Quantitative trait loci for non-race-specific, high-temperature adult-plant resistance to stripe rust in wheat cultivar Express

Wheat cultivar Express has durable, high-temperature adult-plant (HTAP) resistance to stripe rust (Puccinia striiformis f. sp. tritici). To elucidate the genetic basis of the resistance, Express was crossed with 'Avocet Susceptible' (AVS). A mapping population of 146 F(5) recombinant inbred lines (RILs) was developed using single-seed descent. The RILs were evaluated at two sites near Pullman in eastern Washington and one site near Mount Vernon in western Washington in 2005, and were evaluated near Pullman in 2006 under natural stripe rust infection of predominant races virulent on seedlings of Express. Infection type (IT) and disease severity (DS) were recorded three times for each line during each growing season. The DS data were used to calculate relative area under the disease progress curve (rAUDPC) values. Both IT and rAUDPC data showed continuous distributions, indicating that the Express HTAP resistance was controlled by quantitative trait loci (QTL). Resistance gene analog polymorphism (RGAP) and simple sequence repeat (SSR) techniques were used to map the HTAP resistance QTL. Three QTL were detected with significant additive effects, explaining 49.5-69.6% of the phenotypic variation for rAUDPC. Two of the QTL explained 30.8-42.7% of the phenotypic variation for IT. The three QTL were mapped to wheat chromosomes 6AS, 3BL and 1BL, and were designated as QYrex.wgp-6AS, QYrex.wgp-3BL and QYrex.wgp-1BL, respectively. QYrex.wgp-6AS and QYrex.wgp-3BL, which had higher effects than QYrex.wgp-1BL, were different from previously reported QTL/genes for adult-plant resistance. Markers Xgwm334-Xwgp56 and Xgwm299-Xwgp66 flanking the two major QTL were highly polymorphic in various wheat genotypes, suggesting that these markers are useful in marker-assisted selection.     

Combination of all-stage and high-temperature adult-plant resistance QTL confers high-level,durable resistance to stripe rust in winter wheat cultivar Madsen

Key message

Wheat cultivar Madsen has a new gene on the short arm of chromosome 1A and two QTL for all-stage resistance and three QTL for high-temperature adult-plant resistance that in combination confer high-level, durable resistance to stripe rust.

Abstract

Wheat cultivar Madsen has maintained a high-level resistance to stripe rust over 30 years. To map quantitative trait loci (QTL) underlying the high-level, durable resistance, 156 recombinant inbred lines (RILs) developed from cross Avocet S?×?Madsen were phenotyped with selected races of Puccinia striiformis f. sp. tritici in the greenhouse seedling tests, and in naturally infected fields during 2015–2017. The RILs were genotyped by SSR and SNP markers from genotyping by sequencing and the 90 K wheat SNP chip. Three QTL for all-stage resistance were mapped on chromosomes 1AS, 1BS and 2AS, and two QTL for high-temperature adult-plant (HTAP) resistance were mapped on 3BS and 6BS. The most effective QTL on 2AS, explaining 8.97–23.10% of the phenotypic variation in seedling tests and 8.60–71.23% in field tests, contained Yr17 for all-stage resistance and an additional gene for HTAP resistance. The 6BS QTL, detected in all field tests, was identified as Yr78. The 1AS QTL, conferring all-stage resistance, was identified as a new gene, which explained 20.45 and 30.23% of variation in resistance to races PSTv-37 and PSTv-40, respectively, and contributed significantly to field resistance at Pullman in 2015-2017, but was not detected at Mount Vernon. The interactions among QTL were mostly additive, and RILs with all five QTL had the highest level of resistance in the field, similar to Madsen. Genotyping 148 US Pacific Northwest wheat cultivars with markers for the 1AS, 2AS and 6BS QTL validated the genes and markers, and indicated their usefulness for marker-assisted selection.

