CN107418971A - 玉米籽粒类胡萝卜素缺陷基因scd的克隆及应用 - Google Patents
玉米籽粒类胡萝卜素缺陷基因scd的克隆及应用 Download PDFInfo
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Abstract
本发明公开了玉米籽粒类胡萝卜素缺陷基因SCD的克隆及应用。本发明公开的SCD基因编码如下的蛋白质:A1)氨基酸序列是序列表中序列1的蛋白质;A2)氨基酸序列是序列表中序列4的蛋白质;A3)将序列表中序列1或序列4所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有相同功能的蛋白质。实验证明,SCD基因及其编码的蛋白质与玉米白化相关,可用于调控玉米植株、叶片和籽粒颜色,也可用于构建玉米白化模型,可用于植物基因工程、基因编辑技术及分子育种中。
Description
技术领域
本发明涉及植物分子遗传学和植物基因工程领域,具体地,本发明涉及玉米籽粒类胡萝卜素缺陷基因SCD的克隆及功能验证。该基因可用于植物基因工程、基因编辑技术及分子育种中。
背景技术
玉米是重要的粮饲兼用型作物,同时也是研究植物遗传和基因功能的重要模式植物。玉米的种植面积和总产已超过水稻成为我国最重要的作物,因此确保玉米的稳产高产对保证我国粮食安全具有重要的意义。优良的玉米杂交种是影响玉米产量的重要因素之一。但在实际生产中,玉米杂交种还存在很多不足之处,例如有些品种不抗病虫害、易倒伏、产量低等,需要进行遗传改良。随着分子生物学的发展,转基因、基因编辑等技术将是作物遗传改良的主要方法。这两种技术都离不开高效的遗传转化体系。
自1996年第一例转基因作物被批准商业化生产以来,转基因作物的研发与生产得到了迅猛发展,全球转基因作物种植面积从1996年的170万公顷增加到2016年的1.851亿公顷。其中,玉米转基因技术研究始于1986年,到2016年全球单个作物种植面积中转基因玉米的应用率是26%。孟山都、先锋、先正达等国外大公司建立了工厂化流水线式的规模化的相对成熟的玉米转基因技术体系,国外公立机构实验室也实现了较高遗传转化效率玉米转基因技术体系。中国玉米转基因技术体系研究起步较晚,目前初步建立了玉米规模化转基因技术体系,但仍有必要进一步提高玉米遗传转化效率,同时应紧跟玉米转基因技术发展趋势,大力发展多基因转化技术、基因打靶技术及安全转化技术,使转基因玉米技术体系更好服务于玉米基因功能研究、有应用价值转基因玉米新材料产品研发和品种培育等。
尽管玉米规模化转基因体系已初成规模,但玉米转基因技术体系也仍存在许多不足需要完善。首先,基因转化效率低。尽管近些年农杆菌和基因枪等转化方法从多方面都有所提高和优化,但整体而言,玉米基因转化效率相对于水稻、拟南芥等仍然很低。其次,可用于转基因的玉米受体材料有限,农杆菌介导的玉米遗传转化受玉米基因型限制,它需要用于遗传转化的玉米受体材料具有很好的组培特性。目前国际上常用玉米杂交种HiII、玉米自交系B73及中国玉米优良自交系综31等为转化受体,这限制了转化方法在多种基因型应用的普及。除此之外,农杆菌侵染方法以及农杆菌与玉米幼胚共培养后对转化细胞的筛选所需要的选择压等一系列技术细节,都需要进一步的优化,来提高转化效率。在开发新的转化方法和优化已有的转化技术的过程中,需要易观察转化效率的标记基因,即在再生苗阶段就可以直接评价基因转化方法的优劣和转化效率的高低,而不依赖于分子特征的检测。
基因编辑技术产生于20世纪80年代,是一种利用工程核酸酶在体内精准编辑基因组的技术。目前应用最广的三类特异的工程核酸酶有锌指核酸酶(ZFN)、转录激活子内效应子核酸酶(TALEN)和规律成簇间隔短回文重复序列及其核酸酶(CRISPR/Cas9)。这三种技术,尤其是CRISPR/CAS9介导的基因编辑技术,已经开始在植物基因功能验证和遗传改良方面得到应用,但仍然存在一系列的问题需要改进。ZEN是最早被利用,但特异性低、效率不高、脱靶率大,并且操作繁琐、成本高。TALEN技术可以像ZEN一样完成对基因组的编辑,且相对简单费用低。相对于前两种方法,最新的编辑技术CRISPR/Cas9技术具有明显的优势,它的载体构建简单、成本更低,可以同时实现多靶点的定点修饰、靶向效率更高。但自CRISPR/Cas9技术投入使用至今,研究者们仍一直在探索提高该技术靶向效率的方法,如通过提高sgRNA的特异性、改造Cas9蛋白的结构降低脱靶效应以及选择构建合适的载体等。在这些方法的摸索过程中,同样需要在苗期就可以评价相应的突变效率的靶基因。
发明内容
本发明的目的是提供了与玉米白化相关的SCD基因的克隆及应用。
本发明首先提供了SCD基因编码的蛋白质(SCD蛋白质)的下述X1)或X2)的应用:
X1)在调控玉米白化中的应用;
X2)在玉米育种中的应用;
所述SCD蛋白质为如下A1)或A2)或A3):
A1)氨基酸序列是序列表中序列1的蛋白质;
A2)氨基酸序列是序列表中序列4的蛋白质;
A3)将序列表中序列1或序列4所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有相同功能的蛋白质。
为了使A1)或A2)中的蛋白质便于纯化,可在A1)或A2)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A2)中的SCD蛋白质,所述一个或几个氨基酸残基的取代和/或缺失和/或添加为不超过10个氨基酸残基的取代和/或缺失和/或添加。
上述A2)中的SCD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A2)中的SCD蛋白质的编码基因可通过将本发明的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。
本发明还提供了与所述SCD蛋白质相关的生物材料的下述X1)或X2)的应用:
X1)在调控玉米白化中的应用;
X2)在玉米育种中的应用;
所述生物材料为下述B1)至B7)中的任一种:
B1)编码所述SCD蛋白质的核酸分子;
B2)含有B1)所述核酸分子的表达盒;
B3)含有B1)所述核酸分子的重组载体、或含有B2)所述表达盒的重组载体;
B4)含有B1)所述核酸分子的重组微生物、或含有B2)所述表达盒的重组微生物、或含有B3)所述重组载体的重组微生物;
B5)含有B1)所述核酸分子的转基因植物细胞系、或含有B2)所述表达盒的转基因植物细胞系;
B6)含有B1)所述核酸分子的转基因植物组织、或含有B2)所述表达盒的转基因植物组织;
B7)含有B1)所述核酸分子的转基因植物器官、或含有B2)所述表达盒的转基因植物器官。
上述应用中,B1)所述核酸分子,即SCD基因,为如下b1)-b6)中的任一种:
b1)序列表中序列5或序列5的第317-6498位或序列5的第843-6193位所示的DNA分子;
b2)编码序列是序列表中序列6或序列6的第203-2443位的cDNA分子或DNA分子;
b3)序列表中序列2或序列2的第2196-8353位或序列2的第2722-8045位所示的DNA分子;
b4)编码序列是序列表中序列3或序列3的第204-2444位的cDNA分子或DNA分子;
b5)与b1)或b2)或b3)或b4)限定的核苷酸序列具有75%或75%以上同一性,且编码所述SCD蛋白质的cDNA分子或基因组DNA分子;
b6)在严格条件下与b1)或b2)限定的核苷酸序列杂交,且编码所述SCD蛋白质的cDNA分子或基因组DNA分子。
其中,所述核酸分子可以是DNA,如cDNA、基因组DNA或重组DNA;所述核酸分子也可以是RNA,如mRNA或hnRNA等。
本领域普通技术人员可以很容易地采用已知的方法,例如定向进化和点突变的方法,对本发明的编码SCD蛋白质的核苷酸序列进行突变。那些经过人工修饰的,具有与本发明分离得到的SCD蛋白质的核苷酸序列75%或者更高同一性的核苷酸,只要编码SCD蛋白质且具有SCD蛋白质功能,均是衍生于本发明的核苷酸序列并且等同于本发明的序列。
这里使用的术语“同一性”指与天然核酸序列的序列相似性。“同一性”包括与本发明的编码SCD蛋白质的核苷酸序列具有75%或更高,或85%或更高,或90%或更高,或95%或更高同一性的核苷酸序列。同一性可以用肉眼或计算机软件进行评价。使用计算机软件,两个或多个序列之间的同一性可以用百分比(%)表示,其可以用来评价相关序列之间的同一性。
上述应用中,所述严格条件是在2×SSC,0.1%SDS的溶液中,在68℃下杂交并洗膜2次,每次5min,又于0.5×SSC,0.1%SDS的溶液中,在68℃下杂交并洗膜2次,每次15min;或,0.1×SSPE(或0.1×SSC)、0.1%SDS的溶液中,65℃条件下杂交并洗膜。
上述75%或75%以上同一性,可为80%、85%、90%或95%以上的同一性。
上述应用中,B2)所述的含有编码SCD蛋白质的核酸分子的表达盒(SCD基因表达盒),是指能够在宿主细胞中表达SCD蛋白质的DNA,该DNA不但可包括启动SCD基因转录的启动子,还可包括终止SCD基因转录的终止子。进一步,所述表达盒还可包括增强子序列。
可用现有的表达载体构建含有所述SCD基因表达盒的重组载体。
上述应用中,所述载体可为质粒、黏粒、噬菌体或病毒载体。
上述应用中,所述转基因植物细胞系、转基因植物组织和转基因植物器官均不包括繁殖材料。
本发明还提供了所述SCD蛋白质或所述生物材料。
本发明还提供了构建玉米白化模型的方法,所述方法包括降低玉米中所述SCD蛋白质的含量和/或活性得到所述玉米模型。
上述方法可包括抑制编码SCD基因的表达。
上述方法可通过对编码所述SCD蛋白质的核酸分子进行突变实现。
上述方法中,所述突变可采用CRISPR/Cas9方法实现。
上述方法中,所述CRISPR/Cas9方法中所用靶序列为靶序列1和/或靶序列2;
所述靶序列1可为如下m1)、m2)或m3):
m1)序列表中序列6的第288-306位的核苷酸序列;
m2)与m1)限定的DNA序列具有75%或75%以上同一性的由m1)衍生的DNA序列;
m3)在严格条件下与m1)限定的DNA序列杂交的由m1)衍生的DNA序列;
所述靶序列2可为如下n1)、n2)或n3):
n1)序列表中序列6的第354-372位的核苷酸序列;
n2)与n1)限定的DNA序列具有75%或75%以上同一性的由n1)衍生的DNA序列;
n3)在严格条件下与n1)限定的DNA序列杂交的由n1)衍生的DNA序列。
所述核酸分子可为上文B1)所述核酸分子。
上述方法具体可采用表达含有靶向所述靶序列1和/或所述靶序列2的sgRNA和Cas9重组载体实现对编码所述SCD蛋白质的核酸分子进行突变。所述重组载体可为重组CRISPR/Cas9载体,所述重组CRISPR/Cas9载体可为通过BsaⅠ内切酶向pBUE411载体中***靶序列1和靶序列2所示的DNA片段得到的重组载体。
本发明中,所述白化可为叶片白化和/或全植株白化和/或玉米籽粒白化。所述白化可导致玉米苗期死亡。
所述玉米籽粒白化可表现为籽粒由黄色变为浅黄色;
所述玉米籽粒白化可为部分籽粒颜色变化。
实验证明,本发明发现SCD基因及其编码的蛋白质可以控制玉米籽粒和叶片双白化,并进一步验证了其功能,表明SCD基因及其编码的蛋白质可以用来调控玉米植株和叶片颜色以及籽粒颜色,并进一步可用于构建玉米籽粒和叶片双白化的模型。该基因可用于植物基因工程、基因编辑技术及分子育种中。
附图说明
图1为scd突变体的籽粒和叶片表型。
(A)SCD/scd杂合突变体的一个自交果穗。(B)野生型(WT)和scd纯合突变体的成熟籽粒表型。(C)野生型(WT)和scd纯合突变体的发苗7天后苗期植株表型。
图2为SCD基因的克隆。
(A)SCD基因精细定位在5号染色体的14-kb区间。黑色框代表SCD基因型;白色框代表scd基因型;灰色框代表杂合基因型。(B)SCD基因和***其中的7.6kb LTR反转录转座子片段的结构示意图。实线代表内含子;黑框代表外显子;白框代表非翻译区。***位点用倒三角表示。红色椭圆形框:TSR(重复靶点)区域;蓝色TG…CA框:LTR(长片段重复)反转录转座子的典型结构;橘色椭圆形框:PBS(引物结合位点)区域;绿色椭圆形框:In(整合酶)的核心区域;黄色椭圆形框:PPT(多嘌呤序列)区域。(C)SCD基因cDNA全长在授粉后15天胚乳和发苗7天苗期叶片中的PCR产物。(D)scd突变体中SCD基因多种转录本不同剪切模式的示意图。实线代表内含子;黑框代表外显子;白框代表非翻译区。5’UTR区域发生的120-bp***和8号与9号外显子之间发生的111bp-1593bp***用灰框表示。(E)SCD基因在授粉后15天胚乳和发苗7天的苗期叶片中的转录水平的相对表达量。(F)SCD基因在授粉后15天胚乳和发苗7天的苗期叶片中的蛋白水平。
图3为玉米突变体中SCD表达量和表型的检测。(A)SCD基因在3个scd等位突变体中的突变位点,倒三角代表的是突变位点。(B)3个scd等位突变体及其与scd突变体杂交所得F1的表型特征。WT代表野生型。(C)SCD在3个scd等位突变体中授粉后15天胚乳中的蛋白表达水平。
图4为SCD基因的时空表达分析。Em:胚;En:胚乳;DAP(day after pollination):授粉后天数;Root:根;Stem:茎;Leaf:叶;Seed:籽粒;WT:野生型玉米;scd:scd突变体。每个样本都做了3个独立的生物学重复。误差线代表平均值±标准误,**代表P<0.01,*代表0.01<P<0.05,使用的是Student’s t test。
