CN113308479B - SlNAC100基因在提高番茄低温抗性中的应用 - Google Patents
SlNAC100基因在提高番茄低温抗性中的应用 Download PDFInfo
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- CN113308479B CN113308479B CN202110802406.1A CN202110802406A CN113308479B CN 113308479 B CN113308479 B CN 113308479B CN 202110802406 A CN202110802406 A CN 202110802406A CN 113308479 B CN113308479 B CN 113308479B
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Abstract
本发明公开了SlNAC100基因在番茄低温抗性中的应用,该应用为通过基因过表达技术使得所述番茄SlNAC100基因的表达水平上升,所述番茄SlNAC100基因的核苷酸序列:SEQ ID NO:1所示的核苷酸序列。本发明为培育耐低温番茄新品种提供了基因资源,具有潜在的应用价值,为研究番茄应答逆境信号的分子机制和提高对不利环境耐受性的机理奠定了理论基础。
Description
技术领域
本发明涉及基因工程及分子生物学领域,具体地说,涉及一种SlNAC100基因在提高番茄低温抗性中的应用。
背景技术
番茄(Solanum lycopersicum L.)原产于南美洲,属于茄科作物,是世界上广泛栽培的蔬菜,其最适宜的生长温度约20-25℃,对低温较为敏感。然而作为世界面积最大的设施栽培国家,我国各地冬春季节的冷害以及设施环境中早晚的低温极大地影响了番茄等喜温植物的光合作用和生长发育,导致其产量和品质急剧下降。因此,研究番茄对低温胁迫的响应机制及其分子机理,挖掘提高番茄耐低温能力的关键基因具有重要意义。
在低温胁迫响应过程中,应激信号网络的启动通常可以有效、及时地诱导各种应激反应基因的表达,而如此大量的差异基因的表达需要由不同类型的转录因子(TFs)以时间和空间方式协同作用来共同调控。通常,转录因子主要起信号传递者的作用,通过与几种基因启动子区域中的顺式作用元件特异性结合而将外部胁迫信号传递至细胞核,从而影响植物中与胁迫相关的多个基因的转录进而参与多种进程。NAC转录因子是植物中所特有的一类转录调控因子。NAC转录因子编码的蛋白质的N端高度保守,包含A、B、C、D、E 5个亚结构域,其中A、C、D亚结构域保守性高,B和E保守性不强,其C端具有多样性及转录激活的特点。大量研究表明,在植物生长发育、器官形成、激素调节和病害防御等多种胁迫方面NAC转录因子均发挥着重要作用。在植物生长发育方面,拟南芥CUC1,CUC2,CUC3基因与顶端分生组织及花器官形成有关,NST1,NST2,SND1,SND2,VND7等基因与次生壁的形成及加厚有关。然而番茄NAC转录因子在低温胁迫中的作用及其调控机制均鲜见报道。因此,通过对SlNAC100基因的克隆、转基因技术培育SlNAC100不同表达量的番茄材料,在提高番茄对低温胁迫的抗性以及挖掘逆境胁迫基因资源方面,具有很好的应用前景。
发明内容
本申请实施例的目的是提供一种SlNAC100基因在提高番茄低温抗性中的应用。
根据本申请实施例的第一方面,提供一种提高番茄耐低温能力的基因,所述基因为番茄SlNAC100基因,所述番茄SlNAC100基因具有SEQ ID NO:1所示的核苷酸序列。
根据本申请实施例的第二方面,提供番茄SlNAC100基因在提高番茄耐低温能力中的应用,通过基因过表达技术使得所述番茄SlNAC100基因的表达水平上升,所述番茄SlNAC100基因的核苷酸序列:SEQ ID NO:1所示的核苷酸序列。
进一步地,所述基因过表达技术包括:
提取番茄总RNA,反转录获得cDNA,以cDNA为模板,F和R为引物,扩增SlNAC100基因,将扩增产物构建到植物过表达载体上,其中F引物和R引物的核苷酸序列分别如SEQ IDNO:3和4所示;
将植物过表达载体导入宿主细胞中,再利用其侵染目的植株,筛选阳性转基因植株,获得耐低温的转基因番茄植株。
进一步地,所述植物过表达载体为具有35S启动子的表达载体。
进一步地,所述的宿主细胞为大肠杆菌细胞或农杆菌细胞。
进一步地,所述的农杆菌细胞为EHA105。
进一步地,所述植物过表达载体为载体pFGC1008-HA或载体pCAMBIA1301。
根据本申请实施例的第三方面,提供一种提高番茄耐低温能力的蛋白,所述蛋白为番茄SlNAC100蛋白,所述番茄SlNAC100蛋白具有SEQ ID NO.2所示的氨基酸序列。
根据本申请实施例的第四方面,提供番茄SlNAC100蛋白在提高番茄耐低温能力中的应用,通过基因过表达技术使得所述番茄SlNAC100蛋白的表达水平上升,所述番茄SlNAC100蛋白为的氨基酸序列:SEQ ID NO:2所示的氨基酸序列。本申请的实施例提供的技术方案可以包括以下有益效果:
