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.     

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.     

普通小麦DH155抗白粉病基因的分子作图及应用分子标记辅助选择将其转移

普通小麦品系DH155对白粉病菌表现高抗。为明确DH155所携带抗白粉病基因的遗传方式及与抗病基因连锁SSR标记,利用DH155与高感小麦品系SN2890杂交获得的F2和F2:3群体进行接种鉴定和遗传分析,发现DH155对白粉菌菌株E09的抗性受1对显性基因控制,暂命名为Ml DH155。BSA和分子标记分析结果显示,Ml DH155与SSR标记Xcfd81和Xcfd18连锁。利用已发表的中国春和粗山羊草D基因组序列开发新标记,进一步将Ml DH155定位于标记Xsdau K525和Xsdau K527之间,其遗传距离分别为0.2 c M和0.8 c M。将DH155与感白粉病优良品系HB133-4和旱10杂交,在F2~F4代,结合优良农艺性状选择、分子标记辅助选择和抗白粉病鉴定,获得3个高抗白粉病且农艺性状优异的株系(SDAU2100、SDAU2101和SDAU2102)。利用14个白粉菌菌株对DH155进行苗期接种鉴定表明,DH155对13个菌株表现抗病反应型。这些菌株对DH155的毒力谱与已知抗白粉病基因Pm2相似,但DH155对Bg78-3和Bg44-5菌株的反应型与携带Pm2的Ulka/8*Cc不同。结合本试验结果和Pm2基因的相关报道,推测Ml DH155可能是Pm2或其等位基因。     

Genetic analysis of powdery-mildew resistance of a winter-wheat line, RE714, and identification of a new specific-resistance gene

The winter-wheat line RE714, resulting from an intergeneric cross between Aegilops squarrosa and Triticum dicoccum, and implying a sister line of ‘Roazon’ (VM4) in its genealogy, was obtained in order to increase the genetic diversity for disease resistance. The aim of this study was to identify and characterize the powdery-mildew-resistance gene(s) present in this line at the seedling stage on detached leaves. The comparison of the reaction of the RE714 line and of differential hosts to 24 Erysiphe graminis. f.sp. tritici isolates indicated that only Pm4a, Pm4b and/or Mlar genes could be present and that a new resistance factor, different from all known mildew-resistance genes, was present in this line. The powdery-mildew reactions of segregating F₂ populations produced evidence that RE714 carries the Pm4b allelic form and a recessive new resistance gene, tentatively designated mlre. While Ae. squarrosa was resistant to most isolates, T. dicoccum was specifically attacked by REV 14-virulent isolates and was therefore assumed to be the donor of mire. The VM4 parent was confirmed as the Pm4b donor.     

Mapping QTL involved in powdery mildew resistance of the apple clone U 211

Scab and powdery mildew, caused by Venturia inaequalis (Cke.) Wint. and Podosphaera leucotricha (Ellis et Ev.) Salm. are the most important apple diseases. The apple clone U 211 is resistant to scab and is also highly resistant to powdery mildew under field conditions. The interval mapping method was applied for the identification of genomic regions conferring U 211 resistance to powdery mildew. The genetic maps of the ‘Idared’ and U 211 genome sectors were constructed using amplified fragment lenght polymorphism and simple sequence repeat markers and 98 individuals from the progeny of the cross ‘Idared’× U 211. On the basis of the phenotypic and molecular marker data 10 powdery mildew resistance quantitative trait loci (QTL) were identified in U 211 and ‘Idared’. One of the QTL in the clone U 211 explained 48‐72% of the phenotypic variation and its effect was stable over years.     

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.     

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.     

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.     

Genes for Powdery Mildew Resistance in Cultivars of Spring Wheat

Twenty-three cultivars of spring wheat were inoculated with nineteen different powdery mildew isolates; their ruction patterns hive been compared with those of twenty-two cultivars/lines carrying identified powdery mildew resistance genes. Applying the gene-for-gene hypothesis, it is evident that three cultivars have none of the resistance genes used, seven others (including ‘Solo’) may carry Pm4b, only. The resistance pattern of ‘Selpek’ is identical to A/-1 resistant cultivars of winter wheat and may be explained by the presence of Pm5. The resistance pattern of Pm5 (Mt-i) cultivars is very different from a number of ‘Kolibri’-related cultivars of spring wheat. Since either all or nothing of that specific pattern has been transferred to all cross progenies of ‘Kolibri’, a single gene is assumed to oe responsible for it, preliminarily designated as Ml-k. The cultivar ‘Mephisto’ carries the ‘Normandie’ resistance (Pwl 2, 9). In five cultivars to spring wheat the combined effects of at least two of the above-mentioned sources have been found. Despite the fact that ‘Normandie’ and ‘Sappo’ are not closely related. ‘Sappo’ shows the complete ‘Normandie’ resistance pattern plus that of Pm4b. The same is true for ‘Planet’ and ‘Walter’.     

