Resistance toMelampsora larici-epitea leaf rust inSalix: analyses of quantitative trait loci

Quantitative resistance ofSalix toMelampsora larici-epitea leaf rust was studied in 2Salix mapping populations. One population was a backcross between aS. schwerinii ×S. viminalis hybrid andS. viminalis, and the other was an F₂ population betweenS. viminalis andS. dasyclados. A leaf disc bioassay was used to study the components of quantitative resistance (latent period, uredinia number, and uredinia size) to 3 isolates of the leaf rust. The analysis of quantitative trait loci (QTLs) revealed 9 genomic regions in the backcross population and 7 genomic regions in the F₂ population that were important for rust resistance, with QTLs explaining 8–26% of the phenotypic variation. An important genomic region was identified for the backcross population in linkage group 2, where QTLs were identified for all resistance components for 2 of the rust isolates. Four of the QTLs had overlapping mapping intervals, demonstrating a common genetic background for latent period, uredinia diameter, and uredinia number. QTLs specific to some rust isolates and to some resistance components were also found, indicating a combination of common and specific mechanisms involved in the various resistance components. Breeding implications in relation to these findings are discussed.     

Components of Partial Disease Resistance in Wheat Detected in a Detached Leaf Assay Inoculated with Microdochium majus using First, Second and Third Expanding Seedling Leaves

The use of first, second and third expanding seedling leaves of wheat (L1, L2 and L3 respectively), inoculated with conidial suspensions of Microdochium majus (syn. Microdochium nivale var. majus) in a detached leaf assay, for detecting components of partial disease resistance (PDR) was investigated across a range of wheat cultivars. Incubation periods (period from inoculation to first appearance of symptoms; a dull grey–green water‐soaked lesion) and latent periods (period from inoculation to the first appearance of sporodochia) were longest and lesions smallest on L3. The expression of PDR components on L2 was intermediate to those on L1 and L3. The longer latent periods on L3 typically occurred after leaf senescence contrasting with latent periods on L1 and L2 where sporulation most frequently occurred on relatively green leaf tissue. Cultivar differences in the first appearance of symptoms, incubation period, which occurred before any leaf senescence was observed, correlated significantly across all leaf positions. Similarly cultivar differences in latent period were correlated for L1 and L2. However, latent periods on L3, which were the least consistent between cultivars across experiments, were not correlated with those of L1 or L2 in any experiment. The results indicate that due to the delay in sporulation until after leaf senescence, observations on latent period in L3 are less representative of what occurs in the whole plant where infection of living tissue is of greatest interest. This work indicates that the selection of the first or second expanding leaf of wheat is optimal for the use in the detached leaf assay using M. majus for studying components of PDR.     

Effects of Infection Density on the Size of Barley and Wheat Leaf Rust Colonies before and on the Size of Uredia after the Start of Sporulation

Three barley genotypes were exposed to four different inoculum densities of barley leaf rust, Puccinia hordei. Colony area well before first sporulation and uredia size well after the start of sporulation were measured. In a second series two barley and two wheat cultivars were exposed to four different inoculum densities of barley and wheat leaf rust (P. recondita f. sp. tritici), respectively. Before the sporulation is initiated the colony size was independent of the uredia density. This was valid for densities ranging from approximately 7 to over 200 uredia per cm² leaf area. After the sporulation had started the uredia size was strongly dependent on the uredia density. The size of the uredia was approximately halved when the uredia density increased from about 10 to about 150 per cm². The urediospore production per uredium decreased much stronger with increased uredia density.     

