簇毛麦染色体分子核型及染色体结构变异体库的构建
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摘要
簇毛麦(Haynaldia villosa L.)是小麦的一个野生近缘种,抗白粉病、锈病、全蚀病、眼斑病及黄花叶病等多种病害,同时还兼有抗寒耐旱、分蘖力强、密穗多花、籽粒蛋白质含量高等特性,是小麦遗传改良的优良基因源。为了定位、转移和利用簇毛麦的有益基因,本研究以硬粒小麦-簇毛麦双倍体花粉辐射的回交自交后代为材料,首先利用基因组原位杂交筛选小麦背景中只存在一种形式的簇毛麦染色体结构变异体,而后利用簇毛麦各条染色体的特异分子标记对这些结构变异体的身份进行鉴定,以此构建“普通小麦-簇毛麦染色体结构变异体库”
第一部分：普通小麦-簇毛麦二体异附加系的选育与鉴定
目前国际上有2套完整的簇毛麦二体异附加系,其簇毛麦来源与本研究所用的簇毛麦不同,农艺性状或优异特性也具有一定的差异。本研究在(中国春/硬粒小麦-簇毛麦双倍体(60Co-γ射线照射花粉)//中国春)的杂交回交后代中,应用基因组原位杂交和分子标记分析技术,从BCiF2和BC4F2代中选育并鉴定出分别涉及簇毛麦7条染色体的纯合二体附加系,花粉母细胞减数分裂中期Ⅰ染色体构型观察表明添加的一对簇毛麦染色体配对成环状或棒状二价体。利用选育鉴定的一套普通小麦-簇毛麦二体异附加系可以定位簇毛麦的优异基因。
第二部分：簇毛麦染色体分子核型的建立
对导入普通小麦背景中的簇毛麦染色体或染色体片段的准确鉴定是利用簇毛麦优异基因的前提。本研究旨在利用2个重复序列(pSc119.2、pAs1)和2个rDNA多基因家族(45S rDNA、5S rDNA)作为探针建立二倍体簇毛麦(VV,2n=14)染色体的分子核型,以用于鉴定小麦背景中的簇毛麦染色体或染色体片段。荧光原位杂交结果显示45S rDNA和5S rDNA分别只在簇毛麦1VS和5VS上有杂交信号,可分别作为1VS和5VS的细胞学标记；pSc119.2在簇毛麦7条染色体上都有杂交信号,信号分布于染色体的末端或亚末端,但有2对染色体只在短臂上有杂交信号,其信号的分布可区分簇毛麦全部7条染色体；pAs1在簇毛麦7条染色体上也都有杂交信号,信号较为丰富,分布于染色体的末端、亚末端、中部和着丝粒区域。利用2个重复序列pSc119.2、pAs1和2个rDNA多基因家族45S rDNA、5S rDNA对一套簇毛麦二体附加系进行荧光原位杂交分析,建立了由4种重复序列作探针的簇毛麦染色体分子核型。
第三部分：簇毛麦染色体结构变异体库的构建
别同德和曹亚萍先后利用60Co-γ射线照射硬粒小麦-簇毛麦双倍体即将成熟的花粉,然后授给已去雄的普通小麦中国春,在M1代即用基因组原位杂交检测到大量的簇毛麦染色体结构变异,并且大多数结构变异体能在后代中稳定传递。为获得尽可能多的结构变异体,我们首先利用基因组原位杂交筛选在小麦背景中只含有一种类型簇毛麦染色体结构变异的变异体,目前已筛选到140份结构变异体,其中小片段易位41份、大片段易位32份、整臂易位44份、缺失18份和簇毛麦插入型易位5份；随后在这些结构变异体的自交后代中,鉴定出了30份纯合体,包括小片段易位7份、大片段易位9份、整臂易位10份、缺失2份及簇毛麦插入型易位2份。
为了更有效地鉴定这些簇毛麦染色体结构变异体,我们对簇毛麦的特异标记进行了加密,在选取的1576对引物中,共筛选到89个簇毛麦染色体特异标记,其中1V特异标记有2个、2V特异标记有14个、3V特异标记有8个、4V特异标记有8个、5V特异标记有41个、6V特异标记有2个及7V特异标记有14个。由于本实验室缺少涉及3V和7V的端体或整臂易位材料,我们利用pSc119.2和pAs1对2个端体和2个整臂易位材料进行双色荧光原位杂交分析,再与标准的簇毛麦染色体分子核型比较,结果表明单株TV60-1-8-27和TV60-1-8-18分别是3V的短臂和长臂的端体,单株TV54-5-5-22和TV54-5-5-21分别是7V短臂和长臂的整臂易位,利用这几个非整倍体材料我们将3V和7V染色体的特异标记定位到相应的染色体臂上,其中8个标记被定位到3VS上、1个标记被定位到3VL上；另外11个标记被定位到7VS上、7个标记被定位到7VL上。
最后利用现有的分布于簇毛麦各条染色体上的124个特异标记,对这些结构变异体所涉及的簇毛麦的身份及变异区段进行了鉴定。目前,我们明确了67个结构变异体所涉及的簇毛麦染色体身份及变异区段,其中涉及1V的有8个、2V的有11个、3V的有5个、4V的有10个、5V的有9个、6V的有13个及7V的有11个,这些结构变异体涉及了簇毛麦不同的染色体区段,在此基础上初步构建了“普通小麦-簇毛麦染色体结构变异体库”。另外我们还将簇毛麦的特异标记定位到染色体的不同区段上；利用该库中涉及2VS的结构变异体结合表型观察,我们将簇毛麦的控制颖脊刚毛基因定位于2VS FL：0.00-0.33处。
Haynaldia villosa (L.) Schur (syn. Dasypyrum villosum (L.) Candargy) is a wild relative of common wheat(Triticum aestivum L.). It possesses many important agronomic traits, such as resistance to powdery mildew, leaf and stem rusts, take-all, eyespot, and wheat streak mosaic virus, as well as vigorous tillering ability, more spikelets, high grain protein content, and drought and frost tolerance. Therefore, it is a potential gene resource for wheat genetic improvement. To localize, transfer and utilize interested genes of H. villosa, Firstly, we screened common wheat lines carrying structurally changed chromosomes containing single segments of H. villosa by genomic in situ hybridization. Secondly, the specific molecular markers of individual chromosomes of H. villosa were used to identify these structurally changed chromosomes and construct "Triticum aestivum-Haynaldia villosa structural aberrance library"
