CN113874388A - 孤雌生殖基因 - Google Patents
孤雌生殖基因 Download PDFInfo
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Abstract
本发明提供了蒲公英属孤雌生殖基因的核苷酸序列和氨基酸序列及其(功能性)同源物、片段和变体，其提供孤雌生殖作为无融合生殖的一部分。还提供了孤雌生殖的植物及其制备方法，以及分子标记及其使用方法。
Description
发明领域
本发明涉及生物技术领域，尤其涉及包括植物育种的植物生物技术。本发明特别涉及与例如无融合生殖和单倍体诱导相关且有用的基因的鉴定和用途。本发明特别涉及与孤雌生殖相关的基因，及编码的蛋白质，和两者的片段。本发明还涉及在植物和作物中抑制和/或诱导孤雌生殖的方法、基因和/或蛋白质或其片段用于无融合生殖(特别是与无融合生殖基因组合)的用途，或用于产生单倍体植物的用途，其中染色体可以加倍以产生双单倍体。
背景技术
无融合生殖(apomixis)(也称为不完全无配生殖(agamospermy))是通过种子进行的无性植物繁殖。据报道，在大约400种开花植物中发现了无融合生殖(Bicknell andKoltunow,2004)。开花植物中的无融合生殖以两种形式发生：
(1)配子体无融合生殖，其中胚由未减数的、未受精的卵细胞通过孤雌生殖产生；
(2)孢子体无融合生殖，其中胚在体细胞上(somatically)由孢子体细胞产生。
配子体无融合生殖体的实例是蒲公英属(Taraxacum sp.)、山柳菊属(Hieraciumsp.)、早熟禾属(Poa pratensis)和鸭茅状磨擦禾(Tripsacum dactyloides)。孢子体无融合生殖的实例是柑橘属(Citrus sp.)和倒捻子(Garcinia mangostana)。配子体无融合生殖涉及两个发育过程：
(1)避免减数分裂重组和减少(不完全减数分裂)；以及
(2)卵细胞发育成胚，无受精(孤雌生殖)。
无融合生殖产生的种子在遗传上与亲本植物相同。长期以来，已经认识到无融合生殖在植物育种中非常有用(Asker,1979；Hermsen,1980；Asker and Jerling,1990；Vielle-Calzada等,1995)。将无融合生殖引入作物的最明显的优点是杂种优势F1杂种的纯一传代(true breeding)。在大多数作物中，F1杂种是表现最好的品种。然而，在有性作物中，F1杂种必须通过近交纯合亲本的杂交再次产生每一代，因为F1杂种的自体受精通过F2后代植物基因组中的重组导致杂种优势的丧失。生产有性F1种子是一种反复、复杂且成本高昂的方法。相比之下，无融合生殖的F1杂种将不断地纯一传代。换言之，F1杂种的遗传固定和通过种子产生一致的后代植物成为可能。
通过无融合生殖的F1固定(fixation)是无融合生殖的一般性质的一种特殊情况，即任何基因型，无论其遗传复杂性如何，只需一步就能纯一传代。这意味着无融合生殖可用于多基因定量性状的立即固定。应当指出的是，大多数产量性状是多基因的。无融合生殖可用于多个性状(例如各种抗性，几个转基因或多个数量性状基因座)的堆叠(或聚合)。在没有无融合生殖的情况下，为了固定这样一组性状，每个性状基因座必须单独成为纯合子，然后再进行组合。当性状中涉及的基因座数目增加时，通过杂交使这些性状基因座纯合变得耗时、逻辑上具有挑战性并因此成本高昂。此外，等位基因之间的特异性上位相互作用由于纯合性而丧失。通过无融合生殖，可以固定这种类型的非加性遗传变异。因此，无融合生殖(通过种子的无性繁殖)在植物育种、商业种子生产和农业中具有引起范式转变的潜力(vanDijk等2016,Van Dijk and Schauer 2016)。
除了即时固定任何基因型，无论其复杂性如何，无融合生殖还有其他重要的农业用途。由于减数分裂问题，有性种间杂种和同源多倍体经常不育。由于无融合生殖跳过减数分裂，利用无融合生殖可以解决种间杂种和同源多倍体的这些问题。由于无融合生殖阻止了雌性杂交，因此提出了无融合生殖与雄性不育相结合来遏制转基因，防止转基因作物的野生近缘种中的转基因渗入(Daniell,2002)。在昆虫授粉的作物(例如芸苔属植物)中，无融合生殖种子集不会受到传粉昆虫服务不足的限制。鉴于传粉蜂群的健康问题日益严重(瓦螨感染、非洲杀手蜂等)，这一点变得越来越重要。在块茎繁殖的作物(如马铃薯)中，无融合生殖将保持优良的基因型克隆，但降低或甚至消除病毒传播的当前风险以及清洁生产、遏制和认证方面的相关成本。无融合生殖种子的储存成本也远低于块茎或其它无性繁殖的植物部分的储存成本。在观赏植物中，无融合生殖可以替代劳动密集型和昂贵的组织培养繁殖。一般认为无融合生殖大大降低了栽培种发育和植物繁殖的成本。
遗憾的是，在任何主要作物中均不会发生无融合生殖。已有许多尝试在有性作物中引入无融合生殖。例如，无融合生殖基因的渗入、有性模型物种的突变、通过杂交从头产生无融合生殖，以及候选基因的克隆。迄今为止，通过广泛杂交将来自野生无融合生殖体的无融合生殖基因渗入作物物种尚未成功(例如，将来自鸭茅状磨擦禾的无融合生殖渗入玉米-Savidan,Y.,2001；Morgan等,1998；WO97/10704)。关于突变的有性模型物种，WO2007/066214描述了在拟南芥中使用称为Dyad的不完全减数分裂突变体。然而，Dyad是一种外显率非常低的隐性突变。在作物物种中，这种突变用途有限。通过两种有性生殖型之间的杂交从头产生无融合生殖并没有产生农业上感兴趣的无融合生殖体(US2004/0168216A1和US2005/0155111A1)。US2004/0148667中描述了通过转座子标记在玉米中克隆候选无融合生殖基因。延长基因(elongate gene)的直系同源物已被要求保护，其被认为诱导无融合生殖。然而，根据Barrell and Grossniklaus(2005)，延长基因跳过减数分裂II，因此不保持母本基因型，这使其有用性大大降低。
US2006/0179498中已经描述了所谓的反向育种将是无融合生殖的替代方法。然而，这是一种技术上复杂的体外实验室程序，而无融合生殖是一种由植物本身进行的体内程序。此外，对于反向育种，一旦亲本系被重建(双配子纯合子)，仍然必须进行杂交。
天然无融合生殖体中的无融合生殖通常具有遗传基础(由Ozias-Akins和VanDijk,2007综述)。因此，另一种方法可能是从天然无融合生殖物种中分离无融合生殖基因。然而，这并不是一件容易的任务，因为天然无融合生殖体通常具有多倍体基因组，并且在多倍体中定位克隆是非常困难的。其它复杂因素是抑制无融合生殖特异性染色体区域中的重组、重复序列和分离畸变是杂交。
发明内容
如本文所述，需要用于在作物中诱导无融合生殖的方法，其至少没有现有技术的一些限制。特别是，需要生产无融合生殖植物和无融合生殖种子的方法。还需要提供参与无融合生殖(特别是孤雌生殖)过程的基因和蛋白质，其适用于在作物中引入无融合生殖并可以基本上模拟无融合生殖途径。
本发明人现已经鉴定和分离了孤雌生殖基因座和基因，与孤雌生殖表型相关的等位基因(在本文中表示为孤雌生殖等位基因或Par等位基因)和与非孤雌生殖表型(在本文中表示为有性或非孤雌生殖等位基因或par等位基因)相关的等位基因，其遗传序列，即启动子或5'UTR序列，编码序列，3'UTR序列和编码的蛋白质序列。孤雌生殖可以直接引入有性植物，可能通过随机或定向诱变、通过转化或通过体细胞杂交。通过遗传修饰有性植物孤雌生殖基因座的有性等位基因，例如通过诱变、转基因或通过在特定位点引入双链断裂和同源重组进行插入，可以引入Par等位基因，并且植物和/或其后代可以变得能够使卵细胞发育成胚。
定义
如本文所用，术语“基因座”(复数：基因座(loci))是指染色体上发现例如基因或遗传标记的特定位置(或多个位置)或位点。例如，“孤雌生殖基因座”是指基因组中孤雌生殖基因所在的位置，促成孤雌生殖表型的等位基因，即(孤雌生殖等位基因或Par等位基因)和/或其有性对应物，即非孤雌生殖基因(非孤雌生殖等位基因或par等位基因)。“在孤雌生殖中是功能性的”基因、等位基因、蛋白质或核酸在本文中应理解为有助于孤雌生殖表型和/或将植物或植物细胞转化为卵细胞发育为胚的能力。
如本文所用，术语“等位基因”意指特定基因座处基因的一种或多种替代形式中的任一种。在生物体的二倍体和/或多倍体细胞中，给定基因的等位基因位于染色体上的特定位置或基因座，其中在该组同源染色体的每条染色体上存在一个等位基因。二倍体和/或多倍体或植物物种可以在特定基因座包含大量不同的等位基因。
如本文所用，术语“显性等位基因”是指一个基因的等位基因之间的关系，其中对一个等位基因(即显性等位基因)的表型的影响掩盖了相同基因座处的第二等位基因(即隐性等位基因)的贡献。对于常染色体(除性染色体外的任何染色体)上的基因，等位基因及其相关性状是常染色体显性或常染色体隐性的。显性是孟德尔遗传和经典遗传学的一个关键概念。例如，显性等位基因可以编码功能性蛋白质，而隐性等位基因则不编码。在一个实施方案中，本文教导的基因及其片段或变体是指孤雌生殖基因的显性等位基因。
如本文所用，术语“雌性子房”(复数：“子房(ovaries)”)是指其中形成孢子的外壳(enclosure)。它可以由单细胞组成或可以是多细胞的。所有植物、真菌和许多其它谱系在其生命周期中的某个时刻会形成子房。子房可通过有丝分裂或减数分裂产生孢子。通常，在每个子房内，大孢子母细胞的减数分裂产生四个单倍体大孢子。在裸子植物和被子植物中，这四个大孢子中只有一个在成熟时起作用，其他三个退化。所属的大孢子有丝分裂并发育成雌配子体(megagametophyte)，其最终产生一个卵细胞。
如本文所用，术语“雌配子”是指在正常(有性)环境下与另一(“雄性”)细胞在有性繁殖的生物体中受精(受孕)期间融合的细胞。在产生两种形态上不同类型的配子并且其中每个个体仅产生一种类型的物种中，雌性是产生较大类型的配子(称为胚珠(卵)或卵细胞)的任何个体。在植物中，雌性胚珠由花的子房产生。当成熟时，单倍体胚珠产生雌配子，然后准备受精。雄性细胞是(主要是单倍体)花粉并由花药产生。
术语“遗传标记”或“多态性标记”是指基因组DNA上可用于“标记”染色体上特定位置的区域。如果遗传标记与基因紧密连锁或在基因“上”，则它“标记”在其上发现该基因的DNA，并因此可用于(分子)标记测定以选择或对抗该基因的存在，例如在标记辅助育种/选择(MAS)方法中。遗传标记的实例是AFLP(扩增片段长度多态性，EP534858)，微卫星(microsatellite)，RFLP(限制性片段长度多态性)，STS(序列标记位点)，SNP(单核苷酸多态性)，SFP(单特征多态性；参见Borevitz等,2003)，SCAR(序列特征扩增区域)，CAPS标记(切割的扩增多态性序列)等。标记离基因越远，标记和基因之间发生重组(交叉)的可能性越大，由此失去连接(以及标记和基因的共分离)。遗传基因座之间的距离根据重组频率衡量，并以cM(厘摩；1cM是两个标记之间的减数分裂重组频率(1％))给出。由于基因组大小在物种之间差异很大，由1cM表示的实际物理距离(即两个标记之间的千碱基，kb)在物种之间也有很大的差异。
应当理解，当在本文中提及“连锁的(linked)”标记时，这也包括“在”基因本身上的标记。
“MAS”是指“标记辅助选择”，由此筛选植物中一种或多种遗传和/或表型标记的存在与否，以加速包含标记的DNA区(和任选地缺乏侧翼区)转移到(优良)育种品系中。
“分子标记测定”(或测试)是指(基于DNA的)测定，其(直接或间接)指示植物或植物部分中等位基因例如Par或par等位基因的存在与否。优选地，它允许确定特定等位基因在任何单个植物的孤雌生殖基因座是纯合的还是杂合的。例如，在一个实施方案中，使用PCR引物扩增与孤雌生殖基因座连接的核酸，酶消化扩增产物，并基于扩增产物的电泳分辨模式，可以确定哪个(些)等位基因存在于任何个体植物中，以及孤雌生殖基因座处等位基因的接合性(即每个基因座处的基因型)。实例是SCAR标记(序列特征扩增区域)、CAPS标记(切割的扩增多态性序列)和类似的标记测定。
如本文所用，术语“杂合的”是指当两个不同的等位基因位于特定的基因座，但分别位于细胞中相应的同源染色体组时存在的遗传条件。相反，如本文所用，术语“纯合的”是指当两个(或在多倍体的情况下为更多个)相同的等位基因位于特定基因座，但分别位于细胞中相应的同源染色体组时存在的遗传条件。
如本文使用，“变种”符合UPOV公约，并且是指在具有已知最低等级的单个植物分类单位内的植物分组，该分组可以通过特征的表达来定义，并且可以通过所述特征中的至少一种的表达与任何其他植物分组区分开来，并且就其用于无变化地(稳定地)繁殖的适合性而言被认为是一个单位。
术语“蛋白质”或“多肽”可互换使用，是指由氨基酸链组成的分子，而不涉及特定的作用模式、大小、三维结构或来源。因此，蛋白质的“片段”或“部分”仍可称为“蛋白质”。“分离的蛋白质”用于指不再处于其天然环境中的蛋白质，例如在体外或在重组细菌或植物宿主细胞中的蛋白质。
术语“基因”是指包含在细胞中转录成RNA分子(例如，被加工成mRNA的前mRNA)的区域(转录区)的DNA序列，其与合适的调节区(例如，启动子)可操作地连接。因此，基因可包含几个可操作地连接的序列，诸如启动子，包含例如参与翻译起始的序列的5'前导序列，(蛋白质)编码区(cDNA或基因组DNA)和包含例如转录终止位点的3'非翻译序列。
“嵌合基因”(或重组基因)是指在自然界中通常不存在于物种中的任何基因，特别是其中存在核苷酸序列的一个或多个部分的基因，其在自然界中彼此不相关。例如，启动子在本质上不与部分或全部转录区或另一调控区相关。术语“嵌合基因”应理解为包括其中启动子或转录调控序列与一个或多个编码序列或反义(有义链的反向互补序列)或反向重复序列(有义和反义，由此RNA转录物在转录时形成双链RNA)可操作地连接的表达构建体。
“3’UTR”或“3'非翻译序列”(通常也称为3'非翻译区或3'末端)是指在基因编码序列下游发现的核苷酸序列，其包含例如转录终止位点和(在大多数但不是所有真核mRNA中)聚腺苷酸化信号(诸如例如AAUAAA或其变体)。转录终止后，mRNA转录物可在聚腺苷酸化信号的下游被切割，并可添加poly(A)尾，其参与mRNA向细胞质的转运(其中发生翻译)。
“5'UTR”或“前导序列”或“5'非翻译区”是mRNA转录物的区域，以及相应DNA，位于mRNA转录开始的+1位置和编码区的翻译起始密码子之间(通常是mRNA上的AUG或DNA上的ATG)。5'UTR通常含有对翻译、mRNA稳定性和/或周转和其它调控元件重要的位点。
“基因的表达”是指这样的过程，其中与适当的调控区域(特别是启动子)可操作地连接的DNA区被转录为具有生物活性的RNA，即其能够被翻译成具有生物活性的蛋白质或肽(或活性肽片段)或其本身具有活性(例如在转录后基因沉默或RNAi中)。在某些实施方案中，活性蛋白质是指组成型活性的蛋白质。编码序列优选为有义方向，并编码所需的生物活性蛋白质或肽，或活性肽片段。在基因沉默方法中，DNA序列优选以反义DNA或反向重复DNA的形式存在，包括反义或有义和反义方向的靶基因的短序列。
“转录调控序列”在本文中定义为能够调节与转录调控序列可操作连接的(编码)序列的转录速率的核苷酸序列。因此，本文定义的转录调节序列将包含转录起始所必需的所有序列元件(启动子元件)，维持和调控转录所必需的所有序列元件，包括例如衰减子或增强子。尽管大部分涉及编码序列的上游(5')转录调控序列，但在编码序列的下游(3')发现的调控序列也包括在该定义中。
如本文所用，术语“启动子”是指起控制一个或多个基因转录的作用的核酸片段，其位于基因转录起始位点转录方向的上游，并且通过存在DNA依赖性RNA聚合酶的结合位点、转录起始位点和任何其它DNA序列(包括但不限于转录因子结合位点、阻遏蛋白质和激活蛋白质结合位点，以及本领域技术人员已知的直接或间接调控启动子转录量的任何其它核苷酸序列)来在结构上鉴定。任选地，术语“启动子”在本文中还包括5'UTR区域(例如，启动子在本文中可包括基因的翻译起始密码子上游(5')的一个或多个部分，因为此区域可在调控转录和/或翻译中起作用。“组成型”启动子是在大多数生理和发育条件下在大多数组织中具有活性的启动子。“诱导型”启动子是生理上(例如通过外部施用某些化合物)或发育调节的启动子。“组织特异性”启动子仅在特定类型的组织或细胞中具有活性。“在植物或植物细胞中具有活性的启动子”是指启动子在植物或植物细胞内驱动转录的一般能力。它不会对启动子的时空活性产生任何影响。
如本文所用，术语“可操作地连接”是指多核苷酸元件以功能关系连接。当核酸与另一核苷酸序列处于功能关系时，它是“可操作地连接的”。例如，如果启动子或转录调控序列影响编码序列的转录，则其可操作地连接到编码序列。可操作地连接是指被连接的DNA序列通常是连续的，并且当需要连接两个蛋白质编码区时，是连续的并且在阅读框中，以便产生“嵌合蛋白质”。“嵌合蛋白质”或“杂合蛋白质”是由各种蛋白质“结构域”(或基序)组成的蛋白质，其在自然界中没有发现，但其连接形成功能性蛋白质，其显示连接的结构域的功能性。嵌合蛋白质也可以是天然存在的两种或多种蛋白质的融合蛋白质。如本文所用，术语“结构域”是指具有特定结构或功能的蛋白质的任何部分或结构域，其可转移至另一蛋白质以提供具有至少该结构域的功能特性的新杂合蛋白质。
术语“靶向肽”指将蛋白质或蛋白质片段靶向细胞内细胞器诸如质体，优选叶绿体、线粒体或细胞外间隙或质外体(分泌信号肽)的氨基酸序列。编码靶向肽的核苷酸序列可与编码蛋白质或蛋白质片段的氨基末端(N末端)的核苷酸序列融合(框内)，或可用于替代天然靶向肽。
“核酸构建体”或“载体”在本文中应理解为是指使用重组DNA技术产生的人工核酸分子，其用于将外源DNA递送到宿主细胞中。载体骨架可以是例如二元或超二元载体(参见例如US 5591616，US 2002138879和WO95/06722)、共整合载体或T-DNA载体，如本领域已知的和本文别处描述的，基因或嵌合基因整合到其中，或者如果合适的转录调控序列已经存在，则仅所需的核苷酸序列(例如编码序列、反义或反向重复序列)整合到转录调控序列的下游。载体通常包含其它遗传元件以促进它们在分子克隆中的应用，诸如例如可选择的标记、多克隆位点等。
“重组宿主细胞”或“转化的细胞”或“转基因细胞”是指由于至少一种核酸分子而产生的新的单个细胞(或生物体)，尤其包含编码所需蛋白质或核苷酸序列的基因或嵌合基因，所述基因或嵌合基因在转录后产生反义RNA或反向重复RNA(或发夹RNA)以使已经引入所述细胞中的靶基因/基因家族沉默。“分离的核酸”用于指不再处于其天然环境中的核酸，例如在体外或在重组细菌或植物宿主细胞中的核酸。
“宿主细胞”是用转基因转化成为重组宿主细胞的原始细胞。宿主细胞优选为植物细胞或细菌细胞。重组宿主细胞可以含有作为染色体外(附加型)复制分子的核酸构建体，或更优选地，包含整合在宿主细胞的核或质体基因组中的基因或嵌合基因。
“重组植物”或“重组植物部分”或“转基因植物”是包含重组基因或嵌合基因的植物或植物部分(例如，种子或果实或叶子)，即使该基因可能不表达或不在所有细胞中表达。
“优良事件(elite event)”是一种重组植物，其已被选择在基因组中的一个位置包含重组基因，其导致植物具有良好表型和/或农艺学特征。可以对整合位点的侧翼DNA进行测序以表征整合位点并将该事件与在基因组的其它位置包含相同重组基因的其它转基因植物区分开来。
术语“可选择的标记”是本领域普通技术人员熟悉的术语，并且在本文中用于描述任何遗传实体，其在表达时可用于选择含有可选择的标记的一个或多个细胞。可选择的标记基因产物赋予例如抗生素抗性，或更优选地，除草剂抗性或另一种可选性状诸如表型性状(例如色素沉着的变化)或营养需求。术语“报告分子(reporter)”主要用于指可见标记，诸如绿色荧光蛋白质(GFP)、eGFP、荧光素酶、GUS等。
术语基因或蛋白质的“直系同源物(orthologue)”在本文中是指在另一物种中发现的同源基因或蛋白质，其具有与该基因或蛋白质相同的功能，但(通常)在序列上与含有该基因的物种分化时的时间点(即，通过物种形成从共同祖先进化而来)分化。因此，可以基于序列比较(例如，基于整个序列或特定结构域序列同一性百分比)和功能分析，可以在其它植物物种中鉴定Taxaracum孤雌生殖基因的直系同源物。
术语“同源的”和“异源的”是指核酸或氨基酸序列与其宿主细胞或生物体之间的关系，尤其是在转基因生物体的上下文中。因此，同源序列天然存在于宿主物种中(例如用莴苣基因转化的莴苣植物)，而异源序列不天然存在于宿主细胞中(例如用来自马铃薯植物的序列转化的莴苣植物)。根据上下文，术语“同源物”或“同源的”可替代地指来自共同祖先序列的后代的序列(例如，它们可以是直系同源物)。
“严格的杂交条件”可用于鉴定与给定核苷酸序列基本相同的核苷酸序列。严格的条件取决于序列，并且在不同情况下会有所不同。通常，严格的条件被选择为比特定序列在限定的离子强度和pH下的热熔点(Tm)低约5℃。Tm是50％的靶序列与完全匹配的探针杂交的温度(在限定的离子强度和pH)。通常将选择严格的条件，其中盐浓度在pH 7为约0.02摩尔，并且温度为至少60℃。降低盐浓度和/或提高温度可提高严格性。RNA-DNA杂交的严格的条件(使用例如100nt的探针的Northern印迹)是例如包括至少一次在0.2X SSC中在63℃洗涤20分钟的那些，或等同条件。DNA-DNA杂交的严格的条件(使用例如100nt的探针的Southern印迹)是例如包括在0.2X SSC中在至少50℃，通常约55℃的温度至少洗涤一次(通常2次)20分钟的那些，或等同条件。也参见Sambrook等(1989)和Sambrook and Russell(2001)。
“高严格性”条件可例如通过在65℃在水溶液中杂交来提供，该水溶液含有6x SSC(20x SSC含有3.0M NaCl、0.3M柠檬酸钠，pH 7.0)、5x Denhardt's(100X Denhardt's含有2％Ficoll、2％聚乙烯吡咯烷酮、2％牛血清白蛋白质)、0.5％十二烷基硫酸钠(SDS)和20μg/ml变性载体DNA(单链鱼精DNA，平均长度为120-3000个核苷酸)作为非特异性竞争物。杂交后，可以在几个步骤中进行高严格性洗涤，最后在杂交温度在0.2-0.1×SSC、0.1％SDS中洗涤(约30分钟)。
“中等严格性”是指等同于在上述溶液中但在约60-62℃杂交的条件。在这种情况，最后的洗涤在杂交温度在1×SSC、0.1％SDS中进行。
“低严格性”是指等同于在上述溶液中在约50-52℃杂交的条件。在这种情况，最后的洗涤在杂交温度在2×SSC、0.1％SDS中进行。也参见Sambrook等(1989)和Sambrook andRussell(2001)。
“序列同一性”和“序列相似性”可以通过使用全局或局部比对算法对两个肽或两个核苷酸序列来确定，这取决于两个序列的长度。优选使用全局比对算法(例如NeedlemanWunsch)比对相似长度的序列，其在整个长度上最佳地比对序列，而优选使用局部比对算法(例如Smith Waterman)比对基本上不同长度的序列。当序列(当通过例如程序GAP或BESTFIT使用默认参数进行最佳比对时)共享至少某一最小百分比的序列同一性(如本文所定义)时，则可将序列称为“基本上相同”或“基本上相似”。GAP使用Needleman和Wunsch全局比对算法在其整个长度(全长)上比对两个序列，使匹配数最大化并使缺口数最小化。当两个序列具有相似的长度时，全局比对适合用于确定序列同一性。通常，使用GAP默认参数，缺口产生罚分＝50(核苷酸)/8(蛋白质)和缺口延伸罚分＝3(核苷酸)/2(蛋白质)。对于核苷酸，使用的默认评分矩阵是nwsgapdna，而对于蛋白质，默认评分矩阵是Blosum62(Henikoff&Henikoff,1992,PNAS 89,915-919)。序列比对和百分比序列同一性的得分可以使用计算机程序，诸如GCG Wisconsin Package，10.3版，可从Accelrys Inc.(9685Scranton Road,San Diego,CA 92121-3752USA)获得，或使用开源软件，诸如EmbossWIN 2.10.0版中的程序“needle”(使用全局Needleman Wunsch算法)或“water”(使用局部Smith Waterman算法)，使用与上述GAP相同的参数，或使用默认设置(对于“needle”和“water”二者以及对于蛋白质和对于DNA比对二者，默认缺口开放罚分为10.0并且默认缺口延伸罚分为0.5；蛋白质和DNA默认评分矩阵是Blossum62和DNAFull)来确定。当序列具有基本上不同的总长度时，优选局部比对，诸如使用Smith Waterman算法的那些。
或者，可通过使用诸如FASTA、BLAST等算法对公共数据库进行检索来确定百分比相似性或同一性。因此，本发明的核酸和蛋白质序列可进一步用作“查询序列”以对公共数据库进行检索，从而例如鉴定其它家族成员或相关序列。这种检索可以使用Altschul,等(1990)J.Mol.Biol.215:403—10的BLASTn和BLASTx程序(2.0版)进行。BLAST核苷酸检索可以用NBLAST程序进行，得分＝100，字长＝12，以获得与本发明的氧化还原酶核酸分子同源的核苷酸序列。BLAST蛋白质检索可以用BLASTx程序进行，得分＝50，字长＝3，以获得与本发明蛋白质分子同源的氨基酸序列。为了获得用于比较目的的缺口比对，可以使用如Altschul等,(1997)Nucleic Acids Res.25(17):3389-3402中所述的Gapped BLAST。当使用BLAST和Gapped BLAST程序时，可以使用相应程序的默认参数(例如，BLASTx和BLASTn)。参见国家生物技术信息中心(National Center for Biotechnology Information)主页http://www.ncbi.nlm.nih.gov/。
如本文所用，术语“有性植物繁殖”是指一种发育途径，其中被称为“大孢子母细胞”的体细胞(例如二倍体)经历减数分裂以产生四个减少的大孢子。这些大孢子中的一个有丝分裂形成雌配子体(也称为胚囊)，其含有减数的卵细胞(即与母体相比染色体数目减少的细胞)和两个减数的极核。通过花粉粒的一个精子细胞受精卵细胞产生(例如二倍体)胚，而通过第二个精子细胞受精两个极核产生(例如三倍体)胚乳(称为双受精的过程)。
如本文所用，术语“大孢子母细胞(megaspore mother cell)”或“大孢子母细胞(megasporocyte)”是指通过减数(通常为减数分裂)产生大孢子以产生将发育成雌配子体的四个单倍体大孢子的细胞。在被子植物(也称为开花植物)中，大孢子母细胞产生大孢子，该大孢子通过两种不同的过程发育成雌配子体，该过程包括大孢子发生(在珠心或大孢子囊中形成大孢子)和大配子发生(大孢子发育成雌配子体)。
如本文所用，术语“无性植物繁殖”是在没有受精和没有配子融合的情况下实现植物繁殖的过程。无性繁殖产生新的个体，其在遗传上与亲本植物和彼此相同，除了当发生突变或体细胞重组时。植物具有两种主要类型的无性繁殖，包括营养繁殖(即，涉及原始植物的营养部分的出芽、分蘖等)和无融合生殖。
如本文所用，术语“无融合生殖”是指通过无性过程形成种子。无融合生殖的一种形式的特征在于：1)不完全减数分裂，其是指在子房中形成未减数的胚囊，以及2)孤雌生殖，其是指未减数的卵细胞发育成胚。数百种野生植物物种具有无融合生殖和无性繁殖的特征。不完全减数分裂是导致产生未减数卵细胞的过程，其具有与亲本植物的体细胞组织相同的染色体数目和相同或高度相似的基因型。未减数卵细胞可以来自未减数大孢子(二倍体孢子生殖(diplospory))或来自体细胞起始细胞(无孢子生殖)。在二倍体孢子生殖的情况下，大孢子发生被有丝分裂或被修饰的减数分裂代替。经修饰的减数分裂优选是无重组的第一分裂恢复类型。或者，修饰的减数分裂可以是第二分裂重建类型。在一个优选的实施方案中，不完全减数分裂是影响第一次减数分裂的二倍体孢子类型。已知无融合生殖以不同的形式发生，包括至少两种称为配子体无融合生殖和孢子体无融合生殖融合生殖(也称为不定胚)。发生配子体无融合生殖的植物的实例包括蒲公英属、山柳菊属、早熟禾属和鸭茅状磨擦禾等。其中发生孢子体无融合生殖的植物的实例包括柑橘属和倒捻子等。
如本文所用，术语“二倍体孢子生殖(diplospory)”是指未减数的胚囊直接通过有丝分裂或通过中止的减数分裂事件衍生自大孢子母细胞的情况。已经报道了三种主要类型的二倍体孢子生殖，以它们在其中出现的植物命名，其为蒲公英属、苦荬菜属(Ixeris)和蝶须属(Antennaria)类型。在蒲公英属类型中，减数分裂前期已启动，但随后中止该过程，产生两个未减数的二分体，其中一个通过有丝分裂产生胚囊。在苦荬菜属类型中，在减数分裂前期之后，均等分裂，然后细胞核进行两次进一步的有丝分裂以产生八核胚囊。蒲公英属和苦荬菜属类型称为减数分裂二倍体孢子生殖，因为它们涉及减数分裂的修饰。相比之下，在蝶须属类型中，称为有丝分裂二倍体孢子生殖，大孢子母细胞不会启动减数分裂，而是直接分裂三次产生未减数的胚囊。在通过二倍体孢子生殖的配子体无融合生殖中，从未减数的大孢子产生未减数的配子体。这种未减数的大孢子由有丝分裂样分裂(有丝分裂displory)或修饰的减数分裂(减数分裂displory)产生。在通过无孢子生殖的配子体无融合生殖和通过二倍体孢子生殖的配子体无融合生殖的两者中，未减数的卵细胞孤雌生殖发育成胚。蒲公英属中的无融合生殖是二倍体孢子型的，这意味着第一雌性减数分裂(减数分裂I)被跳过，产生与亲本植物相同基因型的两个未减数的大孢子。这些大孢子的其中一个退化，而另一个存活的未减数的大孢子产生未减数的雌配子体(或胚囊)，其含有未减数的卵细胞。这种未减数的卵细胞在不受精的情况发育成与亲本植物具有相同基因型的胚。由配子体无融合生殖过程产生的种子被称为无融合生殖种子。
术语“二倍体孢子生殖功能”是指在植物中，优选在雌性子房中，优选在大孢子母细胞和/或雌配子中诱导二倍体孢子生殖的能力。因此，引入了二倍体孢子生殖功能的植物能够进行二倍体孢子生殖过程，即通过减数分裂I重组产生未减数的配子。
术语“作为配子体无融合生殖的部分的二倍体孢子生殖”是指无融合生殖过程的二倍体孢子生殖组成，即二倍体孢子生殖在通过无性过程形成种子中起的作用。特别是，除了二倍体孢子生殖功能，孤雌生殖功能在建立无融合生殖过程中也是必需的。因此，二倍体孢子生殖和孤雌生殖功能的组合可产生无融合生殖。
如本文所用，术语“二倍体孢子生殖植物”是指通过二倍体孢子生殖进行配子体无融合生殖的植物或已被诱导(例如通过遗传修饰)通过二倍体孢子生殖进行配子体无融合生殖的植物。在两种情况下，当与孤雌生殖因子组合时，二倍体孢子生殖植物产生无融合生殖种子。
如本文所用，术语“无融合生殖种子”是指从无融合生殖植物物种或通过诱导进行无融合生殖，特别是通过二倍体孢子生殖的配子体无融合生殖的植物或作物获得的种子。无融合生殖种子的特征在于它们是克隆的并且在遗传上与能够纯一传代的亲本植物和发芽植物相同。在本发明中，“无融合生殖种子”也指“克隆无融合生殖种子”。
如本文所用，术语“无融合生殖植物”是指无性繁殖而无需受精的植物。无融合生殖植物可以是已经被修饰成为无融合生殖体的有性植物，例如已经例如用本文教导的一种或多种孤雌生殖基因进行遗传修饰以获得无融合生殖植物的有性植物，或作为无融合生殖植物后代的植物。在这种情况下，无融合生殖产生的后代在遗传上与亲本植物相同。
细胞、植物、植物部分或种子的“克隆”的特征在于它们在遗传上与它们的同胞以及它们所来源的亲本植物相同。单个克隆的基因组DNA序列几乎相同，然而，突变可能引起细微的差异。
如本文所用，术语“纯一传代”或“纯一传代生物体”(也称为纯种生物体)是指始终将某种表型性状保持不变或几乎不变地传递给其后代的生物体。对于其适用的每种性状，将生物体称为纯一传代，并且术语“纯一传代”也用于描述个体遗传性状。
如本文所用，术语“F1杂种”(或子代1杂种)是指具有明显不同亲本类型的后代的第一子代。亲本类型可以是或不是自交系。F1杂种用于遗传学和选择性育种，其中它可以表现为F1杂交。明显不同的亲本类型的后代产生具有来自亲本的特征组合的新的、一致的表型。F1杂种与独特的优势如杂种优势相关，因此在农业实践中是非常受欢迎的。在本发明的一个实施方案中，本文教导的方法、基因、蛋白质、其变体或片段可用于固定F1杂种的基因型，而不管其遗传复杂性如何，并允许产生可在一个步骤中纯一传代的生物体。
如本文所用，术语“授粉”或“授粉的”是指将花粉从植物的花药(雄性部分)转移到柱头(雌性部分)，从而能够受精和繁殖的过程。它是被子植物(开花植物)所独有的。每个花粉粒是雄性单倍体配子体，适于被运送到雌配子体，其中通过在双受精过程中产生雄配子(或多个配子)来实现受精。含有雄配子的成功被子植物花粉粒(配子体)被运送到柱头，在柱头处其发芽并且其花粉管沿着花柱向下生长到子房。它的两个配子沿管向下移动到含有雌配子的配子体保持在心皮内的位置。一个核与极体融合产生胚乳组织，而另一个核与胚珠融合产生胚。
如本文所用，术语“孤雌生殖”是指一种无性繁殖的形式，其中胚的生长和发育在没有受精的情况下发生。本发明的基因和蛋白质可以与二倍体孢子生殖的(diplosporous)因子(例如基因或化学因子)组合产生无融合生殖后代。
如本文所用，术语“聚合或堆叠基因”是指将来自不同亲本系的相关或不相关基因组合到一种植物中的过程，其构成所需或有利性状(例如抗病性状、颜色、抗旱性、抗虫性等)的基础。可以使用传统育种方法进行聚合或堆叠基因，或可以通过使用分子标记加速以鉴定和保留含有所需等位基因组合的植物，并舍弃不具有所需等位基因组合的植物。在本发明的一个实施方案中，本文教导的孤雌生殖基因可有利地用于基因聚合或堆叠程序以产生无融合生殖植物或在有性作物中引入无融合生殖。
在本文件及其权利要求书中，动词“包含(to comprise)”及其动词变化以其非限制性意义使用，以表示包括该词之后的项目，但不排除未具体提及的项目。此外，不定冠词“一个”或“一种”对元素的引用不排除存在多于一个元素的可能性，除非上下文清楚地要求存在一个且仅一个元素。因此，不定冠词“一个”或“一种”通常是指“至少一个”。还应理解，当在本文中提及“序列”时，通常提及具有特定亚基序列(例如氨基酸)的实际物理分子。
如本文所用，术语“植物”包括植物细胞、植物组织或器官、植物原生质体、植物细胞组织培养物(植物可由其再生)、植物愈伤组织、植物细胞团块以及在植物或植物部分中完整的植物细胞，诸如胚、花粉、胚珠、果实、花、叶(例如收获的莴苣作物)、种子、根、根尖等。
具体实施方式
根据本发明的核苷酸序列
本发明人首次鉴定了负责孤雌生殖的基因、编码序列、启动子、3'UTR和蛋白质。所述遗传序列、启动子序列、编码序列和3'UTR序列位于Par等位基因上。本发明人还鉴定了位于Par等位基因的有性对应物，即par等位基因上的遗传序列、启动子序列、编码序列和3'UTR序列。作为导致孤雌生殖的显性等位基因的有性对应物，这些par等位基因在本文中也被指示为与孤雌生殖相关，尽管它们的存在对孤雌生殖表型没有贡献，因为par等位基因的存在可以指示有性表型，即非孤雌生殖表型。由于Par等位基因可以是显性等位基因，所以有性表型的确认可能需要将Par基因座的所有等位基因评估为par等位基因和/或需要评估Par等位基因的缺失。换言之，“与…相关”在本文中应理解为指示孤雌生殖或非孤雌生殖表型，并且任选地指示在孤雌生殖中是功能性的。对par等位基因的修饰，例如通过修饰par等位基因的一个或多个表达调控序列，诸如导致所编码蛋白质表达改变的启动子序列，可以将par等位基因赋予能够诱导孤雌生殖表型的Par等位基因。
Par和par等位基因两者均包含具有编码序列的基因，该编码序列编码本文命名为“PAR蛋白质”的蛋白质，其包含锌指C2H2-型结构域(IPR13087)，优选具有以下共有序列的锌指K2-2-样结构域：C.{2}C.{7}[K/R]A.{2}GH.[R/N].H，其也可以被注释为：CXXCXXXXXXX[K/R]AXXGHX[R/N]XH(SEQ ID NO：37)，其中X可以为任何天然存在的氨基酸，其中[K/R]表示在位置12上的氨基酸为赖氨酸或精氨酸，并且其中R/N]表示在位置19上的氨基酸为精氨酸或天冬酰胺(参见Englbrecht等,2004)。除了本文定义的锌指C2H2-型结构域，优选锌指K2-2-样结构域之外，该蛋白质包含具有共有氨基酸序列DLNXXP(SEQ ID NO：58)或DLNXP(SEQ ID NO：59)的EAR基序，其中X可以为任何天然存在的氨基酸(参见Kagale等,2010)。优选地，该蛋白质至多400个氨基酸，其中所述蛋白质包含一个或两个如本文所示的EAR基序和如本文所定义的锌指K2-2-样结构域。优选地，该蛋白质至多400个氨基酸，其中所述蛋白质仅包含一个或两个如本文所述的EAR基序和仅一个如本文所定义的锌指K2-2-样结构域，即没有如本文所定义的其它EAR基序且没有如本文所定义的其它锌指K2-2-样结构域。除了最大大小为400个氨基酸的特征之外，如本文所示的仅一个或两个EAR基序和如所定义的单个锌指K2-2-样结构域之外，PAR蛋白质可仅包含一个具有锌指共有序列C.{2}C.{12}H.{3}H(其也可以被注释为：CXXCXXXXXXXXXXXXHXXXH(SEQ ID NO：38)的另外的锌指结构域，，但更优选不包含具有锌指共有序列C.{2}C.{12}H.{3}H(SEQ ID NO：38)的另外锌指结构域。
因此，本发明提供了与植物孤雌生殖相关的核酸，其中所述核酸包含编码本文定义的PAR蛋白质的核苷酸序列。本发明还提供了与编码所述PAR蛋白质的核苷酸序列可操作连接的启动子序列和3'UTR。药用蒲公英(Taraxacum officinale)包含一个能够诱导孤雌生殖的显性Par等位基因和两个有性对应部分，即par等位基因-1和par等位基因-2，其编码分别具有SEQ ID NO：1、6或11的氨基酸序列的PAR蛋白质。Par等位基因包含具有SEQ IDNO：5的核苷酸序列的基因，par等位基因-1包含具有SEQ ID NO：10的核苷酸序列的par基因，par等位基因-2包含具有SEQ ID NO：15的核苷酸序列的par基因。Par基因包含具有SEQID NO：2的启动子序列、具有SEQ ID NO：3的编码序列和具有SEQ ID NO：4的3’UTR。par基因-1包含具有SEQ ID NO：7的启动子序列、具有SEQ ID NO：8的编码序列和具有SEQ ID NO：9的3’UTR。par基因-2包含具有SEQ ID NO：12的启动子序列、具有SEQ ID NO：13的编码序列和具有SEQ ID NO：14的3'UTR。因此，本发明提供了与植物孤雌生殖相关的核酸，其中所述核酸包含以下中的至少一种：
a)编码具有SEQ ID NO：1、6或11的氨基酸序列的蛋白质的基因；
b)具有SEQ ID NO：2、7或12的核苷酸序列的启动子；
c)具有SEQ ID NO：3、8或13的核苷酸序列的编码序列；
d)具有SEQ ID NO：4、9或14的核苷酸序列的3’UTR；
e)具有SEQ ID NO：5、10或15的核苷酸序列的基因；
f)a)-e)中任一项的变体；以及
g)a)-f)中任一项的片段。
表1提供了本文所用的所有SEQ ID NO的概述。
优选地，所述核酸在孤雌生殖中是功能性的。
在一个实施方案中，本发明的核酸包含以下中的至少一种或由以下中的至少一种组成：
a)编码具有SEQ ID NO：1的氨基酸序列的蛋白质的基因；
b)具有SEQ ID NO：2的核苷酸序列的启动子；
c)具有SEQ ID NO：3的核苷酸序列的编码序列；
d)具有SEQ ID NO：4的核苷酸序列的3'UTR；
e)具有SEQ ID NO：5的核苷酸序列的基因；
f)a)-e)中任一项的变体；以及
g)a)-f)中任一项的片段。
优选地，该实施方案的核酸和/或由其衍生的产物(诸如其RNA转录物或编码的蛋白质)指示孤雌生殖，例如包含所述核酸的植物指示所述植物显示孤雌生殖，这意味着其具有从减数的或未减数的卵细胞发育胚的能力。优选地，所述核酸和/或由其衍生的产物，例如其RNA转录物或编码的蛋白质，在孤雌生殖中是功能性的，甚至更优选诱导或能够诱导孤雌生殖，优选当存在于植物或植物细胞中时。
在另一个实施方案中，本发明的核酸包含以下中的至少一种或由以下中的至少一种组成：
a)编码具有SEQ ID NO：6或11的氨基酸序列的蛋白质的基因；
b)具有SEQ ID NO：7或12的核苷酸序列的启动子；
c)具有SEQ ID NO：8或13的核苷酸序列的编码序列；
d)具有SEQ ID NO：9或14的核苷酸序列的3'UTR；
e)具有SEQ ID NO：10或15的核苷酸序列的基因；
f)a)-e)中任一项的变体；以及
g)a)-f)中任一项的片段。
优选地，该实施方案的所述核酸和/或由其衍生的产物(诸如其RNA转录物或编码的蛋白质)不诱导或不能诱导孤雌生殖，优选当以纯合状态存在于植物或植物细胞中时。换言之，该实施方案的核酸的存在可以指示非孤雌生殖表型或有性表型，例如包含所述核酸的植物指示所述植物具有有性表型，即不能从卵细胞发育胚。
Par等位基因可以是显性等位基因。在Par等位基因是显性的情况下，为了证实植物是非孤雌生殖表型，需要将所述植物中Par基因座的所有等位基因评估为par等位基因，并且单个Par等位基因的存在足以表明该植物能够孤雌生殖。
本发明的核酸可用于筛选和/或基因分型。任选地，推定的核酸或基因和/或其衍生产物的孤雌生殖的功能性，或推定的核酸和/或其衍生产物诱导孤雌生殖的能力，可以用以下方式来评估：通以下来过降低表达、通过沉默或通过敲除孤雌生殖的植物中的所述核酸或基因，例如通过在所述基因的编码序列中引入早期终止。随后孤雌生殖表型的丧失意味着推定的核酸和/或其衍生产物能够诱导孤雌生殖。诱导孤雌生殖的能力也可以通过使功能丧失的无融合生殖植物与推定的核酸和/或其衍生产物(mRNA或蛋白质)互补来评估。这种功能丧失的无融合生殖植物可以是药用蒲公英分离株A68，其已被修饰以通过降低功能性Par等位基因的表达(例如通过缺失或敲除)而丧失无融合生殖表型。这种功能丧失的无融合生殖植物可以是药用蒲公英分离株A68，其包含Par等位基因，其中如本文定义的SEQID NO：23已被修饰为SEQ ID NO：24-27中的任一个(参见表2)。这种药用蒲公英分离株A68的功能丧失的无融合生殖植物可以通过使用CRISPR-Cas9/向导RNA复合物的靶向基因组编辑来获得，其中所述向导RNA(在本文中也表示为gRNA)包含如本文示例的SEQ ID NO：19的靶特异性序列。药用蒲公英分离株A68的Par等位基因的缺失会导致孤雌生殖的丧失，从而导致无融合生殖丧失。在所述推定的核酸或其衍生产物具有诱导孤雌生殖的能力的情况下，在将所述核酸或衍生产物引入所述分离株中时，例如通过用包含所述核酸和/或编码所述产物的载体转染所述分离株，将恢复(或拯救)无融合生殖表型。这种载体优选包含适于驱动分离株中编码产物表达的序列。例如，可能编码本发明PAR蛋白质的推定核酸可以在所述载体内与本文由SEQ ID NO：2定义的启动子和任选地与本文SEQ ID NO：4定义的3'UTR可操作地连接。对于药用蒲公英分离株A68，在没有异花传粉的情况下高结实(High seedset)是无融合生殖的清楚指示。作为另一种解释，可以排除该分离株中的自交，因为由于三倍体雄性和雌性减数分裂的不平衡，有性产生的卵细胞和花粉粒的可育性将非常低。
优选地，本文定义的变体核酸是本文定义的药用蒲公英分离株A68的Par或par等位基因的基因、启动子、编码序列和/或3'UTR的同源物或直系同源物。优选地，所述变体核酸和/或由其衍生的产物，例如其RNA转录物或编码的蛋白质，与如本文定义的孤雌生殖相关，并且任选地诱导或能够诱导孤雌生殖，优选地当存在于植物或植物细胞中时。该变体优选编码本文定义的PAR蛋白质，或与编码本文定义的PAR蛋白质的序列可操作地连接。在其它植物物种的药用蒲公英分离株A68中鉴定的Par和par基因的直系同源物可基于本文定义的PAR蛋白质的特征来鉴定。此类基因可以编码，但不限于选自以下的任何一种PAR蛋白质：菠萝(Ananas comosus)的PAR蛋白质(例如UniProtKB:A0A199URK4)，深圳拟兰(Apostasiashenzhenica)的PAR蛋白质(例如UniProtKB:A0A2I0AZW3)，拟南芥(Arabidopsisthaliana)的PAR蛋白质(例如UniProtKB:Q8GXP9,A0A178V2S4,O81793,A0A178V1Q3,A0MFC1,O81801)，拟南芥亚属Lyrata(Arabidopsis lyrata subsp.Lyrata)的PAR蛋白质(例如UniProtKB:D7MC52或D7MCE8)，花生(Arachis ipaensis)的PAR蛋白质(例如SEQ IDNO：45或SEQ ID NO：49)，二穗短柄草(Brachypodium distachyon)的PAR蛋白质(例如UniProtKB:I1J0D9)，Brassica oleracea var.oleracea的PAR蛋白质(例如UniProtKB:A0A0D3A1Q6或A0A0D3A1Q3)，油菜(Brassica campestris)的PAR蛋白质(例如UniProtKB:A0A398AHT1)，芜菁(Brassica rapa)的PAR蛋白质(例如SEQ ID NO：47)，芜菁亚属Pekinensis的PAR蛋白质(例如UniProtKB:M4D574或M4D571)，甘蓝(Brassica oleracea)的PAR蛋白质(例如UniProtKB:A0A3P6ESB1或A0A3P6F726)，油菜的PAR蛋白质(例如UniProtKB:A0A3P5ZMM3或A0A3P5Z1M1)，木豆(Cajanus cajan)的PAR蛋白质(例如SEQ IDNO：46)，芥菜(Capsella rubella)的PAR蛋白质(例如UniProtKB:R0H2J1或R0H0C2)，土瓶草(Cephalotus follicularis)的PAR蛋白质(例如UniProtKB:A0A1Q3CSK1)，鹰嘴豆(Cicerarietinum)的PAR蛋白质(例如UniProtKB:A0A3Q7YBZ1,A0A1S2YZL9,A0A3Q7Y0Z6或A0A1S2YZM6；或SEQ ID NO：55,56或57)，苦苣(Cichorium endivia)的PAR蛋白质(例如SEQID NO：39)，黄瓜(Cucumis sativus)的PAR蛋白质(例如UniProtKB:A0A0A0KGW4或A0A0A0L0X7)，香瓜(Cucumis melo)的PAR蛋白质(例如UniProtKB:A0A1S3BLF2或A0A1S3B298)，黄瓜的PAR蛋白质(例如UniProtKB:A0A0A0KAW8)，南瓜(Cucurbitamoschata)的PAR蛋白质(例如SEQ ID NO：43)，菟丝子(Cuscuta campestris)的PAR蛋白质(例如UniProtKB:A0A484MGR1)，铁皮石斛(Dendrobium catenatum)的PAR蛋白质(例如UniProtKB:A0A2I0V7N9,A0A2I0X2T2或A0A2I0W0Q8)，旋蒴苣苔(Dorcocerashygrometricum)的PAR蛋白质(例如UniProtKB:A0A2Z7D3Y1)，盐芥(Eutrema salsugineum)的PAR蛋白质(例如UniProtKB:V4LSH0；或SEQ ID NO：44)，欧洲山毛榉(Fagus sylvatica)的PAR蛋白质(例如UniProtKB:A0A2N9E5Y5,A0A2N9HAB9或A0A2N9H993)，Genlisea aurea的PAR蛋白质(例如UniProtKB:S8E1M6)，大豆(Glycine max)的PAR蛋白质(例如SEQ ID NO：51，52，53或54)，陆地棉(Gossypium hirsutum)的PAR蛋白质(例如UniProtKB:A0A1U8LDU9)，向日葵(Helianthus annuus)的PAR蛋白质(例如SEQ ID NO：21)，橡胶树(Hevea brasiliensis)的PAR蛋白质(例如SEQ ID NO：42)，黄山柳菊(Hieraciumaurantiacum)的PAR蛋白质(例如SEQ ID NO：40)，胡桃(Juglans regia)的PAR蛋白质(例如UniProtKB:A0A2I4E6B1)，莴苣(Lactuca sativa)的PAR蛋白质(例如UniProtKB:A0A2J6KZF7；或SEQ ID NO：22)，葫芦(Lagenaria siceraria)的PAR蛋白质(例如SEQ IDNO：48)，蒺藜苜蓿(Medicago truncatula)的PAR蛋白质(例如UniProtKB:G7K024)，川桑(Morus notabilis)的PAR蛋白质(例如UniProtKB:W9SMY3或W9SMQ7)，刺毛黧豆(Mucunapruriens)的PAR蛋白质(例如UniProtKB:A0A371ELJ8)，烟草(Nicotiana attenuata)的PAR蛋白质(例如UniProtKB:A0A1J6IQI6)，美花烟草(Nicotiana sylvestris)的PAR蛋白质(例如UniProtKB:A0A1U7VXJ0)，普通烟草(Nicotiana tabacum)的PAR蛋白质(例如UniProtKB:A0A1S4A651或A0A1S3YHQ2)，水稻(Oryza sativa)亚种Japonica的PAR蛋白质(例如UniProtKB:B9FGH8)，短舌野生稻(Oryza barthii)的PAR蛋白质(例如UniProtKB:A0A0D3FWX3)，黍(Panicum miliaceum)的PAR蛋白质(例如UniProtKB:A0A3L6Q010或A0A3L6T1D6)，山黄麻(Parasponia andersonii)的PAR蛋白质(例如UniProtKB:A0A2P5BMI5)，银白杨(Populus alba)的PAR蛋白质(例如UniProtKB:A0A4U5PSY9)，毛果杨(Populus trichocarpa)的PAR蛋白质(例如UniProtKB:B9H661)，石榴(Punica granatum)的PAR蛋白质(例如UniProtKB:A0A2I0IBB9,A0A218XB85或A0A218W102)，Seneciocambrensis的PAR蛋白质(例如SEQ ID NO：41)，碧桃(Prunus persica)的PAR蛋白质(例如SEQ ID NO：50)，Trema orientale的PAR蛋白质(例如UniProtKB:A0A2P5EB04)，红车轴草(Trifolium pratense)的PAR蛋白质(例如UniProtKB:A0A2K3N851)，地三叶(Trifoliumsubterraneum)的PAR蛋白质(例如UniProtKB:A0A2Z6MYD3或A0A2Z6MDR7)，红车轴草的PAR蛋白质(例如UniProtKB:A0A2K3PR44)，葡萄(Vitis vinifera)的PAR蛋白质(例如UniProtKB:A0A438C778,A0A438ESC4或A0A438DBR4)以及玉米(Zea mays)的PAR蛋白质(例如UniProtKB:A0A1D6HF46,B6UAC5,A0A3L6F4S1,A0A3L6EMC6,A0A3L6EMC6,K7UHQ6或A0A1D6KHZ4)。此类基因还可以编码选自以下的PAR蛋白质：猕猴桃(Actinidia chinensis)的PAR蛋白质(UniProtKB:A0A2R6S2S9)，甜菜(Beta vulgaris)的PAR蛋白质(UniProtKB:XP_010690656.1)，马铃薯(Solanum tuberosum)的PAR蛋白质(UniProtKB:XP_015159151.1)，番茄(Solanum lycopersicum)的PAR蛋白质(UniProtKB:A0A3Q7GXB3)，风铃辣椒(Capsicum baccatum)的PAR蛋白质(UniProtKB:A0A2G2WJR7)，茄子(Solanummelongena)的PAR蛋白质(UniProtKB:AVC18974.1)，野生大豆(Glycine soja)的PAR蛋白质(GeneBank登录号：XP_028201014.1,XP_006596577.1或UniprotKB:A0A445M3M6)，花生(Arachis hypogaea)的PAR蛋白质(UniProtKB:A0A444WUX5)，菜豆(Phaseolus vulgaris)的PAR蛋白质(UniProtKB:V7CIF6)，胡萝卜(Daucus carota)的PAR蛋白质(GeneBank登录号：XP_017245413.1)，小麦(Triticum aestivum)的PAR蛋白质(UniProtKB:A0A3B6RP64)，水稻亚种indica的PAR蛋白质(UniProtKB:A2YH63)，水稻亚种japonica的PAR蛋白质(UniProtKB:Q5Z7P5)以及可可(Theobroma cacao)的PAR蛋白质(UniProtKB:A0A061DL63)。本发明涵盖这些直系同源基因、它们的启动子序列、编码序列(包括cDNA和mRNA序列)和3'UTR。
本发明的核酸可以是，但不限于DNA(诸如基因组DNA、cDNA)或RNA(诸如mRNA)。优选地，本发明的核酸是分离的核酸。优选地，本文定义的变体核酸优选分别与SEQ ID NO：2、3、4、5、7、8、9、10、12、13、14和15的任一序列和/或编码SEQ ID NO：1、6和11的任一序列或其互补序列包含至少约60％，70％，75％，80％，85％，90％，92％，95％，97％，98％，99％或更高的核苷酸序列同一性，优选地，当使用例如Needleman和Wunsch算法(全局序列比对)以默认参数进行成对比对时。例如，SEQ ID NO：3的编码序列的变体优选与SEQ ID NO：3包含至少60％，70％，75％，80％，85％，90％，92％，95％，97％，98％，99％或更高的核苷酸序列同一性；SEQ ID NO：5的编码序列的变体优选与SEQ ID NO：5包含至少约60％，70％，75％，80％，85％，90％，92％，95％，97％，98％，99％或更高的核苷酸序列同一性；等等。
优选地，该变体与SEQ ID NO：2、3、4、5、7、8、9、10、12、13、14和15，以及编码SEQ IDNO：1、6和11的序列或其互补序列中的任一个通过一个或多个核苷酸缺失、插入和/或置换而不同，并且包括天然和/或合成/人工变体。“天然变体”是在自然界，例如在其它蒲公英属物种或其它植物中发现的变体。优选地，该变体为来自不同植物物种的核苷酸序列(基因、启动子序列或编码序列)，例如来自与Taraxacum officinale sensu lato不同的蒲公英属物种，例如不同的栽培品种、种质(accession)或育种系。所述变体还可以在属于蒲公英属的植物之外的植物中发现和/或从所述植物中分离。
