Primer of molecular marker closely linked to wheat powdery mildew resistance gene PmDR363, and application and method thereof

Abstract

This invention belongs to the field of agricultural biotechnology and relates to a wheat powdery mildew resistance gene. PmDR363 Primers for tightly linked molecular markers, their applications, and methods. These molecular markers are linked to genes. PmDR363 The genetic distance is only 1.7 cM, located on wheat chromosome 7BL. The primers include the upstream primer HENU1307-F, whose nucleotide sequence is shown in SEQ ID No. 1; and the downstream primer HENU1307-R, whose nucleotide sequence is shown in SEQ ID No. 2. This invention provides primers for a molecular marker tightly linked to the wheat powdery mildew resistance gene PmDR363 and their applications. This not only allows for rapid and accurate screening of target varieties, unaffected by environmental factors, and with clearly defined selection targets, but also saves production costs and greatly improves the selection efficiency and quality of high-quality powdery mildew-resistant wheat varieties or lines.

Description

Technical Field

[0001] This invention belongs to the field of agricultural biotechnology and relates to wheat genetics and breeding. Background Technology

[0002] Wheat powdery mildew is caused by powdery mildew fungus ( Blumeria graminis f. sp. tritici Powdery mildew is caused by infection with a fungus that is a typical obligate parasitic fungus, exhibiting significant host specificity across different species. Wheat can be infected by powdery mildew throughout its entire growth cycle. This pathogen primarily damages the leaf tissues of wheat, and in severe cases, can extend to the leaf sheaths, stems, and ears, adversely affecting normal growth, development, and yield (Ji et al. Mitochondrial characteristics of the powdery mildew genus Erysiphe revealed an extraordinary evolution in protein-coding genes. International Journal of Biological Macromolecules, 2023, 230: 123-153). Wheat powdery mildew, along with stripe rust and Fusarium head blight, is considered one of the three most serious and widespread fungal diseases affecting wheat production globally.

[0003] Currently, my country mainly uses agricultural control, chemical control, and biological agent control to control powdery mildew. However, each method has its own disadvantages, such as affecting planting density, increasing production costs, and pesticide spraying causing environmental pollution. Discovering superior powdery mildew-resistant genes and using molecular breeding technology to breed disease-resistant varieties is currently the most economical, safe, and effective measure for controlling this disease.

[0004] Currently, over one hundred major disease resistance genes have been discovered in wheat and its closely related species, namely ( Pm1-Pm71 ).in Pm1, Pm2, Pm3, Pm4, Pm5, Pm8, Pm24 These standardized loci carry multiple alleles. Pm1, Pm4 and Pm5 Each locus contains 5 alleles. Pm2 The site is Pm2a-Pm2c, Pm3 The number of sites is... Pm3a arrive Pm3r17 One allele, Pm24There are two alleles (Wang et al. Fighting wheat powdery mildew: fromgenes to fields. Theoretical and Applied Genetics, 2023, 136: 196). The germplasm resource bank of wheat powdery mildew resistance genes is divided into three levels, providing valuable genetic information resources for wheat genetic breeding. The first-level gene bank includes common wheat ( Triticum aestivum L., 2n=6x=42, AABBDD) and its subspecies; the secondary gene pool contains closely related species of one or two of the A, B, and D chromosome sets, such as durum wheat and Aegilops scabra; the tertiary gene pool includes closely related species that are not closely related to wheat, such as Leymus, barley, and rye.

[0005] durum wheat ( Triticum turgidum L. var. durum Desf., 2n=28, AABB) is a tetraploid wheat variety with a predominantly spring-growing habit. It exhibits good resistance to leaf rust, smut, and stripe rust, and is also cold-hardy and tolerant of poor soil. It has been identified as a strategic resource for overcoming bottlenecks in wheat breeding. Pm4b Derived from the durum wheat variety Yuma, research shows Pm4b Two transcripts are generated through alternative splicing. Pm4b_V1 and Pm4b_V2 Both factors together determine wheat's resistance to powdery mildew (Sánchez-Martín et al. Wheat Pm4 Resistance to powdery mildew is controlled by alternative splice variants encoding chimeric proteins. Nature Plants, 2021, 7: 327-341). PmDR147 It is a powdery mildew resistance gene derived from tetraploid wheat material, and PmDR147 Closely linked molecular markers Xgwm311 and Xgwm382 The specific polymorphism observed on chromosome 2AL suggests that... PmDR147 The gene is likely located here (Jia et al. Identification and microsatellite markers of aresistance gene to powdery mildew in common wheat introgressed from Triticum durum. Journal of Integrative Plant Biology, 2004, 46: 867-872). Pm68 Derived from the Greek durum wheat line TRI1796, it is located on 2BS and is linked by a pair of genetically linked lines. NLR (nucleotide-binding leucine repeat sequence) gene ( Pm68-1 and Pm68-2 Co-regulation (He et al. An NLR pair in the Pm68 locusconfers powdery mildew resistance in durum and common wheat. NatureCommunications, 2025, 16: 9039).

