Powdery mildew resistance gene <Emphasis Type="Italic">Pm22</Emphasis> in cultivar Virest is a member of the complex <Emphasis Type="Italic">Pm1</Emphasis> locus in common wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L. em Thell.)

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.     

Chromosomal location of genes for resistance to powdery mildew in common wheat (Triticum aestivum L. em Thell.) 6. Alleles at the Pm5 locus

Genetic characterization of powdery mildew resistance genes were conducted in common wheat cultivars Hope and Selpek possessing resistance gene Pm5, cvs. Ibis and Kormoran expressing resistance gene Mli, a backcross-derived line IGV 1–455 and a Triticum sphaerococcum var. rotundatum Perc. line Kolandi. Monosomic analyses revealed that one major recessive gene is located on chromosome 7B in the lines IGV 1–455 and Kolandi. Allelism tests of the F₂ and F₃ populations involving the tested resistant lines crossed with either cv. Hope or Selpek indicated that their resistance genes are alleles at the Pm5 locus. The alleles are now designated Pm5a in Hope and Selpek, Pm5b in Ibis and Kormoran, Pm5c in T. sphaerococcum var. rotundatum line Kolandi, and Pm5d in backcross-derived line IGV 1–455, respectively. Received: 5 November 1999 / Accepted: 14 April 2000     

Development of functional markers specific for seven Pm3 resistance alleles and their validation in the bread wheat gene pool

In the ideal case, molecular markers used for marker-assisted selection are allele-specific even if the alleles differ only by a few nucleotide polymorphisms within the coding sequence of target genes. Such ‘perfect’ markers are completely correlated with the trait of interest. In hexaploid wheat (Triticum aestivum L.) the Pm3 locus encodes seven alleles (Pm3a–Pm3g) conferring resistance to different races of Blumeria graminis f.sp. tritici, the agent of powdery mildew, a major disease of bread wheat. All Pm3 alleles are known at the molecular level. Here, we generated specific markers for the Pm3 alleles based on nucleotide polymorphisms of coding and adjacent non-coding regions. The specificity of these markers was validated in a collection of 93 modern or historically important cultivars and breeding lines of wheat and spelt (Triticum spelta L.). These markers confirmed the presence of the predicted Pm3 alleles in 31 varieties and lines known to carry Pm3 resistance alleles. In a few varieties, Pm3 alleles different from alleles previously described based on pathogenicity tests or tightly linked markers were observed. In all these cases, the identity of the marker-detected Pm3 alleles was confirmed by DNA sequence analysis. Pm3 markers confirmed the absence of known Pm3 resistance alleles in 54 European wheat and spelt varieties in which Pm3 alleles had not been previously identified. These results indicate that the developed markers are highly diagnostic for specific Pm3 resistance alleles in a wide range of varieties and breeding lines, and will be useful (1) for identifying Pm3 alleles in the wheat gene pool, (2) for efficient marker-assisted selection of these genes, and (3) for combining multiple Pm3 alleles within a single cultivar through transgenic approaches.Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

<Emphasis Type="Italic">Pm37</Emphasis>, a new broadly effective powdery mildew resistance gene from <Emphasis Type="Italic">Triticum </Emphasis><Emphasis Type="Italic">timopheevii</Emphasis>

Powdery mildew is an important foliar disease in wheat, especially in areas with a cool or maritime climate. A dominant powdery mildew resistance gene transferred to the hexaploid germplasm line NC99BGTAG11 from T. timopheevii subsp. armeniacum was mapped distally on the long arm of chromosome 7A. Differential reactions were observed between the resistance gene in NC99BGTAG11 and the alleles of the Pm1 locus that is also located on chromosome arm 7AL. Observed segregation in F_2:3 lines from the cross NC99BGTAG11 × Axminster (Pm1a) demonstrate that germplasm line NC99BGTAG11 carries a novel powdery mildew resistance gene, which is now designated as Pm37. This new gene is highly effective against all powdery mildew isolates tested so far. Analyses of the population with molecular markers indicate that Pm37 is located 16 cM proximal to the Pm1 complex. Simple sequence repeat (SSR) markers Xgwm332 and Xwmc790 were located 0.5 cM proximal and distal, respectively, to Pm37. In order to identify new markers in the region, wheat expressed sequence tags (ESTs) located in the distal 10% of 7AL that were orthologous to sequences from chromosome 6 of rice were targeted. The two new EST-derived STS markers were located distal to Pm37 and one marker was closely linked to the Pm1a region. These new markers can be used in marker-assisted selection schemes to develop wheat cultivars with pyramids of powdery mildew resistance genes, including combinations of Pm37 in coupling linkage with alleles of the Pm1 locus.     

