CN109097372A - 一种调控杨树不定根形成和茎发育的关键基因PeRR12及其表达蛋白和应用 - Google Patents
一种调控杨树不定根形成和茎发育的关键基因PeRR12及其表达蛋白和应用 Download PDFInfo
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Abstract
本发明公开了一种调控杨树不定根形成和茎发育的关键基因PeRR12及其表达蛋白和应用,该关键基因PeRR12的核苷酸序列如SEQ ID NO.1所示。本发明通过将PeRR12基因转入杨树,过量表达PeRR12基因的转基因杨树表现出生根时间推迟,生根率下降,不定根数目减少;添加外源生长素NAA后能部分恢复转基因杨树的生根缺陷,生根时间有所提前,生根率也明显上升,不定根数目也明显增加;同时,与未转基因杨树相比,转基因杨树的株高显著增加,表明过量表达PeRR12基因促进了杨树茎的生长。
Description
技术领域
本发明属于植物基因工程技术领域,具体涉及一种调控杨树不定根形成和茎发育的关键基因PeRR12及其表达蛋白和应用。
背景技术
杨树具有速生、适应力强、用途广等特点,作为用材林、防护林和四旁绿化的主要树种广泛栽培于中纬度地区,具有重要的经济和生态价值。尽管大多数杨树都比较容易扦插生根,然而仍有许多优良无性系扦插生根困难。因此开展杨树不定根发生发育的分子机理研究不仅具有重要的理论意义,而且蕴含潜在应用价值。然而,植物不定根的发育调控十分复杂,会受多重内源激素或环境因素的影响。因此,目前仅有少量基因被鉴定参与调控不定根发生,整个不定根发育的分子机理还十分薄弱。另外,树高直接影响到木材的产量,培育速生的品系一直是林木遗传改良与良种选育的重要目标。但是由于木本植物茎的生长发育机理十分复杂,有关调控杨树茎是生长发育的研究还很有限。
细胞分裂素和生长素能够通过相互作用,共同参与植物生长发育进程以及植物生物和非生物胁迫响应。细胞分裂素可以通过调控细胞分裂素受体AHK(Arabidopsishistidine kinase)激活其响应调节因子ARRs(Arabidopsis response regulator)及其下游基因的表达,进而调控植物生长发育过程。相关ARRs在细胞分裂素和生长素互作网络中扮演着关键角色,ARRs不仅能够调控植物的不定根、侧根、维管组织和茎尖分生组织的发育,还能够参与植物的免疫、盐、干旱、寒冷等多种胁迫响应。ARR基因分为A、B、C三种类型,其中A型包括ARR3、ARR4、ARR5、ARR6、ARR7、ARR8、ARR9,ARR15、ARR16、ARR17这10个家族成员;B型ARR基因包括ARR1,ARR2,ARR10,ARR11,ARR12、ARR13、ARR14,ARR18,ARR19,ARR20,ARR21、ARR23,为非细胞分裂素诱导型;C型包括ARR22和ARR24。A型ARRs的N端含有一个响应调控因子接收结构域(Response regulator receiver domain,RD),RD结构域中含有D-D-K保守序列,而C端较短,功能未知。A型ARRs是细胞分裂素诱导型基因,能够在细胞分裂素的诱导下快速响应,并通过负反馈控制细胞分裂素信号,从而抑制B型ARRs相关基因的表达。在拟南芥胚胎中,生长素通过正调控ARR7和ARR15等A型ARRs的表达,从而对细胞分裂素的积累产生抑制,调控胚胎发育。而在茎尖分生组织中,生长素通过正调控生长素响应因子ARF5,进而负调控ARR7和ARR15的表达,导致WUS基因的表达水平降低,调控茎尖的发育。同时WUS基因能够直接负调控ARR5、ARR6、ARR7和ARR15的表达,降低它们对细胞分裂素的负反馈。B型ARRs在结构上主要包括位于N端富含天冬氨酸及赖氨酸残基的响应调控因子接收结构域、DNA结合域以及富含谷氨酞胺的信号输出结构域。与A型ARRs不同,B型ARRs对细胞分裂素的诱导不敏感。B型ARRs是一类转录因子,在被A型ARRs抑制的同时,也能够能调控包括A型ARRs等下游基因的表达。B型ARRs负调控YUCCA基因的表达,影响芽的再生以及茎端分生组织的维持。
在拟南芥中,细胞分裂素在初生根的转运区(TZ)内能够介导细胞分化,而生长素在TZ区内介导细胞分裂,二者通过拮抗作用共同调节拟南芥根端分生组织。在细胞分裂素作用下,AHK3整调控ARR1/12的表达,ARR1/12与SHY2的启动子结合激活SHY2在TZ区的维管组织中进行特异表达。通过突变arr1发现,向arr1的突变体根部施加细胞分裂素无法导致SHY2的上调,而向野生型根中施加则发现SHY2的表达上调。通过激活SHY2,可以抑制生长素信号的转导,导致PIN在shy2-2突变体的mRNA中表达量较野生型降低,在shy2-31突变体的维管转变区中表达量升高。细胞分裂素可以通过AHK3/ARR1双元信号通路激活SHY2基因的转录抑制生长素信号响应,SHY2蛋白可以负调节PIN基因的表达。
