CN109486838B - 一种调控植物类黄酮合成的转录因子基因及其用途 - Google Patents
一种调控植物类黄酮合成的转录因子基因及其用途 Download PDFInfo
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Abstract
本发明公开了一种调控植物类黄酮合成的转录因子基因，该转录因子基因即为从豆科植物蒺藜苜蓿中鉴定的MtbZIP10基因，其基因序列如SEQ ID NO.1所示。并且，本发明公开了该基因的功能及其应用。本发明的优点在于，本发明从豆科植物蒺藜苜蓿中克隆出可以正调控类黄酮化合物合成的转录因子基因，并对其功能进行了系统鉴定，发现了该基因可以激活类黄酮生物合成途径的多个基因，提高类黄酮化合物的含量。
Description
技术领域
本发明属于基因克隆技术领域，具体为一种调控植物类黄酮合成的转录因子基因及其用途。
背景技术
类黄酮化合物是广泛分布于植物的根、茎、叶、花、果实和种皮中的一大类重要的次生代谢物产物，包括类黄酮、异黄酮、黄酮醇、花青素和原花青素等。其中花青素具有保护植物免受生物侵害和吸引昆虫授粉的功能，花青素还可以响应生物与非生物胁迫、清除氧自由基以及保护植物免受高密度光照的伤害。同时，花青素在抗癌、抗病和抗氧化方面具有一定的保健功能。类黄酮化合物中的另一种类原花青素不仅具有保护植物免受食草动物和病原菌侵害机制方面的作用，而且适量原花青素还可以在牧草中有效防止牲畜牧草胀气并提高反刍牲畜氮营养。同时，原花青素对人体健康有益，因此类黄酮的代谢调控机制是植物次生代谢领域的研究热点。
虽然调控类黄酮生物合成的转录因子如MYB、bHLH和WDR等类的转录因子基因已被报道，但是其它类型的转录因子研究相对较少。bZIP转录因子(basicregion/leucinezippermotif,bZIP)在人、动物、植物、微生物和昆虫中均有发现，是分布最广泛、最保守的一类转录因子。植物bZIP转录因子还具有组织表达特异性，许多植物的bZIP转录因子在根中组成型表达，在茎和叶片的表达量很少或没有表达。植物bZIP转录因子参与多种生物学过程，包括种子成熟、发芽、花发育、植物衰老、光形态建成等。虽然个别其它植物中的bZIP家族的转录因子可以参与调控花青素的生物合成，但是在蒺藜苜蓿中，目前还没有任何有关bZIP转录因子调控类黄酮合成途径的相关研究，限制了对该模式植物类黄酮途径的研究和应用，所以克隆和鉴定具有调控类黄酮合成的新功能、并可在植物植物中提高类黄酮含量的bZIP类转录因子基因具有重要的理论和应用价值。
发明内容
针对现有研究中类黄酮生物合成途径中bZIP类转录因子基因不足、难以大量提高类黄酮含量的问题，本发明提供了一种调控植物类黄酮合成的转录因子基因及其用途，该基因是一个新的基因，在蒺藜苜蓿中还未发现调控类黄酮(花青素和原花青素)合成的bZIP家族转录因子，因此，研究蒺藜苜蓿中调控类黄酮化合物的生物合成的bZIP转录因子十分必要的问题。
为了实现上述目的，本发明提供的技术方案为，本发明提供的一种bZIP转录因子基因，该bZIP转录因子基因即为MtbZIP10，其基因序列如SEQ ID NO.1所示。本发明所发现的MtbZIP10基因是从蒺藜苜蓿中克隆出来的，是一种全新的植物bZIP转录因子基因，丰富了类黄酮bZIP转录因子的种类，同时由于bZIP转录因子可以正调控功类黄酮，特别是花青素和原花青素的功能，所以可以提高植物在抗性和营养品质中的应用。
进一步的，所述该bZIP转录因子基因编码的蛋白质，其氨基酸序列如SEQ ID NO.2所示。
进一步的，所述bZIP转录因子基因的用途为正调控类黄酮的生物合成途径，特别是CHS,DFR,ANS和MATE1等类黄酮途径的关键基因，证明MtbZIP10可以通过调控这些基因进而发挥调控功能，这是蒺藜苜蓿中发现的第一个调控类黄酮途径的bZIP类型的转录因子基因。
进一步的，所述MtbZIP10基因在蒺藜苜蓿中的表达被抑制时，地上部分的花青素和种子中的原花色素减少。
进一步的，所述MtbZIP10基因在蒺藜苜蓿的毛状根中过量表达时，毛状根的花青素含量相对于未转化的对照显著增加，说明过量表达该基因具有提高花青素积累的所用。
本发明采用上述技术方案，具有以下有益效果，本发明从蒺藜苜蓿中克隆出类黄酮异戊烯基转移酶基因，并对其功能进行了系统鉴定，发现了该基因发生突变可以影响花青素和原花色素的含量，该基因的过量表达可以提高花青素的含量。
附图说明
图1 MtbZIP10与其它植物bZIP转录因子氨基酸序列的多重比对分析。Consensus表示序列完全一致的氨基酸，用黑色和灰色来强调保守的氨基酸：黑色字体深灰背景代表100％相同；黑色字体灰白背景代表50％相同；黑色字体白背景代表相似度在50％以下。