     

Identifying QTL for high-temperature adult-plant resistance to stripe rust (<Emphasis Type="Italic">Puccinia striiformis</Emphasis> f. sp. <Emphasis Type="Italic">tritici</Emphasis>) in the spring wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) cultivar ‘Louise’

Over time, many single, all-stage resistance genes to stripe rust (Puccinia striiformis f. sp. tritici) in wheat (Triticum aestivum L.) are circumvented by race changes in the pathogen. In contrast, high-temperature, adult-plant resistance (HTAP), which only is expressed during the adult-plant stage and when air temperatures are warm, provides durable protection against stripe rust. Our objective was to identify major quantitative trait loci (QTL) for HTAP resistance to stripe rust in the spring wheat cultivar ‘Louise’. The mapping population consisted of 188 recombinant inbred lines (RIL) from a Louise (resistant) by ‘Penawawa’ (susceptible) cross. F_5:6 lines were evaluated for stripe rust reaction under natural infection in replicated field trials at five locations in the US Pacific Northwest in 2007 and 2008. Infection type (IT) and disease severity were recorded for each RIL 2–4 times per location. In all environments, Penawawa, the susceptible parent, was rated with an IT ranging from 6 to 8 at all growth stages evaluated. In contrast, Louise, the resistant parent, was rated with an IT of 2 or 3 across growth stages. Distribution of IT values was bimodal, indicating a single major gene was affecting the trait. The parents and RIL population were evaluated with 295 polymorphic simple sequence repeat and one single nucleotide polymorphism markers. One major QTL, designated QYrlo.wpg-2BS, associated with HTAP resistance in Louise, was detected on chromosome 2BS (LOD scores ranging from 5.5 to 62.3 across locations and years) within a 16.9 cM region flanked by Xwmc474 and Xgwm148. SSR markers associated with QYrlo.wpg-2BS are currently being used in marker-based forward breeding strategies to transfer the target region into adapted germplasm to improve the durability of resistance in resulting cultivars.     

Genome-Wide Linkage Mapping of QTL for Adult-Plant Resistance to Stripe Rust in a Chinese Wheat Population Linmai 2 × Zhong 892

Stripe rust is one of the most devastating diseases of wheat (Triticum aestivum) worldwide. Adult-plant resistance (APR) is an efficient approach to provide long-term protection of wheat from the disease. The Chinese winter wheat cultivar Zhong 892 has a moderate level of APR to stripe rust in the field. To determine the inheritance of the APR resistance in this cultivar, 273 F₆ recombinant inbred lines (RILs) were developed from a cross between Linmai 2 and Zhong 892. The RILs were evaluated for maximum disease severity (MDS) in two sites during the 2011–2012, 2012–2013 and 2013–2014 cropping seasons, providing data for five environments. Illumina 90k SNP (single nucleotide polymorphism) chips were used to genotype the RILs and their parents. Composite interval mapping (CIM) detected eight QTL, namely QYr.caas-2AL, QYr.caas-2BL.3, QYr.caas-3AS, QYr.caas-3BS, QYr.caas-5DL, QYr.caas-6AL, QYr.caas-7AL and QYr.caas-7DS.1, respectively. All except QYr.caas-2BL.3 resistance alleles were contributed by Zhong 892. QYr.caas-3AS and QYr.caas-3BS conferred stable resistance to stripe rust in all environments, explaining 6.2–17.4% and 5.0–11.5% of the phenotypic variances, respectively. The genome scan of SNP sequences tightly linked to QTL for APR against annotated proteins in wheat and related cereals genomes identified two candidate genes (autophagy-related gene and disease resistance gene RGA1), significantly associated with stripe rust resistance. These QTL and their closely linked SNP markers, in combination with kompetitive allele specific PCR (KASP) technology, are potentially useful for improving stripe rust resistances in wheat breeding.     

