• We are sorry, but NCBI web applications do not support your browser and may not function properly. More information
Logo of plntphysLink to Publisher's site
Plant Physiol. Jan 2009; 149(1): 142–147.
PMCID: PMC2613708
Focus Issue on the Grasses

The International Barley Sequencing Consortium—At the Threshold of Efficient Access to the Barley Genome1,[W]


Archaeological evidence indicates that barley (Hordeum vulgare) and wheat (Triticum aestivum) were domesticated 10,000 years ago in the Fertile Crescent (Zohary and Hopf, 2001). Among the cereals, barley currently ranks fourth after maize (Zea mays), rice (Oryza sativa), and wheat in terms of total production (Fig. 1). An early maturation and a high level of adaptability to stressful conditions (including cold, drought, alkali, and saline soils) make it well suited for cultivation throughout the world from boreal to equatorial regions. About two-thirds of the global barley crop is used for animal feed, while the remaining third underpins the malting, brewing, and distilling industries. Although the human diet is not a primary use, barley offers a wealth of potential health benefits (Baik and Ullrich, 2008) and is still the major calorie source in several parts of the world (Grando and Macpherson, 2005).

Figure 1.
Statistics for global barley production. Top left section: global barley production parameters for the past 45 years. The top right, bottom left, and bottom right sections give worldwide area, production, and yield parameters for barley for 2006 relative ...


Barley is a diploid inbreeding species with a large genome of >5 Gbp (Bennett and Smith, 1976). Barley, bread wheat, rye (Secale cereale), and their respective wild relatives are closely related and form the Triticeae tribe, which evolved some 12 million years ago within the Pooideae subfamily of the Poaceae (Gaut, 2002). Due to its simpler diploid genome, barley can be considered a good genomic model for hexaploid bread wheat.

The large size and comparatively low gene density of Triticeae genomes provide a substantial challenge to gene isolation. Therefore, model grass species (e.g. rice, Brachypodium) having small genomes may serve as vehicles for synteny-based gene isolation in large genome cereals (for review, see Stein and Graner, 2004). However, barley and rice shared their last common ancestor about 70 million years ago; only 50% of barley genes highly homologous to rice are collinearly arranged in both genomes (Stein et al., 2007). This limitation may not be fully overcome with the more closely related grass model species, Brachypodium distachyon (Draper et al., 2001). Recent, accelerated development of genomic resources for agricultural grass species are poised to alleviate these limitations, concomitantly strengthening our understanding of pan-grass genomics. The revised model of an ancestral grass genome obtained by comparing whole genome sequences and high density transcript maps of model and nonmodel grass species, respectively (Salse et al., 2008), provides a glimpse of the possibilities.


Molecular studies on barley genes and traits has provided (1) a better understanding of Poaceae and plant biology, (2) confirmed and generalized results obtained in other grass species, and (3) revealed barley-specific trait expression controlled by genes highly conserved in related grasses. For example, Poaceae seeds, called caryopses, share a common architecture whereby the embryo is attached to the starchy endosperm via the scutellum—a tissue from which the embryo derives nutrients during seed germination. Due to its paramount importance for the malting and brewing industry (malting is controlled seed germination) and through recent advances in functional genomics, the barley grain is one of the best-studied systems in cereal crops (Gubatz et al., 2007; Sreenivasulu et al., 2008b) and can be regarded as a general model for both Poaceae seed development and germination.

Studies of disease caused by powdery mildew (Blumeria graminis f. sp. hordei) pathogens have been pioneered in barley (Schulze-Lefert and Panstruga, 2003). Factors involved in pathogen recognition, signal transduction, and resistance response have been isolated and characterized, thereby accelerating analyses of host-pathogen interactions across taxonomic boundaries (Jones and Dangl, 2006).

Genes that control responses to important abiotic or environmental factors, such as vernalization requirements (Fu et al., 2005) and photoperiod responses (Turner et al., 2005), have been isolated from barley and either informed previous work on wheat (Vrn1; Cockram et al., 2007) or the barley gene was used as a vehicle to study wheat gene function (e.g. Ppd-D1; Beales et al., 2007).

