CN111235180A - 缩短玉米花期的方法 - Google Patents
缩短玉米花期的方法 Download PDFInfo
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Abstract
本发明属于分子遗传学领域。具体涉及控制玉米花期的基因及其在改变玉米花期性状方面的应用。本发明提供了一个控制玉米花期基因的序列，且公开了利用基因工程手段突变该基因以缩短花期的方法。本发明还提供了花期缩短的玉米突变基因型序列，该突变基因型序列可以用来改良玉米花期性状。
Description
技术领域
本发明属于分子遗传学领域。具体涉及控制玉米花期的基因及其在改变玉米花期性状方面的应用。本发明提供了一个控制玉米花期基因的序列，且公开了利用基因工程手段突变该基因以缩短花期的方法。本发明还提供了花期缩短的玉米突变基因型序列，该突变基因型序列可以用来改良玉米花期性状。
背景技术
在高等植物中，开花代表着从营养生长到生殖生长的转换过程，其在植物体的整个生长发育阶段中具有重要的作用。而何时开花这一生物性状则受到植物体自身遗传因子及外界环境因素的双重作用。在这双重作用的影响下，一系列开花诱导过程在高等植物中具有普遍的规律，即植物体叶片通过感受外界生长条件(光照，温度，湿度等)，在适宜的时间产生成花物质(或叫开花素florigen)，成花物质经过输导组织由叶片运送到茎尖生长点，刺激顶端分生组织成花。
开花期是作物进化和适应过程中的重要性状，理解作物开花期性状的遗传基础、克隆候选基因可以提高作物的环境适应能力和可塑性，这对培育适应不同生态区的优良作物品种具有重要的意义，同时也将促进产量等与开花期密切相关的重要生产性状的遗传改良进程。此外，作物的整体生育期主要决定于开花期的长短，适度缩短开花期能够减少整体的生育期，从而降低种植成本。
发明内容
本发明的目的之一在于提供一个控制玉米花期性状的基因序列。
本发明的目的之二在于公开一种缩短玉米花期的方法。
为实现上述目的，本发明采用如下技术方案：
本发明提供了一种控制玉米花期的基因及其在改变玉米花期性状中的应用，其特征在于：上述基因的核酸序列如SEQ ID NO.1-SEQ ID NO.3任意一个所示。其中，SEQ IDNO.1序列为玉米自交系B73中的基因组序列，SEQ ID NO.2序列为玉米自交系KN5585中的基因组序列，SEQ ID NO.3序列为玉米自交系B73中的cDNA序列。上述基因的核酸序列序列经过取代和/或缺失和/或添加一个或多个氨基酸，且具有相同功能的序列也属于本发明的内容。
在另一方面，本发明还提供了一种缩短玉米花期的方法，其特征在于：抑制玉米中上述基因编码蛋白的表达和/或活性，选择玉米花期缩短的植株。
在一些实施方案中，上述蛋白的氨基酸序列如SEQ ID NO.4所示。
在一些实施方案中，上述抑制蛋白表达和/或活性的方法包括基因编辑、RNA干扰、T-DNA插入、物理或化学诱变中任一种。
在一些实施方案中，上述基因编辑采用CRISPR/Cas9方法。
在一些实施方案中，上述CRISPR/Cas9方法在玉米中的基因组靶标区域的DNA序列如SEQ ID NO.5或SEQ ID NO.6所示。
在另一方面，本发明还提供了一种用于缩短玉米花期的试剂盒，其特征在于：包括如下任一种：
(1)sgRNA分子，其序列如SEQ ID NO.7或SEQ ID NO.8所示；
(2)所述sgRNA的编码DNA分子；
(3)表达所述sgRNA的载体。
在另一方面，本发明还提供了用上述试剂盒得到的一种玉米突变基因型，其特征在于：所述突变基因型序列如SEQ ID NO.9所示。通过基因编辑突变靶基因可以得到很多不同的编辑类型，这些不同的编辑类型所对应的植株表现并不是完全一样的。本发明经过筛选测试，确定SEQ ID NO.9所示的突变基因型能够使玉米花期缩短。该基因型序列可以通过有性杂交的方式导入不同遗传背景的玉米材料中，从而创制花期缩短的玉米新品种。
在另一方面，本发明还提供了用于检测上述突变基因型的引物对，其特征在于：所述引物对为SEQ ID NO:10和SEQ ID NO:11所示的序列或其互补序列。
在另一方面，本发明还提供了检测上述基因型的方法，其特征在于：利用上述引物对检测待测样品中是否存在所述基因型。利用上述引物对对待测样品的基因组DNA进行PCR扩增，并测序分析扩增产物序列，如果扩增产物测序后序列如SEQ ID NO:9所示，则待测样品中包含所述基因型。
本发明的优点及有益效果如下：本发明利用多亲本高世代自交系群体定位到控制玉米株高性状的基因组区域，该区域并不存在已知的影响株高的基因，只存在一个未知功能的注释基因GRMZM2G068943。利用CRISPR/Cas9方法突变GRMZM2G068943基因，意外地发现该基因能够控制玉米花期性状。利用CRISPR/Cas9基因编辑的方法和编辑后的突变基因型序列，可以缩短玉米花期，提高玉米的生态适应性。
附图说明
图1株高QTL定位结果。纵轴表示每个标记关联分析检验的p-value，取-log10，横轴表示染色体的位置。
图2基因编辑载体图。各元件英文及缩写含义列举如下：
RB T-DNA repeat T-DNA右边界重复序列
M13 fwd M13引物序列(正向)
p000204_1F 靶标gRNA序列
Ubi promoter 泛素启动子
3×FLAG 标签序列
SV40NLS 猿猴病毒40核定位信号
Cas9 cas9基因序列
Nucleoplasm in NLS 核定位信号
NOS terminator 胭脂碱合成酶终止子
lac promoter 乳糖启动子
M13 rev M13引物序列(反向)
lac operator 乳糖操纵子
CAP biding site CAP结合位点
CaMV35S promoter(enhanced) 增强的花椰菜花叶病毒35S启动子
BlpR 编码Bar蛋白赋予植物具有草铵膦耐性
CaMV35S polyA single 花椰菜花叶病毒35S多聚腺苷酸序列
LB T-DNA repeat T-DNA左边界重复序列
Kan R 卡那霉素抗性序列
Ori 起始区序列
Bom 骨架区序列
pVS1 RepA pVS1复制子
pVS1 StaA pVS1转录起始区
具体实施方式
提供以下定义和方法用以更好地界定本申请以及在本申请实践中指导本领域普通技术人员。除非另作说明，术语按照相关领域普通技术人员的常规用法理解。本文所引用的所有专利文献、学术论文、行业标准及其他公开出版物等，其中的全部内容整体并入本文作为参考。
如本文所用，“玉米”是任何玉米植物并包括可以与玉米育种的所有植物品种，包括整株植物、植物细胞、植物器官、植物原生质体、植物可以从中再生的植物细胞组织培养物、植物愈伤组织、植物或植物部分中完整的植物细胞，所述植物部分例如胚、花粉、胚珠、种子、叶、花、枝、果实、茎杆、根、根尖、花药等。除非另有所指，核酸以5’至3’方向从左向右书写；氨基酸序列以氨基至羧基方向从左向右书写。氨基酸在本文可以用其通常所知的三字母符号或IUPAC-IUB生物化学命名委员会推荐的单字母符号来表示。同样地，可以用通常接受的单字母码表示核苷酸。数字范围包括限定该范围的数字。如本文所用，“核酸”包括涉及单链或双链形式的脱氧核糖核苷酸或核糖核苷酸多聚物，并且除非另有限制，包括具有天然核苷酸基本性质的已知类似物(例如，肽核酸)，所述类似物以与天然存在的核苷酸类似的方式与单链核酸杂交。如本文所用，术语“编码”或“所编码的”用于特定核酸的上下文时，指该核酸包含指导该核苷酸序列翻译成特定蛋白的必需信息。使用密码子表示编码蛋白的信息。如本文所用，涉及特定多核苷酸或其所编码的蛋白的“全长序列”指具有天然(非合成)内源序列的整个核酸序列或整个氨基酸序列。全长多核苷酸编码该特定蛋白的全长、催化活性形式。本文可互换地使用术语“多肽”和“蛋白”，以指氨基酸残基的多聚物。该术语用于氨基酸多聚物，其中一个或多个氨基酸残基是相应天然存在的氨基酸的人工化学类似物。该术语还用于天然存在的氨基酸多聚物。本文可互换地使用术语“残基”或“氨基酸残基”或“氨基酸”，以指被并入蛋白、多肽或肽(统称“蛋白”)的氨基酸。氨基酸可以是天然存在的氨基酸，并且除非另有限制，可以包括天然氨基酸的已知类似物，所述类似物可以与天然存在的氨基酸相似的方式起作用。
如本文所用，可以互换地使用术语“分离的”和“纯化的”，以涉及核酸或多肽或其生物学活性部分，其基本上或本质上不含如在其天然存在的环境中所发现的通常伴随或反应于该核酸或多肽的组分。因而，用重组技术产生分离的或纯化的核酸或多肽时，分离的或纯化的核酸或多肽基本上不含其它细胞物质或培养基，或者化学合成分离的或纯化的核酸或多肽时，基本上不含化学前体或其它化学品。“分离的”核酸通常不含在该核酸所衍生自的生物体的基因组DNA中天然侧翼于该核酸(即位于该核酸5’和3’端的序列)的序列(诸如，编码蛋白的序列)。例如，在各种实施方案中，所分离的核酸可以包含在该核酸所衍生自的细胞的基因组DNA中天然侧翼于该核酸的少于约0.5kb的核苷酸序列。
在本申请中，将词语“包括”、“包含”或其变体应理解为除所描述的元素、数或步骤外，还包含其它元素、数或步骤。“受试植物”或“受试植物细胞”是指遗传改造已经生效的植物或植物细胞，或者如此改造的植物或细胞的子代细胞，该子代细胞包含所述改造。“对照”或“对照植物”或“对照植物细胞”提供用于测量受试植物或植物细胞表型改变的参考点。对照植物或植物细胞可以包括，例如：(a)野生型植物或细胞，即与遗传改造起始材料具有相同基因型的植物或细胞，所述遗传改造产生受试植物或细胞；(b)与所述起始材料具有相同基因型但已用空构建体(即用对目的性状无已知效果的构建体，诸如包含标物基因的构建体)转化的植物或植物细胞；(c)是受试植物或植物细胞的非转化分离子的植物或植物细胞；(d)与所述受试植物或植物细胞在遗传上一致但未暴露于会诱导目的基因表达的条件或刺激物的植物或植物细胞；或(e)受试植物或植物细胞自身，其处于目的基因不被表达的条件下。
本领域技术人员会容易地认同，诸如位点特异性诱变和随机诱变、聚合酶链式反应方法和蛋白工程化技术的分子生物学领域的进步提供了广泛的适当的工具和操作步骤，以用于改造或者工程化农业上感兴趣的蛋白的氨基酸序列和潜在的基因序列。
在一些实施方案中，可以对本申请的核苷酸序列进行改变，以进行保守氨基酸替换。保守氨基酸替换的原则和实例在下文中进一步描述。在某些实施方案中，可以依照公开的单子叶密码子偏好性对本申请的核苷酸序列进行不改变氨基酸序列的替换，例如可以用单子叶植物偏好的密码子替换编码同一氨基酸序列的密码子，而不改变该核苷酸序列所编码的氨基酸序列。在一些实施方案中，以编码同一氨基酸序列的不同密码子替换本申请中的部分核苷酸序列，从而在改变核苷酸序列的同时不改变其编码的氨基酸序列。保守变体包括由于遗传密码子简并性而编码实施方案的蛋白中的一种的氨基酸序列的那些序列。在一些实施方案中，根据单子叶植物偏好密码子替换本申请中的部分核苷酸序列。本领域技术人员会认识到氨基酸添加和/或取代通常基于氨基酸侧链取代基的相对相似性，例如，所述取代基的疏水性、电荷、大小等等。具有各种前述所考虑性质的示例性氨基酸取代基团为本领域技术人员所公知，并且包括精氨酸与赖氨酸；谷氨酸和天门冬氨酸；丝氨酸和苏氨酸；谷氨酰胺和天冬酰胺；以及缬氨酸、亮氨酸和异亮氨酸。关于不影响目的蛋白生物学活性的适当氨基酸取代的指南可以在Dayhoff等人(1978)Atlas of Protein Sequence andStructure(蛋白序列和结构图集)(Natl.Biomed.Res.Found.,Washington,D.C)(通过引用并入本文)的模型中找到。可以进行诸如将一个氨基酸换作具有相似性质的另一个氨基酸的保守性取代。序列一致性的鉴定包括杂交技术。例如，将已知核苷酸序列的全部或部分用作与其它相应核苷酸序列选择性杂交的探针，所述其它相应核苷酸序列存在于来自所选生物体的已克隆基因组DNA片段或cDNA片段群(即基因组文库或cDNA文库)。
在一些实施方案中，还包括核苷酸序列及其编码的氨基酸序列的片段。如本文所用，术语“片段”指实施方案的多核苷酸的核苷酸序列的一部分或者多肽的氨基酸序列的一部分。核苷酸序列的片段可以编码蛋白片段，所述蛋白片段保留天然或相应全长蛋白的生物学活性，并因而具有蛋白活性。突变体蛋白包括天然蛋白的生物活性片段，其包含保留天然蛋白生物学活性的连续氨基酸残基。一些实施方案还包括转化的植物细胞或转基因植物，其包含至少一种实施方案的核苷酸序列。在一些实施方案中，使用表达载体转化植物，所述表达载体包含至少一种实施方案的核苷酸序列以及与其可操作地连接的在植物细胞中驱动表达的启动子。转化的植物细胞和转基因植物表示基因组内包含异源多核苷酸的植物细胞或植物。一般来说，所述异源多核苷酸在转化的植物细胞或转基因植物的基因组内稳定地整合，以致将所述多核苷酸传递给后代。可以将所述异源多核苷酸单独地或作为表达载体的一部分整合进基因组。在一些实施方案中，本申请涉及的植物包括植物细胞、植物原生质体、可以再生出植物的植物细胞组织培养物、植物愈伤组织、植物团块和植物细胞，其为完整的植物或者植物的部分，诸如胚胎，花粉，胚珠，种子，叶，花，枝，果实，果仁，穗，穗轴，壳，秸秆，根，根尖，花药等等。本申请还包括源于本申请的转基因植物或其子代、并因而至少部分地包含本申请的核苷酸序列的植物细胞、原生质体、组织、愈伤组织、胚胎以及花、茎、果实、叶以及根。
以下实施例用于说明本发明，但不用来限制本发明的范围。在不背离本发明精神和实质的情况下，对本发明方法、步骤或条件所作的修改或替换，均属于本申请的范围。若无特别指明，实施例按照常规实验条件，如Sambrook等人的分子克隆实验手册(SambrookJ&Russell D W,Molecular cloning:a laboratory manual,2001)，或按照制造厂商说明书建议的条件。若未特别指明，实施例中所用的化学试剂均为常规市售试剂，实施例中所用的技术手段为本领域技术人员所熟知的常规手段。
