RESUMEN
Fifteen novel α-gliadin genes were cloned and sequenced from Triticum and related Aegilops genomes by allele-specific polymerase chain reaction (AS-PCR). Sequence comparison displayed high diversities in the α-gliadin gene family. Four toxic epitopes and glutamine residues in the two polyglutamine domains facilitated these α-gliadins to be assigned to specific chromosomes. Five representative α-gliadin genes were successfully expressed in Escherichia coli, and their amount reached a maximum after 4 h induced by isopropyl-ß-D-thiogalactoside (IPTG), indicating a high level of expression under the control of T7 promoter. The transcriptional expression of α-gliadin genes during grain development detected by quantitative real-time polymerase chain reaction (qRT-PCR) showed a similar up-down regulation pattern in different genotypes. A neighbor-joining tree constructed with both full-open reading frame (ORF) α-gliadin genes and pseudogenes further revealed the origin and phylogenetic relationships among Triticum and related Aegilops genomes. The evolutionary analysis demonstrated that α-gliadin genes evolved mainly by synonymous substitutions under strong purifying selection during the evolutionary process.
Asunto(s)
Clonación Molecular , Evolución Molecular , Genoma de Planta , Gliadina/genética , Gliadina/metabolismo , Poaceae/genética , Triticum/genética , Secuencia de Aminoácidos , Genes de Plantas , Gliadina/química , Datos de Secuencia Molecular , Filogenia , Reacción en Cadena en Tiempo Real de la Polimerasa , Alineación de SecuenciaRESUMEN
BACKGROUND: Brachypodium distachyon L. is a newly emerging model plant system for temperate cereal crop species. However, its grain protein compositions are still not clear. In the current study, we carried out a detailed proteomics and molecular genetics study on grain glutenin proteins in B. distachyon. RESULTS: SDS-PAGE and RP-HPLC analysis of grain proteins showed that Brachypodium has few gliadins and high molecular weight glutenin subunits. In contrast the electrophoretic patterns for the albumin, globulin and low molecular weight glutenin subunit (LMW-GS) fractions of the grain protein were similar to those in wheat. In particular, the LMW-C type subunits in Brachypodium were more abundant than the equivalent proteins in common wheat. Southern blotting analysis confirmed that Brachypodium has 4-5 copies of LMW-GS genes. A total of 18 LMW-GS genes were cloned from Brachypodium by allele specific PCR. LMW-GS and 4 deduced amino acid sequences were further confirmed by using Western-blotting and MALDI-TOF-MS. Phylogenetic analysis indicated that Brachypodium was closer to Ae. markgrafii and Ae. umbellulata than to T. aestivum. CONCLUSIONS: Brachypodium possessed a highly conserved Glu-3 locus that is closely related to Triticum and related species. The presence of LMW-GS in B. distachyon grains indicates that B. distachyon may be used as a model system for studying wheat quality attributes.
Asunto(s)
Brachypodium/genética , Glútenes/genética , Filogenia , Proteínas de Plantas/genética , Secuencia de Aminoácidos , Cromatografía Líquida de Alta Presión , Clonación Molecular , Secuencia Conservada , ADN de Plantas/genética , Electroforesis en Gel de Poliacrilamida , Genes de Plantas , Datos de Secuencia Molecular , Peso Molecular , Reacción en Cadena de la Polimerasa , Proteómica , Alineación de Secuencia , Espectrometría de Masa por Láser de Matriz Asistida de Ionización Desorción , Triticum/genéticaRESUMEN
Gliadins were the major components of wheat storage proteins and determine the extensibility properties of gluten dough. In this work, 19 new full-length γ-gliadin genes were isolated from various Aegilops and Triticum species. Sequence characterization showed that a specific octapeptide and celiac disease (CD)-toxic epitope Gliγ-3 (VQGQGIIQPQQPAQL) were present in the rich glutamine domain and C-terminal non-repetitive domain, respectively. Based on the sequence features of both peptides, a new classification system for γ-gliadin gene family was established, in which γ-gliadins were classified into two types (types I and II) with each consisting of two groups. An uneven distribution of different types and groups of γ-gliadin genes was exhibited among 11 Aegilops and Triticum genomes. Phylogenetic analysis revealed that types I and II genes diverged at about 14 MYA while the divergence of 4 γ-gliadin group genes occurred at around 10 MYA almost simultaneously. The γ-gliadin genes from S(l) and B genomes displayed a different transcriptional expression pattern during grain development, and rapid increasing of gliadin mRNA and proteins occurred at 15-20 DPA. In addition, genome-specific variations of CD-toxic epitopes among Aegilops and Triticum genomes were found. The A genome and its related progenitor genomes A(u) and A(m) had fewer CD epitopes than other genomes, suggesting that these genomes might be valuable gene resources to remove CD toxic peptides for wheat quality improvement.
