CN113278630B - 一种改良桑树白藜芦醇生物合成的转录因子基因MaMYB14及其应用 - Google Patents
一种改良桑树白藜芦醇生物合成的转录因子基因MaMYB14及其应用 Download PDFInfo
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- CN113278630B CN113278630B CN202110577231.9A CN202110577231A CN113278630B CN 113278630 B CN113278630 B CN 113278630B CN 202110577231 A CN202110577231 A CN 202110577231A CN 113278630 B CN113278630 B CN 113278630B
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Abstract
本发明公开了一种改良桑树白藜芦醇生物合成的转录因子基因MaMYB14及其应用,所述转录因子基因MaMYB14的核苷酸序列如SEQ ID No.1所示,该基因全长831bp,编码276个氨基酸。亚细胞定位研究表明,该基因蛋白主要定位于细胞核中;酵母激活实验显示该蛋白具有转录激活作用;启动子活性实验表明MaMYB14显著激活ProMaSTS3::LUC的表达,表明MaMYB14能促进桑树白藜芦醇合酶基因MaSTS3的转录,蛋白互作表明MaMYB14能与E3类泛素化酶SIZ1互作,本发明能够应用于改良植物中白藜芦醇的生物合成。
Description
技术领域
本发明涉及功能基因和植物基因育种领域,具体涉及一种改良桑树白藜芦醇生物合成的转录因子基因MaMYB14及其应用。
背景技术
白藜芦醇类物质的生物活性已被广泛认可,具有抗氧化、抗衰老、抑制肿瘤增殖、抑制α-葡萄糖苷酶活性等生物活性。桑树富含白藜芦醇、氧化白藜芦醇和桑皮苷A等芪类物质,前期实验表明在干根皮中桑皮苷A达到9.9mg/g。MaSTS3是桑树白藜芦醇生物合成过程中的关键基因,其催化由4-香豆酰CoA与丙二酰CoA缩合成的中间产物发生分子重排,生成白藜芦醇。
MYB是广泛存在于动物、植物和酵母中的一类转录因子,也是植物中最大的家族蛋白之一,其在植物苯丙氨酸代谢途径中具有重要的调控作用。MYB蛋白上的存在若干重复结构单元,每个单元由50-53个残基组成,编码三个α螺旋,其中第二和第三个螺旋形成螺旋-回转-螺旋结构,与DNA双螺旋的大沟结合,进而调控基因的转录。根据重复结构的数量,MYB可分为1R(R1/2,R3-MYB),2R(R2R3-MYB),3R(R1R2R3-MYB)和4R(4个R1/R2-like)四种类型。不同类型的MYB在植物中参与的生理活动不同,如调控花青素合成、原花青素、黄酮醇等植物次生代谢产物的。
MaSIZ1为类泛素化蛋白SmallUbiquitin-likeModifier(SUMO),其能通过类泛素化作用在受体蛋白上添加泛素,从而改变蛋白的稳定性。
发明内容
本发明的目的在于提供一种改良桑树白藜芦醇生物合成的转录因子基因MaMYB14及其应用,其研究表明桑树MaSIZ1蛋白与MaMYB14互作,通过类泛素化途径改变MaMYB14的稳定性,因此桑树MaMYB14可应用于桑树白藜芦醇生物合成的调控,改良桑品质。
本发明通过以下技术方案来实现上述目的:
一种转录因子基因MaMYB14,所述转录因子基因MaMYB14的核苷酸序列如SEQ IDNo.1所示。
本发明还提供了一种转录因子基因MaMYB14编码的蛋白质,其氨基酸序列如SEQID No.2所示。
本发明还提供了用于克隆获取转录因子基因MaMYB14的引物对,该引物对的引物序列为:
MaMYB14-F:ATGGTAAGAGCACCTTGCTGT;
MaMYB14-R:TCACAATTCCGCTAGTAATTC。
本发明还提供了沉默转录因子基因MaMYB14的沉默载体,所述沉默载体为包含MaMYB14部分特异性序列的VIGS沉默载体,命名为TRV2-ΔMaMYB14,扩增所述MaMYB14部分特异性序列的引物序列为:
VIGS-MYB-F:atcgacgacaagaccctgcagTCCTCAATTAGTGATATCTC;
VIRS-MYB-R:ctgaggagaagagccctgcagCAATTCCGCTAGTAATTCT;
其中,小写字母为载体同源臂。
本发明还提供了一种转录因子基因MaMYB14在改良桑树白藜芦醇生物合成中的应用。
进一步改进在于,所述改良桑树白藜芦醇生物合成指的是调控桑树白藜芦醇生物合成过程中MaSTS基因表达。