QTL mapping of adult‐plant resistance to stripe rust in a ‘Lumai 21 × Jingshuang 16’ wheat population

Stripe rust, caused by Puccinia striiformis f. sp. tritici, is a devastating fungal disease in common wheat (Triticum aestivum L.) worldwide. Chinese wheat cultivars ‘Lumai 21’ and ‘Jingshuang 16’ show moderate levels of adult‐plant resistance (APR) to stripe rust in the field, and they showed a mean maximum disease severity (MDS) ranging from 24 to 56.7% and 26 to 59%, respectively, across different environments. The aim of this study was to identify quantitative trait loci (QTL) for resistance to stripe rust in an F₃ population of 199 lines derived from ‘Lumai 21’ × ‘Jingshuang 16’. The F₃ lines were evaluated for MDS in Qingshui, Gansu province, and Chengdu, Sichuan province, in the 2009–2010 and 2010–2011 cropping seasons. Five QTL for APR were detected on chromosomes 2B (2 QTL), 2DS, 4DL and 5DS based on mean MDS in each environment and averaged values from all three environments. These QTL were designated QYr.caas‐2BS.2, QYr.caas‐2BL.2, QYr.caas‐2DS.2, QYr.caas‐4DL.2 and QYr.caas‐5DS, respectively. QYr.caas‐2DS.2 and QYr.caas‐5DS were detected in all three environments, explaining 2.3–18.2% and 5.1–18.0% of the phenotypic variance, respectively. In addition, QYr.caas‐2BS.2 and QYr.caas‐2BL.2 colocated with QTL for powdery mildew resistance reported in a previous study. These APR genes and their linked molecular markers are potentially useful for improving stripe rust and powdery mildew resistances in wheat breeding.     

Resistance to powdery mildew in selections from Tunisian landraces of barley

Two-hundred and thirty-two accessions of barley landraces collected from Tunisia were screened for resistance to powdery mildew. A number of race-specific genes were detected using the detached leaf technique. Among the 232 accessions tested, 169 were susceptible to powdery mildew, 20 were resistant and 43 showed differential reactions to the three isolates of powdery mildew used. An attempt was made to determine the number of genes, the types of gene, the types of gene action and the gene loci in 20 resistant accessions. Three types of cross were made: (1) the accessions were crossed to the susceptible variety ‘Pallas’, (2) the accessions were crossed with ‘Pallas’ isolines, and (2) accessions with identical powdery mildew reaction patterns were intercrossed. Three isolates of Erysiphe graminis f. sp. hordei were used: Bzm-1, KM 18-75, R13C. A number of different resistance genes were detected among the 19 resistant accessions. Surprisingly, segregation indicating single genes only were detected with the isolates used. Some of these genes could be associated with loci already known. In 19 cases a dominant and in one a recessive mode of inheritance was detected. The recessive gene was not located at the mlo locus. This investigation represents the first systematic study of race-specific genes for powdery mildew resistance in Tunisian landraces. The newly identified sources of resistance may be used in many strategies of breeding for disease resistance.     

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

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

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.     

普通小麦白粉病成株抗性的QTL分析

以抗白粉病的日本小麦品种Fukuho-komugi和以色列小麦Oligoculm杂交F₁的DH（doubled haploid）群体107个系为材料，利用313个SSR标记和37个RFLP标记，对Fukuho-komugi和Oligoculm的白粉病成株抗性进行QTL分析。试验材料于2003－2004年度种植在北京、2003－2004和2004－2005年度种植在安阳，调查白粉病发病情况。构建了由350个位点组成的遗传连锁图，覆盖小麦21个连锁群，全长3 101 cM。采用复合区间作图法进行白粉病成株抗性QTL分析，在1A、2B、4B和7D上发现4个抗白粉病QTL，分别解释13.6%、6.6%、8.9%和12.7%的表型变异。抗白粉病基因及其紧密连锁分子标记的发掘，将为小麦抗白粉病育种的分子标记辅助选择提供理论和技术支持。     