Identification and mapping QTL for high-temperature adult-plant resistance to stripe rust in winter wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) cultivar ‘Stephens’

High-temperature adult-plant (HTAP) resistance from the winter wheat (Triticum aestivum) cultivar 'Stephens' has protected wheat crops from stripe rust caused by Puccinia striiformis f. sp. tritici for 30 years. The objectives of this study were to identify quantitative trait loci (QTL) for HTAP resistance in Stephens through genetic linkage analysis and identify DNA markers linked to the QTL for use in marker-assisted breeding. Mapping populations consisted of 101 recombinant inbred lines (RILs) through single-seed descent from 'Stephens' (resistant) x 'Michigan Amber' (susceptible). F(5), F(6) and F(7) RILs were evaluated for stripe rust resistance at Pullman, WA in 1996, 1997 and 1998, respectively, whereas F(8) RILs were evaluated at Mt Vernon, WA, USA in 2005. The 101 F(8) RILs were evaluated with 250 resistance gene analog polymorphism (RGAP), 245 simple sequence repeat (SSR) and 1 sequence tagged site (STS) markers for genetic linkage map construction. Two QTL, which explained 48-61% of the total phenotypic variation of the HTAP resistance in Stephens, were identified. QYrst.wgp-6BS.1 was within a 3.9-cM region flanked by Xbarc101 and Xbarc136. QYrst.wgp-6BS.2 was mapped in a 17.5-cM region flanked by Xgwm132 and Xgdm113. Both two QTL were physically mapped to the short arm of chromosome 6B, but in different bins. Validation and polymorphism tests of the flanking markers in 43 wheat genotypes indicated that the molecular markers associated with these QTL should be useful in marker-assisted breeding programs to efficiently incorporate HTAP resistance into new wheat cultivars.     

Genome-wide identification of QTL conferring high-temperature adult-plant (HTAP) resistance to stripe rust (<Emphasis Type="Italic">Puccinia striiformis</Emphasis> f. sp<Emphasis Type="Italic">. tritici</Emphasis>) in wheat

High-temperature adult-plant (HTAP) resistance to stripe rust (caused by Puccinia striiformis f. sp. tritici) is a durable type of resistance in wheat (Triticum aestivum L.). This study identified quantitative trait loci (QTL) conferring HTAP resistance to stripe rust in a population consisting of 169 F_8:10 recombinant inbred lines (RILs) derived from a cross between a susceptible cultivar Rio Blanco and a resistant germplasm IDO444. HTAP resistance was evaluated for both disease severity and infection type under natural infection over two years at two locations. The genetic linkage maps had an average density of 6.7 cM per marker across the genome and were constructed using 484 markers including 96 wheat microsatellite (SSR), 632 Diversity Arrays Technology (DArT) polymorphisms, two sequence-tagged-site (STS) from semi-dwarf genes Rht1 and Rht2, and two markers for low molecular-weight glutenin gene subunits. QTL analysis detected a total of eight QTL significantly associated with HTAP resistance to stripe rust with two on chromosome 2B, two on 3B and one on each of 1A, 4A, 4B and 5B. QTL on chromosomes 2B and 4A were the major loci derived from IDO444 and explained up to 47 and 42% of the phenotypic variation for disease severity and infection type, respectively. The remaining five QTL accounted for 7–10% of the trait variation. Of these minor QTL, the resistant alleles at the two QTL QYrrb.ui-3B.1 and QYrrb.ui-4B derived from Rio Blanco and reduced infection type only, while the resistant alleles at the other three QTL, QYrid.ui-1A, QYrid.ui-3B.2 and QYrid.ui-5B, all derived from IDO444 and reduced either infection type or disease severity. Markers linked to 2B and 4A QTL should be useful for selection of HTAP resistance to stripe rust.     