如本文所示，本发明的核酸还包括如本文所定义的Par或par等位基因的确定基因、启动子或编码序列的片段，或其任何变体。“片段”包含或由SEQ ID NO：2、3、4、5、7、8、9、10、12、13、14和15中任一个，和/或编码SEQ ID NO：1、6和11中任一个的序列，或其变体的连续核苷酸序列组成，诸如至少约10，12，15，18，20，30，50，100，150，200，250，300，500，1000，2000或更多个连续核苷酸，或其优选能够与所述序列杂交的互补序列。在一个实施方案中，这种片段可以在如本文所定义的在孤雌生殖中是功能性的(优选能够诱导孤雌生殖)。在另一个实施方案中，这种片段在孤雌生殖中可能不是功能性的，但可能与孤雌生殖相关，例如因为该片段可能与在孤雌生殖中具有功能性的序列杂交，因此可以对其进行指示。这种片段可用作例如PCR引物或杂交探针，因此可用作遗传标记，用于定位分析(mapping assay)或分子分析和/或用于鉴定和/或分离来自其它植物的Par或par等位基因。
优选地，本发明的核酸包含编码如本文定义的PAR蛋白质的基因的调控序列(优选启动子序列)或由其组成，其中所述调控序列(优选启动子序列)包含核酸插入物(优选双链DNA插入物)，其中所述插入物的长度为50至2000bp，100至1900bp，200至1800bp，300至1700bp，400至1600bp，500至1500bp，600至1400bp，1000至1400，1200至1400，或1300至1400bp。甚至更优选地，所述插入物的长度为约1300bp。优选地，插入物与如本文定义的孤雌生殖表型相关，并且任选地在孤雌生殖表型中是功能性的。优选地，所述插入物位于启动子序列内，该启动子序列直接位于编码PAR蛋白质的序列的上游(3')，优选使得所述插入物的3'末端与编码PAR蛋白质的序列的起始密码子之间的距离为50-200bp，优选约50，60，70，80，90，100，110，120，130，140，150，160，170，180，190或200bp，最优选约102bp。优选地，所述插入物被定位为使得插入物的3'末端核苷酸位于与SEQ ID NO：2的核苷酸1798和/或SEQID NO：5的核苷酸1798的位置同源的位置。优选地，所述插入物没有开放阅读框。甚至更优选地，所述插入物为微型反向重复转座元件(MITE)或MITE样序列，其中所述MITE或MITE样序列是一种非自主元件，其特征在于含有缺乏开放阅读框的内部序列，其侧翼为末端反向重复(TIR)，而TIR侧翼为小的同向重复(靶位点重复)。关于MITE、TIR和序列的进一步描述，参考Guo et al,Scientific Reports.2017Jun 1；7(1):2634，其通过引用并入本文。所述插入物，优选所述MITE或MITE样序列，可以与SEQ ID NO：60具有至少约50％，60％，70％，80％，90％，95％，98％，99％或更高的同一性。优选地，所述插入物与如本文定义的孤雌生殖表型相关，并且任选地在孤雌生殖表型中是功能性的。在进一步优选的实施方案中，本发明的核酸包含调控序列(优选启动子序列)或由其组成，该调控序列在如上定义的位置包含所述插入物。优选地，本发明的核酸包含编码本文定义的PAR蛋白质的序列或由其组成，该序列与所述启动子序列可操作地连接，其中优选地，所述启动子序列直接位于编码PAR蛋白质的序列的上游。任选地，本发明的所述核酸可以包含一个或多个另外的转录调控序列。
在一个实施方案中，本发明的核酸可以源自蒲公英属系(例如，Taraxacumofficinale sensu lato)或其它物种。
在一个实施方案中，本发明的核酸来自蒲公英或Taraxacum officinale sensulato以外的来源。
在一个实施方案中，本发明涵盖源自存在孤雌生殖的植物(诸如野生或栽培植物和/或来自其它植物)的同源或直系同源Par等位基因。通过使用提供的核苷酸序列或其部分作为引物或探针，可以容易地分离这种同源物或直系同源物。例如，可以使用中等或严格的核酸杂交方法，例如使用本文定义的核苷酸序列的片段或其互补物。变体还可以使用已知方法如PCR、严格杂交方法等从其它野生或栽培的无融合生殖或非无融合生殖植物(和/或从其它植物)分离。因此，SEQ ID NO：2、3、4、5、7、8、9、10、12、13、14和15中任一个的变体和/或编码SEQ ID NO：1、6和11的序列的变体还包括在其它蒲公英属植物、品系或栽培品种中天然(或在自然界中)发现的和/或在其它植物中天然发现的核酸。
为了在宿主或宿主细胞中最佳表达，本文教导的编码序列可以通过使用可用的密码子选择表(codon usage Table)(例如，更适合于感兴趣的植物中的表达)使密码子选择适应于植物基因中最优选的来进行密码子优化，特别是适应于感兴趣的植物属或种的天然基因(Bennetzen and Hall,1982,J.Biol.Chem.257,3026-3031；Itakura等,1977Science198,1056-1063)。各种植物物种的密码子选择表由例如Ikemura(1993,In"PlantMolecular Biology Labfax",Croy,ed.,Bios Scientific Publishers Ltd.)和Nakamura等人(2000,Nucl.Acids Res.28,292.)以及主要的DNA序列数据库(例如德国海德堡的EMBL)公开。因此，可以构建合成DNA序列，使得可以使用所述合成DNA序列产生相同或基本相同的蛋白质。在专利和科学文献中可以找到几种将密码子选择修饰为宿主细胞优选的密码子选择的技术。密码子选择修饰的确切方法对于本发明并不是关键的。
可以常规地对SEQ ID NO：2、3、4、5、7、8、9、10、12、13、14和15中的任一个和/或编码SEQ ID NO：1、6和11的序列或其变体进行小的修饰，即通过随机或靶向诱变(例如通过化学诱变或CRISPR-核酸内切酶介导的诱变)。如本文所教导的对所述序列的更深刻的修饰可以通过使用可用技术从头DNA合成所需序列来常规进行。
在一个实施方案中，通过在蛋白质的N末端添加或缺失一个或多个氨基酸，可以修饰本发明的核酸，使得由所述核酸编码的本发明蛋白质的N末端具有最佳翻译起始背景。通常优选的是，在植物细胞中表达的本发明的蛋白质以Met-Asp或Met-Ala二肽开始，以获得最佳翻译起始。因此可以在现有的Met之后插入Asp或Ala密码子，或者第二个密码子Val可以被Asp(GAT或GAC)或Ala(GCT、GCC、GCA或GCG)的密码子取代。也可以修饰核苷酸序列以去除非法剪接位点。
在一个实施方案中，本发明的核酸可以具有(遗传)显性功能，优选通过(过)表达具有氨基酸序列SEQ ID NO：1的功能性蛋白质或其变体或功能性片段，例如在另一种植物(即除了蒲公英或Taraxacum officinale sensu lato之外)中发现的直系同源物或其片段来提供。
优选地，本发明的核酸编码蛋白质或其功能性片段，当在植物中产生时，其是功能性的并诱导和/或增强孤雌生殖。例如，当包含SEQ ID NO：3或5的核酸或其变体或片段在合适的植物组织中表达(转录和翻译)并产生适量的本发明的蛋白质时，与仅在它们缺乏所述核酸方面不同的植物相比，孤雌生殖作用显著增强。功能性还可以通过在合适的宿主植物(诸如非孤雌生殖的蒲公英系)中(过)表达本发明的核酸，并在生物测定中分析转化体的孤雌生殖效果(例如如实施例2所述)容易地测试。所述核酸的功能性优选通过将其中一种或多种这些核酸(过)表达的测试植物与对照植物进行比较来评估，所述对照植物与测试植物的区别仅在于对照植物缺乏所述核酸的(过)表达。或者，与孤雌生殖相关的本发明核酸的沉默或破坏可导致功能丧失，即降低孤雌生殖。
本发明的核酸可用于产生载体或质粒，用于在合适的宿主细胞中表达本发明的蛋白质，或用于沉默一个或多个内源性孤雌生殖基因或基因家族。因此，包含本发明核酸和/或沉默构建体的构建体、载体和/或质粒也包括在本发明中。
根据本发明的氨基酸序列
本发明提供了如本文所定义的PAR蛋白质。本发明还提供了与植物孤雌生殖相关的蛋白质，其中所述蛋白质：
a)由本发明的核酸编码；
b)具有SEQ ID NO：1、6或11的氨基酸序列；
c)是a)和/或b)的变体；和/或
d)是a)-c)中任一项的片段，
其中优选地，所述蛋白质在孤雌生殖中是功能性的。
在一个实施方案中，本发明的蛋白质为：
a)由SEQ ID NO：3、8或13中任一项的核酸编码；
b)具有SEQ ID NO：1、6或11的氨基酸序列；
c)是a)和/或b)的变体；和/或
d)是a)-c)中任一项的片段，
其中优选地，本发明的蛋白质适于诱导孤雌生殖。
在一个实施方案中，本发明的蛋白质为：
a)由SEQ ID NO：3或5的核酸编码；
b)具有SEQ ID NO：1的氨基酸序列；
c)是a)和/或b)的变体；和/或
d)是a)-c)中任一项的片段，
其中优选地，本发明的蛋白质适于诱导孤雌生殖。所述变体优选是本文定义的PAR蛋白质。优选地，该蛋白质或蛋白质片段由SEQ ID NO：3或5的核酸或其变体和/或片段编码，或者这种蛋白质包含SEQ ID NO：1或其变体和/或片段。优选地，所述变体包含或由分别与SEQ ID NO：1、6或11具有至少约50％，60％，70％，80％，90％，95％，98％，99％或更高同一性的氨基酸序列组成，优选地，当使用例如Needleman和Wunsch算法(全局序列比对)以默认参数进行成对比对时。变体与提供的序列的不同之处在于一个或多个氨基酸残基缺失、插入和/或置换，并且包括天然和/或合成/人工变体。具有由本发明核酸编码的氨基酸的蛋白质的变体，优选由SEQ ID NO：3、5、8、10、13、15中任一个编码的蛋白质的变体，或具有SEQID NO：1、6或11中任一个的氨基酸序列的蛋白质的变体，可以是同源物或直系同源物。本发明所包括的这种直系同源蛋白质可以是，但不限于选自以下的任何一种PAR蛋白质：菠萝的PAR蛋白质(例如UniProtKB:A0A199URK4)，深圳拟兰的PAR蛋白质(例如UniProtKB:A0A2I0AZW3)，拟南芥的PAR蛋白质(例如UniProtKB:Q8GXP9,A0A178V2S4,O81793,A0A178V1Q3,A0MFC1,O81801)，拟南芥亚属Lyrata的PAR蛋白质(例如UniProtKB:D7MC52或D7MCE8)，花生的PAR蛋白质(例如SEQ ID NO：45或SEQ ID NO：49)，二穗短柄草的PAR蛋白质(例如UniProtKB:I1J0D9)，Brassica oleracea var.oleracea的PAR蛋白质(例如UniProtKB:A0A0D3A1Q6或A0A0D3A1Q3)，油菜的PAR蛋白质(例如UniProtKB:A0A398AHT1)，芜菁的PAR蛋白质(例如SEQ ID NO：47)，芜菁亚属Pekinensis的PAR蛋白质(例如UniProtKB:M4D574或M4D571)，甘蓝的PAR蛋白质(例如UniProtKB:A0A3P6ESB1或A0A3P6F726)，油菜的PAR蛋白质(例如UniProtKB:A0A3P5ZMM3或A0A3P5Z1M1)，木豆的PAR蛋白质(例如SEQ ID NO：46)，芥菜的PAR蛋白质(例如UniProtKB:R0H2J1或R0H0C2)，土瓶草的PAR蛋白质(例如UniProtKB:A0A1Q3CSK1)，鹰嘴豆的PAR蛋白质(例如UniProtKB:A0A3Q7YBZ1,A0A1S2YZL9,A0A3Q7Y0Z6或A0A1S2YZM6；或SEQ ID NO:55,56或57)，苦苣的PAR蛋白质(例如SEQ ID NO：39)，黄瓜的PAR蛋白质(例如UniProtKB:A0A0A0KGW4或A0A0A0L0X7)，香瓜的PAR蛋白质(例如UniProtKB:A0A1S3BLF2或A0A1S3B298)，黄瓜的PAR蛋白质(例如UniProtKB:A0A0A0KAW8)，南瓜的PAR蛋白质(例如SEQ ID NO：43)，菟丝子的PAR蛋白质(例如UniProtKB:A0A484MGR1)，铁皮石斛的PAR蛋白质(例如UniProtKB:A0A2I0V7N9,A0A2I0X2T2或A0A2I0W0Q8)，旋蒴苣苔的PAR蛋白质(例如UniProtKB:A0A2Z7D3Y1)，盐芥的PAR蛋白质(例如UniProtKB:V4LSH0；或SEQ ID NO：44)，欧洲山毛榉的PAR蛋白质(例如UniProtKB:A0A2N9E5Y5,A0A2N9HAB9或A0A2N9H993)，Genlisea aurea的PAR蛋白质(例如UniProtKB:S8E1M6)，大豆的PAR蛋白质(例如SEQ ID NO：51，52，53或54)，陆地棉的PAR蛋白质(例如UniProtKB:A0A1U8LDU9)，向日葵的PAR蛋白质(例如SEQ ID NO：21)，橡胶树的PAR蛋白质(例如SEQ ID NO：42)，黄山柳菊的PAR蛋白质(例如SEQ ID NO：40)，胡桃的PAR蛋白质(例如UniProtKB:A0A2I4E6B1)，莴苣的PAR蛋白质(例如UniProtKB:A0A2J6KZF7；或SEQ ID NO：22)，葫芦的PAR蛋白质(例如SEQ ID NO：48)，蒺藜苜蓿的PAR蛋白质(例如UniProtKB:G7K024)，川桑的PAR蛋白质(例如UniProtKB:W9SMY3或W9SMQ7)，刺毛黧豆的PAR蛋白质(例如UniProtKB:A0A371ELJ8)，烟草的PAR蛋白质(例如UniProtKB:A0A1J6IQI6)，美花烟草的PAR蛋白质(例如UniProtKB:A0A1U7VXJ0)，普通烟草的PAR蛋白质(例如UniProtKB:A0A1S4A651或A0A1S3YHQ2)，水稻亚种Japonica的PAR蛋白质(例如UniProtKB:B9FGH8)，短舌野生稻的PAR蛋白质(例如UniProtKB:A0A0D3FWX3)，黍的PAR蛋白质(例如UniProtKB:A0A3L6Q010或A0A3L6T1D6)，山黄麻的PAR蛋白质(例如UniProtKB:A0A2P5BMI5)，银白杨的PAR蛋白质(例如UniProtKB:A0A4U5PSY9)，毛果杨的PAR蛋白质(例如UniProtKB:B9H661)，石榴的PAR蛋白质(例如UniProtKB:A0A2I0IBB9,A0A218XB85或A0A218W102)，Senecio cambrensis的PAR蛋白质(例如SEQ ID NO：41)，碧桃的PAR蛋白质(例如SEQ ID NO：50)，Trema orientale的PAR蛋白质(例如UniProtKB:A0A2P5EB04)，红车轴草的PAR蛋白质(例如UniProtKB:A0A2K3N851)，地三叶的PAR蛋白质(例如UniProtKB:A0A2Z6MYD3或A0A2Z6MDR7)，红车轴草的PAR蛋白质(例如UniProtKB:A0A2K3PR44)，葡萄的PAR蛋白质(例如UniProtKB:A0A438C778,A0A438ESC4或A0A438DBR4)以及玉米的PAR蛋白质(例如UniProtKB:A0A1D6HF46,B6UAC5,A0A3L6F4S1,A0A3L6EMC6,A0A3L6EMC6,K7UHQ6或A0A1D6KHZ4)。这种直系同源蛋白质还可以是选自以下的PAR蛋白质：猕猴桃的PAR蛋白质(UniProtKB:A0A2R6S2S9)，甜菜的PAR蛋白质(UniProtKB:XP_010690656.1)，马铃薯的PAR蛋白质(UniProtKB:XP_015159151.1)，番茄的PAR蛋白质(UniProtKB:A0A3Q7GXB3)，风铃辣椒的PAR蛋白质(UniProtKB:A0A2G2WJR7)，茄子的PAR蛋白质(UniProtKB:AVC18974.1)，野生大豆的PAR蛋白质(GeneBank登录号：XP_028201014.1,XP_006596577.1或UniprotKB:A0A445M3M6)，花生的PAR蛋白质(UniProtKB:A0A444WUX5)，菜豆的PAR蛋白质(UniProtKB:V7CIF6)，胡萝卜的PAR蛋白质(GeneBank登录号：XP_017245413.1)，小麦的PAR蛋白质(UniProtKB:A0A3B6RP64)，水稻亚种indica的PAR蛋白质(UniProtKB:A2YH63)，水稻亚种japonica的PAR蛋白质(UniProtKB:Q5Z7P5)以及可可的PAR蛋白质(UniProtKB:A0A061DL63)。
因此，本发明包括的SEQ ID NO：1蛋白质的变体可以是，但不限于本文定义的任何一种直系同源物PAR蛋白质。
当存在于植物或植物细胞中时，本发明的PAR蛋白质和/或具有SEQ ID NO：1、6或11的蛋白质的变体能够诱导孤雌生殖。蛋白质的变体可以是所述植物或植物细胞的内源性或非内源性蛋白质。任选地，本发明的PAR蛋白质和/或具有SEQ ID NO：1、6或11的蛋白质的变体，当蛋白质的表达改变，优选增加时，能够诱导孤雌生殖。优选地，这种改变的表达，优选增加的表达，在卵细胞内。改变或增加的表达可以是所述蛋白质在植物或植物细胞中的从头表达，或者可以是内源性蛋白质在植物或植物细胞中的增加的表达。本领域技术人员知晓增加蛋白质表达的方法。蛋白质在植物或植物细胞中的从头表达可以通过例如用编码该蛋白质的构建体或载体转染植物或植物细胞，将编码该蛋白质的基因渗入植物或植物细胞的后代，和/或例如通过遗传修饰来修饰导致编码所述蛋白质的序列的内源序列来诱导。任选地，这种构建体或载体包含与卵细胞启动子可操作连接的编码PAR蛋白质的序列。本领域技术人员知晓卵细胞启动子。能够在植物的卵细胞中驱动表达的示例性卵细胞启动子包括，但不限于卵细胞特异性基因EC1.1、EC1.2、EC1.3、EC1.4或EC1.5的启动子(参见，例如Sprunck等Science,338:1093-1097(2012)；AT2G21740；Steffen et al,Plant Journal51:281-292(2007))，拟南芥DD45启动子(Ohnishi等PlantPhysiology 165:1533-1543(2014))。
优选地，本发明的构建体或载体包含编码PAR蛋白质的序列，该序列与调控序列可操作地连接，该调节序列(优选为启动子序列)包含核酸插入物(优选双链DNA插入物)，其中所述插入物的长度为50至2000bp，100至1900bp，200至1800bp，300至1700bp，400至1600bp，500至1500bp，600至1400bp，1000至1400，1200至1400，或1300至1400bp。甚至更优选地，所述插入物的长度为约1300bp。优选地，插入物与如本文定义的孤雌生殖表型相关，并且任选地在孤雌生殖表型中是功能性的。优选地，所述插入片段位于启动子序列内，该启动子序列直接位于编码PAR蛋白质的序列的上游(3')，优选使得所述插入片段的3'末端与编码PAR蛋白质的序列的起始密码子之间的距离为50-200bp，优选约50，60，70，80，90，100，110，120，130，140，150，160，170，180，190或200bp，最优选约102bp。优选地，所述插入物被定位为使得插入物的3'末端核苷酸位于与SEQ ID NO：2的核苷酸1798和/或SEQ ID NO：5的核苷酸1798的位置同源的位置。优选地，所述插入物没有开放阅读框。甚至更优选地，所述插入物是微型反向重复转座元件(MITE)或MITE样序列，其中所述MITE或MITE样序列是一种非自主元件，其特征在于含有缺乏开放阅读框的内部序列，其侧翼为末端反向重复(TIR)，而TIR侧翼为小的同向重复(靶位点重复)。关于MITE、TIR和序列的进一步描述，参考Guo et al,Scientific Reports.2017Jun 1；7(1):2634，其通过引用并入本文。所述插入物，优选所述MITE或MITE样序列，可以与SEQ ID NO：60具有至少约50％，60％，70％，80％，90％，95％，98％，99％或更高的同一性。优选地，所述插入物与如本文定义的孤雌生殖表型相关，并且任选地在孤雌生殖表型中是功能性的。在进一步优选的实施方案中，本发明的构建体或载体包含调控序列(优选启动子序列)或由其组成，该调控序列在如上定义的位置包含所述插入物。优选地，构建体或载体包含编码本文定义的PAR蛋白质的序列或由其组成，该序列与所述启动子序列可操作地连接，其中优选地，所述启动子序列直接位于编码PAR蛋白质的序列的上游。任选地，本发明的所述构建体或载体可以包含一个或多个另外的转录调控序列。
此外或可选地，这种构建体或载体包含编码PAR蛋白质的序列，该序列与SEQ IDNO：2的启动子可操作地连接。内源性蛋白质的改变或增加的表达可以通过修饰与编码序列可操作地连接的一个或多个调控序列来诱导。例如，可以修饰与编码蛋白质的序列可操作地连接的启动子序列，例如通过遗传修饰。在一个优选的实施方案中，将如上定义的插入物导入启动子序列，优选在如上定义的位置。如本文所述，这种能够诱导孤雌生殖的功能性可以通过使用编码所述变体的核酸的孤雌生殖中的功能性的合适测试来评估。本发明的蛋白质可以是分离的蛋白质。
“天然变体”是在自然界中发现的那些，例如在栽培的或野生的莴苣植物和/或其它植物中发现的那些。还包括片段，即本发明蛋白质的非全长肽，优选功能性片段，即在合适的宿主植物中表达时能够诱导孤雌生殖。本文所教导的蛋白质片段包括包含或由本发明核酸编码的至少约10，20，30，40，50，100，150，200，250或更多个连续氨基酸序列组成的肽，特别是包含或由SEQ ID NO：1、6或11的至少约10，20，30，40，50，100，150，200，250或更多个连续氨基酸或其变体(如本文所定义)组成的肽。在自然界中发现的序列在本文中也表示为“野生型”。
本发明的蛋白质可以从天然来源分离、通过化学合成(使用例如肽合成仪，诸如由Applied Biosystems提供的)从头合成，或通过表达本文教导的编码本发明蛋白质的核苷酸序列由重组宿主细胞产生。本发明的蛋白质也可以通过从本文定义的本发明的核酸表达而产生。
蛋白质变体可包含碱性(例如Arg，His，Lys)、酸性(例如Asp，Glu)、非极性(例如Ala，Val，Trp，Leu，Ile，Pro，Met，Phe，Trp)或极性(例如Gly，Ser，Thr，Tyr，Cys，Asn，Gln)类别内的保守氨基酸取代。此外，非保守氨基酸取代也在本发明的范围内。
嵌合蛋白质，诸如由来自不同来源的结构域组成的蛋白质，诸如SEQ ID NO：1、6或11的蛋白质的N末端(例如获自Taxaracum或植物物种X)和SEQ ID NO：1、6或11的变体的中间结构域和/或C末端结构域(例如获自Taxaracum或植物物种Y或另一植物物种)也包括在本文中。优选地，嵌合蛋白质由来自至少两种直系同源蛋白质的结构域组成。这种嵌合蛋白质可以具有改善的功能性，例如当在植物宿主中表达时，它可以比天然蛋白质更有效地赋予孤雌生殖。
本发明还包括编码本发明蛋白质、蛋白质变体或蛋白质片段的所有核苷酸序列(RNA、cDNA、基因组DNA等)。由于遗传密码的简并性，各种核苷酸序列可能编码相同的氨基酸序列。
孤雌生殖的植物及其制备方法
在另一方面，本发明涉及植物(包括例如植物细胞、器官、种子和植物部分)，以及制备植物的方法，该植物显示出修饰的孤雌生殖，任选地与天然或未修饰的植物相比具有修饰的(优选诱导的)孤雌生殖的转基因植物。这种植物可以使用不同的方法制备，例如如本文进一步描述的。优选地，本发明的植物通过技术手段(优选通过本文所述的方法)获得。这种技术手段是本领域技术人员公知的，并且包括遗传修饰，诸如例如随机诱变、靶向诱变和核酸插入中的至少一种。
优选地，本发明的植物不是通过基本上生物的方法获得的。优选地，本发明的植物不仅仅通过基本上生物的方法获得。优选地，本发明的植物不是通过在植物中引入孤雌生殖的任何基本上生物的方法获得的，优选不是直接获得的。优选地，本发明的植物不仅仅通过在植物中引入孤雌生殖的任何基本上生物的方法获得。优选地，本发明的植物不是天然存在的植物，即不是自然界存在的植物。
特别地，本发明提供一种产生孤雌生殖的植物的方法，其包含以下步骤：
a)在一种或多种植物细胞中引入本发明的核酸和/或其衍生产物，其能够诱导孤雌生殖和/或在孤雌生殖中是功能性的；
b)任选地选择包含所述核酸的植物细胞，其中优选地，所述核酸被整合至所述植物细胞的基因组中；以及
c)从所述植物细胞再生植物，
其中优选地，本发明的所述核酸编码如本文定义的PAR蛋白质(其在孤雌生殖中有功能性的)，或与编码如本文定义的PAR蛋白质(其在孤雌生殖中有功能性的)的序列可操作地连接，和/或是SEQ ID NO：2-5中的任一个，或编码SEQ ID NO：1的蛋白质，或其变体或片段。
本发明进一步提供了一种产生无融合生殖植物的方法，其包含以下步骤：
a)在一个或多个能够进行不完全减数分裂的植物细胞中引入本发明的核酸和/或其衍生产物，其能够诱导孤雌生殖；
b)任选地选择包含所述核酸的植物细胞，其中优选地，所述核酸被整合至所述植物细胞的基因组中；以及
c)从所述植物细胞再生植物，
其中优选地，本发明的所述核酸编码如本文定义的PAR蛋白质(其在孤雌生殖中有功能性的)，或与编码如本文定义的PAR蛋白质(其在孤雌生殖中有功能性的)的序列可操作地连接，和/或是SEQ ID NO：2-5中的任一个，或编码SEQ ID NO：1的蛋白质，或其变体或片段。通过引入能够赋予不完全减数分裂的核酸可以获得能够进行不完全减数分裂的植物细胞。任选地，在引入本发明的核酸之前、同时或之后将所述核酸引入植物细胞中。
本发明的核酸可以通过转化、基因渗入、体细胞杂交和/或原生质体融合被引入一种或多种植物细胞中。这种核酸可以是外源性核酸，即天然不存在于所述植物细胞中的核酸。
本发明的核酸可以通过修饰内源性核酸而被引入一种或多种植物细胞中以获得本发明的核酸。内源基因的修饰优选包含在编码序列和/或调控和/或启动子序列中的一个或多个核苷酸的随机或定向突变，或例如通过同源重组的短或较大序列的插入或缺失，以改变内源性蛋白质的表达。这种方法优选导致一个或多个内源par等位基因修饰为如本文定义的Par等位基因。随机诱变可以是，但不限于化学诱变和γ辐射。化学诱变的非限制性实例包括，但不限于EMS(甲磺酸乙酯)，MMS(甲磺酸甲酯)，NaN3(叠氮化钠)D)，ENU(N-乙基-N-亚硝基脲)，AzaC(氮杂胞苷)和NQO(4-硝基喹啉1-氧化物)。任选地，诱变系统诸如TILLING(定向诱导的局部损伤(Targeting Induced Local Lesions IN Genomics)McCallum等,2000,Nat Biotech 18:455，以及McCallum等2000,Plant Physiol.123,439-442，二者均通过引用并入本文)可用于产生具有本文定义的修饰基因的植物品系。TILLING使用传统的化学诱变(例如EMS诱变)，然后对突变进行高通量筛选。因此，可以使用TILLING获得包含具有一种或多种所需突变的基因的植物、种子和组织。定向诱变是可以设计成改变特定核苷酸或核酸序列的诱变，诸如但不限于寡核苷酸定向诱变，RNA引导的核酸内切酶(例如CRISPR技术)，TALEN或锌指技术。
优选地，修饰是在编码本文定义的PAR蛋白质的基因的启动子序列中的修饰。优选地，该修饰引入或增加如本文的PAR蛋白质的表达。优选地，该修饰在卵细胞中引入或增加本文定义的PAR蛋白质的表达。
因此，本发明的方法可以包含以下步骤：
a)在一种或多种植物细胞中修饰核酸，该核酸是编码与孤雌生殖相关和/或在孤雌生殖中是功能性的蛋白质的序列，或与编码与孤雌生殖相关和/或在孤雌生殖中是功能性的蛋白质的序列可操作地连接，其中优选地，所述核酸在所述一种或多种植物细胞的基因组内；
b)任选地选择包含所述修饰的核酸的植物细胞；以及
c)从所述植物细胞再生植物，
其中优选地，所述与孤雌生殖相关和/或在孤雌生殖中是功能性的蛋白质具有如上文所述的根据本发明的氨基酸序列。优选地，步骤a)中待修饰的核酸是内源性核酸，优选地包含核苷酸序列或由其组成，该核苷酸序列是编码如本文定义的PAR蛋白质和/或具有SEQ ID NO：1、6或11的氨基酸序列的蛋白质的序列，或与编码如本文定义的PAR蛋白质和/或具有SEQ ID NO：1、6或11的氨基酸序列的蛋白质的序列可操作的连接，或其变体或片段。
在特别优选的实施方案中，所述核酸为编码与本文定义的孤雌生殖相关的蛋白质的基因的(5'UTR)启动子序列。优选地，所述修饰为引入核酸插入物，优选双链DNA插入物，其中所述插入物的长度为50至2000bp，100至1900bp，200至1800bp，300至1700bp，400至1600bp，500至1500bp，600至1400bp，1000至1400，1200至1400或1300至1400bp。甚至更优选地，所述插入物的长度为约1300bp。优选地，插入物与如本文定义的孤雌生殖表型相关，并且任选地在孤雌生殖表型中是功能性的。优选地，将所述插入物引入直接位于编码PAR蛋白质的序列上游(3')的启动子序列内，优选使得所述插入物的3'末端与编码PAR蛋白质的序列的起始密码子之间的距离为50-200bp，优选约50，60，70，80，90，100，110，120，130，140，150，160，170，180，190或200bp，最优选约102bp。优选地，所述插入物被引入，使得插入物的3'末端核苷酸在与SEQ ID NO：2的核苷酸1798和/或SEQ ID NO：5的核苷酸1798的位置同源的位置。优选地，所述插入物没有开放阅读框。甚至更优选地，所述插入物是微型反向重复转座元件(MITE)或MITE样序列，其中所述MITE或MITE样序列是一种非自主元件，其特征在于含有缺乏开放阅读框的内部序列，其侧翼为末端反向重复(TIR)，而TIR侧翼为小的同向重复(靶位点重复)。关于MITE、TIR和序列的进一步描述，参考Guo et al,ScientificReports.2017Jun1；7(1):2634，其通过引用并入本文。所述插入物，优选所述MITE或MITE样序列，可以与SEQ ID NO：60具有至少约50％，60％，70％，80％，90％，95％，98％，99％或更高的同一性。优选地，所述插入物与如本文定义的孤雌生殖表型相关，并且任选地在孤雌生殖表型中是功能性的。
优选地，核苷酸序列的修饰导致所述蛋白质的引入或增加的表达，优选在从植物细胞再生的植物的卵细胞中。优选地，修饰的启动子序列包含与SEQ ID NO：2具有至少约75％，80％，85％，90％，95％，96％，97％，98％，99％或100％序列同一性的序列。
此外，本发明的方法可以包含以下步骤：
a)在一个或多个能够进行不完全减数分裂的植物细胞中修饰核酸，该核酸是编码与孤雌生殖相关和/或在孤雌生殖中是功能性的蛋白质的序列或与编码与孤雌生殖相关和/或在孤雌生殖中是功能性的蛋白质的序列可操作地连接，其中优选地，所述核酸在所述一种或多种植物细胞的基因组内；
b)任选地选择包含所述修饰的或改变的核酸的植物细胞；以及
c)从所述植物细胞再生植物，
其中优选地，所述与孤雌生殖相关和/或在孤雌生殖中是功能性的蛋白质具有如上文所述的根据本发明的蛋白质的氨基酸序列。优选地，步骤a)中待修饰的核酸是内源性核酸，优选地包含核苷酸序列或由其组成，该核苷酸序列是编码如本文定义的PAR蛋白质和/或具有SEQ ID NO：1、6或11的氨基酸序列的蛋白质的序列，或与编码如本文定义的PAR蛋白质和/或具有SEQ ID NO：1、6或11的氨基酸序列的蛋白质的序列可操作地连接，或其变体或片段。优选地，步骤a)中待修饰的核酸是内源性核酸。
在一个特别优选的实施方案中，所述核酸是编码与本文定义的孤雌生殖相关和/或在孤雌生殖中是功能性的蛋白质的基因的启动子序列。优选地，核苷酸序列的修饰导致所述蛋白质的引入或增加的表达，优选在从所述植物细胞再生的植物的卵细胞中。优选地，修饰的启动子序列是与本文定义的PAR蛋白质的编码序列可操作连接的启动子序列。优选地，所述经修饰的启动子序列经修饰以包含如上文所定义的插入物，优选地在如上文所定义的位置处。
优选地，修饰的启动子序列包含与SEQ ID NO：2具有至少约75％，80％，85％，90％，95％，96％，97％，98％，99％或100％序列同一性的序列。
本发明还提供了一种产生无融合生殖杂交种子的方法，其包含以下步骤：
a)将有性繁殖的第一植物与第二植物的花粉进行杂交，以产生F1杂交种子；以及
b)任选地从所述F1种子中选择包含无融合生殖表型的种子；
其中所述第一植物和/或第二植物能够进行不完全减数分裂，并且其中所述第二植物包含本发明的核酸，并且其中优选地通过基因分型进行所述选择。优选地，所述第二植物包含本发明的核酸，所述核酸是SEQ ID NO：2-5中的任一个，或编码SEQ ID NO：1的蛋白质，或其变体或片段。
本发明的核酸可以包含在嵌合基因、遗传构建体或核酸载体中。在本发明的一个实施方案中，本发明的核酸可用于制备嵌合基因和/或包含该核酸的载体，用于将该核酸转移到宿主细胞中并在宿主细胞中产生由所述核酸编码的功能性(优选能够诱导孤雌生殖)蛋白质。用于在植物细胞中生产这种蛋白质(或蛋白质片段或变体)的载体在本文中称为“表达载体”。宿主细胞优选为植物细胞。
用于任选地瞬时但优选稳定地将编码蛋白质的核苷酸序列导入宿主细胞基因组的嵌合基因、构建体和/或载体的构建是本领域公知的。为了产生用于诱导孤雌生殖和/或改善孤雌生殖中的功能性的嵌合基因，可以使用标准分子生物学技术将编码SEQ ID NO：1、6或11的蛋白质或其功能性变体和/或功能性片段的核苷酸序列可操作地连接至适于在宿主细胞中表达的启动子序列。启动子序列可以已经存在于载体中，使得编码所述蛋白质的核苷酸序列可以简单地插入到启动子序列下游的载体中。然后可以将载体用于转化宿主细胞，并且可以将本发明的核酸和/或嵌合基因插入核基因组中或插入质体、线粒体或叶绿体基因组中，并且可以使用合适的启动子在宿主细胞中表达(例如，Mc Bride等,1995；US 5,693,507)。在一个实施方案中，本发明的核酸和/或嵌合基因可以包含用于在植物细胞或微生物细胞(例如细菌)中表达的合适启动子，其与编码本发明蛋白质的核苷酸序列可操作地连接，任选地随后是3'非翻译核苷酸序列。编码序列之前任选地有5'UTR序列。启动子、3'UTR和/或5'UTR可以例如来自天然孤雌生殖基因，或者可以来自其它来源。
如本文教导的(编码如本文教导的能够诱导孤雌生殖的蛋白质的)核酸可以稳定地插入单个植物细胞的核基因组中，并且如此转化的植物细胞可以用于产生转化的植物，该转化的植物由于在特定时间在特定细胞中存在所述蛋白质而具有改变的表型。在非限制性实例中，在根癌农杆菌(Agrobacterium tumefaciens)中的T-DNA载体(其包含如本文教导的核酸，该核酸编码如本文教导的在孤雌生殖中是功能性的蛋白质)可用于转化植物细胞，然后，可使用例如EP 0116718、EP 0270822、PCT公布WO84/02913和公布的欧洲专利申请EP0242246以及Gould等(1991)中描述的程序从转化的植物细胞再生转化的植物。用于农杆菌(Agrobacterium)介导的植物转化的T-DNA载体的构建是本领域熟知的。T-DNA载体可以是如EP0120561和EP0120515中描述的二元载体或如EP0116718中描述的可以通过同源重组整合到农杆菌Ti-质粒中的共整合载体。莴苣转化方案已在例如Michelmore等(1987)和Chupeau等(1989)中进行了描述。
优选的T-DNA载体含有与编码本发明蛋白质的核苷酸序列可操作连接的启动子；例如，启动子与SEQ ID NO：3的核苷酸序列或其变体或功能性片段可操作地连接，位于T-DNA边界序列之间，或至少位于右边界序列的左侧。优选地，所述启动子是包含核酸插入物(优选双链DNA插入物)的启动子，其中所述插入物的长度为50至2000bp，100至1900bp，200至1800bp，300至1700bp，400至1600bp，500至1500bp，600至1400bp，1000至1400，1200至1400或1300至1400bp。甚至更优选地，所述插入物的长度为约1300bp。优选地，插入物与如本文定义的孤雌生殖表型相关，并且任选地在孤雌生殖表型中是功能性的。优选地，所述插入物位于启动子序列内，所述启动子序列直接位于编码PAR蛋白质的序列的上游(3')，优选使得所述插入物的3'末端与编码PAR蛋白质的序列的起始密码子之间的距离为50-200bp，优选约50，60，70，80，90，100，110，120，130，140，150，160，170，180，190或200bp，最优选约102bp。优选地，所述插入物被定位成使得插入物的3'末端核苷酸位于与SEQ ID NO：2的核苷酸1798和/或SEQ ID NO：5的核苷酸1798的位置同源的位置。优选地，所述插入物没有开放阅读框。甚至更优选地，所述插入物是微型反向重复转座元件(MITE)或MITE样序列，其中所述MITE或MITE样序列是一种非自主元件，其特征在于含有缺乏开放阅读框的内部序列，其侧翼为末端反向重复(TIR)，而TIR侧翼为小的同向重复(靶位点重复)。关于MITE、TIR和序列的进一步描述，参考Guo et al,Scientific Reports.2017Jun 1；7(1):2634，其通过引用并入本文。所述插入物，优选所述MITE或MITE样序列，可以与SEQ ID NO：60具有至少约50％，60％，70％，80％，90％，95％，98％，99％或更高的同一性。优选地，所述插入物与如本文定义的孤雌生殖表型相关，并且任选地在孤雌生殖表型中是功能性的。在进一步优选的实施方案中，T-DNA载体包含调控序列(优选启动子序列)或由其组成，其在如上定义的位置包含所述插入物。优选地，T-DNA载体包含编码本文定义的PAR蛋白质的序列或由其组成，该序列与所述启动子序列可操作地连接，其中优选地，所述启动子序列直接位于编码PAR蛋白质的序列的上游。任选地，所述T-DNA载体可以包含一个或多个另外的转录调控序列。
Gielen等(1984)描述了边界序列。当然，使用诸如直接基因转移(如描述于例如EP0223247中)、花粉介导的转化(如描述于例如EP0270356和WO85/01856中)、原生质体转化(如描述于例如US4,684,611中)、植物RNA病毒介导的转化(如描述于例如EP0067553和US4,407,956中)、脂质体介导的转化(如描述于例如US4,536,475中)和其它方法的程序，其它类型的载体可用于转化植物细胞。
在另一个实施方案中，本发明的核酸可以通过体细胞杂交导入。体细胞杂交可以通过原生质体融合来进行(例如参见Holmes,2018)。
本发明的核酸还可以整合到基因组中，例如使用一种或多种特异性核酸内切酶(诸如CRISPR-核酸内切酶/向导RNA复合物)在基因组中的适当位点引入双链断裂，以及使用包含本发明的核酸的供体构建体整合到基因组中。本领域技术人员知晓如何设计这种CRISPR-核酸内切酶/向导RNA复合物以引入适于整合的双链断裂和供体构建体(综述参见Bortesi and Fischer,2015)。
或者，可以通过改变内源核苷酸序列来转化植物，从而例如通过随机或定向诱变将包含在植物中的一个或多个par等位基因转化为一个或多个Par等位基因。所述诱变可涉及编码序列的诱变，但也可包括调控序列(诸如启动子序列、5'UTR和/或3'UTR)的诱变。par等位基因的所述内源5'UTR启动子核苷酸序列可以被修饰以包含如本文以上所定义的插入物，优选在如本文以上所定义的位置。
同样，从转化细胞中选择和再生转化植物是本领域熟知的。显然，对于不同物种，甚至对于单一物种的不同品种或栽培种，方案特别适用于高频再生转化体。本发明还包括显示孤雌生殖并包含本发明的核酸和/或蛋白质的转化植物的后代。
除了核基因组的转化，本发明还包括质体基因组(优选叶绿体基因组)的转化。质体基因组转化的一个优点是可以降低转基因扩散的风险。质体基因组转化可以如本领域已知的方法进行，参见例如Sidorov等(1999)或Lutz等(2004)。
所得转化植物可用于常规植物育种方案中以产生更多含有转基因的转化植物。单拷贝转化体可以使用例如Southern印迹分析或基于PCR的方法或
Technology测定(Third Wave Technologies,Inc.)选择。通过本发明的核酸或蛋白质和/或嵌合基因的存在，转化的细胞和植物可以容易地与未转化的细胞和植物区分开来。还可以对转基因插入位点侧翼的植物DNA序列进行测序，由此可以开发“事件特异性”检测方法用于常规用途。参见例如WO0141558，其描述了基于例如整合序列和侧翼(基因组)序列的优良事件检测试剂盒(例如PCR检测试剂盒)。
本发明的核酸可以插入植物细胞基因组中，使得插入的编码序列在启动子的下游(即3')并在其控制下，所述启动子可以指导在植物细胞中的表达。这优选通过在植物细胞基因组中(特别是在细胞核或质体(例如叶绿体)基因组中)插入包含这些元件的嵌合基因来实现。
启动子(其与SEQ ID NO：3或其变体或片段可操作地连接)可以例如是组成型活性启动子，诸如：分离株CM1841(Gardner等,1981)、CabbB-S(Franck等,1980)和CabbB-JI(Hull and Howell,1987)的花椰菜花叶病毒(CaMV)的强组成型35S启动子或增强的35S启动子(“35S启动子”)；Odell等(1985)或US5164316中描述的35S启动子，来自泛素家族的启动子(例如玉米泛素启动子，Christensen等,1992；EP 0342926；也参见Cornejo等,1993)，gos2启动子(de Pater等,1992)，emu启动子(Last等,1990)，拟南芥肌动蛋白质启动子诸如An等(1996)描述的启动子，水稻肌动蛋白质启动子诸如Zhang等(1991)描述的启动子以及US 5,641,876中描述的启动子或如WO070067中描述的水稻肌动蛋白质2启动子；木薯脉花叶病毒(Cassava vein mosaic virus)的启动子(WO97/48819,Verdaguer等1998)，地三叶草矮化病毒(Subterranean Clover Stunt Virus)的pPLEX系列启动子(WO96/06932，特别是S7启动子)，醇脱氢酶启动子，例如pAdh1S(GenBank登录号X04049、X00581)，以及TR1'启动子和TR2'启动子(分别为“TR1'启动子”和“TR2'启动子”)，它们分别驱动T-DNA的1'和2'基因的表达(Velten等,1984)，US6051753和EP426641中描述的玄参花叶病毒(FigwortMosaic Virus)启动子，组蛋白质基因启动子诸如拟南芥的Ph4a748启动子(PMB 8:179-191)等。
或者，可以使用启动子，其不是组成型的，而是对植物的一种或多种组织或器官具有特异性(组织优选/组织特异性，包括发育调控的启动子)，例如卵细胞特异性启动子，由此本发明的蛋白质仅或优先在特定组织或器官的细胞中和/或仅在特定发育阶段期间表达。
由于本发明蛋白质的组成型生产在植物的适合度方面具有高成本，因此在一个实施方案中优选使用其活性是可诱导的启动子。诱导型启动子的实例是创伤诱导型启动子，诸如Cordera等(1994)描述的MPI启动子，其由创伤(诸如由昆虫或物理伤害引起)诱导，或COMPTII启动子(WO0056897)或US6031151中描述的PR1启动子。或者，启动子可被化学物质诱导，诸如Aoyama and Chua(1997)和US6063985中描述的地塞米松或四环素(TOPFREE或TOP 10启动子，参见Gatz,1997和Love等,2000)。
词语“诱导型”不一定要求启动子在没有诱导物刺激时完全失活。可以存在低水平的非特异性活性，只要这不会导致植物的严重产量或质量损失。因此，诱导型优选是指启动子活性的增加，导致与诱导物接触后编码本发明蛋白质的下游编码区的转录增加。
在一个实施方案中，使用天然孤雌生殖基因的启动子。例如，蒲公英Par或par等位基因的启动子可以被分离并与编码本发明蛋白质的编码区可操作地连接。在一个实施方案中，所述启动子(上游转录调控区，例如在翻译起始密码子和/或转录起始密码子上游约2000bp内)可以使用已知方法(诸如TAIL-PCR(Liu等,1995；Liu等,2005)、Linker-PCR或反向PCR(IPCR))从无融合生殖植物和/或其他植物分离。
在一个实施方案中，使用天然孤雌生殖基因的启动子或由其衍生的启动子。例如，可以使用源自SEQ ID NO：2或其变体或片段的启动子。优选地，所述启动子为包含核酸插入物(优选双链DNA插入物)的启动子，其中所述插入物的长度为50至2000bp，100至1900bp，200至1800bp，300至1700bp，400至1600bp，500至1500bp，600至1400bp，1000至1400，1200至1400或1300至1400bp。甚至更优选地，所述插入物的长度为约1300bp。优选地，插入物与如本文定义的孤雌生殖表型相关，并且任选地在孤雌生殖表型中是功能性的。优选地，所述插入物位于启动子序列内，该启动子序列直接位于编码PAR蛋白质的序列的上游(3')，优选使得所述插入物的3'末端与编码PAR蛋白质的序列的起始密码子之间的距离为50-200bp，优选约50，60，70，80，90，100，110，120，130，140，150，160，170，180，190或200bp，最优选约102bp。优选地，所述插入物被定位为使得插入物的3'末端核苷酸位于与SEQ ID NO：2的核苷酸1798和/或SEQ ID NO：5的核苷酸1798的位置同源的位置。优选地，所述插入物没有开放阅读框。甚至更优选地，所述插入物是微型反向重复转座元件(MITE)或MITE样序列，其中所述MITE或MITE样序列是一种非自主元件，其特征在于含有缺乏开放阅读框的内部序列，其侧翼为末端反向重复(TIR)，而TIR侧翼为小的同向重复(靶位点重复)。关于MITE、TIR和序列的进一步描述，参考Guo等,Scientific Reports.2017Jun 1；7(1):2634，其通过引用并入本文。所述插入物，优选所述MITE或MITE样序列，可以与SEQ ID NO：60具有至少约50％，60％，70％，80％，90％，95％，98％，99％或更高的同一性。优选地，所述插入物与如本文定义的孤雌生殖表型相关，并且任选地在孤雌生殖表型中是功能性的。启动子可以具有SEQ ID NO：2的核苷酸序列。也可以使用比本文提及的序列更长的序列。编码区翻译起始密码子上游至多约2000bp的区域可包含转录调控元件(即启动子)。因此，在一个实施方案中，编码本发明蛋白质的序列的翻译起始密码子上游2000bp，1500bp，1000bp，800bp，500bp，300bp或更少的核苷酸序列是分离的，可以测试启动子活性，并且如果是功能性的，该序列可以与编码本发明蛋白质的序列可操作地连接，如本文所教导的。可以通过例如缺失分析测试整个序列及其片段的启动子活性，其中5'和/或3'部分被缺失，并且使用已知方法(例如将启动子或片段与报告基因可操作地连接)测试启动子活性。
本文教导的编码序列优选插入植物基因组，使得编码序列在合适的3'端非翻译区(“3'末端”或3'UTR)的上游(即5')。合适的3'末端包括CaMV35S基因(“3'35S”)，胭脂碱合酶基因(“3'nos”)(Depicker等,1982)，章鱼碱合酶基因(“3'ocs”)(Gielen等,1984)和T-DNA基因7(“3'基因7”)(Velten和Schell,1985)的那些，它们在转化的植物细胞中充当3'-非翻译DNA序列，等等。在一个实施方案中，使用天然孤雌生殖基因的3'UTR或由其衍生的3'UTR。例如，可以使用源自SEQ ID NO：4或其变体或片段的任何3'UTR。3'UTR可具有SEQ ID NO：4的核苷酸序列。
在一个实施方案中，具有SEQ ID NO：2的核苷酸序列或其变体和/或片段的启动子可以与编码本发明蛋白质的核酸可操作地连接，优选地，编码蛋白质的核苷酸序列能够诱导如本文所教导的孤雌生殖，更优选地具有SEQ ID NO：1的氨基酸序列或其变体和/或片段。优选地，所述启动子和编码序列进一步与SEQ ID NO：4或其变体和/或片段的3'UTR可操作地连接。
将T-DNA载体引入农杆菌可使用已知方法(诸如电穿孔或三亲本杂交(triparental mating))进行。
如本文所教导的编码序列可以任选地作为杂合基因序列插入植物基因组中，由此编码序列与编码可选择的或可评分的标记的基因(US 5,254,799；Vaeck等,1987)(诸如例如编码卡那霉素抗性的neo(或nptII)基因(EP0242236))框内连接(linked in-frame)，使得植物表达易于检测的融合蛋白质。
编码本发明蛋白质的序列的全部或部分也可用于转化微生物，诸如细菌(例如大肠杆菌(Escherichia coli)、假单胞菌(Pseudomonas)、农杆菌、芽孢杆菌(Bacillus)等)、真菌或藻类或昆虫，或用于制备重组病毒。这特别适合于蛋白质(优选分离的蛋白质)的生产和随后的纯化。细菌(其具有本文教导的全部或部分编码序列，并入合适的克隆载体中)的转化可以以常规方式进行，优选使用Maillon等(1989)和WO 90/06999中描述的常规电穿孔技术进行。为了在原核宿主细胞中表达，可相应地优化核苷酸序列的密码子使用(如本文对植物所述)。应当除去内含子序列，并且可以如已知的那样进行用于最佳表达的其它调整。包含本发明的核酸和/或表达本发明的蛋白质的这种原核宿主细胞包括在本发明中。这种宿主细胞可用于产生本发明的蛋白质和/或核酸。
本发明核酸的DNA序列可以以翻译中性方式进一步改变，以修饰基因部分中可能存在的抑制性DNA序列和/或通过引入密码子选择的变化，例如，使密码子选择适应植物最优选，优选特定相关植物属最优选的密码子选择，例如如本文所述的宿主植物。
根据本发明的一个实施方案，本发明的蛋白质靶向细胞内细胞器诸如质体，优选叶绿体，线粒体，或从细胞分泌，潜在地优化蛋白质稳定性和/或表达。类似地，该蛋白质可以靶向液泡。为此，在本发明的一个实施方案中，本发明的嵌合基因包含编码信号或靶向肽的编码区，其与编码本发明蛋白质的区域连接。包括在本发明蛋白质中的特别优选的肽是用于叶绿体或其它质体靶向的转运肽，尤其是来自植物基因的重复转运肽区域，其基因产物靶向质体，Capellades等(US 5,635,618)的优化转运肽，菠菜的铁氧还蛋白质-NADP+氧化还原酶的转运肽(Oelmuller等,1993)，Wong等(1992)中描述的转运肽以及已公开的PCT专利申请WO00/26371中的靶向肽。还优选的是在细胞外信号传导与这种肽连接的蛋白质的分泌的肽，诸如马铃薯蛋白质酶抑制剂II的分泌信号(Keil等,1986)，水稻α-淀粉酶3基因的分泌信号(Sutliff等,1991)和烟草PR1蛋白质的分泌信号(Cornelissen等,1986)。根据本发明特别有用的信号肽包括叶绿体转运肽(例如Van Den Broeck等,1985)，或US 5,510,471和US 5,635,618的优化的叶绿体转运肽(其使蛋白质转运至叶绿体)，分泌信号肽或将蛋白质靶向其他质体、线粒体、ER或另一细胞器的肽。靶向细胞内细胞器或植物细胞外分泌或细胞壁的信号序列存在于天然靶向或分泌的蛋白质中，优选
等(1989)，
and Weil(1991)，Neuhaus&Rogers(1998)，Bih等(1999)，Morris等(1999)，Hesse等(1989)，Tavladoraki等(1998)，Terashima等(1999)，Park等(1997)，Shcherban等(1995)描述的那些。
在一个实施方案中，如本文教导的本发明的蛋白质与其它蛋白质(其任选地在不同启动子的控制下在单一宿主中控制(优选增强或诱导)孤雌生殖、不完全减数分裂或无融合生殖)共表达。这种其它基因可以是赋予不完全减数分裂的基因，诸如例如WO2017/039452A1(其通过引用并入本文)中所述的二倍体孢子生殖。
在另一个实施方案中，本发明的蛋白质被渗入种质中，该种质优选包含其它目的基因，诸如赋予不完全减数分裂的基因(例如用于二倍体孢子生殖的基因)。通过杂交和选择，产生杂交体，其中若干目的基因堆叠。
通过转化已经表达本发明蛋白质的植物，或通过与用本发明不同核酸转化的植物杂交，可以容易地获得共表达宿主植物。应当理解，不同的蛋白质可在同一植物中表达，或各自可在单一植物中表达，然后通过使单一植物彼此杂交而在同一植物中组合。例如，在杂交种子生产中，每个亲本植物可以表达需要共表达的每种蛋白质。在杂交亲本植物以产生杂种后，两种蛋白质在杂种植物中组合。本发明包括包含这两种基因和/或表达这两种蛋白质的这种杂种或其后代。
优选地，出于选择目的以及杂草控制选择，本发明的转基因植物还用编码赋予除草剂(诸如广谱除草剂，例如基于草铵膦铵作为活性成分的除草剂(例如
或BASTA；抗性由PAT或bar基因赋予；参见EP 0 242 236和EP 0 242 246)或草甘膦(例如
抗性由EPSPS基因赋予，参见例如EP0 508 909和EP 0 507 698))抗性的蛋白质的DNA转化。使用除草剂抗性基因(或赋予所需表型的其它基因)作为可选择的标记还具有可以避免引入抗生素抗性基因的优点。
可选地或另外，可使用其它可选择的标记基因，诸如抗生素抗性基因。由于通常不接受在转化的宿主植物中保留抗生素抗性基因，因此在选择转化体后可以再次去除这些基因。存在用于去除转基因的不同技术。实现去除的一种方法是在转基因侧翼添加lox位点，并在选择后，将转化的植物与表达CRE重组酶的植物杂交(参见例如EP506763B1)。位点特异性重组导致标记基因的切除。另一种位点特异性重组系统是EP686191和US5527695中描述的FLP/FRT系统。位点特异性重组系统诸如CRE/LOX和FLP/FRT也可用于基因堆叠目的。此外，已经描述了单组件切除系统，参见例如WO9737012或WO9500555)。
优选地，本发明的核酸用于产生具有增强的孤雌生殖表型的转基因植物细胞、植物、植物种子等及其任何衍生物/后代。具有增强孤雌生殖的转基因植物可以通过用本发明的核酸(其优选地编码具有SEQ ID NO：1的氨基酸序列或其变体和/或片段的蛋白质，在合适启动子(如本文所述)的控制下)转化植物宿主细胞，并从所述细胞再生转基因植物来产生。优选地，与未转化的或空载体对照相比，本发明的转基因植物包含增强的孤雌生殖。因此，例如提供了包含增强的孤雌生殖的转基因莴苣植物。因此，与未转化的或空载体转化的对照相比，如果表达本发明蛋白质的转化植物显示孤雌生殖显著增加，则其显示增强的孤雌生殖。增强的孤雌生殖表型可以通过表达合适量的能够在合适的时间和/或位置诱导孤雌生殖的本发明的蛋白质来微调。这种微调可以通过确定最合适的启动子和/或通过选择显示所需表达水平的转基因“事件”来进行。
表达所需水平的本发明蛋白质和/或包含所需或所需水平的本发明核酸的转化体、杂种或近交系通过例如分析拷贝数(Southern印迹分析)、mRNA转录水平(例如使用能够扩增本发明蛋白质的引物对或侧翼引物的RT-PCR)或通过分析各种组织中孤雌生殖蛋白质的存在和水平(例如SDS-PAGE；ELISA测定等)来选择。可以选择单拷贝转化体，例如出于调控原因，并分析转基因插入位点侧翼的序列，优选测序以表征“事件”。选择导致本发明蛋白质高或中等表达的转基因事件用于进一步开发，直到获得具有稳定转基因的高性能优良事件。
表达本发明的蛋白质和/或包含本发明的核酸的转化体还可以包含其它转基因，诸如赋予疾病抗性或赋予对其它生物和/或非生物胁迫的耐受性或赋予二倍体孢子生殖的其它基因。为了获得这种具有“堆叠”转基因的植物，可以将其它转基因引入所述转化体，或者可以随后用一种或多种其它基因转化所述转化体，或者可以用几种嵌合基因转化植物系或变种。例如，几种转基因可以存在于单个载体上，或者可以存在于共转化的不同载体上。