由上述实施例可知,本申请采用基因手段构建番茄SlNAC100过表达植株或基因敲除植株,使番茄SlNAC100基因表达水平增强或抑制,调控所述基因SlNAC100的表达水平来研究其对番茄低温抗性的调控机制。结果发现,SlNAC100过表达植株对番茄营养生长和生殖生长均没有显著影响,但可以提高PSII最大光化学效率(Fv/Fm),以及抗冷相关基因COR47LIKE,COR6.6LIKE表达,降低相对电导率,从而提高番茄植株抗冷性。
本发明首次构建了番茄SlNAC100基因过表达和基因敲除的转基因植株,并进行功能研究。本发明提供的SlNAC100基因为培育耐低温番茄新品种提供了基因资源,具有潜在的应用价值,为研究番茄应答逆境信号的分子机制和提高对不利环境耐受性的机理奠定了理论基础。
应当理解的是,以上的一般描述和后文的细节描述仅是示例性和解释性的,并不能限制本申请。
附图说明
此处的附图被并入说明书中并构成本说明书的一部分,示出了符合本申请的实施例,并与说明书一起用于解释本申请的原理。
图1为本申请实施例中番茄SlNAC100过表达株系#6的植物蛋白Western Blot检测结果;
图2为本申请实施例中SlNAC100基因敲除番茄株系的sgRNA序列的测序结果。
图3为本申请实施例中野生型,SlNAC100过表达植株和SlNAC100 CRISPR/Cas9基因敲除植株在低温处理7天后的表型;
图4为本申请实施例中野生型,SlNAC100过表达植株和SlNAC100 CRISPR/Cas9基因敲除植株在低温处理7天后的相对电导率;
图5为本申请实施例中野生型,SlNAC100过表达植株和SlNAC100 CRISPR/Cas9基因敲除植株在低温处理7天后的PSII最大光化学效率(Fv/Fm)变化。
图6为本申请实施例中野生型,SlNAC100过表达植株和SlNAC100 CRISPR/Cas9基因敲除植株在低温处理12h抗冷基因表达水平的变化;其中,A为番茄COR47like基因的表达水平,B为番茄COR6.6like基因的表达水平。
具体实施方式
下面结合实施例对本发明作进一步描述,但本发明的保护范围并不仅限于此。
实施例1:SlNAC100基因过表达载体的构建
SlNAC100基因来源于番茄,在番茄基因组数据库中的编号为Solyc06g069710,为探究SlNAC100过表达对番茄耐低温能力的影响,首先从番茄基因组中克隆了SlNAC100基因。根据编码区序列分析,设计特异性引物SlNAC100-F和SlNAC100-R,并在引物上分别加上限制性酶切位点(AscI和KpnI),序列如SEQ ID NO:3和4所示。用PrimerSTAR高保真酶PCR扩增SlNAC100片段,然后对PCR扩增片段及载体进行酶切,将SlNAC100片段连接到pFGC1008-HA上,得到植物过表达载体pFGC1008::SlNAC100-HA。将上述重组质粒送到尚亚公司测序确认,所得的基因SlNAC100的核苷酸序列如SEQ ID NO:1所示;该基因编码的蛋白质的氨基酸序列如SEQ ID NO:2所示。结果表明所克隆的序列与Solgenomics中公布的序列(Solyc06g069710)一致,提取阳性质粒备用,命名为pFGC1008::SlNAC100-HA。
实施例2:SlNAC100 CRISPR/Cas9基因敲除载体的构建
为探究SlNAC100缺失对番茄耐低温能力的影响,我们设计SlNAC100的靶基因序列,通过酶切连接构建pCAMBIA1301-U6-26-sgRNA1-SlNAC100-35S-cas9SK载体,利用CRISPR/Cas9技术敲除SlNAC100来进行研究。
首先,利用CRISPR-P网站((http://cbi.hzau.edu.cn/cgi-bin/CRISPR)设计SlNAC100基因的靶序列,具体序列如SEQ ID NO:5所示,为sgRNA1:5’-TGTTAAGGATGATGATCAGA-3’。将合成的sgRNA1序列(单链)进行退火,形成双链sgRNA1,同时其两端具有BbsI限制性内切酶酶切位点。将形成的sgRNA1与BbsI限制性内切酶酶切过的AtU6-26SK载体进行连接,提取阳性质粒备用,命名为U6-26-sgRNA1-SlNAC100-SK。利用KpnI与SalI限制性内切酶同时对U6-26-sgRNA1-SlNAC100-SK和35S-Cas9SK载体进行双酶切,将各自酶切产物回收并将酶切过的U6-26-sgRNA1-SlNAC100-SK片段连接到同样酶切过的35S-Cas9SK载体上。菌液PCR检测引物为,U6-26-F:5’-GACGGCCAGTGAATTGTA-3’,U6-26-R:5’-TATCTAAGCGATGTGGGACT-3',测序验证阳性克隆,提取阳性质粒备用,命名为U6-26-sgRNA1-SlNAC100-35S-cas9SK。利用KpnI与XbaI限制性内切酶同时对U6-26-sgRNA1-SlNAC100-35S-cas9SK和pCAMBIA1301载体进行双酶切,U6-26-sgRNA1-SlNAC100-35S-cas9SK回收约6kb的条带,即U6-26-sgRNA1-SlNAC100-35S-cas9片段,连到酶切过的pCAMBIA1301载体上。连接产物转化大肠杆菌DH5α感受态细胞,挑取单菌落,于含50mg/L卡那霉素(Kan)的液体LB培养基中,37℃,200rpm振荡培养过夜。在pCAMBIA1301载体的5’端设计引物进行菌液PCR检测(~550bp),上下游引物分别为,U6-26-Cas9-F:5'-GCTCGTATGTTGTGTGGAAT-3',U6-26-Cas9-R:5'-TATCTAAGCGATGTGGGACT-3'。测序验证阳性克隆,提取阳性质粒备用,命名为pCAMBIA1301-U6-26-sgRNA1-SlNAC100-35S-cas9。