实施例
实施例1玉米花期QTL的定位过程
1、群体构建
本发明将24个中国的优良玉米自交系，通过两代双列杂交、六代开放授粉及六代自交的方式获得了1404个子代组成的多亲本高世代自交系群体(CUBIC)。这24个中国优良玉米自交系包括LV28(旅28)、E28、DAN340(丹340)、F349、ZI330(自330)、ZONG3(综3)、ZONG31(综31)、HUANGC(黄C)、HZS(黄早四)、HYS(黄野四)、TY4(天涯4)、YUANGFH(原辅黄)、CHANG7-2(昌7-2)、K12、XI502(西502)、LX9801、H21、SHUANG741(双741)、Q1261、JI853(吉853)、JI53(冀53)、5237、81515、NX110(农系110)，该群体的24个亲本及1404子代自交系全部进行基因组测序，获得了超过14M的SNP和InDel变异。
2、表型分析
所有24个亲本和1404子代自交系在黄淮海及东北5个地点种植，表型变异丰富。所考查农艺表型包括植株高度、花期、开花期、穗重等产量性状等。将所有环境重复计算出每个自交系的最佳线性无偏预测因子(BLUP)值用于后续分析，包括基本表型统计、相关性分析和GWAS。通过线性回归估算由给定性状的QTL解释的表型方差或遗传力，并联合估算解释的总方差。
3、基于单变量的GWAS分析(sGWAS)
根据最小等位基因频率(MAF)过滤后，共计1180万个高质量SNP用于下游分析。前十个主成分(PC)共解释了8.76％的遗传方差，同时整合了亲缘关系K矩阵作为随机效应一起用于sGWAS分析。关联的显着性阈值设置为1.23E-8，等于0.05/Ne，其中Ne是根据所有变异计算的独立检验的有效数量。确定QTL区间的标准为：每个性状的显著SNP被首先提取，只保留至少两个连续SNP小于20Kb的位点同时合并作为基本区间单元，20KB以内只有单独显著位点的SNP在下一步分析中被排除；如果相邻区间单元之间的任何显著SNP有较高连锁不平衡关系(LD，r2≥0.2)，则将相邻区间单元进一步合并为QTL候选区间。为了减少由较小的QTL效应而获得的QTL间隔较小而未包含功能基因的可能性，对于那些间隔小于50Kb且最显著SNP的显著性低于阈值100倍的QTL(即1.23E-10)，将对QTL区间向两侧继续延伸25Kb获得最终QTL区间。将区间内最显著的SNP作为sQTL的显著性，将区间间隔内所有基因作为候选基因。
4、获得QTL区间
利用上述的定位方法，本发明找到了玉米株高性状的1个最显著SNP，位于第8染色体126969684-126977958区间内(图1)。该区间内经MaizeGDB数据库(https://www.maizegdb.org/)的查询包含1个有注释的基因，编号为GRMZM5G878561，数据库中并未明确该基因的功能。该基因的基因组序列如SEQ ID NO.1所示，全长9800bp，cDNA序列如SEQID NO.3所示，编码的氨基酸序列如SEQ ID NO.4所示。本发明同时测定了未米生物科技(江苏)有限公司选育的玉米自交系KN5585中该区间的序列，如SEQ ID NO.2所示。
实施例2基因编辑敲除候选基因分析基因功能
本发明利用CRISPR-Cas9基因编辑技术对上述区间内的基因进行定点突变。实施方式包括基因编辑载体的构建、玉米的遗传转化以及编辑效果的功能验证。
具体如下：
1、基因编辑载体的构建
本发明的基因编辑载体为G08943-CPB-ZmUbi-hspCas9，其载体图如图2所示。该载体的基础载体为CPB-ZmUbi-hspCas9。本发明通过OverlapPCR获得双靶U6-sgRNA进而通过同源重组克隆到基础载体中，具体的构建流程如下：
(1)U6启动子的克隆。从B73中克隆U6启动子。
(2)靶标gRNA的设计。将受体材料KN5585基因组序列输入http://cbi.hzau.edu.cn/crispr/进行靶标设计。本发明优选的两个靶标区域的DNA序列如SEQ IDNO.5或SEQ ID NO.6所示。本发明的sgRNA骨架序列通过人工合成得到，其序列信息如SEQID NO.7和SEQ ID NO.8所示。
(3)通过Overlap PCR获得U6-sgRNA。引物对pU6F1/pU6R用于扩增第一个靶标的U6启动子，产物长度为515bp；引物对p1652-1F(3F)/pgRR1用于扩增第一个靶标的sgRNA，产物长度为127bp；引物对pU6F1/gRR1用于进行Overlap PCR第2步扩增(U6-sgRNA)，产物长度为634bp。Overlap PCR第1步分别扩增U6和sgRNA，PCR产物分别稀释50倍后混合作为模板进行Overlap PCR第2步扩增。扩增产物电泳切胶回收并测序确认序列。Overlap PCR体系和条件如下：Overlap PCR第1步的15μL的反应体系如下，模板DNA(U6或sgRNA，≥30ng/μL)：0.5μL，Primer F/R：各1.2μL，灭菌ddH2O：3.7μL，2×phanta max Buffer：7.5μL，dNTP mix：0.6μL，Phanta酶(产品编号：P505-d1/d2/d3)：0.3μL。Overlap PCR第2步的反应体系为30μL体系。U6吸1μL，加49μL ddH2O稀释；sgRNA吸1μL，加49μL ddH2O稀释各吸10μL，混匀。具体如下：混合模板DNA(U6+sgRNA)：1.5μL，Primer F/R：各2.4μL，灭菌ddH2O：6.9μL，2×phanta maxBuffer：15μL，dNTP mix：1.2μL，Phanta酶：0.6μL。Overlap PCR程序如下：(1)94℃5分钟，(2)94℃30秒，(3)62℃35秒，(4)72℃30秒，第(5)步是从(2)步－(4)步循环32次，(6)72℃10分钟，(7)25℃5分钟。载体构建所需的引物序列见表1。
表1载体构建所需的引物序列
(4)通过重组克隆构建到骨架载体。将CPB-Ubi-hspcas9载体用HindIII消化，回收。通过同源重组将U6-gRNA和载体二者连接。配反应液前保证各Overlap产物浓度接近一致，20μL的同源重组体系如下：Cas Hind III：3μL，T-1F Overlap：1μL，灭菌ddH2O：10μL，5×CE MultiS buffer：4μL，Exnase MultiS(产品编号：C113-01/02)：2μL。
2、玉米遗传转化
将载体通过电击法转入农杆菌EHA105中，PCR进行鉴定。以新鲜剥离的1mm左右的玉米自交系KN5585(未米生物科技(江苏)有限公司选育的自交系)的幼胚为材料，将剥取的玉米胚放入含有1.8mL悬浮液的2mL塑料离心管中，30min内大约处理未成熟幼胚150个；吸去悬浮液，余下玉米胚在管中然后加入1.0mL农杆菌悬浮液，放置5min。将离心管中的幼胚悬浮后倒入共培养基上，并用移液器吸去表面多余的农杆菌菌液，于23℃黑暗共培养3天。共培养后，将幼胚转移到休息培养基中，于28℃黑暗培养6天后，放至含5mg/L Bialaphos的筛选培养基上，开始筛选培养2周，然后转到含8mg/L Bialaphos筛选培养基上筛选培养2周。将抗性愈伤组织转移至分化培养基1中，25℃，5000lx，光照培养1周。再将愈伤转移至分化培养基2中，光照培养2周；将分化生出的小苗转移至生根培养基上，25℃，5000lx，光照培养直到生根；将小苗转入小盆中生长，一定生长阶段后移栽于温室中，3-4个月后收获后代种子。
3、基因编辑植株的性状鉴定
共获得两个T0代转化体，他们与对照相比株高性状没有差异。然而其中一个转化体的花期性状与对照相比有变化。对这个花期有变化转化体的T1代材料提取苗期DNA检测基因编辑情况，设计引物，引物序列如SEQ ID NO.10和SEQ ID NO.11所示。扩增靶标编辑区段，扩增体系为：DNA：3μL，双向引物各1μL，2×TaqMix：7.5μL，ddH2O：2.5μL，总体积10μL。PCR反应条件如下：(1)94℃5分钟，(2)94℃40秒，(3)57℃30秒，(4)72℃60秒，(5)从(2)步－(4)步循环35次，(6)72℃7分钟，(7)4℃保存。PCR产物交由武汉擎科生物科技有限公司进行Sanger测序。将转化株测序结果与野生型KN5585的基因组比对，发生碱基替换、插入或缺失的材料即为阳性编辑材料，否则为阴性材料。
PCR扩增靶标编辑区段并对PCR产物进行测序。将PCR产物序列与原始的KN5585中的序列(SEQ ID NO.2)进行比对，发现有2种不同基因型的材料，这两个材料的靶点都发生了突变。其中A1材料的花期明显提早，证明该基因控制花期性状，基因突变后花期提前。而A2材料表型与对照相比无差异(表2)。
A1材料的PCR扩增产物(序列如SEQ ID NO.9所示)在基因靶点处发生了编辑，两个靶点之间的大段序列发生了反转(表2)。
表2基因编辑玉米材料的花期性状数据
数据以“平均值±标准差”表示，字母表示两个材料间的差异显著性(α＝0.05)。“＝”表示序列反转，黑体表示碱基插入或变异，方框表示PAM序列，下划线表示编辑靶标。
虽然，上文中已经用一般性说明及具体实施方案对本发明作了详尽的描述，但在本发明基础上，可以对之作一些修改或改进，这对本领域技术人员而言是显而易见的。因此，在不偏离本发明精神的基础上所做的这些修改或改进，均属于本发明要求保护的范围。
序列表
<110> 未米生物科技（江苏）有限公司，华中农业大学
<120> 缩短玉米花期的方法
<130> 1
<160> 11
<170> SIPOSequenceListing 1.0
<210> 1
<211> 9800
<212> DNA
<213> Zea mays
<400> 1
atctacagta attaagataa gaagttcatt ggaacccaga aatttgagtg gggcatgtgc 60
ccccagtggc cataacgtcg gtccgccctt gagccccgcc cgccccaccg ctggcttcat 120
cactgtctac tgtcgttcca ccataatcta acattctaac ttacggaaac aaacacaccc 180
taaaagctaa gctgaaccca acgctatgat caacgaaagt tagttaactt ttttgagctt 240
agcaattatt ttagttaaca aaaactttga ttttttccta tcgtggctcg tgagtgccat 300
tcaacctaaa atcgagtgga cacgtataat tagaggcatg aagcggtaga gcacacacat 360
aggcacaggt gaacggggat tggtagcatt ccatccacga acataagcca cgtacgctag 420
ctcgtcgtag cggcggtgtg tagctctccg tatcctaacc cacagtcaca agatgctcac 480
ctgcagcctt actcctgcgt tgccaaaaaa tttccccttt taaaaattta aatccgtaca 540
cgtgggttct acaagacagt agcctataaa tgaaggtgta gaaaatctac gctgcaaaac 600
atctaacttt tcgtataagc agcttcttca ataatccatc aacgaccgtt aggagcggtg 660
acaatctagc tgtctagaaa acgcgaaact tgcaagaccc ctccgtcgct cgctcccata 720
aaagcctcaa tccctccacc cacccacctc gcctgcatat cactcgtgtg ctcgcaaaca 780
agcgtccgag ttggaggcgt ggcggcccgt tttgcatgcc gtagctctcc cggtgttcct 840
ccccttccga acccctcgcc gcctctccgt ctcgtgattc gaatcactgc cccattgcga 900
acgagccgcc acagctgctg ccagcctcaa ggaagggccc cggttcccaa tcccagacgg 960
gccagcggat cgatcgaaca ggcggccggc cactcgggat cgggtgggtg agtcgttccg 1020
cttccgaaac cagcctccct gcgtgatggg cctgcttggg tccgcctgtt gccgcctcct 1080
cgtcgtcgct caccgctcgc gtccgcctaa tgctgcaggc cggggccgcg gcggcgcatg 1140
agctctgacg ccgcgagggg gcagcgcggc atcaactgca cgaggggcga cgcggcggcg 1200
atgccaacct catcgccctt tgtcggcgac agcggcggcg cgggctcccc gatccgcgtc 1260
gagcgaatgg tccgcgagcg tagccgccgc tacgacatct tcgcgtcgga cgcgatggat 1320
accgacgccg aggcggcctt cgcgctggat ggggtccagt cgcctggacg tgcgtcaccc 1380
gccaacatgg aggatgctgg cggcgcggcc gcagcgcgac cgccgctcgc cggctcccgc 1440
agcggtttcc gccgcctagg cctccgtggc atgaagcagc gcctcctcgt cgtggccaac 1500
