Association mapping of spot blotch resistance in wild barley

Spot blotch, caused by Cochliobolus sativus, is an important foliar disease of barley. The disease has been controlled for over 40 years through the deployment of cultivars with durable resistance derived from the line NDB112. Pathotypes of C. sativus with virulence for the NDB112 resistance have been detected in Canada; thus, many commercial cultivars are vulnerable to spot blotch epidemics. To increase the diversity of spot blotch resistance in cultivated barley, we evaluated 318 diverse wild barley accessions comprising the Wild Barley Diversity Collection (WBDC) for reaction to C. sativus at the seedling stage and utilized an association mapping (AM) approach to identify and map resistance loci. A high frequency of resistance was found in the WBDC as 95% (302/318) of the accessions exhibited low infection responses. The WBDC was genotyped with 558 Diversity Array Technology (DArT^®) and 2,878 single nucleotide polymorphism (SNP) markers and subjected to structure analysis before running the AM procedure. Thirteen QTL for spot blotch resistance were identified with DArT and SNP markers. These QTL were found on chromosomes 1H, 2H, 3H, 5H, and 7H and explained from 2.3 to 3.9% of the phenotypic variance. Nearly half of the identified QTL mapped to chromosome bins where spot blotch resistance loci were previously reported, offering some validation for the AM approach. The other QTL mapped to unique genomic regions and may represent new spot blotch resistance loci. This study demonstrates that AM is an effective technique for identifying and mapping QTL for disease resistance in a wild crop progenitor.     

Genome-wide association mapping of spot blotch resistance to three different pathotypes of <Emphasis Type="Italic">Cochliobolus sativus</Emphasis> in the USDA barley core collection

Spot blotch, caused by Cochliobolus sativus, is an economically important disease of barley. To identify genetic loci conferring resistance to three different pathotypes of C. sativus, a worldwide barley core collection (BCC) consisting of 1480 accessions from the USDA National Small Grains Collection were genotyped with the barley 9k Illumina Infinium iSELECT assay and phenotyped at the seedling stage with three C. sativus isolates ND85F (pathotype 1), ND90Pr (pathotype 2), and ND4008 (pathotype 7). Association mapping analysis was performed with the Whole_Panel containing 1480 barley accessions, as well as Two-rowed_Panel and Six-rowed_Panel consisting of 621 two-rowed and 857 six-rowed barley accessions, respectively. For resistance to isolate ND4008, one quantitative trait locus (QTL, QRcs-6H-P7) was detected in all three panels. Three other QTL (QRcs-1H-P7, QRcs-2H-P7, and QRcs-3H-P7) were detected in Whole_Panel, Six-rowed_Panel, and Two-rowed_Panel, respectively. For resistance to isolate ND90Pr, one QTL (QRcs-1H-P2) was identified in the Whole_Panel and the Two-rowed_Panel, and the other QTL (QRcs-6H-P2) was only identified in the Six-rowed_Panel. For resistance to isolate ND85F, three QTL (QRcs-1H-P1, QRcs-3H-P1, QRcs-7H-2-P1) were detected in all three panels, and one QTL (QRcs-7H-1-P1) was only detected in the Two-rowed_Panel. Among the ten QTL detected, four (QRcs-1H-P1, QRcs-3H-P1, QRcs-7H-2-P1, and QRcs-1H-P2) were mapped to chromosome regions containing previously identified QTL for spot blotch resistance, while six (QRcs-1H-P7, QRcs-2H-P7, QRcs-3H-P7, QRcs-6H-P7, QRcs-6H-P2, and QRcs-7H-1-P1) were novel. The SNP markers associated with the QTL identified in this study will be useful for breeding barley cultivars with resistance to multiple pathotypes of C. sativus.     

