Increasing wheat yield is a long-term goal for wheat breeders around the world.Exploiting elite genetic resources and dissecting the genetic basis of important agronomic traits in wheat are the necessary methods for h...Increasing wheat yield is a long-term goal for wheat breeders around the world.Exploiting elite genetic resources and dissecting the genetic basis of important agronomic traits in wheat are the necessary methods for high-yield wheat breeding.This study evaluated nine crucial agronomic traits found in a natural population of 156 wheat varieties and77 landraces from Sichuan,China in seven environments over two years.The results of this investigation of agronomic traits showed that the landraces had more tillers and higher kernel numbers per spike (KNS),while the breeding varieties had higher thousand-kernel weight (TKW) and kernel weight per spike (KWS).The generalized heritability (H2) values of the nine agronomic traits varied from 0.74 to 0.95.Structure analysis suggested that the natural population could be divided into three groups using 43 198 single nucleotide polymorphism (SNP) markers from the wheat 55K SNP chip.A total of 67 quantitative trait loci (QTLs) were identified by the genome-wide association study (GWAS) analysis based on the Q+K method of a mixed linear model.Three important QTLs were analyzed in this study.Four haplotypes of QFTN.sicau-7BL.1 for fertile tillers number (FTN),three haplotypes of QKNS.sicau-1AL.2 for KNS,and four haplotypes of QTKW.sicau-3BS.1 for TKW were detected.FTN-Hap2,KNS-Hap1,and TKW-Hap2 were excellent haplotypes for each QTL based on the yield performance of 42 varieties in regional trials from 2002 to 2013.The varieties with all three haplotypes showed the highest yield compared to those with either two haplotypes or one haplotype.In addition,the KASP-AX-108866053 marker of QTL QKNS.sicau-1AL.2 was successfully distinguished between three haplotypes(or alleles) in 63 varieties based on the number of kernels per spike in regional trials between 2018 and 2021.These genetic loci and reliable makers can be applied in marker-assisted selection or map-based gene cloning for the genetic improvement of wheat yield.展开更多
High-molecular-weight glutenin subunits(HMW-GSs) are the most critical grain storage proteins that determine the unique processing qualities of wheat. Although it is a part of the superior HMW-GS pair(Dx5+Dy10), the c...High-molecular-weight glutenin subunits(HMW-GSs) are the most critical grain storage proteins that determine the unique processing qualities of wheat. Although it is a part of the superior HMW-GS pair(Dx5+Dy10), the contribution of the Dy10 subunit to wheat processing quality remains unclear. In this study, we elucidated the effect of Dy10 on wheat processing quality by generating and analyzing a deletion mutant(with the Dy10-null allele), and by elucidating the changes to wheat flour following the incorporation of purified Dy10. The Dy10-null allele was transcribed normally,but the Dy10 subunit was lacking. These findings implied that the Dy10-null allele reduced the glutenin:gliadin ratio and negatively affected dough strength(i.e., Zeleny sedimentation value, gluten index, and dough development and stability times) and the bread-making quality;however, it positively affected the biscuit-making quality. The incorporation of various amounts of purified Dy10 into wheat flour had a detrimental effect on biscuit-making quality. The results of this study demonstrate that the Dy10 subunit is essential for maintaining wheat dough strength. Furthermore, the Dy10-null allele may be exploited by soft wheat breeding programs.展开更多
Waterlogging is a widespread limiting factor for wheat production throughout the world. To identify quantitative trait loci （QTLs） associated with waterlogging tolerance at early stages of growth, survival rate （SR...Waterlogging is a widespread limiting factor for wheat production throughout the world. To identify quantitative trait loci （QTLs） associated with waterlogging tolerance at early stages of growth, survival rate （SR）, germination rate index （GRI）, leaf chlorophyll content index （CCI）, root length index （RLI）, plant height index （PHI）, root dry weight index （RDWI）, shoot dry weight index （SDWI）, and total dry weight index （DWI） were assessed using the International Triticeae Mapping Initiative （ITMI） population W7984/Opata85. Significant and positive correlations were detected for all traits in this population except RLI. A total of 32 QTLs were associated with waterlogging tolerance on all chromosomes except 3A, 3D, 4B, 5A, 5D, 6A, and 6D. Some of the QTLs explained large proportions of the phenotypic variance. One of these is the QTL for GRI on 7A, which explained 23.92% of the phenotypic variation. Of them, 22 alleles from the synthetic hexaploid wheat W7984 contributed positively. These results suggested that synthetic hexaploid wheat W7984 is an important genetic resource for waterlogging tolerance in wheat. These alleles conferring waterlogging tolerance at early stages of growth in wheat could be utilized in wheat breeding for improving waterlogging tolerance.展开更多
Aegiliops tauschii is classified into two subspecies： Ae. tauschii ssp. tauschii and Ae. tauschii ssp. strangulata. Novel genetic variations exist in Ae. tauschii ssp. tauschii that can be utilized in wheat improveme...Aegiliops tauschii is classified into two subspecies： Ae. tauschii ssp. tauschii and Ae. tauschii ssp. strangulata. Novel genetic variations exist in Ae. tauschii ssp. tauschii that can be utilized in wheat improvement. We synthesized a hexaploid wheat genotype（SHW-L1） by crossing an Ae. tauschii ssp. tauschii accession（AS60） with a tetraploid wheat genotype（AS2255）. A population consisting of 171 F8 recombinant inbred lines was developed from SHW-L1 and Chuanmai 32 to identify QTLs associated with agronomic traits. A new genetic map with high density was constructed and used to detect the QTLs for heading date, kernel width, spike length, spikelet number, and thousand kernel weight. A total of 30 putative QTLs were identified for five investigated traits. Thirteen QTLs were located on D genomes of SHW-L1, six of them showed positive effect on agronomic traits. Chromosome region flanked by wPt-6133–wPt-8134 on 2D carried five environment-independent QTLs. Each QTL accounted for more than 10% phenotypic variance. These QTLs were highly consistent across environments and should be used in wheat breeding.展开更多
It was helpful for the wheat improvement to evaluate the genetic resources of Triticum turgidum L. ssp. turgidum landraces. In this study, 68 turgidum landraces accessions, belonging to four geographic populations in ...It was helpful for the wheat improvement to evaluate the genetic resources of Triticum turgidum L. ssp. turgidum landraces. In this study, 68 turgidum landraces accessions, belonging to four geographic populations in China, were investigated by using EST-SSR markers. A total of 63 alleles were detected on 22 EST-SSR loci, and the number of alleles on each locus ranged from 1 to 5, with an average of 2.9. The results of the analysis of molecular variance （AMOVA） indicated that 92.5% of the total variations was attributed to the genetic variations within population, whereas only 7.5% variations among populations. Although the four populations had similar genetic diversity parameters, Sichuan population was yet distinguished from other populations when comparing the population samples in pairs. Significant correlations were detected by the statistic analysis among six genetic diversity parameters among each other. The selection difference between heterozygosty and homozygosty was also observed among different EST-SSR locus. The genetic similarity （GS） ranged from 0.18 to 0.98, with the mean of 0.72, and all accessions could be clustered into 7 groups. The dendrogram suggested that the genetic relationships among turgidum accessions evaluated by EST-SSR markers were unrelated to their geographic distributions. These results implied that turgidum landraces from China had the unique characters of genetic diversity.展开更多