在一个实施方案中，以下基因与本发明的核酸组合：已知的疾病抗性基因，特别是赋予对坏死性病原体的抗性增强的基因，病毒抗性基因，昆虫抗性基因，非生物胁迫抗性基因(例如干旱耐受性、盐耐受性、热或冷耐受性等)，除草剂抗性基因等。因此，堆叠的转化体可以对病原体抗性、昆虫抗性、线虫抗性、盐度、冷胁迫、热胁迫、水胁迫等具有更广泛的生物和/或非生物胁迫耐受性。此外，沉默方法可与单一植物中的表达方法组合，例如Par等位基因的沉默可与par等位基因的表达组合，或反之亦然。
任选地，本发明的核酸可用于抑制孤雌生殖，例如通过沉默、敲低或降低植物或植物细胞中一个或多个Par等位基因上的孤雌生殖基因的表达。这可以通过修饰存在于所述植物或植物细胞中的Par等位基因的编码序列或一个或多个调控序列(例如启动子序列)，或通过引入Par等位基因的RNAi靶向转录物来进行。因此，本发明还提供了一种减少或消除植物或植物细胞中孤雌生殖的方法，其包含以下步骤：
a)在一种或多种植物细胞中减少或消除能够诱导孤雌生殖和/或在孤雌生殖中是功能性的核酸的表达；
b)选择其中所述表达减少或消除的植物细胞；以及
c)从所述植物细胞再生植物。
所述核酸优选是包含或由SEQ ID NO：2-5中的任一个及其变体和/或片段组成的核酸，和/或编码SEQ ID NO：1的蛋白质的核酸，和/或其变体或片段。
本文所述的任何转化植物的完整植物、植物部分(例如种子、细胞、组织)和植物产品(例如果实)和后代均包括在本文中，并且可以通过转基因的存在来鉴定，例如通过使用总基因组DNA作为模板并使用对孤雌生殖基因特异的PCR引物对的PCR分析和/或通过使用基因组变异分析(诸如但不限于基于序列的基因分型(Sequence Based Genotyping，SBG)或
SNPSelect分析。也可以开发“事件特异性”PCR诊断方法，其中PCR引物基于插入的转基因侧翼的植物DNA，参见US6563026。类似地，可以开发事件特异性AFLP指纹或RFLP指纹，其鉴定转基因植物或由其衍生的任何植物、种子、组织或细胞。
应当理解，根据本发明的转基因植物优选不显示不希望的表型，诸如产量降低，对疾病(特别是对necrotrophs)的易感性增加或不希望的结构改变(矮化、变形)等，并且如果在初级转化体中观察到这样的表型，则可以通过常规方法除去。本文所述的任何转基因植物对于转基因可以是杂合的、纯合的或半合的。
本发明还涉及通过本文详述的方法获得或可获得的植物、种子、植物部分(例如植物细胞)和植物产品，优选包含本发明的蛋白质、本发明的核酸和/或本发明的构建体。优选地，所述蛋白质、核酸和/或构建体能够诱导孤雌生殖和/或在孤雌生殖中是功能性的，如本文详述的。本发明的植物优选是本文中作为合适的宿主植物列出的物种。这种方法包括将本发明的核酸从植物渗入后代，和/或用本发明的核酸作为转基因转化植物细胞，以及随后从所述植物细胞再生植物。优选地，植物、植物部分和/或植物产品不是Taraxacumofficinale sensu lato物种，其包含本发明的核酸，其中所述植物或植物细胞优选是本文列出的作为合适的宿主植物的物种，优选来自选自十字花科、葫芦科、豆科、禾本科、茄科和菊科(Asteraceae)(Compositae)的家族。
优选地，植物、植物部分和/或植物产品包含通过遗传修饰或通过基因渗入的本发明的核酸，其中优选地，所述核酸被整合至其基因组中。优选地，所述植物、植物部分和/或植物产品能够孤雌生殖和/或在孤雌生殖中是功能性的。甚至更优选地，所述植物、植物部分和/或植物产品进一步能够进行不完全减数分裂。本发明提供了本发明的植物或植物细胞的种子、植物部分或植物产品。
本发明还涉及源自本发明植物的植物部分和植物产品，其中该植物部分和/或植物产品包含如本文定义的本发明的蛋白质、如本文定义的本发明的核酸和/或如本文定义的本发明的构建体，其可以是如本文所定义的片段，其允许评估这种蛋白质、核酸或构建体在植物产品的植物部分所源自的植物中的存在。这种部分和/或产品可以是种子或果实和/或由其衍生的产品(例如糖或蛋白质)。这种部分、产品和/或由其衍生的产品可以是不繁殖的材料。
任何植物均可为合适的宿主，但最优选地，宿主植物物种应当是将受益于增强的或降低的孤雌生殖的植物物种。合适的宿主包括任何植物物种。特别地，优选具有良好农学特性的栽培种或育种品系。本领域技术人员知晓如何通过产生转基因植物和评估孤雌生殖，以及合适的对照植物，来测试本文教导的核酸和/或蛋白质，和/或其变体或片段是否可以赋予宿主植物所需的孤雌生殖的增加或减少。
合适的宿主植物包括例如属于十字花科、葫芦科、豆科、禾本科(Gramineae)、茄科、菊科(Compositae)、蔷薇科或禾本科(Poaceae)的宿主。
在一个优选的实施方案中，宿主植物可以是选自蒲公英属、莴苣属、豌豆属、辣椒属、茄属、黄瓜属、玉蜀黍属、棉属、大豆属(Glycine)、小麦属、稻属和高粱属的植物物种。
在一个优选的实施方案中，如本文教导的植物、植物部分、植物细胞或种子来自选自以下的物种：蒲公英属、莴苣属、豌豆属、辣椒属、茄属、黄瓜属、玉蜀黍属、棉属、大豆属、小麦属、稻属、葱属、芸苔属、向日葵属、甜菜属、菊苣属、菊属、狼尾草属、黑麦属、大麦属、苜蓿属、菜豆属、蔷薇属、百合属、咖啡属、亚麻属、大麻属、木薯属、胡萝卜属、南瓜属、西瓜属和高粱属。
合适的宿主植物包括例如玉米(maize)/玉米(corn)(玉蜀黍属物种)，小麦(小麦属物种)，大麦(例如Hordeum vulgare)，燕麦(例如Avena sativa)，高粱(Sorghumbicolor)，黑麦(Secale cereale)，大豆(大豆属物种，例如大豆(G.max))，棉花(棉属物种，例如陆地棉(G.hirsutum)，海岛棉(G.barbadense))，芸苔属物种(例如欧洲油菜(B.napus)，印度芥菜(B.juncea)，甘蓝(B.oleracea)，芜菁(B.rapa)等)，向日葵(Helianthus annus)，红花，山药，木薯，苜蓿(Medicago sativa)，水稻(稻属物种，例如籼稻(O.sativa indica)品种群或粳稻(japonica)品种群)，饲草，珍珠稷(狼尾草物种，例如蜡烛稗(P.glaucum))，树种(松属，杨树，冷杉，车前草等)，茶，咖啡，油棕，椰子，蔬菜物种，诸如豌豆，西葫芦，豆类(例如菜豆属物种)，辣椒，黄瓜，朝鲜蓟，芦笋，茄子，西兰花，大蒜，韭菜，莴苣，洋葱，萝卜，芜菁，马铃薯，抱子甘蓝，胡萝卜，花椰菜，菊苣，芹菜，菠菜，菊苣，茴香，甜菜，肉质果实植物(葡萄，桃子，李子，草莓，芒果，苹果，李子，樱桃，杏，香蕉，黑莓，蓝莓，柑橘，猕猴桃，无花果，柠檬，酸橙，油桃，覆盆子，西瓜，橙子，葡萄柚等)，观赏植物(例如玫瑰，矮牵牛，菊花，百合，非洲菊物种)，草本植物(薄荷，欧芹，罗勒，百里香等)，木本树(例如杨属、柳属、栎属、桉属的物种)，纤维物种例如亚麻(Linum usitatissimum)和大麻(Cannabis sativa)。
一种或多种Par等位基因的标记辅助选择和转移或组合
本发明的核酸可用作遗传标记，用于标记辅助选择蒲公英属物种和/或其他植物物种的Par或par等位基因以及用于将不同或相同Par或par等位基因转移和/或组合到/在目的植物中和/或在可用于与其中发现Par或par等位基因(或变体)的植物产生种内或种间杂种的植物中。
可以基于这些序列开发许多不同的标记测定。标记测定的开发通常涉及Par和par等位基因之间的多态性的鉴定，使得多态性是“标记”特定等位基因的遗传标记。然后将多态性用于标记测定。例如，如本文所教导的Par等位基因的序列与孤雌生殖的存在或增强相关。这例如通过针对如本文教导的Par或par等位基因的核苷酸序列(的一部分)筛选孤雌生殖的植物材料和/或非孤雌生殖的植物材料以使特定等位基因与孤雌生殖或非孤雌生殖相关来进行。因此，可以产生PCR引物或探针，其检测从(非)孤雌生殖的植物材料获得的样品(例如RNA、cDNA或基因组DNA样品)中的此类核苷酸序列。比较序列或其部分，并鉴定与孤雌生殖相关的多态性标记。然后可以将多态性标记，例如与Par或par等位基因连接的SNP标记开发成快速分子测定，用于筛选植物材料中孤雌生殖等位基因的存在与否。因此，这些“遗传标记”的存在与否指示与其连接的Par或par等位基因的存在，并且可以用遗传标记的检测代替Par或par等位基因的检测。
优选地，使用容易且快速的标记测定，其能够快速检测样品(例如DNA样品)中的Par或par等位基因或等位基因组合。因此，在一个实施方案中，本文提供了本发明的核酸在用于确定样品中Par或par等位基因的存在与否，和/或用于确定该等位基因的纯合性或杂合性的分子测定中的用途。
这种测定可以例如包括以下步骤：
(a)提供孤雌生殖和非孤雌生殖的植物材料和/或其核酸样品；
(b)确定(a)的所述材料中全部或部分本发明核酸的核苷酸序列。
一方面，提供了用于检测本发明核酸或相关或衍生的RNA序列(诸如转录物)的PCR引物和/或探针、分子标记和试剂盒。从样品中扩增本发明核酸的简并或特异性PCR引物对可基于如本文教导的核苷酸序列或其变体合成，如本领域已知的(参见Dieffenbach和Dveksler,1995；以及McPherson等,2000)。例如，该序列(或互补链)的任何一段9，10，11，12，13，14，15，16，18或更多个连续核苷酸可用作引物或探针。
同样，包含本文教导的Par或par等位基因序列或其互补序列的DNA片段可用作杂交探针。本文提供的检测试剂盒可以包含Par(等位基因-)特异性引物和/或Par(等位基因-)特异性探针，以及使用引物或探针检测样品中本发明核酸的相关方案。这种检测试剂盒可以例如用于确定植物是否已经用本发明的核酸转化，或用于筛选蒲公英属种质和/或其它植物物种种质中Par等位基因的存在以及任选地接合性测定。
因此，在一个实施方案中，提供了一种检测植物组织(例如蒲公英组织)或其核酸样品中编码本发明蛋白质的核苷酸序列存在与否的方法。该方法可包含：
a)从一种或多种植物获得植物组织样品，或其核酸样品，
b)使用分子标记测定分析核酸样品中是否存在一种或多种与Par等位基因连接的标记，其中该标记测定检测与孤雌生殖相关的本发明核酸的存在，以及任选地
c)选择包含一种或多种所述标记的植物用于进一步使用。
可选地或另外地，该方法可包含：
a)从一种或多种植物获得植物组织样品，或其核酸样品，
b)使用分子标记测定分析核酸样品中是否存在一种或多种与par等位基因连接的标记，其中该标记测定检测与非孤雌生殖相关的本发明核酸的存在，以及任选地
c)选择包含一种或多种所述标记的植物用于进一步使用。
优选地，在任何这些方法中使用的一种或多种植物是适合作为本文进一步定义的宿主植物的植物。
孤雌生殖的应用
本发明的核酸和/或蛋白质可用于筛选(例如用于植物或植物细胞中的一个或多个孤雌生殖基因座)、基因分型、赋予孤雌生殖、赋予无融合生殖以增加倍性和/或用于产生双单倍体。优选地，所述用途是在植物生物技术和/或育种中，即在植物或植物细胞中/上。
孤雌生殖是无融合生殖的一项要素，并且孤雌生殖的基因可以与不完全减数分裂(例如二倍体孢子生殖)的基因组合使用以产生无融合生殖，优选将其用于本文列出的应用。这些基因可以通过转化、基因渗入或通过修饰内源合适的基因从而将它们转化为无融合生殖(或二倍体孢子生殖)基因而引入有性作物中。对无融合生殖基因的结构和功能的了解也可用于修饰内源性有性生殖基因，使它们成为无融合生殖基因。优选的用途是将无融合生殖基因置于诱导型启动子下，使得当有性繁殖产生新基因型时可以关闭无融合生殖，而当需要无融合生殖繁殖优良基因型时可以开启无融合生殖。
核酸或其衍生产物可用作无融合生殖的组分。不完全减数分裂和孤雌生殖均为功能性配子体无融合生殖所必需的。可以通过影响减数分裂的突变的组合实现不完全减数分裂(Crismani等,2013)，结果是大孢子中染色体不减少，即有丝分裂而不是减数分裂。通过表观遗传改变呈现配子体命运的体细胞(Grimanelli,2012)也会导致未减数的孢子样细胞，其可能产生未减数的配子(卵细胞)。在另一个实施方案中，通过天然不完全减数分裂基因的转基因或非转基因表达实现不完全减数分裂。无论以何种方式形成未减数卵细胞，能够诱导孤雌生殖的本发明核酸的适当的时间和空间表达可以诱导卵细胞在不受精的情况下表现为受精卵并分裂。
孤雌生殖基因可以以全新的方式使用，例如不直接作为无融合生殖中的工具。例如，尽管在无融合生殖中孤雌生殖和不完全减数分裂在单个植物中组合，但是在一代中使用不完全减数分裂和在下一代中使用孤雌生殖将在二倍体和多倍体水平上将作物的有性基因库联系起来，通过不完全减数分裂提高倍性水平，通过孤雌生殖降低倍性水平。这是非常有用的，因为多倍体群体可以更适合突变诱导，因为它们可以耐受更多的突变。多倍体植物也可以更有活力。但二倍体群体更适合选择，二倍体杂交更适于遗传定位、BAC文库的构建等也较好。多倍体的孤雌生殖可产生单倍体，其可与二倍体杂交。二倍体中的二倍体孢子生殖产生未减数的2n个卵细胞，其可通过来自多倍体的花粉受精以产生多倍体后代。因此，不同育种世代中不完全减数分裂和孤雌生殖的交替将二倍体和多倍体基因库联系起来。
没有不完全减数分裂的核酸衍生产物(转录物或编码的蛋白质)的另一用途是产生单倍体后代，其可用于产生单倍体和通过双单倍体(DH)的基因组加倍(例如自发基因组加倍，秋水仙碱，叠氮化钠或其它化学品)。双单倍体可用作亲本以产生有性F1杂种。双单倍体是使植物纯合的最快方法。双单倍体植物可以是纯合的，而第二种最快的方法是自交，在二倍体植物中达到足够高水平的纯合性需要5-7代。有几种产生双单倍体的方法。在一些植物物种中，单倍体可以通过小孢子培养产生。其它方法是通过用辐射花粉(甜瓜)授粉产生单倍体胚(雌核发育)，或用特定传粉者原种(玉米，马铃薯)授粉。这些方法具有其局限性，诸如成本、基因型的不适应性、劳动强度等。在一些作物中，不存在用于单倍体生产的方法(例如番茄)。孤雌生殖基因的显性等位基因可显著增加雌核发育频率，从而降低单倍体生产的成本。
以下非限制性实施例说明了本发明的不同实施方案。除非在实施例中另有说明，否则所有重组DNA技术均根据Sambrook等(1989)，Sambrook and Russell(2001)；以及Ausubel等(1994)的第1卷和第2卷所述的标准方案进行。用于植物分子工作的标准材料和方法描述于R.D.D.Croy的Plant Molecular Biology Labfax(1993)，由BIOS ScientificPublications Ltd(UK)和Blackwell Scientific Publications,UK联合出版。
表1：本文所用SEQ ID NO的概述
表2：编码Cas9/gRNA-1或Cas9/gRNA-2的T-DNA构建体对种子表型、Par等位基因的影响，更特别是对Par等位基因(SEQ ID NO：23)的核苷酸325-360的片段和所编码的氨基酸片段的影响。
附图说明
图1：来自对照植物(其显示野生型序列(SEQ ID NO：23))和来自包含编码Cas9/RNA-1复合物的载体的转基因植物(其显示修饰序列(SEQ ID NO：24-27))的扩增子的编码序列(Par等位基因编码序列的核苷酸325-360)和编码的氨基酸的多序列比对。向导RNA-1的基因特异性部分用方框表示。修饰以粗体和下划线表示。出于比对的原因，野生型序列包含间隔(-)。
图2：发芽实验。顶行；A68对照，发芽的正常有活力的黑色种子。中行；在基因164中具有3bp缺失的植物pKG10821-6的无活力、浅灰色、不发芽的种子。底行；用FCH72单倍体花粉授粉的植物pKG10821-6的所有四倍体、发芽和有活力的后代。每个培养皿上的种子来自单个种子头。
图3：转基因莴苣品系去雄后75小时，具有胚的透明胚珠(cleared ovule)的实例，该品系含有由拟南芥EC1.1启动子驱动的药用蒲公英Par等位基因。在发现这种胚的情况下，在观察的总和中带上该胚，如表3所示。
图4：转基因莴苣品系去雄后75小时，透明胚珠中的多胚的实例，该品系含有由拟南芥EC1.1启动子驱动的药用蒲公英Par等位基因。每个星号代表胚。
图5：APO、PAR和SEX植物中的Par基因表达的分析。
实施例
实施例1
材料和方法
植物材料
野生型无融合生殖三倍体药用蒲公英A68和有性二倍体药用蒲公英FCH72。
DNA构建体
构建在T-DNA区域编码以下成分的二元载体；欧芹泛素启动子(SEQ ID NO：16)，其驱动Cas9基因(SEQ ID NO：17)，具有35S终止子，和番茄U6启动子(SEQ ID NO：18，Nekrasov等,2013)，其驱动向导RNA-1(具有SEQ ID NO：19的靶特异性序列)，具有TTTTTT终止子序列，以及草铵膦抗性基因用于选择。构建类似的二元载体，其中将向导RNA-1的序列替换为向导RNA-2的序列(具有SEQ ID NO：20的靶特异性序列)。产生这种二元载体的合适技术是
Golden Gate或Gibson
(例如，参见Ma等,2015)。在T-DNA区域编码35S-GUS的载体用作对照构建体。
植物转化方法
农杆菌转化根据Oscarsson(Oscarsson,Lotta."Production of rubber fromdandelion-a proof of concept for a new method of cultivation."2015)的方案的修改版进行。用于植物转化的起始材料是药用蒲公英A68外植体，其从在含有0.8％琼脂的半强度MS20培养基上生长的体外传代培养繁殖的种子衍生植物获得。将50ml在含有二元载体的根癌农杆菌(放射型根瘤菌(Rhizobium radiobacter))诸如菌株C58C1的LB培养基中的过夜培养物以10x稀释度(重悬并在液体MS20中稀释)用于共培养。将外植体切成大约0.5cm2的片并共培养2-3天。接着，将外植体移至愈伤组织诱导培养基(CIM；20g l-1蔗糖，4.4g l-1具有微量和常量营养素的MS，8g l-1琼脂，1mgl-1BAP，0.2mgl-1IAA，3mgl-1草铵膦用于植物选择，100mgl-1万古霉素和100mgl-1头孢噻肟，pH 5.8)。每周将外植体转移到新鲜CIM中。当出现愈伤组织时，将其转移至芽诱导培养基(SIM；20g l-1蔗糖，4.4g l-1具有微量和常量营养素的MS，8g l-1琼脂，2mg l-1玉米素，0.1mg l-1IAA，0.05mg l-1GA3，3mg l-1草铵膦用于植物选择，100mgl-1万古霉素和100mgl-1头孢噻肟，pH 5.8)。最后，形成的直径几厘米的枝条在生根培养基(RM；20g l-1蔗糖，2.2g l-1具有微量和常量营养物的MS，8g l-1琼脂，100mg l-1万古霉素和100mg l-1头孢噻肟，pH 5.8)中生根。将生根的芽在盆栽土壤中转移至温室中。
结果
通过PCR对从农杆菌转化获得的生根植物进行基因分型，以确定植物基因组中编码Cas9和向导RNA-1或向导RNA-2的相应T-DNA的存在。对该试验呈阳性的植物(在本文中表示为转基因植物)生长直至结实。源自包含这些构建体的任一种的单个愈伤组织的单个转基因植物具有正常的有活力的深黑灰色种子，并且这种植物中的一些具有异常的浅灰色种子(参见表2)。这些浅灰色种子被发现是空的，缺乏胚，并且被发现是无活力的并且不发芽。对照植物(T-DNA阴性或用35S-GUS对照构建体转化)从未具有类似的异常浅灰色种子，对照植物均具有正常的种子头(具有可育的黑色灰色种子)。接下来，通过Illumina MiSeqSystem上向导RNA-1靶向基因组DNA区域的扩增子测序对所有转基因植物进行基因分型。发现所有出现异常浅灰色种子的转基因植物在孤雌生殖基因中均有小的缺失或插入，更特别是在由gRNA-1靶向的DNA的片段内。A68是三倍体植物。鉴定了该基因在其它两个等位基因上的序列，并在本文中由SEQ ID NO：10和15表示。这两个等位基因的序列缺乏Cas9/向导RNA诱导DSB所需的PAM序列。
具有正常黑色种子的转基因植物的基因序列没有发生变化。表2总结了观察到的小缺失或插入以及对编码序列翻译为蛋白质序列的影响，图1显示了扩增子的多序列比对。
对于在SEQ ID NO：5的基因中具有小缺失的转基因植物所观察到的结实被解释为无融合生殖表型丧失(在本文中称为无融合生殖丧失或LoA)，以及孤雌生殖表型丧失(孤雌生殖丧失或LoP)的指示。无融合生殖植物总是携带显性Par等位基因。
在没有异花传粉的情况下，三倍体蒲公英属的高结实是无融合生殖的清楚指示。作为另一种解释，可以排除自交，因为由于三倍体雄性和雌性减数分裂不平衡，有性产生的卵细胞和花粉粒的生育力非常低。Par等位基因的缺失导致LoP并因此导致LoA。然而，LoA也可由其它发育过程的干扰引起。因此，LoP植物是LoA植物的一个子集，并且需要进一步的试验以将观察到的表型鉴定为LoP缺失表型。
为了进一步研究观察到的浅灰色种子表型的性质，进行杂交。通过将三倍体转基因A68植物与来自有性FCH72二倍体植物的单倍体花粉进行异花传粉来检测三倍体转基因植物中的LoP。收集、播种这些杂交的种子，并用流式细胞仪测量后代的倍性水平。发现后代均为四倍体，这表明LoA植物具有双倍体孢子生殖和种子繁殖的能力，但缺乏孤雌生殖。
作为对照，播种无融合生殖三倍体A68植物的种子，发现它们都是三倍体。在相同的播种中，从携带具有向导RNA-1的T-DNA的显示浅灰色表型的各种植物取种子，但这些种子从未发芽(图2)。预期携带具有向导RNA-2的T-DNA并显示空种子表型的植物在与FHC72杂交后会出现类似的发芽测试结果(未进行发芽实验)。总而言之，可以得出结论，药用蒲公英A86携带一个显性Par等位基因，其序列为SEQ ID NO：5，其是孤雌生殖所必需的，以及两个隐性有性等位基因，其序列分别为SEQ ID NO：10和15。
实施例2
孤雌生殖所必需的基因可用于将孤雌生殖性状转移至无无融合生殖或无孤雌生殖的植物。可以使用具有SEQ ID NO：5的基因或基因的编码序列或同源基因来实现这一点。用T-DNA制备二元载体，该T-DNA至少具有SEQ ID NO：5的基因或同源基因，由其天然启动子或雌配子特异性启动子驱动。该基因构建体通过农杆菌介导的转化而转化至无孤雌生殖的植物，例如莴苣或拟南芥。对测试转基因存在阳性的植物评估孤雌生殖的发生。由于该性状是显性的，对初级转化植物(T0)进行测试。在非无融合生殖植物中，孤雌生殖可以通过用水杨酸甲酯透明的胚珠的诺马斯基微分干涉显微术(Nomarski Differential InterferenceMicroscopy，DIC)在显微镜下检测(Van Baarlen等2002)。在没有杂交或自体受精的情况下，孤雌生殖的卵细胞发育成胚。在含有上述T-DNA的植物中发现至少一些这样的胚。
植物材料
对于本实验，使用野生型莴苣：Iceberg型，Legacy，Takii Japan和红长叶莴苣(Red Romaine)型，Baker Creek Heirloom Seeds。
DNA构建体
构建在T-DNA区域编码以下成分的二元载体；拟南芥的EC1.1启动子(如Sprunk等2012中所述)，其驱动药用蒲公英的Par等位基因CDS序列(SEQ ID NO：3)的表达，然后是3'UTR的前250个碱基(SEQ ID NO：4的前250个碱基)，随后是35S终止子和用于选择的新霉素磷酸转移酶基因(nptII)。产生这种二元载体的合适技术是
Golden Gate或Gibson 
(例如，参见Ma等,2015)。携带该T-DNA的转基因品系编号为pKG10824。
植物转化方法
农杆菌转化通过使用根癌农杆菌对莴苣进行不依赖于基因型的转化来进行。这些方法是本领域公知的，例如在Curtis等中所教导的。适用于莴苣遗传转化的任何其它方法可用于产生携带所需T-DNA的植物，诸如Michelmore等(1987)或Chupeau等(1989)中所述。
结果
如上文“DNA构建体”部分所述，对测试转基因存在阳性的植物评估孤雌生殖的发生。由于该性状是显性的，对初级转化植物(T0)进行测试。在没有杂交或自体受精的情况下，孤雌生殖的卵细胞发育成胚。为了防止携带转基因的植物的任何受精，使植物在温室中生长，并且在显微镜观察之前，人工去雄所有的花。去雄是通过在花冠长大之前剪除总苞来进行的。在非无融合生殖植物中，孤雌生殖可以通过透明胚珠的诺马斯基微分干涉显微术(DIC)在显微镜下检测。此处采用水合氯醛的透明方法；一种常用于透明植物胚珠以进行显微成像的方法(例如，Franks等2016)。去雄后75小时，收获花芽并用水合氯醛透明胚珠。在所有7个评估的转基因品系中，在这些透明的胚珠中观察到多个胚(参见表3，显示这些品系中5个的数据)。图3显示了这种观察到的胚的实例。在一些单个胚珠中，观察到多个胚(多胚)。图4显示了观察到的多胚的实例。然而，观察到多胚的频率比单胚低得多。在未去雄的转基因品系中，在雄配子体发生完成之前以及因此在受精之前已经可以观察到胚。在这些未去雄的转基因植物中，在一些罕见的情况下也观察到多胚。在以相同方式去雄和成像的未转化对照植物中，根本没有观察到胚。
表3：编码驱动药用蒲公英Par等位基因的EC1.1启动子的T-DNA构建体在转基因莴苣品系中的作用。所示数字来自去雄后75小时的观察。在未转化的对照中，在去雄后75小时未发现胚。在单个花芽中存在约25个胚珠。进一步分析在单个显微平面中可见的胚珠。
这些结果证明，药用蒲公英的Par等位基因本身足以诱导莴苣中的胚形成。这是用药用蒲公英的Par等位基因在莴苣中诱导孤雌生殖的一个明显的实例，在没有杂交或自花受精的情况下，卵细胞发育成胚。当以相同方式将莴苣同源物(SEQ ID NO：22)用于植物转化时，预期类似的结果，例如用包含T-DNA区域的载体转化所述莴苣植物，该T-DNA区域包含拟南芥的EC1.1启动子(如Sprunk等2012中所述)，其驱动编码莴苣同源物的序列(SEQ IDNO：22)的表达，具有35S终止子和用于选择的新霉素磷酸转移酶基因(nptII)。
实施例3
SEQ ID NO：5的基因在孤雌生殖和非孤雌生殖的植物物种中具有同源物。所有这些序列通过多序列比对和变体调用(包括5'和3'调控序列)进行比较。以这样的方式进行，以确定SEQ ID NO：5的基因的孤雌生殖的植物物种版本仅表现出哪些差异。
本发明人鉴定了Par等位基因启动子序列中起始密码子(SEQ ID NO：2)上游(3')102bp处的1335bp的微型反向重复转座因子(MITE)序列或MITE样(本文由SEQ ID NO：60定义)，鉴定其在有性对应物(SEQ ID NO：7和12)中不存在。预期这种MITE或MITE样序列是孤雌生殖表型的指示，并且可能是孤雌生殖表型的原因，例如负责改变编码蛋白质的表达水平。
这些孤雌生殖等位基因特异性多态性、插入或缺失可以通过本发明孤雌生殖基因的有性等位基因同源物的化学诱变或靶向基因编辑引入到非孤雌生殖的植物中。例如，PAR基因的启动子序列可以被蒲公英Par等位基因的启动子(即SEQ ID NO：2)取代，或者MITE序列可以被引入到非孤雌生殖的植物的PAR基因中上述蒲公英Par等位基因中的MITE序列同源的位置。在引入这些孤雌生殖等位基因特异性多态性、插入或缺失后，植物将获得孤雌生殖性状。在非孤雌生殖的植物中，孤雌生殖可以通过用水杨酸甲酯透明的胚珠的诺马斯基微分干涉显微术(DIC)在显微镜下检测(Van Baarlen等2002)。在没有杂交或自花受精的情况下，孤雌生殖的卵细胞发育成胚。在具有上述特定多态性、插入或缺失的植物中发现至少一些这样的胚。
实施例4
三倍体和四倍体蒲公英无融合生殖体作为花粉供体与二倍体橡胶草(Taraxacumkoksaghyz)植物杂交。花粉供体本身是通过将有性橡胶草与无融合生殖的短角蒲公英(Taraxacum brevicorniculatum)花粉供体杂交获得的。因此，无融合生殖基因来源于短角蒲公英(Kirschner等2012)。使用PCR标记物测试三倍体后代植物中是否存在Par等位基因和二倍体孢子生殖(Dip)等位基因(参见WO2017/039452A1)以及无融合生殖种子的产生。无融合生殖结实定义为在没有异花传粉的三倍体植物上产生有活力的种子。
引物DIP_F(SEQ ID NO：33)和DIP_R(SEQ ID NO：34)被设计在二倍体孢子生殖基因VPS13上，以便特异性地扩增Dip等位基因。使用这些引物，Dip等位基因的存在产生PCR829bp的PCR产物，而该等位基因的不存在则不会产生PCR产物。
在SEQ ID：2和SEQ ID：4上设计引物PAR_F(SEQ ID NO：35)和PAR_R(SEQ ID NO：36)以扩增Par、par1和par2等位基因中的任一个。Par等位基因的存在可以通过PCR产物的长度来区分，如表4所示。
表4：使用引物对PAR_F(SEQ ID NO：35)和PAR_R(SEQ ID NO：36)扩增孤雌生殖(Par)等位基因及其有性对应部分(par等位基因1和2)的PCR产物的扩增子长度。
| Par等位基因 | par等位基因1 | par等位基因2 | |
| 扩增子长度(bp) | 2400 | 1071 | 1111 |
测试了56株后代植物，观察到Par等位基因的存在与孤雌生殖之间存在100％的相关性，如下表5所示。未观察到产生无融合生殖种子的并且对DIP和PAR标记是阴性的植物。
表5：作为花粉供体的三倍体和四倍体蒲公英无融合生殖体与二倍体橡胶草植物杂交后代的基因型分析和表型分析。
因此可以得出结论，从药用蒲公英的Par基因座开发的标记也鉴定了在不同物种短角蒲公英中孤雌生殖的存在，这进一步证明了Par等位基因导致孤雌生殖。
实施例5
构建无融合生殖A68的γ辐射缺失群体
将来自克隆A68的约3×2000个种子用三种不同剂量γ辐射：三分之一用250Gy，三分之一用300Gy，三分之一用400Gy。在温室中的花盆中总共种植了3075株来自辐射种子的植物。在低于10℃的2个月的春化期后，植物再次在加热的温室中生长。超过90％的植物开花并产生种子。对植物进行分类，无论其是否显示无融合生殖丧失表型(LoA)。无融合生殖的A68植物自发产生种子并形成具有深棕色中心的大的白色种子头，其中种子(瘦果：单种子果实)附着在花托上。在无融合生殖丧失表型的情况下，种子头的中心较轻，并且通常种子头的直径减小。最后，102株植物被鉴定为具有无融合生殖丧失表型。
使用Wu等(1992)的方法，单剂量显性标记可以在同源多倍体植物中定位。为了发现与Par基因座相关的AFLP标记物(Vos等1995)，使用分群分析(Bulked SegregantAnalysis)法(Michelmore等1991)。构建两个对比DNA库，A库含有来自10个三倍体PAR植物的DNA，B库含有来自10个三倍体非Par植物的DNA，所有后代来自杂交TJX3-20(二倍体有性)×A68。使用Nomarski DIC显微镜对非Par植物仔细地进行表型分析，以确定不存在孤雌生殖(Van Baarlen等2002)。对于Par库，使用无融合生殖植物。筛选了147种AFLP引物组合(Vos等1995)，以确定A库中片段的存在和B库中片段的不存在。在来自库的个体上验证库中的对比片段。基于TJX3-20×A68杂交(76株植物)，使用17种AFLP标记以构建Par基因座染色体区域的遗传图谱。17种AFLP标记中的14种与Par表型严格共分离。这是抑制Par基因座附近重组的指示。
当三个同源染色体之一被部分缺失时，位于缺失区域上的单剂量AFLP标记将丢失。对LoA植物的AFLP分析表明，许多LoA植物丢失了一个或多个与Par基因座遗传连锁的AFLP标记。缺乏Par连锁的AFLP标记的LoA植物在与二倍体花粉供体杂交后产生四倍体后代。这表明这些LoA植物尽管失去了无融合生殖表型，但仍是二倍体孢子生殖的，产生未减数的卵细胞。这些LoA植物可以基于它们缺乏的Par遗传连锁的AFLP标记的数目进行分级。丢失的AFLP标记的数目是缺失大小的指示。在LoA植物中最常丢失的AFLP标记被认为最接近Par基因座。植物i34缺乏最少的PAR连锁的AFLP标记，因此被认为具有最小的缺失。
实施例6
Par基因在无融合生殖体蒲公英属植物vs Par缺失且有性植物中的雌配子体中的基因型和等位基因特异性表达
来自配子体的不同发育阶段的细胞和组织使用SLμCut仪器通过激光辅助显微切割(LAM)分离，该SLμCut仪器利用固态UV-A激光(波长为约350nm)切割组织(2001,MedicalMicro Instruments,Glattbrugg,Switzerland)，如Wuest等(2010)和Florez-Rueda等(2020)中所述。随后，进行转录组分析。使用PicoPureTM RNA分离试剂盒根据制造商(ThermoFisher Scientific)的说明书提取RNA。为了保持样品之间的原始表达差异，mRNA(逆转录为DNA后)使用CEL-seq和CEL-seq2方案进行线性扩增，如Hashimshony等(2012)和Hashimshony等(2016)所述。
对三种植物品系进行了比较：1.三倍体无融合生殖体A68(简称：APO)，源自荷兰，2.三倍体缺失系i34(PAR缺失系，源自A68，参见以上实施例5)与二倍体花粉供体FCH72杂交产生的四倍体PAR缺失后代(简称：DEL)以及3.二倍体有性植物FCH72(简称：SEX)，源自法国。
每株植物系，对5个不同的发育阶段/组织类型进行采样(表6)。对于非常幼小的阶段，分析单个样品。从成熟胚囊、中央细胞和卵母细胞器(oocyte apparatus)(卵细胞和助细胞)一式三份取样。这些一起代表每种植物品系的9个样品(表6)。
表6.每种类型和阶段分析的样品数量
线性扩增的DNA在Illumina Hiseq平台上进行测序。单个读长(reads)被映射到Par基因的序列(图5)。在任何PAR缺失或SEX植物(所有阶段和组织)中均未检测到Par基因的表达。在APO系中，在成熟配子体的所有样品中均发现了Par基因的特异性读长，包括在卵细胞器和中央细胞中。在无融合生殖体的一个较幼的发育阶段也检测到一些转录读长。根据该方法的3'端扩增偏向性，大多数读长映射到编码序列的3'端和基因的3'-UTR。
因此，Par基因在无融合生殖体的7个样品中表达，而在缺失品系的7个样品中不表达，在有性品系的7个样品中也不表达，它们具有相当的发育状态。这进一步强调了该基因在中央细胞和卵细胞器中的异位表达是造成卵细胞停滞丧失的原因，并因此造成胚的孤雌生殖发育。
也如实施例3所示，可能由于启动子区域中MITE序列的影响，这些细胞中无融合生殖体中Par基因的表达可能不会像在有性生殖中那样受到抑制。因为MITE很大，所以它可能在物理上干扰Par基因的转录因子的结合。
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序列表
<110> 主基因有限公司
<120> 孤雌生殖基因
<130> P6069700pct
<150> EP 19177252.4
<151> 2019-05-29
<150> EP 19200872.0
<151> 2019-10-01
<150> EP 20170243.8
<151> 2020-04-17
<160> 60
<170> PatentIn version 3.5
<210> 1
<211> 170
<212> PRT
<213> Taraxacum officinale
<400> 1
Met Ala Asp Asn Gly Asn Thr Gly Arg Gln Lys Asp Asp Asp Gly Gly
1 5 10 15
His Asp Gly Pro Arg Gln Asn Pro Thr Thr Pro Pro Ser Pro Ser Arg
20 25 30
Thr Pro Arg Arg Pro Arg Arg Asn Thr Ser Pro Pro Lys His Ser Pro
35 40 45
Gly Ala Ser Ser Ser Thr Met Pro Ala Pro Pro Thr Pro Pro Ala Pro
50 55 60
Thr Gly Ile Thr Gly Ala Ser Ser Ser Ser Val Gly Thr Asn Ile Ile
65 70 75 80
Ser Phe Ile Pro Pro Lys Thr Lys Arg Thr Lys Ser Val Ile Cys Pro
85 90 95
Ile Cys Asn Lys Asp Met Cys His Glu Lys Ala Leu Cys Gly His Ile
100 105 110
Arg Trp His Thr Gln Glu Glu Arg Leu Ala Ala Ser Ile Ala Ile Ala
115 120 125
Arg Ala Leu Ser Ser Asn Val Val Val Ser Gly Asn Gly Asp Glu Asp
130 135 140
Glu Gly Pro Ser Lys Lys Tyr Lys Leu Pro Asp Leu Asn Lys Ser Pro
145 150 155 160
Pro Pro Glu Glu Glu Asp Glu Asp Ala Ala
165 170
<210> 2
<211> 1900
<212> DNA
<213> Taraxacum officinale
<400> 2
taaggccata acaaggttat ttgaacattc gcctaaaccc taaaccaact atatctattc 60
ggatttgtta cgagttttag agtttctaga aaatcagtta tatatcgatt aggcttggtt 120
ctaggttata agaaaaaatg attctctatc cggtatgaca tagagacaaa ccggtctact 180
tttataagaa aatcatattt gaaacaatgt ctagttaata attaattaat atgtgcatgt 240
ttaaatttag aatatcctga atttatattt agtgcgcaac cctatgtaag atcgaactaa 300
ttttaaaccg taaccagaca tgcatgtttg atcagtaata aaaaggctat tatcgtcgaa 360
aatattaaaa caaaatataa tatcttattt ggcgccattt catgtcaacg gttgaaagca 420
agttaccaac gcctccaacc gatcatttct ccggtaacaa ccaactgcta ccagggccgg 480
ccagaaggct gtgcggcttg ggctgtggca caggggcagc agatttagtc catttattat 540
ggtctgctat ttatttgggt ccaataagat cagggcacta aaactcaaat acatcaattc 600
aagtcggcgg caaaagaata aaaagcgcca atttttagaa gacaaggaga cagcacaata 660
atcatgcttt tccaatggcc aattaatttt agcctttgtg aggtataaat atgaatcact 720
ttttctccaa atatcattat tgtaaactag taaagagcgt aaattgtttt atcgctaatc 780
ttcttctttt tcaaatataa ttgtttgatt ctttgtagta ggtacgttaa gttttatatc 840
attcgtcctg gaaacctttt cttatgcccc ttcaatttat gaatttgtgt tttgtattgt 900
cttgctatta atccaatgca aactaattct gccgcatcga tctcacagag ttggagcagc 960
ttaaaaggag ttttggggca acttaaaatg agggtagctt ttttgttgtt gttggtctac 1020
tatttacttt ggtccttatc gttttagttt tgtgttcatt gaggatttga atgactttga 1080
acacaagttt ataaaactaa gtttttgtga ttctaaattt ttgatgtgtg caaccgacta 1140
aatgaatata aaattctgat ttgtgtattt ctgatttctg atttaccggt tattcattat 1200
gtttttctgg atctcatatg aatttagtat ataaaaatct caggttcttc ttactcagtc 1260
tagtatattt tggcattggg gatcagggtt aaaatcatca tttttctaag cactaacaag 1320
ctttaaaacc atcaattaaa agttagttct ccatacctgg aattgtttat tgctttagta 1380
ttcatttcat agtgtcaata ccttacaact gtttattgtt ttcaaatgtt tactgttttg 1440
cactttagtt gtttttagca tgtttcaact ttcaaagtta attgacgata gtgtcatata 1500
cgaaatatca tcactcatac aaaattaaac tggatagaat tttttccgat gaaatctatt 1560
tttaaatagt tacgaatata ataggagact taatatgttt gtttcaaaat tacacaaaaa 1620
caccatggaa atgggcatta gtttgtataa gcatcctttt aaatataaag ctttgttaca 1680
ttgatgcctt ctaaatatat aatattgtaa tgatttgtag taaacaccat ggaaatgggc 1740
attattaaga tgtttcgccc agggcatgaa aaactatcgg accggccctg actgctacta 1800
ataaccgcca ccgcaaaaac ctataaatac aacttccatc acaattctgt cccctactcc 1860
catttcaatc caaaaaaagg ctacacagac atatacagcg 1900
<210> 3
<211> 513
<212> DNA
<213> Taraxacum officinale
<400> 3
atggcagata atggcaacac cggccgtcaa aaggatgacg acggtggcca tgatggacca 60
cgccaaaacc caactactcc accctcccct tcccgcaccc ctcgaagacc aaggcggaac 120
acatcaccgc ccaaacattc tccgggggcg tcttcaagca ccatgccagc gccgcctact 180
ccccctgcgc cgacgggaat caccggtgct tctagttctt ctgtgggtac taatataatt 240
tcatttattc cacccaaaac caaaagaacg aagtcggtga tctgcccgat ctgcaacaaa 300
gatatgtgcc atgagaaggc gctgtgtggc cacatccggt ggcatacaca ggaggaaaga 360
ttggcggcca gcatcgctat agcaagagcg ctatcttcta acgttgttgt ttctggcaat 420
ggcgatgaag atgaaggtcc atctaaaaag tataaactcc cggacctgaa caaatctcca 480
ccgccggagg aggaggacga ggacgctgcc tga 513
<210> 4
<211> 5356
<212> DNA
<213> Taraxacum officinale
<400> 4
ttttcttgcg ggagaggcat gcatgtatgt gtggcctttt tgcaataata ctatgcatgg 60
gagatgcata aagtttatct ttattttttt tagtttaata atgtgtggca aactttatat 120
catttatgat ttcggccatg tactgctaca tcgttctagg atggttattc cctagaattt 180
gctttatgta tttgttttct ttgttaattc ttctcttcct taaacgttat ttcatgtgaa 240
ttgcgatgtc atgatattgc aggaatgtaa cttagaactc aacgataaaa agtgtgttga 300
gatcgtggaa gttgatcatt tatcttgaat tcataagata agcaacatga gataggataa 360
ttaattactg tggggatttc tttttatatt catattattc gttgtaatgc aaatttttaa 420
gtagaataac gaatgtgtct tattatttac ggttaaatgc aaaagatcat taacatatct 480
ccattgattt gtttactgat gtattctcat gtgttctgtt gctaaattct attttgaaaa 540
gtatattcaa ttaaagcaat gcattttaga tataaagcaa ctccataact tataattgtg 600
tagattttta ccaattcaat gtcttaaaag aaaaataaaa attgaaagtt taattaattg 660
ccatagtgat cccgttcccg tcaattatgt gatcaatttt taccagagca aaaccccaca 720
tttcttgtag tgaacaatct gtggatgata accgatcgaa gacttttgtg gggatttcac 780
taaccaacta tttggctctc attaattact attagttata taaggccatc ccaagtattt 840
tggcaatatt caaaacgagt aaactataaa attacacttt tggttcttgt atttctaact 900
tttagaatta tgtatttggt tcttatagtt ttttatttgt atgaattaca ttacttttgg 960
tccacaatat tttttatttt ctataattcc atttttagtc attgtagttt taatttgcaa 1020
taaaaaatta atttatattt ttaatccata tatatagttg gcaatattct atattttagg 1080
ttcagattgt atcttttata aaaaagagtc ttaaaagact aaaccttttg gatccttata 1140
gttttcacaa tcacttgtga tttttttaat gaacatatca tatttcttaa attacatcag 1200
taatctacgt ataacattta atataatttt atttaatcag aaatatagga ttcgtataat 1260
aactttgctg aacttaggct tcatttttaa caaaggataa tgcatcatgc atgacataaa 1320
aatttgtaat atttctaatg tgcatgcact acaagaacct gacccttcat atacattaat 1380
gaatgattct cttttcttat ttaatcttag caagcggttt ggataattgt ctgcacaata 1440
ttaagaattg cactagctag ctaggggagt ccatgacgta ataagcaaat taactagagg 1500
acttgtagga ggcttgatca cttcattgta tatctaatgg cactatcgat atatgattgg 1560
cttgtgtgta tctatggaat ttaacactat taaaaaatat atctttaatg acaatcaaca 1620
acacacagta atttatgact tacaaaatca tgtgtgatta aaaaaaatgt catgtttttt 1680
aaaattatag aggatttacg acttacataa ttatgtgtca taattatatg tcatcttgtg 1740
taatatttat gacttacaaa aatgtatgtc atgttttatc atcataaaat ctataaaaaa 1800
aaatagaaac acgaggatac tttcagtgta tgttatctgt aaaaaaaaat aaaaactcgg 1860
aggacactaa aagtgtagaa acttgacagc ataaacgaat ttatttacct tcatcaaaac 1920
ttattttact tcaatctctg aaccagctcc cttcattaat ctcttcgaga tgtaggttga 1980
ttttaatgct aaatgaactt ttcaaatgta taaataaaac ttttctgtgc cagatttgac 2040
attgatggag atgaagatac acaagtgaat tcacctttca atgatcatac tgacatacca 2100
agtactccta tcccctctcc acctccacaa aacgaacctc aatttcaaaa acgtaactat 2160
catttaaata tttaatcatt tttcaattgg attcatgcgg taatagaagt tactggactt 2220
taaacttttt gtagctgatg agattcatga gcaacagtcc atggacattt tccaaatcaa 2280
caagtttctg atgaggacaa cgtaaaaaaa gtttttcaat tattaaacat atgattttgg 2340
tttttaaaag ataattaatt atggaaaatc ttcagaaaag tctcaatact ttgggtttat 2400
taatgaaaaa gtctcaaaaa aaaatttgaa taaaaaagtc cataaaattg gataaattat 2460
tcgatttagc ccaatttgcc aagtaaatct ggtaaaatta ggctaaatcg aataatttag 2520
ccagtttggt gggccttttt gcgcaaaaac attattttgg gacttaatcg ttaattttcc 2580
ccaaatttca gtttatttac agtcttggag taaaaccgaa cctagtgcgc taattttggc 2640
ttttggtgtt tggaattatt tgggtttatt gggctcggtt tctgaacgag tttgattttt 2700
ggtttcttga ttcaattttt tggttttggg gttgaaccca tggattttcg atttggtttc 2760
ggctttgata atatttttgt gcaatcgttc aaaatatccg agttagattg ggtttaaatg 2820
ttatcaaaac cgaaaccgag ggtagaaccc caaaactgaa cccaatcgaa aatccatagg 2880
ttgaacccca gaaccgaacc caataaactc taaaaaccaa aagccaaaat taaccgataa 2940
aaaacttaaa atacatgttt gttaaatcta gtaaggtgtt tttgacactt tatgtaaatt 3000
atagaaaaaa atggattaac tgtttgctta ttaagataaa aacgacgtaa aaatagaatt 3060
ggattagaaa tgaaccattt ttttacatcg tttactttat ttactaggtt ataacccgtg 3120
tattacacgg gttgacgata aaaaattgta taaaaaaaca acatataatg gtaagtttgt 3180
aatcgtacat gggttgcttc aattttaatt taaaaattgt aattccaaat gaaatatatg 3240
taggatgata taacatactt gtaaatgtaa atgagagttt gttggcattt catacattta 3300
gcccatcaaa tcaaacgcaa aaaatctttt taatcctata atacatgtgg cacaaatata 3360
cgattagcat ttcgctgaga gagaggcttt aagtgaggat gtgaaaaaga aggaacatag 3420
caacttggaa tttaaagatc aaaaaacctt gaaaggcttt tggaatgtaa aaatatagaa 3480
atgaataaat gtggcttctt gtatgtatca agagtaggac ctgtattaga aacataaaaa 3540
atttgtgtct aataaaaatc aacaaaaaca gaacgtaaac gaaagaagtt attggtaaaa 3600
aaagatttca aaaatagttt ttttttgaac cggttcaaaa atagttattt tacttattta 3660
tccatttaat ccaaactttt ttttgtcaaa tatctcatgg aattagttgg ctgcgtcact 3720
cttgtcactg ggcgttataa cttttgcctt gtggtttgca acaccacctt aaataacata 3780
gacaaataac attaaatatc atgataatac ataataactc acaaaagatg cattctatag 3840
aacggtataa ttgattagaa caaaataatc taaatagttt atgaggatta ccttgataat 3900
tcccttggag tgaatactat ccacgatgtc atcttccaaa ggtattcaat ggcttttatt 3960
agaacccaat ttcagaatta cattgatgcc acagttacat tcactaaata tcttcctctt 4020
aggcttttag tttgactata ttttaaccat tttcacatca cactttaacc accccctttg 4080