Genetics of seedling and adult plant resistance to net blotch (Pyrenophora teres f. teres) and spot blotch (Cochliobolus sativus) in barley

Net blotch (caused by Pyrenophora teres f. teres) and spot blotch (Cochliobolus sativus) are important foliar diseases of barley in the midwestern region of the USA. To determine the number and chromosomal location of Mendelian and quantitative trait loci (QTL) controlling resistance to these diseases, a doubled haploid population (Steptoe/Morex) was evaluated to the pathogens at the seedling stage in the greenhouse and at the adult plant stage in the field. Alleles at two or three unlinked loci were found to confer resistance to the net blotch pathogen at the seedling stage depending on how progeny exhibiting an intermediate infection response were classified. This result was corroborated in the quantitative analysis of the raw infection response data as 2 major QTL were identified on chromosomes 4 and 6M. A third QTL was also identified on chromosome 6P. Seven QTL were identified for net blotch resistance at the adult plant stage and mapped to chromosomes 1P, 2P, 3P, 3M, 4, 6P, and 7P. The 7 QTL collectively accounted for 67.6% of the phenotypic variance under a multiple QTL model. Resistance to the spot blotch pathogen was conferred by a single gene at the seedling stage. This gene was mapped to the distal region of chromosome 1P on the basis of both qualitative and quantitative data analyses. Two QTL were identified for spot blotch resistance at the adult plant stage: the largest QTL effect mapped to chromosome 5P and the other mapped to chromosome 1P near the seedling resistance locus. Together, the 2 QTL explained 70.1% of the phenotypic variance under a multiple QTL model. On the basis of the chromosomal locations of resistance alleles detected in this study, it should be feasible to combine high levels of resistance to both P. teres f. teres and C. sativus in barley cultivars.     

Comprehensive genetic analyses reveal differential expression of spot blotch resistance in four populations of barley

Spot blotch, caused by Cochliobolus sativus, is an important disease of barley in the Upper Midwest region of the United States. The resistance of six-rowed malting cultivars like Morex has remained effective for over 40 years and is considered durable. Previous research on Steptoe/Morex (S/M), a 6×6-rowed doubled haploid (DH) population, showed that seedling resistance is controlled by a single gene (Rcs5) on chromosome 1(7H) and adult plant resistance by two quantitative trait loci (QTL): one of the major effect on chromosome 5(1H) explaining 62% of the phenotypic variance and a second of minor effect on chromosome 1(7H) explaining 9% of the phenotypic variance. To corroborate these results in a 2×6-rowed DH population, composite interval mapping (CIM) was performed on Harrington/Morex (H/M). As in the S/M population, a single major gene (presumably Rcs5) on chromosome 1(7H) conferred resistance at the seedling stage. However, at the adult plant stage, the results were markedly different as no chromosome 5(1H) effect whatsoever was detected. Instead, a QTL at or near Rcs5 on chromosome 1(7H) explained nearly all of the phenotypic variance (75%) for disease severity. To determine whether this result might be due to the genetic background of the two-rowed susceptible parent Harrington, we analyzed another DH population that included the same resistance donor (Morex) and another six-rowed susceptible cultivar Dicktoo (D/M). Three QTL conferred seedling resistance in the D/M population: one near Rcs5 on chromosome 1(7H) explaining 30%, a second near the centromere of chromosome 1(7H) explaining 9%, and a third on the short arm of chromosome 3(3H) explaining 19% of the phenotypic variation. As in the H/M population, no chromosome 5(1H) QTL was detected for adult plant resistance in the D/M population. Instead, three QTL on other chromosomes explained most of the variation: one on the short arm of chromosome 3(3H) explaining 36%, a second on the long arm of chromosome 3(3H) explaining 11%, and a third at or near Rcs5 on chromosome 1(7H) explaining 20% of the phenotypic variation. These data demonstrate the complexity of expression of spot blotch resistance in different populations and have important implications in breeding for durable resistance.     