Inheritance of resistance to yellow rust (Phragmidium rubi-idaei) in red raspberry

The inheritance of resistance in red raspberry (Rubus idaeus) to yellow rust (Phragmidium rubi-idaei) was studied in a diallel cross among the cultivars Boyne, Meeker, Mailing Jewel, Glen Prosen and Glen Clova. The progenies and clonally propagated parents were exposed initially in a rust-infected plantation where the incidence and severity of infection was assessed at the telial stage. The following spring leaves on new canes of the same plants were inoculated with urediniospores in a glasshouse at 18 ± 2 °C. The latent period and number of uredinia per cm² of abaxial leaf surface were determined 13 and 18 days after inoculation. Cv. Boyne developed only chlorotic flecking on inoculated leaves in the glasshouse and no sporulation was observed. It is postulated that this reaction was determined by a major gene which is designated Yr. Some of the other parents showed incomplete resistance of the slow rusting type, and segregants of Boyne selfed that were susceptible also showed low levels of rust, which suggests that Boyne carries both major- and minor-gene resistance. Diallel analyses of both the field and glasshouse data from the susceptible segregates indicated that the degree of incomplete resistance present was determined predominantly by additive gene action, though small but significant non-additive effects also occurred: cv. Meeker was the most resistant parent and contributed the most resistance to the progenies. The Meeker progenies also segregated for gene H, which determines cane hairiness and is known to be associated with resistance to other diseases. Segregants without hairs (genotypes hh) were found to be the more resistant to rust.     

Genetic research as the valid base of strategies for breeding rust resistant wheats

It is known that few wheat cultivars maintain their resistance to rust diseases for a long period of time, particularly when crop populations become genetically more uniform. A number of genetically diverse, so far unexploited, sources of rust resistance in the natural as well as mutagenized population of wheat cultivars were identified. Several of these genes were placed in agronomically superior well-adapted backgrounds so that they could be used as pre-breeding stocks for introducing genetic diversity for resistance in a crop population. Some of these stocks when employed as parents in several cross combinations in a breeding programme have generated a number of promising cultivars with diversity for resistance.Many presently grown wheats in India, near-isogenic lines each with Lr14b, Lr14ab, Lr30 and certain international cultivars were identified as possessing diverse sources of adult plant resistance (APR) to leaf rust. Prolonged leaf rust resistance in some of the Indian cultivars was attributed to the likely presence of Lr34 either alone or in combination with other APR components. Tests of allelism carried out in certain cultivars that continue to show adequate levels of field resistance confirm the presence of Lr34, which explains the role that this gene has played in imparting durability for resistance to leaf rust. Also, Lr34 in combination with other APR components increases the levels of resistance, which suggests that combination of certain APR components should be another important strategy for breeding cultivars conferring durable and adequate levels of resistance. A new adult plant leaf rust resistance source that seems to be associated with durability in Arjun has been postulated. Likewise, cultivars possessing Sr2 in combination with certain other specific genes have maintained resistance to stem rust.Further, non-specific resistances that were transferred across widely different genotypes into two of the popular Indian wheats provided easily usable materials to the national breeding programmes for imparting durable resistance to stripe rust.     

Observed and hypothetical leaf rust progress curves of some genotypes of wheat

Summary The hypothetical leaf rust progress curves of 15 genotypes of wheat were generated by integrating the components of slow rusting resistance determined in the glasshouse. The area under the disease progress curve (AUDPC) from the hypothetical and observed leaf rust progress curves were compared. It was found that the hypothetical AUDPC values of all the genotypes studied were smaller than their respective observed AUDPC values. Possible causes for the discrepancies in the observed and hypothetical values are discussed.     

Wheat leaf photosynthesis loss due to leaf rust, with respect to lesion development and leaf nitrogen status

In wheat (Triticum aestivum cv. Soissons) plants grown under three different fertilisation treatments, we quantified the effect of leaf rust (Puccinia triticina) on flag leaf photosynthesis during the whole sporulation period. Bastiaans' model: Y = (1 - x)beta was used to characterize the relationship between relative leaf photosynthesis (Y) and disease severity (x). The evolution of the different types of symptoms induced by the pathogen (sporulating, chlorotic and necrosed tissues) was evaluated using image analysis. The beta-values varied from 2 to 11, 1.4-2, and 0.8-1 during the sporulation period, when considering the proportion of sporulating, sporulating + necrotic, and total diseased area, respectively. Leaf nitrogen (N) content did not change the effect of the disease on host photosynthesis. We concluded that leaf rust has no global effect on the photosynthesis of the symptomless parts of the leaves and that the large range in the quantification of leaf rust effect on the host, which is found in the literature, can be accounted for by considering the different symptom types. We discuss how our results could improve disease assessments and damage prediction in a wheat crop.     

