CN112126633B - 番茄周期素依赖性激酶SlCDK8基因及应用 - Google Patents

番茄周期素依赖性激酶SlCDK8基因及应用 Download PDF

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CN112126633B
CN112126633B CN202011038566.5A CN202011038566A CN112126633B CN 112126633 B CN112126633 B CN 112126633B CN 202011038566 A CN202011038566 A CN 202011038566A CN 112126633 B CN112126633 B CN 112126633B
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祝英方
许睿
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Abstract

本发明公开了中介体成员番茄周期素依赖性激酶SlCDK8基因及应用,本发明首次从番茄中获得番茄周期素依赖性激酶SlCDK8基因,并且通过利用农杆菌介导的方法将SlCDK8在番茄MT(micro‑Tom)中敲除进行功能验证,得出SlCDK8基因缺失可同时诱导整株番茄各器官的畸形发育,出现株高降低、雄性不育、花器官异常等症状,这也体现了SlCDK8在植物发育的进程中扮演的关键作用。

Description

番茄周期素依赖性激酶SlCDK8基因及应用
技术领域
本发明属于生物技术领域,具体涉及番茄周期素依赖性激酶SlCDK8基因及应用。
背景技术
育种过程中最为复杂的环节之一就是人工去雄,利用雄性不育即可免去人工去雄这一步骤,简化制种程序,降低种子成本的同时,还可提高种子纯度。随着基因工程技术的不断发展,可通过基因工程技术创制雄性不育突变体,对番茄花粉败育相关基因进行精确编辑,加速我国番茄雄性不育发展进程。
番茄(Solanumlycopersicum)是全世界非常重要的果实类作物以及模式作物,越来越受到研究者和育种家的重视。随着2012年,番茄的基因组测序,近年来对番茄的风味、代谢组、变异组和转录组等大数据库的相关研究发展迅速。介体复合体是真核转录机制的重要组成部分。作为转录因子和RNA聚合酶II之间的重要纽带,中介体将不同的信号转导到不同途径中的各类基因中,从而调节各种反应。植物中介体对植物生长、发育、防御、开花、基因组稳定性和代谢动态平衡均会产生不同程度的影响。番茄周期素依赖性激酶SlCDK8是中介体中的细胞周期蛋白依赖性激酶模块的重要成员。植物中的CDK8最开始是从苜蓿中发现的,之后在拟南芥中研究发现与花器官形成相关,所以又被命名为HUA ENHANCER3。目前的研究表明植物中CDK8在叶的细胞增殖,花组织的形成方面发挥着作用。对于植物中的CDK8是否还具有其它方面的应用还没有现有技术进一步报道,尤其是有关番茄花药和花粉发育调控相关机理的深入研究却鲜有报道。因此从基因方面研究花粉不育机制,对番茄雄性不育的发展进程,及番茄花药和花粉发育调控相关机理研究有重要的意义。
发明内容
本发明的目的在于提供番茄周期素依赖性激酶SlCDK8基因及应用。
为实现上述目的,本发明采取如下技术方案:
番茄周期素依赖性激酶SlCDK8基因,其核苷酸序列如SEQ ID NO.1所示。
上述番茄周期素依赖性激酶SlCDK8基因编码的蛋白选自,
(1)其氨基酸序列如SEQ ID NO:2所示;
(2)将SEQ ID NO:2氨基酸序列经过一个或多个((如1-30个;较佳地1-20个;更佳地1-10个;如5个,3个))氨基酸残基的取代、缺失或添加而形成的,且具有(1)蛋白功能的由(1)衍生的蛋白;或
(3)与(1)限定的蛋白序列有80%((较佳地90%以上,如95%,98%,99%或更高))以上同源性且具有(1)蛋白功能的由(1)衍生的蛋白。
也就是说本发明所保护的基因的功能,不仅包括上述番茄周期素依赖性激酶SlCDK8基因,还包括与SEQ ID NO:1具有较高同源性(如同源性高于40%;较佳地高于50%;较佳地高于60%;更佳地高于70%;更佳地高于80%;更佳地高于90%;更佳地高于95%;更佳地高于98%)的同源基因。
其中,序列中的SEQ ID NO.1由1404个碱基组成,自5’端第1位碱基为转录起始位点,第1402-1404位碱基为终止密码子,编码框为1401个碱基,共编码467个氨基酸。
并且含有上述基因的表达载体、重组载体或转基因细胞系以及含有所述载体的宿主细胞的获得方法也落入本发明的保护范围之内。
本发明最主要的目的是在分子水平上对番茄周期素依赖性激酶SlCDK8进行克隆和鉴定,从而为解析其在番茄花药和花粉发育调控相关机理的研究提供理论基础。
本发明还公开了抑制上述番茄周期素依赖性激酶SlCDK8基因的表达在如下方面的应用:
(1)矮化植物;
(2)使植物形成雄性不育表型;
上述应用通过敲除上述基因的方式来获得矮化、雄性不育症状的植物。
其中,矮化表现包括平均株高的高度降低,小叶表型包括叶片的长和宽均变小,雄性不育表型包括花粉活力降低,花粉畸形。