图2植物bZIP转录因子的进化树分析。用DNAMAN构建进化树，MtbZIP10后标注星号。基于1000次重复的bootstrap支持值(％)估算节点的可信度。
图3 MtbZIP10的组织表达谱。MtbZIP10基因在根、茎、叶、花、豆荚和授粉后10天、12天、16天、20天、24天、36天种子中的表达情况。图示数据为平均值±标准误差，每个实验包括三个生物学重复。
图4 R108和NF2620中花青素合成调控关键基因的相对表达量。ANS：花青素合成酶；CHS：查尔酮合酶；CHI：查尔酮异构酶；DFR 1/2：二氢类黄酮醇还原酶；UGT72L1：糖基转移酶；TT8：TT8转录因子；WD40：WD40重复蛋白；MATE1：MATE1转运子。图示数据为平均值±标准误差，每个实验包括三个生物学重复。
图5使用PCR的方法确认植物表达载体的构建。其中左侧为DNA分子量标记；右侧为含有MtbZIP10基因的农杆菌阳性克隆的PCR检测。
图6转基因毛状根中花青素含量测定。A,突变体NF260背景下的毛状根及其阴性对照中的花青素含量。B,野生型A17背景下的毛状根及其阴性对照中的花青素含量.
图7脱落酸和光照对MtbZIP10表达的影响。A，C：在指定的时间间隔内以50μM ABA处理R108幼苗，通过半定量PCR(A)和实时荧光定量PCR(C)检验MtbZIP10的表达水平。未经ABA处理的组分为1。B，D：将R108幼苗在常温下黑暗处理3天，再分别以1、3、6、9、12小时光照处理幼苗，通过半定量PCR(A)和实时荧光定量PCR(C)检验MtbZIP10的表达水平。黑暗处理的组分为1。
具体实施方式
下面将结合本发明的实施例，对本发明实施例中的技术方案进行清楚、完整地描述，显然，所描述的实施例仅仅是本发明一部分实施例，而不是全部的实施例。基于本发明中的实施例，本领域普通技术人员在没有做出创造性劳动前提下所获得的所有其它实施例，都属于本发明保护的范围。
一、主要实验材料和方法
1.1CTAB法提取蒺藜苜蓿基因组DNA
1)取约100mg蒺藜苜蓿组织材料，与钢珠一起置于2mL离心管中；加入液氮不断摇晃，直至组织材料被充分打磨成粉末。
2)加入500μLCTAB提取液(65℃预热)，涡旋振荡充分混匀。
3)混合液65℃水浴30min，期间不断摇晃。
4)待混合液降至室温，加入等体积氯仿，颠倒混匀。
5)在室温情况下12000rpm离心10min。
6)离心后混合液分层，小心吸取上清液置于1.5mL离心管中。
7)加入等体积(约500μL)的异丙醇，颠倒混匀，室温条件下静置30min。
8)4℃12000rpm离心10min。
9)弃上层溶液，用70％乙醇溶液洗涤DNA沉淀。
10)待乙醇蒸发后，加入30-50μL无菌水溶解DNA沉淀。-20℃保存。
1.2TRNzol-A+法提取蒺藜苜蓿RNA
1)取约200mg蒺藜苜蓿组织材料置于于1.5mL RNase-free离心管中，液氮冷冻，并用灭菌冷冻的玻棒迅速研磨至粉状。
2)加入1mL TRNzol-A+提取液，涡旋振荡混匀混合液，室温静置5min。
3)将混合液在4℃条件下12000rmp离心10min。
4)吸取上清液至1.5mL RNase-free离心管中。
5)加入200μL RNase-free氯仿，反复颠倒混匀后室温静置5min。
6)将混合液在4℃条件下12000rmp离心10min。
7)吸取上清液至1.5mL RNase-free新离心管中，加入等体积的RNase-free异丙醇，颠倒混匀后室温放置10min。
8)将混合液在4℃条件下12000rmp离心10min。
9)弃去上层液体，加入75％DEPC处理过的乙醇溶液反复洗涤RNA沉淀。
10)在4℃条件下7500rmp离心5min，完全弃去液体。
11)待乙醇完全蒸发，加入30-50μL DEPC水，溶解RNA沉淀。-80℃保存。
1.3荧光定量qRT-PCR
根据Takara生物医学技术(北京)公司提供的SYBR Premix ExTaqII(TliR NaseHPlus)说明书进行荧光定量qRT-PCR。
根据下表在冰上配制反应液，进行荧光定量qRT-PCR实验。
qRT-PCR反应体系
1.4MtbZIP10基因的克隆和载体构建
根据Takara生物医学技术(北京)公司提供的Prime STAR GXL Premix高保真DNA聚合酶说明书进行PCR反应。根据天根生化科技公司提供的pEASY-T1Cloning Kit说明书进行连接按照表配置反应体系，25℃反应20min。在基因片段两端分别加A，后将pCXSN植物表达载体用XcmΙ限制性内切酶酶切。将各自酶切后的片段进行连接和转化。
1.5转化蒺藜苜蓿毛状根
1)将活化好的MtbZIP10-pCXSN/Arqual I发根农杆菌均匀涂布于含Kan和Strep抗生素的LB固体培养基上，28℃倒置培养24h。