Section I Development and identification of T. aestivum-H. villosa disomic addition lines
Now two sets of T. aestivum-H. villosa disomic addition lines were developed in the world, where from is different with H. villosa of our institute, and difference in agronomic or unique useful traits. Genomic in situ hybridization combined with molecular marker analysis were applied to detect chromosome of H. villosa in backcrossed and selfcrossed generations derived from (Chinese Spring/T. durum-H. villosa amphiploid (irradiated by60Co-γ rays)//Chinese Spring) in this study. A set of disomic addition lines involved in seven H. villosa chromosomes have been identified in BC3F2and BC4F2. The chromosome pairing behavior of H. villosa chromosomes at metaphase I (MI) of pollen mother cells (PMCs) were analyzed through using genomic in situ hybridization, one pair of H. villosa chromosomes paired ring or rod bivalent. Development and identification a set of T. aestivum-H. villosa disomic addition lines were used to localized useful genes of H, villosa.
Section II Construction of molecular karyotyping of H, villosa chromosomes
Identification the chromosomes or chromosome segments of H. villosa in common wheat background is first step to use the interested genes. The aim of the present work was to use two repetitive DNA sequences pSc119.2and pAs1, and two multigene families45S rDNA and5S rDNA as probes to construct molecular karyotyping of diploid H. villosa chromosomes (VV,2n=14), and identified the chromosomes or chromosome segments of H. villosa in wheat background. The results of fluorescence in situ hybridization showed the hybridization sites of the45S rDNA and5S rDNA were localized on1VS and5VS, respectively, and were cytogenetic marker for1VS and5VS. The pSc119.2as probe had a simpler distribution, whose signals were hybridized mainly on terminal or subterminal positions of both arms with the exception of the hybridization point on the short arm of two pair chromosomes. The signals generated by the pScl19.2probe were different on position and signals intensity. Meanwhile, they were uniquely identified. Furthermore, we used the pScl19.2in combination with pAsl as probes labeled with different fluorochromes to hybridize for H. villosa chromosomes. As a result, the pAsl as probe was more widely dispersed than pSc119.2over the H. villosa genome. We found pAs1hybridization sites on all of the chromosomes of H. villosa and practically on all of the chromosome arms. These hybridization points appear in terminal, subterminal, interstitial, and occasionally centromeric positions. The individual chromosomes which exhibited the fluorescence in situ hybridization signals were identified when using the pSc119.2and45S rDNA as probes to a set of T. aestivum-H. villosa disomic addition lines. Combining the genomic in situ hybridization and fluorescence in situ hybridization techniques allowed for all seven chromosomes of H. villosa for identification and constructed molecular karyotyping of H. villosa chromosomes.