gttattaatg tcaaaaggaa caaaactcac aaatcaaatg cgtggtgttt atactattgg 4140
tacaatgttt ataaatttct taaaaatgag cacaatttca aaaaacttaa acaaatgtca 4200
atggtcacca tgtggtatta atattataaa gttctaaaaa taccattcca ccccaacatg 4260
gtactaacca gttagaaaat caagaaacag aaacatactc tttaatctaa aacaatataa 4320
tatagggatg tgttatgagt ttagtttcat ctaaccatga attcatatag gattgtacgt 4380
ttctcctcga aattgcaaag aattttgttc gactaagcaa tctacatatt aatggatttg 4440
tatggtttct cggtttgtag gtaagttaag agctaccagg aaagagaaaa tttgcaccat 4500
aggtactaaa cacattctac aatggtggct ttaacgtgag agagaataga gaggtcgaaa 4560
catatttccg ttaaataagc aaacataagt ttctatttgg gatcaataag tttcttccta 4620
tagtggtaaa tggatctgta cttgttaagc tcttaaacat atttgttgtc actgttcact 4680
tcctattttg cagtgctctt ctattactta cattcttctt atgacattct tttaataagg 4740
aaaaataacc acttattcta gtattctatt gttttgttag agtcaatatc agataagaac 4800
ctaaaaaggt gggacaaatt gataacttaa tcaatatatc aaactgtcat atgtctcaag 4860
tctcaaccca aattgatatg tttaagaaaa attcagaaaa gatatgtcca tgggacatgt 4920
gcatgtatat gtataagtgt acattcctaa aaaaagtagg ctcatatgac atgagcatgt 4980
atgtgcttat tgttataaag attacaaatt ttcttaaaaa tgaacacaat ttcaaaatac 5040
ctataccaaa gcccatgatc aacatagcat ctgatattaa tattataaag ctctaaacaa 5100
atctaccccg acatcaagtg gtacttacaa gttggaaaat caagatacaa aaacatagct 5160
tttaatctaa gtaaaaacac aataatatag ggatatgacc tatatgcatg tttaaaatct 5220
taagtctgta aagattgata ggattggaaa actcttctat tcaacacaag aaaagaatta 5280
cagtaggagg agggactctc acactcaact actacaaaaa aaccaaccct cacttactaa 5340
cactctcaca ctagtg 5356
<210> 5
<211> 7769
<212> DNA
<213> Taraxacum officinale
<400> 5
taaggccata acaaggttat ttgaacattc gcctaaaccc taaaccaact atatctattc 60
ggatttgtta cgagttttag agtttctaga aaatcagtta tatatcgatt aggcttggtt 120
ctaggttata agaaaaaatg attctctatc cggtatgaca tagagacaaa ccggtctact 180
tttataagaa aatcatattt gaaacaatgt ctagttaata attaattaat atgtgcatgt 240
ttaaatttag aatatcctga atttatattt agtgcgcaac cctatgtaag atcgaactaa 300
ttttaaaccg taaccagaca tgcatgtttg atcagtaata aaaaggctat tatcgtcgaa 360
aatattaaaa caaaatataa tatcttattt ggcgccattt catgtcaacg gttgaaagca 420
agttaccaac gcctccaacc gatcatttct ccggtaacaa ccaactgcta ccagggccgg 480
ccagaaggct gtgcggcttg ggctgtggca caggggcagc agatttagtc catttattat 540
ggtctgctat ttatttgggt ccaataagat cagggcacta aaactcaaat acatcaattc 600
aagtcggcgg caaaagaata aaaagcgcca atttttagaa gacaaggaga cagcacaata 660
atcatgcttt tccaatggcc aattaatttt agcctttgtg aggtataaat atgaatcact 720
ttttctccaa atatcattat tgtaaactag taaagagcgt aaattgtttt atcgctaatc 780
ttcttctttt tcaaatataa ttgtttgatt ctttgtagta ggtacgttaa gttttatatc 840
attcgtcctg gaaacctttt cttatgcccc ttcaatttat gaatttgtgt tttgtattgt 900
cttgctatta atccaatgca aactaattct gccgcatcga tctcacagag ttggagcagc 960
ttaaaaggag ttttggggca acttaaaatg agggtagctt ttttgttgtt gttggtctac 1020
tatttacttt ggtccttatc gttttagttt tgtgttcatt gaggatttga atgactttga 1080
acacaagttt ataaaactaa gtttttgtga ttctaaattt ttgatgtgtg caaccgacta 1140
aatgaatata aaattctgat ttgtgtattt ctgatttctg atttaccggt tattcattat 1200
gtttttctgg atctcatatg aatttagtat ataaaaatct caggttcttc ttactcagtc 1260
tagtatattt tggcattggg gatcagggtt aaaatcatca tttttctaag cactaacaag 1320
ctttaaaacc atcaattaaa agttagttct ccatacctgg aattgtttat tgctttagta 1380
ttcatttcat agtgtcaata ccttacaact gtttattgtt ttcaaatgtt tactgttttg 1440
cactttagtt gtttttagca tgtttcaact ttcaaagtta attgacgata gtgtcatata 1500
cgaaatatca tcactcatac aaaattaaac tggatagaat tttttccgat gaaatctatt 1560
tttaaatagt tacgaatata ataggagact taatatgttt gtttcaaaat tacacaaaaa 1620
caccatggaa atgggcatta gtttgtataa gcatcctttt aaatataaag ctttgttaca 1680
ttgatgcctt ctaaatatat aatattgtaa tgatttgtag taaacaccat ggaaatgggc 1740
attattaaga tgtttcgccc agggcatgaa aaactatcgg accggccctg actgctacta 1800
ataaccgcca ccgcaaaaac ctataaatac aacttccatc acaattctgt cccctactcc 1860
catttcaatc caaaaaaagg ctacacagac atatacagcg atggcagata atggcaacac 1920
cggccgtcaa aaggatgacg acggtggcca tgatggacca cgccaaaacc caactactcc 1980
accctcccct tcccgcaccc ctcgaagacc aaggcggaac acatcaccgc ccaaacattc 2040
tccgggggcg tcttcaagca ccatgccagc gccgcctact ccccctgcgc cgacgggaat 2100
caccggtgct tctagttctt ctgtgggtac taatataatt tcatttattc cacccaaaac 2160
caaaagaacg aagtcggtga tctgcccgat ctgcaacaaa gatatgtgcc atgagaaggc 2220
gctgtgtggc cacatccggt ggcatacaca ggaggaaaga ttggcggcca gcatcgctat 2280
agcaagagcg ctatcttcta acgttgttgt ttctggcaat ggcgatgaag atgaaggtcc 2340
atctaaaaag tataaactcc cggacctgaa caaatctcca ccgccggagg aggaggacga 2400
ggacgctgcc tgattttctt gcgggagagg catgcatgta tgtgtggcct ttttgcaata 2460
atactatgca tgggagatgc ataaagttta tctttatttt ttttagttta ataatgtgtg 2520
gcaaacttta tatcatttat gatttcggcc atgtactgct acatcgttct aggatggtta 2580
ttccctagaa tttgctttat gtatttgttt tctttgttaa ttcttctctt ccttaaacgt 2640
tatttcatgt gaattgcgat gtcatgatat tgcaggaatg taacttagaa ctcaacgata 2700
aaaagtgtgt tgagatcgtg gaagttgatc atttatcttg aattcataag ataagcaaca 2760
tgagatagga taattaatta ctgtggggat ttctttttat attcatatta ttcgttgtaa 2820
tgcaaatttt taagtagaat aacgaatgtg tcttattatt tacggttaaa tgcaaaagat 2880
cattaacata tctccattga tttgtttact gatgtattct catgtgttct gttgctaaat 2940
tctattttga aaagtatatt caattaaagc aatgcatttt agatataaag caactccata 3000
acttataatt gtgtagattt ttaccaattc aatgtcttaa aagaaaaata aaaattgaaa 3060
gtttaattaa ttgccatagt gatcccgttc ccgtcaatta tgtgatcaat ttttaccaga 3120
gcaaaacccc acatttcttg tagtgaacaa tctgtggatg ataaccgatc gaagactttt 3180
gtggggattt cactaaccaa ctatttggct ctcattaatt actattagtt atataaggcc 3240
atcccaagta ttttggcaat attcaaaacg agtaaactat aaaattacac ttttggttct 3300
tgtatttcta acttttagaa ttatgtattt ggttcttata gttttttatt tgtatgaatt 3360
acattacttt tggtccacaa tattttttat tttctataat tccattttta gtcattgtag 3420
ttttaatttg caataaaaaa ttaatttata tttttaatcc atatatatag ttggcaatat 3480
tctatatttt aggttcagat tgtatctttt ataaaaaaga gtcttaaaag actaaacctt 3540
ttggatcctt atagttttca caatcacttg tgattttttt aatgaacata tcatatttct 3600
taaattacat cagtaatcta cgtataacat ttaatataat tttatttaat cagaaatata 3660
ggattcgtat aataactttg ctgaacttag gcttcatttt taacaaagga taatgcatca 3720
tgcatgacat aaaaatttgt aatatttcta atgtgcatgc actacaagaa cctgaccctt 3780
catatacatt aatgaatgat tctcttttct tatttaatct tagcaagcgg tttggataat 3840
tgtctgcaca atattaagaa ttgcactagc tagctagggg agtccatgac gtaataagca 3900
aattaactag aggacttgta ggaggcttga tcacttcatt gtatatctaa tggcactatc 3960
gatatatgat tggcttgtgt gtatctatgg aatttaacac tattaaaaaa tatatcttta 4020
atgacaatca acaacacaca gtaatttatg acttacaaaa tcatgtgtga ttaaaaaaaa 4080
tgtcatgttt tttaaaatta tagaggattt acgacttaca taattatgtg tcataattat 4140
atgtcatctt gtgtaatatt tatgacttac aaaaatgtat gtcatgtttt atcatcataa 4200
aatctataaa aaaaaataga aacacgagga tactttcagt gtatgttatc tgtaaaaaaa 4260
aataaaaact cggaggacac taaaagtgta gaaacttgac agcataaacg aatttattta 4320
ccttcatcaa aacttatttt acttcaatct ctgaaccagc tcccttcatt aatctcttcg 4380
agatgtaggt tgattttaat gctaaatgaa cttttcaaat gtataaataa aacttttctg 4440
tgccagattt gacattgatg gagatgaaga tacacaagtg aattcacctt tcaatgatca 4500
tactgacata ccaagtactc ctatcccctc tccacctcca caaaacgaac ctcaatttca 4560
aaaacgtaac tatcatttaa atatttaatc atttttcaat tggattcatg cggtaataga 4620
agttactgga ctttaaactt tttgtagctg atgagattca tgagcaacag tccatggaca 4680
ttttccaaat caacaagttt ctgatgagga caacgtaaaa aaagtttttc aattattaaa 4740
catatgattt tggtttttaa aagataatta attatggaaa atcttcagaa aagtctcaat 4800
actttgggtt tattaatgaa aaagtctcaa aaaaaaattt gaataaaaaa gtccataaaa 4860
ttggataaat tattcgattt agcccaattt gccaagtaaa tctggtaaaa ttaggctaaa 4920
tcgaataatt tagccagttt ggtgggcctt tttgcgcaaa aacattattt tgggacttaa 4980
tcgttaattt tccccaaatt tcagtttatt tacagtcttg gagtaaaacc gaacctagtg 5040
cgctaatttt ggcttttggt gtttggaatt atttgggttt attgggctcg gtttctgaac 5100
gagtttgatt tttggtttct tgattcaatt ttttggtttt ggggttgaac ccatggattt 5160
tcgatttggt ttcggctttg ataatatttt tgtgcaatcg ttcaaaatat ccgagttaga 5220
ttgggtttaa atgttatcaa aaccgaaacc gagggtagaa ccccaaaact gaacccaatc 5280
gaaaatccat aggttgaacc ccagaaccga acccaataaa ctctaaaaac caaaagccaa 5340
aattaaccga taaaaaactt aaaatacatg tttgttaaat ctagtaaggt gtttttgaca 5400
ctttatgtaa attatagaaa aaaatggatt aactgtttgc ttattaagat aaaaacgacg 5460
taaaaataga attggattag aaatgaacca tttttttaca tcgtttactt tatttactag 5520
gttataaccc gtgtattaca cgggttgacg ataaaaaatt gtataaaaaa acaacatata 5580
atggtaagtt tgtaatcgta catgggttgc ttcaatttta atttaaaaat tgtaattcca 5640
aatgaaatat atgtaggatg atataacata cttgtaaatg taaatgagag tttgttggca 5700
tttcatacat ttagcccatc aaatcaaacg caaaaaatct ttttaatcct ataatacatg 5760
tggcacaaat atacgattag catttcgctg agagagaggc tttaagtgag gatgtgaaaa 5820
agaaggaaca tagcaacttg gaatttaaag atcaaaaaac cttgaaaggc ttttggaatg 5880
taaaaatata gaaatgaata aatgtggctt cttgtatgta tcaagagtag gacctgtatt 5940
agaaacataa aaaatttgtg tctaataaaa atcaacaaaa acagaacgta aacgaaagaa 6000
gttattggta aaaaaagatt tcaaaaatag tttttttttg aaccggttca aaaatagtta 6060
ttttacttat ttatccattt aatccaaact tttttttgtc aaatatctca tggaattagt 6120
tggctgcgtc actcttgtca ctgggcgtta taacttttgc cttgtggttt gcaacaccac 6180
cttaaataac atagacaaat aacattaaat atcatgataa tacataataa ctcacaaaag 6240
atgcattcta tagaacggta taattgatta gaacaaaata atctaaatag tttatgagga 6300
ttaccttgat aattcccttg gagtgaatac tatccacgat gtcatcttcc aaaggtattc 6360
aatggctttt attagaaccc aatttcagaa ttacattgat gccacagtta cattcactaa 6420
atatcttcct cttaggcttt tagtttgact atattttaac cattttcaca tcacacttta 6480
accaccccct ttggttatta atgtcaaaag gaacaaaact cacaaatcaa atgcgtggtg 6540
tttatactat tggtacaatg tttataaatt tcttaaaaat gagcacaatt tcaaaaaact 6600
taaacaaatg tcaatggtca ccatgtggta ttaatattat aaagttctaa aaataccatt 6660
ccaccccaac atggtactaa ccagttagaa aatcaagaaa cagaaacata ctctttaatc 6720
taaaacaata taatataggg atgtgttatg agtttagttt catctaacca tgaattcata 6780
taggattgta cgtttctcct cgaaattgca aagaattttg ttcgactaag caatctacat 6840
attaatggat ttgtatggtt tctcggtttg taggtaagtt aagagctacc aggaaagaga 6900
aaatttgcac cataggtact aaacacattc tacaatggtg gctttaacgt gagagagaat 6960
agagaggtcg aaacatattt ccgttaaata agcaaacata agtttctatt tgggatcaat 7020
aagtttcttc ctatagtggt aaatggatct gtacttgtta agctcttaaa catatttgtt 7080
gtcactgttc acttcctatt ttgcagtgct cttctattac ttacattctt cttatgacat 7140
tcttttaata aggaaaaata accacttatt ctagtattct attgttttgt tagagtcaat 7200
atcagataag aacctaaaaa ggtgggacaa attgataact taatcaatat atcaaactgt 7260
catatgtctc aagtctcaac ccaaattgat atgtttaaga aaaattcaga aaagatatgt 7320
ccatgggaca tgtgcatgta tatgtataag tgtacattcc taaaaaaagt aggctcatat 7380
gacatgagca tgtatgtgct tattgttata aagattacaa attttcttaa aaatgaacac 7440
aatttcaaaa tacctatacc aaagcccatg atcaacatag catctgatat taatattata 7500
aagctctaaa caaatctacc ccgacatcaa gtggtactta caagttggaa aatcaagata 7560
caaaaacata gcttttaatc taagtaaaaa cacaataata tagggatatg acctatatgc 7620
atgtttaaaa tcttaagtct gtaaagattg ataggattgg aaaactcttc tattcaacac 7680
aagaaaagaa ttacagtagg aggagggact ctcacactca actactacaa aaaaaccaac 7740
cctcacttac taacactctc acactagtg 7769
<210> 6
<211> 227
<212> PRT
<213> Taraxacum officinale
<400> 6
Met Ser Thr Val Lys Ser Lys Leu Ser Thr Pro Pro Thr Asp His Phe
1 5 10 15
Ser Gly Asn His Gln Leu Leu Leu Ile Thr Ala Thr Ala Lys Thr Tyr
20 25 30
Ile Tyr Asn Phe His His Asn Ser Val Pro Tyr Ser His Phe Asn Pro
35 40 45
Lys Lys Ala Thr Gln Thr Tyr Thr Ala Met Ala Asp Asn Gly Asn Thr
50 55 60
Gly Arg Gln Lys Asp Asp Asp Gly Gly His Asp Gly Gly Arg Pro Asn
65 70 75 80
Pro Thr Thr Pro Pro Ser Pro Ser Arg Thr Pro Arg Arg Pro Arg Arg
85 90 95
Thr Thr Ser Pro Pro Lys His Ser Pro Gly Ala Ser Ser Ser Thr Met
100 105 110
Pro Ala Pro Pro Thr Pro Pro Ala Pro Thr Gly Ile Thr Gly Ala Ser
115 120 125
Ser Ser Ser Val Gly Thr Asn Ile Ile Ser Phe Thr Pro Pro Lys Thr
130 135 140
Lys Arg Thr Lys Ser Val Ile Cys Pro Ile Cys Lys Lys Asp Met Cys
145 150 155 160
His Glu Lys Ala Leu Cys Gly His Ile Arg Trp His Thr Gln Glu Glu
165 170 175
Arg Leu Ala Ala Ser Ile Ala Ile Ala Arg Ala Leu Ser Ser Asn Val
180 185 190
Val Val Ser Gly Asn Gly Asp Glu Asp Glu Gly Pro Ser Lys Lys Tyr
195 200 205
Lys Leu Pro Asp Leu Asn Lys Ser Pro Pro Pro Glu Glu Glu Asp Glu
210 215 220
Asp Ala Ala
225
<210> 7
<211> 14519
<212> DNA
<213> Taraxacum officinale
<400> 7
ctgacttggc caacattttt tgttcttccg atcccttcgt aacaatcagt ttaagcacaa 60
ttgaattata tggagataaa taggtgaaat ttggaagacg aggatgggag agtgcgaaga 120
gccaaagaag agaaatgtgg gatagcggcc ggccggttgt taatagaagt gcttagatct 180
tggagacctt ctaatgctta tggaggaacc agatgcttca atttgtaaat ctgcgctgcc 240
cacgtagaat attgaaaaag aaaatcaaat cgatcaataa tgcgaccata tactttagga 300
tttcccactt gtataattct cttattcatt ttcaaatctt tatctaaatt aaggatatcg 360
gcaaaaatat aatcctaatt gtttttcaaa atgataagga ttataaggta acttaattct 420
tgttttattc atatttaatt attattgttt tttatcaatg ttttaaaacc cgggttttga 480
gtcaacccgg tcttgtgaaa aatcccgggt cagccggtca gctgatctac cggttcaata 540
tatttaaata aaatataaat tttcatatag aatttgagtt ttaggctaag aaaatcgggt 600
tttgggctaa ggaaaatcgg gtttaagttt tgggctaatg gattttcggg tcaataacgt 660
ttttttccgt tttttagatc ggttcgaccg cgttttgtgt gaaacccggc cgggttgatc 720
cgagtcaata attggcaaca acccggtata tgttgacccg ctcttgtccc cgggtcccgg 780
ttcaaccggt tggatcggcc gggttgaccc tggttttaaa acactgtttt ttatacaata 840
tatttgtttt tgtttttagt tttgtttttg tttttgattt tgtagatata tttttaattt 900
tatttttatt ttgtataaat atagtcatta caaccgatta tttgtaagtt gagtcggtca 960
aaacgaactt tctgataatt cttatacatt tttcggcgaa acatgaatat tttgatcaac 1020
caatatgctt ttttctaggt tttttaaata aaccgcacgt tcaattgcat attcgcaaac 1080
aatagaactc gaaaacccta atcgagcttg ttttatagaa atctgggttg agtgcataca 1140
taataattaa taactgtgaa aaagtttgtt ttaacatgct aaacctatcg atagtcggtt 1200
ataataatta tatgtaaaca aaatcagaaa caataatgat atatgtcacc caaaaaaata 1260
gttactaaat atatacaaaa tcaaaaacaa cagtaatttc ttttacaaaa aaaagagaac 1320
aatagtgact aactttagaa agaaaaaaaa agaactaaat tactcttata aatcattatc 1380
ttttatataa attggtaaat tggtcatgta atctagaatg tgaattgcgg aatgctttag 1440
tcacattttg aacaaactgg taattattaa tttcgataaa agttcttcaa gaatagtttg 1500
gtcacatttt ggacgacaat ggtgtgtttt ccatagtaag caatacgtaa tgagttcgct 1560
taataggaaa atagatctaa agcttaatga aatcatatgt ataagtaaca tatatttttc 1620
acacgttaca ttgtttgtaa cgtcctcaat ttttttaata tatataacta ggtacactct 1680
aaggtcgtgc taagcacgac ccacgaatgg taatttaggt aatttttcat ccttgactaa 1740
aatacattat gctaataaca aagaattaag aatacaatgc tataatgcat aagtaaataa 1800
aagtacatga tatttctcac atttagtctt tatacataga aaacttacac tggccatttg 1860
caaaaaaaca ttaaatctcc ctaaaaagta gagtttacaa aatgaaagct tgtttttata 1920
gcaaacaata atcaatatgt gaagctatag tatagtaacc tactcatttc aacttttagg 1980
cagtgcagta attacttact catgttaatg gtgaaatgca ttaagataaa cttcattcaa 2040
cggattaaat ctgggttcat tataaatgta ctcttagtta ctcctccatg taactgcatt 2100
tagcgaacaa caaaagctct aataaataat atgaaacaca tgctaattta taaatgacat 2160
catgggtata gatgctgtaa catcctaaac ttttaaaata aataataaat catatttggg 2220
atttaaagta atattttaag ataattgata aataagagaa tgaaatacaa agaggcattt 2280
tatttttagc ctttgattta taaaaaagaa aaaaccttga aattggatgt attttgaaat 2340
gggtaagtat tttaaaagat gtgtaagtta agggggataa atggaaatta aggagataag 2400
catttgacca aagtcaaaat agaaggaaca ataagtcggt cgattgtttt cctgtgtgca 2460
tcttcggacg acagagagac agaaaagaac gagcgagaga cagagaggaa tcgagacaga 2520
gagattatcg agagagagac agcgaagtcg ggagagggag accgagaacg accgagacag 2580
gaggaaaacc gagagacaag gggaggacga ggcagcagca gcctaggctg ctgcgttttt 2640
ggcgacagaa aaccaccgcc agcggcggtg gtggtgagcg acggtggtgg tggagccgag 2700
aaggggttgt ggtggtgtta gaagtgatgg ttcatctcaa tttcttgata atttgccctt 2760
ttggaacaat aaattgaagg taactttcaa agctttagat gattatctct tgaaagtaga 2820
aagcaagggt aatttagtct tcaattatgt tatggagaag ggtaataaca cccaaatgct 2880
aacaaaatca tatagattaa tgtgatttga agttatatag ttcatgaatg tgaaggttga 2940
ctttctgcaa tggtcaaaca tggacatgga catgattata gcatattagc aatgtaatag 3000
gaatggaatc accaattcca ttgttgaaaa tagaaaattt gatgtgtgca gaattttgta 3060
aaatttccca aaacaccaac tttaaggctg cataagtcat gcatgaagaa tgcaaatcag 3120
tccattcttt cgcctaacat caagtccttg caacatagat tacacaggaa aaagaaacag 3180
ccaaatccga gttcgtatga ggattgtatg actgttcaac gtttgtcaaa agttgctgtc 3240
atgctgtcaa accagaattt ttctaagtct gggaaatatg cagaaaatgg gtttgaccag 3300
tttttggagc ttcataactt gatttataca tatccaaatc agttgattct tgagcctaaa 3360
ctgtagagaa agatgaggag aaggcatagg aaaaagaatc aacacatttg gatatcgtat 3420
acaacctgtg caaagggttg aagtaggctc aaaacccatt tttgacataa ttgtcaatta 3480
tatgaattgt gggtaacttt tgccgtaatg aaatgttttg acatataagt gtttatattg 3540
acttataaac atgttaagac tataaacttg ataaagtaac atgtcaaaac ccgaaaatgc 3600
ctatttaggc aaagtatgat gcttaattga gaaatcggtc aaatagcata aaaactaaaa 3660
gtgttctaaa tggtccaaga agactttata cataacttac acatcattag cacatgatgt 3720
gtagtacatg tgaggtttga acacttttat acttggaaaa ttgtcaagta gtcaaaaatg 3780
ggacaaaagg gtaaaattgt ccaaaatgag tatatggcta attaaagcaa gttgagctac 3840
tgaaatgaac ttagataagt attataagga ctataaaagt ataatacttg gttaactatg 3900
agttttgaag gtaatgtcaa tttaaagtaa gttaaccaaa agaaagtaag ttgctaattt 3960
tgggatatat gacccaaatg agttatgcat tggtttttag gttgtgtaat acctataatg 4020
agatatgaat tgattatgag acatgtattg atgataatac aaatgatgta atgaaatagg 4080
ttcgttgcct atcgtggagc aaggagcatg cggattagtg tagctagtct tacctagcta 4140
ctaaggtgag tacgtgtgga ttgttttccc attttcgggt acatggaaaa ataatatttt 4200
tataatgaag tgcctaatgt tttgaaagaa agaatgaatg aaaatacttg atgcttttga 4260
aagaatgttt taaaagaaag taatgttttg aaagaaaaag gtttgatgct ttacgtttga 4320
aaatgaatat gtcttgatat taatgaatgg aatgggtaaa gcatgaatga tataatatgt 4380
aataataaga tgctaaggat ggttatgttt atgatgttca cgtgaatata agaatgttgc 4440
tcgatctaag atgtcccggg tagggattca gaggagccta tcggggtggt acctcccctt 4500
cgcgagatag gttacctaat gtaaatgatg tcaatgtaat ctgatttgtt ctttttgcat 4560
tggtacagac ttggggtata tcagacccaa gtataatatg atatggccat atgataagat 4620
gattaaaaga gtatgtgaaa tgttaaaggt tatgtaaaat gttatatgaa attattcaaa 4680
gagatcttta tggttattat gtctatgaaa aggaagcata taatgttata gatattttat 4740
gtctaaaccc acgtagctca ccagactagt tgtctgacgt atttattttt atgccatgta 4800
tttcaggtta tacacgagga tagactgatg atcgatagaa gctagatgtt acgacggata 4860
gacggagtgg agctttcaag ttctattgta atgatcttac cgtaaattct tatttgcttt 4920
gctatggttg ctgtttgata tataattggt aacacccgag aattgtttta tatatattca 4980
aaaagttttt atttgggacg gtttttgtaa acatgaccaa gtgtcatgaa atatttttaa 5040
aaccatagcg tttttaaaac gataaaatga ggggtgttac agatgcgata atatatctta 5100
ggattatgtc aaatatttat gccaattata catgagagag caagtttata agagttttag 5160
ttattctatc attggatcaa tatagatacg tattaatagt agacagagat gcatgaatat 5220
tatagcagag aactttttgc aaaacctaac ctcactgcat gttgcattca tcaaccccac 5280
ttgtagttgt aacaccatga atatgccaat taagaaatat catgaggtat attgtactgg 5340
aaatgacaaa aagtactaat ttacaataga acgcagtcaa acttggttat tttgtaaatt 5400
ttaagaagca atttacaaaa aataaaacat aataaagaaa aacataacag taaatcacag 5460
acattgctca tggaagagtg actgatcaaa ctgaccgttt attttcttgt ttcttgtaca 5520
actaccaaac gttcctgtat aaccaccaaa tagtcccaga ctttagaaaa tagaacagaa 5580
aaccacagat atacatacat acagccatac attataattt ttaattgaaa aaaatgcata 5640
gaaaaccagt ctcaaataaa gaaccaaata aagaagatac aaatgaaacc tggttatcga 5700
ataaaagagt tagagataat gatttcagca tttgcactca tgtaaacgat atgccattgg 5760
gatccgttgg aaagggtatc gtgaagtgat taaggagaag cggggtgtta gggttttttt 5820
tgcagtttat tgaacaggtt gatcaaattt tgcgccggta gggttaattc cttttcgaac 5880
gggtttctat ctgggtcttt catcaaaatc actcatgttt gtgtctctaa cctttagttg 5940
cagatggata tcagatcaac ggtttcaggc taaatggggg aacaaataac aagaagaaag 6000
cggtgacatc tgtgttcgta ttcgtatggt ggaaggggaa gagtggcgta tgaattggtc 6060
aatgtaaaac atctggaaac aacttctccc aacatgtaaa aggatccgga aaatatgaaa 6120
atgacgatta gagatttcgt tcggatattg ggaatatggt cgcgtctgat tgccctacgg 6180
tgcccaccgt ttgggtttct tgatttgtcg tttagataga tgtcggaggt ctaaattcac 6240
aatcctgttc ttccgcggta attgaaattg ggtaaggcta attgtttaga tcaaatggca 6300
gaggacgatg tgaagttgcg ttgccctata attggctctc gtgaaattga attgcaaatt 6360
tcacaacgat gttagcaaca ataatcgtcg ctctcgcatc ccataccgtc gcaacgtatg 6420
atggtgatcg agattgagag atttagggct agatgtggcg ttggcggaca taggttgtgg 6480
ttgacggaat gaggagaaga cgtgagcgtc gattgaggta gcaacaggcg gagggttaac 6540
ggtgtcatca tggtgaaagg tcgcgcgtcg ttttatggtc gatagatcat ggacgttgtg 6600
agtgaatgcc ggcgcaaaca cggtggagga aagtctatgc tttagttgac ggggcggttt 6660
tctctcggag acggaaagag gacgtcgaaa gaaaggccgg ttggagaaaa tatattaaac 6720
gcgccgacac acgctatcag agaagaccgg ttttcttata taataatgac taggtacgtg 6780
caagggtcgt gctaagcacg acccatggag gcttcttgag aaaattgtaa acacatatca 6840
gataatagcg acgaacttta ctgttattga tatataaagt gattaagcat ggttggccct 6900
aagttaaaac tcaaactact gtaagtatgt aatggtaata atttatcctt ttcagctatc 6960
gatcttagat tgctctgtag tgttgtgtct tagtgctttt cacggttctt ttatatgcat 7020
atagtcacat aaaatcagac acacacatag aataacccac agacaatcac ataaaaatca 7080
gtctgatgag gtattaattt aagcaataag gattgatatt caagcaatta gaggagaata 7140
atatattaat ttcacgtact cctattgtag ttattgtgac aaaacccaac tgtttaattg 7200
tgccattgca aaaagtccac ctgtttaatt tctttatcaa catgtcgttg catcctttta 7260
catgcaaata tttcagaaca tcttcctata atgacccttg agatgaactt tttagaaata 7320
ttgttgctgg ttacattatc agaaaaaaca gagcttatga aaaaagacca aacatagtaa 7380
ctcgtacttc atgtctggta gttcttttga atatttccgt cttttgttcg aaacaatttg 7440
gttgaactgt agtagatctt tcaaactgta tacaacttca caatatgttt tgtaagttca 7500
ttacttctag atcctttcaa aagaagaggt caaactcaaa atttgcataa accagattta 7560
ttgaaggcta attagattag tgttgtaggg ctttttggat ataaataaaa atgaaacata 7620
ggtgctgcaa caggtgcgaa agaatttggt tggacagtag taggtctttc aaactgtata 7680
caactttaca atatgttttg taagttcatt atttctagat cctttcaaaa aaaaggaggt 7740
caaactaaaa aaatttctaa accagattta ttgaaggctg attagattaa cattgcaggg 7800
ctttttggga ataaatataa atgaaagata ggtccaacag aaacgaagga ggtgtgaatt 7860
atgcaacaaa tggaaacaca caattatcgt agtacacatg taattaacag ggatcaattt 7920
ccatttattg ttccatcaat tgccaaattt aaaagataac aaatgcatcc tttttgaatg 7980
gaaaagtatg atacataaaa ccagagagag gtatacattt agtccagaaa cggtggaaat 8040
tactttacct agagttgcgt gaatcaaatg gaaaatcaga ggaagagacc ctaaatcgca 8100
atccatcttc tattccctat agaatcaaat ggaaatttct tacttccgac ctttttcatt 8160
tccacaagtt tgtaatcgat aagatcataa aagaaaaccg atttacatta gcaaatcttt 8220
aaggcacttg tttgtagttc tcaagtatgc gttacgtatg aaattctaga cagaaaaggg 8280
acgacaggga aaataatggt ggaataaata aacaatcaag aatcaaaaga taataatcct 8340
taaaactaac aaaaattcta tcaataagcc ctacaaatga gaaatatgac ggtggagact 8400
cacctgattg cagtggcgga agctcttggg ggctttggta ggccatggcc ccctgttttt 8460
tggaaaattt tcaaaaaaaa aaaaattata tatattatat cattgaataa gtctagactc 8520
aggaaactat gaaagattag aaatagatca agttatctaa atcaaatctc gaaaaatatg 8580
gaaggaatca aaacacaata tttatctcat tgtgaaatca aaaaccctat aaaaaacaca 8640
aatctaacct acaatttaaa tcaaaaacac cataaaaatc aacctaccgt cggattcagc 8700
cgtcgtcgtc ttcttcaccg gaccaccatc caccgccatt gcttcaatcg tgctttctcg 8760
tcaatagctt agtggcttgt tcagctttct tgctctacca atgattttta tacactttcc 8820
agcaccaccg atttgttgaa cggtaatatt atgctccttt atatttttga aatctttgaa 8880
agttcatctg gttttctgtc tctgtaaact gtaaagaaac ttttatgttt tgttaagaaa 8940
atagagagat agagaagaaa gagtattgac gtgcttttcc catatgtctc gaatgccaca 9000
ttaggaaaag acaaaaagaa aacaattagt acattacgct agtgtgatag tactgtatga 9060
ctattagtat gatttacagc tttaataggg attcatagaa aagaacaaaa taaactactt 9120
ataaattaaa aatcgagctt tatttttaat tagtttaaat aaatatttag ctataacata 9180
atatattata tgttttgtat ttataatcaa atataattat ttgtattaca gattagtttg 9240
tataatcaaa tataatgatt catgtttcga aaagattgtt taaatatttt tatatgccat 9300
gttttttaac tgaattatat ttttttgtat gattttcttt acaatttagt ttttctttac 9360
aatttagttt tggcccccct tgtttttttg tttgtgttcc gcccctgcct gattgatgcg 9420
agaaaaggaa agccgccgac atttgtgctt cttcccgcgt taagaaaaat cgcagatgac 9480
attacgattc tttttctaca gaggaatagg ggggttgcga ctttgcgtga tttgttgtta 9540
atgagtgatg cgtcgtccgt tttgatcggt agcccgtgga gactgattta tttgacagat 9600
agatgtcgtt ctcgttgatg atgcaggtgt aattggcggc gttggtgaag cgaagatggc 9660
atagggaatg gctttgcaga ggagaaaagg agtttcacaa atatggatta ttgaatcgtc 9720
gtttcagagt cgcgatggtt agtaatcgat ggatttgggt ttgcacctgt tcaaggggtg 9780
gtgacgtgtc ttactgtaag aaagagtcgc gatggttagt aatcgacgat ggatttgggt 9840
ttgcaacatc gttgacggaa acgacgcggc gatggagaga ggcggtagag ggaatgaaac 9900
gggagattta ccttcaaccg caaacatgtc ttcgttcctc gaaaatcata tgcgcccttt 9960
atgaaaattc cttcacgcgc cgtacacgcg ctcagaaaaa acatagggtt ttcttatata 10020
ataatgacta ggaacgttta tgcgccgtga taatcacggc ccatgagtgg tacaaatgtt 10080