Asunto(s)
Regulación de la Expresión Génica de las Plantas , Genes de Plantas/genética , Gliadina/genética , Poaceae/genética , Triticum/genética , Secuencia de Aminoácidos , Enfermedad Celíaca/inmunología , Epítopos/inmunología , Evolución Molecular , Perfilación de la Expresión Génica , Gliadina/química , Gliadina/inmunología , Gliadina/metabolismo , Datos de Secuencia Molecular , Péptidos/química , Filogenia , ARN Mensajero/genética , ARN Mensajero/metabolismo , Estándares de Referencia , Semillas/genética , Semillas/crecimiento & desarrollo , Alineación de Secuencia , Transcripción GenéticaRESUMEN
In this study, a novel y-type high molecular weight glutenin subunit (HMW-GS) in wild emmer wheat Triticum turgidum L. var. dicoccoides (Körn.) accession KU1952 was identified by sodium dodecyl sulphate polyacrylamide gel electrophoresis (SDS-PAGE), capillary electrophoresis (CE) and matrix-assisted laser desorption ionisation/time-of-flight/mass spectrometry (MALDI-TOF-MS). Its electrophoretic mobility and molecular weight were similar to those of 1By16 and was designated as 1By16*. The complete coding sequence of the 1By16* gene isolated by allelic-specific polymerase chain reaction (AS-PCR) consists of 2,157 bp, encoding 729 amino acid residues. The real presence and authenticity of the 1By16* gene in KU1952 were further confirmed by liquid chromatography-tandem mass spectrometry (LC-MS/MS), heterologous expression and Western blotting. The molecular structure as well as phylogenetic analysis revealed that 1By16* had 21 single-nucleotide polymorphism (SNP) variations and possessed greater similarity with superior quality subunits 1By15 and 1By16 of common wheat. Secondary structure prediction displayed higher α-helix and ß-strand contents in the 1By16* subunit, which could form a superior gluten structure and, consequently, might have positive effects on dough quality. Our results suggest that 1By16* is expected to be a new potential gene for wheat quality improvement.
Asunto(s)
Glútenes/genética , Subunidades de Proteína/genética , Triticum/genética , Alelos , Secuencia de Aminoácidos , Western Blotting , Cromatografía Liquida , Clonación Molecular , Electroforesis en Gel de Poliacrilamida , Genes de Plantas/genética , Glútenes/química , Datos de Secuencia Molecular , Peso Molecular , Sistemas de Lectura Abierta/genética , Péptidos/química , Filogenia , Proteínas de Plantas/química , Proteínas de Plantas/genética , Reacción en Cadena de la Polimerasa , Polimorfismo de Nucleótido Simple/genética , Células Procariotas/metabolismo , Estructura Secundaria de Proteína , Subunidades de Proteína/química , Espectrometría de Masa por Láser de Matriz Asistida de Ionización DesorciónRESUMEN
The winter wheat cultivar Red Chief has been identified as the wheat cultivar most resistant to Pyrenophora tritici-repentis (Ptr). This study was undertaken to determine the inheritance, chromosomal location and molecular mapping of a tan spot resistance gene in Red Chief. χ² analysis of the F2 segregation data of the hybrids between 21 monosomic lines of the susceptible wheat cultivar Chinese Spring and the resistant cultivar Red Chief revealed that tan spot resistance in cv. Red Chief is controlled by a single recessive gene located on chromosome 3A. Linkage analysis using SSR markers in the Red Chief/Chinese Spring F2 population showed that the tsr4 gene is clustered in the region around Xgwm2a, on the short arm of chromosome 3A. This marker has also been identified as the closest marker to the tsr3 locus on chromosome 3D in synthetic wheat lines. Validation analysis of this marker for the tsr3 and tsr4 genes using 28 resistant and 6 susceptible genotypes indicated that the 120 bp allele (the tsr3 gene) specific fragment was observed in 11 resistant genotypes, including the three synthetic lines XX41, XX45 and XX110, while the 130 bp allele was amplified only in cv. Red Chief and Dashen. Xgwm2a can be used to trace the presence of the target gene in successive backcross generations and pyramiding of the tsr3 & tsr4 genes into a commonly grown and adaptable cultivar.