Mapping quantitative trait loci associated with spot blotch and net blotch resistance in a doubled-haploid barley population

Spot blotch and net blotch are important foliar barley (Hordeum vulgare L.) diseases in Canada and elsewhere. These diseases result in significant yield reduction and, more importantly, loss of grain quality, downgrading barley from malt to feed. Combining resistance to these diseases is a breeding priority but is a significant challenge using conventional breeding methodology. In the present investigation, an evaluation of the inheritance of resistance to spot and net blotch was conducted in a doubled-haploid barley population from the cross CDC Bold (susceptible)?×?TR251 (resistant). The population was screened at the seedling stage in the Phytotron and at the adult-plant stage in the field for several years. Chi-squared analysis indicated one- to four-gene segregation depending on disease, isolate, plant development stage, location and year. A major seedling and adult-plant resistance quantitative trait locus (QTL), designated QRpt6, was re-confirmed for net-form net blotch resistance, explaining 32?C61% of phenotypic variation in different experiments. Additional QTL for seedling and adult-plant resistance to net blotch were identified. For spot blotch resistance, a major seedling resistance QTL (QRcss1) was detected on chromosome 1H for isolate WRS1909, explaining 79% of the phenotypic variation. A highly significant QTL on 3H (QRcs3) was identified for seedling resistance to isolate WRS1908 and adult-plant resistance at Brandon, MB, Canada in 2008. The identification of QTL at only one location or from 1?year suggests spot blotch resistance is complex and highly influenced by the environment. Efforts are being made to combine spot and net blotch resistance in elite barley lines using molecular marker-assisted selection.     

Quantitative trait loci for resistance to spot blotch caused by Bipolarissorokiniana in wheat (T.aestivum L.) lines ‘Ning 8201’ and ‘Chirya 3’

Spot blotch caused by Bipolaris sorokiniana is a destructive disease of wheat in warm and humid wheat growing regions of the world. To identify quantitative trait loci (QTLs) for spot blotch resistance, two mapping populations were developed by making the crosses between common susceptible cultivar ‘Sonalika’ with the resistant breeding lines ‘Ning 8201’ and ‘Chirya 3’. Single seed descent derived F₆, F₇, F₈ lines of the first cross ‘Ning 8201’ × ‘Sonalika’ were evaluated for resistance to spot blotch in three blocks in each of the 3 years. After screening of 388 pairs of simple sequence repeat primers between the two parents, 119 polymorphic markers were used to genotype the mapping population. Four QTLs were identified on the chromosomes 2AS, 2BS, 5BL and 7DS and explained 62.9% of phenotypic variation in a simultaneous fit. The QTL on chromosome 2A was detected only in 1 year and explained 22.7% of phenotypic variation. In the second cross (‘Chirya 3’ × ‘Sonalika’), F₇ and F₈ population were evaluated in three blocks in each of the 2 years. In this population, five QTLs were identified on chromosomes 2BS, 2DS, 3BS, 7BS and 7DS. The QTLs identified in the ‘Chirya 3’ × ‘Sonalika’ population explained 43.4% of phenotypic variation in a simultaneous fit. The alleles for reduced disease severity in both the populations were derived from the respective resistant parent. The QTLs QSb.bhu-2B and QSb.bhu-7D from both populations were placed in the same deletion bins, 2BS1-0.53-0.75 and 7DS5-0.36-0.61, respectively. The closely linked markers Xgwm148 to the QTL on chromosome 2B and Xgwm111 to the QTL on chromosome 7D are potentially diagnostic markers for spot blotch resistance.     

