CN101597329A - 与植物脂肪酸和油脂代谢相关的转录因子及其编码基因与应用 - Google Patents
与植物脂肪酸和油脂代谢相关的转录因子及其编码基因与应用 Download PDFInfo
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Abstract
本发明公开了一种与植物脂肪酸和油脂代谢相关的转录因子及其编码基因与应用。该蛋白是如下a)或b)的蛋白:a)由序列表中序列1所示的氨基酸序列组成的蛋白质;b)在序列表中序列1的氨基酸序列经过取代和/或缺失和/或添加一个或几个氨基酸且与植物脂肪酸或油脂代谢相关的由a)衍生的蛋白质。同时公开了编码该蛋白的基因、含有所述基因的重组表达载体、转基因细胞系和重组菌以及培育脂肪酸和油脂含量提高的转基因植物的方法。本发明的与植物脂肪酸或油脂代谢相关的蛋白及其编码基因在提高植物,特别是油料作物的脂肪酸或油脂含量及相关性状的改良具有重要的应用价值,在农业领域具有广阔的应用和市场前景。
Description
技术领域
本发明涉及与植物脂肪酸和油脂代谢相关的转录因子及其编码基因与应用。
背景技术
植物中脂肪酸又被称为植物油,它广泛存在于植物的种子中。植物油用途广泛,运用生物技术来改良植物体内脂肪酸的组分和含量具有重要经济价值。基因工程方法所选用的操作基因必需依赖于对植物体内脂肪酸生物合成过程的分子调控机制的了解的基础之上。但目前人们对此却了解很少,因此通过基因工程的方法来提高油料作物中含油量的工作目前进展缓慢。
以双子叶模式植物拟南芥为例,植物的胚胎发生主要分为两个阶段:早期形态发生过程和晚期成熟过程。整个胚胎发育过程中伴随着淀粉、蛋白质和脂肪酸等贮藏物质的积累。在胚胎发育早期,主要是淀粉的大量积累,在组织和器官形成以后,淀粉被转化为蛋白质和脂肪酸。在发育的种子中,碳水化合物通过糖酵解途径被分解成磷酸烯醇式丙酮酸((phosphoenolpyruvate,PEP),PEP被运输到质体后再转化成丙酮酸和乙酰CoA,乙酰CoA作为底物参与了脂肪酸的合成(Sari A.Ruuska等,2002,ThePlant Cell,Vol.14,1191-1206.)。
在植物种子成熟过程中,质体内脂肪酸合成酶复合体的酶是负责进行脂肪酸合成的,所合成的脂肪酸被导入胞质酰基CoA库中以维持三酯酰甘油(TAG)积累。植物种子中的TAG的生物合成是在内质网中进行的。前体是甘油-3-磷酸和脂酰COA,合成TAG的过程共需要三种酰基转移酶和一种磷酸水解酶,分别是甘油-3-磷酸酰基转移酶(GPAT),溶血磷脂酸酰基转移酶(LPAT),二脂酰甘油酰基转移酶(DGAT)和磷脂磷酸水解酶(PAPase)(Ohlrogge,J.B等,1979,Proc Natl Acad Sci U S A 76(3):1194-1198.)。这三种酰基转移酶催化甘油骨架的逐步酰基化过程。
近几十年来,科学家们对利用基因工程的方法提高植物体内特别是种子中的脂肪酸含量进行了各种有益的探索。Shintani等(1997)在烟草中过量表达了乙酰辅酶A羧化酶(ACCase)的其中一个亚基-生物素羧化酶,结果发现,在烟草叶片中BC亚基的表达水平提高了三倍,其他三个亚基的表达水平没有发生明显变化,同时脂肪酸的含量和组成也没有明显改变(见Shintani,D.K等,1997,Plant Physiol.114,881-886.)。但是另一项研究表明,增加丙酰COA的含量能够提高脂肪酸的含量。Roeseler等1997年将拟南芥编码的HO-ACCase用种子特异性表达的启动子控制导入油菜中过量表达,结果使ACCase的活性明显提高,并且使种子中油脂的含量提高3-5%(见Roesler,K.等,1997,Plant Physiol.11375-11381.)。Roeseler等的研究结果表明,丙酰CoA水平能够提高脂肪酸的含量,但提高的幅度却很少。Dechesh等(2001)将菠菜的酮酯酰基ACP合酶III在烟草中过量表达,结果使酮酯酰基ACP合酶III的酶活性提高了100-300多倍,但脂肪酸的含量却降低了5-10%(Dehesh,K等,2001,PlantPhysiol.125,1103-1114.)。上述研究结果表明:植物体内脂肪酸代谢途径是一个复杂和高度协调的过程,通过对代谢途径中的个别或单个基因的遗传操作并不能够有效地改变脂肪酸的含量。因此,Girke等人预言在脂肪酸的代谢过程中,很可能存在一种蛋白激酶或其它调控因子(如转录因子)在起控制作用(见Girke,T.等,2000,Plant Physiol 124,1570-1581.)。
Microarray实验表明,在拟南芥种子发育过程中,由糖类物质向脂类物质转化的过程是由多基因协同作用来完成的,因此可能有不止一个基因参与了该过程的调节。其中,转录水平的调节是一个至关重要的环节。在植物的整个基因组中,编码转录因子的基因占了很大一部分,比如,拟南芥中编码转录因子的基因至少有1500个(Riechmann,J.L.等,2000,Science 290,2105-2110.),占整个基因组的5%以上。这些转录因子大多属于大的基因家族,有的基因家族又可包括许多亚族,而且有些转录因子家族是植物所特有的。大量对转录因子研究的结果表明,一个转录因子可能对一类相关性状的很多基因实施调节控制,从而有效改变植物的相关特性。