作为本发明的一种实施方式,作为本发明的一种实施方式,将多核苷酸通过常规的方法克隆到CRISRP载体中,将所述的带有外源基因的重组载体导入到可表达所述SlCDK8蛋白到植物细胞中,使所述的植物细胞中SlCDK8蛋白缺失。可通过将所述植物细胞再生成植物,获得SlCDK8基因缺失的突变体植物。并利用农杆菌转化法将重组质粒转入植物中。
本发明中,对于适用于本发明的植物没有特别的限制,只要其适合进行基因的转化操作,如各种农作物、花卉植物、或林业植物等。所述的植物比如可以是(不限于):双子叶植物、单子叶植物或裸子植物。
作为一种优选方式,所述的“植物”包括但不限于:茄科的番茄,凡是具有该基因或者与之同源的基因均适用。尤其适用于需要矮化的植物,比如蔷薇科苹果树、樱桃树;还尤其适用于需要形成雄性不育表型的植物,比如可以培育少籽番茄品种,无籽石榴等,提高口感;均可通过敲除SlCDK8基因获得矮化植株和雄性不育植株。
本发明具有如下优点:
(1)本发明首次筛选出番茄周期素依赖性激酶,通过克隆得到SlCDK8。
(2)发明人利用农杆菌介导的方法将SlCDK8从番茄MT中敲除进行功能验证,有利于从分子机制上阐明SlCDK8在调控花形态建成方面的作用,对进一步阐明花粉发育机制,培育番茄雄性不育新品种具有积极的指导作用,并且为番茄雄性不育系的研发提供理论基础。
(3)SlCDK8基因缺失可同时诱导整株番茄各器官的畸形发育,出现株高降低、雄性不育、花器官异常等症状,这也体现了SlCDK8在植物发育的进程中扮演的关键作用。
(4)对于一些需要矮化的植物如果树和观赏植物(减少不必要的养分消耗,以便充分利用光能,地力,提早结果、提高产量或增加观赏效果),可以通敲除SlCDK8基因的方式,为植物矮化育种提供一种新的途径。
附图说明
图1是6种不同编辑类型的SlCDK8敲除系;
图2是不同SlCDK8敲除系与野生型番茄WT的表型;
图3为野生型番茄WT、SlCDK8 CRP2(-3bp)和SlCDK8 CRP3(-6bp)的表型观察图;
图4是SlCDK8基因对番茄花粉的发育的影响。
具体实施方式
下面将通过具体实施例对本发明进行详细的描述。提供这些实施例是为了能够更透彻地理解本发明,并且能够将本发明的范围完整的传达给本领域的技术人员。
若未特别指明,实施例中所用技术手段为本领域技术人员所熟知的常规手段。下述实施例中的试验方法,如无特别说明,均为常规方法。如无特殊说明,所采用的试剂及材料,均可以从市场中购买获得。
除非另行定义,文中所使用的所有专业与科学用语与本领域熟练人员所熟悉的意义相同。此外,任何与所记载内容相似或均等的方法及材料皆可应用于本发明中。文中所述的较佳实施方法与材料仅作示范之用。
发明人通过生物信息学技术在番茄基因组基础上得到番茄周期素依赖性激酶SlCDK8基因序列。本发明将其与植物编辑载体连接后导入野生型番茄进行表型鉴定。
实施例番茄周期素依赖性激酶SlCDK8基因的分离和功能鉴定
1、SlCDK8基因的分离
MT番茄幼苗总RNA的提取:按照Trizol提取试剂(Takara)说明进行,第一链cDNA合成按照反转录试剂盒PrimerScriptTM II 1st Strand cDNA Synthesis Kit(Takara)说明进行。以番茄的cDNA为模板,使用Primer STAR Max高保真酶扩增(Takara),退火温度为58℃。
以下述序列为引物,通过PCR扩增获得SlCDK8基因的全长序列。SlCDK8基因的全长序列如序列表中SEQ ID NO:1所示,共1404bp,共编码蛋白的氨基酸序列如序列表中SEQ IDNO:2所示,共467个。
引物序列为:
SlCDK8-F:5′-ATGGGAGATGTCAGAGGAAAT-3′;
SlCDK8-R:5′-GAACCGCCTAGATTTCTGTTG-3′。
2、SlCDK8基因的功能鉴定试验
为研究SlCDK8基因在花粉发育发生过程中是否调控了花粉的形成,通过基因敲除系番茄来鉴定其功能。
2.1构建重组载体
在SlCDK8基因的第一外显子上选取靶位点,即GAGGAAATTCCAATAGCAAT。回收纯化sgRNA-SlCDK8片段并与酶切后的载体链接,从而获得SlCDK8-CRISPR重组质粒。并通过农杆菌介导进行番茄遗传转化,获得番茄SlCDK8-CRISPR的转化苗。
2.2转基因阳性株的筛选及表型分析
取抗性植株叶片100mg,采用CTAB法提取基因组总DNA。并根据靶基因序列设计特异性引物,进行PCR扩增,并送样测序。
引物序列为:
SlCDK8-g1-F:5′-TCCCAACTCTCCCTCGTACAAC-3′;
SlCDK8-g1-R:5′-TAGAGAGACATATCTGCCTGGTTGA-3′。
经测序鉴定,共获得6种不同编辑类型的SlCDK8敲除系,如SlCDK8CRP1(-2bp)、SlCDK8 CRP2(-3bp)、SlCDK8 CRP3(-6bp)、SlCDK8 CRP4(WT/+1bp)、SlCDK8 CRP5(WT/-12bp)、SlCDK8 CRP6(-6bp/-2bp)(图1、2)。图3为野生型番茄、SlCDK8 CRP2(-3bp)和SlCDK8CRP3(-6bp)的表型观察图,SlCDK8基因敲除系的平均株高均显著小于野生型对照,形成矮化表型;SlCDK8基因敲除系叶片大小的生长显著受到抑制,与野生型对照相比,其叶长、叶宽显著降低。从图4可以看出,SlCDK8敲除系与野生型相比,畸形花粉比例明显升高,花粉活力显著降低,形成雄性不育表型。
