Genome-wide association mapping of genetic factors controlling Septoria tritici blotch resistance and their associations with plant height and heading date in wheat

Septoria tritici blotch (STB), caused by the ascomycete fungus Zymoseptoria tritici (also known as Mycosphaerella graminicola), is one of the most devastating foliar wheat diseases worldwide. Host resistance is the most effective strategy for management of the disease. A factor that complicates the determination of resistance is its reported interaction with heading date (Hd) and plant height (Ph). In this study, we report findings from a genome-wide association study of resistance to STB in a world-wide collection of 96 wheat accessions. The collection was evaluated under conditions of artificial infection for seedling and adult plant STB resistance, Hd and Ph in field trials. Marker-trait associations (MTAs) were detected using a mixed linear model. STB disease severities showed significant phenotypic variation. In total, 73 MTAs involving STB resistance were detected. The chromosomal locations of some of them were similar to known Stb genes or quantitative trait loci; whereas others were detected in new genomic regions. The field experiment showed evidence of genetic association between STB resistance and Hd, but only for a few genotypes. This was corroborated at the molecular level, where a total of eight genomic regions associated with STB resistance were located in similar positions to MTAs for Hd. New genomic regions associated with STB resistance found here could be useful in wheat breeding aimed at controlling STB after validation in relevant genetic backgrounds     

The potential of genomic‐assisted breeding to improve Fusarium head blight resistance in winter durum wheat

Durum wheat is the most important tetraploid wheat mainly used for semolina and pasta production, but is notorious for its high susceptibility to Fusarium head blight (FHB). Our objectives were to identify and characterize quantitative trait loci (QTL) in winter durum and to evaluate the potential of genomic approaches for the improvement of FHB resistance. Here, we employed an international panel of 170 winter and 14 spring durum lines, phenotyped for Fusarium culmorum resistance at five environments. Heading date, plant height and mean FHB severity showed significant genotypic variation with high heritabilities and FHB resistance was negatively correlated with both heading date and plant height. The dwarfing gene Rht‐B1 significantly affected FHB resistance and the genome‐wide association scan identified eight additional QTL affecting FHB resistance, explaining between 1% and 14% of the genotypic variation. A genome‐wide prediction approach yielded only a slightly improved predictive ability compared to marker‐assisted selection based on the four strongest QTL. In conclusion, FHB resistance in durum wheat is a highly quantitative trait and in breeding programmes may best be tackled by classical high‐throughput recurrent phenotypic selection that can be assisted by genomic prediction if marker profiles are available.     

Alleles on the two dwarfing loci on 4B and 4D are main drivers of FHB‐related traits in the Canadian winter wheat population “Vienna” × “25R47”

Fusarium head blight (FHB), leaf rust and stem rust are among the most destructive wheat diseases. High‐yielding, native disease resistance sources are available in North America. The objective of this study was to map loci associated with FHB traits, leaf rust, stem rust and plant height in a “Vienna”/”25R47” population. DArT markers were used to generate a genetic map, and quantitative trait loci (QTL) analysis was performed by evaluating 113 doubled haploid lines across three environments in Ontario, Canada. FHB resistance QTL were identified on chromosomes 4D, 4B, 2D and 7A, while a QTL for leaf and stem rust resistance was identified on chromosome 1B. The dwarfing alleles of both Rht‐B1 and Rht‐D1 were associated with increased FHB index and DON content.     

Molecular mapping of quantitative trait loci for field resistance to Fusarium head blight in a European winter wheat population

Fusarium head blight (FHB), one of the most destructive diseases of wheat in many parts of the world, can reduce the grain quality due to mycotoxin contamination up to rejection for usage as food or feed. Objective of this study was to map quantitative trait loci (QTL) associated with FHB resistance in the winter wheat population ‘G16‐92’ (resistant)/‘Hussar’. In all, 136 recombinant inbred lines were evaluated in field trials in 2001 and 2002 after spray inoculation with a Fusarium culmorum suspension. The area under disease progress curve was calculated based on the visually scored FHB symptoms. For means across all environments two FHB resistance QTL located on chromosomes 1A, and 2BL were identified. The individual QTL explained 9.7% and 14.1% of the phenotypic variance and together 26.7% of the genetic variance. The resistance QTL on 1A coincided with a QTL for plant height in contrast to the resistance QTL on 2BL that appeared to be independently inherited from morphological characteristics like plant height and ear compactness. Therefore, especially the QTL on 2BL could be of great interest for breeding towards FHB resistance.     