Asunto(s)
Mapeo Cromosómico/métodos , Cromosomas de las Plantas/genética , Genes de Plantas/genética , Inmunidad Innata/genética , Enfermedades de las Plantas/genética , Estaciones del Año , Triticum/genética , Ascomicetos/aislamiento & purificación , Ascomicetos/fisiología , Segregación Cromosómica/genética , Cruzamientos Genéticos , Ligamiento Genético , Marcadores Genéticos , Genotipo , Patrón de Herencia/genética , Repeticiones de Microsatélite/genética , Enfermedades de las Plantas/inmunología , Enfermedades de las Plantas/microbiología , Reproducibilidad de los Resultados , Triticum/clasificación , Triticum/inmunología , Triticum/microbiologíaRESUMEN
Twelve new LMW-GS genes were characterized from bread wheat (Triticum aestivum L.) cultivar Zhongyou 9507 and five Aegilops species by AS-PCR. These genes belong to the LMW-m type and can be classified into two subclasses designated as 1 and 2, with the latter predominant in both wheat and related wild species. Genes in the two subclasses were significantly different from each other in SNPs and InDels variations. In comparison to subclass 1, the structural features of subclass 2 differs in possessing 21 amino acid residue substitutions, two fragment deletions (each with 7 amino acid residues), and a double-residue deletion and two fragment insertions (12 and 2-5 residues). Phylogenetic analysis revealed that the two subclasses were divergent at about 6.8 MYA, earlier than the divergence of C, M, N, S(s) and U genomes. The S(s) and B genomes displayed a very close relationship, whereas the C, M, N and U genomes appeared to be related to the D genome of bread wheat. The presently characterized genes ZyLMW-m1 and ZyLMW-m2 from Zhongyou 9507 were assigned to the D genome. Moreover, these genes were expressed successfully in Escherichia coli. Their transcriptional levels during grain developmental stages detected by quantitative real-time PCR (qRT-PCR) showed that both genes started to express at 5 days post-anthesis (DPA), reaching the maximum at 14 DPA after which their expressions decreased. Furthermore, the expression level of ZyLMW-m2 genes was much higher than that of ZyLMW-m1 during all grain developmental stages, suggesting that the expression efficiency of LMW-GS genes between the two subclasses was highly discrepant.
Asunto(s)
Regulación de la Expresión Génica de las Plantas , Genes de Plantas/genética , Metionina/genética , Poaceae/genética , Transcripción Genética , Triticum/genética , Secuencia de Aminoácidos , Clonación Molecular , Endospermo/genética , Endospermo/crecimiento & desarrollo , Escherichia coli/metabolismo , Evolución Molecular , Perfilación de la Expresión Génica , Glútenes/química , Glútenes/genética , Glútenes/metabolismo , Datos de Secuencia Molecular , Peso Molecular , Filogenia , Subunidades de Proteína/química , Subunidades de Proteína/genética , Subunidades de Proteína/metabolismo , ARN Mensajero/genética , ARN Mensajero/metabolismo , Reacción en Cadena de la Polimerasa de Transcriptasa Inversa , Homología de Secuencia de Aminoácido , Factores de TiempoRESUMEN
This paper reports cloning and characterisation of four novel low-molecular-weight glutenin subunit (LMW-GS) genes (designated as TzLMW-m2, TzLMW-m1, TdLMW-m1 and AlLMW-m2) from the genomic DNA of Triticum dicoccoides, T. zhukovskyi and Aegilops longissima. The coding regions of TzLMW-m2, TzLMW-m1, TdLMW-m1 and AlLMW-m2 were 1056 bp, 903 bp, 1056 bp and 1050 bp in length, encoding 350, 300, 350 and 348 amino acid residues, respectively. The deduced amino acid sequences showed that the four novel genes were classified as LMW-m types and the comparison results indicated that the four genes had a more similar structure and a higher level of homology with the LMW-m genes than the LMW-s and -i types genes. However, the first cysteine residue's positions of TzLMW-m2, TdLMW-m1 and AlLMW-m2 were different from the others. Moreover, AlLMW-m2, TdLMW-m1 and TzLMW-m2 all possessed a longer repetitive domain, which was considered to be associated with good quality of wheat. The secondary structure prediction revealed that the content of beta-strand in AlLMW-m2 and TdLMW-m1 exceeded the positive control, suggesting that AlLMW-m2 and TdLMW-m1 should be considered as candidate genes that may have positive effect on dough quality. In order to investigate the evolutionary relationship of the novel genes with the other LMW-GSs, a phylogenetic tree was constructed. The results lead to a speculation that AlLMW-m2, TdLMW-m1 and TzLMW-m2 may be the middle types during the evolution of LMW-m and LMW-s.
