斯卑尔脱小麦种质资源遗传评价及育种利用研究
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摘要
斯卑尔脱小麦(Triticum spelta L.,AABBDD)是小麦属中的六倍体带皮栽培种之一,是改良普通小麦(Triticum aestivum L.,AABBDD)产量、品质、抗病性和抗逆性的重要基因资源,对其种质资源进行遗传评价有助于普通小麦育种利用。本文的主要结果如下:
1.对151份斯卑尔脱小麦主要农艺性状进行考察与分析的结果表明,供试材料具有植株高大、分蘖力强、小穗数多和千粒重偏低等特点,但也可从中筛选到植株偏矮、干粒重较高的材料供育种利用。多分蘖及多小穗是斯卑尔脱小麦最具利用价值的农艺性状。相关分析表明,千粒重与穗长、小穗数和穗粒重等多个性状存在复杂的相关关系。基于农艺性状表现供试材料可聚为四类,聚类结果与其地理来源并不完全一致。对农艺性状的综合分析后筛选出5份优异材料,可供育种利用。
2.测定了142份斯卑尔脱小麦12个品质指标,并根据国家现行强筋小麦品种品质标准和专用小麦品种品质标准对所测品质指标进行了分析和评价。结果表明:供试材料中113份为软质,25份为中软质,4份为硬质。除2份材料外,其余140份材料均为角质。平均蛋白质含量高于16%,变幅为12.04%-21.67%;104份材料的湿面筋含量＞28%,达到中筋小麦要求;沉降值大于50ml的材料占总数的56.82%。粉质参数测试表明供试材料面团吸水率高于70%的材料有73份,形成时间大于7.5min的材料有3份;供试材料的淀粉含量低、延展性弱、面包体积偏小。从供试材料中筛选到高蛋白(大于20%)资源材料3份,高沉降值(大于75ml)材料10份,可利用这些优异种质资源改良普通小麦品质。
3.对162份斯卑尔脱小麦贮藏蛋白的遗传多样性进行了评价。结果表明,利用A-PAGE技术共检测出121种醇溶蛋白带型,未发现共有条带,说明斯卑尔脱小麦具有丰富的醇溶蛋白等位变异,且ω/γ-区的变异较α/β-区丰富。利用SDS-PAGE技术,共分离出9种高分子量麦谷蛋白亚基,组成8种亚基组合。其中1、6+8、2+12亚基组合为优势亚基组合,占供试材料的83.00%,高分子量谷蛋白亚基具有物种特异性。在供试材料中,筛选到含有与烘烤品质正相关的7+8、5+10等亚基的材料供育种利用。
4.根据已知的普通小麦α-醇溶蛋白基因序列设计引物,采用PCR方法首次从斯卑尔脱小麦材料NGB5149中克隆得到两个α-醇溶蛋白基因序列Gli-Spelt-1和Gli-Spelt-2(GenBank登录号分别为DQ234066和DQ234067)。它们具有α-醇溶蛋白基因的典型结构特征,但Gli-Spelt-1是一个假基因。Gli-Spelt-2编码区全长849bp,编码263个氨基酸。氨基酸序列比对显示,Gli-Spelt-1和Gli-Spelt-2与已报道的其他小麦族物种中的α-醇溶蛋白序列有较高的一致性,并与小麦氨基酸组分的含量变化趋势一致。斯卑尔脱小麦推测氨基酸序列包含有对腹泻病人的毒性序列,第一次从分子水平证实了对小麦谷蛋白易感人群仍要谨慎食用斯卑尔脱小麦面粉制品。
5.根据已经发布的ARF基因序列设计了一对ARF引物,在斯卑尔脱小麦中能扩增到约1200bp的DNA片断。通过测序首次发现该基因存在内含子,且内含子可能存在独特的剪接方式。氨基酸序列分析表明,斯卑尔脱小麦的ARF同其它物种一样也具有保守的GTP结合区域,即P(GLDAAGKT)、G(NKQD)以及G'(DVGGQ)。利用生物信息学技术对ARF基因进行多序列比对分析后发现该基因在DNA序列上存在大量潜在的SNP位点。本研究根据SNP位点开发了染色体组特异性标记,并利用中国春缺体-四体,第一次将来源于A染色体组和D染色体组的SNP特异性标记定位在2AL和3DL上。首次利用半定量RT-PCR技术对ARF基因在斯卑尔脱小麦中的定向表达进行了分析,发现该基因在根和幼胚中的表达量较茎和旗叶高。
6.利用普通小麦品种川农16与6个斯卑尔脱小麦材料进行杂交。杂种F_1能够正常结实,各考察性状均介于双亲之间。对6个杂种的F_2-F_4代品质性状的测试结果表明,各组合的品质性状较母本川农16均有较大改善,其中组合CN16/P1355625表现最为优异。对组合CN16/NGB4798F_2代的谷蛋白和醇溶蛋白的检测发现,F_2代谷蛋白亚基组成倾向于川农16的亚基组成,亚基组合1、20、5+10在F_2代所占频率最高,且醇溶蛋白谱带表现双亲的共同特征。
Currently, there is considerable interest in the use of spelt (Triticum spelta. L.) for food destined for health and organic food markets. Spelt is an ancient cereal that was once cultivated extensively and is now receiving a renewal interest for its nutritional qualities and, from an agronomic point of view, for its resistance to pests and its ability to grow in soils with limited fertility and in wet and cold climates where it is better than weat. T. spelta, with many useful genes for wheat improvement, is the primary gene pool of bread wheat and easy to be utilized in wheat improvement. Thus, it is important to evaluate the germplasm of spelt wheat. In this paper, the population genetic diversity was estimated based on agronomic traits, quality characters, biochemical and molecular markers. The following results were obtained.
