A novel barley scald resistance gene: genetic mapping of the Rrs15 scald resistance gene derived from wild barley, Hordeum vulgare ssp. spontaneum

Most genes for resistance to barley leaf scald map either to the Rrs1 locus on the long arm of chromosome 3H, or the Rrs2 locus on the short arm of chromosome 7H. Other loci containing scald resistance genes have previously been identified using lines derived from wild barley, Hordeum vulgare ssp. spontaneum. A single dominant gene conditioning resistance to scald was identified in a third backcross (BC3F3) line derived from an Israeli accession of wild barley. The resistance gene is linked to three microsatellite markers that map to the long arm of chromosome 7H; the closest of these loci, HVM49, maps 11.5 cM from the resistance gene. As no other scald resistance genes have been mapped to this chromosome arm, it is considered to be a novel scald resistance locus. As the Acp2 isozyme locus is linked to this scald resistance locus, at 17.7 cM, Acp2 is assigned to chromosome 7H. Molecular markers linked to the novel scald resistance gene, designated Rrs15, can be used in breeding for scald resistance.     

Genetic mapping of the barley Rrs14 scald resistance gene with RFLP, isozyme and seed storage protein markers

The scald susceptible barley cultivar ‘Clipper’ and a third‐backcross (BC₃) line homozygous for the Rrs14 scald resistance gene that originally came from Hordeum vulgare ssp. spontaneum were grown in replicated field trials. The level of resistance that Rrs14 confers against field populations of the pathogen Rhynchosporium secalis, the causal agent of scald disease, was evaluated. The Rrs14 BC₃ line exhibited 80% and 88% less leaf damage than ‘Clipper’ in 1995 and 1996, respectively. Given this effectiveness of Rrs14, research was undertaken to identify a linked marker locus suitable for indirect selection of Rrs14. Based on linkage to a set of previously mapped loci, Rrs14 was positioned to barley chromosome 1H between the seed storage protein (hordein) loci Hor1 and Hor2, approximately 1.8 cM from the latter locus. The Hor2 locus is thus an ideal codominant molecular marker for Rrs14. The tight linkage between Rrs14 and Hor2 and the availability of alternative biochemical and molecular techniques for scoring Hor2 genotypes, permits simple indirect selection of Rrs14 in barley scald resistance breeding programmes.     

Linkage of Rust Resistance Genes from Wild Barley (Hordeum spontaneum) with Isozyme Markers

Eighty-three third backcross lines which comprise a set of near isogenic lines (NIL's) of the barley cultivar ‘Clipper’ but each carrying a different chromosomal segment from Hordeum spontaneum, marked with a distinct isozyme, were tested for resistance to three races of the barley leaf rust pathogen (Puccmia hordei). Fourteen lines showed resistance to at least one race and three showed resistance to all three races. The resistance in two of these lines was controlled by separate, single partially dominant genes. In one case the resistance gene named Rph1O was on chromosome 3 and linked (r = 0.15 ±0.05) with the isozyme locus Est2. In the second case, the gene (Rph11) was on barley chromosome 6 and linked (r = 0.07±0.02) with the isozyme locus Acp3 and (r = 0.11±0.02) with Dip2.     

Potential and limitations of isozymes for chromosomal localization of resistance genes against barley mild mosaic virus (BaMMV)

Summary To assess the possibilities offered by isozymes to locate resistance genes against barley mild mosaic virus (BaMMV), the isozyme patterns of 19 barley (Hordeum vulgare L.) genotypes carrying genes different from ym4 were determined. Of the 15 isozyme systems tested, only three were polymorphic, namely aconitate hydratase, esterase, phosphogluconate dehydrogenase, providing markers on four of the seven barley chromosomes. Studies of F₂ progenies derived from three crosses between resistant genotypes and susceptible varieties failed to reveal linkage between resistance genes and isozymes. Another goal of the experiment was to study the linkage relationships between ym4 and the esterase locus (Est1-Est2-Est4). Our estimates of the recombination rate between these two loci (3.41 and 8.32%) were in the range of those reported between these esterases and one of the resistance genes of the Chinese variety Mokusekko 3.     