实施例3:SlNAC100转基因植株的获得
将植物过表达载体pFGC1008::SlNAC100-HA和基因编辑载体pCAMBIA1301-U6-26-sgRNA1-SlNAC100-35S-cas9,通过电击法转化农杆菌GV3101,并进行野生型(Ailsa Craig)番茄子叶侵染,通过诱导愈伤,抗性诱导分化以及生根培养,获得组培苗,将T1代突变体种子和过表达种子分别进行卡那霉素抗性和氯霉素抗性的测试,选择3/4具有抗性而其余1/4没有抗性的株系,说明在该株系中连有目的基因的过表达载体以单拷贝形式插入。将这些植株移除,再进行单株收种。利用Western Blot验证SlNAC100过表达阳性转基因植株,结果显示野生型没有蛋白条带,而过表达株系有SlNAC100-HA的条带(图1),利用PCR和测序技术验证阳性SlNAC100突变转基因植株,发现SlNAC100#2缺失1个碱基,SlNAC100#4缺失21个碱基(图2)。
实施例4:SlNAC100转基因植株耐低温能力观察
试验选用的番茄品种为野生型Ailsa Craig和实施例3中所得的SlNAC100过表达#6和SlNAC100 CRISPR/Cas9株系#2和#4,将种子播种于盛满3:1草炭和蛭石复合栽培基质的塑料盆中,出苗后按照基质水分情况浇水保持基质湿润,整个过程浇霍格兰营养液,待四叶一心时进行低温处理,处理温度为恒温4℃。
试验共设8个处理:1)WT常温组;2)SlNAC100过表达#6;3)SlNAC100 CRISPR/Cas9#2常温组;4)SlNAC100 CRISPR/Cas9#4常温组;5)WT低温组;6)SlNAC100过表达#6;7)SlNAC100 CRISPR/Cas9#2低温组;8)SlNAC100 CRISPR/Cas9#4低温组。低温处理时间为7d。低温处理结束后进行表型拍摄、光系统II最大光化学效率测定和相对电导率测定。
光系统Ⅱ最大光化学效率具体测定方法为:将植株置于暗环境适应30分钟后,使用叶绿素荧光成像仪(IMAG-PAM;Heinz Walz,Germany)照射检测光(<0.5μmol m-2s-1),测得最小荧光Fo,再照射饱和脉冲光(4000μmol m-2s-1),测得最大荧光Fm。
荧光参数计算方法:PSⅡ最大光化学效率(Fv/Fm)=(Fm-Fo)/Fm。
植株相对电导率测定方法为:将处理结束后的番茄叶片均匀剪成适宜长度的长条(避开主脉),快速称取鲜样3份,每份0.2g,分别置于装有20ml去离子水的刻度离心管中,盖上盖子置于28℃摇床中浸提1.5h-2h。用电导仪测定浸提液电导R1,然后沸水浴加热15min,冷却至室温后摇匀,再次测定浸提液电导R2。相对电导率=R1/R2*100%.
结果显示(图3-6),在低温条件下,与野生型相比,SlNAC100过表达植株表现出更低的电导率以及更高的Fv/Fm和COR基因表达,而SlNAC100CRISPR/Cas9植株反之,这说明SlNAC100基因对提高番茄低温的耐受性具有正调控能力。
需要说明的是,SlNAC100的敲除,可利用CRISPR/Cas9基因编辑方法,还可以采用T-DNA插入、EMS诱变等方法;且载体导入方法不局限于通过农杆菌转化方法,还包括通过花粉管导入作物细胞、愈伤组织、组织或器官中获得的植株。
序列表
<110> 浙江大学
<120> SlNAC100基因在提高番茄低温抗性中的应用
<160> 5
<170> SIPOSequenceListing 1.0
<210> 1
<211> 1020
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<213> 番茄(Solanum lycopersicum)
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atggagaatt attcaggagt tgttaaggat gatgatcaga tggagttacc acctggattt 60
cgatttcatc caactgatga agaattgatc actcattatt tgtctaacaa agttgtggat 120
actaatttcg ttgctattgc tattggtgat gttgatttga acaaagttga accttgggac 180