Asunto(s)
Clonación Molecular , Genes de Plantas , Glútenes/genética , Subunidades de Proteína/genética , Triticum/genética , Secuencia de Aminoácidos , Glútenes/química , Datos de Secuencia Molecular , Filogenia , Poaceae/química , Poaceae/genética , Reacción en Cadena de la Polimerasa , Subunidades de Proteína/químicaRESUMEN
Synthetic wheat lines (2n = 6x = 42, AABBDD), which are amphiploids developed from the hybrid between tetraploid wheat (Triticum turgidum L., 2n = 4x = 28, AABB) and Aegilops tauschii Coss. (2n = 2x = 14, DD), are important sources of resistance against tan spot of wheat caused by Pyrenophora tritici-repentis. In the present study, inheritance, allelism and genetic linkage analysis in synthetic wheat lines have been carried out. Segregation analysis of the phenotypic and molecular data in F(2:3) populations of CS/XX41, CS/XX45, and CS/XX110 has revealed a 1:2:1 segregation ratio indicating that resistance of tan spot in these synthetic lines is controlled by a single gene. Allelism tests detected no segregation for susceptibility among F(1) and F(2) plants derived from intercrosses of the resistance lines XX41, XX45 and XX110 indicating that the genes are either allelic or tightly linked. Linkage analysis using SSR markers showed that all the three genes: tsn3a in XX41, Tsn3b in XX45 and tsn3c in XX110 are clustered in the region around Xgwm2a, located on the short arm of chromosome 3D. The linked markers and genetic relationship of these genes will greatly facilitate their use in wheat breeding and deployment of cultivars resistant to tan spot.
Asunto(s)
Ascomicetos/patogenicidad , Triticum/genética , Triticum/microbiología , Alelos , Mapeo Cromosómico , ADN de Plantas/genética , Genes de Plantas , Hibridación Genética , Repeticiones de Microsatélite , Enfermedades de las Plantas/genética , Enfermedades de las Plantas/microbiología , PoliploidíaRESUMEN
Three novel low molecular weight (LMW) glutenin subunits from cultivated einkorn (Triticum monococcum L., A(m)A(m), 2n = 2x = 14) were characterized by SDS-PAGE and molecular weights determined by MALDI-TOF-MS. Their coding genes were amplified and cloned with designed AS-PCR primers, revealing three complete gene sequences. All comprised upstream, open reading frame (ORF), downstream and no introns were present. The deduced amino acid sequences showed that all three genes, named as LMW-M1, LMW-M3 and LMW-M5, respectively, belonged to the LMW-i type subunits with the predicted molecular weight between 38.5206 and 38.7028 kDa. They showed high similarity with other LMW-i type genes from hexaploid bread wheats, but also displayed unique features. Particularly, LMW-M5 subunit contained an extra cysteine residue in the C-terminus except for eight conserved cysteines, which resulted from a single-nucleotide polymorphism (SNP) of the T-C transition, namely arginine --> cysteine substitution at position 242 from the N-terminal end. This is the first report that the LMW-i subunit contained nine cysteines residues that could result in a more highly cross-linked and more elastic glutenin suggesting that LMW-M5 gene may associates with good quality properties. In addition, a total of 25 SNPs and one insertions/deletions (InDels) were detected among three LMW-i genes, which could result in significant functional changes in polymer formation of gluten. It is anticipated that these SNPs could be used as reliable genetic markers during wheat quality improvement. The phylogenetic analysis indicated that LMW-i type genes apparently differed from LMW-m and LMW-s type genes and diverged early from the primitive LMW-GS gene family, at about 12.92 million years ago (MYA) while the differentiation of A(m) and A genomes was estimated at 3.98 MYA.
