Barley Genetics Newsletter (2005) 35:103-143

REPORTS OF THE COORDINATORS

Overall coordinator’s report

Udda Lundqvist

SvalöfWeibull AB

SE-268 81 Svalöv, Sweden

Since the latest overall coordinator’s report in Barley Genetics Newsletter Volume 34, not too many changes of the coordinators have been reported. I do hope that most of you are willing to continue with this work and provide us with new important information and literature search in the future. In the meantime a replacement was found for Chromosome 3H, namely Luke Ramsey, Cell and Molecular Genetics Department, Scottish Crop Research Institute, Invergowrie, Dundee, DD2 5DA, United Kingdom. Please observe some address changes have taken place since the last volume of BGN.

The report of the ’Barley Genetic Linkage Workshop’ from the 9th International Barley Genetics Symposium in Brno, Czech Republic, 2004, is published in this volume. As it became decided that the current system and trait coordination should continue but with a view towards whole genome coordination, Bill Thomas and Dave Marshall from the Scottish Crop Research Institute, Invergowrie, Dundee, UK, are investigating the potential of modernizing the overall system and integrating all types of current and historic data collections into a single, combined database. More details about this subject are found in the Workshop report in this volume.

Revised and new descriptions of barley genes will be published in this current volume. Also revised lists of BGS descriptions by BGS numbers (Table 2) and by locus symbols in alphabetic order (Table 3) will be republished in this issue. Rules for Nomenclature and Gene Symbolization in Barley with the changed and additional amendments will again be published in this volume.

The AceDB database for ’Barley Genes and Barley Genetic Stocks’ is updated continouosly and some more images are added. Also the germplasm part is under revision.

List of Barley Coordinators

Chromoosome 1H (5): Gunter Backes, Department of Agricultural Sciences, The Royal Vetenary and Agricultural University, Thorvaldsensvej 40, DK-1871 Fredriksberg C, Denmark. e-mail: <guba@kvl.dk>

List of Barley Coordinators (continued)

Chromosome 2H (2): Jerry. D. Franckowiak, Department of Plant Sciences, North Dakota State University, P.O.Box 5051, Fargo, ND 58105-5051, USA. FAX: +1 701 231 8474; e-mail: <j.franckowiak@ndsu.nodak.edu>

Chromosome 3H (3): Luke Ramsey, Cell and Molecular Genetics Department, Scottish Crop Research Institute, Invergowrie, Dundee, DD2 5DA, United Kingdom. FAX: +44 1382 562426. e-mail: <Luke.Ramsey@scri.sari.ac.uk>

Chromosome 4H (4): Brian P. Forster, Cell and Molecular Genetics Department, Scottish Crop Research Institute, Invergowrie, Dundee, DD2 5DA, United Kingdom. FAX: +44 1382 562426. e-mail: <bforst@scri.sari.ac.uk>

Chromosome 5H (7): George Fedak, Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, ECORC, Ottawa, ON, Canada K1A 0C6, FAX: +1 613 759 6559; e-mail: <fedakga@agr.gc.ca>

Chromosome 6H (6): Duane Falk, Department of Crop Science, University of Guelph, Guelph, ON, Canada, N1G 2W1. FAX: +1 519 763 8933; e-mail: <dfalk@uoguelph.ca>

Chromosome 7H (1): Lynn Dahleen, USDA-ARS, State University Station, P.O. Box 5677, Fargo, ND 58105, USA. FAX: + 1 701 239 1369; e-mail: <DAHLEENL@fargo.ars.usda.gov>

Integration of molecular and morphological marker maps: Andy Kleinhofs, Department of Crop and Soil Sciences, Washington State University, Pullman, WA 99164-6420, USA. FAX: +1 509 335 8674; e-mail: <andyk@wsu.edu>

Barley Genetics Stock Center: An Hang, USDA-ARS, National Small Grains Germplasm Research Facility, 1691 S. 2700 W., Aberdeen, ID 83210, USA. FAX: +1 208 397 4165; e-mail: <anhang@uidaho.edu>

Trisomic and aneuploid stocks: An Hang, USDA-ARS, National Small Grains Germplasm Research Facility, 1691 S. 2700 W., Aberdeen, ID 83210, USA. FAX: +1 208 397 4165; e-mail: <anhang@uidaho.edu>

Translocations and balanced tertiary trisomics: Andreas Houben, Institute of Plant Genetics and Crop Plant Research, Corrensstrasse 3, DE-06466 Gatersleben, Germany. FAX: +49 39482 5137; e-mail: <houben@ipk-gatersleben.de>

Desynaptic genes: Andreas Houben, Institute of Plant Genetics and Crop Plant Research, Corrensstrasse 3, DE-06466 Gatersleben, Germany. FAX: +49 39482 5137; e-mail: <houben@ipk-gatersleben.de>

List of Barley Coordinators (continued)

Autotetraploids: Wolfgang Friedt, Institute of Crop Science and Plant Breeding, Justus-Liebig-University, Heinrich-Buff-Ring 26-32, DE-35392 Giessen, Germany. FAX: +49 641 9937429; e-mail: <wolfgang.friedt@agrar.uni-giessen.de>

Disease and pest resistance genes: Brian Steffenson, Department of Plant Pathology, University of Minnesota, 495 Borlaug Hall, 1991 Upper Buford Circle, St. Paul, MN 55108-6030, USA. FAX: +1 612 625 9728; e-mail: <bsteffen@umn.edu>

Eceriferum genes: Udda Lundqvist, Svalöf Weibull AB, SE-268 81 Svalöv, Sweden. FAX:.+46 418 667109; e-mail: <udda@ngb.se>

Chloroplast genes: Mats Hansson, Department of Biochemistry, Lund University, Box 124, SE-221 00 Lund, Sweden. FAX: +46 46 222 4534 e-mail: <mats.hansson@biokem.lu.se>

Genetic male sterile genes: Mario C. Therrien, Agriculture and Agri-Food Canada, P.O. Box 1000A, R.R. #3, Brandon, MB, Canada R7A 5Y3, FAX: +1 204 728 3858; e-mail: <MTherrien@agr.gc.ca>

Ear morphology genes: Udda Lundqvist, Svalöf Weibull AB, SE-268 81 Svalöv, Sweden. FAX: +46 418 667109; e-mail: <udda@ngb.se> and

Antonio Michele Stanca: Istituto Sperimentale per la Cerealicoltura, Sezione di Fiorenzuola d’Arda, Via Protaso 302, Fiorenzuola d’Arda (PC), IT-29017, Italy. FAX: +39 0523 983750, e-mail: <michele@stanca.it>

Semi-dwarf genes: Jerry D. Franckowiak, Department of Plant Sciences, North Dakota State University, P.O. Box 5051, Fargo, ND 58105-5051, USA. FAX: +1 702 231 8474; e-mail: <j.franckowiak@ndsu.nodak.edu>

Early maturity genes: Udda Lundqvist, Svalöf Weibull AB, SE-268 81 Svalöv, Sweden. FAX: +46 418 667109; e-mail: <udda@ngb.se>

Biochemical mutants - Including lysine, hordein and nitrate reductase: Andy Kleinhofs, Department of Crop and Soil Sciences, Washington State University, Pullman, WA 99164-6420, USA. FAX: +1 509 335 8674; e-mail: <andyk@wsu.edu>

Barley-wheat genetic stocks: A.K.M.R. Islam, Department of Plant Science, Waite Agricultural Research Institute, The University of Adelaide, Glen Osmond, S.A. 5064, Australia. FAX: +61 8 8303 7109; e-mail: <rislam@waite.adelaide.edu.au>

Coordinator’s Report: Barley Chromosome 1H (5)

Gunter Backes

The Royal Veterinary and Agricultural University

Department of Agricultural Sciences

Thorvaldsensvej 40
DK-1871 Frederiksberg C, Denmark

e-mail: guba@kvl.dk

Grewal et al., (2005) localised a locus for resistance against covered smut caused by Ustilago hordei indirectly to the chromosome arm 1HS. Earlier (Ardiel et al., 2002), this locus was found to be linked with the RAPD-marker OPJ10₄₅₀ that was converted into a SCAR marker (UhR 450). This SCAR marker was then localized on chromosome 1HS.