[0006] For a long time, the powdery mildew resistance genes used in wheat breeding have been relatively limited, leading to easy adaptation by the pathogen and a gradual loss of resistance in varieties. However, durum wheat, as a valuable genetic resource for wheat breeding, offers the potential to discover new powdery mildew resistance genes, breaking this pattern of limited resistance and improving the durability of wheat varieties' resistance. Although relatively few powdery mildew resistance genes have been discovered in durum wheat so far, it is believed that with advancements in genomics and molecular breeding technologies, more resistance genes will be discovered and utilized in the future. Summary of the Invention

[0007] This invention proposes a gene for resistance to powdery mildew in wheat. PmDR363 Tightly linked molecular markers and their detection applications provide guidance for breeding new wheat varieties resistant to powdery mildew.

[0008] The technical solution of this invention is implemented as follows: This invention provides a wheat powdery mildew resistance gene. PmDR363 Molecular markers HENU1307 This molecular marker is related to genes PmDR363 The genetic distance is only 1.7 cM, located on wheat chromosome 7BL at position 707886879 bp.

[0009] It also provides a gene for amplifying powdery mildew resistance in durum wheat. PmDR363 Primers for tightly linked molecular markers, wherein the molecular markers are codominant SSR markers. HENU1307 Its primers include upstream primers. HENU1307-F and downstream primers HENU1307-R .

[0010] The molecular marker HENU1307 The upstream primer is HENU1307-F Its nucleotide sequence is: 5'-GGCGAGGGAATTACGACGAG-3', as shown in SEQ ID No. 1; The molecular marker HENU1307 The downstream primer is HENU1307-R Its nucleotide sequence is: 5'-GTCAAGCCGCACCACCACTA-3', as shown in SEQ ID No. 2.

[0011] The amplified fruncal wheat powdery mildew resistance gene provided by this invention PmDR363 Primers for tightly linked molecular markers in genes PmDR363 The detection, identification, and auxiliary identification of wheat powdery mildew resistance traits, as well as its application in molecular marker-assisted breeding.

[0012] wheat powdery mildew resistance gene PmDR363 It is located on wheat chromosome 7BL (707,886,495 -708,990,535bp).

[0013] This invention also provides a method for detecting whether a wheat sample carries a flint wheat powdery mildew resistance gene. PmDR363 The method includes the following steps: (1) Extract genomic DNA from the wheat sample to be tested; (2) The genomic DNA of the wheat sample to be tested was amplified by PCR using the primers of the above molecular markers to obtain the amplification products; the molecular marker primers include the upstream primer. HENU1307-F and downstream primers HENU1307-R ; The molecular marker HENU1307 The upstream primer is HENU1307-F Its nucleotide sequence is: 5'-GGCGAGGGAATTACGACGAG-3', as shown in SEQ ID No. 1; The downstream primer of the molecular marker HENU1307 is HENU1307-R Its nucleotide sequence is: 5'-GTCAAGCCGCACCACCACTA-3', as shown in SEQ ID No. 2; (3) Electrophoresis and detection of the amplified products. If a specific band of 239 bp tightly linked can be amplified, it indicates that the wheat sample to be tested carries the flint wheat powdery mildew resistance gene. PmDR363 Otherwise, the wheat sample to be tested does not carry the powdery mildew resistance gene in durum wheat. PmDR363 .

[0014] The PCR amplification system in step (2) of the method is 10 μL, including: 1.0 μL of 50 ng / μL wheat genomic DNA, 4 μL of PCR Master Mix, 0.5 μL of 5 μM upstream primer, 0.5 μL of 5 μM downstream primer, and 4 μL of sterile deionized water.

