取出水稻、大麦、短柄草中位于6EST258和CINAU15所在染色体区间的序列。根据其中一个大麦的序列morex_contig_368490(http://webblast.ipk-gatersleben.de/barley/blastresult.php？jobid＝148384578557&opt＝none)设计引物，在南农9918受白粉菌诱导的cDNA样品中进行扩增，获得LeRLK1-V全长序列。cDNA样品准备及扩增流程如下：
具体流程如下：(1)将抗白粉病小麦南农9918的种子播于培养皿中发芽，露白后移栽到盆钵，一叶期把从感白粉病小麦材料上搜集的白粉菌孢子接种到苗上进行诱导，并于接种不同时间段取样(0h,1h,6h,12h,24h)，分别提取RNA(用Invitrogen公司的Trizol试剂提取)，并且反转录成cDNA样品(用Takara公司的AMV反转录酶反转)。(2)以南农9918经白粉菌诱导24小时的cDNA为模板，以LeRLK1-V基因引物P1(AGGAAGCATGGATCTATGCGT,SEQ IDNO.3)和P2(GCAAGGAACAGACGGAGGAT,SEQ ID NO.4)为引物进行RT-PCR，获得LeRLK1-V基因的cDNA全长片段。对该基因片段进行测序和比对，发现该基因为一个具有lectin和LRR结构域的kinase基因，命名为LeRLK1-V。对LeRLK1-V进行生物信息学分析，发现ORF(开放阅读框)2691bp，其核苷酸序列如SEQ ID NO.1所示，编码896个氨基酸，氨基酸序列如SEQ IDNO.2所示。
实施例2 LeRLK1-V在小麦-簇毛麦易位系6VS/6AL南农9918中受白粉菌诱导表达
以小麦-簇毛麦易位系6VS/6AL南农9918受白粉菌诱导不同时间段的cDNA样品为模板，以根据LeRLK1-V设计的引物P3(TCCGCAGTTATCACATCAGC,SEQ ID NO.5)和P4(CATTCTTGCATCCACAATGC，SEQ ID NO.6)为引物进行Q-PCR分析。PCR程序为：PCR反应在实时荧光定量PCR仪(MyIQ，Bio-Rad公司，美国)上扩增并检测荧光。20uL PCR反应体系中含2×SYBR Green PCR Master Mix 10uL，0.5μM引物P3和P4，反转录cDNA模板2uL。扩增参数为：95℃10分钟，然后95℃15秒、60℃30秒，72℃1分钟，共40个循环。反应结束后，进行熔解曲线的测定。检测基因表达水平用MyiQ***软件进行分析。结果表明：在南农9918中，LeRLK1-V在小麦-簇毛麦易位系6VS/6AL南农9918受白粉菌诱导上调表达(图1)。
实施例3 LeRLK1-V基因瞬间表达载体的构建
以经白粉菌诱导的南农9918中克隆的LeRLK1-V基因cDNA为模板，使用引物对NLR1-V-BamHI-F(GGAGAGAACACGGGGGATCCATGGATCTATGCGTAGTGATTCTCC，SEQ IDNO.7)和NLR1-V-StuI-R(AACGTCGTATGGGTAAGGCCTCTATCTCATTGCAGGACCAGCAG，SEQ IDNO.8)进行PCR扩增，回收扩增片段。用BamHI和StuI对扩增产物进行双酶切，将酶切产物***到BamHI和StuI双酶切后的载体pBI220(Jefferson RA,Kavanagh TA,Bevan MW.GUSfusions:beta-glucuronidase as a sensitive and versatile gene fusion marker in higherplants.EMBO J.1987,6:3901-3907.)，将LeRLK1-V置于35S启动子后面的多克隆位点处。由此将目标基因LeRLK1-V克隆到强启动子35S的下游，获得表达载体pBI220:LeRLK1-V(图1)。经测序验证，表明载体构建成功。
实施例4利用瞬间表达方法将LeRLK1-V基因转入小麦叶片
瞬间表达方法是一种可靠且快速鉴定基因功能的方法(Schweizer,Pokorny etal.A Transient Assay System for the Functional Assessment of Defense-RelatedGenes in Wheat Molecular Plant-Microbe Interactions.1999,12:647-654.)。本研究利用瞬间表达方法，将质粒DNA包裹到金属微粒外层，借助基因枪将金属微粒和基因轰击到小麦叶片的表皮细胞，然后统计轰击TaNACs细胞的白粉菌吸器指数与未轰击LeRLK1-V细胞的白粉菌吸器指数，明确目标基因是否具有白粉病抗病功能。
载体DNA与金属微粒包裹的程序如下：
制备钨粉：称取30mg的钨粉于1.5ml eppendorf管中，加入1ml 70％酒精，涡旋3-5min后静置15min，使钨粉完全沉淀。12000rpm离心1min后弃上清。加入1ml ddH2O水，涡旋混匀后，离心弃上清(重复三次)。最后加入500μl 50％甘油涡旋混匀，以备用。
