偃麦草属着丝粒序列组成分析及小麦—十倍体长穗偃麦草广谱抗锈易位系的鉴定
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摘要
偃麦草属植物是迄今在小麦育种中利用最成功的小麦近缘种属之一，为小麦品种改良提供了优异的基因资源。真核生物的着丝粒保证了细胞分裂中染色体准确分离。外源染色体在细胞中的去留与着丝粒有千丝万缕的关系。了解偃麦草属植物和小麦着丝粒组成的异同，将为小麦远缘杂交育种提供一些理论依据，提高远缘杂交育种的效率。本文对偃麦草着丝粒序列和小麦-十倍体长穗偃麦草杂交后代进行了研究，取得了以下主要结果：
1、构建了偃麦草属祖先种二倍体拟鹅冠草（Pseudoroegneiria stipifolia,2n=14, StSt）的BAC文库，根据禾本科着丝粒特异序列CCS1设计的引物筛选BAC克隆。结合荧光原位杂交，筛选到1个着丝粒特异的BAC克隆和2个着丝粒及近着丝粒区相关的BAC，构建了这3个BAC克隆的Shotgun文库，并完成了测序和组装。
2、发现Ps. stipifolia的着丝粒区包含与小麦着丝粒区特异的反转录转座子CRW、Quinta及unnamedfam6的同源序列，相似性均在85%以上；在Ps. stipifolia的着丝粒区，还发现一种尚未见报道的串联重复序列，基序长度约为500bp，暂命名为CentSt。在小麦中也存在CentSt的同源序列，串联重复单元长度为550bp，两者相似性约85%。
3、利用CRW、Quinta、unnamedfam6和CentSt分析四倍体拟鹅冠草（2n=28, StSt）、四倍体长穗偃麦草（2n=28, EeEe）、中间偃麦草（2n=42, StEbEe）和十倍体长穗偃麦草（2n=70, StStEbEeEx）的着丝粒组成，发现除了四倍体长穗偃麦草中没有检测到unnamedfam6的信号外，其它的三个物种中都含有这些元件，但是它们的分布规律不尽相同。说明在偃麦草多倍体化的过程中，着丝粒的序列组成也在发生着急剧的变化。
4、同源克隆了Ps. stipifolia、Th. elongatum（2n=14, EeEe）、Th. bessarabicum（2n=14, EbEb）、中间偃麦草和十倍体长穗偃麦草中着丝粒功能关键蛋白CENH3的编码基因，它们与小麦的CENH3基因相似性在95%以上。小麦的CENH3蛋白抗体能够结合在偃麦草染色体的着丝粒区。以上结果说明偃麦草属的着丝粒在DNA组成上和着丝粒关键蛋白水平上与小麦有很高的相似性，为偃麦草染色体在小麦背景下的稳定传递提供了可能，也为其与小麦染色体的重组提供了更多机会。
5、从普通小麦与十倍体长穗偃麦草的杂交、回交后代中，获得了对小麦条锈病流行混合小种具有广谱抗性的新种质。通过对其进行条锈病抗性鉴定、农艺性状调查、分子细胞学和品质相关性状分析发现：这批材料在苗期普遍表现为轻度感病，但成株后则完全高抗或免疫，对8份材料进行了连续三年的人工接种鉴定，抗性表现十分稳定，对多个致病小种及混合小种具有很好的抗性，这些材料根尖细胞染色体均为2n=42，农艺性状良好。原位杂交分析显示这些材料中含有同一个易位片段，即在4DS末端有一来自长穗偃麦草St基因组的小片段，其中一个品系（GDR3）还携带一个未能准确定位的小片段易位，该材料对混合小种的抗性优于其它材料。推测这两个小片段易位上均携带抗条锈病基因。高分子量麦谷蛋白分析表明，这批材料均含有优质亚基14+15。新的广谱抗性易位系的育成，将有可能在我国小麦抗锈育种中发挥重要作用。
总之，偃麦草属植物着丝粒在DNA组成和着丝粒关键蛋白CENH3水平上与小麦相似性很高，为偃麦草染色体在小麦背景下的稳定传递提供了可能，也为其与小麦染色体的重组提供了更多的机会。小麦与偃麦草染色体自发抗病易位系的发生和发现，从一个侧面证明了前一结论的可靠性，这些工作为未来的作物远缘杂交育种选材提供了新的视角，即物种间享有共同的着丝粒序列是外源染色体稳定传递的重要基础，也为自发易位和重组的发生创造了机会。
The Thinopyrum specics are the most successful relative species in wheat distanthybridization breeding, providing numerous useful genes in bread wheat improvement. Centromeresmaintain the integrity of the chromosomes during cell division. The deletion and preservation of alienchromosomes in cells related to the centromere. For understanding the centromere sequencecomposition of Thinopyrum and providing theoretical basis of distant hybridization breeding, in thisdissertation, three aspects of studies were performed surrounding the Thinopyrum’s centromere andwheat-Th.ponticum translocation lines.