2)待蒺藜苜蓿的根长至2cm左右时剪掉根尖，将伤口在Aqual I菌膜上轻轻蘸取，然后排列于不含抗生素的F培养基上，光下培养。
3)待新根长出后，换到含有Hyg抗性的B5固体培养基上，光下继续培养。
4)当毛状根生长旺盛时，提取转基因毛状根的DNA，筛选转基因阳性毛状根。
5)观察阳性毛状根表型，并对高表达株系的代谢物含量进行检测。
1.6测定花青素的含量
1)取同一生长环境条件下的蒺藜苜蓿地上部分(茎和叶)新鲜组织样品。
2)向组织材料中加入液氮，研磨后冷冻干燥24h。准确称取10mg，并加入5倍体积的甲醇(含体积比为0.1％的盐酸)，超声30min，放置于4℃条件下12h。
3)再次超声30min，10000rmp离心5min，取上清液于新的离心管中。
4)加入750μL水和750μL氯仿，振荡混匀后10000rmp离心5min，取上清液于新的离心管中。
5)用紫外分光光度计在530nm处检测。以矢车菊素-3-O-葡萄糖的摩尔光吸收作为标准品估算花青素的含量。
1.7测定总类黄酮的含量
1)取同一生长环境下的蒺藜苜蓿新鲜组织样品(包括植株和种子)，加入液氮研磨至粉末状。
2)准确称取50mg粉末，加入1000μL 80％甲醇，超声30min。
3)12000rpm离心20min，取上清液至新的离心管中。
4)取200μL上清液，加入800μL ddH2O和60μL 5％NaNO2，混合均匀后静置5min。
5)加入60μL 10％AlCl3，混匀后静置10min。
6)加入400μL 1mol/L NaOH溶液，再加入ddH2O补至2mL(约480μL)。
7)使用紫外分光光度计于OD 510nm读数。水作为空白对照。以山奈酚标准曲线估算总类黄酮的含量。
二、主要结果
2.1MtbZIP10的克隆和生物信息学分析
本发明对MtbZIP10基因完整ORF进行克隆及测序分析，发现MtbZIP10基因全长为6542bp，由两个完全相同的开放阅读框串联重复组成，含有15个外显子和16个内含子。MtbZIP10基因编码一条含有709个氨基酸的蛋白序列；其转录本长为2127bp。MtbZIP10转录因子的氨基酸序列和另外6个已经验证与花青素和原花青素合成相关的bZIP转录因子的氨基酸序列进行多重序列比对，发现这7个属于不同亚家族的bZIP转录因子在保守结构域都具有两个保守的氨基酸位点，很可能与调控类黄酮生物合成有关(图1)。将MtbZIP10转录因子的氨基酸序列和已知功能的来自拟南芥、大豆和水稻等物种的27个bZIP转录因子的氨基酸序列进行系统进化树分析，发现MtbZIP10转录因子与调控花青素和原花青素合成的bZIP转录因子聚簇(图2)。进一步从系统进化的角度证明了MtbZIP10转录因子很可能参与调控类黄酮代谢途径。
2.2突变体的鉴定
本发明对得到的MtZIP10发生突变的一个突变体NF2620进行表型观察，发现与野生型R108相比，该突变体的幼苗下胚轴部分和植株茎部缺乏花青素积累，而叶的近轴面和远轴面因花青素积累而形成的红色花环以及斑点缺失。此外，与R108种皮由于原花青素积累而表现出的黄棕色不同，NF2620突变体部分种子种皮颜色呈白色。上述结果表明了突变体NF2620由于MtZIP10基因发生突变而造成了花青素和原花青素含量大幅减少。
2.3MtbZIP10的组织表达谱
本发明检测了MtbZIP10基因在蒺藜苜蓿中的表达情况，分别采集了了蒺藜苜蓿根、茎、叶、花和开花后10天、12天、16天、20天、24天、36天收获的种子，提取其RNA。RNA反转录后得到cDNA，以actin为内参进行荧光定量qRT-PCR实验；以野生型蒺藜苜蓿A17各组织中MtbZIP10基因的表达水平为1，检测MtbZIP10基因在突变体NF2620不同组织和种子不同发育时期的相对表达水平。
结果表明，在蒺藜苜蓿的各个部位都能检测到MtbZIP10的表达，但表达量均较低。其中，在茎和12天种子中的表达水平最高(图3)。花青素主要积累于蒺藜苜蓿的茎和叶中，原花青素在种子种皮中积累量最高，即MtbZIP10基因的表达情况与花青素和原花青素积累部位相符。
2.4突变体NF2620中类黄酮合成调控关键基因的表达分析
本发明通过qRT-PCR对野生型和突变体蒺藜苜蓿50d幼苗叶片中参与类黄酮合成途径的几个关键结构基因和转录因子的表达水平进行了定量分析；包括CHS、CHI、ANS、DFR1、DFR2、UGT72L1、TT8、WD40和MATE1。
与野生型相比，突变体中CHS、DFR1、DFR2和ANS基因不表达(图4A，C-E)；UGT72L1、TT8和MATE1的表达量都显著降低(图4F-G、I)，这说明突变体NF2620中与类黄酮生物合成相关的关键结构基因和调控基因的表达量的显著降低造成了突变体中花青素和原花青素的缺乏。
2.5MtbZIP10转化蒺藜苜蓿毛状根