细胞分裂素含量的增加会扰乱侧根中的生长素浓度梯度,破坏细胞的不对称分裂,从而抑制侧根的形成。侧根中细胞分裂素和生长素的互作的分子机理是不同于主根的。细胞分裂素可以通过抑制PIN1/3/7的转录,改变生长素的含量,从而抑制侧根的形成。另外,最近的研究表明,细胞分裂素可以通过控制AHK4和ARR2/12的表达,负调控PIN1的表达。细胞分裂素除了参与主根和侧根的发育外,还通过调控ARR促进茎维管形成层细胞的增殖,并抑制根原生木质部的分化。ARR家族除了调控植物的生长发育,还参与植物的免疫、盐、干旱、寒冷等非生物胁迫以及植物的叶片衰老。
虽然在拟南芥等草本模式植物中已结对ARRs基因进行了较为深入的研究,但是在木本植物中很少有相关具有针对性、系统性的深入研究报道阐述其作用机制。
发明内容
发明目的:针对现有技术中存在的不足,本发明的目的是提供一种调控杨树不定根形成和茎发育的关键基因PeRR12。本发明的另一目的提供一种调控杨树不定根形成和茎发育的关键基因PeRR12的应用。
技术方案:为了实现上述发明目的,本发明采用的技术方案如下:
一种调控杨树不定根形成和茎发育的关键基因PeRR12,其核苷酸序列如SEQ IDNO.1所示。
所述的杨树不定根形成和茎发育的关键基因PeRR12的表达蛋白,其氨基酸序列如SEQ ID NO.2所示。
含有所述的杨树不定根形成和茎发育的关键基因PeRR12的载体。
所述的载体,在PeRR12基因的5’端组装组成型强表达启动子P35S。
所述的载体,在PeRR12基因的3’端组装了强终止子NOS。
所述的载体,组装HPT基因表达盒,作为转基因杨树的筛选标记,可以用潮霉素进行转基因杨树的筛选。
所述的载体,组装LB和RB序列,促使组装于其间的PeRR12基因表达框架和筛选标记基因HPT整合至杨树受体细胞染色体中。
含有所述的杨树不定根形成和茎发育关键基因PeRR12的宿主细胞。
所述的杨树关键基因PeRR12在调控杨树不定根形成和茎发育中的应用。
本发明以南林895杨初生不定根为材料,通过RACE技术克隆了PeRR12基因。同时,采用Gateway克隆技术构建其杨树过量表达载体pH35GS-PeRR12,该基因位于启动子P35S之后,在启动子P35S的驱动下,PeRR12可在杨树体内高效表达,从而调控不定根和茎的形成和发育。其中,所PeRR12基因是杨树不定根发生发育关键基因。
有益效果:与现有技术相比,本发明通过将PeRR12基因转入杨树,过量表达PeRR12基因的转基因杨树表现出生根时间推迟,生根率下降,不定根数目减少;添加外源生长素NAA后能部分恢复转基因杨树的生根缺陷,生根时间有所提前,生根率也明显上升,不定根数目也明显增加;同时,与未转基因杨树相比,转基因杨树的株高显著增加,表明过量表达PeRR12基因促进了杨树茎的生长。
附图说明
图1是植物表达载体pH5GS的结构示意图;
图2为过量表达PeRR12基因的实时定量分子检测结果图;
图3为过量表达PeRR12基因的转基因杨与未转基因杨(CK)的整体形态比较结果图;
图4为过量表达PeRR12基因的转基因杨与未转基因杨(CK)的生根率结果图;
图5是过量表达PeRR12基因的转基因杨与未转基因杨(CK)的不定根数目以及株高的比较结果图;
图6是NAA处理下PeRR12转基因株系表型分析结果图。
具体实施方式
下面结合具体实施例对本发明做进一步的说明。
实施例1通过RACE技术克隆PeRR12基因
基于申请人前期杨树不定根表达谱芯片研究结果,使用Oligo 6设计3’末端的RACE引物,进行3'RACE,获得3’cDNA末端片段,克隆入T-载体,对插入片段进行PCR筛选后进行测序,Blast确认上述片段与其它植物相关的基因同源。以相同的方法,根据获得的正确的3’cDNA末端片段序列设计5’末端的RACE引物,进行5'RACE,获得5’cDNA末端片段,克隆入T-载体,对插入片段进行PCR筛选后进行测序。具体引物和主要过程如下:
PeRR12 3'Outer primer:5′-CCGATCCTTGAACAATCCTCTC-3′;
PeRR12 3'Inner inner:5′-CAACCCTCAGATGCTTCATGTGAACCCA-3′;
PeRR12 5'Outer primer:5′-CCCATCTTTTGACCTCACTAATG-3′;
PeRR12 5'Inner inner:
5′-CAGCTCACTGTGGCCACTCCCACTCGGCATG-3′;
1、3′RACE反应:
1)反转录,将以下成分加入到一个置于冰上的RNase-free的小离心管中:TotalRNA 1μg,dNTP Mix 4μL,3′RACE Adapter 2μL,10×RT Buffer 2μL,RNase Inhibitor 1μL,M-MLV Reverse Transcriptase 1μL,Nuclease-free Water补至20μL;
2)轻轻混匀,短暂离心,42℃温育1h;进入PCR步骤,或-20℃保存反应物;
3)3′RACE巢式PCR;Outer 3′RACE组成:10×LA PCR Buffer(Mg2+Free)5.0μL,MgCl2(25mM)5.0μL,dNTP Mixture(each 2.5mM)8.0μL,3′RACE gene-specific outerprimer 2.0μL,3′RACE Outer Primer(10μM)2.0μL,RT reaction product 1μL,TakaRa LATaq(5U/μl)0.5μL,Nuclease-free Water 26.5μL;Inner 3′RACE组成:5.0μL 10×LA PCRBuffer(Mg2+Free),5.0μL MgCl2(25mM),8.0μL dNTP Mixture(each 2.5mM),2.0μL 3′RACEgene specific inner primer,2.0μL 3′RACE Inner Primer(10μM),1μL Outer 3′RACEPCR product,0.5μL TakaRa LA Taq(5U/μL),26.5μL Nuclease-free Water;反应程序:94℃3min;94℃30sec,60℃30sec,72℃1min,35cycles;72℃7min。
4)纯化片段与克隆载体的连接反应:采用TaKaRa公司的pMD19-T simple Vetor克隆目的DNA分子,参考说明书,反应体系(5μL):2.2μL纯化回收的PCR产物,0.3μLpMD-19Simple Vector,2.5μL Solution I。反应条件:16℃30min;4℃过夜。
5)大肠杆菌转化:将新鲜制备或-70℃冻存的大肠杆菌TOP10感受态细胞在冰上融化;取5μL纯化片段与克隆载体的连接产物,加入到100μL感受态细胞中,并轻轻混匀,冰浴30min左右;42℃水浴中热击90sec,迅速置于冰上3-5min;加入800μL LB液体培养基,37℃&100rmp摇菌1h;4000rmp离心3min,吸掉上层800μL培养基,混匀剩余菌液;将菌液涂抹于含有Amp的LB筛选培养板上,37℃倒置培养过夜。
6)阳性克隆筛选及测序分析
从筛选培养板上挑选单菌落接种于LB液体培养基中,37℃&250rmp摇菌过夜;直接以培养过夜的菌液为模板进行重组转化子的PCR检测。
20.0μL反应体系:10×PCR Buffer(Mg2+free)2.0μL,MgCl2(25mM)1.5μL,dNTPMixture(each 2.5mM)1.3μL,3′RACE gene specific inner primer(10μM)1.0μL,3′RACEInner Primer(10μM)1.0μL,菌液0.1μL,rTaq 1.0μL,Milli-Q Water 12.1μL。
反应程序:94℃3min;94℃30sec,60℃30sec,72℃1min,28cycles;72℃7min。
菌液PCR检测为阳性的克隆送英骏生物技术公司(上海)测序鉴定。
2、5′RACE反应程序
1)RNA处理:CIP反应,将如下成分加入到RNase-free的小离心管中:Total RNA10μg,10×CIP buffer2μL,Calf Intestine Alkaline Phosphatase(CIP)2μL,Nuclease-freeWater加至20μL。轻轻混匀,短暂离心;37℃温育1h;加入以下试剂到CIP反应离心管:Ammonium Acetate Solution15μL,Nuclease-free Water115μL,acid phenol:chloroform150μL;充分涡旋,室温高速离心(≥10000g)5min;;转移上层水相到一个新的离心管中,加入150μL氯仿,充分涡旋,室温高速离心(≥10000g)5min;转移上层水相到一个新的离心管中,加入150μL异丙醇,充分涡旋,冰浴10min;最大转速离心20min,用0.5mL预冷的70%乙醇冲洗沉淀,最大转速离心5min,小心弃乙醇,气干沉淀;以11μLNuclease-freeWater重悬沉淀,即得CIP’RNA,冰上放置进一步用于TAP反应,或者-20℃保存;TAP反应,把以下成分加入到一个RNase-free的小离心管中:CIP'd RNA(from fabove)5μL,10×TAPbuffer1μL,TobaccoAcidPyrophosphatase(TAP)2μL,Nuclease-free Water2μL;轻轻混匀,短暂离心,37℃温育1h,即得CIP/TAP-treated RNA;进入接头连接步骤,或-20℃保存反应物;5′RACE接头连接,把以下成分加入到一个RNase-free的小离心管中:CIP/TAP-treated RNA2μL,5′RACE Adapter1μL,10×RNA Ligase Buffer1μL,T4RNA Ligase(2.5U/μL)2μL,Nuclease-free Water4μL;轻轻混匀,短暂离心,37℃温育1h,即得Ligated RNA;进入反转录步骤,或-20℃保存反应物。