Genetics and molecular mapping of genes for race-specific all-stage resistance and non-race-specific high-temperature adult-plant resistance to stripe rust in spring wheat cultivar Alpowa

Stripe rust, caused by Puccinia striiformis f. sp. tritici, is one of the most widespread and destructive wheat diseases worldwide. Growing resistant cultivars is the preferred control of the disease. The spring wheat cultivar ‘Alpowa’ has both race-specific, all-stage resistance and non-race-specific, high-temperature adult-plant (HTAP) resistances to stripe rust. To identify genes for the stripe rust resistances, Alpowa was crossed with ‘Avocet Susceptible’ (AVS). Seedlings of the parents, and F₁, F₂ and F₃ progeny were tested with races PST-1 and PST-21 of P. striiformis f. sp. tritici under controlled greenhouse conditions. Alpowa has a single partially dominant gene, designated as YrAlp, conferring all-stage resistance. Resistance gene analog polymorphism (RGAP) and simple sequence repeat (SSR) techniques were used to identify molecular markers linked to YrAlp. A linkage group of five RGAP markers and two SSR markers was constructed for YrAlp using 136 F₃ lines. Amplification of a set of nulli-tetrasomic Chinese Spring lines with RGAP markers Xwgp47 and Xwgp48 and the two SSR markers indicated that YrAlp is located on the short arm of chromosome 1B. To map quantitative trait loci (QTLs) for the non-race-specific HTAP resistance, the parents and 136 F₃ lines were tested at two sites near Pullman and one site near Mount Vernon, Washington, under naturally infected conditions. A major HTAP QTL was consistently detected across environments and was located on chromosome 7BL. Because of its chromosomal location and the non-race-specific nature of the HTAP resistance, this gene is different from previously described genes for adult-plant resistance, and is therefore designated Yr39. The gene contributed to 64.2% of the total variation of relative area under disease progress curve (AUDPC) data and 59.1% of the total variation of infection type data recorded at the heading-flowering stages. Two RGAP markers, Xwgp36 and Xwgp45 with the highest R ² values were closely linked to Yr39, should be useful for incorporation of the non-race-specific resistance gene into new cultivars and for combining Yr39 with other genes for durable and high-level resistance.     

Evaluating stripe rust resistance in Indian wheat genotypes and breeding lines using molecular markers

Stripe rust (yellow rust), caused by Puccinia striiformis f. sp. tritici (Pst), is a serious disease of wheat worldwide, including India. Growing resistant cultivars is the most cost-effective and eco-friendly approach to manage the disease. In this study, 70 publically available molecular markers were used to identify the distribution of 35 Yr genes in 68 wheat genotypes. Out of 35 Yr genes, 25 genes amplified the loci associated with Yr genes. Of the 35, 18 were all-stage resistance ASR (All-stage resistance) genes and 7 (Yr16, Yr18, Yr29, Yr30, Yr36, Yr46 & Yr59) were APR (Adult-plant resistance) genes. In the field tests, evaluation for stripe rust was carried out under artificial inoculation of Pst. Fifty-three wheat genotypes were found resistant to yellow rust (ITs 0), accounting for 77.94% of total entries. Coefficients of infection ranged from 0 to 60 among all wheat genotypes. Two genotypes (VL 1099 & VL 3002) were identified with maximum 15 Yr genes followed by 14 genes in VL 3010 and HI8759, respectively. Maximum number of all-stage resistance genes were identified in RKD 292 (11) followed by ten genes in DBW 216, WH 1184 and VL 3002. Maximum number of adult-plant resistance gene was identified in VL 3009 (6), HI 8759 (5) and Lassik (4) respectively. Genes Yr26 (69.2%), Yr2 (69.1%), Yr64 (61.7%), Yr24 (58.9%), Yr7 (52.9%), Yr10 (50%) and Yr 48 (48.5%) showed high frequency among selected wheat genotypes, while Yr9 (2.94%), Yr36 (2.94%), Yr60 (1.47%) and Yr32 (8.8%) were least frequent in wheat genotypes. In future breeding programs, race specific genes and non-race specific genes should be utilised to pyramid with other effective genes to develop improved wheat cultivars with high-level and durable resistance to stripe rust. Proper deployment of Yr genes and utilizing the positive interactions will be helpful for resistance breeding in wheat.     