cttccatgga aggcgaaaat gggggaaaaa gaatggtatt ttttctgtgt gagagacaag 240
aagtatccaa cagggctgag aacaaacagg gcaactgctg cagggtattg gaaagctact 300
ggaaaagaca gagagatttt caggggaaaa tcattggttg gtatgaagaa aactctggtt 360
ttctacaaag ggagagctcc aaaaggtgaa aagacaaatt gggttattca tgaatttaga 420
ttagaaggaa aattgtctct tcaaaatctg ccaaagacag caaagaatga atgggtgatt 480
tgcagagtgt ttcaaaagag cagtggtgga aagaaaatcc acatttcagg gcttttgaaa 540
ctgaattcta atgaaaatga aatggggaat tcatttctgc caccattgac agattctgct 600
actgctactg cttcgaaatc cagccacgtg cactgcttct ccaattttct cactgctcaa 660
aacaactgtt tccctcttct gtcaaatcca atggatagtt accctacaac ttctcttgtt 720
ccaaatacat tttcttgtaa ccaaatagct ccattcacta ctactaataa tccagcttca 780
tttggggttc aagatccttc aattcttcta aggacttcac ttgacagcta tggtctgaat 840
ttcaagaaag aggacatttt taatgtaccc caagaaacag gggtaattag cactgacatg 900
aatactgata tcacctcagt cgtatcaaat cttgaaatga aaagaaggtt tcttgaagat 960
caggtgccat cagcaggtat ggttggatta cagggtcttg attgtctctg gagttgctga 1020
<210> 2
<211> 339
<212> PRT
<213> 番茄(Solanum lycopersicum)
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Pro Pro Gly Phe Arg Phe His Pro Thr Asp Glu Glu Leu Ile Thr His
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Tyr Leu Ser Asn Lys Val Val Asp Thr Asn Phe Val Ala Ile Ala Ile
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Gly Asp Val Asp Leu Asn Lys Val Glu Pro Trp Asp Leu Pro Trp Lys
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Ala Lys Met Gly Glu Lys Glu Trp Tyr Phe Phe Cys Val Arg Asp Lys
65 70 75 80
Lys Tyr Pro Thr Gly Leu Arg Thr Asn Arg Ala Thr Ala Ala Gly Tyr
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Trp Lys Ala Thr Gly Lys Asp Arg Glu Ile Phe Arg Gly Lys Ser Leu
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Val Gly Met Lys Lys Thr Leu Val Phe Tyr Lys Gly Arg Ala Pro Lys
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Gly Glu Lys Thr Asn Trp Val Ile His Glu Phe Arg Leu Glu Gly Lys
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Leu Ser Leu Gln Asn Leu Pro Lys Thr Ala Lys Asn Glu Trp Val Ile
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Cys Arg Val Phe Gln Lys Ser Ser Gly Gly Lys Lys Ile His Ile Ser