cgcctgcctg tatccgccaa ccgccgcgga gaggaccact ggtcgcttga gatcagcgcc 1560
ggcggccttg tgagcgccct ccttggtaat cccgacattt aatctcctcc caaatgcgac 1620
gtggccggtg gcctggacct ggagatgcat gcaatggaat tctgcggtaa catctattac 1680
cttagttcat atgaatgttt tttttctgat gctgaaggcg tgaaggacgt cgacgcgaag 1740
tggattggct gggcaggcgt caacgtaccg gacgaggttg gtcagcgagc cctcactaaa 1800
gctcttgcgg agaaggttgg ttgcaattcc atgttgtctg ggtcaataga acagtgactc 1860
tttgttagta tcactctgat aactcgaggc atttcttgtg tggattgctt gaactttgca 1920
gagatgcata ccagtgttcc tggacgagga gattgtgcac cagtactaca atggatactg 1980
caacaacatc ctgtggccac tgttccacta cctaggacta ccacaggagg acaggctggc 2040
aacaacgagg aactttgagt cacagttcga cgcgtacaag cgtgctaacc agatgttcgc 2100
tgatgtcgtg tacgagcact accaggacgg ggatgtgatc tggtgccatg actaccacct 2160
catgtttctg cccaagtgcc tcaaggacca tgacatcaat atgaaggtcg ggtggttcct 2220
gcacacgccg ttcccgtcat cagagattta ccggacactg ccgtcccgct tggagctgct 2280
tcggtcggtg ctgtgtgccg atttagttgg gtaagcatac ctatggattt tgtgtcggct 2340
gtcctgtcga atttgattgt taaacctgat aagtttaact gagtatcctt tttattcttg 2400
atttacttga agtaacttta tggtttcgat tatgtttttt gtgcatgtaa gtcccaactc 2460
acaacgtccg aaaattgtat atttcccttc cactataaag ttactctctg atcagtgcat 2520
atttaatagt tttttccttc tttttttttt tggggggggg gggggggtca ggccaaaatg 2580
ttctagtttg acagcatgga aaatattgcc tgccagttca ttcactgtat aacatagaaa 2640
gtgttactat agattgtgat gcttttgtta tgtttctgtg acacgtattg ctatcatggt 2700
aggaatgtat tacatacacg attttggtga tgtatttttt tttagtttga actgacttgg 2760
cctatgtatg aagtttgctt tagattcaaa agactaaacc tatttagtaa atttttagtt 2820
ttcttcacat tttcaagaaa tttattctag aagagccaaa ataataaaaa aaaacaggcc 2880
aacacgtcgt gtgaaattat cagttccatt tctatgagtt atttcatggt gctcttatta 2940
attttgttca gatttcatac ttacgactat gcgaggcatt ttgtgagtgc ttgcactaga 3000
atacttggac ttgagggtac ccctgagggc gttgaagatc aaggaaggct aaccagggtt 3060
gcagcggtaa aaatcaaaac tgatgttttt atgagctgtg ttggacacat acttagtgta 3120
ttgttgccat atttccatta tgttgataat tccttatgga catttatttc tcatgttgat 3180
tatttatgag tcatccaaaa cgtatctgcg aaatgagtga atgattcttt tcattcactt 3240
ggctgaaatc ttcagtttcc tattgggata gactctgatc gtttcaagcg agcattggag 3300
cttccagcag tgaaaaggca cgtcagtgaa ttgacagaac gttttgccgg tcgaaaggta 3360
tgaagacact tgtcctttat ttactctata acaaacttaa gatccatcat tgtaagaatc 3420
agatatgaaa agtgaaaggg tctctagccg aattagtcag gtagtctgag tagcactttt 3480
caggtcttgg gttcgactcc ccgtgggagt gatttttcag gccgtggtaa aaaaatcccc 3540
tcgtctgttc cataccagtt cgggggtttc tcgatctgcg tgagaaggtc tttttcttgt 3600
tataatgtcg tgggtgcggt cttaattccc gtaggtcgat tttttttaag aattagattt 3660
gaaaacgagt ttatcaatat atttaggttg caaatggaaa gatttgtatc catttttttt 3720
atttgtatca tcttgaattt tgaatatagc ctttcgagtg tttaatattc tgggctagca 3780
tgagggattc acggtaatcc tctgtcaaat gcaaaatggt atttctggtt atgacttgtt 3840
attaggataa ggatctgtta tgtatggtgc ccaacccttg aatacaagaa taggagaaca 3900
tttctaaaca tttgaattgt tgcagtttgc ttctaggtga ctataacgag aaattcaaaa 3960
atgttcacct ctgaagaata aataccatat cggtctgaag ctttatacta catttggtct 4020
caaaacatgt tgcttatgaa aatatttggt caaaacttaa gaacttcaaa tataatagac 4080
tttcttggag atatgtagga tatttttgtt gatatgttat tagaaaaaca cttgcaaggg 4140
cagaggctgc ttcgaacccg taacctagta actcaatgag atagctttta ccattgcact 4200
aggtctgccc ttcagaaaaa cacttgcaaa atattataaa tttggaggac tctaagaatg 4260
tattctggaa ggtattagcg gcaaaaggta tggatcaaag accctgcaaa gccaagtgtt 4320
acatgttttt gtgactggag gtagtatcta aatgactcta ctgaaaagta gctggtcaat 4380
aatcaatgga ctggtaatag ctgagtggta gtgtgatact acatttccct gcattttata 4440
ttgataagca agcagtattt ctgttgtgct cactaaaaca tgtattatgt actcatcggt 4500
ttaacatacc atgcactcta caatactgtg ggtgatgaat gtcctggaag tctcatcttg 4560
tctaatgcta gctgatcata acctctcttg gtgacaggat actgccactg attttccttt 4620
tattctgctt agttctcctt gtatgttttt agtagtttta agtactcatc agttgttact 4680
tctcttaagc tttagccaat agccatcact attttaaccc cccacccccc ctgatgtatg 4740
ttattgttac ctatcacctg tgtaattcct ttaactttga gttttgtctt ttttgctggc 4800
attgagtata taatgtatat tggtgttcct tcatatagtt ttgtgttatt tgctaatgtg 4860
ataggtaatg cttggtgttg accgacttga catgattaag ggaattccgc aaaagatttt 4920
ggcctttgaa aagtttcttg aggaaaaccc agactggaac aacaaagttg ttctactgca 4980
gattgctgtg ccaacaagaa ctgacgtccc tgaatgtatg agcgctttct gtgttctttc 5040
cttcatttgc taatacttaa tgcaagtctg caagcatgtg tgcagcatag aaggttcaga 5100
aatatgattt gcttactcgc atgagttctt catttacagc ttgatatctt ctatatgaaa 5160
taaattttcc tttttcaaga cgcctgcagt gaaattattt tattgtcata ggaagcttta 5220
tttgggtatt tattcatttc ctttaaagtg tcggcagtac aatatacatg tttactcctc 5280
tgaataatga gacagatgat ccttctgcat ttttgaacgc tattgcattc tttattttgt 5340
aaagagctta ctcaacttcc acagaaacct ttgtggaaat ccaaaacaaa tgtgaccaga 5400
ggaagcactt tgtatgctac catgctgtgt gaaaaaaatc atgttaatga atgcctttat 5460
aagtgtttca tgcatacttg gtcaaagttt taggctgtct acaacttaac gaacctacat 5520
ttgtttgtgt ctggttatta tttataagaa ataagactta ttagaaatca gtaatcactt 5580
ttgctaggtg tgccatgtat cccataacac cacttatttt caaaatgggt ttaatgatct 5640
tatgagaacc ttgattggga gaaatagatt aacatattgc attacaactg attttttgct 5700
ggttttgtac catacttcta tttatctgtt cttcaagttc aacatgattg ttttttatca 5760
ctgaataact actcactaga tttgcagatc aaaagctaac gagccaagtg catgaaattg 5820
ttgggcgcat aaacggtcga tttggaacgt tgactgctgt ccctattcat catctggtag 5880
ttttcacatt tcccattgcg gcctctgttt ctgtattagg acctaacttt tcttaagcgt 5940
tataccacac tgctgagaag gcactaccca tttgtaaggt tcaattttga tttgctgttc 6000
taagtattat gaataagtat caatgtttta aatccttgca taaaaagagc ttccatgcat 6060
tttctgtaca agaatctgaa acagtgtgtt ccttgcagga ccgatctctt gatttccatg 6120
ccttgtgtgc tctttatgca gtcactggta atcatttttg ggcagatgtg ttttcttcca 6180
cttcggtata ccagatttac accagagtta ttggattttt taaaactatc ttaattttca 6240
ctgtttttgt tcagatgttg ctcttgtaac atcactgaga gatgggatga accttgtgag 6300
ctatgaatat gttgcatgcc aagggtctaa gaaaggagtt ctgatactta gcgaggtagc 6360
tgatcgatta tgcatgactt ataatcaagg ggtggaggat gttttgttta tttacttgtt 6420
tttgctatca ctgcaatttc agtttgctgg ggcagcacaa tcacttggag ctggtgccat 6480