Inheritance of the Powdery Mildew Resistance Mlk in Wheat

The inheritance of the Mlk powdery mildew resistance originating from ‘Heine 2174.50’ was analyzed by crossing the Mlk resistant cultivar ‘Ralle’× cv. ‘Amor’ (highly susceptible) and vice versa and by observing the reactions of F₁- and F₂-plants after inoculation with two different Mlk avirulent powdery mildew isolates. In all cases, a 3 (resistant): I (susceptible) segregation was found in F₂. The reactions of the F₂plants against the two powdery mildew isolates were identical in each case. Therefore, it is supposed that one dominant resistant gene is responsible for the resistant reactions against these two isolates. These results support the earlier assumption of Heun and Fischbeck (1987b) that the whole Mlk resistance pattern is controlled by a single gene.     

1BL/1RS易位对小麦产量性状和白粉病抗性的影响及其QTL分析

用PH82-2/内乡188杂交后代240个F_5:6家系,按照α-lattice设计,分别种植在安阳、焦作和泰安,对产量和抗白粉病等性状进行了考察。利用SSR和蛋白标记对群体进行部分连锁作图,分析1BL/1RS易位对产量及其相关性状的遗传效应。结果表明,1BL/1RS易位系对产量、穗数/m²和抽穗期的影响不显著;易位系的千     

小麦白粉病成株抗性和抗倒伏性及穗下节长度的QTL定位

由小麦品种花培3号和豫麦57杂交获得了DH群体168个株系, 利用305个SSR标记对白粉病成株抗性、抗倒伏性和穗下节长度进行了QTL定位研究。DH群体及两亲本于2005年和2006年种植于山东泰安, 2006年种于安徽宿州。利用基于混合线性模型的QTLNetwork 2.0软件, 共检测到12个加性效应位点和10对上位效应位点。在4D染色体上控制白粉病成株抗性的qApr4D, 贡献率为20.0%, 在各环境中稳定表达, 其抗病等位基因来源于抗病亲本豫麦57; 在7D染色体上控制小麦穗下节长度的qIlbs7D, 贡献率为12.9%, 在各环境中稳定表达。加性效应和上位效应对小麦白粉病成株抗性、抗倒伏性和穗下节长度的遗传起重要作用, 并且基因与环境常常具有互作效应。以上两个QTL可分别用于小麦白粉病成株抗性和穗下节长度的分子标记辅助选择。     

Identification of a novel recessive gene for resistance to powdery mildew (Blumeria graminis f. sp. hordei) in barley (Hordeum vulgare)

One of the most important diseases of barley (Hordeum vulgare) is powdery mildew, caused by Blumeria graminis f. sp. hordei. Spring barley line 173-1-2 was selected from a Moroccan landrace and revealed broad-spectrum resistance to powdery mildew. The objective of this study was to map and characterize the gene for seedling powdery mildew resistance in this line. After crossing with the susceptible cultivar ‘Manchuria’, genetic analysis of F₂ and F₃ families at the seedling stage revealed powdery mildew resistance in line 173-1-2 conditioned by a single recessive gene. Molecular analysis of non-segregating homozygous resistant and homozygous susceptible F₂ plants conducted on the DArTseq platform (Diversity Arrays Technology Pty Ltd) identified significant markers which were converted to allele-specific PCR markers and tested among 94 F₂ individuals. The new resistance gene was mapped on the long arm of chromosome 6H. No other powdery mildew recessive resistance gene has been located on 6H so far. Therefore, we concluded that the 173-1-2 barley line carries a novel recessive resistance gene designated as mlmr.     

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.     

Mapping a new source of resistance to powdery mildew in mungbean

Both restriction fragment length polymorphism (RFLP) and amplified fragment length polymorphism (AFLP) analyses were employed to map a new source of resistance to powdery mildew in mungbean. Disease scores of an F₂ population derived from the cross between a moderately resistant breeding line VC1210A and a susceptible wild relative (Vigna radiata var. sublobata, accession TC1966) showed a continuous distribution and was treated as a quantitative trait. Although no significant quantitative trait loci (QTL) that can explain the variation was detected by QTL analysis based on the reconstructed RFLP linkage map, new marker loci associated with resistance were discovered by AFLP analysis. The RFLP loci detected by two of the cloned AFLP bands are associated with resistance and constitute a new linkage group. A major resistance quantitative trait locus was found on this linkage group that accounted for 64.9% of the variation in resistance to powdery mildew. One of the probes developed in this study has the potential to assist in breeding for powdery mildew resistance in mungbean.