The introduction into bread wheat of a major gene for resistance to powdery mildew from wild emmer wheat

Summary A new source of resistance to wheat powdery mildew caused by Erysiphe graminis has been transferred to hexaploid bread wheat, Triticum aestivum, from the wild tetraploid wheat, Triticum dicoccoides. The donor was crossed to bread wheat and the pentaploid progeny was then self-pollinated. Plants having a near stable hexaploid chromosome complement were selected in the F₃ progeny and topcrossing and backcrossing of these to a second wheat cultivar to improve the phenotype was undertaken. Monosomic analysis of early backcross lines showed the transferred gene to be located on chromosome 4A. The gene has been designated Pm16.     

Molecular characterization of a novel powdery mildew resistance gene Pm30 in wheat originating from wild emmer

Powdery mildew caused by Erysiphe graminis f. sp. tritici is one of the most important wheat diseases in many regions of theworld. A powdery mildew resistance gene, originating from wild emmerwheat (Triticum dicoccoides) accession `C20', from Rosh Pinna, Israel,was successfully transferred to hexaploid wheat through crossing andbackcrossing. Genetic analysis indicated that a single dominant genecontrols the powdery mildew resistance at the seedling stage. SegregatingBC₁F₂ progenies of the cross 87-1/C20//2*8866 wereused for bulked segregant analysis (BSA). The PCR approach was used togenerate polymorphic DNA fragments between the resistant and susceptibleDNA pools by use of 10-mer random primers, STS primers, and wheatmicrosatellite primers. Three markers, Xgwm159/430,Xgwm159/460, and Xgwm159/500, were found to be linked tothe resistance gene. After evaluating the polymorphic markers in twosegregating populations, the distance between the markers and the mildewresistance gene was estimated to be 5–6 cM. By means of ChineseSpring nullisomic-tetrasomics and ditelosomics, the polymorphic markersand the resistance gene were assigned to chromosome arm 5BS and werephysically mapped on the gene rich regions of fragment length (FL) 0.41–0.43 by Chinese Spring deletion lines. As no powdery mildew resistancegene has been reported on chromosome arm 5BS, the mildew resistancegene originating from C20 should be a new gene and is designated Pm30.     

Identification and genetic mapping of a powdery mildew resistance gene in wild emmer (<Emphasis Type="Italic">Triticum dicoccoides</Emphasis>) accession IW72 from Israel

Powdery mildew, caused by Blumeria graminis f. sp. tritici (Bgt), is a devastating disease of wheat (Triticum aestivum) in China and worldwide, causing severe yield losses annually. Wild emmer (T. dicoccoides) accession IW72 collected from Israel is resistant to powdery mildew at the seedling and adult stages. Genetic analysis indicated that the resistance was controlled by a single dominant gene, temporarily designated MlIW72. The F₂ population and F₃ families derived from a hybrid between IW72 and susceptible durum wheat line Mo75 were used for molecular mapping of the resistance gene. MlIW72 was linked with SSR loci Xgwm344, Xcfa2040, Xcfa2240, Xcfa2257 and Xwmc525 on the long arm of chromosome 7A. In addition, two STS markers, MAG2185 (derived from RFLP marker PSR680) and MAG1759 (developed from EST CD452874), were mapped close to MlIW72. All these markers were physically located in the terminal bin 0.86–1.00 of 7AL. The chromosome location and genetic mapping results suggested that the powdery mildew resistance gene identified in wild emmer accession IW72 might be a new allele at the Pm1 locus or a new locus closely linked to Pm1.     

Molecular mapping of powdery mildew resistance genes in wheat: A review

Powdery mildew, caused by Blumeria graminis f. sp. tritici (syn. Erysiphe graminis f. sp. tritici), is one of the most important diseases of common wheat (Triticum aestivum L.) worldwide. Molecular mapping and cloning of genes for resistance to powdery mildew in hexaploid wheat will facilitate the study of molecular mechanisms underlying resistance to powdery mildew diseases and help understand the structure and function of powdery mildew resistance genes, and permit marker-assisted selection in breeding programs. So far, 48 genes/alleles for resistance to powdery mildew at 32 loci have been identified and located on 16 different chromosomes, of which 21 resistance genes/alleles have been tagged by restriction fragment length polymorphisms (RFLPs), random-amplified polymorphic DNAs (RAPDs), amplified fragment length polymorphisms (AFLPs), sequence characterized amplified regions (SCARs), sequence-tagged sites (STS) or simple sequence repeats (SSRs). Several quantitative trait loci (QTLs) for adult plant resistance (APR) to powdery mildew have been associated with molecular markers. The detailed information on chromosomal location and molecular mapping of these genes has been reviewed. Isolation of powdery mildew resistance genes and development of valid molecular markers for pyramiding resistance genes in breeding programs is also discussed.     