[0015] The PCR amplification procedure in step (2) of the method is as follows: 94℃ pre-denaturation for 5 min; 94℃ denaturation for 15 s, 55℃ annealing for 15 s, extension for 40 s, 28 cycles; 72℃ extension for 10 min; storage at 12ºC.

[0016] The electrophoresis procedure for the amplification product in step (3) of the method is as follows: electrophoresis is performed on a non-denaturing polyacrylamide gel with a mass-volume percentage concentration of 8%. The amplification product is mixed with 2 μL of 6× loading buffer, and 1.0 μL of the mixture is loaded. Electrophoresis is performed at a constant voltage of 180 V for 1.5-2 h. After staining with silver nitrate, the sample is photographed.

[0017] The present invention has the following beneficial effects: 1. The durum wheat variety Jing DR363 used in this study was immune to powdery mildew at both the seedling and mature stages. Genetic analysis and molecular marker detection of powdery mildew resistance at the seedling stage showed that the resistance of Jing DR363 durum wheat seedlings to different types of wheat powdery mildew is determined by a pair of dominant genes. PmDR363 This novel wheat powdery mildew resistance gene / allele, located on wheat chromosome 7BL, is controlled by this gene. The wheat powdery mildew resistance gene provided by this invention... PmDR363 Molecular markers HENU1307 After genetic segregation population testing, and with genes PmDR363 The genetic distance is only 1.7 cM, which is similar to the gene. PmDR363 Tight linkage allows for accurate detection of genes. PmDR363 Genetic mapping of large populations, applied to genes PmDR363 Fine-grained localization and map-based cloning.

[0018] 2. This invention provides a gene for wheat resistance to powdery mildew. PmDR363 Primers for tightly linked molecular markers and their applications not only enable rapid and accurate screening of target varieties, unaffected by the environment, and with clear selection targets, but also save production costs and greatly improve the efficiency and quality of selecting high-quality powdery mildew-resistant wheat varieties or lines. Attached Figure Description

[0019] To more clearly illustrate the technical solutions in the embodiments of the present invention or the prior art, the drawings used in the description of the embodiments or the prior art will be briefly introduced below. Obviously, the drawings described below are only some embodiments of the present invention. For those skilled in the art, other drawings can be obtained based on these drawings without creative effort.

[0020] Figure 1 molecular markers HENU1307 Results of segregating populations of durum wheat DR363 and Langdon and their hybrid progeny. In the figure: M: D2000; 1: Durum wheat DR363 (powdery mildew resistant variety, carrying the powdery mildew resistance gene). PmDR363 ); 2: Langdon durum wheat (susceptible to powdery mildew); 3: 20 families mixed with resistance; 4: 20 families mixed with susceptibility; 5-24: F1 generation formed by crossing durum wheat Jing DR363 and durum wheat Langdon. 2:3 Family lineage, of which 5-9: homozygous disease-resistant F1 cells 2:3 Family history, 10-19: F strains of immune system separation 2:3 Family history, 20-24: Homozygous F 2:3 Family lineage; white arrows indicate areas where genes can be traced. PmDR363 Specific bands.

[0021] Figure 2 The graph shows the response of the parental strains DR363, Langdon, and the F1 hybrid offspring to strain E09. Detailed Implementation

[0022] The technical solution of the present invention will be clearly and completely described below with reference to the embodiments of the present invention. Obviously, the described embodiments are only some embodiments of the present invention, and not all embodiments. Based on the embodiments of the present invention, all other embodiments obtained by those of ordinary skill in the art without creative effort are within the scope of protection of the present invention.

[0023] Unless otherwise specified, the experimental methods used in the following experimental examples are conventional methods; the materials and reagents used are commercially available unless otherwise specified.

[0024] Example 1: Wheat powdery mildew resistance gene PmDR363 Development of primers for the molecular marker HENU1307 1. Materials Durum wheat DR363 and durum wheat Langdon showed resistance and susceptibility to wheat powdery mildew, respectively. Durum wheat DR363 and durum wheat Langdon were crossed, and the resulting F1 generation was self-crossed to obtain the F2 population and corresponding F1 generation.2:3 The family lineages are shown in Tables 1 and 2.

[0025] Table 1 shows the powdery mildew resistance phenotypes of plant materials.