QTL mapping and marker-assisted selection for Fusarium head blight resistance in wheat: a review

During the past decade, numerous studies have been published on molecular mapping of Fusarium head blight (FHB) resistance in wheat. We summarize the relevant findings from 52 quantitative trait loci (QTL) mapping studies, nine research articles on marker-assisted selection and seven on marker-assisted germplasm evaluation. QTL for FHB resistance were found on all wheat chromosomes except chromosome 7D. Some QTL were found in several independent mapping studies indicating that such QTL are stable and therefore useful in breeding programmes. We summarize and update current knowledge on the genetics of FHB resistance in wheat resulting from QTL mapping investigations and review and suggest FHB breeding strategies based on the available information and DNA markers.     

Short Communication Comparison between QTL analysis of powdery mildew resistance in barley based on detached primary leaves and on field data

A quantitative trait loci (QTL) analysis of quantitative powdery mildew resistance was performed on 216 doubled haploid lines derived from a cross between the winter-barley varieties ‘Igri’ and ‘Danilo’ using 67 RFLP loci. Resistance to powdery mildew was determined in the field with natural infection and on detached primary leaves with a specific isolate. The major QTL found in both sets of analysis mapped to the same chromosomal region. No further QTL could be found in the analysis based on detached leaves and one additional minor QTL was found in the analysis based on field data.     

Identification of novel QTL for resistance to crown rot in the doubled haploid wheat population 'W21MMT70' × 'Mendos'

Crown rot (causal agent Fusarium pseudograminearum) is a fungal disease of major significance to wheat cultivation in Australia. A doubled haploid wheat population was produced from a cross between line ‘W21MMT70’, which displays partial seedling and adult plant (field) resistance to crown rot, and ‘Mendos’, which is moderately susceptible in seedling tests but partially resistant in field trials. Bulked segregant analysis (BSA) based on seedling trial data did not reveal markers for crown rot resistance. A framework map was produced consisting of 128 microsatellite markers, four phenotypic markers, and one sequence tagged site marker. To this map 331 previously screened AFLP markers were then added. Three quantitative trait loci (QTL) were identified with composite interval mapping across all of the three seedling trials conducted. These QTL are located on chromosomes 2B, 2D and 5D. The 2D and 5D QTL are inherited from the line ‘W21MMT70’, whereas the 2B QTL is inherited from ‘Mendos’. These loci are different from those associated with crown rot resistance in other wheat populations that have been examined, and may represent an opportunity for pyramiding QTL to provide more durable resistance to crown rot.     

Inheritance of field resistance to Septoria tritici blotch in the wheat doubled-haploid population Solitär × Mazurka

Breeding for field resistance to Septoria tritici blotch (STB), caused by Mycosphaerella graminicola (anamorph: Septoria tritici), is the most suitable strategy for controlling this important disease of wheat. Although many Stb genes for resistance to single pathogen isolates have been identified in wheat, knowledge of their efficiency against natural fungal populations is lacking. In a quantitative-trait-locus (QTL) mapping approach in six environments and four locations, field resistance to STB was studied in a doubled-haploid population derived from a cross between the field-resistant cultivar Solitär and the susceptible cultivar Mazurka. After plant height as a disease escape trait was accounted for, five QTL with effects on STB response on chromosomes 5A, 6D and 7D explained 20 % of the genotypic variance; QTL × environment interactions were minor. Field resistance was conferred exclusively by alleles from Solitär, which was previously shown to carry the isolate-specific genes Stb6 and Stb11 as well as minor QTL detected with seven fungal isolates. Surprisingly, neither the Stb6 nor Stb11 isolate-specific genes nor minor QTL previously detected in Solitär were found to be involved in its field resistance. The study suggests that resistance breeding for STB should not rest solely on the deployment of Stb genes. Field tests are indispensable to show their efficacy and durability and to identify genes conferring partial field resistance to STB.     