目前,植物体内已有多个转录因子被发现参与了脂肪酸代谢(Santos等,2005,FEBS lett,579,4666-4670.;Cernac,A等,2004,Plant J 40,575-585.;Wang等,2007,planta,226773-783.)。leafy cotyledon 2(LEC2)基因编码了一个B3结构域的转录因子,2005年Santos等报道了过量表达LEC2基因,可以在营养组织中积累种子储存油脂(Santos等,2005,FEBS lett,579,4666-4670.)。WRI1编码了一个推测的AP2/EREB转录因子蛋白,它可能是植物体内由蔗糖向TAG转化的一个关键的调节因子(Cernac,A等,2004,Plant J 40,575-585)。Cernac和Benning 2004年用CaMV-35S启动子过量表达拟南芥中AP2类转录因子WRI1的cDNA,实验结果表明该方法能够提高种子的含油量(Cernac,A.等,2004,Plant J 40,575-585.)。2007年,Baud等人的研究证实WRI1是LEC2的直接靶基因,在植物种子成熟过程中,LEC2调节WRI1基因的表达,通过WRI1基因来完成蔗糖向脂肪酸的转化,完成脂肪酸的积累(Baud等,2007,Plant J 50,825-838.)。FUS3与LEC2一样都是编码了一个B3结构域的转录因子,B3结构域能够与含有RY-motif的DNA序列结合。通过转基因实验,发现在FUS3过量表达的转基因植株中,脂肪酸合成相关的基因如CAC2,KAS2及SSI2等的mRNA水平明显受FUS3诱导(Wang等,2007,Planta,226 773-783.)。上述研究表明,转录因子在植物的脂肪酸代谢尤其是种子中油脂积累的过程中起着重要的作用。
在植物中,转录因子按照其DNA结合域的相似性可以分为几个家族,MYB转录因子家族是其中最大的家族之一。MYB转录因子蛋白的N端具有保守的MYB-DNA结合域;这些DNA结合域是由一个到三个大约50个氨基酸残基的螺旋-转角-螺旋不完全重复结构构成,按照这些不完全重复结构又可以分为MYB1R,MYBR2R3和MYB3R。模式植物拟南芥的MYB因子(AtMYB)参与调控了植物的多种生理活动过程,包括:植物发育,细胞命运和细胞特征决定,对环境因子和植物激素的反应,非生物胁迫,病原菌侵染抗性以及植物次生代谢物合成等信号转导过程(Ralf,S等,2001,Curr Opin PlantBiol 4:447-456.)。
AtMYB转录因子大部分都是R2R3型的MYB因子。目前研究比较清楚的R2R3型MYB因子多参与了植物次生物代谢调控,如AtMYB4,AtMYB75/PAP1,AtMYB90/PAP2,AtMYB123/TT2。除此之外,AtMYB34/ATR1参与了色氨酸的合成调控。然而大部分的已经预测到的AtMYB转录因子功能或者他们参与的信号转动途径未知。
发明内容
本发明的目的是提供与植物脂肪酸和油脂代谢相关的转录因子及其编码基因与应用,以提高商品植物油的产量以实现对植物和植物产品特定的商品性的改良。
本发明所提供的与植物脂肪酸和油脂代谢相关的转录因子,命名为AtMYB118,来源于拟南芥,是如下a)或b)的蛋白:
a)由序列表中序列1所示的氨基酸序列组成的蛋白质;
b)在序列表中序列1的氨基酸序列经过取代和/或缺失和/或添加一个或几个氨基酸且与植物脂肪酸代谢相关的由a)衍生的蛋白质。
其中,序列表中的序列1由437个氨基酸残基组成。其中,自N末端189-235位氨基酸和自N端241-286分别为MYB结构域。
为了使a)的AtMYB118蛋白质便于纯化,可在由序列表中序列1所示的氨基酸序列组成的蛋白质的氨基末端或羧基末端连接上如表1所示的标签。
表1.标签的序列
标签 | 残基 | 序列 |
Poly-Arg | 5-6(通常为5个) | RRRRR |
Poly-His | 2-10(通常为6个) | HHHHHH |
FLAG | 8 | DYKDDDDK |
Strep-tag II | 8 | WSHPQFEK |
c-myc | 10 | EQKLISEEDL |
上述b)中的AtMYB118蛋白质可人工合成,也可先合成其编码基因,再进行生物表达得到。上述b)中的AtMYB118蛋白质的编码基因可通过将序列表中序列3自5′末端第1-1314位所示的DNA序列中缺失一个或几个氨基酸残基的密码子,和/或进行一个或几个碱基对的错义突变,和/或在其5′端和/或3′端连上表1所示的标签的编码序列得到。
AtMYB118蛋白的编码基因也属于本发明的保护范围。
AtMYB118蛋白的编码基因,是如下1)至5)中任一所述的基因;
1)其核苷酸序列是序列表中序列2;
2)其核苷酸序列是序列表中序列3;
3)其编码序列是序列表中序列3自5′末端第1-1314位;
4)在严格条件下与1)或2)或3)限定的DNA片段杂交且编码植物脂肪酸和油脂代谢相关的转录因子的DNA分子;
5)与1)或2)或3)的基因具有90%以上的同源性,且编码植物脂肪酸代谢相关的转录因子的DNA分子。
所述步骤5)中的基因,与1)或2)或3)的基因最好有95%以上的同源性。
序列表中的序列2由2332个碱基组成,包括222个碱基的5′UTR区,开放阅读框架为5′端第222到第2168位碱基,3′UTR区为2169到2332位碱基。自5′端第223-366位碱基为该基因组基因的第一个外显子,自5′端第367-474位碱基为该基因组基因的第一个内含子,自5′端第475-923位碱基为该基因组基因的第二个外显子,自5′端第924-1029位碱基为该基因组基因的第二个内含子,自5′端第1030-1156位碱基为该基因组基因的第三个外显子,自5′端第1157-1574位碱基为该基因组基因的第三个内含子,自5′端第1575-2168位碱基为该基因组基因的第四个外显子,自5′端第2169-2332位碱基为该基因组基因的3′非编码区。
序列表中的序列3由1314个碱基组成,其编码序列为自5′端第1-1314位碱基,编码具有序列表中序列1所示的的氨基酸残基序列的蛋白质,自5′端第567-705位碱基编码MYB结构域,自5′端第723-858位碱基编码MYB结构域。
上述严格条件可为在0.1×SSPE(或0.1×SSC)、0.1%SDS的溶液中,65℃条件下杂交并洗膜。
扩增AtMYB118基因全长或任一片段的引物对也属于本发明的保护范围。
含有上述AtMYB118基因的重组载体、转基因细胞系和重组菌也属于本发明的保护范围。