Analysis of the resistance of Aegilops squarrosa to the wheatgrass mildew fungus by using the gene-for-gene reationship

Summary Pm10 and Pm15, resistance genes to Erysiphe graminis f. sp. agropyri, are located on the D genome of common wheat. It was determined whether or not they were carried by existing lines of the D genome donor, Aegilops squarrosa, using the gene-for-gene relationship. Two lines of Ae. squarrosa tested (one was var. meyeri and the other was var. anathera) were susceptible to culture Tk-1 of E. graminis f. sp. tritici and were highly resistant to culture Ak-1 of E. graminis f. sp. agropyri. The two lines were inoculated with an F₁ population derived from the cross Ak-1 × Tk-1. Comparative analyses of the segregation patterns revealed that Ppm10 and Ppm15, avirulence genes corresponding to Pm10 and Pm15, respectively, are involved in the avirulence of Ak-1 on var. meyeri and var. anathera, respectively. According to the gene-for-gene relationship, var. meyeri and var. anathera were inferred to carry Pm10 and Pm15, respectively. Analysis with a synthetic hexaploid confirmed the inference.     

Location of a gene regulating cold-induced carbohydrate production on chromosome 5A of wheat

 Major changes in osmotic potential during cold acclimation are due to changes in sugar concentration, and there is a good correlation between sugar content and frost tolerance. The objective of the present study was to localize a gene(s) responsible for carbohydrate accumulation during cold acclimation on chromosome 5A of wheat using recombinant lines developed from the cross between the substitution lines Chinese Spring (Cheyenne 5A) and CS(Triticum spelta 5A). Previously, major genes influencing frost resistance (Fr1) and vernalization requirement (Vrn1) had been localized on the long arm of that chromosome. The T. spelta 5A chromosome carrying the Fr1 (frost-sensitive) allele for frost tolerance and the Vrn1 (spring-habit) allele for vernalization requirement did not have a major effect on the sucrose and fructan contents in the Chinese Spring background. On the other hand, the presence of Cheyenne alleles for vernalization requirement, vrn1, and frost tolerance, fr1, significantly increased sugar concentrations. A recombinant line thought to exhibit recombination between the Vrn1 and Fr1 loci suggested that the gene regulating sucrose accumulation was closely associated with, or else represented a pleiotropic effect of, Vrn1, but was separable from the Fr1 locus. Received: 3 March 1997 / Accepted: 7 March 1997     

The powdery mildew resistance gene Pm8 derived from rye is suppressed by its wheat ortholog Pm3

The powdery mildew resistance gene Pm8 derived from rye is located on a 1BL.1RS chromosome translocation in wheat. However, some wheat lines with this translocation do not show resistance to isolates of the wheat powdery mildew pathogen avirulent to Pm8 due to an unknown genetically dominant suppression mechanism. Here we show that lines with suppressed Pm8 activity contain an intact and expressed Pm8 gene. Therefore, the absence of Pm8 function in certain 1BL.1RS‐containing wheat lines is not the result of gene loss or mutation but is based on suppression. The wheat gene Pm3, an ortholog of rye Pm8, suppressed Pm8‐mediated powdery mildew resistance in lines containing Pm8 in a transient single‐cell expression assay. This result was further confirmed in transgenic lines with combined Pm8 and Pm3 transgenes. Expression analysis revealed that suppression is not the result of gene silencing, either in wheat 1BL.1RS translocation lines carrying Pm8 or in transgenic genotypes with both Pm8 and Pm3 alleles. In addition, a similar abundance of the PM8 and PM3 proteins in single or double homozygous transgenic lines suggested that a post‐translational mechanism is involved in suppression of Pm8. Co‐expression of Pm8 and Pm3 genes in Nicotiana benthamiana leaves followed by co‐immunoprecipitation analysis showed that the two proteins interact. Therefore, the formation of a heteromeric protein complex might result in inefficient or absent signal transmission for the defense reaction. These data provide a molecular explanation for the suppression of resistance genes in certain genetic backgrounds and suggest ways to circumvent it in future plant breeding.     