2)反转录:将以下成分加入到一个置于冰上的RNase-free的小离心管中:LigatedRNA2μL,dNTP Mix4μL,Random Decamers2μL,10×RT Buffer2μL,RNase Inhibitor1μL,M-MLV Reverse TranscriptaselμL,Nuclease-free Water补至20μL。轻轻混匀,短暂离心;42℃温育1h,即得RT reaction;进入PCR步骤,或-20℃保存反应物。
3)5′RACE巢式PCR:反应体系、反应条件与3′RACE的巢式PCR一致。
4)PCR产物克隆测序,与3′RACE克隆一致。
采用BioEdit软件对3′RACE和5′RACE序列进行比对拼接,并使用FGENESH(http://mendel.cs.rhul.ac.uk/mendel.php?topic=fgen)预测其阅读框。根据基因全长序列设计引物(扩增子包含起始密码子及终止密码子),再次进行PeRR12基因的全长扩增和测序验证。其中,PeRR12 ORF正向引物为:5'-ATGGGGAAGAATGTGTTGGT-3’,PeRR12 ORF反向引物为:5'-ATGCAGAGAATTGGTTCGAG-3’;高保真PCR反应体系如下:10×LA PCR Buffer(Mg2+free)5.0μL;2.5mM dNTP Mixture 8.0μl;25mM Mg2+5.0μl;LA Taq DNAPolymerase(5U/μL)0.5μL;正向引物(10μM)2μL;反向引物(10μM)2μL;模板(895杨cDNA)1μL;加无菌ddH2O补足50μL。反应程序:预变性94℃3min;94℃40s,55℃30s,72℃30s,35个循环;72℃10min。
最终获得PeRR12全长cDNA序列为2558bp,包含一个2031bp的完整阅读框,序列如SEQ ID NO.1所示,其所表达的蛋白的氨基酸序列如SEQ ID NO.2所示。
实施例2PeRR12基因植物表达载体构建
利用通路克隆技术构建PeRR12基因的过量表达载体。使用特异PCR引物(实施例1的PeRR12ORF引物),以cDNA为模板,进行PCR扩增,将PeRR12基因ORF构建到入门载体。入门载体为pCRTM8/GW/TOPOTM vector(Invitrogen)。反应体系为:Fresh PCR product(purified)10-20ng;Salt solution 1μL;pCRTM8/GW/TOPOTM vector 1μL;加无菌ddH2O补足6μL。反应程序为:室温静置30min。
从筛选培养板上挑取阳性克隆进行PCR检测及测序验证,带PeRR12基因的入门载体与植物表达载体pH35GS进行LR反应。载体质粒如图1所示。反应体系为:linearizedentry clone 100ng;purified destination vector(100ng/μL)1.5μL;LR Clonase IIenzyme mix 2μL;加TE(pH 8.0)补足10μL。反应条件:25℃1h。经LR反应后PeRR12基因导入植物表达载体pH35GS中,在PeRR12基因的5’端组装组成型强表达启动子P35S,它能使PeRR12基因在杨树体内高效表达;在PeRR12基因的3’端组装了强终止子NOS,可有效终止PeRR12基因的转录;在载体质粒上组装HPT基因表达盒,作为转基因杨树的筛选标记,可以用潮霉素进行转基因杨树的筛选;在载体质粒组装LB和RB序列,促使组装于其间的PeRR12基因表达框架和筛选标记基因HPT整合至杨树受体细胞染色体中。通过PCR检测及测序验证,确认过量表达载体构建成功,命名为pH35GS-PeRR12,该基因位于启动子P35S之后,在启动子P35S的驱动下,PeRR12可在杨树体内高效表达。
实施例3PeRR12基因的遗传转化