Genome-wide association mapping reveals genetic architecture of durable spot blotch resistance in US barley breeding germplasm

Spot blotch, an economically important disease of both barley and wheat, is caused by Cochliobolus sativus (anamorph: Bipolaris sorokiniana). The disease has been reported in many regions of the world, but is particularly severe on barley in the Upper Midwest region of the USA and adjacent areas of Canada. For over 50 years, spot blotch has been effectively controlled through the deployment of durable resistance in six-rowed malting cultivars. To characterize loci conferring spot blotch resistance in US barley germplasm, we employed an association mapping approach using 3,840 breeding lines and cultivars. Three quantitative trait loci (QTL), Rcs-qtl-1H-11_10764, Rcs-qtl-3H-11_10565 and Rcs-qtl-7H-11_20162, were found to confer both seedling and adult plant resistance. Together, these three QTL comprise the Midwest Six-rowed Durable Resistant Haplotype (MSDRH), which is present in all Midwest six-rowed cultivars released since the 1960s. Each QTL alone only partially reduced disease levels, but combining all three together reduced the seedling infection response and adult plant disease severity by 47 and 83 %, respectively. The identified MSDRH will be valuable for marker-assisted selection of breeding lines to deploy spot blotch resistance and can also be incorporated into genomic selection as one of the disease resistance traits.     

Mapping of resistance to spot blotch disease caused by Bipolaris sorokiniana in spring wheat

Spot blotch caused by Bipolaris sorokiniana is a destructive disease of wheat in warm and humid wheat growing regions of the world. The development of disease resistant cultivars is considered as the most effective control strategy for spot blotch. An intervarietal mapping population in the form of recombinant inbred lines (RILs) was developed from a cross ‘Yangmai 6’ (a Chinese source of resistance) × ‘Sonalika’ (a spot blotch susceptible cultivar). The 139 single seed descent (SSD) derived F₆, F₇, F₈ lines of ‘Yangmai 6’ × ‘Sonalika’ were evaluated for resistance to spot blotch in three blocks in each of the 3 years. Joint and/or single year analysis by composite interval mapping (CIM) and likelihood of odd ratio (LOD) >2.2, identified four quantitative trait loci (QTL) on the chromosomes 2AL, 2BS, 5BL and 6DL. These QTLs were designated as QSb.bhu-2A, QSb.bhu-2B, QSb.bhu-5B and QSb.bhu-6D, respectively. A total of 63.10% of phenotypic variation was explained by these QTLs based on the mean over years. Two QTLs on chromosomes 2B and 5B with major effects were consistent over 3 years. All QTL alleles for resistance were derived from the resistant parent ‘Yangmai 6’.     

Mapping Rph20: a gene conferring adult plant resistance to Puccinia hordei in barley

A doubled haploid (DH) barley (Hordeum vulgare L.) population of 334 lines (ND24260?×?Flagship) genotyped with DArT markers was used to map genes for adult plant resistance (APR) to leaf rust (Puccinia hordei Otth) under field conditions in Australia and Uruguay. The Australian barley cultivar Flagship carries an APR gene (qRphFlag) derived from the cultivar Vada. Association analysis and composite interval mapping identified two genes conferring APR in this DH population. qRphFlag was mapped to the short arm of chromosome 5H (5HS), accounting for 64?C85% of the phenotypic variation across four field environments and 56% under controlled environmental conditions (CEC). A second quantitative trait locus (QTL) from ND24260 (qRphND) with smaller effect was mapped to chromosome 6HL. In the absence of qRphFlag, qRphND conferred only a low level of resistance. DH lines displaying the highest level of APR carried both genes. Sequence information for the critical DArT marker bPb-0837 (positioned at 21.2?cM on chromosome 5HS) was used to develop bPb-0837-PCR, a simple PCR-based marker for qRphFlag. The 245?bp fragment for bPb-0837-PCR was detected in a range of barley cultivars known to possess APR, which was consistent with previous tests of allelism, demonstrating that the qRphFlag resistant allele is common in leaf rust resistant cultivars derived from Vada and Emir. qRphFlag has been designated Rph20, the first gene conferring APR to P. hordei to be characterised in barley. The PCR marker will likely be effective in marker-assisted selection for Rph20.     