[0026]

[0027] Table 2. Beijing DR363 and Langdon and their F2 and F 2:3 Segregation ratio of Bgt physiological race E09 inoculated at the seedling stage

[0028]

[0029] 2. Extraction of wheat genomic DNA The CTAB method for wheat genomic DNA extraction is as follows: (1) Cut fresh wheat leaves and grind them with liquid nitrogen. Take about 0.5 g and put it into a 2.0 mL centrifuge tube.

[0030] (2) Add 600 μL of CTAB extraction solution, incubate in a water bath at 65℃ for 40 min, and mix thoroughly every 10 min during the process.

[0031] (3) Add 600 μL of chloroform-isoamyl alcohol (24:1, v / v) and shake gently on a shaker for 15 min.

[0032] (4) Centrifuge at 12,000 rpm for 15 min, take the supernatant into another 2 mL centrifuge tube, add 3 times the volume of pre-cooled anhydrous ethanol, and place in a -20℃ refrigerator for precipitation for 60 min.

[0033] (5) Pick out the flocculent DNA precipitate and wash twice with 75% ethanol after pre-cooling.

[0034] (6) Pick out the DNA precipitate and place it in a 1.5 mL centrifuge tube. Let it air dry indoors.

[0035] (7) Add 60 μL of TE buffer (100 mM Tris-HCl, 10 mM EDTA). pH = 8.0) Dissolve the DNA precipitate to prepare a DNA storage solution.

[0036] (8) Dilute the DNA storage solution with ultrapure water to 20-50 ng / μL as a working solution for later use.

[0037] 3. Identification of powdery mildew resistance in wheat seedlings and genetic analysis of disease resistance The disease-resistant parent durum wheat Jing DR363, the disease-susceptible parent durum wheat Langdon, F1 hybrids, F2 populations, and F1 hybrids were compared. 2:3Families were planted in 128-cell trays (2×2cm), with 5 seeds sown per cell. Both parents and F1 strains had 20 seeds each. 2:3 Twenty seeds were sown from each family, while the susceptible control Zhoumai 18 was randomly sown and tagged for identification. After sowing, growth conditions were controlled as a 14-hour light / 10-hour dark cycle, temperature 20°C, and relative humidity 50-80%. When seedlings reached the one-leaf stage, they were inoculated with wheat powdery mildew strain E09. For the first 24 hours after inoculation, conditions were controlled as darkness, temperature 20°C, and relative humidity 100%. Afterward, conditions were controlled as a 14-hour light / 10-hour dark cycle, temperature 18-22°C, and relative humidity 100%. When the first leaf of the susceptible control Zhoumai 18 showed full disease, phenotypes were recorded according to a 0-4 scale, with 0-2 considered resistance and 3-4 considered susceptibility.

[0038] The survey results showed that durum wheat DR363 was immune to strain E09, while durum wheat Langdon (LDN) was highly susceptible to strain E09. When durum wheat DR363 was crossed with durum wheat Langdon, the F1 generation was resistant; of the 230 F2 plants, 174 were resistant and 56 were susceptible, consistent with a 3:1 segregation ratio of resistant to susceptible; 206 F2 plants showed... 2:3 In the family, 51 strains were homozygous resistant, 104 were heterozygous, and 51 were homozygous susceptible. The ratio of homozygous resistant to heterozygous to homozygous susceptible followed a segregation ratio of 1:2:1. Therefore, it is speculated that the resistance of durum wheat Jing DR363 to strain E09 is controlled by a single dominant gene, named gene [gene name missing]. PmDR363 .

[0039] 4. Preliminary molecular marker localization of the powdery mildew resistance gene in durum wheat Jing DR363 Based on phenotypic identification results, 20 homozygous resistant families and 20 homozygous susceptible families were selected to construct resistant and susceptible pools, respectively. Polymorphism detection was performed on durum wheat DR363 and Langdon, as well as the resistant and susceptible pools, using genome-wide uniformly distributed molecular markers. Ten pairs of markers showed consistent polymorphism in the resistant and susceptible parents and the resistant-susceptible pools. Subsequently, these markers were used to analyze the F1 generation of 206 durum wheat DR363 × Langdon durum wheat lines. 2:3 Genotyping of families, gene PmDR363 Preliminary localization to wheat chromosome 7BL.