Rajasekaran et al., (2004) localised QTLs in a 184 RI line population of a cross between the spring barley varieties ’Tankard’ and ’Livet’. They detected one QTL for ‘kernel splitting’, ‘milling energy’, ‘sieve fraction > 2.5 mm’, grain shape, ‘gape between lemma and palea’ and kernel weight between the loci Bmag504 and HvBDG. This QTL had the highest LOD score within the detected QTLs in for kernel splitting and milling energy.

In a DH population (136 lines) from cross between a winter barley (‘Nure’) and spring barley variety (‘Tremois’) winter hardiness and heading date with and without vernalization were determined (Francia et al., 2004). One QTL for heading date with vernalization was found on chromosome 1H nearby the microsatellite marker Bmac0032.

One QTL for kernel weight on the long arm of chromosome 1H and one for kernel number per area on the short arm of chromosome 1H were detected by Verhoeven et al., (2004) in 140 F_2:3-families from the cross of two H. vulgare ssp. spontaeum accessions from contrasting habitats. The aim of the study was to determine QTLs for fitness traits leading to adaptation and population differentiation. The linkage map was purely based on AFLP markers. Based on the same population, Elberse et al., (2004) published a further analysis with growth chamber experiments under high and low nutrient level. On chromosome 1H they detected one QTL for leaf length under high nutrient conditions and a large overlapping area with QTLs for seed mass, leaf length under low nutrient conditions and ‘leaf mass fraction’ and ‘leaf mass area’ both under high and low nutrient conditions.

Chen et al.,(2004) determined the association between SSR marker and kernel weight and kernel colour in wild barley (H. vulgare ssp. spontaneum) populations in Israel. They found associations between the marker loci BMS90 and HVM43.

Edney and Mather (2004) detected two minor QTL for malt friability on chromosome 1H. One was close to the marker locus ABG452, the other one near the marker locus cMWG706A. They carried out the QTL analysis in the ‘Harrington’ x ‘Morex’ population (140 doubled haploid lines).

In order to reveal the relation between grain protein content and malting quality, Emebiri et al., (2004) investigated QTLs for different malting related traits in a double haploid population (180 lines) from a cross between a line with extremely low kernel protein content (VB9524) and a line with poor a-amylase activity (NB11231*1). On chromosome 1H, nearby the marker locus XMXwg912, they detected a QTL for b-glucanase activity.

A QTL for non-parasitic leaf spots was detected on chromosome 1H together with a QTL for heading date by Behn et al., (2004). They performed the analysis in a population of 86 doubled haploid lines from a cross between the spring barley line IPZ24727 and the spring barley variety ‘Barke’. IPZ24727 derives from an Israeli wild barley line and possesses good resistance against non-parasitic leaf spots.

Pillen et al., (2004) used the advanced backcross-strategy to localize QTLs for several agronomic traits in a BC₂F₂-population of 164 plants. The wild parent was the H. vulgare ssp. spontaneum-line ISR101-23. The recurrent parent was the spring barley variety ‘Harry’. BC₂F_2:5-families were used for the phenotyping. The ‘Steptoe’ ´ ‘Morex’ linkage map was used for the marker localization. On chromosome 1H, they detected one QTL for number of spikes per area, heading date, lodging at flowering and grain weight linked to the marker locus HVM20 and one for lodging at flowering and grain weight linked to the marker locus HVM36.

Another advanced backcross-experiment including H. vulgare ssp. spontaneum was carried out by Talemè et al., (2004). They analysed the QTLs in a DH population from a BC1F2. The wild barley parent was the line HOR11508, the recurrent parent was the variety ‘Barke’. They describe two putative QTLs on chromosome 1H: one for heading date, plant height, ear extrusion and grain yield linked to the marker locus Bmac154 and one for growth habit and heading date linked to the marker locus WMCIE8.

By hybridizing DNA probes with DNA from wheat-barley addition lines, Spielmeyer et al., (2004) localized a 2-oxidase (Hv3ox2) from the gibberellin metabolic pathway to the long arm of chromosome 1H. In wheat, the probe hybridised with the chromosomes 1A, 1B and 1D. The orthologous rice gene (Os3ox2) is located on chromosome 1S of rice.

References

Ardiel, G. S., T. Grewal, P. Deberdt, B. G. Rossnagel, and G. J. Scoles. 2002. Inheritance of resistance to covered smut in barley and development of a tightly linked SCAR marker. Theor. Appl. Genet. 104(2-3):457-464.

Behn, A., L. Hartl, G. Schweizer, and G. Wenzel. 2004. QTL mapping for resistance against non-parasitic leaf spots in a spring barley doubled haploid population. Theor. Appl. Genet. 108(7):1229-1235.

Chen, G. X., T. Suprunova, T. Krugman, T. Fahima, and E. Nevo. 2004. Ecogeographic and genetic determinants of kernel weight and colour of wild barley (Hordeum spontaneum) populations in Israel. Seed Sci. Res. 14(2):137-146.

Edney, M. J., and D. E. Mather. 2004. Quantitative trait loci affecting germination traits and malt friability in a two-rowed by six rowed barley cross. J. CerealSci. 39(2):283-290.

Elberse, I. A. M., T. K. Vanhala, J. H. B. Turin, P. Stam, J. M. M. van Damme, and P. H. van Tienderen. 2004. Quantitative trait loci affecting growth-related traits in wild barley (Hordeum spontaneum) grown under different levels of nutrient supply. Heredity 93(1):22-33.

Emebiri, L. C., D. B. Moody, J. F. Panozzo, and B. J. Read. 2004. Mapping of QTL for malting quality attributes in barley based on a cross of parents with low grain protein concentration. Field Crops Res. 87(2-3):195-205.

Francia, E., F. Rizza, L. Cattivelli, A. M. Stanca, G. Galiba, B. Tóth, P. M. Hayes, J. S. Skinner, and N. Pecchioni. 2004. Two loci on chromosome 5H determine low-temperature tolerance in a 'Nure' (winter) x 'Tremois' (spring) barley map. Theor. Appl. Genet. 108(4):670-680.

Grewal, T., B. G. Rossnagel, and G. J. Scoles. 2005. Mapping of a covered smut resistance gene in barley (Hordeum vulgare). Can. J. Plant Pathol. 26(2):156-166.

Pillen, K., A. Zacharias, and J. Léon. 2004. Comparative AB-QTL analysis in barley using a single exotic donor of Hordeum vulgare ssp spontaneum. Theor. Appl. Genet. 18(8):1591-1601.

