Identification of QTLs for partial resistance to leaf rust (Puccinia hordei) in barley

 The partial resistance to leaf rust in barley is a quantitative resistance that is not based on hypersensitivity. To map the quantitative trait loci (QTLs) for partial resistance to leaf rust, we obtained 103 recombinant inbred lines (RILs) by single-seed descent from a cross between the susceptible parent L94 and the partially resistant parent Vada. These RILs were evaluated at the seedling and adult plant stages in the greenhouse for the latent period (LP) of the rust fungus, and in the field for the level of infection, measured as area under the disease progress curve (AUDPC). A dense genetic map based on 561 AFLP markers had been generated previously for this set of RILs. QTLs for partial resistance to leaf rust were mapped using the “Multiple Interval Mapping” method with the putative QTL markers as cofactors. Six QTLs for partial resistance were identified in this population. Three QTLs, Rphq1, Rphq2 and Rphq3, were effective at the seedling stage and contributed approximately 55% to the phenotypic variance. Five QTLs, Rph2, Rphq3, Rphq4, Rphq5, and/or Rphq6 contributed approximtely. 60% of the phenotypic variance and were effective at the adult plant stage. Therefore, only the QTLs Rphq2 and Rhpq3 were not plant-stage dependent. The identified QTLs showed mainly additive effects and only one significant interaction was detected, i.e. between Rphq1 and Rphq2. The map positions of these QTLs did not coincide with those of the race-specific resistance genes, suggesting that genes for partial resistance and genes for hypersensitive resistance represent entirely different gene families. Also, three QTLs for days to heading, of which two were also involved in plant height, were identified in the present recombinant inbred population. These QTLs had been mapped previously on the same positions in different populations. The perspectives of these results for breeding for durable resistance to leaf rust are discussed. Received: 15 July 1997 / Accepted: 30 December 1997     

The evidence for abundance of QTLs for partial resistance to Puccinia hordei on the barley genome

Using AFLP markers, a linkage map was constructed based on a recombinant inbred population of barley derived from a cross between a leaf rust susceptible line, L94, and a partially resistant line, 116-5. The constructed map showed a similar marker distribution pattern as the L94 × Vada map. However, it contained more large gaps, and for some chromosome regions no markers were identified. These regions are most likely derived from L94 because 116-5 was selected from the progeny of a cross of L94 × cv. Cebada Capa. Five QTLs for partial resistance to isolate 1.2.1. were mapped on the L94 × 116-5 map. Three QTLs were effective in the seedling stage, jointly contributing 42% to the total phenotypic variance. Three QTLs were effective in the adult plant stage, collectively explaining 35% of the phenotypic variance. Evidence for two additional linked minor-effect QTLs effective in the adult plant stage was also uncovered. The major-effect QTL, Rphq3, was the only one that was effective in both developmental stages. Moreover, Rphq3, was also identified in the L94 × Vada population, being effective to two rust isolates. The other QTLs were detected in either of the two populations, providing evidence for the existence of many loci for partial resistance to leaf rust on the barley genome. To date, 13 QTLs for partial resistance have been mapped, therefore, a strategy of accumulating many resistance genes in a single cultivar, resulting in a high level of partial resistance, is feasible.     

Localising QTLs for leaf rust resistance and agronomic traits in barley (Hordeum vulgare L.)

The Hordeum vulgare accession ’HOR 1063’ was crossed with the barley cultivar Krona, and 220 doubled haploid lines were produced based on this cross. A molecular map was constructed based on RFLP markers. Field trials were performed over 2 years and at two locations. In field trials, resistance to leaf rust by means of artificial infection, heading date, plant height and Kernel weight were assessed. For leaf rust resistance, 4 QTLs were localised, that explained 96.1% of the genetic variation. One QTL on chromosome 4H confirmed a position found in another genetic background and one mapped to the same position as Rph16 on chromosome 2H. All digenic effects decreased the effects of the respective QTLs. In addition to the denso-locus and the hex-v locus, other QTLs influencing heading date, plant length and kernel weight were found in this cross. Received: 16 July 1999 / Accepted: 7 September 1999     

Introgression of quantitative trait loci (QTLs) determining stripe rust resistance in barley: an example of marker-assisted line development