tctagtaaac ccttggaata ttacagaagt tgcagactca atacggcatg ctttaacgat 6540
gccatccgat gagagagaga aacgacacag acacaactac gcacatgtca caactcacac 6600
ggctcaagat tgggctgaaa cttttgtatt gtgagtttct catttccatg gcccaatatc 6660
attcattcgg tggaccaagc cccttcactt ataattttgt ttgcaactca agttttctgg 6720
tgattatcct tttctttttc atgtagtgag ctaaatgaca cggttgctga agcactactg 6780
aggacaagac aagttcctcc tggtcttcct agtcaaatgg caattcagca atatttgcgc 6840
tctaaaaatc gtctgctcat attggtatta ctatatttct tattgcatac ttgatagtca 6900
tggacatttt tttccatcat aataactggg tgttctagtc agaaaatgac tcacttgtat 6960
ccggtagcaa ctctgttcta tgtagagtca cttcgaacag ttgcttcttt tgctcttctg 7020
cagctactgt cttcattctg gttgttgcct tacattttct ggtagactag atcatcatgg 7080
cacatttgtg gcatatcaag tctccagtaa atctttcact ttcgaatttc cacaagaagg 7140
ctttctggac ccatgaactc tcaactactc tatgagtcac attcgctgtt atttgacagt 7200
aacaatttca acaattcctg ttatctatac atgaccggat gtagggtttc aattcgacat 7260
tgactgagcc agtcgaatcc tctgggagaa ggggtggtga ccaaatcaag gaaatggaac 7320
tcaagttgca tcctgactta aagggtcctc tgagagccct ctgtgaggat gagcgcacta 7380
cagttattgt tcttagcggc agtgacagga gtgttcttga tgaagtaata ctcagagcac 7440
atctctgtta ttataggttt ggttgatatt ttccattaag accaatacat ctctccatgt 7500
gtacagaatt ttggagaatt taaaatgtgg ttggcggcag agcatgggat gtttttacgc 7560
ccgacttacg gagaatggat gacaacaatg cctgagcatc tgaacatgga ttgggttgac 7620
agcgtaaagg tttgtctgtt tcatgaataa tgttctgttg cgtcaaatta atagcaagtt 7680
tctgatattt ttttttctcc agcatgtttt tgaatacttt acagaaagaa ccccaagatc 7740
ccatttcgaa catcgtgaaa catcatttgt gtggaactat aagtatgctg gtaagtgttt 7800
atcttccaaa cattcttttt aagcaaggtc tgtgcagtaa cattttattt ccaccttcta 7860
tctcaaagat gttgagttcg gaaggctaca agcaagagat atgctgcagc acttgtggac 7920
aggtccgatc tcaaatgcag ctgttgatgt tgttcaaggg agtcgatcag ttgaagttcg 7980
gtctgttgga gttaccaagg taagtggtgc tgtatgatgg tccatgacag tagatggttt 8040
gacctagttg aggttttaaa catcatttgt ctccttccta tagggtgctg caattgatcg 8100
tattttaggg gagatagttc acagcgaaaa catgattact ccaattgact atgtcctgtg 8160
catagggcat ttccttggga aggtaatttc caattcttca cgttattgtt ctgatctggt 8220
tttagttctg tggataggat caagaaactt atgaatgaag aaatgaaagg gtttagaaca 8280
ttcaaaatgc attgccaagg aaaggagaag gatgtggact taattagaaa aatcactatg 8340
gtttacttta tatgtttggt ctgtccctat tcttttcgac tttggtctgt tttgtaacat 8400
ctggttcctc gtgtttcagg atgaggacat ctacgtcttc tttgatcccg agtacccttc 8460
tgaatccaaa gtaaagccag agggcggctc agcatcactt gaccggaggc cgaacgggag 8520
gccaccatcg aatggcagga gtaactccag gaacccacag tccaggacac agaaggcgca 8580
gcaggctgca tccgagaggt catcctcatc aagtcacagc agcacgagca gcaaccacga 8640
ctggcgcgaa gggtcctcgg tccttgatct caagggcgag aactacttct cctgcgccgt 8700
cgggaggaag cggtccaacg cccgctacct gctgagctcg tcggaggagg ttgtctcctt 8760
cctcaaagag ttggcgacag cgacagctgg cttccaggcc acctgtgctg actacatgca 8820
tgttcttgga taggcagtaa atagactgaa gttgaagcct ccgtgcttta ccagagacag 8880
agagaagaag aatattcatt cctcgtatgc gcgacagagc tacacccgta gctagtcagc 8940
gtgctgtaca atcatgtaca aaatttatgc tcgtgataaa actgcgagag gggagctagc 9000
aaatgggaaa ggataaagga gtttagttgc ttctggtacg agacacaatc gcctgatttt 9060
gagttctctt cgatttgttg tcctttaaat tgcgcttttg aactggtttg ggtgcacctg 9120
aaatacgcct gcgctatata tgagaccaca tctcatattt ttcatcttaa catcatatga 9180
cacgccaaat gaatctacct gcttattact agtttattta gagactgaaa tcttgtctag 9240
attatgccag ataattttgt ttaaaaaaac ttttctggat gcatgctcag agaaggtgat 9300
gttgaaagga ggtttggagt aacgtcaggc tgggcaagga tagattgtgc ccgtttggtt 9360
gagcttttgg ggggattctg gccgccagaa gctgaagcgg ctaccaaacg cccagcttct 9420
gctgccgctt ctccgcgccg cgcttggatg agaagccgcc cgcccacgcg acgccagaag 9480
cggggctcag atcgcttccg cggaaagcca aacctcattt cgctagggtt tcgcaatcgc 9540
gcccgtgacg cgtccccgtc ccaactcgcc gccgcaggtc tcctcccctt cgcatctccc 9600
gcaaccgcca tcattcgcga cgcccgcggc ggagttcgtg tcccccatcg catctcccgc 9660
cgccgcgctc gtcgtcttcg cagaggtaag ctttgttcct ctcatccatg gcgaacttga 9720
cttgtagctc ctccgccgtg tcggtgattc gatgcattcg tgacccttcc gttctctagg 9780
ggtgtgaccg cggctccgca 9800
<210> 2
<211> 9069
<212> DNA
<213> Zea mays
<400> 2
gcaaaacatc taacttttcg tataagcagc ttcttcaata atccatcaac gaccgttagg 60
agcggtgaca atctagctgt ctagaaaacg cgaaacttgc aagacccctc cgtcgctcgc 120
tcccataaaa gcctcaatcc ctccacccac ccacctcgcc tgcatatcac tcgtgtgctc 180
gcaaacaagc gtccgagttg gaggcgtggc ggcccgtttt gcatgccgta gctctcccgg 240
tgttcctccc cttccgaacc cctcgccgcc tctccgtctc gtgattcgaa tcactgcccc 300
attgcgaacg agccgccaca gctgctgcca gcctcaagga agggccccgg ttcccaatcc 360
cagacgggcc agcggatcga tcgaacaggc ggccggccac tcgggatcgg gtgggtgagt 420
cgttccgctt ccgaaaccag cctccctgcg tgatgggcct gcttgggtcc gcctgttgcc 480
gcctcctcgt cgtcgctcac cgctcgcgtc cgcctaatgc tgcaggccgg ggccgcggcg 540
gcgcatgagc tctgacgccg cgagggggca gcgcggcatc aactgcacga ggggcgacgc 600
ggcggcgatg ccaacctcat cgccctttgt cggcgacagc ggcggcgcgg gctccccgat 660
ccgcgtcgag cgaatggtcc gggagcgtag ccgccgctac gacatcttcg cgtcggacgc 720
gatggatacc gacgccgagg cggccttcgc gctggatggg gtccagtcgc ctggacgtgc 780
gtcacccgcc aacatggagg atgctggcgg cgcggccgca gcgcgaccgc cgctcgccgg 840
ctcccgcagc ggtttccgcc gcctaggcct ccgtggcatg aagcagcgcc tcctcgtcgt 900
ggccaaccgc ctgcctgtat ccgccaaccg ccgcggagag gaccactggt cgcttgagat 960
cagcgccggc ggccttgtga gcgccctcct tggtaatccc gacatttaat ctcctcccaa 1020
atgcgacgtg gcctggacct ggagatgcat gcaatggaat tctgcggtaa catctataac 1080
cttagttcat atgaatgttt tttttttctg atgctgaagg cgtgaaggac gtcgacgcga 1140
agtggattgg ctgggcaggc gtcaacgtac cagacgaggt tggtcagcga gccctcacta 1200
aagctcttgc ggagaaggtt ggttgcaatt ccatgttgtc tgggtcaata gaacagtgac 1260
tctttgttag tatcactctg ataactcgag gcatttttgt atggattgct tgaacttcgc 1320
agagatgcat accagtgttc ctggacgagg agattgtgca ccagtactac aatggatact 1380
gcaacaacat cctgtggcca ctgttccact acctaggact accacaggag gacaggctgg 1440
caacaacgag gaactttgag tcacagttcg acgcgtacaa gcgtgctaac cagatgttcg 1500
ctgatgtcgt gtacgagcac taccaggacg gggatgtgat ctggtgccat gactaccacc 1560
tcatgtttct gcccaagtgc ctcaaggacc atgacatcaa tatgaaggtc gggtggttcc 1620
tgcacacgcc gttcccgtca tcagagattt accggacact gccgtcccgc ttggagctgc 1680