通过液氮冻融法将所构建的pH35GS-PeRR12过量表达载体转入农杆菌菌株EHA105(Invitrogen),通过农杆菌介导将PeRR12基因转入杨树。实验结果如图2~6所示,其中,图2为过量表达PeRR12基因的实时定量分子检测;图3为过量表达PeRR12基因的转基因杨与未转基因杨(CK)的整体形态比较;图4为过量表达PeRR12基因的转基因杨与未转基因杨(CK)的生根时间和生根率;图5是过量表达PeRR12基因的转基因杨与未转基因杨(CK)的不定根数目以及株高的比较;图6是NAA处理下PeRR12转基因株系表型分析。从结果可以明显得出,过量表达PeRR12基因的转基因杨树表现出生根时间推迟,生根率下降,不定根数目减少;添加外源生长素NAA后能部分恢复转基因杨树的生根缺陷,生根时间有所提前,生根率也明显上升,不定根数目也明显增加;同时,与未转基因杨树相比,转基因杨树的株高显著增加,表明过量表达PeRR12基因促进了杨树茎的生长。
序列表
<110> 南京林业大学
<120> 一种调控杨树不定根形成和茎发育的关键基因PeRR12及其表达蛋白和应用
<130> 100
<160> 8
<170> SIPOSequenceListing 1.0
<210> 1
<211> 2558
<212> DNA
<213> 南林895杨(P. deltoides ×P. euramericana‘Nanlin895’)
<400> 1
gaaaatattc caagttacca atgaagatta ttttttgttg tcactttctc tctcaactga 60
ccaattttca agctctgcaa ttattatggt ttctaggttt ttgttaaaag ctccaaactt 120
tgaaacaaaa tccagttaaa gctccaaact ttgaagcaaa ttgtagtacc ccactctgta 180
aattagtctc tttctttcat attttgcttc gtcaaattgt tttgaagagg aaaaaatgac 240
tgtagaacag gggattggag atagcaatat agatcagttt cctattggta tgagagtttt 300
ggctgttgat gatgacccaa cttgtctttt actgttagag actctactcc gaagatgcca 360
gtacaatgtt actacgacga gtcaggcaat cacggcattg aggatgttga gaggaaacaa 420
gaacaagttt gacctggtta tcagtggtgt ccatatgcca gacatggatg gttttaaatt 480
gctggagctc gtggggcttg agatggacct acctgtcatc atgttgtcag caaacggtga 540
tccaaagctt gtgatgaagg ggatcactca tggagcttgt tattatttgc tcaagcctgt 600
tcgaattgag gagctaaaga ccatctggca acatgtgatt aggagaaaga aaagtgataa 660
caaagacaga aacagctcgg ataaccgaga caagcctaat caaggaagta gtgaagcagt 720
acctgaccag aagcttaata aaaaaagaaa agaccagaat ggagacgagg atgaggatca 780
tgatgaggac gaggacgagc atgagcatga aaatgaggat ccaacaaccc aaaagaagcc 840
ccgggttgta tggtcagtgg agctgcatcg caagtttgtt gcagccgtta atcaattggg 900
cgttgacaag gctgttccaa agaagattct tgatttgatg aatgttgaaa agcttactag 960
agaaaatgtg gcaagccatc tccagaaata taggcattac cttaaaagga tcagcactgt 1020
ggcaaaccag caagctaata tggtggcagc tctgggcagt tcagatgcat catacctgca 1080
catgaattct atgagtggac tgggattgca cagcttggct ggatctgtgc agtttcacag 1140
cacacccttc agatctctcc catcttgtgg aatgcttgat aggttgaatt ctcctgctgt 1200
tctgggcatt cacggtcttc cttctcctgg agtgattcaa ttaggtcatg tacagactgc 1260
accccacacg gctaatggtc caagtcacta ccaaccagtt cgacacccag gaaataatgg 1320
