小麦脂肪氧化酶（LOX）活性QTL定位与功能标记开发
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摘要
面粉颜色是决定小麦品质的重要指标,对面条及其它相关制品品质有重要影响。提高脂肪氧化酶(Lipoxygenase, LOX)活性有助于改善中国食品的面粉颜色和面团流变学特性。改良LOX活性已成为小麦品质育种的重要目标。本研究选用中优9507/CA9632的71个双单倍体(DH)系和198份中国冬小麦区的主栽品种和高代品系,进行LOX活性的QTL分析；应用电子克隆结合PCR扩增方法,克隆LOX基因并分析其在不同品种中的等位变异,根据各等位基因之间的序列差异开发可用于分子标记辅助选择的功能标记；在上述工作的基础上分析了这些材料基因型与表型之间的关系。主要结果如下：
1.对中优9507/CA9632 DH系在不同环境下的分析表明,基因型和环境对LOX活性均有显著影响,但基因型与环境互作效应不显著,籽粒LOX活性的广义遗传力为0.94；我国冬小麦品种(系)间籽粒LOX活性存在较大差异,变异范围为41.7-112.7 A234.min~(-1).g(-1)。因此,用高LOX活性的亲本配制杂交组合,在育种早期世代进行较严格的选择,能选育出高LOX活性的品种。
2.利用中优9507/CA9632 DH系和SSR标记对普通小麦LOX活性进行数量性状位点(Quantitative trait loci, QTL)定位。共检测到两个QTL,其中QLpx.caas.1AL位于1AS染色体上,与Xwmc312紧密连锁；另一QTL QLpx.caas-4B位于4BS染色体上,与Xgwm251紧密连锁。QLpx.caas.1AL和QLpx.caas-4B在四个不同的环境下能分别解释表型变异的13.4-25.2%和14.3-27%。此外,分别用Xwmc312和Xgwm251检测198份我国冬小麦材料,不同基因型的LOX活性差异达到1%的显著水平。用两个标记Xwmc312/Xgwm251进一步构建了多重PCR,并对198份冬小麦材料进行检测,表明该多重PCR能有效地应用于分子标记辅助选择育种。
3.利用电子克隆方法克隆了普通小麦4B和5B染色体上LOX基因TaLox-B1和TaLox-B2的全长编码序列。TaLox-B1的基因组DNA序列由4,289bp个碱基对组成,包括7个外显子和6个内含子,以及26bp的5’非翻译区(untranslated region, UTR),并含有2,586bp的开放读码框(Open reading frame, ORF),编码一段861个氨基酸残基的多肽链。TaLox-B2的基因组DNA序列由3,325个碱基对组成,包括6个外显子和5个内含子,以及73bp的3’UTR,并含有一个2,595 bp的开放读码框,编码一段864个氨基酸残基的多肽链。对LOX基因的聚类分析表明,来源于同一同源群的LOX基因在gDNA、cDNA和AA三水平上均具有很高的序列相似性。
4.基于TaLox-B1位点的等位变异TaLox-B1a和TaLox-B1b,开发了两个互补显性标记LOX16和LOX18。LOX16在TaLox-B1a的材料中能扩增出489bp片段,与高LOX活性相关,而在TaLox-B1b类型的材料中没有扩增片段。LOX18则只能在TaLox-B1b类型的材料中扩增出791bp片段,并与低LOX活性相关,在TaLox-B1a类型的材料中没有扩增片段。利用中优9507/CA9632 DH系和一套中国春缺体四体系将LOX16和LOX18定位在小麦4BS染色体。QTL分析表明,互补功能标记LOX16和LOX18与位于4BS染色体的SSR标记Xgwm251紧密连锁,遗传距离为5.4cM。该QTL在4个不同环境中可解释15.9-26.2%的表型变异。用功能标记LOX16和LOX18检测198份中国冬小麦材料,不同基因型的LOX活性差异达到1%显著水平。因此,互补显性标记LOX16和LOX18与LOX基因相关并能有效地用于LOX活性的遗传改良。
5.比较北部冬麦区和黄淮冬麦区小麦品种中TaLox-B1等位变异的分布表明,TaLox-B1b是优势等位变异(65.7%)。此外,在黄淮冬麦区高LOX活性等位变异TaLox-B1a的频率(36.0%)显著高于北部冬麦区(25.0%),表明我国小麦LOX活性选择潜力大,通过分子标记辅助选择将加速LOX活性的遗传改良。
Flour color is an important trait in the assessment of flour quality and exerts significant influence on qualities of noodles and other related products. Lipoxygenase (LOX) activity in grain influences the processing quality of wheat-based products. Characterization of LOX genes and development of functional markers are of importance for marker-assisted selection in wheat breeding. In the present study,71 doubled haploid (DH) lines derived from Zhongyou 9507/CA9632 and 198 Chinese winter wheat cultivars and advanced lines were used and their grain LOX activity was evaluated. LOX gene associated with grain LOX activity, was cloned by the method of in silico cloning in combination with PCR amplification, and functional markers were developed for the genes according to their allelic variants in different wheat cultivars, and their associations with phenotypes were then analyzed. The main results obtained in this study are summarized below.