包裹子弹：吸取5μl涡旋均匀的钨粉于1.5ml的eppendorf管中，加入5μl质粒DNA(总量应为1μg)。边涡旋边向eppendorf管内滴加50μl 2.5M CaCl2，然后加入20μl 0.1M亚精氨(现配先用)，涡旋3min。静置1min后离心2s，弃上清。加入140μl 70％酒精，充分涡旋，离心2s，弃上清。然后加入140μl 100％酒精，充分涡旋，离心2s，弃上清。最后加入15μl100％酒精，充分涡旋，以备使用。
实施GUS基因单转化时，将含有GUS基因的表达载体pAHC25(Christensen A H,Quail P H.Ubiquitin promoter-based vectors for high-level expression ofselectable and/or screenable marker genes in monocotyledonousplants.Transgenic Research,1996,5:213-218.)与钨粉包裹；当实施NLR1-V与GUS基因共转化时，将含有LeRLK1-V基因的表达载体pBI220:LeRLK1-V与含有GUS基因的表达载体pAHC25按摩尔浓度1:1的比例混合，包裹钨粉。当GUS基因与LeRLK1-V基因进行共转化时，Marker基因GUS转入的细胞也是LeRLK1-V转入的细胞。因GUS基因表达的细胞经染色整个细胞呈现蓝色，所以本研究以蓝色细胞作为LeRLK1-V的表达细胞。
基因枪轰击程序如下：剪下长约6cm的小麦幼苗叶片端部，平行贴在载玻片上，每张玻片贴6片叶片左右。基因枪使用PDS1000/He***，采用1350psi的可裂膜片，真空度为28inHg。轰击后将叶片置于垫有润湿滤纸的瓷盘内，罩以打有小孔的保鲜膜，保湿并透气，18-20℃恢复培养4h后，高密度接种白粉菌分生孢子。接种48h后用GUS染液(配方为：0.1mol/L Na2HPO4/NaH2PO4缓冲液(pH7.0)，含10mmol/L EDTA，5mmol/L铁***和亚铁***，0.1mg/ml X-Gluc，0.1％Triton X-100，20％甲醇)真空渗透10min,37℃染色12h，然后用70％酒精脱色2天直至叶片变成白色为止，最后利用浓度为0.6％的考马斯亮蓝对白粉菌孢子染色。
实施例5在单细胞水平对LeRLK1-V抗病功能的分析
白粉菌侵入小麦叶片表皮细胞后，在表皮细胞中产生的指状物称为吸器。吸器不能正常产生是叶片细胞对白粉菌具有抗性的重要指标。在GUS表达的细胞中，吸器会被GUS染色液染成蓝色，在显微镜下容易辨认(图3)。在GUS基因转化细胞后，通过统计与白粉菌互作的GUS表达细胞中，吸器形成的细胞所占的比例(％)，即为“吸器指数”(Schweizer,Pokorny et al.A Transient Assay System for the Functional Assessment ofDefense-Related Genes in Wheat Molecular Plant-Microbe Interactions.1999,12:647-654.)。吸器指数越小，表明抗病性越强。本研究利用“吸器指数”作为抗病强弱的衡量指标。
当单转化GUS基因时，感病小麦扬麦158的平均吸器指数为55.75％(共统计了910个互作细胞，重复1吸器指数为50.00％，重复2吸器指数为47.97％，重复3吸器指数为69.29％)，当GUS基因与LeRLK1-V共转化感病小麦扬麦158后，统计了GUS基因表达(即LeRLK1-V表达)且有白粉菌互作的细胞的吸器指数，结果表明当LeRLK1-V转入后，扬麦158的平均吸器指数为37.15％(共统计了758个细胞，重复1吸器指数为29.22％，重复2吸器指数为35.39％，重复3吸器指数为46.84％)(图4)。该结果说明，LeRLK1-V能显著降低吸器指数，对白粉菌具有抗病作用。
实施例6 LeRLK1-V转化普通小麦扬麦158及LeRLK1-V的白粉病抗性鉴定
挑取预培养7天约2000块扬麦158幼胚愈伤组织，将携有目的基因LeRLK1-V的过量表达载体pBI220:LeRLK1-V通过基因枪轰击法转化到扬麦158，轰击前在高渗培养基(MS+ABA0.5mg/L+水解酪蛋白500mg/L+2，4-D2mg/L+葡萄糖30g/L+0.4mol/L甘露醇，pH5.8)上预处理4–5小时，轰击后在高渗培养基上继续培养16小时。之后将愈伤组织转移至恢复培养基(1/2MS+水解酪蛋白500mg/L+2,4-D2mg/L+蔗糖30g/L，pH5.8)上暗培养2周，再将其转移至含有除草剂的筛选培养基上(1/2MS+ABA0.5mg/L+水解酪蛋白500mg/L+2,4-D1mg/L+蔗糖30g/L+4mg/L Bialaphos，pH5.8)，筛选培养2周。然后将具有除草剂抗性的愈伤组织转移到分化培养基中(1/2MS+L-谷氨酞胺l mmol/L+水解酪蛋白200mg/L+KT 1mg/L+IAA 0.5mg/L+蔗糖30g/L+琼脂0.8％，pH5.8)进行分化，待分化芽长至2–4cm时将其转移至生根培养基(1/2MS+KT 1mg/L+蔗糖30g/L+琼脂0.8％，pH5.8)中。至再生苗长约8cm、根系较健壮时，即可开管炼苗1–2天，最后洗去根系携带的培养基残渣便可移栽入盆钵，获得再生植株共75棵。