 Genome-analysis tools are useful for dissecting complex phenotypes and manipulating determinants of these phenotypes in breeding programs. Quantitative trait locus (QTL)-analysis tools were used to map QTLs conferring adult plant resistance to stripe rust (caused by Puccinia striiformis f.sp. hordei) in barley. The resistance QTLs were introgressed into a genetic background unrelated to the mapping population with one cycle of marker-assisted backcrossing. Doubled-haploid lines were derived from selected backcross lines, phenotyped for stripe-rust resistance, and genotyped with an array of molecular markers. The resistance QTLs that were introgressed were significant determinants of resistance in the new genetic background. Additional resistance QTLs were also detected. The susceptible parent contributed resistance alleles at two of these new QTLs. We hypothesize that favorable alleles were fixed at these new QTLs in the original mapping population. Genetic background may, therefore, have an important role in QTL-transfer experiments. A breeding system is described that integrates single-copy and multiplex markers with confirmation of the target phenotype in doubled-haploid lines phenotyped in field tests. This approach may be useful for simultaneously producing agronomically useful germplasm and contributing to an understanding of quantitatively inherited traits. Received: 6 May 1997 / Accepted: 1 September 1997     

Potential of doubled-haploid lines and localization of quantitative trait loci (QTL) for partial resistance to bacterial leaf streak (Xanthomonas campestris pv. hordei) in barley

 Genetic variability for partial resistance to bacterial leaf streak in barley, caused by Xanthomonas campestris pv. hordei, was investigated in 119 doubled-haploid lines (DH) developed by the Hordeum bulbosum method from the F₁ progeny of the cross between two cultivars, ‘Morex’ (resistant) and ‘Steptoe’ (susceptible). Two experiments were undertaken in a randomized complete block design with three replicates, in a controlled growth chamber. Twenty seeds per replicate were planted in plastic containers (60×40×8 cm) containing moistened vermiculite. At the two-leaf stage seedlings were inoculated with an Iranian strain of the pathogen. Genetic variability was observed among the 119 DH lines for partial resistance to the disease. Some DH lines were significantly more resistant than ‘Morex’ (resistant parent) to bacterial leaf streak. Genetic gain in percentage of resistant parent for 5% of the selected DH lines was significant (47.70% and 33.72% in the first and the second experiment, respectively). A QTL analysis of bacterial leaf streak resistance showed that three QTLs were detected on chromosomes 3 and 7. Multilocus allelic effects of the three QTLs account for almost 54% of the mean difference between the parents and nearly 30% of the phenotypic variation of the trait in the mean experiment. The resistance locus on chromosome 3, near ABG377, apprears to be a major gene. Received: 15 July 1997 / Accepted: 4 August 1997     

Isolate specificity of quantitative trait loci for partial resistance of barley to Puccinia hordei confirmed in mapping populations and near-isogenic lines

Partial resistance is considered race-nonspecific and durable, consistent with the concept of 'horizontal' resistance. However, detailed observations of partial resistance to leaf rust (Puccinia hordei) in barley (Hordeum vulgare) revealed small cultivar x isolate interactions, suggesting a minor-gene-for-minor-gene interaction model, similar to so-called 'vertical' resistance. Three consistent quantitative trait loci (QTLs), labelled Rphq2, Rphq3 and Rphq4, that were detected in the cross susceptible L94 x partially resistant Vada have been incorporated into the L94 background to obtain near-isogenic lines (NILs). Three isolates were used to map QTLs on seedlings of the L94 x Vada population and to evaluate the effect of each QTL on adult plants of the respective NILs under field conditions. Rphq2 had a strong effect in seedlings but almost no effect in adult plants, while Rphq3 was effective in seedlings and in adult plants against all three isolates. However, Rphq4 was effective in seedlings and in adult plants against two isolates but ineffective in both development stages against the third, demonstrating a clear and reproducible isolate-specific effect. The resistance governed by the three QTLs was not associated with a hypersensitive reaction. Those results confirm the minor-gene-for-minor-gene model suggesting specific interactions between QTLs for partial resistance and P. hordei isolates.     