Detection of an earliness per se quantitative trait locus in the proximal region of wheat chromosome 5AL

A homoeologous quantitative trait locus to that of eps5L on barley chromosome 5H was identified in a syntenic region of wheat chromosome 5A. Wheat single chromosome recombinant lines (SCRs) were developed from a cross between ‘Chinese Spring’(‘Cappelle-Desprez’ 5A) and ‘Chinese Spring’(Triticum spelta 5A), these were grown together with the parental controls under different vernalization and photoperiod regimes. The variation for ear emergence time accelerated heading induced by the T. spelta segment indicated an effect associated with the Xcdo412-Xbcd9 interval. Since no differences between the SCRs and controls in responses to vernalization and photoperiod treatments were detected, this effect was identified as an earliness per se gene, Q Eetocs-5 A.2, which may be homoeologous to the eps5L quantitative trait locus of barley. Xbcd926 has been found to be closely linked to the rice flowering time quantitative trait loci, QHd9a or FLTQ2, on chromosome 9, suggesting possible relationships among the quantitative trait loci across wheat, barley and rice genomes.     

Identification of Resistance Genes Against Powdery Mildew in Four Accessions of Hordeum Vulgare SSP. Spontaneum

Summary Four newly detected accessions of wild barley (Hordeum vulgare ssp. spontaneum) resistant to powdery mildew caused by Blumeria graminis f. sp. hordei were studied with the aim of finding the number of genes/loci conferring the resistance of individual accessions, the type of inheritance of the genes and their relationships to the Mla locus. F₂ populations after crosses between the winter variety ‘Tiffany’ and four wild barley accessions and use of microsatellite DNA markers were focused on the identification of individual resistance genes/loci by means of their chromosomal locations. In PI466495, one locus conferring powdery mildew resistance was identified in highly significant linkage with the marker Bmac0213. This location is consistent with the known locus Mla on chromosome 1HS. In the other three accessions the resistance was determined by two independent loci. In PI466197, PI466297 and PI466461, one locus was identified on chromosome 1HS and three new loci were revealed on chromosomes 2HS (highly significant linkage with Bmac0134), 7HS (highly significant linkage with Bmag0021) and 7HL (significant linkage with EBmac0755). Our prospective aim is identification of further linked DNA markers and the exact location of the resistance genes on the barley chromosomes.     

Development of SCAR markers linked to a scald resistance gene derived from wild barley

The F₂ progeny of a third backcross(BC₃) line, BC line 240, derived from a Turkish accession of wild barley (Hordeum vulgare ssp. spontaneum),segregated for resistance to scald (Rhynchosporium secalis) in a manner indicating the presence of a single dominant resistance gene. Two SCAR marker slinked to this resistance were developed from AFLP markers. Screens of disomic and ditelosomic wheat-barley addition lines with the SCAR markers demonstrated that the scald resistance gene is located in the centromeric region of barley chromosome 3H,a region previously reported to contain a major scald resistance locus, Rrs1. Markers that flank the Rrs1 locus were used to screen the wild barley-derivedBC₃F₂ population. These markers also flank the wild barley-derived scald resistance, indicating that it maps to the same locus as Rrs1; it may beallelic, or a separate gene within a complex locus. However, BC line 240 does not respond to treatment with the Rhynchosporium secalis avirulence factorNIP1 in the same way as the Rrs1-carrying cultivar Atlas46. This suggests that the scald resistance gene derived from wild barley confers a different specificity of response to theRrs1 allele in Atlas46.In order to increase the durability of scald resistance in the field, we suggest that at least two scald resistances should be combined into barley cultivars before release. The scald resistance gene described here will be of value in the Australian environment, and the several markers linked to it will facilitate pyramiding. This revised version was published online in July 2006 with corrections to the Cover Date.     

Associations between 23 Quantitative Traits and 10 Genetic Markers in a Barley Cross

Associations of 23 quantitative traits and 10 genetic marker characters were examined in 63 chromosome-doubled lines (DH-lines) derived from the F₁- generation of a cross between an old and a modern spring barley variety. One fourth of the marker × trait combinations showed significant associations. More than two thirds of these associations were to earliness (heading date). Earliness was found to be controlled by two loci: the previously known eak locus and a new locus designated Ea. It is concluded that the associations of quantitative traits with the earliness loci were caused by pleiotropy. Associations were found between two absolutely linked C-bands on chromosome 3 and QTLs (quantitative trait loci) for lodging, straw diameter, and length of top internode of the straw. The three loci on chromosome 5, eak and the two linked powdery mildew loci, Mla9 and Mlk, were associated with a possible QTL for magnesium concentration in grain. Association to the C-band on chromosome 6 suggests QTLs for TGW (thousand grain weight), straw diameter and magnesium concentration in grain. Locus Estl on chromosome 3 was not associated with any of the quantitative traits.     