Three major genes controlling barley flower or inflorescence morphology have been isolated and characterized. The Vrs1 locus controls lateral spikelet fertility, which determines the number of rows in barley ears (Komatsuda et al., 2007). The Nud gene controls lipid biosynthesis in the pericarp, which glues the husks to the kernels and results in either covered or naked barley (Taketa et al., 2008). Dominant mutations in the hooded gene result in an extra flower with inverse polarity on the lemma of the barley spikelet (Müller et al., 1995). Vrs1 and Nud encode, respectively, homeodomain-Leu zipper I and ethylene response family transcription factors. In both cases, highly conserved orthologs with undetermined function are present in rice and likely in other Poaceae. The hooded phenotype is caused by ectopic expression of the barley Knox3 homeobox transcription factor gene in the lemma. In maize, ectopic expression of the orthologous Knotted1 gene results in knotted leaf blade anatomy (Smith et al., 1992). Comparative studies of such genes in barley and other Poaceae will lead to a more comprehensive understanding of plant development and evolution.


Natural and Induced Genetic Variation

Despite the uniformity of modern, high-yielding, elite, inbred varieties, barley morphology is very diverse (Fig. 2). Representative collections of existing natural genetic diversity are maintained worldwide in ex situ genebanks (Supplemental Table S1), which hold an estimated 370,796 accessions of 31 Hordeum species (van Hintum and Menting, 2003). Since the late 1920s, barley has also been widely used to study and utilize induced genetic variability (summarized in Barley Genetics Newsletter Volume 26: http://wheat.pw.usda.gov/ggpages/bgn/26/bgn26tc.html). Recently, the TILLING technique (targeting induced local lesions in genomes; Caldwell et al., 2004; Talame et al., 2008) has reinvigorated mutation research in barley.

Figure 2.
Barley diversity. A sample of barley inflorescences (spikes) is shown to illustrate the highly polymorphic nature of barley genetic resources, in profound contrast to the uniform morphology of modern elite cultivars.

From Early Molecular Genetic Linkage Analysis to High-Throughput Genotyping

Molecular markers couple barley genome diversity with crop plant biology and facilitate positional cloning and knowledge transfer into crop improvement. Since the early RFLP mapping in barley (Graner et al., 1991), numerous genetic maps utilizing the full spectrum of molecular markers have been published (Supplemental Table S2; http://wheat.pw.usda.gov/ggpages/map_summary.html). Altogether, a minimum of 5,000 nonredundant barley genes have been genetically mapped so far (L. Ramsay and A. Druka, personal communication).

Barley Functional and Structural Genomics

Resources for barley functional genomics have improved over the last decade (for review, see Sreenivasulu et al., 2008a), initiated by systematic EST sequencing (Zhang et al., 2004). Currently, 478,734 public ESTs (dbEST summary 08.08.08, http://www.ncbi.nlm.nih.gov/dbEST/dbEST_summary.html) provide partial sequence information for 50,000 tentative unigenes (Sreenivasulu et al., 2008a). Based on this community resource the first comprehensive oligonucleotide array for a crop plant (22 K Barley 1 GeneChip; Close et al., 2004) was designed. PLEXdb (http://plexdb.org/; Wise et al., 2007) provides public, online access to over 1,200 GeneChip hybridizations from barley and wheat of diverse developmental stages and tissues (Druka et al., 2006). The WebComparator (http://contigcomp.acpfg.com.au/) facilitates interspecies comparisons of expression profiles for Triticeae homoeologs.

The current understanding of barley genome structure is based mainly on the sequence of a handful of bacterial artificial chromosome (BAC) clones (summarized in Stein, 2007) selected for map-based cloning. Map-based gene isolation is feasible in barley at the single-gene scale. However, the efficiency of the entire procedure remains limited by the lack of a complete physical map or a reference genome sequence.