Rajasekaran, P., W. T. B. Thomas, A. Wilson, P. Lawrence, G. Young, and R. P. Ellis. 2004. Genetic control over grain damage in a spring barley mapping population. Plant Breed. 123(1):17-23.

Spielmeyer, W., M. Ellis, M. Robertson, S. Ali, J. R. Lenton, and P. M. Chandler. 2004. Isolation of gibberellin and metabolic pathway genes from barley and comparative mapping in barley, wheat and rice. Theor. Appl. Genet. 109(4):847-855.

Talamè, V., M. C. Sanguineti, E. Chiapparino, H. Bahri, M. Ben Salem, B. P. Forster, R. P. Ellis, S. Rhouma, W. Zoumarou, R. Waugh, and R. Tuberosa. 2004. Identification of Hordeum spontaneum QTL alleles improving field performance of barley grown under rainfed conditions. Ann. Appl. Biol. 144(3):309-319.

Verhoeven, K. J. F., T. K. Vanhala, A. Biere, E. Nevo, and J. M. M. van Damme. 2004. The genetic basis of adaptive population differentiation: A quantitative trait locus analysis of fitness traits in two wild barley populations from contrasting habitats. Evolution 58(2):270-283.

Coordinator’s report: Chromosome 2H (2)

J.D. Franckowiak

Department of Plant Sciences

North Dakota State University

Fargo, ND 58105, USA.

e-mail: j.franckowiak@ndsu.nodak.edu

Turuspekov et al., (2004) described and mapped to genes associated with closed flowering in barley. The cleistogamy 1 (cly1) and cleistogamy 2 (Cly2) genes were mapped to loci in the same region of chromosome 2HL near molecular marker MSU21. Plants classified as closed flowering or cleistogamy did not extrude anthers during or after anthesis.

Reinheimer et al., (2004) identified QTL for resistance to frost induced floret sterility in chromosomes 2HL and 5HL. The 2HL QTL maps in the same region as the cly1 gene (Turuspekov et al., 2004). The 5HL QTL is near the Vrn-H1 or Srh2 (spring growth habit 2) locus. A second QTL in 2H, probably the Ppd-H1 or Eam1 gene, was associated with early heading date and escape from frost damage.

Korff et al., (2004) reported that introgression of the Ppd-H1 or Eam1 segment of chromosome 2HS from H. vulgare ssp. spontaneum into European barley cultivars had more effect on heading date and other agronomic traits than other introgressed segments. Daib et al., (2004) reported then this same region of chromosome 2H was associated with QTL for several physiological measurements of drought stress tolerance. Li et al., (2005) reported similar results using backcross-derived lines. Chromosome 2HS was associated with heading date, plant height, yield, lodging, ear length, grain per spike, 1000-kernel weight, and grain protein.

Karsai et al., (2004) reported on heading date variations of Hordeum vulgare ssp. spontaneum accessions caused by photoperiod differences. Vernalized and non-vernalized plants were grown in grown chambers and exposed to various constant day lengths under a constant temperature condition. Early heading under long photoperiods was attributed primarily to the effects of a factor on chromosome 2H, presumably the PpdH1 or Eam1 locus. However, a large number of other genetic factors contributed to the range of responses observed.

Spielmeyer et al., (2004) reported on the barley genes in the metabolic pathway for gibberellic acid (GA) in barley. Characterization of genes involved in GA biosynthesis and its stimulation of cell elongation in barley, wheat and rice is considered the first in determining whether dwarfing genes in barley involve defective GA metabolism. Eleven genes potentially account for the six enzymes in the core GA biosynthetic pathway. Three (HvKSL1, HvKSL2, and Hv3ox1) of those loci were mapped in chromosome 2H.

He et al., (2004) examined molecular markers closely linked to the vrs1 (six-rowed spike 1) locus in chromosome 2HL and reported on progress in positional cloning of alleles at the vrs1 locus using amplified fragment length polymorphism (AFLP) markers and their converted sequence tagged sites (STSs) to screen a bacterial artificial chromosome (BAC library.

Tanno and Takeda, 2004 and Casas et al., (2005) studied the evolution of barley using markers near the vrs1 locus. Tanno and Takeda, 2004, analyzed genetic diversity at the MWG699 marker locus in 10 H. vulgare ssp. spontaneum accessions, 42 six-rowed brittle barley accessions, and 14 six-rowed cultivars. Nine sequence types were found among the H. vulgare ssp. spontaneum accessions, three in brittle barleys, and three in the cultivars. Since the same three sequences were found in the brittle and cultivated barleys, Tanno and Takeda, 2004 concluded that H. vulgare ssp. vulgare f. agriocrithon from the Central Asia is not wild barley, but a weedy outcross from cultivated six-rowed barley. Casas et al., 2005 studied 257 cultivated barleys from the western Mediterranean region using the STS marker MWG699/Taq1. Most two-rowed cultivars had the type K allele. The type D allele was wide spread among winter six-rowed landraces from Spain and cultivars from central Europe. The type A allele was found in both spring and winter six-rowed cultivars. These conclusions agree with information reported by Tanno and Takeda, 2005.

References:

Casas, A.M., S. Yahiaoui, F. Ciudad, and E. Igartua. 2005. Distribution of MWG699 polymorphism in Spanish European barleys. Genome 48:41-45.

Diad, A.A., B. Teulat-Merah, D. This, N.Z. Ozturk, D. Benscher, and M.E. Sorrels. 2004. Identification of drought-induced genes and differentially expressed sequence tags in barley. Theor. Appl. Genet. 109:1417-1425.

He, C., B.E. Sayed-Tabatabael, and T. Komatsuda. 2004. AFLP targing of the 1-cM region conferring the vrs1 gene for six-rowed spike in barley, Hordeum vulgare L. Genome 47:1122-1129.

Karsai, I., P.M. Hayes. J. Kling, I.A. Matus, K. Mészáros, L. Láng, Z. Bedőand K. Sato. 2004. Genetic variation in component traits of heading date in Hordeum vulgare subsp. spontaneum accessions characterized in controlled environments. Crop Sci. 44:1622-1632.

Korff, M. von, H. Wang, J. Léon, and K. Pillen. 2004. Development of candidate introgression lines using an exotic barley accession (Hordeum vulgare spp. spontaneum) as donor. Theor. Appl. Genet. 109:1736-1745.

Li, J.Z., X.Q. Huang, F. Heinrichs, M.W. Ganal, and M.S. Röder. 2005. Analysis of QTLs for yield, yield components, and malting quality in a BC₃-DH population of spring barley. Theor. Appl. Genet. 110:356-363.

Reinheimer, J.L., A.R. Barr, and J.K. Eglinton. 2004. QTL mapping of chromosomal regions conferring reproductive frost tolerance in barley (Hordeum vulgare L.) Theor. Appl. Genet. 109:1267-1274.

Spielmeyer, W., M. Ellis, M. Robertson, S. Ali, J.R. Lenton, and P.M. Chandler. 2004. Isolation of gibberellin metabolic pathway genes from barley and comparative mapping in barley, wheat and rice. Theor. Appl. Genet. 109:847-855.

Tanno, K., and K. Takeda. 2004. On the origin of six-rowed barley with brittle rachis, agriocrithon [Hordeum vulgare ssp. vulgare f. agriocrithon (Åberg) Bowd.], based on a DNA marker closely linked to the vrs1 (six-row gene) locus. Theor. Appl. Genet. 110:145-150.