Three coding sequences of gliadins genes, designed as Gli2_Dul, Gli2_Du2 and Gli2_Du3, were isolated from the genomic DNA of Triticum durum accessions CItr5083. Gli2_Dul and Gli2_Du2 contain 945 and 864 bp, encoding t...Three coding sequences of gliadins genes, designed as Gli2_Dul, Gli2_Du2 and Gli2_Du3, were isolated from the genomic DNA of Triticum durum accessions CItr5083. Gli2_Dul and Gli2_Du2 contain 945 and 864 bp, encoding the mature proteins with 314 and 287 amino acid residues, respectively. Gli2_Du3 is recognized as a pseudogene due to the stop codon occurring in the coding region. The pseudogenes, commonly occurring in gliadins family, are attributed to the single base change C→T. The amino acid sequences deduced from these gene sequences were characterized with the typical structure of α-gliadin proteins, including the toxic sequences （PSQQQP）. The peptide fraction PF（Y）PP（Q）is thought to be an extra unit of repetitive domain, slightly diverging from the previous report. Six cysteine residues were observed within two unique domains. Phylogenetic analysis showed Gli2_Du2 and Gli2_Du3 were closely related to the genes on chromosome 6A, whereas Gli2_Dul seems to be more homologous with the genes on chromosome 6B.展开更多
Micronutrient malnutrition affects over three billion people worldwide, especially women and children in developing countries. Increasing the bioavailable concentrations of essential elements in the edible portions of...Micronutrient malnutrition affects over three billion people worldwide, especially women and children in developing countries. Increasing the bioavailable concentrations of essential elements in the edible portions of crops is an effective resolution to address this issue. To determine the genetic factors controlling micronutrient concentration in wheat, the quantitative trait locus （QTL） analysis for iron, zinc, copper, manganese, and selenium concentrations in two recombinant inbred line populations was performed. In all, 39 QTLs for ifve micronutrient concentrations were identiifed in this study. Of these, 22 alleles from synthetic wheat SHW-L1 and seven alleles from the progeny line of the synthetic wheat Chuanmai 42 showed an increase in micronutrient concentrations. Five QTLs on chromosomes 2A, 3D, 4D, and 5B found in both the populations showed signiifcant phenotypic variation for 2-3 micronutrient concentrations. Our results might help understand the genetic control of micronutrient concentration and allow the utilization of genetic resources of synthetic hexaploid wheat for improving micronutrient efifciency of cultivated wheat by using molecular marker-assisted selection.展开更多
Synthetic hexaploid wheat(SHW),possesses numerous genes for drought that can help breeding for drought-tolerant wheat varieties.We evaluated 10 root traits at seedling stage in 111 F9 recombinant inbred lines derived ...Synthetic hexaploid wheat(SHW),possesses numerous genes for drought that can help breeding for drought-tolerant wheat varieties.We evaluated 10 root traits at seedling stage in 111 F9 recombinant inbred lines derived from a F2 population of a SHW line(SHW-L1)and a common wheat line,under normal(NC)and polyethylene glycol-simulated drought stress conditions(DC).We mapped quantitative trait loci(QTLs)for root traits using an enriched high-density genetic map containing 120370 single nucleotide polymorphisms(SNPs),733 diversity arrays technology markers(DArT)and 119 simple sequence repeats(SSRs).With four replicates per treatment,we identified 19 QTLs for root traits under NC and DC,and 12 of them could be consistently detected with three or four replicates.Two novel QTLs for root fresh weight and root diameter under NC explained 9 and 15.7%of the phenotypic variation respectively,and six novel QTLs for root fresh weight,the ratio of root water loss,total root surface area,number of root tips,and number of root forks under DC explained 8.5–14%of the phenotypic variation.Here seven of eight novel QTLs could be consistently detected with more than three replicates.Results provide essential information for fine-mapping QTLs related to drought tolerance that will facilitate breeding drought-tolerant wheat cultivars.展开更多
A novel HMW glutenin subunit gene 1Dy10. 1 was isolated and characterized from Xinjiang wheat （Triticum petropavlovskyi. Udacz. et Migusch） accession Daomai 2. The complete open reading frame （ORF） of 1Dy10. 1 was...A novel HMW glutenin subunit gene 1Dy10. 1 was isolated and characterized from Xinjiang wheat （Triticum petropavlovskyi. Udacz. et Migusch） accession Daomai 2. The complete open reading frame （ORF） of 1Dy10. 1 was 1 965 bp, encoding 655 amino acids. The numbers and distribution of cysteines in 1Dy 10.1 were similar to those of 1Dy10 and other y-type subunits. In the N-terminal of 1Dy 10.1, an amino acid was changed from L （leucine） to P （proline） at position 55. The repetitive domain of 1Dy10.1 differed from those of known HMW subunits by substitutions, insertions or/and deletions involving single or more amino acid residues. In the repetitive domain of subunit 1Dyl 0.1, the deletion of tripeptide GQQ in the consensus unit PGQGQQ resulted in the appearance of the motif PGQ that have not been observed in other known y-type HMW subunits. In comparison with the subunit 1Dy 12, a deletion of dipeptide GQ, which occurred in subunit IDy10, was also observed in subunit 1Dy10.1. The cloned IDylO. 1 gene had been successfully expressed in Escherichia coli, and the expressed protein had the identical mobility with the endogenous subunit IDy10.1 from seed.展开更多
The effects of auxins and media on callus induction from the mature and immature embryos of Chinese spring wheat （Triticum aestivum L.） varieties were investigated. It was found that genotype, medium, auxin source a...The effects of auxins and media on callus induction from the mature and immature embryos of Chinese spring wheat （Triticum aestivum L.） varieties were investigated. It was found that genotype, medium, auxin source and concentration had the significant effects on the induction of embryogenic callus, explants germination and the increment of callus fresh weight. For immature embryos cultured on MS medium, 2 mg L^-1 of 2, 4-D was optimal, and the highest frequency of embryogenic callus （33.50%） was observed. For the mature embryos on N6 medium, 4 mg L^-1 of 2, 4-D was optimal. The frequency of embryogenic callus and increment of callus fresh weight on 2, 4, 5-T media were higher than those on 2, 4-D media, and in the presence of 2, 4, 5-T the precocious germination of explants for all genotypes were significantly suppressed. These results indicated that 2, 4, 5-T was superior to 2, 4-D and NAA in the culture of immature embryos. This is the first report about the effect of 2, 4, 5-T and NAA on wheat tissue culture, particularly in comparison with 2, 4-D in detail.展开更多