High-temperature adult-plant (HTAP) stripe rust resistance gene Yr36 from Triticum turgidum ssp. dicoccoides is closely linked to the grain protein content locus Gpc-B1

Several new races of the stripe rust pathogen have become frequent throughout the wheat growing regions of the United States since 2000. These new races are virulent to most of the wheat seedling resistance genes limiting the resistance sources that can be used to combat this pathogen. High-temperature adult-plant (HTAP) stripe rust resistance has proven to be more durable than seedling resistance due to its non-race-specific nature, but its use is limited by the lack of mapping information. We report here the identification of a new HTAP resistance gene from Triticum turgidum ssp. dicoccoides (DIC) designated as Yr36. Lines carrying this gene were susceptible to almost all the stripe rust pathogen races tested at the seedling stage but showed adult-plant resistance to the prevalent races in California when tested at high diurnal temperatures. Isogenic lines for this gene were developed by six backcross generations. Field tests in two locations showed increased levels of field resistance to stripe rust and increased yields in isogenic lines carrying the Yr36 gene compared to those without the gene. Recombinant substitution lines of chromosome 6B from DIC in the isogenic background of durum cv. Langdon were used to map the Yr36 gene on the short arm of chromosome 6B completely linked to Xbarc101, and within a 2-cM interval defined by PCR-based markers Xucw71 and Xbarc136. Flanking locus Xucw71 is also closely linked to the grain protein content locus Gpc-B1 (0.3-cM). Marker-assisted selection strategies are presented to improve stripe rust resistance and simultaneously select for high or low Gpc-B1 alleles.     

Genome-Wide Association Mapping for Resistance to Leaf and Stripe Rust in Winter-Habit Hexaploid Wheat Landraces

Leaf rust, caused by Puccinia triticina (Pt), and stripe rust, caused by P. striiformis f. sp. tritici (Pst), are destructive foliar diseases of wheat worldwide. Breeding for disease resistance is the preferred strategy of managing both diseases. The continued emergence of new races of Pt and Pst requires a constant search for new sources of resistance. Here we report a genome-wide association analysis of 567 winter wheat (Triticum aestivum) landrace accessions using the Infinium iSelect 9K wheat SNP array to identify loci associated with seedling resistance to five races of Pt (MDCL, MFPS, THBL, TDBG, and TBDJ) and one race of Pst (PSTv-37) frequently found in the Northern Great Plains of the United States. Mixed linear models identified 65 and eight significant markers associated with leaf rust and stripe rust, respectively. Further, we identified 31 and three QTL associated with resistance to Pt and Pst, respectively. Eleven QTL, identified on chromosomes 3A, 4A, 5A, and 6D, are previously unknown for leaf rust resistance in T. aestivum.     

Comparative genome-wide mapping versus extreme pool-genotyping and development of diagnostic SNP markers linked to QTL for adult plant resistance to stripe rust in common wheat

Key message

A major stripe rust resistance QTL on chromosome 4BL was localized to a 4.5-Mb interval using comparative QTL mapping methods and validated in 276 wheat genotypes by haplotype analysis.

Abstract

CYMMIT-derived wheat line P10103 was previously identified to have adult plant resistance (APR) to stripe rust in the greenhouse and field. The conventional approach for QTL mapping in common wheat is laborious. Here, we performed QTL detection of APR using a combination of genome-wide scanning and extreme pool-genotyping. SNP-based genetic maps were constructed using the Wheat55 K SNP array to genotype a recombinant inbred line (RIL) population derived from the cross Mingxian 169?×?P10103. Five stable QTL were detected across multiple environments. After comparing SNP profiles from contrasting, extreme DNA pools of RILs six putative QTL were located to approximate chromosome positions. A major QTL on chromosome 4B was identified in F_2:4 contrasting pools from cross Zhengmai 9023?×?P10103. A consensus QTL (LOD?=?26–40, PVE?=?42–55%), named QYr.nwafu-4BL, was defined and localized to a 4.5-Mb interval flanked by SNP markers AX-110963704 and AX-110519862 in chromosome arm 4BL. Based on stripe rust response, marker genotypes, pedigree analysis and mapping data, QYr.nwafu-4BL is likely to be a new APR QTL. The applicability of the SNP-based markers flanking QYr.nwafu-4BL was validated on a diversity panel of 276 wheat lines. The additional minor QTL on chromosomes 4A, 5A, 5B and 6A enhanced the level of resistance conferred by QYr.nwafu-4BL. Marker-assisted pyramiding of QYr.nwafu-4BL and other favorable minor QTL in new wheat cultivars should improve the level of APR to stripe rust.