Esterase 1 and Hordein C allelic and genotypic frequencies in CC XXXI,a male sterile barley population

Summary Seeds from the 5th, 6th, 12th and 18th generation of enforced outcrossing in CCXXXXI-B, a barley (Hordeum vulgare L.) population containing genetic male sterility, were assayed for genotypic distribution of Est 1 and Hor C alleles. The result suggest that natural selection was effective in altering allelic frequencies at both loci and that genotypic frequencies were often different from what would be expected if allelic frequencies solely determined genotypic frequencies. Departures of actual from expected genotypic frequencies occurred in that shifts in allelic frequencies were detected.Joint contribution of the USDA-ARS and published with the approval of the Director of the Montana Agric. Exp. Sta. Journal Series No. 2281.     

Quantitative trait loci for scald resistance in barley localized by a non-interval mapping procedure

Three quantitative trait loci (QTL) for scald resistance in barley were identified and mapped in relation to molecular markers using a population of chromosome doubled‐haploid lines produced from the F₁ generation of a cross between the spring barley varieties ‘Alexis’ and ‘Regatta’. Two field experiments were conducted in Denmark and two in Norway to assess disease resistance. The percentage leaf area covered with scald (Rhynchosporium secalis) ranged from 0 to 40% in the 189 doubled‐haploid (DH) lines analysed. One quantitative trait locus was localized in the centromeric region of chromosome 3H, Qryn3, using the MAPQTL program. MAPQTL was unable to provide proper localization of the other two resistance genes and so a non‐interval QTL mapping method was used. One was found to be located distally to markers on chromosome 4H (Qryn4) and the other, Qryn6, was located distally to markers on chromosome 6H. The effects of differences between the Qryn3, Qryn4 and Qryn6 alleles in two barley genotypes for the QTL were estimated to be 8.8%, 7.3% and 7.0%, respectively, of leaf covered by scald. No interactions between the QTLs were found.     

Outcrossing Rates in Autumn and Spring-Sown Barley

Natural outcrossing is a potential cause of the admixtures which are often observed, especially in winter barley. Outcrossing was studied in yield trials with hap laid -derived homozygous offspring, lines from tour crosses between winter and spring barley varieties. Outcrosses were recognised by heterozygosity in storage protein loci Hor1 and Hor2 as revealed by electrophoresis. Outcrossing, was, on average, about 5% in lines sown in the autumn and less than 0.5% after spring sowing. Autumn-sown lines from one of the four crosses studied had about 10% outcrosses.     

Sub-arm location of prolamin and EST-SSR loci on chromosome 1H<Superscript>ch</Superscript> from <Emphasis Type="Italic">Hordeum chilense</Emphasis>

Hordeum chilense Roem. et Schult. is a diploid wild South American barley that contains genes of interest for cereal breeding, many of them located on chromosome 1H^ch. In the current study, two H. chilense-wheat addition lines with deletions in the 1H^ch chromosome were used for sub-arm localization of five prolamin (glutenin and gliadin) loci and 33 EST-SSR marker loci on chromosome 1H^ch. The two sets of markers were distributed across five sub-arm chromosome regions. Three glutenin loci (Glu-H ^ch 2, Glu-H ^ch 3, Glu-H ^ch 4) together with the gliadin locus Gli-H ^ch 1 were located on the distal 20% of the 1H^chS arm, whereas the glutenin locus Glu-H ^ch 1 was on the proximal 88% region of 1H^chL. Among 33 EST-SSR marker loci, 7 (21.2%) were on the 1H^chS arm and, of them, 3 (9.1%) were on the distal 20% end and 4 (12.1%) on the proximal 80% region. The 26 loci (78.8%) on 1H^chL were distributed across three different regions: 18 (78.8%) in the proximal 88%, 3 (9.1%) in the distal 12% and 5 (15.2%) in a region less than 12% from the distal end. The deletions in the 1H^ch chromosome added to the common wheat background were thus shown to be useful for determining the sub-arm location of EST-SSR and prolamin loci. This could facilitate the identification of molecular markers linked to genes of agronomic interest and the isolation of such genes for use in common wheat improvement.     

Candidate markers for powdery mildew resistance genes from wild barley PI284752

PI284752, an accession of wild barley (Hordeum vulgare ssp. spontaneum) resistant to powdery mildew caused by Blumeria graminis f.sp. hordei, was studied with the aim of identifying genes involved in powdery mildew resistance. An F₂ population (456 plants) was established from a cross between the winter barley variety ‘Tiffany’ and PI284752. This cross demonstrated a two-locus model of resistance. Linkage analysis using polymorphic DNA markers was carried out on 180 plants. The RGH1a gene sequence from the Mla locus was used as a source for developing the RGH1aE2I2 marker. By interval mapping on chromosome 1HS, one resistance gene was found to be tightly linked with RGH1aE2I2 and it was found to be located 2 cM from GBMS062. In F₂ plants exhibiting resistance reaction type (RT) 0, specific DNA fragments for the RGH1aE2I2 marker were amplified. In plants with RT1 to RT2-3, the resistance was conferred exclusively by the second R gene that we identified, which is linked with Bmac0134 and GBMS247 on chromosome 2HS. The aforementioned markers may be valuable candidates for marker-assisted selection of resistant genotypes conferred by one or both genes.     