可用现有的植物表达载体构建含有AtMYB118基因的重组表达载体。用于构建所述植物表达载体的出发载体可为任意一种可用于根瘤农杆菌或发根农杆菌转化植物的双元载体或可用于植物微弹轰击的载体等,如pER8、pX6、pBI系列载体、pBin系列载体、pCAMBIA系列载体或其它衍生植物表达载体,所述出发载体还可为可在原核生物中复制的载体,如pUC系列载体或pBluescript系列载体等。
使用AtMYB118基因构建重组植物表达载体时,在其转录起始核苷酸前可加上任何一种增强型、组成型、组织特异型或诱导型启动子。所述组成性表达启动子可为花椰菜花叶病毒(CaMV)35S启动子,玉米Ubiquitin启动子或水稻actin1启动子等;所述组织特异性表达启动子可为种子特异性表达启动子、花特异性表达启动子或根特异性表达启动子;所述诱导型启动子可为受ABA、乙烯、乙醇、***或***等诱导的启动子。它们可单独使用或与其它的植物启动子结合使用。此外,使用本发明的AtMYB118基因构建植物表达载体时,还可使用增强子,包括翻译增强子或转录增强子,这些增强子区域可以是ATG起始密码子或邻接区域起始密码子等,但必需与编码序列的阅读框相同,以保证整个序列的正确翻译。所述翻译控制信号和起始密码子的来源是广泛的,可以是天然的,也可以是合成的。翻译起始区域可以来自转录起始区域或结构基因。
为了便于对转基因植物细胞或植物进行鉴定及筛选,可对所用植物表达载体进行加工,如加入可在植物中表达可产生颜色变化的酶或发光化合物的基因(GUS基因、萤光素酶基因等)、具有抗性的抗生素标记物(新霉素磷酸转移酶基因、潮霉素磷酸转移酶基因、庆大霉素标记物或卡那霉素标记物等)或是抗化学试剂标记基因(如抗除莠剂基因)等。所述含新霉素磷酸转移酶基因的宿主植物细胞、组织或器官可由卡那霉素或其替代衍生物如G418等进行筛选,含潮霉素磷酸转移酶基因的宿主植物细胞、组织或器官可由潮霉素进行筛选。从转基因植物的安全性考虑,可不加任何选择性标记基因,直接以脂肪酸含量或组成的改变筛选转化植株。经上述方法进行筛选后还可采用Southern、PCR或点杂交等分子检测手段对转基因植株进行检测,以确定其是否转化有目的基因。
本发明的另一个目的是提供一种培育脂肪酸和/或油脂含量提高、脂肪酸和/或油脂组分改变的转基因植物的方法。
本发明提供的培育脂肪酸和/或油脂含量提高、脂肪酸和/或油脂组分改变的转基因植物的方法,是将所述AtMYB118基因转入植物细胞、组织或器官中,获得脂肪酸和/或油脂含量提高、脂肪酸和/或油脂组分改变的转基因植物。
其中,所述脂肪酸和/或油脂含量的提高包括植物体内各组织和器官内各种脂肪酸和/或油脂含量(饱和脂肪酸和不饱和脂肪酸)的提高,所述脂肪酸和/或油脂组分的改变包括植物体内各组织和器官内的脂肪酸和/或油脂组成的改善。
所述AtMYB118基因既可为AtMYB118的cDNA序列,也可为AtMYB118的基因组基因序列或与AtMYB118具有90%以上同源性且编码相同蛋白的DNA序列。
与AtMYB118具有90%以上同源性且编码相同蛋白的DNA序列是将AtMYB118的cDNA或基因组基因序列用已知的方法进行分离和/或修饰和/或设计得到的。本领域的技术人员应该理解的是,特定基因序列中核苷酸同一性的微小改变可能会导致该基因效能的降低或者加强,而且在一些应用(例如,反义或共抑制技术)中,部分序列经常会和全长序列同样有效地发挥作用。基因序列变化或缩短的方法,以及测试这些发生变体的基因的有效性的方法均是本领域技术人员熟知的。
所述AtMYB118基因或其同源序列可以正义方向或反义方向导入植物组织、细胞或器官。
所述AtMYB118基因转入植物细胞、组织或器官中可通过使用原生质体-化学介导法(Ca2+、PEG)、Ti质粒、Ri质粒、植物病毒载体、直接DNA转化、花粉管、微注射、电激、基因枪、农杆菌介导等常规生物学方法中的任何一种或几种方法的组合转化植物细胞、组织或器官,并将转化的植物细胞、组织或器官培育成植株;所述组织和器官可包括宿主植物的果荚、愈伤组织、茎尖、叶片和种子等。
本发明的方法对双子叶植物和单子叶植物均适用,因此,所述被转化的植物细胞、组织或器官既可来源于拟南芥、油菜、花生、棉花、大豆、向日葵、棕榈树、橄榄树、蓖麻、马铃薯或烟草等双子叶植物,也可来源于水稻、玉米、小麦、大麦、燕麦、黑麦、高梁、谷子或草坪草等单子叶植物。
本发明用RT-PCR方法从拟南芥(Arabidopsis thaliana)哥伦比亚生态型中分离得到AtMYB118基因。实验证明AtMYB118基因过表达的转基因拟南芥植株体内的油酸(C18:1)和亚油酸(C18:2)的含量明显升高。本发明的与植物脂肪酸和油脂代谢相关的蛋白及其编码基因在提高植物(特别是油料作物)的脂肪酸和/或油脂含量及相关性状的改良具有重要的实际意义,在农业领域具有广阔的应用和市场前景。
附图说明
图1为AtMYB118基因的框架结构。
图2为转AtMYB118基因的拟南芥的表达检测
1:转pER8载体的拟南芥;2:AtMYB118基因过表达的拟南芥#2;3:AtMYB118基因过表达的拟南芥#7。
图3为AtMYB118基因过表达拟南芥在诱导培养基上萌发后10天的表型。
图4为AtMYB118基因过表达拟南芥在诱导培养基上萌发后30天的表型。
图5为苏丹红染色指示AtMYB118基因过表达拟南芥体内脂肪酸或油脂含量。
A:***诱导16小时的转pER8空载体植物;B:***诱导16小时的转pER8-AtMYB118植物。
图6为GC-MS方法检测AtMYB118基因过表达拟南芥在诱导培养基上萌发后体内脂肪酸或油脂含量变化情况。
具体实施方式
下述实施例中所用方法如无特别说明均为常规方法,所用引物及探针均由北京奥科生物公司合成。
实施例1、与植物脂肪酸和油脂代谢相关的基因AtMYB118的克隆
根据AtMYB118的核苷酸序列设计引物P1和P2,用于扩增拟南芥(Arabidopsisthaliana)哥伦比亚生态型(Col-0)中的与植物脂肪酸代谢相关的基因,引物P1和P2的序列如下:
P1(AtMYB118上游引物):5′ATAGGCGCGCCATGGAGTTCGAGTCAGTGTTCA 3′