Asunto(s)
Glútenes/genética , Proteínas de Plantas/genética , Triticum/genética , Secuencia de Aminoácidos , Composición de Base , Clonación Molecular , Eliminación de Gen , Marcadores Genéticos , Glútenes/química , Glútenes/metabolismo , Datos de Secuencia Molecular , Mutagénesis Insercional , Filogenia , Proteínas de Plantas/química , Proteínas de Plantas/clasificación , Proteínas de Plantas/metabolismo , Reacción en Cadena de la Polimerasa , Polimorfismo de Nucleótido Simple , Prolaminas , Subunidades de Proteína/química , Subunidades de Proteína/genética , Subunidades de Proteína/metabolismo , Alineación de Secuencia , Triticum/metabolismoRESUMEN
Segregation analysis of resistance to powdery mildew in a F(2) progeny from the cross Chinese Spring (CS) x TA2682c revealed the inheritance of a dominant and a recessive powdery mildew resistance gene. Selfing of susceptible F(2) individuals allowed the establishment of a mapping population segregating exclusively for the recessive resistance gene. The extracted resistant derivative showing full resistance to each of 11 wheat powdery mildew isolates was designated RD30. Amplified fragment length polymorphism (AFLP) analysis of bulked segregants from F(3)s showing the homozygous susceptible and resistant phenotypes revealed an AFLP marker that was associated with the recessive resistance gene in repulsion phase. Following the assignment of this AFLP marker to wheat chromosome 7A by means of CS nullitetrasomics, an inspection of simple sequence repeat (SSR) loci evenly spaced along chromosome 7A showed that the recessive resistance gene maps to the distal region of chromosome 7AL. On the basis of its close linkage to the Pm1 locus, as inferred from connecting partial genetic maps of 7AL of populations CS x TA2682c and CS x Virest ( Pm1e), and its unique disease response pattern, the recessive resistance gene in RD30 was considered to be novel and tentatively designated mlRD30.
Asunto(s)
Ascomicetos , Mapeo Cromosómico , Inmunidad Innata/genética , Enfermedades de las Plantas/microbiología , Triticum/genética , Cruzamientos Genéticos , Genes Recesivos , Marcadores Genéticos/genética , Repeticiones de Minisatélite , Enfermedades de las Plantas/genética , Polimorfismo de Longitud del Fragmento de RestricciónRESUMEN
A novel y-type high-molecular-weight glutenin subunit possessing a slightly faster mobility than that of subunit 1Dy12 in SDS-PAGE, designated 1Dy12.1(t) in Aegilops tauschi, was identified by one- and two-dimensional gel and capillary electrophoresis. Its coding gene at the Glu-D(t) 1 locus was amplified with allele-specific-PCR primers, and the amplified products were cloned and sequenced. The complete nucleotide sequence of 2,807 bp containing an open reading frame of 1,950 bp and 857 bp of upstream sequence was obtained. A perfectly conserved enhancer sequence and the -300 element were present at positions of 209-246 bp and 424-447 bp upstream of the ATG start codon, respectively. The deduced mature protein of 1 Dy12.1(t) subunit comprised 648 amino acid residues and had a Mr of 67,518 Da, which is slightly smaller than the 1Dy12 (68,695 Da) but larger than the 1Dy10 (67,495 Da) subunits of bread wheat, respectively, and corresponds well with their relative mobilities when separated by acid-PAGE. The deduced amino acid sequence indicated that the 1Dy12.1(t) subunit displayed a greater similarity to the 1Dy10 subunit, with only seven amino acid substitutions, suggesting that this novel gene could have positive effect on bread-making quality. A phenetic tree produced by nucleotide sequences showed that the x- and y-type subunit genes were respectively clustered together and that the Glu-D(t) 1y12.1 gene of Ae. tauschii is closely related to other y-type subunit genes from the B and D genomes of hexaploid bread wheat.