图5为SCD蛋白的亚细胞定位。(A-D)为35S::EGFP空载体的表达,标尺为5μm;(E-H)为35S::SCD-EGFP融合蛋白载体的表达,标尺为5μm。
图6为利用CRISPR/Cas9方法敲除玉米B73中的SCD基因。A为SCD基因结构及靶序列和引物的位置,Target 1和Target 2分别表示靶序列1和靶序列2。B为三株阳性基因敲除玉米幼苗SCD基因序列,Target 1和Target 2分别表示靶序列1和靶序列2,N19为表示核苷酸个数为19个,WT表示野生型SCD基因中的序列,黑框中为PAM序列。C为3个T0代阳性基因敲除玉米植株的基因检测,Cas9基因型的检测结果用加号+表示出。D为SCD基因在3个T0代阳性基因敲除玉米中的相对表达量。E为三株阳性转基因玉米幼苗表型,WT为B73的愈伤组织得到的植株,箭头所示为白化叶片。
具体实施方式
下面结合具体实施方式对本发明进行进一步的详细描述,给出的实施例仅为了阐明本发明,而不是为了限制本发明的范围。下述实施例中的实验方法,如无特殊说明,均为常规方法。下述实施例中所用的材料、试剂、仪器等,如无特殊说明,均可从商业途径得到。以下实施例中的定量试验,均设置三次重复实验,结果取平均值。
下述实施例中的pBUE411载体(Bar)(Hui-Li Xing et al.(2014),A CRISPR/Cas9toolkit for multiplex genome editing in plants.BMC Plant Biology.14:327)公众可从申请人处获得该生物材料,该生物材料只为重复本发明的相关实验所用,不可作为其它用途使用。
下述实施例中的RIL105,为文献(Jihua Tang et al.2007,Gentic dissectionof plant height by molecular markers using a population of recombinant linesin maize.Euphytica.155:117-124)中294个RIL家系中的一家系,公众可从申请人处获得该生物材料,该生物材料只为重复本发明的相关实验所用,不可作为其它用途使用。
下述实施例中的B73(Hui-Li Xing et al.(2014),A CRISPR/Cas9toolkit formultiplex genome editing in plants.BMC Plant Biology.14:327)公众可从申请人处获得该生物材料,该生物材料只为重复本发明的相关实验所用,不可作为其它用途使用。
实施例1:scd基因的定位和克隆
1.研究群体的构建
发明人在繁殖Zong3×Yu87-1重组自交时,发现该群体的RIL105家系有一株系发生了自然突变(图1),将该突变体记为M0代突变体,M0代突变体得到的籽粒记为M1代,M1代籽粒中有部分籽粒颜色由黄色转变成浅黄色,其余颜色仍为黄色。M1代浅黄色籽粒(记为scd纯合突变体)种下得到的M1代玉米植株叶片白化,并在发苗两周后致死。进一步对M1代黄色籽粒自交得到的籽粒(M2代籽粒)进行观察,发现部分M1代植株(纯合)的M2代籽粒全为黄色,部分M1代植株(记为scd杂合突变体)的M2代籽粒为黄色和浅黄色,且黄色和浅黄色的种子呈现3:1分离(黄色籽粒:浅黄色籽粒,χ2<χ2 0.05=3.84;P>0.05;表2)。由此推断这一突变是由隐性单基因控制。以B73和M0代突变体后代中的杂合突变体SCD/scd为双亲,构建B73×SCD/scd的F2群体。
表2杂合RIL105突变果穗籽粒颜色分离比的卡方测验
2.scd基因的精细定位和基因克隆
通过用覆盖玉米10条染色体的366个标记在突变体和B73中筛选到7个多态性标记,扫描509株F2个体,结合这些个体的表型,将目标基因定位在第5号染色体上的两个Indel标记IDP7677和IDP411之间,该区段遗传距离为1.6cM,物理距离为1.86Mb。在此基础上,进一步在区段内开发新的标记筛选重组单株,将目标区间定位在约14kb范围内,包含两个基因GRMZM2G137409和GRMZM2G137399。将这14kb区段进行测序,发现在其中的一个基因GRMZM2G137409的第8号内含子上发生了7.6kb的长片段***。利用NCBI blast进行序列搜索,发现这7.6kb***片段与玉米中CRM着丝粒反转录转座子(AY129008.1)具有98%的相似性;同时也将这7.6kb***片段的序列,利用复旦大学研发的LTR finder网站(http://tlife.fudan.edu.cn/ltr_finder/)进行转座子结构分析。结果发现,这7.6kb***片段的序列含有LTR反转录转座子的典型特征。扩增SCD的cDNA全长,发现在授粉后15天的胚乳和发苗7天的叶片中,这个7.6kb***片段造成了SCD基因由一种剪切模式变成产生多种可变剪切的模式。通过克隆测序,发现这种多可变剪切模式产生至少6种可变剪切模式的转录本。对授粉后15天的胚乳和发苗7天的叶片scd突变体,进行表达分析,也发现SCD基因再转录水平和翻译水平的都发生剧烈的表达下调。这些结构说明,GRMZM2G137409就是SCD的目标候选基因(图2),表明,SCD基因与玉米白化相关,进一步SCD基因与玉米植株、叶片和籽粒的白化相关。RIL105中SCD基因编码序列表中序列1所示的SCD蛋白质,其cDNA为序列3,序列3的第204-2444位编码序列1所示SCD,SCD基因的基因组序列为序列2,序列2的第2196-8353位为自5′UTR至3′UTR的序列,第2722-8045位为SCD基因的外显子和内含子区域。
实施例2、SCD基因突变体的等位性测验
为进一步验证SCD为控制玉米籽粒和叶片双白化的基因,从美国玉米种质资源中心(Maize Genetics Cooperation Stock Center,MGCSC)获得3个突变体——518D lw2、512I lw2-vp12和523B lw2-UFMu-07766,分别记为scd-1、scd-2和scd-3突变体。
通过序列测定,发现:在scd-1突变体中的GRMZM2G137409基因的第12个外显子发生了由G到T的转换,具***置为序列2的第5809位(同时也是序列3的第1584位),使得氨基酸由谷氨酸转换成了终止密码子,造成翻译的提前终止;在scd-2突变体的GRMZM2G137409基因的第1个外显子上(序列2的第2738位和2739之间,同时也是序列3的第220位和第221位之间)发生了8bp的***,造成了移码突变;在scd-3突变体的GRMZM2G137409基因的第1个外显子上(序列2的第2772位和2773之间,同时也是序列3的第254位和第255位之间)发生了Mu转座子的***(图3)。
scd-1、sld-2和scd-3具有与scd突变体相同的表型,即杂合突变植株果穗出现浅黄色籽粒,并且黄色籽粒和浅黄色籽粒的分离比符合3:1的单基因隐性遗传的规律,杂合突变植株得到的浅黄色籽粒种下后得到的植株产生白化叶片,并导致植株死亡(图4)。表明,SCD基因与玉米白化相关,进一步SCD基因与玉米植株、叶片和籽粒的白化相关。
通过Western Blot也发现,在这3个突变体中SCD蛋白水平发生了极显著的下调。所用的Anti-SCD抗体为根据SCD蛋白序列设计的特异性兔多抗,其中所用抗原的序列是CIKDHGRWVDPPAE(即序列1的第733-745位)(注:C为cys,以便-SH偶联载体),由北京康为世纪生物科技公司合成和纯化,其制备方法为:人工合成抗原(10-14mg,纯度85%以上),将合成的抗原进行载体偶联,然后进行动物免疫和抗血清制备,待抗血清验证合格后进行抗原的亲和纯化得到抗体。另一个内参抗体Anti-ACTIN抗体为ABclonal公司的鼠单抗(AC009)
实施例3、SCD的时空表达分析
利用qRT-PCR对SCD在野生型不同组织不同时期的的表达进行分析,发现SCD在玉米发育过程中具有组成性表达的特征。转录水平分析发现:SCD在叶片中表达最高,其次是茎、根。对授粉后不同发育时期中的胚、胚乳和籽粒进行的转录水平表达,发现,SCD在3个组织中均呈现出随着发育时期的进行而呈波动性的表达:在授粉后15天和35天表达出现两次小高峰,在20天和30天表达相对较低。在SCD功能缺失的scd突变体中,其表达水平均呈现明显的下调,但其模式与野生型相似(图4)。
实施例4、SCD蛋白的亚细胞定位
构建表达SCD蛋白与enhanced green fluorescent protein(EGFP)的融合蛋白的载体:将序列3的第176-2441位所示的cDNA片段克隆到连接到PEZS-NL载体(Wei liang Zuoet al.(2015)A maize wall-associated kinase confers quantitative resistance tohead smut.Nat.Genet.47,151–157)上,菌落PCR鉴定并测序,获得35S启动子启动的35S::SCD-EGFP表达载体。
将该35S::SCD-EGFP表达载体,利用PEG化学转化法转化到玉米原生质体中。利用激光共聚焦显微镜进行观察,发现SCD在玉米原生质体的叶绿体中发生表达(图5)。
实施例5、利用CRISPR/Cas9方法敲除SCD基因构建玉米白化苗模型
本实施例通过构建含有SCD基因特异性的双gRNA靶点的重组CRISPR/Cas9载体敲除玉米SCD基因构建玉米模型。SCD基因的序列为序列表中序列5(序列5的第317-6498位为自5′UTR至3′UTR的序列,第843-6193位为SCD基因的外显子和内含子区域),其cDNA序列为序列6,序列6的第203-2443位编码序列4所示的SCD蛋白质。重组CRISPR/Cas9载体编码的sgRNA能特异识别SCD基因,识别序列为序列6的第288-306位(靶序列1)和序列6的第354-372位(靶序列2),靶序列1和靶序列2在基因中的位置如图6中A所示,重组CRISPR/Cas9载体能编码Cas9。重组CRISPR/Cas9载体的具体构建方法如下:
通过BsaⅠ内切酶向pBUE411载体中***两条DNA片段,得到重组载体,该重组载体即为重组CRISPR/Cas9载体。两条DNA片段的序列分别为靶序列1(5′-CGAAGGTCTTGTCTGCTCC-3′)和靶序列2(5′-TGGCGAAGAGCTCTAGTAC-3′)。重组CRISPR/Cas9载体表达含有靶向靶序列1和靶序列2的sgRNA。
将重组CRISPR/Cas9载体导入农杆菌EHA105中,得到重组农杆菌。将该重组农杆菌转染受体玉米品种B73的胚细胞,培养转染后的细胞,得到转基因玉米幼苗,该转基因玉米幼苗为对B73的SCD基因进行改造而其他DNA均未改造得到的植株。将pBUE411载体导入农杆菌EHA105后转染同种玉米胚细胞得到的植株作为对照植株。
利用引物对(F:TGGGTTAGGGCGGTATTG和R:CAGCTCCTTTATCTGCTATCCTC)分别对转基因玉米幼苗叶片cDNA进行PCR扩增,检测SCD基因中靶序列1和靶序列2的上下游序列,得到三株阳性基因敲除玉米幼苗,分别记为CR-scd-1、CR-scd-2和CR-scd-3,这三个植株靶序列1和靶序列2及其上下游序列图6中B所示。结果显示,这三个植株中的两条染色体中的SCD基因均发生了缺失突变,对照植株SCD基因的序列同受体玉米(野生型)一样未发生突变。这三个植株中的缺失突变均导致了SCD基因编码序列的变化,并进一步导致了编码的蛋白质序列的变化。
引物对(F和R)PCR产物的电泳结果如图6中C所示。利用引物对(CTGTTCGTCGAGCAGCACAAGCATT和TTCCCAATGCCATAATACTCAAACTCAG)检测三株阳性基因敲除玉米幼苗,结果(图6中C)显示,这三个植株中均含有Cas9的编码基因。
利用引物对(GTGCTACAGGGACTACGA和CATGCGATCTCCACCTTC)检测三株阳性基因敲除玉米幼苗及野生型玉米(受体玉米,B73)中SCD基因的表达量,结果如图6中D所示。结果显示,与野生型玉米相比,三株阳性基因敲除玉米幼苗中SCD基因的表达量均显著下降,对照植株SCD基因的表达量与野生型玉米无显著差异。
表型鉴定:三株阳性基因敲除玉米幼苗均表现为白化苗,以愈伤组织长出白色叶片记为第1天,CR-scd-1、CR-scd-2和CR-scd-3分别在第6、10和9天死亡,即白色叶片萎蔫枯萎;对照植株的表型与野生型玉米无显著差异。
<110> 中国农业大学
<120> 玉米籽粒类胡萝卜素缺陷基因SCD的克隆及应用
<160> 6
<170> PatentIn version 3.5
<210> 1
<211> 746
<212> PRT
<213> 玉米
<400> 1
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Gly Thr Asn His Gly Ser Leu Ser Asp Arg Ile Met Ser Tyr Tyr Gly
245 250 255
Asp Ser Pro Arg Gly Met Val Glu Ser Ala Leu Glu Phe Ala Arg Ile
260 265 270
Cys Arg Lys Leu Asp Phe His Asn Phe Val Phe Ser Met Lys Ala Ser
275 280 285
Asn Pro Val Ile Met Val Gln Ala Tyr Arg Leu Leu Val Ala Glu Met
290 295 300
Tyr Asn Leu Gly Trp Asp Tyr Pro Leu His Leu Gly Val Thr Glu Ala