进一步改进在于,所述转录因子基因MaMYB14通过激活MaSTS3基因表达、促进MaSTS3基因的转录、与E3类泛素化酶SIZ1互作实现MaSTS基因表达的调控。
本发明的有益效果在于:本发明提供了一个新的MaSTS3的调控基因MaMYB14,该基因全长831bp,编码276个氨基酸。亚细胞定位研究表明,该蛋白主要定位于细胞核中;酵母激活实验显示该蛋白具有转录激活作用;启动子活性实验表明MaMYB14显著激活ProMaSTS3::LUC的表达,表明MaMYB14能促进桑树白藜芦醇合酶基因MaSTS3的转录,蛋白互作表明MaMYB14能与E3类泛素化酶SIZ1互作,本发明能够应用于改良植物中白藜芦醇的生物合成。
附图说明
图1(A-G)为MaMYB14调控的桑树MaSTS3基因表达分析图;
图2(A-C)为MaMYB14转录活性分析图;
图3(A-E)为MaMYB14和MaSIZ1亚细胞定位分析和蛋白互作分析图;
图4(A-D)为MaMYB14和MaSIZ1互作的意义分析图。
具体实施方式
下面结合附图对本申请作进一步详细描述,有必要在此指出的是,以下具体实施方式只用于对本申请进行进一步的说明,不能理解为对本申请保护范围的限制,该领域的技术人员可以根据上述申请内容对本申请作出一些非本质的改进和调整。
实施例
一、MaMYB14的克隆获取
取桑树叶片并提取总RNA,反转录成cDNA,并利用特异性扩增引物进行PCR扩增,克隆MaMYB14所用引物如下:
MaMYB14-F:ATGGTAAGAGCACCTTGCTGT;
MaMYB14-R::TCACAATTCCGCTAGTAATTC。
所得MaMYB14的全长CDS核苷酸序列如SEQ ID No.1所示,编码的氨基酸序列如SEQID No.2所示。
通过NCBI数据库比对MaMYB14特异性序列,构建病毒沉默载体TRV2-ΔMaMYB14,引物序列如下(小写字母为载体同源臂):
VIGS-MYB-F:atcgacgacaagaccctgcagTCCTCAATTAGTGATATCTC;
VIRS-MYB-R:ctgaggagaagagccctgcagCAATTCCGCTAGTAATTCT。
二、基因功能和表达分析
对所构建载体测序,转化农杆菌,挑取单克隆,利用LB液体培养基(含利福平和卡那霉素),220r/min培养过夜,离心收集菌体,利用侵染液重悬菌体,利用注射器将菌液注射进桑叶中。将桑苗放置于16h/8h光周期温室中培养,18天后提取RNA做转录组分析。如图1所示:(A)VIGS-MaMYB144基因组转录变化分析;(B)qRT-PCR检测MaMYB144表达量变化;(C)差异基因GO富集分析;(D)KEGG通路富集分析;(E)差异基因MaSTS3表达验证;(F)差异基因MaCHS1表达验证;(G)差异表达基因MaSTS3和MaCHS1转拟南芥突变体tt4功能分析。
结果表明,沉默MaMYB14导致桑树中313个基因显著发生变化,其中上调179,下调134,MaMYB14表达量显著下调(图1A,B)。差异基因GO富集和KEGG通路富集分析表明MaMYB14与黄酮类物质合成有关(图1C,D),且qRT-PCR验证表明MaSTS3和MaCHS1表达量显著下调(图1E,F)。因白藜芦醇合酶基因STS与查尔酮合酶CHS存在高度的同源性,部分数据库将STS基因注释成CHS。本实验对富集于柚皮素-查尔酮酶活性和黄酮类物质合成的显著差异基因进行功能验证,结果表明在以上通路中的表达差异的CHS基因不能回补chs突变体表型,结合申请人前期发表论文中的酶活实验【Characterization of Stilbene Synthase Genes inMulberry(Morus atropurpurea)and Metabolic Engineering for the Production ofResveratrol in Escherichia coli,Journal of agricultural and food chemistry,2017,65,1659-1668】,表明该CHS基因实际编码STS(图1G),命名为MaSTS3。
三、蛋白定位和互作分析
将MaMYB14连接于pGBKT7载体,开展酵母转录激活实验。如图2所示,结果表明MaMYB14具有转录激活活性(图2A)。将MaSTS3启动子连接于pGreenII-0800-luc载体上构建ProSTS3::luc,与35S::MaMYB14共转烟草,结果显示35S::MaMYB14能显著提高LUC酶活,说明MaMYB14具有激活MaSTS3基因表达的功能(图2B,C)。