Turuspekov, Y., Y. Mano, I. Honda, N. Kawada, Y. Watanabe, and T. Komatsuda. 2004. Identification and mapping of cleistogamy genes in barley. Theor. Appl. Genet. 109:48-487.

Coordinator’s Report: Barley Chromosome 3H.

L. Ramsay

Cell and Molecular Genetics Department

Scottish Crop Research Institute

Invergowrie, Dundee, DD2 5DA, Scotland, UK.

e-mail: Luke.Ramsey@scri.sari.ac.uk

Since the last co-ordinator’s report in BGN 33 there have been a number of publications reporting the mapping of genes and in particular QTL on barley chromosome 3H. Of particular note is the reporting by Chono et al., (2003) of the cloning and functional characterisation of the semi-dwarfing gene uzu on 3HL in Bin6. The authors report that the semi-dwarf phenotype arises from a mutation in a gene encoding a putative brassinosteroid receptor that is possibly homologous to a known rice mutant d61 (Chono et al., 2003).

In contrast synteny with rice did not prove so informative for the detailed molecular mapping of leaf rust resistance gene Rph7 on the distal end of 3HS close to the RFLP marker MWG848 in Bin1 (Brunner et al., 2003). The region genetically delineated as containing the Rph7 locus contained six genes in barley in the cultivar Morex that were not present on the homologous region on rice chromosome 1. Interestingly the characterisation of the same region in the resistance variety Cepada Capa indicated that the colinearity between the barley varieties was restricted to only five genic and two intergenic regions representing less than 35% of the two sequences. The differences were mainly due to the presence of different transposable elements in the intergenic regions but also included the loss of a gene in Cepada Capa (Scherrer et al., 2005).

Two other leaf rust resistance genes Rph5 and Rph6 were also mapped to the distal end of 3HS (Mammadov et al., 2003, Zhong et al., 2003). Detailed mapping work indicated that Rph5 was positioned in the extreme telomeric region of 3HS distal to Rph7 (Mammadov et al., 2003) and that Rph6, maps to a similar location. Indeed segregation analyses indicated that Rph6 is allelic to Rph5 (Zhong et al., 2003). Pellio et al. (2005) report the high-resolution mapping of the Rym4/Rym5 locus conferring resistance to the barley yellow mosaic virus complex (BaMMV, BaYMV, BaYMV-2) on the distal end of the long arm of 3H that has allowed the identification of a candidate gene that has since been confirmed to be involved in the bymovirus resistance (Stein et al., 2005).

Several studies reported QTL for disease resistances on 3H including to scald (Patil et al., 2002, Genger et al., 2003a, 2003b, Bjørnstad et al., 2004, Sayed et al., 2004), net blotch (Cakir et al., 2003a, Raman et al., 2003), leaf stripe (Arru et al., 2003) and powdery mildew (Backes et al., 2003). A new scald resistance gene Rrs4_CI11549 was mapped on 3H located 22cM distally on the long arm corresponding to the region Bin 8-9 (Patil et al., 2003) and the crown rust resistance gene Rpc1 was mapped 6cM distal to the SSR marker Bmag0006 to the long arm of 3H in the Bin 6 region (Agrama et al., 2004).

Quantitative Trait Loci that mapped to chromosome 3H were also reported for hull cracked grain (Kai et al., 2003), kernel discolouration (Li et al., 2003), malt friability (Edney and Mather, 2004) grain shape and damage (Rajasekaran, 2004) as well a range of other agronomic and quality traits (Baum et al., 2003, Cakir et al., 2003b, Collins et al., 2003, Read et al., 2003, Talame et al., 2004). Komatsuda et al., (2004) reported high density mapping of the non-brittle rachis 1 (btr1) and 2 (btr2) genes on 3HS in Bin5. The report confirms the tight linkage of the two loci and a phylogenetic tree based on AFLP markers linked to the genes showed clear separation of occidental and oriental barley lines.

The publications referred to above are in no way an exhaustive list of recently mapped 3H loci which is a reflection of the breadth and vitality of the barley genetics community. Two presentations at the IX. International Barley Genetics Symposium on large scale mapping of barley ESTs highlight the difficulty now in reviewing the mapping of barley genes. Sato et al., (2004) presented results of their EST mapping work that included the novel mapping of 163 genes to 3H and Graner et al., (2004) presented the relationship they had found between high density gene maps of barley 3H and rice chromosome 1 extending the work of Smilde et al., (2001). However neither sets of genetic maps are as yet in the public domain unfortunately but the reports are indicative of the scale of mapping now possible and the increasing importance of barley’s syntenic relationship with the sequenced genome of rice.

References:

Agrama, H.A., L. Dahleen, M. Wentz, Y. Jin, and B. Steffenson. 2004. Molecular mapping of the crown rust resistance gene Rpc1 in barley. Phytopathology 94 (8):858-861.

Arru, L., E. Francia, and N. Pecchioni. 2003. Isolate-specific QTLs of resistance to leaf stripe (Pyrenophora graminea) in the 'Steptoe' x 'Morex' spring barley cross. Theor. Appl. Genet. 106 (4):668-675.

Backes, G., L. H. Madsen, H. Jaiser, J. Stougaard, M. Herz, V. Mohler, and A. Jahoor. 2003. Localisation of genes for resistance against Blumeria graminis f.sp hordei and Puccinia graminis in a cross between a barley cultivar and a wild barley (Hordeum vulgare ssp spontaneum) line. Theor. Appl. Genet. 106 (2):353-362.

Baum, M., S. Grando, G. Backes, A. Jahoor, A. Sabbagh, and S. Ceccarelli. 2003. QTLs for agronomic traits in the Mediterranean environment identified in recombinant inbred lines of the cross ‘Arta’ x H-spontaneum 41-1.Theor. Appl. Genet. 107 (7):1215-1225.

Bjørnstad, A., S. Grønnerød, J. MacKey, A. Tekauz, J. Crossa, and H. Martens. 2004. Resistance to barley scald (Rhynchosporium secalis) in the Ethiopian donor lines 'Steudelli' and 'Jet', analyzed by partial least squares regression and interval mapping. Hereditas 141 (2):166-179.

Brunner, S., B. Keller, and C. Feuillet. 2003. A large rearrangement involving genes and low copy DNA interrupts the microcolinearity between rice and barley at the Rph7 locus. Genetics 164:673-683.

Cakir, M., D. Poulsen, N.W. Galwey, G.A. Ablett, K.J. Chalmers, G.J. Platz, R.F. Park, R.C.M. Lance, J.F. Panozzo, B.J. Read, D.B. Moody, A.R. Barr, P. Johnston, C.D. Li, W.J.R. Boyd, C.R. Grime, R. Appels, M.G.K. Jones, and P. Langridge. 2003a. Mapping and validation of the genes for resistance to Pyrenophora teres f. teres in barley (Hordeum vulgare L.) Austr. J. Agr. Res. 54 (11-12):1369-1377.

Cakir, M., S. Gupta, G.J. Platz, G.A- Ablett, R. Loughman, L.C. Emebiri, D. Poulsen, C.D. Li, R.C.M. Lance, N.W. Galwey, M.G.K. Jones, and R. Appels. 2003b. Mapping and QTL analysis of the barley population Tallon x Kaputar. Austr. J. Agr. Res. 54 (11-12):1155-1162.