gaatatactg caagggatgc caatgccatt agaacttgat caaatacaat ccaacaaggg 1380
tgtgaactac attccggaac ttcctactca tcttgatgac acagccagtt tccctgtttc 1440
cagtggttcc acagatatga aaataattgc gggaagctca aacagccctt ttgttggtgt 1500
ttcaaacaaa cacctgatgt tggaaggaca tggtcaaggg ctccaaggtg gccaaaaatc 1560
tggaaagcaa tcttctcttt cagcagggtc tttaaatcca ggatattctt cccattttcc 1620
agatcatggt agacgtaatg acaactggtc caatgctgtt cagtcaaatg gggcccagtc 1680
agattctttt acgttgaatg actattttaa gcaatctact ctgcatccta gtgccatcgg 1740
agaccgcatg tcaacaatgg tcttgcaaag caggaataat ccttctgatg tttcttctgt 1800
ctctactctt cccatacatt tacaggactc caaagcagat ctaccatgcc gagtgggagt 1860
ggccacagtg agctgtaatg ctggacagct aattaataat tgctctctag agtgggatga 1920
ccatagacag gatgatcctt accactcaaa tggcctatct aactcaataa actcagcaat 1980
tcctattaat ggtaatggga gtacgaatgg gttcaatttg gatccaaata acctgttttt 2040
tcagagaact acaagcttca tttcaactgg accgtcaaat tttgttgata cctcattgat 2100
gaagcataat gaggttgaat gttcagccat ggaaacatta gtgaggtcaa aagatgggaa 2160
tttgctaggc caagagaagc cacaggacag ttctgtttcg aataattttg gctccttgga 2220
aggtttggtg agcgtaatga tcaatcagcc aaccattctt ggttaggaga atagtctggg 2280
aacaagacag gtgaagttac cagaaggaga ctttggatgt ggtggttatt ctctaaggaa 2340
ttgcatataa agcagattta aactgcgaag gtctctaaag aggatgatct gcagaagctt 2400
tgtgctttct tgcagatgag ttattcctag tgtaattgat actcaatata tctatgtatt 2460
ccacttctaa ttttctttag agaactgtat gttaattgca gttgttttgt ccttctcaaa 2520
ttcggcagag tactttgttt ttacctaaaa aaaaaaaa 2558
<210> 2
<211> 676
<212> PRT
<213> 南林895杨(P. deltoides ×P. euramericana‘Nanlin895’)
<400> 2
Met Thr Val Glu Gln Gly Ile Gly Asp Ser Asn Ile Asp Gln Phe Pro
1 5 10 15
Ile Gly Met Arg Val Leu Ala Val Asp Asp Asp Pro Thr Cys Leu Leu
20 25 30
Leu Leu Glu Thr Leu Leu Arg Arg Cys Gln Tyr Asn Val Thr Thr Thr
35 40 45
Ser Gln Ala Ile Thr Ala Leu Arg Met Leu Arg Gly Asn Lys Asn Lys
50 55 60
Phe Asp Leu Val Ile Ser Gly Val His Met Pro Asp Met Asp Gly Phe
65 70 75 80
Lys Leu Leu Glu Leu Val Gly Leu Glu Met Asp Leu Pro Val Ile Met
85 90 95
Leu Ser Ala Asn Gly Asp Pro Lys Leu Val Met Lys Gly Ile Thr His