gtaaaactta gaaaatggga aagttgaatg tacaatgtta tgatttattt tattcttcat 10140
agcacaatac ttttgcttta gtgaaagcag ttaaaaagtg gatgtgtcgt tagcaactat 10200
aaacttttaa caaaatatta catacagaac atagaaataa gataaacact aaaaaaatac 10260
aaaatagccg tgttcaacac ggcccacaga atgcaatttg gatatttcta aggaggtgta 10320
aaaaatattg ctattatggg agctaattct aggtatattt tgttactcac atgacatgaa 10380
tattgagtaa ttttgtaaaa ttggtaattt agcttagatt ttcccacaaa gatgagtgtt 10440
aaaactatca aggaggatat atggtacttc ctcaacaaag atgattttga aatatctata 10500
ttttaaatgt aagaactact atcacatgag acaaatattg atattactgg ttaggattac 10560
aacattgata tgtaaaatat tttggtaatg taaattttct aaaaaagctt ggaagtagaa 10620
tgcaacaaca atatatgcta gtattacatg ttgggtgatt ttatgaatgt acttacgtgt 10680
tgatccttat ttaccgtact tgaataattt tatctgatca atgctttacc tatatggcta 10740
tatctagcaa tgtgcattat tcacaaatta tggagtgtaa attcatatag acgtatatga 10800
cagtggatga aaaggaaaac aagtaatttg attgaatcac tgagaaaaaa gcaaaagata 10860
catgcatata aaatgttaca tcaaaataca gaagataaat gaaagaaaga taataatgtg 10920
gagcattcaa aataagtaag aatgtaagtc cttgatagct cgaaattctt ttaagtcttc 10980
cgtgatagcc taatgtttat aatagaagag acattcaaaa tgaataagaa ttatatcacc 11040
taacagtatt taatcctact ctacttggcc attgagcata ttgctattgt taaaaattat 11100
gtccaaatcg catttaactg ttagatgcaa ttctcattgc cgattacact tatatgtatg 11160
tagtaaatta taattaactc tttgtcctgg tcatttgatg tgtcatggat gtttgtaagg 11220
tacaaataat agattacaat aacttaaata taaaattgag ttaatattcc agaattcaca 11280
aaatgtgtaa taacacttat ttataatttc cacgtaatta atgattaact acttagaatc 11340
ctacaagttt ttgtttatca cctttcatat gatcgtataa atgcaacaac ccttaatgca 11400
tttatagttg ttggtctgcc acgaacacct ttttctgttg attatttgtc attattatgt 11460
taaaaaatgt ttgaaaaatg ttaaatggga ttttgttatg gcaaagagat gaacttctcc 11520
aatattgtag ttgcataaat attgccagtt ccatcataca gaataatacc tcttatattg 11580
ggcaatagaa ttatagaaac aataagggtt aaccgactta catgtattga tctctgattc 11640
ttctttacac actggacaaa ttagtctatc atgtcttgca caaccctgtt aaagaaaaca 11700
gtgaaaattg accacgaagg tgtatcctct agtaccaaca ttacatctat cattttaaaa 11760
gggaaatgta tacttatcat atatatctaa tggacgttgc ctttcctcca attgtcactc 11820
ttacatctaa tggaaattat gatcacaatt tacttttaac agcatagata cattcatttg 11880
ttggtaagac acttttggcg gattcattgt attaatcttt catgtaaaaa ttctcattca 11940
agatattagg tttttgttag aacccttgtt atttgaacat gcaacctgaa ttctgattgt 12000
tcccaacacc aacattaata atacctcttc cactagacca tcattccacc ttcaggccct 12060
tgtgttcatt attataagca ctattcggtt tagttctttt atgagtccta tatttacatc 12120
cttaaccatt ttaacaagtc catcatattg ggaattttaa tataaatacc caatgttatt 12180
catttttgtc gcttcagtag atattgacca tactaaagat gagtatttaa tactcataca 12240
acatgaaaga aaaaaatgaa aaactaacag tcacttatgt agccccaatc tccatacaaa 12300
tagcgcttta agaataaacc aatttcaggt gaaattgtag aaaggacata aatctgtaca 12360
tgaaaaaaca ttaacagtta cacataaatt atttgcacag agataaaaag cacaatatgt 12420
gatgagaaga tttaaaagta ggaaagtgaa caaaaataac catcaaaatc caatgttagt 12480
aataaagaaa atgtacgagc gattcataac attcatccaa aacccatatg cgatgttgaa 12540
tgcatatctg gataaaagaa tttaataaac aataagcgcc tgtaaaatca ataaacaaaa 12600
tcaataaaaa tagacctaaa aatgcaaccg acttattgtt tctatacacc aataaatgat 12660
tttcacgaat tttatccaaa gaaagataaa aagtcataaa attgaacggg cttaccgatt 12720
tgaatcgtgg tctgtatctg ccacgcagat cagataacaa tcataatcta catcaaagat 12780
ttcagaaaaa tatgaaactt ccgccattac taatttgata tgaatttctc agagcctaaa 12840
acaaacctaa attactaaaa atcaaacaca ggtttttgac gtttaccttg atttgtgaaa 12900
ataggcattt gggttttttc gtaggatttt ctatgaaaga agccatcaag tttatgacat 12960
tgtaaattaa agcaaagtat gaatatatga aggataggaa aagagatcgg attgtgatga 13020
tccggagaga aaaatacaca aaccagagag aagcgatgtt gtgattgtgg tgtattaatc 13080
tgtttttaaa cctctaacca cttcttgtaa gtccgtaatc agacgaaatg cgcttgaata 13140
ataagtcttt cacgccagtg tcgaggatta atgccgtcaa gggtggtggg gaggtattta 13200
taggaagaaa tgcggtggag gtgaatggag gcgtcgatca aacccaagag gaaacggttt 13260
gtcgaaatag atgcgttggt cattcgccat aagtttgtga aacggtgagt gtatccttgg 13320
cgtcgtttga aagtttgtga aacggtggcg atgctccgtt aaaggaccgg caacggatcg 13380
agcgatatgt aaatgggagg gcagaaatat tgcatacgcg ccggacacgc gcttatggaa 13440
gtgatagtat ggaagtgata ggattttctt atataataat gatataactt aattttaaat 13500
aacatcgtac tatacgtttc aaaataaatt agagttttaa taaaattacg gaatgattaa 13560
accaattaca aacaaatata agtgtcggtg atataaaatt tgtaacaacg aaaacatttc 13620
aaaaaatctg aatgcggaaa tccataacat gagaatacga ttcgtttgca gtcacgccgg 13680
tttcttcacc ttctgagggt cctgaaaaag gttaataaac aatgggtaag ctaaagttta 13740
gtgatcatac acattacttg tctattatat gagttagata cattcacata tcacatacat 13800
acaagcatac acatattaca aaaaatcctc acacatattg ttggtaccaa ctactattta 13860
gtggaccagg ttattcttgg gggtcaattg tggttaacca taaatgtttg gagtttaggt 13920
tccccttggg ggccgaccat aaacacccat atcatatctc tgtctaggtt ctctttgggg 13980
actgactata gacaactttg ctcagattct ccttggagga taaccatcct cgacgagatt 14040
aacaacggac caccactgtt ttcagcttgt aacctgcaac atcaatagtt gcgatcttcg 14100
gcgacaatga tgaaccaaca tcaacacaac agtagtgccg gtatgttagt cgttggataa 14160
tagaggcatc atctgaccct tttagcggtg acttttgttg cctatgaagg tcttatcagt 14220
tatatatcga ttaggcttgg ttctaggtta taagaaaaaa tgattctcta tccggtatga 14280
catagagaca aaccggtcta cttttataag aaaatcatat ttgaaacaat gtcgtgttaa 14340
taattaatat gtgtatgttt aaatttagaa tatcctgaac ttatatttag ttcgcaaccc 14400
tatgtaagat cgaactaatt ttaaaccgta accagacgtg catgtttgat cagtaataaa 14460
aaggctatca tcgtcgaaaa tattaaaaca aaatataata tcttatttgg cgccatttc 14519
<210> 8
<211> 684
<212> DNA
<213> Taraxacum officinale
<400> 8
atgtcaacgg ttaaaagcaa gttatcaacg cctccaaccg atcatttctc cggtaaccac 60
caactgctac taataaccgc caccgcaaaa acctatatat acaacttcca tcacaattct 120
gtcccctact cccatttcaa tccaaaaaag gctacacaga catatacagc gatggcagat 180
aatggcaaca ccggccgtca aaaggatgac gacggtggcc atgatggggg acgcccaaat 240
ccgaccaccc caccctcccc ttcccgcacc ccgcgaagac ccaggcggac cacatcgccg 300
cctaaacatt ctccgggggc gtcttcaagc accatgccag cgccgcctac tccccctgcg 360
ccgacgggaa tcaccggtgc gtctagttct tctgtgggta ctaatataat ttcatttact 420
ccacccaaaa ccaaaagaac gaagtcggtg atttgtccga tctgtaagaa agatatgtgc 480
catgagaagg cgctgtgtgg ccacatccgg tggcatactc aggaagaaag attggcggcc 540
agcattgcta tagcaagagc gctatcttct aacgttgttg tttctggcaa tggcgatgaa 600
gatgaaggtc catctaaaaa gtataaactc ccggacctga acaagtcacc accgccggag 660
gaggaggacg aggacgctgc ctga 684
<210> 9
<211> 4521
<212> DNA
<213> Taraxacum officinale
<400> 9
ttttcttgcg ggagaggcat gtatgtgtgg cctttttgca ataatactat gcatgggaga 60
tgcataaagt ttatctttat tttgttttag tttaataatg tgtggcaaac tttatatcat 120
ttatgatttc ggccatgtac tgctacatcg ttctaggatg gttattccct agaatttgct 180
tttatatatt ttttttcttc gttaattctt ctcttccttt aacgttattt catgtgaatt 240
tgcgatatca tgatattgca ggaacgtaac ttagaactca acgataaaaa gtgtgttgag 300
atcgtggaag ttgatcgttt atcttgaatt cacaagataa gcaacatgag atagaataat 360
taattactgc ggagatttct ttttagattc atcttattcg ttgtaaattg caatgcaaat 420
ttttaagtag aataacgaaa gcgtgttgtt atttacggtt aaatgcaaaa aatcattaac 480
atatctccat aattgatttg tttagtatgc attctcatgt gttctattgc aaaattctat 540
tttgaaattc aattaaagca atgcatttta gatataaagc aattccatca cttaattata 600
ttgggtagat ttttaccaaa tcactgtcct aaaagaaaaa taaaaaatga aagttcaatg 660
ccatagtgat cccgtcaatt atgtgatcaa tttttaccag agcaaaaccc caaaagttaa 720
agagtataat tgcagaacga caacaatctg aggatgatac taactccggt ataaaccgaa 780
tgcttttgtg gggatttcac taatcaacta tttggctctc atcacaaggc cgtgttaatg 840
gattactatt agttataagg ccatcccaag ttgtttggct atattattac cttttaaaaa 900
ccaaaatgtg taaactgtaa aattacactt ttggttcttg tattttctaa tttttagaat 960
tatgtatttg gttcttgtag tttttttatt tgtatgaatc atattagttt tggtccacga 1020
taacttttat tttctataat tacacttttg gtcattgtag ttttaatttg caagaaaaaa 1080
ataatctatc tttttagtcc atatatatag tttgcaatat tctatatttt aggttcaggt 1140
tatatatctt ttacaaaaat actacttatg attttttaat gaatatatca tatttcgtaa 1200
attacataaa cattttatat aattttattt agtcagaact ataggattcg catgataact 1260
ttgctgaact taattgggct tcatttttaa caaaggataa tgcatcatgc atgacataaa 1320
aatttgtaat gtttctaatg tgcatgcact acaagaacct gacccttcat atacattaat 1380
gaatgattct cttttcttat tatatcttag caaacggttt ggataagagt ctgaacaata 1440
ttaagaattg cactagctag ctagtccatg cagcaagcaa attaattaga ggacttttta 1500
agaggcttga tcacttcatt gtatatctaa tggcactgtg gatatatgat tggattgtgt 1560
gtatgtatgg aatttaatat tactaaaaaa tattaatgac aaacaataac acactaattt 1620
ataacttaca aaagcgtgtg tcattaaaga aaatgtcatg tttcttaaaa tatggagaat 1680
ttactactta cataattatg tgtcataaat ttatgtcatc ttgtgtgata tttatgactt 1740
acaaaaatgt atgttatgtt tatcgtcata aaatctgtaa aaaaaaatag aaacatggag 1800
gatactttca aggtgtagaa acttgacagc ataaacgggt ttattacctt catcaaaact 1860
tatcttactt caatctcttg gagatgcagg ttgattttaa tgctaaatga acttttcaaa 1920
tgtataaata tatatctttt gacggatttg acattgatgg agatgaagat acaccagtga 1980
attcaccttt caatgatcat actgacatac caagtactct tatcccctct ccacctccac 2040
agaacgaacc tcaatttcaa aaacgtaact gccgctgttc taaaaacaga aaattttcaa 2100
aaacgtaact atcatttaaa tatttaatca tttttcaatt ggattcatgc tgtaatagaa 2160
gttactggac tttaaacttc ttgtagctga tgagattcat gaacaacagt ccaaggagat 2220
tttccaaatc aacaagtttc tgatgagaac aaggtaaaaa aaataagttt ttcatttaaa 2280
catatgattt tggttttcaa aagataatta attataggaa ctcttctgaa aagtcccaaa 2340
gtttacccgt aaaattgatt ttggtttttt tttaaattga ataaaaaagt ccataaaatt 2400
ggataaatta ttcgatttag cccaatttac ctagtatatc tggtaaaaat tggttaaatc 2460
gaataattta ggcagtttgt tgggcttttt tgcgcaaaaa cattattttg ggacttaatc 2520
gttaattttc cccaagtttc agtttattta cagtattgga ataaaaccga acctagtgcg 2580
ctaattttgt cttttggtgt ttagaattat ttgggtttat tggtctcggt ttctaaacga 2640
gttcggtttt tggtttcttg attcgatttt tggcttttgg gttcaaccca tggattttcg 2700
atttggtttt ggttttgata atattttttg tgcaatcgtt caaaaaatcc gagttagatt 2760
gggtttaaat gttatcaaaa ccaaaaccga tggtagaacc ccaaaactga acccaatcga 2820
aaatccatgg gttgaacctc aaaaacgaac ccaataaatt ctaaaaacca aaatccaaaa 2880
ttagcggata aaaattttaa aatacatatc tactggtaag gtgtttttga caatttatgt 2940
aaattataga aaaaaatgta ttaagtgttt gctatttaag ataaaaatgg cgttaaaaat 3000
agaattggtt agaaatgaac cattttttta tattttttac ttcatttatt aagtaaatag 3060
tagaggtgaa ttagagatga tctaagatat tttttttctt tcaaaaagag cttcgaaaac 3120
acttttccga cagaaacaag acacaaacct ttttactcgt agaccgacga tttttttaaa 3180
gatcaaagac ctaagaagat ttgtctccca tcttcttaaa acatatgtga gatagacact 3240
tctagcctct tctatgctaa aaaaaagagc tcgcctttag tttggttaat ttgagacaat 3300
ggtctccaca aaaacatagg tagttaggta ctcacgcgcc aagttactaa tctcaaatca 3360
aacataactg tatgatcgtc aacacatttt acaagttaac aaaagagcat tgctttagac 3420
ttaaaaattg ctttgagaaa aaacaaacac ctaagaattt ctaaaagcga tgatttcaac 3480
tctctttaaa atgtatatat atgttacatt ttggtcttta aacttttcaa gaaagtacaa 3540
ttcaactcct attttcaaaa tgttcttaca tgagcgatat gattttaccg tttaaaatgt 3600
aaaccatgat aactatttct aaattccaat ttttgtttcg ttgatcaatc tcattatcat 3660
tcaatgcata caaaacacaa tacaatctgt tttgtcatct tcttcaaggc cgaaagacaa 3720
ctaaacatat gaagactgaa gaggacaatc tagcaaactc aaacttcact ataaatttgc 3780
aattgctatt ggtagccatg aagtaacatt tttgaggagg ttaacaaaat tttaacagta 3840
aattcaataa tccatcacca cgatatgtaa tcatctacac aaacacagat gatgatagtc 3900
atatatgttc gtaatcacac aagcaagaaa tcgcaatgta ctttacaccg tcttctctga 3960
cttgaacacg ggaattggtt gtctaagaag tagattgttt cccacggatt taccaagtac 4020
atttaaatcc ttcacccttt gtatctctct tcctgttttt tgttttgcaa tcaacaaaaa 4080
aaaaagtaaa aatcccatct taaattcacg caaaaaaaca cattgaatca accatcacta 4140
tagtagtttc aagaaaaatc tgcacttcca taatttcttt ctattacacc attatactag 4200
gaaaacttca tagcaatatt atctaaatac aaagcaattt ttactagtaa tattcgtaga 4260
tttgtcaata tatagtgtcc taataaagaa aaaataacgg aactataatg ctctaatagg 4320
tagatttttt ttagtataat gtcctaataa gagaaaaaac aaatgtacaa ttatgtaata 4380
ataagaaaag aaagtaatat gcttaataga caaataaatg aaaccatgtt gctatttctt 4440
gcatctagtc cttatttgaa taatatgtag catattaact acttatgaat tttatatata 4500
aatgataaaa aattccgtga g 4521
<210> 10
<211> 19724
<212> DNA
<213> Taraxacum officinale
<400> 10
ctgacttggc caacattttt tgttcttccg atcccttcgt aacaatcagt ttaagcacaa 60
ttgaattata tggagataaa taggtgaaat ttggaagacg aggatgggag agtgcgaaga 120
gccaaagaag agaaatgtgg gatagcggcc ggccggttgt taatagaagt gcttagatct 180
tggagacctt ctaatgctta tggaggaacc agatgcttca atttgtaaat ctgcgctgcc 240
cacgtagaat attgaaaaag aaaatcaaat cgatcaataa tgcgaccata tactttagga 300
tttcccactt gtataattct cttattcatt ttcaaatctt tatctaaatt aaggatatcg 360
gcaaaaatat aatcctaatt gtttttcaaa atgataagga ttataaggta acttaattct 420
tgttttattc atatttaatt attattgttt tttatcaatg ttttaaaacc cgggttttga 480
gtcaacccgg tcttgtgaaa aatcccgggt cagccggtca gctgatctac cggttcaata 540
tatttaaata aaatataaat tttcatatag aatttgagtt ttaggctaag aaaatcgggt 600
tttgggctaa ggaaaatcgg gtttaagttt tgggctaatg gattttcggg tcaataacgt 660
ttttttccgt tttttagatc ggttcgaccg cgttttgtgt gaaacccggc cgggttgatc 720
cgagtcaata attggcaaca acccggtata tgttgacccg ctcttgtccc cgggtcccgg 780
ttcaaccggt tggatcggcc gggttgaccc tggttttaaa acactgtttt ttatacaata 840
tatttgtttt tgtttttagt tttgtttttg tttttgattt tgtagatata tttttaattt 900
tatttttatt ttgtataaat atagtcatta caaccgatta tttgtaagtt gagtcggtca 960
aaacgaactt tctgataatt cttatacatt tttcggcgaa acatgaatat tttgatcaac 1020
caatatgctt ttttctaggt tttttaaata aaccgcacgt tcaattgcat attcgcaaac 1080
aatagaactc gaaaacccta atcgagcttg ttttatagaa atctgggttg agtgcataca 1140
taataattaa taactgtgaa aaagtttgtt ttaacatgct aaacctatcg atagtcggtt 1200
ataataatta tatgtaaaca aaatcagaaa caataatgat atatgtcacc caaaaaaata 1260
gttactaaat atatacaaaa tcaaaaacaa cagtaatttc ttttacaaaa aaaagagaac 1320
aatagtgact aactttagaa agaaaaaaaa agaactaaat tactcttata aatcattatc 1380
ttttatataa attggtaaat tggtcatgta atctagaatg tgaattgcgg aatgctttag 1440
tcacattttg aacaaactgg taattattaa tttcgataaa agttcttcaa gaatagtttg 1500
gtcacatttt ggacgacaat ggtgtgtttt ccatagtaag caatacgtaa tgagttcgct 1560
taataggaaa atagatctaa agcttaatga aatcatatgt ataagtaaca tatatttttc 1620
acacgttaca ttgtttgtaa cgtcctcaat ttttttaata tatataacta ggtacactct 1680
aaggtcgtgc taagcacgac ccacgaatgg taatttaggt aatttttcat ccttgactaa 1740
aatacattat gctaataaca aagaattaag aatacaatgc tataatgcat aagtaaataa 1800
aagtacatga tatttctcac atttagtctt tatacataga aaacttacac tggccatttg 1860
caaaaaaaca ttaaatctcc ctaaaaagta gagtttacaa aatgaaagct tgtttttata 1920
gcaaacaata atcaatatgt gaagctatag tatagtaacc tactcatttc aacttttagg 1980
cagtgcagta attacttact catgttaatg gtgaaatgca ttaagataaa cttcattcaa 2040
cggattaaat ctgggttcat tataaatgta ctcttagtta ctcctccatg taactgcatt 2100
tagcgaacaa caaaagctct aataaataat atgaaacaca tgctaattta taaatgacat 2160
catgggtata gatgctgtaa catcctaaac ttttaaaata aataataaat catatttggg 2220
atttaaagta atattttaag ataattgata aataagagaa tgaaatacaa agaggcattt 2280
tatttttagc ctttgattta taaaaaagaa aaaaccttga aattggatgt attttgaaat 2340
gggtaagtat tttaaaagat gtgtaagtta agggggataa atggaaatta aggagataag 2400
catttgacca aagtcaaaat agaaggaaca ataagtcggt cgattgtttt cctgtgtgca 2460
tcttcggacg acagagagac agaaaagaac gagcgagaga cagagaggaa tcgagacaga 2520
gagattatcg agagagagac agcgaagtcg ggagagggag accgagaacg accgagacag 2580
gaggaaaacc gagagacaag gggaggacga ggcagcagca gcctaggctg ctgcgttttt 2640
ggcgacagaa aaccaccgcc agcggcggtg gtggtgagcg acggtggtgg tggagccgag 2700
aaggggttgt ggtggtgtta gaagtgatgg ttcatctcaa tttcttgata atttgccctt 2760
ttggaacaat aaattgaagg taactttcaa agctttagat gattatctct tgaaagtaga 2820
aagcaagggt aatttagtct tcaattatgt tatggagaag ggtaataaca cccaaatgct 2880
aacaaaatca tatagattaa tgtgatttga agttatatag ttcatgaatg tgaaggttga 2940
ctttctgcaa tggtcaaaca tggacatgga catgattata gcatattagc aatgtaatag 3000
gaatggaatc accaattcca ttgttgaaaa tagaaaattt gatgtgtgca gaattttgta 3060
aaatttccca aaacaccaac tttaaggctg cataagtcat gcatgaagaa tgcaaatcag 3120
tccattcttt cgcctaacat caagtccttg caacatagat tacacaggaa aaagaaacag 3180
ccaaatccga gttcgtatga ggattgtatg actgttcaac gtttgtcaaa agttgctgtc 3240
atgctgtcaa accagaattt ttctaagtct gggaaatatg cagaaaatgg gtttgaccag 3300
tttttggagc ttcataactt gatttataca tatccaaatc agttgattct tgagcctaaa 3360
ctgtagagaa agatgaggag aaggcatagg aaaaagaatc aacacatttg gatatcgtat 3420
acaacctgtg caaagggttg aagtaggctc aaaacccatt tttgacataa ttgtcaatta 3480
tatgaattgt gggtaacttt tgccgtaatg aaatgttttg acatataagt gtttatattg 3540
acttataaac atgttaagac tataaacttg ataaagtaac atgtcaaaac ccgaaaatgc 3600
ctatttaggc aaagtatgat gcttaattga gaaatcggtc aaatagcata aaaactaaaa 3660
gtgttctaaa tggtccaaga agactttata cataacttac acatcattag cacatgatgt 3720
gtagtacatg tgaggtttga acacttttat acttggaaaa ttgtcaagta gtcaaaaatg 3780
ggacaaaagg gtaaaattgt ccaaaatgag tatatggcta attaaagcaa gttgagctac 3840
tgaaatgaac ttagataagt attataagga ctataaaagt ataatacttg gttaactatg 3900
agttttgaag gtaatgtcaa tttaaagtaa gttaaccaaa agaaagtaag ttgctaattt 3960
tgggatatat gacccaaatg agttatgcat tggtttttag gttgtgtaat acctataatg 4020
agatatgaat tgattatgag acatgtattg atgataatac aaatgatgta atgaaatagg 4080
ttcgttgcct atcgtggagc aaggagcatg cggattagtg tagctagtct tacctagcta 4140
ctaaggtgag tacgtgtgga ttgttttccc attttcgggt acatggaaaa ataatatttt 4200
tataatgaag tgcctaatgt tttgaaagaa agaatgaatg aaaatacttg atgcttttga 4260
aagaatgttt taaaagaaag taatgttttg aaagaaaaag gtttgatgct ttacgtttga 4320
aaatgaatat gtcttgatat taatgaatgg aatgggtaaa gcatgaatga tataatatgt 4380
aataataaga tgctaaggat ggttatgttt atgatgttca cgtgaatata agaatgttgc 4440
tcgatctaag atgtcccggg tagggattca gaggagccta tcggggtggt acctcccctt 4500
cgcgagatag gttacctaat gtaaatgatg tcaatgtaat ctgatttgtt ctttttgcat 4560
tggtacagac ttggggtata tcagacccaa gtataatatg atatggccat atgataagat 4620
gattaaaaga gtatgtgaaa tgttaaaggt tatgtaaaat gttatatgaa attattcaaa 4680
gagatcttta tggttattat gtctatgaaa aggaagcata taatgttata gatattttat 4740
gtctaaaccc acgtagctca ccagactagt tgtctgacgt atttattttt atgccatgta 4800
tttcaggtta tacacgagga tagactgatg atcgatagaa gctagatgtt acgacggata 4860
gacggagtgg agctttcaag ttctattgta atgatcttac cgtaaattct tatttgcttt 4920
gctatggttg ctgtttgata tataattggt aacacccgag aattgtttta tatatattca 4980
aaaagttttt atttgggacg gtttttgtaa acatgaccaa gtgtcatgaa atatttttaa 5040
aaccatagcg tttttaaaac gataaaatga ggggtgttac agatgcgata atatatctta 5100
ggattatgtc aaatatttat gccaattata catgagagag caagtttata agagttttag 5160
ttattctatc attggatcaa tatagatacg tattaatagt agacagagat gcatgaatat 5220
tatagcagag aactttttgc aaaacctaac ctcactgcat gttgcattca tcaaccccac 5280
ttgtagttgt aacaccatga atatgccaat taagaaatat catgaggtat attgtactgg 5340
aaatgacaaa aagtactaat ttacaataga acgcagtcaa acttggttat tttgtaaatt 5400
ttaagaagca atttacaaaa aataaaacat aataaagaaa aacataacag taaatcacag 5460
acattgctca tggaagagtg actgatcaaa ctgaccgttt attttcttgt ttcttgtaca 5520
actaccaaac gttcctgtat aaccaccaaa tagtcccaga ctttagaaaa tagaacagaa 5580
aaccacagat atacatacat acagccatac attataattt ttaattgaaa aaaatgcata 5640
gaaaaccagt ctcaaataaa gaaccaaata aagaagatac aaatgaaacc tggttatcga 5700
ataaaagagt tagagataat gatttcagca tttgcactca tgtaaacgat atgccattgg 5760
gatccgttgg aaagggtatc gtgaagtgat taaggagaag cggggtgtta gggttttttt 5820
tgcagtttat tgaacaggtt gatcaaattt tgcgccggta gggttaattc cttttcgaac 5880
gggtttctat ctgggtcttt catcaaaatc actcatgttt gtgtctctaa cctttagttg 5940
cagatggata tcagatcaac ggtttcaggc taaatggggg aacaaataac aagaagaaag 6000
cggtgacatc tgtgttcgta ttcgtatggt ggaaggggaa gagtggcgta tgaattggtc 6060
aatgtaaaac atctggaaac aacttctccc aacatgtaaa aggatccgga aaatatgaaa 6120
atgacgatta gagatttcgt tcggatattg ggaatatggt cgcgtctgat tgccctacgg 6180
tgcccaccgt ttgggtttct tgatttgtcg tttagataga tgtcggaggt ctaaattcac 6240
aatcctgttc ttccgcggta attgaaattg ggtaaggcta attgtttaga tcaaatggca 6300
gaggacgatg tgaagttgcg ttgccctata attggctctc gtgaaattga attgcaaatt 6360
tcacaacgat gttagcaaca ataatcgtcg ctctcgcatc ccataccgtc gcaacgtatg 6420
atggtgatcg agattgagag atttagggct agatgtggcg ttggcggaca taggttgtgg 6480
ttgacggaat gaggagaaga cgtgagcgtc gattgaggta gcaacaggcg gagggttaac 6540
ggtgtcatca tggtgaaagg tcgcgcgtcg ttttatggtc gatagatcat ggacgttgtg 6600
agtgaatgcc ggcgcaaaca cggtggagga aagtctatgc tttagttgac ggggcggttt 6660
tctctcggag acggaaagag gacgtcgaaa gaaaggccgg ttggagaaaa tatattaaac 6720
gcgccgacac acgctatcag agaagaccgg ttttcttata taataatgac taggtacgtg 6780
caagggtcgt gctaagcacg acccatggag gcttcttgag aaaattgtaa acacatatca 6840
gataatagcg acgaacttta ctgttattga tatataaagt gattaagcat ggttggccct 6900
aagttaaaac tcaaactact gtaagtatgt aatggtaata atttatcctt ttcagctatc 6960
gatcttagat tgctctgtag tgttgtgtct tagtgctttt cacggttctt ttatatgcat 7020
atagtcacat aaaatcagac acacacatag aataacccac agacaatcac ataaaaatca 7080
gtctgatgag gtattaattt aagcaataag gattgatatt caagcaatta gaggagaata 7140
atatattaat ttcacgtact cctattgtag ttattgtgac aaaacccaac tgtttaattg 7200
tgccattgca aaaagtccac ctgtttaatt tctttatcaa catgtcgttg catcctttta 7260
catgcaaata tttcagaaca tcttcctata atgacccttg agatgaactt tttagaaata 7320
ttgttgctgg ttacattatc agaaaaaaca gagcttatga aaaaagacca aacatagtaa 7380
ctcgtacttc atgtctggta gttcttttga atatttccgt cttttgttcg aaacaatttg 7440
gttgaactgt agtagatctt tcaaactgta tacaacttca caatatgttt tgtaagttca 7500
ttacttctag atcctttcaa aagaagaggt caaactcaaa atttgcataa accagattta 7560
ttgaaggcta attagattag tgttgtaggg ctttttggat ataaataaaa atgaaacata 7620
ggtgctgcaa caggtgcgaa agaatttggt tggacagtag taggtctttc aaactgtata 7680
caactttaca atatgttttg taagttcatt atttctagat cctttcaaaa aaaaggaggt 7740
caaactaaaa aaatttctaa accagattta ttgaaggctg attagattaa cattgcaggg 7800
ctttttggga ataaatataa atgaaagata ggtccaacag aaacgaagga ggtgtgaatt 7860
atgcaacaaa tggaaacaca caattatcgt agtacacatg taattaacag ggatcaattt 7920
ccatttattg ttccatcaat tgccaaattt aaaagataac aaatgcatcc tttttgaatg 7980
gaaaagtatg atacataaaa ccagagagag gtatacattt agtccagaaa cggtggaaat 8040
tactttacct agagttgcgt gaatcaaatg gaaaatcaga ggaagagacc ctaaatcgca 8100
atccatcttc tattccctat agaatcaaat ggaaatttct tacttccgac ctttttcatt 8160
tccacaagtt tgtaatcgat aagatcataa aagaaaaccg atttacatta gcaaatcttt 8220
aaggcacttg tttgtagttc tcaagtatgc gttacgtatg aaattctaga cagaaaaggg 8280
acgacaggga aaataatggt ggaataaata aacaatcaag aatcaaaaga taataatcct 8340
taaaactaac aaaaattcta tcaataagcc ctacaaatga gaaatatgac ggtggagact 8400
cacctgattg cagtggcgga agctcttggg ggctttggta ggccatggcc ccctgttttt 8460
tggaaaattt tcaaaaaaaa aaaaattata tatattatat cattgaataa gtctagactc 8520
aggaaactat gaaagattag aaatagatca agttatctaa atcaaatctc gaaaaatatg 8580
gaaggaatca aaacacaata tttatctcat tgtgaaatca aaaaccctat aaaaaacaca 8640
aatctaacct acaatttaaa tcaaaaacac cataaaaatc aacctaccgt cggattcagc 8700
cgtcgtcgtc ttcttcaccg gaccaccatc caccgccatt gcttcaatcg tgctttctcg 8760
tcaatagctt agtggcttgt tcagctttct tgctctacca atgattttta tacactttcc 8820
agcaccaccg atttgttgaa cggtaatatt atgctccttt atatttttga aatctttgaa 8880
agttcatctg gttttctgtc tctgtaaact gtaaagaaac ttttatgttt tgttaagaaa 8940
atagagagat agagaagaaa gagtattgac gtgcttttcc catatgtctc gaatgccaca 9000
ttaggaaaag acaaaaagaa aacaattagt acattacgct agtgtgatag tactgtatga 9060
ctattagtat gatttacagc tttaataggg attcatagaa aagaacaaaa taaactactt 9120
ataaattaaa aatcgagctt tatttttaat tagtttaaat aaatatttag ctataacata 9180
atatattata tgttttgtat ttataatcaa atataattat ttgtattaca gattagtttg 9240
tataatcaaa tataatgatt catgtttcga aaagattgtt taaatatttt tatatgccat 9300
gttttttaac tgaattatat ttttttgtat gattttcttt acaatttagt ttttctttac 9360
aatttagttt tggcccccct tgtttttttg tttgtgttcc gcccctgcct gattgatgcg 9420
agaaaaggaa agccgccgac atttgtgctt cttcccgcgt taagaaaaat cgcagatgac 9480
attacgattc tttttctaca gaggaatagg ggggttgcga ctttgcgtga tttgttgtta 9540
atgagtgatg cgtcgtccgt tttgatcggt agcccgtgga gactgattta tttgacagat 9600
agatgtcgtt ctcgttgatg atgcaggtgt aattggcggc gttggtgaag cgaagatggc 9660
atagggaatg gctttgcaga ggagaaaagg agtttcacaa atatggatta ttgaatcgtc 9720
gtttcagagt cgcgatggtt agtaatcgat ggatttgggt ttgcacctgt tcaaggggtg 9780
gtgacgtgtc ttactgtaag aaagagtcgc gatggttagt aatcgacgat ggatttgggt 9840
ttgcaacatc gttgacggaa acgacgcggc gatggagaga ggcggtagag ggaatgaaac 9900
gggagattta ccttcaaccg caaacatgtc ttcgttcctc gaaaatcata tgcgcccttt 9960
atgaaaattc cttcacgcgc cgtacacgcg ctcagaaaaa acatagggtt ttcttatata 10020
ataatgacta ggaacgttta tgcgccgtga taatcacggc ccatgagtgg tacaaatgtt 10080
gtaaaactta gaaaatggga aagttgaatg tacaatgtta tgatttattt tattcttcat 10140
agcacaatac ttttgcttta gtgaaagcag ttaaaaagtg gatgtgtcgt tagcaactat 10200
aaacttttaa caaaatatta catacagaac atagaaataa gataaacact aaaaaaatac 10260
aaaatagccg tgttcaacac ggcccacaga atgcaatttg gatatttcta aggaggtgta 10320
aaaaatattg ctattatggg agctaattct aggtatattt tgttactcac atgacatgaa 10380