The spikelet number per spike(SNS)contributes greatly to grain yield in wheat.Identifying various genes that control wheat SNS is vital for yield improvement.This study used a recombinant inbred line population genoty...The spikelet number per spike(SNS)contributes greatly to grain yield in wheat.Identifying various genes that control wheat SNS is vital for yield improvement.This study used a recombinant inbred line population genotyped by the Wheat55K single-nucleotide polymorphism array to identify two major and stably expressed quantitative trait loci(QTLs)for SNS.One of them(QSns.sau-2SY-2D.1)was reported previously,while the other(QSns.sau-2SY-7A)was newly detected and further analyzed in this study.QSns.sau-2SY-7A had a high LOD value ranging from 4.46 to 16.00 and explained 10.21-40.78%of the phenotypic variances.QSns.sau-2SY-7A was flanked by the markers AX-110518554 and AX-110094527 in a 4.75-cM interval on chromosome arm 7AL.The contributions and interactions of both major QTLs were further analyzed and discussed.The effect of QSns.sau-2SY-7A was successfully validated by developing a tightly linked kompetitive allele specific PCR marker in an F_(2:3) population and a panel of 101 high-generation breeding wheat lines.Furthermore,several genes including the previously reported WHEAT ORTHOLOG OF APO1(WAPO1),an ortholog of the rice gene ABERRANT PANICLE ORGANIZATION 1(APO1)related to SNS,were predicted in the interval of QSns.sau-2SY-7A.In summary,these results revealed the genetic basis of the multi-spikelet genotype of wheat line 20828 and will facilitate subsequent fine mapping and breeding utilization of the major QTLs.展开更多
Uppermost-internode diameter(UID)is a key morphological trait associated with spike development and yield potential in wheat.Our understanding of its genetic basis remains largely unknown.Here,quantitative trait loci(...Uppermost-internode diameter(UID)is a key morphological trait associated with spike development and yield potential in wheat.Our understanding of its genetic basis remains largely unknown.Here,quantitative trait loci(QTLs)for UID with high-density genetic maps were identified in five wheat recombinant inbred line(RIL)populations.In total,25 QTLs for UID were detected in five RIL populations,and they were located on chromosomes 1A,1D(3 QTL),2B(2),2D(3),3B,3D,4A,4B(3),4D,5A(5),5B(2),6B,and 7D.Of them,five major and stable QTLs(QUid.sau-2CN-1D.1,QUid.sau-2SY-1D,QUid.sau-QZ-2D,QUid.sau-SC-3D,and QUid.sau-AS-4 B)were identified from each of the five RIL populations in multiple environments.QUid.sau-2CN-1D.1,QUid.sau-2SY-1D and QUid.sau-SC-3D are novel QTLs.Kompetitive Allele Specific PCR(KASP)markers tightly linked to them were further investigated for developing near-isogenic lines(NILs)carrying the major loci.Furthermore,candidate genes at these intervals harboring major and stable QTLs were predicted,and they were associated with plant development and water transportation in most cases.Comparison of physical locations of the identified QTL on the‘Chinese Spring’reference genome showed that several QTLs including two major ones,QUid.sau-2CN-1D.1 and QUid.sau-2 SY-1 D,are likely allelic confirming their validity and effectiveness.The significant relationships detected between UID and other agronomic traits and a proper UID were discussed.Collectively,our results dissected the underlying genetic basis for UID in wheat and laid a foundation for further fine mapping and map-based cloning of these QTLs.展开更多
In this paper, the traits of immature embryo culture from 45 genotypes of bread wheat were investigated in detail, and trait correlation analysis in combination with histomorphological examination was used to study th...In this paper, the traits of immature embryo culture from 45 genotypes of bread wheat were investigated in detail, and trait correlation analysis in combination with histomorphological examination was used to study the cytological origin of flower as well as the correlation between it and vegetative shoot development from the calli. The rate of floral development in the subculture was highly significantly and positively related to that of the differentiation culture, which reveals that the genetic system of floral development from immature embryo-derived calli in wheat was highly correspondent and stable. The rate of floral development was related neither to the traits in the dedifferentiation culture, nor to the vegetative shoot formation from the total induced calli, but significantly and positively related to that of the calli without leaf-like green spots in the differentiation culture. These findings indicate the partial correlation as well as differentiation between flower and vegetative shoot development. The floral development might only be associated with the genetic system controlling organ regeneration, while vegetative shoots formed from the two ways, including somatic embryogenesis and organ regeneration.展开更多
The successful worldwide cultivation of hexaploid wheat in a diverse range of environments is because of, in part, breeding and selection for appropriate heading date. To adjust and fine-tune the heading time of hexap...The successful worldwide cultivation of hexaploid wheat in a diverse range of environments is because of, in part, breeding and selection for appropriate heading date. To adjust and fine-tune the heading time of hexaploid wheat to particular geographical regions and specific environment within these, there is an urgent need to evaluate and use alternative alleles for heading time. Aegilops tauschii, the donor species of D-genome of hexaploid wheat, has a wide geographic distribution. The present study revealed a wide variation for heading time among 56 Ae. tauschii accessions. All the accessions with short heading dates belonged to the ssp. tauschii, whereas most of ssp. strangulata accessions showed very long heading date. The heading date was also related to distribution of this species. The monotelosomic and monosomic analysis of a synthetic hexaploid wheat showed that chromosome 2D derived from ssp. tauschii accession AS60 had a major effect on promoting heading time with a reduction of more than 5 days. It is postulated that this Ae. tauschii genotype possess the allele Ppd-D^t1 responsible for the insensitivity to photoperiod. This allele is probably different from Ppd-D1 existing in hexaploid wheat. The new allele Ppd-D^t1 derived from Ae. tauschii might be used as a source for hexaploid wheat breeding on photoperiod response.展开更多