1. The agronomic traits of 151 spelt wheat accessions collected from 18 countries and areas, were investigated based on the correlation, principal component and cluster analysis. Higher variations of the agronomic traits were observed. The spelt wheat investigated in this study had the beneficial traits of more tillers and spikelet per spike, and disadvantage of higher plant height, lower 1000-grains weight and longer growing periods. The correlation analysis indicated that the spike number per plant limited the production of spelt wheat. By clustering analysis, all accessions were divided into four groups, and the genetic distances based on agronomic traits were associated with their geographic distribution. Based on principle elements analysis, five principle elements were obtained, which contributes the variance over 85%. According to these main characters, tweenty-two accessions with one or some elite agronomic performances were screen out.
2. The analysis of quality characters showed that most of spelt kernels were soft and dextrous. Comprared with common wheat, spelt wheat had higher protein content, wet gluten, gluten index and sedimentation values. As rheologic traits were concerned, spelt wheat had higher water absorption and longer development time, whereas spelt wheat showed shorter stable time. Spelt wheat had lower starch content, weaker extension and lower loaf volume. According to these quality characters, all the materials were divided into four groups by clustering analysis. Six accessions with elite agronomic and quality characters were selected for the utilization in wheat improvment.
3. Gliadin and HMW glutenin subunit variations in 162 spelt wheat accessions were detected by A-PAGE and SDS-PAGE. Higher gliadin variation was observed, and 121 gliadin patterns were detected. A total of 9 HMW-GS alleles were found. There were 2, 5 and 2 alleles at Glu-A1, Glu-B1 and Glu-D1 loci, respectively, resulting in 8 HMW-GS combinations. The HMW-GS combination (1, 6+8, 2+12) was the dominant phenotype, which was found in 83.00% accessions. These results suggested that the polymorphism of spelt wheat on Gli-1 loci was higher than that on Glu-1 loci.
4. To get more information onα-gliadin genes of spelt wheat, specific PCR primers were designed based on the known wheatα-gliadin gene sequences, and the coding regions of two gliadin genes Gli-Spelt-1 and Gli-Spelt-2 were isolated from spelt wheat NGB5149. Gli-Spelt-1 was a pseudogene due to the stop codons in its coding region. The full coding region of Spelt-Gli-2 was 849bp, and could be translated into a protein of 263 amino acids. The two cloned gliadin genes had the typical structure ofα-gliadin genes. The deduced amino acid sequences comparison suggested that Spelt-Gli-2 had the high identity with the knownα-gliadin genes.
5. ADP-ribosylation factors (ARFs), a subfamily of the small GTP binding proteins superfamily, are believed to participate in vesicular transport and signal transduction events in the cell. The DNA sequences included four exons and three introns. Its deduced amino acid sequence had consensus with GTP-binding domain P (GLDAAGKT), G'(DVGGQ) and G (NKQD). Moreover, in two ends of introns of ARF gene from diploid species, common wheat and spelt wheat, there were new base sequences which did not comply with typical GU-AG rule. We speculated that these new base sequences might have relation to a new splice process. To assign the chromosomal locations of ARF genes in common wheat, 20 specific primer sets were designed on the specific SNPs, and PCR assay of the genomic DNAs of Chinese Spring ditelosomic lines were carried out. The ARF genes were located on the chromosomes 2AL and 3DL. ARF transcripts were detected mainly in root and immature embryo.
6. High heterosis was obtained in the hybrids between common wheat and spelt wheat. The yield per plant, grains weight per spike and 1000 grain weight had higher heterosis. The heading dates of interspecific hybrids were 0 to 6 days later than female parents (common wheat) and 20 days earlier than male parents (spelt wheat). The analysis the quality characters showed that the the performance of quality characters in interspecific hybrids were superior to the female parents (common wheat), indicating that interspecific hybrids are very useful and efficient for the improvement of bread wheat by exploiting desirable genes in spelt wheat.