100 105 110
Gly Ala Cys Tyr Tyr Leu Leu Lys Pro Val Arg Ile Glu Glu Leu Lys
115 120 125
Thr Ile Trp Gln His Val Ile Arg Arg Lys Lys Ser Asp Asn Lys Asp
130 135 140
Arg Asn Ser Ser Asp Asn Arg Asp Lys Pro Asn Gln Gly Ser Ser Glu
145 150 155 160
Ala Val Pro Asp Gln Lys Leu Asn Lys Lys Arg Lys Asp Gln Asn Gly
165 170 175
Asp Glu Asp Glu Asp His Asp Glu Asp Glu Asp Glu His Glu His Glu
180 185 190
Asn Glu Asp Pro Thr Thr Gln Lys Lys Pro Arg Val Val Trp Ser Val
195 200 205
Glu Leu His Arg Lys Phe Val Ala Ala Val Asn Gln Leu Gly Val Asp
210 215 220
Lys Ala Val Pro Lys Lys Ile Leu Asp Leu Met Asn Val Glu Lys Leu
225 230 235 240
Thr Arg Glu Asn Val Ala Ser His Leu Gln Lys Tyr Arg His Tyr Leu
245 250 255
Lys Arg Ile Ser Thr Val Ala Asn Gln Gln Ala Asn Met Val Ala Ala
260 265 270
Leu Gly Ser Ser Asp Ala Ser Tyr Leu His Met Asn Ser Met Ser Gly
275 280 285
Leu Gly Leu His Ser Leu Ala Gly Ser Val Gln Phe His Ser Thr Pro
290 295 300
Phe Arg Ser Leu Pro Ser Cys Gly Met Leu Asp Arg Leu Asn Ser Pro
305 310 315 320
Ala Val Leu Gly Ile His Gly Leu Pro Ser Pro Gly Val Ile Gln Leu
325 330 335
Gly His Val Gln Thr Ala Pro His Thr Ala Asn Gly Pro Ser His Tyr
340 345 350
Gln Pro Val Arg His Pro Gly Asn Asn Gly Asn Ile Leu Gln Gly Met
355 360 365
Pro Met Pro Leu Glu Leu Asp Gln Ile Gln Ser Asn Lys Gly Val Asn
370 375 380
Tyr Ile Pro Glu Leu Pro Thr His Leu Asp Asp Thr Ala Ser Phe Pro
385 390 395 400
Val Ser Ser Gly Ser Thr Asp Met Lys Ile Ile Ala Gly Ser Ser Asn
405 410 415
Ser Pro Phe Val Gly Val Ser Asn Lys His Leu Met Leu Glu Gly His
420 425 430
Gly Gln Gly Leu Gln Gly Gly Gln Lys Ser Gly Lys Gln Ser Ser Leu
435 440 445
Ser Ala Gly Ser Leu Asn Pro Gly Tyr Ser Ser His Phe Pro Asp His
450 455 460
Gly Arg Arg Asn Asp Asn Trp Ser Asn Ala Val Gln Ser Asn Gly Ala
465 470 475 480
Gln Ser Asp Ser Phe Thr Leu Asn Asp Tyr Phe Lys Gln Ser Thr Leu
485 490 495
His Pro Ser Ala Ile Gly Asp Arg Met Ser Thr Met Val Leu Gln Ser
500 505 510
Arg Asn Asn Pro Ser Asp Val Ser Ser Val Ser Thr Leu Pro Ile His
515 520 525
Leu Gln Asp Ser Lys Ala Asp Leu Pro Cys Arg Val Gly Val Ala Thr