Asunto(s)
Glútenes/análogos & derivados , Glútenes/genética , Poaceae/genética , Subunidades de Proteína/genética , Agricultura/métodos , Secuencia de Aminoácidos , Secuencia de Bases , Análisis por Conglomerados , Cartilla de ADN , Electroforesis en Gel de Agar , Electroforesis en Gel Bidimensional , Electroforesis en Gel de Poliacrilamida , Datos de Secuencia Molecular , Alineación de Secuencia , Análisis de Secuencia de ADNRESUMEN
The allelic compositions of high- and low-molecular-weight subunits of glutenins (HMW-GS and LMW-GS) among European spelt ( Triticum spelta L.) and related hexaploid and tetraploid Triticum species were investigated by one- and two-dimensional polyacrylamide-gel electrophoresis (PAGE) and capillary electrophoresis (CE). A total of seven novel glutenin alleles (designated A1a*, B1d*, B1g*, B1f*, B1j*, D1a* at Glu-1 and A3h at the Glu-3 loci, respectively) in European spelt wheat were detected by SDS-PAGE, which were confirmed further by employing A-PAGE and CE methods. Particularly, two HMW-GS alleles, Glu-B1d* coding the subunits 6.1 and 22.1, and Glu-B1f* coding the subunits 13 and 22*, were found to occur in European spelt with frequencies of 32.34% and 5.11%, respectively. These two alleles were present in cultivated emmer (Triticum dicoccum), but they were not observed in bread wheat (Triticum aestivum L.). The allele Glu-B1g* coding for 13* and 19* subunits found in spelt wheat was also detected in club wheat (Triticum compactum L.). Additionally, two alleles coding for LMW-GS, Glu-A3h and Glu-B3d, occurred with high frequencies in spelt, club and cultivated emmer wheat, whereas these were not found or present with very low frequencies in bread wheat. Our results strongly support the secondary origin hypothesis, namely European spelt wheat originated from hybridization between cultivated emmer and club wheat. This is also confirmed experimentally by the artificial synthesis of spelt through crossing between old European emmer wheat, T. dicoccum and club wheat, T. compactum.
Asunto(s)
Glútenes/análogos & derivados , Glútenes/genética , Ploidias , Triticum/genética , Alelos , Cruzamientos Genéticos , Electroforesis en Gel de Poliacrilamida , Europa (Continente) , Marcadores Genéticos , Peso Molecular , Proteínas de Plantas/genética , Subunidades de ProteínaRESUMEN
Bánkúti 1201, an old Hungarian wheat variety with special quality traits, was analysed to determine the relationships between its storage protein composition and superior quality-attributes for breadmaking. Based on the storage protein composition, the variety appears to have the nature of a population, containing several genotypes with different gluten protein alleles. Using molecular markers, a new mutant x-type HMW glutenin allele was identified, containing an extra cysteine residue and showing a moderate, positive-effect on gluten properties. In lines possessing subunits Bx7+By8 the overexpression of the Bx-type subunit could be detected, resulting in a higher unextractable polymeric protein (UPP) content and increased dough strength. It was found that the presence or absence of subunit Bx7 has an equilibrating effect on the dough extensibility, which is generally characteristic of the Bánkúti 1201 population. The complex good bread-making quality of the variety, which has strong but highly extensible dough, is probably due to the balance between lines which express subunit Bx7 and those which do not.
Asunto(s)
Glútenes/análogos & derivados , Proteínas de Plantas/análisis , Triticum/química , Alelos , Secuencia de Bases , Pan , Cruzamiento , ADN de Plantas/genética , Tecnología de Alimentos , Genes de Plantas , Variación Genética , Gliadina/análisis , Gliadina/química , Gliadina/genética , Glútenes/análisis , Glútenes/química , Glútenes/genética , Hungría , Peso Molecular , Mutación , Proteínas de Plantas/química , Proteínas de Plantas/genética , Subunidades de Proteína , Triticum/genéticaRESUMEN
The powdery mildew resistance gene Pm22, identified in the Italian wheat cultivar Virest and originally assigned to wheat chromosome 1D, was mapped to chromosome 7A with the aid of molecular markers. Mapping of common AFLP and SSR markers in two wheat crosses segregating for Pm22 and Pm1c, respectively, indicated that Pm22 is a member of the complex Pm1 locus. Pm22 also showed a pattern of resistance reaction to a differential set of Blumeria graminis f. sp. tritici isolates that was distinguishable from those from other Pm1 alleles in lines Axminster/8*Cc ( Pm1a), MocZlatka ( Pm1b), Weihenstephan Stamm M1N ( Pm1c) and Triticum spelta var. duhamelianum TRI 2258 ( Pm1d). Based on these results, the gene symbol Pm1e is proposed for the powdery mildew resistance gene in cv. Virest.