305 310 315 320
Gly Glu Gly Glu Asp Gly Arg Met Lys Ser Ala Ile Gly Ile Gly Thr
325 330 335
Leu Leu Met Asp Gly Leu Gly Asp Thr Ile Arg Val Ser Leu Thr Glu
340 345 350
Pro Pro Glu Glu Glu Ile Asp Pro Cys Gln Arg Leu Ala Asn Leu Gly
355 360 365
Thr Gln Ala Ala Asn Leu Gln Ile Gly Val Ala Pro Phe Glu Glu Lys
370 375 380
His Arg Arg Tyr Phe Asp Phe Gln Arg Arg Ser Gly Gln Leu Pro Leu
385 390 395 400
Gln Lys Glu Gly Glu Glu Val Asp Tyr Arg Asn Val Leu His Arg Asp
405 410 415
Gly Ser Val Leu Met Ser Val Ser Leu Asp Gln Leu Met Ala Pro Asp
420 425 430
Leu Leu Tyr Arg Ser Leu Ala Ala Lys Leu Val Val Gly Met Pro Phe
435 440 445
Lys Asp Leu Ala Thr Val Asp Ser Ile Leu Leu Arg Glu Leu Pro Pro
450 455 460
Val Glu Asp Ala Glu Ala Arg Leu Ala Leu Lys Arg Leu Val Asp Ile
465 470 475 480
Ser Met Gly Val Leu Ala Pro Leu Ser Glu Gln Leu Thr Lys Pro Leu
485 490 495
Pro His Ala Ile Val Leu Val Asn Leu Asp Glu Leu Ser Ser Gly Ala
500 505 510
His Lys Leu Leu Pro Glu Gly Thr Arg Leu Ala Val Thr Leu Arg Gly
515 520 525
Asp Glu Ser Tyr Glu Gln Leu Asp Ile Leu Lys Asp Val Asp Asp Ile
530 535 540
Thr Met Leu Leu His Asn Val Pro Tyr Gly Glu Glu Lys Thr Gly Arg
545 550 555 560
Val His Ala Ala Arg Arg Leu Phe Glu Tyr Leu Gln Ala Asn Gly Leu
565 570 575
Asn Phe Pro Val Ile His His Ile Asn Phe Pro Glu Thr Ile Asp Arg
580 585 590
Asp Gly Leu Val Ile Gly Ala Gly Ala Asn Val Gly Ala Leu Leu Val
595 600 605
Asp Gly Leu Gly Asp Gly Val Phe Leu Glu Ala Ala Asp Gln Glu Phe
610 615 620
Glu Phe Leu Arg Asp Thr Ser Phe Asn Leu Leu Gln Gly Cys Arg Met
625 630 635 640
Arg Asn Thr Lys Thr Glu Tyr Val Ser Cys Pro Ser Cys Gly Arg Thr
645 650 655
Leu Phe Asp Leu Gln Glu Ile Ser Ala Glu Ile Arg Glu Lys Thr Ser
660 665 670
His Leu Pro Gly Val Ser Ile Ala Ile Met Gly Cys Ile Val Asn Gly
675 680 685
Pro Gly Glu Met Ala Asp Ala Asp Phe Gly Tyr Val Gly Gly Ala Pro
690 695 700
Gly Lys Ile Asp Leu Tyr Val Gly Lys Thr Val Val Gln Arg Gly Ile
705 710 715 720
Ala Met Glu Gly Ala Thr Asp Ala Leu Ile Gln Leu Ile Lys Asp His
725 730 735
Gly Arg Trp Val Asp Pro Pro Ala Glu Glu
740 745
<210> 2
<211> 11295
<212> DNA
<213> 玉米
<400> 2
ttggcgtgca gcttcctagc ttgggctgct tctgtgcagg ggtatgggct tcattttcat 60
ccgggctcac tcttggtcta ttcgggatgc actgtaccga ggcccatggt aagtggccta 120
accccaagtc cactgttctc acagttgcac actcacttga actatctgca ttcagcattc 180
tacagggcag gcagttcgat ccctcggtga gcgatgcagg cctgactgcc tgagtgacgc 240
tgacgcaggc aggaggcggc cacttctggt tctagacatg cctgacatgt actactgtta 300
tcttggacga aatcgtcact cctcttttct tccccttttt ctttgcacgg cctggtacgc 360
agcacgctct gtcgtgttat ttcgaaaagg caaaaaaaca tgaaagtttg gaccaggtta 420
tttatgaccg tccaatcacg acctaaaaaa taaaaacccg aatcatggtc cggcatctaa 480
aaaataaaaa cccgaagcat ggtccgccct aaatagccgt gttgtgctac gaataggccg 540
tgtttagacc gacatggcat gaaatgaaaa cttatgttta aatgactctt tcatgttcca 600
ataatgtgtt ctctctacaa agtttataga aaaatattga tacttataag catatataat 660
atacatagca gttttaagtc aagttgttag tcatagcaat acatttttaa aggctttctc 720
ttttcggata atttttaatg gcatagttat aaataatttg aaggtatata ttactcatgt 780
catgcttggt ccggtacgac atgtgcgtca tgttttaggg tttggtttgg ctaggcctag 840
ctttttacat gtgggcacga tatgatctaa cttgaaatta ttctgtgcca tactagttca 900
aaaaaaaaaa aaaaatttac cccaaagtat gatgaaccgg gactattctg gcacaaccaa 960
ctttcagctc tagcctgagg gaaccaaata aaaggaacat cactcttttg gacttcatta 1020
cttgcaaaac ggtcagtctt tgctcgtgcc ctggaagtcc caatcgcgga acacgtgtta 1080
aaaattcttc tcccacatgg gataggtggt ttggacgaac aattaagtga cggcacataa 1140
tgtgttttcg acgtttattt tgggtcaata catgtggcgt gacttgattt ccggtctacg 1200
ctcgtcatac agcgcaaaca catgttgacc tgtctgtacc taggctagcc gccgtgggtg 1260
ggagcgaccg agggagttcg tccaagtcgt ccaggaaaaa tacaacagga aattgagcag 1320
atgggtgacc aatgaccagg ggcgcacgtt catcttacct tgaacatgtg ggcttcccaa 1380
ctgacaacac gagcactgtt actacaaccg aagcagcaga ctatgtaggc catgtctaga 1440
ttttatgtgt aaagtatgag ctaaactttg gatcatttta gtttattact ttttagggtc 1500
taataaacta aaactccttt ttaggttagc taaattttaa ccgctcattt atttgaagtc 1560
tcaagagata atataatcct aaatttaata gactatataa acgctgctgt tttttttttt 1620
tggttcttcg taaaaccacg gcttgtgtct catgggaaag cgacggacac gataaaatca 1680
tggcgggcag cgccaggaac tcggccctcg gccgccgcgc ctccccgcat ccggtccaca 1740
gcacccgcga gtattaagcc gaagcgtcag tggcacggca ccgcgccggt gtacatagac 1800
cgggtgcacg ctgtggttca gatggcccct gctgctcgga gataaaaggc gctatgccgc 1860
tatccagtag aaacccacca cgaactggcg ttccactgcc gagcccttcc tcctcttcct 1920
cttcttctga ccccccgctg gcttggaaaa agggagagaa aactcgcgca ctcggttatc 1980
gaaagggagg agcgcgggcg agggtgaggt gtcgcccaca cggagctgcg aggttcggtg 2040
atattttctt ctaccttgct gcccgcctcc gatgtccgat cgcggtcgcc gtgctgtgcg 2100
gccgccgggg cgggttccgt gctcggttta gctgtgaatt attctccctt tcagtctgga 2160
caggtagctg aacggctgaa ccagtggtgg atgctgggtc tgtggttggg gaatttgtgt 2220
tgtgtggatg ggaggatttt gaggtgcacc agctcacagt cgtcggtctg tcaggttgca 2280
gctccatcga attcagtaga atcctccttt gggttagggc ggtattgagc cggccattgc 2340
tcccgttcgt ttgccaataa ggatctacgc cgcttctcgt cttcggattt acttctcctc 2400
tccttgtttc cttcacccgg gaacggcggt cgattcgagt tcgctgaatt ttttatgcgg 2460
tttactatgc gaaacatgtt gtcagcttca ctgcgcgaat tatgctgtcc atttcactgc 2520
acgaattgtt acatgctctg gtttagtagc acacaccata tttgtcctgt tctgtctctt 2580
ccagcgatag ggtatattcc acttgtgccg gttaaaggcc accaggatca gcagccagcg 2640
gtcacctcac ccactcgtgt tggttgctct gcaggtgttt gtaggatctc ctaggtgagc 2700
ccctgctgct tggagacagc catggccacc ggcgtggctc cagctcctct cccacatgtc 2760
agagtgcgtc atgggggcgt cgggttcacc aggagcgtcg attttgcgaa ggtcttgtct 2820
gctcccggtg ccggcacgat gagagcaagc tcctctagag gcagggcgct cgtggcgaag 2880
agctctagta ctggctcgga gaccatggag ctcgagccat cttcagaagg aagcccactt 2940
ttaggtataa tctgcgtgct cttgtgctct tgtgcccttg cacgtgtatt tttcaaggca 3000
agggatctgt cccacgttta atctttgcac tcctttccta actggatgaa ctgtcctttc 3060
cgcgtccttc agtacccagg cagaagtact gtgaatcaac acaccagaca aggaggagga 3120
aaactcgaac tgtgatggtg gggaatgtgc cacttggcag tgatcacccc ataaggattc 3180