Adult plant and seedling resistance to powdery mildew in a Triticum aestivum × Triticum militinae hybrid line

In the progeny of a cross between the common wheat cultivar Tähti and Triticum militinae, a member of the timopheevii group of tetraploid wheats, several hybrid lines were selected that are characterized by improved seedling and adult plant resistance (APR) to powdery mildew. An F₂ single-seed descendant mapping population segregating for seedling resistance and APR to powdery mildew was analysed for the identification of quantitative trait loci (QTL). The main QTL responsible for APR was detected on the long arm of chromosome 4A tightly linked to the Xgwm160 locus on a T. militinae translocation explaining up to 54% of phenotypic variance. The same translocation influenced seedling resistance to powdery mildew upon inoculation of plants with a synthetic population of Blumeria graminis DC. f. sp. tritici, and explained 28–33% of the phenotypic variance.     

Inheritance of field resistance to Stagonospora nodorum leaf and glume blotch and correlations with other morphological traits in hexaploid wheat (Triticum aestivum L.)

Breeding for wheat varieties resistant to Stagonospora nodorum blotch (SNB) is the most sustainable strategy for controlling the disease. In order to map quantitative trait loci (QTLs) for SNB resistance we analysed 204 recombinant inbred lines of the cross between the winter wheat (Triticum aestivum L.) variety Forno and the winter spelt (Triticum spelta L.) variety Oberkulmer. We determined the level of resistance of adult plants to leaf blotch (SNL) and glume blotch (SNG) as well as morphological traits for 2 years after artificial inoculation with S. nodorum. Using composite interval mapping and LOD > 3.7, we detected ten QTLs for SNG blotch resistance (six inherited from the susceptible parent Forno) and 11 QTLs for SNL resistance (four inherited from Forno) across 2 years. Both resistance traits were moderately correlated (r = 0.52) and had only one common QTL. For SNL resistance, seven QTLs were not associated with QTLs for morphological traits. Among them, QSnl.eth-2D, QSnl.eth-4B and QSnl.eth-7B3 had major effects (R(2) > 13%) and were potential candidates for marker-assisted selection. For SNG, the major QTL on chromosome 5A, explaining 36% of the phenotypic variance for resistance, was associated with the q locus conferring the spelt morphology (long lax ear, long culm and hard glumes). Only QSng.eth-1BS, which explained 7% of the variance for resistance to SNG blotch, was not associated with QTLs for morphological traits. The consequences for breeding programmes are discussed.     

QTL dissection of the loss of green colour during post-anthesis grain maturation in two-rowed barley

Ability to genetically manipulate the loss of green colour during grain maturation has potentials for increasing productivity, disease resistance, and drought and heat tolerance in crop plants. Two doubled haploid, two-rowed barley populations (Vlamingh × Buloke and VB9524 × ND11231*12) were monitored over 2 years for loss of green colour during grain filling using a portable active sensor. The aims were to determine the genomic regions that control trait heritability by quantitative trait locus (QTL) analysis, and to examine patterns of QTL-environment interactions under different conditions of water stress. In the Vlamingh × Buloke cross, broad-sense heritability estimate for loss of green colour (measured as the difference in sensor readings taken at anthesis and maturity, ?SRI) was 0.68, and 0.78 for the VB9524 × ND11231*12 population. In the VB9524 × ND11231*12 population, rapid loss of green colour was positively associated with grain yield and percent plump grains, but in the Vlamingh × Buloke population, a slower loss of green colour (low ?SRI) was associated with increased grain plumpness. With the aid of a dense array of single nucleotide polymorphisms (SNPs) and EST-derived SSR markers, a total of nine QTLs were detected across the two populations. Of these, a single major locus on the short arm of barley chromosome 5H was consistently linked with trait variation across the populations and multiple environments. The QTL was independent of flowering time and explained between 5.4 and 15.4 % of the variation observed in both populations, depending on the environment, and although a QTL × E interaction was detected, it was largely due to a change in the magnitude of the effect, rather than a change in direction. The results suggest that loss of green colour during grain maturation may be under the control of a simple genetic architecture, but a careful study of target populations and environments would be required for breeding purposes.     