提取所有再生植株基因组DNA，对转化植株利用基因内部引物P7(TCCAACCACGTCTTCAAAGC，SEQ ID NO.9)和载体上终止子引物P8(CTCCATGCTGAAAGAACCGG，SEQ ID NO.10)进行PCR扩增，以鉴定阳性植株。PCR程序：10-50ng/ul基因组模板，10μM的P7和P8各0.5μl；2.5μl10×buffer；2.5μl 2.5mM的dNTP；1.5μl 25mM的Mg2+；0.25μl(5U/μl)Taq polymerase(TaKaRa)，加水至25μl。PCR反应条件为：94℃预变性3分钟；94℃45秒，55℃45秒，72℃1分钟，33个循环；72℃延伸10分钟。PCR产物经8％的聚丙烯凝胶电泳检测。其中6株可以扩增出与质粒对照一致的约600bp的目的条带，鉴定为阳性植株，株系编号依次为：HP5、HP13、HP48、HP49、HP58、HP59(图5)。
对所有鉴定的阳性植株和对照扬麦158进行白粉病抗性鉴定，鉴定方法为：在植株生长到2叶1心期时，剪取第二叶平铺在琼脂平板上(1％琼脂粉，20mg/L的6-BA)；将感病品种苏麦三号上繁殖的白粉菌孢子(南京地区混合白粉菌孢子)抖落在琼脂平板上的待鉴定叶片上；将接种了白粉菌的平板放置在智能培养箱中(16小时光照、25度/8小时黑暗、22度，80％湿度)，7天后观察拍照。鉴定结果表明：在两个重复鉴定实验中，感病对照扬麦158上白粉菌正常发育，而转基因T0代阳性植株HP5、HP13、HP48、HP49、HP58、HP59的叶片上孢子密度明显降低，与未转化扬麦158相比，转基因植株的白粉病抗性水平显著提高(图6)。
<110> 南京农业大学
<120> 一个kinase基因LeRLK1-V及其表达载体和应用
<160> 10
<210> 1
<211> 2691
<212> DNA
<213> 普通小麦（Triticum asetivum L.）南农9918
<220>
<223> LeRLK1-V基因
<400> 1
atggatctat gcgtagtgat tctcctattc tgcgtcttcc agtttatcca tggccagcct 60
gatgacctag gtttcataag catcgattgt ggcatcccgg tgaactcctc ctaccaagat 120
cccagctcaa aaataatata tgtctcggac tatggatatg tcacctcagg agaaaaccgc 180
aatatcgcct cagtttacat caagccttca ctggcaaagc gcaacctcaa cgtccggttc 240
tttcagcatg gagcacgcaa ttgctacgcc ctgaggtcct tggtggcagg aaacaagtac 300
ttcgtccgtg cgctttttta ctacggaaac tacgatggcc ttaacaaact tcctgtcttt 360
gatctgtaca tgggggcaaa ctactggcat gaagttaata tcagtgctgc aggagcagtc 420
aaatggatgg acatcatagt tgtcgctcct gctgactacc tgcatgtttg tctggtgaac 480
aaagggatgg gaactccatt tatttctggg ctggatctga ggccactgag gattactctt 540
tacccagagg taaatgcaag tcaatctctg gtgctgatta gctccaatcg gttcaatttg 600
gggcccacag gcaatcgcat aatcaggtac cccttggatc cccatgatcg cctatggtcg 660
acttacgaca caatcccaaa ctggaatgaa atatctgcaa catctgtcgt ccagaattac 720
gtcactgatg tgtacgacgt gccatcagct gttatgcaaa atgcagcaac tgtcaacggc 780
tcaaggattg atttctcatg gggtccatcg gatccatcgg agaacatcag ctccagttac 840
ttctttgttt tctacttctc tgagttgcag aatgtacaga gcaatgctgt gcggcagttt 900
gacatcattg tcaataacaa gacatggaac acacaacctt acagccctcg attcctattt 960
gccgattctt tctcaggcac tgtacaaggg ttggcgagtt atagtgtctc acttgttgct 1020