The lack of a physical map and complete sequence of the barley genome is related to the scale of a barley genome project. As this far exceeds the capacity of individual laboratories, it triggered the formation of the International Barley Genome Sequencing Consortium (IBSC; http://barleygenome.org) in 2006. Barley researchers of the eight founding institutions from six different countries comprise a steering committee that elects from its members a chair and a cochair who represent the consortium to the scientific community, funding bodies, and governments. Semiannual steering committee meetings are held to review the progress made toward IBSC goals and to reassess the consortium's strategy for sequencing in view of technological developments and progress made. Similar to previous initiatives, particularly rice and maize (Sasaki and Burr, 2000; Coe et al., 2002), the barley roadmap toward a high-quality genome sequence includes the construction of a genetically anchored physical map. Such a map is not indispensible in some whole-genome sequencing strategies, but in itself it forms an invaluable resource to the scientific community for gene isolation. It represents a milestone that can be reached at a fraction of the cost required for sequencing the whole barley genome.


To initiate physical map development, a set of 83,381 gene-bearing clones were identified by overgo hybridizations to the original public barley BAC library (Yu et al., 2000; Madishetty et al., 2007), then analyzed and assembled through high information content fingerprinting (Luo et al., 2003; preliminary contig information: http://phymap.plantsciences.ucdavis.edu:8080/barley/). Subsequently, a generic approach to building a genome-wide physical map of barley by high information content fingerprinting was initiated and scheduled to be accomplished by the end of 2009 (status September 2008: 308,764 high-quality fingerprints, approximately seven haploid genome equivalents; D. Schulte, N. Stein, P. Langridge, and A. Graner, unpublished data). This effort takes advantage of new BAC library resources (Supplemental Table S3) and aims at fingerprinting BAC clones representing up to 14 genome equivalents (based on an average insert size of 125 kb). Based on previous calculations (Lander and Waterman, 1988; Wendl and Waterston, 2002), this should resolve a total of approximately 7,000 to 10,000 contigs for the entire barley genome (approximately 1,000 contigs/chromosome), with 70% overlap between individual BACs.


A physical map becomes effective only after anchoring to a genetic map. Large numbers of mapped markers are available for the barley genome (Supplemental Table S2) and an IBSC goal is to obtain 10,000 mapped barley genes (theoretical coverage of two mapped genes/megabase). Mapped marker and BAC address relationships can be determined by hybridization to high-density colony filters (Madishetty et al., 2007) or by PCR screening of multidimensional BAC DNA pools (i.e. Yim et al., 2007). Higher anchoring throughput may be achieved if established multiplex marker platforms in barley, e.g. DArT (Wenzl et al., 2006) and the Golden Gate Assay (Rostoks et al., 2006), can be adopted to BAC DNA pool screening.

The integration of physical and genetic maps in barley is challenged by the limited genetic resolution of current mapping populations. High-density maps in barley mostly represent composite maps providing limited genetic resolution (approximately 1 cM). Large recombinant inbred populations of approximately 4,000 individuals are under development (P. Langridge and N. Stein, unpublished data) to increase genetic resolution. Larger populations or alternative population designs, e.g. intermated recombinant inbred populations (Liu et al., 1996), will be unlikely to resolve regions lacking recombination like the genetic centromeres of the barley chromosomes. Ordering of contigs for these will require the use of alternative methods such as HAPPY mapping (Thangavelu et al., 2003) to reveal physical marker order. Colinear gene order between barley and sequenced grass genomes, combined with chromosome evolution models (Salse et al., 2008), will serve to predict the placement of BAC clones and contigs to genomic regions lacking genetic mapping information.