tccggtcggt gctgtgtgcc gatttagttg ggtaagcata actatggatt ttgtgtcggc 1740
tgtcctgtcg aatttgattg ttaaacctga taagtttaac cgagtatcct ttttattctt 1800
gatttacttg aagtaagtcc caactcacaa cgtccggaaa ttggatattt cccttccact 1860
gtaaagttac tctctgatca gtgtatattt aatagttttt tttggtggtg ggaggggggt 1920
ggtcagacag ccaagtccta ctatatttgt aaccccccaa gaaagtttta gaatatgaaa 1980
tcaggtcaaa atgaaggttc taatttgaca gcatggaaaa tattgcctgg cagttcattc 2040
actgtataac atagaaagtg ttactataga ttgtcatgct tttgttatgt ttctgtgacg 2100
tattgctatc atggtaggaa tgtactagat acacgatttt ggtgatgtat tttttttagt 2160
ttgaactgac ttggcctata tatgaagttt gctttagatt caaaagacta aacctattta 2220
gcaaattttt agttttcttc acattttcaa gaaatttatt ctagaagagc caaaataata 2280
aaaaaaaggc caacacgtcg tgtgaaatta tcagtttcat ttctatgagt tatttcatgg 2340
tgctcttatt aattttgttc agatttcata cttacgacta tgcgaggcat tttgtgagtg 2400
cttgcactag aatacttgga cttgagggta cccctgaggg cgttgaagat caaggaaggc 2460
taaccagggt tgcagcggta aaaatcaaaa ctgatgtttt tatgagctgt gttggacaca 2520
tacttaaagt gtattgttgc catatttcca tcgttgataa ttccttatgg acatttattt 2580
ctcatgttga ttatttatga gttatccaaa acgtatctgc gaaatgagtg aatgattttt 2640
tcattcactt ggctgaaatc ttcagtttcc tattgggata gactctgatc gtttcaagcg 2700
agcattggag cttccagcag tgaaaaggca cgtcagtgaa ttgacagaac gttttgccgg 2760
tcgaaaggta tgaaaacact tgtcctttat ttactctaga acaaacttaa aatccatcat 2820
tgtaagaatc agatatgaaa agtgaaagag gctctagccg agttagttag gtggtttgag 2880
tagcactcct taggtcctgg gttcgactcc ctgtgggagc aaatttcaga ccgtggttta 2940
aaaaattctc tcgtctgtct cacgccaaag cacaggttta aggcctggtc ccggtcgcgg 3000
tcgttctcac atgtgctacg gtgctgctgc gtataagtgg ggcaggggtt tgagaggttt 3060
ctcgatctgc gtgagaagtt cttcttgtta taatgtcgtg ggggcggtct taatccccga 3120
aggtcgcgtt ttttttaaga atcagatatg aaaacgagtt tacccatata tttaggttgc 3180
acatggaaag atttgtatcc atttttttat ttgtatcatc ttgaattttg aatatagcct 3240
ttcgagtgtt taatattctg ggctagcatg aggaattcac ggtaatcctc tgtcaaatgc 3300
aaaatggtat ttctggttat gacttgttat taggataagg atcggtttgt atggtgccca 3360
atgcccttga atacaagaat aggagaacat ttctgaacat ttgaattgtt gcagtttgct 3420
tctaggtgac tataacgaga aattcaaaaa tgttcacctc tgaagaataa ataccatatc 3480
ggtctgaaac tttatactac ctttggtctc actttatact acctttggtc tcaaaacatg 3540
ttgcttatga aaatatttgg tcaaaactta agaacttaaa atataataaa ctttcttgga 3600
gatatgtagg atatttttgt tgatatgtat tagaaaaaca cttgcaaggg cagaggctgc 3660
ttcgaaccca taacctagta actcaatgag atatctttca ccattgcact aggtctgtcc 3720
ttcagaaaaa cacttgcaaa atatcataaa tttgaaggac tctaagaatg tattctagaa 3780
ggtattagcg gcagaaggta tggatcaaag accctgcaaa gccaagtgtt acatgttttt 3840
gtgactggag gtagtatcta aatgtctcta ctgaaaaata gctggtcaat aatcaatgga 3900
ctggttatag ctgagtggta gtgtgatact acatttccct gcattttata ttgataagca 3960
agcagtattt ctgttgtgct cactaaaaca tgtattatgc actcatcggt ttaacatacc 4020
atgcactcta caatactgtg cgtgatgaat gtcctggaag tctaatcttt tttcttgtct 4080
aatgctatct gatcataatc tctcttggtg acaggatact gccactgatt ttccttttat 4140
tctgcttagt tatccttgta tgtttttagt agttttaagt actcatcagt tgttacttct 4200
cttaagcttt agccaatagc catcactatt ttaaccccca ccccgttatt gttacctatc 4260
acctatgtaa ttcctttaac tttgagtttt gtcttttttg ctggcattga gtatattatt 4320
gttccttcat atagttttgt gttatttgct aatgtgatag gtaatgcttg gtgttgaccg 4380
acttgacatg attaaaggaa ttccgcaaaa gattttggcc tttgaaaagt ttcttgagga 4440
aaacccagac tggaacaaca aagttgttct actgcagatt gctgtgccaa caagaactga 4500
cgtccctgaa tgtatgagcg ctttctgtgt tcttcccttc atttgctaat actttatgca 4560
agtctgcaag catgtgtgca gcatagaagg ttcagaaata tgatttgctt actcgcatga 4620
gttcttcatt tacagcttga tatcttctat atgaaataaa atttcctttt tcaagacgcc 4680
tgcagtgaaa ttattttatt gtcataggag tgtttcatgc atacttggtc aaagttttag 4740
gctgtctaca acttaacgaa cctacatttg tttgtgtctg ttattattta taagaaataa 4800
gacttattag taatcagtaa tcacttttgc taggtgtgcc atgtatccca taacaccaat 4860
tcttttcaaa atgggtttaa tgatcttatg agaaccttga ttgggagaaa tagattaaca 4920
tattgcatta caactgaatt tttgctggtt ttgtaccata cttctattta tctgttcttc 4980
aagttcaaca tgattgtttt ttatcactga ataactactc actagatttg cagatcaaaa 5040
gctaacgagc caagtgcatg aaattgttgg gcgcataaac ggtcgatttg gaacgttgac 5100
tgctgtccct attcatcatc tggtagtttt cacatttccc attgctgcct ctgtatctgt 5160
attaggacct aacttttctt aagcgttata ccacactgct gagaaggcac tacccatttg 5220
taaggttcaa ttttgatttg ctgttctaag tattatgaat aagtatcaat gttataaatc 5280
cttgcataaa aagagcttcc atgcattttc tgtacaagaa tctgaaacag tgtgttcctt 5340
gcaggaccga tctcttgatt tccatgcctt gtgtgctctt tatgcagtca ctggtaatca 5400
ttaaaattct atttatactg acattttggg cagatgtgtt ttcttccact tcggtatacc 5460
agatttacac cagagttaat ggatttttta aaaaactatc ttaattttca ctgtttttgt 5520
tcagatgttg ctcttgtaac atcactgaga gatgggatga accttgtgag ctatgaatat 5580
gttgcatgcc aagggtctaa gaaaggagtt ctgatactta gcgaggtagc tgatcgatta 5640
tgcatgactt ataatcaagg ggtggaggat gttttgttta tttacttgtt tttgctatca 5700
ctgcaatttc agtttgctgg ggcagcacag tcacttggag ctggtgccat tctagtaaac 5760
ccttggaata ttacagaagt tgcagactca atacggcatg ctttaacgat gccatccgat 5820
gagagagaga aacgacacag acacaactac gcacatgtca caactcacac ggctcaagat 5880
tgggctgaaa cttttgtatt gtgagtttct catttccatg gcccaatatc attcattcgg 5940
tggaccaagc cccttcactt ataattttgt ttgcaactca agttttctgg tgattatcct 6000
tttctttttc atgtagtgag ctaaatgaca cggttgctga agcactactg aggacaagac 6060
aagttcctcc tggtcttcct agtcaaatgg caattcagca atatttgcgc tctaaaaatc 6120
gtctgctcat attggtatta ctatatttct tattgcatac ttgatagtca tggacatttt 6180
tttccatcgt aataactggg cgttctagtc agaaaatgac tcacttgcat ctggtagcaa 6240
ctctgttcta tgtagagtca cttcgaacag ttgcttcttt tgctcttctg cagctactgt 6300
cttcattctg gttgttgcct tacattttct ggtagactag atcatcatgg cacatttgtg 6360
gcatatcaag tctccagtaa aactttcaat ttcgaatttc cataagaagg ctttctggac 6420
ccatgaactc tcaactactc tgagtcacat tcgctgttat ttgacagtaa caatttcaat 6480
aattcttgtt atctatacat gaccggatgt agggtttcaa ttcgacattg actgagccag 6540
tcgaatcctc tgggagaagg ggtggtgacc aaatcaagga aatggaactc aagttgcatc 6600
ctgacttaaa gggtcctctg agagccctct gtgaggatga gcgcactaca gttattgttc 6660
ttagtggcag tgacaggagt gttcttgatg aagtaatact cagagcacat ctctgttatt 6720