Practical breeding strategies to improve resistance to Septoria tritici blotch of wheat

Septoria tritici blotch (STB), caused by fungal agent Zymoseptoria tritici (previously known as Mycosphaerella graminicola) is a devastative foliar wheat diseases globally. Importance and potential threat of STB have been discussed historically and geographically. This paper reviews information on the Z. tritici—wheat pathosystem and proposes approaches to identify resistance genes and to advance in breeding for STB resistance. Screening of resistant lines/cultivars, QTL mapping analysis within genetic populations derived from crosses, detection of new resistance gene(s) and finally application of Stb gene carrier line/cultivar in crosses are the major stages of a practical wheat-breeding program against STB of wheat. Phenotyping and genotyping outputs on the top of each other should confirm each other, so it needs to expose a resistance gene carrier line/cultivar in the epidemic condition at seedling/adult plant stage to confirm resistance performance of detected gene(s) in the real condition. On the other word, detecting an associated QTL to resistance should not be considered as the end of investigation. Climate change resulted geographical disease pattern conversion where some diseases became more important in some area where they had not been serious in the past and vice versa. Hence, a reconsideration of wheat disease importance zone is necessary to predict regions where STB is and will be a limitation for wheat yield improvement.     

Oligogenic control of resistance to soil‐borne viruses SBCMV and WSSMV in rye (Secale cereale L.)

Rye production in European growing areas is constrained by the soilborne cereal mosaic virus (SBCMV) and the wheat spindle streak mosaic virus (WSSMV). To date, no European rye cultivars are known to exhibit resistance against these viruses. In this study, we pursued a quantitative trait locus (QTL) mapping strategy to identify genomic regions for resistance to SBCMV and WSSMV in rye. Three populations, each comprising 100 lines segregating for resistance to SBCMV and/or WSSMV, were evaluated for disease response at two years in three locations in Germany where soils are naturally infested with SBCMV and WSSMV. In the combined analysis across environments, one QTL for SBCMV resistance on chromosome 5R explained 31.9% of the phenotypic variation in one of the populations. For WSSMV resistance, one QTL explaining up to 64.0% of the phenotypic variation was detected on chromosome 7R in each of the three populations. On the Triticeae homoeologous group 5, we found evidence for synteny of the major QTL for SBCMV resistance between the wheat and rye genomes.     

Genome-wide association study reveals loci associated with seed longevity in common wheat (Triticum aestivum L.)

Seed longevity could significantly determine seed regeneration cycle and greatly affect wheat production. With the 90 K chip assays, a genome-wide association study was performed to identify seed longevity-related markers and loci in common wheat. Seed germination ratios (GR) under artificially ageing of 166 wheat accessions across three environments were evaluated to assess seed longevity. Totally, 23 longevity-related loci were identified in the study, explaining 6.7%–11.4% of the phenotypic variations. Of these, QlgGR.cas-1A and QlgGR.cas-2B.2 were deemed as stable loci associated with wheat seed longevity. Fifteen loci were found overlapped with known quantitative trait loci or genes. Besides, QlgGR.cas-1A, QlgGR.cas-2B.2, QlgGR.cas-3D.1, QlgGR.cas-3D.2, QlgGR.cas-4A.2, QlgGR.cas-5A.1, QlgGR.cas-5A.2 and QlgGR.cas-6A.1 were colocated with seed dormancy-related loci. Significant additive effects were obtained for seed longevity by pyramiding favourable alleles. Several candidate genes were found involved in signal transduction and stress resistance pathways by sequencing analysis of significantly longevity-related molecular markers. These results might provide new sights into the genetic architecture of seed longevity.     

Identification of quantitative trait loci associated with resistance to foliar diseases in sorghum [<Emphasis Type="Italic">Sorghum bicolor</Emphasis> (L.) Moench]

Forage sorghum cultivars grown in India are susceptible to various foliar diseases, of which anthracnose, rust, zonate leaf spot, drechslera leaf blight and target leaf spot cause severe damage. We report here the quantitative trait loci (QTLs) conferring resistance to these foliar diseases. QTL analysis was undertaken using 168 F₇ recombinant inbred lines (RILs) of a cross between a female parental line 296B (resistant) and a germplasm accession IS18551 (susceptible). RILs and parents were evaluated in replicated field trials in two environments. A total of twelve QTLs for five foliar diseases on three sorghum linkage groups (SBI-03, SBI-04 and SBI-06) were detected, accounting for 6.9–44.9% phenotypic variance. The morphological marker Plant color (Plcor) was associated with most of the QTL across years and locations. The QTL information generated in this study will aid in the transfer of foliar disease resistance into elite susceptible sorghum breeding lines through marker-assisted selection.     