[0040] 5. Genes PmDR363 Development of tightly linked molecular markers Based on the sequence information of the tetraploid wheat reference genome, *Durum wheat cv. Svevo* genome, Simple Sequence Repeat (SSR) markers were designed using Primer 5.0 software to target the F1 sequence of durum wheat *Jing DR363* × durum wheat *Langdon*. 2:3 Genotyping of the family pedigree yielded results related to the genes. PmDR363 Closely linked SSR tags HENU1307 The genetic distance is only 1.7 cM.

[0041] Molecular markers HENU1307 The primers consist of one upstream primer and one downstream primer: upstream primer HENU1307-F The nucleotide sequence is: 5'-GGCGAGGGAATTACGACGAG-3' (as shown in SEQ ID No. 1); Downstream primer HENU1307-R The nucleotide sequence is: 5'-GTCAAGCCGCACCACCACTA-3' (as shown in SEQ ID No. 2).

[0042] The PCR amplification system was 10 μL, comprising: 1.0 μL of 50 ng / μL wheat genomic DNA, 4 μL of PCR Master Mix, 0.5 μL of 5 μM upstream primer, 0.5 μL of 5 μM downstream primer, and 4 μL of sterile deionized water.

[0043] The PCR amplification program was as follows: 94℃ pre-denaturation for 5 min; 94℃ denaturation for 15 s, 55℃ annealing for 15 s, extension for 40 s, 28 cycles; 72℃ extension for 10 min; storage at 12℃.

[0044] The electrophoretic separation procedure for the amplified products is as follows: electrophoresis is performed on a non-denaturing polyacrylamide gel with a mass-volume percentage concentration of 8%. The amplified products are mixed with 2 μL of 6× loading buffer, and 1.0 μL of the mixture is loaded. Electrophoresis is performed at a constant voltage of 180V for 1.5-2 h. The gel is then stained with silver nitrate and photographed.

[0045] A specific 239 bp tightly linked band was amplified, indicating that the wheat sample carries the powdery mildew resistance gene. PmDR363 No specific band of 239 bp tightly linked was amplified, indicating that the wheat sample did not carry the gene for wheat resistance to powdery mildew. PmDR363 .

[0046] Example 2: Wheat powdery mildew resistance gene PmDR363 Molecular markers HENU1307 Application of primers Using wheat powdery mildew resistance genesPmDR363 Molecular markers HENU1307 Primer amplification detection of durum wheat DR363, durum wheat Langdon, and F1 hybrids formed by their crosses 2:3 The DNA extraction method for materials from the family lineage was the same as in Example 1.

[0047] Samples to be tested: durum wheat DR363, durum wheat Langdon, and their hybrids F1 2:3 Family lineage.

[0048] Genomic DNA was extracted from the above materials and used as a PCR amplification template, utilizing the molecular markers developed in this invention. HENU1307 Amplification was performed using primers: upstream primer HENU1307-F nucleotide sequence: 5'-GGCGAGGGAATTACGACGAG-3' (as shown in SEQ ID No. 1); Downstream primer HENU1307-R nucleotide sequence: 5'-GTCAAGCCGCACCACCACTA-3' (as shown in SEQ ID No. 2).

[0049] The PCR amplification system was 10 μL, comprising: 1.0 μL of 50 ng / μL wheat genomic DNA, 4 μL of PCR Master Mix, 0.5 μL of 5 μM upstream primer, 0.5 μL of 5 μM downstream primer, and 4 μL of sterile deionized water.

[0050] The PCR amplification program is as follows: 94ºC pre-denaturation for 5 min; 94ºC denaturation for 15 s, 55ºC annealing for 15 s, extension for 40 s, 28 cycles; 72ºC extension for 10 min; 12ºC storage.

[0051] The electrophoretic separation procedure for the amplified products is as follows: electrophoresis is performed on a non-denaturing polyacrylamide gel with a mass-volume percentage concentration of 8%. The amplified products are mixed with 2 μL of 6× loading buffer, and 1.0 μL of the mixture is loaded. Electrophoresis is performed at a constant voltage of 180V for 1.5-2 h. The mixture is then stained with silver nitrate and photographed.