IBSC aims at developing a high-quality gold standard reference sequence for barley as the basis for whole-genome single nucleotide polymorphism surveys and genome resequencing. To this end, next generation sequencing (NGS) technologies provide new options by reducing sequencing costs and increasing throughput (Shendure and Ji, 2008). Initial studies demonstrated that NGS technologies can be suitable for determining gene content of barley BAC clones (Wicker et al., 2006) and can support automated repeat annotation of barley genomic DNA (Wicker et al., 2008). New technologies impact sequencing strategies because whole-genome shotgun sequencing (WGS), a proven means of sequencing medium-sized plant genomes such as grapevine (The French-Italian Public Consortium for Grapevine Genome Characterization, 2007), promises to become a rapid, universal approach. However, so far there are no published examples demonstrating that WGS can be applied to large genomes like barley that feature >80% of repetitive DNA. Therefore, a combinatorial approach, combining the strengths of map-based sequencing (classical Sanger or NGS) of a minimal tiling path with paired-end WGS sequencing, can be envisaged currently as a strategy for sequencing the barley genome.

Future challenges lie in the area of barley genome annotation and database construction. The specific aspects of Triticeae genome organization will require new or advanced tools for gene and repeat structure annotation. Here, ongoing sequencing of 25,000 full-length cDNA sequences of barley will provide valuable information (5,000 sequences have been deposited at the DNA Data Bank of Japan; http://www.shigen.nig.ac.jp/barley/; K. Sato and T. Matsumoto, unpublished data). The development of repeat annotation strategies (Wicker et al., 2007) will be critical for structural, functional, and comparative barley genomics with the accumulated knowledge connected wherever possible to public genomics resources (e.g. graingenes: http://wheat.pw.usda.gov/GG2/index.shtml; gramene: http://www.gramene.org/). To achieve this, vigorous links to related grass genome initiatives (rice, maize, Brachypodium, wheat) have been established. Furthermore, IBSC members are committed to raising funding for an international effort to develop a data warehouse environment for Triticeae genomics.


Sequencing the genome of barley, an agriculturally and industrially important cereal crop and a useful diploid model for bread wheat, has become a realistic undertaking. Important steps have been initiated to improve genomics tools, build and anchor a physical map, develop a high-density genetic map, assess new sequencing technologies, and generate substantial datasets of genomic survey information. These are coordinated through an international consortium. A high-quality reference genome sequence will not only further promote our understanding of genome evolution but also blaze the trail toward genomics-based crop improvement. It will inform our understanding of previously sequenced grass and other plant genomes and will become a further milestone toward understanding grass (or even general plant) genomics and systems biology.

Supplemental Data

The following materials are available in the online version of this article.

  • Supplemental Table S1. Ex situ genebanks hosting large collections of barley accessions and genetic stocks.
  • Supplemental Table S2. Overview of recent high-density biparental and consensus genetic maps of barley.
  • Supplemental Table S3. BAC library resources for barley physical mapping.

Supplementary Material

[Supplemental Data]


1This work was supported by grants to K.S. (NBRP, BRAIN, and MAFF, Japan), T.J.C. (National Science Foundation grant no. DBI–0321756, U.S. Department of Agriculture CSREES), and N.S. (WGL-Pakt für Forschung und Innovation, Germany).

The author responsible for distribution of materials integral to the findings presented in this article in accordance with the policy described in the Instructions for Authors (www.plantphysiol.org) is: Nils Stein (ed.nebelsretag-kpi@niets).

[W]The online version of this article contains Web-only data.