Mapping genes for resistance to barley stripe rust (Puccinia striiformis f. sp. hordei)

Two genes conferring resistance to the barley stripe rust found in Mexico and South America, previously identified as race 24, were mapped to the M arms of barley chromosomes 7 and 4 in a doubled haploid population using molecular markers and the quantitative trait loci (QTL) mapping approach. The resistance gene on chromosome 7 had a major effect, accounting for 57% of the variation in disease severity. The resistance gene on chromosome 4 had a minor effect, accounting for 10% of the variation in trait expression. Two pairs of restriction fragment length polymorphism markers are being used to introgress the resistance genes to North American spring barley using molecular marker-assisted backcrossing.Ore. Agric Exp Stn J no. 10283     

Inheritance of resistance to covered smut in barley and development of a tightly linked SCAR marker

Inheritance of resistance to covered smut in the barley line Q21861 was studied using a doubled-haploid population produced by crossing Q21861 with the line SM89010. Based on 3 years of screening in the field and two seasons in the greenhouse, segregation for resistance/susceptibility fits a one-gene ratio, indicating a single major gene for resistance in Q21861. Of 440 random 10-mer primers tested using bulked segregant analysis, one primer (OPJ10) resulted in a reproducible polymorphic band. RAPD marker OPJ10₄₅₀ co-segregated in repulsion with the covered smut resistance. This marker was converted to a sequence-characterized amplified region (SCAR) marker linked in coupling (5.5 cM) with the covered smut resistant gene in Q21861. The SCAR marker was amplified in the line TR640 which is also resistant to covered smut, but not in the other resistant lines. The SCAR marker will be useful for marker-assisted selection for covered smut in barley breeding programs. Received: 9 January 2001 / Accepted: 31 May 2001     

Mapping resistance to cereal aphids in barley

 A set of 150 doubled-haploid (DH) barley (Hordeum vulgare L.) lines derived from the cross of Harrington/TR306 was used to determine the number and chromosomal location of quantitative trait loci (QTLs) controlling resistance to cereal aphids. The experiments were conducted under natural infestation in the field during two growing seasons: 1994 and 1995. Aphid resistance was measured by counting the number of aphids per plot. Counts were made on a weekly basis. Each year at the time of maximum aphid density there was an obvious difference in reaction between the parental genotypes. The DH lines showed continuous variation for aphid density. Simple interval mapping and simplified composite interval mapping revealed that the principal QTL determining cereal aphid resistance is on the distal region of the short arm of chromosome 1. This aphid-resistance QTL is linked with a heading-date QTL. At the time of highest aphid infestation, this QTL accounted for 31% and 22% of the total variance of aphid density in 1994 and 1995, respectively. A QTL on chromosome 5 was also detected but only by simplified composite interval mapping. However, the largest consistent effect was due to the QTL on the short arm of chromosome 1. This QTL may be a useful target for marker-assisted selection for adult plant cereal aphid resistance in barley. Received: 10 September 1996/Accepted: 11 October 1996     

DNA markers for Fusarium head blight resistance QTLs in two wheat populations

Genetic resistance to Fusarium head blight (FHB), caused by Fusarium graminearum, is necessary to reduce the wheat grain yield and quality losses caused by this disease. Development of resistant cultivars has been slowed by poorly adapted and incomplete resistance sources and confounding environmental effects that make screening of germplasm difficult. DNA markers for FHB resistance QTLs have been identified and may be used to speed the introgression of resistance genes into adapted germplasm. This study was conducted to identify and map additional DNA markers linked to genes controlling FHB resistance in two spring wheat recombinant inbred populations, both segregating for genes from the widely used resistance source ’Sumai 3’. The first population was from the cross of Sumai 3/Stoa in which we previously identified five resistance QTLs. The second population was from the cross of ND2603 (Sumai 3/Wheaton) (resistant)/ Butte 86 (moderately susceptible). Both populations were evaluated for reaction to inoculation with F. graminearum in two greenhouse experiments. A combination of 521 RFLP, AFLP, and SSR markers were mapped in the Sumai 3/Stoa population and all DNA markers associated with resistance were screened on the ND2603/Butte 86 population. Two new QTL on chromosomes 3AL and 6AS wer found in the ND2603/Butte 86 population, and AFLP and SSR markers were identified that explained a greater portion of the phenotypic variation compared to the previous RFLP markers. Both of the Sumai 3-derived QTL regions (on chromosomes 3BS, and 6BS) from the Sumai 3/Stoa population were associated with FHB resistance in the ND2603/Butte 86 population. Markers in the 3BS QTL region (Qfhs.ndsu-3BS) alone explain 41.6 and 24.8% of the resistance to FHB in the Sumai 3/Stoa and ND2603/Butte 86 populations, respectively. This region contains a major QTL for resistance to FHB and should be useful in marker-assisted selection. Received: 17 August 2000 / Accepted: 16 October 2000     

Molecular mapping of a new gene in wild barley conferring complete resistance to leaf rust (Puccinia hordei Otth)