     

QTL mapping for adult-plant resistance to stripe rust in Italian common wheat cultivars Libellula and Strampelli

Italian common wheat cultivars Libellula and Strampelli, grown for over three decades in Gansu province of China, have shown effective resistance to stripe rust. To elucidate the genetic basis of the resistance, F₃ populations were developed from crosses between the two cultivars and susceptible Chinese wheat cultivar Huixianhong. The F₃ lines were evaluated for disease severity in Beijing, Gansu and Sichuan from 2005 to 2008. Joint- and single-environment analyses by composite interval mapping identified five quantitative trait loci (QTLs) in Libellula for reduced stripe rust severity, designated QYr.caas-2DS, QYr.caas-4BL, QYr.caas-5BL.1, QYr.caas-5BL.2 and QYr.caas-7DS, and explained 8.1–12.4, 3.6–5.1, 3.4–8.6, 2.6 and 14.6–35.0%, respectively, of the phenotypic variance across four environments. Six interactions between different pairs of QTLs explained 3.2–7.1% of the phenotypic variance. The QTLs QYr.caas-4BL, QYr.caas-5BL.1 and QYr.caas-7DS were also detected in Strampelli, explaining 4.5, 2.9–5.5 and 17.1–39.1% of phenotypic variance, respectively, across five environments. Three interactions between different pairs of QTLs accounted for 6.1–35.0% of the phenotypic variance. The QTL QYr.caas-7DS flanked by markers csLV34 and Xgwm295 showed the largest effect for resistance to stripe rust. Sequence analyses confirmed that the lines with the QYr.caas-7DS allele for resistance carried the resistance allele of the Yr18/Lr34 gene. Our results indicated that the adult-plant resistance gene Yr18 and several minor genes confer effective durable resistance to stripe rust in Libellula and Strampelli.     

Mapping of quantitative adult plant field resistance to leaf rust and stripe rust in two European winter wheat populations reveals co-location of three QTL conferring resistance to both rust pathogens

Key message

We detected several, most likely novel QTL for adult plant resistance to rusts. Notably three QTL improved resistance to leaf rust and stripe rust simultaneously indicating broad spectrum resistance QTL.

Abstract

The rusts of wheat (Puccinia spp.) are destructive fungal wheat diseases. The deployment of resistant cultivars plays a central role in integrated rust disease management. Durability of resistance would be preferred, but is difficult to analyse. The Austrian winter wheat cultivar Capo was released in the 1989 and grown on a large acreage during more than two decades and maintained a good level of quantitative leaf rust and stripe rust resistance. Two bi-parental mapping populations: Capo × Arina and Capo × Furore were tested in multiple environments for severity of leaf rust and stripe rust at the adult plant stage in replicated field experiments. Quantitative trait loci associated with leaf rust and stripe rust severity were mapped using DArT and SSR markers. Five QTL were detected in multiple environments associated with resistance to leaf rust designated as QLr.ifa-2AL, QLr.ifa-2BL, QLr.ifa-2BS, QLr.ifa-3BS, and QLr.ifa-5BL, and five for resistance to stripe rust QYr.ifa-2AL, QYr.ifa-2BL, QYr.ifa-3AS, QYr.ifa-3BS, and QYr.ifa-5A. For all QTL apart from two (QYr.ifa-3AS, QLr.ifa-5BL) Capo contributed the resistance improving allele. The leaf rust and stripe rust resistance QTL on 2AL, 2BL and 3BS mapped to the same chromosome positions, indicating either closely linked genes or pleiotropic gene action. These three multiple disease resistance QTL (QLr.ifa-2AL/QYr.ifa-2AL, QLr.ifa.2BL/QYr.ifa-2BL, QLr.ifa-3BS/QYr.ifa.3BS) potentially contribute novel resistance sources for stripe rust and leaf rust. The long-lasting resistance of Capo apparently rests upon a combination of several genes. The described germplasm, QTL and markers are applicable for simultaneous resistance improvement against leaf rust and stripe rust.     

