簇毛麦6V染色体短臂小片段易位的高效诱导和鉴定
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摘要
由于病虫害、逆境等不利因素对小麦品质和产量的危害不断加重,提高对病虫和逆境的抗性已成为目前小麦育种工作的主要目标。簇毛麦具有抗多种病虫害,抗寒、耐旱等优良性状,是小麦种质改良的重要遗传资源。培育小麦—簇毛麦属间染色体小片段易位,特别是中间插入易位,有助于更好地利用簇毛麦有益基因改良小麦种质。但迄今很少有成功选育小麦—簇毛麦小片段易位、特别是中间插入易位的报道。因此,进一步研究小麦—簇毛麦属间染色体小片段易位的诱导和检测方法对于利用外源优良基因改良小麦种质具有重要意义。
本研究以~(60)Co-γ射线(剂量率:160Rad/min,三种剂量:1600、1920、2240Rad)处理6VS/6AL整臂易位系的成熟雌配子,随后选取适龄穗子去雄套袋,2-3天后再用普通小麦品种“中国春”的新鲜成熟花粉授粉。通过M_1植株根尖细胞有丝分裂中期染色体基因组原位杂交(Genomic In Situ Hybridization,GISH),从534株M_1材料中检测到97株涉及6V染色体短臂(6VS)的小片段结构变异,包括小片段中间插入易位染色体80条、末端易位染色体57条和6VS缺失55条。在2240Rad处理中这三种结构变异的诱变频率分别为21.02%、14.01%和14.65%,远远高于已报道的结果。获得了涉及146条6VS小片段结构变异的74株M_1材料的回交种子。M_1植株中的小片段结构变异可通过回交传递给后代。利用电离辐射处理整臂易位系成熟雌配子是一种高效诱导染色体小片段结构变异、特别是中间插入易位的新方法。本研究获得的6VS小片段涉及不同的片段长度和断裂位点,是精细定位Pm21基因和构建6VS物理图谱的重要材料。
根据定位于小麦第六部分同源群染色体短臂的EST序列,利用PRIMER5.0软件设计PCR引物,成功开发出3个对6VS特异的分子标记(6BS28-386、6DS38-730、6AS6-740)。利用小麦、簇毛麦第六部分同源群染色体短臂的12个缺失系,对簇毛麦6VS的8个特异性分子标记进行了定位。其中标记6VS19-381、CINAU17-1086分别定位在FL值为0.58-0.99、FL值为0.58-1.00的染色体区段,标记CINAU15-902定位在FL值为0.45-0.58的染色体区段,标记CINAU16-1650、6BS28-386、6DS38-730被定位在FL值为0.35-0.45的染色体区段。标记CINAU18-723、6AS6-740被定位在FL值为0.00-0.45的染色体区段。这些特异性分子标记可用于6VS小片段结构变异鉴定、6VS精细作图等研究。
以电离辐射处理获得的M_1变异植株为母本、以普通小麦作父本回交,在M_2中成功筛选到8株只具有单条6VS片段的中间插入易位。利用定位在染色体不同区段的8个6VS特异性分子标记,对8条中间插入的6VS片段进行了鉴定,其中NJ9-30具有的6VS片段为FL值为0.00-0.58的染色体区段,NJ2-2、NJ2-3具有的6VS片段为FL值为0.35-0.58的染色体区段,NJ6-23等材料中的6VS片段为FL值为0.00-0.35的染色体区段。
本研究首次选育出2株高抗小麦白粉病的杂合小片段中间插入易位,可望进一步获得携带Pm21基因的纯合中间插入易位系。利用所获得的6VS结构变异体并结合白粉病抗性鉴定,将Pm21基因定位在FL值为0.35-0.58的染色体区段。
Wheat production and quality are affected by biotic and abiotic stresses sereiously,and the improvement of the resistance to diseases and pests,and the adaptability to various environments are the main objectives of modern wheat breeding.H.villosa(syn. Dasypyrum villosum L.,2n=14,VV) have been proved to be highly resistant to wheat diseases and pests,tolerant to coldness and dryness,and is considered as an important genetic resource for wheat improvement.The translocation lines with small alien fragment, especially the interstitial translocation lines are best for use of alien useful genes in wheat breeding.However,up to now,few Triticum aestivum-H.villosa translocation lines with small alien segment have been reported.Therefore,it is necessary to establish a high efficient strategy for the induction and identification of alien translocation lines.