ttaggtttgg ttgatatttt ccattaagac caatacatct ctccatgtgt acagaatttt 6780
ggagaattta aaatgtggtt ggcggcagag catgggatgt ttttacgccc gacttacgga 6840
gaatggatga caacaatgcc tgagcatctg aacatggatt gggttgacag cgtaaaggtt 6900
tgtctgtttc atgaataatg ttctgttgcg tcaaattaat agcaagtttc tgatatattt 6960
ttttctccag catgtttttg aatactttac agaaagaacc ccaagatccc atttcgaaca 7020
tcgtgaaaca tcatttgtgt ggaactataa gtatgctggt aagtgtttat cttccaaaca 7080
ttctttttaa gcgaggtctg tgcagtaaca ttttatttcc accttctatc tcaaagatgt 7140
tgagttcgga aggctacaag caagagatat gctgcagcac ttgtggacag gtccgatctc 7200
aaatgcagct gttgatgttg ttcaagggag tcgatcagtt gaagttcggt ctgttggagt 7260
taccaaggta agtggtgctg tatgatggtc catgacagta gatggtttgg cctagttgag 7320
gttttaaaca tcatttgtct ccttcctata gggtgctgca attgatcgta ttttagggga 7380
gatagttcac agcgaaaaca tgattactcc aattgactat gtcctgtgca tagggcattt 7440
ccttgggaag gtaatttcca attcttcacg ttattggtct gatctggttt tagttctgtg 7500
gataggatca agaaacttat gaaggaagaa atggaagggt ttagaacatt caaaatgcat 7560
tgccaaggaa aggagaagga tgtggactta attagaaaaa actactatgg tttactttat 7620
atgtttggtc tgtccctatt cttttcgact ttggtctgtt ttgtaacatg tgcttcctcg 7680
tgtttcagga tgaggacatc tacgtcttct ttgatcccga gtacccttct gaatccaaag 7740
taaagccaga gggcggctca gcatcacttg accggaggcc gaacgggagg ccaccatcga 7800
atggcaggag taactccagg aacccacagt ccaggacaca gaaggcgcag caggctgcat 7860
ccgagaggtc atcctcatca agtcacagca gcacgagcag caaccacgac tggcgcgaag 7920
ggtcctcggt ccttgatctc aagggcgaga actacttctc ctgcgccgtc gggaggaagc 7980
ggtccaacgc ccgctacttg ctgagctcgt cggaggaggt tgtctccttc ctcaaagagt 8040
tggcgacagc gacagctggc ttccaggcca cctgtgctga ctacatgcat gttcttggat 8100
aggcagtaaa tagactgaag ttgaagcctc cgtgctttac cagagacaga gagaagaaga 8160
atattcattc ctcgtatgcg cgacagagct acacccgtag ctagtcagcg tgctgtacaa 8220
tcatgtacaa aatttatgct cgtgataaaa ctgcgagagg ggagctagca aatgggaaag 8280
gataaaggag tttagttgct tctggtacga gacacaatcg cctgattttg agttctcttc 8340
gatttgttgt cctttaaatt gcgcttttga actggtttgg gtgcacctga aatacgcctg 8400
cgctatatat gagaccacat ctcatatttt tcatcttaac atcatatgac acgccaaatg 8460
aatctacctg cttattacta gtttatttag agactgaaat cttgtctaga ttatgccaga 8520
taattttgtt taaaaaaact tttctggatg catgctcaga gaaggtgatg ttgaaaggag 8580
gtttggagta acgtcaggct gggcaaggat agattgtgcc cgtttggttg agcttttggg 8640
gggattctgg ccgccagaag ctgaagcggc taccaaacgc ccagcttctg ctgccgcttc 8700
tccgcgccgc gcttggatga gaagccgccc gcccacgcga cgccagaagc ggggctcaga 8760
tcgcttccgc ggaaagccaa acctcatttc gctagggttt cgcaatcgcg cccgtgacgc 8820
gtccccgtcc caactcgccg ccgcaggtct cctccccttc gcatctcccg caaccgccat 8880
cattcgcgac gcccgcggcg gagttcgtgt cccccatcgc atctcccgcc gccgcgctcg 8940
tcgtcttcgc agaggtaagc tttgttcctc tcatccatgg cgaacttgac ttgtagctcc 9000
tccgccgtgt cggtgattcg atgcattcgt gacccttccg ttctctaggg gtgtgaccgc 9060
ggctccgca 9069
<210> 3
<211> 3462
<212> DNA
<213> Escherichia coli
<400> 3
atatcactcg tgtgctcgca aacaagcgtc cgagttggag gcgtggcggc ccgttttgca 60
tgccgtagct ctcccggtgt tcctcccctt ccgaacccct cgccgcctct ccgtctcgtg 120
attcgaatca ctgccccatt gcgaacgagc cgccacagct gctgccagcc tcaaggaagg 180
gccccggttc ccaatcccag acgggccagc ggatcgatcg aacaggcggc cggccactcg 240
ggatcgggtg ggtgagtcgt tccgcttccg aaaccagcct ccctgcgtga tgggcctgct 300
tgggtccgcc tgttgccgcc tcctcgtcgt cgctcaccgc tcgcgtccgc ctaatgctgc 360
aggccggggc cgcggcggcg catgagctct gacgccgcga gggggcagcg cggcatcaac 420
tgcacgaggg gcgacgcggc ggcgatgcca acctcatcgc cctttgtcgg cgacagcggc 480
ggcgcgggct ccccgatccg cgtcgagcga atggtccgcg agcgtagccg ccgctacgac 540
atcttcgcgt cggacgcgat ggataccgac gccgaggcgg ccttcgcgct ggatggggtc 600
cagtcgcctg gacgtgcgtc acccgccaac atggaggatg ctggcggcgc ggccgcagcg 660
cgaccgccgc tcgccggctc ccgcagcggt ttccgccgcc taggcctccg tggcatgaag 720
cagcgcctcc tcgtcgtggc caaccgcctg cctgtatccg ccaaccgccg cggagaggac 780
cactggtcgc ttgagatcag cgccggcggc cttgtgagcg ccctccttgg cgtgaaggac 840
gtcgacgcga agtggattgg ctgggcaggc gtcaacgtac cggacgaggt tggtcagcga 900
gccctcacta aagctcttgc ggagaagaga tgcataccag tgttcctgga cgaggagatt 960
gtgcaccagt actacaatgg atactgcaac aacatcctgt ggccactgtt ccactaccta 1020
ggactaccac aggaggacag gctggcaaca acgaggaact ttgagtcaca gttcgacgcg 1080
tacaagcgtg ctaaccagat gttcgctgat gtcgtgtacg agcactacca ggacggggat 1140
gtgatctggt gccatgacta ccacctcatg tttctgccca agtgcctcaa ggaccatgac 1200
atcaatatga aggtcgggtg gttcctgcac acgccgttcc cgtcatcaga gatttaccgg 1260
acactgccgt cccgcttgga gctgcttcgg tcggtgctgt gtgccgattt agttggattt 1320
catacttacg actatgcgag gcattttgtg agtgcttgca ctagaatact tggacttgag 1380
ggtacccctg agggcgttga agatcaagga aggctaacca gggttgcagc gtttcctatt 1440
gggatagact ctgatcgttt caagcgagca ttggagcttc cagcagtgaa aaggcacgtc 1500
agtgaattga cagaacgttt tgccggtcga aaggtaatgc ttggtgttga ccgacttgac 1560
atgattaagg gaattccgca aaagattttg gcctttgaaa agtttcttga ggaaaaccca 1620
gactggaaca acaaagttgt tctactgcag attgctgtgc caacaagaac tgacgtccct 1680
gaatatcaaa agctaacgag ccaagtgcat gaaattgttg ggcgcataaa cggtcgattt 1740
ggaacgttga ctgctgtccc tattcatcat ctggaccgat ctcttgattt ccatgccttg 1800
tgtgctcttt atgcagtcac tgatgttgct cttgtaacat cactgagaga tgggatgaac 1860
cttgtgagct atgaatatgt tgcatgccaa gggtctaaga aaggagttct gatacttagc 1920
gagtttgctg gggcagcaca atcacttgga gctggtgcca ttctagtaaa cccttggaat 1980
attacagaag ttgcagactc aatacggcat gctttaacga tgccatccga tgagagagag 2040
aaacgacaca gacacaacta cgcacatgtc acaactcaca cggctcaaga ttgggctgaa 2100
acttttgtat ttgagctaaa tgacacggtt gctgaagcac tactgaggac aagacaagtt 2160
cctcctggtc ttcctagtca aatggcaatt cagcaatatt tgcgctctaa aaatcgtctg 2220
ctcatattgg gtttcaattc gacattgact gagccagtcg aatcctctgg gagaaggggt 2280
ggtgaccaaa tcaaggaaat ggaactcaag ttgcatcctg acttaaaggg tcctctgaga 2340
gccctctgtg aggatgagcg cactacagtt attgttctta gcggcagtga caggagtgtt 2400
cttgatgaaa attttggaga atttaaaatg tggttggcgg cagagcatgg gatgttttta 2460
cgcccgactt acggagaatg gatgacaaca atgcctgagc atctgaacat ggattgggtt 2520
gacagcgtaa agcatgtttt tgaatacttt acagaaagaa ccccaagatc ccatttcgaa 2580
catcgtgaaa catcatttgt gtggaactat aagtatgctg atgttgagtt cggaaggcta 2640