1． A BAC library of Pseudoroegneiria stipifolia (2n=14, StSt) was constructed. The BAC librarywas screened via PCR by primers designed based on cereal centromeric sequence (CCS1). Afterfluorescent in situ hybridization (FISH) analysis, one centromere-specific BAC and two centromericassociated BACs were obtained. Shotgun libraries were constructed for the three BAC clones. Theywere sequenced, assembled and annotated.
2． We found that centromeres of Ps.stipifolia contained the homologous sequences of CRW,Quinta and unnamedfam6, which were found in wheat, and their similarity was more than85%. Acetromeric tandem repeat was found in Ps. stipifolia, which was named as CentSt. Homologoussequence of CentSt was also found in wheat and its D genome donor, Aegilops tauschii.
3． CRW, Quinta, unnamedfam6and CentSt were used to analysis the centromere of tetraploid Ps.stipifolia, tetraploid Th. elongatum (2n=28, EeEe), Th. intermedium (2n=42, StEbEe) and Th. ponticum(2n=70, StStEbEeEx). We found that except the tetraploid Th. elongatum which we didn’t detect thesignal of unnamedfam6in its chromosome, all of the other three species contained these centromericelements. But the distribution of the centromeric elements was different from each other. It wassuggested that in the progress of polyploidization inThinopyrum, the sequence compositon of thecentromeres changed a lot.
4． CENH3genes were cloned from Ps. stipifolia, Th. elongatum (2n=14, EeEe), Th.bessarabicum (2n=14, EbEb), Th. intermedium and Th. ponticum. The similarity of CENH3cordingsequences between wheat and genome donors of Thinopyrum genus was more than95%. And wheatCENH3antibody can perfectly attach on the centromere of the Thinopyrum chromosomes. All of theseresults suggested that the wheat and Thinopyrum have high homology centromere sequences andCENH3genes. This provides the base for Thinopyrum chromosomes stably being transmitted in wheatgenetic background, which creats chance for recombination.
4. To transfer stripe rust resistance from Th. ponticum to common wheat, hybrid progeniesbetween them were screened for stripe rust resistance.The progenies were characterized by their diseaseresistance, agronomic traits, genomic in situ hybridization and quality related protein analysis. Eightlines with good resistance or immune to the pathogens at adult stage were obtained. All of them have42chromosomes, in which seven lines convey a new translocation,4DS/4St. In addition, besides this translocation, the GDR3conveys another distal translocation, which may be derived from the E-genomechromosome of Th. ponticum. The GDR3performed very good resistance in whole life. It was immuneto yellow rusts. We estimated that both of the alien segments convey resistant genes to the stripe rusts.All of the eight lines convey the good subunits14+15at Glu-B1, which should be inherited fromXiaoyan6. Wild relatives of wheat usually convey genes with broad spectrum resistance to pathogens.This is the genetic base for their surviving in extreme environments, which also provide excellent generesource for disease resistant breeding in wheat.
