Identification of powdery-mildew-resistance genes in common wheat (Triticum aestivum L.). VI. Wheat cultivars grown in China

A total of 26 common wheat cultivars and advanced breeding lines grown in China were tested with a set of 11 differential powdery-mildew isolates. Seven cultivars were susceptible. Another seven cultivars showed the response pattern of resistance gene Pm2, either individually or in combination with genes Pm3d or Pm4a. Five cultivars expressed the resistance of gene Pm4b singly or in combination with Pm6. Another four cultivars exhibited the response patterns of genes Pm5, Pm6 and Pm8, respectively. Three cultivars, which included one breeding line with a pair of substituted chromosomes from Haynaldia villosa, presumably carrying the resistance gene Pm21, showed resistance-response patterns to all the isolates tested.     

Identification of powdery-mildew-resistance genes in common wheat (Triticum aestivum L. em. Thell.). V. Old German cultivars and cultivars released in the former GDR

A total of 59 old wheat cultivars grown in Germany prior to 1960 were tested for mildew response using a collection of 12 differential isolates of Erysiphe graminis DC f. sp. tritici Marchal (Blumeria graminis (DC) Speer f. sp. tritici). Nineteen cultivars did not possess any major resistance gene and 25 were characterized by susceptible or intermediate responses. Fifteen cultivars revealed isolate-specific response patterns that could not be attributed to known major resistance genes or gene combinations. Many of the old German cultivars inherited a mildew-resistance gene from the Canadian cultivar ‘Garnet’ which is tentatively designated M1-Ga. Cultivars ‘Bretonischer Bartweizen’ (designated M1-Br) and ‘Adlungs Alemannen’ (designated M1-Ad) appeared to carry unknown resistance genes. Among 18 winter wheat cultivars released in the former GDR. eight showed susceptibility to all isolates used. Cv. “Borenos” carries resistance gene Pm3c. Five cultivars possess gene Pm4b. two cultivars gene pm5 and one cultivar a combination of genes Pm2 and Pm4b. Cultivar ‘Zentos’ was resistant to almost all isolates used. Its resistance might be conditioned by different unknown major resistance genes.     

Chromosomal location of genes for resistance to powdery mildew in common wheat (Triticum aestivum L. em Thell.) 7. Gene Pm29 in line Pova

Chromosomal localization and linkage mapping of a powdery mildewresistance gene were conducted in the resistant wheat line Pova, derivedfrom a Triticum aestivum cv. Poros-Aegilops ovata-alien additionline. Monosomic analysis revealed that a major dominant gene was locatedon chromosome 7D. This gene possessed a distinct disease response patternagainst a differential set of Blumeria graminis tritici isolates andsegregated independently from resistance gene Pm19 also located onwheat chromosome 7D. Molecular genetic analysis showed that theresistance gene in Pova was specifically located on the long arm ofchromosome 7D closely linked to one RFLP and three AFLP markers. It isproposed that the new gene be designated Pm29.     

Molecular marker-facilitated pyramiding of different genes for powdery mildew resistance in wheat

Breeding durable resistance to pathogens and pests is a major task for modern plant breeders and pyramiding different resistance genes into a genotype is one way of achieving this. Three powdery mildew resistance gene combinations, Pm2+Pm4a, Pm2+Pm21, Pm4a+Pm21 were successfully integrated into an elite wheat cultivar ‘Yang047′. Double homozygotes were selected from a small F₂ population with the help of molecular markers. As the parents were near‐isogenic lines (NILs) of ‘Yang158′, the progenies showed good uniformity in morphological and other non‐resistance agronomic traits. The present work illustrates the bright prospects for the utilization of molecular markers in breeding for host resistance.     

Chromosomal location of genes for resistance to powdery mildew in common wheat (Triticum aestivum L. em Thell.). 8. Gene Pm32 in a wheat-Aegilops speltoides translocation line

The wheat-Aegilops speltoides translocation line L501 exhibits a disease response pattern distinctive from that of documented powdery mildew genes after inoculation with differential Blumeria graminis tritici isolates. Results based on cytological C-bandings and monosomic analyses reveal that a dominant resistance gene derived from Ae. speltoides is located on a T1BL·1SS chromosome translocation in this line. The new gene is designated Pm32. This revised version was published online in July 2006 with corrections to the Cover Date.     