Comparison of the Wheat Brown and Yellow Rusts for Monocyclic Sporulation and Infection Processes, and their Polycyclic Consequences

Sporulation capacity and infection efficiency of wheat brown and yellow rusts were measured daily in favourable controlled conditions. Monocyclic sporulation capacity for a single lesion of yellow rust was 9 times greater than for an isolated lesion of brown rust, and 40 times greater than for a lesion of brown rust at medium infection density. Infection efficiency fluctuated and reached about 40 % for brown rust but remained under 5% for yellow rust. For both fungi, sporulation capacity and infection efficiency compensated for each other, but their product, the daily multiplication factor, was greater for yellow rust than for brown rust. Progeny/parent ratio was 3 times greater for yellow rust. Effect of daily multiplication factor variations on epidemic progress was simulated using a simple matrix model. Increase in number of lesions was faster in brown rust than in yellow rust because of a latent period shorter by 2 days. Semi-systemic growth of yellow rust fungus reduced, however, the difference between both fungi when sporulating surface was calculated.     

Interaction between barley yellow dwarf virus and rust in wheat, barley and oats, and the effects on grain yield and quality

In three separate experiments, the upper leaf surface of the fifth formed leaf of wheat cv. Highbury, the fourth and fifth leaves of barley cv. Julia and the third and fourth leaves of oat cv. Mostyn were inoculated in a spore settling tower with wheat brown rust (Puccinia recondita f. sp. tritici), barley brown rust (P. hordei) or oat crown rust (P. coronata f. sp. avenae), respectively. Fewer pustules developed on distal portions of leaves of plants infected with barley yellow dwarf virus (BYDV) than on similar portions of leaves from virus-free plants. There were no significant differences in the number of pustules on proximal leaf portions. In barley and oats, the number of pustules on distal leaf portions was negatively correlated with the amount of yellowing of the leaf areas scored. In wheat, symptoms of BYDV were mild and leaves were little affected by yellowing. The latent period of rust on wheat and oats was not affected by BYDV. In barley, BYDV reduced the latent period of rust on leaf 5, but not on leaf 4, and reduced it on proximal, but not distal, leaf portions. In other experiments, BYDV reduced the yield of wheat and oats by 44% and 66%, respectively, while BYDV-infected barley was almost sterile. The appropriate rust reduced the yield of wheat, barley and oats by 33%, 13% and 86%, respectively. When infected with both BYDV and rust, yield of wheat and oats was reduced by 63% and 91%, respectively. Neither BYDV nor rust affected the percentage crude protein content of wheat grain, nor did rust affect that of barley. In oats, BYDV and rust each significantly increased crude protein of grain, but rust infection of BYDV-infected plants tended to reduce it.     

Histology of the Infection and Development of Puccinia recondita f. sp. tritici in a Wheat Line with Lr37

The degree of adult-plant resistance conferred by the gene Lr37 in RL6081 (Thatcher*8/Lr37) to four avirulent leaf rust pathotypes was quantified by assessing histological components as well as latent period, uredinium density and uredinium size. Histological observations on the adaxial flag leaf surfaces revealed significant arrest of fungal structures in RL6081 at early infection stages. Furthermore, host cell necrosis typical of a posthaustorial host-resistance mechanism was conspicuous in sizeably reduced colonies on this genotype. Lr37 significantly decreased the rate of uredinial appearance of all four pathotypes. Compared with Thatcher, fewer uredinia of smaller dimensions developed on flag leaves of RL6081. Characterization of resistance indicated that disease development in genotypes with the gene Lr37 should be extremely limited.     

