CN105492600B - 向日葵中的霜霉菌抗性提供基因 - Google Patents
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Abstract
本发明涉及向日葵中的霜霉菌抗性基因，尤其涉及对霜霉菌具有抗性的向日葵植物。本发明特别涉及对植物病原体霜霉菌具有抗性的向日葵植物，其中，所述植物包括：编码包括如SEQ ID No.2或SEQ ID No.4所示的氨基酸序列的蛋白的霜霉菌抗性赋予基因，或者，编码与SEQ ID No.2或SEQ ID No.4本身具有超过90％序列同一性、优选超过94％序列同一性、更优选超过96％序列同一性的蛋白的霜霉菌抗性赋予基因；并且其中，与对植物病原体霜霉菌没有抗性的向日葵植物中的所述抗性赋予基因的表达相比，所述抗性赋予基因的表达是降低的，或者与对植物病原体霜霉菌没有抗性的向日葵植物中的所述蛋白的酶活性相比，所述蛋白的酶活性是降低的。
Description
技术领域
本发明涉及向日葵中的霜霉菌抗性基因，尤其涉及具有霜霉菌抗性的向日葵植物。本发明进一步涉及获得所述具有霜霉菌抗性的向日葵植物的方法，以及所述基因在向日葵中提供霜霉菌抗性的应用。
背景技术
向日葵属(Helianthus L.)是包括菊(Asteraceae)科中的约52种植物的属。常用名称“向日葵”通常用于表示一年生物种向日葵(Helianthus annuus)。作为食用作物和观赏植物，向日葵(Helianthus annuus)和其他物种，诸如洋姜(菊芋Helianthus tuberosus)被栽培在温和的地区。被驯化的向日葵Helianthus annuus是向日葵属(Helianthus L.)最常见的物种。栽培Helianthus annuus用于观赏，并且用于提供来自种子的植物油。
霜霉病，向日葵中的一种常见且具有毁灭性的疾病，能杀死植物或阻碍植物的正常发育，减少直立(stand)，并且导致显著的产量损失(高达50％～95％)。在种植之后2～3周内降了暴雨的大田里，向日葵最易受到霜霉菌的影响。
霜霉菌(downy mildew)指卵菌植物病原体的植物专性寄生物的几种类型中的任一种。霜霉菌专属于霜霉科(Peronosporaceae)。在栽培的向日葵中常引起霜霉病的霜霉菌病原体，被命名为向日葵霜霉病菌(Plasmopara halstedii)或向日葵轴霜霉(Plasmoparahelianthi)。
在向日葵栽培和育种的技术领域中，始终需要鉴定出抗霜霉菌的新的抗性基因。但是，鉴定出的抗性最大的基因是单基因显性抗性基因，而由这些基因提供的抗性常常很快就被破坏，因为霜霉菌病原体以高频率发生进化并且适应新的情况，从而恢复成功感染宿主植物的能力。因此，在本领域中需要持续寻找新的抗性基因，优选其抗性不易被病原体的适应性破坏的抗性基因。
除了提供病原体抗性之外，已知的向日葵抗性基因的缺点是，这些基因常常伴随不理想的表型，诸如矮化生长或自发出现的细胞死亡。因此，在本领域中需要持续寻找具有抗性而没有不理想表型的新的抗性基因。
发明内容及具体实施方式
除了其他目的，本发明的目的是至少部分(如果不是完全)满足本领域的上述需求。
除了其他目的，本发明的这一目的通过提供如在所附权利要求书中所提出的向日葵植物和抗性基因来满足。
具体地，除了其他目的，根据第一方面，通过对植物病原体霜霉菌具有抗性的向日葵植物来满足本发明的这一目的，其中，所述植物包括：编码(a)包括如SEQ ID No.2和/或SEQ ID No.4所示的氨基酸序列的蛋白的霜霉菌抗性赋予基因，或者编码与SEQ ID No.2和/或SEQ ID No.4本身具有超过90％序列同一性、优选超过94％序列同一性、更优选超过96％序列同一性的蛋白的霜霉菌抗性赋予基因；并且其中，与对植物病原体霜霉菌没有抗性的向日葵植物中的所述抗性赋予基因的表达相比，所述抗性赋予基因的表达是降低的，或者与对植物病原体霜霉菌没有抗性的向日葵植物中的所述蛋白的酶活性相比，所述蛋白的酶活性是降低的。
在得到本发明的研究中，令人惊奇地发现，所述基因的表达降低或所述蛋白的酶活性降低在向日葵植物中提供了对霜霉菌的广谱且持久的抗性。
根据本发明，与对植物病原体霜霉菌没有抗性的向日葵植物中的所述抗性赋予基因表达相比，表达是降低的。术语“没有抗性”指在适当的疾病测试中并且使用适当的参照植物(诸如亲本植物)确定的抗性水平低于在本发明植物中观察到的抗性水平。相应地，所述抗性也可被称为对霜霉菌的提高的抗性。除了亲本植物之外，根据本发明的合适的参照植物也可以是本领域中被普遍称为易受霜霉菌影响的植物。