Chromosomal location of powdery mildew resistance genes in common wheat (Triticum aestivum L. em. Thell.) 3. Gene Pm22 in cultivar Virest

Summary Common wheat cultivar Virest possesses mildew resistance which is different from resistances expressed by currently documented mildew resistance genes, detected by response to eleven differential wheat powdery mildew isolates. F₂ populations from hybrids of the 21 Chinese Spring monosomic lines with Virest revealed one major dominant gene, located on wheat chromosome 1D. The new gene is designated Pm22. Italian cultivars Elia, Est Mottin, Ovest and Tudest also showed the disease response pattern corresponding to Virest.     

Identification of resistance gene analogue markers closely linked to wheat powdery mildew resistance gene Pm31

Specific oligonucleotide primers, designed for the sequences of known plant disease resistance genes, were used to amplify resistance gene analogues (RGAs) from wheat genomic DNA. This method was applied in a bulked segregant analysis to screen for the RGA markers linked to the powdery mildew resistance gene Pm31, introgressed into common wheat from wild emmer. Two RGA markers (RGA200 and RGA390) were found to be closely linked to Pm31 and completely co‐segregating with the marker allele of Xpsp3029 linked to Pm31, with a genetic distance of 0.6 cM. These two RGA markers were then integrated into the formerly established microsatellite map of Pm31 region. The result showed the effectiveness of the RGA approach for developing molecular markers linked to disease resistance genes and demonstrated the efficiency of denaturing polyacrylamide‐gel electrophoresis for detecting polymerase chain reaction polymorphism.     

Chromosomal location of genes for resistance to powdery mildew in common wheat (Triticum aestivum L. em Thell.). 9. Gene MlZec1 from the Triticum dicoccoides-derived wheat line Zecoi-1

The Triticum dicoccoides-derived wheat line Zecoi-1 provides effective protection against powdery mildew. F₃ segregation analysis of Chinese Spring × Zecoi-1 hybrids showed that resistance in line Zecoi-1 is controlled by a single dominant gene. Amplified fragment length polymorphism (AFLP) analysis of bulked segregants from F₃s showing the homozygous resistant and susceptible phenotypes identified eight markers, of which four were associated with the resistance allele in repulsion phase. Following the assignment of these four repulsion phase AFLP markers to wheat chromosome 2B with the aid of Chinese Spring nulli-tetrasomic lines, they were physically mapped in the terminal breakpoint interval 0.89 (2BL-6)–1.00 (telomere) of chromosome 2BL. Genetic and physical mapping of simple sequence repeat markers from the distal half of chromosome 2BL located the wild emmer-derived powdery mildew resistance gene distal of breakpoint 0.89 in deletion line 2BL-6. Based on disease response patterns, genomic origin and chromosomal location the resistance gene in Zecoi-1 is temporarily designated MlZec1.     

Chromosome location of genes for resistance to powdery mildew in common wheat (Triticum aestivum L.) 1. Mlk and other alleles at the Pm3 locus

Summary Several wheat cultivars/lines were inoculated with isolates of Erysiphe graminis tritici to identify new genes/alleles for resistance. The wheats were tested with 13 isolates that had been characterized from responses on differential lines with known resistance genes. Gene Mlk which occurs in cultivars Kolibri, Syros, Ralle and several other European common wheats was found to be an allele at the Pm3 locus and is now designated Pm3d. The mildew resistance in an old Australian wheat, W150, is conferred by a single gene also allelic to Pm3 and now designated Pm3e. The near-isogenic line Michigan Amber/8^*Cc possesses another allele now designated Pm3f. A Syrian land variety of common wheat shows mildew resistance that is conditioned by the combination of genes Pm1 and Pm3a. Finally, two accessions of Triticum aestivum ssp. sphaerococcum appeared to possess the Pm3c allele.     

Growth habit in common wheat (Triticum aestivum L. em. Thell.)

Summary A study of the Vrn genotypes of 642 spring wheats supports the theory that only Vrn1, Vrn2 and Vrn3 exist in Tricticum aestivum. In none of the varieties investigated Vrn4 was present. Seven varieties, which according to literature carry Vrn4, showed to carry Vrn1, Vrn2 and/or Vrn3. Some varieties were mixtures of Vrn-genotypes, which could mislead geneticists in pooled data analysis. Other causes for misinterpretation of the data could be hybrid necrosis, hybrid dwarfness or a wrong determination of plants with a winter habitus. Only Hope was dominant on another Vrn locus. Its haploid Vrn-genotype is Vrn1 vrn2vrn3 Vrn5.     