In the present study,the mature female gametes of wheat-H.villosa 6VS/6AL translocation line were irradiated by ~(60)Co-γray with 160Rad/M dosage rate and three dosages(1600,1920,2240Rad) before flowering.These irradiated florets were emasculated the same day,and then covered with paper bags.After 2-3 days,these florets were pollinated with normal fresh pollens from T.aestivum cv.Chinese Spring.Genomic in situ hybridization(GISH) at mitosis metaphase of root-tip cell of M_1 plants was used to detect the chromosomes structural changes involving 6VS of H.villosa.Among the 534 M_1 plants screened,97 plants contained chromosome structural changes with small segment of 6VS,including 80 interstitial translocated chromosomes,57 terminal translocated chromosomes and 55 deleted chromosomes.For the 2240 Rad dosage treatment,the induction frequencies of interstitial translocation,terminal translocation and deletion were 21.02%,14.01%,and 14.65%respectively,which were much higher than that previously reported.The M_2 seeds were obtained by backcrossing of 74 M_1 plants involving 146 chromosomes with structural changes of 6VS,and it was found that the structure aberrations in the M_1 plants could be transmitted to their progenies.Irradiating mature female gametes of whole arm translocation may be a new and highly efficient approach for creation of small segment chromosome structural changes,especially for interstitial translocations.These small fragments of 6VS obtained in the present research are different in both length of fragment and breakage location,and are useful for fine location of Pm21 gene,physical mapping of 6VS and development of translocation lines with small fragment.
Using a software PRIMER5.0PCR,primers were designed according to the sequences of ESTs located in the different bins of the short arm of homeologous group 6 chromosomes of wheat.Three molecular markers specific for 6VS(6BS28-386,6DS38-730,6AS6-740) were developed.Twelve deletion lines involved in the short arm of homeologous group 6 chromosomes of wheat and H.villosa were used to physically locate 8 specific markers of 6VS.The markers CINAU17-1086,6VS19-381 were located in the chromosome bin of FL (fragment length) value 0.58-1.00 and 0.58-0.99,the marker CINAU15-902 was located in the chromosome bin of FL value 0.45-0.58,the markers CINAU16-1650,6DS38-730, 6BS28-386 were located in the chromosome bin of FL value 0.35-0.45,and the markers CINAU18-723 and 6AS6-740 were located in the bin of FL value 0.00-0.45.These markers are usefull both for identification of structural changes with 6VS and fine mapping of Pm21.
Through the back-crosses between some M_1 plants with chromosome changes(female parent) and common wheat(male parent),8 M_2 plants with single small interstitial translocation chromosomes of 6VS have been obtained.The above 8 6VS specific molecular markers were used to define the fagment of 6VS.The fragment of 6VS in translocation line NJ9-30,NJ2-2(or NJ2-3) had FL value 0.00-0.58,0.35-0.58,respectively. The fragment of 6VS in the other lines(NJ6-23,NJ6-14 etc) was involved in the segment of FL value 0.00-0.35.
Two heterozygous insterstitial translocation lines with small segment of 6VS,which showed high resistance to powdery mildew,were selected,and homozygous interstitial translocation lines with Pm21 will be obtained.The Pm21 gene was further located in the region of 6VS with FL value 0.35-0.58 by using these genetic stocks of 6VS and powdery mildew resistance identification.
本研究以~(60)Co-γ射线(剂量率:160Rad/min,三种剂量:1600、1920、2240Rad)处理6VS/6AL整臂易位系的成熟雌配子,随后选取适龄穗子去雄套袋,2-3天后再用普通小麦品种“中国春”的新鲜成熟花粉授粉。通过M_1植株根尖细胞有丝分裂中期染色体基因组原位杂交(Genomic In Situ Hybridization,GISH),从534株M_1材料中检测到97株涉及6V染色体短臂(6VS)的小片段结构变异,包括小片段中间插入易位染色体80条、末端易位染色体57条和6VS缺失55条。在2240Rad处理中这三种结构变异的诱变频率分别为21.02%、14.01%和14.65%,远远高于已报道的结果。获得了涉及146条6VS小片段结构变异的74株M_1材料的回交种子。M_1植株中的小片段结构变异可通过回交传递给后代。利用电离辐射处理整臂易位系成熟雌配子是一种高效诱导染色体小片段结构变异、特别是中间插入易位的新方法。本研究获得的6VS小片段涉及不同的片段长度和断裂位点,是精细定位Pm21基因和构建6VS物理图谱的重要材料。
根据定位于小麦第六部分同源群染色体短臂的EST序列,利用PRIMER5.0软件设计PCR引物,成功开发出3个对6VS特异的分子标记(6BS28-386、6DS38-730、6AS6-740)。利用小麦、簇毛麦第六部分同源群染色体短臂的12个缺失系,对簇毛麦6VS的8个特异性分子标记进行了定位。其中标记6VS19-381、CINAU17-1086分别定位在FL值为0.58-0.99、FL值为0.58-1.00的染色体区段,标记CINAU15-902定位在FL值为0.45-0.58的染色体区段,标记CINAU16-1650、6BS28-386、6DS38-730被定位在FL值为0.35-0.45的染色体区段。标记CINAU18-723、6AS6-740被定位在FL值为0.00-0.45的染色体区段。这些特异性分子标记可用于6VS小片段结构变异鉴定、6VS精细作图等研究。
以电离辐射处理获得的M_1变异植株为母本、以普通小麦作父本回交,在M_2中成功筛选到8株只具有单条6VS片段的中间插入易位。利用定位在染色体不同区段的8个6VS特异性分子标记,对8条中间插入的6VS片段进行了鉴定,其中NJ9-30具有的6VS片段为FL值为0.00-0.58的染色体区段,NJ2-2、NJ2-3具有的6VS片段为FL值为0.35-0.58的染色体区段,NJ6-23等材料中的6VS片段为FL值为0.00-0.35的染色体区段。