  • Baik BK, Ullrich SE (2008) Barley for food: characteristics, improvement, and renewed interest. J Cereal Sci 48 233–242
  • Beales J, Turner A, Griffiths S, Snape J, Laurie D (2007) A pseudo-response regulator is misexpressed in the photoperiod insensitive Ppd-D1a mutant of wheat (Triticum aestivum L.). Theor Appl Genet 115 721–733 [PubMed]
  • Bennett MD, Smith JB (1976) Nuclear DNA amounts in angiosperms. Philos Trans R Soc Lond B Biol Sci 274 227–274 [PubMed]
  • Caldwell DG, McCallum N, Shaw P, Muehlbauer GJ, Marshall DF, Waugh R (2004) A structured mutant population for forward and reverse genetics in barley (Hordeum vulgare L.). Plant J 40 143–150 [PubMed]
  • Close TJ, Wanamaker SI, Caldo RA, Turner SM, Ashlock DA, Dickerson JA, Wing RA, Muehlbauer GJ, Kleinhofs A, Wise RP (2004) A new resource for cereal genomics: 22K barley GeneChip comes of age. Plant Physiol 134 960–968 [PMC free article] [PubMed]
  • Cockram J, Jones H, Leigh FJ, O'Sullivan D, Powell W, Laurie DA, Greenland AJ (2007) Control of flowering time in temperate cereals: genes, domestication, and sustainable productivity. J Exp Bot 58 1231–1244 [PubMed]
  • Coe E, Cone K, McMullen M, Chen SS, Davis G, Gardiner J, Liscum E, Polacco M, Paterson A, Sanchez-Villeda H, et al (2002) Access to the maize genome: an integrated physical and genetic map. Plant Physiol 128 9–12 [PMC free article] [PubMed]
  • Draper J, Mur LAJ, Jenkins G, Ghosh-Biswas GC, Bablak P, Hasterok R, Routledge APM (2001) Brachypodium distachyon: a new model system for functional genomics in grasses. Plant Physiol 127 1539–1555 [PMC free article] [PubMed]
  • Druka A, Muehlbauer G, Druka I, Caldo R, Baumann U, Rostoks N, Schreiber A, Wise R, Close T, Kleinhofs A, et al (2006) An atlas of gene expression from seed to seed through barley development. Funct Integr Genomics 6 202–211 [PubMed]
  • Fu D, Szűcs P, Yan L, Helguera M, Skinner JS, von Zitzewitz J, Hayes PM, Dubcovsky J (2005) Large deletions within the first intron in VRN-1 are associated with spring growth habit in barley and wheat. Mol Genet Genomics 273 54–65 [PubMed]
  • Gaut BS (2002) Evolutionary dynamics of grass genomes. New Phytol 154 15–28
  • Grando S, Macpherson HG, editors (2005) Food Barley: Importance, Uses and Local Knowledge. ICARDA, Aleppo, Syria
  • Graner A, Jahoor A, Schondelmaier J, Siedler H, Pillen K, Fischbeck G, Wenzel G, Herrmann RG (1991) Construction of an RFLP map of barley. Theor Appl Genet 83 250–256 [PubMed]
  • Gubatz S, Dercksen VJ, Brüß C, Weschke W, Wobus U (2007) Analysis of barley (Hordeum vulgare) grain development using three-dimensional digital models. Plant J 52 779–790 [PubMed]
  • Jones JDG, Dangl JL (2006) The plant immune system. Nature 444 323–329 [PubMed]
  • Komatsuda T, Pourkheirandish M, He C, Azhaguvel P, Kanamori H, Perovic D, Stein N, Graner A, Wicker T, Tagiri A, et al (2007) Six-rowed barley originated from a mutation in a homeodomain-leucine zipper I-class homeobox gene. Proc Natl Acad Sci USA 104 1424–1429 [PMC free article] [PubMed]
  • Lander ES, Waterman MS (1988) Genomic mapping by fingerprinting random clones: a mathematical analysis. Genomics 2 231–239 [PubMed]
  • Liu SC, Kowalski SP, Lan TH, Feldmann IA, Paterson AH (1996) Genome-wide high-resolution mapping by recurrent intermating using Arabidopsis thaliana as a model. Genetics 142 247–258 [PMC free article] [PubMed]