Genetic architecture of qualitative and quantitative Melampsora larici-populina leaf rust resistance in hybrid poplar: genetic mapping and QTL detection

In order to elucidate the genetic control of resistance to Melampsora larici-populina leaf rust in hybrid poplars, a Populus deltoides x P. trichocarpa F(1) progeny was analysed for qualitative and quantitative rust resistances. This progeny was evaluated for three components of quantitative resistance (latent period, uredinia number and uredinia size) to seven M. larici-populina strains in controlled conditions, and for one component of field susceptibility (rust colonization on the most infected leaf). One qualitative resistance locus inherited from P. deltoides, R(1), was localized on the genetic map. It segregates 1 : 1 in the F(1) progeny and is effective against four of the studied strains. QTL analysis was performed separately on R(1) and r(1) genotype subsets. An additional detection was conducted on the entire F(1) progeny for the three strains able to overcome R(1) and for MAX2. A total of nine QTLs were detected. Two had large, broad-spectrum effects. One (R(US)) is inherited from the P. trichocarpa parent; the other is inherited from P. deltoides and colocalized with R(1). Seven QTLs had only limited and specific effects. Significant interaction effects were detected mainly between the two major QTLs. Implications of these results for durable resistance breeding strategies, and possible benefits from the Populus genome sequence, are discussed.     

The mechanism and inheritance of adult plant leaf rust resistance in 12 wheat lines.

Twelve lines of wheat (Triticum aestivum L.) were developed that had susceptible infection types to leaf rust (Puccinia recondita Rob. ex Desm. f.sp. tritici) race UN 15 in the seedling stage but were resistant in the adult plant stage in the field. The lines were developed from four crosses, each involving four parents (eight in total) that had originally been selected for adult plant or field resistance to stem rust (Puccinia graminis Pers. f.sp. tritici Eriks, and Henn.). The objectives of the present study were to determine the mechanism of resistance to leaf rust and its inheritance in the 12 lines. The 12 lines were grown in an artificially inoculated field nursery in Saskatoon, coefficients of infection (CI) were determined at four dates, and the areas under the disease progress curve (AUDPC) were calculated. Four representative lines were grown in a growth chamber to measure the latent period and pustule size at the two-leaf and flag-leaf stages. Eight lines were crossed and backcrossed to a susceptible check and the parents, F1, F2, F3, and BC1F2 generations were grown in a field nursery. The 12 lines showed wide ranges in CI and AUDPC but all were significantly more resistant than the susceptible check. The four lines studied in the growth chamber had longer latent periods and smaller pustules than the susceptible check at both stages. The differences tended to be greater at the flag-leaf stage. The inheritance studied showed that resistance was recessive or partially recessive and was controlled by two or more genes in each line of the eight lines. In three of the eight lines, Lr34 may be one of the genes and in the other five both Lr13 and Lr34 could be present. However, additional genes are clearly involved. A single gene by itself had only a small effect, but in two and three gene combinations the effects appeared to be greater. This type of resistance appears to occur frequently and may be durable because its complex inheritance may make it more difficult for the rust fungus to overcome. It should be used in breeding wheat for areas where leaf rust is a major problem.     

Mapping of QTLs prolonging the latent period of Puccinia triticina infection in wheat

Slow rusting is considered a crucial component of durable resistance to wheat leaf rust caused by Puccinia triticina and is often expressed in the form of a prolonged latent period. Selection for a longer latent period is considered an effective approach to developing wheat cultivars with improved durable resistance to leaf rust. A recombinant inbred line (RIL) population derived from CI 13227 (long latent period) × Suwon 92 (short latent period) was phenotyped for latent period in two greenhouse experiments in separate years, and amplified fragment length polymorphism (AFLP) and simple sequence repeat (SSR) markers were analyzed in the same population. Among the RILs, the frequency distribution for latent period was continuous, and latent period was highly correlated between years (r=0.94, P<0.0001). A quantitative trait locus (QTL) prolonging the latent period of P. triticina, designated as QLrlp.osu-2DS, explained 42.8% and 54.5% of the phenotypic and genetic variance in the two experiments, respectively. QLrlp.osu-2DS was mapped on the distal region of chromosome 2DS. Two other QTLs for latent period, QLrlp.osu-2B and QLrlp.osu-7BL, were localized on chromosome 2B and the long arm of chromosome 7B, respectively. Multiple regression analysis showed that these three QTLs collectively explained 58.0% and 73.8% of the phenotypic and genetic variance over two experiments, respectively. Fourteen RILs that carried all three alleles for long latent period at three AFLP loci flanking QLrlp.osu-2DS, QLrlp.osu-2B, and QLrlp.osu-7BL had a mean latent period of 12.5 days, whereas 13 RILs without any long-latent-period alleles at the corresponding loci had a mean latent period of 7.4 days. Three SSR markers closely linked to these QTLs have potential to be applied in marker-assisted selection for prolonged latent period in wheat.     