530 535 540
Val Ser Cys Asn Ala Gly Gln Leu Ile Asn Asn Cys Ser Leu Glu Trp
545 550 555 560
Asp Asp His Arg Gln Asp Asp Pro Tyr His Ser Asn Gly Leu Ser Asn
565 570 575
Ser Ile Asn Ser Ala Ile Pro Ile Asn Gly Asn Gly Ser Thr Asn Gly
580 585 590
Phe Asn Leu Asp Pro Asn Asn Leu Phe Phe Gln Arg Thr Thr Ser Phe
595 600 605
Ile Ser Thr Gly Pro Ser Asn Phe Val Asp Thr Ser Leu Met Lys His
610 615 620
Asn Glu Val Glu Cys Ser Ala Met Glu Thr Leu Val Arg Ser Lys Asp
625 630 635 640
Gly Asn Leu Leu Gly Gln Glu Lys Pro Gln Asp Ser Ser Val Ser Asn
645 650 655
Asn Phe Gly Ser Leu Glu Gly Leu Val Ser Val Met Ile Asn Gln Pro
660 665 670
Thr Ile Leu Gly
675
<210> 3
<211> 22
<212> DNA
<213> PeRR12 3'Outer primer(Artificial)
<400> 3
ccgatccttg aacaatcctc tc 22
<210> 4
<211> 28
<212> DNA
<213> PeRR12 3'Inner inner(Artificial)
<400> 4
caaccctcag atgcttcatg tgaaccca 28
<210> 5
<211> 23
<212> DNA
<213> PeRR12 5'Outer primer(Artificial)
<400> 5
cccatctttt gacctcacta atg 23
<210> 6
<211> 31
<212> DNA
<213> PeRR12 5'Inner inner(Artificial)
<400> 6
cagctcactg tggccactcc cactcggcat g 31
<210> 7
<211> 20
<212> DNA
<213> PeRR12 ORF正向引物(Artificial)
<400> 7
atggggaaga atgtgttggt 20
<210> 8
<211> 20
<212> DNA
<213> PeRR12 ORF反向引物(Artificial)
<400> 8
atgcagagaa ttggttcgag 20
Claims (9)
1.一种调控杨树不定根形成和茎发育的关键基因PeRR12,其核苷酸序列如SEQ IDNO.1所示。
2.权利要求1所述的杨树不定根形成和茎发育的关键基因PeRR12的表达蛋白,其氨基酸序列如SEQ ID NO.2所示。
3.含有权利要求1所述的杨树不定根形成和茎发育的关键基因PeRR12的载体。
4.根据权利要求3所述的载体,其特征在于:所述载体在PeRR12基因的5’端组装组成型强表达启动子P35S。
5.根据权利要求3所述的载体,其特征在于:所述载体在PeRR12基因的3’端组装了强终止子NOS。
6.根据权利要求3所述的载体,其特征在于:所述载体组装HPT基因表达盒,作为转基因杨树的筛选标记,可以用潮霉素进行转基因杨树的筛选。
7.根据权利要求3所述的载体,其特征在于:所述载体组装LB和RB序列,促使组装于其间的PeRR12基因表达框架和筛选标记基因HPT整合至杨树受体细胞染色体中。
8.含有权利要求1所述的杨树不定根形成和茎发育关键基因PeRR12的宿主细胞。
9.权利要求1所述的杨树关键基因PeRR12在调控杨树不定根形成和茎发育中的应用。
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CN111560381A (zh) * | 2020-05-21 | 2020-08-21 | 扬州大学 | 一种杨树不定根形成关键基因PeSAUR72及其应用 |
CN111560381B (zh) * | 2020-05-21 | 2021-09-07 | 扬州大学 | 一种杨树不定根形成关键基因PeSAUR72及其应用 |
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