Understanding the genetic basis of quality-related traits contributes to the improvement of grain protein concentration(GPC),grain starch concentration(GSC),and wet gluten concentration(WGC)in wheat.In this study,a ge...Understanding the genetic basis of quality-related traits contributes to the improvement of grain protein concentration(GPC),grain starch concentration(GSC),and wet gluten concentration(WGC)in wheat.In this study,a genome-wide association study(GWAS)based on a mixed linear model(MLM)was performed on 236 wheat accessions,including 160 cultivars and 76 landraces,using a 55K single nucleotide polymorphism(SNP)array in multiple environments.A total of 12 stable QTL/SNPs that control different quality traits in this populations in at least two environments under stripe rust stress were identified.Among these 12,three,seven and two QTLs associated with GPC,GSC and WGC were characterized,respectively,and they were located on chromosomes(chr)1B,1D,2A,2B,2D,3B,3D,5D,and 7D with the phenotypic variation explained(PVE)ranging from 4.2 to 10.7%.Compared with the previously reported QTLs/genes,five QTLs(QGsc.sicau-1BL,QGsc.sicau-1DS,QGsc.sicau-2DL.1,QGsc.sicau-2DL.2,and QWgc.sicau-5DL)were potentially novel.KASP markers for the SNPs AX-108770574 and AX-108791420 on chr5D associated with wet gluten concentration were successfully developed.The phenotypes of the cultivars containing the A-allele in AX-108770574and the T-allele in AX-108791420 were extremely significantly(P<0.01)higher than those of the landraces containing the G-or C-allele with respect to the wet gluten concentration in each of the environments.The KASP markers developed and validated in this study could be utilized in molecular breeding aimed at improving the quality of wheat.展开更多
Starch is the major carbohydrate in oat （Avena sativa L.） and starch formation requires the coordinated actions of several synthesis enzymes. In this study, the granule morphology, composition and physicochemical pr...Starch is the major carbohydrate in oat （Avena sativa L.） and starch formation requires the coordinated actions of several synthesis enzymes. In this study, the granule morphology, composition and physicochemical properties of oat starch, as well as the expressions of starch synthesis genes were investigated during oat endosperm development. Under the scanning electron microscopy （SEM）, we observed that the unique compound granules were developed in oat endosperms at 10 days post anthesis （DPA） and then fragmented into irregular or polygonal simple granules from 12 DPA until seed maturity. The amylose content, branch chain length of degree of polymerization （DP=13-24）, gelatinization temperature and percentage of retrogradation were gradually increased during the endosperm development; whereas the distribution of short chains （DP=6-12） were gradually decreased. The relative expressions of 4 classes of 13 starch synthesis genes characterized in this study indicated that three expression pattern groups were significantly different among gene classes as well as among varied isoforms, in which the first group of starch synthesis genes may play a key role on the initiation of starch synthesis in oat endosperms.展开更多
The bread wheat genome harbors a high content of repetitive DNA,which is amenable to detection and characterization using fluorescence in situ hybridization(FISH)karyotyping.An integrated genetic map was derived from ...The bread wheat genome harbors a high content of repetitive DNA,which is amenable to detection and characterization using fluorescence in situ hybridization(FISH)karyotyping.An integrated genetic map was derived from a recombinant inbred population bred from a cross between a synthetic hexaploid wheat and a commercial Chinese bread wheat cultivar,based on 28 variable FISH sites and>150000 single nucleotide polymorphism(SNP)loci.The majority(20/28)of the variable FISH sites were physically located within a chromosomal region consistent with the genetic location inferred from that of their co-segregating SNP loci.The eight exceptions reflected the presence of either a translocation(1 R/1 B,1 A/7 A)or a presumptive intra-chromosomal inversion(4 A).For eight out of the nine FISH sites detected on the Chinese Spring(CS)karyotype,there was a good match with the reference genome sequence,indicating that the most recent assembly has dealt well with the problem of placing tandem repeats.The integrated genetic map produced for wheat is informative as to the location of blocks of tandemly repeated DNA and can aid in improving the quality of the genome sequence assembly in regions surrounding these blocks.展开更多
Novel LMW-GS （low molecular weight glutenin subunit）,α- and γ-gliadin from Triticum macha accessions were characterized via genomic PCR, which can do favor to improve the wheat quality. The complete coding regions...Novel LMW-GS （low molecular weight glutenin subunit）,α- and γ-gliadin from Triticum macha accessions were characterized via genomic PCR, which can do favor to improve the wheat quality. The complete coding regions of two α-gliadin, two γ-gliadin and two LMW-GS gene sequences, which designed as Gli-Mal, Gli-Ma2, Gli-Mrl, Gli-Mr2, Glu-LM1 and Glu-LM2, encoded the mature proteins with 307, 241, 348, 302, 474 and 377 amino acid residues, respectively. Gli-Mal and Gli-Ma2 were recognized as pseudogenes due to the in-frame stop codons. The amino acid sequences deduced from these gene sequences were characterized with the typical structure of α- or γ-gliadin or LMW- m type proteins with the exception of Gli-Ma2. Phylogenetic analysis showed Gli-Mal was closely related to those from T. aestivum, whereas Gli-Ma2 seemed to be more homologous with the gene sequences from Dasypyrum breviaristatum. Gli-Mrl was closely related to those from T. turgidum ssp. dicoccoides, while Gli-Mr2 was the nearest to those from T. aestivum. Glu-LM1 was closely related to those from Aegilops tauschii, whereas GIu-LM2 seemed to be more homologous with those from T. durum.展开更多
基金supported by the Sichuan Science and Technology Program , China ( 2022ZDZX0014 and 2021YFYZ0002)the Plan of Tianfu Qingcheng of Sichuan Province, China。
文摘Increasing wheat yield is a long-term goal for wheat breeders around the world.Exploiting elite genetic resources and dissecting the genetic basis of important agronomic traits in wheat are the necessary methods for high-yield wheat breeding.This study evaluated nine crucial agronomic traits found in a natural population of 156 wheat varieties and77 landraces from Sichuan,China in seven environments over two years.The results of this investigation of agronomic traits showed that the landraces had more tillers and higher kernel numbers per spike (KNS),while the breeding varieties had higher thousand-kernel weight (TKW) and kernel weight per spike (KWS).The generalized heritability (H2) values of the nine agronomic traits varied from 0.74 to 0.95.Structure analysis suggested that the natural population could be divided into three groups using 43 198 single nucleotide polymorphism (SNP) markers from the wheat 55K SNP chip.A total of 67 quantitative trait loci (QTLs) were identified by the genome-wide association study (GWAS) analysis based on the Q+K method of a mixed linear model.Three important QTLs were analyzed in this study.Four haplotypes of QFTN.sicau-7BL.1 for fertile tillers number (FTN),three haplotypes of QKNS.sicau-1AL.2 for KNS,and four haplotypes of QTKW.sicau-3BS.1 for TKW were detected.FTN-Hap2,KNS-Hap1,and TKW-Hap2 were excellent haplotypes for each QTL based on the yield performance of 42 varieties in regional trials from 2002 to 2013.The varieties with all three haplotypes showed the highest yield compared to those with either two haplotypes or one haplotype.In addition,the KASP-AX-108866053 marker of QTL QKNS.sicau-1AL.2 was successfully distinguished between three haplotypes(or alleles) in 63 varieties based on the number of kernels per spike in regional trials between 2018 and 2021.These genetic loci and reliable makers can be applied in marker-assisted selection or map-based gene cloning for the genetic improvement of wheat yield.