P2(AtMYB118下游引物):5′AATAACTAGTCTAAAGACGACCATGAGCAATCA 3′
用TRIZAL试剂(Invitrogen)并参照试剂盒说明书提取拟南芥(Arabidopsisthaliana)哥伦比亚生态型(Col-0)新鲜角果的总RNA,然后用SuperScriptTM IIReverse Transcriptase试剂盒(Invitrogen公司)反转录合成其第一链cDNA,再以所合成的cDNA为模板,在引物P1和P2的引导下进行PCR扩增,反应结束后,对PCR扩增产物进行1%琼脂糖凝胶电泳检测,用DNA回收试剂盒(鼎国公司)回收长度约1000bp的目的片段,将回收片段连接入到载体pGEM-Teasy(Promega公司)中,再将连接产物用热激法转化大肠杆菌DH5α感受态细胞,筛选阳性克隆,将其接种于含50mg/L氨苄青霉素的5mL LB液体培养基中,在37℃、200rpm下培养12-16小时,提质粒,得到含有回收片段的重组质粒,命名为pGEM-T Vector-AtMYB118c,对其进行测序,测序结果表明扩增片段具有序列表中的序列3所示的核苷酸序列组成,由1314个碱基组成,编码序列为自5′端第1-1314位碱基,编码具有序列表中序列1所示的氨基酸残基序列的蛋白质,其中,自5′端第567-705位碱基编码MYB结构域,自5′端第723-858位碱基编码MYB结构域。
利用上述引物P1和P2,从哥伦比亚生态型(Col-0)拟南芥基因组中用高保真DNA聚合酶KOD-Plus(日本东洋纺公司)进行PCR扩增。PCR扩增产物按照上述方法连接到pGEM-Teasy(Promega公司)中,构建重组表达载体pGEM-T Vector-AtMYB118,对其进行测序,测序结果表明扩增片段具有序列表中序列2的核苷酸序列,由2332个碱基组成,自5′端第223-366位碱基为该基因组基因的第一个外显子,自5′端第367-474位碱基为该基因组基因的第一个内含子,自5′端第475-923位碱基为该基因组基因的第二个外显子,自5′端第924-1029位碱基为该基因组基因的第二个内含子,自5′端第1030-1156位碱基为该基因组基因的第三个外显子,自5′端第1157-1574位碱基为该基因组基因的第三个内含子,自5′端第1575-2168位碱基为该基因组基因的第四个外显子,自5′端第2169-2332位碱基为该基因组基因的3’非编码区。该基因的框架结构见图1,将该基因命名为AtMYB118,将其编码蛋白命名为AtMYB118。
用Xho1和Spe1酶切pGEM-T Vector-AtMYB118质粒DNA,酶切得到片段用凝胶回收试剂盒(购自鼎国公司)回收,纯化后的片段用T4连接酶(Roche公司)与经过同样双酶切的的pER8载体(Zuo等,2001年,Plant J.24:265-273.US Patent 6452068)(中国科学院遗传与发育生物学研究所)连接,获得重组表达载体pER8-AtMYB118。重组表达载体pER8-AtMYB118通过热激法转化大肠杆菌(E.coli)DH5α,挑取阳性菌落到5ml含50mg/L潮霉素的LB液体培养基中,37℃、200rpm培养12-16小时,提取质粒,进行PCR鉴定和酶切鉴定。
实施例2、转AtMYB118基因拟南芥的获得
将鉴定正确的重组表达载体pER8-AtMYB118通过电激转入农杆菌GV3101中,挑取农杆菌单菌落接种于20ml LB液体培养基(壮观霉素50mg/L,利福平50mg/L)中,28℃,150rpm振荡培养2天。再按2%的接种量将菌液接种于含有利福平和壮观霉素的300ml LB培养基振荡培养16-18小时,5000rpm,20分钟离心收集菌体,溶于250ml含有5%蔗糖Silwetl-77中,慢慢摇匀。溶液转于250ml烧杯中将已去掉花和果荚的植物倒置于烧杯中,真空抽20秒。为提高转化效率,一周后可再重复一次。所得种子经潮霉素筛选后得到转AtMYB118基因拟南芥,共得到了。
按照上述方法将pER8载体转入哥伦比亚生态型拟南芥中,获得转pER8载体的拟南芥植株,移栽到蛭石中繁殖种子,作为对照。
实施例3、转AtMYB118基因的拟南芥的表达检测
挑选实施例2中14个转AtMYB118基因的拟南芥转基因系中两个不同转基因系#2和#7号检测拟南芥体内AtMYB118基因的表达水平,以转pER8载体的拟南芥植株作为对照。以上植物种子分别播种在1/2MS培养基(Sigma公司)。在16小时光照,22℃培养2周左右。然后,转到含有10μM***的液体1/2MS培养基中,诱导16小时。
用TRIZOL试剂(购自Invitrogen公司)并参照试剂盒说明书提取总RNA,然后用Invitrogen公司的SuperScriptTM II Reverse Transcriptase试剂盒并参照试剂盒说明书反转录合成其第一链cDNA,再以所合成的cDNA为模板,在AtMYB118上游引物和AtMYB118下游引物的引导下进行PCR扩增,反应结束后,对PCR扩增产物进行1%琼脂糖凝胶电泳检测。其中,PCR扩增的反应条件是94℃变性2分钟,94℃30秒,56℃30秒,72℃30秒,共进行25轮反应,最后72℃延伸10分钟。
结果表明,转AtMYB118基因的拟南芥转基因系中两个不同转基因系#2和#7号中的AtMYB118基因内被***诱导表达,而转pER8载体的拟南芥植株中AtMYB118基因的与未转基因的哥伦比亚生态型拟南芥均不能被诱导表达。图2所示为#2和#7转基系AtMYB118基因的表达水平。图2中1代表转pER8载体的拟南芥幼苗中AtMYB118基因没有表达。2代表#2转基因拟南芥中AtMYB118基因被诱导表达,3代表#7转基因拟南芥中AtMYB118基因被诱导表达。
实施例4、转AtMYB118基因的拟南芥在诱导培养基上的萌发