Section Ⅲ Construction of aberrance library of H. villosa chromosomes
To induce as many as T. aestivum-H. villosa intergeneric translocation chromosomes involved in different chromosomes and chromosome segments of H. villosa, T. durum-H. villosa amphiploid was irradiated with60Co-γ rays, and pollens collected from the spikes after irradiation was pollinated to emasculated common wheat cv. Chinese Spring by Bie Tongde and Cao Yaping. Mass of H. villosa chromosomal aberrances were detected using genomic in situ hybridization technique in the M1and many of them could transmit to offspring. Firstly, we screened common wheat lines carrying structurally changed chromosomes containing single segment of H. villosa by genomic in situ hybridization. Up to date,140chromosomal structural changes were screened,41of small alien segment translocation,32of large alien segment translocation,44of whole arm translocation,18of chromosomal deletion and5of intercalary translocation of H. villosa, respectively.30homozygous plants were screened in these chromosomal structural changes,7of small alien segment translocation,9of large alien segment translocation,10of whole arm translocation,2of chromosomal deletion and2of intercalary of H, villosa, respectively.
In order to better identify these structural chromosomal changes, we screened1576primer pairs and89of them could be used for tracing individual chromosomes of H. villosa,2for1V,14for2V,8for3V,8for4V,41for5V,2for6V and14for7V, respectively. These90specific molecular markers add to the existing markers earlier screened and developed in H. villosa. The telosomic chromosome or whole arm translocation of chromosome3V and7V were absent in our laboratory. Two telosomic chromosomes and two whole arm translocations were hybridized with pSc119.2and pAsl as probes. The results showed that the plant of TV60-1-8-27and TV60-1-8-18was telosomic chromosome of3VS and3VL, respectively. Meanwhile, the plant of TV54-5-5-22and TV54-5-5-21was7VS·W and W-7VL, respectively. Furthermore, the specific molecular markers of chromosome3V and7V were localized on corresponding chromosomal arm by using these aneuploid,8on3VS,1on3VL,11on7VS and7on7VL, respectively.
Finally, these structural changes involving different chromosome and region of H. villosa were identified with124specific markers to individual chromosome. Now,67chromosomal structural changes were identified,8involving1V,11involving2V,5involving3V,10involving4V,9involving5V,13involving6V and11involving7V, respectively, and these aberrances involved different chromosomal region of H, villosa. We constructed "T. aestivum-H. villosa chromosomal aberrance library" based on above studies and the specific markers of H. villosa were localized on different chromosomal regions. The gene controlling bristles on the glume ridges was further located on2VS (FL:0.00-0.33) through observed the phenotype of involving chromosomal structural changes of2VS.
第一部分：普通小麦-簇毛麦二体异附加系的选育与鉴定
目前国际上有2套完整的簇毛麦二体异附加系,其簇毛麦来源与本研究所用的簇毛麦不同,农艺性状或优异特性也具有一定的差异。本研究在(中国春/硬粒小麦-簇毛麦双倍体(60Co-γ射线照射花粉)//中国春)的杂交回交后代中,应用基因组原位杂交和分子标记分析技术,从BCiF2和BC4F2代中选育并鉴定出分别涉及簇毛麦7条染色体的纯合二体附加系,花粉母细胞减数分裂中期Ⅰ染色体构型观察表明添加的一对簇毛麦染色体配对成环状或棒状二价体。利用选育鉴定的一套普通小麦-簇毛麦二体异附加系可以定位簇毛麦的优异基因。
第二部分：簇毛麦染色体分子核型的建立