 A dominant gene conferring resistance to all known races of Puccinia hordei Otth was identified in two accessions of Hordeum vulgare ssp. spontaneum. Using restriction fragment length polymorphism (RFLP) markers the gene was mapped on chromosome 2HS in doubled-haploid populations derived from crosses of both accessions to the susceptible cultivar L94. Until now, complete leaf rust resistance was not known to be conditioned by genetic factors on this barley chromosome. Therefore, the designation Rph16 is proposed for the gene described in this study. A series of sequence tagged site (STS) and cleaved amplified polymorphic sequence (CAPS) markers were generated by conversion of RFLP probes which originate from the chromosomal region carrying the resistance gene. Two PCR-based markers were shown to co-segregate with the Rph16 gene in both populations thus providing the basis for marker-assisted selection. Received: 20 May 1998 / Accepted: 9 June 1998     

The maize rp1 rust resistance gene identifies homologues in barley that have been subjected to diversifying selection

A number of agronomically important grasses (sorghum, wheat, panicum, sugar cane, oats, rice and barley) are shown to contain sequences homologous to rp1, a maize gene that confers race-specific resistance to the rust fungus Puccinia sorghi. Mapping of rp1-related sequences in barley identified three unlinked loci on chromosomes 1HL, 3HL and 7HS. The locus located on chromosome 7HS comprises a small gene family of at least four members, two of which were isolated and are predicted to encode nucleotide binding site-leucine-rich repeat (NBS-LRR) proteins that are respectively 58% and 60% identical to the maize rp1 protein. Evidence of positive selection for sequence diversification acting upon these two barley genes was observed; however, diversifying selection was restricted to the carboxy terminal half of the LRR domain. One of these rp1 homologous genes cosegregated with the barley Rpg1 stem rust resistance gene amongst 148 members of the Steptoe × Morex double haploid mapping family. Three other unrelated resistance gene-like sequences, potentially encoding NBS-LRR proteins, are also shown to be linked to the Rpg1 locus but not cosegregating with the gene. Received: 2 August 1999 / Accepted: 28 September 1999     

Inheritance of resistance to southern corn rust in tropical-by-corn-belt maize populations

 The inheritance of resistance to southern rust (caused by Puccinia polysora Underw.) was investigated in two F_2:3 populations derived from crossing two temperate-adapted, 100% tropical maize (Zea mays L.) inbred lines (1416-1 and 1497-2) to a susceptible Corn Belt Dent hybrid, B73Ht×Mo17Ht. The inbred lines possess high levels of resistance to southern rust and may be unique sources of resistance genes. Heritability for resistance was estimated as 30% and 50% in the two populations from regression of F_2:3 family mean scores on F₂ parent scores, and as 65% and 75% from variances among F_2:3 families on a single-plot basis. RFLP loci on three chromosomal regions previously known to possess genes for resistance to either southern rust or common rust (P. sorghi Schw.) were used to localize genes affecting resistance to southern rust in selected genotypes of both populations, and to estimate their genetic effects. A single locus on 10S, bnl3.04, was associated with 82–83% of the variation among field resistance scores of selected F_2:3 families in the two populations. Loci on chromosomes 3 (umc26) and 4 (umc31) were significantly associated with resistance in the 1497-2 population, each accounting for 13–15% of the phenotypic variation for F_2:3 field scores. Multiple-marker locus models, including loci from chromosomes 3, 4, and 10 and their epistatic interactions, accounted for 96–99% of the variation in F_2:3 field scores. Similar results were obtained for resistance measured by counting pustules on juvenile plants in the greenhouse. An attempt was made to determine if the major gene for resistance from 1416-1 was allelic to Rpp9, which is also located on 10S. Testcross families from the cross (1416-1×B37Rpp9)×B14AHt were evaluated for resistance to southern rust in Mexico. Neither source of resistance was completely effective in this environment, preventing determination of allelism of the two genes; however, both sources of resistance had better partial resistance to southern rust than did B14AHt. Received: 6 May 1997/Accepted: 19 September 1997     

QTL analysis of resistance to the rice blast pathogen in barley (Hordeum vulgare)