基金supported by the National Natural Science Foundation of China (31971939, 32072054, and 31901961)the Science and Technology Department of Sichuan Province, China (2019YFH0066 and 2020YFH0150)the Designing Future Wheat Strategic Program of the UK (BB/P016855/1)。
文摘High-molecular-weight glutenin subunits(HMW-GSs) are the most critical grain storage proteins that determine the unique processing qualities of wheat. Although it is a part of the superior HMW-GS pair(Dx5+Dy10), the contribution of the Dy10 subunit to wheat processing quality remains unclear. In this study, we elucidated the effect of Dy10 on wheat processing quality by generating and analyzing a deletion mutant(with the Dy10-null allele), and by elucidating the changes to wheat flour following the incorporation of purified Dy10. The Dy10-null allele was transcribed normally,but the Dy10 subunit was lacking. These findings implied that the Dy10-null allele reduced the glutenin:gliadin ratio and negatively affected dough strength(i.e., Zeleny sedimentation value, gluten index, and dough development and stability times) and the bread-making quality;however, it positively affected the biscuit-making quality. The incorporation of various amounts of purified Dy10 into wheat flour had a detrimental effect on biscuit-making quality. The results of this study demonstrate that the Dy10 subunit is essential for maintaining wheat dough strength. Furthermore, the Dy10-null allele may be exploited by soft wheat breeding programs.
基金supported by the National Basic Research Program of China (2011CB100100)the National High-Tech R&D Program of China (2011AA100103)the National Natural Science Foundation of China (31230053 and 31171556)
文摘Waterlogging is a widespread limiting factor for wheat production throughout the world. To identify quantitative trait loci （QTLs） associated with waterlogging tolerance at early stages of growth, survival rate （SR）, germination rate index （GRI）, leaf chlorophyll content index （CCI）, root length index （RLI）, plant height index （PHI）, root dry weight index （RDWI）, shoot dry weight index （SDWI）, and total dry weight index （DWI） were assessed using the International Triticeae Mapping Initiative （ITMI） population W7984/Opata85. Significant and positive correlations were detected for all traits in this population except RLI. A total of 32 QTLs were associated with waterlogging tolerance on all chromosomes except 3A, 3D, 4B, 5A, 5D, 6A, and 6D. Some of the QTLs explained large proportions of the phenotypic variance. One of these is the QTL for GRI on 7A, which explained 23.92% of the phenotypic variation. Of them, 22 alleles from the synthetic hexaploid wheat W7984 contributed positively. These results suggested that synthetic hexaploid wheat W7984 is an important genetic resource for waterlogging tolerance in wheat. These alleles conferring waterlogging tolerance at early stages of growth in wheat could be utilized in wheat breeding for improving waterlogging tolerance.
基金supported by the National Natural Science Foundation of China(31171556,31171555,31230053)the National High-Tech R&D Program of China(2011AA100103-02)the Key Technologies R&D Program of China during the 12th Five-Year Plan period(2013BAD01B02-9)
文摘Aegiliops tauschii is classified into two subspecies： Ae. tauschii ssp. tauschii and Ae. tauschii ssp. strangulata. Novel genetic variations exist in Ae. tauschii ssp. tauschii that can be utilized in wheat improvement. We synthesized a hexaploid wheat genotype（SHW-L1） by crossing an Ae. tauschii ssp. tauschii accession（AS60） with a tetraploid wheat genotype（AS2255）. A population consisting of 171 F8 recombinant inbred lines was developed from SHW-L1 and Chuanmai 32 to identify QTLs associated with agronomic traits. A new genetic map with high density was constructed and used to detect the QTLs for heading date, kernel width, spike length, spikelet number, and thousand kernel weight. A total of 30 putative QTLs were identified for five investigated traits. Thirteen QTLs were located on D genomes of SHW-L1, six of them showed positive effect on agronomic traits. Chromosome region flanked by wPt-6133–wPt-8134 on 2D carried five environment-independent QTLs. Each QTL accounted for more than 10% phenotypic variance. These QTLs were highly consistent across environments and should be used in wheat breeding.
基金the National High Technology Research and Development Program of China (863 Program,2006AA10Z179 and 2006AA10Z1F8)the National Excellent Doctoral Dissertation of China (200357 and 200458)the Key Technologies R&D Program of China (2006BAD01A02-23 and 2006BAD13B02)
文摘It was helpful for the wheat improvement to evaluate the genetic resources of Triticum turgidum L. ssp. turgidum landraces. In this study, 68 turgidum landraces accessions, belonging to four geographic populations in China, were investigated by using EST-SSR markers. A total of 63 alleles were detected on 22 EST-SSR loci, and the number of alleles on each locus ranged from 1 to 5, with an average of 2.9. The results of the analysis of molecular variance （AMOVA） indicated that 92.5% of the total variations was attributed to the genetic variations within population, whereas only 7.5% variations among populations. Although the four populations had similar genetic diversity parameters, Sichuan population was yet distinguished from other populations when comparing the population samples in pairs. Significant correlations were detected by the statistic analysis among six genetic diversity parameters among each other. The selection difference between heterozygosty and homozygosty was also observed among different EST-SSR locus. The genetic similarity （GS） ranged from 0.18 to 0.98, with the mean of 0.72, and all accessions could be clustered into 7 groups. The dendrogram suggested that the genetic relationships among turgidum accessions evaluated by EST-SSR markers were unrelated to their geographic distributions. These results implied that turgidum landraces from China had the unique characters of genetic diversity.
