Inheritance of the Powdery Mildew Resistance Gene Pm9 in Relation to Pm1 and Pm2 of Wheat

The inheritance of the powdery mildew resistance gene Pm9 originating from the hexaploid spring wheat cultivar ‘Normandie’ was analyzed in relation to Pm1 and Pm2. Two leaf segments of individual P_1?, P_2?, F_1? and F₂-plants of the cross ‘Normandie’ (Pm1, 2, 9) בFederation’ (no known Pm gene) were inoculated separately with two powdery mildew isolates. Using powdery mildew isolate No. 6 virulent for Pm1 and Pm2 but avirulent for Pm9, a 1 resistant (r): 3 susceptible (s) F₂-segregation was found for the Pm9 gene. Using powdery mildew isolate No. 3 virulent for Pm1 and Pm9 but avirulent for Pm2, a 3 (r): 1 (s) F₂-segregation was found for the Pm2 gene. Combining the data of both experiments (leaf segments of identical plants had been used), a 9 (sr): 3 (ss): 3 (rr): 1 (rs) segregation resulted for the F₂ of this cross: therefore, independent inheritance of the genes Pm2 and Pm9 can be concluded. Similarly, the cross ‘Mephisto’ (Pm1, 2, 9) בAmor’ (no known Pm gene) was analyzed. The Pm9 gene again showed a monogenically recessive inheritance, whereas Pm1 showed a monogenically intermediate segregation upon inoculation with powdery mildew isolate No. 9a virulent for Pm2 and Pm9 but avirulent for Pm1. Combining the single gene segregations, linkage between both genes was found among the progenies. A distance of 8.5 cM was calculated. Analyzing a set of spring wheat cultivars with seven defined powdery mildew isolates, the presence of Pm1, Pm2 and Pm9 in these lines was verified; in most cases, Pm1 occurred together with Pm9.     

Race Specific Powdery Mildew Resistance in 31 Northwest European Wheat Cultivars

Race specific powdery mildew resistance in 23 winter wheat cultivars, eight spring wheat cultivars, and 14 lines/cultivars possessing known powdery mildew resistance genes, has been studied by analyzing host/pathogen interactions. The cultivars were tested as intact seedlings, and as detached primary leaf segments on water agar; both methods revealed reproducible and concordant results. The 45 cultivars/lines were divided into 24 resistance spectra according to the patterns of reaction to the powdery mildew isolates used. Of the 31 cultivars investigated, eight did not possess any of the resistance genes detected, and the remaining 23 were divided in 16 resistance spectra. The race specific resistance of nine cultivars was conferred by the single resistance genes Pm2, Pm4b, Pm5/Ml-i: or Pm6, while the race specific resistance of 14 cultivars was conferred by 2, 3, 4 or 5 genes in combination.     

Identification of Wheat Powdery Mildew Resistance Genes by Analysing Host-Pathogen Interactions

Fifty-nine winter wheat cultivars and thirteen lines possessing known powdery mildew resistance genes were inoculated with eleven different isolates. By comparing their resistance patterns the responsible major resistance genes of the above-mentioned cultivars have been determined. The so-called “Blaukorn” resistance is conditioned by gent Pm4b. The resistance patterns of Ml-i and Pm5 being similar, the relationship between them has to be analysed by segregating populations.     

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.     

Chromosomal location of powdery mildew resistance gene Pm16 in wheat using SSR marker analysis

The use of resistant cultivars is a most economical way to control powdery mildew (Blumeria graminis f.sp. tritici) in wheat (Triticum aestivum L.). Identification of molecular markers closely linked to resistance genes can greatly increase the efficiency of pyramiding resistance genes in wheat cultivars. The objective of this study was to identify molecular markers closely linked lo the powdery mildew resistance gene Pm16. An F₂ population with 156 progeny was produced from the cross‘Chancellor’(susceptible) ב70281’ (resistant), A total of 45 SSR markers on chromosomes 4A and 5B of wheat and 15 SSRs on chromosome 3 of rice was used lo lest the parents, as well as the resistant and susceptible bulks: the resulting polymorphic markers were used to genotype the F₂ progeny. Results indicated that the SSR marker Xgwm159, located on the short arm of chromosome 5B, is closely linked to Pm16 (genetic distance: 5.3 CM). The cytogenetical data presented in an original report, in combination with this molecular analysis, suggests that Pm16 may he located on a translocated 4A.5BS chromosome.     