整体看来,与野生型和SlCDK8基因敲除系相比,基因敲除系植株表现出株高降低、叶长和叶宽降低,形成矮化表型。除此之外,SlCDK8基因对番茄花粉的发育存在明显影响,形成雄性不育表型。
以上所述之实施例,只是本发明的较佳实施例而已,仅仅用以解释本发明,并非限制本发明实施范围,对于本技术领域的技术人员来说,当然可根据本说明书中所公开的技术内容,通过置换或改变的方式轻易做出其它的实施方式,故凡在本发明的原理上所作的变化和改进等,均应包括于本发明申请专利范围内。
序列表
<110> 河南大学
<120> 番茄周期素依赖性激酶SlCDK8基因及应用
<130> 2020
<160> 2
<170> SIPOSequenceListing 1.0
<210> 1
<211> 1404
<212> DNA
<213> Solanum lycopersicum
<400> 1
atgggagatg tcagaggaaa ttccaatagc aatcggccag agtggttgca acagtatgat 60
ttgataggga aaattgggga agggacttac gggttggttt ttttggctaa gatcaaagct 120
aaccgtagca aatctattgc tatcaagaag ttcaagcaat ccaaggacgg tgacggcgtt 180
tcccccaccg ccattcgcga aatcatgttg cttcgggaga tatcccatga gaatgttgta 240
aagcttgtca atgtgcatat caaccaggca gatatgtctc tctatctcgc ttttgactat 300
gccgagcatg acctctatga aattatcaga caccatagag acaaggtcat tctttcaatc 360
aatccgtaca ctgtaaagtc tctgttatgg cagctgctta atggtctcaa ttatcttcac 420
agtaattgga ttgttcatcg agacctaaag ccatcaaaca tcttggtaat gggcgaggga 480
gaagaacatg gagttgttaa aattgctgat ttcggccttg caagaattta tcaagctcca 540
ctgaaaccat tagctgagaa tggggttgta gtcactatct ggtatcgtgc gccggagtta 600
ctgcttgggg ctaaacacta cacaagtgca gttgatatgt gggccgttgg ctgtatattt 660
gctgagcttc ttactctaaa gccactattt caagggcaag aagtgaaggg cacaccaaac 720
cctttccagc ttgaccaact ggacaagata tttaaggttc tcggacatcc cacaccagaa 780
aaatggccaa cactggtaaa ccttccgcat tggcaatctg atgtgcaacg aattcaagga 840
cataagtatg acaatcctgc actttatagt gttctgcaca tgtctcccaa aagtcctgca 900
tacgaccttc tttcaaagat gcttgaatat gatcctcgta aaagaataac agcaacacaa 960
gctctagagc atgagtactt ccggatggaa cctctacctg gccgcaatgc acttgtacca 1020
ccacaacctg gagaaaaagt tgtgaattat ccaactcgac ctgtggacac aactacagac 1080
tttgaaggaa ccattagtct tcaaacgtct cagccggtat catctggaaa tgcagcatct 1140
ggtggcatgc atggtcccca tgttatgcca actagatctg taccccgacc aatgcaaatg 1200
gtgaacatgc caaggatgca acctcagggc atgtctgctt ataatcttgc ttctcaagct 1260
ggcatgagtg ctggaatgaa ccctggaaac atgcccatgc agcgtggtgt tgctgctcag 1320
gcacatcagc agcagatgag aaggaaagat ccaggaatgg gaatacctgg atacccttca 1380
caacagaaat ctaggcggtt ctga 1404
<210> 2
<211> 467
<212> PRT
<213> Solanum lycopersicum
<400> 2
Met Gly Asp Val Arg Gly Asn Ser Asn Ser Asn Arg Pro Glu Trp Leu
1 5 10 15
Gln Gln Tyr Asp Leu Ile Gly Lys Ile Gly Glu Gly Thr Tyr Gly Leu
20 25 30
Val Phe Leu Ala Lys Ile Lys Ala Asn Arg Ser Lys Ser Ile Ala Ile