将转AtMYB118基因的拟南芥和转pER8载体的拟南芥植株分别移栽到蛭石中繁殖种子,将收获的种子分别播种在含10μmol/L***的MS培养基上,以转pER8载体的拟南芥作为对照,在培养条件为22℃,光照强度80-120μE-2S-1,16小时光照的温室中培养。转AtMYB118基因的植物的表型为花青素积累、绿根,根部膨大***、植株矮小。图3和图4为转AtMYB118基因的拟南芥植株在诱导培养基上生长分别10天和30天时的表型。转pER8载体的拟南芥在诱导培养基上分别生长10天和30天均未出现异常表型。
图3中左边为在诱导培养基上生长10天的转pER8载体的拟南芥,右边为在诱导培养基上生长10天的转AtMYB118基因的拟南芥;图4中左边为在诱导培养基上生长30天的转pER8载体的拟南芥,右边为在诱导培养基上生长30天的转AtMYB118基因的拟南芥。
实施例6、苏丹红染色指示转基因植株体内脂肪酸或油脂含量的变化
将转AtMYB118基因的拟南芥的种子和转pER8载体的拟南芥植株的种子分别播种在1/2MS培养基(Sigma公司)。在16小时光照,22℃培养20天左右。然后,分别转到含有10μM***的液体1/2MS培养基中,诱导16小时。同时,以分别转到不含***的液体1/2MS培养基中,诱导16小时作为对照。
分别取上述诱导培养基中的转AtMYB118基因的拟南芥植株与转pER8载体的拟南芥植株浸泡于1%苏丹红(Fat Red 7B)(Sigma公司),室温30分钟,用去离子水清洗3遍。
苏丹红染色实验结果如图5所示,转AtMYB118基因的拟南芥植株的根部和子叶处的脂肪酸的积累明显比转pER8载体的拟南芥植株的根部和子叶处的脂肪酸的积累水平高。
实施例7、气质联用(GC-MS)方法检测转AtMYB118基因的拟南芥经诱导后体内脂肪酸含量变化
将转AtMYB118基因的拟南芥种子和转pER8载体的拟南芥植株的种子分别播种在1/2MS培养基(Sigma公司)。在16小时光照,22℃培养20天左右。然后,分别转到含有10μM***的液体1/2MS培养基中,诱导16小时。同时,以分别转到不含***的液体1/2MS培养基中,诱导16小时作为对照。
分别取上述诱导培养基中的转AtMYB118基因的拟南芥植株与转pER8载体的拟南芥植株各100mg,在液氮中研磨成干粉,然后转移到带密封盖的试管中,加入3ml甲醇(含2.5%(体积百分比)浓硫酸),在80℃水浴加热90分钟.再加入4.5ml 0.9g/100ml的NaCL和1ml正己烷,混匀,4000rpm离心10分钟,收集正己烷相。真空抽干,用100ul乙酸乙酯溶解,取1ul上样分析。所用GC-MS仪为TurboMass(PerkinElmer公司),所用GC柱为30m×0.25mm BPX-70柱,GC升温程序为:初始温度120℃,保持1分钟,以每分钟10℃的速率升至150℃,然后以每分钟4℃的速率升温至230℃,保持10分钟.以C17:0的三酯酰甘油作内标。
GC-MS结果如图6所示,转AtMYB118基因的拟南芥植株体内的油酸(C18:1)和亚油酸(C18:2)的含量明显升高,其中油酸的含量与转pER8载体的拟南芥植株相比,转AtMYB118基因的拟南芥植株体内的油酸(C18:1)和亚油酸(C18:2)的含量升高了10倍之多,说明MYB118基因过量表达后可以显著提高植物体内脂肪酸尤其是不饱和脂肪酸的含量。
序列表
<110>中国科学院遗传与发育生物学研究所
<120>与植物脂肪酸和油脂代谢相关的转录因子及其编码基因与应用
<130>CGGNARW81379
<160>3
<210>1
<211>437
<212>PRT
<213>拟南芥属拟南芥(Arabidopsis thaliana)
<400>1
Met Glu Phe Glu Ser Val Phe Lys Met His Tyr Pro Tyr Leu Ala Ala
1 5 10 15
Val Ile Tyr Asp Asp Ser Ser Thr Leu Lys Asp Phe His Pro Ser Leu
20 25 30
Thr Asp Asp Phe Ser Cys Val His Asn Val His His Lys Pro Ser Met
35 40 45
Pro His Thr Tyr Glu Ile Pro Ser Lys Glu Thr Ile Arg Gly Ile Thr
50 55 60
Pro Ser Pro Cys Thr Glu Ala Phe Glu Ala Cys Phe His Gly Thr Ser
65 70 75 80
Asn Asp His Val Phe Phe Gly Met Ala Tyr Thr Thr Pro Pro Thr Ile
85 90 95
Glu Pro Asn Val Ser His Val Ser His Asp Asn Thr Met Trp Glu Asn
100 105 110
Asp Gln Asn Gln Gly Phe Xaa Phe Gly Thr Glu Ser Thr Leu Asn Gln
115 120 125
Ala Met Xaa Asp Ser Asn Gln Phe Asn Met Pro Lys Pro Leu Leu Ser
130 135 140
Ala Asn Glu Asp Thr Ile Met Asn Arg Arg Gln Asn Asn Gln Val Met
145 150 155 160
Ile Lys Thr Glu Gln Ile Lys Lys Lys Asn Lys Arg Phe Gln Met Arg
165 170 175
Arg Ile Cys Lys Pro Thr Lys Lys Ala Ser Ile Ile Lys Gly Gln Trp
180 185 190
Thr Pro Glu Glu Asp Lys Leu Leu Val Gln Leu Val Asp Leu His Gly
195 200 205
Thr Lys Lys Trp Ser Gln Ile Ala Lys Met Leu Gln Gly Arg Val Gly