An STS marker distinguishing the rye-derived powdery mildew resistance alleles at the Pm8/Pm17 locus of common wheat

A sequence‐tagged site marker has been developed from restriction fragment length polymorphism marker probe IAG95 for the rye‐derived powdery mildew resistance Pm8/Pm17 locus of common wheat. This polymerase chain reaction marker enables the amplification of DNA fragments with different sizes from T1AL.1RS and T1BL.1RS wheat‐rye translocation cultivars with chromatin from ‘Insave’ and ‘Petkus’ rye, respectively, and therefore will be very useful in distinguishing Pm8‐carrying cultivars from Pm17‐carrying cultivars. Results obtained with that marker were compared with resistance tests performed on detached primary leaves of 29 wheat lines from two populations derived from doubled haploid production. The molecular assay corresponded well with the resistance tests in all the lines, and therefore will be helpful for the identification of Pm17 in lines in which other Pm genes or quantitative trait loci are present.     

Identification of powdery mildew resistance genes in common wheat (Triticum aestivum L. em Thell.). IX. Cultivars, land races and breeding lines grown in China

The objective of the study was to provide information about the occurrence and distribution of resistance genes in wheat cultivars, including old cultivars, land races and advanced breeding lines grown in China. Ninety-four accessions were analysed with a set of 11 differential powdery mildew isolates. Forty-four cultivars did not possess any major mildew resistance genes. Thirty cultivars revealed the response pattern of individual resistance genes. The most frequently encountered gene was Pm8, which occurred singly in 11 cultivars, combined either with Pm4a in three cultivars or with Pm4b in another three cultivars. However, 12 cultivars possessing the wheat-rye translocated chromosome pair T1BL-1RS did not express Pm8. Gene Pm2 was found in four cultivars and in combination with Pm6 in one cultivar. Genes Pm4a and Pm4b were observed in four and five cultivars, respectively. Another six cultivars carried Pm5. A gene combination of Pm2+Pm4b+Pm6 was found in one cultivar. Twelve cultivars and breeding lines exhibited a response pattern that could not be assigned to resistance genes or gene combinations present in the differential cultivars. Five out of these 12 cultivars/lines showed resistance to all the isolates tested. There is an urgent need to search for novel sources of mildew resistance in order to sustain resistance to existing and emerging powdery mildew pathogens.     

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.     

Powdery mildew resistance in Czech and Slovak barley cultivars

Fifteen powdery mildew resistance genes and the gene MlaN81 derived from ‘Nepal 81’were found in 76 Czech and Slovak spring and winter barley cultivars when tested for reaction to a set of powdery mildew isolates. Nine cultivars (‘Donum’, ‘Expres’, ‘Jubilant’, ‘Orbit’, ‘Primus’, ‘Progres’, ‘Stabil’, ‘Vladan’ and ‘Zlatan’) are composed of lines with different resistance genes. The Mlat gene is present in nine cultivars and was transferred from the Anatolian landrace ‘A‐516′. The resistances derived from ‘KM‐1192’and ‘CI 7672’were identical and designated Ml(Kr). Five winter barley cultivars possess the Ml(Bw) resistance. The winter barley line ‘KM‐2099’carries the mlo gene. The parental cultivar ‘Palestine 10’was also tested in which the genes Mlk1, MlLa were identified. The German cultivar ‘Salome’, a parent of seven cultivars tested, probably carries the gene MlLa in addition to mlo and Mla7. The gene mlo6 may be present in the cultivar ‘Heris’. Most of the results were confirmed by the pedigrees of the cultivars.     