aaaccatgac gacttcagat accaaggatg ttgcgaaaac agtagaggag gtgcatccca 3240
tcccaaattc atgttaactg ttcttagtgt tctctacaca cgtgggagat gaattactca 3300
gcctttgtca gatgatttct attttgtgta tatttaacta atgaacatat cacttgcacc 3360
gtttcaggtg atgaggatag cagataaagg agctgatctt gttagaataa cagtccaggg 3420
taggaaggaa gctgatgcct gctttgagat caagaacact ctggttcaga agaagtaagg 3480
tgccattttt cttcacattc actggggttc acatgcatgg gtcctcaccc ttttcattgt 3540
ttttgtatag ttacaacatt ccactagtgg ccgatattca ttttgctcct acggtagctc 3600
taagggtggc agaatgtttt gacaaaattc gtgtgaaccc aggaaatttt ggtgagtgaa 3660
atcttgacat ccctcatttt agtctgtcat atttttgtta ttgtaaagtg ctgagaactg 3720
tactggcagc tgatcgtcgt gctcaatttg aaaagctgga atatactgac gacgactacc 3780
aaaaagagct agagcatatt gagaaggtcg aaaaattgtc actctagcct tcaaaaaaat 3840
ttcactctta gttttgttac aatcaatgtg aacatatggc atgttgtctt acagagccat 3900
gttaattatg tctcaggtgt tttctccatt agtcgagaaa tgcaagcagt atggaagagc 3960
aatgcgaata ggaacaaatc atggtagtct ttctgaccgc ataatgagct actatggtga 4020
ttctccacgg ggaatggtga ttatctcatt tctgatgatc tacttaaata ttagcacata 4080
aaagtttgac atcttctgca caatgttgca ggttgagtct gctttggaat ttgctagaat 4140
atgtcggaag ttggacttcc acaactttgt attttcgatg aaagctagta accctgttat 4200
catggtccaa gcatatcgcc tgcttgtggc agaaatgtat aacctaggat gggattatcc 4260
cttgcacttg ggtgttacag aagctggaga gggtgaagat ggaaggatga aatctgctat 4320
tggcattggg acactgctaa tggtaattga ttcactactt atttatgctg tatgaaattg 4380
ttaatctgga aacgatgtta acgacatgga atagtagcct catggaatgg taattgcttc 4440
ttttggagct tatagctgga agctttgaga atgaatattt ttaatttttt ttgctattca 4500
gttccatatc atgatatgtt gtgatgccaa attgacaaaa accctagaat caaatagccg 4560
cctgaaaaat gacgtgttta agattattat actttgtttt gtggattaac tgattgggac 4620
atttgccatg ctacgacaca tactagagta caagactatc tgtttcctgc aaaccaattg 4680
aactatggaa actggcaaat tctacaaaaa acacggacta tatcttctgt gtcttctatt 4740
acaggatggt ttgggtgata caatccgtgt ctccctcaca gaaccaccag aagaagagat 4800
tgatccttgc caaaggttgg caaatcttgg gacgcaggct gcaaaccttc aaattggggt 4860
ggtaatggac tttcactttc catactttgc agtcaccata tagtaaatac atgtccttac 4920
tatttgtgtt tcccttaagc atttatttaa ttctcttgga ttttgtatta ttttatcagg 4980
ctccatttga agaaaagcac aggcgttatt ttgatttcca gcgtaggagt ggtcaattgc 5040
ctttgcagaa ggaggtttgt taaaaataat tccacagttt gatcataaga actacactaa 5100
tggtaggttt cttatcggtt gctcatgctc ttatgacagg gtgaggaagt tgactacaga 5160
aatgtcctgc atcgtgatgg ttctgtactg atgtcagttt ccctggatca gttgatggta 5220
atctgcacct tttctattcc tttttgctga gttctggaat acataatata acacaccaaa 5280
agggatttca ccatggggtt ttgatttggg atccatacct agtagttcaa attaatcctc 5340
tggcaggatt cgaacccgag acctctgact ttgttaccat attactaacc acgagattga 5400
caatcatcat ctgtccatag ggactgtagt atatatacag gccagcatgc cttagcaata 5460
ggatgagcct aatgggcctt aacagttagt agtaccattg ttgttttcca tgcatttttc 5520
tattgaaaat tggttgttac tccaaataaa ctttgcagtg accactctat tgattagcca 5580
tctgaataaa tgtgctctgt agtgattaat gctcccttta tttgccaggc tcctgatctc 5640
ctttataggt ctcttgcagc aaagcttgtg gttggcatgc ctttcaaggt ctgttctatg 5700
ttttttgtgt gctagtacac gactattttt agatggattt tagttgaagt tgattattac 5760
tcctatggct accaggatct ggctactgta gattccattc ttttgaggga gctcccccct 5820
gtagaagatg ctgaagctgt gagttgcagc tacatttcta tccctctctc aagtgcatat 5880
gttattcgtt ttattccttc tttagagctc tgctctaatt ttgtgaatct gtacagaggc 5940
ttgcactcaa aagattagtt gacatcagca tgggcgtgct ggccccctta tcggagcagt 6000
tgacaaagcc actcccacat gcaattgtac ttgtcaacct cgacgaattg tcaagtggtg 6060
cacacaaact tttgccagaa ggtagttatt taaattaggc catgtcatta ttgtttcttt 6120
gctaatagtt taaggacaaa tatttcaccg aagacaaata atagaaaatt gaacttttta 6180
gtttacaata gaaaaaggta ctcgctccga tctgatccaa tataaagatt gtttctagtt 6240
tttcatacgt caaacttttc taactttggc aaagttcatt gaaaaacgcc ccaaacatct 6300
acaacatcaa gttatttcat taaatccacc ctgaaatata tcttgataat tcagtttgat 6360
attgtagatg ttaatatatt tttcttataa actcggtcaa agttagagaa gttttgactt 6420
acgataaaac taaaatgact tacattttgg aatggagtga gtgttgtttt gcctaatctc 6480
atcattttgt ccttttttat ggtgtgtata ggcactagac tagctgtcac tcttcgtggt 6540
gatgaatcat acgagcagct agatattctt aaggatgttg atgatataac aatgttgtta 6600
cataatgttc catatggtga ggagaagaca ggcagggtgc atgctgctag gaggtaagaa 6660
aacacagtag tcaattacta ttaaatgaga gatttttagg attcatttat ccaccatgct 6720
caacagcact ttgagtcttt tttgtccttt taggtttaac atatttagta tttttttatg 6780
tcttgcaggt tatttgagta cttacaggcc aatggcttga acttccctgt aattcatcac 6840
ataaatttcc ctgaaaccat tgacaggtga ctgtcaatat tcctattttc ttccccttgg 6900
aattgtgagt cgtggctttc ttgtttattc taattcaact atgtgcagag atggtcttgt 6960
cattggtgct ggggccaacg ttggtgctct cttagtcgat ggtcttggtg atggtgtatt 7020
ccttgaagct gctgaccagg aatttgagtt tctgagggac acatctttca acttgctcca 7080
aggttgcagg atgcgcaaca caaaaactgt aggttcatgt ttgtttggtt tatttaatga 7140
tatggctacc tctttctgta tatctggtgg acaacatagg gataccgcta cttgcagtaa 7200
aaaaaaagaa aaagggaaaa gaaacagcga aacaaaaaaa aatatctgtt gatgtttctc 7260
tctgagatga gaattgattt tgtgagaagt gattttctga ctttgaatgc tgtaggaata 7320
tgtgtcttgt ccttcctgcg gccgaacact ctttgacctt caggaaatca gcgctgagat 7380
tagagaaaag acctctcatc tgccaggtgt ctcggtgagt tttctgtata tggctctttc 7440
aaaaaagatc tactgtaagt ctgtatatat gtatttcctt cttttctatc aattatactg 7500
atcaaccgta tacatatttg cctggcagat cgctatcatg ggctgtattg tcaatggacc 7560
aggagagatg gctgatgccg attttgggta cgttggaggc gctcccggaa agatcgacct 7620
ttacgtcggc aaggtaatca ttccgttgct tggtttccaa gatcatgtac tccctgtgtt 7680
ccgaattata agacattttg acttctctaa atatattgtt tttgttatat atatagacat 7740
atttatatct agatgcatat aaaagctatg tatctagaaa agtaaaaaac atcaaataat 7800
ttggaatgga gggagcatat gcaatggaaa gttttactat gttggccaga actgcctggt 7860
tcagttagca ctacacaaat gtggttatga tcccgtgata gaacaattct ttccgctaaa 7920
acagtgcata tttctttcta cagaccgttg tgcagcgcgg catcgccatg gaaggtgcca 7980
ctgacgcctt gatccagctg atcaaggacc atggccgatg ggtggatcct cctgccgagg 8040
agtagggctg taggctgtag ccaagtctgt agtttatata ttctgtaaag cgaaacacgc 8100