[0052] Molecular marker detection results are shown below Figure 1 . Figure 1 For marking HENU1307 F derived from durum wheat DR363 × durum wheat Langdon 2:3 Partial amplification results from the family. In the figure: M: D2000; 1: Durum wheat Jing DR363 (a powdery mildew-resistant variety carrying the powdery mildew resistance gene).PmDR363 ); 2: Langdon durum wheat (susceptible to powdery mildew); 3: 20 families mixed with resistance; 4: 20 families mixed with susceptibility; 5-24: F1 generation formed by crossing durum wheat Jing DR363 and durum wheat Langdon. 2:3 Family lineage, of which 5-9: homozygous disease-resistant F1 cells 2:3 Family history, 10-19: F strains of immune system separation 2:3 Family history, 20-24: Homozygous F 2:3 Family lineage; white arrows indicate areas where genes can be traced. PmDR363 Specific bands. Amplification results show the markers. HENU1307 A 239 bp tightly linked specific band was amplified in the resistant parent durum wheat DR363 and the resistant family, but no such band was amplified in the susceptible parent durum wheat Langdon and the susceptible family.

[0053] wheat powdery mildew resistance gene PmDR363 Derived from the durum wheat variety Jing DR363, this gene has not yet been reported to be located or map-based cloned. The molecular markers provided in this invention... HENU1307 Detecting large populations for genetic mapping is helpful for gene sequencing. PmDR363 Fine-grained localization and map-based cloning are crucial for gene sequencing. PmDR363 The in-depth analysis of the efficient breeding and disease resistance mechanisms is of great significance.

[0054] The above description is only a preferred embodiment of the present invention and is not intended to limit the present invention. Any modifications, equivalent substitutions, improvements, etc., made within the spirit and principles of the present invention should be included within the protection scope of the present invention.

Claims

1. Wheat powdery mildew resistance gene PmDR363 Primers with tightly linked molecular markers, characterized by: The molecular marker is located on wheat chromosome 7BL at position 707886879 bp.

2. The wheat powdery mildew resistance gene according to claim 1 PmDR363 Primers with tightly linked molecular markers, characterized by: The primers include upstream primers. HENU1307-F and downstream primers HENU1307-R upstream primer HENU1307-F The nucleotide sequence is shown in SEQ ID No. 1; the downstream primer HENU1307-R The nucleotide sequence is shown in SEQ ID No.

2.

3. A reagent kit, characterized in that: It contains the primers as described in claim 2.

4. The primers described in claim 1 or 2 for detecting wheat powdery mildew resistance genes. PmDR363 Applications in [the context of the text].

5. The primers according to claim 1 or 2 in wheat powdery mildew resistance gene PmDR363 Applications in map-based cloning.

6. The application of the primers described in claim 1 or 2 in the breeding of wheat varieties resistant to powdery mildew.

7. A method for detecting whether wheat carries a gene for resistance to powdery mildew. PmDR363 The method is characterized by, Includes the following steps: (1) Extract genomic DNA from the wheat sample to be tested; (2) Using the primers described in claim 2, the genomic DNA of the wheat sample to be tested is detected by PCR amplification to obtain the amplification product; (3) Perform electrophoresis on the amplification product of step (2) and detect whether it carries the wheat powdery mildew resistance gene. PmDR363 .

8. The method for detecting whether wheat carries the powdery mildew resistance gene as described in claim 7 PmDR363 The method is characterized by: The PCR amplification system in step (2) is 10 μL, including 1.0 μL of 50 ng / μL wheat genomic DNA, 4 μL of PCR MasterMix, 0.5 μL of 5 μM upstream primer, 0.5 μL of 5 μM downstream primer, and 4 μL of sterile deionized water.

9. The method for detecting whether wheat carries the powdery mildew resistance gene as described in claim 7. PmDR363 The method is characterized by: The PCR amplification procedure in step (2) is as follows: pre-denaturation at 94℃ for 5 min; denaturation at 94℃ for 15 s, annealing at 55℃ for 15 s, extension for 40 s, 28 cycles; extension at 72℃ for 10 min; storage at 12℃.

10. The method for detecting whether wheat carries a powdery mildew resistance gene according to any one of claims 7-9 PmDR363 The method is characterized by: If the amplification product contains a specific band of 239 bp, it indicates that the wheat sample being tested contains a gene for resistance to powdery mildew. PmDR363 Conversely, the wheat sample tested did not contain the wheat powdery mildew resistance gene. PmDR363 .

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