Identification and mapping of net form of net blotch resistance in South African barley

Net form of net blotch (NFNB) caused by the fungus Pyrenophora teres f. teres is an economically important foliar disease of barley (Hordeum vulgare) in southern and eastern Africa. Little attention has been given to disease resistance breeding, and knowledge about the presence of NFNB resistance in breeding lines is limited. Deploying resistance into varieties used in this region is important for future control of the disease. We have identified NFNB disease resistance in existing South African breeders’ lines and have mapped the resistance in line UVC8. Six different trials, three conducted in South Africa and another three in Australia, were used to identify resistance QTL. A major QTL was identified on chromosome 6H having a LOD score of 40.5 and 55% of the phenotypic variance explained. Kompetitive Allele Specific PCR (KASP?) markers were designed for this QTL region. These and microsatellite markers can now be used to routinely select for NFNB resistance.     

Mapping quantitative trait loci associated with barley net blotch resistance

Net blotch of barley, caused by Pyrenophora teres Drechs., is an important foliar disease worldwide. Deployment of resistant cultivars is the most economic and eco-friendly control method. This report describes mapping of quantitative trait loci (QTL) associated with net blotch resistance in a doubled-haploid (DH) barley population using diversity arrays technology (DArT) markers. One hundred and fifty DH lines from the cross CDC Dolly (susceptible)/TR251 (resistant) were screened as seedlings in controlled environments with net-form net blotch (NFNB) isolates WRS858 and WRS1607 and spot-form net blotch (SFNB) isolate WRS857. The population was also screened at the adult-plant stage for NFNB resistance in the field in 2005 and 2006. A high-density genetic linkage map of 90 DH lines was constructed using 457 DArT and 11 SSR markers. A major NFNB seedling resistance QTL, designated QRpt6, was mapped to chromosome 6H for isolates WRS858 and WRS1607. QRpt6 was associated with adult-plant resistance in the 2005 and 2006 field trials. Additional QTL for NFNB seedling resistance to the more virulent isolate WRS858 were identified on chromosomes 2H, 4H, and 5H. A seedling resistance QTL (QRpts4) for the SFNB isolate WRS857 was detected on chromosome 4H as was a significant QTL (QRpt7) on chromosome 7H. Three QTL (QRpt6, QRpts4, QRpt7) were associated with resistance to both net blotch forms and lines with one or more of these demonstrated improved resistance. Simple sequence repeat (SSR) markers tightly linked to QRpt6 and QRpts4 were identified and validated in an unrelated barley population. The major 6H QTL, QRpt6, may provide adequate NFNB field resistance in western Canada and could be routinely selected for using molecular markers in a practical breeding program.     

QTL for spot blotch resistance in bread wheat line Saar co-locate to the biotrophic disease resistance loci Lr34 and Lr46

Spot blotch caused by Bipolaris sorokiniana is a major disease of wheat in warm and humid wheat growing regions of the world including south Asian countries such as India, Nepal and Bangladesh. The CIMMYT bread wheat line Saar which carries the leaf tip necrosis (LTN)-associated rust resistance genes Lr34 and Lr46 has exhibited a low level of spot blotch disease in field trials conducted in Asia and South America. One hundred and fourteen recombinant inbred lines (RILs) of Avocet (Susceptible) × Saar, were evaluated along with parents in two dates of sowing in India for 3 years (2007–2008 to 2009–2010) to identify quantitative trait loci (QTL) associated with spot blotch resistance, and to determine the potential association of Lr34 and Lr46 with resistance to this disease. Lr34 was found to constitute the main locus for spot blotch resistance, and explained as much as 55 % of the phenotypic variation in the mean disease data across the six environments. Based on the large effect, the spot blotch resistance at this locus has been given the gene designation Sb1. Two further, minor QTL were detected in the sub-population of RILs not containing Lr34. The first of these was located about 40 cM distal to Lr34 on 7DS, and the other corresponded to Lr46 on 1BL. A major implication for wheat breeding is that Lr34 and Lr46, which are widely used in wheat breeding to improve resistance to rust diseases and powdery mildew, also have a beneficial effect on spot blotch.     

