CN114269933A - 通过使用grf1加强基因实现的增强的植物再生及转化 - Google Patents
通过使用grf1加强基因实现的增强的植物再生及转化 Download PDFInfo
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Abstract
本发明涉及植物育种以及生物技术领域，尤其涉及从细胞以及其它组织产生植物。更具体而言，本发明提供了用于改善特别是来自使用GRF1加强基因的转化的或遗传修饰的植物细胞的植物再生的方法及工具。
Description
本发明涉及植物育种以及生物技术领域，尤其涉及从细胞以及其它组织产生植物。更具体而言，本发明提供了用于改善特别是来自转化的或遗传修饰的植物细胞的植物再生的方法及工具。
在植物育种中，为了形成用于特定目的的所需基因型及表型，自接近人类文明开始以来就已经实践了植物物种的操纵过程。随着基因工程的发展，近几十年来，这一农业领域发生了重大变化。已经开发出了用于植物基因工程的各种方法。转化方法的选择取决于许多变量，主要是要转化的植物物种、实验目的以及必需设备的可用性。大多数植物转化技术需要使用具有高再生能力的外植体作为起始材料。另外，基因编辑构成了一种新的分子生物学方法，通过该方法可以将诸如插入、缺失或点突变或其组合之类的特异性修饰引入植物的基因组中。为此，需要特异性分子工具，其首先具有核酸酶活性，但最重要的是可以以足够的特异性引导至待修饰的靶序列以编程并进行特异性的定点诱变。在过去几年中，在植物生物技术中，特异性基因组编辑已经发展成为常规育种以及转基因策略的替代。然而，由于植物的经编辑的起始材料的再生能力有限，因此目前可获得的工具，诸如大范围核酸酶、锌指核酸酶(ZFN)、“转录激活因子样效应物核酸酶”(TALEN)或CRISPR系统，仅在有限程度上用于植物生物技术中。
多种细胞具有发育成胚的潜力，包括单倍体配子体细胞，诸如花粉及胚囊的细胞(参见Forster,B.P.等(2007)Trends Plant Sci.12:368-375以及Seguí-Simarro,J.M.(2010)Bot.Rev.76:377-404)以及衍生自植物的所有三个基本组织层的体细胞(Gaj,M.D(2004)Plant Growth Regul.43:27-47或Rose,R.等(2010)在Plant DevelopmentalBiology-Biotechnological Perspectives中的“Developmental biology of somaticembryogenesis”，Pua E-C以及Davey MR编辑(Berlin Heidelberg:Springer)，第3-26页)。
再生为植物的能力通常限于某些植物物种中的特定基因型，并且在转化以及遗传修饰的植物细胞及其它前体组织中减弱。即使植物细胞的转化及遗传修饰步骤是成功的，这并不一定意味着实际上可以从所修饰的细胞获得所需植物。假定为了实现遗传修饰而使植物细胞及其它前体组织经受的处理影响植物发育及再生。因此，本发明的目的是提高先前已知的用于产生转基因及遗传修饰植物的方法的功效，并支持从修饰的植物细胞及其它植物前体再生植物。
Lowe K等(2016,“形态形成调节因子Baby boom及Wuschel提高单子叶植物转化(Morphogenic regulators Baby boom and Wuschel improve monocottransformation)”Plant Cell 28:1998-2015.)报道了一种利用形态形成基因产生有助于转化效率的生物学背景的转化方法。它们的转化构建体包括玉米(玉蜀黍)Baby boom(BBM)以及Wuschel2(WUS2)。作者将这些构建体用于来自四个玉米自交系的未成熟胚的农杆菌介导的转化。这些系表现出明显较高的转化率，从对照中的0％至2％到在WUS2及BBM存在下的25％至51％。然而，该方法具有缺点并且通常不适用，因为构成性表达ZmWUS2以及ZmBBM的植物通常是不育的或甚至致死的。因此，希望有一种用于提高转化及再生效率的方法，该方法会导致不显示可识别的表型的植物。
在van der Knaap等(2000；“来自水稻的新型赤霉素诱导基因以及它在茎生长中的潜在调节作用(A novel gibberellin-induced gene from rice and its potentialregulatory role in stem growth)”，Plant physiology，122(3)，695-704.)中，作者已经鉴定并表征了水稻中的GRF基因家族的第一成员(OsGRF1)。这是居间分生组织中的赤霉酸诱导基因。拟南芥中的过表达导致茎生长受损、雌性不育以及雄性能育性降低。施用赤霉酸无法恢复转化植物的茎伸长缺陷，表明OsGRF1可以参与GA诱导的茎伸长。在2003年，Kim等(“推定转录因子的AtGRF家族参与拟南芥中的叶子以及子叶生长(The AtGRF family ofputative transcription factors is involved in leaf and cotyledon growth inArabidopsis)”，The Plant Journal，36(1)，94-104.)已经表征了拟南芥GRF家族并确定了基因主要在活跃生长的组织中表达。对过表达At GRF1以及AtGRF2的无效突变体及转基因植物的分析表明，GRF家族的一些成员参与在叶子以及子叶生长期间的细胞扩增的调节。另外，过表达植物显示出延迟的抽苔时间，揭示了在开花中的推定作用。在确定协调正在发育的叶子中的细胞增殖的分子机制的研究中，Rodriguez等发现miR396在拟南芥中拮抗其靶标GRF转录因子的表达模式((2010)，“Control of cell proliferation in Arabidopsisthaliana by microRNA miR396”，Development，137(1)，103-112.)。因此，miR396与GRF之间的平衡控制叶子中的细胞的最终数目。此外，作者显示出miR396靶向的GRF可以调节茎尖分生组织的大小。
Kuijt等(2014，“Interaction between the GROWTH-REGULATING FACTOR andKNOTTED1-LIKE HOMEOBOX Families of Transcription Factors”，Plantphysiology，164(4)，1952-1966.)显示出GRF家族的成员在控制参与茎尖分生组织中细胞分化的限制的KNOTTED1-LIKE HOMEBOX(KNOX)基因的表达的网络中充当参与者。AtGRF4、AtGRF1以及AtGRF6能够结合KNOX基因的启动子，从而抑制其表达。过表达AtGRF4、AtGRF1或AtGRF6的拟南芥幼苗在茎尖分生组织中表现出发育畸变。
Nelissen H等(2015，“ANGUSTIFOLIA3复合物组成中的动态变化揭示了玉米叶子中的生长调节机制。(Dynamic Changes in ANGUSTIFOLIA3 Complex Composition Reveala Growth Regulatory Mechanism in the Maize Leaf.)”，Plant Cell 287:1605-1619.)描述了ZmGRF1在玉米叶子发育期间的功能。
总之，众所周知，几种GRF基因在叶形态发生以及茎发育中起作用。另外，据报道，GRF基因在开花、种子及根发育中起作用，在应激条件下控制植物生长并且调节植物寿命。
然而，在他们的研究中，本发明的发明人惊奇地发现了GRF1(GRF1)的另一种新的功能。GRF1能够提供促进植物再生的积极效果，并且因此允许转基因植物更有效地恢复。本发明允许改善从不同组织或细胞(例如，小孢子)再生，可以克服对植物再生的抗性，特别是基因型依赖性，通过例如目的基因与GRF1的共表达来改善转基因植物的恢复，以及通过例如基因组编辑组分与GRF1的瞬时共表达来改善基因组工程化植物的恢复，并且缩短产生转基因系以及恢复的时间。进一步，发现携带GRF1的植物是健康的并且不显示任何明显的表型。因此，本发明的第一方面是GRF1多肽用于改善植物的再生能力的用途。
下文中对可用于本发明方法的多肽或蛋白质的任何提及都是指如本文所定义的GRF1多肽或GRF1蛋白质。下文中对可用于本发明方法的核酸或多核苷酸的任何提及(除了被转化的感兴趣的核苷酸序列或任选地用作修饰植物基因组的修复模板的核酸分子)都是指能够编码这种GRF1多肽或GRF1蛋白质的核酸或多核苷酸。在一个实施方案中，对可用于本发明方法的多肽/蛋白质或核酸/多核苷酸的任何提及都应理解为是指可用于本发明方法、构建体、表达盒、植物细胞、植物、种子、可收获部分以及产物的蛋白质或核酸。待引入植物细胞或植物中(并且因此可用于执行本发明的方法)的包括mRNA的核酸/多核苷酸是编码现在将描述的多肽/蛋白质类型的任何核酸/多核苷酸，下文也称为“GRF1核酸”、“GRF1多核苷酸”、“GRF1基因”或“GRF1 mRNA”等。
如本文定义的“GRF1多肽”或“GRF1蛋白质”是指转录因子，优选14-3-3样蛋白质GF14-6，当用Interproscan软件(www.ebi.ac.uk/interpro)分析时更优选包含PFAM结构域PF08880(也称为QLQ结构域)以及PFAM结构域PF08879(也称为WRC结构域)，甚至更优选包含在GRF1多肽的N末端处或附近发现至少90％、至少91％、至少92％、至少93％、至少94％、至少95％、至少96％、至少97％、至少98％、至少99％或100％覆盖率的匹配的PFAM结构域PF08880以及在位于GRF1多肽的PFAM结构域PF08880的C末端处发现至少90％、至少91％、至少92％、至少93％、至少94％、至少95％、至少96％、至少97％、至少98％、至少99％或100％覆盖率的匹配的PFAM结构域PF08879，即，匹配PF08879的氨基酸段(stretch)位于匹配PF08880的氨基酸段后面的翻译方向上。优选地，这些匹配中的至少一者具有至少95％、至少96％、至少97％、至少98％、至少99％或100％的覆盖率。
在一个实施方案中，两个匹配的氨基酸段均位于GRF1多肽的N末端一半，优选地，匹配PFAM结构域PF08880的氨基酸段位于GRF1多肽的N末端四分之一。优选地，PFAM结构域PF08880匹配从GRF1多肽的残基5、6、7、8、9、10、11、12、13、14、15、16、17、18、19、20、21、22、23、24或25、优选从残基9、10、11、12、13、14、15、16、17、18、19、20或21开始的GRF1多肽的氨基酸残基，并且PFAM结构域PF08879匹配从GRF1多肽的残基68、69、70、71、72、73、74、75、76、77、78、79、80、81、82、83、84、85、86、87、88、89、90、91、92、93、94、95、96、97、98或99、优选从残基72、73、74、75、76、77、78、79、80、81、82、83、84、85、86、87、88、89、90、91、92、93、94或95开始的GRF1多肽的氨基酸残基。优选地，匹配PFAM结构域PF08880的氨基酸段的起始氨基酸残基与匹配PFAM结构域PF08879的氨基酸段的起始氨基酸残基之间的距离为GRF1多肽内的60至82个氨基酸，更优选为60至75个氨基酸，更优选为61至75个氨基酸，甚至更优选为GRF1多肽内的62至73个氨基酸。
在另一实施方案中，如本文所用的GRF1多肽包含指示基序：[D]-[P]-[E]-[P]-[G]-[R]-[C]-[R]-[R]-[T]-[D]-[G]-[K]-[K]-[W]-[R]-[C]-[AS]-[RK]-[ED]-[A]-[AH]-[PQS]-[D]-[S]-[K]-[Y]-[C]-[E]-[KR]-[H]-[M]-[H]-[R]-[G]-[R]-[N]-[R](SEQ ID NO:25)，其中可允许/可容忍的是，GRF1多肽与指示基序相比表现出最大数目的两个错配(即，在相应GRF1多肽与指示基序之间的序列对比中可出现至多两个错配)，优选地与指示基序相比表现出最大数目的一个错配(即，在相应GRF1多肽与指示基序之间的序列对比中可仅出现一个错配)，并且更优选地与指示基序相比没有错配。错配是指根据指示基序在某一位置的氨基酸被不同的氨基酸取代，或通过插入至少一个附加氨基酸而缺失或移位。方括号内的指示基序的字母表示氨基酸残基(单字母代码)，并且基序表示如本发明的任何GRF1多肽中存在的从N末端到C末端方向上的氨基酸残基的顺序。如果一个方括号内存在两个或三个字母，则它们表示备选。基序已经根据衍生自不同植物物种(包括单子叶以及双子叶植物)的GRF1多肽的序列的全面比较而推导出，并且允许将GRF1多肽与GRF蛋白质家族的其它成员(如GRF5)区分开。优选地，该基序位于GRF1多肽的N末端一半，更优选地，该基序位于GRF1多肽的N末端一半并包含匹配PFAM结构域PF08879的氨基酸段的亚区，即该基序具有与匹配PFAM结构域PF08879的氨基酸段的序列重叠。优选地，指示基序由氨基酸序列SEQ IDNO:26至SEQ ID NO:35中的任一者组成。对应地，GRF1多肽优选包含由氨基酸序列SEQ IDNO:26至SEQ ID NO:35中的任一者组成的一个连续基序。
根据一个方面，本发明提供了一种用于转化植物细胞的方法，该方法包括以下步骤：
(a1)向植物细胞中引入
(i)至少一个感兴趣的核苷酸序列；以及
(ii)包含编码GRF1多肽的多核苷酸的表达盒、编码GRF1多肽的mRNA或GRF1多肽，其中(i)与(ii)可以同时引入或以任何顺序依次引入，或
(a2)向植物细胞中引入至少一个感兴趣的核苷酸序列；并且在所述植物细胞中平行或依次诱导编码GRF1多肽的内源基因的增强的表达水平；以及
(b)任选地在以下条件下培育(a1)或(a2)的植物细胞或衍生自(a1)或(a2)的植物细胞的植物细胞，其中在所述植物细胞中，GRF1多肽从表达盒表达，GRF1多肽从引入的mRNA翻译，GRF1多肽从内源基因增强/增加表达，或优选地，与野生型植物细胞或其中尚未根据步骤(a1)引入包含编码GRF1多肽的多核苷酸的表达盒、编码GRF1多肽的mRNA或GRF1多肽或其中尚未根据步骤(a2)诱导编码GRF1多肽的内源基因的增强的表达水平的植物细胞中的量相比，GRF1多肽存在的量增加。
根据本发明的方法产生修饰的或转化的植物细胞，其由于存在GRF1或存在增加量的GRF1而具有改善的再生能力。然而，优选的是，在修饰的植物细胞中，GRF1的存在或增加量的GRF1的存在是瞬时的。
植物细胞的转化是指将核酸分子以导致稳定整合到植物细胞基因组中或瞬时出现在植物细胞中的方式引入植物细胞中，从而导致例如构成性地、暂时地或特异性地与特定组织或某些发育阶段等等相关的核酸序列的表达。单子叶及双子叶植物细胞的转化及再生现在都是常规的，并且最合适的转化技术的选择将由实践者决定。方法的选择将随待转化的植物的类型或基因型而变化；本领域的技术人员将认识到特定方法对于给定植物类型或基因型的适用性。合适的方法可以包括但不限于：植物原生质体的电穿孔；脂质体介导的转化；聚乙二醇(PEG)介导的转化；使用病毒的转化；植物细胞的微量注射；植物细胞的微射弹轰击；真空渗透；以及农杆菌介导的转化。
可以使用本领域公知的任何合适的方法执行引入所述至少一个感兴趣的核苷酸序列的步骤(a1)(i)或(a2)。许多方法可用于将目的核酸转移到植物细胞中。示例性的载体介导的方法是农杆菌介导的转化，如例如由Lindsay&；Gallois，1990，Journal ofexperimental botany以及Kischenko等，2005，Cell biology international for sugarbeet，Ishida等，2007，针对玉米的(“玉米的农杆菌介导的转化。(Agrobacterium-mediatedtransformation of maize.)”Nature protocols，2(7)，1614-1621)，或由来自日本烟草公司的针对小麦的PureWheat Technology所述。其它合适的技术包括粒子轰击以及电穿孔。
根据本发明的感兴趣的核苷酸序列可以是DNA或RNA序列，例如mRNA、siRNA、miRNA等。更具体而言，感兴趣的核苷酸序列编码至少一种表型性状。优选地，由DNA或RNA赋予的表型性状可以选自以下表型性状：对生物应激的抗性/耐受性，包括病原体抗性/耐受性，其中病原体可以是病毒、细菌、真菌或动物病原体；对非生物应激的抗性/耐受性，包括寒冷抗性/耐受性、干旱应激抗性/耐受性、渗透抗性/耐受性、热应激抗性/耐受性、寒冷或霜冻应激抗性/耐受性、氧化应激抗性/耐受性、重金属应激抗性/耐受性、盐应激或水淹抗性/耐受性、倒伏抗性/耐受性、落粒抗性/耐受性、或对一种或多种除草剂(如草甘膦、草丁膦、2,4-D、麦草畏、ALS抑制剂等)的抗性/耐受性。所述至少一种目的表型性状还可以选自以下表型性状：另一种目的农艺性状的修饰，包括产量增加、开花时间修饰、种子颜色修饰、胚乳组成修饰、营养含量修饰或目的途径的代谢工程。
在本发明的上下文中，GRF1可以作为包含编码GRF1多肽的多核苷酸的表达盒、作为编码GRF1多肽的mRNA(也包括前mRNA或前体mRNA)或作为GRF1多肽引入。上文描述了用于引入核酸分子的示例性技术。可替代地，可通过激活编码GRF1多肽的内源基因的表达而在植物细胞中提供GRF1。这将导致内源GRF1基因的表达水平增强，即导致GRF1多肽以增加的量存在或出现于植物细胞中。内源基因表达的激活可以通过修饰编码GRF1多肽的内源基因的启动子的活性或结构来实现。例如，可以通过基因编辑将增强子元件引入启动子中；或者可以进一步加强调节启动子的增强子元件，或者可以通过例如靶向诱变/修饰而弱化调节启动子的沉默子元件；或可以通过如CRISPR系统之类的基因编辑工具将修饰引入与增强子相关的表观基因组中(Hilton等(2015))。通过基于CRISPR-Cas9的乙酰基转移酶进行的表观基因组编辑激活来自启动子以及增强子的基因。Nature biotechnology，33(5)，510-517)；或基于例如TALE激活因子或dCas9激活因子的合成转录因子可以引入细胞中，其中它们能够结合启动子上或附近的靶向识别位点并激活GRF1基因的转录(Cheng等，(2013).通过RNA引导的转录激活因子系统CRISPR-on进行的内源基因的多重激活，Cell research，23(10)，1163.)；或植物细胞中通过转录后抑制调节GRF1基因表达的微小RNA(miRNA)的量可通过例如敲除(无效突变体)或敲低来降低，以便增加植物细胞中的翻译的GRF1多肽的量。
根据植物物种以及细胞类型，需要不同水平的基因或表达激活，以便当发生再生时在植物细胞中存在足够量的GRF1多肽。本领域的技术人员可利用各种技术来测量内源或引入的基因的实际表达水平，例如qPCR、RT-PCR、Northern印迹或微阵列。这些方法允许本领域的技术人员通过常规工作来调节GRF1基因的表达水平，其使来自不同组织或体细胞及生殖细胞(例如，小孢子)的再生能力得到改善。在优选的实施方案中，在植物细胞中，编码GRF1多肽的内源基因的表达水平增加至少2倍、3倍或5倍，优选10倍、25倍或50倍，更优选100倍、200倍或500倍。
如上文进一步描述，植物细胞中内源基因的增强的表达水平的诱导可以通过应用一种或多种激活因子或其前体来进行。这些可施加到其中培养植物细胞的培养基，然后被植物细胞主动或被动吸收。此外，所述一种或多种激活因子或其前体可以通过显微注射、电穿孔或基因枪轰击而直接导入植物细胞中。除了上述合成转录激活因子之外，根据本领域当前发展状况已知许多其它激活因子，它们可用于增加内源基因的表达水平，特别是内源GRF1基因的表达水平：近年来，化学植物遗传学以及化学植物生物学的技术领域出现，其中生物系统是利用小分子进行处理以特异性干扰细胞功能。小分子在商业上用作不同系统中的药物、除草剂以及杀真菌剂，但近年来它们也越来越多地用作遗传调节的工具。例如，化学遗传学涉及通过化合物库的筛选发现各种细胞功能的小分子效应物(Dejonghe&；Russinova(2017).植物化学遗传学：从基于表型的筛选到合成生物学(Plant ChemicalGenetics:From Phenotype-Based Screens to Synthetic Biology.)Plant Physiology，pp-01805；Kawasumi,M.,&；Nghiem,P.(2007).化学遗传学：用小分子化合物阐明生物系统(Chemical genetics:elucidating biological systems with small-moleculecompounds.)Journal of Investigative Dermatology，127(7)，1577-1584.)。适合于激活如GRF1之类的靶基因的表达的这种小分子效应物可以通过遵循不同策略的化学筛选来鉴定(Dejonghe&；Russinova，2017)。可获得综合化合物库，其允许简单筛选无数小分子并鉴定可用于激活如GRF1之类的基因的基因表达的效应物。如上所述，增强如GRF1之类的内源基因的表达水平的另一种方法是应用所谓的合成转录激活因子。它们通常通过识别结构域与至少一个激活因子结构域的融合来设计。识别结构域可以衍生自如锌指、TAL效应物或CRISPR之类的已知系统；对于激活，例如将单纯疱疹病毒衍生的VP-16或VP-64激活结构域与识别结构域融合可引起转录增加。诸如人NF-κB的AD之类的较弱的激活结构域增加了基因激活的多种选择。此外，如内源启动子所示，激活因子的组合可用于引入协同作用(Moore等(2014).“转录激活因子样效应物：合成生物学工具包。(Transcription activator-likeeffectors:a toolkit for synthetic biology.)”ACS synthetic biology，3(10)，708-716.；US 2002/0046419 A1；Lowder等(2017).“使用基于CRISPR-dCas9的系统在植物中进行多重转录激活或抑制。(Multiplexed transcriptional activation or repression inplants using CRISPR-dCas9-based systems.)”Plant Gene Regulatory Networks:Methods and Protocols，167-184.)。合成转录激活因子也可以作为前体，即作为编码这种人工或合成转录激活因子或其结构域的DNA或RNA分子，或作为稍后在细胞中或在细胞的特定区室中激活的转录激活因子的无活性形式，递送到植物细胞或引入植物细胞中。最后，增强GRF基因的表达还可以通过上游负调节因子的失活或通过产生抗这种负调节因子的GRF1基因的突变形式来实现。
优选地，本发明的GRF1多肽包含选自SEQ ID NO:2、37、39、41、43、45、47、49、51和53的氨基酸序列，或与选自SEQ ID NO:2、37、39、41、43、45、47、49、51和53的序列具有至少70％同一性的氨基酸序列，优选地与选自SEQ ID NO:2、37、39、41、43、45、47、49、51和53的序列具有至少80％、至少85％、至少90％、更优选地至少95％、至少98％或至少99％同一性的氨基酸序列。
编码本发明的GRF1多肽的外源(异源)或内源多核苷酸包含：
(i)选自SEQ ID NO:1、36、38、40、42、44、46、48、50和52的编码核苷酸序列；
(ii)包含如下序列的编码核苷酸序列，所述序列与选自SEQ ID NO:1、36、38、40、42、44、46、48、50和52的核苷酸序列至少70％、优选至少80％、至少85％、至少90％、更优选至少95％、至少98％或至少99％相同；
(iii)编码如上定义的GRF1多肽的核苷酸序列或在遗传密码的简并范围内编码由(i)和/或(ii)编码的多肽的核苷酸序列；
(iv)与(i)、(ii)或(iii)的核苷酸序列互补的核苷酸序列；或/及
(v)在严格条件下与(iv)的核苷酸序列杂交的核苷酸序列。
优选地，编码GRF1多肽的多核苷酸，特别是编码GRF1多肽的多核苷酸，可以包含SEQ ID NO:1的编码核苷酸序列。
为了本发明的目的，以百分比表示的两个相关核苷酸或氨基酸序列的“序列同一性”是指这两个最佳比对的序列中具有相同残基(×100)的位置数除以比较的位置数。缺口，即在比对中其中一个序列中存在残基而另一个序列中不存在残基的位置，被认为是具有不同残基的位置。通过Needleman以及Wunsch算法(Needleman以及Wunsch1970)进行这两种序列的比对。上述计算机辅助序列比对可以使用标准软件程序方便地进行，诸如在欧洲分子生物学开放软件套装(EMBOSS)中实施的程序NEEDLE，例如6.3.1.2版(Trends inGenetics 16(6)，276(2000))，其默认参数例如对于蛋白质基质＝EBLOSUM62、gapopen＝10.0以及gapextend＝0.5。
术语“严格条件”或“在严格条件下杂交”是指这样的条件，在所述条件下彼此具有足够互补性的核苷酸序列通常保持杂交。这些严格条件是本领域的技术人员已知的，并且描述于例如Current Protocols in Molecular Biology，John Wiley&；Sons，纽约(1989)6.3.1-6.3.6。基于例如Sambrook等所著的Molecular Cloning(冷泉港实验室，1989)，技术人员知道如何确定所需的杂交条件。在这方面，术语“杂交条件”不仅指在核酸的实际聚集期间的实际条件，而且指在随后洗涤步骤中的条件。严格杂交条件的示例是这样的条件，在所述条件下，主要是仅那些具有至少80％、优选至少85％、至少90％、至少95％、至少98％或至少99％序列同一性的核酸分子进行杂交。严格杂交条件是例如：在65℃下4×SSC，并且随后在65℃下在0.1×SSC中多次洗涤约1小时。如本文所用的术语“严格杂交条件”还可以指：在68℃下在0.25M磷酸钠，pH 7.2，7％SDS，1mM EDTA以及1％BSA中杂交16小时，并且随后在68℃下用2×SSC以及0.1％SDS洗涤两次。优选地，在严格条件下进行杂交。
表述“可操作地连接”是指嵌合基因的所述因子以这样的方式彼此连接，即它们的功能是协调的并且允许编码序列的表达，即它们是功能性连接的。举例而言，当启动子能够确保另一核苷酸序列的转录以及最终表达时，启动子功能性连接到所述另一核苷酸序列。如果编码核苷酸序列的两个蛋白质以可形成第一与第二蛋白质或多肽的融合蛋白质的方式连接，则这两个蛋白质彼此功能性地或可操作地连接。
当基因导致表达产物形成时，称该基因被表达。表达产物表示由编码这种产物的核酸、DNA或RNA(例如，本文所述的第二核酸)的转录及任选的翻译产生的中间产物或终产物。在转录过程中，受调节区、特别是启动子控制的DNA序列被转录成RNA分子。当RNA分子能够被翻译成肽或蛋白质时，其自身可以形成表达产物或中间产物。当作为基因表达终产物的RNA例如能够与另一种核酸或蛋白质相互作用时，该基因被称为编码作为表达产物的RNA分子。RNA表达产物的示例包括抑制性RNA，诸如例如正义RNA(共抑制)、反义RNA、核酶、miRNA或siRNA、mRNA、rRNA以及tRNA。当基因表达的终产物是蛋白质或肽时，该基因被称为编码作为表达产物的蛋白质。
如本文所用的核酸(分子)或核苷酸(序列)或多核苷酸是指DNA及RNA二者。DNA还包括cDNA以及基因组DNA。核酸分子可以是单链或双链的，并且可以在体外或甚至在体内通过化学合成或通过生物表达产生。
显然，只要RNA分子的核苷酸序列是参照对应DNA分子的核苷酸序列定义的，核苷酸序列中的胸腺嘧啶(T)应被尿嘧啶(U)取代。根据本申请的上下文，参考RNA还是DNA分子将显而易见。
如本文所用，“包含”等应解释为指定所提及的所述特征、整数、步骤或组分的存在，但不排除一个或多个特征、整数、步骤或组分或其组合的存在或添加。因此，例如包含核苷酸或氨基酸序列的核酸或蛋白质可包含比实际引用的核苷酸或氨基酸更多的核苷酸或氨基酸，即嵌入较大的核酸或蛋白质中。包含功能上或结构上限定的DNA区的嵌合基因可以包含附加的DNA区等。