为了进一步证明MtbZIP10的功能，将烟草花叶病毒CaMV 35S启动子驱动的MtbZIP10基因连接到植物表达pCXSN上；再将重组质粒转入发根农杆菌Arqual I中，使用PCR的方法确认该基因在农杆菌中成功表达(图5)。将带有MtbZIP10-pCXSN的发根农杆菌Arqual I转化到突变体NF2620和野生型蒺藜苜蓿A17的毛状根中，获得了转MtbZIP10基因的NF2620毛状根和过量表达MtbZIP10基因的A17毛状根。同时将空载也转化到突变体NF2620和野生型A17毛状根中作为阴性对照，观察并检测阳性基因互补毛状根株系和阳性基因过量表达毛状根株系中花青素的积累情况。
本发明发现转MtbZIP10基因的NF2620毛状根和过量表达MtbZIP10基因的A17毛状根的颜色均比阴性对照深，转基因毛状根中花青素积累多于阴性对照(毛状根生长旺盛期前观察)。选取阳性毛状根，称取10mg提取其花青素进行含量测定。结果表明，转MtbZIP10基因的NF2620毛状根中的花青素含量与阴性对照基本持平，过量表达MtbZIP10基因的A17毛状根中的花青素含量显著高于阴性对照。说明MtbZIP10基因在野生型蒺藜苜蓿中过量表达能够促进花青素积累，同时也进一步说明MtbZIP10转录因子有调控花青素生物合成的功能(图6)。
2.6MtbZIP10在光照和脱落酸诱导条件下的表达情况
本发明还检测了MtbZIP10基因是否受脱落酸诱导，分别在0h、1h、3h、6h、9h和12h对野生型蒺藜苜蓿外援施加100mg/L的脱落酸，利用半定量PCR和荧光定量PCR分别检测MtbZIP10基因表达情况(图7)。结果显示，MtbZIP10基因的表达与外援施加脱落酸没有明显相关性(图7)。本发明还同时对野生型蒺藜苜蓿进行了光照处理。半定量PCR和荧光定量PCR结果表明：随着光照时间的增加MtbZIP10基因的表达量升高(图7)，说明MtbZIP10参与光介导的花青素积累。
三、最终结论
1.蒺藜苜蓿是豆科模式植物，也是优良牧草—紫花苜蓿的近缘种，具有重要的研究价值。本发明首次在蒺藜苜蓿中证明了bZIP转录因子MtbZIP10基因编码一个bZIP型的转录因子，MtbZIP10基因在蒺藜苜蓿中的表达水平与花青素和缩合单宁等类黄酮的积累水平相关，MtbZIP10是调控蒺藜苜蓿类黄酮生物合成的关键转录因子。
2.以花青素和缩合单宁为代表的类黄酮化合物广泛存在于植物中，不仅具有保护植物免受生物与非生物侵害和吸引昆虫授粉的功能，还具有抗癌、抗病和抗氧化方面的生物学特性。蒺藜苜蓿MtbZIP10基因调控类黄酮途径多个基因的表达，其过量表达可以提高黄酮类化合物的含量，为植物类黄酮的代谢工程提供了候选基因和有益的参考，同时为改良苜蓿品质奠定了分子基础。
本实施例中无特殊说明的操作手法均为现有技术，故不在此过多解释。
以上所述仅为本发明的优选实施例而已，并不用于限制本发明，对于本领域的技术人员来说，本发明可以有各种更改和变化。凡在本发明的精神和原则之内，所作的任何修改、等同替换、改进等，均应包含在本发明的保护范围之内。
序列表
<110> 中国农业科学院北京畜牧兽医研究所
<120> 一种调控植物类黄酮合成的转录因子基因及其用途
<160> 2
<170> SIPOSequenceListing 1.0
<210> 1
<211> 2127
<212> DNA
<213> MtbZIP10基因(Medicago truncatula)
<400> 1
atggcagata acagtcaaaa gacagaagat tttgacactg atgataaaaa taatcaatgt 60
ttatcaaccg tgtcttggtg caatggtgtt ggaaatgagg ctcttgtagt tgttgaatcc 120
aaggatcaat gcaagactaa gggtcaatct gatgagcaca agactcttcg tcggctgatg 180
cagaatcgcg aggctgcaag gaaaagtagg ttaaggaaaa aggcttatgt gcaacaattg 240
gagaacagtc gacttaggct tgctcaaata gaacatgagc ttcaacaagt acgtcaacag 300
gtccaacaac ttaaaaagat ttttcatttg gtaaaattgg tttgttcggt agcatttgat 360
atggactatg ctcgttgggt agacgagcat caacgactaa tcaatgacat aagatcagct 420
ataaattctc aaatgggtga taatgaactg catcttcttg ttgatggtgt catggtacat 480