Barley is compatible with the rice blast pathogen (Pyricularia oryzae Cav.). Fiftyfour barley cultivars of diverse geographic origin and pedigree were inoculated with three isolates of the rice blast pathogen. All barley genotypes showed blast disease symptoms when inoculated at the seedling stage with each of the three isolates. However, one genotype showed quantitative resistance to all three isolates and three genotypes showed quantitative resistance to one or two of the isolates. By inoculating a set of doubled-haploid lines derived from the cross ’Harrington’ (susceptible) and ’TR306’ (resistant) with isolate Ken 54–20, we mapped quantitative trait loci (QTLs) determining seedling stage blast resistance. At all QTLs, TR306 contributed the resistance alleles. The four QTLs, when considered jointly, explained 43.6% of the phenotypic variation in blast symptom expression. A comparison of the blast resistance QTLs with other disease resistance QTLs reported in this population revealed a region on chromosome 4 (4H) with multiple disease resistance loci. It will be useful to capitalize on the syntenic relationship of rice and barley and to integrate information on species-specific resistance genes with information on the reaction of the two species to the same pathogen. Received: 7 January 2000 / Accepted: 22 September 2000     

Several QTLs involved in osmotic-adjustment trait variation in barley (Hordeum vulgare L.)

 Osmotic adjustment (OA) was previously demonstrated to be an important adaptive mechanism of drought tolerance in cereals. In order to determine which genomic regions are involved in OA variation, 187 barley (Hordeum vulgare L.) recombinant inbred lines (RILs) derived from a cross between Tadmor (drought tolerant) and Er/Apm (susceptible) were studied in a growth chamber for their OA capacity (through correlated traits and by calculation), at an early growth stage and under two water treatments (soil moisture of 14% and 100% of field capacity). The continuous distribution of the traits and their broad-sense line heritabilities, ranging from 0.04 to 0.44, indicated that OA and related traits should have a polygenic nature. A subset of 167 RILs were also genotyped using 78 RFLP, 32 RAPD and three morphological markers and a linkage map was constructed. Despite strong environmental effects acting on the traits, interval mapping and single-marker ANOVA allowed the detection of three QTLs for relative water content (RWC), four QTLs for osmotic potential (ψ_π), two QTLs of osmotic potential at full turgor (ψ_π100) and one QTL for osmotic adjustment at a soil moisture of 14% field capacity. For the irrigated treatment, only two QTLs were detected: one for RWC and one for ψ_π100. Two chromosomal regions were involved in several OA-related trait variations and could be considered as regions controlling OA; these were present on chromosome 1 (7H) and chromosome 6 (6H), whereas other regions were specific for one trait. No major QTL was found. However, the genomic region involved in OA-related traits on chromosome 1 (7H) in barley seemed to be conserved for OA variation among cereals. Epistatic effects, with or without additive effects, acted on the traits. Received: 15 July 1997 / Accepted: 29 October 1997     

QTLs for a component of partial resistance to cucumber mosaic virus in pepper: restriction of virus installation in host-cells

 Ninety four doubled-haploid (DH) lines obtained from the F₁ between Perennial, a cucumber mosaic virus (CMV)-partially resistant Capsicum annuum line, and Yolo Wonder, a CMV-susceptible C. annuum line, were analysed with 138 markers including mostly RFLPs and RAPDs. Clustering of RAPD markers was observed on five linkage groups of the intraspecific linkage map. These clusters could correspond to the centromeric regions of pepper chromosomes. The same progenies were evaluated for restriction of CMV installation in pepper cells in order to map quantitative trait loci (QTLs) controlling CMV resistance. This component of partial resistance to CMV was quantitatively assessed using a CMV strain that induced necrotic local lesions on the inoculated leaves. The number of local lesions gave an estimation of the density of the virus-infection sites. Genotypic variance among the DH lines was highly significant for the number of local lesions, and heritability was estimated to be 0.94. Using both analysis of variance and non-parametric tests, three genomic regions significantly affecting CMV resistance were detected on chromosomes Noir, Pourpre and linkage group 3, together explaining 57% of the phenotypic variation. A digenic epistasis between one locus that controlled significant trait variation and a second locus that by itself had no demonstrable effect on the trait was found to have an effect on CMV resistance. For each QTL, the allele from Perennial was associated with an increased resistance. Implications of QTL mapping in marker-based breeding for CMV resistance are discussed. Received: 16 September 1996     

QTLs for agronomic traits from a Mediterranean barley progeny grown in several environments