通过GRF1，可以提供具有改善的再生能力的植物细胞或其它植物前体组织。这对于遗传修饰的植物细胞、特别是具有编辑过的基因组的植物细胞特别有用。
根据本发明的优选实施方案，引入所述至少一个感兴趣的核苷酸序列以及GRF1的步骤(a1)或引入所述至少一个感兴趣的核苷酸序列并诱导编码GRF1的内源基因的增强的表达水平的步骤(a2)在植物细胞的瞬时转化中产生。就本发明而言，“瞬时转化”是指插入序列未(稳定地)整合到植物细胞的基因组中。在另一个实施方案中，实现了稳定的转化，其中本文公开的转化方法的步骤(a1)及(a2)中的感兴趣的核苷酸序列和/或本文公开的修饰基因组的方法的步骤(a1)中的编码GRF1多肽的多核苷酸被插入到植物细胞的基因组中。根据本发明的特别优选的实施方案，编码GRF1的核苷酸序列被瞬时转化到细胞中，而感兴趣的核苷酸序列被稳定地转化到细胞的基因组中。
修饰植物细胞的基因组可通过单链DNA断裂(SSB)或双链DNA断裂(DSB)诱导酶或碱基编辑酶实现，所述单链DNA断裂或双链DNA断裂诱导酶或所述碱基编辑酶优选识别所述细胞的基因组中的预定位点。
因此，本发明的另一个实施方案是一种用于修饰植物细胞的基因组的方法，该方法包括以下步骤：
(a1)向植物细胞中引入包含编码GRF1多肽的多核苷酸的表达盒、编码GRF1多肽的mRNA(包括前mRNA)或GRF1多肽；或
(a2)在植物细胞中诱导编码GRF1多肽的内源基因的增强的表达水平；以及
(b)在以下条件下培育(a1)或(a2)的植物细胞或衍生自(a1)或(a2)的植物细胞的植物细胞，其中在植物细胞中，GRF1多肽从表达盒表达，GRF1多肽从引入的mRNA翻译，GRF1多肽从内源基因增强/增加表达，或优选地，与野生型植物细胞或其中尚未根据步骤(a1)引入包含编码GRF1多肽的多核苷酸的表达盒、编码GRF1多肽的mRNA或GRF1多肽或其中尚未根据步骤(a2)诱导编码GRF1多肽的内源基因的增强的表达水平的植物细胞中的量相比，GRF1多肽存在的量增加；
(c)通过单链DNA断裂(SSB)或双链DNA断裂(DSB)诱导酶或碱基编辑酶以及任选地通过修复核酸分子修饰(b)的植物细胞的基因组，所述单链DNA断裂或双链DNA断裂诱导酶或所述碱基编辑酶优选识别所述细胞的基因组中的预定位点，
其中所述基因组在所述预定位点处的修饰选自
i.至少一个核苷酸的取代；
ii.至少一个核苷酸的缺失；
iii.至少一个核苷酸的插入；或
iv.i.-iii.的任何组合；并且
其中步骤(c)与步骤(a1)/(a2)和/或(b)同时进行，在步骤(a1)/(a2)之前，在步骤(a1)/(a2)与(b)之间或在步骤(b)之后进行。
如本文所用，“双链DNA断裂诱导酶”或“DSBI酶”是能够在特定核苷酸序列(称为“识别位点”)处诱导双链DNA断裂的酶。因此，“单链DNA或RNA断裂诱导酶”或“SSBI酶”是能够在特定核苷酸序列处诱导单链DNA或RNA断裂的酶。
为了能够在预定靶位点断裂，酶优选包括结合结构域以及切割结构域。能够诱导双或单链断裂的特定酶是核酸酶及其变体，不再包含核酸酶功能，而是作为与另一种酶组合的识别分子起作用。近年来，已经开发出了许多合适的核酸酶，特别是特制的核酸内切酶，包括大范围核酸酶、锌指核酸酶、TALE核酸酶、Argonaute核酸酶(例如，衍生自格氏嗜盐碱杆菌)以及CRISPR核酸酶，其包括例如Cas、Cpf1、CasX或CasY核酸酶作为成簇的规则间隔的短回文重复(CRISPR)系统的一部分。因此，在本发明的优选方面，DSB或SSB诱导酶选自CRISPR系统，如CRISPR/Cas9、CRISPR/Cpf1、CRISPR/CasX、CRISPR/CasY、CRISPR/Csm1或CRISPR/MAD7a，优选CRISPR/Cas9内切核酸酶或CRISPR/Cpf1内切核酸酶、锌指核酸酶(ZFN)、归巢内切核酸酶、大范围核酸酶以及TAL效应物核酸酶。
罕见切割核酸内切酶是具有优选约14至70个连续核苷酸的识别位点的酶，因此即使在诸如大多数植物基因组之类的较大基因组中，也具有非常低的切割频率。归巢核酸内切酶，也称为大范围核酸酶，构成这种罕见切割核酸内切酶的家族。它们可以由内含子、独立的基因或间插序列编码，并呈现显著的结构及功能特性，这些特性使它们区别于更经典的限制性酶，通常区别于细菌限制性修饰II型系统。它们的识别位点具有一般不对称性，其与大多数限制性酶识别位点的特征性二分对称性截然不同。已经显示由内含子或内含肽编码的几种归巢核酸内切酶促进它们各自的遗传因子归巢到等位基因的无内含子或无内含肽位点中。通过在无内含子或无内含肽等位基因中产生位点特异性双链断裂，这些核酸酶产生重组基因末端，其参与基因转化过程，该基因转化过程复制编码序列并导致在DNA水平上插入内含子或间插序列。在WO 03/004659的表I(第17至20页)中提供了其它罕见切割大范围核酸酶及其各自的识别位点的列表(通过引用并入本文)。
此外，可利用设计定制的罕见切割核酸内切酶的方法，这些核酸内切酶基本上识别任何精选的靶核苷酸序列。简言之，可使用被设计成识别特定核苷酸序列的锌指结构域与来自诸如Fok1之类的天然限制性酶的非特异性DNA切割结构域之间的杂交体来制备嵌合限制性酶。这些方法描述于例如WO 03/080809、WO 94/18313或WO 95/09233中，以及Isalan等(2001).一种通过靶向HIV-1启动子示出的用于设计锌指的快速、普遍适用的方法(Arapid,generally applicable method to engineer zinc fingers illustrated bytargeting the HIV-1promoter.)Nature biotechnology，19(7)，656；Liu等(1997).用于在复杂基因组内唯一定位的多指锌指蛋白的设计(Design of polydactyl zinc-fingerproteins for unique addressing within complex genomes.)Proceedings of theNational Academy of Sciences，94(11)，5525-5530.)。
定制设计的核酸内切酶的另一个示例包括TALE核酸酶(TALEN)，其是基于与核酸酶(例如FokI或其变体)的催化结构域融合的来自细菌属黄单胞菌属的转录激活因子样效应物(TALE)。这些TALE的DNA结合特异性是由串联排列的34/35个氨基酸重复单元的重复可变双残基(RVD)定义，使得一个RVD特异性识别靶DNA中的一个核苷酸。重复单元可以进行装配以基本上识别任何靶序列，并与核酸酶的催化结构域融合，产生序列特异性核酸内切酶(参见例如Boch等(2009).破坏TAL III型效应物的DNA结合特异性的编码(Breaking thecode of DNA binding specificity of TAL-type III effectors.)Science，326(5959)，1509-1512；Moscou&；Bogdanove(2009).A simple cipher governs DNA recognitionby TAL effectors.Science，326(5959)，1501-1501；以及WO 2010/079430、WO2011/072246、WO 2011/154393、WO 2011/146121、WO 2012/001527、WO 2012/093833、WO 2012/104729、WO 2012/138927、WO 2012/138939)。WO 2012/138927进一步描述了具有各种催化结构域的单体(紧密)TALEN及TALE以及其组合。
最近，描述了一种新型可定制的核酸内切酶系统；所谓的CRISPR/Cas系统。在其自然环境中的CRISPR系统描述了一种分子复合物，该分子复合物包含与Cas核酸酶或另一种CRISPR核酸酶(如Cpf1核酸酶)组合的至少一种小的且单独的非编码RNA(Zetsche等，“Cpf1Is a Single RNA-Guides Endonuclease of a Class 2CRISPR-Cas System”，Cell，163，第1-13页，2015年10月)，该分子复合物可以产生特异性DNA双链断裂。目前，CRISPR系统被分为2类，其包括五种类型的CRISPR系统，II型系统例如使用Cas9作为效应子并且V型系统使用Cpf1作为效应分子(Makarova等，Nature Rev.Microbiol.，2015)。在人工CRISPR系统中，合成的非编码RNA及CRISPR核酸酶和/或任选地修饰的CRISPR核酸酶(经修饰以充当切口酶或缺乏任何核酸酶功能)可以与结合crRNA和/或tracrRNA的功能的至少一种合成的或人工的指导RNA或gRNA组合使用(Makarova等，2015，同上)。在天然系统中由CRISPR/Cas介导的免疫应答需要CRISPR-RNA(crRNA)，其中控制CRISPR核酸酶的特异性激活的这种指导RNA的成熟在迄今已经表征的各种CRISPR系统之间显著变化。首先，将侵入的DNA(也称为间隔区)整合在CRISPR基因座近端的两个相邻重复区之间。II型CRISPR系统编码Cas9核酸酶作为干扰步骤的关键酶，该系统含有crRNA以及反式激活RNA(tracrRNA)两者作为引导基序。这些杂交并形成双链(ds)RNA区，其由RNAseIII识别并可被切割以形成成熟的crRNA。然后这些又与Cas分子结合以将核酸酶特异性地导向靶核酸区。重组gRNA分子可以包含可变DNA识别区以及Cas相互作用区两者，并且因此可以被特别设计，而与特定靶核酸以及所希望的Cas核酸酶无关。作为进一步的安全机制，PAM(前间区序列相邻基序)必须存在于靶核酸区中；这些是直接来自Cas9/RNA复合物识别的DNA的DNA序列。来自酿脓链球菌的Cas9的PAM序列已经被描述为“NGG”或“NAG”(标准IUPAC核苷酸代码)(Jinek等，“适应性细菌免疫中的可编程的双RNA引导的DNA核酸内切酶(A programmable dual-RNA-guided DNAendonuclease in adaptive bacterial immunity)”，Science 2012，337：816-821)。来自金黄色酿脓葡萄球菌的Cas9的PAM序列是“NNGRRT”或“NNGRR(N)”。进一步的变体CRISPR/Cas9系统是已知的。因此，脑膜炎奈瑟氏菌Cas9在PAM序列NNNNGATT处切割。嗜热链球菌Cas9在PAM序列NNAGAAW处切割。最近，已经描述了用于弯曲杆菌的CRISPR系统的另外的PAM基序NNNNRYAC(WO 2016/021973 A1)。对于Cpf1核酸酶，已经描述了与通过Cas9系统识别的通常富含G的PAM相反，没有tracrRNA的Cpf1-crRNA复合物有效地识别并切割通过短的富含T的PAM进行的靶DNA(Zetsche等，同上)。此外，通过使用经修饰的CRISPR多肽，可以获得特异性单链断裂。Cas切口酶与各种重组gRNA的组合使用还可以通过双DNA切口形成诱导高度特异性的DNA双链断裂。此外，通过使用两种gRNA，可以优化DNA结合的特异性，从而优化DNA切割。同时，最初针对细菌所描述的其它CRISPR效应物(如CasX及CasY效应物)是可获得的，并且代表可以用于基因组工程目的的其它效应物(Burstein等，“来自野生微生物的新CRISPR-Cas系统(New CRISPR-Cas systems from uncultivated microbes)”，Nature，2017，542，237-241)。
DSBI/SSBI酶的切割位点涉及DNA或RNA上诱导双链断裂的确切位置。切割位点可以包含或不包含在DSBI/SSBI酶的识别位点中(与其重叠)，因此据说DSBI/SSBI酶的切割位点位于其识别位点处或附近。DSBI/SSBI酶的识别位点，有时也称为结合位点，是由DSBI/SSBI酶(特异性地)识别并确定其结合特异性的核苷酸序列。例如，TALEN或ZNF单体具有分别由其RVD重复或ZF重复确定的识别位点，而其切割位点是由其核酸酶结构域(例如，FokI)确定并且通常位于识别位点之外。在二聚体TALEN或ZFN的情况下，切割位点位于相应单体的所述两个识别/结合位点之间，发生切割的该间插DNA或RNA区域称为间隔区。
本领域的技术人员能够或者选择识别某一识别位点并在预选/预定位点处或附近的切割位点诱导DSB或SSB的DSBI/SSBI酶，或者设计这种DSBI/SSBI酶。可替代地，可以使用任何常规转化方法或通过与基因组中具有DSBI/SSBI酶识别位点的生物体杂交来将DSBI/SSBI酶识别位点引入靶基因组中，然后可在该DSBI/SSBI酶的切割位点处或附近引入任何所需核酸。
如本文所用的“碱基编辑酶”或“碱基编辑器”是指与其来源的蛋白质具有相同催化活性的蛋白质或其片段，如果碱基转化不会引起沉默突变而是由包含用碱基编辑器转化的位置的密码子编码的氨基酸的转化，则该蛋白质或其片段单独或当作为分子复合物(在本文中称为碱基编辑复合物)提供时具有介导靶向碱基修饰(即，导致目的点突变的目的碱基的转化)的能力，所述目的点突变又可导致靶向突变。优选地，根据本发明的至少一个碱基编辑器暂时或永久地连接到至少一个位点特异性效应物，或任选地连接到至少一个位点特异性效应物复合物的组分。连接可以是共价的和/或非共价的。
如本文公开的任何碱基编辑器或位点特异性效应物、或其催化活性片段、或碱基编辑器复合物或位点特异性效应物复合物的任何组分均可以作为核酸片段引入细胞中，所述核酸片段代表或编码DNA、RNA或蛋白质效应物，或其可以作为DNA、RNA和/或蛋白质、或其任何组合引入。
存在两种主要且不同的修复断裂的途径－同源重组以及非同源末端连接(NHEJ)。同源重组需要存在同源序列作为模板(例如，“供体”)以指导细胞修复过程，并且修复的结果是无误差的且可预测的。在不存在用于同源重组的模板(或“供体”)序列的情况下，细胞通常试图通过非同源末端连接(NHEJ)的过程来修复断裂。
在该实施方案的特别优选的方面，将修复核酸分子附加地引入植物细胞中。如本文所用，“修复核酸分子”是单链或双链DNA分子或RNA分子，其用作在切割位点附近的预选位点处或在切割位点处修饰基因组DNA的模板。如本文所用，“用作修饰基因组DNA的模板”是指修复核酸分子通过侧翼区与靶基因组中位于预选位点侧翼的对应同源区之间的同源重组在预选位点处复制或整合，任选地在修复核酸分子的两个末端之一处与非同源末端连接(NHEJ)组合(例如，在仅存在一个侧翼区的情况下)。通过同源重组进行的整合将允许修复核酸分子精确连接到靶基因组直至核苷酸水平，而NHEJ可在修复核酸分子与基因组DNA之间的连接处产生小的插入/缺失。
如本文所用，“基因组的修饰”是指基因组已通过至少一个核苷酸改变。这可以通过至少一个核苷酸的取代和/或至少一个核苷酸的缺失和/或至少一个核苷酸的插入而发生，只要与修饰前的预选基因组靶位点的核苷酸序列相比其导致至少一个核苷酸的总改变，从而允许例如通过技术人员熟知的技术(诸如测序或PCR分析等)鉴定修饰。
如本文所用，“预选位点”、“预定位点”或“预定义位点”表示基因组(例如，核基因组或叶绿体基因组)中的期望插入、取代和/或缺失一个或多个核苷酸的特定核苷酸序列。这可以是例如内源基因座、或位于先前导入的外源DNA或转基因中的或连接至先前导入的外源DNA或转基因的特定核苷酸序列。预选位点可以是特定的核苷酸位置，在该位置处(之后)意图进行一个或多个核苷酸的插入。预选位点还可包括欲被交换(取代)或缺失的一个或多个核苷酸的序列。
如在本申请的上下文中所用，术语“约”是指所述值的+/-10％，优选所述值的+/-5％。例如，约100个核苷酸(nt)应理解为90与110nt之间、优选95与105之间的值。
如本文所用，“侧翼区”是具有与预选位点侧翼(即，上游或下游)的DNA区的核苷酸序列同源的核苷酸序列的修复核酸分子的区域。显然，应选择侧翼区的长度及百分比序列同一性，以便能够在所述侧翼区与它们的位于预选位点上游或下游的对应DNA区之间进行同源重组。与修复核酸分子的侧翼DNA区具有同源性的位于预选位点侧翼的DNA区也称为基因组DNA中的同源区。
为了具有用于重组的足够同源性，修复核酸分子的侧翼DNA区域的长度可以变化，并且长度应该为至少约10nt、约15nt、约20nt、约25nt、约30nt、约40nt或约50nt。然而，侧翼区可以尽可能长(例如，高达约100-150kb，诸如完整的细菌人工染色体(BAC))。优选地，侧翼区为约50nt至约2000nt，例如约100nt、200nt、500nt或1000nt。此外，目的DNA侧翼的区域不需要与同源区(预选位点侧翼的DNA区)相同，并且可以与预选位点侧翼的DNA区具有约80％至约100％的序列同一性、优选约95％至约100％的序列同一性。侧翼区越长，对同源性的要求就越不严格。此外，为了在预选位点处交换靶DNA序列而不改变相邻DNA序列的DNA序列，侧翼DNA序列优选应与预选位点侧翼的上游及下游DNA区相同。
如本文所用，“上游”表示核酸分子上更接近所述核酸分子的5”末端的位置。同样，术语“下游”是指核酸分子上更接近所述核酸分子的3"末端的位置。为避免疑问，核酸分子及其序列通常以其5”至3”方向(从左到右)表示。
为了在预选位点处靶向序列修饰，必须选择侧翼区，使得上游侧翼区的3"末端和/或下游侧翼区的5"末端与预定义位点的末端对齐。因此，上游侧翼区的3"末端确定预定义位点的5"末端，而下游侧翼区的5"末端确定预定义位点的3"末端。
如本文所用，所述预选位点位于所述切割(和/或识别)位点外部或远离所述切割(和/或识别)位点是指意图进行基因组修饰的位点(预选位点)不包含DSBI/SSBI酶或碱基编辑酶的切割位点和/或识别位点，即预选位点不与切割(和/或识别)位点重叠。因此，在这方面外部/远离是指切割(和/或识别)位点的上游或下游。
已经根据本发明的方法转化或基因编辑并且可能具有经修饰的基因组的经修饰的植物细胞可以再生为完整(可育)植物。由于在植物细胞中存在附加的GRF1，它们的再生能力显著提高。因此，在本发明的优选方面，植物细胞的转化或植物细胞的基因组的修饰之后分别是再生植物的步骤。因此，本发明提供了一种用于产生转基因植物的方法，该方法包括：
(a)如上文所述转化植物细胞，以及
(b)从(a)的植物细胞或从衍生自(a)的植物细胞的植物细胞再生包含至少一个植物细胞的植物，所述至少一个植物细胞包含作为转基因的至少一个感兴趣的核苷酸序列。
进一步，本发明还提供了一种产生遗传修饰的植物的方法，该方法包括：
(a)如上文所述修饰植物细胞的基因组，以及
(b)从(a)的植物细胞或从衍生自(a)的植物细胞的植物细胞再生在至少一个细胞中包含基因组的修饰或经修饰的植物细胞的植物。
如本文所用，“再生”是指其中单个或多个细胞增殖并发育成组织、器官以及最终整个植物的过程。
再生植物的步骤b)可以例如包括在再生培养基上培养来自步骤a)的转化的或遗传修饰的植物细胞。
再生技术依赖于组织培养生长培养基中的某些植物激素的操作，偶尔依赖于可与所需感兴趣的核苷酸序列一起引入的生物杀灭剂和/或除草剂标记。来自培养的原生质体的植物再生描述于：Evans等，Protoplasts Isolation and Culture，Handbook of PlantCell Culture，第124-176页，MacMillilan出版公司，纽约，1983；以及Binding，Regeneration of Plants，Plant Protoplasts，第21-73页，CRC出版社，波卡拉顿，1985。还可以从植物愈伤组织、外植体、原生质体、未成熟或成熟胚、胚组织、分生组织、器官或其部分获得再生。这种再生技术大体上描述于Klee(1987)Ann.Rev.of Plant Phys.38:467486。为了从诸如未成熟胚之类的转基因组织获得完整植物，可以使它们在包含营养物以及激素的一系列培养基中在受控的环境条件下生长，该过程称为组织培养。一旦产生完整植物并产生种子，就开始评估后代。
根据本发明，不仅可以提高转化的或遗传修饰的植物细胞的再生能力，而且可以改善再生能力差的其它类型的敏感细胞。特别地，可以通过GRF1提高从如未成熟的雄性配子体或小孢子之类的前体产生单倍体植物胚。
因此，本发明的另一个方面是一种产生单倍体植物胚的方法，该方法包括以下步骤：
(a1)将包含编码GRF1多肽的多核苷酸的表达盒、编码GRF1多肽的mRNA或GRF1多肽引入未成熟的雄性配子体或小孢子中；或
(a2)在未成熟的雄性配子体或小孢子中诱导编码GRF1多肽的内源基因的增强的表达水平；以及
(b)在以下条件下培育(a)的未成熟的雄性配子体或小孢子，其中在未成熟的雄性配子体或小孢子中，GRF1多肽从表达盒表达，GRF1多肽从引入的mRNA翻译，GRF1多肽从内源基因增强表达，或优选地，与野生型植物细胞或其中尚未根据步骤(a1)引入包含编码GRF1多肽的多核苷酸的表达盒、编码GRF1多肽的mRNA或GRF1多肽或其中尚未根据步骤(a2)诱导编码GRF1多肽的内源基因的增强的表达水平的植物细胞中的量相比，GRF1多肽存在的量增加；以及
(c)选择衍生自步骤(b)的未成熟的雄性配子体或小孢子的单倍体植物胚。
本发明还包括一种产生单倍体幼苗的方法，该方法包括：将单倍体植物材料暴露于包含编码GRF1多肽的多核苷酸的表达盒、编码GRF1多肽的mRNA或GRF1多肽以产生单倍体胚；以及随后将单倍体胚转化(即，萌发)成幼苗。因此，本发明包括一种制备单倍体植物的方法，该方法包括使根据上述方法产生的幼苗生长。本发明还提供了一种产生双单倍体植物的方法，该方法包括：在GRF1存在下培养单倍体植物材料一段时间；刺激或允许自发的染色体加倍；以及使双单倍体植物材料生长成幼苗、小植物或植物。在某些实施方案中，单倍体胚发生以及染色体加倍可以基本上同时发生。在其它实施方案中，在单倍体胚发生与染色体加倍之间可能存在时间延迟。如果单倍体幼苗、植物或小植物的生长不涉及自发的染色体加倍事件，则可以使用化学染色体加倍剂，例如秋水仙碱。许多程序包括使植物细胞与秋水仙碱、抗微管剂或抗微管除草剂(诸如拿草特)、一氧化二氮或任何有丝分裂抑制剂接触。结果是纯合的双单倍体细胞。当使用秋水仙碱时，培养基中的浓度通常可以是0.01％-0.2％。秋水仙碱浓度的范围可以是约400-600mg/L。
当小孢子暴露于GRF1多肽时，则可形成愈伤组织并且这可经历器官发生以形成胚。因此，本发明包括一种产生单倍体植物愈伤组织的方法，该方法包括：将未成熟的雄性配子体或小孢子暴露于GRF1多肽或向其中引入包含编码GRF1多肽的多核苷酸的表达盒、编码GRF1多肽的mRNA或GRF1多肽。
在培养步骤期间将未成熟的雄性配子体或小孢子暴露于GRF1优选进行足以诱导单倍体胚形成的一段时间。所需的时间段可以取决于所涉及的植物的物种，并且这些都是本领域的普通技术人员容易确定的。GRF1暴露的优选范围为约1至约20小时，更优选约2至约20小时。
在本发明的某些方面，在引入包含编码GRF1多肽的多核苷酸的表达盒、编码GRF1多肽的mRNA或GRF1多肽之前，对单倍体植物材料施加物理应激。物理应激可以是例如温度、黑暗、光或电离辐射、饥饿或渗透应激中的任一种。光可以是全光谱太阳光，或选自可见、红外或UV光谱的一个或多个频率。应激可以是连续的或间断的(周期性的)，随着时间的过去有规律的或随机的。
在本发明的另一方面，发现可以通过组合GRF1与WUS2来进一步提高植物材料的转化及再生能力。因此，除了在用于转化植物细胞的方法、用于修饰植物细胞的基因组的方法、或用于产生用于步骤(a1)的根据本发明的单倍体植物胚的方法中如上所述引入包含编码GRF1多肽的多核苷酸的表达盒、编码GRF1多肽的mRNA或GRF1多肽、或者在用于转化植物细胞的方法、用于修饰植物细胞的基因组的方法、或产生用于步骤(a2)的根据本发明的单倍体植物胚的方法中如上所述诱导编码GRF1多肽的内源基因的增强的表达水平之外，根据该方面的本发明的方法进一步提供将包含编码WUS2多肽的多核苷酸的表达盒、编码WUS2多肽的mRNA或WUS2多肽引入植物细胞中。如果在步骤(a1)中，将包含编码GRF1多肽的多核苷酸的表达盒引入细胞中，则编码WUS2多肽的多核苷酸可以整合到相同的表达盒中并可操作地连接到相同的调节因子。
已经证明该方法在植物细胞的转化及植物细胞的修饰以及在如上所述产生转基因植物、遗传修饰的植物或单倍体植物胚的方法中都是有利的。
优选地，本发明的WUS2多肽包含选自SEQ ID NO:8、10、12、14、16、18、20、22以及24的氨基酸序列，或与SEQ ID NO:8、10、12、14、16、18、20、22以及24具有至少70％同一性、优选地与SEQ ID NO:8、10、12、14、16、18、20、22以及24具有至少80％、至少85％、至少90％、更优选地至少95％、至少98％或至少99％同一性的氨基酸序列。
编码本发明的WUS2多肽的多核苷酸优选地包含：
(i)选自SEQ ID NO:7、9、11、13、15、17、19、21以及23的编码核苷酸序列；
(ii)包含如下序列的编码核苷酸序列，所述序列与选自SEQ ID NO:7、9、11、13、15、17、19、21以及23的编码核苷酸序列至少70％、优选至少80％、至少85％、至少90％、更优选至少95％、至少98％或至少99％相同；
(iii)在遗传密码的简并范围内编码由(i)或(ii)编码的WUS2多肽的核苷酸序列；
(iv)与(i)、(ii)或(iii)的核苷酸序列互补的核苷酸序列；或/及
(v)在严格条件下与(iv)的核苷酸序列杂交的核苷酸序列。
本发明适用于任何植物物种，无论是单子叶植物还是双子叶植物。优选地，可用于本发明的方法及用途的植物是选自以下的属的植物：大麦属、高粱属、甘蔗属、玉蜀黍属、狗尾草属、水稻属、小麦属、黑麦属、黑小麦属、苹果属、短柄草属、山羊草属、胡萝卜属、甜菜属、桉树属、烟草属、茄属、咖啡属、葡萄属、沟酸浆属(Erythrante)、螺旋狸藻属、黄瓜属、桑属、拟南芥属、须弥芥属、碎米荠属、独行菜属、荠属、无苞芥属、筷子芥属、芸苔属、芝麻菜属、萝卜属、柑橘属、麻风树属、杨属、苜蓿属、鹰嘴豆属、木豆属、菜豆属、大豆属、棉属、黄芪属、莲属、蝴蝶草属、葱属或向日葵属。更优选地，植物选自以下组成的组：大麦、球茎大麦、两色高粱、甘蔗、包括玉蜀黍在内的玉蜀黍属物种、小米、小粒稻、水稻、澳洲野生稻、高秆野生稻、小麦、硬粒小麦、黑麦、黑小麦、苹果、紫短柄草、海滨大麦、节节麦、澳大利亚胡萝卜(Daucus glochidiatus)、包括甜菜在内的甜菜属物种、小胡萝卜、胡萝卜(Daucusmuricatus)、野胡萝卜、巨桉、美花烟草、绒毛状烟草、烟草、本氏烟草、番茄、马铃薯、中果咖啡、葡萄、沟酸浆(Erythrante guttata)、螺旋狸藻、黄瓜、川桑、沙生拟南芥、深山南芥、拟南芥、喜马拉雅鼠耳芥、卵叶须弥芥、弯曲碎米荠、北美独行菜、荠菜、无苞芥、筷子芥、欧洲油菜、甘蓝、芜菁、萝卜、芥菜、黑芥、芝麻菜、甜橙、麻风树、毛果杨、蒺藜状苜蓿、山下鹰嘴豆、野生鹰嘴豆、鹰嘴豆、网状鹰嘴豆、鹰嘴豆(Cicer judaicum)、木豆、蔓草虫豆、菜豆、大豆、棉花、紫云英、百脉根、夏堇、洋葱、葱、蒜、向日葵、菊芋和/或韭菜。特别优选的是甜菜、玉蜀黍、小麦、大麦、黑麦、向日葵、马铃薯、两色高粱、芜菁、欧洲油菜、芥菜、甘蓝、萝卜、水稻、大豆和/或棉花。
根据本发明的合适植物细胞尤其是体细胞组织、愈伤组织、分生组织或胚组织、或原生质体、配子体、花粉、胚珠或小孢子的细胞。例如，可以使用愈伤组织、优选易碎愈伤组织、分生组织、生殖组织(例如，小孢子)或胚组织以及原生质体的细胞。
植物的一个或多个部分可以附着于完整植物或与完整植物分离。植物的这些部分包括但不限于植物的器官、组织以及细胞，优选种子。
本发明的主题还是通过上述方法获得或可获得的植物。因此，本发明的一个实施方案是一种通过上述转化植物细胞以及从所述细胞再生植物的方法获得或可获得的转基因植物、以及其后代或部分，其中后代或部分包含至少一个感兴趣的核苷酸序列作为转基因。本发明的另一实施方案是一种通过上述修饰植物细胞的基因组以及从所述细胞再生植物的方法获得或可获得的遗传修饰植物及其后代或部分，其中后代或部分包含通过上述修饰方法引入的基因组中的修饰。