1. LOX activity varies among wheat genotypes and it is also significantly affected by environments. However, the LOX activities of the DH lines across four environments have a high heritability (hl= 0.94), indicating that it was mainly affected by genetypes. Significant differences in grain LOX activity were observed in the Chinese winter wheat cultivars. Grain LOX activity among Chinese winter wheats ranged from 48.4 to 89.6 A234.min-1.g-1. Therefore, selection for LOX activity can be effectively applied at the earlier generation of breeding program. It is possible to breed new wheat cultivars with high LOX activity by choosing parents and making selection in early generations.
2. Quantitative trait loci (QTLs) for LOX activity in common wheat were mapped using 71 doubled haploid (DH) lines derived from a Zhongyou 9507/CA9632 and SSR markers. Two QTLs QLpx.caas.lAL and QLpx.caas-4B were identified on chromosomes 1AL and 4BS, closely related to LOX activity. The SSR loci Xwmc312 and Xgwm251 were proven to be diagnostic and explained 13.4-25.2% of phenotypic variance the 1AL locus and 14.3-27.0% for the 4B locus across four environments. The SSR markers Xwmc312 and Xgwm251 were validated across 198 Chinese wheat cultivars and advanced lines and showed highly significant (P<0.01) association with LOX activity. We further established a multiplexed PCR with SSR marker combination Xwmc312/Xgwm251 to test these wheat cultivars and advanced lines. The results suggested that the marker combination Xwmc312/Xgwm251 is efficient and reliable for evaluating LOX activity and can be used in marker-assisted selection (MAS) targeting for flour color attributes to noodle and other wheat-based products.
3. The full-length genomic DNA sequence of TaLox-B1 and TaLox-B2 on chromosome 4BS and 5B, respectively, were characterized by in silico cloning and experimental validation. The cloned TaLox-B1 comprises seven exons and six introns, with 4,263 bp in total and an open reading frame (ORF) of 2,586 bp, and it encodes a LOX precursor peptide of 861 amino acids with a predicted molecular weight of-96.0 kDa. The genomic DNA sequence of TaLox-B2 comprised 3,325 base pairs, including six exons and five introns, and partial 3'UTR. The gene harbored an ORF of 2,595 bp, encoding a polypeptide of 864 amino acids with a predicted molecular weight of-96.4 kDa. TaLox-B1 and TaLOX1 were mapped on the homologous group 4, and TaLox-B2 and TaLOX2 were mapped on the homologous group 5 in wheat. Cluster analysis of TaLox-B1-TaLox-B2、TaLOXl、TaLOX2 and ZmLOX3 indicated that these LOX genes from the same homologous group showed the highest sequence identities in gDNA, cDNA and AA levels, respectively.
4. Functional markers for the LOX gene on chromosomes 4B based on single nucleotide polymorphism were developed and validated in common wheat. Two complementary dominant STS markers, LOX 16 and LOX 18, were developed based on the single nucleotide polymorphism (SNP) and amplifying 489-bp and 791-bp fragments in cultivars with higher and lower LOX activitives, respectively. The two markers were mapped on chromosome 4BS using a doubled haploid (DH) population derived from Zhongyou 9507/CA9632, and a set of Chinese Spring nullisomic-tetrasomic lines and ditelosomic line 4B. QTL analysis indicated that TaLox-B1 co-segregated with two functional markers and was closely linked to SSR marker Xgwm251 on chromosome 4BS with a genetic distance of 5.4 cM, explaining 15.9-26.2% of the phenotypic variance for LOX activity across four environments. Two complementary dominant functional markers LOX 16 and LOX 18, were validated on 198 Chinese wheat cultivars and advanced lines and showed highly significant (P<0.01) association with LOX activity. The results suggested that these two STS markers are closely related to LOX activity and could be used for the improvement of flour color attributes for noodle and other wheat-based products.