Chromosomal location of genes for resistance to powdery mildew in Chinese wheat lines Jieyan 94-1-1 and Siyan 94-1-2

Two Chinese wheat lines Jieyan 94-1-1 and Siyan 94-1-2 are resistant to all 120 isolates of Blumeria graminis f. sp. tritici maintained in Weihenstephan, Germany. Monosomic analyses employing the susceptible set of 21 Chinese Spring monosomic lines revealed that the line Jieyan 94-1-1 carries one dominant gene on translocated wheat/rye chromosome 1B/1R and one recessive gene on chromosome 7B, whereas line Siyan 94-1-2 possesses one recessive gene on chromosome 7B and one dominant gene on chromosome 5D. Allelism tests in combination with the use of specific isolates comfirmed that the dominant genes in Jieyan 94-1-1 and Siyan 94-1-2 are Pm8 and Pm2, respectively. The recessive genes present in each of the two lines are shown to be new alleles located on chromosome 7B at the pm 5 locus. The two genes are tentatively designated mljy in Jieyan 94-1-1 and mlsy in Siyan 94-1-2, respectively.     

PCR-based markers for the powdery mildew resistance gene<Emphasis Type="Italic"> Pm4a</Emphasis> in wheat

Gene tagging is the basis of marker-assisted selection and map-based cloning. To develop PCR-based markers for Pm4a, a dominant powdery mildew resistance gene of wheat, we surveyed 46 group 2 microsatellite markers between Pm4a near-isogenic line (NIL) CI 14124 and the recurrent parent Chancellor (Cc). One of the markers, gwm356, detected polymorphism and was used for genotyping an F₂ population of 85 plants derived from CI 14124 × Cc. Linkage mapping indicated that Xgwm356 was linked to Pm4a at a distance of 4.8 cM. To identify more PCR-based markers for Pm4a, we also converted the restriction fragment length polymorphism marker BCD1231 linked to it into a sequence-tagged site (STS) marker. The STS primer designed based on the end sequences of BCD1231 amplified an approximately 1.6-kb monomorphic band in both parents. Following digestion of the products with the four-cutter enzymes HaeIII and MspI, it was discovered that the band from CI 14124 consisted of at least two products, one of which had a digestion pattern different from the band from Cc. In the F₂ population, the cleaved polymorphism revealed by the STS marker between the parents co-segregated with powdery mildew resistance. To design Pm4a-specific PCR markers, the 1.6-kb band from Cc and the fragment associated with Pm4a in CI 14124 were sequenced and compared. Based on these sequences a new PCR marker was identified, which detected a 470-bp product only in the Pm4a-containing lines. These PCR-based markers provide a cost-saving option for marker-assisted selection of Pm4a.Communicated by F. Salamini     

Intragenic allele pyramiding combines different specificities of wheat Pm3 resistance alleles

Some plant resistance genes occur as allelic series, with each member conferring specific resistance against a subset of pathogen races. In wheat, there are 17 alleles of the Pm3 gene. They encode nucleotide‐binding (NB‐ARC) and leucine‐rich‐repeat (LRR) domain proteins, which mediate resistance to distinct race spectra of powdery mildew. It is not known if specificities from different alleles can be combined to create resistance genes with broader specificity. Here, we used an approach based on avirulence analysis of pathogen populations to characterize the molecular basis of Pm3 recognition spectra. A large survey of mildew races for avirulence on the Pm3 alleles revealed that Pm3a has a resistance spectrum that completely contains that of Pm3f, but also extends towards additional races. The same is true for the Pm3b and Pm3c gene pair. The molecular analysis of these allelic pairs revealed a role of the NB‐ARC protein domain in the efficiency of effector‐dependent resistance. Analysis of the wild‐type and chimeric Pm3 alleles identified single residues in the C‐terminal LRR motifs as the main determinant of allele specificity. Variable residues of the N‐terminal LRRs are necessary, but not sufficient, to confer resistance specificity. Based on these data, we constructed a chimeric Pm3 gene by intragenic allele pyramiding of Pm3d and Pm3e that showed the combined resistance specificity and, thus, a broader recognition spectrum compared with the parental alleles. Our findings support a model of stepwise evolution of Pm3 recognition specificities.     

Pyramiding of transgenic <Emphasis Type="Italic">Pm3</Emphasis> alleles in wheat results in improved powdery mildew resistance in the field

Key message

The combined effects of enhanced total transgene expression level and allele-specificity combination in transgenic allele-pyramided Pm3 wheat lines result in improved powdery mildew field resistance without negative pleiotropic effects.