本发明的另一主题是一种衍生自上述转基因植物或遗传修饰植物的植物细胞或种子。这种植物细胞优选包含瞬时或稳定整合的编码GRF1多肽的多核苷酸以及优选识别所述细胞的基因组中的预定位点的单链DNA断裂(SSB)或双链DNA断裂(DSB)诱导酶或碱基编辑酶，并任选地包含修复核酸分子。编码GRF1多肽的多核苷酸优选可操作地连接到合适的调节序列，使得植物细胞能够表达GRF1多肽。调节序列是指例如“启动子”，其是指通常在其编码序列上游(5”)的核苷酸序列，其通过提供对RNA聚合酶以及正确转录所需的其它因子的识别来控制编码序列的表达。“构成型启动子”是指在几乎所有组织中且始终指导基因表达的启动子。构成型启动子的示例包括CaMV 35S启动子、双CaMV 35S启动子(70S启动子)、胭脂碱合酶(nos)启动子、BdEF1启动子或泛素启动子如PcUbi4或ZmUbi1。“调节型启动子”是指并非构成型地而是以时间和/或空间调节的方式指导基因表达的启动子，并且包括组织特异性以及诱导型启动子。它包括天然及合成序列以及可以是合成与天然序列的组合的序列。不同的启动子可以指导基因在不同的组织或细胞类型中、或在不同的发育阶段、或响应于不同的环境条件而表达。可用于植物细胞的各种类型的新启动子不断被发现并且是本领域的技术人员公知的。“组织特异性启动子”是指不是在所有植物细胞中表达而是仅在特定器官(诸如叶子或种子)、特定组织(诸如胚或子叶)或特定细胞类型(诸如叶实质细胞或种子贮存细胞)中的一个或多个细胞类型中表达的调节型启动子。这些启动子还包括在发育中的种子或果实的果实成熟期间、在完全分化的叶子中或在衰老开始时被暂时调节(诸如在早期或晚期胚发生中)的启动子。“诱导型启动子”是指可通过外部刺激(诸如化学、光、激素、应激或病原体)而在一个或多个细胞类型中开启的那些调节型启动子。诱导型启动子的示例是由蜕皮激素、地塞米松、乙醇诱导的启动子。这种启动子是本领域熟知的(例如，Samalova等(2005).pOp6/LhGR：用于烟草的严格调控且高度响应性的地塞米松诱导型基因表达系统(a stringently regulated and highly responsive dexamethasone-inducible gene expression system for tobacco)，The Plant Journal,41(6),919-935；Gatz&；Lenk(1998).响应于化学诱导剂的启动子(Promoters that respond tochemical inducers)，Trends in Plant Science，3(9)，352-358.)。
本发明的另一个主题是一种通过本发明的方法获得或可获得的单倍体植物胚。
本发明的另一主题是一种植物细胞，其包含瞬时或稳定整合的编码GRF1多肽的多核苷酸以及优选识别所述细胞的基因组中的预定位点的单链DNA断裂(SSB)或双链DNA断裂(DSB)诱导酶或碱基编辑酶，并任选地包含修复核酸分子，其中优选地，编码GRF1多肽的多核苷酸操作地连接到合适的调节序列，使得植物细胞能够表达GRF1多肽。当进行上述用于修饰植物细胞的基因组的方法时，可以获得这种植物细胞。
本发明的另一方面是编码GRF1多肽的多核苷酸、编码GRF1多肽的mRNA、GRF1多肽或编码GRF1多肽的内源基因的表达的激活因子在植物细胞的转化方法中、优选在如上所述的修饰基因组的方法中、在用于产生植物或单倍体植物胚的方法中、优选在如上所述的用于产生植物或单倍体植物胚的方法中、或在用于再生植物的方法中、优选在如上所述的用于再生植物的方法中的用途。根据优选方面，所述用途与编码WUS2多肽的多核苷酸、编码WUS2多肽的mRNA或WUS2多肽组合。
除非在实施例中另有说明，否则所有重组DNA技术均根据标准方案进行，如描述于：Sambrook等(1989)，Molecular Cloning:A Laboratory Manual，第二版，纽约冷泉港实验室出版社以及Ausubel等(1994)Current Protocols in Molecular Biology，CurrentProtocols，第1卷及第2卷，美国。用于植物分子工作的标准材料及方法描述于R.D.D.Cray的Plant Molecular Biology Labfax(1993)中，其由BIOS科学出版有限公司(英国)与英国布莱克韦尔科学出版公司(Blackwell Scientific Publications)联合出版。标准分子生物学技术的其它参考文献包括Sambrook及Russell(2001)Molecular Cloning:ALaboratory Manual，第三版，纽约冷泉港实验室出版社以及Brown(1998)MolecularBiology LabFax，第I卷及第II卷，第二版，学术出版社(英国)。用于聚合酶链式反应的标准材料及方法可见于：Dieffenbach及Dveksler(1995)PCR Primer:A Laboratory Manual，冷泉港实验室出版社以及McPherson等(2000)PCR-Basics:From Background to Bench，第一版，德国施普林格出版社。
本文提及或引用的所有专利、专利申请以及出版物或公开的公开内容(包括互联网上的出版物)均通过引用整体并入本文。
将参考本文描述的以下附图及实施例进一步描述本发明。然而，应理解，本发明不限于这些实施例。
附图
图1、通过tDT表达评估生物射弹递送(轰击后24小时)。A及C：用tDT构建体轰击的未成熟胚；A、RFP通道以及C、明视野；B及D：用ZmWUS2、ZmBBM及tDT三个构建体轰击的未成熟胚；B、RFP通道以及D、明视野。
图2、通过靶基因表达评估生物射弹递送：A：tDT基因在用tDT构建体轰击的样品中表达，但在野生型A188胚中不表达；B：ZmWUS2基因在用ZmWUS2构建体轰击的样品中表达，但在野生型A188胚或仅用tDT构建体轰击的样品中不表达；C：ZmBBM基因在用ZmBBM构建体轰击的样品中表达，但在野生型A188胚或仅用tDT构建体轰击的样品中不表达。
图3、ZmGRF1的表达在ZmWUS2及ZmBBM表达样品中上调。A：与单独用tDT轰击的样品相比，在用WUS2/BBM/tDT轰击的样品中ZmGRF1表达更高；B及C：与从农杆菌转化获得的tDT对照相比，在转基因WUS/BBM愈伤组织样品(B)以及小芽(C)中ZmGRF1表达显著增加。
图4、ZmGRF1的瞬时递送增强了再生。在仅有tDT的样品(A)、ZmWUS2/ZmGRF1/tDT样品(B)以及ZmWUS2/ZmPLT7/tDT(C)中进行轰击后1个月出现荧光胚结构。
图5、图4的荧光胚结构的定量。
图6、A188玉米品系的稳定转化中的转化频率的增加。将ZmGRF1与标准转化构建体相比，转化频率倍数改变。在实验1(左栏)中，由于事件的生长，比在标准转化中早3周获得转基因植物。
图7、A188玉米品系的稳定转化中的转化频率的增加。愈伤组织选择后的事件数量及大小。标准构建体(左)ZmGRF1(右)
图8、A188玉米品系的稳定转化中的转化频率的增加。GH中的ZmGRF1过表达单拷贝品系没有表型(左)并且它们是可育的(右)
图9、GRF1的共转化对转化效率的影响。2个构建体(A及B)的标准农杆菌介导的转化与利用含有GRF1表达二元载体的农杆菌进行的共转化进行比较。
图10、pAMK-BdUbi10-ZmBbm的载体图谱
图11、pAMK-BdEF1-ZmWus2的载体图谱
图12、pAMK-BdEF1-ZmGRF1的载体图谱
图13、pLH-Pat5077399-70Subi-tDT的载体图谱
实施例
1、生物射弹递送：
基因克隆以及构建体制备
使用从玉米A188未成熟胚分离的总RNA通过RT-PCR来克隆玉米PLT7(ZmPLT7；cDNA＝SEQ ID NO:4；蛋白质＝SEQ ID NO:5)、BBM(ZmBBM；cDNA＝SEQ ID NO:6；蛋白质＝SEQ IDNO:7)、WUS2(ZmWUS2；cDNA＝SEQ ID NO:8；蛋白质＝SEQ ID NO:9)以及GRF1(ZmGRF1；基因组DNA＝SEQ ID NO:1；cDNA＝SEQ ID NO:2；蛋白质＝SEQ ID NO:3)基因。将加强基因片段在克隆位点处克隆到表达载体pAMK-BdUbi10或pAMK-BdEF1中，并且在BdUbi10启动子或BdEF1启动子的控制下表达。两种启动子都是来自短柄草属的强构成型启动子。图10至图12示出了测序确认的构建体图。
制备用于轰击的玉米未成熟胚
授粉后9-12天，收获未成熟胚大小为0.8至1.8mm、优选1.0-1.5mm的玉米穗。用70％乙醇对玉米穗灭菌10-15分钟。在层流通风橱中短暂风干后，用鲨鱼解剖刀从玉米穗中取出核仁的顶部～1/3，并用刮刀小心地将未成熟胚从核仁中拉出。将新鲜分离的胚置于渗透培养基平板(见下文)中的轰击靶区上，其中盾片侧向上。用石蜡膜包裹平板，并在轰击前在黑暗中于25℃下培育4小时。
颗粒共轰击
使用粒子轰击枪以及尺寸为0.4或0.6微米(μm)的金颗粒将DNA递送到玉米未成熟胚的盾片细胞中。将质粒与感兴趣的基因(GOI)预混合。对于10次轰击(shot)，将总体积为100微升(μl)的50％(v/v)甘油中的1mg金粒子(每次注射100μg金粒子)移液到透明的低保留微量离心管中。超声处理15秒以悬浮金颗粒。在低速涡旋的同时，向每100μl金颗粒中依序添加以下各项：
-—最多10μl DNA(预混合1.0-10.0μg总DNA，每次轰击100-1000ng)
-—100μl 2.5M CaCl2(在冰上预冷)
-—40μl 0.1M冷亚精胺
盖上盖子，并且在0-10℃下将管涡旋2-30分钟，并将DNA包被的金粒子旋转减慢。在500μl 100％乙醇中洗涤两次后，将沉淀物重悬于120μl 100％乙醇中。在低速涡旋的同时，用宽开口的20μl尖端将10μl共涂覆的金粒子从管中均匀地移液到大载体的中心上，因为粒子易于在此时形成团块，因此使金粒子尽可能快地到达大载体上。空气干燥。使用Bio-Rad PDS-1000/He粒子枪进行轰击。轰击条件是：28mm/Hg真空，450或650psi破裂盘，6mm间隙距离，样品平台在距室中的底部60mm距离的第二位置，每个样品(玉米未成熟胚)平板三次注射。
轰击后观察及胚培养
在轰击后，使胚在渗透培养基上再保持16小时。在轰击后16-20小时，使用荧光显微镜以最大激发554nm以及最大发射581nm检查瞬时转化的tDT表达。选择在荧光显微镜下具有密集荧光信号的胚，并将其从N6OSM转移到N6-5Ag平板(每个平板约15个胚)上(其中盾片面朝上)以用于愈伤组织诱导。
渗透培养基：N6盐、N6维生素、1.0mg/L 2,4-D、100mg/L酪蛋白、0.7g/L L-脯氨酸、0.2M甘露醇(36.4g/L)、0.2M山梨醇(36.4g/L)、20g/L蔗糖、15g/L细菌培养用琼脂、pH 5.8。
N6-5Ag：N6盐、N6维生素、1.0mg/L 2,4-D、100mg/L酪蛋白、2.9g/L L-脯氨酸、20g/L蔗糖、5g/L葡萄糖、5mg/L AgNO3、8g/L细菌培养用琼脂、pH 5.8。
2、ZmWUS2及ZmBBM靶的识别：
在A188未成熟胚的生物射弹递送中使用以下构建体：pAMK-BdUbi10Zm-Bbm、pAMK-BdEF1ZmWus2、pLH-Pat5077399-70Subi-tDt
实验设置如下：
在生物射弹递送之后的24小时，通过tDT表达评估质粒递送的成功。图1表示对类别2及3的轰击是成功的。进一步，测定了引入的基因tDT、ZmWUS2以及ZmBBM(类别3)的表达(图2)。
3、RNAseq分析：
在生物射弹递送之后的24小时，收集未成熟的胚并在液氮中快速冷冻。随后将所有样品送去进行RNA提取、文库制备以及RNAseq，RNAseq是由外部测序提供者完成。
通过在仅包含tDT的样品(类别2)与包含ZmWUS2、ZmBBM以及tDT的样品(类别3)之间进行比较来鉴定几种候选物。十六个基因响应于ZmWUS2以及ZmBBM的表达而显示出上调的表达，五个基因显示出下调的表达。在通过qRT-PCR进行确认之后，选择候选物ZmGRF1用于进一步分析。图3A显示了在使用包含ZmWUS2、ZmBBM以及TDt的构建体轰击的样品(类别3)中ZmGRF1表达较高。另外，通过农杆菌介导的转化而稳定地整合了ZmWUS2以及ZmBBM。在玉米植物的愈伤组织样品以及小芽中评估了ZmGRF1的表达。与仅通过tDT基因的农杆菌转化获得的对照相比，在WUS2/BBM过表达植物材料中ZmGRF1的表达显著增加(图3B及图C)。
4、用于测试ZmGRF1的瞬时测定
使用以下构建体轰击A188未成熟胚：
在轰击之后的一个月，计数荧光胚结构。这种结构表示增强的再生能力。仅用tDT瞬时转化的样品显示出较不明显的胚结构(类别1；图4A)。根据ZmWUS2与ZmPLT7的组合，已知可以实现再生的增强。因此，该类别3用作阳性对照(图4C)。更令人印象深刻的是，在瞬时表达ZmWUS2及ZmGRF1以及tDT的样品(类别2)中荧光胚结构的出现比在阳性对照中更明显(图4B)。定量证实了视觉结果，并显示出在瞬时表达ZmWUS2及ZmGRF1以及tDT的样品(类别2)中荧光胚结构的百分比比在类别1中高5倍以上，且比在类别3中高2倍以上(图5)。
5、玉米转化方法
按照标准方案(Ishida Y、Saito H、Ohta S、Hiei Y、Komari T、Kumashiro T(1996)High efficiency transformation of maize(Zea mays L.)mediated byAgrobacterium tumefaciens Nat Biotechnol 14:745-750)，使用农杆菌进行玉米转化(品系A188)。简言之，用农杆菌接种A188品系的未成熟胚，并共培养7天。在初始胚性愈伤组织产生后，使用选择培养基，其中针对转化事件选择除草剂。提取推定的转基因植物的DNA并通过qPCR分析可选择的标记基因的存在。转化率被计算为每个接种胚的转基因事件数(％)。当进行共转化时，在接种前将等量的包含构建体与GRF1基因的农杆菌混合在一起。
结果表明，与标准转化相比，即在没有额外递送ZmGRF1的条件下，达到了高达4倍的转化效率。另外，有利的是，由于事件的快速成长，在大约3周前获得转基因植物。图6示出了两个独立实验的定量分析，图7示出了视觉比较。在温室中生长的具有ZmGRF1单拷贝事件的玉米植物品系没有不希望的表型并且是可育的(图8)。
进一步，研究了GRF1的共转化对转化效率的影响。比较了2个构建体(A及B)的标准农杆菌介导的转化与利用含有GRF1表达二元载体的农杆菌进行的共转化。当使用GRF1时，转化率高2.3及1.8倍(图9)。另外，共转化实验比标准协议快36％。
序列表
<110> 科沃施种子欧洲股份两合公司
<120> 通过使用GRF1加强基因实现的增强的植物再生及转化
<130> KWS0320PCT
<150> EP19183486.0
<151> 2019-06-28
<160> 53
<170> PatentIn version 3.5
<210> 1
<211> 1260
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of ZmGRF1
<400> 1
atggctatgc catacgcttc tctgtctcca gccggcgctg ctgatcatag gtcatctacc 60
gctaccgcca gcctggttcc gttctgcagg tctactccac tgtctgctgg cggcggtctt 120
ggcgaggaag atgctcaagc ttctgccagg tggccagctg ctaggccagt ggttcctttc 180
actccggcgc agtaccaaga gttggagcag caggccctga tctacaagta cctggtggct 240
ggcgtgccag tgccaccaga tcttgttgtt ccaatccgca ggggcctcga ctctctggct 300
actaggtttt acggccagcc gacacttggc tacggcccat accttggtag gaagctggac 360
ccagagcctg gtagatgcag aaggaccgac ggtaagaagt ggcgctgcag caaagaggct 420
gccccggact ctaagtactg cgagagacat atgcacaggg gccgcaacag aagccgcaag 480
ccagttgaga ctcagctggc cccacagagc caaccaccag ccgctgctgc tgtttcagct 540
gctccaccac ttgctgctgc agctgctgct accactaacg gctctggctt ccagaaccac 600
tctctgtacc cagctatcgc tggctctact ggcggaggtg gcggcgttgg cggttctggc 660
aacattagca gcccattcag cagctccatg ggcggctcat ctcagctgca catggacagc 720
gctgccagct actcttacgc tgctcttggc ggaggcaccg ctaaggacct taggtacaac 780
gcttacggca tccgcagcct ggctgacgag cataaccagc ttatcgccga ggccatcgac 840
agcagcatcg aatctcaatg gcgcctgccg tccagcagct tcccactttc atcttaccca 900
cacctgggcg ctctgggcga tcttggcggc cagaactcta ctgtgagcag cctgccgaag 960
atggaaaagc agcaaccgcc gtctagcttc ctgggcaacg atactggtgc cggtatggct 1020
atgggcagcg cctctgctaa gcaagagggc cagactctga ggcacttctt cgacgagtgg 1080
ccaaaggcta gggactcttg gccaggcctc tctgacgaga ctgctagcct ggcatctttc 1140
ccaccagcca cgcagctcag catgagcatc ccaatggcta gcagcgactt cagcgtcgcc 1200
agctctcagt ctccaaacgg cgagtctagg accttcctgc tggctaccga taggcgctga 1260
<210> 2
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<212> PRT
<213> Zea mays
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Arg Ser Ser Thr Ala Thr Ala Ser Leu Val Pro Phe Cys Arg Ser Thr
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Pro Leu Ser Ala Gly Gly Gly Leu Gly Glu Glu Asp Ala Gln Ala Ser
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Ala Arg Trp Pro Ala Ala Arg Pro Val Val Pro Phe Thr Pro Ala Gln
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Pro Val Glu Thr Gln Leu Ala Pro Gln Ser Gln Pro Pro Ala Ala Ala
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Ala Val Ser Ala Ala Pro Pro Leu Ala Ala Ala Ala Ala Ala Thr Thr
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Leu Arg Tyr Asn Ala Tyr Gly Ile Arg Ser Leu Ala Asp Glu His Asn
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Gln Leu Ile Ala Glu Ala Ile Asp Ser Ser Ile Glu Ser Gln Trp Arg
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<223> cDNA of ZmPLT7
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atggacatga gctcagctta tccccaccat tggctctcct tctccctctc caacaactac 60
caccatggcc tactcgaggc cttctctaac tcctccggta ctcctcttgg agacgagccg 120
ggcgcagtgg aggagtcccc gaggacggtg gaggacttcc tcggcggcgt cggtggcgcc 180
ggcgccccgc cgcagccggc ggctgctgca gatcaggatc accagcttgt gtgcggcgag 240
ctgggcagca tcacagccag gttcttgcgc cactacccgg cggcgccagc tgggacgacg 300
gtggagaacc ccggcgcggt gaccgtggcg gccatgtcgt cgacggacgt ggcgggggcg 360
gagtccgacc aggcgaggcg gcccgccgag acgttcggcc agcgcacatc catctaccgt 420
ggcgtcacca ggcaccggtg gacagggaga tatgaggcgc acttgtggga caacagctgc 480
cgccgggagg gccaaagccg caaaggacgc caagtctacc taggaggcta tgacaaggag 540
gagaaggcgg ctagagctta cgacctcgcc gcgctcaagt actgggggcc tacaaccacg 600
accaacttcc cggtgtccaa ctacgagaag gagctggagg agatgaagtc catgacgcgg 660
caggagttca tcgcgtcgtt gcgcaggaag agcagcggct tctcacgagg cgcctccatc 720
tacagaggag tcacaaggca tcatcagcac ggccggtggc aggcgaggat cggcagggtg 780
gccggaaaca aggacctgta cttgggcact ttcagtactc aggaagaggc ggcggaggcg 840
tacgacatcg ctgcgatcaa gttccgcggg ctcaacgccg tcaccaactt cgacatgagc 900
cgctacgacg tggagagcat cctcagcagc gacctccccg tcgggggcgg agccaccggg 960
cgcgccgcca agttcccgtt ggactcgctg cagccgggga gcgctgctgc gatgatgctc 1020
gccggggctg ctgccgcttc gcaggccacc atgccgccgt ccgagaagga ctactggtct 1080
ctgctcgccc tgcactacca gcagcagcag gagcaggagc ggcagttccc ggcttctgct 1140
tacgaggctt acggctccgg cggcgtgaac gtggacttca cgatgggcac cagtagcggc 1200
aacaacaaca acaacaccgg cagcggcgtc atgtggggcg ccaccactgg tgcagtagta 1260
gtgggacagc aagacagcag cggcaagcag ggcaacggct atgccagcaa cattccttat 1320
gctgctgctg ctgctatggt ttctggatct gctggctacg agggctccac cggcgacaat 1380
ggaacctggg ttactacgac tattaccagc agcaacaccg gcacggctcc ccactactac 1440
aactatctct tcgggatgga gtag 1464
<210> 4
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<212> PRT
<213> Zea mays
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Met Asp Met Ser Ser Ala Tyr Pro His His Trp Leu Ser Phe Ser Leu
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Ser Asn Asn Tyr His His Gly Leu Leu Glu Ala Phe Ser Asn Ser Ser
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Gly Thr Pro Leu Gly Asp Glu Pro Gly Ala Val Glu Glu Ser Pro Arg
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Thr Val Glu Asp Phe Leu Gly Gly Val Gly Gly Ala Gly Ala Pro Pro
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Gln Pro Ala Ala Ala Ala Asp Gln Asp His Gln Leu Val Cys Gly Glu
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Leu Gly Ser Ile Thr Ala Arg Phe Leu Arg His Tyr Pro Ala Ala Pro
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Ala Gly Thr Thr Val Glu Asn Pro Gly Ala Val Thr Val Ala Ala Met
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Ser Ser Thr Asp Val Ala Gly Ala Glu Ser Asp Gln Ala Arg Arg Pro
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Ala Glu Thr Phe Gly Gln Arg Thr Ser Ile Tyr Arg Gly Val Thr Arg
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His Arg Trp Thr Gly Arg Tyr Glu Ala His Leu Trp Asp Asn Ser Cys