tattgagtaa ttttgtaaaa ttggtaattt agcttagatt ttcccacaaa gatgagtgtt 10440
aaaactatca aggaggatat atggtacttc ctcaacaaag atgattttga aatatctata 10500
ttttaaatgt aagaactact atcacatgag acaaatattg atattactgg ttaggattac 10560
aacattgata tgtaaaatat tttggtaatg taaattttct aaaaaagctt ggaagtagaa 10620
tgcaacaaca atatatgcta gtattacatg ttgggtgatt ttatgaatgt acttacgtgt 10680
tgatccttat ttaccgtact tgaataattt tatctgatca atgctttacc tatatggcta 10740
tatctagcaa tgtgcattat tcacaaatta tggagtgtaa attcatatag acgtatatga 10800
cagtggatga aaaggaaaac aagtaatttg attgaatcac tgagaaaaaa gcaaaagata 10860
catgcatata aaatgttaca tcaaaataca gaagataaat gaaagaaaga taataatgtg 10920
gagcattcaa aataagtaag aatgtaagtc cttgatagct cgaaattctt ttaagtcttc 10980
cgtgatagcc taatgtttat aatagaagag acattcaaaa tgaataagaa ttatatcacc 11040
taacagtatt taatcctact ctacttggcc attgagcata ttgctattgt taaaaattat 11100
gtccaaatcg catttaactg ttagatgcaa ttctcattgc cgattacact tatatgtatg 11160
tagtaaatta taattaactc tttgtcctgg tcatttgatg tgtcatggat gtttgtaagg 11220
tacaaataat agattacaat aacttaaata taaaattgag ttaatattcc agaattcaca 11280
aaatgtgtaa taacacttat ttataatttc cacgtaatta atgattaact acttagaatc 11340
ctacaagttt ttgtttatca cctttcatat gatcgtataa atgcaacaac ccttaatgca 11400
tttatagttg ttggtctgcc acgaacacct ttttctgttg attatttgtc attattatgt 11460
taaaaaatgt ttgaaaaatg ttaaatggga ttttgttatg gcaaagagat gaacttctcc 11520
aatattgtag ttgcataaat attgccagtt ccatcataca gaataatacc tcttatattg 11580
ggcaatagaa ttatagaaac aataagggtt aaccgactta catgtattga tctctgattc 11640
ttctttacac actggacaaa ttagtctatc atgtcttgca caaccctgtt aaagaaaaca 11700
gtgaaaattg accacgaagg tgtatcctct agtaccaaca ttacatctat cattttaaaa 11760
gggaaatgta tacttatcat atatatctaa tggacgttgc ctttcctcca attgtcactc 11820
ttacatctaa tggaaattat gatcacaatt tacttttaac agcatagata cattcatttg 11880
ttggtaagac acttttggcg gattcattgt attaatcttt catgtaaaaa ttctcattca 11940
agatattagg tttttgttag aacccttgtt atttgaacat gcaacctgaa ttctgattgt 12000
tcccaacacc aacattaata atacctcttc cactagacca tcattccacc ttcaggccct 12060
tgtgttcatt attataagca ctattcggtt tagttctttt atgagtccta tatttacatc 12120
cttaaccatt ttaacaagtc catcatattg ggaattttaa tataaatacc caatgttatt 12180
catttttgtc gcttcagtag atattgacca tactaaagat gagtatttaa tactcataca 12240
acatgaaaga aaaaaatgaa aaactaacag tcacttatgt agccccaatc tccatacaaa 12300
tagcgcttta agaataaacc aatttcaggt gaaattgtag aaaggacata aatctgtaca 12360
tgaaaaaaca ttaacagtta cacataaatt atttgcacag agataaaaag cacaatatgt 12420
gatgagaaga tttaaaagta ggaaagtgaa caaaaataac catcaaaatc caatgttagt 12480
aataaagaaa atgtacgagc gattcataac attcatccaa aacccatatg cgatgttgaa 12540
tgcatatctg gataaaagaa tttaataaac aataagcgcc tgtaaaatca ataaacaaaa 12600
tcaataaaaa tagacctaaa aatgcaaccg acttattgtt tctatacacc aataaatgat 12660
tttcacgaat tttatccaaa gaaagataaa aagtcataaa attgaacggg cttaccgatt 12720
tgaatcgtgg tctgtatctg ccacgcagat cagataacaa tcataatcta catcaaagat 12780
ttcagaaaaa tatgaaactt ccgccattac taatttgata tgaatttctc agagcctaaa 12840
acaaacctaa attactaaaa atcaaacaca ggtttttgac gtttaccttg atttgtgaaa 12900
ataggcattt gggttttttc gtaggatttt ctatgaaaga agccatcaag tttatgacat 12960
tgtaaattaa agcaaagtat gaatatatga aggataggaa aagagatcgg attgtgatga 13020
tccggagaga aaaatacaca aaccagagag aagcgatgtt gtgattgtgg tgtattaatc 13080
tgtttttaaa cctctaacca cttcttgtaa gtccgtaatc agacgaaatg cgcttgaata 13140
ataagtcttt cacgccagtg tcgaggatta atgccgtcaa gggtggtggg gaggtattta 13200
taggaagaaa tgcggtggag gtgaatggag gcgtcgatca aacccaagag gaaacggttt 13260
gtcgaaatag atgcgttggt cattcgccat aagtttgtga aacggtgagt gtatccttgg 13320
cgtcgtttga aagtttgtga aacggtggcg atgctccgtt aaaggaccgg caacggatcg 13380
agcgatatgt aaatgggagg gcagaaatat tgcatacgcg ccggacacgc gcttatggaa 13440
gtgatagtat ggaagtgata ggattttctt atataataat gatataactt aattttaaat 13500
aacatcgtac tatacgtttc aaaataaatt agagttttaa taaaattacg gaatgattaa 13560
accaattaca aacaaatata agtgtcggtg atataaaatt tgtaacaacg aaaacatttc 13620
aaaaaatctg aatgcggaaa tccataacat gagaatacga ttcgtttgca gtcacgccgg 13680
tttcttcacc ttctgagggt cctgaaaaag gttaataaac aatgggtaag ctaaagttta 13740
gtgatcatac acattacttg tctattatat gagttagata cattcacata tcacatacat 13800
acaagcatac acatattaca aaaaatcctc acacatattg ttggtaccaa ctactattta 13860
gtggaccagg ttattcttgg gggtcaattg tggttaacca taaatgtttg gagtttaggt 13920
tccccttggg ggccgaccat aaacacccat atcatatctc tgtctaggtt ctctttgggg 13980
actgactata gacaactttg ctcagattct ccttggagga taaccatcct cgacgagatt 14040
aacaacggac caccactgtt ttcagcttgt aacctgcaac atcaatagtt gcgatcttcg 14100
gcgacaatga tgaaccaaca tcaacacaac agtagtgccg gtatgttagt cgttggataa 14160
tagaggcatc atctgaccct tttagcggtg acttttgttg cctatgaagg tcttatcagt 14220
tatatatcga ttaggcttgg ttctaggtta taagaaaaaa tgattctcta tccggtatga 14280
catagagaca aaccggtcta cttttataag aaaatcatat ttgaaacaat gtcgtgttaa 14340
taattaatat gtgtatgttt aaatttagaa tatcctgaac ttatatttag ttcgcaaccc 14400
tatgtaagat cgaactaatt ttaaaccgta accagacgtg catgtttgat cagtaataaa 14460
aaggctatca tcgtcgaaaa tattaaaaca aaatataata tcttatttgg cgccatttca 14520
tgtcaacggt taaaagcaag ttatcaacgc ctccaaccga tcatttctcc ggtaaccacc 14580
aactgctact aataaccgcc accgcaaaaa cctatatata caacttccat cacaattctg 14640
tcccctactc ccatttcaat ccaaaaaagg ctacacagac atatacagcg atggcagata 14700
atggcaacac cggccgtcaa aaggatgacg acggtggcca tgatggggga cgcccaaatc 14760
cgaccacccc accctcccct tcccgcaccc cgcgaagacc caggcggacc acatcgccgc 14820
ctaaacattc tccgggggcg tcttcaagca ccatgccagc gccgcctact ccccctgcgc 14880
cgacgggaat caccggtgcg tctagttctt ctgtgggtac taatataatt tcatttactc 14940
cacccaaaac caaaagaacg aagtcggtga tttgtccgat ctgtaagaaa gatatgtgcc 15000
atgagaaggc gctgtgtggc cacatccggt ggcatactca ggaagaaaga ttggcggcca 15060
gcattgctat agcaagagcg ctatcttcta acgttgttgt ttctggcaat ggcgatgaag 15120
atgaaggtcc atctaaaaag tataaactcc cggacctgaa caagtcacca ccgccggagg 15180
aggaggacga ggacgctgcc tgattttctt gcgggagagg catgtatgtg tggccttttt 15240
gcaataatac tatgcatggg agatgcataa agtttatctt tattttgttt tagtttaata 15300
atgtgtggca aactttatat catttatgat ttcggccatg tactgctaca tcgttctagg 15360
atggttattc cctagaattt gcttttatat attttttttc ttcgttaatt cttctcttcc 15420
tttaacgtta tttcatgtga atttgcgata tcatgatatt gcaggaacgt aacttagaac 15480
tcaacgataa aaagtgtgtt gagatcgtgg aagttgatcg tttatcttga attcacaaga 15540
taagcaacat gagatagaat aattaattac tgcggagatt tctttttaga ttcatcttat 15600
tcgttgtaaa ttgcaatgca aatttttaag tagaataacg aaagcgtgtt gttatttacg 15660
gttaaatgca aaaaatcatt aacatatctc cataattgat ttgtttagta tgcattctca 15720
tgtgttctat tgcaaaattc tattttgaaa ttcaattaaa gcaatgcatt ttagatataa 15780
agcaattcca tcacttaatt atattgggta gatttttacc aaatcactgt cctaaaagaa 15840
aaataaaaaa tgaaagttca atgccatagt gatcccgtca attatgtgat caatttttac 15900
cagagcaaaa ccccaaaagt taaagagtat aattgcagaa cgacaacaat ctgaggatga 15960
tactaactcc ggtataaacc gaatgctttt gtggggattt cactaatcaa ctatttggct 16020
ctcatcacaa ggccgtgtta atggattact attagttata aggccatccc aagttgtttg 16080
gctatattat taccttttaa aaaccaaaat gtgtaaactg taaaattaca cttttggttc 16140
ttgtattttc taatttttag aattatgtat ttggttcttg tagttttttt atttgtatga 16200
atcatattag ttttggtcca cgataacttt tattttctat aattacactt ttggtcattg 16260
tagttttaat ttgcaagaaa aaaataatct atctttttag tccatatata tagtttgcaa 16320
tattctatat tttaggttca ggttatatat cttttacaaa aatactactt atgatttttt 16380
aatgaatata tcatatttcg taaattacat aaacatttta tataatttta tttagtcaga 16440
actataggat tcgcatgata actttgctga acttaattgg gcttcatttt taacaaagga 16500
taatgcatca tgcatgacat aaaaatttgt aatgtttcta atgtgcatgc actacaagaa 16560
cctgaccctt catatacatt aatgaatgat tctcttttct tattatatct tagcaaacgg 16620
tttggataag agtctgaaca atattaagaa ttgcactagc tagctagtcc atgcagcaag 16680
caaattaatt agaggacttt ttaagaggct tgatcacttc attgtatatc taatggcact 16740
gtggatatat gattggattg tgtgtatgta tggaatttaa tattactaaa aaatattaat 16800
gacaaacaat aacacactaa tttataactt acaaaagcgt gtgtcattaa agaaaatgtc 16860
atgtttctta aaatatggag aatttactac ttacataatt atgtgtcata aatttatgtc 16920
atcttgtgtg atatttatga cttacaaaaa tgtatgttat gtttatcgtc ataaaatctg 16980
taaaaaaaaa tagaaacatg gaggatactt tcaaggtgta gaaacttgac agcataaacg 17040
ggtttattac cttcatcaaa acttatctta cttcaatctc ttggagatgc aggttgattt 17100
taatgctaaa tgaacttttc aaatgtataa atatatatct tttgacggat ttgacattga 17160
tggagatgaa gatacaccag tgaattcacc tttcaatgat catactgaca taccaagtac 17220
tcttatcccc tctccacctc cacagaacga acctcaattt caaaaacgta actgccgctg 17280
ttctaaaaac agaaaatttt caaaaacgta actatcattt aaatatttaa tcatttttca 17340
attggattca tgctgtaata gaagttactg gactttaaac ttcttgtagc tgatgagatt 17400
catgaacaac agtccaagga gattttccaa atcaacaagt ttctgatgag aacaaggtaa 17460
aaaaaataag tttttcattt aaacatatga ttttggtttt caaaagataa ttaattatag 17520
gaactcttct gaaaagtccc aaagtttacc cgtaaaattg attttggttt ttttttaaat 17580
tgaataaaaa agtccataaa attggataaa ttattcgatt tagcccaatt tacctagtat 17640
atctggtaaa aattggttaa atcgaataat ttaggcagtt tgttgggctt ttttgcgcaa 17700
aaacattatt ttgggactta atcgttaatt ttccccaagt ttcagtttat ttacagtatt 17760
ggaataaaac cgaacctagt gcgctaattt tgtcttttgg tgtttagaat tatttgggtt 17820
tattggtctc ggtttctaaa cgagttcggt ttttggtttc ttgattcgat ttttggcttt 17880
tgggttcaac ccatggattt tcgatttggt tttggttttg ataatatttt ttgtgcaatc 17940
gttcaaaaaa tccgagttag attgggttta aatgttatca aaaccaaaac cgatggtaga 18000
accccaaaac tgaacccaat cgaaaatcca tgggttgaac ctcaaaaacg aacccaataa 18060
attctaaaaa ccaaaatcca aaattagcgg ataaaaattt taaaatacat atctactggt 18120
aaggtgtttt tgacaattta tgtaaattat agaaaaaaat gtattaagtg tttgctattt 18180
aagataaaaa tggcgttaaa aatagaattg gttagaaatg aaccattttt ttatattttt 18240
tacttcattt attaagtaaa tagtagaggt gaattagaga tgatctaaga tatttttttt 18300
ctttcaaaaa gagcttcgaa aacacttttc cgacagaaac aagacacaaa cctttttact 18360
cgtagaccga cgattttttt aaagatcaaa gacctaagaa gatttgtctc ccatcttctt 18420
aaaacatatg tgagatagac acttctagcc tcttctatgc taaaaaaaag agctcgcctt 18480
tagtttggtt aatttgagac aatggtctcc acaaaaacat aggtagttag gtactcacgc 18540
gccaagttac taatctcaaa tcaaacataa ctgtatgatc gtcaacacat tttacaagtt 18600
aacaaaagag cattgcttta gacttaaaaa ttgctttgag aaaaaacaaa cacctaagaa 18660
tttctaaaag cgatgatttc aactctcttt aaaatgtata tatatgttac attttggtct 18720
ttaaactttt caagaaagta caattcaact cctattttca aaatgttctt acatgagcga 18780
tatgatttta ccgtttaaaa tgtaaaccat gataactatt tctaaattcc aatttttgtt 18840
tcgttgatca atctcattat cattcaatgc atacaaaaca caatacaatc tgttttgtca 18900
tcttcttcaa ggccgaaaga caactaaaca tatgaagact gaagaggaca atctagcaaa 18960
ctcaaacttc actataaatt tgcaattgct attggtagcc atgaagtaac atttttgagg 19020
aggttaacaa aattttaaca gtaaattcaa taatccatca ccacgatatg taatcatcta 19080
cacaaacaca gatgatgata gtcatatatg ttcgtaatca cacaagcaag aaatcgcaat 19140
gtactttaca ccgtcttctc tgacttgaac acgggaattg gttgtctaag aagtagattg 19200
tttcccacgg atttaccaag tacatttaaa tccttcaccc tttgtatctc tcttcctgtt 19260
ttttgttttg caatcaacaa aaaaaaaagt aaaaatccca tcttaaattc acgcaaaaaa 19320
acacattgaa tcaaccatca ctatagtagt ttcaagaaaa atctgcactt ccataatttc 19380
tttctattac accattatac taggaaaact tcatagcaat attatctaaa tacaaagcaa 19440
tttttactag taatattcgt agatttgtca atatatagtg tcctaataaa gaaaaaataa 19500
cggaactata atgctctaat aggtagattt tttttagtat aatgtcctaa taagagaaaa 19560
aacaaatgta caattatgta ataataagaa aagaaagtaa tatgcttaat agacaaataa 19620
atgaaaccat gttgctattt cttgcatcta gtccttattt gaataatatg tagcatatta 19680
actacttatg aattttatat ataaatgata aaaaattccg tgag 19724
<210> 11
<211> 227
<212> PRT
<213> Taraxacum officinale
<400> 11
Met Ser Thr Val Lys Ser Lys Leu Ser Thr Pro Pro Thr Asp His Phe
1 5 10 15
Ser Gly Asn His Gln Leu Leu Leu Ile Thr Ala Thr Ala Lys Thr Tyr
20 25 30
Ile Tyr Asn Phe His His Asn Ser Val Pro Tyr Ser His Phe Asn Pro
35 40 45
Lys Lys Ala Thr Gln Thr Tyr Thr Ala Met Ala Asp Asn Gly Asn Thr
50 55 60
Gly Arg Gln Lys Asp Asp Asp Gly Gly His Asp Gly Gly Arg Pro Asn
65 70 75 80
Pro Thr Thr Pro Pro Ser Pro Ser Arg Thr Pro Arg Arg Pro Arg Arg
85 90 95
Thr Thr Ser Pro Pro Lys His Ser Pro Gly Ala Ser Ser Ser Thr Met
100 105 110
Pro Ala Pro Pro Thr Pro Pro Ala Pro Thr Gly Ile Thr Gly Ala Ser
115 120 125
Ser Ser Ser Leu Gly Thr Asn Ile Ile Ser Phe Thr Pro Pro Lys Thr
130 135 140
Lys Arg Thr Lys Ser Val Ile Cys Pro Ile Cys Lys Lys Asp Met Cys
145 150 155 160
His Glu Lys Ala Leu Cys Gly His Ile Arg Trp His Thr Gln Glu Glu
165 170 175
Arg Leu Ala Ala Ser Ile Ala Ile Ala Arg Ala Leu Ser Ser Asn Val
180 185 190
Val Val Ser Gly Asn Gly Asp Glu Asp Glu Gly Pro Ser Lys Lys Tyr
195 200 205
Lys Leu Pro Asp Leu Asn Lys Ser Pro Pro Pro Glu Glu Glu Asp Glu
210 215 220
Asp Ala Ala
225
<210> 12
<211> 3217
<212> DNA
<213> Taraxacum officinale
<400> 12
gttgaaagga aggatcggca gagaacaaga ttgagagagt tttatcgaag agaagagatg 60
gttgtgttga ttgacggcag agttcagcaa tggcagatat gcgtgtttaa agcgcagttt 120
gtggacagtt tgtccaaagg ccttcaagtg ggagattgtt gagatgaaga cccttttgac 180
caagtactta aatgtgattg gttgcatggg aaagagcttt acacttcatt ggttgaaaat 240
gaagcatgtt atttaatagt acgtctccac ttgttcttta tgaaggatga agacagccac 300
caagtcttta tgaatgtttc aaggagaaga caatgattag atgctgcctg ctcccttact 360
tcttcactat aaataggtgc ttaaggcacc tctcttctta caccaaacaa cacacaagtc 420
acagattaca aagattgaga gagttttcat tttctctcta gagtttgttt tgtcaaaagt 480
tttgtgtgca gattaattag ttgagagaca ctcctaatta atataaccac ttacacttgt 540
cattgattag cattctagtg tcagagagtt ctgagtgatt gtaacctttg tgtgatttat 600
atataaatct ctctcttgtt cgccagtgga cgtagctgat tatcctgatc agtgaaccac 660
ttaaaatatc gtgttgatat tatttatctg cagattattt taccagatag ttttattccg 720
caccgcaaag cataccgtcc attatttgtc agacgtacct actctcagct catcccagca 780
ggagttagac ctgcaggcag ccagccgtat ttcacaacag tgaattatgc aacaaatgga 840
aacacacaat tatcgtagta cacatgtaat taacagggat caatttccat ttattgttcc 900
atcaattgcc aaatttaaaa gataacaaat gcatcctttt tgaatggaaa agtatgatac 960
ataaaaccag agagaggtat acatttagtc cagaaacggt ggaaattact ttacctagag 1020
ttgcgtgaat caaatggaaa atcagaggaa gagaccctaa atcgcaatcc atcttctatt 1080
ccctatagaa tcaaatggaa atttcttact tccgaccttt ttcatttcca caagtttgta 1140
atcgataaga tcataaaaga aaaccgattt acattagcaa atctttaagg cagtttgtag 1200
ttctcaagta tgcgttacgt atgaaattct agacagaaaa gggacgacag ggaaaataat 1260
ggtggaataa ataaacaatc aagaatcaaa agataataat ccttaaaact aacaaaaatt 1320
ctatcaataa gccctacaaa tgagaaatat gacggtggag actcacctga ttgatgcgag 1380
aaaaggaaag ccgccgacat ttgtgcttct tcccgcgtta agaaaaatcg cagatgacat 1440
tacgattctt tttctacaga ggaatagggg ggttgcgact ttgcgtgatt tgttgttaat 1500
gagtgatgcg tcgtccgttt tgatcggtag cccatggaga ctgatttatt tgacagatag 1560
atgtcgttct cgttgatgat gcaggtgtaa ttggcggcgt tggtgaagcg aagatggcat 1620
agggaatgga tttgcagagg agaaaaggag tttcacaaat atggattatt gaattgtcgt 1680
ttcagagtcg cgatggttag taatcgatgg atttgggttt gcacctgttc aaggggtggt 1740
gacgtgtctt actgtaagaa agagtcgcga tggttagtaa tcgacgatgg atttgggttt 1800
gcaacatcgt tgacggaaac gacgcggcga tggagagagg cggtagaggg aatgaaacgg 1860
gagatttacc ttcaaccgca aacatgtctt cgttcctcga aaatcatatg cgccctttat 1920
gaaaattcct tcacgcgccg tacacgcgct cagaaaaagc atagggtttt cttatataat 1980
aatgatataa cttaatttta aataacatcg tactatacgt ttcaaaataa attagagttt 2040
taataaaatt acggaatgat taaacctatt acaaacaaat ataagtgtcg gtaatataaa 2100
atttgtaaca acgaaaacat ttcaaaaaat ctgaatgcgg aaatccataa catgagaata 2160
cgattcgttt gcagtcacgc cggtttcttc accttctgag ggtcctgaaa aaggttaata 2220
aacaatgggt aagctaaagt ttagggatca tacacattac ttgtctatta tatgagttag 2280
atacattcac atatcacata catacaagca tacacatatt acaaaaaatc ctaacacata 2340
tggttggtac caactactat ttagtggacc aggttattct tgggggtcaa ttgtggttaa 2400
ccataaatgt ttggagttta ggttctcttt gggggccgac cataaacacc catatcatat 2460
ctctgtctag gttctctttg gggactgact atagacaact ttgctcagat tctccttgga 2520
ggataaccat cctcgacgag attaacaacg gaccaccact gttttcagct tgtaacctgc 2580
aacatcaata gttgcgatct tcggcgacaa tgatgaacca acatcaacac aacagtagtg 2640
ccggtatgtt agtcgttgga taatagaggc atcatctgac ccttttagcg gtgacttttg 2700
ttgcctatga aggtcttatc agttatatat cgattaggct tggttctagg ttataagaaa 2760
aaatgattct ctatccggta tgacatagag acaaaccggt ctacttttat aagaaaatca 2820
tactcgtatt tgaaacgaaa atcatatttg aaacaatgtc gtgttaataa ttaatatgtg 2880
tatgtttaaa tttagaatat cctgaactta tatttagttc gcaaccctat gtaagatcga 2940
actaatttta aaccgtaacc agacatgcat gtttgatcag taataaaaag gctatcatcg 3000
tcgaaaatat taaaacaaaa tataatttct tatttggcgc catttcatgt caacggttaa 3060
aagcaagtta tcaacgcctc caaccgatca tttctccggt aaccaccaac tgctactaat 3120
aaccgccacc gcaaaaacct atatatacaa cttccatcac aattctgtcc cctactccca 3180
tttcaatcca aaaaaggcta cacagacata tacagcg 3217
<210> 13
<211> 684
<212> DNA
<213> Taraxacum officinale
<400> 13
atgtcaacgg ttaaaagcaa gttatcaacg cctccaaccg atcatttctc cggtaaccac 60
caactgctac taataaccgc caccgcaaaa acctatatat acaacttcca tcacaattct 120
gtcccctact cccatttcaa tccaaaaaag gctacacaga catatacagc gatggcagat 180
aatggcaaca ccggccgtca aaaggatgac gacggtggcc atgatggggg acgcccaaat 240
ccgaccaccc caccctcccc ttcccgcacc ccgcgaagac ccaggcggac cacatcgccg 300
cctaaacatt ctccgggggc gtcttcaagc accatgccag cgccgcctac tccccctgcg 360
ccgacgggaa tcaccggtgc gtctagttct tctctgggta ctaatataat ttcatttact 420
ccacccaaaa ccaaaagaac gaagtcggtg atttgtccga tctgtaagaa agatatgtgc 480
catgagaagg cgctgtgtgg ccacatccgg tggcatactc aggaagaaag attggcggcc 540
agcattgcta tagcaagagc gctatcttct aacgttgttg tttctggcaa tggcgatgaa 600
gatgaaggtc catctaaaaa gtataaactc ccggacctga acaagtcacc accgccggag 660
gaggaggacg aggacgctgc ctga 684
<210> 14
<211> 3925
<212> DNA
<213> Taraxacum officinale
<400> 14
ttttcttgcg ggagaggcat gtatgtgtgg cctttttgca ataatactat gcatgggaga 60
tgcataaagt ttatctttat tttgttttag tttaataatg tgtggcaaac tttatatcat 120
ttatgatttc ggccatgtac tgctacatcg ttctaggatg gttattccct agaatttgct 180
tttatatatt ttttttcttc gttaattctt ctcttccttt aacgttattt catgtgaatt 240
tgcgatatca tgatattgca ggaacgtaac ttagaactca acgataaaaa gtgtgttgag 300
atcgtggaag ttgatcgttt atcttgaatt cacaagataa gcaacatgag atagaataat 360
taattactgc ggagatttct ttttagattc atcttattcg ttgtaaattg caatgcaaat 420
ttttaagtag aataacgaaa gcgtgttgtt atttacggtt aaatgcaaaa aatcaataac 480
atatctccat aattgatttg tttagtatgc attctcatgt gttctattgc aaaattctat 540
tttgaaattc aattaaagca atgcatttta gatataaagc aattccatca cttaattata 600
ttgggtagat ttttaccaaa tcactgtcct aaaagaaaaa taaaaaatga aagttcaatg 660
ccatagtgat cccgtcaatt atgtgatcaa tttttaccag agcaaaaccc caaaagttaa 720
agagtataat tgcagaacga caacaatctg aggatgatac taactccggt ataaaccgaa 780
tgcttttgtg gggatttcac taatcaacta tttggctctc atcacaaggc cgtgttaatg 840
gattactatt agttataagg ccatcccaag ttgtttggct atattattac cttttaaaaa 900
ccaaaatgtg taaactgtaa aattacactt ttggttcttg tattttctaa tttttagaat 960
tatgtatttg gttcttgtag tttttttatt tgtatgaatc atattagttt tggtccacga 1020
taacttttat tttctataat tacacttttg gtcattgtag ttttaatttg caagaaaaaa 1080
ataatctatc tttttagtcc atatatatag tttgcaatat tctatatttt aggttcaggt 1140
tatatatctt ttacaaaaat actacttatg attttttaat gaatatatca tatttcgtaa 1200
attacataaa cattttatat aattttattt agtcagaact ataggattcg catgataact 1260
ttgctgaact taattgggct tcatttttaa caaaggataa tgcatcatgc atgacataaa 1320
aatttgtaat gtttctaatg tgcatgcact acaagaacct gacccttcat atacattaat 1380
gaatgattct cttttcttat tatatcttag caaacggttt ggataagagt ctgaacaata 1440
ttaagaattg cactagctag ctaggggagt ccatgcagca agcaaattaa ttagaggact 1500
ttttaagagg cttgatcact tcattgtata tctaatggca ctgtggatat atgattggat 1560
tgtgtgtatg tatggaattt aatattacta aaaaatatac atttaatgac aaacaataac 1620
acactaattt ataacttaca aaagcgtgta tcattaaaga aaatttcatg tttcttaaaa 1680
tatggagaat ttactactta cataattatg tgtcataaat ttatgtcatc ttgtgtgata 1740
tttatgactt acaaaaatgt atgtgatatt tatgacttac aaaaatgtat gttatgttta 1800
tcgtcataaa atctgtaaaa aaaaatagaa acatggagga tactttcaat gtgtagaaac 1860
ttgacagcat aaacaggttt attaccttca tcaaaactta tcttacttca atctcttgga 1920
gatgcaggtt gattttaatg ctaaatgaac ttttcaaatg tataaatata tatcttttga 1980
cggatttgac attgatggag atgaagatac accagtgaat tcacctttca atgatcatac 2040
tgacatacca agtactctta tcccctctcc acctccacag aacgaacctc aatttcaaaa 2100
acgtaactgc cgctgttcta aaaacagaaa attttcaaaa acgtaactat catttaaata 2160
tttaatcatt tttcaattgg attcatgctg taatagaagt tactggactt taaacttctt 2220
gtagctgatg agattcatga acaacagtcc aaggagattt tccaaatcaa caagtttctg 2280
atgagaacaa ggtaaaaaaa ataagttttt catttaaaca tatgattttg gttttcaaaa 2340
gataattaat tataggaact cttctgaaaa gtcccaaagt ttacccgtaa aattgatttt 2400
ggtttttttt ttttaaattg aataaaaaag tccataaaat tggataaatt attcgattta 2460
gcccaattta cctagtatat ctggtaaaaa tgggttaaat cgaataattt aggcagtttg 2520
ttgggctttt ttgcgcaaaa acattatttt gggacttaat cgttaatttt ccccaagttt 2580
cagtttattt acagtattgg aataaaaccg aacctagtgc gctaattttg tcttttggtg 2640
tttagaatta tttgggttta ttggtctcgg tttctaaacg agttcggttt ttggtttctt 2700
gattcgattt ttggcttttg ggttcaaccc atggattttc gatttggttt tggttttgat 2760
aatatttttt gtgcaatcgt tcaaaaaatc cgagttagat tgggtttaaa tgttatcaaa 2820
accaaaaccg atggtagaac cccaaaactg aacccaatcg aaaatccatg ggttgaacct 2880
caaaaacgaa cccaataaat tctaaaaacc aaaatccaaa attagcggat aaaaatttta 2940
aaatacatat ctactggtaa ggtgtttttg acaatttatg taaattatag aaaaaaatgt 3000
attaagtgtt tgctatttaa gataaaaatg gcgttaaaaa tacaattggt tagaaatgaa 3060
ccattttttt gttgtaaatg agtggtgcat caacgatggt ggatgccatc ttttccttcc 3120
accatataca tctttgtcaa acaccataca cacaaattct catatgtcaa agatattaat 3180
tatcatgtcc actgagtttg taatataaat agatgatcaa ttacaacgag aaatatacac 3240
tgaattctca tttctaaatt gaagaatctc tctctctctc tctcatctct cagttttcaa 3300
agttttataa gttactagtt tatagtgttt ctaattactt catatttata acacgttatc 3360
agcacgagag ctctaagcaa gatatgcaat tacttaataa agtttataaa aataaacaaa 3420
tcataataaa caacatagat ataaaattaa ttattaccat ggacatgatg gactactaac 3480
attttatttc ttatactaac atttatttat gttaactaac atttatttat gtttctaaca 3540
tttatttatg catactaaca tttatttatg ttaactaaca tttatttatg tttctaacat 3600
ttatttatgc atactaacat ttatttatgt tactaacttt tatttatgta tactaatgtt 3660
tatttatcaa atttatttaa catttactat tttttgatta tattttttaa ttagccgata 3720
gaatttattt atttaattat tctatttatt aaacttattt ttataaatat atatatatat 3780
ataaccgtta tatatataac ggctatatat ataacgtcta tatatatata acggctatat 3840
ttttacacta taaataaaca catcattctc catttttata ccacaacaaa taatttctct 3900
catcttccaa atcatcaaga gaaaa 3925
<210> 15
<211> 7826
<212> DNA
<213> Taraxacum officinale
<400> 15
gttgaaagga aggatcggca gagaacaaga ttgagagagt tttatcgaag agaagagatg 60
gttgtgttga ttgacggcag agttcagcaa tggcagatat gcgtgtttaa agcgcagttt 120
gtggacagtt tgtccaaagg ccttcaagtg ggagattgtt gagatgaaga cccttttgac 180
caagtactta aatgtgattg gttgcatggg aaagagcttt acacttcatt ggttgaaaat 240
gaagcatgtt atttaatagt acgtctccac ttgttcttta tgaaggatga agacagccac 300
caagtcttta tgaatgtttc aaggagaaga caatgattag atgctgcctg ctcccttact 360
tcttcactat aaataggtgc ttaaggcacc tctcttctta caccaaacaa cacacaagtc 420
acagattaca aagattgaga gagttttcat tttctctcta gagtttgttt tgtcaaaagt 480
tttgtgtgca gattaattag ttgagagaca ctcctaatta atataaccac ttacacttgt 540
cattgattag cattctagtg tcagagagtt ctgagtgatt gtaacctttg tgtgatttat 600
atataaatct ctctcttgtt cgccagtgga cgtagctgat tatcctgatc agtgaaccac 660
ttaaaatatc gtgttgatat tatttatctg cagattattt taccagatag ttttattccg 720
caccgcaaag cataccgtcc attatttgtc agacgtacct actctcagct catcccagca 780
ggagttagac ctgcaggcag ccagccgtat ttcacaacag tgaattatgc aacaaatgga 840
aacacacaat tatcgtagta cacatgtaat taacagggat caatttccat ttattgttcc 900
atcaattgcc aaatttaaaa gataacaaat gcatcctttt tgaatggaaa agtatgatac 960
ataaaaccag agagaggtat acatttagtc cagaaacggt ggaaattact ttacctagag 1020
ttgcgtgaat caaatggaaa atcagaggaa gagaccctaa atcgcaatcc atcttctatt 1080
ccctatagaa tcaaatggaa atttcttact tccgaccttt ttcatttcca caagtttgta 1140
atcgataaga tcataaaaga aaaccgattt acattagcaa atctttaagg cagtttgtag 1200
ttctcaagta tgcgttacgt atgaaattct agacagaaaa gggacgacag ggaaaataat 1260
ggtggaataa ataaacaatc aagaatcaaa agataataat ccttaaaact aacaaaaatt 1320
ctatcaataa gccctacaaa tgagaaatat gacggtggag actcacctga ttgatgcgag 1380
aaaaggaaag ccgccgacat ttgtgcttct tcccgcgtta agaaaaatcg cagatgacat 1440
tacgattctt tttctacaga ggaatagggg ggttgcgact ttgcgtgatt tgttgttaat 1500
gagtgatgcg tcgtccgttt tgatcggtag cccatggaga ctgatttatt tgacagatag 1560
atgtcgttct cgttgatgat gcaggtgtaa ttggcggcgt tggtgaagcg aagatggcat 1620
agggaatgga tttgcagagg agaaaaggag tttcacaaat atggattatt gaattgtcgt 1680
ttcagagtcg cgatggttag taatcgatgg atttgggttt gcacctgttc aaggggtggt 1740