Dynamics of changes in the races and virulence of wheat powdery mildew in Hungary between 1971 and 1999

Important microevolutional processes have taken place in the wheatpowdery mildew population over the last thirty years. There has been aconsiderable change in the race composition of the pathogen populationand in the prevalent races. Of the 78 races identified, only eleven have`lived' for more than 15 years. Many races were only isolated in one ortwo years. The number of virulence genes rose from 2.03 in 1973 to 5.63in 1993. On the basis of race composition and virulence the wheatpowdery mildew population between 1971 and 1999 can be divided intofour distinct groups. A large proportion of the powdery mildew isolates arevirulent to most resistance genes. Complete resistance is provided byresistance genes Pm4a (Khapli) and partial resistance by Pm2 + Mld (Halle st. 13471), Pm4b+ (TP 315/2) and Pm1 + 2 + 9 (Normandie). The majority of cultivated varieties carry theresistance gene Pm8 due to the presence of the 1B/1R translocation.     

Effectiveness and environmental stability of quantitative powdery mildew (Blumeria graminis) resistance among winter wheat cultivars

Powdery mildew in wheat (Blumeria graminis f. sp. tritici) is a major disease in Northern and Central Europe. The aim of the study was to analyse the effectiveness and environmental stability of quantitative powdery mildew resistance under high epidemic pressure in the field across years in the absence/presence of ineffective race‐specific resistances. Cultivars with and without Pm (major) genes were inoculated in three experiments with a genetically broad mildew population with all matching virulences. Resistance was measured three times by assessing the percentage of leaf area covered by powdery mildew on a plot basis (0–100%). Mean powdery mildew severity of the highly susceptible cv. ‘Kanzler’ varied across 10 years from 24% to 66% (Exp. 1). Means of three cultivars without Pm genes, ‘Ramiro’, ‘Miras’ and ‘Zentos’, and several cultivars with ineffective Pm genes varied quantitatively from 4% to 13%. Environmental stability of the quantitative resistances was on average higher than that of susceptible genotypes, as revealed by a regression approach. In the second experiment, all groups of cultivars with ineffective Pm gene(s) contained a large proportion of entries with a similar low mildew rating as the quantitatively resistant standard ‘Miras’. Mildew severity of pairs of cultivars with the same Pm gene(s) was significantly different across 6 years (Exp. 3) indicating the presence of additional quantitative resistances in some of these cultivars. In the analysis of variance, genotypic variance had a high impact (P < 0.01) with low importance of genotype × environment interaction. Consequently, heritabilites were high (0.95–0.97). In conclusion, breeders have already accumulated effective minor genes for powdery mildew resistance in many of the released German winter wheat cultivars. These quantitative resistances are long lasting, environmentally stable and provide a high level of protection to powdery mildew.     

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.     

Temperature and resistance gene interactions in the expression of resistance to Blumeria Graminis f. sp. Tritici

The influence of temperature on the resistance to powdery mildew, caused by Blumeria graminis (DC.) E.O. Speer f. sp. tritici Em. Marchal, was evaluated on six wheat cultivars/lines, Axminster/8^* Chancellor (Cc) (Pm1), Chul/8^*Cc (Pm3b), Yuma/8^*Cc (Pm4a), VPM1 (Pm4b), Kavkaz (Pm8), and Chancellor, at two temperatures of 25EC and 15EC with two isolates of powdery mildew. Resistance, based on the components of latency period, infection type, colony number per unit leaf area, per cent infected leaf area, and AUDPC, was much less effective at 15EC than at 25EC for VPM1 and Yuma/8^*Cc, but no resistance shift was observed for Kavkaz and Axminster/8^*Cc, indicating the difference of temperature-sensitivity among Pm genes or among the interactions of individual host genes with corresponding powdery mildew pathogen genes. This revised version was published online in July 2006 with corrections to the Cover Date.     