基金supported by the National High Technology Research and Development Program of China(863 Program,2006AA10Z179,2006AA10ZEF87)National Natural Science Foundation of China(30300219 and 30571163)+2 种基金the Foundation for the Authors of National Excellent Doctoral Dissertation of China(200357 and 200458)supported by the Program for New Century Excellent Talents in Universifies of Chinasupported by the Program for Changjiang Scholars and Innovative Research Teams in Universities of China(IRT0453).
文摘Three coding sequences of gliadins genes, designed as Gli2_Dul, Gli2_Du2 and Gli2_Du3, were isolated from the genomic DNA of Triticum durum accessions CItr5083. Gli2_Dul and Gli2_Du2 contain 945 and 864 bp, encoding the mature proteins with 314 and 287 amino acid residues, respectively. Gli2_Du3 is recognized as a pseudogene due to the stop codon occurring in the coding region. The pseudogenes, commonly occurring in gliadins family, are attributed to the single base change C→T. The amino acid sequences deduced from these gene sequences were characterized with the typical structure of α-gliadin proteins, including the toxic sequences （PSQQQP）. The peptide fraction PF（Y）PP（Q）is thought to be an extra unit of repetitive domain, slightly diverging from the previous report. Six cysteine residues were observed within two unique domains. Phylogenetic analysis showed Gli2_Du2 and Gli2_Du3 were closely related to the genes on chromosome 6A, whereas Gli2_Dul seems to be more homologous with the genes on chromosome 6B.
基金supported by the National Natural Science Foundation of China (31301318, 31230053 and 31171556)the National Basic Research Program of China (2011CB100100)
文摘Micronutrient malnutrition affects over three billion people worldwide, especially women and children in developing countries. Increasing the bioavailable concentrations of essential elements in the edible portions of crops is an effective resolution to address this issue. To determine the genetic factors controlling micronutrient concentration in wheat, the quantitative trait locus （QTL） analysis for iron, zinc, copper, manganese, and selenium concentrations in two recombinant inbred line populations was performed. In all, 39 QTLs for ifve micronutrient concentrations were identiifed in this study. Of these, 22 alleles from synthetic wheat SHW-L1 and seven alleles from the progeny line of the synthetic wheat Chuanmai 42 showed an increase in micronutrient concentrations. Five QTLs on chromosomes 2A, 3D, 4D, and 5B found in both the populations showed signiifcant phenotypic variation for 2-3 micronutrient concentrations. Our results might help understand the genetic control of micronutrient concentration and allow the utilization of genetic resources of synthetic hexaploid wheat for improving micronutrient efifciency of cultivated wheat by using molecular marker-assisted selection.
基金supported by the National Natural Science Foundation of China(31771794,91731305 and 31560388)the outstanding Youth Foundation of the Department of Science and Technology of Sichuan Province,China(2016JQ0040)+1 种基金the Key Technology Research and Development Program of the Department of Science and Technology of Sichuan Province,China(2016NZ0057)the International Science&Technology Cooperation Program of the Bureau of Science and Technology of Chengdu,China(2015DFA306002015-GH03-00008-HZ)。
文摘Synthetic hexaploid wheat(SHW),possesses numerous genes for drought that can help breeding for drought-tolerant wheat varieties.We evaluated 10 root traits at seedling stage in 111 F9 recombinant inbred lines derived from a F2 population of a SHW line(SHW-L1)and a common wheat line,under normal(NC)and polyethylene glycol-simulated drought stress conditions(DC).We mapped quantitative trait loci(QTLs)for root traits using an enriched high-density genetic map containing 120370 single nucleotide polymorphisms(SNPs),733 diversity arrays technology markers(DArT)and 119 simple sequence repeats(SSRs).With four replicates per treatment,we identified 19 QTLs for root traits under NC and DC,and 12 of them could be consistently detected with three or four replicates.Two novel QTLs for root fresh weight and root diameter under NC explained 9 and 15.7%of the phenotypic variation respectively,and six novel QTLs for root fresh weight,the ratio of root water loss,total root surface area,number of root tips,and number of root forks under DC explained 8.5–14%of the phenotypic variation.Here seven of eight novel QTLs could be consistently detected with more than three replicates.Results provide essential information for fine-mapping QTLs related to drought tolerance that will facilitate breeding drought-tolerant wheat cultivars.
基金This work was supported by the National High Technology Research and Development Program of China （2003AA207100）the grants from the National Natural Science Foundation of China （30300219, 30370882, and 30571163） the Foundation for the Author of National Excellent Doctoral Dissertation of China （200357 and 200458）.
文摘A novel HMW glutenin subunit gene 1Dy10. 1 was isolated and characterized from Xinjiang wheat （Triticum petropavlovskyi. Udacz. et Migusch） accession Daomai 2. The complete open reading frame （ORF） of 1Dy10. 1 was 1 965 bp, encoding 655 amino acids. The numbers and distribution of cysteines in 1Dy 10.1 were similar to those of 1Dy10 and other y-type subunits. In the N-terminal of 1Dy 10.1, an amino acid was changed from L （leucine） to P （proline） at position 55. The repetitive domain of 1Dy10.1 differed from those of known HMW subunits by substitutions, insertions or/and deletions involving single or more amino acid residues. In the repetitive domain of subunit 1Dyl 0.1, the deletion of tripeptide GQQ in the consensus unit PGQGQQ resulted in the appearance of the motif PGQ that have not been observed in other known y-type HMW subunits. In comparison with the subunit 1Dy 12, a deletion of dipeptide GQ, which occurred in subunit IDy10, was also observed in subunit 1Dy10.1. The cloned IDylO. 1 gene had been successfully expressed in Escherichia coli, and the expressed protein had the identical mobility with the endogenous subunit IDy10.1 from seed.
基金the National High Tech R&D Program of China(863 Program,2003AA207100)the Foundation for the Author of National Excellent Doctoral Dissertation of P R China(200357) the Program forChangjiang Scholars and Innovative Research Team inUniversity(IRT0453).