tatgatgaat tatacaagtt gaagagcata ggtgcaaagg ctgatgtatt tcacatactt 540
tctgggttgt ggaagacacc tgcagaaaga tgtttcatgt ggcttggtgg attccgttca 600
tccgaacttc tcaagataat tagaaaccac ctcgaggcgt taacggatca gcaattgatg 660
gcgatcttca atctgcagca ttcttgtcaa caggctgaag atgcattatc tcaaggaatg 720
gaaggtttgc aacaatctct ttcagagaca ctttcctcca cgtccactgg atctggaaat 780
gttgttgagt atatgggtca aatggctctt tcaatggcca agctttccac actggagact 840
ttcattcatc aggcagatat cttgaggcaa caaacactgc aacagatgcg tcgaattttg 900
actgcgcacc aagctgctcg tgctctcctt gtcataaatg atttcatttc acgaatcaga 960
gctcttaatt cattatggca tgtgtttggt gcaaagtcta acaatgtcac tgttgttgca 1020
agtaacttgc agtatggaac attcaacagt aacattggtt cagcttcttc ttcaggcatg 1080
gcagataaca gtcaaaagac agaagatttt gacactgatg ataaaaataa tcaaagtggg 1140
aatgtcctta tttttctgtg tttatcaacc gtgtcttggt gcaatggtgt tggaaatgag 1200
gctcttgtag ttgttgaatc caaggatcaa tgcaagacta agggtcaatc tgatgagcac 1260
aagactcttc gtcggctgat gcagaatcgc gaggctgcaa ggaaaagtag gttaaggaaa 1320
aaggcttatg tgcaacaatt ggagaacagt cgacttaggc ttgctcaaat agaacatgag 1380
cttcaacaag tacgtcaaca gggtacattt gttgcacctg gagttacagc ggatcatggt 1440
cattcaattg ttggaaacag taatgcaggt tcggtagcat ttgatatgga ctatgctcgt 1500
tgggtagacg agcatcaacg actaatcaat gacataagat cagctataaa ttctcaaatg 1560
ggtgataatg aactgcatct tcttgttgat ggtgtcatgg tacattatga tgaattatac 1620
aagttgaaga gcataggtgc aaaggctgat gtatttcaca tactttctgg gttgtggaag 1680
acacctgcag aaagatgttt catgtggctt ggtggattcc gttcatccga acttctcaag 1740
ataattagaa accacctcga ggcgttaacg gatcagcaat tgatggcgat cttcaatctg 1800
cagcattctt gtcaacaggc tgaagatgca ttatctcaag gaatggaagg tttgcaacaa 1860
tctctttcag agacactttc ctccacgtcc actggatctg gaaatgttgt tgagtatatg 1920
ggtcaaatgg ctctttcaat ggccaagctt tccacactgg agactttcat tcatcaggca 1980
gatatcttga ggcaacaaac actgcaacag atgcgtcgaa ttttgactgc gcaccaagct 2040
gctcgtgctc tccttgtcat aaatgatttc atttcacgaa tcagagctct taattcatta 2100
tggtcagcat ttcctaaaga atactaa 2127
<210> 2
<211> 708
<212> PRT
<213> MtbZIP10蛋白(Medicago truncatula)
<400> 2
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1 5 10 15
