Introgression of useful genes from Thinopyrum intermedium to wheat for improvement of bread‐making quality

To study the influence of genes from Thinopyrum intermedium on traits affecting the bread‐making quality of wheat, two derivatives from a putative disomic addition line in cultivar ‘Vilmorin 27’ were used in cytological, biochemical and molecular characterization. Cytological analysis suggested that one of the derivatives (Line‐1) had a terminal deletion involving the long arm of chromosome 1D (2n = 42, Del‐1DL”), and the other (Line‐2) was a conventional addition line, but also carried the same deletion on chromosome 1D (2n = 44, Thi”+Del‐1DL”). Amplification and sequencing of high‐molecular‐weight glutenin subunit (HMW‐GS) genes coded by the Th. intermedium chromosome in Line‐2 indicated the presence of one x‐type with an extra cysteine and four (rather than one) unique y‐type genes. Rheological studies of Line‐1 showed significantly lower dough strength compared to ‘Vilmorin 27’, confirming the recognized role of Glu‐1D coded HMW‐GSs. Line‐2 showed significantly higher dough strength compared to the background cultivar, indicating a significant potential of Th. intermedium for improvement of bread‐making quality in wheat.     

Characterization of high-molecular-weight glutenin subunit genes from Elytrigia elongata

Three high‐molecular‐weight glutenin subunit (HMW‐GS) genes from Elytrigia elongata (Host) Nevski were characterized by determining the coding sequences of two x‐type subunit genes Ee2.1 and Ee1.9, and one y‐type subunit gene Ee1.8 with 2082, 1938 and 1788 bp, respectively. The numbers of amino acids in the central repeat domains of Ee2.1, Ee1.9 and Ee1.8 were considerably fewer than the other known HMW‐GSs with substitutions, insertions and/or deletions involving a single or more amino acid residues. Moreover, an extra cysteine (Cys) was found in the repeated domain of x‐type subunit Ee2.1. The difference in the number and position of Cys residues might be associated with the good dough quality. The extra Cys in the good‐quality subunit Dx5 was at the beginning of the repetitive domain, while the extra Cys in Ee2.1 was at the central repetitive domain. Evolutionary relationships between the HMW‐GSs from E. elongata and the known HMW‐GSs were estimated. It transpired that Ee1.9, Ee2.1 and Bx7 were clustered into one subgroup with high bootstrap values, while Ee1.8 was closely related to Ay.     

Development and molecular cytogenetic identification of a new wheat–Leymus mollis Lm#6Ns disomic addition line

We developed a new disomic addition line M11028‐1‐1‐5 (2n = 44 = 21” + 1”) from a cross between wheat cv. ‘7182’ and octoploid Tritileymus M47 (2n = 8x = 56, AABBDDN sNs ). Cytological observations demonstrated that M11028‐1‐1‐5 contained 44 chromosomes and formed 22 bivalents during meiotic metaphase I. The genomic in situ hybridization (GISH) investigations showed this line contained 42 wheat chromosomes and a pair of L. mollis chromosomes. SSR, EST and PCR‐based landmark unique gene (PLUG) markers were screened to determine the homoeologous relationships of the introduced L. mollis chromosomes in wheat background. Nine markers, i.e. Xwmc256, Xgpw312, Swes123, CD452568, BF483643, BQ169205, TNAC1748, TNAC1751 and TNAC1752, all of which were located on the homoeologous group 6 chromosomes of common wheat, amplified bands unique to L. mollis in M11028‐1‐1‐5. Gliadin analysis also confirmed that the added chromosomes in M11028‐1‐1‐5 were correlated with the sixth group chromosome. This indicated that M11028‐1‐1‐5 contained a pair of introduced L. mollis chromosome belonging to homoeologous group 6, which we designated it as Lm#6 Ns disomic addition line. This is the first report of a common wheat–L. mollis disomic addition line.     