Abstract

Allelic Pm3 resistance genes of wheat confer race-specific resistance to powdery mildew (Blumeria graminis f. sp. tritici, Bgt) and encode nucleotide-binding domain, leucine-rich repeat (NLR) receptors. Transgenic wheat lines overexpressing alleles Pm3a, b, c, d, f, and g have previously been generated by transformation of cultivar Bobwhite and tested in field trials, revealing varying degrees of powdery mildew resistance conferred by the transgenes. Here, we tested four transgenic lines each carrying two pyramided Pm3 alleles, which were generated by crossbreeding of lines transformed with single Pm3 alleles. All four allele-pyramided lines showed strongly improved powdery mildew resistance in the field compared to their parental lines. The improved resistance results from the two effects of enhanced total transgene expression levels and allele-specificity combinations. In contrast to leaf segment tests on greenhouse-grown seedlings, no allelic suppression was observed in the field. Plant development and yield scores of the pyramided lines were similar to the mean scores of the corresponding parental lines, and thus, the allele pyramiding did not cause any negative effects. On the contrary, in pyramided line, Pm3b × Pm3f normal plant development was restored compared to the delayed development and reduced seed set of parental line Pm3f. Allele-specific RT qPCR revealed additive transgene expression levels of the two Pm3 alleles in the pyramided lines. A positive correlation between total transgene expression level and powdery mildew field resistance was observed. In summary, allele pyramiding of Pm3 transgenes proved to be successful in enhancing powdery mildew field resistance.

     

Compatibility of Erysiphe graminis Hybrids f. sp. secalis X f. sp. Tritici with Wheat Lines Involving Resistance Genes from Rye

Compatibility of hybrid cultures Erysiphe graminis ff. sp. secalis (SI) ×tritici (t2) was tested in the laboratory with wheat cultivars involving different resistance genes and with two rye cultivars. Segregation was observed on wheat without resistance gene and with resistance genes Pm1, Pm3b and Pm3c compatible with t2, but not on wheat with resistance gene Pm2, Pm 3a, Pm 4a and Pm 5 incompatible with t2, nor on rye. It was obvious that S1 involves avirulence genes to Pm1, Pm2, Pm 3a, pm 3b, Pm 3c, Pm 4a, Pm 5. Segregation was found on wheat cultivars involving rye resistance genes Pm 7 (Transfed) and Pm 8 (Kavkaz), but cv. Transec (Pm7) was incompatible with all cultures used, because Transec involves another gene for resistance. The results indicate that hybridization between formae speciales secalis and tritici of the fungus can be a source of fungus compatibility with wheat with rye resistance, even in field conditions.     

Selection of human cells having two different types of mutations in individual cells (genetic/artificial mutants)

Summary We have previously reported the establishment and characterization of B cell lines from patients and family members with various types of adenine phosphoribosyltransferase (APRT) deficiencies. These cell lines contain, at the APRT locus, three different alleles (APRT ^* 1, APRT ^* Q0, and APRT ^* J) that are clearly distinguishable from each other. From five genetically heterozygous cell lines with two different genotypes (APRT ^* 1/APRTQ0 and APRT ^* 1/APRT ^* J), we have selected 48 clones resistant to 2,6-diaminopurine. Resistance to this adenine analogue is a characteristic of cells having defects in both of the APRT alleles in individual cells. The mutant clones from a cell line from a complete-type heterozygote had APRT activities close to zero (mean=0.04 nmol/min per milligram protein) in the cell extracts, while 15 clones from four cell lines from the four Japanese-type heterozygotes had significant enzyme activities (mean=3.88 nmol/min per milligram protein). Kinetic studies on two of the mutants from two Japancse-type heterozygous cell lines have shown that affinity to substrate 5-phosphoribosyl-1-pyrophosphate was reduced, indicating that APRT in those clones reflected the characteristics of the Japanese-type enzyme. The data presented here indicate that clones we obtained are genetic/artificial mutants, each having a genetic mutation in a single allele (APRT ^* J or APRT ^* Q0) and an artificially produced mutation in the other previously functional allele (APRT ^*1). The present procedure provided the only diagnostic method for Japanese-type APRT heterozygotes (APRT ^* 1/APRT ^* J).     