gacgtgtctt actgtaagaa agagtcgcga tggttagtaa tcgacgatgg atttgggttt 1800
gcaacatcgt tgacggaaac gacgcggcga tggagagagg cggtagaggg aatgaaacgg 1860
gagatttacc ttcaaccgca aacatgtctt cgttcctcga aaatcatatg cgccctttat 1920
gaaaattcct tcacgcgccg tacacgcgct cagaaaaagc atagggtttt cttatataat 1980
aatgatataa cttaatttta aataacatcg tactatacgt ttcaaaataa attagagttt 2040
taataaaatt acggaatgat taaacctatt acaaacaaat ataagtgtcg gtaatataaa 2100
atttgtaaca acgaaaacat ttcaaaaaat ctgaatgcgg aaatccataa catgagaata 2160
cgattcgttt gcagtcacgc cggtttcttc accttctgag ggtcctgaaa aaggttaata 2220
aacaatgggt aagctaaagt ttagggatca tacacattac ttgtctatta tatgagttag 2280
atacattcac atatcacata catacaagca tacacatatt acaaaaaatc ctaacacata 2340
tggttggtac caactactat ttagtggacc aggttattct tgggggtcaa ttgtggttaa 2400
ccataaatgt ttggagttta ggttctcttt gggggccgac cataaacacc catatcatat 2460
ctctgtctag gttctctttg gggactgact atagacaact ttgctcagat tctccttgga 2520
ggataaccat cctcgacgag attaacaacg gaccaccact gttttcagct tgtaacctgc 2580
aacatcaata gttgcgatct tcggcgacaa tgatgaacca acatcaacac aacagtagtg 2640
ccggtatgtt agtcgttgga taatagaggc atcatctgac ccttttagcg gtgacttttg 2700
ttgcctatga aggtcttatc agttatatat cgattaggct tggttctagg ttataagaaa 2760
aaatgattct ctatccggta tgacatagag acaaaccggt ctacttttat aagaaaatca 2820
tactcgtatt tgaaacgaaa atcatatttg aaacaatgtc gtgttaataa ttaatatgtg 2880
tatgtttaaa tttagaatat cctgaactta tatttagttc gcaaccctat gtaagatcga 2940
actaatttta aaccgtaacc agacatgcat gtttgatcag taataaaaag gctatcatcg 3000
tcgaaaatat taaaacaaaa tataatttct tatttggcgc catttcatgt caacggttaa 3060
aagcaagtta tcaacgcctc caaccgatca tttctccggt aaccaccaac tgctactaat 3120
aaccgccacc gcaaaaacct atatatacaa cttccatcac aattctgtcc cctactccca 3180
tttcaatcca aaaaaggcta cacagacata tacagcgatg tcaacggtta aaagcaagtt 3240
atcaacgcct ccaaccgatc atttctccgg taaccaccaa ctgctactaa taaccgccac 3300
cgcaaaaacc tatatataca acttccatca caattctgtc ccctactccc atttcaatcc 3360
aaaaaaggct acacagacat atacagcgat ggcagataat ggcaacaccg gccgtcaaaa 3420
ggatgacgac ggtggccatg atgggggacg cccaaatccg accaccccac cctccccttc 3480
ccgcaccccg cgaagaccca ggcggaccac atcgccgcct aaacattctc cgggggcgtc 3540
ttcaagcacc atgccagcgc cgcctactcc ccctgcgccg acgggaatca ccggtgcgtc 3600
tagttcttct ctgggtacta atataatttc atttactcca cccaaaacca aaagaacgaa 3660
gtcggtgatt tgtccgatct gtaagaaaga tatgtgccat gagaaggcgc tgtgtggcca 3720
catccggtgg catactcagg aagaaagatt ggcggccagc attgctatag caagagcgct 3780
atcttctaac gttgttgttt ctggcaatgg cgatgaagat gaaggtccat ctaaaaagta 3840
taaactcccg gacctgaaca agtcaccacc gccggaggag gaggacgagg acgctgcctg 3900
attttcttgc gggagaggca tgtatgtgtg gcctttttgc aataatacta tgcatgggag 3960
atgcataaag tttatcttta ttttgtttta gtttaataat gtgtggcaaa ctttatatca 4020
tttatgattt cggccatgta ctgctacatc gttctaggat ggttattccc tagaatttgc 4080
ttttatatat tttttttctt cgttaattct tctcttcctt taacgttatt tcatgtgaat 4140
ttgcgatatc atgatattgc aggaacgtaa cttagaactc aacgataaaa agtgtgttga 4200
gatcgtggaa gttgatcgtt tatcttgaat tcacaagata agcaacatga gatagaataa 4260
ttaattactg cggagatttc tttttagatt catcttattc gttgtaaatt gcaatgcaaa 4320
tttttaagta gaataacgaa agcgtgttgt tatttacggt taaatgcaaa aaatcaataa 4380
catatctcca taattgattt gtttagtatg cattctcatg tgttctattg caaaattcta 4440
ttttgaaatt caattaaagc aatgcatttt agatataaag caattccatc acttaattat 4500
attgggtaga tttttaccaa atcactgtcc taaaagaaaa ataaaaaatg aaagttcaat 4560
gccatagtga tcccgtcaat tatgtgatca atttttacca gagcaaaacc ccaaaagtta 4620
aagagtataa ttgcagaacg acaacaatct gaggatgata ctaactccgg tataaaccga 4680
atgcttttgt ggggatttca ctaatcaact atttggctct catcacaagg ccgtgttaat 4740
ggattactat tagttataag gccatcccaa gttgtttggc tatattatta ccttttaaaa 4800
accaaaatgt gtaaactgta aaattacact tttggttctt gtattttcta atttttagaa 4860
ttatgtattt ggttcttgta gtttttttat ttgtatgaat catattagtt ttggtccacg 4920
ataactttta ttttctataa ttacactttt ggtcattgta gttttaattt gcaagaaaaa 4980
aataatctat ctttttagtc catatatata gtttgcaata ttctatattt taggttcagg 5040
ttatatatct tttacaaaaa tactacttat gattttttaa tgaatatatc atatttcgta 5100
aattacataa acattttata taattttatt tagtcagaac tataggattc gcatgataac 5160
tttgctgaac ttaattgggc ttcattttta acaaaggata atgcatcatg catgacataa 5220
aaatttgtaa tgtttctaat gtgcatgcac tacaagaacc tgacccttca tatacattaa 5280
tgaatgattc tcttttctta ttatatctta gcaaacggtt tggataagag tctgaacaat 5340
attaagaatt gcactagcta gctaggggag tccatgcagc aagcaaatta attagaggac 5400
tttttaagag gcttgatcac ttcattgtat atctaatggc actgtggata tatgattgga 5460
ttgtgtgtat gtatggaatt taatattact aaaaaatata catttaatga caaacaataa 5520
cacactaatt tataacttac aaaagcgtgt atcattaaag aaaatttcat gtttcttaaa 5580
atatggagaa tttactactt acataattat gtgtcataaa tttatgtcat cttgtgtgat 5640
atttatgact tacaaaaatg tatgtgatat ttatgactta caaaaatgta tgttatgttt 5700
atcgtcataa aatctgtaaa aaaaaataga aacatggagg atactttcaa tgtgtagaaa 5760
cttgacagca taaacaggtt tattaccttc atcaaaactt atcttacttc aatctcttgg 5820
agatgcaggt tgattttaat gctaaatgaa cttttcaaat gtataaatat atatcttttg 5880
acggatttga cattgatgga gatgaagata caccagtgaa ttcacctttc aatgatcata 5940
ctgacatacc aagtactctt atcccctctc cacctccaca gaacgaacct caatttcaaa 6000
aacgtaactg ccgctgttct aaaaacagaa aattttcaaa aacgtaacta tcatttaaat 6060
atttaatcat ttttcaattg gattcatgct gtaatagaag ttactggact ttaaacttct 6120
tgtagctgat gagattcatg aacaacagtc caaggagatt ttccaaatca acaagtttct 6180
gatgagaaca aggtaaaaaa aataagtttt tcatttaaac atatgatttt ggttttcaaa 6240
agataattaa ttataggaac tcttctgaaa agtcccaaag tttacccgta aaattgattt 6300
tggttttttt tttttaaatt gaataaaaaa gtccataaaa ttggataaat tattcgattt 6360
agcccaattt acctagtata tctggtaaaa atgggttaaa tcgaataatt taggcagttt 6420
gttgggcttt tttgcgcaaa aacattattt tgggacttaa tcgttaattt tccccaagtt 6480
tcagtttatt tacagtattg gaataaaacc gaacctagtg cgctaatttt gtcttttggt 6540
gtttagaatt atttgggttt attggtctcg gtttctaaac gagttcggtt tttggtttct 6600
tgattcgatt tttggctttt gggttcaacc catggatttt cgatttggtt ttggttttga 6660
taatattttt tgtgcaatcg ttcaaaaaat ccgagttaga ttgggtttaa atgttatcaa 6720
aaccaaaacc gatggtagaa ccccaaaact gaacccaatc gaaaatccat gggttgaacc 6780
tcaaaaacga acccaataaa ttctaaaaac caaaatccaa aattagcgga taaaaatttt 6840
aaaatacata tctactggta aggtgttttt gacaatttat gtaaattata gaaaaaaatg 6900
tattaagtgt ttgctattta agataaaaat ggcgttaaaa atacaattgg ttagaaatga 6960
accatttttt tgttgtaaat gagtggtgca tcaacgatgg tggatgccat cttttccttc 7020
caccatatac atctttgtca aacaccatac acacaaattc tcatatgtca aagatattaa 7080
ttatcatgtc cactgagttt gtaatataaa tagatgatca attacaacga gaaatataca 7140
ctgaattctc atttctaaat tgaagaatct ctctctctct ctctcatctc tcagttttca 7200
aagttttata agttactagt ttatagtgtt tctaattact tcatatttat aacacgttat 7260
cagcacgaga gctctaagca agatatgcaa ttacttaata aagtttataa aaataaacaa 7320
atcataataa acaacataga tataaaatta attattacca tggacatgat ggactactaa 7380
cattttattt cttatactaa catttattta tgttaactaa catttattta tgtttctaac 7440
atttatttat gcatactaac atttatttat gttaactaac atttatttat gtttctaaca 7500
tttatttatg catactaaca tttatttatg ttactaactt ttatttatgt atactaatgt 7560
ttatttatca aatttattta acatttacta ttttttgatt atatttttta attagccgat 7620
agaatttatt tatttaatta ttctatttat taaacttatt tttataaata tatatatata 7680
tataaccgtt atatatataa cggctatata tataacgtct atatatatat aacggctata 7740
tttttacact ataaataaac acatcattct ccatttttat accacaacaa ataatttctc 7800
tcatcttcca aatcatcaag agaaaa 7826
<210> 16
<211> 972
<212> DNA
<213> Artificial Sequence
<220>
<223> parsley ubiquitin promoter sequence
<400> 16
aaaaattacg gatatgaata taggcatatc cgtatccgaa ttatccgttt gacagctagc 60
aacgattgta caattgcttc tttaaaaaag gaagaaagaa agaaagaaaa gaatcaacat 120
cagcgttaac aaacggcccc gttacggccc aaacggtcat atagagtaac ggcgttaagc 180
gttgaaagac tcctatcgaa atacgtaacc gcaaacgtgt catagtcaga tcccctcttc 240
cttcaccgcc tcaaacacaa aaataatctt ctacagccta tatatacaac ccccccttct 300
atctctcctt tctcacaatt catcatcttt ctttctctac ccccaatttt aagaaatcct 360
ctcttctcct cttcattttc aaggtaaatc tctctctctc tctctctctc tgttattcct 420
tgttttaatt aggtatgtat tattgctagt ttgttaatct gcttatctta tgtatgcctt 480
atgtgaatat ctttatcttg ttcatctcat ccgtttagaa gctataaatt tgttgatttg 540
actgtgtatc tacacgtggt tatgtttata tctaatcaga tatgaatttc ttcatattgt 600
tgcgtttgtg tgtaccaatc cgaaatcgtt gatttttttc atttaatcgt gtagctaatt 660
gtacgtatac atatggatct acgtatcaat tgttcatctg tttgtgtttg tatgtataca 720
gatctgaaaa catcacttct ctcatctgat tgtgttgtta catacataga tatagatctg 780
ttatatcatt ttttttatta attgtgtata tatatatgtg catagatctg gattacatga 840
ttgtgattat ttacatgatt ttgttattta cgtatgtata tatgtagatc tggacttttt 900
ggagttgttg acttgattgt atttgtgtgt gtatatgtgt gttctgatct tgatatgtta 960
tgtatgtgca gc 972
<210> 17
<211> 4140
<212> DNA
<213> Artificial Sequence
<220>
<223> Cas9
<400> 17
atggataaga agtactctat cggactcgat atcggaacta actctgtggg atgggctgtg 60
atcaccgatg agtacaaggt gccatctaag aagttcaagg ttctcggaaa caccgatagg 120
cactctatca agaaaaacct tatcggtgct ctcctcttcg attctggtga aactgctgag 180
gctaccagac tcaagagaac cgctagaaga aggtacacca gaagaaagaa caggatctgc 240
tacctccaag agatcttctc taacgagatg gctaaagtgg atgattcatt cttccacagg 300
ctcgaagagt cattcctcgt ggaagaagat aagaagcacg agaggcaccc tatcttcgga 360
aacatcgttg atgaggtggc ataccacgag aagtacccta ctatctacca cctcagaaag 420
aagctcgttg attctactga taaggctgat ctcaggctca tctacctcgc tctcgctcac 480
atgatcaagt tcagaggaca cttcctcatc gagggtgatc tcaaccctga taactctgat 540
gtggataagt tgttcatcca gctcgtgcag acctacaacc agcttttcga agagaaccct 600
atcaacgctt caggtgtgga tgctaaggct atcctctctg ctaggctctc taagtcaaga 660
aggcttgaga acctcattgc tcagctccct ggtgagaaga agaacggact tttcggaaac 720
ttgatcgctc tctctctcgg actcacccct aacttcaagt ctaacttcga tctcgctgag 780
gatgcaaagc tccagctctc aaaggatacc tacgatgatg atctcgataa cctcctcgct 840
cagatcggag atcagtacgc tgatttgttc ctcgctgcta agaacctctc tgatgctatc 900
ctcctcagtg atatcctcag agtgaacacc gagatcacca aggctccact ctcagcttct 960
atgatcaaga gatacgatga gcaccaccag gatctcacac ttctcaaggc tcttgttaga 1020
cagcagctcc cagagaagta caaagagatt ttcttcgatc agtctaagaa cggatacgct 1080
ggttacatcg atggtggtgc atctcaagaa gagttctaca agttcatcaa gcctatcctc 1140
gagaagatgg atggaaccga ggaactcctc gtgaagctca atagagagga tcttctcaga 1200
aagcagagga ccttcgataa cggatctatc cctcatcaga tccacctcgg agagttgcac 1260
gctatcctta gaaggcaaga ggatttctac ccattcctca aggataacag ggaaaagatt 1320
gagaagattc tcaccttcag aatcccttac tacgtgggac ctctcgctag aggaaactca 1380
agattcgctt ggatgaccag aaagtctgag gaaaccatca ccccttggaa cttcgaagag 1440
gtggtggata agggtgctag tgctcagtct ttcatcgaga ggatgaccaa cttcgataag 1500
aaccttccaa acgagaaggt gctccctaag cactctttgc tctacgagta cttcaccgtg 1560
tacaacgagt tgaccaaggt taagtacgtg accgagggaa tgaggaagcc tgcttttttg 1620
tcaggtgagc aaaagaaggc tatcgttgat ctcttgttca agaccaacag aaaggtgacc 1680
gtgaagcagc tcaaagagga ttacttcaag aaaatcgagt gcttcgattc agttgagatt 1740
tctggtgttg aggataggtt caacgcatct ctcggaacct accacgatct cctcaagatc 1800
attaaggata aggatttctt ggataacgag gaaaacgagg atatcttgga ggatatcgtt 1860
cttaccctca ccctctttga agatagagag atgattgaag aaaggctcaa gacctacgct 1920
catctcttcg atgataaggt gatgaagcag ttgaagagaa gaagatacac tggttgggga 1980
aggctctcaa gaaagctcat taacggaatc agggataagc agtctggaaa gacaatcctt 2040
gatttcctca agtctgatgg attcgctaac agaaacttca tgcagctcat ccacgatgat 2100
tctctcacct ttaaagagga tatccagaag gctcaggttt caggacaggg tgatagtctc 2160
catgagcata tcgctaacct cgctggatct cctgcaatca agaagggaat cctccagact 2220
gtgaaggttg tggatgagtt ggtgaaggtg atgggaaggc ataagcctga gaacatcgtg 2280
atcgaaatgg ctagagagaa ccagaccact cagaagggac agaagaactc tagggaaagg 2340
atgaagagga tcgaggaagg tatcaaagag cttggatctc agatcctcaa agagcaccct 2400
gttgagaaca ctcagctcca gaatgagaag ctctacctct actacctcca gaacggaagg 2460
gatatgtatg tggatcaaga gttggatatc aacaggctct ctgattacga tgttgatcat 2520
atcgtgccac agtcattctt gaaggatgat tctatcgata acaaggtgct caccaggtct 2580
gataagaaca ggggtaagag tgataacgtg ccaagtgaag aggttgtgaa gaaaatgaag 2640
aactattgga ggcagctcct caacgctaag ctcatcactc agagaaagtt cgataacttg 2700
actaaggctg agaggggagg actctctgaa ttggataagg caggattcat caagaggcag 2760
cttgtggaaa ccaggcagat cactaagcac gttgcacaga tcctcgattc taggatgaac 2820
accaagtacg atgagaacga taagttgatc agggaagtga aggttatcac cctcaagtca 2880
aagctcgtgt ctgatttcag aaaggatttc caattctaca aggtgaggga aatcaacaac 2940
taccaccacg ctcacgatgc ttaccttaac gctgttgttg gaaccgctct catcaagaag 3000
tatcctaagc tcgagtcaga gttcgtgtac ggtgattaca aggtgtacga tgtgaggaag 3060
atgatcgcta agtctgagca agagatcgga aaggctaccg ctaagtattt cttctactct 3120
aacatcatga atttcttcaa gaccgagatt accctcgcta acggtgagat cagaaagagg 3180
ccactcatcg agacaaacgg tgaaacaggt gagatcgtgt gggataaggg aagggatttc 3240
gctaccgtta gaaaggtgct ctctatgcca caggtgaaca tcgttaagaa aaccgaggtg 3300
cagaccggtg gattctctaa agagtctatc ctccctaaga ggaactctga taagctcatt 3360
gctaggaaga aggattggga ccctaagaaa tacggtggtt tcgattctcc taccgtggct 3420
tactctgttc tcgttgtggc taaggttgag aagggaaaga gtaagaagct caagtctgtt 3480
aaggaacttc tcggaatcac tatcatggaa aggtcatctt tcgagaagaa cccaatcgat 3540
ttcctcgagg ctaagggata caaagaggtt aagaaggatc tcatcatcaa gctcccaaag 3600
tactcactct tcgaactcga gaacggtaga aagaggatgc tcgcttctgc tggtgagctt 3660
caaaagggaa acgagcttgc tctcccatct aagtacgtta actttcttta cctcgcttct 3720
cactacgaga agttgaaggg atctccagaa gataacgagc agaagcaact tttcgttgag 3780
cagcacaagc actacttgga tgagatcatc gagcagatct ctgagttctc taaaagggtg 3840
atcctcgctg atgcaaacct cgataaggtg ttgtctgctt acaacaagca cagagataag 3900
cctatcaggg aacaggcaga gaacatcatc catctcttca cccttaccaa cctcggtgct 3960
cctgctgctt tcaagtactt cgatacaacc atcgatagga agagatacac ctctaccaaa 4020
gaagtgctcg atgctaccct catccatcag tctatcactg gactctacga gactaggatc 4080
gatctctcac agctcggtgg tgattcaagg gctgatccta agaagaagag gaaggtttga 4140
<210> 18
<211> 80
<212> DNA
<213> Artificial Sequence
<220>
<223> tomato U6 promoter
<400> 18
ggagtgatca aaagtcccac atcgatcagg tgatatatag cagcttagtt tatataatga 60
tagagtcgac atagcgattg 80
<210> 19
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> target specific sequence of RNA-1
<400> 19
catccggtgg catacacagg 20
<210> 20
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> target specific sequence of RNA-2
<400> 20
ggagtagttg ggttttggcg 20
<210> 21
<211> 89
<212> PRT
<213> Helianthus annuus
<400> 21
Met Ser Ser Pro Ser Lys Asn Lys Ser Lys Ser Ser Ile Cys Pro Val
1 5 10 15
Cys Lys Arg Asp Leu Cys His Glu Lys Ala Leu Asn Gly His Ile Arg
20 25 30
Trp His Thr Gln Gln Glu Arg Glu Ala Ala Gly Ile Gly Asn Ala Lys
35 40 45
Ala Val Ala Ser Ala Ser Val Val Ile Arg Asp Ala Asn Val Arg Arg
50 55 60
Pro Glu Thr Ser Lys Pro Val Lys Leu Pro Asp Leu Asn Lys Ser Pro
65 70 75 80
Pro Arg Glu Glu Asp Lys Asp Ala Ala
85
<210> 22
<211> 157
<212> PRT
<213> Lactuca sativa
<400> 22
Met Ala Asp Asp Gly Asn Thr Ala Arg Gln Gln Ala Asp Ala Gly Gly
1 5 10 15
His Gly Asn Ala Arg Pro Asn Ser Ser Thr Pro Pro Ser Ser Pro Ser
20 25 30
Gln Pro Pro Arg Arg Pro Arg Arg Ala Gly Ala Thr Thr Pro Ser Lys
35 40 45
Leu Ser Gln Ala Ala Ser Ser Ser Thr Asn Leu Pro Pro Pro Pro Thr
50 55 60
Pro Thr Pro Thr Pro Pro Thr Pro Ser Ala Asp Gly Ile Leu Leu Gly
65 70 75 80
Thr Ala Arg Arg Pro Val Ile Cys Pro Ile Cys Lys Lys Asp Met Tyr
85 90 95
His Glu Lys Ala Leu Cys Gly His Ile Arg Trp His Thr Gln Glu Glu
100 105 110
Arg Leu Ala Ala Ser Arg Asp Ile Ala Arg Ala Leu Ser Ala Asn Val
115 120 125
Val Ser Gly Gln Arg Gly Asp Gly Glu Gln Gly Pro Ser Lys Arg Phe
130 135 140
Lys Leu Pro Asp Leu Asn Glu Pro Pro Pro Ser Glu Asp
145 150 155
<210> 23
<211> 36
<212> DNA
<213> Taraxacum officinale
<400> 23
tgtggccaca tccggtggca tacacaggag gaaaga 36
<210> 24
<211> 37
<212> DNA
<213> Artificial Sequence
<220>
<223> pKG10821-1
<400> 24
tgtggccaca tccggtggca tacaccagga ggaaaga 37
<210> 25
<211> 37
<212> DNA
<213> Artificial Sequence
<220>
<223> pKG10821-4
<400> 25
tgtggccaca tccggtggca tacacaagga ggaaaga 37
<210> 26
<211> 35
<212> DNA
<213> Artificial Sequence
<220>
<223> pKG10821-5
<400> 26
tgtggccaca tccggtggca tacaaggagg aaaga 35
<210> 27
<211> 33
<212> DNA
<213> Artificial Sequence
<220>
<223> pKG10821-6
<400> 27
tgtggccaca tccggtggca taaggaggaa aga 33
<210> 28
<211> 12
<212> PRT
<213> Artificial Sequence
<220>
<223> pKG10821-1
<400> 28
Cys Gly His Ile Arg Trp His Thr Pro Gly Gly Lys
1 5 10
<210> 29
<211> 12
<212> PRT
<213> Artificial Sequence
<220>
<223> pKG108121-4
<400> 29
Cys Gly His Ile Arg Trp His Thr Gln Gly Gly Lys
1 5 10
<210> 30
<211> 11
<212> PRT
<213> Artificial Sequence
<220>
<223> pKG10821-5
<400> 30
Cys Gly His Ile Arg Trp His Thr Arg Arg Lys
1 5 10
<210> 31
<211> 11
<212> PRT
<213> Artificial Sequence
<220>
<223> pKG10821-6
<400> 31
Cys Gly His Ile Arg Trp His Lys Glu Glu Arg
1 5 10
<210> 32
<211> 12
<212> PRT
<213> Taraxacum officinale
<400> 32
Cys Gly His Ile Arg Trp His Thr Gln Glu Glu Arg
1 5 10
<210> 33
<211> 22
<212> DNA
<213> Artificial Sequence
<220>
<223> forward primer
<400> 33
gaaaccgaag caaactctac ca 22
<210> 34
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> reversed primer
<400> 34
gcgctttcta caatcttaca 20
<210> 35
<211> 20
<212> DNA
<213> Artificial Sequence
<220>
<223> forward primer
<400> 35
gatcgaacta attttaaacc 20
<210> 36
<211> 22
<212> DNA
<213> Artificial Sequence
<220>
<223> reversed primer
<400> 36
gatcaacttc cacgatctca ac 22
<210> 37
<211> 21
<212> PRT
<213> Artificial Sequence
<220>
<223> K2-2 like zinc finger domain
<220>
<221> misc_feature
<222> (2)..(3)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> misc_feature
<222> (5)..(11)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> MISC_FEATURE
<222> (12)..(12)
<223> Xaa can be K or R
<220>
<221> misc_feature
<222> (14)..(15)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> misc_feature
<222> (18)..(18)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> MISC_FEATURE
<222> (19)..(19)
<223> Xaa can be R or N
<220>
<221> misc_feature
<222> (20)..(20)
<223> Xaa can be any naturally occurring amino acid
<400> 37
Cys Xaa Xaa Cys Xaa Xaa Xaa Xaa Xaa Xaa Xaa Xaa Ala Xaa Xaa Gly
1 5 10 15
His Xaa Xaa Xaa His
20
<210> 38
<211> 21
<212> PRT
<213> Artificial Sequence
<220>
<223> consensus zinc finger domain
<220>
<221> misc_feature
<222> (2)..(3)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> misc_feature
<222> (5)..(16)
<223> Xaa can be any naturally occurring amino acid
<220>
<221> misc_feature
<222> (18)..(20)
<223> Xaa can be any naturally occurring amino acid
<400> 38
Cys Xaa Xaa Cys Xaa Xaa Xaa Xaa Xaa Xaa Xaa Xaa Xaa Xaa Xaa Xaa
1 5 10 15
His Xaa Xaa Xaa His
20
<210> 39
<211> 163
<212> PRT
<213> cichorium endivia
<400> 39
Met Val Asp Asp Gly Thr Thr Ala Arg Gln Gln Val Asp Asp Gly Gly
1 5 10 15
His Gly Asn Pro Arg Pro Asn Pro Thr Thr Pro Pro Pro Ser Pro Ser
20 25 30
Arg Thr Pro Arg Arg Thr Arg Arg Ala Gly Ala Thr Thr Pro Ser Lys
35 40 45
Leu Ser Pro Ala Val Ser Ser Thr Thr Leu Pro Pro Pro Pro Thr Pro
50 55 60
Pro Thr Pro Pro Ala Asp Arg Leu Val Thr Gly Thr Ser Ser Ser Leu
65 70 75 80
Gly Thr Gln Arg Val Arg Arg Ser Lys Ser Val Ile Cys Pro Ile Cys
85 90 95
Lys Lys Asp Met Cys His Glu Lys Ala Leu Cys Gly His Ile Arg Trp
100 105 110
His Thr His Glu Glu Arg Gln Ala Ala Ser Ser Asp Ile Ala Arg Ala
115 120 125
Leu Ser Ser Asn Phe Ser Ser Gly His Gly Gly Glu Glu Gln Gly Pro
130 135 140
Ser Lys Arg Phe Lys Val Pro Asp Leu Asn Lys Pro Pro Pro Pro Glu
145 150 155 160
Glu Asp Asp
<210> 40
<211> 161
<212> PRT
<213> Hieracium aurantiacum
<400> 40
Met Val Asp Asp Gly Thr Ala Ala Arg Gln Gln Val Asp Asp Gly Gly
1 5 10 15
His Gly Glu Arg Arg Pro Thr Pro Thr Thr Pro Pro Pro Ser Pro Ser
20 25 30
Arg Thr Pro Arg Arg Ser His Arg Pro Gly Thr Thr Pro Pro Ser Lys
35 40 45
Leu Ser Pro Ala Ala Ser Ser Thr Ala Leu Pro Pro Pro Pro Thr Pro
50 55 60
Pro Thr Pro Thr Val Val Ala Gly Ser Ser Ser Ser Leu Thr Thr Pro
65 70 75 80
Arg Ile Lys Arg Ser Lys Ser Val Ile Cys Pro Ile Cys Lys Lys Asp
85 90 95
Met Cys His Glu Lys Ala Leu Cys Gly His Ile Arg Trp His Thr Gln
100 105 110
Glu Glu Arg Leu Ala Ala Ser Gly Asp Ile Ala Arg Ala Leu Ser Ala
115 120 125
Asn Phe Val Ser Gly Gly Asn Gly Asp Glu Glu Gln Gly Ser Ser Lys
130 135 140
Arg Phe Lys Val Pro Asp Leu Asn Lys Pro Pro Pro Pro Glu Glu Asp
145 150 155 160
Asp
<210> 41
<211> 166
<212> PRT
<213> Senecio cambrensis
<400> 41
Met Val Asn Ser Gly Asp Ala Ser Arg Ser Asp Asp Asn Pro Asn Thr
1 5 10 15
Pro Asn Arg Arg Ala Gly Lys Ala His Ala Ala Pro Ser Thr Ser Pro
20 25 30
Ser Ser Pro Pro Pro Pro Ile Gly Arg Leu Ser Ala Thr Thr Pro Leu
35 40 45
Ser Gly Ser Lys Arg Pro His Ser Pro Thr Lys Lys Ser Asn Phe Ile
50 55 60
Cys Pro Ile Cys Gly Lys Asp Leu Tyr His Ser Lys Ala Leu Asn Gly
65 70 75 80
His Ile Arg Trp His Ser Gln Met Glu Arg Glu Leu Lys Lys Lys Ala
85 90 95
Leu Arg Asp Ile Arg Glu Arg Asp Val Ala Val Val Ala Ala Ala Asp
100 105 110
Ala Val Gln Val Ala Pro Val Pro Ala Thr Val Pro Val Pro Val Pro
115 120 125
Thr Ile Val Arg Thr Arg Leu Phe His Glu Gln Pro Gln Pro Gln Pro
130 135 140
Gln Glu Glu Val Pro Tyr Asn Tyr Ala Tyr Leu Leu Pro Asp Leu Asn
145 150 155 160
Ala Ser Ala Asp Glu Glu
165
<210> 42
<211> 203
<212> PRT
<213> Hevea brasiliensis
<400> 42
Met Val Asn Ser Ser Pro Ser Ser Ser Ser Ser Ser Ser Ser Pro Thr
1 5 10 15
Pro Pro Pro Glu Val Gln Val Gly Pro Gly Gly Ser Ser Val Gly Gly
20 25 30
Glu Ser Glu Glu Gly Cys Ser Ser Arg Lys Lys Lys Ala Gly Ser Asn
35 40 45
Asp Gln Gln Gly Asp Glu Glu Gly Val Gln Lys Arg Leu Lys Arg Gly
50 55 60
Glu Met Asp Ser Pro Ile Ser Glu Pro Val Cys Cys Ile Cys Asn Lys
65 70 75 80
Lys Phe Gly Ser Trp Lys Gly Val Phe Gly His Met Arg Ala His Pro
85 90 95
Glu Arg Glu Trp Arg Gly Ala Phe Pro Pro Pro Lys Glu Lys Ala Gly
100 105 110
Ala Ile Asn Ile Asn His Gln Gln Ile Leu Gln Gln Gln Leu Ala Pro
115 120 125
Thr Leu Leu Ser Leu Gly Arg Glu Ala Leu Ala Arg Met Thr Asn His
130 135 140
His Gly Ser Gly Ala Ala Asn Ala Ala Ala Pro Ser Arg Arg Ala Gly
145 150 155 160
Asp Thr Glu Leu Asn Arg Glu Gln Gln Glu Leu Ala Gly Pro Ser Thr
165 170 175
Ile Ile His Pro Ser Leu Arg Gly Phe Asp Leu Asn Leu Pro Pro Pro
180 185 190
Pro Glu Glu Ala Asn Glu Asp Asp Arg Asn Asn
195 200
<210> 43
<211> 230
<212> PRT
<213> Cucurbita moschata
<400> 43
Met Glu Lys Asn Ala Asn Thr Asn Val Pro Ser Glu Thr Ser Ser Asp
1 5 10 15
Gln Arg His Gly Gly Glu Arg Ser Pro Thr Gly Pro Thr Ser Pro Pro
20 25 30
Pro Thr His Asn Arg Asp Asn Asn Pro Asn Pro Thr Thr Glu His Ser
35 40 45
Ser Tyr Ser Asp Thr Leu Arg Val Pro Pro Ala Ala Glu Thr Ser Arg
50 55 60
Arg Asp Ala Leu Leu Asp Val Gly Thr Ser Tyr Gln Asn Ile Val Gly
65 70 75 80
Ser Ser Glu Gly Glu Pro Gly Lys Lys Arg Gly Arg Gly Asp Gly Glu
85 90 95
Glu Glu Glu Glu Gln Gln Gln Pro Pro Gln Gln Gln Val Lys Ala Ala
100 105 110
Lys Lys Lys Gly Glu Leu Thr Glu Val Pro Lys Gly Glu Pro Arg Cys
115 120 125
Ala Thr Cys Asn Lys Val Phe Lys Ser Trp Lys Ala Leu Phe Gly His
130 135 140
Leu Arg Ser His Pro Glu Arg Thr Tyr Arg Gly Ala Leu Pro Pro Pro
145 150 155 160
Thr Ala Ala Glu Leu Asp Ile Arg Arg Cys Gln Gln Gln Leu Ala Ser
165 170 175
Thr Leu Leu Thr Val Ala Gln Glu Val Ala Ala Ser Arg Arg Gly Leu
180 185 190
Asp Ile Asp Leu Asn Gln Pro Ser Thr Ala Asp Asp Gly Asp Ser Pro
195 200 205
Glu Lys Thr Gly Gly Val Gly Phe Asp Leu Asn Ala Asp Pro Pro Pro
210 215 220
Glu Ser Asp Asp Glu Asp
225 230
<210> 44
<211> 242
<212> PRT
<213> Eutrema salsugineum
<400> 44
Met Val Thr Glu Glu Glu Ile Asn Glu Ile Ser Glu Asp Ser Gly Tyr
1 5 10 15
Leu Ser Ser Gly Glu Glu Gly Glu Ile Arg Glu Leu Val Cys Ala Leu
20 25 30
Pro Ala Leu Asn Val Thr Glu Arg Leu Asn Met Glu Glu Ala Gly Leu
35 40 45
His Asn Ala Val Val Ala Ala Glu Phe Val Met Val Ala Ser Glu Glu
50 55 60
Ala Pro Val Ser Lys Glu Asn Met Gln Thr Val Asp Ala Ile Asp Asp
65 70 75 80
Asp Ala Lys Asn Thr Ala Leu Gly Arg Arg Pro Arg Ile Asp Glu Leu
85 90 95
Asp Asp Ala Ala Gly Ala Ser Gly Ser Thr Ala Ile Gly Ser Gly Gly
100 105 110
Glu Asp Glu Pro Val Lys Lys Ala Lys Lys Lys Gly Ser Ser Gln Leu
115 120 125
His Ala Pro Pro Gln Gly Pro Pro Gln Cys Asn Ile Cys Gly Arg Asn
130 135 140
Phe Thr Ser Trp Lys Ala Val Phe Gly His Leu Arg Ala His Lys Asp
145 150 155 160
Arg Gly Tyr Gln Gly Phe Leu Pro Pro Pro Thr Phe Asn Ala Thr Glu
165 170 175
Glu Gly Phe Ser Ala Val Val Leu Ala Ser Asp Ser Ser Arg Gly Gly
180 185 190
Val Leu Gly Phe Gly Ser Ser Gly Ile Asp Leu Asn Leu Asp Pro Met
195 200 205
Glu Glu Glu Glu Glu Ala Val Ser Gly Phe Ile Pro Lys Phe Asp Leu