Methodology for creating alloplasmic soybean lines by using Glycine tomentella as a maternal parent

Soybean breeders have not exploited the diversity of the 26 wild perennial species of the subgenus Glycine Willd. that are distantly related to soybean [Glycine max (L.) Merr.] and harbour useful genes. The objective of this study was to develop a methodology for introgressing cytoplasmic and genetic diversity from Glycine tomentella PI 441001 (2n = 78) into cultivated soybean using ‘Dwight’ (2n = 40) as the male parent. Immature seeds (19–21 days post‐pollination) were cultured in vitro to produce F₁ plants (2n = 59). Amphidiploid (2n = 118) plants, induced by colchicine treatment, were vigorous and produced mature pods and seeds after backcrossing with ‘Dwight’. The BC₁ plants (2n = 79) produced mature seeds in crosses with ‘Dwight’. Chromosome numbers in BC₂F₁ plants ranged from 2n = 41–50. From BC₂F₂ to BC₃F₁, the number of plants in parentheses with 2n = 40 (275), 2n = 41 (208), 2n = 42 (80), 2n = 43 (27), 2n = 44 (12) and 2n = 45 (3) were identified. Fertile lines were grown in the field during 2012 and 2013. This is the first report of the successful development of new alloplasmic soybean lines with cytoplasm from G. tomentella.     

Diversity of five glutenin loci within durum wheat (Triticum turgidum L. ssp. durum (Desf.) Husn.) germplasm grown in Algeria

The HMW and B‐LMW glutenin subunits composition of 120 durum wheat germplasm grown in Algeria was examined using SDS‐PAGE. All together, 39 glutenin patterns were detected, including eight for HMW and 21 for B‐LMW glutenin subunits. Twenty‐six different alleles were identified for the five glutenin loci studied, that is, Glu‐A1 (3), Glu‐B1 (7), Glu‐A3 (5), Glu‐B3 (9) and Glu‐B2 (2). Two new alleles were found at Glu‐B3 locus: Glu‐B3new₁ encodes for five subunits (7 + 8 + 14 + 16 + 18) and Glu‐B3new₂ codes for five subunits (4 + 6* + 12 + 15 + 15*), of which subunit 15* with mobility between bands 15–16 was not described previously. At the Glu‐1 loci, the Glu‐A1c/Glu‐B1e allelic composition was predominant. For the B‐LMW glutenins, the most common allelic composition was Glu‐A3a/Glu‐B3a/Glu‐B2a. The collection analysed shows glutenin alleles and allele combinations related to high gluten strength. This information could be useful to select varieties with improved quality and also as a source of genes to develop new lines when breeding for quality.     

Survey and new insights in the application of PCR‐based molecular markers for identification of HMW‐GS at the Glu‐B1 locus in durum and bread wheat

Wheat, among all cereal grains, possesses unique characteristics conferred by gluten; in particular, high molecular weight glutenin subunits (HMW‐GS) are of considerable interest as they strictly relate to bread‐making quality and contribute to strengthening and stabilizing dough. Thus, the identification of allelic composition, in particular at the Glu‐B1 locus, is very important to wheat quality improvement. Several PCR‐based molecular markers to tag‐specific HMW glutenin genes encoding Bx and By subunits have been developed in recent years. This study provides a survey of the molecular markers developed for the HMW‐GS at the Glu‐B1 locus. In addition, a selection of molecular markers was tested on 31 durum and bread wheat cultivars containing the By8, By16, By9, Bx17, Bx6, Bx14 and Bx17 Glu‐B1 alleles, and a new assignation was defined for the ZSBy9_aF1/R3 molecular marker that was specific for the By20 allele. We believe the results constitute a practical guide for results that might be achieved by these molecular markers on populations and cultivars with high variability at the Glu‐B1 locus.     

Pollen viability and meiotic behaviour in intraspecific hybrids of Hydrangea aspera subsp. aspera Kawakami group x subsp. sargentiana

Pollen viability and male meiosis in intraspecific hybrids of Hydrangea aspera subsp. aspera Kawakami group (2n = 2x = 36) and subsp. sargentiana (2n = 2x = 34) were investigated. Although it had been assumed that they were sterile, pollen viability was observed; it varied from 2.5% to 12.1%. The production of gametes with different chromosome numbers was implied by the analysis of the dispersion of the diameter distribution of pollen grains. Analysis of male meiosis made it possible to identify the origins with two major categories of meiotic aberrations: abnormal chromosome distribution (early chromosome migration at metaphase, lagging chromosomes at anaphase, micronuclei at telophase), leading to the formation of unbalanced tetrads and/or ones with supernumerary microspores; and abnormal spindle orientation in metaphase II (tripolar, fused and parallel spindles), leading to the formation of dyads or triads. The mode of 2n pollen formation is of the First Division Restitution type. The high level of parental heterozygosity that is normally associated with them should facilitate the transfer of a polygenic trait in breeding programme.     