Phenotyping at hot spots and tagging of QTLs conferring spot blotch resistance in bread wheat

Spot blotch is a major foliar disease of wheat caused by Bipolaris sorokiniana in warm and humid environments of the world including South Asian countries. In India, it has a larger impact in Indo-Gangetic plains of the country. Therefore, the present study was undertaken to phenotype a mapping population at different hot spots of India and to detect quantitative trait loci (QTL) for resistance to spot blotch in wheat. For this study, 209 single seed descent (SSD) derived F₈, F₉, F₁₀ recombinant inbred lines (RILs) of the cross ‘Sonalika’ (an Indian susceptible cultivar)/‘BH 1146’ (a Brazilian resistant cultivar) were assessed for spot blotch resistance at two hot spot locations (Coochbehar and Kalyani) for three years and for two years under controlled conditions in the polyhouse (Karnal). The population showed large variation in spot blotch reaction for disease severity in all the environments indicating polygenic nature of the disease. Microsatellite markers were used to create the linkage maps. Joint and/or individual year analysis by composite interval mapping (CIM) and likelihood of odds ratio (LOD) >2.1, detected two consistent QTLs mapped on chromosome 7BL and 7DL and these explained phenotypic variation of 11.4 percent and 9.5 percent over the years and locations, respectively. The resistance at these loci was contributed by the parent ‘BH 1146’ and shown to be independent of plant height and earliness. Besides, association of some agro-morphological traits has also been observed with percent disease severity. These identified genomic regions may be used in future wheat breeding programs through marker assisted selection for developing spot blotch resistant cultivars.     

Genomewide analysis of epistatic effects for quantitative traits in barley

The doubled-haploid (DH) barley population (Harrington x TR306) developed by the North American Barley Genome Mapping Project (NABGMP) for QTL mapping consisted of 145 lines and 127 markers covering a total genome length of 1270 cM. These DH lines were evaluated in approximately 25 environments for seven quantitative traits: heading, height, kernel weight, lodging, maturity, test weight, and yield. We applied an empirical Bayes method that simultaneously estimates 127 main effects for all markers and 127(127-1)/2=8001 interaction effects for all marker pairs in a single model. We found that the largest main-effect QTL (single marker) and the largest epistatic effect (single pair of markers) explained approximately 18 and 2.6% of the phenotypic variance, respectively. On average, the sum of all significant main effects and the sum of all significant epistatic effects contributed 35 and 6% of the total phenotypic variance, respectively. Epistasis seems to be negligible for all the seven traits. We also found that whether two loci interact does not depend on whether or not the loci have individual main effects. This invalidates the common practice of epistatic analysis in which epistatic effects are estimated only for pairs of loci of which both have main effects.     

Two stable QTL involved in adult plant resistance to powdery mildew in the winter wheat line RE714 are expressed at different times along the growing season

Powdery mildew (Blumeria graminis f. sp. tritici) is one of the major diseases of wheat (Triticum aestivum). Adult plant resistance (APR) to powdery mildew is considered more durable than resistance conferred by major race-specific resistance genes. The objective of the present study was a better understanding of the genetic basis of APR in RE714 by means of QTL analysis of several resistance scores along the growing season. A population of 160 recombinant inbred lines obtained from the cross between RE714 and Hardi (susceptible) was assessed for APR under natural infection conditions during 3 years and a genetic map with whole genome coverage was developed with microsatellite and AFLP markers in this population. Two major QTL on chromosomes 5D and 6A were detected each year, and 6 minor QTL were detected only in 1 or 2 years. The QTL on chromosome 5D was detected during all the growing season each year and its R ² value varied between 8.5 and 56.3%, whereas the QTL on chromosome 6A was detected at 1–4 scoring dates in the 3 years, and its R ² value varied between 6.1 and 20.5%. The two QTL explained between 24.4 and 52.1% of the phenotypic variance for AUDPC, depending on the year. The models including QTL and cofactors in the composite interval mapping explained between 29 and 72% of the variance. The molecular markers linked to the two major QTL could be used in marker-assisted selection for adult plant resistance to powdery mildew. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Inheritance and localisation of resistance to <Emphasis Type="Italic">Mycosphaerella graminicola</Emphasis> causing septoria tritici blotch and plant height in the wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) genome with DNA markers