210 215 220
Lys Gln Cys Arg Glu Arg Trp His Asn His Leu Arg Pro Asp Ile Lys
225 230 235 240
Lys Asp Gly Trp Thr Glu Glu Glu Asp Ile Ile Leu Ile Lys Ala His
245 250 255
Lys Glu Ile Gly Asn Arg Trp Ala Glu Ile Ala Arg Lys Leu Pro Gly
260 265 270
Arg Thr Glu Asn Thr Ile Lys Asn His Trp Asn Ala Thr Lys Arg Arg
275 280 285
Gln His Ser Arg Arg Thr Lys Gly Lys Asp Glu Ile Ser Leu Ser Leu
290 295 300
Gly Ser Asn Thr Leu Gln Asn Tyr Ile Arg Ser Val Thr Tyr Asn Asp
305 310 315 320
Asp Pro Phe Met Thr Ala Asn Ala Asn Ala Asn Ile Gly Pro Arg Asn
325 330 335
Met Arg Gly Lys Gly Lys Asn Val Met Val Ala Val Ser Glu Tyr Asp
340 345 350
Glu Gly Glu Cys Lys Tyr Ile Val Asp Gly Val Asn Asn Leu Gly Leu
355 360 365
Glu Asp Gly Arg Ile Lys Met Pro Ser Leu Ala Ala Met Ser Ala Ser
370 375 380
Gly Ser Ala Ser Thr Ser Gly Ser Ala Ser Gly Ser Gly Ser Gly Val
385 390 395 400
Thr Met Glu Ile Asp Glu Pro Met Thr Asp Ser Trp Met Val Met His
405 410 415
Gly Cys Asp Glu Val Met Met Asn Glu Ile Ala Leu Leu Glu Met Ile
420 425 430
Ala His Gly Arg Leu
435
<210>2
<211>2332
<212>DNA
<213>拟南芥属拟南芥(Arabidopsis thaliana)
<400>2
ctaatcaatt actttcctcg ttatcctttt ttgcagtcaa gttaaattgc tctctttcaa 60
gacttgtgtt ctttaaacca aaaaaaaaaa aaaagtatct gtgttcatca ccaactcatt 120
cttctttcag atctagggtt tcatgcttca ctcaattttt ttttgtttag gtagttccat 180
cttctaaacg ttgtattttt ttttttttgc catcataatc atatggagtt cgagtcagtg 240
ttcaaaatgc attatccgta tctcgcagcc gttatctacg atgatagctc cactttaaaa 300
gattttcatc catctcttac cgatgatttt tcttgtgtac acaatgtgca tcacaaacca 360
tcgatggtaa actctaatcc tgatcttttt ttcacaagaa tctcacatta aagatctcaa 420
tcctatcact ttactcaaaa tcttaatatt gttccctatc actttgatat gcagcctcac 480
acatatgaaa taccatcaaa agaaaccatt aggggcatca ctccttctcc atgcactgaa 540
gctttcgagg catgttttca tggcacatcc aacgaccatg ttttttttgg catggcctat 600
accaccccac caactattga acccaacgtt tcacatgtct cacatgacaa tactatgtgg 660
gaaaacgatc aaaaccaagg attcatcttt ggaaccgagt caaccctcaa tcaagccatg 720
gcggactcta atcaattcaa tatgccaaaa ccactcttga gcgcaaacga agacaccatc 780
atgaatcgac gtcaaaataa ccaggtaatg atcaagaccg agcagatcaa gaagaagaac 840
aagagatttc agatgaggag gatatgtaaa cccacaaaaa aagctagcat catcaaagga 900
caatggactc ctgaagaaga caagtaaatt caacatctaa ttttttggaa ttctcataaa 960
taattttctt aatctagtat gatattgttt ttaatattaa tattttctaa tgattctacc 1020
tatatttagg ttattggtgc agctagtgga ccttcacgga actaaaaaat ggtctcagat 1080
tgctaagatg cttcaaggac gagttggaaa acagtgcaga gaaaggtggc ataaccatct 1140
ccgtcccgat atcaaggtct cttattttat tttattttca ttttcttatt ttcataggag 1200