5. TaLox-B1 b in Northern China Plain Winter Wheat Rgion and the Yellow and the Huai River Valley Winter Wheat Region was dominant. TaLox-B1a was a superior allele for flour color and dough rheology. TaLox-Bla associated with higher LOX activitive had low frequency (34.3%) in these two regions. The frequency of TaLox-Bla in Yellow and Huai River Valley Winter Wheat Region (36.0%) were higher than those in Northern China Plain Winter Wheat Region (25.0%) (P0.01). These results indicated the preferred TaLox-B1a has not yet widely used in Chinese winter wheat regions. Therefore, it has a large potential to select new cultivars with high LOX activity.
1.对中优9507/CA9632 DH系在不同环境下的分析表明,基因型和环境对LOX活性均有显著影响,但基因型与环境互作效应不显著,籽粒LOX活性的广义遗传力为0.94；我国冬小麦品种(系)间籽粒LOX活性存在较大差异,变异范围为41.7-112.7 A234.min~(-1).g(-1)。因此,用高LOX活性的亲本配制杂交组合,在育种早期世代进行较严格的选择,能选育出高LOX活性的品种。
2.利用中优9507/CA9632 DH系和SSR标记对普通小麦LOX活性进行数量性状位点(Quantitative trait loci, QTL)定位。共检测到两个QTL,其中QLpx.caas.1AL位于1AS染色体上,与Xwmc312紧密连锁；另一QTL QLpx.caas-4B位于4BS染色体上,与Xgwm251紧密连锁。QLpx.caas.1AL和QLpx.caas-4B在四个不同的环境下能分别解释表型变异的13.4-25.2%和14.3-27%。此外,分别用Xwmc312和Xgwm251检测198份我国冬小麦材料,不同基因型的LOX活性差异达到1%的显著水平。用两个标记Xwmc312/Xgwm251进一步构建了多重PCR,并对198份冬小麦材料进行检测,表明该多重PCR能有效地应用于分子标记辅助选择育种。
3.利用电子克隆方法克隆了普通小麦4B和5B染色体上LOX基因TaLox-B1和TaLox-B2的全长编码序列。TaLox-B1的基因组DNA序列由4,289bp个碱基对组成,包括7个外显子和6个内含子,以及26bp的5’非翻译区(untranslated region, UTR),并含有2,586bp的开放读码框(Open reading frame, ORF),编码一段861个氨基酸残基的多肽链。TaLox-B2的基因组DNA序列由3,325个碱基对组成,包括6个外显子和5个内含子,以及73bp的3’UTR,并含有一个2,595 bp的开放读码框,编码一段864个氨基酸残基的多肽链。对LOX基因的聚类分析表明,来源于同一同源群的LOX基因在gDNA、cDNA和AA三水平上均具有很高的序列相似性。
4.基于TaLox-B1位点的等位变异TaLox-B1a和TaLox-B1b,开发了两个互补显性标记LOX16和LOX18。LOX16在TaLox-B1a的材料中能扩增出489bp片段,与高LOX活性相关,而在TaLox-B1b类型的材料中没有扩增片段。LOX18则只能在TaLox-B1b类型的材料中扩增出791bp片段,并与低LOX活性相关,在TaLox-B1a类型的材料中没有扩增片段。利用中优9507/CA9632 DH系和一套中国春缺体四体系将LOX16和LOX18定位在小麦4BS染色体。QTL分析表明,互补功能标记LOX16和LOX18与位于4BS染色体的SSR标记Xgwm251紧密连锁,遗传距离为5.4cM。该QTL在4个不同环境中可解释15.9-26.2%的表型变异。用功能标记LOX16和LOX18检测198份中国冬小麦材料,不同基因型的LOX活性差异达到1%显著水平。因此,互补显性标记LOX16和LOX18与LOX基因相关并能有效地用于LOX活性的遗传改良。
5.比较北部冬麦区和黄淮冬麦区小麦品种中TaLox-B1等位变异的分布表明,TaLox-B1b是优势等位变异(65.7%)。此外,在黄淮冬麦区高LOX活性等位变异TaLox-B1a的频率(36.0%)显著高于北部冬麦区(25.0%),表明我国小麦LOX活性选择潜力大,通过分子标记辅助选择将加速LOX活性的遗传改良。
Flour color is an important trait in the assessment of flour quality and exerts significant influence on qualities of noodles and other related products. Lipoxygenase (LOX) activity in grain influences the processing quality of wheat-based products. Characterization of LOX genes and development of functional markers are of importance for marker-assisted selection in wheat breeding. In the present study,71 doubled haploid (DH) lines derived from Zhongyou 9507/CA9632 and 198 Chinese winter wheat cultivars and advanced lines were used and their grain LOX activity was evaluated. LOX gene associated with grain LOX activity, was cloned by the method of in silico cloning in combination with PCR amplification, and functional markers were developed for the genes according to their allelic variants in different wheat cultivars, and their associations with phenotypes were then analyzed. The main results obtained in this study are summarized below.