Characterization and molecular mapping of Yr52 for high-temperature adult-plant resistance to stripe rust in spring wheat germplasm PI 183527

Stripe rust, caused by Puccinia striiformis f. sp. tritici, is one of the most destructive diseases of wheat worldwide. Resistance is the best approach to control the disease. High-temperature adult-plant (HTAP) stripe rust resistance has proven to be race non-specific and durable. However, genes conferring high-levels of HTAP resistance are limited in number and new genes are urgently needed for breeding programs to develop cultivars with durable high-level resistance to stripe rust. Spring wheat germplasm PI 183527 showed a high-level of HTAP resistance against stripe rust in our germplasm evaluations over several years. To elucidate the genetic basis of resistance, we crossed PI 183527 and susceptible wheat line Avocet S. Adult plants of parents, F(1), F(2) and F(2:3) progeny were tested with selected races under the controlled greenhouse conditions and in fields under natural infection. PI 183527 has a single dominant gene conferring HTAP resistance. Resistance gene analog polymorphism (RGAP) and simple sequence repeat (SSR) markers in combination with bulked segregant analysis (BSA) were used to identify markers linked to the resistance gene. A linkage map consisting of 4 RGAP and 7 SSR markers was constructed for the resistance gene using data from 175 F(2) plants and their derived F(2:3) lines. Amplification of nulli-tetrasomic, ditelosomic and deletion lines of Chinese Spring with three RGAP markers mapped the gene to the distal region (0.86-1.0) of chromosome 7BL. The molecular map spanned a genetic distance of 27.3?cM, and the resistance gene was narrowed to a 2.3-cM interval flanked by markers Xbarc182 and Xwgp5258. The polymorphism rates of the flanking markers in 74 wheat lines were 74 and 30?%, respectively; and the two markers in combination could distinguish the alleles at the resistance locus in 82?% of tested genotypes. To determine the genetic relationship between this resistance gene and Yr39, a gene also on 7BL conferring HTAP resistance in Alpowa, a cross was made between PI 183527 and Alpowa. F(2) segregation indicated that the genes were 36.5?±?6.75?cM apart. The gene in PI 183527 was therefore designed as Yr52. This new gene and flanking markers should be useful in developing wheat cultivars with high-level and possible durable resistance to stripe rust.     

QTL characterization of resistance to leaf rust and stripe rust in the spring wheat line Francolin#1

Growing resistant wheat varieties is a key method of controlling two important wheat diseases, leaf rust and stripe rust. We analyzed quantitative trait loci (QTL) to investigate adult plant resistance (APR) to these rusts, using 141 F₅ RILs derived from the cross ‘Avocet-YrA/Francolin#1’. Phenotyping of leaf rust resistance was conducted during two seasons at Ciudad Obregon, Mexico, whereas stripe rust was evaluated for two seasons in Toluca, Mexico, and one season in Chengdu, China. The genetic map was constructed with 581 markers, including diversity arrays technology and simple sequence repeat. Significant loci for reducing leaf rust severity were designated QLr.cim-1BL, QLr.cim-3BS.1, QLr.cim-3DC, and QLr.cim-7DS. The six QTL that reduced stripe rust severity were designated QYr.cim-1BL, QYr.cim-2BS, QYr.cim-2DS, QYr.cim-3BS.2, QYr.cim-5AL, and QYr.cim-6AL. All loci were conferred by Francolin#1, with the exception of QYr.cim-2DS, QYr.cim-5AL, and QYr.cim-6AL, which were derived from Avocet-YrA. Closely linked markers indicated that the 1BL locus was the pleiotropic APR gene Lr46/Yr29. QYr.cim-2BS was a seedling resistance gene designated as YrF that conferred intermediate seedling reactions and moderate resistance at the adult plant stage in both Mexican and Chinese environments. Significant additive interactions were detected between the six QTL for stripe rust, but not between the four QTL for leaf rust. Furthermore, we detected two new APR loci for leaf rust in common wheat: QLr.cim-3BS.1 and QLr.cim-7DS.     