  • Luo MC, Thomas C, You FM, Hsiao J, Ouyang S, Buell CR, Malandro M, McGuire PE, Anderson OD, Dvorak J (2003) High-throughput fingerprinting of bacterial artificial chromosomes using the snapshot labeling kit and sizing of restriction fragments by capillary electrophoresis. Genomics 82 378–389 [PubMed]
  • Madishetty K, Condamine P, Svensson JT, Rodriguez E, Close TJ (2007) An improved method to identify BAC clones using pooled overgos. Nucleic Acids Res 35 e5. [PMC free article] [PubMed]
  • Müller K, Romano N, Gerstner O, Garcia-Maroto F, Pozzi C, Salamini F, Rohde W (1995) The barley Hooded mutation caused by a duplication in a homeobox gene intron. Nature 374 727–730 [PubMed]
  • Rostoks N, Ramsay L, MacKenzie K, Cardle L, Bhat PR, Roose ML, Svensson JT, Stein N, Varshney RK, Marshall DF, et al (2006) Recent history of artificial outcrossing facilitates whole-genome association mapping in elite inbred crop varieties. Proc Natl Acad Sci USA 103 18656–18661 [PMC free article] [PubMed]
  • Salse J, Bolot S, Throude M, Jouffe V, Piegu B, Quraishi UM, Calcagno T, Cooke R, Delseny M, Feuillet C (2008) Identification and characterization of shared duplications between rice and wheat provide new insight into grass genome evolution. Plant Cell 20 11–24 [PMC free article] [PubMed]
  • Sasaki T, Burr B (2000) International Rice Genome Sequencing Project: the effort to completely sequence the rice genome. Curr Opin Plant Biol 3 138–141 [PubMed]
  • Schulze-Lefert P, Panstruga R (2003) Establishment of biotrophy by parasitic fungi and reprogramming of host cells for disease resistance. Annu Rev Phytopathol 41 641–667 [PubMed]
  • Shendure J, Ji H (2008) Next-generation DNA sequencing. Nat Biotechnol 26 1135–1145 [PubMed]
  • Smith LG, Greene B, Veit B, Hake S (1992) A dominant mutation in the maize homeobox gene, Knotted-1, causes its ectopic expression in leaf cells with altered fates. Development 116 21–30 [PubMed]
  • Sreenivasulu N, Graner A, Wobus U (2008. a) Barley genomics: an overview. Int J Plant Genomics 2008 486258 [PMC free article] [PubMed]
  • Sreenivasulu N, Usadel B, Winter A, Radchuk V, Scholz U, Stein N, Weschke W, Strickert M, Close TJ, Stitt M, et al (2008. b) Barley grain maturation and germination: metabolic pathway and regulatory network commonalities and differences highlighted by new MapMan/PageMan profiling tools. Plant Physiol 146 1738–1758 [PMC free article] [PubMed]
  • Stein N (2007) Triticeae genomics: advances in sequence analysis of large genome cereal crops. Chromosome Res 15 21–31 [PubMed]
  • Stein N, Graner A (2004) Map-based gene isolation in cereal genomes. In P Gupta, R Varshney, eds, Cereal Genomics. Kluwer Academic Publishers, Dordrecht, The Netherlands, pp 331–360
  • Stein N, Prasad M, Scholz U, Thiel T, Zhang H, Wolf M, Kota R, Varshney RK, Perovic D, Graner A (2007) A 1000 loci transcript map of the barley genome—new anchoring points for integrative grass genomics. Theor Appl Genet 114 823–839 [PubMed]
  • Taketa S, Amano S, Tsujino Y, Sato T, Saisho D, Kakeda K, Nomura M, Suzuki T, Matsumoto T, Sato K, et al (2008) Barley grain with adhering hulls is controlled by an ERF family transcription factor gene regulating a lipid biosynthesis pathway. Proc Natl Acad Sci USA 105 4062–4067 [PMC free article] [PubMed]
  • Talame V, Bovina R, Sanguineti MC, Tuberosa R, Lundqvist U, Salvi S (2008) TILLMore, a resource for the discovery of chemically induced mutants in barley. Plant Biotechnol J 6 477–485 [PubMed]
  • Thangavelu M, James AB, Bankier A, Bryan GJ, Dear PH, Waugh R (2003) HAPPY mapping in a plant genome: reconstruction and analysis of a high-resolution physical map of a 1.9 Mbp region of Arabidopsis thaliana chromosome 4. Plant Biotechnol J 1 23–31 [PubMed]