12个小麦品种(系)白粉病抗性的遗传分析

利用17个不同来源和毒力的白粉菌菌株对12个小麦品种(系)进行苗期抗性鉴定和抗病性遗传分析,同时利用Pm2和Pm8基因的特异分子标记检测了相应基因。供试的12个品种至少能够抗11个白粉菌菌株。用E09、E20和Bg2菌株接种F₂群体,抗感植株分离比例和适合性测验证明这12个品种对不同白粉菌菌株的抗性均受1对显性基因控制。抗谱分析和基因紧密连锁分子标记(Xcfd81)分析表明良星66很可能含有Pm2或其等位基因。ω-黑麦碱基因(1RS染色体)和Glu-B1基因(1BS染色体)特异分子标记分析结果证明,山农20和郑麦9962含有T1BL·1RS易位染色体,即可能携带Pm8基因。由于Pm8基因对大多数菌株表现感病,所以这2个品种除Pm8外,还具有其他抗病基因。偃展4110与天民668对参试菌株的反应型表现一致,其他材料对不同菌株的反应型表现不同。     

Identification of a microsatellite marker associated with Pm3 resistance alleles to powdery mildew in wheat

The Pm3 resistance locus, located on chromosome 1A in wheat, confers race‐specific resistance to the obligate biotrophic fungus Blumeria graminis (DC) E.O. Speer f. sp. tritici, the causal agent of powdery mildew. Several Pm3 alleles are still effective in controlling the disease in Europe. A genetic map was constructed to map the Pm3g allele in the recombinant inbred line progeny from the cross ‘RE9001’ (susceptible) בCourtot’ (resistant). Two microsatellite markers were closely mapped to Pm3g. The PSP2999 marker, which cosegregates with this allele, was shown to detect the presence of the Pm3g resistance allele in other cultivars. A collection of 56 wheat cultivars or advanced lines carrying one Pm3 allele was used to assess the allele‐specific amplification of the PSP2999 marker. The same amplification pattern was obtained for lines with Pm3a, Pm3b, Pm3e, Pm3f and Pm3g alleles. Twenty genotypes carrying Pm3d showed a specific amplification pattern. This marker allowed the detection of the Pm3d allele in highly resistant lines whose resistance gene combinations were unknown. It was concluded that PSP2999 is a useful marker to detect Pm3 alleles in parents and to manage them in breeding programmes.     

兼抗型成株抗性小麦品系的培育、鉴定与分子检测

小麦条锈病、叶锈病和白粉病是我国小麦的重要真菌病害,培育兼抗型成株抗性品种是控制病害最为经济有效和持久安全的方法。本研究选用由成株抗性育种方法培育的21份冬小麦高代品系和96份春小麦高代品系,在多个环境下进行这3种病害的成株期抗性鉴定,并利用紧密连锁的分子标记检测了兼抗型基因Lr34/Yr18/Pm38、Lr46/Yr29/Pm39和Sr2/Yr30的分布。田间鉴定表明,21份冬小麦品系中有17份兼抗3种病害,占80.9%;96份春小麦品系中有85份兼抗3种病害,占88.5%。分子标记检测发现,21份冬小麦品系均含QPm.caas-4DL,其中7份还含QPm.caas-2BS,9份还含QPm.caas-2BL;96份春小麦品系中,18份含Lr34/Yr18/Pm38,37份含Lr46/Yr29/Pm39,29份含Sr2/Yr30。以上结果表明,分子标记与常规育种相结合,可有效培育兼抗型成株抗性品种,为我国小麦抗病育种提供了新思路。     

A major QTL effect controlling resistance to powdery mildew in winter wheat at the adult plant stage