Quantitative trait loci for dry matter digestibility and particle size traits in two-rowed × six-rowed barley population

More than half of the barley grown in the USA is used for livestock feed, with the remaining stocks diverted for human food and malting purposes. The use of barley grain as a major source of cattle feed has been criticized because of its rapid digestion in the rumen, which can result in digestive disorders in cattle. In sacco dry matter digestibility (ISDMD) and particle size (PS) after dry rolling have been found to play a role in the feedlot performance of barley as a feed grain. Reducing the rate of ISDMD is predicted to result in significantly improved animal health and average daily gain. A recombinant inbred line population derived from a cross between a high ISDMD, two-rowed barley cultivar (Valier) and a six-rowed Swiss landrace line (PI370970) exhibiting far slower ISDMD has been developed for studying the underlying genetic locations and mechanisms of these traits. To detect associated quantitative trait loci (QTLs), we collected and analyzed data from irrigated and rain-fed environments. A significant negative correlation was observed between ISDMD and PS. High heritability estimates for ISDMD and PS suggest that early selection for these traits during breeding would be achievable. Four QTLs were identified on chromosomes 2H, 6H, and 7H, explaining 73–85% of ISDMD phenotypic variation, while three QTLs on 2H and 7H were associated with variation in PS and explained 58–77% of its variation. A major QTL on chromosome 2H tightly linked to the morphology-modifying gene vrs1 was found to dramatically control 35–62% of the phenotypic variation of ISDMD and 26–53% of that of PS. The impact of the vrs1 locus on ISDMD was validated in two populations representing different genetic backgrounds. Our results suggest that it may also be advantageous to simultaneously overlap these QTLs around the vrs1 locus.     

Differentiation of resistance genes at the M1-a locus in six pairs of isogenic barley lines

Summary Five pairs of lines, nearly isogenic except for genes for resistance and susceptibility to culture CR3 of Erysiphe graminis (DC). Merat hordei Em. Marchal, developed from the barley (Hordeum vulgare L.) varieties Algerian, Franger, Durani, Rupee, and Multan, had five different genes at the M1-a locus on chromosome 5 conditioning resistance to culture CR3. The resistant isogenic lines developed from Durani, Rupee, and Multan each had one additional resistance gene, closely linked in coupling to their resistance gene at the M1-a locus, conditioning resistance to culture 63.5 or 64.54. The sixth pair of isogenic lines, developed from the variety Long Glumes, had three resistance genes: one at the M1-a locus, a second closely linked in coupling to the first, and a third gene. The first and the third genes in Long Glumes appear to be the same as the two genes in Multan. The outstanding resistance to E. graminis hordei of most of the donor varieties for the isogenic lines is explained by their possessing two or more resistance genes. The presence of approximately 20 different genes in and around the M1-a locus suggests that there is a cluster of closely linked resistance loci in that region of chromosome 5.Contribution from Applied Plant Genetics Laboratory, Northeastern Region, Agricultural Research Service, U.S. Department of Agriculture, in cooperation with the Agricultural Research Department, Danish Atomic Energy Commission.     

Identification and genetic mapping of a powdery mildew resistance gene in wild emmer (<Emphasis Type="Italic">Triticum dicoccoides</Emphasis>) accession IW72 from Israel

Powdery mildew, caused by Blumeria graminis f. sp. tritici (Bgt), is a devastating disease of wheat (Triticum aestivum) in China and worldwide, causing severe yield losses annually. Wild emmer (T. dicoccoides) accession IW72 collected from Israel is resistant to powdery mildew at the seedling and adult stages. Genetic analysis indicated that the resistance was controlled by a single dominant gene, temporarily designated MlIW72. The F₂ population and F₃ families derived from a hybrid between IW72 and susceptible durum wheat line Mo75 were used for molecular mapping of the resistance gene. MlIW72 was linked with SSR loci Xgwm344, Xcfa2040, Xcfa2240, Xcfa2257 and Xwmc525 on the long arm of chromosome 7A. In addition, two STS markers, MAG2185 (derived from RFLP marker PSR680) and MAG1759 (developed from EST CD452874), were mapped close to MlIW72. All these markers were physically located in the terminal bin 0.86–1.00 of 7AL. The chromosome location and genetic mapping results suggested that the powdery mildew resistance gene identified in wild emmer accession IW72 might be a new allele at the Pm1 locus or a new locus closely linked to Pm1.     