Chono, M., L. Honda, H. Zeniya, K. Yoneyama, D. Saisho, K. Takeda, S. Takatsuto, T. Hoshino, and Y. Watanabe. 2003. A semidwarf phenotype of barley uzu results from a nucleotide substitution in the gene encoding a putative brassinosteroid receptor. Plant Physiology 133:1209-1219.

Collins, H.M., J.F. Panozzo, S.J. Logue, S.P. Jefferies, and A.R. Barr. 2003. Mapping and validation of chromosome regions associated with high malt extract in barley (Hordeum vulgare L.). Austr. J. Agr. Res. 54 (11-12):1223-1240.

Edney, M.J., and D.E. Mather. 2004. Quantitative trait loci affecting germination traits and malt friability in a two-rowed by six rowed barley cross. J. Cereal Sci. 39 (2):283-290.

Genger, R.K., A.H.D. Brown, W. Knogge, K. Nesbitt, and J.J. Burdon. 2003a. Development of SCAR markers linked to a scald resistance gene derived from wild barley. Euphytica 134 (2):149-159.

Genger, R.K., K.J. Williams, H. Raman, B.J. Read, H. Wallwork, J.J. Burdon, and A.H.D. Brown. 2003b. Leaf scald resistance genes in Hordeum vulgare and Hordeum vulgare ssp spontaneum: parallels between cultivated and wild barley. Austr. J. Agr. Res. 54 (11-12):1335-1342.

Graner, A., R. Kota, D. Perovic, E. Potokina, M. Prasad, U. Scholz, N. Stein, T. Thiel, R.K. Varshney, and H. Zhang. 2004. Molecular mapping: shifting from the structural to the functional level. In: J. Spunar and J. Janikova (eds.), pp 49-57. Barley Genetics IX. Proc. Ninth Int. Barley Genet. Symp., Brno, Czech Republic, June 20-26 2004.(in press).

Kai. H., T. Baba, M. Tsukazaki, Y. Uchimura, and M. Furusho. 2003. The QTL analysis of hull-cracked grain in Japanese malting barley. Breeding Sci 53 (3): 225-230.

Komatsuda, T., P. Maxinm, N. Senthil, and Y. Mano. 2004. High-density AFLP map of nonbrittle rachis 1 (btr1) and 2 (btr2) genes in barley (Hordeum vulgare L.). Theor. Appl. Genet. 109: 986-995.

Li, C.D., R.C.M. Lance, H.M.Collins, A. Tarr, S. Roumeliotis, S. Harasymow, M. Cakir, G.P. Fox, C.R. Grime, S. Broughton, K.J. Young, H. Raman, A.R. Barr, D.B. Moody, and B.J. Read. 2003. Quantitative trait loci controlling kernel discoloration in barley (Hordeum vulgare L.). Austr. J. Agr. Res. 54 (11-12):1251-1259.

Mammadov, J.A., J.C. Zwonitzer, R.M. Biyashev, C.A. Griffey, Y. Jin, B.J. Steffenson, and M.A.S. Maroof. 2003. Molecular mapping of leaf rust resistance gene Rph5 in barley. Crop Sci. 43 (1):388-393.

Patil V., A. Bjørnstad, and J. MacKey. 2003. Molecular mapping of a new gene Rrs4_(CI11549) for resistance to barley scald (Rhynchosporium secalis). Mol. Breeding 12 (2):169-183.

Patil V., A. Bjørnstad, H. Magnus, and J. MacKey. 2002. Resistance to scald (Rhynchosporium secalis) in barley (Hordeum vulgare L.). II. Diallel analysis of near-isogenic lines. Hereditas 137 (3):186-197.

Pellio, B., S. Streng, E. Bauer, N. Stein, D. Perovic, A. Schiemann, W. Friedt, F. Ordon, and A. Graner. 2005. High-resolution mapping of the Rym4/Rym5 locus conferring resistance to the barley yellow mosaic virus complex (BaMMV, BaYMV, BaYMV-2) in barley (Hordeum vulgare ssp vulgare L.). Theor. Appl. Genet. 110:283-293.

Rajasekaran, P., W.T.B. Thomas, A. Wilson, P. Lawrence, G. Young, and R.P. Ellis. 2004. Genetic control over grain damage in a spring barley mapping population. Plant Breeding 123:17-23.

Raman, H., G.J. Platz, K.J. Chalmers, R. Raman, B.J. Read, A.R. Barr, and D.B. Moody. 2003. Mapping of genomic regions associated with net form of net blotch resistance in barley. Austr. J. Agr. Res. 54 (11-12):1359-1367.

Read, B.J., H. Raman, G. McMichael, K.J. Chalmers, G.A. Ablett, G.J. Platz, R. Raman, R.K. Genger, W.J.R. Boyd, C.D. Li, C.R. Grime, R.F. Park, H. Wallwork, R. Prangnell, and R.C.M. Lance. 2003. Mapping and QTL analysis of the barley population Sloop x Halcyon. Austr. J.Agr. Res. 54 (11-12):1145-1153.

Sato, K., N. Nankaku, Y. Motoi, and K. Takeda. 2004. A large scale mapping of ESTs on barley genome. In: J. Spunar and J, Janikova (eds.), pp. 79-85. Barley Genetics IX. Proc.Ninth Int. Barley Genet. Symp., Brno, Czech Republic, June 20-26 2004. (in press).

Sayed, H., G. Backes, H. Kayyal, A. Yahyaoui, S. Ceccarelli, S. Grando, A. Jahoor, and M. Baum. 2004. New molecular markers linked to qualitative and quantitative powdery mildew and scald resistance genes in barley for dry areas. Euphytica 135 (2):225-228.

Scherrer, B., E. Isidore, P. Klein, J-S. Kim, A. Bellec, B. Chalhoub, B. Keller, and C. Feuillet. 2005. Large intraspecific haplotype variability at the Rph7 locus results from rapid and recent divergence in the barley genome. Plant Cell 17:361-374.

Smilde, W.D., J. Haluskova, T. Sasaki, and A. Graner. 2001. New evidence for the synteny of rice chromosome 1 and barley chromosome 3H from rice expressed sequence tags. Genome 44:361-367.

Stein, N., D. Perovic, B. Pellio, J. Kumlehn, T. Wicker, S. Stracke, F. Ordon, and A. Graner. 2005. The gene elF4E is a common determinant of recessive virus resistance in dicot and monocot plants as revealed by map-based cloning of Rym4 conferring bymovirus resistance in barley. Plant and Animal Genomes XIII, San Diego, CA, USA, January 15-19 2005. http://www.intl-pag.org/abstracts/PAG13_W040.html

Talame, V., M.C. Sanguineti, E. Chiapparino, H. Bahri, M. Ben Salem, B.P. Forster, R.P. Ellis, S. Rhouma, W. Zoumarou, R. Waugh, and R. Tuberosa. 2004. Identification of Hordeum spontaneum QTL alleles improving field performance of barley grown under rainfed conditions. Ann. Appl. Biol. 144 (3):309-319.

Zhong, S. B., R. J. Effertz, Y. Jin, J.D. Franckowiak, and B.J. Steffenson. 2003. Molecular mapping of the leaf rust resistance gene Rph6 in barley and its linkage relationships with Rph5 and Rph7. Phytopathology 93 (5):604-609.