  • The French-Italian Public Consortium for Grapevine Genome Characterization (2007) The grapevine genome sequence suggests ancestral hexaploidization in major angiosperm phyla. Nature 449 463–467 [PubMed]
  • Turner A, Beales J, Faure S, Dunford R, Laurie D (2005) The pseudo-response regulator Ppd-H1 provides adaptation to photoperiod in barley. Science 310 1031–1034 [PubMed]
  • van Hintum T, Menting F (2003) Diversity in ex situ genebank collections of barley. In R von Bothmer, T van Hintum, H Knüpffer, K Sato, eds, Diversity in Barley (Hordeum vulgare). Elsevier Science B.V., Amsterdam, pp 247–257
  • Wendl MC, Waterston RH (2002) Generalized gap model for bacterial artificial chromosome clone fingerprint mapping and shotgun sequencing. Genome Res 12 1943–1949 [PMC free article] [PubMed]
  • Wenzl P, Li H, Carling J, Zhou M, Raman H, Paul E, Hearnden P, Maier C, Xia L, Caig V, et al (2006) A high-density consensus map of barley linking DArT markers to SSR, RFLP and STS loci and agricultural traits. BMC Genomics 7 206. [PMC free article] [PubMed]
  • Wicker T, Narechania A, Sabot F, Stein J, Vu GTH, Graner A, Ware D, Stein N (2008) Low-pass shotgun sequencing of the barley genome facilitates rapid identification of genes, conserved non-coding sequences and novel repeats. BMC Genomics 9 518. [PMC free article] [PubMed]
  • Wicker T, Sabot F, Hua-Van A, Bennetzen JL, Capy P, Chalhoub B, Flavell A, Leroy P, Morgante M, Panaud O, et al (2007) A unified classification system for eukaryotic transposable elements. Nat Rev Genet 8 973–982 [PubMed]
  • Wicker T, Schlagenhauf E, Graner A, Close TJ, Keller B, Stein N (2006) 454 sequencing put to the test using the complex genome of barley. BMC Genomics 7 275. [PMC free article] [PubMed]
  • Wise R, Caldo R, Hong L, Shen L, Cannon E, Dickerson J (2007) BarleyBase/PLEXdb: A unified expression profiling database for plants and plant pathogens. In D Edwards, ed, Methods in Molecular Biology, Plant Bioinformatics—Methods and Protocols, Vol 406. Humana Press, Totowa, NJ, pp 347–363 [PubMed]
  • Yim YS, Moak P, Sanchez-Villeda H, Musket TA, Close P, Klein PE, Mullet JE, McMullen MD, Fang Z, Schaeffer ML, et al (2007) A BAC pooling strategy combined with PCR-based screenings in a large, highly repetitive genome enables integration of the maize genetic and physical maps. BMC Genomics 8 47. [PMC free article] [PubMed]
  • Yu Y, Tomkins JP, Waugh R, Frisch DA, Kudrna D, Kleinhofs A, Brueggeman RS, Muehlbauer GJ, Wise RP, Wing RA (2000) A bacterial artificial chromosome library for barley (Hordeum vulgare L.) and the identification of clones containing putative resistance genes. Theor Appl Genet 101 1093–1099
  • Zhang H, Sreenivasulu N, Weschke W, Stein N, Rudd S, Radchuk V, Potokina E, Scholz U, Schweizer P, Zierold U, et al (2004) Large-scale analysis of the barley transcriptome based on expressed sequence tags. Plant J 40 276–290 [PubMed]
  • Zohary D, Hopf M (2001) Domestication of Plants in the Old World, Ed 3. Oxford University Press, Oxford, UK, pp 1–12

Articles from Plant Physiology are provided here courtesy of American Society of Plant Biologists
PubReader format: click here to try


Related citations in PubMed

See reviews...See all...

Cited by other articles in PMC

See all...


Recent Activity

Your browsing activity is empty.

Activity recording is turned off.

Turn recording back on

See more...