Inter-subspecific maps of non-brittle rachis genes btr1/btr2 using occidental, oriental and wild barley lines

Brittle rachis of wild barley is controlled by two dominant complementary genes, Btr1 and Btr2, and mutation in either locus (btr1 or btr2) results in the non-brittle rachis of cultivated barley. In this study, a simple monogenic inheritance of non-brittle rachis was demonstrated, and moreover differentiation of multiple dominant alleles for either Btr1 or Btr2 among cultivated and wild barley lines was suggested. Two amplified-fragment-length polymorphism (AFLP) linkage maps of the genes were constructed using wild × btr1-type cultivar and wild × btr2-type cultivar F₂ populations. The order of AFLPs and the btr1/btr2 locus was constant between the wild × cultivar maps and a cultivar × cultivar map previously constructed. No suppression of recombination due to the inter-subspecific crosses was noticed in the interval studied. The btr1 locus and all AFLP loci were separated in the wild × btr1-type cultivar F₂ map, but the btr2 locus and eight AFLP loci did not recombine in the wild × btr2-type cultivar F₂ map, thus slightly different levels of affinity between parental cultivars with the wild line was suggested at the btr1/btr2 locus.     

Allozymes and growth habit of Aegilops tauschii: genetic control and linkage patterns

Aegilops tauschii line of spring type growth habit with theearliest heading among all the VIR world germplasm collection of thisspecies was crossed with three Ae. tauschii lines of winter type growthhabit with low, intermediate and the highest vernalization requirement. 12enzyme loci were involved in genetic analysis. The growth habit was foundto be encoded by single codominant major gene, Vrn-D2. Thefollowing linkages were found: Est5 – Nadhd2 in chromosome 3, Vrn-D2 – Aco2 – Cat2 and Pgm – Nadhd1 in chromosome 4, Est2 – Got2 in chromosome 6.     

Genes for scald resistance from wild barley (Hordeum vulgare ssp spontaneum) and their linkage to isozyme markers

Summary Accessions of Hordeum vulgare ssp. spontaneum, the wild progenitor of barley, collected in Israel (70), Iran (15) and Turkey (6) were screened for seedling response to four isolates of Rhynchosporium secalis, the pathogen causing leaf scald in barley. Resistance was very common in the collection (77%) particularly among accessions from the more mesic sites (90%). The genetics of this resistance were investigated in fifteen backcross (BC₃) lines that contained an isozyme variant from H.v. ssp. spontaneum in a H.v. ssp. vulgare (cv. Clipper) background and were resistant to scald. Segregation in the BC₃F₂ families conformed with a single dominant resistance gene in 9 of the 15 lines. Scald resistance and the isozyme marker were closely linked in three of the BC₃-lines, loosely linked in four and unlinked in the remaining eight. Scald resistance genes were identified on barley chromosomes 1, 3, 4 and 6. Crosses between several of the scald resistant BC-lines together with the linkage data indicated that at least five genetically independent resistances are available for combining together for deployment in barley. The linkage of scald resistance in several BC₃-lines to the isozyme locus Acp2 is of special interest as this locus is highly polymorphic in wild barley.     

Mapping of a gene for leaf scald resistance in barley line 'B87/14' and validation of microsatellite and RFLP markers for marker-assisted selection

Seedlings of the barley line ‘B87/14’ were resistant to 22 out of 23 Australian isolates of Rhynchosporium secalis, the causal agent of leaf scald.‘B87/14’‐based populations were developed to determine the location of the resistance locus. Scald resistance segregated as a single dominant trait in BC₁F₂ and BC₁F₃ populations. Bulked segregant analysis identified amplified fragment length polymorphisms (AFLPs) with close linkage to the resistance locus. Fully mapped populations not segregating for scald resistance located these AFLP markers on chromosome 3H, possibly within the complex Rrs1 scald locus. Microsatellite and restriction fragment length polymorphism markers adjacent to the AFLP markers were identified and validated for their linkage to scald resistance in a second segregating population, with the closest marker 2.2 cM from the resistance locus. These markers can be used for selection of the Rrs.B87 scald‐resistance locus, and other genes at the chromosome 3H Rrs1 locus.