caagcaagag atatgctgca gcacttgtgg acaggtccga tctcaaatgc agctgttgat 2700
gttgttcaag ggagtcgatc agttgaagtt cggtctgttg gagttaccaa gggtgctgca 2760
attgatcgta ttttagggga gatagttcac agcgaaaaca tgattactcc aattgactat 2820
gtcctgtgca tagggcattt ccttgggaag gatgaggaca tctacgtctt ctttgatccc 2880
gagtaccctt ctgaatccaa agtaaagcca gagggcggct cagcatcact tgaccggagg 2940
ccgaacggga ggccaccatc gaatggcagg agtaactcca ggaacccaca gtccaggaca 3000
cagaaggcgc agcaggctgc atccgagagg tcatcctcat caagtcacag cagcacgagc 3060
agcaaccacg actggcgcga agggtcctcg gtccttgatc tcaagggcga gaactacttc 3120
tcctgcgccg tcgggaggaa gcggtccaac gcccgctacc tgctgagctc gtcggaggag 3180
gttgtctcct tcctcaaaga gttggcgaca gcgacagctg gcttccaggc cacctgtgct 3240
gactacatgc atgttcttgg ataggcagta aatagactga agttgaagcc tccgtgcttt 3300
accagagaca gagagaagaa gaatattcat tcctcgtatg cgcgacagag ctacacccgt 3360
agctagtcag cgtgctgtac aatcatgtac aaaatttatg ctcgtgataa aactgcgaga 3420
ggggagctag caaatgggaa aggataaagg agtttagttg ct 3462
<210> 4
<211> 960
<212> PRT
<213> Zea mays
<400> 4
Met Ser Ser Asp Ala Ala Arg Gly Gln Arg Gly Ile Asn Cys Thr Arg
1 5 10 15
Gly Asp Ala Ala Ala Met Pro Thr Ser Ser Pro Phe Val Gly Asp Ser
20 25 30
Gly Gly Ala Gly Ser Pro Ile Arg Val Glu Arg Met Val Arg Glu Arg
35 40 45
Ser Arg Arg Tyr Asp Ile Phe Ala Ser Asp Ala Met Asp Thr Asp Ala
50 55 60
Glu Ala Ala Phe Ala Leu Asp Gly Val Gln Ser Pro Gly Arg Ala Ser
65 70 75 80
Pro Ala Asn Met Glu Asp Ala Gly Gly Ala Ala Ala Ala Arg Pro Pro
85 90 95
Leu Ala Gly Ser Arg Ser Gly Phe Arg Arg Leu Gly Leu Arg Gly Met
100 105 110
Lys Gln Arg Leu Leu Val Val Ala Asn Arg Leu Pro Val Ser Ala Asn
115 120 125
Arg Arg Gly Glu Asp His Trp Ser Leu Glu Ile Ser Ala Gly Gly Leu
130 135 140
Val Ser Ala Leu Leu Gly Val Lys Asp Val Asp Ala Lys Trp Ile Gly
145 150 155 160
Trp Ala Gly Val Asn Val Pro Asp Glu Val Gly Gln Arg Ala Leu Thr
165 170 175
Lys Ala Leu Ala Glu Lys Arg Cys Ile Pro Val Phe Leu Asp Glu Glu
180 185 190
Ile Val His Gln Tyr Tyr Asn Gly Tyr Cys Asn Asn Ile Leu Trp Pro
195 200 205
Leu Phe His Tyr Leu Gly Leu Pro Gln Glu Asp Arg Leu Ala Thr Thr
210 215 220
Arg Asn Phe Glu Ser Gln Phe Asp Ala Tyr Lys Arg Ala Asn Gln Met
225 230 235 240
Phe Ala Asp Val Val Tyr Glu His Tyr Gln Asp Gly Asp Val Ile Trp
245 250 255
Cys His Asp Tyr His Leu Met Phe Leu Pro Lys Cys Leu Lys Asp His
260 265 270
Asp Ile Asn Met Lys Val Gly Trp Phe Leu His Thr Pro Phe Pro Ser
275 280 285
Ser Glu Ile Tyr Arg Thr Leu Pro Ser Arg Leu Glu Leu Leu Arg Ser
290 295 300
Val Leu Cys Ala Asp Leu Val Gly Phe His Thr Tyr Asp Tyr Ala Arg
305 310 315 320
His Phe Val Ser Ala Cys Thr Arg Ile Leu Gly Leu Glu Gly Thr Pro
325 330 335
Glu Gly Val Glu Asp Gln Gly Arg Leu Thr Arg Val Ala Ala Phe Pro
340 345 350
Ile Gly Ile Asp Ser Asp Arg Phe Lys Arg Ala Leu Glu Leu Pro Ala
355 360 365
Val Lys Arg His Val Ser Glu Leu Thr Glu Arg Phe Ala Gly Arg Lys
370 375 380
Val Met Leu Gly Val Asp Arg Leu Asp Met Ile Lys Gly Ile Pro Gln
385 390 395 400
Lys Ile Leu Ala Phe Glu Lys Phe Leu Glu Glu Asn Pro Asp Trp Asn
405 410 415
Asn Lys Val Val Leu Leu Gln Ile Ala Val Pro Thr Arg Thr Asp Val
420 425 430
Pro Glu Tyr Gln Lys Leu Thr Ser Gln Val His Glu Ile Val Gly Arg
435 440 445
Ile Asn Gly Arg Phe Gly Thr Leu Thr Ala Val Pro Ile His His Leu
450 455 460
Asp Arg Ser Leu Asp Phe His Ala Leu Cys Ala Leu Tyr Ala Val Thr
465 470 475 480
Asp Val Ala Leu Val Thr Ser Leu Arg Asp Gly Met Asn Leu Val Ser
485 490 495
Tyr Glu Tyr Val Ala Cys Gln Gly Ser Lys Lys Gly Val Leu Ile Leu
500 505 510
Ser Glu Phe Ala Gly Ala Ala Gln Ser Leu Gly Ala Gly Ala Ile Leu
515 520 525
Val Asn Pro Trp Asn Ile Thr Glu Val Ala Asp Ser Ile Arg His Ala
530 535 540
Leu Thr Met Pro Ser Asp Glu Arg Glu Lys Arg His Arg His Asn Tyr
545 550 555 560
Ala His Val Thr Thr His Thr Ala Gln Asp Trp Ala Glu Thr Phe Val
565 570 575
Phe Glu Leu Asn Asp Thr Val Ala Glu Ala Leu Leu Arg Thr Arg Gln
580 585 590
Val Pro Pro Gly Leu Pro Ser Gln Met Ala Ile Gln Gln Tyr Leu Arg
595 600 605
Ser Lys Asn Arg Leu Leu Ile Leu Gly Phe Asn Ser Thr Leu Thr Glu
610 615 620
Pro Val Glu Ser Ser Gly Arg Arg Gly Gly Asp Gln Ile Lys Glu Met
625 630 635 640
Glu Leu Lys Leu His Pro Asp Leu Lys Gly Pro Leu Arg Ala Leu Cys
645 650 655
Glu Asp Glu Arg Thr Thr Val Ile Val Leu Ser Gly Ser Asp Arg Ser
660 665 670
Val Leu Asp Glu Asn Phe Gly Glu Phe Lys Met Trp Leu Ala Ala Glu
675 680 685
His Gly Met Phe Leu Arg Pro Thr Tyr Gly Glu Trp Met Thr Thr Met
690 695 700
Pro Glu His Leu Asn Met Asp Trp Val Asp Ser Val Lys His Val Phe
705 710 715 720
Glu Tyr Phe Thr Glu Arg Thr Pro Arg Ser His Phe Glu His Arg Glu
725 730 735
Thr Ser Phe Val Trp Asn Tyr Lys Tyr Ala Asp Val Glu Phe Gly Arg
740 745 750
Leu Gln Ala Arg Asp Met Leu Gln His Leu Trp Thr Gly Pro Ile Ser
755 760 765
Asn Ala Ala Val Asp Val Val Gln Gly Ser Arg Ser Val Glu Val Arg
770 775 780
Ser Val Gly Val Thr Lys Gly Ala Ala Ile Asp Arg Ile Leu Gly Glu
785 790 795 800
Ile Val His Ser Glu Asn Met Ile Thr Pro Ile Asp Tyr Val Leu Cys
805 810 815
Ile Gly His Phe Leu Gly Lys Asp Glu Asp Ile Tyr Val Phe Phe Asp
820 825 830
Pro Glu Tyr Pro Ser Glu Ser Lys Val Lys Pro Glu Gly Gly Ser Ala
835 840 845
Ser Leu Asp Arg Arg Pro Asn Gly Arg Pro Pro Ser Asn Gly Arg Ser
850 855 860
Asn Ser Arg Asn Pro Gln Ser Arg Thr Gln Lys Ala Gln Gln Ala Ala
865 870 875 880
Ser Glu Arg Ser Ser Ser Ser Ser His Ser Ser Thr Ser Ser Asn His
885 890 895
Asp Trp Arg Glu Gly Ser Ser Val Leu Asp Leu Lys Gly Glu Asn Tyr
900 905 910
Phe Ser Cys Ala Val Gly Arg Lys Arg Ser Asn Ala Arg Tyr Leu Leu
915 920 925
Ser Ser Ser Glu Glu Val Val Ser Phe Leu Lys Glu Leu Ala Thr Ala
930 935 940
Thr Ala Gly Phe Gln Ala Thr Cys Ala Asp Tyr Met His Val Leu Gly
945 950 955 960
<210> 5
<211> 20
<212> DNA
<213> Zea mays
<400> 5
gttataccac actgctgaga 20
<210> 6
<211> 20
<212> DNA
<213> Zea mays
<400> 6
ggggagctag caaatgggaa 20
<210> 7
<211> 103
<212> RNA
<213> unknown(人工合成)
<400> 7
guuauaccac acugcugaga guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu uuu 103
<210> 8
<211> 103