1. LOX activity varies among wheat genotypes and it is also significantly affected by environments. However, the LOX activities of the DH lines across four environments have a high heritability (hl= 0.94), indicating that it was mainly affected by genetypes. Significant differences in grain LOX activity were observed in the Chinese winter wheat cultivars. Grain LOX activity among Chinese winter wheats ranged from 48.4 to 89.6 A234.min-1.g-1. Therefore, selection for LOX activity can be effectively applied at the earlier generation of breeding program. It is possible to breed new wheat cultivars with high LOX activity by choosing parents and making selection in early generations.
2. Quantitative trait loci (QTLs) for LOX activity in common wheat were mapped using 71 doubled haploid (DH) lines derived from a Zhongyou 9507/CA9632 and SSR markers. Two QTLs QLpx.caas.lAL and QLpx.caas-4B were identified on chromosomes 1AL and 4BS, closely related to LOX activity. The SSR loci Xwmc312 and Xgwm251 were proven to be diagnostic and explained 13.4-25.2% of phenotypic variance the 1AL locus and 14.3-27.0% for the 4B locus across four environments. The SSR markers Xwmc312 and Xgwm251 were validated across 198 Chinese wheat cultivars and advanced lines and showed highly significant (P<0.01) association with LOX activity. We further established a multiplexed PCR with SSR marker combination Xwmc312/Xgwm251 to test these wheat cultivars and advanced lines. The results suggested that the marker combination Xwmc312/Xgwm251 is efficient and reliable for evaluating LOX activity and can be used in marker-assisted selection (MAS) targeting for flour color attributes to noodle and other wheat-based products.
3. The full-length genomic DNA sequence of TaLox-B1 and TaLox-B2 on chromosome 4BS and 5B, respectively, were characterized by in silico cloning and experimental validation. The cloned TaLox-B1 comprises seven exons and six introns, with 4,263 bp in total and an open reading frame (ORF) of 2,586 bp, and it encodes a LOX precursor peptide of 861 amino acids with a predicted molecular weight of-96.0 kDa. The genomic DNA sequence of TaLox-B2 comprised 3,325 base pairs, including six exons and five introns, and partial 3'UTR. The gene harbored an ORF of 2,595 bp, encoding a polypeptide of 864 amino acids with a predicted molecular weight of-96.4 kDa. TaLox-B1 and TaLOX1 were mapped on the homologous group 4, and TaLox-B2 and TaLOX2 were mapped on the homologous group 5 in wheat. Cluster analysis of TaLox-B1-TaLox-B2、TaLOXl、TaLOX2 and ZmLOX3 indicated that these LOX genes from the same homologous group showed the highest sequence identities in gDNA, cDNA and AA levels, respectively.
4. Functional markers for the LOX gene on chromosomes 4B based on single nucleotide polymorphism were developed and validated in common wheat. Two complementary dominant STS markers, LOX 16 and LOX 18, were developed based on the single nucleotide polymorphism (SNP) and amplifying 489-bp and 791-bp fragments in cultivars with higher and lower LOX activitives, respectively. The two markers were mapped on chromosome 4BS using a doubled haploid (DH) population derived from Zhongyou 9507/CA9632, and a set of Chinese Spring nullisomic-tetrasomic lines and ditelosomic line 4B. QTL analysis indicated that TaLox-B1 co-segregated with two functional markers and was closely linked to SSR marker Xgwm251 on chromosome 4BS with a genetic distance of 5.4 cM, explaining 15.9-26.2% of the phenotypic variance for LOX activity across four environments. Two complementary dominant functional markers LOX 16 and LOX 18, were validated on 198 Chinese wheat cultivars and advanced lines and showed highly significant (P<0.01) association with LOX activity. The results suggested that these two STS markers are closely related to LOX activity and could be used for the improvement of flour color attributes for noodle and other wheat-based products.
5. TaLox-B1 b in Northern China Plain Winter Wheat Rgion and the Yellow and the Huai River Valley Winter Wheat Region was dominant. TaLox-B1a was a superior allele for flour color and dough rheology. TaLox-Bla associated with higher LOX activitive had low frequency (34.3%) in these two regions. The frequency of TaLox-Bla in Yellow and Huai River Valley Winter Wheat Region (36.0%) were higher than those in Northern China Plain Winter Wheat Region (25.0%) (P0.01). These results indicated the preferred TaLox-B1a has not yet widely used in Chinese winter wheat regions. Therefore, it has a large potential to select new cultivars with high LOX activity.
引文
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