Mapping quantitative trait loci associated with grain filling duration and grain number under terminal heat stress in bread wheat (Triticum aestivum L.)

Terminal heat stress has the potential negative impact on wheat production across the world, especially in South Asia. Under the threat of terminal heat stress, wheat genotypes with stay green trait would suffer from high temperature stress during their long grain filling duration (GFD). The genotypes with short GFD would be advantageous. To identify quantitative trait loci (QTL) for heat tolerance, a RIL population of K 7903 (heat tolerant) and RAJ 4014 (heat sensitive) wheat genotypes was investigated under timely and late‐sown conditions. Heat susceptibility index of GFD, yield components and traits under late‐sown condition were used as phenotypic data for QTL identification. Stable QTLs associated with these traits were identified on chromosomes 1B, 2B, 3B, 5A and 6B. The LOD value ranged from 2.9 to 5.0 and the corresponding phenotyping variation explained ranged from 12.0–22%. QTL for heat susceptibility index for the grain filling duration were colocalized with QTL for productive tillers under late sown and GFD under late‐sown condition on chromosomes 1B and 5A, respectively. These genomic regions could be exploited for molecular wheat breeding programmes targeting heat tolerance.     

玉米抗穗粒腐病QTL定位

用已构建的包括88个AFLP标记和151个SSR标记的遗传图谱和230个F₂植株用于抗病QTL定位研究,在四川雅安、绵阳对F2株系进行抗病性鉴定,采用复合区间定位法进行抗病QTL检测。在雅安检测到位于第2、3、4、6和9染色体上的抗病QTL 6个,解释表型变异的8.3%~25.7%;在绵阳检测到位于第1、6、7和9染色体上的抗病QTL 4个,解释表型变异的11.3%~26.4%。在10个抗病QTL中,位于第6和第9染色体上的2个同时在两点被检测到,贡献率均超过15%,表明玉米穗粒腐病确实存在遗传抗病性。利用2个环境抗病指数的平均值进行抗性QTL检测,共检测到位于第1、6和7连锁群上的3个抗性QTL,单个QTL的贡献率在8.9%~17.2%之间。结果有助于了解玉米穗粒腐病的抗性机制,并为分子标记辅助选择提供理论支撑。     

Changes in allele frequencies in landraces, old and modern barley cultivars of marker loci close to QTL for grain yield under high and low input conditions

Changes in alleles frequencies of marker loci linked to yield quantitative trait loci (QTL) were studied in 188 barley entries (landraces, old and modern cultivars) grown in six trials representing low and high yielding conditions in Spain (2004) and Syria (2004, 2005). A genome wise association analysis was performed per trial, using 811 DArT^® markers of known map position. At the first stage of analysis, spatially adjusted genotypic means were created per trial by fitting mixed models. At the second stage, single QTL models were fitted with correction for population substructure, using regression models. Finally, multiple QTL models were constructed by backward selection from a regression model containing all significant markers from the single QTL analyses. In addition to the association analyses per trial, genotype by environment interaction was investigated across the six trials. Landraces seemed best adapted to low yielding environments, while old and modern entries adapted better to high yielding environments. The number of QTL and the magnitude of their effects were comparable for low and high input conditions. However, none of the QTL were found within a given bin at any chromosome in more than two of the six trials. Changes in allele frequencies of marker loci close to QTL for grain yield in landraces, old and modern barley cultivars could be attributed to selection exercised in breeding, suggesting that modern breeding may have increased frequencies of marker alleles close to QTL that favour production particularly under high yield potential environments. Moreover, these results also indicate that there may be scope for improving yield under low input systems, as breeding so far has hardly changed allele frequencies at marker loci close to QTL for low yielding conditions.     