文摘The effects of auxins and media on callus induction from the mature and immature embryos of Chinese spring wheat （Triticum aestivum L.） varieties were investigated. It was found that genotype, medium, auxin source and concentration had the significant effects on the induction of embryogenic callus, explants germination and the increment of callus fresh weight. For immature embryos cultured on MS medium, 2 mg L^-1 of 2, 4-D was optimal, and the highest frequency of embryogenic callus （33.50%） was observed. For the mature embryos on N6 medium, 4 mg L^-1 of 2, 4-D was optimal. The frequency of embryogenic callus and increment of callus fresh weight on 2, 4, 5-T media were higher than those on 2, 4-D media, and in the presence of 2, 4, 5-T the precocious germination of explants for all genotypes were significantly suppressed. These results indicated that 2, 4, 5-T was superior to 2, 4-D and NAA in the culture of immature embryos. This is the first report about the effect of 2, 4, 5-T and NAA on wheat tissue culture, particularly in comparison with 2, 4-D in detail.
基金supported by the projects from the Applied Basic Research Programs of Science and Technology Department of Sichuan Province, China (2020YJ0140 and 2021YJ0503)the International Science and Technology Cooperation and Exchanges Program of Science and Technology Department of Sichuan Province, China (2021YFH0083 and 2022YFH0053)+1 种基金the National Natural Science Foundation of China (31971937 and 31970243)the Key Projects of Scientific and Technological Activities for Overseas Students of Sichuan Province, China
文摘The spikelet number per spike(SNS)contributes greatly to grain yield in wheat.Identifying various genes that control wheat SNS is vital for yield improvement.This study used a recombinant inbred line population genotyped by the Wheat55K single-nucleotide polymorphism array to identify two major and stably expressed quantitative trait loci(QTLs)for SNS.One of them(QSns.sau-2SY-2D.1)was reported previously,while the other(QSns.sau-2SY-7A)was newly detected and further analyzed in this study.QSns.sau-2SY-7A had a high LOD value ranging from 4.46 to 16.00 and explained 10.21-40.78%of the phenotypic variances.QSns.sau-2SY-7A was flanked by the markers AX-110518554 and AX-110094527 in a 4.75-cM interval on chromosome arm 7AL.The contributions and interactions of both major QTLs were further analyzed and discussed.The effect of QSns.sau-2SY-7A was successfully validated by developing a tightly linked kompetitive allele specific PCR marker in an F_(2:3) population and a panel of 101 high-generation breeding wheat lines.Furthermore,several genes including the previously reported WHEAT ORTHOLOG OF APO1(WAPO1),an ortholog of the rice gene ABERRANT PANICLE ORGANIZATION 1(APO1)related to SNS,were predicted in the interval of QSns.sau-2SY-7A.In summary,these results revealed the genetic basis of the multi-spikelet genotype of wheat line 20828 and will facilitate subsequent fine mapping and breeding utilization of the major QTLs.
基金supported by the projects from the National Natural Science Foundation of China(31971937 and 31970243)the Key Projects of Scientific and Technological Activities for Overseas Students of Sichuan Province,Chinathe Applied Basic Research Programs of Science and Technology Department of Sichuan Province,China(2020YJ0140)。
文摘Uppermost-internode diameter(UID)is a key morphological trait associated with spike development and yield potential in wheat.Our understanding of its genetic basis remains largely unknown.Here,quantitative trait loci(QTLs)for UID with high-density genetic maps were identified in five wheat recombinant inbred line(RIL)populations.In total,25 QTLs for UID were detected in five RIL populations,and they were located on chromosomes 1A,1D(3 QTL),2B(2),2D(3),3B,3D,4A,4B(3),4D,5A(5),5B(2),6B,and 7D.Of them,five major and stable QTLs(QUid.sau-2CN-1D.1,QUid.sau-2SY-1D,QUid.sau-QZ-2D,QUid.sau-SC-3D,and QUid.sau-AS-4 B)were identified from each of the five RIL populations in multiple environments.QUid.sau-2CN-1D.1,QUid.sau-2SY-1D and QUid.sau-SC-3D are novel QTLs.Kompetitive Allele Specific PCR(KASP)markers tightly linked to them were further investigated for developing near-isogenic lines(NILs)carrying the major loci.Furthermore,candidate genes at these intervals harboring major and stable QTLs were predicted,and they were associated with plant development and water transportation in most cases.Comparison of physical locations of the identified QTL on the‘Chinese Spring’reference genome showed that several QTLs including two major ones,QUid.sau-2CN-1D.1 and QUid.sau-2 SY-1 D,are likely allelic confirming their validity and effectiveness.The significant relationships detected between UID and other agronomic traits and a proper UID were discussed.Collectively,our results dissected the underlying genetic basis for UID in wheat and laid a foundation for further fine mapping and map-based cloning of these QTLs.
基金the National Natural Science Foundation of China(39970445)Program for Changjiang Scholars and Innovativc Rescarch Team in University(PCSIRT,IRT0453)+1 种基金 the Natural Science Foundation of Education Committee Scienceand Technology Committee in Sichuan Province of China.
文摘In this paper, the traits of immature embryo culture from 45 genotypes of bread wheat were investigated in detail, and trait correlation analysis in combination with histomorphological examination was used to study the cytological origin of flower as well as the correlation between it and vegetative shoot development from the calli. The rate of floral development in the subculture was highly significantly and positively related to that of the differentiation culture, which reveals that the genetic system of floral development from immature embryo-derived calli in wheat was highly correspondent and stable. The rate of floral development was related neither to the traits in the dedifferentiation culture, nor to the vegetative shoot formation from the total induced calli, but significantly and positively related to that of the calli without leaf-like green spots in the differentiation culture. These findings indicate the partial correlation as well as differentiation between flower and vegetative shoot development. The floral development might only be associated with the genetic system controlling organ regeneration, while vegetative shoots formed from the two ways, including somatic embryogenesis and organ regeneration.