Transfer of powdery mildew resistance from Aegilops variabilis into bread wheat

Winter hexaploid wheat (Triticum aestivum L.) was crossed with Aegilops variabilis to transfer resistance to powdery mildew into wheat. Following two backcrosses to wheat and from 5 to 9 generations of selfing, several disomic addition and substitution lines of hexaploid wheat resistant to the mildew pathogen were isolated. A pair of short satellited chromosomes was always observed in the resistant lines. Further evidence utilizing as markers for homoeologous group 1 HMW glutenin subunits and DNA hybridization with probe pGBX 3076 showed that an alien substitution involved this homoeologous group.     

Resistance to powdery mildew in wild emmer (Triticum dicoccoides Körn.)

Summary Wild emmer from 73 collection sites, including 107 accessions from Israel, two from Lebanon and one from Turkey, were evaluated for resistance to powdery mildew in field nurseries in Israel and the Netherlands.The wild emmer entries displayed a diversity of responses to powdery mildew infection, ranging from high resistance to complete susceptibility. Most entries were resistant in at least one of the nurseries; several entries proved to be resistant in all the tests.Comparing the reactions of 47 wild emmer accessions tested in six nurseries, 11 markedly different patterns were discerned, indicating the probable presence of several different resistance genes.Genes for resistance to powdery mildew appear to be very common in wild emmer indigenous to Israel. Resistance was found in accessions from most collection sites, in all the geographic regions represented in the collection.The common occurrence of resistance and the apparent diversity of genotypes makes wild emmer a rich gene-pool for resistance to powdery mildew. Since genes for resistance to wheat pathogens can be quite readily transferred to cultivated wheat, wild emmer may be utilized as a valuable source of powdery mildew resistance in wheat breeding.     

STS markers for powdery mildew resistance gene Pm6 in wheat

The powdery mildew resistance gene Pm6, transferred to common wheat from the tetraploid Triticum timopheevii, is effective in most epidemic areas for powdery mildew in China. RFLP probe BCD135 was previously associated with Pm6. In the present research, four STS primers (NAU/STS_BCD135-1, NAU/STS_BCD135-2, STS003 and STS004) were designed from the sequence data of BCD135. These primers were used for PCR amplification using the genomic DNA of resistant near-isogenic lines with Pm6 and their recurrent parent, cv. Prins. No polymorphic product was observed using primers STS003 and STS004; however, primers NAU/STS_BCD135-1 and NAU/STS_BCD135-2 amplified two and one bands, respectively, polymorphic between the resistant near-isogenic-lines and Prins. The two primers were then used to amplify the F₂ population from the cross IGV1-465 (FAO163b/7*Prins) × Prins. The amplification and the powdery mildew resistance identification data were analyzed using the software Mapmaker 3.0. The results indicated that both NAU/STS_BCD135-1 and NAU/STS_BCD135-2 were closely linked to Pm6 with a genetic distance of 0.8 cM. A total of 175 commercial varieties without Pm6 from different ecological areas of China were tested using marker NAU/STS_BCD135-2 and none of them amplified the 230 bp-specific band. This marker thus has high practicability and can be used in MAS of Pm6 in wheat breeding programs for powdery mildew resistance. Jianhui Ji and Bi Qin contributed equally to this work.     

Characterization of a wheat–Thinopyron intermedium substitution line with resistance to powdery mildew

Among the progenies of a hybrid between common wheat Triticum aestivum L. cv. Yannong 15 and Thinopyron intermedium, plant E99018 was identified with the chromosome number 2n = 42 and stable agronomic traits. An analysis of the metaphase chromosome pairing indicated that it formed 21 bivalents but that 2 univalents were present in the F₁ hybrid of this plant with common wheat. Resistance verification by race 15 and with mixed races of Blumeria graminis f. sp. tritici at the seedling and adult stages showed that at both stages, the plant was immune to powdery mildew. In situ hybridization with the genomic Th. intermedium and the St genome DNAs as probes and wheat DNA as a block has shown that it contained a pair of Th. intermedium chromosomes. On the basis of the hybridization pattern of the St genome probe to the critical chromosome, a conclusion was reached that this pair of chromosomes belonged to the E genome. Therefore, plant E99018 was a spontaneously formed substitution line. An analysis by 116 SSR markers indicated that the substituted wheat chromosome was 2D and the most likely substitution in E99018 is 2E(2D).     