In summary, the Triticum aestivum and Thinopyrum have simliar centromere sequences and CENH3genes. This provides the possibility that Thinopyrum chromosomes can stay and transmission stably inwheat genetic background, which creates chance for recombination between wheat and alienchromosomes. The occurrence of spontaneous translocations between wheat and Th. ponticumillustrates the reliability of ourspeculation. This work sheds light for chose of donor species in cropwide hybridization breeding.
1、构建了偃麦草属祖先种二倍体拟鹅冠草（Pseudoroegneiria stipifolia,2n=14, StSt）的BAC文库，根据禾本科着丝粒特异序列CCS1设计的引物筛选BAC克隆。结合荧光原位杂交，筛选到1个着丝粒特异的BAC克隆和2个着丝粒及近着丝粒区相关的BAC，构建了这3个BAC克隆的Shotgun文库，并完成了测序和组装。
2、发现Ps. stipifolia的着丝粒区包含与小麦着丝粒区特异的反转录转座子CRW、Quinta及unnamedfam6的同源序列，相似性均在85%以上；在Ps. stipifolia的着丝粒区，还发现一种尚未见报道的串联重复序列，基序长度约为500bp，暂命名为CentSt。在小麦中也存在CentSt的同源序列，串联重复单元长度为550bp，两者相似性约85%。
3、利用CRW、Quinta、unnamedfam6和CentSt分析四倍体拟鹅冠草（2n=28, StSt）、四倍体长穗偃麦草（2n=28, EeEe）、中间偃麦草（2n=42, StEbEe）和十倍体长穗偃麦草（2n=70, StStEbEeEx）的着丝粒组成，发现除了四倍体长穗偃麦草中没有检测到unnamedfam6的信号外，其它的三个物种中都含有这些元件，但是它们的分布规律不尽相同。说明在偃麦草多倍体化的过程中，着丝粒的序列组成也在发生着急剧的变化。
4、同源克隆了Ps. stipifolia、Th. elongatum（2n=14, EeEe）、Th. bessarabicum（2n=14, EbEb）、中间偃麦草和十倍体长穗偃麦草中着丝粒功能关键蛋白CENH3的编码基因，它们与小麦的CENH3基因相似性在95%以上。小麦的CENH3蛋白抗体能够结合在偃麦草染色体的着丝粒区。以上结果说明偃麦草属的着丝粒在DNA组成上和着丝粒关键蛋白水平上与小麦有很高的相似性，为偃麦草染色体在小麦背景下的稳定传递提供了可能，也为其与小麦染色体的重组提供了更多机会。
5、从普通小麦与十倍体长穗偃麦草的杂交、回交后代中，获得了对小麦条锈病流行混合小种具有广谱抗性的新种质。通过对其进行条锈病抗性鉴定、农艺性状调查、分子细胞学和品质相关性状分析发现：这批材料在苗期普遍表现为轻度感病，但成株后则完全高抗或免疫，对8份材料进行了连续三年的人工接种鉴定，抗性表现十分稳定，对多个致病小种及混合小种具有很好的抗性，这些材料根尖细胞染色体均为2n=42，农艺性状良好。原位杂交分析显示这些材料中含有同一个易位片段，即在4DS末端有一来自长穗偃麦草St基因组的小片段，其中一个品系（GDR3）还携带一个未能准确定位的小片段易位，该材料对混合小种的抗性优于其它材料。推测这两个小片段易位上均携带抗条锈病基因。高分子量麦谷蛋白分析表明，这批材料均含有优质亚基14+15。新的广谱抗性易位系的育成，将有可能在我国小麦抗锈育种中发挥重要作用。
总之，偃麦草属植物着丝粒在DNA组成和着丝粒关键蛋白CENH3水平上与小麦相似性很高，为偃麦草染色体在小麦背景下的稳定传递提供了可能，也为其与小麦染色体的重组提供了更多的机会。小麦与偃麦草染色体自发抗病易位系的发生和发现，从一个侧面证明了前一结论的可靠性，这些工作为未来的作物远缘杂交育种选材提供了新的视角，即物种间享有共同的着丝粒序列是外源染色体稳定传递的重要基础，也为自发易位和重组的发生创造了机会。
The Thinopyrum specics are the most successful relative species in wheat distanthybridization breeding, providing numerous useful genes in bread wheat improvement. Centromeresmaintain the integrity of the chromosomes during cell division. The deletion and preservation of alienchromosomes in cells related to the centromere. For understanding the centromere sequencecomposition of Thinopyrum and providing theoretical basis of distant hybridization breeding, in thisdissertation, three aspects of studies were performed surrounding the Thinopyrum’s centromere andwheat-Th.ponticum translocation lines.
