黄河中游地区粗山羊草遗传多样性、高分子量谷蛋白亚基基因的克隆及纹枯病抗性分析
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摘要
粗山羊草(Aegilops tauschii, 2n = 2x = 14,DD)是六倍体普通小麦D基因组的供体种,由于仅有特定区域的少数粗山羊草基因型参与了普通小麦的起源,其遗传变异类型远比普通小麦D染色体组丰富。我国黄河中游地区自然分布着粗山羊草,但对其系统收集及规模性深入研究尚未见报导。本研究利用SSR标记比较了我国黄河中游(河南、陕西)粗山羊草与新疆、中东粗山羊草的多样性及遗传分化状况,并针对黄河中游居群粗山羊草的高分子量谷蛋白亚基(high molecular weight glutenin subunit, HMW-GS)的组成、克隆及纹枯病抗性进行了系统研究。
(1)选用分布在粗山羊草14条染色体上的32对SSR引物,对来自中国河南、陕西、新疆和中东地区共147份粗山羊草材料进行遗传分化及多样性分析,结果表明在26个多态性位点中,等位基因数平均为4.15,Nei基因多样性指数(He)平均为0.243,多态性信息含量指数(PIC)平均为0.226;居群间遗传变异差异明显,中东粗山羊草居群具有丰富的遗传变异(He =0.607,PIC =0.551),而来自陕西和河南的粗山羊草资源遗传多样性较低(He =0.055,PIC =0.047)和(He =0.024,PIC =0.021)。AMOVA分子变异分析显示,居群间遗传变异占总变异的52%,达到显著水平;河南粗山羊草和陕西粗山羊草间发生了一定的遗传分化(Fst =0.210),为研究黄河中游粗山羊草资源的起源与分化问题提供了有用的信息与证据,并为小麦品质改良育种提供丰富的遗传资源。
(2)利用十二烷基硫酸钠聚丙烯酰胺凝胶电泳(SDS-PAGE)技术分析了黄河中游地区161份粗山羊草的HMW-GS,发现3种编码序列未知的y-型亚基,即Dy10.5t、Dy10.4t和Dy10.5*t亚基。通过AS-PCR扩增、克隆、测序和氨基酸序列推导,发现3种未知序列均具有典型HMW-GS的序列结构特征且较为相似,仅Dy10.4t与Dy10.5t亚基存在一个氨基酸重复单元的缺失,Dy10.5t与Dy10.5*t亚基在信号肽部位有一个氨基酸的替换(L-F)。通过对这3种HMW-GS体外表达及与32个已知氨基酸序列的HMW-GS多序列比对和系统进化关系分析,证实Dy10.5t、Dy10.4t和Dy10.5*t 3个亚基是D基因组编码的高分子量谷蛋白y-型亚基家族的新成员,同时为研究小麦族的起源和进化提供有用的信息和证据。深入研究黄河中游地区粗山羊草HMW-GS,特别是对其编码基因的测序,对于进一步研究其结构功能以及在小麦品质改良中的应用具有非常重要的意义。
(3)利用改进的纹枯病菌接种法,于2007-2008年两个小麦种植年度,对我国黄河流96份粗山羊草材料纹枯病抗性进行田间鉴定,结果显示,稳定表现近免疫(I)的材料28个,高抗(HR)材料37个,中抗(MR)材料30个,仅SX-6表现中感(MS)。并根据所有供试材料在不同生长期和不同年份的相对抗病指数,对所调查的材料进行抗性多样性聚类分析,综合分析得出共有76份材料表现为较强的纹枯病抗性,是我国小麦改良育种很好的纹枯病菌抗源。系统地调查黄河中游地区粗山羊草的纹枯病特性,从中筛选出的抗性种质为小麦的改良提供了抗源材料。
Aegilops tauschii (2n = 2x = 14,DD), the donor of the D genome of hexaploid wheat (Triticum aestivum, AABBDD), has been found to possess many valuable genes what have been used to improve the quality and biotic stress resistance of bread wheat. Ae. tauschii ,which has a few of region-specific genotypes involved in the origin of hexaploid wheat, contains much wider genetic background than wheat’s. There is no a large-scale investigation of Ae. tauschii in the Middle reaches of the Yellow River in China where many kinds of Ae. tauschii accessions have been found. In this work, the genetic diversity of different Ae. tauschii populations were investigated by SSR assay, and then the molecular cloning of high molecular weight glutenin subunit genes and resistance to wheat sharp eyespot were focused on the Ae. tauschii from the middle reaches of the Yellow River.