Asn Asn Gln Cys Leu Ser Thr Val Ser Trp Cys Asn Gly Val Gly Asn
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Glu Ala Leu Val Val Val Glu Ser Lys Asp Gln Cys Lys Thr Lys Gly
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Gln Ser Asp Glu His Lys Thr Leu Arg Arg Leu Met Gln Asn Arg Glu
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Ala Ala Arg Lys Ser Arg Leu Arg Lys Lys Ala Tyr Val Gln Gln Leu
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Glu Asn Ser Arg Leu Arg Leu Ala Gln Ile Glu His Glu Leu Gln Gln
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Val Arg Gln Gln Val Gln Gln Leu Lys Lys Ile Phe His Leu Val Lys
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Leu Val Cys Ser Val Ala Phe Asp Met Asp Tyr Ala Arg Trp Val Asp
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Glu His Gln Arg Leu Ile Asn Asp Ile Arg Ser Ala Ile Asn Ser Gln
130 135 140
Met Gly Asp Asn Glu Leu His Leu Leu Val Asp Gly Val Met Val His
145 150 155 160
Tyr Asp Glu Leu Tyr Lys Leu Lys Ser Ile Gly Ala Lys Ala Asp Val
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Phe His Ile Leu Ser Gly Leu Trp Lys Thr Pro Ala Glu Arg Cys Phe
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Met Trp Leu Gly Gly Phe Arg Ser Ser Glu Leu Leu Lys Ile Ile Arg
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Asn His Leu Glu Ala Leu Thr Asp Gln Gln Leu Met Ala Ile Phe Asn
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Leu Gln His Ser Cys Gln Gln Ala Glu Asp Ala Leu Ser Gln Gly Met
225 230 235 240
Glu Gly Leu Gln Gln Ser Leu Ser Glu Thr Leu Ser Ser Thr Ser Thr
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Gly Ser Gly Asn Val Val Glu Tyr Met Gly Gln Met Ala Leu Ser Met
260 265 270
Ala Lys Leu Ser Thr Leu Glu Thr Phe Ile His Gln Ala Asp Ile Leu
275 280 285
Arg Gln Gln Thr Leu Gln Gln Met Arg Arg Ile Leu Thr Ala His Gln
290 295 300
Ala Ala Arg Ala Leu Leu Val Ile Asn Asp Phe Ile Ser Arg Ile Arg
305 310 315 320
Ala Leu Asn Ser Leu Trp His Val Phe Gly Ala Lys Ser Asn Asn Val
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Thr Val Val Ala Ser Asn Leu Gln Tyr Gly Thr Phe Asn Ser Asn Ile