Coordinator’s Report: Barley Chromosome 5H(7)

George Fedak

Eastern Cereal & Oilseed Research Centre

Agriculture & Agri-Food Canada

Ottawa, Ontario, K1A 0C6

e-mail: fedakga@agr.gc.ca

Seed dormancy and it’s release during after-ripening are important traits in barley harvesting and the malting industry. The traits are quantitatively inherited and under environmental influence.

In yet another study, a DH mapping population was made from two cultivars with elite malting quality; Triumph (European two row, prone to dormancy) and Morex (North America 6 row, non dormant) (Prada et al., 2004).

The QTL for GP3 (7) (germination percentage at 3 days of incubation at 7 days post harvest) obtained from Triumph was located near the centromere of chromosome 7(5H) and explained 52% of the phenotypic variance. The QTL for GP7 (7) (germination percentage at 7 days of incubation at 7 days post harvest) obtained from Triumph was located in the centromere region of chromosome 7(5H) and explained 33% of the phenotypic variance; a second GP7 (7) QTL from the variety Morex was located at the long arm telomere of chromosomes 3(3H) and explained 13% of phenotypic variation. It is not yet known if the common position of

GP3 (7) and GP7 (7) is due to linkage or pleiotropy.

In the same population a DR QTL (dormancy release through after-ripening) obtained from Morex, was located on the long arm telomere of chromosome 7(5H) and another on chromosome 2(2H) and explained 19 & 9% of phenotypic variability respectively. It is assumed that a moderate level of dormancy could be maintained by manipulating the balance between the GP and DR loci.

It is interesting to note that in previous studies major dormancy QTL, SD1 and SD2 in a Steptoe/Morex cross were located in the centromeric and long arm telomeric regions respectively of chromosome 7(5H) (Han et al., 1996). In addition a major dormancy QTL in the Harrington/TR306 mapping population (Ullrich et. al., 2002) and one in the Chebec/Harrington population (Karakousis et al. 1996) have been mapped to the long arm telomere of chromosome 7(5H). It is suggested that the QTL located in the Triumph/Morex population could be allelic with those detected in the Steptoe/Morex cross and those detected in the Harrington / TR306 and the Chebec/Harrington populations.

Dormancy was also tested in an F₂ population derived from a cross between Triumph and Steptoe, both cultivars with some degree of dormancy. The distribution of the F₂ population was continuous but a large number of transgressive segregants were obtained, indicating that although the two parents come from distinct gene pools, there is probably some difference in the genetic control of dormancy. Minor genes could be affecting the expression of dormancy in both cultivars.

Winter hardiness is a complex trait involving aspects of low temperature tolerance, vernalization requirement and photoperiod sensitivity. To map the QTL controlling some of those traits a DH mapping population consisting of 136 lines was developed from a hybrid between the cultivars Nure (winter two rowed feed barley) x Tremois (spring two rowed malting barley), (Francia et al., 2004).

A total of nine QTL for the various cold-hardiness-related traits were mapped on the long arm of chromosome 5H. These included 2 QTL for winter-field survival, 2 QTL for a controlled field test plus two for functionality of photosystem II. In addition, for COR genes (cold regulated genes) QTL were located on chromosome 5H and 6H, heading date QTL were located on chromosomes 5H, 1H, 2H and 6H and a vernalization QTL was also located on chromosome 5H (long arm).

The vernalization QTL coincided with previously described vernalization loci in the Triticeae and other QTL for all measures of cold hardiness coincided with this locus. In summary all of the QTL reported above coincided with the two major loci on the long arm of chromosome 5H.

References:

Francia, E., F. Rezza, L. Cattivelli, A.M. Stanca, G. Galiba,B. Toth, P.M. Hayes, J.S.Skinner, and N. Pecchioni. 2004. Two loci on chromosome 5H determine low-temperature tolerance in a ‘Nure’ (winter) x Tremois (spring) barley maps. Theor. Appl. Genet. 108:670-680.

Han, F., S.E. Ullrich, J.A. Clancy, V. Jitkou, A Kilian, and I. Ramagosa. 1996. Verification of barley seed dormancy loci via linked molecular markers. Theor. Appl. Genet. 92:87-91.

Ullrich, S. E., F. Han, W. Gao, D, Prada, J.A. Clancy, A. Kleinhofs, I. Ramagosa, and J.L. Molina-Cano. 2002. Summary of QTL analysis of seed dormancy trait in barley. Barley Newsl. 45:39-41.

Karakousis, A, J. Kretschmer, S. Manning, K. Chalmers,and P. Langridge. 1996. The Australian genome mapping project. Available on-line at: http://greengenes. cit. Cornell.edu/Waite QTL.

Prada, D., S.E. Ullrich, J.L. Molina-Cano, L. Cistice, J.A. Clancey, and I. Ramagosa. 2004. Genetic control of dormancy in a Triumph/Morex cross in barley. Theor. Appl. Genet.109:62-70.

Coordinator’s Report: Chromosome 7H.

Lynn S. Dahleen

USDA-Agricultural Research Service

Fargo, ND 58105, USA

e-mail: DAHLEENL@fargo.ars.usda.gov

As usual, progress in gene and QTL mapping covered numerous traits. In addition to the peer-reviewed papers described here, many reports were presented at the International Barley Genetics Symposium in Brno, Czech Republic in June 2004, available in the Proceedings and the book of abstracts published in the Czech Journal of Genetics and Plant Breeding.

New microarray technology has been applied to mapping. (Potokina et al., 2004) used a 1,400 EST array to identify genes for malting quality traits. Functional associations in ten genotypes identified two ESTs on chromosome 7H. HY05O13 for sucrose synthase1 was associated with malt extract and b-amylase activity. HY03C01 for catalase (cat1) was associated with extract, b-amylase, Kolbach index and final attenuation. QTL mapping in the Steptoe x Morex population identified QTLs for extract and a-amylase at HY05O13 but no QTLs at HY03C01.

Rajasekaran et al., (2004) mapped QTLs for grain damage traits in a Tankard x Livet population. Nine QTLs were located in chromosome 7H in 3-4 locations. Traits included sieve fraction greater than 2.5 mm (2 QTLs), skinning less than 25% (2 QTLs), grain width by image analysis, ratio of grain width to length, height and grain milling energy (2 QTLs). The major region on chromosome 7H was at the SSR marker Bmag507.

QTLs affecting germination and malt friability were located by Edney and Mather, (2004), using the Harrington x Morex mapping population. A QTL for germination of 100 seeds in 4 ml of water was found in chromosome 7H by composite interval mapping. Morex provided the favorable allele.

Han et al., (2004) developed 39 isolines from a Steptoe x Morex cross differing for marker genotype in a 28 cM malting quality QTL region in chromosome 7H. QTL analysis after micromalting identified one QTL for malt extract, and two QTLs each for a-amylase activity, diastatic power and malt b-glucan. Resolutions of 2.0 cM or less were achieved, providing good markers for breeding.

Validation of the many QTL regions identified for Fusarium head blight resistance, kernel discoloration and deoxynivalenol concentration has begun with two populations derived from Chevron (Canci et al., 2004). Of the fifteen QTLs identified in the original mapping population, only five were validated in the new populations. None of the QTLs on chromosome 7H were detected in either new population.

Nonbrittle rachis has been a key locus in domestication of barley. Komatsuda et al., (2004) compared nonbrittle rachis loci in occidental and oriental barley lines. In addition to the major gene btr2, the oriental barley contained two QTLs, one of which was located in chromosome 7H.