合适的疾病测试是，使用霜霉菌病原体接种植物，随后观察疾病症状的出现，诸如在叶子的上表面或被破坏的叶组织上出现可见的大、有角或块状、黄色的区域。
可使用任何合适的且众所周知的分子生物学技术，诸如定量聚合酶链式反应(PCR)或mRNA杂交，来确定本发明的植物和参照植物中的表达水平
根据本发明，与对植物病原体霜霉菌没有抗性的向日葵植物中的本发明蛋白的活性相比，酶活性是降低的。术语“没有抗性”指在适当的疾病测试中并且使用适当的参照植物(诸如亲本植物)确定的抗性水平低于在本发明植物中观察到的抗性水平。相应地，所述抗性也可被称为对霜霉菌的提高的抗性。除了亲本植物之外，根据本发明的合适的参照植物也可以是本领域中被普遍称为易受霜霉菌影响的植物。合适的疾病测试是，使用霜霉菌病原体接种植物，随后观察疾病症状的出现，诸如在叶子的上表面或被破坏的叶组织上出现可见的大、有角或块状、黄色的区域。
本发明所述的蛋白具有2-酮戊二酸FE(II)依赖性氧合酶活性。所述酶绝对需要Fe(II)，并且催化双电子氧化，包括羟基化、脱饱和和氧化闭环反应。“最初”底物的氧化与2OG转化成琥珀酸盐和CO2相关。氧分子的一个氧并入到琥珀酸盐中。在脱饱和反应的情况中，其他氧分子来源的氧可能转化成水。在羟基化反应中，来自氧分子的氧部分并入醇产物中，同时观察到来自水的氧发生显著水平的交换。相应地，所述降低的活性可使用酶反应的起始化合物或生成化合物的试验测量化合物来测定。作为合适的选择，可通过例如ELISA或蛋白杂交(二者都是技术人员公知的技术)，来确定所述蛋白的蛋白水平(固有地指示降低的活性)
在本发明的上下文内，通过使所述向日葵植物的所述蛋白或编码所述蛋白的基因的表达或活性降低，以单独地或者组合地提供对霜霉菌的抗性。
可通过对易受霜霉菌影响的植物或霜霉科真菌抗性植物进行诱变，从而提高其抗性，来获得本发明的向日葵植物。例如，通过使用诸如甲基磺酸乙酯(EMS)的诱变化学品，或使用γ射线或快中子对植物材料进行辐照，可在这些植物中引入表达水平或蛋白水平上的突变。产生的突变可以是定向的或者是随机的。当产生随机突变时，则使用TILLING(定向诱导基因组局部突变)法(McCallum et al.(2000)Targeted screening for inducedmutations.Nat.Biotechnol.18,455-457,and Henikoff et al.(2004)TILLING.Traditional mutagenesis meets functional genomics.Plant Physiol.135,630-636)，可以很容易地鉴定出在所述抗性赋予基因中携带突变的诱变植物。简单来讲，该方法基于：在M2代中，对一大批诱变植物的基因组DNA的感兴趣基因进行PCR扩增。通过使用单链特异性核酸酶，诸如CEL-I核酸酶(Till et al.(2004)Mismatch cleavage bysingle-strand specific nucleases.Nucleic Acids Res.32,2632-2641)进行DNA测序或者扫描点突变，鉴定出在所述基因中具有突变的个体植物。
根据本发明的该第一方面的优选实施方式，所述霜霉菌病原体是向日葵霜霉病菌(Plasmopara halstedii)和/或向日葵轴霜霉(Plasmopara helianthi)。但是，属于霜霉科(Peronosporaceae)且能在向日葵中引起霜霉病的其他病原体也在本发明的范围内。
根据本发明的该第一方面的另一优选实施方式，通过在所述基因的编码序列中的产生截短或非功能性蛋白的一个或多个突变，来提供所述降低的酶活性。通过分析在mRNA或cDNA水平上的基因转录物，可以很容易地确定截短的蛋白，并且，在酶测定中或使用构象依赖性抗体，可以确定非功能性蛋白。可以在转录水平上测定的突变例如是氨基酸置换、移码突变或提取出现的终止密码子。
根据本发明的该第一方面的特别优选的实施方式，使所述蛋白的活性降低的突变是，使所述抗体提供基因的编码序列缺乏序列基元“WRDYLR”或Trp-Arg-Asp-Tyr-Leu-Arg，或在序列基元“WRDYLR”或Trp-Arg-Asp-Tyr-Leu-Arg中产生氨基酸置换的突变。所述序列基元位于SEQ ID No.2的氨基酸第107～112位和SEQ ID No.4的氨基酸第116～121位。发明人发现了，在这一区域中的突变尤其影响观察到的霜霉菌抗性表型，即抗性水平。涉及Y(Tyr)和/或R(Arg)的突变尤其与观察到的霜霉菌抗性表型(即抗性水平)高度相关。