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Arg Arg Glu Gly Gln Ser Arg Lys Gly Arg Gln Val Tyr Leu Gly Gly
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Lys Tyr Trp Gly Pro Thr Thr Thr Thr Asn Phe Pro Val Ser Asn Tyr
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Ala Ser Leu Arg Arg Lys Ser Ser Gly Phe Ser Arg Gly Ala Ser Ile
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Tyr Arg Gly Val Thr Arg His His Gln His Gly Arg Trp Gln Ala Arg
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Thr Gln Glu Glu Ala Ala Glu Ala Tyr Asp Ile Ala Ala Ile Lys Phe
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Arg Gly Leu Asn Ala Val Thr Asn Phe Asp Met Ser Arg Tyr Asp Val
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Arg Ala Ala Lys Phe Pro Leu Asp Ser Leu Gln Pro Gly Ser Ala Ala
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Ala Met Met Leu Ala Gly Ala Ala Ala Ala Ser Gln Ala Thr Met Pro
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Pro Ser Glu Lys Asp Tyr Trp Ser Leu Leu Ala Leu His Tyr Gln Gln
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Gln Gln Glu Gln Glu Arg Gln Phe Pro Ala Ser Ala Tyr Glu Ala Tyr
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Gly Ser Gly Gly Val Asn Val Asp Phe Thr Met Gly Thr Ser Ser Gly
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Asn Asn Asn Asn Asn Thr Gly Ser Gly Val Met Trp Gly Ala Thr Thr
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Gly Ala Val Val Val Gly Gln Gln Asp Ser Ser Gly Lys Gln Gly Asn
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Gly Tyr Ala Ser Asn Ile Pro Tyr Ala Ala Ala Ala Ala Met Val Ser
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Gly Ser Ala Gly Tyr Glu Gly Ser Thr Gly Asp Asn Gly Thr Trp Val
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Thr Thr Thr Ile Thr Ser Ser Asn Thr Gly Thr Ala Pro His Tyr Tyr
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Asn Tyr Leu Phe Gly Met Glu
485
<210> 5
<211> 2133
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of ZmBBM
<400> 5
atggccactg tgaacaactg gctcgctttc tccctctccc cgcaggagct gccgccctcc 60
cagacgacgg actccacact catctcggcc gccaccgccg accatgtctc cggcgatgtc 120
tgcttcaaca tcccccaaga ttggagcatg aggggatcag agctttcggc gctcgtcgcg 180
gagccgaagc tggaggactt cctcggcggc atctccttct ccgagcagca tcacaaggcc 240
aactgcaaca tgatacccag cactagcagc acagtttgct acgcgagctc aggtgctagc 300
accggctacc atcaccagct gtaccaccag cccaccagct cagcgctcca cttcgcggac 360
tccgtaatgg tcgcctcctc ggccggtgtc cacgacggcg gtgccatgct cagcgcggcc 420
gccgctaacg gtgtcgctgg cgctgccagt gccaacggcg gcggcatcgg gctgtccatg 480
attaagaact ggctgcggag ccaaccggcg cccatgcagc cgagggtggc ggcggctgag 540
ggcgcgcagg ggctctcttt gtcgatgaac atggcgggga cgacccaagg cgctgctggc 600
atgccacttc tcgctggaga gcgcgcacgg gcgcccgaga gtgtatcgac gtcagcacag 660
ggtggagccg tcgtcgtcac ggcgccgaag gaggatagcg gtggcagcgg tgttgccggc 720
gctctagtag ccgtgagcac ggacacgggt ggcagcggcg gcgcgtcggc tgacaacacg 780
gcaaggaaga cggtggacac gttcgggcag cgcacgtcga tttaccgtgg cgtgacaagg 840
catagatgga ctgggagata tgaggcacat ctttgggata acagttgcag aagggaaggg 900
caaactcgta agggtcgtca agtctattta ggtggctatg ataaagagga gaaagctgct 960
agggcttatg atcttgctgc tctgaagtac tggggtgcca caacaacaac aaattttcca 1020
gtgagtaact acgaaaagga gctcgaggac atgaagcaca tgacaaggca ggagtttgta 1080
gcgtctctga gaaggaagag cagtggtttc tccagaggtg catccattta caggggagtg 1140
actaggcatc accaacatgg aagatggcaa gcacggattg gacgagttgc agggaacaag 1200
gatctttact tgggcacctt cagcacccag gaggaggcag cggaggcgta cgacatcgcg 1260
gcgatcaagt tccgcggcct caacgccgtc accaacttcg acatgagccg ctacgacgtg 1320
aagagcatcc tggacagcag cgccctcccc atcggcagcg ccgccaagcg cctcaaggag 1380
gccgaggccg cagcgtccgc gcagcaccac cacgccggcg tggtgagcta cgacgtcggc 1440
cgcatcgcct cgcagctcgg cgacggcgga gccctggcgg cggcgtacgg cgcgcactac 1500
cacggcgccg cctggccgac catcgcgttc cagccgggcg ccgccagcac aggcctgtac 1560
cacccgtacg cgcagcagcc aatgcgcggc ggcgggtggt gcaagcagga gcaggaccac 1620
gcggtgatcg cggccgcgca cagcctccag gacctccacc acctgaacct gggcgcggcc 1680
ggcgcgcacg actttttctc ggcagggcag caggccgccg ccgctgcgat gcacggcctg 1740
ggtagcatcg acagtgcgtc gctcgagcac agcaccggct ccaactccgt cgtctacaac 1800
ggcggggtcg gcgacagcaa cggcgccagc gccgtcggcg gcagtggcgg tggctacatg 1860
atgccgatga gcgctgccgg agcaaccact acatcggcaa tggtgagcca cgagcaggtg 1920
catgcacggg cctacgacga agccaagcag gctgctcaga tggggtacga gagctacctg 1980
gtgaacgcgg agaacaatgg tggcggaagg atgtctgcat gggggactgt cgtgtctgca 2040
gccgcggcgg cagcagcaag cagcaacgac aacatggccg ccgacgtcgg ccatggcggc 2100
gcgcagctct tcagtgtctg gaacgacact taa 2133
<210> 6
<211> 710
<212> PRT
<213> Zea mays
<400> 6
Met Ala Thr Val Asn Asn Trp Leu Ala Phe Ser Leu Ser Pro Gln Glu
1 5 10 15
Leu Pro Pro Ser Gln Thr Thr Asp Ser Thr Leu Ile Ser Ala Ala Thr
20 25 30
Ala Asp His Val Ser Gly Asp Val Cys Phe Asn Ile Pro Gln Asp Trp
35 40 45
Ser Met Arg Gly Ser Glu Leu Ser Ala Leu Val Ala Glu Pro Lys Leu
50 55 60
Glu Asp Phe Leu Gly Gly Ile Ser Phe Ser Glu Gln His His Lys Ala
65 70 75 80
Asn Cys Asn Met Ile Pro Ser Thr Ser Ser Thr Val Cys Tyr Ala Ser
85 90 95
Ser Gly Ala Ser Thr Gly Tyr His His Gln Leu Tyr His Gln Pro Thr
100 105 110
Ser Ser Ala Leu His Phe Ala Asp Ser Val Met Val Ala Ser Ser Ala
115 120 125
Gly Val His Asp Gly Gly Ala Met Leu Ser Ala Ala Ala Ala Asn Gly
130 135 140
Val Ala Gly Ala Ala Ser Ala Asn Gly Gly Gly Ile Gly Leu Ser Met
145 150 155 160
Ile Lys Asn Trp Leu Arg Ser Gln Pro Ala Pro Met Gln Pro Arg Val
165 170 175
Ala Ala Ala Glu Gly Ala Gln Gly Leu Ser Leu Ser Met Asn Met Ala
180 185 190
Gly Thr Thr Gln Gly Ala Ala Gly Met Pro Leu Leu Ala Gly Glu Arg
195 200 205
Ala Arg Ala Pro Glu Ser Val Ser Thr Ser Ala Gln Gly Gly Ala Val
210 215 220
Val Val Thr Ala Pro Lys Glu Asp Ser Gly Gly Ser Gly Val Ala Gly
225 230 235 240
Ala Leu Val Ala Val Ser Thr Asp Thr Gly Gly Ser Gly Gly Ala Ser
245 250 255
Ala Asp Asn Thr Ala Arg Lys Thr Val Asp Thr Phe Gly Gln Arg Thr
260 265 270
Ser Ile Tyr Arg Gly Val Thr Arg His Arg Trp Thr Gly Arg Tyr Glu
275 280 285
Ala His Leu Trp Asp Asn Ser Cys Arg Arg Glu Gly Gln Thr Arg Lys
290 295 300
Gly Arg Gln Val Tyr Leu Gly Gly Tyr Asp Lys Glu Glu Lys Ala Ala
305 310 315 320
Arg Ala Tyr Asp Leu Ala Ala Leu Lys Tyr Trp Gly Ala Thr Thr Thr
325 330 335
Thr Asn Phe Pro Val Ser Asn Tyr Glu Lys Glu Leu Glu Asp Met Lys
340 345 350
His Met Thr Arg Gln Glu Phe Val Ala Ser Leu Arg Arg Lys Ser Ser
355 360 365
Gly Phe Ser Arg Gly Ala Ser Ile Tyr Arg Gly Val Thr Arg His His
370 375 380
Gln His Gly Arg Trp Gln Ala Arg Ile Gly Arg Val Ala Gly Asn Lys
385 390 395 400
Asp Leu Tyr Leu Gly Thr Phe Ser Thr Gln Glu Glu Ala Ala Glu Ala
405 410 415
Tyr Asp Ile Ala Ala Ile Lys Phe Arg Gly Leu Asn Ala Val Thr Asn
420 425 430
Phe Asp Met Ser Arg Tyr Asp Val Lys Ser Ile Leu Asp Ser Ser Ala
435 440 445
Leu Pro Ile Gly Ser Ala Ala Lys Arg Leu Lys Glu Ala Glu Ala Ala
450 455 460
Ala Ser Ala Gln His His His Ala Gly Val Val Ser Tyr Asp Val Gly
465 470 475 480
Arg Ile Ala Ser Gln Leu Gly Asp Gly Gly Ala Leu Ala Ala Ala Tyr
485 490 495
Gly Ala His Tyr His Gly Ala Ala Trp Pro Thr Ile Ala Phe Gln Pro
500 505 510
Gly Ala Ala Ser Thr Gly Leu Tyr His Pro Tyr Ala Gln Gln Pro Met
515 520 525
Arg Gly Gly Gly Trp Cys Lys Gln Glu Gln Asp His Ala Val Ile Ala
530 535 540
Ala Ala His Ser Leu Gln Asp Leu His His Leu Asn Leu Gly Ala Ala
545 550 555 560
Gly Ala His Asp Phe Phe Ser Ala Gly Gln Gln Ala Ala Ala Ala Ala
565 570 575
Met His Gly Leu Gly Ser Ile Asp Ser Ala Ser Leu Glu His Ser Thr
580 585 590
Gly Ser Asn Ser Val Val Tyr Asn Gly Gly Val Gly Asp Ser Asn Gly
595 600 605
Ala Ser Ala Val Gly Gly Ser Gly Gly Gly Tyr Met Met Pro Met Ser
610 615 620
Ala Ala Gly Ala Thr Thr Thr Ser Ala Met Val Ser His Glu Gln Val
625 630 635 640
His Ala Arg Ala Tyr Asp Glu Ala Lys Gln Ala Ala Gln Met Gly Tyr
645 650 655
Glu Ser Tyr Leu Val Asn Ala Glu Asn Asn Gly Gly Gly Arg Met Ser
660 665 670
Ala Trp Gly Thr Val Val Ser Ala Ala Ala Ala Ala Ala Ala Ser Ser
675 680 685
Asn Asp Asn Met Ala Ala Asp Val Gly His Gly Gly Ala Gln Leu Phe
690 695 700
Ser Val Trp Asn Asp Thr
705 710
<210> 7
<211> 909
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of ZmWUS2
<400> 7
atggcggcca atgcgggcgg cggtggagcg ggaggaggca gcggcagcgg cagcgtggct 60
gcgccggcgg tgtgccgccc cagcggctcg cggtggacgc cgacgccgga gcagatcagg 120
atgctgaagg agctctacta cggctgcggc atccggtcgc ccagctcgga gcagatccag 180
cgcatcaccg ccatgctgcg gcagcacggc aagatcgagg gcaagaacgt cttctactgg 240
ttccagaacc acaaggcccg cgagcgccag aagcgccgcc tcaccagcct cgacgtcaac 300
gtgcccgccg ccggcgcggc cgacgccacc accagccaac tcggcgtcct ctcgctgtcg 360
tcgccgccgc cttcaggcgc ggcgcctccc tcgcccaccc tcggcttcta cgccgccggc 420
aatggcggcg gatcggctgt gctgctggac acgagttccg actggggcag cagcggcgct 480
gccatggcca ccgagacatg cttcctccag gactacatgg gcgtgacgga cacgggcagc 540
tcgtcgcagt ggccacgctt ctcgtcgtcg gacacgataa tggcggcggc cgcggcgcgg 600
gcggcgacga cgcgggcgcc cgagacgctc cctctcttcc cgacctgcgg cgacgacggc 660
ggcagcggta gcagcagcta cttgccgttc tggggtgccg cgtccacaac tgccggcgcc 720
acttcttccg ttgcgatcca gcagcaacac cagctgcagg agcagtacag cttttacagc 780
aacagcaaca gcacccagct ggccggcacc ggcaaccaag acgtatcggc aacagcagca 840
gcagccgccg ccctggagct gagcctcagc tcatggtgct ccccttaccc tgctgcaggg 900
agtatgtga 909
<210> 8
<211> 302
<212> PRT
<213> Zea mays
<400> 8
Met Ala Ala Asn Ala Gly Gly Gly Gly Ala Gly Gly Gly Ser Gly Ser
1 5 10 15
Gly Ser Val Ala Ala Pro Ala Val Cys Arg Pro Ser Gly Ser Arg Trp
20 25 30
Thr Pro Thr Pro Glu Gln Ile Arg Met Leu Lys Glu Leu Tyr Tyr Gly
35 40 45
Cys Gly Ile Arg Ser Pro Ser Ser Glu Gln Ile Gln Arg Ile Thr Ala
50 55 60
Met Leu Arg Gln His Gly Lys Ile Glu Gly Lys Asn Val Phe Tyr Trp
65 70 75 80
Phe Gln Asn His Lys Ala Arg Glu Arg Gln Lys Arg Arg Leu Thr Ser
85 90 95
Leu Asp Val Asn Val Pro Ala Ala Gly Ala Ala Asp Ala Thr Thr Ser
100 105 110
Gln Leu Gly Val Leu Ser Leu Ser Ser Pro Pro Pro Ser Gly Ala Ala
115 120 125
Pro Pro Ser Pro Thr Leu Gly Phe Tyr Ala Ala Gly Asn Gly Gly Gly
130 135 140
Ser Ala Val Leu Leu Asp Thr Ser Ser Asp Trp Gly Ser Ser Gly Ala
145 150 155 160
Ala Met Ala Thr Glu Thr Cys Phe Leu Gln Asp Tyr Met Gly Val Thr
165 170 175
Asp Thr Gly Ser Ser Ser Gln Trp Pro Arg Phe Ser Ser Ser Asp Thr
180 185 190
Ile Met Ala Ala Ala Ala Ala Arg Ala Ala Thr Thr Arg Ala Pro Glu
195 200 205
Thr Leu Pro Leu Phe Pro Thr Cys Gly Asp Asp Gly Gly Ser Gly Ser
210 215 220
Ser Ser Tyr Leu Pro Phe Trp Gly Ala Ala Ser Thr Thr Ala Gly Ala
225 230 235 240
Thr Ser Ser Val Ala Ile Gln Gln Gln His Gln Leu Gln Glu Gln Tyr
245 250 255
Ser Phe Tyr Ser Asn Ser Asn Ser Thr Gln Leu Ala Gly Thr Gly Asn
260 265 270
Gln Asp Val Ser Ala Thr Ala Ala Ala Ala Ala Ala Leu Glu Leu Ser
275 280 285
Leu Ser Ser Trp Cys Ser Pro Tyr Pro Ala Ala Gly Ser Met
290 295 300
<210> 9
<211> 783
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of artificial regernation booster protein 1
<400> 9
atggagtcgg gctccgggac ggctgctggc tctggctatg tttacagaca gccaggatca 60
acgcggtgga acccgacagc tgaacaactg tccttgctta gagaaatcta ctaccgcaac 120
ggattgcgga ccccgaccgc ggacgaaatc agacaaatca gctcaaagct ctcaaggtac 180
ggaaaaatag agggcaaaaa cgtttacaac tggttccaga atagacgcgc aagagaaaag 240
cgcaagcaac ggctctctac aatcggctgt gatccagcac tgatcgagat ggggaatgtc 300
gcttcactgg aattcggtac tgagagcgcc ctggaatcgc tgtcgtcagg accatcctca 360
gaactccgcg aagcgccaac gagaaaattt tacgaaaaaa agacggttgg agagaactca 420
actataataa acccagtgga acaaaactgt accctttcct gcggaacgtc ccaagagttc 480
cagtatgcgg tcgattctcg gcgcgtcatg aaagctatgg aggaaaagca ggcgacggac 540
gatgaacccg acggaaataa atggactgag tcaaacagac acgtcaagat tctccagctt 600
ttcccgctcc acaataacga ggatcagaca ttgataaaga gcgacaaaga aatctattgt 660
ttgggctcgt gcgagaagaa aatggatttg tcaccgctgg gtcattcagg ctctcagcgc 720
gcttcggccc ttgacttgtg cctttcattg ggcaacgaat cttgtgggct gcatgataat 780
tga 783
<210> 10
<211> 260
<212> PRT
<213> Artificial Sequence
<220>
<223> artificial regernation booster protein 1
<400> 10
Met Glu Ser Gly Ser Gly Thr Ala Ala Gly Ser Gly Tyr Val Tyr Arg
1 5 10 15
Gln Pro Gly Ser Thr Arg Trp Asn Pro Thr Ala Glu Gln Leu Ser Leu
20 25 30
Leu Arg Glu Ile Tyr Tyr Arg Asn Gly Leu Arg Thr Pro Thr Ala Asp
35 40 45
Glu Ile Arg Gln Ile Ser Ser Lys Leu Ser Arg Tyr Gly Lys Ile Glu
50 55 60
Gly Lys Asn Val Tyr Asn Trp Phe Gln Asn Arg Arg Ala Arg Glu Lys
65 70 75 80