Diab et al. (2004) mapped 68 QTLs involved in drought tolerance traits using the cross Tadmor x Er/Apm. Ten were located in chromosome 7H, including three for relative water content in stressed and irrigated plots and seven for water soluble carbohydrate concentration traits. Two genes were located in these QTLs, acyl carrier protein III (Acl3) and sucrose synthase (bSS1B), and an EST for a copper binding protein (BM816463).

The new tetra-primer ARMS-PCR technique was used to validate previously identified single nucleotide polymorphisms (SNPs) in five of nine RFLP clones (Chiapparino et al., 2004). Two of these five were located in chromosome 7H, MWG2062 and ABC465. They then screened 132 varieties and determined the frequency of each nucleotide at the polymorphic site. This technique has potential in low to medium throughput laboratories.

Varshney et al., 2004 tested transferability of 165 barley EST-SSR markers to wheat, rye and rice, to expand our knowledge of cereal synteny. Four of the barley chromosome 7H EST-SSR markers tested were homologous to group 7 chromosome wheat ESTs. The barley electronic comparisons to rice showed synteny between eleven of the 21 barley 7H EST-SSRs examined and rice chromosomes 2, 5, 6, 8, 9, and 12. The two chromosome 7H barley EST-SSRs mapped in rye were located on rye chromosome 4RL.

Comparisons of sequence-based polymorphisms in barley EST-derived markers were examined by Russell et al., 2004. The one sequence evaluated in chromosome 7H was Best 1239, with homology to sucrose synthase. Two polymorphic sites were located in landraces, producing two haplotypes. Spring barley cultivars and H. spontaneum both showed no diversity, producing a single haplotype. Markers for other ESTs had as many as nine different haplotypes. In general, cultivated barley showed less diversity than the landraces and H. spontaneum.

Wenzl et al., (2004) developed and tested diversity arrays technology (DarT) using polymorphism-enriched microarray hybridization. They then applied DarT to the Steptoe x Morex mapping population and mapped 42 markers to chromosome 7H. With this technique, it is possible to create a medium density linkage map in a few days.

Fluorescent in situ hybridization with RFLP clones was used for comparing physical and genetic maps by Stephens et al., 2004. One landmark plasmid, p18S5Shor, was used to identify and orient all seven chromosome pairs. Six of the fourteen cDNA clones used mapped to chromosome 7H, Amy2, Brz, Chi, Glx, His3, and Ubi. Physical mapping showed that barley genetic maps do not cover large areas of the genome.

References:

Canci P.C., L.M. Nduulu, G.J. Muehlbauer, R. Dill-Macky, D.C. Rasmusson, and K.P. Smith. 2004. Validation of quantitative trait loci for Fusarium head blight and kernel discoloration in barley. Molec. Breed. 14:91-104.

Chiapparino E., D. Lee. and P. Donini. 2004. Genotyping single nucleotide polymorphisms in barley by tetra-primer ARMS-PCR. Genome 47:414-420.

Diab A.A., B. Teulat-Merah, D. This, N.Z. Ozturk, D. Benscher, and M.E. Sorrells. 2004. Identification of drought-inducible genes and differentially expressed sequence tags in barley. Theor. Appl. Genet. 109:1417-1425.

Edney M.J., and D.E Mather. 2004. Quantitative trait loci affecting germination traits and malt friability in a two-rowed by six-rowed barley cross. J. Cereal Sci. 39:283-290.

Han, F., J.A. Clancy, B.L. Jones, D.M. Wesenberg, A. Kleinhofs, and S.E. Ullrich. 2004. Dissection of a malting quality QTL region on chromosome 1 (7H) of barley. Molec. Breed. 14:339-347.

Komatsuda, T., P. Maxim, N. Senthil, and Y. Mano. 2004. High-density AFLP map of nonbrittle rachis 1 (btr1) and 2 (btr2) genes in barley (Hordeum vulgare L.). Theor. Appl. Genet. 109:986-995.

Potokina, E., M. Caspers, M. Prasad, R. Kota, H. Zhang, N. Sreenivasulu, M. Wang, and A. Graner. 2004. Functional association between malting quality trait components and cDNA array based expression patterns in barley (Hordeum vulgare L.). Molec. Breed. 14:153:170.

Rajasekaran, P., W.T.B. Thomas, A. Wilson, P. Lawrence, G. Young, and R.P. Ellis. 2004. Genetic control over grain damage in a spring barley mapping population. Plant Breed. 123:17-23.

Russell, J., A. Booth, J. Fuller, B. Harrower, P. Hedley, G. Machray, and W. Powell. 2004. A comparison of sequence-based polymorphism and haplotype content in transcribed and anonymous regions of the barley genome. Genome 47:389-398.

Stephens, J.L., S.E. Brown, N.L.V. Lapitan, and D.L. Knudson. 2004. Physical mapping of barley genes using an ultrasensitive fluorescence in situ hybridization technique. Genome 47:179-189.

Varshney, R.K., R. Sigmund, A. Börner, V. Korzun, N. Stein, M.E. Sorrells, P. Langridge, and A. Graner. 2004. Interspecific transferability and comparative mapping of barley EST-SSR markers in wheat, rye and rice. Plant Sci. 168:195-202.

Wenzl, P., J. Carling, D. Kudrna, D. Jaccoud, E. Huttner, A. Kleinhofs, and A. Kilian. 2004. Diversity arrays technology (DArT) for whole-genome profiling of barley. Proc. Natl. Acad. Sci. USA 101:9915-9920.

Integrating Molecular and Morphological/Physiological Marker Maps

A. Kleinhofs

Dept. Crop and Soil Sciences and

School of Molecular Biosciences

Washington State University

Pullman, WA 99164-6420, USA

e-mail: andyk@wsu.edu

Barley gene mapping and cloning is progressing, albeit slowly. The Uzu and Nec1 genes were identified by homology to Arabidopsis and rice genes (Chono et al., Plant Phys. 133:1209, ’03; Rostoks et al., in press). The Uzu gene encodes a brassinosteroid receptor and maps to chromosome 3H Bin6. The Nec1 gene encodes a cyclic nucleotide-gated ion channel 4 protein and maps to chromosome 5(1H) Bin9. There should be ample opportunities for the identification of other barley genes by homology to the model dicot and monocot plants, Arabidopsis and rice. For example the rym4,5,6 gene (resistance to yellow mosaic virus) was identified as translation initiation factor eIF4E by recognition of a similar virus resistance pathway in dicotyledonous species (Kanyuka et al., in press). The location of rym4 on chromosome 3H Bin016 was previously known (Graner et al., TAG 86:689 ´93). The locus is now marked by the CAPS marker BGK105 derived from the eIF4E sequence identified as the Rym4 gene. The Vrs1 gene was mapped at high resolution and shown to co-segregate with markers e40m36-1110 and e34m13-260 (He et al., Genome 47:1122, ’04). The previously closest marker MWG699 is located 0.1 cM proximal from Vrs1.

Please advise me if you have additions or corrections to this information.