acagaaaatg caactcttcc acctatcctc aatgccatgg agatgtacct ggtggaaccg 1080
ataactgagg ccactactga tactggagat gctagagcca tgatgtcaat ccaagagaat 1140
tttggcgtgg agaaaaactg gatgggtgat ccatgtgctc caaaagcttt tgcgtggata 1200
ggattaacct gctcctatcc tccagcttat gcctctagaa taacagcatt aaatgtgtct 1260
tcctttgggt tggctggtac catctctact gattttggag atctgaacgc acttcagtac 1320
ctggatctgt caaacaacaa cttgtctggc accattccaa attttcttgg gcagcttcca 1380
tttctaatat ttctggatct gtccagcaat gatctacatg gaacaatccc tgacaatctt 1440
cttaaaaaat cccaaaatgg gactctgtcg ttaagggttg gtaataatgc aaatatatgt 1500
ggcaatggta ctgcctgtgg atcaagtcga aagaaaatta atggggcact tcttactgca 1560
atagttattc cgatcattgc tgtcattgca atatttcttg tcttgtttct tctgctgcgc 1620
caaaagctca agggaaaaga taagagaaag gctactggtc ctgaagatga atcagcgtta 1680
cttgagaacc gagaattttc ttacagagaa ctgaagtata ttacgagcaa ctttaggcaa 1740
gggattggca agggagggtt tggagctgtc ttccttggct acctgaagaa cggaaactca 1800
gttgctgtga aagtgcgttc tgattcatct tcacaagggg gtaaagagtt tctggccgag 1860
gctcaacact tgacaaggat tcatcacaag aacttggttt ccttgattgg ctactgcaag 1920
gacaaaaatc atctagccct tgtttacgag tacatgcccg aagggaacct gcaggatcat 1980
ctgagaggtt ctactagtaa accactcact tgggagcagc gtcttcacat cgctcttgat 2040
gctgcacaag gtctggagta tctgcacatc gcgtgtaagc cagcattgat ccacagagat 2100
gtgaagagta gcaacatcct gctgaccaca gatcttgggg ctaagattgc tgattttggc 2160
ctgaccaagg tttttggcga ctcgaaaaca catataacca ctgagccagc tggtactatg 2220
ggctacttag atcccgagta cttccgcagt tatcacatca gtgagaagag cgacgtgtac 2280
agcttcggcg tcgtactcct ggagctcatc acaggccgtc ctcctgtcat ccctatcagc 2340
gacagcgtaa acgtccacgt cggcgagtgg gtacagcaga gcctcgacca cggcaccatg 2400
cagagcattg tggatgcaag aatggtaggg gactatgaca tcaactctgt ctggaaagct 2460
gctgacctgg cactgcattg caagcgagat gtctcgaggg agcgtccggc gatggcggag 2520
gtggtggcgc agcttaagga gtgcttggag ctcgagaacc gtctcgacag gaagcaaaga 2580
agcttgggtt cgaacattcc tagggaggga agtgcacttg aggcagaaga agaagaagaa 2640
gaacaaggtg gggatataca agctgttgct gctggtcctg caatgagata g 2691
<210> 2
<211> 896
<212> PRT
<213> 普通小麦（Triticum asetivum L.）南农9918
<220>
<223> 蛋白LeRLK1-V
<400> 2
Met Asp Leu Cys Val Val Ile Leu Leu Phe Cys Val Phe Gln Phe Ile
1 5 10 15
His Gly Gln Pro Asp Asp Leu Gly Phe Ile Ser Ile Asp Cys Gly Ile