Development of SCAR markers linked to the Pm21 gene conferring resistance to powdery mildew in common wheat

Powdery mildew is an important disease in most of the wheat production areas of the world. The resistance gene Pm21 (6AL/6VS trans-location) derived from Haynaldia villosa confers resistance to all available isolates of Erysiphe (Blumeria) graminis f. sp. tritici in China and Europe. The objective of this study was to develop fast and reliable sequence characterized amplified region (SCAR) markers linked to the Pm21 gene. A random amplified polymorphic DNA (RAPD) marker for Pm21, OPH17₁₄₀₀, was converted to SCAR markers after sequencing the two ends of the polymorphic DNA fragment. Two SCAR markers, SCAR₁₂₆₅ and SCAR₁₄₀₀, were developed to detect the Pm21 gene in different genetic backgrounds. The specific SCAR₁₂₆₅ marker enable large-scale accurate screening for the presence/absence of Pm21 allele.     

Identification of Powdery Mildew Resistance Genes in Common Wheat (Triticum aestivum L.)

Major genes for resistance to powdery mildew were analysed in 24 Czechoslovakian wheat cultivars and, in part, in their parents. For this purpose individual isolates of the pathogen, able to differentiate host lines with known resistance genes, were selected. Eight of nineteen winter wheat cultivars do not possess any major resistance gene. Three cultivars have one and seven have two genes. One cultivar carries a combination of three genes (Pm2, Pm4b, Pm8). The most common resistance genes are Pm4b, Pm5 and Pm8. Pm2 is once combined with Pm6. Only one of five spring cultivars lacked a major resistance gene. Mlk is once present alone and twice combined with Pm5. There is one spring cultivar with a novel combination of three genes: Pm1, Pm5 and another gene needing further characterization. The observations are discussed with additional results of parent lines and further information on pedigrees.     

Genetic studies of powdery mildew resistance in common oat (avena sativa L.) I. Cultivars and breeding lines grown in Western Europe and North America

Common oat (Avena sativa L.) cultivars and breeding lines grown in Western Europe and North America were tested against twelve Erysiphe graminis f. sp. avenae mildew isolates collected in Germany and Denmark. These isolates were selected for their ability to produce differential response patterns permitting characterization of five oat mildew resistance (OMR) groups. From a total of 259 cultivars and lines tested 173 accessions showed susceptible responses, 48 accessions were characterized by susceptible or intermediate responses and 38 accessions revealed isolate-specific resistance response patterns. Eight cultivars and lines had resistance patterns of OMR group 1, six cultivars of OMR group 2, and eleven cultivars of OMR group 3. Two cultivars, each possessed resistance patterns corresponding to OMR groups 1 + 3 and 2 + 3 in combination, respectively. Three cultivars exhibited the response pattern of OMR group 3 in association with an additional unknown resistance. Eight cultivars and lines showed a resistance response pattern not yet detected in the documented OMR groups. This revised version was published online in July 2006 with corrections to the Cover Date.     

The Growth in Culture of Detached Ovaries of Wheat (Triticum aestivum L. em. Thell)

Viable embryos have been recovered from fertilized wheat (Triticum aestivum) ovaries detached on the day of, or up to 7 days post-, anthesis and cultured aseptically fur up to 18 days. The most significant factors in determining the yield of embryos was the plating density, age and complexity of the explants. 26% of ovaries excised on the day of anthesis produced viable embryos if cultured as pairs of florets. The potential use of detached ovary cultures in gametophyte microinjection experiments, rescuing embryos from wide crosses and in chemically manipulating the early stages of embryo development are discussed.     

Potential for effective marker-assisted selection of three quantitative trait loci conferring adult plant resistance to powdery mildew in elite wheat breeding populations

Three quantitative trait loci (QTL) associated with adult plant resistance (APR) to powdery mildew (Blumeria graminis) in wheat (Triticum aestivum) cultivar ‘Massey’ were mapped in a previous study. The three QTL were located on chromosomes 2A, 2B and 1B, and explained 50% of the total phenotypic variation. A 293 recombinant inbred line (RIL) breeding population (UJ) derived from the cross of ‘USG 3209’, a derivative of ‘Massey’, and ‘Jaypee’ was used to evaluate the potential effectiveness of marker‐assisted selection (MAS) for APR. Powdery mildew severities of the 293 UJ RILs were evaluated in 2002 (F_{5 : 6}) and 2003 (F_{6 : 7}) under natural disease pressure in the field. The 293 RILs were also evaluated for disease severity in a 2004 (F_{7 : 8}) greenhouse experiment using a composite of five different isolates of B. graminis. Selection of RILs possessing the QTL on chromosome 2A, and to a lesser extent, the one on chromosome 1B was effective in identifying powdery mildew resistance in both greenhouse and field experiments. Overall, selecting RILs with QTL on chromosomes 2A and 2B was most successful in identifying highly resistant RILs, which had mean mildew severities of 4.4% and 3.2% in 2002 and 2003 field experiments, respectively. Breeders implementing MAS programs for APR to powdery mildew via selection of RILs containing the two QTL on chromosomes 2A and 2B likely will obtain RILs having high levels of resistance in the field, however combining all three QTL may ensure greater durability.     