Bin Assignments for Morphological Map Markers and closest molecular marker

Chr.1(7H)

BIN1 *Rpg1 RSB228 Brueggeman et al., PNAS 99:9328, ‘02

Run1

Rdg2a MWG851A Bulgarelli et al., TAG 108:1401, ‘04

Rrs2 MWG555A Schweizer et al., TAG 90:920, ‘95

mlt

brh1 MWG2074BLi et al., 8^th IBGS 3:72, ‘00

BIN2 Est5 iEst5 Kleinhofs et al., TAG 86:705, ‘93

fch12 BCD130 Schmierer et al., BGN 31:12, ‘01

*wax Wax Kleinhofs BGN 32:152, ‘02

gsh3 His3A Kleinhofs BGN 32:152, ‘02

BIN3 fch5 ABC167A Kleinhofs BGN 32:152, ‘02

Rcs5 KAJ185 Johnson & Kleinhofs, unpublished

yvs2

cer-ze ABG380 Kleinhofs BGN 27:105, ‘96

BIN4 wnd

Lga BE193581 Johnson & Kleinhofs, unpublished

abo7

BIN5 ant1

nar3 MWG836 Kleinhofs BGN 32:152, ‘02

ert-m

ert-a

BIN6 ert-d

fch8

fst3

cer-f

dsp1

msg14

BIN7 msg10

rsm1 ABC455 Edwards & Steffenson, Phytopath. 86:184,’96 sex6

seg5

seg2

pmr ABC308 Kleinhofs BGN 27:105, ‘96

mo6b Hsp17 Soule et al., J Her. 91:483, ‘00

nud CDO673 Heun et al., Genome 34:437, ‘91

fch4 MWG003 Kleinhofs BGN 27:105, ‘96

BIN8 *Amy2 Amy2 Kleinhofs et al., TAG 86:705, ‘93

lks2 WG380B Costa et al., TAG 103:415, ‘01

Rpt4 Psr117D Williams et al., TAG 99:323, ‘99

ubs4

blx2

BIN9 lbi3

xnt4

lpa2 ? Larson et al., TAG 97:141, ‘98

msg50

Rym2

seg4

BIN10 Xnt1 BF626025 Hansson et al., PNAS 96:1744, ‘99

xnt-h BF626025 Hansson et al., PNAS 96:1744, ‘99

BIN11 Rph3 Tha2 Toojinda et al., TAG 101:580, ‘00

BIN12 Mlf

xnt9

seg1

msg23

BIN13 Rph19 Rlch4(Nc) Park & Karakousis Plt. Breed. 121:232. ‘02

BIN14 none

Chr.2(2H)

BIN1 sbk

BIN2 none

BIN3 gsh6 MWG878A Kleinhofs BGN 32:152, ‘02

gsh1

gsh8

BIN4 Eam1

Ppd-H1 MWG858 Laurie et al., Heredity 72:619, ‘94

sld2

rtt

flo-c

sld4

BIN5 fch15

brc1

com2

BIN6 msg9

abo2

Rph15 P13M40 Weerasena et al., TAG 108:712 ‘04

rph16 MWG874 Drescher et al., 8thIBGS II:95, ‘00

BIN7 yst4 CDO537 Kleinhofs BGN 32:152, ‘02

Az94 CDO537 Kleinhofs BGN 32:152, ‘02

gai MWG2058 Börner et al., TAG 99:670, ‘99

msg33

msg3

fch1

BIN8 Eam6 ABC167b Tohno-oka et al., 8thIBGS III:239, ‘00

gsh5

msg2

eog ABC451 Kleinhofs BGN 27:105, ‘96

abr

cer-n

BIN9 Gth

hcm1

wst4

vrs1 MWG699 Komatsuda et al., Genome 42:248, ‘00

BIN10 cer-g

Lks1

mtt4

Pre2

msg27

ant2

BIN11 Rha2 AWBMA21 Kretschmer et al., TAG 94:1060, ‘97

*Rar1 AW983293BFreialdenhoven et al., Plt. Cell 6:983, ’94

fol-a

gal MWG581A Börner et al., TAG 99:670, ‘99

fch14

Pau

BIN12 Pvc

BIN13 lig BCD266 Pratchett & Laurie Hereditas 120:35, ‘94

nar4 Gln2 Kleinhofs BGN 27:105, ‘96

Zeo1 cnx1 Costa et al., TAG 103:415, ‘01

lpa1 ABC157 Larson et al., TAG 97:141, ‘98

BIN14 none

BIN15 gpa CDO036 Kleinhofs BGN 27:105, ‘96

wst7 MWG949A Costa et al., TAG 103:415, ‘01

MlLa Ris16 Giese et al., TAG 85:897, ‘93

trp

Chr. 3(3H)

BIN1 BE216031; BF264341; BF623053

Rph5

Rph6 BCD907 Zhong et al., Phytopath. 93:604, ‘03

Rph7 MWG848 Brunner et al., TAG 101:783, ‘00

BIN2 BI958652; BF631357; BG369659

ant17

sld5

mo7a ABC171A Soule et al., J. Hered. 91:483, ‘00

brh8

BIN3 ARD1769.1 Druka et al., PNAS 99:850, ‘02

xnt6

BIN4 btr1

btr2

lzd

BIN5 alm ABG471 Kleinhofs BGN 27:105, ‘96

abo9

sca

yst2

dsp10

BIN6 BI956389; BE456118B

Rrs1 Graner et al., TAG 93: 421 ´96

BG418711

Rh/Pt ABG396 Smilde et al., 8^th IBGS 2:178, ‘00

BE455901

Rrs.B87 BCD828 Williams et al., Plant Breed. 120:301, ‘01

AtpbB

abo6

xnt3

msg5

ari-a

yst1

zeb1

ert-c

ert-ii

cer-zd

Ryd2 WG889B Collins et al., TAG 92:858, ‘96

*uzu AB088206 Saisho et al., Breeding Sci. 54:409, ‘04

BIN7 cer-r

BIN8 wst6

cer-zn

sld1

BIN9 wst1

BIN10 vrs4

Int1

gsh2

BIN11 als

sdw1 PSR170 Laurie et al., Plant Breed. 111:198, ‘93

BIN12 sdw2

BIN13 Pub ABG389 Kleinhofs et al., TAG 86:705, ‘93

BIN14 cur2

BIN15 Rph10

fch2

BIN16 eam10

Est1/2/3

*rym4 eIF4E Kanyuka et al., in press ‘05

*rym5 eIF4E Kanyuka et al., in press ’05

Stein et al., Plt.J. 42:912, ‘05

Est4

ant28

Chr.4(4H)

BIN1 none

BIN2 fch9

sln

BIN3 int-c MWG2033 Komatsuda, TAG 105:85, ‘02

Zeo3

Dwf2 Ole1 Ivandic et al., TAG 98:728, ‘99

Ynd

glo-a

rym1 MWG2134 Okada et al., Breeding Sci. 54:319, ‘04

BIN4 *Kap X83518 Muller et al., Nature 374:727, ‘95

lbi2

zeb2

lgn3

BIN5 lgn4

lks5

eam9

msg24

BIN6 glf1

rym11 MWG2134 Bauer et al., TAG 95:1263, ‘97

Mlg MWG032 Kurth et al., TAG 102:53, ‘01

cer-zg

brh2

BIN7 glf3

frp

min1

blx4

sid

blx3

BIN8 blx1

BIN9 ert1

BIN10 *mlo P93766 Bueschges et al., Cell 88:695, ‘97

BIN11 none

BIN12 Hsh HVM067 Costa et al., TAG 103:415, ‘01

Hln