Genetic characterization of powdery mildew resistance in U.S. hard winter wheat

Powdery mildew significantly affects grain yield and end-use quality of winter wheat in the southern Great Plains. Employing resistance resources in locally adapted cultivars is the most effective means to control powdery mildew. Two types of powdery mildew resistance exist in wheat cultivars, i.e., qualitative and quantitative. Qualitative resistance is controlled by major genes, is race-specific, is not durable, and is effective in seedlings and in adult plants. Quantitative resistance is controlled by minor genes, is non-race-specific, is durable, and is predominantly effective in adult plants. In this study, we found that the segregation of powdery mildew resistance in a population of recombinant inbred lines developed from a cross between the susceptible cultivar Jagger and the resistant cultivar 2174 was controlled by a major QTL on the short arm of chromosome 1A and modified by four minor QTLs on chromosomes 1B, 3B, 4A, and 6D. The major QTL was mapped to the genomic region where the Pm3 gene resides. Using specific PCR markers for seven Pm3 alleles, 2174 was found to carry the Pm3a allele. Pm3a explained 61% of the total phenotypic variation in disease reaction observed among seedlings inoculated in the greenhouse and adult plants grown in the field and subjected to natural disease pressure. The resistant Pm3a allele was present among 4 of 31 cultivars currently being produced in the southern Great Plains. The genetic effects of several minor loci varied with different developmental stages and environments. Molecular markers associated with these genetic loci would facilitate incorporating genetic resistance to powdery mildew into improved winter wheat cultivars.     

Transgenic Pm3b wheat lines show resistance to powdery mildew in the field

Plant resistance (R) genes are highly effective in protecting plants against diseases, but pathogens can overcome such genes relatively easily by adaptation. Consequently, in many cases R genes do not confer durable resistance in agricultural environments. One possible strategy to make the use of R genes more sustainable depends on the modification of R genes followed by transformation. To test a possible transgenic use of R genes, we overexpressed in wheat the Pm3b resistance gene against powdery mildew under control of the maize ubiquitin promoter. Four independent transgenic lines were tested in the greenhouse and the field during 3 years. The four lines showed a five‐ to 600‐fold transgene overexpression compared with the expression of the endogenous Pm3b gene in the landrace ‘Chul’. Powdery mildew resistance was significantly improved in all lines in the greenhouse and the field, both with naturally occurring infection or after artificial inoculation. Under controlled environmental conditions, the line with the strongest overexpression of the Pm3b gene showed a dramatic increase in resistance to powdery mildew isolates that are virulent on the endogenous Pm3b. Under a variety of field conditions, but never in the greenhouse, three of the four transgenic lines showed pleiotropic effects on spike and leaf morphology. The highest overexpressing line had the strongest side effects, suggesting a correlation between expression level and phenotypic changes. These results demonstrate that the successful transgenic use of R genes critically depends on achieving an optimal level of their expression, possibly in a tissue‐specific way.     

Allelic variation of the HMW glutenin subunits in Spanish accessions of spelt wheat (Triticum aestivum ssp. spelta L. em. Thell.)

Spelt wheat (Triticum aestivum ssp. spelta L. em. Thell.) is a hulled wheat of Germanic origin that survives at marginal areas in Asturias (Spain). The HMW glutenin subunit composition of 403 accessions of spelt wheat from Spain has been analysed by SDS-PAGE. Three allelic variants were detected for Glu-A1. For the Glu-B1 locus, two of seven alleles detected have not been found before; while four of nine alleles detected for the Glu-D1 are not previously described. Considering the three loci, twenty five combinations were found among all the evaluated lines. This wide polymorphism could be used to transfer new quality genes to wheat, and widen the genetic basis of them. Received: 19 September 2000 / Accepted: 20 October 2000     

RFLP markers linked to powdery mildew resistance genes Pm1, Pm2, Pm3, and Pm4 in wheat.

Near-isogenic lines (NILs) and their recurrent parent Chancellor (Cc) were used to identify restriction fragment length polymorphic markers linked to powdery mildew (Blumeria graminis (DC.) E.O. Speer f.sp. tritici) resistance genes Pm1, Pm2, Pm3, and Pm4 in wheat (Triticum aestivum L. em. Thell). By mapping these polymorphic markers in F2 progenies from crosses of the NILs with Cc, it was found that Pm1 cosegregated with a polymorphic locus detected by DNA probe CDO347; Pm2 was linked to a locus detected by probe BCD1871 with a distance of 3.5 cM; Pm3b was linked to a locus detected by probe BCD1434 with a distance of 1.3 cM; Pm4a cosegregated with Xbcd1231-2A(2) and Xcdo678-2A, and was closely flanked by Xbcd1231-2A(1) and Xbcd292-2A both with a distance of 1.5 cM. Aneuploid mapping of these markers indicated that locus Xcdo347-7A is on 7AL, Xbcd1871-5D on 5DS, Xbcd1434-1A on 1AS, and loci Xbcd292-2A and Xcdo678-2A are on 2AL. The same polymorphic fragments detected in the Pm3b NIL by Xbcd1434-1A were found in Pm3a NIL using several enzyme digestions.     