aaacaaaaac aaaatatcat cttcgatact ttcaaattca ttacatgata cattgagaat 1260
cttctaagag aatattaaca aacttttttc acacttgtca acatgtatac ggttttattt 1320
catgcatgaa tactttaaac ttttgtttgt atatacatat atgtctatga caattcttta 1380
aggcattttg tcttcttgtt gcggacaaaa cttcacaatt ggtaagacat ttaaactaaa 1440
gtcattgtca cttggattct ctcatataat tcaacaaaca taaaagagaa aaaaacatat 1500
aaaaaataaa aatatttgtt taatattgta tgtattattt acatataaag catattttat 1560
gtggtatgtt acagaaagat ggatggactg aagaagagga tataatactg ataaaagccc 1620
ataaggagat tgggaacaga tgggctgaga tagctcgaaa actcccggga cgcactgaaa 1680
atacgatcaa gaaccattgg aacgcgacta aacgtcgaca acactcgagg aggactaaag 1740
gaaaagatga aatttccctt tcacttggta gcaacactct tcagaactac attaggtctg 1800
ttacctacaa tgatgatcct ttcatgaccg caaatgcaaa cgcaaacatt ggtccaagaa 1860
acatgagagg taaaggtaag aatgtaatgg ttgcggtctc ggagtatgat gagggtgaat 1920
gtaagtatat tgtggatggt gtgaataact tgggtttaga agatggaagg atcaagatgc 1980
cgtcattggc ggctatgtcg gcctccggat cagcgtctac ttctggttct gcgtctggtt 2040
ctggaagtgg tgtgaccatg gagattgatg agccgatgac tgatagctgg atggtgatgc 2100
atggatgtga tgaagttatg atgaacgaga ttgctttgct ggagatgatt gctcatggtc 2160
gtctttagac cgtgcaatat aaacaagtcg aacttgatta tatctaccta tataattatg 2220
ctatttatga aattatgttt gtgcttttaa ttgaagaatt ggacatgtaa tatatattgt 2280
ggtttaattg aaggtttctt ttggactaaa caaaaattga aggttttctt gc 2332
<210>3
<211>1314
<212>DNA
<213>拟南芥属拟南芥(Arabidopsis thaliana)
<400>3
atggagttcg agtcagtgtt caaaatgcat tatccgtatc tcgcagccgt tatctacgat 60
gatagctcca ctttaaaaga ttttcatcca tctcttaccg atgatttttc ttgtgtacac 120
aatgtgcatc acaaaccatc gatgcctcac acatatgaaa taccatcaaa agaaaccatt 180
aggggcatca ctccttctcc atgcactgaa gctttcgagg catgttttca tggcacatcc 240
aacgaccatg ttttttttgg catggcctat accaccccac caactattga acccaacgtt 300
tcacatgtct cacatgacaa tactatgtgg gaaaacgatc aaaaccaagg attcatcttt 360
ggaaccgagt caaccctcaa tcaagccatg gcggactcta atcaattcaa tatgccaaaa 420
ccactcttga gcgcaaacga agacaccatc atgaatcgac gtcaaaataa ccaggtaatg 480
atcaagaccg agcagatcaa gaagaagaac aagagatttc agatgaggag gatatgtaaa 540
cccacaaaaa aagctagcat catcaaagga caatggactc ctgaagaaga caagttattg 600
gtgcagctag tggaccttca cggaactaaa aaatggtctc agattgctaa gatgcttcaa 660
ggacgagttg gaaaacagtg cagagaaagg tggcataacc atctccgtcc cgatatcaag 720
aaagatggat ggactgaaga agaggatata atactgataa aagcccataa ggagattggg 780
aacagatggg ctgagatagc tcgaaaactc ccgggacgca ctgaaaatac gatcaagaac 840
cattggaacg cgactaaacg tcgacaacac tcgaggagga ctaaaggaaa agatgaaatt 900
tccctttcac ttggtagcaa cactcttcag aactacatta ggtctgttac ctacaatgat 960
gatcctttca tgaccgcaaa tgcaaacgca aacattggtc caagaaacat gagaggtaaa 1020