Eyespot resistance in wheat ×Aegilops kotschyi backcross lines

In wheat, eyespot caused by PseudoCercosporella herpotrichoides, is one of the main foot‐rot diseases. Yield losses up to 40% occur in some years. Plant protection by fungicide application is possible, but a better way is through resistance breeding. Two resistance sources are currently used: Aegilops ventricosa and the old French variety ‘Cappelle Desprez’. A new source of resistance has been found in the accession AE120 of Ae. kotschyi from the Gatersleben gene bank with the genome constitution UUSvSv. This accession has been crossed and backcrossed twice to susceptible wheat varieties, and in each generation, plants with a relatively high level of resistance have been selected. From this material, lines have been developed and tested in F₆ to F₈. Finally, several lines could be classified as moderately resistant, such as the French variety ‘Cappelle Desprez’ after resistance determination during milk ripeness (DC75). No line reached the high resistance level achieved with Pch‐1 from Ae. ventricosa. The yield of these lines under infection conditions was higher compared with ‘Cappelle Desprez’. The line 6018‐96‐3 showed a high yield of 64.3 dt/ha compared with 59.6 dt/ha, on the average, in combination with the best expression of eyespot resistance in the adult growth stage over 3 years.     

First-division restitution in hybrids of Langdon durum disomic substitution lines with rye and Aegilops squarrosa

Durum wheat ‘Langdon’(LDN) caused a high frequency of first‐division restitution (FDR) and partial fertility in hybrids with rye, Secale cereale L., and Aegilops squarrosa L. In order to determine the genetic control of FDR, a complete set of 14 Langdon durum D‐genome disomic substitution lines (LDNDS) was crossed with ‘Gazelle’ rye and one accession (RL5286) of A. squarrosa. The microsporogenesis and fertility of the hybrids were studied. The results showed that most of the hybrids expressed a high frequency of FDR and partial fertility. However, the hybrids of 2D(2A), 4D(4A), 5D(5B) and 6D(6B) crossed with both rye and A. squarrosa, as well as 1D(1A) with A. squarrosa, had either little or no FDR and were completely sterile. These hybrids had different types of first meiotic divisions compared with LDN control hybrids. The hybrids with 2D(2A), 4D(4A) and 6D(6B) had a high frequency of random segregation of chromosomes at the first division. The hybrids with 5D(5B), as expected, showed high homoeologous pairing. The hybrid of 1D(1A) with A. squarrosa had a high frequency of equational division at first division. These results suggest that the reduced or absent FDR in such hybrids might be related to the substitution of chromosomes with an FDR gene and poor compensation ability of the D‐genome chromosomes for their homoeologous A‐ or B‐ genome chromosomes. Cytological analysis suggested that chromosome 4A in LDN most likely carries a gene for high frequency of FDR in hybrids. In addition, some monads were observed at the end of meiosis in the hybrids of 3D(3A) and 6D(6A) crossed with rye. They were formed from FDR cells that failed to divide at second division, suggesting that the LDN 3A and 6A chromosomes might carry genes for normal second division of FDR cells in the rye crosses.     

Development of a complete set of disomic rape-radish chromosome-addition lines

After crosses of multiple rape-radish chromosome additions with rape, nine different monosomic additions (2n = 4x = 38 + 1) of individual radish chromosomes, a–i, were isolated, having the genomic background of winter oil seed rape and radish cytoplasm. Extra chromosomes were identified with RAPD markers, specific for individual radish chromosomes. All radish-plasmic monosomic additions, except that of chromosome f, had low seed production due to pistilloid stamens. Rape cytoplasm was substituted by crossing the monosomic additions as pollinators to rape followed by reselection of the radish chromosome additions. The monosomic additions in rape cytoplasm showed normal male fertility. Average transmission rates of radish chromosomes via egg cells and pollen cells were 37% and 27%, respectively. Lowest male transmission was found for chromosomes e and f with 0.04 and 0.01. Monosomic additions were self-pollinated to produce disomic additions. Monosomic and disomic addition progenies were discriminated by a prescreening with quantitative double primer (dp) RAPDs and cytological confirmation of preselected candidates by FISH analysis using a Raphanus-specific probe. Usability of this complete set of nine disomic rape-radish additions (2n = 4x = 38 + 2) was discussed.     