Genetic and physical characterization of theLR1 leaf rust resistance locus in wheat (Triticum aestivum L.)

The objective of this study was to characterize the leaf rust resistance locusLr1 in wheat. Restriction fragment length polymorphism (RELP) analysis was performed on the resistant lineLr1/6^*Thatcher and the susceptible varieties Thatcher and Frisal, as well as on the segregating F₂ populations. Seventeen out of 37 RFLP probes mapping to group 5 chromosomes showed polymorphism betweenLr1/6^*Thatcher and Frisal, whereas 11 probes were polymorphic between the near-isogenic lines (NILs)Lr1/6^*Thatcher and Thatcher. Three of these probes were linked to the resistance gene in the segregating F₂ populations. One probe (pTAG621) showed very tight linkage toLr1 and mapped to a single-copy region on chromosome 5D. The map location of pTAG621 at the end of the long arm of chromosome 5D was confirmed by the absence of the band in the nulli-tetrasomic line N5DT5B of Chinese Spring and a set of deletion lines of Chinese Spring lacking the distal part of 5DL. Twenty-seven breeding lines containing theLr1 resistance gene in different genetic backgrounds showed the same band asLr1/6^*Thatcher when hybridized with pTAG621. The RFLP marker was converted to a sequence-tagged-site marker using polymerase chain reaction (PCR) amplification. Sequencing of the specific fragment amplified from both NILs revealed point mutations as well as small insertion/deletion events. These were used to design primers that allowed amplification of a specific product only from the resistant lineLr1/6^*Thatcher. This STS, specific for theLr1 resistance gene, will allow efficient selection for the disease resistance gene in wheat breeding programmes. In addition, the identification of a D-genome-specific probe tightly linked toLr1 should ultimately provide the basis for positional cloning of the gene.     

Chromosomal location of a gene suppressing powdery mildew resistance genes Pm8 and Pm17 in common wheat (Triticum aestivum L. em. Thell.)

The chromosomal location of a suppressor for the powdery mildew resistance genes Pm8 and Pm17 was determined by a monosomic set of the wheat cultivar Caribo. This cultivar carries a suppressor gene inhibiting the expression of Pm8 in cv Disponent and of Pm17 in line Helami-105. In disease resistance assessments, monosomic F₁ hybrids (2n=41) of Caribo x Disponent and Caribo x Helami-105 lacking chromosome 7D were resistant, whereas monosomic F₁ hybrids involving the other 20 chromosomes, as well as disomic F₁ hybrids (2n=42) of all cross combinations, were susceptible revealing that the suppressor gene for Pm8 and Pm17 is localized on chromosome 7D. It is suggested that genotypes without the suppressor gene be used for the exploitation of genes Pm8 and Pm17 in enhancing powdery mildew resistance in common wheat.     

Multiple Avirulence Loci and Allele-Specific Effector Recognition Control the Pm3 Race-Specific Resistance of Wheat to Powdery Mildew

In cereals, several mildew resistance genes occur as large allelic series; for example, in wheat (Triticum aestivum and Triticum turgidum), 17 functional Pm3 alleles confer agronomically important race-specific resistance to powdery mildew (Blumeria graminis). The molecular basis of race specificity has been characterized in wheat, but little is known about the corresponding avirulence genes in powdery mildew. Here, we dissected the genetics of avirulence for six Pm3 alleles and found that three major Avr loci affect avirulence, with a common locus_1 involved in all AvrPm3-Pm3 interactions. We cloned the effector gene AvrPm3^a2/f2 from locus_2, which is recognized by the Pm3a and Pm3f alleles. Induction of a Pm3 allele-dependent hypersensitive response in transient assays in Nicotiana benthamiana and in wheat demonstrated specificity. Gene expression analysis of Bcg1 (encoded by locus_1) and AvrPm3 ^a2/f2 revealed significant differences between isolates, indicating that in addition to protein polymorphisms, expression levels play a role in avirulence. We propose a model for race specificity involving three components: an allele-specific avirulence effector, a resistance gene allele, and a pathogen-encoded suppressor of avirulence. Thus, whereas a genetically simple allelic series controls specificity in the plant host, recognition on the pathogen side is more complex, allowing flexible evolutionary responses and adaptation to resistance genes.