基金supported by the New Century Excellent Talents in University (NCET-04-0908)Changjiang Scholars and Innovative Research Team in University (IRT0453) of Ministry of Education of China+2 种基金National Natural Science Foundation of China(30700495)Key Technologies R&D Program of China(2006BAD13B02)Education Department,and Scienceand Technology Department of Sichuan Province(07ZZ025)
文摘The successful worldwide cultivation of hexaploid wheat in a diverse range of environments is because of, in part, breeding and selection for appropriate heading date. To adjust and fine-tune the heading time of hexaploid wheat to particular geographical regions and specific environment within these, there is an urgent need to evaluate and use alternative alleles for heading time. Aegilops tauschii, the donor species of D-genome of hexaploid wheat, has a wide geographic distribution. The present study revealed a wide variation for heading time among 56 Ae. tauschii accessions. All the accessions with short heading dates belonged to the ssp. tauschii, whereas most of ssp. strangulata accessions showed very long heading date. The heading date was also related to distribution of this species. The monotelosomic and monosomic analysis of a synthetic hexaploid wheat showed that chromosome 2D derived from ssp. tauschii accession AS60 had a major effect on promoting heading time with a reduction of more than 5 days. It is postulated that this Ae. tauschii genotype possess the allele Ppd-D^t1 responsible for the insensitivity to photoperiod. This allele is probably different from Ppd-D1 existing in hexaploid wheat. The new allele Ppd-D^t1 derived from Ae. tauschii might be used as a source for hexaploid wheat breeding on photoperiod response.
基金supported by the National Key Research and Development Program of China(2017YFD0100900,2016YFD0102000 and 2016YFD0100100)the International Science and Technology Cooperation and Exchanges Programs of Science and Technology Department of Sichuan Province,China(2019YFH0063)the Sichuan Science and Technology Program,China(2022ZDZX0014)。
文摘Understanding the genetic basis of quality-related traits contributes to the improvement of grain protein concentration(GPC),grain starch concentration(GSC),and wet gluten concentration(WGC)in wheat.In this study,a genome-wide association study(GWAS)based on a mixed linear model(MLM)was performed on 236 wheat accessions,including 160 cultivars and 76 landraces,using a 55K single nucleotide polymorphism(SNP)array in multiple environments.A total of 12 stable QTL/SNPs that control different quality traits in this populations in at least two environments under stripe rust stress were identified.Among these 12,three,seven and two QTLs associated with GPC,GSC and WGC were characterized,respectively,and they were located on chromosomes(chr)1B,1D,2A,2B,2D,3B,3D,5D,and 7D with the phenotypic variation explained(PVE)ranging from 4.2 to 10.7%.Compared with the previously reported QTLs/genes,five QTLs(QGsc.sicau-1BL,QGsc.sicau-1DS,QGsc.sicau-2DL.1,QGsc.sicau-2DL.2,and QWgc.sicau-5DL)were potentially novel.KASP markers for the SNPs AX-108770574 and AX-108791420 on chr5D associated with wet gluten concentration were successfully developed.The phenotypes of the cultivars containing the A-allele in AX-108770574and the T-allele in AX-108791420 were extremely significantly(P<0.01)higher than those of the landraces containing the G-or C-allele with respect to the wet gluten concentration in each of the environments.The KASP markers developed and validated in this study could be utilized in molecular breeding aimed at improving the quality of wheat.
基金supported by the National Natural Science Foundation of China (31230053)the Ministry of Education (MOE) of China and Agriculture and Agri-Food Canada (AAFC) Ph D Research Program
文摘Starch is the major carbohydrate in oat （Avena sativa L.） and starch formation requires the coordinated actions of several synthesis enzymes. In this study, the granule morphology, composition and physicochemical properties of oat starch, as well as the expressions of starch synthesis genes were investigated during oat endosperm development. Under the scanning electron microscopy （SEM）, we observed that the unique compound granules were developed in oat endosperms at 10 days post anthesis （DPA） and then fragmented into irregular or polygonal simple granules from 12 DPA until seed maturity. The amylose content, branch chain length of degree of polymerization （DP=13-24）, gelatinization temperature and percentage of retrogradation were gradually increased during the endosperm development; whereas the distribution of short chains （DP=6-12） were gradually decreased. The relative expressions of 4 classes of 13 starch synthesis genes characterized in this study indicated that three expression pattern groups were significantly different among gene classes as well as among varied isoforms, in which the first group of starch synthesis genes may play a key role on the initiation of starch synthesis in oat endosperms.
基金financially supported by the National Key Research and Development Program of China(2016YFD0102000)。
文摘The bread wheat genome harbors a high content of repetitive DNA,which is amenable to detection and characterization using fluorescence in situ hybridization(FISH)karyotyping.An integrated genetic map was derived from a recombinant inbred population bred from a cross between a synthetic hexaploid wheat and a commercial Chinese bread wheat cultivar,based on 28 variable FISH sites and>150000 single nucleotide polymorphism(SNP)loci.The majority(20/28)of the variable FISH sites were physically located within a chromosomal region consistent with the genetic location inferred from that of their co-segregating SNP loci.The eight exceptions reflected the presence of either a translocation(1 R/1 B,1 A/7 A)or a presumptive intra-chromosomal inversion(4 A).For eight out of the nine FISH sites detected on the Chinese Spring(CS)karyotype,there was a good match with the reference genome sequence,indicating that the most recent assembly has dealt well with the problem of placing tandem repeats.The integrated genetic map produced for wheat is informative as to the location of blocks of tandemly repeated DNA and can aid in improving the quality of the genome sequence assembly in regions surrounding these blocks.
基金supported by the National High Technology Research and Development Program of China(863 Program, 2006AA10Z179, 2006AAl0Z1F8)the Key Technologies R&D Program (2006BAD01A02,2006 BAD13B02)
文摘Novel LMW-GS （low molecular weight glutenin subunit）,α- and γ-gliadin from Triticum macha accessions were characterized via genomic PCR, which can do favor to improve the wheat quality. The complete coding regions of two α-gliadin, two γ-gliadin and two LMW-GS gene sequences, which designed as Gli-Mal, Gli-Ma2, Gli-Mrl, Gli-Mr2, Glu-LM1 and Glu-LM2, encoded the mature proteins with 307, 241, 348, 302, 474 and 377 amino acid residues, respectively. Gli-Mal and Gli-Ma2 were recognized as pseudogenes due to the in-frame stop codons. The amino acid sequences deduced from these gene sequences were characterized with the typical structure of α- or γ-gliadin or LMW- m type proteins with the exception of Gli-Ma2. Phylogenetic analysis showed Gli-Mal was closely related to those from T. aestivum, whereas Gli-Ma2 seemed to be more homologous with the gene sequences from Dasypyrum breviaristatum. Gli-Mrl was closely related to those from T. turgidum ssp. dicoccoides, while Gli-Mr2 was the nearest to those from T. aestivum. Glu-LM1 was closely related to those from Aegilops tauschii, whereas GIu-LM2 seemed to be more homologous with those from T. durum.