Isolation, chromosomal location, and expression analysis of putative powdery mildew resistance genes in wheat (Triticum aestivum L.)

Special and degenerate primers are designed according to the conservative sequence of barley powdery mildew resistance genes Mla1, Mla6, and Mla13. Two wheat Mla-like orthologs, TaMla-2 and TaMla-3 are cloned and sequenced from the cDNA of wheat resistant-powdery mildew line TAM104R by RT-PCR method. TaMla-2 and TaMla-3 encode distinct but highly related coiled-coil nucleotide-binding site leucine-rich repeat type (NBS-LRR) resistant disease proteins and both reveal about 74 and 81% identity with amino acid sequence of Mla1, respectively. They are multiple copies in wheat genomes, one copy of them is mapped on wheat chromosome 1AL and two on 1BL using Chinese Spring nulli-tetra-somic lines and ditelosomic lines of 1A, 1B and 1D in southern analysis. This result suggests that may be the two Mla-like genes originated from the two diploid ancestral genomes, respectively. The expression pattern analysis of semi-quantitative PCR shows the TaMla genes are mainly expressed in leaf and sheath, and expression level is enhanced in organs infected by Erysiphe graminis, suggesting that TaMla-2 and TaMla-3 are powdery mildew resistance related-genes in wheat. Electronic Supplementary Material Supplementary material is available for this article at     

Reciprocal substitutions analysis of freezing resistance in wheat (Triticum aestivum L.)

To investigate the effects of individual chromosomes on freezing resistance, as well as their interactions with the genetic background, reciprocal sets of chromosome substitution lines between two hard red winter wheat cultivars, ‘Cheyenne’ and ‘Wichita’, were used in this study. Duplicate lines for each chromosome were included to check background homogeneity. Two experiments were carried out in complete block designs with two replications for each duplicate. Crown and leaf water content and leaf wet weight were measured in the field experiments. Crown survival, electrolyte leakage and 50% lethality temperature (LT50) were measured in the laboratory. The results showed that ‘Cheyenne’ was more resistant than ‘Wichita’. Crown survival was significantly correlated with crown water content, crown wet weight and electrolyte leakage. Chromosomes 6A, 3B and 5D substituted from ‘Wichita’ into ‘Cheyenne’ (‘CNN‐WI’), decreased the crown survival, and increased membrane stability, crown water content and crown wet weight of ‘Cheyenne’. Thus, these chromosomes from ‘Wichita’ decreased freezing resistance in ‘Cheyenne’. Reciprocally, chromosomes 5A, 5D, 3B and 4D from ‘Cheyenne’ into ‘Wichita’ increased crown survival and decreased crown water content and crown wet weight of ‘Wichita’. It was concluded that these chromosomes from ‘Cheyenne’ cause freezing resistance in ‘Wichita’ and carry freezing‐resistance genes.     

Evidence of allelism between genes Pm8 and Pm17 and chromosomal location of powdery mildew and leaf rust resistance genes in the common wheat cultivar'Amigo'

TIBL-1RS wheat-rye translocation cultivars utilized in wheat programmes worldwide carry powdery mildew resistance gene Pm8. Cultivar‘Amigo’possesses resistance gene Pm17 on its TIAL-1RS translocated chromosome. To be able to use Pm17efficiently in breeding programmes, this gene was transferred to a TIBL-1RS translocation in line Helami-105, and allelism between Pm8 and Pm17was studied. The progenies of the hybrids in the F₂ generation and F₃ families provided evidence that the two genes are allelic. Genetic studies using monosomic analyses confirmed that in cultivar‘Amigo', Pm17 and leaf rust resistance gene Lr24 are located on a translocated chromosome involving 1 A and 1B, respectively.