1.对151份斯卑尔脱小麦主要农艺性状进行考察与分析的结果表明,供试材料具有植株高大、分蘖力强、小穗数多和千粒重偏低等特点,但也可从中筛选到植株偏矮、干粒重较高的材料供育种利用。多分蘖及多小穗是斯卑尔脱小麦最具利用价值的农艺性状。相关分析表明,千粒重与穗长、小穗数和穗粒重等多个性状存在复杂的相关关系。基于农艺性状表现供试材料可聚为四类,聚类结果与其地理来源并不完全一致。对农艺性状的综合分析后筛选出5份优异材料,可供育种利用。
2.测定了142份斯卑尔脱小麦12个品质指标,并根据国家现行强筋小麦品种品质标准和专用小麦品种品质标准对所测品质指标进行了分析和评价。结果表明:供试材料中113份为软质,25份为中软质,4份为硬质。除2份材料外,其余140份材料均为角质。平均蛋白质含量高于16%,变幅为12.04%-21.67%;104份材料的湿面筋含量＞28%,达到中筋小麦要求;沉降值大于50ml的材料占总数的56.82%。粉质参数测试表明供试材料面团吸水率高于70%的材料有73份,形成时间大于7.5min的材料有3份;供试材料的淀粉含量低、延展性弱、面包体积偏小。从供试材料中筛选到高蛋白(大于20%)资源材料3份,高沉降值(大于75ml)材料10份,可利用这些优异种质资源改良普通小麦品质。
3.对162份斯卑尔脱小麦贮藏蛋白的遗传多样性进行了评价。结果表明,利用A-PAGE技术共检测出121种醇溶蛋白带型,未发现共有条带,说明斯卑尔脱小麦具有丰富的醇溶蛋白等位变异,且ω/γ-区的变异较α/β-区丰富。利用SDS-PAGE技术,共分离出9种高分子量麦谷蛋白亚基,组成8种亚基组合。其中1、6+8、2+12亚基组合为优势亚基组合,占供试材料的83.00%,高分子量谷蛋白亚基具有物种特异性。在供试材料中,筛选到含有与烘烤品质正相关的7+8、5+10等亚基的材料供育种利用。
4.根据已知的普通小麦α-醇溶蛋白基因序列设计引物,采用PCR方法首次从斯卑尔脱小麦材料NGB5149中克隆得到两个α-醇溶蛋白基因序列Gli-Spelt-1和Gli-Spelt-2(GenBank登录号分别为DQ234066和DQ234067)。它们具有α-醇溶蛋白基因的典型结构特征,但Gli-Spelt-1是一个假基因。Gli-Spelt-2编码区全长849bp,编码263个氨基酸。氨基酸序列比对显示,Gli-Spelt-1和Gli-Spelt-2与已报道的其他小麦族物种中的α-醇溶蛋白序列有较高的一致性,并与小麦氨基酸组分的含量变化趋势一致。斯卑尔脱小麦推测氨基酸序列包含有对腹泻病人的毒性序列,第一次从分子水平证实了对小麦谷蛋白易感人群仍要谨慎食用斯卑尔脱小麦面粉制品。
5.根据已经发布的ARF基因序列设计了一对ARF引物,在斯卑尔脱小麦中能扩增到约1200bp的DNA片断。通过测序首次发现该基因存在内含子,且内含子可能存在独特的剪接方式。氨基酸序列分析表明,斯卑尔脱小麦的ARF同其它物种一样也具有保守的GTP结合区域,即P(GLDAAGKT)、G(NKQD)以及G'(DVGGQ)。利用生物信息学技术对ARF基因进行多序列比对分析后发现该基因在DNA序列上存在大量潜在的SNP位点。本研究根据SNP位点开发了染色体组特异性标记,并利用中国春缺体-四体,第一次将来源于A染色体组和D染色体组的SNP特异性标记定位在2AL和3DL上。首次利用半定量RT-PCR技术对ARF基因在斯卑尔脱小麦中的定向表达进行了分析,发现该基因在根和幼胚中的表达量较茎和旗叶高。
6.利用普通小麦品种川农16与6个斯卑尔脱小麦材料进行杂交。杂种F_1能够正常结实,各考察性状均介于双亲之间。对6个杂种的F_2-F_4代品质性状的测试结果表明,各组合的品质性状较母本川农16均有较大改善,其中组合CN16/P1355625表现最为优异。对组合CN16/NGB4798F_2代的谷蛋白和醇溶蛋白的检测发现,F_2代谷蛋白亚基组成倾向于川农16的亚基组成,亚基组合1、20、5+10在F_2代所占频率最高,且醇溶蛋白谱带表现双亲的共同特征。
Currently, there is considerable interest in the use of spelt (Triticum spelta. L.) for food destined for health and organic food markets. Spelt is an ancient cereal that was once cultivated extensively and is now receiving a renewal interest for its nutritional qualities and, from an agronomic point of view, for its resistance to pests and its ability to grow in soils with limited fertility and in wet and cold climates where it is better than weat. T. spelta, with many useful genes for wheat improvement, is the primary gene pool of bread wheat and easy to be utilized in wheat improvement. Thus, it is important to evaluate the germplasm of spelt wheat. In this paper, the population genetic diversity was estimated based on agronomic traits, quality characters, biochemical and molecular markers. The following results were obtained.