Arg Lys Gln Arg Leu Ser Thr Ile Gly Cys Asp Pro Ala Leu Ile Glu
85 90 95
Met Gly Asn Val Ala Ser Leu Glu Phe Gly Thr Glu Ser Ala Leu Glu
100 105 110
Ser Leu Ser Ser Gly Pro Ser Ser Glu Leu Arg Glu Ala Pro Thr Arg
115 120 125
Lys Phe Tyr Glu Lys Lys Thr Val Gly Glu Asn Ser Thr Ile Ile Asn
130 135 140
Pro Val Glu Gln Asn Cys Thr Leu Ser Cys Gly Thr Ser Gln Glu Phe
145 150 155 160
Gln Tyr Ala Val Asp Ser Arg Arg Val Met Lys Ala Met Glu Glu Lys
165 170 175
Gln Ala Thr Asp Asp Glu Pro Asp Gly Asn Lys Trp Thr Glu Ser Asn
180 185 190
Arg His Val Lys Ile Leu Gln Leu Phe Pro Leu His Asn Asn Glu Asp
195 200 205
Gln Thr Leu Ile Lys Ser Asp Lys Glu Ile Tyr Cys Leu Gly Ser Cys
210 215 220
Glu Lys Lys Met Asp Leu Ser Pro Leu Gly His Ser Gly Ser Gln Arg
225 230 235 240
Ala Ser Ala Leu Asp Leu Cys Leu Ser Leu Gly Asn Glu Ser Cys Gly
245 250 255
Leu His Asp Asn
260
<210> 11
<211> 783
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of artificial regernation booster protein 2
<400> 11
atggaatcgg gctccggcac ggcggcaggg tctggttatg tctatcggca gagcggaagc 60
acccggtgga atccaacagc agaacagttg tcgctgctca aggaacttta ttaccggaat 120
ggaattcgga caccgtcggc agatcaaatt aggcaaattt cggcccggct gtccagatac 180
ggcaaaatag aagggaaaaa cgtcttttac tggtttcaaa atcataaagc acgggaacgg 240
cagaagaaaa gactttccac ggtcggctgc gaccctgctc tcatagaaat gggtaacgtc 300
gcgagcttgg aatttgggac cgaaagcgct cttgaatctc tcagctcagg cccgtccagc 360
gagttgcgcg aggctcctac ccgcaagttt tatgagaaga aaaccgttgg tgagaacagc 420
accataatca atcctgttga gcagaactgc acactttctt gcggtacttc gcaggaattt 480
cagtatgctg ttgatagccg ccgggtgatg aaggcaatgg aagagaagca agcaacggat 540
gatgaaccgg acggaaacaa atggacggag tcgaacaggc atgtgaagac cctccctctt 600
ttccccttgc ataataatga agatcagacc ttgatcaagt cggacaagga aatttattgc 660
cttgggagct gtgaaaaaaa aatggatctg tccccattgg gacactcggg ctctcagagg 720
gcgtcggcac tggatttgtg cctgtctttg ggtaatgaat cttgtggcct ccacgacaat 780
tga 783
<210> 12
<211> 260
<212> PRT
<213> Artificial Sequence
<220>
<223> artificial regernation booster protein 2
<400> 12
Met Glu Ser Gly Ser Gly Thr Ala Ala Gly Ser Gly Tyr Val Tyr Arg
1 5 10 15
Gln Ser Gly Ser Thr Arg Trp Asn Pro Thr Ala Glu Gln Leu Ser Leu
20 25 30
Leu Lys Glu Leu Tyr Tyr Arg Asn Gly Ile Arg Thr Pro Ser Ala Asp
35 40 45
Gln Ile Arg Gln Ile Ser Ala Arg Leu Ser Arg Tyr Gly Lys Ile Glu
50 55 60
Gly Lys Asn Val Phe Tyr Trp Phe Gln Asn His Lys Ala Arg Glu Arg
65 70 75 80
Gln Lys Lys Arg Leu Ser Thr Val Gly Cys Asp Pro Ala Leu Ile Glu
85 90 95
Met Gly Asn Val Ala Ser Leu Glu Phe Gly Thr Glu Ser Ala Leu Glu
100 105 110
Ser Leu Ser Ser Gly Pro Ser Ser Glu Leu Arg Glu Ala Pro Thr Arg
115 120 125
Lys Phe Tyr Glu Lys Lys Thr Val Gly Glu Asn Ser Thr Ile Ile Asn
130 135 140
Pro Val Glu Gln Asn Cys Thr Leu Ser Cys Gly Thr Ser Gln Glu Phe
145 150 155 160
Gln Tyr Ala Val Asp Ser Arg Arg Val Met Lys Ala Met Glu Glu Lys
165 170 175
Gln Ala Thr Asp Asp Glu Pro Asp Gly Asn Lys Trp Thr Glu Ser Asn
180 185 190
Arg His Val Lys Thr Leu Pro Leu Phe Pro Leu His Asn Asn Glu Asp
195 200 205
Gln Thr Leu Ile Lys Ser Asp Lys Glu Ile Tyr Cys Leu Gly Ser Cys
210 215 220
Glu Lys Lys Met Asp Leu Ser Pro Leu Gly His Ser Gly Ser Gln Arg
225 230 235 240
Ala Ser Ala Leu Asp Leu Cys Leu Ser Leu Gly Asn Glu Ser Cys Gly
245 250 255
Leu His Asp Asn
260
<210> 13
<211> 477
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of artificial regernation booster protein 3
<400> 13
atggttagca agggggagga ggttattaaa gaattcccag gaagcacccg gtggaatcca 60
acggcggagc aactttcgct ccttcgggaa atttattaca ggaatgggct gagaactccc 120
accgctgacg agatcagaca gatttcatca aaattgtcgc ggtatggtaa gattgaaggg 180
aaaaatgtgt acaactggtt ccagaacaga agagctaggg aaaagagaaa acagcggttg 240
tcaaccatag gctgtggttc ggcggggtcg gccgcaggga gcggagagtt tcgccatgtg 300
aagaccctgc cccttttccc tttgcataat aatgaggatt caggttcgtt ggatctggag 360
ctggaattgg gttttgcctc tgctaccgca gccgcgactt ctggttcaca aagggcgtcg 420
gcattggatc tctgcctgtc ccttggaaat gagtcatgcg gtttgcatga caactga 477
<210> 14
<211> 158
<212> PRT
<213> Artificial Sequence
<220>
<223> artificial regernation booster protein 3
<400> 14
Met Val Ser Lys Gly Glu Glu Val Ile Lys Glu Phe Pro Gly Ser Thr
1 5 10 15
Arg Trp Asn Pro Thr Ala Glu Gln Leu Ser Leu Leu Arg Glu Ile Tyr
20 25 30
Tyr Arg Asn Gly Leu Arg Thr Pro Thr Ala Asp Glu Ile Arg Gln Ile
35 40 45
Ser Ser Lys Leu Ser Arg Tyr Gly Lys Ile Glu Gly Lys Asn Val Tyr
50 55 60
Asn Trp Phe Gln Asn Arg Arg Ala Arg Glu Lys Arg Lys Gln Arg Leu
65 70 75 80
Ser Thr Ile Gly Cys Gly Ser Ala Gly Ser Ala Ala Gly Ser Gly Glu
85 90 95
Phe Arg His Val Lys Thr Leu Pro Leu Phe Pro Leu His Asn Asn Glu
100 105 110
Asp Ser Gly Ser Leu Asp Leu Glu Leu Glu Leu Gly Phe Ala Ser Ala
115 120 125
Thr Ala Ala Ala Thr Ser Gly Ser Gln Arg Ala Ser Ala Leu Asp Leu
130 135 140
Cys Leu Ser Leu Gly Asn Glu Ser Cys Gly Leu His Asp Asn
145 150 155
<210> 15
<211> 477
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of artificial regernation booster protein 4
<400> 15
atggtctcga aaggagaaga ggtcatcaaa gaattttcag gcagcacccg ctggaatccg 60
actgcagaac aactttctct cctcaaagag ctttattaca gaaatgggat cagaacccct 120
tcagcggacc agatacgcca gataagcgcc cgcttgtcga ggtacggcaa aatcgagggg 180
aagaatgttt tttactggtt ccaaaaccat aaagcgaggg agcggcagaa gaaaagactg 240
agcactgtgg gttgtggatc ggcgggatcg gctgccgggt caggtgaatt tcgccatgtt 300
aaaacgcttc cactgtttcc cttgcataat aatgaagaca gcggatcact tgacctggag 360
ctggagttgg gcttcgcttc cgcgacggct gctgcaacat ccggctcgca gcgggcgtcg 420
gcgctggatt tgtgcctttc gctgggaaat gagagctgcg gtcttcatga caactga 477
<210> 16
<211> 158
<212> PRT
<213> Artificial Sequence
<220>
<223> artificial regernation booster protein 4
<400> 16
Met Val Ser Lys Gly Glu Glu Val Ile Lys Glu Phe Ser Gly Ser Thr
1 5 10 15
Arg Trp Asn Pro Thr Ala Glu Gln Leu Ser Leu Leu Lys Glu Leu Tyr
20 25 30
Tyr Arg Asn Gly Ile Arg Thr Pro Ser Ala Asp Gln Ile Arg Gln Ile
35 40 45
Ser Ala Arg Leu Ser Arg Tyr Gly Lys Ile Glu Gly Lys Asn Val Phe
50 55 60
Tyr Trp Phe Gln Asn His Lys Ala Arg Glu Arg Gln Lys Lys Arg Leu
65 70 75 80
Ser Thr Val Gly Cys Gly Ser Ala Gly Ser Ala Ala Gly Ser Gly Glu
85 90 95
Phe Arg His Val Lys Thr Leu Pro Leu Phe Pro Leu His Asn Asn Glu
100 105 110
Asp Ser Gly Ser Leu Asp Leu Glu Leu Glu Leu Gly Phe Ala Ser Ala
115 120 125
Thr Ala Ala Ala Thr Ser Gly Ser Gln Arg Ala Ser Ala Leu Asp Leu
130 135 140
Cys Leu Ser Leu Gly Asn Glu Ser Cys Gly Leu His Asp Asn
145 150 155
<210> 17
<211> 501
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of artificial regernation booster protein 5
<400> 17
atggtgtcta aaggggaaga ggtcataaaa gagttctcag gttctactcg ctggaatcca 60
actgcagaac aactctctct cctgaaggag ctctactatc ggaacggaat acgcacgcct 120
tcggcggacc aaataaggca gatttcagcc cgcctgtcta gatacgggaa aatcgaggga 180
aagaacgtgt tttattggtt ccaaaaccat aaggcaagag agcggcagaa aaaacggctc 240
agcacagtcg gctgtgggtc agccggttcg gcggctggct ctggtgaatt ccgccacgtc 300
aaaaccctgc ctctgttccc gctccacaat aatgtcgaga cacttccctt gttcccggag 360
gattccgggt ctctcgactt ggagctcgaa ttgggctttg cctctgccac tgcggcagcg 420
accagcggat cacagagagc ctccgcactg gatctgtgcc tgtcacttgg aaatgaatcg 480
tgcggattgc acgacaattg a 501
<210> 18
<211> 166
<212> PRT
<213> Artificial Sequence
<220>
<223> artificial regernation booster protein 5
<400> 18
Met Val Ser Lys Gly Glu Glu Val Ile Lys Glu Phe Ser Gly Ser Thr
1 5 10 15
Arg Trp Asn Pro Thr Ala Glu Gln Leu Ser Leu Leu Lys Glu Leu Tyr
20 25 30
Tyr Arg Asn Gly Ile Arg Thr Pro Ser Ala Asp Gln Ile Arg Gln Ile
35 40 45
Ser Ala Arg Leu Ser Arg Tyr Gly Lys Ile Glu Gly Lys Asn Val Phe
50 55 60
Tyr Trp Phe Gln Asn His Lys Ala Arg Glu Arg Gln Lys Lys Arg Leu
65 70 75 80
Ser Thr Val Gly Cys Gly Ser Ala Gly Ser Ala Ala Gly Ser Gly Glu
85 90 95
Phe Arg His Val Lys Thr Leu Pro Leu Phe Pro Leu His Asn Asn Val
100 105 110
Glu Thr Leu Pro Leu Phe Pro Glu Asp Ser Gly Ser Leu Asp Leu Glu
115 120 125
Leu Glu Leu Gly Phe Ala Ser Ala Thr Ala Ala Ala Thr Ser Gly Ser
130 135 140
Gln Arg Ala Ser Ala Leu Asp Leu Cys Leu Ser Leu Gly Asn Glu Ser
145 150 155 160
Cys Gly Leu His Asp Asn
165
<210> 19
<211> 771
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of artificial regernation booster protein 6
<400> 19
atggtgtcta aaggcgaaga ggtcattaag gagttcagcg gctctactag atggaatcca 60
acggccgagc aacttagcct gctcaaagaa ctctattatc gcaatgggat acgcacgccc 120
tctgcagatc agatacgcca aatctcagcg cgcctgtcca ggtatgggaa gattgagggt 180
aaaaacgtct tttactggtt ccaaaatcac aaagcgcgcg agagacagaa gaaaagactc 240
agcaccgttg gttgtgaccc tgctctcatc gagatgggga atgttgcttc cctggagttt 300
ggcacagaga gcgctttgga atcactctct tccggcccct caagcgaact cagagaggct 360
cccactcgga agttttacga gaaaaaaacc gtcggagaaa acagcacgat tatcaacccc 420
gtcgagcaga actgcaccct ctcatgcggc acgtcccagg agtttcagta cgccgtcgac 480
tcgaggagag tgatgaaggc gatggaggaa aagcaagcaa ctgacgacga gcccgacgga 540
aacaagtgga ctgaatccaa tcgccatgtt aaaacgttgc ctttgttccc gctccataac 600
aatgtggaga cgttgccact cttcccggag gactcgggat ctctggactt ggagctggaa 660
ctgggctttg cttcggctac cgcagcagcc acctcgggtt cccagagagc ctcggctctt 720
gatctctgcc tgtcactggg taacgaatct tgcggactgc acgataatta g 771
<210> 20
<211> 256
<212> PRT
<213> Artificial Sequence
<220>
<223> artificial regernation booster protein 6
<400> 20
Met Val Ser Lys Gly Glu Glu Val Ile Lys Glu Phe Ser Gly Ser Thr
1 5 10 15
Arg Trp Asn Pro Thr Ala Glu Gln Leu Ser Leu Leu Lys Glu Leu Tyr
20 25 30
Tyr Arg Asn Gly Ile Arg Thr Pro Ser Ala Asp Gln Ile Arg Gln Ile
35 40 45
Ser Ala Arg Leu Ser Arg Tyr Gly Lys Ile Glu Gly Lys Asn Val Phe
50 55 60
Tyr Trp Phe Gln Asn His Lys Ala Arg Glu Arg Gln Lys Lys Arg Leu
65 70 75 80
Ser Thr Val Gly Cys Asp Pro Ala Leu Ile Glu Met Gly Asn Val Ala
85 90 95
Ser Leu Glu Phe Gly Thr Glu Ser Ala Leu Glu Ser Leu Ser Ser Gly
100 105 110
Pro Ser Ser Glu Leu Arg Glu Ala Pro Thr Arg Lys Phe Tyr Glu Lys
115 120 125
Lys Thr Val Gly Glu Asn Ser Thr Ile Ile Asn Pro Val Glu Gln Asn
130 135 140
Cys Thr Leu Ser Cys Gly Thr Ser Gln Glu Phe Gln Tyr Ala Val Asp
145 150 155 160
Ser Arg Arg Val Met Lys Ala Met Glu Glu Lys Gln Ala Thr Asp Asp
165 170 175
Glu Pro Asp Gly Asn Lys Trp Thr Glu Ser Asn Arg His Val Lys Thr
180 185 190
Leu Pro Leu Phe Pro Leu His Asn Asn Val Glu Thr Leu Pro Leu Phe
195 200 205
Pro Glu Asp Ser Gly Ser Leu Asp Leu Glu Leu Glu Leu Gly Phe Ala
210 215 220
Ser Ala Thr Ala Ala Ala Thr Ser Gly Ser Gln Arg Ala Ser Ala Leu
225 230 235 240
Asp Leu Cys Leu Ser Leu Gly Asn Glu Ser Cys Gly Leu His Asp Asn
245 250 255
<210> 21
<211> 840
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of artificial regernation booster protein 7
<400> 21
atggtttcaa agggggagga ggtcataaaa gaattttctg gcagcactag atggaatccg 60
actgctgagc agttgtctct cctcaaagaa ctttactata ggaacggtat ccgcactccg 120
tccgctgatc aaattcgcca aatttcggcc agactgtcca gatacgggaa aattgaggga 180
aaaaatgttt tctactggtt ccagaatcat aaggcgcgcg aaaggcagaa aaagagactt 240
tcgaccgtcg gatgtgaccc tgcacttatc gaaatgggca atgtcgcgtc gttggagttt 300
ggcaccgaat ccgctctcga atctcttagc tcgggcccct catcggaact cagagaggcc 360
cctaccagaa aattttatga gaaaaagaca gtgggagaga actctacaat cattaacccc 420
gtcgagcaga actgcacatt gagctgtgga acatcgcagg agtttcaata cgcggtggat 480
tcgcgccgcg ttatgaaggc aatggaggag aagcaggcaa cagatgatga gccagacggc 540
aataaatgga ccgaatctaa cagacatgtc aagacactgc cacttttccc tctccacaac 600
aatgtcgaga cactccctct gtttcccgag gattcgggaa gcctggatct tgagctggaa 660
cttggctttg catcaggtgg atctgccggg tcagttgaca cgctgccatt gtttcctaga 720
gaggctgcgg gttctgggga atttgccact gcggctgcga ctagcggatc acagcgcgca 780
tcggcgctcg acctctgcct ttctttgggt aacgagtctt gcggcctgca tgacaactga 840
<210> 22
<211> 279
<212> PRT
<213> Artificial Sequence
<220>
<223> artificial regernation booster protein 7
<400> 22
Met Val Ser Lys Gly Glu Glu Val Ile Lys Glu Phe Ser Gly Ser Thr
1 5 10 15
Arg Trp Asn Pro Thr Ala Glu Gln Leu Ser Leu Leu Lys Glu Leu Tyr
20 25 30
Tyr Arg Asn Gly Ile Arg Thr Pro Ser Ala Asp Gln Ile Arg Gln Ile
35 40 45
Ser Ala Arg Leu Ser Arg Tyr Gly Lys Ile Glu Gly Lys Asn Val Phe
50 55 60
Tyr Trp Phe Gln Asn His Lys Ala Arg Glu Arg Gln Lys Lys Arg Leu
65 70 75 80
Ser Thr Val Gly Cys Asp Pro Ala Leu Ile Glu Met Gly Asn Val Ala
85 90 95
Ser Leu Glu Phe Gly Thr Glu Ser Ala Leu Glu Ser Leu Ser Ser Gly
100 105 110
Pro Ser Ser Glu Leu Arg Glu Ala Pro Thr Arg Lys Phe Tyr Glu Lys
115 120 125
Lys Thr Val Gly Glu Asn Ser Thr Ile Ile Asn Pro Val Glu Gln Asn
130 135 140