Genetic analysis of durable resistance to yellow rust in bread wheat

Yellow rust, caused by Puccinia striiformis, is one of the most damaging diseases affecting bread wheat in temperate regions. Although resistance to yellow rust is frequently overcome by new virulent races, a durable form of resistance in the French bread wheat Camp Rémy (CR) has remained effective since its introduction in 1980. We used 217 F₇ recombinant inbred lines (RILs) derived from the cross between CR and the susceptible cultivar Récital to identify and map quantitative trait loci (QTLs) involved in durable yellow rust resistance. Six significant QTLs that were stable over a 4-year period were detected. Two QTLs, denoted QYr.inra-2DS and QYr.inra-5BL.2, were located on the short arm of chromosome 2D and the long arm of chromosome 5B, respectively. Each explained on average 25–35% of the observed phenotypic variation and were probably inherited from Cappelle Desprez, a parent of CR that confers durable adult plant resistance to yellow rust. QYr.inra-2DS probably corresponds to the Yr16 gene. The most consistent QTL, designated QYr.inra-2BL, was located on the centromeric region of chromosome 2B and explained 61% of the phenotypic variation in 2003. This QTL was responsible for seedling-stage resistance and may correspond to a cluster of genes, including Yr7. The remaining QTLs were mapped to the short arm of chromosome 2B (R²=22–70%) and to the long arm of chromosomes 2A (R²=0.20–0.40) and 5B (R²=0.18–0.26). This specific combination of seedling and adult plant resistance genes found in CR and CD may constitute the key to their durable resistance against yellow rust.     

Two stripe rust effectors impair wheat resistance by suppressing import of host Fe–S protein into chloroplasts

Several effectors from phytopathogens usually target various cell organelles to interfere with plant defenses, and they generally contain sequences that direct their translocation into organelles, such as chloroplasts. In this study, we characterized a different mechanism for effectors to attack chloroplasts in wheat (Triticum aestivum). Two effectors from Puccinia striiformis f. sp. tritici (Pst), Pst_4, and Pst_5, inhibit Bax-mediated cell death and plant immune responses, such as callose deposition and reactive oxygen species (ROS) accumulation. Gene silencing of the two effectors induced significant resistance to Pst, demonstrating that both effectors function as virulence factors of Pst. Although these two effectors have low sequence similarities and lack chloroplast transit peptides, they both interact with TaISP (wheat cytochrome b6–f complex iron–sulfur subunit, a chloroplast protein encoded by nuclear gene) in the cytoplasm. Silencing of TaISP impaired wheat resistance to avirulent Pst and resulted in less accumulation of ROS. Heterogeneous expression of TaISP enhanced chloroplast-derived ROS accumulation in Nicotiana benthamiana. Co-localization in N. benthamiana and western blot assay of TaISP content in wheat chloroplasts show that both effectors suppressed TaISP from entering chloroplasts. We conclude that these biotrophic fungal effectors suppress plant defenses by disrupting the sorting of chloroplast protein, thereby limiting host ROS accumulation and promoting fungal pathogenicity.

Despite the lack of chloroplast transit peptide, rust effectors affect chloroplast-mediated defenses by suppressing import of host Fe–S protein to chloroplast to promote pathogenicity of stripe rust.