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Gly Leu Leu Lys Leu Asn Ser Asn Glu Asn Glu Met Gly Asn Ser Phe
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Leu Pro Pro Leu Thr Asp Ser Ala Thr Ala Thr Ala Ser Lys Ser Ser
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His Val His Cys Phe Ser Asn Phe Leu Thr Ala Gln Asn Asn Cys Phe
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Pro Leu Leu Ser Asn Pro Met Asp Ser Tyr Pro Thr Thr Ser Leu Val
225 230 235 240
Pro Asn Thr Phe Ser Cys Asn Gln Ile Ala Pro Phe Thr Thr Thr Asn
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Asn Pro Ala Ser Phe Gly Val Gln Asp Pro Ser Ile Leu Leu Arg Thr
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Ser Leu Asp Ser Tyr Gly Leu Asn Phe Lys Lys Glu Asp Ile Phe Asn
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Val Pro Gln Glu Thr Gly Val Ile Ser Thr Asp Met Asn Thr Asp Ile
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Thr Ser Val Val Ser Asn Leu Glu Met Lys Arg Arg Phe Leu Glu Asp
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Gln Val Pro Ser Ala Gly Met Val Gly Leu Gln Gly Leu Asp Cys Leu
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Trp Ser Cys
<210> 3
<211> 51
<212> DNA
<213> 人工序列(Artificial Sequence)
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ttacaattac catggggcgc gccatggaga attattcagg agttgttaag g 51
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<212> DNA
<213> 人工序列(Artificial Sequence)
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<210> 5
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
tgttaaggat gatgatcaga 20
Claims (7)
1.番茄SlNAC100基因在提高番茄耐低温能力中的应用,通过基因过表达技术使得所述番茄SlNAC100基因的表达水平上升,所述番茄SlNAC100基因的核苷酸序列:SEQ ID NO:1所示的核苷酸序列。
2.根据权利要求1所述的应用,所述基因过表达技术包括:
提取番茄总RNA,反转录获得cDNA,以cDNA为模板,F和R为引物,扩增SlNAC100基因,将扩增产物构建到植物过表达载体上,其中F引物和R引物的核苷酸序列分别如SEQ ID NO:3和4所示;
将植物过表达载体导入宿主细胞中,再利用其侵染目的植株,筛选阳性转基因植株,获得耐低温的转基因番茄植株。
3.根据权利要求2所述的应用,所述植物过表达载体为具有35S启动子的表达载体。
4.根据权利要求2所述的应用,所述的宿主细胞为农杆菌细胞。
5.根据权利要求4所述的应用,所述的农杆菌细胞为EHA105。
6.根据权利要求5所述的应用,所述植物过表达载体为载体pFGC1008-HA或载体pCAMBIA1301。
7.番茄SlNAC100蛋白在提高番茄耐低温能力中的应用,通过基因过表达技术使得所述番茄SlNAC100蛋白的表达水平上升,所述番茄SlNAC100蛋白的氨基酸序列:SEQ ID NO:2所示的氨基酸序列。
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