Cys Thr Leu Ser Cys Gly Thr Ser Gln Glu Phe Gln Tyr Ala Val Asp
145 150 155 160
Ser Arg Arg Val Met Lys Ala Met Glu Glu Lys Gln Ala Thr Asp Asp
165 170 175
Glu Pro Asp Gly Asn Lys Trp Thr Glu Ser Asn Arg His Val Lys Thr
180 185 190
Leu Pro Leu Phe Pro Leu His Asn Asn Val Glu Thr Leu Pro Leu Phe
195 200 205
Pro Glu Asp Ser Gly Ser Leu Asp Leu Glu Leu Glu Leu Gly Phe Ala
210 215 220
Ser Gly Gly Ser Ala Gly Ser Val Asp Thr Leu Pro Leu Phe Pro Arg
225 230 235 240
Glu Ala Ala Gly Ser Gly Glu Phe Ala Thr Ala Ala Ala Thr Ser Gly
245 250 255
Ser Gln Arg Ala Ser Ala Leu Asp Leu Cys Leu Ser Leu Gly Asn Glu
260 265 270
Ser Cys Gly Leu His Asp Asn
275
<210> 23
<211> 498
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of artificial regernation booster protein 8
<400> 23
atggtttcga agggggaaga agtcattaaa gagttttcag ggtctacgcg ctggaaccct 60
accgctgaac aagtcaaggt tttgagggaa atatattacc gcgcgggtct taggacgcca 120
tccgcggatc agatccgcca aatatcctca cagctgaaac ggtatggaaa ggtggaagga 180
aaaaacgtgt tttattggtt ccagaaccat aaagcgcggg aaagacaaaa aaagcggctc 240
agcaccgttg ggagcgctgg ctcagcagcg ggttcaggtg agttcaggca cgttaaaaca 300
ctgcccttgt tccctttgca caataatgaa gactcaggga gcggtttgga cttggatctg 360
aaccttgagt tgcgcctgtc ggcgactgct gctgccactt cgggctcaca gcgggcgtct 420
gccctcgatc tggacttgga gctcagactc ggtttcgcgc ttggtaatga gtcgtgtgga 480
ctgcacgaca actagtga 498
<210> 24
<211> 164
<212> PRT
<213> Artificial Sequence
<220>
<223> artificial regernation booster protein 8
<400> 24
Met Val Ser Lys Gly Glu Glu Val Ile Lys Glu Phe Ser Gly Ser Thr
1 5 10 15
Arg Trp Asn Pro Thr Ala Glu Gln Val Lys Val Leu Arg Glu Ile Tyr
20 25 30
Tyr Arg Ala Gly Leu Arg Thr Pro Ser Ala Asp Gln Ile Arg Gln Ile
35 40 45
Ser Ser Gln Leu Lys Arg Tyr Gly Lys Val Glu Gly Lys Asn Val Phe
50 55 60
Tyr Trp Phe Gln Asn His Lys Ala Arg Glu Arg Gln Lys Lys Arg Leu
65 70 75 80
Ser Thr Val Gly Ser Ala Gly Ser Ala Ala Gly Ser Gly Glu Phe Arg
85 90 95
His Val Lys Thr Leu Pro Leu Phe Pro Leu His Asn Asn Glu Asp Ser
100 105 110
Gly Ser Gly Leu Asp Leu Asp Leu Asn Leu Glu Leu Arg Leu Ser Ala
115 120 125
Thr Ala Ala Ala Thr Ser Gly Ser Gln Arg Ala Ser Ala Leu Asp Leu
130 135 140
Asp Leu Glu Leu Arg Leu Gly Phe Ala Leu Gly Asn Glu Ser Cys Gly
145 150 155 160
Leu His Asp Asn
<210> 25
<211> 38
<212> PRT
<213> Artificial Sequence
<220>
<223> indicator motif for GRF1
<220>
<221> VARIANT
<222> (18)..(18)
<223> X is A (alanine) or S (serine)
<220>
<221> VARIANT
<222> (19)..(19)
<223> X is R (arginine) or K (lysine)
<220>
<221> VARIANT
<222> (20)..(20)
<223> X is E (glutamic acid) or D (aspartic acid)
<220>
<221> VARIANT
<222> (22)..(22)
<223> X is A (alanine) or H (histidine)
<220>
<221> VARIANT
<222> (23)..(23)
<223> X is P (proline), Q (glutamine) or S (serine)
<220>
<221> VARIANT
<222> (30)..(30)
<223> X is K (lysine) or R (arginine)
<400> 25
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
1 5 10 15
Cys Xaa Xaa Xaa Ala Xaa Xaa Asp Ser Lys Tyr Cys Glu Xaa His Met
20 25 30
His Arg Gly Arg Asn Arg
35
<210> 26
<211> 38
<212> PRT
<213> Zea mays
<400> 26
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
1 5 10 15
Cys Ser Lys Glu Ala Ala Pro Asp Ser Lys Tyr Cys Glu Arg His Met
20 25 30
His Arg Gly Arg Asn Arg
35
<210> 27
<211> 38
<212> PRT
<213> Brassica napus
<400> 27
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
1 5 10 15
Cys Ser Arg Glu Ala His Pro Asp Ser Lys Tyr Cys Glu Lys His Met
20 25 30
His Arg Gly Arg Asn Arg
35
<210> 28
<211> 38
<212> PRT
<213> Brassica napus
<400> 28
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
1 5 10 15
Cys Ser Arg Glu Ala His Pro Asp Ser Lys Tyr Cys Glu Lys His Met
20 25 30
His Arg Gly Arg Asn Arg
35
<210> 29
<211> 38
<212> PRT
<213> Brassica napus
<400> 29
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
1 5 10 15
Cys Ser Arg Glu Ala His Pro Asp Ser Lys Tyr Cys Glu Lys His Met
20 25 30
His Arg Gly Arg Asn Arg
35
<210> 30
<211> 38
<212> PRT
<213> Solanum tuberosum
<400> 30
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
1 5 10 15
Cys Ser Lys Asp Ala His Pro Asp Ser Lys Tyr Cys Glu Arg His Met
20 25 30
His Arg Gly Arg Asn Arg
35
<210> 31
<211> 38
<212> PRT
<213> Solanum tuberosum
<400> 31
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
1 5 10 15
Cys Ser Lys Asp Ala His Pro Asp Ser Lys Tyr Cys Glu Arg His Met
20 25 30
His Arg Gly Arg Asn Arg
35
<210> 32
<211> 38
<212> PRT
<213> Secale cereale
<400> 32
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
1 5 10 15
Cys Ser Lys Glu Ala Ala Gln Asp Ser Lys Tyr Cys Glu Arg His Met
20 25 30
His Arg Gly Arg Asn Arg
35
<210> 33
<211> 38
<212> PRT
<213> Triticum aestivum
<400> 33
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
1 5 10 15
Cys Ala Lys Glu Ala Ala Ser Asp Ser Lys Tyr Cys Glu Arg His Met
20 25 30
His Arg Gly Arg Asn Arg
35
<210> 34
<211> 38
<212> PRT
<213> Sorghum bicolor
<400> 34
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
1 5 10 15
Cys Ser Lys Glu Ala Ala Pro Asp Ser Lys Tyr Cys Glu Arg His Met
20 25 30
His Arg Gly Arg Asn Arg
35
<210> 35
<211> 38
<212> PRT
<213> Sorghum bicolor
<400> 35
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
1 5 10 15
Cys Ser Lys Glu Ala Ala Pro Asp Ser Lys Tyr Cys Glu Arg His Met
20 25 30
His Arg Gly Arg Asn Arg
35
<210> 36
<211> 1080
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of Brassica napus GRF1 isoform
<400> 36
atgatgaacc ttagcggaac tagtgggaga acaatagaca ggcctccatt tacaccaaca 60
caatggcaag aaatggagca tcaagcccta atttacaagt acatggtctc aggtgttcct 120
gtcgctcctg agctcatctt ttcgattaga agaagcttgg actcttcctt ggtctctaga 180
ctcctccctc accaatccat tgggtgggga tgctaccaga tgggattagg gagaaaacca 240
gatccagaac caggaagatg cagaagaaca gatggtaaga agtggagatg ctcaagagaa 300
gcacacccag attcaaagta ttgtgaaaaa cacatgcaca gaggaagaaa ccgtgccaag 360
aaagctcttg atcagaatca gacaactgct ccttcaacat caccatctct ttctttcccc 420
aacaaagaca acccaataag ccccaccttg tcttcttctt cctctagttt cattgacact 480
tacagtaggt ttggtgttgg tagtaggaat agtagaggct acttcaacaa ccattctctt 540
gattatcctt atccttcttc ttgttcacct aaacaacaac aaactcttga gcatgcttct 600
gctttgtcac ttcatcaaaa cacatctact tctcagttca atgtcttagc ttctgaaact 660
gaccacaaag acttcagaca ttttcaaggg attggggaga gaactttctt tccagaagct 720
tctaaaagct ttcaagagtc tccataccat caccaagaag cgctagcaac agcattgaat 780
gatccctacc acggtagtag ttctgatcat catcacacat actcatcctc atcatctcga 840
catcgagatt atcaccaaca acaacaacat ttcgttttgg gggccgacga gttcaacaag 900
tccacaagaa ccgtcttctc aaacacatct agacaaggtc atcaggaaga ggagaaagat 960
tcaccgaaaa taaagaagtc tcttcatcat ttctttggtg aggactgggc acagatcaag 1020
aacgcttcag attcttggct tgacctttct tcccagtcaa gactcgacac tgctgaataa 1080
<210> 37
<211> 359
<212> PRT
<213> Brassica napus
<400> 37
Met Met Asn Leu Ser Gly Thr Ser Gly Arg Thr Ile Asp Arg Pro Pro
1 5 10 15
Phe Thr Pro Thr Gln Trp Gln Glu Met Glu His Gln Ala Leu Ile Tyr
20 25 30
Lys Tyr Met Val Ser Gly Val Pro Val Ala Pro Glu Leu Ile Phe Ser
35 40 45
Ile Arg Arg Ser Leu Asp Ser Ser Leu Val Ser Arg Leu Leu Pro His
50 55 60
Gln Ser Ile Gly Trp Gly Cys Tyr Gln Met Gly Leu Gly Arg Lys Pro
65 70 75 80
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
85 90 95
Cys Ser Arg Glu Ala His Pro Asp Ser Lys Tyr Cys Glu Lys His Met
100 105 110
His Arg Gly Arg Asn Arg Ala Lys Lys Ala Leu Asp Gln Asn Gln Thr
115 120 125
Thr Ala Pro Ser Thr Ser Pro Ser Leu Ser Phe Pro Asn Lys Asp Asn
130 135 140
Pro Ile Ser Pro Thr Leu Ser Ser Ser Ser Ser Ser Phe Ile Asp Thr
145 150 155 160
Tyr Ser Arg Phe Gly Val Gly Ser Arg Asn Ser Arg Gly Tyr Phe Asn
165 170 175
Asn His Ser Leu Asp Tyr Pro Tyr Pro Ser Ser Cys Ser Pro Lys Gln
180 185 190
Gln Gln Thr Leu Glu His Ala Ser Ala Leu Ser Leu His Gln Asn Thr
195 200 205
Ser Thr Ser Gln Phe Asn Val Leu Ala Ser Glu Thr Asp His Lys Asp
210 215 220
Phe Arg His Phe Gln Gly Ile Gly Glu Arg Thr Phe Phe Pro Glu Ala
225 230 235 240
Ser Lys Ser Phe Gln Glu Ser Pro Tyr His His Gln Glu Ala Leu Ala
245 250 255
Thr Ala Leu Asn Asp Pro Tyr His Gly Ser Ser Ser Asp His His His
260 265 270
Thr Tyr Ser Ser Ser Ser Ser Arg His Arg Asp Tyr His Gln Gln Gln
275 280 285
Gln His Phe Val Leu Gly Ala Asp Glu Phe Asn Lys Ser Thr Arg Thr
290 295 300
Val Phe Ser Asn Thr Ser Arg Gln Gly His Gln Glu Glu Glu Lys Asp
305 310 315 320
Ser Pro Lys Ile Lys Lys Ser Leu His His Phe Phe Gly Glu Asp Trp
325 330 335
Ala Gln Ile Lys Asn Ala Ser Asp Ser Trp Leu Asp Leu Ser Ser Gln
340 345 350
Ser Arg Leu Asp Thr Ala Glu
355
<210> 38
<211> 1080
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of Brassica napus GRF1 isoform
<400> 38
atgatgaacc ttagcggaac tagtgggaga acaatagaca ggcctccatt tacaccaaca 60
caatggcaag aaatggagca tcaagcccta atttacaagt acatggtctc aggtgttcct 120
gtcgctcctg agctcatctt ttcgattaga agaagcttgg actcttcctt ggtctctaga 180
ctcctccctc accaatccat tgggtgggga tgctaccaga tgggattagg gagaaaacca 240
gatccagaac caggaagatg cagaagaaca gatggtaaga agtggagatg ctcaagagaa 300
gcacacccag attcaaagta ttgtgaaaaa cacatgcaca gaggaagaaa ccgtgccaag 360
aaagctcttg atcagaatca gacaactgct ccttcaacat caccatctct ttctttcccc 420
aacaaagaca acccaataag ccccaccttg tcttcttctt cctctagttt cattgacact 480
tacagtaggt ttggtgttgg tagtaggaat agtagaggct acttcaacaa ccattctctt 540
gattatcctt atccttcttc ttgttcacct aaacaacaac aaactcttga gcatgcttct 600
gctttgtcac ttcatcaaaa cacatctact tctcagttca atgtcttagc ttctgaaact 660
gaccacaaag acttcagaca ttttcaaggg attggggaga gaactttctt tccagaagct 720
tctaaaagct ttcaagagtc tccataccat caccaagaag cgctagcaac agcattgaat 780
gatccctacc acggtagtag ttctgatcat catcacacat actcatcctc atcatctcga 840
catcgagatt atcaccaaca acaacaacat ttcgttttgg gggccgacga gttcaacaag 900
tccacaagaa ccgtcttctc aaacacatct agacaaggtc atcaggaaga ggagaaagat 960
tcaccgaaaa taaagaagtc tcttcatcat ttctttggtg aggactgggc acagatcaag 1020
aacgcttcag attcttggct tgacctttct tcccagtcaa gactcgacac tgctgaataa 1080
<210> 39
<211> 359
<212> PRT
<213> Brassica napus
<400> 39
Met Met Asn Leu Ser Gly Thr Ser Gly Arg Thr Ile Asp Arg Pro Pro
1 5 10 15
Phe Thr Pro Thr Gln Trp Gln Glu Met Glu His Gln Ala Leu Ile Tyr
20 25 30
Lys Tyr Met Val Ser Gly Val Pro Val Ala Pro Glu Leu Ile Phe Ser
35 40 45
Ile Arg Arg Ser Leu Asp Ser Ser Leu Val Ser Arg Leu Leu Pro His
50 55 60
Gln Ser Ile Gly Trp Gly Cys Tyr Gln Met Gly Leu Gly Arg Lys Pro
65 70 75 80
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
85 90 95
Cys Ser Arg Glu Ala His Pro Asp Ser Lys Tyr Cys Glu Lys His Met
100 105 110
His Arg Gly Arg Asn Arg Ala Lys Lys Ala Leu Asp Gln Asn Gln Thr
115 120 125
Thr Ala Pro Ser Thr Ser Pro Ser Leu Ser Phe Pro Asn Lys Asp Asn
130 135 140
Pro Ile Ser Pro Thr Leu Ser Ser Ser Ser Ser Ser Phe Ile Asp Thr
145 150 155 160
Tyr Ser Arg Phe Gly Val Gly Ser Arg Asn Ser Arg Gly Tyr Phe Asn
165 170 175
Asn His Ser Leu Asp Tyr Pro Tyr Pro Ser Ser Cys Ser Pro Lys Gln
180 185 190
Gln Gln Thr Leu Glu His Ala Ser Ala Leu Ser Leu His Gln Asn Thr
195 200 205
Ser Thr Ser Gln Phe Asn Val Leu Ala Ser Glu Thr Asp His Lys Asp
210 215 220
Phe Arg His Phe Gln Gly Ile Gly Glu Arg Thr Phe Phe Pro Glu Ala
225 230 235 240
Ser Lys Ser Phe Gln Glu Ser Pro Tyr His His Gln Glu Ala Leu Ala
245 250 255
Thr Ala Leu Asn Asp Pro Tyr His Gly Ser Ser Ser Asp His His His
260 265 270
Thr Tyr Ser Ser Ser Ser Ser Arg His Arg Asp Tyr His Gln Gln Gln
275 280 285
Gln His Phe Val Leu Gly Ala Asp Glu Phe Asn Lys Ser Thr Arg Thr
290 295 300
Val Phe Ser Asn Thr Ser Arg Gln Gly His Gln Glu Glu Glu Lys Asp
305 310 315 320
Ser Pro Lys Ile Lys Lys Ser Leu His His Phe Phe Gly Glu Asp Trp
325 330 335
Ala Gln Ile Lys Asn Ala Ser Asp Ser Trp Leu Asp Leu Ser Ser Gln
340 345 350
Ser Arg Leu Asp Thr Ala Glu
355
<210> 40
<211> 1092
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of Brassica napus GRF1 isoform
<400> 40
atgatgaacc taagtggaac tagtgggaga acaatagaga ggccaccatt tacaccaaca 60
caatggcaag aactagagca tcaagcccta atctacaagt acatggtctc aggtgttcct 120
gtccctcctg agctcatctt ctccattaga agaagcttgg actcttcctt ggtctctaga 180
ctcctccctc accaatccat tgggtgggga tgctaccaaa tgggattagg gagaaaacca 240
gatccagaac caggaagatg cagaagaaca gatggtaaga agtggagatg ctcaagagaa 300
gcacacccag attcaaagta ttgtgaaaaa cacatgcaca gaggaagaaa ccgtgccaag 360
aaagctcttg atcagaatca gacaactgct ccttcaacat caccatctct ttctttcccc 420
aacaaagaca acccaataag ccccaccttg tcttcttctt cctctagttt cattgacact 480