Number of genes controlling slow rusting resistance to leaf rust in five spring wheat cultivars

The number of genes controlling slow rusting resistance to leaf rust (Puccinia triticina) was estimated in five spring wheat (Triticum aestivum) cultivars using quantitative formulae. Parents and F₆ families were evaluated in replicated field trials under epidemics initiated by artificial inoculation. The F₆ families resulted from a diallel cross involving the fast-rusting cultivar Yecora 70 and five slow-rusting wheat cultivars: Sonoita 81, Tanager ‘S’, Galvez 87, Ures 81, and Moncho ‘S’. The area under the disease progress curve (AUDPC) was used to measure leaf rust severity over time. Results indicate that cultivar Sonoita 81 has three or four genes, Tanager ‘S’ has two or three genes, Galvez 87 has three genes, and both Ures 81 and Moncho ‘S’ have two genes for slow rusting resistance to leaf rust. Based on this result and previously reported moderate to high narrow-sense heritability estimates for slow rusting resistance in these materials, early-generation selection for slow leaf rusting would be effective.     

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     

Race non-specific resistance to Bremia lactucae

Three ways of measuring race non-specific resistance to Bremia lactucae in lettuce were studied to determine the most important components of this form of resistance. Tests were conducted on leaf discs from 4–6 wk-old plants that were quantitatively inoculated and incubated under controlled conditions. Observations were made of disease intensity, latent period and production of conidia. Inoculum concentration markedly influenced disease intensity and latent period. Cv. Iceberg exhibited race non-specific resistance and this was characterised by a longer latent period, a lower disease intensity and reduced production of conidia compared to the susceptible control cv. Cobham Green. On older, intact plants, the differences in latent period and disease intensity between cultivars were even greater. Production of conidia was a particularly good criterion for discriminating levels of resistance.     

Evaluation of Macroscopic and Microscopic Components of Partial Resistance to Leaf Rust in Durum Wheat

Leaf rust, caused by the fungus Puccinia triticina, is considered one of the most important foliar diseases in durum wheat. Hypersensitive resistance (HR) may be rapidly overcome by the pathogen when resistant cultivars are grown on a large acreage or following changes in virulence in the pathogen population. Prolonging the durability of the resistance requires uses of other types of resistance such as partial resistance (PR). In this study, six durum wheat lines provided by the International Center for Corn and Wheat Improvement (CIMMYT) with a high level of PR to leaf rust were studied in monocyclic tests in a growth chamber. Inoculations were performed on both primary and fifth leaves using the Spanish race DGB/BN. UV fluorescence microscopy was employed to determine microscopic components of the resistance, such as the number of early aborted infection units not associated with plant cell necrosis (EA?) and relative colony size (RCS) of the established infection units. Macroscopic components of PR such as latency period, infection frequency and uredinium size were measured as well. All six resistant lines were characterized by a higher EA? and smaller RCS respect to the susceptible control ‘Don Rafael’. Line 3 showed the highest level of PR. It had 22% of EA? compared with 4% in the susceptible control, and the smallest RCS (17% respect to RCS of ‘Don Rafael’) at adult plant stage. Both EA? and RCS had a high heritability (more than 97%) and the correlation with macroscopic parameters (latency period and uredinium size) was also high (significant at 0.001 level). Hence, PR to leaf rust in these durum wheat genotypes has been revealed at microscopic level (higher EA? and smaller RCS).     