对导入普通小麦背景中的簇毛麦染色体或染色体片段的准确鉴定是利用簇毛麦优异基因的前提。本研究旨在利用2个重复序列(pSc119.2、pAs1)和2个rDNA多基因家族(45S rDNA、5S rDNA)作为探针建立二倍体簇毛麦(VV,2n=14)染色体的分子核型,以用于鉴定小麦背景中的簇毛麦染色体或染色体片段。荧光原位杂交结果显示45S rDNA和5S rDNA分别只在簇毛麦1VS和5VS上有杂交信号,可分别作为1VS和5VS的细胞学标记；pSc119.2在簇毛麦7条染色体上都有杂交信号,信号分布于染色体的末端或亚末端,但有2对染色体只在短臂上有杂交信号,其信号的分布可区分簇毛麦全部7条染色体；pAs1在簇毛麦7条染色体上也都有杂交信号,信号较为丰富,分布于染色体的末端、亚末端、中部和着丝粒区域。利用2个重复序列pSc119.2、pAs1和2个rDNA多基因家族45S rDNA、5S rDNA对一套簇毛麦二体附加系进行荧光原位杂交分析,建立了由4种重复序列作探针的簇毛麦染色体分子核型。
第三部分：簇毛麦染色体结构变异体库的构建
别同德和曹亚萍先后利用60Co-γ射线照射硬粒小麦-簇毛麦双倍体即将成熟的花粉,然后授给已去雄的普通小麦中国春,在M1代即用基因组原位杂交检测到大量的簇毛麦染色体结构变异,并且大多数结构变异体能在后代中稳定传递。为获得尽可能多的结构变异体,我们首先利用基因组原位杂交筛选在小麦背景中只含有一种类型簇毛麦染色体结构变异的变异体,目前已筛选到140份结构变异体,其中小片段易位41份、大片段易位32份、整臂易位44份、缺失18份和簇毛麦插入型易位5份；随后在这些结构变异体的自交后代中,鉴定出了30份纯合体,包括小片段易位7份、大片段易位9份、整臂易位10份、缺失2份及簇毛麦插入型易位2份。
为了更有效地鉴定这些簇毛麦染色体结构变异体,我们对簇毛麦的特异标记进行了加密,在选取的1576对引物中,共筛选到89个簇毛麦染色体特异标记,其中1V特异标记有2个、2V特异标记有14个、3V特异标记有8个、4V特异标记有8个、5V特异标记有41个、6V特异标记有2个及7V特异标记有14个。由于本实验室缺少涉及3V和7V的端体或整臂易位材料,我们利用pSc119.2和pAs1对2个端体和2个整臂易位材料进行双色荧光原位杂交分析,再与标准的簇毛麦染色体分子核型比较,结果表明单株TV60-1-8-27和TV60-1-8-18分别是3V的短臂和长臂的端体,单株TV54-5-5-22和TV54-5-5-21分别是7V短臂和长臂的整臂易位,利用这几个非整倍体材料我们将3V和7V染色体的特异标记定位到相应的染色体臂上,其中8个标记被定位到3VS上、1个标记被定位到3VL上；另外11个标记被定位到7VS上、7个标记被定位到7VL上。
最后利用现有的分布于簇毛麦各条染色体上的124个特异标记,对这些结构变异体所涉及的簇毛麦的身份及变异区段进行了鉴定。目前,我们明确了67个结构变异体所涉及的簇毛麦染色体身份及变异区段,其中涉及1V的有8个、2V的有11个、3V的有5个、4V的有10个、5V的有9个、6V的有13个及7V的有11个,这些结构变异体涉及了簇毛麦不同的染色体区段,在此基础上初步构建了“普通小麦-簇毛麦染色体结构变异体库”。另外我们还将簇毛麦的特异标记定位到染色体的不同区段上；利用该库中涉及2VS的结构变异体结合表型观察,我们将簇毛麦的控制颖脊刚毛基因定位于2VS FL：0.00-0.33处。
Haynaldia villosa (L.) Schur (syn. Dasypyrum villosum (L.) Candargy) is a wild relative of common wheat(Triticum aestivum L.). It possesses many important agronomic traits, such as resistance to powdery mildew, leaf and stem rusts, take-all, eyespot, and wheat streak mosaic virus, as well as vigorous tillering ability, more spikelets, high grain protein content, and drought and frost tolerance. Therefore, it is a potential gene resource for wheat genetic improvement. To localize, transfer and utilize interested genes of H. villosa, Firstly, we screened common wheat lines carrying structurally changed chromosomes containing single segments of H. villosa by genomic in situ hybridization. Secondly, the specific molecular markers of individual chromosomes of H. villosa were used to identify these structurally changed chromosomes and construct "Triticum aestivum-Haynaldia villosa structural aberrance library"
Section I Development and identification of T. aestivum-H. villosa disomic addition lines
Now two sets of T. aestivum-H. villosa disomic addition lines were developed in the world, where from is different with H. villosa of our institute, and difference in agronomic or unique useful traits. Genomic in situ hybridization combined with molecular marker analysis were applied to detect chromosome of H. villosa in backcrossed and selfcrossed generations derived from (Chinese Spring/T. durum-H. villosa amphiploid (irradiated by60Co-γ rays)//Chinese Spring) in this study. A set of disomic addition lines involved in seven H. villosa chromosomes have been identified in BC3F2and BC4F2. The chromosome pairing behavior of H. villosa chromosomes at metaphase I (MI) of pollen mother cells (PMCs) were analyzed through using genomic in situ hybridization, one pair of H. villosa chromosomes paired ring or rod bivalent. Development and identification a set of T. aestivum-H. villosa disomic addition lines were used to localized useful genes of H, villosa.