tacagtaggt ttggtgttgg tagtagtaat actagaggtt acttcaacaa ccattctctt 540
gattatcctt atccttcttc ttgttcacct aaacaacaac aaactcttga gcatgcttct 600
gctttgtcac ttcatcaaaa caaatcatct atttctcagt tcaatgtctt agcttctgaa 660
actgaccaca aagacttcag acattttcaa gggattgggg agagaacttt ttttccggaa 720
gcttctaaaa gctttcaaga gtctccatac catcaccaag aagcgctagc aacagcaatg 780
aacgatccct accatggtag tagttctgat catcatcaca catactcatc cccatcatca 840
tcatctcgac atcgagatca tcaccaacaa caacaacatt tcgttttggg ggctgacgag 900
tttaacaagc ccacaagaac cgtcttctca aacacatcta gacaaggtca tcaagaagag 960
gaggagaaag attcaccgaa aataaagaag tctcttcatc atttctttgg tgaggactgg 1020
gcacagatca agaacacttc agattcttgg cttgatcttt cttcccagtc aagactcgac 1080
actggtagct ga 1092
<210> 41
<211> 363
<212> PRT
<213> Brassica napus
<400> 41
Met Met Asn Leu Ser Gly Thr Ser Gly Arg Thr Ile Glu Arg Pro Pro
1 5 10 15
Phe Thr Pro Thr Gln Trp Gln Glu Leu Glu His Gln Ala Leu Ile Tyr
20 25 30
Lys Tyr Met Val Ser Gly Val Pro Val Pro Pro Glu Leu Ile Phe Ser
35 40 45
Ile Arg Arg Ser Leu Asp Ser Ser Leu Val Ser Arg Leu Leu Pro His
50 55 60
Gln Ser Ile Gly Trp Gly Cys Tyr Gln Met Gly Leu Gly Arg Lys Pro
65 70 75 80
Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg
85 90 95
Cys Ser Arg Glu Ala His Pro Asp Ser Lys Tyr Cys Glu Lys His Met
100 105 110
His Arg Gly Arg Asn Arg Ala Lys Lys Ala Leu Asp Gln Asn Gln Thr
115 120 125
Thr Ala Pro Ser Thr Ser Pro Ser Leu Ser Phe Pro Asn Lys Asp Asn
130 135 140
Pro Ile Ser Pro Thr Leu Ser Ser Ser Ser Ser Ser Phe Ile Asp Thr
145 150 155 160
Tyr Ser Arg Phe Gly Val Gly Ser Ser Asn Thr Arg Gly Tyr Phe Asn
165 170 175
Asn His Ser Leu Asp Tyr Pro Tyr Pro Ser Ser Cys Ser Pro Lys Gln
180 185 190
Gln Gln Thr Leu Glu His Ala Ser Ala Leu Ser Leu His Gln Asn Lys
195 200 205
Ser Ser Ile Ser Gln Phe Asn Val Leu Ala Ser Glu Thr Asp His Lys
210 215 220
Asp Phe Arg His Phe Gln Gly Ile Gly Glu Arg Thr Phe Phe Pro Glu
225 230 235 240
Ala Ser Lys Ser Phe Gln Glu Ser Pro Tyr His His Gln Glu Ala Leu
245 250 255
Ala Thr Ala Met Asn Asp Pro Tyr His Gly Ser Ser Ser Asp His His
260 265 270
His Thr Tyr Ser Ser Pro Ser Ser Ser Ser Arg His Arg Asp His His
275 280 285
Gln Gln Gln Gln His Phe Val Leu Gly Ala Asp Glu Phe Asn Lys Pro
290 295 300
Thr Arg Thr Val Phe Ser Asn Thr Ser Arg Gln Gly His Gln Glu Glu
305 310 315 320
Glu Glu Lys Asp Ser Pro Lys Ile Lys Lys Ser Leu His His Phe Phe
325 330 335
Gly Glu Asp Trp Ala Gln Ile Lys Asn Thr Ser Asp Ser Trp Leu Asp
340 345 350
Leu Ser Ser Gln Ser Arg Leu Asp Thr Gly Ser
355 360
<210> 42
<211> 1221
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of Sorghum bicolor GRF1 isoform
<400> 42
atggcgatgc cgtatgcctc tctttccccg gcaggcgccg accaccgctc ctccacggcc 60
acggcggcgt cgctcctccc cttctgccgc tccaccccgc tctccgcggg cggcggcggc 120
ggcctggggg aggacgccca gttgagctcg cggtggccgg ccgcgaggcc ggtggtgccg 180
ttcacgccgg cgcagtacga ggagctggag cagcaggcgc tcatatacaa gtacctggtg 240
gccggcgtgc ccgtcccgcc ggatctcgtg gttccaatcc gccgcggtct cgactccctc 300
gcaacccgct tctacggcca tcccacactt ggtgggtacg ggacgtacta cttaggcaag 360
aaactggatc cggagccggg gcggtgccgg cgtacggacg gcaagaagtg gcggtgctcc 420
aaggaggccg ccccagactc caagtactgc gagcgccaca tgcaccgcgg ccgcaaccgt 480
tcaagaaagc ctgtggaaac gcagctcgtg ccccagtccc aaccgcccgc caccgccgct 540
gccgtctccg ccgctccgcc cttggccttg gccgccgcca ccaccaccac caacggcagc 600
tgcttccaga atcactctct ttacccggcc attgcaggca gcaccggtgg aggtggcggg 660
gccagcaata tctctacccc gttctcctcg tcgatggggt cgtctcagct gcacatggac 720
aatgctgcca gctacgcagc tcttggtggt ggaactgcaa aggatctcag gtacaacgcc 780
tacggaataa gatctttggc ggaggagcac aaccagctga ttgcagaagc cattgactca 840
tcaatggaga accagtggcg cctcccgcca tcccaaacct cttcgtttcc gctctcgagc 900
tacccccagc ttggggcgct gagcaacctg ggtcagagca cagtcacctc gctgtcgaag 960
atggagcggc agcagccact ctccttccta gggaactccg agttcggggc catggaatcc 1020
gccgccaagc agcaggagaa ccagacgctg cggcccttct tcgacgagtg gcccaaggcg 1080
agggactcct ggccgggcct ctccgacgac aacgccgcaa gcctcgctcc gtcgttcccg 1140
gcgacccagc tgtcgatgtc cataccgatg gcgtcctcgg acttctccgt ggccagctcc 1200
cagtcgccca acgatgacta a 1221
<210> 43
<211> 406
<212> PRT
<213> Sorghum bicolor
<400> 43
Met Ala Met Pro Tyr Ala Ser Leu Ser Pro Ala Gly Ala Asp His Arg
1 5 10 15
Ser Ser Thr Ala Thr Ala Ala Ser Leu Leu Pro Phe Cys Arg Ser Thr
20 25 30
Pro Leu Ser Ala Gly Gly Gly Gly Gly Leu Gly Glu Asp Ala Gln Leu
35 40 45
Ser Ser Arg Trp Pro Ala Ala Arg Pro Val Val Pro Phe Thr Pro Ala
50 55 60
Gln Tyr Glu Glu Leu Glu Gln Gln Ala Leu Ile Tyr Lys Tyr Leu Val
65 70 75 80
Ala Gly Val Pro Val Pro Pro Asp Leu Val Val Pro Ile Arg Arg Gly
85 90 95
Leu Asp Ser Leu Ala Thr Arg Phe Tyr Gly His Pro Thr Leu Gly Gly
100 105 110
Tyr Gly Thr Tyr Tyr Leu Gly Lys Lys Leu Asp Pro Glu Pro Gly Arg
115 120 125
Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg Cys Ser Lys Glu Ala Ala
130 135 140
Pro Asp Ser Lys Tyr Cys Glu Arg His Met His Arg Gly Arg Asn Arg
145 150 155 160
Ser Arg Lys Pro Val Glu Thr Gln Leu Val Pro Gln Ser Gln Pro Pro
165 170 175
Ala Thr Ala Ala Ala Val Ser Ala Ala Pro Pro Leu Ala Leu Ala Ala
180 185 190
Ala Thr Thr Thr Thr Asn Gly Ser Cys Phe Gln Asn His Ser Leu Tyr
195 200 205
Pro Ala Ile Ala Gly Ser Thr Gly Gly Gly Gly Gly Ala Ser Asn Ile
210 215 220
Ser Thr Pro Phe Ser Ser Ser Met Gly Ser Ser Gln Leu His Met Asp
225 230 235 240
Asn Ala Ala Ser Tyr Ala Ala Leu Gly Gly Gly Thr Ala Lys Asp Leu
245 250 255
Arg Tyr Asn Ala Tyr Gly Ile Arg Ser Leu Ala Glu Glu His Asn Gln
260 265 270
Leu Ile Ala Glu Ala Ile Asp Ser Ser Met Glu Asn Gln Trp Arg Leu
275 280 285
Pro Pro Ser Gln Thr Ser Ser Phe Pro Leu Ser Ser Tyr Pro Gln Leu
290 295 300
Gly Ala Leu Ser Asn Leu Gly Gln Ser Thr Val Thr Ser Leu Ser Lys
305 310 315 320
Met Glu Arg Gln Gln Pro Leu Ser Phe Leu Gly Asn Ser Glu Phe Gly
325 330 335
Ala Met Glu Ser Ala Ala Lys Gln Gln Glu Asn Gln Thr Leu Arg Pro
340 345 350
Phe Phe Asp Glu Trp Pro Lys Ala Arg Asp Ser Trp Pro Gly Leu Ser
355 360 365
Asp Asp Asn Ala Ala Ser Leu Ala Pro Ser Phe Pro Ala Thr Gln Leu
370 375 380
Ser Met Ser Ile Pro Met Ala Ser Ser Asp Phe Ser Val Ala Ser Ser
385 390 395 400
Gln Ser Pro Asn Asp Asp
405
<210> 44
<211> 1314
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of Sorghum bicolor GRF1 isoform
<400> 44
atgcctctct ttccccggca ggcgccgacc accgctcctc cacggccacg gcggcgtcgc 60
tcctcccctt ctgccgctcc accccgctct ccgcgtaagt actccaccac ctccactcgc 120
cgtcgtcgtt tcgctcgtgg ctctcccatt gttgccgtct ggttcgcggt tgtgacgcgg 180
tggtatgtgt gctggtgcag gggcggcggc ggcggcctgg gggaggacgc ccagttgagc 240
tcgcggtggc cggccgcgag gccggtggtg ccgttcacgc cggcgcagta cgaggagctg 300
gagcagcagg cgctcatata caagtacctg gtggccggcg tgcccgtccc gccggatctc 360
gtggttccaa tccgccgcgg tctcgactcc ctcgcaaccc gcttctacgg ccatcccaca 420
cttggtgggt acgggacgta ctacttaggc aagaaactgg atccggagcc ggggcggtgc 480
cggcgtacgg acggcaagaa gtggcggtgc tccaaggagg ccgccccaga ctccaagtac 540
tgcgagcgcc acatgcaccg cggccgcaac cgttcaagaa agcctgtgga aacgcagctc 600
gtgccccagt cccaaccgcc cgccaccgcc gctgccgtct ccgccgctcc gcccttggcc 660
ttggccgccg ccaccaccac caccaacggc agctgcttcc agaatcactc tctttacccg 720
gccattgcag gcagcaccgg tggaggtggc ggggccagca atatctctac cccgttctcc 780
tcgtcgatgg ggtcgtctca gctgcacatg gacaatgctg ccagctacgc agctcttggt 840
ggtggaactg caaaggatct caggtacaac gcctacggaa taagatcttt ggcggaggag 900
cacaaccagc tgattgcaga agccattgac tcatcaatgg agaaccagtg gcgcctcccg 960
ccatcccaaa cctcttcgtt tccgctctcg agctaccccc agcttggggc gctgagcaac 1020
ctgggtcaga gcacagtcac ctcgctgtcg aagatggagc ggcagcagcc actctccttc 1080
ctagggaact ccgagttcgg ggccatggaa tccgccgcca agcagcagga gaaccagacg 1140
ctgcggccct tcttcgacga gtggcccaag gcgagggact cctggccggg cctctccgac 1200
gacaacgccg caagcctcgc tccgtcgttc ccggcgaccc agctgtcgat gtccataccg 1260
atggcgtcct cggacttctc cgtggccagc tcccagtcgc ccaacgatga ctaa 1314
<210> 45
<211> 437
<212> PRT
<213> Sorghum bicolor
<400> 45
Met Pro Leu Phe Pro Arg Gln Ala Pro Thr Thr Ala Pro Pro Arg Pro
1 5 10 15
Arg Arg Arg Arg Ser Ser Pro Ser Ala Ala Pro Pro Arg Ser Pro Arg
20 25 30
Lys Tyr Ser Thr Thr Ser Thr Arg Arg Arg Arg Phe Ala Arg Gly Ser
35 40 45
Pro Ile Val Ala Val Trp Phe Ala Val Val Thr Arg Trp Tyr Val Cys
50 55 60
Trp Cys Arg Gly Gly Gly Gly Gly Leu Gly Glu Asp Ala Gln Leu Ser
65 70 75 80
Ser Arg Trp Pro Ala Ala Arg Pro Val Val Pro Phe Thr Pro Ala Gln
85 90 95
Tyr Glu Glu Leu Glu Gln Gln Ala Leu Ile Tyr Lys Tyr Leu Val Ala
100 105 110
Gly Val Pro Val Pro Pro Asp Leu Val Val Pro Ile Arg Arg Gly Leu
115 120 125
Asp Ser Leu Ala Thr Arg Phe Tyr Gly His Pro Thr Leu Gly Gly Tyr
130 135 140
Gly Thr Tyr Tyr Leu Gly Lys Lys Leu Asp Pro Glu Pro Gly Arg Cys
145 150 155 160
Arg Arg Thr Asp Gly Lys Lys Trp Arg Cys Ser Lys Glu Ala Ala Pro
165 170 175
Asp Ser Lys Tyr Cys Glu Arg His Met His Arg Gly Arg Asn Arg Ser
180 185 190
Arg Lys Pro Val Glu Thr Gln Leu Val Pro Gln Ser Gln Pro Pro Ala
195 200 205
Thr Ala Ala Ala Val Ser Ala Ala Pro Pro Leu Ala Leu Ala Ala Ala
210 215 220
Thr Thr Thr Thr Asn Gly Ser Cys Phe Gln Asn His Ser Leu Tyr Pro
225 230 235 240
Ala Ile Ala Gly Ser Thr Gly Gly Gly Gly Gly Ala Ser Asn Ile Ser
245 250 255
Thr Pro Phe Ser Ser Ser Met Gly Ser Ser Gln Leu His Met Asp Asn
260 265 270
Ala Ala Ser Tyr Ala Ala Leu Gly Gly Gly Thr Ala Lys Asp Leu Arg
275 280 285
Tyr Asn Ala Tyr Gly Ile Arg Ser Leu Ala Glu Glu His Asn Gln Leu
290 295 300
Ile Ala Glu Ala Ile Asp Ser Ser Met Glu Asn Gln Trp Arg Leu Pro
305 310 315 320
Pro Ser Gln Thr Ser Ser Phe Pro Leu Ser Ser Tyr Pro Gln Leu Gly
325 330 335
Ala Leu Ser Asn Leu Gly Gln Ser Thr Val Thr Ser Leu Ser Lys Met
340 345 350
Glu Arg Gln Gln Pro Leu Ser Phe Leu Gly Asn Ser Glu Phe Gly Ala
355 360 365
Met Glu Ser Ala Ala Lys Gln Gln Glu Asn Gln Thr Leu Arg Pro Phe
370 375 380
Phe Asp Glu Trp Pro Lys Ala Arg Asp Ser Trp Pro Gly Leu Ser Asp
385 390 395 400
Asp Asn Ala Ala Ser Leu Ala Pro Ser Phe Pro Ala Thr Gln Leu Ser
405 410 415
Met Ser Ile Pro Met Ala Ser Ser Asp Phe Ser Val Ala Ser Ser Gln
420 425 430
Ser Pro Asn Asp Asp
435
<210> 46
<211> 747
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of Triticum aestivum GRF1 isoform
<400> 46
atggcgatgc cgtatgcctc tctttccccg gcaggcgacc gccgctcctc cccggccgcc 60
accgccaccg cctccctcct ccccttctgc cgctcctccc ccttctccgc cggcggcaat 120
ggcggcatgg gggaggaggc gccgatggac gggaggtgga tggcgaggcc ggtgcccttc 180
acggcggcgc agtacgagga gctggagcac caggcgctca tatacaagta cctggtggcc 240
ggcgtgcccg tcccgccgga tctcgtgctc cccatccgcc gcggcatcga gtccctcgcc 300
gcccgcttct accacaaccc cctcgccatc gggtacggat cgtacctggg caagaaggtg 360
gatccggagc cgggccggtg ccggcgcacg gacggcaaga agtggcggtg cgccaaggag 420
gccgcctccg actccaagta ctgcgagcgc cacatgcacc gcggccgcaa ccgttcaaga 480
aagcctgtgg aaacgcagct cgtgccccac tcccagccgc cggccgcctc cgccgtgccg 540
cccctcgcca ccggcttcca cggccactcc ctctaccccg ccgtcggcgg cggcaccaac 600
ggtggtggag gcggggggaa caacggcatg tccatgcccg gcacgttctc ctccgcgctg 660
gggccgcctc agcagcacat gggcaacaat gccgcctctc cctacgcggc tctcggcggc 720
gccggaacat gcaaagattt caggtga 747
<210> 47
<211> 248
<212> PRT
<213> Triticum aestivum
<400> 47
Met Ala Met Pro Tyr Ala Ser Leu Ser Pro Ala Gly Asp Arg Arg Ser
1 5 10 15
Ser Pro Ala Ala Thr Ala Thr Ala Ser Leu Leu Pro Phe Cys Arg Ser
20 25 30
Ser Pro Phe Ser Ala Gly Gly Asn Gly Gly Met Gly Glu Glu Ala Pro
35 40 45
Met Asp Gly Arg Trp Met Ala Arg Pro Val Pro Phe Thr Ala Ala Gln
50 55 60
Tyr Glu Glu Leu Glu His Gln Ala Leu Ile Tyr Lys Tyr Leu Val Ala
65 70 75 80
Gly Val Pro Val Pro Pro Asp Leu Val Leu Pro Ile Arg Arg Gly Ile
85 90 95
Glu Ser Leu Ala Ala Arg Phe Tyr His Asn Pro Leu Ala Ile Gly Tyr
100 105 110
Gly Ser Tyr Leu Gly Lys Lys Val Asp Pro Glu Pro Gly Arg Cys Arg
115 120 125
Arg Thr Asp Gly Lys Lys Trp Arg Cys Ala Lys Glu Ala Ala Ser Asp
130 135 140
Ser Lys Tyr Cys Glu Arg His Met His Arg Gly Arg Asn Arg Ser Arg
145 150 155 160
Lys Pro Val Glu Thr Gln Leu Val Pro His Ser Gln Pro Pro Ala Ala
165 170 175
Ser Ala Val Pro Pro Leu Ala Thr Gly Phe His Gly His Ser Leu Tyr
180 185 190
Pro Ala Val Gly Gly Gly Thr Asn Gly Gly Gly Gly Gly Gly Asn Asn
195 200 205
Gly Met Ser Met Pro Gly Thr Phe Ser Ser Ala Leu Gly Pro Pro Gln
210 215 220
Gln His Met Gly Asn Asn Ala Ala Ser Pro Tyr Ala Ala Leu Gly Gly
225 230 235 240
Ala Gly Thr Cys Lys Asp Phe Arg
245
<210> 48
<400> 48
000
<210> 49
<211> 385
<212> PRT
<213> Secale cereale
<400> 49
Met Ala Met Pro Phe Ala Ser Leu Ser Pro Ala Ala Asp His His Arg