Postulation of resistance genes to yellow rust in wild emmer wheat derivatives and advanced wheat lines from Nepal

Summary Seedlings of 38 wild emmer derivatives, and a total of 53 advanced wheat varieties/lines introduced from the International Maize and Wheat Improvement Centre (CIMMYT) or other sources, Nepalese breeding lines and local cultivars were inoculated with 18 different yellow rust isolates to postulate yellow rust resistance genes (Yr). Many wild emmer wheat derivatives used were resistant to all isolates indicating the presence of undescribed genes. Some derivatives carried Yr9, Yr6 and/or YrSU. Genes Yr1, Yr2, Yr6, Yr7, Yr8, Yr15, YrSU and YrA+ are no longer effective in Nepal; Yr4, Yr5, Yr9, Yr10, YrSP and YrSD are still effective; the effectiveness of Yr3 remains unclear. This study shows that stripe rust resistance in seedling stage of most Nepalese cultivars and advanced materials is based on Yr9 with combinations of Yr2, Yr6, Yr7, and YrA+, of which only Yr9 is still effective in Nepal. In many countries Yr9 has lost its effectiveness. Therefore the introduction of new Yr-genes from wild emmer wheat in Nepalese cultivars is highly important.     

Microsatellite mapping of powdery mildew resistance allele <Emphasis Type="Italic">Pm5d</Emphasis> from common wheat line IGV1-455

The powdery mildew resistance allele Pm5d in the backcross-derived wheat lines IGV1-455 (CI10904/7*Prins) and IGV1-556 (CI10904/7*Starke) shows a wide spectrum of resistance and virulent pathotypes have not yet been detected in Germany. Although this allele may be distinguished from the other documented Pm5 alleles by employing a differential set of Blumeria graminis tritici isolates, the use of linked molecular markers could enhance selection, especially for gene pyramiding. Pm5d was genetically mapped relative to six microsatellite markers in the distal part of chromosome 7BL using 82 F₃ families of the cross Chinese Spring × IGV1-455. Microsatellite-based deletion line mapping placed Pm5d in the terminal 14% of chromosome 7BL. The closely linked microsatellite markers Xgwm577 and Xwmc581 showed useful variation for distinguishing the different Pm5 alleles except the ones originating from Chinese wheat germplasm. Their use, however, would be limited to particular crosses because they are not functional markers. The occurrence of resistance genes closely linked to the Pm5 locus is discussed. Ghazaleh Nematollahi and Volker Mohler equally contributed to this work.     

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.     

小麦新种质CH1357抗白粉病遗传分析及染色体定位

白粉病是影响小麦产量和品质的一种主要病害。小偃麦衍生品系CH1357对白粉病具有较好的成株抗性,苗期对27个菌株表现为免疫或高抗,是一个高抗白粉病的优异抗源。为了明确其抗白粉病基因在染色体上的位置,对台长29/CH1357和绵阳11/CH1357的F_1、BC_1及F_(2:3)家系进行了遗传分析,并利用分离群体分组分析法(bulked segregantanalysis,BSA)将其初步定位。CH1357的白粉病抗性受1对显性核基因控制,位于染色体5DS,暂命名为PmCH1357。其侧翼连锁标记为Xcfd81和Xbwm8,在2个作图群体台长29/CH1357和绵阳11/CH1357中的遗传距离分别为2.0 cM/11.3 cM和1.5 cM/8.9 cM。PmCH1357与5DS染色体上已报道的其他抗白粉病基因抗谱不同,可能是一个新的抗源。     

A new resistance gene to powdery mildew identified in <Emphasis Type="Italic">Solanum neorossii</Emphasis> has been localized on the short arm of potato chromosome 6

A new resistance (R) gene to powdery mildew has been identified and characterized in a population derived from the wild potato species, Solanum neorossii under natural infection in the greenhouse. The segregation of resistance has revealed that this R gene is controlled by a single monogenic and dominant gene designated Rpm-nrs1. Analysis of the DNA sequence on an internal transcribed spacer (ITS) region of the pathogen genome suggests that the pathogen causing the powdery mildew disease is either Golovinomyces orontii or G. cichoracearum. The resistance locus was localized to the short arm of chromosome 6 where several disease R genes already identified in potato and tomato are known to reside. The resistance locus cosegregated in 96 progeny with three AFLP markers and one PCR marker. The sequences of the two cosegregating AFLP markers are highly homologous to Mi-1 conferring resistance to nematode, potato aphid and whitefly and Rpi-blb2 conferring resistance to late blight. The results in this study will facilitate the cloning of this gene conferring resistance to powdery mildew.     