本研究首次选育出2株高抗小麦白粉病的杂合小片段中间插入易位,可望进一步获得携带Pm21基因的纯合中间插入易位系。利用所获得的6VS结构变异体并结合白粉病抗性鉴定,将Pm21基因定位在FL值为0.35-0.58的染色体区段。
Wheat production and quality are affected by biotic and abiotic stresses sereiously,and the improvement of the resistance to diseases and pests,and the adaptability to various environments are the main objectives of modern wheat breeding.H.villosa(syn. Dasypyrum villosum L.,2n=14,VV) have been proved to be highly resistant to wheat diseases and pests,tolerant to coldness and dryness,and is considered as an important genetic resource for wheat improvement.The translocation lines with small alien fragment, especially the interstitial translocation lines are best for use of alien useful genes in wheat breeding.However,up to now,few Triticum aestivum-H.villosa translocation lines with small alien segment have been reported.Therefore,it is necessary to establish a high efficient strategy for the induction and identification of alien translocation lines.
In the present study,the mature female gametes of wheat-H.villosa 6VS/6AL translocation line were irradiated by ~(60)Co-γray with 160Rad/M dosage rate and three dosages(1600,1920,2240Rad) before flowering.These irradiated florets were emasculated the same day,and then covered with paper bags.After 2-3 days,these florets were pollinated with normal fresh pollens from T.aestivum cv.Chinese Spring.Genomic in situ hybridization(GISH) at mitosis metaphase of root-tip cell of M_1 plants was used to detect the chromosomes structural changes involving 6VS of H.villosa.Among the 534 M_1 plants screened,97 plants contained chromosome structural changes with small segment of 6VS,including 80 interstitial translocated chromosomes,57 terminal translocated chromosomes and 55 deleted chromosomes.For the 2240 Rad dosage treatment,the induction frequencies of interstitial translocation,terminal translocation and deletion were 21.02%,14.01%,and 14.65%respectively,which were much higher than that previously reported.The M_2 seeds were obtained by backcrossing of 74 M_1 plants involving 146 chromosomes with structural changes of 6VS,and it was found that the structure aberrations in the M_1 plants could be transmitted to their progenies.Irradiating mature female gametes of whole arm translocation may be a new and highly efficient approach for creation of small segment chromosome structural changes,especially for interstitial translocations.These small fragments of 6VS obtained in the present research are different in both length of fragment and breakage location,and are useful for fine location of Pm21 gene,physical mapping of 6VS and development of translocation lines with small fragment.
Using a software PRIMER5.0PCR,primers were designed according to the sequences of ESTs located in the different bins of the short arm of homeologous group 6 chromosomes of wheat.Three molecular markers specific for 6VS(6BS28-386,6DS38-730,6AS6-740) were developed.Twelve deletion lines involved in the short arm of homeologous group 6 chromosomes of wheat and H.villosa were used to physically locate 8 specific markers of 6VS.The markers CINAU17-1086,6VS19-381 were located in the chromosome bin of FL (fragment length) value 0.58-1.00 and 0.58-0.99,the marker CINAU15-902 was located in the chromosome bin of FL value 0.45-0.58,the markers CINAU16-1650,6DS38-730, 6BS28-386 were located in the chromosome bin of FL value 0.35-0.45,and the markers CINAU18-723 and 6AS6-740 were located in the bin of FL value 0.00-0.45.These markers are usefull both for identification of structural changes with 6VS and fine mapping of Pm21.
Through the back-crosses between some M_1 plants with chromosome changes(female parent) and common wheat(male parent),8 M_2 plants with single small interstitial translocation chromosomes of 6VS have been obtained.The above 8 6VS specific molecular markers were used to define the fagment of 6VS.The fragment of 6VS in translocation line NJ9-30,NJ2-2(or NJ2-3) had FL value 0.00-0.58,0.35-0.58,respectively. The fragment of 6VS in the other lines(NJ6-23,NJ6-14 etc) was involved in the segment of FL value 0.00-0.35.
Two heterozygous insterstitial translocation lines with small segment of 6VS,which showed high resistance to powdery mildew,were selected,and homozygous interstitial translocation lines with Pm21 will be obtained.The Pm21 gene was further located in the region of 6VS with FL value 0.35-0.58 by using these genetic stocks of 6VS and powdery mildew resistance identification.
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