构建35S::MaMYB14-GFP超表达载体,通过农杆菌注射桑树叶片,培养3天后收集叶片,液氮低温研磨,进行蛋白提取,并利用GFP beads富集MaMYB14蛋白复合体,对所提取蛋白进行SDS-PAGE电泳,待溴酚蓝指示剂刚跑出浓缩胶后切胶做质谱分析。对质谱结果进行肽段分析,发现MaSIZ1部分片段,该蛋白为E3类泛素蛋白连接酶SIZ1(E3 SUMO-proteinligase SIZ1)。
构建亚细胞定位载体35S::MaSIZ1-GFP,转化农杆菌并注射烟草。如图3所示,结果表明MaMYB14和MaSIZ1都定位于细胞核中(图3A)。构建酵母双杂交(Y2H)载体AD-MaMYB14和BK-MaSIZ1进行验证,结果表明MaMYB14和MaSIZ1能相互作用(图3B)。同理,将MaMYB14和MaSIZ1分别连接于35S::YNE和35S::YCE载体上,转化农杆菌后注射烟草(方法同上),结果显示YNE-MaMYB14能与YCE-MaSIZ1互作(图3C)。同理,Nluc-MaMYB14和Cluc-MaSIZ1也能发生相互作用(图3D)。此外,蛋白pull-down实验表明MaMYB14能与MaSIZ1直接作用(图3E)。MaSIZ1具有类泛素化功能,其通过在目的蛋白上添加泛素,使蛋白更稳定。
四、MaMYB14和MaSIZ1互作的意义分析
如图4所示,为MaMYB14和MaSIZ1互作的意义分析图,结果表明,过表达MaSIZ1使MaMYB14-GFP蛋白更稳定(图4A,B);过表达MaSIZ1促进MaMYB14-Luc蛋白稳定(图4C,D)。
以上所述实施例仅表达了本发明的几种实施方式,其描述较为具体和详细,但并不能因此而理解为对本发明专利范围的限制。应当指出的是,对于本领域的普通技术人员来说,在不脱离本发明构思的前提下,还可以做出若干变形和改进,这些都属于本发明的保护范围。
序列表
<110> 安徽农业大学
<120> 一种改良桑树白藜芦醇生物合成的转录因子基因MaMYB14及其应用
<141> 2021-05-26
<160> 2
<170> SIPOSequenceListing 1.0
<210> 1
<211> 831
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<213> 桑树(Morus alba)
<400> 1
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gaagatcaga ttctcatctc ttacattcaa agatatggcc atgccaattg gcgtgcactc 120
cccaaacaag cgggtttatt aaggtgtgga aagagttgca gattgcgatg gataaactac 180
ttgagaccgg acattaaacg aggaaacttt agcactgaag aagaggagac catcattaag 240
ctacacgaaa tgttaggaaa taggtcggca attgcagcga gactgccggg acgaacagac 300
aatgaaataa aaaatgtgtg gcatacccat ttgaagaaaa ggctcacaca atccaatcat 360
aatgtcgctc caaaaaatac aaaacgacag gatcatgaag gagcagaagt agtacgtggc 420
atgaattctt caaaatacga cagcatggag aataggccaa tcacttcgcc ccaacagagt 480
tcctcaatta gtgatatctc ctcagcaatc agcactgatc gtgataacaa taacaacaac 540
aaccacacat tcatgaaagc cgattctccc gggaaattgt ccgaaacggc cgacgagagt 600
ttctggtcgg aagttttcac aaccgatgag agtaattatt ctgaggtgct ggatgatttt 660
tcagcaatta gtagtaaccc aatagttcaa tttccatcct ctcctctagt gtccagcgaa 720
ccggtagtcc atgcctttcg ttccaatatt catggtgatg atatggattt ttggcacaat 780
ttcttcacaa aagctgggga tgatctacaa gaattactag cggaattgtg a 831
<210> 2
<211> 276
<212> PRT