1． A BAC library of Pseudoroegneiria stipifolia (2n=14, StSt) was constructed. The BAC librarywas screened via PCR by primers designed based on cereal centromeric sequence (CCS1). Afterfluorescent in situ hybridization (FISH) analysis, one centromere-specific BAC and two centromericassociated BACs were obtained. Shotgun libraries were constructed for the three BAC clones. Theywere sequenced, assembled and annotated.
2． We found that centromeres of Ps.stipifolia contained the homologous sequences of CRW,Quinta and unnamedfam6, which were found in wheat, and their similarity was more than85%. Acetromeric tandem repeat was found in Ps. stipifolia, which was named as CentSt. Homologoussequence of CentSt was also found in wheat and its D genome donor, Aegilops tauschii.
3． CRW, Quinta, unnamedfam6and CentSt were used to analysis the centromere of tetraploid Ps.stipifolia, tetraploid Th. elongatum (2n=28, EeEe), Th. intermedium (2n=42, StEbEe) and Th. ponticum(2n=70, StStEbEeEx). We found that except the tetraploid Th. elongatum which we didn’t detect thesignal of unnamedfam6in its chromosome, all of the other three species contained these centromericelements. But the distribution of the centromeric elements was different from each other. It wassuggested that in the progress of polyploidization inThinopyrum, the sequence compositon of thecentromeres changed a lot.
4． CENH3genes were cloned from Ps. stipifolia, Th. elongatum (2n=14, EeEe), Th.bessarabicum (2n=14, EbEb), Th. intermedium and Th. ponticum. The similarity of CENH3cordingsequences between wheat and genome donors of Thinopyrum genus was more than95%. And wheatCENH3antibody can perfectly attach on the centromere of the Thinopyrum chromosomes. All of theseresults suggested that the wheat and Thinopyrum have high homology centromere sequences andCENH3genes. This provides the base for Thinopyrum chromosomes stably being transmitted in wheatgenetic background, which creats chance for recombination.
4. To transfer stripe rust resistance from Th. ponticum to common wheat, hybrid progeniesbetween them were screened for stripe rust resistance.The progenies were characterized by their diseaseresistance, agronomic traits, genomic in situ hybridization and quality related protein analysis. Eightlines with good resistance or immune to the pathogens at adult stage were obtained. All of them have42chromosomes, in which seven lines convey a new translocation,4DS/4St. In addition, besides this translocation, the GDR3conveys another distal translocation, which may be derived from the E-genomechromosome of Th. ponticum. The GDR3performed very good resistance in whole life. It was immuneto yellow rusts. We estimated that both of the alien segments convey resistant genes to the stripe rusts.All of the eight lines convey the good subunits14+15at Glu-B1, which should be inherited fromXiaoyan6. Wild relatives of wheat usually convey genes with broad spectrum resistance to pathogens.This is the genetic base for their surviving in extreme environments, which also provide excellent generesource for disease resistant breeding in wheat.
In summary, the Triticum aestivum and Thinopyrum have simliar centromere sequences and CENH3genes. This provides the possibility that Thinopyrum chromosomes can stay and transmission stably inwheat genetic background, which creates chance for recombination between wheat and alienchromosomes. The occurrence of spontaneous translocations between wheat and Th. ponticumillustrates the reliability of ourspeculation. This work sheds light for chose of donor species in cropwide hybridization breeding.
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