20 25 30
Pro Val Asn Ser Ser Tyr Gln Asp Pro Ser Ser Lys Ile Ile Tyr Val
35 40 45
Ser Asp Tyr Gly Tyr Val Thr Ser Gly Glu Asn Arg Asn Ile Ala Ser
50 55 60
Val Tyr Ile Lys Pro Ser Leu Ala Lys Arg Asn Leu Asn Val Arg Phe
65 70 75 80
Phe Gln His Gly Ala Arg Asn Cys Tyr Ala Leu Arg Ser Leu Val Ala
85 90 95
Gly Asn Lys Tyr Phe Val Arg Ala Leu Phe Tyr Tyr Gly Asn Tyr Asp
100 105 110
Gly Leu Asn Lys Leu Pro Val Phe Asp Leu Tyr Met Gly Ala Asn Tyr
115 120 125
Trp His Glu Val Asn Ile Ser Ala Ala Gly Ala Val Lys Trp Met Asp
130 135 140
Ile Ile Val Val Ala Pro Ala Asp Tyr Leu His Val Cys Leu Val Asn
145 150 155 160
Lys Gly Met Gly Thr Pro Phe Ile Ser Gly Leu Asp Leu Arg Pro Leu
165 170 175
Arg Ile Thr Leu Tyr Pro Glu Val Asn Ala Ser Gln Ser Leu Val Leu
180 185 190
Ile Ser Ser Asn Arg Phe Asn Leu Gly Pro Thr Gly Asn Arg Ile Ile
195 200 205
Arg Tyr Pro Leu Asp Pro His Asp Arg Leu Trp Ser Thr Tyr Asp Thr
210 215 220
Ile Pro Asn Trp Asn Glu Ile Ser Ala Thr Ser Val Val Gln Asn Tyr
225 230 235 240
Val Thr Asp Val Tyr Asp Val Pro Ser Ala Val Met Gln Asn Ala Ala
245 250 255
Thr Val Asn Gly Ser Arg Ile Asp Phe Ser Trp Gly Pro Ser Asp Pro
260 265 270
Ser Glu Asn Ile Ser Ser Ser Tyr Phe Phe Val Phe Tyr Phe Ser Glu
275 280 285
Leu Gln Asn Val Gln Ser Asn Ala Val Arg Gln Phe Asp Ile Ile Val
290 295 300
Asn Asn Lys Thr Trp Asn Thr Gln Pro Tyr Ser Pro Arg Phe Leu Phe
305 310 315 320
Ala Asp Ser Phe Ser Gly Thr Val Gln Gly Leu Ala Ser Tyr Ser Val
325 330 335
Ser Leu Val Ala Thr Glu Asn Ala Thr Leu Pro Pro Ile Leu Asn Ala
340 345 350
Met Glu Met Tyr Leu Val Glu Pro Ile Thr Glu Ala Thr Thr Asp Thr
355 360 365
Gly Asp Ala Arg Ala Met Met Ser Ile Gln Glu Asn Phe Gly Val Glu
370 375 380
Lys Asn Trp Met Gly Asp Pro Cys Ala Pro Lys Ala Phe Ala Trp Ile
385 390 395 400
Gly Leu Thr Cys Ser Tyr Pro Pro Ala Tyr Ala Ser Arg Ile Thr Ala
405 410 415
Leu Asn Val Ser Ser Phe Gly Leu Ala Gly Thr Ile Ser Thr Asp Phe
420 425 430
Gly Asp Leu Asn Ala Leu Gln Tyr Leu Asp Leu Ser Asn Asn Asn Leu
435 440 445
Ser Gly Thr Ile Pro Asn Phe Leu Gly Gln Leu Pro Phe Leu Ile Phe
450 455 460
Leu Asp Leu Ser Ser Asn Asp Leu His Gly Thr Ile Pro Asp Asn Leu
465 470 475 480