aatgcagtaa ctgagcatac gatgtacatc atccctacac attttgtaca tttattgaga 8160
tctgagaaat aaactagaaa ccgtcggtga cggcatgtct gcagcacccc aaatgaagca 8220
catacgattt acatattagt agtagcccat gtgagcaaga cacaatccta tacaatcttg 8280
aagtctgcat tatatatttt ttccaacaga ggtagcttag aaagtttttc gagcaatgca 8340
attttgttcg ttactttccg tttacttttt ctttttttta gcgttgaggg ttatataaat 8400
agatttataa tacaaagaga aaaacggtcg tgattacaca acaccaaaat ttttattgtg 8460
gtcttgagag tttgtgatga tctattcata tccttactag agcattaatc atcgtcatat 8520
aagagttcta gctggcagcc tgtcattacc agggtgtaaa gataagatta tgctagtatc 8580
tgtgctaatt tccattcttg tttggtttgt gaaaccaatt tgctctgtct tttctgacat 8640
agatttgcta tgtgcttgtt aagaccgact ctggttggcg agtcagttgg cttaactagt 8700
gatcatttgt aacacttcgg tttgctccac tactgatatg ctcatcccat caattgaacg 8760
tgcgatattg tggagaacct agctgaatgg ggatggctca gctgctgttt agtttcctcg 8820
ccacttccat agcatcggat ggcaaacgtt gcggtcaggg cttcaagcct aacaaatctg 8880
cagtgataca taaccactat gaaccagtat tttttaacca gatatgcaag agtgttgtat 8940
atcattatat taaaatagaa tgatagtatc aaaacgcaaa cacatcatgt gactatcttc 9000
aacatccaac agactctcta tttcactcca tattttattc actattttaa attctactat 9060
gcaaacagtg caaaaacagt actttgtatg accgtttaca cagcctatag gagatgacta 9120
tcctagtgca aacataacct gtgctatggt gaatagacaa cccatttata tttcgaaaaa 9180
tcttatgcat gccatcgatt tttataccca ttctttgtgt gctattgaat gacaactaaa 9240
ggataatttt tatgtgtatg atatgtaggc tagtggcata ttacgcagtc agtatgtcac 9300
tgtcgccaac ctaccaaagg cgacgaacaa tttggcctgg tcctcgtgtt gctgggcaag 9360
aattccaagg gcatctccga gttgcagagc ttgatgtcgg agatgcgagc cttgtgtcca 9420
gatatcaacg tttggaaacc tcaagtggac aatcgtgttg atgagctgga gaacgtcgcg 9480
cgtgatgtca gcgagtgggt ggaggtaaaa cgctcggcag tcggtacatg gaaggatcag 9540
gagggggagg gtggattggg ttttcctaaa aattctcata ataattaaaa cctaagaaca 9600
aattcagctt caactgattg tctaatgtgt taataaagag ttatgcaccc taacttcaac 9660
ccatgctatc gtgttaattc tagagaagta aaaatccaaa gtaaatttct caaataaaat 9720
agcgtaacgt aaataagggt ttagaagact caagtttttg atgaggttct caaggagttg 9780
gctttccaca ctaatccttg ttagagcact cacagaaggg tattgcttct tagtcagtgt 9840
aatatcaagt actctaggga tggcaacggg ttgagtttat gtcgggtgca gtggagcaaa 9900
atctcgtccg cgatcacacc agtgaagtct acttaaaagt taaacccgag cgcgatgaca 9960
cttgcgggtg caatttcaaa cccgcactag gtcacccgtg agtttttact cagtatacac 10020
ttttaaacaa tagttcaact ataagcaatt aaatagagca ataattgagt tatatatata 10080
catattgtaa gcattctcac gttgagcagc aacaaaatct ttcatgaatc attagccaag 10140
ttattcattc aaaataatta ctattgtatc ttaatcattt ttcgtaaaat aaagaatgaa 10200
ccaaattttg ggttgggtgc gggccacccg cgagtagaaa tagaaactca cacccatatc 10260
cgcgaaactt tgtgaaaggg aaataggctt acaccttttc ctaaatgatt ttggtggttg 10320
aattgcccaa ctcaaataat tggactaact agtttgctct agatcataag ttctacaggt 10380
gccaaaggtt caacacaaac caataaaaag tccaagaaag ggttcaaata aaaagagcaa 10440
aagacaaccg aagactgccc tggtctggcg caccggactg tccggtgtgc caccggacag 10500
tgtccggtgc acctccggac agtgtccggt gcaccaggga gatcaactcc aaacttgcca 10560
ccttcgggaa ttctgggagc cactccacta taattcaccg gactgtccga tgtgccaagc 10620
ggagcaacag ctacctgcgc caacggtcgt ctgcaacagc tacaggaaca gtgaacagtg 10680
cgcgactgcg cgcgtagagt cagagcaggc gctagatggc gcaccggata gtgaacagaa 10740
gctgtccggt gcaccaccgg actgtccggt ggcccacatg tcagaagctc caacggttga 10800
actctaatgg ttgggtgacg tggctggcgc accggactgt ccagtgcgcc atacgacaac 10860
agccttcccc aacgaccatt ttggtggttg gagctataaa tacccccaac caccactctt 10920
caagacatcc aagcattcat tagtcctcat tcaataaaag agcaatacac aacactccaa 10980
gacacaaatc aaagccaccg atcgaatcaa agtccacaat tcaactctag tttttaggac 11040
ttgtgagaag atcgacttgt gttcttttgt tgttcttgtt gcttggttgg ccttccttct 11100
ttctctcatt cttgctctca aagccttgta agcgaagcaa gagacaccaa ttgtatgatg 11160
gtccttgtgg ggtctaagtg actcgggaaa tcaaggaagg aagctcactc ggtctaagtg 11220
accgtttgag agagggaaag ggttgaaaga gacccggtct ttgtgaccac ctcaacgggg 11280
agtaggttgc aagaa 11295
<210> 3
<211> 2609
<212> DNA
<213> 玉米
<400> 3
accacgaact ggcgttccac tgccgagccc ttcctcctct tcctcttctt ctgacccccc 60
gctggcttgg aaaaagggag agaaaactcg cgcactcggt tatcgaaagg gaggagcgcg 120
ggcgagggtg aggtgtcgcc cacacggagc tgcgaggtgt ttgtaggatc tcctaggtga 180
gcccctgctg cttggagaca gccatggcca ccggcgtggc tccagctcct ctcccacatg 240
tcagagtgcg tcatgggggc gtcgggttca ccaggagcgt cgattttgcg aaggtcttgt 300
ctgctcccgg tgccggcacg atgagagcaa gctcctctag aggcagggcg ctcgtggcga 360
agagctctag tactggctcg gagaccatgg agctcgagcc atcttcagaa ggaagcccac 420
ttttagtacc caggcagaag tactgtgaat caacacacca gacaaggagg aggaaaactc 480
gaactgtgat ggtggggaat gtgccacttg gcagtgatca ccccataagg attcaaacca 540
tgacgacttc agataccaag gatgttgcga aaacagtaga ggaggtgatg aggatagcag 600
ataaaggagc tgatcttgtt agaataacag tccagggtag gaaggaagct gatgcctgct 660
ttgagatcaa gaacactctg gttcagaaga attacaacat tccactagtg gccgatattc 720
attttgctcc tacggtagct ctaagggtgg cagaatgttt tgacaaaatt cgtgtgaacc 780
caggaaattt tgctgatcgt cgtgctcaat ttgaaaagct ggaatatact gacgacgact 840
accaaaaaga gctagagcat attgagaagg tgttttctcc attagtcgag aaatgcaagc 900
agtatggaag agcaatgcga ataggaacaa atcatggtag tctttctgac cgcataatga 960
gctactatgg tgattctcca cggggaatgg ttgagtctgc tttggaattt gctagaatat 1020
gtcggaagtt ggacttccac aactttgtat tttcgatgaa agctagtaac cctgttatca 1080
tggtccaagc atatcgcctg cttgtggcag aaatgtataa cctaggatgg gattatccct 1140
tgcacttggg tgttacagaa gctggagagg gtgaagatgg aaggatgaaa tctgctattg 1200
gcattgggac actgctaatg gatggtttgg gtgatacaat ccgtgtctcc ctcacagaac 1260
caccagaaga agagattgat ccttgccaaa ggttggcaaa tcttgggacg caggctgcaa 1320
accttcaaat tggggtggct ccatttgaag aaaagcacag gcgttatttt gatttccagc 1380
gtaggagtgg tcaattgcct ttgcagaagg agggtgagga agttgactac agaaatgtcc 1440
tgcatcgtga tggttctgta ctgatgtcag tttccctgga tcagttgatg gctcctgatc 1500
tcctttatag gtctcttgca gcaaagcttg tggttggcat gcctttcaag gatctggcta 1560
ctgtagattc cattcttttg agggagctcc cccctgtaga agatgctgaa gctaggcttg 1620
cactcaaaag attagttgac atcagcatgg gcgtgctggc ccccttatcg gagcagttga 1680