Powdery mildew is one of the major diseases of wheat in regions with a maritime or semi‐continental climate which can strongly affect grain yield. The objective of the study was to identify and compare quantitative resistance to powdery mildew of line RE9001 at the adult plant and vernalized seedling stages. RE9001 has no known Pm gene and shows a high level of adult plant resistance in the field. Using 104 recombinant inbred lines (RILs) of an RE9001 × ‘Courtot’ F8 population, a genetic map was developed with 363 markers distributed over 26 linkage groups and covering 3825 cM. The global map density was 1 locus/10.3 cM. RILs were assessed under field and tunnel greenhouse conditions for 2 years in two locations. Eleven quantitative trait loci (QTL) were detected at the adult stage and they explained 63% of the variation, depending on the environment. Three QTLs were found, at least, in the two environments. One QTL from RE9001, mapped on chromosome 2B, was stable in each environment. This QTL, QPm.inra.2B, explained 10.3–36.6% of the variation and could be mapped in the vicinity of the Pm6 gene. At the vernalized seedling stage, one QTL detected by the isolate 93‐27 could be an allele of the Pm3g gene present in ‘Courtot’. No residual effect of the Pm3g gene was detected at either stage. Markers flanking the QTL 2B could be useful tools to combine resistance to powdery mildew in wheat cultivars.     

Mapping QTL involved in adult plant resistance to powdery mildew in the winter wheat line RE714 in two susceptible genetic backgrounds

The objective was to study the genetic basis of adult plant resistance to powdery mildew of the winter wheat line RE714 by quantitative trait loci (QTL) analysis and to investigate the stability of the QTL detected in two different genetic backgrounds. Two DH populations from the crosses between RE714 and the susceptible parents ‘Festin’ and ‘Hardi’ were used. Reaction of the DH lines to powdery mildew was assessed in different environments in Belgium under natural disease infection. Considering both populations and according to the environment tested, one to seven QTL were detected. Among them, residual effects of the race‐specific resistance genes Pm4b and MIRE were found. Two major QTL were very stable (on chromosome 5D and at the MIRE locus), since they were detected in both populations and over all environments tested. The QTL detected varied according to the susceptible parent used, and a residual effect at the Pm4b gene was not observed with the genetic background of ‘Hardi’.     

Race-Specific Resistance Genes Against Erysiphe graminis f. sp. hordei in Old and Recent French Barley Accessions

Leaf segments from seedlings of 68 old barley varieties and 38 more recent cultivars of past or current importance in France were infected with 6—9 powdery mildew isolates in order to identify race-specific resistance genes. No resistance gene was apparent in the 21 old winter varieties, 43 of 47 old spring varieties, 10 recent winter and 3 recent spring cultivars. Two old spring varieties (‘Colmar S 142’ and ‘Johanna’) were postulated to have Mlg and Ml(CP), and two others (‘Pontrieux’ and ‘Finistère 62-5’) had an unidentified, weakly effective gene. Mlg, Ml(CP), Mlb, Mla6 and Mlal3 were the only genes detected in recent winter cultivars. Recent spring cultivars presented the greatest diversity; the presence of one or several genes among Mlal, Mla6, Mla9, Mla12, Mlg, Ml(CP), Ml(La), Mlk and mlo was postulated in several lines.     

Phenotypic expression of quantitative and qualitative components of partial resistance to powdery mildew (Sphaerotheca fuliginea race 1) in melon (Cucumis melo) germplasm

A study was conducted to investigate the expression of four components of partial resistance to Sphaerotheca fuliginea race 1 in selected melon (Cucumis melo L.) lines viz. infection frequency, latent period, spore production, and disease-severity score. Those components were evaluated at two developmental stages of the host: the cotyledon stage and the stage of the first two true leaves. Detached plant parts (disks of cotyledons and true leaves) were inoculated using a vacuum-operated settling tower. All four components showed significant variation among genotypes, and correlations between components at both developmental stages were large and significant. The line ‘CNPH 83–095’ (without any major resistance gene to powdery mildew) presented the highest level of partial resistance in both vegetative stages for almost all components evaluated. The lines ‘W-6’ (Pm1Pm1, Pm2Pm2), ‘Cinco’ (Pm1Pm1, Pm2Pm2), and CNPH ‘84–147’ (Pm1Pm1), even though carrying the major gene Pm1 for complete resistance to race 1 of the fungus, showed slight but significant differences for quantitative components of partial resistance at the cotyledonal stage. Different levels of partial resistance may be expressed, even in lines with a major race-specific resistance gene to powdery mildew, in specific developmental stages of the melon plants.