(1)The genetic diversity and differentiation of 147 Aegilops tauschii accessions from Henan, Shannxi, Xinjiang in China and the Middle East, were investigated by SSR assay with 32 pairs of primers homogeneously distributed on 14 chromosomes of Ae. tauschii. The results demonstrated that 26 loci were polymorphic, with an average of 4.15 alleles on each locus. Nei’s genetic diversity index (He) and polymorphism information content (PIC) value were 0.243 and 0.226, respectively. Genetic variations among different populations varied greatly. The maximal diversity values were found in Middle East population (He =0.607, PIC =0.551), which was much higher than those in Shannxi (He =0.055, PIC =0.047) and Henan (He =0.024, PIC =0.021) populations. Analysis of molecular variance (AMOVA) showed that the genetic differentiation among the geographical populations was significant (P<0.05) and accounted for about 52% of the total genetic variation. Meanwhile,the genetic differentiation was obvious between the populations of Henan and Shannxi(Fst =0.210). These results would offer useful information and evidence for further understanding the origin and evolution of the Ae. tauschii from the Middle reaches of the Yellow River, and provided genetic resources for wheat breeding.
(2)To detect HMW-GS genes in Ae. tauschii resources in the Middle reaches of the Yellow River and to understand its structure and function by sequencing, 161 Ae. tauschii accessions collected in July 2006 were used in this study with Yumai 49 as a control. Through sodium dodecyl sulphate polyacrylamide gel electrophoresis (SDS-PAGE) technique, three y-type subunits were observed, named as Dy10.5t, Dy10.4t , and Dy10.5*t. Genes of the three subunits were amplified, cloned and sequenced by allele-specific PCR. The results revealed that the deduced amino acid sequence had typical characters of HMW-GS and a general resemblance to each other, but Dy10.4t had a repeated amino acid motif deletion to Dy10.5t, and Dy10.5*t had an amino acid substitution sites in signal peptide domain to Dy10.5t (L-F). From multiple sequence alignment and phylogenetic analysis of storage protein gene family, it is concluded that the subunits Dy10.5t、Dy10.4t and Dy10.5*t of Ae. tauschii are similar to the Glu-1 locus found in wheat and related species, and are new members of y-type HWM glutenin family. It would offer useful information and evidence for further understanding the origin and evolution of Triticum species, and it is potentially a elite gene resources for quality improvement of bread wheat.
(3)The disease resistance of 128 Ae. tauschii to R. cerealis in middle reaches of yellow river of China was identified in fields by an improved method of inoculations during 2007,2008 planting seasons. The resistance to R. cerealis of varieties were different. 28 immunity, 37 high resistant, 30 middle resistant, only SX-6 is middle susceptible were identified among 96 Ae. tauschii varieties. The diversification analysis of Ae. tauschiis resistance to wheat sharp eyespot according to the relative resestance index of four phases revealed that, seventy-six varieties have a higher resistance to R. cerealis and would be used in wheat engineering in future. Investigating the resistance of Ae. tauschii in the middle reaches of yellow river to R.cerealis, will increase the genetic variability and germplasm resources for the improvement of wheat.
(1)选用分布在粗山羊草14条染色体上的32对SSR引物,对来自中国河南、陕西、新疆和中东地区共147份粗山羊草材料进行遗传分化及多样性分析,结果表明在26个多态性位点中,等位基因数平均为4.15,Nei基因多样性指数(He)平均为0.243,多态性信息含量指数(PIC)平均为0.226;居群间遗传变异差异明显,中东粗山羊草居群具有丰富的遗传变异(He =0.607,PIC =0.551),而来自陕西和河南的粗山羊草资源遗传多样性较低(He =0.055,PIC =0.047)和(He =0.024,PIC =0.021)。AMOVA分子变异分析显示,居群间遗传变异占总变异的52%,达到显著水平;河南粗山羊草和陕西粗山羊草间发生了一定的遗传分化(Fst =0.210),为研究黄河中游粗山羊草资源的起源与分化问题提供了有用的信息与证据,并为小麦品质改良育种提供丰富的遗传资源。
(2)利用十二烷基硫酸钠聚丙烯酰胺凝胶电泳(SDS-PAGE)技术分析了黄河中游地区161份粗山羊草的HMW-GS,发现3种编码序列未知的y-型亚基,即Dy10.5t、Dy10.4t和Dy10.5*t亚基。通过AS-PCR扩增、克隆、测序和氨基酸序列推导,发现3种未知序列均具有典型HMW-GS的序列结构特征且较为相似,仅Dy10.4t与Dy10.5t亚基存在一个氨基酸重复单元的缺失,Dy10.5t与Dy10.5*t亚基在信号肽部位有一个氨基酸的替换(L-F)。通过对这3种HMW-GS体外表达及与32个已知氨基酸序列的HMW-GS多序列比对和系统进化关系分析,证实Dy10.5t、Dy10.4t和Dy10.5*t 3个亚基是D基因组编码的高分子量谷蛋白y-型亚基家族的新成员,同时为研究小麦族的起源和进化提供有用的信息和证据。深入研究黄河中游地区粗山羊草HMW-GS,特别是对其编码基因的测序,对于进一步研究其结构功能以及在小麦品质改良中的应用具有非常重要的意义。
(3)利用改进的纹枯病菌接种法,于2007-2008年两个小麦种植年度,对我国黄河流96份粗山羊草材料纹枯病抗性进行田间鉴定,结果显示,稳定表现近免疫(I)的材料28个,高抗(HR)材料37个,中抗(MR)材料30个,仅SX-6表现中感(MS)。并根据所有供试材料在不同生长期和不同年份的相对抗病指数,对所调查的材料进行抗性多样性聚类分析,综合分析得出共有76份材料表现为较强的纹枯病抗性,是我国小麦改良育种很好的纹枯病菌抗源。系统地调查黄河中游地区粗山羊草的纹枯病特性,从中筛选出的抗性种质为小麦的改良提供了抗源材料。