caaagccact cccacatgca attgtacttg tcaacctcga cgaattgtca agtggtgcac 1740
acaaactttt gccagaaggc actagactag ctgtcactct tcgtggtgat gaatcatacg 1800
agcagctaga tattcttaag gatgttgatg atataacaat gttgttacat aatgttccat 1860
atggtgagga gaagacaggc agggtgcatg ctgctaggag gttatttgag tacttacagg 1920
ccaatggctt gaacttccct gtaattcatc acataaattt ccctgaaacc attgacagag 1980
atggtcttgt cattggtgct ggggccaacg ttggtgctct cttagtcgat ggtcttggtg 2040
atggtgtatt ccttgaagct gctgaccagg aatttgagtt tctgagggac acatctttca 2100
acttgctcca aggttgcagg atgcgcaaca caaaaactga atatgtgtct tgtccttcct 2160
gcggccgaac actctttgac cttcaggaaa tcagcgctga gattagagaa aagacctctc 2220
atctgccagg tgtctcgatc gctatcatgg gctgtattgt caatggacca ggagagatgg 2280
ctgatgccga ttttgggtac gttggaggcg ctcccggaaa gatcgacctt tacgtcggca 2340
agaccgttgt gcagcgcggc atcgccatgg aaggtgccac tgacgccttg atccagctga 2400
tcaaggacca tggccgatgg gtggatcctc ctgccgagga gtagggctgt aggctgtagc 2460
caagtctgta gtttatatat tctgtaaagc gaaacacgca atgcagtaac tgagcatatg 2520
atgtacatca tccctacaca ttttgtacat ttattgagat ctgagaaata aactagaaac 2580
cgtcggtgac ggcatgtctg caacacccc 2609
<210> 4
<211> 746
<212> PRT
<213> 玉米
<400> 4
Met Ala Thr Gly Val Ala Pro Ala Pro Leu Pro His Val Arg Val His
1 5 10 15
His Gly Gly Val Gly Phe Thr Arg Ser Val Asp Phe Ala Lys Val Leu
20 25 30
Ser Ala Pro Gly Ala Gly Thr Met Arg Ala Ser Ser Ser Arg Gly Arg
35 40 45
Ala Leu Val Ala Lys Ser Ser Ser Thr Gly Ser Glu Thr Met Glu Leu
50 55 60
Glu Pro Ser Ser Glu Gly Ser Pro Leu Leu Val Pro Arg Gln Lys Tyr
65 70 75 80
Cys Glu Ser Thr His Gln Thr Arg Arg Arg Lys Thr Arg Thr Val Met
85 90 95
Val Gly Asn Val Pro Leu Gly Ser Asp His Pro Ile Arg Ile Gln Thr
100 105 110
Met Thr Thr Ser Asp Thr Lys Asp Val Ala Lys Thr Val Glu Glu Val
115 120 125
Met Arg Ile Ala Asp Lys Gly Ala Asp Leu Val Arg Ile Thr Val Gln
130 135 140
Gly Arg Lys Glu Ala Asp Ala Cys Phe Glu Ile Lys Asn Thr Leu Val
145 150 155 160
Gln Lys Asn Tyr Asn Ile Pro Leu Val Ala Asp Ile His Phe Ala Pro
165 170 175
Thr Val Ala Leu Arg Val Ala Glu Cys Phe Asp Lys Ile Arg Val Asn
180 185 190
Pro Gly Asn Phe Ala Asp Arg Arg Ala Gln Phe Glu Lys Leu Glu Tyr
195 200 205
Thr Asp Asp Asp Tyr Gln Lys Glu Leu Glu His Ile Glu Lys Val Phe
210 215 220
Ser Pro Leu Val Glu Lys Cys Lys Gln Tyr Gly Arg Ala Met Arg Ile
225 230 235 240
Gly Thr Asn His Gly Ser Leu Ser Asp Arg Ile Met Ser Tyr Tyr Gly
245 250 255
Asp Ser Pro Arg Gly Met Val Glu Ser Ala Leu Glu Phe Ala Arg Ile
260 265 270
Cys Arg Lys Leu Asp Phe His Asn Phe Val Phe Ser Met Lys Ala Ser
275 280 285
Asn Pro Val Ile Met Val Gln Ala Tyr Arg Leu Leu Val Ala Glu Met
290 295 300
Tyr Asn Leu Gly Trp Asp Tyr Pro Leu His Leu Gly Val Thr Glu Ala
305 310 315 320
Gly Glu Gly Glu Asp Gly Arg Met Lys Ser Ala Ile Gly Ile Gly Thr
325 330 335
Leu Leu Met Asp Gly Leu Gly Asp Thr Ile Arg Val Ser Leu Thr Glu
340 345 350
Pro Pro Glu Glu Glu Ile Asp Pro Cys Gln Arg Leu Ala Asn Leu Gly
355 360 365
Thr Gln Ala Ala Asn Leu Gln Ile Gly Val Ala Pro Phe Glu Glu Lys
370 375 380
His Arg Arg Tyr Phe Asp Phe Gln Arg Arg Ser Gly Gln Leu Pro Leu
385 390 395 400
Gln Lys Glu Gly Glu Glu Val Asp Tyr Arg Asn Val Leu His Arg Asp
405 410 415
Gly Ser Val Leu Met Ser Val Ser Leu Asp Gln Leu Lys Ala Pro Asp
420 425 430
Leu Leu Tyr Arg Ser Leu Ala Ala Lys Leu Val Val Gly Met Pro Phe
435 440 445
Lys Asp Leu Ala Thr Val Asp Ser Ile Leu Leu Arg Glu Leu Pro Pro
450 455 460
Val Glu Asp Ala Glu Ala Arg Leu Ala Leu Lys Arg Leu Val Asp Ile
465 470 475 480
Ser Met Gly Val Leu Ala Pro Leu Ser Glu Gln Leu Thr Lys Pro Leu
485 490 495
Pro His Ala Ile Val Leu Val Asn Leu Asp Glu Leu Ser Ser Gly Ala
500 505 510
His Lys Leu Leu Pro Glu Gly Thr Arg Leu Ala Val Thr Leu Arg Gly
515 520 525
Asp Glu Ser Tyr Glu Gln Leu Asp Ile Leu Lys Asp Val Asp Asp Ile
530 535 540
Thr Met Leu Leu His Asn Val Pro Tyr Gly Glu Glu Lys Thr Gly Arg
545 550 555 560
Val His Ala Ala Arg Arg Leu Phe Glu Tyr Leu Gln Ala Asn Gly Leu
565 570 575
Asn Phe Pro Val Ile His His Ile Asn Phe Pro Glu Thr Ile Asp Arg
580 585 590
Asp Gly Leu Val Ile Gly Ala Gly Ala Asn Val Gly Ala Leu Leu Val
595 600 605
Asp Gly Leu Gly Asp Gly Val Phe Leu Glu Ala Ala Asp Gln Glu Phe
610 615 620
Glu Phe Leu Arg Asp Thr Ser Phe Asn Leu Leu Gln Gly Cys Arg Met
625 630 635 640
Arg Asn Thr Lys Thr Glu Tyr Val Ser Cys Pro Ser Cys Gly Arg Thr
645 650 655
Leu Phe Asp Leu Gln Glu Ile Ser Ala Glu Ile Arg Glu Lys Thr Ser
660 665 670
His Leu Pro Gly Val Ser Ile Ala Ile Met Gly Cys Ile Val Asn Gly
675 680 685
Pro Gly Glu Met Ala Asp Ala Asp Phe Gly Tyr Val Gly Gly Ala Pro
690 695 700
Gly Lys Ile Asp Leu Tyr Val Gly Lys Thr Val Val Gln Arg Ala Ile
705 710 715 720
Ala Met Glu Gly Ala Thr Asp Ala Leu Ile Gln Leu Ile Lys Asp His
725 730 735
Gly Arg Trp Val Asp Pro Pro Ala Glu Glu
740 745
<210> 5
<211> 6498
<212> DNA
<213> 玉米
<400> 5
acgaactggc gtgccagtgc cagtgccgag cgcccgagcc cttactcctc ttcttccgac 60
cccccgctgg cttggaaaaa aggggaagaa aactcgcgca gtcggtcacc gaagggagga 120
gcgcggtcga ggtgccgccc acacggagct gcgaggtgcg gtgatatttt ctaccttgct 180
gcccgcatcc gatgtccgat cgcgcggtcg tcgtgctgtg cggctggggc gggattcgtg 240
atcgttttag ctgtgtggat tattctcccg ttcagcctgg gcaggtagct gaacggctga 300
accagtggtg gatgctgggt ctgtggttgg ggaatttgtg ttgtgtggat gggaggattt 360
tgaggtgcac cagctcacag tcgtcggtct gtcaggttgc agctccatcg aattcagtat 420
aatcctcctt tgggttaggg cggtattgag ccggccattg ctcccgttcg tttgccaata 480
aggatctacg ccgcttctcg tcttcggatt tacttctcct ctccttgttt ccttcacccg 540
ggaacggcgg tcgattcgag ttcgctgaat tttttatgcg gtttactatg cgaaacatgt 600
tgtcagcttc actgcgcgaa ttatgctgtc catttcactg cacgaattgt tacatgctct 660
ggtttagtag cacacaccat atttgtcctg ttctgtctct tccagcgata gggtatattc 720
cacttgtgcc ggttaaaggc caccaggatc agcagccagt ggtcacctca cccactcgtg 780
ttggttgctc tgcaggtgtt tgtaggatct cctaggtgag cccctgctgc ttggagacag 840
ccatggccac cggcgtggct ccagctcctc tcccacatgt cagagtgcat catgggggcg 900
tcgggttcac caggagcgtc gattttgcga aggtcttgtc tgctcccggt gccggcacga 960