Resistance to the disease septoria tritici blotch of wheat (Triticum aestivum L.), caused by the fungus Mycosphaerella graminicola (Fuckel.) J. Schrot in Cohn (anamorph Septoria tritici Roberge in Desmaz.) was investigated in a doubled-haploid (DH) population of a cross between the susceptible winter wheat cultivar Savannah and the resistant cultivar Senat. A molecular linkage map of the population was constructed including 76 SSR loci and 244 AFLP loci. Parents and DH progeny were tested for resistance to single isolates of M. graminicola in a growth chamber at the seedling stage, and to an isolate mixture at the adult plant stage, in field trials. A gene located at or near the Stb6 locus mapping to chromosome 3A provided seedling resistance to IPO323. Two complementary genes, mapping to chromosome 3A, one of which was the IPO323 resistance gene, were needed for resistance to the Danish isolate Ris?97-86. In addition, a number of minor loci influenced the expression of resistance in the growth chamber. In the field, four QTLs for resistance to septoria tritici blotch were detected. Two QTLs, located on chromosomes 3A and 6B explained 18.2 and 67.9% of the phenotypic variance in the mean over two trials. Both these QTLs were also detected at the seedling stage with isolate Ris?97-86, whereas isolate IPO323 only detected the QTL on 3A. Additionally, two QTLs identified in adult plants on chromosomes 2B and 7B were not detected at the seedling stage. Four QTLs were detected for plant height located on chromosomes 2B, 3A, 3B and on a linkage group not assigned to a chromosome. The major QTLs on 3A and on the unassigned linkage group were consistent over two trials, and the QTL on 3A seemed to be linked to a QTL for septoria tritici blotch resistance.     

Detection of QTLs for Stagonospora glume blotch resistance in Swiss winter wheat

Stagonospora nodorum is the causal agent of the Stagonospora glume blotch disease in hexaploid wheat. The Swiss winter bread wheat cv. 'Arina' has a highly effective, durable and quantitative glume blotch resistance. We studied 240 single seed descent (SSD)-derived lines of an 'Arina × Forno' F_5:7 population to identify and map quantitative trait loci (QTLs) for glume blotch resistance under natural infestation. Using composite interval mapping (CIM) and LOD>4.5, we detected two chromosomal regions on chromosome arms 3BS and 4BL which were specifically associated with glume blotch resistance. These identified QTLs were designated QSng.sfr-3BS and QSng.sfr-4BL, respectively. QSng.sfr-3BS peaked at the locus Xgwm389 in the telomeric region of the short arm of chromosome 3B and explained 31.2% of the observed phenotypic variance for the resistance within the population. The responsible QSng.sfr-3BS allele originated from the resistant parent 'Arina'. The QTL QSng.sfr-4BL (19.1%) mapped to chromosome arm 4BL ('Forno' allele) very close to two known genes, TaMlo and a catalase (Cat). Both QTL alleles combined could enhance the resistance level by about 50%. Additionally, they showed significant epistatic effects (4.4%). We found PCR-based microsatellite markers closely linked to QSng.sfr-3BS (gwm389) and QSng.sfr-4BL (gwm251) which make marker-assisted selection (MAS) for Stagonospora glume blotch resistance feasible. We also found one resistance QTL, QSng.sfr-5BL, on the long arm of chromosome 5B which overlapped with QTLs for plant height as well as heading time.Communicated by H. C. Becker