In order to identify quantitative trait loci (QTLs) controlling agronomic trait variation and their consistency under Mediterranean conditions in barley, a progeny of 167 recombinant inbred lines (RILs) and the parents Tadmor and Er/Apm, originating from the Mediterranean basin, were grown under Mediterranean conditions in 1995, 1996, 1997 and 1999. For the 2 first years (M95 and G96), one replicate was grown, but for the latter (M97 and M99) two rainfed (rain) and two irrigated (ir) replicates were produced. M95, G96, M97rain, M97ir, M99rain and M99ir were considered as six different environments and were compared in terms of their meteorological conditions and water supply. Grain yield and yield components were assessed, as well as heading date and plant height. Highly significant differences were noted between environments. QTLs were obtained from each environment separately and from a multiple environment analysis (simple interval mapping and simplified composite interval mapping). Despite heterogeneity between environments, numerous QTLs were common to several environments. This was particularly true for traits like plant height and thousand-grain weight. The most reliable QTLs which explained the largest part of the phenotypic variation were obtained for plant height on chromosomes 3 (3H) and 6 (6H). The multiple-environment analysis provided an opportunity to identify consistent QTLs for agronomic traits over six Mediterranean environments. A total of 24 consistent QTLs were detected. Out of these, 11 presented main effects, seven presented QTL×E interaction, and six presented both effects. In addition, 18 of the consistent QTLs were common to other published work and six seemed specific to this study. These latter QTLs could be involved in Mediterranean adaptive specificities or could be specific to the studied genetic background. Finally, when the rainfed and the irrigated environments of M97 were considered separately, a total of 16 QTLs presenting main effects over the two water conditions were identified, whereas five QTLs seemed dependent on the water conditions. Received: 31 January 2001 / Accepted: 19 February 2001     

Identification and mapping of a new leaf stripe resistance gene in barley (Hordeum vulgare L.)

Pyrenophora graminea is the seed-borne pathogen causal agent of barley leaf stripe disease. Near-isogenic lines (NILs) carrying resistance of the cv ”Thibaut” against the highly virulent isolate Dg2 were obtained by introgressing the resistance into the genetic background of the susceptible cv ”Mirco”. The segregation of the resistance gene was followed in a F₂ population of 128 plants as well as on the F₃ lines derived from the F₂ plants; the segregation fitted the 1:2:1 ratio for a single gene. By using NILs, a RAPD marker associated with the resistance gene was identified; sequence-specific (STS) primers were designed on the basis of the amplicon sequence and a RILs mapping population with an AFLP-based map were used to position this molecular marker to barley chromosome 1 S (7HS). STS and CAPS markers were developed from RFLPs mapped to the telomeric region of barley chromosome 7HS and three polymorphic PCR-based markers were developed. The segregation of these markers was followed in the F₂ population and their map position with respect to the resistance gene was determined. Our results indicate that the Thibaut resistance gene, which we designated as Rdg2a, maps to the telomeric region of barley chromosome 7HS and is flanked by the markers OPQ-9₇₀₀ and MWG 2018 at distances of 3.1 and 2.5 cM respectively. The suitability of the PCR-based marker MWG2018 in selection- assisted barley breeding programs is discussed. Received: 22 June 2000 / Accepted: 16 October 2000     

Dynamic gene action at QTLs for resistance to Setosphaeria turcica in maize

 Cultivars with quantitative resistance are widely used to control Setosphaeria turcica (Luttrell) Leonard & Suggs, the causal organism of northern corn leaf blight (NCLB). Here the effectiveness of quantitative trait loci (QTLs) for NCLB resistance was investigated over the course of host plant development in inoculated field trials. A population of 194–256 F_2:3 lines derived from a cross between a susceptible Italian (Lo951) and a highly resistant African inbred line (CML202) was tested in three environments in Kenya. The traits assessed were the incubation period (IP), the percentage disease severity (DS 1 to 5, taken biweekly), and the area under the disease progress curve (AUDPC). Considering all resistance traits and environments, a total of 19 putative QTLs were detected by composite interval mapping using a linkage map with 110 RFLP markers. In the combined analysis across environments, nine QTLs were significant (LOD >3.0) for DS 3, recorded around flowering time, explaining 71% of the genotypic variance. Four of these nine QTLs displayed significant (P<0.05) QTL×environment (QTL×E) interaction. Most QTLs were already significant in the juvenile stage (IP) and became less effective after flowering. Across environments, three QTLs conditioned adult-plant resistance, in the sense that they were only significant after flowering. Six QTL alleles on chromosomes 2, 4, 5, 8, and 9 of CML202 should be useful for marker-assisted backcrossing. Received: 24 August 1998 / Accepted: 29 September 1998