Development of chloroplast‐specific microsatellite markers for molecular characterization of alloplasmic lines and phylogenetic analysis in wheat

Wheat (Triticum aestivum L.) is strictly a self‐pollinated crop, where hybrid breeding requires well‐characterized cytoplasmic male sterile (CMS) lines. The CMS has mostly been developed by substituting nuclear genome of wheat into the cytoplasm from wild relatives. Molecular characterization of 90 genotypes including 82 CMS lines originating from five different species, namely Aegilops speltoides, Ae. kotschyi, Ae. variabilis, Triticum araraticum and T. timopheevii, and eight popular varieties was carried out. Consequently, a set of 25 microsatellite markers specific to chloroplast (cpSSRs) were designed and successfully validated for specificity of amplification. A total of 15 cpSSRs (60%) were found polymorphic, of which three cpSSRs (TaCM7, TaCM8 and TaCM11) in genic region and twelve cpSSRs were located in intergenic region. Phylogenetic analysis of genotypes using cpSSRs revealed two major groups well in accordance with respective origin. A set of cpSSRs and phylogeny of CMS belonging to different origins developed, which will be helpful for the improvement in CMS system in wheat. The genic cpSSRs can be used for the allele mining and evolutionary studies.     

Detection of HMW glutenin subunit variations among 205 cultivated emmer accessions (Triticum turgidum ssp. dicoccum)

The high molecular weight glutenin subunits (HMW‐GS) encoded by Glu‐1 loci among 205 accessions of cultivated emmer wheat (Triticum turgidum ssp. dicoccum Schrank) collected from different regions of Europe and China were separated and characterized by SDS‐PAGE in combination with two‐dimensional gel electrophoresis (A‐PAGE × SDS‐PAGE) and acidic capillary electrophoresis. High genetic polymorphisms in HMW‐GS compositions were found. A total of 40 alleles (6 for Glu‐A1 and 34 for Glu‐B1) and 62 subunit combinations (genotypes) were detected, some of which were not previously described. At Glu‐A1 locus, two novel alleles, designated Glu‐A1x coding for the subunit 1A × 1.1 and Glu‐A1y coding for the subunit 1A × 2.1′ were found while seven new subunits (1B × 17*, 1B × 6′, 1B × 13′, 1B × 20*, 1By9*, 1By14.1 and 1By8.1) and 20 novel alleles at Glu‐B1 locus were detected. In particular, some additional protein components were detected, which probably were 1Ay subunits encoded by Glu‐A1 locus. The introduction of both Ax and Ay subunits from tetraploid wheats into hexaploid wheats may increase the genetic variability of gluten genes and consequently improve flour technological properties.     

Genetic polymorphisms at Gli-Dt gliadin loci in Aegilops tauschii as revealed by acid polyacrylamide gel and capillary electrophoresis

Gliadin variation at Gli‐D^t1 and Gli‐D^t2 loci in 198 Aegilops tauschii accessions was studied by acid polyacrylamide gel electrophoresis (A‐PAGE) and capillary electrophoresis (CE). High genetic polymorphisms were found at both gliadin coding loci, revealing a total of 184 and 169 gliadin variants at the Gli‐D^t1 and Gli‐D^t2 loci, respectively. In particular, 12 gliadin blocks encoded by different alleles were apparently expressed and readily identified in six synthetic hexaploids produced by hybridization between Triticum durum and Ae. tauschii accessions. Compared with Ae. tauschii ssp. eusquarrosa, the gliadin profile of the D genome in Ae. tauschii ssp. strangulata more resembles that of T. aestivum, supporting the view that the subspecies strangulata is the most likely progenitor of bread wheat. Capillary electrophoresis analysis showed that the method is capable of separating and characterizing gliadins with speed, in high resolution using small sample amounts, and is well‐suited to detect protein alleles and to identify desirable genotypes in wheat quality improvement.     

Influence of high molecular weight glutenin subunit substitution on rheological behaviour and bread-baking quality of near-isogenic lines developed from Chinese wheats