A quantitative trait locus on chromosome 5B controls resistance of <Emphasis Type="Italic">Triticum turgidum</Emphasis> (L.) var. <Emphasis Type="Italic">diccocoides</Emphasis> to Stagonospora nodorum blotch

Stagonospora nodorum blotch (SNB) is an important foliar disease of durum wheat (Triticum turgidum var. durum) worldwide. The combined effects of SNB and tan spot, considered as components of the leaf spotting disease complex, result in significant damage to wheat production in the northern Great Plains of North America. The main objective of this study was the genetic analysis of resistance to SNB caused by Phaeosphaeria nodorum in tetraploid wheat, and its association with tan spot caused by Pyrenophora tritici-repentis race 2. The 133 recombinant inbred chromosome lines (RICL) developed from the cross LDN/LDN(Dic-5B) were evaluated for SNB reaction at the seedling stage under greenhouse conditions. Molecular markers were used to map a quantitative trait locus (QTL) on chromosome 5B, explaining 37.6% of the phenotypic variation in SNB reaction. The location of the QTL was 8.8 cM distal to the tsn1 locus coding for resistance to P. tritici-repentis race 2. The presence of genes for resistance to both SNB and tan spot in close proximity in tetraploid wheat and the identification of molecular markers linked to these genes or QTLs will be useful for incorporating resistance to these diseases in wheat breeding programs.     

Quantitative trait loci for resistance to Fusarium head blight in recombinant inbred wheat lines from the cross Huapei 57-2 / Patterson

Fusarium head blight (FHB), primarily caused by Fusarium graminearum in North America, often results in significant losses in yield and grain quality of wheat (Triticum aestivum L.). Evaluation of FHB resistance is laborious and can be affected by environmental conditions. The development of DNA markers associated with FHB quantitative trait loci (QTL) and their use in breeding programs could greatly enhance selection. The objective of this study was to identify the location and effect of QTLs for FHB resistance using simple sequence repeat (SSR) markers. A population of wheat recombinant inbred lines derived from the cross ‘Huapei57-2’/‘Patterson’ was characterized for type II resistance in one field experiment and two tests under controlled conditions in the greenhouse. Bulked segregant analysis followed by QTL mapping was used to identify the major segregating QTLs. Results indicate that ‘Huapei 57-2’ may have the same resistance allele as ‘Sumai3’ at a QTL located on the short arm of chromosome 3B. Other QTLs of lower effect size were identified on the long arm of 3Band on chromosomes 3A and 5B. Our findings along with results from other studies demonstrate that the effect of the QTL on3BS is large and consistent across a wide range of genetic backgrounds and environments. Pyramiding this QTL with other FHB QTLs using marker-assisted selection should be effective in improving FHB resistance in a wheat breeding program. This revised version was published online in August 2006 with corrections to the Cover Date.     

小麦水分利用效率及相关性状QTL研究进展

随着水资源的日趋短缺,生物节水在节水农业发展中的地位越来越受到重视。近年来, 随着分子标记技术和数量性状统计分析方法的改进,对调控水分利用效率等复杂数量性状的位点进行作图和定位的研究很多。本文综述了鉴定小麦高WUE的常用指标以及相关性状的QTL定位研究进展,并对目前在QTL定位研究中存在的问题及发展前景进行了探讨。     

Identification and validation of a quantitative trait locus associated with wheat yellow mosaic virus pathotype I resistance in a Japanese wheat variety

Yellow mosaic disease, caused by wheat yellow mosaic virus (WYMV), is one of the most serious diseases of winter wheat (Triticum aestivum L.) in Japan. The three pathotypes of WYMV are distributed in different geographical areas: pathotype I is found mainly in western and central Japan (Kanto), pathotype II in northern Japan (Tohoku and Hokkaido) and pathotype III on the southern island of Japan (Kyushu). A total of 246 doubled‐haploid (DH) lines, derived from a cross between ‘Yumechikara’ (resistant) and ‘Kitahonami’ (susceptible), were evaluated for 2 years for their resistance to WYMV pathotype I. A single major quantitative trait locus, Q.Ymym, mapping to chromosome 2D was associated with resistance to pathotype I in ‘Yumechikara’. This is the first time a QTL responsible for pathotype I resistance has been identified. Fine mapping of Q.Ymym indicated that it was on a tight linkage block originating from ‘Yumechikara’, and the markers associated with this block will accelerate the development of varieties resistant to WYMV pathotype I.     

Mapping a new source of resistance to powdery mildew in mungbean

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