tgagggcaag ctcctctaga ggcagggcgc tcgtggcgaa gagctctagt actggctcgg 1020
agaccatgga gctcgagcca tcttcagaag gaagcccact tttaggtata atctgcgtgc 1080
tcttgtgccc ttgcacgtgt atttttcaag gcaagggatc tgtcccacgt ttaatctttg 1140
cactcctttc ctaactggat gaactgtcct ttccgcgtcc ttcagtaccc aggcagaagt 1200
actgtgaatc aacacaccag acaaggagga ggaaaactcg aactgtgatg gtggggaatg 1260
tgccacttgg cagtgatcat cccataagga ttcaaaccat gacgacttca gataccaagg 1320
atgttgcaaa aacagtagag gaggtgcatc ccatccgaaa ttcatgttaa ctgttcttag 1380
tgttctctac acacgtggga gatgaattac tcagcctttg tcagatgatt tctgttttgt 1440
gtatatttaa ctaatgaaca tatcacttgc accgtttcag gtgatgagga tagcagataa 1500
aggagctgat cttgttagaa taacagtcca gggtaggaag gaagctgatg cctgctttga 1560
gatcaagaac actctggttc agaagaagta aggtgccatt tttcttcaca ttcactgggg 1620
ttcacatgca tgggtcctca cccttttcat tgtttttgta tagttacaac attccactag 1680
tggccgatat tcattttgct cctacggtag ccctaagggt ggcagaatgt tttgacaaaa 1740
ttcgtgtgaa cccaggaaat tttggtgagt gaaatcttga catccctcat tttcgtctgt 1800
catgtttttg ttattgtaaa gtgctgagaa ctgtactggc agctgatcgt cgtgctcaat 1860
ttgaaaagct ggaatatact gacgacgact accaaaaaga gctagagcat attgagaagg 1920
tcgaaaaatt gtcactctag ccttaaaaaa aatcactctt agttttgtta caatcaatgt 1980
gaacatatgg catgttatct tacagagcca tgttaattat gtctcaggtg ttttctccat 2040
tagttgagaa atgcaagcag tatggaagag caatgcgaat aggaacaaat catggtagtc 2100
tttctgaccg cataatgagc tactatggtg attctccacg gggaatggtg attatctcat 2160
ttctgatgat ctacttaaat attagcacat aaaagtttga catcttctgc acaatgttgc 2220
aggttgagtc tgctttggaa tttgctagaa tatgtcggaa gttggacttc cacaactttg 2280
tattttcgat gaaagctagt aaccctgtta tcatggtcca agcatatcgc ctgcttgtgg 2340
cagaaatgta taacctagga tgggattatc ccttgcactt gggtgttaca gaagctggag 2400
agggtgaaga tggaaggatg aaatctgcta ttggcattgg gacactgcta atggtaattg 2460
atccactact tatttatgct gtatgaaatt gttaatctgg aaacgatgtt aacgacatgg 2520
aatagtagcc tcatggaatg gtaaatgctt cttttggagc ttatagctgg aagctttgag 2580
aatgaatatt ttttaaaaat attttttgct attcagttcc atagcatgat atgttgtgat 2640
gccaaattga caaaaaccct agaatcaaat agccgcctga aaaatgacgt gtttaagatt 2700
attaaacttt gttttgtgga ttaactgatt gggacatttg ccatgctacg acacatacta 2760
gagtacaaga ctatctgttt cctgcaaacc aattgaacta tggaaacagg caaattctac 2820
aaaaaacacg gactatatct tctgtgtctt ctattacagg atggtttggg tgatacaatc 2880
cgtgtctccc tcacagaacc accagaagaa gagattgatc cttgccaaag gttggcaaat 2940
cttgggacgc aggccgcaaa ccttcaaatt ggggtggtaa tggactttca ctttccatac 3000
tttgcagtca ccatatagta aatacatgtc cttactattt gtgttttccc ttaagcattt 3060
atttaattct cttggatttt gtattatttt atcaggcccc atttgaagaa aagcacaggc 3120
gttattttga tttccagcgt aggagtggtc aattgccttt gcagaaggag gtttgttaaa 3180
aataattcca cagtttgatc ataagaacta cactaatggt aggtttctta tcggttgctc 3240
atgctcttat gacagggtga ggaagttgac tacagaaatg tcctgcatcg tgatggttct 3300
gtactgatgt cagtttccct ggatcagttg aaggtaatct gcaccttttc tattcctttt 3360
tgctgagttc tggaatacat aatataacac caaaagggat ttcaccatgg ggttttgctt 3420
tgggatccat acctagtagt tcaaattaat cctctggcag gattcgaacc cgagtcctct 3480
gactctgaat ctgataccat attactaacc acgagattga caatcatcat ctgtccatag 3540
ggactgtagt atatatacag gccagcatgc cttagcaata ggatgagcct aatgggcctt 3600
aacagttagt agtaccattg ttgttttcca tgtatttttc tattgaaaat tggttgttac 3660
tccaaataaa ctttgcagtg accactctat tgattagcca tctgaataaa tgtgctctgt 3720
agtgattaat gctcccttta tttgccaggc tcctgatctc ctttataggt ctcttgcagc 3780
aaagcttgtg gttggcatgc ctttcaaggt ctgttctatg ttttttgtgt gctagtacac 3840
gactattttt agatggattt tagttgaagt tgattattac tcctatggct accaggatct 3900
ggctactgta gattccattc ttttgaggga gctcccccct gtagaagatg ctgaagctgt 3960
gagttgcagc tacatttcta tccctctctc aagtgcatat gttattcgtt ttattccttc 4020
tttagagctc tgctctaatt ttgtgaatct gtacagaggc ttgcactcaa aagattagtt 4080
gacatcagca tgggcgtact ggccccctta tcggagcagt tgacaaagcc actcccacat 4140
gcaattgtac ttgtcaacct cgacgaattg tcaagtggtg cacacaaact tttgccagaa 4200
ggtagttatt taaattaggc catgtcatta ttgtttcttt gctaatagtt taaggacaaa 4260
tatttcactg aagacaaata atagaaaatt gaacttttta gtttacaata gaaaaaggta 4320
ctcgctccga tctgatccaa tataaagatt gtttctagtt tttcatacgt caaacttttc 4380
taactttggc aaagttcatt gaaaaacgcc ccaaacatct acaacatcaa gttatttcat 4440
taaatccacc ctgaaatata tcttgataat tcagtttgat attgtagatg ttaatatatt 4500
tttcttataa actcggtcaa agttagagaa gttttgactt acgataaaac taaaatgact 4560
tacattttgg aatggagtga gtgttgtttt gcctaatctc atcattttgt ccttttttat 4620
ggtgtgtata ggcactagac tagctgtcac tcttcgtggt gatgaatcat acgagcagct 4680
agatattctt aaggatgttg atgatataac aatgttgtta cataatgttc catatggtga 4740
ggagaagaca ggcagggtgc atgctgctag gaggtaagaa aacacagtag tcaattacta 4800
ttaaatgaga gatttttagg attcatttat ccaccatgct caacagcact ttgagtcttt 4860
tttgtccttt taggtttaac atatttaata ttttttttat gtcttgcagg ttatttgagt 4920
acttacaggc caatggcttg aacttccctg taattcatca cataaatttc cctgaaacca 4980
ttgacaggtg actgtcaata ttcctatttt cttccccttg gaattgtgag tcgtggcttt 5040
cttgtttatt ctaattcaac tatgtgcaga gatggtcttg tcattggtgc tggggccaac 5100
gttggtgctc tcttagtcga tggtcttggt gatggtgtat tccttgaagc tgctgaccag 5160
gaatttgagt ttctgaggga cacatctttc aacttgctcc aaggttgcag gatgcgcaac 5220
acaaaaactg taggttcatg ttttttggtt tatttaatga tatggctacc tctttctgta 5280
tatccagtgg acgacatata ccactacttg caggagttgc agtaaaaaaa aagaaaaagg 5340
gaaaagaaac agcgaaacaa aaaaatatct gttgatgttt ctcactgaga tgagaattga 5400
ttttgtgaga agtgattttc tgactttgaa tgctgtagga atatgtgtct tgtccttcct 5460
gcggccgaac actctttgac cttcaggaaa tcagcgctga gattagagaa aagacctctc 5520
atctgccagg tgtctcggtg agttttctga ctcaaatata cgtggtctgg tcaactgtat 5580
aaggctcttt aaaaaaagat caactgtaag tctgtatata catttccttc ttttctatca 5640
attatactga tcaaccgtat acatatttgc ctggcagatc gctatcatgg gctgtattgt 5700
caatggacca ggagagatgg ctgatgccga ttttgggtac gttggaggcg ctcccggaaa 5760
gatcgacctt tacgtcggca aggtaatcat accgttgctt ggttgccaaa atcatgtact 5820
ccctgtgttc cgaattataa gacattttga cttttctaaa tatattgttt ttgttatata 5880