35 40 45
Lys Lys Phe Lys Gln Ser Lys Asp Gly Asp Gly Val Ser Pro Thr Ala
50 55 60
Ile Arg Glu Ile Met Leu Leu Arg Glu Ile Ser His Glu Asn Val Val
65 70 75 80
Lys Leu Val Asn Val His Ile Asn Gln Ala Asp Met Ser Leu Tyr Leu
85 90 95
Ala Phe Asp Tyr Ala Glu His Asp Leu Tyr Glu Ile Ile Arg His His
100 105 110
Arg Asp Lys Val Ile Leu Ser Ile Asn Pro Tyr Thr Val Lys Ser Leu
115 120 125
Leu Trp Gln Leu Leu Asn Gly Leu Asn Tyr Leu His Ser Asn Trp Ile
130 135 140
Val His Arg Asp Leu Lys Pro Ser Asn Ile Leu Val Met Gly Glu Gly
145 150 155 160
Glu Glu His Gly Val Val Lys Ile Ala Asp Phe Gly Leu Ala Arg Ile
165 170 175
Tyr Gln Ala Pro Leu Lys Pro Leu Ala Glu Asn Gly Val Val Val Thr
180 185 190
Ile Trp Tyr Arg Ala Pro Glu Leu Leu Leu Gly Ala Lys His Tyr Thr
195 200 205
Ser Ala Val Asp Met Trp Ala Val Gly Cys Ile Phe Ala Glu Leu Leu
210 215 220
Thr Leu Lys Pro Leu Phe Gln Gly Gln Glu Val Lys Gly Thr Pro Asn
225 230 235 240
Pro Phe Gln Leu Asp Gln Leu Asp Lys Ile Phe Lys Val Leu Gly His
245 250 255
Pro Thr Pro Glu Lys Trp Pro Thr Leu Val Asn Leu Pro His Trp Gln
260 265 270
Ser Asp Val Gln Arg Ile Gln Gly His Lys Tyr Asp Asn Pro Ala Leu
275 280 285
Tyr Ser Val Leu His Met Ser Pro Lys Ser Pro Ala Tyr Asp Leu Leu
290 295 300
Ser Lys Met Leu Glu Tyr Asp Pro Arg Lys Arg Ile Thr Ala Thr Gln
305 310 315 320
Ala Leu Glu His Glu Tyr Phe Arg Met Glu Pro Leu Pro Gly Arg Asn
325 330 335
Ala Leu Val Pro Pro Gln Pro Gly Glu Lys Val Val Asn Tyr Pro Thr
340 345 350
Arg Pro Val Asp Thr Thr Thr Asp Phe Glu Gly Thr Ile Ser Leu Gln
355 360 365
Thr Ser Gln Pro Val Ser Ser Gly Asn Ala Ala Ser Gly Gly Met His
370 375 380
Gly Pro His Val Met Pro Thr Arg Ser Val Pro Arg Pro Met Gln Met
385 390 395 400
Val Asn Met Pro Arg Met Gln Pro Gln Gly Met Ser Ala Tyr Asn Leu
405 410 415
Ala Ser Gln Ala Gly Met Ser Ala Gly Met Asn Pro Gly Asn Met Pro
420 425 430
Met Gln Arg Gly Val Ala Ala Gln Ala His Gln Gln Gln Met Arg Arg
435 440 445
Lys Asp Pro Gly Met Gly Ile Pro Gly Tyr Pro Ser Gln Gln Lys Ser
450 455 460
Arg Arg Phe
465

Claims (3)

1.抑制番茄周期素依赖性激酶SlCDK8基因的表达在矮化植物和/或使植物形成雄性不育表型中的应用,其特征在于,其核苷酸序列如SEQ ID NO.1所示。
2.权利要求1所述的应用,其特征在于,所述番茄周期素依赖性激酶SlCDK8基因的氨基酸序列如SEQ ID NO:2所示。
3.根据权利要求1所述的应用,其特征在于,通过敲除SlCDK8基因的方式来获得矮化和/或雄性不育症状的植物。
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