Aegilops tauschii (2n = 2x = 14,DD), the donor of the D genome of hexaploid wheat (Triticum aestivum, AABBDD), has been found to possess many valuable genes what have been used to improve the quality and biotic stress resistance of bread wheat. Ae. tauschii ,which has a few of region-specific genotypes involved in the origin of hexaploid wheat, contains much wider genetic background than wheat’s. There is no a large-scale investigation of Ae. tauschii in the Middle reaches of the Yellow River in China where many kinds of Ae. tauschii accessions have been found. In this work, the genetic diversity of different Ae. tauschii populations were investigated by SSR assay, and then the molecular cloning of high molecular weight glutenin subunit genes and resistance to wheat sharp eyespot were focused on the Ae. tauschii from the middle reaches of the Yellow River.
(1)The genetic diversity and differentiation of 147 Aegilops tauschii accessions from Henan, Shannxi, Xinjiang in China and the Middle East, were investigated by SSR assay with 32 pairs of primers homogeneously distributed on 14 chromosomes of Ae. tauschii. The results demonstrated that 26 loci were polymorphic, with an average of 4.15 alleles on each locus. Nei’s genetic diversity index (He) and polymorphism information content (PIC) value were 0.243 and 0.226, respectively. Genetic variations among different populations varied greatly. The maximal diversity values were found in Middle East population (He =0.607, PIC =0.551), which was much higher than those in Shannxi (He =0.055, PIC =0.047) and Henan (He =0.024, PIC =0.021) populations. Analysis of molecular variance (AMOVA) showed that the genetic differentiation among the geographical populations was significant (P<0.05) and accounted for about 52% of the total genetic variation. Meanwhile,the genetic differentiation was obvious between the populations of Henan and Shannxi(Fst =0.210). These results would offer useful information and evidence for further understanding the origin and evolution of the Ae. tauschii from the Middle reaches of the Yellow River, and provided genetic resources for wheat breeding.
(2)To detect HMW-GS genes in Ae. tauschii resources in the Middle reaches of the Yellow River and to understand its structure and function by sequencing, 161 Ae. tauschii accessions collected in July 2006 were used in this study with Yumai 49 as a control. Through sodium dodecyl sulphate polyacrylamide gel electrophoresis (SDS-PAGE) technique, three y-type subunits were observed, named as Dy10.5t, Dy10.4t , and Dy10.5*t. Genes of the three subunits were amplified, cloned and sequenced by allele-specific PCR. The results revealed that the deduced amino acid sequence had typical characters of HMW-GS and a general resemblance to each other, but Dy10.4t had a repeated amino acid motif deletion to Dy10.5t, and Dy10.5*t had an amino acid substitution sites in signal peptide domain to Dy10.5t (L-F). From multiple sequence alignment and phylogenetic analysis of storage protein gene family, it is concluded that the subunits Dy10.5t、Dy10.4t and Dy10.5*t of Ae. tauschii are similar to the Glu-1 locus found in wheat and related species, and are new members of y-type HWM glutenin family. It would offer useful information and evidence for further understanding the origin and evolution of Triticum species, and it is potentially a elite gene resources for quality improvement of bread wheat.
(3)The disease resistance of 128 Ae. tauschii to R. cerealis in middle reaches of yellow river of China was identified in fields by an improved method of inoculations during 2007,2008 planting seasons. The resistance to R. cerealis of varieties were different. 28 immunity, 37 high resistant, 30 middle resistant, only SX-6 is middle susceptible were identified among 96 Ae. tauschii varieties. The diversification analysis of Ae. tauschiis resistance to wheat sharp eyespot according to the relative resestance index of four phases revealed that, seventy-six varieties have a higher resistance to R. cerealis and would be used in wheat engineering in future. Investigating the resistance of Ae. tauschii in the middle reaches of yellow river to R.cerealis, will increase the genetic variability and germplasm resources for the improvement of wheat.
引文
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