1 5 10 15
Ser Ser Pro Ile Phe Pro Phe Cys Arg Ser Ser Pro Leu Tyr Ser Val
20 25 30
Gly Gly Gly Thr Ala His Gln Gln Gln Gln Gln His Thr Met Ser Gly
35 40 45
Ala Arg Trp Ala Ala Arg Pro Ala Pro Phe Thr Ala Ala Gln Tyr Glu
50 55 60
Glu Leu Glu Gln Gln Ala Leu Ile Tyr Lys Tyr Leu Val Ala Gly Val
65 70 75 80
Pro Val Pro Pro Asp Leu Leu Leu Pro Ile Arg Arg Gly Phe Asp Ser
85 90 95
Leu Ala Ser Arg Phe Tyr His His His Ala Leu Gly Tyr Gly Ser Tyr
100 105 110
Phe Gly Lys Lys Leu Asp Pro Glu Pro Gly Arg Cys Arg Arg Thr Asp
115 120 125
Gly Lys Lys Trp Arg Cys Ser Lys Glu Ala Ala Gln Asp Ser Lys Tyr
130 135 140
Cys Glu Arg His Met His Arg Gly Arg Asn Arg Ser Arg Lys Pro Val
145 150 155 160
Glu Thr Gln Leu Val Ala Ala Pro His Ser His Ser Asn Gln Leu Gln
165 170 175
Gln Gln Ala Ala Ala Ala Phe His Gly His Ser Pro Tyr Pro Ala Ile
180 185 190
Ala Ser Gly Gly Gly Gly Gly Gly Ala Ala Gly Ser Phe Ala Leu Gly
195 200 205
Ser Ala Gln Leu His Met Asp Asn Ala Ala Ala Pro Tyr Ala Thr Ala
210 215 220
Gly Ala Ala Gly Asn Lys Asp Phe Arg Tyr Ser Ala Tyr Gly Phe Arg
225 230 235 240
Thr Ser Ala Met Glu Glu His Asn Gln Phe Ile Thr Ala Ala Met Asp
245 250 255
Ser Ala Met Glu Asn Tyr Ser Trp Arg Leu Met Pro Ala Gln Asn Ser
260 265 270
Ala Phe Ser Leu Ser Ser Tyr Pro Met Leu Gly Thr Leu Gly Asp Leu
275 280 285
Asp Gln Ser Thr Ile Cys Ser Leu Ala Lys Thr Glu Arg Glu Pro Leu
290 295 300
Ser Ser Ser Gly Gly Gly Gly Gly Phe Glu Asp Asp Glu Ser Val Val
305 310 315 320
Lys Gln Glu Asn Gln Thr Leu Arg Pro Phe Phe Asp Glu Trp Pro Lys
325 330 335
Asp Arg Asp Ser Trp Pro Glu Leu Gln Glu His Asp Ala Asn Ser Asn
340 345 350
Ala Phe Ser Ala Thr Lys Leu Ser Ile Ser Ile Pro Val Thr Ser Ser
355 360 365
Asp Phe Ser Thr Thr Ala Gly Ser Arg Ser Pro Asn Gly Ile Tyr Ser
370 375 380
Arg
385
<210> 50
<211> 1059
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of Solanum tuberosum GRF1 isoform
<400> 50
atgagtggga cctcaacgac atcagtggtg ggggtggggt tggtgggagg tggaggggag 60
gtggggtata atgattatgg attccggcca ccgtttacag cggtgcagtg gcaggaattg 120
gagcatcaag ctatgattta taagtattta gtagctgggc tacctgtgcc accggaccta 180
gtagtaccta ttcgccggag ttttgatgct atctcagcca ggttcttcaa tcatcctagc 240
ttgggttatt gttcctatta tgggaagaag tttgaccccg agccaggaag atgtagaagg 300
acagatggaa agaagtggag gtgctccaaa gatgcacatc ctgactccaa atattgtgaa 360
cgacacatgc atcgaggccg caaccgttca agaaagcctg tggaatctca aactacttcc 420
cagtccttgt cgacaagtat atcacacact gctactggga gcagcaatag aagtggaagt 480
ttccaaagca atagcagtgg aagcttccaa aatatgccat tatattccgt tgctaattca 540
gatggaataa gttacggaag caccaccacg aaactgcaga tggagcctgc ctcctatggg 600
ataaataata aggagtatag gtacggaatg gctcctgatg cggatgacca caatttcttg 660
ccagtagctt ccgcaagtgt gagaggacta ggaggtacgg gttctaacac agacaacatg 720
tggcgtctga tgccatcaca agttccgtca aaacccaatc tgaaaaatga ttcccagctg 780
ttgggtagtt cacctaatcc gtttgagcct gtgatcgaat caaaacagca accccaacat 840
tgcttcttca gcagtgacat agattcacct ggtacagtaa agcaggagac acagcatcca 900
atgcgttcat tctttgacga gtggcctaca tccaaagaat catggtccaa tcttgatgag 960
ggatctggca aaaataattt ctccaccact cagctgtcca tatccattcc gaatgctcct 1020
tctggattct cttcaaggag tgcttcttcc cctaaatga 1059
<210> 51
<211> 352
<212> PRT
<213> Solanum tuberosum
<400> 51
Met Ser Gly Thr Ser Thr Thr Ser Val Val Gly Val Gly Leu Val Gly
1 5 10 15
Gly Gly Gly Glu Val Gly Tyr Asn Asp Tyr Gly Phe Arg Pro Pro Phe
20 25 30
Thr Ala Val Gln Trp Gln Glu Leu Glu His Gln Ala Met Ile Tyr Lys
35 40 45
Tyr Leu Val Ala Gly Leu Pro Val Pro Pro Asp Leu Val Val Pro Ile
50 55 60
Arg Arg Ser Phe Asp Ala Ile Ser Ala Arg Phe Phe Asn His Pro Ser
65 70 75 80
Leu Gly Tyr Cys Ser Tyr Tyr Gly Lys Lys Phe Asp Pro Glu Pro Gly
85 90 95
Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg Cys Ser Lys Asp Ala
100 105 110
His Pro Asp Ser Lys Tyr Cys Glu Arg His Met His Arg Gly Arg Asn
115 120 125
Arg Ser Arg Lys Pro Val Glu Ser Gln Thr Thr Ser Gln Ser Leu Ser
130 135 140
Thr Ser Ile Ser His Thr Ala Thr Gly Ser Ser Asn Arg Ser Gly Ser
145 150 155 160
Phe Gln Ser Asn Ser Ser Gly Ser Phe Gln Asn Met Pro Leu Tyr Ser
165 170 175
Val Ala Asn Ser Asp Gly Ile Ser Tyr Gly Ser Thr Thr Thr Lys Leu
180 185 190
Gln Met Glu Pro Ala Ser Tyr Gly Ile Asn Asn Lys Glu Tyr Arg Tyr
195 200 205
Gly Met Ala Pro Asp Ala Asp Asp His Asn Phe Leu Pro Val Ala Ser
210 215 220
Ala Ser Val Arg Gly Leu Gly Gly Thr Gly Ser Asn Thr Asp Asn Met
225 230 235 240
Trp Arg Leu Met Pro Ser Gln Val Pro Ser Lys Pro Asn Leu Lys Asn
245 250 255
Asp Ser Gln Leu Leu Gly Ser Ser Pro Asn Pro Phe Glu Pro Val Ile
260 265 270
Glu Ser Lys Gln Gln Pro Gln His Cys Phe Phe Ser Ser Asp Ile Asp
275 280 285
Ser Pro Gly Thr Val Lys Gln Glu Thr Gln His Pro Met Arg Ser Phe
290 295 300
Phe Asp Glu Trp Pro Thr Ser Lys Glu Ser Trp Ser Asn Leu Asp Glu
305 310 315 320
Gly Ser Gly Lys Asn Asn Phe Ser Thr Thr Gln Leu Ser Ile Ser Ile
325 330 335
Pro Asn Ala Pro Ser Gly Phe Ser Ser Arg Ser Ala Ser Ser Pro Lys
340 345 350
<210> 52
<211> 1059
<212> DNA
<213> Artificial Sequence
<220>
<223> cDNA of Solanum tuberosum GRF1 isoform
<400> 52
atgagtggga cctcaacgac atcagtggtg ggggtggggt tggtgggagg tggaggggag 60
gtggggtata atgattatgg attccggcca ccgtttacag cggtgcagtg gcaggaattg 120
gagcatcaag ctatgattta taagtattta gtagctgggc tacctgtgcc accggaccta 180
gtagtaccta ttcgccggag ttttgatgct atctcagcca ggttcttcaa tcatcctagc 240
ttgggttatt gttcctatta tgggaagaag tttgaccccg agccaggaag atgtagaagg 300
acagatggaa agaagtggag gtgctccaaa gatgcacatc ctgactccaa atattgtgaa 360
cgacacatgc atcgaggccg caaccgttca agaaagcctg tggaatctca aactacttcc 420
cagtccttgt cgacaagtat atcacacact gctactggga gcagcaatag aagtggaagt 480
ttccaaagca atagcagtgg aagcttccaa aatatgccat tatattccgt tgctaattca 540
gatggaataa gttacggaag caccaccacg aaactgcaga tggagcctgc ctcctatggg 600
ataaataata aggagtatag gtacggaatg gctcctgatg cggatgacca caatttcttg 660
ccagtagctt ccgcaagtgt gagaggacta ggaggtacgg gttctaacac agacaacatg 720
tggcgtctga tgccatcaca agttccgtca aaacccaatc tgaaaaatga ttcccagctg 780
ttgggtagtt cacctaatcc gtttgagcct gtgatcgaat caaaacagca accccaacat 840
tgcttcttca gcagtgacat agattcacct ggtacagtaa agcaggagac acagcatcca 900
atgcgttcat tctttgacga gtggcctaca tccaaagaat catggtccaa tcttgatgag 960
ggatctggca aaaataattt ctccaccact cagctgtcca tatccattcc gaatgctcct 1020
tctggattct cttcaaggag tgcttcttcc cctaaatga 1059
<210> 53
<211> 352
<212> PRT
<213> Solanum tuberosum
<400> 53
Met Ser Gly Thr Ser Thr Thr Ser Val Val Gly Val Gly Leu Val Gly
1 5 10 15
Gly Gly Gly Glu Val Gly Tyr Asn Asp Tyr Gly Phe Arg Pro Pro Phe
20 25 30
Thr Ala Val Gln Trp Gln Glu Leu Glu His Gln Ala Met Ile Tyr Lys
35 40 45
Tyr Leu Val Ala Gly Leu Pro Val Pro Pro Asp Leu Val Val Pro Ile
50 55 60
Arg Arg Ser Phe Asp Ala Ile Ser Ala Arg Phe Phe Asn His Pro Ser
65 70 75 80
Leu Gly Tyr Cys Ser Tyr Tyr Gly Lys Lys Phe Asp Pro Glu Pro Gly
85 90 95
Arg Cys Arg Arg Thr Asp Gly Lys Lys Trp Arg Cys Ser Lys Asp Ala
100 105 110
His Pro Asp Ser Lys Tyr Cys Glu Arg His Met His Arg Gly Arg Asn
115 120 125
Arg Ser Arg Lys Pro Val Glu Ser Gln Thr Thr Ser Gln Ser Leu Ser
130 135 140
Thr Ser Ile Ser His Thr Ala Thr Gly Ser Ser Asn Arg Ser Gly Ser
145 150 155 160
Phe Gln Ser Asn Ser Ser Gly Ser Phe Gln Asn Met Pro Leu Tyr Ser
165 170 175
Val Ala Asn Ser Asp Gly Ile Ser Tyr Gly Ser Thr Thr Thr Lys Leu
180 185 190
Gln Met Glu Pro Ala Ser Tyr Gly Ile Asn Asn Lys Glu Tyr Arg Tyr
195 200 205
Gly Met Ala Pro Asp Ala Asp Asp His Asn Phe Leu Pro Val Ala Ser
210 215 220
Ala Ser Val Arg Gly Leu Gly Gly Thr Gly Ser Asn Thr Asp Asn Met
225 230 235 240
Trp Arg Leu Met Pro Ser Gln Val Pro Ser Lys Pro Asn Leu Lys Asn
245 250 255
Asp Ser Gln Leu Leu Gly Ser Ser Pro Asn Pro Phe Glu Pro Val Ile
260 265 270
Glu Ser Lys Gln Gln Pro Gln His Cys Phe Phe Ser Ser Asp Ile Asp
275 280 285
Ser Pro Gly Thr Val Lys Gln Glu Thr Gln His Pro Met Arg Ser Phe
290 295 300
Phe Asp Glu Trp Pro Thr Ser Lys Glu Ser Trp Ser Asn Leu Asp Glu
305 310 315 320
Gly Ser Gly Lys Asn Asn Phe Ser Thr Thr Gln Leu Ser Ile Ser Ile
325 330 335
Pro Asn Ala Pro Ser Gly Phe Ser Ser Arg Ser Ala Ser Ser Pro Lys
340 345 350
Claims (15)
1.一种用于转化植物细胞的方法，包括以下步骤：
(a1)平行或依次向植物细胞中引入
i.至少一个感兴趣的核苷酸序列；以及
ii.包含编码GRF1多肽的多核苷酸的表达盒，编码GRF1多肽的mRNA，或GRF1多肽；或
(a2)向植物细胞中引入至少一个感兴趣的核苷酸序列；并且
在所述植物细胞中平行或依次诱导编码GRF1多肽的内源基因的增强的表达水平；以及
(b)任选地，在以下条件下培育(a1)或(a2)的植物细胞或衍生自(a1)或(a2)的植物细胞的植物细胞：其中在所述植物细胞中，所述GRF1多肽从所述表达盒表达，GRF1多肽从引入的mRNA翻译，GRF1多肽从所述内源基因增强表达，或存在GRF1多肽。
2.一种用于修饰植物细胞的基因组的方法，包括以下步骤：
(a1)向植物细胞中引入包含编码GRF1多肽的多核苷酸的表达盒，编码GRF1多肽的mRNA或GRF1多肽；或
(a2)在植物细胞中诱导编码GRF1多肽的内源基因的增强的表达水平；以及
(b)在以下条件下培育(a1)或(a2)的植物细胞或衍生自(a1)或(a2)的植物细胞的植物细胞：其中在所述植物细胞中，所述GRF1多肽从所述表达盒表达，GRF1多肽从引入的mRNA翻译，GRF1多肽从所述内源基因增强表达，或存在GRF1多肽；
(c)通过单链DNA断裂(SSB)或双链DNA断裂(DSB)诱导酶或碱基编辑酶以及任选地通过修复核酸分子修饰(b)的植物细胞的基因组，所述SSB或DSB诱导酶或所述碱基编辑酶优选识别所述细胞的基因组中的预定位点，
其中所述基因组在所述预定位点处的修饰选自
i.至少一个核苷酸的取代；
ii.至少一个核苷酸的缺失；
iii.至少一个核苷酸的插入；或
iv.i.-iii.的任何组合；以及
其中步骤(c)与步骤(a1)/(a2)和/或(b)同时进行，在步骤(a1)/(a2)之前进行，在步骤(a1)/(a2)与(b)之间进行或在步骤(b)之后进行。
3.一种产生转基因植物的方法，包括以下步骤：
(a)根据权利要求1所述的方法转化植物细胞，以及
(b)从(a)的植物细胞或从衍生自(a)的植物细胞的植物细胞再生包含至少一个细胞的植物，所述至少一个细胞包含作为转基因的至少一个感兴趣的核苷酸序列。
4.一种产生遗传修饰的植物的方法，包括以下步骤：
(a)根据权利要求2的方法修饰植物细胞的基因组，以及
(b)从(a)的植物细胞或从衍生自(a)的植物细胞的植物细胞再生在至少一个细胞中包含所述基因组的修饰的植物。
5.一种产生单倍体植物胚的方法，包括以下步骤：
(a1)将包含编码GRF1多肽的多核苷酸的表达盒、编码GRF1多肽的mRNA或GRF1多肽引入未成熟的雄性配子体或小孢子中；或
(a2)在未成熟的雄性配子体或小孢子中诱导编码GRF1多肽的内源基因的增强的表达水平；以及
(c)在以下条件下培育(a)的未成熟的雄性配子体或小孢子：其中在所述未成熟的雄性配子体或所述小孢子中，所述GRF1多肽从所述表达盒表达，GRF1多肽从引入的mRNA翻译，GRF1多肽从所述内源基因增强表达，或存在GRF1多肽；以及
(d)选择衍生自步骤(b)的所述未成熟的雄性配子体或所述小孢子的单倍体植物胚。
6.权利要求1至5中任一项的方法，其中所述GRF1多肽包含PFAM结构域PF08880以及PFAM结构域PF08879，优选地其中所述PFAM结构域PF08880在所述GRF1多肽的N末端处或附近发现至少90％覆盖率的匹配，并且所述PFAM结构域PF08879在位于所述GRF1多肽中的所述PFAM结构域PF08880的C末端处发现至少90％的匹配。
7.权利要求6的方法，其中两个匹配的氨基酸段均位于所述GRF1多肽的N末端一半。
8.权利要求1至7中任一项的方法，其中所述GRF1多肽包含的基序[D]-[P]-[E]-[P]-[G]-[R]-[C]-[R]-[R]-[T]-[D]-[G]-[K]-[K]-[W]-[R]-[C]-[AS]-[RK]-[ED]-[A]-[AH]-[PQS]-[D]-[S]-[K]-[Y]-[C]-[E]-[KR]-[H]-[M]-[H]-[R]-[G]-[R]-[N]-[R](SEQ ID NO:25)，最多有两个错配，其中优选地所述基序由氨基酸序列SEQ ID NO:26至SEQ ID NO:35中任一条组成，和/或其中优选地所述基序含有匹配PFAM结构域PF08879的氨基酸段的亚区。
9.权利要求1至8中任一项的方法，其中所述GRF1多肽包含：
(i)选自SEQ ID NO:2、37、39、41、43、45、47、49、51和53的氨基酸序列；或
(ii)包含如下序列的氨基酸序列，所述序列与选自SEQ ID NO:2、37、39、41、43、45、47、49、51和53的氨基酸序列至少70％相同。
10.权利要求1至9中任一项的方法，其中编码所述GRF1多肽的所述多核苷酸包含：
(i)选自SEQ ID NO:1、36、38、40、42、44、46、48、50和52的编码核苷酸序列；
(ii)包含如下序列的编码核苷酸序列，所述序列与选自SEQ ID NO:1、36、38、40、42、44、46、48、50和52的核苷酸序列至少70％相同；
(iii)在遗传密码的简并范围内编码由(i)或(ii)编码的所述多肽的核苷酸序列；
(iv)与(i)、(ii)或(iii)的核苷酸序列互补的核苷酸序列；或
(v)在严格条件下与(iv)的核苷酸序列杂交的核苷酸序列。
11.权利要求1至10中任一项的方法，其中将包含编码GRF1多肽的多核苷酸的表达盒引入植物细胞中导致其稳定整合到所述植物细胞的基因组中，或其中将包含编码GRF1多肽的多核苷酸的表达盒、编码GRF1多肽的mRNA或GRF1多肽引入植物细胞中或在植物细胞中诱导编码GRF1多肽的内源基因的增强的表达水平导致GRF1多肽在所述植物细胞中或在其后代细胞中的瞬时出现。
12.权利要求1至11中任一项的方法，其中编码所述GRF1多肽的所述多核苷酸与适于在植物细胞中表达所述GRF1多肽的至少一个调节序列可操作连接。
13.权利要求1至4中任一项的方法，其中步骤(a1)或(a2)的植物细胞是体组织、愈伤组织、分生组织或胚组织的细胞、原生质体、配子体、花粉、胚珠或小孢子。
14.一种通过权利要求3或4的方法获得或可获得的植物或其部分或其后代植物。
15.一种权利要求14的植物的植物细胞或种子，其中所述植物细胞或所述种子包含作为转基因的所述至少一个感兴趣的核苷酸序列或在基因组中包含所述修饰。
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| EP19183486.0A EP3757219A1 (en) | 2019-06-28 | 2019-06-28 | Enhanced plant regeneration and transformation by using grf1 booster gene |
| EP19183486.0 | 2019-06-28 | ||
| PCT/EP2020/068209 WO2020260682A1 (en) | 2019-06-28 | 2020-06-29 | Enhanced plant regeneration and transformation by using grf1 booster gene |
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| EP (2) | EP3757219A1 (zh) |
| CN (1) | CN114269933A (zh) |
| BR (1) | BR112021026002A2 (zh) |
| CA (1) | CA3144888A1 (zh) |
| WO (1) | WO2020260682A1 (zh) |
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| CN114672513A (zh) * | 2022-04-12 | 2022-06-28 | 北京大学现代农业研究院 | 一种基因编辑系统及其应用 |
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| CN116179590B (zh) * | 2022-12-28 | 2023-08-15 | 南京林业大学 | 一种春兰miR396基因在调控植物茎增粗中的应用 |
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| CN114672513A (zh) * | 2022-04-12 | 2022-06-28 | 北京大学现代农业研究院 | 一种基因编辑系统及其应用 |
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Also Published As
| Publication number | Publication date |
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| BR112021026002A2 (pt) | 2022-02-08 |
| EP3757219A1 (en) | 2020-12-30 |
| WO2020260682A1 (en) | 2020-12-30 |
| CA3144888A1 (en) | 2020-12-30 |
| US20220235363A1 (en) | 2022-07-28 |
| EP3990642A1 (en) | 2022-05-04 |
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