Three near‐isogenic lines (NILs) of wheat involving Glu‐B1 and Glu‐D1 alleles were used to study the genetic contribution of high molecular weight glutenin subunits (HMW‐GS) to gluten strength. The HMW‐GS composition of each NILs was determined by SDS‐PAGE. No significant differences were found in grain protein contents among the NILs. Gluten strength and dough‐mixing properties were measured by the Farinograph, the Extensograph, and SDS‐sedimentation (SDS‐SE). Results indicated that line 2, containing the Glu‐1B 14 + 15 and Glu‐1D 5 + 10 combination of subunits, had higher values for flour quality, dough rheological parameters, and bread‐baking quality when compared with lines 8 and 13. Line 8, containing Glu‐1B 7 + 9 and Glu‐1D 5 + 10, was better than line 13 with the Glu‐1B 14 + 15 and Glu‐1D 10 combination. Some major parameters appeared significantly different. The presence of Glu‐1B 14 + 15 was associated with higher dough strength based on SDS‐SE volume and several rheological parameters when compared with Glu‐1B 7 + 9. Lines with subunit 10 at Glu‐D1 performed significantly worse than those with 5 + 10 in gluten index, SDS‐SE volume, Farinograph stability time, Extensograph area and bread‐baking quality.     

Unreduced 3x gamete formation of allotriploid hybrid derived from the cross of Primula denticulata (4x)?×?P. rosea (2x) as a causal factor for producing pentaploid hybrids in the backcross with pollen of tetraploid P. denticulata

A triploid hybrid, which was obtained from interspecific crosses between tetraploid Primula denticulata (2n = 4x = 44) and P. rosea (2n = 2x = 22), successfully produced 11 plants by backcrossing with pollen of tetraploid P. denticulata. Analysis of ploidy level using flow cytometry and chromosome counting in the 11 BC₁ plants revealed that all progeny had much larger DNA contents and chromosome number than both parents. In this triploid-tetraploid (3x–4x) crossing, progeny was predominantly true or near pentaploid presumably produced by the fertilization between true or near triploid female gamete produced from triploid hybrid and diploid pollen of tetraploid P. denticulata. These results suggest that unreduced (3x) or near triploid female gametes were partially produced by single step meiosis, either first-division restitution or second-division restitution process. The zygotes formed by the fertilization between true or near triploid egg produced by single step meiosis in triploid hybrid and diploid pollen produced by normal meiosis of tetraploid P. denticulata might be the only survivors in embryogenesis.     

Expression of morphological and flowering time variation through allopolyploidization: an empirical study with 27 wheat synthetics and their parental Aegilops tauschii accessions

It was recently shown that allopolyploidy brings novel epistatic interactions to genes belonging to different genomes. However, systematic studies of the phenotypic relationships between synthetic hexaploid wheats and their parental lines have not been conducted. In this study, 27 synthetic hexaploid wheats were produced by crossing the tetraploid wheat cultivar ‘Langdon’ with 27 accessions of Aegilops tauschii. Variations in 20 morphological and flowering traits were analysed in both the synthetic wheat lines and the parental Ae. tauschii accessions. The 20 traits exhibited large variations in the wheat lines. For many of the traits, the degree of variation in the parental accessions was reduced in the hexaploid derivatives. Principal component analysis of floret‐related traits divided the Ae. tauschii accessions into two subspecies, ssp. tauschii and ssp. strangulata, but this parental pattern of subspecific division was not detectable in the hexaploids. Our results suggest that the ‘Langdon’ genome may have an alleviating effect on the expression of D‐genome‐derived variations in derived synthetics.     

The study of introgressive lines of Triticum aestivum × Aegilops speltoides by in situ and SSR analyses

Fluorescence in situ hybridization (FISH) with a genome‐specific repeat, Spelt1, and wheat simple sequence repeat (SSR) markers were used to analyse the chromosome constitution of two Triticum aestivum×Aegilops speltoides introgressive lines. The lines 170/98i and 178/98i carried one and two subtelomeric regions of Ae. speltoides (per haploid genome), respectively, marked by Spelt1 repeats according to FISH data. SSR analysis detected homoeologous substitution of wheat chromosome 7D with Ae. speltoides chromosome 7S in the lines 178/98i and 170/98i as well as the assumed terminal translocation in the short arm of chromosome 3A in the line 178/98i. Anthocyanin pigmentation of the coleoptiles was found in the lines 170/98i and 178/98i and resulted from the 7S (7D) substitution. It was demonstrated that Spelt1 could be effectively used for the rapid identification (without DNA isolation) of terminal translocations of T. aestivum×Ae. speltoides introgressive lines as well as for further analysis of the stability of the hybrid plants.     