<212> RNA
<213> unknown(人工合成)
<400> 8
ggggagcuag caaaugggaa guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu uuu 103
<210> 9
<211> 3555
<212> DNA
<213> Zea mays
<400> 9
tgctaggtgt gccatgtatc ccataacacc aattcttttc aaaatgggtt taatgatctt 60
atgagaacct tgattgggag aaatagatta acatattgca ttacaactga atttttgctg 120
gttttgtacc atacttctat ttatctgttc ttcaagttca acatgattgt tttttatcac 180
tgaataacta ctcactagat ttgcagatca aaagctaacg agccaagtgc atgaaattgt 240
tgggcgcata aacggtcgat ttggaacgtt gactgctgtc cctattcatc atctggtagt 300
tttcacattt cccattgctg cctctgtatc tgtattagga cctaactttt cttaagcgtt 360
ataccacact gctcatttgc tagctcccct ctcgcagttt tatcacgagc ataaattttg 420
tacatgattg tacagcacgc tgactagcta cgggtgtagc tctgtcgcgc atacgaggaa 480
tgaatattct tcttctctct gtctctggta aagcacggag gcttcaactt cagtctattt 540
actgcctatc caagaacatg catgtagtca gcacaggtgg cctggaagcc agctgtcgct 600
gtcgccaact ctttgaggaa ggagacaacc tcctccgacg agctcagcaa gtagcgggcg 660
ttggaccgct tcctcccgac ggcgcaggag aagtagttct cgcccttgag atcaaggacc 720
gaggaccctt cgcgccagtc gtggttgctg ctcgtgctgc tgtgacttga tgaggatgac 780
ctctcggatg cagcctgctg cgccttctgt gtcctggact gtgggttcct ggagttactc 840
ctgccattcg atggtggcct cccgttcggc ctccggtcaa gtgatgctga gccgccctct 900
ggctttactt tggattcaga agggtactcg ggatcaaaga agacgtagat gtcctcatcc 960
tgaaacacga ggaagcacat gttacaaaac agaccaaagt cgaaaagaat agggacagac 1020
caaacatata aagtaaacca tagtagtttt ttctaattaa gtccacatcc ttctcctttc 1080
cttggcaatg cattttgaat gttctaaacc cttccatttc ttccttcata agtttcttga 1140
tcctatccac agaactaaaa ccagatcaga ccaataacgt gaagaattgg aaattacctt 1200
cccaaggaaa tgccctatgc acaggacata gtcaattgga gtaatcatgt tttcgctgtg 1260
aactatctcc cctaaaatac gatcaattgc agcaccctat aggaaggaga caaatgatgt 1320
ttaaaacctc aactaggcca aaccatctac tgtcatggac catcatacag caccacttac 1380
cttggtaact ccaacagacc gaacttcaac tgatcgactc ccttgaacaa catcaacagc 1440
tgcatttgag atcggacctg tccacaagtg ctgcagcata tctcttgctt gtagccttcc 1500
gaactcaaca tctttgagat agaaggtgga aataaaatgt tactgcacag acctcgctta 1560
aaaagaatgt ttggaagata aacacttacc agcatactta tagttccaca caaatgatgt 1620
ttcacgatgt tcgaaatggg atcttggggt tctttctgta aagtattcaa aaacatgctg 1680
gagaaaaaaa tatatcagaa acttgctatt aatttgacgc aacagaacat tattcatgaa 1740
acagacaaac ctttacgctg tcaacccaat ccatgttcag atgctcaggc attgttgtca 1800
tccattctcc gtaagtcggg cgtaaaaaca tcccatgctc tgccgccaac cacattttaa 1860
attctccaaa attctgtaca catggagaga tgtattggtc ttaatggaaa atatcaacca 1920
aacctaaaat aacagagatg tgctctgagt attacttcat caagaacact cctgtcactg 1980
ccactaagaa caataactgt agtgcgctca tcctcacaga gggctctcag aggacccttt 2040
aagtcaggat gcaacttgag ttccatttcc ttgatttggt caccacccct tctcccagag 2100
gattcgactg gctcagtcaa tgtcgaattg aaaccctaca tccggtcatg tatagataac 2160
aagaattatt gaaattgtta ctgtcaaata acagcgaatg tgactcagag tagttgagag 2220
ttcatgggtc cagaaagcct tcttatggaa attcgaaatt gaaagtttta ctggagactt 2280
gatatgccac aaatgtgcca tgatgatcta gtctaccaga aaatgtaagg caacaaccag 2340
aatgaagaca gtagctgcag aagagcaaaa gaagcaactg ttcgaagtga ctctacatag 2400
aacagagttg ctaccagatg caagtgagtc attttctgac tagaacgccc agttattacg 2460
atggaaaaaa atgtccatga ctatcaagta tgcaataaga aatatagtaa taccaatatg 2520
agcagacgat ttttagagcg caaatattgc tgaattgcca tttgactagg aagaccagga 2580
ggaacttgtc ttgtcctcag tagtgcttca gcaaccgtgt catttagctc actacatgaa 2640
aaagaaaagg ataatcacca gaaaacttga gttgcaaaca aaattataag tgaaggggct 2700
tggtccaccg aatgaatgat attgggccat ggaaatgaga aactcacaat acaaaagttt 2760
cagcccaatc ttgagccgtg tgagttgtga catgtgcgta gttgtgtctg tgtcgtttct 2820
ctctctcatc ggatggcatc gttaaagcat gccgtattga gtctgcaact tctgtaatat 2880
tccaagggtt tactagaatg gcaccagctc caagtgactg tgctgcccca gcaaactgaa 2940
attgcagtga tagcaaaaac aagtaaataa acaaaacatc ctccacccct tgattataag 3000
tcatgcataa tcgatcagct acctcgctaa gtatcagaac tcctttctta gacccttggc 3060
atgcaacata ttcatagctc acaaggttca tcccatctct cagtgatgtt acaagagcaa 3120
catctgaaca aaaacagtga aaattaagat agttttttaa aaaatccatt aactctggtg 3180
taaatctggt ataccgaagt ggaagaaaac acatctgccc aaaatgtcag tataaataga 3240
attttaatga ttaccagtga ctgcataaag agcacacaag gcatggaaat caagagatcg 3300
gtcctgcaag gaacacactg tttcagattc ttgtacagaa aatgcatgga agctcttttt 3360
atgcaaggat ttataacatt gatacttatt cataatactt agaacagcaa atcaaaattg 3420
aaccttacaa atgggtagtg ccttctcgga aaggataaag gagtttagtt gcttctggta 3480
cgagacacaa tcgcctgatt ttgagttctc ttcgatttgt tgtcctttaa attgcgcttt 3540
tgaactggtt tgggt 3555
<210> 10
<211> 21
<212> DNA
<213> unknown(人工合成)
<400> 10
tgctaggtgt gccatgtatc c 21
<210> 11
<211> 21
<212> DNA
<213> unknown(人工合成)
<400> 11
acccaaacca gttcaaaagc g 21
Claims (10)
1.一种控制玉米花期的基因及其在改变玉米花期性状中的应用，其特征在于：所述基因的核酸序列如SEQ ID NO.1-SEQ ID NO.3任意一个所示；或所述基因的核酸序列序列如SEQ ID NO.1-SEQ ID NO.3任意一个所示的序列经过取代和/或缺失和/或添加一个或多个氨基酸，且具有与SEQ ID NO.1-SEQ ID NO.3所示的序列相同功能的序列。
2.一种缩短玉米花期的方法，其特征在于：抑制玉米中权利要求1所述基因编码蛋白的表达和/或活性，选择玉米花期缩短的植株。
3.根据权利要求2所述缩短花期的方法，其特征在于：所述蛋白的氨基酸序列如SEQ IDNO.4所示。
4.根据权利要求2所述缩短花期的方法，其特征在于：所述抑制蛋白表达和/或活性的方法包括基因编辑、RNA干扰、T-DNA插入、物理或化学诱变中任一种。
5.根据权利要求4所述缩短花期的方法，其特征在于：所述基因编辑采用CRISPR/Cas9方法。
6.根据权利要求5所述缩短玉米花期的方法，其特征在于：所述CRISPR/Cas9方法在玉米中的基因组靶标区域的DNA序列如SEQ ID NO.5或SEQ ID NO.6所示。
7.一种用于缩短花期的试剂盒，其特征在于：包括如下任一种：
(1)sgRNA分子，其序列如SEQ ID NO.7或SEQ ID NO.8所示；
(2)所述sgRNA的编码DNA分子；
(3)表达所述sgRNA的载体。
8.一种玉米突变基因型在缩短玉米花期中的应用，其特征在于：所述突变基因型序列如SEQ ID NO.9所示。
9.用于检测权利要求8所述基因型的引物对，其特征在于：所述引物对为SEQ ID NO:10和SEQ ID NO:11所示的序列或其互补序列。
10.检测权利要求8所述基因型的方法，其特征在于：利用权利要求9所述的引物对检测待测样品中是否存在所述基因型。
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| CN112646016A (zh) * | 2021-01-22 | 2021-04-13 | 华中农业大学 | 改变玉米开花期的基因及方法 |
| CN112646014A (zh) * | 2021-01-22 | 2021-04-13 | 华中农业大学 | 改变玉米开花期的基因及方法 |
| CN112661823A (zh) * | 2021-01-22 | 2021-04-16 | 华中农业大学 | 改变玉米开花期的基因及方法 |
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| CN112662687A (zh) * | 2021-01-22 | 2021-04-16 | 华中农业大学 | 推迟玉米花期的方法、试剂盒、基因 |
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| CN112662687B (zh) * | 2021-01-22 | 2023-09-26 | 华中农业大学 | 推迟玉米花期的方法、试剂盒、基因 |
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