Determination of Rust Resistance Gene Complex Lr34/Yr18 in Spring Wheat and its Effect on Components of Partial Resistance

The non‐durable nature of hypersensitive (race‐specific) resistance has stimulated scientists to search for other options such as race‐non‐specific resistance to provide long‐lasting protection against plant diseases. Adult plant resistance gene complex Lr34/Yr18 confers a dual race‐non‐specific type of resistance to wheat against stripe rust (Puccinia striiformis f. sp. tritici) and leaf rust (P. triticina Eriks). This study was conducted to evaluate 59 spring bread wheat (Triticum aestivum L.) genotypes for the presence of the Lr34/Yr18‐linked csLV34 allele using STS marker csLV34 and to determine the effect of this gene complex on the components of partial resistance in wheat to leaf/stripe rust. Lr34/Yr18‐linked csLV34 allele was detected only in 12 genotypes, namely Iqbal 2000, NR‐281, NR 354, NR 363, NR 364, NR 366, NR 367, NR 370, NR 376, 4^thEBWYT 509, 4^thEBWYT 510 and 4^thEBWYT 518. Eleven genotypes showing the amplified Lr34/Yr18‐linked allele were further studied for the assessment of the effect of Lr34/Yr18 on components of partial resistance along with nine genotypes lacking this gene complex. Both stripe and leaf rusts were studied separately. The components of partial resistance including latency period (LP) and infection frequency (IF) were studied on primary leaf (seedling stage), fourth leaf and fully expanded young flag leaf (adult plant stage). Both the stripe and leaf rust fungi showed a prolonged LP and reduced IF on genotypes carrying Lr34/Yr18 gene complex. Generally, a longer LP was associated with a reduced IF at all growth stages. Although significant effect of Lr34/Yr18 gene complex on LP and IF was observed almost at all three growth stages, the effect was more pronounced at flag leaf. This suggested that Lr34/Yr18 gene complex is more effective at later stages of plant growth.     

Diallel analysis of the latent period of stripe rust in wheat

A half diallel was made amongst five wheat (Triticum aestivum L.) genotypes of which one was susceptible, while the others had adult-plant resistance, to stripe rust (Puccinia striiformis West.). The five parent and ten F₁ progeny were grown in the glasshouse and were inoculated with three rust pathotypes at the seedling stage. The latent period was measured on the first leaf. Two procedures were used to analyze the half diallel. Both methods showed that the average effects of alleles were of much greater importance than was dominance in conditioning resistance in response to two of the pathotypes, while for the third pathotype dominance was important. Resistance was conditioned by partial dominance for two pathotypes whereas for the third it was determined by full dominance. Broad-sense heritabilities range from 60–73% and the number of genes involved was different (from 1 to 4), depending on the pathotype.     

QTL mapping of resistance to leaf rust in Salix

Genetic mapping of quantitative trait loci (QTL) for resistance to Melampsora leaf rust was performed in two willow families: the progeny from a backcross between Salix viminalis and a hybrid S. viminalis × Salix schwerinii (population S₁), and the F₁ progeny of a cross between two S. viminalis (population S₃). Disease levels were scored in the field for three consecutive years. In the laboratory, five different rust strains were sprayed onto leaf disks and the following resistance components were scored: latent period, diameter and number of uredinia, and flecking. One major QTL and 14 smaller were identified in the S₁ host population. One rust strain, that represents a Melampsora form with limited incidence on S. viminalis, showed lower aggressiveness to the S₁ host population together with a different pattern in QTLs. In the S₃ host population, we detected 13 QTLs for rust resistance, of which two were located to the same genomic regions as those found for the S₁ population. We showed that the strongest QTL co-segregated with a gene homologous to a known Toll/interleukin receptor-nucleotide binding site-leucine-rich repeat resistance gene in poplar. The importance of the identified QTLs is discussed in relation to breeding for durable resistance.