tatagacata tttatatcta gatgcatata aaagctatgt atgtagaaaa gtcaaaacat 5940
caaataattt ggaatggagg gagcatatgc aatggaaagt tttactatgt tggccagaac 6000
tgcctggttc agttagcact acacaaatgt ggttatgatc ccgtgataga acaattcttt 6060
ccgctaaaac agtgcatctt tctttctaca gaccgttgtg cagcgcgcca tcgccatgga 6120
aggtgccact gacgccttga tccagctgat caaggaccat ggccgatggg tggatcctcc 6180
tgccgaggag tagggctgta ggctgtagcc aagtctgtag tttatatatt ctgtaaagcg 6240
aaacacgcaa tgcagtaact gagcatacga tgtacatccc tacacatttt gtacatttat 6300
tgagatctga gaaataaact agaaaccgtc ggtgacggca tgtctgcagc accccaaatg 6360
aagcacatac gatttacata ttagtagtag cccatgtgag caagacataa tcctatacaa 6420
tcttgaagtc tgcattatat attttttcca acagaggtag cttagaaagt ttttcgagca 6480
atgcaatttt gttcgtta 6498
<210> 6
<211> 2698
<212> DNA
<213> 玉米
<400> 6
acgaactggc gtgccagtgc cagtgccgag cgcccgagcc cttactcctc ttcttccgac 60
cccccgctgg cttggaaaaa aggggaagaa aactcgcgca gtcggtcacc gaagggagga 120
gcgcggtcga ggtgccgccc acacggagct gcgaggtgtt tgtaggatct cctaggtgag 180
cccctgctgc ttggagacag ccatggccac cggcgtggct ccagctcctc tcccacatgt 240
cagagtgcat catgggggcg tcgggttcac caggagcgtc gattttgcga aggtcttgtc 300
tgctcccggt gccggcacga tgagggcaag ctcctctaga ggcagggcgc tcgtggcgaa 360
gagctctagt actggctcgg agaccatgga gctcgagcca tcttcagaag gaagcccact 420
tttagtaccc aggcagaagt actgtgaatc aacacaccag acaaggagga ggaaaactcg 480
aactgtgatg gtggggaatg tgccacttgg cagtgatcat cccataagga ttcaaaccat 540
gacgacttca gataccaagg atgttgcaaa aacagtagag gaggtgatga ggatagcaga 600
taaaggagct gatcttgtta gaataacagt ccagggtagg aaggaagctg atgcctgctt 660
tgagatcaag aacactctgg ttcagaagaa ttacaacatt ccactagtgg ccgatattca 720
ttttgctcct acggtagccc taagggtggc agaatgtttt gacaaaattc gtgtgaaccc 780
aggaaatttt gctgatcgtc gtgctcaatt tgaaaagctg gaatatactg acgacgacta 840
ccaaaaagag ctagagcata ttgagaaggt gttttctcca ttagttgaga aatgcaagca 900
gtatggaaga gcaatgcgaa taggaacaaa tcatggtagt ctttctgacc gcataatgag 960
ctactatggt gattctccac ggggaatggt tgagtctgct ttggaatttg ctagaatatg 1020
tcggaagttg gacttccaca actttgtatt ttcgatgaaa gctagtaacc ctgttatcat 1080
ggtccaagca tatcgcctgc ttgtggcaga aatgtataac ctaggatggg attatccctt 1140
gcacttgggt gttacagaag ctggagaggg tgaagatgga aggatgaaat ctgctattgg 1200
cattgggaca ctgctaatgg atggtttggg tgatacaatc cgtgtctccc tcacagaacc 1260
accagaagaa gagattgatc cttgccaaag gttggcaaat cttgggacgc aggccgcaaa 1320
ccttcaaatt ggggtggccc catttgaaga aaagcacagg cgttattttg atttccagcg 1380
taggagtggt caattgcctt tgcagaagga gggtgaggaa gttgactaca gaaatgtcct 1440
gcatcgtgat ggttctgtac tgatgtcagt ttccctggat cagttgaagg ctcctgatct 1500
cctttatagg tctcttgcag caaagcttgt ggttggcatg cctttcaagg atctggctac 1560
tgtagattcc attcttttga gggagctccc ccctgtagaa gatgctgaag ctaggcttgc 1620
actcaaaaga ttagttgaca tcagcatggg cgtactggcc cccttatcgg agcagttgac 1680
aaagccactc ccacatgcaa ttgtacttgt caacctcgac gaattgtcaa gtggtgcaca 1740
caaacttttg ccagaaggca ctagactagc tgtcactctt cgtggtgatg aatcatacga 1800
gcagctagat attcttaagg atgttgatga tataacaatg ttgttacata atgttccata 1860
tggtgaggag aagacaggca gggtgcatgc tgctaggagg ttatttgagt acttacaggc 1920
caatggcttg aacttccctg taattcatca cataaatttc cctgaaacca ttgacagaga 1980
tggtcttgtc attggtgctg gggccaacgt tggtgctctc ttagtcgatg gtcttggtga 2040
tggtgtattc cttgaagctg ctgaccagga atttgagttt ctgagggaca catctttcaa 2100
cttgctccaa ggttgcagga tgcgcaacac aaaaactgaa tatgtgtctt gtccttcctg 2160
cggccgaaca ctctttgacc ttcaggaaat cagcgctgag attagagaaa agacctctca 2220
tctgccaggt gtctcgatcg ctatcatggg ctgtattgtc aatggaccag gagagatggc 2280
tgatgccgat tttgggtacg ttggaggcgc tcccggaaag atcgaccttt acgtcggcaa 2340
gaccgttgtg cagcgcgcca tcgccatgga aggtgccact gacgccttga tccagctgat 2400
caaggaccat ggccgatggg tggatcctcc tgccgaggag tagggctgta ggctgtagcc 2460
aagtctgtag tttatatatt ctgtaaagcg aaacacgcaa tgcagtaact gagcatacga 2520
tgtacatccc tacacatttt gtacatttat tgagatctga gaaataaact agaaaccgtc 2580
ggtgacggca tgtctgcagc accccaaatg aagcacatac gatttacata ttagtagtag 2640
cccatgtgag caagacataa tcctatacaa tcttgaagtc tgcattatat attttttc 2698
Claims (10)
1.蛋白质的下述X1)或X2)的应用:
X1)在调控玉米白化中的应用;
X2)在玉米育种中的应用;
所述蛋白质为如下A1)或A2)或A3):
A1)氨基酸序列是序列表中序列1的蛋白质;
A2)氨基酸序列是序列表中序列4的蛋白质;
A3)将序列表中序列1或序列4所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有相同功能的蛋白质。
2.与权利要求1所述蛋白质相关的生物材料的下述X1)或X2)的应用:
X1)在调控玉米白化中的应用;
X2)在玉米育种中的应用;
所述生物材料为下述B1)至B7)中的任一种:
B1)编码权利要求1中所述蛋白质的核酸分子;
B2)含有B1)所述核酸分子的表达盒;
B3)含有B1)所述核酸分子的重组载体、或含有B2)所述表达盒的重组载体;
B4)含有B1)所述核酸分子的重组微生物、或含有B2)所述表达盒的重组微生物、或含有B3)所述重组载体的重组微生物;
B5)含有B1)所述核酸分子的转基因植物细胞系、或含有B2)所述表达盒的转基因植物细胞系;
B6)含有B1)所述核酸分子的转基因植物组织、或含有B2)所述表达盒的转基因植物组织;
B7)含有B1)所述核酸分子的转基因植物器官、或含有B2)所述表达盒的转基因植物器官。
3.根据权利要求2所述的应用,其特征在于:B1)所述核酸分子为如下b1)-b6)中的任一种:
b1)序列表中序列5或序列5的第317-6498位或序列5的第843-6193位所示的DNA分子;
b2)编码序列是序列表中序列6或序列6的第203-2443位的cDNA分子或DNA分子;
b3)序列表中序列2或序列2的第2196-8353位或序列2的第2722-8045位所示的DNA分子;
b4)编码序列是序列表中序列3或序列3的第204-2444位的cDNA分子或DNA分子;
b5)与b1)或b2)或b3)或b4)限定的核苷酸序列具有75%或75%以上同一性,且编码权利要求1中所述蛋白质的cDNA分子或基因组DNA分子;
b6)在严格条件下与b1)或b2)或b3)或b4)限定的核苷酸序列杂交,且编码权利要求1中所述蛋白质的cDNA分子或基因组DNA分子。
4.权利要求1中所述蛋白质或权利要求2或3中所述生物材料。
5.构建玉米白化模型的方法,包括降低玉米中权利要求1中所述蛋白质的含量和/或活性得到所述玉米模型。
6.根据权利要求5所述的方法,其特征在于:所述方法包括抑制编码所述蛋白质的基因的表达。
7.根据权利要求6所述的方法,其特征在于:所述方法通过对编码权利要求1中所述蛋白质的核酸分子进行突变实现。
8.根据权利要求7所述的方法,其特征在于:所述突变采用CRISPR/Cas9方法实现。
9.根据权利要求8所述的方法,其特征在于:所述CRISPR/Cas9方法中所用靶序列为靶序列1和/或靶序列2;
所述靶序列1为如下m1)、m2)或m3):
m1)序列表中序列6的第288-306位的核苷酸序列;
m2)与m1)限定的DNA序列具有75%或75%以上同一性的由m1)衍生的DNA序列;
m3)在严格条件下与m1)限定的DNA序列杂交的由m1)衍生的DNA序列;
所述靶序列2为如下n1)、n2)或n3):
n1)序列表中序列6的第354-372位的核苷酸序列;
n2)与n1)限定的DNA序列具有75%或75%以上同一性的由n1)衍生的DNA序列;
n3)在严格条件下与n1)限定的DNA序列杂交的由n1)衍生的DNA序列。
10.根据权利要求1-3中任一所述的应用或权利要求5-9中任一所述的方法,其特征在于:所述白化为叶片白化和/或全植株白化和/或玉米籽粒白化;
进一步,所述玉米籽粒白化表现为籽粒由黄色变为浅黄色;
更进一步,所述玉米籽粒白化为部分籽粒颜色变化。
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CN113801870A (zh) * | 2021-10-09 | 2021-12-17 | 中国农业科学院作物科学研究所 | SiLCYB调控番茄红素等谷子类胡萝卜素合成代谢的功能及应用 |
CN115216455A (zh) * | 2021-04-19 | 2022-10-21 | 中国农业大学 | Enb1基因及其编码蛋白在调控植物籽粒大小和粒重中的应用 |
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