根据本发明的该第一方面的又一优选实施方式，通过所述基因的调控区或非编码序列中的一个或多个突变，提供所述降低的表达。所述基因的调控区的实例是启动子区和终止子区，而非编码区的实例是内含子，尤其是其中影响剪接的基序。
根据第二方面，本发明提供了如上所述的向日葵植物的种子、植物组织或植物部分，或可从如上所述的向日葵植物获得的种子、植物组织或植物部分，包括：编码包括如SEQID No.2和/或SEQ ID NO.4所示的氨基酸序列的蛋白的霜霉菌抗性赋予基因，或者编码与SEQ ID No.2和/或SEQ ID NO.4本身具有超过90％序列同一性、优选超过94％序列同一性、更优选超过96％序列同一性的蛋白的霜霉菌抗性赋予基因；并且，与对植物病原体霜霉菌没有抗性的向日葵植物中的所述抗性赋予基因的表达相比，所述抗性赋予基因的表达是降低的，或者，与对植物病原体霜霉菌没有抗性的向日葵植物中的所述蛋白的酶活性相比，所述蛋白的酶活性是降低的。
根据第三方面，本发明涉及提供对植物病原体霜霉菌具有抗性的向日葵植物的方法，或者提高向日葵植物对植物病原体霜霉菌的抗性的方法，其中，所述方法包括以下步骤：在向日葵植物中引入编码包括如SEQ ID No.2和/或SEQ ID No.4所示的氨基酸序列的蛋白的霜霉菌抗性赋予基因，或者编码与SEQ ID No.2和/或SEQ ID No.4本身具有超过90％序列同一性、优选超过94％序列同一性、更优选超过96％序列同一性的蛋白的霜霉菌抗性赋予基因；并且，与初始向日葵植物中的所述抗性赋予基因的表达相比，所述抗性赋予基因的表达是降低的，或者，与对植物病原体霜霉菌没有抗性的初始向日葵植物中的所述蛋白的酶活性相比，所述蛋白的酶活性是降低的。
根据第四方面，本发明涉及编码包括如SEQ ID No.2或SEQ ID NO.4所示的氨基酸序列的蛋白的基因或其cDNA序列，或者编码与SEQ ID No.2或SEQ ID NO.4本身具有超过90％序列同一性、优选超过94％序列同一性、更有选超过96％序列同一性的蛋白的霜霉菌抗性赋予基因，在提供对植物病原体霜霉菌具有抗性或提高的抗性的向日葵植物中的应用。
根据第五方面，本发明涉及具有包括SEQ ID No.2或SEQ ID No.4的氨基酸序列的蛋白。
根据第六方面，本发明涉及包括SEQ ID No.1或SEQ ID No.3的核酸序列。
根据第七方面，本发明涉及编码具有包括SEQ ID No.2或SEQ ID No.4的氨基酸序列的蛋白的基因，或者包括SEQ ID No.1或SEQ ID No.3的核酸序列。
Claims (5)
1.一种提供对植物病原体霜霉菌具有抗性的向日葵植物的方法，所述方法包括以下步骤：在向日葵植物中引入编码由如SEQ ID No. 2或SEQ ID No. 4所示的氨基酸序列组成的蛋白的霜霉菌抗性赋予基因，并且其中，与对植物病原体霜霉菌没有抗性的向日葵植物中的蛋白的酶活性相比，所述蛋白的酶活性是降低的，其中，所述酶活性的降低是由所述抗性赋予基因所编码的序列的序列基元“WRDYLR”中第二个R置换为Q提供的。
2.编码由如SEQ ID No. 2或SEQ ID No. 4所示的氨基酸序列组成的蛋白的基因或其cDNA序列在提供对植物病原体霜霉菌具有抗性或具有提高的抗性的向日葵植物中的应用，其中所述抗性赋予基因所编码的序列的序列基元“WRDYLR”中第二个R置换为Q。
3.一种蛋白，其中，所述蛋白的氨基酸序列为经突变的SEQ ID No. 2或经突变的SEQID No. 4，所述突变是序列基元“WRDYLR”中的第二个R置换为Q。
4.一种由经突变的SEQ ID No. 1或经突变的SEQ ID No. 3组成的核酸序列，其中所述突变是所述核酸序列所编码的序列的序列基元“WRDYLR”中的第二个R置换为Q。
5.一种编码由经突变的SEQ ID No. 2或经突变的SEQ ID No. 4的氨基酸序列组成的蛋白的基因，其中所述突变是指所述氨基酸序列的序列基元“WRDYLR”的第二个R置换为Q。
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| RU2672335C2 (ru) | 2013-07-22 | 2018-11-13 | Шьенца Байотекнолоджис 5 Б.В. | Гены, обеспечивающие устойчивость подсолнечника к ложной мучнистой росе |
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