Leu Lys Lys Ser Gln Asn Gly Thr Leu Ser Leu Arg Val Gly Asn Asn
485 490 495
Ala Asn Ile Cys Gly Asn Gly Thr Ala Cys Gly Ser Ser Arg Lys Lys
500 505 510
Ile Asn Gly Ala Leu Leu Thr Ala Ile Val Ile Pro Ile Ile Ala Val
515 520 525
Ile Ala Ile Phe Leu Val Leu Phe Leu Leu Leu Arg Gln Lys Leu Lys
530 535 540
Gly Lys Asp Lys Arg Lys Ala Thr Gly Pro Glu Asp Glu Ser Ala Leu
545 550 555 560
Leu Glu Asn Arg Glu Phe Ser Tyr Arg Glu Leu Lys Tyr Ile Thr Ser
565 570 575
Asn Phe Arg Gln Gly Ile Gly Lys Gly Gly Phe Gly Ala Val Phe Leu
580 585 590
Gly Tyr Leu Lys Asn Gly Asn Ser Val Ala Val Lys Val Arg Ser Asp
595 600 605
Ser Ser Ser Gln Gly Gly Lys Glu Phe Leu Ala Glu Ala Gln His Leu
610 615 620
Thr Arg Ile His His Lys Asn Leu Val Ser Leu Ile Gly Tyr Cys Lys
625 630 635 640
Asp Lys Asn His Leu Ala Leu Val Tyr Glu Tyr Met Pro Glu Gly Asn
645 650 655
Leu Gln Asp His Leu Arg Gly Ser Thr Ser Lys Pro Leu Thr Trp Glu
660 665 670
Gln Arg Leu His Ile Ala Leu Asp Ala Ala Gln Gly Leu Glu Tyr Leu
675 680 685
His Ile Ala Cys Lys Pro Ala Leu Ile His Arg Asp Val Lys Ser Ser
690 695 700
Asn Ile Leu Leu Thr Thr Asp Leu Gly Ala Lys Ile Ala Asp Phe Gly
705 710 715 720
Leu Thr Lys Val Phe Gly Asp Ser Lys Thr His Ile Thr Thr Glu Pro
725 730 735
Ala Gly Thr Met Gly Tyr Leu Asp Pro Glu Tyr Phe Arg Ser Tyr His
740 745 750
Ile Ser Glu Lys Ser Asp Val Tyr Ser Phe Gly Val Val Leu Leu Glu
755 760 765
Leu Ile Thr Gly Arg Pro Pro Val Ile Pro Ile Ser Asp Ser Val Asn
770 775 780
Val His Val Gly Glu Trp Val Gln Gln Ser Leu Asp His Gly Thr Met
785 790 795 800
Gln Ser Ile Val Asp Ala Arg Met Val Gly Asp Tyr Asp Ile Asn Ser
805 810 815
Val Trp Lys Ala Ala Asp Leu Ala Leu His Cys Lys Arg Asp Val Ser
820 825 830
Arg Glu Arg Pro Ala Met Ala Glu Val Val Ala Gln Leu Lys Glu Cys
835 840 845
Leu Glu Leu Glu Asn Arg Leu Asp Arg Lys Gln Arg Ser Leu Gly Ser
850 855 860
Asn Ile Pro Arg Glu Gly Ser Ala Leu Glu Ala Glu Glu Glu Glu Glu
865 870 875 880
Glu Gln Gly Gly Asp Ile Gln Ala Val Ala Ala Gly Pro Ala Met Arg
885 890 895
<210> 3
<211> 21
<212> DNA
<213> 人工序列
<220>
<223> 引物P1
<400> 3
aggaagcatg gatctatgcg t 21
<210> 4
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物P2
<400> 4
gcaaggaaca gacggaggat 20
<210> 5