210 215 220
Asn Arg Ser Pro Pro Gln Glu Gly Gln Glu Glu Glu Glu Glu Asp Lys
225 230 235 240
Ala Lys
<210> 45
<211> 244
<212> PRT
<213> Arachis ipaensis
<400> 45
Met Glu Lys Asn His Ser Thr Ser Ala Ser Thr Thr Ser Leu Pro Ser
1 5 10 15
Leu Ser Leu Ser Thr Pro Val Thr Glu Pro Pro Ser Ser Pro Leu Val
20 25 30
Asp Gln Pro Asn Leu Ser Gly Glu Glu Glu Lys Leu Asn Ile Leu Lys
35 40 45
Thr Pro Glu Lys Ile Leu Thr Asp Leu Thr Thr Asn Asn Asp Ala Asn
50 55 60
Gly Gly Ser Ala Leu Pro Pro Ser Ser Pro Pro Pro Ser Pro Pro Gln
65 70 75 80
Met Met Val Ser Gly Ala Ile Ala Thr Ala Val Gly Gly Gly Ala Val
85 90 95
Val Leu Gly Gly Asn Asn Ile Ile Gly Pro Ile Lys Arg Arg Lys Ala
100 105 110
Ser Glu Ile Met Asp Pro Pro Ser Gly Thr Pro Thr Cys Pro Ile Cys
115 120 125
Ser Lys Thr Phe Ser Ser Trp Lys Gly Ala Phe Gly His Met Arg Lys
130 135 140
His Pro Asp Arg Gln Tyr Arg Gly Phe Phe Lys Pro Pro Thr Phe Ala
145 150 155 160
Pro Ser Ser Ser Lys Ala Ala Thr Asn Ala Ser Gly Asn Arg His Glu
165 170 175
Ala Ala Gly Glu Glu Gly Ser Ser Ala Ala Ser Gly Glu Val Gly Glu
180 185 190
Gly Gly Ala Ser Pro Ala Val Arg Asn Leu Leu Phe Asp Leu Asn His
195 200 205
Pro Val Glu Gly Val Lys Asp Glu Glu Glu Gly Lys Ile Ile Leu Gly
210 215 220
Phe Asp Leu Asn Glu Pro Ala Val Glu Glu Glu Glu Glu Glu Lys Glu
225 230 235 240
Asp Lys Pro Phe
<210> 46
<211> 202
<212> PRT
<213> Cajanus cajan
<400> 46
Met Gln Glu Gln Asn Pro Ser Glu Asn Ser Pro Ser Asn Asp Asp Ala
1 5 10 15
Pro Asn Pro Gln Gly Gly Glu Gly Glu Ile Met Lys Thr Pro Trp Gln
20 25 30
Val Leu Glu Glu Leu Cys Gly Gly Ser Ser Thr Pro Val Ala Val Ala
35 40 45
Pro Val Ser Glu Gly Val Leu Gln Pro Lys Lys Arg Lys Thr Ser Glu
50 55 60
Val Arg Asp Pro Pro Arg Gly Lys Pro Ile Cys Pro Val Cys Asn Lys
65 70 75 80
Glu Phe Thr Ser Trp Lys Ala Ala Phe Gly His Met Arg Ala His Pro
85 90 95
Asn Arg Asp Tyr Arg Gly Phe Tyr Lys Pro Pro Ala Phe Pro Ser Ser
100 105 110
Ser Ser Ser Lys Asp Gln Pro Pro Pro Ser Ala Asn Asn Asn Lys Gly
115 120 125
Gly Gly Ala Lys Lys Thr Asn Ala Ala Ser Asp Gly Thr Asp Glu Val
130 135 140
Asp Ser Gly Gly Gly Lys Glu Val Thr Pro Ser Pro Asn Gln Leu Phe
145 150 155 160
Gly Phe Asp Leu Asn Glu Pro Val Glu Gly Leu Gly Ser Ser His Ala
165 170 175
Val Glu Glu Gly Val Gln Glu Glu Lys Asp Leu Gly Phe Asp Leu Asn
180 185 190
Glu Met Pro Pro Ala Glu Asp His Glu Gly
195 200
<210> 47
<211> 265
<212> PRT
<213> Brassica rapa
<400> 47
Met Thr Asn Pro Glu Lys Met Lys Val Asp Gly Asp Thr Glu Lys Glu
1 5 10 15
Lys Glu Asp Ser Ser Asp Glu Gln Trp Ser Asp Glu Glu Ser Ala Met
20 25 30
Arg Glu Ile Val Leu Gly Leu Pro Ala Leu Ser Ile Ser Ser Ala Thr
35 40 45
Phe Gly Val Ser Ile Ala Val Asp Glu Glu Glu Glu Ala Arg Leu Asn
50 55 60
Glu Gln Ala Val Val Ala Ala Glu Leu Val Ile Ala Ala Ala Glu Glu
65 70 75 80
Ala Val Met Lys Glu Lys Ser Asp Gly Lys Lys Lys Lys Val Arg Arg
85 90 95
Gln Arg Lys Thr Met Lys Leu Asn Asn Asp Asp Glu Ala Gly Gly Ser
100 105 110
Ser Lys Gly Gly Glu Ala Ala Lys Lys Pro Arg Lys Lys Thr Ser Glu
115 120 125
Phe Thr Asn Leu Pro Arg Gly Pro Pro Val Cys Asn Val Cys Gly Arg
130 135 140
Ala Phe His Ser Trp Lys Ala Val Phe Gly His Leu Arg Ser His Lys
145 150 155 160
Asp Arg Ser Tyr Arg Gly Phe Leu Pro Pro Pro Arg Phe Ser Ala Pro
165 170 175
Ile Arg Gly Val Met Ile Pro Gly Pro Asn Ser Ala Phe Val Arg Val
180 185 190
Val Ala Arg Gly Gly Ser Ser Gly Val Val Val Ala Ser Gly Gly Gly
195 200 205
Gly Gly Gly Gly Ala Ser Gly Gly Glu Gly Gly Arg Gly Val Gly Ile
210 215 220
Val Met Asp Val Asp Arg Val Glu Lys Glu Lys Gln Glu Val Thr Glu
225 230 235 240
Ser Gly Thr Ala Ala Lys Phe Asp Leu Asn Lys Ser Pro Pro Lys Asp
245 250 255
Asp Asp Glu Glu Glu Asp Lys Thr Lys
260 265
<210> 48
<211> 237
<212> PRT
<213> Lagenaria siceraria
<400> 48
Met Glu Lys Asn Thr Asn Ala Asn Ala Asn Asp Asn Thr Val Ser Glu
1 5 10 15
Thr Ser Pro Asp Gln Arg His Gly Gly Glu Arg Ser Pro Met Ala Ser
20 25 30
Ser Pro Pro Pro Ser Ala Arg Asn Asp Asp Val Asn Asn Pro Thr Pro
35 40 45
Asn Val Val Ile Glu Leu Ser Ser Ser Val Val Val Ala Ala Pro Glu
50 55 60
Ser Thr Thr Thr Gly Asp Val Arg Arg Arg Gln Thr Val Asp Ala Thr
65 70 75 80
Leu Leu Asp Val Gly Thr Ser Ser His Asp His Val Gly Gly Ser Ser
85 90 95
Asp Ile Glu Gln Gly Lys Lys Arg Gly Arg Gly Asp Gly Gly Glu Gln
100 105 110
Gln Gln Gln Val Lys Ala Ala Lys Lys Lys Gly Glu Leu Thr Glu Val
115 120 125
Pro Lys Gly Glu Pro Arg Cys Ala Thr Cys Asn Lys Val Phe Lys Ser
130 135 140
Trp Lys Ala Leu Phe Gly His Leu Arg Ser His Pro Glu Arg Thr Tyr
145 150 155 160
Arg Gly Ala Leu Pro Pro Pro Thr Ala Ala Glu Leu Asp Ile Arg Arg
165 170 175
Cys Gln Gln Gln Leu Ala Ser Thr Leu Leu Thr Val Ala Gln Gln Val
180 185 190
Ala Thr Ser Arg Arg Gly Leu Asn Ile Asp Leu Asn Gln Pro Ser Ala
195 200 205
Ala Asp Asp Gly Asp Ser Pro Glu Lys Thr Gly Ser Val Gly Phe Asp
210 215 220
Leu Asn Val Glu Pro Pro Pro Glu Ser Asp Asp Glu Lys
225 230 235
<210> 49
<211> 243
<212> PRT
<213> Arachis ipaensis
<400> 49
Met Asp Lys Asn Ala Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser
1 5 10 15
Ser Ser Pro Ser Thr Ser Thr Thr Glu Gln Pro Pro Pro Pro Ser Pro
20 25 30
Gln Asp Val Pro Val Glu Glu Pro Pro Val Ile Leu Lys Thr Pro Glu
35 40 45
Glu Val Leu Ala Glu Leu Thr Glu Ser Gly Glu Ser Pro Pro Pro Pro
50 55 60
Pro Pro Ser Ala Ala Glu Glu Gly Gly Ile Gly Gly Ser Ala Met Ala
65 70 75 80
Ile Ala Val Gly Val Gly Gly Glu Gly Val Val Leu Gly Glu Ala Gly
85 90 95
Ser Gly Gly Ser Gly Ala Val Gly Ala Ser Arg Lys Arg Lys Ala Ser
100 105 110
Glu Val Lys Asp Pro Pro Thr Gly Thr Pro Thr Cys Pro Val Cys Asn
115 120 125
Lys Thr Phe Ala Ser Trp Lys Gly Ala Phe Gly His Met Arg Lys His
130 135 140
Pro Asp Arg Gln Tyr Arg Gly Phe Phe Lys Pro Pro Thr Phe Thr Ser
145 150 155 160
Ser Val Ala Ala Ala Ala Thr Ser Gly Ala Thr Thr Gln Gly Phe Val
165 170 175
Glu Gly Gly Ser Ser Ala Ala Ser Gly Glu Gly Gly Ser Val Gly Ala
180 185 190
Ala Gly Gln Val Gly Glu Gly Gly Ser Phe Asp Leu Asn Gln Pro Val
195 200 205
Met Glu Arg Gly Gly Ala Gly Asp Gly Asp Glu Gly Arg Asn Gly Asp
210 215 220
Glu Gly Lys Asp Leu Gly Phe Asp Leu Asn Lys Pro Ala Asp Glu Glu
225 230 235 240
Asp Lys Ala
<210> 50
<211> 271
<212> PRT
<213> Prunus persica
<400> 50
Met Ala Asn Gln Gln Asn Thr Thr Thr Asn Asn Thr Asn Arg Leu Ala
1 5 10 15
Ser Ser Ala Glu Gly Gln Gln Gln Pro Gly Asn Asp Asn Val Asn Val
20 25 30
Asn Val Thr Gly Val Ala Ala Thr Gly Thr Pro Ser Phe Pro Asn Pro
35 40 45
Pro Pro Pro Ser Thr Val Gln Leu Asn Val Ala Arg Gly Ser Ser Val
50 55 60
Ser Pro Pro Gly Leu Asn Arg Gly Pro Ser Gly Arg Gly Ser Ser Asn
65 70 75 80
Pro Arg Arg Val Ser Pro Ser Gly Arg Lys Ser Pro Pro Gly Gly Arg
85 90 95
Gly Ser Pro Ser Ser Gly Arg Gly Ser Ser Ser Phe Ala Ser Ser Pro
100 105 110
Gly Ala Gly Gln Ile Thr Gln Ile Gly Gly Arg Asn Phe Thr Pro Leu
115 120 125
Ser Val Glu Ile Thr Ile Gly Gly Gly Ser Gly Thr Ala Gly Gly Ser
130 135 140
Met Ile Pro Gly Ser Ser Ser Gly Ser Pro Pro Arg Ala Lys Arg Lys
145 150 155 160
Ala Glu Ile Thr Ala Pro Glu Gly Val Ser Pro Val Cys Ser Val Cys
165 170 175
Lys Arg Ser Asn Phe Ser Ser Trp Lys Ala Leu Phe Gly His Met Arg
180 185 190
Cys His Pro Glu Arg Gln Trp Arg Gly Cys Phe Pro Pro Pro Gly Phe
195 200 205
Glu Glu Ala Gln Arg Ala Leu Gln Gln Gly Glu Arg Leu Gly Ala Leu
210 215 220
Arg Gly Phe Asp Leu Asn Glu Ala Ser Asp Pro Glu Glu Glu Asn Glu
225 230 235 240
Gly Gly Gly Gly Gly Gly Phe Asp Leu Asn Met Leu Pro Pro Asp Glu
245 250 255
Asp Lys Asp Gly Gly Gly Ser Ser Gly Gly Ala Ala Lys Thr Gly
260 265 270
<210> 51
<211> 321
<212> PRT
<213> Glycine max
<400> 51
Met Gln Asn Gln Asn Pro Ser Glu Asp Asp Ser Thr Asn Lys Ala Pro
1 5 10 15
Ser Asp Asp Thr Pro His Pro Thr Asn Asn Asp Asn Asn Asn Pro Pro
20 25 30
Gly Glu Ser Leu Ile Met Lys Thr Val Arg Gln Val Leu Glu Glu Leu
35 40 45
Gly Gly Ser Arg Asp Asp Asn Asp Gly Gly Val Gly Asp Gly Thr Pro
50 55 60
Val Ala Val Ala Leu Glu Val Gly Gly Gly Gly Asp Gly Ser Gly Gly
65 70 75 80
Asp Gly Gly Gly Val Gly Gly Gly Pro Pro Val Ala Met Glu Ala Ser
85 90 95
Asp Gly Gly Asp Glu Asp Asn Asp Asn Glu His Ser Asp Ser Ile Gly
100 105 110
Asp Ser Val Pro Val Ala Val Ala Val Lys Val Gly Gly Arg Gly Gly
115 120 125
Asp Asn Ser Gly Gly Gly Ala Thr Val Thr Val Ser Val Ala Leu Ser
130 135 140
Asp Gly Asp Tyr Gly Gly Ser Asn Gly Ser Gly Gly Ser Pro Val Ala
145 150 155 160
Val Ala Val Ala Val Gly Gly Gly Ser Thr Val Met Glu Ser Val Val
165 170 175
Val Gly Thr Glu Gly Val Gln Gly Arg Lys Arg Lys Thr Pro Leu Val
180 185 190
Arg Asp Pro Pro Thr Gly Arg Pro Thr Cys Pro Leu Cys Gln Lys Glu
195 200 205
Phe Gln Thr Trp Lys Gly Ala Phe Gly His Met Arg Ala His Pro Asp
210 215 220
Arg Asp Tyr Arg Gly Phe Phe Lys Pro Pro Val Phe Gly Ser Pro Ser
225 230 235 240
Ser Thr Gln Asp Gln Pro Pro Ser Asp Gly Lys Gly Asp Asp Ser Ala
245 250 255
Lys Lys Ser Thr Gly Glu Asp Asn Thr Ala Glu Lys Gly Ser Ala Ser
260 265 270
Leu Pro Val Arg Val Pro Met Phe Asp Leu Asn Glu Leu Ile Glu Glu
275 280 285
Asp Gly Ser Ser His Ala Ala Glu Pro Ala Glu Asp Met Ser Thr Gly
290 295 300
Glu Gly Lys Gly Ser Gly Phe Asp Leu Asn Glu Met Pro Pro Ala Glu
305 310 315 320
Asp
<210> 52
<211> 355
<212> PRT
<213> Glycine max
<400> 52
Met Asn His His Lys Tyr Ser Ser His Arg His Asn Leu Phe Asn Ser
1 5 10 15
Phe Leu Thr Leu Ala Ile Phe Ile Ile Leu Gly Phe Ser Phe Ser Ala
20 25 30
Thr Gly Met Gln Asn Gln Asn Pro Ser Glu Asp Asp Ser Thr Asn Lys
35 40 45
Ala Pro Ser Asp Asp Thr Pro His Pro Thr Asn Asn Asp Asn Asn Asn
50 55 60
Pro Pro Gly Glu Ser Leu Ile Met Lys Thr Val Arg Gln Val Leu Glu
65 70 75 80
Glu Leu Gly Gly Ser Arg Asp Asp Asn Asp Gly Gly Val Gly Asp Gly
85 90 95
Thr Pro Val Ala Val Ala Leu Glu Val Gly Gly Gly Gly Asp Gly Ser
100 105 110
Gly Gly Asp Gly Gly Gly Val Gly Gly Gly Pro Pro Val Ala Met Glu
115 120 125
Ala Ser Asp Gly Gly Asp Glu Asp Asn Asp Asn Glu His Ser Asp Ser
130 135 140
Ile Gly Asp Ser Val Pro Val Ala Val Ala Val Lys Val Gly Gly Arg
145 150 155 160
Gly Gly Asp Asn Ser Gly Gly Gly Ala Thr Val Thr Val Ser Val Ala
165 170 175
Leu Ser Asp Gly Asp Tyr Gly Gly Ser Asn Gly Ser Gly Gly Ser Pro
180 185 190
Val Ala Val Ala Val Ala Val Gly Gly Gly Ser Thr Val Met Glu Ser
195 200 205
Val Val Val Gly Thr Glu Gly Val Gln Gly Arg Lys Arg Lys Thr Pro
210 215 220
Leu Val Arg Asp Pro Pro Thr Gly Arg Pro Thr Cys Pro Leu Cys Gln
225 230 235 240
Lys Glu Phe Gln Thr Trp Lys Gly Ala Phe Gly His Met Arg Ala His
245 250 255
Pro Asp Arg Asp Tyr Arg Gly Phe Phe Lys Pro Pro Val Phe Gly Ser
260 265 270
Pro Ser Ser Thr Gln Asp Gln Pro Pro Ser Asp Gly Lys Gly Asp Asp
275 280 285
Ser Ala Lys Lys Ser Thr Gly Glu Asp Asn Thr Ala Glu Lys Gly Ser
290 295 300
Ala Ser Leu Pro Val Arg Val Pro Met Phe Asp Leu Asn Glu Leu Ile
305 310 315 320
Glu Glu Asp Gly Ser Ser His Ala Ala Glu Pro Ala Glu Asp Met Ser
325 330 335
Thr Gly Glu Gly Lys Gly Ser Gly Phe Asp Leu Asn Glu Met Pro Pro
340 345 350
Ala Glu Asp
355
<210> 53
<211> 357
<212> PRT
<213> Glycine max
<400> 53
Met Asn His His Lys Tyr Ser Ser His Arg His Asn Leu Phe Asn Ser
1 5 10 15
Phe Leu Thr Leu Ala Ile Phe Ile Ile Leu Gly Phe Ser Phe Ser Ala
20 25 30
Thr Gly Met Gln Asn Gln Asn Pro Ser Glu Asp Asp Ser Thr Asn Lys
35 40 45
Ala Pro Ser Asp Asp Thr Pro His Pro Thr Asn Asn Asp Asn Asn Asn
50 55 60
Pro Pro Gly Glu Ser Leu Ile Met Lys Thr Val Arg Gln Val Leu Glu
65 70 75 80
Glu Leu Gly Gly Ser Arg Asp Asp Asn Asp Gly Gly Val Gly Asp Gly
85 90 95
Thr Pro Val Ala Val Ala Leu Glu Val Gly Gly Gly Gly Asp Gly Ser
100 105 110
Gly Gly Asp Gly Gly Gly Val Gly Gly Gly Pro Pro Val Ala Met Glu
115 120 125
Ala Ser Asp Gly Gly Asp Glu Asp Asn Asp Asn Glu His Ser Asp Ser
130 135 140
Ile Gly Asp Ser Val Pro Val Ala Val Ala Val Lys Val Gly Gly Arg
145 150 155 160
Gly Gly Asp Asn Ser Gly Gly Gly Ala Thr Val Thr Val Ser Val Ala
165 170 175
Leu Ser Asp Gly Asp Tyr Gly Gly Ser Asn Gly Ser Gly Gly Ser Pro
180 185 190
Val Ala Val Ala Val Ala Val Gly Gly Gly Ser Thr Val Met Glu Ser
195 200 205
Val Val Val Gly Thr Glu Gly Val Gln Gly Arg Lys Arg Lys Thr Pro
210 215 220
Leu Val Arg Asp Pro Pro Thr Gly Arg Pro Thr Cys Pro Leu Cys Gln
225 230 235 240
Lys Glu Phe Gln Thr Trp Lys Gly Ala Phe Gly His Met Arg Ala His
245 250 255
Pro Asp Arg Asp Tyr Arg Gly Phe Phe Lys Pro Pro Val Phe Gly Ser
260 265 270
Pro Ser Ser Thr Gln Asp Gln Pro Pro Ser Asp Gly Lys Val Ala Gly
275 280 285
Asp Asp Ser Ala Lys Lys Ser Thr Gly Glu Asp Asn Thr Ala Glu Lys
290 295 300
Gly Ser Ala Ser Leu Pro Val Arg Val Pro Met Phe Asp Leu Asn Glu
305 310 315 320
Leu Ile Glu Glu Asp Gly Ser Ser His Ala Ala Glu Pro Ala Glu Asp
325 330 335
Met Ser Thr Gly Glu Gly Lys Gly Ser Gly Phe Asp Leu Asn Glu Met
340 345 350
Pro Pro Ala Glu Asp
355
<210> 54
<211> 324
<212> PRT
<213> Glycine max
<400> 54
Met Gln Asn Gln Asn Pro Ser Glu Asp Asp Ser Thr Lys Lys Ser Pro
1 5 10 15
Ser Ser Pro Ser Asp Asp Thr Ser His Pro Thr Asn Asn Asn Asn Asn
20 25 30
Pro Gln Gly Glu Ser Leu Val Met Lys Thr Val Trp Gln Val Leu Glu
35 40 45
Glu Leu Gly Gly Ser Gly Asp Asp Ser Asp Gly Asp Ser Gly Phe Gly
50 55 60
Asp Gly Ala Pro Val Ala Val Ala Val Gly Gly Gly Gly Asp Asp Asn
65 70 75 80
Gly Asp Gly Asp Gly Ser Val Gly Gly Gly Ala Thr Val Ala Met Glu
85 90 95
Lys Ser Asp Asp Asp Ser Asp Asn Asn Gly Asp Asn Ser Gly Ser Asn
100 105 110
Leu Ala Val Ala Val Val Val Gly Gly Gly Asp Gly Ser Gly Ser Ser
115 120 125
Gly Val Gly Ala Thr Val Thr Val Ser Val Ala Val Ser Gly Gly Gly
130 135 140
Gly Asn His Gly Gly Gly Ser Ser Gly Gly Gly Gly Gly Gly Pro Val
145 150 155 160
Ala Val Ser Val Ala Val Gly Gly Gly Ser Thr Val Val Glu Ser Val
165 170 175
Val Val Ser Thr Glu Gly Val Gln Gly Arg Lys Arg Lys Thr Ser Ile
180 185 190
Val Arg Asp Pro Pro Thr Gly Arg Pro Thr Cys Pro Leu Cys Gln Lys
195 200 205
Glu Phe Pro Thr Trp Lys Gly Ala Phe Gly His Met Arg Ala His Pro
210 215 220
Asp Arg Asp Tyr Arg Gly Phe Phe Lys Pro Pro Val Phe Gly Ser Pro
225 230 235 240
Ser Ser Thr Gln Asp Gln Pro Pro Arg Asp Asp Lys Val Ala Gly Asp
245 250 255
Asp Ser Ala Lys Lys Ser Ser Gly Glu Val Asn Asn Gly Glu Lys Gly
260 265 270
Ser Ala Ser Ser His Val Arg Val Pro Met Phe Asp Leu Asn Glu Pro
275 280 285
Ile Glu Glu Glu Gly Ser Ser His Ala Ala Glu Pro Ala Glu Asp Met
290 295 300
Ser Ser Glu Glu Gly Lys Gly Phe Gly Phe Asp Leu Asn Glu Met Pro
305 310 315 320
Pro Ala Glu Asp
<210> 55
<211> 280
<212> PRT
<213> Cicer arietinum
<400> 55
Met Lys Arg Phe Cys His Ser Phe His Lys Thr Glu Lys His Asp Asp
1 5 10 15
Tyr Val Trp Pro Leu Leu Ala His Glu Met Lys Lys Asn Gly Ser Arg
20 25 30
Ile Phe Ile Cys Asp Leu Cys Asp Lys Ser Phe Ser Ser Gly Asn Ala
35 40 45
Leu Gly Gly His Lys Ala Ser His Asn Arg Ser Asp Leu Leu Gln Pro
50 55 60
Pro Ile Lys Lys His Lys Leu Thr Ile Asp Ser Cys Ser Leu Ser Ser
65 70 75 80
Pro Asn Asp His Asp Asp Val Lys His Lys His Ala Cys Val Leu Cys
85 90 95
His Lys Val Phe Pro Ser Asn Lys Ala Leu Tyr Gly His Met Arg Ser
100 105 110
His Ser Gln Lys Asp Ser Lys Ala Ile Gln Pro Pro Leu Ile Thr Thr
115 120 125
Thr Ser Arg Asp Ser Lys Ile Gln Ser Asn Asn Thr Asp Asp Gln Pro
130 135 140
Ile Leu Pro Ala Ile Asp Leu Glu Lys Tyr Phe Pro Pro Arg Ser His
145 150 155 160
Gln Thr Lys Lys Arg Cys Ser Lys Ser Thr Ile Asp Tyr Glu Leu Ile
165 170 175
Asn Val Ala Gln Ile Leu Cys Asp Met Ser Arg Ser Asp Pro Thr Lys
180 185 190
Phe Thr Thr Asn Ile Asp Asn Gln Arg Asn Lys Glu His Val Thr Ile
195 200 205
Val Lys Asn Asn Asn Asn Pro Lys Lys Leu Val Val Thr Phe Lys Ile
210 215 220
Pro Lys Asp Lys Thr Phe Lys Ile Pro Lys Asp Lys Glu Glu Lys Asn
225 230 235 240
Lys Glu Ala Glu Met Glu Lys Glu Ser Ser His Gln Leu Gly Ser Arg
245 250 255
Ala Val Ala Lys Asp Phe Asp Leu Asn Glu Ile Pro Val Asp Leu Asp
260 265 270
Leu Val Ala Asp Glu Gln Ala Pro
275 280
<210> 56
<211> 290
<212> PRT
<213> Cicer arietinum
<400> 56
Met Asp Val Asp Glu Leu Gln Ala Ala Glu Arg Glu Asp Arg Arg Ser
1 5 10 15
Ile Ile Ile Met Lys Arg Ser Cys Gln Ser Phe Asp Lys Pro Met Lys
20 25 30
Asn Asp Asp Phe Val Trp Pro Leu Leu Ala Asn Glu Met Lys Lys Asn
35 40 45
Gly Ser Arg Ile Phe Ile Cys Asp Leu Cys Gly Lys Ser Phe Ser Ser
50 55 60
Gly Asn Ala Leu Gly Gly His Lys Ala Ser His Asn Arg Ser Asn Leu
65 70 75 80
Leu Gln Pro Pro Ile Lys Lys His Lys Leu Thr Ile Asp Ser Cys Ser
85 90 95
Leu Ser Ser Pro His Asp His Asp Asp Val Lys His Lys His Ala Cys
100 105 110
Val Leu Cys His Lys Val Phe Pro Ser Asn Lys Val Leu Tyr Gly His
115 120 125
Met Arg Ser His Ser Gln Lys Asp Ser Lys Ala Ile Gln Pro Pro Pro
130 135 140
Ile Thr Thr Thr Ser Pro Asp Ser Lys Lys Gln Ser Asn Asn Thr His
145 150 155 160
Asp Gln Pro Ile Leu Pro Thr Ile Asp Leu Glu Lys Tyr Phe Pro Pro
165 170 175
Arg Ser His Gln Thr Lys Lys Arg Arg Ser Lys Ser Ile Ile Asp Tyr
180 185 190
Glu Leu Ile Asn Val Ala Gln Ile Leu Cys Asp Met Ser Arg Ser Asp
195 200 205
Pro Lys Arg Leu Lys Leu Ser Thr Asn Ile Asp Asn Gln Lys Asn Lys
210 215 220
Glu His Val Thr Ile Val Lys Asn Asn Asn Asn Glu Lys Lys Leu Val
225 230 235 240
Val Arg Phe Lys Ile Pro Lys Asp Lys Glu Val Lys Asn Asn Glu Gly
245 250 255
Glu Thr Lys Lys Glu Ser Ser His Gln Leu Gly Ser Arg Gly Val Val
260 265 270
Lys Asp Phe Asp Leu Asn Glu Ile Pro Val Asp Val Ala Asp Glu Gln
275 280 285
Ala Asn
290
<210> 57
<211> 246
<212> PRT
<213> Cicer arietinum
<400> 57
Met Lys Lys Asn Gly Ser Arg Ile Phe Ile Cys Asp Leu Cys Gly Lys
1 5 10 15
Ser Phe Ser Ser Gly Asn Ala Leu Gly Gly His Lys Ala Ser His Asn
20 25 30
Arg Ser Asn Leu Leu Gln Pro Pro Ile Lys Lys His Lys Leu Thr Ile
35 40 45
Asp Ser Cys Ser Leu Ser Ser Pro His Asp His Asp Asp Val Lys His
50 55 60
Lys His Ala Cys Val Leu Cys His Lys Val Phe Pro Ser Asn Lys Val
65 70 75 80
Leu Tyr Gly His Met Arg Ser His Ser Gln Lys Asp Ser Lys Ala Ile
85 90 95
Gln Pro Pro Pro Ile Thr Thr Thr Ser Pro Asp Ser Lys Lys Gln Ser
100 105 110
Asn Asn Thr Asp Asp Gln Pro Ile Leu Pro Thr Ile Asp Leu Glu Lys
115 120 125
Tyr Phe Pro Pro Arg Ser His Gln Thr Lys Lys Arg Arg Ser Lys Ser
130 135 140
Ile Ile Asp Tyr Glu Leu Ile Asn Val Ala Gln Ile Leu Cys Asp Met
145 150 155 160
Ser Arg Ser Asp Pro Lys Arg Leu Lys Leu Ser Thr Asn Ile Asp Asn
165 170 175
Gln Lys Asn Lys Glu His Val Thr Ile Val Lys Asn Asn Asn Asn Glu
180 185 190
Lys Lys Leu Val Val Arg Phe Lys Ile Pro Lys Asp Lys Glu Val Lys
195 200 205
Asn Asn Glu Gly Glu Thr Lys Lys Glu Ser Ser His Gln Leu Gly Ser
210 215 220
Arg Gly Val Val Lys Asp Phe Asp Leu Asn Glu Ile Pro Val Asp Val
225 230 235 240
Ala Asp Glu Gln Ala Asn
245
<210> 58
<211> 6
<212> PRT
<213> Artificial Sequence
<220>
<223> EAR motif
<220>
<221> misc_feature
<222> (4)..(5)
<223> Xaa can be any naturally occurring amino acid
<400> 58
Asp Leu Asn Xaa Xaa Pro
1 5
<210> 59
<211> 5
<212> PRT
<213> Artificial Sequence
<220>
<223> EAR motif
<220>
<221> misc_feature
<222> (4)..(4)
<223> Xaa can be any naturally occurring amino acid
<400> 59
Asp Leu Asn Xaa Pro
1 5
<210> 60
<211> 1335
<212> DNA
<213> Artificial Sequence
<220>
<223> Tar-MITE insert
<400> 60
actgctacca gggccggcca gaaggctgtg cggcttgggc tgtggcacag gggcagcaga 60
tttagtccat ttattatggt ctgctattta tttgggtcca ataagatcag ggcactaaaa 120
ctcaaataca tcaattcaag tcggcggcaa aagaataaaa agcgccaatt tttagaagac 180
aaggagacag cacaataatc atgcttttcc aatggccaat taattttagc ctttgtgagg 240
tataaatatg aatcactttt tctccaaata tcattattgt aaactagtaa agagcgtaaa 300
ttgttttatc gctaatcttc ttctttttca aatataattg tttgattctt tgtagtaggt 360
acgttaagtt ttatatcatt cgtcctggaa accttttctt atgccccttc aatttatgaa 420
tttgtgtttt gtattgtctt gctattaatc caatgcaaac taattctgcc gcatcgatct 480
cacagagttg gagcagctta aaaggagttt tggggcaact taaaatgagg gtagcttttt 540
tgttgttgtt ggtctactat ttactttggt ccttatcgtt ttagttttgt gttcattgag 600
gatttgaatg actttgaaca caagtttata aaactaagtt tttgtgattc taaatttttg 660
atgtgtgcaa ccgactaaat gaatataaaa ttctgatttg tgtatttctg atttctgatt 720
taccggttat tcattatgtt tttctggatc tcatatgaat ttagtatata aaaatctcag 780
gttcttctta ctcagtctag tatattttgg cattggggat cagggttaaa atcatcattt 840
ttctaagcac taacaagctt taaaaccatc aattaaaagt tagttctcca tacctggaat 900
tgtttattgc tttagtattc atttcatagt gtcaatacct tacaactgtt tattgttttc 960
aaatgtttac tgttttgcac tttagttgtt tttagcatgt ttcaactttc aaagttaatt 1020
gacgatagtg tcatatacga aatatcatca ctcatacaaa attaaactgg atagaatttt 1080
ttccgatgaa atctattttt aaatagttac gaatataata ggagacttaa tatgtttgtt 1140
tcaaaattac acaaaaacac catggaaatg ggcattagtt tgtataagca tccttttaaa 1200
tataaagctt tgttacattg atgccttcta aatatataat attgtaatga tttgtagtaa 1260
acaccatgga aatgggcatt attaagatgt ttcgcccagg gcatgaaaaa ctatcggacc 1320
ggccctgact gctac 1335
Claims (15)
1.一种与植物孤雌生殖相关的核酸，其中所述核酸包含以下中的至少一种：
a)编码具有SEQ ID NO：1、6或11的氨基酸序列的蛋白质的基因；
b)具有SEQ ID NO：2、7或12的核苷酸序列的启动子；
c)具有SEQ ID NO：3、8或13的核苷酸序列的编码序列；
d)具有SEQ ID NO：4、9或14的核苷酸序列的3'UTR；
e)具有SEQ ID NO：5、10或15的核苷酸序列的基因；
f)a)-e)中任一项的变体或片段；
其中优选地，所述核酸在孤雌生殖中是功能性的。
2.权利要求1的核酸，其中所述核酸包含在嵌合基因、遗传构建体或核酸载体中。
3.一种与植物孤雌生殖相关的蛋白质，其中所述蛋白质：
a)由权利要求1所述的核酸编码；
b)具有SEQ ID NO：1、6或11的氨基酸序列；和/或
c)是a)和/或b)的变体或片段；
其中优选地所述蛋白质在孤雌生殖中是功能性的。
4.一种不属于Taraxacum officinale sensu lato物种的植物或植物细胞，其包含权利要求1所述的核酸和/或权利要求3所述的蛋白质，其中所述植物或植物细胞优选来自选自以下的家族：十字花科(Brassicaceae)、葫芦科(Cucurbitaceae)、豆科(Fabaceae)、禾本科(Gramineae)、茄科(Solanaceae)、菊科(Asteraceae)、蔷薇科(Rosaceae)和禾本科(Poaceae)。
5.权利要求4的植物或植物细胞，其中所述植物或植物细胞通过遗传修饰或通过基因渗入包含权利要求1所述的核酸，其中优选地所述核酸被整合至其基因组中。
6.权利要求4或5的植物或植物细胞，其中所述植物或植物细胞能够进行孤雌生殖。
7.权利要求4-6中任一项的植物或植物细胞，其中所述植物或植物细胞还能够进行不完全减数分裂，优选地其中所述植物或植物细胞能够进行无融合生殖。
8.权利要求4-7中任一项的植物或植物细胞的种子、植物部分或植物产品。
9.一种用于产生孤雌生殖的植物的方法，其包含以下步骤：
a)在一种或多种植物细胞中引入能够诱导孤雌生殖的权利要求1的核酸；
b)选择包含所述核酸的植物细胞，其中优选地所述核酸被整合至所述植物细胞的基因组中；以及
c)从所述植物细胞再生植物。
10.一种用于产生无融合生殖植物的方法，其包含权利要求9所述的步骤a)至c)，其中步骤a)的所述一种或多种植物细胞能够进行不完全减数分裂。
11.一种用于产生无融合生殖的F1杂种种子的方法，其包含以下步骤：
a)将有性繁殖的第一植物与第二植物的花粉进行杂交育种以产生F1杂种种子，其中所述第二植物包含权利要求1的核酸，并且其中所述第一植物和/或第二植物能够进行不完全减数分裂。
12.权利要求11所述的方法，其中所述方法还包含以下步骤：
b)从所述的F1种子中选择包含无融合生殖表型的，优选通过基因分型选择。
13.一种用于产生无融合生殖的杂种植物的方法，其包含权利要求11或12所述的步骤，并且还包含以下步骤：
c)从所述F1杂种种子培育至少一株F1植物。
14.可通过权利要求9-13中任一项的方法获得的植物、种子、植物部分或植物产品。
15.权利要求1或2的核酸或权利要求3的蛋白质在筛选植物或植物细胞中的孤雌生殖基因、对植物或植物细胞进行孤雌生殖的基因分型和/或赋予植物或植物细胞孤雌生殖中的用途。
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| EP19177252.4 | 2019-05-29 | ||
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| EP19200872.0 | 2019-10-01 | ||
| EP20170243 | 2020-04-17 | ||
| EP20170243.8 | 2020-04-17 | ||
| PCT/EP2020/064991 WO2020239984A1 (en) | 2019-05-29 | 2020-05-29 | Gene for parthenogenesis |
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| CN113874388A true CN113874388A (zh) | 2021-12-31 |
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| CN202080039188.4A Pending CN113874388A (zh) | 2019-05-29 | 2020-05-29 | 孤雌生殖基因 |
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| US (1) | US20220106607A1 (zh) |
| EP (1) | EP3976633A1 (zh) |
| JP (1) | JP2022533813A (zh) |
| CN (1) | CN113874388A (zh) |
| AU (1) | AU2020285344A1 (zh) |
| BR (1) | BR112021023769A2 (zh) |
| CA (1) | CA3138988A1 (zh) |
| IL (1) | IL287956A (zh) |
| WO (1) | WO2020239984A1 (zh) |
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| CN116746490A (zh) * | 2023-07-03 | 2023-09-15 | 安徽农业大学 | 一种提高乌菜游离小孢子出胚率及胚性培养物增殖的方法 |
| CN117660525A (zh) * | 2024-01-30 | 2024-03-08 | 三亚中国农业科学院国家南繁研究院 | 一种水稻单倍体诱导方法 |
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| AU2021363100A1 (en) * | 2020-10-13 | 2023-05-25 | Keygene N.V. | Modified promoter of a parthenogenesis gene |
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| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN116589550A (zh) * | 2023-06-27 | 2023-08-15 | 中农集团种业控股有限公司 | 固定水稻杂种优势的方法 |
| CN116589550B (zh) * | 2023-06-27 | 2023-10-03 | 中农集团种业控股有限公司 | 固定水稻杂种优势的方法 |
| CN116746490A (zh) * | 2023-07-03 | 2023-09-15 | 安徽农业大学 | 一种提高乌菜游离小孢子出胚率及胚性培养物增殖的方法 |
| CN116746490B (zh) * | 2023-07-03 | 2024-02-02 | 安徽农业大学 | 一种提高乌菜游离小孢子出胚率及胚性培养物增殖的方法 |
| CN117660525A (zh) * | 2024-01-30 | 2024-03-08 | 三亚中国农业科学院国家南繁研究院 | 一种水稻单倍体诱导方法 |
| CN117660525B (zh) * | 2024-01-30 | 2024-04-26 | 三亚中国农业科学院国家南繁研究院 | 一种水稻单倍体诱导方法 |
Also Published As
| Publication number | Publication date |
|---|---|
| WO2020239984A1 (en) | 2020-12-03 |
| IL287956A (en) | 2022-01-01 |
| AU2020285344A1 (en) | 2021-11-11 |
| BR112021023769A2 (pt) | 2022-01-11 |
| CA3138988A1 (en) | 2020-12-03 |
| JP2022533813A (ja) | 2022-07-26 |
| US20220106607A1 (en) | 2022-04-07 |
| EP3976633A1 (en) | 2022-04-06 |
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