340 345 350
Gly Ser Ala Ser Ser Ser Gly Met Ala Asp Asn Ser Gln Lys Thr Glu
355 360 365
Asp Phe Asp Thr Asp Asp Lys Asn Asn Gln Ser Gly Asn Val Leu Ile
370 375 380
Phe Leu Cys Leu Ser Thr Val Ser Trp Cys Asn Gly Val Gly Asn Glu
385 390 395 400
Ala Leu Val Val Val Glu Ser Lys Asp Gln Cys Lys Thr Lys Gly Gln
405 410 415
Ser Asp Glu His Lys Thr Leu Arg Arg Leu Met Gln Asn Arg Glu Ala
420 425 430
Ala Arg Lys Ser Arg Leu Arg Lys Lys Ala Tyr Val Gln Gln Leu Glu
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Asn Ser Arg Leu Arg Leu Ala Gln Ile Glu His Glu Leu Gln Gln Val
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Arg Gln Gln Gly Thr Phe Val Ala Pro Gly Val Thr Ala Asp His Gly
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Glu Leu Leu Lys Ile Ile Arg Asn His Leu Glu Ala Leu Thr Asp Gln
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Gln Leu Met Ala Ile Phe Asn Leu Gln His Ser Cys Gln Gln Ala Glu
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Asp Ala Leu Ser Gln Gly Met Glu Gly Leu Gln Gln Ser Leu Ser Glu
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Thr Leu Ser Ser Thr Ser Thr Gly Ser Gly Asn Val Val Glu Tyr Met
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Gly Gln Met Ala Leu Ser Met Ala Lys Leu Ser Thr Leu Glu Thr Phe
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Ile His Gln Ala Asp Ile Leu Arg Gln Gln Thr Leu Gln Gln Met Arg
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Arg Ile Leu Thr Ala His Gln Ala Ala Arg Ala Leu Leu Val Ile Asn
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Asp Phe Ile Ser Arg Ile Arg Ala Leu Asn Ser Leu Trp Ser Ala Phe
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Pro Lys Glu Tyr
705
Claims (2)
1.一种调控植物类黄酮合成的转录因子基因MtbZIP10的用途，其特征在于，所述基因MtbZIP10的核苷酸序列如SEQ ID NO.1所示，所述用途为在蒺藜苜蓿中抑制MtbZIP10的表达导致蒺藜苜蓿地上部分类黄酮中花青素减少，种子中原花色素减少。
2.一种调控植物类黄酮合成的转录因子基因MtbZIP10的用途，其特征在于，所述基因MtbZIP10的核苷酸序列如SEQ ID NO.1所示，所述用途为将MtbZIP10基因在蒺藜苜蓿毛状根中过量表达，从而提高花青素的含量。
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