Genomic affinities between Brassica napus and Raphanus raphanistrum as revealed by meiotic GISH

An intergeneric hybrid between Brassica napus (AC, n = 19) and Raphanus raphanistrum (Rr, n = 9) was developed using the sequential ovary–ovule culture. Morphological, molecular, cytological studies and fluorescent genomic in situ hybridization were conducted to establish the hybridity and genomic relatedness. F₁ hybrid plants were male sterile and morphologically intermediate between the two parents. Cytological analysis of the hybrid plants revealed 10II + 8I as the predominant meiotic configuration with the mean bivalent frequency of 8.83. A maximum of 13II were observed in 8.24% of the pollen mother cells (PMCs). GISH studies facilitated the identification of two allopairs, confirming homoeologous pairing between A/C and Rr genomes. This may have significant implications in terms of transgene flow and possible introgression into R. raphanistrum, which is a common canola weed in Canada and Australia. The intergeneric hybrid B. napus × R. raphanistrum so produced also has the potential to be used as a bridging species for the transfer of desirable genes to amphiploid crop Brassica species carrying A/C genomes.     

Allelic variation at the <Emphasis Type="Italic">Glu-1</Emphasis> and <Emphasis Type="Italic">Glu-3</Emphasis> loci,presence of the 1B.1R translocation,and their effects on mixographic properties in Chinese bread wheats

Allelic variations at the Glu-1 and Glu-3 loci play an important role in determining dough properties and bread-making quality. Two hundred and fifty-one cultivars and advanced lines from four major Chinese wheat-producing zones in the autumn-sown wheat regions were used to investigate the high-molecular-weight glutenin subunits (HMW GS) and low-molecular-weight glutenin subunit (LMW GS) composition controlled by the Glu-1 and Glu-3 loci, respectively, as well as the presence of the 1B.1R translocation, and to determine the association of storage protein composition with protein content, SDS sedimentation value, and dough-mixing properties measured by mixograph. Three, nine, and four allelic variations were present at Glu-A1, Glu-B1, and Glu-D1, respectively. Subunits 1, N, 7+8, 7+9, and 2+12 are the dominant HMW GS, with frequencies of 51.3, 39.4, 38.2, 45.0, and 59.8%, respectively. Five and eight allelic variations were present at the Glu-A3 and Glu-B3 loci (data of Glu-D3 were not available), Glu-A3a, Glu-A3d, Glu-B3j (presence of the 1B.1R translocation), and Glu-B3d are the dominant LMW GS, with frequencies of 37.1, 31.7, 44.6, and 20.3%, respectively. The frequencies of allelic variation at Glu-1 and Glu-3 differ greatly in different regions. The effects of HMW GS and LMW GS on SDS sedimentation value, mixing time, and mixing tolerance were significant at P = 0.01, with Glu-D1 and Glu-B3 showing the largest contributions to mixing time and mixing tolerance. Averaged data from two locations showed that the quality effects of glutenin loci could be ranked as Glu-B3 > Glu-B1 > Glu-A1 > Glu-D1 > Glu-A3 for SDS sedimentation value, Glu-D1 > Glu-B3 > Glu-A1 = Glu-B1 = Glu-A3 for mixing time, and Glu-D1 > Glu-B3 = Glu-B1 > Glu-A3 > Glu-A1 for mixing tolerance, respectively. The significant and negative effect of the 1B.1R translocation on dough properties was confirmed. It was concluded that the high frequency of undesirable HMW GS and LMW GS and the presence of the 1B.1R translocation are responsible for the weak gluten property of Chinese germplasm; hence, reducing the frequency of the 1B.1R translocation and integration of desirable subunits at Glu-1 and Glu-3 such as 1, 7+8, 14+15, 5+10, Glu-A3d, and Glu-B3d, could lead to the improvement of gluten quality in Chinese wheats.     

欧山羊草高分子谷蛋白亚基向小麦的转入及其遗传规律

欧山羊草具有两个容易用聚丙烯酰胺凝胶电泳与小麦区别的高分子谷蛋白亚基,分别命名为Glu-Aeb1和Glu-Aeb2。采用聚丙烯酰胺凝胶电泳（SDS-PAGE）方法对21份欧山羊草与小麦杂交后代BC2F6和15份BC2F5与小麦杂交F0种子的谷蛋白亚基组成进行了鉴定。实验结果表明,36份材料全部都有欧山羊草高分子谷蛋白亚基的转入。Glu-Aeb1在BC2F6和杂交组合F0中的传递频率分别为59.52%和20.94%。而Glu-Aeb2在BC2F6和杂交组合F0中的传递频率分别为40.48%和22.96%。