Characterisation of new oat germplasm for resistance to powdery mildew

Summary Wild oat accessions from the Institute of Plant Genetics and Crop Plant Research (IPK) Gatersleben were selected as potential sources of resistance to powdery mildew (Erysiphe graminis f.sp. avenae). The Avena strigosa accessions AVE 128, AVE 488 and AVE 264, along with A. occidentalis accession CAV 3889 showed high levels of mildew resistance in tests at early and late growth stages over two years; the latter is a particularly useful source because it is a hexaploid species. Moderate levels of resistance were also found in A. fatua accession AVE 1981, AVE 2032 and A.sterilis accession AVE 1373 but this could be due to the later maturity of these genotypes.     

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     

Screening wild Lycopersicon species for resistance to powdery mildew (Oidium lycoperiscum)

Since the late 1980s powdery mildew, designated Oidium lycopersicum, frequently invaded the tomato crop in Western Europe. All commercial cultivars are susceptible. To screen for resistance in wild species a reliable and efficient disease test was developed. Young plants with two to three true leaves are inoculated at high relative humidity by spraying with a freshly prepared suspension of 2×10⁴ conidia, ml^–1. Symptoms are periodically evaluated according to a scale based on the percentage of leaf area with mycelium.One hundred and twenty seven accessions, representing eight wild Lycopersicon species, were screened for resistance to O. lycopersicum. A large variation in resistance was found between species. L. hirsutum was the most resistant species; L. pennellii was moderately resistant; species of the subgeneric group of L. esculentum and of the peruvianum-complex were all susceptible. L. parviflorum was classified separately due to a large variation between accessions. Except for this species, a low variation was found between accessions within species. High levels of resistance were observed in four accessions of L. hirsutum, in one of L. parviflorum and in one of L. peruvianum. This resistance is characterized by a very low disease incidence and a strongly restricted mycelium growth and lack of sporulation.     

Characterisation and origin of rust and powdery mildew resistance genes in VPM1 wheat

Summary The expression of rust resistances conferred by closely linked genes derived from VPM1 varied with environmental conditions and with genetic backgrounds. Under low light and low temperature conditions seedlings carrying Yr17 showed susceptible responses. Stem rust and leaf rust resistance genes Sr38 and Lr37 tended to confer more resistance at 17±2° C than at normal temperatures above > 20° C. These studies supported the hypothesis that Yr17, Lr37 and Sr38 were derived from Aegilops ventricosa, whereas Pm4b was probably derived from T. persicum. Studies on certain addition lines and parental stocks indicated that wheat cytoplasm may enhance the expression of Sr38.     

Isolation of wheat mutants with increased resistance to powdery mildew from small induced variant populations

Small variant wheat populations created by induced mutagenesis (n = 69) or adventitious regeneration (n = 66) were intensively screened for an altered response (compared to the parent variety ‘Guardian’) to the causal pathogen of powdery mildew in wheat, Blumeria graminis f. sp. tritici. Intensive field screening following natural infection of replicated plots of wheat lines over two years revealed a total of 13 mutants exhibiting significantly greater resistance than ‘Guardian’: eight from induced mutagenesis (11.6%) of the M2 population and five from adventitious regeneration (7.6%). Complete resistance was identified in two lines, (one (M66) developed following induced mutagenesis, and the other (SC240) by adventitious regeneration). The complete resistance in the induced mutant was stable over two generations and was associated with a high frequency of leaf flecking, and consequently a low grain yield. Resistance in SC240 proved to be unstable; SC240 exhibited complete resistance to powdery mildew in the SC₂ and SC₃ generations, but only 20% of the SC₄ plants were completely resistant, while the remainder were indistinguishable in mildew response to ‘Guardian’. The mildew response of all the SC₅ generation of SC240 was not significantly different from ‘Guardian’. Yield analysis of the thirteen mutants with increased resistance in the presence of powdery mildew indicated that eleven exhibitedgrain yields at least as high as that of ‘Guardian’, while the mutant M19 exhibited a yield significantly higher than that of ‘Guardian’. This revised version was published online in July 2006 with corrections to the Cover Date.