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物P3
<400> 5
tccgcagtta tcacatcagc 20
<210> 6
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物P4
<400> 6
cattcttgca tccacaatgc 20
<210> 7
<211> 45
<212> DNA
<213> 人工序列
<220>
<223> 引物NLR1-V-BamHI-F
<400> 7
ggagagaaca cgggggatcc atggatctat gcgtagtgat tctcc 45
<210> 8
<211> 44
<212> DNA
<213> 人工序列
<220>
<223> 引物NLR1-V-StuI-R
<400> 8
aacgtcgtat gggtaaggcc tctatctcat tgcaggacca gcag 44
<210> 9
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物P7
<400> 9
tccaaccacg tcttcaaagc 20
<210> 10
<211> 20
<212> DNA
<213> 人工序列
<220>
<223> 引物P8
<400> 10
ctccatgctg aaagaaccgg 20
Claims (6)
1.一个kinase基因LeRLK1-V，来自小麦-簇毛麦易位系6VS/6AL南农9918的6VS染色体臂上，其ORF序列如SEQ ID NO.1所示。
2.权利要求1所述的基因LeRLK1-V编码的蛋白质，其氨基酸序列为SEQ ID NO.2。
3.LeRLK1-V的重组表达载体pBI220:LeRLK1-V。
4.根据权利要求3所述的重组表达载体，其特征在于所述的LeRLK1-V的表达载体pBI220:LeRLK1-V是以pBI220为出发载体，将LeRLK1-V基因***pBI220的BamHI和StuI酶切位点间所得。
5.权利要求1所述的LeRLK1-V在构建抗白粉病小麦品种中的应用。
6.权利要求3或4所述的LeRLK1-V的表达载体pBI220:LeRLK1-V在构建抗白粉病小麦品种中的应用。
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| CN107699630A (zh) * | 2017-10-27 | 2018-02-16 | 南京农业大学 | 与小麦抗病基因Pm21连锁的分子标记及其在育种上的应用 |
| CN109535236A (zh) * | 2018-11-16 | 2019-03-29 | 南京农业大学 | 一个血红素结合蛋白基因TaHBP1及其重组干扰载体和应用 |
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| CN102533812A (zh) * | 2012-01-16 | 2012-07-04 | 南京农业大学 | 一个类受体蛋白激酶基因及其表达载体和应用 |
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| JEYARAMAN RAJARAMAN ET AL.: "An LRR/Malectin Receptor-Like Kinase Mediates Resistance to Non-adapted and Adapted Powdery Mildew Fungi in Barley and Wheat", 《FRONTIERS IN PLANT SCIENCE》 * |
| TINGTING CHEN ET AL.: "Two members of TaRLK family confer powdery mildew resistance in common wheat", 《BMC PLANT BIOLOGY》 * |
Cited By (4)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN107699630A (zh) * | 2017-10-27 | 2018-02-16 | 南京农业大学 | 与小麦抗病基因Pm21连锁的分子标记及其在育种上的应用 |
| CN107699630B (zh) * | 2017-10-27 | 2021-06-01 | 南京农业大学 | 与小麦抗病基因Pm21连锁的分子标记及其在育种上的应用 |
| CN109535236A (zh) * | 2018-11-16 | 2019-03-29 | 南京农业大学 | 一个血红素结合蛋白基因TaHBP1及其重组干扰载体和应用 |
| CN109535236B (zh) * | 2018-11-16 | 2022-04-01 | 南京农业大学 | 一个血红素结合蛋白基因TaHBP1及其重组干扰载体和应用 |
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