<213> 桑树(Morus alba)
<400> 2
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Cys Gly Lys Ser Cys Arg Leu Arg Trp Ile Asn Tyr Leu Arg Pro Asp
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Ile Lys Arg Gly Asn Phe Ser Thr Glu Glu Glu Glu Thr Ile Ile Lys
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Leu His Glu Met Leu Gly Asn Arg Ser Ala Ile Ala Ala Arg Leu Pro
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Gly Arg Thr Asp Asn Glu Ile Lys Asn Val Trp His Thr His Leu Lys
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Lys Arg Leu Thr Gln Ser Asn His Asn Val Ala Pro Lys Asn Thr Lys
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Arg Gln Asp His Glu Gly Ala Glu Val Val Arg Gly Met Asn Ser Ser
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Lys Tyr Asp Ser Met Glu Asn Arg Pro Ile Thr Ser Pro Gln Gln Ser
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Ser Ser Ile Ser Asp Ile Ser Ser Ala Ile Ser Thr Asp Arg Asp Asn
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Asn Asn Asn Asn Asn His Thr Phe Met Lys Ala Asp Ser Pro Gly Lys
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Pro Val Val His Ala Phe Arg Ser Asn Ile His Gly Asp Asp Met Asp
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Claims (7)
1.一种转录因子基因MaMYB14,其特征在于:所述转录因子基因MaMYB14的核苷酸序列如SEQ ID No.1所示。
2.一种如权利要求1所述转录因子基因MaMYB14编码的蛋白质,其特征在于:其氨基酸序列如SEQ ID No.2所示。
3.用于克隆获取如权利要求1所述转录因子基因MaMYB14的引物对,其特征在于:该引物对的引物序列为:
MaMYB14-F:ATGGTAAGAGCACCTTGCTGT;
MaMYB14-R:TCACAATTCCGCTAGTAATTC。
4.沉默转录因子基因MaMYB14的沉默载体,其特征在于:所述沉默载体为包含MaMYB14部分特异性序列的VIGS沉默载体,命名为TRV2-ΔMaMYB14,扩增所述MaMYB14部分特异性序列的引物序列为:
VIGS-MYB-F:atcgacgacaagaccctgcagTCCTCAATTAGTGATATCTC;
VIRS-MYB-R:ctgaggagaagagccctgcagCAATTCCGCTAGTAATTCT;
其中,小写字母为载体同源臂。
5.一种如权利要求1所述转录因子基因MaMYB14在改良桑树白藜芦醇生物合成中的应用。
6.根据权利要求5所述的应用,其特征在于:所述改良桑树白藜芦醇生物合成指的是调控桑树白藜芦醇生物合成过程中MaSTS基因表达。
7.根据权利要求6所述的应用,其特征在于:所述转录因子基因MaMYB14通过激活MaSTS3基因表达、促进MaSTS3基因的转录、与E3类泛素化酶SIZ1互作实现MaSTS基因表达的调控。
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