Section II Construction of molecular karyotyping of H, villosa chromosomes
Identification the chromosomes or chromosome segments of H. villosa in common wheat background is first step to use the interested genes. The aim of the present work was to use two repetitive DNA sequences pSc119.2and pAs1, and two multigene families45S rDNA and5S rDNA as probes to construct molecular karyotyping of diploid H. villosa chromosomes (VV,2n=14), and identified the chromosomes or chromosome segments of H. villosa in wheat background. The results of fluorescence in situ hybridization showed the hybridization sites of the45S rDNA and5S rDNA were localized on1VS and5VS, respectively, and were cytogenetic marker for1VS and5VS. The pSc119.2as probe had a simpler distribution, whose signals were hybridized mainly on terminal or subterminal positions of both arms with the exception of the hybridization point on the short arm of two pair chromosomes. The signals generated by the pScl19.2probe were different on position and signals intensity. Meanwhile, they were uniquely identified. Furthermore, we used the pScl19.2in combination with pAsl as probes labeled with different fluorochromes to hybridize for H. villosa chromosomes. As a result, the pAsl as probe was more widely dispersed than pSc119.2over the H. villosa genome. We found pAs1hybridization sites on all of the chromosomes of H. villosa and practically on all of the chromosome arms. These hybridization points appear in terminal, subterminal, interstitial, and occasionally centromeric positions. The individual chromosomes which exhibited the fluorescence in situ hybridization signals were identified when using the pSc119.2and45S rDNA as probes to a set of T. aestivum-H. villosa disomic addition lines. Combining the genomic in situ hybridization and fluorescence in situ hybridization techniques allowed for all seven chromosomes of H. villosa for identification and constructed molecular karyotyping of H. villosa chromosomes.
Section Ⅲ Construction of aberrance library of H. villosa chromosomes
To induce as many as T. aestivum-H. villosa intergeneric translocation chromosomes involved in different chromosomes and chromosome segments of H. villosa, T. durum-H. villosa amphiploid was irradiated with60Co-γ rays, and pollens collected from the spikes after irradiation was pollinated to emasculated common wheat cv. Chinese Spring by Bie Tongde and Cao Yaping. Mass of H. villosa chromosomal aberrances were detected using genomic in situ hybridization technique in the M1and many of them could transmit to offspring. Firstly, we screened common wheat lines carrying structurally changed chromosomes containing single segment of H. villosa by genomic in situ hybridization. Up to date,140chromosomal structural changes were screened,41of small alien segment translocation,32of large alien segment translocation,44of whole arm translocation,18of chromosomal deletion and5of intercalary translocation of H. villosa, respectively.30homozygous plants were screened in these chromosomal structural changes,7of small alien segment translocation,9of large alien segment translocation,10of whole arm translocation,2of chromosomal deletion and2of intercalary of H, villosa, respectively.
In order to better identify these structural chromosomal changes, we screened1576primer pairs and89of them could be used for tracing individual chromosomes of H. villosa,2for1V,14for2V,8for3V,8for4V,41for5V,2for6V and14for7V, respectively. These90specific molecular markers add to the existing markers earlier screened and developed in H. villosa. The telosomic chromosome or whole arm translocation of chromosome3V and7V were absent in our laboratory. Two telosomic chromosomes and two whole arm translocations were hybridized with pSc119.2and pAsl as probes. The results showed that the plant of TV60-1-8-27and TV60-1-8-18was telosomic chromosome of3VS and3VL, respectively. Meanwhile, the plant of TV54-5-5-22and TV54-5-5-21was7VS·W and W-7VL, respectively. Furthermore, the specific molecular markers of chromosome3V and7V were localized on corresponding chromosomal arm by using these aneuploid,8on3VS,1on3VL,11on7VS and7on7VL, respectively.
Finally, these structural changes involving different chromosome and region of H. villosa were identified with124specific markers to individual chromosome. Now,67chromosomal structural changes were identified,8involving1V,11involving2V,5involving3V,10involving4V,9involving5V,13involving6V and11involving7V, respectively, and these aberrances involved different chromosomal region of H, villosa. We constructed "T. aestivum-H. villosa chromosomal aberrance library" based on above studies and the specific markers of H. villosa were localized on different chromosomal regions. The gene controlling bristles on the glume ridges was further located on2VS (FL:0.00-0.33) through observed the phenotype of involving chromosomal structural changes of2VS.
引文
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