ggtaagaatg taatggttgc ggtctcggag tatgatgagg gtgaatgtaa gtatattgtg 1080
gatggtgtga ataacttggg tttagaagat ggaaggatca agatgccgtc attggcggct 1140
atgtcggcct ccggatcagc gtctacttct ggttctgcgt ctggttctgg aagtggtgtg 1200
accatggaga ttgatgagcc gatgactgat agctggatgg tgatgcatgg atgtgatgaa 1260
gttatgatga acgagattgc tttgctggag atgattgctc atggtcgtct ttag 1314
Claims (10)
1、一种蛋白质,是如下a)或b)的蛋白:
a)由序列表中序列1所示的氨基酸序列组成的蛋白质;
b)在序列表中序列1的氨基酸序列经过取代和/或缺失和/或添加一个或几个氨基酸且与植物脂肪酸和/或油脂代谢相关的由a)衍生的蛋白质。
2、权利要求1所述蛋白的编码基因。
3、根据权利要求2所述的基因,其特征在于:所述基因是如下1)或2)或3)或4)或5)的基因:
1)其核苷酸序列是序列表中序列2;
2)其核苷酸序列是序列表中序列3;
3)其编码序列是序列表中序列3的自5′末端第1-1314位;
4)在严格条件下与1)或2)或3)限定的DNA片段杂交且编码植物脂肪酸和/或油脂代谢相关蛋白的DNA分子;
5)与1)或2)或3)的基因具有90%以上的同源性,且编码植物脂肪酸和/或油脂代谢相关蛋白的DNA分子。
4、含有权利要求2或3所述基因的重组表达载体。
5、根据权利要求4所述的重组表达载体,其特征在于:所述重组表达载体的出发载体为pER8系列载体或pX6系列载体或pBI系列载体或pBin系列载体或pCAMBIA系列载体或pUC系列载体或pBluescript系列载体。
6、含有权利要求2或3所述基因的转基因细胞系或重组菌;扩增权利要求2或3所述基因的全长及其任意片段的引物对。
7、一种培育脂肪酸和/或油脂含量提高和/或脂肪酸和油脂组分改变的转基因植物的方法,是将权利要求2或3所述编码基因转入植物细胞、组织或器官中,获得脂肪酸和油脂含量提高和/或脂肪酸和油脂组分改变的转基因植物。
8、一种培育脂肪酸和/或油脂组分改变的转基因植物的方法,是将权利要求2或3所述编码基因转入植物细胞、组织或器官中,获得脂肪酸和/或油脂组分改变的转基因植物。
9、根据权利要求7或8所述的方法,其特征在于:所述脂肪酸为饱和脂肪酸和/或不饱和脂肪酸。
10、根据权利要求9所述的方法,其特征在于:所述植物为所有单子叶植物和所有双子叶植物。
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Cited By (4)
Publication number | Priority date | Publication date | Assignee | Title |
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CN102532291A (zh) * | 2012-01-10 | 2012-07-04 | 中国科学院遗传与发育生物学研究所 | Nf-ya1蛋白及其编码基因在培育脂肪酸含量提高的植物中的应用 |
CN104877021A (zh) * | 2015-06-02 | 2015-09-02 | 南京农业大学 | 与植物脂肪酸和油脂代谢相关的油菜转录因子BnFUS3及其编码基因与应用 |
CN110951769A (zh) * | 2011-12-27 | 2020-04-03 | 联邦科学技术研究组织 | 产生脂质的方法 |
CN111574603A (zh) * | 2020-05-14 | 2020-08-25 | 安徽农业大学 | 一种与脂肪酸合成有关的转录因子及其dna分子、提高油料作物油脂含量的方法、及应用 |
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Cited By (7)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
CN110951769A (zh) * | 2011-12-27 | 2020-04-03 | 联邦科学技术研究组织 | 产生脂质的方法 |
CN102532291A (zh) * | 2012-01-10 | 2012-07-04 | 中国科学院遗传与发育生物学研究所 | Nf-ya1蛋白及其编码基因在培育脂肪酸含量提高的植物中的应用 |
CN102532291B (zh) * | 2012-01-10 | 2014-05-14 | 中国科学院遗传与发育生物学研究所 | Nf-ya1蛋白及其编码基因在培育脂肪酸含量提高的植物中的应用 |
CN104877021A (zh) * | 2015-06-02 | 2015-09-02 | 南京农业大学 | 与植物脂肪酸和油脂代谢相关的油菜转录因子BnFUS3及其编码基因与应用 |
CN104877021B (zh) * | 2015-06-02 | 2018-09-11 | 南京农业大学 | 与植物脂肪酸和油脂代谢相关的油菜转录因子BnFUS3及其编码基因与应用 |
CN111574603A (zh) * | 2020-05-14 | 2020-08-25 | 安徽农业大学 | 一种与脂肪酸合成有关的转录因子及其dna分子、提高油料作物油脂含量的方法、及应用 |
CN111574603B (zh) * | 2020-05-14 | 2022-10-14 | 安徽农业大学 | 一种与脂肪酸合成有关的转录因子及其dna分子、提高油料作物油脂含量的方法、及应用 |
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