1. The agronomic traits of 151 spelt wheat accessions collected from 18 countries and areas, were investigated based on the correlation, principal component and cluster analysis. Higher variations of the agronomic traits were observed. The spelt wheat investigated in this study had the beneficial traits of more tillers and spikelet per spike, and disadvantage of higher plant height, lower 1000-grains weight and longer growing periods. The correlation analysis indicated that the spike number per plant limited the production of spelt wheat. By clustering analysis, all accessions were divided into four groups, and the genetic distances based on agronomic traits were associated with their geographic distribution. Based on principle elements analysis, five principle elements were obtained, which contributes the variance over 85%. According to these main characters, tweenty-two accessions with one or some elite agronomic performances were screen out.
2. The analysis of quality characters showed that most of spelt kernels were soft and dextrous. Comprared with common wheat, spelt wheat had higher protein content, wet gluten, gluten index and sedimentation values. As rheologic traits were concerned, spelt wheat had higher water absorption and longer development time, whereas spelt wheat showed shorter stable time. Spelt wheat had lower starch content, weaker extension and lower loaf volume. According to these quality characters, all the materials were divided into four groups by clustering analysis. Six accessions with elite agronomic and quality characters were selected for the utilization in wheat improvment.
3. Gliadin and HMW glutenin subunit variations in 162 spelt wheat accessions were detected by A-PAGE and SDS-PAGE. Higher gliadin variation was observed, and 121 gliadin patterns were detected. A total of 9 HMW-GS alleles were found. There were 2, 5 and 2 alleles at Glu-A1, Glu-B1 and Glu-D1 loci, respectively, resulting in 8 HMW-GS combinations. The HMW-GS combination (1, 6+8, 2+12) was the dominant phenotype, which was found in 83.00% accessions. These results suggested that the polymorphism of spelt wheat on Gli-1 loci was higher than that on Glu-1 loci.
4. To get more information onα-gliadin genes of spelt wheat, specific PCR primers were designed based on the known wheatα-gliadin gene sequences, and the coding regions of two gliadin genes Gli-Spelt-1 and Gli-Spelt-2 were isolated from spelt wheat NGB5149. Gli-Spelt-1 was a pseudogene due to the stop codons in its coding region. The full coding region of Spelt-Gli-2 was 849bp, and could be translated into a protein of 263 amino acids. The two cloned gliadin genes had the typical structure ofα-gliadin genes. The deduced amino acid sequences comparison suggested that Spelt-Gli-2 had the high identity with the knownα-gliadin genes.
5. ADP-ribosylation factors (ARFs), a subfamily of the small GTP binding proteins superfamily, are believed to participate in vesicular transport and signal transduction events in the cell. The DNA sequences included four exons and three introns. Its deduced amino acid sequence had consensus with GTP-binding domain P (GLDAAGKT), G'(DVGGQ) and G (NKQD). Moreover, in two ends of introns of ARF gene from diploid species, common wheat and spelt wheat, there were new base sequences which did not comply with typical GU-AG rule. We speculated that these new base sequences might have relation to a new splice process. To assign the chromosomal locations of ARF genes in common wheat, 20 specific primer sets were designed on the specific SNPs, and PCR assay of the genomic DNAs of Chinese Spring ditelosomic lines were carried out. The ARF genes were located on the chromosomes 2AL and 3DL. ARF transcripts were detected mainly in root and immature embryo.
6. High heterosis was obtained in the hybrids between common wheat and spelt wheat. The yield per plant, grains weight per spike and 1000 grain weight had higher heterosis. The heading dates of interspecific hybrids were 0 to 6 days later than female parents (common wheat) and 20 days earlier than male parents (spelt wheat). The analysis the quality characters showed that the the performance of quality characters in interspecific hybrids were superior to the female parents (common wheat), indicating that interspecific hybrids are very useful and efficient for the improvement of bread wheat by exploiting desirable genes in spelt wheat.
引文
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